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INTEENATIONAL  SCIENTIFIC  SEEIES 

VOL.  LXXXVII 


KEGAN  PAUL,  TRENCH,  TRUBNER  &  CO.,  Ltd.'s 
NEW  SCIENTIFIC  WORKS. 


IMPORTANT  ADDITIONS 
TO  THE  ''INTERNATIONAL  SCIENTIFIC  SERIES." 

ON    BUDS    AND    STIPULES. 

By  the  Right  Hon.  Sir  John  Lubbock,  Bart.,  M.P.,  F.R.S. 
"With  4  coloured  plates  and  many  cuts  in  the  text.    Crown  8vo.    68. 

EVOLUTION    BY    ATROPHY: 

IN   BIOLOGY  AND  SOCIOLOGY. 

By  Jban  Demook,  Jean  Massaet,  and  Emile  Vandeeveldk. 

Translated  by  Mrs  Chalmers  Mitchell. 

With  84  figures.    Crown  Svo.    6s. 

SEISMOLOGY. 

By  John  Milne,  F.R.S.,  F.G.S., 

Late  Professor  of  Mining,  Geology,  and  Seismology  in  the  Imperial  College 

of  Engineering,  Tokio,  Japan ;  Hon.  Fellow  of  King's  College,  London. 

With  63  figures.    Crown  Svo.    6s. 


In  one  Volume  Demy  Svo,  with  many  Illustrations  from  wood-blocks 
especially  executed  for  this  work. 

THE    GEOGRAPHY   OF    MAMMALS. 

By  W.  L.  ScLATER,  Director  of  the  South  African  Museum,  and 

P.  L.  ScLATER,  Secretary  of  the  Zoological  Society  of  London. 

With  8  maps.    Demy  Svo. 


LONDON :  PATERNOSTER  HOUSE,  CHARING  CROSS  ROAD,  W.C. 


EVOLUTION  BY  ATROPHY 

IN  BIOLOGY  AND  SOCIOLOGY 


BY 

JEAN    DEMOOR  JEAN    MASSART 

AQRilG^    OF  THK  FBEE  UNIVER8ITT  CHARG^   DE   COUBS   OF    THK  FREE 

OF  BRUSSELS  UNIVEESITT  OF  BRUSSELS 

Emile  vandervelde 

PROFBSSOB  AT  THE  IN8TITDTB  OF  HAUTES  ilVDES  OF  BBDSSELS 


TRANSLATED   BY 

Mrs    CHALMERS    MITCHELL 


LONDON 
KEGAN  PAUL,  TRENCH,  TRUBNER  &  CO.,  Ltd, 

PATKRNOSTER  HOUSE,    CHARING  CROSS  ROAD 

J899 


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CONTENTS 


Prbface 


PAOB 

5 


INTRODUCTION 

1.  Societies  and  organisms.  2.  Individuals,  colonies  and 
societies.  3.  Communities  and  societies.  4.  Distinc- 
tive characters  of  societies  of  which  the  members  are 
imited  by  social  contract  .  .  .  . 


BOOK  I 

UNIVERSALITY   OF  DEGENERATIVE   EVOLUTION 

Part  I.  Degeneration  in  the  development  of  institutions 

and  organs             ......  21 

Cfmpter  I.  In  the  evolution  of  organs  all  modification  is 

necessarily  attended  by  degeneration     ...  22 

§  1.  Preliminary  considerations       ....  22 

(1)  Original  formation  of  organs  in  ancestral  forms   .  23 

(2)  Development  of  organs  in  the  individual  .            .  23 

(3)  Philogenetic  evolution  of  function             .            .  25 

(4)  Individual  adaptation       •.            .            .            .  26 
Section  I.  Transformation  of  organs  of  animals           .  30 

§  2.  Transformation  of  hemodynamic  organs  in  the  indivi- 
dual (metameric  appendages)  ...  30 
The  Cray -fish      ......              31 

§  3.  Transformation  of  homologous  organs  in  individuals 

of  different  species  (limbs) ....  41 

Limbs  adapted  to  an  aquatic  life  {Ceratodus,  Qrtha- 
canthus,  Protoptervs  amphibius,  Protopterus  annec- 
tens,  Lepidosiren        .....  43 

Limbs  adapted  to  a  terrestrial  life       ...  43 


285662 


CONTENTS 


1.  Adaptation  to  walking  on  two  legs  (man, 
birds).  2.  Adaptation  to  leaping  {Dlptis  cegyp- 
tins,  kangaroo,  Tardus  spectrum,  Rana  esculenta. 
3.  Adaptation  to  running  (horse,  ruminants).  4. 
Adaptation  to  flight  (birds,  Pterosaurians,  bats). 
5.  Adaptation  to  arboreal  life  (Arctocebus  cala- 
harensis,  Chamceleo).  6.  Adaptation  to  swimming 
(Cetaceans,  Sirenia).  7.  Adaptation  to  burrowing 
{Talpa  curopcea,  Heterocephalus)       ...  46 

Section  II.  Modification  of  the  organs  of  plants 

§  4.  Modification  of  hemodynamic  organs  in  the  indivi 

dual  (basilar  and  apical  leaves)      ...  68 

1.   Rosa    rugosa.     2.    Serratula   centauroides. 

Sagittaria   sagittifolia.     4.    Lathy ra   Aphaca. 

Nymphaea  dentata     .....  70 

§  5.  Modification  of  organs  which    are    homologous    in 

individuals  of  different  species  (foliage  leaves)     .  78 

1.  Adaptation  to  climbing  {Cobaeca  scandens,  Vicia 
pyrenaica,  Cucumis  sativus).  2.  Adaptation  to 
carnivorous  nutrition  {Utricularia,  Nepenthes, 
Drosera).  3.  Adaptation  to  an  aquatic  life  (Sagit- 
taria, Nymphaea,  Vallisneria,,  Poiamogeton,  Ranun- 
culvs,  Ouvirandra  fenestralis).  4.  Adaptation  to 
defence  against  ants  {Acacia  sphaerocephala).  5. 
Adaptation  to  drought  {Sempervivum).  6.  Adapta- 
tion to  defence  against  herbivorous  animals 
{Caragarva,  Ilex,  Mamillaria,  etc.)  .  .  78 

Chapter  11.   In  the  evolution  of  institutions  all  modificatioh 

is  necessarily  accompanied  by  degeneration        .  .  90 

§  1.  Modifications    of    similar    institutions    in  the  same 

society  ......  91 

(1)  The  communal  budgets  of  Belgium  .  .  92 

(2)  Budget  of  the  States  of  the  German  Empire         .  95 

(3)  The  budgets  of  Germany,  France,  and  England  .  97 

§  2.  Modification  of  similar  institutions  in  different  so(   J 

groups  (the  development  of  landed  property)         .  98 

1.  Family  property  (Montenegro).      2.  Village  pro- 
perty (Russia).     3.  Feudal  property  (England). 


CONTENTS 


4.  Public  or  collective  property  (Switzerland). 

5.  Corporative  property  (Belgium).     6.  Private 
property  (Switzerland).     7.  Summary     .  .  100 

Pakt  II.  Degeneration  in  the  evolution  of  organisms  and 

societies      .  .  .  .  .  .  .  115 

Chanter  1.  All  organisms  exhibit  rudimentary  organs  .  115 

Section  I.  Rudimentary  organs  of  animals      .  .  117 

§  1.  Rudimentary  organs  in  man     ....  117 

1.  Integumentary  system.  2.  Skeleton.  3.  Mus- 
cular system.  4.  Nervous  system.  5.  Digestive 
system.  6.  Vascular  system.  7.  Sense  organs. 
8.  Genito-urinary  system  .  ,  .  117 

§  2.  Rudimentary  organs  in  various  groups            .  .  121 

1.  Ccelenterates.      2.  Worms.      3.  Echinoderms.  4. 

Mollusca.    5.  Artbropodes.     6.  Vertebrates  .  121 

Section  II.  Rudimentary  organs  in  plants       .  .  145 

§  3.  Rudimentary  organs  in  various  groups  of  plants  .  145 
1.  Algae.     2.  Mushrooms.     3.  Bryophyta.     4.  Pteri- 

dophyta.    5.  Phanerogams  .            .  .  145 

§  4.  Reduced  organs  in  the  vegetative  apparatus  of  the 

Phanerogams       ,  .  .  .  .  .  149 

Chapter  II.  Survivals  exist  in  all  kinds  of  societies    .  .  151 

§  1 .  Instances  of  survival  in  various  groups  .  .  153 

(1)  Instances  of  survival  in  the  most  modem  social 
groups  (the  United  States)   ....  155 

(2)  Instances  of  survival  in  less  civilized  social  groups 
(Veddahs,  Fuegoes,  Australian  tribes  .  .  156 

§  2.  Survivals  of  ancient  forms  of  marriage,  and  of  the 

far  lily  in  Modern  Europe  ....  161 

1.  Forms  of  mm-rmge. — (1)  Marriage  by  capture.  (2) 
Marriage  by  purchase.  (3)  Marriage  by  consent 
of  both  parties  (marriage  by  simple  consent,  mar- 
"  iage  in  fade  Eccledoe)  ....  161 

2. '  The  Family  System. — (1)  Matriarchy.  (2)  Patri- 
archy .......  167 

Pabt  III.  Simimary  and  conclusions  ....  170 


CONTENTS 


BOOK  II 

THE  PATH  OP  DEGENERATIVE  EVOLUTION 


Part  I.  The  supposed  law  that  degeneration  retraces  the 

steps  of  progress    ..... 
Chapter  I.  The  path  of  degeneration  in  biology 

Section  I.  The  path  of  degeneration  in  animals 
1.  Morphology  and  embryology  ;  the  law  of  recapitu 
lation.    2.  Degeneration  of  the  third  eye  in  lizards, 
3.  Degeneration  of  the  organs  of  sight  in  deep-sea 
Crustacea.     4.  Atrophy  of  the  branchial  vessels  in 
man 
Section  II.  The  path  of  degeneration  in  plants. 
1.  Rarity  of  cases  of  recapitulation  in  the  organ 
ogeny  of  leaves  (Vicia,   Acacia  with  phyllodes) 
2.  Organogeny  of   flowers   {Brassica  oleracea  var, 
Botri/tis).    3.  Progressive  degeneration  of  the  pro 
thallus  in  phanerogams         ... 
Chapter  II.  The  path  of  degeneration  in  sociology    . 
§  1.  Investigation  of  facts     .... 

1.  Tithings,  hundreds  and  counties  in  England.  2, 
Order  of  elimination  of  various  racial  elements  in 
a  country.  3.  The  degenerative  evolution  of  pol 
tical  organizations.  4.  Degeneration  in  monetary 
systems.  5.  Degenerative  adaptation  in  colonial 
legislation.  6.  Degenerative  evolution  of  the  cor 
porations  of  Western  Flanders 
§  2.  A  criticism  of  the  supposed  inverse  path  of  degenera 
tion  ...... 

Part  II.  The  irreversibility  of  degenerative  evolution. 
Chapter  I.  Do  institutions  or  organs  which  have  disappeared 
reappear?       ..... 
Section  I.  Disappeared  organs  . 

1.  Plants  ..... 

2.  Animals.  Teratology  of  the  horse,  Hypertrichosis, 
etc.  Swimming  limbs  in  Stomatopoda  and  De 
capoda  Macroura       .... 

Section  II.  Disappeared  institutions    . 


175 
178 
179 


179 
192 


192 
205 
205 


207 

217 
221 

222 
222 
222 

223 
227 


CONTENTS  * 

PAGE 

(1)  Apparent  revival  of  bygone  institutions    .  .  227 

(2)  Apparent  disappearance  of  institutions     .  .  229 

(3)  Instances  of  convergence    ....  230 
Chapter  II.  Can  rudimentary  institutions  or  organs  reassume 

their  primitive  functions  ?  .  .  .  .  232 

Section  1.  Rudimentary  organs  .  .  .  232 

1.  Animals.  (1)  Muscles  of  the  ear  in  man.  (2)  The 
abdomen  and  appendages  in  deep-sea  hermit  crabs  232 

2.  Plants.  (1)  Hermaphrodite  flowers  in  Melaiidryum. 
(2)  Branches  of  Colletia  cruciaia,  Crataegus,  Vicia 

Faba,  Sempervivum,  Veronica,  etc.    .  .  .  233 

Section  II.  Rudimentary  institutions   .  .  .  237 

(1)  Truck  system  and  clearing-house.  (2)  Corpora- 
tions and  syndicates.  (3)  Archaic  collectivism  and 
modern  collectivism.  (4)  The  survival  of  elective 
sovereignty  in  England         ....  239 

Chapter  III.  Can  rudimentary  organs  or  institutions   re- 
develop and  assume  new  functions  ?  .  .  .  242 
Section  I.  Rudimentary  organs             .             .             .  243 

1.  Animals.  Respiratory  organs  in  Birgus  latro; 
Mesonephric  spaces  in  the  higher  vertebrates        .  243 

2.  Plants.     Staminodes  of  Pentstemon  .  .  244 
Section  II.  Rudimentary  institutions  .            .             .  245 

Levirat  .  .  .  .  .  .         _  .  245 

Part  III,  Summary  and  conclusions  .  .  .  247 

BOOK  III 

CAUSES  OF  DEGENERATIVE  EVOLUTION 

Part  I.  Atrophy  of  organs  and  institutions  .  .  .  251 

Section  7.  The  factors  of  atrophy     ....  251 

(1)  Biology  (accidental,  individual,  normal  and  specific 
atrophy).      (2)    Sociology    (accidental    and    normal 

atrophy)  ......  251 

Section  II.  Causes  producing  atrophy  .  .  .  260 

Chapter  I.  Atrophy  of  organs  .....  260 

§  1,  Atrophy  from  lack  of  space       ....  261 


CONTENTS 


(1)  Development  of  the  teeth.  (2)  Atrophy  of  the 
superior  ghime  {Lolhivi).  (3)  Degeneration  of 
Paleae  (composite  flowers)  and  of  stamens 
(Scrophulariaceae)  ....  261 

§  2.  Atrophy  from  lack  of  use  .  .  .  .  263 

1.  Functional  inutility  :    (1)  Etiolated  plants  and 

immobile  limbs.  (2)  Epicotyl  and  primary  leaf 
of  Nyviphaea.  (3)  Eoots  of  Utricularia  ;  coty- 
ledons of  parasitic  plants  {Guscuta,  Orobanche) ; 
leaves  transformed  to  spines  in  Phyllocactus 
a-eriatus.  (4)  Atrophy  of  the  branchial  arches  in 
mammals.  {5) Atro-phy of  yentrsd^ns {Pediculati, 
Protopttims).  (6)  Atrophy  of  muscles  {Cetacea, 
Sirenia).  (7)  Atrophy  of  the  tail  in  man.  (8) 
Degeneration  of  the  hyoid  apparatus  in  man 
and  birds  ......  263 

2.  Transference  of  function  :  (1)  Atrophy  of  the  tail 

in  Batrachia  anura,  and  the  larval  gills  in  some 
insects.  (2)  Disappearance  of  limbs  (Slow- 
worms,  Amphisbsena,  Snakes,  Eels,  and  Saccu- 
lina).  (3)  Atrophy  of  the  leaf  (acacia  with  phyl- 
\odes,  Xplophplla).  (4)  Atrophy  of  the  pro tonema 
in  mosses,  and  of  the  leaves  in  some  xerophilus 
plants  {MueMenhechia  plafpclados,  Genista, 
Spartium,  Alhagi).  (5)  Disappearance  of  the 
calyx.  (6)  Atrophy  of  roots  (Pine,  Beech, 
Corallorhyza,  Myrmechis,  Tillandsia  usneoides), 
or  of  leaves  and  stems  (TcBniophyllum  Zollin- 
geHy  Podostemacece)  ....  268 

§  3.  Atrophy  from  lack  of  nutrition  .  .  .  274 

1.  Parasitic    castration    {Melandryum    album).     2. 
Severe  or  prolonged  compression  of  a  limb. 

3.  Atrophy  of   the  genitalia  in  neuter  bees. 

4.  Atrophy  of  the  superior  flowers  in  Carex. 

5.  Atrophy  of  pistils  and  stamens  {Fritillaria 
persica,  Vibui-navi  tomentomm,  Viburnam  Opulus)  274 

§  4.  Atrophy  without  apparent  cause  .  .  .  278 

Atrophy  of  perianth  (J  rtemis/a,  Poterium).  Atrophy 
of  the  eyes  (Myriopodes,  C'ymotkoe).  Correlative 
atrophy        ......  278 


CONTENTS 

PAGE 

Chapter  II.  Atrophy  of  institutions    .  .  ■         .  .  281 

§  1.  Atrophy  from  lack  of  use  ....  282 

1.  Functional  inviility :  (1)  OflBces  of  the  port  of 

Bruges.     (2)  The  forest-courts  of  England         .  282 

2.  Transference  of  function  :  (1)  Republican  institu- 

tions under  the  Roman  Empire.     (2)  Special 
jurisdiction  in  Eogland    ....  284 

§  2.  Atrophy  from  lack  of  resource  .  ,  .  287 

1.  Local  administration  at  the  close  of  the  Roman 

Empire.   2.  Degeneration  of  societies  in  general  287 

Part  II.  Causes  of  the  persistence  of  organs  or  institutions 

without  function    ......  292 

Chjapter  I.  Survival  of  organs  .....  292 

§  1.  Unfunctional  organs  that  are  not  rudimentary  .  292 

Absence  of  variation  (flowers  of  Ficaria  ranuncu- 
loides,  Lysimachia  Nummularia,  Elodeaca)iadensis, 
stratiotes  aloides,  cleistogamous  flowers,  eyes  in 
the  male  Machaerites  ....  292 

§  2.  Unfunctional  organs  which  persist  as  rudiments         .  295 

1.  Absence  of  variation.    Insignificance  of  the  rudi- 
mentary organ  (stipules  of  Tropaeolum  majics, 
accessory  rudiments  of  enamel  organs  in  the 
development  of  teeth      ....  295 

Chapter  11  Survival  of  institutions     ....  298 

§  1.  Integral  persistence  of  an  institution  .  .  .  299 

(1)  Maintenance  by  compulsion  (rotten  boroughs  in 
England ;   the   States  of   Normandy  and  the 
Dauphiny  in  France).     (2)  Indirect  usefulness 
(English  monarchy).     (3)  Respect  for  tradition 
(royal  prerogatives  exercised  by  the  praetor; 
institution  of  sherififs  in  England)  .  .  299 

§  2.  Survival  of  institutions  in  a  reduced  condition  .  306 

1.  Insignificance  of  the  institution  (jurisdiction  at 
da  Martinique  ;  summons  fpio  wairanto ;  Diocese 
of  Cambrai ;  marsh-land  in  Artois ;  tribute  paid 
to  Spain  by  France).     Respect  for  traditions 
(survival  of  the  old  regime  in  England ;  instances 
of  survival  in  law  and  religion    .  .  .  307 

Part  III.  Summary  and  conclusions  ....  317 

General  conclusions     ......  320 


.    »  ». 


PREFACE 

This  treatise,  compiled  in  connection  with  a  scheme 
for  research  work  in  general  sociology,  elaborated 
in  June  1894,  was  presented  to  the  Institute  of 
Sociology  at  Brussels.  In  drawing  up  the  pro- 
gramme of  the  Institute,  the  founder,  M.  Ernest 
Solvay,  after  having  mentioned  the  questions  which 
especially  called  for  the  investigation  of  his  col- 
leagues, added  the  following  statement : — 

"  The  Institute  of  Sociology  will  take  part  in  the 
labours  of  the  modern  school  of  Sociology,  the  object 
of  which  is  to  ascertain  the  normal  conditions  under 
which  societies  exist,  and  the  laws  which  govern  their 
evolution.  But  the  advances  of  Natural  Science  in 
this  century  have  not  yet  been  sufficiently  assimilated 
by  those  Sciences  most  closely  related  to  it,  and  it 
is  from  such  assimilation  that  the  most  important 
additions  to  knowledge  may  be  expected." 

In  stating  that  the  results  of  Natural  Science 
have  not  been  sufficiently  assimilated  by  Sociology, 
M.  Solvay  is  only  apparently  at  variance  with 
those  who  rightly  protest  against  exaggerated  and 


•  ,«••• 


•6* .,  *  i  S.J  A  ^.'  PREFACE 

premature  comparisons  between  social  organizations 
and  animal  or  vegetable  organisms. 

The  existence  of  such  exaggerations,  which  have 
caused  a  reaction  such  as  recently  induced  an  emi- 
nent American  economist  to  declare  the  bankruptcy 
of  biological  sociology,  is  perhaps  due  to  the  fact 
that,  with  a  few  distinguished  exceptions,  bio- 
sociological  investigations  have  hitherto  been  con- 
ducted either  by  naturalists  with  a  limited  know- 
ledge of  social  questions,  or  by  sociologists  whose 
training  in  biology  was  incomplete  and  superficial. 

To  prevent  this  danger,  our  researches  in  the 
same  subject  have  been  made  separately  from  the 
social  side,  and  from  the  biological  side,  and  have 
now  been  co-ordinated  and  combined. 

This  work  was  commenced  in  May  1893,  with 
the  collaboration  of  our  friend  M.  Dollo,  the  curator 
of  the  Natural  History  Museum  at  Brussels.  In 
June  1894,  however,  M.  Dollo's  many  occupations 
no  longer  permitted  of  his  collaboration.  The  zoo- 
logical part  was  therefore  completed  by  M.  Jean 
Demoor,  to  whom  most  of  the  facts  quoted  in  the 
first  book  were  given  by  M.  Dollo,  whose  assistance 
we  most  gratefully  acknowledge. 


INTKODUCTION 

Ever  since  the  application  of  theories  of  evolution 
to  social  phenomena,  there  has  been  a  constant  inter- 
change in  terminology  between  biology  and  sociology ; 
societies  have  been  called  organisms,  and  organisms 
societies  of  cells.  There  is  an  actual  division  of 
labour  among  the  organs  of  a  living  body,  while 
institutions  have  been  called  the  organs,  or  parts 
of  organs,  of  Society.  The  interchange  of  matter 
effected  among  the  organs  of  an  individual  has  been 
called  a  "  physiological  contract,"  while  the  circula- 
tion of  money  may  be  compared  to  the  circulation 
of  blood  and  lymph.  ,  Questions  arise  as  to  what 
extent  such  comparisons  are  legitimate,  if  they 
should  be  taken  in  any  other  than  a  metaphorical 
sense,  and  if  it  is  possible  to  set  a  precise  boundary 
between  the  provinces  of  biology  and  sociology. 

Much  has  already  been  written  on  such  problems 
as  these,  and  no  doubt  much  more  will  yet  be 
written.  We  shall  not  attempt  either  to  discuss 
or  to  solve  them  in  these  few  introductory  pages  ; 
they  are  merely  touched  upon  here,  and  will  only 
be  alluded  to  when  absolutely  necessary  for  the 
careful  investigation  of  facts  bearing  upon  our  work, 

I.  Societies  and   organisms. — The  analogy  exist- 


8  EVOLUTION  BY  ATROPHY 

ing,  from  the  point  of  view  of  evolution,  between 
biology  and  sociology,  arises  from  the  fact  that  the 
evolution  of  societies  as  well  as  that  of  organisms, 
is  the  result  of  the  co-operation  of  two  factors — 
similarity  and  adaptation. 

In  biology,  the  similarity  between  organisms 
springing  from  the  same  stock,  is  due  to  heredity, 
while  adaptation  is  the  result  of  individual  variation. 

In  sociology,  societies  are  the  descendants  of 
former  societies,  in  that  the  new  are  modelled 
upon  the  old.  Similarity  is  the  result  of  imita- 
tion, while  adaptation  is  the  result  of  invention 
— i.e.  of  all  improvement  and  innovation  tending 
to  make  a  new  society  different  from  that  which 
preceded  it.^ 

These  fundamental  analogies  suffice  in  themselves 
to  justify  our  collaboration,  whatever  may  be  the 
solution  of  the  question — which  is  really  only  a 
question  of  terms — as  to  whether  societies  should 
be  regarded  as  organisms,  or  organisms  as  particular 
kinds  of  societies.^ 

1  Invention  is  frequently  a  combination  of  several  imitations. 
When  a  society  is  formed,  its  characters  are  not  necessarily 
borrowed  from  those  societies  from  which  it  more  or  less  directly 
proceeds.  It  may  be  modelled  upon  other  social  structures  with 
which  it  had  no  hereditary  connection. — V.  Tarde,  Les  Lois  dc 
V  imitation. 

^See  Les  Soci4tds  animaleSf  p.  128  (Espinas).  "Integration, 
or  grouping  together,  is  a  universal  law  common  to  all  organic 
or  inorganic  existence.  Society,  properly  speaking,  is  only  a 
complex  and  important  instance  of  this  universal  law." 

See  La  Science  Sociale,  p.  97  (Fouillee).     **  All  purely  physio- 


INTRODUCTION  ^9 

In  either  case,  it  is  certain  that  organisms  and 
societies — used  in  the  sociological  acceptation  of 
the  term — exhibit  some  characters  in  common,  and 
some  distinctive  characters. 

The  common  characters  may  be  accounted  for 
by  the  co-operation  which  exists  in  both  cases 
between  the  units  of  which  they  are  composed 
(individuals,  or  cells). 

The  dissimilar  characters  are  probably  connected 
with  what .  constitutes  the  essential  difference  be- 
tween social  aggregates  and  organic  aggregates. 
With  the  former  there  is  a  physiological  continuity 
between  the  composing  units,  while  with  the  latter 
co-operation  is  entirely  due  to  mental  relations. 

II.  Individuals,  colonies  and  societies. — Our  view 
in  this  matter  is  considerably  at  variance  with  the 
current  opinion.  Many  authors,  and  M.  Espinas 
among  them,  regard  colonies,  whether  animal  or  vege- 
table, as  societies,  even  when  the  members  of  these 
colonies  are  connected  by  physiological  bonds.^ 

logical  characters  of  life,  viz.:  1.  Correlation  of  parts  ;  2.  Relation 
between  structure  and  function  ;  3.  Division  of  all  living  parts 
into  other  living  parts  ;  4.  Spontaneity  of  movement ;  5.  Partic- 
ulate existence  ;  6.  Development  and  degeneration,  1  revolution, 
are  to  be  found  in  a  greater  extent  in  animal  and  human  societies." 
^  See  Les  Societes  animales,  section  2,  pp.  207  and  foil. — 
(Espinas),  Cours  de  philosophie  positive,  vol.  iv.  (A  Comte). 
**  These  strange  societies  are  to  be  found  among  the  lower 
animals,  an  involuntary  co-operation  being  the  result  of  an  un- 
severable  organic  union,  which  is  either  a  mere  adherence  or 
actual  continuity." 


10  EVOLUTION  BY  ATROPHY 

We  shall  justify  the  distinction  we  make  by 
showing  that  colonies  are  entirely  distinct  from 
societies.  The  two  are  divergent  branches  which 
spring  from  the  same  source,  a  solitary  indi- 
vidual. These  divergent  branches  of  organic  life 
may  be  distinguished  in  the  following  manner : 
An  individual  may  be  either  unicellular  or  mul- 
ticellular. Every  cell  capable  of  sustaining  life 
and   reproducing   its   own   kind  is  an  individual.^ 

In  the  case  of  multicellular  individuals  there  is 
an  unbroken  physiological  continuity,  while  life 
lasts,  between  the  cells  of  which  they  are  com- 
posed. All  these  units  spring  from  one  primary 
cell  (i.e.  a  fertilized  egg)  which  sprang  itself  from 
two  cells  (male  and  female),  proceeding  usually 
from  two  separate  individuals.  From  the  moment 
that  this  union  is  effected,  the  continuity  remains 
unbroken.^     When  such  an  individual  reproduces 

^  We  regard  the  cell  as  the  unit  of  life,  although,  according  to 
Altmann,  the  unit  of  life  is  a  still  simpler  structure — the  bioblast. 
He  maintains  that  the  bioblast,  or  living  granule,  is  the  ultimate 
element  of  the  cell ;  it  is  born  from  a  pre-existing  granule  ;  it 
lives,  feeds,  reproduces,  and  is,  in  fact,  an  organism  (see  Die 
Elementarorganismcn  und  ihre  Beziehungen  zu  den  Zellen ; 
Leipzig,  1890. — Studien  Uber  die  Zellen;  Leipzig,  1886. — Zur 
Geschichte  'der  Zelltheorien ;  Leipzig,  1889  (Altmann).  The 
existence,  however,  of  the  bioblast  is  a  pure  hypothesis,  no 
definite  proof  having  been  established  of  its  existence  ;  and  to 
assume  that  it  is  the  simplest  unit  of  life  is  to  abandon  oneself 
to  pure  imagination. 

^  Although  this  theory  is  in  agreement  with  the  observed  facts 
of  biology,  it  is  necessary  to  point  out  the  complexity  of  the 
ideas  of  individual  and  organism.     Boveri  {Uher  die  BefrucMungs 


INTRODUCTION  ll- 

asexually,  giving  rise  to  offspring  which  remain 
attached  to  it,  the  whole  forms  a  colony.  All 
the  cells  of  this  colony  spring  then  from  one 
single  cell  (a  fertilized  egg  cell),  and  an  inter- 
change of  nutriment  goes  on  incessantly  among 
these  cells.  This  is  what  Spencer  designates  as 
a  "  physiological  contract."  Societies,  on  the  other 
hand,  consist  of  units  springing  from  various  sources, 
whose  connection  is  merely  mental.  It  is  only  in 
the  communities  which  Tonnies  calls  "  Gemein- 
schaften"  in  distinction  from  societies  bound  by 
mental  agreement  {Gesellscliaften),  that  all  the  in- 
dividuals spring  from  one  and  the  same  couple. 

From  our  point  of  view,  a  colony  must  not  be 
regarded  as  an  intermediate  condition  between  an 
individual  and  a  society.  No  known  society  has 
passed  through  a  colonial  stage,  and  the  members  of 
a  colony  could  not,  on  separating,  become  a  society. 
Among  the  simplest  aggregates,  all  the  units  are 

und  EntwicklungsfdhigkeU  kernloser  Seeigel-Eier,  und  iiher  the 
Moglichkeit  ihrer  Bastardirung.  Arch.  /.  Eiriwicklungsmechanik 
der  Organismen,  Bd.  ii.,  pp.  394-443),  for  instance,  has  shown 
that  it  is  possible  to  fertilize  the  eggs  of  Echiuoderms,  from 
which  the  nuclei  have  been  removed,  by  the  spermatozoa  of 
other  species  of  Echinoderms.  The  egg  is  a  typical  cell,  an 
organism,  an  individual.  All  its  parts  are  essential  to  it,  and 
arc  incapable  of  separate  existence,  at  any  rate  for  any  length 
of  time  ;  close  physiological  bonds  unite  the  component  parts, 
yet  it  is  possible  to  substitute  for  an  essential  part  of  one 
individual  a  part  taken  from  another  individual.  It  may  there- 
fore be  concluded  that  the  idea  of  an  organism  no  longer  necessarily 
implies  the  idea  of  continuous  functional  unity  Avhich  one  was 
formerly  tempted  to  ascribe  to  it. 


12  EVOLUTION  BY  ATROPHY 

equivalent,  and  each  retains  all  the  functions  of 
life  {Spirogyra,  Choanoflagellates,  Hydra).  The 
colony  may  disintegrate,  but  the  cells  or  indi- 
viduals set  at  liberty  are  capable  of  living  alone, 
and  do  not  form  a  society.^ 

As  the  organisation  of  a  colony  becomes  more 
complicated,  the  units  proceed  to  differentiate.  Each 
one  assumes  certain  functions  in  particular,  and 
becomes  less  adapted  to  discharge  others.  If  such 
a  colony  is  dispersed,  the  individuals  are  incapable 
of  maintaining  a  separate  existence.^ 

^  (a)  A  thread  of  Spirocpjra  consists  of  cells  placed  eud  to  end, 
physiologically  connected  with  one  another.  Under  certain 
conditions,  however,  quite  apart  from  the  phenomenon  of  cellular 
reproduction,  all  these  cells  are  capable  of  isolating  themselves. 
The  colony  is  thus  transformed  into  a  set  of  individuals  capable 
of  maintaining  a  separate  existence,  and  no  longer  in  connection 
with  one  another. 

[h)  Among  the  Choanoflagellates,  some  consist  of  free  individuals, 
while  in  others  all  the  individuals  are  united  by  a  common  stalk, 
and  intercommunicate  by  protoplasmic  threads  in  the  stalk.  When, 
for  some  cause,  these  individuals  separate,  they  never  form  them- 
selves into  a  society. 

(c)  A  colony  of  Hydra  is  formed  by  the  budding  of  a  single  indi- 
vidual. While  nourishment  is  abundant,  all  the  individuals  of  the 
colony  retain  their  connection  with  each  other,  and  may  themselves 
give  rise  to  buds.  On  the  other  hand,  when  food  is  scarce,  the 
colony  disintegi-ates  and  each  individual  lives  a  free  life  without 
entering  into  social  relations  with  its  neighbours. 

^  In  some  cases  the  terms  individual  and  colony  become  ex- 
tremely involved.  Although  one  accept  our  view,  it  is  difficult  to 
rigorously  apply  our  generalizations  to  such,  facts  as  the  following  : 
Certain  male  cephalopods,  at  the  period  of  reproduction,  separate 
from  their  bodies  a  specialized  tentacle  (hectocotyle)  in  which  is 
stored  the  seminal  fluid.     This  organ,  set  at  liberty,  swims  in  the 


INTRODUCTION  13 

III.  Communities  and  societies. — The  bond  ex- 
isting between  all  the  constituent  parts  of  a  society- 
is  not  of  the  same  nature  as  that  which  unites  the 
members  of  a  colony  or  the  cells  of  an  animal  or 
plant.     These  physiological  bonds  are  not  compar- 

sea,  and  after  living  there  a  certain  time,  enters  the  female  oepha- 
lopod,  to  effect  impregnation. 

The  individuality  of  this  hectocotyle  appears  so  obvious  that 
for  a  long  time  it  was  regarded  as  a  distinct  species— some  kind 
of  worm.  At  first  sight,  then,  the  hectocotyle  seems  to  be  an 
individual.  This,  however,  is  not  the  case,  as  it  reproduces,  not 
a  hectocotyle  but  a  cephalopod. 

Many  Echinoderms,  by  a  spontaneous  act  of  J)rotection,  can 
separate  their  arms  from  their  bodies.  Such  a  separated  arm 
may  live,  feed  and  slowly  build  up  again  a  whole  Echinoderm. 
Plainly,  it  would  be  impossible  to  indicate  the  precise  point  at 
which  such  an  organ  should  be  regarded  as  an  individual.  The 
vegetable  kingdom  abounds  in  analogous  facts.  A  strawberry 
plant,  for  instance,  gives  off  runners  in  the  course  of  the  Summer, 
which  take  root,  and  themselves  become  sti*awberry  plants.  So 
long  as  these  young  plants  are  insufficiently  developed  to  main- 
tain themselves,  the  mother-plant  continues  to  supply  them  with 
nourishment.  As  soon,  however,  as  the  young  shoot  can  dispense 
with  this  support,  the  runner  atrophies,  and  the  little  plant  begins 
a  separate  existence.  By  prematurely  cutting  the  runner,  the 
new  plant  may  be  compelled  to  live  alone  sooner  than  it  naturally 
would  have  done. 

With  some  other  plants  (Phalangium  viviparum)  the  young 
shoots  frequently  retain  their  connection  with  the  mother-plant, 
although  quite  capable  of  maintaining  themselves.  Under  these 
circumstances  it  is  obviously  impossible  to  say  if  these  plants 
represent  colonies  or  free  individuals.  Speaking  generally,  it 
may  be  said  that  no  precise  line  can  be  drawn  between  colonies 
and  individuals.  Many  writers  on  the  subject,  and  Perrier  among 
the  number,  consider  that  every  colony  where  there  is  a  physical 
continuity  among  the  members,  should  be  regarded  as  an  in- 
dividual, 


14  EVOLUTION  BY  ATROPHY 

able,  without  forcing  the  analogy,  although  they  have 
been  so  compared  by  some  sociologists,  to  such  means 
of  communication  between  individuals  and  societies 
as  exchange,  traffic,  roads,  railways,  telegrams  and 
telephones. '  It  is  merely  a  matter  of  definition,  and 
if  societies  are  to  be  termed  organisms,  they  should 
be  distinguished  as  organisms  by  social  contract, 
organismes  contractuels  (Fouill^e). 

This  definition,  however,  only  applies  to  those 
societies  which  owe  their  existence  to  a  formal 
contract  with  definite  objects  in  view,  and  not  to 
ready-made  communities  consisting  of  individuals 
already  united  together  without  any  preliminary 
contract.  The  latter  is  the  case,  for  instance,  in 
societies  of  ants  or  bees,  and  in  human  societies  in 
those  social  groups  in  which  the  individuals  are 
united  by  the  bonds  of  consanguinity.  The  char- 
acters of  such  communities  partly  approach  the 
characters  of  organic  associations,  but  precisely  as 
such  natural  communities  approach  societies  by 
social  contract,  the  differences  between  social  groups 
and  actual  organisms  become  more  marked.  In  the 
more  complex  forms  of  societies  the  results  of  the 
characters  we  have  distinguished  become  most  ac- 
centuated. 

IV.  Distinctive  characters  of  societies  of  which 
the  memhers  are  united  hy  social  contract. — (1)  A 
cell  cannot  be  part  of  two  organisms  or  of  two 


INTRODUCTION  15 

organs  at  the  same  time.'  On  the  other  hand, 
there  is  no  reason  why  the  members  of  one  social 
community  should  not  belong  to  other  communities 
at  the  same  time. 

2.  Speaking  generally,  the  biological  conception 
of  an  organism  denotes  a  definite  thing — a  plant  or 
an  animal  in  itself,  quite  distinct  from  similar 
organisms.  In  sociology,  however,  there  is  no 
precise  line  between  co-existing  social  communities. 
Are  we  to  regard,  for  instance,  the  families,  com- 
munes and  cantons  of  a  state  as  distinct  organisms, 
or  merely  as  organs  ?  Does  a  free  town  such  as 
Hamburg  or  Frankfort  cease  to  be  an  organism 
when  it  loses  its  independence  ?  Take  the  various 
Swiss  cantons,  which  are  now  mere  organs  of 
the  Helvetic  Confederation,  like  the  Provinces 
of  Belgium,  or  the  Departments  of  France,  would 
they  become  organisms  on  the  rupture  of  the 
Federal  bond  ?  On  the  other  hand,  with  the 
growth  of  international  treaties  between  the 
states  of  Eastern  Europe,  will  their  social  indi- 
viduality disappear,  and  will  they  come  to  be 
regarded  as  are  the  United  States  of  America, 
as  the  organs  of  an  organism  in  process  of 
formation  ?      These  few  examples  suffice  to  show 

1  When  two  organs  are  united  into  one  whole,  the  cell  exercises 
two  totally  different  functions.  The  liver,  as  we  now  know,  is  a 
double  organ  consisting  of  a  bile-forming  liver,  and  the  glycogen- 
producing  liver,  two  organs  which  are  embryologically  distinct. 
The  cells  of  which  the  liver  is  composed  are  both  bile-secreting 
and  glycogen-forming  organs, 


1  6  EVOLUTION  BY  ATROPHY 

that,  SO  far  as  social  matters  are  concerned,  the 
conception  of  organisms  is  a  pure  convention. 
In  the  course  of  this  treatise,  we  may  therefore 
regard  families,  societies  and  nations  as  distinct 
organisms,  or,  with  regard  to  their  connection  with 
other  and  vaster  organizations,  as  organs  of  the 
latter. 

3.  The  structure  of  an  animal  or  plant  depends 
upon  the  physical  arrangements  of  its  parts,  and  on 
the  physiological  links  between  those  parts.  The 
structure  of  a  society  depends  upon  the  links  of 
social  contract  existing  among  its  members.  We 
regard  these  as  two  very  different  things,  and  we 
cannot  follow  Tarde  in  pressing  the  analogy  between 
them  in  the  following  way :  "  The  length,  breadth, 
and  height  of  an  organism  are  never  very  much  out 
of  proportion.  With  snakes  and  poplars  the  height 
or  length  preponderates ;  among  flat  fish  the  thick- 
ness is  very  small  compared  to  the  other  dimensions, 
but  in  each  instance  the  disproportion  exhibited  in 
extreme  cases  is  not  comparable  to  that  shown  by 
any  social  aggregate — such  as  China  for  instance, 
which  is  3000  kilometres  in  length  and  breadth, 
and  only  one  or  two  yards  in  average  height,  for 
the  Chinese  being  a  short  race,  build  their  edifices 
correspondingly  low."  ^ 

4.  Organic  modifications  are  effected  more  slowly 
and  with  greater  difficulty  than  are  social  modifica- 

^  Lcs  Monades  et  la  Science  sodale.    (Tarde.    Eemie  de  Sodologie, 
1893,  p.  169.) 


iNTKODUCTiON  1 7 

tions.^     The  result  is  an  important  one  from  the 
point  of  view  of  method. 

In  biology,  excepting  in  the  case  of  individual 
adaptation  of  artificial  selection,  direct  observation 
— the  historical  method,  if  we  may  so  call  it — is 
not  available  for  the  study  of  the  origin  and 
modification  of  organisms.  Phylogeny,  the  science 
of  organic  kinship,  resorts  to  other  methods,  and 
particularly  to  the  comparative  method  in  its 
various  forms : — 

(a)  Morphology,  the  science  of  determining  the 
phylogeny  of  organs  by  comparing  them  with  the 
organs  of  other  creatures  belonging  to  the  same 
systematic  group. 

(b)  Palaeontology,  which  determines  the  direct 
ancestors  of  living  creatures. 

(c)  Embryology,  which,  so  far  as  it  is  founded 
upon  the  principle  of  recapitulation,  investigates 
the  development  of  organs  in  the  individual,  and 
draws  conclusions  therefrom  bearing  upon  its 
descent. 

*  Among  animals  there  is  a  special  factor  which  gives  a,stability 
to  the  specific  characters  not  found  elsewhere — this  is  reproduction. 
Specific  characters  being  common  to  the  whole  line  of  descent,  arc 
very  deeply  enrooted  in  the  organism.  They  are  not  easily  modi- 
fied by  the  influence  of  new  environments,  but  maintain  their 
likeness  to  one  another  in  spite  of  external  conditions.  They  are 
regulated  by  an  internal  force,  notwithstanding  the  importunities 
to  variation  off'ered  from  outside.  This  force  is  heredity,  and 
heredity  accounts  for  the  precise  way  in  which  specific  characters 
may  be  defined.  In  society  this  internal  force  is  wanting.  {Lcs 
Regies  de  la  metlwde  sociologique.  Durckheim,  Paris,  F.  Alcan ,  1 895 . ) 

B 


18  EVOLUTION  BY  ATROPHY 

{d)  Teratology,  which  compares  normal  with 
abnormal  forms  for  the  same  purpose. 

In  sociology  these  various  systems  of  comparison 
are  only  of  a  secondary  value,  owing  to  the  great 
variability  of  social  forms,  while  the  historical 
method  of  investigation  assumes  a  greater  impor- 
tance. In  sociology,  however,  there  are  methods 
analogous  to  the  comparative  methods  of  biology : 

{a)  Archaeology  corresponds  to  palaeontology. 

(Jb)  Social  morphology,  by  comparison  of  series 
of  institutions,  makes  up  to  a  certain  extent  for 
the  absence  of  direct  observation  of  their  origin 
and  development.  Thus,  to  use  the  phrase  of 
Bagehot,  by  studying  the  customs  and  institu- 
tions of  modern  savages,  the  prehistoric  living 
may  be  made  to  throw  light  upon  the  pre- 
historic dead.  If  an  institution  be  found  in  full 
operation  among  savages,  of  which  a  vestige  still 
exists  among  more  civilized  people,  it  may  be 
assumed  that  the  vestige  was  at  some  time  fully 
functional  among  the  latter.  It  must  be  borne 
in  mind,  however,  that  in  many  instances  such 
vestiges  are  the  result  of  imitation.  Theodore 
Reinach  has  shown  that  this  applies  in  the  case 
of  circumcision.^ 

This  gravely  weakens  the  conclusions  drawn  by 
Spencer  from  the  survival  of  this  custom  among 

^  De  quelqucs  fails  relatifs  d  I'Mstoire  de  la  circoncision  cJiez  Ics 
peuplcs  dc  la  Syrie  (Th.  Reinach,  V Anthropologies  1893,  vol.  iv., 
pp.  28  and  following). 


INTKODUCTION  19 

certain  Australian  tribes.  Granting,  he  says,  that 
circumcision,  the  removal  of  teeth,  and  other 
similar  mutilations  imply  a  condition  of  political 
or  religious  subjection — or  both — no  longer  exist- 
ing among  these  tribes,  the  custom  is  obviously  the 
vestige  of  a  more  complex  social  condition.  This 
conclusion  seems  the  less  reliable,  since,  according 
to  the  Eev.  J.  Matthew,  the  rite  of  circumcision 
was  probably  introduced  into  Australia  by  natives 
of  Sumatra,  and  this  view  is  confirmed  by  the 
local  distribution  of  the  custom  and  by  other  evi- 
dences of  the  same  origin,  such  as  the  paintings 
which  have  been  discovered  in  certain  caves.^ 

(c)  Teratology  and  social  embryology  also  play  a 
part  in  sociology,  but  it  is  of  less  importance  than 
the  others. 

Certain  customs  among  criminals  show  a  re- 
semblance to  the  habits  of  primitive  man.  On 
the  other  hand,  we  find  cases  where  the  individual 
development  of  an  institution  or  society  is  a  mere 
repetition  of  the  development  through  which  similar 
institutions  and  societies  of  other  epochs  and  places 
have  passed.  Thus,  for  instance,  there  still  exists 
in  some  parts  of  modern  Russia  a  voluntary  agri- 
cultural commune,  for  the  periodical  division  of  the 
land,  an  institution  which  existed  more  universally 
in  the  sixteenth  and  seventeenth  centuries  during 

^  The  Cave  Paintings  of  Australia  {Journal  of  the  Anthropo- 
logical Institute  of  Great  Britain  and  Ireland,  April  1893,  pp.  51 
and  following). 


20  EVOLUTION  BY  ATROPHY 

the  existence  and  after  the  disappearance  of  family 
communities.^ 

In  studying  the  development  of  these  new  com- 
munities, one  can,  in  a  measure,  picture  to  oneself 
the  development  of  laws  of  property,  which  were 
current  in  other  countries  at  other  times,  and  of 
which  we  possess  little  or  no  direct  information. 
It  is,  however,  hardly  necessary  to  insist  upon  the 
hypothetical  quality  of  such  conclusions. 

Our  methods  show,  then,  that  organisms  and 
societies  exhibit  considerable  differences  as  well 
as  analogies,  a  necessary  result  of  their  different 
natures.  These  few  remarks  must  suffice ;  to  add 
to  them,  we  should  have  to  overstep  the  limits  we 
have  set  to  this  treatise,  enter  into  well-worn  con- 
troversies, and  anticipate  our  own  conclusions. 
Having  merely  explained  our  terminology,  and 
indicated  our  general  views,  we  will  proceed  to 
the  subject  of  our  investigations. 

1  Tableau  des  orighus  et  dc  revolution  de  la  propridi  et  de  la 
famille,  p.  170  (Kowalevsky). 


BOOK  I 

UNIVERSALITY  OF  DEGENERATIVE 

EVOLUTION 

PART  I 

DEGENEKATION  IN  THE  DEVELOPMENT  OF  INSTITUTIONS 
AND  ORGANS 

The  term  "  Evolution "  does  not  in  itself  convey 
an  idea  of  either  progress  or  degeneration.  It 
comprises  all  the  changes  undergone  by  an 
organism  or  society  independently  of  the  ques- 
tion as  to  whether  these  changes  are  favourable 
or  otherwise.  The  evolution  of  an  organ,  or  of 
the  different  parts  of  an  organ,  is  degenerative  if 
it  tends  to  the  ultimate  decay  of  that  organ  or  of 
its  parts,  and  the  facts  are  shewn  by  means  of 
arranging  series  of  fossils  or  living  forms,  and  com- 
paring them.  Evolution  is  progressive  if  it  tends 
to  the  development  of  an  organ  or  to  the  formation 
of  a  new  organ. 

These  definitions  may  be  applied  —  mutatis 
mutandis — to  the  changes  undergone  by  societies 
and  institutions  or  their  constituent  parts. 

The  ideas  of  progress  and  of  degeneration  seem 
at  first  sight  to  relate  to  diametrically  opposite 
phenomena.       The    term    "  progressive    evolution " 


22        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

conveys  the  ideas  of  progress,  development,  im- 
provement, of  increasing  differentiation,  and  of  the 
progressive  co-ordination  of  the  functions  or  organs 
thus  dififerentiated. 

Degenerative  evolution,  on  the  other  hand,  con- 
veys the  ideas  of  decline,  of  decay,  and  of  de- 
generation, such  as  the  atrophy  of  the  organs  of 
locomotion  in  Sacculina,  the  degeneration  of  the 
leaves  of  parasitic  plants,  or  the  dissolution  of 
corporate  bodies  in  a  declining  state. 

We  propose  to  show,  however,  that  these  two 
ideas,  which  at  first  sight  seem  contradictory  and 
mutually  exclusive,  are  found,  on  a  strict  examina- 
tion of  the  facts,  to  interpenetrate  and  complete 
each  other.  Degeneration  and  progress  will  appear 
as  the  two  sides  of  one  whole,  or  as  two  aspects  of 
the  same  evolution,  and  it  will  be  seen  that  all 
progress  must  necessarily  be  attended  by  degenera- 
tion. 


CHAPTER  I 

IN  THE  EVOLUTION  OF  ORGANS  ALL  MODIFICATION  IS 
NECESSARILY  ATTENDED  BY  DEGENERATION 

Section  I. — Preliminary  Considerations 

In  order  to  effect  the  demonstration  which  is  the 
object  of  this  chapter,  only  the  phylogenetic  modifi- 
cations  of   organs  will  be  discussed,  setting  aside 


EVOLUTION  AND  DEGENEEATION  OF  ORGANS        23 

the  original  formation  of  these  organs  in  their 
ancestral  forms,  their  development  in  the  indivi- 
dual, the  phylogenetic  evolution  of  function,  and  the 
great  variation  resulting  from  individual  adaptation. 
Before  entering  upon  our  immediate  subject,  it 
would  be  as  well  to  define  its  limits  by  saying 
a  few  words  concerning  the  questions  not  under 
immediate  consideration. 

1.  Original  formation  of  orgatis  in  ancestral 
forms. — Little  is  known  concerning  the  primary 
origin  of  organs,  and  their  development  before  their 
assumption  of  the  aspect  and  function  by  which 
we  know  them.  What  were  the  rudimentary 
leaves  like  in  the  ancestors  of  flowering  plants? 
What  were  the  eyes  of  the  first  vertebrates  like  ? 
Did  these  organs  develop  from  existing  organs  ful- 
filling other  functions,  or  were  they  formed  inde- 
pendently ?  However  that  may  be,  if  they  arose 
from  other  organs,  we  know  nothing  of  the  modifi- 
cations which  they  underwent ;  and  if  they  were 
formed  independently,  we  need  not  discuss  the 
fact  here,  for  in  that  case  the  organ  would  not 
have  developed  by  transformation.  Having  once 
been  formed,  it  would  develop  and  improve,  and 
this  process  would  not  necessarily  have  been 
attended  by  partial  degeneration. 

2.  Development  of  organs  in  the  individual. — In 
the  course  of  embryological  development,  organs 
do     not    exactly     repeat    the    successive    phases 


24       UNIVEKSALITY  OF  DEGENERATIVE  EVOLUTION 


through  which  they  passed  during  their  ancestral 
evohition.  Generally  speaking,  and  especially  in 
the  case  of  plants,  the  development  of  organs  in 
an  individual  is  direct,  and  gives  no  clue  to  its 
ancestral  history.  Moreover,  when  there  is  a  re- 
capitulation of  ancestral  stages,  it  often  happens 
that  evolution  takes  place  without  leaving  traces 

of  the  various 
stages.  This  is 
especially  the 
case  in  complex 
organs  which 
have  been  pro- 
duced by  many 
lines  of  evolu- 
tion converging 
in  a  single  struc- 
ture—  a  struc- 
ture which  thus 
becomes  the  seat 

Fig.  1.— DiaRiam  showing  the  evolution  of  pyramidal  ^f  „  Qr»pf>iQl  fnnp- 
cells  in  the  animal  series.  "^  "^  bpecidl  lUIlO 

The  upper  series  of  cells  represents  the  psychic  cell  in  tiou      Or      SCt     of 
various  vertebrates:   A,  the  frog;    5,  the  lizard; 
C,  the  rat;  D,  man.    Tlie  lower  series  shows  the  fuUCtlOUS. 
progressive  stages  in  the  evolution  of  the  pyramidal 

cell  in  the  human  brain  :  a,  the  neuroblast  without  lllC       UCUrOn, 

protoplasmic  processes;    6,  the  appearance  of  the  .  . 

nerve  process  and  of   the  terminal  ramifications ;  for  iustaUCC,  the 
c,  the  nerve  more  fully  developed ;  d,  appearance  of  t       • 

lateral  branches  of  the  axis  cylinder ;  e,  development  ganojllOniC        CCll 
of  protoplasmic  outgrowths  of   the  protoplasm  of  ° 

nerve-cell  and  nerve.    (Ramon  y  Cajal.)  of    the   COrtCX  01 

the  human  brain,  passes  successively  through  stages 
corresponding  to  those  which  are  to  be  found  in  the 
adult  fish,  frog,  bird,  and  mammal.     In  this  case 


EVOLUTION  AND  DEGENERATION  OF  ORGANS       25 

the  development  consists  in  an  increasing  com- 
plexity of  the  cell  with  no  formation  of  unnecessary- 
rudimentary  parts. 

3.  Phylogenetic  evolution  of  function. — Evolution 
may  be  regarded  from  a  physiological,  as  well  as 
from  an  anatomical  standpoint,^  but,  in  the  former 
case,  evolution  is  less  a  set  of  changes  of  function 
than  an  increasing  specialization  and  division  of 
labour,  and  under  these  circumstances  it  is  often  diffi- 
cult to  recognize  a  degenerative  element  in  the  evolu- 
tion.    A  few  examples  will  demonstrate  this  point : 

Self-mutilation  is  a  very  common  phenomenon 
among  Echinoderms.  Among  brittle  star-fish  this 
reaction  is  controlled  by  some  region  of  the  nervous 
system ;  in  some  star-fish  the  reaction  follows  more 
quickly  because  the  stimulus  can  act  upon  the 
ganglion  at  the  root  of  each  arm  near  the  circum- 
oral  nerve-collar.  In  Asteracanthum  riihen^,  there 
is  a  complete  localization  of  this  function,  and  self- 
mutilization  only  results  when  an  exact  region  of 
the  nervous  system  is  stimulated.^  In  the  Medusa 
we  find  an  equally  interesting  example  of  functional 
evolution  (Romanes).-'^     With   some  of   these   (the 

*  Evohdion  fcmctionnelle  du  sysUme  nerveux.  J.  Demoor,  Revue 
universitaire,  Bmxelles,  1892. 

2  Contribution  d  la  physiologie  nerveuse  des  Echinodermes.  J. 
Demoor  and  M.  Chapeaux.  Tijds.  Ned.  DierTc.  Vereen.  (2)  III. 
2  Nov.  1891. 

^  Preliminary  Observations  an  the  Locomotor  System  of  Medusa  ; 
Jelly-fish,  Star-fish,  and  Sea-urchins.  Romanes,  Int.  Scient.  Series, 
1885, 


26        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

Acraspedotc,  on  the  outside  of  the  umbrella  being 
separated  from  the  central  part,  the  two  separate 
parts  continue  to  lash  the  water,  the  outer  part 
with  even  strokes,  the  central  mass  more  slowly 
and  feebly.  With  the  Graspedote,  on  the  other 
hand,  the  central  part,  under  the  same  conditions 
seems  quite  paralyzed  and  immovable,  while  the 
outer  part  continues  to  move  in  a  perfectly  normal 
manner.  The  causative  function  of  the  movement, 
the  spontaneity  of  the  movement  as  it  was  formerly 
called  in  physiology,  is  incompletely  specialized  in 
the  Acraspedote,  whereas  in  the  Crasrpedote  it  is 
entirely  localized. 

Individual  adaptation. — The  individual  is  by 
no  means  a  slave  to  heredity.  It  is  capable  of 
certain  modifications  under  the  influence  of  certain 
external  conditions.  These  phenomena  of  individual 
adaptation  may  be  arranged  in  three  groups. 

(a)  When  an  organism,  either  animal  or  vegetable, 
is  placed  under  new  conditions  of  existence,  when 
for  instance,  it  relinquishes  a  terrestrial  for  an 
aquatic  life,  light  for  darkness,  or  fresh  water  for 
salt  or  esturine  water,  its  external  aspect,  and  in- 
ternal structure,  undergo  variations  of  considerable 
importance  if  it  succeeds  in  adapting  itself  to  the 
new  conditions.^ 

^  Examples  :  {a)  The  leaves  of  the  water  Ranunculus  with 
lacinated  leaves  (llaniLnculus  aquatilis  fluitans,  etc.),  are  of  normal 
structure  when  cultivated  on  dry  land.  The  epidermis  is  furnished 
with  stomata  and  the  constituent  cells  contain  no  chlorophyll. 


EVOLUTION  AND  DEGENERATION  OF  ORGANS       27 

The  organ  does  not,  however,  lose  its  primitive 
and  typical  characters.  Actual  organic  transforma- 
tion cannot,  therefore,  be  said  to  take  place  in  the 
case  of  individual  adaptation. 

The  same  leaves  of  the  same  plant  when  grown  in  water  are 
much  longer  than  those  of  the  terrestrial  type  ;  the  leaves  have  no 
stomata,  and  the  epidermic  cells  are  full  of  chlorophyll  (Askenasy, 
Ueher  den  Einfiuss  des  Waclistumsmediums  auf  die  Gcstalt  der 
Pflanzen,  Bot.  Zeit.,  1870,  pp.  193  and  following).  Among  the 
Stratiotes  aloides  it  is  not  uncommon  for  the  upper  part  of  a  leaf  to 
rise  above  the  surface  of  the  water  while  the  base  is  submerged  ;  the 
epidermis  at  the  base  contains  chlorophyll,  but  has  no  stomata, 
while  the  part  of  the  same  leaf  which  rises  out  of  the  water  is 
furnished  with  stomata,  but  has  no  chlorophyll  in  the  epidermis. 

(6)  A  good  example  of  this  individual  adaptation  may  be  obtained 
by  cultivating  Cacti  alternately  in  the  light  and  in  the  dark. 
Goebel  has  shown  that  when  a  specimen  of  Phyllocactus  is  culti- 
vated in  the  dark,  the  stems  are  prismatic  and  thorny ;  if  the  plant 
is  afterwards  placed  in  the  light,  the  thorns  disappear  and  the 
stems  become  quite  smooth.  (K.  Goebel,  Ueher  die  Einicirkung  des 
Lichtes  auf  die  GestaUung  der  Kacteen  und  anderer  PJlanzen,  Flora, 
vol.  80,  p.  96,  1895.) 

(c)  The  animal  kingdom  furnishes  numerous  examples  of  indi- 
vidual adaptation. 

The  gi-adual  drying  up  of  Lake  Aral  caused  the  formation  of  a 
number  of  basins  containing  water  at  various  stages  of  concentration. 
The  Cardium  of  this  region  exhibits  a  whole  series  of  adaptive  varia- 
tions. The  shells  become  thinner  and  horny,  their  shape  elongates, 
the  openings  contract,  and  their  colour  becomes  duller  {Bateson). 

Mytilus  edidis  (the  edible  mussel),  exhibits  three  different  kinds 
of  shells.  It  lives  either  in  salt  water,  deep  water,  or  shallow 
water  visited  by  the  tide.  In  each  of  these  three  vicinities  the 
shells  exhibit  typical  aspects. 

The  direction  of  boney  lamellae  is  known  to  agi-ee  with  that  in 
which  the  greatest  strain  is  habitually  applied,  and  the  entire 
structure  of  a  bone  is  dominated  by  the  incidences  of  the  forces 
applied  to  it.    When,  after  a  badly-mended  fracture,  the  two  broken 


28       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

(h)  Our  second  group  comprises  the  cases  of 
cellular  adaptation  which  may  be  produced  in  the 
course  of  embryonic  life  and  may  result  in  the  for- 
mation of  an  embryo  from  half  or  the  quarter  of 
an  egg} 

Here  again  the  formation  of  organs  takes  place 
under  new  and  abnormal  conditions,  but  there  is 
no  "  transformation "  of  these  organs  into  fresh 
structures  exhibiting  other  characters  than  those 
of  primitive  and  typical  organs. 

pieces  are  joined  by  an  oblique  segment,  the  trabeculse  in  this  seg- 
ment follow  the  direction  from  whence  comes  the  greatest  strain,  a 
different  direction  to  that  which  they  would  have  taken  had  the 
inserted  segment  been  placed  parallel  with  the  two  broken  pieces. 

Without  entering  upon  explanatory  theories  concerning  these 
facts,  it  is  important  to  notice  that  the  adaptations  of  plants  exhibit 
different  characters  to  those  of  animals. 

With  plants  it  is  only  when  a  young  organ  is  born  under  new 
conditions  that  it  exhibits  new  characters.  This  is  not  necessarily 
the  case  with  animals.  In  an  animal  organism  the  separation  of 
the  young  tissues  from  the  old  is  not  so  noticeable  as  with  plants, 
as  the  organs  undergo  a  continuous  renewal  and  can  always  adapt 
themselves  more  or  less  to  new  conditions. 

1  When  the  two  cells  resulting  from  the  first  division  of  a  fertilized 
egg  of  Amphioxus  are  artificially  separated,  each  cell  may  develop 
directly  into  a  complete  individual.  The  same  happens  even  whea 
the  first  four  cells  are  separated  from  each  other  artificially. 

When  this  is  effected  in  the  case  of  Echinus  or  of  an  Amphioxus 
embryo  of  eight  cells,  each  cell  developes  as  if  it  had  remained  a 
part  of  the  whole  ;  but  when  the  blastular  stage  is  reached,  that 
stage  slowly  completes  itself. 

In  the  first  case  the  cell  adapts  itself  to  the  new  conditions,  in 
the  second  case  it  is  the  blastular  which  does  so.  (See  Analytische 
Theorie  der  organischen  Entioickelung.  .  Hans  Driesch,  Leipzig, 
1894.) 


EVOLUTION  AND  DEGENERATION  OF  ORGANS   29 

In  an  adult  cell,  adaptation  may,  in  a  more  or 
less  normal  way,  cause  the  formation  of  new  organs. 

Even  completely  specialized  cells  of  an  organism 
may,  when  placed  under  certain  conditions  of  life, 
adapt  themselves  to  those  new  conditions  and  give 
rise  to  successive  generations  of  different  cells,  thus 
generating  other  kinds  of  organs  than  those  to 
which  they  would  have  given  birth  under  normal 
conditions.  This  happens,  for  instance,  with  all  the 
cells  of  Salix,  which  are  capable  of  reproducing  the 
whole  organism  by  means  of  budding.  This  organic 
plasticity  is  shared  with  the  Begonia  Rex,  but  in  that 
case  it  is  only  thoroughly  developed  in  the  epidermic 
cells.  Analogous  examples  abound  among  animals, 
especially  in  the  lower  zoological  groups.^ 

*  Loeb  has  shown  that  in  certain  groups,  when  the  organism, 
after  having  been  wounded,  is  subjected  to  unusual  light,  position, 
or  pressure,  organs  are  formed  at  the  wounded  part,  which  are 
essentially  different  from  those  which  would  have  been  formed 
in  a  normal  recovery.  When  this  heteromorphosis  occurs  in 
Tubularia  mesembri/anthenuim,  Aglaophenia  pluma,  Anthennu- 
laria  rosa,  etc.,  the  positions  of  tlie  oral  and  apical  poles  may 
be  transposed.  When  Antennularia  is  placed  so  that  gravity 
acts  upon  it  in  a  contrary  direction  to  the  ordinary  one,  there 
are  formed  oral  and  apical  branches,  where  they  would  not  have 
been  formed  had  the  organism  been  kept  in  a  normal  position. 
(Jacques  Loeb,  Unters.  zur  physiol.  Morplwlogie  der  Thiere :  L 
Ueher  Heteroinorplwse,  Wiirzburg,  1891  ;  IL  Organhildung  und 
Wcvchstlmm,  Wurzburg,  1892.) 

Similarly  there  is  a  true  heteromorphosis  when  in  a  case  of 
club-foot  the  faces  and  articular  surfaces  of  the  bones  and 
cartilages  assume  characters  adapted  to  the  new  work  thrown 
on  them  as  the  abnormality  of  the  joint  increases.  Another 
such  case  is  in  the  false  joints  sometimes  formed  when  the  two 


30       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

These  various  instances  of  cellular  adaptation 
called  "  heteromorphosis "  by  Loeb,  are  outside 
the  limits  of  our  present  researches,  for  reasons 
which  we  have  already  given. 

The  law  of  universal  degenerative  evolution 
applies  only  to  the  transformations  of  organs,  and 
not  to  their  original  formation  in  either  individuals 
or  species,  under  normal  or  abnormal  conditions. 

Section  I. 
Transformation  of  organs  of  animals. 

§  2.   Transformation  of  homodynamic  organs  in 
the  individual. 

In  order  to  study  the  degenerative  evolution 
exhibited  in  the  specific  modifications  of  the 
organs  of  an  individual,  it  is  necessary  to  choose 
homodynamic  organs  that  are  numerous,  and  conse- 
quently small  enough  to  undergo  different  kinds  of 
transformation  fitted  to  the  functions  they  may 
have  to  serve  in  the  different  parts  of  the  body. 

The  numerous  appendages  of  the  cray-fish  form 
an  extremely  interesting  study  of  this.  kind.     The 

parts  of  a  bone  do  not  unite  after  fracture.  In  that  case  a  com- 
pleted joint  with  cartilage  ligament  and  synovial  membrane  may 
be  formed  in  the  neighbouring  tissues  under  the  influence  of  the 
new  stimuli.  In  extra  uterine  gestation  the  placenta  is  formed 
upon  some  abdominal  organ.  When  this  occurs,  the  unusual 
stimulus  is  sufficient  to  cause  the  cells  of  an  abdominal  organ 
to  form  a  perfectly  specialized  organ  of  a  nature  foreign  to  their 
normal  life. 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS         31 

appendages  of  the  different  metameres  (the  somites, 
or  constituent  segments  of  the  body)  of  the  cray- 
fish are  constructed  on  the  same  plan.  Tlie 
typical  organ  (fig.  2),  when  complete,  consists  of 
three  parts — the  protopodite  (pr.), 
which  is  inserted  into  the  body  ^^^  _ 
and  carries  a  gill  (hr.) ;  the  endo- 
podite  (en.)  and  the  exopodite 
(ex.),  each  of  which  consists  of  a 
series  of  joints  which  are  attached 
to  the  end  of  the  protopodite 
— the  endopodite  to  the  inner 
side,  the  exopodite  to  the  outer. 

^  .  Fig.  2.— Diagram  of  typical 

Let        us        now       examine      the      and  complete  appendage 

of  Astacus  Jluviatilis  :  pr. 

twenty  metameres  of  the  animal    protopodite;    br,  gui; 

"'  ex,  exopodite;  en,  endo- 

in  succession.  podite. 

In  each  segment  of  the  body  we  find  a  pair  of 
appendages,  the  structure  of  which  is 
based  upon  that  of  a  complete  ap- 
pendage, but  in  which  many  different 
adaptations  have  brought  about  the 
conservation  and  increase  of  some  of 
the  typical  parts,  and  the  partial  or 
Fig.  3.  -  Astacus  total  atrophy  of  other  parts. 
^peS^oMhe  Take  first  the  abdominal  segments, 
segmenf'r'rhetliey  Carry  appendages  (fig.  3)  formed 
Ir™%topodite;of  the  three  fundamental  parts:  the 
Tn]  llTm^  protopodite  (pr.),  the  exopodite  (ex.),  and 
(Huxley).  ^^^   endopodite   (en.).      Each  of  these 

parts  is   itself    divided    into    different   joints,    the 


en — , 


32        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

description  of  which  is  not  of  immediate  impor- 
tance. The  portion  of  the  appendage  which  has 
degenerated  is  the  podobranch,  which  has  completely 
disappeared.  The  reason  is  obvious ;  these  ap- 
pendages, having  become  a  support  for  the  eggs, 
have  lost  their  respiratory  function.  The  part 
specially  adapted  for  that  function  has  been  allowed 
to  atrophy,  the  more  readily  because  the  presence 
of  gills  on  the  ventral  surface  of  the  abdomen 
would  be  incompatible  with  free  movement  of  that 

part  of  the  body  when  the 
cray-fish  is  swimming. 

The  appendages  of  the 
sixth  abdominal  segment 
are  greatly  modified  into 
a  caudal  fin  (fig.  4). 
This  also  exhibits  the  three 
fundamental    parts   of    a 

¥iG.  4.— Astacus  Jluviatilis.— Left  ap- 

pendage  of  the  fith  abdominal  seg-  COmpletC  appCndaffC.  ihc 

ment.    Fins  (1,5/1)  :  ^r,  protopodite;                                ~,              .  i  •    i 

ex,exopodite;  e»,endopodite(Huxley).^r0^02?Oat^e     [pr.)    IS  thlCK 

and  short  and  has  no  gill,  degeneration  being  ex- 
hibited in  the  loss  of  the  gill,  and  by  the  reduced 
length  of  the  part.  The  exopodite  (ex.),  and  the 
endopodite  (en.)  are  modified  into  two  large  oval 
plates  which  serve  as  propellers.  The  actual  de- 
velopment of  these  two  parts  is  accompanied  by 
degeneration.  In  a  typical  appendage,  the  exopodite 
and  the  endopodite  terminate  in  slender  parts  divided 
into  several  rings  by  false  joints ;  in  the  propellers, 
which  should  offer  the  maximum  resistance  to  the 


TRANTSFORMATION  OF  ORGANS  OF  ANIMALS         33 

pressure  of  the  water,  the  segmentation  has  dis- 
appeared, but  across  the  part  corresponding  to  the 
exopodite  there  still  remains  a  transverse  groove. 

The  second  abdominal  segment  of  the  female 
carries  appendages  similar  to  those  which  have 
just  been  described.  In  the  male, "  the  organs  of 
this  segment,  as  also  those  of  the  segment  in  front 
of  it,  have  become  organs  used  in  fertilization. 
We  must  consider  what  new  structures 
have  appeared  here,  and  to  what  extent 
these  new  modifications  have  been 
attended  by  degeneration.  The  ap- 
pendage of  the  second  segment  (fig.  5) 
is  longer  than  that  of  the  other  seg-  f  ^ 
ments ;  pressed  against  the  ventral 
surface  of  the  back  part  of  the  thorax, 
it  stretches  out  as  tar  as  the  space  be-  fluviatms.  Left 
tween  the  second  and  third  walking    2nd  abdominal  seg- 

_  -  .  1  •   1  1       J         ™6nt  of  the  male 

legs,  the  part   which  corresponds  to    (front view)(i,6/i): 

^1  '  ^      ^      £  ^x,      £  1  T4.  ^'      Protopodite; 

the  oviduct  of  the  female,  it  serves  as  ex,  exopodite ;  en, 
a  channel  to  conduct  the  fluid  from    roiied  piate  of  the 

_  1  -o  ^      ,^      D  ±^^        p  ^  endopodite;     b, 

the  male  orince  to  that  or  tne  lemale.    the   jointed   ex- 

-r  .  (.  J  T,         /         \       tremity     of      the 

It      consists     01      a     protopoaite      {pr.)      same  (Huxley). 

and  an  exopodite  (ex.)  similar  to  the  corresponding 
parts  of  the  appendages  we  have  just  described. 
The  endopodite  is  profoundly  modified.  The  inner 
border  of  its  proximal  region,  which  is  not  jointed, 
is  extended  into  a  thin  plate  rolled  into  a  hollow 
horn  (a)  while  the  outer  border  is  represented  by 
an   annulated    part   (h).     The   part   in   process   of 

C 


34       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

degeneration  is  the  exopodite.     On  considering  the 

development  of  the  various  parts  of  this  appendage, 

it   may   be  concluded   that  the  exopodite  has  not 

undergone  the  same  modification  as  the  endopodite. 

It   now   remains    to    conclude    this    study  of    the 

.  abdominal  appendages  by  an  ex- 

H'  amination    of    the    first   segment. 

Q-pf  Immense  variation  occurs  in  the 

female.     Sometimes  there  are  two 

Fig.  6.  —  Astacus  fluvia-  .  .       . 

tiiis.    Left  appendage  appendages,  somctimcs  one  or  both 

of    the    4th  abdominal  .      .  .  , 

segment  of  the  female    are     miSSlUff  I       lU     aUV     CaSC      thC 
(3/1) :  ^r,  protopodite ;  .      .  ^  '' 

c»,endopodite  (Huxley),  cxistiug  orgaus  are  vcry  small 
(tig.  6).  The  protopodite  (pr.)  is  small,  the  exopodite 
is  missing,  and  the  endopodite  (en.)  is  represented 
by  an  imperfectly  jointed  thread.  The  appendages 
of  this  first  segment  being  no  longer  necessary, 
atrophy,  and,  as  has  just  been 
pointed  out,  there  is  no  regularity 
or  uniformity  in  the  order  of  their 
disappearance.  In  the  male,  the 
appendages  of  this  segment  (fig.  7) 
possess  an  unjointed  rod  (t)  cor- 
responding to  the  endopodite  of  the    fw  7  _  Astacus  flwia- 

second  segment,  and  this  rod  Xe  K  ffiS 
stretches  out  to  a  considerable  aK" V  "nJoJ^n'tedTod 
distance.  The  exopodite  is  missing,  (^^u^iey)- 
and  the  articulations  between  the  different  parts 
have  disappeared.  This  may  reasonably  be  regarded 
as  the  result  of  degeneration,  and  confirms  what  has 
already  been  said  concerning  the  appendage  of  the 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS         35 

second  metamere,  where  degeneration  was  exhibited 
by  the  relatively  small  size  of  the  exopodite. 

It  may  be  concluded  from  this  examination  that 
in  the  abdominal  appendages  there  is  a  degenera- 
tion common  to  all — this  is  the  absence  of  the 
podohranch.  In  accordance  with  the  particular 
adaptations  of  each  pair  of  appendages,  special 
degeneration  accompanies  special  adaptive  develop- 
ments as  we  have  shown  above. 

We  now  pass  to  the  fourteen  anterior  metameres 
of  the  cray-tish.  The  first  six  of  these  constitute 
the  head,  and  carry  on  the  first  segment  the  stalked 
eyes ;  on  the  second,  the  antennules ;  on  the  third, 
the  antennae ;  on  the  fourth,  the  mandibles ;  on  the 
fifth  and  sixth,  the  two  pairs  of  maxillae.  The 
eight  metameres  of  the  middle  of  the  body  form 
the  thorax,  and  carry  on  the  seventh,  eighth,  and 
ninth,  the  maxillipedes ;  on  the  tenth,  the  claws ; 
and  on  the  eleventh,  twelfth,  thirteenth,  and  four- 
teenth, the  walking  legs. 

The  structure  of  all  these  appendages  may  be 
referred  to  that  of  the  typical  primitive  appendage. 
First  take  the  thoracic  appendages,  commencing 
with  the  third  maxillipede  (fig.  8)  which  is  the 
most  complete  appendage.  It  consists  of  a  proto- 
podite  formed  of  two  parts  (coxopodite  and  basipodite) 
and  carries  a  podohranch  (br.),  a  well-developed 
endopodite  (en.)  consisting  of  five  jointed  parts,  and 
of  a  small  exopodite  segmented  like  those  of  the 
abdominal  appendages. 

c  2 


36        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


The  third  maxillipede  may  be  regarded  as  the 
mean  type  of  the  average  thoracic  appendages. 
Compared  with  it  the  other  appendages  exhibit  a 
true  alternative  development  of  their  parts.  When 
the  exopodite  is  large  the  endopodite  is  small,  and 
vice  versd.  Besides  these  general  modifications  in 
which  the  degenerative  evolution  of  the  entire 
appendages  is  exhibited,  each  one  of  the  segments 
undergoes  certain  modifications  of  its  own. 


Fig.  S^.—Astacus  fluviatUis.  Third  left  maxillipede 
(1,5/1):  CO,  coxopodite,and  ba,  basipodite,  forming 
pr,  protopodite ;  br,  gill ;  ear,  exopodite ;  en,  en- 
dopodite (Huxley). 


Fig.  9.— Astacus  fluviatUis.  Second 
left  maxillipede  (1,5/1) :  co,  coxo- 
podite,  and  6a,  basipodite,  forming 
pr,  protopodite  ;  en,  endopodite, 
ex,  exopodite  ;  br,  gill  (Huxley). 


In  the  second  maxillipede  (fig.  9),  which  much 
resembles  the  third,  the  eocopodite  {ex)  is  large,  and 
the  endopodite  {en)  is  small,  the  prodopodite  (pr.)  is 
better  developed,  and  the  podohranch  {hr.)  has 
begun  to  atrophy. 

The    same  evolution  is  exhibited    by    the   first 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS 


37 


maxillipede  (fig.  10),  but  here  it  is  more  striking, 

and    considerably    modifies 

the    general    appearance   of  en 

the  appendage.     The  exopo-         , 

dite  (ex.)  is  well  developed,       j 

especially  at  its   base  ;  the   "^  '' 

endopodite  (en.)  is  small  and 

consists  of  only  two  joints ; 

in  the  prodopodite  (  pr.)  the  .  .    „ 

^  ^  xx-     /  FiQ.lO.—AstacusJIuviatilis.   First 

two       component       segments       leftmaxillipede(l,5/l):co,coxo- 

■^  "  podite,  and  6a,  basipodite,  foiin- 

p  ing  pr,  protopodite ;  en,  endo- 

/'  .ca  podite;   ex,  exopodlte ;   ep,  epi- 

podite  (Huxley). 

are  transformed  into 
two  long  thin  plates, 
and  the  podohranch 
is  replaced  by  a  mem- 
braneous plate  (the 
epipodite)  (ep).  Behind 
the  maxillipedes  are 
the  claws  and  the  four 
walking  legs.  In  these 
five  pairs  of  appen- 
dages the  exopodite 
shows  most  signs  of 
degeneration.    We  will 

Fio.   11. — Astacus  fluviatilii.    Second  left 

walking  lee  (1,5/1) :  co,  coxopodite  and  nOW  CXamilie  the  COm- 
ba,  basipodite,  forming  pr,  protopodite; 

6r,  gill;  t«,  ischiopodite; /7je,meropodite;  pOUCIlt    partS    of    thcSC 

ca,  carpopodite; />,  propodite;   rfu,  dacty-  ^  -"^ 

lopodite  (Huxley).  appCudagCS  (fig.    11). 

T\\Q  protopodite  (pr.)  consists  of  two  parts  (ba.  and 
CO.),  and  carries  a  gill  (br.)  except  in  the  last  walking 


38       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

leg.  The  endopodite  consists  of  five  primitive  joints, 
which  are  well  developed  and  form  the  ordinary 
claw ;  these  five  parts  are  the  ischiopodite  (is.),  the 
mesopodite  (me.),  the  carpopodite  (ca.),  the  propodite 
(pr.),  and  the  dactylopodite  (da.).  The  exopodite  is 
missing.  No  vestige  or  rudiment  of  it  is  to  be 
found  in  any  phase  of  the  development  of  the  cray- 
fish. In  the  lobster,  however,  which  is  closely 
allied  to  the  cray-fish,  the  exopodite  is  still  to  be 
found  during  the  larval  period.  The  third  funda- 
mental part  of  the  primitive  member  persists  also 
in  prawns  throughout  the  entire  period  of  life,  but 
the  organ  is  very  small.  At  the  extremity  of  the 
first  and  second  pairs  of  walking  legs  there  is  an 
apparatus  consisting  of  a  fixed  part — an  elongation 
of  the  protopodite — and  of  a  moveable  part — the 
dactylopodite.  This  furnishes  the  walking  leg  with 
a  prehensile  organ  which  is  well  developed  in  the 
first  pair  of  walking  legs,  and  which  is  enormously 
increased  in  the  true  claws.  In  this  evolution 
degeneration  is  exhibited  by  the  disappearance  of  a 
joint,  for  in  these  appendages  the  basipodite  and  the 
ischiopodite  are  immovably  united.  This  morpho- 
logical degeneration  corresponds  to  a  functional 
change  in  the  appendage.  So  long  as  the  claw  was 
used  for  locomotion  a  joint  at  this  point  was  in- 
dispensable for  progression.  It  is  this  joint  which, 
in  six  or  eight-footed  beasts  allows  of  the  horizontal 
motion  of  the  member  necessary  for  locomotion, 
which  in  six-footed  beasts  results  from  the  general 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS        39 


structure  of  the  parts  and  their  auricular  combina- 
tions. On  the  claw  becoming  prehensile,  the  joint 
hinge  consolidates,  the  lever 
thus  becoming  much  stronger 
and  permitting  the  claw  to  be 


sc 


Fio.  12.  —  Aslacus  Jiuvialilis. 
Second  left  maxilla  (1,5/1): 
CO,  coxopodite,  and  6a,  basipo- 
dlte,  forming  pr,  protopodite ; 
e»,  endopodite;  sc,  scaphog- 
nathlde  (Huxley). 


used  to  greater  advantage. 
It  now  remains  to  examine 

the    appendages    situated    in 

front  of  the  maxillipedes,  i.e. 

the  appendages  of  the  head. 

In  the  second  maxilla  (fig.  12) 

a  special  transformation  may 

be  observed.     The  coxopodite 

(co.),    and    the  basipodite  (ha.),  are  flat  plates  ;  the 

endopodite  {en.),  which  is  small  and  undivided,  ex- 
hibits signs  of  degeneration  in  its 
size,  and  in  the  absence  of  all 
articulations.  The  exopodite,  accord- 
ing   to    some    authorities    on    the 

FiG.is.—Astacusfluvia-  subicct,    HO    longcr    cxists,    while 

mis.    First  left  max-  "^     _  ^^  .  . 

ilia  (1,5/1):  CO,  coxo-  accordinsj    to   others  it   constitutes 

podite,  and  ba,  basi-  ,  .        7 .         /  ^  i 

podite,  forming  pr.  With   the    evipoditc   (the  aualogous 

protopodite;    en,  en-  i  i  •    i  i 

dopodite  (Huxley),  part  to  that  which  wc  regard  as 
representing  the  gill  in  the  maxillipede),  a  large 
peculiarly-shaped  blade,   the  scaphognathide  (sc.). 

In  the  first  maxilla  (fig.  13),  a  partial  degenera- 
tion of  the  organ  is  very  marked :  the  exopodite  and 

^  See  J.  Demoor,  Recherches  sur  la  Marche  des  Crustac6s  (Arch, 
de  Zool.  exp.  etgeii.,  2°  serie,  t.  iv. ,  1891). 

T.  List,  Bewegungsapparat  des  Arthropoden,  1.  Theil,  Astacus 
Fluviatilis.     Morphol.  Jahrbuch.,  xxii.  Bd.  3.  Heft,  1895. 


40       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


the  epipodite  are  missing ;  the  endopodite  is  reduced 
to  a  mere  unjointed  stem,  and  only  the  protopodite 
retains  its  two  normal  component  parts. 

The  mandibles  (fig.  14),  the  appendages  of  the 
fourth  segment,  are  modified  entirely  for  mastica- 
tion. They  consist  of  a  strong  transverse  piece 
(pr.)  provided  at  the  extremity  with  an  inner 
surface  (st)  for  grinding  and  sawing,  and  of  a 
three-jointed  piece  (en.)  with  bristles  which  point 
outwards.     The  first    piece   is   the    result    of    the 


en 


r><j^ 


p 

Fio.   a.  —  Astacuit  fluvia-  Fig.    15.— As tacus  fluvia-  Fio.  16.  —  Astacus  ^uvia- 

tilis.        Left     mandible  tilts.         Left      antenna  tilts.       Left     antennule 

(1,6/1) :  pr,  protopodite;  (1,5/1)  :  pr,  protopodite  ;  (1,5/1)  :  pr,  protopodite; 

en,  endopodite;  st,  rasp-  ex,  exopodite;  en,  endo-  ex,  exopodite;  en,  endo- 

ing  surface  of  the  proto-  pod  ite  (Huxley).  podite  (Huxley), 
podite  (Huxley). 

modification  of  the  two  parts  of  the  protopodite, 
which  have  united  to  form  the  organ  of  mastica- 
tion ;  the  second,  which  represents  the  endopodite, 
is  the  feeler,  an  organ  of  sensation.  All  the  other 
parts  of  the  appendage  have  disappeared. 

The  antenna  (fig.  15),  which  is  a  tectile  ap- 
paratus, is  formed  of  two  parts  representing  the 
segments  of  the  protopodite  (pr.).  The  long-ringed 
process  is  the  eiidopodite,  while  the  lateral  scale  of 
the  antenna  represents  a  much  reduced  exopodite  (ex.). 


TBANSFORMATION  OF  ORGANS  OF  ANIMALS        41 

The  antennule  (fig.  1 6)  consists  of  a  protopodite 
(pr.),  furnished  with  an  annulated  endopodite  (en.) 
and  exopodite  (ex.). 

The  eye-stalk  (fig.  17)  consists  of  a  two-jointed 
protopodite  (pr.).  This  is  all  that  remains,  the 
endopodite  and  the  exopodite  being  ya^ 

absent  altogether.  kt)'?^ 

This  examination  of  the  appen-  „     ,,^^,       ^   . 

^i^  Fig.  n.—Aitacus  fluvia- 

dages  of  the  cray-fish  clearly  shows     ff  5*/i,  .^^."  rX"S 
•that  all  fresh  adaptation    in   the     (Huxiey).  ' 
appendage  entails  the  modification  of  some  parts 
and    the    degeneration    of    others.      In    each    case 
evolution  is  accompanied  by  degeneration. 

§   3.    Transformation    of  homologous    organs    in 
individuals  of  different  species. 

The  limitations  of  our  present  knowledge  make 
it  difficult  to  determine  definitely  the  origin  of 
limbs  among  vertebrates,  but  they  are  universally 
supposed  to  have  developed  from  the  lateral  folds 
which  still  persist  in  Amphioxus,  and  which  pro- 
bably existed  in  the  ancestors  of  vertebrates.^ 

^  A.  Morphological  proofs : 

(a)  Lateral  folds  of  amphioxus. 

(&)  Identity  of  the  skeletons  of  paired  and  unpaired  fins. 

(c)  The  number  of  spinal  nerves  passing  to  the  fins. 

{d)  The  mode  of  entrance  of  these  nerves  into  the  fins. 
B.  Emhryological  proof s : 

(a)  Continuous  lateral  folds  in  the  embryoes  of  fish. 

(6)  The  formation  of  metameric  pouches  (coelomic  in- 
vaginations not  only  at  the  point  of  origin  of  the 
limbs,  but  between  them). 


42       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

At  first  the  only  skeleton  of  these  lateral  rods 
consisted  of  parallel  rods  of  a  tough  material.  By 
the  transformation  of  these  rods,  the  skeleton  of 
the  limbs  of  vertebrates  was  ultimately  formed. 

The  skeleton  of  limbs  then  consisted  originally 
of  a  certain  number  of  parallel  rods.  One  of  these 
rods,  lying  in  the  long  axis  of  the  future  limb, 
became  longer  than  the  others,  while  the  neigh- 
bouring rods  began  to  slant,  so  that  those  nearest 
to  the  elongated  rod  spread  out  like  a  fan,  and 
gradually  moved  outwards  along  the  principal  rod. 
This  phenomenon  was  repeated  several  times,  so 
that  eventually  those  rods  nearest  to  the  principal 
rod  passed  towards  the  free  end  of  it,  and  as  the 
others  followed  in  the  same  direction,  the  fin  finally 
acquired  a  feather-like  structure. 

This  transformation  of  the  continuous  folds  into 
limbs — a  progressive  transformation,  since  new  and 
more  perfectly  adapted  organs  were  formed — was 
accompanied  by  degeneration,  for  a  considerable 
part  of  the  folds  disappeared.  In  the  same  way, 
although  the  transformation  of  the  parallel  rods 
into    bipinnated    fins    constituted    a    development, 

C.  Palocontological  proofs  : 

{a)  The  series  of  paired  spines  in  Diplacanthus  and  in 
Climatius  between  the  pectoral  and  ventral  spines. 
(6)  The  skeleton  of  the  fins  of  Cladoselache. 

D.  PMlogenetic  proof:  The  necessity  of  the  bifurcation  of  the 

unpaired  folds  at  the  anal  region  {Stemarchus). 

E.  Physiological  proof:  The  lateral  folds  are  the  undifferenti- 

ated condition  of  organs  of  e(juilibriuin  in  modern  fish. 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS         43 

the  rods  nearest  to  the  free  extremity  of  the 
principal  rod  were  considerably  reduced  in  size. 

Two  series  of  transformations  should  now  be 
followed — that  of  limbs  which  have  not  yet 
ceased  to  be  aquatic,  and  that  of  limbs  adapted 
to  terrestrial  existence. 

Among  aquatic  creatures,  the  Pleuracanthides 
and  the  Dipneusti  {Xenacanthus  and  Ceratodus) 
have  best  preserved  the  bipinnated  fin.  In 
Ceratodus  especially  it  is  exhibited  in  almost  the 
primitive  condition.  Gradually,  however  (as  in 
Orthacanthus),  the  rods  situated  along  one  edge 
of  the  principal  rod  disappeared,  and  the  fin,  no 
longer  bipinnated,  became  unilateral.  This  pro- 
gressive transformation  entailed  the  disappearance 
of  nearly  half  of  the  fin. 

With  creatures  which  have  become  adapted  to  a 
terrestrial  life,  the  limbs — so  far  as  can  be  judged 
from  what  is  known  at  present — appear  to  have 
undergone  the  following  transformations :  The  bi- 
pinnated fin  (such  as  that  of  Ceratodus)  (fig.  18) 
is  always  the  starting-point.  Then  the  lateral  rods 
of  one  half  almost  completely  disappear  (as  in 
(Protopterus  Amphihius).  Next,  the  other  half 
follows  (as  in  P.  anneeteus)  (fig.  19),  and  finally 
only  the  principal  rod  remains  (as  in  Zepidosiren) 
(fig.  20). 

At  this  point  the  limbs  are  reduced  to  mere 
lopped  stems ;  they  have  not,  however,  atrophied ; 
the  degeneration  which  has  accompanied  this   de- 


Fig.  lii.—Ceralodus  Forsteri,  Krefft.  Bipinnate  fins  (after  Giinther).    {See  Dollo,  Su 


Fig.  20. — Lepidosiren  paradoxa,  I 


I^ente  det  Dipneu$tes,  Bull.  Soc,  beige  de  GSol.,  de  Pal^ont.,  et  dCHydrol.,i.  IX.,  1895. 


(after  T.  E.  Gray).    {See  Dollo.) 


(tfter  Ray  Lankester).    {See  DotLo.) 


46        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

velopQient  has  reached  its  ultimate  limit;  the 
limb  has  now  to  disappear  altogether,  or  else  to 
provide  itself  v^ith  new  elements. 

Judging  from  the  embryology  of  the  Urodele 
Amphibia,  the  primitive  terrestrial  limb  had  only 
one  toe  at  first,  and  then  a  second  appeared,  then 
a  third,  a  fourth,  and  a  fifth.  In  this  way  was 
formed  the  primitive  five- toed  limb. 

Let  us  now  trace  the  adaptation  of  this  five-toed 
limb  to  the  principal  functions  it  fulfils  in  the 
animal  kingdom — walking  upon  two  legs,  running, 
jumping,  climbing,  burrowing,  swimming,  and  flying. 
We  shall  find  that  the  transformations  it  has  under- 
gone for  all  these  functions  have  been  accompanied 
by  degeneration. 

1.  Adaptation  to  walking  on  two  legs. — Adapta- 
tion to  the  upright  position  may  be  effected  in  two 
different  ways,  by  walking  or  jumping.  We  will 
first  investigate  the  former. 

This  adaptation  to  bipedal  progression  may  be 
effected  with  or  without  the  intervention  of  arboreal 
life,  as  may  be  seen  by  the  following  examination, 
the  adaptations  of  man  and  of  birds. 

(a)  Man. — The  first  sign,  among  mammals,  of  a 
definite  adaptation  to  arboreal  life,  is  the  oppos- 
ability  of  the  great  toe.  The  great  toes  of  human 
beings  are  not  opposable,  the  cases  of  "  prehension  " 
exhibited  among  savage  races  being  due  to  lateral 
movements ;  we  shall  see  presently,  however,  that 
this  opposability  of  the  great  toe  which  is  exhibited 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS         47 

by  all  modern  monkeys,  probably  existed  at  some 
time  in  man  or  liis  immediate  ancestors. 

The  terrestrial  five-toed  foot  must  have  acquired 
an  opposable  great  toe  in  the  course  of  adaptation 
to  arboreal  life.  Without  attempting  to  enumerate 
all  the  attendant  phenomena  of  degeneration  en- 
tailed by  this  modification,  we  will  merely  examine 
those  exhibited  by  the  nails  of  the  hind  foot. 
Comparison  with  the  supposed  ancestors  of  the 
Primates  (Insectivores)  shows  that  in  the  primitive 
terrestrial  foot  the  nails  completely  covered  the 
extremities  of  the  terminal  phalanges.  With  the 
lowest  Lemur  (Tarsius)  the  nails  of  the  second  and 
third  toes  are  in  the  form  of  claws,  while  all  the 
others  are  degenerating;  they  cover  only  the 
anterior  face  of  the  terminal  phalanges,  and  are, 
in  fact,  flat  nails.  In  all  the  other  Lemurs,  only 
the  second  toe  has  a  claw,  whereas  in  true  monkeys 
all  the  nails  are  flat.  Finally,  in  the  Orang  the 
great  toe  often  has  no  nail  at  all. 

We  then  come  to  limbs  adapted  to  arboreal  life, 
such  as  those  of  the  Gorilla.  With  some  anthropoids, 
however,  and  especially  the  Orang,  the  modification 
has  gone  further,  and  is  accompanied  by  phenomena 
of  degeneration  in  other  parts  of  the  limb.  While, 
in  the  case  of  the  Gorilla  (fig.  21)  the  great  toe 
has  a  true  articulation  between  the  metatarsal  and 
the  first  phalanx,  and  also  between  the  first  and 
second  phalanges,  with  the  Orang  (fig.  22)  the  great 
toe  is  much  reduced  so  that  it  has  no  articulation 


48       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

between  the  metatarsals  and  phalanges,   and    fre- 
quently the  last  phalange  is  absent  altogether. 

The  development  of  another  foot,  that  of  man, 
adds  confirmation  to  the  heading  of  this  chapter. 
The  great  toes  of  man  or  his  immediate  ancestors 
must  have  been  opposable.  This  may  clearly  be 
seen  by  a  careful  study  of  the  muscles.  We  know 
what  a  resemblance  there  is  between  the  muscles 


r 


-f-'Sv. 


^TneJtaXan&e., 


Figs.  21  and  22.— Skelt-tons  of  the  feet  of  the  Goi-illa  and  Orang-outanR. 
ca   calcaneum;    as,  astragulus;    sc,  scaphoid;    Ire  cu,  first  cuneiform;    2e  cv, 
second  cuneiform ;  3e  cu,  third  cuneiform  ;  c,  cuboid  ;  J/,  metatarsals  ;   phal, 
phalanges;  ph  andp,  distal  and  terminal  joints  (after  Waterhouse  and  Hawkins). 
See  Huxley,  Man's  Place  in  Nature. 

of  the  soles  of  the  feet  and  those  of  the  palmar 
region.  With  the  exception  of  man,  all  mammals 
which  have  opposable  thumbs,  have  also  opposable 
great  toes.  On  the  other  hand,  there  are  various 
mammals  (prehensile  marsupials,  rodents,  bats,  mar- 
mosettes,  etc.)  which  have  opposable  great  toes  but 


TKANSFORMATION  OF  ORGANS  OF  ANIMALS         49 

not  opposable  thumbs.  We  may  presume  then 
that  the  hind  limbs  were  the  first  to  become 
adapted  to  arboreal  life,  and,  since  the  great  toe 
of  man  is  not  opposable,  it  may  be  concluded  that 
it  ceased  to  be  so  when  terrestrial  life  was  assumed. 
This  modification  is  not  wholly  degenerative,  for 
the  functions  of  the  lower  limbs  are  no  less  im- 
portant in  man  than  in  apes.  There  are,  however, 
undoubted  signs  of  degeneration  in  the  changes 
entailed.  The  muscles  which  rendered  the  great 
toe  opposable  were  probably  the  first  to  degenerate, 
but  by  referring  to  a  skeleton  it  may  be  seen  that 
whereas  the  great  toe  and  the  metatarsals  have 
gained  in  length,  the  first  phalanges,  and  still  more 
the  other  phalanges,  are  reduced.  In  some  cases 
the  last  phalange  of  the  little  toe  has  even  dis- 
appeared altogether. 

(h)  Birds. — First  take  the  reptile-footed  Dino- 
saurians.  These  are  four-footed  beasts,  and  the 
hind  limbs  are  plantigrade  and  five-toed.  Counting 
from  the  inner  side  the  phalanges  of  the  toes  are : 
2,  3,  4,  5,  4  in  number. 

In  passing  on  to  the  bird-footed  Dinosaurians 
we  find  that  with  Hypsilophodon,  the  foot  is  func- 
tionally four-toed.  From  the  first  to  the  fourth 
toe  the  phalanges  are  the  same  in  number  as  with 
the  reptile-footed  Dinosaurians,  but  the  fifth  toe  is 
greatly  reduced,  being  merely  represented  by  a 
pointed  metatarsal.  In  Camptosaurus,  the  fifth  toe 
has  disappeared,  the  second,  third,  and  fourth  are 

D 


50       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

fully  functional,  but  the  first,  although  retaining  the 
usual  number  of  phalanges,  is  greatly  reduced  in 
size.  Finally,  in  Iguanodon,^  which  is  fully  adapted 
to  the  upright  position,  the  second,  third,  and  fourth 
toes  are  still  fully  functional,  but  not  only  has  the 
fifth  toe  disappeared,  but  the  first  is  merely  repre- 
sented by  a  pointed  metatarsal. 

Thus  it  is  apparent  that  each  new  phase  in  the 
transformation  from  quadruped  to  biped  is  attended 
by  partial  degeneration,  although  the  hind  limbs, 
having  alone  to  fulfil  the  function  of  locomotion, 
necessarily  acquire  a  fuller  development.  The 
IgvAiTwdon  is  too  specialized  in  its  development 
to  have  been  the  ancestor  of  birds ;  but  this  is 
not  so  with  other  bird-footed  Dinosaurians,  such 
as  Camptosaurus  and  Hypsilophodon,  where  the 
hind  foot  is  sufficiently  primitive  to  have  been 
the  forerunner  of  the  bird's  claw. 

Most  birds  have  four  functional  toes,  which  have 
retained  the  primitive  number  of  phalanges,  but  the 
legs  of  running  birds  exhibit  more  or  less  important 
variations.  The  modifications  they  have  undergone 
have  obviously  not  diminished  their  functional  im- 
portance, but  quite  the  contrary.  These  modifications 
have,  however,  been  accompanied  by  degeneration 
with  running  birds,  and  the  first  toe  disappeared 
first,  as  in  the  Cassowary  and  the  American  Ostrich, 

^  DoUo  :  First,  second,  third,  fourth,  and  fifth  Notes  sur  les 
Dinosauriens  de  Bernissart.  Bull.  Mils,  royal  d'Hist.  nat.  de  Belg. 
1882,  1883,  1884. 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS         51 

to  be  followed  in  the  second  stage  by  the  second 
toe.  Then  the  fourth  began  to  atrophy  until  the 
whole  weight  of  the  body  was  supported  by  the 
third  toe  alone,  as  in  the  ostrich. 

2.  Adaptation  to  leaping — We  now  come  to  the 
various  instances  of  adaptation  to  leaping.  This 
adaptation  is  one  necessarily  undergone  either  by  an 
animal  which  originally  lived  a  terrestrial  life,  such 
as  the  Jerboa,  or  by  one  which  had  ceased  to  be 
arboreal  in  adapting  itself  to  leaping,  such  as 
the  kangaroo ;  or,  again,  by  an  animal  which 
formerly  lived  an  arboreal  life,  and  which  con- 
tinued to  lead  such  a  life  with  the  additional 
adaptation  to  leaping,  such  as  the  tarsius  ;  and, 
finally,  by  an  animal  which  first  lived  and  swam 
in  the  water,  and  which  continued  to  do  so  after 
becoming  adapted  to  leaping,  such  as  the  frog. 

In  each  of  these  cases  adaptation  to  leaping  has 
resulted  in  the  fuller  development  of  the  hind  limbs, 
the  size  of  which,  as  well  as  the  functional  impor- 
tance, has  greatly  increased,  which  shows  that  the 
modification  is  of  a  progressive  nature.  All  these 
modifications  are,  however,  attended  by  degeneration. 

{a)  The  Jerhoa  (Dipics). — Some  rodents,  such  as 
rats,  porcupines  and  squirrels,  are  five-toed.  This 
five-toed  foot  represents  the  primitive  one  from 
which,  after  passing  through  various  still  existing 
phases,  was  derived  the  three-toed  foot  of  the  jerboa. 

With  the  hare  there  is  no  first  toe,  so  that  the 
foot  is  functionally  four-toed. 


52       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


The  fifth  toe  next  disappeared,  leaving  the  foot 
three-toed,  as  in  the  viscacha,  the 
capybara  and  agouti.  In  the  foot 
of  the  agouti  the  metatarsals  have 
considerably  gained  in  length,  and 
are  closely  pressed  together. 

In  the  jerboa  (fig.  23)  the  three 
metatarsals  (if.)  are  united,  and 
the  intervening  tissues  have  dis- 
appeared. Besides  this  modifica- 
tion, the  hind  limbs  of  the  animal, 
being  used  for  leaping  only,  have 
become  thinner,  while  gaining  in 
—Hind  limb  of  length  hv  the   elongation   of   the 

(Jerboa).    I,  first  ■, 

metatarsals.^ 

(h)    The  Kangaroo. — Although 


Fio.  23. 
Dipus 

rudimentary  digit;  II, 
second  digit;  III,  third 
digit;  IV,  fourth  digit; 
if,    bone    formed    bv  .    ,     ,  .  p        ^  ^    • 

fusion  of  2nd,  3rd,  and  ROW    inhabitants     of     thc    plain, 

4th    metatarsals;      ca,  i     i        « 

caieaneum;  as,  astra-  kauffaroos    are     desccndcd     from 

gulus;     ra,    navicular;  *="  •    i  t       i        i 

c3,  third  cuneiform ;  c2,  arborcal  marsupials,  which  them- 

second  cuneiform ;    cb, 

cuboid.  (After  Flower,  sclvcs  havc  gradually  ceased  to  be 

Introduction  to  the  Oste-  in         ■,  . 

oiogyo/the  Mammalia.)  arborcal.^  All  tlic  intervening 
phases  of  this  transition  from  type  to  type  are  known. 

^  The  last  segment  of  the  leg  of  the  jerboa — the  foot  being 
formed  by  the  union  of  the  second,  third  and  fourth  metatarsals 
— corresponds  more  or  less  to  the  formation  of  the  bird's  foot. 
This,  however,  is  a  case  of  convergent  modification  more  apparent 
than  real,  for  in  the  cannon  bone  of  birds,  besides  the  second,  third 
and  fourth  metatarsals,  there  is  the  whole  set  of  distal  tarsals, 
whereas  in  the  jerboa  the  distal  tarsals  remain  free  and  isolated 
bones. 

^  L.  Dollo,  autographs  of  the  course  given  at  the  Institute  of 
Solvay,  1891-1892,  5th  lecture,  The  Origin  of  Kangaroos. 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS 


53 


In  the  Opossum  (Didelphys)  we  find  a  normal 
five-toed  foot,  in  which  the  third  toe  predominates, 
bnt  which  is  fully  adapted  to  arboreal  life,  the  first 
toe  being  opposable  and  provided  with  a  flat  nail. 

The  foot  of  the  Phalanger 
(Fhalangista)  (fig.  24)  is 
also  five-toed,  and  the  great 
toe  is  opposable,  but  here  it 
is  the  fourth  toe  (iv.)  which 
predominates  owing  to  the  ^^ 
degeneration  of  the  second 
(ii.)  and  third  (iii.) 

The  Kaola  (Phascolardos) 
exhibits  the  same  type  of 
foot,  but  with  an  even  more 
marked  diminution  of  the 
second  and  third  toes. 

In  Hypsiprymnodon  the 
first  toe  has  become  rudi- 
mentary. 


In  Ferameles  the  first  toe 


Fig.  24.— Right  fore-paw  of 
Phalanger. 
c,    calcaneum;    a,    astragalus; 


OS  centrale ;  cb,  cuboid ;  ci,  c2,  c^, 
flrst.seoond,  and  third  cuneiforms ; 

is    only   represented    by    a     l^i^i^lTh^a^I^'S^^'^; 
metatarsal  with  phalanges.       ?rfrC\tS.'otS^^^^ 

In   the  kangaroo  (fig.  25)       osteology  o/ the  Mammalia.) 

it  has  completely  disappeared. 

Finally,  in  Chceropus  (fig.  26)  the  fifth  toe  is 
reduced  to  a  mere  thread,  as  are  also  the  second 
(ii.)  and  third  (iii.)  toes. 

This  last  stage,  which  represents  the  present  and 
most  perfect  adaptation  of  marsupials  to  leaping, 


54       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


was  attained  by  the  predominance  of  the  fourth  toe, 
accompanied  by  the  almost  total  extinction  of  the 
other    toes.      The    persistence    of    the   fourth   toe 


Fig.  25.— Righf  fore-paw 
of  Kangaroo. 

c,  calcaneuni;  a,  astragu- 
lus;  cb,  cuboid;  n,  os 
centrale;  cn.third  cunei- 
form. II,  III,  IV,  V,— 
second,  tiiird,  fourth 
and  fifth  digits,  the 
second  and  third  being 
degenerate.  (After 

Flower's  Introduction 
to  the  Osteology  of  the 
Mammalia.) 


Fig.  26, — Choeropm. 
Right  fore-paw. 
c,  calcaneum  ;  a,  astraga- 
lus ;  n,  OS  centrale ;  cb, 
cuboid;  c^,  3e,  third 
cuneiform;  II,  III.  IV, 
V,  —  second,  third, 
fourth  and  fifth  digits, 
of  which  the  second, 
third  and  fifth  are  de- 
generating. (After 
Flower's  Introduction 
tfl  the  Osteology  of  the 
Mam,malia.) 


Fig.  27.  —Right  hind-paw 
of  Tarsius  Spectrum. 

c,  calcaneum ;  a,  astragu- 
lus;  r,  navicular;  ci,  c'\ 
c^,  first,  second,  and 
third  "cuneiforms  ;  cb, 
cuboid  ;  I,  II,  III,  IV, 
V,— first,  second,  third 
and  fourth  digits.  (After 
Flower's  Introduction 
to  the  Osteology  of  the 
Mammalia.) 


instead  of  the  third  is  accounted  for .  by  the 
previous  arboreal  existence  of  marsupials  before 
they  became  adapted  to  leaping. 

(c)  The  Tarsius  (Tarsius  spectrum). — The  tarsius, 
which  is  a  small  lemur,  is  to  be  found  in  Celebes. 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS         55 

It  is  so-called  owing  to  the  remarkable  length  of 
its  tarsus  (fig.  27),  a  very  exceptional  formation, 
for  the  elongation  of  an  animal's  foot  is  usually 
effected  by  means  of  a  lengthened  metatarsal. 
This  abnormal  formation  in  the  foot  of  the 
tarsius  can,  however,  be  accounted  for.  Like 
most  of  the  Primates,  the  tarsius  is  adapted  to 
an  arboreal  life,  and,  among  other  characters,  it 
exhibits  an  opposable  great  toe.  When  on  the 
ground,  however,  instead  of  running  on  two  or 
four  legs,  it  progresses  by  leaping ;  and  this 
may  also  be  noticed  with  some  of  the  lemurs 
of  Senegal  (Galago).  The  tarsius  has  become 
adapted  to  leaping  by  the  lengthening  of  the 
hind  limbs,  the  thigh,  leg  and  foot  being  equally 
elongated.  The  foot,  however,  being  of  fan-like 
structure,  it  is  clear  that  if  all  the  toes  were 
elongated,  the  feet  would  overlap  one  another 
in  the  middle  line,  and  therefore  with  most 
animals  that  leap  the  elongation  of  the  foot 
involves  a  reduction  in  size  of  the  outer  and 
inner  toes  to  prevent  overlapping.  The  tarsius, 
however,  has  remained  arboreal,  although  it  has 
taken  to  leaping,  and  it  therefore  requires  a  well- 
developed  foot  with  an  opposable  great  toe  in  order 
to  obtain  a  firm  grip  on  branches.  To  avoid  over- 
lapping of  the  feet,  the  length  has  been  augmented 
at  the  ankle,  which,  not  being  shaped  like  a  fan, 
does  not  consequently  gain  in  length. 

A   more   detailed   examination   would    doubtless 


56        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

show  that  this  modification  of  the  foot  has  been 
attended  by  the  degeneration  of  some  parts  of  it, 
but,  as  we  have  to  restrict  ourselves  here  to  the 
data  with  which  we  are  already  furnished,  it  is  the 
degenerative  phenomena  exhibited  by  other  parts  of 
the  limb  consequent  to  adaptation  to  leaping,  which 
we  shall  proceed  to  enumerate. 

There  are  two  leg  bones,  the  fibula,  and  a  larger 
one,  the  tibia.  These  two  bones  are  usually  held 
together  by  muscles,  or  by  an  interosseous  membrane, 
the  latter  being  merely  degenerated  muscular  tissue. 
As  a  rule,  the  tibia  and  the  fibula  reach  the  entire 
length  of  the  leg  and  are  quite  separate.  In  the 
process  of  adaptation  to  arboreal  life,  these  bones 
have  remained  apart,  as  may  be  observed  in  most 
arboreal  animals  of  the  present  day.  Moreover,  in 
order  to  promote  prehension  by  the  foot,  the  tibia  and 
fibula  in  prehensile-footed  marsupials  may  turn  upon 
one  another,  as  is  the  case  with  our  radius  and  ulna. 
With  the  tarsius  it  is  quite  otherwise.  Here  the 
lower  halves  of  the  tibia  and  fibula  are  united,  and 
the  muscles  which  formerly  connected  the  two  bones 
have  consequently  disappeared.  There  is  further 
evidence  of  degeneration  in  the  upper  half  of  the 
fibula  which  has  become  reduced  to  a  mere  thread. 
There  is  no  question  that  these  phenomena  are  the 
result  of  adaptation  to  leaping,  for  they  are  not 
exhibited  among  exclusively  arboreal  animals,  while 
many  parallel  cases  of  modification  are  known  to 
exist  amongst  other  species. 


TKANSFORMATION  OF  ORGANS  OF  ANIMALS    57 

(d)  The  Frog  (Bana  Esculenta). — Adaptation  to 
leaping  entails,  as  a  rule,  the  elongation  of  the  hind 
limbs.  When  a  large  blade-like  swimming  foot  is 
required,  the  number  of  toes  cannot  be  diminished 
and  length  must  be  augmented,  as  in  the  tarsius,  in 
the  region  of  the  ankle  (fig.  28). 
But,  in  the  frog,  it  is  the 
astragulus  {a.)  and  the  calcan- 
eum  (c.)  which  have   elongated,  l\  /     ^.^ 

whereas    in    the  tarsius    it  was  |(?!oa^3^B 

the    calcaneum    and    navicular  />p>^^<r\ 

bones. 

Besides  the  union  of  the 
calcaneum  and  the  astragulus 
bones — which  may  be  regarded 
as  a  phenomenon  of  degenera- 
tion, as  it  entails  the  disappear- 
ance of  the  intervening  tissues 
— the  adaptation  of  the  frog  to 
leaping  has  been  attended  by  a  FiG.28.-Kight  foot  of  izana 
yet  more  characteristic  sign  of  a,  ^strngSsTc,  caican- 
degeneration :  the  tarsus,  which  ^""„,U;ta?sa1sfi,n^^^^^^ 
is  a  very  complicated  structure  ^Zker,' ISmy  i/^^l 
in  tailed  amphibia,  is  reduced,  ^'"''^•^ 
especially  in  the  distal  row,  to  a  few  minute 
bones. 

3.  Adaptation  to  running. — The  ruminants  and 
the  horse  furnish  us  with  two  excellent  examples 
among  mammals  of  adaptation  to  running. 

Their  common  ancestor  is  known  to  have  been  a 


58        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

clumsy  five-toed  animal  which  lived  in  swamps.  As 
the  horse  type  was  evolved,  the  legs,  which  originally 
only  served  to  support  the  weight  of  the  body  in  a 
slow  progress,  were  gradually  adapted  to  running. 
This  modification  was  certainly  advantageous  to  the 
animal,  as  it  enabled  it  to  escape  more  easily  from 
its  enemies.  It  was,  however,  accompanied  by  de- 
generation, for  the  ancestor  of  the  horse  lost  first  the 
great  toe,  then  the  fifth,  and  next  the  second  and 
third  toes  became  so  reduced  that  eventually  only 
one  toe  remained  functional. 

Adaptation  to  running  was  effected  differently 
with  the  ruminants,  but  it  was  equally  accompanied 
by  degeneration.  With  them,  first  the  great  toe 
disappeared,  then  the  second  and  fifth  became  simul- 
taneously much  smaller,  while  the  third  and  fourth 
continued  to  increase  equally  in  length.  In  process 
of  time  the  second  and  third  toes  entirely  disap- 
peared, and  the  metatarsals  of  the  third  and  fourth 
joined  together  to  form  the  cannon  bone.  In  this 
way  mammals  were  evolved  with  cloven  hoofs  and 
adapted  to  running. 

4.  Adaptation  to  flying. — There  are  three  types 
of  animals  among  the  vertebrates :  birds,  ptero- 
saurians,  and  flying  bats,  by  which  is  meant,  not 
those  bats  which  are  merely  able  to  float  in  the 
air  like  a  parachute,  but  those  which  can  both  raise 
and  guide  themselves  during  flight. 

{a)  Birds. — We  have  seen  that  the  ancestor  of 
all  terrestrial  vertebrates  was  a  five-toed  animal. 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS    59 

The  hand  of  the  oldest  known  bird,  however, — 
Archceopteryx  —  had  only  three  digits,  and  the 
question  arises  as  to  whether  this  atrophy  of  the 
two  outer  digits  was  effected  in  the  process  of 
the  animal's  gradual  adaptation  to  flight  or  if  it 
dates  from  a  still  earlier  period. 

Archmopteryx  had  four  toes.  The  fore  legs  were 
smaller  than  the  hind  legs.  Now,  with  bipedal  or 
quadrupedal  animals  which  make  no  special  use 
of  their  fore  legs,  the  contrary  may  invariably  be 
noticed.  It  is  of  small  importance,  therefore, 
whether  Arclimopteryx  was  quadrupedal  or  bipedal 
before  it  took  to  flying,  for  neither  of  these  modes 
of  progression  would  have  entailed  the  loss  of  two  of 
its  digits,  and  the  disappearance  may  reasonably  be  at- 
tributed as  due  to  the  process  of  adaptation  to  flight. 

The  three  remaining  digits  of  Archceopteryx  were 
complete  in  their  structure.  The  phalanges,  which 
were  of  different  lengths,  are  furnished  with  claws, 
and  were  of  the  same  number  as  is  normal  among 
reptiles,  i.e.  2,  3,  4,  counted  from  the  inner  side. 
The  wing  of  Archceopteryx  had  not  therefore  under- 
gone much  modification  beyond  the  toes  of  the  two 
outer  digits,  but  it  must  be  remembered  that  the 
bird  was  a  weak  flyer ;  instead  of  having  a  large 
boney  sternum  necessary  for  powerful  flight,  the 
sternum  of  Archceopteryx  was  formed  of  cartilage, 
and  the  wings  were  short  and  rounded  in  shape. 

With  good  flyers,  on  the  other  hand — such  as 
pigeons  or   sea-birds — the   adaptation   to  flight   is 


60        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


more  perfect,  the  modifications  entailed  are  much 

greater  and  have  been  accompanied  by  further  signs 
of  degeneration.  In  the  fowl 
(fig.  29)  all  the  claws  have 
disappeared  {degeneration  No.  1)  ; 
the  thumb  (i.)  and  the  index 
finger  (ii.)  have  each  lost  a 
phalanx  ;  the  middle  finger  (iii.) 
has  lost  three  (degeneration  No. 
2) ;  finally,  the  metacarpals  of 
the  index  and  middle  fingers 
have  fused  (a.)  thus  entailing 
the  degeneration  of  the  muscles 
between  those  parts  {degeneration 
No.  3). 

(h)  The  Pterosaurians. — These 
reptiles,  which  are  now  extinct, 
were  able  to  fly,  like  bats,  by 
means  of  a  membrane.  There 
is  no  connection  between  them 

Fig.  29.-skeieton  of  wing  and  birds  which  fly  in  quite  a 

i,Fir8tdigfto7thumb;  II,  different  way. 
nnSirS^r^grStdlliE;       The  mcmbrauc  of  the  Ptero- 
?h/TeconrrrlVi  saurians  (fig.  30)  was  supported 
digits.  (After  Huxley.)    ^^   ^   framcwork   of    which  the 

most  important  part  was  the  fifth  finger  (v.),  the 
digit  corresponding  to  our  little  finger,  which  was 
greatly  elongated. 

The  patagium  (the  flight  membrane)  consisted  of 
two  kinds  of  membrane,  the  antebrachial  membrane 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS        61 


which  extended  from  the  neck  to  the  wrist,  and  the 
true  wing  membrane,  which  reached  from  the  free 
end  of  the  little  finger  to  the  ankle. 

There  are  three  essential  types  of  Pterosaurians. 
Fterodadylus,  which  had  a  short  tail  and  teeth  all 
along  the  gape; 
Rham'phory  -  \ 
hnchus,  which 
had  a  long  tail 
and  a  beak  set 
in  the  front 
part  of  the  jaw 
with  teeth  be- 
hind it ;  and 
Pteranodon, 
which  had 
neither  teeth 
nor  tail.  This 
last  type  was 
the  most  spe- 
cialized of  the 
Pterosaurians, 
and  the  strongest  flyer  of  the  whole  group.  They 
attained  to  a  huge  size,  the  skull  measurement  averag- 
ing 1  metre  20,  and  the  wing  extension  8  metres  30. 
The  modification  which  the  fore-limbs  of  the  Ptero- 
saurians underwent  in  the  course  of  their  adapta- 
tion to  flight,  was  accompanied  by  degenerative 
phenomena  in  proportion  to  the  extent  of  the 
specialization  (see  Pterodactylus  sfpectahilis)  (fig.  30). 


Fig.  30. — Pterodactylus  spectabilis  (after  Huxley, 
Anatomy  of  Vtrtebrated  Animals). 


62       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


In  the  process  of  adaptation  to  flight  the  thumb  (i.), 
and  the  little  finger  (v.)  first  lost  their  nails  and 
became  useless.  Next,  the  metacarpal  and  the 
proximal  phalange  of  the  thumb  became  much 
smaller,  and  united  with  the  bones  of  the  wrist, 
while  the  distal  phalange  degenerated  to  a  mere 
thread.  Fresh  evidences  of  degeneration  presented 
themselves  as  the  specialization  became  more  ad- 
vanced. In  Ptera- 
nodon,  for  instance, 
the  metacarpals  of 
the  clawed  digits 
were  considerably 
reduced  in  size,  the 
arms  having  almost 
ceased  to  exercise 
any  function  but 
that  of  flight.    There 

Fig.  3l.-Pteropus  (a  bat).    Skeleton  of  fore-  '^^^     ^     SpCCial     prO- 

limb:  r,  radius;  c,  degenerating  ulna  ;  I   II,  visiOU  for  the  SUpport 
III,  IV,  V,  digits.    (After  Huxley,  ^n^^omy  ^^ 

of  Vertebrated  Animals.)  of  thcSC  great  wingS, 

the  development  of  which  was  only  equalled  among 
vertebrates  by  that  of  the  pectoral  fins  of  the 
skate.  Whereas  the  legs  were  upheld  by  the  pelvis, 
which  was  in  its  turn  supported  by  the  sacrum,  the 
arms  of  Pteranodon  were  attached  to  the  middle  of 
the  shoulder  blade,  the  latter  being  supported  by 
the  vertebral  column — an  absolutely  unique  con- 
dition. This  kind  of  pectoral  sacrum  implied  the 
fusion  of  a  number  of  vertebrae,  and  involved  a 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS    63 

consequent  degeneration  of  the  muscles  which  in 
earlier  Pterosaurians  caused  the  movement  of  one 
vertebra  upon  another. 

Two  phases  may  thus  be  observed  in  the  evohi- 
tion  of  the  Pterosaurians :  first,  Pterodactylus  and 
Bhamiolwrynchus,  and  next  Pteranodon,  while  de- 
generation attended  the  modifications  of  both. 

(c)  Bats. — Bats,  like  the  Pterosaurians,  fly  by 
means  of  a  membrane,  only  that  the  membrane, 
instead  of  principally  extending  from  the  little 
finger  to  the  body,  is  equally  developed  between 
all  the  fingers. 

Let  us  consider  what  degeneration  is  involved  by 
this  modification. 

Firstly,  all  the  digits  have  lost  their  nails,  except- 
ing the  thumb  and  the  index  finger  in  fruit-eating 
bats,  and  the  thumb  alone  in  insect-eating  bats, 
while  some  of  the  phalanges  of  the  digits  are 
missing,  the  usual  number  being  two  instead  of 
three.  Further,  instead  of  the  ulna  and  radius 
being  equally  developed  in  the  fore-arm,  the  ulna 
is  greatly  reduced  in  size. 

6.  Adaptation  to  arboreal  life. — We  have  seen 
that,  as  a  rule,  the  first  evidence  of  adaptation  to 
arboreal  life  is  the  opposability  of  the  great  toe. 
This  modification  is  accompanied  by  the  degeneration 
of  the  nail,  for,  instead  of  consisting  of  a  claw  cover- 
ing the  entire  extremity  of  the  last  phalange,  it 
becomes  a  small  nail  covering  only  the  upper  side. 

Another  modification  is  exhibited  at  a  slightly 


64       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

later  stage.  In  the  primitive  foot  of  the  aminota, 
the  third  toe  is  the  longest,  but  among  animals 
which  have  become  adapted  to  an  arboreal  life  (fig. 
32)  the  fourth  toe  preponderates,  thus  allowing  of 
a  wider  grasp,  and  the  second  (ii.)  and  the  second 
and  third  toes  gradually  degenerate  {Arctocebus 
calaharensis,  Potto).  A  clear  proof  that  this  modi- 
.  fication  is  due  to  adaptation  to 

arboreal  life,  lies  in  this  same 
tendency  being  exhibited  by  two 
groups  not  closely  allied  to  one 
another — the  marsupials  and  the 
lemurs. 

In  a  still  more  advanced  stage 
the  phalanges  of  the  second  digit 
^?;rS^'*'Sght1fanT^""  disappear  almost  completely,  and, 
''fi"«e"Vof''vLtna:d  for    all    practical    purposes,    no 

(Tfte/^SuxferSS    longer  e™t- 

of  Vertebrated  Animals.)  AmOng      rCptilcS,      chamclcOnS 

furnish  a  striking  example  of  adaptation  to  climbing. 
With  them  three  digits  are  opposable  to  the  other 
two  (figs.  33  and  34),  instead  of  one  being  oppos- 
able to  the  other  four.  To  further  promote  this 
opposability,  there  is  a  fusion  of  digits  (syndactilism), 
i.e.  each  set  of  digits  is  enclosed  up  to  the  nails 
within  a  common  integument.  The  result  of  this  is 
that  the  enclosed  digits  are  no  longer  capable  of 
lateral  movement,  and  the  muscles  have  degenerated 
in  consequence.  The  wrist  and  tarsus  also  show 
signs  of  partial  degeneration. 


TEANSFOEMATION  OF  ORGANS  OF  ANIMALS    65 

6.  Adaptation  to  swimming. — Among  mammals, 
adaptation  to  swimming  entails  the  functional  dis- 
appearance of  the  lower  limbs,  and  the  modification 
of  the  fore-limbs  into  fins. 

This  modification  involves  considerable  degener- 
ation. The  alteration  in  the  number  of  digits 
varies ;  the  whale-bone  whales  have  kept  all  five ; 
in  rorquals  only  four  remain ;   and,  after  a  series 


U    % 


Fio.  33.— Skeleton  of  hand  of 
Chameleon.    (After  Cuvier.) 


Fig.  34.— Chameleon.    Skele- 
ton of  foot.   (After  Cuvier.) 


of  intermediate  conditions,  the  dolphin  is  evolved, 
the  fins  of  which  are  functionally  two-toed.  If  the 
Cetaceans  vary  as  to  the  number  of  digits  they  ex- 
hibit, they  all  have  this  character  in  common — the 
joints  have  disappeared.  The  arm  no  longer  articu- 
lates at  the  elbow,  and  neither  the  wrist  nor  the 
phalanges  are  jointed.  The  simple  stratum  of 
cartilage  which  takes  the  place  of  the  true  joints 
allows  of  a  slight  movement  of  the  various 
segments,    but    not    of    true    articular    movement. 


66        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

This  results  in  the  degeneration  of  all  the 
muscles  which  caused  this  mobility  in  their 
terrestrial  ancestors. 

A  similar  degeneration  may  be  observed  among 
Plesiosaurians,  Ichthyosaurians,  and  Mosasaurians.^ 
"With  regard  to  Sirenia,  the  Dugong  is  unable  to 
move  the  arm  at  the  elbow 
joint,  but  the  Manatee  can 
grasp   things   with   its    fins. 
Besides  many  other  modifi- 
cations,   the    adaptation    of 
these  animals  to  an  aquatic 
life  has  entailed  the  fusion 
of  the  two  bones  of  the  fore- 
arm, thus  involving  the  de- 
generation of  those  muscles 
which     formerly     prevailed 
Fio.  35.-Fore-iimb  of  yazpa     oveY  the  movcmcnt  of    the 
R,rJ::;TsJ^XTl,serni.  two  boucs  whcu  Separate. 
"^Z^^^T^/^^        ^.  Adaptation  to  hirrow- 
%  ^7SSy;^rfSS  m^.— There   are   two   types 
XdTsSjoir^f'digt'Y/S  of   adaptation  to  burrowing 
^^"^^^'•^  —that    of    the    Mole    and 

that  of  the  HeterocepJialus. 

In  the  mole  (fig.  35),  the  fore-limb,  although  it 
has  become  shorter  in  order  that  it  may  better  fulfil 


^  L.  Dollo,  Premiere  note  sur  les  Mosasauriens  de  Mesvin  {Bull. 
Soc.  helg.  04ol.  PaUonL  HydroL,  vol,  iii.,  1889)  ;  Nouvelle  note 
sur  V ost6ologie  de  Mosasauriens  [Bull.  Soc.  helg.  Gdol.  Pal^ont. 
HydroL,  vol.  v.,  1892). 


TRANSFORMATION  OF  ORGANS  OF  ANIMALS        67 


its  function,  is  obviously  in  process  of  development, 
for  it  has  not  only  retained  all  five  digits,  but,  what 
is  functionally  a  sixth  digit,  has  made  its  appear- 
ance (rs.).  The  phalanges  are  very  well  developed 
(phal.),  but,  on  the  other  hand,  the  distal  phalanges 
(ph.)  have  degenerated  and  are  very  short. 

Heterocephalus  (fig.  36)  is  a  burrowing  rodent, 
only,  instead  of  being  talpoide  as  is  the  Bathyergiis, 
i.e.   instead    of   exhibiting  a   body   furnished   with 


Fig.  Z&.—Heterocephalus  Philippi.    (After  Oldfield  Thomas.) 

strong  short  legs,  Heterocephalus  looks  more  like 
an  ordinary  quadruped.  In  process  of  adaptation 
to  an  underground  life,  the  hair,  especially  that  on 
the  legs,  has  disappeared.  This  is  the  more  re- 
markal)le,  as  in  no  other  case  does  it  occur  among 
mammals  unless  as  an  adaptation  to  aquatic  life. 
Here  it  is  in  the  atrophy  of  the  roots  of  the  hair 
that  degeneration  is  manifested. 

One  form  of  hair,  however,  is  still  to  be  found 
on  Heterocephalus y  for  there  are  a  few  bristles  on 
the  outer  sides  of  the  feet.  These  serve  as  brushes 
to  sweep  away  the  sand  while  burrowing. 


68 


UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


Section  II. 
Modification  of  the  organs  of  plants. 

§  4.  Modification  of  homodynamic  organs  in  the 
Individual. 


For    the 
pointed 


m 


same  reasons  as  we  have  already 
the  case  of  vertebrates,  those  leaves 
which  grow  out  of 
different  parts  of  the 
main  stem,  and  have 
undergone  many 
modifications  in 
order  to  adapt  them- 
selves to  their  vari- 
ous functions,  will 
best  serve  as  ex- 
amples for  our  pre- 
sent demonstration. 
In  descriptive 
botanical  treatises 
the  term  "  leaf "  is 
applied  without  dis- 
crimination. As  a 
rule,    however,    this 

term  should  only  be  applied  to  the  true  leaves,  i.e. 

those    of    which    the    parts    are    fully    developed 

throughout.^ 

^  The  embryological  development  of  the  leaf  among  the  Angio- 
sperms  (fig.  37)  exhibits  the  following  phases  : — The  leaf  arises 
from  the  growing  point  in  the  form  of  a  small  bud,  which  grows 


Fia.  37. — Growing  point  of  Vicia  varia, 
showing  the  origin  of  leaves. 
1,  2,  3,  4,  leaves  respectively  older;  leaf  1  is 
still  in  the  primordial  condition ;  leaf  2  is 
divided  into  epipodium  and  hypopodium  (Ji). 
In  leaves  3  and  4  the  epipod  is  divided  into 
petiole  and  leaflets,  and  the  hypopod  fornis 
stipules. 


"  MODIFICATION  OF  THE  ORGANS  OF  PLANTS    69 

These  foliage  leaves,  which  are  solely  assimilative, 
are  philogenetically  the  oldest,  for  the  assimilative 
function  of  leaves  was  certainly  an  earlier  function 
than  any  they  may  now  exercise ;  but  it  is  not  un- 
common to  find  both  basilar  {NiederUdtter)  and 
apical  leaves  {HoclibloMer)  as  well  as  foliage  leaves 
{Laubhldtter)  in  the  same  plant,  and  even  growing 
from  the  same  branch.  These  leaves,  which  arise 
directly  from  the  foliage  leaves,  have  more  or  less 
completely  lost  their  primitive  function,  and  have 
assumed  others.  Their  structure  has  undergone 
corresponding  modifications  which  are  all  attended 

for  some  time  without  any  evidences  of  differentiation,  and  con- 
stitutes what  is  called  the  primordial  leaf.  Next  the  leaf  is 
differentiated  into  a  proximal  part,  which  almost  surrounds  the 
stem  (the  hypopod),  a  distal  part  (the  epipod),  and  an  inter- 
mediate part  (the  mesopod).  From  this  point  the  various  parts 
of  the  leaf  begin  to  develop  individually. 

Little  foliacious  lamina  (the  stipules)  frequently  arise  laterally 
from  the  hypopod,  and,  while  still  in  the  bud  these  stipules,  being 
often  much  larger  than  the  rest  of  the  leaf,  serve  as  a  very  effective 
protection  to  the  young  organs. 

The  way  in  which  the  epipod  differentiates  varies  very  much  in 
different  cases.  The  lamina  is  developed  from  it,  and  little  buds 
are  gradually  formed  around  the  point  until  the  whole  is  ready  to 
branch  out.  At  this  stage  the  young  leaf  is  still  rolled  up  within 
itself,  and  protected  by  stipules,  when  there  are  any. 

Next,  the  mesopod  grows  into  the  petiole,  and  the  gi-owth  of  the 
petiole  causes  the  separation  of  the  different  parts  of  the  leaf. 

The  leaves  of  many  plants  are  much  less  complicated  than 
these.  Sometimes  the  mesopod  is  missing,  and  the  hypopod  and 
the  epipod  are  left  in  contact.  Sometimes  there  is  practically  no 
hypopod,  and  in  some  cases  the  primordial  leaf  develops  with- 
out any  differentiation  into  hypopod  and  epipod.  See  fig.  49 
{Sempervivum  arachnoideum)* 


70       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


by  degeneration.  A  good  demonstration  of  this  will 
be  to  examine  the  leaves  of  the  following  types  : 
Rosa  rngosa,  Serratula  centauroides,  Sagittaria  sagitti- 
folia,  Lathyrus  Aphaca,  and  Nymphaea  dentata} 

1.  Rosa  rugosa. — In  the  foliage  leaves  (fig.  38,  f) 
of  a  rose-branch,  such  as  of  Rosa  rvgosa,  there  are  two 


Fijj.  38.— Leaves  from  the  same  branch  of  Rosa  rugosa. 
A— E,  successive  basilar  leaves  ;  F,  foliage  leaf;  G— I,  successive  apical  leaves. 

lateral  stipules  which  arise  from  the  hypopod  and 
are  fused  with  the  base  of  the  petiole.  The  petiole, 
which  arises  from  the  mesopod,  has  from  two  to  six 
pairs  of  leaflets,  and  also  a  terminal  leaflet — all  of 
which  are  derived  from  the  epipod. 

^  In  studying  these  types  we  will  confine  our  attention  to  tlie 
leaves  arising  directly  from  the  foliage  leaves,  to  the  exclusion  of 
the  floral  leaves. 


MODIFICATION  OF  THE  ORGANS  OF  PLANTS        7l 


The  basilar  leaves  (fig.  38,  A  to  e),  which  are 
usually  withered  in  an  adult  branch,  have  more  or 
less  completely  lost  their  assimilative  function,  and 
merely  serve  as  a  protection  to 
the  bud.  The  hypopod  plays  an 
important  part  in  this  modifica- 
tion, but  there  are  evidences  of  a 
partial  degeneration,  the  epipod 
being  reduced  in  size  in  propor- 
tion as  the  assimilating  function 
is  lost.^ 

The  apical  leaves  also  have 
partially  lost  their  assimilative 
function,  and  have  assumed,  like 
the  basilar  leaves,  a  protective 
function  which  they  exercise  on 
the  floral  buds.  This  modification 
is  effected  differently  to  that  of 
the  basilar  leaves,  but  is  equally  ^'/utSlTlikan^fve?^ 
attended  with  evidences  of  degener-  ^'^™«*  °^  '^^  '"^«"^*='^«- 
ation.  The  hypopod  continues  to  gain,  as  the  epipod 
loses,  in  importance,  but  the  leaflets,  instead  of  de- 


FiG.  39. — Leaves  from  the 
same  branch  of  Serra- 
tula  centauroides. 


^  On  examining  the  foliage  leaves  of  a  plant  from  above  down- 
wards, one  finds  successively  leaves  with  large  stipules  and  small 
leaflets  (fig  38,  e),  then  leaves  with  very  small  leaflets  crowded  at 
the  end  of  the  hypopod  and  which  have  only  the  basilar  part  of  the 
petiole  (fig.  38,  d),  and  then  leaves  of  which  the  epipod  and  the 
free  part  of  the  petiole  have  disappeared  (fig.  38,  c).  Finally,  at 
the  base  of  the  plant  there  are  leaves  in  which  the  hypopod  is 
markedly  reduced,  and  which  carry  nothing  but  small  stipules 
(fig.  38,  A  and  b). 


72        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


creasing  in  size,  decrease  in  number.  At  first  there 
may  be  three  leaflets,  and  then  only  a  single  large 
leaflet  (fig.  38,  g). 
Finally,  towards  the 
floral  end  of  the 
branch  the  leaflet  be- 
comes smaller  and 
smaller  until  it  dis- 
appears altogether 
(fig.  38,  H  and  i). 
The  apical  leaves,  like 
the  basilar  leaves,  are 
eventually  reduced 
simply  to  the  hypopod. 

2.  Serratula  cen- 
tauroides.— The  foli- 
age leaves  of  this 
species  are  close  to 
the  ground,  their  bases 
are  ensheathed,  and 
the  long  petioles  ter- 
minate in  rhomboidal 
blades. 

In  passing  on  from 
these  assimilative 
leaves  to  the  basilar 
and  apical  leaves,  a 
double  adaptation  becomes  apparent,  as  in  Hosa 
Tugosa.  In  Rosa  rugosa,  however,  both  adaptations 
were  protective,  while  in  Serratula  centauroides  this 


Fig.  AQ.—Sagittaria  sagittifolia. 
A,  winter  bud  with  membraneous  leaves  en- 
closing the  swollen  portion  and  ro  led  leaves 
protecting  the  bud ;  B,C,  submerged  leaves; 
D,E,  floating  leaves;  F,  aerial  foliage  leaf; 
G,  leaf  protecting  the  flower-bud ;  H, 
seedling  grown  from  a  seed  planted  in  the 
water  on  the  surface  of  the  mud.  I,  seed- 
ling from  seed  planted  in  ihe  water  under 
a  layer  of  mud.  In  flgures  H  and  I,  c  is 
the  cotyledon,  I  the  first  leaves. 


MODIFICATION  OF  THE  ORGANS  OF  PLANTS        73 

is  ouly  the  case  with  the  basilar  leaves,  which  sur- 
round the  winter  buds. 

The  modifications  of  the  basilar  and  apical  leaves 
of  Serratula  centauroides  are  similar  to  those  of  Bosa 
rtcgosa,  and  it  is  therefore  unnecessary  to  repeat  the 
description. 

Those  of  the  apical  leaves  tend  towards  their 
protection  from  rain.  When  rain  falls  upon  the 
radical  leaves,  they  are  flattened  out  upon  the 
ground,  which  prevents  them  from  being  torn. 
This  is  not  so  with  the  apical  leaves,  and  had 
their  primitive  structure  remained  unchanged,  they 
would  have  been  constantly  exposed  to  destruction. 
M.  Stahl^  has  shown  how  these  leaves  have  pro- 
tected themselves  by  shortening  their  petioles  and 
fringing  their  edges.  In  this  way  the  leaf  has 
acquired  greater  resistance,  and  the  water  is  better 
enabled  to  run  off  it.  This  modification,  however, 
although  one  advantageous  to  the  plant,  has  in- 
volved the  partial  degeneration  of  the  petiole  and 
the  blade.2 

3.   Sagittaria  Sagittifolia. — The  Sagittaria  is  an 

*  E.  Stahl,  Regenfall  und  Blattgestalt  {Ann.  du  Jard.  hot.  dc 
Buitenzorg,  vol.  xi.,  1893). 

2  In  the  leaves  (fig.  39,  b  and  c),  immediately  above  the  foliage 
leaves,  the  petiole  gradually  disappears  altogether,  while  the  blade 
has  become  deeply  serrated,  thus  providing  for  the  water  drainage 
while  retaining  an  assimilative  surface. 

Further  up  the  stem  the  leaf  blades  are  more  and  more  reduced, 
until  eventually  they  are  no  longer  differentiated  into  hypopod  and 
epipod,  but  arise  directly  from  the  growth  of  the  primordial  leaf 
(fig.  39,  D  to  F). 


74       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

aquatic  plant,  the  foliage  leaves  of  which  emerge 
from  the  water.  The  blade  of  these  leaves  (fig. 
40,  f)  is  shaped  like  an  arrow-head,  and  the  barbs 
facilitate  rain-water  drainage.  The  petiole  is  very 
long,  and  terminates  in  a  sheathed  base  derived 
from  the  hypopod. 

The  apical  leaves  (fig.  4,  g)  are  represented  by 
transparent  membraneous  scales,  which  serve  as  a 
protection  to  the  floral  buds. 

The  basilar  leaves,  which  either  float  on  the 
surface  of  the  water  or  else  are  entirely  sub- 
merged, have  undergone  both  development  and 
degeneration.  Degeneration  is  more  noticeable  on 
glancing  downwards  from  the  foliage  leaves  to 
those  which  protect  the  winter  buds.^ 

*  {a)  The  leaves  nearest  to  the  foliage  leaves  float  upon  the 
surface  of  the  water  (fig.  40,  e).  The  lateral  barbs, 
being  no  longer  required,  have  begun  to  degenerate. 

(&)  The  floating  leaves  immediately  beneath  exhibit  barbs 
which  are  still  more  reduced  (fig.  40,  d),  and  there  are 
no  stomata  except  those  upon  the  upper  surface. 

(c)  The  next  leaves  are  completely  submerged  and  lengthened 
out  into  long  ribbons  which  broaden  into  flat  blades 
at  their  upper  ends,  which  are  not  separated  from  the 
hypopod  by  petioles  (fig.  40,  c).  In  this  case  the  blade 
is  very  much  smaller,  and  the  petiole  has  disappeared. 

{d)  The  next  leaves  to  these  are  submerged,  and  are  ribbon- 
like in  shape.  They  spring  immediately  from  the 
primordial  leaf  without  differentiating  into  hypopod, 
mesopod,  and  epipod. 

(c)  After  these  leaves  follow  leaves  consisting  solely  of  a  hypo- 
pod, which  is  much  reduced  in  size  (fig.  40,  A).  These 
leaves  are  folded  round,  and  serve  as  a  protection  to  the 
bud. 


MODIFICATION  OF  THE  ORGANS  OF  PLANTS        75 

Lathyrus  Aphaca. — The  foliage  leaf  (fig.  4,  leaves 
4  and  5),  which  most  nearly  resembles  the  primi- 
tive type,^  consists  of  two  stipules  and  a  petiole 
which  terminates  in  a  point  and  is  provided  with  a 
pair  of  lateral  leaflets. 

Each  individual  plant  is  furnished  with  from  gne 
to  three  of  these  foliage  leaves. 

Above  them  are  leaves  (fig.  41,  leaf  6)  consisting 
of  only  a  hypopod  which  forms  two  large  stipules 
that  partially  enclose  a  small  point  arising  from 
the  epipod.  The  entire  function  of  assimilation  is 
exercised  by  these  stipules,  and  consequently  both 

(/)  Finally,  there  are  some  scaly  leaves  which  also  consist 

merely  of  a  hypopod.     These   protect  the  tubers  (fig. 

40,  a),  and  are  practically  devoid  of  chlorophyll.     These 

various  leaves  follow  one  another  in  the  course  of  the 

growth  of  the  plant,  in  the  inverse   order  to   that  in 

which  they  are  here  described.     As  Avinter  approaches, 

some  upright  stems  appear,  the  inflated  ends  of  which 

are   stored  with   reserve  nutrition   (fig.    40,    a).     It  is 

round  these  tubers  that  the  scaly  leaves  are  to  be  found. 

The  bud  which   does  not  develop  until   the   following 

Spring  is  at  the  top,  surrounded  by  folded  leaves.     The 

ribbon-like  leaves  which  grow  under  water  and  also  those 

broadened  out  at  the  ends  make  their  appearance  in  the 

course  of  the  Summer.     Next  come  the  floating  leaves 

with  slightly  developed  barbs,  then  the  leaves  which  rise 

well  above  the  surface  of  the  water,  and  finally  the  scaly 

leaves  which  grow  out  of  the  flowering  stalk. 

^  The  foliage  leaves  of  Lathyrus  Aphaca  may  be  regarded  as  the 

best  representatives  of  the  primitive  leaf,  for  in  most  species  of 

the  genus  Lathyrus,  each  leaf  is  furnished  with  leaves  and  with 

one  or  more  tendrils.     L.  ApJiaca  is  probably  specialized  from 

other  species,  for  the  majority  of  the  leaves  have  lost  their  lateral 

leaflets. 


76        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


the  petiole  and  the  lateral  leaflets  have  atrophied. 

Higher  up  on  the  stem 
(fig,  41,  leaves  7  and  8) 
are  some  leaves  consisting 
of  two  stipules  and  a  non- 
branching  tendril.  The 
^Sl^  latter  represents  the  meso- 
pod  and  the  epipod  from 
which  the  lateral  leaflets 
have  disappeared.  Below 
the  foliage  leaves,  there 
are  usually  three  leaves 
which  have  considerably 
degenerated  (fig.  41,  leaves 
1  to  3).  The  hypopod 
is  represented  by  two 
very  small  stipules,  and 
the  epipod  by  the  tiny 
point  in  the  centre.  The 
portion  of  the  stem  out 
of  which  the  basilar  leaves 
would  grow  does  not, 
under  normal  conditions, 
emerge  from  the  soil, 
which  accounts  for  the 
way  in  which  these  leaves 
have  degenerated. 

5.  Nympli<Xea  dentata. 
— Here  the  foliage  leaves 
are  large,  floating,  fleshy. 


Fig.  41. — SecdliiiK  of  Lathyrus 

Aphaca. 

1,  2,  3,  very  rudimentary  leaves ;  4,  6. 

foliage    leaves;    6,    leaf    composed 

only  of  a  pair  of  stipules.     7,  8, 

terminal  leaves  turned  into  tendrils. 


MODIFICATION  OF  THE  ORGANS  OF  PLANTS    77 


and  toothed  at  the  edges.  During  germination  the 
seed-leaves  remain  within  the  seed  as  with  Lathyriis 
Aphaca. 

The  first  leaf  is  upright  and  reed-like  in  shape 
(fig.  42,  A  1).  Its  principal  function  is  to  pierce 
the  layer  of  soil  which 
covers  the  grain,  thus 
allowing  the  little  ter- 
minal bud  to  emerge 
into  the  light.  Com- 
pared with  the  other 
leaves,  this  needle-like 
leaf  seems  to  have  con- 
siderably degenerated. 
The  plant  next  pro- 
duces a  series  of  little 
slender  short-stemmed 
leaves  which  grow 
under  the  water.  The 
first  of  these  leaves 
are  narrow  and  ribbon- 
like (fig.  42,  A  2);  the 
next  are  broader  (fig. 
42,  b),  then  follow 
more  slender  sub- 
merged leaves,  but  with  no  lateral  barbs  (fig.  42,  c) ; 
then  floating  leaves,  the  edges  of  which  are  not 
dentated,  and  last  of  all  there  are  some  leaves  with 
dentated  edges.  Owing  to  their  submergence,  the 
basilar  leaves  of  Nymphaea  have  greatly  degener- 


Fig.  42. — Nymphaea  dentata. 
A,  seedling  with  two  leaves  (1,  2)  ; 
leaves  taken  fi'om  older  plants. 


B,  C, 


78       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

ated ;  the  blade  is  very  thin,  being  composed  of  a 
few  layers  of  cells,  and  there  are  no  stomata. 

The  basilar  leaves  and  the  apical  leaves  fre- 
quently lose  either  partially  or  entirely  the  as- 
similative function  which  they  previously  exercised. 
Some  of  these  leaves  serve  as  a  protection  to  the 
buds  of  both  leaves  and  flowers.  Others  are 
adapted  as  protection  from  rain,  and  others  have 
undergone  considerable  modifications  owing  to  their 
existence  either  under  water  or  under  ground. 

In  all  these  instances  of  modification  it  can  be 
seen  that  some  degeneration  has  invariably  attended 
each  change  which  has  taken  place. 

§  5. — Modification  of  organs  which  are  homologous 
in  individuals  of  different  species. 

Having  investigated  the  modifications  of  leaves 
in  the  individual,  we  will  now  give  our  exclusive 
attention  to  the  various  modifications  undergone 
by  the  foliage  leaves  ;  we  shall  see  among  the 
various  species  we  examine  that  degeneration  has 
played  a  part  in  each  instance  of  modification.    • 

Of  these  adaptations  the  following  are  the  most 
characteristic :  adaptation  to  climbing,  to  a  carni- 
vorous diet,  to  aquatic  life,  to  defence  against  ants, 
against  drought,  and  against  herbivorous  animals. 

1.  Adaptation  to  climbing.  —  Climbing  plants 
attach  themselves  by  means  of  tendrils  to  the 
nearest  support  within  their  reach,     These  tendrils, 


MODIFICATION  OF  THE  ORGANS  OF  PLANTS        79 

which  are  thread-like  and  sensitive  to  contact,  are 


Fig.  43. — Cobaea  scandens. 
A,  B,  seedlings ;  the  leaves  of  the  second  end  in  tendrils ; 
udult  plant. 


C,  young  leaf  of 


modifications  of  either  stems,  leaves,  or  roots,  the 
latter  being  the  most  rare. 
We  will  confine  our  atten- 
tion at  present  to   those 
adapted  from  leaves. 

In  Cobaea  scandens  the 
first  two  leaves  of  the 
seedling  usually  terminate 
in  a  leaflet  (fig.  43,  a)  ; 
the  leaves  which  come 
after  are  much  finer  and 
are  modified  into  tendrils 
(fig.  43,  c). 

This  modification  of 
leaflets  into  tendrils,  which 
entails  the  almost  com- 
plete disappearance  of  the  original  assimilative  func- 


FiG.    44. — Basal  portion  of  a  young 
plant  of  Vicia  Pyrenaica. 


80       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

tion  of  the  leaflet,  is  by  no  means  uncommon.  Vicia 
Pyrenaica  (fig.  44)  furnishes  a  good  example  of 
this  change,  of  which  the  degeneration  of  the  blade 
is  the  necessary  consequence. 

In  Cucumis  sativus  (the  cucumber)  some  of  the 
leaves  are  the  shape  of  ordinary  assimilative  leaves, 
while  others  are  entirely  modified  into  tendrils  ;  in 
the  case  of  the  latter  the  blade  has  completely 
degenerated. 

2*  Adaptation  to  carnivorous  nutrition. — Some 
plants,  instead  of  obtaining  nutriment  exclusively 
from  minerals  and  carbon  dioxide,  are  capable  of 
assimilating  animal  matter  from  insects  and  other 
small  organisms,  which  they  capture  by  means  of  a 
special  function  exercised  by  the  leaves. 

(a)  Utricularia. — In  the  aquatic  Utricularia  the 
leaves  are  minutely  sub-divided  like  many  other 
submerged  plants.  The  leaves  of  Utricularia 
vulgaris  exhibit  leaves  of  this  type.  Some  of 
their  ramifications  carry  pouches,  which  serve  for 
the  capture  of  minute  organisms.  These  pouches 
are  formed  by  the  modification  of  part  of  the 
blade,  and  have  almost  completely  lost  their 
chlorophyll. 

Utricularia  intermedia  (fig.  45)  exhibits  two 
kinds  of  branches ;  the  one  kind  stretches  out 
horizontally  and  carries  green  leaves ;  the  others 
are  devoid  of  chlorophyll,  and  the  leaves  are 
purely  carnivorous. 

(6)  Nepenthe. — The  distal  extremity  of  the  leaf 


MODIFICATION  OF  THE  ORGANS  OF  PLANTS 


81 


terminates  in  a  receptacle  of  complicated  structure 
which  is  adapted  for  the  capture  of  carnivorous 
nutriment.  Degeneration  has  attended  this  change, 
for    the    modified    portion   of    the    leaf   is    almost 


Fig.  ^b.—Utricularia  intermedia  (after  Goebel,  Pflanzenbiologitche  Sehilder- 
ungen,  vol.  ii.,  p.  136). 
The  plant  has  grown  from  a  winter  bud  of  which  the  remains  are  visible  to  the 
left.    The  horizontal,  assimilating  branch  bears  two  pendulous  twigs  with 
carnivorous  urns. 

entirely  without  chlorophyll.  A  carnivorous  diet, 
however,  for  these  plants,  is  only  supplementary  to 
the  typical  plant  nutrition,  and  therefore  the  assimi- 
lative function  remains  active. 


82       UNIVERSALITY  OF  DEGENERATIVE   EVOLUTION 

In  Utricularia  this  function  is  exercised  by  the 
leaves  (as  in  U.  intermedia),  or  by  certain  parts 
of  the  leaves  {U.  vulgaris).  In  Nepenthe,  however, 
another  and  different  assimilative  organ  is  exhibited. 
This  consists  of  two  lateral  herbaceous  growths,  which 
are  not  formed  from  the  blade,  but  arise  from  the 
petiole.  In  between  the  receptacle  and  the  en- 
larged portion  of  the  petiole  is  a  part  in  which 
the  petiole  has  assumed  the  function  of  a  tendril. 
While  the  receptacle  is  in  process  of  formation,  this 
tendril  twines  itself  round  a  support  in  order  to 
obtain  the  additional  strength  that  will  be  required 
later  on  to  uphold  the  receptacle  when  full  of  diges- 
tive juice. 

(c)  Drosera. — The  leaves  of  the  sundews  (Drosera 
rotundifolia,  D.  longifolia,  etc.)  are  furnished  with 
a  great  number  of  emergences,  each  of  which  ter- 
minates in  a  digestive  gland.  There  is  a  large 
drop  of  sticky  fluid  at  the  end  of  each  gland, 
and  these  'drops,  which  sparkle  in  the  sunlight, 
have  given  the  plant  its  name  {Bos  solis  —  sundew) 
(fig.  46). 

These  emergences  contain  very  little  chlorophyll. 

"  A  plant  of  Drosera,  with  the  edges  of  its  leav^es  curled 
inwards  so  as  to  form  a  temporary  stomach,  with  the  glands 
of  the  closely-inflected  tentacles  pouring  forth  their  acid 
secretion,  which  dissolves  animal  matter  afterwards  to  be 
absorbed,  may  be  said  to  feed  like  an  animal." 

— Darwin,  Insectivorous  Plants,  p.  18. 

This    adaptation    to    a    carnivorous    diet    has    not 


MODIFICATION  OF  THE  ORGANS  OF  PLANTS 


83 


come  about  without  a  certain  amount  of  conse- 
quent degeneration.  The  chlorophyll  has  dis- 
appeared, excepting  a  small  quantity  exhibited  in 
the  upper  and  under 
surfaces  of  the  blades 
of  the  leaves,  in  the 
flower  stalks,  in  the 
central  tentacles,  and 
in  the  petioles. 

3.  Adaptation  to 
an  aquatic  life, — The 
adaptation  of  leaves, 
particularly  of  sub- 
merged leaves,  to  an 
aquatic  life  entails 
important  changes, 
and  these  changes 
are  invariably  at- 
tended by  degenera- 
tion. 

1.  Owing  to  the 
buoyancy  of  water, 
submerged  leaves  are 

Tint        Hpnpnrlpnl-        art  fiG.  4G.— Leaf  of  Drosera  longi/olia. 

not        uepeuutJD  l        on  ^ear  the  top,  an  insect  is  in  the  act  of  being 

nnv  hiahl V   rlpVplnnpH  capt^^ed  and  the  suckers  are  bencUng  over 

any  nigniy   ueveiopeu  jt     Lowtr  down  is  an  insect  reduced  to  a 

organic     support,    and  c^^itonous  skeleton. 

the  fibres  and  other  thick-walled  cells  have  con- 
sequently degenerated  to  a  great  extent. 

2.  There  being  no  transpiration   in.  submerged 
leaves     and    very    little    in    floating    leaves,    the 


84       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

former  are  very  seldom  provided  with  stomata, 
and  the  latter  only  exhibit  stomata  on  their 
upper  surfaces  (see  page  26,  note  on  Ranunculus 
and  Stratiotes).  The  conducting  apparatus  too,  i.e. 
the  wood  and  the  roots,  has  greatly  diminished- in 
size  owing  to  the  diminution  of  transpiration. 

3.  Very  little  light  penetrates  to  submerged 
leaves,  and  oxygen  is  only  partially  soluble  in 
water.  The  blade  has  therefore  become  much 
smaller.  Very  little  importance,  however,  is  at- 
tached to  this  last  modification,  as  the  size  of  the 
blade  varies  greatly  in  different  specimens,  whereas 
the  other  modifications  are  similar  in  all. 

{a)  In  the  species  already  mentioned — Sagittaria 
(p.  72)  and  Nympliaea  (p.  76) — the  first  leaves 
which  are  formed  are  submerged,  with  petioles  either 
short  or  entirely  absent,  and  very  slender  blades. 
In  Vallisneria  and  many  species  of  Potamogeton  there 
are  only  slender  submerged  leaves. 

All  these  leaves  augment  their  surface  contact 
with  the  water  by  ridding  themselves  of  the  deeper 
layers  of  assimilative  cells.  Very  little  light  could 
penetrate  to  these  cells,  and  any  oxygen  formed 
would  be  difficult  to  get  rid  of. 

(b)  In  other  plants,  especially  in  the  Dicotyledons, 
another  kind  of  modification  takes  place  for  the 
same  end ;  the  blade  is  divided  into  very  thin  seg- 
ments which  in  some  cases  present  a  hair -like 
appearance. 

The    Banunculus   is    a    good    example    of    this. 


MODIFICATION  OF  THE  ORGANS  OF  PLANTS        85 

Besides  the  many  species  found  on  moors  and  culti- 
vated land  and  in  woods,  there  are  other  kinds 
of  Ranunculus — B.  sceleratus,  for  instance — which 
grow  by  the  water-side.  In  this  species  it  is  not 
uncommon  to  find  floating  leaves,  and  leaves  which 
wholly  emerge  from  the  water,  growing  on  the  same 
stem.  In  other  species,  such  as  Banunculus 
hederaceus,  there  are  seldom  any  but  floating  leaves. 
Among  the  numerous  varieties  of  Banunculus  aqua- 
tilis  there  are  some  which,  in  addition  to  the  floating 
leaves,  have  also  some  leaves  which  are  completely 
submerged  and  deeply  dentated ;  in  other  varieties 
of  the  same  species  there  are  only  slightly  dentated 
leaves,  while  in  Banunculus  fluitans  the  leaves  are 
all  fringed.^ 

{c)  In  a  submerged  plant  which  is  to  be  found  in 
Madagascar,  Oitmrandra  fenestralis  (fig.  47),  the 
diminution  in  volume,  compared  to  the  surface,  is 
effected  in  a  different  manner.  After  the  leaf  has 
developed  in  the' ordinary  way,  all  the  parts  of  the 
leaf  in  between  the  veins  of  the  blade  disappear. 
This  results  in  the  blade  being  reduced  to  a  delicate 
piece  of  network,  consisting  entirely  of  communicat- 
ing veins  accompanied  by  adjacent  assimilative  cells. 

4.  Adaptation   to   defence   against   ants. — In  the 

1  Ranuncfiilus  sceleratus  grows  by  the  water-side  ;  R.  hederaceus 
is  to  be  found  on  marshy  ground,  the  intern  odes  and  petioles  being 
quite  short ;  11.  aquatilis  grows  in  shallow  water,  and  the  internodes 
are  of  sufficient  length  to  raise  the  flowers  out  of  the  water ;  R. 
jluitans  gi'ows  in  strongly-running  streams,  so  that  floating  leaves 
would  be  useless. 


MODIFICATION  OF  THE  ORGANS  OF  PLANTS 


87 


tropics,  especially  in  South  America,  plants  are 
much  exposed  to  the  ravages  of  leaf- eating  ants 
(Atta).  In  order  to  protect  themselves,  they  provide 
other  kinds  of  ants  with  shelter  and  nourishment, 
and   this  necessarily  entails  several  modifications. 


Fio.  48.--Part  of  a  leaf  of  Acacia  sphaerocephala  (after  Schimper,  Die  Wechsel- 
beziehungen  zwischen  Pflanzenund  Ameisen  im  tropischen  Amerika.  Fasc.  1 
of  Schimper's  botanische  Mittheilungen  aus  den  Tropen.    Jena,  1888), 

In  Acacia  sfphaerocepliala  (fig.  48),  for  instance,  the 
blades  of  the  leaves  are  bipinnated  and  the  proximal 
leaflets  terminate  in  small  cavities  filled  with  a  nutri- 
tive secretion.  The  stipules,  which  no  longer  contain 
chlorophyll,  are  modifiecl  into  hollow  thorns  in  which 


88       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


the  ants  live  and  obtain  nutriment,  not  only  at  the 

ends  of  the  leaflets,  but  also  in  a  thorny  gland  which 

is  situated  upon  the  petiole. 

5.  Adaptation  to  drought 
—  In  places  where  rain 
seldom  falls,  plants  are 
provided  with  natural 
reservoirs  of  water.  These 
reservoirs  are  situated 
either  in  the  roots  or  the 
stems  and  occasionally  in 
the  leaves.  Where  the 
leaves  are  fleshy,  these 
reservoir  leaves  are  gener- 
ally very  simple  in  forma- 
tion. Those  of  Sempervi- 
vuin  and  of  several  other 

genera    arise    directly   from    the    primordial,   non- 
differentiated    leaf, 

and    there    is    no 

differentiation  into 

hypopod,       epipod 

and     petiole     (fig. 

49).      See  further 

on  (fig.  73),  p.  236 

(Sempervivum    tec- 

tOTUTR^  Fig.  50. — Branch  of  Caragana. 

6.  Adaptation  to  defence  against  herbivorous 
animals. — Thorns  are  an  adaptation  which  serve  as 
a   protection   against    herbivorous    animals.     They 


Fig.  49.— Growing  point  of  Semper- 
vivum arachnoideum. 


MODIFICATION  OF  THE  ORGANS  OF  PLANTS 


89 


are  derived  from  the  modifications  of  various  parts 
of  the  plant,  either  the  roots,  stems,  leaves,  or 
even  of  the  floral  stems. 

This  modification  is  always  accompanied  by  some 
degeneration  ;  the  thorn,  which  is  quite  hard,  ex- 


TiG.  51.— Mamillaria  elephantident  (after  Lemaire?     The  figure  is  taken  from 
Goebel,  Pflanzenbiologische  Schilderungen,  vol.  i.,  p.  71). 

cepting  at  the  point,  is  made  up  of  thick-walled 
cells  from  which  the  protoplasm  has  disappeared. 

Leaf  thorns  arise  from  the  modification  of  various 
parts  of  the  leaves — either  of  the  stipules  (as  in 
Cciragana)^  the   petioles   {id.)   (fig.    50),  or   of   the 


90        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

blade  (as  in  Ilex).  In  each  case  the  leaf  par- 
tially retains  its  assimilating  function  and  its 
chlorophyll. 

In  other  plants,  however,  and  especially  in  Cacti, 
and  the  fleshy-leaved  Euphorbias,  the  leaves  exhibit 
further  evidences  of  degeneration,  their  function 
being  exclusively  one  of  defence  (fig.  51), 


CHAPTEE  II 

IN    THE    EVOLUTION    OF    INSTITUTIONS    ALL    MODIFICA- 
TION   IS    NECESSARILY    ACCOMPANIED    BY 
DEGENERATION 

The  distinction  we  have  drawn  between  the 
homodynamic  organs  of  an  organism,  and  the  homo- 
logous organs  of  organisms  belonging  to  other 
species,  is  not  applicable  in  sociology,  as  we  have 
already  pointed  out  in  the  introduction.  Institu- 
tions, however,  may  be  regarded  from  two  distinct 
standpoints — from  a  statical  point  of  view,  as  they 
exist  in  tlie  same  society,  and  from  the  dynamical 
point  of  view,  as  existing  from  epoch  to  epoch,  and 
from  society  to  society.  In  both  cases,  we  shall 
arrive  at  the  same  conclusion  as  in  biology,  that 
all  modification  entails  degeneration. 

In  order  to  demonstrate  this  we  will  examine 
in  succession  the  modifications  undergone  by  tlie 
principal  types  of  financial  organizations  now  exist- 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    91 

ing  in  Europe,  and  the  most  important  stages  in 
the  evolution  of  landed  property  amongst  various 
peoples. 

§  1. — Modifications  of  similar  institutions  in  the 
same  society. 

The  financial  organization  of  European  towns 
and  states  has  undergone  very  important  changes 
since  the  middle  ages.  Taxes  and  duties  have 
attained  a  now  universal  importance,  as  substitution 
for  the  revenue  from  crown  lands,  which  constituted 
the  principal,  if  not  the  only,  resources  of  the 
sovereigns  of  the  feudal  ages.^ 

*  There  are  three  stages  in  the  evohition  of  tinancial  systems  in 
countries  (such  as  England,  for  instance)  where  the  question  of 
finance  has  been  most  successfully  dealt  ^Wth. 

1.  The  Feudal  System,  wherein  the  king  had  no  separate  revenue 

apart  from  the  nation,  and  wherein  the  revenue  of  the 
sovereign  was  principally  derived  from  crown  land,  the 
cultivation  and  administration  of  which  was  carried  on  as 
a  source  of  private  income  to  the  king. 

2.  The  Co-existence  of  the  Feudal  System  and  the  Modem  System, 

viz.,  the  disappearance  of  personally  held  land  and  its 
attendant  institutions,  the  development  of  the  public  pro- 
perty of  the  State  or  Township,  and  the  imposition  of 
duties  and  taxes. 

3.  The  Modern  System,   viz.,  the  complete  separation   of  the 

personal  property  of  the  sovereign  from  the  property  of 
the  nation,  the  increasing  importance  of  taxes  and  duties, 
and  the  almost  complete  disappearance  of  State  and  town 
lands.  Industries  taken  over  by  the  State — such  as  rail- 
ways, postal  and  telegraphic  arrangements,  etc. — and  by 
the  towns — such  as  gas,  water,  etc. — constitute  monopolies, 
and  are  no  longer  subjected  to  the  law  of  competition  which 
is  always  active  in  private  business.  — Wagnek. 


92        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

Here,  too,  we  find  that  all  modification  is 
attended  by  degeneration.  As  a  demonstration 
of  this,  we  will  examine  in  succession — firstly, 
the  Communal  budgets  of  Belgium  ;  secondly,  the 
State  budgets  of  the  States  which  compose  the 
German  Empire ;  thirdly,  the  budgets  of  Germany, 
England,  and  France,  regarded  as  belonging  to  one 
group — the  Western  Republic  of  Auguste  Comte. 

I.   The  Communal  Budgets  of  Belgium, 

Collective  property  in  its  archaic  form  still  exists, 
in  spite  of  considerable  modifications  in  certain 
parts  of  the  Ardennes,  of  the  Fagne,  and  of 
Lower  Luxembourg.^ 

At  Wanlin,  for  instance,  the  collective  property 
constitutes  nearly  one-half  of  the  land — 220  hec- 
tares, of  which  130  is  arable  land,  and  90  is  wood- 
land. The  arable  land  is  divided  into  allotments, 
apportioned  among  the  heads  of  the  various  families 
every  eighteen  years,  for  which  they  pay  an  annual 
rent  of  1 0  francs.  There  are  no  taxes  or  communal 
duties,  the  revenue  derived  from  this  public  property 
being  sufficient  without  them. 

This  economic  and  fiscal  system,  suitable  to  a 
scattered  and  agricultural  people,  existed  formerly 
throughout  the  whole  country,  but  the  increase  in 
population  and  cultivation  of  the  land  have  almost 

^  Paul  Errera,  Les  Masuirs.  Historical  and  judicial  researches 
in  the  vestiges  of  the  old  territorial  system  of  Belgium.  Brussels, 
Weissenbruch,  1891, 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    93 

put  an  end  to  this  archaic  condition  of  things. 
Private  property  has  encroached  upon  collective 
property,  and  taxes  and  duties  are  now  substi- 
tuted for  the  revenue  which  was  formerly  derived 
from  land  alone. 

This  modification  was  of  a  progressive  nature, 
the  extended  requirements  of  the  population  de- 
manding a  corresponding  augmentation  of  the  com- 
munal revenue.  Degeneration,  however,  is  exhibited 
in  the  disappearance  of  the  collective  land  of  the 
communes  and  of  the  attendant  administrative 
institutions. 

The  monographs  of  Paul  Errera  and  the  docu- 
ments collected  in  1875  by  the  Statistical  Commis- 
sion relating  to  the  expenditure  and  receipts  of  the 
communes,  demonstrate  the  various  stages  of  this 
degenerative  evolution : — 

1.  The    vestiges    of    the    old    system    may    be 

observed  to  decrease  in  number  and  im- 
portance on  approaching  the  North -West 
from  the  South.  The  proportion  of  re- 
ceipts from  land  compared  with  those 
derived  from  taxes  and  duties  is  greatest 
in  Upper  Belgium ;  then  follow  those  of 
Condroz,  Hainault,  Brabant,  Campine,  and 
finally,  of  the  two  Flanders. 

2.  The  constitution  of  the  two  Flanders  passed 

through  the  following  stages: — 
(a)  In  certain  localities  round  about  Bruges  the 
old  and  the  new  system  co-exist.     Side  by  side,  for 


94       UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

instance,  with  the  communes  of  Oedelem,  Beernem 
and  Oostcamp  we  find,  though  in  a  degenerate  con- 
dition, the  old  collective  property  of  the  aenhorgers 
of  Beverhoutsveld,  with  its  administrative  college  of 
Veldheeren} 

(h)  In  Sysseele,  and  in  several  other  communes, 
the  assembly  of  Veldheeren  has  been  abolished,  and 
the  property  which  they  administrated  has  become 
the  property  of  the  commune  by  whom  the  rents 
are  collected. 

(c)  At  Gand,  the  grazing  land  of  the  Heernisse  of 
Saint-Bavon,  which  was  declared  collective  property 
by  a  decree  of  November  16,  1887,  was  afterwards 

^  Beverhoutsveld  is  a  vast  domain  situated  outside  the  gates  of 
Bruges,  in  the  territory  of  the  commune  of  Oedelem.  According  to 
the  terms  of  a  decision  of  the  13th  August  1859,  it  constitutes  a 
collective  property  belonging  to  three  sections  of  the  communes  of 
Oedelem,  of  Beernem,  and  of  Oostcamp.  The  family  representatives 
of  these  three  sections  are  alone  privileged  to  rent  these  lands, 
and  the  revenue  accruing  from  them  is  divided  among  the  three 
sections. 

The  old  administrative  authority,  the  college  of  Veldheeren, 
which  was  in  existence  in  the  thirteenth  century,  still  exists, 
though  in  a  new  form.  According  to  the  terms  of  the  decision 
of  1859,  the  police  are  provided  for  from  the  taxes  of  Oedelem,  as 
the  property  is  within  that  district ;  "but  in  other  matters,  and 
in  the  administration  of  the  rights  and  revenues  accruing  from 
it,  the  property  shall  be  under  the  control  of  a  commissioner 
appointed,  so  far  as  possible,  by  the  interested  parties — i.e.  the 
aenborgers — in  conformity  with  the  laws  and  regulations  relating 
to  the  matter." 

The  Vrij-Geweid,  also  situated  in  the  environs  of  Bruges,  is 
similarly  administrated.  (See  Errera,  des  Masuirs,  chaps,  xix., 
XX.   and  following.) 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    95 

transformed  into  private  estates,  and  has  been  so 
built  over  as  to  become  almost  unrecognizable. 

{d)  In  some  towns  the  receipts  from  land  only 
represent  a  very  small  part  of  the  communal  revenue, 
3  per  cent,  at  Eoulers,  0*1  at  Saint-Nicolas.^ 

It  is  thus  evident  that  the  evolution  of  com- 
munal finance,  which  has  been  especially  char- 
acterized by  the  development  of  duties  and  taxes, 
has  been  accompanied  by  the  degeneration  of  the 
old  system  of  collective  property. 


II.  Bvdget  of  the  States  of  the  German  Empire. 

The  feudal  system  of  finance  has  left  its  mark 
upon  Germany,  especially  in  the  eighteen  subsidiary 
states  where  archaic  institutions  have  been  best 
preserved.^  The  principal  stages  in  the  progressive 
evolution  of  the  modern  system,  and  the  corre- 
sponding degeneration  of  the  feudal  system  may  be 
enumerated  as  follows  : — 

1.  Mecklembourg- Strelitz  still  exhibits  the  primi- 
tive system  in  a  comparatively  little  altered 
form.  The  Grand  Duke  is  the  sole  adminis- 
trator ;  the  budget  is  not  separated  from  the 
civil  list;  the  revenue  amounts  to  an  average 

^  Hector  Denis,  VImpCL,  pp.  55.     Brussels,  V®.  Monnom,  1889. 

2  Leroy-Beaiilieu,  TraiU  de  la  science  des  Finances,  1,  21,  and 
following. 

Adolf  Wagner,  Lehr-und  Handhuch  der  Politischen  Oekonomie  ; 
Finanzwisscnschaft.     Erster  Theil,  sees.  214-216. 


96        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

of  seven  millions,  of  which  five  are  derived 
from  land. 

2.  The  Duchy  of  Mecklembourg-Schwerin  is  ad- 

ministered partly  on  the  modern  and  partly 
on  the  feudal  system,  but  the  former  pre- 
dominates. There  are  two  public  budgets. 
That  of  the  Duke  is  of  the  greater  im- 
portance, and  amounted  to  14,500,000 
marks  in  1887,  of  which  7,000,000  were 
derived  from  landed  property.^  The  second 
budget  is  that  of  the  municipal  expenditure 
(Gemeinsamen  Finanzverwcdtung),  which  is 
defrayed  by  means  of  rates  and  taxes,  the 
receipts  of  which  amounted  in  1887  to 
4,175,000  marks. 

3.  In  the  larger  States  pf  the  Empire  the  feudal 
system  has  completely  disappeared,  but  the  revenue 
obtained  from  collective  property  forms  an  im- 
portant part  of  the  public  income. 

Bavaria       .  .  .        17-3  % 

Wiirtemberg  .  .        13-2  % 

Saxony        .  .  .          9-7  % 

Baden          .  .  .          7*1  % 

Prussia        .  .  .          8'4  % 

4.  In  the  budget  of  the  Empire,  which  is  of 
recent  construction,  naturally  no  traces  of  the  feudal 
system  remain. 

^  These  lands  comprise  more  than  99  German  square  miles  with 
over  206,000  inhabitants — about  37  per  cent,  of  the  population. 
L.  Beaulieu,  i.,  p.  43. 


EVOLUTION  AND  DEGENEEATION  OF  INSTITUTIONS   97 

III.   The  Budgets  of  Germany ,  France,  and  England. 

(a)  Germany. — We  have  just  seen  that  in  the 
principal  States  of  the  Empire  there  is  still  an 
extensive  amount  of  collective  property.  In 
Prussia,  for  instance,  besides  forest  land,  there 
are  nearly  1,500,000  acres  of  arable  land,  an  area 
equal  in  extent  to  one  of  the  smaller  departments 
of  France. 

(h)  France. — Of  the  old  collective  property  of 
France  only  forest  land  remains,  all  grazing  and 
arable  land  having  long  since  been  alienated. 

The  budgets  of  modern  times,  however,  still 
exhibit  traces  of  the  old  feudal  system.  The 
State,  for  instance,  up  to  the  last  few  years, 
continued  to  receive  quit-rents  for  properties 
under  the  old  system.^  The  budget  shows  an 
annual  decrease  in  these  receipts,  having  fallen 
from  100,000  francs  in  1857  to  32  francs  in 
1869.  In  1876,  however,  there  was  a  rise  to 
2000   francs. 

(c)  England.  —  Here  the  decline  of  collective 
property  is  still  further  exhibited.  In  the 
Statistical  Abstract  published  in  1877,  the  net 
revenue  from  Crown  lands  for  Great  Britain  and 
Ireland  figures  at  £40,000  net.  Among  the  "  mis- 
cellaneous receipts"  we  find  only  £200,000  which 
can  be  regarded  as  revenue  derived  from  collec- 
tive    property,     i.e.     15     millions    in    a    budget 

^  Jjeroy-Beaulieu,  TraiUde  la  science  des  finances,  i.,  p.  35, 
G 


98        UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

receipt  of  1,956,000,000  francs.  Later  on,  in 
1884-1885,  these  revenues  became  rather  larger 
(principally  owing  to  the  purchase  of  the  shares 
in  the  Suez  Canal  which  had  belonged  to  the 
Viceroy  of  Egypt),  but  the  proportion  remains 
the  same — 22  to  23  millions  in  a  budget  of 
2  milliards  300  millions.  We  see,  then,  that  in 
all  the  financial  systems  of  modern  Europe  the 
progressive  evolution  of  the  modern  system  is 
taking  the  place  of  the  more  or  less  rapidly- 
degenerating  feudal  system. 

§  2.  Modification  of  similar  institutions  in  dijfferent 
social  groups. 

According  to  the  primitive  constitution  of  things, 
the  land  occupied  by  a  tribe  or  clan  was  regarded 
as  res  nulliiis,  and  consequently  at  the  free  disposal 
of  all  the  members  of  the  community  (the  Feld- 
Walt-  und  Weidegemeinschaft  of  V.  Maurer). 

With  the  increase  of  population,  the  value  of 
land  rose,  and  the  state  of  things  became  modified, 
the  rights  of  groups  and  individuals  becoming  con- 
solidated and  at  the  same  time  limited.  Then  arose 
gradually  or  simultaneously  the  following  various 
forms  of  landed  property  : — (1)  Land  held  by 
families ;  (2)  by  villages ;  (3)  feudal  property ; 
(4)  communal  or  public  property ;  (5)  property 
belonging  to  corporations;  (6)  private  property. 

Family,   village,  and  feudal  property  represent, 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    99 

among  certain  peoples,  three  successive  stages  in 
the  evolution  of  property.  When  the  old  system 
of  land  tenure  was  abolished,  private  and  communal 
or  public  property  began  to  develop  simultaneously. 

While  certain  lands  which  were  free  to  all  the 
inhabitants  became  transformed  into  collective 
property,  other  such  lands  lost  their  public  char- 
acter and  became  private  property.  In  the  first 
case,  the  communes,  on  being  called  upon  to  fulfil 
functions  of  increasing  complexity,  proceeded  to 
transform  all  or  part  of  the  properties  concerned 
into  patrimonial  property  or  property  for  the  use  of 
the  people  (communaux,  allmenden)} 

In  the  second  case  the  property  of  the  old 
community  became  the  joint  but  undivided  property 
of  the  members  of  the  corporation ;  when,  however, 
for  purposes  of  cultivation  it  became  necessary  to 
divide  it,  the  corporative  property  became  trans- 
formed into  private  property. 

*  Giron,  Le  droit  ndministratif  de  la  Belgique,  No.  683.  ' '  There 
were  three  kinds  of  commimal  property  : 

"(a)  Property  directly  appropriated  to  the  use  of  the  public  such 
as  public  squares,  streets,  churches,  &c. 

"  (6)  Cmnmunal  property  properly  speaking — i.e.  the  real  estate 
and  rights  belonging  to  the  tribune  and  to  which  the  people 
were  entitled  to  a  personal  share.  These  consisted  of  the 
forest  land,  rights  of  appanage,  waste  land,  moorland,  and 
the  rights  of  pasturing. 

"(c)  Patrimonial  property,  i.e.  that  held  by  the  commune, 
the  revenue  from  which  went  to  the  commune  to  defray  the 
expenses  of  administration.  It  included  timber  land,  arable 
land,  house  property,  market  places,  &c." 


100     UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

On  reviewing  in  succession  these  various  phases 
in  the  evolution  of  landed  property,  it  will  clearly 
be  seen  that  modification  has  everywhere  been 
attended  by  degeneration. 

1.  Family  property  {Montenegro). — Of  all  the 
Balkan  States,  Montenegro — owing  to  the  natural 
barriers  which  separate  it  from  the  rest  of  Europe — - 
has  best  preserved  its  archaic  institutions.  Here, 
side  by  side  with  modern  institutions,  may  be  found 
the  old  system  of  division  into  forty-two  tribes 
(pleme)  which  are  sub-divided  into  clans  or  confrater- 
nities (brastvo)  and  into  patriarchal  families  (zad- 
rugas  and  inokosnas).^  The  development  of  modern 
political  and  judicial  institutions  has,  however,  con- 
siderably lessened  the  importance  of  the  plemes 
and  the  brastvos,  so  that  progression  in  this  direc- 
tion has  not  been  effected  without  accompanying 
degeneration. 

With  regard  to  property,  the  two  different  forms 
of  family  tenure  have  been  substituted  for  what 
was  formerly  the  tribal  or  clan  system.  Of  the 
former  collective  property  of  the  clan,  there  only 
remain  the  following  traces  : — 

1.  Property  rights  held  over  certain  portions  of 

land — generally  forest  or  waste  land. 

2.  The  right  of  pre-emption  in  favour  of  mem- 

bers of  the  brastvo  or  of  those  related  to 

^  For  information  about  the  common  or  differential  characters 
of  the  zadrnga  and  the  inokosna  see  Ardent,  La  Famille  zoiigo- 
slave  au  Mont^6gro,    {Mforme  sociale,  17th  October  1888.) 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    101 

a  member  within   the  first  six  degrees  of 

lineal  descent.^ 

3.  The  right  of  allotting  to  relatives  their  share 

in  the  duties  of  helping  widows  and  paupers 

in  their  work.      The  workers  in  this  case 

receive  no  payment,  neither  have  they  any 

right  to  demand  maintenance. 

Still  rarer  are  vestiges  of  the  collective  property 

of  the  pleme.    A  few  portions  of  land,  however,  still 

belong  to  that  body,  and  it  is  probably  a  survival  of 

this  ancient  condition  of  things  that  foreigners  are 

^Article  48  of  the  Civil  Code  of  1888,  drawn  up  by  Bogisic  in 
all  possible  accordance  with  "the  excellent  customs"  of  Monte- 
negro, begins  with  the  statement  that  "the  right  of  pre-emption, 
a  privilege  which  has  so  long  been  enjoyed  by  the  members  of  the 
brastvo,  by  persons  whose  lands  adjoin,  and  by  the  members  of  the 
village  and  pleme,  still  flourishes,  and  will  probably  continue  to 
do  so." 

Bogisic  adds  that,  in  accordance  with  this  right,  "any  person 
desiring  to  sell  his  land,  or  any  kind  of  real  estate  belonging  to 
him,  is  constrained,  according  to  the  established  custom  in  such 
cases,  to  first  off'er  it  in  legal  order  to  those  persons  who  enjoy  the 
right  of  pre-emption,  in  order  to  give  them  an  opportunity  of  pur- 
chasing it  at  the  price  at  which  it  is  to  be  offered  to  the  public. 

Article  49,  sec.  1,  gives  a  list  setting  out  the  order  of  precedence 
of  those  who  enjoy  the  right  of  pre-emption. 

1.  Members  of  the  brastvo  within  the  first  six  lineal  degrees  of 

descent. 

„    r>  .        J-  •  •      1     J       ^  Transference,    of    recent 

2.  Persons  owning  adjoining  lands.  .  .  '      .  , , 

3.  The  other  members  of  the  village.  J-       f  •  iV       •  •     ii 

4.  The  other  members  of  the  pleme.    I      „  "f  \  ^  ongma  y  con- 

^  ^      fined  to  relatives. 

If  none  of  those  entitled  to  the  first  oflFer  desire  to  purchase,  the 
owner  may  then  sell  his  property  to  any  other  Montenegiiii.  ;  ,    ,  , 


102     UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

not  permitted  to  acquire  landed  property  in 
Montenegro,^  and  that  the  public  are  unrestricted 
in  the  right  to  hunt  over  any  ground  they 
choose.^ 

2.  Village  'property  {Russia). — Village  communes 
and  the  periodical  division  of  land — the  mer  of 
Kussia  or  the  dessah  of  Java,  for  instance — do 
not  represent  types  of  a  primitive  system,  but 
are  the  outcome  of  a  whole  series  of  modifica- 
tions. Kowalevsky  traces  the  evolution  of  the 
present  system  in  Eussia  through  the  following 
principal  stages : — 

1.  The  joint  use  of  land  by  the  members  of  one 

family  group  (pechische),  corresponding  to 
the  zadruga  of  the  Southern  Servians,  and 
sometimes  comprised  of  more  than  forty 
persons. 

2.  The     division     of    the     mother-family     into 

separate  households,  thus  forming  a  village 
community,  and  the  temporary  allotment 
of  the  land  of  the  community  among  the 
separate  families. 


^  The  old  Montenegrin  law  relating  to  landed  property,  which 
prescribes  the  purchase  of  land  in  Montenegro  by  any  but 
Montenegrins,  is  still  in  full  force.  No  transaction  in  violation 
of  it  is  legally  binding  (Dickel,  £ti(de  sur  le  nouveau  Code  civil 
monUn^grin). 

^  Throughout  the  Southern  Slavonic  countries,  hunting  is  the 
free  right  of  all.  Anyone  may  hunt  where  and  how  they  please, 
not  only  on  public  ground,  among  the  mountains  and  forests,  but 
■upop  private  property,  whether  cultivated  or  not  (Dickel,  p.  36). 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    103 

3.  The   alienation   of   all   or  part  of  the  allot- 

ments assigned  to  the  families  and  the 
constitution  of  agricultural  communes  no 
longer  necessarily  consisting  exclusively 
of  persons  akin. 

4.  The  periodical  division  of  land,  which,  as  the 

population  increased,  was  instituted  with  a 
view  to  re-establishing  an  equal  distribu- 
tion.     This  system  of  division,  which  was 
established  gradually,  only  dealt  with  the 
more  valuable  sort  of  land,  such  as  meadows 
and  arable  land.     The  forest  land  and  pas- 
turage, that  at  least  which  was  not  already 
annexed  by  the  Lords  of  the  Manor,  was 
free  to  all. 
This  transformation  of  family  communities  into 
village     communities     was     not    effected     without 
accompanying     degeneration.     The    administrative 
institutions  of  the  family  group  disappeared,  and 
the  rights  of  pre-emption  in  favour  of  blood  rela- 
tions  were    gradually   replaced    by   village   rights. 
The    importance    of    the    family,    regarded    as    an 
economic  group,  decreased  in  proportion  with  the 
increase  of  the  importance  of  the  village. 

In  some  places,  however,  and  especially  among 
the  Ossetes  who  inhabit  the  valleys  of  the 
Caucasus,  the  old  system  may  still  be  found. 
There,  at  any  rate  up  to  within  the  last  few 
years,  the  aouls  (villages)  are  principally  com- 
prised   of    families   holding   land  in   joint   tenure, 


104     UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

frequently  sharing  all  things  in  common.  These 
aonls  are  very  rarely  met  with  nowadays.^ 

Besides  these  family  communities,  there  yet 
remain  among  the  Ossetes,  as  in  Montenegro, 
numerous  vestiges  of  the  primitive  system  of 
clan  property,  i.e.  the  appropriation  of  certain 
portions  of  land  by  the  members  of  the  clan, 
the  common  use  of  pasturage  and  forest  land,  the 
enforced  participation  in  certain  public  works,  and 
the  rights  of  heritage  over  unclaimed  land,  or  un- 
appropriated property  which  had  become  so  owing 
to  the  lapse  of  some  "  feu  "  or  by  the  extinction  of 
a  family  community.^ 

3.  Feudal  property  {EnglaTid). — The  introduction 
of  the  feudal  system  into  England  resulted  in  the 
substitution  of  a  new  method  of  grouping,  in  place 
of  the  old  agrarian  communities.  These  new 
groups,  like  the  townships  of  earlier  times,  con- 
sisted of  a  complete  organization  occupying  definite 
boundaries.  Instead,  however,  of  being  a  family 
group  administrated  by  a  democratic  organization 
and  government,  this  new  system,  which  Stimner- 
Maine  calls  a  "  manorial  group,"  consisted  of 
a  tenantry  autocratically  grouped  together  and 
governed  by  a  feudal  chief — the  Lord  or  Seignior.^ 

^  See  Kowalevsky,  Coutume  contemporaiiu  ct  hi  a7icienne,  p.  42. 
Paris,  Larose,  1893. 

2  See  Kowalevsky,  Coutume  contemporaiiu  et  loi  ancienne,  pp.  68 
and  following.     Paris,  Larose,  1893. 

^  See  Sumner-Maine,  Les  Communaut4s  de  village;  1  Z>.,  Etudes 
8ur  VHistoire  du  Droit;  1  D.,  VAncien  Droit  et  la  coutume  ancienne. 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    105 

Although  it  cannot  definitely  be  asserted  that 
each  one  of  these  manorial  groups  was  developed 
from  what  had  formerly  been  a  village  community, 
yet  it  is  evident  that  such  was  frequently  the  case, 
and  that  the  transformation  was  accompanied  by 
partial  degeneration.     For  instance  : — 

1.  The  assembly  of  the  inhabitants  of  the  town- 

ship, which  formerly  exercised  complete  con- 
trol over  all  legal  and  administrative  affairs, 
disappeared,  and  in  its  place  sprang  up  the 
Manorial  Court  which  was  comprised  of  a 
limited  number  of  tenants  and  presided  over 
by  the  Lord  of  the  Manor  or  his  representa- 
tive. 

2.  The  collective  property  became  absorbed,  or 

was  at   least  considerably  reduced  by  the 
acquisitions  of  the  Lord  of  the  Manor,  or  by 
divisions  effected  by  members  of  the  com- 
munes.    The  old  system  of  collective  pro- 
perty held  by  townships  did  not,  however, 
wholly  disappear. 
(a)  The  "  rights  of  usage "  in   regard  to  waste 
land,  forest  and  moors  (such  as  the  use  of  unclaimed 
pasturage,  the   cutting   of   timber,  etc.),  were  still 
enjoyed  by  the  old  inhabitants,  and  even  in  some 
instances  by  other  persons  upon  whom  these  "rights" 
had  been  conferred. 

Emile  Laveleye,    Les   Communautes   de  famille   et   de   village 
{Remie  d' ^oiwmie 'politique y  1888,  pp.  350  and  following). 

Vinogradoff,  Villainage  in  EnglaTid,  Oxford  and  London,  1892. 


106     UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

(h)  With  regard  to  meadow  land,  sometimes  the 
Lord  of  the  Manor  put  up  enclosures  for  his  own 
benefit  from  Candlemas  till  Midsummer,  the  rights 
of  the  community  being  established  during  the 
remainder  of  the  year  only ;  sometimes  it  was  the 
community  who  put  up  the  enclosures,  when  the 
Lord  of  the  Manor  was  only  entitled  to  the  use  of 
the  land  during  the  intervals  ;  sometimes  pasturage 
was  held  as  the  joint  property  of  the  old  community, 
or  rather  of  their  descendants  the  tenants ;  but  as  a 
rule  it  was  regarded  as  more  or  less  common  pro- 
perty. The  best  meadow  land  was  divided  up  into 
what  were  termed  "  deals "  and  apportioned  by 
drawing  lots. 

(c)  With  regard  to  arable  land,  the  method  of 
appropriating  and  cultivating  the  land  occupied  by 
the  tenants  retained  many  traces  of  the  village 
system  of  collective  property.  For  instance :  the 
enforced  rotation  of  crops ;  the  periodical  division 
of  land  in  certain  parts  of  the  country ;  the 
division  of  land  into  three  breaks  in  other  places ; 
and  the  destruction  after  the  harvest  of  the  en- 
closures surrounding  the  crops,  after  which  the  land 
was  used  for  the  herding  of  cattle. 

These  survivals  may  yet  be  found  in  some  districts 
of  modern  England,  in  spite  of  all  the  great  changes 
in  the  English  system  of  property  ;  changes  such  as 
the  disappearance  of  the  serf  and  the  appearance  of 
yeomen  in  the  course  of  the  thirteenth  to  the  six- 
teenth century,  and  the  dispossession  of  the  yeoman 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    107 

in  favour  of  the  growth  of  large  properties  in  the 
course  of  the  sixteenth  to  the  eighteenth  century. 

4.  Public  or  collective  property  (Switzerland). — 
It  is  easy  in  Switzerland  to  follow  the  course  of 
the  transformation  into  political  communes  of  the 
old  system  of  economic  communes,  whether  village 
or  manorial. 

In  the  mountainous  parts  of  Switzerland  this 
transformation  is  still  incomplete,  and  side  by 
side  with  the  modern  commune  may  be  seen 
the  old  form  of  collective  property,  though  in  a 
more  or  less  advanced  stage  of  degeneration. 

The  successive  stages  of  this  evolution  may  be 
enumerated  as  follows  : — 

1.  The  village  communities  (the  Felcl-Walt-und- 

Weidegemeinschaft  of  von  Maurer). 

2.  The   collective    property   of  the   inhabitants, 

whether  feudal,  free,  or  partly  both 
(Gemischte  Gemeinde). 
The  Feldgemeinschaft  completely  disappeared 
after  the  Eeformation,  the  collective  land 
of  the  community,  the  joint  use  of  which 
was  the  right  of  all  the  inhabitants,  being 
restricted  to  mere  waste  land,  forest  land, 
and  pasturage  (Allmend)} 

^  The  Allmend,  in  the  primitive  sense  of  the  word,  meant  that 
part  of  the  old  collective  property  held  in  joint  tenancy  by  a 
community  of  inhabitants  or  any  other  public  body,  the  use  of 
which  was  limited  to  those  who  had  a  personal  title  to  it.  This 
primitive  meaning  has  changed  in  Switzerland — excepting  in  the 


108     UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

3.  The    institution    of    Bilrgergemeinde  —  public 

corporations  —  the  members  of  which 
enjoyed  the  sole  use  of  the  Allmend, 
and  upon  whom  fell  the  costs  of  local 
administration ;  as  the  population  increased, 
the  conditions  of  admission  into  the  Burger- 
gemeinde  became  more  and  more  strict. 

4,  The  creation  of  Mnwohnergemeinde,  or  political 

communes,  established  side  by  side  with  the 
old  system,  and  in  many  localities  eventu- 
ally taking  its  place.  In  these  the  responsi- 
bilities of  the  former  system  were  assumed, 
and  part  or  the  whole  of  the  collective 
property  was  appropriated.  Where  the 
old  communities  still  survive,  they  have, 
as  a  rule,  ceased  to  exercise  the  greater 
part  of  their  original  functions.  In  the 
Cantons  of  Berne  and  Saint-Gall,  for 
instance,  the  old  communities  have  de  - 
livered  up  the  greater  part  of  their 
possessions  to  the  political  communes  to 
provide  for  the  expenses  of  general  ad- 
ministration. Their  only  now  remaining 
function  is  the  administration  of  the  small 
remainder  of  their  patrimony  and  the  main- 
tenance of  the  indigent  members  of  the 
community. 

Canton  of  Schwyz — owing  to  the  changes  in  the  institution  itself 
(see  Miaskowski,  Die  Schweitzerische  Allmend  in  ihrer  geschicht- 
lichen  Entwickelung,  von  xiii.  Jalirhwndcrt  his  zum  Gegenwart). 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    109 

Independently  of  this  decay  of  the  old  system 
of  communities,  the  formation  of  political  com- 
munes was  attended  by  other  phenomena  of 
degeneration : — 

1.  The  suppression  of  all  or  part  of  the  "  rights 
of  usage  "  enjoyed  by  the  inhabitants. 

The  Einwohnergemeinde,  being  called  upon  to  dis- 
charge more  and  more  onerous  and  complex  func- 
tions, were  obliged  to  either  partially  or  wholly 
transform  the  communal  possessions,  to  the  per- 
sonal use  of  which  the  people  were  entitled,  into 
property  appropriated  to  the  use  of  the  public, 
either  directly  {i.e.  into  churches,  teaching  insti- 
tutions, etc.)  or  indirectly,  as  a  means  of  obtaining 
a  revenue  {Erwerbsquelle). 

2.  The  decrease  in  collective  property. 

Many  of  the  Bilrgergemeinde,  although  no  longer 
discharging  public  functions,  retained  part  of  their 
estates,  which  were  held  by  the  members  in  joint 
tenancy.  On  the  other  hand,  as  the  increased 
population  necessitated  the  cultivation  of  the 
Allmend,  the  original  "  right  of  usage  "  re- 
sulted in  many  instances  in  a  transformation  of 
the  land  into  private  property. 

This  transformation,  however,  was  not  always 
complete,  and  all  the  intermediate  stages  may  be 
traced  between  the  old  collective  tenure  and  the 
appropriation  by  individuals. 

5.  Corporative  property  (Belgium). — Here  we  will 
limit  the  sphere  of  our  observations  to  Belgium,  in 


110    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

order  to  avoid  repetition,  similar  examples  being 
almost  universally  exhibited  throughout  Europe.-^ 

Our  information  is  obtained  from  the  work  of 
Paul  Errera,  entitled  Les  Masuirs,  recherches  his- 
toriques  et  juridiques  sur  quelques  vestiges  des  formes 
anciennes  de  la  'proprUU  en  Belgique. 

The  masuirs  (the  amborgers  of  Flanders)  were 
the  mansuarii  or  maiisoarii  of  the  Merovingian 
period,  originally  serfs,  afterwards  tenants  and 
copy-holders,  and  finally  freemen. 

Their  history  exhibits  the  following  stages : — 

1.  The  feudal  epoch  in  which  the  masuirs — i.e. 

all  the  members  of  the  manorial  group — 
enjoyed  "rights  of  usage"  over  all  waste 
lands,  forests  and  pasturage  adjoining  their 
holdings.  These  rights  appear  to  have  been 
conferred  by  the  Lord  of  the  Manor,  but 
they  really  dated  from  a  much  earlier 
period. 

2.  As  the  increasing  population  necessitated  the 

regulation  and  limitation  of  these  rights, 
certain  conditions  of  property  and  residence 
were  stipulated  for  in  those  seeking  admis- 
•  sion  to  the  rights  of  the  masuirs,  and  these 
privileged  persons  organized  themselves  into 
corporations  which  were  more  or  less  ex- 
clusive and  separate  from  the  general 
community. 

*  With  regard  to  corporative  property  in  Switzerland,  see  von 
Miaskowski,  Die  schweizerische  Allmend,  pp.  37  and  following. 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    111 

3.  By  degrees — by  means  of  cantonments,  pur- 

chases, prescriptive  claims,  &c. — these  cor- 
porations absorbed  the  best  part  of  the 
land,  and  became  almost  independent  of 
the  Lord  of  the  Manor;  as  a  rule,  the  latter 
gave  up  half  of  the  common  territory  to 
them,  and  freed  the  surplus  from  all  rights 
of  usage.  In  the  corporations  of  masuirs, 
however,  there  were  still  a  few  remaining 
vestiges  of  some  of  the  institutions  of  the 
old  manorial  group  from  which  they  had 
gradually  developed.  The  Lord  of  the 
Manor,  for  instance,  himself  being  an  in- 
habitant and  a  masuir,  had  a  right  to  a 
share  in  the  property  of  the  community, 
and  further,  in  his  seigniorial  capacity, 
certain  privileges  accrued  to  him  such  as 
"  la  haute  fleur  des  bois,"  i.e.  tithes  and 
pannage  (crops  of  acorns). 

4.  The   Ee volution   put   an   end    to    all    feudal 

rights,  and  removed  the  last  remaining 
traces  of  the  origin  of  the  masuirs. 
Throughout  this  long  series  of  transformations,  it 
is  evident  that  degeneration  has  followed  in  the 
track  of  progress.  Besides  the  disappearance  of 
the  manorial  group  and  its  attendant  institutions, 
the  rights  of  the  masuirs  may  be  said  to  have 
become  more  restricted  as  they  became  more  defined 
and  secure.  In  the  early  days,  all  the  inhabitants 
enjoyed  joint  rights  over  a  vast  common  territory, 


112    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

at  the  close  of  the  old  system  this  territory  had 
become  much  reduced  in  extent,  and  had  become 
the  property  of  a  more  or  less  large  group  of 
privileged  persons. 

6.  Private  'pro'perty  {Switzerland). — After  the 
Eevolution,  the  communities  of  masuirs  and  other 
similar  corporations  ceased  to  have  any  recognized 
legal  existence.  Those  which  still  survived  in 
spite  of  the  irregularity  of  their  legal  position, 
owed  their  existence  to  their  insignificance.  The 
others  dispersed  themselves,  or  were  dispersed,  and 
the  property  which  had  belonged  to  them  was 
either  incorporated  with  the  communal  estate,  or 
divided  up  among  the  members  of  the  old  com- 
munity. 

In  each  of  these  cases  the  transformation  was 
attended  by  degeneration,  for  the  archaic  adminis- 
trative organization  disappeared. 

We  saw  in  the  allmend  of  Switzerland,  this 
same  divergent  evolution  of  public  and  private  pro- 
perty, part  of  the  common  land  being  transformed 
into  communal  property,  while  the  use  of  the  sur- 
plus ended  in  some  instances  in  the  land  becoming 
ultimately  the  private  property  of  individuals. 
This  frequently  occurred  where  land  was  cultivated 
as  orchards.  In  early  times  both  fruit  and  fruit- 
trees  belonged,  like  the  land  itself,  to  the  community, 
and  in  certain  parts  of  the  Cantons  of  XJri  and 
Schwyz  this  is  still  the  case.  By  degrees,  however, 
individual  rights  over   fruit-trees    planted  on  the 


EVOLUTION  AND  DEGENERATION  OF  INSTITUTIONS    113 

allmend  came  to  be  recognized.  These  rights, 
whether  temporary  or  held  for  a  life-time,  eventually 
became  perpetual,  and  finally  this  right  to  the 
private  acquisition  of  trees  led  to  a  right  to  acquire 
the  land  itself.  This  last  transformation  was  not 
effected  without  a  struggle  and  occasionally  the 
land  was  reclaimed  by  the  community,  the  pro- 
prietor of  the  trees  receiving  compensation.  Now- 
a-days  the  possession  of  trees  and  land  usually  go 
together.  Duality  of  this  kind,  however,  is  still  to 
be  met  with  in  certain  localities.  In  the  Scrnfthal 
(in  the  Canton  of  Glaris)  a  still  stranger  custom 
prevails  with  regard  to  the  maple  forests.  There, 
the  soil,  the  trees,  and  the  fallen  leaves  (the  latter 
being  used  as  litter  for  cattle)  all  belong  to  different 
persons.!  With  regard  to  house  property  there  are 
more  intermediary  conditions  between  use  and 
possession.  In  some  villages,  the  chalets  as  well 
as  the  ground  upon  which  they  are  built,  belong  to 
the  whole  community  ;  in  other  villages,  both  are 
part  of  the  collective  property.  Sometimes  private 
possession  is  restricted  to  the  house  or  chalet,  the 
right  to  the  ground  upon  which  it  is  built  lapsing 
with  the  existence  of  the  house.  In  order  to  limit 
the  number  and  durability  of  these  buildings,  many 
restrictions  are  imposed,  such  as  the  prohibition  to 
build  houses  of  stones,  or  chalets  of  wood  cut  from 
trees  not  belonging  to  the  builder  himself  or  to  the 
corporation  to  which  he  belongs,  etc. 

^  Miaskowski,  Die  schweizerische  Allmendy  pp.  18  and  following. 
H 


/ 


114    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

7.  Summary, — This  long  series  of  modifications, 
the  result  of  which  was  the  transformation  of 
primitive  communities  of  goods  into  the  modern 
forms  of  public  and  private  property,  was  ac- 
companied throughout  by  degenerative  changes. 
The  establishment  of  family  property  entailed 
the  curtailment  of  tribal  and  clan  rights.  Family 
property  passed  into  property  held  by  the  village ; 
next  the  development  of  feudal  tenure  involved  the 
degeneration  of  the  old  agrarian  communities ; 
finally,  the  primitive  organization' of  property  with 
the  administrative  and  political  institution  depen- 
dent on  it,  disintegrated  and  disappeared  as  the 
primitive  community  of  goods  lapsed  into  the 
personal  enjoyment  of  these  by  individuals,  and 
as  the  primitive  method  of  land  tenure  passed 
into  the  rights  of  private  property. 

We  see  then  that  degeneration  has  always 
accompanied  evolution :  the  destruction  of  old 
institutions  is  involved  in  the  formation  of  new 
institutions. 


PART  II 

DEGENERATION    IN    THE    EVOLUTION    OF    ORGANISMS 
AND    SOCIETIES 

We  have  seen  that  modification  of  organs  and  of 
institutions  is  always  associated  with  partial  de- 
generation. We  have  now  to  show  that,  similarly, 
when  organisms  and  societies  become  modified,  de- 
generation is  shown  in  some  of  their  organs  or 
institutions.  This  shows  again  the  universality  of 
decjenerative  evolution. 


CHAPTEK  I 

ALL    ORGANISMS    EXHIBIT    RUDIMENTARY    ORGANS 

All  existing  organisms  have  lost  some  organs  in 
the  course  of  their  phylogenetic  development. 

This  may  be  proved  in  two  ways  :  either  there 
are  remaining  vestiges  of  these  organs,  or  else  they 
are  to  be  found  in  other  creatures  which  may  be 
regarded  as  ancestors. 

MS 


116    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

1.  Rudimentary  organs,  signs  of   a  degenerative 

transformation  in  the  organism  itself,  are 
either  organs  which  have  ceased  to  be 
functional,  or  which  have  so  diminished  in 
importance  that  their  total  disappearance 
would  be  unattended  by  any  appreciable 
loss  to  the  organism.  In  the  majority  of 
cases  this  cessation  of  function  is  attended 
by  a  corresponding  structural  decay. 

2.  The   system   of   comparing   living   organisms 

with   their   presumptive   ancestors   equally 

demonstrates   the   retrogression    of   certain 

organs. 

Among  the  Orobanchacete  for  instance,  parasitic 

plants  derived  from  normal  green  plants,  no  trace 

of    cotyledons    is    observable   from   the    period    of 

germination.^ 

Among  animals,  .taking  the  horse  as  an  example, 
several  organs  have  wholly  disappeared.  In  the 
genealogy  of  the  horse,  which  is  well  known,  the 
earliest  ancestor  Eohippus,  possessed  five  functional 
fingers  on  the  fore-feet,  and  four  toes  on  the  hind- 
feet.  The  horse  still  possesses  one  functional  finger 
and  one  functional  toe,  two  rudimentary  fingers  and 
two  rudimentary  toes.  Two  fingers  and  two  toes 
have  entirely  disappeared. 

It  is  hardly  necessary  to  point  out  that  this 
system   of   comparison    does    not   demonstrate   the 

^  L.  Koch,  Die  Entwickeliingsgeschichtc  der  OrobancJien.    Heidel- 
berg, C.  Winter,  1887. 


RUDIMENTARY  ORGANS  OF  ANIMALS  1 1 7 

degenerative  changes  attending  the  phylogenetic 
development  of  an  organisra  with  such  incontestable 
certainty  as  does  the  existence  of  vestiges  of  rudi- 
mentary organs. 

It  is  our  belief  that  all  organisms  contain 
vestiges  of  organs,  either  more  or  less  apparent. 
In  our  present  condition  of  knowledge,  however, 
it  is  quite  impossible  —  particularly  as  regards 
plants — to  prove  this  theory  universally.  It  is 
to  be  hoped,  however,  that  future  researches  will 
ultimately  succeed  in  establishing  it. 

In  the  meantime,  we  will  point  out  the  most 
typical  among  the  cases  known  to  us. 

With  animals,  as  with  plants,  our  investigations 
have  extended  not  only  to  every  kind  of  group,  but 
to  the  most  varied  systems  of  organs,  thus  giving 
our  theory  an  extremely  wide  application. 

Section  I. 

Budimentary  Organs  of  Animals. 
§  1.  Budimentary  Organs  in  Man. 

Throughout  the  whole  human  organic  systems 
signs  of  degeneration  abound. 

1.  The  Integumentary  System. — In  the  ancestors 
of  man,  the  entire  surface  of  the  skin  was  covered 
with  hairs.  Man's  clothing  of  hair  is  far  from 
perfect,  the  hairs  of  which  it  is  composed  being 
rudimentary. 


118    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

According  to  Her  twig,  the  teeth  should  be 
regarded  as  part  of  the  tegumentary  system,  as 
they  really  represent  the  scales  of  the  skate, 
situated  within  the  buccal  cavity. 

In  man,  the  last  molar,  or  wisdom  tooth,  is  a 
rudimentary  tooth.  The  small- sized  shallow  crown, 
the  diminished  number  of  tubercles,  the  fusion  of 
the  roots,  the  tardy  appearance  and  occasional 
absence  altogether,  are  all  indications  of  a  rudi- 
mentary condition. 

2.  The  Skeleton.  —  With  few  exceptions,  the 
articular  surfaces  of  the  bodies  of  mammalian 
vertebrates  are  covered  in  youth  with  bony 
plates.  These  sometimes  become  very  thick,  and 
are  called  terminal  epiphyses.  In  some  mammals 
— the  Sirenians,  for  instance — the  terminal  epi- 
physes have  disappeared.  In  man  they  still 
exist,  but  in  an  advanced  stage  of  degeneration. 
In  the  lower  vertebrates,  sucli  as  the  crocodile, 
many  more  ribs  are  functional  than  in  man. 
In  the  crocodile  all  the  ribs  connected  with  the 
cervical  vertebrae  are  functional,  whereas  in  man 
they  have  degenerated.  Of  one  entire  section  of 
the  human  vertebral  column — the  tail — so  fully 
developed  in  the  majority  of  other  vertebrates, 
only  a  vestige  now  remains. 

Other  rudimentary  skeletal  pieces  are  the  lesser 
horn  of  the  hyoid  bone,  the  stylo-hyoidean  liga- 
ment, the  coracoid  process,  and  the  interclavicular 
ligament. 


KUDIMENTARY  ORGANS  OF  ANIMALS     119 

3.  The  Muscular  System. — The  cutaneous  muscles, 
those  of  the  shell  of  the  ear,  and  those  that  move 
the  tail,  which  in  most  mammals  are  well  developed, 
are  still  present  in  man,  but  have  degenerated. 

Further,  there  is  to  be  found  in  man  the 
intra-acetabular  part  (the  round  ligament)  of  the 
deep  flexor  muscle  of  the  toes  which  is  functional 
in  some  animals — in  young  ostriches,  for  instance. 
In  the  adult  ostrich  the  intra-acetabular  part  is 
separated  from  the  rest  of  the  muscle,  which  is 
attached  to  the  pelvis.  Traces  remain  in  the 
horse  of  a  connection  between  the  intra-  and  the 
extra-acetabular  parts ;  the  muscle  itself  is  divided 
into  two  parts,  the  pectineal  muscle  in  the  thigh, 
and  the  deep  flexor  muscle  of  the  toe  situated  in 
the  leg.  In  the  orang-outang,  this  degeneration 
has  made  further  advances  than  in  man,  the  intra- 
acetabular  part  of  the  muscle  having  entirely 
disappeared.^ 

4.  The  Nervous  System. — Here  we  find  numerous 
signs  of  degeneration,  of  which  the  following  are 
a  few  examples  : 

In  the  brain  the  pineal  gland,  the  last  remaining 
vestige  of  what  was  formerly  a  functional  eye,  is 
present. 

In  the  spinal  cord  the  filu^n  terminale  still 
exists.  We  know  that  the  spinal  cord  in  man  does 
not  retain  its  normal  thickness  to  the  extremity  of 

^  See  Sutton,  Ligaments,  their  mature  and  morphology.  London, 
1887, 


120    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

the  vertebral  column  but  is  arrested  at  the  first 
lumbar  vertebra.  There  a  considerable  number  of 
special  nerves  leave  it,  forming  a  mass  of  branches 
like  a  horse's  tail.  Along  the  centre  of  these 
nerves,  in  the  middle  line,  a  slender  filament 
represents  the  spinal  cord  to  the  extremity  of  the 
coccyx.  This  filament  is  the  filum  terminale,  the 
spinal  cord  in  a  condition  of  degeneration. 

5.  The  Digestive  System. — The  caecum  and  its 
vermiform  appendage,  are  well  known  to  be  organs 
which  have  degenerated. 

6.  The  Vascular  System. — In  quadrupeds  the 
intercostal  veins  are  vertical,  the  blood  consequently 
flowing  against  gravity.  These  veins  contain  valves 
which  indirectly  facilitate  the  upward  and  onward 
flow  of  the  blood  by  preventing  it  from  running 
back.  Man,  being  a  biped  with  a  vertical  thorax, 
is  provided  with  intercostal  veins  that  are  almost 
horizontal.  The  ancestral  valves  being  no  longer 
indispensable  are  in  a  condition  of  degeneration. 

7.  Sense  Organs, — In  the  olfactory  organ  there 
remains  a  degenerate  Jacohson's  organ.  In  the 
organ  of  sight  there  is  a  third  eyelid  in  a  state 
of  degeneration.  In  the  organ  of  hearing  there 
remains  on  the  shell  of  the  ear  a  kind  of  point 
(Darwin's  point)  which  is  the  last  remaining  vestige 
of  the  ancestral  elongated  and  pointed  ear. 

8.  Genito-urinary  System. — There  is  a  whole 
series  of  rudimentary  organs  in  the  genito-urinary 
system  of  the  higher  animals.     As  is  well  known, 


RUDIMENTARY  ORGANS  OF  ANIMALS  121 

the  Wolffian  body  plays  a  considerable  part  in  the 
formation  of  the  system.  This  body,  the  primitive 
kidney,  loses  its  urinary  function  at  a  certain 
stage  of  embryonic  development,  and  the  permanent 
kidney  which  gradually  develops  alongside,  assumes 
the  urinary  function.  Later  on,  the  Wolffian  body 
assumes  new  functions,  becoming  an  important  part 
of  the  genital  apparatus. 

In  this  transformation  partial  degeneration  occurs, 
resulting  in  such  reduced  structures  as  the  epi- 
didymis, the  organ  of  Rosenmiiller,  the  vas  aherrans, 
etc.  (see  fig.  57). 

§  2.  Budimentary  organs  in  various  groups. 

1.  Ccdenterates. — The  Coelenterates  comprise  three 
great  groups.^  The  Anthozoa,  of  which  the  coral  is 
a  type,  the  Hydrazoa,  which  include  fresh  water 
Hydra  and  the  common  jelly-fish  of  our  seas,  and 
the  Ctenophora,  of  which  the  chief  representative 
in  our  seas  is  Cydippe,  a  globular  transparent  animal 
frequently  to  be  found  floating  in  large  numbers  on 
the  surface  of  the  water. 

The  colonies  of  Anthozoa  are  usually  composed 
of  individuals  all  exactly  alike.  In  some  species, 
however,  in  the  Pennatulidoe  and  the  Alcyonaria 
for  instance,  there  is  a  distinct  differentiation 
amongst  the  numerous  individuals  composing  the 
colony.       Side    by    side    with    sexual    individuals 

^  See  C.  Vogt  and  Emile  Yung,  Traits  d'anatomie  comparie,  vol.  i. 


122    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

provided  with  tentacles  and  the  eight  mesenteric 
folds,  are  other  far  simpler  individuals  :  the  zooides, 
the  function  of  which  is  respiratory  and  of  which 
the  greater  part  of  the  organs  have  degenerated ; 
the  generative  organs  are  lacking,  the  tentacles  are 
very  small,  and  the  mesenteric  folds  only  number 
two  instead  of  eight.  Degeneration,  then,  is 
exhibited  side  by  side  with  specialization. 

Apiong  the  Hydromedusce  similar  examples 
abound.  It  is  known  that  the  polyp-like  or 
medusa-like  forms  of  this  group  which  may  live 
independently,  frequently  associate  themselves 
together  to  form  colonies,  sometimes  predominantly 
polypoid,  sometimes  completely  medusoid,  and  occa- 
sionally a  mixture  of  the  two. 

In  these  cases  a  marked  polymorphism  is  often 
apparent.  The  different  individuals  become  adapted 
to  definite  functions,  and  the  corresponding  organs 
undergo  special  development ;  the  other  parts  of 
the  body  having  become  either  unnecessary  or 
merely  accessory,  begin  to  degenerate  and  finally 
disappear.  Thus  we  see  in  Hydroid  colonies,  not 
only  the  hydra-like  members,  nutritive,  fixed  and 
sterile,  and  the  medusa-like  members  which  are 
reproductive  and  become  free  from  the  colony,  but 
also  certain  individuals  which  are  termed  gono- 
phores.  These  gonophores  are  really  medusa-like 
members  which  have  lost  their  independent  move- 
ment, and  have  consequently  more  or  less  lost 
both  their  tentacles  and  their  umbrella-like  discs, 


RUDIMENTARY  ORGANS  OF  ANIMALS 


123 


i.e.  their  organs  of  locomotion.  In  some  colonies 
of  Hydroids,  polymorphism  has  made  such  advances 
that  there  are  tactile  individuals  of  which  the 
digestive  tube  lacks  both  mouth  and  tentacles,  and 
other  purely  defensive  individuals  of  which  the 
internal  organs  are  almost  all  in  a  state  of  atrophy. 

Opinions  differ  regarding 
the  complicated  question  of 
the  structure  of  the  Siphon - 
ophora.  The  organism  (fig. 
52)  may  be  regarded  as  a 
simple  medusa  of  which 
the  different  appendages — 
the  pneumatophore  (pn.), 
the  swimming  bells  (cL), 
the  siphons  (s.),  the  shield 
(b.),  the  tentacle  (t),  the 
palp  (pa.),  the  gonophores 
(go.),  etc. — constitute  the 
organs,  or  as  a  colony  each 
part  of  which  is  represented 
by  an  individual  polypoid 
adapted  to  fulfil  a  special 
function.  Whatever  theory 
is  accepted,  it  is  clear 
that  a  whole  series  of  parts  of  the  creature  must  be 
regarded  as  organs  in  a  condition  of  degeneration. 

We  accept  the   second    of    these  theories,^  and 

'  Haeckel,  System  der  Medusen :  Jena,  1880-1881.      A.  Lang, 
Traits  d'aimtomie  compar^e. 


Fig.  52.— Diagram  of  the  structure 
of  one  of  the  Siphonophora. 

pn,  pneumatophore  or  float ;  cl, 
swimming  bell;  6,  protective 
polyp  ;  ^  tentacle;  pa,  palp;  g, 
gonophore;  p,  peduncle  bearing 
the  mouth;  «,  iDdivlduals  for  nu- 
trition. 


124    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

regard  the  whole  creature  as  formed  from  a 
Craspedote  Medusa  which  has  become  mother  of  a 
colony  and  of  which  the  umbrella,  developed  into 
the  pneumatophore  {jpn.)  has  had  its  radial  canals 
greatly  simplified  and  its  tentacles  reduced  to  one 
during  the  growth  of  the  colony.  The  stalk-like 
stomach  (p.)  of  this  medusa  has  increased  in  length, 
but  this  development  is  attended  by  corresponding 
degeneration,  the  buccal  aperture,  which  is  situated 
at  the  free  end  of  the  peduncle  in  normal 
Craspedote  Medusae,  having  entirely  disappeared. 
The  stalk,  formed  in  this  way,  serves  as  a  support 
to  the  number  of  other  individuals  of  which  the 
colony  is  constituted,  and  which  are  remarkable 
for  the  great  morphological  variation  they  exhibit. 
Among  these  individuals,  those  at  the  top,  i.e. 
those  nearest  to  the  air-sac,  fulfil  the  function  of 
locomotion.  They  become  transformed  into  swim- 
ming bells  (cl.)  and  contain  no  organs  whatever. 
Below  these  locomotory  organs  are  the  sexual 
individuals,  or  gonophores  {g),  and  the  sterile 
individuals  (s.).  The  former  are  of  medusoid 
structure,  the  umbrella  is  more  or  less  perfect,  and 
they  are  sometimes  provided  with  tentacles,  and 
possess  a  peduncle  or  manubrium  which  some- 
times has  a  buccal  aperture.  The  sterile  individuals 
provide  nutrition  for  the  others.  The  organs  no 
longer  essential  to  them  atrophy  in  a  variable 
degree.  In  the  case  of  some  the  umbrella  is 
present,  in  others  it  is  absent,  and  between  these 


RUDIMENTARY  ORGANS  OF  ANIMALS 


125 


two  extreme  types  come  intermediate  types  which 
exhibit  every  possible  stage  of  degeneration. 

In  the  Ctenophore  group  development  and  de- 
generation are  exhibited  simultaneously  in  the 
organs  of  locomotion.  The  fundamental  and  typical 
shape  of  the  Ctenophore  is  round  or  oval,  and  the 
eight  sides  are  provided  with  swimming  plates, 
originally  uniform — as  in  Bero'e. 
The  individuals  belonging  to 
this  group  exhibit  important 
evidences  of  modification  in 
their  external  morphology.  The 
body  being  sometimes  com- 
pressed in  various  directions, 
the  shape  is  altered  from  the 
original,  and  assumes  a  more 
or  less  irregular  appearance. 
The  organs  of  locomotion 
undergo  a  corresponding 
change.  Take  for  example 
an  adult  specimen  of  Bolina 
norvegica  (fig.  53);  the  body  is  lobate,  although  it 
was  round  during  the  larval  period ;  the  swimming 
plates  are  not  uniform,  four  being  long  and  reaching 
along  the  whole  length  of  the  body,  the  other  four 
being  only  developed  in  the  upper  half  of  the  body 
as  far  as  where  the  lobes  are  inserted,  where  they 
end  as  degenerate  hair-like  processes.  By  referring 
to  the  Cestidge,  which  are  ribbon-like  in  shape,  it 
will  be  seen  that  by  means  of  compression  the  body 


Fig.  53.— Bolina  Norvegica, 
seen  from  the  broad  side. 

C,  short  rows  of  swimming 
plates;  A,  long  rows;  L, 
lobes.  (After  Vogt  and 
Yung,  Trait e  dfanatomie 
comparie.) 


126    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

of  the  animal  is  lengthened  out  into  the  shape  of  a 
narrow  ribbon.  Of  the  primitive  lateral  row  of 
plates,  four  are  represented  by  mere  vestiges,  and 
the  other  four,  which  continue  to  be  functional,  are 
situated,  closely  coupled  together,  on  the  two  edges 
of  the  animal. 

2.  Worms. — We  will  next  take  in  succession  the 
Plathelminthes  or  fiat  worms,  the  Eotifers,  the 
Nemathelminthes,  or  round  worms,  the  Annelids 
and  the  Gephyreans. 

Among  the  Plathelminthes,  the  group  of  Cestodes 
•contains  the  common  tape- worm  of  man  {Taenia 
solium).  In  the  course  of  its  parasitic  existence 
this  worm  has  undergone  considerable  morphological 
changes.  The  digestive  tube  is  lacking,  and  the 
whole  nervous  system  has  become  greatly  simplified. 
The  degeneration  of  the  nervous  system  is  not, 
however,  complete,  for  important  vestiges  still  per- 
sist. The  degeneration  of  the  digestive  tube  is 
much  more  thorough.  In  Tcenia  solium  it  is  alto- 
gether absent.  In  species  closely  allied  to  the  Tcenia 
some  slight  vestiges  of  the  digestive  apparatus  yet 
remain.  The  head  or  scolex  of  some  species  of 
Tetrarhynchus  contain  glandular  cells  which  have 
been  homologized  with  the  salivary  glands  of  other 
flat  worms  {Trematodcs).  In  other  species  of  Tetra- 
rhynchus there  is  a  rudimentary  organ  which  repre- 
sents the  oval  sucker  of  the  Treinatodcs,  and  in 
Anthrocephalus  eloTigatus  the  orifices  of  the  salivary 
glands  are  in  the  region  of  this  vestige  of  the 
digestive  tube. 


RUDIMENTARY  ORGANS  OF  ANIMALS  127 

Many  facts  in  support  of  our  argument  may  be 
drawn  from  a  study  of  the  excretory  organs  of  the 
Cestodes. 

The  condition  of  these  organs  in  Caryophylleus 
mutabilis  best  represents  the  primitive  condition. 
Within  the  body  of  this  worm  are  a  large  number 
of  narrow  ducts  with  ciliated  funnels  communicating 
with  the  spaces  in  the  parenchyma  (Fraipont). 
These  organs  communicate  with  canals  which 
gradually  reunite  and  anastomose  to  find  a  vent  in 
one  single  aperture,  the  foramen  caudale,  which  is 
situated  in  the  posterior  part  of  the  body  where 
there  is  a  bladder.  In  the  Cestodes,  however,  where 
the  body  is  very  long,  the  action  of  the  bladder 
is  insufficient  to  secure  a  complete  evacuation. 
Secondary  apertures  are  therefore  formed  at  in- 
tervals along  the  main  ducts.  This  new  structure 
entails  the  degeneration  of  the  terminal  bladder 
which  has  become  superfluous.  In  Botryoce/phalus 
punctatus,  which  possesses  a  great  number  of 
excretory  apertures,  the  primitive  evacuatory  appa- 
ratus— i.e.  the  contractile  cavity — has  completely 
disappeared. 

Rotifers  are  minute  animals,  usually  living  in 
fresh  water,  a  few  being  marine.  One  of  them,  an 
inhabitant  of  damp  earth  or  moss,  has  been  supposed 
to  possess  the  power  of  revivifying  after  complete 
dessication.  At  the  anterior  end  of  the  body,  a 
rotifer  possesses  a  complicated  ciliary  apparatus 
which  fulfils  the  function  of  locomotion,  and  from 


128    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

the  rotatory  movements  of  which  the  name  of  the 
group  have  been  given.  This  organ  is  well  developed 
in  those  types  which  lead  an  independent  existence, 
but  in  those  where  movement  is  more  restricted  or 
where  the  character  of  the  organ  has  changed,  it  is 
considerably  modified  and  reduced  in  size.  In 
Philodinoe,  (crawling  Eotifers)  the  organ  of  rota- 
tion has  lost  the  central  part,  and  in  its  place  is 
substituted  a  very  complicated  organ  of  prehension. 

In  sessile  forms  such  as  Floscularia,  the  organ  is 
modified  and  only  the  primitive  character  of  the 
inner  ring  is  retained,  while  the  outer  ring  is 
segmented  and  becomes  a  series  of  arms  or  lobes, 
furnished  with  stiff  bristles.  In  Apsilus,  another 
sessile  form,  the  organ  of  locomotion  has  disap- 
peared ;  this  is  obviously  an  instance  of  true  retro- 
gression, for  in  young  specimens  a  vibratory  crown 
still  persists. 

The  Nemathelminthes  contain  such  round  worms 
as  these  common  intestinal  parasites :  Ascaris, 
Oxyuris,  Strongylus,  etc.  These  all  belong  to 
the  Nematode  group,  and  possess  a  complete 
digestive  tube.  Gordius,  however,  exhibits  orgegis 
which  are  reduced  in  a  marked  degree ;  in  the 
adult  animal  the  buccal  orifice  of  the  digestive 
tube  is  closed,  and  the  posterior  part  of  the 
intestine  has  disappeared,  although  in  the  young 
worm  the  alimentary  canal  is  complete.  This 
modification,  though  incomplete  and  appearing 
only    in    the    adult    life    of    Gordius,  is    complete 


EUDIMENTARY  ORGANS  OF  ANIMALS  129 

and  permanent  in  some  other  Nematodes.  In 
Echinorhynclius,  for  instance,  the  digestive  tube 
is  absent,  and  nourishment  is  obtained  by  mean^ 
of  osmotic  soaking  through  the  body  walls. 

The  Annelids  comprise  the  annulated  sea-worms 
and  forms  like  the  common  earth-worm  (Zumbricus 
terricola).  In  these  creatures  we  will  take,  first, 
the  development  of  the  eyes.  In  Oligochcetes,  which 
for  the  most  part  live  in  soil  or  mud,  the  organs  of 
sight  are  greatly  reduced.  The  Naidonwrphce  alone 
have  eyes.  The  Archiannelida — Histrior  (a  para- 
site), for  instance — possess  eyes  when  young,  but 
in  the  adult  state  the  eyes  have  greatly  degener- 
ated. As  a  rule,  the  eyes  of  the  Polychcetes  are 
well  developed,  and  in  some  of  them  quite  re- 
markably so.  In  species,  however,  which  do  not 
move  about  much,  the  eyes  are  merely  represented 
by  small  pigmented  spots. 

We  may  mention,  too,  the  Gephyreans,  without 
pledging  ourselves  as  to  their  exact  relationship. 
Bon^llia  viridis,  the  history  of  which  is  well  known 
and  of  great  interest,  belongs  to  this  group.  The 
male  Bonellia  lives  as  a  parasite  on  the  proboscis,  or 
in  the  gullet  or  the  nephridium  of  the  female.  It 
is  flat  and  small,  and  has  neither  mouth,  arms,  nor 
circulatory  system.  All  the  organs  remain  as  in 
the  larval  condition,  with  the  exception  of  the 
genital  organs,  which  are  fully  developed. 

Bonellia  and  Dinophilus,  a  rotifer,  of  which  the 
male  is  degenerate,  exhibit  a  progressive  degenera- 

I 

; 


130    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

tion  of  all  the  organs  not  connected  with  reproduc- 
tion. Degeneration  has  made  furthest  advances  in 
Bonellia,  which  affords  a  striking  example  of  retro- 
gression side  by  side  with  development. 

Investigations  of  animal  series  such  as  these 
might  well  be  continued  throughout  the  various 
classes  and  groups,  showing  the  existence  of  rudi- 
mentary organs  in  all.  We  will  restrict  ourselves 
here,  however,  to  mentioning  the  larger  subdivisions 
only,  taking  one  example  from  each  group. 

3.  The  EchiTwderms.  —  This  order  comprises 
star-fish  or  asterids,  sea-urchins,  Crinoids  and 
Holothuria  or  trepangs. 

Of  these  we  will  take  the  star-fish,  and  proceed 
to  examine  its  digestive  tube.  Under  normal 
circumstances,  the  intestine  terminates  in  a  dorsal 
anus,  centrally,  or  slightly  excentrically  placed,  pre- 
ceded by  a  very  short  but  well-developed  rectum. 
Asteracanthion  and-  Solaster  furnish  good  examples 
of  this.  In  some  kinds  of  Asterids — in  Astropecten 
aurantiaeus,  for  instance — the  anus  no  longer  exists, 
and  the  rectum,  having  become  useless,  is  greatly 
reduced,  though  still  exhibiting  signs  of  its  original 
condition. 

4.  Mollusca.  —  Of  the  group  of  Mollusca  we 
will  take  the  Gastropods  and  the  Cephalopods  as 
examples. 

There  are  two  kinds  of  Gastropods,  straight 
and  twisted,  the  former  representing  the  primitive 
type.     The  straight  types — such  as  Chiton,  Patella^ 


RUDIMENTARY  ORGANS  OF  ANIMALS 


131 


HalioteSj  and  Fissurella — are  bilaterally  symmetri- 
cal, while  in  other  Gastropods  the  spiral  twisting 
of  the  body  causes  a  progressive  diminution  of  the 
organs  situated  on  the  side  towards  which  the 
twisting  occurs ;  the  organs  of  the  left  side  may 
therefore  become  smaller,  and  finally  atrophy 
almost  completely. 

The  internal  shell  of  the  Cephalopods  furnishes 
a  striking  example  of 
a  rudimentary  organ.^ 
The  Nautilus  (fig.  54) 
has  a  shell,  the  spiral 
coils  of  which  are 
pressed  tightly  against 
one  another.  The  spiral 
is  divided  into  a  series 
of  chambers  by  means 
of  partitions,  each  par- 
tition being  provided 
with    an    aperture   for  f.^.  ^A.-Nautnu,  pompuius. 

fhp  flHmiQQinn  nf  f>iP  ^  terminal  chamber  of  the  shell  ;  a,  body 
Cne     aamiSSlOn     or     tne        of  the  anlmal;  /.siphon;  r,mantlefold. 

siphon  (Z.).  The  shell  <Af*«^ ^>«^«°) 
of  Spirula  (fig.  55  and  56,  a),  a  creature  still  existing, 
is  only  partially  curled  round ;  the  last  chamber  of 
the  shell  is  very  small,  and  only  encloses  a  part  of 
the  animal,  most  of  which  remains  outside  the  shell, 
and  partially  covers  it  by  the  mantle  (p.),  the 
shell  being  therefore  partly  external  and  partly 
internal.      On  examining  the  fossil  species  ;  Spiru- 

1  W.  Boas,  Lehrhuch  der  Zoologie.     Jena,  1894. 


132    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


lirostra  (fig.  56,  B),  Belemnites  (fig.  56,  c),  and 
Conoteuthis  (fig.  56,  d),  a  progressive  simplifi- 
cation of  the  shell  may  be  observed,  the  latter 
becoming  less  and  less  coiled,  until  finally  the 
original  shell  is  transformed  into  a  straight 
chambered  portion  surmounted  by  a  stiliform 
process.     Degeneration  is  principally  exhibited  in 


Fig.  55.—Spirula 
prototypos. 
,  body  of  the  animal; 
p,  mantle  fold ;  c, 
shell,  partly  internal, 
partly  external. 
(After  Owen.) 


Fig.  66  —Shells  of  various  Cephalopods 
A,  Spirula;  B,  Spirvlirostra  ;  C,  Belemnites; 
teuthis;   E,    Ommatostrephes ;    F,  Loligopsis 
Boas.) 


D,  Cmo. 
.    (After 


the  segmented  part,  which  becomes  more  and  more 
reduced.  In  Ommatostrephes  (fig.  56,  e),  and  in 
all  existing  species  (such  as  Loligopsis,  fig.  56,  f), 
with  the  exception  of  those  just  mentioned  above, 
degeneration  has  become  complete.  The  shell  no 
longer  contains  a  cavity  for  the  reception  of  the 
animal,  and  the  phenomenon  already  mentioned 
with  regard  to  Spirula — the  development  of  the 


RUDIMENTARY  ORGANS  OF  ANIMALS  133 

organism  round  about  the  shell — has  become 
more  marked  in  character.  The  shell  has 
become  internal  instead  of  external,  and  forms 
the  so-called  cuttle-bone.  This  structure,  being 
only  the  vestige  of  what  was  originally  an 
external  shell,  must  be  regarded  as  a  reduced 
organ. 

5.  Arthropoda. — The  group  of  Arthropods  com- 
prises the  Myriopoda,  the  Crustacea,  the  Arachnida, 
and  the  Insecta. 

Instances  of  rudimentary  organs  are  very  common 
among  the  Crustacea,  but  our  investigations  with 
regard  to  the  appendages  of  the  cray-fish  were  so 
thorough  that  we  will  give  examples  from  another 
group,  that  of  the  Insecta,  instead. 

Insects  are  characterized  by  the  possession  of 
three  pairs  of  legs  and  two  pairs  of  wings.  The 
organs  of  flight  exhibit  a  multitude  of  special 
adaptations,  and  numerous  instances  of  degenera- 
tion are  exhibited.  In  the  Neuroptera  (dragon 
flies),  Hymenoptera  (saw  flies  and  bees),  and  Lepi- 
doptera  (moths  and  butterflies),  the  four  wings  are 
generally  all  alike  and  fully  developed.  In  the 
Diptera  (flies),  the  posterior  wings  have  atrophied 
and  are  represented  by  two  reduced  organs,  the 
"  balancers,"  these  being  absent  in  certain  types. 
The  Strepsiptera  form  a  group  of  which  compara- 
tively little  is  yet  known;  the  larvae  live  in  the  nests 
of  bees ;  the  females  have  no  wings ;  in  the  males 
the  anterior  wings  are  rudimentary  and  the  pos- 


134    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

terior  wings  are  fully  developed.  In  the  group  of 
Coleoptera  (scarabs,  cockchafers,  longicorns,  and 
beetles),  the  anterior  wings  ("wing  covers")  have 
become  very  resistant,  and  constitute  a  perfect 
cuirass  covering  and  protecting  the  abdomen,  and 
the  posterior  wings,  the  organs  of  flight,  are  folded 
beneath  the  wing  covers  when  in  repose.  In  the 
Scarabsei  and  the  Calosoma  the  posterior  wings 
are  much  reduced.  In  other  Coleoptera  the  wing 
covers  are  united  by  their  inner  edges,  thus  render- 
ing all  movement  of  the  underlying  wings  ineffectual 
and  useless.  The  result  may,  as  in  Gibbium, 
be  the  total  atrophy  of  these  under  wings.  In 
the  Staphylinidse,  the  upper  wall  of  the  abdomen 
is  so  strong  that  the  protection  of  wing  covers  is 
unnecessary;  consequently  the  wing  covers  have 
degenerated  into  little  lamella  which  cover  only 
the  anterior  quarter  of  the  body.  In  the  female 
Lampyris  (glow-worm),  the  wings  have  totally  dis- 
appeared. The  Orthoptera  (cockroaches  and  ear- 
wigs) exhibit  a  great  number  of  variations  in  the 
organs  of  flight.  In  cockroaches  all  four  wings 
are  usually  fully  developed  ;  in  some  specimens, 
however,  and  these  principally  female,  a  very  pro- 
nounced degeneration  of  all  the  wings  may  be 
observed.  The  anterior  wings  of  the  Forficulides 
are  reduced.  In  the  group  of  Phasmidae  we  find 
side  by  side  with  species  of  the  genus  Bacillus, 
which  have  no  wings  and  look  like  dried  twigs, 
Phyllmm  siccifolium  to  which  the  large  green  and 


RUDIMENTAKY  ORGANS  OF  ANIMALS  135 

yellow  wings  give  the  appearance  of  a  leaf.      The 
neuters  of  Termites  have  no  wings. 

6.  Vertebrates. — Among  Vertebrates  the  genito- 
urinary system  contains  a  great  number  of  rudi- 
mentary organs.  In  order  to  fully  understand  the 
nature  of  these  organs,  it  will  be  necessary  to 
glance  through  the  ontogenetic  and  phylogenetic 
development  of  the  system. 

The  first  stage  in  the  formation  of  the  kidney- 
system  is  the  pronephros  (fig.  57,  a).  This  primitive 
organ  consists  of  intricate  canals  (a,  a,  a),  opening 
into  the  body  cavity  at  the  point  where  the  glomeruli 
are  formed  on  the  sub-intestinal  vein.  All  these 
canals  originally  had  apertures  to  the  exterior. 
Later  on,  however,  these  uriniferous  tubules  became 
connected  with  one  single  excretory  canal  (c.)  opening 
into  the  cloaca  (ce.).  The  primitive  genital  gland  was 
situated  close  to  the  pronephros.  In  process  of  time 
the  mesonephros  replaced  the  pronephros  (fig.  57,  b); 
in  origin  it  was  quite  distinct  from  the  pronephros, 
its  appearance  being  that  of  a  secretory  urinary 
gland  (g.)  and  its  secretory  canal  (c.)  (segmental 
duct),  was  the  same  as  that  of  the  pronephros. 
The  urinary  system  thus  formed,  became  and .  still 
continues  to  be,  closely  connected  with  the  genital 
gland,  the  discharging  canals  of  which  passed 
through  the  mesonephric  kidney  in  order  to  find 
a  passage  to  the  exterior  through  the  segmental 
duct.  During  the  mesonephric  stage,  another  canal 
was   formed   which   started   from   the   cloaca    and 


136    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

opened  out  into  the  general  body  cavity,  this  was 
Mtiller's  duct  (m.). 


ce 


Fig.  57. — Development  of  the  Urino-genital  system  in  higher  Vertebrates. 

A.  Pronephric  stage,    a,  tubules;  c,  excreting  duct;  ce,  cloaca. 

B.  Mesonephric  stage.  G,  mesonephros;  r,  remains  of  pronephros  ;  c,  excreting 
canal;  w,  neutral  genital  gland ;  if,  Mtiller's  duct;  ce,  cloaca, 

C.  Metanephric  or  adult  stHge,  in  the  male.  R,  permanent  kidney;  U,  ureter; 
F,  bladder;  T,  testis;  e and  r,  epididymis  and  vas  deferens  ;  s,  vas  deferens; 
hm,  hydatid  of  Morgagni;  h,  hydatid ;  p,  paradidymis;  urn,  uterus  masculinus. 

D.  Metanephric  or  adult  stage  in  the  female.  R,  permanent  kidney ;  U,  ureter  ; 
F,  bladder;  0>  ovary;  p  and  p\  parovarium  and  paraophorcn;  w,  Weber's 
organ  ;  F,  vagina;  «,  uterus;  t,  aperture  of  Fallopian  tube;  h,  hydatid. 

The  mesonephros,  however,  was  not  the  perma- 
nent kidney.  In  the  course  of  time  the  metanephros, 
the  permanent  excretory  gland,  was  developed  (fig. 


KUDIMENTARY  ORGANS  OF  ANIMALS  137 

57,  c  and  d).  This  development  was  attended  by 
important  modifications,  further  instances  of  de- 
generation taking  place,  and  fresh  organic  connec- 
tions being  established.  An  examination  of  the 
male  and  female  sexes  is  necessary  in  order  to 
explain  what  really  took  place. 

In  the  male  (fig.  57,  c)  the  mesonephros  began 
to  atrophy ;  that  part  which  was  connected  with 
the  testes  was  transformed  into  the  epididymis  and 
the  vas  deferens  (e.  and  v.)  of  the  true  male  genital 
organs ;  the  remaining  part  atrophied,  and  when  the 
permanent  organization  was  attained,  only  persisted 
in  the  form  of  a  paradidymis  (p.)  and  a  hydatid 
(h.),  organs  which  are  quite  without  function  in  the 
adult  state. 

The  discharging  canal  which,  during  the  mesone- 
phric  stage  was  common  to  both  urinary  and  genital 
glands,  remained  simply  in  connection  with  the 
testes,  and  then  became  the  vas  deferens  (s.)  of 
which  the  cloaca  having  disappeared,  the  terminal 
extremity  became  gradually  individualized.  The 
permanent  kidney  (R.)  became  connected  with  a 
freshv  canal — the  ureter  (u.)  which  was  formed  by 
degrees  at  the  expense  of  the  primitive  discharging 
canal,  and  subsequently  became  separated  from  the 
latter  in  order  to  empty  itself  into  the  bladder  (v.). 

These  changes  were  attended  by  a  remarkable 
evolution  of  Muller's  canal,  which  first  increased 
in  size,  and  then  at  a  certain  point  proceeded  to 
atrophy  until  all  that  remained  were  the  distal  and 


138    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

proximal  extremities  in  the  shape  of  reduced  organs, 
the  hydatid  of  Morgagni  (hm),  and  the  uterus 
masculinus  (um),  neither  of  these  being  functional. 
The  intervening  part  of  the  canal  remained,  and 
formed  a  canal  which  has  been  described  by  Gasser. 

We  see  then,  that  in  the  genito-urinary  apparatus 
of  an  adult  male  there  are : 

(1)  Organs  which  have  come  into  existence  at 
different  times,  but  which  have  retained  their 
original  functions,  viz. :  the  testes,  the  kidney 
(metanephros),  and  the  ureter;  (2)  Organs  which 
are  functional,  but  of  which  the  ultimate  function 
differs  from  the  original,  viz. :  the  epididymis  and 
the  vas  deferens ;  (3)  Reduced  organs,  vestiges  of 
what  were  formerly  active  organs,  viz. :  the  hydatid 
and  the  paradidymis  ;  (4)  Reduced  organs,  vestiges 
of  Mlillerian  canal  which  only  became  active  in 
the  female,  viz.  :  the  hydatid  of  Morgagni  and  the 
male  uterus. 

In  the  female  (fig.  57,  d),  the  development  of 
the  renal  part  is  similar  to  that  we  have  just 
described  in  the  male.  Taking  first  that  part  of 
the  mesonephros  which  became  connected  with  the 
genital  gland  and  the  corresponding  discharging 
canal,  we  find  that  the  canal  disappeared  with  but 
rare  exceptions  in  which  it  formed  Gartner's  duct  (^.), 
the  lower  part  persisting  in  the  form  of  a  rudiment 
(Weber's  organ  {w)) ;  the  upper  part  became  reduced 
to  a  small  tissue  which  surrounded  the  paraovarium 
(29.),  and  the  paraophoron  (p.)  vestiges  of  what  was 


RUDIMENTARY  ORGANS  OF  ANIMALS  139 

formerly  the  mesonephros.  Mliller's  canal  became 
considerably  enlarged  ;  it  formed  the  vagina,  the 
uterus,  and  the  Fallopian  tubes ;  at  the  upper  end 
it  was  connected  with  the  hydatid,  a  vestige  of  the 
mesonephros. 

It  is  plain  then  that  the  genital-urinary  apparatus 
of  the  female  comprises  some  organs  of  which  the 
functions  remain  unchanged :  the  ovaries,  the  per- 
manent kidney,  the  Fallopian  tubes,  the  uterus,  the 
vagina,  and  the  ureter,  and  some  rudimentary  organs, 
vestiges  of  what  were  once  active  organs ;  the 
paraovarium,  the  paraophoron,  hydatid  and  Weber's 
organ. 

The  complicated  development  of  this  system 
becomes  clear  if  a  careful  study  is  made  of  the 
history  of  the  genito-urinary  apparatus  of  the  entire 
series  of  vertebrates. 

It  appears  that  the  various  phases  through  which 
the  embryos  of  the  higher  vertebrates  pass  are 
stages  similar  to  those  which  may  be  observed  in 
the  adult  lower  vertebrates. 

The  principle  of  recapitulation,  that  the  embry- 
onic stages  of  higher  animals  recapitulate  successive 
stages  attained  by  the  adults  of  lower  animals, 
receives  a  full  corroboration  from  the  facts  we  have 
been  displaying. 

AmpMoxus,  for  instance,  remains  still  at  the 
pronephric  stage :  fish  as  a  rule  have  a  mesonephric 
or  permanent  kidney.  Some  lizards  (Lacerta)  up 
to  the  age  of  two  years  make  use  of  the  mesone- 


140    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

phros  as  the  organ  for  eliminating  urine,  but,  at 
the  same  time  they  make  use  of  the  metanephros 
which  is  also  functional. 

In  Chamceles  the  mesonephros  remains  partially 
active  throughout  life.  Both  birds  and  mammals 
completely  lose  the  mesonephros,  and  in  the  adult 
stage  the  metanephros  is  the  only  active  kidney. 

This  is  not  the  place  in  which  to  complete  our 
study  of  the  recapitulation  theory,  and  we  shall 
have  to  recur  to  it  later  on ;  but  it  was  impossible 
to  describe  the  numerous  rudimentary  organs  of 
this  system  without  taking  a  comprehensive  glance 
at  the  individual  and  specific  development  of  the 
whole.  This  investigation,  moreover,  raises  another 
question. 

It  has  just  been  shown  that  the  epididymis  is 
only  a  vestige  of  the  mesonephros,  but  in  this  case 
it  cannot  be  said  that  there  has  been  degeneration ; 
what  has  happened  is  that  an  organ  has  been  trans- 
formed, and  that  one  function  has  been  replaced  by 
another.  According  to  some  authorities  the  supra- 
renal capsule,  an  organ  of  unknown,  but  doubtless 
essential  function,  is  the  result  of  the  transformation 
of  the  pronephros.  If  this  theory  be  ultimately 
established,  it  will  furnish  a  second  example  of  what 
we  have  stated  above.  The  thyroid  gland  may 
and  ought  to  be  investigated  from  this  standpoint. 
The  various  component  parts  of  this  organ  had 
no  original  connection.  In  higher  vertebrates  only 
the  central  part  of  the  organ  appears  to  be  similar 


RUDIMENTARY  ORGANS  OF  ANIMALS  141 

to  the  same  organ  in  the  whole  series  of  vertebrates. 
In  man  the  mesial  part  of  the  thyroid  gland  is 
reduced,  but  it  cannot  correctly  be  said  that  the 
organ  itself  is  degenerating.  The  contrary  may 
even  be  asserted,  for  all  we  know  upon  the  subject 
goes  to  prove  that  in  mammals  the  thyroid  gland 
is  formed  and  established  at  the  expense  of  the 
primitive  rudimentary  organ  of  which  all  the 
morphological  and  embryological  connections  are 
changed.  This  secondary  development,  entailing 
the  loss  of  a  reduced  ancestral  organ,  is  attended 
by  a  functional  modification  of  great  importance. 
The  primitive  function  probably  discharged  by  the 
gland  has  given  place  to  another  and  rather  vague 
function,  but  one  which  is  connected  with  the 
breaking  down  of  toxic  matter  formed  by  living 
tissues. 

An  examination  of  the  vertebrae  of  vertebrates 
will  show  the  existence  of  rudimentary  organs 
throughout  the  whole  group,  each  type  and  each 
individual  among  the  vertebrates  exhibiting  special 
degeneration.  In  order  to  briefly  demonstrate  this 
point,  take  two  quite  different  types  of  which  all 
the  vertebrae  are  well  known  and  can  therefore 
be  examined  without  difficulty :  man  and  the 
frog. 

We  know  that  the  construction  of  the  primitive 
bony  vertebra  was  as  follows  : — a  centrum,  carrying 
neurapophyses,  an  intercentrum,  carrying  haema- 
pophyses,  and  a  pair  of  unforked  ribs. 


/ 


142    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

Each  phase  in  the   evolution   of  this   primitive 
vertebra  has  been  attended  by  degeneration. 
In  man  the  vertebral  column  consists  of : — 

(a)  Seven  cervical  vertebrae. 

(b)  Three  dorsal  vertebrae. 

(c)  Five  lumbar  vertebrae. 

(d)  Three  sacral  vertebrae. 

(e)  Six  coccygeal  vertebrae. 

Each  one  of  these  vertebrae  exhibits  important 
modifications,  and  shows  signs  of  degeneration. 
The  proatlas  is  represented  by  its  intercentrum  only. 
The  atlas  consists  of  a  centrum  and  neurapophyses, 
but  there  is  no  zygapophyses ;  it  possesses  one  pair 
of  ribs  and  hsemapophyses  in  a  reduced  condition. 
The  axis  consists  of  the  same  elements  but  carries 
postzygapophyses. 

The  four  following  cervical  vertebrae  consist  of 
the  same  elements,  but  carry  both  zygapophyses 
and  postzygapophyses.  In  man  all  the  cervical 
vertebrae,  with  the  exception  of  the  first  (the 
proatlas)  have  lost  the  intercentrum.  The  next 
vertebrae  which  is  generally  regarded  as  the  seventh 
cervical  vertebra,  consists  of  the  same  elements,  but 
it  ought  to  be  regarded  as  the  first  dorsal  vertebra. 
The  vertebral  artery  and  the  sympathetic  nerve 
trunks  accompanying  it,  do  not  pass  through  the 
vertebral  canals ;  in  some  cases  these  do  not  exist. 
The  so-called  seventh  cervical  vertebra  has  occasion- 
ally one  fully  developed  rib  which  articulates  with 
the  sternum  like  the  ribs  of  true  dorsal  vertebrae. 


RUDIMENTAKY  OKGANS  OF  ANIMALS  143 

There  are  thirteen  dorsal  vertebrae.  With  the 
exception  of  that  which  has  just  been  described, 
they  consist  of : — a  centrum,  neurapophyses,  and  a 
pair  of  fully  developed  ribs ;  the  intercentra  has 
completely  disappeared,  and  the  hsemapophyses, 
which  form  the  head  and  neck  of  the  rib,  are  in  a 
reduced  condition.  In  the  five  lumbar  vertebrae 
which  follow,  the  rib  disappears,  or  to  speak  more 
accurately,  the  transverse  processes  are  all  that 
remain  of  what  were  the  ribs,  and  have  ossififed 
with  the  vertebra. 

The  sacrum  is  a  region  profoundly  modified  to 
support  the  pelvic  basin.  It  is  formed  by  the 
fusion  of  five  vertebrae,  each  consisting  of  a 
centrum,  neurapophyses,  and  short  bicipital  ribs. 
The  .first  three  vertebrae  are  the  true  sacrals,  as 
these  alone  support  the  basin ;  the  two  following  are 
really  caudal  vertebrae  in  process  of  fusion  with  the 
sacrum.  In  monkeys  other  than  anthropoids  there 
are  really  only  three  sacral  vertebrae  and  these  are 
at  once  succeeded  by  the  tail.  In  man,  the  taiP 
consists  of  six  vertebrae  of  which  the  two  first — 
which  consist  of  a  centrum  and  neurapophyses  and 
of  bicipital  ribs — have  fused  with  the  sacrum, 
while  the  four  remaining  lower  vertebrae,  which 
consist  of  only  the  centra  (the  first  still  exhibits 
rudimentary  neurapophyses)  have  fused,  and  form 
the  coccyx. 

^  See  Albrecht,  La  queue  chez  Vhomme  (Bull.  soc.    Anthrop. 
Briix.,  vol.  iii.,  1884-1885, 


144    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

2.  The  Frog. — The  vertebral  column  in  the 
frog  consists  of  nine  vertebrae  and  the  coccyx 
(urostyle). 

The  first  vertebra  (the  proatlas),  which  is  fully 
functional,  retains  a  centrum  and  two  well-developed 
neurapophyses,  but  the  transverse  processes,  the 
intercentrum,  the  hsemapophyses  and  the  ribs  have 
all  disappeared. 

The  eight  following  vertebrae  each  have  a 
centrum,  neurapophyses  and  transverse  processes 
which  at  least  partially  represent  the  ribs. 

The  coccyx,  whether  it  be  formed  by  the 
lengthening  out  of  the  last  caudal  vertebra  or  by 
the  fusion  of  several,  is  undoubtedly  part  of  the 
vertebral  column  which  has  been  transformed. 
The  coccyx  of  the  frog  is  equal  in  length  to  the 
whole  of  the  remaining  part  of  the  vertebral 
column,  and  is  fully  functional ;  it  serves  as  a 
support  to  the  pelvic  region  and  fulfils  the  part  of 
a  sacrum  from  the  physiological  point  of  view. 
The  coccyx,  excepting  at  its  commencement,  con- 
sists of  only  one  centrum,  or  of  several  fused 
centra,  all  other  elements  of  the  vertebrae  have 
disappeared.  Here  then  is  an  animal  in  which  the 
modifications  of  the  vertebral  column  have  been 
attended  by  the  following  retrogressive  phenomena  : 
in  the  upper  part  of  the  vertebral  column  from 
three  to  five  parts  of  the  vertebrae  have  disappeared  ; 
in  the  lower  half,  all  the  parts,  excepting  one,  are 
gone. 


RUDIMENTARY  ORGANS  IN  PLANTS  145 

Section  II. 
Rudimentary  Orgatis  in  Plants. 

We  have  just  glanced  through  a  series  of 
rudimentary  organs  in  animals,  and  many  more 
examples  might  easily  have  been  furnished,  but, 
when  dealing  with  rudimentary  organs  in  the 
vegetable  world,  much  greater  difficulty  is  met 
with.  In  plants,  the  elimination  of  non-functional 
organs  is  usually  complete,  and  the  vestiges  left  are 
insignificant  and  hard  to  recognize.  We  can  find, 
however,  amongst  the  various  groups  of  the  vege- 
table world,  and  especially  among  the  Phanerogams, 
some  instances  of  reduced  organs. 

§   3.  Budimentary  organs  in  various  groups  of 
plants. 

1.  Algae. — On  the  surface  of  sea-wrack  (Fucus) 
may  be  found,  distributed  in  large  numbers,  little 
crypts  (conceptacles)  with  hairs  growing  out  of 
them.  On  certain  parts  of  the  plant,  these  crypts 
represent  the  organs  of  reproduction,  producing 
eggs  and  spermatozoa ;  in  other  parts  they  fulfil 
no  known  function  and  may  be  regarded  as  con- 
ceptacles arrested  in  the  course  of  development. 
The  fact  that  in  other  Fucacise  {Splachnidium\ 
fertile  conceptacles  are  distributed  over  the  entire 
surface  of  the  plant  adds  support  to  this  theory. 

K 


146    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

2.  Mushrooms. — Among  the  Peronospora  and 
the  Saprolegnese  there  originally  existed,  besides 
the  various  sexual  means  of  propagation,  a  typical 
reproductive  process,  including  eggs,  and  antheridia, 
consisting  of  male  branches  separated  by  a  cell- 
wall  from  the  rest  of  the  organism. 

In  PythiuDi,  for  instance,  in  wliich  this  primitive 
stage  may  be  observed,  an  actual  fecundation  takes 
place,  the  protoplasmic  contents  of  the  antheridia 
being  injected  into  the  ova. 

In  other  species,  the  organs  of  reproduction  have 
undergone  a  more  or  less  complete  degeneration. 
In  Phytophthora,  a  small  part  only  of  the  male 
protoplasm  passes  into  the  ova.  In  some  species 
of  Saprolegnia  and  Achlya,  the  male  branch  con- 
tinues to  attach  itself  to  the  ova,  but  the  membrane 
between  them  remains  intact,  and  consequently 
protoplasmic  communication  is  not  established. 

In  other  species,  the  antheridia  are  very  short, 
and  do  not  even  touch  the  female  cells. 

In  Leptomitios,  which  exhibits  an  advanced  stage 
of  atrophy,  the  female  organs  are  not  discernible, 
and  reproduction  is  carried  on  completely  asexu- 
ally.i 

3.  Bryophyta. — In  the  germination  of  a  certain 
number   of    Hepaticse,   belonging    to    such    widely 

^  For  further  details  see  A.  de  Bary,  Vergleicheiulc  Morpliologic 
tend  Biologic  der  Pilzc,  Leipzig,  Engelraan,  1884,  W.  Zopf,  Die 
Pilze.  In  Schenk's  Handbuch  der  Botaniky  4.  Bd.,  Breslau, 
Treweiidt.  1890. 


RUDIMENTARY  ORGANS  IN  PLANTS  147 

separated  genera  as  Blasia,  Radula,  and  Preissia, 
four  cells  of  equal  size  are  formed,  arranged  round  a 
centre.  One  only  of  these  cells  proceeds  to  develop 
into  a  plant,  and  the  others  simply  atrophy.  In 
all  probability  the  Hepaticse  have  sprung  from 
some  ancestor,  in  which  each  spore  gave  rise  to 
four  individuals. 

4.  Pteridophyta. — According  to  Farlow,^  in- 
stances of  apogamy — the  loss  of  sex — such  as  have 
just  been  described  as  existing  among  mushrooms, 
are  also  exhibited  in  certain  ferns.  In  some 
species,^  the  eggs  are  not  fertilized,  but  the  organs 
of  reproduction  still  persist  in  a  reduced  condition ; 
in  other  species  there  are  no  spores,  and  the 
prothalli  spring  directly  from  the  leaves  (apospory). 

5.  Phanerogams. — Some  of  the  Phanerogams — 
Bilene  (fig.  58),  Melandryum,  (fig.  59),  Aspara- 
gus, etc.  —  exhibit  unisexual  flowers,  but  have 
obviously  sprung  from  species  of  which  the 
flowers  were  hermaphrodite. 

In  Silene  maritima  (fig.  58)  there  are  herma- 
phrodite flowers  (fig.  58,  b),  and  also  unisexual 
flowers.  The  female  flowers  (fig.  58,  A)  still  possess 
some  tiny  stamens,  each  of  which  is  provided  with 
filaments  and  anthers  in  a  state  of  degeneration. 
The    male    flowers    have    non  -  functional   pistils, 

^  W.  Farlow,  Ueher  ungeschlecMliche  Erzeugung  von  Keimpflanz- 
clien  an  FarnprotJiaUien.     Bot.  Zeit.,  1874,  p.  180. 

2  A.  de  Bary,  Ueber  apogame  Fame  u.s.w.  Bot.  Zeit.,  1878, 
p.  449. 


148    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 


consisting  of  an  ovary  stylus  and  stigma  in  reduced 
conditions. 

In  Melandryum  (fig.  59,  a) 


Fig.  58. — Flowtrs  of  Sileiie  mariiima. 

A,  female  flower  with  rudimentary  stamens ; 

B,  hennapluodite  flower. 


the  female  flowers 
retain  only 
vestiges  of 
stamens,  and 
the  pistils  of 
the  male  flowers 
are  reduced  to 
mere  filaments. 
In  Asparagus 
officinalis  all  the 
transition  stages 
between  herma- 
phrodite and  unisexual  flowers  may  be  observed : 
in  the  unisexual 
flowers,  the  organs 
of  the  opposite  sex 
still  exist,  though 
in  various  stages  of 
degeneration.  In 
many  unisexual 
flowers  which  have 
sprung  from  herma- 
phrodite flowers  — 
Valeriana  dioica, 
for  instance  —  no 
traces  of  the  non- 
functional organs  remain. 

Among    the    Phanerogams,   rudimentary 


Fig.  59. — Flowe-'S  of  Melatidryttm  album. 
A  (to  the  left),  a  female  flower;  to  the  right,  a 
male  flower  ;  e,  rudimentary  stamens  form- 
ing a  circle  at  the  base  of  the  ovary. 


RUDIMENTARY  ORGANS  IN  PLANTS  149 

appear  not  only  in  the  reproductive  organs,  but  in  the 
accessory  organs  of  the  flower — the  calix  and  corolla. 
Many  of  the  Umbelliferae  exhibit  a  reduced  calyx. 
The  corolla  persists,  though  in  a  very  reduced 
state  in  cleistogamous  flowers  {i.e.  flowers  which 
never  open,  and  which  are  self -fertilizing) — such  as 
the  Oxalis,  Impatiens,  Violet,  etc.  The  corollas  of 
the  winter-opening  flowers  of  Stellaria  media  are 
much  reduced,  and  for  a  very  obvious  reason — 
the  corolla  exists  only  for  the  attraction  of  insects, 
and  there  are  no  insects  at  that  time  of  the  year. 

§  4.  Reduced  organs  in  the  vegetative  apparatus 
of  the  Phanerogams. 

We  have  seen  that  reduced  sexual  organs  are 
exhibited  among  the  various  groups  of  plants,  and 
we  will  now  mention  a  few  instances  of  reduced 
organs  in  the  vegetative  apparatus  of  the  Phan- 
erogams. 

1.  The  embryo  within  the  ripe  seed  of  Phanerogam's 
contains  a  rudimentary  root  which  develops  during 
germination.  In  certain  Nympheacese — Nelumhiwm 
Euryale  and  Victoria — this  root  never  properly 
develops. 

In  other  aquatic  plants  degeneration  has  gone 
further  ;  in  the  embryo  of  Utricularia,  for  instance, 
the  root  has  entirely  disappeared.^ 

^  For  further  details  relating  to  the  roots  of  the  Nympheaceae  and 
the  Utricularia  see  Goebel  in  Pflanzenhiologische  Sohilderungen, 
vol.  ii.,  Marburg,  Elwert,  1891-1893. 


150    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

2.  As  a  general  rule,  the  cotyledons,  which  are 
the  two  first  leaves  to  appear  after  germination,  are 
formed  within  the  embryo.  The  ripe  seeds  of  some 
Anemones,  however,  contain  no  traces  of  cotyledons.^ 
These  are  formed,  nevertheless, 
after  germination,  sometimes  sprout- 
ing up  out  of  the  ground  and 
becoming  functional,  and  occasion- 
ally remaining  underground,  in 
which  case  they  are  quite  small, 
without  chlorophyll  and  non- 
functional {Anemone  nemorosa); 
these  underground  leaves  may 
fairly  be  regarded  as  rudimentary 
organs. 

3.  In  Lathyrus  Nissolia  {^g.  60) 
there  are  some  very  small  stipules 
of  unimportant  function,  at  the 
base  of  the  simplified  leaves  ;  these 
reduced  stipules  are  occasionally 
absent  altogether. 

4.  The  foliage  organs  in  the 
adult     Oxalis     bulpeurifolia     are 

Fig.  60.— Seedling  of  i       i  i  i 

Lathyrus  Nissoiia.  merely  represented  by  enlarged 
leaflets.  These  phyllodes  bear  reduced  leaflets 
which  rapidly  disappear.  In  an  adult  specimen  of 
the  Acacia  which  has  phyllodes,  these  reduced 
leaves  are  absent. 

^  E.  de  Janczewsky,  ^titdes  morphologiques  sur  le  genre  Anemone. 
{Revue  g4n4raU  de  hotaniquey  t.  iv.,  p.  241.) 


SURVIVALS  EXIST  IN  ALL  KINDS  OF  SOCIETIES    1 5  1 

5.  In  several  plants  the  assimilative  function 
of  the  leaves  is  lost,  either  because  the  plant  is 
parasitic  or  saprophytic,  such  as  Corallorhiza, 
Rafflesia,  Cuscuta  and  Orobanche,  or  because  the 
assimilative  function  is  relegated  to  the  stem  alone 
as  in  the  Euphorhia  of  the  desert,  Busctos,  Mamil- 
laria}  Fhyllocachcs,  Phyllanthus,  and  Miihlenbeckia, 
or  to  the  roots  as  in  Tceniophyllum} 

In  each  of  these  cases  the  leaves  are  greatly 
reduced,  and  only  serve  as  a  means  of  protection  to 
the  functional  organs,  principally  to  the  flowers 
and  buds,  but  though  very  minute  they  may  often 
be  discerned  quite  easily  on  the  young  shoots.^ 


CHAPTER  II 

SURVIVALS    EXIST   IN    ALL    KINDS    OF    SOCIETIES 

It  may  be  said  as  certainly  of  societies  as  of  other 
organisms  that  certain  modifications  have  taken 
place,  and  that  no  society  actually  represents  a 
primitive  social  organization.  All  have  been 
submitted  to  more  or  less  important  modifications 
and  have  lost  some  of   their  early  institutions  in 

1  See  fig.  51. 

2  See  further  on  the  figs,  of  PhyllocactMS  (fig.  78),  Phyllanthus 
(fig.  84),  Miihlenbeckia  (fig.  80),  and  Tceniophyllum  (fig.  81). 

^  Goebel,  Pflayizenhiologische  Sehilderungenj  Bd.  i. 


152    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

process  of  their  development.     In  many  cases  this 
can  be  historically  demonstrated. 

It  may  fairly  be  asserted  that  in  all  societies 
there  are  instances  of  survival,  i.e.  survival  of 
customs,  beliefs  and  institutions,  the  original 
character  of  which  has  so  completely  disappeared 
that  they  might  well  be  dispensed  with  altogether. 
We  shall  deal  only  with  such  survivals  as 
correspond — mutatis  mutandis — to  the  rudimentary 
organs  of  animals  and  plants. 

These  survivals  are  of  two  kinds,  the  institu- 
tion itself,  such  as  the  various  corporations  of  the 
city  of  London,  which  may  still  persist,  though  in 
a  modified  condition,  or  there  may  remain  only 
traces  of  the  institution  in  forms,  ceremonies, 
symbols,  public  games  and  fetes,  customs  and 
legislative  formula. 

In  order  to  demonstrate  this  point,  it  will  not 
be  necessary  to  make  a  complete  enumeration, 
furnishing  examples  from  all  countries  of  the 
world,  or  to  draw  up  a  complete  list  of  survivals  in 
any  given  country.  It  will  be  enough  to  establish 
two  points  : 

1.  That  instances  of  survival  shall  be  shown  to 

exist  in  all  societies,  even  where  they  are 
least  likely  to  be  found. 

2.  That,  in  any  institution — that  of  the  family, 

for  instance — survivals  may  be  found  of 
all  the  former  stages  through  which  it 
passed  into  its  present  condition. 


SURVIVALS  EXIST  IN  ALL  KINDS  OF  SOCIETIES    153 

These  two  points  being  established  our  conclu- 
sions drawn  from  them  may  be  given  a  very  wide 
application. 

§  1.  Instances  of  survival  in  various  groups. 

**  It  is  a  well-known  fact,"  says  Kowalevsky, 
"that  as  the  past  gives  place  to  the  present  it 
leaves  traces  which  vary  in  number  and  impor- 
tance." ^ 

This  is  obviously  the  case  with  regard  to  most 
customs,  but  it  is  unnecessary  to  point  out  all  the 
instances  of  survival  which  abound  among  the 
peoples  of  the  countries  round  about  us.  They  are 
naturally  most  common  in  barbarous  societies 
where  the  servile  imitation  of  the  ancestor  plays  a 
much  greater  part  than  with  us.  This  is  strongly 
urged  by  Bagehot  in  the  following  passage ; — 

"  Man,"  he  says,  "  may  be  defined  as  a  creature 
of  habit.  As  he  has  done  a  thing  once,  so  he  will 
probably  do  it  again,  and  the  oftener  he  has  done 
a  thing  the  more  likely  he  will  be  to  repeat  it  in 
the  same  way,  and,  what  is  more,  to  insist  upon 
others  doing  the  same. 

"  By  means  of  counsel  and  example  he  transmits 
to  his  offspring  the  customs  he  himself  originated. 
This  is  true  of  the  human  beings  to-day  and  will 
doubtless  hold  good  for  all  time.     It  is  character- 

^  Kowalevsky,    Tableau  des  origines  de   la  propriiti  et  de   la 
familley  p.  7. 


154    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

istic  of  primitive  societies  that  sooner  or  later  most 
of  these  customs  come  to  be  regarded  as  having  a 
supernatural  sanction,  and  the  whole  community  is 
impressed  with  the  belief  that  if  the  old  tribal 
customs  are  violated,  incalculable  misfortune  will 
follow."  1 

Social  modifications  are  therefore  effected  very 
slowly  and  with  great  difficulty — stagnation  is  the 
rule,  and  progress  but  a  rare  exception,  innovators 
being  forced  to  retain  the  greater  part  of  the  old 
institutions,  introducing  only  an  inevitable  minimum 
of  change.  A  course  of  history,  or  a  careful  study 
of  the  conditions  of  social  institutions  at  an  earlier 
stage  of  development  than  our  own,  will  furnish 
numerous  instances  of  survivals.  It  now  remains 
to  be  seen  if  there  are  no  rudimentary  social  groups 
wherein  all  the  primitive  institutions  have  been 
retained,  and  which,  having  undergone  no  modifica- 
tions, exhibit  no  traces  of  degeneration.  We  shall 
further  see  if  no  form  of  society  exists  uninfluenced 
by  the  spirit  of  tradition,  and  where  institutions 
which  have  come  to  be  regarded  as  no  longer  useful, 
disappear  suddenly  and  entirely  either  by  voluntary 
dissolution  or  by  legislative  measures.  Only  in  these 
two  extreme  cases  can  the  existence  of  survivals 
be  questioned.  We  have  then  to  establish  two 
points: 

(a)  That  all  societies,  even  those  to  be  regarded  as 

^  Bagehot,  Lois  scientifiques  du  developpement  des  natioris,  p.  154, 
Bibl.  scient.  intern.,  Paris,  F.  Alcan,  1885. 


SURVIVALS  EXIST  IN  ALL  KINDS  OF  SOCIETIES    155 

primitive,  have  undergone  certain  modifica- 
tions. 
(h)  That  all  societies,  even  the  least  conservative, 
exhibit    instances   of    reduced   institutions, 
and  of  vestiges  of  institutions  which  have 
disappeared. 
We  will  take  the  second  point  first,  as  it  can  be 
more  briefly  dealt  with. 

1.  We  know  that  imitation  of  the  past  and 
respect  for  tradition  arid  custom  are  reduced  to  a 
minimum  in  modern  societies,  especially  in  the 
countries  of  the  New  World.  Even  in  these 
recently  formed  States,  however,  instance  of  sur- 
vival may  be  found. 

In  the  first  place,  there  are  legal  and  religious 
survivals  of  European  origin.  Jews  settling  in  the 
United  States  kept  up  the  practice  of  cir^imcision, 
while  Christians  introduced  the  Eucharist.  Spencer  ^ 
has  shown  that  forms  of  greeting  are  vestiges  of  a 
primitive  ceremonial  demonstrating  submission  to 
the  omnipotence  of  others.  Then  take  the  Calendar 
system  which  is  universal ;  we  know  that  the  names 
of  months  and  days  of  the  weeks  are  survivals  from 
Polytheistic  times,  and  it  seems  certain  that  both 
circumcision  and  the  celebration  of  the  Mass  are 
true  survivals  which  originated  in  religious  sacrifices. 
Independently  of  these  imitative  survivals  vestiges 
remain  in  the  United  States  of  reduced  institutions 
which  were  fully  functional  in  the  last  century. 
^  Essays  on  Progress. 


'/  ' 


156    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

Take  for  instance  the  town  meetings  of  Boston  and 
Newhaven.-^ 

When  the  inhabitants  of  a  town  attained  to  a 
certain  number,  the  town  became  a  city,  and  the 
Assembly  of  inhabitants  was  transformed  into  a 
Common  Council.  In  some  instances,  this  old 
system  persists  though  in  a  reduced  condition. 

In  Boston,  which  continued  to  be  a  town, 
governed  by  an  Assembly  of  all  the  inhabitants  up 
to  1821,  the  present  Charter  of  the  city  authorizes 
the  convocation  of  a  town  meeting  wherever  the 
Mayor  and  Aldermen  consider  it  advisable;  the 
latter,  however,  never  make  use  of  this  privilege. 

In  Newhaven  (Connecticut),  the  old  town  meeting 
continues  to  exist  side  by  side  with  the  Common. 
Council  which  was  established  in  1784,  but, 
Levermore  says,  "  This  ancient  institution  nowadays 
is  a  meeting  together  of  a  small  number  of  citizens 
to  conduct  the  business  of  several  thousands.  The 
few  people  connected  with  the  affairs  of  the  town 
(which  is  very  poor),  meet  together  to  discuss 
matters  in  a  friendly  way,  decide  what  money  is 
required  for  current  expenses,  and  then  adjourn. 
Not  one  in  seventy  of  the  inhabitants  attends  these 
meetings.  Few  know  when  they  take  place,  and 
the  papers  make  brief,  if  any,  mention  of  them." 

2.  We  have  now  to  show  that  the  simplest 
societies  have  undergone  modifications,  and  exhibit 

1  Bryce,  The  American  Commonwealth,  i.,  pp.  598  and  following. 
London,  Macmillan,  1893. 


SURVIVALS  EXIST  IN  ALL  KINDS  OF  SOCIETIES    157 

instances  of  survival.  We  will  take  as  examples 
those  rudimentary  types  most  nearly  approaching  to 
the  primitive  type,i  i.e.  the  Yeddahs  of  Ceylon,  the 
Fuegoes  of  Cape  Horn,  and  the  Australian  tribes. 

(a)  The  Veddahs,  who  have  lived  in  the  jungles 
of  Ceylon  for  centuries,  either  as  separate  families, 
or  in  groups  of  two  or  three  families,  appear  to 
have  formerly  possessed  a  much  more  complicated 
social  organization.  According  to  Max  Muller, 
they  were  not  formerly  so  low  in  the  scale  of 
humanity ;  he  says  that  their  language,  if  not 
their  blood,  betrays  their  "  distant  connection  with 
Plato,  Newton,  and  Goethe." 

In  their  language,  folk-lore,  and  clothing,  these 
retain  characteristic  vestiges  of  a  former  condition. 
Take  for  instance  the  carefully  observed  practice  of 
piercing  the  ears  of  cliildren  at  the  age  of  three  or 
four  years,  although  eventually  only  a  small  number 
of  them  could  wear  ornamenrs  in  them,  others 
having  to  be  content  with  small  pieces  of  twig, 
coiled  leaves,  or  bits  of  straw. 

"  This  custom,"  says  Deschamps,  "  is  extremely 
old,  and  we  may  suppose — as  there  is  no  other 
signification  in  it  than  the  prospect  of  ultimately 
wearing  jewels — that  it  dates  back  from  a  time 
when  the  people  were  not  in  so  low  and  destitute 

1  "Aggregates  formed  by  a  simple  repetition  of  hordes  or  clans 
without  any  such  interrelations  between  them  as  to  form  inter- 
mediate groups  between  the  whole  collection  and  the  individual 
clans."  Durckheim,  les  Regies  de  la  m&lwde  sociologique,  Paris, 
F.  Alcan,  1895. 


158    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

a  condition  as  they  are  now.  Having  in  more 
civilized  times  worn  jewels  in  their  ears,  the 
custom  of  piercing  the  ears  in  youth  persists, 
though  the  jewels  may  be  lacking."  ^ 

(b)  Bridges  says  that  according  to  a  tradition 
which  is  probably  true,  the  Fuegoes,  until  quite 
recently,  submitted  their  young  men  to  a  sort  of 
initiatory  trial  when  they  attained  to  adolescence. 
They  were  taken  into  a  hut  (the  Mna)  set  apart 
for  the  purpose,  and  there  underwent  certain  tests, 
including  a  rigorous  fast.  Bridges  adds  that  the 
Mna  was  also  the  theatre  of  mysterious  and  bizarre 
scenes  of  very  ancient  origin,  the  roles  of  which, 
now  relegated  to  men  only,  were  entirely  performed 
by  women.  Contrary  to  Giraud-Teulon  who  cites 
these  facts  as  evidence  of  the  former  existence  of 
a  matriarchy,  the  fetes  of  the  kina  seem  to  have  quite 
disappeared  from  among  the  natives  of  Orange  Bay. 

Dr  Hyades,  however,  mentions  a  survival  of  the 
old  custom.  "  The  custom  is  still  observed  of 
submitting  young  girls  to  a  fast  at  the  time  of 
puberty,  but  this  fast  is  less  severe  than  that 
already  mentioned  as  undergone  by  the  boys ;  the 
same  good  advice  is  then  given  them  by  their 
parents,  as  was  formerly  bestowed  upon  the  boys 
in  the  Kina."  ^ 

^  Emile  Deschamps,  V Anthropologic,  1891,  t.  ii.,  pp.  297  and 
following. 

2  Mission  scientifiquc  du  cap  Horn,  1882-1883,  t.  vii.  Anthro- 
pologic, Ethnographic,  by  P.  Hyades  and  J.  Deniker ;  Paris, 
Gauthier-Villars,  1891,  p.  377. 


SURVIVALS  EXIST  IN  ALL  KINDS  OF  SOCIETIES    159 

(c)  Eecent  researches  into  the  family  system 
among  the  Australian  tribes  has  brought  a  number 
of  survivals  to  light.  This  is  especially  the  case 
with  regard  to  the  careful  researches  of  Fison  and 
Howitt^  who  have  shown  that,  independently  of 
their  tribal  divisions — which  are  really  territorial 
groups — the  Australians  are  divided  up  into  clans 
or  sexual  groups  comprising  all  the  individuals  with 
the  same  Kobong.^ 

The  members  of  these  groups  are  regarded  as 
members  of  the  same  family,  and  may  never, 
under  any  circumstance,  intermarry,  under  pain 
of  being  driven  out  of  the  clan  and  hunted  like 
wild  beasts.  Sometimes  individuals  of  antagonistic 
tribes  living  at  several  hundred  miles'  distance  from 
one  another  and  speaking  different  languages  liave 
the  same  Koboruj.  The  law  of  classes  remains 
active ;  a  captor  may  not  violate  a  prisoner  belong- 
ing to  his  group,  but  a  stranger  may  enter  into 
relations  with  the  women  of  another  tribe,  so  long 
as  the  tribe  belongs  to  a  class  related  to  his  own. 
This  system  of  relationship  can  only  be  explained 
as  being  a  survival  from  a  former  period  in  which 
all  persons  with  the  same  Kohong  belonged  to  the 
same  group.     This  is  a  disputed   point,^  however, 

^  Fison  and  Howitt,  Kurnai  and  Kamilaro'i  {Journal  of  the 
Anthropological  Institute,  1884). 

^  '*The  Kohong  of  a  man  is  the  animal  or  plant,  the  name  of 
which  he  bears  and  reveres  as  a  protecting  spirit"  (Starcke). 

*  Starcke,  la  Famille  primitive  (Bibl.  sciens.  intern.,  Paris, 
F.  Alcan,  1891,  p.  22). 


160    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

for  besides  this  very  likely  hypothesis,  undoubted 
survivals  remain  of  intermarriage  by  groups  or 
sexual  groups.  In  the  writings  of  Fison  and  Ho  wit  t, 
we  find  the  two  following  instances  of  this  in  two 
tribes  which,  according  to  them,  severally  represent 
the  highest  and  lowest  in  the  scale  of  civilization, 
among  those  with  which  they  came  in  contact.^ 

{d)  The  tribe  called  Kunandaburi  was  divided 
into  two  exogamous  classes  :  Mattara  and  Yungo. 
Theoretically  all  the  Yungos  whether  male  or 
female  were  regarded  as  the  males  of  the  Mattaras, 
and  vice  versd.  In  point  of  fact,  however,  only 
one  vestige  of  the  primitive  communal  marriage 
remained — the  jus  primce  noctis  which  was  the 
prerogative  of  all  the  contemporaries  of  the 
husband  belonging  to  the  same  group. 

(e)  The  tribe  called  Narrinyeri  which  repre- 
sented a  more  advanced  stage  of  civilization,  was 
equally  divided  into  two  sexual  groups,  but  in 
reality,  marriage  was  strictly  individual.  One 
survival  remained,  however,  of  the  former  system. 
When  a  man  captured  an  alien  bride,  all  the  men 
of  his  own  generation  and  belonging  to  the  same 
group  possessed  the  right  of  jus  jprimm  noctis. 

3.  We  have  seen  that  instances  of  survival  are 
rare  in  some  countries  because  modifications  are 
only  effected  slowly,  and  in  others  because  changes 
are  effected  very  quickly  and  useless  institutions 

^  Fison  and  Howitt,  Journal  of  the  Anthropological  Inatitutef 
1882,  p.  35. 


SURVIVALS  EXIST  IN  ALL  KINDS  OF  SOCIETIES    1 6  1 

are  eliminated  root  and  branch.  It  is  in  countries 
like  England,  where  modifications  are  brought  about 
with  a  due  respect  for  old  customs  and  traditions 
that  ceremonies,  institutions  and  customs  exhibit 
the  greatest  number  of  survivals. 

§  2.  Survivals  of  ancient  forms  of  marriage  and  of 
the  family  in  Modern  Europe, 

We  think  we  may  regard  it  as  proved  that 
all  societies  exhibit  instances  of  survival,  but  in 
order  to  further  demonstrate  the  universal  character 
of  retrogressive  evolution,  we  shall  show,  by  means 
of  a  careful  study  of  one  particular  institution,  that 
vestiges  of  former  institutions  are  neither  rare  nor 
exceptional,  taking  as  examples  the  various  forms 
of  marriage  and  of  the  family  throughout  Modern 
Europe. 

1.  Forms  of  Marriage, — From  archaic  times  up 
to  our  own,  we  find  that  among  modern  nations, 
marriage  by  capture,  marriage  by  purchase,  and 
marriage  with  the  consent  of  the  woman  have  been 
successively  followed  by  marriage  by  simple  consent, 
religious  marriage  {in  facie  Ecclesice)  and  civil 
marriage,^  and  that  survivals  remain  of  all  the 
forms  of  marriage  anterior  to  civil  marriage.^ 

^  Paul  Viollet,  Histoire  du  droit  civil  Frangais,  p.  424  and 
following.     Paris,  Larose  &  Forcel,  1893. 

2  Westermarck,  The  History  of  Human  Marriage,  1892,  p.  418 
(when  the  mode  of  contracting  a  marriage  altered,  the  earlier  mode, 
from  having  been  a  reality,  survived  as  "  ceremony  "). 

L 


162    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

Marriage  hy  Capture. — Traces  of  this  early  form 
of  marriage,  which  is  still  to  be  met  with  in  certain 
parts  of  Bulgaria,^  are  exhibited  in  the  nuptial  rites 
and  customs  of  Ukraine,  where  the  ceremonial  is 
quite  a  museum  of  reduced  institutions. 

Vestiges  of  what  originally  was  marriage  by 
capture  occur  too  in  France  and  Belgium,  and  other 
countries.  The  nuptial  games  of  Lower  Brittany 
and  Chimay  have  obviously  been  derived  from  it.^ 

Marriage  hy  purchase. — The  retrogressive  evolu- 
tion of  this  form  of  marriage  exhibits  the  following 
stages  : 

^  Th.  Volvok,  Rites  et  usuges  nuptiaitx  de  V  Ukraine  [VAnthro- 
pologie,  1891,  p.  169).  "  In  certain  parts  of  Bulgaria  (Kustendil), 
the  capture  of  young  girls  takes  place  even  in  the  present  day  and 
constitutes  a  form  of  marriage  {Vlatcheny  monny)." 

2  Monseur,  Bulletin  du  Folklore^  January  and  March  1895, 
Coutumes,  p.  1  ;  les  noces,  p.  18. 

"  In  some  villages  of  Chimay  {Hainault)^  when  a  young  man 
chooses  a  bride  from  a  neighbouring  village,  the  young  people 
proceed  in  cavalcade  to  the  home  of  the  bride.  The  leader  of 
them  presents  her  with  a  whip  and  a  large  cake  crowned  with 
a  bouquet.  The  bride  then  takes  up  her  position  on  the  door- 
steps Avhile  the  bridegroom  and  his  friends  pass  and  repass  her 
at  full  gallop  seeking  to  dispossess  her  of  the  whip  with  which 
she  lashes  at  them.  All  this  may  be  regarded  as  the  last  remain- 
ing phase  of  marriage  by  capture,  the  bridegroom  arriving  at  the 
dwelling  of  the  bride  with  a  cortege,  being  more  or  less  figurative 
of  a  marauding  expedition." 

Baudrillard  {in  Stances  et  travaux  de  VAcaddmie  des  sciences 
morales,  Jan.  1884,  p.  36)  connects  the  customary  nuptial  games 
of  Lower  Brittany  with  the  primitive  marriage  by  capture.  In 
these  games  the  bride  hides  before  going  to  church,  and  the 
bridegroom  has  to  search  till  he  finds  her. 


SURVIVALS  EXIST  IN  ALL  KINDS  OF  SOCIETIES    163 

1.  The  payment    made   was   originally  a  pecu- 

niary compensation  to  the  family  of 
the  captured  bride.  By  degrees,  actual 
capture  gave  place  to  mere  symbolical 
capture  and  then  the  system  of  compensa- 
tion became  transformed  into  the  purchase 
system. 

2.  The  price. paid  for  the  bride  which  was  origin- 

ally the  property  of  the  whole  clan  under 
the  name  of  wergelt,  became  the  perquisite 
of  the  bride's  father. 

3.  It    next    became    modified   into    a    marriage 

dowry  given  to  the  wife  by  her  husband 
and  the  ceremony  of  purchase  became 
purely  symbolical.  In  the  Merovingian 
period,  for  instance,  the  future  husband 
presented  the  father  of  the  bride  with  a 
sou  and  a  denier  (marriage  per  solidum  et 
denarium)} 
Other  survivals  still  exist  of  these  three  stages 
in  the  evolution  of  marriage. 

(a)  In  Ukraine,  where  the  signification  of  the 
purchase  system  was  purely  one  of  the  vira 
or  compensation  (wergelt),  the  bridegroom 
has  to  give  a  present  to  each  member  of 
the  bride's  family ;  this  is  the  custom  too 
in  the  valleys  of  the  Caucasus  where  the 

^  Vanderkindere,  Condition  de  lafemme  d  1^4-poque  raerovvngienne, 
p.  12.  A.  Heussler,  Inslitidionen  des  deutschen  Privatreehts,n.,  p. 
280. 


164    UNIVEESALITY  OF  DEGENERATIVE  EVOLUTION 

relations  make  a  further  demand  of  pay- 
ment in  money.  ^ 
{b)  The  custom  of  purchasing  the  bride  from  the 
father  still  exists — although  with  no  legal 
recognition  among  the  Ossetes  and  in  certain 
Eussian  villages, 
(c)  Lastly,  the  marriage  per  solidum  et  denarium. 
Traces   of    this    are    left   even   in   modern 
France.     This  particular  purchase  signified 
the    payment    of    thirteen    silver    deniers 
according  to  a  certain  value  of  the  sou. 
The  introduction  of  these  thirteen  deniers  into 
the  ceremony  of  marriage  can  be  traced  through 
the  middle  ages  up  to  the  marriage  of  Louis  XVI. 
when  they  still  appeared.     In  some  parts  of  France 
such   as  Dijon,  Bordeaux   and   Barrois,  they   may 
still  be  met  with  even  nowadays.     With  the  excep- 
tion of  this  figurative  number  of  thirteen,  character- 
istic of  the  primitive  origin  of  the  ceremony,  this 
form  of  marriage  has  undergone  such  changes  as 
almost  to  entirely  obliterate  its  primitive  character. 
This  is  so,  too,  with  other  survivals  of  the  system 
of  marriage  by  purchase — they  have  come  to  be 
only    intelligible    by    means    of    the    comparative 
method,    or    by    a    knowledge    of    their    historical 
antecedents.^ 

^  Volvok,  Journal  V Anthropologic,  1892,  p.  579.  Kowalevsky, 
Droit  coutumier  ossitien,  p.  176. 

2  See  Paul  Viollet,  Histoire  du  droit  civil  frarypis  ;  Paris,  1893, 
p.  403. 


SURVIVALS  EXIST  IN  ALL  KINDS  OF  SOCIETIES     165 

Marriage  fairs — now  transformed  into  kermesses, 
and  still  held  at  Lierre  in  Belgium  and  other 
places — appear  to  have  been  originally  regular 
markets  for  the  purchase  of  young  women.^ 

The  custom  of  offering  the  wrong  young  woman 
to  the  bridegroom  as  his  future  wife — a  custom 
still  in  vogue  in  the  department  of  Landes  in 
France  —  formed  part  of  the  nuptial  ceremonies 
among  the  ancient  Hindoos,  and  is  probably  a 
vestige  of  the  tricks  which  were  played  upon  the 
bridegroom  after  the  purchase  of  the  bride.  Several 
vestiges  of  this  kind  may  be  noticed  among  the 
customs  of  modern  peasants.  It  is  by  no  means 
infrequent  for  a  wife  to  be  sold  by  her  husband, 
on  the  principle  that  what  has  been  bought  may 
fairly  be  sold,  and  the  transaction  rendered  legally 
binding  by  being  drawn  up  on  stamped  paper. 
This  is  merely  a  survival  of  the  old  system  of 
marriage  by  purchase. 

^  "The  second  Sunday  of  this  fair,  which  commences  on  the 
Sunday  after  All  Saint's  Day,  is  called  the  veersensmarkt  (Heifer- 
fair  or  market),  and  the  third  Sunday  is  called  the  hrullenmerkt, 
a  name  derived  from  the  word  hrul,  and  signifying  a  heifer  lowing 
noisily.  The  veersensmerkt  is  the  day  especially  set  apart  for  the 
young  girls  who  attend  the  fair  to  find  husbands.  The  hrullen- 
merkt is  the  day  dedicated  to  older  women  as  a  sort  of  forlorn  hope 
for  those  who  have  hitherto  failed  to  get  married.  No  one  seems 
to  know  when  these  "markets "  first  came  to  be  held.  I  am  more 
than  sixty  years  old,  and  they  were  old  when  I  was  young,  with 
this  difference  only,  that  in  my  youth,  a  young  girl  who  respected 
herself  would  not  have  been  seen  at  the  hrullenmerkt^  which  is  not 
the  case  nowadays."  (Taken  from  a  letter  from  the  Secretary  of 
the  Commune  at  Lierre.) 


16()    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

Marriage  hy  consent  of  both  parties. — In  French 
civil  marriages  of  the  present  day,  traces  remain  of 
the  two  other  forms  of  marriage  by  common  con- 
sent which  preceded  it. 

{a)  The  marriage  hy  simple  agreement,  which  held 
good  throughout  Christian  Europe  until  the  Council 
of  Trent,  is  still  valid  in  Scotland.  Gretna  Green 
marriages  (Gretna  Green  being  a  village  situated  on 
the  Border,  near  Carlisle)  were  notorious.  In  ac- 
cordance with  an  old  custom,  the  blacksmith  of 
Gretna  Green  kept  the  register  of  these  marriages, 
and  the  union  was  contracted  in  his  presence.^ 

After  the  Council  of  Trent,  marriages  in  facie 
Ecclesioi  came  to  be  alone  recognized  by  the 
Church :  all  marriage  to  be  valid  must  be  sanc- 
tioned (if  not  actually  celebrated)  by  the  parish 
priest  of  one  contracting  party,  in  the  presence  of 
one  or  more  witnesses.  The  legislation  of  the 
Council  of  Trent  became  French  law  after  the 
Ordinance  of  Blois  in  1574,  until  the  law  was 
revised  in  September  1792,  when  civil  marriage 
was  definitely  established  in  France. 

^  Viollet,  Histoire  du  droit  civil  fran^ais,  p.  428. 

"These  marriages,  about  which  everyone  has  heard,  were  not 
an  invention  of  Scotch  law.  Like  most  legal  curiosities,  they 
find  an  explanation  in  survival  from  a  former  condition  of 
things,  persisting  in  a  singularly  original  and  peculiar  form. 

"The  blacksmith  and  his  register  were  not  necessary  in  them- 
selves to  contract  a  marriage  ;  they  merely  constituted  evidence 
that  it  had  taken  place.  In  1804  a  Scotch  man  and  woman 
merely  declared  themselves  man  and  wife  in  writing,  and  in 
1811  they  were  recognized  as  legally  married  by  Scotch  law." 


SURVIVALS  EXIST  IN  ALL  KINDS  OF  SOCIETIES    167 

From  that  time  forth,  religious  marriage  may  be 
regarded  in  the  light  of  a  survival,  having  lost  all 
legal  importance,  while  civil  marriages  are  greatly 
on  the  increase.  Priests,  then,  the  "  groomsmen  " 
of  Lower  Brittany,  and  the  blacksmith  of  Gretna 
Green  may  alike  be  regarded  as  plain  evidences  of 
an  institution  in  process  of  decline. 

II.  The  family  system. — Archaic  forms  of  the 
family  still  exist,  sometimes  as  mere  vestiges, 
sometimes  as  exceptional  cases,  in  countries  where 
only  separate  families  are  legally  recognized. 

1.  The  Matriarchy. — Traces  of  the  matriarchy, 
i.e.  of  the  exogamous  family  of  blood  relations 
through  the  mother,  abound  among  the  customs 
of  the  inhabitants  of  the  valleys  of  the  Caucasus. 
Kowelevsky,  in  his  book  on  the  customs  of  the 
Ossetes,  has  dealt  with  this  subject.  The  vestiges 
remaining  even  in  France  of  marriage  by  capture 
and  the  prohibition  of  certain  marriages  in  Monte- 
negro, derived  doubtless  from  an  exogamous  period, 
are  alike  survivals  of  this  primitive  family  system. 
Further,  although  both  facts  and  their  interpreta- 
tion are  rather  doubtful,  some  authorities  regard 
the  couvade,  a  custom  which  is  still  in  practice 
among  the  Basques  and  also  in  the  Isle  of  Mark 
(in  Holland),  as  a  vestige  of  the  transition  period 
between  the  matriarchy  and  paternal  affiliation.^ 

^  See  Viollet,  Pricis  de  Vhistoire  clu  droit  francais,  ii.  326,  and 
Giraud-Teulon  in  Origines  du  marriage  et  de  la  famille  primitive, 
Paris,  1884,  p.  138  ;  and  Starcke,  Famille  primitive ,  p.  49. 


168    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

2.  The  Patriarchy. — The  Patriarchy,  which  was 
a  system  of  family  community,  is  still  exhibited  in 
the  farticvpanze  of  Italy,  the  companias  de  Galicia 
of  Spain,  the  parsonneries  of  France,  the  Haus- 
genossenschaften  of  Germany,  and  the  zadrugas  of 
the  Balkan  peninsular.  Besides  these,  the  family 
system  of  to-day,  according  to  Sumner-Maine, 
affords  constant  evidences  of  a  patria  protestas 
in  process  of  decay  and  of  declining  male  property 
rights. 

Survivals  from  the  old  patriarchal  system  abound 
in  modern  legislation.  Take  the  following  examples  : 

(a)  The  limitations  imposed  on  a  testator  with 

regard  to  leaving  his  property  away  from 
his  family  (Civ.  Code,  213  and  following). 

(b)  The  legal  opposition  to  a  woman's  equality 

in  succession. 
The  more  barbarous  laws,  if  not  actually 
excluding  women  from  succession,  at  least 
excluded  their  succession  to  landed  estate, 
in  order  to  keep  the  family  property  intact. 
In  France  the  privilege  of  sex  was  main- 
tained in  some  respects,  even  among  the 
peasants,  up  to  the  close  of  the  old 
regime,  and  it  still  exists  in  the  present 
day  in  different  degrees  in  Scandinavia, 
Russia,  Servia,  and  some  of  the  Swiss 
Cantons. 
c)  The  inequality  of  the  sexes  as  regards  con- 
jugal fidelity. 


SURVIVALS  EXIST  IN  ALL  KINDS  OF  SOCIETIES    169 

The  privileges  of  the  husband  over  the  wife 
in  this  matter  are  survivals  from  the  time 
when  these  obligations  were  wholly  on  one 
side. 
{d)  Affiliation  folloiving  on  a  double  marriage. — 
In  some  countries,  under  the  old  system,  a 
child  could  only  succeed  to  the  property  of 
his  parents  if  he  resided  with  them  (ex- 
cepting in  quite  exceptional  cases).     Hence 
there   arose   marriages   by   exchange.       In 
order  to  compensate  the  children,  the  two 
families,  if  each  had  a  son  and  daughter, 
exchanged  them,  and  bestowed  the  rights 
of  one  upon  the  other.     Under  the  new  law 
these  rights  could  not  be  legally  claimed, 
and  yet  even  in  the  present  century  they 
are  not  wholly  unknown.     The  last  mar- 
riage   by    exchange    was    probably    that 
mentioned  by  Dupin^  which  took  place  at 
Gacogne  (Ni^vre)  in  1889. 
In  conclusion  then,  we  have  found  that  while 
the   legislative   systems   of   modern   races   tend   to 
become  more  and  more  alike  in  main  principle,  we 
can  yet  find  vestiges,  more  or  less  faint  and  dis- 
torted,   but    quite    recognizable,    of    the    different 
institutions  which  dominated  the  earlier  conditions 
of  the  different  races. 

^  VioUet,  Histoire  du  droit  civil  frangais,  p.  491. 


PART  III 

SUMMARY    AND    CONCLUSIONS 

The  examples  we  have  been  able  to  give  in  the 
first  part  of  this  volume  make  it  plain  that  de- 
generative evolution  exists  everywhere.  It  must  be 
noticed,  however,  that  biological  investigation  shows 
that  in  the  evolution  of  organs  certain  parts  may 
disappear  completely,  but  also  that  in  the  evolution 
of  organisms  certain  organs  may  also  disappear. 
This  last  phenomenon  is  most  common  in  embryo- 
logical  development,  when  it  is  known  as  ontological 
abbreviation.^  Sometimes  it  is  the  adult  stage  that 
is  suppressed,  this  being  possible  by  what  is  called 
psedogenesis  a  precocious  appearance  and  ripening 
of  the  sexual  organs.^ 

^  In  Scalpellum  Stroini,  a  deep  sea  Cirripede,  the  nauplius  stage 
of  the  larval  life  is  suppressed  at  least  so  far  as  tliat  is  a  free 
swimming  larva.  Here  is  at  least  a  physiologically  complete  sup- 
pression of  a  whole  larval  stage. 

^  Axolotl.  Most  salamanders  pass  through  a  larval  stage  in 
which  they  are  aquatic  and  perform  their  respiration  by  means  of 
external  gills.  In  this  condition  they  are  incapable  of  reproduc- 
tion, and  must  undergo  metamorphosis  to  secure  propagation  of 
the  species.  In  the  case  of  Amblystoma,  however,  a  Mexican 
salamander,  the  larval  form  of  which  is  called  the  Axolotl,  repro- 
duction is  possible  in  the  larval  stage.  Thus  most  individuals  of 
170 


SUMMARY  AND  CONCLUSIONS  l7l 

Sometimes  a  degenerative  transformation  becomes 
still  more  complete  and  wonderful ;  not  only  may 
a  larval  stage  or  an  adult  stage  be  completely 
suppressed,  but  a  multicellular  organism  may  even 
lose  its  power  of  dying.  It  is  known  that  the 
simplest  forms  of  life  are  practically  immortal : 
when  a  microbe  like  micrococcics  divides  nothing 
dies,  and  throughout  the  whole  series  of  successive 
divisions  the  primitive  life  is  preserved.  On  the 
other  hand,  in  the  case  of  higher  animals  such  as 
man  there  are  both  mortal  somatic  cells  and 
reproductive  cells  which  by  means  of  conjugation 
become  practically  immortal.  The  mortality  of  the 
somatic  cells  is  evidently  an  acquisition,  an  advant- 
age fixed  by  natural  selection ;  but  there  exist 
multicellular  organisms  evidently  derived  from 
creatures  which  had  acquired  the  division  into 
mortal  somatic  and  immortal  reproductive  cells  and 
which  have  lost  it  since.  All  the  cells  of  their 
body  are  able  to  avoid  death  by  conjugation. 
This  occurs  in  many  conjugate  algae  like  spirogyra 

this  species  do  not  actually  reach  the  adult  stage.  According  to 
Boas  writing  on  Neotenie  in  Gegenbaur's  Festschrift,  1896,  this 
probably  happens  in  the  case  of  all  the  perennibranchiate  urodeles. 
Eanunculacece.  On  page  85  we  showed  that  in  Ranunculus 
aqvMLlis  there  are  produced  first  submerged  leaves,  and  afterwards 
floating  lobed  leaves,  and  that  the  flowers  are  produced  in  the 
axils  of  the  floating  leaves.  Some  forms  of  the  plant  living  in 
deep  water  produce  only  lacinated  leaves,  in  the  axils  of  which  by 
a  kind  of  psedogenesis  the  flowers  are  produced.  In  other  species 
{Ranunculus  fluitans  and  R.  divarkatus)  the  psedogenesis  has 
become  definitely  established  and  no  floating  leaves  are  formed. 


172    UNIVERSALITY  OF  DEGENERATIVE  EVOLUTION 

and  in  some  of  the  Volvocinere  (Stephanosphsera, 
Eudorina). 

Plainly  then,  the  further  one  examines  the  facts, 
the  more  enlarged  becomes  the  conception  of  de- 
generative evolution.  It  is  not  confined  to  unusual, 
abnormal  or  pathological  cases.  Degeneration  is 
not  an  accident  in  evolution :  it  is  the  obverse  of 
progressive  evolution  and  the  necessary  complement 
of  every  transformation  whether  anatomical  or 
social. 

Whatever  transformation  may  be  studied,  it  will 
be  found  that  change  is  always  accompanied  by  an 
elimination  of  some  parts  and  that  in  the  interests 
of  the  organism  as  a  whole  these  useless  parts 
gradually  degenerate.  When  a  whole  organization 
begins  to  undergo  retrogressive  evolution  and  to 
decay,  it  is  frequently  in  the  interests  of  some  still 
larger  organization.  Individuals  or  species  out  of 
harmony  with  their  surroundings  disappear  to  make 
room  for  others.  August  Comte  has  shown  how 
death  is  a  progressive  agency  in  the  social  organi- 
zation removing  the  worn-out  tissues  and  leaving 
room  for  new  and  more  plastic  intelligences.  All 
progress  implies  necessary  eliminations.  In  the 
domain  of  society,  those  who  are  victims  and  who 
from  birth,  education,  or  interests,  attach  themselves 
to  the  decaying  institutions  naturally  see  only  the 
degenerative  side  of  the  change ;  but  those  who 
regard  the  process  as  a  whole  and  do  not  concentrate 
their    attention    upon    the    injured    interests    and 


SUMMARY  AND  CONCLUSIONS  173 

individual  sufferings  will  see  the  other  side  of  the 
movement. 

When  a  social  organism  is  degenerating  there  is 
considerable  opposition  to  its  complete  disappearance, 
and  so  as  Houzeau  has  said  (see  the  summary  of 
Book  III.)  it  is  to  be  expected  that  living  and 
superior  civilizations  drag  behind  them  a  trail  of 
debris  from  dead  civilizations. 


BOOK  II 

THE  PATPT  OF  DEGENERATIVE 
EVOLUTION 

PART  I 

THE  SUPPOSED  LAW  THAT  DEGENERATION  RETRACES 
THE  STEPS  OF  PROGRESS 

It  is  a  common  opinion,  supported  partly  by  the 
etymology  of  the  word,  that  retrogression  is  a  tracing 
backwards  of  progression. 

"  In  the  degeneration  of  organizations  fallen  out 
of  use,'*  M.  A.  Lameere  has  said,  "  it  is  to  be  ob- 
served that  the  structures  formed  most  recently 
and  most  specialized  are  the  first  to  disappear,  and 
that  the  most  fundamental  characters  are  those 
which  persist  longest :  that  in  fact  degenerative 
evolution  retraces  the  steps  made  by  progressive 
evolution.  Peculiarities  recently  acquired,  if  they 
become  disused,  rapidly  disappear,  while  dispositions 
of  a  more  ancient  kind  have  a  persistence  almost 
exactly  proportioned  to  their  age.^ 

This  supposed  biological  law  of  retracement  has 

1  A.  Lameere,  Esquisse  de  la  Zoologie,  Bruxelles,  Rosez,  p.  184. 

175 


/ 


176        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

penetrated  to  psychology  and  sociology.  In  1868 
Hughlings  Jackson,  in  the  study  of  certain  maladies 
of  the  nervous  system,  had  arrived  at  the  conclusion 
that,  "In  the  degeneration  of  this  system  the  higher 
functions,  those  more  complex,  specialized  and  volun- 
tary, disappear  more  quickly  than  the  lower,  simpler, 
less  specialized  and  more  automatic  functions."  ^ 

Starting  from  this  point,  and  expressing  it  in 
terms  of  physiology,  Eibot  formulates  as  follows 
the  law  of  degeneration  of  will  and  memory : 
"  The  dissolution  of  the  will  occurs  in  a  retro- 
grade fashion,  from  the  more  voluntary  and  com- 
plex to  the  less  voluntary  and  simpler — that  is  to 
say,  towards  automatism."  ^ 

So  also  in  progressive  loss  of  memory,  the 
degeneration  proceeds  from  the  less  stable  to  the 
more  stable.  "  It  begins  with  recent  acquisitions 
not  firmly  rooted  in  the  brain,  rarely  repeated,  and 
so  not  firmly  associated  with  others,  in  fact  with 
the  least  organized  parts  of  memory.  It  ends 
with  sensory  memory  which  is  instinctive,  and  is 
deeply  rooted  in  the  organism,  or  is  indeed  a 
part  of  the  organism  itself."^ 

These  retrograde  transformations  of  the  nervous 
centres  have  their  echoes  in  the  modes  in  which 
ideas  and  feelings  are  expressed.     Paul  Heger,  in 

^  Ribot,  Maladies  de  la  Miinoire,  p.  29.     Dallemague  {D6gen4r4s 
et  D^4quilibr^)y  p.  430. 

^  Maladies  de  la  Volonti,  p.  150.     Paris,  F.  Alcan. 
^  Maladies  de  la  M^noire,  p.  94. 


DEGENERATION  RETRACES  STEPS  OF  PROGRESS  177 

particular,  has  shown  this  in  his  investigations  into 
the  degeneration  of  writing  and  speech.^ 

In  a  lecture  on  the  evolution  of  language,  de- 
livered at  the  University  of  Brussels,  he  said  as 
follows  :  "  For  several  years  I  have  studied  the 
degenerative  evolution  of  writing,  and  I  have 
shown  how  the  writing  of  the  insane  resembles 
that  of  children.  All  that  I  have  said  with 
regard  to  writing  applies  to  speech,  and  just  as 
drawing  lasts  longer  than  writing,  so  rhythm  and 
music  survive  after  the  power  of  expressing  ideas 
by  words  has  been  lost."  "  The  gradual  degenera- 
tion of  speech  may  be  traced  in  the  case  of  old 
men  who  gradually  pass  down  the  incline  into 
senility.  Study  of  the  speech  of  such  persons 
shows  that  the  degeneration  of  their  faculty  re- 
traces the  steps  by  which  it  had  been  progressively 
acquired." 

The  labours  of  Heger  were  in  a  field  where  the 
social  element  was  important.  It  is  a  small  step 
from  them  to  social  affairs  themselves.  A  number 
of  authors,  including  Ferrero,  Colajanni  and  Degreef, 
base  their  ideas  upon  this  law  of  retrogression,  which 
they  regard  as  established  and  applying  to  sociologi- 
cal phenomena.^ 

^  Sur  revolution  aggressive  de  Vilcriture  chez  certains  aliinis 
{Bull,  de  la  Soei6t6  d' Anthropologic  de  Bruxelles,  v.,  1885-1886). 
Sur  revolution  du  Langage  {Eevue  Universitaire,  Bruxelles,  1892- 
1893,  p.  143). 

2  Degi-eef,  Le  Transformisme  Social,  F.  Alcan,  Paris,  1895, 
p.  365. 

M 


^ 


1*78   THE  PATH  OF  DEGENERATIVE  EVOLUTION 

Before  describing  and  discussing  the  special  facts 
to  which  they  apply  their  theory,  it  is  necessary  to 
examine  its  biological  foundation. 


CHAPTEE  I 

THE    PATH    OF    DEGENERATION    IN    BIOLOGY 

We  have  now  to  consider  if  the  degeneration  of 
organs  retraces  the  steps  taken  in  their  progres- 
sive evolution.  According  to  Hughlings  Jackson 
and  Eibot,  in  the  cases  mentioned  in  the  preceding 
chapter,  degeneration  proceeds  by  successive  atro- 
phies occurring  in  the  order  opposite  to  that  of 
ontological  formation.  Is  the  same  order  to  be 
found  when  we  compare  the  degeneration  of 
organs  or  of  individuals  with  their  phylogenetic 
development  ?  To  answer  this  question,  we  must 
employ  both  morphology  and  embryology.  Using 
the  morphological  method,  we  shall  study  the  re- 
duction of  a  homologous  organ  in  several  species 
descended  from  the  same  type,  and  compare  the 
different  stages  of  reduction  with  the  different 
stages  of  phylogenetic  development  of  the  organ.^ 

^  In  this  investigation  it  will  be  necessary  to  compare  absolutely 
identical  organs — for  instance,  not  to  compare  the  pineal  eye  with 
the  paired  eyes.  It  will  be  necessary  also  to  choose  animals  of 
common  parentage — to  avoid,  for  instance,  comparing  a  Vertebrate 
eye  with  a  Crustacean  eye. 


THE  PATH  OF  DEGENEKATION  IN  ANIMAL>S       179 

Using  the  method  of  embryology  and  the  prin- 
ciple, so  far  as  it  can  be  followed,  that  ontogeny 
recapitulates  phylogeny,  we  shall  investigate  the 
mode  of  origin  of  some  reduced  structures.  We 
shall  thus  learn  if  the  organs  in  degenerating 
resume  any  of  their  ancestral  stages. 

Reduced  organs  may  appear  in  two  different 
forms.  They  may  be  atrophied  after  having 
reached  a  more  or  less  complete  development, 
and  in  this  case  we  shall  have  to  compare  the 
course  of  the  atrophy  with  the  course  of  the 
development.  They  may  be  rudimentary,  that 
is  to  say.,  their  development  may  have  been 
arrested  at  a  given  point,  the  adult  state  never 
being  attained.  In  this  case,  so  far  as  ontogeny 
repeats  phylogeny,  the  arrest  of  these  organs  at 
diflerent  stages  in  different  species  should  furnish 
a  series  with  greater  or  smaller  lacunae,  but  a 
series  which  will  be  the  reversed  series  of  the 
original  stages  in  phylogeny. 

Section  I. 
The  path  of  degeneration  in  animals. 

1.  Morphology  and  Embryology.  The  law  of  re- 
capitulation. — It  has  been  so  often  repeated  that  the 
individual  development  of  an  organ  is  a  resume  more 
or  less  compressed  of  its  historical  evolution,  that 
people  are  apt  to  attempt  too  exact  an  application 


180        THE  PATH  or  DEGENEKATIYE  EVOLUTION 

of  this  principle  to  every  individual  case.  Such  an 
exact  application  is,  however,  impossible.  Every 
living  organism  is  plastic,  and  in  its  development 
presents  individual  variations  which  serve  as 
material  for  the  operation  of  natural  selection. 
In  consequence,  the  recapitulation  cannot  be  more 
than  a  repetition  more  or  less  vague  of  the  essential 
phases  of  phylogeny.^ 

Moreover,  there  is  nothing  inevitable  in  the  law 
of  recapitulation,  for  most  plants  develop  directly. 

With  these  limitations,  however,  we  may  state 
that  among  animals,  the  ontogeny  usually  repeats 
in  a  modified  fashion  the  main  ancestral  stages. 
This  is  certainly  the  case  when  we  compare  the 
development  of  the  brain  of  man  with  the  probable 
ancestral  stages  as  displayed  in  the  series  of  verte- 
brates. 

^  Lang,  Anatomic  comparee.  As  throughout  the  whole  course  of 
time,  adaptation,  that  is  to  say,  the  preservation  of  wliat  is  most 
useful  in  the  struggle  for  existence,  is  a  force  modifying  heredity, 
it  is  plain  that  a  species  instead  of  resting  stable  must  change. 
According  to  its  circumstances,  moreover,  the  successive  stages  in 
the  ontogeny  of  a  creature  are  under  the  influence  of  conditions 
different  from  those  that  affected  the  corresponding  ancestral  stages. 
We  shall  call  the  process  of  embryology  palingenetic  so  far  as  it  is 
based  upon  inherited  legacies,  and  ccenogenetic  so  far  as  it  is  modi- 
fied by  adaptation. 

Baldwin  in  his  Treatise  on  Mental  development  in  the  child  and 
in  the  race  (London,  1895),  also  shows  that  the  development  of  the 
individual  is  not  an  exact  repetition  of  ancestral  stages.  The 
development  of  the  child  exhibits  "short  cuts"  and  phases  of 
direct  development  due  to  adaptation  and  destroying  the  exactness 
of  the  parallel  with  phylogeny. 


THE  PATH  OF  DEGENERATION  IN  ANIMALS       1  8 1 


In  Fish         The  cerebral  hemispheres 

and  Batrachia.  do  not  cover  the  region 

of    the    third    ventricle 

from  which  the  eyes  arise 

(thalamencephalon). 


In  the  human    Same 
embryo     (fig.    aspect. 
61,  A)  of  the 
seventh  week. 


In  Reptiles.  The  hemispheres  cover 
the  thalamenaphalon  Init 
leave  uncovered  the  region 
of  the  optic  lobes  (mesen- 
cephalon). 


In  the  human    Same 
embryo      (fig.    aspect. 
61,   b)  of  the 
middle  of  the 
third  month. 


In  mammals. 


The  hemispheres  cover 
the  thalamencephalon, 
the  mesencephalon,  some- 
times the  metencephalon 
(cerebellum  and  medulla), 
and  the  olfactory  lobes. 


In  the  human    Same 
embryo     (fig.    aspect. 
61,   c)  of  the 
fifth  month. 


In  some  mam-     The      hemispheres      are 

mals  even   of    smooth. 

higher  orders 

(c.  g.      some 

Hapalidse). 


In  the  human    Same 
embryo     (fig.    aspect. 
61,  d)  of  the 
midddle  of  the 
fifth  month. 


Within  such  limits,  the  law  of  recapitulation 
may  be  applied,  and  the  embryonic  history  of  an 
individual  may  be  considered  roundly  as  a  repeti- 
tion of  the  essential  phases  of  its  ancestral  history. 
We  have  now  to  consider  how  far  a  reduction  by 
atrophy  or  by  arrest  represents  a  retracing  of  steps 
in  evolution  (fig.  61). 

From   this   point    of   view,   we   may   study  the 


182        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

degeneration  of  the  pineal  eye  in  the  slow-worm, 
and  in  a  series  of  lizards. 

The  pineal  or  median  eye  in  the  slow  worm  and 


Fig.  61. — A,  Brain  of  a  human  embryo  of  seven  weeks  ;  h,  cerebi-al  hemispheres; 
ci,  intei-mediate  brain  or  thalairencpphalon;  cm,  raid-brain  ;  cp,  hind-brain. 
B,  Brain  of  a  human  embryo  about  the  beginning  of  the  third  month;  h,  cere- 
bral hemisplieres;  tg,  region  of  the  corpora  quadrigemina ;  cm,  mid-brain. 
(7,  Brain  of  a  human  embryo  at  tlie  middle  of  the  third  month;  h,  cerebral 
liemispheres  ;  (q,  coi'pora  quadrigemina;  ce,  cerebellum,  Z>,  Human  brain 
of  the  fifth  embryonic  month  ;  /i,  cerebral  hemispheres;  ol/,  olfactory  lobes ; 
S,  fissure  of  Sylvius;  ce,  cerebellum.  (After  Mihalkovics,  Entwickelungs- 
gcschichte  des  Gehirns.    Leipzig,  1877.) 

the  lizard  passes  through  the   following  stages  in 
its  individual  development.^ 

(1)  Formation  of  a  hollow  outgrowth  from  the 
roof  of  the  third  ventricle  of  the  brain 
(fig.  62,  D). 

^  P.  Francotte,  Recherches  sur  le  d4veloppement  de  V6pipliyse. 
{These  preseyiUe  d  la  Facidie  de  mMecine  de  Bruxelles.)  Arch,  de 
Biologie,  1888. 


THE  PATH  OF  DEGENEKATION  IN  ANIMALS   183 


(2)  This  little  sac  elongates,  changes  its  direction, 


B 


'C  r 


C/f 


Fig.  62.— Diagram  indicating  the  progressive  evolution  and  the  degeneration  of 
the  pineal  eye. 

A.  Perfect  pineal  eye,  as  found  in  the  slow-worm  before  birth,  or  in  the  adult 
Sphenodon  (Hatteria)  ;  c,  lens;  r,  retina ;  n,  optic  nerve ;  rf,  divertialum  of  the 
thalamencephalon.  B.  Pineal  eye  in  first  stage  of  degeneration  as  it  exists  in 
Chamceleo  and  as  it  was  in  the  slow-worm  before  stage  A.  The  lens  (c),  and 
the  retina  (r),  are  not  differentiated.  C.  Pineal  eye  In  the  degenerate  form 
found  in  Valotes  and  Leiodera;  c,  lens;  r,  retina;  n,  optic  nerve  in  fatty  de- 
generation. D.  Very  degenerate  pineal  eye  as  in  Cyclodus  and  lilie  the  earliest 
stage  in  the  slow-worm ;  there  is  no  differentiation  of  the  divertialum  from  the 
thalamencephalon.  E,  F,  G.  Other  modes  of  degeneration  of  the  pineal  eye. 
The  eye  lies  within  the  skull  and  there  is  no  parietal  foramen ;  cr,  cranial 
membranes;  E.  Ceratophora.  F.  Birds;  g,  mammals.  (After  Baldwin 
Spencer.) 

and  becomes  divided  into  a  proximal  and 
distal  portion.     The  cells  lining  the  distal 


184         THE  PATH  OF  DEGENERATIVE  EVOLUTION 

part,  that  farthest  from  the  brain,  become 

differentiated    into    the    cells    which    will 

form   the   lens,  and   the   cells   which   will 

form  the  retina. 
(8)  The  distal  part  becomes  specialized,  the  lens, 

the  retina,  and  the  stalk  of  the  optic  nerve 

are  mapped  out. 
(4)  The  lens,  the   retina,  and  the   optic   nerve 

become  fully  formed  (fig.  62,  a). 
At  this  stage  the  third  eye  has  reached  its  limit 
of  development. 

There  is  a  well-formed  retina  connected  with  the 
brain  by  a  special  optic  nerve.  The  organ  projects 
strongly  from  the  surface  of  the  head,  but  from 
this  point,  owing  to  the  development  of  the 
cerebral  hemispheres,  degeneration  begins.  The 
nerve  (fig.  52,  c),  becomes  broken  and  fatty,  and 
pigmentary  degeneration  occurs  in  it.  At  the 
same  time,  the  pineal  eye  having  become  useless 
or  even  harmful  to  the  animal  possessed  of  it, 
before  the  power  of  receiving  perceptions  of  light 
has  been  lost,  and  before  the  organ  has  been  far 
reduced  by  phylogenetic  destruction,  a  veil  of 
black  pigment  is  formed  over  it,  completely  shut- 
ting it  off  from  the  outer  light.  The  nerve 
disappears  completely  before  birth,  its  degenerate 
cells  becoming  lost  in  the  mesoblastic  skeletal 
tissue  of  that  region.  At  the  time  of  birth 
the  whole  eye  is  enclosed  in  a  thick  membrane 
which  isolates  it.       The  deposition  of  pigment  has 


THE  PATH  OF  DEGENERATION  IN  ANIMALS       185 

destroyed  any  functional  activity  in  the  lens  and  the 
retina,  but  these  parts  none  the  less  retain  traces 
of  a  complicated  structure  recalling  their  condition 
when  functional. 

In  the  Ehynchocephala  and  Lizards  examined 
by  W.  B.  Spencer,^  there  is  to  be  found  a  series  of 
types  representing  the  various  stages  of  the 
degeneration  of  the  eye  in  the  slow-worm. 

In  the  type  Sphenodon  {Hatteria,  fig.  62,  a),  the 
organ  in  the  adult  is  in  the  complete  form.  The 
eye  possesses  a  lens  (c),  a  retina,  (r),  with  com- 
plicated histological  structure.  A  nerve  {n)  places 
the  retina  in  communication  with  the  brain. 

In  Ghamceleo  (fig.  62,  b),  the  degradation  of  the 
organ  has  reached  the  following  stage  :  the  epi- 
physis has  a  distal  portion  corresponding  to  the 
eye,  but  the  histological  differentiation  of  this  is 
incomplete,  neither  the  retina  nor  the  lens  being 
distinct.  Nervous  fibres  connect  this  with  the 
proximal  portion  which  is  hollow  and  in  coni- 
munication  with  the  brain.  It  thus  represents 
the  second  stage  in  the  formation  of  the  eye  in 
the  slow-worm. 

In  the  types  Leiodera  and  Calotes  (fig.  62,  c),  the 
chief  degeneration  is  in  the  optic  nerve,  which  has 
partially  disappeared,  and  no  longer  connects  the 
eye  with  the  brain.      The  eye  itself  is  not  quite 

^  W.  B.  Spencer,  On  the  presence  and  structure  of  the  pineal 
eye  in  Lacertilia  (Quarterly  Journal  of  Microscopical  Science, 
1886). 


186        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

SO  degenerate  :  the  retina  has  not  a  complex 
structure,  but  both  retina  and  lens  are  present. 

In  the  type  Gyclodus  (fig.  62,  d),  the  degeneration 
is  still  greater :  the  epiphysis  is  a  vesicle  attached 
to  the  thalamencephalon.  The  walls  of  this 
vesicle  show  only  the  smallest  symptom  of  primi- 
tive differentiation  into  lens  and  retina.  The 
proximal  part  remains  hollow,  and  shows  no  trace 
of  differentiation  at  all. 

This,  however,  is  not  the  only  fashion  in  which 
degeneration  of  the  pineal  eye  proceeds.  In 
another  series  of  creatures  it  retains  its  connec- 
tion with  the  thalamencephalon,  but  remains  inside 
the  skull.  The  parietal  foramen  closes,  thus 
completely  shutting  off  the  eye  from  the  light; 
the  eye  becomes  useless,  degenerates,  and  the 
optic  nerve  loses  its  function  as  a  conducting 
channel,  Ceratophora  (fig.  62,  E).  The  pineal 
organ  then  becomes  a  degenerate  structure  in 
which  it  is  exceedingly  difficult  to  see  traces  of 
its  original  condition,  and  which  is  usually  marked 
by  an  abundance  of  blood-vessels  :  Birds  (fig.  62,  F), 
Mammals  (fig.  62,  g). 

Thus,  the  degeneration  of  the  pineal  eye  shows 
that  the  optic  nerve,  the  last  organ  to  be  completed, 
is  the  first  to  disappear.^ 

In  studying  the  degeneration  of  the  pineal  eye, 

1  The  same  happens  in  the  case  of  the  ordinary  paired  eyes.  In 
the  degenerate  eyes  of  the  mole,  the  optic  nerve  is  more  reduced 
than  are  the  other  structures. 


THE  PATH  OF  DEGENERATION  IN  ANIMALS       187 

we  have  Seen  the  path  of  progressive  evolution 
being  retraced,  at  least  in  the  case  of  these  reptiles. 
The  present  state  of  knowledge  does  not  permit  an 
extension  of  the  enquiry  to  the  conditions  found  in 
higher  animals.  However,  according  to  Flesch, 
traces  of  a  sensory  epithelium  have  been  found  in 
the  pineal  body  of  man,  the  horse,  the  sheep,  and 
the  bat :  that  is  to  say,  that  in  these  creatures  too 
the  oldest  parts  of  the  structure  have  resisted 
degeneration  longest.  We  cannot  refrain  from  the 
conclusion  that  in  this  series  degeneration  retraces 
to  a  large  extent  the  steps  of  original  advance. 

2.  Degeneration  of  the  organs  of  sight  in  deep- 
sea  Crustacea. — We  cannot  however  establish  the 
conclusion  of  the  last  paragraph  as  a  general 
principle. 

The  fauna  of  the  deep  sea  includes  a  large 
number  of  Crustacea,  and  in  these  the  eyes,  which 
are  relatively  useless,  are  often  degenerate.  The 
course  of  the  degeneration  is  generally  definite,  and 
of  all  the  structural  parts  the  most  long-lived  are 
the  eye-stalks,  although  we  know  that  these  are 
a  recent  formation.  A  number  of  examples  chosen 
from  Decapod  Crustaceans,  which  are  specially 
abundant,  will  illustrate  this  point.^ 

Nephropsis,  which  lives  in  the  Atlantic  and 
Indian  Oceans  at  moderate  depths,  is  a  relative  of 
the  Lobster.     The  optic  stalk  is  short  and  carries 

*  See  Pelseneer,    VExploration   zoologique   des    mers   profmides 
(Conferences  Universitaire  de  Bruxelles).     2  Annee,  1890. 


188    THE  PATH  OF  DEGENERATIVE  EVOLUTION 


a  rudimentary  eye  which  has  neither  pigment  nor 
cornea  and  is  coloured  like  the  general  surface  of 
the  body. 

Eryonicus  (fig.  63)  belongs  to  the  same  group, 
and  comes  from  the  region  of  Saint-Thomas  in  the 
Antilles,  where  it  lives  at  a  depth  of  about  825 
yards.  This  animal  has  a  reduced  optic  stalk, 
but  at  the  extremity  of  this,  where  in  littoral 
J.  forms   the  eye 

Y~     ^^-^A  is  borne,  there 

"~^  is    only  a   de- 

pression as  if 
the  eye  had 
been  carefully 
scooped  out. 

Willemoesia 
(fig.  64),  a  re- 
lative of  the 
marine      cray- 

FiG.  63.    Evyonicus  coecus.    Bate?    (After  W.  Faxon,    -fioVi    qrifl    qti   in- 
The  Stalk-eyed  Crustacea,  Mem.  of  Mus.  of  Comp., 
Zool.  Harvard  College,  vol.  xviii.,  1895.)  habitant  of  the 

Atlantic  at  a  depth  of  about  3500  yards,  is  com- 
pletely devoid  of  eyes  in  the  adult  condition, 
although  it  possesses  them  in  the  larval  stage. 

Scolophthalmus  (fig.  65),  which  lives  down  to 
4000  yards,  is  quite  devoid  of  eyes,  but  possesses 
eye-stalks  which  terminate  in  spines. 

It  seems,  then,  that  different  species  of  deep-sea 
Crustacea  may  present  different  degrees  of  degenet*a- 
tion  of  the  eye.      One  species  in  itself  exhibits  all 


Fig.  Gi.  —  WUlemcesia.    (After  Pelseneer.) 


190        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

grades  of  degradation  according  to  the  depth  at 
which  it  lives.  This  creature — Cymonomus — 
which,  when  near  the  surface,  has  fully  formed 
eyes  upon  movable  stalks,  at  a  depth  of  a  few 
hundred  yards  exhibits  movable  stalks  without 
eyes;  and  at  1500  yards  the  stalks  are  fixed  and 
end  in  spines. 

Isopod  Crustacea,   which  live  in  the  deep    sea, 
present  similarly  degenerate  eyes.     Many  are  blind 


Fig.  &i).—Scolophthalmus  luci/ngus,  Fax. 

a,  optic  peduncle  li-ansformed  to  a  spine.      (After  W.  Faxon,  TTie  Stalk-eyed 

Crustacea,  Mem.  of  Mus.  of  Comp.  Zool.  Harvard  College,  vol.  xviii.,  1895.) 

and  display  all  kinds  of  optic  degeneration.  Nocsa, 
for  instance,  simply  has  eyes  devoid  of  pigment. 
Thus,  in  abysmal  Crustacea,  the  degeneration  of  the 
eyes  is  in  no  sense  a  retracing  of  developmental 
stages. 

Another  instance  chosen  from  examples  of  the 
atrophy  of  organs  in  individuals,  shows  that  the 
supposed  law  of  retracing  cannot  be  made  uni- 
versal. 

4.  Atrophy  of  the  hranchial  vessels  in  man. — 
Examination  of   a  human   embryo  of  about  three 


THE  PATH  OF  DEGENEBATION  IN  ANIMALS       191 


if 


C 

G 
C 

C 


weeks  old  shows  the  presence  of  a  series  of  slits  on 
the  sides  of  the  neck,  the  slits  not  being  parallel, 
but  converging  towards  the  ventral  surface.  Be- 
tween these  slits  are  swellings,  or  pads,  which-  pass 
up  towards  the  dorsal  surface  and  appear  like  the 
beginnings  of  hoops  or  ribs  enclosing  the  visceral 
cavity ;  the  elevations  are  the  branchial  arches,  the 
slits  are  the  gill-slits. 

In  the  human  embryo  (fig.  66,  A),  as  in  fish, 
these  slits  appear  « 

from  above 
downwards,  and 
as  they  are 
formed,  the  cor- 
responding blood- 
vessels arise. 

These  vessels, 
or  aortic  arches, 
arise  from  a  ven- 
tral aorta  (a.) 
which  gives  off 
six  lateral 
branches  (c.)  at  each  side.  These  lateral  branches 
run  up  between  the  gill- slits  and  form  two  main 
trunks  on  the  dorsal  side  which  converge  to  form 
the  descending  aorta  (ad.). 

In  man  the  branchial  arches  are  transformed, 
parts  of  them  entering  into  the  structure  of  the 
face,  and  during  the  transformation  parts  of  the 
aortic  arches  atrophy  (fig.   66,  b).     But  the  order 


Fig.  66. — Diagram  of  branchial  arches  in  mammals. 

A.  Embryonic  stage,  a,  aorta;  c,  aortic  arches; 
AD,  dorsal  aorta.  B.  Adult  stage.  The  parts 
represented  by  dotted  lines  have  degenerated. 
A,  aorta;  v,  carotid;  ap,  pulmonary  artery. 


192        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

of  this  atrophy  does  not  correspond  in  any  way  to 
the  order  of  the  formation  of  the  vessels. 

The  median  parts  of  the  anterior  two  lateral 
branches  (w}  and  m^)  disappear,  and  the  vertical 
parts  remain  as  the  internal  and  external  carotid 
vessels.  The  vertical  piece  which  joined  the 
posterior  parts  of  the  third  and  fourth  arches  dis- 
appears :  the  internal  and  external  carotids  thus 
acquire  a  stem  of  their  own.  The  parts  of  the 
fourth  arch  remain ;  the  fifth  arch  disappears  at 
each  side,  and  the  sixth  arch  forms  the  pulmonary 
artery  (ap.). 

Thus  the  degeneration  of  these  vessels  represents 
in  no  way  whatever  a  retracing  of  their  develop- 
mental history.  AH  that  occurs  is  that  the  useless 
parts  disappear  and  the  useful  parts  persist.  A 
comparative  study  of  this  example  would  only 
enforce  our  conclusion. 

In  ontogeny  the  neurapophyses  are  more  ancient 
than  the  vertebral  centres.  None  the  less,  as  we 
have  already  seen,  the  examination  of  any  vertebral 
column  from  head  toward  tail  shows  a  gradual 
disappearance  of  all  parts  except  the  centra, 
although  the  centra  are  the  last  to  be  formed. 

Section  II. 

The  path  of  degeneration  in  plants. 

].  Rarity  of  cases  of  recapitulation  in  the 
organogeny  of  leaves. — We  have  already  said  that 


THE  PATH  OF  DEGENERATION  IN  PLANTS         193 

recapitulation  seldom  occurs  in  plants,  the  develop- 
ment of  the  whole  and  of  its  organs  being  usually 
direct.  When  it  does  occur,  it  is  generally  limited 
to  characters  coming  from  comparatively  recent 
ancestors  and  not  even  in  the  most  transitory 
form  entering  into  the  formation  of  the  fundamental 
parts  of  the  plant.^ 

^  The  rarity  of  recapitulation  among  vegetables  is  the  result 
partly  of  their  fixed  condition  in  the  soil,  and  partly  of  the  more 
rigid  nature  of  their  cells. 

The  immobility  of  a  plant  forces  the  adult  to  live  in  the  same 
place  as  the  embryo.  Among  animals,  on  the  other  hand,  it 
frequently  happens  that  the  young  pursue  a  manner  of  life 
different  from  that  of  the  adult  and  resembling  that  of  the 
ancestor.  Young  Cirrepedes  are  vagrant  and  have  the  same 
needs  and  use  the  same  organs  as  other  vagrant  Crustacea  ;  larval 
frogs  inhabit  the  water  like  their  fish-like  ancestors.  In  plants 
there  is  nothing  similar  ;  all  the  aquatic  flowering  plants  are 
derived  from  terrestrial  ancestors,  but  if  at  the  beginning  of  their 
existence  these  aquatic  plants  were  to  bear  leaves  adapted  to 
aerial  life  they  would  ensure  their  own  destruction.  The  ex- 
ceedingly rare  ancestral  traits  to  be  found  in  'a  few  species  are 
naturally  of  a  kind  not  to  incommode  their  possessors.  It  is 
improbable  that  these  are  a  legacy  from  distant  ancestors  ;  they 
would  not  have  been  spared  by  natural  selection  had  they  not 
come  from  ancestors  of  very  much  the  same  habit.  The  absence 
of  locomotion  in  plants  has  also  produced  a  greater  adaptability 
than  among  animals.  Animals,  \vhen  conditions  are  unfavourable 
can  remove  in  search  of  more  suitable  localities,  plants  being  fixed 
in  the  soil  must  become  modified  or  perish.  Plants,  therefore, 
ofler  numerous  cases  of  individual  adaptation.  We  do  not  know 
if  these  adaptations  are  transmitted  by  heredity,  but  natural 
selection  has  at  least  secured  the  widest  range  of  plasticity. 
Thus  plants  rapidly  rid  themselves  of  ancestral  legacies  which 
have  become  useless. 

The  transitory  organs  of  animals  arc  employed  for  the  service 
N 


/ 


194   THE  PATH  OF  DEGENERATIVE  EVOLUTION 

In  consequence,  vegetable  embryology  is  of  little 
use  for  investigation  of  the  supposed  backward  path 
of  degeneration,  for  the  rudimentary  or  reduced 
organs  of  plants  do  not  generally  represent  ancestral 
stages. 

The  seedling  of  Lathyrus  tenuifolius  (fig.  67),  a 
vetch,  possesses  rudimentary  organs  which  cannot 
be  ancestral  stages  as  their  development  is  direct. 
In  this  plant  a  whole  series  of  leaves  are  formed 
between  those  arising  at  germination  and  the  adult 
leaves.  This  intermediate  series  displays  many 
arrests  of  development. 

The  adult  leaf  has  a  pair  of  stipules,  foliage 
leaflets,  and  tendrils  (fig.  67,  j).  The  leaves  just 
before  these,  have  a  pair  of  stipules  (fig.  67,  i), 
which  are  absent  in  the  leaves  next  before  (fig.  67, 
h).  Still  earlier  leaves  are  produced  with  fewer 
leaflets  and  tendrils  (fig.  67,  d-g),  leaves  without 
leaflets  and  with  a  single  tendril  (fig.  67,  c),  and 
leaves  entirely  without  tendrils  (fig.  67,  b).  Lastly, 
at  germination  very  rudimentary  leaves  are  pro- 
of the  whole  body,  the  branchial  arches  of  mammals  are  employed 
in  the  formation  of  important  parts  of  the  head  and  neck.  The 
tail  of  the  tadpole  is  reabsorbed  by  phagocytes  and  its  substance 
used  for  the  nutrition  of  the  body.  In  the  case  of  plants,  such 
occurrences  are  rare  and  limited  ;  the  cells  are  enclosed  in  a  rigid 
wall  which  resists  displacement  or  alteration  ;  the  protoplasmic 
contents  may  be  absorbed  and  used  as  nutritive  material  by 
another  part,  but  the  cellulose  cell-wall  remains.  A  useless  organ 
can  be  eliminated  only  at  the  expense  of  loss  of  material. 
J.  Massart,  La  Recapitulation  et  V Innovation  en  emhryologie 
v6gdtale  {Bull.  Soc.  Boy.  Bot.  Belg.,  t.  xxxiii.,  p.  150,  1894). 


Fig.  67.— Lathyrus  tenuifolius. 
A,  B,VeedUng8  in  two  stages  of  growth.    C  to  J,  diflferent  forms  of  successive  leaves. 


196        THE  PATH  OF  DEGENERATIVE  EVOLUTION 


duced,  the  sole  function  of  which  is  to  protect 
the  axillary  bud  (fig.  67,  B,  1,  2).  We  have  now 
to  show  that  this  series  is  by  no  means  a  retracing 

of  ancestral  stages. 
All,  or  nearly  all, 
the  Papilionace- 
ous plants  have 
stipulate  leaves ; 
this  shows  that 
the  exstipulate 
leaves  of  L.  tenui- 
folius  do  not  re- 
present an  ances- 
tral stage.  More- 
over, before  the 
acquisition  of 
tendrils,  vetches 
had  a  terminal 
leaflet  (see  Vicia 
Pyrenaica,  fig. 
44) ;  none  of  the 
reduced  leaves  in 
L.  tenuifolius  re- 
produce       this 

Fig.  08. — SeeilmgVicia  of  TDonanthos.  Sta<^e  '       mOrCOVCr 

the  simple  leaves  without  leaflets  do  not  represent 
an  ancestral  condition ;  the  winged  petiole  is  not  an 
ancestral  character. 

The     primary    leaves    of     another    vetch     Vicia 
monanthos   confirm   our    conclusion.      In   this   case 


THE  PATH  OF  DEGENERATION  IN  PLANTS         197 

the  first  few  leaves  formed  in  the  seedling  remain 
rudimentary,  aud  serve  only  to  protect  the  axillary 
buds  (iig.  68,  leaves  1  and  2).  Contrasted  with 
the  condition  in  L.  tenuifolius,  although  all  the 
vetches  had  probably  a  common  ancestor,  these 
primary  leaves  have  three  little  projections,  the 
two  lateral  of  which  are  reduced  stipules. 

We  do  not  know  of  any  vegetable  example  of 
recapitulation  in  the  case  of  an  organ  reduced  by 
arrest  of  development.  Plants  exist,  however, 
which  after  having  produced  leaves  of  typical 
structure  begin  to  produce  leaves  the  development 
of  which  remains  arrested.  Such  a  plant  is  the 
Acacia  which  bears  phyllodes.  At  first  the  leaves 
are  like  those  of  other  Acacias ;  next  it  bears  leaves, 
the  blades  of  which  are  rudimentary ;  and  finally 
leaves  with  normal  stipules,  but  with  no  trace  of 
lateral  leaflets  on  the  petiole. 

In  this  case  the  ancestral  conditions  are  known 
and  are  quite  different.^ 

2.  Organogeny  of  flowers. — What  we  have  said 
about  leaves  applies  to  the  organogeny  of  flowers. 
Here  also  in  the  cases  of  atrophy  produced  by 
arrest    of    development   there   is   no   indication   of 

^  Many  other  reduced  plant  organs  might  be  instanced,  such  as 
the  stipules  of  Sambucus  or  the  teeth  of  the  calyx,  in  many  Com- 
positse  and  Umbelliferse.  After  their  formation  such  organs  grow 
very  slowly  and  exhibit  no  trace  of  recapitulation.  This  happens 
with  the  leaves  of  Sempervivum  (fig.  49).  Without  doubt  these 
leaves  are  derived  from  leaves  normally  divided  into  hypopodium 
and  epipodium,  but  they  show  no  trace  of  this  division. 


198    THE  PATH  OF  DEGENERATIVE  EVOLUTION 


ancestral  stages,  and  thus  there  is  no  evidence  that 
degeneration  retraces  the  path  of  progressive  evolu- 
tion. 

In  the  cauliflower  (Brassica  oleracea,  var.  Botry- 

tis),  a  cultivated 
A         ^^N  _  variety,   the 

inflorescence 
branches  ex- 
uberantly; most 
of  the  flowers 
produced  on 
these  branches 
are  arrested  in 
their  develop- 
ment. Of  the 
immense  num- 
ber of  flowers 
produced  on 
each  plant  only 
a  few  attain 
sexual  maturity 
and  produce 
seeds;  the  others 
abort   at    differ- 


FiG.  69. — Development  of  flowers  of  Ihe  Cauliflower. 

sa,  anterior  sepal;  sp,  posterior  sepal;  si,  lateral 
sepal ;  p,  petal;  e,  long  stamen ;  ec,  short  stamen  ; 
o,  ovary.  A,  very  young  flower  with  only  rudi- 
ments of  two  sepals;  B,  flower  with  rudiments  of 
four  sepals  ;  C,  older  flower  with  rudiments  of 
petals;  D,  still  older  flower  with  rudiments  of 
stamens  and  ovary.  (The  flowers  A— D  are  seen 
from  the  f'ide.  In'  the  figures  C  and  D  the  lateral 
sepal  has  been  removed.)  E,  flower  seen  from 
behind;  the  posterior  sepal  has  been  removed. 


ent  stages. 


stage 


Most  of  these  remain  in  a  very  primitive 
and  do  not  develop  sepals ;  but,  scattered 
among  them,  may  be  found  more  fully  developed 
flowers,  so  that  the  same  plant  presents  almost 
every  possible  stage  of  flower  development. 

The  orgaaogeny  of  these  flowers  is   quite  like 


THE  PATH  OF  DEGENERATION  IN  PLANTS         199 

that  of  other  Cruciferse.  If  in  the  cauliflower  the 
arrested  flowers  corresponded  to  different  ancestral 
stages  the  case  would  be  striking ;  but  this  does 
not  occur.  Primitively  the  flower  contained  only 
the  stamens  and  pistils,  the  essential  organs  of 
reproduction.  But  we  see  that  the  cauliflower 
produces  first  the  anterior  and  posterior  sepals 
(fig.  69,  a),  then  the  lateral  sepals  (fig.  69,  B),  then 
successively  the  petals  (fig.  69,  c),  the  four  larger 
stamens  (fig.  69,  d),  and  the  two  shorter  stamens 
(fig.  69,  e).  Moreover,  the  flowers  display  a  special 
readiness  to  the  suppression  of  certain  parts  such 
as  the  petals,  as  in  the  flower  d  (fig.  69). 

3.  Progressive  degeneratio7i  of  the  prothallus  in 
phanerogams. — Although  embryology  gives  us  few 
examples,  morphology  proves  clearly  enough  that 
in  plant  degeneration  there  is  no  return  to  ancestral 
types. 

This  appears  clearly  from  a  comparison  of  the 
progressive  evolution  of  the  prothallus  in  crypto- 
gams with  its  degeneration  in  phanerogams. 

Terrestrial  vegetation  has  been  derived  entirely 
from  aquatic  life.  The  Bryophyta  (mosses  and 
liver-worts),  the  Vascular  Cryptogams  (ferns),  and 
the  flowering  plants  all  have  sprung  from  aquatic 
algae  probably  not  very  different  from  Coleochcete. 
Such  aquatic  forms  are  reproduced  by  means  of 
true  ova  and  spermatozoa.  The  terrestrial  plants 
which  were  derived  from  them  had  also  spermatozoa 
with   vibratile   locomotor    hairs    and  impregnation 


200        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

took  place  in  a  fluid.  A  special  organ,  the  arche- 
gonium,  was  developed ;  this  contained  the  ovum 
and  made  the  approach  of  the  spermatozoa  more 
easy. 

For  the  present  purpose  we  may  omit  considera- 
tion of  the  mosses  and  liver-worts  as  it  is  improb- 
able that  they  are  in  the  line  of  ancestry  of  the 
flowering  plants.  It  is  necessary  only  to  say  that 
in  them,  while  a  single  egg  is  produced  in  the 
archegonium  a  large  number  of  spermatozoa  are 
produced  in  the  antheridium. 

The  same  condition  is  found  among  the  fern-like 
plants,  but  in  their  case,  owing  to  the  development 
of  special  channels  for  the  passage  of  nutritive 
materials,  it  is  possible  for  a  much  greater  size  to 
be  reached.  In  these  circumstances  it  would  be 
unlikely  that  the  spermatozoa  should  find  at  the 
summit  of  a  comparatively  lofty  plant  the  drop 
of  water  necessary  for  the  task  of  fertilization.-^ 
Accordingly  the  sexual  cells  are  produced  on 
prothalli,  which  hardly  reach  above  the  surface  of 
the  ground,  and  are  in  a  favourable  position  for  the 
necessary  moisture. 

In  the  less  specialized  ferns  (fig.  70)  a  single 
prothallus  bears  both  male  and  female  organs 
(antheridia    and    archegonia).     In  the  Equisetums 

^  The  smaller  forms  like  the  Selaginellidse  and  the  club-mosses 
were  represented  in  the  past  by  plants  of  much  greater  size,  as  is 
seen  from  fossil  remains,  and  it  is  probable  that  the  modern  forms 
have  descended  from  these  giants. 


THE  PATH  OF  DEGENERATION  IN  PLANTS 


201 


the  prothalli  are  unisexual,  but  the  spores  from 
which  they  are  produced  are  alike.  Finally,  in  the 
allies  of  Selaginella  which  are  the  Cryptogams  most 
nearly  approaching  flowering  plants,  the  prothalli 
which  bear  male  organs 
are  quite  distinct  from 
the  prothalli  which  bear 
female  organs  ;  more- 
over, the  spores  which 
give  rise  to  the  two 
kinds  of  prothalli  are 
quite  different.  A 
small  number  of  large 
spores  arise  in  special 
sporangia,  termed 
macrosporangia,  and  it 
is  from  these  that  the 
female  prothalli  arise. 
A    larger    number    of 

smaller  spores    are  pro-      p,Q    70.— Seedllng  of  Fern  {Asplmtum 

duced       in       micrOSpOr-    The  prothallusO>Tl)Siis"antheridiaat  its 
prima     onrl   tliPCP   miPrn  edges  and  towards  the  centre  an  arche- 

angia    ana  tnese  micro-        gonium  which  has  been  fertUized.    The 

Qnnrpcinrnrlnpp  tliP  nmlp       5"^""^    seedling    developed    from    the 
spuitJb  piUUUOe  me  Illdlt;        fertilized  ovum   has  already  given  rise 

■nrnt'li  a  111  ^^  ^^'^'''^  ^^^  **^  ^"^^  leaves. 

The  spores  of  ferns  are  very  small  and  contain 
only  the  nutritive  matter  required  for  germination ; 
the  spores  of  Selaginella,  even  the  microspores,  are 
very  much  larger  and  contain  a  large  supply  of 
reserve  material  at  the  expense  of  which  the 
development  of  the  prothallus  takes  place. 


202        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

We  may  give  a  brief  account  of  the  formation  of 
the  sexual  cells  in  Selaginella.  The  first  stages  in 
the  appearance  of  microsporangia  and  macro- 
sporangia  are  identical.  Later  on  in  the  one  case 
each  of  the  spore  mother-cells  breaks  up  into  four 
microspores,  in  the  other  case  only  one  mother-cell 
breaks  up  into  four  macrospores,  the  others  dis- 
appear. On  germination  the  macrospore  becomes  a 
fairly  large  prothallus  with  several  archegonia :  the 
microspore  forms  only  one  cell  as  the  representative 
of  the  prothallus,  the  rest  of  the  structure  becoming 
the  antheridium. 

In  Phanerogams  (flowering  plants)  a  drop  of 
water  is  no  longer  necessary  for  fertilization.  The 
spermatozoon  reaches  the  ovum,  not  by  swimming 
movements,  but  by  a  kind  of  growth.  This 
difference  has  resulted  in  profound  modifications. 
Fertilization  may  occur  in  the  air,  and  it  is  no 
longer  necessary  for  the  macrospore  to  become 
detached  :  it  remains  fixed  on  the  individual  from 
which  it  arose  and  there  proceeds  to  develop. 
In  the  Gymnosperms  the  archegonia  are  fully 
formed  in  a  prothallus  of  considerable  size,  but  the 
Angiosperms  which  are  more  specialized,  retain 
only  traces  of  these  structures.  The  microspores 
(pollen  grains)  exactly  as  in  Selaginella  are  pro- 
duced in  great  numbers,  but  when  they  leave  the 
parent  plant  they  develop  only  if  they  reach  suit- 
able conditions.  In  germination  there  is  shut  off, 
as  in  Selaginella,  a   single   cell    to    represent  the 


THE  PATH  OF  DEGENERATION  IN  PLANTS         203 

prothallns ;  later  on  in  a  few  Gymnosperms  it 
undergoes  a  few  divisions  representing  the  forma- 
tion of  the  antheridium.  In  Angiosperms  the 
reduction  has  proceeded  further,  and  each  pollen 
grain  besides  the  prothallus  cell  produces  only  one 
or  at  most  two  reproductive  nuclei.  In  all 
Phanerogams  the  reproductive  nucleus  reaches  the 
ovum  by  being  carried  in  a  long  tube  which  grows 
out  from  the  pollen  grain  at  the  expense  of 
nutriment  derived  from  the  female  tissues. 

Thus,  the  prothallus  may  be  traced  through 
marked  stages  of  reduction  from  the  condition  in 
ferns,  through  Selaginella  to  Gymnosperms  and 
higher  flowering  plants.  But  these  phases  of 
reduction  are  by  no  means  to  be  regarded  as 
repetitions  of  ancestral  conditions. 

In  conclusion,  it  is  plain  that  we  cannot  assert 
as  a  general  law,  that  degeneration  retraces  the 
steps  of  evolution.  In  the  vegetable  kingdom  no 
facts  support  such  a  conclusion.  In  certain  special 
cases  in  the  animal  kingdom,  the  most  recently 
developed  structures  are  the  first  to  disappear  when 
atrophy  sets  in,  but  this  proves  no  more  than  that 
these  particular  structures  happened  to  be  less 
stable,  and  to  offer  less  resistance.  It  is  quite 
impossible  to  make  such  facts  support  the  generally 
held  view,  that  degeneration  is  a  kind  of  inverse  of 
evolution. 

It  is  very  seldom  that  a  living  apparatus  with 
complex   functions  loses  all  of  them,   but  usually 


204        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

preserves  one  or  other ;  for  instance,  the  leaves  of 
parasitic  flowering  plants  continue  to  serve  as 
protectors  of  the  buds,  and  degeneration  does  not 
affect  the  part  that  has  remained  functional.  It 
is  highly  probable  that  the  original  function  of 
leaves  was  assimilative  rather  than  protective,  and 
yet  here  it  is  the  later  function  that  is  retained. 

Even  when  all  function  is  lost,  and  the  whole 
structure  degenerates,  there  is  no  reason  why  the 
degeneration  should  retrace  the  evolution.  In  the 
case  of  the  atrophy  of  an  organ  in  an  individual 
especially  in  such  pathological  instances  as  those 
mentioned  by  Ribot,  it  may  be  that  the  latest 
formed  parts  are  the  most  fragile,  and  the  most 
ready  to  disappear ;  but  the  path  of  atrophy  is 
quite  different  in  the  case  of  the  gradual  reduction 
of  an  organ  in  a  species.  When  an  organ  becomes 
useless  to  a  species,  as  in  the  case  of  the  eyes  of 
deep-sea  Crustacea,  the  only  thing  that  matters  to 
the  species  is  that  it  may  be  got  rid  of.  Any 
individual  variation  tending  towards  reduction  will 
be  of  advantage,  and  may  be  retained  by  natural 
selection.  There  is  no  reason  to  suppose  that  such 
individual  variations  appear  in  any  inverse  order ; 
in  fact  we  do  not  know  that  the  appearance  of 
variations  follows  any  law  at  all.  Perhaps  the 
apparent  inverse  order  of  the  degeneration  of  the 
pineal  eye  in  lizards  may  be  explained  from  the 
fact  that  the  most  recently  acquired  characters  are 
frequently  the  most  variable. 


THE  PATH  OF  DEGENERATION  IN  SOCIOLOGY    206 

However,  even  when  the  most  recent  organs 
disappear  first,  we  cannot  speak  safely  of  a  law 
of  degeneration  inverse  to  evolution.  In  the 
actual  evolution  of  many  organs,  parts  have 
appeared  and  then  completely  disappeared.  If 
degeneration  were  a  true  inverse  of  evolution, 
it  would  be  necessary  that  such  lost  parts  should 
reappear  only  to  disappear  again.  Such  observa- 
tions apply  both  to  ontogeny  and  phylogeny. 


CHAPTEE  II 

THE  PATH  OF  DEGENERATION  IN  SOCIOLOGY 

§  1.  Investigation  of  facts. 

We  have  now  to  see  if  degenerative  evolution  in 
social  matters  retraces  the  steps  of  progressive 
evolution. 

In  the  first  place  the  question  cannot  be  even 
entertained  with  regard  to  some  cases,  and  this  for 
a  general  reason  which  will  be  dealt  with  later  on. 

When  a  complex  institution — such  as  a  com- 
mercial society  or  an  administrative  organization — 
becomes  useless  and  ceases  to  be  fimctional,  it  usually 
disappears  either  by  voluntary  dissolution  or  else  it 
is  legally  suppressed.  In  either  case  there  is  no 
slow  retrogressive  degeneration  retracing  inversely 
the   steps    of    progress,  for  all  the  parts  cease  to 


206        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

exist  simultaneously.  If  certain  parts  of  the  sup- 
pressed institution  are  allowed  to  persist,  these  are 
by  no  means  necessarily  the  oldest  parts,  but  quite 
the  contrary. 

When,  for  instance,  the  Provincial  States  of 
Dauphiny  and  Normandy  were  suppressed  by  the 
French  monarchy,  only  the  titles  with  their  cor- 
responding emoluments  were  allowed  to  remain,  and 
they  were  obviously  of  more  recent  origin  than  the 
States  themselves. 

It  must  be  borne  in  mind  that  all  the  parts  of 
an  institution  rarely  become  simultaneously  useless 
and  non-functional.  Those  which  retain  their  utility 
longest  are  by  no  means  always  the  most  ancient  in 
origin. 

English  sheriffs  have  gradually  become  of  less 
and  less  functional  importance,  and  now  fulfil  no 
other  role  than  that  of  presiding  over  elections  and 
accompanying  the  judges  when  on  circuit.  Both  of 
these  functions  have  been  acquired  recently  com- 
pared with  all  those  which  the  sheriff  discharged  in 
the  days  when  the  care  and  protection  of  the  whole 
county  practically  devolved  upon  him. 

The  question  then  of  the  pathway  of  degeneration 
only  arises  in  those  cases  where  the  same  cause 
of  dissolution  simultaneously  affects  all  parts  of 
the  institution,  and  where,  without  sudden  inter- 
ruption, degeneration  is  effected  slowly  but  surely 
through  many  successive  stages.  This,  of  course, 
happens  in  the  degenerative  evolution  of  individual 


THE  PATH  OF  DEGENERATION  IN  SOCIOLOGY    207 

societies  or  institutions,  and  not  in  the  disappearance 
of  complete  classes  of  institutions. 

These  reservations  being  understood,  we  will 
mention  a  few  more  or  less  obvious  cases  in  which 
degeneration  does  retrace  the  footsteps  of  progressive 
evolution. 

1.  Tithings,  hundreds  and  counties  in  England. — 
In  tbe  chapter  dealing  with  the  pathway  of  degenera- 
tion in  Transformisme  social,  G.  Degreef  mentions 
the  following  interesting  facts  : — 

"  Mr  Herbert  Spencer,  after  describing  the  forma- 
tion of  tithings,  hundreds  and  counties  in  England 
under  the  Anglo-Saxon  regime,  observes  that  the 
tithings  along  with  their  courts  of  justice  were  the 
first  to  disappear,  then  the  hundreds  followed,  though 
some  vestiges  of  their  old  courts  of  justice  remained, 
and  only  the  counties  and  the  county  courts  were 
left  intact.  Now  we  have  historical  proofs  that 
English  counties  along  with  their  courts  of  justice 
were  created  before  the  hundreds,  and  the  hundreds 
before  the  tithings."  ^ 

2.  Order  of  elimination  of  various  racial  elements 
in  a  country. — In  his  interesting  work  Civiliza- 
tion et  d^'po'pulation,^  Dumont  mentions  certain 
facts  which  go  to  show  that  the  inhabitants  of  poor 
districts,  who  are  nevertheless  of  pure  racial  descent, 
have  a  birth-rate  higher  than  that  of  the  members 
of  the  population  who  are  not  aboriginal,  and  who 

1  Degreef,  Le  transformisme  social^  p.  450. 
•^  P.  156. 


208        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

for  the  most  part  dwell  in  the  towns  and  fertile 
plains.  From  this  he  concludes  that  the  various 
racial  elements  of  a  nation  are  eliminated  in  inverse 
order  to  that  in  which  they  were  introduced.  In 
France,  for  instance,  the  Frank  has  been  com- 
pletely absorbed  in  the  Gaul. 

8.   The  degenerative  evolution  of  political  organiza-^ 
tions. — The  progressive  and  degenerative  evolution 
of    political    organizations  has   been   described   by 
Herbert  Spencer  as  follows  ^ : — 

"  Political  integration,  as  it  advances,  obliterates 
the  original  divisions  among  the  united  parts.  In 
the  first  there  is  the  slow  disappearance  of  those 
non-topographical  divisions  arising  from  relation- 
ship, as  seen  in  separate  gentes  and  tribes — 
gradual  intermingling  destroys  them.  In  the 
second  place,  the  smaller  local  societies  united 
into  a  larger  one,  which  at  first  retains  their 
separate  organizations,  lose  them  by  long  co- 
operation ;  a  common  organization  begins  to 
ramify  through  them.  And,  in  the  third  place, 
there  simultaneously  results  a  fading  of  their 
topographical  bounds,  and  a  replacing  of  them  by 
the  new  administrative  bounds  of  the  common 
organization. 

"  Hence,  naturally,  results  the  converse  truth  that 
in  the  course  of  social  dissolution  the  great  groups 
separate  first,  and  afterwards,  if  dissolution  con- 

^  Herbert  Spencer,  "  Political  Institutions,"  Part  iv.  of  Principles 
of  Sociology,  p.  286. 


THE  PATH  OF  DEGENERATION  IN  SOCIOLOGY    209 

tinues,  these  separate  into  their  component  smaller 
groups.  Instance  the  ancient  empires  successively 
formed  in  the  East,  the  united  kingdoms  of  which 
severally  resumed  their  autonomies  when  the  coer- 
cion of  keeping  them  together  ceased.  Instance 
again  the  Carlo vingian  Empire  which,  first  parting 
into  its  large  divisions,  became,  in  course  of  time, 
further  disintegrated  by  subdivisions  of  these.  And 
when,  as  in  this  last  case,  the  process  of  dissolution 
goes  very  far,  there  is  a  return  to  something  like 
the  primitive  condition,  under  which  small  preda- 
tory societies  are  engaged  in  continuous  warfare 
with  like  small  societies  around  them." 

We  may  conclude  then  that  political  integration 
is  attended  by  degeneration ;  primitive  institutions 
disappear  and  make  way  for  fresh  institutions,  and 
their  disappearance  is  permanent.  In  the  course 
of  the  dissolution  of  the  Carlovingian  Empire 
there  was  no  reappearance  either  of  the  gentes 
or  of  the  primitive  tribal  system ;  but  when  this 
vast  organization  broke  down,  it  was  natural  that 
the  more  recently  formed  social  bonds,  having  had 
the  least  opportunity  of  becoming  consolidated, 
should  be  the  first  to  be  sundered. 

4.  Degeneration  in  monetary  systems. — The  prin- 
ciple that  degeneration  retraces  the  steps  of  pro- 
gress applies  equally  to  a  very  different  range  of 
ideas, — the  evolution  of  monetary  systems.  Stanley 
Jevdns  says  that  there  is  little  doubt  that  every 
system  of  coinage  was  originally  identical  with  a 

0 


210    THE  PATH  OF  DEGENERATIVE  EVOLUTION 

system  of  weights.  A  survival  of  this  primitive 
condition  existed  in  Eoman  law,  and  even  when  no 
use  was  made  of  them,  the  custom  of  bringing  a 
pair  of  scales  survived  as  a  legal  formality  in  the 
sale  of  slaves  at  Kome. 

After  the  time  of  the  Punic  wars,  the  aes, 
which  originally  equalled  a  Roman  pound  in 
weight,  diminished  rapidly,  until  it  became  re- 
duced to  the  weight  of  an  ounce.  The  Eomans 
had  naturally  reverted  to  weighing  the  metal,  and 
the  oes  grave  was  money  reckoned  by  weight,  and 
not  by  tale.  Generally  speaking,  whatever  be  the 
inconveniences  of  the  method,  currency  by  weight 
is  yet  the  natural  and  necessary  system  to  which 
people  revert  whenever  the  abrasion  of  coins,  the 
intermixture  of  currencies,  the  downfall  of  a  State, 
or  other  causes,  destroy  the  public  confidence  in  a 
more  highly  organized  system.-^ 

It  is  plain  then  that  the  more  recent  developments 
in  the  coinage  system  are  the  first  to  disappear.^ 

The  disappearance  of  money  altogether  and  the 
return  to  a  system  of  exchange  would  represent  a 
much  farther  advanced  stage  in  degeneration. 

^  See  Stanley  Jevons  in  Money,  International  Scientific  Series. 

2  There  is  no  silver  money  and  only  a  little  copper  in  China. 
Nowadays,  Mexican  piastres,  on  reaching  the  country  in  payment 
of  commercial  transactions,  are  melted  down  into  bars  as 'soon  as 
they  fall  into  the  hands  of  the  merchants,  and  these  bars  are  then 
imprinted  with  the  Chinese  stamp.  This  was  the  usual  system 
employed  amongst  civilized  peoples  before  the  invention  of  money r 
See  Thorold  Rogers  in  ' '  The  Economic  Interpretation  of  History. " 


THE  PATH  OF  DEGENERATION  IN  SOCIOLOGY    211 

5.  Degenerative  adaptation  in*colonial  legislation. 
— In  his  treatise  (Annalisi  delta  proprieta  capita- 
lista),  Loria  furnishes  another  striking  example  of 
the  law  of  degeneration :  "  When  English  colonies 
were  first  formed  in  America,  the  colonists  hesitated 
to  establish  any  legislation  other  than  that  of  the 
mother-country.  They  were  habituated  to  it ;  it 
was  written  in  their  own  language,  and  therefore 
seemed  best  to  correspond  with  their  national 
characteristics.  But,  from  the  outset,  the  greatest 
difficulties  were  met  with  in  the  application  of  this 
legislation  to  the  colonies. 

"  In  the  first  place  the  Statute  law  of  England,  the 
most  recent  addition  to  the  legislation,  was  found 
to  be  quite  unsuited  to  the  economic  condition  of 
a  colony,  and  so  common  law  alone  came  to  be 
established,  which,  being  the  more  ancient,  was 
better  suited  to  the  social  organization  of  a  newly- 
formed  society.  But  even  this  form  of  legislation 
did  not  remain  permanent  under  social  conditions 
profoundly  diflferent  to  those  in  which  it  had  been 
originally  established,  and  the  construction  of  a 
special  legislation  was  found  to  be  necessary.  In 
this  way  the  common  law  of  England  came  to  be 
regarded  as  unsuited  to  her  colonies,  excepting  in 
such  cases  as  were  unprovided  for  in  the  new 
colonial  law."  -^ 

Loria  then  proceeds  to  give  numerous  examples 
of  how  these  colonial  statutes — owing  to  the  simi- 

^  Loria,  Annalisi  delta  proprieta  capitalista,  ii.  48. 


212        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

larity  of  circumstances  between  those  for  whom 
they  were  severally  fashioned — came  to  resemble 
the  primitive  law  of  England. 

6.  Degenerative  evolution  of  the  corporations  of 
Western  Flanders. — In  a  treatise  published  by  one 
of  our  number  in  May  1892,  entitled  VEvohUion 
regressive  des  corporations  de  la  West-Flandre}  it 
was  shown  that  among  these  associations,  the  insti- 
tutions the  first  to  degenerate  were  those  most 
recently  established. 

The  twelve  corporations  which  still  exist,  though 
in  more  or  less  degenerate  conditions  in  Bruges, 
Furnes,  Eeghem,  and  Iseghem,  were  formerly  con- 
structed on  similar  principles  and  fulfilled  the 
following  functions : — 

1.  The  furtherance  of  sociability  {i.e.  the  holding 

of  banquets  and  fetes). 

2.  The   encouragement  of   religious  feeling  {i.e. 

frequent  celebrations  of  the  mass  and  the 
building  of  new  churches). 

3.  Mutual  assistance  {i.e.  insurance  against  loss 

of  work  through  illness,  or  against  funeral 
expenses). 

4.  Mutual  protection  of  professional  interests. 

5.  The  furtherance  of  certain  political  and  mili- 

tary interests. 
These    various    functions    were    established    not 
simultaneously    but    in    succession.     The    Flemish 
corporation  of  the  fourteenth  or  fifteenth  century 
1  La  SocUt6  nouvelle,  Mai  1892,  Bruxelles,  Monnom. 


THE  PATH  OF  DEGENERATION  IN  SOCIOLOGY    213 

represents  the  last  stage  in  a  long  series  of  corpora- 
tions of  different  kinds,  and  of  increasing  complexity. 
It  will  be  as  well  to  glance  briefly  through  these 
various  kinds  of  corporations  in  order  to  compare 
the  different  stages  of  dissolution  through  which  the 
corporative  system  eventually  passed  with  the  pro- 
gressive evolution  exhibited  in  the  course  of  its 
establishment. 

{a)  Associations  formed  for  the  holding  of 
banquets  and  fetes  (convivia)  which  were  origin- 
ally distributed  throughout  Northern  and  Western 
Europe. 

(h)  The  convivia  assumed  a  religious  character 
when  the  Church,  unable  to  suppress  them,  deter- 
mined to  transform  them. 

(c)  Guilds  (etymologically  significant,  according 
to  Brentano,  of  repasts  where  all  expenses  were 
shared)  where  the  original  convivium  was  accom- 
panied by  religious  ceremonies,  but  a  feature  of 
which  was  the  addition  of  institutions  of  mutual 
insurance. 

(d)  Corporations  (amhachten). — These  embodied 
all  the  primitive  institutions  that  had  gone  before : 
(1)  the  banquets;  (2)  religious  ceremonies;  (3) 
mutual  assistance ;  (4)  fresh  means  for  the  protec- 
tion and  development  of  professional  interests. 

(e)  Finally,  at  the  commencement  of  the  four- 
teenth century,  these  corporations  assumed  both 
political  and  military  functions.  The  concession  of 
ke^tre  to  the  people  in    1304  by  Philip  of  Thiette 


214        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

was  the  first  act  by  which  was  recognized  the  right 
of  the  corporations  of  Bruges  to  take  part  in  com- 
munal administration  and  to  provide  a  miUtary 
contingent  of  their  own. 

When  these  corporations  began  to  decline  mili- 
tary and  political  functions  were  the  first  to  be 
eliminated,  in  other  respects  the  corporation  con- 
tinued to  exist  till  the  close  of  the  eighteenth 
century  in  the  form  of  economic  groups,  and  groups 
for  mutual  aid  and  religious  development. 

An  investigation  of  such  corporations  as  still 
survive  shows  that  it  is  in  the  economic  functions 
formerly  discharged  that  degeneration  has  made 
furthest  strides,  while  the  religious  character  is 
maintained  and  banquets  are  still  held.  Of  the 
twelve  corporations  of  Western  Flanders  six  preserve 
the  old  corporative  institutions  almost  intact ;  seven 
maintain  a  scale  of  charges  and  regulations  con- 
nected with  them  ;  ten  provide  organizations  for 
assurance  against  illness  ;  eleven  hold  annual  re- 
ligious ceremonies  ;  twelve,  ix.  all  of  them,  continue 
to  hold  banquets. 

These  figures  alone  show  that  the  original  and 
earlier  functions  have  remained  longest  in  force,  but 
to  further  demonstrate  this  point  it  will  be  necessary 
to  enter  more  fully  into  details  and  to  study  each 
group  separately. 

The  first  group — consisting  of  associations  con- 
tinuing to  protect  professional  interests — consists  of 
three  branches :  the  four  offices  of  Bruges  (including 


THE  PATH  OF  DEGENERATION  IN  SOCIOLOGY     215 

vendors  of  lime,  coal,  seeds  and  beer),  the  two  offices 
of  Furnes  (street  porters  and  vendors  of  beer),  and 
the  community  of  the  bakers  of  Bruges. 

All  the  ancient  statutes  of  the  four  offices  have 
been  preserved,  including  trade  monopolies,  the 
freedom  of  the  city,  and  scales  of  charges,  help 
in.  times  of  illness,  accident,  or  when  out  of  work ; 
religious  ceremonies  and  banquets.  With  the  com- 
munity of  bakers,  however,  this  is  not  the  case; 
here  the  economic  functions  discharged  are  reduced 
to  a  minimum.  The  association  continues  to  exist 
in  a  triple  capacity :  as  a  syndicate  to  keep  up  a 
fair  price  in  wheat  and  bread,  as  a  mutual  assur- 
ance association  in  times  of  distress  among  the 
members,  and  as  a  confraternity  imposing  religious 
obligations  and  holding  an  annual  banquet  and  f^te. 

In  the  second  group  consisting  of  mutual  aid 
associations  such  as  the  corporations  of  tailors,  shoe- 
makers and  weavers  of  Bruges,  all  trade  interests 
have  disappeared,  and  the  corporations  only  exist 
in  the  capacities  of  confraternities  and  mutual  aid 
societies.  At  this  particular  stage  of  degeneration 
these  corporations  resemble  in  a  striking  degree  the 
old  guilds  which  preceded  the  amhachten  en  neringen. 
It  is  interesting  in  connection  with  this,  that  the 
corporation  of  wool  weavers  (wollewevers)  in  Bruges 
has  lost  its  orginal  professional  character,  and  quite 
heterogeneous  elements  have  been  introduced  ;  there 
are  only  twenty-five  weavers  in  Bruges,  and  their 
society  numbers  nearly  two  hundred  members.     The 


216    THE  PATH  OF  DEGENERATIVE  EVOLUTION 

guild  of  weavers  in  Eegliem  represents  the  third 
stage  only — that  of  a  religious  confraternity.  The 
office  of  mutual  aid  has  disappeared,  and  only  the 
Saints'  Day  Fete  and  the  convivmm  remain :  "  The 
first  Sunday  after  the  fete  of  the  Trois-Eois,  which 
is  the  annual  fete  day  of  the  weavers,  the  guild 
proceeds  to  the  parish  church,  headed  by  a  banner, 
accompanied  by  a  jester,  and  with  drums  beating, 
where  mass  is  celebrated  in  honour  of  Saint  Severin. 
The  rest  of  the  day  is  passed  in  diversions." 

The  guilds  of  Saint  Crispin  at  Iseghem,  all  that 
remains  of  the  old  corporation  of  shoemakers,  are 
representative  of  the  final  stage  of  degeneration : 
the  association  has  resumed  its  most  primitive 
character,  and  is  reduced  to  a  mere  dining  society — 
the  primitive  convivium. 

There  are  from  1500  to  2000  shoemakers  in 
Iseghem.  At  the  time  of  the  Ee volution  their 
ambaclit,  having  resumed  its  archaic  form,  was 
divided  up  into  six  or  seven  guilds.  Some  years 
ago  a  vain  attempt  was  made  to  reconstruct  these 
guilds  and  adapt  them  to  modern  requirements. 
At  a  time  when  success  in  this  project  was  still 
hoped  for,  one  of  the  promoters  of  the  reconstruction 
wrote  as  follows  : — 

"  The  members  of  bhe  guild  still  recognize  Saint 
Crispin  as  their  patron  saint,  but  they  no  longer 
assemble  in  church  to  do  him  honour  by  the  cele- 
bration of  mass.  They  keep  the  anniversary  instead 
by   going  from  one  public-house  to  another,  with 


THE  PATH  OF  DEGENERATION  IN  SOCIOLOGY    21 7 

flying  banner  and  drums  beating,  and  the  day  has 
become  merely  a  day  of  copious  libations,  and  serves 
as  a  pretext  for  poor  workmen,  the  fathers  of  families, 
to  spend  all  the  week  drinking  in  public-houses." 

In  conclusion  then  the  cycle  is  complete ;  a 
corporation,  unless  suddenly  dispersed,  ends  as  it 
began  ;  in  the  last  stage  of  decline  it  resembles 
the  associations  from  which  it  originally  developed, 
the  most  recently  established  functions  having  been 
the  first  to  decay  and  disappear. 

§2.-4  criticism  of  the  supposed  inverse  path  of 
degeneration. 

These  few  examples  suffice  to  show  that  in 
certain  cases  the  more  recently  formed  institu- 
tio'bs  are  the  first  to  decline  and  disappear,  while 
the  older  persist  to  the  end. 

It  must  be  remembered,  however,  that  the  con- 
trary is  at  least  as  frequently  the  case.  AU  changes 
of  legislation,  either  juridical  or  religious,  follow, 
but  never  precede,  the  economic  transformations  to 
which  they  relate,  whether  these  be  social  or  ethical, 
unless  the  transformations  are  ephemeral.  "  Imita- 
tion," says  Tarde,  "  proceeds  from  the  more  obvious 
to  the  less  obvious ;  that  is  to  say,  ends  and  feel- 
ings are  imitated  sooner  than  their  means  and 
expressions." 

Title-deeds  and  armorial  bearings  survive  no- 
bility ;  houses  continued  to  be  held  as  personal 
or  moveable  property  long  after  the  disappearance 


218   THE  PATH  OF  DEGENERATIVE  EVOLUTION 

of  nomadic  tribes,  which,  living  as  they  did  in 
tents,   originated  the  conception.^ 

Among  peoples  where  the  system  of  marriage  by 
groups  has  existed,  family  nomenclatures  continued 
to  persist  long  after  the  disappearance  of  the  family 
system  to  which  they  owed  their  origin.  "  The 
family,"  says  Morgan,  "  is  an  active  element,  never 
stationary  ;  it  keeps  pace  with  the  development  of 
society  in  the  march  of  progress.  On  the  other 
hand,  the  reckoning  of  kinship  changes  very 
slowly ;  only  after  long  lapses  of  time  does  it 
register  the  progress  actually  made  by  the  family 
in  the  course  of  ages,  and  does  not  undergo  any 
radical  transformation  until  long  after  the  family 
itself  has  been  completely  changed."  "And,"  adds 
Karl  Marx,  whose  critical  annotations  on  Morgan's 
book  were  carefully  preserved  by  Engels,  "  this  also 
applies  to  systems  of  politics,  law,  religion,  or 
philosophy." 

These  systems,  formed  after  the  completion  of 
the  social  organization  which  they  express,  survive 
after  the  organization  itself  has  disappeared.  Their 
elimination  is  not  of  such  importance  to  society 
as  is  that  of  the  economic  or  family  institutions 
themselves,  as  these,  when  they  become  useless  and 
disadvantageous,  are  a  drag  on  future  development. 

^  Viollet,  Histoire  du  droit  civil  fr.,  p.  617. 

' '  Although  houses  were  for  centuries  treated  as  moveable  pro- 
perty, they  dbntinued  to  be  legally  treated  as  such  for  a  still 
longer  period  of  time  ;  it  is  characteristic  of  judicial  ideas  that 
they  lag  far  behind  economic  progress." 


THE  PATH  OF  DEGENERATION  IN  SOCIOLOGY    219 

It  cannot  be  established,  however,  as  a  general 
principle,  that  the  pathway  of  degeneration  as 
regards  societies  or  institutions,  is  inverse  to  that 
pursued  by  their  progressive  evolution.  In  the 
first  place,  the  mere  explanation  of  this  supposed 
law  shows  that  it  is  quite  untenable. 

What  jeason  is  given  for  supposing  the  decline 
of  memory  or  will  power,  the  degeneration  of  writ- 
ing and  speech,  the  decadence  of  societies  and 
institutions,  to  be  a  retracing  of  the  steps  of 
progressive  development  ?  The  reason  given  is 
that,  other  things  being  equal,  the  more  fragile, 
unsteady  and  complicated  structures  are  the  first 
to  fall. 

Now,  although  the  most  fragile  structures  are 
frequently  those  most  recently  formed,  and  which 
have  not  had  time  to  settle  down  and  firmly  estab- 
lish themselves,  it  is  also  true  that  in  many  cases 
the  more  recent  acquisitions  and  structures  attain 
a  more  solid  basis  than  those  which  have  preceded 
them. 

There  is  nothing  invariable  about  the  pathway  of 
degeneration.  It  can  no  more  be  said  to  retrace  the 
pathway  of  progress  in  an  inverse  direction  than  it 
could  be  said  that  in  a  country  abandoned  by  its 
inhabitants  the  more  recently  formed  paths  of  com- 
munication would  be  the  first  to  become  effaced.  It 
is  quite  true  that  the  broader  roads,  which  would 
naturally  last  longer,  are  frequently  the  oldest 
paths    of    communication ;  whereas   the    footpaths, 


220    THE  PATH  OF  DEGENERATIVE  EVOLUTION 

which  are  the  first  to  disappear,  are  usually  of 
more  recent  origin.  Very  frequently,  however, 
the  new  roads  follow  a  rather  dififerent  direction, 
and,  although  more  recently  constructed,  are  not 
the  first  to  disappear. 

It  is  the  same  with  great  commercial  crises.  It 
is  quite  inexact  to  say  with  Ribot,  who  is  respon- 
sible for  the  analogy  :  "  Old  houses  ofifer  the  best 
resistance  to  the  storm ;  it  is  the  new  houses 
which,  being  less  solid,  crumble  and  fall."  ^ 

After  the  time  of  the  cotton  famine,  during  the 
American  war  of  Independence,  the  greater  part  of 
the  old  firms  of  Gand  became  bankrupt,  whereas 
most  of  the  large,  newly-established  joint-stock 
companies  survived  the  crisis. 

Moreover,  in  those  cases  where  the  most  recently 
formed  structures  are  the  first  to  decay,  it  cannot 
be  deduced  that  evolution  is  reversed,  and  that 
the  institution  returns  to  its  primitive  condition, 
for  there  is  no  reappearance  of  the  intermediate 
structures. 

^  Ribot,  Les  Maladies  de  la  Memoire,  p.  99. 


PART  II 

The  irreversibility  of  degenerative  evolution 

Most  authorities  on  the  subject  are  agreed  that 
evolution  is  not  reversible/  and  that  institutions 
or  organs  which  have  disappeared  or  been  reduced 
to  rudiments  do  not  reappear  and  develop  afresh. 
It  would  be  a  useless  extension  of  this  volume 
to  cite  many  facts  in  favour  of  a  view  which  is 
almost  without  supporters,  but  it  will  be  useful  to 
examine  the  exceptions,  real  or  apparent,  and  to 
discuss — 

1.  If    an   institution   or   organ    which   has   dis- 

appeared may  reappear. 

2.  If  an  institution   or  organ  which   has  been 

reduced  may  resume  its  primitive  function. 

3.  If   an  institution  or  organ  which  has  been 

reduced  may  redevelop  and  assume  a  func- 
tion other  than  its  original  function. 

*  L.   DoUo,  Les  his  de  VJ^voluti&ii  {Soc.  Belg.   04ol.  Fal^ont. 
Hydr.,  t.  vii.,  1893,  proces-verbaux,  pp.  164-166. 


222    THE  PATH  OF  DEGENERATIVE  EVOLUTION 


CHAPTER  I 

DO    INSTITUTIONS    OR    ORGANS    WHICH    HAVE    DIS- 
APPEARED   REAPPEAR  ? 

Section  I 
Disappeared  organs 

In  biology  we  are  almost  unaware  of  indisputable 
examples  of  the  normal  reappearance  of  disappeared 
organs. 

1.  Plants. — As  the  embryonic  development  of 
plants  is  usually  direct,  it  is  impossible  to  decide 
whether  an  organ  which  forms  a  component  part 
of  the  embryological  history  represents  an  ancestral 
organ.  However,  in  a  few  rare  cases,  artificial 
selection  causes  an  actual  reversion  of  evolution. 

Typical  geraniums  possess  two  whorls  of  five 
stamens,  as,  for  instance,  in  Geranium.  In  Erodium 
there  is  only  one  cycle  of  five.  In  Pelargonium  one 
cycle  of  five  is  complete ;  the  other  is  represented 
by  two  stamens  and  three  filaments  which  have 
lost  their  anthers.  But  in  certain  varieties  with 
very  large  flowers  the  two  complete  cycles  re- 
appear, five  stamens  having  long,  and  five  short, 
filaments.  In  this  case  there  is  no  doubt  as  to 
the  reappearance  of  the  three  stamens  lost  in 
typical  Pelargonimns. 


DO  INSTITUTIONS  OK  ORGANS  REAPPEAR?       223 

This  reappearance  is  the  result  of  artificial 
selection.  The  typical  Pelargoniums  have  a  bi- 
lateral symmetry,  but  horticulturists  set  a  higher 
value  on  flowers  with  radial  symmetry,  and  in 
consequence  have  produced  flowers  with  such  a 
symmetry.  As  a  matter  of  fact,  they  have  paid 
attention  only  to  the  symmetry  of  the  petals,  but 
in  modifying  that,  they  have  also  modified  the 
symmetry  of  the  stamens.^ 

The  Privets  (Ligustrum),  like  most  of  the  Oleacese, 
possess  only  two  stamens.  Nevertheless,  it  is  not 
uncommon  to  find  among  normal  flowers  of  the 
common  Privet  specimens  with  three  or  four 
stamens.  In  this  case,  however,  it  is  uncertain 
whether  there  is  a  real  reappearance  of  lost  organs, 
or  if  the  loss  has  not  actually  become  complete. 

Animals. — In  the  case  of  animals,  teratology  and 
embryology  furnish  a  few  exceptional  cases  of  an 
apparent  reversion  of  degenerative  evolution. 

As  an  abnormality  in  the  horse,  the  first,  second, 
fourth,  and  fifth  digits  may  reappear. 

Adult  man  has  lost  the  complete  covering  of 
downy  hair.  According  to  Ecker,  however,  hyper- 
trichosis is  an  abnormality  really  due  to  the  re- 
appearance of  this  ancestral  condition,  as  may  be 
seen  from  the  mode  of  its  distribution  in  whorls. 
Hypertrichosis  is  a  trait  frequently  inherited,  and 
in  this  connection  the  Mauchamp  variety  of  Merino 

^  The  flowers  upon  which  these  observations  were  made  were 
kindly  provided  by  H.  Cannell  of  Swanley. 


224   THE  PATH  OF  DEGENERATIVE  EVOLUTION 

sheep  are  interesting.  This  variety  was  obtained 
by  breeding  from  a  sport  which  appeared  in  a 
normal  flock,  and  which  transmitted  its  peculiarity 
to  its  descendants.  From  time  to  time,  in  normal 
flocks,  variations  occur  which  are  similarly  capable 
of  giving  rise  to  Mauchamp  breeds. 

As  recorded  by  Willett  and  Walsham,  there 
have  been  found  in  human  children  cases  of  a 
bone  stretching  from  the  scapula  to  the  sixth  and 
seventh  cervical  vertebrae.  According  to  these 
authors,  the  bone  represents  the  suprascapula  of 
the  tailless  amphibia,  which  the  normal  homologue 
in  man  is  the  merest  edge  of  the  scapula  ossified 
from  a  separate  centre. 

Such  cases,  as  well  as  cases  of  polydactylism  and 
of  supernumerary  mammae,  are  usually  set  down  as 
atavistic.  However,  the  attempt  to  explain  by 
atavism  such  pathological  and  teratological  peculi- 
arities must  be  made  with  caution.  Such  inherited 
anomalies  occur  very  frequently  in  degenerate 
families — the  neuropathic  families  of  F^re^ — and 
are  associated  with  other  abnormalities  equally 
heritable  and  certainly  not  due  to  atavism.  Such 
are  pigmented  retinitis,  congenital  cataract,  chro- 
matic asymmetry  of  the  iris,  asymmetry  of  the 
pupil,  ichthyosis,  pigmented  erectile  spots  on  the 
skin,  and  congenital  disposition  to  bleeding. 

As  in  a  degenerate  line  of  heredity  these 
abnormalities    may    replace    one    another    iudiffer- 

^  F^r^,  La  Famille  ndvropathiqiie,  1894.     Paris,  F.  Alcan. 


DO  INSTITUTIONS  OR  ORGANS  REAPPEAR  ?       225 

ently  it  is  very  doubtful  if  any  of  them  ought 
to  be  regarded  as  the  inheritance  of  ancestral 
traits.  Teratology  contains  no  undoubtful  case 
of  the  reversion  of  evolution. 

On  the  other  hand,  it  seems  certain  that  in  the 
individual  development  of  some  species  there  is  a 
real  reappearance  of  lost  organs.  The  larval  his- 
tory of  certain  Crustacea  Malacostraca  ^  (cray-fish, 
shrimps,  etc.),  seems  to  provide  instances. 

In  Stomatopoda,  the  youngest  erychtheus  larvae 
(fig.  71,  a)  are  formed  of  three  parts  :  the  head, 
five  anterior  thoracic  segments,  each  bearing  a  pair 
of  biramous  swimming  limbs  (i.-v.),  the  three  last 
decreasing  in  size  from  before  backwards  (these  five 
pairs  of  appendages  represent  the  five  pairs  of  buccal 
appendages  of  the  adult) ;  three  terminal  posterior 
segments  (vi.-viii.),  and  a  caudal  fin,  all  without 
appendages.  In  older  larva?  the  first  and  second 
pairs  are  profoundly  modified,  losing  a  joint  and 
acquiring  gills  (fig.  71,  b,  i.-ii.) ;  the  third,  fourth, 
and  fifth  pairs  disappear  completely,  or  at  most 
occur  as  minute  saccules  (iii.,  iv.,  v.).  New  thoracic 
segments  are  formed,  and  later  on  in  the  third 
larval  stage  the  thoracic  appendages  reappear  in 
their  final  form. 

Similar  facts  occur  in  the  development  of  the 
Decapoda  Macroura,  such  as  Palimterus  and 
Scyllarus.  While  within  the  egg  the  creature 
passes   successively  through  nauplius  and   phyllo- 

^  Lang,  Anatomie  comparde,  vol.  i.,  p.  458. 
P 


226   THE  PATH  OF  DEGENERATIVE  EVOLUTION 


some  stages,  and  possesses  all  its  thoracic  appen- 
dages (three  pairs  of  jaws  and  five  pairs  of 
ambulatory  appendages).  In  this  larval  life  the 
exopodites  of  the  second  and  third  pairs  of  jaws 


Fig.  71. — Erychtheus  larvce  of  Stomatopoda. 
A,  The  younjcest  known  Erychtheus  larva  :  I — V,  buccal  appendages  well  developed.^ 
B,  YounK  Erychtheus  larva  :  I  and  11  modified  buccal  appendages  ;  III— V,  depreneratinf; 
buccal  appendages;  VI— VIII,  segments  devoid  of  appendages.— C,  Older  Erychtheus 
larva:  I — If,  modified  buccal  appendages;  III — V,  reappearing  buccal  appendages; 
VI — VIII,  ambulatory  appendages  in  course  of  formation.  (After  Claus,  in  Lang's  Traitl 
(Tanatomie  comparee,  vol.  i.) 

atrophy.  Shortly  before  hatching,  the  first  pair 
of  jaws  atrophies  completely ;  the  two  pairs  of 
antennae  and  the  two  posterior  pairs  of  ambula- 
tory legs  become  very  degenerate.  In  the  young 
phyllosome  stage  these  organs  are  rudimentary, 
but  in  the  older  phyllosome  larvae   the  first  pair 


DO  INSTITUTIONS  OR  ORGANS  REAPPEAR  ?       227 

of  jaws  and  the  two  posterior  pairs  of  ambulatory 
limbs  are  reformed ;  the  posterior  two  pairs  of  jaws 
reacquire  exopodites,  and  gills  are  formed  on  the 
ambulatory  limbs. 

As  a  matter  of  fact,  in  most  Crustacea,  the 
ambulatory  appendages  appear  when  they  become 
necessary,  that  is  to  say  at  the  end  of  larval  life,  but 
in  the  Stomatopoda  and  in  the  Decapoda  Macroura, 
owing  to  inheritance,  they  appear  much  sooner. 
But  when  these  appendages  are  useless  during  the 
larval  life  they  disappear  again  to  reappear  at  the 
end  of  larval  life  as  in  most  Crustacea.  This 
adaptation  of  the  larva  to  special  conditions  is  of 
great  importance,  as  the  larval  life  is  most  im- 
portant from  the  point  of  view  of  the  species. 


Section  II. 
Disappeared  Institutions. 

The  apparent  revival  of  hygone  institutions. — It 
seems,  at  first  sight,  as  if  there  were  many  instances 
of  the  subsequent  revival  of  bygone  institutions. 

Those  of  ancient  Rome  and  Greece,  for  instance, 
appear  from  time  to  time  to  have  been  recon- 
structed. In  feudal  Rome  of  the  fourteenth  century, 
Cola  di  Rienzi,  by  turns  tribune  and  senator  of  the 
people,  re-established  the  old  republican  constitu- 
tion.    During  the  Renaissance  period  the  ancient 


228   THE  PATH  OF  DEGENERATIVE  EVOLUTION 

schools  of  science  were  resuscitated,  and  during  the 
French  Revolution  similar  attempts  at  revival  were 
made,  especially  in  the  department  of  politics.  When 
Herault  de  S^chelle,  being  ordered  to  draw  up  a 
scheme  of  legislation,  revived  the  laws  of  Minos, 
in  the  constitution  of  the  year  VIII.,  the  Tribunal, 
Senate  and  Consuls  reappeared.  During  the  First 
Empire,  Napoleon,  in  imitation  of  Augustus,  affected 
a  respect  for  republican  institutions,  and  had  the 
coinage  stamped  with  his  own  effigy  and  that  of 
the  Republic.  In  Germany,  the  Holy  Empire 
which  nominally  ceased  to  exist  in  1806,  reap- 
peared in  1871.  In  Greece,  the  Olympic  games, 
suppressed  in  1525,  were  re-established  in  1896. 

It  is  hardly  necessary  to  insist  upon  the  essen- 
tially superficial  nature  of  these  revivals.  It  is 
always  possible  to  bestow  upon  new  social  systems 
the  ceremonial  code  of  an  institution  long  since 
abandoned,  but  it  is  quite  impossible  that  the 
institution  itself  should  be  resuscitated  in  the 
midst  of  surroundings  which  have  been  completely 
transformed.  The  consuls  of  the  year  VIII.  and 
the  emperors  of  modern  times  do  not  resemble  the 
consuls  and  imperatores  of  ancient  Rome  more  than 
the  Christian  societies  of  the  present  day  resemble 
those  of  the  middle  ages.  With  regard  to  outer 
form  in  the  drawing  up  of  statutes,  in  all  which 
constitutes,  so  to  speak,  the  decorative  side  of  the 
institution,  the  organizers  imitated  minutely  the 
keures  of  the  old  amhachtcn ;  beneath  this  appar- 


DO  INSTITUTIONS  OR  ORGANS  REAPPEAR  ?       229 

ent  similitude,  however,  were  structures  of  an 
essentially  different  nature.  Just  as  the  new  rail- 
way station  at  Bruges,  in  spite  of  its  towers  and  its 
pointed  arches,  is  far  more  like  any  other  railway 
station  than  a  Gothic  cathedral,  so  the  Christian 
societies  of  to-day,  in  spite  of  the  archaic  caprices 
of  their  founders,  resemble  more  closely  modern 
associations  than  ancient  associations.  We  see  then 
that  it  cannot  be  said  in  any  of  these  cases  that 
the  actual  revival  of  a  decayed  institution  took 
place.  The  empty  form  reappeared,  but  the  founda- 
tions and  the  essential  parts  had  become  completely 
transformed. 

2.  The  apparent  disappearance  of  institutions. — 
There  are  other  instances,  however,  showing  the 
opposite  of  this  phenomenon.  The  essential  parts 
remain  unchanged,  but  the  form  itself  is  modified ; 
the  institution  persists,  but  its  existence  is  dis- 
sembled. Sometimes  even  when  the  dissolution 
of  an  institution  has  been  enforced,  there  has  been 
a  reconstruction  on  the  first  opportunity. 

An  instance  of  this  is  the  reappearance  of  jjoly- 
gamy  among  the  Mormons,  the  last  traces  of 
polygamy  having  disappeared  during  the  middle 
ages  from  the  people  of  the  West.^ 

1  In  Bigorre  there  was  maintained  up  to  the  fifteenth  century, 
a  kind  of  system  of  concubinage  called  massipia,  which,  though 
not  actually  polygamy,  was  the  contraction  of  an  inferior  union  in 
conjunction  with  real  marriage. 

In  Marseilles,  too,  polygamy  seems  to  have  reappeared  in  the 
middle  ages,  owing  to  the  frequent  intercommunication  between 


230        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

In  ceasing  to  be  legal,  however,  polygamy  has 
by  no  means  ceased  to  exist,  even  in  the  present 
day,  and  the  Mormons  in  instituting  polygamy 
have  only  given  official  recognition  to  what  had 
never  really  disappeared. 

In  the  process  of  social  transformation,  periods  of 
transition  are  frequently  characterized  by  reactions 
in  favour  of  bygone  institutions,  which  reappear 
although  apparently  permanently  abolished.  This 
has  been  the  case  with  certain  corporations,  and 
with  numerous  other  institutions  of  the  old  regime 
which  reappeared  after  the  revolutionary  crisis. 
Some  years  ago  a  large  landed  proprietor  in  the 
domain  of  Peterhoff,  in  Eussia,  attempted,  in  the 
interests  of  his  serfs,  to  introduce  the  rural  system 
of  European  countries.  He  divided  up  the  land 
into  independent  allotments,  and  built  at  his  own 
expense  a  separate  house  for  each  family ;  but  no 
sooner  was  the  abolition  of  serfdom  declared  than 
the  peasants  proceeded  to  re-establish  the  primitive 
community,  and  to  rebuild  their  houses  on  the  old 
sites  in  spite  of  the  considerable  labour  which  this 
entailed.^ 

3.  InstaTices  of  convergence. — It  sometimes  happens 

that  town  and  the  East.  It  was  never,  however,  established  there 
officially  as  was  the  massipia  in  Bigorre.  The  municipality 
promptly  suppressed  it  by  issuing  the  following  mandate  :  ' '  Quod 
(vir)  non  habeat  duas  uxores,  vel  mulier  duos  viros."  (Viollet, 
Histoire  du  droit  civil,  fr.  p.  388. ) 

^  De  Laveleye,  De  la  Propri4U  et  ses  Formes  primitives.     Paris, 
F.  Alcan,  1882,  p.  23. 


DO  INSTITUTIONS  OR  ORGANS  REAPPEAR  ?       231 

that  after  the  lapse  of  several  centuries,  an  institu- 
tion seems  to  reappear. 

An  exact  analogy  to  the  primitive  contuhernium 
(the  community  of  the  cabin)  is  exhibited  in  modern 
slavery.  Only  a  few  years  ago,  in  the  Spanish 
Antilles,  marriage  between  slaves  was  recognized  by 
neither  church  nor  state.  When  a  negro  wished  to 
become  united  to  a  particular  negress  he  asked  per- 
mission of  his  owner  to  share  his  cabin  with  her, 
and  these  unions  could  only  be  dissolved  with  the 
consent  of  the  master.  It  is  hardly  likely  that 
these  slave  marriages  of  the  Spanish  Antilles  are 
survivals  or  rather  resuscitations  of  the  Roman 
contuhernium.  They  rather  represent  a  case  of 
convergence :  identity  in  circumstance  has  been 
productive  of  identity  in  institution.  In  this  case, 
as  in  all  others  of  the  same  kind,  it  cannot  be  said 
that  a  bygone  institution  has  reappeared,  for  the 
new  institution  has  quite  a  different  origin.  Further, 
in  the  other  instances  which  have  been  mentioned, 
an  institution  which  has  reappeared  has  never  really 
ceased  to  exist ;  a  real  dissolution  has  never  been 
followed  by  a  resuscitation.  For  this  to  happen,  the 
whole  social  surroundings  would  have  to  be  trans- 
formed into  their  former  condition,  which  is  obviously 
impossible. 


232        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

CHAPTEE  II 

CAN  RUDIMENTARY  INSTITUTIONS  OR  ORGANS  RESUME 
THEIR  PRIMITIVE  FUNCTIONS  ? 

There  is  no  break  of  continuity  between  a  rudi- 
mentary organ  and  the  complete  ancestral  organ: 
the  rudimentary  organ  is  the  ancestral  organ  trans- 
mitted by  inheritance.  It  is  the  same  organ  not 
only  because  it  has  the  same  form,  but  because  it  is 
actually  a  part  of  it.  For  the  reduced  organ  to 
return  to  its  ancestral  condition  and  resume  its 
ancestral  function,  it  must  retrace  the  series  of 
steps  in  degeneration  along  which  it  has  passed. 

So  also  in  sociology  there  are  many  instances  of 
institutions  which  are  rudimentary,  but  which  retain 
their  original  form  because  of  uninterrupted  imitative 
transmission.  In  their  case  also  resumption  of  the 
primitive  functional  activity  would  imply  a  retracing 
of  the  degenerate  steps. 

This  necessity  of  retracing  shows  at  once  that 
after  a  certain  amount  of  degeneration  resumption 
becomes  impossible.  We  shall  find  in  the  few  cases 
we  are  able  to  adduce,  that  when  organs  retrace 
their  steps  and  resume  their  ancestral  functions, 
degeneration  had  not  gone  very  far  in  them. 

Section  I. 
Rudimentary  organs. 

I.  Animals. — Among  animals  it  is  very  unusual 
for  a  rudimentary  organ   to  become  active  again. 


CAN  ORGANS  RESUME  THEIR  PRIMITIVE  FUNCTIONS?  233 

The  only  cases  show  that  in  them  no  great  amount 
of  degeneration  had  taken  place. 

1.  Muscles  of  the  ear  in  man. — It  is  known  that 
the  human  ear  possesses  a  number  of  intrinsic  and 
extrinsic  muscles  reduced  to  delicate  fibres,  and 
incapable  of  producing  movement  of  the  whole  ear 
or  of  one  part  of  the  ear  on  another  part.  In  some 
abnormal  persons,  however,  certain  of  these  muscles 
may  be  well  developed,  making  movements  of  the 
ear  possible. 

2.  The  abdomen  and  appendages  in  deep- sea 
hermit-crabs. — The  hermit-crabs  of  the  deep  sea  are 
another  instance  of  reversion  to  an  ancestral  form. 
Littoral  hermit-crabs  inhabit  the  spiral  shells  of 
Gastropods,  and  to  suit  this  mode  of  life  the  body  is 
unsymmetrical,  the  appendages  of  one  side  being 
rudimentary.  In  the  depths  of  the  ocean  such 
spiral  shells  are  rare,  and  the  crabs  either  abandon 
this  mode  of  life  or  live  in  straighter  shells.  In 
consequence  the  limbs  and  the  abdomen  become 
nearly  symmetrical  again.  It  is  plain,  however,  that 
in  the  littoral  crabs  these  structures  are  not  truly 
rudimentary. 

II.  Plants. — In  plants  it  is  very  difficult  to 
distinguish  between  the  reappearance  of  lost  organs 
and  the  formation  of  new  organs. 

1.  Hermaphrodite  flowers  in  Melandryum. — The 
Hermaphrodite  flowers  of  melandryum  (fig.  59)  may 
be  flowers  which  after  being  unisexual  have  again 
become  hermaphrodite,  or  they  may  have  retained 


234        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

the  primitive  type.  We  cannot  decide  between  the 
alternatives. 

2.  Branches  of  Colletia  cruciata,  Crataegus,  Vicia 
Faba,  etc. —  Some  cases,  however,  point  clearly  to 
a  renewed  development  of  rudimentary  organs. 
Here  are  some  examples.  (See  also,  further  on, 
page  244  on  hybrid  individuals  of  Pentstemon.) 

Colletia  cricciata  (fig.  72)  in  the  normal  adult 
condition  bears  large  flattened  branches,  which 
serve  for  assimilation  and  possess  only  very  rudi- 
mentary leaves.  Sometimes,  however,  the  plant  may 
give  rise  to  more  slender  branches  with  normal 
assimilating  leaves.  These  branches  and  leaves  are 
probably  the  reappearance  of  the  ancestral  condition. 

Wild  pear  and  apple  trees  produce  small  lateral 
branches  which  are  transformed  into  spines.  These 
thorns  have  evidently  arisen  from  normal  lateral 
branches  which  originally  bore  leaves.  In  the 
cultivated  varieties  these  lateral  branches  have 
resumed  the  leaf-bearing  habit. 

In  the  hawthorn  (Crataegus)  the  lateral  branches 
are  similarly  modified  into  spines.  None  the  less, 
while  these  spines  are  still  young  they  may  be 
artificially  stimulated  to  produce  leaves  by  cutting 
the  principal  stem. 

The  branches  of  Vicia  fdba  bear  low  down  a  set 
of  rudimentary  leaves.  If  the  main  stem  be  lopped 
while  still  quite  young,  the  usually  rudimentary 
leaves  grow  to  the  normal  size.^ 

^  Goebel,  Beitrdge  zur  Morphologie  und  Physiologie  des  Blattes. 
Bot.  Zeit.,  1880. 


kC{M/^ 


FiQ.  72.— Colletia  cruciata  (after  Goebel,  Pflanzenhiologische  Schilderungen,  vol.  I,,  p.  17). 
A,  normal  branch.    B,  branch  recurring  to  ancestral  form ;  (natural  size). 


236 


THE  PATH  OF  DEGENERATIVE  EVOLUTION 


Many  plants  such  as  Semper vivum'^  (fig.  73) 
normally  possess  a  very  much  shortened  stem. 
However,  if  the  plant  be  grown  in  a  saturated 
atmosphere  the  internodes  of  the  stem  lengthen  out. 


Fig.  73.—Sempervivum  tectorum. 
A  (left  figure),  normal  branch;  B  (right  figure),  branch  grawn  for  several  months  in  a 
saturated  atmosphere. 

Certain  Veronicas  (fig.  74)  bear  only  small  scaly 
leaves.  Cultivation  of  these  plants  in  an  atmosphere 
saturated  with  water,  results  in  the  appearance  of 
normal  leaves. 

Lastly,  we  may  quote  again  the  instances  given 
by  Goebel  of  the  production  by  Equisetuvi  arvense 
of  normal  leaves  under  special  conditions.^ 

^  The  branch  jfigured  here  was  gi-own  bj'^  G.  Clautriau,  in  the 
Brussels  Botanical  Institute. 

2  Goebel,  Ueber  die  Fruchtsprosse  der  Equiseten.  {Ber.  d.  d.  Bot. 
Ges.),  vol.  iv.  p.  184,  1886. 


CAN  ORGANS  RESUME  THEIR  PRIMITIVE  FUNCTIONS?  237 


Section  II. 
Rudimentary  institutions. 

It  seems,  at  first  sight,  as  if  some  societies  of 
the  present  day  furnished  instances  of  a  return  to 
primitive  conditions. 

Modern  developments  in  finance  seemed  to  tend 
towards  a  return  to  the  exchange  system.  Politi- 
cal institutions, 
after  a  period  of 
absolutism,  point 
anew  towards  de- 
mocratic equality. 
Corporations  re- 
appear in  the  form 
of  syndicates  or 
religious  societies. 
Landed  property, 
formerly  collective 
and  now  individual,  seems  to  be  again  tending 
towards  collectivism. 

The  same  phenomenon  occurs  in  the  evolution 
of  matters  relating  to  maritime  rights.  The  sea, 
according  to  Eoman  law,  was  equally  open  to  all 
maritime  nations.  Later  on  it  has  been  from  time 
to  time  practically  in  the  hands  of  a  few  nations, 
and  we  have  now  returned  to  a  condition  in  which 
it  is  equally  open  to  all.^ 

1  Tarde,  Transformation  dii  droit,  pp.  161-162.   Paris,  F.  Alcan, 
X890. 


Fig.  74. — Veronica  cupressoides  (after  Goebel, 
Pflanzenbiologische  Schilderungen,  vol.  i.  p.  19). 
End  of  a  branch  grown  under  a  bell-jar  in  a 

saturated  atmosphere.    In  the  oldfr  parts  (B), 

the  leaves  are  small  and  applied  to  the  stem ; 

in  the  younger  parts  (A),  the  leaves  are  larger 

and  protiude  from  the  stem. 


238        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

These,  however,  are  not  cases  of  true  revival. 
The  resemblance  goes  little  farther  than  the  name. 

A  comparison  of  modern  institutions  with  such 
survivals  of  primitive  institutions  as  continue  to 
exist  will  demonstrate  this  point.  The  difference 
between  them  is  so  wide  that  it  would  be  hardly 
possible  to  utilize  the  old  as  a  basis  upon  which  to 
form  the  new.^ 


^  Cf.  Durckheim,  Les  Regies  de  la  mdhode  sociologique.  Paris, 
F.  Alcan,  1895. 

In  sociology,  dealing  as  it  does  with  things  familiar  to  us  all, 
such  as  the  family,  property,  crime,  etc.,  it  is  useless  to  attempt 
to  adhere  to  exact  definitions.  The  exact  meaning  of  some  words 
in  common  use  in  conversation  cannot  be  defined  with  any  pre- 
cision ;  the  common  acceptation  of  these  words  cannot  be  avoided. 
Now  this  common  acceptation  is  frequently  very  ambiguous,  so 
that  two  totally  diflFerent  things  are  often  referred  to  under  the 
same  name,  causing  hopeless  confusion. 

There  are,  for  instance,  two  diff'erent  kinds  of  monogamous 
unions — those  so  only  in  point  of  fact,  and  those  which  are 
legally  so.  In  the  first  case,  a  man  has  only  one  wife,  though 
legally  entitled  to  several  ;  in  the  second  he  is  only  legally 
entitled  to  one.  These  two  kinds  of  conjugal  conditions  are 
quite  different,  and  yet  the  same  word  serves  to  express  both  ; 
it  is  commonly  said  of  some  animals  that  they  are  monogamous, 
although  there  can  be  nothing  approaching  to  a  legal  contract 
between  them.  Spencer,  when  dealing  with  the  subject  of 
marriage,  makes  use  of  the  terra  monogamy  without  defining 
it  in  its  common  and  equivocal  sense.  The  result  of  this  is 
that  the  evolution  of  marriage  seems  to  him  to  represent  an 
incomprehensible  anomaly.  It  seems,  according  to  him,  that 
the  superior  or  monogamous  form  of  union  was  prevalent  during 
the  primitive  phases  of  historic  development ;  that  it  then  dis- 
appeared during  an  intermediate  period,  to  subsequently  reappear. 
From  this  he  concludes  that  there  is  no  regular  connection  between 


CAN  ORGANS  RESUME  THEIR  PRIMITIVE  FUNCTIONS?  239 

It  may  be  definitely  asserted  then  that  a  re- 
duced, but  still  persistent,  institution  never  again 
becomes  actively  functional.  The  following  are  a 
few  examples  which  will  serve  to  illustrate  this 
point : — 

1.  The  truck  system  and  cleariTig-house. — Some 
forms  of  the  primitive  system  of  exchange  sur- 
vive, not  only  in  countries  where  money  is 
unknown,  but  in  certain  industries  where  the 
workers  continue  to  be  paid  in  kind  (the  truck 
system). 

On  the  other  hand,  there  seems  to  be  a  modern 
tendency  towards  the  elimination  of  money  as  an 
instrument  of  exchange.  The  clearing-house  system 
is  singularly  analogous  to  the  old  exchange  system. 
"The  truck  system,"  says  Stanley  Jevons,  "represents 
the  first  and  the  last  stage  ;  but  it  appears  for  the 
second  time  in  a  very  different  form.  Gold  and 
silver  money  continue  theoretically  to  be  the 
instrument  for  buying  and  selling,  but  practically 
metal  no  longer  constitutes  the  real  medium  of 
exchange,  and  has  ceased  to  pass  from  the  hands 
of  the  purchaser  into  those  of  the  vendor." 

In  this  transformation  there  is  obviously  no 
return  to  primitive  systems,  the  last  vestiges  of 
which,  far  from  being  revived,  are  rapidly  dis- 
appearing. 

social  development  in  general  and  a  progressive  advance  towards 
an  improved  system  of  family  life.  A  more  exact  definition  would 
have  prevented  this  erroneous  conclusion. 


240    THE  PATH  OF  DEGENERATIVE  EVOLUTION 

The  suppression  of  the  truck  system  coincides 
with  the  development  of  the  clearing-house  system. 

2.  Corporations  and  syndicates. — The  radical 
differences  existing  between  the  corporations  of 
former  days  and  the  greater  part  of  modern  pro- 
fessional associations  has  already  been  pointed  out. 
The  ecclesiastical  associations,  however,  of  the 
present  day  are  modelled  as  closely  as  possible 
upon  mediaeval  institutions.  It  does  not  follow 
that  the  last  remaining  vestiges  of  the  latter  have 
been  revived.  There  seems  to  be  evidence  that 
quite  the  contrary  has  taken  place. 

At  Bruges  no  attempt  was  made  by  the  founders 
of  the  guild  of  ambachten  to  resuscitate  such 
mediaeval  corporations  as  continue  to  exist  in  a 
state  of  decline. 

At  Iseghem,  a  small  town  in  the  west  of 
Flanders,  we  have  already  seen  that  the  corpora- 
tion of  shoemakers  was  divided  up  into  six  or 
seven  guilds  at  the  time  of  the  Eevolution.  An 
attempt  was  made  a  few  years  ago  to  reconstruct 
and  modernize  these  guilds,  but  the  scheme  fell 
through.  A  new  corporation — wholly  disconnected 
with  the  guilds  of  Saint  Crispin,  and  with  no 
structural  resemblance  to  them — was  established 
instead.^ 

3.  Archaic  collectivism  and  modern  collectivism, — 
Societies    of    the    present    day    exhibit    numerous 

^  Emile  Vandervelde,  EnquUe  sur  les  Associations  professionelles 
d'ouvriers  el  d' artisans  en  Belgique,  i.,  p.  17,  Bruxelles,  1891. 


CAN  ORGANS  EESUME  THEIR  PRIMITIVE  FUNCTIONS?  241 

vestiges  of  archaic  collectivism.  The  question 
arises  as  to  whether  there  is  a  tendency  in  the 
modern  school  of  collectivism  to  resuscitate  such 
vestiges  as  remain  of  the  old  archaic  form  of  col- 
lectivism. Far  from  this  being  the  case,  collective 
property,  as  conceived  by  the  modern  socialist, 
implies  the  suppression  of  the  few  existing  remnants 
of  archaic  collectivism. 

Inheritance,  ah  intestat,  for  instance,  is  a  survival 
from  the  days  of  the  family  community,  which 
itself  arose,  as  we  have  already  seen,  from  the 
primitive  community.  If  the  modern  coUectivist 
school  had  any  desire  for  a  return  to  the  old 
primitive  community,  it  would  make  for  the  re- 
construction of  the  family  community  by  re-estab- 
lishing the  law  of  collateral  succession.  Now  it 
is  just  the  opposite  with  the  collectivists.  In 
order  to  establish  a  universal  system  of  collective 
property  they  demand  among  other  things,  the 
suppression  of  inherited  succession,  ah  intestat,  at 
least  as  regards  the  collateral  line  of  descent. 

4.  The  survival  of  elective  sovereignty  in  Eng- 
land.— The  above  examples  apply  to  institutions 
which  have  degenerated  without  having  completely 
ceased  to  be  functional. 

It  very  rarely  happens,  however,  that  having 
arrived  at  that  condition,  they  renew  their  vitality 
and  all  their  former  functions,  and  this  still  more 
rarely  occurs  in  cases  of  genuine  survival. 

In    the    English    coronation  -ceremony   vestiges 


I 


242        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

remain  of  the  old  democratic  system  in  which  the 
king  was  elected  by  the  people.^ 

The  English  sovereignty  of  the  present  day  is 
merely  a  decorative  institution,  the  real  head  of  the 
Government  being  the  Prime  Minister,  who  is 
nominated  in  fact  if  not  in  theory  by  the  public. 
This  system  may  almost  be  regarded  as  a  return 
to  bygone  democracy.  Nobody  would  wish,  how- 
ever, to  revive  the  old  system  of  elective  sovereignty, 
and  to  retrace  in  an  inverse  direction  the  various 
stages  of  its  degeneration. 

CHAPTER  III 

CAN  RUDIMENTARY  ORGANS  OR  INSTITUTIONS  RE- 
DEVELOP AND  ASSUME  NEW  FUNCTIONS  ? 

The  few  facts  which  we  are  able  to  cite  on  this 
subject  must  be  received  with  considerable  caution. 

^  The  formality  of  an  election  disappeared  during  the  Tudor 
period.  The  coronation  of  Henry  VIII.  was  the  last  occasion  on 
which  the  formula  was  read  which  set  forth  the  national  agree- 
ment with  and  recognition  of,  the  succession.  The  king  was,  in 
fact,  declared  chosen  and  elected.  This  formula  of  election,  which 
disappeared  after  the  coronation  of  Henry  VIII, ,  is  recalled  to  mind 
by  the  conclusion  of  the  coronation  ceremony  of  the  present  day. 
The  archbishop,  walking  in  succession  to  all  four  corners  of  the 
platform  upon  which  the  throne  is  placed,  addresses  the  people  in 
the  following  terms  :  ' '  Gentlemen,  I  herewith  present  to  you  the 
undisputed  sovereign  of  the  realm.  Come  all  who  are  present  and 
offer  homage  to  him.  Are  you  prepared  to  offer  it?"  and  the 
people  signifying  their  assent  by  acclamation,  cry,  "God  save  the 
Queen  "  or  "God  save  the  King."  (De  Franqueville,  Le  gouverne- 
ment  ct  Ic  parleinent  Britanniques,  i.,  p.  291.) 


CAN  ORGANS  REDEVELOP  NEW  FUNCTIONS  ?     243 

Section  I. 
Bvdimentary  organs, 

1.  Animals. — In  Birgus  latro  (a  land-crab  of  the 
Philippines),  the  gills  are  atrophied  and  the  bronchial 
chamber  is  very  richly  supplied  with  blood  vessels, 
while  a  kind  of  incipient  lung  is  formed  from  the 
lining  membrane  of  the  reduced  bronchial  chamber.^ 

However,  it  is  by  no  means  certain  that  the 
atrophy  of  the  bronchial  apparatus  has  preceded 
this  development  of  a  pulmonary  apparatus.  In 
the  following  case  it  rather  seems  to  be  one  in 
which  a  rudimentary  structure  has  redeveloped  in 
order  to  assume  a  new  function.  In  the  develop- 
ment of  the  urinary  organs,  it  appears  that  the 
ducts  of  the  mesonephros  are  quite  independent  of 
those  of  the  pronephros,  although  these  mesonephric 
ducts  become  functional  later  in  the  embryonic  life 
than  the  pronephric  ducts.  They  are,  neverthe- 
less, formed  at  an  earlier  stage,  and  their  rudiments 
have  appeared  before  there  is  any  trace  of  the  others. 
From  this  fact  it  would  appear  that  in  some 
ancestors  of  existing  vertebrates  there  existed  simul- 
taneously mesonephric  canaliculi  and  canals  homo- 
logous with  them,  but  exercising  a  different  function. 
Such  a  condition  actually  exists  in  Amphioxus :  in 
the  branchial  region  of  that  animal  there  are  pro- 
nephric urinary  canals  and  genital  chambers  which 

^  Semper,  "The  Natural  Conditions  of  Existence  as  they  affect 
Animal  Life."     (International  Scientific  Series.) 


244        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

are  homologous  with  mesonephric  spaces ;  but  these 
latter  do  not  exist  as  genital  chambers  in  higher 
vertebrates/ 

It  must  be  noted  that  the  homology  between 
mesonephric  spaces  and  the  genital  spaces  of 
Amjohioxus,  as  made  by  these  writers,  is  not  uni- 
versally accepted. 

2.  Plants.  —  The  Scophulariaceae,  which  have 
usually  four  stamens,  are  derived  from  ancestors 
which  possessed  five.  Usually  the  fifth  stamen  is 
only  represented  by  a  tiny  process  which  rapidly 
atrophies.  However,  in  Pentstemon  the  fifth  or 
posterior  stamen  is  developed,  not  as  a  functional 
stamen,  but  as  a  staminode,  the"  function  of  which  is 
to  stretch  open  the  flower  to  make  it  accessible  to 
hymenopterous  insects  with  short  probosces. 

Can  it  be  said  that  in  such  cases  a  rudimentary 
organ  has  really  become  redeveloped  to  assume  a 
new  function?  To  establish  this  it  would  be 
necessary  to  show  in  the  case  we  have  just 
mentioned,  that  the  stamen  did  not  become  trans- 
formed directly  into  a  staminode,  but  that  it  first 
became  rudimentary  and  then  developed  afresh  into 
a  staminode. 

An  interesting  fact  is,  that  in  some  hybrid 
varieties  of  Pentstemon  the  staminode  becomes  fertile 


^  Boveri,  Die  Nierencanalclien  der  Amphioxus.  {Zool.  Jdhrhuch. 
Ahth.  Anat.  und  Ontogenie  der  Thiere,  vol.  v.,  1892.) 

Wiedersheim,  Grundziige  der  Vergleichenden  Anatomie  der  Wir- 
helthiere.     Jena,  1893. 


CAN  ORGANS  REDEVELOP  NEW  FUNCTIONS  ?     245 

again.  In  some  flowers  sent  to  us  by  Mr  Cannell 
of  Swanley,  the  number  of  petals  was  increased  to 
six,  seven,  eight,  or  nine.  In  some  of  these  the 
posterior  stamen  was  sterile  and  like  a  staminode ; 
in  others  there  were  five  fertile  stamens.  It  is 
obvious  that  in  this  case  the  staminode  had  resumed 
its  original  function  after  having  lost  it. 

Section  II. 
Rudimentary  institutions. 

The  Levirat. — In  his  work  entitled  Tableau  des 
origines  et  de  revolution  de  la  famille  et  de  la 
propriety,  Kovalewsky  mentions  an  instance  of  a 
reduced  institution  which,  without  having  first 
ceased  to  be  functional,  became  transformed  into 
another  institution. 

"  The  custom  mentioned  in  the  Bible  of  alloting 
a  woman  to  the  brother  of  her  deceased  husband,  is 
explained  by  the  primitive  condition  of  things  with 
regard  to  the  relations  between  the  sexes ;  all  the 
women  were  the  common  property  of  the  men 
belonging  to  one  group  of  relations.  Under  the 
name  of  levirat,  this  custom  survived  for  several 
centuries,  owing  to  the  idea  which  arose  later  on 
that  a  wife  was  property.  Consequently,  on  the 
death  of  the  husband,  the  widow,  along  with  his 
other  belongings,  was  treated  as  the  inheritance  of 
the  person  whom  the  death  promoted  to  the  rank 
of  chief  or  head  of  the  family  community." 


246        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

The  levirat,  a  family  institution,  thus  derived 
from  the  old  system  of  marriage  by  groups,  was 
transformed  by  degrees  into  an  economic  institution. 
It  is  important  to  notice,  however,  that  this  trans- 
formation was  effected  without  the  institution  having 
even  been  reduced  to  the  condition  of  a  mere 
survival. 


PART  III 

SUMMARY  AND  CONCLUSIONS 

From  all  the  facts  that  we  have  brought  together, 
the  general  conclusion  becomes  plain  that  retrogres- 
sion, notwithstanding  the  etymology  of  the  word, 
does  not  imply  a  return  to  the  ancestral  condition. 

Eudimentary  organs  and  institutions  resemble  the 
primitive  states  of  these,  in  so  far  as  they  no  longer 
possess  certain  parts  which  the  primitive  stages  did 
not  yet  possess.  None  the  less,  profound  differences 
exist  between  the  primitive  and  the  reduced  forms. 
In  the  primitive  condition  the  institution  or  organ 
is  capable  of  varying  in  the  direction  of  new  uses ; 
in  the  reduced  form,  after  a  certain  degree  of 
atrophy,  there  is  no  longer  the  possibility  of 
redevelopment  to  resume  old  or  to  acquire  new 
functions.  These  observations  apply  equally  to 
biology  and  to  sociology. 

Magnan  and  Legrain,  in  their  work  on  de- 
generate persons,  came  to  similar  conclusions. 
They  came  to  regard  degenerate  persons  as 
abnormal,  chiefly  because  they  were  devoid  of 
the  power  to  reacquire  the  normal  condition  and 
quite  unlike  their  primitive  ancestors,  who,  although 

247 


248        THE  PATH  OF  DEGENERATIVE  EVOLUTION 

possibly  brutal  and  unintelligent,  were  normal 
beings  with  the  activity  and  stamina  necessary 
for  future  progress. 

The  following  two  diagrams,  borrowed  from  these 
authors,  represent  clearly  the  differences  between  the 
initial  and  reduced  condition  of  an  organ  or  institu- 
tion : — 


In  the  diagrams  the  ascending  lines  represent 
the  progressive  evolution  of  an  organ  or  institu- 
tion ;  the  descending  lines  represent  the  degenera- 
tive evolution.  From  the  point  a,  representing 
the  primitive  condition  progressive  evolution  passes 
towards  o,  an  imaginary  perfect  condition  of  the 
organ.  Along  the  upward  line,  however,  the  points 
a,  b,  c,  d,  etc.,  represent  obstacles  to  further  pro- 
gress— that  is  to  say,  factors  tending  towards 
degeneration.  From  these  points  lines  of  de- 
generation pass  towards  z,  and  the  condition  at 
z,  although  representing  that  at  a,  is  not  identical 


SUMMARY  AND  CONCLUSIONS  249 

with  a,  and  is  not  reached  by  a  sliding  backwards 
down  the  line  o,  a. 

Thus,  although  the  most  recently  acquired 
features  may  disappear  first,  degeneration  is  not 
an  actual  retracing  of  steps  until  the  point  of 
departure  is  reached.  The  degenerate  condition 
is  a  new  point,  and  really  the  term  retrogressive 
evolution  is  misleading. 


BOOK  III 

CAUSES  OF  DEGENEKATIVE 
EVOLUTION 

PART  I 

ATROPHY  OF  ORGANS  AND  INSTITUTIONS 

The  factors  of  atrophy 

The  causes  which  are  active  in  producing  degenera- 
tion are  various,  but  they  may  all  be  referred  to  the 
limited  nature  of  the  means  of  subsistence,  that  is 
to  say,  of  nourishment  in  the  case  of  organisms, 
and  of  capital  and  labour  in  the  case  of  institu- 
tions. This  limitation  produces  a  struggle  between 
the  individuals  (societies  or  organisms)  and  between 
their  component  parts. 

In  the  course  of  the  perpetual  struggle  for 
existence  among  the  different  parts  of  an  indivi- 
dual, the  institutions  or  organs  which  have  ceased 
to  be  functional  tend  to  disappear,  their  nourish- 
ment being  absorbed  by  the  active  parts. 

1.  Biology. — In  biology  the  struggle  for  existence 
among  component  parts  appears  clearly  as  a  factor 
of  degeneration  in  the  case  of  accidental  atrophy. 
This  is  to  be  seen,  for  instance,  in  the  atrophy  of 

ass 


252         CAUSES  OF  DEGENERATIVE  EVOLUTION 

the  leaves  of  an  etiolated  plant  ^  or  of  the  muscles 
of  a  limb  which  has  been  immobile  for  long,  or  in 
the  case  of  muscles  which  have  become  inactive 
from  disease  of  the  central  nervous  system.  The 
results  are  similar  in  cases  of  normal  atrophy. 
In  frogs,  toads,  and  other  Batrachia  Anura,^  the 
disappearance  of  the  tail  before  the  adult  state  is 
reached  is  the  result  of  a  struggle  amongst  the 
cells.  The  active  protoplasm  of  the  muscular 
fibres  develops  specially,  and  gives  rise  to  many 
cells,  which  enter  the  contractile  material  and 
separate  its  elements.  Gradually  all  the  contrac- 
tile material  is  absorbed  by  these  isolated  cells. 

Many  plants,  especially  Sempervivum  (see  fig. 
73,  A,  p.  236)  possess  a  reduced  stem  with  the 
leaves  closely  massed  upon  it.  This  reduction 
of    the    stem,    which    is    nearly    constant    in    the 

^  When  a  cutting  from  a  potato  or  a  seed  (fig.  75)  is  allowed  to 
sprout  in  the  dark,  the  young  stems  assume  characters  different 
from  those  of  plants  grown  in  light.  The  absence  of  chlorophyll 
produces  important  modifications  of  growth.  In  light  the  stem 
is  short,  and  the  leaves  are  large  and  expanded  ;  in  darkness  the 
stem  is  very  long,  and  the  leaves  are  much  reduced.  This  atrophy 
of  the  leaves  is  the  result  of  the  struggle  for  existence  amongst  the 
organs  of  the  plant.  Light  increases  the  rate  of  transpiration, 
which  is  chiefly  due  to  the  presence  of  chlorophyll.  As  chloro- 
phyll is  most  abundant  in  the  leaves,  the  transpiratory  current 
sets  strongly  towards  them,  carrying  in  it  the  nutritive  materials 
for  the  formation  of  new  cells.  On  the  other  hand,  in  etiolated 
plants,  transpiration  is  slower,  and  the  nutritive  materials  delayed 
in  the  stem  give  that  the  opportunity  for  specially  active  growth, 
which  takes  place  at  the  expense  of  the  leaves. 

2  Metchnikoff,  Annates  Inst.  Fasteur,  January  1892. 


THE  FACTORS  OF  ATROPHY 


253 


species,  is  the  result  of  a  struggle  amongst  the 
organs  produced  by  a  scanty  water  supply.  The 
leaves  attract  to  themselves  the  greater  part  of 
the  water  absorbed  by  the  roots, 
and  thus  retard  the  growth  of 
the  stem.  But  if  the  plant  be 
cultivated  in  an  atmosphere  satur- 
ated with  water,  the  struggle  be- 
tween the  leaves  and  stem  is 
stopped,  and  the  stem  grows  to 
a  much  greater  extent  (fig.  73, 
B,  p.  236). 

Vicia  Faba,  like  most  vetches, 
produces  at  germination  rudi- 
mentary leaves,  and  similar  leaves 
are  borne  at  the  base  of  each 
branch.  Goebel  has  shown  that 
these  rudimentary  leaves  may  be 
made  to  grow  by  cutting  away 
the  terminal  buds  at  a  young 
stage.  In  this  way  the  struggle 
between  the  first  formed  and  later 
leaves  is  suppressed. 

In  the  case  of  the  individual, 
atrophy  results  from  the  struggle 
for  existence  amongst  the  organs. 
In  atrophy  throughout  a  species  it  is  the  struggle 
for  existence  amongst  individuals  that  plays  the 
chief  part.  Clearly  in  this  struggle,  useless  organs 
become  impediments  and  burdens.     If  any  organs 


Fig.  75.  —Two  seedlings 
of  Cicer  arietinum. 

The  seeds  were  planted  at 
the  same  time.  A, 
seedling  grown  in  light. 
B,  seedling  grown  in 
darkness. 


254         CAUSES  OF  DEGENERATIVE  EVOLUTION 

are  useless  they  are  harmful  as  they  use  nutrition 
without  conferring  any  advantage  upon  the  whole 
organism.  Darwin  pointed  out  that  when  through 
changed  environment  a  structure  became  useless, 
its  degeneration  became  certain,  as  it  was  a  dis- 
advantage to  the  individual  to  squander  nourish- 
ment upon  a  useless  part.  Weismann  has  shown 
that  such  a  reduction  or  disappearance  of  a  useless 
organ  is  the  result  of  variation  and  natural  selec- 
tion.-^ 

Variation  results  in  the  appearance  of  individuals 
with  the  useless  organ  in  various  stages  of  imperfect 
development ;  natural  selection  perpetuates  these 
advantageous  stages  by  giving  advantage  to  indi- 
viduals which  tend  to  produce  the  organ  in  the 
most  degenerate  condition.^ 

^  See,  however,  Herbert  Spencer's  Social  atid  Moral  Problems,  as 
he  differs  from  Weismann  on  this  point. 

2  The  following  are  good  examples  of  the  operation  of  variation 
and  selection  in  producing  atrophy  in  a  species  : — 

(1)  Loss  of  constant  colour  ainong  domesticated  animals. — Wild 
animals,  especially  birds  and  mammals,  have  a  colour  which  is 
constant  for  a  whole  species.  Frequently  the  colour  is  protective 
in  rendering  the  animal  little  distinguishable  from  the  environ- 
ment in  which  it  lives.  As  soon  as  such  a  species  has  been 
domesticated  man  becomes  its  protector,  and  protective  colouration 
is  no  longer  necessary,  and  soon  disappears.  In  the  wild  state, 
the  colour  is  quite  as  variable  as  in  the  domesticated  state,  but 
the  abnormal  individuals  become  the  prey  of  enemies  and  are 
removed  from  the  species.  This  applies  not  only  to  animals 
which  are  preyed  on  by  others,  but  to  predatory  animals  them- 
selves. The  wild-cat,  for  instance,  will  have  less  difficulty  in 
stalking  its  prey  if  its  colour  makes  concealment  easy.     In  the 


THE  FACTORS  OF  ATROPHY  255 

2.  Sociology. — With  societies  this  elimination  of 
useless  structures  is  effected  much  more  easily  than 
with  organisms  for  several  reasons : — 

In  biology  a  special  factor,  heredity,  gives  to 
specific  characters  a  force  which  does  not  exist  in 
the  same  degree  with  social  institutions.  Now 
functional  organs  common  to  a  whole  line  of 
descent  are  not  easily  effected  by  the  influence 
of  individual  surroundings.  Further,  the  trans- 
domesticated  condition  man  provides  food  and  the  colour  being 
unimportant  all  variations  may  survive. 

(2)  Loss  of  spines  in  plants  on  oceanic  islands. — It  is  well  known 
that  the  presence  of  spines  protects  plants  from  the  ravages  of 
herbivorous  animals,  particularly  mammals.  But  in  oceanic  islands 
bats  are  generally  the  only  mammalian  inhabitants,  and  so,  accord- 
ing to  "Wallace  {Darwinism),  there  are  no  spiny  plants  in  the 
indigenous  flora  of  St  Helena.  The  much  richer  flora  of  the 
Hawaian  islands  includes  only  a  very  few  prickly  plants.  All 
the  endemic  genera  are  unarmed,  as  also  are  most  of  the  endemic 
species  of  other  genera ;  even  genera  like  Xanthoxi/lum,  Acacia^ 
Xylosoma,  Lycium,  and  Solanum,  which  are  so  frequently  armed 
in  other  countries,  are  there  represented  by  unarmed  species. 
The  two  species  of  Ruhus  bear  prickles  reduced  to  the  merest 
points  and  the  two  palms  are  devoid  of  spines.  How  is  the 
absence  of  spines  to  be  explained  in  these  plants  ?  The  plants 
have  been  derived  from  the  mainland,  the  seeds  being  brought  by 
the  wind,  by  currents,  or  by  birds,  and  having  found  soil  have 
germinated.  In  their  new  country  they  are  not  attacked  by 
herbivorous  animals,  and  it  is  immaterial  to  them  whether  or  no 
they  bear  spines.  The  individuals  badly  armed  are  at  no  dis- 
advantage compared  with  those  possessing  the  normal  armature  ; 
on  the  other  hand,  they  have  the  advantage  of  being  without 
useless  organs  to  support.  Spines,  in  consequence,  gradually 
disappear. 

This  struggle  for  existence  may  cause  the  disappearance  of  some 
organisms  themselves,  and  not  only  the  atrophy  of  parts  of  them. 


256    CAUSES  OF  DEGENERATIVE  EVOLUTION 

mission  of  acquired  characters  is,  to  say  the  least, 
doubtful.  There  is  no  proof  of  individual  atrophy- 
being  hereditary,  while  with  societies  modification 
may  be  transmitted  by  imitation.  Institutions 
which  have  fallen  into  disuse  rarely  recur  in 
freshly- formed  societies.  Natural  selection  plays 
an  all-important  part  in  biology,  but  it  is  artificial 
selection  which  almost  exclusively  governs  the  social 
domain. 

Many  vegetables,  as  for  instance  the  carrot  {Daucus  Carota)  are 
natives  of  France.  The  seeds  of  the  cultivated  carrot  must  fre- 
quently be  carried  to  waste  lands  or  uncultivated  soil.  The 
domesticated  variety,  however,  is  never  found  wild  although  the 
wild  variety  is  abundant.  This  vegetable  has  lost  the  power  of 
struggling  against  weeds ;  it  flourishes  only  when  it  is  protected 
by  man  and  when  by  repeated  weedings  its  wild  competitors  are 
removed.  When  it  is  returned  to  its  original  wild  haunts  the 
plant  dies  out  at  once. 

Most  cereals,  although  we  may  not  know  their  wild  ancestors, 
are  in  a  similar  condition.  For  instance,  if  man  were  to  cease 
cultivating  Wheat  {Triticum  sativum),  or  Rye  {Secede  cereale), 
there  is  no  doubt  but  that  these  would  completely  disappear. 
Their  fate  would  be  shared  in  Belgium  at  least  by  many  species 
which  are  reaped  with  them  at  harvest,  such  as  Centaurea  cyanus 
the  Corn-flower,  Agrostemma  Githago  the  Rose-Campion,  Specie- 
alaria  speculum,  and  others.  If  a  corn-field  were  left  to  the  free 
operation  of  nature,  weeds  would  soon  intrude  and  cause  the  dis- 
appearance of  the  plants  usually  present  in  it.  What  would 
happen  in  Belgium  would  happen  with  other  plants  in  other 
countries.  Thus,  near  Bergen  in  Norway,  some  plants,  such  as 
Melandryum  album,  Silcne  inflata,  Vicia  cracca,  etc.,  occur  only 
in  cultivated  fields.  In  Java,  many  aquatic  plants  such  as  Limno- 
charis  Plumieri,  LiLdwigia  perennis,  Jussiaea  suffruticosa^  etc., 
live  only  in  the  rice-fields  which  are  artificially  watered  and 
manured.  The  cessation  of  tillage  would  cause  the  disappearance 
of  all  these  plants  from  those  localities. 


THE  FACTORS  OF  ATROPHY         257 

An  institution  which  has  become  useless  and 
burdensome  is  generally  suppressed  before  its- 
complete  degenerative  evolution  is  accomplished. 
This  suppression  may  be  either  voluntary — as  in 
the  liquidation  of  a  commercial  company,  for 
instance — or  it  may  be  enforced.  By  the  terms 
of  Article  73  of  the  Belgian  Company's  Act, 
"  the  dissolution  must  be  declared  upon  the 
demand  of  all  those  interested  at  the  termina- 
tion of  six  months  from  the  time  when  the 
number  of  shareholders  has  been  reduced  to  less 
than  seven." 

The  downfall  is  generally  effected  in  this  sudden 
way,  either  voluntarily  by  the  interested  parties,  or 
by  the  intervention  of  legislative  means.^ 

Sometimes,  however,  artificial  selection  does  occur, 

^  The  occurrence  of  autotomy  or  self-mutilation  in  animals,  as 
in  crabs,  has  analogies  with  what  we  have  been  discussing. 
Similarly  some  plants  brought  into  a  new  locality  suddenly 
shed  their  leaves.  Ranunculus  aqucUilis,  cultivated  in  water, 
produces  long  divided  leaves  without  stomata  and  with  chloro- 
phyll in  the  epidermic  cells.  If,  from  some  chance,  the  water 
falls  below  the  level  of  the  plants,  the  adult  leaves  become  dry 
and  perish.  The  very  young  leaves  growing  unsubmerged  are 
still  divided,  but  to  a  lesser  extent  ;  they  have  stomata,  and  the 
epidermis  is  devoid  of  chlorophyll.  If  the  plants  be  again  sub- 
merged, this  form  of  leaf  dies,  and  there  is  a  new  development 
from  the  youngest  leaves  of  the  normal  aquatic  type. 

Other  plants  show  similar  occurrences.  Thus,  when  a  Fuchsia 
that  has  been  cultivated  in  a  conservatory  is  exposed  to  the  air, 
all  its  leaves  are  shed  and  are  replaced  by  new  leaves.  These  new 
leaves  again  fall  at  once  if  the  plant  be  brought  back  into  the 
conservatory.     This  is  a  real  case  of  autotomy  in  plants. 

B 


^ 


258    CAUSES  OF  DEGENERATIVE  EVOLUTION 

and  then  the  degenerative  evolution  of  useless  insti- 
tutions is  brought  about  in  a  similar  way  as  that 
of  non-functional  organs.  Atrophy  of  this  kind  may 
be,  as  in  biology,  either  accidental  or  normal.^ 

In  a  besieged  town  cut  off  from  all  outside  com- 
munication, all  train  service  is  necessarily  stopped 
and  the  railway  staff  rendered  useless.  On  the 
other  hand,  the  defence  of  the  city  requires  both 
men  and  money.  Under  these  circumstances  the 
resources  of  the  railway  naturally  come  to  be 
absorbed  in  the  service  of  the  defence. 

As  a  good  example  of  normal  atrophy  may  be 
mentioned    the   disappearance,   at   a   certain   point 

^  See  Durckheim  on  the  difference  between  normal  and  acci- 
dental sociological  phenomena  in  Les  Regies  de  la  m^thode 
sociologique. 

"All  sociological  phenomena,  like  biological  phenomena,  are 
liable,  while  remaining  the  same  individuals,  to  revert  to 
different  forms.  Now,  of  these  forms,  there  are  two  kinds  of 
reversion : — 

*  *  The  one  is  common  to  the  whole  species,  and  is  to  be  found, 
if  not  in  each  individual,  at  least  in  the  greater  part  of  them. 
The  cases  are  not  always  identical,  varying  slightly  with  the 
individual,  but  individual  variation  is  restricted  to  very  narrow 
limits. 

"The  other  kind  of  reversion  is  exceptional,  being  of  a  nature 
rarely  met  with,  and,  when  occurring,  is  seldom  permanent 
throughout  the  life  of  the  individual.  Cases  of  this  kind  are  excep- 
tional in  point  of  duration  as  in  other  respects. 

"Here  then  are  two  distinct  varieties  of  phenomena  which 
should  be  distinguished  from  one  another  by  different  terms. 
Individuals  exhibiting  only  common  characters  are  called  '  nor- 
mal, '  while  those  exhibiting  exceptional  characters  are  designated 
as  '  morbid '  or  '  pathological. ' " 


THE  FACTORS  OF  ATROPHY  259 

of  social  development,  of  the  popular  assemblies 
in  which  lay  the  origin  of  future  societies,  i.e. 
the  comitia,  assemblies  at  the  market-place,  the 
witenagemot,  May  Day  games,  rustic  assemblies, 
the  LaThdsgemeinde  of  the  Swiss  cantons,  the 
parochial  assemblies  of  the  Andora  Valley,  and  the 
town  meetings  of  New  England,  etc.  Some  of 
these  were  suppressed,  but  some  of  them  merely 
fell  into  disuse  as  other  institutions  arose  which 
were  better  adapted  to  more  modern  conditions  of 
society.  This  happened  with  regard  to  the  comitia 
curiaia  of  the  Eomans  during  the  period  of  the 
Empire,  which  were  gradually  supplanted  by  the 
comitia  centuriata  and  tribunal  comitia.  At  the 
time  when  they  are  first  mentioned  in  history,  they 
fulfilled  only  one  function :  that  of  ridding  the 
laws  of  all  traces  of  extrinsic  customs.^ 

To  sum  up  then,  it  is  plain  that  although  social 
degeneration  is  brought  about  by  the  same  general 
causes  as  organic  degeneration,  the  comparative 
importance  of  the  factors  in  degenerative  evolution 
is  far  from  being  identical  in  the  two  cases.  The 
autotomy  of  organs,  a  protective  self-mutilation, 
exhibits  only  a  far-fetched  analogy  with  the 
conscious  and  voluntary  suppression  of  social 
structures  which  have  become  either  useless  or 
prejudicial. 

Direct  individual  adaptation,  which  plays  a  part 

1  Mommsen,  Droit  public  romain.     Le  peuple  et  le  Sdnat,  vol.  i. , 
p.  364. 


260    CAUSES  OF  DEGENERATIVE  EVOLUTION 

of  only  secondary  importance  in  the  development  of 
animals  and  plants,  is  a  dominating  element  in 
sociology.  Institutions  are  able  to  modify  their 
structure  by  assimilating  new  inventions  and  im- 
provements, and  by  getting  rid  of  the  useless  parts. 
On  the  other  hand,  indirect  adaptation  spread  over 
a  species  plays  no  part  whatever  in  sociology,  for 
societies  seldom  reproduce  the  structures  of  the 
societies  from  which  they  sprang,  when  the  latter 
have  ceased  to  exist ;  whereas,  in  animals,  when  a 
useless  organ  is  reproduced  by  hereditary  repetition, 
variability  and  natural  selection  become  agents  in 
its  suppression. 


CHAPTER  I 

ATROPHY  OF  ORGANS 

Part  II 

Causes  producitig  atrophy 

The  ultimate  cause  of  the  atrophy  of  organs  is  the 
limitation  in  the  quantity  of  nourishment.  We 
have  shown  that  if  there  were  an  indefinite  supply 
of  food  there  would  be  no  struggle,  and,  as  a  result, 
no  degeneration.  We  have  now  to  examine  more 
minutely  the  course  of  atrophy  and  the  nature  of 
its  immediate  causes. 

The   atrophy    of    an    organ    is    a    reduction    in 


ATROPHY  OF  ORGANS  FROM  LACK  OF  SPACE    261 

structure,  in  nutrition,  and  in  functional  activity, 
but  the  succession  of  these  three  events  varies  with 
the  nature  of  the  exciting  cause. 

Eeduction  begins  with  structure  when  the  exciting 
cause  is  lack  of  space,  due,  for  instance,  to  increase 
in  another  organ  {atrophy  from  lack  of  space). 

Atrophy  begins  with  function  when  an  organ  has 
become  useless  (atrophy  from  lack  of  utility).  This 
uselessness  may  arise  from  two  causes  ;  the  function 
may  be  no  longer  useful  to  the  individual  or  to  the 
species,  or  it  may  be  assumed  by  another  organ. 

Lastly,  atrophy  may  begin  with  a  diminution  in 
the  supply  of  nutritive  materials  {atrophy  from  lack 
of  nutrition).  This  defective  nutrition  may  be  the 
result  of  a  general  cause  such  as  feebleness  of  the 
whole  organism,  or  it  may  be  due  to  the  hypertrophy 
of  another  organ. 

§  1.  Atrophy  from  lack  of  space. 

Cases  of  this  kind  are  rare. 

1.  Development  of  the  teeth. — Among  animals,  the 
development  of  the  teeth  furnishes  an  excellent 
example.  The  number  of  the  teeth  in  human 
beings  is  reduced  compared  with  the  number 
present  in  Lemurs  and  in  Platyrhine  Monkeys. 
These  have  six  grinding  teeth  while  in  man  five 
is  the  maximum  number.  Our  posterior  molar, 
however,  appears  late  in  life ;  it  is  smaller  than 
the  others,  so  that  it  may  be  useless  for  chewing ; 


I 


262 


CAUSES  OF  DEGENERATIVE  EVOLUTION 


frequently    it    is    absent, 


in    the    lower 
nineteen    per 


races 
cent. 


m 

of 


cases  examined  ;  in  the 
higher  races  in  forty-two 
per  cent. 

The  reduction  in  num- 
ber and  size  of  these 
teeth  is  due  to  a  re- 
duction in  size  of  the 
jaw-bones,  a  cause  which 
also  frequently  produces 
a  distortion  in  the 
arrangement  of  the  other 
teeth.  The  rudiments 
of  the  wisdom  teeth 
appear  on  the  maxillary 
tuberosity  and  on  the 
coronoid  process ;  it  is 
only  after  eruption  that 
they  come  into  normal 
connection  with  the  jaw- 

FiG.  76. — Tip  of  an  car  of  Lolium  perenne,    , 
with  two  earlets,  the  superior  bearing     DOnCS. 

two  glumes,  the  lateral  earlet  with  ^         .  ,  -,  n    ±i 

only    the  superior  glume.       In   the  I.    AtVOpliy     OJ     t llC 

lateral  earlet  the  lower  flower  is  open  .  ,  . 

and  has  two  glumules  ;  all  the  other    SUperiOV  gtumc. AUlOUg 

flowers  are  closed  and  exhibit  only  the       i        ,      ,  i  c 

inferior  giumuie.  plauts  thcrc  are  tew  cases 

GI,  inferior  glume;  GS,  superior  glume;        p      i  i  li.      r 

gi,    inferior    glumule;     gs,    superior    01  atropliy  aS  a  rCSUlt  01 
glumule.  1      1       £  T 

lack  of  space,  in  grasses 
of  the  genus  Lolium,  the  earlets  are  arranged  in  a 
spike,  but  in  such  a  fashion  that  only  the  terminal 
earlet  has  space  for   both  glumes  (fig.   76).     The 


ATROPHY  OF  ORGANS  FROM  LACK  OF  USE   263 

superior  glumes  remain  ;  the  inferior,  pressed  against 
the  axis,  disappear  after  the  embryonic  develop- 
ment of  the  flower. 

3.  Degeneration  of  jpalece  and  of  stamens. — Lack 
of  space  is  probably  the  immediate  cause  of  the 
disappearance  of  palese  in  the  receptacles  of  some 
composite  flowers  and  of  the  posterior  stamen  in 
the  flowers  of  some  Scrophulariacese  and  Labiates. 

In  normal  racemose  inflorescences  each  floret 
grows  in  the  axil  of  a  reduced  leaf  called  a 
bract.  When  the  axis  of  the  inflorescence  is 
shortened  and  the  florets  crowded,  as  in  the 
capitula  of  composite  flowers,  it  frequently  hap- 
pens that  the  bracts  of  the  florets  (termed 
palese)  disappear.  This  absence  is  most  usual 
when  the  capitulum  is  small  and  the  florets  are 
large. 

In  Labiates  and  most  Scrophulariacese,  although 
the  ancestral  number  of  stamens  was  five,  there  are 
not  more  than  four  present ;  when  only  one  is 
absent,  it  is  the  original  posterior  stamen  which 
was  pressed  against  the  axis  of  the  inflorescence. 

§  2.  Atrophy  from  lack  of  use. 

1.  Functional  Inutility. 

(1)  Etiolated  plants  and  immobile  limhs. — We 
have  already  quoted  as  instances  of  accidental 
atrophy,  cases   of   degeneration   of   leaves  in  etio- 


264    CAUSES  OF  DEGENERATIVE  EVOLUTION 


lated  plants,  and  of  muscles  in  unused  limbs  (see 
fig.  75,  p.  253). 


Fig  .77. — Nymphaea  alba  sown  on  tlie  mud  and  at  different  depths  in  it.     1,  2,  3,  successive 
stages  of  the  same  seedling.    Eau,  water;  limon,  mud. 


•  ATROPHY  OF  ORGANS  FROM  LACK  OF  USE   265 

(2)  Epicotyl  and  frimary  leaf  of  Nymfhaea. — 
These  structures  in  the  water-lily  are  good 
examples  of  normal  atrophy  in  individuals.  Dur- 
ing germination  (fig.  77)  the  cotyledons  of  the 
water-lily  remain  inside  the  seed,  and  a  new 
organ  (at  right  angles  to  them)  grows  vertically 
upwards.  The  lower  part  of  this  is  the  first 
internode  of  the  stem  (epicotyl),  and  the  upper 
part  is  a  primary  acicular  leaf.  It  grows  up- 
wards through  the  mud  until  the  summit  of  the 
leaf  reaches  light.  The  growth  of  the  epicotyl  is 
then  much  slower,  and  its  terminal  bud  begins  to 
shoot  out  horizontally.  The  use  of  this  growth  of 
the  epicotyl  and  primary  leaf  is  to  carry  the  bud 
to  the  light.  When  that  purpose  is  achieved,  these 
structures  atrophy.  A  similar  occurrence  may  be 
found  in  Sagittaria  sagittifolia  (fig.  40,  H,  I,  p.  72). 
In  that  case,  however,  it  is  the  hypocotyl  which  elon- 
gates, until  light  is  reached,  and  then  degenerates. 

(3)  Roots  of  Utricular ia,  cotyledons  of  imrasitic 
plants,  leaves  transformed  to  spines  in  Phyllocactus 
crenatus. — As  instances  of  atrophy  throughout  a 
species  produced  by  inutility  of  the  parts  con- 
cerned, we  have  already  mentioned  the  roots  of 
Utricularia  and  the  cotyledons  of  the  parasitic 
plants   Cuscuta,   Orobanche,  etc. 

The    spines    of    Phyllocactus    crenatus    produced  . 
from  modified  leaves  are  another  example  (fig.  78). 
Above   the   rounded   base   by   which  a   branch   of 
Phyllocactus  is   attached   to   older   branches,  there 


266 


CAUSES  OF  DEGENERATIVE  EVOLUTION 


\ 


/ 


/ 


\ 


// 


is  an  angular  region,  the  sides  of  which  are 
prominent,  and  bear  leaves  modified  into  spines 
to  serve  as  protecting  organs,  as  in  the  similar 
case  of  Cereiis.  Higher  up  the  branch  the  pro- 
minent sides  become  flat,  and  the 
spines  are  replaced  by  minute 
scales.  This  degeneration  is  the 
result  of  loss  of  utility.  The 
Phyllocacti  are  epiphytes,  and 
their  situation  consequently  places 
them  out  of  the  reach  of  crop- 
ping animals.  The  spines  near 
the  basis  of  the  branches  are  a 
survival  from  the  terrestrial  an- 
cestors of  Phyllocdctus.  The 
spines  higher  up  have  degenerated. 
Animals  offer  many  instances 
of  atrophy  as  a  result  of  inutility, 
both  in  individuals  and  in  species. 
(4)  Atrophy  of  the  h^anchial 
arches  in  mammals. — As  they  are 
no  longer  functional,  most  of 
the  mammalian  branchial  arches 
atrophy.  Three  pairs  alone  persist, 
and  of  these  it  is  only  those  parts  which  are  useful. 
(5)  Atrophy  of  ventral  fins. — Instances  of  atrophy 
through  uselessness  in  species  are  to  be  found  in 
the  ventral  fins  of  fishes  like  the  Pediculati,  which 
live  in  the  mud,  or  in  Frotopterus,  which  for  a  part 
of  the  year  is  completely  buried  in  mud  (fig.  1 9,  p.  44). 


Fig.  78.— Branch  of 
Phyllocactus  crenatus. 


ATROPHY  OF  ORGANS  FROM  LACK  OF  USE       267 

(6)  Atrophy  of  mvsclcs. — Cessation  of  use  is  also 
the  cause  of  the  degeneration  of  the  flexor  and 
extensor  muscles  of  the  fore-limb  in  Cetacea,  and 
of  the  imperfection  of  the  finger  joint  articulations 
in  Cetacea  and  Sirenia  (fig.  79).  In  the  latter 
cases  the  surfaces  of  the  articu- 
lation which  make  flexor  and 
extensor  movements  easy  dis- 
appear. When  a  limb  becomes 
a  paddle,  it  is  necessary  that  it 
should  be  flexible,  but  that  the 
articulations  should  be  immobile. 

(7)  Atrophy  of  the  tail  in 
man. — The  caudal  region  of  the 
human  vertical  column  is  com- 
posed of  four  or  five  very  de- 
generate vertebrae.  The  whole 
of  this  organ  is  degenerate. 
When  the  tail  is  formed  at  an 
early    stage    of    embryonic     life,  fig.  79— aioMocephaius 

,1  J.   1        1  1  '   1.         £       melas.     Right  anterior 

the    vertebral    column     consists    01       An  showing  absence  of 
,,.,  •!,  ,1  ,1        1  articular  facets  for  the 

thirty-eight  vertebrae ;  the  lesser  joints  of  the  fingers. 
number  of  vertebrae  in  the  adult 
is  due  to  reduction  of  the  tail,  which  in  man  is 
quite  useless.  Later  on  in  life  a  further  instance 
of  atrophy  may  occur  in  individuals.  In  old  men 
the  caudal  vertebrae  are  frequently  fused,  and  the 
whole  region  is  smaller. 

(8)  Degeneration  of  the  hyoid  appear atus  in  man 
and    birds. — This    case    shows    a   close  correspon- 


/ 


268         CAUSES  OF  DEGENERATIVE  EVOLUTION 

dence  between  atrophy  and  loss  of  function.  The 
second  arch  becomes  connected  with  the  third,  the 
parts  of  which  are,  in  the  adult,  the  styloid  pro- 
cess, the  stylo-hyoidean  ligament,  and  the  lesser 
horn  of  the  hyoid  bone.  According  to  the  weight 
of  the  tongue,  the  parts  of  the  second  arch  become 
more  or  less  developed.  In  man  the  suspensory 
apparatus  of  the  hyoid  bone  is  extremely  simple, 
and  it  is  still  more  reduced  in  birds.  "  The 
tongue,  reduced  to  a  minute  cartilage,  no  longer 
requires  the  support  of  a  bony  base  so  that  the 
hyoidean  apparatus  might  almost  be  removed  from 
the  anatomy  of  a  bird.  It  is  present,  but  in  a 
rudimentary  condition "  (Geoffroy-Saint-Hilaire). 

In  the  horse,  which  has  a  heavy  tongue,  the 
second  arch  is  strong  and  completely  bony. 

In  fish  the  hyoidean  system  is  still  more  im- 
portant, although  in  them  this  is  associated  not 
with  any  importance  of  the  tongue,  but  with  the 
branchial  apparatus.  The  parts  of  the  second  arch 
are  very  strong,  as  they  form  a  fulcrum  against 
which  the  branchial  system  works,  but  its  main 
parts  are  recognizable.  "  The  hyoidean  apparatus 
is  the  same  in  all  vertebrates ;  its  functions  are  at 
a  maximum  in  fish,  and  at  a  minimum  in  birds, 
while  in  mammals  the  condition  is  intermediate  " 
(Geofifroy-Saint-Hilaire). 

II.  Transference  of  function. — Among  plants 
and  animals  there  are  many  instances  of  organs  well 
developed    in    younger   stages    of    life,  but   which 


ATROPHY  OF  ORGANS  FROM  TRANSFERENCE      269 

become  rudimentary  in  later  stages  on  account  of 
their  functions  being  assumed  by  other  organs. 

(1)  Atrophy  of  the  tail  in  Batrachia  Anura,  and 
of  the  larval  gills  in  some  insects. — The  tadpole  of 
the  frog  has  a  well-developed  tail  which  acts  as 
the  organ  of  locomotion ;  the  adult  animal  moves 
by  its  limbs,  and  the  tail,  useless  in  the  adult,  has 
been  removed  by  phagocytosis. 

The  aquatic  larv£e  of  many  terrestrial  insects 
possess  tracheal  gills,  that  i^  to  say,  membranous 
expansions  of  the  skin,  within  which  tracheae 
ramify.  In  the  adult  insect  respiration  is  con- 
ducted by  normal  tracheae  communicating  with  the 
air,  and  the  larval  organs  of  respiration  atrophy. 

In  addition  to  such  cases  of  atrophy  occurring 
normally  in  the  life-history  of  individuals,  there 
are  known  many  cases  where  the  organs  of  a 
species  have  disappeared  on  account  of  the  trans- 
ference of  their  functions. 

(2)  Disappearance  of  limbs. — Vertebrates  which 
move  by  general  undulations  of  the  body  have 
lost  their  limbs  for  this  reason.  Such  cases  are 
Slow-worms,  Amphisbaena,  Snakes,  Eels  and  Cse- 
cilians. 

Many  parasitic  creatures  have  similarly  lost  their 
organs  of  progression,  as  they  depend  upon  their 
host  for  movement  from  place  to  place. 

Sacculina,  a  parasite  on  the  carapace  of  crabs, 
has  completely  lost  its  organs  of  locomotion.  More- 
over, as  it  takes  its  food  by  processes  passing  into 


270    CAUSES  OF  DEGENERATIVE  EVOLUTION 

the  tissues  of  the  crab,  its  digestive  canal  has  been 
lost. 

(3)  Atrophy  of  the  leaf. — In  many  plants  the 
leaves  have  disappeared,  their  function  having 
been  assumed  by  some  other  part  of  the  plant, 
as,  for  instance,  by  the  phyllodes  in  Acacia  and 
in  Phyllanthus  (fig.  84). 

(4)  Atrophy  of  the  protonema  in  mosses,  and  of 
the  leaves  in  some  xerophilous  plants. — At  germina- 
tion, mosses  produce  a  much  branched  filamentous 
structure  which  serves  as  the  organ  of  nutrition, 
and  is  termed  the  protonema.  Later  on  this  gives 
rise  to  buds  which  develop  into  the  normal  leafy 
shoots  of  the  plants.  As  soon  as  the  leaves  are 
large  enough  to  manufacture  food  for  the  plant,  the 
protonema  begins  to  degenerate,  and  disappears 
completely,  except  in  a  few  rare  forms  (Ephe- 
meracese)  where  the  leafy  shoots  remain  very 
small. 

In  Muehlenhechia  platy dados  (fig.  80),  which  has 
become  adapted  to  arid  regions,  and  in  consequence 
has  the  surface  from  which  evaporation  may  take 
place  much  reduced,  the  branches  which  do  not 
bear  assimilating  leaves  are  rounded  at  their  bases, 
but  higher  up  flatten  into  broad  blades.  These 
blades  contain  chlorophyll  and  fulfil  the  functions 
of  leaves,  these  latter  being  present  only  as  minute 
scales. 

Similar  phenomena  occur  in  many  Papilion- 
aceous   plants    belonging    to    the    genera    Genista 


ATROPHY  OF  ORGANS  FROM  TRANSFERENCE     271 


(broom),   Spartium,   Alhagi,   etc.     The  leaves  dis- 
appear, and  the  stems  assume  their  functions. 

(5)  The  reduction  or  disa/ppearance  of  the  calyx. 
— The  disappearance  of  the  calyx  in  many  com- 
posite flowers  and  of  roots  in 
epiphytes  are  instances  of  species 
losing  organs  because  of  the 
transference  of  function  to  other 
organs. 

In     the    simplest    Compositse, 
the   fruits  (achenes)  are    dissemi- 
nated   by    the    wind,    the    calyx 
usually    being    modified     into    a 
feathery    tuft.       In    other    Com- 
positse,    although    wind    dispersal 
still   occurs,    the    feathery    calyx 
is  lost,  its  purpose  being  fulfilled 
by  wings  on  the  sides  of  the  fruit 
(Florestinia  pedata),  or  it  is  united 
to     a     large     membranous     scale 
developed     from     the     receptacle 
(Dahlia),   or    the     sterile     florets 
may  be  turned  into  wings  (Lind-  fig- 
heimeri/i  texana).     In  other  cases 
the  fruits  are  dispersed,  not  by  the 
animals    which  eat  them  (Clibadium 
they  adhere  to  the  fur  of  animals  by 
on   the   achenes   (Calendula),  or  by 
from  the  involucre  of  bracts  (Zappa). 
cases,  and  the  list  might  have  been 


80.— Branch  oiMueh- 
nbeckia  platyclados. 

wind,  but  by 

asperum),   or 

hooks  formed 

hooks  formed 

In  all  these 

made  longer, 


272         CAUSES  OF  DEGENERATIVE  EVOLUTION 

the  function  of  the  calyx  has  been  reduced,  or  has 
disappeared  on  account  of  the  transference  of  its 
function. 

(6)  Atrophy  of  roots. — Eoots  in  most  plants  per- 
form two  functions  :  they  fix  the  plant  in  the  soil, 
and,  chiefly  by  means  of  the  delicate  hairs  on  their 
youngest  parts,-  they  absorb  water  and  dissolved 
mineral  substances.  Sometimes,  however,  instead 
of  having  root-hairs,  the  rootlets  enter  into  a 
kind  of  partnership  with  a  fungus,  which  lives 
in  their  tissues,  and  absorbs  by  its  processes  the 
necessary  food  materials  from  the  soil.  The  pine 
and  beech  are  examples  of  this. 

In  some  orchids  {Corallorhyza^  Myrmechis)  the 
fungi  lodge  in  the  subterranean  part  of  the  plant, 
and  the  branches,  having  no  function,  disappear. 

In  most  of  the  epiphytic  Bromeliacese  the  roots 
are  useful  only  to  anchor  the  plants ;  the  absorption 
of  water  takes  place  entirely  through  new  organs 
developed  from  the  leaves.  The  roots  are  few  in 
number  and  small,  and  after  the  plant  has  obtained 
a  firm  position  their  growth  almost  ceases.  In 
another  species  (Tillandsia  icsneo'ides)  of  the  same 
family  the  roots  have  completely  disappeared.-^ 
This  plant,  called  by  the  Brazilians,  "  the  plant  of 
the  air,"  fixes  itself  to  branches  of  trees  by  its 
leaves,  and  is  easily  transported  by  the  wind.     Its 

1  For  further  details  concerning  the  Bromeliacese  see  Schimper, 
Die  epiphytische  Vegetation  Amerikas,  in  Schimper^ s  Botanische 
MittheiluTigen  aus  den  Tropen.     Jena,  1891. 


ATKOPHY  OF  ORGANS  FROM  TRANSFERENCE      273 

absorption   takes  place  entirely  through  hairs  de- 
veloped upon  the  leaves. 

In  addition  to  such  examples  of  plants  which 
have  lost  their  roots  on  account  of  the  functions  of 
the  roots  being  assumed  by  other  organs,  there  are 
also  plants  in  which  the  roots  represent  the  sole 


Fig.  Sl.—Tceniophyllum  Zollingeri  with  branches  adhering  to  bark.    The  plant 
bears  a  flower  and  a  bud. 

vegetative  organs.  In  Tmniophyllium  Zollingeri 
(fig.  81),  an  epiphytic  orchid,  the  leaves  are  reduced 
to  minute  scales  and  are  of  no  importance  in  the 
nutrition  of  the  plant,  that  function  being  trans- 
ferred to  the  roots  which  in  the  form  of  flat  green 
ribbons,  apply  themselves  closely  to  the  bark  of 
trees.      In  this  case  the  roots  have  assumed  the 

s 


/ 


274    CAUSES  OF  DEGENERATIVE  EVOLUTION 

functions  of    leaves,  while  in   Tillandsia  usneoldes 
the  leaves  have  taken  on  the  functions  of  roots.    . 

The  Podostemaceae  are  aquatic  plants  which  live 
in  warm  regions  attached  to  rocks  in  cascades. 
The  stems  and  leaves  are  completely  absent,  the 
flowers  even  being  produced  on  the  roots.  Some 
of  the  roots  become  closely  attached  to  the  stones. 
Others  which  are  green  and  ribbon-like,  float  in  the 
stream,  and  serve  for  assimilation. 

§  3.  Atrophy  from  lack  of  nutrition. 

In  some  cases  of  degeneration,  the  organs  do  not 
lose  their  functions,  but  become  reduced,  merely 
because  adjacent  parts  rob  them  of  their  nourish- 
ment. This  kind  of  lack  of  nutrition  which  results 
in  local  atrophy,  must  be  distinguished  from  the 
general  limitation  of  food-supply  which  is  the 
ultimate  cause  of  all  degeneration.  The  possible 
amount  of  food  within  the  reach  of  any  organism 
is  limited,  but  besides  this,  the  share  of  the  absorbed 
food  obtained  by  any  particular  organ  or  part  of 
an  organ  may  be  limited  with  a  resulting  atrophy 
of  that  organ  or  part.  Atrophy  of  this  kind  may 
be  accidental  or  normal. 

1.  Parasitic  castration. — A  good  instance  of  acci- 
dental atrophy  of  this  kind  is  found  in  Melandryum 
album  where  the  ravages  of  a  fungus  Ustilago 
antheridarum    may    produce    parasitic    castration.^ 

^  See  Giard,  La  Castration  parasitaire  in  the  Bull,  Sclent,  de  la 
France  et  de  la  Belgique.     1887,  1888,  1889. 


ATROPHY  OF  ORGANS  FROM  LACK  OF  NUTRITION    275 

Under  the  influence  of  this  parasitic  fungus  which 
makes  its  way  to  the  anthers,  the  stamens  of  the 
female  flowers  assume  the  form  of  those  in  male 
flowers  and  in  consequence  the  pistils  of  these 
flowers  abort  from  defect  in  nutrition. 

2.  Severe  or  prolonged  compression  of  a  limh. — A 
long  continued  or  forcible  compression  of  a  limb 
results  in  atrophy  of  its  extremity  on  account  of 
lack  of  nutrition  from  the  compression  of  the  blood- 
vessels. 

3.  Atrophy  of  the  genitalia  in  neuter  bees. — Lack 
of  nutrition  is  also  the  cause  of  the  arrested  de- 
velopment of  the  genitalia  normal  in  neuter  bees. 
The  neuters  of  bees  and  of  some  of  their  allies  are 
females  in  a  state  of  arrested  development.  In 
wasps,  humble-bees,  and  hive-bees,  it  sometimes 
occurs  that  the  genitalia  of  these  forms  develop 
sufficiently  to  be  functional,  thus  resulting  in  the 
appearance  of  small  females.  In  most  honeycombs 
two  kinds  of  cells  are  formed :  in  the  smaller  and 
more  numerous  cells  are  placed  the  larvae  destined 
to  become  neuters,  in  the  larger  and  less  numerous 
those  destined  to  become  queens  or  perfect  females. 
The  food  of  the  two  sets  of  larvae  is  different ; 
those  in  the  larger  cells  are  given  "  royal  food  "  a 
more  nutritious  substance.  When  some  of  the 
royal  food  by  an  accident  gets  into  a  worker  cell 
the  sexual  organs  of  that  larva  are  developed  so 
that  a  small  female  is  formed.  In  this  way  as 
many  females  as  may  be  desired  can  be  produced, 


276         CAUSES  OF  DEGENERATIVE  EVOLUTION 

and  when  a  hive   has  lost   its  queen,  the  worker 
bees  produce  another i  (Lacordaire). 

Plants  afford  many  instances  of  degeneration  due 
to  defect  of  nutrition. 

4.  Atrophy  of  tJie  superior  flowers  in  Car  ex. — In 
the  tall  spikes  of  Carex,  it  frequently  happens  that 
the  flowers  towards  the  summit  are  rudimentary, 
and  authorities  are  agreed  in  regarding  this  condi- 
tion as  the  result  of  defect  in  nutrition. 

5.  Atrophy  of  pistils  and  stamens. — Cases  of 
atrophy  of  the  stamens  or  pistils  normal  in  species 
may  be  given. 

In  Fritillaria  persica  the  flowers  are  disposed  in 
bunches.  The  lower  flowers  possess  six  perianth 
members,  two  cycles  of  three  stamens  each,  and  a 
pistil.  In  the  median  flowers  the  pistil  is  smaller, 
and  rarely  capable  of  being  fertilized.  In  the 
superior  flowers  the  degeneration  is  complete,  the 
pistil   hardly  being   formed.     It   might   be   shown 

^  Tlie  transference  of  a  function  is  not  invariably  accompanied 
by  degeneration.  Thus,  in  the  functional  development  of  an 
individual's  nervous  system  voluntary  acts  which  have  been  re- 
peated frequently  become  reflex  actions,  and  have  their  seat  in  a 
different  region  of  the  nervous  system — as,  for  instance,  walking 
and  acquired  professional  dexterities.  It  has  been  sought  to  ex- 
plain the  development  of  the  instincts  of  species  in  this  Avay  by 
supposing  that  frequently  repeated  voluntary  acts  have  become 
inherited  reflexes.  As  pathological  degeneration  in  man  frequently 
affects  the  higher  regions  of  the  brain,  reflexes  and  instincts  may 
persist  after  loss  of  voluntary  action.  In  the  cases  of  transference 
of  nervous  functions  to  lower  centres  the  higher  centres  do  not 
degenerate  but  remain  able  to  acquire  new  voluntary  functions. 


ATROPHY  OF  ORGANS  FROM  LACK  OF  NUTRITION    277 


that  this  degeneration  is  due  to  lack  of  nutrition, 
by  removing  the  inferior  flowers  from  a  young 
bunch,  but  we  are  unaware  that  the  experiment  has 
been  made. 

In  Viburnum  tomentosum  (fig.  82)  the  flowers 
are  arranged  in  a  compound  corymb  and  occur  in 
two  forms.  The  central  flowers  have  a  small 
corolla,  five   stamens,  and  a   well-developed   pistil 


Fig.  82.— Flowers  from  the  same  inflorescence  of  Viburnum  tometitosum. 
A,  sterile  flower  witli  rudimentary  pistil  and  no  stamens. — B,  sterile  flower  with 
very  rudimentary  pistil  and  stamens.— C,  flower  witli  two  fertile  stamens;  the 
other  three  stamens  and  the  pistil  are  rudimentary. — D,  fertile  hermaphrodite 
flower. 

(fig.  82,  d).  The  eight  or  nine  peripheral  flowers 
of  each  inflorescence  have  the  corolla  much  larger, 
but  the  stamens  are  absent  and  the  pistil  is  rudi- 
mentary (fig.  82,  a).  The  petals  turned  towards 
the  outer  side  are  much  the  largest,  and  it  is  not 
rare  for  the  petals  turned  towards  the  inner  side  to 
be  hardly  larger  than  those  of  the  central  flowers. 
What  is  important  to  notice,  as  it  bears  out  the 
relation  between  degeneration  and  lack  of  nourish- 


278  CAUSES  OF  DEGENERATIVE  EVOLUTION 

ment,  is  that  a  smaller  petal  is  always  associated 
with  a  larger  stamen  (fig.  82,  b,  c). 

In  Vihurnitm  o^pulus,  the  wild  guelder-rose,  there 
are  also  two  kinds  of  flowers,  the  central  flowers 
which  are  sexual  and  hermaphrodite,  and  the 
peripheral  flowers  which  are  sterile  and  possess  a 
very  large  corolla.  The  five  petals  are,  however, 
equal  in  size,  and  there  are  no  forms  transitional 
between  the  sterile  and  sexual  flowers.  The 
cultivated  guelder-rose  is  a  variety  of  the  wild 
form  in  which  all  the  flowers  are  sterile  and 
possess  a  large  corolla  :  the  plant  has  completely 
lost  the  power  of  sexual  reproduction. 

§  4.  Atrophy  without  apparent  cause. 

In  many  cases  it  is  impossible  to  determine  the 
cause  of  atrophy.  Why,  for  instance,  has  half  the 
inflorescence  disappeared  from  a  unilateral  cyme  ? 
Why  have  some  composites  like  Artemisia  or 
Eosaceae  like  Poterium  lost  their  habit  of  being 
fertilized  by  the  agency  of  insects,  and  become 
anemophilous  without  conspicuous  perianth  ?  Nor 
can  we  explain  why  many  Myriapods  are  blind,  al- 
though they  live  in  association  with  species  possess- 
ing eyes  and  in  conditions  where  vision  apparently 
would  be  useful.  Semper  discovered,  in  brackish 
water  in  the  Philippines,  Crustacea  (Gymothoe) 
which  were  completely  blind,  although  they  lived 
in  light.     The  cause  of  such  atrophies  is  unknown. 


ATKOPHY  OF  ORGANS  WITHOUT  APPARENT  CAUSE  279 

Moreover,  we  know  that  organs  may  atrophy- 
through  correlation  with  other  degenerating  organs, 
or  because  the  whole  organism  is  degenerating 
although  there  are  no  obvious  anatomical  bonds 
present  between  the  related  parts.  As  instances  of 
such  atrophies  we  may  mention  degeneration  of  the 
brain  resulting  on  degeneration  of  the  supra-renal 
capsules ;  or  of  the  condition  of  myxodoema 
resulting  from  mechanical  or  functional  disturbance 
of  the  thyroid  gland.  When  the  essential  sexual 
organs  of  the  male  or  female  are  congenitally  or 
accidentally  absent,  the  secondary  sexual  characters 
— the  beard,  voice,  hair,  and  whole  male  or  female 
aspect — of  the  body  degenerate.  When  one  eye 
becomes  injured  or  diseased,  a  frequent  consequence 
is  sympathetic  degeneration  of  the  undamaged  eye. 

To  these  cases  of  correlative  atrophy  we  may 
add  the  cases  of  leaves  on  many  plants  with 
drooping  or  horizontal  branches.  On  these,  the 
leaves  on  the  upper  aspect  are  frequently  rudi- 
mentary. The  appearance  is  most  striking  where 
the  leaves  are  opposite  the  pairs  being  at  right 
angles  to  one  another.  In  a  branch  of  Acer 
campestre  which  is  upright,  all  the  leaves  are  of  the 
same  size.  In  a  drooping  branch  the  leaves  turned 
towards  the  sky  are  small.  In  plants  belonging  to 
very  different  families  (Acanthaceae,  Melastomacese 
Urticacese,  etc.)  oblique  branches  exist,  and  the 
failure  in  development  of  the  dorsal  upper  leaves  is 
invariable.     In  Procris  laevigata  (fig.  83)  the  reduc^ 


I 


280 


CAUSES  OF  DEGENERATIVE  EVOLUTION 


tion    of    these   leaves  has   gone   very   far,   but   in 
Klugia  notoniana  the  leaves  in  the  corresponding 


Fig.  83.— Branch  of  Procris  laevigata. 

F'/',  F"/")  F'"/"'.     Tlie  successive  pairs  of  leaves  in  the  axil  of  a  large  leaf  F' 

is  a  female  inflorescence ;  in  the  axil  of  a  small  leaf  /"  is  a  male  inflorescence. 

(The  original  drawing  was  made  at  Buitenzorg  in  Java  by  Mas  Kromohardjo,  a 

Malay  draughtsman.) 


position  are  absent.  In  another  representative  of 
the  same  family  (Strejptocmym  monophylleia),  the 
degeneration  has  gone  still  further ;  all  the  leaves 


ATROPHY  OF  INSTITUTIONS  281 

are  absent  and  a  single  greatly  enlarged  cotyledon 
is  the  sole  assimilating  organ/ 

It  is  evident  that  in  these  oblique  and  horizontal 
branches,  the  leaves  directed  vertically  towards  the 
sky  are  in  the  most  unfavourable  position  for  assi- 
milation, and  that,  in  addition,  their  presence  would 
shade  the  leaves  lying  under  them.  In  these 
species  the  more  or  less  complete  reduction  is  an 
inherited  fixation  of  a  sacrifice  of  these  particular 
leaves  for  the  benefit  of  the  whole  organism. 


CHAPTEE    II 

ATROPHY    OF    INSTITUTIONS 

The  causes  of  atrophy  in  institutions  are  more  or 
less  analogous  to  those  which  bring  about  atrophy 
of  organs.  First,  there  is  atrophy  from  ivant  of 
use,  when  function  either  becomes  useless  or  is 
transferred  to  another  institution.  Atrophy  from 
lack  of  resources  corresponds  precisely  with 
atrophy  from  lack  of  nutrition.  There  is  no- 
thing, however,  in  the  atrophy  of  institutions  quite 
analogous  to  that  which,  in  organisms,  results  from 
lack  of  space.  But  if  the  development  of  an 
institution  cannot  be  actually  impeded  by  the  co- 

1  See  the  figures  of  plants  of  this  family  in  Fritsch,  Gesneriacece. 
{Engler  und  PrantVs  natiirliclien  Pflanzenfamilie^i.    Leipzig,  1891. ) 


282         CAUSES  OF  DEGENERATIVE  EVOLUTION 

existence  of  another  institution,  certain  instances 
of  atrophy  do  occur  which  "cannot  be  said  to  arise 
from  either  want  of  use  or  lack  of  resources.  The 
institution  degenerates  merely  from  hindrance 
offered  to  the  exercise  of  its  functions.  Thus,  for 
instance,  the  laws  of  exclusion  voted  in  Germany 
against  socialists  brought  about  the  decadence,  and, 
ultimately,  the  downfall  of  a  number  of  professional 
and  trade  associations  which  cannot  be  said  to  have 
been  actually  suppressed  by  law.  It  is  the  same 
with  the  corporations  of  Western  Flanders ;  these 
have  survived  the  revolutionary  laws,  but  are  no 
longer  legally  recognized,  a  state  of  things  leading 
to  many  difficulties  ;  the  properties  of  several  of 
them  have  been  confiscated,  as  having  no  owners, 
and  allotted  to  benevolent  institutions.  This  want 
of  legal  recognition  must  inevitably  lead  in  the 
near  future  to  the  entire  disappearance  of  these 
vestiges  of  the  corporative  system. 

Cases  such  as  these,  however,  must  be  regarded 
as  exceptional,  and  it  may  be  said  that,  on  the 
broad  average,  institutions  atrophy  either  from 
want  of  use,  or  from  lack  of  resources. 

§  1 .  Atrophy  from  want  of  use. 
I.  Functional  inutility. 

(1)  Offices  in  connection  with  the  Fort  of  BrugeSf 
— The  disorganization  of  the  train  service  in"  a 
besieged     city    has    already    been    alluded    to    as 


ATROPHY  OF  INSTITUTIONS  FROM  LACK  OF  USE  283 

furnishing  an  example  of  degeneration  by  sus- 
pension of  function. 

Another  incidence  of  accidental  degeneration  is 
furnished  by  the  office  of  wharf -porter,  which  was 
formerly  exercised  in  the  port  of  Bruges.  When 
Bruges,  consequent  upon  the  blocking  up  of  the 
Zwyn,  ceased  to  be  a  sea-port  town,  the  wharf- 
porters  who  were  formerly  employed  to  carry 
grain,  lime,  and  coal,  etc.,  were  no  longer  required 
and  so  abandoned  their  calling. 

(2)  The  forest-courts  of  England. — As  an  instance 
of  normal  degeneration  due  to  the  transformation 
of  an  agricultural  country  into  a  commercial  country, 
take  the  old  English  forest-courts.  In  the  middle 
ages  there  still  existed  in  England  great  tracts  of 
forest  land  which  were  Crown  property,  and  sub- 
jected to  special  legislation  conducted  by  three 
separate  courts  of  justice  :  (a)  The  Court  of  Attach- 
ment ^  which  instituted  proceedings  ;  (Jb)  the  Court 
of  Swainmote  ^  before  which  the  culprits  were  tried, 
and  the  Judge's  Court  presided  over  by  the  Lord 
Chief  Justice,  who  pronounced  sentence,  and  from 
whose  decision  there  was  no  appeal.  These  courts 
have  lost  all  importance  since  the  seventeenth 
century,  and  the  forest  laws  are  now  only  functional 

^  When  a  forest  law  was  infriuged,  it  was  the  duty  of  the  forester 
to  "attach"  the  culprit — i.e.  constrain  him  to  appear  either  by 
seizing  his  person  or  his  goods.  These  attachments  were  then 
submitted  to  the  Court  of  Attachment. 

'■^  The  judges  were  called  verderers ;  the  jury  was  composed  of 
foresters  of  the  reeve,  and  of  four  men  out  of  each  forest  hundred. 


284    CAUSES  OF  DEGENERATIVE  EVOLUTION 

with  regard  to  the  Forest  of  Dean  and  the  New 
Forest.  The  office  of  Forest  Judge  has  disappeared, 
and  but  few  vestiges  remain  of  the  courts  of  attach- 
ment and  Swainmote.  Those  persons  who  are  con- 
nected with  either  the  Forest  of  Dean  or  the  New 
Forest,  meet  together  still  to  transact  business  in 
connection  with  them,  but  these  meetings  are  of  no 
great  importance. 

II.  Teansference  of  function. 

Instances  abound  of  the  transference  of  a  function 
from  one  institution  to  another  ;  but  as  a  rule,  when 
this  change  is  effected,  it  is  attended  by  the  sup- 
pression of  the  old  institution.  Sometimes,  how- 
ever, this  is  not  the  case,  and  the  institution  merely 
atrophies  without  having  been  suppressed. 

(1)  BepuUican  institutions  under  the  Boman 
Empire. — As  instances  of  this  form  of  degenera- 
tion take  the  institutions  of  the  Eoman  Republic, 
after  the  establishment  of  the  Empire — or  the 
decadence  of  the  functions  formerly  discharged 
by  the  Privy  Council  in  England,  the  political 
functions  of  which  have  been  transferred  to  Parlia- 
ment, and  the  judicial  functions  to  the  Court  of 
Common  Pleas  and  the  Court  of  Exchequer.^ 

(2)  Special  jurisdiction  in  England. — It  is  the 

^  De  Franqueville,  Le  Gouvenument  et  le  Parlement  hrittanique, 
i.,  p.  431. 


ATROPHY  OF  INSTITUTIONS  FROM  LACK  OF  USE  285 

same  with  special  jurisdiction,  such  as  the  Eccle- 
siastical Court,  and  the  University  Courts ;  they 
have  completely  degenerated,  their  functions  having 
been  transferred  to  the  jurisdiction  of  the  Common 
Law  Courts.  In  England,  however,  special  juris- 
diction has  not  wholly  ceased  to  be  functional,  for 
there  still  exist,  side  by  side  with  the  modern 
Courts  of  Justice,  a  few  local  Courts  and  other 
exceptional  forms  of  jurisdiction  which  are  still 
maintained  in  support  of  certain  ancient  acquired 
rights  and  traditions.^  By  far  the  greater  part  of 
these  special  jurisdictions  have,  however,  fallen 
into  disuse,  owing  to  the  creation  of  the  modern 
Courts  of  Justice.  This  happened,  for  instance, 
with  the  following  institutions  : — 

{a)  The  Local  Courts  of  Feudal  origin. — These 
Courts  have  decreased  both  in  number  and  impor- 
tance since  the  close  of  the  thirteenth  century ;  the 
only  vestiges  now  remaining  of  them  are  the  Court- 
leets  of  certain  manors.  Sir  James  Stephen  makes 
special  mention  of  the  Court-leet  of  the  Manor  of 
Savoy,  which  extended  from  near  the  old  city  gate 
of  London  (Temple  Bar)  up  to  Cecil  Street.  Some 
of  the  old  functions  of  this  Court  are  maintained 
in  the  present  day. 

{b)   Country   Town    Courts. — Twenty-seven   local 

courts  are  mentioned  in  modern  judicial  statistics 

before  six  of  which  no   case  has   been   heard   for 

twelve  years.       On   the   application   of   a   litigant, 

^  Idem,  Le  sysUme  judiciaire  de  la  Grande- Bretagne,  i.,  p.  216. 


286         CAUSES  OF  DEGENERATIVE  EVOLUTION 

however,  any  one  of  these  would  resume  its 
functions.^ 

(c)  The  Court  of  Stannaries,  the  vice-warden 
of  which  is  appointed  by  the  Prince  of  Wales. 
This  Court  has  greatly  degenerated  in  importance, 
and  only  a  few  minor  cases  are  now  heard  before  it. 

{d)  The  Court  of  Piejpoivder. — This  was  a  Court 
of  summary  jurisdiction,  dealing  promptly  with 
disputes  arising  during  fairs  and  markets.  It  has 
almost  disappeared  from  having  fallen  into  disuse. 
Practically  only  one  example  remains ;  that  is  at 
Bristol,  and  is  becoming  merged  in  another  local 
court  called  the  Tolzey  Court. 

{e)  The  Hunting's  Court  (or  folkmote,  scirmote, 
a  kind  of  County  Court  of  the  city  of  London). — 
The  old  Husting's  Court  has  completely  lost  all  its 
former  attributes,  in  favour  of  the  Court  of  the 
Lord  Mayor  and  Sheriffs.  Up  to  1860,  however, 
it  reserved  to  itself  the  right  of  jurisdiction  with 
regard  to  matters  relating  to  landed  property  in 
the  city ;  the  only  cases  which  come  before  this 
Court  nowadays  are  those  of  replevy .^ 

Finally,   there    is    the    House    of    Lords    which 

^  According  to  jurisprudence,  the  fact  that  a  Court  still  existing 
by  virtue  of  Royal  Charter,  has  ceased  to  be  functional  for  two 
hundred  years,  is  no  reason  against  a  citizen  having  his  case  tried 
before  it,  if  he  so  chooses,  even  though  the  town  authorities  declare 
the  funds  to  be  insuificient  for  payment  of  the  judge. 

{Case  of  Rex  v.  Mayor  of  Wells,  Dowling  Practise  Cases,  p.  562), 
mentioned  by  Franqueville. 

2  Franqueville,  Systeme  jud. ,  i.,  pp.  235,  and  following. 


ATROPHY  OF  INSTITUTIONS  FROM  LACK  OF  USE  287 

formerly  held  full  powers  of  jurisdiction  with 
regard  to  matters  relating  to  members  of  the 
peerage ;  this  jurisdiction  is  now  limited  to  cases 
of  high  treason  or  crimes  committed  by  peers  to 
the  exclusion  of  mere  acts  of  misdemeanour  which 
are  tried  before  the  ordinary  Courts. 

§  2.  Atrophy  from  lack  of  resources. 

The  instances  we  have  mentioned  are  those  of 
institutions  decaying  because  they  had  become 
useless,  and  their  resources  were  transferred  to 
other  objects.  The  lack  of  resources  was  the  result 
of  the  lack  of  function.  Sometimes,  however,  it  is 
the  cause  and  not  the  result,  in  which  case  atrophy 
may  be  due  either  to  an  abnormal  development, 
such  as  the  hypertrophy  of  another  institution,  or  to 
poverty .  ensuing  on  the  general  decline  of  society 
in  general.  Of  course  both  factors  may  act 
simultaneously. 

1.  Local  admhiistration  at  the  close  of  the  Roman 
Empire. — The  decline  of  local  government  at  the 
close  of  the  Eoman  Empire  is  an  instance  of 
atrophy  ensuing  from  the  ultra-development  of 
another  institution. 

As  the  demands  of  the  central  powers  grew 
more  and  more  excessive,  the  fiscal  rates  had  to  be 
augmented  in  order  to  meet  them,  and  the  curiales, 
which  consisted  of  members  of  the  City  Council 
who  were  made   responsible  for    the    payment    of 


288         CAUSES  OF  DEGENERATIVE  EVOLUTION 

taxes,  finally  found  it  was  quite  impossible  to  meet 
their  engagements,  and  made  every  effort  to  leave 
the  curia.^ 

2.  The  degeneration  of  Societies  in  all  their  parts. 
— A  number  of  instances  might  be  mentioned  of 
general  social  degeneration  bringing  about  tlie 
atrophy  of  some  one  or  other  institution  in 
particular. 

Besides  giving  classical  examples,  such  as  the 
Konians,  Peruvians  and  Astecs,  V.  Lilienfeld  men- 
tions the  decline  of  the  Negro  kingdoms  which 
existed  during  the  sixteenth  and  seventeenth 
centuries  in  Southern  and  Western  Africa,  and 
which  are  merely  represented  nowadays  by 
wretched  little  tribes.^ 

There  are,  according  to  Waitz,  at  some  distance 
from  Carimango  (the  equatorial  Eepublic)  some 
people  of  pure  Spanish  blood  who  have  fallen  back 
into  absolute  barbarism.  Their  language  is  de- 
formed pa'fet  recognition,  and  their  manners  and 
customs  exhibit  no  traces  of  their  former  condi- 
tion.^ 

Space  precludes  us  from  dwelling  further  upon 
the  various  causes — often  complex  and  obscure — 
which  bring  about  the  downfall  of  societies, 
suffice    it   to    say    that    they  are   connected    with 

1  Lavisse  and  Rambeaud,  Histoire  g6n4rale,  I.,  ch.  i.,  pp.  14  and 
folloAving. 

2  Von    Lilienfeld,    Gedanke    iiber    de    Soziahvissenscha/t    der 
Zukimft,  ii.,  p.  241. 

^  Waitz,  Anthropologie  der  Nahcrvolker,  1.  B.,  p.  369. 


ATROPHY  OF  INSTITUTIONS  FROM  LA.CK  OF  USE    289 

territory  or    with    population,  the    two   factors  of 
social  evolution. 

Either  the  physical  surroundings  of  institutions 
undergo  unfavourable  transformations,  or  else  the 
population  itself  degenerates. 

(1)  The  almost  complete  disappearance  of  the 
great  family  communities  {zadrugas)  of  Montenegro 
is  a  characteristic  instance  of  atrophy  from  lack  of 
resources  caused  by  the  impoverishment  of  physical 
surroundings.  The  persistent  cutting  down  of  trees 
in  the  Black  Mountain  has  had  a  disastrous  effect 
on  the  water  supply,  and  consequently  upon  the 
fertility  of  the  ground.  Most  of  the  zadrugas ^ 
having  found  it  impossible  to  continue  their 
existence  in  common  upon  the  same  territory, 
have  split  up  into  small  families  (inoJcosnas). 
These  latter  represent,  in  a  reduced  state,  the 
old  family  system  from  which  they  have  sprung. 
Bogisic  has  shown  that  these  in  no  way  resemble 
our  modern  families,  but  are  to  be  regarded,  from 
the  judicial  point  of  view,  as  reduced  family  com- 
munities, each  comprised  of  only  a  few  persons. 

(2)  Other  cases  occur  where  the  atrophy  of  an 
institution — of  an  artistic  or  scientific  society,  for 
instance — is  brought  about  by  the  degeneration  of 
a  population  which  ceases  to  be  interested  in  the 
society  and  no  longer  contributes  to  its  support. 
A  large  number  of  cases  of  this  kind  might  easily 
be  mentioned,  especially  as  occurring  during  the 
period  of  the  Byzantine  Empire,  but  it  is  difficult 

T 


290    CAUSES  OF  DEGENERATIVE  EVOLUTION 

to  account  for  the  sudden  degeneration  of  a 
people  where  the  physical  surroundings  had  re- 
cently undergone  no  special  modifications,  and 
when  there  had  been  no  sudden  and  violent 
check  upon  social  development.  According  to 
Lapouge  and  other  sociologists  of  the  Darwinian 
school,  this  social  degeneration  was  merely  the 
outcome  of  hereditary  influences.  The  destiny  of 
a  nation  is  dependent  upon  the  quality  of  the 
elements  of  which  it  is  composed  and  by  which 
it  is  directed.  If  a  nation  is  rich  in  energetic 
and  intelligent  qualities,  the  greatest  of  disasters 
can  only  have  a  transitory  and  limited  influence. 
When  the  contrary  is  the  case,  the  same  circum- 
stances may  produce  an  arrest  in  development 
or  a  complete  decline  and  fall.  Up  to  the 
present  time,  and  especially  in  antiquity  and  the 
middle  ages,  these  favourable  qualities  were  gene- 
rally supplied  by  a  dominating  minority  establish- 
ing itself  in  a  conquered  country.  In  the  common 
course  of  evolution,  these  superior  elements,  which 
are  indispensable  to  social  progress,  are  eventually 
eliminated.  The  inferior  elements  regain  greater 
power,  and  each  step  of  their  progress  is  attended 
by  a  backward  step  towards  barbarism.  Although, 
at  first  sight,  this  seems  contrary  to  the  Darwinian 
theory,  it  is  strictly  in  accordance  with  it.  The 
superior  individuals  are  relatively  inferior  when 
their  chances  of  success  or  of  posterity  diminish. 
The  superior  individuals  may  not  only  be  swamped 


ATKOPHY  OF  INSTITUTIONS  FROM  LACK  OF  USE  291 

by  a  diminution  in  their  birth-rate,  but  in  some 
cases  there  may  be  a  direct  elimination  of  them.^ 

The  tendency  of  decadence  is  always  towards  the 
degenerative  and  eliminatory  selection  of  superior 
elements. 

It  may  be  said  in  conclusion  that  there  are 
constant  calls  upon  the  capital  and  labour  of  a 
society  from  its  various  institutions,  and  the  conse- 
quence is  that,  the  resources  not  being  unlimited,  a 
regular  struggle  for  existence  goes  on  amongst  the 
institutions.  In  the  course  of  this  struggle,  the 
decline  of  an  institution  may  be  brought  about 
in  two  different  ways.  It  either  begins  to  de- 
generate from  lack  of  sufficient  means  of  support, 
or  degeneration  sets  in  consequent  upon  the  insti- 
tution having  ceased  to  be  functional  by  inutility, 
by  transference  of  function  to  another  institution, 
or  by  obstacles  placed  in  the  way  of  exercising 
that  function.  In  either  case  the  institution  dis- 
appears. It  is  only  in  exceptional  cases,  which 
will  be  alluded  to  further  on,  that  existence  is 
still  maintained. 

^  G.  de  Lapouge,  La  Vie  et  la  Mort  des  nations  {Mvue  int.  de 
Sociologie,  1894,  pp.  421  and  following).  Several  terms  used 
in  this  treatise  were  borrowed  from  the  above  article. 

See  also  Hovelaeque  and  Herve,  Precis  d' Anthropologie,  p.  189  : 
' '  War,  in  its  double  consequence  of  the  elimination  of  the  strong 
and  the  survival  of  the  weak,  is  for  the  more  civilized  races  a 
powerful  factor  in  the  cause  of  degeneration  and  downfall," 


PART  II 

THE    CAUSES    OF    THE    PERSISTENCE    OF    ORGANS 
OR    INSTITUTIONS    WITHOUT    FUNCTION 


CHAPTEK    I 

SURVIVAL    OF    ORGANS 

We  have  shown  how  and  why  organs  may  become 
rudimentary  and  tend  to  disappear.  In  many 
cases  the  disappearance  is  complete ;  and  the  organ 
may  not  even  reappear  temporarily  in  the  course  of 
the  individual  development.  This  disappearance 
is,  however,  by  no  means  universal.  Even  apart 
from  the  phenomena  of  recapitulation,  rudimentary 
organs  may  persist  in  the  adult,  and  sometimes,  even 
although  organs  have  ceased  to  be  functional,  they 
persist  without  degenerating.  We  have  now  to  con- 
sider why  in  such  cases  degenerative  evolution  does 
not  result  in  complete  obliteration  of  such  organs. 

§   1 .    UnfunctioTial  orgaiis  that  are  not  rudimentary. 

Absence  of  Variations. — There  are  some  plants 
such    as    Ficaria'^   ranunculo'ides    and    Zysimachia 

1  Lysimachia  Nummularia  occasionally  produce  seeds  in  some 
valleys  of  the  Pyrenees,  and  Errera  has  shown  us  specimens  grown 
from^seeds  coming  from  the  shores  of  the  lake  of  Quatre-Cantons, 


SURVIVAL  OF  ORGANS  293 

Nummularia,  the  flowers  of  which  hardly  ever 
produce  seeds.  How  is  it  that  in  such  species 
flowers  are  still  produced  ?  The  probable  ex- 
planation of  this  anomaly  is,  that  for  the  disappear- 
ance of  flowers  there  would  have  to  be  produced 
individuals  with  this  advantageous  variation.  It 
is  the  case,  however,  that  the  Ficaria  and  the 
Lysimachia  reproduce  most  actively  by  asexual 
methods,  and  variations  are  extremely  rare  in 
cases  of  these  modes  of  reproduction.  The  result 
is  that  these  species  having  begun  to  form  sterile 
flowers  continue  to  produce  them  through  simple 
lack  of  variation. 

An  analogous  case  is  presented  by  Elodea 
Canadensis.  This  unisexual  plant  is  represented 
in  Europe  by  only  female  plants.  These  plants 
have  multiplied  asexual ly  so  luxuriantly  that  in 
Holland  they  began  to  choke  up  the  canals,  and 
it  became  necessary  to  make  provision  in  the 
budget  of  that  country  for  the  extermination  of 
the  pest.  The  plants  are,  of  course,  able  to  multiply 
only  asexually,  as  the  female  flowers  cannot  be 
fertilized,  and  these  useless  flowers  have  been 
maintained  simply  from  the  absence  of  variations. 

Stratiotes  alo'ides,  a  plant  belonging  to  the 
same  family  as  Elodea,  is  practically  only  repre- 
sented by  male  individuals.  Females  are  extremely 
rare,  and  none  the  less  the  male  flowers  are  pro- 
duced, although  in  the  vast  majority  of  cases  they 
must  be  useless. 


294    CAUSES  OF  DEGENERATIVE  EVOLUTION 

W.  Burck  has  called  attention  to,  without 
endeavouring  to  explain,  other  instances  of  the 
persistence  of  functionless  organs.  A  large 
number  of  Anonaceae  bear  flowers  which  do  not 
open,  and  which  are  self -fertilizing  (cleistogamous 
flowers).  None  the  less,  they  have  retained  the 
corolla,  the  original  purpose  of  which  was  to 
attract  insects.^ 

Burck  has  called  our  attention  to  the  circum- 
stance that  several  species  belonging  to  the  same 
genus  produce  cleistogamous  flowers,  and  that  it  is 
improbable  that  this  condition  has  been  acquired 
independently  by  these  species.  One  would  thus 
have  to  admit  that  the  original  type  must  be  very 
remote,  as  it  has  given  rise  to  descendants  of 
specific  distinction,  and  yet  the  useless  corolla  has 
persisted  through  the  long  series. 

Parallel  examples  may  be  found  among  animals. 

Machaerites  is  an  insect  which  inhabits  caves 
in  North  America.  The  females  are  quite  blind  ; 
the  males,  on  the  other  hand,  have  preserved,  or 
seem  to  have  preserved,  well- developed  eyes  ;  but 
are  these  eyes  real?  An  abyssal  fish,  Ijnuyps,^ 
seems  as  if  it  had  enormous  eyes  extending  from 
the  corner  of  the  snout  some  distance  along  the 
neck,  but  these  organs  are  not  really  eyes ;  they 

^  W.  Burck,  Ueber  Kleistogamie  im  weileren  Sinne  und  das 
Knight- DarwiTische  Gesetz  {Anii.  Jard.  Bot.  Buitenzorg,  viii.,  p. 
122,  1889). 

2  DoUo,  La  Vie  au  sein  des  mers,  p.  242,  Paris,  Bailliere,  1891. 


SURVIVAL  OF  ORGANS  295 

are  light-producing  organs,  and  the  fish  are  in 
reality  blind.  This  may  be  the  case  also  in  the 
males  of  Machaerites.  It  may  also  be  the  case 
that  the  male  has  an  opportunity  for  using  eyes 
absent  in  the  case  of  the  female,  the  males  some- 
times leaving  the  caves,  the  females  remaining 
within  them.  Something  analogous  to  this  occurs 
in  the  case  of  eels :  the  males  remain  always  in 
the  sea  while  the  females  rejoin  them  only  for 
purposes  of  reproduction.  Moreover,  there  is  still 
a  third  hypothesis,  that  the  male  of  Machaerites  be- 
came an  inhabitant  of  caves  later  than  the  female, 
and  has  not  yet  had  time  for  the  loss  of  its  eyes. 

§  2.    Unfunctional  organs  which  persist  as  rudiments. 

It  is  outside  our  purpose  to  discuss  here  the 
numerous  cases  of  organs  reduced  through  adapta- 
tion, such  as  the  leaves  reduced  to  serve  as 
protectors  of  young  buds  (Phyllocactus,  fig.  78),  or 
the  wings  of  the  ostrich  which,  although  much 
reduced,  are  supposed  to  assist  the  bird  in  running. 
The  utility  of  such  organs  explains  their  persist- 
ence ;  we  are  concerned  here  with  organs  which, 
although  useless,  persist  in  a  reduced  form. 

1.  Absence  of  Variations. — In  discussing  organs 
which,  although  without  function,  have  persisted  in 
a  complete  state,  we  attributed  the  persistence  to 
absence  of  variations.  It  is  probable  that  the 
same  cause  operates  in  maintaining  useless  vestiges. 


296    CAUSES  OF  DEGENERATIVE  EVOLUTION 

It  is  to  be  noticed,  however,  that  the  variability 
of  vestiges  is  frequently  considerable.  The  flowers 
of  Asparagus  officinalis  are  sometimes,  although 
rarely,  hermaphrodite.  Usually  they  are  uni- 
sexual and  exhibit  the  organs  of  the  other  sex  in 
every  conceivable  stage  of  degeneration.  It  is 
probable  that  the  unisexual  condition  has  been 
acquired  recently,  and  that  there  has  not  yet  been 
time  for  the  operation  of  natural  selection  to  cause 
the  disappearance  of  the  useless  organs.^ 

^  We  have  actual  knowledge  of  the  mode  of  disappearance  of  an 
organ  in  one  case,  and  can  see  the  part  played  by  variability.  In 
Phyllanthus  speciosus  (Xylophylla  arhuscula)  the  adult  plant  has 
three  kinds  of  branches  :  vertical  branches,  with  rudimentary 
leaves  ;  oblique  branches  which  spring  from  the  axils  of  these 
leaves  and  themselves  bear  in  two  rows  very  rudimentary  leaves  ; 
and  flat  branches  which  are  the  chief  organs  of  assimilation  of  the 
plant  and  which  also  bear  rudimentary  leaves  in  two  rows.  In  the 
seedling,  on  the  other  hand,  there  are  formed  after  the  two  coty- 
ledons, one  or  two  completely  developed  assimilating  leaves  upon 
an  upright  stem  (fig.  84,  a). 

The  flat  branches  grow  from  the  axils  of  these,  and  bear, 
unlike  the  flat  branches  of  the  adult,  assimilating  leaves  ;  higher 
up  the  vertical  stem  bears  only  rudimentary  leaves  with  flat 
branches  in  their  axils. 

It  may  happen,  however,  that  the  seedlings  bear,  directly  after 
their  cotyledons,  rudimentary  leaves  (fig.  84,  b)  and  in  this  case 
the  reduction  of  the  leaves  is  present  not  only  in  the  leaves  borne 
upon  the  vertical  stem  but  in  those  borne  on  the  flat  branches. 
These  latter  bear  only  a  small  number  of  assimilating  leaves. 
Thus,  we  have  in  this  plant  a  remarkable  example  of  incomplete 
recapitulation  :  the  seedling  preserves  in  a  functional  condition 
organs  that  are  rudimentary  in  the  adult,  but  the  species  furnishes 
instances  where  these  leaves  cease  to  be  functional  even  in  the 


SURVIVAL  OF  ORGANS 


297 


2.  Insignificance  of  the  Rudimentary  Organ. — 
It  frequently  happens  that  rudimentary  organs  are 
preserved  simply  on  account  of  their  insignificance  : 
the  absence  of  organs  so  small  would  not  be  an 
advantage  to  the  plant  sufficiently  great  to  be  laid 
hold  of  by  natural  selection. 

Many  species  of  Tropaeolum  bear  leaves  without 


Fig.  84. — Two  seedlings  of  Phyllanthus  speciosus. 
1,  2,  3,  4,  successive  assimilating  branches ;  c,  cotyledons. 

stipules.  In  Tropaeolum  majus  there  are  stipules 
only  in  the  case  of  the  first  two  leaves  of  the 
seedling,  and  the  position  of  these  stipules  is  very 
variable.  Sometimes  both  stipules  are  at  the  base 
of  the  petiole;  sometimes  only  one  is  present; 
sometimes  both  are  several  millimetres  from  the 
base — a  position  in  which  functional  stipules  never 
occur. 


298         CAUSES  OF  DEGENERATIVE  EVOLUTION 

In  the  same  way  may  be  explained  the  persist- 
ence of  accessory  rudiments  of  enamel  organs  in 
;the  development  of  teeth.  Besides  the  rudiments 
of  the  enamel  organs  for.  the  milk  teeth  and 
permanent  teeth,  there  are  additional  organs 
present  in  a  very  variable  condition  and  number, 
nearer  the  external  surface.  They  are,  however, 
very  generally  present,  and  are  exceedingly  similar 
to  the  youngest  stages  of  the  normal  enamel  organs. 
Kollmann  and  Gegenbaur  believe  that  they  are 
abortive  rudiments  surviving  from  an  ancestral 
condition  in  which  teeth  were  more  numerous. 


CHAPTER  II 

THE    SURVIVAL    OF    INSTITUTIONS 

We  have  seen"  that  an  institution  which  ceases  to 
be  functional  without  dissolution — either  voluntary 
or  enforced — ensuing,  tends  to  atrophy  and  dis- 
appear, while  its  resources  are  appropriated  by 
other  institutions.  It  now  remains  to  account  for 
the  fact  that  this  atrophy  does  not  generally  end  in 
total  disappearance.  The  two  following  hypotheses 
may  be  made : — 

1.  The  structure  of  a  non-functional  institution 

may  remain  intact. 

2.  The  institution  may  survive,  but  in  a  rudi- 

mentary condition. 


SURVIVAL  OF  INSTITUTIONS  299 

§    1.    The  integral  persistence  of  an  institution. 

A  non-functional  institution  may  survive  and 
retain  its  structure  and  resources  in  the  following 
cases : — 

1.  By  the  intervention  of  some  superior  authority 

to  prevent  its  suppression. 

2.  When,  while  ceasing  to  be  functional,  it  con- 

tinues to  be  useful,  though  indirectly  so. 

3.  When  its  existence  is  maintained  out  of  respect 

for  old  traditions. 

We  will  take  these  three  hypotheses  in  succession, 
but  it  must  be  borne  in  mind  that  when  a  non- 
functional institution  is  maintained  out  of  respect 
to  tradition,  or  by  virtue  of  an  indirect  usefulness, 
it  is  always  by  the  intervention  of  legal  authority. 
This  legal  authority,  however,  amounts  to  an  ex- 
pression of  the  public  will,  whereas  an  institution 
may  also  be  maintained  by  the  exercise  of  personal 
influence  on  the  part  of  some  one  person. 

It  sometimes  happens,  too,  that  a  non-functional 
institution  continues  to  survive  because  its  suppres- 
sion would  entail  important  changes  in  other  parts 
of  the  social  organization. 

1.  Maintenance  by  compulsion. — A  useless  in- 
stitution is  frequently  maintained  by  compulsion, 
when  its  conservation  is  advantageous  to  those  con- 
nected with  it,  or  even  to  other  persons.^ 

The  following  are  a  few  examples  of  this  : — 

^  Spencer,  Principles  of  Sociology,  vol.  iii. 


300    CAUSES  OF  DEGENERATIVE  EVOLUTION 

{a)  A  long  list  might  be  made  of  all  the  sinecures, 
now  quite  useless,  that  some  governments  insist 
upon  maintaining  for  the  advantage  of  those  occu- 
pying the  posts.  Such  were  certain  offices  in  con- 
nection with  the  Court  in  former  days  or  the 
avoueries  of  the  end  of  the  middle  ages. 

"  Like  the  Fief  system,"  says  Errera,  when  writing 
about  the  Massuirs,  "  The  avouerie  afforded  an  effec- 
tive protection — military  as  well  as  judicial — 
against  the  various  dangers  arising  in  a  still  barbaric 
age.  But,  in  the  course  of  the  last  centuries  of  the 
middle  ages,  the  obligations  of  feudal  chiefs  and 
the  condition  of  avouerie  disappeared ;  the  reasons 
were  that  relative  security  was  attained ;  militia 
was  established,  and  the  army,  under  the  command 
of  the  sovereign  himself,  became  better  disciplined ; 
and  that  there  arose  the  organization  of  the  justices 
scabinales,  of  bailiwicks,  and  of  superior  courts  of 
justice.  However,  although  the  ancient  offices  dis- 
appeared, the  emoluments  attached  to  them  con- 
tinued to  be  drawn.^ 

(h)  It  often  happens  that  institutions  which  have 
ceased  to  be  functional  are  yet  maintained  as  being 
a  source  of  profit  not  only  to  those  in  direct  con- 
nection with  them,  but  to  a  considerable  number  of 
other  persons. 

Before  the  Keform  Bill  of  1832,  when  large 
towns  like  Leeds,  Birmingham,  and  Manchester 
were   unrepresented   in   Parliament,  the   House   of 

*  Errera,  Les  Massuirs,  p.  75.     Brussels,  Weissenbruch,  1892. 


SUEVIVAL  OF  INSTITUTIONS  301 

Commons  contained  seventy  members,  nominated 
by  thirty-five  rotten  boroughs  in  which  there 
were  no  electors,  and  ninety  members  nominated 
by  forty-six  boroughs,  containing  less  than  fifty 
electors. 

The  borough  of  Old  Sarum  was  a  mere  hillock 
belonging  to  Lord  Canalford  ;  Gatton  and  S.  Michael 
had  only  seven  electors ;  the  borough  of  Dumwich 
had  been  long  since  submerged  by  the  encroach- 
ment of  the  sea;  Beeralston,  belonging  to  Lord 
Beverley,  consisted  of  one  house,  and  Castlerising  of 
two.  In  the  county  of  Bute  there  were  twenty- 
one  electors,  only  one  of  whom  was  a  resident  and 
who  nominated  himself. 

The  preservation  of  this  system  of  representation 
which  had  long  ceased  to  be  adequate,  was  eminently 
favourable  to  the  few  persons  who  benefited  by  it, 
and  they  vigorously  resisted  the  passing  of  the 
Reform  Bill. 

(c)  After  the  provincial  states  of  Kormandy  and 
the  Dauphiny  had  been  suppressed,  the  state  func- 
tionaries retained  their  titles  and  emoluments. 

In  the  Dauphiny  the  representative  institutions 
ceased  to  be  functional  in  1627,  but  at  the  close  of 
the  eighteenth  century  the  Bishop  of  Grenoble  con- 
tinued to  receive  6000  livres  per  annum  as  primate 
of  the  dominion.  Two  barons,  delegates  of  the 
nobility,  shared  a  similar  salary,  and  the  Syndic 
of  the  province  and  two  secretaries  received  lesser 
emoluments    from  the  province    which   continued 


302    CAUSES  OF  DEGENERATIVE  EVOLUTION 

to  make  these  payments  for  services  no  longer 
rendered.^ 

In  each  of  the  above  cases  of  survival  it  is  plain 
that  compulsion  intervened — more  or  less  directly 
— to  secure  the  maintenance  of  non-functional 
institutions. 

In  the  first  cases  referred  to,  the  privileged 
persons  concerned,  took  advantage  of  their  influ- 
ential positions  to  enforce  the  maintenance  of  their 
sinecures. 

In  the  second  case — that  of  the  rotten  boroughs 
— the  institution  was  not  only  advantageous  to  the 
member  himself  but  to  the  whole  of  his  party,  so 
that  naturally  its  maintenance  met  with  the  cordial 
support  of  the  latter. 

In  the  third  case,  those  in  authority  maintained 
part  of  the  institution — that  of  the  mere  titles 
and  emoluments — in  order  to  suppress  the  remainder 
more  easily.  In  this  case  it  was  not  an  institution 
which  atrophied,  but  an  institution  which  was  caused 
to  atrophy  by  compulsion. 

2.  Indirect  usefulness. — It  sometimes  happens 
that  an  institution,  although  ceasing  to  be  functional, 
yet  retains  a  certain  usefulness.  This  is  so  in 
England,  with  the  office  of  the  Privy  Seal.  All 
the  functions  formerly  discharged  by  the  holder 
of  this  office  have  long  since  disappeared,  but  the 
post  is  reserved  as  a  sinecure  for  persons  who  have 

^  Babeau,  Les  assemblies  des  pays  d'Etat  sous  Vancien  regime; 
Jl^forme  sociale,  1893,  p.  704, 


SURVIVAL  OF  INSTITUTIONS  303 

distinguished  themselves  in  poHtics,  but  who  from 
advanced  age  are  unable  to  take  an  active  part  in 
public  afiairs.  A  great  many  sinecures  are  main- 
tained for  a  similar  purpose — that  of  furnishing 
practical  though  not  nominal  pensions  to  distin- 
guished persons  in  art  or  science. 

"  It  may  well  be,"  says  Viollet,  "  that  an 
institution,  which  at  first  sight  seems  to  be  a 
mere  useless  wreck,  is  really  of  immense  service 
to  society.  The  deep  roots  of  a  dead  tree  may 
continue  to  furnish  support  to  a  new  structure." 

It  is  by  reason  of  this  negative  usefulness  that 
— according  to  Bagehot — the  English  monarchy 
has  been  preserved,  forming  as  it  does  a  symbol 
of  unity  and  coherence  amidst  the  electoral  changes 
of  power.  The  English  monarchy  offers  a  char- 
acteristic instance  of  an  institution  which  has  lost 
nearly  all  its  former  functional  importance,  while 
nominally  retaining  its  power.  According  to 
Bagehot,  the  Queen  would  now  have  to  sign  her 
own  death  warrant  if  condemned  by  both  Houses 
of  Parliament.  The  outer  form,  however,  remains 
almost  the  same  as  in  the  days  of  absolute  monarchy, 
when  the  sovereign  took  an  active  part  in  public 
affairs.  At  within  the  last  few  years,  the 
Queen's  signature  was  required  to  all  official 
documents.  It  was  only  in  1862  that  a  law 
was  passed  deciding  that  for  the  future,  promotions 
in  the  Army  or  Navy  should  be  signed  either  by 
the  Commander-in-Chief,  or  by  the   Secretary  of 


304         CAUSES  OF  DEGENERATIVE  EVOLUTION 

State.  At  that  time,  by  dint  of  hard  work, 
the  Queen  had  signed  all  commissions  up  to 
1858,  and  there  remained  15,931  documents  of 
this  kind  still  unsigned.^ 

3.  Respect  for  Tradition. — The  persistence  of 
some  institutions  can  only  be  accounted  for  by  a 
lack  of  invention.  Mommsen  calls  attention  to 
a  remarkable  instance  of  this  kind  in  the  history 
of  early  Rome  : 

When  a  government  by  praetors — as  consuls 
were  first  called — was  substituted  for  a  govern- 
ment by  kings,  the  new  system  remained  the  same 
in  idea  though  nominally  different.  The  old  idea 
of  royal  authority  survived  for  a  long  period,  and 
the  praetors  enjoyed  all  the  old  kingly  powers,  even 
those  in  plain  contradiction  to  the  temporary 
character  of  their  office :  the  king  could  not  be 
deposed,  but  neither  could  the  praetor  be  con- 
strained to  depose  himself ;  the  king,  when  dying, 
nominated  his  successor  himself,  and  this  power 
remained  to  the  praetor — although  the  system  of 
election  by  the  comitia  had  been  introduced — 
for  the  praetor  had  the  right  of  excluding  whoever 
he  chose  from  the  list  of  candidates,  and  also  of 
annulling  the  votes  given  to  those  who  displeased 
him.     It  was  only  at  a  later  period  that  a  logical 

*  Bagehot,  The,  English  Constitution,  pp.  57  and  following ; 
London,  1891. 

De  Franqueville,  Gouvemement  et  ParUment  BritanniqueSj  i., 
p.  251. 


SURVIVAL  OF  INSTITUTIONS  305 

and  consistent  conception  of  the  consular  authority- 
came  to  be  formed.  1 

In  the  same  way  the  maintenance  of  the  insti- 
tution of  sheriffs  in  modern  England  can  only  be 
attributed  to  an  exaggerated  respect  for  tradition. 

In  the  commission  which  sat  in  1888  to  inquire 
into  judicial  organization,  one  of  the  commissioners 
expressed  himself  as  follows  :  "  I  cannot  see  that 
sheriffs  are  of  any  use  whatever,  unless  it  be  for 
show ;  there  is  not  one  single  function  attached  to 
the  post  which  a  sheriff  can  fulfil  himself.  I  am 
a  sheriff,  but  I  do  not  know  what  a  sheriff's  duties 
are."  The  institution,  however,  remains  intact. 
The  sheriff  is  credited  with  the  discharge  of  several 
functions,  which  are  executed  in  his  name  and  on 
his  responsibility.  All  he  does  is  to  receive  the 
judges,  accompany  them  on  circuit,  and  preside  over 
parliamentary  elections.^ 

The  mode  of  nominating  the  sheriffs  has  remained 
unchanged  since  1340.  The  Queen,  by  means  of  a 
traditional  gold  pin,  is  supposed  to  prick  by  chance 
in  the  list  of  candidates  which  is  presented  to  her, 
the   name   of   him   upon  whom   she   confers   "  the 

1  Ferrero,  Simbolii,  p.  53,  Torino,  1893. 

2  *'  The  two  sheriffs  of  London,  who  are  elected  from  among  the 
liverymen  of  the  various  city  companies,  have  not  to  go  on  circuit, 
but  are  supposed  to  attend  at  the  Central  Criminal  Court.  Their 
functions  chiefly  consist  in  escorting  the  Lord  Mayor  to  all  city 
ceremonies,  and  in  attending  numerous  banquets,  some  of  which 
are  given  at  their  expense."  De  Franqueville,  Systeme  judiciaire  de 
la  Grand- Br etagne,  i.,  p.  611. 

U 


306         CAUSES  OF  DEGENERATIVE  EVOLUTION 

charge  and  keeping  of  the  county."     This  ceremony 
is  known  as  the  "  Pricking  of  Sheriffs." 

§  2.  The  Survival  of  Institutions  in  a  reduced 

STATE. 

It  has  been  shown  that  a  number  of  institutions 
in  a  state  of  decline  continue  to  be  maintained, 
because  they  are  still  useful  in  spite  of  their  reduced 
condition. 

This  is  the  case  with  the  symbolic  ceremonies 
which  in  former  days  accompanied  the  drawing  up 
of  solemn  contracts.^ 

We  now  come  to  institutions  which  persist  in  a 
reduced  condition,  but  which  are  not  directly  use- 
ful to  any  one.  In  this  case  their  persistence 
may  be  attributed  to  one  of  two  causes:  either  to 
respect  for  tradition  or  to  the  insignificance  of  the 
vestiges  which  remain. 

^  According  to  Viollet  in  Histoire  du  droit  civil  fr.y  p.  607,  the 
primitive  assembly  of  the  people  still  survives,  though  in  a  reduced 
condition,  in  the  Roman  mancipation  and  in  Scandinavia  in  a  solemn 
form  of  sale  called  the  scotatio, 

•*  I  believe,"  he  says,  "  that  it  was  the  primitive  sale  of  German 
law  a  sale  concluded  and  ratified  in  a  popular  assembly,  that  gave 
rise,  in  the  middle  ages,  to  the  Scandinavian  scotatio.  So  also  in 
the  case  of  the  Roman  inancipatio  there  has  long  been  believed  to 
exist  the  remnants  of  a  popular  assembly.  The  dumb  witnesses  in 
the  scotatio  appear  to  me  the  petrified  representatives  of  the 
German  tribe  or  village  ;  and,  in  the  opinion  of  good  judges,  the 
witnesses  in  mancipatio  are  no  other  than  symbolic  statues  of  the 
five  classes  of  the  Roman  people.  However,  this  is  mere 
hypothesis. 


SURVIVAL  OF  INSTITUTIONS  307 

It  is  difficult,  however,  to  distinguish  between 
these  two  causes,  for  insignificant  vestiges  of 
institutions  are  especially  numerous  in  very  con- 
servative environments,  and,  on  the  other  hand, 
mere  respect  for  tradition  very  rarely  ensures  the 
maintenance  of  either  a  harmful  or  an  expensive 
institution. 

1.  Insignificance  of  the  Institution. — Some 
of  these  insignificant  institutions  are  reduced  to  mere 
vestiges,  no  longer  functional,  or  of  use  to  any  one, 
others — representing  traces  of  a  former  system — 
retain  some  local  vitality,  although  not  in  keeping 
with  the  new  conditions  ;  these  are  of  such  small 
importance  that  their  very  insignificance  ensures 
their  survival. 

(a)  The  two  French  laws  passed  in  1835  and 
1849  relating  to  entail  have  never  become  func- 
tional  at  La  Martinique.  The  old  law  of  entail  is 
still  active  there,  and  this  state  of  things  is  allowed 
to  continue  without  authoritative  interference.^ 

(b)  In  England,  the  sovereign,  up  to  the  close  of 
the  sixteenth  century,  reserved  to  himself  the  right 
of  presiding  over  the  Courts  of  Justice,  and  of 
pronouncing  sentence.  Since  the  Ee volution,  no 
sovereign  has  essayed  to  render  justice  personally, 
and  any  attempt  of  the  kind  would  be  regarded  as 
unconstitutional  nowadays.  Vestiges  of  the  old 
system  remain,  however,  in  the  formula  of  certain 
legal  proceedings,  such  as  the  serving  of  warrants, 

^  Viollet,  Hist,  du  dr.  civ.fr.,  p.  883. 


/ 


308         CAUSES  OF  DEGENERATIVE  EVOLUTION 

in  which  an  order  is  given  to  appear  before  the 
Queen  herself,  and  it  has  never  been  suggested  that 
the  old  formula  should  be  altered.^ 

The  following  are  a  few  examples  of  local  sur- 
vivals which  owe  their  existence  to  their  insignifi- 
cance : — 

(a)  Although  Cambray  has  belonged  to  France 
since  the  close  of  the  seventeenth  century, 
its  diocese  includes  a  small  part  of  Belgium; 
the  ancient  religious  organization  has  in 
this  instance  survived  political  changes.^ 
(Jb)  It  is  not  generally  known  that  with  certain 
properties  situated  in  Artois — consisting 
chiefly  of  marsh  land — the  law  of  primo- 
geniture still  holds  good.^ 
(c)  Public  attention  has  recently  been  directed 
to  a  very  curious  survival  of  the  political 
connections  which  existed  during  the  middle 
ages  between  Beam  and  Spain.  Every 
year,  on  the  13  th  of  July,  the  inhabitants 
of  the  French  valley  of  Baretous  in  the 
Pyrenees,  solemnly  pay  a  tribute — for  the 

^  De  Franqueville,  Systeme  judiciaire  de  la  Grande- Bretagne, 
p.  23. 

2  Viollet,  Hist.  du.  dr.  civ.  fr.,  p.  882. 

^  Errera,  Les  Masuirs,  p.  290.  Decision  of  the  Council  of  25th 
February  1779,  as  regards  the  castle-wards  of  Lille,  Douai,  and 
Orchies.  *'  Those  portions  of  land  which  fall  or  have  fallen  to  each 
inhabitant  as  the  result  of  division,  shall  be  inalienable ;  no  person 
shall  possess  two  portions.  The  eldest  male  of  each  family,  or  in 
default  of  the  eldest  male,  the  eldest  female,  shall  alone  inherit 
the  said  portions  of  land." 


SURVIVAL  OF  INSTITUTIONS  309 

maintenance  of  peace — to  the  inhabitants 

of  the  Spanish  valley  of  Eoncal.^ 

It  is  plain  that  this  custom,  which  has  gone  on 

so  long  as  to  pass  unnoticed,  is  too  inconsistent 

with    the    present    relations    between  France   and 

1  The  ceremony  commences  at  nine  o'clock  in  accordance  with 
instructions  laid  down  in  a  document  which,  according  to  the 
mention  made  of  it  in  the  Proces-verbal,  dates  from  1375. 
First,  the  French  mayors  don  their  scarves  of  office  ;  next  the 
Spanish  mayors,  advancing  from  a  group  of  compatriots,  proceed 
towards  the  frontier  boundary  line,  accompanied  by  a  peasant 
carrying  a  lance  with  a  red  flag  tied  to  it — a  symbol  of  justice — 
and  draw  up  within  six  yards  of  the  boundary.  The  French 
mayors  follow  suit,  but  the  flag  hoisted  on  the  lance  which 
precedes  them,  carried  by  a  peasant,  is  white,  as  signifying 
their  pacific  intentions.  The  Mayor  of  Isaba  then  says  to 
them,  "Is  it  peace?"  The  French  mayors  reply  in  the  affirma- 
tive, and,  as  a  proof  of  their  sincerity,  couch  their  lance  upon  the 
mile-stone  marking  the  boundary.  The  Spaniards  then  first  plant 
their  lance  in  French  soil  leaning  against  the  stone,  and  after- 
wards place  it  so  as  to  form  a  cross  with  the  French  lance. 
Next  the  Mayor  of  Arette  places  one  hand  upon  the  crossed 
lances,  and  the  Mayor  of  Isaba  does  the  same,  and  together 
they  utter  the  formal  declaration  of  peace,  which  all  those 
present  swear  to  observe.  After  the  vow,  the  Mayor  of  Isaba 
cries  three  times,  "Paz  davans  !  "  which  means,  *'  May  peace  con- 
tinue." 

Peace  being  thus  declared,  the  Roncalais,  in  order  to  ratify 
their  abandonment  of  hostilities,  order  the  guards  to  remove 
their  arms  from  the  French  side.  The  ceremony  being  over, 
it  only  remains  to  pay  the  blood-tax.  This  formerly  consisted 
of  three  white  mares,  all  exactly  alike  ;  but,  owing  to  the  great 
difficulty  of  matching  them,  three  unblemished  heifers  were  sub- 
stituted, all  of  same  colour  and  with  the  same  markings.  Tliese 
three  heifers  cost  about  600  francs,  which  is  a  large  sum  for  that 
part  of  the  world. 


310  CAUSES  OF  DEGENERATIVE  EVOLUTION 

Spain  to  be  continued  much  longer.  "  It  is  to 
be  expected,"  says  the  paper  from  which  this 
information  was  obtained,  "  that  the  French 
Foreign  Minister  will  in  the  near  future  come 
to  an  understanding  with  the  Spanish  authorities 
to  put  an  end  to  this  iniquitous  custom,  and  it  is 
to  be  hoped  that  this  year  is  the  last  occasion  upon 
which  a  blood-tax  will  be  paid  by  the  valley  of 
Bare  tons  to  the  valley  of  Roncal." 

2.  Respect  for  tradition. — In  his  "  Essays  on  Pro- 
gress, Manners  and  Customs  "  (  Westminster  Review, 
1854),  Spencer  points  out  the  connection  between 
respect  for  tradition  and  custom,  and  the  con- 
servatism of  those  in  authority.  He  says  that 
certain  customs,  which  have  elsewhere  died  out, 
survive  in  some  departments  of  the  government. 
The  Secretary  of  State  in  ratifying  acts  passed  in 
Parliament  uses  old  Norman  French,^  and  certain 
legal  terms  in  old  Norman  French  are  still  used. 
The  wigs  now  worn  by  judges  and  barristers  are 
identical  with  those  seen  in  old  portraits,  while  the 
"  Beef -eaters "  of   the  Tower  of  London  wear   the 

1  *'  For  financial  Acts  the  formula  is :  La  Reyne  remercie  ses  bons 
sujets,  accepte  leur  benevolence  et  ainsi  le  veult ;  for  general  Acts  : 
La  Reyne  le  veult ;  for  private  bills :  Soit  fait  comme  il  est  desire  ; 
for  petitions :  Soit  droit  fait  comme  il  est  desire.  The  veto  is 
announced  :  La  Reyne  s'avisera.  Cromwell  had  changed  these  old 
forms ;  he  gave  his  consent  to  Bills  in  English  ;  the  old  custom 
was  resumed  at  the  Restoration,  and  the  House  of  Commons  in 
1706  rejected  a  Bill  passed  by  the  Lords  to  abolish  the  French 
phraseology."  De  Franqueville,  Gmivernement  et  Parlement  Britan- 
niques,  vol.  i.,  p.  279. 


SURVIVAL  OF  INSTITUTIONS  311 

same  costume  as  that  once  worn  by  the  body-guard 
of  Henry  YIU 

Two  similar  examples  may  be  added : — 
(a)  At  the  coronation  of  English  sovereigns  two 
gentlemen  of  the  Privy  Council,  chosen  "  on  account 
of  their  appearance,"  and  created  knights  for  the 
occasion,  are  appointed  by  the  Lord  Chamberlain  to 
represent  the  Dukes  of  Aquitaine  and  Normandy. 

(h)  The  First  Officer  of  the  Crown  was  formerly 
the  Lord  High  Steward,  which  title  having,  in  the 
course  of  time,  become  purely  honorary,  was 
hereditary  in  the  family  of  the  Earls  of  Leicester. 
The  post  is  now  in  abeyance,  but,  on  the  coronation 
of  a  sovereign,  or,  on  the  occasion  of  a  peer  being 
placed  upon  his  trial,  this  dignity  is  conferred  upon 
some  important  person  nominated  solely  for  the 
occasion.  It  is  not  only  in  ceremonial — which, 
according  to  Viollet,  is  the  museum  of  history — that 
reduced  institutions,  which  are  completely  useless, 
are  tenaciously  maintained  ;  it  is  the  same  with 
judicial  and  religious  institutions.  A  few  examples 
will  suffice  to  show  that  this  is  so  : — 

(a)  In  its  primitive  form,  the  Assembly  of  the 
People  included  the  whole  army,  and  was 
necessarily  held  in  some  large  open  space. 
The  custom  survived  the  necessity  of  the 
choice  of  some  such  spot,  and  up  to  the 
sixteenth  century,  whenever  a  new  Emperor 
was   proclaimed   in   Germany,   it   was   the 

^  H.  Spencer,  Morals^  Science  and  Art^ 


312    CAUSES  OF  DEGENERATIVE  EVOLUTION 

custom  for  the  electors  to  proceed  to  some 
mountain  for  the  purpose,  probably  because 
it  was  the  custom  in  former  days  to  hold  a 
general  meeting  there,  before  the  election. 

In  Iceland,  the  althing — consisting  of 
two  chambers  now — was  formerly  one  large 
assembly  held  in  the  open  air  upon  the 
Logberg  (the  Mountain  of  the  Law)  near  to 
the  Lake  of  Thingvellir.  In  the  Eepublic 
of  Andora,  criminal  sentences  are  still 
pronounced  with  great  solemnity  from  the 
Market-Place.^ 
(b)  Among  the  Ossetes,  where  the  family  com- 
munity still  flourishes,  the  only  inalienable 
and  unsaleable  property  is  not  the  real 
property,  but  certain  personal  possessions, 
such  as  the  great  cooking-pot  with  the 
chain  by  which  it  is  hung  over  the  fire. 
"  At  first,"  says  Kowalewsky,  "  this  may 
appear  a  strange  custom,  but  it  must  be 
remembered  that  possessions  of  that  kind 
were  of  equal  importance  to  the  Ossetic 
*  family '  as  were  their  tombs  to  the 
ancient  Greeks  and  Eomans,  which  ex- 
plains    how     any     infringement     of     the 


^  In  Montenegro,  it  was  the  custom,  till  recently,  for  the  Prince 
himself  to  render  justice,  sitting  under  a  tree  in  front  of  his  palace 
at  Cettigne.  This  mode  of  jurisdiction,  which  was  probably  a 
survival  of  the  old  system  of  which  we  have  been  speaking,  has  not 
completely  disappeared. 


SUKVrVAL  OF  INSTITUTIONS  313 

custom  is  regarded  as  an  infamy  by  the 
Ossetes."  ^ 

We  now  come  to  cases  of  purely  religious 
survival,  which  offer  the  strongest  resistance  to 
the  inroads  of  change.  Spencer  instances  the 
custom  of  circumcising  with  a  knife  made  of 
flint,  and  the  vestiges  remaining  in  Catholic 
worship  of  former  primitive  religions.  The 
Eucharist,  as  we  have  already  pointed  out,  is 
reminiscent  of  real  sacrifices,  and  the  symbolic 
representation  by  a  dove  of  the  Holy  Ghost  is 
only  a  rudimentary  form  of  zoolatry.^ 

In  Belgium  there  are  still  traces  of  the  old 
custom  of  sacrificing  an  animal  upon  the  com- 
pletion of  a  new  building,  with  the  idea  that  the 
animal's  spirit  will  protect  the  edifice  from  harm,^ 
and  if  the  observer  of  the  following  facts  is  correct 
in  his  interpretation  of  them,  there  also  remain  in 
Belgium  traces  of  the  ancient  sacrifices  to  the 
genius  of  the  earth  : — 

"  It  is  the  custom,  round  about  Florenville 
(in  the  Belgian  Ardennes),  to  offer  a  sacrifice  to 
the  presiding  genius  of  the  road  upon  the  con- 
struction of  a  new  road  or  railway.  It  is  usually 
a  fowl,  or  a  rabbit,  or  even  a  calf  which  is  sacri- 
ficed. ...  In  some  parts  of  Luxembourg  animals 

^  Kowalevsky,  Droit  coutumier  Ossetien,  p.  105. 

2  Spencer,  Principles  of  Sociology. 

3  Folklore,  Wallon,  No.  1526,  p.  115  {Bulletin  de  Folklore,  ii., 
177). 


314         CAUSES  OF  DEGENERATIVE  EVOLUTION 

are  also  offered  in  sacrifice  to  the  genius  supposed 
to  preside  over  a  newly-purchased  field,  with  a 
view  to  ensuring  abundant  crops.^ 

Eemains,  either  undoubted  ^  or  only  probable,^ 
of  phallic  worship,  are  scattered  throughout  Europe. 

In  Brittany*  and  in  Belgium,^  for  instance, 
strange  old  customs  still  exist  showing  that  here, 

^  Rivue  des  traditions  pojnclaires,  1893,  p.  394. 

^  Th.  Volkov,  Eiies  et  usages  nuptiaux  en  Ukraine  {VAnthro- 
pologie,  1891,  p.  167). 

Only  a  short  time  ago  it  was  the  custom  in  Tver,  on  the  day 
dedicated  to  Yarilo  (the  phallic  God  of  Spring),  for  the  parents 
of  young  daughters  to  send  them  to  join  in  games  similar  to 
those  of  the  ancient  Slavs,  with  a  view  to  their  getting  married. 

^  Note  sur  un  vestige  du  culte  de  la  terre  mere  (phallism)  en 
Provence^  by  Berenger-Feraud  {Rdime  d'Anthropologie,  1888,  p. 
563). 

"At  Luc,  in  Provence,  upon  the  1st  of  May,  which  is  a  country 
holiday,  the  young  girls  proceeded  to  a  place  where  two  roads  met. 
Hero  they  assembled  around  an  olive  tree,  and  after  each  dance 
they  struck  the  olive  tree  three  times  with  their  backs. 

"  This  fete,  a  survival  of  the  floral  fetes  of  the  month  of  May 
which  are  still  celebrated  in  Provence  and  Italy,  continued  to  be 
lield  until  quite  recently,  and  appears  to  have  been  a  lingering 
vestige  of  the  ancient  worship  of  creative  Nature,  Mother-earth 
—  in  short,  of  phallic  worship. 

*'  The  three  knocks  given  by  the  young  girls  to  the  tree  trunk 
is  a  survival  of  the  ancient  virginal  sacrifice  to  the  phallic 
emblem.  The  original  meaning  was  not  quite  lost,  for  the  Pro- 
ven9als  still  realized,  though  vaguely,  that  the  three  knocks  were 
somehow  connected  with  the  idea  of  marriage." 

^  ' '  Les  M^galithes  de  Locmariaquer  et  de  Carnac,  et  les  amours,  by 
Bonnemere  {R4tme  des  traditions  populaires,  1894,  p.  123).  **In 
former  days  it  was  the  custom  for  all  the  young  women  who  wished 
to  get  maiTied  to  climb  (on  the  night  of  May  1),  to  the  top  of  the 
great  menhir  where  they  lifted  up  their  clothing  that  their  bodies 


SURVIVAL  OF  INSTITUTIONS  815 

as  elsewhere,  a  belief  in  the  influence  of  fetiches 
once  prevailed,  and  particularly  in  the  form  of 
megalithic  monuments  relating  to  the  fecundity  of 
women. 

The  custom  still  prevalent  in  the  African  Congo, 
of  driving  a  nail  into  a  fetich,  with  the  view  of 
reminding  it  of  a  request,  has  not  disappeared  from 
Europe.  The  young  men  of  Couvin  (Namur)  still 
stick  pins  into  the  wooden  saints  of  the  little 
chapels  round  about  in  order  to  draw  a  lucky 
number  in  the  military  lottery,  and  young  girls  in 
Brittany    do    the    same    with    a   view    to    getting 

might  come  in  direct  contact  with  the  stone,  and  then  slid  from 
the  top  to  the  bottom." 

At  Carnac,  young  girls  wishing  to  marry  undressed  completely 
upon  the  same  night,  and  proceeded  to  rub  their  abdomen  against 
a  special  menhir.  In  that  part  of  Brittany,  where  only  French  is 
spoken,  similar  customs  have  equally  prevailed. 

°  Fete  de  Notre-Dame  de  liide-cul  (Jules  Lemoine,  in  the  journal 
Le  petit  bleu,  of  October  18,  1896). 

Similar  to  the  above  Breton  custom  is  that  of  sliding  down  the 
Rocher  Ride-cul,  which  is  situated  near  to  Landelies,  in  the  valley 
of  the  Sambre.  Here,  as  in  many  other  places,  old  customs  have 
been  Christianized,  and  a  Christian  chapel  now  stands  close  to  the 
ancient  shrine.  Young  people  of  both  sexes  used  to  seat  them- 
selves upon  the  top  of  the  stone  upon  little  fagots  of  boxwood 
gathered  in  the  neighbourhood,  and  then  slide  down  to  the  bottom. 

According  to  the  old  saying  an  upset  meant  waiting  ;  an  em- 
brace signified  mutual  affection  ;  a  collision,  indifference  ;  and  an 
embrace  followed  by  rolling  over  indicated  matrimonial  suitability. 
Similar  customs  prevailed  at  Trou-deux-Trous,  situated  near  to 
the  Rocher  Ride-cul.  These  two  megalithic  temples  disappeared 
about  forty  years  ago,  their  materials  have  been  used  as  fluxes  in 
smelting  works. 


316         CAUSES  OF  DEGENERATIVE  EVOLUTION 

married,^  and  in  Belgium,^  as  in  Brittany,^  if  the 
image  prove  too  hard,  the  reminding  pin  is  stuck 
instead  into  a  fissure,  or  into  the  door  of  the  niche 
containing  the  image.  Besides  these  mere  vestiges 
of  pre-historic  customs  and  belief,  by  referring  to 
certain  illustrated  documents  bearing  upon  the 
subject,  it  would  be  easy  to  work  out  the  connec- 
tion between  the  worship  of  saints  which  is  pre- 
valent in  Belgium  in  the  present  day,  and  the 
pagan  worship  of  the  ancient  Celts  and  Germans.* 

^Bulletin  de  Folklore,  i.,  250-251. 

2  Ibidem. 

^  The  menhir  of  the  Pierre-Frite  in  the  valley  of  Lunain.  In 
nearly  every  hole  or  fissure  of  this  monument,  a  nail  or  pin  has 
been  stuck  by  the  young  people  of  that  part  in  the  belief  that  it 
will  ensure  them  a  speedy  marriage.  {Revue  des  traditions  popu- 
laires,  1893,  p.  448.) 

''See  for  Saint  Eloi,  Mdusine,  viii.,  122-132  ;  for  Saint  Martin, 
Bulletin  de  Folklore,  i,,  309-315  ;  for  Saint  Hubert,  Gaidez,  La 
Bage  et  Saint  Hubert. 


PAET  III 

RESUME    AND    CONCLUSIONS 

When  an  institution  or  an  organ  ceases  to  be 
functional  or  in  any  way  useful,  it  very  soon 
disappears  altogether.  If,  as  happens  in  some 
exceptional  cases,  it  persists,  it  is  because  neither 
of  the  chief  factors  in  causing  atrophy,  variability 
or  selection,  have  intervened. 

Sometimes  the  vestiges  are  of  too  insignificant 
a  nature  to  call  for  their  removal  by  either 
artificial  or  natural  selection,  and  sometimes  their 
existence  is  ensured  by  the  lack  of  variability,  as 
in  the  case  of  the  persistence  of  flowers  in  plants 
which  multiply  asexually.  This  absence  of  varia- 
tion occurs  equally  in  the  social  domain,  especially 
in  matters  connected  with  religion,  wherein  ancient 
customs  are  credited  with  a  divine  origin.  Eeligions 
may  pass  away,  philosophies  may  be  transformed, 
and  old  beliefs  cease  to  prevail,  but  the  remnants 
of  old  creeds,  conveyed  by  popular  tradition 
through  the  centuries,  defy  destruction  by  modern 
innovations. 

The  ancient  winter  festival,  on  which  day  the 

3*7 


318    CAUSES  OF  DEGENERATIVE  EVOLUTION 

dead  were  supposed  to  leave  their  graves  and  join 
the  living  in  a  feast  around  the  family  hearth,  is 
still  celebrated  in  the  keeping  of  Christmas  and  in 
the  various  customary  practices  on  the  first  two 
days  of  November. 

The  May-Day  festivals — pagan  festivals  held  in 
honour  of  vegetable  and  human  fecundity — are 
still  held  in  their  early  form  round  about 
Locmariaquer  and  in  the  village  of  Campine. 
Traces  also  remain  in  the  picking  and  wearing 
of  flowers  on  the  1st  of  May,  and  the  same  day 
is  selected  by  the  socialists  for  the  celebration  of 
their  near  approach  to  a  life  under  freer  and 
happier  conditions. 

This  survival  of  festivals,  customs  and  traditions, 
while  the  religions  and  civilizations  which  produced 
them  have  passed  away,  is  the  principal  link  which 
connects  us  with  bygone  generations. 

"  Their  value  lies,"  says  Houzeau  in  his  Etude  de 
la  Nature,  "  in  the  establishment  of  a  chain  between 
successive  generations.  The  memory  of  an  indivi- 
dual may  be  regarded  as  constituting  his  personality. 
Take  from  him  the  memory  of  his  past,  and  he  is  left 
at  a  point  in  time  wherein  there  is  no  stability  and 
complete  isolation.  To  be  himself,  a  man  requires 
not  only  his  recollections,  but  a  knowledge  of  his  past 
habits  and  traditions.  When  a  savage  is  removed 
from  his  fellows  and  transported  to  new  surround- 
ings in  a  distant  country,  he  loses  all  knowledge  of 
his  former  condition.      Society  itself,  made  up  as  it 


SUMMARY  AND  CONCLUSIONS  319 

is  of  customs  and  prejudices,  constitutes  history. 
The  mirror  of  the  past  is  exhibited  in  the  conscious- 
ness of  the  collective  individual  which  is  called 
a  nation.  What  link  shall  we  have  with  former 
generations  if  not  a  heritage  of  their  ideas — i.e.  of 
their  discoveries  and  their  mistakes  ?  Nations, 
like  individuals,  are  continually  modifying  this 
inherited  legacy,  but,  like  the  individual,  they 
cannot  get  away  from  it  without  breaking  the 
thread  which  has  made  them  themselves." 


GENERAL  CONCLUSIONS 


All  evolution  is  at  once  progressive  and  retro- 
gressive. 

All  modifications  of  organs  and  institutions  are 
attended  by  retrogression.  This  occurs  equally 
in  the  modifications  of  organisms  and  of  societies. 
All  existing  forms,  whether  organic  or  social,  have 
undergone  certain  modifications,  and,  as  a  result, 
have  lost  some  parts  of  their  structure.  This 
universality  of  degenerative  evolution  may  be 
proved  either  by  the  comparative  method,  or  by 
showing  that  all  organisms  contain  rudimentary 
organs,  and  that  all  societies  contain  survivals. 

II 

Degenerative  evolution  follows  no  definite  path, 
and  can  in  no  way  be  regarded  as  constituting  a 
return  to  the  primitive  condition. 

In  some  cases — when  one  cause  of  dissolution 
equally  and  simultaneously  affects  all  the  parts  of  an 
institution  or  an  organism — the  most  complicated 
and  delicate  structures  are  the  first  to  disappear ; 
but  it  must  not  be  taken  as  a  general  principle 
that  the  most  complicated  structures  are  necessarily 
320 


GENERAL  CONCLUSIONS  321 

the  most  recent,  and  that  consequently  degeneration 
always  retraces  the  path  of  progress.  Evolution 
is  irreversible,  and  accordingly,  with  a  few  more 
or  less  obvious  exceptions,  we  draw  the  following 
conclusions : — 

1.  That  an  institution  or  an  organ  which   has 

once  disappeared  never  reappears. 

2.  That  an   institution  or   organ   once   reduced 

to  the  condition  of  a  vestige  cannot  be 
re-established  and  resume  its  former  func- 
tions. 

3.  Neither  can  they  assume  fresh  functions. 


III 

Degenerative  evolution  is  brought  about  by  a 
limitation  in  means  of  subsistence  —  either  in 
nutriment,  capital  or  labour.  In  biology  the  prin- 
cipal if  not  the  sole  agents  in  its  accomplishment 
are  the  struggle  for  existence  between  the  various 
organs,  and  the  struggle  for  existence  between  the 
various  organisms. 

In  sociology  it  is  artificial  selection  which  is  the 
dominating  agent,  and  natural  selection  plays  only  a 
secondary  part. 

The  occasional  causes  of  degenerative  evolution 
are  inutility  of  function,  insufficiency  of  nutri- 
ment or  resource,  and  (in  biology  only)  lack  of 
space. 

X 


/ 


322  GENERAL  CONCLUSIONS 

An  institution  or  an  organ  which  has  ceased  to 
be  functional,  and  has  also  ceased  to  be  useful  either 
directly  or  indirectly,  continues  to  exist  if  neither 
variability  or  selection  intervene. 


TCRNBUI-L    AND   SPEARS,    PRINTERS.   EDIXBURGH. 


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H.  Waij.is  Kkw,  F.Z.S.  With  Preface  by  A.  R.  Waltack,  P.R.S.,  and 
Illustrations. 

LXXVI.  RACE  and  LANGUAGE.     By  Andre  Lefevrr,  Professor 

in  the  Anthropological  School,  Paris. 

LXXV^II.  The  ORIGIN  of  PLANT   STRUCTURES  by   SBLP- 

ADAPTATION  TO  THE  ENVIRONMENT.  By  Rev.  G.  Henslow, 
M.A.,  F.L.S,,  F.G.S.,  &c.,  author  of  'The  Origin  of  Floral  Structures,'  &c. 

LXXVIII.  ICE- WORK    PRESENT   and   PAfaT.      By   Rev.  T.  G. 

BoNNRY,  D.Sc,  LL.D.,  F.R.S.,  &c.,  Professor  of  Geology  at  University 
College,  Loudon  ;  Fellow  of  St.  John's  College,  Cambridge.  Second 
Edition 

LXXTX.  A     CONTRIBUTION    to     our     KNOWLEDGE     of 

SEEDLINGS.    By  Lord  Avkrury. 

LXXX.  The  ART  of  MUSIC.  By  Sir  C.  Hubert  H.  Parry,  Mus.  Doc. 

LXXXI.  The  POLAR  AURORA.    By  Alfred  Angot.  Illustrated. 

LXXXII.  WHAT  is  ELECTRICITY  P  By  J.  Trowbridge.  Illustrated. 

LXXXIII.  MEMORY.  By  F.  W.  Ed  ridge-Green,  M.D.  With 
Frontispiece. 

LXXXIV.  The    ELEMENTS    of    HYPNOTISM.      By  R.   Harry 

Vincent.    With  Diagrams.    Second  Edition. 

LXXXV.  SEISMOLOGY.  By  John  Milne,  F.R.S.,  RG.S.,  &c., 
Author  of  '  Earthquakes.'    With  53  Figures. 

LXXXVI  On  BUDS  and  STIPULES.  By  Lord  Avebury, 
F.R.S.,  D.C.L.,  LL.D.  With  4  Coloured  Plates  and  340  Figures  in  the 
Text. 

LXXX  VII.  EVOLUTION  by  ATROPHY,  in  Biology  and  Sociology. 

By  Jkan  Demoor,  Jkan  Mapsaht,  and  Emii,k  Vandbrvrldk.  Translated 
by  Mrs.  Cualmkhs  MrrciiKr^u    With  84  Figures. 

LXXXVIII.  VARIATION     in     ANIMA.LS     and     PLANTS.     By 

H.  M.  Vkhnox,  M.A.,  M.D. 

LXXXIX.  THE  MIND  AND  THE  BRAIN.  By  Alfred  Binet, 
Directeur  in  Laboratoire  de  Psychologie  h  la  Sorbonne. 


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