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UNIVERiSITY  OF   PITTSBURGH 

DoM. 


Jjarlington  .Alemorial  Library 


THE   DESCEE'T    OF   MAN, 


SELECTION  IN  RELATION  TO  SEX. 


THE 


DESCEN^T   OF   MAI^, 


AND 


SELECTION  IN  RELATION  TO  SEX. 


BY 

CHAELES  DAEWIN,   M.  A. 

FELLOW  OF  THE   KOYAL   SOCIETY,    ETC. 


m\t^  lllastratiflits; 


NEW  EDITION,  REVISED   AND   AUGMENTED. 


COilPLETE  IN  ONE  VOLUME. 


NEW    YOKE  : 
D.     APPLETON     AND     COMPANY, 

549    AND   551    BROADWAY. 

1875. 


X. 


.^-3- 


^ 


PEEFACE  TO  THE  SECOND  EDITION. 


During  the  successive  reprints  of  the  first  edition  of  this  work, 
published  in  1871,  I  was  able  to  introduce  several  important 
corrections;  and  now  that  more  time  has  elapsed,  I  have 
endeavoured  to  profit  by  the  fiery  ordeal  through  which  the 
book  has  passed,  and  have  taken  advantage  of  all  the  criticisms 
which  seem  to  me  sound.  I  am  also  greatly  indebted  to  a  large 
number  of  correspondents  for  the  communication  of  a  surprising 
number  of  new  facts  and  remarks.  These  have  been  so  numerous, 
that  I  have  been  able  to  use  only  the  more  important  ones ;  and 
of  these,  as  well  as  of  the  more  important  corrections,  I  will 
append  a  list.  Some  new  illustrations  have  been  introduced, 
and  four  of  the  old  drawings  have  been  replaced  by  better  ones, 
done  from  life  by  Mr.  T.  W.  Wood.  I  must  especially  call 
attention  to  some  observations  which  I  owe  to  the  kindness  of 
Prof.  Huxley  (given  as  a  supplement  at  the  end  of  Part  I.),  on 
the  nature  of  the  differences  between  the  brains  of  man  and  the 
higher  apes.  I  have  been  particularly  glad  to  give  these  obser- 
vations, because  during  the  last  few  years  several  memoirs  on  the 
subject  have  appeared  on  the  Continent,  and  their  importance 
has  been,  in  some  cases,  greatly  exaggerated  by  popular  writers. 
I  may  take  this  opportunity  of  remarking  that  my  critics 
frequently  assume  that  I  attribute  all  changes  of  corporeal 
structure  and  mental  power  exclusively  to  the  natural  selection 
of  such  variations  as  are  often  called  spontaneous;  whereas, 
even  in  the  first  edition  of  the  '  Origin  of  Species,'  I  distinctly 
stated  that  great  weight  must  be  attributed  to  the  inherited 
effects  of  use  and  disuse,  with  respect  both  to  the  body  and 
mind.  I  also  attributed  some  amount  of  modification  to  the 
direct  and  prolonged  action  cf  changed  conditions  of  life.  Some 
aUowance.  too,  must  be    made    for   occasional    reversions  of 


vi  Preface  to  the  Second  Edition. 

structure ;  nor  must  we  forget  what  I  liaTe  called  "  correlated  *' 
growth,  meaning,  thereby,  that  various  parts  of  the  organisation 
are  in  some  unknown  manner  so  connected,  that  when  one  part 
varies,  so  do  others;  and  if  variations  in  the  one  are  accu- 
mulated by  selection,  other  parts  will  be  modified.  Again,  it 
has  been  said  by  several  critics,  that  when  I  found  that  many 
details  of  structure  in  man  could  not  be  explained  through 
natural  selection,  I  invented  sexual  selection ;  I  gave,  however, 
a  tolerably  clear  sketch  of  this  principle  in  the  first  edition  ol 
the  '  Origin  of  Species,'  and  I  there  stated  that  it  was  applicable 
to  man.  This  subject  of  sexual  selection  has  been  treated  at 
full  length  in  the  present  work,  simply  because  an  opportunity  was 
here  first  afforded  me.  I  have  been  struck  with  the  likeness  oi 
many  of  the  half-favourable  criticisms  on  sexual  selection,  with 
those  which  appeared  at  first  on  natural  selection;  such  as, 
that  it  would  exj)lain  some  few  details,  but  certainly  was  not 
apphcable  to  the  extent  to  which  I  have  employed  it.  My 
conviction  of  the  power  of  sexual  selection  remains  unshaken ; 
but  it  is  probable,  or  almost  certain,  that  several  of  my  con- 
clusions will  hereafter  be  found  erroneous ;  this  can  hardly  fail 
to  be  the  case  in  the  first  treatment  of  a  subject.  When 
naturalists  have  become  familiar  with  the  jdea  of  sexual  selection^ 
it  will,  as  I  believe,  be  much  more  largely  accepted;  and  it 
has  already  been  fully  and  favourably  received  by  several  cajDable 
judges. 


Down,  Beckexham,  Kent, 
September  1874. 


TABLE 


PEINCIPAL  ADDITIONS  AND  COEEECTIONS  TO  THE 
PRESENT  EDITION. 


First 

Edition. 

Trcsent 

Edition. 

Vol.  I. 

Page 

22 

26 

27,  note. 

29 

32,  note. 

40 
42 

44 

47 
50 
53 

58 

64 

78 
79 
81 

90,  note. 

91 

94 

97 
112 
122 

124,  note. 

125,  note. 

128-9 


146 


Page 
15-17 

19 

20,  note. 
23 

24,  note, 

70 
71 

72  3 

75 
80 
83 

88-9 

92 

104 

104 

105-6 

ril2-113,] 

\    note.     ] 

114 

117,  note. 

120,  note. 
28 
35-G 

39,  note. 
3G-8,  note. 

41-2 


55,  note. 


f  Discussion  on  the  rudimentary  points  in  the 
\     human  ear  revised. 
Cases  of  men  born  with  hairy  bodies. 
Muntegazza  on  the  last  molar  tooth  in  man. 
Tiie  rudiments  of  a  tail  in  man. 

IBianconi  on  homologous  structures,  as  ex- 
plained by  adaptation  on  mcciianieal 
principles. 

Intelligence  in  a  biiboon. 

Sense  of  humour  in  dogs. 
(Further   facts    on    imitation    in     man    and 
\     animals. 

Reasoning  power  in  the  lower  animals. 

Acquisition  of  experience  by  animals. 

Power  of  abstraction  in  animals. 
JPower  of  forming   concepts   in   relation   to 
\     language.  .  ,         , 

(Pleasure  from  certain  sounds,  colours,  and 
\     forms. 

Fidelity  in  (he  elephant. 

Galton  on  gregariousncss  of  cattle. 

Parental  atfection. 

Persistence  of  enmity  and  hatred. 

^Nature  and  strength  of  shame,  regret,  and 
\     remorse. 

Snicide  amongst  savages. 
The  motives  of  conduct. 
Selection,  as  applied  to  primeval  man. 
Resemblances  between  idiots  and  animals. 
Division  of  the  malar  bone. 
Supernumerary  mamnuo  and  digits. 
Further  cases  of  mutcles  projier  to  aninials 

appiaring  in  man. 
Broca  :  average  oftpacity  of  slaill  diniinislied 

by  the  preservation  of  the  inferior  members 

of  society. 


viii  Table  of  the  Principal  A  dditions  and 


First 
Edition. 


Vol. 


Page 
149 

150 

169 

180 

193 

208.  note. 
209 

239 
245 
250 
256 

275-6 
290 


Present 
Edition. 


Page 
57 

58-9 

134-5 
143 

151 

161,  note. 
163 

188-190 
195-6 
199-206 
209-210 

224-5 
235 


301 
314 

243-4 
254 

315 

255-6 

327 

264 

338 

272 

339 

273 

345 

277 

349 

280 

350 

281 

351 

282 

854 

284-5 

359 

288,  note 

366 

292-3 

387 

308 

397 

315 

401 

319 

412 

324-5 

117 

326 

I  Belt  on  advantages  to  man  from  his  hair- 

\     lessness. 

(Disappearance  of  the  tail  in  man  and  certain 

\    monkeys. 

("Injurious  forms    of    selection    in    civilised 

\     nations. 

(Indolence  of  man,  when  free  from  a  struggle 

\     for  existence, 

jGorilla  protecting  himself  from  rain  with  his 

\     hands. 

Hermaphroditism  in  fish. 

Eudimentary  mammae  in  male  mammals. 
(Changed  conditions  lessen  fertility  and  cause 
\     ill-health  amongst  savages. 
JDarkness  of  skin  a  protection  against  the 
\     sun. 

jNote  by  Professor  Huxley  on  the  develop- 
\     ment  of  the  brain  in  man  and  apes. 
(Special  organs  of  male  parasitic  worms  for 
\     holding  the  female. 

Greater  variability  of  male  than  female; 
direct  action  of  the  environment  in  causing 
differences  between  the  sexes. 

Period  of  development  of  protuberances 
on  birds'  htads  determines  their  trans- 
mission to  one  or  both  sexes. 

Causes  of  excess  of  male  births. 

Proportion  of  the  sexes  in  the  bee  family. 
(Excess  of  males  perhaps  sometimes  deter- 
(    mined  by  selection. 

Bright  colours  of  lowly  organised  animals. 

Sexual  selection  amongst  spiders. 

Cause  of  smallness  of  male  spiders. 

Use  of  phosphorescence  of  the  glow-worm. 

The  humming  noises  of  flies. 

Use  of  bright  colours  to  Hemiptera  (bugs). 

Musical  apparatus  of  Homoptera. 

(Development   of  stridulating   apparatus   in 
\     Orthoptera. 

(Hermann  Miiller   on   sexual   differences  of 
\    bees. 

Sounds  produced  by  moths. 

Display  of  beauty  by  butterflies. 

(Female  butterflies,  taking  the  more  active 
\     part  in  courtship,  brighter  than  their  males. 

(Further  cases  of  mimicry  in  butterflies  and 
\     moths. 

(Cause  of  bright  and  diversified  colours  of 
\    caterpillars. 


Corrections  to  tJie  Present  Edition. 


IX 


First 
Edition. 


Vol.  II. 


Pape 
2 

M 

23 

26 
30 
82 
3G 
72 

91 

108 
118 

120 

124 

147-150 

152 

157 

232 

^47 

248 
256 

260 

266 
286 
299 

316 

337 

356 

359  et  seq. 

373 

380 


rropont 
Kdition. 


Page 
33  i 

341 

347 
349 
352 
353 
357 
383 

398 

411 
417 

419 

423 

438-441 

443 

446 

495-6 

505 

506 
513-514 

516 

521 
534 
542-3 

556 

572 

586 

588  et  seq. 

59S-9 

603 


Brush-liko  scales  of  male  Mall(.)tu.s. 
|Furtlier  facts  on  courtship  of  tishcs,  and  the 
\     spawning  of  Macropus. 

Dufosse'  on  the  sounds  made  by  fl.slics. 

Belt  on  a  frog  protected  by  bright  colouring. 

Further  facts  on  mental  powers  of  snakes. 

Sounds  produced  by  snakes ;  the  rattlesnake. 

Cond)ats  of  Chameleons. 

Marshall  on  protuberances  on  birds'  heads. 
^Further    facts    on    display    by   the    Argus 
\     pheasant. 

Attachment  between  paired  birds. 

Female  pigeon  rejecting  certain  males. 
jAlbino  birds  not  finding  partners,  in  a  state 
\     of  nature. 

Direct  action  of  climate  on  birds'  colours. 
jFurther   facts  on  the  ocelli   in  the  Argus 
\     pheasant. 

Display  by  humming-birds  in  courtship. 
jCases  with  pigeons  of  colour  transmitted  to 
\     one  sex  alone. 

I  Taste  for  the  beautiful  permament  enough 
to  allow  of  sexual  selection  with  the  lower 
animals. 
I  Horns    of    sheep    originally    a    masculine 
\     character. 

Castration  afiecting  horns  of  animals. 

Prong-horned  variety  of  Cervus  vir'jimaiius. 
|Ilelative  sizes  of  male  and  female  whales  aud 
\     seals. 

Absence  of  tusks  in  male  miocene  pigs. 

Dobson  on  sexual  differences  of  bats. 

Eeekson  advantage  from  peculiar  colouring. 
|Difterence  of  comj)lexion  in  men  and  women 
\     of  an  African  tribe. 

Speech  subsequent  to  singing, 
|Schopenhauer  on  importance  of  courtship  to 
\     mankind. 

^Revision  of  discussion  on  communal  marriages 
\     and  promiscuity. 

|Fower    of    choice   of  woman    in   marriage, 
\     amongst  savages. 

|Long-continued  habit  of  plucking  out  hairs 
\     may  produce  an  inherited  cfiect. 


CONTENTS. 

Inteoduotion Pages  1-4 

PART  I. 
THE  DESCENT  OR  ORIGIN  OF  MAN. 


CHAPTER  I. 

The  Evidence  of  the  Descent  of  Man  from  some  Lower  Form. 

PAGE 

Nature 'of  the  evidence  bearing  on  the  origin  of  man — Homologous 
structures  in  man  and  the  lower  animals — IMiscellaneous  points 
of  correspondence  —  Development  —  Rudimentary  structures, 
muscles,  sense-organs,  hair,  bones,  reproductive  organs,  Arc. — 
The  bearing  of  these  three  great  classes  of  facts  on  the  origin  of 
man      ........••  ^ 

CHAPTER  11. 

On  the  Manner  of  Development  of  Man  fkom  some  Lower 

Form. 

Variability  of  body  and  mind  in  man— Inheritance— Causes  of 
variability— Laws  of  variation  the  t-ame  in  man  as  in  the  lower 
animals— Direct  action  of  the  conditions  of  life— Etl'ects  of  the 
increased  use  and  disuse  of  parts— Arrested  development— Re- 
version—Cnrrc  luted  variation— Rate  of  Increase  — Checks  to 
increase— Natural  selection- Man  the  most  dominant  animal  in 
tlie  world— Im})ortance  of  his  corporeal  structure— Tlie  causes 
which  have  kd  to  his  becoming  erect— Consequent  clianges  of 
structure— Decrease  in  size  of  the  canine  teeth— Increased  size 
and  altered  shape  of  the  skull— Nakedness— Absence  of  a  tail- 
Defenceless  condition  of  man 2G 

CHAPTER  III. 

Comparison  of  the  Mental  Powers  of  Man  and  the  Lowkr 
Animals. 

The  difftrence  in  mental  power  between  the  highest  ape  and  the 
lowest  savage,  immense— Certain  instincts  in  common— The 
emotions— Curiosity— Imitation— Att(ntion  — Memory— Imagi- 
nation—Reason— Progressive  improvement- Tools  and  weapons 


xii  Contents. 


PAGE 

used  by  animals — Abstraction,  Sell-consciousness — Language 
— Sense  of  beauty — Belief  in  God,  spiritual  agencies,  super- 
stitions    .........  65 

CHAPTEll  IV. 

COMPAKISON    OP   THE   MeNTAL   PoWERS   OF   MaN   AND   THE   LoWER 

Andials — continued. 

The  moral  sense — Fundamental  proposition-^The  qualities  of 
social  animals — Origin  of  sociability*— Struggle  between  opposed 
instincts — Man  a  social  animal — The  more  endm-ing  social  in- 
stincts conquer  other  less  persistent  instincts — The  social  virtues 
alone  regarded  by  savages — The  self-regarding  virtues  acquired 
at  a  later  stage  of  development — The  importance  of  the  judg- 
ment of  the  members  of  the  same  community  on  conduct — 
Transmission  of  moral  tendencies — Summary        .  .  97 

CHAPTEK  V. 

On  the  Development  of  the  Intellectual  and  Moral 
Faculties  during  Primeval  and  Civilised  Times. 

Advancement  of  the  intellectual  powers  through  natural  selec- 
tion— Importance  of  imitation — Social  and  moral  faculties — 
Their  development  within  the  limits  of  the  same  tribe — Natural 
selection  as  affecting  civilised  nations — Evidence  that  civilised 
nations  were  once  barbarous  .  ....  127 

CHAPTER  VI. 

On  the  Affinities  and  Genealogy  of  Man. 

Position  of  man  in  the  animal  series — The  natural  system  genea- 
logical— Adaptive  characters  of  slight  value — Various  small 
points  of  resemblance  between  man  and  the  Quadrumana — 
Bank  of  man  in  the  natural  system — Birthplace  and  antiquity 
of  man — Absence  of  fossil  connecting-links — Lower  stages  in 
the  genealogy  of  man,  as  inferred,  tirstly  from  his  aiSnities  and 
secondly  from  his  structure — Early  androgynous  condition  of 
the  Vertebrata — Conclusion         .  .          .  .  .  14G 

CHAPTER  VII. 
On  the  Eaces  of  Man. 

The  nature  and  value  of  specific  characters — Application  to  the 
races  of  man — Arguments  in  favour  of,  and  opposed  to,  ranking 
the  so-called  races  of  man  as  distinct  species — Sub-species — 
Monogenists    and    polygenists — Convergence    of    character  — 


Contents.  xiii 


Numerous  points  of  resemblance  in  body  and  mind  bet\Ycen  the 
most  distinct  races  of  man — The  state  of  man  when  he  lirst 
spread  over  the  earth — Each  race  not  descended  from  a  single 
pair — The  extinction  of  races — The  formation  of  races — The 
eftects  of  crossing — Slight  influence  of  the  direct  action  of  the 
conditions  of  life — Slight  or  no  influence  of  natural  selection — 
Sexual  selection         .......  166 


PAET  II. 
SEXUAL  SELECTION. 

CHAPTEE  VIII. 

Principles  of  Sexual  Selection. 

Secondary  sexual  characters — Sexual  selection — Manner  of  action 
— Excess  of  males  —  Polygamy  —  The  male  alone  generally 
modified  through  sexual  selection — Eagerness  of  the  male — 
Variability  of  the  male — Choice  exerted  by  the  female— Sexual 
compared  with  natural  selection — Inheritance  at  corresponding 
periods  of  life,  at  corresponding  seasons  of  the  year,  and  as 
limited  by  sex — Relations  between  the  several  forms  of  inheri- 
tance— Causes  why  one  sex  and  the  young  are  not  modititd 
through  sexual  selection — Supplement  on  the  proportional  num- 
bers of  the  two  sexes  throughout  the  animal  kingdom — The 
proportion  of  the  sexes  in  relation  to  natural  selection  .  .       207 

CHAPTER  IX. 

Secondary  Sexual  Characters  in  the  Lower  Classes  of 
THE  Animal  Kingdom. 

These  characters  absent  in  the  lowest  classes — Brilliant  colours — 
Mollusca — Annelids— Crustacea,  secondary  sexual  characters 
strongly  developed ;  dimorphism;  colour;  characters  not  ac- 
quired before  maturity — Spiders,  sexual  colours  of;  stridulation 
by  the  males — Myriapoda 2G0 

CHAPTER  X. 

Secondary  Sexual  Characters  of  Insects. 

Diversified  structures  possessed  by  the  males  for  seizing  the 
females— Differences  between  the  sexes,  of  which  the  mean- 
ing is  not  understood— Difference  in  size  between  the  sexes — 
Thysanura — Diptera— llemiptera — Homoptera,  musical  powers 


xiv  Contents. 


PAGS 

possessed  by  the  males  alone — Orthoptera,  musical  instruments 
of  the  males,  nuicli  diversified  in  structure;  pugnacity;  colours — 
Neuroptera  sexual  differences  in  colour — Hymenoptera,  pug- 
nacity and  colours — Culenptera,  colours ;  furnished  with  great 
horns,  apparently  as  an  ornament ;  battles ;  stridulating  organs 
generally  common  to  both  sexes  ....  274 

CHAPTER  XL 

Insects,  continued. — Order  Lepidoptera. 

(butterflies  and  moths.) 

Courtship  of  butterflies — Battles — Ticking  noise — Colours  common 
to  both  sexes,  or  more  brilliant  iti  the  males — Examples — Not 
due  to  the  direct  action  of  the  conditions  of  life  —  Colours 
adapted  for  protection — Colours  of  moths — Distday — Perceptive 
powers  of  the  Lepidoptera — Variability — Causes  of  the  difference 
in  colour  between  the  males  and  females — Mimicry,  female 
butterflies  more  brilliantly  coloured  than  the  males — Bright 
colours  of  caterpillars — Summary  and  concluding  remarks  on 
the  secondary  sexual  characters  of  insects — Birds  and  insects 
compared  ........  307 

CHAPTEK  XII. 

Secondary  Sexual  Charactters  of  Fishes,  Amphibians,  and 
Reptiles. 

Fishes:  Courtship  and  battles  of  the  males — Larger  size  of  the 
females — Males,  bright  colours  and  ornamental  appendages; 
other  strange  characters — Colours  and  appenilages  acquired  by 
the  males  during  the  breeding-season  alone — Fishes  with  both 
sexes  brilliantly  coloured — Protective  colours — The  less  con- 
spicuous colours  of  the  female  cannot  be  accounted  for  on  the 
principle  of  protection — Male  fishes  building  nests,  and  taking 
charge  of  the  ova  and  young.  Amphibians:  Differences  in 
structure  and  colour  between  the  sexes — Vocal  organs.  Rep- 
tiles :  Chelonians — Crocodiles — Snakes,  colours  m  some  cases 
protective  —  Lizards,  battles  of — Ornamental  appendages  — 
Strange  differences  in  structure  between  the  sexes — Colours — 
Sexual  difterences  almost  as  great  as  with  birds    .  .  330 

CHAPTER  XIII. 

Secondary  Sexual  Characters  of  Birds. 

Sexual  differences  —  Law  of  battle — Special  weapons  —  Vocal 
organs — Instrumental  music — Love-antics  and  dances — Deco- 
rations, permanent  and  seasonal — Double  and  single  annual 
moults — Display  of  ornaments  by  the  males  .         .  358 


Contents.  xv 


CHAPTEK  XIV. 

BiUDS — continued. 

PAOB 

Choice  exerted  by  the  female — Length  of  eourtsliip — Unpaired 
birds — jMental  qualities  and  ta^^te  for  the  beautiful — Preference 
or  antipatliy  shewn  by  tlie  female  for  particular  males — Vari- 
ability of  birds — Variations  sometimes  abrupt — Laws  of  varia- 
tion— Formation  of  ocelli— Gradations  of  character — Case  of 
Peacock,  Argus  phea.sant,  nnd  Uro^ticte        .  .          .  404 

CEL\PTER  XV. 

Birds — continued. 

Discussion  as  to  why  the  males  alone  of  some  species,  and  both 
sexes  of  others  are  briglilly  coloured — On  sexually-limited 
inheritance,  as  applied  to  various  structures  and  to  brightly- 
coloured  plumage — Nidification  in  relation  to  colour — Loss  of 
nuptial  plumage  during  the  winter      ....  444 

CHAPTER  XVI. 

Birds — concluded. 

The  immature  plumage  in  relation  to  the  character  of  the  plumage 
in  both  sexes  when  adult — Six  classes  of  cases — Sexual  diflfer- 
ences  between  the  males  of  closelj-allied  or  representative 
species — The  female  assuming  the  characters  of  the  male — 
Plumage  of  the  young  in  relation  to  the  summer  and  winter 
plumage  of  the  adults — On  the  increase  of  beauty  in  the  birds 
of  the  woi-ld  —  Protective  colouring  —  Conspicuously-coloured 
birds — Novelty  appreciated — Summary  of  the  four  chapters  on 
birds 463 

CHAPTER  XVII. 

Secondary  Sexual  Characters  of  Mammals. 

The  law  of  battle — Special  weapons,  contined  to  the  males — Cause 
of  absence  of  weapons  in  the  female — Weapons  common  to  both 
sexf  s,  yet  primarily  acquired  by  the  male — Other  uses  of  such 
weapons- -Their  high  importance — Greater  size  of  the  male — 
Means  of  defence — On  the  preference  sliewn  by  either  sex  in  the 
pairing  of  quadrupeds         ......  500 

CHAPTER  XVIII. 

Secondary  Sexual  Characters  of  Masimals — continued. 

Voice — Remarkable  sexual  peculiarities  in  seals — Odour — Develop- 
ment of  the  hair — Colour  of  the   hair   and   skin— Anomalous 


xvi  Contents. 


case  of  the  female  being  more  ornamented  than  the  male — 
Colour  and  ornaments  due  to  sexual  selection — Colour  acquired 
for  the  sake  of  protection — Colour,  though  common  to  both 
sexes,  often  due  to  sexual  selection — On  the  disappearance  of 
spots  and  stripes  in  adult  quadrupeds — On  the  colours  and  orna- 
ments of  the  Quadrumana — Summary        .  .         .  525 


PAET  in. 

SEXUAL  SELECTION  IN  KELATION  TO  MAN, 

AND   CONCLUSION. 


CHAPTER  XIX. 

Secondary  Sexual  Characters  op  Man. 

Differences  between  man  and  woman — Causes  of  such  differences, 
and  of  certain  characters  common  to  both  sexes — Law  of  battle 
— Differences  in  mental  powers,  and  voice — On  the  influence 
of  beauty  in  determining  the  marriages  of  mankind — Attention 
paid  by  savages  to  ornaments — Their  ideas  of  beauty  in  woman 
— The  tendency  to  exaggerate  each  natural  peculiarity         .         556 

CHAPTER  XX. 

Secondary  Sexual  Characters  of  Man — continued. 

On  the  effects  of  the  continued  selection  of  women  according  to  a 
different  standard  of  beauty  in  each  race — On  the  causes  which 
interfere  with  sexual  selection  in  civilised  and  savage  nations 
— Conditions  favourable  to  sexual  selection  during  primeval 
times — On  the  manner  of  action  of  sexual  selection  with  man- 
kind— On  the  women  in  savage  tribes  having  some  power  to 
choose  theix  husbands — Absence  of  hair  on  the  body,  and 
development  of  the  beard — Colour  of  the  skin — Summary      .         585 

CHAPTER  XXI. 

General  Summary  and  Conclusion. 

Main  conclusion  that  man  is  descended  from  some  lower  form — 
Manner  of  development — Genealogy  of  man — Intellectual  and 
moral  faculties — Sexual  selection — Concluding  remarks       .         603 


Index 620 


THE  DESCENT  OF  MAN; 

AND 

SELECTION  IN  KELATION  TO  SEX. 


INTRODUCTION. 

The  nature  of  the  following  work  will  be  best  imderstood  by  a 
brief  account  of  how  it  came  to  be  written.  During  many  years 
I  collected  notes  on  the  origin  or  descent  of  man,  without  any 
intention  of  publishing  on  the  subject,  but  rather  with  the 
determination  not  to  publish,  as  I  thought  that  I  should  thus 
only  add  to  the  prejudices  against  my  views.  It  seemed  to  me 
sufficient  to  indicate,  in  the  first  edition  of  my  'Origin  of 
Species,'  that  by  this  work  "light  would  be  thrown  on  the 
"  origin  of  man  and  his  history ;"  and  this  imphes  that  man  must 
be  included  with  other  organic  beings  in  any  general  conclusion 
respecting  his  manner  of  appearance  on  this  earth.  Now  the 
case  wears  a  wholly  different  aspect.  When  a  naturalist  like 
Carl  Vogt  ventures  to  say  in  his  address  as  President  of  the 
National  Institution  of  Geneva  (1869),  "  pcrsonne,  en  Europe 
"  au  moins,  n'ose  plus  soutenir  la  creation  independante  et  de 
"  toutes  pieces,  des  especes,"  it  is  manifest  that  at  least  a  largo 
number  of  naturalists  must  admit  that  species  are  the  modified 
descendants  of  other  species ;  and  this  especially  holds  good  with 
the  younger  and  rising  naturalists.  The  greater  number  accept 
the  agency  of  natural  selection ;  though  some  urge,  whether  with 
justice  the  future  must  decide,  that  I  have  greatly  overrated  its 
importance.  Of  the  older  and  honoured  chiefs  in  natural  science, 
many  unfortunately  are  still  opposed  to  evolution  in  every 
form. 

In  consequence  of  the  views  now  adopted  by  most  naturalists, 
and  which  will  ultimately,  as  in  every  other  case,  be  followed  by 


Introduction. 


others  who  are  not  scientific,  I  have  been  led  to  put  together 
my  notes,  so  as  to  see  how  far  the  general  conclusions  arrived  at 
in  my  former  works  were  applicable  to  man.  This  seemed  all 
the  more  desirable,  as  I  had  never  deliberately  applied  these 
views  to  a  species  taken  singly.  When  we  confine  our  attention 
to  any  one  form,  we  are  •  dej^rived  of  the  weighty  arguments 
derived  from  the  nature  of  the  affinities  which  connect  together 
whole  groups  of  organisms— their  geographical  distribution  in 
past  and  present  times,  and  their  geological  succession.  The 
homological  structure,  embryological  development,  and  rudi- 
mentary organs  of  a  species  remain  to  be  considered,  whether  it 
be  man  or  any  other  animal,  to  which  our  attention  may  be 
directed  ;  but  these  great  classes  of  facts  afford,  as  it  appears  to 
me,  ample  and  conclusive  evidence  in  favour  of  the  principle  of 
gradual  evolution.  The  strong  support  derived  from  the  other 
arguments  should,  however,  always  be  kept  before  the  mind. 
.  The  sole  object  of  this  work  is  to  consider,  firstly,  whether 
man,  like  every  other  species,  is  descended  from  some  pre- 
existing form  ;  secondly,  the  manner  of  his  development ;  and 
thirdly,  the  value  of  the  differences  between  the  so-called  races 
of  man.  As  I  shall  confine  myself  to  these  jDoints,  it  will  not  be 
necessary  to  describe  in  detail  the  differences  between  the  several 
races— an  enormous  subject  which  has  been  fully  discussed  in 
many  valuable  works.  The  high  antiquity  of  man  has  recently 
been  demonstrated  by  the  labours  of  a  host  of  eminent  men, 
beginning  with  M.  Boucher  de  Perthes ;  and  this  is  the  indis- 
pensable basis  for  understanding  his  origin.  I  shall,  therefore, 
take  this  conclusion  for  granted,  and  may  refer  my  readers  to 
the  admirable  treatises  of  Sir  Charles  Lyell,  Sir  John  Lubbock, 
and  others.  Nor  shall  I  have  occasion  to  do  more  than  to  allude 
to  the  amount  of  difference  between  man  and  the  anthropomor- 
phous apes  ;.for  Prof.  Huxley,  in  the  opinion  of  most  competent 
judges,  has  conclusively  shewn  that,  in  every  visible  character 
man  differs  less  from  the  higher  apes,  than  these  do  from  the 
lower  members  of  the  same  order  of  Primates. 

This  work  contains  hardly  any  original  facts  in  regard  to  man ; 
but  as  the  conclusions  at  which  I  arrived,  after  drawing  up  a 
rough  draft,  appeared  to  me  interesting,  I  thought  that  they 
might  interest  others.  It  has  often  and  confidently  been  asserted, 
that  man's  origin  can  never  be  known  :  but  ignorance  more 
frequently  begets  confidence  than  does  knowledge :  it  is  those 
who  know  little,  and  not  those  who  know  much,  who  so 
positively  assert  that  this  or  that  problem  will  never  be  solved 
by  science.  The  conclusion  that  man  is  the  co-descendant  with 
other  species  of  some  ancient,  lower,  and  extinct  form,  is  not  in 


Introductioji. 


any  degree  new.  Lamarck  long  ago  came  to  this  conclusion, 
which  has  lately  been  maintained  by  several  eminent  naturalists 
and  philosophers ;  for  instance,  by  Wallace,  Hnxley,  Lyell,  Vogt, 
Lubbock,  Biichner,  Rolle,  &c.,^  and  especially  by  Hackel.  Tlj^s 
last  naturalist,  besides  his  great  work,  *  Gencrclle  Morphologic ' 
(1866),  has  recently  (1868,  with  a  second  edit,  in  1870),  pub- 
lished his  *  Naturliche  Schopfungsgeschichte,'  in  which  he  fully 
discusses  the  genealogy  of  man.  If  this  work  had  appeared 
before  my  essay  had  been  written,  I  should  probably  never  have 
completed  it.  Almost  all  the  conclusions  at  which  I  have 
arrived  I  find  confirmed  by  this  naturalist,  whose  knowledge  on 
many  points  is  much  fuller  than  mine.  Wherever  I  have  added 
any  fact  or  view  from  Prof.  Hiickel's  writings,  I  give  his  autho- 
rity in  the  text ;  other  statements  I  leave  as  they  originally  stood 
in  my  manuscript,  occasionally  giving  in  the  foot-notes  references 
to  his  works,  as  a  confirmation  of  the  more  doubtful  or  interesting 
points. 

During  many  years  it  has  seemed  to  me  highly  probable  that 
sexual  selection  has  played  an  important  part  in  differentiating 
the  races  of  man  ;  but  in  my  '  Origin  of  Species  '  (first  edition,  p. 
199)  I  contented  myself  by  merely  alluding  to  this  belief.  "When 
I  came  to  apply  this  view  to  man,  I  found  it  indispensable  to 
treat  the  whole  subject  in  full  detail.^  Consequently  the  second 
part  of  the  present  work,  treating  of  sexual  selection,  has  ex- 
tended to  an  inordinate  length,  compared  with  the  first  part; 
but  this  could  not  be  avoided. 

I  had  intended  adding  to  the  present  volumes  an  essay  on  the 
expression  of  the  various  emotions  byman  and  the  lower  animals. 
My  attention  was  called  to  this  subject  many  years  ago  by 
Sir  Charles  Bell's  admirable  work.     This  illustrious  anatomist 

'  As  the  works  of  the  first-named  JSat.,' Modena,  ]8(37,  p.   81)  a  very 

authors  are  so  well  known,  I  need  curious  paper  on  rudimentary  cha- 

not  give  the  titles;  but  as  those  of  racters,  as  bearing  on  the  ori'^in  of 

the   latter  are   less  well   known  in  man.      Another    work    has    (1869) 

England,  I  will  give  them  : — '  Sechs  been    published    by    Dr.    Francesco 

Vorlesungen   iiber   die  Darwin'sche  Barrago,  bearing  in  Italian  the  title 

Theorie:'  zweite  Auflage,  1868,  von  of  "Man,  made  in  the  image  of  God, 

Dr.    L.    Buchner;     translated    into  "  was  also  made  in  the  image  of  the 

French  under  the  title  'Conferences  "ape." 

sur  la  Theorie  Darwinienne,'  1869.  '•^    Prof.    Hiickel    was    the     only 

*Der  Mensch,   im   Lichte    der   Dar-  author  who,  at  the  time  when  this 

win'sche    Lehre,'   1865,   von  Dr.   F.  work   first  appeared,  had   di.-,cussed 

Rolle.     I  will   not  attempt  to  give  the  subject  of  sexual  selection,  and 

references  to   all    the    authors  who  had  seen   its  full   importance,  since 

have    taken   the    same    side   of  the  the  publication  of  the  'Origin ';  and 

question.     Thus   G.   Canestrini    has  this  he  did  in  a  very  able  manner  io 

published  ('  Annuario  della   Soc.  d.  his  various  works. 


Introduction. 


maintains  that  man  is  endowed  with  certain  muscles  solely  for 
the  sake  of  expressing  his  emotions.  As  this  view  is  obviously 
opposed  to  the  belief  that  man  is  descended  from  some  other  and 
l4)wer  form,  it  was  necessary  for  me  to  consider  it.  I  likewise 
wished  to  ascertain  how  far  the  emotions  are  expressed  in  the 
same  manner  by  the  different  races  of  man.  But  owing  to.  the 
length  of  the  present  work,  I  have  thought  it  better  to  reser\^e 
my  essay  for  separate  publication. 


(    5    ) 


Part  I. 
THE  DESCENT  OR  ORIGIN  OF  MAN. 


CHAPTER  I. 

The  Evidence  of  the  Descent  of  Man  from  some 
Lower  Form. 

Nature  of  the  evidence  bearing  on  the  origin  of  man  —  Homologous 
structures  in  man  and  the  lower  animals — Miscellaneous  points  of 
correspondence — Development — Rudimentary  structures,  muscles,  sense- 
organs,  hair,  bones,  reproductive  organs,  &c. — The  bearing  of  these  three 
great  classes  of  tacts  on  the  origin  of  man. 

He  who  wishes  to  decide  whether  man  is  the  modified  descendant 
of  some. pre-existing  form,  would  probably  first  enquire  whether 
man  varies,  however  slightly,  in  bodily  structure  and  in  mental 
faculties;  and  if  so,  whether  the  variations  are  transmitted  to 
his  offspring  in  accordance  with  the  laws  which  prevail  with  tlie 
lower  animals.  Again,  are  the  variations  the  result,  as  far  as 
our  ignorance  permits  us  to  judge,  of  the  same  general  causes, 
and  are  they  governed  by  the  same  general  laws,  as  in  the  case 
of  other  organisms ;  for  instance,  by  correlation,  the  inherited 
effects  of  use  and  disuse,  &c.  ?  Is  man  sulycct  to  similar  mal- 
couformations,  the  result  of  arrested  development,  of  reduplication 
of  parts,  &c.,  and  does  he  display  in  any  of  his  anomalies  rever- 
sion to  some  former  and  ancient  typo  of  structure  ?  It  might 
also  naturally  be  enquired  whether  man,  like  so  many  other 
animals,  has  given  rise  to  varieties  and  sub-races,  differing  but 
slightly  from  each  other,  or  to  races  differing  so  much  that  they 
must  be  classed  as  doubtful  species?  How  are  such  races 
distributed  over  the  world;  and  how,  when  crossed,  do  they 
react  on  each  other  in  the  first  and  succeeding  generations  ? 
And  so  with  many  other  points. 

The  enquirer  would  next  come  to  the  important  point, 
whether  man  tends  to  increase  at  so  rapid  a  rate,  as  to  lead  to 
occasional  severe  struggles  for  existence;   and  consequently  to 


6  The  Descent  of  Man.  Part  I. 

beneficial  variations,  whether  in  body  or  mind,  being  preserved, 
and  injurious  ones  eliminated.  Do  the  races  or  species  of  men, 
whichever  term  may  be  applied,  encroach  on  and  replace  one 
another,  so  that  some  finally  become  extinct  ?  We  shall  see  that 
all  these  questions,  as  indeed  is  obvious  in  respect  to  most  of 
them,  must  be  answered  in  the  affirmative,  in  the  same  manner 
as  with  the  lower  animals.  But  the  several  considerations  just 
referred  to  may  be  conveniently  deferred  for  a  time :  and  we 
will  first  see  how  far  the  bodily  structure  of  man  shows  traces, 
more  or  less  plain,  of  his  descent  from  some  lower  form.  In 
succeeding  chapters  the  mental  powers  of  man,  in  comparison 
w^"th  those  of  the  lower  animals,  will  be  considered. 

The  Bol'dy  Structure  of  Man. — It  is  notorious  that  man  is 
constructed  on  the  same  general  type  or  model  as  other  mam- 
mals. All  the  bones  in  his  skeleton  can  be  compared  with 
corresponding  bones  in  a  monkey,  bat,  or  seal.  So  it  is  with  his 
muscles,  nerves,  blood-vessels  and  internal  viscera.  The  brain, 
the  most  important  of  all  the  organs,  follows  the  same  law,  as 
shewn  by  Huxley  and  other  anatomists.  Bischoff,^  who  is  a 
hostile  witness,  admits  that  every  chief  fissure  and  fold  in  the 
brain  of  man  has  its  analogy  in  that  of  the  orang  ;  but  he  adds 
that  at  no  period  of  development  do  their  brains  perfectly  agree  ; 
nor  could  perfect  agreement  be  expected,  for  otherwise  their 
mental  powers  would  have  been  the  same.  Vulpian  ^  remarks  : 
"  Les  differences  reelles  qui  existent  entre  I'encephale  de 
"  t'homme  et  celui  des  singes  superieurs,  sont  bien  minimes.  II 
"  ne  faut  pas  se  faire  d'illusions  a  cet  egard.  L'homme  est  bien 
"  plus  pres  des  singes  anthropomorphes  par  les  caracteres 
"  anatomiques  de  son  cerveau  que  ceux-ci  ne  le  sont  nou- 
"  seulement  des  autres  mammiferes,  mais  meme  de  certains 
"  quadrumanes,  des  guenons  et  des  macaques."  But  it  would 
be  superfluous  here  to  give  further  details  on  the  correspondence 
between  man  and  the  higher  manmials  in  the  structure  of  the 
brain  and  all  other  i^arts  of  the  body. 

It  may,  however,  be  worth  while  to  specify  a  few  points,  not 
directly  or  obviously  connected  with  structure,  by  which  this 
correspondence  or  relationship  is  well  shewn. 

Man  is  liable  to  receive  from  the  lower  animals,  and  to  com- 

1  '  Grosshirnwiadungea  des  Men-  in  the  Preface  to  this  edition, 
scheu,'  1868,  s.  96.     The  conchisions  ^  t  Le^_  g^^.  i^  Phj-s.'  1866,  p.  890, 

of  this  author,  as  well  as  those  of  as  quoted  by  M.  Dally,  '  L'Ordre  des 

Gratiolet  and  Aeby,  concerning  the  Primates  et  leTranslbrmisme,' 1868, 

brain,   will    be    discussed   by  Prof.  p.  29. 
Huxley  in  the  Appendix  alluded  to 


Chap.  I.  Hoinological  Structures,  7 

mnnicatc  to  them,  certain  diseases,  as  hydrophobia,  varioUi,  tho 
glanders,  syphilis,  cholera,  herpes,  etc. ;  ^  and  this  fact  proves  tho 
close  similarity*  of  their  tissues  and  blood,  both  in  minntc 
structure  and  composition,  far  more  plainly  than  does  their 
comparison  under  the  best  microscope,  or  by  the  aid  of  the  best 
chemical  analysis.  Monkeys  are  liable  to  many  of  tho  same  non- 
contagious diseases  as  we  are;  thus  Eengger,^  who  carefully 
observed  for  a  long  time  the  Cthm  Azarai  in  its  native  land, 
found  it  liable  to  catarrh,  with  the  usual  symptoms,  and  which, 
when  often  recurrent,  led  to  consumption.  These  monkeys 
suffered  also  from  apoph^xy,  infiannnation  of  the  bowels,  and 
cataract  in  the  eye.  The  younger  ones  when  shedding  their 
milk-teeth  often  died  from  fever.  Medicines  produced  the  same 
effect  on  them  as  on  us.  Many  kinds  of  monkeys  have  a  strong 
taste  for  tea,  coffee,  and  spirituous  liquors :  they  will  also',  as  I 
have  myself  seen,  smoke  tobacco  with  pleasure."  Brehm  asserts, 
that  the  natives  of  north-eastern  Africa  catch  the  wild  baboons 
by  exposing  vessels  with  strong  beer,  by  which  they  are  made 
drunk.  He  has  seen  some  of  these  animals,  which  he  kept  in 
confinement,  in  this  state ;  and  he  gives  a  laughable  account  of 
their  behaviour  and  strange  grimaces.  On  the  following 
morning  they  were  very  cross  and  dismal ;  they  held  their  aching 
heads  with  both  hands,  and  wore  a  most  pitiable  expression : 
when  beer  or  wine  was  offered  them,  they  turned  away  with 
disgust,  but  relished  the  juice  of  lemons.^  An  American  monkey, 
an  Ateles,  after  getting  drunk  on  brandy,  would  never  touch  it 
again,  and  thus  was  wiser  than  many  men.  These  trifling  facts* 
prove  how  similar  the  nerves  of  taste  must  be  in  monkeys  and 
man,  and  how  similarly  their  whole  nervous  system  is  affected. 
Man  is  infested  with  internal  parasites,  sometimes  causing 

'    Dr.    W.    Lauder    Lindsay    has  tinct   fluids   by   the   same   chemical 

treated  this  subject  at  some  length  reagent. 

in  the  '  Journal  of  Mental  Science,'  *    '  Naturgescliichte    der    Siiuge- 

July  1871;  and  in  the  'Edinburgh  thiere  von  Paraguay,'  1830,  s.  50. 
Veterinary  lieview,'  July  1858.  ®  The  same  tastes  are  common  to 

*     A     Rev.ewer     has      criticised  some    animals    much    lower  in    the 

('  British    Quarterly    Review,'    Oct.  scale.     Mr.    A.    Nicols    informs   me 

1st,  1871,  p.  472)  what  I  have  here  that  he  kept  in  Queensland,  in  Aus- 

said  with  much    severity  and    con-  tralia,    three    individuals     of     the 

tempt;  but  as  I  do  not  use  the  term  Fhaseolarctus    cinereus  ;    and    that, 

identity,  I    cannot    see    that    I    am  without  having  been  taught  in  any 

greatly  in  error.     There  apjiears  to  way,  they  acquired   a  strong  taste 

me  a  strong   analogy  between   the  for  rum,  and  for  smoking  tobacco, 
"same    infection    or   contagion    pro-  ^  Brehm, 'Thicrleben,' B.  i.  1864, 

ducing    the    same    result,    or    one  s.  75,  86.     On    the  Ateles,  s.   105. 

closely  similar,  in  two  distinct  ani-  For  other  analogous  statements,  see 

mals,  and   the   testing  of  two   dis-  s.  25,  107. 


8  The  Descent  of  Man.  Part  I. 

fatcal  effects ;  and  is  plagued  by  external  parasites,  all  of  which 
belong  to  the  same  genera  or  families  as  those  infesting  other 
mammals,  and  in  the  case  of  scabies  to  the  same  species.^  Man 
is  subject,  like  other  mammals,  birds,  and  even  insects,^  to  that 
mysterious  law,  which  causes  certain  normal  processes,  such  as 
gestation,  as  well  as  the  maturation  and  duration  of  various 
diseases,  to  follow  lunar  periods.  His  wounds  are  repaired  by 
the  same  process  of  healing;  and  the  stumps  left  after  the 
amputation  of  his  limbs,  especially  during  an  early  embryonic 
period,  occasionally  possess  some  power  of  regeneration,  as  in 
the  lowest  animals.'" 

The  whole  process  of  that  most  important  function,  the 
reproduction  of  the  species,  is  strikingly  the  same  in  all  mam- 
mals, from  the  first  act  of  courtship  by  the  male,"  to  the  birth 
and  nurturing  of  the  young.  Monkeys  are  born  in  almost  as 
helpless  a  condition  as  our  own  infants ;  and  in  certain  genera 
the  young  differ  fully  as  much  in  appearance  from  the  adults,  as 
do  our  children  from  their  full-grown  parents.'^  It  has  been 
urged  by  some  writers,  as  an  important  distinction,  that  with 
man  the  young  arrive  at  maturity  at  a  much  later  age  than  with 
any  other  animal :  but  if  we  look  to  the  races  of  mankind  which 
inhabit  tropical  countries  the  difference  is  not  great,  for  the 
orang  is  believed  not  to  be  adult  till  the  age  of  from  ten  to  fifteen 
years.'^  Man  differs  from  woman  in  size,  bodily  strength, 
hairiness,  &c.,  as  well  as  in  mind,  in  the  same  manner  as  do  the 

.  ^  Dr.  W.  Lauder  Lindsay,  '  Edrn-  "  sagax,  hoc  mihi  certissime  pro- 
burgh  Vet.  Review,'  July  1858,  "  bavit,  et  curatores  ejusdem  loci  et 
p.  13.  "  alii    e    ministris    confirmaverunt. 

^  With  respect  to  insects  see  Dr.  "  Sir  Andrew  Smith  et  Brehm  no- 

Laycock,  "  On  a  General  Law  of  Vital  "  tabant  idem  in  Cynocephalo.     II- 

Periodicity,"   '  British   Association,'  "  lustrissimus  Cuvier  etiam  narrat 

1842.     Dr.  Macculloch,  '  Silliman's  *'  multa  de   hac  re,  qua  ut  opinor, 

North  American  Journal  of  Science,'  "  nihil  turpius  potest  indicari  inter 

vol.   xvii.  p.   305,  has    seen   a  dog  "  omnia  hominibus  et  Quadrumanis 

suffering  from  tertian  ague.     Here-  "  communia.      Narrat    enim   Cyno- 

after  I  shall  return  to  this  subject.  "  cephalum  quendam  in  furorem  in- 

^o  I   haA-e   given  the  evidence  on  "  cidere     aspectu    feminarum     ali- 

this  head  in  my  '  Variation  of  Ani-  "  quarum,  sed   nequaquam  accendi 

mals  and  Plants  under  Domestica-  "  tanto   furore   ab    omnibus.     Sem- 

tion,'  Tol.  ii.  p.  15,  and  more  could  "  per  eligebat  juniores,  et   dignos- 

be  added.  "  cebat  in  turba,  et  advocabat  voce 

•^    "  Mares  e    diversis    generibus  "  gestuque." 

"  Quadrumanorum   sine    dubio    di-  '^  This  remark  is  made  with  re- 

"  gnoscunt  feminas  humanas  a  ma-  spect  to  Cynocephalus  and  the  an- 

"  ribus.       Primum,  credo,  odoratu,  thropomorphous    apes   by   Geoffrey 

"  postea  aspectu.     Mr.  Youatt,  qui  Saint-Hilaire  and  F.  Cuvier,  '  Hist, 

*'  diu  in    Hortis   Zoologicis   (Besti-  Nat.  des  Mammiferes,'  tom.  1.  1824. 

"  ariis)    medicus    animalium    erat,  ^^  Huxley,  '  Man's   Place  in   Na- 

"  vir  in  rebus  observandis  cautus  et  ture,'  1863,  p.  34. 


(^.iiAP.  1.  Homological  Structures.  9 

two  sexes  of  many  mammals.  So  that  the  correspondence  in 
general  structure,  in  the  minute  structure  of  the  tissues,  in 
chemical  composition  and  in  constitution,  between  man  and  tlie 
liigher  animals,  especially  the  anthropomorphous  apes,  is  ex- 
tremely close. 

Emhnjonlc  Dcv^Ioj-mcnt. — Man  is  developed  from  an  ovule, 
about  the  125th  of  an  inch  in  diameter,  which  diifcrs  in  no 
respect  from  the  ovules  of  other  animals.  The  embryo  itself  at 
a  very  early  period  can  hardly  be  distinguished  from  that  of 
other  members  of  the  vertebrate  kingdom.  At  this  period  the 
arteries  run  in  arch-like  branches,  as  if  to  carry  the  blood  to 
branchite  which  are  not  present  in  the  higher  vertebrata,  though 
the  slits  on  the  sides  of  the  neck  still  remain  (/,  g,  fig.  1), 
marking  their  former  position.  At  a  somewhat  later  period, 
when  the  extremities  are  developed,  "  the  feet  of  lizards  and 
"  mammals,"  as  the  illustrious  Von  Baer  remarks,  "  the  wings 
"  and  feet  of  birds,  no  less  than  the  hands  and  feet  of  man,  all 
"  arise  from  the  same  fundamental  form."  It  is,  says  Prof. 
Huxley,^'*  "  quite  in  the  later  stages  of  development  that  the 
young  human  being  presents  marked  differences  from  the  young 
"  ape,  while  the  latter  departs  as  much  from  the  dog  in  its 
"  developments,  as  the  man  does.  Startling  as  this  last  assertion 
"  may  appear  to  be,  it  is  demonstrably  true." 

As  some  of  my  readers  may  never  have  seen  a  drawing  of  an 
embryo,  I  have  given  one  of  man  and  another  of  a  dog,  at  about 
the  same  early  stage  of  development,  carefully  copied  from  two 
works  of  undoubted  accuracy.'^ 

After  the  foregoing  statements  made  by  such  high  autho- 
rities, it  would  be  superfluous  on  my  part  to  give  a  number  of 
borrowed  details,  shewing  that  the  embryo  of  man  closely 
resembles  that  of  other  mammals.  It  may,  however,  be  added, 
that  the  human  embryo  likewise  resembles  certain  low  forms 
when  adult  in  various  points  of  structure.  For  instance,  the 
heart  at  first  exists  as  a  simple  pulsating  vessel;  the  excreta 
are  voided  through  a  cloacal  passage  ;  and  the  os  coccyx  projects 

•*  '  Man's  Place  in  Nature,'  1863,  magnified,  the  embryo  being  twenty- 

p,  67.  five  days  old.     The  internal  viscera 

'^    The    human     embryo    (upper  havebeenomitted, and  theuterineap- 

fig.)  is  from  Ecker,  '  Icones  Phys.,'  pendages  in  both  drawings  removed. 

1851-1859,    tab.   xxx.  fig.   2.     This  I  was  directed  to  these  figures    by 

embryo  was  ten   lines  in  length,  so  Prof.    Huxley,    from     whose     woj-k, 

that  the  drawing  is  much  magnified.  '  Man's  Place  in  Nature,'  the  idea  of 

The    embryo    of    the    dog    is    from  givmg  them  was  taken.    Hiickcl  has 

Jiischoff,      '  Entwicklungsgeschichte  also  given  analogous  drawings  in  his 

des  Humle-Eies,'  1845,  tab.  xi.   fig.  '  Schoufuugsgescliichte.' 
42  B       This  drawing  is  five  time? 
2 


10 


The  Descent  of  Man. 


Part  I. 


Fig.  1. 


Upper  figure  human  embryo,  from  'Ecker. 
from  BischofF. 


Lower  figure  that  of  a  dog. 


a.  Fore-brain,  cerebral  hemispheres,  &c. 
6.  Mid-brain,  corpora  quadrigeniina. 

c.  Hind-brain,  cerebellum,  medulla  ob- 
longata. 

d.  Eye. 

e.  Ear. 

f.  First  visceral  arch. 


<7.  Second  visceral  arch. 
H.  Vertebral  columns  and  muscles 
process  of  development. 
i    Anterior 
K.  Posterior 
L.  Tail  or  os  coccyx. 


extremities. 


Chap.  I.  Riidivioits.  1 1 

like  a  true  tail,  "extending  considerably  beyond  the  rudi- 
"  mentary  legs.""^  In  the  embryos  of  all  air-breathing  vertebrates, 
certain  glands,  called  the  corpora  Wolffiana,  correspond  with, 
and  act  like  the  kidneys  of  mature  fishes.''^  Even  at  a  later 
embryonic  period,  some  striking  resemblances  between  man  and 
the  lower  animals  may  be  observed.  Bischoflf  says  that  the 
convolutions  of  the  brain  in  a  human  foetus  at  the  end  of  the 
seventh  month  reach  about  the  same  stage  of  develoj^ment  as  in 
a  baboon  when  adult.^**  The  great  toe,  as  Prof.  Owen  rcmarks,^^ 
"  which  forms  the  fulcrum  when  standing  or  w\alking,  is 
"  perhaps  the  *  most  characteristic  peculiarity  in  the  human 
structure ;"  but  in  an  embryo,  about  an  inch  in  length.  Prof. 
Wyman  '^"^  found  "  that  the  great  toe  was  shorter  than  the  others  ; 
"  and,  instead  of  being  parallel  to  them,  projected  at  an  angle 
"  from  the  side  of  the  foot,  thus  corresponding  with  the  per- 
*'  manent  condition  of  this  part  in  the  quadrumana."  I  will 
conclude  with  a  quotation  from  Huxley ,^^  who  after  asking, 
does  man  originate  in  a  different  way  from  a  dog,  bird,  frog  or 
fish?  says,  "the  reply  is  not  doubtful  for  a  moment;  without 
"  question,  the  mode  of  origin,  and  the  early  stages  of  the 
"  development  of  man,  are  identical  with  those  of  the  animals 
"  immediately  below  him  in  the  scale :  without  a  doubt  in 
"  these  respects,  he  is  far  nearer  to  apes  than  the  apes  are  to 
"  the  dog." 

lludimenU.  —  This  subject,  though  not  intrinsically  more 
important  than  the  two  last,  will  for  several  reasons  be  treated 
here  more  fully.-^  Kot  one  of  the  higher  animals  can  be  named 
which  does  not  bear  some  part  in  a  rudimentary  condition  ;  and 
man  forms  no  exception  to  the  rule.  Eudimentary  organs  must 
be  distinguished  from  those  that  are  nascent ;  though  in  some 
cases  the  distinction  is  not  easy.  The  former  are  either  abso- 
lutely useless,  such  as  the  mammae  of  male  quadrupeds,  or  the 
incisor  teeth  of  ruminants  which  never  cut  through  the  gums ; 
or  they  are  of  such  slight  service  to  their  present  possessors, 
that  w^e  can  hardly  suppose  that  they  were  developed  under  the 

'*    Prof.    Wyman    in    *  Proc.     of  ^^  I  had  written  a  rough  copy  of 

American  Acad,  of  Sciences,'  rol.  iv.  this  chapter  before  reading  a  valu- 

1860,  p.  .17,  able   paper,  "  Caratteri  rudimentali 

*'    Owen,    '  Anatomy    of    Verte-  in    ordine    all'    origine    del    iiomo " 

brates,'  vol.  i.  p.  533.  ('  Annuario  della  Soc.  d.  Nat.,'  Mo- 

'*  'Die    Grosshirnwinduugen  des  dena,  1867,  p.  81),  by  G.  Canestrini, 

Menschen,'  1868,  s.  95.  to   which  paper  I  am   considerably 

'^  'Anatomy  of  Vertebrates,' vol.  indebted.     Hiickel  has  given  admir- 

ji.  p.  553.  able  discussions  on  this  whole  sub- 

'^°  '  Proc.  Soc.  Nat.  Hist.'  Boston,  ject,  under  the  title  of  Dysteleology, 

1863,  vol.  ix.  p.  185.  in  his  '  Generelle  Morphologic'  and 

'*   '  Man's  Place  in  Nature,'  p.  65.  '  Schopfungsgeschichta.' 


1 2  The  Descent  of  Man.  Part  I. 


conditions  wliicli  now  exist.  Organs  in  this  latter  state  are  not 
strictly  rudimentary,  but  they  are  tending  in  this  direction. 
Nascent  organs,  on  the  other  hand,  though  not  fully  developed, 
are  of  high  service  to  their  possessors,  and  are  capable  of  further 
development.  Rudimentary  organs  are  eminently  variable ;  and 
this  is  partly  intelligible,  as  they  are  useless,  or  nearly  useless, 
and  consequently  are  no  longer  subjected  to  natural  selection. 
They  often  become  wholly  suppressed.  When  this  occurs,  they 
are  nevertheless  liable  to  occasional  reappearance  through 
reversion— a  circumstance  well  worthy  of  attention. 

The  chief  agents  in  causing  organs  to  become  rudimentary 
seem  to  have  been  disuse  at  that  period  of  life  when  the  organ 
is  chiefly  used  (and  this  is  generally  during  maturity),  and  also 
inheritance  at  a  corresponding  period  of  life.  The  term 
"  disuse "  does  not  relate  merely  to  the  lessened  action  of 
muscles,  but  includes  a  diminished  flow  of  blood  to  a  part  or 
organ,  from  being  subjected  to  fewer  alternations  of  pressure,  or 
from  becoming  in  any  way  less  habitually  active.  Eudiments, 
however,  maj  occur  in  one  sex  of  those  parts  which  are  normally 
present  in  the  other  sex;  and  such  rudiments,  as  we  shall 
hereafter  see,  have  often  originated  in  a  way  distinct  from  those 
here  referred  to.  In  some  cases,  organs  have  been  reduced  by 
means  of  natural  selection,  from  having  become  injurious  to  the 
species  under  changed  habits  of  life.  The  process  of  reduction 
is  probably  often  aided  through  the  two  principles  of  compensa- 
tion and  economy  of  growth  ;  but  the  later  stages  of  reduction, 
after  disuse  has  done  all  that  can  fairly  be  attributed  to  it,  and 
when  the  saving  to  be  effected  by  the  economy  of  growth  would  be 
very  small/^  are  difl&cult  to  understand.  The  final  and  complete 
suppression  of  a  part,  already  useless  and  much  reduced  in  size, 
in  which  case  neither  compensation  nor  economy  can  come  into 
play,  is  perhaps  intelligible  by  the  aid  of  the  hypothesis  of 
pangenesis.  But  as  the  wliole  subject  of  rudimentary  organs 
has  been  discussed  and  illustrated  in  my  former  works,^^  I  need 
here  say  no  more  on  this  head. 

Eudiments  of  various  muscles  have  been  observed  in  many 
parts  of  the  human  body  ;^^  and  not  a  few  muscles,  which  are 

^^  Some  good  criticisms  on  this  Zoolog.  1852,  torn,  xviii.  p.  13)  de- 
subject  have  been  given  by  Messrs.  scribes  and  figures  rudiments  of 
Murie  and  Mivart,  in  'Transact.  what  he  calls  the  "  muscle  pe'dieux 
Zoolog.  Soc'  1869,  vol.  vii.  p.  92.  de  la  main,"  which  he  says  is  some- 

2*    '  Variation    of    Animals    and  times  "  ipfiniment  petit."     Another 

Plrnts  under  Domestication,'  vol.  ii.  muscle,  called  "  le  tibial  posterieur," 

pp.  317  and  397.     See  also  '  Oj-igin  is    generally    quite    absent    in    the 

of  Species,'  5th  edit.  p.  535,  hand,  but  appears  from  time  to  time 

2*  For  instance  M.  Richard  ('  An-  in  a  more  or  less  rudimentary  con- 

uales  des  Sciem'.es  Nat.'  3i-d  sex'ies,  dltion. 


Chap.  I.  Riidii}ie]its.  13 

regularly  present  in  some  of  the  lower  animals  can  occasionally 
be  detected  in  man  in  a  greatly  reduced  condition.  Every  one 
must  have  noticed  the  power  which  many  animals,  especially 
horses,  possess  of  moving  or  twitchiDg  their  skin;  and  this  is 
effected  by  the  panniculns  carnosus.  Remnants  of  this  muscle 
in  an  efficient  state  are  found  in  various  parts  of  our  bodies ;  for 
instance,  the  muscle  on  the  forehead,  by  which  the  eyebrows  are 
raised.  The  /^A/Zysywa  myoides,  which  is  well  developed  on  the 
neck,  belongs  to  this  system.  Prof.  Turner,  of  Edinburgh,  has 
occasionally  detected,  as  he  informs  me,  muscular  lasciculi  in 
five  different  situations,  namely  in  the  axillae,  near  the  scapulae, 
&c.,  all  of  which  must  be  referred  to  the  system  of  the  jianni- 
culus.  He  has  also  shewn ^'^  that  the  muscnlus  stemalis  or  stemalis 
hrutorum,  which  is  not  an  extension  of  the  rectus  abdominal  is, 
but  is  closely  allied  to  the  pannicidas,  occurred  in  the  proportion 
of  about  three  per  cent,  in  upwards  of  600  bodies  :  he  adds,  that 
this  muscle  affords  "an  excellent  illustration  of  the  statement 
"  that  occasional  and  rudimentary  structures  are  especially 
"  liable  to  variation  in  arrangement." 

Some  few  persons  have  the  power  of  contracting  the  super- 
ficial muscles  on  their  scalps ;  and  these  muscles  are  in  a 
variable  and  partially  rudimentary  condition.  M.  A.  de  Candolle 
has  communicated  to  me  a  curious  instance  of  the  long-continued 
persistence  or  inheritance  of  this  power,  as  well  as  of  its  unusual 
development.  He  knows  a  family,  in  which  one  member,  the 
present  head  of  the  family,  could,  when  a  youth,  pitch  several 
heavy  books  from  his  head  by  the  movement  of  the  scalp  alone ; 
and  he  won  wagers  by  performing  this  feat.  His  father,  uncle, 
grandfather,  and  his  three  children  possess  the  same  power  to 
the  same  unusual  degree.  This  family  became  divided  eight 
generations  ago  into  two  branches;  so  that  the  head  of  the 
above-mentioned  branch  is  cousin  in  the  seventh  degree  to  the 
head  of  the  other  branch.  This  distant  cousin  resides  in 
another  part  of  France  ;  and  on  being  asked  whether  he  possessed 
the  same  faculty,  immediately  exhibited  his  power.  This  case  offers 
a  good  illustration  how  persistent  may  be  the  transmission  of  an 
absolutely  useless  faculty,  probably  derived  from  our  remote  semi- 
human  progenitors;  since  many  monkeys  have,  and  frequently 
use  the  power,  of  largely  moving  their  scalps  up  and  down." 

The  extrinsic  muscles  which  serve  to  move  the  external  ear, 
and  the  intrinsic  muscles  which  move  the  different  parts,  are  in  a 
rudimentary  condition  in  man,  and  they  all  belong  to  the  system 

'«  Prof.  W.  Turner,  '  Proc.  Royal  Emotions  in  Man  and  Animals,' 
Soc.  Edinburgh,'  1866-67,  p.  65.  1872,  p.  144. 

-^    See    my    '  Expression    of   the 


14  The  Descent  of  Man.  Part  1 


of  the  panniculus ;  they  are  also  variable  in  development,  or  at 
least  in  function.  I  have  seen  one  man  who  could  draw  the 
whole  ear  forwards ;  other  men  can  draw  it  upwards ;  another 
who  could  draw  it  backwards  f^  and  from  what  one  of  these 
persons  told  me,  it  is  probable  that  most  of  us,  by  often  touching 
our  ears,  and  thus  directing  our  attention  towards  them,  could 
recover  some  power  of  movement  by  repeated  trials.  The  power 
of  erecting  and  directing  the  shell  of  the  ears  to  the  various 
points  of  the  compass,  is  no  doubt  of  the  highest  service  to 
many  animals,  as  they  thus  perceive  the  direction  of  danger; 
but  I  have  never  heard,  on  sufficient  evidence,  of  a  man  who 
possessed  this  power,  the  one  which  might  be  of  use  to  him. 
The  whole  external  shell  may  be  considered  a  rudiment,  together 
with  the  various  folds  and  prominences  (helix  and  anti-helix, 
tragus  and  anti-tragus,  &c.)  which  in  the  lower  animals 
strengthen  and  support  the  ear  when  erect,  without  adding 
much  to  its  weight.  Some  authors,  however,  suppose  that  the 
cartilage  of  the  shell  serves  to  transmit  vibrations  to  the 
acoustic  nerve;  but  Mr.  Toynbee,^^  after  collecting  all  the 
known  evidence  on  this  head,  concludes  that  the  external  shell 
is  of  no  distinct  use.  The  ears  of  the  chimpanzee  and  orang  are 
curiously  like  those  of  man,  and  the  proper  muscles  are  likewise 
but  very  slightly  developed. "°  I  am  also  assured  by  the  keepers  in 
the  Zoological  Gardens  that  these  animals  never  move  or  erect 
their  ears ;  so  that  they  are  in  an  equally  rudimentary  condition 
with  those  of  man,  as  far  as  function  is  concerned.  Why  these 
animals,  as  well  as  the  progenitors  of  man,  should  have  lost  the 
power  of  erecting  their  ears,  we  cannot  say.  It  may  be,  though 
I  am  not  satisfied  with  this  view,  that  owing  to  their  arboreal 
habits  and  great  strength  they  were  but  little  exposed  to  danger, 
and  so  during  a  lengthened  period  moved  their  ears  but  little, 
and  thus  gradually  lost  the  power  of  moving  them.  This 
would  be  a  parallel  case  with  that  of  those  large  and  heavy 
birds,  which,  from  inhabiting  oceanic  islands,  have  not  been 
exposed  to  the  attacks  of  beasts  of  prey,  and  have  consequently 
lost  the  power  of  using  their  wings  for  flight.  The  inability  to 
move  the  ears  in  man  and  several  apes  is,  however,  partly  com- 
pensated by  the  freedom  with  which  they  can  move  the  head  in 

2*  Canestrini  quotes  Hyrtl.  ('  An-  lately   been   experimenting    on   the 

nuario    della    Soc.    dei  Naturalist!, '  function    of   the   shell    of  the    ear, 

Modena,  1867,  p.  97)  to   the  same  and   has   come    to  nearly  the  same 

effect.  conclusion  as  that  given  here. 

"^^  '  The  Diseases  of  the  Ear,'  by  ^o  p^-of.   A.    Macalister,    '  Annals 

J.    Toynbee,    F.R.S.,    1860,    p.    12.  and  Mag.  of  Nat.  History,'  vol,  vii.. 

A  distinguished   physiologist.  Prof.  1871,  p.  342. 
Preyer,    informs    me    that    he    had 


Chap.  I. 


Rudiments. 


15 


a  horizontal  plane,  so  as  to  catch  sounds  from  all  directions.  It 
lias  been  asserted  that  the  ear  of  man  alone  possesses  a  lobule ; 
but  "a  rudiment  of  it  is  found  in  the  gorilla ;"^^  and,  as  I  hear 
from  Prof.  Prefer,  it  is  not  rarely  absent  in  the  negro. 

The  celebrated  sculptor,  Mr.  Woolner,  informs  me  of  one  little 
peculiarity  in  the  external  ear,  which  he  has  often  observed  both 
in  men  and  women,  and  of  which  he  perceived  the  lull  signi- 
ficance. His  attention  was  first  called  to  the  subject  whilst  at 
work  on  his  figure  of  Puck,  to  which  he  had  given  pointed  ears. 
He  was  thus  led  to  examine  the  ears  of  various  monkeys,  and  sub- 
sequently more  carefully  those  of  man.  The  peculiarity  consists 
in  a  little  blunt  point,  projecting  from  the  inwardly  folded  margin, 
or  helix.  WHien  present,  it  is  developed  at  birth,  and,  iiccording 
to  Prof.  Ludwig  Meyer,  more  frequently  in  man  than  in  woman. 
Mr.  Woolner  made  an  exact  model  of  one  such  case,  and  sent  me 
the  accompanying  drawing.  (Fig.  2.) 
These  points  not  only  project  inwards 
towards  the  centre  of  the  ear,  but  often 
a  little  outwards  from  its  plane,  so  as 
to  be  visible  when  the  head  is  viewed 
from  directly  in  front  or  behind.  They 
are  variable  in  size,  and  somewhat  in 
position,  standing  either  a  little  higher 
or  lower;  and  they  sometimes  occur 
on  one  ear  and  not  on  the  other.  They 
are  not  confined  to  mankind,  for  I  ob- 
served a  case  in  one  of  the  spider- 
monkeys  (Ateles  beehfhufh)  in  our 
Zoological  Gardens;  and  Dr.  E.  Pay 
Lankester  informs  me  of  another  case 
in  a  chimpanzee  in  the  gardens  at 
Hamburg.  The  helix  obviously  con- 
sists of  the  extreme  margin  of  the  ear  folded  inwards;  and 
this  folding  appears  to  be  in  some  manner  connected  with  the 
whole  external  ear  being  permanently  pressed  backwards.  In 
many  monkeys,  which  do  not  stand  high  in  the  order,  as  baboons 
and  some  species  of  macacus,^^  the  upper  portion  of  the  ear  is 
slightly  pointed,  and  the  margin  is  not  at  all  folded  inwards ; 
but  if  the  margin  were  to  be  thus  folded,  a  slight  point  would 
necessarily  project  inwards  towards  the  centre,  and  probably  a 
little  outwards  from  the  plane  of  the  ear ;  and  this  I  believe  to 

^*  Mr.  St.  George    Mivart,   '  Ele-  Lemuroidea,    in    Messrs.   Murie  and 

mentary  Anatomy,'  1873,  p.  396.  ]\Iivart's   excellent  paper   in  'Tran« 

32  See    also    some    remarks,    and  sact.  Zoolog.  Soc'  vol.  vii.  1869,  pp. 

the  drawings    of    the    ears    of   the  6  and  90. 


Fig  2.     Unman  Ear.  morielled 

and  drawn  by  Mr.  Woolner. 

a.  The  projecting  point. 


1 6  TJie  Descent  of  Man.  Part  I. 

be  their  origin  in  many  cases.  On  the  other  hand,  Prof.  L.  Meyer, 
in  an  able  paper  recently  pubhshed,^  maintains  that  the  whole 
case  is  one  of  mere  variability ;  and  that  the  projections  are  not 
real  ones,  but  are  due  to  the  internal  cartilage  on  each  side  of 
the  points  not  having  been  fully  developed.  I  am  quite  ready 
to  admit  that  this  is  the  correct  explanation  in  many  instances, 
as  in  those  figured  by  Prof.  Meyer,  in  which  there  are  several 
minute  points,  or  the  whole  margin  is  sinuous.  I  have  myself 
seen,  through  the  kindness  of  Dr.  L.  Down,  the  ear  of  a  micro- 
cephalous idiot,  on  which  there  is  a  projection  on  the  outside 
of  the  helix,  and  not  on  the  inward  folded  edge,  so  that  this 
point  can  have  no  relation  to  a  former  apex  of  the  ear.  Never- 
theless in  some  cases,  my  original  view,  that  the  points 
are  vestiges  of  the  tips  of  formerly  erect  and  pointed  ears, 
still  seems  to  me  probable.  I  think  so  from  the  frequency  of 
their  occurrence,  and  from  the  general  correspondence  in 
position  with  that  of  the  tip  of  a  pointed  ear.  In  one  case,  of 
which  a  photograph  has  been  sent  me,  the  projection  is  so  large, 
that  supposing,  in  accordance  with  Prof.  Meyer's  view,  the  ear 
to  be  made  perfect  by  the  equal  development  of  the  cartilage 
throughout  the  whole  extent  of  the  margin,  it  would  have 
covered  fully  one-third  of  the  whole  ear.  Two  cases  have  been 
communicated  to  me,  one  in  North  America,  and  the  other  in 
England,  in  wliich  the  upper  margin  is  not  at  all  folded  inwards, 
but  is  pointed,  so  that  it  closely  resembles  the  pointed  ear  of  an 
ordinary  quadruped  in  outline.  In  one  of  these  cases,  which  was 
that  of  a  young  child,  the  father  compared  the  ear  with  the 
drawing  which  I  have  given^^  of  the  ear  of  a  monkey,  the 
Cynopithecus  rdger,  and  says  that  their  outlines  are  closely 
similar.  If,  in  these  two  cases,  the  margin  had  been  folded 
inwards  in  the  normal  manner,  an  inward  projection  must  have 
been  formed.  I  may  add  that  in  two  other  cases  the  outline  still 
remains  somewhat  pointed,  although  the  margin  of  the  upper 
part  of  the  ear  is  normally  folded  inwards — in  one  of  them, 
however,  very  narrowly.  The  following  woodcut  (No.  3)  is  an 
accurate  copy  of  a  photograph  of  the  foetus  of  an  orang  (kindly 
sent  me  by  Dr.  Nitsche),  in  which  it  may  be  seen  how  different  the 
pointed  outline  of  the  ear  is  at  this  period  from  its  adult  condition, 
when  it  bears  a  close  general  resemblance  to  that  of  man.  It  is 
evident  that  the  folding  over  of  the  tip  of  such  an  ear,  unless  it 
changed  greatly  during  its  further  development,  would  give  rise 
to  a  point  projecting  inwards.     On  the  whole,  it  still  seems  to 

'^  Ueber  das  Darwin'sche  Spitzohr,  ^*  'The  Expression  of   the   Emo- 

Archiv  fiir  Path.     Anat.  und  Phys.      tions,'  p.  136. 
1871,  p.  485. 


Chap.  I. 


Riidiinents. 


17 


me  probable  that  the  points  in  question  are  in  some  cases,  both 
in  man  and  apes,  vestiges  of  a  former  condition. 


Fig  3.     Foetus  of  an  OranR.    Fxact  copy  of  a  photograph,  shewing  the  form  of 
the  ear  at  this  early  age. 

The  nictitating  membrane,  or  third  eyelid,  with  its  accessory 
mnscles  and  other  structures,  is  especially  well  developed  in 
birds,  and  is  of  much  functional  importance  to  them,  as  it  can 
be  rapidly  drawn  across  the  whole  eye-ball.  It  is  found  in  some 
reptiles  and  amphibians,  and  in  certain  fishes,  as  in  sharks.  It 
is  fairly  well  developed  in  the  two  lower  divisions  of  the  mam- 
malian series,  namely,  in  the  monotreraata  and  marsupials,  and 
in  some  few  of  the  higher  mammals,  as  in  the  walrus.  But  in 
man,  the  quadruraana,  and  most  other  mammals,  it  exists,  as  is 
admitted  by  all  anatomists,  as  a  mere  rudiment,  called  the 
semilunar  fold.^^ 

The  sense  of  smell  is  of  the  highest  importance  to  the  greater 
number  of  mammals— to  some,  as  the  ruminants,  in  warning 
them  of  danger;  to  others,  as  the  carnivora,  in  finding  their 
prey;  to  others,  again,  as  the  wild  boar,  for  both  purposes 
combined.  But  the  sense  of  smell  is  of  extremely  slight  service, 
if  any,  even  to  the  dark  coloured  races  of  men,  in  whom  it  is 


"  Miillev's  'Elements  of  Physi- 
ology,* Eng.  translat.,  1842.  vol.  ii. 
p.  1117.  Owen,  *  Anatomy  of  Verte- 
brates,' vol.  iii.  p.  260;  ibid,  on 
the  Walrus,  *  Proo.  Zoolog.  Soc' 
November   8th,   1854.     See  also  R. 


Knox,  'Great  Artists  and  Anato- 
mists,' p.  106.  This  rudiment  ap- 
parently is  somewhat  larger  in 
Negroes  and  Australians  than  in 
Europeans,  see  Carl  Vogt,  '  Lectures 
on  Man,'  Eng.  translat.  p.  129. 


1 8  The  Descent  of  Man.  Part  I. 


much  more  highly  deyeloped  than  in  the  white  and  civilised 
races."^  Nevertheless  it  does  not  warn  them  of  danger,  nor  guide 
them  to  their  food ;  nor  does  it  prevent  the  Esquimaux  from 
sleeping  in  the  most  fetid  atmosphere,  nor  many  savages  from 
eating  half-putrid  meat.  In  Europeans  the  power  differs  greatly 
in  different  individuals,  as  I  am  assured  by  an  eminent  naturalist 
who  possesses  this  sense  highly  developed,  and  who  has  at- 
tended to  the  subject.  Those  who  believe  in  the  principle 
of  gradual  evolution,  will  not  readily  admit  that  the  sense  of 
smell  in  its  present  state  was  originally  acquired  by  man,  as 
he  now  exists.  He  inherits  the  power  in  an  enfeebled  and 
so  far  rudimentary  condition,  from  some  early  progenitor,  to 
whom  it  was  highly  serviceable,  and  by  whom  it  was  con- 
tinually used.  In  those  animals  which  have  this  sense  lughly 
developed,  such  as  dogs  and  horses,  the  recollection  of  persons 
and  of  places  is  strongly  associated  with  their  odour ;  and  we  can 
thus  perhaps  understand  how  it  is,  as  Dr.  Maudsley  has  truly 
remarked,^^  that  the  sense  of  smell  in  man  "  is  singularly  effective 
"  in  recalHng  vividly  the  ideas  and  images  of  forgotten  scenes 
"  and  places." 

Man  differs  conspicuously  from  all  the  other  Primates  in  being 
almost  naked.  But  a  few  short  straggling  hairs  are  found  over 
the  greater  part  of  the  body  in  the'mau,  and  fine  down  on  that 
of  the  woman.  The  different  races  differ  much  in  hairiness ;  and 
in  the  individuals  of  the  same  race  the  hairs  are  highly  variable, 
not  only  in  abundance,  but  likewise  in  position :  thus  in  some 
Europeans  the  shoulders  are  quite  naked,  whilst  in  others  they 
bear  thick  tufts  of  hair.^^  There  can  be  little  doubt  that  the 
hairs  thus  scattered  over  the  body  are  the  rudiments  of  the 
uniform  hairy  coat  of  the  lower  animals.  This  view  is  rendered 
all  the  more  probable,  as  it  is  known  that  fine,  short,  and  pale- 
coloured  hairs  on  the  limbs  and  other  parts  of  the  body,  occasion- 

^^  The  account  given  by  Humboldt  olfactory  region,  as  well  as  of  the 
of  the  power  of  smell  possessed  by  skin  of  the  body.  I  have,  therefore, 
the  natives  of  South  America  is  well  spoken  in  the  text  of  the  dark- 
known,  and  has  been  confirmed  by  coloured  races  having  a  finer  sense 
others.  M.  Hoiizeau  ('  Etudes  sur  of  smell  than  the  white  races.  See 
les  Faculte's  Mentales,'  &c.,  torn.  i.  his  paper, '  Medico-Chirurgical  Tran- 
1872,  p.  91)  asserts  that  he  re-  sactions,'  London,  vol.  liii.,  1870, 
peatedly     made     experiments,     and  p.  276. 

proved     that    Negroes    and    Indians  ^^  '  The  Physiology  and  Pathology 

could  recognise  persons  in  the  dark  of  Mind,'  2nd  edit.  1868,  p.  134-. 

by  their  odour.     Dr.    W.  Ogle   has  ^s  Eschricht,  Ueber  die  Kichtung 

made  some  curious  observations  on  der  Haare  am  menschlichen  Korper, 

the   connection  between   the  power  'Miiller's  Archivfur  Anat.und  Phys.' 

of  smell  and  the  colouring  matter  1837,  s.  47.     I  shall  often  have  to 

of    the   mucous   membrane    of  the  refer  to  this  very  curious  paper. 


Chap.  I.  Rudiments.  19 


ally  become  developed  into  "  thickset,  long,  and  rather  coarse 
"  dark  hairs,"  when  abnormally  nourished  near  old-standing 
inflamed  surfaccs.^^ 

I  am  informed  by  Sir  James  Paget  that  often  several  members 
of  a  family  have  a  few  hairs  in  their  eyebrows  much  longer  than 
the  others;  so  that  even  this  slight  peculiarity  seems  to  be 
inherited.  These  hairs,  too,  seem  to  have  their  repiesentatives ; 
for  in  the  chimpanzee,  and  in  certain  sjDecies  of  Macacus,  there 
are  scattered  hairs  of  considerable  length  rising  from  the  naked 
skin  above  the  eyes,  and  corresponding  to  our  eyebrows  ;  similar 
long  hairs  project  from  the  hairy  covering  of  the  superciliary 
ridges  in  some  baboons. 

The  fine  wool-like  hair,  or  so-called  lanugo,  with  which  the 
human  foetus  during  the  sixth  month  is  thickly  covered,  ofifers  a 
more  curious  case.  It  is  first  developed,  daring  the  fifth  month, 
on  the  eyebrows  and  face,  and  especially  round  the  mouth, 
where  it  is  much  longer  than  that  on  the  head,  A  moustache 
of  this  kind  was  observed  by  Eschriclit^°  on  a  female  foetus ;  but 
this  is  not  so  surprising  a  circumstance  as  it  may  at  first  appear, 
for  the  two  sexes  generally  resemble  each  other  in  all  external 
characters  during  an  early  period  of  growth.  The  direction  and 
arrangement  of  the  hairs  on  all  parts  of  the  foetal  body  are  the 
same  as  in  the  adult,  but  are  subject  to  much  variability.  The 
whole  surface,  including  even  the  forehead  and  ears,  is  thus 
tliickly  clothed ;  but  it  is  a  significant  fact  that  the  palms  of  the 
hands  and  the  soles  of  the  feet  are  quite  naked,  like  the  inferior 
surfaces  of  all  four  extremities  in  most  of  the  lower  animals.  As 
this  can  hardly  be  an  accidental  coincidence,  the  woolly  cover- 
ing of  the  foetus  probably  repres(;nts  the  first  permanent  coat  of 
hair  in  those  mammals  which  are  born  hairy.  Three  or  four 
cases  have  been  recorded  of  persons  born  with  their  whole  bodies 
and  faces  thickly  covered  with  fine  long  hairs ;  and  this  strange 
condition  is  strongly  inherited,  and  is  correlated  with  an  abnor- 
mal condition  of  the  teeth.'*^  Prof.  Alex.  Brandt  informs  me  that 
he  has  compared  the  hair  from  the  face  of  a  man  thus  charac- 
terised, aged  thirty-five,  with  the  lanugo  of  a  foetus,  and  finds  it 
quite  similar  in  texture ;  therefore,  as  he  remarks,  the  case  may 
be  attributed  to  an  arrest  of  development  in  the  hair,  together 
with  its  continued  growth.     Many  delicate  children,  as  I  have 

^'  Paget,  'Lectures    on   Sui'gical  has  recently  sent  me  an  additional 

Pathology,'  1853,  vol.  i.  p.  71.  case   of  a   father  and  son,  born  in 

■*°  Eschrichi,  ibid.  s.  40,  47.  Piussia,  with   these  peculi;irities.     I 

**  See  my  '  Variation  of  Animals  have  received  drawings  of  both  from 

and    Plants    under    Domestication,'  Paris. 

vol.  ii.  p.  327.     Prof.  Alex.   Brandt 


20  The  Descent  of  Man.  Part  I. 

been  assured  by  a  surgeon  to  a  hospital  for  children,  have  their 
backs  covered  by  rather  long  silky  hairs ;  and  such  cases  pro- 
bably come  under  the  same  head. 

It  appears  as  if  the  posterior  molar  or  wisdom-teeth  were 
tending  to  become  rudimentary  in  the  more  civilised  races  of 
man.  These  teeth  are  rather  smaller  than  the  other  molars,  as 
is  likewise  the  case  with  the  corresponding  teeth  in  the  chim- 
panzee and  orang ;  and  they  have  only  two  separate  fangs. 
They  do  not  cut  through  the  gums  till  about  the  seventeenth 
year,  and  I  have  been  assured  that  they  are  much  more  liable  to 
decay,  and  are  earlier  lost  than  the  other  teeth ;  but  this  is  denied 
by  some  eminent  dentists.  They  are  also  much  more  liable  to 
vary,  both  in  structure  and  in  the  period  of  their  development, 
than  the  other  teeth.'*'^  In  the  Melanian  races,  on  the  other 
hand,  the  wisdom-teeth  are  usually  furnished  with  three 
separate  fangs,  and  are  generally  sound ;  they  also  differ  from 
the  other  molars  in  size,  less  than  in  the  Caucasian  races.*^ 
Prof.  Schaaffhausen  accounts  for  this  difference  between  the 
races  by  "  the  posterior  dental  portion  of  the  jaw  being  always 
"  shortened"  in  those  that  are  civilised,^*  and  this  shortening  may, 
I  presume,  be  attributed  to  civilised  men  habitually  feeding  on 
soft,  cooked  food,  and  thus  using  their  jaws  less.  I  am  informed 
by  Mr.  Brace  that  it  is  becoming  quite  a  common  practice  in  the 
United  States  to  remove  some  of  the  molar  teeth  of  children,  as 
the  jaw  does  not  grow  large  enough  for  the  perfect  development 
of  the  normal  number.^^ 

With  respect  to  the  alimentary  canal,  I  have  met  with  an 
account  of  only  a  single  rudiment,  namely  the  vermiform  append- 
age of  the  caecum.  The  caecum  is  a  branch  or  diverticulum  of 
the  intestine,  ending  in  a  cul-de-sac,  and  is  extremely  long  in 
many  of  the  lower  vegetable-feeding  mammals.  In  the  marsupial 
koala  it  is  actually  more  than  thrice  as  long  as  the  whole  body.^^ 
It  is  sometimes  produced  into  a  long  gradually-tapering  point, 
and  is  sometimes  constricted  in  parts.  It  appears  as  if,  in  con- 
sequence of  changed  diet  or  habits,  the  caecum  had  become  much 

42  Dr.  Webb,  'Teeth   in  Man  and  from  Florence,  that  he    has   lately 

the  Anthropoid  Apes,'  as  quoted  by  been  studying  the  last  molar  teeth 

Di*.  C.  Carter  Blake  in  '  Anthropo-  in   the  different  races  of  man,  and 

logical  Review,'  July  1867,  p.  299.  has  come  to  the  same  conclusion  as 

■*3  Owen,     '  Anatomy     of    Verte-  that  given  in  my  text,  viz.,  that  in 

brates,'  vol.  iii.  pp.   320,  321,  and  the  higher  or   civilised   races    they 

325.  are  on  the  road  towards  atrophy  or 

**  'On  the  Primitive  Form  of  the  elimination. 

Skull,' Eng.  translat.  in  'Anthropo-  ^^  Owen,    'Anatomy    of    Verte- 

logical  Review,'  Oct.  1868,  p.  426.  brates,'  vol.  iii.  pp.  416,  434,  441. 

^=  Prof.  Montegazza  writes  to  me 


Chap.  I.  Rudiments.  21 

shortened  in  various  animals,  the  vermiform  appendage  being 
left  as  a  rudiment  of  the  shortened  part.  That  this  appendage 
is  a  rudiment,  we  may  infer  from  its  small  size,  and  from  the 
evidence  which  Prof.  Canestrini*^  has  collected  of  its  variability 
in  man.  It  is  occasionally  quite  absent,  or  again  is  largely 
developed.  The  passage  is  sometimes  completely  closed  for  half 
or  two-thirds  of  its  length,  with  the  terminal  part  consisting  of 
a  flattened  solid  expansion.  In  the  orang  this  appendage  is  long 
and  convoluted:  in  man  it  arises  from  the  end  of  the  short 
caecum,  and  is  commonly  from  four  to  five  inches  in  length, 
being  only  about  the  third  of  an  inch  in  diameter.  Kot  only  is 
it  useless,  but  it  is  sometimes  the  cause  of  death,  of  which  fact 
I  have  lately  heard  two  instances :  this  is  due  to  small  hard 
bodies,  such  as  seeds,  entering  the  passage,  and  causing  inflam- 
mation."*^ 

In  some  of  the  lower  Quadrumana,  in  the  Lemurida3  and 
Carnivora,  as  well  as  in  many  marsupials,  there  is  a  passage  near 
the  lower  end  of  the  humerus,  called  the  supra-condyloid  fora- 
men, through  which  the  great  nerve  of  the  fore  limb  and  often 
the  great  artery  pass.  Now  in  the  humerus  of  man,  there  is 
generally  a  trace  of  this  passage,  which  is  sometimes  fairly  well 
developed,  being  formed  by  a  depending  hook -like  process  of 
bone,  completed  by  a  band  of  ligament.  Dr.  Struthers,''^  who  has 
closely  attended  to  the  subject,  has  now  shewn  that  this 
peculiarity  is  sometimes  inherited,  as  it  has  occurred  in  a  father, 
and  in  no  less  than  four  out  of  his  seven  children.  When  pre- 
sent, the  great  nerve  invariably  passes  through  it;  and  this 
clearly  indicates  that  it  is  the  homologue  and  rudiment  of  the 
supra-condyloid  foramen  of  the  lower  animals.  Prof.  Turner 
estimates,  as  he  informs  me,  that  it  occurs  in  about  one  per  cent, 
of  recent  skeletons.  But  if  the  occasional  development  of  this 
structure  in  man  is,  as  seems  probable,  due  to  reversion,  it  is  a 
return  to  a  very  ancient  state  of  things,  because  in  the  higher 
Quadrumana  it  is  absent. 

There  is  another  foramen  or  perforation  in  the  humerus, 

<^  'Annuario   della  Soc,  d.  Nat'  Feb.    15,    1873,    and    another    im- 

Modena,  1867,  p.  94.  portant  paper,  ibid.,  Jan.  24,  1863, 

■"8  M.   C.    Martins    (*'  De    I'Unite  p.  83.     Dr.  Knox,  as  I  am  informed, 

Orgauique,"    in    '  Revue    des    Deux  was  the  first  anatomist  who    drew 

Moudes,'  June  15,  1862,  p.  16),  and  attention  to  this  peculiar  structure 

Hackel    ('  Generelle     Morphologic,'  in  man  ;  see  his  '  Great  Artists  and 

B.  ii.  s.  278),  have   both    remarked  Anatomists,'  p.  63.     See  also  an  im- 

on  the   singular   fact  of   this  rudi-  portant  memoir  on  this  process  by 

ment  sometimes  causing  death.  Dr.    Gruber,    in    the    '  Bulletin    de 

**  With    respect    to    inheritance,  I'Acad.    Imp.   de    St.    Pe'tersbourg,' 

see  Dr.  Struthers  in  the  '  Lancet,'  tom.  xii.  1867,  p.  448. 


22  The  Descent  of  Man.  Part  I. 

occasionally  present  in  man,  which  may  be  called  the  inter- 
condyloid.  This  occurs,  but  not  constantly,  in  various  anthro- 
poid and  other  apes,^"  and  likewise  in  many  of  the  lower  animals. 
It  is  remarkable  that  this  perforation  seems  to  have  been  present 
in  man  much  more  frequently  during  ancient  times  than 
recently.  Mr.  Busk^^  has  collected  the  following  evidence  on 
this  head :  Prof.  Broca  "  noticed  the  perforation  in  four  and  a 
"half  per  cent,  of  the  arm-bones  collected  in  the  '  Cimetiere  du 
"  Sud,'  at  Paris ;  and  in  the  Grotto  of  Orrony,  the  contents  of 
"  which  are  referred  to  the  Bronze  period,  as  many  as  eight 
"  humeri  out  of  thirty-two  were  perforated ;  but  this  extraordi- 
"  nary  proportion,  he  thinks,  might  be  due  to  the  cavern  having 
"  been  a  sort  of  '  family  vault.'  Again,  M.  Dupont  found  thirty 
"  per  cent,  of  perforated  bones  in  the  caves  of  the  Valley  of  the 
''  Lesse,  belonging  to  the  Eeindeer  period  ;  whilst  M.  Leguay,  in 
"  a  sort  of  dolmen  at  Argenteuil,  observed  twenty -five  per  cent, 
"to  be  perforated;  and  M.  Pruner-Bey  found  twenty-six  per 
"  cent,  in  the  same  condition  in  bones  from  Yaureal.  Nor  should 
''it  be  left  unnoticed  that  M.  Pruner-Bey  states  that  this  con- 
'*  dition  is  common  in  G-uanche  skeletons."  It  is  an  interesting 
fact  that  ancient  races,  in  this  and  several  other  cases,  more 
frequently  present  structures  which  resemble  those  of  the  lower 
animals  than  do  the  modern.  One  chief  cause  seems  to  be  that 
the  ancient  races  stand  somewhat  nearer  in  the  long  line  of 
descent  to  their  remote  animal-like  progenitors. 

In  man,  the  os  coccyx,  together  with  certain  other  vertebrae 
hereafter  to  be  described,  though  functionless  as  a  tail,  plainly 
represent  this  part  in  other  vertebrate  animals.  At  an  early 
embryonic  period  it  is  free,  and  projects  beyond  the  lower 
extremities ;  as  may  be  seen  in  the  drawing  (Fig.  1.)  of  a  human 
embryo.  Even  after  birth  it  has  been  known,  in  certain  rare 
and  anomalous  cases,^^  to  form  a  small  external  rudiment  of  a 
tail.  The  os  coccyx  is  short,  usually  including  only  four 
vertebrEe,  all  anchylosed  together:    and  these  are  in  a  rudi- 

*"  Mr.  St.  George  Mivart, '  Trans-  ^2  Quatrefages  has  lately  collected 

act.  Phil.  Soc'  1867,  p.  310.  the  evidence  on  this  subject.    '  Revue 

"  "On  the  Caves  of  Gibraltar,"  des  Cours  Scientifiques,'  1867-1868, 

'Transact.    Internat.    Congress     of  p.  625.     In   1840    Fleischmann  ex- 

Prehist.  Arch.'  Third  Session,  1869,  hibited    a    human   foetus    bearing  a 

p.    159.     Prof.    Wyman    has    lately  free  tail,  which,  as  is  not  always  the 

shewn  (Fourth  Annual  Report,  Pea-  case,  included  vertebral  bodies ;  and 

body  Museum,  1871,  p.  20),  that  this  this  tail  was  critically  examined  by 

perforation  is  present  in  thirty-one  the  many  anatomists  present  at  the 

per    cent,    of  some  human  remains  meeting   of  naturalists  at  Erlangen 

from  ancient  mounds  in  the  Western  (see    Marshall    in    Niederltindischen 

United  States,  and  in  Florida.  It  Archivfiir  Zoologie,  Decemberl871). 
frequently  occurs  in  the  negro. 


Chap.  I.  Rudimejits.  23 


mentary  condition,  for  they  consist,  with  the  exception  of  tlie 
basal  one,  of  the  centrum  alone.^^  They  are  furnished  with 
some  small  muscles ;  one  of  which,  as  I  am  informed  by  Prof. 
Turner,  has  been  exj^ressly  described  by  Theile  as  a  rudimentary 
repetition  of  the  extensor  of  the  tail,  a  muscle  which  is  so 
largely  developed  in  many  mammals. 

The  spinal  cord  in  man  extends  only  as  far  downwards  as  the 
last  dorsal  or  first  lumbar  vertebra;  but  a  thread-like  struc- 
ture {i\\Q.  filum  ttrminah-^  runs  down  the  axis  of  the  sacral  part 
of  the  spinal  canal,  and  even  along  the  back  of  the  coccygeal 
bones.  The  upper  part  of  this  filament,  as  Prof.  Turner 
informs  me,  is  undoubtedly  homologous  with  the  spinal  cord ; 
but  the  lower  part  apparently  consists  merely  of  the  pia  mater, 
or  vascular  investing  membrane.  Even  in  this  case  the  os 
coccyx  may  be  said  to  possess  a  veslige  of  so  important  a 
structure  as  the  spinal  cord,  though  no  longer  enclosed  within 
a  bony  canal.  The  following  fact,  for  which  I  am  also  in- 
debted to  Prof.  Turner,  shews  how  closely  the  os  coccyx  corre- 
sponds with  the  true  tail  in  the  lower  animals  :  Luschka  has 
recently  discovered  at  the  extremity  of  the  coccygeal  bones  a 
very  peculiar  convoluted  body,  which  is  continuous  with  the 
middle  sacral  artery ;  and  this  discovery  led  Krause  and  Meyer 
to  examine  the  tail  of  a  monkey  (Macacus),  and  of  a  cat,  in  both 
of  which  they  found  a  similarly  convoluted  body,  though  not  at 
the  extremity. 

The  reproductive  system  offers  various  rudimentary  struc- 
tures; but  these  differ  in  one  important  respect  from  the 
foregoing  cases.  Here  we  are  not  concerned  with  the  vestige  of 
a  part  which  does  not  belong  to  the  species  in  an  efficient  state, 
but  with  a  part  efficient  in  the  one  sex,  and  represented  in  the 
other  by  a  mere  rudiment.  Nevertheless,  the  occurrence  of 
such  rudiments  is  as  difficult  to  explain,  on  the  belief  of  the 
separate  creation  of  each  species,  as  in  the  foregoing  cases. 
Hereafter  I  shall  have  to  recur  to  these  rudiments,  and  shall 
shew  that  their  presence  generally  depends  merely  on  inheri- 
tance, that  is,  on  parts  acquired  by  one  sex  having  been 
partially  transmitted  to  the  other.  I  will  in  this  place  only  give 
somQ  instances  of  such  rudiments.  It  is  well  known  that  in  the 
males  of  all  mammals,  including  man,,  rudimentary  mammae 
exist.  These  in  several  instances  have  become  well  developed, 
and  have  yielded  a  copious  supply  of  milk.  Their  essential 
identity  in  the  two  sexes  is  likewise  shewn  by  their  occasional 
sympathetic  enlargement  in  both    during  an    attack  of    the 

"  Owen,  'On  the  Nature  of  Limbs,'  1849,  p.  114. 


24  The  Desceiit  of  Mmi.  Paet  I. 

measles.  The  vesicula  prostatlca,  which  has  been  observed  in 
many  male  mammals,  is  now  universally  acknowledged  to  bo 
the  homologne  of  the  female  uterus,  together  with  the  con- 
nected passage.  It  is  impossible  to  read  Leuckart's  able 
de'scription  of  this  organ,  and  his  reasoning,  without  admitting 
the  justness  of  his  conclusion.  This  is  especially  clear  in  the 
case  of  those  mammals  in  which  the  true  female  uterus 
bifurcates,  for  in  the  males  of  these  the  vesicula  likewise 
bifurcates.^*  Some  other  rudimentary  structui-es  belonging  to 
the  reproductive  system  might  have  been  here  adduced.  ^^ 

The  bearing  of  the  three  great  classes  of  facts  now  given  is 
unmistakeable.  But  it  would  be  superfluous  fully  to  recapitulate 
the  line  of  argument  given  in  detail  in  my  '  Origin  of  Species.' 
The  homological  construction  of  the  whole  frame  in  the  members 
of  the  same  class  is  intelligible,  if  we  admit  their  descent  from 
a  common  progenitor,  together  with  their  subsequent  adaptation 
to  diversified  conditions.  On  any  other  view,  the  similarity  of 
pattern  between  the  hand  of  a  man  or  monkey,  the  foot  of  a 
horse,  the  flipper  of  a  seal,  the  wing  of  a  bat,  &c.,  is  utterly 
inexplicable,^^  It  is  no  scientific  explanation  to  assert  that  they 
have  all  been  formed  on  the  same  ideal  plan.  With  respect  to 
development,  we  can   clearly  understand,  cm  the  principle  of 

'*  Leuckart,    in   Todd's    'Cyclop.  words)  a   mere  metaphysical  prin- 

vf  Anat.'  1849-52,  vol.  iv.  p.  1415.  ciple,  namely,  the  preservation  "in 

In    man    this   organ    is   only    from  "  its    integrity   of  the    mammalian 

three   to   six   lines  in   length,   but,  "  nature  of  the  animal."     In  only  a 

like    so    many    other    rudimentary  few  cases  does  he  discuss  rudiments, 

parts,  it  is  variable  in  development  and  then  only  those  parts  which  are 

as  well  a^  in  other  characters.  partially  rudimentary,  such  as   the 

5^    See,    on    this    subject,    Owen,  little  hoofs  of  the  pig  and  ox,  which 

'Anatomy  of  Vertebrates,'  vol.  iii.  do  not  touch  the  ground;  these  he 

pp.  675,  676,  706.  shews  clearly  to  be  of  service  to  the 

^s  Prof.  Bianconi,  in  a  recently  animal.  It  is  unfortunate  that  he 
published  work,  illustrated  by  ad-  did  not  consider  such  cases  as  the 
mirable  engravings  ('  La  The'orie  minute  teeth,  which  never  cut 
Darwiuienne  et  la  creation  dite  in-  through  the  jaw  in  the  ox,  or  the 
de'pendante,'  1874),  endeavours  to  mammae  of  male  quadrupeds,  or  the 
show  that  homological  structures,  in  wings  of  certain  beetles,  existing 
the  above  and  other  cases,  can  be  under  the  soldered  wing-covers,  or 
fully  explained  on  mechanical  prin-  the  A'estiges  of  the  pistil  and  stamens 
ciples,  in  accordance  with  their  uses,  in  various  flowers,  and  man}f  other 
No  one  has  shewn  so  well,  how  ad-  such  cases.  Although  1  greatly 
mirably  such  structures  are  adapted  admire  Prof.  Bianconi's  work,  yet 
for  their  final  purpose;  and  this  the  belief  now  held  by  most  natural- 
adaptation  can,  as  I  believe,  be  ists  seems  to  me  left  unshaken, 
explained  through  natural  selection.  that  homological  structures  are  in- 
In  considering  the  wing  of  a'bat,  he  explicable  on  the  principle  of  mere 
brings  forward  (p.  218)  what  appears  adaptation. 
to    me    (to    use    Auguste    Comte's 


Chap.  I.  Rjidimetits.  2$ 


■variations  supervening  at  a  rather  late  embryonic  period,  and 
being  inherited  at  a  corresponding  period,  how  it  is  that  the 
embryos  of  wonderfully  different  forms  should  still  retain,  more 
or  less  perfectly,  the  structure  of  their  common  progenitor. 
No  other  explanation  has  ever  been  given  of  the  marvellous  fact 
that  the  embryos  of  a  man,  dog,  seal,  bat,  reptile,  &c.,  can  at  first 
hardly  be  distinguished  from  each  other.  In  order  to  understand 
the  existence  of  rudimentary  organs,  we  have  only  to  suppose 
that  a  former  progenitor  possessed  the  parts  in  question  in  a 
perfect  state,  and  that  under  changed  habits  of  life  thoy  became 
greatly  reduced, either  from  simple  disuse,  or  through  the  natural 
selection  of  those  individuals  which  were  least  encumbered  with 
a  superfluous  part,  aided  by  the  other  means  previously  in- 
dicated. 

Thus  we  can  understand  how  it  has  come  to  pass  that  man  and 
all  other  vertebrate  animals  have  been  constructed  on  the  same 
general  model,  why  they  pass  through  the  same  early  stages  of 
development,  and  why  they  retain  certain  rudiments  in  common. 
Consequently  we  ought  frankly  to  admit  their  community  of 
descent;  to  take  any  other  view,  is  to  admit  that  our  own 
structure,  and  that  of  all  the  animals  around  us,  is  a  mere  snare 
laid  to  entrap  our  judgment.  This  conclusion  is  greatly 
strengthened,  if  we  look  to  the  members  of  the  whole  animal 
series,  and  consider  the  evidence  derived  from  their  affinities 
or  classification,  their  geographical  distribution  and  geolo- 
gical succession.  It  is  only  our  natural  prejudice,  and  that 
arrogance  which  made  our  forefathers  declare  that  they  were 
descended  from  demi-gods,  which  leads  us  to  demur  to  this 
conclusion.  But  the  time  will  before  long  come,  when  it  will  be 
thought  wonderful  that  naturalists,  who  were  well  acquainted 
with  the  comparative  structure  and  development  of  man,  and 
other  mammals,  should  have  believed  that  each  was  the  work 
of  a  separate  act  of  creation. 


26  The  Descent  of  Man.  Part  L 


CEAPTEE  n. 

On  the  Manner  of  Development  of  Man  from  some 
Lower   Form. 

Variability  of  body  and  mind  in  man — Inheritance — Causes  of  variability 
— Laws  of  variation  the  same  in  man  as  in  the  lower  animals — Direct 
action  of  the  conditions  of  life — Effects  of  the  increased  use  and  disuse 
of  parts — Arrested  development — Reversion — Correlated  variation  — 
Rate  of  increase — Checks  to  increase — Natural  selection — Man  the  most 
dominant  animal  in  the  world — Importance  of  his  corporeal  structure — 
The  causes  which  have  led  to  his  becoming  erect — Consequent  changes 
of  structure — Decrease  in  size  of  the  canine  teeth — Increased  size  and 
altered  shape  of  the  skull — Nakedness — Absence  of  a  tail — Defenceless 
condition  of  man. 

It  is  manifest  that  man  is  now  subject  to  much  yariability. 
No  two  individuals  of  the  same  race  are  quite  alike.  We  may 
compare  millions  of  faces,  and  each  will  be  distinct.  There  is 
an  equally  great  amount  of  diversity  in  the  proportions  and 
dimensions  of  the  various  parts  of  the  body ;  the  length  of  the 
legs  being  one  of  the  most  variable  points.^  Although  in  some 
quarters  of  the  world  an  elongated  skull,  and  in  other  quarters 
a  short  skull  prevails,  yet  there  is  great  diversity  of  shape  even 
within  the  limits  of  the  same  race,  as  with  the  aborigines  of 
America  and  South  Australia — the  latter  a  race  *'  probably  as 
"  pure  and  homogeneous  in  blood,  customs,  and  language  as  any 
"  in  existence" — and  even  with  the  inhabitants  of  so  confined 
an  area  as  the  Sandwich  Islands. ^  An  eminent  dentist  assures 
me  that  there  is  nearly  as  much  diversity  in  the  teeth  as  in  the 
features.  The  chief  arteries  so  frequently  run  in  abnormal 
courses,  that  it  has  been  found  useful  for  surgical  purposes  to 
calculate  from  1040  corpses  how  often  each  course  prevails.' 
The  muscles  are  eminently  variable :  thus  those  of  the  foot 
were  found  by  Prof.  Turner  *  not  to  be  strictly  alike  in  any  two 
out  of  fifty  bodies ;  and  in  some  the  deviations  were  considerable. 

*  '  Investigations  in  Military  and  Huxley,  in  Lyell's  '  Antiquity  of 
Anthropolog.  Statistics  of  American  Man,'  1863,.  p.  87.  On  the  Sand- 
Soldiers,'  by  B.  A.  Gould,  1869,  p.  wich  Islanders,  Prof.  J.  Wyman, 
256.  '  Observations    on    Crania,'    Boston, 

2  With  respect    to    the  "Cranial  1868,  p.  18. 

forms  of  the  American  aborigines,"  '  'Anatomy  of  the  Arteries,'  by 

see    Dr.    Aitken    Meigs    in    '  Proc.  R.  Quain.     Preface,  vol.  i.  1844. 

Acad.  Nat.  Sci.'  Philadelphia,  May,  *  'Transact.    Royal    Soc.     Edin- 

1868.       On     the    Australians,    see  burgh,'  vol.  xxiv.  pp.  175,  189. 


Chap.  II.  Maimer  of  Development.  2^ 

He  adds,  that  the  power  of  performing  the  appropriate  move- 
ments must  have  been  modified  in  accordance  with  the  several 
deviations.  Mr.  J.  Wood  has  recorded^  the  occurrence  of  295 
muscular  variations  in  thirty-six  subjects,  and  in  another  set  of 
the  same  number  no  less  than  558  variations,  those  occurring  on 
both  sides  of  the  body  being  only  reckoned  as  one.  In  the  last 
set,  not  one  body  out  of  the  thirty-six  was  "  found  totally 
"  wanting  in  departures  from  the  standard,  descriptions  of  the 
"  muscular  sj^stem  given  in  anatomical  text  books."  A  single 
body  presented  the  extraordinary  number  of  twenty-five  distinct 
abnormalities.  The  same  muscle  sometimes  varies  in  many 
ways:  thus  Prof.  Macalister  describes*^  no  less  than  twenty 
distinct  variations  in  the  jpalmaru  accessor  ins. 

The  famous  old  anatomist,  Wolff,'^  insists  that  the  internal 
viscera  are  more  variable  than  the  external  parts :  Nulla  jparti- 
cala  est  qux  non  aliter  et  aliter  in  aliis  se  habeat  hominibus.  He 
has  even  written  a  treatise  on  the  choice  of  typical  examples  of 
the  viscera  for  representation.  A  discussion  on  the  beau-ideal 
of  the  liver,  lungs,  kidneys,  &c.,  as  of  the  human  face  divine, 
sounds  strange  in  our  ears. 

The  variability  or  diversity  of  the  mental  faculties  in  men  of 
the  same  race,  not  to  mention  the  greater  differences  between 
the  men  of  distinct  races,  is  so  notorious  that  not  a  word  need 
here  be  said.  So  it  is  with  the  lower  animals.  All  who  have 
had  charge  of  menageries  admit  this  fact,  and  we  see  it  plainly 
in  our  dogs  and  other  domestic  animals.  Brehm  especially 
insists  that  each  individual  monkey  of  those  which  he  kept  tame 
in  Africa  had  its  own  peculiar  disposition  and  temper ;  he  men- 
tions one  baboon  remarkable  for  its  high  intelligence ;  and  the 
keepers  in  the  Zoological  Gardens  pointed  out  to  me  a  monkey, 
belonging  to  the  New  World  division,  equally  remarkable  for 
intelligence.  Eengger,  also,  insists  on  the  diversity  in  the 
various  mental  characters  of  the  monkeys  of  the  same  species 
which  he  kept  in  Paraguay ;  and  this  diversity,  as  he  adds,  is 
partly  innate,  and  partly  the  result  of  the  manner  in  which  they 
have  been  treated  or  educated.^ 

I  have  elsewhere^  so  fully  discussed  the  subject  of  Inheritance, 
that  I  need  here  add  hardly  anything.     A  greater  number  of 

5  'Pioc.     Royal    Soc'     1867,    p.  »  Brehm,    '  Thierleben,'    B.    i.    s. 

544  ;  also  ]  868,  pp.  48.'3, 524.    There  58,  87.     Kengger,  '  Saugethiere  von 

is  a  previous  paper,  1866,  p.  229.  Paraguay,'  s.  57. 

*  '  Proc.  R.  Irish  Academy,'  vol.  '    '  Variation     of    Animals     an^ 

X.  1868,  p.  141.  Plants    under    Domestication,'    vol 

^  'Act.    Acad.    St.    Petersburg,*  ii.  chap.  xii. 
1778,  part  ii.  p.  217 


28  The  Descent  of  Man.  Part  L 

facts  have  been  collected  with  respect  to  the  transmission  of  the 
most  trifling,  as  well  as  of  the  most  important  characters  in 
man,  than  in  any  of  the  lower  animals;  though  the  facts  are 
copious  enough  with  respect  to  the  latter.  So  in  regard  to 
mental  qualities,  their  transmission  is  manifest  in  our  dogs, 
horses,  and  other  domestic  animals.  Besides  special  tastes  and 
habits,  general  intelligence,  courage,  bad  and  good  temper,  &c., 
are  certainly  transmitted.  With  man  we  see  similar  facts  in 
almost  every  family;  and  we  now  know,  through  the  admirable 
labours  of  Mr.  Galton,^°that  genius  which  implies  a  wonderfully 
complex  combination  of  high  faculties,  tends  to  be  inherited; 
and,  on  the  other  hand,  it  is  too  certain  that  insanity  and  deteri- 
orated mental  powers  likewise  run  in  families. 

With  respect  to  the  causes  of  variability,  we  are  in  all  cases 
very  ignorant;  but  we  can  see  that  in  man  as  in  the  lower 
animals,  they  stand  in  some  relation  to  the  conditions  to  which 
each  species  has  been  exposed,  during  several  generations. 
Domesticated  animals  vary  more  than  those  in  a  state  of  nature ; 
and  this  is  apparently  due  to  the  diversified  and  changing  nature 
of  the  conditions  to  which  they  have  been  subjected.  In  this 
respect  the  different  races  of  man  resemble  domesticated  animals, 
and  so  do  the  individuals  of  the  same  race,  when  inhabiting  a 
very  wide  area,  like  that  of  America.  We  see  the  influence  of 
diversified  conditions  in  the  more  civilised  nations;  for  the 
members  belonging  to  different  grades  of  rank,  and  following 
different  occupations,  present  a  greater  range  of  character  than 
do  the  members  of  barbarous  nations.  But  the  uniformity  of 
savages  has  often  been  exaggerated,  and  in  some  cases  can  hardly 
be  said  to  exist.^^  It  is,  nevertheless,  an  error  to  speak  of  man, 
even  if  we  look  only  to  the  conditions  to  which  he  has  been 
exposed,  as  "  far  more  domesticated  "^^  than  any  other  animal. 
Some  savage  races,  such  as  the  Australians,  are  not  exposed  to 
more  diversified  conditions  than  are  many  species  which  have 
a  wide  range.  In  another  and  much  more  important  respect, 
man  differs  widely  from  any  strictly  domesticated  animal ;  for 
his  breeding  has  never  long  been  controlled,  either  by  methodical 
or  unconscious  selection.     No  race  or  body  of  men  has  been  so 

'"  *  Hereditary    Genius  :    an    In-  "  man  had  an  oval  visage  with  fine 

quiry    into    its    Laws    and    Conse-  "  features,  and    another  was    quite 

quences,'  1869.  "  Mongolian    in    breadth    and    pro- 

'*  Mr.  Bates  remarks  (' The  Natu-  "  minence  of  cheek,  spread  of  nos- 

ralist  on  the  Amazons,'  1863,  vol.  ii.  "  trils,  and  obliquity  of  eyes." 

p.  159),  with  respect  to  the  Indians  ^^  Blumenbach,  'Treatises  on  An- 

of  the  same  South  American  tribe,  thropolog.'  Eng.  translat.,  1865,  p. 

*'  no  two  of  them  were  at  all  similar  205. 
"  in   the   shape    of  the   head ;    one 


Chap.  II.  Manner  of  Developmefit  29 

completely  siibjugcated  by  other  men,  as  that  certain  individuals 
should  be  preserved,  and  thus  unconsciously  selected,  from  some- 
how excelling  in  utility  to  their  masters.  Nor  have  certain 
male  and  female  individuals  been  intentionally  picked  out  and 
matched,  except  in  the  well-known  case  of  the  Prussian  grena- 
diers ;  and  in  this  case  man  obeyed,  as  miglit  have  been  ex- 
pected, the  law  of  methodical  selection ;  for  it  is  asserted  that 
many  tall  men  were  reared  in  the  villages  inhabited  by  the 
grenadiers  and  their  tall  wives.  In  Sparta,  also,  a  form  of  selec- 
tion was  followed,  for  it  was  enacted  that  all  children  should  be 
examined  shortly  after  birth ;  the  well-formed  and  vigorous 
being  preserved,  the  others  left  to  perish.^' 

If  we  consider  all  the  races  of  man  as  forming  a  single  species, 
his  range  is  enormous ;  but  some  separate  races,  as  the  Americans 
and  Polynesians,  have  very  wide  ranges.  It  is  a  well-known 
law  that  widely-ranging  species  are  much  more  variable  than 
species  with  restricted  ranges  ;  and  the  variability  of  man  may 
with  more  truth  be  compared  with  that  of  widely-ranging  species, 
than  with  that  of  domesticated  animals. 

Not  only  does  variability  appear  to  be  induced  in  man  and 
the  lower  animals  by  the  same  general  causes,  but  in  both  the 
same  parts  of  the  body  are  affected  in  a  closely  analogous 
manner.    This,  has  been  proved  in  such  full  detail  by  Godron  and 

^'  Mitford's  'History  of  Greece,'  vigour  of  their  children.     The  Gre- 

vol.  i.  p.  282.     It  appears  also  from  cian  poet,  Theognis,  who  lived  550 

a  passage  in  Xenophon's  '  Memora-  B.C.,    clearly    saw    how    important 

bilia,'  B.  ii.  4  (to  which  my  atten-  selection,  if  carefully  applied,  would 

tion  has    been    called    by  the    Rev.  be  for  the  improvement  of  mankind. 

J.  N.  Hoare),    that    it  was    a   well  He  saw,  likewise,  that  wealth  often 

recognised  principle  with  the  Greeks,  checks    the  proper  action  of  sexual 

that  men  ought  to  select  their  wives  selection.     He  thus  writes  : 
with     a    view    to     the    health    and 

"  With  kine  and  horses,  Kurnus  !  we  proceed 
By  reasonable  rules,  and  choose  a  breed 
For  profit  and  increase,  at  any  price ; 
Of  a  sound  stock,  without  detect  or  vice. 
But,  in  the  daily  matches  that  we  make, 
The  price  is  everything :  for  money's  sake, 
Men  marry  :  women  are  in  marriage  given  ; 
The  churl  or  ruffian,  that  in  wealth  has  thriven, 
May  match  his  oflspring  with  the  proudest  race  : 
Thus  everytliing  is  mix'd,  noble  and  base  ! 
If  then  in  outward  manner,  form,  and  mind, 
You  find  us  a  degraded,  motley  kind, 
Wonder  no  more,  my  friend  !  the  cause  is  plain, 
And  to  lament  the  consequence  is  vain." 
(The  Works  of  J.  Hookham  Frere,  vol.  ii.  1872,  p.  334.) 


30  TJie  Descent  of  Mmi.  Part  I, 

Quatrefages,  that  I  need  here  only  refer  to  their  works.^^  Mon- 
strosities, which  graduate  into  slight  variations,  are  likewise  so 
similar  in  man  and  the  lower  animals,  that  the  same  classification 
and  the  same  terms  can  be  used  for  both,  as  has  been  shewn  by 
Isidore  Geoffroy  St.-Hilaire.'^  In  my  work  on  the  yariation  of 
domestic  animals,  I  have  attemjDted  to  arrange  in  a  rude  fashion 
the  laws  of  variation  under  the  following  heads  : — The  direct  and 
definite  action  of  changed  conditions,  as  exhibited  by  all  or  nearly 
all  the  individuals  of  the  same  species,  varying  in  the  same  manner 
under  the  same  circumstances.  The  effects  of  the  long-continued 
use  or  disuse  of  parts.  The  cohesion  of  homologous  parts.  The 
variability  of  multiple  parts.  Compenpation  of  growth ;  but  of 
this  law  I  have  found  no  good  instance  in  the  case  of  man.  The 
effects  of  the  mechanical  pressure  of  one  part  on  another ;  as  of 
the  pelvis  on  the  cranium  of  the  infant  in  the  womb.  Arrests  of 
development,  leading  to  the  diminution  or  suppression  of  parts. 
The  reappearance  of  long-lost  characters  through  reversion. 
And  lastly,  correlated  variation.  All  these  so-called  laws  apply 
equally  to  man  and  the  lower  animals ;  and  most  of  them  even 
to  plants.  It  would  be  superfluous  here  to  discuss  all  of  them  ;^^ 
but  several  are  so  important,  that  they  must  be  treated  at  con- 
siderable length. 

The  direct  and  definite  action  of  changed  conditions. — This  is  a 
most  perplexing  subject.  It  cannot  be  denied  that  changed  con- 
ditions produce  some,  and  occasionally  a  considerable  effect,  on 
organisms  of  all  kinds ;  and  it  seems  at  first  probable  that  if 
sufficient  time  were  allowed  this  would  be  the  invariable  result. 
But  1  have  failed  to  obtain  clear  evidence  in  favour  of  this  con- 
clusion ;  and  valid  reasons  may  be  urged  on  the  other  side,  at 
least  as  far  as  the  innumerable  structures  are  concerned,  which 
are  adapted  for  special  ends.  There  can,  however,  be  no  doubt 
that  changed  conditions  induce  an  almost  indefinite  amount  of 
fluctuating  variability,  by  which  the  whole  organisation  is  rend- 
ered in  some  degree  plastic. 

In  the  United  States,  above  1,000,000  soldiers,  who  served  in 
the  late  war,  were  measured,  and  the  States  in  which  they  were 

^'*  Godron,    'De    I'Espece,'    1859,  '^  I    have    fully    discussed    these 

torn.  ii.  livre  3.     Quatrefages, '  Unite  laws  in  my  *  Variation  of  Animals 

de    I'Espece  Humaine,'  1861.     Also  and    Plants    under    Domestication,' 

Lectures  on  Anthropology,  given  in  vol.  ii.  chap.  xxii.  and  xxiii.     M.  J. 

the  '  Revue  des  Cours  Scientifiques,'  P.  Durand    has  lately   (1868)  pub- 

1886-1868.  lished    a    valuable    essay    '  De    I'ln- 

'*  '  Hist.  Gen.  et  Part,  des  Ano-  fluence  des  Milieux,'  &c.     He   lays 

malies   de   I'Organisation,'  in  three  much  stress,  in  the  case  of  plants,  on 

volumes,  tom.  i.  1832.  the  nature  of  the  soil. 


Chap.it.  Manner  of  Dcvelopiuciit.  31 

born  and  reared  were  recorded.'^  From  this  astonishing  number 
of  observations  it  is  proved  tlmt  local  influences  of  some  kind 
act  directly  on  stature ;  and  we  further  learn  that  "  the  State 
"  where  the  physical  growth  has  in  great  measure  taken  place, 
"  and  the  State  of  birth,  which  indicates  the  ancestry,  seem  to 
"  exert  a  marked  influence  on  the  stature."  For  instance,  it  is 
established,  "  that  residence  in  the  Western  States,  during  the 
"  years  of  growth,  tends  to  produce  increase  of  stature."  On  the 
other  hand,  it  is  certain  that  with  sailors,  their  life  delays  growth, 
as  shewn  ''  by  the  great  difference  between  the  statures  of  soldiers 
"  and  sailors  at  the  ages  of  seventeen  and  eighteen  years."  Mr.  B. 
A.  Gould  endeavoured  to  ascertain  the  nature  of  the  influences 
which  thus  act  on  stature ;  but  he  arrived  only  at  negative  results, 
namely,  that  they  did  not  relate  to  climate,  the  elevation  of  the 
land,  soil,  nor  even  "  in  any  controlling  degree  "  to  the  abundance 
or  the  need  of  the  comforts  of  life.  This  latter  conclusion  is 
directly  opposed  to  that  arrived  at  by  Villerme,  from  the  statistics 
of  the  height  of  the  conscripts  in  different  parts  of  France.  When 
we  compare  the  differences  in  stature  between  the  Polynesian 
chiefs  and  the  lower  orders  within  the  same  islands,  or  between 
the  inhabitants  of  the  fertile  volcanic  and  low  barren  coral  islands 
of  the  same  ocean,^^  or  again  between  the  Fuegians  on  the  eastern 
and  western  shores  of  their  country,  where  the  means  of  subsis- 
tence are  very  different,  it  is  scarcely  possible  to  avoid  the  con- 
clusion that  better  food  and  greater  comfort  do  influence  stature. 
But  the  preceding  statements  shew  how  difficult  it  is  to  arrive 
at  any  precise  result.  Dr.  Beddoe  has  lately  proved  that,  with 
the  inhabitants  of  Britain,  residence  in  towns  and  certain  occupa- 
tions have  a  deteriorating  influence  on  height ;  and  he  infers  that 
the  result  is  to  a  certain  extent  inherited,  as  is  likewise  the  case 
in  the  United  States.  Dr.  Beddoe  further  believes  that  wherever 
a  "  race  attains  its  maximum  of  physical  development,  it  rises 
"  highest  in  energy  and  moral  vigour."  ^^ 

Whether  external  conditions  produce  any  other  direct  effect 
on  man  is  not  known.  It  might  have  been  expected  that  dif-^ 
ferences  of  climate  would  have  had  a  marked  influence,  in  as  much 
as  the  lungs  and  kidneys  are  brought  into  activity  under  a  low 

"  'Investigations  in  Military  and  289.     There    is    also  a   remarkable 

Anthrop.  Statistics,'  &c.    1869,    by  ditlerence    in    appearance    between 

B.  A.  Gould,  p.  93,  107,  126,  131,  the  closely-allied  Hindoos  inhabiting 

134.  the   Upper  Ganges  and  Bengal ;  see 

'*  For  the  Polynesians,  see  Prich-  Elphinstone's  '  History  of  India,'  vol. 

ard's  '  Physical  Hist,  of  Mankind,'  i.  p.  324. 

vol.   v.    1847,    p.    145,    283.     Also  '^  '  Memoirs,    Anthropolog.    Soc. 

Godron,    '  De    I'Espfece,'   torn.  ii.  p.  vol.  iii.  1867-69,  pp.  561,  565,  567. 


32  The  DesceJtt  of  Man.  Part  I. 

temperature,  and  the  liver  and  skin  under  a  high  one.^  It  was 
formerly  thought  that  the  colour  of  the  skin  and  the  character 
of  the  hair  were  determined  by  light  or  heat ;  and  although  it 
can  hardly  be  denied  that  some  effect  is  thus  produced,  almost 
all  observers  now  agree  that  the  effect  has  been  very  small,  even 
after  exposure  during  many  ages.  But  this  subject  will  be  more 
properly  discussed  when  we  treat  of  the  different  races  of  man- 
kind. With  our  domestic  animals  there  are  grounds  for 
believing  that  cold  and  damp  directly  affect  the  growth  of  the 
hair ;  but  I  have  not  met  with  any  evidence  on  this  head  in  the 
case  of  man. 

Effects  of  the  iucnased  Use  and  Disuse  of  Paris. — It  is  well 
known  that  use  strengthens  the  muscles  in  the  individual,  and 
complete  disuse,  or  the  destruction  of  the  proper  nerve,  weakens 
them.  When  the  eye  is  destroyed,  the  optic  nerve  often  becomes 
atrophied.  When  an  artery  is  tied,  the  lateral  channels  increase 
not  only  in  diameter,  but  in  the  thickness  and  strength  of  their 
coats.  When  one  kidney  ceases  to  act  from  disease,  the  other 
increases  in  size,  and  does  double  work.  Bones  increase  not 
only  in  thickness,  but  in  length,  from  carrying  a  greater  weight.^^ 
Different  occupations,  habitually  followed,  lead  to  changed 
proportions  in  various  parts  of  the  body.  Thus  it  was  ascertained 
by  the  United  States  Commission  ^^  that  the  legs  of  the  sailors 
employed  in  the  late  war  were  longer  by  0'217  of  an  inch  than 
those  of  the  soldiers,  though  the  sailors  were  on  an  average 
shorter  men ;  whilst  their  arms  were  shorter  by  I  09  of  an  inch, 
and  therefore,  out  of  proportion,  shorter  in  relation  to  their 
lesser  height.  This  shortness  of  the  arms  is  apparently  due  to 
their  greater  use,  and  is  an  unexi^ected  result :  but  sailors 
chiefly  use  their  arms  in  pulling,  and  not  in  supporting  weights. 
With  sailors,  the  girth  of  the  neck  and  the  depth  of  the  instep 
are  greater,  whilst  the  circumference  of  the  chest,  waist,  and 
hips  is  less,  than  in  soldiers. 

Whether  the  several  foregoing  modifications  would  become 
hereditary,  if  the  same  habits  of  life  were  followed  during  many 
generations,  is  not  known,  but  it  is  probable.  Eengger^^  attri- 
butes the  thin  legs  and  thick  arms  of  the  Payaguas  Indians  to 

20  Dr.    Brakenridge,    '  Theory    of  Dr.    Jaeger,    "  Ueber    das    Langen- 

Diathesis,' '  Medical  Times,' June  19  wachsthum    der    Knochen,"  '  Jena- 

and  July  17,  1869.  ischen  Zeitschrift,'  B.  v.  Heft  i. 

2*  I   have    given    authorities    f(ir  "  '  Investigations,'  &c.     By  B.  A. 

these     several    statements    in    my  Gould,  1869,  p.  288. 

*  Variation    of  Animals    under   Do-  ^3    t  Saugethiere    von    Paraguay, 

mestication,'  vol.    ii.    pp.  297-300.  1830,  s.  4. 


Chap.  II.  Manner  of  Development.  33 


successive  generations  liaring  passed  nearly  their  whole  lives  in 
canoes,  with  their  lower  extremities  motionless.  Other  writers 
have  come  to  a  similar  conclusion  in  analogous  cases.  According 
to  Cranz/-^  who  lived  for  a  long  time  with  the  Esquimaux,  "  the 
"  natives  believe  that  ingenuity  and  dexterity  in  seal-catching 
"  (their  highest  art  and  virtue)  is  hereditary ;  there  is  really 
"  somethiag  in  it,  for  the  son  of  a  celebrated  seal-catcher  will 
"  distinguish  himself,  though  he  lost  his  father  in  childhood." 
But  in  this  case  it  is  mental  aptitude,  quite  as  much  as  bodily 
structure,  which  appears  to  be  inherited.  It  is  asserted  that 
the  hands  of  English  labourers  are  at  birth  larger  than  those  ot 
the  gentry.-^  From  the  correlation  which  exists,  at  least  in 
some  cases,-^  between  the  development  of  the  extremities  and  of 
the  jaws,  it  is  possible  that  in  those  classes  which  do  not  labour 
much  with  their  hands  and  feet,  the  jaws  would  be  reduced  in 
size  from  this  cause.  That  they  are  generally  smaller  in  refined 
and  civilised  men  than  in  hard-working  men  or  savages,  is  certain. 
But  with  savages,  as  Mr.  Herbert  Spencer  '""^  has  remarked,  the 
greater  use  of  the  jaws  in  chewing  coarse,  uncooked  food,  would 
act  in  a  direct  manner  on  the  masticatory  muscles,  and  on  the 
bones  to  which  they  are  attached.  In  infants,  long  before  birth, 
the  skin  on  the  soles  of  the  feet  is  thicker  than  on  any  other  part 
of  the  body;-^  and  it  can  hardly  be  doubted  that  this  is  due 
to  the  inherited  effects  of  pressure  during  a  long  series  of 
generations. 

It  is  familiar  to  every  one  that  watchmakers  and  engravei-s 
are  liable  to  be  short-sighted,  whilst  men  living  much  out  of 
doors,  and  especially  savages,  are  generally  long-sighted.^^  Short- 
sight  and  long-sight  certainly  tend  to  be  inherited.^''  The 
inferiority  of  Europeans,  in  comparison  w^ith  savages,  in  eye- 
sight and  in  the  other  senses,  is  no  doubt  the  accumulated  and 
transmitted  effect  of  lessened  use  during  many  generations ;  for 
Eengger^^   states  that  he  has  repeatedly  observed  Europeans, 

^*  'History    of  Greenland,'    Eng.  ('Sanitary  Memoirs   of  the  War  of 

translat.  17G7,  vol.  i.  p.  230.  the   Rebellion,'  1869,   p.   530),    has 

2^  '  Intermarriage.'        By      Alex,  proved  this  to  be  the  case ;  and  h«' 

Walker,  1838,  p.  377.  accounts    for    it    by    the    ordinal  v 

^°  'The    Variation     of     Animals  range  of  vision  in  sailors  being  "  n-- 

under  Domestication,'  vol.  i,  p.  173.  "  stricted  to  the  length  of  the  vcsm;! 

27  I.  Principles  of  Biology,'  vol.  i.  "  and  the  height  of  tlie  masts." 

p.  455.  ^°  'The     Variation     of     Animals 

28  Paget,   '  Lectures    on    Surgical      under  Domestication,'  vol.  i.  p.  8. 
Pathology,'  vol.  ii.  1853,  p.  209.  ^'  '  Siingethiere    von    Paraguay,' 

2^  It    is    a    singular    and     unex-  s.  8,  10.     I  have  had  good  opportuui- 

pected  fact  that  sailors  are  interior  ties  for  observing  the  extraordinarv 

to  landsmen  in  their  mean  distance  power  of  eyesight  in  the  Fuegians. 

of  distinct  vision.     Dr.  B.  A.  Gould  See    also    Lawrence    ('  lectures    on 


34  T^he  Descent  of  Alan.  Part  I, 

who  had  been  brought  up  and  spent  their  whole  lives  with  the 
wild  Indians,  who  nevertheless  did  not  equal  them  in  the  sharp- 
ness of  their  senses.  The  same  naturahst  observes  that  the 
cavities  in  the  skull  for  the  reception  of  the  several  sense-organs 
are  larger  in  the  American  aborigines  than  in  Europeans ;  and 
this  probably  indicates  a  corresponding  difference  in  the  dimen- 
sions of  the  organs  themselves.  Blumenbach  has  also  remarked 
on  the  large  size  of  the  nasal  cavities  in  the  skulls  of  the 
American  aborigines,  and  connects  this  fact  with  their  remarkably 
acute  power  of  smell.  The  Mongolians  of  the  plains  of  Northern 
Asia,  according  to  Pallas,  have  wonderfully  perfect  senses ;  and 
Prichard  believes  that  the  great  breadth  of  their  skulls  across 
the  zygomas  follows  from  their  high]y-develox)ed  sense-organs.^^ 
The  Quechua  Indians  inhabit  the  lofty  plateaux  of  Peru ;  and 
Alcide  d'Orbigny  states  ^^  that,  from  continually  breathing  a 
highly  rarefied  atmosphere,  they  have  acquired  chests  and  lungs 
of  extraordinary  dimensions.  The  cells,  also,  of  the  lungs  are 
larger  and  more  numerous  than  in  Europeans.  These  observa- 
tions have  been  doubted ;  but  Mr.  D.  Forbes  carefully  measured 
many  Aymaras,  an  allied  race,  living  at  the  height  of  between 
10,000  and  15,000  feet;  and  he  informs  me 2*  that  they 
differ  conspicuously  from  the  men  of  all  other  races  seen  by  him 
in  the  circumference  and  length  of  their  bodies.  In  his  table  of 
measurements,  the  stature  of  each  man  is  taken  at  1000,  and  the 
other  measurements  are  reduced  to  this  standard.  It  is  here 
seen  that  the  extended  arms  of  the  Aymaras  are  shorter  than 
those  of  Europeans,  and  much  shorter  than  those  of  Negroes. 
The  legs  are  likewise  shorter  ;  and  they  present  this  remarkable 
peculiarity,  that  in  every  Aymara  measured,  the  femur  is  actually 
shorter  than  the  tibia.  On  an  average,  the  length  of  the  femur 
to  that  of  the  tibia  is  as  211  to  252 ;  whilst  in  two  Europeans, 
measured  at  the  same  time,  the  femora  to  the  tibiae  were  as  2'44 
to  230;  and  in  three  Negroes  as  258  to  211.  The  humerus  is 
likewise  shorter  relatively  to  the  forearm.  This  shortening  of 
that  part  of  the  limb  which  is  nearest  to  the  body,  appears  to  be, 
as  suggested  to  me  by  Mr.  Forbes,  a  case  of  compensation  in 


Physiology,'   &c.,   1822,  p.  404)  on  bach,  vol.  i.  1851,  p.  311;  for  the 

this  same  subject.  M.  Giraud-Teulon  statement  by  PaHas,  a-o1.  iv.   1844, 

has   recently  collected  ('  Revue  des  p.  407. 

Cours  Scieutifiques,'  1870,  p.  625)  ^^  Quoted  by  Prichard,  '  Re- 
a  large  and  valuable  body  of  evidence  searches  into  the  Phys.  Hist,  of  Man- 
proving  that  the  cause  of  short-  kind,'  vol.  v.  p.  463. 
sight,  "  Cest  le  travail  assidu,  de  34  ]\jj._  Forbes'  valuable  paper  is 
pres."  now  published  in  the  '  Journal  of 
'2  Prichard,  '  Phys.  Hist,  of  Man-  the  Ethnological  Sec.  of  London,* 
kind,'  on  the  authoritv  of  F>lumen-  new  series,  vnl.  ii.  1870,  p.  193. 


Chap.  II.  Manner  of  Devdopnient.  35 


relation  with  the  greatly  increased  length  of  the  trunk.  The 
Aymaras  present  some  other  singular  points  of  structure,  for 
instance,  the  very  small  projection  of  the  heel. 

These  men  are  so  thoroughly  acclimatised  to  their  cold  and 
lofty  abode,  that  when  formerly  carried  down  by  the  Spaniards 
to  the  low  eastern  plains,  and  when  now  tempted  down  by  high 
wages  to  the  gold-washings,  they  suifer  a  frightful  rate  of  mor- 
tality. Nevertheless  Mr.  Forbes  found  a  few  pure  families 
which  had  survived  during  two  generations :  and  he  observed  that 
they  still  inherited  their  characteristic  peculiarities.  But  it  was 
manifest,  even  without  measurement,  that  these  pecaharities 
had  all  decreased  ;  and  on  measurement,  their  bodies  were  found 
not  to  be  so  much  elongated  as  those  of  the  men  on  the  high 
plateau  ;  wliilst  their  femora  had  become  somewhat  lengthened, 
as  had  their  tibite,  although  in  a  less  degree.  The  actual 
measurements  may  be  seen  by  consulting  Mr.  Forbes's  memoir. 
From  these  observations,  there  can,  I  think,  be  no  doubt  that 
residence  during  many  generations  at  a  great  elevation  tends, 
both  directly  and  indirectly,  to  induce  inherited  modifications 
in  the  proportions  of  the  body.^^ 

Although  man  may  not  have  been  much  moiified  during 
the  latter- stages  of  his  existence  through  the  increased  or  de- 
creased use  of  parts,  the  facts  now  given  shew  that  his  liability  in* 
this  respect  has  not  been  lost ;  and  we  positively  know  that  the 
same  law  holds  good  with  the  lower  animals.  Consequently  we 
may  infer  that  when  at  a  remote  epoch  the  progenitors  of  man 
were  in  a  transitional  state,  and  were  changing  from  quadrupeds 
into  bipeds,  natural  selection  would  probably  have  been  greatly 
aided  by  the  inherited  effects  of  the  increased  or  diminished  use 
of  the  different  parts  of  the  body. 

A  rrests  of  Developmen  f. — There  is  a  difference  between  arrested 
development  and  arrested  growth,  for  parts  in  the  former  state 
continue  to  grow  whilst  still  retaining  their  early  condition. 
Various  riionstrosities  come  under  this  head ;  and  some,  as  a 
cleft-palate,  are  known  to  be  occasionally  inherited.  It  will 
suffice  for  our  purpose  to  refer  to  the  arrested  brain-development 
of  microcephalous  idiots,  as  described  in  Yogt's  memoir.''^ 
Their  skulls  are  smaller,  and  the  convolutions  of  the  brain 
are  less  complex  than  in  normal  men.     The  frontal  sinus,  or  the 

'^  Dr.  Wilckens  ('  Landwirth-  regions,  have  their  frames  modified, 
schaft.  Wochenblatt,'  No.  10,  1869)  ^<*  '  Me'moire     sur     les     Microc6- 

has  lately  published    an    interesting  phales,'  1867,  jip.  50,  125,  169,  171, 

Essav  shewing    how    domestic    ani-  18-1-198. 
mals,    which    live    in     mountainous 


36  The  Descent  of  Man.  Part  I. 

projection  over  the  eye-brows,  is  largely  developed,  and  the  jaws 
are  prognathous  to  an  "  effrayant'^  degree ;  so  that  these  idiots 
somewhat  resemble  the  lower  types  of  mankind.  Their  in- 
telligence, and  most  of  their  mental  faculties,  are  extremely 
feeble.  They  cannot  acquire  the  power  of  speech,  and  are 
wholly  incapable  of  prolonged  attention,  but  are  much  given  to 
imitation.  They  are  strong  and  remarkably  active,  continually 
gambohng  and  jumping  about,  and  making  grimaces.  They 
often  ascend  stairs  on  all-fours;  and  are  curiously  fond  of 
climbing  up  furniture  or  trees.  We  are  thus  reminded  of  the 
delight  shewn  by  almost  all  boys  in  climbing  trees;  and  this 
again  reminds  us  how  lambs  and  kids,  originally  alpine  animals, 
delight  to  frisk  on  any  hillock,  however  small.  Idiots  also 
resemble  the  lower  animals  in  some  other  respects ;  thus  several 
cases  are  recorded  of  their  carefully  smelling  every  mouthful  of 
food  before  eating  it.  One  idiot  is  described  as  often  using  his 
mouth  in  aid  of  his  hands,  whilst  hunting  for  lice.  They  are 
often  filthy  in  their  habits,  and  have  no  sense  of  decency ;  and 
several  cases  have  been  published  of  their  bodies  being  re- 
markably hairy  ."'^ 

Reversion. — Many  of  the  cases  to  be  here  given,  might  have 
been  introduced  under  the  last  heading.  When  a  structure 
is  arrested  in  its  development,  but  still  continues  growing, 
until  it  closely  resembles  a  corresponding  structure  in  some 
lower  and  adult  member  of  the  same  group,  it  may  in  one  sense 
be  considered  as  a  case  of  reversion.  The  lower  members  in  a 
group  give  us  some  idea  how  the  common  progenitor  was 
probably  constructed ;  and  it  is  hardly  credible  that  a  complex 
part,  arrested  at  an  early  phase  of  embryonic  development,  should 
go  on  growing  so  as  ultimately  to  perform  its  proper  function, 
unless  it  had  acquired  such  power  during  some  earher  state  of 
existence,  when  the  present  exceptional  or  arrested  structure 
was  normal.  The  simple  brain  of  a  microcephalous  idiot,  in  as 
far  as  it  resembles  that  of  an  ape,  may  in  this  sense  be  said  to 
offer  a  case  of  reversion.^^    There  are  other  cases  which  come 

3^  Prof.    Laycock    sums    up    the  pp.  46-51.     Pinel  has  also  given  a 

character    of    brute-like    idiots    by  striking    case    of    hairiness    in    an 

calling   them    theroid ;  'Journal    of  idiot. 

Mental    Science,'    July    1 863.      Dr.  ^*  In  my  '  Variation  of  Animals 

Scott  ('The  Deaf  and  Dumb,'  2nd  under  Domestication  '  (vol.  ii.  p.  57), 

edit.,    1870,   p.    10)   has    often    ob-  I  attributed  the  not  very  rare  cases 

served  the  imbecile  smelling  their  of  supernumerary  mammae  in  women 

food.     See,    on    this    same    subject,  to  reversion.     I  was  led  to  this  as  a 

and   on   the   hairmess  of  idiots,  Dr.  probable  conclusion,  by  the  additional 

Maudsley,  '  Body  and  Mind,'   1870,  mammae     being     generally     placed 


Chap.  II. 


Manner  of  Development, 


Z7 


more  strictly  under  oiir  jiresent  head  of  reversion.     Certain 
structures,  regularly  occurring  in  the  lower  members  of  the  group 


symmetrical!}'  on  the  breast ;  and 
more  especially  from  one  case,  in 
whicii  a  single  elHcient  mamma 
occurred  in  the  inguinal  region  of 
a  woman,  the  daughter  of  another 
woman  with  supernumerary  mam- 
mas. But  I  now  find  (see,  for  in- 
stance, Prof.  Preyer,  '  Der  Kampf 
um  das  Dasein,'  1869,  s.  45)  that 
mamma:  crraticcc  occur  in  other 
situations,  as  on  the  back,  in  the 
armpit,  and  on  the  thigh ;  the 
mammaj  in  this  latter  instance 
having  given  so  much  milk  that  the 
child  was  thus  nourished.  The  pro- 
bability that  the  additional  mammaj 
are  due  to  reversion  is  thus  much 
weakened ;  nevertheless,  it  still 
seems  to  me  probable,  because  two 
pairs  are  often  found  symmetrically 
on  the  breast ;  and  of  this  I  myself 
have  received  information  in  several 
cases.  It  is  well  known  that  some 
Lemurs  normally  have  two  pairs  of 
mammje  on  the  breast.  Five  cases 
have  been  recorded  of  the  presence 
of  more  than  a  pair  of  mammte  (of 
course  rudimentary)  in  the  male 
sex  of  mankind  ;  see  '  Journal  of 
Anat.  and  Physiology,'  1872,  p.  56, 
for  a  case  given  by  Dr.  Handyside, 
in  which  two  brothers  exhibited 
this  peculiarity ;  see  also  a  paper  by 
Dr.  Bartels,  in  Pieichert's  and  du 
Bois  Reymond's  Archiv.,  1872,  p. 
304.  In  one  of  the  cases  alluded  to 
by  Dr.  Bartels,  a  man  bore  five 
mammas,  one  being  medial  and 
placed  above  the  navel ;  Meckel 
von  Hemsbach  thinks  that  this 
latter  case  is  illustrated  by  a 
medial  mamma  occurring  in  certain 
Cheiroptera.  On  the  whole,  we  may 
well  doubt  if  additional  mamma) 
would  ever  have  been  developed  in 
both  sexes  of  mankind,  had  not  his 
early  progenitors  been  provided  with 
more  than  a  single  pair. 

In  the  above  work  (vol.  ii.  p,  12), 
I  also  attributed,  though  with  much 
hesitation,    the    frequent    cases    of 


polydactylism  in  men  to  reversion. 
I  was  partly  led  to  this  through 
Prof.  Owen's  statement,  that  some 
of  the  Ichthyopterygia  possess  more 
than  five  digits,  and  therefore,  as  I 
supposed,  had  retained  a  primordial 
condition ;  but  Prof.  Gegenbaur 
(^  Jenaischen  Zeitschrift,'  B.  v.  Heft 
o,  s.  341),  disputes  Owen's  conclu- 
sion. On  the  other  hand,  according 
to  the  opinion  lately  advanced  by 
Dx'.  Gunther  on  the  paddle  of  Ceia- 
todus,  which  is  provided  with  ar- 
ticulated bony  rays  on  both  sides  of 
a  central  chain  of  bones,  there 
seems  no  great  difficulty  in  admit- 
ting that  six  or  more  digits  on  one 
side,  or  double  the  number  on  both 
sides,  might  reappear  through  re- 
version. I  am  informed  by  Dr. 
Zouteveen  that  there  is  a  case  on 
record  of  a  man  having  twenty-four 
fingers  and  twenty-four  toes !  I 
was  chiefly  led  to  the  conclusion 
that  the  presence  of  supernumerary 
digits  is  due  to  reversion  from  the 
fact  that  such  digits  not  only  are 
strongly  inherited,  but  have  the 
power  of  regrowth  after  amputa- 
tion, like  the  normal  digits  of  the 
lower  vertebrata.  This  fact  of  their 
regrowth  remains  inexplicable,  if 
the  belief  in  reversion  to  some  ex- 
tremely remote  progenitor  is  re- 
jected. Arrested  development  and 
reversion  are  intimately  connected, 
and  thus  the  belief  in  reversion  in 
the  present  case  is  to  a  certain  ex- 
tent supported  by  the  frequent,  or 
almost  constant,  coincidence  insisted 
on  by  Meckel  and  I.  Geotfroy  St.- 
Hilaire,  between  various  arrests  of 
development,  such  as  cleft-palate, 
bifid  uterus,  cyclopean  state  of  the 
eyes,  &c.,  and  the  presence  of  ad- 
ditional digits  (see,  on  this  head, 
M.  A.  Pionjou,  'Types  Primitifs  des 
Mammiferes,'  p.  Gl  ;  and  M.  Ber- 
tillon,  '  Valeur  Phil.  Hyp.  du  Trans- 
formisme ').  It  is,  on  the  other  hand, 
no  real  objection  to  the  view  here 


2,S  The  Descent  of  Man.  Pakt  1. 

to  which  man  belongs,  occasionally  make  their  appearance  in 
him,  though  not  found  in  the  normal  human  embryo;  or,  if 
normally  present  in  the  human  embryo,  they  become  abnormally 
developed,  although  in  a  manner  which  is  normal  in  the  lower 
members  of  the  group.  These  remarks  will  be  rendered  clearer 
by  the  following  illustrations. 

In  yarious  mammals  the  uterus  graduates  from  a  double 
organ,  with  two  distinct  orifices  and  two  passages,  as  in  the 
marsupials,  into  a  single  organ,  which  is  in  no  way  double, 
except  from  having  a  slight  internal  fold,  as  in  the  higher  apes 
and  man.  The  rodents  exhibit  a  perfect  series  of  gradations 
between  these  two  extreme  states.  In  all  mammals  the  uterus 
is  developed  from  two  simple  primitive  tubes,  the  inferior 
portions  of  wliich  form  the  cornua ;  and  it  is,  in  the  words  of 
I)r.  Farre,  "  by  the  coalescence  of  the  two  cornua  at  their  lower 
"  extremities  that  the  body  of  the  uterus  is  formed  in  man ; 
"  while  in  those  animals  in  which  no  middle  portion  of  body 
'•'  exists,  the  cornua  remain  un-united.  As  the  development  of 
"  the  uterus  proceeds,  the  two  cornua  become  gradually  shorter, 
"  until  at  length  they  are  lost,  or,  as  it  were,  absorbed  into  the 
"  body  of  the  uterus."  The  angles  of  the  uterus  are  still 
produced  into  cornua,  even  so  high  up  in  the  scale  as  the  lower 
apes,  and  lemurs. 

Now  in  women,  anomalous  cases  are  not  very  infrequent,  in 
which  the  mature  uterus  is  furnished  with  cornua,  or  is  partially 
divided  into  two  organs;  and  such  cases,  according  to  Owen, 
repeat  "the  grade  of  concentrative  development,"  attained  by 
certain  rodents.  Here  perhaps  we  have  an  instance  of  a  simple 
arrest  of  embryonic  development,  with  subsequent  growth  and 
perfect  functional  development ;  for  either  side  of  the  partially 
double  uterus  is  capable  of  performing  the  proper  office  of 
gestation.  In  other  and  rarer  cases,  two  distinct  uterine  cavities 
are  formed,  each  having  its  proper  orifice  and  passage.^^  No 
such  stage  is  passed  through  during  the  ordinary  development 
of  the  embryo,  and  it  is  difficult  to  beheve,  though  perhaps  not 
impossible,  that  the  two  simple,  minute,  primitive  tubes  should 
know  how  (if  such  an  expression  may  be  used)  to  grow  into  two 


maintained  that  supernumerary  di-  ultimately  prevail, 

gits  are  often  present  without  any  ^^  See   Dr.  A.  Farre's  well-known 

other  structure  being  affected  ;  for  article  in  the  '  Cyclop,  of  Anat.  and 

numerous     cases     could    be     given  Phys.'  vol.  v.  1859,  p.  642.     Owen 

of    single     characters    reappearing  '  Anatomy  of  Vertebrates,'  vol.  iii. 

through  reversion.     On  the  whole,  1868,    p.    687.      Prof.    Turner    in 

I  cannot   but  think   that  the  view  '  Edinburgh  Medical  Journal,'  Feb. 

originally   propounded    by    me  %vill  1805. 


Chap.  II.  Manner  of  Development.  39 

distinct  uteri,  each  with  a  well-constructed  orifice  and  passage, 
and  each  furnished  with  numerous  muscles,  nerves,  glands  and 
vessels,  if  they  had  not  formerly  passed  through  a  similar  course 
of  development,  as  in  the  case  of  existing  marsupials.  No  one 
will  pretend  that  so  perfect  a  structure  as  the  abnormal  double 
uterus  in  woman  could  be  the  result  of  mere  chance.  But  the 
principle  of  reversion,  by  which  a  long-lost  structure  is  called 
back  into  existence,  might  serve  as  the  guide  for  its  full  develop- 
ment, even  after  the  lapse  of  an  enormous  interval  of  time. 

Professor  Canestrini,  after  discussing  the  foregoing  and  various 
analogous  cases,  arrives  at  the  same  conclusion  as  that  just 
given.  He  adduces  another  instance,  in  the  case  of  the  malar 
bone,*"  which,  in  some  of  the  Quadrumana  and  other  mammals, 
normally  consists  of  two  portions.  This  is  its  condition  in  the 
human  foetus  when  two  months  old ;  and  through  arrested  develop- 
ment, it  sometimes  remains  thus  in  man  when  adult,  more 
especially  in  the  lower  prognathous  races.  Hence  Canestrini 
concludes  that  some  ancient  progenitor  of  man  must  have  had 
this  bone  normally  divided  into  tvro  portions,  wiiich  afterwards 
became  fused  together.  In  man  the  frontal  bone  consists  of  a 
single  jDiece,  but  in  the  embryo,  and  in  children,  and  in  almost 
all  the  lower  mammals,  it  consists  of  two  pieces  separated  by  a 
distinct  suture.  This  suture  occasionally  persists  more  or  less 
distinctly  in  man  after  maturity ;  and  more  frequently  in  ancient 
than  in  recent  crania,  especially,  as  Canestrini  has  observed,  in 
those  exhumed  from  the  Drift,  and  belonging  to  the  brachyce- 
phalic  type.  Here  again  he  comes  to  the  same  conclusion  as  in 
the  analogous  case  of  the  malar  bones.  In  this,  and  other  instances 
presently  to  be  given,  the  cause  of  ancient  races  approaching  the 
lower  animals  in  certain  characters  more  frequently  than  do  the 
modern  races,  appears  to  be,  that  the  latter  stand  at  a  somewhat 

*°  '  Annuario  della  Soc.  dei  Xatu-  tected    in    about    two    per    cent,  of 

ralisti    in    Modena,'    1867,    p.    83.  adult  skulls;  he  also  remarks  that 

Prof.   Canestrini    gives    extracts   on  it  more  frequently    occurs  in    pro- 

this  subject  from  various  authorities.  gnathous  skulls,  not  of  the  Aryan 

Laurillard  remarks,  that  as  he  has  race,  than   in   others.     See   also  G. 

found  a  complete  similarity  in  the  Delorenzi  on  the  same  subject ;  '  Tre 

form,  proportions,  and  connection  of  nuovi     cast     d' auomalia    dell'osso, 

the    two    malar   bones    in    several  malare,'   Torino,     1872.  .    Also,    E. 

human  subjects  and  in  certain  apes,  Morselli,  '  Sopra  una  rara  anomalia 

he   cannot  consider   this  disposition  dell'  osso    malare,'    Modena,    1872. 

of  the    parts  as  simply  accidental.  Still     more     recently    Gruber    has 

Another  paper  on  this  same  anomaly  written  a  pamjjlilet  on  the  division 

has  been  published  by  Dr.  Saviotti  ofthisboue.     1  give  these  references 

in    the    '  Gazzetta   delle     Cliniche,'  because    a    reviewer,    without    any 

Turin,    1871,    where    he    says    that  grounds    or    scruples,    has    thrown 

traces   of   the    divij,ion   mav  be  de-  doubts  on  mv  statements. 


40  The  Descent  of  Man.  Part  I. 

greater  distance  in  the  long  line  of  descent  from  their  early  semi- 
human  progenitors. 

Various  other  anomalies  in  man,  more  or  less  analogous  to  the 
foregoing,  have  been  advanced  by  different  authors,  as  cases  of 
reversion;  but  these  seem  not  a  little  doubtful,  for  we  have  to 
descend  extremely  low  in  the  mammalian  series,  before  we  find 
such  structures  normally  present.*^ 

In  man,  the  canine  teeth  are  perfectly  efficient  instruments  for 
mastication.  But  their  true  canine  character,  as  Owen*^  re- 
marks, "  is  indicated  by  the  conical  form  of  the  crown,  which 
**  terminates  in  an  obtuse  point,  is  convex  outward  and  flat  or 
"  sub-concave  within,  at  the  base  of  which  surface  there  is  a 
"  feeble  prominence.  The  conical  form  is  best  expressed  in  the 
"  Blelanian  races,  especially  the  Australian.  The  canine  is  more 
"  deeply  implanted,  and  by  a  stronger  fang  than  the  incisors." 
Nevertheless,  this  tooth  no  longer  serves  man  as  a  special  weapon 
for  tearing  his  enemies  or  prey ;  it  may,  therefore,  as  far  as  its 
proper  function  is  concerned,  be  considered  as  rudimentary.  In 
every  large  collection  of  human  skulls  some  may  be  found,  as 
Hackel^^  observes,  with  the  canine  teeth  projecting  considerably 
beyond  the  others  in  the  same  manner  as  in  the  anthropomorphous 
apes,  but  in  a  less  degree.  In  these  cases,  open  spaces  between 
the  teeth  in  the  one  jaw  are  left  for  the  reception  of  the  canines 
of  the  opposite  jaw.  An  interspace  of  this  kind  in  a  Kaffir 
skull,  figured  by  Wagner,  is  surprisingly  wide.^*  Considering 
how  few  are  the  ancient  skulls  which  have  been  examined, 
compared  to  recent  skulls,  it  is  an  interesting  fact  that  in  at 
least  three  cases  the  canines  project  largely;  and  in  the  Naulette 
jaw  they  are  spoken  of  as  enormous.*^ 

*'  A  whole  series  of  cases  is  given  if  in    any  way  serviceable,  for   in- 

by  Isid.  GeofFroy  St.-Hilaire,  '  Hist,  stance,  in  shortening  and  simplifying 

des   Anomalies,'    torn.    iii.    p.    437.  the    course    of  development  ?     And 

A  reviewer  ('  Journal  of  Anat.  and  again,  why  should  not  injurious  ab- 

Physiology,'  1871,  p.    366)    blames  normalities,  such  as  atrophied  or  hj-- 

me  much  for   not  having  discussed  pertrophied    parfs,  which    have    no 

the    numerous    cases,    which    have  relation  to  a  former  state  of  exist- 

been  recorded,  of  various  parts  ar-  ence,  occur  at  an   early  period,  as 

rested    in    their    development.     He  well  as  during  maturity  ? 

says  that,  according  to  my  theory,  4-  '  Anatomy  of  Vertebrates,'  vol. 

"  every   transient    condition    of  an  iii.  1868,  p.  323. 

"  organ,  during  its  development,  is  *^  '  Generelle  Morphologie,'  1866, 

"  not  only  a  means  to  an   end,  but  B.  ii.  s.  civ. 

"  once  was  an  end  in  itself."     This  *^  Carl  Vogt's  'Lectures  on  Man,' 

does  not  seem  to  me  necessarily  to  Eng.  translat.  1864,  p.  151. 

hold  good.     Why  should  not  varia-  ^^  C.    Carter    Blake,    on    a    jaw 

tions  occur  during  an  early  period  from    La    Naulette,    '  Anthropolog. 

of  development,  having  no  relation  Keview,'     1867,     p.     295.     Schaaff- 

to   -"pvprsion ;    yet   such    variations  hausen,  iliid.  1868,  p.  426. 
might  be  preserved  and  accumul.ite'l. 


Chap.  II.  Manner  of  Development.  41 

Of  the  anthropomorphous  apes  the  males  alone  have  their 
canines  fully  developed ;  but  in  the  female  gorilla,  and  in  a  less 
degree  in  the  female  orang,  these  teeth  project  considerably 
beyond  the  others;  therefore  the  fact,  of  which  I  have  been 
assured,  that  women  sometimes  have  considerably  projecting 
canines,  is  no  serious  objection  to  the  belief  that  their  occasional 
great  development  in  man  is  a  case  of  reversion  to  an  ape-hke 
progenitor.  He  who  rejects  with  scorn  the  belief  that  the  shape 
of  his  own  canines,  and  their  occasional  great  development  in 
other  men,  are  due  to  our  early  forefathers  having  been  pro- 
vided with  these  formidable  weapons,  will  probably  reveal,  by 
sneering,  the  line  of  his  descent.  For  though  he  no  longer 
intends,  nor  has  the  power,  to  use  these  teeth  as  weapons,  he  will 
unconsciously  retract  his  "  snarling  muscles"  (thus  named  by 
Sir  C.  Bell),""'  so  as  to  expose  them  ready  for  action,  like  a  dog 
prepared  to  fight. 

Many  muscles  are  occasionally  developed  in  man,  which  are 
proper  to  the  Quadrumana  or  other  mammals.  Professor 
Ylacovich*'  examined  forty  male  subjects,  and  found  a  muscle, 
called  by  him  the  iscliio-pubic,  in  nineteen  of  them ;  in  three 
others  there  was  a  ligament  which  represented  this  muscle ;  and 
in  the  remaining  eighteen  no  trace  of  it.  In  only  two  out  of 
thirty  female  subjects  was  this  muscle  developed  on  both  sides, 
but  in  three  others  the  rudimentary  ligament  was  present.  This 
muscle,  therefore,  appears  to  be  much  more  common  in  the 
male  than  in  the  female  sex ;  and  on  the  belief  in  the  descent 
of  man  from  some  lower  form,  the  fact  is  intelligible ;  for  it 
has  been  detected  in  several  of  the  lower  animals,  and  in  all 
of  these  it  serves  exclusively  to  aid  the  male  in  the  act  of 
reproduction. 

Mr.  J.  Wood,  in  his  valuable  series  of  papers,"*^  has  minutely 
described  a  vast  number  of  muscular  variations  in  man,  which 
resemble  normal  striPctures  in  the  lower  animals.     The  muscles 

<«  'The  Anatomy  of  Expression,'  pp.  241,  24-2;  vol.  xv.  1867,  p.  544; 

1844,  pp.  110,  131.  vol.  xvi.  1868,  p.  524.     I  may  here 

*'   Quoted  by  Prof.  Canestrini  in  add    that    Dr.    Murie    and    Mr.   St. 

the  'Annuario,'  &c.,  1867,  p.  90.  George  Mivart  have  shewn  in  their 

^^  These    papers    deserve    careful  Memoir  on  the  Lemuroidea  (•  Tran- 

study  by   any    one   who    desires    to  sact.    Zoolog.   Soc'   vol.    vii.    1869, 

learn    how  frequently  our   muscles  p.  '^%\  how  extraordinarily  variable 

vary,  and    in  varying    come  to   re-  some   of  the  muscles   are    in  these 

semble    those    of   the  Quadrumana.  animals,  the  lowest  members  of  the 

The  following  references    relate    to  Primates.     Gi-adations,  also,  in  the 

the  few  points    touched    on   in   my  muscles  leading  to  structures  found 

text:    '  Proc.    Royal    Soc.    vol.   xiv.  in  animals  still  lower  in  the  scale, 

1865,  pp.  379-384;    vol.  xv.    186G,  are  numerous  in  tin?  Lemuroidea. 


42  The  Descent  of  Man.  Part  I. 

■which  closely  resemble  those  regularly  present  in  our  nearest 
allies,  the  Quadrumana,  are  too  numerous  to  be  here,  even 
specified.  In  a  single  male  subject,  having  a  strong  bodily 
frame,  and  well-formed  skull,  no  less  than  seven  muscular  varia- 
tions were  observed,  all  of  which  plainly  represented  muscles 
proper  to  various  kinds  of  apes.  This  man,  for  instance,  had  on 
both  sides  of  his  neck  a  true  and  powerful  "  levator  claviculce,'' 
such  as  is  found  in  all  kinds  of  apes,  and  which  is  said  to  occur 
in  about  one  out  of  sixty  human  subjects."^  Again,  this  man 
had  "  a  special  abductor  of  the  metatarsal  bone  of  the  fifth 
"  digit,  such  as  Professor  Huxley  and  Mr.  Flower  have  shewn 
"  to  exist  uniformly  in  the  higher  and  lower  apes."  I  will  give 
only  two  additional  cases;  the  acromio-basilar  muscle  is  found 
in  all  mammals  below  man,  and  seems  to  be  correlated  with  a 
quadrupedal  gait,^°  and  it  occurs  in  about  one  out  0/  sixty 
human  subjects.  In  the  lower  extremities  Mr.  Bradley  ^^  found 
an  abductor  ossis  metatarsi  quinti  in  both  feet  of  man ;  this  muscle 
had  not  up  to  that  time  been  recorded  in  mankind,  but  is 
always  present  in  the  anthropomorphous  apes.  The  muscles  of 
the  hands  and  arms— parts  which  are  so  eminently  characteristic 
of  man— are  extremely  liable  to  vary,  so  as  to  resemble  the 
corresponding  muscles  in  the  lower  animals.^^  Such  resem- 
blances are  either  perfect  or  imperfect;  yet  in  the  latter  case 
they  are  manifestly  of  a  transitional  nature.  Certain  variations 
are  more  common  in  man,  and  others  in  woman,  without  our 
being  able  to  assign  any  reason.  Mr.  Wood,  after  describing 
numerous  variations,  makes  the  following  pregnant  remark: 
"  Notable  departures  from  the  ordinary  type  of  the  muscular 
"  structures  run  in  grooves  or  directions,  which  must  be  taken 
"  to  indicate  some  unknown  factor,  of  much  importance  to  a 
"  comprehensive  knowledge  of  general  and  scientific  anatomy."*^ 

*^  See    also    Prof.   Macalister  in  able  case  of  variation  in  the  human 

'  Proc.    R.    Irish  Academ}-,'  vol.   x.  flexor  poUicis   longus,   adds,    "  This 

1868,  p.  124.  "  remarkable    example    shews    that 

^"  Mr.  Champneys  in  '  Journal  of  "  man  may  sometimes    possess   the 

Anat.  and  Phys.'  Nov.,  1871,  p.  178.  "  arrangement  of  tendons  of  thumb 

51   'Journal  of  Anat.  and  Phys.'  "and  fingers   characteristic  of   the 

May,  1872,  p.  421.  "macaque;    but    whether    such    a 

*2  Prof,  Macalister  (ibid.  p.  121)  "  case  should  be  regarded  as  a 
has  tabulated  his  observations,  and  "  macaque  passing  upwards  into  a 
finds  that  muscular  abnormalities  "  man,  or  a  man  passing  downwards 
are  most  frequent  in  the  fore-arms,  "  into  a  macaque,  or  as  a  congenital 
secondly,  in  the  face,  thirdly,  in  the  "  freak  of  nature,  I  cannot  under- 
foot  &c.  "  take  to  say."     It    is    satisfactory 

*^  The  Rev.  Dr.  Haughton,  after  to   hear   so    capable    an    anatomist, 

giving    ('  Proc.    R.  Irish  Academy,'  and  so    embittered   an    opponent  of 

June°27,  1864,  p.  71o)  a  remark-  evolutionism,    admitting    even  the 


Chap.  II.  Manner  of  Development.  43 

That  this  unknown  factor  is  reTcrsion  to  a  former  state  of 
existence  may  be  admitted  as  in  the  highest  degree  probable.'^* 
It  is  quite  incredible  that  a  man  should  through  mere  accident 
abnormally  resemble  certain  apes  in  no  less  than  seven  of  his 
muscles,  if  there  had  been  no  genetic  connection  between  them. 
On  the  other  hand,  if  man  is  descended  from  some  ape-like 
creature,  no  valid  reason  can  be  assigned  why  certain  muscles 
should  not  suddenly  reappear  after  an  interval  of  many  thou- 
sand generations,  in  the  same  manner  as  with  horses,  asses,  and 
mules,  dark-coloured  stripes  suddenly  reappear  on  the  legs, 
and  shoulders,  after  an  interval  of  hundreds,  or  more  probably 
of  thousands  of  generations. 

These  various  cases  of  reversion  are  so  closely  related  to  those 
of  rudmientary  organs  given  in  the  first  chapter,  that  many  of 
them  might  have  been  indifferently  introduced  either  there  or 
here.  Thus  a  human  uterus  furnished  with  cornua  may  be  said 
to  represent,  in  a  rudimentary  condition,  the  same  organ  in  its 
normal  state  in  certain  mammals.  Some  parts  which  are  rudi- 
mentary in  man,  as  the  os  coccyx  in  both  sexes,  and  the  mammee 
in  the  male  sex,  are  always  present ;  whilst  others,  such  as  the 
supracondyloid  foramen,  only  occasionally  appear,  and  therefore 
might  have  been  introduced  under  the  head  of  reversion.  These 
several  reversionary  structures,  as  well  as  the  strictly  rudi- 
mentary ones,  reveal  the  descent  of  man  from  some  lower  form 
in  an  unmistakable  manner. 

Correlated  Variation. — In  man,  as  in  the  lower  animals,  many 
structures  are  so  intimately  related,  that  when  one  part  varies 
so  does  another,  without  our  being  able,  in  most  cases,  to -assign 
any  reason.  We  cannot  say  whether  the  one  i^art  governs  the 
other,  or  whether  both  are  governed  by  some  earlier  developed 


possiibility  of  either  of  his  first  pro-  closely  the  variations  resemble  the 

positions.     Prof.  Macalister  has  also  normal  muscles    of  the    lower  ani- 

describeJ    ('  Proc.    R.    Irish    Acad.'  mals.     He  sums  up  by  remarking, 

vol.  X.  1864,  p.   138)  variations  in  "  It  will  be  enough  for  my  purpose 

the/ej:or/)oWiciS  /on^M5,  remarkable  "if  I   have    succeeded    in   shewing 

from    their   relations    to    the    same  "  the  more  important  forms  whicli, 

muscle  in  the  Quadrumann.  "  when  occurring  as  varieties  in  the 

^■*  Since  the  first  edition  of  this  *'  human  subject,  tend  to  exhibit  in 

book  appeared,  Mr.  Wood  has  pub-  "  a  sufficiently  marked  manner  what 

lished  another  memoir  in  the  '  Phil.  "  may  be  considered  as  proofs  and 

Transactions,'    1870,   p.  83,  on   the  "  examples  of  the  Darwinian  prin- 

varieties  of  the  muscles  of  the  human  "  ciple  of  reversion,  or  law  of  in- 

neck,  shoulder,  and  chest.     He  here  "  heritance,  in    this  department  ot 

shews  how  extremely  variable  these  "  anatomical  science." 
muscles  are,  and  how  often  and  how 


44  The  Descent  of  Man.  Part  I. 


part.  Various  monstrosities,  as  I.  Geoffroy  repeatedly  insists,  are 
thus  intimately  connected.  Homologous  structures  are  par- 
ticularly liable  to  change  together,  as  we  see  on  the  opposite 
sides  of  the  body,  and  in  the  upper  and  lower  extremities. 
Meckel  long  ago  remarked,  that  when  the  muscles  of  the  arm 
depart  from  their  proper  type,  they  almost  always  imitate  those 
of  the  leg ;  and  so,  conversely,  with  the  muscles  of  the  legs.  The 
organs  of  sight  and  hearing,  the  teeth  and  hair,  the  colour  of  the 
skin  and  of  the  hair,  colour  and  constitution,  are  more  or  less  cor- 
related.^^ Professor  Schaaffhausen  first  drew  attention  to  the 
relation  apparently  existing  between  a  muscular  frame  and 
the  strongly-pronounced  supra-orbital  ridges,  which  are  so 
characteristic  of  the  lower  races  of  man. 

Besides  the  variations  which  can  be  grouped  with  more  or 
less  probability  under  the  foregoing  heads,  there  is  a  large  class 
of  variations  which  may  be  provisionally  called  spontaneous,  for 
to  our  ignorance  they  appear  to  arise  without  any  exciting 
cause.  It  can,  however,  be  shewn  that  such  variations,  whether 
consisting  of  slight  individual  differences,  or  of  strongly-marked 
and  abrupt  deviations  of  structure,  depend  much  more  on  the 
constitution  of  the  organism  than  on  the  nature  of  the  condi- 
tions to  which  it  has  been  subjected.^^ 

Bate  of  Increase. — Civilised  populations  have  been  known 
under  favourable  conditions,  as  in  the  United  States,  to  double 
their  numbers  in  twenty-five  years ;  and,  according  to  a  calcula- 
tion by  Euler,  this  might  occur  in  a  little  over  twelve  years.^'^  At 
the  former  rate,  the  present  population  of  the  United  States 
(thirty  millions),  would  in  657  years  cover  the  whole  terraqueous 
globe  so  thickly,  that  four  men  would  have  to  stand  on  each 
square  yard  of  surface.  The  primary  or  fundamental  check  to 
the  continued  increase  of  man  is  the  difficulty  of  gaining 
subsistence,  and  of  living  in  comfort.  We  may  infer  that  this  is 
the  case  from  what  we  see,  for  instance,  in  the  United  States, 
where  subsistence  is  easy,  and  there  is  plenty  of  room.  If  such 
means  were  suddenly  doubled  in  Great  Britain,  our  number  would 
be  quickly  doubled.  With  civilised  nations  this  primary  check 
acts  chiefly  by  restraining  marriages.  The  greater  death-rate  of 
infants  in  the  poorest  classes  is  also  very  important;  as  well  as 

'*  The  authorities  for  these  seve-  my  '  Variation  of  Animals  and  Plants 

ral    statements    are    given    in    my  under  Domestication.' 
'Variation    of  Animals   under    Do-  *^  See  the  ever  memorable 'Essay 

mestication,'  vol.  ii.  pp.  320-335,  on  the  Principle  of  Population,'  by 

^^  This  whole    subject    has    been  the  Rev.  T.  Malthus,  vol.  i.  1826,  p, 

discussed    in  chap,  xxiii.  vol.  ii.  of  6,  517. 


Chap.  U.  jMajiucr  of  Dcvelopnieiit.  45 

the  gi-eatcr  mortality,  from  various  diseases,  of  the  inhabitants  of 
crowded  and  miserable  houses,  at  all  ages.  The  effects  of  severe 
epidemics  and  wars  are  soon  counterbalanced,  and  more  than 
counterbalanced,  in  nations  placed  under  favourable  conditions. 
Emigration  also  comes  in  aid  as  a  temporary  check,  but,  with 
the  extremely  poor  classes,  not  to  any  great  extent. 

There  is  reason  to  suspect,  as  Malthus  has  remarked,  that  the 
reproductive  power  is  actually  less  in  barbarous,  than  in  civilised 
races.  We  know  nothing  positively  on  this  head,  for  with 
savages  no  census  has  been  taken;  but  from  the  concurrent 
testimony  of  missionaries,  and  of  others  who  have  long  resided 
with  such  people,  it  appears  that  their  families  are  usually  small, 
and  large  ones  rare.  This  may  be  partly  accounted  for,  as  it  is 
believed,  by  the  women  suckling  their  infants  during  a  long 
time ;  but  it  is  highly  probable  that  savages,  who  often  suffer 
much  hardship,  and  who  do  not  obtain  so  much  nutritious  food 
as  civilised  men,  would  be  actually  less  prolific.  I  have  shewn 
in  a  former  work,^^  that  all  our  domesticated  quadrupeds  and 
birds,  and  all  our  cultivated  plants,  are  more  fertile  than  the 
corresponding  species  in  a  state  of  nature.  It  is  no  valid 
objection  to  this  conclusion  that  animals  suddenly  supplied  with 
an  excess  of  food,  or  when  grown  very  fat ;  and  that  most  plants 
on  sudden  removal  from  very  poor  to  very  rich  soil,  are 
rendered  more  or  less  sterile.  We  might,  therefore,  expect  that 
civilised  men,  who  in  one  sense  are  highly  domesticated,  would 
be  more  prolific  than  wild  men.  It  is  also  probable  that  the 
increased  fertility  of  civilised  nations  would  become,  as  with  our 
domestic  animals,  an  inherited  character :  it  is  at  least  known 
that  with  mankind  a  tendency  to  produce  twins  runs  in 
families.^^ 

Notwithstanding  that  savages  appear  to  be  less  prolific  than 
civilised  people,  they  would  no  doubt  rapidly  increase  if  their 
numbers  were  not  by  some  means  rigidly  kept  down.  The  San- 
tali,  or  hill-tril)es  of  India,  have  recently  afforded  a  good  illustra- 
tion of  this  fact ;  for,  as  shewn  by  Mr.  Hunter,*^  they  have 
increased  at  an  extraordinary  rate  since  vaccination  has  been 
introduced,  other  pestilences  mitigated,  and  war  sternly  repressed. 
This  increase,  however,  would  not  have  been  possible  had  not 
these  rude  people  spread  into  the  adjoining  districts,  and  worked 
for  hire.  Savages  almost  always  marry  ;  yet  there  is  some 
prudential  restraint,  for  they  do  not  commonly  marry  at  the 

**  'Variation     of    Animals,  and  Foreign    Medico-Chirurg.     Pieview,' 

Plants  under  Domestication,'  vol.  ii.  July,  1863,  p.  170. 
pp.  111-113,  163.  ««  '  The  Annals  of  Rural  Bengal,' 

"  Mr.    Sedgwick,    'British     and  bv  \V.  W.  Hunter,  1868,  p.  259. 


46  The  Descent  of  Man.  Part  I. 

earliest  possible  age.  The  young  men  are  often  required  to  shew 
that  they  can  support  a  wife ;  and  they  generally  have  first  to 
earn  the  price  with  which  to  purchase  her  from  her  parents. 
With  savages  the  difficulty  of  obtaining  subsistence  occasionally 
limits  their  number  in  a  much  more  direct  manner  than  with 
civilised  people,  for  all  tribes  periodically  suffer  from  severe 
famines.  At  such  times  savages  are  forced  to  devour  much  bad 
food,  and  their  health  can  hardly  fail  to  be  injured.  Many 
accounts  have  been  published  of  their  protruding  stomachs  and 
emaciated  limbs  after  and  during  famines.  They  are  then,  also, 
compelled  to  wander  much,  and,  as  I  was  assured  in  Australia, 
their  infants  perish  in  large  numbers.  As  famines  are  period- 
ical, depending  chiefly  on  extreme  seasons,  all  tribes  must 
fluctuate  in  number.  They  cannot  steadily  and  regularly 
increase,  as  there  is  no  artificial  increase  in  the  supply  of  .food. 
Savages,  when  hard  pressed,  encroach  on  each  other's  territories, 
and  war  is  the  result ;  but  they  are  indeed  almost  always  at  war 
with  their  neighbours.  They  are  liable  to  many  accidents  on 
land  and  water  in  their  search  for  food ;  and  in  some  countries 
they  sufi"er  much  from  the  larger  beasts  of  prey.  Even  in 
India,  districts  have  been  depopulated  by  the  ravages  of 
tigers. 

Malthus  has  discussed  these  several  checks,  but  he  does  not 
lay  stress  enough  on  what  is  probably  the  most  important  of  all, 
namely  infanticide,  especially  of  female  infants,  and  the  habit  of 
procuring  abortion.  These  practices  now  prevail  in  many 
quarters  of  the  world ;  and  infanticide  seems  formerly  to  have 
prevailed,  as  Mr.  ]\['Lennan'^^  has  shewn,  on  a  still  more  extensive 
scale.  These  practices  appear  to  have  originated  in  savages  re- 
cognising the  difficulty,  or  rather  the  impossibility  of  supporting 
all  the  infants  that  are  born.  Licentiousness  may  also  be  added 
to  the  foregoing  checks;  but  this  does  not  follow  from  failing 
means  of  subsistence ;  though  there  is  reason  to  believe  that  in 
some  cases  (as  in  Japan)  it  has  been  intentionally  encouraged 
as  a  means  of  keeping  down  the  population. 

If  we  look  back  to  an  extremely  remote  epoch,  before  man  had 
arrived  at  the  dignity  of  manhood,  he  would  have  been  guided 
more  by  instinct  and  less  by  reason  than  are  the  lowest  savages 
at  the  present  time.  Our  early  semi-human  progenitors  would 
not  have  practised  infanticide  or  polyandry ;  for  the  instincts  of 
the  lower  animals  are  never  so  perverted  ^-  as  to  lead  them  re- 

**  '  Primitive  Marriage,'  1865.  ments  as  follows  on  this  passage  : — 

^^  A    writer   in   the    'Spectator'      "  Mr.  Darwin  finds  himself  compelled 

(March   12th,  1871,    p.   320)    com-      "  to  reintroduce  a  new  doctrine  of  the 


Chap.  II.  Manner  of  Devclopuient.  47 

gularly  to  destroy  their  own  offspring,  or  to  be  quite  devoid  of 
jealousy.  There  would  have  been  no  prudential  restraint  from 
marriage,  and  the  sexes  would  have  freely  united  at  an  early  age. 
Hence  the  progenitors  of  man  would  have  tended  to  increase 
rapidly;  but  checks  of  some  kind,  either  periodical  or  constant, 
must  have  kept  down  their  numbers,  even  more  severely  than 
with  existing  savages.  What  the  precise  nature  of  these  checks 
were,  we  cannot  say,  any  more  than  with  most  other  animals. 
We  know  that  horses  and  cattle,  which  are  not  extremely  prolific 
animals,  when  first  turned  loose  in  South  America,  increased  at 
an  enormous  rate.  The  elephant,  the  slowest  breeder  of  all 
known  animals,  would  in  a  few  thousand  years  stock  the  whole 
world.  The  increase  of  every  species  of  monkey  must  be 
checked  by  some  means;  but  not,  as  Brehm  remarks,  by  the 
attacks  of  beasts  of  prey.  No  one  w^ill  assume  that  the  actual 
power  of  reproduction  in  the  wild  horses  and  cattle  of  America, 
was  at  first  in  any  sensible  degree  increased ;  or  that,  as  each 
district  became  fully  stocked,  this  same  power  was  diminished. 
No  doubt  in  this  case,  and  in  all  others,  many  checks  concur, 
and  different  checks  under  different  circumstances ;  periodical 
dearths,  depending  on  unfavourable  seasons,  being  probably  the 
most  important  of  all.  So  it  will  have  been  with  the  early  pro- 
genitors of  man. 

Natural  Selection. — We  have  now  seen  that  man  is  variable  in 
body  and  mind;  and  that  the  variations  are  induced,  either 
directly  or  indirectly,  by  the  same  general  causes,  and  obey  the 
same  general  laws,  as  with  the  low^er  animals.  Man  has  spread 
widely  over  the  face  of  the  earth,  and  must  have  been  exposed, 
during  his  incessant  migrations,^^  to  the  most  diversified  con- 
ditions. The  inhabitants  of  Tierra  del  Fuego,  the  Cape  of  Good 
Hope,  and  Tasmania  in  the  one  hemisphere,  and  of  the  Arctic 
regions  in  the  other,  must  have  passed  through  many  climates, 


"  fiill  of  man.     He  shews  that  the  *'  by    the    many    foul   customs,    es- 

"  instincts    of   the    higher    animals  "  pecially  as  to  marriage,  of  savage 

"  are  far  nobler  than  the  habits  of  "  tribes.     What     does     the    Jewish 

"  savage  races  of  men,  and  he  finds  "  tradition  of  the  moral  degonera- 

"  himself,    therefore,    compelled    to  "  tion  of  man  through  his  snatching 

"  re-introduce, — in    a   form    of  the  "  at    a    knowledge    forbidden    him 

"  substantial  orthodoxy  of  which  he  "  by    his     highest    instinct    a'^sert 

"  appears  to  be  quite  unconscious, —  "  beyond  this?" 

"  and   to   introduce    as   a   scientific  ®^  See  some  good  remarks  to  this 

"  hypothesis  the  doctrine  that  man's  effect    by  W.    Stanley    Jevons,    "  A 

"  gain  of  knoicledje  was  the  cause  of  "  Deduction  from  Darwin's  Theory," 

"a    temporary    but    long-enduriug  '  Nature,'  1869,  p.  231. 
"  moral    deterioration,   as  indicated 


48  TJie  Descent  of  Man.  Part  I. 

and  changed  their  habits  many  times,  before  they  reached  their 
present  homes.®*  The  early  progenitors  of  man  must  also  have 
tended,  like  all  other  animals,  to  have  increased  beyond  their 
means  of  subsistence  they  must,  therefore,  occasionally  have 
been  exposed  to  a  struggle  for  existence,  and  consequently  to  the 
rigid  law  of  natural  selection.  Beneficial  variations  of  all  kinds 
will  thus,  either  occasionally  or  habitually,  have  been  preserved, 
and  injurious  ones  eliminated.  I  do  not  refer  to  strongly-marked 
deviations  of  structure,  which  occur  only  at  long  intervals  of 
time,  but  to  mere  individual  differences.  We  know,  for  instance, 
that  the  muscles  of  our  hands  and  feet,  which  determine  our 
powers  of  movement,  are  liable,  hke  those  of  the  lower  animals,^^ 
to  incessant  variability.  If  then  the  progenitors  of  man  inhabit- 
ing any  district,  especially  one  undergoing  some  change  in  its 
conditions,  were  divided  into  two  equal  bodies,  the  one  half 
which  included  all  the  individuals  best  adapted  by  their  powers 
of  movement  for  gaining  subsistence,  or  for  defending  themselves, 
would  on  an  average  survive  in  greater  numbers,  and  procreate 
more  offspring  than  the  other  and  less  well  endowed  half. 

Man  in  the  rudest  state  in  which  he  now  exists  is  the  most 
dominant  animal  that  has  ever  appeared  on  this  earth.  He  has 
spread  more  widely  that  any  other  highly  organised  form :  and 
all  others  have  yielded  before  him.  He  manifestly  owes  this 
immense  superiority  to  his  intellectual  faculties,  to  his  social 
habits,  wliich  lead  him  to  aid  and  defend  his  fellows,  and  to  his 
corporeal  structure.  The  supreme  importance  of  these  characters 
has  been  proved  by  the  final  arbitrament  of  the  battle  for  life. 
Through  his  powers  of  intellect,  articulate  language  has  been 
evolved;  and  on  this  his  wonderful  advancement  has  mainly 
depended.  As  Mr.  Chauncey  Wright  remarks  :®s  "  a  psychological 
"  analysis  of  the  faculty  of  language  shews,  that  even  the  smallest 
"  proficiency  in  it  might  require  more  brain  power  than  the 
"  greatest  proficiency  in  any  other  direction.''  He  has  invented 
and  is  able  to  use  various  weapons,  tools,  traps,  &c.,  with  which 
he  defends  himself,  kills  or  catches  prey,  and  otherwise  obtains 
food.  He  has  made  rafts  or  canoes  for  fishing  or  crossing  over 
to  neighbouring  fertile  islands.     He  has  discovered  the  art  of 

^*  Latham,  '  Man  and  his  Migra-  "  classed     in     any    of    the     above 

tions,'  1851,  p.  135.  "  groups."      These    muscles    differ 

•^^  Messrs.  Murie    and  Mivart  in  even   on    the    opposite  sides  of  the 

their  'Anatomy  of  the  Lemuroidea'  same  individual. 

('  Transact.    Zoolog.    Soc'    vol.    vii.  ^^  Limits    of    Natural    Selection, 

1869,  pp.  96-98)  say,  "some- muscles  'North     American     Review,'     Oct. 

"  are  so  irregular  in  their  distribu-  1870,  p.  295. 
"  tion    that"  they    cannot    be    well 


Chap.  II.  Manner  of  Develcpmeiit.  49 

making  fire,  by  wliicli  hard  and  stringy  roots  can  be  rendered 
digestible,  and  poisonous  roots  or  herbs  innocuous.  This  dis- 
covery of  fire,  probably  the  greatest  ever  made  by  man,  excepting 
language,  dates  from  before  the  dawn  of  history.  These  several 
inventions,  by  wliich  man  in  the  rudest  state  has  become  so  pre- 
eminent, are  the  direct  results  of  the  development  of  his  powers 
of  observation,  memory,  curiosity,  imagination,  and  reason.  I 
cannot,  therefore,  understand  how  it  is  that  Mr.  Wallace"  main- 
tains, that  **  natural  selection  could  only  have  endowpd  the 
"  savage  with  a  brain  a  little  superior  to  that  of  an  ape." 

Although  the  intellectual  powers  and  social  habits  of  man  are 
of  paramount  importance  to  him,  we  must  not  underrate  the 
importance  of  his  bodily  structure,  to  which  subject  the  remain- 
der of  this  chapter  will  be  devoted ;  the  development  of  the  in- 
tellectual and  social  or  moral  faculties  being  discussed  in  a  later 
chapter. 

Even  to  hammer  with  precision  is  no  easy  matter,  as  every 
one  who  has  tried  to  learn  carpentry  will  admit.  To  throw^  a 
stone  with  as  true  an  aim  as  a  Fuegian  in  defending  himself,  or 
in  killing  birds,  requires  the  most  consummate  perfection  in  the 
correlated  action  of  the  muscles  of  the  hand,  arm,  and  shoulder, 
and,  further,  a  fine  sense  of  touch.  In  throwing  a  stone  or  spear, 
and  in  many  other  actions,  a  man  must  stand  firmly  on  his  feet ; 
and  this  again  demands  the  perfect  co-adaptation  of  numerous 
muscles.  To  chip  a  flint  into  the  rudest  tool,  or  to  form  a 
barbed  spear  or  hook  from  a  bone,  demands  the  use  of  a  perfect 
hand ;  for,  as  a  most  capable  judge,  Mr.  Schoolcraft,^^  remarks, 
the  shaping  fragments  of  stone  into  knives,  lances,  or  arrows-heads, 

°^  '  Quarterly      Review,'      April  hoi-e    resist    quoting    a    most   just 

1869,  p.  392.  'This  subject  is  more  remark  by  Sir  J.  Lubbock  (' Pre- 
fuUy  discussed  in  Mr.  Wallace's  historic  Times,'  1865,  p.  479)  in 
'  Contributions  to  the  Theory  of  reference  to  this  paper,  namely,  that 
Natural  Selection,'  1870,  in  which  Mr.  Wallace,  "with  characteristic 
all  the  essays  referred  lo  in  this  "  unselfishness,  ascribes  it  (i.  e.  the 
work  are  republished.  The  '  Essay  "  idea  of  natural  selection)  unre- 
on  Man '  has  been  ably  criticised  by  "  servedly  to  Mr.  Darwin,  although, 
Prof.  Claparede,  one  of  the  most  "  as  is  well  known,  he  struck  out 
distinguished  zoologists  in  Europe,  "  the  idea  independently,  and  pub- 
in  an  article  published  iu  the  "  lished  it,  though  not  with  the 
'  Bibliotheque      Universelle,'     June  "  same    elaboration,    at    the    same 

1870.  The   remark  quoted   in    my      "  time." 

text  will  surprise  every  one  who  ®*  Quoted  by  Mr.  Lawson  Tait  in 
has  read  Mr.  Wallace's  celebrated  his  '  Law  of  Natural  Selection,' — 
paper  on  'The  Origin  of  Human  '  Dublin  Quarterly  Journal  of  Modi- 
Races  deduced  from  the  Theory  of  cal  Science,'  Feb.  1869.  Dr.  Keller 
Natural  Selection,'  originally  pub-  is  likewise  quoted  to  the  same 
lished  in  the  '  Anthropological  Re-  effect, 
view,' !^L^y  1864,  p.  chili.     I    cannot 


50  The  Descent  of  Man.  Part  I. 

shews  "  extraordinary  ability  and  long  practice."  This  is  to  a 
great  extent  proved  by  the  fact  that  primeyal  men  practised  a 
division  of  labour;  each  man  did  not  manufacture  his  own  flint 
tools  or  rude  pottery,  but  certain  individuals  appear  to  have 
devoted  themselves  to  such  work,  no  doubt  receiving  in  exchange 
the  produce  of  the  chase.  Archaeologists  are  convinced  that  an 
enormous  interval  of  time  elapsed  before  our  ancestors  thought 
of  grinding  chipped  flints  into  smooth  tools.  One  can  hardly 
doubt,  that  a  man-like  animal  who  possessed  a  hand  and  arm 
sufficiently  perfect  to  throw  a  stone  with  precision,  or  to  form  a 
flint  into  a  rude  tool,  could,  with  sufficient  joractice,  as  far  as 
mechanical  skill  alone  is  concerned,  make  almost  anything 
which  a  civilised  man  can  make.  The  structure  of  the  hand  in 
this  respect  may  be  compared  with  that  of  the  vocal  organs, 
which  in  the  apes  are  used  for  uttering  various  signal-cries,  or, 
as  in  one  genus,  musical  cadences;  but  in  man  the  closely 
similar  vocal  organs  have  become  adapted  through  the  inherited 
eifects  of  use  for  the  utterance  of  articulate  language. 

Turning  now  to  the  nearest  aUies  of  men,  and  therefore  to  the 
best  representatives  of  our  early  progenitors,  we  find  that  the 
hands  of  the  Quadrumana  are  constructed  on  the  same  general 
pattern  as  our  own,  but  are  far  less  perfectly  adapted  for  diver- 
sified uses.  Their  hands  do  not  serve  for  locomotion  so  well 
as  the  feet  of  a  dog ;  as  may  be  seen  in  such  monkeys  as  the 
chimpanzee  and  orang,  which  walk  on  the  outer  margins  of 
the  palms,  or  on  the  knuckles. ^^  Their  hands,  however,  are 
acbnirably  adapted  for  climbing  trees.  Monkeys  seize  thin 
branches  or  ropes,  with  the  thumb  on  one  side  and  the  fingers 
and  jDalm  on  the  other,  in  the  same  manner  as  we  do.  They  can 
thus  also  lift  rather  large  objects,  such  as  the  neck  of  a  bottle,  to 
their  mouths.  Baboons  turn  over  stones,  and  scratch  up  roots 
with  their  hands.  They  seize  nuts,  insects,  or  other  small 
objects  with  the  thumb  in  opposition  to  the  fingers,  and  no 
doubt  they  thus  extract  eggs  and  the  young  from  the  nests  of 
bii'ds.  American  monkeys  beat  the  wild  oranges  on  the  branches 
until  the  rind  is  ci-acked,  and  then  tear  it  off  with  the  fingers  of 
the  two  hands.  In  a  wild  state  they  break  open  hard  fruits 
with  stones.  Other  monkeys  open  mussel-shells  with  the  two 
thumbs-  With  their  fingers  they  pull  out  thorns  and  burs,  and 
hunt  for  each  other's  parasites.  They  roll  down  stones,  or  throw 
them  at  their  enemies  :  nevertheless,  they  are  clumsy  in  these 
various  actions,  and,  as  I  have  myself  seen,  are  quite  unable  to 
tlirow  a  stone  with  precision, 

*^  Owen,     Anatomy  of  Vertebrattis/  vol.  iii.  p.  71. 


CiiAi   II.  Manner  of  Development.  51 


It  seems  to  me  far  from  true  that  because  "  objects  are  grasped 
"  clumsily "  by  monkeys,  "  a  much  less  specialised  organ  of 
"  prehension "  would  have  served  them ''"  equally  "well  with 
their  present  hands.  On  the  contrary,  I  see  no  reason  to  doubt 
that  more  perfectly  constructed  hands  would  have  been  an 
advantage  to  them,  provided  that  they  were  not  thus  rendered 
less  fitted  for  climbing  trees.  We  may  suspect  that  a  hand  as 
perfect  as  that  of  man  would  have  been  disadvantageous  for 
climbing;  for  the  most  arboreal  monkeys. iii  the  world,  namely, 
Ateles  in  America,  Colobus  in  Africa,  and  Hylobates  in  Asia, 
are  either  thumbless,  or  their  toes  partially  cohere,  so  that  their 
limbs  are  converted  into  mere  grasping  hooks,'^ 

As  soon  as  some  oncient  member  in  the  great  series  of  the 
Primates  came  to  be  less  arboreal,  owing  to  a  change  in  its 
manner  of  procuring  subsistence,  or  to  some  change  in  the 
surrounding  conditions,  its  habitual  manner  of  progression  would 
have  been  modified  :  and  thus  it  would  have  been  rendered  more 
strictly  quadrupedal  or  bipedal.  Baboons  frequent  hilly  and 
rocky  districts,  and  only  from  necessity  climb  high  trees ; '-  and 
they  have  acquired  almost  the  gait  of  a  dog.  Man  alone  has 
become  a  biped ;  and  we  can,  I  think,  partly  see  how  he  has 
come  to  assume  his  erect  attitude,  which  forms  one  of  his  most 
conspicuous  characters.  Man  could  not  have  attained  his  present 
dominant  posijtion  in  the  world  without  the  use  of  his  hands, 
which  are  so  admirably  adapted  to  act  in  obedience  to  his  will. 
Sir  C.  Bell  "^  insists  that  "  the  hand  supplies  all  instruments, 
"  and  by  its  correspondence  with  the  intellect  gives  him  univer- 
"  sal  dominion."  But  the  hands  and  arms  could  hardly  have 
become  perfect  enough  to  have  manufactured  weapons,  or  to 
have  hurled  stones  and  sj)ears  with  a  true  aim,  as  long  as  they 
were  habitually  used  for  locomotion  and  for  supporting  the 
whole  weight  of  the  body,  or,  as  before  remarked,  so  long  as  they 
were  especially  fitted  for  climbing  trees.  Such  rough  treatment 
would  also  have  blunted  the  sense  of  touch,  on  which  their 
delicate  use  largely  depends.  From  these  causes  alone  it  would 
have  been  an  advantage  to  man  to  become  a  biped;  but  for 

^0  '  Quarterly     Review,'       April  but  whether  a  better  climber  than 

1869,  p.  392.  the  species  of  the  allied  genera,  1  do 

'1  In    IrLjlohntcs    siindactylus,    as  not  know.     It  deserves  notice  that 

the  name  expresses,  two  of  the  toes  the    feet    of  the    sloths,    the    most 

regularly  cohere  ;  and  this,  as  Mr.  arboreal  animals  in  the  world,  are 

Blyth    informs   me,  is   occasionally  wonderfully  hook-like, 

the  case  with  the  toes  of  H.  a(]ilis,  "  Brehm,    '  Thierlobon,'   B.   i.   s, 

/a;-,  and /eucj'scMS.    Colobus  is  strictly  80. 

arboreal  and  extraordinarily  active  "'  ''The    Hand,"    <&:c.      *  Bridge- 

(Brehm,   '  Thierleben,'  B.  i.  s.  50),  water  Treatise,'  1833,  p.  38. 


52  The  Descent  of  Man.  Part  I. 

many  actions  it  is  indispensable  that  the  arms  and  whole  upper 
part  of  the  body  should  be  free  ;  and  he  must  for  this  end  stand 
firmly  on  his  feet.  To  gain  this  great  advantage,  the  feet  have 
been  rendered  flat ;  and  the  great  toe  has  been  peculiarly  modi- 
fied, though  this  has  entailed  the  almost  complete  loss  of  its 
power  of  prehension.  It  accords  with  the  principle  of  the 
division  of  physiological  labour,  prevailing  throughout  the 
animal  kingdom,  that  as  the  hands  became  perfected  for  i^re- 
hension,  the  feet  should  have  become  perfected  for  support  and  ■ 
locomotion.  "With  some  savages,  however,  the  foot  has  not 
altogether  lost  its  prehensile  power,  as  shewn  by  their  manner 
of  climbing  trees,  and  of  using  them  in  other  ways.'^* 

If  it  be  an  advantage  to  man  to  stand  firmly  on  his  feet  and  to 
have  his  hands  and  arms  free,  of  which,  from  his  pre-eminent 
success  in  the  battle  of  life,  there  can  be  no  doubt,  then  I  can  see 
no  reason  why  it  should  not  have  been  advantageous  to  the 
progenitors  of  man  to  have  become  more  and  more  erect  or 
bipedal.  They  would  thus  have  been  better  able  to  defend 
themselves  with  stones  or  clubs,  to  attack  their  prey,  or  other- 
wise to  obtain  food.  The  best  built  individuals  would  in  the 
long  run  have  succeeded  best,  and  have  survived  in  larger 
numbers.  If  the  gorilla  and  a  few  allied  forms  had  becomn. 
extinct,  it  might  have  been  argued,  with  great  force  and  apparent 
truth,  that  an  animal  could  not  have  been  gradually  converted 
from  a  quadruped  into  a  biped,  as  all  the  individuals  in  an 
intermediate  condition  would  have  been  miserably  ill-fitted 
for  progression.  But  we  know  (and  this  is  well  worthy  of 
reflection)  that  the  anthropomorphous  apes  are  now  actually  in 
an  intermediate  condition ;  and  no  one  doubts  that  they  are  on 
the  whole  well  adapted  for  their  conditions  of  life.  Thus  the 
gorilla  runs  with  a  sidelong  shambling  gait,  but  more  commonly 
progresses  by  resting  on  its  bent  hands.  The  long-armed  apes 
occasionally  use  their  arms  like  crutches,  swinging  their  bodies 
forward  between  them,  and  some  kinds  of  Hylobates,  without 
having  been  taught,  can  walk  or  run  upright  with  tolerable 
quickness ;  yet  they  move  awkwardly,  and  much  less  securely 
than  man.  We  see,  in  short,  in  existing  monkeys  a  manner  of 
progression  intermediate  between  that  of  a  quadruped  and  a 

^*  Hackel    has   an   excellent   dis-  foot  as  a  prehensile  organ  by  man ; 

cussion  on  the  steps  by  which  man  and  has  also  written  on  the  manner 

became  a  biped:  '  Natiirliche  Schop-  of  progression  of  the  higher  apes,  to 

fungsgeschichte,'  1868,  s.  507.     Dr.  Avhich    I    allude    in    the    following 

Biichner  ('  Conferences  sur  la  Theorie  paragraph  :  see  also  Owen  ('  Anatomy 

Darwinienne,'    1869,    p.     135)    has  of  Vertebrates,'  vol,  iii.  p.  71)  on 

given  good  cases  of  the  use  of  the  this  latter  subject. 


C:  H  A  p.  I  r .  Manner  of  Developnien  t.  53 

biped ;  but,  as  an  unprejudiced  judge  '^  insists,  the  anthropomor- 
phous apes  approach  in  structure  more  nearly  to  the  bipedal 
than  to  the  quadrupedal  type. 

As  the  progenitors  of  man  became  more  and  more  erect,  with 
their  hands  and  arms  more  and  more  modified  for  prehension 
and  other  purposes,  with  their  feet  and  legs  at  the  same  time 
transformed  for  firm  support  and  progression,  endless  other 
changes  of  structure  would  have  become  necessary.  The  pelvis 
•would  have  to  be  broadened,  the  spine  peculiarly  curved,  and  the 
head  fixed  in  an  altered  position,  all  which  changes  have  been 
attained  by  man.  Prof.  Schaaflihausen  "  maintains  that  "the 
"  powerful  mastoid  processes  of  the  human  skull  are  the  result  of 
"  his  erect  position  ;"  and  these  processes  are  absent  in  the  orang, 
chimpanzee,  &c.,  and  are  smaller  in  the  gorilla  than  in  man. 
Various  other  structures,  which  appear  connected  with  man's 
erect  position,  might  here  have  been  added.  It  is  very  difficult 
to  decide  how  far  these  correlated  modifications  are  the  result  of 
natural  selection,  and  how  far  of  the  inherited  effects  of  the 
increased  use  of  certain  parts,  or  of  the  action  of  one  part  on 
another.  No  doubt  these  means  of  change  often  co-operate  :  thus 
when  certain  muscles,  and  the  crests  of  bone  to  which  they  are 
attached,  become  enlarged  by  habitual  use,  this  shews  that 
certain  actions  are  habitually  performed  and  must  be  serviceable. 
Hence  the  individuals  which  performed  them  best,  would  tend 
to  survive  in  greater  numbers. 

The  free  use  of  the  arms  and  hands,  partly  the  cause  and 
partly  the  result  of  man's  erect  position,  appears  to  have  led  in  an 
indirect  manner  to  other  modifications  of  structure.  The  early 
male  forefathers  of  man  were,  as  previously  stated,  jDrobably 
furnished  with  great  canine  teeth;  but  as  they  gradually 
acquired  the  habit  of  using  stones,  clubs,  or  other  weaj^ons,  for 
fighting  with  their  enemies  or  rivals,  they  would  use  their  jaws 
and  teeth  less  and  less.  In  this  case,  the  jaws,  together  with  the 
teeth,  w^ould  become  reduced  in  size,  as  we  may  feel  almost  sure 
from  innumerable  analogous  cases.  In  a  future  chapter  we 
shall  meet  with  a  closely  parallel  case,  in  the  reduction  or  com- 
plete disappearance  of  the  canine  teeth  in  male  ruminants, 
apparently  in  relation  with  the  development  of  their  horns ;  and 
in  horses,  in  relation  to  tlieii'  habit  of  fighting  with  their  incisor 
teeth  and  hoofs. 

"  Prof.    Broca,    La    Constitution  the  Skull,'  translated  iu    '  Authro- 

des  Vertfebres   ciiudales;  'La  Revue  pological    Review,'    Oct.     18*38,    p. 

d'Anthropologie,'      1872,      p.      iG,  428.     Owen  ('  Anatomy   of  Verte- 

(separate  copy)  brates,'  vol.  ii.  1866,  p.  551)  on  the 

'•^  'On    the    rrimitive    Form     of  mastoid  processes  in  the  higher  apes. 


54  The  Descent  of  Man.  Tart  I. 

In  the  adult  male  anthropomorphous  apes,  as  Eutimeyer/''" 
and  others,  have  insisted,  it  is  the  effect  on  the  skull  of  the  great 
development  of  the  jaw-muscles  that  causes  it  to  differ  so  greatly 
in  many  respects  from  that  of  man,  and  has  given  to  these 
animals  "  a  truly  frightful  physiognomy."  Therefore,  as  the  jaws 
and  teeth  in  man's  progenitors  gradually  became  reduced  in  size, 
the  adult  skull  would  have  come  to  resemble  more  and  more 
that  of  existing  man.  As  we  shall  hereafter  see,  a  great  reduction 
of  the  canine  teeth  in  the  males  would  almost  certainly  affect  the 
teeth  of  the  females  through  inheritance. 

As  the  various  mental  faculties  gradually  developed  themselves 
the  brain  would  almost  certainly  become  larger.  No  one,  I 
presume,  doubts  that  the  large  proportion  which  the  size  of 
man's  brain  bears  to  his  body,  compared  to  the  same  proportion 
in  the  gorilla  or  orang,  is  closely  connected  vfith  his  higher 
mental  powers.  We  meet  with  closely  analogous  facts  with 
insects,  for  in  ants  the  cerebral  ganglia  are  of  extraordinary 
dimensions,  and  in  all  the  Hymenoptera  these  ganglia  are  many 
times  larger  than  in  the  less  intelligent  orders,  such  as  beetles. '^^ 
On  the  other  hand,  no  one  supposes  that  the  intellect  of  any 
two  animals  or  of  any  two  men  can  be  accurately  gauged  by  the 
cubic  contents  of  their  skulls.  It  is  certain  that  there  may  be 
extraordinary  mental  activity  with  an  extremely  small  absolute 
mass  of  nervous  matter:  thus  the  wonderfully  diversified 
instincts,  mental  powers,  a-nd  affections  of  ants  are  notorious, 
yet  their  cerebral  ganglia  are  not  so  large  as  the  quarter  of  a 
small  pin's  head.  Under  this  point  of  view,  the  brain  of  an  ant  is 
one  of  the  most  marvellous  atoms  of  matter  in  the  world,  perhaps 
more  so  than  the  brain  of  a  man. 

The  belief  that  there  exists  in  man  some  close  relation  between 
the  size  of  the  brain  and  the  development  of  the  intellectual 
faculties  is  supported  by  the  comparison  of  the  skulls  of  savage 
and  civilised  races,  of  ancient  and  modern  people,  and  by  the 
analogy  of  the  whole  vertebrate  series.  Dr.  J.  Barnard  Davis  has 
proved,'^^  by  many  careful  measurements,  that  the  mean  internal 
capacity  of  the  skull  in  Europeans  is  92-3  cubic  inches;  in 
Americans  87-5;  in  Asiatics  871 ;  and  in  Australians  only  81-9  - 
cubic  inches.   Professor  Broca  ^^  found  that  the  nineteenth  century 

^^  'Die    Grenzen    der  Thierwelt,  vomitoria*    1870,    p.    14.     Mv   son, 

oine  Betrachtung  zu  Darwin's  Lehre,'  Mr.  F.  Darwin,  dissected  for  ine  the 

1868,  s.  51.  cerebral    ganglia    of    the    Formica 

^^  Dujardin,     '  Annales    des     Sc.  rttfa. 

Nat,'  3rd    series   Zoolog.   torn,   xiv.  ''^    <  Philosophical     Transactions, 

1850,  p.  203.     See  also  Mr.  Lowne,  1869,  p.  513. 

'Anatomy  and  Phys.  of  the  3Imca  8<>  '  Les  Selections,'  M.  P.  Broca, 


Cji A  P.  I  [ .  Manner  of  Development,  5  5 

skulls  from  graves  in  Paris  were  larger  than  those  from  -vaults 
of  the  twelfth  century,  in  the  proportion  of  1-184  to  1426 ;  and 
that  the  increased  size,  as  ascertained  by  measurements,  was 
exclusively  in  the  frontal  part  of  the  skull — the  seat  of  the 
intellectual  faculties.  Prichard  is  persuaded  that  the  present 
inhabitants  of  Britain  have  "  much  more  capacious  brain-cases  " 
than  the  ancient  inhabitants.  Nevertheless,  it  must  be  admitted 
that  some  skulls  of  very  high  antiquity,  such  as  the  famous  one 
of  Neanderthal,  are  well  developed  and  capacious.^^  With 
respect  to  the  lower  animals,  M.  E.  Lartet,^-  by  comparing  the 
crania  of  tertiary  and  recent  mammals  belonging  to  the  same 
groups,  has  come  to  the  remarkable  conclusion  that  the  brain  is 
generally  larger  and  the  convolutions  are  more  complex  in  the 
more  recent  forms.  On  the  other  hand,  I  have  shewn^^  that  the 
brains  of  domestic  rabbits  are  considerably  reduced  in  bulk,  in 
comparison  with  those  of  the  wild  rabbit  or  hare ;  and  this  may  be 
attributed  to  their  having  been  closely  confined  during  many 
generations,  so  that  they  have  exerted  their  intellect,  instincts, 
senses  and  voluntary  movements  but  little. 

The  gi'adually  increasing  weight  of  the  brain  and  skull  in 
man  must  have  influenced  the  development  of  the  supporting 
spinal  column,  more  especially  whilst  he  was  becoming  erect. 
As  this  change  of  iDOsition  was  being  brought  about,  the  internal 
pressure  of  the  brain  will  also  have  influenced  the  form  of  the 
skull ;  for  many  facts  show  how  easily  the  skull  is  thus  affected. 
Ethnologists  believe  that  it  is  modified  by  the  kind  of  cradle  in 
which  infants  sleep.  Habitual  spasms  of  the  muscles,  and  a 
cicatrix  from  a  severe  burn,  have  permanently  modified  the  facial 
bones.  In  young  persons  whose  heads  have  become  fixed  either 
sideways  or  backwards,  o\ving  to  disease,  one  of  the  two  eyes  has 
changed  its  position,  and  the  shape  of  the  skull  has  been  altered 


'  Revue      d'Anthropologies,'     1873  ;  the  other  hand,  with  savages,  the  ave- 

see    also,    as   quoted    in    C.    Vogt's  rage  includes  only  the  more  capable 

'  Lectures    on    Man,'  Eng.   translat.  individuals,  who  have   been  able  to 

1864,  pp.  88,  90.     Prichard,  '  Phys.  survive  under   extremely  hard  con- 

Hist.  of  Mankind,'  vol.  i.   1838,  p.  ditions  of  lite.     Broca  thus  explains 

305,  the  otherwise  inexplicable  fact,  that 

*'  In  the  interecting  article  just  the  mean  capacity  of  the  skull  of 

referred    to.   Prof.    Broca    has  well  the  ancient  Troglodytes  of  Lozere  is 

remarked,  that  in  civilised  nations,  greater  than  that  ot  modern  French- 

the  average   capacity  of  the   skull  men. 

must  be  lowered   by  the   preserva-  "-   '  Comptes-rendus  des  Sciences,' 

tion    of  a    considerable    number    of  (Sec.  June  1,  18G8. 

individuals,  weak  in  mind  aud  body,  ^^  'The  Variation  of  Animals  and 

who    would    have    been    promptly  Plants  under  Domestication,'  vol.  i. 

eliminated  in  the  savage  state.     On  pp.  124-129. 


56  The  Descent  of  Man.  Part  I. 

apparently  by  the  pressure  of  the  brain  in  a  new  direction.^* 
I  have  shewn  that  with  long-eared  rabbits  even  so  trifling  a  cause 
as  the  lopping  forward  of  one  ear  drags  forward  almost  every 
bone  of  the  skull  on  that  side ;  so  that  the  bones  on  the  opposite 
side  no  longer  strictly  correspond.  Lastly,  if  any  animal  were 
to  increase  or  diminish  much  in  general  size,  without  any  change 
in  its  mental  powers,  or  if  the  mental  powers  were  to  be  much 
increased  or  diminished,  without  any  great  change  in  the  size  of 
the  body,  the  shape  of  the  skull  would  almost  certainly  be 
altered.  I  infer  this  from  my  observations  on  domestic  rabbits, 
some  kinds  of  which  have  become  very  much  larger  than  the 
wild  animal,  whilst  others  have  retained  nearly  the  same  size, 
but  in  both  cases  the  brain  has  been  much  reduced  relatively  to 
the  size  of  the  body.  Now  I  was  at  first  much  sui'prised  on 
finding  that  in  all  these  rabbits  the  skull  had  become  elongated 
or  dolichocephalic;  for  instance,  of  two  skulls  of  nearly  equal 
breadth,  the  one  from  a  wild  rabbit  and  the  other  from  a  large 
domestic  kind,  the  former  was  3-15  and  the  latter  4-3  inches  in 
length.^5  One  of  the  most  marked  distinctions  in  difierent  races 
of  men  is  that  the  skull  in  some  is  elongated,  and  in  others 
rounded;  and  here  the  explanation  suggested  by  the  case  of  the 
rabbits  may  hold  good;  for  Welcker  finds  that  short  "men  incline 
more  "  to  brachycephaly,  and  tall  men  to  dolichocephaly  ;"^^  and 
tall  men  may  be  compared  with  the  larger  and  longer-bodied 
rabbits,  all  of  which  have  elongated  skulls,  or  are  dolicho- 
cephalic. 

From  these  several  facts  we  can  understand,  to  a  certain 
extent,  the  means  by  which  the  great  size  and  more  or  less 
rounded  form  of  the  skull  have  been  acquired  by  man  ;  and  these 
are  characters  eminently  distinctive  of  him  in  comparison  with 
the  lower  animals. 

Another  most  conspicuous  difference  between  man  and  the 
lower  animals  is  the  nakedness  of  his  skin.  Whales  and 
porpoises  (Cetacea),  dugongs  (Sirenia)  and  the  hippopotamus  are 
naked;    and   this  may  be  advantageous  to  them  for  gliding 

**  SchaafFhausen  gives  fi-om  Blu-  maker,  where  the  head  is  habitually 

menbach  and  Busch,  the  cases  of  the  held  forward,  the  forehead  becomes 

spasms    and    cicatrix,    in    'Anthro-  more  rounded  and  })rominent. 

polog.    Review,'  Oct.   1868,  p.  420.  ^^  'Variation    of    Animals,'    &c., 

Dr.  Jarrold  (' Anthropologia,'  1808,  vol.  i.  p.   117,  on  the  elongation  of 

pp.  115,  116)  adduces  from  Camper  the  skull;  p.  119,  on  the  efi'ect  of 

and  from  his  own  observations,  cases  the  lopping  of  one  ear. 

of  the  modification  of  the  skull  from  ^"^  Quoted    by    Schaaffhausen,    in 

the  head  being  fixed  in  an  unnatural  '  Anthropolog.    Review,'  Oct.    18G8, 

position.     He  believes  that  in   cer-  p.  419. 
t.iin  trades,  such  as  that  of  a  shoe- 


Chap.  II.  Maimer  of  Development.  57 

through  the  water ;  nor  would  it  be  injurious  to  them  from  the 
loss  of  warmth,  as  the  species,  which  inhabit  the  colder  regions, 
are  protected  by  a  thick  layer  of  blubber,  serving  the  same 
purpose  as  the  fur  of  seals  and  otters.  Elephants  and  rhino- 
ceroses are  almost  hairless ;  and  as  certain  extinct  species, 
which  formerly  lived  under  an  Arctic  climate,  were  covered  with 
long  wool  or  hair,  it  would  almost  appear  as  if  the  existing 
species  of  both  genera  had  lost  their  hairy  covering  from  exposure 
to  heat.  This  appears  the  more  probable,  as  the  elephants  in 
India  which  live  "on  elevated  and  cool  districts  are  more  hairy  ^^ 
than  those  on  the  lowlands.  May  we  then  infer  that  man 
became  divested  of  hair  from  having  aboriginally  inhabited  some 
tropical  land  ?  That  the  hair  is  chiefly  retained  in  the  male  sex  on 
the  chest  and  face,  and  in  both  sexes  at  the  junction  of  all  four 
limbs  with  the  trunk,  favours  this  inference — on  the  assumption 
that  the  hair  was  lost  before  man  became  erect ;  for  the  parts 
which  now  retain  most  hair  would  then  have  been  most  protected 
from  the  heat  of  the  sun.  The  crown  of  the  head,  however, 
offers  a  curious  exception,  for  at  all  times  it  must  have  been  one 
of  the  most  exposed  parts,  yet  it  is  thickly  clothed  with  hair. 
The  fact,  however,  that  the  other  members  of  the  order  of 
Primates,  to  which  man  belongs,  although  inhabiting  various  hot 
regions,  are  well  clothed  with  hair,  generally  thickest  on  the  upper 
surface,*^  is  o^Dposed  to  the  supposition  that  man  became  naked 
through  the  action  of  the  sun.  Mr.  Belt  believes  ^^  that  within 
the  tropics  it  is  an  advantage  to  man  to  be  destitute  of  hair,  as 
he  is  thus  enabled  to  free  himself  of  the  multitude  of  ticks  (acari) 
and  other  jDarasites,  with  which  he  is  often  infested,  and  which 
sometimes  cause  ulceration.  But  whether  this  evil  is  of  sufficient 
magnitude  to  have  led  to  the  denudation  of  his  body  through 
natural  selection,  may  be  doubted,  since  none  of  the  many 
quadrupeds  inhabiting  the  tropics  have,  as  far  as  I  know, 
acquired  any  specialised  means  of  relief.  The  view  which  seems 
to  me  the  most  probable  is  that  man,  or  rather  primarily  woman, 

*^    Osven,    'Anatomy    of    Yerte-  ever,  states  that  in  the  Gorilla  the 

brates,'  vol.  iii.  p.  619.  hair  is  thinner  on  the  back,  where 

**  Isidore  Geotfroy  St.-Hilaire  re-  it  is  partly  rubbed  oil",  than  on  the 

marks  ('  Hist.  Nat.  Generale,'  torn,  lower  surface. 

ii.  1859,  pp.  215-217)  on  the  head  of  ^^  The  '  Naturalist  in  Nicaragua,' 

man  being  covered  with  long  hair;  1874,  p.  209.     As   some    confirma- 

also  on  the  upper  surfaces  of  mon-  tion  of  Mr.  Belt's  view,  I  may  quote 

keys  and  of  other  mammals  being  the   following  passage  from  Sir  VV. 

more  thickly  clothed  than  the  lower  Denison    ('  Varieties    of  Vice-Kegal 

surfaces.      This    has    likewise    been  Life,' vol.  i.  1870,  p.  440):  "  It  is  said 

observed  by  various  authors.     Prof.  "  to   be   a  })ractice  with    the  Aus- 

P.  Gervais  ('  Hist.  Nat.   des  Mam-  "  tralians,    when    the    vermin    get 

mifbres,'  torn.  i.  1854,  p.  28),  how-  "  troublesome,  to  twinge  themselves." 
4 


58  The  Descent  of  Man.  Part  I. 

became  divested  of  hair  for  omamenial  purposes,  as  we  sliall  see 
under  Sexual  Selection ;  and,  according  to  this  belief,  it  is  not 
surprising  that  man  should  diifer  so  greatly  in  hairiness  from  all 
other  Primates,  for  characters,  gained  through  sexual  selection, 
often  differ  to  an  extraordinary  degree  in  closely-related  forms. 

According  to  a  popular  impression,  the  absence  of  a  tail  is 
eminently  distinctive  of  man ;  but  as  those  apes  which  come 
nearest  to  him  are  destitute  of  this  organ,  its  disapi^earance  does 
not  relate  exclusively  to  man.  The  tail  often  differs  remarkably 
in  length  within  the  same  genus :  thus  in  some  species  of  Macacus 
it  is  longer  than  the  whole  body,  and  is  formed  of  twenty-four 
vertebrre;  in  others  it  consists  of  a  scarcely  visible  stumj^, 
containing  only  three  or  four  vertebi-se.  In  some  kinds  of 
baboons  there  are  twenty-five,  whilst  in  the  mandrill  there  are 
ten  very  small  stunted  caudal  vertebrae,  or,  according  to  Cuvier,®° 
sometimes  only  five.  The  tail,  whether  it  be  long  or  short,  almost 
always  tapers  towards  the  end ;  and  this,  I  presume,  results  from 
the  atrophy  of  the  terminal  muscles,  together  with  their  arteries 
and  nerves,  through  disuse,  leading  to  the  atrophy  of  the  terminal 
bones.  But  no  explanation  can  at  present  be  given  of  the  great 
diversity  which  often  occurs  in  its  length.  Here,  however,  we 
are  more  specially  concerned  with  the  complete  external  dis- 
appearance of  the  tail.  Professor  Broca  has  recently  shewn  ^^ 
that  the  tail  in  all  quadrupeds  consists  of  two  portions,  generally 
separated  abruptly  from  each  other ;  the  basal  portion  consists 
of  vertebras,  more  or  less  perfectly  channelled  and  furnished  with 
apophyses  like  ordinary  vertebrae  ;  whereas  those  of  the  terminal 
portion  are  not  channelled,  are  almost  smooth,  and  scarcely 
resemble  true  vertebrae,  A  tail,  though  not  externally  visible,  is 
really  present  in  man  and  the  anthropomorphous  apes,  and  is 
constructed  on  exactly  the  same  pattern  in  both.  In  the  terminal 
l^ortion  the  vertebrae,  constituting  the  os  coccyx,  are  quite 
rudimentary,  being  much  reduced  in  size  and  nimiber.  In  the 
basal  portion,  the  vertebrae  are  likewise  few,  are  united  firmly 
together,  and  are  arrested  in  development;  but  they  have  been 
rendered  much  broader  and  flatter  than  the  corresponding 
vertebrae  in  the  tails  of  other  animals :  they  constitute  what 
Broca  calls  the  accessory  sacral  vertebrte.  These  are  of  functional 
importance  by  supporting  certain  internal  parts  and  in  other 
ways ;  and  theii'  modification  is  directly  connected  with  the  erect 

^^  Ml ,  St.  George  Mivart,  '  Proc.  Geoffroy,  '  Hist.  Xat.  Gen.'  torn.  ii. 

Zoolog.    Soc'   1865,   pp.    562,    583.  p.  244. 

Dr.  J.E.  Gray,  'Cat.   Brit.   Mus.:  ^^  'Revue  d'Anthropologie,' 1872  ; 

Skeletons.'      Owen,     '  Anatomy     of  '  La  Constitution  des  Yertebres  cau- 

Vertebrates,'  vol.  ii.  p.  517.     Isidore  dales.' 


Chap.  II.  Manner  of  Development.  59 

or  semi-erect  attitude  of  man  and  the  anthropomorphous  apes. 
This  conclusion  is  the  more  trustworthy,  as  Broca  formerly  held 
a  different  view,  which  he  has  now  abandoned.  The  modifica- 
tion, therefore,  of  the  basal  caudal  vertebrae  in  man  and  the 
higher  apes  may  have  been  effected,  directly  or  indirectly, 
through  natural  selection. 

But  what  arc  we  to  say  about  the  rudimentary  and  variable 
vertebrae  of  the  terminal  portion  of  the  tail,  forming  the  os  coccyx  ? 
A  notion  which  has  often  been,  and  will  no  doubt  again  be 
ridiculed,  namely,  that  friction  has  had  something  to  do  with 
the  disappearance  of  the  external  portion  of  the  tail,  is  not 
so  ridiculous  as  it  at  first  appears.  Dr.  Anderson '-"^  states 
that  the  extremely  short  tail  of  Macacus  hrnnneus  is  formed  of 
eleven  vertebrae,  including  the  imbedded  basal  ones.  The 
extremity  is  tendinous  and  contains  no  vertebrae ;  this  is  suc- 
ceeded by  five  rudimentary  ones,  so  minute  that  together  they 
are  only  one  line  and  a  half  in  length,  and  these  are  permanently 
bent  to  one  side  in  the  shape  of  a  hook.  The  free  part  of  the 
tail,  only  a  little  above  an  inch  in  length,  includes  only  four  more 
small  vertebrae.  This  short  tail  is  carried  erect;  but  about  a 
quarter  of  its  total  length  is  doubled  on  to  itself  to  the  left ;  and 
tliis  terminal  part,  which  includes  the  hook-like  portion,  serves 
"  to  fill  up  the  interspace  between  the  upper  divergent  portion 
"  of  the  callosities ;"  so  that  the  animal  sits  on  it,  and  thus  renders 
it  rough  and  callous.  Dr.  Anderson  thus  sums  up  his  observa- 
tions :  "  These  facts  seem  to  me  to  have  only  one  explanation ; 
"  this  tail,  from  its  short  size,  is  in  the  monkey's  way  when  it 
"  sits  down,  and  frequently  becomes  placed  under  the  animal 
*'  while  it  is  in  this  attitude  ;  and  from  the  circumstance  that  it 
"  does  not  extend  beyond  the  extremity  of  the  ischial  tuberosities 
"  it  seems  as  if  the  tail  originally  had  been  bent  round,  by  the 
"  will  of  the  animal,  into  the  interspace  between  the  callosities,  to 
"  escape  being  pressed  between  them  and  the  ground,  and  that 
"  in  time  the  curvature  became  permanent,  fitting  in  of  itself 
"  when  the  organ  happens  to  be  sat  upon."  Under  these  circum- 
stances it  is  not  surprising  that  the  surface  of  the  tail  should 
liave  been  roughened  and  rendered  callous;  and  Dr.  Murie,^^  who 
carefully  observed  this  species  in  the  Zoological  Gardens,  as  well 
as  three  other  closely  allied  forms  with  slightly  longer  tails,  says 
that  when  the  animal  sits  down,  the  tail  "  is  necessarily  thrust 
"  to  one  side  of  the  buttocks ;  and  whether  long  or  short  its  root 
"  is  consequently  liable  to  be  rubbed  or  chafed."    As  we  now 

»2  'Proc.  Zoolog.  Soc.,'  1872,   p.  "  *  Proc.    Zoolog.    Sue.    1872,    p. 

210.  786. 


6o  The  Descent  of  Man.  Part  I 

have  evidence  that  mutilations  occasionally  produce  an  inherited 
effect/*  it  is  not  very  improbable  that  in  short-tailed  monkeys,  the 
projecting  part. of  the  tail,  being  functionally  useless,  should  after 
many  generations  have  become  rudimentary  and  distorted,  from 
being  continually  rubbed  and  chafed.  We  see  the  projecting  part  in 
this  condition  in  the  Macacushruimeus,  and.  absolutely  aborted  in 
the  if.  ecaudatus  and  in  several  of  the  higher  apes.  Finally,  then, 
as  far  as  we  can  judge,  the  tail  has  disappeared  in  man  and  the 
anthropomorphous  apes,  owing  to  the  terminal  portion  having 
been  injured  by  friction  during  a  long  lapse  of  time ;  the  basal 
and  embedded  portion  having  been  reduced  and  modified,  so  as 
to  become  suitable  to  the  erect  or  semi-erect  position. 

I  have  now  endeavoured  to  shew  that  some  of  the  most 
distinctive  characters  of  man  have  in  all  probabiHty  been 
acquii-ed,  either  directly,  or  more  commonly  indirectly,  through 
natural  selection.  We  should  bear  in  mind  that  modifications 
in  structure  or  constitution,  which  do  not  serve  to  adapt  an 
organism  to  its  habits  of  life,  to  the  food  which  it  consumes,  or 
passively  to  the  surrounding  conditions,  cannot  have  been  thus 
acquired.  We  must  not,  however,  be  too  confident  in  decidnig 
what  modifications  are  of  service  to  each  being:  we  should 
remember  ho^^  little  we  know  about  the  use  of  many  parts,  or 
what  changes  in  the  blood  or  tissues  may  serve  to  fit  an 
organism  for  a  new  climate  or  new  kinds  of  food.  Nor  must  we 
forget  the  principle  of  con-elation,  by  which,  as  Isidore  Geoffroy 
has  shewn  in  the  case  of  man,  many  strange  deviations  of 
structure  are  tied  together.  Independently  of  correlation,  a 
change  in  one  part  often  leads,  through  the  increased  or  decreased 
use  of  other  parts,  to  other  changes  of  a  quite  unexpected 
nature.  It  is  also  well  to  reflect  on  such  facts,  as  the  wonderful 
growth  of  galls  on  plants  caused  by  the  poison  of  an  insect,  and 
on  the  remarkable  changes  of  colour  in  the  plumage  of  parrots 
when  fed  on  certain  fishes,  or  inoculated  with  the  poison  of 
toads  ;^^  for  we  can  thus  see  that  the  fluids  of  the  system,  if 
altered  for  some  special  purpose,  might  induce  other  changes. 
We  should  especially  bear  in  mind  that  modifications  acquired 

5*  I  allude  to  Dr.  Brown-Sequard's  inherited  effects  of  mot-mots  biting 
observations  on  the  transmitted  off  the  barbs  of  their  own  tail- 
effect  of  an  operation  causing  epi-  feathers.  See  jlso  on  the  general 
lepsy  in  guinea-pigs,  and  likewise  subject  '  Variation  of  Animals  and 
more  recently  on  the  analogous  Plants  under  Domestication,'  vol. 
effects    of  cutting  the  sympathetic  ii.,  pp.  22-24. 

nerve  in  the  neck.    I  shall  hereafter  ^^  '  The  Variation  of  Animals  and 

have  occasion  to  refer  to  Mr.  Salvin's  Plants  under  Domestication/  vol.  ii. 

interesting   case  of  the  apparently  pp.  280,  282. 


CiiAi'.  II.  Manner  of  Development.  61 

and  continually  used  during  past  ages  for  some  useful  purpose, 
would  probably  become  firmly  fixed,  and  might  be  long  inherited. 

Thus  a  large  yet  undefined  extension  may  safely  be  given  to 
the  direct  and  indirect  results  of  natural  selection ;  but  I  now 
admit,  after  reading  the  essay  by  NUgeli  on  plants,  and  the 
remarks  by  yarious  authors  with  respect  to  animals,  more 
especially  those  recently  made  by  Professor  Broca,  that  in  the 
earlier  editions  of  my  *  Origin  of  Species '  I  perhaps  attributed 
too  much  to  the  action  of  natural  selection  or  the  survival  of 
the  fittest.  I  have  altered  the  fifth  edition  of  the  '  Origin '  so  as  to 
confine  my  remarks  to  adaptive  changes  of  structure ;  but  I  am 
convinced,  from  the  light  gained  during  even  the  last  few  years, 
that  very  many  structures  which  now  appear  to  us  useless,  will 
hereafter  be  proved  to  be  useful,  and  will  therefore  come  within 
the  range  of  natui'al  selection.  Nevertheless,  I  did  not  formerly 
consider  sufficiently  the  existence  of  structures,  which,  as  far  as 
we  can  at  present  judge,  are  neither  beneficial  nor  injurious ; 
and  this  I  believe  to  be  one  of  the  greatest  oversights  as  yet 
detected  in  my  work.  I  may  be  permitted  to  say,  as  some 
excuse,  that  I  had  two  distinct  objects  in  view;  firstly,  to 
shew  that  species  had  not  been  separately  created,  and  secondly, 
that  natural  selection  had  been  the  chief  agent  of  change, 
though  largely  aided  by  the  inherited  effects  of  habit,  and  slightly 
by  the  dii-ect  action  of  the  surrounding  conditions.  I  was 
not,  however,  able  to  annul  the  influence  of  my  former  belief, 
then  almost  universal,  that  each  species  had  been  purposely 
created ;  and  this  led  to  my  tacit  assumption  that  every  detail 
of  structure,  excepting  rudiments,  was  of  some  special,  though 
unrecognised,  service.  Any  one  with  this  assimiption  in  his 
mind  would  naturally  extend  too  far  the  action  of  natural 
selection,  either  during  past  or  present  times.  Some  of  those 
who  admit  the  principle  of  evolution,  but  reject  natural  selec- 
tion, seem  to  forget,  when  criticising  my  book,  that  I  had  the 
above  two  objects  in  view ;  hence  if  I  have  erred  in  giving  to 
natural  selection  great  power,  which  I  am  very  far  from 
admitting,  or  in  having  exaggerated  its  power,  which  is  in  itself 
probable,  I  have  at  least,  as  I  hope,  done  good  service  in  aiding 
to  overthrow  the  dogma  of  separate  creations. 

It  is,  as  I  can  now  see,  probable  that  all  organic  beings, 
including  man,  possess  peculiarities  of  structure,  which  neither 
are  now,  nor  were  formerly  of  any  service  to  them,  and  which, 
therefore,  are  of  no  physiological  importance.  Wo  know  not 
what  produces  the  numberless  slight  difierenccs  between  the 
individuals  of  each  species,  for  reversion  only  carries  the 
problem  a  few  steps  backwards;   but  each  peculiarity  must 


62  The  Descent  of  Man.  Taut  I. 

have  had  its  efficient  cause.  If  these  causes,  whatever  they 
may  be,  were  to  act  more  -uniformly  and  energetically  during  a 
lengthened  period  (and  against  this  no  reason  can  be  assigned), 
the  result  w^ould  probably  be  not  a  mere  slight  individual 
difference,  but  a  well-marked  and  constant  modification,  though 
one  of  no  physiological  importance.  Chauged  structures,  which 
are  in  no  w^ay  beneficial,  cannot  be  kept  uniform  through  natural 
selection,  though  the  injurious  will  be  thus  eliminated.  Uni- 
formity of  character  would,  however,  naturally  follow  from  the 
assumed  nniformity  of  the  exciting  causes,  and  likewise  from 
the  free  intercrossing  of  many  individuals.  During  successive 
periods,  the  same  organism  might  in  this  manner  acquire 
successive  modifications,  w^hich  would  be  transmitted  in  a  nearly 
uniform  state  as  long  as  the  exciting  causes  remained  the  same 
and  there  w^as  free  intercrossing.  With  respect  to  the  exciting 
causes  we  can  only  say,  as  when  speaking  of  so-called  spon- 
taneous variations,  that  they  relate  much  more  closely  to  the 
constitution  of  the  varying  organism,  than  to  the  nature  of  the 
conditions  to  which  it  has  been  subjected. 

Conclusion.— In  this  chapter  we  have  seen  that  as  man  at  the 
present  day  is  liable,  like  every  other  animal,  to  multiform 
individual  differences  or  slight  variations,  so  no  doubt  were  the 
early  progenitors  of  man ;  the  variations  being  formerly  induced 
by  the  same  general  causes,  and  governed  by  the  same  general 
and  complex  laws  as  at  present.  As  all  animals  tend  to  multiply 
beyond  their  means  of  subsistence,  so  it  must  have  been  with 
the  progenitors  of  man;  and  this  would  inevitably  lead  to  a 
struggle  for  existence  and  to  natural  selection.  The  latter 
process  would  be  greatly  aided  by  the  inherited  effects  of  the 
increased  use  of  parts,  and  these  two  processes  would  incessantly 
react  on  each  other.  It  appears,  also,  as  we  shall  hereafter  see, 
that  various  unimportant  characters  have  been  acquired  by  man 
through  sexual  selection.  An  unexplained  residuum  of  change 
must  be  left  to  the  assumed  uniform  action  of  those  unknown 
agencies,  which  occasionally  induce  strongly  marked  and  abrupt 
deviations  of  structure  in  our  domestic  productions. 

Judging  from  the  habits  of  savages  and  of  the  greater  number 
of  the  Quadrumana,  primeval  men,  and  even  their  ape-hke 
progenitors,  probably  lived  in  society.  With  strictly  social 
animals,  natural  selection  sometimes  acts  on  the  individual, 
through  the  preservation  of  variations  which  are  beneficial  to 
the  community.  A  community  which  includes  a  large  number 
of  well-endowed  individuals  increases  in  number,  and  is  victo- 
rious over  other  less  favoured  ones ;  even  although  each  separate 


Cjiai'.  II.  Manner  of  Devdopmcjit.  6^ 

member  gains  no  advantage  over  the  others  of  the  same  com- 
munity. Associated  insects  have  thus  acquired  many  remark- 
able structures^  M'hich  are  of  little  or  no  service  to  the  individual, 
such  as  the  pollen-collecting  apparatus,  or  tlie  sting  of  the 
worker-bee,  or  the  great  jaws  of  soldier-ants.  With  the  liigher 
social  animals,  I  am  not  aware  that  any  structure  has  been 
modified  solely  for  the  good  of  the  community,  though  some  are 
of  secondary  service  to  it.  For  instance,  the  horns  of  ruminants 
and  the  great  canine  teeth  of  baboons  appear  to  liave  been 
acquired  by  the  males  as  weapons  for  sexual  strife,  but  they  are 
used  in  defence  of  the  herd  or  troop.  In  regard  to  certain 
mental  powers  the  case,  as  we  shall  see  in  the  fifth  chapter,  is 
wholly  dificrent ;  for  these  faculties  have  been  chiefly,  or  even 
exclusively,  gained  for  the  benefit  of  the  community,  and  the 
individuals  thereof,  have  at  the  same  time  gained  an  advantage 
indirectly. 

It  has  often  been  objected  to  such  views  as  the  foregoing,  that 
man  is  one  of  the  most  helpless  and  defenceless  creatures  in  the 
uorld;  and  that  during  his  early  and  less  well-developed 
condition  he  would  have  been  still  more  helpless.  The  Duke  of 
Argyll,  for  instance,  insists ^*^  that  "the  human  frame  has 
"  diverged  from  the  structure  of  brutes,  in  the  direction  of 
"  greater  physical  helplessness  and  weakness.  That  is  to  say,  it 
"  is  a  divergence  which  of  all  others  it  is  most  impossible  to 
"  ascribe  to  mere  natural  selection."  He  adduces  the  naked  and 
unprotected  state  of  the  body,  the  absence  of  great  teeth  or 
claws  for  defence,  the  small  strength  and  speed  of  man,  and  his 
shght  power  of  discovering  food  or  of  avoiding  danger  by  smell. 
To  these  deficiencies  there  might  be  added  one  still  more 
serious,  namely,  that  he  cannot  climb  quickly,  and  so  escape 
from  enemies.  The  loss  of  hair  would  not  have  been  a  great 
injury  to  the  inhabitants  of  a  warm  country.  For  we  know  that  the 
unclothed  Fuegians  can  exist  under  a  wretched  climate.  When 
we  compare  the  defenceless  state  of  man  with  that  of  apes,  we 
must  remember  that  the  great  canine  teeth  with  which  the  latter 
are  provided,  are  possessed  in  their  full  development  by  the  males 
alone,  and  are  chiefly  used  by  them  for  fighting  with  their  rivals ; 
yet  the  females,  which  are  not  thus  provided,  manage  to  survive. 

In  regard  to  bodily  size  or  strength,  we  do  not  know  whether 
man  is  descended  from  some  small  species,  hke  the  chimpanzee, 
or  from  one  as  powerful  as  the  gorilla;  and,  therefore,  we  cannot 
say  whether  man  has  become  lai-ger  and  stronger,  or  smaller 

»«  '  Primeval  Man,'  1869,  p.  ^<o. 


64  The  Descent  of  Man.  Takt  I. 

and  weaker,  than  his  ancestors.  We  should,  however,  bear  in 
mind  that  an  animal  possessing  great  size,  strength,  and  ferocity, 
and  which,  like  the  gorilla,  could  defend  itself  from  all  enemies, 
would  not  perhajjs  have  become  social;  and  this  would  most 
effectually  have  checked  the  acquirement  of  "the  higher  mental 
qualities,  such  as  sympathy  and  the  love  of  his  fellows.  Hence  it 
might  have  been  an  immense  advantage  to  man  to  have  s^^rung 
from  some  comparatively  weak  creature. 

The  small  strength  and  speed  of  man,  his  want  of  natural 
weapons,  &c.,  are  more  than  counterbalanced,  firstly,  by  his 
intellectual  powers,  through  which  he  has  formed  for  himself 
weapons,  tools,  &c.,  though  still  remaining  in  a  barbarous  state, 
and,  secondly,  by  his  social  qualities  which  lead  him  to  give  and 
receive  aid  from  his  fellow-men.  No  country  in  the  world  abounds 
in  a  gi-eater  degree  with  dangerous  beasts  than  Southern  Africa ; 
no  country  presents  more  fearful  physical  hardships  than  the 
Arctic  regions;  yet  one  of  the  puniest  of  races,  that  of  the 
Bushmen,  maintains  itself  in  Southern  Africa,  as  do  the  dwarfed 
Esquimaux  in  the  Arctic  regions.  The  ancestors  of  man  were,  no 
doubt,  inferior  in  intellect,  and  probably  in  social  disposition,  to 
the  lowest  existing  savages ;  but  it  is  quite  conceivable  that  they 
might  have  existed,  or  even  flourished,  if  they  had  advanced  in 
intellect,  whilst  gradually  losing  their  brute-like  powers,  such 
as  that  of  climbing  trees,  &c.  But  these  ancestors  would  not 
have  been  exposed  to  any  special  danger,  even  if  far  more 
helpless  and  defenceless  than  any  existing  savages,  had  they 
inhabited  some  warm  continent  or  large  island,  such  as 
Austraha,  New  Guinea,  or  Borneo,  which  is  now  the  home  of  the 
orang.  And  natural  selection  arising  from  the  competition  of 
tribe  with  tribe,  in  some  such  large  area  as  one  of  these,  together 
with  the  inherited  effects  of  habit,  would,  under  favourable 
conditions,  have  sufficed  to  raise  man  to  his  present  high  position 
in  the  organic  scale. 


Chap.  III.  Mental  Poivers.  65 


CHAPTEE  III. 

Comparison  of  the  Mental  Powers  of  Man  and  the 
Lower  Animals. 

The  difference  in  mental  power  between  the  highest  ape  and  the  lowest 
savage,  immense  —  Certain  instincts  in  common  —  The  emotions  — 
Curiosity — Imitation — Attention  —  Memory  —  Imagination  —  Reason — 
Progressive  improvement  —  Tools  and  weapons  used  by  animals  — 
Abstraction,  self-consciousness — Language — Sense  of  beauty — Belief  in 
God,  spiritual  agencies,  superstitions. 

We  have  seen  in  the  last  two  chapters  that  man  bears  in  his 
bodily  structure  clear  traces  of  his  descent  from  some  lower 
form ;  but  it  may  be  urged  that,  as  man  differs  so  greatly  in 
his  mental  power  from  all  other  animals,  there  must  be  some 
error  in  this  conclusion.  No  doubt  the  difference  in  this 
respect  is  enormous,  even  if  we  compare  the  mind  of  one  of  the 
lowest  savages,  who  has  no  words  to  express  any  number  higher 
than  four,  and  who  uses  hardly  any  abstract  terms  for  common 
objects  or  for  the  affections,^  with  that  of  the  most  highly 
organised  ape.  The  difference  would,  no  doubt,  still  remain 
immense,  even  if  one  of  the  higher  apes  had  been  improved  or 
civilised  as  much  as  a  dog  has  been  in  comparison  with  itt> 
parent-form,  the  wolf  or  jackal.  The  Fuegians  rank  amongst 
the  lowest  barbarians;  but  I  was  continually  struck  with 
surprise  how  closely  the  three  natives  on  board  H.M.S. ''  Beagle," 
who  had  lived  some  years  in  England,  and  could  talk  a  little 
English,  resembled  us  in  disposition  and  in  most  of  our  mental 
faculties.  If  no  organic  being  excepting  man  had  possessed  any 
mental  power,  or  if  his  powers  had  been  of  a  wholly  different 
nature  from  those  of  the  lower  animals,  then  we  should  never 
have  been  able  to  convince  ourselves  that  our  high  faculties 
had  been  gradually  developed.  But  it  can  be  shewn  that  there 
is  no  fundamental  difference  of  this  kind.  We  must  also  admit 
that  there  is  a  much  wider  interval  in  mental  power  between 
one  of  the  lowest  fishes,  as  a  lamprey  or  lancelet,  and  one  of  the 
higher  apes,  than  between  an  ai)e  and  man ;  yet  this  interval 
is  filled  up  by  numberless  gradations. 

Nor  is  the  difference  slight  in  moral  disposition  between  a 
barbarian,   such  as  the  man  described  by  the  old  navigator 

*  See  the  evidence  on  those  points,  as  given  by  Lubboclc,  '  Prehistoric 
Times,'  p.  354,  &c. 


66  TJie  Descent  of  Man.  Part  I. 

Byron,  -who  dashed  his  child  on  the  rocks  for  dropping  a  basket 
of  sea-urchins,  and  a  Howard  or  Clarkson;  and  in  intellect, 
between  a  savage  who  uses  hardly  any  abstract  terms,  and  a 
Newton  or  Shakspeare.  Differences  of  this  kind  between  the 
highest  men  of  the  highest  races  and  the  lowest  savages,  are 
connected  by  the  finest  gradations.  Therefore  it  is  possible  that 
they  might  pass  and  be  developed  into  each  other. 

My  object  in  this  chapter  is  to  shew  that  there  is  no  funda- 
mental difference  between  man  and  the  higher  mammals  in  their 
mental  faculties.  Each  division  of  the  subject  might  have  been 
extended  into  a  separate  essay,  but  must  here  be  treated  briefly. 
As  no  classification  of  the  mental  i:>owers  has  been  universally 
accepted,  I  shall  arrange  my  remarks  in  the  order  most  con- 
venient for  my  purpose  ;  and  will  select  those  facts  which  have 
struck  me  most,  with  the  hope  that  they  may  produce  some 
effect  on  the  reader. 

With  respect  to  animals  very  low  in  the  scale,  I  shall  give 
some  additional  facts  under  Sexual  Selection,  shewing  that  their 
mental  powers  are  much  higher  than  might  have  been  expected. 
The  variability  of  the  faculties  in  the  individuals  of  the  same 
species  is  an  important  point  for  us,  and  some  few  illustrations 
will  here  be  given.  But  it  would  be  superfluous  to  enter  into 
many  details  on  this  head,  for  I  have  found  on  frequent  enquiry, 
that  it  is  the  unanimous  opinion  of  all  those  who  have  long 
attended  to  animals  of  many  kinds,  including  birds,  that  the 
individuals  difibr  greatly  in  every  mental  characteristic.  In 
what  manner  the  mental  powers  were  first  developed  in  the 
lowest  organisms,  is  as  hojoeless  an  enquiry  as  how  life  itself 
first  originated.  These  are  problems  for  the  distant  future,  if 
they  are  ever  to  be  solved  by  man. 

As  man  possesses  the  same  senses  as  the  lower  animals,  his 
fundamental  intuitions  must  be  the  same.  Man  has  also  some 
few  instincts  in  common,  as  that  of  self-preservation,  sexual  love, 
the  love  of  the  mother  for  her  new-born  offspring,  the  desire 
possessed  by  the  latter  to  suck,  and  so  forth.  But  man,  perhaps, 
has  somewhat  fewer  instincts  than  those  possessed  by  the 
animals  which  come  next  to  him  in  the  series.  The  orang  in 
the  Eastern  islands,  and  the  chimpanzee  in  Africa,  build  plat- 
forms on  which  they  sleep ;  and,  as  both  species  follow  the  same 
habit,  it  might  be  argued  that  this  was  due  to  instinct,  but  we 
cannot  feel  sure  that  it  is  not  the  result  of  both  animals  having 
similar  wants,  and  possessing  similar  powers  of  reasoning. 
These  apes,  as  we  may  assume,  avoid  the  many  poisonous  fruits 
of  the  tropics,  and  man  has  no  such  knowledge :  but  as  our 
domestic  animals,  when  taken  to  foreign  lands,  and  when  first 


Chap.  II  r.  Mental  Powers.  6y 

turned  out  in  the  spring,  often  eat  poisonous  lierbs,  which  they 
afterwards  avoid,  we  cannot  feel  sure  that  the  apes  do  not  learn 
from  their  own  experience  or  from  that  of  their  parents  what 
fruits  to  select.  It  is,  however,  certain,  as  we  shall  presently  see, 
that  apes  have  an  instinctive  dread  of  serpents,  and  probably  of 
other  dangerous  animals. 

The  fewness  and  the  comparative  simplicity  of  the  instincts  in 
the  higher  animals  are  remarkable  in  contrast  with  those  of  the 
lower  animals.  Cuvicr  maintained  that  instinct  and  intelligence 
stand  in  an  inverse  ratio  to  each  other  ;  and  some  have  thought 
that  the  intellectual  f\iculties  of  the  higher  animals  have  been 
gradually  developed  from  their  instincts.  But  Pouchet,  in  an 
interesting  essay,-  has  shewn  that  no  such  inverse  ratio  really 
exists.  Those  insects  which  possess  the  most  wonderful  instincts 
are  certainly  the  most  intelligent.  In  the  vertebrate  series,  the 
least  intelligent  members,  namely  fishes  and  amphibians,  do  not 
possess  complex  instincts;  and  amongst  mammals  the  animal 
most  remarkable  for  its  instincts,  namely  the  beaver,  is  highly 
intelligent,  as  will  be  admitted  by  every  one  who  has  read  Mr. 
Morgan's  excellent  work.^ 

Although  the  first  dawnings  of  intelligence,  according  to  jMr. 
Herbert  Spencer,*  have  been  developed  through  the  multiplica- 
tion and  co-ordination  of  reflex  actions,  and  although  many  of 
the  simpler  instincts  graduate  into  reflex  actions,  and  can  hardly 
be  distinguished  from  them,  as  in  the  cnse  of  young  animals 
sucking,  yet  the  more  complex  instincts  seem  to  have  originated 
independently  of  intelligence.  I  am,  however,  very  far  from 
wishing  to  deny  that  instinctive  actions  may  lose  their  fixed  and 
untaught  character,  and  be  replaced  by  others  performed  by  the 
aid  of  the  free  will.  On  the  other  hand,  some  intelligent  actions, 
after  being  performed  during  several  generations,  become  con- 
verted into  instincts  and  are  inherited,  as  when  birds  on  oceanic 
islands  learn  to  avoid  man.  These  actions  may  then  be  said 
to  be  degraded  in  character,  for  they  are  no  longer  performed 
through  reason  or  from  experience.  13ut  the  greater  number  of 
the  more  complex  instincts  appear  to  have  been  gained  in  a 
wholly  different  manner,  through  the  natural  selection  of  varia- 
tions of  simjilcr  instinctive  actions.  Such  variations  ai')pear  to 
arise  from  the  same  unknown  causes  acting  on  the  cerebral 
organisation,  which  induce  slight  variations  or  individual  dif- 
ferences in  other  parts  of  the  body ;  and  these  variations,  owing 

^  *  L'Instinct    chez    les    Inscctos.'  '  '  The  American  Beaver  and  his 

'  Revue  des  Deux  Mondes,'  Feb.  1870,      Works,'  1868. 

p.  G90.  ■*  '  The  Principles  of  Psychology, 

2nd  edit.  1870,  pp.  418-443. 


68  The  Descent  of  Man.  Part  I. 

to  our  ignorance,  are  often  said  to  arise  spontaneously.  We  can, 
I  think,  come  to  no  other  conclusion  with  respect  to  the  origin  of 
the  more  complex  instincts,  when  we  reflect  on  the  marvellous 
instincts  of  sterile  worker-ants  and  bees,  which  leave  no  off- 
spring to  inherit  the  effects  of  experience  and  of  modified  habits. 

Although,  as  we  learn  from  the  above-mentioned  insects  and 
the  beaver,  a  high  degree  of  intelligence  is  certainly  compatible 
with  complex  instincts,  and  although  actions,  at  first  learnt 
voluntarily  can  soon  through  habit  be  performed  with  the 
quickness  and  certainty  of  a  reflex  action,  yet  it  is  not  improbable 
that  there  is  a  certain  amount  of  interference  between  the 
development  of  free  intelligence  and  of  instinct, — which  latter 
imphes  some  inherited  modification  of  the  brain.  Little  is 
known  about  the  functions  of  the  brain,  but  we  can  perceive 
that  as  the  intellectual  powers  become  highly  developed,  the 
various  parts  of  the  brain  must  be  connected  by  very  intricate 
channels  of  the  freest  intercommunication;  and  as  a  conse- 
quence, each  separate  part  would  perhaps  tend  to  be  less  well  fitted 
to  answer  to  particular  sensations  or  associations  in  a  definite 
and  inherited— that  is  instinctive— manner.  There  seems  even 
to  exist  some  relation  between  a  low  degTce  of  intelligence  and  a 
strong  tendency  to  the  formation  of  fixed,  though  not  inherited 
habits ;  for  as  a  sagacious  physician  remarked  to  me,  persons 
who  are  slightly  imbecile  tend  to  act  in  everything  by  routine 
or  habit ;  and  they  are  rendered  much  happier  if  this  is  en- 
couraged. 

I  have  thought  this  digression  worth  giving,  because  we  may 
easily  underrate  the  mental  powers  of  the  higher  animals,  and 
especially  of  man,  when  we  compare  their  actions  founded  on  the 
memory  of  past  events,  on  foresight,  reason,  and  imagination, 
with  exactly  similar  actions  instinctively  performed  by  the  lower 
animals;  in  this  latter  case  the  capacity  of  performing  such 
actions  has  been  gained,  step  by  step,  through  the  variability  of 
the  mental  organs  and  natural  selection,  without  any  conscious 
inteUigenCe  on  the  part  of  the  animal  during  each  successive 
generation.  No  doubt,  as  Mr.  Wallace  has  argued,^  much  of  the 
intelligent  work  done  by  man  is  due  to  imitation  and  not  to 
reason;  but  there  is  this  great  difference  between  his  actions 
and  many  of  those  performed  by  the  lower  animals,  namely,  that 
man  cannot,  on  his  first  trial,  make,  for  instance,  a  stone  hatchet 
or  a  canoe,  through  his  power  of  imitation.  He  has  to  learn  his 
work  by  practice ;  a  beaver,  on  the  other  hand,  can  make  its 
dam  or  canal,  and  a  bird  its  nest,  as  well,  or  nearly  as  well,  and 

'  'Contributions  to  the  Theory  of  Natural  Selection,'  1870,  p.  212. 


Chap.  III.  Mental  Powers.  69 

a  spider  its  wonderful  web,  quite  as  well,''  the  fii-st  time  it  tries, 
as  when  old  and  experienced. 

To  return  to  our  immediate  subject :  the  lower  animals,  like 
man,  manifestly  feel  pleasure  and  pain,  happiness  and  misery. 
Haj^piness  is  never  better  exhibited  than  by  young  animals,  such 
as  i^uiDpies,  kittens,  lambs,  &c.,  when  playing  together,  like  our 
own  childi'en.  Even  insects  play  together,  as  has  been  described 
by  that  excellent  observer,  P.  Huber,'^  who  saw  ants  chasing  and 
pretending  to  bite  each  other,  like  so  many  puppies. 

The  fact  that  the  lower  animals  are  excited  by  the  same 
emotions  as  ourselves  is  so  well  established,  that  it  will  not  be 
necessary  to  weary  the  reader  by  many  details.  Terror  acts  in 
the  same  manner  on  them  as  on  us,  causing  the  muscles  to 
tremble,  the  heart  to  palpitate,  the  sphincters  to  be  relaxed,  and 
the  hair  to  stand  on  end.  Suspicion,  the  offspring  of  fear,  is 
eminently  characteristic  of  most  wild  animals.  It  is,  I  think, 
impossible  to  read  the  account  given  by  Sir  E.  Tennent,  of  the 
behaviour  of  the  female  elephants,  used  as  decoys,  without 
admitting  that  they  intentionally  practise  deceit,  and  well  know 
what  they  are  about.  Courage  and  timidity  are  extremely 
variable  qualities  in  the  individuals  of  the  same  species,  as  is 
plainly  seen  in  our  dogs.  Some  dogs  and  horses  are  ill-tempered, 
and  easily  turn  sulky ;  others  are  good-temi>ered ;  and  these 
qualities  are  certainly  inherited.  Every  one  knows  how  liable 
animals  are  to  furious  rage,  and  how  plainly  they  show  it. 
Many,  and  probably  true,  anecdotes  have  been  published  on  the 
long-delayed  and  artful  revenge  of  various  animals.  The 
accurate  Eengger,  and  Brehm^  state  that  the  American  and 
African  monkeys  which  they  kept  tame,  certainly  revenged 
themselves.  Sir  Andrew  Smith,  a  zoologist  whose  scrupulous 
accuracy  was  known  to  many  persons,  told  me  the  following 
story  of  which  he  was  himself  an  eye-witness;  at  the  Cape 
of  Good  Hope  an  officer  had  often  i)lagned  a  certain  baboon, 
and  the  animal,  seeing  him  approaching  one  Sunday  for 
parade,  poured  water  into  a  hole  and  hastily  made  some  thick 
mud,  which  he  skilfully  dashed  over  the  officer  as  he  passed 
by,  to  the  amusement  of  many  bystanders.  For  long  after- 
wards the  baboon  rejoiced  and  triumi^hcd  whenever  he  saw  his 
victim. 

6  For    the     evidence    on    this  «  All    the   following    statements, 

heail,    see    Mr.    J.    Traherue    Mog-  given  on  the  authority  of  these  two 

gridge's     most     interesting     work,  naturalists,  are  taken  from  Rongger's 

'Harvesting    Ants    and    Trap-door  'Naturgesch.  der  Siingethiere    von 

Spiders,'  1873,  p.  126,  128.  Paraguay,'  1830,  s.  41-57,  and  from 

^  '  Recherches  sur  les  Mceurs  dcs  Brehm's  'Thierlehen,'  B.  i.  s.  10-87. 
Fourmis,'  1810,  p.  173. 


70  The  Descent  of  Man.  Tart  I. 

The  love  of  a  dog  for  his  master  is  notorious ;  as  an  old 
writer  quaintly  says/  "  A  dog  is  the  only  thing  on  this  earth 
"  that  luvs  you  more  than  he  luvs  himself." 

In  the  agony  of  death  a  dog  has  been  known  to  caress 
his  master,  and  every  one  has  heard  of  the  dog  suffering 
under  vivisection,  who  licked  the  hand  of  the  operator;  this 
man,  unless  the  oiDeration  was  fully  justified  by  an  increase 
of  our  knowledge,  or  unless  he  had  a  heart  of  stone,  must  have 
felt  remorse  to  the  last  hour  of  his  life. 

As  WhewelP^  has  well  asked,  "  who  that  reads  the  touching 
"  instances  of  maternal  affection,  related  so  often  of  the  women  of 
"  all  nations,  and  of  the  females  of  all  animals,  can  doubt  that  the 
"  principle  of  action  is  the  same  in  the  two  cases  ?  "  We  see  mater- 
nal affection  exliibited  in  the  most  trifling  details  ;  thus  Eengger 
observed  an  American  monkey  (a  Cebus)  carefully  driving  away 
the  flies  which  plagued  her  infant;  and  Duvaucel  saw  a 
Hylobates  washing  the  faces  of  her  young  ones  in  a  stream.  So 
intense  is  the  grief  of  female  monkeys  for  the  loss  of  their 
young,  that  it  invariably  caused  the  death  of  certain  kinds  kept 
under  confinement  by  Brehm  in  N.  Africa.  Orphan  monkeys 
were  always  adopted  and  carefully  guarded  by  the  other  monkeys, 
both  males  and  females.  One  female  baboon  had  so  capacious 
a  heart  that  she  not  only  adopted  young  monkeys  of  other 
species,  but  stole  young  dogs  and  cats,  which  she  continually 
carried  about.  Her  kindness,  however,  did  not  go  so  far  as  to 
share  her  food  with  her  adopted  offspring,  at  which  Brehm  was 
surprised,  as  his  monkeys  always  divided  everythiDg  quite 
fairly  with  their  own  young  ones.  An  adopted  kitten  scratched 
this  affectionate  baboon,  who  certainly  had  a  fine  intellect,  for 
she  was  much  astonished  at  being  scratched,  and  immediately 
examined  the  kitten's  feet,  and  without  more  ado  bit  off  the 
claws.^^  In  the  Zoological  Gardens,  I  heard  from  the  keeper 
that  an  old  baboon  (C,  chacmci)  had  adopted  a  Ehesus  monkey; 
but  when  a  young  drill  and  mandrill  were  placed  in  the  cage, 
she  seemed  to  perceive  that  these  monkeys,  though  distinct 
species,  were  her  nearer  relatives,  for  she  at  once  rejected  the 
Ehesus  and  adopted  both  of  them.  The  young  Ehesus,  as  I  saw, 
was  greatly  discontented  at  being  thus  rejected,  and  it  would, 
like  a  naughty  child,  annoy  and  attack  the  young  drill  and  mandrill 

3  Quoted  by  Dr.  Lauder  Lindsay,  72),  disputes  the  possibility  of  this. 

u  his  '  Physiology  of  Mind  in  the  act  as  described  by  Brehm,  for  the 

Lower  Animals ;'  '  Journal  of  Mental  sake     of    discrediting     my     work. 

Science,'  April  1871,  p.  38.  Therefore  I  tried,  and  found  that  I 

10  '  BridiT-ewater  Treatise,'  p.  263.  could    readily  seize    with    my  own 

^1  A  critic,  without  any  grounds  teeth  the    sharp   little   claws    of  a 

('Quarterly  Review,'  July  1871,  p.  kitten  nearly  five  weeks  old. 


CuAP.  III.  Mental  Powers,  yi 

whenever  it  could  do  so  with  safety ;  this  conduct  exciting  great 
indignation  in  the  old  baboon.  Monkeys  will  also,  according  to 
Brehm,  defend  their  master  when  attacked  by  any  one,  as  well  as 
dogs  to  whom  they  are  attached,  from  the  attacks  of  other  dogs. 
But  we  here  trench  on  the  subjects  of  sympathy  and  fidelity,  to 
which  I  shall  recur.  Some  of  Brehm's  monkeys  took  much 
delight  in  teasing  a  certain  old  dog  whom  they  disliked,  as 
well  as  other  animals,  in  various  ingenious  ways. 

Most  of  ihe  more  complex  emotions  arc  common  to  the 
higher  animals  and  ourselves.  Every  one  has  seen  how  jealous 
a  dog  is  of  his  master's  affection,  if  lavished  on  any  other 
creature ;  and  I  have  observed  the  same  fact  with  monkeys. 
Tliis  shews  that  animals  not  only  love,  but  have  desire  to  bo 
loved.  Animals  manifestly  feel  emulation.  They  love  ajipro- 
bation  or  praise ;  and  a  dog  carrying  a  basket  for  his  master 
exhibits  in  a  high  degree  self-complacency  or  pride.  There  can, 
I  think,  be  no  doubt  that  a  dog  feels  shame,  as  distinct  from 
fear,  and  something  very  like  modesty  when  begging  too  often 
for  food.  A  great  dog  scorns  the  snarling  of  a  httle  dog,  and 
this  may  be  called  magnanimity.  Several  observers  have  stated 
that  monkeys  certainly  dislike  being  laughed  at ;  and  they 
sometimes  invent  imaginary  offences.  In  the  Zoological  Gardens 
I  saw  a  baboon  who  always  got  into  a  furious  rage  v/hen  his 
keeper  took  out  a  letter  or  book  and  read  it  aloud  to  him ;  and 
his  rage  was  so  violent  that,  as  I  witnessed  on  one  occasion,  he 
bit  his  own  leg  till  the  blood  flowed.  Dogs  show  what  may  be 
fairly  called  a  sense  of  humour,  as  distinct  from  mere  jilay ;  if 
a  bit  of  stick  or  other  such  object  be  thrown  to  one,  he  will  often 
carry  it  away  for  a  short  distance ;  and  then  squatting  down 
with  it  on  the  ground  close  before  him,  will  wait  until  his 
master  comes  quite  close  to  take  it  away.  The  dog  will  then 
seize  it  and  rush  away  in  triumph,  repeating  the  same  manoeuvre, 
and  evidently  enjoying  the  practical  joke. 

Wo  will  now  turn  to  the  more  intellectual  emotions  and 
faculties,  which  are  very  important,  as  forming  the  basis  for  the 
development  of  the  higher  mental  powers.  Animals  manifestly 
enjoy  excitement,  and  suffer  from  ennui,  as  may  be  seen  with 
dogs,  and,  according  to  Eengger,  with  monkeys.  All  animals 
feel  Wonder,  and  many  exhibit  Curiositij.  They  sometimes 
suffer  from  this  latter  quality,  as  when  the  hunter  plays  antics 
and  thus  attracts  them ;  I  have  witnessed  this  with  doer,  and  so 
it  is  with  the  wary  chamois,  and  with  some  kinds  of  wild-ducks. 
Brehm  gives  a  curious  account  of  the  instinctive  dread,  which 
his  monkeys  exhibited,  for  snakes;  but  their  curiosity  was 
so  great  that  they  could  not  desist  from  occasionally  satiating 


72  The  Descent  of  Man.  Part  I. 

their  horror  in  a  most  human  fashion,  by  lifting  up  the  lid  of  the 
box  in  which  the  snakes  were  kept.  I  was  so  much  surprised  at 
his  account,  that  I  took  a  stuffed  and  coiled-up  snake  into  the 
monkey-house  at  the  Zoological  Gardens,  and  the  excitement 
thus  caused  was  one  of  the  most  curious  spectacles  which  I  ever 
beheld.  Three  species  of  Cercopithecus  were  the  most  alarmed ; 
they  dashed  about  their  cages,  and  uttered  sharp  signal  cries  of 
danger,  which  were  understood  by  the  other  monkeys.  A  few 
young  monkeys  and  one  old  Anubis  baboon  alone  took  no  notice 
of  the  snake.  I  then  placed  the  stuffed  specimen  on  the  ground 
in  one  of  the  larger  comx)artments.  After  a  time  all  the  monkeys 
collected  round  it  in  a  large  ch'cle,  and  staring  intently, 
presented  a  most  ludicrous  appearance.  They  became  extremely 
nervous ;  so  that  when  a  wooden  ball,  with  which  they  were 
familiar  as  a  plaything,  was  accidentally  moved  in  the  straw, 
under  which  it  was  partly  hidden,  they  all  instantly  started 
away.  These  monkeys  behaved  very  differently  when  a  dead 
fish,  a  mouse,^^  a  living  turtle,  and  other  new  objects  were  placed 
in  their  cages;  for  though  at  fkst  frightened,  they  soon 
approached,  handled  and  examined  them.  I  then  placed  a  live 
snake  in  a  paper  bag,  with  the  mouth  loosely  closed,  in  one  of 
the  larger  compartments.  One  of  the  monkeys  immediately 
approached,  cautiously  opened  the  bag  a  little,  peeped  in,  and 
instantly  dashed  away.  Then  I  witnessed  what  Brehm  has 
described,  for  monkey  after  monkey,  with  head  raised  high  and 
turned  on  one  side,  could  not  resist  taking  a  momentary  peep 
into  the  upright  bag,  at  the  dreadful  object  lying  quietly  at  the 
bottom.  It  would  almost  appear  as  if  monkeys  had  some 
notion  of  zoological  affinities,  for  those  kept  by  Brehm  exhibited 
a  strange,  though  mistaken,  instinctive  di-ead  of  innocent  lizards 
and  frogs..  An  orang,  also,  has  been  known  to  be  much  alarmed 
at  the  first  sight  of  a  turtle.^^ 

The  principle  of  Imitation  is  strong  in  man,  and  especially,  as 
I  have  myself  observed,  with  savages.  In  certain  morbid  states 
of  the  brain  this  tendency  is  exaggerated  to  an  extraordinary 
degree ;  some  hemiplegic  patients  and  others,  at  the  commence- 
ment of  inflammatory  softening  of  the  brain,  unconsciously 
imitate  every  word  which  is  uttered,  whether  in  their  own  or  in 
a  foreign  language,  and  every  gesture  or  action  which  is  per- 
formed near  them.-*     Desor^^  has  remarked  that  no  animal 

^2  I  have  given  a  short  account  of  Mammalia,'  1841,  p.  405. 
of  their  behaviour  on  this  occasion  '*   Dr.    Bateman  '  On    Aphasia,' 

in  my  'Expression  of  the  Emotions,'  1870,  p.  110. 
p.  43.  *'  Quoted  by  Vogt,  'Me'moire  sur 

»3  \V.  C.  L.  Martin,  'Nat.  Hist,  les  Microcephales,'  1867,  p.  168. 


CiL^.p.  III.  Mental  Powers.  73 

volunt<arily  imitates  an  action  performed  by  man,  until  in  the 
ascending  scale  we  come  to  monkeys,  which  are  well  known  to 
be  ridiculous  mockers.  Animals,  however,  sometimes  imitate 
each  other's  actions  :  thus  two  species  of  wolves,  wliich  had  been 
reared  by  dogs,  learned  to  bark,  as  docs  sometimes  the  jackal,"^ 
but  whether  this  can  bo  called  voluntary  imitation  is  another 
question.  Birds  imitate  the  songs  of  their  parents,  and  some- 
times of  other  birds ;  and  parrots  are  notorious  imitators  of  any 
sound  which  they  often  hear.  Bureau  de  la  Malic  gives  an 
account  ^^  of  a  dog  reared  by  a  cat,  who  learnt  to  imitate  the 
well-known  action  of  a  cat  licking  her  paws,  and  thus  washing 
her  ears  and  face;  this  was  also  witnessed  by  the  celebrated 
naturalist  Audouin.  I  have  received  several  confirmatory  ac- 
counts ;  in  one  of  these,  a  dog  had  not  been  suckled  by  a  cat, 
but  had  been  brought  up  with  one,  together  with  kittens,  and 
had  thus  acquired  the  above  habit,  which  he  ever  afterwards 
practised  during  his  life  of  thirteen  years.  Dureau  de  la  Malle's 
dog  likewise  learnt  from  the  kittens  to  play  with  a  ball  by  roll- 
ing it  about  with  his  fore  paws,  and  springing  on  it.  A  corre- 
spondent assures  me  that  a  cat  in  his  house  used  to  put  her  paws 
into  jugs  of  milk  having  too  narrow  a  mouth  for  her  head.  A 
kitten  of  tliis  cat  soon  learned  the  same  trick,  and  practised  it 
ever  afterwards,  whenever  there  was  an  opportunity. 

The  parents  of  many  animals,  trusting  to  the  principle  of 
imitation  in  their  young,  and  more  especially  to  their  instinctive 
or  inherited  tendencies,  may  be  said  to  educate  them.  "We  see 
this  when  a  cat  brings  a  live  mouse  to  her  kittens ;  and  Dureau 
de  la  Malle  has  given  a  curious  account  (in  the  paper  above 
quoted)  of  his  observations  on  hawks  which  taught  their  young 
dexterity,  as  well  as  judgment  of  distances,  by  first  chopping 
through  the  air  dead  mice  and  sparrows,  which  the  young 
generally  failed  to  catch,  and  then  bringing  them  live  birds 
and  letting  them  loose. 

Hardly  any  faculty  is  more  important  for  the  intellectual 
progress  of  man  than  Attention.  Animals  clearly  manifest  this 
power,  as  when  a  cat  watches  by  a  hole  and  prepares  to  spring 
on  its  prey.  Wild  animals  sometimes  become  so  absorbed  when 
thus  engaged,  that  they  may  be  easily  approached.  Mr.  Bartlett 
has  given  me  a  curious  proof  how  variable  this  faculty  is  in 
monkeys.  A  man  who  trains  monkeys  to  act  in  plays,  used  to 
purchase  common  kinds  from  the  Zoological  Society  at  the  price 
of  five  pounds  for  each  ;  but  he  offered  to  give  double  the  price, 

*^   'The  Variation  of  Animals  and  ''  'Annales    des    Sc.    Nat.'    (1st 

Plants  under  Domestication,'  vol.  i.      Series),  torn.  xxii.  p.  397. 
p.  27. 


74  The  Descent  of  Man.  Part  I. 

if  he  might  keep  three  or  four  of  them  for  a  few  days,  in  order 
to  select  one.  When  asked  how  he  could  possibly  learn  so  soon, 
whether  a  particular  monkey  would  tui'n  out  a  good  actor,  he 
answered  that  it  all  depended  on  their  power  of  attention.  Jf, 
when  he  was  talking  and  explaining  anything  to  a  monkey,  its 
attention  was  easily  distracted,  as  by  a  fly  on  the  wall  or  other 
trifling  object,  the  case  was  hopeless.  If  he  tried  by  punishment 
to  make  an  inattentive  monkey  act,  it  turned  sulky.  On  the 
other  hand,  a  monkey  which  carefully  attended  to  him  could 
always  be  trained. 

It  is  almost  superfluous  to  state  that  animals  have  excellent 
Meinories  for  persons  and  places.  A  baboon  at  the  Cape  of  Good 
Hope,  as  I  have  been  informed  by  Sir  Andrew  Smith,  recognised 
him  with  joy  after  an  absence  of  nine  months.  I  had  a  dog  who 
was  savage  and  averse  to  all  strangers,  and  I  purposely  tried  his 
memory  after  an  absence  of  five  years  and  two  days.  I  went 
near  the  stable  where  he  lived,  and  shouted  to  him  in  my  old 
manner ;  he  shewed  no  joy,  but  instantly  followed  me  out  walk- 
ing, and  obeyed  me,  exactly  as  if  I  had  parted  with  him  only 
half  an  hour  before.  A  train  of  old  associations,  dormant  during 
five  years,  had  thus  been  instantaneously  awakened  in  his  mind. 
Even  ants,  as  P.  Huber  ^^  has  clearly  shewn,  recognised  their 
fellow-ants  belonging  to  the  same  community  after  a  separation 
of  four  months.  Animals  can  certainly  by  some  means  judge  of 
the  intervals  of  time  between  recurrent  events. 

The  Imagination  is  one  of  the  highest  prerogatives  of  man. 
By  this  faculty  he  unites  former  images  and  ideas,  independently 
of  the  will,  and  thus  creates  brilliant  and  novel  results.  A  poet, 
as  Jean  Paul  Eichter  remarks,^^  "  who  must  reflect  whether  he 
"  shall  make  a  character  say  yes  or  no — to  the  devil  with  him ; 
"  he  is  only  a  stupid  corpse."  Dreaming  gives  us  the  best  notion 
of  this  power ;  as  Jean  Paul  again  says,  "  The  dream  is  an  in- 
"  voluntary  art  of  poetry."  The  value  of  the  products  of  our 
imagination  depends  of  course  on  the  number,  accuracy,  and 
clearness  of  our  impressions,  on  our  judgment  and  taste  in 
selecting  or  rejecting  the  involuntary  combinations,  and  to  a 
certain  extent  on  our  power  of  voluntarily  combining  them.  As 
dogs,  cats,  horses,  and  probably  all  the  higher  animals,  even 
birds  ^°  have  vivid  dreams,  and  this  is  shewn  by  their  movements 
and  the  sounds  uttered,  we  must  admit  that  they  possess  some 

'^    '  Les    Mceurs     des    Fourmis,'  20  jyy    Jerdon,   *  Birds   of  India,' 

1810,  p.  150.  vol.  i.  1862,  p.  xxi.     Houzeau   says 

^^  Quoted  in  Dr.  Maudsley's  '  Phy-  that  his  pavokeets  and  canary-birds 

siology  and  Pathology  of  Mind,' 18G8,  dreamt:  *  Facultes   Mentales/  torn, 

pp.  19,  220.  ii.  p.  13G. 


Chap.  111.  Mental  Poivcrs.  75 


power  of  imagination.  There  must  be  something  special,  which 
causes  dogs  to  howl  in  the  night,  and  especially  during  moonlight, 
in  that  remarkable  and  melancholy  manner  called  baying. 
All  dogs  do  not  do  so  ;  and,  according  to  Houzcau,-^  they  do  not 
then  look  at  the  moon,  but  at  some  fixed  point  near  the  horizon. 
Houzeau  thinks  that  their  imaginations  arc  disturbed  by  tlie 
vague  outlines  of  the  surrounding  objects,  and  conjure  up  before 
them  fantastic  images :  if  this  be  so,  their  feelings  may  almost 
be  called  superstitious. 

Of  all  the  faculties  of  the  human  mind,  it  will,  I  presume,  be 
admitted  that  Reason  stands  at  the  summit.  Only  a  few  persons 
now  dispute  that  animals  possess  some  power  of  reasoning. 
Animals  may  constantly  be  seen  to  pause,  deliberate,  and  resolve. 
It  is  a  significant  fiict,  that  the  more  the  habits  of  any  particular 
animal  are  studied  by  a  naturalist,  the  more  he  attributes  to 
reason  and  the  less  to  unlearnt  instincts.-^  In  future  chapters 
we  shall  see  that  some  animals  extremely  low  in  the  scale  appar- 
ently display  a  certain  amount  of  reason.  No  doubt  it  is  often 
difficult  to  distinguish  between  the  power  of  reason  and  that  of 
instinct.  For  instance.  Dr.  Hayes,  in  his  work  on  '  The  Open 
Polar  Sea,'  repeatedly  remarks  that  his  dogs,  instead  of  continu- 
ing to  draw  the  sledges  in  a  compact  body,  diverged  and  separ- 
ated when  they  came  to  thin  ice,  so  that  their  weight  might  be 
more  evenly  distributed.  This  was  often  the  first  warning 
which  the  travellers  received  that  the  ice  was  becoming  thin  and 
dangerous.  Kow,  did  the  dogs  act  thus  from  the  experience  of 
each  individual,  or  from  the  example  of  the  older  and  wiser  dogs, 
or  from  an  inherited  habit,  that  is  from  instinct  ?  This  instinct, 
may  possibly  have  arisen  since  the  time,  long  ago,  when  dogs 
were  first  employed  by  the  natives  in  drawing  their  sledges ;  or 
the  Arctic  wolves,  the  parent-stock  of  the  Esquimaux  dog,  may 
have  acquired  an  instinct,  impelling  them  not  to  attack  their 
prey  in  a  close  pack,  when  on  thin  ice. 

We  can  only  judge  by  the  circumstances  under  which  actions 
are  performed,  whether  they  are  due  to  instinct,  or  to  reason,  or 
to  the  mere  association  of  ideas :  this  latter  principle,  however, 
is  intimately  connected  with  reason.  A  curious  case  has  been 
given  by  Prof.  Mobius,'''^  of  a  pike,  separated  by  a  plate  of  glass 
from  an  adjoining  aquarium  stocked  with  fish,  and  who  often 
dashed  himself  with  such  violence  against  the  glass  in  trying  to 

^*    '  Facultes    Mentales   des   Ani-  I    cannot    help    thinking,  however, 

maux,'  1872,  torn,  ii,  p.  181.  that  he  goes  too  far  in  underrating 

^*  Mr.   L.  H.  Morgan's  work    on  the  power  of  Instinct. 
'The  American  Beaver,'  1868,  offers  -^  'Die  Bewegungou  der  Tliiere, 

a  good  illustration  of  this  remark.  &c.,  187.'J,  p.  11. 


76  The  Descent  of  Man.  Part  I. 


catch  tlie  other  fishes,  that  he  was  sometimes  completely 
stunned.  The  pike  went  on  thus  for  three  months,  but  at  last 
learnt  caution,  and  ceased  to  do  so.  The  plate  of  glass  was  then 
removed,  but  the  pike  would  .not  attack  these  particular  fishes, 
though  he  would  devour  others  which  were  afterwards  intro- 
duced ;  so  strongly  was  the  idea  of  a  violent  shock  associated 
in  his  feeble  mind  with  the  attempt  on  his  former  neighboiu's. 
If  a  savage,  who  had  never  seen  a  large  plate-glass  window, 
were  to  dash  himself  even  once  against  it,  he  would  for  a  long 
time  afterwards  associate  a  shock  with  a  window-frame ;  but 
very  differently  from  the  pike,  he  would  probably  reflect  on  the 
nature  of  the  impediment,  and  be  cautious  under  analogous 
circumstances.  Now  with  monkeys,  as  we  shall  presently  see,  a 
painful  or  merely  a  disagreeable  impression,  from  an  action  once 
performed,  is  sometimes  sufficient  to  prevent  the  animal  from 
repeating  it.  If  we  attribute  this  difference  between  the  monkey 
and  the  pike  solely  to  the  association  of  ideas  being  so  much 
stronger  and  more  persistent  in  the  one  than  the  other,  though 
the  pike  often  received  much  the  more  severe  injury,  can  we 
maintain  in  the  case  of  man  that  a  similar  difference  implies  the 
possession  of  a  fundamentally  different  mind  ? 

Houzeau  relates  ^^  that,  whilst  crossing  a  wide  and  arid  plain 
in  Texas,  his  two  dogs  suffered  greatly  from  thirst,  and  that 
between  thii'ty  and  forty  times  they  rushed  down  the  hollows 
to  search  for  water.  These  hollows  were  not  valleys,  and  there 
were  no  trees  in  them,  or  any  other  difference  in  the  vegetation, 
and  ^s  they  were  absolutely  diy  there  could  have  been  no 
smell  of  damp  earth.  The  dogs  behaved  as  if  they  knew  that 
a  dip  in  the  ground  offered  them  the  best  chance  of  finding 
water,  and  Houzeau  has  often  witnessed  the  same  behaviour  in 
other  animals. 

I  have  seen,  as  I  daresay  have  others,  that  when  a  small 
object  is  thrown  on  the  ground  beyond  the  reach  of  one  of  the 
elephants  in  the  Zoological  Gardens,  he  blows  through  his  trunk 
on  the  ground  beyond  the  object,  so  that  the  current  reflected 
on  all  sides  may  drive  the  object  within  his  reach.  Again  a  well- 
known  ethnologist,  LIr.  Westroj)p,  informs  me  that  he  observed  in 
Vienna  a  bear  deliberately  making  with  his  paw  a  current  in 
some  water,  which  was  close  to  the  bars  of  his  cage,  so  as  to 
draw  a  piece  of  floating  bread  within  his  reach.  These  actions  of 
the  elephant  and  bear  can  hardly  be  attributed  to  instinct  or 
inherited  habit,  as  they  would  be  of  little  use  to  an  animal  in  a 
state  of  nature.    Now,  what  is  the  difference  between  such 

2*  'Facultcs  Mentales  des  Animaux,'  1872,  torn.  ii.  p.  265. 


Chap.  III.  Mental  Powers.  77 

actions,  when  performed  by  an  uncultivated  man,  and  by  one  of 
tlie  higlier  animals  ? 

The  savage  and  the  dog  have  often  found  water  at  a  low  level, 
and  the  coincidence  under  such  circumstances  has  become  asso- 
ciated in  their  minds.  A  cultivated  man  would  perhaps  make 
some  general  proposition  on  the  subject ;  but  from  all  that  wc 
know  of  savages  it  is  extremely  doubtful  whether  tliey  would  do 
so,  and  a  dog- certainly  would  not.  But  a  savage,  as  well  as  a 
dog,  would  search  in  the  same  way,  though  frequently  dis- 
appointed; and  in  both  it  seems  to  be  equally  an  act  of  reason, 
whether  or  not  any  general  proposition  on  the  subject  is 
consciously  placed  before  the  mind.^  The  same  would  apply  to 
tlie  elephant  and  the  bear  making  currents  in  the  air  or  water. 
The  savage  would  certainly  neither  know  nor  care  by  what  law 
the  desired  movements  were  effected;  yet  his  act  would  be 
guided  by  a  rude  process  of  reasoning,  as  surely  as  would  a 
philosopher  in  his  longest  chain  of  deductions.  There  would  no 
doubt  be  this  difference  between  him  and  one  of  the  higher 
animals,  that  he  would .  take  notice  of  much  slighter  circum- 
stances and  conditions,  and  would  observe  any  connection 
between  them  after  much  less  experience,  and  this  would  be  of 
paramount  importance.  I  kept  a  daily  record  of  the  actions  of 
one  of  my  infants,  and  when  he  was  about  eleven  months  old, 
and  before  he  could  speak  a  single  word,  I  was  continually 
struck  with  the  greater  quickness,  with  which  all  sorts  of  objects 
and  sounds  were  associated  together  in  his  mind,  compared  with 
that  of  the  most  intelligent  dogs  I  ever  knew.  But  the  higher 
animals  differ  in  exactly  the  same  way  in  this  power  of  associa- 
tion from  those  low  in  the  scale,  such  as  the  pike,  as  well  as  in 
that  of  drawing  inferences  and  of  observation. 

The  promptings  of  reason,  after  very  short  experience,  are  well 
shewn  by  the  following  actions  of  American  monkeys,  which 
stand  low  in  their  order.  Eengger,  a  most  careful  observer, 
states  that  when  he  flrst  gave  eggs  to  his  monkeys  in  Paraguay, 
they  smashed  them,  and  thus  lost  much  of  their  contents ;  after- 
wards they  gently  hit  one  end  against  some  hard  body,  and 
picked  off  the  bits  of  shell  with  their  fingers.  After  cutting 
themselves  only  once  with  any  sharp  tool,  they  would  not  toucli 
it  again,  or  would  handle  it  with  the  greatest  caution.  Lumps 
of  sugar  were  often  given  them  wi-apped  up  in  paper;   and 

-*  Prof.  Huxley  has  analysed  with  See     his     article,     *  Mr.     Darwin's 

admirable  clearness  the  mental  steps  Critics,'  in    the  *  Contemporary  Ke- 

by  which  a  man,  as  well  as  a  dog,  view,'  Nov.  1871,  p.  462,  and  in  his 

arrives  at  a  conclusion  in  a  case  '  Critiques  and  Essays,'  1873,  p.  279. 
analogous  to  that  given  in  my  text. 


78  The  Descent  of  Man.  Part  I. 

Eengger  sometimes  put  a  live  wasp  in  the  paper,  so  that  in 
hastily  unfolding  it  they  got  stung ;  after  this  had  once  happened, 
they  always  first  held  the  packet  to  their  ears  to  detect  any 
movement  within."^ 

The  following  cases  relate  to  dogs.  Mr,  Colquhoun^^  winged 
two  wild-ducks,  which  fell  on  the  further  side  of  a  stream ;  his 
retriever  tried  to  bring  over  both  at  once,  but  could  not  succeed; 
she  then,  though  never  before  known  to  ruffle  a  feather, 
deliberately  killed  one,  brought  over  the  other,  and  returned 
for  the  dead  bird.  Col.  Hutchinson  relates  that  two  jDartridges 
were  shot  at  once,  one  being  killed,  the  other  wounded ;  the 
latter  ran  away,  and  was  caught  by  the  retriever,  who  on  her 
return  came  across  the  dead  bird;  "she  stopped,  evidently 
"  gTeatly  puzzled,  and  after  one  or  two  trials,  finding  she  could 
"  not  take  it  up  without  permitting  the  escape  of  the  winged 
"  bird,  she  considered  a  moment,  then  deliberately  murdered  it 
"  by  giving  it  a  severe  crunch,  and  afterwards  brought  away 
"  both  together.  This  was  the  only  known  instance  of  her 
"  ever  having  wilfully  injured  any  game."  Here  we  have  reason 
though  not  quite  perfect,  for  the  retriever  might  have  brought 
the  wounded  bird  first  and  then  retui-ned  for  the  dead  one,  as  in 
the  case  of  the  two  wild-ducks.  I  give  the  above  cases,  as 
resting  on  the  evidence  of  two  independent  witnesses,  and 
because  in  both  instances  the  retrievers,  after  deliberation, 
broke  through  a  habit  which  is  inherited  by  them  (that  of  not 
killing  the  game  retrieved),  and  because  they  shew  how  strong 
their  reasoning  faculty  must  have  been  to  overcome  a  fixed 
habit. 

I  will  conclude  by  quoting  a  remark  by  the  illustrious 
Humboldt.-^  "  The  muleteers  in  S.  America  say,  '  I  will  not  give 
"  '  you  the  mule  whose  step  is  easiest,  but  la  mas  racional, — the 
" '  one  that  reasons  best ;' "  and  as  he  adds,  "  this  popular  expres- 
"  sion,  dictated  by  long  experience,  combats  the  system  of 
"  animated  machines,  better  perhaps  than  all  the  arguments  of 
"  speculative  philosophy."  Nevertheless  some  wiiters  even  yet 
deny  that  the  higher  animals  possess  a  trace  of  reason ;  and  they 
endeavour  to  explain  away,  by  what  appears  to  be  mere 
verbiage,^^  all  such  facts  as  those  above  given. 

26  Mr.  Belt,  in  his  most  interest-  p.  45.  Col.  Hutchinson  on  '  Dog 
ing   work,  '  The   Naturalist    in   IS'i-      Breaking,' 1850,  p.  46. 

caragua,'    1874   (p.    119),    likewise  ^^    '  Personal     Karrative,'     Eng. 

describes  various  actions  of  a  tamed  translat.,  vol.  iii.  p.  106. 

Cebus,  which,  I  think,  clearly  shew  ^^  I  am  glad  to  find  that  so  acute 

that    this    animal    possessed    some  a    reasoner   as   Mr.   Leslie    Stephen 

reasoning  power.  ('  Darwinism   and    Divinity,    Essavs 

27  'The    Moor     and     the    Loch,'  on    Free-thinking,'  1873,  p.  80),  in 


Chap.  II  r.  Menial  Poiucrs.  79 


It  has,  I  think,  now  been  slicwn  that  man  and  the  higher 
animals,  especially  the  Primates,  have  some  few  instincts  in 
commoift-  All  have  the  same  senses,  intuitions,  and  sensations, — 
similar  passions,  affections,  and  emotions,  even  the  more  complex 
ones,  such  as  jealousy,  suspicion,  emulation,  gratitude,  and 
magnanimity  ;  they  j)ractise  deceit  and  arc  revengeful ;  they  are 
sometimes  susceptible  to  ridicule,  and  even  have  a  sense  of 
humour ;  they  feel  wonder  and  curiosity  ;  they  possess  the  same 
faculties  of  imitation,  attention,  deliberation,  choice,  memory, 
imagination,  the  association  of  ideas,  and  reason,  though  in  very 
different  degrees.  The  individuals  of  the  same  species  graduate 
in  intellect  from  absolute  imbecility  to  high  excellence.  They 
are  also  liable  to  insanity,  though  far  less  often  than  in  the  case 
of  man.'^'^  Nevertheless,  many  authors  have  insisted  that  man  is 
divided  by  an  insuperable  barrier  from  all  the  lower  animals  in 
his  mental  faculties.  I  formerly  made  a  collection  of  above  a 
score  of  such  aphorisms,  but  they  are  almost  worthless,  as  their 
wide  difference  and  number  prove  the  difficulty,  if  not  the  im- 
possibility, of  the  attempt.  It  has  been  asserted  that  man  alone 
is  capable  of  progressive  improvement ;  that  he  alone  makes  use 
of  tools  or  fire,  domesticates  other  animals,  or  possesses  property; 
that  no  animal  has  the  power  of  abstraction,  or  of  forming 
general  concepts,  is  self-conscious  and  comprehends  itself ;  that 
no  animal  employs  language;  that  man  alone  has  a  sense  of 
beauty,  is  liable  to  caprice,  has  the  feeling  of  gratitude,  mystery, 
&c.;  believes  in  God,  or  is  endowed  with  a  conscience.  I  will 
hazard  a  few  remarks  on  the  more  important  and  interesting  of 
these  points. 

Archbishop  Sumner  formerly  maintained '^^  that  man  alone  is 
capable  of  progressive  improvement.  That  he  is  capable  of 
incomj)arably  greater  and  more  rapid  improvement  than  is  any 
other  animal,  admits  of  no  dispute;  and  this  is  mainly 
due  to  his  power  of  speaking  and  handing  down  his  acquired 
knowledge.  With  animals,  looking  first  to  the  individual,  every 
one  who  has  had  any  experience  in  setting  traps,  knows  that 


speaking  of  the  supposed  impassable  "  lures.     It    is   diflicult    to    umU'r- 

barrier  between  the  miuds   of  man  "  stand  how  anybody  wiio   has  ever 

and  the  lower  animals,  says,  "The  "  kept  a  dog,  or  seen  an   elephant, 

"  distinctions,    indeed,    which    have  "  can    have    any   doubts    as    to    an 

"  been   drawn,  seem   to   us   to   rest  "  animal's  power  of  performing  tlio 

'•  upon  no  better  founJation  than  a  "  essential  ]»rocesses  of  reasoning," 

"  great    many    other    metaphysical  ^°  See   '  Gladness  in  Animals,'  by 

"  distinctions;  that  is,  the  assump-  Dr.  W.  Lauder  Lindsay,  in  'Journal 

"  tion   that    because   you    can    give  of  Mental  Science,' July  1871. 

"  two   things   ditl'erent  names,  they  "   Quoted  by  Sir  C.  i^ycU,  '  Auti- 

**  must  therefore  have  ditlerent  na-  quity  of  Man,'  p.  497. 


'80  The  Descent  of  Man.  Part  I. 

young  animals  can  be  caught  much  more  easily  than  old  ones ; 
and  they  can  be  much  more  easily  approached  by  an  enemy. 
Even  with  respect  to  old  animals,  it  is  impossible  to  catch"  many  in 
the  same  place  and  in  the  same  kind  of  trap,  or  to  destroy  them 
by  the  same  kind  of  poison ;  yet  it  is  improbable  that  all  should 
have  partaken  of  the  poison,  and  impossible  that  all  should  have 
been  caught  in  a  trap.  They  must  learn  caution  by  seeing  their 
brethren  caught  or  poisoned.  In  North  America,  where  the  fur- 
bearing  animals  have  long  been  pursued,  they  exhibit,  according 
to  the  unanimous  testimony  of  all  observers,  an  almost  incredible 
amount  of  sagacity,  caution  and  cunning ;  but  trapping  has  been 
there  so  long  carried  on,  that  inheritance  may  possibly  have  come 
into  play.  I  have  received  several  accounts  that  when  telegraphs 
are  first  set  up  in  any  district,  many  birds  kill  themselves  by 
flying  against  the  wires,  but  that  in  the  course  of  a  very  few 
years  they  learn  to  avoid  this  danger,  by  seeing,  as  it  would 
appear,  their  comrades  killed.^^ 

If  we  look  to  successive  genprations,  or  to  the  race,  there  is  no 
doubt  that  birds  and  other  animals  gradually  both  acquire  and 
lose  caution  in  relation  to  man  or  other  enemies  ;^  and  this 
caution  is  certainly  in  chief  part  an  inherited  habit  or  instinct, 
but  in  part  the  result  of  individual  experience.  A  good  observer, 
Leroy,^*  states,  that  in  districts  where  foxes  are  much  hunted, 
the  young,  on  first  leaving  their  burrows,  are  incontestably  much 
more  wary  than  the  old  ones  in  districts  where  they  are  not  much 
disturbed. 

Our  domestic  dogs  are  descended  from  wolves  and  jackals,^^ 
and  though  they  may  not  have  gained  in  cunning,  and  may  have 
lost  in  wariness  and  suspicion,  yet  they  have  progressed  in 
certain  moral  qualities,  such  as  in  affection,  trust-worthiness, 
temper,  and  probably  in  general  intelligence.  The  common  rat 
has  conquered  and  beaten  several  other  species  throughout 
Europe,  in  parts  of  North  America,  New  Zealand,  and  recently  in 
Formosa,  as  well  as  on  the  mainland  of  China.  Mr.  Swinhoe,^® 
who  describes  these  two  latter  cases,  attributes  the  victory  of  the 
common  rat  over  the  large  Mus  coninga  to  its  superior  cunning ; 
and  this  latter  quality  may  probably  be  attributed  to  the  habitual 

'2'  For   additional    evidence,  with  -        ^*  '  Lettres  Phil,  sur  I'lntelligence 

details,     see     M.     Houzeau,     '  Les  des  Animaux,'  nouvelle  e'dit.   1802, 

Facultes   Mentales,'    torn.   ii.  1872,  p.  86. 
p.  147.  ^^  See  the  evidence  on  this  head 

^'  See,  with  respect  to  birds  on  in  chap.  i.  vol.  i.  '  On  the  Variation 
oceanic  islands,  my  '  Journal  of  of  Animals  and  Plants  under  Do- 
Researches  during  the  voyage  of  the  mestication.' 

"Beagle,"' 1845,  p.  398.     'Origin  ^e  c^^-qq,   Zoolog.  Soc'    1864,  p. 

of  Species,' 5th  edit.  p.  260.  186. 


CuAP.  III.  Mental  Powers. 


exercise  of  all  its  faculties  in  avoiding  extirpation  b}^  man,  as 
well  as  to  nearly  all  the  loss  cunning  or  weak-minded  rats  having 
been  continuously  destroyed  by  him.  It  is,  however,  possible 
that  the  success  of  the  common  rat  may  be  due  to  its  having 
possessed  greater  cunning  than  its  fellow-species,  before  it 
became  associated  with  man.  To  maintain,  independently  of  any 
direct  evidence,  that  no  animal  during  the  course  of  ages  has 
progressed  in  intellect  or  other  mental  faculties,  is  to  beg  the 
question  of  the  evolution  of  siDccies.  We  have  seen  that,  ac- 
cording to  Lartet,  existing  mammals  belonging  to  several  orders 
have  larger  brains  than  their  ancient  tertiary  prototypes. 

It  has  often  been  said  that  no  animal  uses  any  tool;  but 
the  chimpanzee  m  a  state  of  nature  cracks  a  native  fruit,  some- 
what like  a  wiilnut,  with  a  stone.^  Rengger^^  easily  taught  an 
American  monkey  thus  to  break  open  hard  palm-nuts  ;  and 
afterwards  of  its  own  accord,  it  used  stones  to  open  other  kinds 
of  nuts,  as  well  as  boxes.  It  thus  also  removed  the  soft  rind  of 
fruit  that  had  a  disagreeable  flavour.  Another  monkey  was 
taught  to  open  the  lid  of  a  large  box  with  a  stick,  and  after- 
wards it  used  the  stick  as  a  lever  to  move  heavy  bodies ;  and  1 
have  myself  seen  a  young  orang  put  a  stick  into  a  crevice,  slip 
his  hand  to  the  other  end,  and  use  it  in  the  proper  manner  as  a 
lever.  The  tamed  elephants  in  India  are  well  known  to  break 
off  branches  of  trees  and  use  them  to  drive  away  the  flies ;  and 
this  same  act  has  been  observed  in  an  elejihant  in  a  state  ot 
nature.^^  I  have  seen  a  young  orang,  when  she  thought  she  was 
going  to  be  whipped,  cover  and  protect  herself  with  a  blanket  or 
straw.  In  these  several  cases  stones  and  sticks  were  emjiloyed 
as  imjDlements ;  but  they  are  likewise  used  as  weapons.  Brehm'*° 
states,  on  the  authority  of  the  well-known  traveller  Schiniper, 
that  in  Abyssinia  w^heu  the  baboons  belonging  to  one  species 
(  C.  ydada)  descend  in  troops  from  the  mountains  to  plunder  the 
fields,  they  sometimes  encounter  troops  of  another  species 
(C.  hamadryas),  and  then  a  fight  ensues.  The  Geladas  roll 
down  great  stones,  which  the  Hamadryas  try  to  avoid,  and  then 
both  species,  making  a  great  uproar,  rush  furiously  against  each 
other.  Erehm,  when,  accompanying  the  Duke  of  Coburg-Golha, 
aided  in  an  attack  with  fire-arms  on  a  troop  of  baboons  in  the 
pass  of  Mensa  in  Abyssinia.  The  baboons  in  return  rolled  so 
many  stones  down  the  mountain,  some  as  large  as  a  man's  head, 
that  the  attackers  had  to  beat  a  hasty  retreat ;  and  the  pass  was 

^'  Savage  and  Wyman  in  '  Boston  1830,  s.  51-5G. 

Journal  of  Nat.  Hist.' vol.  iv.  1843-  ^a  'pji^   'Indian  iMeld,'   iMarch  4, 

44,  p.  383.  1871. 

'«  'Sau'-ethiere    von     Paracruav,'  *<>  '  ThiorlcdxMi,'  P..  i.  s.  79,  82. 


82  The  Descent  of  Man.  Tart  I. 

actually  closed  for  a  time  against  the  caravan.  It  deserves 
notice  that  these  baboons  thus  acted  in  concert.  Mr.  Wallace*^ 
on  three  occasions  saw  female  orangs,  accompanied  by  their 
young,  "  breaking  off  branches  and  the  great  spiny  fruit  of  the 
"  Durian  tree,  with  every  appearance  of  rage;  causing  such  a 
"  shower  of  missiles  as  effectually  kept  us  from  approaching  too 
"  near  the  tree."  As  I  have  repeatedly  seen,  a  chimpanzee  will 
throw  any  object  at  hand  at  a  person  who  offends  him ;  and  the 
before  mentioned  baboon  at  the  Cape  of  Good  Hope  prepared 
mud  for  the  purpose. 

In  the  Zoological  Gardens,  a  monkey,  which  had  weak  teeth, 
used  to  break  open  nuts  with  a  stone ;  and  I  was  assui'ed  by  the 
keepers  that  after  using  the  stone,  he  hid  it  in  the  straw,  and 
would  not  let  any  other  monkey  touch  it.  Here,  then,  we  have 
the  idea  of  property  ;  but  this  idea  is  common  to  every  dog  with 
a  bone,  and  to  most  or  all  birds  with  their  nests. 

The  Duke  of  Argyll'*-  remarks,  that  the  fashioning  of  an 
implement  for  a  special  purpose  is  absolutely  peculiar  to  man ; 
and  he  considers  that  this  forms  an  immeasiu'able  gulf  between 
him  and  the  brutes.  This  is  no  doubt  a  very  important  dis- 
tinction ;  but  there  appears  to  me  much  truth  in  Sir  J.  Lubbock's 
suggestion,'^  that  when  primeval  man  first  used  flint-stones  for 
any  purpose,  he  would  have  accidentally  splintered  them,  and 
would  then  have  used  the  sharp  fragments.  From  this  step  it 
would  be  a  small  one  to  break  the  tlints  on  purpose,  and  not  a 
very  wide  step  to  fashion  them  rudely.  This  latter  advance, 
however,  may  have  taken  long  ages,  if  we  may  judge  by  the 
immense  interval  of  time  which  elai^sed  before  the  men  of  the 
neolithic  period  took  to  grinding  and  polishing  their  stone  tools. 
In  breaking  the  flints,  as  Sir  J.  Lubbock  likewise  remarks, 
sparks  would  have  been  emitted,  and  in  grinding  them  heat 
would  have  been  evolved:  thus  the  two  usual  methods  of 
"  obtaining  fire  may  have  originated."  The  nature  of  fire  would 
have  been  known  in  the  many  volcanic  regions  where  lava 
occasionally  flows  through  forests.  The  anthropomoriihous 
apes,  guided  probably  by  instinct,  build  for  themselves  tem- 
porary platforms ;  but  as  many  instincts  are  largely  controlled 
by  Teason,  the  simpler  ones,  such  as  this  of  buildiog  a  platform, 
might  readily  pass  into  a  voluntary  and  conscious  act.  The 
orang  is  known  to  cover  itself  at  night  with  the  leaves  of  the 
Pandanus ;  and  Brehm  states  that  one  of  his  baboons  used  to 
protect  itself  from  the  heat  of  the  sun  by  throwing  a  straw-mat 

41  'The  Malay  Archipelago,'  vol.      14-5,147. 
1.  -1869.  p.  87.  ■'^  '  Trehistoric   Times,'   1865,    p. 

«   '  Primeval    Mau,'     1869,    pp.      473,  &c 


Chap.  II  r.  Mental  Powers.  %i 

over  its  head.  In  tliese  several  habits,  we  probably  see  the  first 
steps  towards  some  of  the  simpler  arts,  such  as  rude  architecture 
and  dress,  as  they  arose  amongst  the  early  progenitors  of  man. 

Abstraction,  General  Conceptions,  ^elf -consciousness,  Mtntd 
Individuality.— It  would  be  very  difficult  for  any  one  with  even 
much  more  knowledge  than  I  possess,  to  determine  how  far 
animals  exhibit  any  traces  of  these  high  mental  powers.  This 
difficulty  arises  from  the  impossibility  of  judging  what  passes 
through  the  mind  of  an  animal ;  and  again,  the  fact  that  writers 
differ  to  a  great  extent  in  the  meaning  which  they  attribute  to 
the  above  terms,  causes  a  further  difficulty.  If  one  may  judge 
from  various  articles  wliich  have  been  published  lately,  the 
greatest  stress  seems  to  be  laid  on  the  supposed  entire  absence 
in  animals  of  the  power  of  abstraction,  or  of  forming  general 
concepts.  But  when  a  dog  sees  another  dog  at  a  distance,  it  is 
often  clear  that  he  perceives  that  it  is  a  dog  in  the  abstract ;  for 
when  he  gets  nearer  his  whole  manner  suddenly  changes,  if  the 
other  dog  be  a  friend.  A  recent  writer  remarks,  that  in  all  such 
cases  it  is  a  pure  assumption  to  assert  that  the  mental  act  is 
not  essentially  of  the  same  nature  in  the  animal  as  in  man.  If 
either  refers  what  he  perceives  with  his  senses  to  a  mental 
concept,  then  so  do  both.*^  When  I  say  to  my  terrier,  in  an 
eager  voice  (and  I  have  made  the  trial  many  times),  "  Hi,  hi, 
where  is  it  ?  "  she  at  once  takes  it  as  a  sign  that  something  is  to 
be  hunted,  and  generally  first  looks  quickly  all  around,  and 
then  rushes  into  the  nearest  thicket,  to  scent  for  any  game,  but 
finding  nothing,  she  looks  up  into  any  neighbouring  tree  for  a 
squirrel.  Now  do  not  these  actions  clearly  shew  that  she  had  in 
lier  mind  a  general  idea  or  concept  that  some  animal  is  to  bo 
discovered  and  hunted  ? 

It  may  be  freely  admitted  that  no  animal  is  self-conscious, 
if  by  this  term  it  is  implied,  that  he  reflects  on  such  points,  as 
whence  he  comes  or  whither  he  will  go,  or  what  is  life  and  deatli, 
and  so  forth.  But  how  can  we  feel  sure  that  an  old  dog  with  an 
excellent  memory  and  some  power  of  imagination,  as  shewn  by 
his  dreams,  never  reflects  on  his  past  pleasures  or  pains  in  the 
chase  ?  And  this  would  be  a  form  of  self-consciousness.  On  tlie 
other  hand,  as  Buchner^^  has  remarked,  how  little  can  the  hard- 
worked  wife  of  a  degraded  Australian  savage,  who  uses  very 
few  abstract  words,  and  cannot  count  above  four,  exert  her  self- 
consciousness,  or  reflect  on  the  nature  of  her  own  existence.  It 
is  generally  admitted,  that  the  higher  animals  possess  memory, 

**  ^Ir.   Hookham,  in   a  lettei-  to  *^  'Conferences    sui-    la    Tli(^orie 

Prof.  Max  Miiller,  in  the  '  Biimiug-  Darwinienne,'  French  translat. 
ham  News,'  May  1873.  18f;9,  p.  1:52. 


84  The  Descent  of  Man.  Past  I. 

attention,  association,  and  even  some  imagination  and  reason. 
If  these  powers,  which  differ  much  in  different  animals,  are 
capable  of  improvement,  there  seems  no  great  improbability  in 
more  comj^lex  faculties,  such  as  the  higher  forms  of  abstraction, 
and  self-consciousness,  &c.,  having  been  evolved  through  the 
development  and  combination  of  the  simpler  ones.  It  has  been 
urged  against  the  views  here  maintained,  that  it  is  impossible 
to  say  at  what  point  in  the  ascending  scale  animals  become 
capable  of  abstraction,  &c. ;  but  who  can  say  at  what  age  this 
occurs  in  our  young  children  ?  We  see  at  least  that  such  powers 
are  developed  in  children  by  imperceptible  degrees. 

That  animals  retain  their  mental  individuality  is  unquestion- 
able. When  my  voice  awakened  a  train  of  old  associations  in 
the  mind  of  the  before-mentioned  dog,  he  must  have  retained 
his  mental  individuality,  although  every  atom  of  his  brain  had 
probably  undergone  change  more  than  once  during  the  interval 
of  five  years.  This  dog  might  have  brought  forward  the 
argument  lately  advanced  to  crush  all  evolutionists,  and  said, 
"  I  abide  amid  all  mental  moods  and  all  material  changes.  .  .  . 
"  The  teaching  that  atoms  leave  their  impressions  as  legacies  to 
''  other  atoms  falling  into  the  places  they  have  vacated  is  con- 
"  tradictory  of  the  utterance  of  consciousness,  and  is  therefore 
''  false  ;  but  it  is  the  teaching  necessitated  by  evolutionism,  con- 
''  sequently  the  hypothesis  is  a  false  one."'^*^ 

Language. — This  faculty  has  justly  been  considered  as  one  of 
the  chief  distinctions  between  man  and  the  lower  animals.  But 
man,  as  a  highly  competent  judge,  Archbishop  Whately  remarks, 
"  is  not  the  only  animal  that  can  make  use  of  language  to  express 
"  what  is  passing  in  his  mind,  and  can  understand,  more  or  less, 
"what  is  so  expressed  by  another."^'  In  Paraguay  the  Cebus, 
azaroi  when  excited  utters  at  least  six  distinct  sounds,  which 
excite  in  other  monkeys  similar  emotions."^^  The  movements  of 
the  features  and  gestures  of  monkeys  are  understood  by  us,  and 
they  joartly  understand  ours,  as  Pengger  and  others  declare.  It 
is  a  more  remarkable  fact  that  the  dog,  since  being  domesticated, 
has  learnt  to  bark^^  in  at  least  four  or  five  distinct  tones. 
Although  barking  is  a  new  art,  no  doubt  the  wild  parent-species 
of  the  dog  expressed  their  feelings  by  cries  of  various  kinds. 
"With  the  domesticated  dog  we  have  the  bark  of  eagerness,  as  in 
the  chase ;  that  of  anger,  as  well  as  growling  ;  the  yelp  or  howl  of 
despair,  as  when  shut  up  ;  the  baying  at  night ;  the  bark  of  joy,  as 

*^  The  Rev.  Dr.  J.  M'Canu,  '  Auti-  "^  Rengger,  ibid.  s.  45. 

Darwinism,' 1869,  p.  13.  ^^  See    mj    'Variation    of    Ani- 

*''  Quoted  in  'Anthropological  Re-  mak-    and  Plants  under  Domestica- 

view  '  18.'34,  p.  158.  tion,'  vol.  i.  p.  27. 


Ch ap.  III.  Mental  Powers.  8  5 

when  starting  on  a  walk  with  his  master ;  and  the  very  distinct 
one  of  demand  or  snppheation,  as  when  wishing  for  a  door  or 
window  to  be  opened.  According  to  Houzean,  who  paid  par- 
ticular attention  to  the  subject,  the  domestic  fowl  utters  at  least 
a  dozen  significant  sounds.^*^ 

The  habitual  use  of  articulate  language  is,  however,  iieculiar 
to  man ;  but  he  uses,  in  common  with  the  lower  animals,  inarti- 
culate cries  to  express  his  meaning,  aided  by  gestures  and  the 
movements  of  the  muscles  of  the  face/'^  This  especially  holds 
good  with  the  more  simple  and  vivid  feelings,  which  are  but 
little  connected  with  our  higher  intelligence.  Our  cries  of  pain, 
fear,  surprise,  anger,  together  with  tlieir  appropriate  actionS; 
and  the  murmur  of  a  mother  to  her  beloved  child,  are  more 
expressive  than  any  words.  That  which  distinguishes  man 
from  the  lower  animals  is  not  the  understanding  of  articulate 
sounds,  for,  as  every  one  knows,  dogs  understand  many  words 
and  sentences.  In  this  respect  they  are  at  the  same  stage  of 
development  as  infants,  between  the  ages  of  ten  and  twelve 
months,  who  understand  many  words  and  short  sentences,  but 
cannot  yet  utter  a  single  word.  It  is  not  the  mere  articulation 
which  is  our  distinguishing  character,  for  parrots  and  other 
birds  possess  this  power.  Nor  is  it  the  mere  capacity  of  con- 
necting definite  sounds  with  definite  ideas ;  for  it  is  certain  that 
some  parrots,  which  have  been  taught  to  speak,  connect  un- 
erringly words  with  things,  and  persons  with  events.^^  The 
lower  animals  differ  from  man  solely  in  his  almost  infinitely 
larger    power    of   associating    together    the    most    diversified 

*•  'Facultes   Meutales    des   Ani-  to  add   to  the  "good  morning"    a 

maux,'  torn.  ii.  1872,  p.  346-349.  short    sentence,    which    was    nevcM- 

^^  See  a  discussion  on  this  subject  once     repeated    after     his    father's 

in  Mr.  E.  B.  Tylor's  very  interesting  death.       He     scolded     violently     a 

work,    '  Researches    into    the  Early  strange   dog  which    came    into   the 

History  of  Mankind,'   18(35,   chaps.  room    through    the    open   window; 

ii.  to  iv.  and  he  scolded  another  parrot  (say- 

"  I  have  received  several  detailed  ing   "you    naughty  polly  ")    which 

accounts    to    this    effect.     Admiral  had  got  out   of  its    cage,  and  was 

Sir  J.  Sulivan,  whom  I  know  to  be  eating  apples  on  the  kitchen  table, 

a  careful  observer,  assures  me  that  See  also,  to  the  same  elfect,  Houzeau 

an  African  parrot,  long  kept  in  his  on     parrots,    '  Facultes     Mentales,' 

lather's     house,    invariably    called  torn,  ii,  p.  309.     Dr.  A.  Moschkau 

certain  persons  of  the  household,  as  informs  me  that  he  knew  a  starling 

well  as  visitors,  by  their  names.     He  which    never    made    a   mistake    in 

said  "good  morning"  to  every  one  at  saying  in  German  "  good  morning  " 

breakfast,  and  "good  night "  to  each  to    persons    arriving,    and    "  good- 

as  they  left  the  room  at  night,  and  bye,  old  follow,"  to  those  departing, 

never    reversed    these    salutations.  I    could    add    several    other    such 

To  Sir  J.  Sulivan's  father,  he   used  cases. 


86  TJie  Descent  of  Man.  Paet  L 


sounds  and  ideas;  and  this  obviously  depends  on  the  high 
development  of  his  mental  powers. 

As  Home  Took,  one  of  the  founders  of  the  noble  science  of 
philology,  observes,  language  is  an  art,  like  brewing  or  baking  ; 
but  writing  would  have  been  a  better  simile.  It  certainly  is  not 
a  true  instinct,  for  every  language  has  to  be  learnt.  It  differs, 
however,  widely  from  all  ordinary  arts,  for  man  has  an  in- 
stinctive tendency  to  speak,  as  we  see  in  the  babble  of  our 
young  children ;  whilst  no  child  has  an  instinctive  tendency  to 
brew,  bake,  or  write.  Moreover,  no  philologist  now  supposes 
that  any  language  has  been  deliberately  invented ;  it  has  been 
slowly  and  unconsciously  developed  by  many  steps.^^  The 
sounds  uttered  by  birds  offer  in  several  respects  the  nearest 
analogy  to  language,  for  all  the  members  of  the  same  species  utter 
the  same  instinctive  cries  expressive  of  their  emotions ;  and  all 
the  kinds  which  sing,  exert  their  power  instinctively;  but  the 
actual  song,  and  even  the  call  notes,  are  learnt  from  their 
parents  or  foster-imrcnts.  These  sounds,  as  Daines  Barrington^* 
has  proved,  "  are  no  more  innate  than  language  is  in  man." 
The  first  attempts  to  sing  "  may  be  compared  to  the  imperfect 
"  endeavour  in  a  child  to  babble."  The  young  males  continue 
practising,  or  as  the  bird-catchers  say,  "  recording,"  for  ten  or 
eleven  months.  Their  first  essays  show  hardly  a  rudiment  of 
the  future  song ;  but  as  they  grow  older  we  can  perceive  what 
they  are  aiming  at ;  and  at  last  they  are  said  "  to  sing  their 
"  song  round."  Nestlings  which  have  learnt  the  song  of  a  distinct 
species,  as  with  the  canary-birds  educated  in  the  Tyrol,  teach 
and  transmit  their  new  song  to  their  offspring.  The  slight 
natural  differences  of  song  in  the  same  species  inhabiting 
different  districts  may  be  appositely  compared,  as  Barrington 
remarks,  "  to  provincial  dialects ;"  and  the  songs  of  allied, 
though  distinct  species  may  be  compared  with  the  languages  of 
distinct  races  of  man.  I  have  given  the  foregoing  details  to 
shew  that  an  instinctive  tendency  to  acquire  an  art  is  not 
peculiar  to  man. 

With  respect  to  the  origin  of  articulate  language,  after  having 
read  on  the  one  side  the  highly  interesting  works  of  Mr.  Hens- 

^^  See  some  good  remarks  on  this  "  gards   the   immediate    end    to   be 

head     by     Prof.    Whitney,    in    his  "  attained ;  unconsciously  as  regards 

'  Oriental    and    Linguistic    Studies,'  "  the  further   consequences    of  the 

1873,  p.  354.     He  observes  that  the  "  act." 

desire    of    communication    between  ^^    Hon.     Daines    Barrington     in 

man    is    the    living    force,    which,  '  Philosoph.   Transactions,'  1773,  p. 

in    the    development   of    language,  262.     See  also  Dureau  de  la  Malle, 

"  works   both    consciously  and   un-  in  *  Ann.  des.  Sc.   Nat.*  3rd  series, 

"  consciously;     consciously    as    re-  Zoolog.  torn.  x.  p.  119. 


Chap.  111.  Mental  Poivers.  87 


leigh  Wedgwood,  the  Eev.  F.  Farrar,  and  Prof.  Schleicher,^^  and 
the  celebrated  lectures  of  Prof.  Max  Mtiller  on  the  other  side,  I 
cannot  doubt  that  language  owes  its  origin  to  the  imitation  and 
modification  of  various  natural  sounds,  the  voices  of  other 
animals,  and  man's  own  instinctive  cries,  aided  by  signs  and 
gestures.  AVhen  we  treat  of  sexual  selection  we  shall  sec  that 
primeval  man,  or  rather  some  early  progenitor  of  man,  probably 
first  used  his  voice  in  producing  true  musical  cadences,  that  is 
in  singing,  as  do  some  of  the  gibbon-apes  at  the  present  day ; 
and  we  may  conclude  from  a  widely-spread  analogy,  that  this 
power  would  have  been  especially  exerted  during  the  courtship 
of  the  sexes, — would  have  expressed  various  emotions,  such  as 
love,  jealousy,  triumph, — and  would  have  served  as  a  challenge  to 
rivals.  It  is,  therefore,  probable  that  the  imitation  of  musical 
cries  by  articulate  sounds  may  have  given  rise  to  words  expres- 
sive of  various  complex  emotions.  The  strong  tendency  in  our 
nearest  allies,  the  monkeys,  in  microcephalous  idiots,"^'^  and  in 
the  barbarous  races  of  mankind,  to  imitate  whatever  they  hear 
deserves  notice,  as  bearing  on  the  subject  of  imitation.  Since 
monkeys  certainly  understand  much  that  is  said  to  them  by 
man,  and  when  wild,  utter  signal-cries  of  danger  to  their 
fellows ;  ^^  and  since  fowls  give  distinct  warnings  for  danger  on 
the  gi-ound,  or  in  the  sky  from  hawks  (both,  as  well  as  a  third 
cry,  intelligible  to  dogs),^*^  may  not  some  unusually  wise  ape-like 
animal  have  imitated  the  growl  of  a  beast  of  prey,  and  thus 
told  his  fellow-monkeys  the  nature  of  the  expected  danger  ?  This 
would  have  been  a  first  step  in  the  formation  of  a  language. 

As  the  voice  was  used  more  and  more,  the  vocal  organs  would 
have  been  strengthened  and  perfected  through  the  principle  of 
the  inherited  effects  of  use ;  and  this  would  have  reacted  on  the 
power  of  speech.  But  the  relation  between  the  continued  use  of 
language  and  the  development  of  the  brain,  has  no  doubt  been 
far  more  important.  The  mental  powers  in  some  early  pro- 
genitor of  man  must  have  been  more  highly  developed  than  in 

"  <0n  the  Origin  of  Language,'  ^e  y^g^^  *  Memoire  sur  les  Micro- 

by  H.  Wedgwood,  18GG.     'Chapters  cephales,'   1867,  p.   1(39.     With  re- 

on   Language,'    by   the   Rev.  F.   W.  spect  to  savages,  I  have  given  some 

Farrar,     1865.      These    works    are  facts  in  my  '  Jouru.il  of  llesearches,' 

most  interesting.     See  also  '  De    la  &c.,  1845/p.  206. 
Phys.    et     de    i'arole,'    par    Albert  ^7  g^g  ^.\^,.^y^.  evidence  on  this  head 

Lemoine,  1865,  p.   190.     The  work  in   the  two  works   so  often   quoted, 

on    this    subject,  by  the    Lite  Prot".  by  Brehm  and  Ilunggei-. 
Aug.  Schleicher,  has  been  translated  ^^  Houzeau   gives  a  very  curious 

by  Dr.  Bikkers  into  English,  under  account  of  his  oljservations  on  *liis 

the   title  of  '  Darwinism   tested    by  subject    in    liis   '  Facultes    Meutales 

the  Science  of  Language,'  1869  dcs  Aniniaux,'  toni.  ii.,  p.  348. 


88  The  Descent  of  Man.  Part  I. 

any  existing  ape,  before  even  the  most  imperfect  form  of  speech 
could  have  come  into  use ;  but  we  may  confidently  believe  that 
the  continued  use  and  advancement  of  this  power  Avould  have 
reacted  on  the  mind  itself,  by  enabling  and  encouraging  it  to 
carry  on  long  trains  of  thought.  A  complex  train  of  thought 
can  no  more  be  carried  on  without  the  aid  of  words,  whether 
spoken  or  silent,  than  a  long  calculation  without  the  use  of 
figures  or  algebra.  It  ajopears,  also,  that  even  an  ordinary  train 
of  thought  almost  requires,  or  is  greatly  facilitated  by  some 
form  of  language,  for  the  dumb,  deaf,  and  blind  girl,  Laura 
Bridgman,  was  observed  to  use  her  fingers  whilst  dreaming.^^ 
Nevertheless,  a  long  succession  of  vivid  and  connected  ideas  may 
pass  through  the  mind  without  the  aid  of  any  form  of  language, 
as  we  may  infer  from  the  movements  of  dogs  during  their 
dreams.  We  have,  also,  seen  that  animals  are  able  to  reason 
to  a  certain  extent,  manifestly  without  the  aid  of  language. 
The  intimate  connection  between  the  brain,  as  it  is  now 
developed  in  us,  and  the  faculty  of  speech,  is  well  shewn  by 
those  curious  cases  of  brain-disease  in  which  speech  is  specially 
affected,  as  when  the  power  to  remember  substantives  is  lost, 
whilst  other  words  can  be  correctly  used,  or  where  substantives 
of  a  certain  class,  or  all  except  the  initial  letters  of  substantives 
and  i^roper  names  are  forgotten.^  There  is  no  more  improb- 
ability in  the  continued  use  of  the  mental  and  vocal  organs 
leading  to  inherited  changes  in  their  structure  and  functions, 
than  in  the  case  of  handwriting,  which  depends  partly  on  the 
form  of  the  hand  and  partly  on  the  disposition  of  the  mind ;  and 
hand- writing  is  certainly  inherited.''^ 

Several  writers,  more  especially  Prof.  Max  Muller,^^  have 
lately  insisted  that  the  use  of  language  implies  the  power  of 
forming  general  concepts ;  and  that  as  no  animals  are  supposed 
to  possess  this  power,  an  impossible  barrier  is  formed  between 
them  and  man.*^^     With   respect  to  animals,  I  have  already 

53  See  remarks  on   tins   head   by  ^-  Lectures  on 'Mr.  Darwin's  Phi- 

Dr.  Maudsley,  'The  Physiology  and  losophy  of  Language,'  1873. 

Pathology  of  Mind,'  2ud  edit.  1868,  ^'^    The    judgment    of    a    distin- 

p.  199.  guished    philologist,    such    as    Prof. 

^^  Many  curious  cases  have  been  Whitney,  will  have  far  more  weight 

recorded.     See,    for     instance,     Dr.  on    this    point  than    anything    that 

Bateman 'On  Aphasia,' 1870,  p.  27,  I  can  say.     He  remarks  ('Oriental 

31,  53,    100,  &c.     Also,   'Inquiries  and    Linguistic    Studies,'    1873,    p.. 

Concei-ning  the  Intellectual  Powers,'  297),  in  speaking  of  Bleek's  views  : 

by  Dr.  Abercrombie,  1838,  p.  150.  "Because   on  the    grand  scale  lan- 

^^    'The    Variation    of    Animals  "  guage  is  the   necessary  auxiliary 

and    Plants    under   Domestication,'  "  of  thought,  indispensable   to   tlie 

vol.  ii.  p.  6.  "  development     of     the     power    of 


Chap.  II  r.  Mental  Pozvc7'S,  89 

endeavoured  to  show  that  they  have  this  power,  at  least  in  a 
riule  and  incipient  degree.  As  for  as  concerns  infants  of  from 
ten  to  eleven  months  old,  and  deaf-mutes,  it  seems  to  mo  in- 
credible, that  they  should  be  able  to  connect  certain  sounds  with 
certain  general  ideas  as  quickly  as  they  do,  unless  such  ideas 
were  already  formed  in  their  minds.  The  same  remark  may  be 
extended  to  the  more  intelligent  animals ;  as  Mr.  Leslie  Stephen 
observes,'^'*  "  A  dog  frames  a  general  concept  of  cats  or  sheep, 
"  and  knows  the  corresponding  words  as  well  as  a  philosopher. 
"  And  the  capacity  to  understand  is  as  good  a  j^roof  of  vocal 
"  intelligence,  though  in  an  inferior  degree,  as  the  capacity  to 
"  speak> 

Why  the  organs  now  used  for  speech  should  have  been 
originally  perfected  for  this  purpose,  rather  than  any  other 
organs,  it  is  not  difficult  to  see.  Ants  have  considerable  powers 
of  intercommunication  by  means  of  their  antennjc,  as  shewn  by 
Huber,  who  devotes  a  whole  chapter  to  their  language.  We 
might  have  used  our  fingers  as  efficient  instruments,  for  a 
person  with  practice  can  report  to  a  deaf  man  every  word  of  a 
speech  rapidly  delivered  at  a  public  meeting;  but  the  loss  of 
our  hands,  whilst  thus  employed,  would  have  been  a  serious 
inconvenience.  As  all  the  higher  mammals  possess  vocal  organs, 
constructed  on  the  same  general  plan  as  ours,  and  used  as  a 
means  of  communication,  it  was  obviously  probable  that  these 
same  organs  would  be  still  further  developed  if  the  power  of 
communication  had  to  be  improved ;  and  this  has  been  cftected  by 
the  aid  of  adjoining  and  well  adapted  parts,  namely  the  tongue 
and  lips.^5  The  fact  of  the  higher  apes  not  using  their  vocal 
organs  for  speech,  no  doubt  depends  on  their  intelligence  not 
having  been  sufficiently  advanced.    The  possession  by  them  of 


"  thinking,  to  the  distinctness  and  "  fingers  into    imitation    of  spoken 

"  variety  and  complexity  of  cogni-  "  words."      Jlax    Miillcr    gives    in 

"  tions  to  the  full  mastery  of  con-  italics  ('Lectures    on  ]\Jr.  Darwin's 

'•  sciousness ;    therefore    he    woukl  Philosophy     of     Language,'     187.'>, 

"  fain  make  thought  absolutely  im-  third  lecture)  the  following  aphor- 

"  possible  without  speech,  identify-  ism:    "There  is    no   thought  with- 

"  ing   the  faculty  with  its    instru-  "  out  words,  as  little   as  tiiere  are 

"  ment.     He  might  just  as  reason-  "  words  without    thought."     What 

"  ably  assert  that  the  liuman  hand  a   strange    definition  must   here  be 

"  cannot  act  without  a  tool.     With  gi\-cn  to  the  word  thought ! 

"  such  a  doctrine  to  start  from,  he  ^^  '  Essays  on  Free-thinking,'  tS;c., 

"  cannot    stop    short    of     IMiiiler's  1873,  p.  82. 

"  worst    paradoxes,  that    an  infant  ^^  See  some  ggod  remarks  to  this 

"  (m  /«»5,  not   speaking)  is  not    a  effect  by  Dr.  Maudslcy,  '  The  I'hy- 

•*  human  being,  and  that  deaf-mutes  siology    and     Pathology    of    Mind,' 

"  do  not  become  possessed  of  reason  18G8,  p.  l'J9. 
"  until    tliey  learn    to    twist    their 


90  '  The  Descent  of  Man.  Part  1. 

organs,  wliicli  with  long- continued  practice  might  have  been 
used  for  speech,  although  not  thus  used,  is  paralleled  by  the 
case  of  many  birds  which  possess  organs  fitted  for  singing, 
though  they  never  sing.  Thus,  the  nightingale  and  crow  have 
vocal  organs  similarly  constructed,  these  being  used  by  the 
former  for  diversified  song,  and  by  the  latter  only  for  croaking.*^' 
If  it  be  asked  why  apes  have  not  had  their  intellects  developed 
to  the  same  degree  as  that  of  man,  general  causes  only  can  be 
assigned  in  answer,  and  it  is  unreasonable  to  expect  anything 
more  definite,  considering  our  ignorance  with  respect  to  the 
successive  stages  of  development  through  which  each  creature 
has  passed.  • 

The  formation  of  different  languages  and  of  distinct  species, 
and  the  proofs  that  both  have  been  developed  through  a  gradual 
process,  are  curiously  parallel.^^  But  we  can  trace  the  formation 
of  many  words  further  back  than  that  of  species,  for  we  can 
perceive  how  they  actually  arose  from  the  imitation  of  various 
sounds.  We  find  in  distinct  languages  striking  homologies  duo 
to  community  of  descent,  and  analogies  due  to  a  similar  process 
of  formation.  The  manner  in  which  certain  letters  or  sounds 
change  when  others  change  is  very  like  correlated  growth.  We 
have  in  both  cases  the  reduplication  of  parts,  the  effects  of  long- 
continued  use,  and  so  forth.  The  frequent  presence  of  rudi- 
ments, both  in  languages  and  in  species,  is  still  more  remarkable. 
The  letter  m  in  the  word  am,  means  / ;  so  that  in  the  expres- 
sion /  am,  a  superfluous  and  useless  rudiment  has  been  retained. 
In  the  spelling  also  of  words,  letters  often  remain  as  the  rudi- 
ments of  ancient  forms  of  pronunciation.  Languages,  like 
organic  beings,  can  be  classed  in  groups  under  groups ;  and  they 
can  be  classed  either  naturally  according  to  descent,  or  arti- 
ficially by  other  characters.  Dominant  languages  and  dialects 
spread  widely,  and  lead  to  the  gradual  extinction  of  other^ 
tongues.  A  language,  like  a  species,  when  once  extinct,  never, 
as  Sir  C.  Lyell  remarks,  reappears.  The  same  language  never 
has  two  birth-places.  Distinct  languages  may  be  crossed  or 
blended  together.''^   We  see  variability  in  every  tongue,  and  new 

^^  Macgillivray,  '  Hist,  of  British  display   any   -anusual    capacity   for 

Birds,'   A'ol.    ii.    1839,    p.    29.      An  imitation.     '  Researches  in  Zoology,' 

excellent   observer,  Mr.  Blackwall,  1834,  p.  158. 

remarks  that  the  magpie  learns  to  ''^  See  the  very  interesting  pa- 
pronounce  single  words,  and  even  rallelism  between  the  development 
short  sentences,  more  readily  than  of  species  and  languages,  given  by 
almost  any  other  British  bird ;  yet,  Sir  C.  Lyell  in  '  The  Geolog.  Evi- 
as  he  adds,  after  long  and  closely  dences  of  the  Antiquity  of  Man,' 
investigating  its  habits,  he  has  1863,  chap,  xxiii. 
Lever  known  it,  in  a  state  of  nature,  ^^  See  remarks  to  this  effect  by 


CiLvp.  III.  Mental  Powers,    •  91 

words  are  continually  cropping  np ;  but  as  there  is  a  limit  to 
the  powers  of  the  memory,  single  words,  like  whole  languages, 
gradually  become  extinct.  As  Max  Miiller'^^  has  well  re- 
marked : — '''  A  struggle  for  life  is  constantly  going  on  amongst 
*Hhe  words  and  grammatical  forms  in  each  language.  The 
"  better,  the  shorter,  the  easier  forms  are  constantly  gaining  the 
"  upper  hand,  and  they  owe  their  success  to  their  own  inherent 
"  virtue."  To  these  more  important  causes  of  the  survival  of 
certain  words,  mere  novelty  and  fashion  may  be  added ;  for 
there  is  in  the  mind  of  man  a  strong  love  for  slight  changes  in  all 
things.  The  survival  or  preservation  of  certain  favoured  words 
in  the  struggle  for  existence  is  natural  selection. 

The  perfectly  regular  and  wonderfully  complex  construction 
of  the  languages  of  many  barbarous  nations  has  often  been 
advanced  as  a  proof,  either  of  the  divine  origin  of  these  lan- 
guages, or  of  the  high  art  and  former  civilisation  of  their 
founders.  Thus  F.  von  Schlegel  writes :  "  In  those  languages 
"  which  appear  to  be  at  the  lowest  grade  of  intellectual  culture, 
*'  we  frequently  observe  a  very  high  and  elaborate  degree  of  art 
"in  their  gi-ammatical  structure.  This  is  especially  the  case 
"  with  the  Basque  and  the  Lapponian,  and  many  of  the  Ame- 
"  rican  languages.'"''®  But  it  is  assuredly  an  error  to  speak  of 
any  language  as  an  art,  in  the  sense  of  its  having  been  elabor- 
ately and  methodically  formed.  Philologists  now  admit  that 
conjugations,  declensions,  &c.,  originally  existed  as  distinct 
words,  since  joined  together;  and  as  such  words  express  the 
most  obvious  relations  between  objects  and  persons,  it  is  not 
surprising  that  they  should  have  been  used  by  the  men  of  most 
races  during  the  earliest  ages.  With  respect  to  perfection,  the 
following  illustration  will  best  shew  how  easily  we  may  err :  a 
Crinoid  sometimes  consists  of  no  less  than  150,000  pieces  of 
shell,'^  all  arranged  with  i^erfect  symmetry  in  radiating  lines ; 
but  a  naturalist  does  not  consider  an  animal  of  tliis  kind  as 
more  perfect  than  a  bilateral  one  with  comparatively  few  jDarts, 
and  with  none  of  these  parts  alike,  excepting  on  the  opposite  sides 
of  the  body.  He  justly  considers  the  differentiation  and  special- 
isation of  organs  as  the  test  of  perfection.  So  with  languages  ; 
the  most  symmetrical  and  complex  ought  not  to  be  ranked  above 
irregular,  abbreviated,  and  bastardised  languages,  which  have 


the    Rev.  F.  W.  Fanar,  in    an    in-  '«  Quoted  by  C.  S.  Wake,  '  Cliap- 

terostiiig     article,    eutitleJ    '  Philo-  ters  on  Mau,'  1868,  p.  101, 
logy  and  Darwinism'  in  'Nature,'  ''   Buckland,   'Bridgewatcr  Trca- 

March  2-tth,  1870,  p.  528.  ti.se.'  p.  411. 
«3  '  Nature,'  Jan.  tith,  1870,  p.  2:)7. 


92  The  Descent  of  Man.  Part  I. 

borrowed  expressive  words  and  useful  forms  of  construction  from 
various  conquering,  conquered,  or  immigrant  races. 

From  these  few  and  imperfect  remarlis  I  conclude  tliat  the 
extremely  complex  and  regular  construction  of  many  barbarous 
languages,  is  no  proof  that  they  owe  their  origin  to  a  special 
act  of  creation.'^^  Nor,  as  we  have  seen,  does  the  faculty  of 
articulate  speech  in  itself  offer  any  insuperable  objection  to 
the  belief  that  man  has  been  developed  from  some  lower 
form. 

Bense  of  Beauty. — This  sense  has  been  declared  to  be  peculiar 
to  man.  I  refer  here  only  to  the  pleasure  given  by  certain 
colours,  forms,  and  sounds,  and  which  may  fairly  be  called  a 
sense  of  the  beautiful ;  with  cultivated  men  such  sensations  arc, 
however,  intimately  associated  with  complex  ideas  and  trains  of 
thought.  When  we  behold  a  male  bird  elaborately  displaying 
his  graceful  plumes  or  splendid  colours ,  before  the  female, 
whilst  other  birds,  not  thus  decorated,  make  no  such  display, 
it  is  impossible  to  doubt  that  she  admires  the  beauty  of  her 
male  partner.  As  women  everywhere  deck  themselves  with 
these  plumes,  the  beauty  of  such  ornaments  cannot  be  disputed. 
As  we  shall  see  later,  the  nests  of  humming-birds,  and  the 
playing  passages  of  bower-birds  are  tastefully  ornamented 
with  gaily-coloured  objects;  and  this  shews  that  they  must 
receive  some  kind  of  pleasure  from  the  sight  of  such  things. 
With  the  great  majority  of  animals,  however,  the  taste  for  the 
beautiful  is  confined,  as  far  as  we  can  judge,  to  the  attractions 
of  the  opposite  sex.  The  sweet  strains  poured  forth  by  many 
male  birds  during  the  season  of  love,  are  certainly  admired  by 
the  females,  of  which  fact  evidence  will  hereafter  be  given.  If 
female  birds  had  been  incapable  of  appreciating  the  beautiful 
colours,  the  ornaments,  and  voices  of  their  male  partners,  all  the 
labour  and  anxiety  exhibited  by  the  latter  in  disi^laying  their 
charms  before  the  females  would  have  been  thrown  away ;  and 
this  it  is  impossible  to  admit.  Why  certain  bright  .colours 
should  excite  pleasure  cannot,  I  presume,  be  explained,  any 
more  than  why  certain  flavours  and  scents  are  agreeable ;  but 
habit  has  something  to  do  with  the  result,  for  that  which  is  at 
first  unpleasant  to  our  senses,' ultimately  becomes  pleasant,  and 
habits  are  inherited.  With  respect  to  sounds,  Helmholtz  has 
explained  to  a  certain  extent  on  physiological  principles,  why 
harmonies  and  certain  cadences  are  agreeable.  But  besides 
this,  sounds   frequently  recurring  at  irregular   intervals   are 

''-  Soe  some  gooil  remarks  on  the  J.  Lubbock,  'Origin  of  Civilisatiou,* 
simplification  of   languages^  by  Sir      1870,  p.  278. 


CuAp.  111.  Mental  Powers,  93 

highly  disagreeable,  as  every  one  will  admit  who  has  listened  at 
night  to  the  irregular  flapping  of  a  rope  on  board  ship.  The 
same  princii^le  seems  to  come  into  play  with  vision,  as  the 
eye  prefers  symmetry  or  figures  with  some  regular  recurrence. 
Patterns  of  this  kind  are  employed  by  even  the  lowest  savages 
as  ornaments;  and  they  have  been  developed  through  sexual 
selection  for  the  adornment  of  some  male  animals.  Whether  we 
can  or  not  give  any  reason  for  the  pleasure  thus  derived  from 
vision  and  hearing,  yet  man  and  many  of  the  lower  animals  are 
alike  pleased  by  the  same  colours,  graceful  shading  and  forms, 
and  the  same  sounds. 

The  taste  for  the  beautiful,  at  least  as  far  as  female  beauty  is 
concerned,  is  not  of  a  special  nature  in  the  human  mind ;  for  it 
differs  widely  in  the  different  races  of  man,  and  is  not  quite  the 
same  even  in  the  different  nations  of  the  same  race.  Judging 
from  the  hideous  ornaments,  and  the  equally  hideous  music 
admired  by  most  savages,  it  might  be  urged  that  their  jesthetic 
faculty  was  not  so  highly  developed  as  in  certain  animals,  for 
instance,  as  in  birds.  Obviously  no  animal  would  be  capable  of 
admiring  such  scenes  as  the  heavens  at  night,  a  beautiful  land- 
scape, or  refined  music;  but  such  high  tastes  are  acquired 
through  culture,  and  depend  on  complex  associations ;  they  are 
not  enjoyed  by  barbarians  or  by  uneducated  persons. 

Many  of  the  faculties,  which  have  been  of  inestimable  service 
to  man  for  his  progressive  advancement,  such  as  the  powers  of 
the  imagination,  wonder,  curiosity,  an  undefined  sense  of  beauty, 
a  tendency  to  imitation,  and  the  love  of  excitement  or  novelty, 
could  hardly  fail  to  lead  to  capricious  changes  of  customs  and 
fashions.  I  have  alluded  to  this  point,  because  a  recent  writer  '^ 
has  oddly  fixed  on  Caprice  "  as  one  of  the  most  remarkable  and 
"typical  differences  between  savages  and  brutes."  But  not 
only  can  we  partially  understand  how  it  is  that  man  is  from 
various  conflicting  influences  rendered  capricious,  but  that 
the  lower  animals  are,  as  we  shall  hereafter  see,  likewise  capri- 
cious in  their  affections,  aversions,  and  sense  of  beauty.  There 
is  also  reason  to  suspect  that  they  love  novelty,  for  it  own  sake. 

Bdief  in  God — BeJiyirm. — There  is  no  evidence  that  man  was 
aboriginally  endowed  with  the  ennobling  belief  in  the  existence 
of  an  Omnipotent  God.  On  the  contrary  there  is  ample  evidence, 
derived  not  from  hasty  travellers,  but  from  men  who  have  long 
resided  with  savages,  that  numerous  races  have  existed,  and  still 
exist,  who  have  no  idea  of  one  or  more  gods,  and  who  have  no 

"  'The  Spectator,'  Dec.  -tth,  1SG9,  p.  1430. 


94  The  Descent  of  Man.  Part  I. 

words  in  their  languages  to  express  snch  an  idea/*    The  question 

is  of  course  wholly  distinct  from  that  higher  one,  whether  there 
exists  a  Creator  and  Euler  of  the  universe ;  and  this  has  been 
answered  in  the  affirmative  by  some  of  the  highest  intellects  that 
have  ever  existed. 

If,  however,  we  include  under  the  term  "  religion ''  the  belief 
in  miseen  or  spiritual  agencies,  the  case  is  wholly  different ;  for 
this  belief  seems  to  be  universal  with  the  less  civilised  races. 
Nor  is  it  difficult  to  comprehend  how  it  arose.  As  soon  as  the 
important  faculties  of  the  imagination,  wonder,  and  curiosity, 
together  with  some  power  of  reasoning,  had  become  partially 
developed,  man  would  naturally  crave  to  understand  what  was 
passing  around  him,  and  would  have  vaguely  speculated  on  his 
own  existence.  As  Mr.  M'Lennan'^  has  remarked,  "  Some  explan- 
"  ation  of  the  phenomena  of  life,  a  man  must  feign  for  himself; 
"  and  to  judge  from  the  universality  of  it,  the  simplest  hypothesis, 
"  and  the  first  to  occur  to  men,  seems  to  have  been  that  natural 
"  phenomena  are  ascribable  to  the  presence  in  animals,  plants, 
"  and  things,  and  in  the  forces  of  nature,  of  such  spirits  prompting 
"  to  action  as  men  are  conscious  they  themselves  possess."  It 
is  also  probable,  as  Mr.  Tylor  has  shewn,  that  dreams  may  have 
first  given  rise  to  the  notion  of  spirits ;  for  savages  do  not  readily 
distinguish  between  subjective  and  objective  impressions.  When 
•a  savage  dreams,  the  figures  which  appear  before  him  arc 
believed  to  have  come  from  a  distance,  and  to  stand  over  him ; 
or  "  the  soul  of  the  dreamer  goes  out  on  its  travels,  and  comes 
"  home  with  a  remembrance  of  what  it  has  seen."^^"    But  until 

^■*  See  an  excellent  article  on  this  forms  of  religious  belief  throughout 
subject  by  the  Re\^.  F.  W.  Farrar,  the  world,  by  man  being  led  through 
in  the  'Anthropological  Review,'  dreams,  shadows,  and  other  causes, 
Aug.  1864,  p.  ccxvii.  For  further  to  look  at  himself  as  a  double 
facts  see  Sir  J.  Lubbock,  '  Pre-  essence,  corporeal  and  spiritual.  As 
historic  Times,'  2nd  edit.  1869,  p.  the  spiritual  being  is  supposed  to 
.564 ;  and  especially  the  chapters  on  exist  after  death  and  to  be  power- 
Religion  in  his  'Origin  of  Civilisa-  ful,  it  is  propitiated  by  various  gifts 
tion,'  1870.  and  ceremonies,  and  its  aid  involved. 

"''"  '  The  Worship  of  Animals  and  He  then  further  shews  that  names 

Plants,'  in  the  '  Fortnightly  Review,'  or    nicknames     given     from     some 

Oct.  1,  1869,  p.  422.  animal  or  other  object,  to  the  early 

"^  Tylor,  '  Early  History  of  Man-  progenitors  or  founders  of  a  tribe, 

kind,'    1865,    p.    6.     See    also    the  arc  supposed  after  a  long   interval 

three  striking  chapters  on  the  De-  to  represent  the  real  progenitor  of 

velopment  of  Religion,  in  Lubbock's  the  tribe;  and  such  animal  or  object 

'  Origin  of  Civilisation,'  1870.     In  a  is   then   naturally  believed  still  to 

like    manner    Mr,  Herbert  Spencer,  exist  as  a  spirit,  is  held  sacred,  and 

in  his  ingenious  essay  in  the  '  Fort-  worshipped  as  a  god.     Nevertheless 

nightly    Review'   (May    1st,    1870,  I  cannot  but  suspect  that  there  is 

p.    535),    accounts   for    the  earliest  a  still  earlier  and  ruder  stage,  when 


Chap.  II  f.  Mental  Powers,  95 

the  faculties  of  imagination,  curiosity,  reason,  &c.,  had  been 
fairly  well  developed  in  the  mind  of  man,  his  dreams  would  not 
have  led  him  to  believe  in  s^Dirits,  any  more  than  in  the  case  of 
a  dog. 

The  tendency  in  savages  to  imagine  that  natural  objects  and 
agencies  are  animated  by  spiritual  or  living  essences,  is  perha]')s 
illustrated  by  a  little  fact  which  I  once  noticed  :  my  dog,  a  full- 
grown  and  very  sensible  animal,  was  lying  on  the  lawn  during  a 
liot  and  still  day ;  but  at  a  little  distance  a  slight  breeze  occa- 
sionally moved  an  open  parasol,  which  would  have  been  wholly 
disregarded  by  the  dog,  had  any  one  stood  near  it.  As  it  was, 
every  time  that  the  parasol  slightly  moved,  the  dog  growled 
fiercely  and  barked.  He  must,  I  think,  have  reasoned  to  himself 
in  a  rapid  and  unconscious  manner,  that  movement  without  any' 
apparent  cause  indicated  the  presence  of  some  strange  living 
agent,  and  that  no  stranger  had  a  right  to  be  on  his  territory. 

The  belief  in  spiritual  agencies  would  easily  pass  into  the 
belief  in  the  existence  of  one  or  more  gods.  For  savages  would 
naturally  attribute  to  spirits  the  same  passions,  the  same  love  of 
vengeance  or  simplest  form  of  justice,  and  the  same  affections 
which  they  themselves  feel.  The  Fuegians  appear  to  be  in  this 
respect  in  an  intermediate  condition,  for  when  the  surgeon  on  board 
the  "Beagle"  shot  some  young  ducklings  as  specimens,  York 
Minster  declared  in  the  most  solemn  manner,  "  Oh,  Mr.  Bynoe, 
"  much  rain,  much  snow,  blow  much ;"  and  this  was  evidently 
a  retributive  punishment  for  wasting  human  food.  So  again  he 
related  how,  when  his  brother  killed  a  "  wild  man,"  storms  long 
raged,  much  rain  and  snow  fell.  Yet  we  could  never  discover 
that  the  Fuegians  believed  in  what  we  should  call  a  God,  or 
practised  any  religious  rites ;  and  Jemmy  Button,  with  justifiable 
pride,  stoutly  maintained  that  there  was  no  devil  in  his  land. 
This  latter  assertion  is  the  more  remarkable,  as  with  savages  the 
belief  in  bad  spirits  is  far  more  common  than  that  in  good 
ones. 

The  feehog  of  religious  devotion  is  a  highly  complex  one, 
consisting  of  love,  complete  submission  to  an  exalted  and 
mysterious  superior,  a  strong  sense  of  dependence,^^  fear, 
reverence,  gratitude,  hope  for  the  future,  and  perhaps  other 
elements.     No  being  could  cxiDcricnce  so  complex  an  emotion 


anything  which  manifests  power  or  ^^  See    an    able    article    on    the 

movement  is  thought  to  be  endowed  '  Physical  Elements  of  ileligion,'  by 

with  some  form  of  life,   and   with  !Mr.  L.  Owen  Pike,  in  '  Anthropolog. 

mental  faculties  analogous    to    our  Kevicw,'  April,  1870,  p.  Ixiii. 


g6  The  Desceitt  of  Man.  Tart  I, 

until  advanced  in  his  intellectual  and  moral  faculties  to  at  least 
a  moderately  high  level.  Nevertheless,  we  see  some  distant 
approach  to  this  state  of  mind  in  the  deep  love  of  a  dog  for  his 
master,  associated  with  complete  submission,  some  fear,  and 
perhaps  other  feelings.  The  behaviour  of  a  dog  when  returning 
to  his  master  after  an  absence,  and,  as  I  may  add,  of  a  monkey 
to  his  beloved  keeper,  is  widely  diiferent  from  that  towards  their 
fellows.  In  the  latter  case  the  transports  of  joy  appear  to  be 
somewhat  less,  and  the  sense  of  equality  is  shewn  in  every  action. 
Professor  Braubach  goes  so  far  as  to  maintain  that  a  dog  looks 
on  his  master  as  on  a  god.^^ 

The  same  high  mental  faculties  which  first  led  man  to  believe 
in  miseen  spiritual  agencies,  then  in  fetishism,  polytheism,  and 
ultimately  in  monotheism,  would  infallibly  lead  him,  as  long  as 
his  reasoning  powers  remained  poorly  developed,  to  various 
strange  superstitions  and  customs.  Many  of  these  are  terrible 
to  think  of — such  as  the  sacrifice  of  human  beings  to  a  blood- 
loving  god ;  the  trial  of  innocent  persons  by  the  ordeal  of  poison 
or  fire ;  witchcraft,  &c. — yet  it  is  well  occasionally  to  reflect  on 
these  superstitions,  for  they  shew  us  what  an  infinite  debt  of 
gratitude  we  owe  to  the  improvement  of  our  reason,  to  science, 
and  to  our  accumulated  knowledge.  As  Sir  J.  Lubbock''^  has  well 
observed,  "  it  is  not  too  much  to  say  that  the  horrible  dread  of 
"  unknown  evil  hangs  like  a  thick  cloud  over  savage  life,  and 
"  embitters  every  pleasure."  These  miserable  and  indirect 
consequences  of  our  highest  faculties  may  be  compared  with  the 
incidental  and  occasional  mistakes  of  the  instincts  of  the  lov/er 
animals. 

^*  '  Religion,  Moral,  &c.,  del*  Dar-  ^^  'Prehistoric  Times,'  2iid    edit, 

win'schen  Art-Lehre,'    1869,  s.  53.  p.     571.      In    this    work    (p.  571) 

It  is  said  (Dr.  W.  Lauder  Lindsay,  there    will    be    found   an    excellent 

'Journal  of  Mental  Science,'  1871,  account    of  the    many  strange  and 

p.  43),  that  Bacon  long  ago,  and  the  capricious  customs  of  savages, 
poet  Burns,  held  the  same  notion. 


Chap.  IV.  Moral  Sense.  97 


CHAPTEK  IV. 

CoMrAEisoN  OF  THE  Mental  Powers  of  Man  and  the 
Lower  Animals— co??^mz(ecZ. 

Thf  moral  sense  —  Fundamental  proposition  —  The  qualities  of  social 
animals — Origin  of  sociability — Strutrgle  between  op])osed  instincts — 
Man  a  social  animal — Tlie  more  enduring  social  instincts  conquer  otlicr 
less  persistent  instincts — The  social  virtues  alone  regarded  by  savages — 
The  self-regarding  virtues  acquired  at  a  later  stage  of  development — 
The  importance  of  the  judgment  of  the  members  of  the  same  community 
on  conduct — Transmission  of  moral  tendencies — Summary. 

I  FULLY  subscribe  to  the  judgment  of  those  writers^  who 
maintain  that  of  all  the  differences  between  man  and  the 
lower  animals,  the  moral  sense  or  conscience  is  by  far  the 
most  important.  This  sense,  as  Mackintosh^  remarks,  "  has  a 
"  rightful  supremacy  over  every  other  principle  of  human 
"  action ;"  it  is  summed  up  in  that  short  but  imperious  word 
ouglit,  so  full  of  high  significance.  It  is  the  most  noble  of  all 
the  attributes  of  man,  leading  him  without  a  moment's  hesita- 
tion to  risk  his  life  for  that  of  a  fellow-creature ;  or  after  due 
deliberation,  impelled  simply  by  the  deep  feeling  of  right  or 
duty,  to  sacrifice  it  in  some  great  cause.  Immanucl  Kant 
exclaims,  "  Duty !  Wondrous  thought,  that  workest  neither  by 
"  fond  insinuation,  flattery,  nor  by  any  threat,  but  merely  by 
"  holding  np  thy  naked  law  in  the  soul,  and  so  extorting  for 
"  thyself  always  reyerence,  if  not  always  obedience ;  before 
"  whom  all  appetites  are  dumb,  however  secretly  they  rebel ; 
•'  whence  thy  original?"^ 

This  great  question  has  been  discussed  by  many  writers^  of 
consummate  ability ;  and  my  sole  excuse  for  touching  on  it,  is 
the  impossibility  of  here  passing  it  over ;  and  because,  as  far  as  I 
know,  no  one  has  approached  it  exclusively  from  the  side  of 
natural  history.    The  investigation  possesses,  also,  some  in- 

*  See,  for  instance,  on  this  subject,  and  Moral  Science,'  18G8,  p.  543- 
Quatrefages,  '  Llnite  de  i'Espece  725)  of  twenty-six  British  authors 
Humaine,'  1861,  p.  21,  &c.  who  have  written  on  this  subject, 

-  '  Dissertation  on  Ethical   Philo-  and  whose    names    are    familiar   to 

sophy,'  1837,  p.  231,  &c.  every  reader  ;  to  these,  ^Ir.  Bain's 

'  '  Metaphysics  of  Ethics,'  trans-  own  name,  and  those  of  Mr.  Lecky, 

lated  by  J.  W.  Semple,  Edinburgh,  Mr,    Shadworth    Hodgson,    Sir    J. 

1836,  p.  136.  Lubbock,    and     others,    mi|ht     be 

*  Mr.  Bain  gives  a  list  ('  Mental  added. 


98  The  Descent  of  Man,  Paet  1. 

dependent  interest,  as  an  attempt  to  see  how  far  the  study  of 
the  lower  animals  throws  light  on  one  of  the  highest  psychical 
faculties  of  man. 

The  folio v\dng  proposition  seems  to  me  in  a  high  degree 
probable — namely,  that  any  animal  whatever,  endowed  with 
well-marked  social  instincts,^  the  parental  and  filial  affections 
being  here  included,  would  inevitably  acquire  a  moral  sense  or 
conscience,  as  soon  as  its  intellectual  powers  had  become  as  well, 
or  nearly  as  well  developed,  as  in  man.  For,  firstly,  the  social 
instincts  lead  an  animal  to  take  pleasure  in  the  society  of  its 
fellows,  to  feel  a  certain  amount  of  sympathy  with  them,  and  to 
perform  various  services  for  them.  The  services  may  be  of  a 
definite  and  evidently  instinctive  nature  ;  or  there  may  be  only 
a  wish  and  readiness,  as  with  most  of  the  higher  social  animals, 
to  aid  their  fellows  in  certain  general  ways.  But  these  feelings 
and  services  are  by  no  means  extended  to  all  the  individuals  of 
the  same  species,  only  to  those  of  the  same  association.  Secondly, 
as  soon  as  the  mental  faculties  had  become  highly  developed, 
images  of  all  past  actions  and  motives  would  be  incessantly 
passing  through  the  brain  of  each  individual ;  and  that  feeling 
of  dissatisfaction,  or  even  misery,  which  invariably  results,  as  we 
shall  hereafter  see,  from  any  unsatisfied  instinct,  would  arise, 
as  often  as  it  was  perceived  that  the  enduring  and  always 
present  social  instinct  had  yielded  to  some  other  instinct,  at  the 
time  stronger,  but  neither  enduring  in  its  nature,  nor  leaving 

^  Six-  B.  Brodie,  aftei    observing  all   this,  he  also  remarks,  "  if,  as  is 

that  man  is  a  social  animal  ('  Psy-  "  my  own  belief,  the  moral  feelings 

chological  Enquiries,'  1854,  p.  192),  "  are  not  innate,  but  acquired,  they 

asks  the  pregnant  question,  "  ought  "  are  not  for  that  reason  less  natu- 

"  not    this    to    settle    the    disputed  "  ral."     It  is  with  hesitation  that  I 

"  question  as  to  the  existence  of  a  venture    to   differ    at   all    from    so 

"  moral  sense  ?"    Similar  ideas  have  profound    a    thinker,    but    it     can 

probably  occurred  to  many  persons,  hardly  be   disputed   that   the  social 

as   they   did   long   ago    to    Marcus  feelings  are  instinctive  or  innate  in 

Aurelius.     Mr.  J.  S.  Mill  speaks,  in  the  lower  animals ;  and  why  should 

his    celebrated    work,    *  Utilitarian-  they  not  be  so  in  man  ?     Mr.  Bain 

ism,'  (1864,  pp.  45,  46),  of  the  social  (see,  for  instance,  'The  Emotions  and 

feelino-s    as    a    "powerful    natural  the  Will,'  1865,  p.  481)  and  others 

"  sentiment,"  and  as    "  the  natural  believe  that  the  moral  sense  is  ac- 

"  basis  of  sentiment  for  utilitarian  quired    by  each    individual    during 

"  morality."     Again  he  says,  "  Like  his  lifetime.     On  the  general  theory 

"  the  other  acquired  capacities  above  of    evolution    this    is    at    least    ex- 

"  referred  to,  the  moral  faculty,  if  tremely  improbable.     The  ignoring 

'•  not    a    part    of    our    nature,    is  of  all  transmitted  mental  qualities 

'•  a   natural    out-growth    from   it ;  will,  as  it  seems  to  me,  be  hereafter 

"  capable,  like    them,  in   a  certain  judged  as  a  most  serious  blemish  iu 

"  small  flegree  of  springing  up  spon-  the  works  of  Mr.  Mill. 
"  taneously."     But  in  opposition  to 


CiiAi'.  IV.  Moral  Sense.  99 

behind  it  a  very  vivid  impression.  It  is  clear  that  many  in- 
stinctive desires,  sucli  as  that  of  hunger,  are  in  their  nature  of 
short  duration;  and  after  being  satisfied,  are  not  readily  or 
vividly  recalled.  ThirdJy,  after  the  power  of  language  had  been 
acquired,  and  the  wishes  of  the  community  could  be  expressed, 
the  common  opinion  how  each  member  ought  to  act  for  the 
public  good,  would  naturally  become  in  a  paramount  degree 
the  guide  to  action.  But  it  should  be  borne  in  mind  that  how- 
ever great  weight  we  may  attribute  to  public  oi)inion,  our  regard 
for  the  approbation  and  disapprobation  of  our  fellows  depends 
on  sympathy,  which,  as  we  shall  see,  forms  an  essential  part  of 
the  social  instinct,  and  is  indeed  its  foundation-stone,  Lastly, 
habit  in  the  individual  would  ultimately  play  a  very  important 
part  in  guiding  the  conduct  of  each  member ;  for  the  social  in- 
stinct, together  with  sympathy,  is,  like  any  other  instinct,  greatly 
strengthened  by  habit,  and  so  consequently  would  be  obedience 
to  the  wishes  and  judgment  of  the  community.  These  several 
subordinate  propositions  must  now  be  discussed,  and  some  of 
them  at  considerable  length. 

It  may  be  well  first  to  premise  that  I  do  not  wish  to  maintain 
that  any  strictly  social  animal,  if  its  intellectual  faculties  were 
to  become  as  active  and  as  highly  developed  as  in  man,  would 
acquire  exactly  the  same  moral  sense  as  ours.  In  the  same 
manner  as  various  animals  have  some  sense  of  beauty,  though 
they  admire  widely  different  objects,  so  they  might  have  a  sense 
of  right  and  wrong,  though  led  by  it  to  follow  widely  different 
lines  of  conduct.  If,  for  instance,  to  take  an  extreme  case,  men 
were  reared  under  precisely  the  same  conditions  as  hive-bees, 
there  can  hardly  be  a  doubt  that  our  unmarried  females  would, 
like  the  worker-bees,  think  it  a  sacred  duty  to  kill  their  brothers, 
and  mothers  would  strive  to  kill  their  fertile  daughters  ;  and  no 
one  would  think  of  interfering.*^     Nevertheless,  the  bee,  or  any 

^  Mr.  H.   Sidgwick   i-emarks,    in  in    Morals,'   'Theological    Review,' 

an   able  discussion  on    this  subject  April,    1872,    p.    188-191)   on    the 

(the   'Academy,'  June   15th,    1872,  same    illustration,    says,    the  j)rin- 

(p.  231),  "a  superior  bee,  we  may  ciplcs  of  social  duty  would  be  thus 

"  feel  sure,  would  aspire  to  a  milder  reversed  ;  and    by  this,    I  presume, 

"  solution  of  the  population   ques-  she  means  that  the  fulfilment  of  a 

"  tion."      Judging,    however,   from  social  duty  would  tend  to  tlie  injury 

the  habits  of  many  or  most  savages,  ot"  individuals  ;    but    she    overlooks 

man  solves  the  problem  by  famale  the  fact,  which  she  would  doubtless 

infanticide,  polyandry  and    promis-  admit,  that  tlie  instincts  of  the  bee 

cuous  intercourse  ;  therefore  it  may  have  been  acquired  for  the  good  of 

well   be  doubted  whether  it  would  the  community.     She  goes  so  far  as 

be     by    a     milder     method.     Jliss  to  say  that  if  the  theory  of  ethics 

Cobbe,  in  commenting  ('Darwinism  advocated  in  this  chapter  were  ever 


100  The  Descent  of  Ma]i.  Part  I. 

other  social  animal^  would  gain  in  our  supposed  case,  as  it 
appears  to  me,  some  feeling  of  right  or  wrong,  or  a  conscience. 
For  each  individual  would  have  an  inward  sense  of  possessing 
certain  stronger  or  more  enduring  instincts,  and  others  less 
strong  or  enduring ;  so  that  there  would  often  be  a  struggle  as  to 
which  impulse  should  be  followed;  and  satisfaction,  dissatis- 
faction, or  even  misery  would  be  felt,  as  past  impressions  were 
compared  during  their  incessant  passage  through  the  mind.  In 
this  case  an  inward  monitor  would  tell  the  animal  that  it  would 
have  been  better  to  have  followed  the  one  impulse  rather  than 
the  other.  The  one  course  ought  to  have  been  followed,  and  the 
other  ought  not ;  the  one  would  have  been  right  and  the  other 
wrong ;  but  to  these  terms  I  shall  recur. 

Sociahility. — Animals  of  many  kinds  are  social ;  we  find  even 
distinct  species  living  together ;  for  example,  some  American 
monkeys ;  and  united  flocks  of  rooks,  jackdaws,  and  starlings. 
Man  shews  the  same  feeling  in  his  strong  love  for  the  dog,  which 
the  dog  returns  with  interest.  Every  one  must  have  noticed  how 
miserable  horses,  dogs,  sheep,  &c.,  are  when  separated  from 
their  companions,  and  what  strong  mutual  affection  the  two 
former  kinds,  at  least,  shew  on  theii*  reunion.  It  is  curious  to 
speculate  on  the  feelings  of  a  dog,  who  will  rest  peacefully  for 
hours  in  a  room  with  his  master  or  any  of  the  family,  without 
the  least  notice  being  taken  of  him ;  but  if  left  for  a  short  time 
by  himself,  barks  or  howls  dismally.  "We  will  confine  our 
attention  to  the  higher  social  animals ;  and  pass  over  insects, 
although  some  of  these  are  social,  and  aid  one  another  in  many 
important  ways.  The  most  common  mutual  service  in  the 
higher  animals  is  to  warn  one  another  of  danger  by  means  of 
the  united  senses  of  all.  Every  sportsman  knows,  as  Dr.  Jaeger 
remarks,"  how  difficult  it  is  to  ajDproach  animals  in  a  herd  or 
troop.  Wild  horses  and  cattle  do  not,  I  believe,  make  any 
danger-signal;  but  the  attitude  of  any  one  of  them  who  first 
discovers  an  enemy,  warns  the  others.  Eabbits  stamp  loudly  on 
the  ground  with  their  hind-feet  as  a  signal :  sheep  and  chamois 
do  the  same  with  their  forefeet,  uttering  likewise  a  whistle. 
Many  birds,  and  some  mammals,  post  sentinels,  which  in  the 
case  of  seals  are  said^  generally  to  be  the  females.    The  leader 


generally  accepted,   "  I  cannot  but  earth  is  not  held  by  many  persons 

"  believe  that  in  the  hour  of  their  on  so  weak  a  tenure. 

"  triumph    would   be    sounded    the  "  '  Die   Darwin'sche    Theorie,'    s, 

*'  knell  of  the  virtue  of  mankind  '."  101, 

It  is  to  be  hoped  that  the  belief  in  *  Mr.  R.  Brown  in  '  Proc.  Zoolog. 

the  permanence   of  virtue  on  this  %<c.'  1868,  p.  409. 


Chap.  IV.  Moral  Sense.  lOi 


of  a  troop  of  monkeys  acts  as  the  sentinel,  and  utters  cries 
expressive  both  of  danger  and  of  safety.''  Social  animals  perform 
jnany  little  services  for  each  other :  horses  nibble,  and  cows  lick 
each  other,  on  any  spot  which  itches:  monkeys  search  each 
other  for  external  parasites ;  and  Brelim  sta,tcs  that  after  a  troop 
of  the  Cercopithecus  griseo-vlridls  has  rushed  through  a  thorny 
brake,  each  monkey  stretches  itself  on  a  branch,  and  another 
monkey  sitting  by,  "conscientiously"  examines  its  fur,  and 
extracts  every  thorn  or  burr. 

Animals  also  render  more  important  services  to  one  another : 
thus  wolves  and  some  other  beasts  of  prey  hunt  in  packs,  and 
aid  one  another  in  attacking  their  victims.  Pelicans  fish  in 
concert.  The  Hamadrj^as  baboons  turn  over  stones  to  find 
insects,  &c. ;  and  when  they  come  to  a  large  one,  as  many  as  can 
stand  round,  turn  it  over  together  and  share  the  booty.  Social 
animals  mutually  defend  each  other.  Bull  bisons  in  N.  America, 
when  there  is  danger,  drive  the  cows  and  calves  into  the  middle 
of  the  herd,  whilst  they  defend  the  outside.  I  shall  also  in  a 
future  chapter  give  an  account  of  two  young  wild  bulls  at 
Chillingham  attacking  an  old  one  in  concert,  and  of  two  stallions 
together  trying  to  drive  away  a  third  stallion  from  a  troop  of 
mares.  In  Abyssinia,  Brehm  encountered  a  great  troop  of 
baboons,  who  were  crossing  a  valley  :  some  had  already  ascended 
the  opposite  mountain,  and  some  were  still  in  the  valley :  the 
latter  were  attacked  by  the  dogs,  but  the  old  males  immediately 
hurried  down  from  the  rocks,  and  with  mouths  widely  opened, 
roared  so  fearfully,  that  the  dogs  quickly  drew  back.  They 
were  again  encouraged  to  the  attack ;  but  by  this  time  all  the 
baboons  had  reascendcd  the  heights,  excepting  a  young  one, 
about  six  months  old,  who,  loudly  calling  for  aid,  climbed  on  a 
block  of  rock,  and  was  surrounded.  Now  one  of  the  largest 
males,  a  true  hero,  came  down  again  from  the  mountain,  slowly 
went  to  the  young  one,  coaxed  him,  and  triumi:)hantly  led  him 
away — the  dogs  being  too  much  astonished  to  make  an  attack. 
I  cannot  resist  giving  another  scene  which  was  witnessed  by  this 
same  naturalist ;  an  eagle  seized  a  young  Ccrcopithecus,  which, 
by  clinging  to  a  branch,  was  not  at  once  carried  off;  it  cried 
loudly  for  assistance,  upon  which  the  other  members  of  the  troop, 
with  much  uproar,  rushed  to  the  rescue,  surrounded  the  eagle, 

9  Brehm,  'Thierlebcn,' B.  i.  18G4,  the  ovi  lence  of  Alvarez,  whose  ob- 
s.  52,  79.  For  the  cnse  of  the  servatious  Brehm  thinks  quite  trust- 
monkeys  extracting  thorns  from  worthy.  For  the  cases  of  the  okl 
each  other,  sec  s.  54-.  With  respect  male  baboons  attacking  the  dogs, 
to  the  Hamadryas  turning  over  see  s.  79 ;  and  with  respect  to  the 
stones,  the  fact  is  given  (s.  76)  on  eagle,  s.  5(3. 


102  The  Descent  of  Man.  Part  I. 

and  pnlled  out  so  many  feathers,  tliat  he  no  longer  thought 
of  his  prey,  but  only  how  to  escai)e.  This  eagle,  as  Brehm 
remarks,  assiu'edly  would  never  again  attack  a  single  monkey  of 
a  troop. ^'^ 

It  is  eei-tain  that  associated  animals  have  a  feeling  of  love  for 
each  other,  which  is  not  felt  by  non-social  adult  animals.  How 
far  in  most  cases  they  actually  sympathise  in  the  pains  and 
pleasures  of  others,  is  more  doubtful,  especially  with  respect  to 
])Ieasures.  Mr.  Buxton,  however,  who  had  excellent  means  of 
observation,"  states  that  his  macaws,  which  lived  free  in  Norfolk, 
took  "  an  extravagant  interest "  in  a  pair  with  a  nest ;  and  when- 
ever the  female  left  it,  she  was  surrounded  by  a  troop  '"  scream- 
"  ing  horrible  acclamations  in  her  honour."  It  is  often  diflBcult 
to  judge  whether  animals  have  any  feeling  for  the  sufferings  of 
others  of  their  kind.  Who  can  say  what  cows  feel,  when  they 
surround  and  stare  intently  on  a  dying  or  dead  companion ;  ap- 
parently, however,  as  Houzeau  remarks,  they  feel  no  pity.  That 
animals  sometimes  are  far  from  feeling  any  sympathy  is  too 
certain;  for  they  will  expel  a  wounded  animal  from  the  herd,  or 
gore  or  worry  it  to  death.  This  is  almost  the  blackest  fact  in 
natural  liistory,  unless,  indeed,  the  explanation  which  has  been 
suggested  is  true,  that  their  instinct  or  reason  leads  them  to 
expel  an  injured  companion,  lest  beasts  of  prey,  including  man, 
should  be  tempted  to  follow  the  troop.  In  this  case  their  con- 
duct is  not  much  worse  than  that  of  the  North  American  Indians, 
who  leave  their  feeble  comrades  to  perish  on  the  plains ;  or  the 
Fijians,  who,  when  their  parents  get  old,  or  fall  ill,  bury  them 
alive.^^ 

Many  animals,  however,  certainly  sympathise  with  each  other's 
distress  or  danger.  This  is  the  case  even  with  birds.  Capt. 
Stansbury  ^^  found  on  a  salt  lake  in  Utah  an  old  and  completely 
blind  pelican,  which  was  very  fat,  and  must  have  been  well  fed 
for  a  long  time  by  his  companions.    Mr.  Blyth,  as  he  informs 

''  Ml-.  Belt  gives   the   case   of  a  "  '  Annals  and  Mag.  of  Nat.  Hist.,' 

spider-monkey  (Ateles)    in    Nicara-  Kovember,  1868,  p.  382. 

gua,  which  was  heard  screaming  for  ^-  Sir    J.    Lubbock,    '  Prehistoric 

nearly  two  hours  in  the  forest,  and  Times,'  2nd  edit.  p.  446. 

was    found  with   an    eagle  perched  ^^  As  quoted  by  Mr.  L.  H.  Morgan, 

close   by  it.     The    bird    apparently  '  The  American   Beaver,'    1868,   p. 

feared   to   attack  as  long  as  it  re-  272.     Capt.  Stansbury  also  gives  an 

mained  f\\ce  to  face;  and  Mr.  Belt  interesting  account  of  the  manner 

believes,  from  what  he  has  seen  of  in   which    a   very    young    pelican, 

the  habits   of  these  monkeys,  that  carried    away  by  a    strong  stream, 

they  protect  themselves  from  eagles  was  guided   and  encouraged   in  its 

by  "keeping  two  or  three  together,  attempts  to  reach  the  shore  by  half 

'The  Naturalist  in  Nicaragua,'  1874,  a  dozen  old  birds 
p.  118. 


Chap.  IV.  Moral  Sense.  103 

me,  saw  Indian  crows  feeding  two  or  three  of  their  companions 
which  were  blind ;  and  I  have  heard  of  an  analogous  case  with 
the  domestic  cock.  We  may,  if  we  choose,  call  these  actions 
instinctive  ;  but  such  cases  are  much  too  rare  for  the  develop- 
ment of  any  special  instinct.^*  I  have  myself  seen  a  dog,  who 
never  passed  a  cat  who  lay  sick  in  a  basket,  and  was  a  great 
friend  of  his,  without  giving  her  a  few  licks  with  his  tongue,»tlie 
surest  sign  of  kind  feeling  in  a  dog. 

It  must  be  called  sympathy  that  leads  a  courageous  dog  to 
fly  at  any  one  who  strikes  his  master,  as  he  certainly  will.  I 
saw  a  person  pretending  to  beat  a  lady,  who  had  a  very  timid 
little  dog  on  her  lap,  and  the  trial  had  never  been  made  before  ; 
the  little  creature  instantly  jumped  away,  but  after  the  pretended 
beating  was  over,  it  was  really  pathetic  to  see  how  perseveringly 
he  tried  to  lick  his  mistress's  face,  and  comfort  her.  Brehm  '^ 
states  that  when  a  baboon  in  confinement  was  pursued  to  be 
punished,  the  others  tried  to  protect  him.  It  must  have  been 
sympathy  in  the  cases  above  given  which  led  the  baboons  and 
Cercopitheci  to  defend  their  young  comrades  from  the  dogs  and 
the  eagle.  I  will  give  only  one  other  instance  of  sympathetic 
and  heroic  conduct,  in  the  case  of  a  little  American  monkey. 
Several  years  ago  a  keeper  at  the  Zoological  Gardens  shewed  me 
some  deep  and  scarcely  healed  wounds  on  the  nape  of  his  own  neck, 
inflicted  on  him,  whilst  kneeling  on  the  floor,  by  a  fierce  baboon. 
The  little  American  monkey,  who  was  a  warm  friend  of  this 
keeper,  lived  in  the  same  large  compartment,  and  was  dreadfully 
afraid  of  the  great  baboon.  Nevertheless,  as  soon  as  he  saw  his 
friend  in  peril,  he  rushed  to  the  rescue,  and  by  screams  and  bites 
so  distracted  the  baboon  that  the  man  was  able  to  escape,  after, 
as  the  surgeon  thought,  running  great  risk  of  his  life. 

Besides  love  and  symi:>athy,  animals  exhibit  other  qualities 
connected  with  the  social  instincts,  which  in  us  would  be  called 
moral ;  and  I  agree  with  Agassiz  ^^  that  dogs  possess  something 
very  hke  a  conscience. 

Dogs  possess  some  power  of  self-command,  and  this  does  not 
appear  to  be  wholly  the  result  of  fear.  As  Braubach'^  remarks, 
they  will  refrain  from  stealing  food  in  the  absence  of  their 
master.  They  have  long  been  accepted  as  the  very  type  of 
fidelity  and  obedience.  But  the  elei)hant  is  likewise  very  faith- 
ful to  his  driver  or  keeper,  and  probably  considers  him  as  the 

'*  As  Mr.  Bain  states,  "effective  ^^  '  De   I'Esp&ce   ct  dc  la  Classe,' 

"  aid  to  a  sufferer  springs  from  sym-  18(59,  p.  97. 

"  pathy  proper  :" 'Mental  and  Moral  *'  '  Die    Darwin'sclic    Art-Lehre,* 

Science,'  1868,  p.  245.  18G9,  s.  54. 

*^  'Thierleben,   B.  i.  s,  85. 


T04  The  Descent  of  Man.  Pari  I. 

leader  of  the  lierd.  Dr.  Hooker  informs  me  that  an  elephant, 
which  he  was  riding  in  India,  became  so  deeply  bogged  that  he 
remained  stnck  fast  until  the  next  day,  when  he  was  extricated 
by  men  with  ropes.  Under  snch  circumstances  elephants  will 
seize  with  their  trunks  any  object,  dead  or  alive,  to  place  under 
their  knees,  to  prevent  their  sinking  deeper  in  the  mud  ;  and  the 
driver  was  dreadfully  afraid  lest  the  animal  should  have  seized 
Dr.  Hooker  and  crushed  him  to  death.  But  the  driver  himself, 
as  Dr.  Hooker  was  assured,  ran  no  risk.  This  forbearance  under 
an  emergency  so  dreadful  for  a  heavy  animal,  is  a  wonderful 
proof  of  noble  fidelity .^^ 

All  animals  living  in  a  body,  which  defend  themselves  or  attack 
their  enemies  in  concert,  must  indeed  be  in  some  degree  faithful 
to  one  another ;  and  those  that  follow  a  leader  must  be  in  some 
degree  obedient.  "When  the  baboons  in  Abyssinia  ^^  plunder  a 
garden,  they  silently  follow  their  leader ;  and  if  an  imprudent 
young  animal  makes  a  noise,  he  receives  a  slap  from  the  others 
to  teach  him  silence  and  obedience.  j\Ir.  Galton,  who  has  had 
excellent  opportunities  for  observing  the  half-wild  cattle  in  S. 
Africa,  says,-"  that  they  cannot  endure  even  a  momentary  separa- 
tion from  the  herd.  They  are  essentially  slavish,  and  accept  the 
common  determination,  seeking  no  better  lot  than  to  be  led  by 
any  one  ox  who  has  enough  self-reliance  to  accept  the  position. 
The  men  who  break  in  these  animals  for  harness,  watch  assidu- 
ously for  those  who,  by  grazing  apart,  shew  a  self-reliant  dis- 
position, and  these  they  train  as  fore-oxen.  Mr.  Galton  adds 
that  such  animals  are  rare  and  valuable  ;  and  if  many  were  born 
they  would  soon  be  eliminated,  as  lions  are  always  on  the  look- 
out for  the  individuals  which  wander  from  the  herd. 

"With  respect  to  the  impulse  which  leads  certain  animals  to 
associate  together,  and  to  aid  one  another  in  many  ways,  we 
may  infer  that  in  most  cases  they  are  impelled  by  the  same 
sense  of  satisfaction  or  pleasure  which  they  experience  in  per- 
forming other  instinctive  actions;  or  by  the  same  sense  of 
dissatisfaction  as  when  other  instinctive  actions  are  checked. 
^Ye  see  this  in  innumerable  instances,  and  it  is  illustrated  in  a 
striking  manner  by  the  acquired  instincts  of  our  domesticated 
animals ;  thus  a  young  shepherd-dog  delights  in  driving  and 
running  round  a  flock  of  sheep,  but  not  in  worrying  them ;  a 
young  fox-hound  delights  in  hunting  a  fox,  whilst  some  other 
kinds  of  dogs,  as  I  have  witnessed,  utterly  disregard  foxes.   What 

^^  See  also  Hooker's  '  Himalayan  20  See  his  extremely  interesting 

Journals,'  vol.  ii.,  1854,  p.  333.  paper  on  '  Gregariousness  in  Cattle, 

**  Brehm,  '  Thierleben,'    B.   i.  s.  and  in  Man,' '  Macmiilan's  Mag.'  Feb. 

7G.  1871,  p.  353. 


Chap.  IV.  Moral  Sense.  105 


a  strong  feeling  of  inward  satisfaction  must  impel  a  bird,  so  full 
of  activity,  to  brood  day  after  day  over  her  eggs.  Migratory 
birds  are  quite  miserable  if  stopped  from  migrating;  perliajjs 
they  enjoy  starting  on  their  long  flight ;  but  it  is  hard  to  believe 
that  the  poor  pinioned  goose,  described  by  Audubon,  which 
started  on  foot  at  the  proper  time  for  its  journey  of  probably 
more  than  a  thousand  miles,  could  have  felt  any  joy  in  doing  so. 
Some  instincts  are  determined  solely  by  painful  feelings,  as  by 
fear,  which  leads  to  self-preservation,  and  is  in  some  cases  directed 
towards  special  enemies.  No  one,  I  presume,  can  analyse  the 
sensations  of  pleasure  or  pain.  In  many  instances,  however,  it 
is  probable  that  instincts  are  persistently  followed  from  the 
mere  force  of  inheritance,  without  the  stimulus  of  either 
l">leasure  or  pain.  A  young  pointer,  when  it  tirst  scents  game, 
apparently  cannot  help  pointing.  A  squirrel  in  a  cage  who  pais 
the  nuts  which  it  cannot  eat,  as  if  to  bury  them  in  the  ground, 
can  hardly  be  thought  to  act  thus,  either  from  pleasure  or  pain. 
Hence  the  common  assumption  that  men  must  be  impelled  to 
every  action  by  experiencing  some  pleasure  or  pain  may  be  erro- 
neous. Although  a  habit  may  be  blindly  and  implicitly 
followed,  independently  of  any  j^lcasure  or  pain  felt  at  the 
moment,  yet  if  it  be  forcibly  and  abruptly  checked,  a  vague 
sense  of  dissatisfaction  is  generally  experienced. 

It  has  often  been  assumed  that  animals  were  in  the  first  place 
rendered  social,  and  that  they  feel  as  a  consequence  uncomfort- 
able when  separated  from  each  other,  and  comfortable  whilst 
together ;  but  it  is  a  more  probable  view  that  these  sensations 
were  first  developed,  in  order  that  those  animals  which  would 
profit  by  living  in  society,  should  be  induced  to  live  together, 
in  the  same  manner  as  the  sense  of  hunger  and  the  pleasure  of 
eating  were,  no  doubt,  first  acquired  in  order  to  induce  animals 
to  eat.  The  feeling  of  j^lcasure  from  society  is  probably  an 
extension  of  the  parental  or  filial  affections,  since  the  social 
instinct  seems  to  be  developed  by  the  young  remaining  for  a 
long  time  with  their  parents;  and  this  extension  may  be  attri- 
buted in  part  to  habit,  but  chiefly  to  natural  selection.  With 
tliose  animals  which  were  benefited  by  living  in  close  association, 
the  individuals  which  took  the  greatest  pleasure  in  society 
would  best  escape  various  dangers;  whilst  those  that  cared 
least  for  their  comrades,  and  lived  solitary,  would  perish  in 
greater  numbers.  With  respect  to  the  origin  of  the  parental 
and  fihal  affections,  which  apparently  lie  at  the  base  of  the 
social  instincts,  we  know  not  the  steps  by  which  they  have 
been  gained;  but  we  may  infer  that  it  has  been  to  a  large 
extent  through  natural  selection.  So  it  has  almost  certainly 
6 


io6  The  Descent  of  Man.  Part  T. 


been  with  the  unusual  aiid  opposite  feeling  of  hatred  between 
the  nearest  relations,  as  with  the  worker-bees  which  kill  their 
brother-drones,  and  with  the  queen-bees  which  kill  their 
daughter-queens;  the  desire  to  destroy  their  nearest  relations 
having  been  in  this  case  of  service  to  the  community.  Parental 
affection,  or  some  feeling  which  replaces  it,  has  been  developed 
in  certain  animals  extremely  low  in  the  scale,  for  example,  in 
star-fishes  and  spiders.  It  is  also  occasionally  present  in  a  few 
members  alone  in  a  whole  group  of  animals,  as  in  the  genus 
Forficula,  or  earwigs. 

The  all-important  emotion  of  sympathy  is  distinct  from  that 
of  love.  A  mother  may  passionately  love  her  sleeping  and 
passive  infant,  but  she  can  hardly  at  such  times  be  said  to  feel  • 
sympathy  for  it.  The  love  of  a  man  for  his  dog  is  distinct  from 
sympathy,  and  so  is  that  of  a  dog  for  his  master.  Adam  Smith 
formerly  argued,  as  has  Mr.  Bain  recently,  that  the  basis  of 
sympathy  Hes  in  our  strong  retentiveness  of  former  states  of 
pain  or  pleasure.  Hence,  "the  sight  of  another  person  en- 
"  during  hunger,  cold,  fatigue,  revives  in  us  some  recollection  of 
"  these  states,  which  are  painful  even  in  idea."  We  are  thus 
impelled  to  relieve  the  sufferings  of  another,  in  order  that  our 
own  painful  feelings  may  be  at  the  same  time  relieved.  In  Hke 
manner  we  are  led  to  participate  in  the  pleasures  of  others.^' 
But  I  cannot  see  how  this  view  explains  the  fact  that  sympathy 
is  excited,  in  an  immeasurably  stronger  degree,  by  a  beloved, 
than  by  an  indifferent  person.  The  mere  sight  of  suffering, 
independently  of  love,  would  suffice  to  call  up  in  us  vivid 
recollections  and  associations.  The  explanation  may  lie  in  the 
fact  that,  with  all  animals,  sympathy  is  dhected  solely  towards 
the  members  of  the  same  community,  and  therefore  towards 
known,  and  more  or  less  beloved  members,  but  not  to  all  the 
individuals  of  the  same  species.  This  fact  is  not  more  sur- 
prising than  that  the  fears  of  many  animals  should  be  directed 
against  special  enemies.  Species  which  are  not  social,  such  as 
lions  and  tigers,  no  doubt  feel  sympathy  for  the  suffering  of 
tbeir  own  young,  but  not  for  that  of  any  other  animal.     With 

21     See    the     first    and    striking  "  or  others  in  his  stead,  may  make 

chapter    in  Adam  Smith's   '  Theory  "  up,  by  sympathy  and  good   offices 

of    Moral    Sentiments.'      Also    Mr.  "  returned,    for   all    the    sacrifice." 

Bain's  '  Mental  and  Moral  Science,'  But  if,  as  appears  to  be  the  case, 

1868,    p.   244,  and    275-282.     Mr.  sympathy   is    strictly   an    instinct, 

Bain  states,  that  "sympathy  is,  in-  its  exercise  would  give  direct  plea- 

"  directly,  a  source  of  pleasure   to  sure,  in   the   same   manner  as    the 

"  the  sympathiser  ;"  and  he  accounts  exercise,  as  before  remarked,  of  al- 

f(ir   this    through    reciprocity.     He  most  every  other  instinct, 
remarks  that  "  the  person  benefited, 


CtiAP.  IV.  Moral  Sense.  107 


mankind,  sclfislincss,  experience,  and  imitation,  probably  add, 
as  Mr.  Bain  has  shown,  to  the  power  of  sympathy ;  for  we  are 
led  by  the  hope  of  receiving  good  in  return  to  perform  acts 
of  sympathetic  kindness  to  others ;  and  sympathy  is  much 
strengthened  by  habit.  In  however  complex  a  manner  this 
feeling  may  have  originated,  as  it  is  one  of  high  importance  to 
all  those  animals  which  aid  and  defend  one  another,  it  will  have 
been  increased  through  natural  selection ;  for  those  commu- 
nities, which  included  the  greatest  number  of  the  most  sympa- 
thetic members,  would  flourish  best,  and  rear  the  greatest 
number  of  offspring. 

It  is,  however,  impossible  to  decide  in  many  cases  whether 
certain  social  instincts  have  been  acquired  through  natural 
selection,  or  are  the  indirect  result  of  other  instincts  and 
faculties,  such  as  sympathy,  reason,  experience,  and  a  tendency 
to  imitation ;  or  again,  whether  they  are  simply  the  result  of 
long-continued  habit.  So  remarkable  an  instinct  as  the  placing 
sentinels  to  warn  the  community  of  danger,  can  hardly  have 
been  the  indirect  result  of  any  of  these  faculties  ;  it  must,  there- 
fore, have  been  directly  acquired.  On  the  other  hand,  the  habit 
followed  by  the  males  of  some  social  animals  of  defending  the 
community,  and  of  attacking  their  enemies  or  their  prey  in 
concert,  may  perhaps  have  originated  from  mutual  sym^Dathy ; 
but  courage,  and  in  most  cases  strength,  must  have  been 
previously  acquired,  probably  through  natural  selection. 

Of  the  various  instincts  and  habits,  some  are  mnch  stronger 
than  others ;  that  is,  some  either  give  more  pleasure  in  their 
performance,  and  more  distress  in  their  prevention,  than  others ; 
or,  ^hich  is  probably  quite  as  important,  they  are,  through 
inheritance,  more  persistently  followed,  without  exciting  any 
special  feeling  of  pleasure  or  pain.  We  are  ourselves  conscious 
that  some  habits  are  much  more  difficult  to  cure  or  change  than 
others.  Hence  a  struggle  may  often  be  observed  in  animals 
between  different  instincts,  or  between  an  instinct  and  some 
habitual  disposition;  as  when  a  dog  rushes  after  a  hare,  is 
rebuked,  pauses,  hesitates,  pursues  again,  or  returns  ashamed  to 
his  master  ;  or  as  between  the  love  of  a  female  dog  for  her  young 
l-)uppies  and  for  her  master,— for  she  may  be  seen  to  slink  away 
to  them,  as  if  half  ashamed  of  not  accompanying  her  master. 
But  the  most  carious  instance  known  to  me  of  one  instinct 
getting  the  better  of  another,  is  the  migratory  instinct  conquer- 
ing the  maternal  instinct.  The  former  is  wonderfully  strong;  a 
confined  bird  will  at  the  proper  season  beat  her  breast  against 
the  wires  of  her  cage,  until  it  is  bare  and  bloody.  It  causes 
young  salmon  to  leap  out  of  the  fresh  water,  in  which  they  could 


io8  The:  Descent  of  Man.  Part  T. 

continue  to  exist,  and  thus  unintentionally  to  commit  suicide. 
Every  one  knows  how  strong  the  maternal  instinct  is,  leading 
even  timid  birds  to  face  great  danger,  though  with  hesitation, 
and  in  opposition  to  the  instinct  of  self-preservation.  Neverthe- 
less, the  migratory  instinct  is  so  powerful,  that  late  in  the  autumn 
swallows,  house-martins,  and  swifts  frequently  desert  their 
tender  young,  leaving  them  to  perish  miserably  in  their  nests.^^ 

We  can  perceive  that  an  instinctive  impulse,  if  it  be  in  any 
way  more  beneficial  to  a  species  than  some  other  or  ojDposed 
instinct,  would  be  rendered  the  more  potent  of  the  two  through 
natural  selection  ;  for  the  individuals  which  had  it  most  strongly 
developed  would  survive  in  larger  numbers.  Whether  this  is  the 
case  with  the  migratory  in  comparison  with  the  maternal  instinct, 
may  be  doubted.  The  great  persistence,  or  steady  action  of  the 
former  at  certain  seasons  of  the  year  during  the  whole  day,  may 
give  it  for  a  time  paramount  force. 

Man  a  social  animal. — Every  one  will  admit  that  man  is  a 
social  being.  We  see  this  in  his  dislike  of  solitude,  and. in  his 
wish  for  society  beyond  that  of  his  own  family.  Solitary  con- 
finement is  one  of  the  severest  punishments  which  can  be 
inflicted.  Some  authors  suppose  that  man  primevally  lived  in 
single  families  ;  but  at  the  present  day,  though  single  families, 
or  only  two  or  three  together,  roam  the  solitudes  of  some  savage 
lands,  they  always,  as  far  as  I  can  discover,  hold  friendly 
relations  with  other  families  inhabiting  the  same  district.  Such 
families  occasionally  meet  in  council,  and  unite  for  their  common 
defence.  It  is  no  argument  against  savage  man  being  a  social 
animal,  that  the  tribes  inhabiting  adjacent  districts  are  almost 
always  at  war  with  each  other  ;  for  the  social  instincts  never 
extend  to  all  the  individuals  of  the  same  species.  Judging  from 
the  analogy  of  the  majority  of  the  Quadrumana,  it  is  probable 
that  the  early  ape-like  progenitors  of  man  were  likewise  social ; 
but  thLs  is  not  of  much  importance  for  us.     Although  man,  as 

^*  This  fact,  the  Rev.  L.  Jenyns  hatched.     Many  birds,  not  yet  old 

states  (see  his  edition  of  '  White's  enough    for  a  prolonged  flight,  are 

Nat.    Hist,    of   Selborne,'    1853,    p.  likewise    deserted   and   left    behind. 

204)  was  first  recorded  by  the  illus-  See  Blackwall,  'Researches  in  Zoo- 

trious  Jenner,  in   'Phil.  Transact,'  logy,' 1834,  pp.  108,  118.     For  some 

1824,  and  has  since  been  confirmed  additional    evidence,    although    this 

by  several   observers,  especially  by  is    not  wanted,  see  Leroy,  '  Lettres 

Mr.  Blackwall.     This  latter  careful  Phil.'    1802,    p.    217.     For    Swifts, 

observer     examined,    late     in     the  Gould's  '  Introduction  to  the  Birds 

autumn,  during  two  years,  thirty-  of  Great  Britain,'  1823,  p.  o.    Simi- 

six    nests ;    he    found    that    twelve  lar    cases    have    been    observed    in 

contained   young    dead    birds,    five  Canada     by     Mr.     Adams ;     '  Pop. 

contained  eggs  on  the  point  of  being  Science     Review,'    July     1873,     p. 

hatched,  and  three,  eggs  not  nearly  283. 


CiiAP.  IV.  Moral  Sense.  109 


ho  now  exists,  has  few  special  instincts,  having  lost  any  which 
his  early  progenitors  may  have  possessed,  tliis  is  no  reason  wliy 
he  should  not  have  retained  from  an  extremely  remote  period 
some  degree  of  instinctive  love  and  sympathy  for  his  fellows. 
We  are  indeed  all  conscious  that  we  do  possess  such  sympathetic 
feelings ;  -^  but  our  consciousness  does  not  tell  ns  whether  they 
are  instinctive,  having  originated  long  ago  in  the  same  manner 
as  with  the  lower  animals,  or  whether  tliey  have  been  acquired 
by  each  of  us  during  our  early  years.  As  man  is  a  social  animal, 
it  is  almost  certain  that  he  would  inherit  a  tendency  to  be 
faithful  to  his  comrades,  and  obedient  to  the  leader  of  his  tribe ; 
for  these  qualities  are  connnon  to  most  social  animals.  lie  would 
consequently  possess  some  capacity  for  self-command.  lie 
would  from  an  inherited  tendency  be  willing  to  defend,  in 
concert  with  others,  his  fellow-men ;  and  would  be  ready  to  aid 
them  in  any  way,  which  did  not  too  greatly  interfere  with  his 
own  welfare  or  his  own  strong  desires. 

The  social  animals  which  stand  at  the  bottom  of  the  scale  are 
guided  almost  exclusively,  and  those  which  stand  higher  in  the 
scale  are  largely  guided,  by  special  instincts  in  the  aid  which 
they  give  to  the  members  of  the  same  community ;  but  they  arc 
likewise  in  part  impelled  by  mutual  love  and  sympathy,  assisted 
apparently  by  some  amount  of  reason.  Although  man,  as  just 
remarked,  has  no  special  instincts  to  tell  him  how  to  aid  his 
fellow-men,  he  still  has  the  impulse,  and  with  his  improved 
intellectual  faculties  would  naturally  be  much  guided  in  this 
respect  by  reason  and  experience.  Instinctive  sympathy  would 
also  cause  him  to  value  highly  the  approbation  of  his  fello^^s  ; 
for,  as  Mr.  Bain  has  clearly  shewn,^^  the  love  of  praise  and 
the  strong  feeling  of  glory,  and  the  still  stronger  horror  of  scorn 
and  infamy,  "  are  due  to  the  workings  of  sympathy."  Conse- 
quently man  would  be  influenced  in  the  highest  degree  by  the 
wishes,  approbation,  and  blame  of  his  fellow-men,  as  expressed 
by  their  gestures  and  language.  Thus  the  social  instincts, 
which  must  have  been  acquired  by  man  in  a  very  rude  state, 
and  probably  even  by  his  early  ape-like  progenitors,  still  give 
the  impulse  to  some  of  his  best  actions :  but  his  actions  are  in  a 
higher  degree  determined  by  the  expressed  wishes  and  judgment 

2'  Hume  remarks  ('An  Enquiiy  *' of  the  former  .  .  .  communicates 

Concerning  the  Principles  of  Morals,'  "a  secret  joy;    the  appearance  of 

edit,  of  1751,  p.  lo2),  "There  seems  "the   latter  .  .  .  throws  a  melau- 

"  a  necessity  for  confessing  that  the  "  choly    damp    ov6r    the    imagiua- 

"  happiness    and    misery    of    others  "  tion." 

"are  not  spectacles    altogether  in-  '^■'  'Mental   ami    Moral    Science,' 

"  different  to  us.  but  that  the  view  ISGS,  p.  2.04 


no  TJie  Descent  of  Man.  Tart  1. 

of  his  fellow-men,  and  unfortunately  very  often  by  his  own  strong 
selfish  desires.  But  as  love,  sympathy  and  self-command  become 
strengthened  by  habit,  and  as  the  power  of  reasoning  becomes 
clearer,  so  that  man  can  value  justly  the  judgments  of  his 
fellows,  he  will  feel  himself  impelled,  apart  from  any  transitory 
pleasure  or  pain,  to  certain  lines  of  conduct.  He  might  then 
declare — not  that  any  barbarian  or  uncultivated  man  could 
thus  think — I  am  the  supreme  judge  of  my  own  conduct,  and  in 
the  words  of  Kant,  I  will  not  in  my  own  person  violate  the 
dignity  of  humanity. 

The  more  enduring  Socid  Instincts  conquer  the  less  2yersisfent 
Instincts. — We  have  not,  however,  as  yet  considered  the  main 
point,  on  which,  from  our  present  point  of  view,  the  whole 
question  of  the  moral  sense  turns.  Why  should  a  man  feel  that 
he  ought  to  obey  one  instinctive  desii*e  rather  than  another  ? 
"Why  is  he  bitterly  regretful,  if  he  has  yielded  to  a  strong  sense 
of  self-preservation,  and  has  not  risked  his  life  to  save  that  of  a 
fellow-creature  ?  or  why  does  he  regret  having  stolen  food  from 
hunger  ? 

It  is  evident  in  the  first  place,  that  with  mankind  the  instinc- 
tive impulses  have  different  degi-ees  of  strength ;  a  savage  will 
risk  his  own  life  to  save  that  of  a  member  of  the  same  community, 
but  will  be  wholly  indifferent  about  a  stranger  :  a  young  and 
timid  mother  urged  by  the  maternal  instinct  will,  without  a 
moment's  hesitation,  run  the  greatest  danger  for  her  own  infant, 
but  not  for  a  mere  fellow-creature.  Nevertheless  many  a 
civilized  man,  or  even  boy,  who  never  before  risked  his  life  for 
andther,  but  full  of  courage  and  sympathy,  has  diregarded 
the  instinct  of  self-preservation,  and  plunged  at  once  into  a 
torrent  to  save  a  drowning  man,  though  a  stranger.  In  this  case 
man  is  impelled  by  the  same  instinctive  motive,  which  made  the 
heroic  little  American  monkey,  formerly  described,  save  his 
keeper,  by  attacking  the  great  and  dreaded  baboon.  Such 
actions  as  the  above  appear  to  be  the  simple  result  of  the  gi-eater 
strength  of  the  social  or  maternal  instincts  than  that  of  any 
other  instinct  or  motive;  for  they  are  perfonned  too  instan- 
taneously for  reflection,  or  for  pleasure  or  pain  to  be  felt  at  the 
time  ;  though,  if  prevented  by  any  cause,  distress  or  even  misery 
might  be  felt.  In  a  timid  man,  on  the  other  hand,  the  instinct 
of  self-preservation  might  be  so  strong,  that  he  would  be  unable 
to  force  himself  to  run  any  such  risk,  perhaps  not  even  for  his 
own  child. 

I  am  aware  that  some  persons  maintain  that  actions  performed 
impulsively,  as  in  the  above  cases,  do  not  come  under  the 
domijiion  of  the  moral  sense,  and  cannot  be  called  moral.    They 


Chap.  iV.  Mora/  Sense.  1 1 1 

confine  this  term  to  actions  done  deliberately,  after  a  victory 
over  opposing  desires,  or  when  prompted  l)y  some  exalted 
motive.  But  it  appears  scarcely  possible  to  draw  any  clear  line 
of  distinction  of  this  kind.'-^^  As  far  as  exalted  motives  are 
concerned,  many  instances  have  been  recorded  of  savages, 
destitute  of  any  feeling  of  general  benevolence  towards  mankind, 
and  not  guided  by  any  religious  motive,  who  have  deliberately 
sacrificed  their  lives  as  i)risoners,^''  rather  than  betray  their 
comrades;  and  surely  their  conduct  ought  to  be  considered  as 
moral.  As  far  as  deliberation,  and  the  victory  over  opposing 
motives  are  concerned,  animals  may  be  seen  doubting  between 
opposed  instincts,  in  rescuing  their  offspring  or  comrades 
from  danger ;  yet  their  actions,  though  done  for  the  good  of 
others,  are  not  called  moral.  Moreover,  anything  performed 
very  often  by  us,  will  at  last  be  done  without  deliberation  or 
hesitation,  and  can  then  hardly  be  distinguished  from  an 
instinct;  yet  surely  no  one  will  pretend  that  such  an  action 
ceases  to  be  moral.  On  the  contrary,  w^e  all  feel  that  an  act 
cannot  be  considered  as  perfect,  or  as  performed  in  the  most 
noble  manner,  unless  it  be  done  impulsively,  without  deliberation 
or  effort,  in  the  same  manner  as  by  a  man  in  whom  the  requisite 
qualities  are  innate.  He  who  is  forced  to  overcome  his  fear  or 
want  of  sympathy  before  he  acts,  deserves,  however,  in  one  way 
higher  credit  than  the  man  whose  innate  disposition  leads  him 
to  a  good  act  without  effort.  As  we  cannot  distinguish  between 
motives,  we*  rank  all  actions  of  a  certain  class  as  moral,  if 
performed  by  a  moral  being.  A  moral  being  is  one  who  is 
capable  of  comparing  his  past  and  future  actions  or  motives,  and 
of  approving  or  disapproving  of  them.  "We  have  no  reason  to 
suppose  that  any  of  the  lower  animals  have  this  capacity ; 
therefore,  wdien  a  Newfoundland  dog  drags  a  child  out  of  the 
water,  or  a  monkey  faces  danger  to  rescue  its  comrade,  or  takes 
charge  of  an  orphan  monkey,  we  do  not  call  its  conduct  moral. 
But  in  the  case  of  man,  who  alone  can  with  certainty  be  ranked 
as  a  moral  being,  actions  of  a  certain  class  are  called  moral, 
whether  performed  deliberately,  after  a  struggle  with  oj^posing 

"^^  I  refer  here  to  the  distinction  "  material    and    formal  morality  is 

between  what  has  been  called  ma-  "  as    irrelevant  as  other  siuh    dis- 

terial  and  formal  morality.     I   am  "  tinctions." 

glad  to  find  that  Prof.  Huxley  ('Cri-  ^g  j  have    given    one   such    case, 

tiques  and  Addresses,'  1873,  p.  287)  namely  of  three  Patagonian  Indians 

takes  the  same  view  on  this  subject  who  jtreferred  being  shot,  one  after 

as    I    do.     Mr.    Leslie    Stc])hen    re-  the  other,  to  betraying  the  ]»lans  of 

marks  ('  Essavs  on  Freethinking  and  their  companions  in  war  ('  Journal 

Plain  Speaking,'  1873,  p.  8;'.)/"  the  of  Researches,'  1845,  p.  103). 
"  metaphysical  distinction   betwceu 


112  The  Descent  of  Man.  Part  I. 

motives,  or  impulsively  through  instinct,  or  from  the  effects  of 
slowly-gained  habit. 

But  to  return  to  our  more  immediate  subject.  Although 
some  uistincts  are  more  powerful  than  others,  and  thus  lead  to  cor- 
responding actions,  yet  it  is  untenable,  that  in  man  the  social 
instincts  (including  the  love  of  praise  and  fear  of  blame)  possess 
greater  strength,  or  have,  through  long  habit,  acquired  greater 
strength  than  the  instincts  of  self-preservation,  hunger,  lust, 
vengeance,  &c.  Why  then  does  man  regret,  even  though  trying 
to  banish  such  regret,  that  he  has  followed  the  one  natural 
impulse  rather  than  the  other ;  and  why  does  he  further  feel 
that  he  ought  to  regret  his  conduct  ?  Man  in  this  respect  differs 
profoundly  from  the  lower  animals.  Nevertheless  we  can,  I  think, 
see  with  some  degree  of  clearness  the  reason  of  this  difference. 

Man,  from  the  activity  of  his  mental  faculties,  cannot  avoid 
reflection :  past  impressions  and  images  are  incessantly  and 
clearly  passing  through  his  mind.  Now  with  those  animals 
which  live  permanently  in  a  body,  the  social  instincts  are  ever 
present  and  persistent.  Such  animals  are  always  ready  to  utter 
the  danger-signal,  to  defend  the  community,  and  to  give  aid  to 
their  fellows  in  accordance  w^ith  their  habits ;  they  feel  at  all 
times,  without  the  stimulus  of  any  special  passion  or  desire, 
some  degree  of  love  and  sympathy  for  them ;  they  are  unhappy 
if  long  separated  from  them,  and  always  happy  to  be  again  m 
their  company.  So  it  is  with  ourselves.  Even  when  we  are 
quite  alone,  how  often  do  we  think  with  pleasure  or  pain  of 
what  others  think  of  us, — of  their  imagined  aijprobation  or 
disapprobation;  and  this  all  follows  from  sympathy,  a  funda- 
mental element  of  the  social  instincts.  A  man  who  possessed 
no  trace  of  such  instincts  would  be  an  unnatural  monster.  On 
the  other  hand,  the  desire  to  satisfy  hunger,  or  any  passion  such 
as  vengeance,  is  in  its  nature  temporary,  and  can  for  a  time  be 
fully  satisfied.  Nor  is  it  easy,  jDcrhaps  hardly  possible,  to  call 
up  with  complete  vividness  the  feeling,  for  instance,  of  hunger ; 
nor  indeed,  as  has  often  been  remarked,  of  any  suffering.  The 
instinct  of  self-preservation  is  not  felt  except  in  the  presence  of 
danger;  and  many  a  coward  has  thought  himself  brave  until  he 
has  met  his  enemy  face  to  face.  The  wish  for  another  man's 
property  is  jDsrhaps  as  persistent  a  desire  as  any  that  can  be 
named;  but  even  in  this  case  the  satisfaction  of  actual  pos- 
session is  generally  a  weaker  feeling  than  the  desire  :  many  a 
thief,  if  not  a  habitual  one,  after  success  has  wondered  why  he 
stole  some  article.^ 

^^  Enmity  or  hatred  seems  also  perhaps  more  so  than  auy  other 
to   be    a   highly  persistent   feeling,      that   can   be   named.     Envy  is  de- 


Chap.  IV.  Moral  Sense.  1 1 3 

A  man  cannot  jircYcnt  past  impressions  often  repassing  tlirongli 
liis  mind  ;  lie  will  thns  be  driven  to  make  a  comparison  between 
tbo  imjiressions  of  past  Imnger,  vengeance  satisfied,  or  danger 
shunned  at  other  men's  cost,  with  the  almost  ever-present 
instinct  of  sympathy,  and  with  his  early  knowledge  of  what 
others  consider  as  praise v/Orthy  or  blameable.  This  knowledge 
cannot  be  banished  from  his  mind,  and  from  instinctive  sympathy 
is  esteemed  of  great  moment.  He  will  then  feel  as  if  he  had 
been  baulked  in  following  a  present  instinct  or  habit,  and  this 
with  all  animals  causes  dissatisfaction,  or  even  misery. 

The  above  case  of  the  swallow  affords  an  illustration,  though 
of  a  reversed  nature,  of  a  temporary  though  for  the  time  strongly 
persistent  instinct  con(iuering  another  instinct,  which  is  usually 
dominant  over  all  others.  At  the  proper  season  these  birds 
seem  all  day  long  to  be  impressed  with  the  desire  to  migrate ; 
their  habits  change ;  they  become  restless,  are  noisy,  and  con- 
gregate in  flocks.  Whilst  the  mother-bird  is  feeding,  or  brooding 
over  her  nestlings,  the  maternal  instinct  is  probably  stronger 
than  the  migratory ;  but  the  instinct  wiiich  is  the  more  persis- 
tent gains  the  victory,  and  at  last,  at  a  moment  when  her  young 
ones  are  not  in  sight,  slic  takes  flight  and  deserts  them.  When 
arrived  at  the  end  of  her  long  journey,  and  the  migratory 
instinct  has  ceased  to  act,  what  an  agony  of  remorse  the  bird 
would  feel,  if,  from  being  endowed  with  great  mental  activity, 
she  could  not  prevent  the  image  constantly  passing  through  her 
mind,  of  her  young  ones  perishing  in  the  bleak  north  from  cold 
and  hunger. 


fined  as  hatred  of  another  for  some  had  done  him  an  injiirv  and  liad 
excellence  or  success;  and  Bacon  become  his  enemy,  iv'or  is  it  pro- 
insists  (Essay  ix.),  "  Of  all  otlier  bable  that  the  primitive  conscience 
"  affections  envy  is  the  most  im-  would  reproach  a  man  lor  injurin;-- 
"  port  une  and  continual."  Dogs  ai-e  his  enemy:  ratlier  it  would  re- 
very  apt  to  hate  both  strange  men  proach  him,  if  he  had  not  revenged 
and  strange  dogs,  esjiecially  if  they  himself.  To  do  good  in  return  lor 
live  near  at  hand,  but  do  not  beU)ng  evil,  to  love  your  enemy,  is  a  height 
to  the  same  family,  tribe,  or  clan;  of  morality  to  which  it  mav^be 
this  feeling  would  thus  seem  to  be  doubted  whether  the  social  instincts 
innate,  and  is  certainly  a  most  per-  would,  by  themselves,  have  ever  led 
sistent  one.  It  seems  to  be  the  us.  It  is  necessary  that  these  in- 
complement  and  converse  of  the  stincts,  together  with  sympathy, 
true  social  instinct.  From  what  should  have  been  highly  cultivated 
we  hear  of  savages,  it  would  appear  and  extended  l)y  the  aid  of  reason, 
that  something  of  the  same  kind  instruction,  and  the  love  or  fear  of 
holds  good  with  them.  If  this  be  God,  before  any  such  golden  rule 
so,  it  would  be  a  small  step  in  would  ever  be  thought  of  and 
any  one  to  transfer  such  feelings  to  obeyed. 
any  member  of  the  same  tribe  if  he 


1 14  The  Descent  of  Man.  Part  I. 

A-t  the  moment  of  action,  man  will  no  doubt  be  apt  to  follow 
the  stronger  impulse;  and  though  this  may  occasionally 
prompt  him  to  the  noblest  deeds,  it  will  more  commonly  lead 
him  to  gratify  his  own  desires  at  the  expense  of  other  men. 
But  after  their  gratification,  when  past  and  weaker  impressions 
are  judged  by  the  ever-enduring  social  instinct,  and  by  his  deep 
regard  for  the  good  opinion  of  his  fellows,  retribution  will  surely 
come.  He  will  then  feel  remorse,  repentance,  regret,  or  shame; 
this  latter  feeling,  however,  relates  almost  exclusively  to  the 
judgment  of  others.  He  will  consequently  resolve  more  or  less 
firmly  to  act  differently  for  the  future ;  and  this  is  conscience ;  for 
conscience  looks  backwards,  and  seiwes  as  a  guide  for  the  future. 

The  nature  and  strength  of  the  feelings  which  we  call  regret, 
shame,  repentance  or  remorse,  depend  apparently  not  only  on 
the  strength  of  the  violated  instinct,  but  partly  on  the  strength 
of  the  temptation,  and  often  still  more  on  the  judgment  of 
our  fellows.  How  far  each  man  values  the  appreciation  of 
others,  depends  on  the  strength  of  his  innate  or  acquired 
feeling  of  sympathy ;  and  on  his  own  capacity  for  reasoning  out 
the  remote  consequences  of  his  acts.  Another  element  is  most 
important,  although  not  necessary,  the  reverence  or  fear  of  the 
Gods,  or  Spirits  believed  in  by  each  man :  and  this  applies 
especially  in  cases  of  remorse.  Several  critics  have  objected 
that  though  some  shght  regret  or  repentance  may  be  explained 
by  the  view  advocated  in  this  chapter,  it  is  impossible  thns  to 
account  for  the  soul-shaking  feeling  of  remorse.  But  I  can  see 
little  force  in  this  objection.  My  critics  do  not  define  what 
they  mean  by  remorse,  and  I  can  find  no  definition  implying 
more  than  an  overwhelming  sense  of  repentance.  Eemorse 
seems  to  bear  the  same  relation  to  rej^entance,  as  rage  does  to 
anger,  or  agony  to  pain.  It  is  far  from  strange  that  an  instinct 
so  strong  and  so  generally  admired,  as  maternal  love,  should,  if 
disobeyed,  lead  to  the  deepest  misery,  as  soon  as  the  impression 
of  the  past  cause  of  disobedience  is  weakened.  Even  when  an 
action  is  opposed  to  no  special  instinct,  merely  to  know  that  our 
friends  and  equals  despise  ns  for  it  is  enough  to  cause  great 
misery.  Who  can  doubt  that  the  refusal  to  fight  a  duel  tlu-ough 
fear  has  caused  many  men  an  agony  of  shame  ?  Many  a  Hindoo, 
it  is  said,  has  been  stirred  to  the  bottom  of  his  soul  by  having 
partaken  of  unclean  food.  Here  is  another  case  of  what  must,  I 
think,  be  called  remorse.  Dr.  Landor  acted  as  a  magistrate  in 
West  Australia,  and  relates,"*  that  a  native  on  his  farm,  after 
losing  one  of  his  wives  from  disease,  came  and  said  that  "  he  was 

23  '  Insanity  in  Relation  to  Law;'  Ontario,  United  States,  1871,  p.  14. 


CuAP.  IV.  Moral  Soisc.  1 1 5 


"  going  to  a  distant  tribe  to  spear  a  woman,  to  satisfy  liis  sense 
''  of  duty  to  liis  wife.  I  told  liim  that  if  lie  did  so,  I  would 
"  send  him  to  prison  for  life.  He  remained  about  the  farm  for 
"  some  months,  but  got  exceedingly  thin,  and  complained  that 
"  he  could  not  rest  or  cat,  that  his  wife's  spirit  was  haunting 
"  him,  because  he  had  not  taken  a  life  for  hers.  I  was  in- 
"  exorable,  and  assured  him  that  nothing  should  save  him  if  he 
"  did."  Nevertheless  the  man  disappeared  for  more  than  a  year, 
and  then  returned  in  high  condition ;  and  his  other  wife  told 
Dr.  Landor  that  her  husband  had  taken  the  life  of  a  woman 
belonging  to  a  distant  tribe ;  but  it  was  impossible  to  obtain 
legal  evidence  of  the  act.  The  breach  of  a  rule  held  sacred  by 
the  tribe,  will  thus,  as  it  seems,  give  rise  to  the  deepest  feelings, 
—  and  this  quite  apart  from  the  social  instincts,  excepting  in  so 
far  as  the  rule  is  grounded  on  the  judgment  of  the  community. 
How  so  many  strange  superstitions  have  arisen  throughout  the 
world  we  know  not ;  nor  can  wc  tell  how  some  real  and  great 
crimes,  such  as  incest,  have  come  to  be  held  in  an  abhorrence 
(which  is  not  however  quite  universal)  by  the  lowest  savages.  It 
is  even  doubtful  whether  in  some  tribes  incest  would  be  looked  on 
with  greater  horror,  than  would  the  marriage  of  a  man  with  a 
woman  bearing  the  same  name,  though  not  a  relation.  "  To 
"  violate  this  law  is  a  crime  which  the  Australians  hold  in  the 
"  greatest  abhorrence,  in  this  agreeing  exactly  with  certain 
"  tribes  of  North  America.  When  the  question  is  put  in  either 
"  district,  is  it  worse  to  kill  a  girl  of  a  foreign  tribe,  or  to  marry 
"  a  girl  of  one's  own,  an  answer  just  opposite  to  oui's  would  be 
given  without  hesitation."  ^^  We  may,  therefore,  reject  the 
belief,  lately  insisted  on  by  some  writers,  that  the  abhorrence  of 
incest  is  due  to  our  possessing  a  special  God -implanted  con- 
science. On  the  whole  it  is  intelligible,  that  a  man  urged  by 
so  powerful  a  sentiment  as  remorse,  though  arising  as  above 
explained,  should  be  led  to  act  in  a  manner,  which  he  has  been 
taught  to  believe  serves  as  an  expiation,  such  as  delivering 
himself  up  to  justice. 

Man  prompted  by  his  conscience,  will  through  long  habit 
acquire  such  perfect  self-command,  that  his  desires  and  passions 
will  at  last  yield  instantly  and  without  a  struggle  to  his  social 
sympathies  and  instincts,  including  his  feeling  for  the  judgment  of 
his  fellows.  The  still  hungry,  or  the  still  revengeful  man  will  not 
think  of  stealing  food,  or  of  wreaking  his  vengeance.  It  is  possible, 
or  as  we  shall  hereafter  see,  even  probable,  that  the  habit  of  self- 
command  may,  like  other  habits,  be  inherited.   Thus  at  last  man 

'*  E.  B.  Tylor  iu  'Contemporary  Review,'  April,  1873,  p.  707. 


1 1 6  The  Descent  of  Man.  Part  I 

comes  to  feel,  through  acquired  and  perhaps  inherited  habit,  that 
it  is  best  for  him  to  obey  his  more  persistent  impulses.  The 
imperious  word  oz((/A^  seems  merely  to  imply  the  consciousness  of 
the  existence  of  a  rule  of  conduct,  however  it  may  have 
originated.  Formerly  it  must  have  been  often  vehemently 
urged  that  an  insulted  gentleman  ought  to  fight  a  duel.  We 
even  say  that  a  pointer  ouglit  to  point,  and  a  retriever  to 
retrieve  game.  If  they  fail  to  do  so,  they  fail  in  their  duty 
and  act  wrongly. 

If  any  desire  or  instinct  leading  to  an  action  opposed  to  the 
good  of  others  still  appears,  when  recalled  to  mind,  as  strong 
as,  or  stronger  than,  the  social  instinct,  a  man  will  feel  no  keen 
regret  at  having  followed  it ;  but  he  will  be  conscious  that  if  his 
conduct  were  known  to  his  fellows,  it  would  meet  with  their 
disapprobation ;  and  few  are  so  destitute  of  sympathy  as  not  to 
feel  discomfort  when  this  is  realised.  If  he  has  no  such 
sympathy,  and  if  his  desires  leading  to  bad  actions  are  at  the 
time  strong,  and  when  recalled  are  not  over-mastered  by  the 
persistent  social  instincts,  and  the  judgment  of  others,  then  he 
is  essentially  a  bad  man ;  ^"  and  the  sole  restraining  motive  left 
is  the  fear  of  punishment,  and  the  conviction  that  in  the  long 
run  it  would  be  best  for  his  own  selfish  interests  to  regard  the 
good  of  others  rather  than  his  own. 

It  is  obvious  that  every  one  may  with  an  easy  conscience 
gratify  his  own  desires,  if  they  do  not  interfere  with  his  social 
instincts,  that  is  with  the  good  of  others ;  but  in  order  to  be  quite 
free  from  self-rejDroach,  or  at  least  of  anxiety,  it  is  almost  neces- 
sary for  him  to  avoid  the  disapprobation,  whether  reasonable  or 
not,  of  his  fellow-men.  Nor  must  he  break  through  the  fixed 
habits  of  his  life,  especially  if  these  are  supported  by  reason; 
for  if  he  does,  he  will  assuredly  feel  dissatisfaction.  He  must 
likewise  avoid  the  reprobation  of  the  one  God  or  gods  in  whom, 
according  to  his  knowledge  or  superstition,  he  may  believe ;  but 
in  this  case  the  additional  fear  of  divine  punishment  often 
supervenes. 

The  strictly  Social  Virtues  at  first  alone  regarded. — The  above 
view  of  the  origin  and  nature  of  the  moral  sense,  which  tells  us 
what  we  ought  to  do,  and  of  the  conscience  which  reproves  us  if 
we  disobey  it,  accords  well  with  what  we  see  of  the  early  and 
undeveloped  condition  of  this  faculty  in  mankind.  The  virtues 
which  must  be  practised,  at  least  generally,  by  rude  men,  so 

*"  Dr.  Prosper  Despine,  in  his  many  curious  cases  of  the  worst 
'  Psychologic  Naturelle,'  1868  (torn.  criminals,  who  apparently  have  been 
I.  p,   243;    torn.    ii.  p.   1G9)    gives      entirely  destitute  of  conscience. 


CiiAp  IV.  Moral  Sense.  117 

tliat  they  may  associate  in  a  body,  arc  those  which  arc  still 
recognised  as  the  most  important.  But  they  are  practised 
almost  exclusively  in  relation  to  the  men  of  the  same  tribe ;  and 
their  opposites  are  not  regarded  as  crimes  in  relation  to  the  men 
of  other  tribes.  No  tribe  could  hold  together  if  murder, 
robbery,  treachery,  &c.,  were  common;  consequently  such 
crimes  within  the  limits  of  the  same  tribe  "are  branded  with 
"everlasting  infamy  ;"^^  but  excite  no  such  sentiment  beyond 
these  limits.  A  North-American  Indian  is  well  pleased  with 
himself,  and  is  honoured  by  others,  when  he  scalps  a  man  of 
another  tribe ;  and  a  Dyak  cuts  off  the  head  of  an  unoflfending 
person,  and  dries  it  as  a  trophy.  The  murder  of  infants  has 
prevailed  on  the  largest  scale  throughout  the  world,^^  and  has 
met  with  no  reproach  ;  but  infcinticide,  especially  of  females,  has 
been  thought  to  be  good  for  the  tribe,  or  at  least  not  injurious. 
Suicide  during  former  times  was  not  generally  considered  as  a 
crime,^^  but  rather,  from  the  courage  displayed,  as  an  honourable 
act ;  and  it  is  still  practised  by  some  semi-civilised  and  savage 
nations  without  reproach,  for  it  does  not  obviously  concern 
others  of  the  tribe.  It  has  been  recorded  that  an  Indian  Thug 
conscientiously  regretted  that  he  had  not  robbed  and  strangled 
as  many  travellers  as  did  his  father  before  him.  In  a  rude  state 
of  civilisation  the  robbery  of  strangers  is,  indeed,  generally 
considered  as  honourable. 

Slavery,  although  in  some  ways  beneficial  during  ancient 
times,^'*  is  a  great  crime  ;  yet  it  was  not  so  regarded  until  quite 
recently,  even  by  the  most  civilized  nations.  And  this  was 
especially  the  case,  because  the  slaves  belonged  in  general  to  a 
race  different  from  that  of  their  masters.  As  barbarians  do  not 
regard  the  opinion  of  their  women,  wives  are  commonly  treated 
like  slaves.  Most  savages  are  utterly  indiifcrcnt  to  the  sufferings 
of  strangers,  or  even  delight  in  witnessicg  them.     It  is  well 

3'  See    an    able    article    in    the  of   European    Morals,'  vol.  i.   18G9, 

*  North    British    Review,'  1867,    p.  p.  223.     With  respect    to    savages, 

395.     See    also    Mr.   W.    Bagehot/s  ^Mr.    Winwood    Keade    informs    me 

articles  on  the  Importance  of  Obe-  that    the    negroes    of    West    Africa 

dience   and  Coherence  to  Primitive  often    commit    suicide.     It    is    well 

Man,  in  the  'Fortnightly  Review,'  known  how  common  it  was  amongst 

1867,  p.  529,  and  1868,  p.  457,  &c.  the  miserable  aborigines    of  South 

^2  The    fullest    account  which    I  America,  after  the  Spanish  conquest, 

have  met  with  is  by  Dr.  Gerland,  in  For  New  Zealand,  see  the  voyage  of 

his    'Ueber     dan    Aussterben     der  the  "  Novara,"  and  for  the  Aleutian 

Naturvolker,'    1868;    but     I    shall  Islands,  Miiiler,  as  quoted  by  Hou- 

have    to   recur    to    the    subject    of  zeau,   '  Les  Facultes  Mentalcs,'  &c., 

infanticide  in  a  future  chapter.  torn.  ii.  p.  136. 

'^  See  the  very  interesting  discus-  ^^  Sec  Mr.  Bagehot,  '  Physics  and 

sicn  on  Suicide  in  Lecky's  'History  Politics,'  1872,  p.  72. 


1 18  The  Descent  of  Man,  Part  I. 

known  that  the  women  and  children  of  the  North- American 
Indians  aided  in  torturing  their  enemies.  Some  savages  take  a 
horrid  pleasure  in  cruelty  to  animals/^  and  humanity  is  an 
unknown  •virtue.  Nevertheless,  besides  the  family  affections, 
kindness  is  common,  especially  during  sickness,  between  the 
members  of  the  same  tribe,  and  is  sometimes  extended  beyond 
these  limits.  Mungo  Park's  touching  account  of  the  kindness  of 
the  negro  women  of  the  interior  to  him  is  well  known.  Many 
instances  could  be  given  of  the  noble  fidehty  of  savages  towards 
each  other,  but  not  to  strangers ;  common  experience  justifies 
the  maxim  of  the  Spaniard,  "Never,  never  trust  an  Indian." 
There  cannot  be  fidelity  without  truth  ;  and  this  fundamental 
virtue  is  not  rare  between  the  members  of  the  same  tribe  :  thus 
Mungo  Park  heard  the  negi-o  women  teaching  their  young 
childi-en  to  love  the  truth.  This,  again,  is  one  of  the  virtues 
which  becomes  so  deeply  rooted  in  the  mind,  that  it  is  sometimes 
practised  by  savages,  even  at  a  high  cost,  towards  strangers ;  but 
to  lie  to  your  enemy  has  rarely  been  thought  a  sin,  as  the  history 
of  modern  diplomacy  too  plainly  shews.  As  soon  as  a  tribe  has 
a  recognised  leader,  disobedience  becomes  a  crime,  and  even 
abject  submission  is  looked  at  as  a  sacred  virtue. 

As  during  rude  times  no  man  can  be  useful  or  faithful  to  his 
tribe  without  courage,  this  quality  has  universally  been  placed 
in  the  highest  rank;  and  although  in  civihsed  countries  a 
good  yet  timid  man  may  be  far  more  useful  to  the  community 
than  a  brave  one,  we  cannot  helj)  instinctively  honouring  the 
latter  above  a  coward,  however  benevolent.  Prudence,  on  the 
other  hand,  which  does  not  concern  the  welfare  of  others,  though 
a  very  useful  virtue,  has  never  been  highly  esteemed.  As  no 
man  can  practise  the  virtues  necessary  for  the  welfare  of  his 
tribe  without  self-sacrifice,  self-command,  and  the  power  of 
endurance,  these  qualities  have  been  at  all  times  highly  and 
most  justly  valued.  The  American  savage  voluntarily  submits 
to  the  most  horrid  tortui-es  without  a  groan,  to  prove  and 
strengthen  his  fortitude  and  courage;  and  we  cannot  help 
admiring  him,  or  even  an  Indian  Fakir,  who,  from  a  foolish 
religious  motive,  swings  suspended  by  a  hook  buried  in  his 
flesh. 

The  other  so  called  self-regarding  virtues,  which  do  not 
obviously,  though  they  may  really,  affect  the  welfare  of  the  tribe, 
have  never  been  esteemed  by  savages,  though  now  highly 
ax^preciated  by  civilised  nations.    The  greatest  intemperance 

^^  See,  for  instance,  Mi'.  Hamilton's  account  of  the  Kaffirs,  '  Anthro- 
pological Review,'  1870,  p.  xv. 


CiiAi'.  IV.  Moral  Sense.  119 

is  no  rcproach  with  savages.  Utter  licentiousness,  and  un- 
natural crimes,  prevail  to  an  astounding  extent.^^  As  soon, 
however,  as  marriage,  whether  polygamous,  or  monogamous, 
becomes  common,  jealousy  will  lead  to  the  inculcation  of  female 
virtue;  and  this,  being  honoured,  will  tend  to  spread  to  tlie 
unmarried  females.  How  slowly  it  spreads  to  the  male  sex, 
we  see  at  the  present  day.  Chastity  eminently  requires  self- 
command;  therefore  it  has  been  honoured  from  a  very  early 
period  in  the  moral  history  of  civilised  man.  As  a  consequence 
of  this,  the  senseless  practice  of  celibacy  has  been  ranked  from  a 
remote  period  as  a  virtue.^^  The  hatred  of  indecency,  which 
appears  to  us  so  natural  as  to  be  thought  innate,  and  which  is 
so  valuable  an  aid  to  chastity,  is  a  modern  virtue,  appertaining 
exclusively,  as  Sir  G.  Staunton  remarks,^^  to  civilised  life.  This 
is  shewn  by  the  ancient  religious  rites  of  various  nations,  by  the 
drawings  on  the  walls  of  Pompeii,  and  by  the  practices  of  many 
savages.  » 

We  have  now  seen  that  actions  are  regarded  by  savages,  and 
were  probably  so  regarded  by  primeval  man,  as  good  or  bad, 
solely  as  they  obviously  affect  the  welfare  of  the  tribe,— not  that 
of  the  si^ecies,  nor  that  of  an  individual  member  of  the  tribe. 
Tliis  conclusion  agrees  well  with  the  belief  that  the  so-called 
moral  sense  is  aboriginally  derived  from  the  social  instincts,  for 
both  relate  at  first  exclusively  to  the  community.  The  chief 
causes  of  the  low  morality  of  savages,  as  judged  by  our 
standard,  are,  firstly,  the  confinement  of  sympathy  to  the  same 
tribe.  Secondly,  powers  of  reasoning  insufficient  to  recognise 
the  bearing  of  many  virtues,  especially  of  the  self-regarding 
virtues,  on  the  general  welfare  of  the  tribe.  Savages,  for 
instance,  fail  to  trace  the  multiplied  evils  consequent  on  a 
want  of  temperance,  chastity,  &c.  And,  thirdly,  weak  power 
of  self-command ;  for  this  jDower  has  not  been  strengthened 
through  long-continued,  perhaps  inherited,  habit,  instruction  and 
religion. 

I  have  entered  into  the  above  details  on  the  immorality  of 
savages,"^  because  some  authors  have  recently  taken  a  high  view 
of  their  moral  nature,  or  have  attributed  most  of  their  crimes  to 
mistaken  benevolence.^^     These  authors  appear  to  rest  their 

^^      Jlr.     M'l.ennaa     has     given  ''^  '  Embassy  to  China,' vol.  ii.  p. 

('  Primitive     Marriage,'     1865,     p.  348. 

176)  a  good  collection  of  facts  on  ^^  See    on    this    subject    copious 

this  head.  evidence  in  Chap.  vii.  ol'  Sir  J.  Lub- 

•*'  Lecky,    'History   of    European  bock,  '  Origin  of  Civilisation,'  1870. 
Morals,' vol.  i.  1869,  p.  109.  "    For     instance    Lecky,    'Hist. 

European  Morals,'  vol.  i.  p.  12}-. 


1 20  The  Descent  of  Man.  Paet  T. 

conclusion  on  savages  possessing  those  virtues  which  are  ser- 
viceable, or  even  necessary,  for  the  existence  of  the  family  and  of 
the  tribe,  —qualities  which  they  undoubtedly  do  possess,  and  often 
in  a  high  degree. 

Concluding  Bemarhs. — It  was  assumed  formerly  by  philosophers 
of  the  derivative*^  school  of  morals  that  the  foundation  of  morality 
lay  in  a  form  of  Selfishness ;  but  more  recently  the  "  Greatest 
"  happiness  principle  "  has  been  brought  prominently  forward. 
It  is,  however,  more  correct  to  speak  of  the  latter  principle  as 
the  standard,  and  not  as  the  motive  of  conduct.  Nevertheless,  all 
the  authors  whose  works  I  have  consulted,  with  a  few  excep- 
tions,*^ write  as  if  there  must  be  a  distinct  motive  for  every 
action,  and  that  this  must  be  associated  with  some  pleasure  or 
displeasure.  But  man  seems  often  to  act  impulsively,  that  is 
from  instinct  or  long  habit,  without  any  consciousness  of  pleasure, 
in  the  same  manner  as  does  probably  a  bee  or  ant,  when  it 
blindly  follows  its  instincts.  Under  circumstances  of  extreme 
peril,  as  during  a  fire,  when  a  man  endeavours  to  save  a  fellow- 
creature  without  a  moment's  hesitation,  he  can  hardly  feel 
pleasure ;  and  still  less  has  he  time  to  reflect  on  the  dissatisfaction 
which  he  might  subsequently  experience  if  he  did  not  make  the 
attempt.  Should  he  afterwards  reflect  over  his  own  conduct,  he 
would  feel  that  there  lies  within  him  an  impulsive  power  widely 
different  from  a  search  after  pleasure  or  happiness;  and  this 
seems  to  be  the  deeply  planted  social  instinct. 

In  the  case  of  the  lower  animals  it  seems  much  more  appro- 
priate to  speak  of  their  social  instincts,  as  having  been  developed 

•*^  This  term  is  used   in  an  able  "  noss  extra -regarding  impulse,  di- 

article  in  the  'Westminster  Review,'  "  recied  towards  something  that  is 

Oct.  1869,  p.  498.   For  the  "  Greatest  "not  pleasure;  that  in  many  cases 

"  happiness  principle,"  see  J.  S.  Mill,  "  the  impulse  is  so  far  incompatible 

'  Utilitarianism,'  p.  17.  "  with  the  self-regarding  that  the 

^2    Mill    recognises   ('  Sj'stem    of  "  two  do  not  easily  co-exist  in  the 

Logic,'  vol.  ii.,  p.  422)  in  the  clearest  "  same   moment    of  consciousness." 

manner,   that    actions  mny  be  per-  A  dim  feeling  that  our  impulses  do 

formed  through  habit  without  the  not  by  any  means  always  arise  from 

anticipation    of  pleasure.      Mr.    H.  any  contemporaneous  or  anticipated 

Sidgwick    also,     in     his     Essay     on  pleasure,  has,  I  cannot  but   think. 

Pleasure    and    Desire    ('The    Con-  been  one  chief  cause  of  the  accept- 

temporary  Review,'  April  1872,  p.  ance    of    the    intuitive    theory    of 

671),  remarks:     "To    sum    up,  in  morality,  and  of  the  rejection  of  the 

"  contravention  of  the  doctrine  that  utilitarian  or  "  Greatest  happiness  '' 

"  our  conscious  active  impulses  are  theory.     With  respect  to  the  latter 

"  always  directed  towards  the  pro-  theory,  the  standard  and  the  motive 

"  duction  of  agreeable  sensations  in  of  conduct  have  no  doubt  often  been 

"  ourselves,  I  would  maintain  that  confused,    but    they   are   really   in 

"  we  find  everywhere  in  conscious-  some  degree  blended. 


Chap.  IV.  Mo7'al  Sense.  1 2 1 


for  the  general  good  rather  than  for  the  general  happiness  of  the 
species.  The  term,  general  good,  may  be 'defined  as  the  rearing 
of  the  greatest  number  of  individuals  in  full  vigour  and  health, 
with  all  their  faculties  perfect,  under  the  conditions  to  which 
they  are  subjected.  As  the  social  instincts  both  of  man  and  the 
lower  animals  have  no  doubt  been  developed  by  nearly  the  same 
steps,  it  would  be  advisable,  if  found  practicable,  to  use  the 
same  definition  in  both  cases,  and  to  take  as  the  standard  of 
morality,  the  general  good  or  welfare  of  tlie  community,  rather 
than  the  general  happiness ;  but  this  definition  would  perhaps 
require  some  limitation  on  account  of  political  ethics. 

When  a  man  risks  his  life  to  save  that  of  a  fellow-creature,  it 
seems  also  more  correct  to  say  that  he  acts  for  the  general  good, 
rather  than  for  the  general  happiness  of  mankind.  No  doubt 
the  welfare  and  the  happiness  of  the  individual  usually  coincide ; 
and  a  contented,  happy  tribe  will  flourish  better  than  one  that 
is  discontented  and  unhappy.  We  have  seen  that  even  at  an 
early  period  in  the  history  of  man,  the  expressed  wishes  of  the 
community  will  have  naturally  influenced  to  a  large  extent  the 
conduct  of  each  member;  and  as  all  wish  for  happiness,  the 
"greatest  happiness  principle"  will  have  become  a  most  im- 
portant secondary  guide  and  object ;  the  social  instinct,  however, 
together  with  sympathy  (which  leads  to  our  regarding  the 
approbation  and  disapprobation  of  others),  having  served  as  the 
primary  impulse  and  guide.  Thus  the  reproach  is  removed  of 
laying  the  foundation  of  the  noblest  part  of  our  nature  in  the 
base  principle  of  selfishness;  unless,  indeed,  the  satisfaction 
which  every  animal  feels,  when  it  follows  its  proper  instincts, 
and  the  dissatisfaction  felt  when  prevented,  be  called  selfish. 

The  wishes  and  opinions  of  the  members  of  the  same  community, 
expressed  at  first  orally,  but  later  by  writing  also,  either  form 
the  sole  guides  of  our  conduct,  or  greatly  reinforce  the  social 
instincts ;  such  opinions,  however,  have  sometimes  a  tendency 
directly  opposed  to  these  instincts.  This  latter  fact  is  well 
exemplified  by  the  Law  of  Honour,  that  is,  the  law  of  the  opinion 
of  our  equals,  and  not  of  all  our  countrymen.  The  breach  of 
this  law,  even  when  the  breach  is  known  to  be  strictly  accordant 
with  true  morality,  has  caused  many  a  man  more  agony  than  a 
real  crime.  We  recognise  the  same  influence  in  the  burning 
sense  of  shame  which  most  of  us  have  felt,  even  after  the  interval 
of  years,  when  calling  to  mind  some  accidental  breach  of  a 
trifling,  though  fixed,  rule  of  etiquette.  The  judgment  of  the 
community  will  generally  be  guided  by  some  rude  experience  of 
what  is  best  in  the  long  run  for  all  the  members ;  but  this  judg- 
ment will  not  rarely  err  from  ignorance  and  weak  i^owers  of 


122  The  Descent  of  Man.  Part  I. 


reasoning.  Hence  the  strangest  cnstoms  and  superstitions,  in 
complete  opposition  to*  tlie  true  welfare  and  happiness  of  man- 
kind, have  become  all-powerful  throughout  the  world.  We  see 
this  in  the  horror  felt  by  a  Hindoo  who  breaks  his  caste,  and 
in  many  other  such  cases.  It  would  be  difficult  to  distinguish 
between  the  remorse  felt  by  a  Hindoo  who  has  yielded  to  the 
temptation  of  eating  unclean  food,  from  that  felt  after  committing 
a  theft ;  but  the  former  would  probably  be  the  more  severe. 

How  so  many  absurd  rules  of  conduct,  as  well  as  so  many 
absurd  religious  beliefs,  have  originated,  we  do  not  know ;  nor 
how  it  is  that  they  have  become,  in  all  quarters  of  the  world,  so 
deeply  impressed  on  the  mind  of  men;  but  it  is  worthy  of 
remark  that  a  belief  constantly  inculcated  during  the  early  years 
of  life,  whilst  the  brain  is  impressible,  appears  to  acquire  almost 
the  nature  of  an  instinct ;  and  the  very  essence  of  an  instinct  is 
that  it  is  followed  independently  of  reason.  Neither  can  we  say 
why  certain  admirable  Yirtues;  such  as  the  love  of  truth,  are 
much  more  highly  appreciated  by  some  savage  tribes  than  by 
others  ;^^  nor,  again,  why  similar  diiferences  prevail  even  amongst 
highly  civihsed  nations.  Knowing  how  firmly  fixed  many 
strange  customs  and  superstitions  have  become,  we  need  feel  no 
surprise  that  the  self-regarding  virtues,  supported  as  they  are  by 
reason,  should  now  appear  to  us  so  natural  as  to  be  thought 
innate,  although  they  were  not  valued  by  man  in  his  early 
condition. 

Notwithstanding  many  sources  of  doubt,  man  can  generally 
and  readily  distinguish  between  the  higher  and  lower  moral 
rules.  The  higher  are  founded  on  the  social  instincts,  and  relate 
to  the  welfare  of  others.  They  are  supported  by  the  approbation 
of  our  fellow-men  and  by  reason.  The  lower  rules,  though  some 
of  them  when  implying  self-sacrifice  hardly  deserve  to  be  called 
lower,  relate  chiefly  to  self,  and  arise  from  public  opinion,  ma- 
tured by  experience  and  cultivation ;  for  they  are  not  practised 
by  rude  tribes. 

As  man  advances'  in  civilisation,  and  small  tribes  are  united 
into  .  larger  communities,  the  simplest  reason  would  tell  each 
individual  that  he  ought  to  extend  his  social  instincts  and 
sympatlries  to  all  the  members  of  the  same  nation,  though 
personally  unknown  to  him.  This  point  being  once  reached, 
there  is  only  an  artificial  barrier  to  prevent  his  sympathies 
extending  to  the  men  of  all  nations  and  races.  If,  indeed,  such 
men  are  separated  from  him  by  great  differences  in  appearance 

*^  Good  instances  are  given  by  in  his 'Contributions  to  the  Theory 
Mr.  Wallace  in  'Scientific  Opinion/  of  Natural  Selection,'  1870,  p.  353. 
Sept.    15,    1869  ;    and    more   fully 


Chap.  IV.  Moral  Sense.  123 


or  habits,  experience  ■unfortunately  shews  us  how  long  it  is, 
before  we  look  at  them  as  our  fellow-creatures.  Sympathy 
beyond  the  confines  of  man,  that  is,  humanity  to  the  lower 
animals,  seems  to  be  one  of  the  latest  moral  acquisitions.  It  is 
apparently  unfelt  by  savages,  except  towards  their  pets.  How 
little  the  old  Romans  knew  of  it  is  shewn  by  their  abhorrent 
gladiatorial  exhibitions.  The  very  idea  of  humanity,  as  tar  as  I 
could  observe,  w^as  new  to  most  of  the  Gauchos  of  the  Pampas. 
This  virtue,  one  of  the  noblest  with  which  man  is  endowed, 
seems  to  arise  incidentally  from  our  sympathies  becoming  more 
tender  and  more  widely  diffused,  until  they  are  extended  to  all 
sentient  beings.  As  soon  as  this  virtue  is  honoured  and  practised 
by  some  few  men,  it  spreads  through  instruction  and  example 
to  the  young,  and  eventually  becomes  incorporated  in  public 
opinion . 

The  highest  possible  stage  in  moral  culture  is  when  we  re- 
cognise that  w^c  ought  to  control  our  thoughts,  and  "  not  even  in 
"  inmost  thought  to  think  again  the  sins  that  made  the  past  so 
"  pleasant  to  us."  " '  Whatever  makes  any  bad  action  familiar  to 
the  mind,  renders  its  performance  by  so  much  the  easier.  As 
Marcus  Aurelius  long  ago  said,  "Such  as  are  thy  habitual 
"  thoughts,  such  also  will  be  the  character  of  thy  mind  ;  for  the 
"  soul  is  dyed  by  the  thoughts."  ^^ 

Our  great  philosopher,  Herbert  Spencer,  has  recently  explained 
his  views  on  the  moral  sense.  He  says,'*'^  "  I  believe  that  the 
"  experiences  of  utility  organised  and  consolidated  through  all 
"  past  generations  of  the  human  race,  have  been  producing. 
"  corresponding  modifications,  which,  by  continued  transmission 
"  and  accumulation,  have  become  in  us  certain  faculties  of 
"  moral  intuition— certain  emotions  responding  to  right  and 
"  wrong  conduct,  w^hich  have  no  apparent  basis  in  the  individual 
"  experiences  of  utility."  There  is  not  the  least  inherent 
improbability,  as  it  seems  to  me,  in  virtuous  tendencies  being 
more  or  less  strongly  inherited;  for,  not  to  mention  the  various 
dispositions  and  habits  transmitted  by  many  of  our  domestic 
animals  to  their  offspring,  I  have  heard  of  authentic  cases  in 
which  a  desire  to  steal  and  a  tendency  to  lie  appeared  to  run 
in  families  of  the  upper  ranks ;  and  as  stealing  is  a  rare  crime  in 
the  wealthy  classes,  we  can  hardly  account  by  accidental  coinci- 
dence for  the  tendency  occurring  in  two  or  three  members  of 

^^  Tennyson,  'Idylls  of  the  King,*  Aurelius  was  born  a.d.  121. 

p.  244.  ••«  Letter    to  Mr.   Mill    in  B;tiu's 

**  '  The  Thoughts  of  the  Emperor  '  Mental  and  Moral  Science,'  18(J8, 

M.  Aurelius  Antoninus,'  Eng.  trans-  p.  722. 
lat.,  2ud  edit.,  1869,  p.  112.    Marcus 


124  The  Descejtt  of  Man.  Paet  I. 

the  same  family.  If  bad  tendencies  are  transmitted,  it  is  pro- 
bable that  good  ones  are  likewise  transmitted.  That  the  state 
of  the  body  by  affecting  the  brain,  has  great  influence  on  the 
moral  tendencies  is  known  to  most  of  those  who  have  suffered 
from  chronic  derangements  of  the  digestion  or  liver.  The  same 
fact  is  likewise  shewn  by  the  "  perversion  or  destruction  of  the 
"  moral  sense  being  often  one  of  the  earliest  symptoms  of  mental 
"derangement;"^^  and  insanity  is  notoriously  often  inherited. 
Except  through  the  principle  of  the  transmission  of  moral  ten- 
dencies, we  cannot  understand  the  differences  believed  to  exist  in 
this  respect  between  the  various  races  of  mankind. 

Even  the  partial  transmission  of  virtuous  tendencies  would 
be  an  immense  assistance  to  the  primary  impulse  derived  directly 
and  indirectly  from  the  social  instincts.  Admitting  for  a  moment 
that  virtuous  tendencies  are  inherited,  it  appears  probable,  at 
least  in  such  cases  as  chastity,  temperance,  humanity  to  animals, 
&c.,  that  they  become  first  impressed  on  the  mental  organization 
through  habit,  instruction  and  example,  continued  during 
several  generations  in  the  same  family,  and  in  a  quite  subordinate 
degree,  or  not  at  all,  by  the  individuals  possessing  such  virtues 
having  succeeded  best  in  the  struggle  for  life.  My  chief  source 
of  doubt  with  respect  to  any  such  inheritance,  is  that  senseless 
customs,  superstitions,  and  tastes,  such  as  the  horror  of  a  Hindoo 
for  unclean  food,  ought  on  the  same  principle  to  be  transmitted. 
I  have  not  met  with  any  evidence  in  support  of  the  transmission 
of  superstitious  customs  or  senseless  habits,  although  in  itself  it 
is  perhaps  not  less  probable  than  that  animals  should  acquire 
inherited  tastes  for  certain  kinds  of  food  or  fear  of  certain  foes. 

Finally  the  social  instincts,  Avhich  no  doubt  were  acquired  by 
man  as  by  the  lower  a-nimals  for  the  good  of  the  community, 
will  from  the  first  have  given  to  him  some  wish  to  aid  his 
fellows,  some  feeling  of  sympathy,  and  have  compelled  him  to 
regard  their  approbation  and  disapprobation.  Such  impulses 
will  have  served  him  at  a  very  early  period  as  a  rude  rule  of 
right  and  wrong.  But  as  man  gradually  advanced  in  intellectual 
power,  and  was  enabled  to  trace  the  more  remote  consequences 
of  his  actions ;  as  he  acquired  sufficient  knowledge  to  reject 
baneful  customs  and  superstitions;  as  he  regarded  more  and 
more,  not  only  the  welfare,  but  the  happiness  of  his  fellow-men ; 
as  from  habit,  following  on  beneficial  experience,  instruction 
and  example,  his  sympathies  became  more  tender  and  widely 
diffused,  extending  to  men  of  all  races,  to  the  imbecile,  maimed, 

*^  Maudsley,  'Body  and  Mind,'  1870,  p.  60. 


Chap.  IV.  Summary.  125 

and  otlier  useless  members  of  society,  and  finally  to  the  lower 
animals,— so  would  the  standard  of  his  morality  rise  higher  and 
higher.  And  it  is  admitted  by  moralists  of  the  derivative 
school  and  by  some  intuitionists,  that  the  standard  of  morality 
has  risen  since  an  early  period  in  the  history  of  man/** 

As  a  struggle  may  sometimes  be  seen  going  on  between  the 
various  instincts  of  the  lower  animals,  it  is  not  surprising  that 
there  should  be  a  struggle  in  man  between  his  social  instincts, 
with  their  derived  virtues,  and  his  lower,  though  momentarily 
stronger  impulses  or  desires.  This,  as  Mr.  Galton*^  has  remarked, 
is  all  the  less  surprising,  as  man  has  emerged  from  a  state  of 
barbarism  within  a  comparatively  recent  period.  After  having 
yielded  to  some  temptation  we  feel  a  sense  of  dissatisfaction, 
shame,  repentance,  or  remorse,  analogous  to  the  feelings  caused 
by  other  iwwcrful  instincts  or  desires,  when  left  unsatisfied  or 
baulked.  We  compare  the  weakened  impression  of  a  past 
temptation  with  the  ever  present  social  instincts,  or  with  habits, 
gained  in  early  youth  and  strengthened  during  our  whole  lives, 
until  they  have  become  almost  as  strong  as  instincts.  If  with 
the  temptation  still  before  us  we  do  not  yield,  it  is  because 
either  the  social  instinct  or  some  custom  is  at  the  moment 
predominant,  or  because  we  have  learnt  that  it  will  appear  to  us 
hereafter  the  stronger,  when  comj)ared  with  the  weakened  im- 
pression of  the  temptation ,  and  we  realise  that  its  violation  would 
cause  us  suffering.  Looking  to  future  generations,  there  is  no 
cause  to  fear  that  the  social  instincts  will  grow  weaker,  and  we 
may  expect  that  virtuous  habits  will  grow  stronger,  becoming 
perhaps  fixed  by  inheritance.  In  this  case  the  struggle  between 
our  higher  and  lower  impulses  will  be  less  severe,  and  virtue 
will  be  triumphant. 

Summary  of  the  last  two  Chapters.— Thci'O  can  be  no  doubt  that 
the  difference  between  the  mind  of  the  lowest  man  and  that  of 
the  highest  animal  is  immense.  An  anthropomorphous  ape,  if 
he  could  take  a  dispassionate  view  of  his  own  case,  would  admit 
that  though  he  could  form  an  artful  plan  to  plunder  a  garden — 
though  he  could  use  stones  for  fighting  or  for  breaking  open 
nuts,  yet  that  the  thought  of  fashioning  a  stone  into  a  tool  was 

■•'  A  writer  in  the  'North  British  coincide  therein. 

Review'  (July  1869,  p.  o.'jI),  well  *"  See    his    remarkable  work    on 

capable   of   forming  a  sound   judg-  '  Hereditary  Genius,*  18G9,  p.  349. 

ment,  expresses  himself  strongly  in  The    Duke    of    Argyll    ('  Primeval 

favour    of     this    conclusion.      Mr.  Man,'  1809,  p.  188)  has  some  good 

Lecky  ('Hist,  of  Morals,'   vol.  i.  p.  remarks   on    the    contest    in    man's 

143)  seems  to  a  certain  extent  to  nature  between  right  and  wrong. 


1 26  The  Descent  of  Man.  Part  I. 


quite  beyond  liis  scope.  Still  less,  as  he  would  admit,  could  lie 
follow  out  a  train  of  metaphysical  reasoning,  or  solve  a  mathe- 
matical problem,  or  reflect  on  God,  or  admire  a  grand  natural 
scene.  Some  apes,  however,  would  probably  declare  that  they 
could  and  did  admire  the  beauty  of  the  coloured  skin  and  fur  of 
their  partners  in  marriage.  They  would  admit,  that  though  they 
could  make  other  apes  understand  by  cries  some  of  their  per- 
ceptions and  simpler  wants,  the  notion  of  expressing  definite  ideas 
by  definite  sounds  had  never  crossed  theii'  minds.  They  might 
insist  that  they  were  ready  to  aid  their  fellow-apes  of  the  same 
troop  in  many  ways,  to  risk  their  lives  for  them,  and  to  take 
charge  of  their  orphans ;  but  they  would  be  forced  to  acknow- 
ledge that  disinterested  love  for  all  living  creatures,  the  most 
noble  attribute  of  man,  was  quite  beyond  their  comprehension. 

Nevertheless  the  difference  in  mind  between  man  and  the 
higher  animals,  great  as  it  is,  certainly  is  one  of  degree  and  not 
of  kind.  We  have  seen  that  the  senses  and  intuitions,  the 
various  emotions  and  faculties,  such  as  love,  memory,  attention, 
curiosity,  imitation,  reason,  &c.,  of  w^hich  man  boasts,  may  be 
found  in  an  incipient,  or  even  sometimes  in  a  well- developed 
condition,  in  the  lower  animals.  They  are  also  capable  of  some 
inherited  improvement,  as  we  see  in  the  domestic  dog  compared 
with  the  wolf  or  jackal.  If  it  could  be  proved  that  certain  high 
mental  powers,  such  as  the  formation  of  general  concepts,  self-con- 
sciousness, &c.,  were  absolutely  peculiar  to  man,  which  seems 
extremely  doubtful,  it  is  not  improbable  that  these  qualities  are 
merely  the  incidental  results  of  other  highly-advanced  intel- 
lectual faculties;  and  these  again  mainly  the  result  of  the 
continued  use  of  a  perfect  language.  At  what  age  does  the 
new-born  infant  possess  the  power  of  abstraction,  or  become 
self-conscious,  and  reflect  on  its  own  existence  ?  We  cannot 
answer ;  nor  can  we  answer  in  regard  to  the  ascending  organic 
scale.  The  half-art,  half-instinct  of  language  still  bears  the 
stamp  of  its  gradual  evolution.  The  ennobhng  belief  in  God  is 
not  universal  with  man;  and  the  belief  in  spiritual  agencies 
naturally  follows  from  other  mental  powers.  The  moral  sense 
perhaps  affords  the  best  and  highest  distinction  between  man  and 
the  lower  animals ;  but  I  need  say  nothing  on  this  head,  as  I 
have  so  lately  endeavoured  to  shew  that  the  social  instincts, — 
the  prime  principle  of  man's  moral  constitution  ^" — with  the  aid 
of  active  intellectual  powers  and  the  effects  of  habit,  naturally  lead 
to  the  golden  rule,  "  As  ye  would  that  men  should  do  to  you,  do 
ye  to  them  likewise ;''  and  this  lies  at  the  foundation  of  morality. 

so  '  The  Tlioughts  of  Marcus  Aurelius,'  &c.,  p.  139. 


TiiAP.  V.  Ijitellcctnal  Faculties.  1 27 

In  the  next  chapter  I  shall  make  some  few  remarks  on  the 
probable  steps  and  means  by  which  the  several  mental  and  moral 
faculties  of  man  have  been  gradually  evolved.  That  such  evolu- 
tion is  at  least  possible,  ought  not  to  be  denied,  for  we  daily  sec 
these  faculties  developing  in  every  infant ;  and  we  may  trace  a 
perfect  gradation  from  tlie  mind  of  an  utter  idiot,  lower  than 
that  of  an  animal  low  in  the  scale,  to  the  mind  of  a  Newton. 


CHAPTER  V. 

On  the  Development  of  the  Intellectual  and  Moral 
Faculties  during  Primeval  and  Civilised  Times. 

Ailvanceniont  of  the  intellectual  powers  through  natural  selection — 
Importance  of  imitation — Social  and  moral  faculties — Their  develop- 
ment within  the  limits  of  the  same  tribe — Natural  selection  as  affecting 
civilised  nations — Evidence  that  civilised  nations  were  once  barbarous. 

The  subjects  to  be  discussed  in  this  chapter  are  of  the  highest 
interest,  but  are  treated  by  me  in  an  imperfect  and  fragmentary 
manner.  Mr.  Wallace,  in  an  admirable  paper  before  referred  to,' 
argues  that  man,  after  he  had  partially  acquired  those  intel- 
tectual  and  moral  faculties  which  distinguish  him  from  the 
lower  animals,  would  have  been  but  little  liable  to  bodily 
modifications  through  natural  selection  or  any  other  means. 
For  man  is  enabled  through  his  mental  faculties  ''  to  keep  with 
"  an  unchanged  body  in  harmony  with  the  changing  universe." 
He  has  great  power  of  adapting  his  habits  to  new  conditions  of 
life.  He  invents  weapons,  tools,  and  various  stratagems  to 
procure  food  and  to  defend  himself.  When  he  migrates  into  a 
colder  climate  he  nses  clothes,  builds  sheds,  and  makes  fires ; 
and  by  the  aid  of  fire  cooks  food  otherwise  indigestible.  Ho 
aids  his  fellow-men  in  many  ways,  and  anticipates  future  events 
Even  at  a  remote  period  he  practised  some  division  of  labour. 

The  lower  animals,  on  the  other  hand,  must  have  their  bodily 
structure  modified  in  order  to  survive  under  greatly  changed 
conditions.  They  must  be  rendered  stronger,  or  acquire  more 
effective  teeth  or  claws,  for  defence  against  new  enemies;  or 
they  must  be  reduced  in  size,  so  as  to  escape  detection  and 
danger.  When  they  migrate  into  a  colder  climate,  they  must 
become  clothed  with  thicker  fur,  or  have  their  constitutions 
altered.  If  they  fail  to  be  thus  modified,  they  will  cease  to 
exist 

•  '  Antliropological  Review,'  ^'^ay  13G4,  p.  clviii. 


128  The  Descent  of  Man.  Part  I. 


The  case,  however,  is  widely  different,  as  Mr.  Wallace  has 
with  justice  inssted,  in  relation  to  the  intellectnal  and  moral 
faculties  of  man.  These  faculties  are  variable;  and  we  have 
every  reason  to  believe  that  the  variations  tend  to  be  inherited. 
Therefore,  if  they  were  formerly  of  high  importance  to  primeval 
man  and  to  his  ape-like  progenitors,  they  would  have  been 
perfected  or  advanced  through  natural  selection.  Of  the  high 
importance  of  the  intellectual  faculties  there  can  be  no  doubt, 
for  man  mainly  owes  to  them  his  predominant  position  in  the 
world.  We  can  see,  that  in  the  rudest  state  of  society,  the 
individuals  who  were  the  most  sagacious,  who  invented  and  used 
the  best  weapons  or  traps,  and  who  were  best  able  to  defend 
themselves,  would  rear  the  gi-eatest  number  of  offspring.  The 
tribes,  which  included  the  largest  number  of  men  thus  endowed, 
would  increase  in  number  and  supplant  other  tribes.  Numbers 
depend  primarily  on  the  means  of  subsistence,  and  this  depends 
partly  on  the  physical  nature  of  the  country,  but  in  a  much  higher 
degree  on  the  arts  which  are  there  practised.  As  a  tribe  increases 
and  is  victorious,  it  is  often  still  further  increased  by  the  ab- 
sorption of  other  tribes.^  The  stature  and  strength  of  the  men 
of  a  tribe  are  likewise  of  some  importance  for  its  success,  and 
these  depend  in  part  on  the  nature  and  amount  of  the  food  which 
can  be  obtained.  In  Europe  the  men  of  the  Bronze  period  were 
supplanted  by  a  race  more  powerful,  and,  judging  from  their 
sword-handles,  with  larger  hands  f  but  their  success  was  pro- 
bably still  more  due  to  their  superiority  in  the  arts. 

All  that  we  Imow  about  savages,  or  may  infer  from  their 
traditions  and  from  old  monuments,  the  history  of  which  is  quite 
forgotten  by  the  present  inhabitants,  shew  that  from  the  remotest 
times  successful  tribes  have  supplanted  other  tribes.  Eelics  of 
extinct  or  forgotten  tribes  have  been  discovered  throughout  the 
civilised  regions  of  the  earth,  on  the  wild  plains  of  America,  and 
on  the  isolated  islands  in  the  Pacific  Ocean.  At  the  i^resent  day 
civilised  nations  are  everywhere  supplanting  barbarous  nations, 
excepting  where  the  climate  opposes  a  deadly  barrier;  and  they 
succeed  mainly,  though  not  exclusively,  through  their  arts,  which 
are  the  products  of  the  intellect.  It  is,  therefore,  highly  probal^le 
that  with  mankind  the  intellectual  faculties  have  been  mainly 
and  gradually  perfected  through  natural  selection;  and  this  con- 
clusion is  sufficient  for  our  purpose.  Undoubtedly  it  would 
be  interesting  to  trace  the  development  of  each  separate  faculty 

2  After    a   time  the  members  or  1861,  p.  131),  that  they  are  the  co- 
tribes     which     are     absorbed     into  descendants  of  the  same  ancestors, 
another  tribe  assume,  as  Sir  Henry  ^  Morlot,   '  Soc.   Vaud.   Sc.    Nat.' 
Maine    remarks     ('Ancient     Law,'  1860,  p.  294. 


Chap.  V.  Moral  Faculties.  129 


from  the  state  in  which  it  exists  in  the  lower  animals  to  that  in 
which  it  exists  in  man ;  but  neither  my  ability  nor  knowledge 
permits  the  attempt. 

It  deserves  notice  that,  as  soon  as  the  progenitors  of  man 
became  social  (and  this  probably  occurred  at  a  very  early  period), 
the  principle  of  imitation,  and  reason,  and  experience  would 
have  increased,  and  much  modified  the  intellectual  powers  in  a 
way,  of  which  we  see  only  traces  in  the  lower  animals.  Apes  are 
much  given  to  imitation,  as  are  the  lowest  savages;  and  the 
simple  fact  previously  referred  to,  that  after  a  time  no  animal 
can  be  caught  in  the  same  place  by  the  same  sort  of  trap,  shews 
that  animals  learn  by  experience,  and  imitate  the  caution  of 
others.  Now,  if  some  one  man  in  a  tribe,  more  sagacious  than 
the  others,  invented  a  new  snare  or  weapon,  or  other  means  of 
attack  or  defence,  the  plainest  self-interest,  without  the  assistance 
of  much  reasoning  power,  would  prompt  the  other  members  to 
imitate  him ;  and  all  would  thus  profit.  The  habitual  practice 
of  each  new  art  must  likewise  in  some  slight  degree  strengthen 
the  intellect.  If  the  new  invention  were  an  important  one,  the 
tribe  would  increase  in  number,  spread,  and  sujDplant  other 
tribes.  In  a  tribe  thus  rendered  more  numerous  there  would 
always  be  a  rather  greater  chance  of  the  birth  of  other  superior 
and  inventive  members.  If  such  men  left  children  to  inherit 
their  mental  superiority,  the  chance  of  the  birth  of  still  more 
ingenious  members  would  be  somewhat  better,  and  in  a  very 
small  tribe  decidedly  better.  Even  if  they  left  no  children,  the 
tribe  would  still  include  their  blood- relations ;  and  it  has  been 
ascertained  by  agriculturists  *  that  by  preserving  and  breeding 
from  the  family  of  an  animal,  which  when  slaughtered  was 
found  to  be  valuable,  the  desired  character  has  been  obtained. 

Turning  now  to  the  social  and  moral  faculties.  In  order  that 
primeval  men,  or  the  ape-like  i)regenitors  of  man,  should  become 
social,  they  must  have  acquired  the  same  instinctive  feelings, 
which  impel  other  animals  to  live  in  a  body ;  and  they  no  doubt 
exhibited  the  same  general  disposition.  They  would  have  felt 
uneasy  when  separated  from  their  comrades,  for  whom  they 
would  have  felt  some  degree  of  love ;  they  would  have  warned 
each  other  of  danger,  and  have  given  mutual  aid  in  attack  or 
defence.  All  this  implies  some  degree  of  sympathy,  fidelity,  and 
courage.  Such  social  qualities,  the  paramount  importance  of 
which  to  the  lower  animals  is  disputed  by  no  one,  were  no  doubt 

*  I  have  given  iustanccs  in  my  *  Variation  of  Animals  ujiJer  Domestica- 
tion,' vol.  ii.  p.  196. 

7 


1 30  The  Descent  of  Man.  Part  1. 

acquired  by  the  progenitors  of  man  in  a  similar  manner^  namely, 
through  natural  selection,  aided  by  inherited  habit.  When  two 
tribes  of  primeval  man,  living  in  the  same  country,  came  into 
competition,  if  (other  circumstances  being  equal)  the  one  tribe 
included  a  gi-eat  number  of  courageous,  sjTnpathetic  and  faithful 
members,  who  were  always  ready  to  warn  each  other  of  danger, 
to  aid  and  defend  each  other,  this  tribe  would  succeed  better  and 
conquer  the  other.  Let  it  be  borne  in  mind  how  all-important 
in  the  never-ceasing  wars  of  savages,  fidehty  and  courage  must 
be.  The  advantage  which  disciplined  soldiers  have  over  undis- 
ciplined hordes  follows  chiefly  from  the  confidence  which  each 
man  feels  in  his  comrades.  Obedience,  as  IMr.  Bagehot  has  well 
shewTi,^  is  of  the  highest  value,  for  any  form  of  government  is 
better  than  none.  Selfish  and  contentious  people  will  not  cohere, 
and  without  coherence  nothing  can  be  effected.  A  ti*ibe  rich  in 
the  above  qualities  would  spread  and  be  victorious  over  other 
tribes  :  but  in  the  course  of  time  it  would,  judging  from  all  past 
history,  be  in  its  turn  overcome  by  some  other  tribe  still  more 
highly  endowed.  Thus  the  social  and  moral  qualities  would 
tend  slowly  to  advance  and  be  diffused  throughout  the  world. 

But  it  may  be  asked,  how  within  the  limits  of  the  same  tribe 
did  a  large  number  of  members  first  become  endowed  with  these 
social  and  moral  qualities,  and  how  was  the  standard  of  ex- 
cellence raised  ?  It  is  extremely  doubtful  whether  the  offspring 
of  the  more  sympathetic  and  benevolent  parents,  or  of  those 
who  were  the  most  faithful  to  their  comrades,  would  be  reared 
in  greater  numbers  than  the  children  of  selfish  and  treacherous 
parents  belonging  to  the  same  tribe.  He  who  was  ready  to 
sacrifice  his  life,  as  many  a  savage  has  been,  rather  than  betray 
his  comrades,  would  often  leave  no  offspring  to  inherit  his  noble 
nature.  The  bravest  men,  who  were  always  willing  to  come  to 
the  front  in  war,  and  who  freely  risked  their  lives  for  others, 
would  on  an  average  perish  in  larger  numbers  than  other  men. 
Therefore  it  hardly  seems  probable,  that  the  number  of  men 
gifted  with  such  virtues,  or  that  the  standard  of  their  excellence, 
could  be  increased  through  natural  selection,  that  is,  by  the 
survival  of  the  fittest ;  for  we  are  not  here  speaking  of  one  tribe 
being  victorious  over  another. 

Although  the  circumstances,  leading  to  an  increase  in  the 
number  of  those  thus  endowed  within  the  same  tribe,  are  too 
complex  to  be  clearly  followed  out,  we  can  trace  some  of  the 
probable  steps.    In  the  first  place,  as  the  reasoning  powers  and 

*  See  a  remarkable  series  of  arti-      April  1,  1868  ;  July  1,  1869,  since 
cles  on  '  Physics  and  Politics '  in  the      separately  published. 
'  Fortnightly  Review,'    Nov.    1867  ; 


Chap.  V.  Moral  Facilities.  1 3 1 

foresight  of  the  members  became  improved,  each  man  would' 
soon  learn  that  if  ho  aided  his  fellow-men,  he  would  commonly 
receive  aid  in  return.  From  this  low  motive  he  might  acquire 
the  habit  of  aiding  his  fellows ;  and  the  habit  of  performing 
benevolent  actions  certainly  strengthens  the  feeling  of  sympathy 
which  gives  the  first  impulse  to  benevolent  actions.  Habits, 
moreover,  followed  during  many  generations  probably  tend  to 
be  inherited. 

But  another  and  much  more  powerful  stimulus  to  the  de- 
velopment of  the  social  virtues,  is  afforded  by  the  praise  and  the 
blame  of  our  fellow-men.  To  the  instinct  of  sympathy,  as  we  have 
already  seen,  it  is  primarily  due,  that  we  habitually  bestow  both 
praise  and  blame  on  others,  whilst  we  love  the  former  and  dread 
the  latter  when  applied  to  ourselves ;  and  this  instinct  no  doubt 
was  originally  acquired,  like  all  the  other  social  instincts,  through 
natural  selection.  At  how  early  a  period  the  progenitors  of  man 
in  the  course  of  their  development,  became  capable  of  feehngand 
being  impelled  by,  the  praise  or  blame  of  their  fellow-creatures, 
we  caijnot  of  course  say.  But  it  appears  that  even  dogs  appre- 
ciate encouragement,  praise,  and  blame.  The  rudest  savages 
feel  the  sentiment  of  glory,  as  they  clearly  show  by  preserving 
the  trophies  of  their  prowess,  by  their  habit  of  excessive  boasting, 
and  even  by  the  extreme  care  which  they  take  of  their  per- 
sonal appearance  and  decorations ;  for  unless  they  regarded  the 
opinion  of  their  comrades,  such  habits  would  be  senseless. 

They  certainly  feel  shame  at  the  breach  of  some  of  their  lesser 
rules,  and  apparently  remorse,  as  shewn  by  the  case  of  the 
Australian  who  grew  thin  and  could  not  rest  from  having 
delayed  to  murder  some  other  woman,  so  as  to  propitiate  his  dead 
wife's  spirit.  Though  I  have  not  met  with  any  other  recorded 
case,  it  is  scarcely  credible  that  a  savage,  who  will  sacrifice  his 
life  rather  than  betray  his  tribe,  or  one  who  will  deliver  himself 
up  as  a  prisoner  rather  than  break  his  parole,*^  would  not 
feel  remorse  in  his  inmost  soul,  if  he  had  failed  in  a  duty, 
which  he  held  sacred.         » 

We  may  therefore  conclude  that  primeval  man,  at  a  very 
remote  period,  was  influenced  by  the  praise  and  blame  of  his 
fellows.  It  is  obvious,  that  the  members  of  the  same  tribe  would 
approve  of  conduct  which  appeared  to  them  to  be  for  the  general 
good,  and  would  reprobate  that  which  appeared  evil.  To  do 
good  unto  others — to  do  unto  others  as  ye  would  they  should  do 
unto  you— is  the  foundation-stone  of  morality.  It  is,  therefore, 
hardly  possible  to  exaggerate  the  importance  during  rude  times 

^  Mr.  Wallace  gives  cases  in  of  Natural  Seleotioa,'  1870,  p. 
bla    'Contributions   to    the   Theory      354. 


132  The  Descent  of  Man.  Part  1. 

of  the  loYe  of  praise  and  the  dread  of  blame.  A  man  who  was 
not  impelled  by  any  deep,  instinctive  feeling,  to  sacrifice  his 
life  for  the  good  of  others,  yet  was  roused  to  such  actions  by  a 
sense  of  glory,  would  by  his  example  excite  the  same  wish  for 
glory  in  other  men,  and  would  strengthen  by  exercise  the  noble 
feeling  of  admiration.  He  might  thus  do  far  more  good  to  his 
tribe  than  by  begetting  offspring  with  a  tendency  to  inherit  his 
own  high  character. 

With  increased  experience  and  reason,  man  perceives  the 
more  remote  consequences  of  his  actions,  and  the  self-regarding 
virtues,  such  as  temperance,  chastity,  &c.,  which  during  early 
times  are,  as  we  have  before  seen,  utterly  disregarded,  come  to 
be  highly  esteemed  or  even  held  sacred.  I  need  not,  however, 
repeat  what  I  have  said  on  this  head  in  the  fourth  chapter. 
Ultimately  our  moral  sense  or  conscience  becomes  a  highly 
complex  sentiment— originating  in  the  social  instincts,  largely 
guided  by  the  approbation  of  our  fellow-men,  ruled  by  reason, 
self-interest,  and  in  later  times  by  deep  religious  feelings,  and 
confirmed  by  instruction  and  habit.       '  * 

It  must  not  be  forgotten  that  although  a  high  standard  of 
morality  gives  but  a  slight  or  no  advantage  to  each  individual 
man  and  his  children  over  the  other  men  of  the  same  tribe,  yet 
that  an  increase  in  the  number  of  well-endowed  men  and  an 
advancement  iii  the  standard  of  morality  will  certainly  give  an 
immense  advantage  to  one  tribe  over  another.  A  tribe  including 
many  members  who,  from  possessing  in  a  high  degTee  the  spmt 
of  patriotism,  fidelity,  obedience,  courage,  and  sympathy,  were 
always  ready  to  aid  one  another,  and  to  sacrifice  themselves  for 
the  common  good,  would  be  victorious  over  most  other  tribes ; 
and  this  would  be  natural  selection.  At  all  times  throughout 
the  world  tribes  have  supplanted  other  tribes ;  and  as  morahty 
is  one  important  element  in  their  success,  the  standard  of 
morality  and  the  number  of  well- en  do  wed  men  will  thus  every- 
where tend  to  rise  and  increase. 

It  is,  however,  very  difficult  to  form  any  judgment  why  one 
particular  tribe  and  not  another  has  been  successful  and  has 
risen  in  the  scale  of  civilisation.  Many  savages  are  in  the  same 
condition  as  when  first  discovered  several  centuries  ago.  As  Mr. 
Bagehot  has  remarked,  we  are  apt  to  look  at  progress  as  normal 
in  human  society ;  but  history  refutes  this.  The  ancients  did 
not  even  entertain  the  idea,  nor  do  the  Oriental  nations  at  the 
present  day.  According  to  another  high  authority.  Sir  Henry 
Maine,^  "  the  greatest  part  of  mankind  has  never  shewn   a 

'  'Ancient  Law,'  1861,  p.  22.  nightly  Review,'  April  1,  1S68,  p. 
For  Mr.  Bagehot's  remarks,  '  Fort-      452. 


Chap.  V.  Civilised  Nations. 


00 


"  particlo  of  desire  that  its  civil  institutions  should  be  iui- 
"  proved."  Progress  seems  to  depend  on  many  concurrent 
favourable  conditions,  far  too  complex  to  be  followed  out.  But 
it  has  often  been  remarked,  that  a  cool  climate,  from  leading  to 
industry  and  to  the  various  arts,  has  been  highly  favourable 
thereto.  The  Esquimaux,  pressed  by  hard  necessity,  have 
succeeded  in  many  ingenious  inventions,  but  their  climate  has 
been  too  severe  for  continued  progress.  Nomadic  habits,  whether 
over  wide  plains,  or  through  the  dense  forests  of  the  tropics,  or 
along  the  shores  of  the  sea,  have  in  every  case  been  highly 
detrimental.  "Whilst  observing  the  barbarous  inhabitants  of 
Tierra  del  Fuego,  it  struck  me  .that  the  possession  of  some 
property,  a  fixed  abode,  and  the  union  of  many  families  under  a 
chief,  were  the  indispensable  requisites  for  civilisation.  Such 
habits  almost  necessitate  the  cultivation  of  the  ground ;  and  the 
first  steps  in  cultivation  would  j^robably  result,  as  I  have  else- 
where shewu,^  from  some  such  accident  as  the  seeds  of  a  fruit- 
tree  falling  on  a  heap  of  refuse,  and  producing  an  unusually  fine 
variety.  The  problem,  however,  of  the  first  advance  of  savages 
towards  civilisation  is  at  present  much  too  difficult  to  be  solved. 

Natural  Selection  as  affecting  Civilised  Nations. — I  have  hitherto 
only  considered  the  advancement  of  man  from  a  semi-human 
condition  to  that  of  the  modern  savage.  But  some  remarks  on 
the  action  of  natural  selection  on  civilised  nations  may  be  worth 
adding.  This  subject  has  been  ably  discussed  by  Mr.  W.  E. 
Greg,^  and  previously  by  Mr.  Wallace  and  Mr.  Galton.^**  Most 
of  my  remarks  are  taken  from  these  three  authors.  With 
savages,  the  weak  in  body  or  mind  are  soon  eliminated ;  and 
those  that  survive  commonly  exhibit  a  vigorous  state  of  health. 
We  civilised  men,  on  the  other  hand,  do  our  utmost  to  check  the 
process  of  elimination ;  we  build  asylums  for  the  imbecile,  the 
maimed,  and  the  sick ;  we  institute  poor-laws ;  and  our  medical 

"  'The  Variation  of  Animals  and  1869,  and  by  Mr.  E.  Ray  Lankester 

Plants  under  Domestication,' vol.  i.  in     his     'Comparative    Longevitv,' 

p.  309.  1870,    p.    128.     Similar   views   ap- 

*  'Eraser's  Magazine,' Sept.  1868,  peared   previously  in   the   'Austra- 

p.  353.     This  article  seems  to  have  lasian,'    July*   13,    1867.      I    have 

struck  many  persons,  and  has  given  borrowed  ideas  from  several  of  these 

rise  to  two  remarkable  essays  and  a  writers. 

rejoinder    in    the  '  Spectator,'    Oct.  ^'^  For  Mr.  Wallace,  see  '  Anthro- 

3rd    and   17th,  1868.     It    has    also  polog.  Review,'  as  before  cited.     Mr. 

been  discussed  in  the  'Q.  Journal  of  Galton    in  '  Macmillan's  Magazine,' 

Science,'  1869,  p.  152,  and  by  Mr.  Aug.  1865,  p.  318  ;   also  his  great 

Lawson    Tait    in    the    'Dublin    Q.  work,  '  Hereditary  Genius,'  1870. 
Journal   of  Medical   Science,'    Feb. 


1 34  The  Descent  of  Man.  Part  I. 


men  exert  their  utmost  skill  to  save  the  Kfe  of  every  one  to  the 
last  moment.  There  is  reason  to  belieye  that  vaccination  has 
preserved  thousands,  who  from  a  weak  constitution  would 
formerly  have  succumbed  to  small-pox.  Thus  the  weak  members 
of  civilised  societies  propagate  their  kind.  No  one  who  has 
attended  to  the  breeding  of  domestic  animals  will  doubt  that 
this  must  be  highly  injurious  to  the  race  of  man.  It  is  surprising 
how  soon  a  want  of  care,  or  care  wrongly  directed,  leads  to  the 
degeneration  of  a  domestic  race ;  but  excepting  in  the  case  of 
man  himself,  hardly  any  one  is  so  ignorant  as  to  allow  his 
worst  animals  to  breed. 

The  aid  which  we  feel  impelled  to  give  to  the  helpless  is  mainly 
an  incidental  result  of  the  instinct  of  sympathy,  which  was 
originally  acquired  as  pai*t  of  the  social  instincts,  but  sub- 
sequently rendered,  in  the  manner  previously  indicated,  more 
tender  and  more  widely  diifused.  Nor  could  we  check  our 
sympathy,  even  at  the  urging  of  hard  reason,  without  deteriora- 
tion in  the  noblest  part  of  our  nature.  The  surgeon  may  harden 
himself  whilst  performing  an  operation,  for  he  knows  that  he  is 
acting  for  the  good  of  his  patient ;  but  if  we  were  intentionally 
to  neglect  the  weak  and  helpless,  it  could  only  be  for  a  con- 
tingent benefit,  with  an  overwhelming  present  evil.  We  must 
therefore  bear  the  undoubtedly  bad  eflfects  of  the  weak  surviving 
and  propagating  their  kind ;  but  there  appears  to  be  at  least  one 
check  in  steady  action,  namely  that  the  weaker  and  inferior 
members  of  society  do  not  marry  so  freely  as  the  sound ;  and 
this  check  might  be  indefinitely  increased  by  the  weak  in 
body  or  mind  refraining  from  marriage,  though  this  is  more  to  be 
hoped  for  than  expected. 

In  every  country  in  which  a  large  standing  army  is  kept  up, 
the  finest  young  men  are  taken  by  the  conscription  or  are 
enlisted.  They  are  thus  exposed  to  early  death  during  war,  are 
often  tempted  into  vice,  and  are  prevented  from  marrying  during 
the  prime  of  life.  On  the  other  hand  the  shorter  and  feebler  men, 
with  poor  constitutions,  are  left  at  home,  and  consequently  have 
a  much  better  chance  of  marrying  and  jDropagating  their  kind.^^ 

Man  accumulates  property  and  bequeaths  it  to  his  children, 
so  that  the  children  of  the  rich  have  an  advantage  over  the  poor 
in  the  race  for  success,  independently  of  bodily  or  mental  su- 
periority. On  the  other  hand,  the  children  of  parents  who  are 
short-lived,  and  are  therefore  on  an  average  deficient  in  health 
and  vigour,  come  into  their  property  sooner  than  other  children, 

"  Prof.   H,    Fick  (' Einfluss    der      on    this   head,   and    on   other   such 
Naturwissenschaft  auf  das    Eecht,'      points. 
June,  1872)  has  some  good  remarks 


Chap.  V.  Civilised  Nations.  1 3  5 

and  will  be  likely  to  marry  earlier,  and  leave  a  larger  number  of 
offspring  to  inherit  their  inferior  constitutions.  But  the  in- 
heritance of  property  by  itself  is  very  far  from  an  evil ;  for 
without  the  accumulation  of  capital  the  arts  could  not  progress ; 
and  it  is  chiefly  through  their  power  that  the  civilised  races  have 
extended,  and  arc  now  everywhere  extending  their  range,  so  as 
to  take  the  place  of  the  lower  races.  Nor  does  the  moderate 
accumulation  of  wealth  interfere  with  the  process  of  selection. 
When  a  poor  man  becomes  moderately  rich,  his  children  enter 
trades  or  professions  in  which  there  is  struggle  enough,  so  that 
the  able  in  body  and  mind  succeed  best.  The  presence  of  a  body 
of  well-instructed  men,  who  have  not  to  labour  for  their  daily 
bread,  is  important  to  a  degree  which  cannot  be  over-estimated ; 
as  all  high  intellectual  work  is  carried  on  by  them,  and  on  such 
work,  material  jirogress  of  all  kinds  mainly  depends,  not  to 
mention  other  and  higher  advantages.  No  doubt  wealth  when 
very  gi-eat  tends  to  convert  men  into  useless  drones,  but  their 
number  is  never  large;  and  some  degree  of  elimination  here 
occurs,  for  we  daily  see  rich  men,  who  happen  to  be  fools  or 
profligate,  squandering  away  their  wealth. 

Primogeniture  with  entailed  estates  is  a  more  direct  evil, 
though  it  may  formerly  have  been  a  great  advantage  by  the 
creation  of  a  dominant  class,  and  any  government  is  better 
than  none.  Most  eldest  sons,  though  they  may  be  weak  in  body 
or  mind,  marry,  whilst  the  younger  sons,  however  superior 
in  these  respects,  do  not  so  generally  marry.  Nor  can  worth- 
less eldest  sons  with  entailed  estates  squander  their  wealth. 
But  here,  as  elsewhere,  the  relations  of  civilised  life  are  so 
complex  that  some  compensatory  checks  intervene.  The  men 
who  are  rich  through  primogeniture  are  able  to  select  genera- 
tion after  generation  the  more  beautiful  and  charming  women ; 
and  these  must  generally  be  healthy  in  body  and  active  in 
mind.  The  evil  consequences,  such  as  they  may  be,  of  the 
continued  preservation  of  the  same  line  of  descent,  without  any 
selection,  are  checked  by  men  of  rank  always  wishing  to  increase 
their  wealth  and  power;  and  this  they  effect  by  marrying 
heiresses.  But  the  daughters  of  parents  who  have  produced 
single  children,  are  themselves,  as  Mr.  Galton^^  has  shewn,  apt  to 
be  sterile ;  and  thus  noble  families  are  continually  cut  off  in  the 
direct  line,  and  their  wealth  flows  into  some  side  channel ;  but 
unfortunately  this  channel  is  not  determined  by  superiority  of 
any  kind. 

Although  civilisation  thus  checks  in  many  ways  the  action  of 

>2  'Hereditary  Genius,'  1870,  pp.  132-140. 


136  The  Desce7it  of  Man.  Part  I. 

natural  selection,  it  apparently  favours  the  better  development 
of  the  body,  by  means  of  good  food  and  the  freedom  from  occa- 
sional hardships.  This  may  be  inferred  from  civilised  men 
having  been  found,  wherever  compared,  to  be  physically 
stronger  than  savages.'^  They  appear  also  to  have  equal  powers 
of  endurance,  as  has  been  proved  in  many  adventurous  ex- 
peditions. Even  the  great  luxury  of  the  rich  can  be  but  little 
detrimental ;  for  the  expectation  of  life  of  our  aristocracy,  at  all 
ages  and  of  both  sexes,  is  very  httle  inferior  to  that  of  healthy 
English  lives  in  the  lower  classes.^* 

We  will  now  look  to  the  intellectual  faculties.  If  in  each 
grade  of  society  the  members  were  divided  into  two  equal 
bodies,  the  one  including  the  intellectually  superior  and  the 
other  the  inferior,  there  can  be  little  doubt  that  the  former 
would  succeed  best  in  all  occupations,  and  rear  a  greater  number 
of  children.  Even  in  the  lowest  walks  of  life,  skill  and  ability 
must  be  of  some  advantage;  though  in  many  occupations, 
owing  to  the  great  division  of  labour,  a  very  small  one.  Hence 
in  civilised  nations  there  will  be  some  tendency  to  an  increase 
both  in  the  number  and  in  the  standard  of  the  intellectually 
able.  But  I  do  not  wish  to  assert  that  this  tendency  may  not  be 
more  than  counterbalanced  in  other  ways,  as  by  the  multiplica- 
tion of  the  reckless  and  improvident ;  but  even  to  such  as  these, 
ability  must  be  some  advantage. 

It  has  often  been  objected  to  views  like  the  foregoing,  that  the 
most  eminent  men  who  have  ever  lived  have  left  no  offspring  to 
inherit  their  great  intellect.  Mr.  Galton  says,^^  "  I  regret  I  am 
"  unable  to  solve  the  simple  question  whether,  and  how  far, 
''  men  and  women  who  are  prodigies  of  genius  are  infertile.  I 
"  have,  however,  shewn  that  men  of  eminence  are  by  no  means 
"  so."  Great  lawgivers,  the  founders  of  beneficent  rehgions, 
great  philosophers  and  discoverers  in  science,  aid  the  progress  of 
mankind  in  a  far  higher  degree  by  their  works  than  by  leaving 
a  numerous  progeny.  In  the  case  of  corporeal  structures,  it  is 
the  selection  of  the  slightly  better-endowed  and  the  elimination 
of  the  slightly  less  well-endowed  individuals,  and  not  the  pre- 
servation of  strongly-marked  and  rare  anomalies,  that  leads  to 
the  advancement  of  a  species.^''  So  it  will  be  with  the  intellectual 
faculties,  since  the  somewhat  abler  men  in  each  grade  of  society 

'^  Quatrefages,  'Revue  des  Cours  1870,  p.  115. 
Scientifiques,'  1867-68,  p.  659.  i^  <  Hereditary  Genius,'  1870,   p. 

^■^  See  the  fifth  and  sixth  columns,  330. 
compiled  from  good  authorities,  in  ^*^  '  Origin  of  Species  '  (fifth  edi- 

the  table  given   in  Mr.  E.  R,  Lan-  tion,  1869),  p.  lO-l. 
kester's    '  Comparative    Longevity,' 


Chap.  V.  •      Civilised  Nations.  ■  137 

succeed  rather  better  than  the  less  able,  and  consequently 
increase  in  number,  if  not  otherwise  prevented.  When  in 
any  nation  the  standard  of  intellect  and  the  number  of  intel- 
lectual men  have  increased,  we  may  expect  from  the  law  of 
the  deviation  from  an  average,  that  prodigies  of  genius  will,  as 
shewn  by  Mr.  Galton,  appear  somew^hat  more  frequently  than 
before. 

In  regard  to  the  moral  qualities,  some  elimination  of  the 
worst  dispositions  is  always  in  progress  even  in  the  most  civilised 
nations.  Malefactors  are  executed,  or  imprisoned  for  long 
periods,  so  that  they  cannot  freely  transmit  their  bad  qualities. 
Melancholic  and  insane  persons  are  confined,  or  commit  suicide. 
Violent  and  quarrelsome  men  often  come  to  a  bloody  end.  The 
restless  who  will  not  follow  any  steady  occupation— and  this 
relic  of  barbarism  is  a  great  check  to  civilisation  ^^  —  emigrate  to 
newly-settled  countries,  where  they  prove  useful  pioneers.  In- 
temperance is  so  highly  destructive,  that  the  expectation  of  life 
of  the  intemperate,  at  the  age  of  thirty  for  instance,  is  only  13-8 
years ;  whilst  for  the  rural  labourers  of  England  at  the  same  age 
it  is  40*59  years.^^  Profligate  women  bear  few  children,  and 
profligate  men  rarely  marry ;  both  suifer  from  disease.  In  the 
breeding  of  domestic  animals,  the  elimination  of  those  individuals, 
though  few  in  number,  which  are  in  any  marked  manner  inferior, 
is  by  no  means  an  unimportant  element  towards  success.  This 
especially  holds  good  with  injurious  characters  which  tend  to 
reappear  through  reversion,  such  as  blackness  in  sheep;  and 
with  mankind  some  of  the  worst  dispositions,  W'hich  occasionally 
without  any  assignable  cause  make  their  appearance  in  families, 
may  perhaps  be  reversions  to  a  savage  state,  from  which  we  are 
not  removed  by  very  many  generations.  This  view  seems 
indeed  recognised  in  the  common  expression  that  such  men  are 
the  black  sheep  of  the  family. 

With  civilised  nations,  as  far  as  an  advanced  standard  of 
morality,  and  an  increased  number  of  fairly  good  men  are  con- 
cerned, natural  selection  apparently  effects  but  little ;  though 
the  fundamental  social  instincts  w^ere  originally  thus  gained. 
But  I  have  already  said  enough,  whilst  treating  of  the  lower 
races,  on  the  causes  which  lead  to  the  advance  of  morality, 
namely,  the  approbation  of  our  fellows-men — the  strengthening 

^^  'Hereditary  Genius,'   1870,   p.  Xeison's  *  Vital   Statistics.'     In    re- 

347.  gard    to   profligacy,    see    Dr.    Farr, 

"  E.  Ray  Lankester,  '  Compara-  'Influence  of  Marriage  on  Mor- 
tive  Longevity,'  1870,  p.  115,  Tiie  tality,'  'Nat.  Assoc,  for  the  Promo- 
table    of  the    intemperate    is    from  tion  of  Social  Science,'  1858. 


138  The  Descent  of  Man.  Part  T. 

of  our  sympathies  by  habit — example  and  imitation — reason — 
experience,  and  even  self-interest — instruction  during  youth,  and 
rehgious  feelings. 

A  most  important  obstacle  in  civilised  conntries  to  an  increase 
in  the  number  of  men  of  a  superior  class  has  been  strongly  insisted 
on  by  Mr.  Greg  and  Mr.  Galton/^  namely,  the  fact  that  the  very 
poor  and  reckless,  who  are  often  degraded  by  vice,  almost  invari- 
ably marry  early,  whilst  the  careful  and  frugal,  who  are  generally 
otherwise  virtuous,  marry  late  in  life,  so  that  they  may  be  able 
to  support  themselves  and  their  children  in  comfort.  Those  who 
marry  early  produce  within  a  given  period  not  only  a  greater 
number  of  generations,  but,  as  shewn  by  Dr.  Duncan,^*^  they  pro- 
duce many  more  children.  The  children,  moreover,  that  are 
born  by  mothers  during  the  prime  of  life  are  heavier  and  larger, 
and  therefore  probably  more  vigorous,  than  those  born  at  other 
periods.  Thus  the  reckless,  degraded,  and  often  vicious  members 
of  society,  tend  to  increase  at  a  quicker  rate  than  the  provident 
and  generally  virtuous  members.  Or  as  Mr.  Greg  puts  the  case  : 
"  The  careless,  squalid,  unaspiring  Irishman  multiplies  like 
"  rabbits  :  the  frugal,  foreseeing,  self-respecting,  ambitious  Scot, 
"  stern  in  his  morality,  spiiitual  in  his  faith,  sagacious  and  dis- 
"  ciplined  in  his  intelligence,  passes  his  best  years  in  struggle 
"  and  in  celibacy,  marries  late,  and  leaves  few  behind  him. 
"  Given  a  land  originally  peopled  by  a  thousand  Saxons  and  a 
"  thousand  Celts — and  in  a  dozen  generations  five-sixths  of  the 
"  population  would  be  Celts,  but  five-sixths  of  the  property,  of 
"  the  power,  of  the  intellect,  would  belong  to  the  one-sixth  of 
''  Saxons  that  remained.  In  the  eternal '  struggle  for  existence,' 
"  it  would  be  the  inferior  and  le»8  favoured  race  that  had  pre- 
"  vailed— and  prevailed  by  virtue  not  of  its  good  qualities  but  of 
"  its  faults." 

There  are,  however,  some  checks  to  this  downward  tendency. 
We  have  seen  that  the  intemj)erate  suffer  from  a  high  rate  of 
mortality,  and  the  extremely  profligate  leave  few  offspring.  The 
poorest  classes  crowd  into  towns,  and  it  has  been  proved  by  Dr. 
Stark  from  the  statistics  of  ten  years  in  Scotland,^^  that  at  all 

'^     *  Fraser's      Magazine,'     Sept.  title    of   '  Fecundity,  Fertility,  and 

1868,  p.  353.     'Macmillan's  Maga-  Sterility,'    1871.       See,    also,    Mr. 

zine,  Aug.  1865,  p.  318.     The  Rev.  Galton,    'Hereditary    Genius,'    pp. 

F.  W.  Farrar  ('  Fraser's  Mag.'  Aug.  352-357,    for    observations    to    the 

1870,  p.  264)  takes  a  different  view.  above  effect. 

2*^  '  On  the  Laws  of  the  Fertility  2'     '  Tenth     Annual     Report     of 

of    Women,'    in    '  Transact.    Royal  Births,    Deaths,    &c.,   in   Scotland,' 

Soc'  Edinburgh,  vol.  xxiv.  p.  287  ;  1867,  p.  xxix. 
now  published  separately  under  the 


(jiiAP.  V.  Civilised  Nations.  1 39 

ages  the  death-rate  is  higher  in  towns  tlian  in  rural  districts, 
"  and  during  the  first  five  years  of  life  the  town  death-rate  is 
*'  almost  exactly  double  that  of  the  rural  districts."  As  these  re- 
tui-ns  include  both  the  rich  and  the  poor,  no  doubt  more  than 
twice  the  number  of  births  would  be  requisite  to  keep  up  the 
number  of  the  very  poor  inhabitants  in  the  towns,  relatively  to 
those  in  the  country.  With  women,  marriage  at  too  early  an 
age  is  highly  injurious ;  for  it  has  been  found  in  France  that, 
"  twice  as  many  wives  under  twenty  die  in  the  year,  as  died  out 
*'  of  the  same  number  of  the  unmarried."  The  mortality,  also, 
of  husbands  under  twenty  is  "  excessively  high,"  ^^  but  what  the 
cause  of  this  may  be,  seems  doubtful.  Lastly,  if  the  men  who 
prudently  delay  marrying  until  they  can  bring  up  their  families 
in  comfort,  were  to  select,  as  they  often  do,  women  in  the  prime 
of  life,  the  rate  of  increase  in  the  better  class  w^ould  be  only 
slightly  lessened. 

It  was  established  from  an  enormous  body  of  statistics,  taken 
during  1853,  that  the  unmarried  men  throughout  France, 
between  the  ages  of  twenty  and  eighty,  die  in  a  much  larger 
proportion  than  the  married :  for  instance,  out  of  every  1000 
unmarried  men,  between  the  ages  of  twenty  and  thirty,  11-3 
annually  died,  whilst  of  the  married  only  6'5  died.^^  A  similar 
law  was  proved  to  hold  good,  during  the  years  18G3  and  1864, 
with  the  entire  population  above  the  age  of  twenty  in  Scotland : 
for  instance,  out  of  every  1000  unmarried  men,  between  the  ages 
of  twenty  and  thirty,  14-97  annually  died,  whilst  of  the  married 
only  7-21:  died,  that  is  less  than  half.^*  Dr.  Stark  remarks  on 
this, "  Bachelorhood  is  more  destructive  to  life  than  the  most 
"unwholesome  trades,  or  than  residence  in  an  unwholesome 
"  house  or  district  where  there  has  never  been  the  most  distant 
"  attempt  at  sanitary  improvement."  He  considers  that  the 
lessened  mortality  is  the  direct  result  of  "  marriage,  and  the 
"  more  regular  domestic  habits  which  attend  that  state."  He 
admits,  however,  that  the  intemperate,  profligate,  and  criminal 
classes,  whose  duration  of  life  is  low,  do  not  commonly  marry ; 
and  it  must  likewise  be  admitted  that  men  with  a  weak  constitu- 

^    These    quotations    are    taken  from  the  same  striking  paper, 

from  our  highest  authority  on  such  -*  1  have  taken  the  mean  of  the 

questions,  namely,  Dr.  Farr,  in  his  quinquennial  means,  given  in  '  The 

paper   'On    the  Influence    of  Mar-  Tenth    Annual    Report    of    Births, 

riage  on  the  Mortality  of  the  French  Deaths,    &c.,    in    Scotland,'     18G7. 

People,'  read  before  the  Nat.  Assoc.  The    quotation   from    Dr.   Stark    is 

for  the  Promotion  of  Social  Science,  copied  from  an  article  in  the  '  Daily 

1858.  News,'  Oct.   17th,   18G8,  which  Dr. 

23  Dr.    Farr,    ibid.      The    quota-  Farr  considers  very  carefully  writ- 

tioas    given    below    are    extracted  ten. 


140  The  Descent  of  Man.  Part  I. 

tion,  ill  health,  or  any  great  infirmity  in  body  or  mind,  will  often 
not  wish  to  marry,  or  will  be  rejected.  L)r.  Stark  seems  to  have 
come  to  the  conclusion  that  marriage  in  itself  is  a  main  cause  of 
prolonged  life,  from  finding  that  aged  married  men  still  have  a 
considerable  advantage  in  this  respect  over  the  unmarried  of  the 
same  advanced  age  ;  but  every  one  must  have  known  instances 
of  men,  who  with  weak  health  during  youth  did  not  marry,  and 
yet  have  survived  to  old  age,  though  remaining  weak,  and  there-  ' 
fore  always  with  a  lessened  chance  of  life  or  of  marrying.  There 
is  another  remarkable  circumstance  which  seems  to  support  Dr. 
Stark's  conclusion,  namely,  that  widows  and  widowers  in  France 
suffer  in  comparison  with  the  married  a  very  heavy  rate  of  mor- 
tality ;  but  Dr.  Farr  attributes  this  to  the  poverty  and  evil  habits 
consequent  on  the  disruption  of  the  family,  and  to  grief.  On 
the  whole  we  may  conclude  with  Dr.  Farr  that  the  lesser  mortahty 
of  married  than  of  unmarried  men,  which  seems  to  be  a  general 
law,  "is  mainly  due  to  the  constant  elimination  of  imperfect 
"  types,  and  to  the  skilful  selection  of  the  finest  individuals  out 
"  of  each  successive  generation ;"  the  selection  relating  only  to 
the  marriage  state,  and  acting  on  all  corporeal,  intellectual,  and 
moral  qualities.^^  We  may,  therefore,  infer  that  sound  and 
good  men  who  out  of  prudence  remain  for  a  time  unmarried,  do 
not  suffer  a  high  rate  of  mortality. 

If  the  various  checks  specified  in  the  two  last  paragraphs,  and 
perhaps  others  as  yet  unknown,  do  not  prevent  the  reckless,  the 
vicious  and  otherwise  inferior  members  of  society  from  increas- 
ing at  a  quicker  rate  than  the  better  class  of  men,  the  nation  will 
retrograde,  as  has  too  often  occurred  in  the  history  of  the  world. 
We  must  remember  that  progress  is  no  invariable  rule.  It  is 
very  diflScult  to  say  why  one  civilised  nation  rises,  becomes  more 
l^owerful,  and  sjDreads  more  widely,  than  another ;  or  why  the 
same  nation  progresses  more  quickly  at  one  time  than  at  another. 
We  can  only  say  that  it  depends  on  an  increase  in  the  actual 
number  of  the  population,  on  the  number  of  the  men  endowed 
with  high  intellectual  and  moral  faculties,  as  well  as  on  their 
standard  of  excellence.  Corporeal  structure  appears  to  have 
little  influence,  except  so  far  as  vigour  of  body  leads  to  vigour  of 
mind. 

It  has  been  urged  by  several  writers  that  as  high  intellectual 
powers  are  advantageous  to  a  nation,  the  old  Greeks,  who  stood 
some  grades  higher  in  intellect  than  any  race  that  has  ever 

25  Dr.  Duncan  remarks  ('  Fecund-  '■'■  from  the  unmarried   side  to   the 

ity,  Fertility,'  &c.,  1871,  p.  334)  on  "  married,    leaving   the    unmarried 

this  subject;    "At   every   age    the  "columns  crowded  with  the  sickly 

"  healthy    and    beautiful    go    over  "  and  unfortunate." 


Chap  V.  Civilised  Nations.  1 41 


existed,"^  ought,  if  the  power  of  natural  selection  were  real,  to 
have  risen  still  higher  in  the  scale,  increased  in  number,  and 
stocked  the  whole  of  Europe.  Here  we  have  the  tacit  assump- 
tion, so  often  made  with  respect  to  corporeal  structures,  that 
there  is  some  innate  tendency  towards  continued  development  in 
mind  and  body.  But  development  of  all  kinds  depends  on  many 
concurrent  favourable  circumstances.  Natural  selection  acts 
only  tentatively.  Individuals  and  races  may  have  acquired  cer- 
tain indisputable  advantages,  and  yet  have  perished  from  failing 
in  other  characters.  The  Greeks  may  have  retrograded  from  a 
want  of  coherence  between  the  many  small  states,  from  the  small 
size  of  their  whole  country,  from  the  practice  of  slavery,  or  from 
extreme  sensuality ;  for  they  did  not  succumb  until  "  they  were 
"  enervated  and  corrupt  to  the  very  core."  '^  The  western  nations 
of  Europe,  who  now  so  immeasurably  surpass  their  former  savage 
progenitors,  and  stand  at  the  summit  of  civilisation,  owe  little 
or  none  of  their  superiority  to  direct  inheritance  from  the  old 
Greeks,  though  they  owe  much  to  the  written  works  of  that 
wonderful  people. 

Who  can  positively  say  why  the  Spanish  nation,  so  dominant 
at  one  time,  has  been  distanced  in  the  race.  The  awakening  of 
the  nations  of  Europe  from  the  dark  ages  is  a  still  more  perplex- 
ing problem.  At  that  early  period,  as  Mr.  Galton  has  remarked, 
almost  all  the  men  of  a  gentle  nature,  those  given  to  meditation 
or  culture  of  the  mind,  had  no  refuge  except  in  the  bosom  of 
a  Church  which  demanded  celibacy  ;^^  and  this  could  hardly 
fail  to  have  had  a  deteriorating  influence  on  each  successive 
generation.  During  this  same  period  the  Holy  Inquisition 
selected  with  extreme  care  the  freest  and  boldest  men  in  order 
to  burn  or  imprison  them.  In  Spain  alone  some  of  the  best 
men — those  who  doubted  and  questioned,  and  without  doubting 
there  can  be  no  progress — were  eliminated  during  three  cen- 
turies at  the  rate  of  a  thousand  a  year.  The  evil  which  the 
Catholic  Church  has  thus  effected  is  incalculable,  though  no 
doubt  counterbalanced  to  a  certain,  perhaps  to  a  large,  extent 
in  other  ways;  nevertheless,  Europe  has  progressed  at  an  un- 
paralleled rate. 

2®  See  the  ingenious  and  original  257)   advances   arguments    on    the 

argument    on   this    subject    by  Mr.  othi'r  side.    Sir  C.  Lyell  had  already 

Galton,     'Hereditary    Genius,'    i)p.  ('Principles    of    Geology,'    vol.     .1. 

340-342.  1868,  p.  489)  in  a  striking  passage 

^^  Mr.  Greg,  'Eraser's  Magazine,'  called  attention  to  the  evil  inHucnce 

Sept.  1868,  p.  357.  of  the  Holy  Inquisition  in   having, 

^®  'Hereditary  Genius,'  1870,  pp.  through  selection,  lowered  the  gene- 

357-359.     The   Rev.   F.  W.  Farrar  ral  standard  of  intelligence  in  Eu- 

('  Fraser's    Mag.,'    Aug.     1870,     p.  rope. 


142  The  Descent  of  Alan.  Pabt  I. 

The  remarkable  success  of  the  English  as  colonists,  compared 
to  other  European  nations,  has  been  ascribed  to  their  "  daring 
"  and  persistent  energy ; "  a  result  which  is  xvell  illustrated  by 
comparing  the  progress  of  the  Canadians  of  English  and  French 
extraction ;  but  who  can  say  how  the  EngUsh  gained  their  energy  ? 
There  is  apparently  much  truth  in  the  belief  that  the  wonderful 
progress  of  the  United  States,  as  well  as  the  character  of  the 
people,  are  the  results  of  natural  selection ;  for  the  more  ener- 
getic, restless,  and  courageous  men  from  all  parts  of  Europe 
have  emigrated  during  the  last  ten  or  twelve  generations  to  that 
great  country,  and  have  there  succeeded  best.-^  Looking  to  the 
distant  future,  I  do  not  think  that  the  Eev.  Mr.  Zincke  takes  an 
exaggerated  view  when  he  says  :^  ''  All  other  series  of  events — 
"  as  that  which  resulted  in  the  cultui'e  of  mind  in  Greece,  and 
"that  which  resulted  in  the  emj^ire  of  Piome— only  appear  to 
"  have  purpose  and  value  when  viewed  in  connection  with,  or 
"  rather  as  subsidiary  to  ...  .  the  gi'eat  stream  of  Anglo-Saxon 
"  emigration  to  the  west."  Obscure  as  is  the  problem  of  the 
advance  of  civilisation,  we  can  at  least  see  that  a  nation 
which  produced  during  a  lengthened  period  the  greatest 
number  of  highly  intellectual,  energetic,  brave,  patriotic,  and 
benevolent  men,  would  generally  prevail  over  less  favoured 
nations. 

Natural  selection  follows  from  the  struggle  for  existence  ;  and 
this  from  a  rajDid  rate  of  increase.  It  is  impossible  not  to  regret 
bitterly,  but  whether  wisely  is  another  question,  the  rate  at 
which  man  tends  to  increase ;  for  this  leads  in  barbarous  tribes 
to  infanticide  and  many  other  evils,  and  in  civilised  nations  to 
abject  poverty,  celibacy,  and  to  the  late  marriages  of  the  prudent. 
But  as  man  suffei-e  from  the  same  physical  evils  as  the  lower 
animals,  he  has  no  right  to  expect  an  immunity  from  the  evils 
consequent  on  the  struggle  for  existence.  Had  he  not  been  sub- 
jected during  primeval  times  to  natural  selection,  assuredly  he 
would  never  have  attained  to  his  present  rank.  Since  we  see  in 
many  parts  of  the  world  enormous  areas  of  the  most  fertile  land 
capable  of  supporting  numerous  happy  homes,  but  peopled  only 
by  a  few  wandering  savages,  it  might  be  argued  that  the  struggle 
for  existence  had  not  been  sufficiently  severe  to  force  man  up- 
wards to  his  highest  standard.  Judging  from  all  that  we  know 
of  man  and  the  lower  animals,  there  has  always  been  sufficient 
variability  in  their  intellectual  and  moral  faculties,  for  a  steady 
advance  through   natural  selection.     No  doubt  such  advance 

29  Mr.  Gallon,  '  Macmillau's  and  National  Life,'  Dec.  1869,  p.  184. 
Magazine,'   August,    1865,   p.    32  o.  ^o  ^Last    Winter   in  the     United 

See   also,  'Nature,'  'On  Darwinism      States,'  1868,  p.  29. 


Chap.  V.  Civilised  Nations.  143 


demands  many  favourable  concurrent  circumstances ;  but  it  may 
well  be  doubted  whether  the  most  favourable  would  have  sufficed, 
had  not  the  rate  of  increase  been  rapid,  and  the  consequent 
struggle  for  existence  extremely  severe.  It  even  appears  from 
what  we  see,  for  instance,  in  parts  of  S.  America,  that  a  people 
which  may  be  called  civilised,  such  as  the  Spanish  settlers,  is 
liable  to  become  indolent  and  to  retrograde,  when  the  con- 
ditions of  life  are  very  easy.  With  highly  civilised  nations  con- 
tinued progress  depends  in  a  subordinate  degree  on  natural 
selection-;  for  such  nations  do  not  supplant  and  exterminate  one 
another  as  do  savage  tribes.  Nevertheless  the  more  intelligent 
members  within  the  same  community  will  succeed  better  in  the 
long  run  than  the  inferior,  and  leave  a  more  numerous  progeny, 
and  this  is  a  form  of  natural  selection.  The  more  efficient 
causes  of  progress  seem  to  consist  of  a  good  education  during 
youth  whilst  the  brain  is  impressible,  and  of  a  high  standard  of 
excellence,  inculcated  by  the  ablest  and  best  men,  embodied  in 
the  laws,  customs  and  traditions  of  the  nation,  and  enforced  by 
public  opinion.  It  should,  however,  be  borne  in  mind,  that  the 
enforcement  of  public  opinion  depends  on  our  appreciation  of 
the  approbation  and  disapprobation  of  others ;  and  this  apprecia- 
tion is  founded  on  our  sympathy,  which  it  can  hardly  be  doubted 
was  originally  developed  through  natural  selection  as  one  of  the 
most  important  elements  of  the  social  instincts.^^ 

On  the  evidence  that  all  civUked  nations  ivere  once  harlarotis. — 
The  present  subject  has  been  treated  in  so  full  and  admirable  a 
manner  by  Sir  J.  Lubbock,"^  Mr.  Tylor,  Mr.  M'Lenuan,  and 
others,  that  I  need  here  give  only  the  briefest  summary  of  their 
results.  The  arguments  recently  advanced  by  the  Duke  of 
ArgylP^  and  formerly  by  Archbishop  Whately,  in  favour  of  the 
belief  that  man  came  into  the  world  as  a  civilised  being,  and 
that  all  savages  have  since  undergone  degradation,  seem  to  me 
weak  in  comparison  with  those  advanced  on  the  other  side. 
Many  nations,  no  doubt,  have  fallen  away  in  civilisation,  and 
some  may  have  lapsed  into  utter  barbarism,  though  on  this 
latter  head  I  have  met  with  no  evidence.  The  Fuegians  were 
probably  compelled  by  other  conquering  hordes  to  settle  in  their 
inhospitable  country,  and  they  may  have  become  in  consequence 
somewhat  more  degraded;   but  it  would  be  difficult  to  prove 

"  I    am    much   iudebted  to    Mr.  '^  'On  the  Origin  of  Civilisation," 

John  Morley   for    some   good   criti-  *  Proc.  Ethnological    Soc'  Nov.  26 

cisms   on   this    subject:     see,    also,  1867. 

Broca, 'Les  Selections,' 'Kevue  d'Au-  ^■^  '  Primeval  Man,'  18G9. 
thropologie,'  1872. 


144  ^^^^  Descent  of  Man.  Part  I. 

that  they  have  fallen  much  below  the  Botocudos,  who  inhabit 
the  finest  parts  of  Brazil. 

The  evidence  that  all  civilised  nations  are  the  descendants  of 
barbarians,  consists,  on  the  one  side,  of  clear  traces  of  their 
former  low  condition  in  still-existing  customs,  beliefs,  language, 
&c.;  and  on  the  other  side,  of  proofs  that  savages  are  inde- 
pendently able  to  raise  themselves  a  few  steps  in  the  scale  of 
civilisation,  and  have  actually  thus  risen.  The  evidence  on  the 
first  head  is  extremely  curious,  but  cannot  be  here  given  :  I  refer 
to  such  cases  as  that  of  the  art  of  enumeration,  which,  as  Mr. 
Tylor  clearly  shews  by  reference  to  the  words  still  used  in  some 
places,  originated  in  counting  the  fingers,  first  of  one  hand  and 
then  of  the  other,  and  lastly  of  the  toes.  We  have  traces  of  this 
in  our  own  decimal  system,  and  in  the  Eoman  numerals,  where, 
after  the  V.,  which  is  supposed  to  be  an  abbreviated  picture  of  a 
human  hand,  we  pass  on  to  VI.,  &c.,  when  the  other  hand  no 
doubt  was  used.  So  again,  "  when  we  speak  of  three-score  and 
"  ten,  we  are  counting  by  the  vigesimal  system,  each  score  thus 
"  ideally  made,  standing  for  20— for  '  one  man '  as  a  Mexican  or 
"  Carib  would  put  it."^^  According  to  a  large  and  increasing 
school  of  philologists,  every  language  bears  the  marks  of  its  slow 
and  gradual  evolution.  So  it  is  with  the  art  of  writing,  for 
letters  are  rudiments  of  pictorial  representations.  It  is  hardly 
possible  to  read  Mr.  M'Lennan's  work^^  and  not  admit  that 
almost  all  civilised  nations  still  retain  traces  of  such  rude  habits 
as  the  forcible  capture  of  wives.  What  ancient  nation,  as  the 
same  author  asks,  can  be  named  that  was  originally  mono- 
gamous ?  The  primitive  idea  of  justice,  as  shewn  by  the  law  of 
battle  and  other  customs  of  which  vestiges  still  remain,  was 
likewise  most  rude.  Many  existing  superstitions  are  the 
remnants  of  former  false  religious  beliefs.  The  highest  form  of 
religion — the  gi-and  idea  of  God  hating  sin  and  loving  right- 
eousness— was  unknown  during  primeval  times.       •  . 

Turning  to  the  other  kind  of  evidence :  Sir  J.  Lubbock  has 
shewn  that  some  savages  have  recently  improved  a  little  in 
some  of  their  simpler  arts.  From  the  extremely  curious 
account  which  he  gives  of  the  weapons,  tools,  and  arts,  in  use 

3*  '  Royal  Institution  of  Great  '  A  Conjectural  Solution  of  the 
Britain,'  March  15,  1867.  Also,  Origin  of  the  Class,  System  of 
'  Researches  into  the  Early  History  Relationship,'  in  '  Proc.  American 
of  Mankind,'  1865.  Acad,    of    Sciences,'   vol.    vii.    Feb. 

35  'Primitive  Marriage,'  1865.  1868.  Prof.  Schaaffhausen  ('An- 
See,  likewise,  an  excellent  article,  thropolog.  Review,'  Oct.  1869,  p. 
evidently  by  the  same  author,  in  373)  remarks  on  "  the  vestiges  of 
the  'North  British  Review,'  July,  "human  sacrifices  found  both  in 
1869.      Also,    Mr,    L.    H.    Morgjin,      "  Homer  and  the  Old  Testament." 


CiiAr.  V.  Nations.  1 45 

amongst  savages  in  yarious  parts  of  the  world,  it  cannot  be 
doubted  that  these  have  nearly  all  been  independent  discoveries, 
excepting  perhaps  the  art  of  making  fire.^'^  The  Australian 
boomerang  is  a  good  instance  of  one  such  independent  discovery. 
The  Tahitians  when  first  visited  had  advanced  in  many  respects 
beyond  the  inhabitants  of  most  of  the  other  Polynesian  islands. 
There  are  no  just  grounds  for  the  belief  that  the  high  culture  of 
the  native  Peruvians  and  Mexicans  was  derived  from  abroad  ;  ^^ 
many  native  plants  were  there  cultivated,  and  a  few  native 
animals  domesticated.  We  should  bear  in  mind  that,  judging 
from  the  small  influence  of  most  missionaries,  a  wandering  crew 
from  some  semi-civilised  land,  if  washed  to  the  shores  of 
America,  would  not  have  produced  any  marked  eifect  on  the 
natives,  unless  they  had  already  become  somewhat  advanced. 
Looking  to  a  very  remote  period  in  the  history  of  the  world,  we 
find,  to  use  Sir  J.  Lubbock's  well-known  terms,  a  paleolithic  and 
neolithic  period;  and  no  one  will  pretend  that  the  art  of 
grinding  rough  flint  tools  was  a  borrowed  one.  In  all  parts  of 
EurojDe,  as  far  east  as  Greece,  in  Palestine,  India,  Japan,  New 
Zealand,  and  Africa,  including  Egypt,  flint  tools  have  been 
discovered  in  abundance;  and  of  their  use  the  existing  in- 
habitants retain  no  tradition.  There  is  also  indirect  evidence  of 
their  former  use  by  the  Chinese  and  ancient  Jews.  Hence  there 
can  hardly  be  a  doubt  that  the  inhabitants  of  these  countries, 
which  include  nearly  the  whole  civilised  world,  were  once 
in  a  barbarous  condition.  To  believe  that  man  was  abori- 
ginally civilised  and  then  suffered  utter  degradation  in  so  many 
regions,  is  to  take  a  pitiably  low  view  of  human  nature.  It  is 
apparently  a  truer  and  more  cheerful  view  that  progress  has 
been  much  more  general  than  retrogression ;  that  man  has  risen, 
though  by  slow  and  interrupted  steps,  from  a  lowly  condition  to 
the  highest  standard  as  yet  attained  by  him  in  knowledge, 
morals  and  reUgion. 

3«  Sir    J.    Lubbock,    'Prehistoric  edit.,  1870. 

Times,'   2ad    edit.   1869,    chap.   xv.  ^r  j^j..  F.  Miiller  has  made  some 

and   xvi.   et  passim.     See    also    the  good  remarks   to  this   effect  in  the 

excellent    9th    chapter    in    Trior's  '  Reise    der    Novara :    Anthropolog. 

Early    History   of   Mankind,''  2nd  Theil,'  Abtheil.  iii.  18G8,  s.  127. 


146  The  Descent  of  Maiu  Pakt  I. 


CHAPTER  YI. 
On  the  Affinities  and  Genealogy  cf  Man. 

Position  of  man  in  the  animal  series — The  natui'al  system  genealogical — ■ 
Adaptive  characters  of  slight  value — Various  small  points  of  resem- 
blance between  man  and  the  Quadrumana — Rank  of  man  in  the  natural 
system — Birthplace  and  antiquity  of  man — Absence  of  fossil  connecting- 
links — Lower  stages  in  the  genealogy  of  man,  as  inferred,  firstly  from 
his  affinities  and  secondly  from  his  structure — Early  androgynous  con- 
dition of  the  Vertebrata — Conclusion. 

Even  if  it  be  granted  that  the  diflference  between  man  and  his 
nearest  allies  is  as  great  in  cori)oreal  structure  as  some  natu- 
ralists maintain,  and  although  we  must  grant  that  the  differ- 
ence between  them  is  immense  in  mental  power,  yet  the  facts 
given  in  the  earlier  chapters  appear  to  declare,  in  the  plainest 
manner,  that  man  is  descended  from  some  lower  form,  notwith- 
standing that  connecting-links  have  not  hitherto  been  dis- 
covered. 

Man  is  liable  to  numerous,  slight,  and  diversified  variations, 
which  are  induced  by  the  same  general  causes,  are  governed 
and  transmitted  in  accordance  with  the  same  general  laws,  as  in 
the  lower  animals.  Man  has  multiplied  so  rapidly,  that  ho  has 
necessarily  been  exposed  to  struggle  for.  existence,  and  con- 
sequently to  natural  selection.  He  has  given  rise  to  many  races^ 
some  of  which  differ  so  much  from  each  other,  that  they  have 
often  been  ranked  by  naturalists  as  distinct  species.  His  body 
is  constructed  on  the  same  homological  jDlan  as  that  of  other 
mammals.  He  passes  thi'ough  the  same  phases  of  embryo- 
logical  development.  He  retains  many  rudimentary  and  useless 
structures,  which  no  doubt  were  once  serviceable.  Characters 
occasionally  make  their  re-appearance  in  him,  which  we  have 
reason  to  believe  were  possessed  by  his  early  progenitors.  If  the 
origin  of  man  had  been  wholly  different  from  that  of  all  other 
animals,  these  various  appearances  would  be  mere  empty 
deceptions ;  but  such  an  admission  is  incredible.  These  appear- 
ances, on  the  other  hand,  are  intelligible,  at  least  to  a  large 
extent,  if  man  is  the  co-descendant  with  other  mammals  of  some 
unkno\\Ti  and  lower  form. 

Some  natm-alists,  from  being  deeply  impressed  with  the 
mental  and  spiritual  powers  of  man,  have  divided  the  whole 
organic  world  into  three  kingdoms,  the  Human,  the  Animal, 


CiiAi'.  VI.  Affinities  and  Genealogy.  147 

and  the  Vegetable,  thus  giving  to  man  a  separate  kingdom.' 
Spiritual  powers  cannot  be  compared  or  classed  by  the  natu- 
ralist: but  he  may  endeavour  to  shew,  as  I  have  done,  that  the 
mental  faculties  of  man  and  the  lower  animals  do  not  differ  in 
kind,  although  immensely  in  degree.  A  difference  in  degree, 
however  great,  does  not  justify  us  in  placing  man  in  a  distinct 
kingdom,  as  will  perhaps  be  best  illustrated  by  comparing  the 
mental  j^owers  of  two  insects,  namely,  a  coccus  or  scale-insect 
and  an  ant,  which  undoubtedly  belong  to  the  same  class.  The 
difference  is  here  greater  than,  though  of  a  somewhat  different  kind 
from,  that  between  man  and  the  highest  mammal.  The  female 
coccus,  whilst  young,  attaches  itself  by  its  proboscis  to  a  plant ; 
sucks  the  sap,  but  never  moves  again ;  is  fertilised  and  lays  eggs ; 
and  this  is  its  whole  history.  On  the  other  hand,  to  describe  the 
habits  and  mental  powers  of  worker-ants,  would  require,  as 
Pierre  Huber  has  shewn,  a  large  volume ;  I  may,  however,  briefly 
specify  a  few  points.  Ants  certainly  communicate  information  to 
each  other,  and  several  unite  for  the  same  work,  or  for  games  of 
play.  They  recognise  their  fellow-ants  after  months  of  absence, 
and  feel  sympathy  for  each  other.  They  build  gi'eat  edifices, 
keep  them  clean,  close  the  doors  in  the  evening,  and  post 
sentries.  They  make  roads  as  well  as  tunnels  under  rivers,  and 
temporary  bridges  over  them,  by  clinging  together.  They 
collect  food  for  the  community,  and  when  an  object,  too  large  for 
entrance,  is  brought  to  the  nest,  they  enlarge  the  door,  and 
afterwards  build  it  up  again.  They  store  up  seeds,  of  which 
they  prevent  the  germination,  and  which,  if  damp,  are  brought 
up  to  the  surface  to  dry.  They  keep  aphides  and  other  insects  as 
milch-cows.  They  go  out  to  battle  in  regular  bands,  and  freely 
sacrifice  their  lives  for  the  common  weal.  They  emigrate  ac- 
cording to  a  preconcerted  plan.  They  capture  slaves.  They  move 
the  eggs  of  their  aphides,  as  well  as  their  own  eggs  and  cocoons, 
into  warm  parts  of  the  nest,  in  order  that  they  may  be  quickly 
hatched ;  and  endless  similar  facts  could  be  given.^  On  the 
whole,  the  difference  in  mental  power  between  an  ant  and  a 
coccus  is  immense ;  yet  no  one  has  ever  dreamed  of  placing  these 
insects  in  distinct  classes,  much  less  in  distinct  kingdoms.     IS'o 

*  Isidore  GeofFroy  St.-Hilaire  gives  of  auts   are  given  by  Mr.  Belt,  in 

a  detailed  account   of   the   position  in    his    'Naturalist    in    Kicaragua,' 

assigned  to  man  by  various  natural-  1874.      See    also    Mr.    Sloggridge's 

ists  in   their  classifications:   'Hist,  admirable  work,  '  Harvesting  Ants,' 

Nat.  Gen,'   torn.  ii.  1859,  pp.  170-  &e.,  1873,  also  '  L'Instinct  chez  les 

189.  Insectes,'    by    M.    George    Pouchet, 

-  Some   of  the  most    interesting  'Revue    des    Deux    Moudes,'    Feb. 

facts  ever  published  on  the  habits  1870  p.  682. 


148  The  Descent  of  Man.  Part  T. 

doubt  the  difference  is  bridged  over  by  other  insects ;  and  this 
is  not  the  case  with  man  and  the  higher  apes.  But  we  have 
every  reason  to  believe  that  the  breaks  in  the  series  are  simply 
the  results  of  many  foi-ms  having  become  extinct. 

Professor  Owen,  relying  chiefly  on  the  structure  of  the  brain, 
has  divided  the  mammalian  series  into  four  sub-classes.  One  of 
these  he  devotes  to  man;  in  another  he  places  both  the 
Marsupials  and  the  Monotremata;  so  that  he  makes  man  as 
distinct  from  all  other  mammals  as  are  these  two  latter  groups 
conjoined.  This  view  has  not  been  accepted,  as  far  as  I  am 
aware,  by  any  naturalist  capable  of  forming  an  independent 
judgment;  and  therefore  need  not  here  be  further  considered. 

We  can  understand  why  a  classification  founded  on  any  single 
character  or  organ— even  an  organ  so  wonderfully  complex  and 
important  as  the  brain — or  on  the  high  development  of  the 
mental  faculties,  is  almost  sure  to  prove  unsatisfactory.  This 
principle  has  indeed  been  tried  with  hymenopterous  insects; 
but  when  thus  classed  by  their  habits  or  instincts,  the  arrange- 
ment proved  thoroughly  artificial.^  Classifications  may,  of 
course,  be  based  on  any  character  whatever,  as  on  size,  colour, 
or  the  element  inhabited;  but  naturalists  have  long  felt  a 
profound  conviction  that  there  is  a  natural  system.  This 
system,  it  is  now  generally  admitted,  must  be,  as  far  as  possible, 
genealogical  in  arrangement, — that  is  the  co-descendants  of  the 
same  form  must  be  kept  together  in  one  group,  apart  from  the 
co-descendants  of  any  other  form ;  but  if  the  parent-forms  are 
related,  so  will  be  their  descendants,  and  the  two  groups  to- 
gether will  form  a  larger  group.  The  amount  of  difference 
between  the  several  groups — that  is  the  amount  of  modification 
which  each  has  undergone— is  expressed  by  such  terms  as 
genera,  families,  orders,  and  classes.  As  we  have  no  record  of 
the  lines  of  descent,  the  pedigree  can  be  discovered  only  by 
observing  the  degrees  of  resemblance  between  the  beings  which 
are  to  be  classed.  For  this  object  numerous  points  of  resem- 
blance are  of  much  more  importance  than  the  amount  of 
similarity  or  dissimilarity  in  a  few  points.  If  two  languages 
were  found  to  resemble  each  other  in  a  multitude  of  words  and 
points  of  construction,  they  would  be  universally  recognised  as 
having  sprung  from  a  common  source,  notwithstanding  that 
they  differed  greatly  in  some  few  words  or  points  of  construction. 
But  with  organic  beings  the  points  of  resemblance  must  not 
consist  of  adaptations  to  similar  habits  of  fife  :  two  animals  may, 
for  instance,  have  had  their  whole  frames  modified  for  living  in 

'  Westwood,  '  Modern  Class  of  Insects/  vol.  ii.  1840,  p.  87. 


Chap.  XL  Affinities  and  Genealogy.  149 

the  ■water,  and  yet  they  will  not  be  bronglit  any  nearer  to  each 
other  in  the  natural  system.  Hence  "sve  can  see  how  it  is  that 
resemblances  in  several  unimportant  structures,  in  useless  and 
rudimentary  organs,  or  not  now  functionally  active,  or  in  an 
embryological  condition,  are  by  far  the  most  serviceable  for  clas- 
sification ;  for  they  can  hardly  be  due  to  adaptations  within  a 
late  period ;  and  thus  they  reveal  the  old  lines  of  descent  or  of 
true  affinity. 

We  can  further  see  why  a  gi-eat  amount  of  modification  in 
some  one  character  ought  not  to  lead  us  to  separate  widely  any 
two  organisms.  A  part  which  already  differs  much  from  the 
same  part  in  other  allied  forms  has  already,  according  to  the 
theory  of  evolution,  varied  much ;  consequently  it  would  (as  long 
as  the  organism  remained  exposed  to  the  same  exciting  con- 
ditions) be  liable  to  further  variations  of  the  same  kind ;  and 
these,  if  beneficial,  would  be  preserved,  and  thus  be  continually 
augmented.  In  many  cases  the  continued  development  of  a  part, 
for  instance,  of  the  beak  of  a  bird,  or  of  the  teetli  of  a  mammal, 
would  not  aid  the  species  in  gaining  its  food,  or  for  any  other 
object ;  but  with  man  we  can  see  no  definite  limit  to  the  con- 
tinued development  of  the  brain  and  mental  faculties,  as  far  as 
advantage  is  concerned.  Therefore  in  determining  the  position 
of  man  in  the  natural  or  genealogical  system,  the  extreme  de- 
velopment  of  his  brain  ought  not  to  outweigh  a  multitude  of 
resemblances  in  other  less  important  or  quite  unimportant 
points. 

The  greater  number  of  naturalists  who  have  taken  into  con- 
sideration the  whole  structure  of  man,  including  his  mental 
faculties,  have  followed  Blumenbach  and  Cuvier,  and  have  placed 
man  in  a  separate  Order,  under  the  title  of  the  Bimana,  and 
therefore  on  an  equality  with  the  orders  of  the  Quadrumana, 
Carnivora,  &c.  Recently  many  of  our  best  naturalists  have 
recurred  to  the  view  first  propounded  by  Linnscus,  so  remarkable 
for  his  sagacity,  and  have  placed  man  in  the  same  Order  with 
the  Quadrumana,  under  the  title  of  the  Primates.  The  justice  of 
this  conclusion  will  be  admitted :  for  in  the  first  place,  we  must 
bear  in  mind  the  comparative  insignificance  for  classification 
of  the  great  development  of  the  brain  in  man,  and  that  the 
strongly-marked  differences  between  the  skulls  of  man  and  tlK^ 
Quadrumana  (lately  insisted  upon  by  Bischoff,  Aeby,  and  others) 
apparently  follow  from  their  differently  developed  brains.  In 
the  second  place,  we  must  remember  that  nearly  all  the  other 
and  more  important  difi'crences  Ixitween  man  and  the  Quadrumana 
are  manifestly  adaptive  in  their  nature,  and  relate  cliiefly  to  tljo 
erect  position  of  man ;  such  as  the  structure  of  his  hand,  /oot, 


1 50  The  Descent  of  Man.  Part  I. 

and  pelvis,  the  curvature  of  his  spine,  and  the  position  of  his 
head.  The  family  of  Seals  offers  a  -good  illustration  of  the  small 
importance  of  adaptive  characters  for  classification.  These 
animals  differ  from  all  other  Carnivora  in  the  form  of  their 
bodies  and  in  the  structure  of  their  limbs,  far  more  ihan  does 
man  from  the  higher  apes ;  yet  in  most  systems,  from  ihat  of 
Cuvier  to  the  most  recent  one  by  Mr.  Flower/  seals  are  ranked 
as  a  mere  family  in  the  Order  of  the  Carnivora.  If  man  had  not 
been  his  own  classifier,  he  would  never  have  thought  of  founding 
a  separate  order  for  his  own  reception.  < 

It  would  be  beyond  my  limits,  and  quite  beyond  my  knowledge, 
even  to  name  the  innumerable  points  of  structure  in  which  man 
agrees  with  the  other  Primates.  Our  great  anatomist  and 
l^hilosopher.  Prof.  Huxley,  has  fully  discussed  this  subject,^  and 
concludes  that  man  in  all  parts  of  his  organisation  differs  less 
from  the  higher  apes,  than  these  do  from  the  lower  members  of 
the  same  group.  Consequently  there  "is  no  justification  for 
"  placing  man  in  a  distinct  order." 

In  an  early  part  of  this  work  I  brought  forward  various 
facts,  shewing  how  closely  man  agrees  in  constitution  with  the 
higher  mammals ;  and  this  agreement  must  depend  on  our 
close  similarity  in  minute  structure  and  chemical  composition. 
I  gave,  as  instances,  our  liability  to  the  same  diseases,  and  to  the 
attacks  of  allied  parasites ;  our  tastes  in  common  for  the  same 
stimulants,  and  the  similar  effects  produced  by  them,  as  well  as 
by  various  drugs,  and  other  such  facts. 

As  small  unimportant  points  of  resemblance  between  man  and 
the  Quadrumana  are  not  commonly  noticed  in  systematic  works, 
and  as,  when  numerous,  they  clearly  reveal  our  relationship,  I 
will  specify  a  few  such  points.  The  relative  position  of  our 
features  is  manifestly  the  same ;  and  the  various  emotions  are 
displayed  by  nearly  similar  movements  of  the  muscles  and  skin, 
chiefly  above  the  eyebrows  and  round  the  mouth.  Some  few 
expressions  are,  indeed,  almost  the  same,  as  in  the  weeping  of 
certain  kinds  of  monkeys  and  in  the  laughing  noise  made  by 
others,  during  which  the  corners  of  the  mouth  are  drawn  back- 
wards, and  the  lower  eyelids  wrinkled.  The  external  ears  are 
curiously  alike.  In  man  the  nose  is  much  more  prominent  than 
in  most  monkeys ;  but  we  may  trace  the  commencement  of  an 
aquiline  curvature  in  the  nose  of  the  Hoolock  Gibbon  ;  and  this 
in  the  Semnopitliecus  nasica  is  carried  to  a  ridiculous  extreme. 

The  faces  of  many  monkeys  are  ornamented  with  beards, 
whiskers,  or  moustaches.  The  hair  on  the  head  grows  to  a  great 
"»  'Pvoc.  Zoolog.  Soc'  1863,  p.  4. 

.   *  '  Evidence  as  to  Man's  Piace  in  Nature,'  1863,  p.  70,  et  passim. 


CH4P.  vr.  Affinities  and  Genealogy.  151 

length  in  some  species  of  Scmnopithccus ;''  and  in  the  Bonnet 
monkey  (Macacus  radiatus)  it  radiates  from  a  point  on  the  crown, 
with  a  parting  down  the  middle.  It  is  commonly  said  that  the 
forehead  gives  to  man  his  noble  and  intellectual  appearance ;  but 
the  thick  hair  on  the  head  of  the  Bonnet  monkey  terminates 
downwards  abruptly,  and  is  succeeded  by  hair  so  short  and  fine 
that  at  a  little  distance  the  forehead,  with  the  exception  of  the 
eyebrows,  appears  quite  naked.  It  has  been  erroneously  asserted 
that  eyebrows  are  not  present  in  any  monkey.  In  the  species 
just  named  the  degree  of  nakedness  of  the  forehead  differs  in 
different  individuals ;  and  Eschricht  states  ^  that  in  our  children 
the  limit  between  the  hairy  scalp  and  the  naked  forehead  is 
sometimes  not  well  defined;  so  that  here  we  seem  to  have  a 
trifling  case  of  reversion  to  a  yjrogenitor,  in  whom  the  forehead 
had  not  as  yet  become  quite  naked. 

It  is  well  known  that  the  hair  on  our  arms  tends  to  converge 
from  above  and  below  to  a  point  at  the  elbow.  Tliis  curious 
arrangement,  so  unlike  that  in  most  of  the  lower  mammals,  is 
common  to  the  gorilla,  chimpanzee,  orang,  some  species  of 
Hylobates,  and  even  to  some  few  American  monkeys.  But  in 
IJylohates  agilis  the  hair  on  the  fore-arm  is  directed  downwards 
or  towards  the  wrist  in  the  ordinary  manner ;  and  in  //.  lar  it  is 
nearly  erect,  with  only  a  very  slight  forward  inclination ;  so  that 
in  this  latter  species  it  is  in  a  transitional  state.  It  can  hardly 
be  doubted  that  with  most  mammals  the  thickness  of  the  hair  on 
the  back  and  its  direction,  is  adapted  to  throw  off  the  rain ;  even 
the  transverse  hairs  on  the  fore- legs  of  a  dog  may  serve  for  this 
end  when  he  is  coiled  up  asleep.  Mr.  Wallace,  who  has  carefully 
studied  the  habits  of  the  orang,  remarks  that  the  convergence  of 
the  hair  towards  the  elbow  on  the  arms  of  the  orang  may  be 
explained  as  serving  to  throw  off  the  rain,  for  this  animal  during 
rainy  weather  sits  with  its  arms  bent,  and  with  the  hands  clasped 
round  a  branch  or  over  its  head.  According  to  Livingstone,  the 
gorilla  also  "  sits  in  pelting  rain  with  his  hands  over  his  head."" 
If  the  above  explanation  is  correct,  as  seems  probable,  the  direc- 
tion of  the  hair  on  our  own  arms  offers  a  curious  record  of  our 
former  state ;  for  no  one  supposes  that  it  is  now  of  any  use  in 
throwing  off  the  rain ;  nor,  in  our  jDresent  erect  condition,  is  it 
properly  directed  for  this  purpose. 

It  would,  however,  be  rash  to  trust  too  much  to  the  principle 
of  adaptation  in  regard  to  the  direction  of  the  hair  in  man  or  his 

«  Isid.  GeoflFroy,  '  Hist.  Xiit.  Gen.'  Anat.  und  Phys.'  1837,  s.  51. 

torn.  ii.  1859,  p.  217.  "  Quoted  by  Reade,  'The  African 

^    '  Ueber      die     ILichtung       der  Sketch  Book, ''vol.  i.,  1873,  p.  152. 
Haare,'    kc,  Muller's   'Archiv   I'iir 


152  The  Descent  of  Man.    '  Paut  I. 

early  iDrogenitors ;  for  it  is  impossible  to  study  the  figures  given 
by  Escbricht  of  the  arrangement  of  the  hair  on  the  human  foetus 
(this  being  the  same  as  in  the  adult)  and  not  agree  with  this 
excellent  observer  that  other  and  more  complex  causes  have 
intervened.  The  points  of  convergence  seem  to  stand  in  some 
relation  to  those  points  in  the  embryo  which  are  last  closed  in 
duiing  development.  There  appears,  also,  to  exist  some  relation 
between  the  arrangement  of  the  hair  on  the  limbs,  and  the  course 
of  the  medullary  arteries.^ 

It  must  not  be  supposed  that  the  resemblances  between  man 
and  certain  apes  in  the  above  and  many  other  points — such  as  in 
having  a  naked  forehead,  long  tresses  on  the  head,  &c. — are  all 
necessarily  the  result  of  unbroken  inheritance  from  a  common 
progenitor,  or  of  subsequent  reversion.  Many  of  these  resem- 
blances are  more  probably  due  to  analogous  variation,  which 
follows,  as  I  have  elsewhere  attempted  to  shew,^"  from  co-descended 
organisms  having  a  similar  constitution,  and  having  been  acted  on 
by  like  causes  inducing  similar  modifications.  "With  respect  to 
the  similar  direction  of  the  hair  on  the  fore-arms  of  man  and 
certain  monkeys,  as  this  character  is  common  to  almost  all  the 
anthropomorphous  apes,  it  may  probably  be  attributed  to  in- 
heritance; but  this  is  not  certain,  as  some  very  distinct  American 
monkeys  are  thus  characterised. 

Although,  as  we  have  now  seen,  man  has  no  just  right  to  form 
a  separate  Order  for  his  own  reception,  he  may  perhaps  claim  a 
distinct  Sub-order  or  Family.  Prof.  Huxley,  in  his  last  work," 
divides  the  Primates  into  three  Sub-orders;  namely,  the  An- 
thropidse  with  man  alone,  the  Simiadae  including  monkeys  of  all 
kinds,  and  the  Lemuridse  with  the  diversified  genera  of  lemurs. 
As  far  as  differences  in  certain  important  points  of  structure  are 
concerned,  man  may  no  doubt  rightly  claim  the  rank  of  a  Sub- 
order ;  and  this  rank  is  too  low,  if  we  look  chiefly  to  his  mental 
faculties.  Nevertheless,  from  a  genealogical  point  of  view  it 
appears  that  this  rank  is  too  high,  and  that  man  ought  to  form 
merely  a  Family,  or  possibly  even  only  a  Sub-family.  If  we 
imagine  three  lines  of  descent  proceeding  from  a  common  stock, 
it  is  quite  conceivable  that  two  of  them  might  after  the  lapse  of 

^  On  the  hair  iu  Hylobates,  see  the    Theory    of  Natural    Selection,' 

'Nat.  Hist,  of  Mammals,'  by  C.  L.  1870,  p.  344. 

JMartin,    1841,  p.  415.     Also,   l^iJ.  ^^  'Origin    of  Species,'    5th  edit. 

Geoffroy  on  the  American  monkeys  1869,   p.    194.     'The    Variation    of 

and  other  kinds,  '  Hist.  Xat.   Gen.'  Animals  and  Plants  under  Domesti- 

vol.   ii.   1859,  p.    216,    243.     Esch-  cation,'  vol.  ii.  1868,  p.  348. 
richt,   ibiil.    s.   46,   55,   61.     Owen,  ^^   '  An  Introduction  to  the  Classi- 

'  Aunt,   of  Vertebrates,'  vol.   iii.   p.  fication  of  Animals,' 1869,  p.  99. 
619.      Wallace,    'Contributions    to 


Chap.  Y I.  Affinities  and  Genealogy.  153 

ages  be  so  slightly  changed  as  still  to  remain  as  species  of  the 
same  genus,  whilst  the  third  line  might  become  so  greatly 
modified  as  to  deserve  to  rank  as  a  distinct  Sub- family,  Family, 
or  even  Order.  But  in  this  case  it  is  almost  certain  that  the 
third  line  would  still  retain  through  inheritance  numerous  small 
points  of  resemblance  with  the  other  two.  Here,  then,  would 
occur  the  difficulty,  at  present  insoluble,  how  much  weight  we 
ought  to  assign  in  our  classifications  to  strongly-marked  dif- 
ferences in  some  few  points, — that  is,  to  the  amount  of  modifi- 
cation undergone ;  and  how  much  to  close  resemblance  in 
numerous  unimportant  points,  as  indicating  the  lines  of  descent 
or  genealogy.  To  attach  much  weight  to  the  few  but  strong 
differences  is  the  most  obvious  and  perhaps  the  safest  course, 
though  it  appears  more  correct  to  pay  great  attention  to  the 
many  small  resemblances,  as  giving  a  truly  natural  classification. 

In  forming  a  judgment  on  this  head  with  reference  to  man,  we 
must  glance  at  the  classification  of  the  Simiadae.  This  family  is 
divided  by  almost  all  naturalists  into  the  Catarhine  group,  or 
Old  World  monkeys,  all  of  which  are  characterised  (as  their 
name  exjDresses)  by  the  peculiar  structure  of  their  nostrils,  and  by 
having  four  premolars  in  each  jaw;  and  into  the  Platyrhine 
group  or  New  World  monkeys  (including  two  very  distinct 
sub-groups),  all  of  which  are  characterised  by  differently 
constructed  nostrils,  and  by  having  six  j^remolars  in  each  jaw. 
Some  other  small  differences  might  be  mentioned.  Now  man 
unquestionably  belongs  in  his  dentition,  in  the  structure  of  his 
nostrils,  and  some  other  respects,  to  the  Catarhine  or  Old  World 
division ;  nor  does  he  resemble  the  Platyrhines  more  closely  than 
the  Catarhines  in  any  characters,  excepting  in  a  few  of  not  much 
importance  and  apparently  of  an  adaptive  nature.  It  is  therefore 
against  all  probability  that  some  New  World  species  should  have 
formerly  varied  and  produced  a  man-like  creature,  with  all  the 
distinctive  characters  proper  to  the  Old  World  division ;  losing 
at  the  same  time  all  its  own  distinctive  characters.  There  can, 
consequently,  hardly  be  a  doubt  that  man  is  an  off-shoot  from  the 
Old  World  Simian  stem ;  and  that  under  a  genealogical  point  of 
view,  he  must  be  classed  with  the  Catarhine  division.^^ 

The  anthropomorphous  apes,  namely  the  gorilla,  chimpanzee, 

^2  This  is  nearly  the  same  classifi-  adse  which  answer  to  the  Catarhines, 

cation  as  that  provisionally  adopted  the    Cebidae,    and    the    Ilapalida?, — 

by  ]Mr.  St.  George   Mivart   ('  Tran-  these  two  latter   groups  answering 

sact.  Philosoph.  Soc'  1867,  p.  300),  to    the    Platyrhines.      Mr.    Mivart 

who,  after  separating  the  Lemuridffi,  still  abides  by  the  same  view;  see 

divides   the  remainder    of   the   Pri-  '  Nature,'  1871,  p.  481. 
mates  into  the  Hominidac,  the  Simi- 

8 


154  The  Descent  of  Man.  Part  I. 

orang,  and  hylobates,  are  by  most  naturalists  separated  from  the 
other  Old  ^Yorld  monkeys,  as  a  distinct  sub-group.  I  am  aware 
that  Gratiolet,  relying  on  the  structure  of  the  brain,  does  not 
admit  the  existence  of  this  sub-group,  and  no  doubt  it  is  a  broken 
one.  Thus  the  orang,  as  l\Ir.  St.  G.  Mivart  remarks,^^ ''  is  one  of  the 
"  most  peculiar  and  aberrant  forms  to  be  found  in  the  Order." 
The  remaining  non-anthropomorphous  Old  World  monkeys,  are 
again  divided  by  some  naturalists  into  two  or  three  smaller  sub- 
groups ;  the  genus  Semnopithecus,  with  its  pecuhar  sacculated 
stomach,  being  the  type  of  one  such  sub-group.  But  it  appears 
from  M.  Gaudry's  wonderful  discoYeries  in  Attica,  that  during 
the  Miocene  period  a  form  existed  there,  which  connected 
Semnopithecus  and  Macacus ;  and  this  probably  illustrates  the 
manner  in  which  the  other  and  higher  grouj^s  were  once  blended 
together. 

If  the  anthropomorphous  apes  be  admitted  to  form  a  natural 
sul>group,  then  as  man  agrees  with  them,  not  only  in  all  those 
characters  which  he  possesses  in  common  with  the  whole 
Catarhine  groui^,  but  in  other  peculiar  characters,  such  as  the 
absence  of  a  tail  and  of  callosities,  and  in  general  appearance,  we 
may  infer  that  some  ancient  member  of  the  anthropomorphous 
sub-group  gave  birth  to  man.  It  is  not  probable  that,  through 
the  law  of  analogous  yariation,  a  member  of  one  of  the  other 
lower  sub-groups  should  have  given  rise  to  a  man-like  creature, 
resembling  the  higher  anthropomorphous  apes  in  so  many 
respects.  No  doubt  man,  in  comparison  with  most  of  his  allies, 
has  undergone  an  extraordinary  amount  of  modification,  chiefly 
in  consequence  of  the  great  development  of  his  brain  and  his 
erect  position ;  nevertheless,  we  should  bear  in  mind  that  he  "  is 
"  but  one  of  several  exceptional  forms  of  Primates."  ^^ 
-  Every  naturalist,  who  believes  in  the  principle  of  evolution, 
will  grant  that  the  two  main  divisions  of  the  Simiadte,  namely 
the  Catarhine  and  Platyrhine  monkeys,  with  their  sub-groups, 
have  all  proceeded  from  some  one  extremely  ancient  progenitor. 
The  early  descendants  of  this  progenitor,  before  they  had 
diverged  to  any  considerable  extent  from  each  other,  would  still 
have  formed  a  single  natural  group ;  but  some  of  the  species  or 
inci]oient  genera  would  have  already  begun  to  indicate  by  their 
diverging  characters  the  future  distinctive  marks  of  the  Catarhine 
and  Platyrhine  divisions.  Hence  the  members  of  this  supposed 
ancient  group  would  not  have  been  so  uniform  in  their  den- 
tition, or  in  the  structure  of  their  nostrils,  as  are  the  existing 

13  '  Transact,  Zoolog.  Soc'  vol.  vi.  ^^  Mr.  St.  G.  Mivart,  '  Transact. 

IS67,  p.  2U.  Phil.  Soc'  1867,  p.  410. 


Chap.  VI.  Affinities  and  Genealogy.  155 

Catarhine  monkeys  in  one  way  and  the  Platyrhinos  in  another 
way,  but  would  have  resembled  in  this  respect  the  allied  Lemu- 
ridx,  which  differ  greatly  from  each  other  in  the  form  of  their 
muzzles/^  and  to  an  extraordinary  degree  in  their  dentition. 

The  Catarhine  and  Platyrhine  monkeys  agree  in  a  multitude 
of  characters,  as  is  shewn  by  their  unquestionably  belonging 
to  one  and  the  same  Order.  The  many  characters  which 
they  possess  in  common  can  hardly  have  been  independently 
acquired  by  so  many  distinct  species ;  so  that  these  characters 
must  have  been  inherited.  But  a  naturalist  would  undoubtedly 
have  ranked,  as  an  ape  or  a  monkey,  an  ancient  form  which 
possessed  many  characters  common  to  the  Catarhine  and 
Platyrhine  monkeys,  other  characters  in  an  intermediate  con- 
dition, and  some  few,  perhaps,  distinct  from  those  now  found  in 
either  group.  And  as  man  from  a  genealogical  point  of  view 
belongs  to  the  Catarhine  or  Old  World  stock,  we  must  conclude, 
however  much  the  conclusion  may  revolt  our  pride,  that  our 
early  progenitors  would  have  been  properly  thus  designated.'" 
But  we  must  not  fall  into  the  error  of  supposing  that  the  early 
progenitor  of  the  whole  Simian  stock,  including  man,  was  iden- 
tical with,  or  even  closely  resembled,  any  existing  ape  or  monkey. 

0)1  the  Dirthi.lace  and  Antiquify  of  Man. — We  are  naturally 
led  to  enquire,  where  was  the  birthplace  of  man  at  that  stage  of 
descent  when  our  progenitors  diverged  from  the  Catarhine 
stock  ?  The  fact  that  they  belonged  to  this  stock  clearly  shews 
that  they  inhabited  the  Old  World ;  but  not  Australia  nor  any 
oceanic  island,  as  we  may  infer  from  the  laws  of  geographical 
distribution.  In  each  great  region  of  the  world  the  living 
mammals  are  closely  related  to  the  extinct  sjDecies  of  the  same 
region.  It  is  therefore  probable  that  Africa  was  formerly  in- 
habited by  extinct  apes  closely  allied  to  the  gorilla  and  chim- 
panzee ;  and  as  these  two  species  are  now  man's  nearest  allies,  it 
is  somewhat  more  probable  that  our  early  progenitors  lived  on 
the  African  continent  than  elsewhere.  But  it  is  useless  to 
speculate  on  this  subject;  for  two  or  three  anthropomorphous 
apes,  one  the  Dryopithccus ^^  of  Lartet,  nearly  as  large  as  a  man, 

•^  Messrs.  ]\Iurie  and   Mivart   on  his       '  Natiirliche       S-hopfungsge- 

the  Lemuroidea,   'Transact.  Zoolui;.  schichte,'   18G8,  iu  which  he  gives 

Sue'  vol.  vii.  18G9,  p.  5.  iu  detail  his  views  on  the  genealogy 

'^  Hilckel  has  come  to  this  same  of  man. 

conclusion.      See    '  Ueber    die    Ent-  "  Dr.  C.  Forsyth  Major,  '  Sur  les 

stehung    des   Menschengeschlechts,'  Singes  Fossiles    trouves   en   Italie  :* 

in    Virchow's    '  Sammlung.  gemein.  '  Soc.Ital.  des  Sc.  Nat.' torn.  xv.  1872. 
wissen.  Vortriige,'  18G8,  s.  Gl.    Also 


156  The  Descent  of  Man.  Part  1. 

and  closely  allied  to  Hylobates,  existed  in  Europe  during  the 
Miocene  age ;  and  since  so  remote  a  period  the  earth  has 
certainly  undergone  many  great  revolutions,  and  there  has  been 
ample  time  for  migration  on  the  largest  scale. 

At  the  period  and  place,  wheneyer  and  whereTer  it  was,  when 
man  first  lost  his  hairy  coYering,  he  probably  inhabited  a  hot 
country ;  a  circumstance  favourable  for  the  frugiferous  diet  on 
wliich,  judging  from  analogy,  he  subsisted.  We  are  far  from 
knowing  how  long  ago  it  was  when  man  first  diverged  from  the 
Catarhine  stock;  but  it  may  have  occurred  at  an  epoch  as  remote 
as  the  Eocene  period;  for  that  the  higher  apes  had  diverged 
from  the  lower  apes  as  early  as  the  Upper  Miocene  period  is 
shewn  by  the  existence  of  the  Dryopithecus.  We  are  also  quite 
ignorant  at  how  rajDid  a  rate  organisms,  whether  high  or  low  in 
the  scale,  may  be  modified  under  favourable  circumstances  ;  we 
know,  however,  that  some  have  retained  the  same  form  during 
an  enormous  lapse  of  time.  From  what  we  see  going  on  under 
domestication,  we  learn  that  some  of  the  co-descendants  of  the 
same  species  may  be  not  at  all,  some  a  little,  and  some  greatly 
changed,  all  within  the  same  period.  Thus  it  may  have  been 
with  man,  who  has  undergone  a  great  amount  of  modification 
in  certain  characters  in  comparison  with  the  higher  apes. 

The  great  break  in  the  organic  chain  between  man  and  his 
nearest  allies,  which  cannot  be  bridged  over  by  any  extinct  or 
living  species,  has  often  been  advanced  as  a  grave  objection  to 
the  belief  that  man  is  descended  from  some  lower  form ;  but  this 
objection  will  not  appear  of  much  weight  to  those  who,  from 
general  reasons,  believe  in  the  general  princij)le  of  evolution. 
Breaks  often  occur  in  all  parts  of  the  series,  some  being  wide, 
sharp  and  defined,  others  less  so  in  various  degrees ;  as  between 
the  orang  and  its  nearest  allies — betw^een  the  Tarsius  and  the 
other  Lemuridse — between  the  elephant,  and  in  a  more  striking 
manner  between  the  Ornithorhynchus  or  Echidna,  and  a.l  other 
mammals.  But  these  breaks  depend  merely  on  the  number  of 
related  forms  which  have  become  extinct.  At  some  future 
period,  not  very  distant  as  measured  by  centuries,  the  civilised 
races  of  man  will  almost  certainly  exterminate,  and  replace,  the 
savage  races  throughout  the  world.  ^  At  the  same  time  the  anthro- 
pomorphous apes,  as  Professor  Schaafifliausen  has  remarked,^^ 
will  no  doubt  be  exterminated.  The  break  between  man  and  his 
nearest  allies  will  then  be  wider,  for  it  will  intervene  between 
man  in  a  more  civilised  state,  as  we  may  hope,  even  than  the 
Caucasian,  and  some  ai)e  as  low  as  a  baboon,  instead  of  as  no  «v 
between  the  negro  or  Australian  and  the  gorilla. 

"  '  Anthropological  Review/  April,  1867,  p.  236. 


Chap.  YI.  Affinities  and  Genealogy.  1 57 

"With  respect  to  the  absence  of  fossil  remains,  serving  to 
connect  man  with  his  ape-like  progenitors,  no  one  will  lay  much 
stress  on  this  fact  who  reads  Sir  C.  Ly ell's  discussion,'^  where 
he  shews  that  in  all  the  vertebrate  classes  the  discovery  of  fossil 
remains  has  been  a  very  slow  and  fortuitous  process.  Nor 
should  it  be  forgotten  that  those  regions  which  are  the  most 
hkely  to  afford  remains  connecting  man  with  some  extinct  ape- 
like creature,  have  not  as  yet  been  searched  by  geologists. 

Lower  Stages  in  the  Genealogy  of  Man. — We  have  seen  that 
man  appears  to  have  diverged  from  the  Catarhine  or  Old  World 
division  of  the  Simiadae,  after  these  had  diverged  from  the  New 
World  division.  We  will  now  endeavour  to  follow  the  remote 
traces  of  his  genealogy,  trusting  principally  to  the  mutual 
affinities  between  the  various  classes  and  orders,  with  some 
slight  reference  to  the  periods,  as  far  as  ascertained,  of  their 
successive  appearance  on  the  earth.  The  Lemuridae  stand 
below  and  near  to  the  Simiadse,  and  constitute  a  very  distinct 
family  of  the  Primates,  or,  according  to  Hackel  and  others,  a 
distinct  Order.  This  group  is  diversified  and  broken  to  an 
extraordinary  degree,  and  includes  many  aberrant  forms.  It 
has,  therefore,  i:)robably  suffered  much  extinction.  Most  of  the 
remnants  survive  on  islands,  such  as  Madagascar  and  the 
Malayan  archipelago,  where  they  have  not  been  exposed  to  so 
severe  a  competition  as  they  would  have  been  on  well-stocked 
continents.  This  group  likewise  presents  many  gradations, 
leading,  as  Huxley  remarks,-*' ''  insensibly  from  the  crown  and 
"  summit  of  the  animal  creation  down  to  creatures  from  which 
"  there  is  but  a  step,  as  it  seems,  to  the  lowest,  smallest,  and 
"  least  intelligent  of  the  placental  mammalia."  From  these 
various  considerations  it  is  probable  that  the  Simiadai  were 
originally  developed  from  the  progenitors  of  the  existing 
Lemnridaj ;  and  these  in  their  turn  from  forms  standing  very 
low  in  the  mammalian  series. 

The  Marsupials  stand  in  many  important  characters  below  the 
placental  mammals.  They  appeared  at  an  earlier  geological 
period,  and  their  range  was  formerly  much  more  extensive 
than  at  present.  Hence  the  Placentata  are  generally  supposed 
to  have  been  derived  from  the  Implacentata  or  Marsui)ials; 
not,  however,  from  forms  closely  resembling  the  existing  Mar- 
supials, l)ut  from  their  early  progenitors.  The  Monotremata  are 
plainly  allied  to  the  Marsupials,  forming  a  third  and  still  lower 

'»  'Elements    of  Geology,'  1865,  20  <  Man's    Place    in    Nature,'   p. 

pp.  583-585.     '  Antiquity  of  Man,'      105. 
1863,  p.  145. 


158  The  Descent  of  Man,  Paut  I 

division  in  tlie  great  mammalian  series.  They  are  represented 
at  the  present  day  solely  by  the  Ornithorhynchus  and  Echidna  ; 
and  these  two  fonns  may  be  safely  considered  as  relics  of  a 
much  larger  group,  representatives  of  which  have  been  preserved 
in  Australia  through  ^ome  favourable  concurrence  of  circum- 
stances. The  Monotremata  are  eminently  interesting,  as  leading 
in  several  important  points  of  structure  towards  the  class  of 
reptiles. 

In  attempting  to  trace  the  genealogy  of  the  Mammalia,  and 
therefore  of  man,  lower  down  in  the  series,  we  become  involved 
in  greater  and  greater  obscurity ;  but  as  a  most  capable  judge, 
Mr.  Parker,  has  remarked,  we  have  good  reason  to  believe,  that 
no  true  bird  or  reptile  intervenes  in  the  direct  line  of  descent. 
He  who  wishes  to  see  what  ingenuity  and  knowledge  can  effect, 
may  consult  Prof.  Hackel's  works.^^  I  will  content  myself  with 
a  few  general  remarks.  Every  evolutionist  will  admit  that  the 
five  great  vertebrate  classes,  namely,  mammals,  birds,  reptiles, 
amphibians,  and  fishes,  are  descended  from  some  one  prototype ; 
for  they  have  much  in  common,  especially  during  their  embryonic 
state.  As  the  class  of  fishes  is  the  most  lowly  organised,  and 
appeared  before  the  others,  we  may  conclude  that  all  the 
members  of  the  vertebrate  kingdom  are  derived  from  some  fish- 
like animal.  The  belief  that  animals  so  distinct  as  a  monkey, 
an  elephant,  a  humming-bird,  a  snake,  a  frog,  and  a  fish,  &c.,  could 
all  have  sprung  from  the  same  parents,  will  appear  monstrous 
to  those  who  have  not  attended  to  the  recent  progress  of  natural 
history.  For  this  belief  implies  the  former  existence  of  links 
binding  closely  together  all  these  forms,  now  so  utterly  unlike. 

Nevertheless,  it  is  certain  that  groups  of  animals  have  existed, 
or  do  now  exist,  which  serve  to  connect  several  of  the  great 
vertebrate  classes  more  or  less  closely.  We  have  seen  that  the 
Ornithorhynchus  graduates  towards  reptiles ;  and  Prof.  Huxley 
has  discovered,  and  is  confirmed  by  Mr.  Cope  and  others,  that 
the  Dinosaurians  are  in  many  important  characters  intermediate 
between  certain  reptiles  and  certain  birds— the  birds  referred 
to  being  the  ostrich- tribe  (itself  evidently  a  widely-diffused 
remnant  of  a  larger  group)  and  the  Archeopteryx,  that  strange 
Secondary  bird,  with  a  long  lizard-like  tail.   Again,  according  to 

21  Elaborate  tables  are   given  in  the   phylum  or  lines  of  descent  of 

his  *  Generelle  Morphologie '  (B.  ii.  the  Vertebrata  to  be  admirabl}'  dis- 

s.  cliii.  and  s.  425)  ;  and  with  more  cussed  by  Hackel,  although  he  differs 

especial    reference    to   man    in    his  on    some    points.       He     expresses, 

'Natiirliche    Schopfuugsgeschichte,'  also,     his    high     estimate     of    the 

1868.     Prof.  Huxley,  in   reviewing  general    tenor    and    spirit    of    the 

this   latter  work    ('The   Academj-/  whole  work. 

18(39,  p.  42)  says,  that  he"  considers  22     Palaeontology,'  1860,  p.  199. 


Chap.  VI.  Affinities  and  Genealogy.  159 


Prof.  Owen,-2  the  Tchthyosanrians— great  sea-lizards  furnished 
with  paddles— present  many  affinities  with  fishes,  or  rather, 
according  to  Huxley,  with  amphibians  ;  a  class  which,  including 
in  its  highest  division  frogs  and  toads,  is  plainly  allied  to  the 
Ganoid  fishes.  These  latter  fishes  swarmed  during  the  earlier 
geological  periods,  and  were  constructed  on  what  is  called  a 
generalised  type,  that  is,  they  presented  diversified  affinities  with 
other  groups  of  organisms.  The  Lepidosiren  is  also  so  closely 
allied  to  amphibians  and  fishes,  that  naturalists  long  disputed  in 
which  of  these  two  classes  to  rank  it ;  it,  and  also  some  few 
Ganoid  fishes,  have  been  preserved  from  ntter  extinction  by 
inhabiting  rivers,  which  are  harbours  of  refuge,  and  are  related 
to  the  great  waters  of  the  ocean  in  the  same  way  that  islands 
are  to  continents. 

Lastly,  one  single  member  of  the  immense  and  diversified  class 
of  fishes,  namely,  the  lancelet  or  amphioxus,  is  so  different  from 
all  other  fishes,  that  Hackel  maintains  that  it  ought  to  form  a 
distinct  class  in  the  vertebrate  kingdom.  This  fish  is  remarkable 
for  its  negative  characters ;  it  can  hardly  be  said  to  possess  a 
brain,  vertebral  column,  or  heart,  &c. ;  so  that  it  was  classed  by 
the  older  naturalists  amongst  the  worms.  Many  years  ago  Prof. 
Goodsir  perceived  that  the  lancelet  presented  some  affinities  with 
the  Ascidians,  which  are  invertebrate,  hermaphrodite,  marine 
creatures  permanently  attached  to  a  support.  They  hardly 
appear  like  animals,  and  consist  of  a  simple,  tough,  leathery 
sack,  with  two  small  projecting  orifices.  They  belong  to  the 
Molluscoida  of  Huxley— a  lower  division  of  the  great  kingdom 
of  the  Mollusca ;  but  they  have  recently  been  placed  by  some 
naturahsts  amongst  the  Vermes  or  worms.  Their  larvae  some- 
what resemble  tadpoles  in  shape,-^  and  have  the  power  of 
swimming  freely  about.  M.  Kovalevsky  -■*  has  lately  observed  that 
the  larvae  of  Ascidians  are  related  to  the  Vertebrata,  in  their 
manner  of  development,  in  the  relative  position  of  the  nervous 
system,  and  in  possessing  a  structure  closely  like  the  chorda 
dorsalis  of  vertebrate  animals;    and  in  this  he  has  been  since 

-^  At  the  Falkland  Islands  I  had  under  a  simple  microscope,  plainly 

tlie  satisfaction  of  seeing,  in  April  divided  by  transverse  opaque  parti- 

1833,  and  therefore  some  years  be-  tions,    which    I    presume    represent 

tore  any  other  naturalist,  the  loco-  the  great  cells  figured  by  Kovalev- 

motive   larvae  of  a  compound  Asci-  sky.     At  an  early  stage  of  develop- 

dian,    closely    allied    to    Synoicum,  raent    the    tail    was    closely    coiled 

but  apparently  renerically  distinct  round  the  head  of  the  larva, 

from   it.     The   tail  was    about  five  ^^    '  Memoiros     de     I'Acad.     des 

times  as  long  as  the  oblong  head,  Sciences  de  St.  Pe'tersliourg,'  torn.  x. 

and  terminated   in  a  very  fine   fila-  No.  15,  186G. 
ment.     It  was,  as  bketclied   by  me 


i6o  The  Descent  of  Man.  Pakt  I. 

confirmed  by  Prof.  Kupffer.  M.  Kovalevsky  writes  to  me  from 
Naples,  that  he  has  now  carried  these  observations  yet  further ; 
and  should  his  results  be  well  established,  the  whole  will  form  a 
discovery  of  the  very  greatest  value.  Thus,  if  we  may  rely  on 
embryology,  ever  the  safest  guide  in  classification,  it  seems  that 
we  have  at  last  gained  a  clue  to  the  source  whence  the  Yertebrata 
were  derived.^^  We  should  then  be  justified  in  believing 
that  at  an  extremely  remote  period  a  group  of  animals  existed, 
resembling  in  many  respects  the  larvae  of  our  present  Ascidians, 
which  diverged  into  two  great  branches— the  one  retrograding  in 
development  and  producing  the  present  class  of  Ascidians,  the 
other  rising  to  the  crown  and  summit  of  the  animal  kingdom  by 
giving  birth  to  the  Yertebrata. 

We  have  thus  far  endeavoured  rudely  to  trace  the  genealogy 
of  the  Yertebrata  by  the  aid  of  their  mutual  aflBnities.  We  will 
now  look  to  man  as  he  exists ;  and  we  shall,  I  think,  be  able 
partially  to  restore  the  structure  of  our  early  progenitors,  during 
successive  iDeriods,  but  not  in  due  order  of  time.  This  can  be 
eff'ected  by  means  of  the  rudiments  which  man  still  retains,  by 
the  characters  which  occasionally  make  their  appearance  in  him 
through  reversion,  and  by  the  aid  of  tlie  principles  of  morphology 
and  embryology.  The  various  facts,  to  which  I  shall  here  allude, 
have  been  given  in  the  previous  chapters. 

The  early  progenitors  of  man  must  have  been  once  covered 
with  hair,  both  sexes  having  beards ;  their  ears  were  probably 
pointed,  and  capable  of  movement ;  and  their  bodies  were  pro- 
vided with  a  tail,  having  the  proper  muscles.  Their  limbs  and 
bodies  were  also  acted  on  by  many  muscles  which  now  only 
occasionally  reappear,  but  are  normally  present  in  the  Quadru- 
mana.  At  this  or  some  earlier  period,  the  great  artery  and  nerve 
of  the  humerus  ran  thi'ough  a  supra-condyloid  foramen.  The 
intestine  gave  forth  a  much  larger  diverticulum  or  caecum  than 
that  now  existing.  The  foot  was  then  prehensile,  judging  from 
the  condition  of  the  great  toe  in  the  foetus ;  and  our  progenitors, 
no  doubt,  were  arboreal  in  their  habits,  and  frequented  some 
warm,  forest-clad  land.    The  males  had  great  canine  teeth,  which 

25  But  I  am   bound  to  add  that  "  peut  produire  la  disposition  fonda- 

some  competent  judges  dispute  this  "  mentale    du    type  A-ertebre    (I'ex- 

couclusion ;    for   instance,  M.  Giard,  "  istence  d'une  corde  dorsale)  chez 

in  a  series  of  papers  in  the 'Archives  "  un   iuvertebre   par  la   seule    con- 

dc  Zoologie  Experimentale,'  for  1872.  "  dition     vitale      de      I'adaptation, 

Nevertheless,     this     naturalist    re-  "  et     cette    simple     possibilite     du 

marks,  p.  28i,  "  L'organisaiion  de  la  "passage    supprirae    I'abime    entre 

"  larve    ascidienne     en     dehors    de  "  les  deux  sous-regnes,  encore    bien 

"  toute  hypothfese  etde  toutethebrie,  *' qu'en   ignore    par   oil    le    passage 

"  nous  montre  comment  la  nature  '•  s'est  fait  en  reality." 


Chap.  VI.  Affinities  and  Ge7iealogy.  i6i 

served  them  as  formidable  weapons.  At  a  miicli  earlier  period 
the  uterus  was  double;  the  excreta  were  voided  through  a  cloaca; 
and  the  eye  was  protected  by  a  third  eyelid  or  nictitating  mem- 
brane. At  a  still  earlier  period  the  progenitors  of  man  must  have 
been  aquatic  in  their  habits;  for  morjAology  plainly  tells  us  that 
our  lungs  consist  of  a  modified  swim-bladder,  which  once  served 
as  a  float.  The  clefts  on  the  neck  in  the  embryo  of  man  show 
where  the  branchiss  once  existed.  In  the  lunar  or  weekly  re- 
current periods  of  some  of  our  functions  we  apparently  still  retain 
traces  of  our  primordial  birthplace,  a  shore  washed  by  the  tides. 
At  about  this  same  early  period  the  true  kidneys  were  replaced 
by  the  corpora  wolffiana.  The  heart  existed  as  a  simple  pulsating 
vessel;  and  the  chorda  dorsalis  took  the  place  of  a  vertebral 
column.  These  early  ancestors  of  man,  thus  seen  in  the  dim 
recesses  of  time,  must  have  been  as  simply,  or  even  still  more 
simply  organised  than  the  lancelet  or  amphioxus. 

There  is  one  other  point  deserving  a  fuller  notice.  It  has  long 
been  known  that  in  the  vertebrate  kingdom  one  sex  bears 
rudiments  of  various  accessory  parts,  appertaining  to  the  re- 
productive system,  which  properly  belong  to  the  opposite  sex ; 
and  it  has  now  been  ascertained  that  at  a  very  early  embryonic 
period  both  sexes  possess  true  male  and  female  glands.  Hence 
some  remote  progenitor  of  the  whole  vertebrate  kingdom  appears 
to  have  been  hermaphrodite  or  androgynous.^*^  But  here  we 
encounter  a  singular  difficulty.  In  the  mammalian  class  the 
males  possess  rudiments  of  a  uterus  with  the  adjacent  passage, 
in  their  vesiculse  prostaticse ;  they  bear  also  rudiments  of 
mammae,  and  some  male  Marsupials  have  traces  of  a  marsupial 
sack.^^  Other  analogous  facts  could  be  added.  Are  we,  then,  to 
suppose  that  some  extremely  ancient  mammal  continued  andro- 
gynous, after  it  had  acquired  the  chief  distinctions  of  its  class, 
and  therefore  after  it  had  diverged  from  the  lower  classes  of  the 
vertebrate  kingdom  ?  This  seems  very  improbable,  for  we  have 
to  look  to  fishes,  the  lowest  of  all  the  classes,  to  find  any  still 
existent  androgynous  forms.'-^^      That  various  accessory  parts, 

-^  This  is  the  conclusion  of  Prof.  *'  brata  a]-e,  in  their  early  condition. 

Gegenbaur,  one  of  the  highest  au-  "  hermaphrodite."      Similar    views 

thorities    in  comparative  anatomy;  have  long  been  held  by  som«  authors, 

sec'Grundziigeder  vergleich.  Anat.'  though    until    recently    without    a 

1870,  s.  876.     The  result  has  been  firm  basis. 

arrived  at  chiefly  from  the  study  of  ^^  The  male  Thylacinus  offers  the 

the  Amphibia;  but  it  appears  from  best  instance.     Owen,  'Anatomy  of 

the    researches     of     Waldeyer.  (as  Vertebrates,'  vol.  iii.  p.  771. 

quoted    in    'Journal    of  Anat.    and  ^^  Hermaphroditism  has  been  ob- 

Phys.'  1869,^  p.  161),  that  the  sexual  served  in  several  species  of  Serranus, 

oriT-ms  of   even  "the  higher  verte-  as    well    a^    in    some    other    fishes, 


1 62  The  Descent  of  Man.  Pakt  I. 

proper  to  each  sex,  are  found  in  a  rudimentary  condition  in  the 
opposite  sex,  may  he  explained  by  such  organs  having  been 
gradually  acquired  by  the  one  sex,  and  then  transmitted  in  a 
more  or  less  imperfect  state  to  the  other.  When  we  treat  of 
sexual  selection,  we  shall  meet  with  innumerable  instances  of 
this  form  of  transmission, — as  in  the  case  of  the  spurs,  plumes, 
and  brilliant  colours,  acquired  for  battle  or  ornament  by  male 
birds,  and  inherited  by  the  females  in  an  imperfect  or  rudimentary 
condition. 

The  possession  by  male  mammals  of  functionally  imperfect 
mammary  organs  is,  in  some  respects,  especially  curious.  The 
Monotremata  have  the  proper  milk-secreting  glands  with  orifices, 
but  no  nipples ;  and  as  these  animals  stand  at  the  very  base  of 
the  mammalian  series,  it  is  probable  that  the  progenitors  of 
the  class  also  had  milk-secreting  glands,  but  no  nipples.  This 
conclusion  is  supported  by  what  is  known  of  their  manner  of 
development ;  for  Professor  Turner  informs  me,  on  the  authority 
of  Kolliker  and  Langer,  that  in  the  embryo  the  mammary  glands 
can  be  distinctly  traced  before  the  nipples  are  in  the  least 
visible;  and  the  development  of  successive  parts  in  the  indi- 
vidual generally  represents  and  accords  with  the  development  of 
successive  beings  in  the  same  line  of  descent.  The  Marsupials 
differ  from  the  Monotremata  by  possessing  nipples;  so  that 
probably  these  organs  were  first  acquired  by  the  Marsupials, 
after  they  had  diverged  from,  and  risen  above,  the  Monotremata, 
and  were  then  transmitted  to  the  placental  mammals.^^  No  one 
will  suppose  that  the  Marsupials  still  remained  androgynous 
after  they  had  approximately  acquired  their  present  structure. 
How  then  are  we  to  account  for  male  mammals  possessing 
mammae  ?  It  is  possible  that  they  were  first  developed  in  the 
females  and  then  transferred  to  the  males ;  but  from  what 
follows  this  is  hardly  probable. 

where  it  is  either  normal  and  sym-  delle    Scienze,'    Bologna,    Doc.     28, 

metrical,    or    abnormal    and     uni-  1871)  that  eels  ai-e  androgynous, 

lateral.     Dr.    Zouteveen   has    given  ^^   Prof.    Gegenbaur    has    shewn 

me  references  on  this  subject,  more  (*  Jenaische   Zeitschrift,'  Bd.  vii.  p. 

especially  to  a  paper  by  Prof  Hal-  212)   that    two    distinct    types    of 

bertsma,  in    the    '  Transact,   of  the  nipples     prevail      throughout     the 

Dutch  Acad,  of  Sciences,'  vol.  xvi.  several     mammalian     orders,      but 

Dr.  Giinther   doubts    the    fact,  but  that  it  is  quite  intelligible  how  both 

it    has    now  been   recorded   by  too  could  have  been  derived  from    the 

many    good    observers     to    be    any  nipples  of  the  Marsupials,  and  the 

longer    disputed.       Dr.    M.   Lessona  latter  from   those  of  the  Monotre- 

writes   to  me,    that    he    has   veri-  mata.     See,  also,  a  memoir  by  Dr. 

fied     the     observations     made      by  Max  Huss,  on  the  mammary  glands, 

Cavolini   on  Serranus.     Prof    Ereo-  ibid.  B.  viii.  p.  176. 
lani    has    recently    shewn    (' Accad. 


Chap.  VI.  Affinities  and  Genealogy.  163 


It  may  be  suggested,  as  another  view,  that  long  after  the 
progenitors  of  the  whole  mammalian  class  had  ceased  to  be 
androgynous,  both  sexes  yielded  milk,  and  thus  nourished  their 
young ;  and  in  the  case  of  the  Marsupials,  that  both  sexes  carried 
their  young  in  marsupial  sacks.  This  will  not  appear  altogether 
improbable,  if  we  reflect  that  the  males  of  existing  syngnathous 
fishes  receive  the  eggs  of  the  females  in  their  abdominal  pouches, 
hatch  them,  and  afterwards,  as  some  believe,  nourish  the 
young  ;^ — that  certain  other  male  fishes  hatch  tlie  eggs  within 
their  mouths  or  branchial  cavities ; — that  certain  male  toads 
take  the  chaplets  of  eggs  from  the  females,  and  wind  them  round 
their  own  thighs,  keeping  them  there  until  the  tad])oles  are 
born ; — that  certain  male  birds  undertake  the  whole  duty  of 
incubation,  and  that  male  pigeons,  as  well  as  the  females,  feed 
their  nestlings  with  a  secretion  from  their  crops.  But  the  above 
suggestion  first  occurred  to  me  from  the  mammary  glands  of 
male  mammals  being  so  much  more  perfectly  developed  than 
the  rudiments  of  the  other  accessory  reproductive  parts,  which 
are  found  in  the  one  sex  though  proper  to  the  other.  The 
mammary  glands  and  nipples,  as  they  exist  in  male  mammals, 
can  indeed  hardly  be  called  rudimentary ;  they  are  merely  not 
f Lilly  developed,  and  not  functionally  active.  They  are  sympa- 
thetically affected  under  the  influence  of  certain  diseases,  like 
the  same  organs  in  the  female.  They  often  secrete  a  few  drops 
of  milk  at  birth  and  at  puberty  :  this  latter  fact  occurred  in  the 
curious  case,  before  referred  to,  where  a  young  man  possessed 
two  pairs  of  mammae.  In  man  and  some  other  male  mammals 
these  organs  have  been  known  occasionally  to  become  so  well 
developed  during  maturity  as  to  yield  a  fair  supply  of  milk. 
Now  if  we  suppose  that  during  a  former  prolonged  jDcriod  male 
mammals  aided  the  females  in  nursing  their  offspring,^^  and  that 
afterwards  from  some  cause  (as  from  the  production  of  a  smaller 
number  of  young)  the  males  ceased  to  give  this  aid,  disuse  of  the 
organs  during  maturity  would  lead  to  their  becoming  inactive  ; 
and  from  two  well-known  principles  of  inheritance,  this  state  of 
inactivity  would  probably  be  transmitted  to  the  males  at  the 
corresponding  age  of  maturity.     But  at  an  earlier  age  these 

'°  Mr.  Lockwood  believes  (ns  by  Prof.  Wyman,  in  '  Proc.  Boston 
quoted  in 'Quart.  Journal  of  Science,'  Soc.  of  Nat.  Hist.'  Sept.  15,1857; 
April,  1868,  p.  269),  from  what  he  also  Prof.  Turner,  in  'Journal  of 
has  observed  of  the  development  of  Anat.  and  Phys.'  Nov.  1,  186(3,  p. 
Hippocampus,  that  the  walls  of  the  78.  Dr.  Giinther  has  likewi.se  de- 
abdominal  pouch  of  the  male  in  scribed  similar  cases, 
some  way  aflbrd  nourishment.  On  ''  Madlle.  C.  Pvoyer  has  suggested 
male  fishes  hatching  the  ova  in  their  a  similar  view  in  her  '  Origine  de 
mouths,  see  a  very  interesting  paper  I'Homme,'  (S:c.,  1870. 


1 64  The  Descent  of  Man.  Part  I. 

organs  would  be  left  unaffected,  so  that  they  would  be  almost 
equally  well  developed  in  the  young  of  both  sexes. 

Conclusion. — Yon  Baer  has  defined  advancement  or  progress  in 
the  organic  scale  better  than  any  one  else,  as  resting  on  the 
amount  of  differentiation  and  specialisation  of  the  several  parts 
of  a  being, — when  arrived  at  maturity,  as  I  should  be  inclined  to 
add.  Now  as  organisms  have  become  slowly  adapted  to  diver- 
sified lines  of  life  by  means  of  natural  selection,  their  parts  will 
have  become  more  and  more  differentiated  and  specialised  for 
various  functions,  from  the  advantage  gained  by  the  division  of 
physiological  labour.  The  same  part  appears  often  to  have  been 
modified  first  for  one  purpose,  and  then  long  afterwards  for 
some  other  and  quite  distinct  purpose ;  and  thus  all  the  parts 
are  rendered  more  and  more  complex.  But  each  organism  still 
retains  the  general  type  of  structure  of  the  progenitor  from 
which  it  was  aboriginally  derived.  In  accordance  with  this 
view  it  seems,  if  we  turn  to  geological  evidence,  that  organisa- 
tion on  the  whole  has  advanced  throughout  the  world  by  slow 
and  interrupted  steps.  In  the  great  kingdom  of  the  Vertebrata 
it  has  culminated  in  man.  It  must  not,  however,  be  supposed 
that  groups  of  organic  beings  are  always  supplanted,  and  dis- 
appear as  soon  as  they  have  given  birth  to  other  and  more 
perfect  groups.  The  latter,  though  victorious  over  their  pre- 
decessors, may  not  have  become  better  adapted  for  all  places  in 
the  economy  of  nature.  Some  old  forms  appear  to  have  survived 
from  inhabiting  protected  sites,  where  they  have  not  been 
exposed  to  very  severe  competition ;  and  these  often  aid  us  in 
constructing  our  genealogies,  by  giving  us  a  fair  idea  of  former 
and  lost  populations.  But  we  must  not  fall  into  the  error  of 
looking  at  the  existing  members  of  any  lowly-organised  group  as 
perfect  representatives  of  their  ancient  predecessors. 

The  most  "ancient  progenitors  in  the  kingdom  of  the  Vertebrata, 
at  which  we  are  able  to  obtain  an  obscure  glance,  apparently 
consisted  of  a  group  of  marine  animals,^^  resembling  the  larvae  of 
existing  Ascidians.     These  animals    probably   gave  rise  to  a 

3^  The  inhabitants  of  the  sea-  ditions  for  many  generations,  can 
.shore  must  be  greatly  affected  by  hardly  fail  to  run  their  course  in 
the  tides;  animals  living  either  regular  weekly  periods.  Now  it  is  a 
aboixt  the  mean  high-water  marlv,  mysterious  fact  that  in  the  higher 
or  about  the  mean  low-water  mark,  and  now  terrestrial  Vertebrata,  as 
pass  through  a  complete  cycle  of  well  as  in  other  classes,  many  nor- 
tidal  changes  in  a  fortnight.  Con-  mal  and  abnormal  processes  have 
sequently,  their  food  supply  will  one  or  more  whole  weeks  as  their 
undergo  marked  changes  week  by  periods ;  this  would  be  rendered 
week.  The  A'ital  functions  of  such  intelligible  if  the  Vertebrata  are  de- 
animals,    living    under    these    con-  scended  from    an   animal    allied   to 


Chap.  VJ,  Affinities  and  Genealogy.  165 

group  of  fishes,  as  lowly  organised  as  the  lancelet ;  and  from 
these  the  Ganoids,  and  other  fishes  like  the  Lcpidosiren,  must 
have  been  developed.  From  such  fish  a  very  small  advance 
would  carry  us  on  to  the  Amphibians.  AYe  have  seen  that  birds 
and  reptiles  were  once  intimately  connected  together ;  and  the 
Monotremata  now  connect  mammals  with  reptiles  in  a  slight 
degree.  But  no  one  can  at  present  say  by  what  line  of  descent 
the  three  higher  and  related  classes,  namely,  mammals,  birds, 
and  reptiles,  were  derived  from  the  two  lower  vertebrate  classes, 
namely,  amphibians  and  fishes.  In  the  class  of  mammals  the 
steps  are  not  difficult  to  conceive  which  led  from  the  ancient 
Monotremata  to  the  ancient  Marsupials ;  and  from  these  to  the 
early  progenitors  of  the  placental  mammals.  "We  may  thuy 
ascend  to  the  Lemuridse  ;  and  the  interval  is  not  very  wide  from 
these  to  the  Simiadae.  The  Simiadse  then  branched  off  into  two 
great  stems,  the  New  World  and  Old  World  monkeys ;  and  from 
the  latter,  at  a  remote  period,  Man,  the  wonder  and  glory  of  the 
Universe,  i^roceeded. 

Thus  we  have  given  to  man  a  pedigree  of  prodigious  length,  but 
not,  it  may  be  said,  of  noble  quality.  The  world,  it  has  often 
been  remarked,  appears  as  if  it  had  long  been  preparing  for  the 
advent  of  man :  and  this,  in  one  sense  is  strictly  true,  for  he 
owes  his  birth  to  a  long  line  of  progenitors.  If  any  single  link 
in  this  chain  had  never  existed,  man  would  not  have  been  exactly 
what  he  now  is.  Unless  we  wilfully  close  our  eyes,  we  may,  with 
our  present  knowledge,  approximately  recognise  our  parentage ; 
nor  need  we  feel  ashamed  of  it.  The  most  humble  organism  is 
something  much  higher  than  the  inorganic  dust  under  our  feet ; 
and  no  one  with  an  unbiassed  mind  can  study  any  living 
creature,  however  humble,  without  being  struck  with  enthusiasm 
at  its  marvellous  structure  and  properties. 


the  existing  tidal  Asoirlians.  ^Many  cess  or  function,  would  not,  when 
instances  ot"  such  periodic  processes  once  gained,  be  liable  to  change; 
might  be  given,  as  the  gestation  of  consequently  it  might  be  thus  trans- 
mammals,  the  duration  ot"  fevers,  &c.  mitted  through  almost  any  number 
The  hatching  of  eggs  affords  also  a  of  geneivitions.  But  if  the  function 
good  example,  for,  according  to  Mr.  changed,  the  period  would  have  to 
Bartlett  ('  Land  and  Water,'  Jan.  7,  change,  and  would  be  apt  to  change 
1871),  the  eggs  of  the  pigeon  are  almost  abruptly  by  a  whole  week, 
hatched  in  two  weeks  ;  those  of  the  This  conclusion',  if  sound,  is  highly 
fowl  in  three  ;  those  of  the  duck  in  remarkable  ;  for  the  period  of  gesta- 
four ;  those  of  the  goose  in  five ;  tion  in  each  mammal,  and  the 
and  those  of  the  ostrich  in  seven  hatching  of  each  bird's  eggs,  and 
weeks.  As  far  as  we  can  judge,  a  many  other  vital  processes,  thus 
recurrent  period,  if  approximately  betray  to  us  the  primordial  birth- 
of  the  right  duration  tor  any  pro-  place  of  these  animals. 


1 66  The  Descent  of  Man.  Tabt  I. 


CHAPTER  VII. 
On  the  Eaces  of  Man. 

The  nature  and  value  of  specific  characters — Application  to  the  races  of 
ii:an — Arguments  in  favour  of,  and  opposed  to,  ranking  the  so-called 
races  of  man  as  distinct  species — Sub-species — Monogenists  and  poly- 
genists — Convergence  of  character — Numerous  points  of  resemblance  in 
body  and  mind  between  the  most  distinct  races  of  man — The  state  of 
man  when  he  first  spread  over  the  earth — Each  race  not  descended  from 
a  single  pair — The  extinction  of  races — The  formation  ot  races — The 
effects  of  crossing — Slight  influence  of  the  direct  action  of  the  con- 
ditions of  life — Slight  or  no  influence  of  natural  selection — Sexual 
selection. 

It  is  not  iny  intention  here  to  describe  the  several  so-called  races 
of  men ;  but  I  am  about  to  enquire  what  is  the  value  of  the  dif- 
ferences between  them  under  a  classificatory  point  of  view,  and 
how  they  have  originated.  In  determining  whether  two  or  more 
allied  forms  ought  to  be  ranked  as  species  or  varieties,  naturalists 
are  practically  guided  by  the  following  considerations;  namely,  the 
amount  of  difference  between  them,  and  whether  such  differences 
relate  to  few  or  many  points  of  structure,  and  whether  they  are 
of  physiological  importance ;  but  more  especially  whether  they 
are  constant.  Constancy  of  character  is  what  is  chiefly  valued 
and  sought  for  by  naturalists.  "Whenever  it  can  be  shewn,  or 
rendered  probable,  that  the  forms  in  question  have  remained 
distinct  for  a  long  period,  this  becomes  an  argument  of  much 
weight  in  favour  of  treating  them  as  species.  Even  a  slight 
degree  of  sterility  between  any  two  forms  when  first  crossed,  or 
in  their  offspring,  is  generally  considered  as  a  decisive  test  of 
their  specific  distinctness;  and  their  continued  persistence 
without  blending  witliin  the  same  area,  is  usually  accepted  as 
sufficient  evidence,  either  of  some  degree  of  mutual  steiility,  or 
in  the  case  of  animals  of  some  mutual  repugnance  to  pairing. 

Independently  of  fusion  from  intercrossing,  the  complete 
absence,  in  a  well-investigated  region,  of  varieties  linking 
together  any  two  closely-allied  forms,  is  probably  the  most 
important  of  all  the  criterions  of  their  specific  distinctness  ;  and 
this  is  a  somewhat  different  consideration  from  mere  constancy 
of  character,  for  two  forms  may  be  highly  variable  and'  yet  not 
yield  intermediate  varieties.  Geographical  distribution  is  often 
brought  into  play  unconsciously  and  sometimes  consciously ;  so 
that  forms  living  in  two  widely  separated  areas,  in  which  most 


Ohap.  TIT.  The  Races  of  Man,  167 

of  the  other  inhabitants  are  specifically  distinct,  are  themselves 
usually  looked  at  as  distinct ;  but  in  truth  this  affords  no  aid  in  dis- 
tinguishing geographical  races  from  so-called  good  or  true  species. 

Now  let  us  apply  these  generally- admitted  principles  to  the 
races  of  man,  viewing  him  in  the  same  spirit  as  a  naturalist  would 
any  other  animal.  In  regard  to  the  amount  of  difference  between 
the  races,  we  must  make  some  allowance  for  our  nice  powers  of 
discrimination  gained  by  the  long  habit  of  observing  ourselves. 
In  India,  as  Elphinstone  remarks,  although  a  newly-arrived 
European  cannot  at  first  distinguish  the  various  native  races, 
yet  they  soon  appear  to  him  extremely  dissimilar  ■}  and  the 
Hindoo  cannot  at  first  perceive  any  difference  between  the  several, 
European  nations.  Even  the  most  distinct  races  of  man  are 
much  more  like  each  other  in  form  than  would  at  first  be  sup- 
posed ;  certain  negro  tribes  must  be  excepted,  whilst  others,  as 
Dr.  Eohlfs  writes  to  me,  and  as  I  have  myself  seen,  have 
Caucasian  features.  This  general  similarity  is  well  shewn  by 
the  French  photographs  in  the  Collection  Anthropologique  du 
]\Iuseum  de  Paris  of  the  men  belonging  to  various  races,  the 
greater  number  of  which  might  pass  for  Europeans,  as  many 
persons  to  whom  I  have  shewn  them  have  remarked.  Neverthe- 
less, these  men,  if  seen  alive,  would  undoubtedly  appear  very 
distinct,  so  that  we  are  clearly  much  influenced  in  our  judgment 
by  the  mere  colour  of  the  skin  and  hair,  by  slight  differences  in 
the  features,  and  by  expression. 

There  is,  however,  no  doubt  that  the  various  races,  when 
carefully  compared  and  measured,  differ  much  from  each  other, 
— as  in  the  texture  of  the  hair,  the  relative  proportions  of  all  parts 
of  the  body,^  the  capacity  of  the  lungs,  the  form  and  capacity  of 
the  skull,  and  even  in  the  convolutions  of  the  brain.^  But  it 
would  be  an  endless  task  to  specify  the  numerous  points  of 
difference.  The  races  differ  also  in  constitution,  in  acclimatisation 
and  in  liability  to  certain  diseases.  Their  mental  characteristics 
are  likewise  very  distinct ;  chiefly  as  it  would  appear  in  their 
emotional,  but  partly  in  their  intellectual  faculties.  Every  one 
who  has  had  the  opportunity  of  comparison,  must  have  been 

*  'History  of  India,'  184-1,  vol.  i.  'On  the  capacity  of  the  lungs,' p.  471. 

p.  323.     Father  Ripa  makes  exactly  See  also  the  numerous  and  valuable 

the  same    remark   with   respect    to  tables,  by  Dr.  Weisbach,   fi-om    the 

the  Chinese.  observations    of    Dr.    Scherzer   and 

2  A  vast  number  of  measure-  Dr.  Schwarz,  in  the  '  Reise  der 
ments  of  Whites,  Bhicks,  and  In-  Kovara :  Anthropolog.  Theil,'  1867. 
dians,  are  given  in  the  'Investiga-  •*  See,  for  instance,  Mr.  Marshall's 
tions  in  the  Military  and  Anthropo-  account  of  the  brain  of  a  Bush- 
log.  Statistics  of  American  Soldiers,'  woman,  in  'Phil.  Transact.'  186-1-, 
by  B.  A.  Gould,  1869,  pp.  298-358  ;  p.  519. 


1 68  The  Descent  of  Man.  Part  I. 

struck  with  the  contrast  between  the  taciturn,  even  morose, 
aborigines  of  S.  America  and  the  light-hearted,  talkative  negroes. 
There  is  a  nearly  similar  contrast  between  the  IMalays  and  the 
Papuans,*  who  live  under  the  same  physical  conditions,  and  are 
separated  from  each  other  only  by  a  narrow  space  of  sea. 

We  will  first  consider  the  arguments  which  may  be  advanced 
in  favour  of  classing  the  races  of  man  as  distinct  species,  and 
then  the  arguments  on  the  other  side.  If  a  naturalist,  who  had 
never  before  seen  a  Negro,  Hottentot,  Australian,  or  Mongolian, 
were  to  compare  them,  he  would  at  once  perceive  that  they 
differed  in  a  multitude  of  characters,  some  of  slight  and  some  of 
considerable  importance.  On  enquiry  he  would  find  that  they 
were  adapted  to  live  under  widely  different  cUmates,  and  that 
they  differed  somewhat  in  bodily  constitution  and  mental  dis- 
position. If  he  were  then  told  that  hundreds  of  similar  specimens 
could  be  brought  from  the  same  countries,  he  would  assuredly 
declare  that  they  were  as  good  species  as  many  to  which  he  had 
been  in  the  habit  of  affixing  specific  names.  This  conclusion 
would  be  greatly  strengthened  as  soon  as  he  had  ascertained  that 
these  forms  had  all  retained  the  same  character  for  many 
centuries ;  and  that  negroes,  apparently  identical  with  existing 
negroes,  had  lived  at  least  4000  years  ago.^  He  would  also  hear, 
on  the  authority  of  an  excellent  observer,  Dr.  Lund,^  that  the 
human  skulls  'found  in  the  caves  of  Brazil,  entombed  with  many 
extinct  mammals,  belonged  to  the  same  tyj)e  as  that  now  pre- 
vaibng  throughout  the  American  Continent. 

<  Wallace,    'The     Malay    Archi-  man  (' Races  of  Man,'  1850,  p.  201), 

pelago,'  vol.  ii.  1869,  p.  178.  speaking    of    young    Men?.non    (the 

^  With  respect  to  the  figures  in  same    as  Rameses   II.,  as  I  am    in- 

the  famous  Egyptian  caves  of  Abou-  formed  by  Mr.  Birch),  insists  in  the 

Simbel,    M.     Pouchet     says    ( '  The  strongest  manner  that  he  is  identical 

Plurality  of  the  Human  Races,' Eng.  in  character  with  the  Jev/s  of  Ant- 

translat.  1861:,  p.  50),  that  he  was  werp.     Again,  when  I  looked  at  the 

far  from  finding  recognisable  repre-  statue  of  Amunoph  III.,  I  agreed  with 

sentations    of    the    dozen    or    more  two    officers    of   the    establishment, 

nations  which  some  authors  beliere  both  competent  judges,  that  he  had 

that  they  can  recognise.     Even  some  a    strongly  marked    negro    type    of 

of  the  most  strongly-marked  races  features;     but     Messrs.     Nott    and 

cannot   be   identified  with   that  de-  Gliddon    (ibid.  p.   146,  fig.  53)  de- 

gi"oe  of  unanimity  which  might  have  scribe  him  as  a  hybrid,  but  not   of 

been  expected   from  what   has   been  "  negro  intermixture." 

written     on     the     subject.       Thus  ^  As  quoted  by  Nott  and  Gliddon, 

Messrs.  Nott   and   Gliddon    ('Types  'Types  of  Mankind,'  1854,  p.  439. 

of    Mankind,'    p.    148)    state    that  They    give    also    corroborative    evi- 

Rameses    II.,    or     the    Great,    has  dence ;  but  C.  Vogt  thinks  that  the 

features  suped'bly  European  ;  where-  subject  requires    further  investiga- 

as   Knox,  another  firm   believer   in  tion. 
the  specific  distinctness  of  the  races  of 


(Jhap.  VJI.  The  Races  of  Man.  169 

Our  naturalist  would  then  perhaps  turn  to  geographical  dis- 
tribution, and  he  would  probably  declare  that  those  forms  must 
be  distinct  species,  which  differ  not  only  in  aj^pcarance,  but 
are  fitted  for  hot,  as  well  as  damp  or  dry  countries,  and  for  the 
Arctic  regions.  He  might  appeal  to  the  fact  that  no  species  in 
the  group  next  to  man,  namely  the  Quadrumana,  can  resist  a  low 
temperature,  or  any  considerable  change  of  climate ;  and  that 
the  species  which  come  nearest  to  man  have  never  been  reared 
to  maturity,  even  under  the  temperate  climate  of  Europe.  He 
would  be  deeply  impressed  with  the  fact,  first  noticed  by  Agassiz," 
that  the  different  races  of  man  are  distributed  over  the  world  in 
the  same  zoological  provinces,  as  those  inhabited  by  undoubtedly 
distinct  species  and  genera  of  mammals.  This  is  manifestly  the 
case  with  the  Australian,  Mongolian,  and  Negro  races  of  man ;  in 
a  less  well-marked  manner  with  the  Hottentots;  but  plainly 
with  the  Papuans  and  Malays,  who  are  separated,  as  Mr.  Wallace 
has  shewn,  by  nearly  the  same  line  which  divides  the  great 
Malayan  and  Australian  zoological  provinces.  The  Aborigines 
of  America  range  throughout  the  Continent ;  and  this  at  first 
appears  opposed  to  the  above  rule,  for  most  of  the  productions  of 
the  Southern  and  Northern  halves  differ  widely :  yet  some  few 
living  forms,  as  the  opossum,  range  from  the  one  into  the  other, 
as  did  formerly  some  of  the  gigantic  Edentata.  The  Esquimaux, 
like  other  Arctic  animals,  extend  round  the  whole  polar  regions. 
It  should  be  observed  that  the  amount  of  difference  between  the 
mammals  of  the  several  zoological  provinces  does  not  correspond 
with  the  degree  of  separation  between  the  latter ;  so  that  it  can 
hardly  be  considered  as  an  anomaly  that  the  Negro  differs  more, 
and  the  American  much  less  from  the  other  races  of  man,  than 
do  the  mammals  of  the  African  and  American  continents  from 
the  mammals  of  the  other  provinces.  Man,  it  may  be  added, 
does  not  appear  to  have  aboriginally  inhabited  any  oceanic  island ; 
and  in  this  respect  he  resembles  the  other  members  of  his  class. 

In  determining  whether  the  supposed  varieties  of  the  same 
kind  of  domestic  animal  should  be  ranked  as  such,  or  as  spe- 
cifically distinct,  that  is,  whether  any  of  them  are  descended  from 
distinct  wild  species,  every  naturalist  would  lay  much  stress  on 
the  fact  of  their  external  parasites  being  specifically  distinct. 
All  the  more  stress  would  be  laid  on  this  fact,  as  it  would  be  an 
exceptional  one ;  for  I  am  informed  by  Mr.  Denny  that  the  most 
different  kinds  of  dogs,  fowls,  and  pigeons,  in  England,  are 
infested  by  the  same  species  of  Pediculi  or  lice.  Now  Mr.  A. 
Murray  has  carefully  examined  the  Pediculi  collected  in  different 

'  'Diversity  of  Origin  of  the  Pluman  Races,'  in  the  'Christian 
Examiner,'  July  1850. 


I/O  The  Descent  of  Man.  Tart  T. 


countries  from  the  different  races  of  man ;  *  and  he  finds  that 
they  differ,  not  only  in  coloiir,  but  in  the  structure  of  their 
claws  and  limbs.  In  every  case  in  -svliicli  many  specimens  were 
obtained  the  differences  were  constant.  The  surgeon  of  a  whaling 
ship  in  the  Pacific  assured  me  that  when  the  Pediculi,  with 
which  some  Sandwich  Islanders  on  board  swarmed,  strayed  on 
to  the  bodies  of  the  Enghsh  sailors,  they  died  in  the  course  of 
three  or  four  days.  These  Pediculi  were  darker  coloured,  and 
appeared  different  from  those  proper  to  the  natives  of  Chiloe  in 
South  America,  of  which  he  gave  me  Sjoecimens.  These,  again, 
appeared  larger  and  much  softer  than  European  lice.  Mr. 
Murray  procured  four  kinds  from  Africa,  namely  from  the  Negroes 
of  the  Eastern  and  Western  coasts,  from  the  Hottentots  and 
Kaflirs ;  two  kinds  from  the  natives  of  Australia ;  two  from  North 
and  two  from  South  America.  In  these  latter  cases  it  may  be 
presumed  that  the  Pediculi  came  from  natives  inhabiting  different 
districts.  With  insects  shght  structural  differences,  if  constant, 
are  generally  esteemed  of  specific  value :  and  the  fact  of  the 
races  of  man  being  infested  by  parasites,  which  appear  to  be 
specifically  distinct,  might  fairly  be  urged  as  an  argument  that 
the  races  themselves  ought  to  be  classed  as  distinct  species. 

Our  supposed  naturalist  having  proceeded  thus  far  in  his 
investigation,  would  next  enquire  whether  the  races  of  men,  when 
crossed,  were  in  any  degree  sterile.  He  might  consult  the  work* 
of  Professor  Broca,  a  cautious  and  i^bilosophical  observer,  and  in 
this  he  would  find  good  evidence  that  some  races  were  quite 
fertile  together,  but  evidence  of  an  opposite  nature  in  regard  to 
other  races.  Thus  it  has  been  asserted  that  the  native  women  of 
Australia  and  Tasmania  rarely  produce  children  to  European 
men  ;  the  evidence,  however,  on  this  head  has  now  been  shewn 
to  be  almost  valueless.  The  half-castes  are  killed  by  the  pure 
blacks :  and  an  account  has  lately  been  published  of  eleven  half- 
caste  youths  murdered  and  burnt  at  the  same  time,  whose 
remains  were  found  by  the  police.^'^  Again,  it  has  often  been 
said  that  when  mulattoes  intermarry  they  produce  few  children; 
on  the    other  hand,  Dr.  Bachman   of  Charleston"  positively 

*  'Transact.  R.  Soc.  of  Edinburgh,'  who  have  borne  children  to  a  white 
vol.  xxiL  1861,  p.  567.  man    are    afterwards    sterile    with 

*  '  On  the  Phenomena  of  Hybridity  their  own  race,  is  disproved.  M.  A. 
m  the  Genus  Homo,'  Eng.  transhit.  de  Quatrefages  has  also  collected 
1864.  ('  Revue    des    Cours    Scientifiques,' 

'"  See  the  interesting    letter   by  March  1869,  p.  239)  much   evidence 

Mr.  T.  A.  Murray,  in  the  '  Anthro-  that  Australians  and  Europeans  are 

polog.   Review,'  April  1868,  p.  liii.  not  sterile  when  crossed. 
la    this    letter   Count    Strzelecki's  *^    '  An     Examination     of    Prof, 

statement,  that    Australian  women  Agassiz's  Sketch    of  the  Nat.   Pro- 


Chap.  VII.  The  Races  of  Man.  17 1 


asserts  that  lie  lias  known  mulatto  families  wliioli  have  inter- 
married for  several  generations,  and  have  continued  on  an 
average  as  fertile  as  either  pure  whites  or  pure  blacks.  Enquiries 
formerly  made  by  Sir  C.  Lyell  on  this  subject  led  him,  as  he 
informs  me,  to  the  same  conclusion.'-  In  the  United  States  the 
census  for  the  year  1851  included,  according  to  Dr.  Bachman, 
405,751  mulattoes  ;  and  this  number,  considering  all  the  circum- 
stances of  the  case,  seems  small ;  but  it  may  2:»art]y  be  accounted 
for  by  the  degraded  and  anomalous  position  of  the  class,  and  by 
the  profligacy  of  the  women.  A  certain  amount  of  absorption  of 
mulattoes  into  negroes  must  always  be  in  progress ;  and  this 
would  lead  to  an  apparent  diminution  of  the  former.  The  inferior 
vitality  of  mulattoes  is  spoken  of  in  a  trustworthy  work^^  as  a 
well-known  phenomenon ;  and  this,  although  a  different  considera- 
tion from  their  lessened  fertility,  may  perhaps  be  advanced  as 
a  proof  of  the  specific  distinctness  of  the  jDarent  races.  No  doubt 
both  animal  and  vegetable  hybrids,  when  j)roduced  from  extremely 
distinct  species,  are  liable  to  premature  death;  but  the  parents 
of  mulattoes  cannot  be  put  under  the  category  of  extremely 
distinct  species.  The  common  Mule,  so  notorious  for  long  life 
and  vigour,  and  yet  so  sterile,  shews  how  little  necessary  con- 
nection there  is  in  hybrids  between  lessened  fertility  and  vitality ; 
other  analogous  cases  could  be  cited. 

Even  if  it  should  hereafter  be  proved  that  all  the  races  of 
men  were  perfectly  fertile  together,  he  who  was  inclined  from 
other  reasons  to  rank  them  as  distinct  species,  might  with  justice 
argue  that  fertility  and  sterility  are  not  safe  criterions  of  specific 
distinctness.  We  know  that  these  qualities  are  easily  affected 
by  changed  conditions  of  life,  or  by  close  inter-breeding,  and  that 
they  are  governed  by  highly  complex  laws,  for  instance,  that  of 
the  unequal  fertility  of  converse  crosses  between  the  same  two 
species.  With  forms  which  must  be  ranked  as  undoubted 
species,  a  perfect  series  exists  from  those  which  are  absolutely 
sterile  when  crossed,  to  those  which  are  almost  or  completely 


viucesof  the  Animal  World,' Charles-  the    children   are    i^^ii   and    sickly, 

ton,  1855,  p.  44.  This  belief,  as  Mr.  Reade  remarks, 

^2  Dr.  Rohlt's  writes  to  me  that  deserves    attention,    as    white    men 

he    found    the   mixed   races    in  the  have  visited  and  resided  on  the  Gold 

Great  Sahara,  derived   from  Arabs,  Coast    for    four    hundred    years,    so 

Berbers,  and  Negroes  of  three  tribes,  that   the    natives   have    had    ample 

extraordinarily  fertile.    On  the  other  time    to    gain    knowledge    through 

hand,  Mr.  VViuwood  Reade   informs  experience. 

me  that  the   Negroes  ou  the   Gold  '^    '  Military    and    Anthropolog. 

Coast,  though   admiring  white  men  Statistics  of  American  Soldiers,'  by 

and  mulattoes,  have  a  maxim  that  B.  A.  Gould,  18G9,  p.  319. 
mulattoes  should  not  intermarrv,  as 


172 


The  Descent  of  Man. 


Part  T. 


fertile.  The  degrees  of  sterility  do  not  coincide  strictly  with 
the  degrees  of  difference  between  the  parents  in  external  structure 
or  habits  of  life.  Man  in  many  respects  may  be  compared  with 
those  animals  which  have  long  been  domesticated,  and  a  large 
body  of  evidence  can  be  advanced  in  favour  of  the  Pallasian 
doctrine/*  that  domestication  tends  to  eliminate  the  sterility 
which  is  so  general  a  result  of  the  crossing  of  species  in  a  state 
of  nature.  From  these  several  considerations,  it  may  be  justly 
urged  that  the  perfect  fertility  of  the  intercrossed  races  of  man, 
if  estabnshed,  would  not  absolutely  preclude  us  from  ranking 
them  as  distinct  species. 

Independently  of  fertility,  the  characters  presented  by  the  off- 
spring from  a  cross  have  been  thought  to  indicate  whether  or  not 
the  parent-forms  ought  to  be  ranked  as  species  or  varieties ;  but 
after  carefully  studying  the  evidence,  I  have  come  to  the  con- 
clusion that  no  general  rules  of  this  kind  can  be  trusted.  The 
ordinary  result  of  a  cross  is  the  production  of  a  blended  or 


'*  '  The  Variation  of  Animals  and 
Plants  under  Domestication,'  vol.  ii. 
p.  109.  I  may  here  remind  the 
reader  that  the  sterility  of  species 
when  crossed  is  not  a  specially- 
acquired  quality,  but,  like  the  in- 
capacity of  certain  trees  to  be  graft- 
ed together,  is  incidental  on  other 
acquired  differences.  The  nature 
of  these  differences  is  unknown,  but 
they  relate  more  especially  to  the  re- 
productive system,  and  much  less  so 
to  external  structure  or  to  ordinary 
differences  in  constitution.  One 
important  element  in  the  sterility 
of  crossed  species  apparently  lies  in 
one  or  both  having  been  long  habi- 
tuated to  fixed  conditions  ;  for  we 
know  that  changed  conditions  have 
a  special  influence  on  the  repro- 
diictive  system,  and  we  have  good 
reason  to  believe  (as  before  re- 
marked) that  the  fluctuating  con- 
ditions of  domestication  tend  to 
eliminate  that  sterility  which  is  so 
general  with  species,  in  a  natural 
state,  when  crossed.  It  has  else- 
where been  shewn  by  me  (ibid.  vol. 
ii.  p.  185,  and  'Origin  of  Species' 
5th  edit.  p.  317),  that  the  sterility 
of  crossed  species  has  not  been  ac- 
quired through  natural  selection : 
we  can  see  that  when  two  forms 
have    already    been    rendered    very 


sterile,  it  is  scarcely  possible  that 
their  sterility  should  be  augmented 
by  the  preservation  or  survival  of 
the  more  and  more  sterile  indi- 
viduals ;  for  as  the  sterility  in- 
creases, fewer  and  fewer  offspring 
will  be  produced  from  which  to 
breed,  and  at  last  only  single  in- 
dividuals will  be  produced,  at  the 
rarest  intervals.  But  there  is  even 
a  higher  grade  of  sterility  than 
this.  Both  Gartner  and  Kolreuter 
have  proved  that  in  genera  of  plants 
including  many  species,  a  series 
can  be  formed  from  species  which 
when  crossed  yield  fewer  and  fewer 
seeds,  to  species  which  never  pro- 
duce a  single  seed,  but  yet  are 
affected  by  the  pollen  of  the  other 
species,  as  shewn  by  the  swelling 
of  the  germen.  It  is  here  mani- 
festly impossible  to  select  the  more 
sterile  individuals,  which  have  al- 
ready ceased  to  yield  seeds ;  so  that 
the  acme  of  sterility,  when  the 
germen  alone  is  affected,  cannot 
have  been  gained  through  selection. 
This  acme,  and  no  doubt  the  other 
grades  of  steriiity,  are  the  incidental 
results  of  certain  unknown  differ- 
ences in  the  constitution  of  the  re- 
productive system  of  the  species 
which  are  crossed. 


Chap.  Yll.  The  Races  of  Man.  173 

intermediate  form  ;  but  in  certain  cases  some  of  the  ofiFspring  take 
closely  after  one  parent-form,  and  some  after  the  other.  This  is 
especially  apt  to  occur  when  the  parents  differ  in  characters 
which  first  appeared  as  sudden  variations  or  monstrosities.^^  I 
refer  to  this  point,  because  Dr.  Rohlfs  informs  me  that  he  has 
frequently  seen  in  Africa  the  offspring  of  negroes  crossed  with 
members  of  other  races,  either  completely  black  or  completely 
white,  or  rarely  piebald.  On  the  other  hand,  it  is  notorious 
that  in  America  mulattocs  commonly  present  an  intermediate 
appearance. 

We  have  now  seen  that  a  naturalist  might  feel  himself  fully 
justified  in  ranking  the  races  of  man  as  distinct  species ;  for  he 
has  found  that  they  are  distinguished  by  many  differences  in 
structure  and  constitution,  some  being  of  importance.  These 
differences  have,  also,  remained  nearly  constant  for  very  long 
periods  of  time.  Our  naturalist  will  have  been  in  some  degree 
influenced  by  the  enormous  range  of  man,  which  is  a  great 
anomaly  in  the  class  of  mammals,  if  mankind  be  viewed  as  a 
single  species.  He  will  have  been  struck  w^ith  the  distribution  of 
the  several  so-called  races,  which  accords  with  that  of  other 
undoubtedly  distinct  species  of  mammals.  Finally,  he  might 
urge  that  the  mutual  fertility  of  all  the  races  has  not  as  yet  been 
fully  proved,  and  even  if  proved  would  not  be  an  absolute  proof 
of  their  specific  identity. 

On  the  other  side  of  the  question,  if  our  supposed  naturalist 
were  to  enquire  whether  the  forms  of  man  keep  distinct  like 
ordinary  species,  when  mingled  together  in  large  numbers  in  the 
same  country,  he  would  immediately  discover  that  this  was  by 
no  means  the  case.  In  Brazil  he  w^ould  behold  an  immense 
mongrel  population  of  Negroes  and  Portuguese ;  in  Chiloe,  and 
other  parts  of  South  America,  he  would  behold  the  whole  popu- 
lation consisting  of  Indians  and  Spaniards  blended  in  various 
degrees.^"  In  many  parts  of  the  same  continent  he  would  meet 
with  the  most  complex  crosses  between  Negroes,  Indians,  and 
Europeans ;  and  judging  from  the  vegetable  kingdom,  such  triple 
crosses  afford  the  severest  test  of  the  mutual  fertility  of  the 
parent-forms.  In  one  island  of  the  Pacific  he  w^ould  find  a 
small  population  of  mingled  Polynesian  and  English  blood;  and 
m  the  Fiji  Archipelago  a  population  of  Polynesian  and  Negritos 

'*  '  The    Yariatiou    of  Animals,'  success  and  energy  of  the  Paulistas 

&c.,  vol.  ii.  p.  92.  in  Brazil,  who  are  a  much  crossed 

'*  M.  de  Quatrefages    has    given  race  of  Portuguose  and  Indians,  with 

(' Anthropolog.  Keview,'  Jan.   1869,  a   mixture    of   the    blood    of  other 

p.  22)  an  interesting  account  of  the  races. 


1/4  T lie  Descent  of  Man.  Part  I. 

crossed  in  all  degrees.  Many  analogous  cases  could  be  be  added  ; 
for  instance,  in  Africa.  Hence  the  races  of  man  are  not  suf- 
ficiently distinct  to  inhabit  the  same  country  without  fusion ; 
and  the  absence  of  fusion  affords  the  usual  and  best  test  of 
si^ecific  distinctness. 

Our  naturalist  would  likewise  be  much  disturbed  as  soon  as 
he  perceived  that  the  distinctive  characters  of  all  the  races  were 
highly  variable.  This  fact  strikes  every  one  on  first  beholding 
the  negTO  slaves  in  Brazil,  who  have  been  imported  from  all 
parts  of  Africa.  The  same  remark  holds  good  with  the 
Polynesians,  and  with  many  other  races.  It  may  be  doubted 
whether  any  character  can  be  named  which  is  distinctive  of  a 
race  and  is  constant.  Savages,  even  within  the  limits  of  the 
same  tribe,  are  not  nearly  so  uniform  in  character,  as  has  been 
often  asserted.  Hottentot  women  offer  certain  peculiarities, 
more  strongly  marked  than  those  occurring  in  any  other  race, 
but  these  are  known  not  to  be  of  constant  occurrence.  In  the 
several  American  tribes,  colour  and  hairiness  differ  considerably; 
as  does  colour  to  a  certain  degree,  and  the  shape  of  the  features 
greatly,  in  the  Negroes  of  Africa.  The  shape  of  the  skull  varies 
much  in  some  races  ;^^  and  so  it  is  with  every  other  character. 
Now  all  naturalists  have  learnt  by  dearly-bought  experience,  how 
rash  it  is  to  attempt  to  define  species  by  the  aid  of  inconstant 
chara,cters. 

But  the  most  weighty  of  all  the  arguments  against  treating 
the  races  of  man  as  distinct  si)ecies,  is  that  they  graduate  into 
each  other,  independently  in  many  cases,  as  far  as  we  can  judge, 
of  their  having  intercrossed.  Man  has  been  studied  more 
carefully  than  any  other  animal,  and  yet  there  is  the  greatest 
possible  diversity  amongst  capable  judges  whether  he  should  be 
classed  as  a  single  species  or  race,  or  as  two  (Virey),  as  three 
(Jacquinot),  as  four  (Kant),  five  (Blumenbach),  six  (Buffon), 
seven  (Hunter),  eight  (Agassiz),  eleven  (Pickering),  fifteen 
(Bory  St.  Vincent),  sixteen  (Desmoulins),  tw^enty-two  (Morton), 
sixty  (Crawfurd),  or  as  sixty-three,  according  to  Burke.^^  This 
diversity  of  judgment  does  not  prove  that  the  races  ought  not 
to  be  ranked  as  sj^ecies,  but  it  shews  that  they  graduate  into  each 

^^  For    instance  with    the  abori-  ^*  See   a   good  discussion  on  this 

gines    of    America    and    Australia.  subject    in     Waitz,    '  Introduct.     to 

Prof.  Huxley  says  (' Transact.  Inter-  Anthropology,'  Eng.  translat.  1863, 

nat.     Congress    of    Prehist.    Arch.'  pp.    198-208,    227.     I    have    taken 

1868,   p.    105)    that    the    skulls    of  some  of  the  above  statements  from 

many  South  Germans  and  Swiss  are  H.  Tuttle's  '  Origin    and  Antiquity 

"  as  short  and  as  broad  as  those  of  of  Physical  Man,'  Boston,  1866,  p. 

"the  Tartars,"  &c.  35. 


Cii^?.  VII.  The  Races  of  Man.  175 

other,  and  tbat  it  is  hardly  possible  to  discover  clear  distinctive 
characters  between  tliem. 

Every  naturalist  who  has  had  the  misfortune  to  undertake  the 
description  of  a  group  of  highly  varying  organisms,  has  en- 
countered cases  (I  speak  after  experience)  precisely  like  that  of 
man ;  and  if  of  a  cautious  disposition,  he  will  end  by  uniting  all 
the  forms  which  graduate  into  each  other,  under  a  single 
species  ;  for  he  will  say  to  himself  that  he  has  no  right  to  give 
names  to  objects  which  he  cannot  define.  Cases  of  this  kind 
occur  in  the  Order  which  includes  man,  namely  in  certain  genera 
of  monkeys;  whilst  in  other  genera,  as  in  Cercopithecus,  most  of 
the  species  can  be  determined  with  certainty.  In  the  American 
genus  Cebus,  the  various  forms  are  ranked  by  some  naturalists 
as  species,  by  others  as  mere  geographical  races.  Now  if 
numerous  sjiecimens  of  Cebus  were  collected  from  all  parts  of 
South  America,  and  those  forms  which  at  present  appear  to  be 
specifically  distinct,  were  found  to  graduate  into  each  other  by 
close  steps,  they  would  usually  be  ranked  as  mere  varieties  or 
races ;  and  this  course  has  been  followed  by  most  naturalists 
with  respect  to  the  races  of  man.  Nevertheless,  it  must  be 
confessed  that  there  are  forms,  at  least  in  the  vegetable  king- 
dom,'^ which  we  cannot  avoid  naming  as  species,  but  which  are 
connected  .  together  by  numberless  gradations,  independently  of 
intercrossing. 

Some  naturalists  have  lately  employed  the  term  "  sub-species" 
to  designate  forms  which  possess  many  of  the  characteristics  of 
true  species,  but  which  hardly  deserve  so  high  a  rank.  Now  if 
we  reflect  on  the  weighty  arguments  above  given,  for  raising  the 
races  of  man  to  the  dignity  of  species,  and  the  insuperable  diffi- 
culties on  the  other  side  in  defining  them,  it  seems  that  the  term 
"  sub-species  "  might  here  be  used  with  propriety.  But  from 
long  habit  the  term  *'  race  "  will  perhaps  always  be  employed. 
The  choice  of  terms  is  only  so  far  imi3ortant  in  that  it  is  desirable 
to  use,  as  far  as  possible,  the  same  terms  for  the  same  degrees  of 
difiFerence.  Unfortunately  this  can  rarely  be  done :  for  the  larger 
genera  generally  include  closely-aUied  forms,  which  can  be 
distinguished  only  with  much  difficulty,  whilst  the  smaller 
genera  within  the  same  family  include  forms  that  are  perfectly 
distinct ;  yet  all  must  be  ranked  equally  as  species.  So  again, 
species  within  the  same  large  genus  by  no  means  resemble 
each  other  to  the  same  degree  r  on  the  contrary,  some  of  them 

^^  Prof.  Nagfli  has  carefully  de-  has    made    analogous    remarks    on 

Bcribed  several  striking  cases  in  his  some    intermediate     forms    in    the 

'  Botanische    Mittheilungen,'    B.    ii.  Composita;  of  N.  America 
18G6,  s.  294-369.     Prof.  Asa  Gray 


176.  The  Descent  of  Man.  Part  I. 

can  generally  be  arranged  in  little  groups  round  other  species, 
like  satellites  round  planets.'^" 

The  question  whether  mankind  consists  of  one  or  several 
species  has  of  late  years  been  much  discussed  by  anthropologists, 
who  are  divided  into  the  two  schools  of  monogenists  and 
polygenists.  Those  who  do  not  admit  the  principle  of  evolution, 
must  look  at  species  as  separate  creations,  or  as  in  some  manner 
as  distinct  entities ;  and  they  must  decide  what  forms  of  man  they 
will  consider  as  species  by  the  analogy  of  the  method  commonly 
pursued  in  ranking  other  organic  beings  as  species.  But  it  is  a 
hopeless  endeavour  to  decide  this  point,  until  some  definition  of 
the  term  "  species "  is  generally  accepted ;  and  the  definition 
must  not  include  an  indeterminate  element  such  as  an  act  of 
creation.  We  might  as  well  attempt  without  any  definition  to 
decide  whether  a  certain  number  of  houses  should  be  called  a 
village,  town,  or  city.  "We  have  a  practical  illustration  of  the 
diflBcultyin  the  never-ending  doubts  whether  many  closely-allied 
mammals,  birds,  insects,  and  plants,  which  represent  each 
other  respectively  in  North  America  and  Europe,  should  be 
ranked  as  species  or  geographical  races  ;  and  the  like  holds  true 
of  the  productions  of  many  islands  situated  at  some  little  distance 
from  the  nearest  continent. 

Those  naturalists,  on  the  other  hand,  who  admit  the  principle 
of  evolution,  and  this  is  now  admitted  by  the  majority  of  rising 
men,  will  feel  no  doubt  that  all  the  races  of  man  are  descended 
from  a  single  primitive  stock ;  whether  or  not  they  may  think 
fit  to  designate  the  races  as  distinct  species,  for  the  sake  of  ex- 
pressing their  amount  of  difference. ^^  With  our  domestic 
animals  the  question  whether  the  various  races  have  arisen  from 
one  or  more  species  is  somewhat  different.  Although  it  may  be 
admitted  that  all  the  races,  as  well  as  all  the  natural  species 
within  the  same  genus,  have  sprung  from  the  same  primitive 
stock,  yet  it  is  a  fit  subject  for  discussion,  whether  all  the 
domestic  races  of  the  dog,  for  instance,  have  acquired  their 
present  amount  of  difference  since  some  one  species  was  first 
domesticated  by  man ;  or  whether  they  owe  some  of  their 
characters  to  inheritance  from  distinct  species,  which  had 
already  been  differentiated  in  a  state  of  nature.  With  man  no 
such  question  can  arise,  for  he  cannot  be  said  to  have  been 
domesticated  at  any  particular  period. 

During  an  early  stage  in  the  divergence  of  the  races  of  man 

20  <  Origin  of  Species,'  5th  edit,  in  the  '  Fortnightly  Review,'  18G5 
p.  68.  p.  275. 

*•  See  Prof.  Huxley  to  this  effect 


Chap.  VII.  The  Races  of  Man.  lyj 

from  a  coramon  stock,  tlie  differences  between  the  races  and 
their  number  must  have  been  small ;  consequently  as  far  as 
their  distinguishing  characters  are  concerned,  they  tht  n  had  less 
claim  to  rank  as  distinct  species  than  the  existing  so-called  races. 
Nevertheless,  so  arbitrary  is  the  term  of  species,  that  such  early 
races  would  perhaps  have  been  ranked  by  some  naturalists  as 
distinct  species,  if  their  differences,  although  extremely  slight, 
had  been  more  constant  than  they  are  at  present,  and  had  not 
graduated  into  each  other. 

It  is  however  possible,  though  far  from  probable,  that  the 
early  progenitors  of  man  might  formerly  have  diverged  much  in 
character,  until  they  became  more  unlike  each  other  than  any 
now  existing  races;  but  that  subsequently,  as  suggested  by 
Vogt,"  they  converged  in  character.  "When  man  selects  the  off- 
spring of  two  distinct  species  for  the  same  object,  he  sometimes 
induces  a  considerable  amount  of  convergence,  as  far  as  general 
appearance  is  concerned.  This  is  the  case,  as  shewn  by  Von 
Nathusius,-^  with  the  improved  breeds  of  the  pig,  which  are 
descended  from  two  distinct  species;  and  in  a  less  marked 
manner  with  the  improved  breeds  of  cattle.  A  great  anatomist, 
Gratiolet,  maintains  that  the  anthropomorphous  apes  do  not 
form  a  natural  sub-group ;  but  that  the  orang  is  a  highly 
developed  gibbon  or  semnopithecus,  the  chimpanzee  a  highly 
developed  macacus,  and  the  gorilla  a  highly  developed  mandrill. 
If  this  conclusion,  which  rests  almost  exclusively  on  brain - 
characters,  be  admitted,  w^e  should  have  a  case  of  convergence 
at  least  in  external  characters,  for  the  anthropomorphous  apes 
are  certainly  more  like  each  other  in  many  points,  than  they  are 
to  other  apes.  All  analogical  resemblances,  as  of  a  whale  to  a 
fish,  may  indeed  be  said  to  be  cases  of  convergence;  but  this 
term  has  never  been  applied  to  superficial  and  adaptive  resem- 
blances. It  would,  however,  be  extremely  rash  to  attril)ute  to 
convergence  close  similarity  of  character  in  many  points  of 
structure  amongst  the  modified  descendants  of  widely  distinct 
beings.  The  form  of  a  crystal  is  determined  solely  by  the  mole- 
cular forces,  and  it  is  not  surprising  that  dissimilar  substances 
should  sometimes  assume  the  same  form ;  but  with  organic 
beings  we  should  bear  in  mind  that  the  form  of  each  depends  on 
an  infinity  of  complex  relations,  namely  on  variations,  due  to 
causes  far  too  intricate  to  be  followed,— on  the  nature  of  the 
variations  preserved,  these  depending  on  the  physicial  condi- 

^-  '  Lectures  oa  ^lan,' Eng.  trans-  schichte,      &c.,      Schweineschildel,' 

lat.  186+,  p.  4fi8.  1864,  s.  104.    With  respect  to  cattle, 

23    '  Die    Racen    des    Schwcines,'  see  M.    de    Quatretages,    '  Unit<5  de 

I860,  s.  46.     'Voistudien   fiir  Ge-  I'Espece  Humaine,' 1861,  p.  119. 


178  The  Descent  of  Man,  Part  1. 

tions,  and  still  more  on  the  surrounding  organisms  which  com- 
pete with  each,— and  lastly,  on  inheritance  (in  itself  a  fluctuating 
element)  from  innumerable  progenitors,  all  of  which  have  had 
their  forms  determined  through  equally  complex  relations.  It 
appears  incredible  that  the  modified  descendants  of  two  organ- 
isms, if  these  differed  from  each  other  in  a  marked  manner, 
should  ever  afterwards  converge  so  closely  as  to  lead  to  a  near 
approach  to  identity  throughout  their  whole  organisation.  In 
the  case  of  the  convergent  races  of  pigs  above  referred  to,  evi- 
dence of  their  descent  from  two  primitive  stocks  is,  according  to 
Von  Nathusius,  still  plainly  retained,  in  certain  bones  of  their 
skulls.  If  the  races  of  man  had  descended,  as  is  supposed  by 
some  naturalists,  from  two  or  more  species,  which  differed  from 
each  other  as  much,  or  nearly  as  much,  as  does  the  orang  from 
the  gorilla,  it  can  hardly  be  doubted  that  marked  differences  in 
the  structure  of  certain  bones  would  still  be  discoverable  in  man 
as  he  now  exists. 

Although  the  existing  races  of  man  differ  in  many  respects,  as 
in  colour,  hair,  shape  of  skull,  proportions  of  the  body,  &c.,  yet 
if  their  whole  structure  be  taken  into  consideration  they  are 
found  to  resemble  each  other  closely  in  a  multitude  of  points. 
Many  of  these  are  of  so  unimportant  or  of  so  singular  a  nature, 
that  it  is  extremely  improbable  that  they  should  have  been  inde- 
pendently acquired  by  aboriginally  distinct  species  or  races. 
The  same  remark  holds  good  with  equal  or  greater  force  with 
respect  to  the  numerous  points  of  mental  similarity  between  the 
most  distinct  races  of  man.  The  American  aborigines,  Negroes 
and  Europeans  are  as  different  from  each  other  in  mind  as  any 
three  races  that  can  be  named;  yet  I  was  incessantly  struck, 
whilst  living  with  the  Fuegians  on  board  the  "  Beagle,"  with  the 
many  little  traits  of  character,  shewing  how  similar  their  minds 
were  to  ours ;  and  so  it  was  with  a  full-blooded  negro  with  whom 
I  happened  once  to  be  intimate. 

He  who  will  read  Mr.  Tylor's  and  Sir  J.  Lubbock's  interesting 
works  ^^  can  hardly  fail  to  be  deeply  impressed  with  the  close 
similarity  between  the  men  of  all  races  in  tastes,  dispositions  and 
habits.  This  is  shewn  by  the  pleasure  which  they  all  take  in 
dancing,  rude  music,  acting,  painting,  tattooing,  and  otherwise 
decorating  themselves ;  in  their  mutual  comprehension  of  gesture- 
language,  by  the  same  expression  in  their  features,  and  by  the 
same  inarticulate  cries,  when  excited  by  the  same  emotions. 
This  similarity,  or  rather  identity,  is  striking,  when  contrasted 

^*  Tylor's  '  Early  History  of  Man-  language,  see  p.  54.  Lubbock's 
kind,'  1865  :  with  respect  to  gesture-      '  Prehistoric  T'mes,'  2nd  edit.  1869. 


Chap.  VI  r.  TJie  Races  of  Man.  179 

with  the  different  expressions  and  erics  made  by  distinct  species 
of  monkeys.  There  is  good  evidence  that  tlie  art  of  sliooting 
with  bows  and  arrows  has  not  been  lianded  down  from  any 
common  progenitor  of  mankind,  yet  as  Westroi)p  and  Nilsson 
have  remarked/^  the  stone  arrow-heads,  brought  from  the  most 
distant  parts  of  the  world,  and  manufactured  at  the  most  remote 
periods,  are  almost  identical ;  and  this  fact  can  only  be  accounted 
for  by  the  various  races  having  similar  inventive  or  mental 
powers.  The  same  observation  has  been  made  by  archaeologists  ^^ 
with  respect  to  certain  widely-prevalent  ornaments,  such  as  zig- 
zags, &c. ;  and  with  respect  to  various  simple  beliefs  and  cus- 
toms, such  as  the  burying  of  the  dead  under  megalithic  struc- 
tures. I  remember  observing  in  South  America,^^  that  there,  as 
in  so  many  other  parts  of  the  world,  men  have  generally  chosen 
the  summits  of  lofty  hills,  to  throw  uj)  piles  of  stones,  either  as 
a  record  of  some  remarkable  event,  or  for  burying  their  dead. 

Now  when  naturalists  observe  a  close  agreement  in  numerous 
small  details  of  habits,  tastes,  and  dispositions  between  two  or 
more  domestic  races,  or  between  nearly-allied  natural  forms, 
they  use  this  fact  as  an  argument  that  they  are  descended  from  a 
common  progenitor  who  was  thus  endowed ;  and  consequently 
that  all  should  be  classed  under  the  same  species.  The  same 
argument  may  be  applied  with  much  force  to  the  races  of  man. 

As  it  is  improbable  that  the  numerous  and  unimportant  points 
of  resemblance  between  the  several  races  of  man  in  bodily  struc- 
ture and  mental  faculties  (I  do  not  here  refer  to  similar  customs) 
should  all  have  been  independently  acquired,  they  must  have  been 
inherited  from  progenitors  who  had  these  same  characters.  We 
thus  gain  some  insight  into  the  early  state  of  man,  before  he  had 
spread  step  by  step  over  the  face  of  the  earth.  The  spreading 
of  man  to  regions  widely  separated  by  the  sea,  no  doubt,  pre- 
ceded any  great  amount  of  divergence  of  character  in  the  several 
races ;  for  otherwise  we  should  sometimes  meet  with  the  same 
race  in  distinct  continents ;  and  this  is  never  the  case.  Sir  J. 
Lubbock,  after  comparing  the  arts  now  practised  by  savages  in 
all  parts  of  the  world,  sjDecifies  those  which  man  could  not  have 
known,  when  he  first  wandered  from  his  original  birth-place ; 
for  if  once  learnt  they  would  never  have  been  forgotten.'-**     He 

2^  'On  Analogous  Forms  of  Im-  'Journal    of   Ethnological    Soc'    as 

plenients,' in 'Memoirs  of  Anthropo-  given  in  *  Scientific   Opinion,'  June 

log.  Soc.,'  by  H.  M.  Westropp.    'The  2ud,  1869,  p.  3. 

Primitive    Inhabitants    of    Scandi-  ^^ 'Journal  of  Researches:  Voyage 

navia,'  Eng.  transFat.  edited  by  Sir  of  the  "Beagle,"'  p.  46. 

J.  Lubbock,  1868,  p.  104-.  "-*  'Prehistoric    Times,'    1809,    p. 

2«  Westropp, '  On  CDmlechs/ &c.,  574. 


l8o  The  Descent  of  Man.  Part  I. 

tlms  shews  that  "  the  spear,  which  is  but  a  development  of  the 
"  knife-point,  and  the  club,  which  is  but  a  long  hammer,  are  the 
"  only  things  left."  He  admits,  however,  that  the  art  of  making- 
lire  probably  had  been  already  discovered,  for  it  is  common  to 
all  the  races  now  existing,  and  was  known  to  the  ancient  cave- 
inhabitants  of  Europe.  Perhaps  the  art  of  making  rude  canoes 
or  rafts  was  likewise  known;  but  as  man  existed  at  a  remote 
epoch,  when  the  land  in  many  places  stood  at  a  very  different 
level  to  what  it  does  now,  he  would  have  been  able,  without  the 
aid  of  canoes,  to  have  spread  widely.  Sir  J.  Lubbock  further 
remarks  how  improbable  it  is  that  our  earliest  ancestors  could 
have  "  counted  as  high  as  ten,  considering  that  so  many  races 
"now  in  existence  cannot  get  beyond  four."  Nevertheless,  at 
this  early  p)eriod,  the  intellectual  and  social  faculties  of  man 
could  hardly  have  been  inferior  in  any  extreme  degree  to  those 
possessed  at  present  by  the  lowest  savages ;  otherwise  primeval 
man  could  not  have  been  so  eminently  successful  in  the  struggle 
for  life,  as  proved  by  his  early  and  wide  diffusion. 

From  the  fundamental  differences  between  certain  languages, 
some  philologists  have  inferred  that  when  man  first  became 
widely  diffused,  he  was  not  a  speaking  animal ;  but  it  may  be 
suspected  that  languages,  far  less  perfect  than  any  now  spoken, 
aided  by  gestures,  might  have  been  used,  and  yet  have  left  no 
traces  on  subsequent  and  more  highly-developed  tongues.  With- 
out the  use  of  some  language,  however  imperfect,  it  appears 
doubtful  whether  man's  intellect  could  have  risen  to  the 
standard  implied  by  his  dominant  position  at  an  early  period. 

"Whether  primeval  man,  when  he  possessed  but  few  arts,  and 
those  of  the  rudest  kind,  and  when  his  power  of  language  was 
extremely  imperfect,  would  have  deserved  to  be  called  man,  must 
depend  on  the  definition  wliich  we  employ.  In  a  series  of  forms 
graduating  insensibly  from  some  ape-like  creature  to  man  as  he 
now  exists,  it  would  be  impossible  to  fix  on  any  definite  point  when 
the  term  "  man  "  ought  to  be  used.  But  this  is  a  matter  of  very 
little  importance.  So  again,  it  is  almost  a  matter  of  indifference 
whether  the  so-called  races  of  man  are  thus  designated,  or  are 
ranked  as  species  or  sub-species  ;  but  the  latter  term  apj^ears  the 
more  appropriate.  Finally,  we  may  conclude  that  when  the 
principle  of  evolution  is  generally  accepted,  as  it  surely  will  be 
l3efore  long,  the  dispute  between  the  monogenists  and  the  poly- 
genists  will  die  a  silent  and  unobserved  death. 

One  other  question  ought  not  to  be  passed  over  without  notice, 
namely,  whether,  as  is  sometimes  assumed,  each  sub-sj)ecies  or 
race  of  man  has  sprung  from  a  single  pair  of  progenitors.     With 


Chap.  VII.  TJie  Extinction  of  Races.  1 8 1 


oui*  domestic  animals  a  new  race  can  readily  be  formed  by  care- 
fully matching  the  varying  ofiFspring  from  a  single  pair,  or  even 
from  a  single  individual  possessing  some  new  character;  but 
most  of  our  races  have  been  formed,  not  intentionally  from  a 
selected  pair,  but  unconsciously  by  the  preservation  of  many  in- 
dividuals which  have  varied,  however  slightly,  in  some  useful  or 
desired  manner.  If  in  one  country  stronger  and  heavier  horses, 
and  in  another  country  lighter  and  fleeter  ones,  were  habitually 
preferred,  we  may  feel  sure  that  two  distinct  sub-breeds  would 
be  produced  in  the  course  of  time,  without  any  one  pair  having 
been  separated  and  bred  from,  in  either  country.  Many  races 
have  been  thus  formed,  and  their  manner  of  formation  is  closely 
analogous  to  that  of  natural  species.  We  know,  also,  that  the 
horses  taken  to  the  Falkland  Islands  have,  during  successive 
generations,  become  smaller  and  weaker,  whilst  those  which  have 
run  wild  on  the  Pampas  have  acquired  larger  and  coarser  heads ; 
and  such  changes  are  manifestly  due,  not  to  any  one  pair,  but  to 
all  the  individuals  having  been  subjected  to  the  same  conditions, 
aided,  perhaps,  by  the  principle  of  reversion.  The  new  sub- 
breeds  in  such  cases  are  not  descended  from  any  single  pair,  but 
from  many  individuals  which  have  varied  in  different  degrees, 
but  in  the  same  general  manner ;  and  we  may  conclude  that  the 
races  of  man  have  been  similarly  produced,  the  modifications 
being  either  the  direct  result  of  exposure  to  different  conditions, 
or  the  indirect  result  of  some  form  of  selection.  But  to  this 
latter  subject  we  shall  presently  return. 

On  the  Extinction  of  the  Eacesof  Man. — The  partial  or  complete 
extinction  of  many  races  and  sub-races  of  man  is  historically 
known.  Humboldt  saw  in  South  America  a  parrot  which  was 
the  sole  living  creature  that  could  speak  a  word  of  the  language 
of  a  lost  tribe.  Ancient  monuments  and  stone  implements 
found  in  all  parts  of  the  world,  about  which  no  tradition  has  been 
])reserved  by  the  present  inhabitants,  indicate  much  extinction, 
Some  small  and  broken  tribes,  remnants  of  former  races,  still 
survive  in  isolated  and  generally  mountainous  districts.  In 
Europe  the  ancient  races  were  all,  according  to  Schaaffhausen,-" 
"  lower  in  the  scale  than  the  rudest  living  savages ;"  they  must 
therefore  have  differed,  to  a  certain  extent,  from  any  existing 
race.  The  remains  described  by  Professor  Broca  from  Les  Eyzies, 
though  they  unfortunately  appear  to  have  belonged  to  a  single 
family,  indicate  a  race  with  a  most  singular  combination  of  low 
or  simious,  and  of  high  characteristics.     This  race  is  "  entirely' 

29  Translation  in  'Anthropological  Review,'  Oct.  1868,  p.  431 


1 82  The  Descent  of  Man.  Part  I. 


"  different  from  any  other,  ancient  or  modern,  that  we  have  ever 
"heard  of."^°  It  differed,  therefore,  from  the  quaternary  race  of 
the  caverns  of  Belgium. 

Man  can  long  resist  conditions  which  appear  extremely  un- 
favourable for  his  existence.^^  He  has  long  lived  in  the  extreme 
regions  of  the  North,  with  no  wood  for  his  canoes  or  implements, 
and  with  only  blubber  as  fuel,  and  melted  snow  as  drink.  In 
the  southern  extremity  of  America  the  Fuegians  survive  with- 
out the  protection  of  clothes,  or  of  any  building  worthy  to  be 
called  a  hovel.  In  South  Africa  the  aborigines  wander  over  arid 
plains,  where  dangerous  beasts  abound.  Man  can  withstand  the 
deadly  influence  of  the  Terai  at  the  foot  of  the  Himalaya,  and 
the  pestilential  shores  of  tropical  Africa. 

Extinction  follows  chiefly  from  the  competition  of  tribe  with 
tribe,  and  race  with  race.  Various  checks  are  always  in  action, 
serving  to  keep  down  the  numbers  of  each  savage  tribe, — such 
as  periodical  famines,  nomadic  habits  and  the  consequent  deaths 
of  infants,  prolonged  suckling,  wars,  accidents,  sickness,  licen- 
tiousness, the  stealing  of  women,  infanticide,  and  especially 
lessened  fertility.  If  any  one  of  these  checks  increases  in  power, 
even  slightly,  the  tribe  thus  aft'ected  tends  to  decrease ;  and 
when  of  two  adjoining  tribes  one  becomes  less  numerous  and  less 
powerful  than  the  other,  the  contest  is  soon  settled  by  war, 
slaughter,  cannibalism,  slavery,  and  absorption.  Even  when  a 
weaker  tribe  is  not  thus  abruptly  swept  away,  if  it  once  begins 
to  decrease,  it  generally  goes  on  decreasing  until  it  becomes 
extinct.^^ 

When  civilised  nations  come  into  contact  with  barbarians  the 
struggle  is  short,  except  where  a  deadly  climate  gives  its  aid  to 
the  native  race.  Of  the  causes  which  lead  to  the  victory  of 
civilised  nations,  some  are  plain  and  simple,  others  complex  and 
obscure.  We  can  see  that  the  cultivation  of  the  land  will 
be  fatal  in  many  ways  to  savages,  for  they  cannot,  or  will  not, 
change  their  habits.  New  diseases  and  vices  have  in  some  cases 
proved  highly  destructive;  and  it  appears  that  a  new  disease 
often  causes  much  death,  until  those  who  are  most  susceptible 
to  its  destructive  influence  are  gradually  weeded  out  f^  and  so  it 
may  be  with  the  evil  effects  from  spirituous  liquors,  as  well  as 
with  the  unconquerably  strong  taste  for  them  shewn  by  so  many 

^^  'Transact,   Internat.   Congress  terben  der  NatuvA^olker,' 1868,  s.  82. 

of  Prehistoric  Arch,'  1868,  pp.  172-  ^^  Gerland  (ibid.  s.  12)  gives  facts 

•  175.      See  also  Broca  (translation)  in  support  of  this  statement. 

in   '  Anthropological    Review,'   Oct.  ^^  See   remarks  to   this  effect  in 

1868,  p.  410.  Sir  H.  Holland's  '  Medical  Notes  and 

31  Dr.  Gerland  '  Ueber  das  Auss-  Reflections,'  1839,  p.  390. 


Chap.  VI [.  The  Extinction  of  Races.  1S3 


savages.  It  further  appears,  mysterious  as  is  the  fact,  that 
the  first  meeting  of  distinct  and  separated  people  generates 
disease.^^  Mr.  Sproat,  wlio  in  Vancouver  Island  closely  attended 
to  the  subject  of  extinction,  believed  that  changed  habits  of  life, 
consequent  on  the  advent  of  Europeans,  induces  much  ill  health. 
He  lays,  also,  great  stress  on  the  apparently  trifling  cause  that 
the  natives  become  ''  bewildered  and  dull  by  the  new  life  around 
"  them ;  they  lose  the  motives  for  exertion,  and  get  no  new  ones 
"  in  their  place."  ^^ 

The  grade  of  their  civilisation  seems  to  be  a  most  important 
element  in  the  success  of  competing  nations.  A  few  centuries 
ago  Europe  feared  the  inroads  of  Eastern  barbarians ;  now  any 
such  fear  would  be  ridiculous.  It  is  a  more  curious  fact,  as 
Mr.  Bagehot  has  remarked,  that  savages  did  not  formerly  waste 
away  before  the  classical  nations,  as  they  now  do  before  modern 
civilised  nations;  had  they  done  so,  the  old  moralists  would 
have  mused  over  the  event ;  but  there  is  no  lament  in  any  writer 
of  that  period  over  the  perishing  barbarians.^^  The  most  po- 
tent of  all  the  causes  of  extinction,  appears  in  many  cases  to  be 
lessened  fertility  and  ill-health,  especially  amongst  the  children, 
arising  from  changed  conditions  of  life,  notwithstanding  that  the 
new  conditions  may  not  be  injurious  in  themselves.  I  am  much 
indebted  to  Mr.  H.  H.  Howorth  for  having  called  my  attention  to 
this  suliject,  and  for  having  given  me  information  respecting  it. 
I  have  collected  the  following  cases. 

When  Tasmania  was  first  colonised  the  natives  were  roughly 
estimated  by  some  at  7000  and  by  others  at  20,000.  Their 
number  was  soon  greatly  reduced,  chiefly  by  fighting  with  the 
English  and  with  each  other.  After  the  famous  hunt  by  all  the 
colonists,  when  the  remaining  natives  delivered  themselves  up 
to  the  government,  they  consisted  only  of  120  individuals,^'^ 
who  were  in  18o2  transported  to  Flinders  Island.  This  island, 
situated  between  Tasmania  and  Australia,  is  forty  miles  long, 
and  from  twelve  to  eighteen  miles  broad:  it  seems  healthy, 
and  the  natives  were  well  treated.  Nevertheless,  they  suffered 
greatly  in  health.  In  1834  they  consisted  (Bon wick,  p.  250)  of 
forty-seven  adult  males,  forty-eight  adult  females,  and  sixteen 
children,  or  in  all  of  111  souls.   In  1835  only  one  hundred  were  left. 

"  I  have   collected  ('Journal  of  Savage  Life,'  18G8,  p.  284-. 
Researches,  Voyage  of  the  "Beagle,"'  ^'^  Bagehot,    '  I'hysics    and    Poli- 

p.  435)  a  good  many  cases  bearing  tics,'    '  Fortnightly    Review,'   April 

on   this  subject:    see  also  Geriand,  1,  18<J8,  p.  455. 
ibid.  s.  8.     Poepidg    speaks    of  the  ^^  All  the  statements  here  given 

"  breath  of  civilisation  as  poisonous  are    taken   from    'The    last    of  the 

"to  savages."  Tasmanians,'  by  J.  Bonwick,  1870. 

'•^  Sproat,  '  Scenes  and  Studies  ot 


1 84  The  Descent  of  Man.  Part  I. 

As  tliey  continued  rapidly  to  decrease,  and  as  they  themselves 
thought  that  they  should  not  perish  so  quickly  elsewhere,  they  were 
removed  in  1847  to  Oyster  Cove  in  the  southern  part  of  Tasmania. 
They  then  consisted  (Dec.  20th,  1847)  of  fourteen  men,  twenty- 
two  women  and  ten  children.^^  But  the  change  of  site  did  no  good. 
Disease  and  death  still  pursued  them,  and  in  1864  one  man  (who 
died  in  1869),  and  three  elderly  women  alone  survived.  The 
infertility  of  the  women  is  even  a  more  remarkable  fact  than 
the  liability  of  all  to  ill-health  and  death.  At  the  time  when 
only  nine  women  were  left  at  Oyster  Cove,  they  told  Mr.  Bonwick 
(p.  386),  that  only  two  had  ever  borne  children  :  and  these  two 
had  together  produced  only  three  children ! 

With  resj^ect  to  the  cause  of  this  extraordinary  state  of  things. 
Dr.  Story  remarks  that  death  followed  the  attemj)ts  to  civilise 
the  natives.  "  If  left  to  themselves  to  roam  as  they  were  wont 
"  and  undisturbed,  they  would  have  reared  more  children,  and 
''  there  would  have  been  less  mortality."  Another  careful 
observer  of  the  natives,  Mr.  Davis,  remarks,  "  The  births  have 
"  been  few  and  the  deaths  numerous.  This  may  have  been  in  a 
''  great  measure  owing  to  their  change  of  living  and  food ;  but 
"  more  so  to  their  banishment  from  the  mainland  of  Van  Diemen's 
"Land,  and  consequent  depression  of  spirits"  (Bonwick,  jDp. 
388,  390). 

Similar  facts  have  been  observed  in  two  widely  different 
parts  of  Australia.  The  celebrated  explorer,  Mr.  Gregory,  told 
Mr.  Bonwick,  that  in  Queensland  "the  want  of  reproduction 
"was  being  already  felt  with  the  blacks,  even  in  the  most 
"  recently  settled  parts,  and  that  decay  would  set  in."  Of 
thirteen  aborigines  from  Shark's  Bay  who  visited  Murchison 
Eiver,  twelve  died  of  consumption  within  three  months.^^ 

The  decrease  of  the  Maories  of  New  Zealand  has  been  carefully 
investigated  by  Mr.  Teuton,  in  an  admirable  Eeport,  from  which 
all  the  following  statements,  with  one  exception,  are  taken."*" 
The  decrease  in  number  since  1830  is  admitted  by  every  one, 
including  the  natives  themselves,  and  is  still  steadily  progress- 
ing. Although  it  has  hitherto  been  found  impossible  to  take  an 
actual  census  of  the  natives,  their  numbers  were  carefully 
estimated  by  residents  in  many  districts.  The  result  seems 
trustworthy,  and  shows  that  during  the  fourteen  years,  previous 

38  This   is   the  statement  of  the  1870,  p.  90;  and  the  'Last  of  the 

Governor  of  Tasmania,  Sir  W.  Deni-  Tasmanians,'  1870,  p.  386. 
.son,  '  Varieties  of  Vice-Regal  Life,'  ^^  'Observations  on  the  Aboriginal 

1870,  vol.  i.  p.  67.  Inhabitants  of  New  Zealand,'  pub- 

3^  For  these  cases,  see  Bonwick 's  lished  by  the  Government,  1859. 
*  Daily    Life    of    the    Tasmanians,' 


Chap.  VII.  The  Extinction  of  Races.  185 

to  1858,  the  decrease  was  19.42  per  cent.  Some  of  the  tribes, 
thus  carefully  examined,  lived  above  a  Inmdrcd  miles  apart, 
some  on  the  coast,  some  inland ;  and  their  means  of  subsistence 
and  habits  differed  to  a  certain  extent  (p.  28).  The  total 
number  in  1858  was  believed  to  be  53,700,  and  in  1872,  after  a 
second  interval  of  fourteen  years,  another  census  was  taken, 
and  the  number  is  given  as  only  36,359,  shewing  a  decrease  of 
32-29  per  cent. !  *^  Mr.  Fenton,  after  shewing  in  detail  the  in- 
sufficiency of  the  various  causes,  usually  assigned  in  explana- 
tion of  this  extraordinary  decrease,  such  as  new  diseases,  the 
profligacy  of  the  women,  drunkenness,  wars,  &c.,  concludes  on 
weighty  grounds  that  it  depends  chiefly  on  the  unproductiveness 
of  the  women,  and  on  the  extraordinary  mortality  of  the  young 
children  (pp.  31,  34).  In  proof  of  this  he  shews  (p.  33)  that  in 
1844  there  was  one  non-adult  for  every  2*57  adults;  whereas  in 
1858  there  was  only  one  non-adult  for  every  3-27  adults.  The 
mortality  of  the  adults  is  also  great.  He  adduces  as  a  further 
cause  of  the  decrease  the  inequality  of  the  sexes ;  for  fewer  females 
are  born  than  males.  To  this  latter  point,  depending  perhaps 
on  a  widely  distinct  cause,  I  shall  return  in  a  future  chapter. 
Mr.  Fenton  contrasts  with  astonishment  the  decrease  in  New 
Zealand  with  the  increase  in  Ireland ;  countries  not  very  dis- 
similar in  climate,  and  where  the  inhabitants  now  follow  nearly 
similar  habits.  The  Maories  themselves  (p.  35)  "  attribute  their 
"decadence,  in  some  measure,  to  the  introduction  of  new  food 
"■  and  clothing,  and  the  attendant  change  of  habits;"  and  it  will 
be  seen,  when  we  consider  the  influence  of  changed  conditions 
on  fertility,  that  they  are  probably  right.  The  diminution  began 
between  the  years  1880  and  1840 ;  and  Mr.  Fenton  shews  (p.  40) 
that  about  1830,  the  art  of  manufacturing  putrid  corn  (maize), 
by  long  steeping  in  water,  was  discovered  and  largely  practised ; 
and  this  proves  that  a  change  of  habits  was  beginning  amongst 
the  natives,  even  when  New  Zealand  was  only  thinly  inhabited 
by  Europeans.  When  I  visited  the  Bay  of  Islands  in  1835, 
the  dress  and  food  of  the  inhabitants  had  already  been  much 
modified :  they  raised  potatoes,  maize,  and  other  agricultural 
produce,  and  exchanged  them  for  English  manufactured  goods 
and  tobacco. 

It  is  evident  from  many  statements  in  the  life  of  Bishop 
Patteson,^^  that  the  Melanesians  of  the  New  Hebrides  and 
neighbouring  archipelagoes,  suffered  to  an  extraordinary  degree 
in  health,  and    perished  in  large  numbers,  when  they  were 

*'  'New  Zealand,'  by  Alex.  Ken-  C.  M.  Younge,  1874;  see  more 
nedy,  1873,  p.  47.  especially  vol.  i.  p.  530. 

*2  « Life   of  J.    C.    Patteson,'   by 


i86 


The  Descent  of  Man. 


Part  I. 


removed  to  New  Zealand,  Norfolk  Island,  and  other  salubrious 
places,  in  order  to  be  educated  as  missionaries. 

The  decrease  of  the  native  population  of  the  Sandwich  Islands 
is  as  notorious  as  that  of  New  Zealand.  It  has  been  roughly 
estimated  by  those  best  capable  of  judging,  that  when  Cook 
discovered  the  Islands  in  1779,  the  population  amounted  to 
about  300,000.  According  to  a  loose  census  in  1828,  the 
numbers  then  were  142,050.  In  1832,  and  at  several  subsequent 
periods,  an  accurate  census  was  officially  taken,  but  I  have 
been  able  to  obtain  only  the  following  returns  : 


Annual  rate  of  decrease 

Native  ropuLATiON. 

per    cent.,  assuming    it 

to  have  been  uniform  be- 

Year. 

(Except    during    1832 

tween  the  successive  cen- 

and   1836,  when   the 

suses  ;  these  censuses  be- 

few foreigners  in  the 

ing    taken    at  irregular 

islands  were  included). 

intervals. 

1832 

130,313^ 

4-46 

1836 

108,579| 
71,019. 

2-47 

1853 

. 

0-81 

1860 

67,084: 
58,765| 
51,531^ 

2-18 

1866 

2-17 

1872 

We  here  see  that  in  the  interval  of  forty  years,  between  1832  and 
1872,  the  population  has  decreased  no  less  than  sixty-eight  per 
cent. !  This  has  been  attributed  by  most  writers  to  the  profligacy 
of  the  women,  to  former  bloody  wars,  and  to  the  severe  labour 
imposed  on  conquered  tribes  and  to  newly  introduced  diseases, 
which  have  been  on  several  occasions  extremely  destructive.  No 
doubt  these  and  other  such  causes  have  been  highly  efficient, 
and  may  account  for  the  extraordinary  rate  of  decrease  between 
the  years  1832  and  1836  ;  but  the  most  potent  of  all  the  causes 
seems  to  be  lessened  fertility.  According  to  Dr.  Euschenberger 
of  the  U.S.  Navy,  who  visited  these  islands  between  1835  and 
1837,  in  one  district  of  Hawaii,  only  twenty-five  men  out  of  1134, 
and  in  another  district  only  ten  out  of  637,  had  a  family  with  as 
many  as  three  children.  Of  eighty  married  women,  only  thirty- 
nine  had  ever  borne  children  ;  and  "  the  official  report  gives  an 
"  average  of  half  a  cliild  to  each  married  couple  in  the  whole 


Chap.  VII.  The  Extinction  of  Races.  187 

"  island."  This  is  almost  exactly  the  same  average  as  with  the 
Tasmanians  at  Oyster  Cove.  Jarves,  who  published  his  History 
in  1843,  says  that "  families  who  have  three  children  are  freed  from 
"  all  taxes  ;  those  having  more,  arc  rewarded  by  gifts  of  land  and 
*' other  encouragements."  This  unparalleled  enactment  by  the 
government  well  shews  how  infertile  the  race  had  become.  The 
Eev.  A.  Bishop  stated  in  the  Hawaiian  '  Spectator'  in  1839,  that  a 
large  proportion  of  the  children  die  at  early  ages,  and  Bishop 
Staley  informs  me  that  this  is  still  the  case,  just  as  in  New 
Zealand.  This  has  been  attributed  to  the  neglect  of  the  children 
by  the  women,  but  it  is  probably  in  large  part  due  to  innate  weak- 
ness of  constitution  in  the  children,  in  relation  to  the  lessened 
fertility  of  their  parents.  There  is,  moreover,  a  further  resem- 
blance to  the  case  of  New  Zealand,  in  the  fact  that  there  is  a 
large  excess  of  male  over  female  births :  the  census  of  1872 
gives  31,650  males  to  25,247  females  of  all  ages,  that  is  125-36 
males  for  every  100  females;  whereas  in  all  civilised  countries 
the  females  exceed  the  males.  No  doubt  the  profligacy  of  the 
women  may  in  part  account  for  their  small  fertility ;  but  their 
changed  habits  of  life  is  a  much  more  probable  cause,  and  which 
will  at  the  same  time  account  for  the  increased  mortality, 
especially  of  the  children.  The  islands  were  visited  by  Cook  in 
1779,  by  Vancouver  in  1794,  and  often  subsequently  by  w^halers. 
In  1819  missionaries  arrived,  and  found  that  idolatry  had  been 
already  abolished,  and  other  changes  effected  by  the  king.  After 
this  period  there  was  a  rapid  change  in  almost  all  the  habits  of 
life  of  the  natives,  and  they  soon  became  "  the  most  civilised  of 
"  the  Pacific  Islanders."  One  of  my  informants,  Mr.  Coan,  who 
was  born  on  the  islands,  remarks  that  the  natives  have  undergone 
a  greater  change  in  their  habits  of  life  in  the  course  of  fifty  years 
than  Englishman  during  a  thousand  years.  From  information 
received  from  Bishop  Staley,  it  does  not  appear  that  the 
poorer  classes  have  ever  much  changed  their  diet,  although 
many  new  kinds  of  fruit  have  been  introduced,  and  the  sugar- 
cane is  in  universal  use.  Owing,  however,  to  their  passion  for 
imitating  Europeans,  they  altered  their  manner  of  dressing  at 
an  early  period,  and  the  use  of  alcoholic  drinks  became  very 
general.  Although  these  changes  appear  inconsiderable,  I  can 
well  believe,  from  what  is  known  with  respect  to  animals,  that 
tliey  might  suffice  to  lessen  the  fertility  of  the  natives."*^ 

*'  The  foregoing   statements  are  Islands,'    1851,  p.    277.      Ruschen- 

taken    chiefly    from    the    following  berger  is  quoted  by  Bonwick,  'Last 

works:     'Jarves'    Histoi-y    of    the  of  the   Tasmanians,'   1870,   p.    378. 

Hawaiian  Islauds,'  1843,  p.  400-407.  Bishop  is  quoted  by  Sir  E.  Belcher, 

Cheever,    '  Life    in    the    Sandwich  '  Voyage   Round   the  World,'  1843, 


1 88  The  Descent  of  Man.  Part  1. 

Lastly,  Mr.  Macnamara  states**  tliat  the  low  and  degraded 
inhabitants  of  the  Andaman  Islands,  on  the  eastern  side  of  the 
Gulf  of  Bengal,  are  "  eminently  susceptible  to  any  change  of 
"  climate  :  in  fact,  take  them  away  from  their  island  homes,  and 
"  they  are  almost  certain  to  die,  and  that  independently  of  diet 
"  or  extraneous  influences."  Ee  further  states  that  the  inhabit- 
ants of  the  Valley  of  Nepal,  which  is  extremely  hot  in  summer, 
and  also  the  various  hill-tribes  of  India,  suffer  from  dysentery 
and  fever  when  on  the  plains ;  and  they  die  if  they  attempt  to 
pass  the  whole  year  there. 

We  thus  see  that  many  of  the  wilder  races  of  man  are  apt  to 
suffer  much  in  health  when  subjected  to  changed  conditions 
or  habits  of  life,  and  not  exclusively  from  being  transported  to 
a  new  climate.  Mere  alterations  in  habits,  which  do  not  appear 
injurious  in  themselves,  seem  to  have  this  same  effect ;  and  in 
several  cases  the  children  are  particularly  liable  to  suffer.  It 
has  often  been  said,  as  Mr.  Macnamara  remarks,  that  man  can 
resist  with  impunity  the  greatest  diversities  of  climate  and  other 
changes;  but  this  is  true  only  of  the  civilised  races.  Man  in 
his  wild  condition  seems  to  be  in  this  respect  almost  as  sus- 
ceptible as  his  nearest  allies,  the  anthropoid  apes,  which  have 
never  yet  survived  long,  when  removed  from  their  native 
country. 

Lessened  fertility  from  changed  conditions,  as  in  the  case  of  the 
Tasmanians,  Maories,  Sandwich  Islanders,  and  apparently  the 
Australians,  is  still  more  interesting  than  their  liability  to 
ill-health  and  death;  for  even  a  slight  degree  of  infertility, 
combined  with  those  other  causes  which  tend  to  check  the 
increase  of  every  population,  would  sooner  or  later  lead  to 
extinction.  The  diminution  of  fertility  may  be  explained  in 
some  cases  by  the  j)rofligacy  of  the  women  (as  until  lately  with 
the  Tahitians),  but  Mr.  Fenton  has  shewn  that  this  explanation 
by  no  means  suffices  with  the  New  Zealanders,  nor  does  it  with 
the  Tasmanians. 

In  the  paper  above  quoted,  Mr.  Macnamara  gives  reasons  for 
believing  that  the  inhabitants  of  districts  subject  to  malaria  arc 
apt  to  be  sterile;  but  this  cannot  apply  in  several  of  the  abovo 
cases.  Some  writers  have  suggested  that  the  aborigines  of 
islands  have  suffered  in  fertility  and  health  from  long  continued 


vol,  i.,  p.  272.     I  owe  the  census  of  of  the  above-named  works.     I  have 

the  several  years  to  the  kindness  of  omitted   the   census  for  1850,  as  I 

Mr.  Coan,  at  the  request  of  Dr.  You-  have  seen  two  widely  different  num- 

mans  of  New   York  ;  and   in   most  bers  given. 

cases  I  have  compared  the  Youmans  ■**  '  The  Indian  Medical  Gazette, 

iigures  with  those  given  in  several  Nov.  1,  1871,  p.  240. 


Chap.  VII.  The  Extinction  of  Races,  1 89 


inter-breeding ;  but  in  the  above  cases  infertility  has  coincided 
too  closely  with  the  arrival  of  Europeans  for  us  to  admit  this 
explanation.  Nor  have  we  at  present  any  reason  to  believe 
that  man  is  highly  sensitive  to  the  evil  effects  of  inter-breeding, 
especially  in  areas  so  large  as  New  Zealand,  and  the  Sandwich 
archipelago  with  its  diversified  stations.  On  the  contrary,  it  is 
known  that  the  present  inhabitants  of  Norfolk  Island  are  nearly 
all  cousins  or  near  relations,  as  are  the  Todas  in  India,  and  the 
inhabitants  of  some  of  the  Western  Islands  of  Scotland;  and 
yet  they  seem  not  to  have  suffered  in  fertility.'*'' 

A  much  more  probable  view  is  suggested  by  the  analogy  of 
the  lower  animals.  The  reproductive  system  can  be  shewn  to  be 
susceptible  to  an  extraordinary  degree  (though  why  we  know 
not)  to  changed  conditions  of  life ;  and  this  susceptibility  leads 
both  to  beneficial  and  to  evil  results.  A  large  collection  of  facts 
on  this  subject  is  given  in  chap,  xviii.  of  vol.  ii.  of  my  '  Variation 
of  Animals  and  Plants  under  Domestication/  I  can  here  give  only 
the  briefest  abstract;  and  every  one  interested  in  the  subject 
may  consult  the  above  work.  Very  slight  changes  increase  the 
health,  vigour  and  fertility  of  most  or  all  organic  beings, 
whilst  other  changes  are  known  to  render  a  large  number  of 
animals  sterile.  One  of  the  most  familiar  cases,  is  that  of  tamed 
elephants  not  breeding  in  India;  though  they  often  breed  in 
Ava,  where  the  females  are  allowed  to  roam  about  the  forests  to 
some  extent,  and  are  thus  placed  under  more  natural  conditions. 
The  case  of  various  American  monkeys,  both  sexes  of  which 
have  been  kept  for  many  years  together  in  their  own  countries, 
and  yet  have  very  rarely  or  never  bred,  is  a  more  apposite  in- 
stance, because  of  their  relationship  to  man.  It  is  remarkable 
how  slight  a  change  in  the  conditions  often  induces  sterility  in  a 
wild  animal  when  captured  ;  and  this  is  the  more  strange  as  all 
our  domesticated  animals  have  become  more  fertile  than  they 
were  in  a  state  of  nature;  and  some  of  them  can  resist  the 
most  unnatural  conditions  with  undiminished  fertility .^^  Certain 
groups  of  animals  are  much  more  liable  than  others  to  be 
affected  by  captivity ;  and  generally  all  the  species  of  the  same 
group  are  affected  in  the  same  manner.  But  sometimes  a  single 
species  in  a  grouiD  is  rendered  sterile,  whilst  the  others  are  not 
so ;  on  the  other  hand,  a  single  species  may  retain  its  fertility 

••^  On  the  close  relationship  of  the  Scotland,  Dr.  Mitchell,  'Edinburgh 

Norfolk  Islanders,  see  Sir  W.  Deni-  Medical   Journal,'  March    to  June, 

son,  'Varieties  of  Vice-Kegal  Life,'  1865. 

vol.  i.  1870,  p.  410.    For  the  Todas,  ^°  For  the  evidence  on  this  head, 

see  Col.  Marshall's  work,  1873,  p.  see    'Variation    of    Animals'    &c., 

110.     For    the   Western   Ishinds    of  vol.  ii.  p.  111. 


1 90  The  Descent  of  Man.  Part  1. 

whilst  most  of  the  others  fail  to  breed.  The  males  and  females 
of  some  species  when  confined,  or  when  allowed  to  live  almost, 
but  not  quite  free,  in  their  native  country,  never  unite ;  others 
thus  circumstanced  frequently  unite  but  never  produce  offspring ; 
others  again  produce  some  offspring,  but  fewer  than  in  a  state 
of  nature ;  and  as  bearing  on  the  above  cases  of  man,  it  is 
important  to  remark  that  the  young  are  apt  to  be  weak  and 
sickly,  or  malformed,  and  to  jDerish  at  an  early  age. 

Seeing  how  general  is  this  law  of  the  susceptibility  of  the 
reproductive  system  to  changed  conditions  of  life,  and  that  it 
holds  good  with  our  nearest  allies,  the  Quadrimaana,  I  can 
hardly  doubt  that  it  applies  to  man  in  his  primeval  state.  Hence 
if  savages  of  any  race  are  induced  suddenly  to  change  their 
habits  of  hfe,  they  become  more  or  less  sterile,  and  their  young 
offspring  suffer  in  health,  in  the  same  manner  and  from  the  same 
cause,  as  do  the  elephant  and  hunting-leopard  in  India,  many 
monkeys  in  America,  and  a  host  of  animals  of  all  kinds,  on  removal 
from  their  natural  conditions. 

We  can  see  why  it  is  that  aborigines,  who  have  long  inha- 
bited islands,  and  who  must  have  been  long  exposed  to  nearly 
uniform  conditions,  should  be  specially  affected  by  any  change 
in  their  habits,  as  seems  to  be  the  case.  Civilised  races  can 
certainly  resist  changes  of  all  kinds  far  better  than  savages; 
and  in  this  respect  they  resemble  domesticated  animals,  for 
though  the  latter  sometimes  suffer  in  health  (for  instance 
European  dogs  in  India),  yet  they  are  rarely  rendered  sterile, 
though  a  few  such  instances  have  been  recorded.*^  The 
immunity  of  civilised  races  and  domesticated  animals  is 
probably  due  to  their  having  been  subjected  to  a  greater  extent, 
and  therefore  having  grown  somewhat  more  accustomed,  to 
diversified  or  varying  conditions,  than  the  majority  of  wild 
animals ;  and  to  their  having  formerly  immigrated  or  been 
carried  from  country  to  country,  and  to  different  families  or 
sub-races  having  inter-crossed.  It  appears  that  a  cross  with 
civilised  races  at  once  gives  to  an  aboriginal  race  an  immunity 
from  the  evil  consequences  of  changed  conditions.  Thus  the 
crossed  offspring  from  the  Tahitians  and  English,  when  settled 
in  Pitcairn  Island,  increased  so  rapidly  that  the  island  was  soon 
overstocked ;  and  in  June  1856  they  were  removed  to  Norfolk 
Island.  They  then  consisted  of  60  married  persons  and  134 
children,  making  a  total  of  194.  Here  they  likewise  in- 
creased so  rapidly,  that  although  sixteen  of  them  returned  to  Pit- 
cairn Island  in  1859,  they  numbered  in  January  1868,  300  souls ; 

*^  '  Variation  of  Animals,'  &c.,  vol.  ii.,  p  16, 


Chap.  VII.  The  Extinction  of  Races.  191 


the  males  and  females  being  in  exactly  equal  numbers.  What  a 
contrast  does  tliis  case  present  with  that  of  the  Tasmanians; 
the  Norfolk  Islanders  increased  in  only  twelve  and  a  lialf  years 
from  194  to  300;  whereas  the  Tasmanians  decreased  during 
fifteen  years  from  120  to  46,  of  which  latter  number  only  ten 
were  children.^^ 

So  again  in  the  interval  between  the  census  of  18G6  and  1872 
the  natives  of  full  blood  in  the  Sandwich  Islands  decreased  by 
8081,  whilst  the  half-castes,  who  are  believed  to  be  healthier,  in- 
creased by  847 ;  but  I  do  not  know  whether  the  latter  number 
includes  tlie  offspring  from  the  half-castes,  or  only  the  half-castes 
of  the  first  generation. 

The  cases  which  I  have  here  given  all  relate  to  aborigines, 
wlio  have  been  subjected  to  new  conditions  as  the  result  of  the 
immigration  of  civilised  men.  But  sterility  and  ill-health  would 
probably  follow,  if  savages  were  compelled  by  any  cause,  such 
as  the  inroad  of  a  conquering  tribe,  to  desert  their  homes  and 
to  change  their  habits.  It  is  an  interesting  circumstance  that 
the  chief  check  to  wild  animals  becoming  domesticated,  which 
implies  the  power  of  their  breeding  freely  when  first  cai)tured, 
and  one  chief  check  to  wild  men,  when  brought  into  contact 
with  civilisation,  surviving  to  form  a  civilised  race,  is  the  same, 
namely,  sterility  from  changed  conditions  of  life. 

Finally,  although  the  gradual  decrease  and  ultimate  extinction 
of  the  races  of  man  is  a  highly  complex  problem,  depending  on 
many  causes  which  differ  in  different  places  and  at  different 
times ;  it  is  the  same  problem  as  that  presented  by  the  extinc- 
tion of  one  of  the  higher  animals — of  the  fossil  horse,  for  in- 
stance, which  disappeared  from  South  America,  soon  afterwards 
to  be  replaced,  within  the  same  districts,  by  countless  troops 
of  the  Spanish  horse.  The  New  Zealander  seems  conscious  of 
this  parallelism,  for  he  compares  his  future  fate  with  that  of 
the  native  rat  now  almost  exterminated  by  the  European  rat. 
Though  the  difficulty  is  great  to  our  imagination,  and  really 
great,  if  we  wish  to  ascertain  the  precise  causes  and  their 
manner  of  action,  it  ought  not  to  be  so  to  our  reason,  as  long  as 
we  keep  steadily  in  mind  that  the  increase  of  each  species  and 
each  race  is  constantly  checked  in  various  ways  ;  so  that  if  any 
new  check,  even  a  slight  one,  be  superadded,  the  race  will  surely 
decrease  in  number;  and  decreasing  numbers  will  sooner  or 

««  These  details  are  taken   from  Jlay    29th,    1863.      The    following 

'The  Mutineers  of  the  "Bounty,"'  statements  about  the  Sandwich  Is- 

by  Lady  Belcher,  1870;    and  from  landers    are    from    the    'Honolulu 

♦Titcairn    Island,'    ordered    to    be  Gazette,'  and  from  Mr.  Coan. 
printed  by  the  House  of  Commons, 


192  The  Descent  of  Man.  Part  I. 

later  lead  to  extinction ;  the  end,  in  most  cases,  being  promptly 
determined  by  the  inroads  of  conquering  tribes. 

On  the  Formation  of  the  Races  of  Man. — In  some  cases  the 
crossing  of  distinct  races  has  led  to  the  formation  of  a  new  race. 
The  singular  fact  that  Europeans  and  Hindoos,  who  belong  to 
the  same  Aryan  stock,  and  speak  a  language  fundamentally  the 
same,  differ  widely  in  appearance,  whilst  Europeans  differ  but 
little  from  Jews,  who  belong  to  the  Semitic  stock,  and  speak 
quite  another  language,  has  been  accounted  for  by  Broca,^^ 
through  certain  Aryan  branches  having  been  largely  crossed 
by  indigenous  tribes  during  their  wide  diffusion.  When  two 
races  in  close  contact  cross,  the  first  result  is  a  heterogeneous 
mixture :  thus  Mr.  Hunter,  in  describing  the  SantaH  or  hill-tribes 
of  India,  says  that  hundreds  of  imperceptible  gradations  may  be 
traced  "  from  the  black,  squat  tribes  of  the  mountains  to  the  tall 
"  olive-coloured  Brahman,  with  his  intellectual  brow,  calm  eyes, 
*'and  high  but  narrow  head;"  so  that  it  is  necessary  in  courts 
of  justice  to  ask  the  witnesses  whether  they  are  Santalis  or 
Hindoos.^"  Whether  a  heterogeneous  people,  such  as  the  inhabi- 
tants of  some  of  the  Polynesian  islands,  formed  by  the  crossing 
of  two  distinct  races,  with  few  or  no  pure  members  left,  would 
ever  become  homogeneous,  is  not  kno\vn  from  direct  evidence. 
But  as  with  our  domesticated  animals,  a  cross-breed  can  certainly 
be  fixed  and-  made  uniform  by  careful  selection^^  in  the  course  of 
a  few  generations,  we  may  infer  that  the  free  intercrossing  of  a 
heterogeneous  mixture  during  a  long  descent  would  supply  the 
place  of  selection,  and  overcome  any  tendency  to  reversion ;  so 
that  the  crossed  race  would  ultimately  become  homogeneous, 
though  it  might  not  partake  in  an  equal  degree  of  the  characters 
of  the  two  parent-races. 

Of  all  the  differences  between  the  races  of  man,  the  colour  of 
the  skin  is  the  most  conspicuous  and  one  of  the  best  marked.  It 
was  formerly  thought  that  differences  of  this  kind  could  be 
accounted  for  by  long  exposure  to  different  climates;  but 
Pallas  first  shewed  that  this  is  not  tenable,  and  he  has  since  been 
followed  by  almost  all  anthropologists.^^  This  view  has  been 
rejected  chiefly  because  the  distribution  of  the  variously 
coloured  races,  most  of  whom  must  have  long  inhabited  their 

*9  *0n     Anthropology,'    transla-  "  Pallas,  'Act.  Acad.  St.  Peters- 

tion    '  Aathropolog.    Keview,'    Jan.  burg,'  1780,  part  ii.  p.  69.     He  was 

1868,  p.  38.  followed  by  Kudolphi,  in  his  '  Bcy- 

^•^  'The  Annals  of  Rural  Bengal,'  trage     zur     Anthropologic,'    1812. 

1868,  p.  134.  An  excellent  summary  of   the  evi- 

^'  '  The  Variation  of  Animals  and  dence    is    given    by    Godron,    '  De 

Plants  under  Domestication,'  vol.  ii.  I'Espece,'  1859,  vol.  ii.  p.  246,  &c. . 
p.  95. 


CiiAi'.  YII.  The  Formation  of  Races.  193 

present  homes,  does  not  coincide  with  corresponding  differences 
of  climate.  Some  httle  weight  may  be  given  to  such  cases  as 
that  of  the  Dutch  fomihes,  who,  as  we  hear  on  excellent  autho- 
rity," have  not  undergone  the  least  change  of  colour  after 
residing  for  three  centuries  in  South  Africa.  An  argument  on 
the  same  side  may  likewise  be  drawn  from  the  uniform  appear- 
ance in  various  parts  of  the  world  of  gipsies  and  Jews,  though 
the  uniformity  of  the  latter  has  been  somewhat  exaggerated.^* 
A  very  damp  or  a  very  dry  atmosphere  has  lx)on  supposed  to  be 
more  influential  in  modifying  the  colour  of  the  skin  than  mere 
heat ;  but  as  D'Orbigny  in  South  America,  and  Livingstone  in 
Africa,  arrived  at  diametrically  opposite  conclusions  with  respect 
to  dampness  and  dryness,  any  conclusion  on  this  head  must  be 
considered  as  very  doubtful. ^^ 

Various  facts,  which  I  have  given  elsewhere,  prove  that  the 
colour  of  the  skin  and  hair  is  sometimes  correlated  in  a  surpris- 
ing manner  with  a  complete  immunity  from  the  action  of  certain 
vegetable  poisons,  and  from  the  attacks  of  certain  parasites. 
Hence  it  occurred  to  me,  that  negroes  and  other  dark  races 
might  have  acquired  their  dark  tints  by  the  darker  individuals 
escaping  from  the  deadly  influence  of  the  miasma  of  their 
native  countries,  during  a  long  series  of  generations. 

I  afterwards  found  that  this  same  idea  had  long  ago  occurred 
to  Dr.  Wells.^^  It  has  long  been  known  that  negroes,  and  even 
mulattoes,  are  almost  completely  exempt  from  the  yellow -fever, 
so  destructive  in  tropical  America.^^  They  likewise  escape  to  a 
large  extent  the  fatal  intermittent  fevers,  that  prevail  along  at 
least  2600  miles  of  the  shores  of  Africa,  and  which  annually 
cause  one-fifth  of  the  white  settlers  to  die,  and  another  fifth  to 
return  home  invalided.^*  This  immunity  in  the  negro  seems  to 
be  partly  inherent,  depending  on  some  unknown  peculiarity  of 
constitution,  and  partly  the  result  of  acclimatisation.   Pouchet** 

*'  Sir  Andrew  Smith,  as  quoted  the  Historical  Sketch  (p.  xvi.)  to  niy 

by  Knox,  '  Kaces  of  Man  '  1850,  p.  '  Origin   of  Species.'     Various  cases 

473.  of  colour  correlated  with  constitu- 

^*  See    De    Quatrefages    on    this  tional  peculiarities  are  given  in  my 

head,    '  Revue    des    Cours    Scienti-  '  Variation    of  Animals    under   Do- 

fiques,'  Oct.  17,  1868,  p.  731.  mestication,'  vol.  ii.  pp.  227,  335. 

^*  Livingstone's  '  Travels  and  Re-  ^^  See,    for    instance,    Nott    and 

searches    in    S.    Africa,'    1857,    pp.  Gliddon,  '  Types  of  Mankind,' p.  08. 

338,  329.     D'Orbigny,  as  quoted  by  *'  Major  Tulloch,  in  a  i)ai)er  read 

Godron,    '  De    I'Espfece,'    vol.    ii.    p.  before  the  Statistical  Sociotv,  April 

206.  20th,  1840,  and  given  in  the' '  Athe- 

^®  See  a  paper    read    before    the  uaium,'  1840,  p.  353. 

Royal  Soc.   in  1813,  and  published  *"  'The  Plurality  of  the  Human 

in  his  Essays  in  1818.     I  have  given  Race  '  (translat.),  1864,  p.  60. 
an  account  of  Dr.  Wells'   views   in 


194  The  Descent  of  Man.  Part  I. 


states  that  the  negro  regiments  recruited  near  the  Soudan,  and 
borrowed  from  the  Viceroy  of  Egypt  for  the  Mexican  war. 
escaped  the  yellow-fever  almost  equally  with  the  negroes  origin- 
ally brought  from  various  parts  of  Africa  and  accustomed  to  the 
climate  of  the  West  Indies.  That  acclimatisation  plays  a  part, 
is  shewn  by  the  many  cases  in  which  negroes  have  become  some- 
what liable  to  tropical  fevers,  after  having  resided  for  some  time 
in  a  colder  climate.'^°  The  nature  of  the  climate  under  which  the 
white  races  have  long  resided,  likewise  has  some  influence  on 
them;  for  during  the  fearful  epidemic  of  yellow-fever  in 
Demerara  during  1837,  Dr.  Blair  found  that  the  death-rate  of  the 
immigrants  was  proportional  to  the  latitude  of  the  country 
whence  they  had  come.  With  the  negro  the  immunity,  as  far  as 
it  is  the  result  of  acclimatisation,  implies  exposure  during  a 
prodigious  length  of  time ;  for  the  aborigines  of  tropical  America 
who  have  resided  there  from  time  immemorial,  are  not  exemjDt 
from  yellow  fever;  and  the  Eev.  H.  B.  Tristram  states,  that 
there  are  districts  in  Northern  Africa  which  the  native  inhabit- 
ants are  compelled  annually  to  leave,  though  the  negroes  can 
remain  with  safety. 

That  the  immunity  of  the  negro  is  in  any  degree  correlated 
with  the  colour  of  his  skin  is  a  mere  conjecture :  it  may  be 
correlated  with  some  difference  in  his  blood,  nervous  system,  or 
other  tissues.  Nevertheless,  from  the  facts  above  alluded  to,  and 
from  some  connection  apparently  existing  between  complexion 
and  a  tendency  to  consumption,  the  conjecture  seemed  to  me 
not  improbable.  Consequently  I  endeavoured,  with  but  little 
success,^^  to  ascertain  how  far  it  holds  good.     The  late  Dr. 

^^  Quatrefjiges,  *  Unite  de  I'Espece  "  is  some  limited  degree  of  relation 

Humiiine,'    1861,    p.    205.      Waitz,  "  between  the  colour  of  the  races  of 

'  Introduct.  to  Anthropology,'  trans-  "  man  and  the  climate  inhabited  bv 

lat.  vol.  i.   1863,   p.   124.     Living-  "  them ;    the    following    invostiga- 

stone  gives   analogous  cases   in  his  "'tion    seems    worth    consideration. 

'  Travels.'  "  Namely,  whether  there  is  any  re- 

"1  In  the    spring  of   1862  I   ob-  "  lation  in  Europeans  between  the 

tained  permission  from  the  Director-  "  colour    of  their    hair,    and    their 

General  of  the  Medical  department  "  liability  to  the  diseases  of  tropical 

of  the  Army,  to    transmit    to    the  "countries.     If  the  surgeons  of  the 

surgeons  of  the  various  regiments  "  several  regiments,  when  stationed 

on    foreign    service    a    blank   tabL"!,  "  in    unhealthy    tropical    districts, 

with    the   ibllowing   appended    re-  "  would  be  so  good  as  first  to  count, 

marks,  but  I  have  received  no  re-  "  as  a  standard  of  comparison,  how 

turns.      "  As    several    well-marked  "  many  men,  in    the  force  whence 

"  cases    have    been    recorded    with  "  the    sick    are    drawn,   have    dark 

*'  our  domestic  animals  of  a  relation  "  and  light-coloured  hair,  and  hair 

"  between  the  colour  of  the  dermal  "  of  intermediate  or  doubtful  tints  ; 

"  appendages  and  the  constitution ;  "  and    if    a   similar   account   were 

"  and  it  being  notorious  that  there  "  kept  by  the  same  medical  gentle- 


Chap.  Vir.  The  Format  ion  vf  Races.  195 

Daniell,  who  had  long  lived  on  the  West  Coast  of  Africa,  told  me 
that  he  did  not  believe  in  any  such  relation.  He  was  himself 
unusually  fair,  and  had  withstood  the  climate  in  a  wonderful 
manner.  ^Vhen  he  first  arrived  as  a  boy  on  the  coast,  an  old  and 
experienced  negro  chief  predicted  from  his  appearance  that  this 
would  prove  the  case.  Dr.  Nicholson,  of  Antigua,  after  having 
attended  to  this  subject,  writes  to  me  that  he  does  not  think  that 
dark-coloured  Europeans  escape  the  yellow-fever  more  than 
those  that  are  light-coloured.  Mr.  J.  ]\I.  Harris  altogether 
denies  that  Europeans  with  dark  hair  withstand  a  hot  climate 
better  than  other  men  :  on  the  contrary,  experience  lias  taught 
him  in  making  a  selection  of  men  for  service  on  the  coast  of 
Africa,  to  choose  those  with  red  hair.*'-  As  far,  therefore,  as 
these  slight  indications  go,  there  seems  no  foundation  for  the 
hypothesis,  that  blackness  has  resulted  from  the  darker  and 
darker  individuals  having  survived  better  during  long  exposure 
to  fever-generating  miasma. 

Dr.  Sharpe  remarks,^^  that  a  tropical  sun,  which  burns  and 
blisters  a  white  skin,  does  not  injure  a  black  one  at  all ;  and,  as 
he  adds,  this  is  not  due  to  habit  in  the  individual,  for  children 
only  six  or  eight  months  old  are  often  carried  about  naked,  and 
are  not  affected.  I  have  been  assured  by  a  medical  man,  that 
some  years  ago  during  each  summer,  but  not  during  the  winter, 
his    hands   became   marked  with  light  brown  patches,   like. 


"  men.  of  all  the  men  who  suffered  "  of  generations." 

"  from  malarious  and  yellow  fevers,  ^-  'Anthropological  RcA'iew,' Jan. 

"  or  from  dysentery,  it  would  soon  1866,  p.  xxi.     Dr.  Sharpe- also  says, 

*'  he  apparent,  after  some  thousand  with  respect  to  India  ('  Man  a  Spe- 

*' cases  had  been  tabulated,  whether  cial  Creation,'  1873,  p.    118),  that 

"  there  exists  any  relation  between  "  it  has  been  noticed  by  some  medi- 

"  the  colour  of  the  hair  and  consti-  "  cal   otHcers  that  Europeans   with 

"  tutional  liability  to  tropical  dis-  "  light  hair  and  florid  complexions 

"eases.     Perhaps  no  such  relation  "  sutler  less  fi'om  diseases  of  tropical 

*'  would  be  discovered,  but  the  in-  "  countries  than  persons  with  dark 

*'  vestigation  is  well  worth  making.  "  hair     and     sallow     complexions; 

*'  In  case   any  positive   result  were  "  and,  so  far  as  1  know,  there  ap- 

"  obtained,    it    might    be    of    some  "  pear  to  be  good  grounds  for  this 

"  practical  use  in  selecting  men  for  "  remark."      On    the    other    hand, 

"  any  particular  service.     Theoreti-  ]\Ir.   Heddle,  of  Sierra  Leone  "  who 

"  cally  the  result  would  be  of  high  "  has  had  more  clerks  killed  under 

"  interest,  as  indicating  one  means  "  him  than  any  other  man,"  by  the 

"  by  which  a  race  of  men  inhabiting  climate  of  the  West   African  Coast 

"from    a    remote    period    an    un-  (\V.  Reade,  'African  Sketch  Book,' 

'■  healthy    tropical    climate,    might  vol.    ii.    p.    52'J),    holds  a   directly 

"  have    become    dark-coloured    by  opposite  view,  as  does  Capt.  Burton, 

"the  better  preservation  of  dark-  ^^  '  Man  a  Special  Creation,' 1873, 

"  haired    or   dark-complexioned  in-  p.  119. 
*'  dividuals  duriusr  a  long  succession 


196  The  Descent  of  Ma7t.  Part  I. 

although  larger  than  freckles,  and  that  these  patches  were  never 
affected  by  sun-burning,  -whilst  the  white  parts  of  his  skin 
have  on  several  occasions  been  much  inflamed  and  blistered. 
With  the  lower  animals  there  is,  also,  a  constitutional  difference 
in  liability  to  the  action  of  the  sun  between  those  parts  of  the 
skin  clothed  with  white  hair  and  other  parts.^*  Whether  the 
saving  of  the  skin  from  being  thus  burnt  is  of  sufficient  impor- 
tance to  account  for  a  dark  tint  having  been  gradually  acquired 
by  man  through  natural  selection,  I  am  unable  to  judge.  If  it 
be  so,  we  should  have  to  assume  that  the  natives  of  tropical 
America  have  lived  there  for  a  much  shorter  time  than  the 
negroes  in  Africa,  or  the  PajDuans  in  the  southern  parts  of  the 
Malay  archipelago,  just  as  the  lighter-coloured  Hindoos  have 
resided  in  India  for  a  shorter  time  than  the  darker  aborigines  of 
the  central  and  southern  jDarts  of  the  peninsula. 

Although  with  our  present  knowledge  we  cannot  account  for 
the  differences  of  colour  in  the  races  of  man,  through  any 
advantage  thus  gained,  or  from  the  direct  action  of  climate ;  yet 
we  must  not  quite  ignore  the  latter  agency,  for  there  is  good 
reason  to  believe  that  some  inherited  effect  is  thus  produced.*^^ 

We  have  seen  in  the  second  chapter  that  the  conditions  of  life 
affect  the  development  of  the  bodily  frame  in  a  direct  manner, 
and  that  the  effects  are  transmitted.  Thus,  as  is  generally 
admitted,  the  European  settlers  in  the  United  States  undergo  a 
slight  but  extraordinarily  rapid  change  of  appearance.  Their 
bodies  and  limbs  become  elongated;  and  I  hear  from  Col. 
Bernys  that  during  the  late  war  in  the  United  States,  good 
evidence  was  afforded  of  this  fact  by  the  ridiculous  appearance 
presented  by  the  German  regiments,  when  dressed  in  ready-made 
clothes  manufactured  for  the  American  market,  and  which  were 
much  too  loug  for  the  men  in  every  way.  There  is,  also,  a  con- 
siderable body  of  evidence  shewing  that  in  the  Southern  States 
the  house-slaves  of  the  third  generation  present  a  markedly 
different  appearance  from  the  field-slaves.^^ 

^*  '  Variation    of    Animals    and  settled  in  Georgia,  have  acquired  in 

Plants  under  Domestication,'  vol.  ii.  the  course  of  two  generations  dark 

pp.  336,  337.  hair  and  eyes.     Mr.  D.  Forbes  in- 

^^  See,  for  instance,  Quatrefages  forms  me  that  the  Quichuas  in  the 

('  Revue    des    Cours    Scientifiques,'  Andes  vary  greatly  in   colour,  ac- 

Oct.  10,  1868,  p.  724)  on  the  effects  cording  to  the  position  of  the  valleys 

of  residence  in  Abyssinia  and  Arabia,  inhabited  by  them, 
and    other    analogous    cases.      Dr.  ^^  Harlan,   '  Medical   Piesearches,' 

liolle  ('  Dcr  Mensch,  seine  Abstam-  p.     532.     Quatrefages    ('  Unite'    de 

mung,'  &c.,  1865,  s.  99)  states,  on  I'Espece    Humaine,'    1861,    p.    128) 

the  authority  <ii  Khanikof,  that  the  has  collected  much  evidence  on  this 

greater  number  of  German  families  head. 


Chap.  VII.  The  Formation  of  Races.  1 97 

If,  however,  we  look  to  the  races  of  man  as  distributed  over 
the  world,  we  iiuist  infer  that  their  characteristic  differences  can- 
not be  accounted  for  by  the  direct  action  of  different  conditions 
of  life,  even  after  exposure  to  them  for  an  enormous  period  of 
time.  The  Esquimaux  live  exclusively  on  animal  food ;  they  are 
clothed  in  thick  fur,  and  are  exposed  to  intense  cold  and  to 
prolonged  darkness;  yet  they  do  not  differ  in  any  extreme 
degree  from  the  inhabitants  of  Southern  China,  who  live  entirely 
on  vegetable  food,  and  are  exposed  almost  naked  to  a  hot,  glaring 
climate.  The  unclothed  Fuegians  live  on  the  marine  produc- 
tions of  their  inhospitable  shores ;  the  Botocudos  of  Brazil 
wander  about  the  hot  forests  of  the  interior  and  live  chiefly  on 
vegetable  productions ;  yet  these  tribes  resemble  each  other  so 
closely  that  the  Fuegians  on  board  the  "  Beagle  "  were  mistaken 
by  some  Brazilians  for  Botocudos.  The  Botocudos  again,  as 
well  as  the  other  inhabitants  of  tropical  America,  are  wholly 
different  from  the  Negroes  who  inhabit  the  opposite  shores  of 
the  Atlantic,  are  exposed  to  a  nearly  similar  climate,  and  follow 
nearly  the  same  habits  of  life. 

Nor  can  the  differences  between  the  races  of  man  be  accounted 
for  by  the  inherited  effects  of  the  increased  or  decreased  use  of 
parts,  except  to  a  quite  insignificant  degree.  Men  who  habitu- 
ally live  in  canoes,  may  have  their  legs  somewhat  stunted ; 
those  who  inhabit  lofty  regions  may  have  their  chests  enlarged  ; 
and  those  who  constantly  use  certain  sense-organs  may  have  the 
cavities  in  which  they  are  lodged  somewhat  increased  in  size,  and 
their  features  consequently  a  little  modified.  AVith  civilised 
nations,  the  reduced  size  of  the  jaws  from  lessened  use — the 
habitual  play  of  different  muscles  serving  to  express  different 
emotions — and  the  increased  size  of  the  brain  from  greater 
intellectual  activity,  have  together  produced  a  considerable 
effect  on  their  general  appearance  when  compared  with 
savages.*''  Increased  bodily  stature,  without  any  corresponding 
increase  in  the  size  of  the  brain,  may  (judging  from  the  pre- 
viously adduced  case  of  rabbits),  have  given  to  some  races  an 
elongated  skull  of  the  dolichocephalic  type. 

Lastly,  the  little-understood  principle  of  correlated  develop- 
ment has  sometimes  come  into  action,  as  in  the  case  of  great 
muscular  development  and  strongly  projecting  supra-orbital 
ridges.  The  colour  of  the  skin  and  hair  are  plainly  correlated,  as 
is  the  texture  of  the  hair  with  its  colour  in  the  INlandans  of 
North  America.'^**  The  colour  also  of  the  skin,  and  the  odour 

«'  See  Prof.  SchaanTiaiison,  trans-  ^^  ^\^,^  Catlin  states  (' N.  Ameri- 

lat.    in    'Anthropological    Keview,'      can   Indians,'  3rd  edit.  1812,  vol.  I. 
Oct.  1868,  p.  429.  p.  49)  that  in  the  whole  tribe  of 


198  The  Descent  of  Man.  Part  1. 

emitted  "by  it^  are  likewise  in  some  maimer  connected.  With  the 
breeds  of  sheep  the  number  of  hairs  within  a  given  space  and  the 
number  of  the  excretory  pores  are  related.*'^  If  we  may  judge 
from  the  analogy  of  our  domesticated  animals,  many  modifica- 
tions of  structure  in  man  probably  come  under  this  principle  of 
correlated  development. 

We  have  now  seen  that  the  external  characteristic  differences 
between  the  races  of  man  cannot  be  accounted  for  in  a  satisfac- 
tory manner  by  the  direct  action  of  the  conditions  of  life,  nor  by 
the  effects  of  the  continued  use  of  parts,  nor  through  the 
principle  of  correlation.  We  are  therefore  led  to  inquire 
whether  slight  individual  differences,  to  which  man  is  emi- 
nently liable,  may  not  have  been  preserved  and  augmented 
during  a  long  series  of  generations  through  natural  selection. 
But  here  we  are  at  once  met  by  the  objection  that  beneficial 
variations  alone  can  be  thus  preserved ;  and  as  far  as  we  are 
enabled  to  judge,  although  always  liable  to  err  on  this  head,  none 
of  the  differences  between  the  races  of  man  are  of  any  direct  or 
special  service  to  him.  The  intellectual  and  moral  or  social 
faculties  must  of  course  be  excepted  from  this  remark.  The  great 
variability  of  all  the  external  differences  between  the  races  of  man, 
likewise  indicates  that  they  cannot  be  of  much  importance ;  for 
■if  important,  they  would  long  ago  have  been  either  fixed  and 
preserved,  or  eliminated.  In  this  resj^ect  man  resembles  those 
forms,  called  by  naturalists  protean- or  polymorphic,  which  have 
remained  extremely  variable,  owing,  as  it  seems,  to  such  varia- 
tions being  of  an  indifferent  nature,  and  to  their  having  thus 
escaped  the  action  of  natural  selection. 

We  have  thus  far  been  baffled  in  all  our  attempts  to  account 
for  the  differences  between  the  races  of  man  ;  but  there  remains 
one  important  agency,  namely  Sexual  Selection,  which  appears  to 
have  acted  powerfully  on  man,  as  on  many  other  animals.  I  do 
not  intend  to  assert  that  sexual  selection  will  account  for  all  the 
differences  between  the  races.  An  unexplained  residuum  is  left, 
about  which  we  can  only  say,  in  our  ignorance,  that  as  indivi- 
duals are  continually  born  with,  for  instance,  heads  a  little 
rounder  or  narrower,  and  with  noses  a  little  longer  or  shorter, 
such  slight  differences  might  become  fixed  and  uniform,  if  the 


the  Maodans,  about  one    in  ten   or  fine  and  soft. 

twelve  of  the  members,  of  all  ages  ®^  On    the    odour    of    the    skin, 

and  both  sexes,  have  bright  silvery  Godron,  '  Sur  I'Espece,'  torn.  ii.   p. 

grey    hair,    wliich     is    hereditary.  217.     On    the    pores   in    the    skin, 

Now    this    hair    is    as    coarse    and  Dr.   Wilckens,   '  Die   Aufgaben    der 

harsh   as  that   of  a    horse's   mane,  Landwirth.    Zootechnik,'  1869,  s.  7. 
whilst  the  hair  of  other  colours  is 


Chap.  YII.  Structure  of  the  Brain.  199 


unknown  agencies  which  induced  them  were  to  a^  in  a  more 
constant  manner,  aided  by  long-continued  intercrossing.  Such 
variations  come  under  the  provisional  class,  alluded  to  in  our 
second  chai)ter,  which  for  the  want  of  a  better  term  arc  often 
called  spontaneous.  Nor  do  I  pretend  that  the  etfects  of  sexual 
selection  can  be  indicated  with  scientific  precision ;  but  it  can  bo 
shewn  that  it  would  be  an  inexplicable  fact  if  man  had  not  been 
modified  by  this  agency,  which  appears  to  have  acted  powerfully 
on  innumerable  animals.  It  can  further  be  shewn  that  the 
differences  between  the  races  of  man,  as  in  colour,  hairiness, 
form  of  features,  &c.,  are  of  a  kind  which  might  have  been 
expected  to  come  under  the  influence  of  sexual  selection.  But  in 
order  to  treat  this  sul»ject  i)roperly,  I  have  found  it  necessary  to 
pass  the  whole  animal  kingdom  in  review.  I  have  therefore 
devoted  to  it  the  Second  Part  of  this  work.  At  the  close  I  shall 
return  to  man,  and,  after  attempting  to  shew  how  far  he  has 
been  modified  through  sexual  selection,  will  give  a  brief  summary 
of  the  chapters  in  this  First  Part. 


Note  ox  the  Resemblances  and  Differences  in  the  Structure 
AND  THE  Development  of  the  Brain  in  Man  and  Apes.  By 
Professor  Huxlev,  F.R.S. 

The  controversy  respecting  the  nature  and  the  extent  of  the  differ- 
ences in  the  slructme  of  the  brain  in  man  and  the  apes,  which  arose 
some  fifteen  years  ago,  has  not  yet  come  to  an  end,  though  the  subject 
matter  of  the  dispute  is,  at  present,  totally  different  fronr  what  it  was 
formerly.  It  was  originally  asserted  and  re-asserted,  with  singular 
pertinacity,  that  the  biain  of  all  the  apes,  even  the  highest,  differs  from 
that  of  man,  in  the  absence  of  such  conspicuous  structures  as  the 
posterior  lobes  of  the  cerebral  licmifepheres,  with  the  posterior  cornu  of 
the  lateral  ventricle  and  the  hi ppocampus  minor,  contained  in  those 
lobes,  which  are  so  obvious  in  man. 

But  the  truth  that  tlie  three  structm-es  in  question  are  as  well  deve- 
loped in  apes'  as  in  human  brains,  or  even  b.  ttei  ;  and  that  it  is  character- 
istic of  all  the  rrhnaits  (if  we  exclude  the  Lennu's)  to  have  these  parts 
well  developed,  stands  at  present  on  as  secure  a  basis  as  any  profiosition 
in  comparative  anatomy.  Moreover,  it  is  admitted  by  every  one  of  the 
long  series  of  anatomihts  who,  of  late  years,  have  paid  special  attention  to 
the  arrangtment  of  the  complicated  sulci  and  gyri  which  sippcar  up;in 
the  surface  of  the  cerebral  hemispheres  in  man  and  the  higher  ape-', 
that  ihey  are  di?posed  after  the  very  same  pattern  in  him,  a>  in  them. 
Every  principal  gyrus  and  sulcus  of  a  chim])anzee's  bram  is  ( h  arly 
rcpiesentLd  in  that  of  a  man,  so  that  the  terminology  which  ajjpHcs  to 
the  one  answers  lor  the  other.  On  this  point  there  is  no  ditlVrenrc  of 
opinion.  Some  years  since,  Professor  Bischoff  published  a  memoir  '"  on 
the  C('rebral  convolutions  of  man  and  apes;  and  as  the  ptupnse  of 
my  learned  colleague  was  certainly  not  to  diminish  the  value  of  the 


*"  'Die  Gi'osshirn-Wmdungen  des   Menschen ;'    '  AbhaQJluugea    dev  K, 
Bayerischeu  Akademie.*  Bd.  x.,  18G8. 


200  •  The  Descent  of  Man.  Part  1. 

differences  between  apes  and  men  in  this  respect,  I  am  glad  to  make  a 
citation  from  him. 

"  That  the  apes,  and  especially  the  orang,  chimpanzee  and  gorilla, 
"  come  very  close  to  man  in  their  organisation,  much  nearer  than  to  any 
"  otlier  animal,  is  a  well  known  fact,  disputed  by  nobody.  Looking  at 
"  the  matter  from  the  point  of  view  of  organisation  alone,  no  one  probably 
"  would  ever  have  disputed  Ihe  view  of  Linnaeus,  that  man  should  be 
"  placed,  merely  as  a  peculiar  species,  at  the  hs  ad  of  the  mammalia  and  of 
*'  those  apes.  Both  shew,  in  all  their  organs,  so  close  an  affinity,  that  the 
"  most  exact  anatomical  investigation  is  needed  in  order  to  demonstrate 
"  those  differences  which  really  exist.  So  it  is  with  the  brains.  The 
"  brains  of  man,  the  orang,  the  chimpanzee,  the  gorilla,  in  spite  of  all 
''  the  important  differences  which  they  present,  cume  very  close  to  one 
"  another  '  (1.  c.  p.  101). 

Tiiere  remains,  then,  no  dispute  as  to  the  resemblance  in  fundamental 
characters,  between  the  ape's  brain  and  man's  ;  nor  any  as  to  the  won- 
derfully close  similarity  between  the  chimpanzee,  orang  and  man,  in 
even  the  details  of  the  arrangement  of  the  gyri  and  sulci  of  the  cerebral 
hemispheres.  Nor,  turning  to  the  differences  between  the  brains  of 
the  highest  apes  and  that  of  man,  is  there  any  serious  question  as  to 
the  nature  and  extent  of  these  differences.  It  is  admitted  that  the  man's 
cerebral  hemispheres  are  absolutely  and  relatively  larger  than  those  of 
the  orang  and  chimpanzee  ;  that  his  frontal  lobes  are  less  excavated  by 
the  upward  protrusion  of  the  roof  of  the  orbits  ;  that  his  gyri  and  sulci 
are.  as  a  rule,  less  symmetrically  disposed,  and  present  a  greater  num^jer 
of  secondary  plications.  And  it  is  admitted  that,  as  a  rule,  in  man,  the 
temporo-occipital  or  "external  perpendicular"  fissure,  which  is  usually 
so  strongly  marked  a  feature  of  the  ape's  brain  is  but  faintly  marked. 
But  it  is  also  clear,  that  none  of  these  differences  constitutes  a  sharp 
demarcation  between  the  man's  and  the  ape's  brain.  In  respect  to  the 
external  perpendicular  fissure  of  Gratiolet,  in  the  human  brain,  for 
instance.  Professor  Turner  remarks  :'^ 

"  In  some  brains  it  appears  simply  as  an  indentation  of  the  margin  of 
"  the  hemisphere,  but,  in  others,  it  extends  for  some  distance  more  or  less 
"  transversely  outwards.  I  saw  it  in  the  right  hemisphere  of  a  female 
"  brain  pass  more  than  two  inches  outwards  ;  and  in  another  specimen, 
"  also  the  right  hemisphere,  it  proceeded  for  four-tenths  of  an  inch  out- 
"  wards,  and  then  extended  downwards,  as  far  as  the  lower  margin  of  the 
"  outer  smface  of  the  hemisphere.  The  imperfect  definition  of  this  fissure 
"  in  the  majority  of  human  brains,  as  compared  with  its  remarkable  dis- 
"  tinctness  in  the  brain  of  most  Quadrumana,  is  owing  to  the  presence,  in 
"  the  former,  of  certain  superficial,  well  marked,  secondary  convolutions 
"  which  bridge  it  over  and  connect  the  parietal  with  the  occipital  lobe. 
"  The  closer  the  first  of  these  bridging  gyri  lies  to  the  longitudinal 
"  fissure,  the  shorter  is  the  external  parieto-occipital  fissure."  (1.  c  p.  12.) 

The  obliteration  of  the  external  perpendicular  fissure  of  Gratiolet, 
therefore,  is  not  a  constant  character  of  the  human  brain.  On  the  other 
hand,  its  full  development  is  not  a.  constant  character  of  the  higher 
ape's  brain.  For,  in  the  chimpanzee,  the  more  or  less  extensive  oblitera- 
tion of  the  exteraal  perpt-mUeular  sulcus  by  "  bridging  convolutions,"  on 
one  side  or  the  other,  has  been  noted  over  and   over  again  bv  Prof 


'*  'Convolutions  of  the  Human   Cerebrum  Topographically  Considered, 
1866,  p.  12. 


Chap.  VII.  Siriictiire  of  the  Brain.  20 1 

Rolleston,  Mr.  Marshall,  -M.  Brora  and  Professor  Turner.  At  the 
conclusion  of  a  special  paper  on  this  subject  the  latter  writes :' - 

•'  The  three  specimens  of  the  brain  of  a  chimpanzee  just  described, 
*'  prove,  that  the  generalisation  which  Gratiolet  has  attempted  to  draw  of 
"  the  complete  absence  of  the  first  ccmnecting  convolution  and  the 
"  concealment  of  the  second,  as  essentially  characteristic  features  in  th(; 
"  brain  of  this  animal,  is  by  no  means  universally  applicable.  In  only  one 
"  specimen  did  the  brain,  in  these  particulars,  follow  the  law  which 
"  Gratiolet  has  expressed.  As  regards  the  presence  of  the  su])eriur  britlg- 
"  ing  convolution,  I  am  inclined  to  think  that  it  has  existed  in  one  hemi- 
"  sphere,  at  least,  in  a  majority  of  the  brains  of  this  animal  which  have,  up 
"  to  this  time,  been  figured  or  described.  The  superficial  position  of  the 
"  second  bridging  convolution  is  evidently  less  frequent,  and  has  as  yet, 
•'  I  believe,  only  been  seen  in  the  brain  (A)  recorded  in  this  communi- 
"  cation.  The  asymmetrical  arrangement  in  the  convolutions  of  the 
"  two  hemispheres,  which  previous  observers  have  referred  to  in  their 
"  descriptions  is  also  well  illustrated  in  these  specimens."  (pp.  8,  9.) 

Even  were  the  presence  of  the  temporo-occipital,  or  external  per- 
pendicular, sulcus  a  mark  of  distinction  between  the  higher  apes  an  1 
man,  the  value  of  such  a  distinctive  character  would  be  rendered  very 
doubtful  by  the  structure  of  the  brain  in  the  Piatyrhine  apes.  In  fact 
wliile  the  temporo-occipital  is  one  of  the  most  con.-itant  of  sulci  in 
the  Catarhine,  or  Old  World,  apes,  it  is  never  very  strongly  developed 
in  the  New  World  apes ;  it  is  absent  in  the  smaller  Platyrhini  ; 
rudimentary  in  Tithecia ;"  and  more  or  less  obliterated  by  bridging 
convolutions  in  Ateles. 

A  character  which  is  thus  variable  within  the  limits  of  a  single  group 
cfin  have  no  great  taxouomic  value. 

It  is  further  established,  that  the  degree  of  asymmetry  of  the  convolu- 
tion of  the  two  sides  in  the  human  brain  is  subject  to  nnieh  individual 
variation,;  and  that,  in  those  individuals  of  the  Bushman  race  who  have 
been  examined,  the  gyri  and  sulci  of  the  two  hemispheres  are  consider- 
ably less  complicated  and  more  symmetrical  than  in  the  European 
brain,  while,  in  some  individuals  of  the  chimpanzee,  their  complexity 
and  asymmetiy  become  notable.  This  is  particularly  tlie  case  in  the 
brain  of  a  young  male  chimpanzee  figured  by  M.  Broca.  ('  Lorch-e 
des  Primates,"  p.  165,  fig.  11.) 

Again,  as  respects  the  question  of  absolute  size,  it  is  established  that 
the  difterence  between  the  largest  and  the  smallest  healtiiy  human 
brain  is  greater  than  the  ditference  between  the  smallest  healthy 
human  brain  and  the  largest  chimpanzees  or  orang's  brain. 

Moreover,  there  is  one  circumstance  in  which  the  orang's  and  chim- 
panzee's brains  resemble  man's,  but  in  which  they  ditter  from  the  lower 
apes,  and  that  is  the  presence  of  two  corpora  candicantia — the 
t'ynomorpha  having  but  one. 

In  view  of  these  facts  I  do  not  hesitate  in  this  year  1874,  to  repeat 
and  insist  upon  the  proptjsition  which  I  inundated  in  ISGS.^'' 

"  So  far  as  cerebral  structure  goes,  therefore,  it  is  clear  that  man 


"  Notes    more   especially  on  the  "  Flower    'On    the  Anatomy    of 

bridging  convolutions  in  the  Brain  I'ithecia  Monackns,''  '  I'roceedings  of 

of  the  Chimpanzee,  'Proceedings  of  the  2lk)ol<)gical  Society,'  18G2. 

the    Royal    Society    of    Edinburgh,'  '*  '  Man's  Place  in  Nature,' p.  102. 
1865-6. 

10 


202  The  Descent  of  Man.  Part  L 

"  diffirs  less  from  the  chimpanzee  or  the  orang,  than  these  do  even 
"from  the  monkeys,  and  that  the  diflference  between  the  brain  of  the 
"•chimpanzee  and  of  man  is  almost  insignificant,  when  compared  with 
"that  between  the  chimpanzee  brain  and  that  of  a  Lemur." 

In  the  paper  to  which  I  have  referred,  Professor  Bischofif  does  not 
deny  tlie  second  part  of  this  statement,  but  he  first  makes  the  irrelevant 
remark  that  it  is  not  wonderful  if  the  brains  of  an  orang  and  a  Lemur 
are  very  different ;  and  secondly,  goes  on  to  assert  that,  "  If  we  succes- 
"  sively  compare  the  brain  of  a  man  with  that  of  an  orang ;  the  brain  of 
"  this  with  that  of  a  chimpanzee  ;  of  this  with  that  of  a  "orilla,  and  so 
"on  of  a  Hylobates,  Semnopithecus,  Cynocephalus,  Cercopithecus,  Macacus, 
"  Cehus,  Callithrix,  Lemur,  Stenops,  Hapale,  we  shall  not  meet  with  a 
"  greater,  or  even  as  great  a,  break  in  the  degree  of  development  of  the 
"convolutions,  as  we  find  between  the  brain  of  a  man  and  that  of  au 
"  orang  or  chimpanzee." 

To  which  I  reply,  firstly,  that  whether  this  assertion  be  true 
or  false,  it  has  nothing  whatever  to  do  with  the  proposition  enunciated 
in  'Man's  Place  in  Nature,'  which  refers  not  to  the  development  of  the 
convolutions  alone,  but  to  the  structure  of  the  whole  brain.  It  Professor 
Bischoff  had  taken  the  trouble  to  refer  to  p.  96  of  the  work  he  criticises, 
in  fact,  he  would  have  found  the  following  passage:  "And  it  is  a 
"  remarkable  circumstance  that  though,  so  far  as  our  present  know-  ■ 
"  ledge  extends,  there  is  one  true  structural  break  in  the  series  of  forms 
"  of  Simian  brains,  this  hiatus  does  not  lie  between  man  and  the 
"  manlike  apes,  but  between  the  lower  and  the  lowest  Simians,  or  in 
"  other  words,  between  the  Old  and  New  World  apes  and  monkeys  and 
"  the  Lemurs.  Every  Lemur  which  has  yet  been  examined,  in  fact, 
"  has  its  cerebellum  partially  visible  from  above;  and  its  posterior  lobe, 
"  with  the  contained  posterior  cornu  and  hippocampus  minor,  moie  or 
"less  rudimentary.  Every  marmoset,  American  monkey,  Old  World 
"  monkey,  baboon,  or  manlike  ape,  on  the  contrary,  has  its  cerebellum 
"  entirely  hidden,  posteriorly,  by  the  cerebral  lobes,  and  possesses  a 
"  large  posterior  cornu  with  a  well-developed  hippocampus  minor." 

This  statement  was  a  strictly  accurate  account  of  what  w^as  known 
when  it  was  made;  and  it  does  not  appear  to  me  to  be  more  than 
apparently  weakened  by  the  subsequent  discovery  of  the  relatively 
small  development  of  the  posterior  lobes  in  the  Siamang  and  in  the 
Howling  monkey.  Notwithstanding  the  exceptional  brevity  of  the 
posterior  lobes  in  these  two  species,  no  one  will  pretend  that  their 
brains,  in  the  slightest  degree,  approach  those  of  the  Lemurs.  And 
if,  instead  of  putting  Hapale  out  of  its  natural  place,  as  Professor 
Bischoff  most  unaccountably  does,  we  write  the  series  of  animals 
he  has  chosen  to  mention  as  follows  :  Homo,  Pithecus,  Troglodytes, 
Hylobates,  Semnopithecus,  Cynocephalus,  CercopWieeus,  Macacus,  Cebus, 
Callithrix,  Hapale,  Lemur,  Stenops,  I  venture  to  reaffirm  that  the 
great  break  in  this  series  lies  between  Hapale  and  Lemur,  and  that 
tills  break  is  considerably  greater  than  that  between  any  other  two 
terms  of  that  series.  Professor  Bischoff"  ignores  the  fact  that  long 
before  he  wrote,  Gratiolet  had  suggested  the  separation  of  the  Lemurs 
from  the  other  Primates  on  the  very  ground  of  the  difference  in  their 
cerebral  characters  ;  and  that  Professor  Flower  had  made  the  following 
observations  in  the  course  of  his  description  of  the  brain  of  the  Javan 
Lor  is  J  ^ 

'^  'Transactions  of  the  Zoological  Society,'  vol.  v.  1862. 


(JHAP.  VII.  Structure  of  the  Brain.  203 

"And  it  is  especially  remarkable  that,  in  the  development  of  the 
"  posterior  lobes,  there  is  no  approximation  to  the  Leinurine,  short 
"hemisphered,  brain,  in  those  monkeys  which  are  commonly  supposed 
'•  to  approach  this  family  in  other  respects,  viz.,  the  lower  members  of 
"  tlie  Fiatyrhine  gronp." 

So  far  as  the  structure  of  the  adult  brain  is  concerned,  then,  the  very 
considerable  additions  to  oar  knowledge,  which  have  been  made  by  the 
researches  of  so  many  investigators,  during  tlie  past  ten  years,  fully 
justify  the  statement  which  I  made  in  1SG3.  But  it  has  been  said 
that,  admitting  the  similarity  between  the  adult  brains  of  man  and 
apes,  they  are  nevertheless,  in  reality,  widely  diiFerent,  because  th(  y 
exhibit  fundamental  dilierences  in  the  mode  of  their  development.  No 
one  would  be  more  ready  tlian  I  to  admit  the  force  of  this  argument,  if 
8uch  fundamental  diiierenccs  of  development  really  exist,  liut  I  deny 
that  they  do  exist.  On  the  contrary,  there  is  a  fundamental  agree- 
ment in  the  developmeut  of  the  brain  in  men  and  apes. 

Gratiolet  originated  the  statement  that  there  is  a  fundamental 
difference  in  the  development  of  the  brains  of  apes  and  that  of  man — 
consisting  in  this;  that,  in  the  apes,  the  sulci  which  first  make  their 
appearance  are  situated  on  the  posterior  region  of  the  cerebral  hemi- 
spheres, while,  in  the  liuman  foatus,  the  sulci  first  become  visible  on  the 
frontal  lobes."* 

This  general  statement  is  based  upon  two  observations,  the  one  of  a 
Gibbon  almost  ready  to  be  born,  in  which  the  posterior  gyri  were  "  well 
"  developed,"  while  those  of  the  frontal  lobes  were'*  hardly  indicated  "''" 
(1.  c.  p.  39),  and  the  other  of  a  human  foetus  at  the  22nd  or  23rd  week 
of  uterogestation,  in  which  Gratiolet  notes  that  the  insula  was  un- 
covered, but  that  nevertheless  "  des  incisures  sement  le  lobe  anterieur, 
"  une  scissure  pen  profonde  indique  la  separation  du  lobe  occipital,  tres- 


''  "  Chez  tous  les  singes,  les  plis  Rolando,    and    one    of    the    frontal 

"  posterieurs  se  developpent  les  pre-  sulci,  plainly  enough.    Nevertheless, 

*'  miers ;     les     plis     anterieurs     se  M.    Alix,    in    his    '  Xotice    sur    les 

"  developpent    plus    tard,  aussi    la  travaux  anthropologiques  de  Gratio- 

*'  vert^bre  occipitale  et  la  parietale  let '  (Mem.  de  la  Societe  d'Anthro- 

**sont-ellesrelativemeuttres-grandes  pologie  de  Paris,'  1868,  p.    xxxii.), 

"  chez  le  foetus.    L'Horame  presente  writes  thus  :  "  Gratiolet  a  eu  outre 

*'  une  exception  remarquable  quant  "  les  mains  le  cerveau  d'un  fa>tus  de 

"  i  I'epoque  de  Tapparitioa  des  plis  "  Gibbon,     singe    emiuemment     su- 

"  frontaux,    qui    sont    les    premiers  "  perieur,  et  tellement  rapproche  de 

*'  indiquus  ;  niais  le  developpement  "  I'orang,  que  des  naturalistes  tres- 

*•  general  du  lobe  frontal,   envisage  "  competeuts  I'ont  range  parmi  les 

"  seulement     par     rapport    a     son  "  anthropoides.    M.  Huxley,  par  ex- 

*' volume,  suit  les  memeslois  que  dans  "  emple,  u'hesite  pas  sur  ce   point. 

"  les  singes  :"    Gratiolet,  '  Memoire  "  Eh  bien,  c'est  sur  le  cerveau  d'un 

sur  les  plis  cerebraux  de  I'Homme  *'  foetus  de  Gibbon  que  Gratiolet  a 

et    des    Primates,'    p.    39,    tab.   iv.  "  vu  les  circonvolutiuns  du  lobe  tem- 

fig.  3.  "  poro-Sfjhenoidal    deja    deceloppecs 

''''  Gratiolet's  words  are  (1.  c.  p.  '■'■  lorsqu'il  n  existent  pas  encore  de  plis 

39):   "Dans  le  foetus  don  t  11  s'agit  "  s/^r  le  lobe  frontal.     II  etait  done 

"  les  plis  cerebraux  posterieurs  sont  "  bien    autorise  a   dire    que,    cliez 

"  bien    developpes,    tandis    que    les  "  I'homme  les  circonvolutious  aj)pa- 

"  plis  du  lobe  frontal  sont  i  peine  "  raissent  d'a  eu  w,  tandis  que  chez 

"  indiques."     The    figure,    however  "  les   singes    elles    se    developpent 

(PI.  iv.  fig.  3),  shews   the  fissure  of  "  d'w  en  a." 


204  The  Descent  of  Man.  Part  I. 

"  rediiit,  d'ailleurs  des  cctte  epoque.  Le  reste  de  la  siuface  cerebrale 
"  eit  encore  absolument  lisse." 

Three  views  of  this  brain  are  given  in  Plate  11.  figs,  1,  2,  3,  of  the 
work  cited,  shewing  the  upper,  lateral  and  inferior  views  of  the  hemi- 
spheres, but  not  the  inner  view.  It  is  worthy  of  note  that  the  figure 
by  no  means  bears  out  Gratiolet's  description,  inasmuch  as  tlie  fissure 
(anterotempoi  al)  on  the  posterior  half  of  the  face  of  the  hemisphere,  is 
more  marked  than  any  of  those  vaguely  indicated  in  the  anterior  half. 
If  the  figure  is  correct  it  in  no  way  justifies  Gratiolet's  conclusion  : 
"  II  y  a  done  entre  ces  cerveaux  [those  of  a  Callithrix  and  of  a  Gibbon]  et 
"  celui  du  foetus  humain  une  ditierence  fondamental.  Chez  celui-ci,  long- 
"  temps  avant  que  les  plis  temporaux  apparaissent,  les  plis  frontaux 
"  essayent  d'exister." 

Since  Gratiolet's  time,  however,  the  development  of  the  gyri  and 
sulci  of  the  brain  has  been  made  the  subject  of  renewed  investigation 
by  Schmidt,  Bischoft',  Pansch,"*  and  more  particularly  by  Ecker,^^  whose 
work  is  not  only  the  latest,  but  by  far  the  most  complete,  memoir  on 
the  subject. 

The  final  resiilts  of  their  inquiries  may  b©  summed  up  as  follows  : — 

1.  In  the  human  foetus,  the  sylvian  fissure  is  formed  in  the  course  of 
the  third  month  of  uterogestation.  In  this,  and  in  the  fourth  month, 
the  cerebral  hemispheres  are  smooth  and  rounded  (with  the  exception 
of  the  sylvian  depression),  and  they  project  backwards  far  beyond  the 
cerebellum. 

2.  The  sulci,  properly  so  called,  begin  to  appear  in  the  interval 
between  the  end  of  the  fourth  and  the  beginning  of  the  sixth  month  of 
foetal  life,  but  Ecker  is  careful  to  point  out  that,  not  only  the  time,  but 
the  order,  of  their  appearance  is  subject  to  considerable  individual 
variation.  In  no  case,  however,  are  either  the  frontal  or  the  temporal 
sulci  the  earliest. 

The  fir;.t  which  appears,  in  fact,  lies  on  the  inner  face  of  the  hemi- 
sphere (whence  doubtless  Gratiolet,  who  does  not  seem  to  have  examined 
that  face  in  his  foetus,  overlooked  it),  and  is  either  the  internal  perpen- 
dicular (occipito-parietal),  or  the  calcarine  sulcus,  these  two  being  close 
together  and.  eventually  running  into  one  another.  As  a  rule  the 
occipito-parietal  is  the  earlier  of  the  two. 

3.  At  the  latter  part  of  this  period,  another  sulcus,  the  "  posterio, 
parietal,"  or  "  Fissure  of  Rolando "'  is  developed,  and  it  is  followed,  in 
the  course  of  the  sixth  month,  by  the  other  principal  sulci  of  the 
fiontal,  parietal,  tempoial  and  occipital  lobes.  There  is,  however,  no 
clear  evidence  that  one  of  these  constantly  appears  before  the  otlier; 
and  it  is  remarkable  that,  in  the  brain  at  the  period  described  and 
figured  by  Ecker  (I.  c.  p.  212-13,  Taf.  II.  figs.  1,  2,  3,  4),  the  antero- 
temporal  sulcus  {scissure  parallele)  so  characteristic  of  the  ape's  brain, 
IS  as  well,  if  not  better  developed  than  the  fissure  of  llolando,  and  is 
much  more  marked  than  the  proper  frontal  sulci. 

Taking  the  facts  as  they  now  stand,  it  appears  to  me  that  the  order 
of  the  appearance  of  the  sulci  and  gyri  in  the  foetal  human  brain  is  iu 
perfect  harmony  with  the  general  tloctrine  of  evolution,  and  with  the 


'*  '  Ueber  die  typische  Anordnung  ^^  '  Zur  Entwickelungs  Geschichte 

der   Furchen    und    Windungea   auf  der   Furchen    und    Windungen    der 

den      Grosshirn-Hemispharen       des  Grosshirn-Hemispharen    im    Foetus 

Menschen  und  der  Aften.'    'Archiv  des  MeHschen.'  '  Archiv  fiir  Anthro- 

fiir  Anthropologie,'  iii.,  1868.  pologie,'  iii.,  1868. 


Chap.  Y 1 1.  StructtLve  of  tJie  Brain.  205 


view  that  man  has  been  evolved  from  some  ape- like  form  ;  though  there 
can  be  no  doubt  that  that  form  was,  in  many  respects,  ditieient  from 
any  member  of  the  Frimates  now  living. 

Von  Buer  taught  us,  half  a  century  ago,  that,  in  the  course  of  their 
development,  allied  animals  put  on,  at  first,  the  characters  of  the  greater 
groups  to  which  they  belong,  and,  by  degrees,  assume  those  which  restrict 
them  within  the  limits  of  their  "family,  genus,  and  species ;  and  he 
proved,  at  the  same  time,  that  no  developmental  stage  of  a  higher 
ituimal  is  precisely  similar  to  the  adult  condition  of  any  lower  animal, 
it  is  quite  correct  to  say  that  a  frog  passes  through  the  condition  of  a 
fish,  inasmuch  as  at  one  period  of  its  life  the  tadpole  has  all  the  cha- 
racters of  a  fish,  and,  if  it  went  no  further,  would  have  to  be  grouped 
among  fishes.  But  it  is  equally  true  that  a  tadpole  is  very  different 
from  any  known  fish. 

In  like  manner,  the  brain  of  a  human  foetus,  at  the  fifth  month,  may 
correctly  be  said  to  be,  not  only  the  braiu  of  an  ape,  but  that  of  a:i 
Arctopitheciue  or  marmoset-like  ape ;  for  its  hemispheres,  with  their 
great  posterior  lobster,  and  with  no  sulci  but  the  sylvian  and  the 
calcarine,  present  the  characteristics  found  only  in  the  group  of  the 
Arctopilhecine  Primates.  But  it  is  equally  true,  as  Gratiolet  remarks, 
that,  in  its  widely  open  sylvian  fissure,  it  differs  from  the  biain  of  any 
actual  marmoset.  No  doubt  it  would  be  much  more  similar  to  the  brain 
of  an  advanced  foetus  of  a  marmoset.  But  we  know  nothing  whatever 
of  the  development  of  the  brain  in  the  marmosets.  In  the  Platyrhini 
proper,  the  only  observation  with  which  I  am  acquainted  is  due  to 
Pansch,  who  found  in  the  brain  of  a  foetal  Cehus  Apella,  in  addition  to 
the  sylvian  fissure  and  the  deep  calcarine  fissure,  only  a  very  shallow 
anterotemporal  fissure  (sc/ssare  parallele  of  Gratiolet.) 

Now  this  fact,  taken  together  with  the  circumstance  that  the  antnro- 
temporal  sulcus  is  present  in  such  Platyrhini  as  the  Saimiri,  which 
present  mere  traces  of  sulci  on  the  anterior  half  of  the  exterior  of  the 
cerebral  hemispheres,  or  none  at  all,  undoubtedly,  so  far  as  it  goes, 
affords  fair  evidence  in  favour  of  Graliolet's  hypothesis,  that  the 
])osterior  sulci  appear  before  the  anterior,  in  the  brains  of  the 
riatyrldui.  But,  it  by  no  means  follows,  that  the  rule  which  may  hold 
good  for  the  PlatyrJuni  extends  to  the  Catarhini.  We  have  no  in- 
formation whatever  respecting  the  development  of  the  brain  in  the 
Cyiunnorpha;  and,  as  regards  the  Anthropomorplta,  nothing  but  the 
account  of  the  brain  of  the  Gibbon,  near  biitli,  already  referred  to. 
At  the  present  moment,  there  is  not  a  shadow  of  evidence  to  shew 
that  the  sulci  of  a  chiiupanzee's,  or  orang's,  brain  do  not  appear  in  the 
same  order  as  a  man's. 

Gratiolet  opens  his  preface  with  the  aphorism.  "  II  est  dangereux 
"  dans  les  sciences  de  conclure  trop  vite."  I  fear  he  must  have  for- 
goiten  this  sound  maxim  by  the  time  he  had  reached  the  discussion  of 
the  differences  between  men  and  apes,  in  the  body  of  his  work.  No 
doubt,  the  excellent  author  of  one  of  the  most  remarkable  contributions 
to  the  just  understanding  of  the  mammalian  brain  which  has  ever  been 
made,  would  have  been  the  first  to  admit  the  insufficiency  of  his  data 
had  he  lived  to  profit  by  the  advance  of  inquiry.  The  misfortune  is 
tliat  his  conclusions  have  been  employed  by  persons  incompetent  to 
appreciate  their  foundation,  as  arguments  in  favour  of  obscurantism." 


*"  For  example,  !M.  I'Abbe  Lecomte      winisme  et   I'origine   de  THomme. 
ia  his  terrible  pamphlet  '  Le   Dar-      1873. 


2o6  T]ie  Descent  of  Man.  Part  I. 

But  it  is  important  to  remark  that,  whether  Gratiolet  was  right  or 
wrong  in- his  hypothesis  respecting  the  relative  order  of  appeaiance  of 
the  temporal  and  frontal  sulci,  the  fact  remains ;  that,  before  either 
temj)oral  or  frontal  sulci,  appear,  the  foetal  brain  of  man  presents 
characters  which  are  found  only  in  the  lowest  group  of  the  Primates 
(leaving  out  the  Lemurs) ;  and  that  this  is  exactly  what  we  shouLl 
expect  to  be  the  case,  if  man  has  resulted  from  the  gradual  modifica- 
tion of  the  same  form  as  that  from  which  the  other  Primates  have 
sprung. 


(      207       ) 


Part  LL 
SExXUAL  SELECTION. 


CHAPTEE  VIII. 

Principles  of  Sexual  Selection. 

Secondary  sexual  characters — Sexual  selection — Manner  of  action— Ex- 
cess of  males — Polygamy — The  male  alone  generally  modified  through 
sexual  selection — Eagerness  of  the  male — Variability  of  the  male — 
Choice  exerted  by  the  female — Sexual  compared  with  natural  selection 
— Inheritance,  at  corresponding  periods  of  life,  at  corresponding  seasons 
of  the  year,  and  as  limited  by  sex — Relations  between  the  several  forms 
of  inheritance — Causes  why  one  sex  and  the  young  are  not  modified 
through  sexual  selection — Supplement  on  the  proportional  numbers  of 
the  two  sexes  throughout  the  animal  kingdom — The  proportion  of  the 
sexes  in  i-elation  to  natural  selection. 

"^'iTH  animals  which  have  their  sexes  separated,  the  males 
necessarily  differ  from  the  females  in  their  organs  of  reproduction; 
and  these  are  the  primary  sexual  characters.  But  the  sexes 
often  differ  in  what  Hunter  has  called  secondary  sexual  charac- 
ters, which  are  not  directly  connected  with  the  act  of  reproduc- 
tion ;  for  instance,  the  male  possesses  certain  organs  of  sense  or 
locomotion,  of  which  the  female  is  quite  destitute,  or  has  them 
more  highly-developed,  in  order  that  he  may  readily  find  or 
reach  her ;  or  again  the  male  has  special  organs  of  prehension  for 
holding  her  securely.  These  latter  organs,  of  infinitely  diversified 
kinds,  graduate  into  those  which  are  commonly  ranked  as 
primary,  and  in  some  cases  can  hardly  be  distinguished  from 
them ;  we  see  instances  of  this  in  the  complex  appendages  at  the 
apex  of  the  abdomen  in  male  insects.  Unless  indeed  we  confine 
the  term  "primary"  to  the  reproductive  glands,  it  is  scarcely 
possible  to  decide  which  ought  to  be  called  primary  and  which 
secondary. 

The  female  often  differs  from  tlie  male  in  having  organs  for  the 
nourishment  or  protection  of  her  young,  such  as  the  mammary 


2o8  The  Descent  of  Man.  Part  II. 


glands  of  mammals,  and  tlie  abdominal  sacks  of  the  marsupials. 
In  some  few  cases  also  the  male  possesses  similar  organs,  which 
are  wanting  in  the  female,  snch  as  the  receptacles  for  the 
ova  in  certain  male  fishes,  and  those  temporarily  deyeloped  in 
certain  male  frogs.  The  females  of  most  bees  are  provided  with 
a  special  apparatus  for  collecting  and  carrying  pollen,  and  their 
ovipositor  is  modified  into  a  sting  for  the  defence  of  the  larvae 
and  the  community.  Many  similar  cases  could  be  given,  but 
they  do  not  here  concern  us.  There  are,  however,  other  sexual 
differences  quite  unconnected  with  the  primary  reproductive 
organs,  and  it  with  is  these  that  we  are  more  especially  concerned 
— such  as  the  greater  size,  strength,  and  pugnacity  of  the  male, 
his  weapons  of  offence  or  means  of  defence  against  rivals,  his 
gaudy  colouring  and  various  ornaments,  his  power  of  song,  and 
other  such  characters. 

Besides  the  primary  and  secondary  sexual  differences,  such  as 
the  foregoing,  the  males  and  females  of  some  animals  differ  in 
structures  related  to  different  habits  of  life,  and  not  at  all,  or 
only  indirectly,  to  the  reproductive  functions.  Thus  the  females 
of  certain  flies  (Cuhcidse  and  Tabanidse)  are  blood-suckers, 
whilst  the  males,  living  on  flowers,  have  mouths  destitute  of 
mandibles.^  The  males  of  certain  moths  and  of  some  crustaceans 
(e.  (J.  Tanais)  have  imperfect,  closed  mouths,  and  cannot  feed. 
The  complemeutal  males  of  certain  Cirripedes  live  like  epiphytic 
plants  either  on  the  female  or  the  hermaphrodite  form,  and  are 
destitute  of  a  mouth  and  of  prehensile  limbs.  In  these  cases  it  is 
the  male  which  has  been  modified,  and  has  lost  certain  important 
organs,  which  the  females  possess.  In  other  cases  it  is  the  female 
which  has  lost  such  parts;  for  instance,  the  female  glow- worm  is 
destitute  of  wings,  as  also  are  many  female  moths,  some  of  which 
never  leave  theii-  cocoons.  Many  female  parasitic  crustaceans 
have  lost  their  natatory  legs.  In  some  weevil-beetles  (Curcu- 
lionidse)  there  is  a  great  difference  between  the  male  and  female 
in  the  length  of  the  rostrum  or  snout ;  ^  but  the  meaning  of  this 
and  of  many  analogous  differences,  is  not  at  all  understood. 
Differences  of  structure  between  the  two  sexes  in  relation  to 
different  habits  of  life  are  generally  confined  to  the  lower 
animals ;  but  with  some  few  birds  the  beak  of  the  male  differs 
from  that  of  the  female.  In  the  Huia  of  New  Zealand  the 
difference  is  wonderfully  great,  and  we  hear  from  Dr.  Duller  ^ 

^  Westwood,    'Modern    Class,    of  -  Kirby  and    Spence,   '  Introduc- 

Insects,'  vol.  ii.  1840,  p.  541.     For  tion  to  Entomology,'  vol.  iii.  1826, 

the    statement    about  Tanais,  men-  p.  309. 

tioned  below,  I  am  indebted  to  Fritz  ^  '  Bii^jg  ^f  New  Zealand,'  1872, 

Miiller.  p.  66. 


Chap.  VIII.  Sexual  Selection.  209 

that  the  male  uses  his  strong  beak  in  chiselling  the  larvre  of 
insects  out  of  decayed  wood,  whilst  the  female  probes  the  softer 
parts  with  her  far  longer,  much  curved  and  pliant  beak :  and 
thus  they  mutually  aid  each  other.  In  most  cases,  differences  of 
structure  between  the  sexes  are  more  or  less  directly  connected 
with  the  propagation  of  the  species  :  thus  a  female,  which  has  to 
nourish  a  multitude  of  ova,  requires  more  food  than  the  male, 
and  consequently  requires  special  means  for  procuring  it.  A  male 
animal,  which  lives  for  a  very  short  time,  might  lose  its  organs 
for  procuring  food  through  disuse,  without  detriment ;  but  he 
would  retain  his  locomotive  organs  in  a  perfect  state,  so  that 
he  might  reach  the  female.  The  female,  on  the  other  hand, 
might  safely  lose  her  organs  for  flying,  swimming,  or  walking, 
if  she  gradually  acquired  habits  which  rendered  such  powers 
useless. 

We  are,  however,  here  concerned  only  with  sexual  selection. 
This  depends  on  the  advantage  which  certain  individuals  have 
over  others  of  the  same  sex  and  species  solely  in  respect  of 
reproduction.  When,  as  in  the  cases  above  mentioned,  the  two 
sexes  differ  in  structure  in  relation  to  different  habits  of  life, 
they  have  no  doubt  been  modified  through  natural  selection,  and 
by  inheritance  limited  to  one  and  the  same  sex.  So  again  the 
primary  sexual  organs,  and  those  for  nourishing  or  protecting  the 
young,  come  under  the  same  influence ;  for  those  individuals  which 
generated  or  nourished  their  offspring  best,  would  leave,  cctteris 
paribus,  the  greatest  number  to  inherit  their  superiority  ;  whilst 
those  which  generated  or  nourished  their  offspring  badly,  would 
leave  but  few  to  inherit  their  weaker  powers.  As  the  male  has  to 
find  the  female,  he  requires  organs  of  sense  and  locomotion,  but 
if  these  organs  are  necessary  for  the  other  purposes  of  life,  as  is 
generally  the  case,  they  will  have  been  developed  through 
natural  selection.  When  the  male  has  found  the  female,  he 
sometimes  absolutely  requires  prehensile  organs  to  hold  her; 
thus  Dr.  Wallace  informs  me  that  the  males  of  certain  moths 
cannot  unite  with  the  females  if  their  tarsi  or  feet  are  broken. 
The  males  of  many  oceanic  crustaceans,  when  adult,  have  their 
legs  and  antennae  modified  in  an  extraordinary  manner  for  the . 
prehension  of  the  female ;  hence  we  may  suspect  that  it  is  be- 
cause these  animals  are  washed  about  by  the  waves  of  the 
open  sea,  that  they  require  these  organs  in  order  to  propagate 
their  kind,  and  if  so,  their  development  has  been  the  result  of 
ordinary  or  natural  selection.  Some  animals  extremely  low  in 
the  scale  have  been  modified  for  this  same  purpose ;  thus  the 
males  of  certain  parasitic  worms,  when  fully  grown,  have  the 
lower  surface  of  the  terminal  part  of  their  bodies  roughened 


2  lo  The  Descent  of  Alan.  Pakt  II, 


like  a  rasp,  and  with  this  they  coil  round  and  permanently  hold 
the  females.* 

•  When  the  two  sexes  follow  exactly  the  same  habits  of  life,  and 
the  male  has  the  sensory  or  locomotive  organs  more  highly 
developed  than  those  of  the  female,  it  may  be  that  the  perfection 
of  these  is  indispensable  to  the  male  for  finding  the  female  ;  but 
in  the  vast  majority  of  cases,  they  serve  only  to  give  one  male 
an  advantage  over  another,  for  with  sufficient  time,  the  less  well- 
endowed  males  would  succeed  in  pairing  with  the  females ;  and 
judging  from  the  structure  of  the  female,  they  would  be  in  all 
other  respects  equally  well  adapted  for  their  ordinary  habits  of 
life.  Since  in  such  cases  the  males  have  acquired  their  present 
structure,  not  from  being  better  fitted  to  survive  in  the  struggle 
for  existence,  but  from  having  gained  an  advantage  over  other 
males,  and  from  having  transmitted  this  advantage  to  their  male 
offspring  alone,  sexual  selection  must  here  have  come  into  action. 
It  was  the  importance  of  this  distinction  which  led  me  to 
designate  this  form  of  selection  as  Sexual  Selection.  So  again, 
if  the  chief  service  rendered  to  the  male  by  his  prehensile  organs 
is  to  prevent  the  escape  of  the  female  before  the  arrival  of  other 
males,  or  when  assaulted  by  them,  these  organs  '^ill  have  been 
perfected  through  sexual  selection,  that  is  by  the  advantage 
acquired  by  certain  individuals  over  their  rivals.  But  in  most 
cases  of  this  kind  it  is  impossible  to  distinguish  between  the 
effects  of  natural  and  sexual  selection.  Whole  chapters  could 
be  filled  with  details  on  the  differences  between  the  sexes  in  their 
sensory,  locomotive,  and  prehensile  organs.  As,  however,  these 
structures  are  not  more  interesting  than  others  adapted  for  the 
ordinary  purposes  of  life  I  shall  pass  them  over  almost  entirely, 
giving  only  a  few  instances  under  each  class. 

There  are  many  other  structures  and  instincts  which  must 
have  been  developed  through  sexual  selection— such  as  the 
weapons  of  offence  and  the  means  of  defence  of  the  males  for 
fighting  with  and  driving  away  their  rivals— their  courage  and 
pugnacity — their  various  ornaments — their  contrivances  for  pro- 

*  M.  Perrier  advances  this  case  claspers  of  certain  male  animals 
('Revue  Scientifique,' Feb.  1,  1873,  could  not  have  been  developed 
p.  865)  as  one  fatal  to  the  belief  in  through  the  choice  of  the  female  I 
sexual  selection,  inasmuch  as  he  Had  J  not  met  with  this  remark,  I 
supposes  that  I  attribute  all  the  should  not  have  thought  it  possible 
differences  between  the  sexes  to  for  any  one  to  have  read  this  chapter 
sexual  selection.  This  distinguished  and  to  have  imagined  that  I  main- 
naturalist,  therefore,  like  so  many  tain  that  the  choice  of  the  female 
other  Frenchmen,  has  not  taken  the  had  anything  to  do  with  the  develop- 
trouble  to  understand  even  the  first  meut  of  the  prehensile  organs  in  the 
principles  of  sexual  selection.  An  male. 
English  naturalist  insists   that  the 


Chap.  VII  r.  Sexual  Selection.  211 


duciiig  vocal  or  instrumental  music — and  their  glands  for 
emitting  odours,  most  of  these  latter  structures  serving  only  to 
allure  or  excite  the  female.  It  is  clear  that  these  characters  are 
the  result  of  sexual  and  not  of  ordinary  selection,  since  unarmed, 
unornamented,  or  unattractive  males  would  succeed  equally  well 
in  the  battle  for  life  and  in  leaving  a  numerous  jDrogeny,  but  for 
the  presence  of  better  endowed  males.  We  may  infer  that  this 
would  be  the  case,  because  the  females,  which  arc  unarmed  and 
unornamented,  are  able  to  survive  and  procreate  their  kind. 
Secondary  sexual  characters  of  the  kind  just  referred  to,  will  be 
fully  discussed  in  the  following  chapters,  as  being  in  many 
respects  interesting,  but  especially  as  depending  on  the  will, 
choice,  and  rivalry  of  the  individuals  ot  either  sex.  When  we 
behold  two  males  fighting  for  the  possession  of  the  female,  or 
several  male  birds  displaying  their  gorgeous  plumage,  and  per- 
forming strange  antics  before  an  assembled  body  of  females,  we 
cannot  doubt  that,  though  led  by  instinct,  they  know  what  they 
are  about,  and  consciously  exert  then'  mental  and  bodily  powers. 

Just  as  man  can  improve  the  breed  of  his  game-cocks  by  the 
selection  of  those  birds  which  are  victorious  in  the  cockpit,  so  it 
appears  that  the  strongest  and  most  vigorous  males,  or  those 
provided  with  the  best  weapons,  have  prevailed  under  nature, 
and  have  led  to  the  improvement  of  the  natural  breed  or  species. 
A  slight  degree  of  variability  leading  to  some  advantage,  how- 
ever slight,  in  reiterated  deadly  contests  would  suffice  for  the 
work  of  sexual  selection;  and  it  is  certain  that  secondary  sexual 
characters  are  eminently  variable.  Just  as  man  can  give  beauty, 
according  to  his  standard  of  taste,  to  his  male  poultry,  or  more 
strictly  can  modify  the  beauty  originally  acquired  by  the  parent 
species,  can  give  to  the  Sebright  bantam  a  new  and  elegant 
plumage,  an  erect  and  peculiar  carriage— so  it  appears  that 
female  bii'ds  in  a  state  of  nature,  have  by  a  long  selection  of  the 
more  attractive  males,  added  to  their  beauty  or  other  attractive 
quahties.  No  doubt  this  implies  powers  of  discrimination  and 
taste  on  the  part  of  the  female  wliich  will  at  first  appear 
extremely  improbable;  but  by  the  facts  to  be  adduced  here- 
after, I  hope  to  be  able  to  shew  that  the  females  actually 
have  these  powers.  When,  however,  it  is  said  that  the  lower 
animals  have  a  sense  of  beauty,  it  must  not  be  supposed  that 
such  sense  is  comparable  with  that  of  a  cultivated  man,  with  his 
multiform  and  complex  associated  ideas.  A  more  just  com- 
parison would  be  between  the  taste  for  the  beautiful  in  animals, 
and  that  in  the  lowest  savages,  who  admire  and  deck  themselves 
with  any  brilliant,  glittering,  or  curious  object. 

From  our  ignorance  on  several  points,  the  precise  manner  in 


212  The  Descent  of  Man.  Part  IL 

which  sexual  selection  acts  is  somewhat  "uncertain.  Neverthe- 
less if  those  naturalists  who  already  believe  in  the  mutability  of 
species,  will  read  the  following  chapters,  they  will,  I  think,  agree 
with  me,  that  sexual  selection  has  played  an  important  part  in 
the  history  of  the  organic  world.  It  is  certain  that  amongst 
almost  all  animals  there  is  a  struggle  between  the  males  for  the 
possession  of  the  female.  This  fact  is  so  notorious  that  it  would 
be  superfluous  to  give  instances,  Hence  the  females  have  the 
opportunity  of  selecting  one  out  of  several  males,  on  the  suppo- 
sition that  their  mental  capacity  suffices  for  the  exertion  of  a 
choice.  In  many  cases  special  circumstances  tend  to  make  the 
struggle  between  the  males  particularly  severe.  Thus  the  males 
of  our  migratory  birds  generally  arrive  at  their  places  of  breeding 
before  the  females,  so  that  many  males  are  ready  to  contend  for 
each  female.  I  am  informed  by  Mr.  Jenner  Weir,  that  the  bird- 
catchers  assert  that  this  is  invariably  the  case  with  the  nightin- 
gale and  blackcap,  and  with  respect  to  the  latter  he  can  himself 
confirm  the  statement. 

Mr.  Swaysland  of  Brighton  has  been  in  the  habit,  during  the 
last  forty  years,  of  catching  our  migratory  birds  on  their  first 
arrival,  and  he  has  never  known  the  females  of  any  species  to 
arrive  before  their  males.  During  one  spring  he  shot  thirty -nine 
males  of  Eay's  wagtail  {Budytes  Rail)  before  he  saw  a  single 
female.  Mr.  Gould  has  ascertained  by  the  dissection  of  those 
snipes  which  arrive  the  first  in  this  country,  that  the  males  come 
before  the  females.  And  the  like  holds  good  with  most  of  the 
migratory  birds  of  the  United  States.^  The  majority  of  the  male 
salmon  in  our  rivers,  on  coming  up  from  the  sea,  are  ready  to 
breed  before  the  females.  So  it  appears  to  be  with  frogs  and 
toads.  Throughout  the  great  class  of  insects  the  males  almost 
always  are  the  first  to  emerge  from  the  pupal  state,  so  that  they 
generally  abound  for  a  time  before  any  females  can  be  seen.*^ 
The  cause  of  tliis  difference  between  the  males  and  females  in 
their  periods  of  arrival  and  maturity  is  sufficiently  obvious. 
Those  males  which  annually  first  migrated  into  any  country,  or 
T\^hich  in  the  spring  were  first  ready  to  breed,  or  were  the  most 
eager,  would  leave  the  largest  number  of  ofl"spring ;  and  these 

'  J.  A.  Allen,  on  the  'Mammals  rodite  plants  are  dichogamous ;  that 

and  Winter  Birds  of  Florida,'  Bull.  is,  their  male  and  female  organs  are 

Comp.  Zoology,  Harvard  College,  p.  not  ready  at  the  same  time,  so  that 

268.  they  cannot  be  self-fertilised.     Now 

^  Even  with  those  plants  in  which  in    such    flowers,   the    pollen    is    in 

the    sexes    are    separate,    the    male  general  matured  before  the  stigma, 

flowers    are    generally    mature    be-  though  there  are  exceptional  cases 

fore    the    fernale.      As   first   shewn  in    Avhich    the    female    organs   are 

by  C.  K.  Sprengel,  many  hermaph-  beforehaad. 


Chap.  VIII.  Sexual  Selection.  213 


would  tend  to  inherit  similar  instincts  and  constitutions.  It 
must  be  borne  in  mind  that  it  would  have  been  impossible  to 
change  very  materially  the  time  of  sexual  maturity  in  the 
females,  without  at  the  same  time  interfering  with  the  period  of 
the  production  of  the  young— a  period  which  must  bo  determined 
by  the  seasons  of  the  year.  On  the  whole  there  can  be  no  doubt 
that  with  almost  all  animals,  in  which  the  sexes  are  separate, 
there  is  a  constantly  recurrent  struggle  between  the  males  for 
the  possession  of  the  females. 

Our  difficulty  in  regard  to  sexual  selection  lies  in  understand- 
ing how  it  is  that  the  males  which  conquer  other  males,  or  those 
which  prove  the  most  attractive  to  the  females,  leave  a  greater 
number  of  offspring  to  inherit  their  superiority  than  their 
beaten  and  less  attractive  rivals.  Unless  this  result  does  follow, 
the  charactei-s  which  give  to  certain  males  an  advantage  over 
others,  could  not  be  perfected  and  augmented  through  sexual 
selection.  When  the  sexes  exist  in  exactly  equal  numbers,  the 
vrorst-endowed  males  will  (except  where  polygamy  prevails), 
ultimately  find  females,  and  leave  as  many  offspring,  as  well 
fitted  for  theu'  general  habits  of  life,  as  the  best-endowed  males. 
From  various  facts  and  considerations,  I  formerly  inferred  that 
with  most  animals,  in  which  secondary  sexual  characters  are 
well  developed,  the  males  considerably  exceeded  the  females  in 
number;  but  this  is  not  by  any  means  always  true.  If  the 
males  were  to  the  females  as  two  to  one,  or  as  three  to  two,  or 
even  in  a  somewhat  lower  ratio,  the  whole  affair  would  be 
simple;  for  the  better-armed  or  more  attractive  males  would 
leave  the  largest  number  of  offspring.  But  after  investigating, 
as  far  as  possible,  the  numerical  proportion  of  the  sexes,  I  do  not 
beheve  that  any  great  inequality  in  number  commonly  exists. 
In  most  cases  sexual  selection  appears  to  have  been  effective  in 
the  following  manner. 

Let  us  take  any  species,  a  bird  for  instance,  and  divide  the 
females  inhabiting  a  district  into  two  equal  bodies,  the  one 
consisting  of  the  more  vigorous  and  better-nourished  individuals, 
and  the  other  of  the  less  vigorous  and  healthy.  The  former, 
there  can  be  little  doubt,  would  be  ready  to  breed  in  the  spring 
before  the  others ;  and  this  is  the  opinion  of  ]\Ir.  Jenncr  Weir, 
who  hasT^arefully  attended  to  the  habits  of  birds  during  many 
years.  There  can  also  be  no  doubt  that  the  most  vigorous, 
best-nourished  and  earhest  breeders  would  on  an  average 
succeed  in  rearing  the  largest  number  of  fine  offspring.'^  The 
males,  as  we  have  seen,  are  generally  ready  to  breed  before  the 

*  Here  is  excellent  evidence  on  an  experienced  ornithologist.  Mr. 
the  character  of  the  ofispring  from       J.  A.  Allen,  in  speaking  ('Mammals 


214  The  Descent  of  Man.  Part  I L 

females ;  the  strongest,  and  with  some  species  the  best  armed  of 
the  males,  drive  away  the  weaker  ;  and  the  former  would  then 
unite  with  the  more  vigorous  and  better-nourished  females,  be- 
cause they  are  the  first  to  breed.^  Such  vigorous  pairs  would 
surely  rear  a  larger  number  of  offspring  than  the  retarded 
females,  which  would  be  compelled  to  unite  with  the  conquered 
and  less  powerful  males,  supposing  the  sexes  to  be  numerically 
equal ;  and  this  is  all  that  is  wanted  to  add,  in  the  course  of 
successive  generations,  to  the  size,  strength  and  courage  of  the 
males,  or  to  improve  their  weajDons. 

But  in  very  many  cases  the  males  which  conquer  their  rivals, 
do  not  obtain  possession  of  the  females,  independently  of  the 
choice  of  the  latter.  The  courtship  of  animals  is  by  no  means 
so  simple  and  short  an  affair  as  might  be  thought.  The 
females  are  most  excited  by,  or  prefer  pairing  with,  the  more 
ornamented  males,  or  those  which  are  the  best  songsters,  or  play 
the  best  antics;  but  it  is  obviously  probable  that  they  would 
at  the  same  time  prefer  the  more  vigorous  and  lively  males,  and 
this  has  in.  some  cases  been  confirmed  by  actual  observation.^ 
Thus  the  more  vigorous  females,  which  are  the  first  to  breed,  will 
have  the  choice  of  many  males  ;  and  though  they  may  not  always 
select  the  strongest  or  best  armed,  they  will  select  those  which 
are  vigorous  and  well  armed,  and  in  other  respects  the  most  at- 
tractive. Both  sexes,  therefore,  of  such  early  pairs  would  as  above 
explained,  have  an  advantage  over  others  in  rearing  offspring ;  and 
this  apparently  has  suflficed  during  a  long  course  of  generations 
to  add  not  only  to  the  strength  and  fighting  powers  of  the  males, 
but  Hkewise  to  their  various  ornaments  or  other  attractions. 

In  the  converse  and  much  rarer  case  of  the  males  selecting 
particular  females,  it  is  plain  that  those  which  were  the  most 
vigorous  and  had  conquered  others,  would  have  the  freest 
choice  ;  and  it  is  almost  certain  that  they  would  select  vigorous 
as  well  as  attractive  females.     Such  pairs  would  have  an  advan- 


and    Winter    Birds    of  E.    Florida,'  to  those  female  bees  which  are  the 

p.  229)  of  the  later  broods,  after  the  first  to  emerge  from  the   puj^a  each 

accidental  destruction  of   the  first,  year.      See     his    remarkable    essaj-, 

bays,  that  these  "  are   found   to  be  'Anwendung  den  Darwin'schen  Lehra 

"  smaller    and  paler-coloured    than  auf  Bienen,'  '  Verh.    d.  ^ .  Jahrg.' 

"  those  hatched  earlier  in  the  sea-  xxix.  p.  45. 

*'  son.    In  cases  where  several  broods  ^  With  respect  to  poultry,  I  have 

"  are  reared  each  year,  as  a  general  received    information,    hereafter    to 

"  rule  the  birds  of  the  earlier  broods  be  given,  to  this  effect.     Even  with 

"  seem  iji  all  respects  the  most  per-  birds,  such   as  pigeons,  which  pair 

*'  feet  and  vigorous."  for  life,  the  female,  as  I  hear  from 

^  Hermann    Miiller  has    come    to  Mr.  Jenner  Weir,  will    desert   her 

this    same    conclusion  with   respect  mate  if  he  is  injured  or  grows  weak. 


Uh.\  1'.  VIII.  Scxnal  Selection.  2 1 5 


tage  in  rearing  offspring,  more  especially  if  the  male  had  the 
power  to  defend  the  female  during  the  pairing-season  as  occurs 
with  some  of  the  higher  animals,  or  aided  her  in  providing  for 
the  young.  The  same  principles  would  apply  if  each  sex  pre- 
ferred and  selected  certain  individuals  of  the  opposite  sex ; 
supposing  that  they  selected  not  only  the  more  attractive,  but 
likewise  the  more  vigorous  individuals. 

Numerical  Proportion  of  the  Two  Sexes. — I  have  remarked  that 
sexual  selection  would  be  a  simple  affair  if  the  males  were  con- 
siderably more  numerous  than  the  females.  Hence  I  was  led  to 
investigate,  as  far  as  I  could,  the  proportions  between  the  two 
sexes  of  as  many  animals  as  possible;  but  the  materials  are 
scanty.  I  will  here  give  only  a  brief  abstract  of  the  results, 
retaining  the  details  for  a  sujiplementary  discussion,  so  as  not 
to  interfere  with  the  course  of  my  argument.  Domesticated 
animals  alone  afford  the  means  of  ascertaining  the  propor- 
tional numbers  at  birth;  but  no  records  have  been  specially 
kept  for  this  purpose.  By  indirect  means,  however,  I  have 
collected  a  considerable  body  of  statistics,  from  which  it  appears 
that  with  most  of  our  domestic  animals  the  sexes  are  nearly 
equal  at  birth.  Thus  25,560  births  of  race-horses  have  been 
recorded  during  twenty-one  years,  and  the  male  births  were 
to  the  female  births  as  99"7  to  100.  In  greyhounds  the  in- 
equality is  greater  than  with  any  other  animal,  for  out  of  6878 
births  dui-ing  twelve*  years,  the  male  births  were  to  the  female 
as  110-1  to  100.  It  is,  however,  in  some  degree  doubtful 
whether  it  is  safe  to  infer  that  the  proportion  would  be  the  same 
under  natural  conditions  as  under  domestication ;  for  slight  and 
unknown  differences  in  the  conditions  affect  the  proi^ortion  of 
the  sexes.  Thus  with  mankind,  the  male  births  in  England 
are  as  101'5,  in  Russia  as  108'9,  and  with  the  Jews  of  Livonia  as 
120,  to  100  female  births.  But  I  shall  recur  to  this  curious  point 
of  the  excess  of  male  births  in  the  supplement  to  this  chapter.  At 
the  Cape  of  Good  Hope,  however,  male  children  of  Euroj^ean 
extraction  have  been  born  during  several  years  in  the  proportion 
of  between  90  and  99  to  100  female  children. 

For  our  present  purpose  we  are  concerned  with  the  proportion 
of  the  sexes,  not  only  at  birth,  but  also  at  maturity,  and  this 
adds  another  element  of  doubt ;  for  it  is  a  well-ascertained  fact 
that  with  man  the  number  of  males  dying  before  or  during  birth, 
and  during  the  first  few  years  of  infancy,  is  considerably  larger 
than  that  of  females.  So  it  almost  certainly  is  with  male  lambs, 
and  probably  with  some  other  animals.  The  males  of  some  species 
kill  one  another  by  fighting ;  or  they  drive  one  another  about 


2 1 6  The  Descent  of  Man.  Tart  II. 

until  they  become  greatly  emaciated.  Tliey  must  also  be  often 
exposed  to  yarious  dangers,  whilst  wandering  about  in  eager 
search  for  the  females.  In  many  kinds  of  fish  the  males  are- 
much  smaller  than  the  females,  and  they  are  beheved  often  to  be 
devoured  by  the  latter,  or  by  other  fishes.  The  females  of 
some  birds  appear  to  die  earlier  than  the  males;  they  are 
also  liable  to  be  destroyed  on  their  nests,  or  whilst  in  cliarge 
of  their  young.  With  insects  the  female  laryse  are  often  larger 
than  those  of  the  males,  and  w^ould  consequently  be  more  likely 
to  be.  devoured.  In  some  cases  the  mature  females  are  less 
active  and  less  rapid  in  their  movements  than  the  males,  and 
could  not  escape  so  well  from  danger.  Hence,  with  animals  in  a 
state  of  nature,  wo  must  rely  on  mere  estimation,  in  order  to 
judge  of  the  proportions  of  the  sexes  at  maturity ;  and  this  is 
but  little  trustworthy,  except  when  the  inequality  is  strongly 
marked.  Nevertheless,  as  far  as  a  judgment  can  be  formed,  we 
may  conclude  from  the  facts  given  in  the  supplement,  that  the 
males  of  some  few  mammals,  of  many  birds,  of  some  fish  and 
insects,  are  considerably  more  numerous  than  the  females. 

The  proportion  between  the  sexes  fluctuates  slightly  during 
successive  years :  thus  with  race-horses,  for  every  100  mares  born 
the  stallions  varied  from  107'1  in  one  year  to  92-6  in  another  year, 
and  with  greyhounds  from  116"3  to  95*3.  But  had  larger  num- 
bers been  tabulated  throughout  an  area  more  extensive  than 
England,  these  fluctuations  would  probably  have  disappeared ; 
and  such  as  they  are,  would  hardly  suffice  to  lead  to  efi'ective 
sexual  selection  in  a  state  of  nature.  Nevertheless,  in  the  cases 
of  some  few  wild  animals,  as  shewn  in  the  supi3lement,  the 
proportions  seem  to  fluctuate  either  during  different  seasons 
or  in  different  localities  in  a  sufficient  degree  to  lead  to  such 
selection.  For  it  should  be  observed  that  any  advantage, 
gained  during  certain  years  or  in  certain  localities  by  those  males 
which  were  able  to  conquer  their  rivals,  or  were  the  most 
attractive  to  the  females,  would  probably  be  transmitted  to  the 
offspring,  and  would  not  subsequently  be  eliminated.  During 
the  succeeding  seasons,  when,  from  the  equality  of  the  sexes, 
every  male  was  able  to  procure  a  female,  the  stronger  or  more  at- 
tractive males  previously  produced  would  si  ill  have  at  least  as 
good  a  chance  of  leaving  offspring  as  the  weaker  or  less  attractive. 

Polygamy. — The  practice  of  polygamy  leads  to  the  same  results 
as  would  follow  from  an  actual  inequality  in  the  number  of  the 
sexes ;  for  if  each  male  secures  two  or  more  females,  many  males 
cannot  pair  ;  and  the  latter  assuredly  will  be  the  weaker  or  less 
attractive  individuals.     Many  mammals  and  some  few  birds  are 


Chap.  VI II.  Sexual  Selection.  217 

polygamous,  but  with  animals  belonging  to  the  lower  classes  I 
have  found  no  evidence  of  this  habit.  The  intellectual  powers 
of  such  animals  are,  perhaps,  not  sufificicnt  to  lead  them  to 
collect  and  guard  a  harem  "of  females.  That  some  relation  exists 
between  polygamy  and  the  development  of  secondary  sexual 
characters,  appears  nearly  certain ;  and  this  supports  the  view 
that  a  numerical  preponderance  of  males  would  be  eminently 
favourable  to  the  action  of  sexual  selection.  Nevertheless  many 
animals,  which  are  strictly  monogamous,  especially  birds,  display 
strongly-marked  secondary  sexual  characters ;  whilst  some  few 
animals,  which  are  polygamous,  do  not  have  such  characters. 

\Ye  will  jfirst  briefly  run  through  the  mammals,  and  then  turn 
to  birds.  The  gorilla  seems  to  be  polygamous,  and  the  male 
differs  considerably  from  the  female ;  so  it  is  with  some  baboons, 
which  live  in  herds  containing  twice  as  many  adult  females  as 
males.  In  South  America  the  Mycetts  caraya  presents  well- 
marked  sexual  differences,  in  colour,  beard,  and  vocal  organs ; 
and  the  male  generally  lives  with  two  or  three  wives :  the  male 
of  the  Cehus  capucinus  differs  somewhat  from  the  female,  and 
appears  to  be  polygamous.^*^  Little  is  known  on  this  head  with 
respect  to  most  other  monkeys,  but  some  species  are  strictly 
monogamous.  The  ruminants  are  eminently  polygamous,  and 
they  present  sexual  differences  more  frequently  than  almost  any 
other  group  of  mammals;  this  holds  good,  especially  in  their 
weapons,  but  also  in  other  characters.  Most  deer,  cattle,  and 
sheep  are  polygamous ;  as  are  most  antelopes,  though  some  are 
monogamous.  Sir  Andrew  Smith,  in  speaking  of  the  antelopes 
of  South  Africa,  says  that  in  herds  of  about  a  dozen  there  was 
rarely  more  than  one  mature  male.  The  Asiatic  Amilope  saiga 
appears  to  be  the  most  inordinate  polygamist  in  the  world  ;  for 
Pallas ^^  states  that  the  male  drives  away  all  rivals,  and  collects  a 
herd  of  about  a  hundred  females  and  kids  together ;  the  female 
is  hornless  and  has  softer  hair,  but  does  not  otherwise  differ 
much  from  the  male.  The  wild  horse  of  the  Falkland  Islands  and 
of  the  Western  States  of  N.  America  is  polygamous,  but,  except 
in  his  greater  size  and  in  the  proportions  of  his  body,  differs  but 
little  from  the  mare.  The  wild  boar  presents  well-marked  sexual 

"  On    the    Gorilla,    Savage    and  Fasc.  xii.  1777,  p.  29,     Sir  Andrew 

Wyman.   '  Boston   Journal    of  Nat.  Smith,  '  Illustrations  of  the  Zoology 

Hist.*  vol.  V.  1845-47,  p.  423.     On  of  S.  Africa,'  1849,  pi.  29,  on   the 

Cynocephalus,  Brehm, 'lUust.Thier-  Kobus.     Owen,  in  his  '  Anatomy  of 

leben,'  B.  i.   1864,  s.  77.     On  My-  Vertebrates'  (vol.  iii.  1868,  p.  6":33) 

cetes,  Rengger, 'Xaturgesch.:  Siiuge-  gives  a   table  shewing   incidentally 

thiere  von   Paraguay,'  1830,  s.   14,  which  species  of  antelopes  are  gre- 

20.     Cebus,  Brehm, 'ibid.  s.  108,  garious. 

"  Pallas,      'S])icilegia     Zoolog.,' 


2 1 8  The  Descent  of  Man.  Pakt  II. 

characters,  in  his  gi'eat  tusks  and  some  other  points.  In  Europe 
and  in  India  he  leads  a  solitary  hfe,  except  during  the  breeding- 
season  ;  but  as  is  beUeved  by  Sir  W.  Elliot,  who  has  had  many 
opportunities  in  India  of  observing  this  animal,  he  consorts  at 
this  season  "with  several  females.  "Whether  this  holds  good 
in  Europe  is  doubtful,  but  it  is  supported  by  some  evidence. 
The  adult  male  Indian  elephant,  like  the  boar,  passes  much  of 
Ms  time  in  solitude ;  but  as  Dr.  Campbell  states,  when  with 
others,  "  it  is  rare  to  find  more  than  one  male  with  a  whole  herd 
"  of  females ;"  the  larger  males  expelling  or  killing  the  smaller 
and  weaker  ones.  The  male  differs  from  the  female  in  his  immense 
tusks,  greater  size,  strength,  and  endurance;  so  great  is  the 
difference  in  these  respects,  that  the  males  when  caught  are 
valued  at  one-fifth  more  than  the  females.^^  The  sexes  of  other 
pachydermatous  animals  differ  very  little  or  not  at  all,  and,  as 
far  as  known,  they  are  not  polygamists.  Nor  have  I  heard  of  any 
species  in  the  Orders  of  Cheiroptera,  Edentata,  Insectivora  and 
Piodents  being  polygamous,  excepting  that  amongst  the  Eodents, 
the  common  rat,  according  to  some  rat-catchers,  lives  with  several 
females.  Nevertheless  the  two  sexes  of  some  sloths  (Edentata) 
differ  in  the  character  and  colour  of  certain  patches  of  hair  on 
their  shoulders.^^  And  many  kinds  of  bats  (Chehoptera)  present 
well-marked  sexual  differences,  chiefly  in  the  males  possessing 
odoriferous  glands  and  pouches,  and  by  their  being  of  a  lighter 
colour.^^  In  the  great  order  of  Eodents,  as  far  as  I  can  learn, 
the  sexes  rarely  differ,  and  when  they  do  so,  it  is  but  shghtly  in 
the  tint  of  the  fur. 

As  I  hear  from  Sir  Andrew  Smith,  the  lion  in  South  Africa 
sometimes  lives  with  a  single  female,  but  generally  with  more, 
and,  in  one  case,  was  found  with  as  many  as  five  females ;  so 
that  he  is  polygamous.  As  far  as  I  can  discover,  he  is  the  only 
polygamist  amongst  all  the  terrestrial  Carnivora,  and  he  alone 
presents  well-marked  sexual  characters.  If,  however,  we  turn 
to  the  marine  Carnivora,  as  we  shall  hereafter  see,  the  case  is 
widely  different ;  for  many  species  of  seals  offer  extraordinary 
sexual  differences,  and  they  are  eminently  polygamous.  Thus, 
according  to  Peron,  the  male  sea-elephant  of  the  Southern  Ocean 
always  possesses  sever-ai  females,  and  the  sea-lion  of  Forster  is 
said  to  be  surrounded  by  from  twenty  to  thirty  females.  In  the 
North,  the  male  sea-bear  of  Steller  is  accompanied  by  even  a 

^2  Dr.  Campbell,  in   '  Proc.  Zoo-  '    '^   Dr.     Gray,    in    *  Annals    and 

log.  Soc'  1869,  p.  138.     See  also  an  Mag.  of  Nat.  Hist.'  1871,  p.  302. 

interesting   paper,  by  Lieut.  John-  ^'^    See     Dr.    Dobson's     excellent 

stone,    in    '  Proc.    Asiatic    Soc.    of  paper,  in  '  Proc.  Zoolog.  Soc'  1873, 

Bengal,'  May,  1868.  p.  ii41. 


CiiAP.  VIII.  Sexual  Selection.  219 

greater  number  of  females.  It  is  an  interesting  fact,  as  Dr. 
Gill  remarks,^^  that  in  the  monogamous  species,  "  or  those 
"  living  in  small  communities,  there  is  little  difference  in  size 
"  between  the  males  and  females ;  in  tlie  social  species,  or  rather 
"  those  of  which  the  males  have  harems,  the  males  are  vastly 
"  larger  than  the  females." 

Amongst  birds,  many  species,  the  sexes  of  which  differ  greatly 
from  each  other,  are  certainly  monogamous.  In  Great  Britain 
we  see  well-marked  sexual  differences,  for  instance,  in  the  wild- 
duck  which  pairs  with  a  single  female,  the  common  blackbird, 
and  the  bullfinch  which  is  said  to  pair  for  life.  I  am  informed 
by  Mr.  Wallace  that  the  like  is  true  of  the  Chatterers  or 
Cotingidcie  of  South  America,  and  of  many  other  birds.  In  several 
groups  I  have  not  been  able  to  discover  whether  the  species  are 
polygamous  or  monogamous.  Lesson  says  that  birds  of  paradise, 
so  remarkable  for  their  sexual  differences,  are  polygamous,  but  Mr. 
Wallace  doubts  wdiether  he  had  sufficient  evidence.  IMr.  Salviu 
tells  me  he  has  been  led  to  believe  that  humming-bii'ds  are 
polygamous.  The  male  widow-bird,  remarkable  for  his  caudal 
plumes,  certainly  seems  to  be  a  polygamist.^^  I  have  been 
assured  by  Mr.  Jenner  Weir  and  by  others,. that  it  is  somewhat 
common  for  three  starlings  to  frequent  the  same  nest ;  but 
whether  this  is  a  case  of  polygamy  or  polyandry  has  not  been 
ascertained. 

The  Gaihnacese  exhibit  almost  as  strongly  marked  sexual 
differences  as  birds  of  paradise  or  humming-birds,  and  many  of 
the  species  are,  as  is  well  known,  polygamous ;  others  being 
strictly  monogamous.  What  a  contrast  is  presented  between  the 
sexes  of  the  polygamous  peacock  or  pheasant,  and  the  mono- 
gamous guinea-fowl  or  partridge !  Many  similar  cases  could  be 
given,  as  in  the  grouse  tribe,  in  which  the  males  of  the  poly- 
gamous capercailzie  and  black-cock  differ  greatly  from  the 
females;  whilst  the  sexes  of  the  monogamous  red  grouse  and 
ptarmigan  differ  very  little.  In  the  Cursores,  except  amongst 
the  bustards,  few  species  offer  strongly-marked  sexual  dif- 
ferences, and  the  great  bustard  {0th  tarda)  is  said  to  be  poly- 
gamous. With  the  Grallatores,  extrem.ely  few  species  differ 
sexually,  but    the  ruff  {Machetes  jniunax)    affords    a   marked 

"  The    Eared    Seals,    '  American  Great  Bustard,  see  L.  Lloyd,  '  Game 

Naturalist,'  vol.  iv.,  Jan.  1871.  Birds   of  Sweden,'  18G7,  p.  19,  and 

'^  'The   Ibis,'  vol.   iii.    1861,    p.  182.     Montagu  and  Selby  speak  of 

133,    on    the    Progne    Widow-bird.  the    Black    Grouse    as    polygamous 

See    also    on    the     Vidua    axillaris^  and    of   the  Red   Grouse    as   mono- 

ibid.  vol.  ii.   1860,  p.  211.     On  the  gamous. 
polygamy  of   the   Capercailzie    and 


220  The  Descejit  of  Man.  Part  II. 

exception,  and  this  species  is  belieyed  by  Montagu  to  be  a 
polygamist.  Hence  it  appears  that  amongst  bii'ds  there  often 
exists  a  close  relation  between  polygamy  and  the  development  of 
strongly-marked  sexual  differences.  I  asked  Mr.  Bartlett,  of  the 
Zoological  Gardens,  who  has  had  yery  large  experience  with 
birds,  whether  the  male  tragopan  (one  of  the  Gallinacese)  was 
polygamous,  and  I  was  struck  by  his  answering,  "  I  do  not 
"  know,  but  should  think  so  from  his  splendid  colours." 

It  deserves  notice  that  the  instinct  of  pairing  with  a  single 
female  is  easily  lost  under  domestication.  The  wild-duck  is 
strictly  monogamous,  the  domestic-duck  highly  polygamous. 
The  Eev.  W.  D.  Fox  informs  me  that  out  of  some  half-tamed 
wild-ducks,  on  a  large  pond  in  his  neighbourhood,  so  many 
mallards  were  shot  by  the  gamekeeper  that  only  one  was  left  for 
every  seven  or  eight  females ;  yet  unusually  large  broods  were 
reared.  The  guinea-fowl  is  strictly  monogamous  ;  but  Mr.  Fox 
finds  that  his  birds  succeed  best  when  he  keeps  one  cock  to  two 
or  three  hens.  Canary-birds  pair  in  a  state  of  nature,  but  the 
breeders  in  England  successfully  put  one  male  to  four  or  five 
females.  I  have  noticed  these  cases,  as  rendering  it  probable 
that  wild  monogamous  species  might  readily  become  either 
temporarily  or  permanently  polygamous. 

Too  little  is  known  of  the  habits  of  reptiles  and  fishes  to  enable 
us  to  speak  of  their  marriage  arrangements.  The  stickle -back 
(Gasterosteus),  however,  is  said  to  be  a  polygamist ;"  and  the 
male  during  the  breeding  season  differs  conspicuously  from  the 
female. 

To  sum  up  on  the  means  through  which,  as  far  as  we  can 
judge,  sexual  selection  has  led  to  the  development  of  secondary 
sexual  characters.  It  has  been  shewn  that  the  largest  number 
of  vigorous  offspring  will  be  reared  from  the  pairing  of  the 
strongest  and  best-armed  males,  victorious  in  contests  over 
other  males,  with  the  most  vigorous  and  best-nourished  females, 
which  are  the  first  to  breed  in  the  spring.  If  such  females  select 
the  more  attractive,  and  at  the  same  time  vigorous  males,  they 
will  roar  a  larger  number  of  offspring  than  the  retarded  females, 
which  must  pair  with  the  less  vigorous  and  less  attractive 
males.  So  it  will  be  if  the  more  vigorous  males  select  the  more 
attractive  and  at  the  same  time  healthy  and  vigorous  females ; 
and  this  will  especially  hold  good  if  the  male  defends  the 
female,  and  aids  in  providing  food  for  the  young.  The  ad- 
vantage thus  gained  by  the  more  vigorous  pairs  in  rearing  a 
larger  number  of  offspring  has  apparently  sufficed  to  render 
sexual  selection  efficient.    But  a  large  numerical  preponderance 

1^  Noel  Humphreys,  '  River  Gardens/  1857. 


Chap.  VIII.  Sexual  Selection.  221 


of  males  over  females  will  bo  still  more  efiBcient;  whether  the 
preponderance  is  only  occasional  and  local,  or  permanent; 
whether  it  occurs  at  birth,  or  afterwards  from  the  greater  de- 
struction of  the  females ;  or  whether  it  indirectly  follows  from 
the  practice  of  polygamy. 

The  Male  generally  more  raodified  than  the  Female. — Throughout 
the  animal  kingdom,  when  the  sexes  differ  in  external  appearance, 
it  is,  with  rare  exceptions,  the  male  which  has  been  the  more 
moditied ;  for,  generally,  the  female  retains  a  closer  resemblance 
to  the  young  of  her  own  species,  and  to  other  adult  members  of 
the  same  group.  The  cause  of  this  seems  to  lie  in  the  males 
of  almost  all  animals  having  stronger  passions  than  the  females. 
Hence  it  is  the  males  that  tight  together  and  sedulously  display 
their  charms  before  the  females ;  and  the  victors  transmit  their 
superiority  to  their  male  offspring.  Why  both  sexes  do  not  thus 
acquire  the  characters  of  their  fathers,  will  be  considered  here- 
after. That  the  males  of  all  mammals  eagerly  pursue  the 
females  is  notorious  to  every  one.  So  it  is  with  birds ;  but  many 
cock  birds  do  not  so  much  pursue  the  hen,  as  display  their 
plumage,  perform  strange  antics,  and  pour  forth  their  song  in 
her  presence.  The  male  in  the  few  fish  observed  seems  much 
more  eager  than  the  female;  and  the  same  is  true  of  aUigators, 
and  apparently  of  Batrachians.  Throughout  the  enormous  class  of 
insects,  as  Kirby  remarks, ^^  "  the  law  is,  that  the  male  shall  seek 
"  the  female."  Two  good  authorities,  Mr.  Blackwall  and  Mr.  C. 
Spence  Bate,  tell  me  that  the  males  of  spiders  and  crustaceans 
are  more  active  and  more  erratic  in  their  habits  than  the  females. 
When  the  organs  of  sense  or  locomotion  are  present  in  the  one 
sex  of  insects  and  crustaceans  and  absent  in  the  other,  or  when, 
as  is  frequently  the  case,  they  are  more  highly  developed  in  the 
one  than  in  the  other,  it  is,  as  far  as  I  can  discover,  almost 
invariably  the  male  which  retains  such  organs,  or  has  them  most 
developed;  and  this  shews  that  the  male  is  the  more  active 
member  in  the  courtship  of  the  sexes.^^ 

'^  Kirby  and   Spence,  *  Introduc-  females  of  this  species   are  impreg- 

tion  to  Entomology,'  vol.  iii.   1826,  nated  by  the  males  which  are   born 

p.  342.  in  the  same  cells  with  them  ;   but 

^^  One    parasitic    Hymenopterous  it  is  much  more  probable  that  the 

insect  (Westwood,  '  Modern  Class,  of  females    visit   other    cells,    so    that 

Insects,'  vol.  ii.   p.   160)    forms   an  close  interbreeding  is  thus  avoided, 

exception  to   the  rule,  as  the  male  We  shall  hereafter  meet  in  various 

has  rudimentary  wings,  and  never  classes,  with  a  few  exceptional  cases, 

quits  the  cell  in  which  it  is  born,  in  which  the  female,  instead  of  the 

whilst  the  female  has  well-develcped  male,  is  the  seeker  and  wooer, 
wings.     Audouin  believes  that  the 


222  The  Descent  of  Man.  Part  XL 

The  female,  on  the  other  hand,  with  the  rarest  exceptions,  is 
less  eager  than  the  male.  As  the  illustrious  Hunter  ^^  long  ago 
observed,  she  generally  "  requires  to  be  courted ;"  she  is  coy,  and 
may  often  be  seen  endeavouring  for  a  long  time  to  escape  from 
the  male.  Every  observer  of  the  habits  of  animals  will  be  able 
to  call  to  mind  instances  of  this  kind.  It  is  shown  by  various 
facts,  given  hereafter,  and  by  the  results  fairly  attributable  to 
sexual  selection,  that  the  female,  though  comparatively  passive, 
generally  exerts  some  choice  and  accepts  one  male  in  preference 
to  others.  Or  she  may  accept,  as  appearances  would  sometimes 
lead  us  to  beheve,  not  the  male  which  is  the  most  attractive  to 
her,  but  the  one  which  is  the  least  distasteful.  The  exertion  of 
some  choice  on  the  part  of  the  female  seems  a  law  almost  as 
general  as  the  eagerness  of  the  male. 

We  are  naturally  led  to  enquire  why  the  male,  in  so  many  and 
such  distinct  classes,  has  become  more  eager  than  the  female,  so 
that  he  searches  for  her,  and  plays  the  more  active  part  in  court- 
ship. It  would  be  no  advantage  and  some  loss  of  power  if  each 
sex  searched  for  the  other;  but  why  should  the  male  almost 
always  be  the  seeker?  The  ovules  of  plants  after  fertili- 
sation have  to  be  noiuished  for  a  time;  hence  the  pollen  is 
necessarily  brought  to  the  female  organs — being  placed  on  the 
stigma,  by  means  of  insects  or  the  wind,  or  by  the  spontaneous 
movements  of  the  stamens ;  and  in  the  Algas,  &c.,  by  the  loco- 
motive power  of  the  antherozooids.  With  lowly-organised 
aquatic  animals,  permanently  affixed  to  the  same  spot  and  having 
their  sexes  separate,  the  male  element  is  invariably  brought  to 
the  female ;  and  of  this  we  can  see  the  reason,  for  even  if  the 
ova  were  detached  before  fertilisation,  and  did  not  require 
subsequent  nourishment  or  protection,  there  would  yet  be  greater 
difficulty  in  transporting  them  than  the  male  element,  because, 
being  larger  than  the  latter,  they  are  produced  in  far  smaller 
numbers.  So  that  many  of  the  lower  animals  are,  in  this  re- 
spect, analogous  with  plants.^^  The  males  of  affixed  and  aquatic 
animals  having  been  led  to  emit  their  fertilising  element  in 
this  way,  it  is  natui'al  that  any  of  their  descendants,  which 
rose  in  the  scale  and  became  locomotive,  should  retain  the  same 
habit ;  and  they  would  approach  the  female  as  closely  as  jdos- 
sible,  in  order  not  to  risk  the  loss  of  the  fertilising  element  in  a 
long  passage  of  it  thi'ough  the  water.   With  some  few  of  the  lower 

'"  'Essays       and      Observations.'  of  the  male  and  female  reproductive 

edited  .by  Owen,   vol.    i.    1861,    p.  cells,    remarks,    "  verhalt    sich    die 

194.  "  eine    bei    der  Vereinigung    activ, 

2^  Prof.    Sachs    ('  Lehrbuch    der  "...  die  andere  erscheint  bei  der 

Botanik,'  1870,  s.  633)  in  speaking  "  Vereinigung  passiv." 


Chap.  VIII.  Seximl  Selection.  223 

animals,  the  females  alone  are  fixed,  and  the  males  of  these  must 
be  the  seekers.  But  it  is  difficult  to  understand  why  the  males  of 
species,  of  which  the  progenitors  wxre  primordially  free,  should 
invariably  have  acquired  the  habit  of  approaching  the  females, 
instead  of  being  approached  by  them.  But  in  all  cases,  in  order 
that  the  males  should  seek  efiSciently,  it  would  be  necessary  that 
they  should  be  endowed  with  strong  passions ;  and  the  acquire- 
ment of  such  passions  would  naturally  follow  from  the  more 
eager  leaving  a  larger  number  of  offspring  than  the  less  eager. 
'  The  great  eagerness  of  the  males  has  thus  indirectly  led  to  their 
much  more  frequently  developing  secondary  sexual  characters 
than  the  females.  But  the  development  of  such  characters 
would  be  much  aided,  if  the  males  were  more  liable  to  vary  than 
the  females — as  I  concluded  they  were — after  a  long  study  of 
domesticated  animals.  Yon  Nathusius,  who  has  had  very  wide 
experience,  is  strongly  of  the  same  opinion.^^  Good  evidence  also 
in  favour  of  this  conclusion  can  be  produced  by  a  comparison 
of  the  two  sexes  in  mankind.  During  the  Novara  Expedition  ^^ 
a  vast  number  of  measurements  was  made  of  various  parts  of  the 
body  in  different  races,  and  the  men  were  found  in  almost  every 
case  to  present  a  greater  range  of  variation  than  the  women  ;  but  I 
shall  have  to  recur  to  this  subject  in  a  future  chapter.  Mr.  J. 
Wood,^*  who  has  carefully  attended  to  the  variation  of  the  muscles 
in  man,  puts  in  italics  the  conclusion  that  "  the  greatest  number  of 
''  abnormalities  in  each  subject  is  found  in  the  males,"  He  had 
previously  remarked  that  "altogether  in  102  subjects,  the  varieties 
"  of  redundancy  were  found  to  be  half  as  many  again  as  in 
"  females,  contrasting  widely  with  the  greater  frequency  of 
"  deficiency  in  females  before  described."  Professor  Macalister 
likewise  remarks  ^^  that  variations  in  the  m.uscles  "  are  probably 
"  more  common  in  males  than  females."  Certain  muscles  which 
are  not  normally  present  in  mankind  are  also  more  frequently 
developed  in  the  male  than  in  the  female  sex,  although  excei^tions 
to  this  rule  are  said  to  occur.  Dr.  Burt  Wilder  ^"^  has  tabulated 
the  cases  of  152  individuals  with  supernumerary  digits,  of  which 
86  were  males,  and  39,  or  less  than  half,  females,  the  remaining 
27  being  of  unknown  sex.     It  should  not,  however,  be  overlooked 

22  *  Vortrage     liber     Viehzucht,'  my    '  Variation     of    Animals    and 

1872,  p.  63.  Plants  under  Domestication,'  vol,  ii. 

-^  '  Reise    der   Kovara  :    Anthro-  1868,  p.  75. 

polog.    Theil,'     1867,    s.     216-269.  ^4  'Proceedings   Royal   Soc'  vol. 

The  results  were  calculated  by  Dr.  xvi.  July  1868,  pp.  519  and  524. 

Weisbach  from  measurements  made  -^  '  Troc.   Royal    h-ish   Academy,' 

by  Drs.  K.  Scherzer   and  Schwarz.  vol.  x.  1868,  p.  123. 

On  the    greater   variability   of  the  2"  'Massachusetts    Me<lical    Soc' 

males  of  domesticated  animals,  see  vol.  ii.  No.  3,  1868,  p.  9. 


224  The  Descent  of  Man.  Part  1  J. 

that  women  would  more  freqnently  endeaTour  to  conceal  a 
deformity  of  this  kind  than  men.  Again,  Dr.  L.  Meyer  asserts  that 
the  ears  of  man  are  more  variable  in  form  than  those  of  woman.^'^ 
Lastly  the  temperature  is  more  variable  in  man  than  in  woman.^^ 

The  cause  of  the  greater  general  variability  in  the  male  sex, 
than  in  the  female  is  unknown,  except  in  so  far  as  secondary 
sexual  characters  are  extraordinarily  variable,  and  are  usually 
confined  to  the  males ;  and,  as  we  shall  presently  see,  this  fact  is, 
to  a  certain  extent,  intelligible.  Through  the  action  of  sexual 
and  natural  selection  male  animals  have  been  rendered  in  very 
many  instances  widely  different  from  their  females;  but  in- 
dependently of  selection  the  two  sexes,  from  differing  constitu- 
tionally, tend  to  vary  in  a  somewhat  different  manner.  The 
female  has  to  expend  much  organic  matter  in  the  formation  of 
her  ova,  whereas  the  male  expends  much  force  in  fierce  contests 
with  his  rivals,  in  wandering  about  in  search  of  the  female,  in 
exerting  his  voice,  pouring  out  odoriferous  secretions,  &c, :  and 
this  expenditure  is  generally  concentrated  within  a  short  period. 
The  great  vigour  of  the  male  during  the  season  of  love  seems 
often  to  intensify  his  colours,  independently  of  any  marked  dif- 
ference /rom  the  female.^  In  mankind,  and  even  as  low  down 
in  the  organic  scale  as  in  the  Lepidoptera,,  the  temperature  of  the 
body  is  higher  in  the  male  than  in  the  female,  accompanied  in  the 
case  of  man  by  a  slower  pulse.^^  On  the  whole  the  expenditure 
of  matter  and  force  by  the  two  sexes  is  probably  nearly  equal, 
though  effected  in  very  different  ways  and  at  different  rates. 

From  the  causes  just  specified  the  two  sexes  can  hardly  fail  to 
differ  somewhat  in  constitution,  at  least  during  the  breeding 
season;  and,  although  they  may  be  subjected  to  exactly  the 
same  conditions,  they  will  tend  to  vary  in  a  different  manner. 
If  such  variations  are  of  no  service  to  either  sex,  they  will  not  be 
accumulated  and  increased  by  sexual  or  natural  selection.  Never- 
theless, they  may  become  ^Dermanent  if  the  exciting  cause  acts 

^'^  'Archiv   fiir  Path.  Anat.  und  and  retention  by  them  of  the  sperm- 

Phys.'  1871,  p.  488.  atic   fluid;  but  this   can  hardly  be 

^'^  The    conclusions    recently    ar-  the  case;  for  many  male  birds,  for 

rived  at  by  Dr.  J.  Stockton  Hough,  instance    young    pheasants,    become 

on    the    temperature    of    man,    are  brightly  coloured  in  the  autumn  of 

given  in  the  '  Pop.  Science  Review,'  their  first  year. 
Jan.  1st,  1874,  p.  97.  ^**  For  mankind,  see  Dr.  J.  Stock- 

-^  Prof.    Mantegazza    is    inclined  ton   Hough,   whose   conclusions   are 

to  believe  (' Lettei-a  a  Carlo  Darwin,'  given  in  the  'Pop.  Science  Review,' 

•Archivio      per     1' Anthr  opologia,'  1874,  p.  97.     See  Girard's  observa- 

1871,    p.    306)    that     the     bright  tions  on  the  Lepidoptera,  as   given 

colours,  common    in  so  many  male  in  the  '  Zoological  Record,'  1869,  p. 

animals,  are   due     to   the   presence  347. 


Chap.  VIll.  Sexual  Selection.  225 

permanently ;  and  in  accordance  witli  a  frequent  form  of  inheri- 
tance they  may  be  transmitted  to  that  sex  alone  in  which  they 
first  appeared.  In  this  case  the  two  sexes  will  come  to  present 
permanent,  yet  unimportant,  differences  of  character.  For 
instance,  Mr,  Allen  shews  that  with  a  large  number  of  birds 
inhabiting  the  northern  and  southern  United  States,  the  s])cci- 
mens  from  the  south  are  darker-coloured  than  those  from  the 
north ;  and  this  seems  to  be  the  direct  result  of  the  difference  in 
temperature,  light,  &c.,  between  the  two  regions.  Now,  in  some 
few  cases,  the  two  sexes  of  the  same  species  appear  to  have  been 
differently  affected  ;  in  the  Agdcms  2'>hoeniceus  the  males  have  had 
their  colours  gi-eatly  intensified  in  the  south;  w^hereas  with  Cur- 
diualis  Virginia nus  it  is  the  females  which  have  been  thus  affected  ; 
with  Quiscahis  major  the  females  have  been  rendered  extremely 
variable  in  tint,  whilst  the  reales  remain  nearly  uniform.^^ 

A  few  exceptional  cases  occur  in  various  classes  of  animals,  in 
which  the  females  instead  of  the  males  have  acquired  well 
pronounced  secondary  sexual  characters,  such  as  brighter  colours, 
greater  size,  strength,  or  pugnacity.  With  birds  there  has  some- 
times been  a  complete  transposition  of  the  ordinary  characters 
proper  to  each  sex ;  the  females  having  become  the  more  eager 
in  courtship,  the  males  remaining  comparatively  passive,  but 
apparently  selecting  the  more  attractive  females,  as  we  may  infer 
from  the  results.  Certain  hen  birds  have  thus  been  rendered 
more  highly  coloured  or  otherwise  ornamented,  as  well  as  more 
powerful  and  jDugnacious  than  the  cocks ;  these  characters  being 
transmitted  to  the  female  offspring  alone. 

It  may  be  suggested  that  in  some  cases  a  double  process  of 
selection  has  been  carried  on;  that  the  males  have  selected 
the  more  attractive  females,  and  the  latter  the  more  attractive 
males.  This  process,  however,  though  it  might  lead  to  the 
modification  of  both  sexes,  would  not  make  the  one  sex 
different  from  the  other,  unless  indeed  their  tastes  for  the  beauti- 
ful differed ;  but  this  is  a  supposition  too  improbable  to  be  worth 
considering  in  the  case  of  any  animal,  excepting  man.  There 
are,  however,  many  animals  in  which  the  sexQs  resemble  each 
other,  both  being  furnished  with  the  same  ornaments,  which 
analogy  would  lead  us  to  attribute  to  the  agency  of  sexual 
selection.  In  such  cases  it  may  be  suggested  with  more  plausi- 
bility, that  there  has  teen  a  double  or  mutual  process  of  sexual 
solection ;  the  more  vigorous  and  precocious  females  selecting 
the  more  attractive  and  vigorous  males,  the  latter  rejecting  all 
except  tho  more  attractive  females.     But  from  what  we  know 

*'  '  Mamtuals  and  Birds  of  E.  Florida,'  pp.  234,  280,  295. 
11 


226  The  Descent  of  Man.  Paet  II. 

of  the  habits  of  animals,  this  view  is  hardly  probable,  for  the 
male  is  generally  eager  to  pair  Tvith  any  female.  It  is  more 
probable  that  the  ornaments  common  to  both  sexes  were  acquired 
by  one  sex,  generally  the  male,  and  then  transmitted  to  the  off- 
spring of  both  sexes.  If,  indeed,  during  a  lengthened  period  the 
males  of  any  species  were  gi'eatly  to  exceed  the  females  in 
number,  and  then  during  another  lengthened  period,  but  under 
different  conditions,  the  reverse  were  to  occur,  a  double,  but 
not  simultaneous,  process  of  sexual  selection  might  easily  be 
carried  on,  by  which  the  two  sexes  might  be  rendered  widely 
different. 

We  shall  hereafter  see  that  many  animals  exist,  of  which 
neither  sex  is  brilliantly  coloured  or  provided  with  special  orna- 
ments, and  yet  the  members  of  both  sexes  or  of  one  alone  have 
probably  acquired  simple  colour's,  such  as  white  or  black,  through 
sexual  selection.  The  absence  of  bright  tints  or  other  ornaments 
may  be  the  result  of  yariations  of  the  right  kind  never  having 
occurred,  or  of  the  animals  themselves  having  preferred  iDlain 
black  or  white.  Obscure  tints  have  often  been  developed 
through  natural  selection  for  the  sake  of  protection,  and  the 
acquii-ement  through  sexual  selection  of  conspicuous  colours, 
appears  to  have  been  sometimes  checked  from  the  danger  thus 
incurred.  But  in  other  cases  the  males  during  long  ages  may 
have  struggled  together  for  the  possession  of  the  females,  and 
yet  no  effect  will  have  been  produced,  unless  a  larger  number  of 
offspring  were  left  by  the  more  successful  males  to  inherit  their 
superiority,  than  by  the  less  successful :  and  this,  as  previously 
shewn,  depends  on  many  complex  contingencies. 

Sexual  selection  acts  in  a  less  rigorous  manner  than  natural 
selection.  The  latter  produces  its  effects  by  the  life  or  death  at 
all  ages  of  the  more  or  less  successful  individuals.  Death,  indeed, 
not  rarely  ensues  from  the  conflicts  of  rival  males.  But  generally 
the  less  successful  male  merely  fails  to  obtain  a  female,  or  obtains  a 
retarded  and  less  vigorous  female  later  in  the  season,  or,  if  poly- 
gamous, obtains  fewer  females  ;  so  that  they  leave  fewer,  less  vigor- 
ous, or  no  offspring.  In  regard  to  structures  acquired  through 
ordinary  or  natural  selection,  there  is  in  most  cases,  as  long  as  the 
conditions  of  hfe  remain  the  same,  a  limit  to  the  amount  of 
advantageous  modification  in  relation  to  certain  special  purposes ; 
but  in  regai'd  to  structures  adapted  to  make  one  male  victorious 
over  another,  either  in  fighting  or  in  charming  the  female,  there 
is  no  definite  limit  to  the  amount  of  advantageous  modification ; 
so  that  as  long  as  the  proper  variations  arise  the  work  of  sexual 
selection  will  go  on.  This  circumstance  may  partly  account  for 
tlie  frequent  and  extraordinary  amount  of  variability  presented 


CfiAP.  VIII.  Sexual  Selection.  227 

by  secondary  sexual  characters.  Nevertheless,  natural  selection 
-will  determine  that  such  characters  shall  not  be  acquired  by  the 
Tictorious  males,  if  they  would  be  highly  injurious,  either  by 
expending  too  much  of  their  vital  powers,  or  by  exposing  them 
to  any  great  danger.  The  development,  however,  of  certain 
structures— of  the  horns,  for  instance,  in  certain  stags — has  been 
carried  to  a  wonderful  extreme ;  and  in  some  cases  to  an  extreme 
which,  as  far  as  the  general  conditions  of  life  are  concerned, 
must  be  slightly  injurious  to  the  male.  From  this  fact  we  learn 
that  the  advantages  which  favoured  males  derive  from  conquer- 
ing other  males  in  battle  or  courtship,  and  thus  leaving  a 
numerous  progeny,  are  in  the  long  run  greater  than  those  derived 
from  rather  more  perfect  adaptation  to  their  conditions  of  life. 
We  shall  further  see,  and  it  could  never  have  been  anticipated, 
that  the  power  to  charm  the  female  has  sometimes  been  more 
important  than  the  power  to  conquer  other  males  in  battle. 


LAWS   OF   INHEIIITANCE. 

In  order  to  understand  how  sexual  selection  has  acted  on  many 
animals  of  many  classes,  and  in  the  course  of  ages  has  produced 
a  conspicuous  result,  it  is  necessary  to  bear  in  mind  the  laws  of 
inheritance,  as  far  as  they  are  known.  Two  distinct  elements 
are  included  under  the  term  "  inheritance  " — the  transmission, 
and  the  development  of  characters ;  but  as  these  generally  go 
together,  the  distinction  is  often  overlooked.  We  see  this  dis- 
tinction in  those  characters  which  are  transmitted  through 
the  early  years  of  life,  but  are  developed  only  at  maturity 
or  during  old  age.  We  see  the  same  distinction  more  clearly 
with  secondary  sexual  characters,  for  these  are  transmitted 
through  both  sexes,  though  developed  in  one  alone.  That  they 
are  present  in  both  sexes,  is  manifest  when  two  species,  having 
strongly-marked  sexual  characters,  are  crossed,  for  each  trans- 
mits the  characters  proper  to  its  own  male  and  female  sex  to  the 
liybrid  offspring  of  either  sex.  The  same  fact  is  likewise  mani- 
fest, when  characters  proper  to  the  male  are  occasionally  deve- 
loped in  the  female  when  she  grows  old  or  becomes  diseased, 
as,  for  instance,  when  the  common  hen  assumes  the  flowing  tail- 
feathers,  hackles,  comb,  spurs,  voice,  and  even  i^ugnacity  of  the 
cock.  Conversely,  the  same  thing  is  evident,  more  or  less  plainly, 
with  castrated  mal  es.  Again,  independently  of  old  age  or  disease, 
characters  are  occasionally  transferred  from  the  male  to  the 
female,  as  when,  in  certain  breeds  of  the  fowl,  spurs  regularly 
appear  in  the  young  and  healthy  females.  But  in  truth  they  are 
simply  developed  in  the  female ;  for  in  every  breed  each  detail 


228  The  Descent  of  Man.  Part  II. 

in  the  structure  of  tlie  spur  is  transmitted  through  the  female 
to  her  male  offspring.  Many  cases  will  hereafter  be  given,  where 
the  female  exhibits,  more  or  less  perfectly,  characters  proper  to 
the  male,  in  whom  they  must  have  been  first  developed,  and  then 
transferred  to  the  female.  The  converse  case  of  the  first  de- 
velopment of  characters  in  the  female  and  of  transference  to  the 
male,  is  less  frequent ;  it  will  therefore  be  well  to  give  one  strik- 
ing instance.  With  bees  the  pollen-collecting  apparatus  is  used 
by  the  female  alone  for  gathering  pollen  for  the  larvae,  yet  in 
most  of  the  species  it  is  partially  developed  -in  the  males 
to  whom  it  is  quite  useless,  and  it  is  perfectly  developed 
in  the  males  of  Bombus  or  the  humble-bee.^^  As  not  a 
single  other  Hymenopterous  insect,  not  even  the  wasp,  which  is 
closely  allied  to  the  bee,  is  provided  with  a  pollen-collecting 
apparatus,  we  have  no  grounds  for  supposing  that  male  bees 
primordially  collected  pollen  as  .well  as  the  females ;  although 
we  have  some  reason  to  suspect  that  male  mammals  primordially 
suckled  their  young  as  well  as  the  females.  Lastly,  in  all  cases  of 
reversion,  characters  are  transmitted  through  two,  three,  or  many 
more  generations,  and  are  then  developed  under  certain  unknown 
favourable  conditions.  This  important  distinction  between 
transmission  and  development  will  be  best  kept  in  mind  by  the 
aid  of  the  hypothesis  of  pangenesis.  According  to  this  hypothesis, 
every  unit  or  cell  of  the  body  throws  off  gemmules  or  undeveloped 
atoms,  which  are  transmitted  to  the  offspring  of  both  sexes,  and 
are  multiphed  by  self-division.  They  may  remain  undeveloped 
during  the  early  years  of  life  or  during  successive  generations; 
and  their  development  into  units  or  cells,  like  those  from  which 
they  were  derived,  depends  on  their  affinity  for,  and  union 
with  other  units  or  cells  previously  developed  in  the  due  order 
of  growth. 

Inheritance  at  corresponding  Periods  of  Life. — This  tendency 
is  well  established.  A  new  character,  appearing  in  a  young 
animal,  whether  it  lasts  throughout  life  or  is  only  transient,  will,' 
in  general,  reappear  in  the  offspring  at  the  same  age  and  last 
for  the  same  time.  If,  on  the  other  hand,  a  new  character 
appears  at  maturity,  or  even  during  old  age,  it  tends  to  re- 
appear in  the  offspring  at  the  same  advanced  age.  When  devia- 
tions from  this  rule  occur,  the  transmitted  characters  much 
oftenor  appear  before,  than  after  the  corj-esponding  age.  As  I 
have  dwelt  on  this  subject  sufficiently  in  another  work,^^  I  will 

^-  H.    Miiller,    '  Anweudung    der  ^^    'The    Variation    of    Animals 

Darwin'schen     Lehre,'     &c.     Verb.      and    Plants    under    Domestication,' 
d.  n.  V.  Jahrg.  xxix.  p.  42.  vol.  ii.   1868,  p.  75.     In   the   last 


Chap.  Vlll.  Sexual  Selection.  229 

here  merely  give  two  or  three  instances,  for  the  sake  of  recalling 
the  subject  to  the  reader's  mind.  In  several  breeds  of  the  Fowl, 
the  down-covered  chickens,  the  young  birds  in  their  first  true 
X)limiage,  and  the  adults  differ  greatly  from  one  another,  as  well 
as  from  their  common  parent-form,  the  Gallus  hankiva ;  and 
these  characters  are  faithfully  transmitted  by  each  breed  to  their 
offspring  at  the  corresponding  periods  of  life.  For  instance,  the 
chickens  of  spangled  Hamburg's,  whilst  covered  with  down,  have 
a  few  dark  spots  on  the  head  and  rump,  but  are  not  striped 
longitudinally,  as  in  many  other  breeds ;  in  their  first  true  plu- 
mage, "  they  are  beautifully  pencilled,"  that  is  each  feather  is 
transversely  marked  by  numerous  dark  bars  ;  but  in  their  second 
plumage  the  feathers  all  become  spangled  or  tipped  with  a  dark 
round  spot.^^  Hence  in  this  breed  variations  have  occurred  at, 
and  been  transmitted  to,  three  distinct  periods  of  life.  The 
Pigeon  offers  a  more  remarkable  case,  because  the  aboriginal 
parent  species  does  not  undergo  any  change  of  plumage  with 
advancing  age,  excepting  that  at  maturity  the  breast  becomes 
more  iridescent ;  yet  there  are  breeds  which  do  not  acquire  their 
characteristic  colours  until  they  have  moulted  two,  three,  or 
four  times ;  and  these  modifications  of  plumage  are  regularly 
transmitted. 

Inheritance  at  corresponding  Seasons  of  the  Year. — With  animals 
in  a  state  of  nature,  innumerable  instances  occur  of  characters 
appearing  periodically  at  different  seasons.  We  see  this  in  the 
horns  of  the  stag,  and  in  the  fur  of  arctic  animals  which  becomes 
thick  and  white  during  the  winter.  Many  birds  acquire  bright 
colours  and  other  decorations  during  the  breeding-season  alone. 
Pallas  states,^^  that  in  Siberia  domestic  cattle  and  horses  become 
lighter-coloured  during  the  winter ;  and  I  have  myself  observed, 
and  heard  of  similar  strongly  marked  changes  of  colour,  that  is, 
from  brownish  cream-colour  or  reddish-brown  to  a  perfect  white, 
in  several  ponies  in  England.  Although  I  do  not  know  that  this 
tendency  to  change  the  colour  of  the  coat  during  different  seasons 


chapter    but    one,    the   provisional  mals,'    &c.,    vol.    i.    pp.    160,    249  ; 

hypothesis     of     pangenesis,     above  vol.  ii.  p.  77. 

alluded  to,  is  fully  explained.  "  '  Novte  species  Quadrupeduui  e 
^*  These  Aicts  are  given  on  the  Glirium  ordine,'  177<S,  p.  7.  On 
high  authority  of  a  great  breeder,  the  transmission  of  colour  by  the 
Mr.  Teebay  ;  see  Tegetmeier's  '  Poul-  horse,  see  '  Variation  of  Animals, 
try  Book,'  1868,  p.  158.  On  the  &c.,  under  Domestication,'  vol.  i.  p. 
characters  of  chickens  of  different  51.  Also  vol.  ii.  p.  71,  for  a  gene- 
breeds,  and  on  the  bi-eeds  of  the  ral  discussion  on  '  Inheritance  as 
pigeon,  alluded  to  in  the  following  limited  by  Sex.' 
paragraph,  see  '  Variation   of  Ani- 


230  The  Descent  of  Man.  Part  II. 

is  transmitted,  yet  it  probably  is  so,  as  all  shades  of  colour  are 
strongly  inherited  by  the  horse.  Nor  is  this  form  of  inheritance, 
as  limited  by  the  seasons,  more  remarkable  than  its  limitation 
by  age  or  sex. 

Inheritance  as  Limited  :ly  Sex. — The  equal  transmission  of 
characters  to  both  sexes  is  the  commonest  form  of  inheritance, 
at  least  with  those  animals  which  do  not  present  strongly-marked 
sexual  differences,  and  indeed  with  many  of  these.  But  characters 
are  somewhat  commonly  transferred  exclusively  to  that  sex,  in 
which  they  first  appear.  Ample  evidence  on  tMs  head  has  been 
advanced  in  my  work  on  *  Variation  under  Domestication/  but  a 
few  instances  may  here  be  given.  There  are  breeds  of  the  sheep 
and  goat,  in  which  the  horns  of  the  male  differ  greatly  in  shape 
from  those  of  the  female ;  and  these  differences,  acquired  under 
domestication,  are  regularly  transmitted  to  the  same  sex.  As  a 
rule,  it  is  the  females  alone  in  cats  which  are  tortoise-shell, 
the  corresponding  colour  in  the  males  being  rusty-red.  With 
most  breeds  of  the  fowl,  the  characters  proper  to  each  sex 
are  transmitted  to  the  same  sex  alone.  So  general  is  this  form 
of  transmission  that  it  is  an  anomaly  when  variations  in  certain 
breeds  are  transmitted  equally  to  both  sexes.  There  are  also 
certain  sub-breeds  of  the  fowl  in  which  the  males  can  hardly  be 
distinguished  from  one  another,  whilst  the  females  differ  con- 
siderably in  colour.  The  sexes  of  the  pigeon  in  the  parent-species 
do  not  differ  in  any  external  character;  nevertheless,  in  certain 
domesticated  breeds  the  male  is  coloured  differently  from  the 
female.^^  The  wattle  in  the  English  Carrier  pigeon,  and  the  crop 
in  the  Pouter,  are  more  highly  developed  in  the  male  than  in  the 
female ;  and  although  these  characters  have  been  gained  through 
long-continued  selection  by  man,  the  slight  differences  between 
the  sexes  are  wholly  due  to  the  form  of  inheritance  which  has 
prevailed ;  for  they  have  arisen,  not  from,  but  rather  in  opposi- 
tion to,  the  wish  of  the  breeder. 

Most  of  our  domestic  races  have  been  formed  by  the  accumula- 
tion of  many  slight  variations;  and  as  some  of  the  successive 
steps  have  been  transmitted  to  one  sex  alone,  and  some  to  both 
sexes,  we  find  in  the  different  breeds  of  the  same  species  all 
gradations  between  great  sexual  dissimilarity  and  complete 
similarity.  Instances  have  already  been  given  with  the  breeds 
of  the  fowl  and  pigeon,  and  under  nature  analogous  cases  are 

^^  Dr.  Chapnis,  '  Le  Pigeon  Voya-  similar  differences  in  certain  breeds 

geur  Beige,'  1865,  p.  87.     Boitard  at     Modena,     '  Le     variazioni     dei 

et  Corbie,  '  Les  Pigeons  de  Voliere,'  Colombi   domestici,'   del    Paolo    Bo- 

&c.,   1824,    p.    173.     See,    also,    on.  nizzi,  1873. 


Chap,  y HI.  Sexual  Selection  231 

comraon.  With  auimals  under  domestication,  but  whether  in 
nature  I  will  not  venture  to  say,  one  sex  may  lose  characters 
jjropcr  to  it,  and  may  thus  come  somewhat  to  resemble  the 
opposite  sex  ;  for  instance,  the  males  of  some  breeds  of  the  fowl 
have  lost  their  masculine  tail-plumes  and  hackles.  On  the 
other  hand,  the  differences  between  the  sexes  may  be  increased 
under  domestication,  as  with  merino  sheep,  in  which  the 
e^^•es  have  lost  their  horns.  Again,  characters  proper  to  one 
sex  may  suddenly  aj^pear  in  the  other  sex;  as  in  those  sub- 
breeds  of  the  fowl  in  which  the  hens  acquire  spurs  whilst  young  ; 
or,  as  in  certain  Polish  sub-breeds,  in  which  the  females,  as 
there  is  reason  to  believe,  originally  acquired  a  crest,  and  sub- 
sequently transferred  it  to  the  males.  All  these  cases  are  in- 
telligible on  the  hypothesis  of  pangenesis ;  for  they  depend  on 
the  gemmules  of  certain  parts,  although  present  in  both  sexes, 
becoming,  through  the  influence  of  domestication,  either  dormant 
or  developed  in  either  sex. 

There  is  one  difficult  question  which  it  will  be  convenient  to 
defer  to  a  future  chapter ;  namely,  whether  a  character  at  first 
developed  in  both  sexes,  could  through  selection  be  limited  in 
its  development  to  one  sex  alone.  If,  for  instance,  a  breeder 
observed  that  some  of  his  pigeons  (of  which  the  characters  are 
usually  transferred  in  an  equal  degree  to  both  sexes)  varied  into 
pale  blue,  could  he  by  long-continued  selection  make  a  breed, 
in  which  the  males  alone  should  be  of  this  tint,  whilst  the  females 
remained  unchanged  ?  I  will  here  oftly  say,  that  this,  though 
jDcrhaps  not  impossible,  would  be  extremely  difficult;  for  the 
natural  result  of  breeding  from  the  pale-blue  males  would  be 
to  change  the  whole  stock  of  both  sexes  to  this  tint.  If,  how- 
ever, variations  of  the  desired  tint  appeared,  which  were  from 
the  fijst  limited  in  their  development  to  the  male  sex,  there  would 
not  be  the  least  difficulty  in  making  a  breed  with  the  two  sexes 
of  a  different  colour,  as  indeed  has  been  effected  with  a  Belgian 
breed,  in  which  the  males  alone  are  streaked  with  black.  In  a 
similar  manner,  if  any  variation  appeared  in  a  female  pigeon, 
which  was  from  the  first  sexually  limited  in  its  development  to 
the  females,  it  would  be  easy  to  make  a  breed  with  the  females 
alone  thus  characterised ;  but  if  the  variation  was  not  thus 
originally  limited,  the  process  would  be  extremely  difficult,  per- 
haps impossible.^^ 

^'  Since  the  publication  of  the  perieaced  a  breeder  as  Mr,  Teget- 
first  edition  of  this  work,  it  has  meier.  After  describing  some  cu- 
been  highly  satisfactory  to  me  to  rious  cases  in  pigeons,  of  the  trans- 
find  the  following  remarks  (the  mission  of  colour  by  one  sex  alone, 
'  Field,'   Sept.    1872)    from   so   ex-  and  the  formation  of  a   sub-breed 


232  The  Descent  of  Man.  Part  II. 


On  the  Relation  between  the  Period  of  Development  of  a  Character 
and  its  Transmission  to  one  Sex  or  to  both  Sexes. — "Why  certain 
characters  should  be  inherited  by  both  sexes,  and  other  charac- 
ters by  one  sex  alone,  namely  by  that  sex  in  which  the  character 
first  appeared,  is  in  most  cases  qiiite  unknown.  We  cannot  even 
conjectui'e  why  with  certain  sub-breeds  of  the  pigeon,  black 
striae,  though  transmitted  through  the  female,  should  be  deve- 
loped in  the  male  alone,  whilst  every  other  character  is  equally 
transferred  to  both  sexes.  Why,  again,  witli  cats,  the  tortoise- 
shell  colour  should,  with  rare  exceptions,  be  develoj^ed  in  the 
female  alone.  The  very  same  character,  such  as  deficient  or  su- 
pernumerary digits,  colour-blindness,  &c.,  may  with  mankind  be 
inherited  by  the  males  alone  of  one  family,  and  in  another  family 
by  the  females  alone,  though  in  both  cases  transmitted  through 
the  opposite  as  well  as  through  the  same  sex.^^  Although  we  are 
thus  ignorant,  the  two  following  rules  seem  often  to  hold  good — 
that  variations  which  first  appear  in  either  sex  at  a  late  period  of 
life,  tend  to  be  developed  in  the  same  sex  alone  ;  whilst  variations 
which  first  appear  early  in  life  in  either  sex  tend  to  be  developed  in 
both  sexes.  I  am,  however,  far  from  supposing  that  this  is  the 
sole  determining  cause.  As  I  have  not  elsewhere  discussed  this 
subject,  and  as  it  has  an  important  bearing  on  sexual  selection, 
I  must  here  enter  into  lengthy  and  somewhat  intricate  details. 

It  is  in  itself  probable  that  any  character  aj^pearing  at  an 
early  age  would  tend  to  be  inherited  equally  by  both  sexes,  for 
the  sexes  do  not  differ  much  in  constitution  before  the  jDower 
of  reproduction  is  gained.  On  the  other  hand,  after  this  power 
has  been  gained  and  the  sexes  have  come  to  differ  in  constitution, 
the  gemmules  (if  I  may  again  use  the  language  of  pangenesis) 
which  are  cast  off  from  each  varying  part  in  the  one  sex  would 
be  much  more  likely  to  possess  the  proper  affinities  for  uniting 
with  the  tissues  of  the  same  sex,  and  thus  becoming  developed, 
than  with  those  of  the  ojDposite  sex. 

I  was  first  led  to  infer  that  a  relation  of  this  kind  exists,  from 
the  fact  that  whenever  and  in  whatever  manner  the  adult  male 
differs  from  the  adult  female,  he  differs  in  the  same  manner  from 
the  young  of  both  sexes.  The  generality  of  this  fact  is  quite 
remari?:able :  it  holds  good  with  almost  all  mammals,  birds, 


with  this  character,  he  says  :  "  It  is  *'  facts  that  I  have  rehited ;  but  it 

"  a  singular  circumstance  that  Mr.  "  is    remarkable    how  very  closely 

"  Darwin  should  have  suggested  the  "  he  suggested  the  right  method  of 

"  possibility  of  modifying  the  sexual  "  procedure." 

"  colours    of  birds  by  a  course  of  ^*  References    are    given   in    my 

"  artificial  selection.     When  he  did  Variation  of  Animals  under  Domes- 

'•  so,  he  was  in  ignorance  of  these  tication,'  vol.  ii.  p.  72. 


Chap.  Vlll.  Scxnal  Selection.  233 

amphibians,  and  fislies  ;  also  with  many  crustaceans,  spiders,  and 
some  few  insects,  such  as  certain  orthoptera  and  libellul?e.  In 
all  these  cases  the  variations,  through  the  accumulation  of  which 
the  male  acquired  his  proper  masculine  characters,  must  have 
occurred  at  a  somewhat  late  period  of  life ;  otherwise  the  young 
males  would  have  been  similarly  characterised ;  and  conformably 
with  our  rule,  the  variations  are  transmitted  to  and  developed  in 
the  adult  males  alone.  When,  on  the  other  hand,  the  adult  male 
closely  resembles  the  young  of  both  sexes  (these,  with  rare 
exceptions,  being  alike),  he  generally,  resembles  the  adult  female; 
and  in  most  of  these  cases  the  variations  through  which  the  young 
and  old  acquired  their  present  characters,  probably  occurred, 
according  to  our  rule,  during  youth.  But  there  is  here  room  for 
doubt,  for  characters  are  sometimes  transferred  to  the  offspring 
at  an  earlier  age  than  that  at  which  they  first  appeared  in  the 
parents,  so  that  the  parents  may  have  varied  when  adult,  and 
have  transferred  their  characters  to  their  offspring  whilst  young. 
There  are,  moreover,  many  animals,  in  which  the  two  sexes  closely 
resemble  each  other,  and  yet  both  differ  from  their  young ;  and 
here  the  characters  of  the  adults  must  have  been  acquired  late  in 
life ;  nevertheless,  these  characters,  in  apparent  contradiction  to 
our  rule,  are  transferred  to  both  sexes.  We  must  not,  however, 
overlook  the  possibility  or  even  probability  of  successive  varia- 
tions of  the  same  nature  occurring,  under  exposure  to  similar 
conditions,  simultaneously  in  both  sexes  at  a  rather  late  i)eriod 
of  life ;  and  in  this  case  the  variations  would  be  transferred  to 
the  offspring  of  both  sexes  at  a  corresponding  late  age ;  and  there 
would  then  be  no  real  contradiction  to  the  rule  that  variations 
occurring  late  in  life  are  transferred  exclusively  to  the  sex  in 
which  they  first  appeared.  This  latter  rule  seems  to  hold  true 
more  generally  than  the  second  one,  namely,  that  vai'iations 
which  occur  in  either  sex  early  in  hfe  tend  to  be  transferred  to 
both  sexes.  As  it  was  obviously  impossible  even  to  estimate  in 
how  large  a  number  of  cases  throughout  the  animal  kingdom 
these  two  propositions  held  good,  it  occurred  to  me  to  investigate 
some  striking  or  crucial  instances,  and  to  rely  on  the  result. 

An  excellent  case  for  investigation  is  afforded  by  the  Deer 
family.  In  all  the  species,  but  one,  the  horns  are  developed 
only  in  the  males,  though  certainly  transmitted  through  the 
females,  and  capable  of  abnormal  develoiDment  in  them.  In  the 
reindeer,  on  the  other  hand,  the  female  is  provided  with  horns ; 
so  that  in  this  species,  the  horns  ought,  according  to  our  rule, 
to  api^ear  early  in  life,  long  before  the  two  sexes  are  mature 
and  have  come  to  differ  much  in  constitution.  In  all  the 
other  species  the  horns  ought  to  appear  later  in   life,  which 


234  The  Descent  of  Man.  Part  II. 

would  lead  to  their  development  in  that  sex  alone,  in  which 
they  first  appeared  in  the  progenitor  of  the  whole  Family.  Now 
in  seven  species,  belonging  to  distinct  sections  of  the  family  and 
inhabiting  different  regions,  in  which  the  stags  alone  bear  horns, 
I  find  that  the  horns  first  appear  at  periods,  varying  from  nine 
months  after  birth  in  the  roebnck,  to  ten,  twelve  or  even  more 
months  in  the  stags  of  the  six  other  and  larger  species.^^  But 
with  the  reindeer  the  case  is  widely  different ;  for,  as  I  hear  from 
Prof.  Nilsson,  who  kindly  made  special  enquiries  for  me  in 
I^apland,  the  horns  appear. in  the  young  animals  within  four  or 
five  weeks  after  birth,  and  at  the  same  time  in  both  sexes.  So 
that  here  we  have  a  structure,  developed  at  a  most  unusually 
early  age  iu  one  species  of  the  family,  and  likewise  common  to 
both  sexes  in  this  one  species  alone. 

In  several  kinds  of  antelopes,  only  the  males  are  provided  with 
horns,  whilst  in  the  greater  number  both  sexes  bear  horns. 
With  respect  to  the  period  of  development,  Mr.  Blyth  informs 
me  that  there  was  at  one  time  in  the  Zoological  Gardens  a  young 
koodoo  (^4;^^.  strepsiceros),  of  which  the  males  alone  are  horned, 
and  also  the  young  of  a  closelj^-allied  species,  the  eland  (Ant. 
oreas),  in  which  both  sexes  are  horned.  Now  it  is  in  strict 
conformity  with  our  rule,  that  in  the  young  male  koodoo, 
although  ten  months  old,  the  horns  were  remarkably  small,  con- 
sidering the  size  ultimately  attained  by  then] ;  whilst  in  the 
young  male  eland,  although  only  three  months  old,  the  horns 
were  already  very  much  larger  than  in  the  koodoo.  It  is 
also  a  noticeable  fact  that  in  the  prong-horned  antelope,'*° 
only  a  few  of  the  females,  about  one  in  five,  have  horns,  and 
these  are  in  a  rudimentary  state,  though  sometimes  above  four 
inches  long ;  so  that  as  far  as  concerns  the  possession  of  horns 
by  the«males  alone,  this  species  is  in  an  intermediate  condition, 
and  the  horns  do  not  ai^pear  until  about  five  or  six  months  after 
birth.  Therefore  in  comparison  with  what  little  we  know  of 
the  development  of  the  horns  in  other  antelopes,  and  from  what 

2^  I    am    much    obliged    to    Mr.  tinent,  see  J.  D.  Caton,  in  'Ottawa 

Cupples  for  having  made  enquiries  Acad,  of  Kat.  Sc.  1868,  p.  13.      For 

for   me   in  regard   to   the    Roebuck  Cervus    EkH   of    Pegu,    see    Lieut, 

and  Red  Deer  of  Scotland  from  Mr.  Beavan,    '  Proc.  Zoolog.  Soc'   1867, 

Robertson,    the    experienced    head-  p.  762. 

forester  to  the  Marquis  of  Breadal-  ^°  Antilocapra  Americana.    I  have 

bane.     In  regard  to  Fallow-deer,   I  to  thank  Dr.  Canfield  for  informa- 

hare     to     thank     Mr.     Eyton    and  tion  Avith  respect  to  the  horns  of  the 

others    for    information.     For    the  female  :  see  also  his  paper  in  '  Proc. 

Cervus    alces    of    N.    America,    see  Zoolog.    Soc'    1866,    p.    109.     Also 

•Land    and  Water,*  1868,  pp.    221  Owen,    'Anatomy    of    Vertebrates,' 

and  254;  and  for  the  C.  Yirgbuanus  vol.  iii.  p.  627. 
and  stronjyLceros  of  the  same  con- 


Chap.  VJII.  Sexual  Selection.  235 


ve  do  know  with  respect  to  the  horns  of  deer,  cattle,  &c.,  those 
of  the  prong-horned  antelope  appear  at  an  intermediate  period 
of  life,— that  is,  not  very  early,  as  in  cattle  and  sheep,  nor  very 
late,  as  in  the  larger  deer  and  antelopes.  The  horns  of  sheep, 
goats,  and  cattle,  which  are  well  developed  in  both  sexes,  thongh 
not  quite  equal  in  size,  can  be  felt,  or  even  seen,  at  birth  or  soon 
afterwards.'*^  Our  rule,  however,  seems  to  fail  in  some  breeds 
of  sheep,  for  instance  merinos,  in  which  the  rams  alone  are 
horned;  for  I  cannot  find  on  enquiry,"*^  that  the  horns  are 
developed  later  in  life  in  this  breed  than  in  ordinary  sheep  in 
■which  both  sexes  are  horned.  But  with  domesticated  sheep  the 
presence  or  absence  of  horns  is  not  a  firmly  fixed  character ;  for 
a  certain  proportion  of  the  merino  ewes  bear  small  horns,  and 
some  of  the  rams  are  hornless;  and  in  most  breeds  hornless 
ewes  are  occasionally  produced. 

Dr.  W.  Marshall  has  lately  made  a  special  study  of  the  pro- 
tuberances so  common  on  the  heads  of  birds,^^  and  he  comes 
to  the  following  conclusion ; — that  with  those  species  in  which 
they  are  confined  to  the  males,  they  are  developed  late  in 
life ;  whereas  with  those  species  in  which  they  are  common  to 
the  two  sexes,  they  are  developed  at  a  very  early  period.  This  is 
certainly  a  striking  confirmation  of  my  two  laws  of  inheritance. 

In  most  of  the  species  of  the  splendid  family  of  the  Pheasants, 
the  males  difi'er  conspicuously  from  the  females,  and  they  acquire 
their  ornaments  at  a  rather  late  period  of  life.  The  eared 
pheasant  (^Crossoptilon  auritum),  however,  offers  a  remarkable 
exception,  for  both  sexes  possess  the  fine  caudal  plumes,  the 
large  ear-tufts  and  the  crimson  velvet  about  the  head ;  I  find 
that  all  these  characters  appear  very  early  in  life  in  accordance 
with  rule.  The  adult  male  can,  however,  be  distinguished  from 
the  adult  female  by  the  presence  of  spui's;    and  conformably 

<i  I  have   been  assured  that  the  however,  a  breed  of  sheej:)  in  which, 

horns  of  the  sheep  in  North  Wales  as   with    merinos,    the    rams   alone 

can  always  be  felt,  and  are  some-  bear     horns  ;     and    Mr.     Winwood 

times   even   an    inch    in   length,  at  Reade  informs  me  that  in  one  case 

birth.     Youatt  says  ('Cattle,' 1834,  observed    by   him,    a    young    ram. 

p.    277),   that    the    prominence    of  born    on    Feb.    10th,    iirst    shewed 

the  frontal  bone  in  cattle  penetrates  horns  on  March  6th,  so  that  in  this 

the    cutis    at    birth,    and    that    the  instance,  in    conformity  with    rule, 

horny  matter   is   soon   formed  over  the   development  of  the  horns  oc- 

it.  curred  at  a  later  period  oi  life  than 

*2  I  am  greatly  indebted  to  Prof.  in  Welsh  sheep,  in  which  both  sexes 

Victor  Carus  for  having  made  en-  are  horned. 

quiries    for   me,   from    the    highest  *^  *  Ueber  die  knocheruen  Schadel- 

authorities,    with    respect    to    the  hocker  der  Vogel'  in  the  '  Kieder- 

merino   sheep    of   Saxony.     On   the  landischen    Archiv    fur     Zoologie,' 

Guinea    coast    of    Africa    there    is,  Baud  I.  Heft  2,  1872. 


236  The  Descent  of  Man.  Part  II. 

with  our  rule,  tliese  do  not  begin  to  be  developed  before  the  age 
of  six  months,  as  I  am  assured  by  Mr.  Bartlett,  and  even  at  this 
age,  the  two  sexes  can  hardly  be  distinguished.^*  The  male  and 
female  Peacock  diifer  conspicuously  from  each  other  in  almost 
every  part  of  their  plumage,  except  in  the  elegant  head-crest, 
which  is  common  to  both  sexes ;  and  this  is  developed  very  early 
in  life,  long  before  the  other  ornaments,  which  are  confined  to  the 
male.  The  wild-duck  offers  an  analogous  case,  for  the  beautiful 
green  speculum  on  the  wings  is  common  to  both  sexes,  though 
duller  and  somewhat  smaller  in  the  female,  and  it  is  developed 
early  in  life,  whilst  the  curled  tail-feathers  and  other  ornaments- 
of  the  male  are  developed  later.'*^  Between  such  extreme  cases 
of  close  sexual  resemblance  and  wide  dissimilarity,  as  those  of 
the  Crossoptilon  and  peacock,  many  intermediate  ones  could  be 
given,  in  which  the  characters  follow  our  two  rules  in  their  order 
of  development. 

As  most  insects  emerge  from  the  pupal  state  in  a  mature 
condition,  it  is  doubtful  whether  the  period  of  development  can 
determine  the  transference  of  their  characters  to  one  or  to  both 
sexes.  But  we  do  not  know  that  the  coloured  scales,  for  instance, 
in  two  species  of  butterflies,  in  one  of  which  the  sexes  differ  in 
colour,  whilst  in  the  other  they  are  ahke,  are  developed  at  the 
same  relative  age  in  the  cocoon.  Nor  do  we  know  whether  all 
the  scales  are  simultaneously  developed  on  the  wings  of  the  same 
species  of  butterfly,  in  which  certain  coloured  marks  are  confined 

•**  In  the  common  peacock  {Paxio  sexes;  but  I  have  not  been  able  to 

cristatus)  the  male  alone  possesses  discover  whether  its   full  develop- 

spurs,  whilst  both  sexes  of  the  Java  ment   occurs    later   in    life    in    the 

Peacock  (P.  muticus)  offer  the  un-  males  of  such  species,  than  in  the 

usual  case  of  being  furnished  with  male  of  the  common  duck,  as  ought 

spurs.     Hence  I  fully  expected  that  to   be    the    case   according    to    our 

in   the    latter   species    they    would  rule.     With  the  allied  Mergus  ctt- 

have  been  developed  earlier  in  life  cullatus  we  have,  however,  a  case  of 

than  in  the  common  peacock;  but  this  kind:  the  two  sexes  differ  con- 

M.  Hegt  of  Amsterdam  informs  me,  spicuously  in  general  plumage,  and 

that  with  young  birds  of  the   pre-  to    a    considerable    degree    in    the 

vious  year,   of  both    species,    com-  speculum,  which  is  pure  white    in 

pared    on   April   23rd,    1869,   there  the  male  and  greyisli-white  in  the 

was    no   difference   in  the   develop-  female.     Now  the  young   males  at 

ment    of    the    spurs.      The    spurs,  first  entirely  resemble  the  females, 

however,   were   as  yet    represented  and  have  a  greyish-white  speculum, 

merely   by  slight    knobs    or    eleva-  which  becomes    pure    white    at    an 

tions.      1    presume    that    I    should  earlier  age  than  that  at  which  the 

have  been  informed  if  any  difference  adult  male  acquires  his  other  and 

in   the    rate    of    development    had  more    strongly-marked  sexual    dif- 

been  observed  subsequently.  ferences  :    see   Audubon,  '  Ornitho- 

*^  In  some  other  species  of  the  logical   Biography,'    vol.    iii.   1835, 

Duck    family  the   speculum   differs  pp.  249-250. 
in    a    greater    degree    in     the    two 


Chap.  VIII.  Sexual  Selection. 


'J/ 


to  one  sex,  whilst  others  are  common  to  both  sexes.  A  difference 
of  this  kind  in  the  period  of  development  is  not  so  improbable  as 
it  may  at  first  appear ;  for  with  the  Orthoptera,  which  assume 
their  adult  state,  not  by  a  single  metamorphosis,  but  by  a  suc- 
cession of  moults,  the  young  males  of  some  species  at  first 
resemble  the  females,  and  acquire  their  distinctive  masculine 
characters  only  at  a  later  moult.  Strictly  analogous  cases  occur 
at  the  successive  moults  of  certain  male  crustaceans. 

We  have  as  yet  considered  the  transference  of  characters,  re- 
latively to  their  period  of  development,  only  in  species  in  a 
natural  state;  we  will  now  turn  to  domesticated  animals,  and 
first  touch  on  monstrosities  and  diseases.  The  presence  of  super- 
numerary digits,  and  the  absence  of  certain  phalanges,  must  be 
determined  at  an  early  embryonic  period— the  tendency  to  profuse 
bleeding  is  at  least  congenital,  as  is  probably  colour-blindness — 
yet  these  peculiarities,  and  other  similar  ones,  are  often  limited 
in  their  transmission  to  one  sex;  so  that  the  rule  that 
characters,  developed  at  an  early  period,  tend  to  be  trans- 
mitted to  both  sexes,  here  wholly  fails.  But  this  rule  as 
before  remarked,  does  not  appear  to  be  nearly  so  general  as  the 
converse  one,  namely,  that  characters  which  appear  late  in  life 
in  one  sex  are  transmitted  exclusively  to  the  same  sex.  From 
the  fact  of  the  above  abnormal  peculiarities  becoming  attached 
to  one  sex,  long  before  the  sexual  functions  are  active,  we  may 
infer  that  there  must  be  some  difference  between  the  sexes  at  an 
extremely  early  age.  With  respect  to  sexually-limited  diseases, 
we  know  too  little  of  the  period  at  which  they  originate,  to  draw 
any  safe  conclusion.  Gout,  however,  seems  to  fall  under  our 
rule,  for  it  is  generally  caused  by  intemperance  during  manhood, 
and  is  transmitted  from  the  father  to  his  sons  in  a  much  inore 
marked  manner  than  to  his  daughters. 

In  the  various  domestic  breeds  of  sheep,  goats,  and  cattle,  the 
males  differ  from  theh  respective  females  in  the  shape  or  develop- 
ment of  their  horns,  forehead,  mane,  dewlap,  tail,  and  humjD  on 
the  shoulders ;  and  these  peculiarities,  in  accordance  with  our 
rule,  are  not  fully  developed  until  a  rather  late  period  of  life. 
The  sexes  of  dogs  do  not  differ,  except  that  in  certain  breeds, 
especially  in  the  Scotch  deer-hound,  the  male  is  much  larger 
and  heavier  than  the  female ;  and,  as  we  shall  see  in  a  future 
chapter,  the  male  goes  on  increasing  in  size  to  an  unusually  late 
period  of  life,  which,  according  to  rule,  will  account  for  liis  in- 
creased size  being  transmitted  to  his  male  offspring  alone.  On 
the  other  hand,  the  tortoise-shell  colour,  which  is  confined  to 
female  cats,  is  ({uite  distinct  at  bhth,  and  this  case  violates  the 
rule.     There  is  a  breed  of  pigeons  in  which  the  males  alone  are 


238  The  Descent  of  Man.  Part  II. 

streaked  with  black,  and  the  streaks  can  be  detected  even  in  the 
nestlings ;  but  they  become  more  conspicuous  at  each  successive 
moult,  so  that  this  case  partly  opposes  and  partly  supports  the 
rule.  With  the  English  Carrier  and  Pouter  pigeons,  the  full 
development  of  the  wattle  and  the  croj)  occurs  rather  late  in  life, 
and  conformably  with  the  rule,  these  characters  are  transmitted 
in  full  perfection  to  the  males  alone.  The  following  cases  perhaps 
come  within  the  class  previously  alluded  to,  in  which  both  sexes 
have  varied  in  the  same  manner  at  a  rather  late  period  of  life, 
and  have  consequently  transferred  their  new  characters  to  both 
sexes  at  a  corresponding  late  period ;  and  if  so,  these  cases  are 
not  opposed  to  our  rule  : — there  exist  sub-breeds  of  the  pigeon, 
described  by  Neumeister,^''  in  which  both  sexes  change  their 
colour  during  two  or  three  moults  (as  is  likewise  the  case  with 
the  Almond  Tumbler),  nevertheless,  these  changes,  though 
occurring  rather  late  in  life,  are  common  to  both  sexes.  One 
variety  of  the  Canary-bud,  namely  the  London  Prize,  offers  a 
nearly  analogous  case. 

With  the  breeds  of  the  Fowl  the  inheritance  of  various  charac- 
ters by  one  or  both  sexes,  seems  generally  determined  by  the 
period  at  which  such  characters  are  developed.  Thus  in  all  the 
many  breeds  in  which  the  adult  male  differs  greatly  in  colour 
from  the  female,  as  well  as  from  the  wild  parent-species,  he 
differs  also  from  the  young  male,  so  that  the  newly-acquired 
characters  must  have  appeared  at  a  rather  late  period  of  Hfe. 
On  the  other  hand,  in  most  of  the  breeds  in  which  the  two  sexes 
resemble  each  other,  the  young  are  coloured  in  nearly  the  same 
manner  as  their  parents,  and  this  renders  it  probable  that  their 
colours  first  aj^peared  early  in  life.  We  have  instances  of  this 
fact  in  all  black  and  white  breeds,  in  which  the  young  and  old 
of  both  sexes  are  alike ;  nor  can  it  be  maintained  that  there  is 
something  pecuhar  in  a  black  or  white  plumage,  which  leads  to 
its  transference  to  both  sexes ;  for  the  males  alone  of  many 
natural  species  are  either  black  or  white,  the  females  being 
differently  coloured.  With  the  so-called  Cuckoo  sub-breeds  of 
the  fowl,  in  which  the  feathers  are  transversely  pencilled  with 
dark  stripes,  both  sexes  and  the  chickens  are  coloured  in  nearly 
the  same  manner.  The  laced  plumage  of  the  Sebright  bantam 
is  the  same  in  both  sexes,  and  in  the  young  chickens  the  wing- 
feathers  are  distinctly,  though  imperfectly  laced.  Spangled 
Hamburgs,  however,  offer  a  partial  exception ;  for  the  two  sexes, 
though  not  quite  alike,  resemble  each  other  more  closely  than 

^^  'DasGanze  der  Taubonzucht,'      puis,     *  Le  pigeon  voyageui'  Beige,' 
1837,  s.   21,   24.     For  the   case  of      1865,  p.  87. 
the  streaked  pigeons,  see  Dr.  Cha- 


Chap.  VIII.  Sexual  Selection.  239 

do  the  sexes  of  the  aborlgiDal  parent-species ;  yet  they  acquu-e 
their  characteristic  phimage  late  in  life,  for  the  chickens  are 
distinctly  pencilled.  With  respect  to  other  characters  besides 
colour,  in  the  wild-parent  species  and  in  most  of  the  domestic 
breeds,  the  males  alone  possess  a  well-developed  comb ;  but  in 
the  young  of  the  Spanish  fowl  it  is  largely  developed  at  a  very 
early  age,  and,  in  accordance  with  this  early  development  in  the 
male,  it  is  of  unusual  size  in  the  adult  female.  In  the  Game 
breeds  pugnacity  is  developed  at  a  wonderfully  early  age,  of 
which  curious  proofs  could  be  given ;  and  this  character  is  trans- 
mitted to  both  sexes,  so  that  the  hens,  from  their  extreme 
pugnacity,  are  now  generally  exhibited  in  separate  pens.  With 
the  Polish  breeds  the  bony  protuberance  of  the  skull  which 
supports  the  crest  is  partially  developed  even  before  the  chickens 
are  hatched,  and  the  crest  itself  soon  begins  to  grow,  though  at 
iirst  feebly  ;*'^  and  in  this  breed  the  adults  of  both  sexes  are 
characterised  by  a  great  bony  protuberance  and  an  immense  crest. 

Finally,  from  what  we  have  now  seen  of  the  relation  which 
exists  in  many  natural  species  and  domesticated  races,  between 
the  period  of  the  develox)ment  of  their  characters  and  the 
manner  of  their  transmission — for  example,  the  striking  fact  of 
the  early  growth  of  the  horns  in  the  reindeer,  in  which  both 
sexes  bear  horns,  in  comparison  with  their  much  later  growth 
in  the  other  species  in  which  the  male  alone  bears  horns — we 
may  conclude  that  one,  though  not  the  sole  cause  of  characters 
being  exclusively  inherited  by  one  sex,  is  their  development  at 
a  late  age.  And  secondly,  that  one,  though  apparently  a  less 
efficient  cause  of  characters  being  inherited  by  both  sexes,  is 
their  development  at  an  early  age,  whilst  the  sexes  differ 
but  little  in  constitution.  It  appears,  however,  that  some 
difference  must  exist  between  the  sexes  even  during  a  very 
early  embryonic  period,  for  characters  developed  at  this  age  not 
rarely  become  attached  to  one  sex. 

Summary  and  concluding  remarls. — From  the  foregoing  dis- 
cussion on  the  various  laws  of  inheritance,  we  learn  that  the 
characters  of  the  parents  often,  or  even  generally,  tend  to  become 
developed  in  the  offspring  of  the  same  sex,  at  the  same  age,  aud 
periodically  at  the  same  season  of  the  year,  in  which  they  Iirst 

*'  For    full    particulars    and    re-  250,  256.     In  regard  to  the  higher 

ferences  on  all  these  points  respect-  animals,  the  sexual  differences  which 

ing  the  several  breeds  of  the  Fowl,  have  arisen  under  domestication  are 

see 'Variation  of  Animals  and  Plants  described  in   the   same  work   under 

under    Domestication,'    vol.    i.    pp.  the  head  of  each  species. 


240  TJie  Descent  of  Man.  Part  II. 


appeared  in  the  parents.  But  these  rules,  owing  to  unknown 
causes,  are  far  from  being  fixed.  Hence  during  the  modification 
of  a  species,  the  successive  changes  may  readily  be  transmitted 
in  different  ways ;  some  to  one  sex,  and  some  to  both ;  some  to 
the  offspring  at  one  age,  and  some  to  the  offspring  at  all  ages. 
Not  only  are  the  laws  of  inheritance  extremely  complex,  but  so 
are  the  causes  which  induce  and  goyem  variability.  The 
variations  thus  induced  are  preserved  and  accumulated  by 
sexual  selection,  which  is  in  itself  an  extremely  complex  affair, 
depending,  as  it  does,  on  the  ardour  in  love,  the  courage,  and 
the  rivab-y  of  the  males,  as  well  as  on  the  powers  of  perception, 
the  taste,  and  will  of  the  female.  Sexual  selection  will  also 
be  largely  dominated  by  natural  selection  tending  towards 
the  general  welfare  of  the  species.  Hence  the  manner  in  which 
the  individuals  of  either  or  both  sexes  have  been  affected 
through  sexual  selection  cannot  fail  to  be  complex  in  the  highest 
degree. 

When  variations  occur  late  in  life  in  one  sex,  and  are  trans- 
mitted to  the  same  sex  at  the  same  age,  the  other  sex  and  the 
young  are  left  unmodified.  When  they  occur  late  in  life,  but 
are  transmitted  to  both  sexes  at  the  same  age,  the  young  alone 
are  left  unmodified.  Variations,  however,  may  occur  at  any 
period  of  hfe  in  one  sex  or  in  both,  and  be  transmitted  to  both 
sexes  at  all  ages,  and  then  all  the  individuals  of  the  species 
are  similarly  modified.  In  the  following  chapters  it  will  be  seen 
that  all  these  cases  frequently  occur  in  nature. 

Sexual  selection  can  never  act  on  any  animal  before  the  age 
for  reproduction  arrives.  From  the  great  eagerness  of  the  male 
it  has  generally  acted  on  this  sex  and  not  on  the  females.  The 
males  have  thus  become  provided  with  weapons  for  fighting 
with  their  rivals,  with  organs  for  discovering  and  securely 
holding  the  female,  and  for  exciting  or  charming  her.  When 
the  sexes  differ  in  these  respects,  it  is  also,  as  we  have  seen,  an 
extremely  general  law  that  the  adult  male  differs  more  or  less 
from  the  young  male ;  and  we  may  conclude  from  this  fact  that 
the  successive  variations,  by  which  the  adult  male  became  modi- 
fied, did  not  generally  occur  much  before  the  age  for  reproduction. 
Whenever  some  or  many  of  the  variations  occurred  early  in 
life,  the  young  males  would  partake  more  or  less  of  the  charac- 
ters of  the  adult  males ;  and  differences  of  this  kind  between 
the  old  and  young  males  may  be  observed  in  many  species  of 
animals. 

It  is  probable  that  young  male  animals  have  often  tended  to 
vary  in  a  manner  which  would  not  only  have  been  of  no  use  to 
them  at  an  early  age,  but  would  have  been  actually  injui'ious— 


Chap.  YIII.  ScxicjI  Selcct^n.  24 1 

as  by  acquiring  bright  colours,  which  would  render  them  con- 
spicuous to  their  enemies,  or  by  acquiring  structures,  such  as 
great  horns,  which  would  exj^end  much  vital  force  in  their 
development.  Variations  of  this  kind  occurring  in  the  young 
males  would  almost  certainly  be  eliminated  through  natural 
selection.  With  the  adult  and  experienced  males,  on  the  other 
hand,  the  advantages  derived  from  the  acquisition  of  such 
characters,  would  more  than  counterbalance  some  exposure  to 
danger,  and  some  loss  of  vital  force. 

As  ■  variations  which  give  to  the  male  a  better  chance  of 
conquering  other  males,  or  of  finding,  securing,  or  charming  the 
opposite  sex,  would,  if  they  happened  to  arise  in  the  female,  be 
of  no  service  to  her,  they  would  not  be  preserved  in  her  through 
sexual  selection.  We  have  also  good  evidence  with  domesticated 
animals,  that  variations  of  all  kinds  are,  if  not  carefully  selected, 
soon  lost  through  intercrossing  and  accidental  deaths.  Conse- 
quently in  a  state  of  nature,  if  variations  of  the  above  kind  chanced 
to  arise  in  the  female  line,  and  to  be  transmitted  exclusively  in 
this  line,  they  would  be  extremely  liable  to  be  lost.  If,  however, 
the  females  varied  and  transmitted  their  newly  acquired 
characters  to  their  offspring  of  both  sexes,  the  characters  which 
were  advantageous  to  the  males  would  be  preserved  by  them 
through  sexual  selection,  and  the  two  sexes  would  in  consequence 
be  modified  in  the  same  manner,  although  such  characters  were  of 
no  use  to  the  females ;  but  I  shall  hereafter  have  to  recur  to  these 
more  intricate  contingencies.  Lastly,  the  females  may  acquire,  and 
apparently  have  often  acquired  by  transference,  characters  from 
the  male  sex. 

As  variations  occurring  late  in  life,  and  transmitted  to  one 
sex  alone,  have  incessantly  been  taken  advantage  of  and  accumu- 
lated through  sexual  selection  in  relation  to  the  reproduction  of 
the  species ;  therefore  it  appears,  at  first  sight,  an  unaccountable 
fact  that  similar  variations  have  not  frequently  been  accumu- 
lated through  natural  selection,  in  relation  to  the  ordinary 
habits  of  life.  If  this  had  occurred,  the  two  sexes  would  often 
have  been  differently  modified,  for  the  sake,  for  instance,  of 
capturing  prey  or  of  escaping  from  danger.  Differences  of  this 
kind  between  the  two  sexes  do  occasionally  occur,  especially  in 
the  lower  classes.  But  this  implies  that  the  two  sexes  follow 
different  habits  in  their  struggles  for  existence,  which  is  a  rare 
circumstance  with  the  higher  animals.  The  case,  however,  is 
widely  different  with  the  reproductive  functions,  in  which  respect 
the  sexes  necessarily  differ.  For  variations  in  structure  which 
are  related  to  these  functions,  have  often  proved  of  value  to  one 
sex,  and  from  having  arisen  at  a  late  period  of  hfe,  have  been 


242  Thi^esccnt  of  Man.  Tart  II. 

transmitted  to  one  sex  alone ;  and  such  variations,  thus  preserved 
and  transmitted,  have  given  rise  to  secondary  sexual  characters. 
In  the  following  chapters,  I  shall  treat  of  the  secondary 
sexual  characters  in  animals  of  all  classes,  and  shall  endeavour  in 
each  case  to  apply  the  principles  explained  in  the  present 
chapter.  The  lowest  classes  will  detain  us  for  a  very  short  time, 
but  the  higher  animals,  especially  birds,  must  be  treated  at 
considerable  length.  It  should  be  borne  in  mind  that  for 
reasons  already  assigned,  I  intend  to  give  only  a  few  illustrative 
instances  of  the  innumerable  structures  by  the  aid  of  which  the 
male  finds  the  female,  or,  when  found,  holds  her.  On  the  other 
hand,  all  structures  and  instincts  by  the  aid  of  which  the  male 
conquers  other  males,  and  by  w^hich  he  allures  or  excites  the 
female,  will  be  fully  discussed,  as  these  are  in  many  ways  the 
most  interesting. 

Supplemtnt  on  the  proportional  numbers  of  the  two  sexes  in  animals 
belonging  to  various  classes. 

As  no  one,  as  far  as  I  can  discover,  has  paid  attention  to  the 
relative  numbers  of  the  two  sexes  throughout  the  animal 
kingdom,  I  will  here  give  such  materials  as  I  have  been  able  to 
collect,  although  they  are  extremely  imperfect.  They  consist  in 
only  a  few  instances  of  actual  enumeration,  and  the  numbers  are 
not  very  large.  As  the  proportions  are  known  with  certainty  only 
in  mankind,  I  will  first  give  them  as  a  standard  of  comparison. 

Man.— In  England  during  ten  years  (from  1857  to  1866)  the 
average  number  of  children  born  alive  yearly  was  707,120,  in 
the  proportion  of  104-5  males  to  100  females.  But  in  1857  the 
male  births  throughout  England  were  as  105-2,  and  in  1865  as 
104-0  to  100.  Looking  to  separate  districts,  in  Buckingham- 
shire (where  about  5000  children  are  annually  born)  the  mean 
proportion  of  male  to  female  births,  during  the  whole  period  of 
the  above  ten  years,  was  as  102-8  to  100;  w^hilst  in  N.  Wales 
(where  the  average  annual  births  are  12,873)  it  w^as  as  high 
as  106-2  to  100.  Taking  a  still  smaller  district,  viz.,  Eut- 
landshire  (where  the  annual  births  average  only  739),  in  1864 
the  male  births  were  as  114*6,  and  in  1862  as  only  97-0^  to 
100 ;  but  even  in  this  small  district  the  average  of  the  7385 
births  during  the  whole  ten  years,  was  as  104-5  to  100 ;  that  is  in 
the  same  ratio  as  throughout  England.'*^  The  proportions  are 
sometimes  shghtly  disturbed  by  unlmown  causes;    thus  Prof. 

*^  'Twenty-ninth  Annual  Report  In  this  report  (p.  xii.)  a  special  de- 
of  the  Ilegistnir-Geueral  for  1866.*      ceunial  table  is  given. 


Chap.  Till.  Proportion  of  the  Sexes.  243 

Faye  states  "  that  in  some  districts  of  Norway  there  has  been 
"  during  a  decennial  jDeriod  a  steady  deficiency  of  boys,  whilst 
'*  in  others  the  opposite  condition  has  existed."  In  France 
during  forty-four  years  the  male  to  the  female  births  have  been 
as  I06'2  to  100;  but  during  this  period  it  has  occurred  five 
times  in  one  department,  and  six  times  in  another,  that  the 
female  births  have  exceeded  the  males.  In  Kussia  the  average 
proportion  is  as  high  as  lOS'U,  and  in  Philadelphia  in  the  United 
States  as  llO'S  to  lOO."*^  The  average  for  Europe,  deduced  by 
Bickes  from  about  seventy  million  births,  is  106  males  to  100 
females.  On  the  other  hand,  with  w^hite  children  born  at  the 
Cape  of  Good  Hope,  the  proportion  of  males  is  so  low  as  to  fluctuate 
during  successive  years  bctw^een  90  and  99  males  for  every  100 
females.  It  is  a  singular  fact  that  with  Jews  the  proportion  of 
male  births  is  decidedly  larger  than  with  Christians  :  thus  in 
Prussia  the  proportion  is  as  113,  in  Breslau  as  114,  and  in 
Livonia  as  120  to  100 ;  the  Christian  births  in  these  countries 
being  the  same  as  usual,  for  instance,  in  Livonia  as  104  to  lOO."*' 
Prof.  Faye  remarks  that  "  a  still  greater  preponderance  of 
"  males  would  be  met  with,  if  death  struck  both  sexes  in  equal 
*'  proportion  in  the  womb  and  during  birth.  But  the  fact  is,  that 
*'  for  every  100  still-born  females,  we  have  in  several  countries 
"  from  134-6  to  144'9  still-born  males.  During  the  first  four  or 
"  five  years  of  life,  also,  more  male  children  die  than  females ; 
"  for  example  in  England,  during  the  first  year,  126  boys  die  for 
"  every  100  girls — a  proportion  which  in  France  is  still  more 
"  unfavourable."^^  Dr.  Stockton-Hough  accounts  for  these  facts 
in  part  by  the  more  frequent  defective  development  of  males 
than  of  females.    We  have  before  seen  that  the  male  sex  is  more 

^^  For    Norway  and    Russia,    see  343.        Dr.     Stark     also     remarks 

abstract  of  Prof.  Faye's  researches,  ('Tenth  Annual  Report    of   Births, 

in    'British    and    Foreign    Medico-  Deaths,  &c.,   in  Scotland,'  1867,  p. 

Chirurg.  Review,'  April,  1867,  pp.  xxviii.)  that  "These  examples  may 

343,    345.     For    France,    the   '  An-  "  suffice    to    shew    that,   at    almost 

nuaire     pour    I'An    1867,'    p.    213.  "  every  stage   of  life,  the   males   in 

For     Phihidelphia,     Dr.     Stockton-  "  Scotland  have  a  greater  liability 

Hough,  'Social  Science  Assoc'  1874.  "  to  death  and  a  higher  death-rate 

For  the  Cape  of  Good  Hope,  Quetelet  "  than  the  females.     The  foct,  how- 

as  quoted  by  Dr.  H.  H.  Zouteveen,  "  ever,    of    this    peculiarity    being 

in    the    Dutch  Translation    of  this  "  most  strongly  developed   at   that 

work   (vol.  i.  p.  417),  where   much  "  infantile  period  of  life  when  the 

information  is  given  on  the  propor-  "  dress,  food,  and  genei-al  treatment 

tion  of  the  sexes.  "  of  both  sexes  are  alike,  seems  to 

^*>  In  regard  to  the  Jews,  see  M.  "  prove  that  the  higher  male  death- 

Thury,  'La  Loi  de  Production  des  "  rate  is  an  impressed,  natural,  and 

Sexes,'  1863,  p.  25.  "  constitutional   peculiarity  due   to 

*^  '  British    and  Foreign  Medico-  "  sex  alone." 
Chirurg.    Review,'  April,    1867,   p. 


244  The  Descent  of  Man.  Part  II. 

variable  in  structure  than  the  female;  and  variations  in  im- 
portant organs  would  generally  be  injurious.  But  the  size  of 
the  body,  and  especially  of  the  head,  being  greater  in  male  than 
female  infants  is  another  cause;  for  the  males  are  thus  more 
liable  to  be  injured  during  parturition.  Consequently  the  still- 
born males  are  more  numerous ;  and,  as  a  higlily  competent  judge. 
Dr.  Crichton  Browne,^^  believes,  male  infants  often  suffer  in  health 
for  some  years  after  birth.  Owing  to  this  excess  in  the  death- 
rate  of  male  children,  both  at  birth  and  for  some  time  sub- 
sequently, and  owing  to  the  exposure  of  grown  men  to  various 
dangers,  and  to  their  tendency  to  emigrate,  the  females  in  all 
old-settled  countries,  where  statistical  records  have  been  kept,^^ 
are  found  to  preponderate  considerably  over  the  males. 

It  seems  at  first  sight  a  mysterious  fact  that  in  different 
nations,  under  different  conditions  and  cHmates,  in  Naples, 
Prussia,  Westphalia,  Holland,  France,  England  and  the  United 
States,  the  excess  of  male  over  female  births  is  less  when  they 
are  illegitimate  than  when  legitimate.^*  This  has  been  explained  by 
different  writers  in  many  different  ways,  as  from  the  mothers 
being  generally  young,  from  the  large  proportion  of  first  preg- 
nancies, &c.  But  we  have  seen  that  male  infants,  from  the  large 
size  of  their  heads,  suffer  more  than  fehiale  infants  diu'ing 
parturition ;  and  as  the  mothers  of  illegitimate  children  must  be 
more  liable  than  other  women  to  undergo  bad  labours,  from 
various  causes,  such  as  attempts  at  concealment  by  tight  lacing, 
hard  work,  distress  of  mind,  &c.,  their  male  infants  would 
proportionably  suffer.  And  this  probably  is  the  most  eflaeient 
of  all  the  causes  of  the  proportion  of  males  to  females  born 
alive  being  less  amongst  illegitimate  children  than  amongst  the 
legitimate.  With  most  animals  the  greater  size  of  the  adult 
male  than  of  the  female,  is  due  to  the  stronger  males  having 
conquered  the  weaker  in  their  struggles  for  the  possession  of 
the  females,  and  no  doubt  it  is  owing  to  this  fact  that  the  two 
sexes  of  at  least  some  animals  differ  in  size  at  birth.     Thus 

*2  <  West  Riding  Lunatic  Asylum  Paraguay,  according  to  the  accurate 

Reports,' vol.  i.  1871,  p.  8.     Sir  J.  Azara  ('Voyages   dans    I'Amerique 

Simpson  has  proved  that  the  head  merid.'  tom.  ii.  1809,  p.  60,   179), 

of  the  male  infant  exceeds  that  of  the  women  are  to  the  men  in  the' 

the  female  by  3-8ths  of  an  inch  in  proportion  of  14  to  13. 
circumference,    and     by    l-8th    in  ^^  Babbage,   '  Edinburgh  Journal 

transverse  diameter.     Quetelet  has  of  Science,'  1829,  vol.  i.  p.  88;  also 

shewn  that  woman  is  born  smaller  p.   90,   on   still-born  children.     On 

than   man;    see    Dr.   Duncan,   '  Fe-  illegitimate    children    in    England, 

cundity,  Fertility,  Sterility,'  1871,  see    'Report    of    Registrar-General 

P-  382.  for  1866,'  p.  xv. 

*^  With  the  savage  Guaranys  of 


Chap.  VJII.  Proportion  of  the  Sexes.  245 

M^e  have  the  ctirions  fact  that  we  may  attribute  the  more 
frequent  deaths  of  male  than  female  infants,  especially  amongst 
the  illegitimate,  at  least  in  part  to  sexual  selection. 

It  has  often  been  supposed  that  the  relative  age  of  the  two 
parents  determines  the  sex  of  the  offspring ;  and  Prof.  Leuckart^'^ 
has  advanced  what  he  considers  sufficient  evidence,  with  respect 
to  man  and  certain  domesticated  animals,  that  this  is  one  impor- 
tant though  not  the  sole  factor  in  the  result.  So  again  the  period 
of  impregnation  relatively  to  the  state  of  the  female  has  been 
thought  by  some  to  be  the  efficient  cause ;  but  recent  observa- 
tions discountenance  this  belief.  According  to  Dr.  Stockton- 
nough,^6  the  season  of  the  year,  the  poverty  or  wealth  of  the 
parents,  residence  in  the  country  or  in  cities,  the  crossing  of 
foreign  immigrants,  &c.,  all  influence  the  proportion  of  the 
sexes.  With  mankind,  polygamy  has  also  been  supposed  to  lead 
to  the  birth  of  a  greater  proportion  of  female  infants ;  but  Dr.  J. 
Campbell  ^^  carefully  attended  to  this  subject  in  the  harems  of 
Siam,  and  concludes  that  the  proportion  of  male  to  female  births 
is  the  same  as  from  monogamous  unions.  Hardly  ^ny  animal 
has  been  rendered  so  highly  polygamous  as  the  English  race- 
horse, and  we  shall  immediately  see  that  his  male  and  female 
offspring  are  almost  exactly  equal  in  number.  I  will  now  give 
the  facts  which  I  have  collected  with  respect  to  the  proportional 
numbers  of  the  sexes  of  various  animals ;  and  will  then  briefly 
discuss  how  far  selection  has  come  into  play  in  determining  the 
result. 

Horses. — Mr.  Tegetraeier  Las  been  so  kind  as  to  tabulate  for  me  from 
the  '  Kacing  Calendar '  the  births  of  race-horses  during  a  period  of 
twenty-one  years,  viz.,  from  184G  to  1867;  1849  being  omitted,  as  no 
returns  were  that  year  published.  The  total  births  were  25,560,^*  con- 
sisting of  12,763  males  aiid  12,797  females,  or  iti  tlie  proportion  of  997 
males  to  100  females.  As  these  numbers  are  tolerably  large,  and  as 
they  are  drawn  from  all  parts  of  England,  during  several  years,'we  may 
with  much  confidence  conclude  that  with  the  domestic  horse,  or  at 
least  with  the  race-horse,  the  two  sexes  are  produced  in  almost  equal 
numbers.     The  fluctuations  in  the  proportions  during  successive  years 

"  Leuckart  (in  Wagner   'Hand-  notice,    as    shewing    how    infertile 

wortorbuch  der  Phys.'  B.  iv.  1853,  these    highly-nurtured    and    rather 

s.  774.  closely-inteibred   animals    have  be- 

5"  Social  Science  Assoc,  of  Phila-  come,  that  not  far  from  one-third  of 

delphia,  1874.  the  mares  failed  to  produce  living 

"     '  Anthropological        Review,'  foals.     Thus  during  186G,  809  male 

April,  1870,  p.  cviii.  colts  and  816  female  colts  were  born, 

^8  During  eleven   years  a  record  and    743    mares   failed   to    produce 

was   kept   of  the  number  of  mares  offspring.     During  1867,  836  males 

which  proved  barren  or  prematurely  and  902  females  were  born,  and  794 

slipped  their  foals ;  and  it  deserves  mares  failed. 


246  The  Descent  of  Man.  Part  11. 


are  closely  like  those  which  occur  with  mankind,  when  a  small  and 
thinly-populated  area  is  considered;  thus  in  1856  the  male  horses  were 
as  107"1,  and  in  18o7  as  only  92-6  to  100  females.  In  the  tabulated 
returns  the  proportions  vary  in  cycles,  for  the  males  exceeded  the 
females  during  six  successive  years ;  and  the  females  exceeded  the 
males  during  two  periods  each  of  four  years :  this,  however,  may  be 
accidental ;  at  least  I  can  detect  nothing  of  the  kind  with  man  in  the 
decennial  table  in  the  Eegistrar's  Report  for  1866. 

DoQs. — During  a  period  of  twelve  years,  from  1857  to  1868,  the  births 
of  a  large  number  of  greyhounds,  throughout  England,  were  sent  to 
the  '  Field'  newspaper ;  and  I  am  again  indebted  to  Mr.  Tegetmeier  for 
carefully  tabulating  the  results.  The  recorded  births  were  6878, 
consisting  of  3605  males  and  3273  females,  that  is,  in  the  proportion  of 
llO'l  males  to  100  females.  The  greatest  fluctuations  occurred  in 
1864,  when  the  proportion  was  as  95-3  males,  and  in  1867,  as  116-3 
males  to  100  females.  The  above  average  proportion  of  llO'l  to  100  ia 
probably  nearly  correct  in  the  case  of  the  greyhound,  but  whether  it 
would  hold  with  other  domesticated  breeds  is  iu  some  degree  doubtful. 
Mr.  Cupples  has  enquired,  from  several  great  breeders  of  dogs,  and  iinds 
that  all  without  exception  believe  that  females  are  produced  in  excess; 
but  he  suggests  that  this  belief  may  have  arisen  from  females  being 
less  valued,  and  from  the  consequent  disappointment  producing  a 
stronger  impression  on  the  mind. 

QUee-p. — The  sexes  of  sheep  are  not  ascertained  by  agriculturists  until 
several  months  after  birth,  at  the  period  when  the  males  are  castrated ; 
so  that  the  following  returns  do  not  give  the  proportions  at  birth. 
Moreover,  I  find  that  several  great  breeders  in  Scotland,  who  annually 
raise  some  thousand  sheep,  are  firmly  convinced  that  a  larger  proportion 
of  males  than  of  females  die  during  the  first  year  or  two.  Therefore  the 
proportion  of  males  would  be  somewhat  larger  at  birth  than  at  the  age  of 
castration.  This  is  a  remarkable  coincidence  with  what,  as  we  ijave 
seen,  occurs  with  mankind,  and  both  cases  probably  depend  on  the 
same  cause.  I  have  received  returns  from  four  gentlemen  in  England 
who  have  bred  Lowdand  sheep,  chiefly  Leicesters,  during  the  last  ten  to 
sixteen  years;  they  amount  altogether  to  8965  births,  consisting  of 
4407  males  and  4558  females;  that  is  in  the  proportion  of  96  7  males  to 
100  females.  With  respect  to  Cheviot  and  black-faced  sheep  bred  in 
Scotland,  I  have  received  returns  from  six  breeders,  two  of  them  on  a 
large  scale,  chiefly  for  the  years  1867-1869,  but  some  of  the  returns 
extend  back  to  1862.  The  total  number  recorded  amounts  to  50,685, 
consisting  of  25,071  males  and  25,614  females,  or  in  the  proportion  of 
97.9  males  to  100  females.  If  we  take  the  English  and  Scotch  returns 
together,  the  total  number  amounts  to  59,650,  consisting  of  29,478 
males  and  30,172  females,  or  as  97*7  to  100.  So  that  with  sheep  at  the 
age  of  castration  the  females  are  certainly  in  excess  of  the  males,  but 
probably  this  would  not  hold  good  at  birth.^^ 

Of  Cattle  I  have  received  returns  from  nine  gentlemen  of  982  births, 
too  few  to  be  trusted ;  these  consisted  of  477  bull-calves  and  505  cow- 


^3  I    am    much    indebted    to  Mr.  tion  to  the  premature  deaths  of  the 

Cupples  for  having  procured  for  me  males, — a    statement    subsequently 

the  above  returns  from  Scotland,  as  confirmed    by    Mr.    Aitcliison    and 

well  as  some  of  the  following  re-  others.     To   this  latter    gentleman, 

turns  on  cattle.     Mr.  R.  Elliot,  of  and  to  Mr.  Payan,  I  owe  my  thanks 

Laio-hwood,  first    called    my  atten-  for  large  returns  as  to  sheep. 


Chap.  VIII.  Proportion  of  the  Sexes.  l.X'j 


calves ;  i.e.,  in  the  proportion  of  91-4  males  to  100  females.  The  Rev. 
W.  D.  Fox  informs  me  that  in  18G7  out  of  84  calves  born  on  a  farm  in 
Derbyshire  only  one  was  a  bull.  Mr.  Harrison  Weir  has  enquired  from 
several  breeders  of  P/(/.*,  and  most  of  them  estimate  the  male  to  the 
female  births  as  about  7  to  6.  This  same  gentleman  has  bred  liahhits 
for  many  years,  and  has  noticed  that  a  far  greater  number  of  bucks  aie 
produced  than  does.     But  estimations  are  of  little  value. 

Of  mammalia  in  a  state  of  nature  I  have  been  able  to  learn  very 
little.  In  regard  to  the  common  rat,  I  have  received  conflicting 
statements.  Mr,  R.  Elliot,  of  Laighwood,  informs  me  that  a  rat-catchei 
assured  him  that  he  had  always  found  the  males  in  great  excess,  even 
with  the  young  in  the  nest.  In  consequence  of  this,  Mr.  Elliot 
Jiimself  subsequently  examined  some  hundred  old  ones,  and  found  the 
statement  true.  Mr.  F.  Buckland  has  bred  a  large  number  of  while 
rats,  and  he  also  believes  that  the  males  greatly  exceed  the  females. 
Tn  regard  to  Moles,  it  is  said  that  "  the  males  are  much  more  numerous 
"  than  the  females  ;"^''  and  as  the  catching  of  these  animals  is  a  special 
occupation,  the  statement  may  perhaps  be  trusted.  Sir  A.  Smith,  in 
describing  an  antelope  of  S.  Africa*'^  (Kobus  ellipsiprymnus),  remarks, 
that  in  the  herds  of  this  and  other  species,  the  males  are  few  in  number 
compared  with  the  females  :  the  natives  believe  that  they  are  born  in 
this  proportion ;  others  believe  that  the  younger  males  arc  expelled 
from  the  herds,  and  Sir  A.  Smith  says,  that  though  he  has  himself 
never  seen  herds  consisting  of  young  males  alone,  others  affirm  tiiat 
this  does  occur.  It  appears  probable  that  the  young  when  expelled 
from  the  herd,  would  often  fall  a  prey  to  the  many  beasts  of  prey  of  the 
country. 

BIRDS. 

With  respect  to  the  Fowl,  I  have  received  only  one  account,  namely, 
that  out  of  1001  chickens  of  a  highly-bred  fctock  of  Cochins,  reared 
during  eight  years  by  Mr.  Stretch,  487  proved  males  and  514  females; 
i.e.,  as  94-7  to  100.  In  regard  to  domestic  pigeons  tliere  is  good 
evidence  either  that  the  males  are  produced  in  excess,  or  that  they  live 
longer ;  for  these  birds  invariably  pair,  and  single  males,  as  Mr.  Teget- 
raeier  informs  me,  can  always  be  purchased  cheaper  than  females. 
Usually  the  two  birds  reared  from  the  two  eggs  laid  in  the  san\e  nest 
are  a  male  and  a  female  ;  but  Mr.  Harrison  Weir,  who  has  been  so  lar-^e 
a  breeder,  says  that  he  has  often  bred  two  cocks  from  the  same  nest, 
and  seldom  two  hens  ;  moreover,  the  hen  is  generally  the  weaker  of  the 
two,  and  more  liable  to  perish. 

With  respect  to  birds  in  a  state  of  nature,  Mr.  Gould  and  others^^ 
are  convinced  that  the  males  are  generally  the  more  numerous ;  and 
as  the  young  males  of  many  species  resemble  the  females,  the  latter 
would  naturally  appear  to  be  the  more  numerous.  Large  numbers  of 
pheasants  are  reared  by  Mr.  Baker  of  Leadenhall  from  eggs  laid  by  wild 
birds,  and  he  informs  Mr.  Jenner  Weir  that  four  or  five  males  to  one 
female  are  generally  produced.     An  experienced  observer  remarks." 


so  Bell,  *  History  of  Briti.sh  Quad-  iv.  s.  990)  comes  to   the  same  oon- 

rupeds,'  p.  lUO.  clu.sioa. 

61  '  Illustrations    of   the  Zoology  e^  Qq  j-j^g  authority  of  L.  Lloyd, 
of  S.  Africa,'  1849,  pi.  29.  'Game  Bird.s  of  Sweden,'  18G7,  pp. 

62  Brehm  ('  Illust.  Thierleben,'  B.  12,  133 


248  T]ie  Descent  of  Man.  Paet  II. 

that  in  Scandinavia  the  broods  of  the  capercailzie  and  black-cock 
contain  more  males  than  females ;  and  that  with  the  Dal-ripa  (a  kind 
of  ptarmigan)  more  males  than  females  attend  the  lelis  or  places  of 
courtship ;  but  this  latter  circumstance  is  accounted  for  by  some 
observers  by  a  greater  number  of  hen  birds  being  killed  by  vermin. 
From  various  facts  given  by  White  of  Selborne,"*  it  seems  clear  that 
the  males  of  the  partridge  must  be  in  considerable  excess  in  the  south 
of  England ;  and  1  have  been  assured  that  this  is  the  case  in  Scotland. 
Mr.  Weir  on  enquiring  from  the  dealers,  who  receive  at  certain  seasons 
large  numbers  of  rufl's  {Machetes  pugnax),  was  told  that  the  males  are 
much  the  more  numerous.  This  same  naturalist  has  also  enquired  for 
me  from  the  birdcatchers,  who  annually  catch  an  astonishing  number 
of  various  small  species  alive  for  the  London  market,  and  he  was  un- 
hesitatingly answered  by  an  old  and  trustworthy  man,  that  with  the 
chaffinch  the  males  are  in  large  excess;  he  thought  as  high  as  2  males  to 
1  female,  or  at  least  as  high  as  5  to  3.^^  The  males  of  the  blackbird, 
he  likewise  maintained,  were  by  far  the  more  numerous,  whether  • 
caught  by  traps  or  by  netting  at  night.  These  statements  may 
apparently  be  trusted,  because  this  same  man  said  that  the  sexes  are 
about  equal  with  the  lark,  the  twite  (^Linaria  montana)^  and  goldfinch. 
On  the  other  hand,  he  is  certain  that  with  the  common  linnet,  the 
females  preponderate  greatly,  but  unequally  during  different  years  ; 
during  some  years  he  has  foimd  the  females  to  the  males  as  four  to  one. 
It  should,  however,  be  borne  in  mind,  that  the  chief  season  for  catching 
birds  does  net  begin  till  September,  so  that  with  some  species  partial 
migrations  may  have  begun,  and  the  flocks  at  this  period  often  consist 
of  hens  alone.  Mr.  Salvin  paid  particular  attention  to  the  sexes  of  the 
humming-birds  in  Central  America,  and  he  is  convinced  that  with 
most  of  the  species  the  males  are  in  excess;  thus  one  year  he  procured 
204  specimens  belonging  to  ten  species,  and  these  consisted  of  1G6 
males  and  of  only  38  females.  With  two  other  species  the  females  were 
in  excess :  but  the  proportions  apparently  vary  either  during  ditferent 
seasons  or  in  difterent  localities;  for  on  one  occasion  the  males  of 
Campylopterus  hemileucurus  were  to  the  females  as  5  to  2,  and  on 
another  occasion "^"^  in  exactly  the  reversed  ratio.  As  bearing  on  this 
latter  point,  I  may  add,  that  Mr.  Powys  found  in  Corfu  and  Epirus 
the  sexes  of  the  chaffinch  keeping  apart,  and  "  the  females  by  far  the 
"  most  numerous;"  whilst  in  Palestine  Mr.  Tristram  found  "the  male 
'•  flocks  appearing  greatly  to  exceed  the  female  in  number.""^'  So 
again  with  the  Quiscalus  major,  Mr.  G.  Taylor***  says,  that  in  Florida 
there  were  "  very  few  females  in  proportion  to  the  males,"  whilst  in 
Honduras  the  proportion  was  the  other  way,  the  species  there  having 
the  character  of  a  poiygamiot. 


"  'Nat.  Hist,  of  Selborne,'  letter  ever  caught  by  one  man  in  a  single 

xxix.  edit,  of  1825,  vol.  i.  p.  139.  day  was  70. 

®5  Mr.     Jenner     Weir     received  ^^  '  Ibis,'  a'oI.  ii.  p.  260,  as  quoted 

similar  information,  on  making  en-  in    Gould's    '  Trcchilidae,'    1861,    p. 

quiries  during   the   following  year.  5'2.     For  the  foregoing  proportions, 

To  shew  the  number  of  living  chaf-  I  am   indebted  to  Mr.  Salvin   for   a 

finches  caught,  I  may  mention  that  table  of  his  results. 
in  1869  the^re  was  a  match  between  "^  'Ibis,'  1860,  p.  137;  and  1867, 

two  experts,   and    one  man  caught  p.  369. 
in  a  day  62,  and   another  40,  male  "*'  '  Ibis,'  1862,  p.  137. 

chaffinches.     The    greatest    number 


Chap.  VIII.  Proportion  of  the  Sexes,  249 


FISH. 

With  Fish  the  proportional  numbers  of  the  sexes  can  be  asfertnined 
only  by  catching  them  in  the  adult  or  neaily  adult  state  ;  and  there 
are  many  difficulties  in  arriving  at  any  just  conclusion.'^^  Infertile 
fojuales  might  readily  be  mistaken  for  males,  as  Dr.  Giinther  has 
remarked  to  me  in  regard  to  trout.  With  some  species  the  males  are 
believed  to  die  soon  after  fertilising  the  ova.  Witli  many  species  the 
mules  are  of  much  smaller  size  than  the  females,  so  that  a  large 
number  of  males  would  escape  from  the  same  net  by  which  the  females 
were  caught.  M.  Carbounier/"  who  has  especially  attended  to  the 
natural  history  of  the  pike  {E^x^x  lucins),  states  that  many  males,  owing 
to  tiieir  small  size,  are  devoured  by  the  larger  females  ;  and  he  believes 
that  the  males  of  almost  all  fish  are  exposed  from  ihis  same  cause  to 
greaier  danger  than  the  females.  Nevertheless,  in  the  few  cases  in 
which  the  proportional  numbers  have  been  actually  observed,  the 
males  appear  tj  be  largely  in  excess.  Thus  Mr.  R.  Uuist,  the  sujierin- 
tendent  of  the  Stormontfield  experiments,  says  that  in  1865,  out  of  70 
salmon  first  landed  for  the  purpose  of  oblaining  the  ova,  upwards  of  60 
were  males.  In  1867  he  again  "  calls  attention  to  the  vast  disproportion 
•'  of  the  males  to  the  females.  We  had  at  the  outset  at  least  ten  males 
"  to  one  female."  Afterwards  females  sufficient  for  obtaining  ova  were 
procured.  He  adds,  "  from  the  great  proportion  of  the  males,  they  are 
"constantly  fighting  and  tearing  each  other  on  the  spawning-beds."^' 
This  disproportion,  no  doubt,  can  be  accounted  for  in  part,  but  whether 
wholly  is  doubtful,  by  the  males  ascending  the  rivers  before  the 
females.  Mr.  F.  Buckland  remarks  in  regard  to  trout,  that  "  it  is  a 
"curious  fact  that  the  males  preponderate  very  largely  in  number  over 
*'  the  females.  It  invariably  happens  that  when  the  first  rush  of  fish  is 
"  made  to  the  net,  there  will  be  at  least  seven  or  ei^ht  males  to  one 
"female  found  captive.  I  cannot  quite  account  for  this;  either  the 
*'  males  nre  more  numerous  than  the  females,  or  the  latter  seek  safety 
"  by  concealment  rather  than  flight."  He  then  adds,  that  by  carefully 
searching  the  banks,  sufficient  females  for  obtaining  ova  can  be  found.'^ 
Mr.  H.  Lee  informs  me  that  out  of  212  trout,  taken  for  this  purpose  in 
Lord  Portsmouth's  park,  150  were  males  and  62  females. 

The  males  of  the  Cyprinidae  likewise  seem  to  be  in  excess ;  but 
several  members  of  this  Family,  viz.,  the  carp,  tench,  brtam  and 
minnow,  appear  regularly  to  follow  the  practice,  rare  in  the  animal 
kingdom,  of  polyandry ;  for  the  female  whilst  spawning  is  always 
attended  by  two  males,  one  on  each  side,  and  in  the  case  of  the  bream 
by  three  or  four  males.  This  fact  is  so  well  known,  that  it  is  always 
recommended  to  stock  a  pond  with  two  male  tenches  to  one  female,  or 
at  least  with  three  males  to  two  females.  With  the  minnow,  an 
excellent  observer  states,  that  on  the  spawning-beds  the  males  are  ten 
times  as  numerous  as  the  females;  when  a  female  comes  amongst  the 


«3  Leuckart  quotes  Bloch  (Wag-  18,  1869,  p.  3G9. 

ner,    '  Handworterbuch    der    Phys.'  "  '  The     Stormontfield    Piscicul- 

B.  iv.  1853,  s.  775),  that  with  fish  tural    Experiments,'    186G,    p.    23. 

there   are  twice  as  many  males   as  The  '  Field '  newspaper,  June  29th, 

females.  1867. 

^0  Quoted  in  the  'Farmer,'  March  "  '  Land  and  Water,'  1868,  p.  41. 
12 


250  The  Descent  of  Man.  Part  11. 

males,  *'  she  is  imraediatelj'  pressed  closely  by  a  male  oa  each  side ; 
"and  when  they  have  been  in  that  situation  for  a  time,  are  superseded 
"  by  other  two  males."  "^ 

INSECTS. 

In  this  great  Class,  the  Lepidoptera  almost  alone  afford  means  for 
judging  of  the  proportional  numbers  of  the  sexes  ;  for  they  have  been 
collected  with  special  care  by  many  good  observers,'  and  have  been 
largely  bred  from  the  egg  or  caterpillar  state.  I  had  hoped  that  some 
breeders  of  silk-moths  might  have  kept  an  exact  record,  but  after 
writing  to  France  and  Italy,  and  consulting  various  treatises,  I  cannot 
hud  that  this  has  ever  been  done.  Tiie  general  opinion  appears  to  be 
that  the  sexes  are  nearly  equal,  but  in  Italy,  as  I  hear  from  Professor 
Canestrini,  many  breeders  are  convinced  that  the  females  are  produced 
in  excess.  This  same  naturalist,  however,  informs  me,  that  in  the  two 
yearly  broods  of  the  Ailanthus  silk-moth  (Bomhijx  cynthia),  the  males 
greatly  preponderate  in  the  first,  whilst  in  the  second  the  two  sexes  are 
nearly  equal,  or  the  females  rather  in  excess. 

In  regard  to  Butterflies  in  a  state  of  nature,  several  obseivers  have 
been  much  struck  by  the  apparently  enormous  proponderance  of  the 
males.'^  Thus  Mr.  JBates,'^  in  speaking  of  several  species,  about  a 
hundred  in  number,  which  inhabit  the  Upper  Amazons,  says  that  the 
males  are  much  more  numerous  than  the  females,  even  in  the  propor- 
tion of  a  hundred  to  one.  In  North  America,  Edwards,  who  had  great 
experience,  estimates  in  the  genus  Papilio  the  males  to  the  females  as 
four  to  one ;  and  Mr.  Walsh,  who  informed  me  of  this  statement,  says 
that  with  P.  turims  this  is  certainly  the  case.  In  South  Africa,  Mr.  K. 
Trimen  found  the  males  in  excess  in  19  species ;""  and  in  one  of  these, 
which  swarms  in  optn  places,  he  estimated  the  number  of  males  as 
fifty  to  one  female.  With  another  speoes,  in  which  the  males  are 
numerous  in  certain  localities,  lie  collected  only  five  feuiales  during 
seven  years.  In  the  island  of  Bourbon,  M.  IVLdllard  states  that  the 
males  of  one  species  of  Papilio  are  twenty  times  as  numerous  as  the 
females.^^  Mr.  Trimen  informs  me  that  as  far  as  he  has  himself  seen, 
or  heard  from  others,  it  is  rare  for  the  females  of  any  butterfly  to 
exceed  the  males  in  number ;  but  three  South  African  species  per- 
haps oifer  an  exception.  Mr.  Wallace  '*  states  that  the  females  of 
Orintlwjjtera  croesus,  in  the  Malay  archipelago,  are  more  common  and 
more  easily  caught  than  the  males ;  but  this  is  a  rare  butterfly.     I  may 


'  "  Yarrell,  '  Hist.  British  Fishes,'  or   four  times   as   numerous  as  the 

vol.  i.  1826,  p.  307  ;  on  the  Cyprhvis  females. 

ca/'jDiO,  p.  331;  on  the  Tmcavu/gram,  "  'The  Naturalist  on  the  Ama- 

p.  331  ;   on  the  Ahmmis  braina,  p.  zons,'  vol.  ii.  18B3,  p.  228,  347. 
336.     See,   for    the    minnow   (Leu-  "«  Four  of  the.se  cases  are  given 

ciscus  phoxin'is),  '  Loudon's  Mag.  of  by  Mr.  Trimen   in  his  '  Rhopalocera 

Nat.  Hist.'  vol.  v.  1832,  p.  682.  Africae  Australis.' 

7*     Leuckart      quotes     Meinecke  "  Quoted  by  Trimen,  '  Transact. 

(Wagner,     '  Handworterbuch      der  Ent.  Soc' vol.  v.  part  iv.  1866,  p.  330. 
Phys.'    B.    iv.    1853,    s.    775)   that  '*  '  Transact.  Liun.  Soc' vol.  xxv. 

the  males  of  Butterflies  are  three  p.  37. 


Chap.  VIII.  Proportion  of  the  Sexes.  251 

here  add,  that  in  Hyperytlira,  a  genus  of  moths,  Guenee  says,  that 
from  four  to  five  females  are  sent  in  collections  from  India  for  one 
male. 

When  this  subject  of  the  proportional  numbi^rs  of  the  sexes  of  insects 
was  brought  before  the  Entomological  Society,'"  it  was  generally 
admitted  that  the  males  of  most  Leiiidoptera,  in  the  adult  or  ima^io 
state,  are  caugijt  in  greater  numbers  than  the  females :  but  this  fact 
was  attributed  by  various  observers  to  the  more  retiring  habits  of  the 
females,  and  to  the  males  emerging  earlier  from  the  cocoon.  This 
latter  circumstance  is  well  known  to  occur  with  most  Lepidoptcra.  ns 
well  as  with  other  insects.  So  that,  as  M.  Pcr.-^onnat  remarks,  the 
males  of  the  domesticated  Bombyx  Yamamai,  aie  useless  at  the  begin- 
ning of  the  Season,  and  the  females  at  the  end,  from  the  want  of 
mates.*"  I  cannot,  however,  persuade  myself  that  these  causes  suffice  to 
explain  tl.e  great  excess  of  males,  in  the  above  cases  of  ccrtnin  butter- 
flies which  are  extremely  common  in  their  native  countries.  Mr. 
Stainton,  who  has  paid  very  close  attention  during  many  years  to  the 
smaller  moths,  informs- me  that  when  he  collected  them  in  the  imago 
state,  he  thought  that  the  males  were  ten  times  as  numerous  as  the 
females,  but  that  since  ho  has  reared  them  on  a  huge  scale  from  the 
caterpillar  state,  he  is  convinced  that  the  females  are  the  liiore 
numerous.  Several  entomologists  concur  in  this  view.  Mr.  Double- 
day,  however,  and  some  others,  take  an  opposite  view,  and  are  con- 
vinced that  they  have  reared  from  tlie  eggs  and  caterpillars  a  larger 
proportion  of  males  than  of  females. 

Besides  the  more  active  habits  of  the  males,  their  earlier  emergence 
from  the  cucoon,  and  in  some  cases  their  frequenting  more  open 
stations,  other  causes  may  be  assigned  for  an  apparent  or  real  difference 
in  the  propnrtional  numbers  of  the  sexes  of  Lepidoptera,  when  cap- 
tured in  the  imago  state,  and  when  reared  from  the  egg  or  caterpillar 
state.  I  hear  from  Professor  Canestrini,  that  it  is  believed  by  many 
breeders  in  Italy,  that  tiie  female  caterpillar  of  the  silk-moth  suffers 
more  from  the  recent  disease  than  the  male  ;  and  Dr.  Staudinger 
informs  me  that  in  rearing  Lepidoptera  more  females  die  in  the 
cocoon  than  males.  With  many  species  the  female  caterpilliir  is  la'ger 
than  the  male,  and.  a  collector  would  naturally  choose  the  finest 
specimens,  and  thus  unintentionally  collect  a  larger  number  of  females. 
Three  collectors  have  told  me  that  this  was  their  practice ;  but  Dr. 
Wallace  is  sure  that  most  collectors  take  all  the  specimens  which  they 
can  find  of  the  rarer  kinds,  which  alone  are  worth  the  trouble  of 
rearing.  Birds  when  surrounded  by  caterpillars  would  probably 
devour  the  largest ;  and  Professor  Canestrini  inlbrms  me  that  in  Italy 
some  breeders  believe,  though  on  insufficient  evidence,  that  in  the  first 
broods  of  the  Ailanthus  siik-moth,  the  wasps  destroy  a  larger  number  of 
the  female  than  of  the  male  caterpillars.  Dr.  Wallace  further  remarks 
that  female  caterpillars,  from  being  larger  than  the  males,  require 
more  time  for  their  development,  and  consume  more  food  and  mois- 
ture ;  and  thus  they  would  be  exposed  during  a  longer  time  to 
danger  from  ichneumons,  birds,  &c.,  and  in  times  of  scarcity  would 
perish  in   greater  numbers.     Hence   it    appears   quite  possible    that 


^^  *  Proc.    Eutomolog.    Soc'  Feb.      '  Proc.  Eat.  Soc'  3rd  series,  vol.  v. 
I7th,  1868.  1867,  p.  487. 

8f   Quoted    by    Dr.     Wallace    in 


252  T J le  Descent  of  Man,  Pakt  II. 

in  a  fctate  of  nature,  fewer  female  LepidoptDra  may  reach  maturity 
tiian  males ;  and  for  our  special  object  we  are  concerned  with  their 
relative  numbers  at  maturity,  when  the  sexes  are  leady  to  propagate 
their  land. 

The  manner  in  which  Iho  males  of  certain  moths  congregjate  in 
extraordinary  numbers  round  a  single  female,  apparently  indicates  a 
great  excess  of  males,  though  this  fact  may  perhaps  bo  accounted  for 
by  ihe  earlier  emergence  of  the  males  from  their  cocoons.  Mr. 
Stainton  informs  me  that  from  twelve  to  twenty  males,  may  often  be 
seen  congregattd  round  a  female  Elachida  rufocinerea.  It  is  well 
known  that  if  a  virgin  Lasiocampa  quercus  or  Saiurnia  carpiiti 
be  exposed  in  a  cage,  vast  numbers  of  males  collect  round  her,  and  if 
confined  in  a  room  will  even  come  down  the  chimney  to  htr.  Mr. 
Doubleday  believes  that  he  has  seen  from  iifty  to  a  hundred  males  of 
both  these  species  attracted  in  the  course  of  a  single  day  by  a  ft  male 
in  confinement.  In  the  Isle  of  Wight  Mr.  Trimen  exposed  a  box 
in  which  a  female  of  the  Lasiocampa  hid.  been  confined  on  the 
previous  day,  and  five  males  soon  endeavoured  to  gain  admittance. 
In  Australia,  M.  Yerreuux,  having  placed  the  female  of  a  small 
Bombyx  in  a  box  in  his  pocket,  was  followed  by  a  crowd  of  males,  so 
that  about  200  entered  the  house  with  him.*^ 

Mr.  Doubleday  has  called  my  attention  to  M.  Staudinger's  ®-  litt 
of  Lepidoptera,  which  gives  the  prices  of  the  males  and  females  of 
300  species  or  well-marked  varieties  of  butterflies  (Ehopalocera).  The 
prices  for  both  sexes  of  the  very  common  species  are  of  course  the  same ; 
but  in  114  of  the  rarer  species  they  ditier;  the  males  being  in  all  cases, 
excepting  one,  the  cheaper.  On  an  average  of  the  prices  of  the  113  species, 
the  price  of  the  male  to  that  of  the  female  is  as  100  to  149  ;  and  this 
apparently  indicates  that  inversely  the  males  exceed  the  females  in 
the  same  proportion.  About  2000  species  or  varieties  of  moths 
(Heterocem)  are  catalogued,  those  with  wingless  females  being  here 
excluded  on  account  of  the  difference  in  habits  between  the  two  sexes  : 
of  these  2000  species,  141  differ  in  price  according  t)  i-ex,  the  males 
of  130  being  cheaper,  and  those  of  oirly  11  being  dearer  than  the 
females.  The  average  price  of  the  males  of  the  130  species,  to  that  of 
the  females,  is  as  lOu  to  143.  With  respect  to  the  buttertiies  in  this 
priced  litt,  Mr.  Doubleday  thinks  (and  no  man  in  England  has  had 
more  experience),  that  there  is  nothing  in  the  habit:,  of  the  species 
which  tan  account  for  the  difference  in  the  prices  of  the  two  sexes, 
and  that  it  Ciin  be  accounted  for  only  by  an  excess  in  the  number  of 
the  malts.  But  I  am  bound  to  add  that  Dr.  Staudinger  informs  me, 
that  he  is  h'mself  of  a  different  opinion.  He  thinks  that  the  less  active 
habits  of  the  females  and  the  earlier  emergence  of  the  males  will 
account  for  his  collectors  securing  a  larger  number  of  males  than 
of  females,  and  consequently  for  the  lower  prit-es  of  the  former.  With 
respect  to  specimens  rtared  from  the  caterpillar-state.  Dr.  Staudinger 
believes,  as  previously  stated,  that  a  greater  number  of  females  than  of 
males  die  whilst  confined  in  the  cocoons.  He  adds  that  with  certain 
species  one  sex  seems  to  preponderate  over  the  oth(  r  during  certain 
ye.irs. 

Of  direct  observations   on  the  sexes  of  Lepidoptera,  reared  either 

'^    Blanchard,     '  Me'taraorphoses,  ^^         '  Lepidopteren  -  Doubletten 

Moeurs  des  lasectes,'  1868,  pp.  225-      Liste,'  Berlin,  Ko.  x.  1866. 
226. 


Chap.  VII  I. 


Propo7'tion  of  the  Sexes. 


253 


from  eggs   or  caterpillars,  I   have   received  only  the   few   following 


Males.     !  Females. 


The  Rev.  J.  Hellins*^  of  Exeter  reared,  during  1868,|i 

imagos  of  73  species,  which  consisted  of.  .  ./j 
Mr.  Albert  Jones  of  Eltham   reared,   during    1868,^ 

imagos  of  9  species,  which  consisted  of  .  .  .  )\ 
During   18G9  he  reared   imagos  from  4  species,  con-i! 

sitting  of /I 

Mr.    Buckler    of    Emsworth,    Hants,    during    1869,11 

reared  imagos  from  74:  species,  consisting  of.  .  J 1 
Dr.  Wallace  of  Colchester  reared  from  one  brood  of ll 

Bombyx  cynthia j  i 

Dr.  Wallace  raised,  from  cocoons  of  Bombyx  Pernyin 

sent  from  China,  during  1869 jj 

Dr.  Wallace  raised,  during  1868  and  1869,  from  twolj 

lots  of  cocoons  of  Bombyx  yama-mai        .       .      .Jl 


Total 


153  I 

159  I 

114  j 

ISO  ; 

224 

i 

934  I 


137 
126 
112 
169 

48 
123 

46 

761 


So  that  in  these  eight  lots  of  cocoons  and  eggs,  males  were  produced 
in  excess.  Taken  together  the  proportion  of  males  is  as  \1tl 
to  100  females.  Bat  the  numbers  are  hardly  large  enough  to  be 
trustworthy. 

On  the  whole,  from  these  various  sources  of  evidence,  all  pointing 
in  the  same  direction,  I  infer  that  with  most  species  of  Lepidopteia, 
the  mature  males  generally  exceed  the  females  in  number,  whatever 
the  proportions  may  be  at  their  first  emergence  from  tlie  o.^^^. 

\s\\\\  reference  to  the  other  Orders  of  insects,  I  have  been  able 
to  collect  very  little  reliable  information.  With  the  stag-l^ee  le 
{Lucaiuis  cervus)  "tlie  males  appear  to  be  much  more  numerous 
"than  the  females;"  but  when,  as  Cornelius  remarked  during  18(37, 
sin  unusual  number  of  these  beetles  appeared  in  one  i)art  of  C'erinany, 
the  females  appeared  to  exceed  the  males  as  six  to  one.  With  one  of 
the  Elateridae,  the  males  are  said  to  be  much  more  numerous  than  the 
females,  and  "two  or  thiee  are  often  found  united  with  one  female  ;^^ 
"  so  that  here  polyandry  seems  to  prevail."  With  Siagonium  (!St  iphy- 
linidoe),  in  which  the  males  are  furnished  with  horns,  "  t  .e  females  are 
"  far  more  numerous  than  the  opposite  sex."  Mr.  Janson  stated  at  the 
Entumological  Society  that  the  females  of  the  btrk- feeding  Tomicns 
villosus  are  so  common  as  to  be  a  plague,  whilst  the  males  are  so  rare 
jis  to  be  hardly  known. 


'^  This  naturalist  has  been  so 
kind  as  to  send  me  some  results 
from  former  years,  in  which  the 
females  seemed  to  preponderate  ; 
but  so  many  of  the  figures  were 
estimates,  that  I  found  it  impossible 
to  tabulate  them. 


**  Giinther's  '  Record  of  Zoo- 
logical Literature,'  1867,  p.  260. 
On  the  excess  of  female  Liicanus, 
ibid.  p.  250.  On  the  males  of  Luca- 
nus  in  England,  Westwood,  'Modern 
Class,  of  Insects,'  vol.  i.  p.  187.  On 
the  Siagouium,  ibid.  p.  172. 


254  1^^^  Desce7it  of  Man.  Part  II. 

It  is  liardly  worth  while  saymg  anything  about  the  proportion  of 
the  sexes  in  certain  spe 'ies  and  even  groups  of  insects,  for  the  males 
are  unknown  or  very  rare,  and  the  females  are  parthenogenetic,  that 
is,  fertile  without  sexual  rmion ;  examples  of  this  are  afforded  by 
several  of  the  Cynipida^.*^  In  all  the  gall-making  Cynipidfe  known 
to  INIr.  Walth,  the  females  are  four  or  live  times  as  numerous  as  the 
males  ;  and  so  it  i.s,  as  he  informs  me,  with  the  gall-making  Cecidomyiire 
(Diptera).  With  some  common  i-pecies  of  Saw-flies  (Tenthredinai) 
Mr.  F.  Smith  has  reared  hundreds  of  specimens  from  larvae  of  ail 
sizes,  but  has  never  reared  a  single  male :  c^n  the  other  hand,  Curtis 
say:<,*^  that  with  certain  species  (Athalia),  bred  by  him,  the  males  were 
t(»  the  females  as  six  to  one;  whilst  exactly  the  reverse  occurred  with 
the  matiue  insects  of  the  same  species  caught  in  the  fields.  In  the 
family  of  Bees,  Hermann  iMiiller.*^  collected  a  large  number  of 
specimens  of  mfiny  s|  ecies,  and  reared  others  from  tl.e  cocouns,  and 
counted  the  texes.  He  found  thut  the  males  of  some  species  gieatly 
exceeded  the  females  in  number  ;  in  others  the  reverse  occurred ;  and 
in  others  the  two  sexes  were  nearly  equal.  But  as  in  most  cases  the 
males  emerge  from  the  cocoons  before  the  females,  they  are  at  the 
comme.'icement  ,ot"  the  breeding  seaton  practically  in  excess.  Miiller 
also  observed  that  the  relative  number  of  the  two  sexes  in  s  ^me 
species  diftered  much  in  different  localities.  But  as  H.  Miiller  has 
himself  remarked  to  me,  these  remarks  must  be  received  with 
some  caution,  as  one  sex  might  more  easily  escape  observation  than 
the  other.  Thus  his  brother  Fritz  INIiiller  has  noticed  in  Brazil  that 
the  two  sexes  of  the  same  species  of  bee  sometimes  frequent  different 
kinds  of  lluwers.  "With  respect  to  the  Orthoptera,  I  know  hardly 
anything  about  the  relative  number  cf  the  sexe^ :  Korte,®^  however, 
says  that  out  of  500  locusts  which  he  examine  1,  the  males  were  to 
the  females  as  five  to  six.  With  the  Neuroptera,  Mr.  Walth  states 
that  in  many,  but  by  no  means  in  all  the  species  of  the  Odonatous 
group,  there  is  a  great  overplus  of  males  :  in  thg  genus  Heta^rina,  also, 
the  males  are  generally  at  least  four  times  as  numerous  as  the  females. 
In  certaiii  species  in  the  genus  Gomphus  the  males  are  equally  in 
excess,  whilbt  in  two  other  spe  'ies,  the  females  are  twice  or  thrice 
as  numerous  as  the  males.  In  some  European  species  of  Psocus 
thousands  of  females  may  be  collected  without  a  single  male,  whilst 
with  oilier  species  of  the  same  genus  both  sexes  are  common,^^  In 
England,  Mr.  ]\IacLachlan  has  captured  hundreds  of  the  female 
Apatania  muliebris,  but  has  never  seen  t!,e  male;  and  of  Boreus 
hyfrtKilis  on\j  four  or  five  males  have  been  seen  here."**  With  m<;8t 
of  tin  se  species  (excepting  the  Tenthredinre)  there  is  at  present  no 
evidence  that  the  females  are  subject  to  parthenogenesis;  and  thus  we 
see  how  ignorant  we  are  of  the  causes  of  the  apparent  discrepancy  in 
the  proportion  of  the  two  sexes. 

In  ti.e  other  Classes  of  the  Articulata  I  have  been  able  to  collect  still 


8^  Walsh,  in  'The  American  En-  derhenschrecke,'  1828,  p.  20. 

tomologist,'    vol.    i.    1869,    p.    103.  *®  '  Observations  on  X.  American 

F.    Smith,    'Record    of    Zoological  Neuroptera,' by  H.  Hagen  and  B.  D. 

Literature,'  1867,  p.  328.  Walsh,     '  Proc.     Ent.     Soc.    Phila- 

'"'  '  Farm  Insects,'  pp.  45-46.  delphia,'   Oct.   1863,  pp.   168,  223, 

*'  *  Anwendung  der  Darwinschen  239. 

Lehre  Verb.  d.  n.  V.  Jahrg.  x.xiv.'  ^°  '  Proc.  Ent.  Soc.  London,'  Feb. 

«8  'Die  Strich,   Zuir    oder   Wan-  17,1868. 


Chap.  VIII.  Proportion  of  the  Sexes.  255 

loss  information.  With  Spiilers,  Mr.  Blackwall,  who  has  carefully 
attended  to  this  class  during  many  years,  writes  to  me  that  the  male's 
from  their  more  erratic  habits  are  more  commonly  seen,  and  therefore 
appeur  more  numerous.  This  is  actually  the  case  with  a  few  species  ; 
but  he  mentions  several  species  in  six  genera,  in  wiiich  the  females 
ap[)ear  to  be  much  more  numerous  than  the  males.**'  'I'he  small  size  of 
the  males  in  comparison  with  the  females  (a  peculiarity  which  is  some- 
times carried  to  an  extreme  degree),  and  their  widely  ditierent  appear- 
ance, may  account  in  some  instances  for  their  rarity  in  collections.''^ 

Some  of  the  lower  Crustaceans  ai-e  able  to  propigate  their  kind 
nsexually,  and  this  v/ill  account  for  the  extreme  larity  of  the  males  : 
thus  Von  SiebokP^  carefully  examined  no  less  than  1:3,000  specimens  of 
Apus  from  twenty-one  localities,  and  amongst  these  he  found  only 
319  males.  AV^itii  some  other  forms  (as  Tanais  and  Cypris),  as  Fritz 
Miillt-r  informs  me,  there  is  reason  to  believe  that  the  males  are  much 
shorter-lived  than  the  females ;  and  this  would  explain  their  scarcity, 
supposing  the  two  sexes  to  be  at  first  equal  in  number.  On  the  other 
hand,  Miiller  has  invariably  taken  far  more  males  than  females  of  the 
Diastylidc-e  and  of  Cypridina  on  the  shores  of  Brazil;  thus  witli  a 
species  in  the  latter  genus,  (33  specimens  caught  the  same  day  included 
57  males ;  but  he  suggests  that  this  preponderance  may  be  due  to 
some  unknown  difference  in  the  habits  of  the  two  sexes.  With  one 
of  the  higher  Brazilian  crabs,  n;tmely  a  Celasimus,  Fritz  Miiller 
found  the  males  to  be  more  numerous  than  the  females.  According 
to  the  Inrge  experience  of  Mr.  C.  Spence  Bate,  the  reverse  seems  to 
be  the  case  with  six  common  British  crabs,  the  names  of  which  he 
has  given  me. 

The  proportion  of  the  sexes  in  relation  to  natural  selection. 

There  is  reason  to  suspect  that  in  some  cases  man  has  by 
selection  indirectly  influenced  his  own  sex-producing  jwwers. 
Certain  women  tend  to  produce  during  their  whole  lives  more 
children  of  one  sex  than  of  the  other  :  and  the  same  holds  good 
of  many  animals,  for  instance,  cows  and  horses  ;  thus  Mr.  Wright 
of  Yelderslcy  House  informs  me  that  one  of  his  Arab  mares, 
though  put  seven  times  to  different  horses,  produced  seven 
fillies.  Though  I  have  very  little  evidence  on  this  head,  analogy 
would  lead  to  the  belief,  that  the  tendency  to  produce  either 
sex  would  be  inherited  like  almost  every  other  peculiarity,  for 
instance,  that  of  producing  twins ;  and  concerning  the  above 
tendency  a  good  authority,  Mr.  J.  Downing,  has  communicated 
to  me  facts  which  seem  to  prove  that  this  does  occur  in  certain 
families  of  short-horn  cattle.  Col.  Marshall  ^^  has  recently  found 
on  careful  examination  that  the  Todas,  a  hill-tribe  of  India, 

^*  Another  great  authority  with  0.  P.  Cambridge,  as  quoted  in 
respect  to  this  class,  Prof. Thorell  of  'Quarterly  Journal  of  Science,' 
Upsala     ('  On     European     Spiders,'       18G8,  p.  429. 

1869-70,  part  i.  p.  205)  speaks  as  if  ^^  i  Beitrage  zur  Parthenogenesis,' 

female  spiders  wei'e  generally  com-      p.  174. 
moner  than  the  males.  9*    *  The    Todas,'    1873,  pp.    100, 

»2  See,     on     this     subject,     Mr,       111,  194,  196, 


256  The  Descent  of  Man.  Part  II. 

consist  of  112  males  and  84  females  of  all  ages— that  is  in  a  ratio 
of  133-3  males  to  100  females.  The  Todas,  who  are  polyandrous 
in  their  marriages,  during  former  times  invariably  practised 
female  infanticide  ;  bnt  this  practice  has  now  been  discontinued 
for  a  considerable  period.  Of  the  children  born  within  late  years, 
the  males  are  more  numerous  than  the  females,  in  the  proportion 
of  121:  to  100.  Colonel  Marshall  accounts  for  this  fact  in  the 
following  ingenious  manner.  "  Let  us  for  the  purpose  of  illustra- 
"  tion  take  tln'ee  families  as  I'epresenting  an  average  of  the 
"  entire  tribe  ;  say  that  one  mother  gives  birth  to  six  daughters 
'*  and  no  sons ;  a  second  mother  has  six  sons  only,  whilst  the 
"  third  mother  has  three  sons  and  three  daughters.  The  first 
"  mother,  following  the  tribal  custom,  destroys  four  daughters 
"  and  preserves  two.  The  second  retains  her  six  sons.  The  third 
"  kills  two  daughters  and  keeps  one,  as  also  her  three  sons.  We 
"  have  then  from  the  three  families,  nine  sons  and  three  daughters, 
"  with  which  to  continue  the  breed.  But  whilst  the  males 
"  belong  to  families  in  which  the  tendency  to  produce  sons  is 
"  great,  the  females  are  of  those  of  a  converse  inclination.  Thus 
"  the  bias  strengthens  with  each  generation,  until,  as  we  find, 
"  families  grow  to  have  habitually  more  sons  than  daughters." 

That  this  result  would  follow  from  the  above  form  of  infanticide 
seems  almost  certain  ;  that  is  if  we  assume  that  a  sex-producing 
tendency  is  inherited.  But  as  the  above  numbers  are  so  ex- 
tremely scanty,  I  have  searched  for  additional  evidence,  but 
cannot  decide  whether  wliat  I  have  found  is  trustworthy ; 
nevertheless  the  facts  are,  i)erhaps,  worth  giving.  The  Maories  of 
New  Zealand  have  long  practised  infanticide  ;  and  Mr.  Fenton^^ 
states  that  he  "  has  met  with  instances  of  women  who  have  de- 
"  stroyed  four,  six,  and  even  seven  children,  mostly  females. 
"  However,  the  universal  testimony  of  those  best  qualified  to 
"  judge,  is  conclusive  that  this  custom  has  for  many  years  been 
"  almost  extinct.  Probably  the  year  1885  may  be  named  as  the 
"  period  of  its  ceasing  to  exist.'^  Now  amongst  the  New  Zea- 
landers,  as  with  the  Todas,  male  births  are  considerably  in  excess. 
Mr.  Fenton  remarks  (p.  30),  "  One  fact  is  certain,  although  the 
"  exact  period  of  the  commencement  of  this  singular  condition  of 
"  the  disproportion  of  the  sexes  cannot  be  demonstratively  fixed, 
"  it  is  quite  clear  that  this  course  of  decrease  was  in  full  opera- 
"  tion  during  the  years  1S30  to  1844,  when  the  non-adult 
''  population  of  1844  was  being  produced,  and  has  continued 
"  with  great  energy  up  to  the  present  time."  The  following 
statements  are  taken  from  Mr.  Fenton  (jd,  26),  but  as  the  numbers 

^^  'Aboriginal  Inhabitants  of  New  Zealand  ;  Government  Report,'  1859, 
p.  36. 


Chap.  VIII.  Proportion  of  the  Sexes.  257 

are  not  large,  and  as  the  census  was  not  accurate,  uniform 
results  cannot  be  expected.  It  should  be  borne  in  mind  in  this 
and  the  following  cases,  that  the  normal  state  of  every  population 
is  an  excess  of  women,  at  least  in  all  civilised  countries,  chiefly 
owing  to  the  greater  mortality  of  the  male  sex  during  youth,  and 
partly  to  accidents  of  all  kinds  later  in  life.  In  1858,  the 
native  population  of  New  Zealand  was  estimated  as  consisting 
of  31,607  males  and  21,303  females  of  all  ages,  that  is  in  the 
ratio  of  1303  males  to  100  females.  But  during  this  same  year, 
and  in  certain  limited  districts,  the  numbers  were  ascertained 
with  much  care,  and  the  males  of  all  ages  were  here  753 
and  the  females  616 ;  that  is  in  the  ratio  of  122'2  males  to  100 
females.  It  is  more  important  for  us  that  during  this  same 
year  of  1858,  the  non-adult  males  within  the  same  district 
were  found  to  be  178,  and  the  uon-adidt  females  142,  that  is  in 
the  ratio  of  125*3  to  100.  It  may  be  added  that  in  1844,  at 
which  pariod  female  infanticide  had  only  lately  ceased,  the 
nou-adult  males  in  one  district  were  281,  and  the  non-adult 
females  only  191,  that  is  in  the  ratio  of  144-8  males  to  100  females. 

In  the  Sandwich  Islands,  the  males  exceed  the  females  in 
number.  Infanticide  was  formerly  practised  there  to  a  frightful 
extent,  but  was  by  no  means  confined  to  female  infants,  as 
is  shewn  by  Mr.  EUis,^''  and  as  I  have  been  informed  by  Eishop 
Staley  and  the  Eev.  Mr.  Coan.  Nevertheless,  another  apparently 
trustworthy  writer,  Mr.  Jarves,^^  whose  observations  apply  to 
the  whole  archiiDelago,  remarks: — "Numbers  of  women  are  to 
"  be  found,  who  confess  to  the  murder  of  from  three  to  six  or  eight 
"  children ;"  and  he  adds,  "  females  from  being  considered  less 
"  useful  than  males  were  more  often  destroyed."  From  what  is 
kno^vn  to  occur  in  other  parts  of  the  world,  this  statement  is 
probable ;  but  must  be  received  with  much  caution.  The 
practice  of  infanticide  ceased  about  the  year  1819,  when  idolatry 
was  abolished  and  missionaries  settled  in  the  Islands.  A  careful 
census  in  1839  of  the  adult  and  taxable  men  and  women  in  the 
island  of  Kauai  and  in  one  district  of  Oahu  (Jarves,  p.  404), 
gives  4723  males  and  3776  females;  that  is  in  the  ratio  of 
125'08  to  100.  At  the  same  time  the  number  of  males  under 
fourteen  years  in  Kauai  and  under  eighteen  in  Oahu  was  1797, 
and  of  females  of  the  same  ages  1429 ;  and  here  we  have  the 
ratio  of  12575  males  to  100  females. 

In  a  census  of  all  the  islands  in  1850,^*^  the  males  of  all  ages 

^^  'Narrative  of  a  Tour  through  ^*  This  is  given  in  the  Rev.  H.  T. 

Hawaii,'  1826,  p.  298.  Cheever's  '  Life  in  the  Sandwich  Is- 

»^     '  History    of     the     Sandwich  lands,'  1851,  p.  277. 
Islands,'  1843,  p.  93. 


258 


The  Descent  of  Man. 


Paet  II. 


amount  to  36,272,  and  the  females  to  33,128,  or  as  109-49  to 
100.  The  males  under  seventeen  years  amounted  to  10,773,  and 
the  females  under  the  same  age  to  9593,  or  as  112*3  to  100. 
From  the  census  of  lfe72,  the  proportion  of  males  of  all  ages 
(including  half-castes)  to  females,  is  as  125*36  to  100.  It  must 
be  borne  in  mind  that  all  these  returns  for  the  Sandwich 
Islands  give  the  proportion  of  living  males  to  living  females, 
and  not  of  the  births ;  and  judging  from  all  civilised  countries 
the  proportion  of  males  would  have  been  considerably  higher  it 
the  numbers  had  referred  to  births.^^ 

From  the  several  foregoing  cases  we  have  some  reason  to 
believe  that  infanticide  practised  in  the  manner  above  explained, 
tends  to  make  a  male-producing  race ;  but  I  am  far  from  sup- 
posing that  this  practice  in  the  case  of  man,  or  some  analogous 
process  with  other  species,  has  been  the  sole  determining  cause 
of  an  excess  of  males.  There  may  be  some  unknown  law  leading 
to  this  result  in  decreasing  races,  which  have  already  become 
somewhat  infertile.      Besides    the    several    causes    previously 


^^  Dr.  Coulter,  in  describing 
('  Journal  R.  Geograph.  Soc.,'  vol. 
V.  1835,  p.  67)  the  state  of  Cali- 
fornia about  the  year  1830,  says 
that  the  natives,  reclaimed  by  the 
Spanish  missionaries,  have  nearly 
all  perished,  or  are  perishing,  al- 
though well  treated,  not  driven 
from  their  native  land,  and  kept 
from  the  use  of  spirits.  He  at- 
tributes this,  in  great  part,  to  the 
undoubted  fact  that  the  men  greatly 
exceed  the  women  in  number ;  but 
he  does  not  know  whether  this  is 
due  to  a  failure  of  female  offspring, 
or  to  more  females  dying  during 
early  youth.  The  latter  alternative, 
according  to  all  analogy,  is  very 
improbable.  He  adds  tiiat  "  in- 
"  fanticide,  properly  so  called,  is 
"  not  common,  though  very  fre- 
"  quent  recourse  is  had  to  abor- 
"  tion."  If  Dr.  Coulter  is  correct 
about  infjinticide,  this  case  cannot 
be  advanced  in  support  of  Col. 
Marshall's  view.  From  the  rapid 
decrease  of  the  reclaimed  natives, 
we  may  suspect  that,  as  in  the 
cases  lately  given,  their  fertility 
has  been  diminished  from  changed 
habits  of  life. 

I  had  hoped  to  gain  some  light 


on  this  subject  from  the  breeding 
of  dogs;  inasmuch  as  in  most  breeds, 
with  the  exception,  perhaps,  of 
greyhounds,  many  more  female 
puppies  are  destroyed  than  males, 
just  as  with  the  Toda  infants.  Mr. 
Cupples  assures  me  that  this  is 
usual  with  Scotch  deer-hounds. 
Unfortunately,  I  know  nothing  of 
the  proportion  of  the  sexes  in  any 
breed,  excepting  greyhounds,  and 
there  the  male  births  are  to  the 
female  as  110-1  to  100.  Now  from 
enquiries  made  from  many  breeders, 
it  seems  that  the  females  are 
in  some  respects  more  esteemed, 
though  otherwise  troublesome  ;  and 
it  does  not  appear  that  the  female 
puppies  of  the  best-bred  dogs  are 
systematically  destroyed  more  than 
the  males,  though  this  does  sometimes 
take  place  to  a  limited  extent.  There- 
fore 1  am  unable  to  decide  whether 
we  can,  on  the  above  principles,  ac- 
count for  the  preponderance  of  male 
births  in  greyhounds.  On  the  other 
hand,  we  have  seen  that  with 
horses,  cattle,  and  sheep,  which  are 
too  valuable  for  the  young  of  either 
sex  to  be  destroyed,  if  (;here  is  any 
difference,  the  females  are  slightly 
in  excess. 


Chap.  VIII.  Proportion  of  tJie  Sexes.  259 


alluded  to,  the  greater  facility  of  parturition  amongst  savages, 
and  the  less  consequent  injury  to  their  male  infants,  would 
tend  to  increase  the  proportion  of  live-born  males  to  females. 
There  does  not,  however,  seem  to  be'  any  necessary  connection 
between  savage  life  and  a  marked  excess  of  males ;  that  is  if  we 
may  judge  by  the  character  of  the  scanty  offspring  of  the  lately 
existing  Tasmanians  and  of  the  crossed  offspring  of  the  Tahitians 
now  inhabiting  Norfolk  Island. 

As  the  males  and  females  of  many  animals  differ  somewhat  in 
habits  and  are  exposed  in  different  degrees  to  danger,  it  is 
probable  that  in  many  cases,  more  of  one  sex  than  of  the  other 
are  habitually  destroyed.  But  as  far  as  I  can  trace  out  the  com- 
plication of  causes,  an  indiscriminate  though  large  destruction 
of  either  sex  would  not  tend  to  modify  the  sex-producing  power 
of  the  species.  With  strictly  social  animals,  such  as  bees  or  ants, 
which  produce  a  vast  number  of  sterile  and  fertile  females  in 
comparison  with  the  males,  and  to  whom  this  preponderance  is 
of  paramount  importance,  we  can  see  that  those  communities 
would  flourish  best  which  contained  females  having  a  strong 
inherited  tendency  to  produce  more  and  more  females ;  and  in 
such  cases  an  unequal  sex-producing  tendency  would  be  ulti- 
mately gained  through  natural  selection.  With  animals  living 
in  herds  or  troops,  in  which  the  males  come  to  the  front  and 
defend  the  herd,  as  with  the  bisons  of  North  America  and  certain 
baboons,  it  is  conceivable  that  a  male-producing  tendency  might 
be  gained  by  natural  selection ;  for  the  individuals  of  the  better 
defended  herds  would  leave  more  numerous  descendants.  In 
the  case  of  mankind  the  advantage  arising  from  having  a  pre- 
ponderance of  men  in  the  tribe  is  supposed  to  be  one  chief  cause 
of  the  practice  of  female  infanticide. 

In  no  case,  as  far  as  we  can  see,  would  an  inherited  tendency 
to  produce  both  sexes  in  equal  numbers  or  to  produce  one  sex 
in  excess,  be  a  direct  advantage  or  disadvantage  to  certain 
individuals  more  than  to  others ;  for  instance,  an  individual 
with  a  tendency  to  produce  more  males  than  females  would  not 
succeed  better  in  the  battle  for  life  than  an  individual  with  an 
opposite  tendenc}'" ;  and  therefore  a  tendency  of  this  kind  could 
not  be  gained  through  natural  selection.  Nevertheless,  there  are 
certain  animals  (for  instance,  fishes  and  cirripedes)  in  which  two 
or  more  males  appear  to  be  necessary  for  the  fertilisation  of  the 
female ;  and  the  males  accordingly  largely  preponderate,  but  it 
is  by  no  means  obvious  how  this  male-producing  tendency  could 
have  been  acquired.  I  formerly  thought  that  when  a  tendency 
to  produce  the  two  sexes  in  equal  numbers  was  advantageous  to 
the  species,  it  would   follow  from  natural  selection,  but  I  now 


26o  The  Descent  of  Man.  Part  II. 

see  that  the  whole  problem  is  so  intricate  that  it  is  safer  to  leave 
its  solution  for  the  future. 


CHAPTER  IX. 

Secondary  Sexual  Characters  in  the  Lower  Classes  of 
THE  Animal  Kingdom. 

These  characters  absent  in  the  lowest  classes — Brilliant  colours — MoUusca 
— Annelids — Crustacea,  secondary  sexual  characters  strongly  developed  ; 
dimorphism;  colour;  characters  not  acquired  before  maturity — Spiders, 
sexual  colours  of;  stridulation  by  the  males — Myriapoda. 

With  animals  belonging  to  the  lower  classes,  the  two  sexes 
are  not  rarely  united  in  the  same  individual,  and  therefore 
secondary  sexual  characters  cannot  be  developed.  In  many 
cases  where  the  sexes  are  separate,  both  are  permanently  at- 
tached to  some  support,  and  the  one  cannot  search  or  struggle 
for  the  other.  Moreover  it  is  almost  certain  that  these  animals 
have  too  imperfect  senses  and  much  too  low  mental  powers,  to 
appreciate  each  other's  beauty  or  other  attractions,  or  to  feel 
rivalry. 

Hence  in  these  classes  or  sub-kingdoms,  such  as  the  Protozoa, 
Coelenterata,  Echinodermata,  Scolecida,  secondary  sexual  cha- 
racters, of  the  kind  which  we  have  to  consider,  do  not  occur ;  and 
this  fact  agrees  with  the  belief  that  such  characters  in  the 
higher  classes  have  been  acquired  through  sexual  selection, 
which  depends  on  the  will,  desire,  and  choice  of  either  sex. 
Nevertheless  some  few  apparent  exceptions  occur ;  thus,  as  I 
hear  from  Dr.  Eaird,  the  males  of  certain  Entozoa,  or  internal 
parasitic  worms,  differ  slightly  in  colour  from  the  females ;  but 
we  have  no  reason  to  suppose  that  such  differences  have  been 
augmented  through  sexual  selection.  Contrivances  by  which  the 
male  holds  the  female,  and  which  are  indispensable  for  the 
propagation  of  the  species,  are  independent  of  sexual  selection, 
and  have  been  acquired  through  ordinary  selection. 

Many  of  the  lower  animals,  whether  hermaphrodites  or  with 
separate  sexes,  are  ornamented  with  the  most  brilliant  tints,  or 
are  shaded  and  striped  in  an  elegant  manner ;  for  instance,  many 
corals  and  sea-anemones  (Actiniae),  some  jelly-fish  (Medusae, 
Porpita,  &c.),  some  Planarise,  many  star-fishes.  Echini,  Ascidians, 
&c. ;  but  we  may  conclude  from  the  reasons  already  indicated, 
namely  the  union  of  the  two  sexes  in  some  of  these  animals,  the 
permanently  affixed  condition  of  others,  and  the  low  mental 
powers  of  all,  that  such  colours  do  not  serve  as  a  sexual 
attraction,  and  have  not  been  acquired  through  sexual  selection. 


Chap.  IX.  Sexual  Selection.  26 1 

It  should  be  borne  in  mind  that  in  no  case  have  we  suffi- 
cient evidence  that  colours  have  been  thus  acquired,  ex- 
cept where  one  sex  is  much  more  brilliantly  or  conspicuously 
coloured  than  the  other,  and  where  there  is  no  difference 
in  habits  between  the  sexes  sufficient  to  account  for  their 
different  colours.  But  the  evidence  is  rendered  as  complete 
as  it  can  ever  be,  only  when  the  more  ornamented  indivi- 
duals, almost  always  the  males,  voluntarily  display  their 
attractions  before  the  other  sex  ;  for  we  cannot  believe  that  such 
display  is  useless,  and  if  it  be  advantageous,  sexual  selection 
will  almost  inevitably  follow.  AVe  may,  however,  extend  this 
conclusion  to  both  sexes,  when  coloured  alike,  if  their  colours  are 
plainly  analogous  to  those  of  one  sex  alone  in  certain  other 
species  of  the  same  group. 

How,  then,  are  we  to  account  for  the  beautiful  or  even 
gorgeous  colours  of  many  animals  in  the  lowest  classes?  It 
appears  doubtful  whether  such  colours  often  serve  as  a  protec- 
tion ;  but  that  we  may  easily  err  on  this  head,  will  be  admitted 
by  every  one  who  reads  Mr.  Wallace's  excellent  essay  on  this 
subject.  It  would  not,  for  instance,  at  first  occur  to  auy  one 
that  the  transparency  of  the  Medusae,  or  jelly-fishes,  is  of  the 
highest  service  to  them  as  a  protection;  but  when  we  are 
reminded  by  Hackel  that  not  only  the  medusae,  but  many 
floating  mollusca,  crustaceans,  and  even  small  oceanic  fishes 
partake  of  this  same  glass-like  appearance,  often  accompanied 
by  prismatic  colours,  we  can  hardly  doubt  that  they  thus 
escape  the  notice  of  pelagic  birds  and  other  enemies.  M. 
Girard  is  also  convinced  ^  that  the  bright  tints  of  certain 
sponges  and  ascidians  serve  as  a  protection.  Conspicuous 
colours  are  likewise  beneficial  to  many  animals  as  a  warning  to 
their  would-be  devourers  that  they  are  distasteful,  or  that  they 
possess  some  special  means  of  defence ;  but  this  subject  will  be 
discussed  more  conveniently  hereafter. 

We  can,  in  our  ignorance  of  most  of  the  lowest  animals,  only 
say  that  their  bright  tints  result  either  from  the  chemical 
nature  or  the  minute  structure  of  their  tissues,  independently  of 
any  benefit  thus  derived.  Hardly  any  colour  is  finer  than  that 
of  arterial  blood;  but  there  is  no  reason  to  suppose  that  the 
colour  of  the  blood  is  in  itself  any  advantage;  and  though  it 
adds  to  the  beauty  of  the  maiden's  cheek,  no  one  will  pretend 
that  it  has  been  acquired  for  this  purpose.  So  again  with  many 
animals,  especially  the  lower  ones,  the  bile  is  richly  coloured ; 
thus,  as  I  am  informed  by  Mr.  Hancock,  the  extreme  beauty  of 
the  Eolidae  (naked  sea-slugs)  is  chiefly  due  to  the  biliary  glands 
'  'Archives  dc  Zoolog.  Exper.,'  Oct.  1872,  p.  563. 


262  The  Descejit  of  Man,  Part  U. 

being  seen  through  the  translucent  integuments — this  beauty 
being  probably  of  no  service  to  these  animals.  The  tints  of  the 
decaying  leaves  in  an  American  forest  are  described  by  every 
one  as  gorgeous;  yet  no  one  supposes  that  these  tints  are 
of  the  least  advantage  to  the  trees.  Bearing  in  mind  how  many 
substances  closely  analogous  to  natural  organic  compounds  have 
been  recently  formed  by  chemists,  and  which  exhibit  the  most 
splendid  colours,  it  would  have  been  a  strange  fact  if  substances 
similarly  coloured  had  not  often  originated,  independently  of 
any  useful  end  thus  gained,  in  the  complex  laboratory  of  living 
organisms. 

The  sub-kingdom  of  the  Mollusca. — Throughout  this  great 
division  of  the  animal  kingdom,  as  far  as  I  can  discover, 
secondary  sexual  characters,  such  as  we  are  here  considering, 
never  occur.  Nor  could  they  be  expected  in  the  three  lowest 
classes,  namely  in  the  Ascidians,  Polyzoa,  and  Brachiopods 
(constituting  the  Molluscoida  of  some  authors),  for  most  of 
these  animals  are  permanently  affixed  to  a  support  or  have  their 
sexes  united  in  the  same  individual.  In  the  Lamellibranchiata, 
or  bivalve  shells,  hermaphroditism  is  not  rare.  In  the  next 
higher  class  of  the  Gasteropoda,  or  univalve  shells,  the  sexes  are 
either  united  or  separate.  But  in  the  latter  case  the  males 
never  possess  special  organs  for  finding,  securing,  or  charming 
the  females,  or  for  fighting  with  other  males.  As  I  am  informed 
by  Mr.  Gwyn  Jeffreys,  the  sole  external  difference  between  the 
sexes  consists  in  the  shell  sometimes  differing  a  little  in  form ; 
for  instance,  the  shell  of  the  male  periwinkle  {LUtorina  littorea) 
is  narrower  and  has  a  more  elongated  spire  than  that  of  the 
female.  But  differences  of  this  nature,  it  may  be  presumed,  are 
directly  connected  with  the  act  of  reproduction,  or  with  the 
development  of  the  ova. 

The  Gasteropoda,  though  capable  of  locomotion  and  furnished 
with  imperfect  eyes,  do  not  appear  to  be  endowed  with  sufficient 
mental  powers  for  the  members  of  the  same  sex  to  struggle 
together  in  rivalry,  and  thus  to  acquire  secondary  sexual 
characters.  Nevertheless  with  the  pulmoniferous  gasteropods,  or 
land-snails,  the  pairing  is  preceded  by  courtship;  for  these 
animals,  though  hermaphrodites,  are  compelled  by  their  structure 
to  pair  together.  Agassiz  remarks,^  "  Quiconque  a  eu  I'occasion 
"  d'observer  les  amours  des  lima9ons,  ne  saurait  mettre  en  doute 
"  la  seduction  deployee  dans  les  mouvements  et  les  allures  qui 
"  preparent  et  accomplissent  le  double  embrassement  de  ces 
"  hermaphrodites."  These  animals  appear  also  susceptible  of 
some  degree  of  permanent  attachment :  an  accurate  observer, 
*  '  De  I'Espece  et  de  la  Class.'  &c.,  1869,  p.  106. 


Chap.  IX. 


Molluscs.  263 


Mr.  Lonsdale,  informs  me  that  he  placed  a  pair  of  land-snails, 
{Hdix  j>omatia),  one  of  which  was  weakly,  into  a  small  and  ill- 
provided  garden.  After  a  short  time  the  strong  and  healthy 
individual  disappeared,  and  was  traced  by  its  track  of  slime 
over  a  wall  into  an  adjoining  well-stocked  garden.  Mr. 
Lonsdale  concluded  that  it  had  deserted  its  sickly  mate;  but 
after  an  absence  of  twenty-four  hours  it  returned,  and  apparently 
communicated  the  result  of  its  successful  exploration,  for  both 
then  started  along  the  some   track  and  disappeared  over  the 

wall. 

Even  in  the  highest  class  of  the  Mollusca,  the  Cephalopoda  or 
cuttlefishes,  in  which  the  sexes  are  separate,  secondary  sexual 
characters  of  the  present  kind  do  not,  as  far  as  I  can  discover, 
occur.  This  is  a  surprising  circumstance,  as  these  animals 
possess  highly-developed  sense-organs  and  have  considerable 
mental  powers,  as  will  be  admitted  by  every  one  who  has  watched 
their  artful  endeavours  to  escape  from  an  enemy.^  Certain 
Cephalopoda,  however,  are  characterised  by  one  extraordinary 
sexual  character,  namely,  that  the  male  clement  collects  within 
one  of  the  arms  or  tentacles,  which  is  then  cast  off,  and  clinging 
by  its  sucking-discs  to  the  female,  lives  for  a  time  an  independent 
life.  So  completely  does  the  cast-off  arm  resemble  a  separate 
animal,  that  it  was  described  by  Cuvier  as  a  parasitic  worm 
under  the  name  of  Plectocolyle.  But  this  marvellous  structure 
may  be  classed  as  a  primary  rather  than  as  a  secondary  sexual 
character. 

Although  with  the  Mollusca  sexual  selection  does  not  seem  to 
have  come  into  play;  yet  many  univalve  and  bivalve  shells, 
such  as  volutes,  cones,  scallops,  &c.,  are  beciutifully  coloured 
and  shaped.  The  colours  do  not  appear  in  most  cases  to  be  of 
any  use  as  a  protection ;  they  are  probably  the  direct  result,  as 
in  the  lowest  classes,  of  the  nature  of  the  tissues ;  the  patterns 
and  the  sculpture  of  the  shell  depending  on  its  manner  of 
growth.  The  amount  of  light  seems  to  be  influential  to  a  certain 
extent;  for  although,  as  repeatedly  stated  by  Mr.  Gwyn  Jeffreys, 
the  shells  of  some  species  living  at  a  profound  depth  are  brightly 
coloured,  yet  we  generally  see  the  lower  surfaces,  as  well  as  the 
parts  covered  by  the  mantle,  less  highly-coloured  than  the 
upper  and  exposed  surfaces.-*    In  some  cases,  as  with  shells 

3  See,  for  instance,  the    account  influence  of  light  on  the  colours  of 

which  I  have  given  in  my  'Journal  a       I'rondescent     incrustation,     de- 

of  Researches,' 1845,  p.  7.  posited  by  the  surf  on  the    coast- 

*  I  have  given  (' Geolog.  Obser-  rocks  of  Ascension,  and   formed  by 

vations  on  Volcanic  Islands,'  18+4,  the  solution  of  triturated  sea-shells, 
p.   53)   a    curious    instance    of   the 


264  The  DesccJit  of  Man.  Fart  II. 

living  amongst  corals  or  briglitly-tinted  sea-weeds^  the  bright 
colours  may  serve  as  a  protection.^  But  that  many  of  the  nudi- 
brancli  mollusca,  or  sea-slugs,  are  as  beautifully  coloured  as  any 
shells,  may  be  seen  in  Messrs.  Alder  and  Hancock's  magnificent 
work ;  and  from  information  kindly  given  me  by  Mr.  Hancock, 
it  seems  extremely  doubtful  whether  these  colours  usually  serve 
as  a  protection.  With  some  species  this  may  be  the  case,  as  with 
one  kind  which  lives  on  the  green  leaves  of  algse,  and  is  itself 
bright-green.  But  many  brightly-coloured,  white  or  otherwise 
conspicuous  species,  do  not  seek  concealment ;  whilst  again  some 
equally  conspicuous  species,  as  well  as  other  dull-coloured  kinds, 
live  under  stones  and  in  dark  recesses.  So  that  with  these  nudi- 
branch  molluscs,  colour  apparently  does  not  stand  in  any  close 
relation  to  the  nature  of  the  places  which  they  inhabit. 

These  naked  sea-slugs  are  hermaphrodites,  yet  they  pair 
together,  as  do  land-snails,  many  of  which  have  extremely 
pretty  shells.  It  is  conceivable  that  two  hermaphrodites, 
attracted  by  each  other's  greater  beauty,  might  unite  and  leave 
offspring  which  would  inherit  their  parents'  greater  beauty. 
But  with  such  lowly-organised  creatures  this  is  extremely 
improbable.  Nor  is  it  at  all  obvious  how  the  offspring  from  the 
more  beautiful  pairs  of  hermaphrodites  would  have  any  ad- 
vantage over  the  offspring  of  the  less  beautiful,  so  as  to  increase 
in  number,  unless  indeed  vigour  and  beauty  generally  coincided. 
We  have  not  here  the  case  of  a  number  of  males  becoming 
mature  before  the  females,  with  the  more  beautiful  males 
selected  by  the  more  vigorous  females.  If,  indeed,  brilliant 
colours  were  beneficial  to  a  hermaphrodite  animal  in  relation 
to  its  general  habits  of  life,  the  more  brightly-tinted  individuals 
would  succeed  best  and  would  increase  in  number ;  but  this 
would  be  a  case  of  natural  and  not  of  sexual  selection. 

Sub-kingdom  of  the  Vermes:  Class,  Annelida  (or  Sea-ivorms). — 
In  this  class,  although  the  sexes,  when  separate,  sometimes 
differ  from  each  other  in  characters  of  such  importance  that  they 
have  been  placed  under  distinct  genera  or  even  families,  yet  the 
differences  do  not  seem  of  the  kind  which  can  be  safely  at- 
tributed to  sexual  selection.  These  animals  are  often  beauti- 
fully coloured,  but  as  the  sexes  do  not  differ  in  this  respect,  we 
are  but  little  concerned  with  them.  Even  the  Nemertians, 
though  so  lowly  organised,  "  vie  in  beauty  and  variety  of 
"  colouring  with  any  other  group  in  the  invertebrate  series;"  yet 

^  Dr.  Morse  has  lately  discussed      '  Proc.    Boston    Soc.    of  Nat.  Hist.' 
this   subject    in   his    paper    on    the      vol.  xiv.,  April,  1871. 
Adaptive    Coloration    of    Mollusca, 


Chap.  IX.  Crustaceaiis,  265 

Dr.  Mcintosh"  cannot  discoTcr  that  these  colours  are  of  any 
service.  The  sedentary  annelids  become  dnller-colonred,  ac- 
cording to  M.  Quatrefages/  after  the  period  of  reproduction ;  and 
this  I  presume  may  be  attributed  to  their  less  vigorous  condition 
at  that  time.  All  these-worm-like  animals  apparently  stand  too 
low  in  the  scale  for  the  individuals  of  either  sex  to  exert  any 
choice  in  selecting  a  partner,  or  for  the  individuals  of  the  same 
sex  to  struggle  together  in  rivalry. 

Suh-hingdom  of  the  Arthropoda :  Class,  Crustacea, — In  this  great 
class  we  first  meet  with  undoubted  secondary  sexual  characters, 
often  developed  in  a  remarkable  manner.  Unfortunately  the 
habits  of  crustaceans  are  very  imx^erfectly  known,  and  we  cannot 
explain  the  uses  of  many  structures  i3eculiar  to  one  sex.  With  the 
lower  i^arasitic  species  the  males  are  of  small  size,  and  they 
alone  are  furnished  with  perfect  swimming-legs,  antennse  and 
sense-organs ;  the  females  being  destitute  of  these  organs,  with 
their  bodies  often  consisting  of  a  mere  distorted  mass.  But 
these  extraordinary  differences  between  the  two  sexes  are  no 
doubt  related  to  their  widely  different  habits  of  life,  and  con- 
sequently do  not  concern  us.  In  various  crustaceans,  belonging  to 
distinct  families,  the  anterior  antennae  are  furnished  with  Xjeculiar 
thread-like  bodies,  which  are  believed  to  act  as  smelling-organs, 
and  these  are  much  more  numerous  in  the  males  than  in  the 
females.  As  the  males,  without  any  unusual  development  of 
their  olfactory  organs,  would  almost  certainly  be  able  sooner  or 
later  to  find  the  females,  the  increased  number  of  the  smelling- 
threads  has  probably  been  acquired  through  sexual  selection,  by 
the  better  provided  males  having  been  the  more  successful  in 
finding  partners  and  in  producing  offspring.  Fritz  Miiller  lias 
described  a  romarkable  dimorphic  species  of  Tanais,  in  which  the 
male  is  represented  by  two  distinct  forms,  which  never  graduate 
into  each  other.  In  the  one  form  the  male  is  furnished  with 
more  numerous  smelling-threads,  and  in  the  other  form  with 
more  powerful  and  more  elongated  chelse  or  pincers,  which  serve 
to  hold  tlie  female.  Fritz  Miiller  suggests  that  these  differences 
between  the  two  male  forms  of  the  same  species  may  have 
originated  in  certain  individuals  having  varied  in  the  number  of 
the  smelling-threads,  whilst  other  individuals  varied  in  the 
fthape  and  size  of  their  chelae ;  so  that  of  the  former,  those  which 
were  best  able  to  find  the  female,  and  of  the  latter,  those  v/hich 

•  See  his  beautiful  monograph  on  ^  See    M.   Perrier,    '  I'Origine   de 

*  British  Annelids,'  part  i.  1873,  I'Homme  d'apres  Darwin,'  '  Kevue 
p.  3.  Scientifique,'  Feb.  1873,  p.  8G6. 


266 


The  Descent  of  Man. 


Part  II. 


were  best  able  to  hold  her,  have  left  the  greatest  number  of 
progeny  to  inherit  their  respective  advantages.^ 
In  some  of  the  lower  crustaceans,  the  right  anterior  antenna 
of  the  .male  differs  greatly  in  structure 
from  the  left,  the  latter  resembling  in 
its  simple  tapering  joints  the  antennsB 
of  the  female.  In  the  male  the 
modified  antenna  is  either  swollen  in 
the  middle  or  angularly  bent,  or 
converted  (fig.  4)  into  an  elegant, 
and  sometimes  wonderfully  complex, 
prehensile  organ.^  It  serves,  as  I  hear 
from  Sir  J.  Lubbock,  to  hold  the 
female,  and  for  this  same  purpose  one 
of  the  two  posterior  legs  (6)  on  the 
same  side  of  the  body  is  converted 
into  a  forceps.  In  another  family  the 
inferior  or  .  posterior  antennsB  are 
''  curiously  zigzagged "  in  the  males 
alone. 

In  the  higher  crustaceans  the  an- 
terior legs  are  developed  into  chelae 
or  pincers;  and  tlicse  are  generally 
larger  in  the  male  than  in  the  female, 
— so  much  so  that  the  market  value  of 
the  male  edible  crab  (^Caiicer  jpagurus), 
according  to  IMr.  C.  Spence  Bate,  is 
five  times  as  great  as  that  of  the  fe- 
male. In  many  species  the  chelse  are 
of  unequal  size  on  the  opposite  side  of 
the  body,  the  right-hand  one  being,  as 
I  am  informed  by  Mr.  Bate,  generally, 
though  not  invariably,  the  largest.  This  inequality  is  also  often 
much  greater  in  the  male  than  in  the  female.  The  two  chelae 
of  the  male  often  differ  in  structure  (figs.  5,  6,  and  7),  the 
smaller  one  resembling  that  of  the  female.  What  advantage  is 
gained  by  their  inequality  in  size  on  the  opposite  sides  of  the 


4.     LabWocera    Darwinii 
(from  Lubbock). 

a.  Part  of   right    anterior    an- 

tenna of  male,  forming  a 
prehensile  organ. 

b.  Posterior  pair  of  thoracic  legs 

of  male. 

c.  Ditto  of  temale. 


*  *  Facts  and  Arguments  for 
Darwin,'  English  translat.  1869,  p. 
20.  See  the  pi-evious  discussion  on 
the  olfactory  threads.  Sars  has 
described  a  somewhat  analogous 
case  (as  quoted  in  *  Nature,'  1870, 
p.  455)  in  a  Norwegian  crustacean, 
the  Poiitoporeia  affinis. 

^  See  Sir  J.  Lubbock  in  '  Annals 


and  Mag.  of  Nat.  Hist.'  vx)l.  xi. 
1853,  pi.  i.  and  x. ;  and  vol.  xii. 
(1853)  pi.  vii.  See  also  Lubbock  in 
'Transact.  Ent.  Soc'  vol.  iv.  new 
series,  1856-1858,  p.  8.  With  re- 
spect to  the  zig-zagged  antennjE 
mentioned  below,  see  Fritz  Miiller, 
'  Facts  and  Arguments  for  Darwin,' 
1869,  p.  40,  foot-note. 


Chap.  IX. 


Crjistaceans. 


267 


body,  and  by  the  inequality  being  much  greater  in  the  male  than 
in  the  female ;  and  why,  when  they  are  of  equal  size,  both  aro 


Fig.  5.    Anterior  part  of  body  of  Callianassa  (from  Milne-Edwards),  showing  the  un- 
equal and  dififerently-coustructed  right  and  left-hand  chela?  of  the  male. 

N.B— The  artist  by  mistake  has  reversed  the  drawing,  and  made  the  left-hand  chela 
the  largest. 


Fig.  6. 
Fig.  7. 


Fig.  6.  Fig.  Y. 

Second  leg  of  male  Orchestia  Tucuratinga  (from  Fritz  MUller). 
Ditto  of  female. 


often  much  larger  in  the  male  than  in  the  female,  is  not  known. 
As  1  hear  from  Mr.  Bate,  the  chelae  are  sometimes  of  such  length 
and  size  that  they  cannot  possibly  be  used  for  carrying  food  to  the 
mouth.  In  the  males  of  certain  fresh-water  prawns  (Palsemon) 
the  right  leg  is  actually  longer  than  the  whole  body.^°  The 
gr3at  size  of  the  one  leg  with  its  chela3  may  aid  the  male  in 
fighting  with   his  rivals;   but  this  will  not  account  for  their 


*"  See  a  paper  by  ^Ir.  C.  Spence 
Bate,  with  figures,  in  '  Proc.  Zoolosj. 
See'  1868,  p.  363  ;  and  on  the 
nomenclature  of  the  genus,  ibid.  p. 


585.  I  am  greatly  indebted  to  Mr. 
Spence  Bate  for  nearly  all  the  above 
statements  with  respect  to  the  chelae 
of  the  higher  crustaceans. 


268  T]ie  Descent  of  Man.  Part  II. 


inequality  in  the  female  on  the  opposite  sides  of  of  the  body.  In 
Gelasimus,  according  to  a  statement  quoted  by  Milne-Edwards," 
the  male  and  the  female  live  in  the  same  bnrrow,  and  this 
shews  that  they  pair ;  the  male  closes  the  month  of  the  burrow 
with  one  of  its  chelre,  which  is  enormously  developed ;  so  that 
here  it  indirectly  serves  as  a  means  of  defence.  Their  main  use, 
however,  is  probably  to  seize  and  to  secure  the  female,  and  this 
in  some  instances,  as  with  Gammarus,  is  known  to  be  the  case. 
The  male  of  the  hermit  or  soldier  crab  {Fafiurus)  for  weeks 
together,  carries  about  the  shell  inhabited  by  the  female. ^^  The 
sexes,  however,  of  the  common  shore-crab  (Carcinus  manas),  as 
Mr.  Bate  informs  me,  unite  directly  after  the  female  has  moulted 
her  hard  shell,  when  she  is  so  soft  that  she  would  be  injured  if 
seized  by  the  strong  pincers  of  the  male ;  but  as  she  is  caught 
and  carried  about  by  the  male  before  moulting,  she  could  then  be 
seized  with  impunity. 

Fritz  Muller  states  that  certain  species  of  Melita  are  distin- 
guished from  all  other  amphipods  by  the  females  having  "  the 
"  coxal  lamellae  of  the  penultimate  pair  of  feet  produced  into 
"  hook-like  processes,  of  which  the  males  lay  hold  with  the 
*'  hands  of  the  first  pair."  The  develojDment  of  these  hook-like 
processes  has  probably  followed  from  those  females  which  were 
the  most  securely  held  during  the  act  of  reproduction,  having 
left  the  largest  number  of  offspring.  Another  Brazilian  amphi- 
pod  (Oichestia  Darwinii,  fig.  8)  presents  a  case  of  dimorphism, 
like  that  of  Tanais ;  for  there  are  two  male  forms,  which  differ 
in  the  structure  of  their  chelse.^^  As  either  chela  would  certainly 
suffice  to  hold  the  female, — for  both  are  now  used  for  this  purpose, 
— the  two  male  forms  probably  originated  by  some  having  varied 
in  one  manner  and  some  in  another ;  both  forms  having  derived 
certain  special,  but  nearly  equal  advantages,  from  their  differently 
shaped  organs. 

It  is  not  known  that  male  crustaceans  fight  together  for  the 
possession  of  the  females,  but  it  is  probably  the  case  ;  for  with 
most  animals  when  the  male  is  larger  than  the  female,  he  seems 
to  owe  his  greater  size  to  his  ancestors  having  fought 
with  other  males  during  many  generations.  In  most  of  the 
orders,  especially  in  the  highest  or  the  Brachyura,  the  male  is 
larger  than  the  female  ;  the  parasitic  genera,  however,  in  which 
the  sexes  follow  different  habits  of  life,  and  most  of  the  Ento- 
mostraca  must  be  excei)ted.     The  cheljB  of  many  crustaceans  are 

"  *Hist.  Nat.  des  Crust.'  torn.  ii.  of  S.  Devon.' 
1837,  p.  50.  ^^  Fritz  Miiller, '  Facts  and  Argu- 

>2  Mr.  C.  Spence  Bate,  *  Brit.  ments  for  Darwin,'  1869,  pp.  25-28 
Assoc,  Fourth  Report  on  the  Fauna 


Chap.  IX. 


Criistacemis. 


269 


■weapons  well  adapted  for  fighting.  Tluis  when  a  Devil-crab 
{Portunus  pubcr)  was  seen  by  a  son  of  Mr.  Bate  fighting  with  a 
Carcinus  mcenas,  the  latter  was  soon  thrown  on  its  back,  and  had 
every  limb  torn  from  its  body.  When  several  males  of  a  Brazilian 
Gelasimns,  a  species  furnished  with  immense  pincers,  were 
placed  together  in  a  glass  vessel  by  Fritz  Miiller,  they  mutilated 
and  killed  one  another.     Mr.  Bate  put  a  large  male  Carcmus 


Fig.  8.     Orchestia  Darwinii  (from  Fritz  Miiller),  showing  the  differently-constructed 
cheltB  uf  the  two  male  forms. 


mcenas  into  *a  pan  of  water,  inhabited  by  a  female  which  was 
paired  with  a  smaller  male ;  but  the  latter  was  soon  dispossessed. 


2/0  The  Descent  of  Man.  Part  II. 

Mr.  Bate  adds,  ''  if  they  fought,  the  victory  was  a  bloodless  one, 
"  for  I  saw  110  wounds."  This  same  naturalist  separated  a  male 
sand-skipper  (so  common  on  our  sea-shores),  Gammarus  marmus, 
from  its  female,  both  of  whom  were  imprisoned  in  the  same 
vessel  with  many  individuals  of  the  same  species.  The  female, 
when  thus  divorced,  soon  joined  the  others.  After  a  time  the 
male  Avas  put  again  into  the  same  vessel ;  and  he  then,  after 
swimming  about  for  a  time,  dashed  into  the  crowd,  and  without 
any  fighting  at  once  took  away  his  wife.  This  fact  shews  that 
in  the  Amphipoda,  an  order  low  in  the  scale,  the  males  and 
females  recognise  each  other,  and  are  mutually  attached. 

The  mental  powers  of  the  Crustacea  are  i^robably  higher  than 
at  first  sight  appears  probable.  Any  one  who  tries  to  catch  one 
of  the  shore-crabs,  so  common  on  tropical  coasts,  will  perceive 
how  wary  and  alert  they 'are.  There  is  a  large  crab  {Binjus 
latro),  found  on  coral  islands,  which  makes  a  thick  bed  of  the 
picked  fibres  of  the  cocoa-nut,  at  the  bottom  of  a  deep  burrow. 
It  feeds  on  the  fallen  fruit  of  this  tree  by  tearing  off  the  husk, 
fibre  by  fibre ;  and  it  always  begins  at  that  end  where  the  three 
eye-like  depressions  are  situated.  It  then  breaks  through  one  of 
these  eyes  by  hammering  with  its  heavy  front  pincers,  and 
turning  round,  extracts  the  albuminous  core  with  its  narrow 
posterior  pincers.  But  these  actions  are  probably  instinctive,  so 
that  they  would  be  performed  as  well  by  a  young  animal  as  by 
an  old  one.  The  following  case,  however,  can  hardly  be  so  con- 
sidered :  a  trustworthy  naturalist,  Mr.  Gardner,^^  whilst  watching 
a  shore-crab  (Gelasimus)  making  its  burrow,  threw  some  shells 
towards  the  hole.  One  rolled  in,  and  three  other  shells  remained 
within  a  few  inches  of  the  mouth.  In  about  five  minutes  the 
crab  brought  out  the  shell  which  had  fallen  in,  and  carried  it 
away  to  the  distance  of  a  foot ;  it  then  saw  the  three  other  shells 
lying  near,  and  evidently  thinking  that  they  might  likewise  roll 
in,  carried  them  to  the  spot  where  it  had  laid  the  first.  It 
would,  I  think,  be  difi&cnlt  to  distinguish  this  act  from  one 
performed  by  man  by  the  aid  of  reason. 

Mr.  Bate  does  not  know  of  any  well-marked  case  of  difference 
of  colour  in  the  two  sexes  of  our  British  crustaceans,  in  which 
respect  the  sexes  of  the  higher  animals  so  often  differ.  In  some 
cases,  however,  the  males  and  females  differ  slightly  in  tint,  but 
Mr.  Bate  thinks  not  more  than  may  be  accounted  for  by  their 
different  habits  of  life,  such  as  by  the  male  wandering  more 
about,  and  being  thus  more  exposed  to  the  light.    Dr.  Power 

**  '  Travels    in    the    Interior    of     463,  un  account  of  the  hiibits  of  the 
Brazil,'  1846,  p.  111.    I  have  given,      Birgus. 
in  my  '  Joui-nal  of  Researches,'  p. 


Chap.  IX.  Crustaceans.  2/1 

tried  to  distinguish  by  colour  the  sexes  of  the  several  species 
which  inhabit  the  Mauritius,  but  failed,  except  with  one  species 
of  Squilla,  probably  .S'.  stylifera,  the  male  of  which  is  described  as 
being"  of  a  beautiful  bluish-green,"  with  some  of  the  appendages 
cherry-red,  whilst  the  female  is  clouded  with  brown  and  grey, 
"  with  the  red  about  her  much  less  vivid  than  in  the  male."  '^ 
In  this  case,  we  may  suspect  the  agency  of  sexual  selection. 
From  M.  Bert's  observations  on  Daphnia,  when  placed  in  a  vessel 
illuminated  by  a  prism,  we  have  reason  to  believe  that  even  the 
lowest  crustaceans  can  distinguish  colours.  With  Saphirina  (an 
oceanic  genus  of  Entomostraca),  the  males  are  furnished  with 
minute  shields  or  cell-hke  bodies,  which  exhibit  beautiful 
changing  colours ;  these  are  absent  in  the  females,  and  in 
both  sexes  of  one  species.^*^  It  would,  however,  be  extremely 
rash  to  conclude  that  these  curious  organs  serve  to  attract  the 
females.  I  am  informed  by  Fritz  Miiller,  that  in  the  female  of  a 
Brazilian  species  of  Gelasimus,  the  whole  body  is  of  a  nearly 
uniform  greyish-brown.  In  the  male  the  posterior  part  of  the 
cephalo-thorax  is  pure  white,  with  the  anterior  part  of  a  rich 
green,  shading  into  dark  brown ;  and  it  is  remarkable  that  these 
colours  are  liable  to  change  in  the  course  of  a  few  minutes — the 
white  becoming  dirty  grey  or  even  black,  the  green  "  losing  much 
"  of  its  brilliancy."  It  deserves  especial  notice  that  the  males  do 
not  acquire  their  bright  colours  until  they  become  mature.  They 
appear  to  be  much  more  numerous  than  the  females;  they 
differ  also  in  the  larger  size  of  their  chelae.  In  some  species  of 
the  genus,  probably  in  all,  the  sexes  pair  and  inhabit  the  same 
burrow.  They  are  also,  as  we  have  seen,  highly  intelligent 
animals.  From  these  various  considerations  it  seems  probable 
that  the  male  in  this  species  has  become  gaily  ornamented  in 
order  to  attract  or  excite  the  female. 

It  has  just  been  stated  that  the  male  Gelasimus  does  not 
acquire  his  conspicuous  colours  until  mature  and  nearly  ready 
to  breed.  This  seems  a  general  rule  in  the  whole  class  in  respect 
to  the  many  remarkable  structural  differences  between  the  sexes. 
We  shall  hereafter  find  the  same  law  prevailing  throughout  the 
great  sub-kingdom  of  the  Vertebrata ;  and  in  all  cases  it  is 
eminently  distinctive  of  characters  which  have  been  acquired 
through  sexual  selection.  Fritz  Miiller  ^^  gives  some  striking 
instances  of  this  law ;  thus  the  male  sand-hopper  (Orchestia) 
does  not,  until  nearly  full  grown,  acquire  his  large  claspers, 

*^  Mr.  Ch.  Fraser,  in  *  Proc.  Zoo-  '*  Claus,  '  Die  freilebenden  Cope- 
log.  Soc'  1869,  p.  3.     I  am  indebted  poden,'  1863,  s.  35. 
to  Mr.  Bate   for  Dr.  Power's  state-  '^  '  Facts    and   Arguments,'   &c., 
ment.  p.  79. 


2/2  TJie  Descent  of  Man.  Part  II. 

which  are  yery  differently  constructed  from  those  of  the  female ; 
whilst  young,  his  claspers  resemble  those  of  the  female. 

Class,  Arachnida  (Spiders). — The  sexes  do  not  generally  differ 
much  in  colour,  but  the  males  are  often  darker  than  the  females, 
as  may  be  seen  in  Mr.  Blackwall's  magnificent  work.^**  In  some 
species,  however,  the  difference  is  conspicuous  :  thus  the  female 
of  Sjmrassus  smaragdidus  is  dullish  green,  whilst  the  adult  male 
has  the  abdomen  of  a  fine  yellow,  with  three  longitudinal  stripes 
of  rich  red.  In  certain  species  of  Thomisus  the  sexes  closely 
resemble  each  other,  in  others  they  differ  much ;  and  analogous 
cases  occur  in  many  other  genera.  It  is  often  difficult  to  say 
which  of  the  two  sexes  departs  most  from  the  ordinary  coloration 
of  the  genus  to  which  the  species  belong ;  but  Mr.  Blackwall 
thinks  that,  as  a  general  rule,  it  is  the  male ;  and  Canestrini  ^^ 
remarks  that  in  certain  genera  the  males  can  be  specifically  dis- 
tinguished with  ease,  but  the  females  with  great  difficulty.  I  am 
informed  by  Mr.  Blackwall  that  the  sexes  whilst  young  usually 
resemble  each  other ;  and  both  often  undergo  gi-eat  changes  in 
colour  during  their  successive  moults,  before  arriving  at  maturity. 
In  other  cases  the  male  alone  appears  to  change  colour.  Thus 
the  male  of  the  above  bright-coloured  Sparassus  at  first  re- 
sembles the  female,  and  acquires  his  peculiar  tints  only  when 
nearly  adult.  SjDiders  are  j)ossessed  of  acute  senses,  and  exliibit 
much  intelligence ;  as  is  well  known,  the  females  often  shew 
the  strongest  affection  for  their  eggs,  wliich  they  carry  about 
enveloped  in  a  silken  web.  The  males  search  eagerly  for  the 
females,  and  have  been  seen  by  Canestrini  and  others  to  fight  for 
possession  of  them.  This  same  author  says  that  the  union  of  the 
two  sexes  has  been  observed  in  about  twenty  species ;  and  he 
asserts  positively  that  the  female  rejects  some  of  the  males  who 
court  her,  threatens  them  with  open  mandibles,  and  at  last  after 
long  hesitation  accepts  the  chosen  one.  From  these  several 
considerations,  we  may  admit  with  some  confidence  that  the 
well-marked  differences  in  colour  between  the  sexes  of  certain 
species  are  the  results  of  sexual  selection ;  though  we  have  not 
here  the  best  kind  of  evidence, — the  display  by  the  male  of  his 
ornaments.  From  the  extreme  variability  of  colour  in  the  male 
of  some  sjDecies,  for  instance  of  Theridion  llneatum,  it  would 
appear  that  these  sexual  characters  of  the  males  have  not  as  yet 
become  well  fixed.     Canestrini  draws  the  same  conclusion  from 

'*  •  A.  History  of   the  Spiders  of  '  Caratteri  sessuali  secondarii  degli 

Great   Britain,'   1861-G4-.     For  the  Arachnidi,'  in  the  'Atti  della  See. 

fcllowiug  facts,  see  pp.  77,  88,  102.  Veneto-Trentiua  di  Sc.  Nat.  Padova, 

'^  This  author  has  recentfy  pub-  vol.  i.  Fasc.  3,  1873. 
lishod     a    valuable    essay    on    the 


Chap.  IX.  Spiders.  273 

the  fact  that  the  males  of  certain  species  present  two  forms, 
differing  from  each  other  in  the  size  and  length  of  their  jaws ;  and 
this  reminds  us  of  the  above  cases  of  dimorphic  crustaceans. 

The  male  is  generally  much  smaller  than  the  female,  sometimes 
to  an  extraordinary  degree,-"  and  he  is  forced  to  be  extremely 
cautious  in  making  his  advances,  as  the  female  often  carries  her 
coyness  to  a  dangerous  pitch.  De  Geer  saw  a  male  that  "  in  the 
"  midst  of  his  preparatory  caresses  was  seized  by  the  object  of 
"  his  attentions,  enveloped  by  her  in  a  web  and  then  devoured,  a 
"  sight  which,  as  he  adds,  filled  him  with  horror  and  indignation."  ^^ 
The  Eev.  0.  P.  Cambridge  ^^  accounts  in  the  following  manner 
for  the  extreme  smalhiess  of  the  male  in  the  genus  Nephila. 
"  M.  Vinson  gives  a  gTaphic  account  of  the  agile  way  in  which 
"  the  diminutive  male  escapes  from  the  ferocity  of  the  female,  by 
"  gliding  about  and  playing  hide  and  seek  over  her  body  and 
"  along  her  gigantic  limbs  :  in  such  a  pursuit  it  is  evident  that 
"  the  chances  of  escape  would  be  in  favour  of  the  smallest  males, 
"  while  the  larger  ones  would  fall  early  victims ;  thus  gradually 
"  a  diminutive  race  of  males  would  be  selected,  until  at  last  they 
"  would  dwindle  to  the  smallest  possible  size  compatible  with  the 
"  exercise  of  their  generative  functions, — in  fact  probably  to  the 
"  size  we  now  see  them,  i.e.,  so  small  as  to  be  a  sort  of  parasite 
"  upon  the  female,  and  either  beneath  her  notice,  or  too  agile  and 
"  too  small  for  her  to  catch  without  great  difficulty." 

Westring  has  made  the  interesting  discovery  that  the  males 
of  several  species  of  Theridion^  have  the  power  of  making 
a  stridulating  sound,  whilst  the  females  are  mute.  The  ap- 
paratus consists  of  a  serrated  ridge  at  the  base  of  the  abdomen, 
against  wliich  the  hard  hinder  part  of  the  thorax  is  rubbed ;  and 
of  this  structure  not  a  trace  can  be  detected  in  the  females.  It 
deserves  notice  that  several  writers,  including  the  well-known 
arachnologist  Walckenaer,  have  declared  that  spiders  are  attracted 
by  music.^*    From  the  analogy  of  the  Orthoptera  and  Homoptera, 

-"  Aug.  Vinson  (' Araneides    des  tion    to   Entomology,'  vol.  i.  1818, 

lies    de    la  Reunion/  pi.  vi.  figs.  1  p.  280. 

and  2)  gives  a  good  instance  of  the  ^^  'Proc.Zoolog.  Soc'  1871,  p.  621. 

small   size   of  the  male,  in  Epeira  ^s    Jheridion      (Asagena,     Sund.) 

nigra.     In    this    species,  as    I    may  serratipes,    A-punctatum    ct    gutta- 

add,  the  male  is  testaceous  and  the  turn ;     see    Westring,    in     Kroyer, 

female  black  with  legs  banded  with  '  Naturhist.  Tidslcrift,'  vol.  iv.  1842- 

red.      Other    even    more    striking  1843,   p.   349;    and   vol.   ii.    1846-- 

cases  of  inequality  in  size  between  1849,  p.  342.     See,  also,  for  other 

the      sexes      have     been     recorded  species,  'Aranege  Suecica?,' p.  184. 

('  Quarterly    Journal    of    Science,'  ^^  Dr.  H.  H.  van    Zouteveen,   in 

1868,  July,    p.    429) ;    but   I   have  his  Dutch  translation  of  this  work 

not  seen  the  original  accounts.  (vol.  i.  p.  444),  has  collected  several 

2^  Kirby  and  Speuce,  '  Introduc-  cases 

13 


2/4  The  Descent  of  Man.  Part  II. 

to  be  described  in  the  next  chapter,  we  may  feel  almost  sure  that 
the  stridulation  serves,  as  Westring  also  believes,  to  call  or  to 
excite  the  female ;  and  this  is  the  first  case  known  to  me  in  the 
ascending  scale  of  the  animal  kingdom  of  sounds  emitted  for 
this  purpose.^" 

Class,  Myriaj;oda. — In  neither  of  the  two  orders  in  this  class, 
the  millipedes  and  centipedes,  can  I  find  any  well-marked 
instances  of  such  sexual  differences  as  more  particularly  concern 
us.  In  Glomeris  li.mhata,  however,  and  perhaps  in  some  few 
other  species,  the  males  differ  slightly  in  colour  from  the  females ; 
but  this  Glomeris  is  a  highly  variable  species.  In  the  males  of 
the  Diplopoda,  the  legs  belonging  either  to  one  of  the  anterior  or 
of  the  posterior  segments  of  the  body  are  modified  into  pre- 
hensile hooks  which  serve  to  secure  the  female.  In  some  species 
of  lulus  the  tarsi  of  the  male  are  furnished  with  membranous 
suckers  for  the  same  purpose.  As  we  shall  see  when  we  treat 
of  Insects,  it  is  a  much  more  unusual  circumstance,  that  it  is 
the  female  in  Lithobius,  which  is  furnished  with  prehensile 
appendages  at  the  extremity  of  her  body  for  holding  the  male.^^ 


CHAPTER  X. 
Secondary  Sexual  Characters  of  Insects. 

Diversified  structures  possessed  by  the  males  for  seizing  the  females — 
Differences  between  the  sexes,  of  which  the  meaning  is  not  understood — 
Difference  in  size  between  the  sexes — Thysanura — Diptera — Hemiptera 
— Homoptera,  musical  powers  possessed  by  the  males  alone — Orthoptera, 
musical  instruments  of  the  males,  much  diversified  in  structure  ; 
pugnacity  ;  colours — Neuroptera,  sexual  differences  in  colour — Hyme- 
noptera,  pugnacity  and  colours — Coleoptera,  colours ;  furnished  with 
great  horns,  apparently  as  an  ornament ;  battles ;  stridulating  organs 
generally  common  to  both  sexes. 

In  the  immense  class  of  insects  the  sexes  sometimes  differ  in 
their  locomotive-organs,  and  often  in  their  sense-organs,  as  in 
the  pectinated  and  beautifully  plumose  antennae  of  the  males  of 
many  species.  In  Chloeon,  one  of  the  EjDhemerse,  the  male  has 
great  pillared  eyes,  of  which  the  female  is  entirely  destitute.* 
The  ocelli  are  absent  in  the  females  of  certain  insects,  as  in  the 

2^  Hilgendorf,  however,  has  lately  '  Hist.  Nat.  des  Insectes  :   Apteres,' 

called    attention    to    an    analogous  torn.  iv.  1847,  pp.  17,  19,  (58. 
structure   in    some    of    the    higher  *  Sir     J.     Lubbock,     '  Transact, 

crustaceans,   which    seems    adapted  Linnean    Soc'    vol.    xxv.    1866,    p. 

to  produce  sound ;   see   '  Zoological  484.     With    respect    to    the    Mu- 

Record,'  1869,  p.  603.  tillidae     see     West  wood,     '  Modern 

^^  Walckenaer     et     P.    Gervais,  Class,  of  Insects,' vol.  ii.  p.  213. 


Chap.  X.  Insects.  275 

Mutillidie ;  and  here  the  females  are  hkewise  wingless.  But 
we  are  chiefly  concerned  with  structures  by  which  one  male  is 
enabled  to  conquer  another,  either  in  battle  or  courtship,  through 
his  strength,  pugnacity,  ornaments,  or  music.  The  innumerable 
contrivances,  therefore,  by  which  the  male  is  able  to  seize  the 
female,  may  be  briefly  passed  over.  Besides  the  complex  structures 
at  the  apex  of  the  abdomen,  which  ought  perhaps  to  be  ranked 
as  primary  organs,^  "  it  is  astonishing,"  as  Mr.  B.  D.  Walsh  ^  has 
remarked,  "  how  many  different  organs  are  worked  in  by  nature 
"  for  the  seemingly  insignificant  object  of  enabling  the  male  to 
"  grasp  the  female  firmly."  The  mandibles  or  jaws  are  some- 
times used  for  this  purpose;  thus  the  male  Corydalis  cornutus  (a 
neuropierous  insect  in  some  degree  allied  to  the  Dragon-flies,  &c.) 
lias  immense  curved  jaws,  many  times  longer  than  those  of  the 
female ;  and  they  are  smooth  instead  of  being  toothed,  so  that 
he  is  thus  enabled  to  seize  her  without  injury.'^  One  of  the 
stag-beetles  of  North  America  {Lucanus  ela/Juis)  uses  his  jaws, 
which  are  much  larger  than  those  of  the  female,  for  the  same 
purpose,  but  probably  likewise  for  fighting.  In  one  of  the 
sand-wasps  {Ammophild)  the  jaws  in  the  two  sexes  are  closely 
alike,  but  are  used  for  widely  different  purposes  :  the  males,  as 
Professor  Westwood  observes,  "  are  exceedingly  ardent,  seizing 
"  their  partners  round  the  neck  with  their  sickle-shaped  jaws;"^ 
whilst  the  females  use  these  organs  for  burrowing  in  sand-banks 
and  making  their  nests. 

The  tarsi  of  the  front-legs  are  dilated  in  many  male  beetles,  or 
are  furnished  with  broad  cushions  of  hairs  ;  and  in  many  genera 
of  water-beetles  they  are  armed  with  a  round  flat  sucker,  so  that 
the  male  may  adhere  to  the  slippery  body  of  the  female.     It  is  a 

"^  These  organs  in  the  male  often  species    having    been    observed    in 

differ    in   closely-allied    species,  and  union.      Mr.     MacLachlan    informs 

afford  excellent  specific  characters.  me    (vide    '  Stett.    Eut.    Zeitung,' 

But  their  importance,  from  a  func-  1867,    s,    155)   that    when    several 

tional    point    of    view,    as    Mr.    R.  species   of  Phryganidje,  which   ])re- 

MacLachlan    has    remarked    to  me,  sent  strongly-pronounced  differences 

has    probably    been    overrated.      It  of  this  kind,  were  confined  together 

has  been  suggested,  that  slight  dif-  by   Dr.   Aug.   Meyer,    theji   coupled, 

ferences     in     these    organs    would  and  one  pair  produced  fertile  ova. 

suffice  to  prevent  the  intercrossing  ^  t  -pj^g    Practical    Entomologist,' 

_  of  well-marked  varieties  or  incipient  Philadelphia,    vol.    ii.    May,    1807, 

species,  and  would  thus  aid  in  their  p.  88. 

development.  That  this  can  hardly  ''  Mr.  Walsh,  ibid.  p.  107. 
be  the  case,  we  may  infer  from  the  ^  '  Modern  classification  of  In- 
many  recorded  cases  (see,  for  in-  sects,'  vol.  ii.  1840,  pp.  205,  206. 
instance,  Bronn,  '  Geschichte  der  Mr.  Walsh,  who  called  my  attention 
Natur,'  B.  ii.  18-13,  s.  164;  and  to  the  double  use  of  the  jaws,  says 
Westwood,  '  Transact.  Ent.  Soc'  that  he  has  repeatedly  observed 
vol.   iii.   1842,  p.    195)  of  distinct  this  fact. 


2/6 


The  Descent  of  Man. 


Part  II. 


much  more  unusual  circumstance  that  the  female  of  some  water- 
beetles  (Dytiscus)  have  their  elytra  deeply  grooved,  and  in 
Acilius  sulcatus  thickly  set  with  hairs,  as  an  aid  to  the  male. 

The  females  of  some  other  water- 
beetles  (Hydroporus)  have  their 
elytra  punctured  for  the  same 
purpose."  In  the  male  of  Crahro 
cribrurius  (fig.  9),  it  is  the  tibia 
which  is  dilated  into  a  broad 
horny  plate,  with  minute  mem- 
braneous dots,  giving  to  it  a  sin- 
gular appearance  like  that  of  a 
riddle.''  In  the  male  of  Penthe 
(a  genus  of  beetles)  a  few  of  the 
middle  joints  of  the  antennae  are 
dilated  and  furnished  on  the  in- 
ferior surface  with  cushions  of  hair, 
exactly  like  those  on  the  tarsi  of 
the  Carabidse,  "  and  obviously  for 
"  the  same  end."  In  male  dragon- 
flies,  "  the  appendages  at  the  tip 
"  of  the  tail  are  modified  in  an 
"  almost  infinite  variety  of  curious 
"  patterns  to  enable  them  to  em- 
^''-  'm^ai:!TowS^Sgure,fYEL"'"^     "  brace  the  neck  of  the  female." 

Lastly,  in  the  males  of  many  in- 
sects, the  legs  are  furnished  with  peculiar  spines,  knobs  or 
spurs ;  or  the  whole  leg  is  bowed  or  thickened,  but  this  is  by  no 
means  invariably  a  sexual  character ;  or  one  pair,  or  all  three 
IDairs  are  elongated,  sometimes  to  an  extravagant  length.^ 

The  sexes  of  many  species  in  all  the  orders  present  differences, 
of  which  the  meaning  is  not  understood.  One  curious  case  is 
that  of  a  beetle  (fig.  10),  the  male  of  which  has  the  left  mandible 
much  enlarged;  so  that  the  mouth  is  greatly  distorted.  In 
another  Carabidous  beetle,   Eurygnathus,^  we  have  the  case. 


^  We  have  here  a  curious  and 
inexplicable  case  of  dimorphism,  for 
some  of  the  females  of  four  Euro- 
pean species  of  Dytiscus,  and  of 
cei'tain  species  of  Hydroporus,  have 
their  elytra  smooth ;  and  no  inter- 
mediate gradations  between  the 
sulcated  or  punctured,  and  the  quite 
smooth  elytra  have  been  observed. 
See  Dr.  11.  Schaum,  as  quoted  in 
the  'Zoologist,'  vol.  A-.-vi.  1847-48, 
p.  1896.     Also   Kirby  and   Spence, 


'  Introduction  to  Entomology,'  vol. 
iii.  1826,  p.  305. 

^  Westwood,  '  Modern  Class.'  vol. 
ii.  p.  193.  The  following  state- 
ment about  Penthe,  and  others  in 
inverted  commas,  are  taken  from 
Mr.  Walsh,  '  Practical  Entomolo- 
gist,' Philadelphia,  vol.  ii.  p.  88. 

^  Kirby  and  Spence,  'Introduct.' 
&c.,  vol.  iii.  pp.  332-336. 

^  '  Insecta  Madcrensia,'  1854,  p. 
20. 


Chap.  X. 


Insects. 


277 


unique  as  ftir  as  known  to  Mr.  Wollaston,  of  the  head  of  the 
female  being  much  broader  and  larger,  though  in  a  variable 
degree,  than  that  of  the  male.  Any  number 
of  such  cases  could  be  given.  They  abound 
in  the  Lepidoptera:  one  of  the  most  extra- 
ordinary is  that  certain  male  butterflies 
have  their  fore-legs  more  or  less  atrojiliied, 
with  the  tibise  and  tarsi  reduced  to  mere  ru- 
dimentary knobs.  The  wings,  also,  in  the  two 
sexes  often  differ  in  neuration,^*'  and  some- 
times considerably  in  outline,  as  in  the  Ari- 
coris  epitiis,  wiiicli  was  shewn  to  me  in  the 
British  Museum  by  Mr.  A.  Butler.  The  males 
of  certain  South  American  butterflies  have 
tufts  of  hair  on  the  margins  of  the  wings, 
and  horny  excrescences  on  the  discs  of  the 
posterior  pair.^^  In  several  British  butter- 
flies, as  shewn  by  Mr.  Wonfor,  the  males  alone 
are  in  parts  clothed  with  peculiar  scales. 

The  use  of  the  bright  light 'of  the  female 
glow-worm  has  been  subject  to  much  discus- 
sion. The  male  is  feebly  luminous,  as  are  the 
larva3  and  even  the  eggs.  It  has  been  sup- 
posed by  some  authors  that  the  light  serves  to 
frighten  away  enemies,-  and  by  others  to 
guide  the  male  to  the  female.  At  last,  Mr. 
Belt  ^2  appears  to  have  solved  the  difficulty : 
he  finds  that  all  the  Lampyridse  which  he  has 
tried  are  highly  distasteful  to  insectivorous 
mammals  and  birds.  Hence  it  is  in  accordance 
with  Mr.  Bates'"  view,  hereafter  to  be  explained, 
that  many  insects  mimic  the  Lampyridae 
closely,  in  order  to  be  mistaken  for  them,  and 
thus  to  escape  destruction.  He  further  be- 
lieves that  the  luminous  species  profit  by 
being  at  once  recognised  as  unpalatable. 
It  is  i^robable  that  the  same  explanation  may  be  extended  to  the 

'^^  E.  Doubleday,  '  Annals  and  74.  Mr.  Wonfor's  observations  are 
Mag.  of  Xat.  Hist.'  vol.  i.  1848,  p.  quoted  in  '  Popular  Science  Review,* 
379.     I  may  add  that  the  wings  in      1868,  p.  343. 

^-  'The  Naturalist  in  Nicaragua,' 
1874,  pp.  316-320.  On  the  phos- 
phorescence of  the  eggs,  see  'Annals 
and  Mag.  of  Nat.  Hist.'  1871,'  Nov., 
p.  372. 


Fig.  10.  Taphroderes 
distortus  (much  en- 
larged). Upper  fig- 
ure, male  ;  lower 
figure,  female. 


certain  Hymenoptera  (see  Shuclvard, 
'  Fossorial  Hynienop.'  1837,  pp.  39- 
43)  differ  in  neuration  according  to 
sex. 

"  H.   W.  Bates,   in   'Journal    of 
Proc.  Linn.  Soc'  vol.  vi.  1862,  p. 


2/8  The  Descent  of  Man,  Part  11. 

Elaters,  both  sexes  of  which  are  highly  luminous.  It  is  not 
known  why  the  wings  of  the  female  glow-worm  have  not  been 
developed ;  but  in  her  present  state  she  closely  resembles  a 
larva,  and  as  larvae  are  so  largely  preyed  on  by  many  animals, 
we  can  understand  why  she  has  been  rendered  so  much  more 
luminous  and  conspicuous  than  the  male ;  and  why  the  larvae 
themselves  are  likewise  luminous. 

Difference  in  Size  hetween  the  Sexes. — "With  insects  of  all  kinds 
the  males  are  commonly  smaller  than  the  females;  and  this 
difference  can  often  be  detected  even  in  the  larval  state.  So 
considerable  is  the  difference  between  the  male  and  female 
cocoons  of  the  silk-moth  (Bomhyx  mori),  that  in  France  they  are 
separated  by  a  particular  mode  of  weighing.^^  In  the  lower 
classes  of  the  animal  kingdom,  the  greater  size  of  the  females 
seems  generally  to  depend  on  their  developing  an  enormous 
number  of  ova  ;  and  this  may  to  a  certain  extent  hold  good  with 
insects.  But  Dr.  Wallace  has  suggested  a  much  more  probable 
explanation.  He  finds,  after  carefully  attending  to  the  develop- 
ment of  the  caterpillars  of  Bombyx  cynthia  and  yamamai,  and 
especially  to  that  of  some  dwarfed  caterjDillars  reared  from  a 
second  brood  on  unnatural  food,  "  that  in  proportion  as  the  in- 
"  dividual  moth  is  finer,  so  is  the  time  required  for  its  metamor- 
"  pilosis  longer ;  and  for  this  reason  the  female,  which  is  the 
"  larger  and  heavier  insect,  from  having  to  carry  her  numerous 
"  eggs,  will  be  preceded  by  the  male,  which  is  smaller  and  has 
"  less  to  mature,"  ^^  Now  as  most  insects  are  short-lived,  and  as 
they  are  exposed  to  many  dangers,  it  would  manifestly  be  ad- 
vantageous to  the  female  to  be  impregnated  as  soon  as  possible. 
This  end  would  be  gained  by  the  males  being  first  matured  in 
large  numbers  ready  for  the  advent  of  the  females ;  and  this 
again  would  naturally  follow,  as  Mr  A.  E.  "Wallace  has  re- 
marked,^^  through  natural  selection;  for  the  smaller  males 
would  be  first  matured,  and  thus  would  procreate  a  large 
number  of  offspring  which  would  inherit  the  reduced  size  of 
then*  male  parents,  whilst  the  larger  males  from  being  matured 
later  would  leave  fewer  offspring. 

There  are,  however,  exceptions  to  the  rule  of  male  insects 
being  smaller  than  the  females  :  and  some  of  these  exceptions  are 
intelligible.  Size  and  strength  would  be  an  advantage  to  the 
males,  which  fight  for  the  jDossession  of  the  females ;  and  in 
those  cases,  as  with  the  stag-beetle  (Lucanus),  the  males  are 
larger  than  the  females.      There   are,  however,  other    beetles 

"  Robinet,  'Vers   a  Soie,'  1848,      vol.  y.  p.  486. 
p.  207.  '5  i  Journal    of  Proc.    Ent.    Soc. 

^^  'Transact.  Ent.  Soc'  3rd  series,      Feb.  4th,  1867,  p.  Ixxi. 


Chap.  X.  TJiysanura.  279 

which  are  not  known  to  fight  together,  of  which  the  males 
exceed  the  females  in  size ;  and  the  meaning  of  this  fact  is  not 
known ;  but  in  some  of  these  cases,  as  with  the  huge  Dynastes 
and  Megasoma,  we  can  at  least  see  that  there  would  be  no 
necessity  for  the  males  to  be  smaller  than  the  females,  in  order 
to  be  matured  before  them,  for  these  beetles  are  not  short-lived, 
and  there  would  be  ample  time  for  the  pairing  of  the  sexes.  So 
again,  male  dragon-flies  (Libellulidse)  are  sometimes  sensibly 
larger,  and  never  smaller,  than  the  females;^*'  and  as  Mr. 
MacLachlan  believes,  they  do  not  generally  pair  with  the  females 
until  a  week  or  fortnight  has  elapsed,  and  until  they  have 
assumed  their  proper  masculine  colours.  But  the  most  curious 
case,  shewing  on  what  comj^lex  and  easily-overlooked  relatioDS, 
so  trifling  a  character  as  difference  in  size  between  the  sexes 
may  depend,  is  that  of  the  aculeate  Hymenoptera ;  for  Mr.  F. 
Smith  informs  me  that  throughout  nearly  the  whole  of  this 
large  group,  the  males,  in  accordance  with  the  general  rule,  are 
smaller  than  the  females,  and  emerge  about  a  week  before  them  ; 
but  amongst  the  Bees,  the  males  oi  Aipis  melUfica,  Anthidium 
manicatum,  and  Anthopliora  ac&rvorum,  and  amongst  the  Fossores, 
the  males  of  the  Methoca  ichneumonides,  are  larger  than  the 
females.  The  explanation  of  this  anomaly  is  that  a  marriage 
flight  is  absolutely  necessary  with  these  species,  and  the  male 
requires  great  strength  and  size  in  order  to  carry  the  female 
through  the  air.  Increased  size  has  here  been  acquired  in  op- 
position to  the  usual  relation  between  size  and  the  period  of 
development,  for  the  males,  though  larger,  emerge  before  the 
smaller  females. 

We  will  now  review  the  several  Orders,  selecting  such  facts 
as  more  particularly  concern  us.  The  Lepidoptera  (Butterflies 
and  Moths)  will  be  retained  for  a  separate  chapter. 

Order,  Thysanura. — The  members  of  this  lowly  organized 
order  are  wingless,  dull-coloured,  minute  insects,  with  ugly, 
almost  misshapen  heads  and  bodies.  Their  sexes  do  not  difi"er ; 
but  they  are  interesting  as  shewing  us  that  the  males  pay 
sedulous  court  to  the  females  even  low  down  in  the  animal 
scale.  Sir  J.  Lubbock  ^^  says  :  "  it  is  very  amusing  to  see  these 
"  little  creatures  (SmyntJmrus  luteus)  coquetting  together.  The 
"  male,  which  is  much  smaller  than  the  female,  runs  round  her, 
"  and  they  butt  one  another,  standing  face  to  face,  and  moving 

'®  For  this  and  other  statements  see  p.  344, 
on  the  size  of  the  sexes,  see  Kirby  ^'  '  Transact.   Linnean    Soc'  vol. 

and  Spence,  ibid.   vol.   iii.  p.  300  ;  xxvi,  1868,  p.  296. 
on  the  dui'ation  of  life  in  insects, 


28o  The  Descent  of  Man.  Part  II. 

"  backward  and  forward  like  tw'o  playful  lambs.  Then  the 
"  female  pretends  to  run  away  and  the  male  runs  after  her  with 
"  a  queer  appearance  of  anger,  gets  in  front  and  stands  facing 
"  her  again ;  then  she  turns  coyly  round,  but  he,  quicker  and 
"  more  active,  scuttles  round  too,  and  seems  to  whip  her 
"  with  his  antennae ;  then  for  a  bit  they  stand  face  to  face, 
"  play  with  their  antennae,  and  seem  to  be  all  in  all  to  one 
"  another." 

Order,  Dlptera  (Flies). — The  sexes  differ  little  in  colour.  The 
greatest  difference,  known  to  Mr.  F.  Walker,  is  in  the  genus 
Bibio,  in  which  the  males  are  blackish  or  quite  black,  and  the 
females  obscure  brownish-orange.  The  genus  Elaphomyia,  dis- 
covered by  Mr.  Wallace  ^^  in  New  Guinea,  is  highly  remarkable, 
as  the  males  are  furnished  with  horns,  of  which  the  females  are 
quite  destitute.  The  horns  spring  from  beneath  the  eyes,  and 
curiously  resemble  those  of  a  stag,  being  either  branched  or  pal- 
mated.  In  one  of  the  species,  they  equal  the  whole  body  in 
length.  They  might  be  thought  to  be  adapted  for  fighting,  but 
as  in  one  species  they  are  of  a  beautiful  pink  colour,  edged  with 
black,  with  a  pale  central  stripe,  and  as  these  insects  have 
altogether  a  very  elegant  appearance,  it  is  perhaps  more  probable 
that  they  serve  as  ornaments.  That  the  males  of  some  Diptera 
fight  together  is  certain ;  for  Prof.  Westw^ood  ^^  has  several  times 
seen  this  with  the  Tipulse.  The  males  of  other  Diptera  ap- 
parently try  to  win  the  females  by  their  music  :  H.  Miiller  ^ 
watched  for  some  time  two  males  of  an  Eristalis  courting  a 
female;  they  hovered  above  her,  and  flew  from  side  to  side, 
making  a  high  humming  noise  at  the  same  time.  Gnats  and 
mosquitoes  (Culicidse;  also  seem  to  attract  each  other  by  hum- 
ming ;  and  Prof.  Mayer  has  recently  ascertained  that  the  hairs 
on  the  antennae  of  the  male  vibrate  in  unison  with  the  notes  of  a 
tuning-fork,  within  the  range  of  the  sounds  emitted  by  the  female. 
The  longer  hairs  vibrate  sympathetically  with  the  graver  notes, 
and  the  shorter  hairs  with  the  higher  ones.  Landois  also  asserts 
that  he  has  repeatedly  drawn  down  a  whole  swarm  of  gnats  by 
uttering  a  particular  note.  It  may  be  added  that  the  mental 
faculties  of  the  Diptera  are  probably  higher  than  in  most  other  in- 
sects, in  accordance  with  their  highly  developed  nervous  system.^^ 

^*  *  The  Malay  Archipelago,'  vol.  ^i  ggg  ^ly^  -^  f.  Lowne's  interest- 

ii.  1869,  p.  313.  ing  work,  '  On  the  Anatomy  of  the 

'^  'Modern  Classification    of   In-  Blow-fly,  Musca  vomitoria,' 1870,  p. 

sects,'  A'ol.  ii.  1840,  p.  526.  14.     He  remarks  (p.  33)  that,  "  the 

2"  Anwendung,  &c.,  '  Verh.  d.  n.  "  captured    flies    utter    a    peculiar 

V.   Jahrg.'  xxix.  p.  80.     Mayer,  in  "  plaintive  note,  and  that  this  sound 

'American  Naturalist,'  1874, p.  236.  "  causes  other  flies  to  disappear." 


Chap.  X.  Hemiptera  and  Homoptera,  281 

Order,  Hemiptera  (Field-Bugs).— Mr.  J.  W.  Douglas,  who  has 
particularly  attended  to  the  British  species,  has  kindly  given  me 
an  account  of  their  sexual  differences.  The  males  of  some  species 
are  furnished  with  wings,  whilst  the  females  are  wingless ;  the 
sexes  differ  in  the  form  of  their  bodies,  elytra,  antenn£e  and  tarsi ; 
but  as  the  signification  of  these  differences  are  unknown,  they 
may  be  here  passed  over.  The  females  are  generally  larger  and 
more  robust  than  the  males.  With  British,  and,  as  far  as 
Mr.  Douglas  knows,  with  exotic  species,  the  sexes  do  not 
commonly  differ  much  in  colour;  but  in  about  six  British 
species  the  male  is  considerably  darker  than  the  female,  and 
in  about  four  other  species  the  female  is  darker  than  the  male. 
Both  sexes  of  some  species  are  beautifully  coloured;  and  as 
these  insects  emit  an  extremely  nauseous  odour,  their  con- 
spicuous colours  may  serve  as  a  signal  that  they  are  unpalat- 
able to  insectivorous  animals.  In  some  few  cases  their  colours 
appear  to  be  directly  protective :  thus  Prof.  Hoffmann  informs 
me  that  he  could  hardly  distinguish  a  small  pink  and  green 
species  from  the  buds  on  the  trunks  of  lime-trees,  which  this 
insect  frequents. 

Some  species  of  Eeduvidae  make  a  stridulating  noise;  and,  in 
the  case  of  Pirates  stridulus,  this  is  said  ^^  to  be  effected  by  the 
movement  of  the  neck  within  the  pro-thoracic  cavity.  Accord- 
ing to  Westring,  Beduvius  jiersonatus  also  stridulates.  But  I 
have  no  reason  to  suppose  that  this  is  a  sexual  character,  ex- 
cepting that  with  non-social  insects  there  seems  to  be  no  use 
for  sound-producing  organs,  unless  it  be  as  a  sexual  call. 

Order,  Homoptera. — Every  one  who  has  wandered  in  a  tropi- 
cal forest  must  have  been  astonished  at  the  din  made  by  the 
male  Cicadae.  The  females  are  mute;  as  the  Grecian  poet 
Xenarchus  says,  "  Happy  the  Cicadas  live,  since  they  all  have 
*'  voiceless  waives."  The  noise  thus  made  could  be  plainly  heard 
on  board  the  "  Beagle,"  when  anchored  at  a  quarter  of  a  mile 
from  the  shore  of  Brazil  ;•  and  Captain  Hancock  says  it  can  be 
heard  at  the  distance  of  a  mile.  The  Greeks  formerly  kept,  and 
the  Chinese  now  keep  these  insects  in  cages  for  the  sake  of 
their  song,  so  that  it  must  be  pleasing  to  the  ears  of  some  men.^^ 
The  Cicadidse  usually  sing  during  the  day,  whilst  the  FulgoridsB 
appear  to  be  night-songsters.  The  sound,  according  to  Landois,^* 

"  Westwood,   'Modern  Class,   of  also,  on  the  Fulgorida?,  Kirby  and 

Insects,'  vol.  ii.  p.  473.  Spence,  '  Introduct.'  vol.  ii.  p.  401. 

"^  These    particulars    are    taken  ^4  <  Zeitschrift    fiir   wissenschaft. 

from  Westwood's  '  Modern  Class,  of  Zoolog.'  B.  xvii.  1867,  s.  152-158. 
Insects,'  vol.  ii.  1840,  p.  422.     See, 


282  The  Descent  of  Man.  Part  II. 

is  produced  by  the  vibration  ot  the  lips  of  the  spiracles,  which 
are  set  into  motion  by  a  current  of  air  emitted  from  the  trachese ; 
but  tills  view  has  lately  been  disputed.  Dr.  Powell  appears  to 
have  proved 2^  that  it  is  produced  by  the  vibration  of  a  mem- 
brane, set  into  action  by  a  speci-al  muscle.  In  the  living  insect, 
whilst  stridulating,  this  membrane  can  be  seen  to  vibrate ;  and 
in  the  dead  insect  the  proper  sound  is  heard,  if  the  muscle, 
when  a  little  dried  and  hardened,  is  pulled  with  the  point  of  a 
pin.  In  the  fem.ale  the  whole  complex  musical  apparatus  is 
present,  but  is  much  less  developed  than  in  the  male,  and  is 
never  used  for  producing  sound. 

With  respect  to  the  object  of  the  music,  Dr.  Hartman,  in 
speaking  of  the  Cicada  septemdecim  of  the  United  States,  says,  ^s 
"  the  drums  are  now  (June  6th  and  7th,  1851)  heard  in  all 
"  directions.  This  I  believe  to  be  the  marital  summons  from 
"  the  males.  Standing  in  thick  chestnut  sprouts  about  as  high 
"  as  my  head,  where  hundreds  were  around  me,  I  observed  the 
"  females  coming  around  the  drumming  males,"  Ho  adds,  "  this 
"  season  (Aug.  1868)  a  dwarf  pear-tree  in  my  garden  produced 
"  about  fifty  larvsB  of  Cic.  prm7iosa ;  and  I  several  times  noticed 
"  the  females  to  alight  near  a  male  while  he  was  uttering  his 
"  clanging  notes."  Fritz  Miiller  writes  to  me  from  S.  Brazil 
that  he  has  often  listened  to  a  musical  contest  between  two  or 
three  males  of  a  species  with  a  particularly  loud  voice,  seated 
at  a  considerable  distance  from  each  other :  as  soon  as  one  had 
finished  his  song,  another  immediately  began,  and  then  another. 
As  there  is  so  much  rivalry  between  the  males,  it  is  probable 
that  the  females  not  only  find  them  by  their  sounds,  but  that, 
like  female  birds,  they  are  excited  or  allured  by  the  male  with 
the  most  attractive  voice. 

I  have  not  heard  of  any  well-marked  cases  of  ornamental 
differences  between  the  sexes  of  the  Homoptera.  Mr.  Douglas 
informs  me  that  there  are  three  British  species,  in  which  the 
male  is  black  or  marked  with  black  bands,  whilst  the  females  are 
pale-coloured  or  obscure. 

Order,  Orthoptera  (Crickets  and  Grasshoppers).— The  males  in 
the  three  saltatorial  families  in  this  Order  are  remarkable  for 
their  musical  powers,  namely  the  Achetidse  or  crickets,  the 
Locustidse  for  which  there  is  no  equivalent  English  name,  and  the 
Acridiidae  or  grasshoppers.     The  stridulation  produced  by  some 

"  'Transact.    New   Zealand    In-  from   a  'Journal  of  the  Doings  of 

stitute,'  vol.  V.  1873,  p.  286.  Cicada  septemdecim  '  by  Dr.  Hart- 

2^  I  am   indebted  to  Mr.  Walsh  man. 
for   havmg   sent    me    this   extract 


Chap.  X. 


Orthoptera. 


283 


of  the  Locustidse  is  so  loud  that  it  can  be  heard  during  tlie  night 
at  the  distance  of  a  mile ;  -"  and  that  made  by  certain  species  is 
not  unmusical  even  to  the  human  ear,  so  that  the  Indians  on  the 
Amazons  keep  them  in  wicker  cages.  All  observers  agree  tliat 
the  sounds  serve  either  to  call  or  excite  the  mute  females.  With 
respect  to  the  migratory  locusts  of  Russia,  Korte  has  given  ^^  an 
interesting  case  of  selection  by  the  female  of  a  male.  The  males 
of  this  species  {Pachytylus  miyratorius)  whilst  coupled  with  the 
female  stridulate  from  anger  or  jealousy,  if  approached  by  other 
males.  The  house-cricket  when  surprised  at  night  uses  its  voice 
to  warn  its  fellows.-^  In  North  America  the  Katy-did  {Phity- 
2>hyllum  coneavum,onQoii\\Q  Locustidse) is  described^"  as  mount- 
ing on  the  upper  branches  of  a  tree,  and  in  the  evening  beginning 
"  his  noisy  babble,  while  rival  notes  issue  from  the  neighbouring 
"  trees,  and  the  groves  resound  with  the  call  of  Kafydid-she-did 
"  the  hve-long  night."  Mr. 
Bates,  in  speaking  of  the  Euro- 
pean field-cricket  (one  of  the 
Achetidse),  says, "  the  male  has 
"  been  observed  to  jDlace  him- 
"  self  in  the  evening  at  the 
"  entrance  of  his  burrow,  and 
"  stridulate  until  a  female  ap- 
"  preaches,  when  the  louder 
"  notes  are  succeeded  by  a 
"  more  subdued  tone,  whilst 
"  the  successful  musician  ca- 
"  resses  with  his  antennae  the 
"  mate  he  has  won."^^  Dr. 
Scudder  was  able  to  excite  one 
of  these  insects  to  answer  him, 
by  rubbing  on  a  file  with  a 
quill.^  In  both  sexes  a  re- 
markable auditory  apparatus 
has  been  discovered  by  Von  Siebold,  situated  in  the  front  legs 


Fig.  11,   Gryllus  carapestris  (from  Landois). 
Right-hand  figure,  under  side  of  part  of  a 

wing-nervure,  much  magnified,  showing 

the  teeth,  st. 
Left-hand  figure,   upper  surface  of  wing- 

cover,  with  the  projecting,  smooth  nervure, 

r,  across  which  the  teeth  {st)  are  scraped. 


2^  L.  Guilding,  'Transact  Linn. 
See'  vol.  XV.  p.  154. 

28  I  state  this  on  the  authority 
of  Koppen, '  Ueber  die  Heuschrecken 
in  Siidrussland,'  1866,  p.  32,  for  I 
have  in  vain  endeavoured  to  procure 
Korte's  work. 

29  Gilbert  White,  'Nat.  Hist,  of 
Selborne,'  vol.  ii.  1825,  p.  262. 

^"  Harris,  '  Insects  of  New  Eng- 
land/ 1842,  p.  128. 

31  'The  Naturalist  on  the  Ama- 


zons,' vol.  i.  1863,  p.  252.  Mr. 
Bates  gives  a  very  interesting  dis- 
cussion on  the  gfadations  in  the 
musical  apparatus  of  the  three 
families.  See  also  Westwood, 
*  Modern  Class.'  vol.  ii.  pp.  445 
and  453. 

32  'Proc.  Boston  Soc.  of  Nat. 
Hist.'  vol.  xi.  April,  1868. 

33  '  Nouveau  Manuel  d'Anat. 
Comp.'  (French  translat.),  torn.  i. 
1850,  p.  567. 


284  The  Descent  of  Man.  Part  II. 

In  the  three  ramilies  the  sounds  are  differently  produced.  In 
the  males  of  the  Achetidse  both  wing-covers  have  the  same 
apparatus ;  and  this  in  the  field-cricket  {Gryllus  campestris, 
fig.  11)  consists,  as  described  by  Landois,^*  of  from  131  to  138 
sharp,  transverse  ridges  or  teeth  (.s^)  on  the  under  side  of  one  of 
the  nervures  of  the  wing-cover.  This  toothed  nervure  is  rapidly 
scraped  across  a  projecting,  smooth,  hard  ner- 
vure (r)  on  the  upper  surface  of  the  opposite 
wing.  First  one  wing  is  rubbed  over  the 
other,  and  then  the  movement  is  reversed. 
Both  wings  are  raised  a  little  at  the  same 
time,  so  as  to  increase  the  resonance.  In 
some  species  the  wing-covers  of  the  males  are 
furnished  at  the  base  with  a  talc-like  plate.^^ 
I  here  give  a  drawing  (fig.  Vl)  of  the  teeth  on 
the  under  side  of  the  nervure  of  another 
Fig.  12.  Teeth  of  Ner-  spccics  of  Grvllus,  viz.,  (J.  domesticus.  With 
SrcS^SoS)!'  'respect  to  the  formation  of  these  teeth.  Dr. 
Gruber  has  shewn  ^*^  that  they  have  been  de- 
veloped by  the  aid  of  selection,  from  the  minute  scales  and  hairs 
with  which  the  wings  and  body  are  covered,  and  I  came  to  the 
same  conclusion  with  respect  to  those  of  the  Coleoptera.  But 
Dr.  Gruber  further  shews  that  their  development  is  in  part 
directly  due  to  the  stimulus  from  the  friction  of  one  wing  over 
the  other. 

In  the  Locustidse  the  opposite  wing-covers  differ  from  each 
other  in  structure  (fig.  13),  and  the  action  caonot,  as  in  the 
last  family,  be  reversed.  The  left  wing,  which  acts  as  the 
bow,  lies  over  the  right  wing  which  serves  as  the  fiddle.  One 
of  the  nervures  (a)  on  the  under  surface  of  the  former  is 
finely  serrated,  and  is  scraped  across  the  prominent  nervures 
on  tlie  upper  surface  of  the  opposite  or  right  wing.  In  our 
British  fhasgonura  viridissima  it  appeared  to  me  that  the 
serrated  nervure  is  rubbed  against  the  rounded  hind-corner 
of  the  opposite  wing,  the  edge  of  which  is  thickened,  coloured 
brown,  and  very  sharp.  In  the  right  wing,  but  not  in  the  left, 
there  is  a  little  plate,  as  transparent  as  talc,  surrounded  by 
nervures,  and  called  the  speculum.  In  Epliiiiinger  vitium,  a 
member  of  this  same  family,  we  have  a  curious  subordinate 
modification ;  for  the  wing-covers  are  greatly  reduced  in  size, 
but  "  the  posterior  part  of  the  pro-thorax  is  elevated  into  a  kind 

^*  '  Zeitschi-ift    fiir   wissenschaft.  ^^    '  Ueber    der    Tonapparat    der 

Zoolocf.'  B.  xvii.  1867,  s.  117.  Locustiden,  ein  Beitrag   zum   Dar- 

^^  Westwood,   '  Modern  Class,    of  wiaismus,'  '  Zeitsch.  fiir  wissensch. 

Insects,'  vol.  i.  p.  440.  Zoolog.'  B.  xxii.  1872,  p.  100. 


Chap.  X. 


Orthoptera, 


285 


"of  dome  over  tlie  wing-covers,  and  which  hxis  probably  the 
"  effect  of  increasing  the  sound."  '^'^ 


Fig.  13.    ChloroccElus  Tanana  (from  Bates),    a,  b.  Lobes  ux"  opposite  wing-covers. 

We  thus  see  that  the  musical  apparatus  is  more  differentiated 
or  specialised  in  the  Locustidse  (which  include,  I  believe,  the 
most  powerful  performers  in  the  Order),  than  in  the  Achetidre, 
in  which  both  wing-covers  have  the  same  structure  and  the 
same  function.'^  Landois,  however,  detected  in  one  of  the 
Locustidae,  namely  in  Decticus,  a  short  and  narrow  row  of  small 
teeth,  mere  rudiments,  on  the  inferior  surface  of  the  right  wing- 
cover,  which  underlies  the  other  and  is  never  used  as  tlje  bow. 
I  observed  the  same  rudimentary  structure  on  the  under  side  of 
the  right  wing-cover  in  Phasyonura  viridissima.  Hence  we  may 
infer  with  confidence  that  the  Ijocustidsa  are  descended  from  a 
form,  in  which,  as  in  the  existing  Achetidse,  both  wing-covers 
had  serrated  nervures  on  the  under  surface,  and  could  be 
indifferently  used  as  the  bow ;  but  tliat  in  the  Locustidse  the 
two  wing-covers  gradually  became  differentiated  and  perfected. 


2'  Westwood,  'Modern  Class,  of  Insects,'  vol.  i.  p.  453. 

^^  Landois,  'Zeitsch.  f.  wiss.  Zoolog.'  B.  xvii.  1867,  s.  121,  122. 


286 


The  Descent  of  Man, 


Part  II. 


on  the  principle  of  the  division  of  labour,  the  one  to  act  ex- 
clusively as  the  bow,  and  the  other  as  the  fiddle.  Dr.  Gruber 
takes  the  same  view,  and  has  shewn  that  rudimentary  teeth  are 
commonly  found  on  the  inferior  surface  of  the  right  wing.  By 
what  steps  the  more  simple  apparatus  in  the  Achetidse  originated, 
we  do  not  know,  but  it  is  probable  that  the  basal  portions  of 
the  wing-covers  originally  overlapped  each  other  as  they  do  at 
present;  and  that  the  friction  of  the  nervures  produced  a 
gTating  sound,  as  is  now  the  case  with  the  wing-covers  of  tlie 
females.^^  A  grating  sound  thus  occasionally  and  accidentally 
made  by  the  males,  if  it  served  them  ever  so  little  as  a  love-call 
to  the  females,  might  readily  have  been  intensified  through 
sexual  selection,  by  variations  in  the  roughness  of  the  nervures 
having  been  continually  preserved. 

In  the  last  and  third  Family,  namely  the  Acridiidse  or 
grasshoppers,  the  stridulation  is  produced  in  a  very  different 
manner,  and  according  to  Dr.  Scudder,  is  not  so  shrill  as  in  the 
prec.eding  Families.  The  inner  surface  of  the  femur  (fig.  14,  r) 
is  furnished  with  a  longitudinal  row  of  minute,  elegant,  lancet- 
shaped,  elastic  teeth,  from  85  to  93  in  number  ;^*'  and  these  are 
scraped  across  the  sharp,  projecting  nervures  on  the  wing-covers, 
which  are  thus  made  to  vibrate  and  resound.  Karris'*^  says 
_  that  when  one  of  the  males 

begins  to  play,  he  first "  bends 
"  the  shank  of  the  hind-leg 
"  beneath  the  thigh,  where  it 
"  is  lodged  in  a  furrow  de- 
"  signed  to  receive  it,  and 
**  then  draws  the  leg  briskly 
"  up  and  down.  He  does  not 
"  play  both  fiddles  together, 
"  but  alternately,  first  upon 
"  one  and  then  on  the  other." 
In  many  species,  the  base 
of  the  abdomen  is  hollowed 
out  into  a  great  cavity  which 
is  believed  to  act  as  a  re- 
sounding board.  In  Pneu- 
mora  (fig.  15),  a  S.  African 
same  family,  we    meet  with  a  new 


Fig.  14.  Hind-leg  of  Stenobothrus  pratorum : 
r,  the  stridulating  ridge ;  lower  figure,  the 
teeth  forming  the  ridge,  much  magnified 
(from  Landois). 


genus   belonging  to  the 


39  Mr.  Walsh  also  informs  me 
that  he  has  noticed  that  the  female 
of  the  Platyphyllum  concavum, 
*'  when  captured  makes  a  feeble 
"grating    noise    by    shuffling   her 


"  wing-covers  together." 

*"  Landois,  ibid.  s.  113. 

*i    '  Insects    of    New    England, 
1842,  p.  133. 


Chap.  X. 


Orthoptera, 


2^7 


and  remarkable  modification;  in  the  males  a  small  notched 
ridge  projects  obliquely  from  each  side  of  the  abdomen, 
against  which  the  hind  femora  are  rubbed.^^  As  the  male  is 
furnished  with  wings  (the  female  being  wingless),  it  is  re- 
markable that  the  thighs  are  not  rubbed  in  tlie  usual  manner 
against  the  "wing-covers ;  but  this  may  perhaps  be  accounted  for 
by  the  unusually  small  size  of  the  hind-legs.  I  have  not  been 
able  to  examine  the  inner  surface  of  the  thighs,  which,  judging 


Fig.  15.    Pneumora  (from  specimens  in  the  British  Museum).    Upper  figure,  male; 
lower  figure,  female. 

from  analogy,  would  be  finely  serrated.  The  species  of  Pneumora 
have  been  more  profoundly  modified  for  the  sake  of  stridulation 
than  any  other  orthopterous  insect ;  for  in  the  male  the  whole 
body  has  been  converted  into  a  musical   instrument,  being 

*•  Westwood,  '  Mode]  u  Classification,'  vol.  i.  p.  462. 


288  The  Descent  of  Man.  Part  II. 

distended  witli  air,  like  a  great  pellucid  bladder,  so  as  to 
increase  the  resonance.  Mr.  Trimen  informs  me  that  at  the 
Cape  of  Good  Hope  these  insects  make  a  wonderful  noise 
during  the  night. 

In  the  three  foregoing  families,  the  females  are  almost  always 
destitute  of  an  efficient  musical  apparatus.  But  there  are  a  few 
exceptions  to  this  rule,  for  Dr.  Gruber  has  shewn  that  both 
sexes  of  Eplujypiger  vitium  are  thus  provided;  though  the  organs 
differ  in  the  male  and  female  to  a  certain  extent.  Hence  we 
cannot  suppose  that  they  have  been  transferred  from  the  male 
to  the  female,  as  appears  to  have  been  the  case  with  the  secondary 
sexual  characters  of  many  other  animals.  They  must  have  been 
independently  developed  in  the  two  sexes,  which  no  doubt 
mutually  call  to  each  other  during  the  season  of  love.  In  most 
other  Locustidse  (but  not  according  to  Landois  in  Decticus)  the 
females  have  rudiments  of  the  stridulatory  organs  proper  to  the 
male;  from  whom  it  is  probable  that  these  have  been  transferred. 
Landois  also  found  such  rudiments  on  the  under  surface  of  the 
wing-covers  of  the  female  Achetidse,  and  on  the  femora  of  the 
female  Acridiidte.  In  the  Homoptera,  also,  the  females  have  the 
proper  musical  apparatus  in  a  functionless  state ;  and  we  shall 
hereafter  meet  in  other  divisions  of  the  animal  kingdom  with 
many  instances  of  structures  proper  to  the  male  being  present 
in  a  rudimentary  condition  in  the  female. 

Landois  has  observed  another  important  fact,  namely,  that  in 
the  females  of  the  Acridiidte,  the  stridulating  teeth  on  the 
femora  remain  throughout  life  in  the  same  condition  in  which 
they  first  appear  during  the  larval  state  in  both  sexes.  In  the 
males,  on  the  other  hand,  they  become  further  developed,  and 
acquire  their  perfect  structure  at  the  last  moult,  when  the  insect 
is  mature  and  ready  to  breed. 

From  the  facts  now  given,  we  see  that  the  means  by  which 
the  males  of  the  Orthoptera  produce  their  sounds  are  extremely 
diversified,  and  are  altogether  different  from  those  employed  by 
the  Homoptera.^^  But  throughout  the  animal  kingdom  we 
often  find  the  same  object  gained  by  the  most  diversified  means ; 
this  seems  due  to  the  whole  organisation  having  undergone  mul- 
tifarious changes  in  the  course  of  ages,  and  as  part  after  part 
varied  different  variations  were  taken  advantage  of  for  the 
same  general  purpose.  The  diversity  of  means  for  producing 
sound  in  the  three  families    of   the    Orthoj^tera   and  in  the 

*^  Landois  has  recently  found  in  moptera ;   and    this   is  a   surprising 

certain     Orthoptera       rudimentary  fact.      See  '  Zeitschr.  fiir  wissensch. 

structures    closely    similar    to    the  Zoolog.'  B.   xxii.  Heft    3,  1871,  p. 

sound-producing  organs  in  the  Ho-  348. 


Chap.  X.-  Or t hop t era.  289 

Bomoptera,  impresses  the  mind  with  the  high  importance  of  these 
structures  to  the  males,  for  the  sake  of  calling  or  alluring  the 
females.  We  need  feel  no  surprise  at  the  amount  of  modification 
which  the  Orthoptera  have  undergone  in  this  respect,  as  we  now 
know,  from  Dr.  Scudder's  remarkable  discovery/*  that  there  has 
been  more  than  ample  time.  This  naturalist  has  lately  found 
a  fossil  insect  in  the  Devonian  formation  of  New  Brunswick, 
which  is  furnished  with  "  the  well-known  tympanum  or  stridu- 
"  lating  apparatus  of  the  male  Locustidse."  The  insect,  though 
in  most  respects  related  to  the  Neuroptera,  appears,  as  is  so  often 
the  case  with  very  ancient  forms,  to  connect  the  two  related 
Orders  of  the  Neuroptera  and  Orthoptera. 

I  have  but  little  more  to  say  on  the  Orthoptera.  Some  of  the 
species  are  very  pugnacious:  when  two  male  field-crickets 
(GryUus  camjoestris)  are  confined  together,  they  fight  till  one 
kills  the  other;  and  the  species  of  Mantis  are  described  as 
mancBuvriug  with  their  sword-like  front-limbs,  like  hussars  with 
their  sabres.  The  Chinese  keep  these  insects  in  little  bamboo 
cages,  and  match  them  like  game-cocks.'*^  With  respect  to 
colour,  some  exotic  locusts  are  beautifully  ornamented ;  the 
posterior  wings  being  marked  with  red,  blue,  and  black ;  but  as 
throughout  the  Order  the  sexes  rarely  differ  much  in  colour,  it 
is  not  probable  that  they  owe  their  bright  tints  to  sexual 
selection.  Conspicuous  colours  may  be  of  nse  to  these  insects, 
by  giving  notice  that  they  are  unpalatable.  Thus  it  has  been 
observed  ^°  that  a  bright-coloured  Indian  locust  was  invariably 
rejected  when  offered  to  birds  and  lizards.  Some  cases,  however, 
are  known  of  sexual  differences  in  colour  in  this  Order.  The 
male  of  an  American  cricket-*^  is  described  as  being  as  white  as 
ivory,  whilst  the  female  varies  from  almost  white  to  greenish- 
yellow  or  dusky.  Mr.  Walsh  informs  me  that  the  adult  male  of 
Spectrum  femoratum  (one  of  the  Phasmidse)  "  is  of  a  shining 
"  brownish-yellow  colour ;  the  adult  female  being  of  a  dull, 
"  opaque,  cinereous  brown ;  the  young  of  both  sexes  being  green." 
Lastly,  I  may  mention  that  the  male  of  one  curious  kind  of 
cricket*^  is  furnished  with  "a  long  membranous  appendage, 
"  which  falls  over  the  face  like  a  veil ;"  but  what  its  use  may  be, 
is  not  known. 

**  '  Transact.  Ent.  Soc'  3r J  series,  *^  The  CEcantlms  nivalis.    Harris, 

vol.  ii.     ('Journal  of  Proceedings,'  'Insects  of  New  England,'  1842,  p.. 

p.  117.)  "  124.    The  two  sexes  of  (E.  pcllucichts ' 

*^  Westwood,   '  Modern  Class,   of  of   Europe  differ,  as    I    hear    from 

Insects,'  vol.  i.  p.  427  ;  for  crickets,  Victor  Carus,   in   nearly  the   same 

p.  445.  manner. 

*^  Mr.  Ch.  Home,  in  '  Proc.  Ent.  **      Platyblemnus  :       Westwood, 

Soc '  May  3,  1869,  p.  xii.  '  Modern  Class.'  vol.  i.  p.  447. 


2QO  The  Descent  of  Man.  Part  II. 

Order,  Neuroj)tera. — Little  need  here  be  said,  except  as  to 
colour.  In  the  Ephemeridse  the  sexes  often  differ  slightly  in 
their  obscure  tints ;  *^  but  it  is  not  probable  that  the  males  are 
thus  rendered  attractive  to  the  females.  The  Ijibellulidae,  or 
dragon-flies,  are  ornamented  with  sjDlendid  green,  blue,  yellow, 
and  vermilion  metallic  tints ;  and  the  sexes  often  differ.  Thus. 
as  Prof.  Westwood  remarks,^"  the  males  of  some  of  the 
Agrionidae,  "  are  of  a  rich  blue  with  black  wings,  whilst  the 
'*  females  are  fine  green  with  colourless  wings."  But  in  Agrion 
Bamhurii  these  colours  are  exactly  reversed  in  the  two  sexes.^^ 
In  the  extensive  N.  American  genus  of  Hetserina,  the  males  alone 
have  a  beautiful  carmine  spot  at  the  base  of  each  wing.  In 
Anax  Junius  the  basal  part  of  the  abdomen  in  the  male  is  a  vivid 
ultramarine  blue,  and  in  the  female  grass-green.  In  the  allied 
genus  Gomphus,  on  the  other  hand,  and  in  some  other  genera, 
the  sexes  differ  but  little  in  colour.  In  closely-allied  forms 
throughout  the  animal  kingdom,  similar  cases  of  the  sexes 
differing  greatly,  or  very  little,  or  not  at  all,  are  of  frequent 
occurrence.  Although  there  is  so  wide  a  difference  in  colour 
between  the  sexes  of  many  Libellulidse,  it  is  often  difficult  to  say 
which  is  the  more  brilliant ;  and  the  ordinary  coloration  of  the 
two  sexes  is  reversed,  as  we  have  just  seen,  in  one  species  of 
Agrion.  It  is  not  probable  that  their  colours  in  any  case  have 
been  gained  as  a  protection.  Mr.  MacLachlan,  who  has  closely 
attended  to  this  family,  writes  to  me  that  dragon-flies— the 
•tyi-ants  of  the  insect-world — are  the  least  liable  of  any  insect  to 
be  attacked  by  birds  or  other  enemies,  and  he  believes  that  their 
bright  colours  serve  as  a  sexual  attraction.  Certain  dragon-flies 
apparently  are  attracted  by  particular  colours :  Mr.  Patterson 
observed  ^^  that  the  Agrionidrs,  of  which  the  males  are  blue, 
settled  in  numbers  on  the  blue  float  of  a  fishing  line ;  whilst  two 
other  species  were  attracted  by  shining  white  colours. 

It  is  an  interesting  fact,  first  noticed  by  Schelver,  that,  in 
several  genem  belonging  to  two  sub-families,  the  males  on  first 
emergence  from  the  pupal  state,  are  coloured  exactly  like  the 
females;  but  that  their  bodies  in  a  short  time  assume  a  con- 
spicuous milky-blue  tint,  owing  to  the  exudation  of  a  kind  of  oil, 
soluble  in  ether  and  alcohol.  Mr.  MacLachlan  believes  that  in 
the  male  of  Lihelkda  depressa  this  change  of  colour  does  not  occur 
until  nearly  a  fortnight  after  the  metamorphosis,  when  the  sexes 
are  ready  to  pair. 

*^  B.  D.  Walsh,  the  '  Pseudo-neu-  indebted  to  this  naturalist  for  the 

roptera  of  Illinois,'  in  '  Proc.  Ent.  following  facts   on  Hetarina,  Anax, 

Soc.  of  Philadelphia,'  1862,  p,  361.  and  Gomphus. 

5*  'Modern  Class.' vol.  ii.  p.  37.  ^-  'Transact.    Ent.    Soc'    vol.    i. 

"  Walsh,   ibid.    p.    381.      I  am  1836,  p.  Ixxxi. 


CiiAP.  X.  HynieiiGptera.  291 

Certain  si3ecies  of  Neurotliemis  present,  according  to  Brauer,  °^ 
a  curious  case  of  dimorphism,  some  of  the  females  having  ordinary 
wings,  whilst  others  have  them  "  very  richly  netted,  as  in  the 
"  males  of  the  same  species."  Brauer  "  exjilains  the  phenomenon 
"  on  Darwinian  principles  by  the  suj^position  that  the  close 
"  netting  of  the  veins  is  a  secondary  sexual  character  in  the 
"  males,  which  has  been  abruptly  transferred  to  some  of  the 
"  females,  instead  of,  as  generally  occurs,  to  all  of  them."  Mr. 
MacLachlan  informs  me  of  another  instance  of  dimorphism 
in  several  species  of  Agrion,  in  which  some  individuals  are  of 
an  orange  colour,  and  these  are  invariably  females.  This  is 
probably  a  case  of  reversion ;  for  in  the  true  Libellulse,  when 
the  sexes  differ  in  colour,  the  females  are  orange  or  yellow; 
so  that  supposing  Agrion  to  be  descended  from  some  primordial 
form  which  resembled  the  typical  Libellulse  in  its  sexual  cha- 
racters, it  would  not  be  surprising  that  a  tendency  to  vary  in 
this  manner  should  occur  in  the  females  alone. 

Although  many  dragon-flies  are  large,  powerful,  and  fierce 
insects,  the  males  have  not  been  observed  by  Mr.  MacLachlan  to 
fight  together,  excepting,  as  he  believes,  in  some  of  the  smaller 
sj^ecies  of  Agrion.  In  another  group  in  this  Order,  namely,  the 
Termites  or  white  ants,  both  sexes  at  the  time  of  swarming  may 
be  seen  running  about,  "  the  male  after  the  female,  sometimes 
"  two  chasing  one  female,  and  contending  with  great  eagerness 
"  who  shall  win  the  prize."^^  The  Atropos  pulsatorius  is  said 
to  make  a  noise  with  its  jaws,  which  is  answered  by  other 
individuals.^^ 

Order,  Hymenoptera. — That  inimitable  observer,  M.  Fabre,^"  in 
describing  the  habits  of  Cerceris,  a  wasp-like  insect,  remarks  that 
"  fights  frequently  ensue  between  the  males  for  the  possession  of 
"  some  particular  female,  who  sits  an  apparently  unconcerned 
"  beholder  of  the  struggle  for  supremacy,  and  when  the  victory 
"  is  decided,  quietly  flies  away  in  company  with  the  conqueror." 
Westwood  ^'  says  that  the  males  of  one  of  the  saw-flies  (Tenthre- 
dinse)  "  have  been  found  fighting  together,  with  their  mandibles 
"  locked."  As  M.  Fabre  speaks  of  the  males  of  Cerceris  striving 
to  obtain  a  particular  female,  it  may  be  well  to  bear  in  mind 
that  insects  belonging  to  this  Order  have  the  power  of  recognising 

^'  See  abstract  in  the  '  Zoological  ^^   See     an     interesting     article. 

Record'  for  1867,  p.  450.  'The  Writings  of  Fabre,'  in  'Nat. 

^■*  Kirby  and  Spence,  'Introduct.  Hist.  Review,'  April  1862,  p.  122. 
to  Entomology,'  vol.  ii.  1818,  p.  35.  ^'  'Journal  of  Proc.  of  Entoraolog. 

"  Houzeau,  '  Les    Facultes  Men-  Soc'  Sept,  7th,  1863,  p.  169. 
tales,'  &c.     Tom.  1.  p.  104. 


292  The  Descent  of  Man.  Fart  II. 


each  other  after  long  intervals  of  time,  and  are  deeply  attached. 
For  instance,  Pierre  Hnber,  whose  accuracy  no  one  doubts, 
separated  some  ants,  and  when,  after  an  interval  of  four  months, 
they  met  others  which  had  formerly  belonged  to  the  same- 
community,  they  recognised  and  caressed  one  another  with  their 
antennae.  Had  they  been  strangers  they  would  have  fought 
together.  Again,  when  two  communities  engage  in  a  battle,  the 
ants  on  the  same  side  sometimes  attack  each  other  in  tlie  general 
confusion,  but  they  soon  perceive  their  mistake,  and  the  one  ant 
soothes  tbe  other.^^ 

In  this  Order  slight  differences  in  colour,  according  to  sex,  are 
common,  but  conspicuous  differences  are  rare  except  in  the 
family  of  Bees ;  yet  both  sexes  of  certain  groups  are  so  brilliantly 
coloured— for  instance  in  Chrysis,  in  which  vermilion  and 
metallic  greens  prevail — that  we  are  tempted  to  attribute  the 
result  to  sexual  selection.  In  the  Ichneumonids^,  according  to 
Mr.  Walsli,^^  the  males  are  almost  universally  lighter-coloured 
than  the  females.  On  the  other  hand,  in  the  Tenthredinidse  the 
males  are  generally  darker  than  the  females.  In  the  Siricidse 
the  sexes  frequently  differ ;  thus  the  male  of  Sirex  juvencus  is 
banded  with  orange,  whilst  the  female  is  dark  purple  ;  but  it  is 
difficult  to  say  which  sex  is  the  more  ornamented.  In  Tremex 
cohnnhop.  the  female  is  much  brighter-coloured  than  the  male. 
I  am  informed  by  Mr.  F.  Smith,  that  the  male  ants  of  several 
species  are  black,  the  females  being  testaceous. 

In  the  family  of  Bees,  especially  in  the  solitary  species,  as  I 
hear  from  the  same  entomologist,  the  sexes  often  differ  in  colour. 
The  males  are  generally  the  brighter,  and  in  Bombus  as  well  as  in 
Apathus,  much  more  variable  in  colour  than  the  females.  In 
Anthophora  return  the  male  is  of  a  rich  fulvous-brown,  whilst 
the  female  is  quite  black  :  so  are  the  females  of  several  species 
of  Xylocopa,  the  males  being  bright  yellow.  On  the  other  hand 
the  females  of  some  species,  as  of  Andrcena  fiilva,  are  much 
brighter-coloured  than  the  males.  Such  differences  in  colour' 
can  hardly  be  accounted  for  by  the  males  being  defenceless  and 
thus  requiring  protection,  whilst  the  females  are  well  defended 
by  their  stings.  H.  Miiller,  *^°  who  has  particularly  attended  to 
the  habits  of  bees,  attributes  these  differences  in  colour  in  chief 
part  to  sexual  selection.  That  bees  have  a  keen  perception  of 
colour  is  certain.  He  says  that  the  males  search  eagerly  and 
fight  for  the  possession  of  the  females ;  and  he  accounts  thi'ough 

58  p.  Huber,  '  Recherches  sur  les  Philadelphia,'  1866,  pp.  238-239. 

Moeurs  des  Fourmis,'  1810,  pp.  150,  ^^  '  Anwendung  der  Darwinschen 

165.  Lehi-e    auf    Bienen.'      Verh.    d.    li. 

^^  '  Proc.     Entomolog.     Soc.     of  Jahrg.  xxix. 


Chap.  X.  Hyniienoptera.  293 

such  contests  for  the  mandibles  of  the  males  being  in  certain 
species  larger  than  tliose  of  the  females.  In  some  cases  tlie 
males  are  far  more  numerous  than  the  females,  either  early 
in  the  season,  or  at  all  times  and  places,  or  locally ;  whereas  the 
females  in  other  cases  are  apparently  in  excess.  In  some  species 
the  more  beautiful  males  appear  to  have  been  selected  by  the 
females  ;  and  in  others  the  more  beautiful  females  by  the  males. 
Consequently  in  certain  genera  (Miiller,  p.  42),  the  males  of  the 
several  species  differ  much  in  appearance,  vs^hilst  the  females  are 
almost  indistinguishable;  in  other  genera  the  reverse  occurs. 
H.  Miiller  believes  (p.  82)  that  the  colours  gained  by  one  sex 
through  sexual  selection  have  often. been  transferred  in  a  variable 
degree  to  the  other  sex,  just  as  the  pollen-collecting  apparatus 
of  the  female  has  often  been  transferred  to  the  male,  to  whom 
it  is  absolutely  useless.''^ 

Mutilla  Earopxa  makes  a  stridulating  noise ;  and  according  to 
Goureau  ^'^  both  sexes  have  this  power.  He  attributes  the  sound 
to  the  friction  of  the  third  and  preceding  abdominal  segments, 
and  I  find  that  these  surfaces  are  marked  with  very  fine  con- 
centric ridges ;  but  so  is  the  projecting  thoracic  collar,  into  which 
the  head  articulates,  and  this  collar,  when  scratched  with  the 
point  of  a  needle,  emits  the  proper  sound.  It  is  rather  surprising 
that  both  sexes  should  have  the  power  of  stridulating,  as  the 
male  is  winged  and  the  female  wingless.  It  is  notorious  that 
Bees  express  certain  emotions,  as  of  anger,  by  the  tone  of  their 
humming;  and  according  to  H.  Miiller  (p.  80),  the  males  of 
some  species  make  a  peculiar  singing  noise  whilst  pursuing  the 
females. 

®^  M.  Perrier  in  his  article  'la  Se-  male  grandfathers?     To  take  a  case 

lection  sexuelled'aprfes  Darwin '('Re-  with    ordinary    animals    as     nearly 

vue  Scientitiqiae,'  Feb.  1873,  p.  868),  parallel  as  possible:  if  a  female  of 

without  apparently  having  reflected  any  white  quadruped  or  bird  were 

much  on  the  subject,  objects  that  as  crossed  by  a  male  of  a  black  breed, 

the  males  of  social  bees  are  known  and  the  male  and   female  offspring 

to    be    produced    from    unfertilised  were    paired    together,    will    it    be 

ova,  they  could    not   transmit  new  pretended    that    the    grandchildren 

characters  to  their  male  offspring.  would    not    inherit   a   tendency    to 

This  is  an  extraordinary  objection,  blackness   from    their  male   grand- 

A  female  bee  fertilised  by  a  male,  father?      The  acquirement  of  new 

which  presented  some  character  fa-  characters  by  the  sterile  worker-bees 

cilitating  the  union  of  the  sexes,  or  is  a  much  more  difficult  case,  but  I 

rendering    him    more   attractive   to  have    endeavoured   to    show  in  my 

the  female,  would   lay  eggs  which  '  Origin  of  Species,' how  these  sterile 

would   produce    only  females  ;    but  beings  are  subjected  to  the  power  of 

these    young    females    would    next  natural  selection, 

year  produce  males  ;   and  will  it  be  ^^  Quoted  by  Westwood,  '  Modern 

pretended    that    such    males  would-  Class,  of  Insects,' vol.  ii.  p.  214-. 
not  inherit  the  characters  of  their 


294  The  Descent  of  Man.  Part  II. 

Order,  Coleoj)tera  (Beetles). — Many  beetles  are  coloured  so  as 
to  resemble  the  surfaces  which  they  habitually  frequent,  and 
they  thus  escape  detection  by  their  enemies.  Other  species,  for 
instance  diamond-beetles,  are  ornamented  with  siDlendid  colours, 
which  are  often  arranged  in  stripes,  spots,  crosses,  and  other 
elegant  patterns.  Such  colours  can  hardly  serve  directly  as  a 
protection,  except  in  the  case  of  certain  flower-feeding  species ; 
but  they  may  serve  as  a  warning  or  means  of  recognition,  on  the 
same  principle  as  the  phosphorescence  of  the  glow-worm.  As 
with  beetles  the  colours  of  the  two  sexes  are  generally  alike,  we 
have  no  evidence  that  they  have  been  gained  through  sexual 
selection;  but  this  is  at  least  possible,  for  they  may  have  been 
developed  in  one  sex  and  then  transferred  to  the  other ;  and 
this  view  is  even  in  some  degree  probable  in  those  groups  which 
possess  other  well-marked  secondary  sexual  characters.  Blind 
beetles,  which  cannot  of  course  behold  each  other's  beauty, 
never,  as  I  hear  from  Mr.  Waterhouse,  jun.,  exhibit  bright 
colours,  though  they  often  have  polished  coats ;  but  the  expla- 
nation of  their  obscurity  may  be  that  they  generally  inhabit 
caves  and  other  obscure  stations. 

Some  Longicorns,  especially  certain  Prionidse,  offer  an  excep- 
tion to  the  rule  that  the  sexes  of  beetles  do  not  differ  in  colour. 
Most  of  these  insects  are  large  and  splendidly  coloured.  The 
males  in  the  genus  Pyrodes,*^^  which  I  saw  in  Mr.  Bates's  col- 
lection, are  generally  redder  but  rather  duller  than  the  females, 
the  latter  being  coloured  of  a  more  or  less  splendid  golden-green. 
On  the  other  hand,  in  one  species  the  male  is  golden-green,  the 
female  being  richly  tinted  with  red  and  purple.  In  the  genus 
Esmeralda  the  sexes  differ  so  greatly  in  colour  that  they  have 
been  ranked  as  distinct  species ;  in  one  species  both  are  of  a 
beautiful  shining  green,  but  the  male  has  a  red  thorax.  On  the 
whole,  as  far  as  I  could  judge,  the  females  of  those  Prionidse,  in 

"^    Pyrodes       pulcherrimus,       in  the  family  of  Longicorns.     Messrs. 

which  the  sexes  differ  conspicuously,  R.    Trimen   and   Waterhouse,   jun., 

has  been  described  by  Mr.  Bates  in  inform    me    of    two    Lamellicorns, 

'Transact.  Ent.   Soc'  1869,   p.    50.  viz.,  a  Peritrichia  and  Trichius,  the 

I  will  specify  the  few  other  cases  in  male     of     the     latter    being    more 

which  1  have  heard  of  a  difference  obscurely  coloured  than  the  female. 

m    colour    between    the    sexes    of  In  Tillus  elongatus  the  male  is  black, 

beetles.      Kirby    and    Spence    ('  In-  and    the    female    always,    as    it    is 

troduct.  to  Entomology,'  vol.  iii.  p.  believed,  of  a  dark  blue  colour,  with 

301)   mention    a  Cantharis,  Meloc,  a   red   thorax.     The    male,  also,  of 

Rhagium,  and  the  Leptura  testacca ;  Orsodacna  atra,  as  I  hear  from  Mr. 

the  male  of  the   latter   being    tes-  Walsh,   is    black,    the    female    (the 

taceous,  with   a  black   thorax,  and  so-called     0.    ruficollis)    having   .a 

the   female  of  a  dull  red  all  over,  rufous  thorax. 
The.se  two  latter  beetles  belong  to 


Chap.  X. 


Coleoptera. 


295 


which  the  sexes  differ,  are  coloured  more  richly  than  the  males, 
and 'this  does  not  accord  with  the  common  rule  in  regard  to 
colour,  when  acquired  through  sexual  selection. 

A  most  remarkable  distinction  between  the  sexes  of  many 
beetles  is  presented  by  the  great  horns  which  rise  from  the  head, 
thorax,  and  clypeus  of  the  males ;  and  in  some  few  cases  from 
the  under  surface  of  the  body.  These  horns,  in  the  great  family 
of  the  Lamellicorns,  resemble  those  of  various  quadrupeds,  such 
as  stags,  rhinoceroses,  &c.,  an-d  are  wonderful  both  from  their 
size  and  diversified  shapes.  Instead  of  describing  them,  I  have 
given  figures  of  the  males  and  females  of  some  of  the  more  re- 
markable forms.  (Figs.  16  to  20.)  The  females  generally  ex- 
liibit  rudiments  of  the  horns  in  the  form  of  small  knobs  or 
ridges ;  but  some  are  destitute  of  even  the  slightest  rudiment. 
On  the  other  hand,  the  horns  are  nearly  as  well  developed  in  the 
female  as  in  the  male  of  Phanceus  lancifer ;  and  only  a  little  less 
well  developed  in  the  females  of  some  other  species  of  this  genus 
and  of  Copris.  I  am  informed  by  Mr.  Bates  that  the  horns  do 
not  differ  in  any  manner  corresponding  with  the  more  important 
characteristic  differences  between  the  several  subdivisions  of  the 
family :  thus  within  the  same  section  of  the  genus  Onthophagus, 
there  are  species  which  have  a  single  horn,  and  others  which 
have  two. 


Fig.  16.    Chalcosoma  atlas.    Upper  figure,  male  (roduced) ;  lower  figure,  female 
(nat.  size). 


296 


'The  Descent  of  Man.  Part  II. 


Copris  isidis.    (Left-hand  figures,  males.) 


Plianaeus  faunus. 


Fisr.  19 


ripelicus  cantoii. 


Fig.  20.  Onthophagiis  rangifer,  enlarged 


Chap.  X.  '  Coleoptera.  '  297 

In  almost  all  cases,  the  horns  are  remarkable  from  their  ex- 
cessive variability;  so  that  a  graduated  series  can  be  formed, 
from  the  most  highly  developed  males  to  others  so  degenerate 
that  they  can  barely  bo  distinguished  from  the  females.  Mr. 
Walsh ^^  found  that  in  Phcuueus  caniifex  the  horns  were  thrice  as 
long  in  some  males  as  in  others.  Mr.  Bates,  after  examining 
above  a  hundred  males  of  Ontliophagus  runcjifer  (fig.  20),  thought 
that  he  had  at  last  discovered  a  species  in  which  the  horns  did 
not  vary ;  but  further  research  proved  the  contrary. 

The  extraordinary  size  of  the  horns,  and  their  widely  different 
structure  in  closely-allied  forms,  indicate  that  they  have  been 
formed  for  some  purpose ;  but  their  excessive  variability  in  the 
males  of  the  same  species  leads  to  the  inference  that  this  purpose 
cannot  be  of  a  definite  nature.  The  horns  do  not  show  marks  of 
friction,  as  if  used  for  any  ordinary  work.  Some  authors  sup- 
pose "^^  that  as  the  males  wander  about  much  more  than  the 
females,  they  require  horns  as  a  defence  against  tJieir  enemies ; 
but  as  the  horns  are  often  blunt,  they  do  not  seem  well  adapted 
for  defence.  The  most  obvious  conjecture  is  that  they  are  used 
by  the  males  for  fighting  together ;  but  the  males  have  never 
been  observed  to  fight;  nor  could  Mr.  Bates,  after  a  careful 
examination  of  numerous  species,  find  any  sufficient  evidence,  in 
their  mutilated  or  broken  condition,  of  their  having  been  thus 
used.  If  the  males  had  been  habitual  fighters,  the  size  of  their 
bodies  would  probably  have  been  increased  through  sexual 
selection,  so  as  to  have  exceeded  that  of  the  females;  but 
Mr.  Bates,  after  comparing  the  two  sexes  in  above  a  hundred 
species  of  the  Copridse,  did  not  find  any  marked  difference  in 
this  respect  amongst  well-developed  individuals.  In  Lethrus, 
moreover,  a  beetle  belonging  to  the  same  great  division  of  the 
Lamellicorns,  the  males  are  known  to  fi^'ht,  but  are  not  provided 
with  horns,  though  their  mandibles  are  much  larger  than  those 
of  the  female. 

The  conclusion  that  the  horns  have  been  acquired  as  ornaments 
is  that  which  best  agrees  with  the  fact  of  their  having  been  so 
immensely,  yet  not  fixedly,  developed, — as  shewn  by  their  extreme 
variability  in  the  same  species,  and  by  their  extreme  diversity  in 
closely- allied  species.  This  view  will  at  first  appear  extremely 
improbable;  but  we  shall  hereafter  find  with  many  animals 
standing  much  higher  in  the  scale,  namely  fishes,  amphibians, 
reptiles  and  birds,  that  various  kinds  of  crests,  knobs,  horns  and 
combs  have  been  developed  apparently  for  this  sole  purpose. 

The  males  of  Onitls  furciftr  (fig.   21),  and  of  some  other 

^■*    '  Pi-oc.    Entomolog.     Soc.     of  ^^  Kirby  and  Spence,  '  latroduct. 

rhiladelphia,'  1864,  p.  228.  Entomolog.'  vol.  iii.  p.  300. 

14 


298 


^he  Descent  of  Man. 


Part  II. 


species  of  tlie  genus,  are  furnished  with  singular  projections  on 
their  anterior  femora,  and  with  a  great  fork  or  pair  of  horns  on 
the  lower  surface  of  the  thorax.  Judging 
from  other  insects,  these  may  aid  the  male 
in  clinging  to  the  female.  Although  the 
males  have  not  even  a  trace  of  a  horn  on 
the  upper  surface  of  the  body,  yet  the  fe- 
males plainly  exhibit  a  rudiment  of  a  single 
horn  on  the  head  (fig.  22,  a),  and  of  a  crest 
(&)  on  the  thorax.  That  the  slight  thoracic 
crest  in  the  female  is  a  rudiment  of  a  pro- 
jection proper  to  the  male,  though  entirely 
absent  in  the  male  of  this  jDarticular  species, 
is  clear :  for  the  female  of  Buhas  bison  (a 
genus  which  comes  next  to  Onitis)  has  a 
similar  slight  crest  on  the  thorax,  and  the  male  bears  a  great 
projection  in  the  same  situation.  So,  again,  there  can  hardly  be 
a  doubt  that  the  little  point  (a)  on  the  head  of  the  female  Onitis 


Fig  21.  Onitis  furcifer, 
male  viewed  from  be- 
neath. 


Fig.  22.     Left-hand  fignre,  male  of  Onitis  furcifer,  viewed  laterally.     Right-hand 
figure,  female,    a.  Rudiment  of  cephalic  horn.    b.  Truce  of  thoracic  horn  or  crest. 

furcifer,  as  well  as  on  the  head  of  the  females  of  two  or  three 
allied  species,  is  a  rudimentary  representative  of  the  cephalic 
horn,  which  is  common  to  the  males  of  so  many  Lameilicorn 
beetles,  as  in  Phanseus  (fig.  18). 

The  old  belief  that  rudiments  have  been  created  to  complete 
the  scheme  of  nature  is  here  so  far  from  holding  good,  that  we 
have  a  complete  inversion  of  the  ordinary  state  of  things  in  the 
family.  We  may  reasonably  suspect  that  the  males  originally 
bore  horns  and  transferred  them  to  the  females  in  a  rudimentary 
condition,  as  in  so  many  othej*  Lamellicorns.  Why  the  males 
subsequently  lost  their  horns,  we  know  not ;  but  this  may  have 
been  caused  through  the  principle  of  compensation,  owing  to 
the  development  of  the  large  horns  and  projections  on  the  lower 
surface ;  and  as  these  are  confined  to  the  males,  the  rudiments 
of  the  upper  horns  on  the  females  would  not  have  been  thus 
obliterated. 


Chap.  X.  Coleoptera,  299 


The  cases  hitherto  given  refer  to  the  Lamellicorns,  but  the 
males  of  some  few  other  beetles,  belonging  to  two  widely  distinct 
groups,  namely,  the  Cui'culionidre  and  Staphylinidce,  are  fur- 
nished with  homs — in  the  former  on  the  lower  surface  of  the 
body,*"^  in  the  latter  on  the  upi^er  surface  of  the  head  and  thorax. 
In  the  Staphylinid?e,  the  horns  of  the  males  are  extraordinarily 
variable  in  the  same  species,  just  as  we  have  seen  with  the 
Lamellicorns.  In  Siagonium  we  have  a  case  of  dimorphism, 
for  the  males  can  be  divided  into  two  sets,  differing  greatly 
in  tiie  size  of  their  bodies  and  in  the  development  of  their 
horns,  without  intermediate  gradations.  In  a  species  of  Bledius 
(fig.  23),  also  belonging  to  the  Staphylinidse,  Professor  Westwood 


Fig.  23.    Bledius  taurus,  magnified.    Left-hand  figure,  male ;  right-hand  figure  female. 

states  that,  "male  specimens  can  be  found  in  the  same  locality 
"  in  which  the  central  horn  of  the  thorax  is  very  large,  but  the 
"  horns  of  the  head  quite  rudimental ;  and  others,  in  which  the 
"thoracic  horn  is  much  shorter,  whilst  the  protuberances  on 
"  the  head  are  long."  *^^  Here  we  apparently  have  a  case  of 
compensation,  which  throws  light  on  that  just  given  of  the 
supposed  loss  of  the  upper  horns  by  the  males  of  Onitis. 

Law  of  Battle. — Some  male  beetles,  which  seem  ill-fitted  .for 
fighting,  nevertheless  engage  in  conflicts  for  the  possession  of 
the  females.  Mr.  Wallace  *^^  saw  two  males  of  Leptorhynchus 
angustatus,  a  linear  beetle  with  a  much  elongated  rostrum, 
"  fighting  for  a  female,  who  stood  close  by  busy  at  her  boring. 
"  They  pushed  at  each  other  with  their  rostra,  and  clawed  and 
"  thumped,  apparently  in  the  greatest  rage."  The  smaller  male, 
however,  "  soon  ran  away,  acknowledging  himself  vanquished." 
In  some  few  cases  male  beetles  are  well  adapted  for  fighting,  by 
possessing  great  toothed  mandibles,  much  larger  than  those  of 
the  females.  This  is  the  case  with  the  common  stag-beetle 
{Lucanus  cervus),  the  males  of  which  emerge  from  the  pupal 
state  about  a  week  before  the  other  sex,  so  that  several  may 
often  be  seen  pursuing  the  same  female.     At  this  season  they 

®®  Kirby  and  Spence,  '  Introduct.  gonium   in   an   intermediate    condi- 

Entomolog.'  vol.  iii.  p.  329.  tion,  .so  that  the  dimorphism  is  not 

®^  '  Modern  Classification  of  In-  strict, 
sects,'  vol.    i.   p.   172:    Siagonium,  ^^  'The  Malay  Archijjelago,' vol. 

p.  172.     In  the  British   Museum  I  ii.  1869,  p.  276.  Riley,  Sixth "*  Report 

noticed  one  male  specimen  of  Sia-  on  insects  of  Missouri/ 1874,  p.  115. 


300  The  Descent  of  Mmi.  Part  II. 


engage  in  fierce  conflicts.  When  Mr.  A.  H.  Davis  *^^  enclosed 
two  males  with  one  female  in  a  box,  the  larger  male  severely 
I.)inched  the  smaller  one,  until  he  resigned  his  pretensions.  A 
friend  informs  me  that  when  a  boy  he  often  iDut  the  males 
together  to  see  them  fight,  and  he  noticed  that  they  were  much 
bolder  and  fiercer  than  the  females,  as  with  the  higher  animals. 
The  males  would  seize  hold  of  his  finger,  if  held  in  front  of 
them,  but  not  so  the  females,  although  they  have  stronger 
jaws.  The  males  of  many  of  the  Lucanidse,  as  well  as  of  the 
above-mentioned  Leptorhynchus,  are  larger  and  more  powerful 
insects  than  the  females.  The  two  sexes  of  Lethnis  cepJialotes 
(one  of  the  Lamellicorns)  inhabit  the  same  burrow;  and  the 
male  has  larger  mandibles  than  the  female.  If,  during  the 
breeding-season,  a  strange  male  attempts  to  enter  the  burrow, 
he  is  attacked ;  the  female  does  not  remain  passive,  but  closes 
the  mouth  of  the  burrow,  and  encourages  her  mate  by  con- 
tinually pushing  him  on  from  behind ;  and  the  battle  lasts  until 
the  aggressor  is  killed  or  runs  away. '**  The  two  sexes  of  another 
Lamellicorn  beetle,  the  Ateuchus  cicafricosus,  live  in  pairs,  and 
seem  much  attached  to  each  other ;  the  male  excites  the  female 
to  roll  the  balls  of  dung  in  which  the  ova  are  deposited ;  and  if 
she  is  removed,  he  becomes  much  agitated.  If  the  male  is 
removed  the  female  ceases  all  work,  and  as  M.  Brulerie  '^^  believes, 
would  remain  on  the  same  spot  until  she  died. 

The  great  mandibles  of  the  male  Lucanidse  are  extremely 
variable  both  in  size  and  structure,  and  in  this  respect  resemble 
the  horns  on  the  head  and  thorax  of  many  male  Lamellicorns 
and  Staphylinidse.  A  perfect  series  can  be  formed  from  the 
best-provided  to  the  worst-provided  or  degenerate  males.  Al- 
though the  mandibles  of  the  common  stag-beetle,  and  probably 
of  many  other  species,  are  used  as  efficient  weapons  for  fighting, 
it  is  doubtful  whether  their  great  size  can  thus  be  accounted 
for.  We  have  seen  that  they  are  used  by  the  Lucanus  elaphus 
of  N.  America  for  seizing  the  female.  As  they  are  so  con- 
spicuous and  so  elegantly  branched,  and  as  owing  to  their  great 
length  they  are  not  well  adapted  for  pinching,  the  suspicion 
has  crossed  my  mind  that  they  may  in  addition  serve  as  an 
ornament,  like  the  horns  on  the  head  and  thorax  of  the  various 
species  above  described.  The  male  Chiaf^ognathus  Grant ii  of 
S.  Chile — a  splendid  beetle  belonging  to  the  same  family — has 

^^  'Entomological  Magazine,'  vol.  ""^  Quoted  from  Fischer,  in  '•Diet, 

i.    1833,    p.   82.     See    also    on    the  Class.  d'Hist.  Nat.'  torn.  x.  p.  324. 
conflicts  of  this  species,  Kirby  and  ^>  '  Ann.  Soc.  Entomolog.  France,' 

Spence,  ibid.  vol.  iii.  p.  314 ;    and  1866,    as    quoted    in    '  Journal    of 

Westwood,  ibid.  vol.  i.  p.  187.  Travel,'  by  A.  Murray,  1868,  p.  135. 


Chap.  :^. 


Coleoptera. 


301 


enormously  developed  mandibles  (fig.  '24) ;  he  is  bold  and  pug- 
nacious ;  when  threatened  he  faces  round,  opens  his  great  jaws, 
and  at  the  same  time  stridulates  loudly. 
But  the  mandibles  were  not  strong 
enough  to  pinch  my  finger  so  as  to 
cause  actual  pain. 

Sexual  selection,  which  implies  the 
possession  of  considerable  perceptive 
powers  and  of  strong  passions,  seems 
to  have  been  more  effective  with  the 
Lamellicorns  than  with  any  other 
family  of  beetles.  With  some  species 
the  males  are  provided  with  weapons 
for  fighting;  some  live  in  pairs  and 
show  mutual  affection ;  many  have 
the  power  of  stridulating  when  excited  ; 
many  are  furnished  with  the  most  ex- 
traordinary horns,  apparently  for  the 
sake  of  ornament;  and  some,  which 
are  diurnal  in  their  habits,  are  gor- 
geously coloured.  Lastly,  several  of 
the  largest  beetles  in  the  world  belong 
to  this  family,  which  was  placed  by 
Linnseus  and  Fabricius  at  the  head  of 
the  Order.^^ 

Stridulating  organs. — Beetles  belong- 
ing to  many  and  widely  distinct 
families  possess  these  organs.  The 
sound  thus  produced  can  sometimes 
be  heard  at  the  distance  of  several  feet 
or  even  yards,'^^  but  it  is  not  comparable 
with  that  made  by  the  Orthoptera. 
The  rasp  generally  consists  of  a  narrow, 
slightly-raised  surface,  crossed  by  very 
fine,  parallel  ribs,  sometimes  so  fine  as 
to  cause  iridescent  colours,  and  having 
a  very  elegant  appearance  under  the 
microscope.  In  some  cases,  as  Avith 
Typhosus,  minute,  bristly  or  scale-like 
prominences,  with  which  the  whole 
surrounding  surface  is  covered  in  approximately  parallel 
lines,  could  be  traced  passing  into  the  ribs  of  the  rasp.     The 


Fi°r.  24.  Chiasognathus  grantii, 
reduced.  U  ppcr  figure,  male ; 
lower  figure,  female. 


^^    Westwood, 
vol.  i.  p.  184. 


Modern    Class.' 


"  Wollaston,  '  On  certain  Musical 


Curculionidaj,'  'Annals  and  Mag.  of 
Nat.  Hist.'  vol.  vi.  18G0,  p.  14. 


302 


The  Descent  of  Man. 


Part  II. 


transition  takes  place  by  their  becoming  confluent  and  straight, 
and  at  the  same  time  more  prominent  and  smooth.  A  hard 
ridge  on  an  adjoining  part  of  the  body  serves  as  the  scraper 
for  the  rasp,  but  this  scraper  in  some  cases  has  been  specially 
modified  for  the  purpose.  It  is  rapidly  moved  across  the  rasp, 
or  conversely  the  rasp  across  the  scraper. 


Fig.  25.    Necrophorus  (from  Landois).    r.  The  two  rasps, 
the  rasp  highly  magnified. 


Left-hand  figure,  part  of 


These  organs  are  situated  in  widely  different  positions.  In 
the  carrion-beetles  (Necrophorus)  two  parallel  rasps  (r,  fig.  25) 
stand  on  the  dorsal  surface  of  the  fifth  abdominal  segment,  each 
rasp^*  consisting  of  126  to  140  fine  ribs.  These  ribs  are  scraped 
against  the  posterior  margins  of  the  elytra,  a  small  portion  of 
which  projects  beyond  the  general  outline.  In  many  Crioceridas, 
and  in  Clylhra  ^-punctata  (one  of  the  Chrysomehdse),  and  in  some 
Tenebrionidse,  &c.,'^^  the  rasp  is  seated  on  the  dorsal  apex  of  the 
abdomen,  on  the  pygidium  or  pro-pygidium,  and  is  scraped  in 
the  same  manner  by  the  elytra.  In  Heterocerus,  which  belongs 
to  another  family,  the  rasps  are  placed  on  the  sides  of  the 
first  abdominal  segment,  and  are  scraped  by  ridges  on  the 
femora.'^^    In  certain  Curculionidse  and  Carabidae,''^  the  parts 

^^  Landois,  '  Zeitschrift  fiir  wiss. 
Zoolog.'  B.  xvii.  1867,  s.  127. 

^^  I  am  greatly  indebted  to  Mr. 
G.  R.  Crotch  for  having  sent  me 
man}^  prepared  specimens  of  various 
beetles  belonging  to  these  three 
families  and  to  others,  as  well  as  for 
valuable  information.  He  believes 
that  the  power  of  stridulation  in 
the  Clythra  has  not  been  previously- 
observed.  I  am  also  much  indebted 
to  Mr.  E.  W.  Janson,  for  informa- 
tion and  specimens.  I  may  add 
that  my  son,  Mr.  F.  Darwin,  finds 
that  Dermestes  murinus  stridulates, 
but  he  searched  in  vain  for  the 
apparatus.       Scolytus     has     lately 


been  described  by  Dr.  Chapman  as 
a  stridulator,  in  the  'Entomolo- 
gist's Monthly  Magazine,'  vol.  vi.  p. 
130. 

'^  Schiodte,  translated  in  *  Annals 
and  Mag.  of  Nat.  Hist.'  vol.  xx. 
1867,  p.  37. 

^^  Westringhas  described  (Krover, 
'  Naturhist.  Tidskrift,'  B.  ii.  1848- 
49,  p.  334)  the  stridulating  organs 
in  these  two,  as  well  as  in  other 
families.  In  the  Carabidae  I  have 
examined  Elaphrus  uliginosus  and 
Blethisa  multipiinctata,  sent  to  me 
by  Mr.  Crotch.  In  Blethisa  the 
transverse  ridges  on  the  furrowed 
border  of  the  abdominal  segment  do 


Chap.  X. 


Coleoptera. 


303 


are  completely  reversed  in  position,  for  the  rasps  are  seated  on 
the  inferior  surface  of  the  elytra,  near  their  apices,  or  along 
their  outer  margins,  and  the  edges  of  the  abdominal  segments 
serve  as  the  scrapers.  In  Pelohius  Ileriminni  (one  of  Dytiscidje 
or  water-beetles)  a  strong  ridge  runs  parallel  and  near  to  the 
sutural  margin  of  the  elytra,  and  is  crossed  by  ribs,  coarse  in 
the  middle  part,  but  becoming  gradually  finer  at  both  ends, 
especially  at  the  upper  end;  when  this  insect  is  held  under 
water  or  in  the  air,  a  stridulating  noise  is  produced  by  the 
extreme  horny  margin  of  the  abdomen  being  scraped  against 
the  rasps.  In  a  great  number  of  long-horned  beetles  (Longi- 
cornia)  the  organs  are  situated  quite  otherwise,  the  rasp  being 
on  the  meso-thorax,  which  is  rubbed  against  the  pro-thorax; 
Landois  counted  238  very  fine  ribs  on  the  rasp  of  Ceramhyx 
heros. 

Many  Lamellicorns  have  the  power  of  stridulating,  and  the 
organs  differ  greatly  in  position.  Some  species  stridulate  very 
loudly,  so  that  when  Mr.  F.  Smith  caught  a  Trox  sahidosus,  a 
gamekeeper,  who  stood  by,  thought  he  had  caught  a  mouse ; 
but  I  failed  to  discover  the  proper  organs  in  ^ 

this  beetle.  In  Geotrupes  and  Typhoeus  a 
narrow  ridge  runs  obliquely  across  (r,  fig.  2G) 
the  coxa  of  each  hind-leg  (having  in  G.  ster- 
corarius  84  ribs),  which  is  scraped  by  a 
specially  projecting  part  of  one  of  the  ab- 
dominal segments.  In  the  nearly  allied  Copris 
lunaris,  an  excessively  narrow  fine  rasp  runs 
along  the  sutural  margin  of  the  elytra,  with 
another  short  rasp  near  the  basal  outer  mar- 
gin; but  in  some  other  Coprini  the  rasp  is 
seated,  according  to  Leconte,'^^  on  the  dorsal 
surface  of  the  abdomen.  In  Oryctes  it  is 
seated  on  the  pro-pygidium ;  and,  according  to 
the  same  entomologist,  in  some  other  Dynastini, 
on  the  under  surface  of  the  elytra.  Lastly, 
Westring  states  that  in  Omaloplia  hrunnea  the 
rasp  is  placed  on  the  pro-sternum,  and  the 
scraper  on  the  meta-sternum,  the  parts  thus 
occupying  the  under  surface  of  the  body,  instead  of  the  upper 
surface  as  in  the  Longicorns. 

We  thus  see  that  in  the  different  coleopterous  families  the 

not,  as  far  as  I  could  judge,  come  of  Illinois,  for  having  sent  me  ex- 

into  play  in  scraping  the  rasps  on  tracts  from  Leconte's  '  Introduction 

the  elytra.  to  Entomology,'  pp.  101,  143. 
"  1  am  indebted  to  Mr.  Walsh, 


ip.  26.  Hind-leg  of 
Gp(jtrnpcs  storcora- 
rius  (from  Landois). 

,  Rasp.  c.  Coxa.  /. 
Femur,  t.  Tibia 
tr.  Tarsi. 


304  The  Descent  of  Man.  Paet  II. 

stridulating  organs  are  wonderfully  diversified  in  position,  but 
not  much  in  structure.  Within  the  same  family  some  species 
are  provided  with  these  organs,  and  others  are  destitute  of  them. 
This  diversity  is  intelligible,  if  we  suppose  that  originally  various 
beetles  made  a  shuffiing  or  hissing  noise  by  the  rubbing  together 
of  any  hard  and  rough  parts  of  their  bodies,  which  happened  to 
be  in  contact ;  and  that  from  the  noise  thus  produced  being  in 
some  way  useful,  the  rough  surfaces  were  gradually  developed 
into  regular  stridulating  organs.  Some  beetles  as  they  move, 
now  produce,  either  intentionally  or  unintentionally,  a  shuffling 
noise,  without  possessing  any  proper  organs  for  the  purpose. 
Mr.  Wallace  infoims  me  that  the  Euchirus  longimanus  (a 
Lamellicorn,  with  the  anterior  legs  wonderfully  elongated  in  the 
male)  "  makes,  whilst  moving,  a  low  hissing  sound  by  the  pro- 
'*  trusion  and  contraction  of  the  abdomen ;  and  when  seized  it 
"  produces  a  grating  sound  by  rubbing  its  hind-legs  against  the 
"  edges  of  the  elytra."  The  hissing  sound  is  clearly  due  to  a 
narrow  rasp  running  along  the  sutural  margin  of  each  elytron  ; 
and  I  could  likewise  make  the  grating  sound  by  rubbing  the 
shagreened  surface  of  the  femur  against  the  granulated  margin 
of  the  corresponding  elytron ;  but  I  could  not  here  detect  any 
proper  rasp ;  nor  is  it  hkely  that  I  could  have  overlooked  it  in 
so  large  an  insect.  After  examining  Cychrus,  and  reading  what 
Westring  has  written  about  this  beetle,  it  seems  very  doubtful 
whether  it  possesses  any  true  rasp,  though  it  has  the  power  of 
emitting  a  sound. 

From  the  analogy  of  the  Orthoptera  and  Homoptera,  I 
expected  to  find  the  stridulating  organs  in  the  Coleoptera 
differing  according  to  sex;  but  Landois,  who  has  carefully 
examined  several  sjDecies,  observed  no  such  difference ;  nor  did 
Westring;  nor  did  Mr.  G.  E.  Crotch  in  preparing  the  many 
specimens  which  he  had  the  kindness  to  send  me.  Any  difference 
in  these  organs,  if  slight,  would,  however,  be  difficult  to  detect, 
on  account  of  their  great  variability.  Thus,  in  the  first  pair  of 
specimens  of  Necrophorus  humafor  and  of  Pelohius  which  I  ex- 
amined, the  rasp  was  considerably  larger  in  the  male  than  in 
the  female;  but  not  so  with  succeeding  specimens.  In  Geo- 
triipes  stercorarius  the  rasp  appeared  to  me  thicker,  opaquer, 
and  more  prominent  in  three  males  than  in  the  same  number  of 
females;  in  order,  therefore,  to  discover  whether  the  sexes 
differed  in  their  power  of  stridulating,  my  soa,  Mr.  F.  Darwin, 
collected  fifty-seven  living  specimens,  which  he  separated  into  two 
lots,  according  as  they  made  a  greater  or  lesser  noise,  when  held 
in  the  same  manner.  He  then  examined  all  these  specimens, 
and  found  that  the  males  were  very  nearly  in  the  same  proportion 


Chap.  X.  Coleoptera.  305 

to  the  females  in  both  the  lots.  Mr.  F.  Smith  has  kept  alive 
inimerous  specimens  of  Monoijnclms  pseudacoH  (Cnrculiomda}), 
and  is  convinced  that  both  sexes  stridulate,  and  apparently  in 

an  equal  degree.  .         ,   .  ^  ^ 

Nevertheless,  the  power  of  stridulating  is  certainly  a  sexual 
character  in  some  few  Coleoptera.  Mr.  Crotch  discovered  that 
the  males  alone  of  two  species  of  Heliopathes  (TenebrionidsB) 
possess  stridulating  organs.  I  examined  live  males  of  11.  (jlhhiis, 
and  in  all  these  there  was  a  well-developed  rasp,  partially 
divided  into  two,  on  the  dorsal  suriace  of  the  terminal  abdominal 
•sef'-ment ;  whilst  in  the  same  number  of  females  there  was  not 
even  a  rudiment  of  the  rasp,  the  membrane  of  this  segment 
being  transparent,  and  much  thinner  than  in  the  male.  In 
H  cribratostriatus  the  male  has  a  similar  rasp,  excepting  that  it 
is  not  partially  divided  into  two  portions,  and  the  female  is 
completely  destitute  of  this  organ;  the  male  in  addition  has  on 
the  apical  margins  of  the  elytra,  on  each  side  of  the  suture, 
three  or  four  short  longitudinal  ridges,  which  are  crossed  by 
extremely  fine  ribs,  parallel  to  and  resembling  those  on  the 
abdominal  rasp;  whether  these  ridges  serve  as  an  independent 
rasp,  or  as  a  scraper  for  the  abdominal  rasp,  I  could  not  decide: 
the  female  exhibits  no  trace  of  this  latter  structure. 

Ao-ain,  in  three  species  of  the  Lamellicorn  genus  Oryctes,  we 
have  a  nearly  parallel  case.  In  the  females  of  0.  gryphus  and 
nasicornis  the  ribs  on  the  rasp  of  the  pro-pygidmm  are  less 
continuous  and  less  distinct  than  in  the  males;  but  the  chief 
difference  is  that  the  whole  upper  surface  of  this  segment,  when 
held  in  the  proper  light,  is  seen  to  be  clothed  with  hairs,  which 
are  absent  or  are  represented  by  excessively  fine  down  m  the 
males.  It  should  be  noticed  that  in  all  Coleoptera  the  effective 
part  of  the  rasp  is  destitute  of  hairs.  In  0.  senegalensts  the 
difference  between  the  sexes  is  more  strongly  marked,  and  this 
is  best  seen  when  the  proper  abdominal  segment  is  cleaned  and 
viewed  as  a  transparent  object.  In  the  female  the  whole  surface 
is  covered  with  little  separate  crests,  bearing  spines ;  whilst  m 
the  male  these  crests  in  proceeding  towards  the  apex,  become 
more  and  more  confluent,  regular,  and  naked;  so  that  three- 
fourths  of  the  segment  is  covered  with  extremely  fine  parallel 
ribs  which  are  quite  absent  in  the  female.  In  the  females, 
however,  of  all  three  species  of  Oryctes,  a  slight  grating  or 
Gtridulating  sound  is  produced,  when  the  abdomen  of  a  softened 
.specimen  is  pushed  backwards  and  forwards. 

In  the  case  of  the  Heliopathes  and  Oryctes  there  can  hardly 
be  a  doubt  that  the  males  stridulate  in  order  to  call  or  to 
excite  the  females;    but  with  most  beetles  the    stridulation 


306  The  Desceftt  of  Man.  Pakt  II. 

apparently  serves  both  sexes  as  a  mntual  call.  Beetles  striclu- 
late  under  various  emotions,  in  the  same  manner  as  birds  use 
their  voices  for  many  purposes  besides  singing  to  their  mates. 
The  great  Chiasognathus  stridulates  in  anger  or  defiance ;  many 
species  do  the  same  from  distress  or  fear,  if  held  so  that  they 
cannot  escape;  by  striking  the  hollow  stems  of  trees  in  the 
Canary  Islands,  Messrs.  Wollaston  and  Crotch  were  able  to 
discover  the  presence  of  beetles  belonging  to  the  genus  Acalles 
by  their  stridulation.  Lastly,  the  male  Ateuchus  stridulates  to 
encourage  the  female  in  her  work,  and  from  distress  when  she 
is  removed.'^^  Some  naturalists  believe  that  beetles  make  this 
noise  to  frighten  away  their  enemies ;  but  I  cannot  think  that 
a  quadruped  or  bird,  able  to  devour  a  large  beetle,  would 
be  frightened  by  so  slight  a  sound.  The  belief  that  the  stridu- 
lation serves  as  a  sexual  call  is  supported  by  the  fact  that  death- 
ticks  {Anohium  tessellatuni)  are  well  known  to  answer  each 
other's  ticking,  and,  as  I  have  myself  observed,  a  tapping  noise 
artificially  made.  Mr.  Doubleday  also  informs  me  that  he 
has  sometimes  observed  a  female  ticking,^"  and  in  an  hour  or 
two  afterwards  has  found  her  united  with  a  male,  and  on  one 
occasion  surrounded  by  several  males.  Finally,  it  is  probable 
that  the  two  sexes  of  many  kinds  of  beetles  were  at  first 
enabled  to  find  each  other  by  the  slight  shuffling  noise  produced 
by  the  rubbing  together  of  the  adjoining  hard  parts  of  their 
bodies;  and  that  as  those  males  or  females  which  made  the 
greatest  noise  succeeded  best  in  finding  partners,  rugosities  on 
various  parts  of  their  bodies  were  gradually  developed  by  means 
of  sexual  selection  into  true  stridulating  organs. 

^®  M.  P.  de  la  Brulerie,  as  quoted  Landois,    '  Zeitschrift    fiir    wissen. 

in  'Journal  of  Travel,' A.  Murray,  Zoolog.'    B.    xvii.    s.    131.      Oliver 

vol.  i.  1868,  p.  135.  says    (as     quoted     by    Kirby    and 

'"  According  to   Mr.   Doubleday,  Spence,  '  Introduct.'  vol.  ii.  p.  395) 

"  the  noise   is  produced  by  ihe  in-  that  the   female   of  Pimelia  striata 

"  sect  raising  itself  on  its  legs  as  produces    a   rather    loud    sound  by 

"  high  as  it  can,  and  then  striking  striking  her  abdomen   against  any 

"  its   thorax  five   or    six    times,  in  hard  substance,  "  and  that  the  male, 

*'  rapid  succession,  against  the  sub-  "  obedient  to  this  call,  soon  attends 

"  stance  upon  which  it  is  sitting."  "  her,  and  they  pair." 
For  references   on   this  subject  see 


Chap.  XI.  Biitterjiies  and  Moths.  307 


CHAPTER  XI. 
Insects,  continued. — Order  Lepidoptera. 

(butterflies  and  moths.) 

Courtship  of  butterflies— Battles — Ticking  noise— Colours  conimon  to 
both  feexes,  or  more  brilliant  in  the  males — Examples — Not  due  to  the 
direct  action  of  the  conditions  of  life — Colours  adapted  for  protection — 
Colours  of  moths— Display — Perceptive  powers  of  the  Lepidoptera — 
Variability — Causes  of  the  difference  in  colour  between  the  males  and 
females— Mimicry,  female  butterflies  more  brilliantly  coloured  than 
the  males — Bright  colours  of  caterpillars — Summary  and  concluding 
remarks  on  the  secondary  sexual  characters  of  insects — Birds  and  insects 
compared. 

In  this  great  Order  the  most  interesting  points  for  us  are  the 
differences  in  colour  between  the  sexes  of  the  same  species,  and 
between  the  distinct  species  of  the  same  genus.  Nearly  the 
whole  of  the  following  chapter  will  be  devoted  to  this  subject ; 
but  I  will  first  make  a  few  remarks  on  one  or  two  other  points. 
Several  males  may  often  be  seen  pursuing  and  crowding  round 
the  same  female.  Their  courtship  appears  to  be  a  prolonged 
affair,  for  I  have  frequently  watched  one  or  more  males  pirouet- 
ting round  a  female  until  I  was  tired,  without  seeing  the 
end  of  the  courtship.  Mr.  A.  G.  Butler  also  informs  me  that 
he  has  several  times  watched  a  male  courting  a  female  for  a  full 
quarter  of  an  hour ;  but  she  pertinaciously  refused  bim,  and  at 
last  settled  on  the  ground  and  closed  her  wings,  so  as  to  escape 
from  his  addresses. 

Although  butterflies  are  weak  and  fragile  creatures,  they  are 
pugnacious,  and  an  Emperor  butterfly  ^  has  been  captured  with 
the  tips  of  its  wings  broken  from  a  conflict  with  another  male. 
Mr.  Collingwood,  in  speaking  of  the  frequent  battles  between  the 
butterflies  of  Borneo,  says,  "  They  whirl  round  each  other  with 
"  the  greatest  rapidity,  and  appear  to  be  incited  by  the  greatest 
"  ferocity." 

The  Ageronia  feronia  makes  a  noise  like  that  produced  by  a 
toothed  wheel  passing  under  a  spring  catch,  and  which  can  be 
heard  at  the  distance  of  several  yards :  I  noticed  this  sound  at 
Eio  de  Janeiro,  only  when  two  of  these  butterflies  were  chasing 
each  other  in  an  irregular  course,  so  that  it  is  probably  made 
during  the  courtship  of  the  sexes.^ 

1  Apatura  Iris:  'The  Entomolo-      Natura-list,'  18G8,  p.  IS.'), 
gist's  Weekly  Intelligence,'  1859,  p.  ^  gee  my  'Journal  of  Researches,' 

139.     For  the  Bornean  Butterflies,      1845,   p.    33.      Mr.  Doubleday  has 
6e«  C.  Collingwood,  '  Rambles  of  a      detected    ('  Proc.  Ent.   See'  March 


3o8  The  Descent  of  Man.  Part  II. 

Some  moths  also  produce  sounds ;  for  instance,  the  males  of 
Thecophora  fovea.  On  two  occasions  Mr.  F.  Buchanan  White  ^ 
heard  a  sharp  quick  noise  made  by  the  male  of  Hylophila 
jjrasinana,  and  which  he  believes  to  be  produced,  as  in  Cicada, 
by  an  elastic  membrane,  furnished  with  a  muscle.  He  quotes, 
also,  Guenee,  that  Setina  produces  a  sound  like  the  ticking 
of  a  watch,  apparently  by  the  aid  of  "  two  large  tympaniform 
"  vesicles,  situated  in  the  pectoral  region ; "  and  these  "  are  much 
"  more  developed  in  the  male  than  in  the  female."  Hence  the 
sound-producing  organs  in  the  Lepidoptera  appear  to  stand  in 
some  relation  with  the  sexual  functions.  I  have  not  alluded 
to  the  well-known  noise  made  by  the  Death's  Head  Sphinx,  for 
it  is  generally  heard  soon  after  the  moth  has  emerged  from 
its  cocoon. 

Girard  has  always  observed  that  the  musky  odour,  which  is 
emitted  by  two  species  of  Sphinx  moths,  is  peculiar  to  the  males  ;^ 
and  in  the  higher  classes  we  shall  meet  with  many  instances  of 
the  males  alone  being  odoriferous. 

Every  one  must  have  admired  the  extreme  beauty  of  many 
butterflies  and  of  some  moths ;  and  it  may  be  asked,  are  their 
colours  and  diversified  patterns  the  result  of  the  direct  action  of 
the  i)hysical  conditions  to  which  these  insects  have  been  exposed, 
without  any  benefit  being  thus  derived  ?  Or  have  successive 
variations  been  accumulated  and  determined  as  a  protection, 
or  for  some  unknown  purpose,  or  that  one  sex  may  be  at- 
tractive to  the  other  ?  And,  again,  what  is  the  meaning  of  the 
colours  being  widely  different  in  the  males  and  females  of 
certain  siDccies,  and  alike  in  the  two  sexes  of  other  species  of  the 
same  genus  ?  Before  attempting  to  answer  these  questions  a 
body  of  facts  must  be  given. 

With  our  beautiful  English  butterflies,  the  admiral,  peacock, 
and  painted  lady  (Vanessse),  as  well  as  many  others,  the  sexes 
are  alike.  This  is  also  the  case  with  the  magniflcent  Heliconidae, 
and  most  of  the  Danaidse  in  the  tropics.  But  in  certain  other 
tropical  groups,  and  in  some  of  our  English  butterflies,  as  the 
purple  emperor,  orange-tip,  &c.  {Apatura  Iris  and  Anthocharis 
cardamines),  the  sexes  differ  either  greatly  or  slightly  in  colour. 
No  language  suffices  to  describe  the  splendour  of  the  males  of 


3rd,  1845,  p.  123)  a  peculiar  mem-  observations,  *  The  Scottish  Natural- 

branous    sac    at    the    base    of    the  ist,*  July  1872,  p.  214. 

front  wings,  which  is  probably  con-  ^  '  The  Scottish  Naturalist,'  July 

nected  with  the  production  of  the  1872,  p.  213. 

sound.    For  the  case  of  Thecophora,  *   '  Zoological    Record,'    1869,   p. 

see    'Zoological    Record,'    1869,    p.  347. 

401.     For    Mr.    Buchanan   White's 


Chap.  XL  Butterflies  and  Moths.  309 

some  tropical  species.  Even  within  the  same  genus  Ave  often 
find  species  presenting  extraordinary  differences  between  the 
sexes,  whilst  others  have  their  sexes  closely  alike.  Thns  in  the 
South  American  genus  Epicalia,  Mr.  Bates,  to  whom  I  am 
indebted  for  most  of  the  following  facts,  and  for  looking  over 
this  whole  discussion,  informs  me  that  he  knows  twelve  species, 
the  two  sexes  of  which  haunt  the  same  stations  (and  this  is  not 
always  the  case  with  butterflies),  and  which,  therefore,  cannot 
have  been  differently  affected  by  external  conditions.^  In  nine  of 
these  twelve  species  the  males  rank  amongst  tlie  most  brilliant  of 
all  butterflies,  and  differ  so  greatly  from  the  comparatively  plain 
females  that  they  were  formerly  placed  in  distinct  genera.  The 
females  of  these  nine  species  resemble  each  other  in  their  general 
type  of  coloration;  and  they  likewise  resemble  both  sexes  of 
the  species  in  several  alhed  genera,  found  in  various  parts  of 
the  world.  Hence  we  may  infer  that  these  nine  species,  and 
probably  all  the  others  of  the  genus,  are  descended  from  an 
ancestral  form  which  was  coloured  in  nearly  the  same  manner. 
In  the  tenth  species  the  female  still  retains  the  same  general 
colouring,  but  the  male  resembles  her,  so  that  he  is  coloured  in 
a  much  less  gaudy  and  contrasted  manner  than  the  males  of  the 
previous  species.  In  the  eleventh  and  twelfth  species,  the 
females  depart  from  the  usual  type,  for  they  are  gaily  decorated 
almost  like  the  males,  but  in  a  somewhat  less  degree.  Hence  in 
these  two  latter  species  the  bright  colours  of  the  males  seem  to 
have  been  transferred  to  the  females ;  whilst  in  the  tenth 
species  the  male  has  either  retained  or  recovered  the  plain 
colours  of  the  female,  as  well  as  of  the  parent-form  of  the  genus. 
The  sexes  in  these  three  cases  have  thus  been  rendered  nearly 
alike,  though  in  an  opposite  manner.  In  the  allied  genus  Eubagis, 
both  sexes  of  some  of  the  species  are  plain-coloured  and  nearly 
alike ;  whilst  with  the  greater  number  the  males  are  decorated 
with  beautiful  metallic  tints  in  a  diversified  manner,  and  differ 
much  from  their  females.  The  females  throughout  the  genus 
retain  the  same  general  style  of  colouring,  so  that  they  resemble 
one  another  much  more  closely  than  they  resemble  their  own 
males. 

In  the  genus  Papilio,  all  the  species  of  the  ^neas  group 
are  remarkable  for  their  conspicuous  and  strongly  contrasted 
colours,  and  they  illustrate  the  frequent  tendency  to  gradation 
in  the  amount  of  difference  between  the  sexes.  In  a  few  species, 
for  instance  in  P.  ascauius,  the  males  and  females  are  alike ;  in 

*  See  also  Mr.  Bates's  paper  in  the  same  subject,  in  regard  to 
♦  Proc.  Ent.  Soc.  of  Philadelphia,'  Diadema,  in  '  transact.  Entomolog. 
1865,  p.  206.     Also  Mr.  Wallace  on      Soc.  of  London,'  1869,  p.  278. 


3IO  The  Descent  of  Mail.  Part  II. 

others  the  males  are  either  a  little  brighter,  or  very  much  more 
superb  than  the  females.  The  genus  Juuouia,  allied  to  our 
Vaness?e,  offers  a  nearly  parallel  case,  for  although  the  sexes  of 
most  of  the  species  resemble  each  other,  and  are  destitute  of 
rich  colours,  yet  in  certain  species,  as  in  J.  cenone,  the  male  is 
rather  more  bright- coloured  than  the  female,  and  in  a  few  (for 
instance  J.  andremiaja)  the  male  is  so  different  from  the  female 
that  he  might  be  mistaken  for  an  entirely  distinct  species. 

Another  striking  case  was  pointed  out  to  me  in  the  British 
Museum  by  Mr.  A.  Butler,  namely,  one  of  the  tropical  American 
Theclse,  in  which  both  sexes  are  nearly  alike  and  wonderfully 
splendid ;  in  another  species  the  male  is  coloured  in  a  similarly 
gorgeous  manner,  whilst  the  whole  upper  surface  of  the 
female  is  of  a  dull  uniform  brown.  Our  common  little  English 
blue  butterflies  of  the  genus  Lyc^na,  illustrate  the  various  dif- 
ferences in  colour  between  the  sexes,  almost  as  well,  though  not 
in  so  striking  a  manner,  as  the  above  exotic  genera.  In  Lycxna 
agestis  both  sexes  have  wings  of  a  brown  colour,  bordered  with 
small  ocellated  orange  spots,  and  are  thus  alike.  In  L.  csgon 
the  wings  of  the  male  are  of  a  fine  blue,  bordered  with  black  ; 
whilst  those  of  the  female  are  brown,  with  a  similar  border, 
closely  resembling  the  wings  of  X.  agestis.  Lastly,  in  L.  arion  both 
sexes  are  of  a  blue  colour  and  are  very  like,  though  in  the  female 
the  edges  of  the  wings  are  rather  duskier,  with  the  black  spots 
plainer ;  and  in  a  bright  blue  Indian  sj^ecies  both  sexes  are  still 
more  alike. 

I  have  given  the  foregoing  details  in  order  to  show,  in  the  first 
place,  that  when  the  sexes  of  butterflies  differ,  the  male  as  a 
general  rule  is  the  more  beautiful,  and  departs  more  from  the 
usual  type  of  colouring  of  the  group  to  which  the  species 
belongs.  Hence  in  most  groups  the  females  of  the  several 
species  resemble  each  other  much  more  closely  than  do  the 
males.  In  some  cases,  however,  to  which  I  shall  hereafter 
allude,  the  females  are  coloured  more  splendidly  than  the 
males.  In  the  second  place,  these  details  have  been  given  to 
bring  clearly  before  the  mind  that  within  the  same  genus,  the 
two  sexes  frequently  present  every  gradation  from  no  difference 
in  colour,  to  so  great  a  difference  that  it  was  long  before  the  two 
were  placed  by  entomologists  in  the  same  genus.  In  the  third 
l^lace,  we  have  seen  that  when  the  sexes  nearly  resemble  each 
other,  this  appears  due  either  to  the  male  having  transferred 
his  colours  to  the  female,  or  to  the  male  having  retained,  or 
perhaps  recovered,  the  jr-rimordial  colours  of  the  group.  It  also 
deserves  notice  that  in  those  groups  in  which  the  sexes  differ, 
the  females  usually  somewhat  resemble  the  males,  so  that  when 


Chap.  XI.  Btittei'Jlies  and  Aloths.  311 

the  males  cire  beautiful  to  an  extraordinary  degree,  the  females 
almost  invariably  exhibit  some  degree  of  beauty.  From  the 
many  cases  of  gradation  in  the  amount  of  difference  between 
the  sexes,  and  from  the  prevalence  of  the  same  general  type  of 
coloration  throughout  the  whole  of  the  same  group,  we  may  con- 
clude that  the  causes  have  generally  been  the  same  which  have 
determined  the  brilliant  colouring  of  the  males  alone  of  some 
species,  and  of  both  sexes  of  other  species. 

As  so  many  gorgeous  butterflies  inhabit  the  tropics,  it  has 
often  been  supposed  that  they  owe  their  colours  to  the  great 
heat  and  moisture  of  these  zones  ;  but  Mr.  Bates  °  has  shewn  by 
the  comparison  of  various  closely-allied  groups  of  insects  from 
the  temperate  and  tropical  regions,  that  this  view  cannot  be 
maintained ;  and  the  evidence  becomes  conclusive  wlien  bril- 
liantly-coloured males  and  plain-coloured  females  of  the  same 
species  inhabit  the  same  district,  feed  on  the  same  food,  and 
follow  exactly  the  same  habits  of  life.  Even  when  the  sexes 
resemble  each  other,  we  can  hardly  believe  that  their  brilliant 
and  beautifully-arranged  colours  are  the  purposeless  result  of 
the  nature  of  the  tissues  and  of  the  action  of  the  surrounding 
conditions. 

With  animals  of  all  kinds,  whenever  colour  has  been  modified 
for  some  special  purpose,  this  has  been,  as  far  as  we  can  judge, 
either  for  direct  or  indirect  protection,  or  as  an  attraction  between 
the  sexes.  With  many  species  of  butterflies  the  upper  surfaces 
of  the  wings  are  obscure;  and  this  in  all  probability  leads 
to  their  escaping  observation  and  danger.  But  butterflies 
would  be  particularly  liable  to  be  attacked  by  their  enemies 
when  at  rest ;  and  most  kinds  whilst  resting  raise  their  wings 
vertically  over  their  backs,  so  that  the  lower  surface  alone  is 
exposed  to  view.  Hence  it  is  this  side  which  is  often  coloured 
so  as  to  imitate  the  objects  on  which  these  insects  commonly 
rest.  Dr.  Bossier,  I  believe,  first  noticed  the  similarity  of  the 
closed  wings  of  certain  Vanesste  and  other  butterflies  to  the 
bark  of  trees.  Many  analogous  and  striking  facts  could  be 
given.  The  most  interesting  one  is  that  recorded  by  Mr. 
Wallace ''  of  a  common  Indian  and  Sumatran  butterfly  (Kallima), 
which  disappears  like  magic  when  it  settles  on  a  bush ;  for  it 
hides  its  head  and  antennae  between  its  closed  wings,  which, 
in  form,  colour  and  veining,  cannot  be  distinguished  from  a 
withered  leaf  with  its  footstalk.     In  some  other  cases  the  lower 

«  'The  Naturalist    on  the  Ama-  18G7,    p.    10.      A    woodcut    of  the 

zons,'  vol.  i.  1863,  p.  19.  Kallima  is  given  by  Mr.  Wallace  in 

^  See    the   interesting  article    in  '  Hardwicke's  Science  Gossip,'  Sept. 

the    '  Westminster    Review,*     July  18G7,  p.  19G. 


312  The  Descent  of  Man.  Part  11. 

surfaces  of  the  wings  are  brilliantly  coloured,  and  yet  are 
protective  ;  thus  in  Theda  ruhi  the  Avings  when  closed  are  of  an 
emerald  green,  and  resemble  the  young  leaves  of  the  bramble, 
on  which  in  spring  this  butterfly  may  often  be  seen  seated.  It  is 
also  remarkable  that  in  very  many  species  in  which  the  sexes 
differ  greatly  in  colour  on  their  upper  surface,  the  lower  surface 
is  closely  similar  or  identical  in  both  sexes,  and  serves  as  a 
IDrotection.^ 

Although  the  obscure  tints  both  of  the  up])er  and  under 
sides  of  many  butterflies  no  doubt  serve  to  conceal  them,  yet  we 
cannot  extend  this  view  to  the  brilliant  and  conspicuous  colours 
on  the  upper  surface  of  such  species  as  our  admiral  and  peacock 
Yanessge,  our  white  cabbage-butterflies  (Pieris),  or  the  great 
swallow-tail  Papilio  which  haunts  the  open  fens — for  these 
butterflies  are  thus  rendered  visible  to  every  living  creature. 
In  these  species  both  sexes  are  alike  ;  but  in  the  common  brim- 
stone butterfly  (Gonepteryx  rhamni),  the  male  is  of  an  intense 
yellow,  whilst  the  female  is  much  paler;  and  in  the  orange- 
tip  (Anthocharis  cardamines)  the  males  alone  have  their  wings 
tipped  with  bright  orange.  Both  the  males  and  females  in 
these  cases  are  conspicuous,  and  it  is  not  credible  that  their 
difference  in  colour  should  stand  in  any  relation  to  ordinary 
protection.  Prof.  Weismann  remarks,^  that  the  female  of  one  of 
the  Lycsense  expands  her  brown  wings  when  she  settles  on 
the  ground,  and  is  then  almost  invisible ;  the  male,  on  the  other 
hand,  as  if  aware  of  the  danger  incurred  from  the  bright  blue  of 
the  upper  surface  of  his  wings,  rests  with  them  closed ;  and  this 
shews  that  the  blue  colour  cannot  be  in  any  way  protective. 
Nevertheless,  it  is  probable  that  conspicuous  colours  are  in- 
directly beneficial  to  many  species,  as  a  warning  that  they  are 
unpalatable.  For  in  certain  other  cases,  beauty  has  been  gained 
through  the  imitation  of  other  beautiful  species,  which  inhabit 
the  same  district  and  enjoy  an  immunity  from  attack  by  being 
in  some  way  offensive  to  their  enemies ;  but  then  we  have  to 
account  for  the  beauty  of  the  imitated  species. 

As  Mr.  Walsh  has  remarked  to  me,  the  females  of  our  orange- 
tip  butterfly,  above  referred  to,  and  of  an  American  species 
{Anth.  genutia)  probably  shew  us  the  primordial  colours  of  the 
parent-species  of  the  genus;  for  both  sexes  of  four  or  five 
widely-distributed  species  are  coloured  in  nearly  the  same 
manner.  As  in  several  previous  cases,  we  may  here  infer  that 
it  is  the  males  of  Anth.  cai'damines  and  genutia  which  have 
departed  from  the  usual  type  of  the  genus.    In  the  Anth.  sara 

8  Mr.    G.    Fraser,    in    '  Nature,'  »  '  Einfluss  der  Isolirung  auf  die 

April  1871,  p.  489.  Artbildung,'  1872,  p.  58. 


Chap.  XI.  BiUterflies  and  Moths.  313 

from  California,  the  orange-tips  to  the  wings  have  been  partially 
developed  in  the  female  ;  but  they  are  paler  than  in  the  male,  and 
slightly  different  in  some  other  respects.  In  an  allied  Indian 
form,  the  Iphias  glaucippe,  the  orange-tips  are  fully  developed  in 
both  sexes.  In  this  Iphias,  as  pointed  out  to  me  by  Mr.  A.  Butler, 
the  under  surface  of  the  wings  marvellously  resembles  a  pale- 
coloured  leaf;  and  in  our  English  orange-tip,  the  under  surface 
resembles  the  flower-head  of  the  wild  parsley,  on  which  the 
butterfly  often  rests  at  night.^*^  The  same  reason  which  compels 
us  to  believe  that  the  lower  surfaces  have  here  been  coloured  for 
the  sake  of  protection,  leads  us  to  deny  that  the  wings  have 
been  tipped  with  bright  orange  for  the  same  purpose,  especially 
when  this  character  is  confined  to  the  males. 

Most  Moths  rest  motionless  during  the  whole  or  gi-eater  pai-t 
of  the  day  with  their  wiDgs  depressed ;  and  the  whole  upper 
surface  is  often  shaded  and  coloured  in  an  admirable  manner,  as 
Mr.  Wallace  has  remarked,  for  escaping  detection.  The  front- 
wings  of  the  Bombycidae  and  Noctuidse,^^  when  at  rest,  generally 
overlap  and  conceal  the  hind- wings ;  so  that  the  latter  might  be 
brightly  coloured  without  much  risk;  and  they  are  in  fact 
often  thus  coloured.  During  flight,  moths  would  often  be  able 
to  escape  from  their  enemies ;  nevertheless,  as  the  hind-wings 
are  then  fully  exposed  to  view,  their  bright  colours  must 
generally  have  been  acquired  at  some  little  risk.  But  the 
following  fact  shews  how  cautious  we  ought  to  be  in  drawing 
conclusions  on  this  head.  The  common  Yellow  Under- wings 
(Triphsena)  often  fly  about  during  the  day  or  early  evening,  and 
are  then  conspicuous  from  the  colour  of  their  hind-wings.  It 
would  naturally  be  thought  that  this  would  be  a  source  of 
danger ;  but  Mr.  J.  Jenner  Weir  believes  that  it  actually  serves 
them  as  a  means  of  escape,  for  birds  strike  at  these  brightly 
coloured  and  fragile  surfaces,  instead  of  at  the  body.  For  in- 
stance, Mr.  Weir  turned  into  his  aviary  a  vigorous  specimen  of 
Triphcena  pronuba,  which  was  instantly  pursued  by  a  robin; 
but  the  bird's  attention  being  caught  by  the  coloured  wings,  the 
moth  was  not  captured  until  after  about  fifty  attempts,  and 
small  portions  of  the  wings  were  repeatedly  broken  off.  He  tried 
the  same  experiment,  in  the  open  air,  with  a  swallow  and  T. 
fimbria',  but  the  large  size  of  this  moth  probably  interfered 
with  its  capture.^^    We  are  thus  reminded  of  a  statement  made 

'0  See  the  interesting  observatious  Science  Gossip,'  Sept.  18G7,  p.  193. 
hy  Mr.  T.  W.  Wood,  '  The  Student,'  i^  g^g  j^jgo,  on  this  subject,  Mr. 

Sept.  1868,  p.  81.  Weir's  paper  in  'Transact.  Eut.  Soc' 

•1  Mr.  Walhice    in   '  Hardwicke's  1869,  p.  23. 


314  The  Descent  of  Man.  Part  II. 

by  Mr.  Wallace,^^  namely,  that  in  the  Brazihan  forests  and 
Malayan  islands,  many  common  and  highly-decorated  butterflies 
are  weak  flyers,  though  furnished  with  a  broad  expanse  of  wing ; 
and  they  "  are  often  captured  with  pierced  and  broken  wings, 
"  as  if  they  had  been  seized  by  birds,  from  which  they  had 
"  escaped :  if  the  wings  had  been  much  smaller  in  proportion 
"  to  the  body,  it  seems  probable  that  the  insect  would  more 
"  frequently  have  been  struck  or  pierced  in  a  vital  part,  and 
"  thus  the  increased  expanse  of  the  wings  may  have  been  in- 
"  directly  beneficial." 

Display. — The  bright  colours  of  many  butterflies  and  of  some 
moths  are  specially  arranged  for  display,  so  that  they  may  be 
readily  seen.  During  the  night  colours  are  not  visible,  and 
there  can  be  no  doubt  that  the  nocturnal  moths,  taken  as  a 
body,  are  much  less  gaily  decorated  than  butterflies,  all  of 
which  are  diurnal  in  their  habits.  But  the  moths  of  certain 
families,  such  as  the  Zygsenidas,  several  Sphingidse,  Uraniidai 
some  Arctiidse  and  Saturniidse,  fly  about  during  the  day  or 
early  evening,  and  many  of  these  are  extremely  beautiful,  being 
far  brighter  coloured  than  the  strictly  nocturnal  kinds.  A 
few  exceptional  cases,  however,  of  bright-coloured  nocturnal 
species  have  been  recorded.^* 

There  is  evidence  of  another  kind  in  regard  to  display. 
Butterflies,  as  before  remarked,  elevate  their  wings  when  at 
rest,  but  whilst  basking  in  the  sunshine  often  alternately  raise 
and  depress  them,  thus  exposing  both  surfaces  to  full  view ;  and 
although  the  lower  surface  is  often  coloured  in  an  obscure 
manner  as  a  protection,  yet  in  many  species  it  is  as  highly 
decorated  as  the  upper  surface,  and  sometimes  in  a  very 
different  manner.  In  some  tropical  species  the  lower  surface  is 
even  more  brilliantly  coloured  than  the  upper.  ^^  In  the  Eng- 
lish fritillaries  (Argynnis)  the  lower  surface  alone  is  orna- 
mented with  shining  silver.  Nevertheless,  as  a  general  rule, 
the  upjxir  surface,  which  is  probably  more  fully  exposed,  is 
coloured  more  brightly  and  diversely  than  the  lower.  Hence 
the  lower  surface  generally  affords  to  entomologists  the  more 

13  <  Westminster  Review,'  July  Insects  of  New  England,' 184-2,  p.  315. 
1867,  p.  16.  ^^  Such  differences    between   the 

^*  For    instance,    Lithosia ;    but  upper    and    lower    surfoces    of    the 

Prof.  Westwood  ('  Modern  Class,  of  wings  of  several  species  of  Papilio, 

Insects,'  vol.  ii.  p.  390)  seems   sur-  may  be  seen  in  the  beautiful  plates 

prised  at  this  case.    On  the  relative  to    Mr.  Wallace's    '  Memoir  on  the 

colours    of    diurnal   and   nocturnal  Papilionidae  of  the  Malayan  Region," 

Lepidoptera,  see  ibid.  pp.  333    and  in  'Transact.   Linn.  Soc'  vol.  xxv. 

392;  also  Harris,  'Treatise  on  the  part  i.  1865. 


Chap.  XL  Butterflies  and  Moths.  3 1 5 

useful  character  for  detecting  the  affinities  of  the  various 
species,  Fritz  Mliller  informs  me  that  three  species  of  Castnia 
are  found  near  his  house  in  S.  Brazil :  of  two  of  them  the  hind- 
wings  are  obscure,  and  are  always  covered  by  the  front- wings 
when  these  butterflies  are  at  rest ;  but  the.  third  species  has 
black  hind-wings,  beautifully  spotted  with  red  and  white,  and 
these  are  fully  expanded  and  displayed  whenever  the  butterfly 
rests.     Other  such  cases  could  be  added. 

If  we  now  turn  to  the  enormous  group  of  moths,  which,  as 
I  hear  from  Mr.  Stainton,  do  not  habitually  expose  the  under 
surface  of  their  wings  to  full  view,  we  find  this  side  very  rarely 
coloured  with  a  brightness  greater  than,  or  even  equal  to,  that 
of  the  upper  side.  Some  exceptions  to  the  rule,  either  real  or 
apparent,  must  be  noticed,  as  the  case  of  Hypopyra.^^  Mr. 
Trimen  informs  me  that  in  Guenee's  great  work,  three  moths 
are  figured,  in  which  the  under  surface  is  much  the  more 
brilliant.  For  instance,  in  the  Australian  Gastrophora  the 
upper  surface  of  the  fore-wing  is  pale  greyish-ochreous,  while 
the  lower  surface  is  magnificently  ornamented  by  an  ocellus  of 
cobalt-blue,  placed  in  the  midst  of  a  black  mark,  surrounded 
by  orange-yellow,  and  this  by  bluish-wliite.  But  the  habits  of 
these  three  moths  are  unknown ;  so  that  no  explanation  can  be 
given  of  their  unusual  style  of  colouring.  Mr.  Trimen  also 
informs  me  that  the  lower  surface  of  the  wings  in  certain  other 
Geometrse"  and  quadrifid  Noctuse  are  either  more  variegated 
or  more  brightly-coloured  than  the  upper  surface ;  but  some  of 
these  species  have  the  habit  of  "  holding  their  wings  quite  erect 
"  over  their  backs,  retaining  them  in  this  position  for  a  con- 
"  siderable  time,"  and  thus  exposing  the  under  surface  to  view. 
Other  species,  when  settled  on  the  ground  or  herbage,  now  and 
then  suddenly  and  slightly  lift  up  their  wings.  Hence  the  lower 
surface  of  the  wings  being  brighter  than  the  upper  surface 
in  certain  moths  is  not  so  anomalous  as  it  at  first  appears. 
The  Saturniidae  include  some  of  the  most  beautiful  of  all 
moths,  their  wings  being  decorated,  as  in  our  British  Emperor 
moth,  with  fine  ocelli ;  and  Mr.  T.  W.  "VVood^*  observes  that 
they  resemble  butterflies  in  some  of  their  movements;  "for 
"  instance,  in  the  gentle  waving  up  and  down  of  the  wings  as  if 
"  for  display,  which  is  more  characteristic  of  diurnal  tlian  of 
**  nocturnal  Lepidoptera." 

"    See    Mr.    Wormald    on    this  the    Geometra})    in  'Transact.  Eut. 

moth  :  '  Proc.  Ent.  Soc'  March  2nd,  Soc'  new  series,  vol.  v.  pi.  xv.  and 

1868.  xvi. 

1'  See  also  an  account  of  the  S.  ^^  '  Proc.    Ent.  Soc.    of  London,' 

American    genus    Erateina   (one    of  July  6,  18(38,  p.  xxvii. 


3i6  T lie  Descent  of  Man.  Part  II. 

It  is  a  singular  fact  that  no  British  moths  which  are  bril- 
liantly coloured,  and,  as  far  as  I  can  discover,  hardly  any  foreign 
species,  differ  much  in  colour  according  to  sex ;  though  this  is 
the  case  with  many  brilliant  butterflies.  The  male,  however,  of 
one  American  moth,  the  Saturnia  lo,  is  described  as  having  its 
fore-wings  deep  yellow,  curiously  marked  with  purplish-red 
spots  ;  whilst  the  wings  of  the  female  are  purple-brown,  marked 
with  grey  lines.^^  The  British  moths  which  differ  sexually  in 
colour  are  all  brown,  or  of  various  dull  yellow  tints,  or  nearly 
white.  In  several  species  the  males  are  much  darker  than  the 
females,^*^  and  these  belong  to  grouj^s  which  generally  fly  about 
during  the  afternoon.  On  the  other  hand,  in  many  genera,  as 
Mr.  Stainton  informs  me,  the  males  have  the  hind- wings  whiter 
than  those  of  the  female — of  which  fact  Agrotis  exdamationis 
offers  a  good  instance.  In  the  Ghost  Moth  {Hepialus  humuli) 
the  difference  is  more  strongly  marked ;  the  males  being  white, 
and  the  females  yellow  with  darker  markings.^^  It  is  probable 
that  in  these  cases  the  males  are  thus  rendered  more  conspicuous, 
and  more  easily  seen  by  the  females  whilst  flying  about  in 
the  dusk. 

From  the  several  foregoing  facts  it  is  impossible  to  admit 
that  the  brilliant  colours  of  butterflies,  and  of  some  few  moths, 
have  commonly  been  acquired  for  the  sake  of  protection.  We 
have  seen  that  their  colours  and  elegant  patterns  are  arranged 
and  exhibited  as  if  for  display.  Hence  I  am  led  to  believe  that 
the  females  prefer  or  are  most  excited  by  the  more  brilliant 

^^  Harris,  '  Treatise,'  <S:c.,  edited  which  were    fond    of  eating    other 

by  Flint,  1862,  p.  395.  moths ;    so  that  if  the  Cycnia  was 

2°  For  instance,  I  observe  in  my  commonly  mistaken  by  British  birds 

son's    cabinet    that    the   males    are  for  the  Spilosoma,  it  would  escape 

darker    than    the    females    in    the  being  devoured,  and  its  white   de- 

Lasiocampa  quercus,  Odonestis  pota-  ceptive  colour  would  thus  be  highly 

toria,  Hypogymnt  dispar,  Dasychira  beneficial. 

pudibunda,  and  Cycnia  mendica.  In  ^^  It  is  remarkable,  that  in  the 
this  latter  species  the  difference  in  Shetland  Islands  the  male  of  this 
colour  between  the  two  sexes  is  moth,  instead  of  differing  widely 
strongly  marked ;  and  Mr.  Wallace  from  the  female,  frequently  re- 
informs  me  that  we  here  have,  as  sembles  her  closely  in  colour  (see 
he  believes,  an  instance  of  protective  Mr.  MacLachlan,  '  Transact.  Ent. 
mimicry  confined  to  one  sex,  as  Soc'  vol.  ii.  1866,  p.  459).  Mr. 
will  hereafter  be  more  fully  ex-  G.  Fraser  suggests  ('Nature,'  April 
plained.  The  white  female  of  the  1871,  p.  489)  that  at  the  season  of 
Cycnia  resembles  the  very  common  the  year  when  the  ghost-moth  ap- 
Spilosoma  menthrasti,  both  sexes  of  pears  in  these  northern  islands,  the 
which  are  white ;  and  Mr.  Stainton  whiteness  of  the  males  would  not 
observed  that  this  latter  moth  was  be  needed  to  render  them  visible  tc 
rejected  with  utter  disgust  by  a  the  females  in  the  twilight  night, 
whole     brood    of    young    turkeys, 


Chap.  XL  Biittcrjiies  and  MotJis.  317 

males ;  for  on  any  other  supposition  the  males  would,  as  far  as 
sve  can  see,  be  ornamented  to  no  purpose.  We  know  that  ants 
and  certain  Lamellicorn  beetles  are  capable  of  feeling  an  attach- 
ment for  each  other,  and  that  ants  recognise  their  tellows  after 
an  interval  of  several  months.  Hence  there  is  no  abstract 
improbability  in  the  Lepidoptcra,  which  probably  stand  nearly 
or  quite  as  high  in  the  scale  as  these  insects,  having  sufficient 
mental  capacity  to  admire  bright  colours.  They  certainly 
discover  flowers  by  colour.  The  Humming-bird  Sphinx  may 
often  be  seen  to  swoop  down  from  a  distance  on  a  bunch  of 
flowers  in  the  midst  of  green  foliage ;  and  I  have  been  assured 
by  two  persons  abro.ad,  that  these  moths  repeatedly  visit  flowers 
painted  on  the  walls  of  a  room,  and  vainly  endeavour  to  insert 
their  proboscis  into  them.  Fritz  Miiller  informs  me  that  several 
kinds  of  butterflies  in  S.  Brazil  shew  an  unmistakable  prefer- 
ence for  certain  colours  over  others :  he  observed  that  they 
very  often  visited  the  brilliant  red  flowers  of  five  or  six  genera  of 
plants,  but  never  the  white  or  yellow  flowering  species  of  the 
same  and  other  genera,  growing  in  the  same  garden ;  and  I 
have  received  other  accounts  to  the  same  efl"ect.  As  I  hear 
from  Mr.  Doubleday,  the  common  white  butterfly  often  flies 
down  to  a  bit  of  paper  on  the  ground,  no  doubt  mistaking  it 
for  one  of  its  own  species.  Mr.  Collingwood^^  in  speaking  of 
the  difficulty  in  collecting  certain  butterflies  in  the  Malay 
Archipelago,  states  that  "  a  dead  specimen  pinned  upon  a 
"  conspicuous  twig  will  often  arrest  an  insect  of  the  same  species 
"  in  its  headlong  flight,  and  bring  it  down  within  easy  reach  of 
*'  the  net,  especially  if  it  be  of  the  opposite  sex." 

The  courtship  of  butterflies  is,  as  before  remarked,  a  prolonged 
affair.  The  males  sometimes  fight  together  in  rivalry;  and 
many  may  be  seen  pursuing  or  crowding  round  the  same 
female.  Unless,  then,  the  females  prefer  one  male  to  another, 
the  pairing  must  be  left  to  mere  chance,  and  this  does  not 
appear  probable.  If,  on  the  other  hand,  the  females  habitually, 
or  even  occasionally,  prefer  the  more  beautiful  males,  the  colours 
of  the  latter  will  have  been  rendered  brighter  by  degrees,  and 
will  have  been  transmitted  to  both  sexes  or  to  one  sex,  according 
to  the  law  of  inheritance  which  has  prevailed.  The  process  of 
sexual  selection  will  have  been  much  facilitated,  if  the  conclusion 
can  be  trusted,  arrived  at  from  various  kinds  of  evidence  in  the 
supplement  to  the  ninth  chapter;  namely,  that  the  males  of 
many  Lepidoptera,  at  least  in  the  imago  state,  greatly  exceed 
the  females  in  number. 

Some  facts,  however,  are  opposed  to  the  belief  that  female 

"  'Rambles  of  a  Naturalist  in  the  Chinese  Seas,'  1868,  p.  182. 


3 1 8  The  Descent  of  Man.  Part  II. 


butterflies  prefer  tlio  more  beautiful  males ;  thus,  as  1  have 
been  assured  by  several  collectors,  fresh  females  may  frequently 
be  seen  paired  with  battered,  faded,  or  dingy  males ;  but  this  is 
a  circumstance  which  could  hardly  fail  often  to  follow  from  the 
males  emerging  from  their  cocoons  earlier  than  the  females. 
With  moths  of  the  family  of  the  Bombycidae,  the  sexes  pair 
immediately  after  assuming  the  imago  state;  for  they  cannot 
feed,  owing  to  the  rudimentary  condition  of  their  mouths.  The 
females,  as  several  entomologists  have  remarked  to  me,  lie  in  an 
almost  torpid  state,  and  appear  not  to  evince  the  least  choice  in 
regard  to  their  partners.  This  is  the  case  with  the  common 
silk-moth  (^B.  morl),  as  I  have  been  told  by  some  continental 
and  English  breeders.  Dr.  Wallace,  who  has  had  great 
experience  in  breeding  Bomhyx  cynthia,  is  convinced  that  the 
females  evince  no  choice  or  preference.  He  has  kept  above 
300  of  these  moths  together,  and  has  often  found  the 
most  vigorous  females  mated  with  stunted  males.  The  reverse 
appears  to  occur  seldom ;  for,  as  he  believes,  the  more  vigorous 
males  pass  over  the  weakly  females,  and  are  attracted  by  those 
endowed  with  most  vitality.  Nevertheless,  the  Bombycidse, 
though  obscurely-coloured,  are  often  beautiful  to  our  eyes  from 
their  elegant  and  mottled  shades. 

I  have  as  yet  only  referred  to  the  species  in  which  the  males 
are  brighter  coloured  than  the  females,  and  I  have  attributed 
their  beauty  to  the  females  for  many  generations  having  chosen 
and  paired  with  the  more  attractive  males.  But  converse 
cases  occur,  though  rarely,  in  which  the  females  are  more 
brilliant  than  the  males ;  and  here,  as  I  believe,  the  males  have 
selected  the  more  beautiful  females,  and  have  thus  slowly  added 
to  their  beauty.  We  do  not  know  why  in  various  classes  of 
animals  the  males  of  some  few  sjDecies  have  selected  the  more 
beautiful  females  instead  of  having  gladly  accepted  any  female, 
as  seems  to  be  the  general  rule  in  the  animal  kingdom ;  but  if, 
contrary  to  what  generally  occurs  with  the  Lepidoptera,  the 
females  were  much  more  numerous  than  the  males,  the  latter 
would  be  likely  to  pick  out  the  more  beautiful  females.  Mr. 
Butler  shewed  me  several  species  of  Callidryas  in  the  British 
Museum,  in  some  of  which  the  females  equalled,  and  in  others 
gi-eatly  surpassed  the  males  in  beauty;  for  the  females  alone 
have  the  borders  of  their  wings  suffused  with  crimson  and 
orange,  and  spotted  with  black.  The  plainer  males  of  these 
species  closely  resemble  each  other,  showing  that  here  the 
females  have  been  modified ;  whereas  in  those  cases,  where  the 
males  are  the  more  ornate,  it  is  these  which  have  been  modified, 
the  females  remaining  closely  alike. 


Chap.  XL  Butterflies  a7id  Moths,  319 


In  England  we  have  some  analogous  cases,  though  not  so 
marked.  The  females  alone  of  two  species  of  Thecla  have  a 
bright-purple  or  orange  patch  on  their  fore-wings.  In  Hip- 
parchia  the  sexes  do  not  differ  much ;  but  it  is  the  female  of 
H.  janira  which  has  a  conspicuous  light-brown  patch  on  her 
wings ;  and  the  females  of  some  of  the  other  species  are  brighter 
coloured  than  their  males.  Again,  the  females  of  CoJias  ednsa 
and  hyale  have  "  orange  or  yellow  spots  on  the  black  marginal 
"  border,  represented  in  the  males  only  by  thin  streaks ; "  and 
in  Pieris  it  is  the  females  which  "  are  ornamented  with  black 
"  spots  on  the  fore-wings,  and  these  are  only  partially  present 
"  in  the  males."  Now  the  males  of  many  butterflies  are  known 
to  support  the  females  during  their  marriage  flight ;  but  in  the 
species  just  named  it  is  the  females  which  support  the  males ; 
so  that  the  part  which  the  two  sexes  play  is  reversed,  as  is  their 
relative  beauty.  Throughout  the  animal  kingdom  the  males 
commonly  take  the  more  active  share  in  wooing,  and  their 
beauty  seems  to  have  been  increased  by  the  females  having 
accepted  the  more  attractive  individuals ;  but  with  these  but- 
terflies, the  females  take  the  more  active  part  in  the  final  mar- 
riage ceremony,  so  that  we  may  suppose  that  they  likewise  do 
so  in  the  wooing ;  and  in  this  case  we  can  understand  how  it  is 
that  they  have  been  rendered  the  more  beautiful.  Mr.  Meldola, 
from  whom  the  foregoing  statements  have  been  taken,  sajs  in 
conclusion ;  "  Though  I  am  not  convinced  of  the  action  of 
"  sexual  selection  in  producing  the  colours  of  insects,  it  cannot 
"  be  denied  that  these  facts  are  strikingly  corroborative  of 
"  Mr.  Darwin's  views."  ^^ 

As  sexual  selection  primarily  depends  on  variability,  a  few 
words  must  be  added  on  this  subject.  In  respect  to  colour 
there  is  no  difficulty,  for  any  number  of  highly  variable  Lepi- 
doptera  could  be  named.  One  good  instance  will  suffice.  Mr. 
Bates  shewed  me  a  whole  series  of  specimens  of  Papilio  sesostris 
and  F.  childrence :  in  the  latter  the  males  varied  much  in  the 
extent  of  the  beautifully  enamelled  green  patch  on  the  fore- 
wings,  and  in  the  size  of  the  white  mark,  and  of  the  splendid 
crimson  stripe  on  the  hind- wings ;  so  that  there  was  a  great 
contrast  amongst  the  males  between  the  most  and  the  least 
gaudy.  The  male  of  Papilio  sesostris  is  much  less  beautiful 
than  of  P.  childrence:  and  it  likewise  varies  a  little  in  the  size  of 

23  'Nature,' April  27th,  1871,  p.  whilst    pairing.     See    also    Mr.    G. 

508,     Mr.   Meldola  quotes    Donzel,  Fraser,    in    '  Nature,'   April    20th, 

in  'Soc.  Ent.  de   France,'  1837,   p.  1871,  p.  489,  on  the  sexual  differ- 

77,    on    the    flight    of    butterflies  ences  of  several  British  butterflies. 


320  The  Descent  of  Man.  Pakt  II. 


the  green  patcli  on  the  fore-wings,  and  in  the  occasional  ap- 
pearance of  the  small  crimson  stripe  on  the  hind-wings, 
borrowed,  as  it  would  seem,  from  its  own  female ;  for  the  females 
of  this  and  of  many  other  species  in  the  ^neas  group  possess 
this  crimson  stripe.  Hence  between  the  brightest  specimens 
of  P.  sesostris  and  the  dullest  of  P.  chiklrence,  there  was  but  a 
small  interval ;  and  it  was  evident  that  as  far  as  mere  varia- 
bility is  concerned,  there  would  be  no  difficulty  in  permanently 
increasing  the  beauty  of  either  species  by  means  of  selection. 
The  variability  is  here  almost  confined  to  the  male  sex;  but 
Mr.  Wallace  and  ]\Ir.  Bates  have  shewn  ^^  that  the  females  of 
some  species  are  extremely  variable,  the  males  being  nearly 
constant.  In  a  future  chapter  I  shall  have  occasion  to  shew 
that  the  beautiful  eye-like  spots,  or  ocelli,  found  on  the  wings  of 
many  Lepidoptera,  are  eminently  variable.  I  may  here  add 
that  these  ocelli  offer  a  difficulty  on  the  theory  of  sexual 
selection ;  for  though  appearing  to  us  so  ornamental,  they  are 
never  present  in  one  sex  and  absent  in  the  other,  nor  do  they 
ever  differ  much  in  the  two  sexes.^^  This  fact  is  at  present 
inexphcable ;  but  if  it  should  hereafter  be  found  that  the  for- 
mation of  an  ocellus  is  due  to  some  change  in  the  tissues 
of  the  wings,  for  instance,  occui'ring  at  a  very  early  period  of 
development,  we  might  expect,  from  what  we  know  of  the  laws 
of  inheritance,  that  it  would  be  transmitted  to  both  sexes,  though 
arising  and  perfected  in  one  sex  alone. 

On  the  whole,  although  many  serious  objections  may  be 
urged,  it  seems  probable  that  most  of  the  brilliantly  coloured 
species  of  Lepidoptera  owe  their  colours  to  sexual  selection, 
excepting  in  certain  cases,  presently  to  be  mentioned,  in  which 
conspicuous  colours  have  been  gained  through  mimicry  as 
a  protection.  From  the  ardour  of  the  male  throughout  the 
animal  kingdom,  he  is  generally  willing  to  accept  any  female ; 
and  it  is  the  female  which  usually  exerts  a  choice.  Hence,  if 
sexual  selection  has  been  efficient  with  the  Lepidoptera,  the 
male,  when  the  sexes  differ,  ought  to  be  the  more  brilliantly 
coloured,  and  this  undoubtedly  is  the  case.  When  both  sexes 
are  brilliantly  coloured  and  resemble  each  other,  the  characters 
acquired  by  the  males  appear  to  have  been  transmitted  to  both. 

2*  Wallace  on  the  Papilionida;  of  tomolog.  Soc'  Nov.  19th,  1866,  p. 

the  Malayan  Region,  in  '  Transact,  xl. 

Linn.  Soc'  vol.   xxv.    1865,   pp.   8,  "^  ]\Ir.  Bates  was  so  kind   as   to 

36.     A    striking    case    of    a    rare  lay  this  subject  before  the  Entomo- 

variety,    strictly    intermediate    be-  logical  Society,  and  I  hare  received 

tween  two  other  well-marked  female  answers  to  this  eftect  from  several 

varieties,  is  given  by  Mr.  Wallace,  entomologists. 
See   also  Mr.  Bates,  in  '  Proc.  En- 


Chap.  XI.  Butterflies  and  Moths.  321 

We  are  led  to  this  conclusion  by  cases,  even  within  the  same 
genus,  of  gradation  from  an  extraordinary  amount  of  difference 
to  identity  in  colour  between  the  two  sexes. 

But  it  may  be  asked  whether  the  differences  in  colour  between 
the  sexes  may  not  be  accounted  for  by  other  means  besides 
sexual  selection.  Thus  the  males  and  females  of  the  same 
species  of  butterfly  are  in  several  cases  known  ^"^  to  inhabit 
dilferent  stations,  the  former  commonly  basking  in  the  sunshine, 
the  latter  haunting  gloomy  forests.  It  is  therefore  possible  that 
different  conditions  of  life  may  have  acted  directly  on  the  two 
sexes ;  but  this  is  not  probable,^^  as  in  the  adult  state  tlicy  are 
exposed  to  different  conditions  during  a  very  short  period ;  and 
the  larvae  of  both  are  exposed  to  the  same  conditions.  Mr. 
Wallace  believes  that  the  difference  between  the  sexes  is  due 
not  so  much  to  the  males  having  been  modified,  as  to  the  females 
having  in  all  or  almost  all  cases  acquired  dull  colours  for  the 
sake  of  protection.  It  seems  to  me,  on  the  contrary,  far  more 
probable  that  it  is  the  males  which  have  been  chiefly  modified 
through  sexual  selection,  the  females  having  been  comparatively 
little  changed.  We  can  thus  understand  how  it  is  that  the 
females  of  allied  species  generally  resemble  one  another  so  much 
more  closely  than  do  the  males.  They  thus  shew  us  ap- 
proximately the  primordial  colouring  of  the  parent-species  of 
the  group  to  which  they  belong.  They  have,  however,  almost 
always  been  somewhat  modified  by  the  transfer  to  them  of  some 
of  the  successive  variations,  through  the  accumulation  of  which 
the  males  were  rendered  beautiful.  But  I  do  not  wish  to  deny 
that  the  females  alone  of  some  species  may  have  been  specially 
modified  for  protection.  In  most  cases  the  males  and  females  of 
distinct  species  will  have  been  exposed  during  their  prolonged 
larval  state  to  different  conditions,  and  may  have  been  thus 
affected;  though  with  the  males  any  slight  change  of  colour 
thus  caused  will  generally  have  been  masked  by  the  brilliant 
tints  gained  through  sexual  selection.  When  we  treat  of  Birds, 
I  shall  have  to  discuss  the  whole  question,  as  to  how  far  the 
differences  in  colour  between  the  sexes  are  due  to  the  males 
having  been  modified  through  sexual  selection  for  ornamental 
purposes,  or  to  the  females  having  been  modified  through 
natural  selection  for  the  sake  of  protection,  so  that  I  will  here 
say  but  little  on  the  subject. 

In  all  the  cases  in  which  the  more  common  form  of  equal 

2«  H.  W.  Bates,  'The  Naturalist  "  Qu  this  whole  subject  see  'The 

on    the  Amazons,'  vol.  ii.   1863,  p.  Variation    of    Animals    and    Plants 

228.     A.  R.  Wallace,  in  'Transact,  under  Domestication,'  18(J8,  vol.  ii. 

Linn.  Soc'  vol.  xxv.  1865,  p.  10.  chap,  xxiii. 

15 


322  The  Descent  of  Mmi.  Part  II. 

inheritance  by  both  sexes  has  prevailed,  the  selection  of  bright- 
coloured  males  would  tend  to  make  the  females  bright-coloured; 
and  the  selection  of  dull-coloured  females  would  tend  to  make 
the  males  dull.  If  both  processes  were  carried  on  simultaneously, 
they  would  tend  to  counteract  each  other ;  and  the  final  result 
would  depend  on  whether  a  greater  number  of  females  from 
being  well  protected  by  obscure  colours,  or  a  greater  number  of 
males  by  l>eing  brightly-coloured  and  thus  finding  partners, 
succeeded  in  leaving  more  numerous  oflfspring. 

In  order  to  account  for  the  frequent  transmission  of  characters 
to  one  sex  alone,  Mr.  Wallace  expresses  his  belief  that  the  more 
common  form  of  equal  inheritance  by  both  sexes  can  be  chan.2:ed 
through  natural  selection  into  inheritance  by  one  sex  alone,  but 
in  favour  of  this  view  I  can  discover  no  evidence.  We  know 
from  what  occurs  under  domestication  that  new  characters  often 
appear,  which  from  the  first  are  transmitted  to  one  sex  alone ; 
and  by  the  selection  of  such  variations  there  would  not  be  the 
slightest  difficulty  in  giving  bright  colours  to  the  males  alone, 
and  at  the  same  time  or  subsequently,  dull  colours  to  the  females 
alone.  In  this  manner  the  females  of  some  butterflies  and  moths 
have,  it  is  probable,  been  rendered  inconspicuous  for  the  sake  of 
protection,  and  widely  different  from  their  males. 

I  am,  however,  unwilling  without  distinct  evidence  to  admit 
that  two  complex  processes  of  selection,  each  requiring  the 
transference  of  new  characters  to  one  sex  alone,  have  been 
carried  on  with  a  multitude  of  species, — that  the  males  have 
been  rendered  more  brilliant  by  beating  their  rivals,  and  the 
females  more  dull-coloured  by  having  escaped  from  their 
enemies.  The  male,  for  instance,  of  the  common  brimstone 
butterfly  (Gonepteryx),  is  of  a  far  more  intense  yellow  than  the 
female,  though  she  is  equally  conspicuous ;  and  it  does  not  seetn 
probable  that  she  specially  acquired  her  pale  tints  as  a  protec- 
tion, though  it  is  probable  that  the  male  acquired  his  bright 
colours  as  a  sexual  attraction.  The  female  of  Anthocharis  car- 
damines  does  not  possess  the  beautiful  orange  wing-tips  of  the 
male;  consequently  she  closely  resembles  the  white  butterflies 
(Pieris)  so  common  in  our  gardens ;  but  we  have  no  evidence 
that  this  resemblance  is  beneficial  to  her.  As,  on  the  other  hand, 
she  resembles  both  sexes  of  several  other  species  of  the  genus 
inhabiting  various  quarters  of  the  world,  it  is  probable  that  she 
has  simply  retained  to  a  large  extent  her  primordial  colours. 

Finally,  as  we  have  seen,  various  considerations  lead  to  the 
conclusion  that  with  the  greater  number  of  brilliantly-coloured 
Lepidoptera  it  is  the  male  which  has  been  chiefly  modified 
through   sexual   selection;    the  amount  of  difference  between 


Chap.  XT.  Btitterjiies  and  Moths.  323 

the  sexes  mostly  depending  on  tlie  form  of  inheritance  which 
has  prevailed.  Inheritance  is  governed  by  so  many  unknown 
laws  or  conditions,  that  it  seems  to  us  to  act  in  a  capricious 
manner  ;  ^*  and  we  can  thus,  to  a  certain  extent,  understand  how 
it  is  that  with  closely  allied  species  the  sexes  either  differ  to  an 
astonishing  degree,  or  are  identical  in  colour.  As  all  the  suc- 
cessive steps  in  the  process  of  variation  are  necessarily  trans- 
mitted through  the  female,  a  greater  or  less  number  of  such 
steps  might  readily  become  developed  in  her;  and  thus  we  can 
understand  the  frequent  gradations  from  an  extreme  difference 
to  none  at  all  between  the  sexes  of  allied  species.  These  cases  of 
gradation,  it  may  be  added,  are  much  too  common  to  favour  the 
supposition  that  we  here  see  females  actually  undergoing  the 
]3rocess  of  transition  and  losing  their  brightness  for  the  sake  of 
protection ;  for  w^e  have  every  reason  to  conclude  that  at  any 
one  time  the  greater  number  of  speci(;s  are  in  a  fixed  condition. 

Mimicry. — This  principle  was  first  made  clear  in  an  admirable 
paper  by  Mr.  Bates,-^  who  thus  threw  a  flood  of  light  on  many 
obscure  problems.  It  had  previously  been  observed  that  certain 
butterflies  in  S,  America  belonging  to  quite  distinct  families, 
resembled  the  Heliconidse  so  closely  in  every  stripe  and  shade  of 
colour,  that  they  could  not  be  distinguished  save  by  an  ex- 
perienced entomologist.  As  the  Heliconidse  are  coloured  in 
their  usual  manner,  whilst  the  others  depart  from  the  usual 
colouring  of  the  groups  to  which  they  belong,  it  is  clear  that 
the  latter  are  the  imitators,  and  the  Heliconidee  the  imitated. 
Mr.  Bates  further  observed  that  the  imitating  species  are  com- 
paratively rare,  whilst  the  imitated  abound,  and  that  the  two 
sets  live  mingled  together.  From  the  fact  of  the  HeliconidsB 
being  conspicuous  and  beautiful  insects,  yet  so  numerous  in 
individuals  and  species,  he  concluded  that  they  must  be  pro- 
tected from  the  attacks  of  enemies  by  some  secretion  or  odour  ; 
and  this  conclusion  has  now  been  amply  confirmed,-*'  especially 
by  Mr.  Belt.  Hence  Mr.  Bates  inferred  that  the  butterflies 
which  imitate  the  protected  species  have  acquired  their  present 
marvellously  deceptive  appearance  through  variation  and  natural 
selection,  in  order  to  be  mistaken  for  the  protected  kinds,  and 
thus  to  escape  being  devoured  No  explanation  is  here  attempted 
of  the  brilliant  colours  of  the  imitated,  but  only  of  the  imitating 
butterflies.  We  must  account  for  the  colours  of  the  former  in 
the  same  general  manner,  as  in  the  cases  previously  discussed 

"*  '  The  Variation  of  Animals  and  xxiii.  1802,  p.  495. 
Plants  under  Domestication,'  vol.  ii.  ^o  <  Pyoc.    Ent.    Soc'     Dec.    3rd, 

chap.  xii.  p.  17.  1866,  p.  xlv. 

29    '  Transact.    Linn.    Soc'    vol. 


324  The  Descent  of  Man.  Part  II. 


in  this  chapter.  Since  the  publication  of  Mr.  Bates'  paper, 
similar  and  equally  striking  facts  have  been  observed  by 
]^[r.  Wallace  in  the  Malayan  region,  by  Mr,  Trimen  in  South 
Africa,  and  by  Mr.  Eiley  in  the  United  States.^^ 

As  some  writers  have  felt  much  difficulty  in  understanding 
how  the  first  steps  in  the  process  of  mimicry  could  have  been 
effected  through  natural  selection,  it  may  be  well  to  remark  that 
the  process  probably  commenced  long  ago  between  forms  not 
widely  dissimilar  in  colour.  In  this  case  even  a  slight  variation 
would  be  beneficial,  if  it  rendered  the  one  species  more  Like 
the  other ;  and  afterwards  the  imitated  species  might  be  modi- 
fied to  an  extreme  degree  through  sexual  selection  or  other 
means,  and  if  the  changes  were  gradual,  the  imitators  might 
easily  be  led  along  the  same  track,  until  they  differed  to  an 
equally  extreme  degree  from  their  original  condition ;  and  they 
would  thus  ultimately  assume  an  appearance  or  colouring  wholly 
unlike  that  of  the  other  members  of  the  group  to  which  they 
belonged.  It  should  also  be  remembered  that  many  species  of 
Lepidoptera  are  liable  to  considerable  and  abrupt  variations  in 
colour.  A  few  instances  have  been  given  in  this  chapter ;  and 
many  more  may  be  found  in  the  papers  of  Mr.  Bates  and 
BIr.  Wallace. 

With  several  species  the  sexes  are  alike,  and  imitate  the  two 
sexes  of  another  species.  But  Mr.  Trimen  gives,  in  the  paper 
already  referred  to,  three  cases  in  which  the  sexes  of  the  imitated 
form  differ  from  each  other  in  colour,  and  the  sexes  of  the 
imitating  form  differ  in  a  like  manner.  Several  cases  have  also 
been  recorded  where  the  females  alone  imitate  brilliantly- 
coloured  and  protected  species,  the  males  retaining  "the 
"  normal  aspect  of  their  immediate  congeners."  It  is  here  obvious 
that  the  successive  variations  by  which  the  female  has  been 
modified  have  been  transmitted  to  her  alone.  It  is,  however, 
probable  that  some  of  the  many  successive  variations  would 
have  been  transmitted  to,  and  developed  in,  the  males  had 
not  such  males  been  eliminated  by  being  thus  rendered  less 
attractive  to  the  females;  so  that  only  those  variations  were 
preserved  which  were  from  the  fii'st  strictly  limited  in  their 
transmission  to  the  female  sex.  We  have  a  partial  illus- 
tration of  these  remarks  in  a  statement  by  Mr.  Belt;='-  that 

'^  Wallace,  'Transact.  Linn.  Soc'  163-168.     This  latter  essay  is  valu- 

vol.  XXV.  1865,  p.  1 ;  also  '  Transact,  able,  as  Mr.  Riley  here  discusses  all 

Ent.  Soc'  vol.  iv.  (3rd  series),  1867,  the    objections    which    have    been 

p.  301.     Trimen,  '  Linn.  Transact.'  raised  against  Mr.  Bates'  theory, 

vol.    xxvi.     1869,    p.    497.      Riley,  ,      ^-  '  The  Naturalist  in  Nicaragua, 

'Third  Annual  Report  on  the  Noxi-  1874,  p.  385. 
ous   Insects  of  Missouri,'  1871,  pp 


OiiAP.  Xr.  Butterjiies  and  Moths.  325 


the  males  of  some  of  the  Leptalides,  which  imitate  protected 
species,  still  retain  in  a  concealed  manner  some  of  their  original 
characters.  Thus  in  the  males  ''the  npper  half  of  the  lower 
"  wing  is  of  a  pure  white,  whilst  all  the  rest  of  the  wings  is 
"  barred  and  sjDottcd  with  black,  red  and  yellow,  like  the  species 
"  they  mimic.  The  females  have  not  this  white  patch,  and  the 
"  males  usually  conceal  it  by  covering  it  with  the  ^^pper  wing, 
"  so  that  I  cannot  imagine  its  being  of  any  other  use  to  them 
"  than  as  an  attraction  in  courtship,  when  they  exhibit  it  to  the 
"  females,  and  thus  gratify  their  deep-seated  preference  for  the 
"  normal  colour  of  the  Order  to  which  the  Leptalides  belong." 

Briijlit  Colours  of  Caterpillars.  —  Whilst  reflecting  on  the 
beauty  of  many  butterflies,  it  occurred  to  me  that  some  cater- 
pillars were  splendidly  coloured ;  and  as  sexual  selection  could 
not  possibly  have  here  acted,  it  appeared  rash  to  attribute  the 
beauty  of  the  mature  insect  to  this  agency,  unless  the  bright 
colours  of  their  larvas  could  be  somehow  explained.  In  the 
first  place,  it  may  be  observed  that  the  colours  of  caterpillars  do 
not  stand  in  any  close  correlation  with  those  of  the  mature 
insect.  Secondly,  their  bright  colours  do  not  serve  in  any 
ordinary  manner  as  a  protection.  Mr.  Bates  informs  me,  as  an 
instance  of  this,  that  the  most  conspicuous  caterpillar  which  he 
ever  beheld  (that  of  a  Sphinx)  lived  on  the  large  green  leaves  of 
a  tree  on  the  open  llanos  of  South  America ;  ]t  was  about  four 
inches  in  length,  transversely  banded  with  black  and  yellow, 
and  with  its  head,  legs,  and  tail  of  a  bright  red.  Hence  it 
caught  the  eye  of  any  one  who  passed  by,  even  at  the  distance  of 
many  yards,  and  no  doubt  that  of  every  passing  bird. 

I  then  applied  to  Mr.  Wallace,  who  has  an  innate  genius  for 
solving  difficulties.  After  some  consideration  he  replied  :  "  Most 
*'  caterpillars  require  protection,  as  may  be  inferred  from  some 
*'  kinds  being  furnished  with  spines  or  irritating  hairs,  and 
"  from  many  being  coloured  green  like  the  leaves  on  which  they 
"  feed,  or  being  curiously  like  the  twigs  of  the  trees  on  which  they 
"  live."  Another  instance  of  protection,  furnished  me  by  Mr,  J. 
Mansel  Weale,  may  be  added,  namely,  that  there  is  a  caterpillar 
of  a  moth  which  lives  on  the  mimosas  in  South  Africa,  and 
fabricates  for  itself  a  case  quite  indistinguishable  from  the 
surrounding  thorns.  From  such  considerations  Mr.  Wallace 
thought  it  probable  that  conspicuously-coloured  oateri^illars 
were  protected  by  having  a  nauseous  taste ;  but  as  their  skin 
is  extremely  tender,  and  as  their  intestines  readily  protrude 
from  a  wound,  a  slight  peck  from  the  beak  of  a  bird  would 
be  as  fatal  to  them  as  if  they  had  been  devoured.     Hence,  as 


326  The  Descent  of  Man.  Paet  II 

Mr.  Wallace  remarks,  "  distastefulness  alone  would  be  insufficient 
"  to  protect  a  caterpillar  unless  some  outward  sign  indicated  to 
"  its  would-be  destroyer  that  its  prey  was  a  disgusting  morsel." 
Under  these  circumstances  it  would  be  highly  advantageous  to 
a  caterpillar  to  be  instantaneously  and  certainly  recognised  as 
unpalatable  by  all  birds  and  other  animals.  Thus  the  most 
gaudy  colours  would  be  serviceable,  and  might  have  been 
gained  by  variation  and  the  sui'vival  of  the  most  easily-re- 
cognised individuals. 

This  hypothesis  appears  at  first  sight  very  bold,  but  when  it 
was  brought  before  the  Entomological  Society  ^^  it  was  supported 
by  various  statements ;  and  Mr.  J.  Jenner  Weir,  who  keeps  a 
large  number  of  birds  in  an  aviary,  informs  me  that  he  has 
made  many  trials,  and  finds  no  exception  to  the  rule,  that  all 
caterpillars  of  nocturnal  and  retiring  habits  with  smooth  skins, 
all  of  a  green  colour,  and  all  which  imitate  twigs,  are  greedily 
devoured  by  his  birds.  The  hairy  and  spinose  kinds  are 
in  variably  rejected,  as  were  four  conspicuously-coloured  species. 
When  the  birds  rejected  a  caterpillar,  they  plainly  shewed,  by 
shaking  their  heads,  and  cleansing  their  beaks,  that  they  were 
disgusted  by  the  taste.^  Three  conspicuous  kinds  of  cater- 
pillars and  moths  were  also  given  to  some  lizards  and  frogs,  by 
Mr.  A.  Butler,  and  were  rejected,  though  other  kinds  were 
eagerly  eaten.  Thus  the  probability  of  Mr.  Wallace's  view  is 
confirmed,  namely,  that  certain  caterpillars  have  been  made 
conspicuous  for  their  own  good,  so  as  to  be  easily  recognised  by 
their  enemies,  on  nearly  the  same  principle  that  poisons  are  sold 
in  coloured  bottles  by  druggists  for  the  good  of  man.  We 
cannot,  however,  at  present  thus  explain  the  elegant  diversity 
in  the  colours  of  many  caterpillars ;  but  any  species  which  had 
at  some  former  period  acquired  a  dull,  mottled,  or  strijDed  appear- 
ance, either  in  imitation  of  surrounding  objects,  or  from  the 
direct  action  of  climate,  &c.,  almost  certainly  would  not  become 
uniform  in  colour,  when  its  tints  were  rendered  intense  and 
bright ;  for  in  order  to  make  a  caterpillar  merely  conspicuous, 
there  would  be  no  selection  in  any  definite  direction. 

Summary  and  Concluding  Bemarhs  on  Insects. — Looking  back 

23  *  Proc.    Entomolog.    Soc*   Dec.  analogous  facts  in  the  '  Third  An- 

3rd,  1866,  p.  xlv.,  and  March  4th,  nual  Report  on  the  Noxious  Insects 

1867,  p.  Ixxx.  of  Missouri,'  1871,  p.   148.     Some 

^'^  See     Mr.     J.    Jenner     Weir's  opposed  cases  are,  however,  given  hy 

paper  on  Insects   and  Insectivorous  Dr.  Wallace    and   M,  H.  d'Orville ; 

Birds,  in  '  Transact.  Ent.  Soc' 1869,  see    'Zoological    Record,'    1869,    p. 

p.    21  ;    also    Mr.    Butler's    paper,  349. 
ibid.  p.   27.     Mr.  Riley  has  given 


Chap.  XI.  Stivimary  on  Insects.  327 

to  the  several  Orders,  we  see  that  the  sexes  often  differ  in 
various  characters,  the  meaning  of  which  is  not  in  the  least 
understood.  The  sexes,  also,  often  differ  in  their  organs  of 
sense  and  means  of  locomotion,  so  that  the  males  may  quickly 
discover  and  reach  the  females.  They  differ  still  oftener  in 
the  males  possessing  diversified  contrivances  for  retaining  the 
females  when  found.  We  are,  however,  here  concerned  only  in 
a  secondary  degree  with  sexual  differences  of  these  kinds. 

In  almost  all  the  Orders,  the  males  of  some  species,  even  of 
weak  and  delicate  kinds,  are  known  to  be  highly  pugnacious ; 
and  some  few  are  furnished  with  special  weapons  for  fighting 
with  their  rivals.  But  the  law  of  battle  does  not  prevail  nearly 
so  widely  with  insects  as  with  the  higher  animals.  Hence  it 
probably  arises,  that  it  is  in  only  a  few  cases  that  the  males  have 
been  rendered  larger  and  stronger  than  the  females.  On  the 
contrary,  they  are  usually  smaller,  so  that  they  may  be  developed 
within  a  shorter  time,  to  be  ready  in  large  numbers  for  the 
emergence  of  the  females. 

In  two  families  of  the  Homoptera  and  in  three  of  the  Orthop- 
tera,  the  males  alone  possess  sound-producing  organs  in  an 
efficient  state.  These  are  used  incessantly  during  the  breeding- 
season,  not  only  for  calling  the  females,  but  apparently  for 
charming  or  exciting  them  in  rivalry  with  other  males.  No 
one  who  admits  the  agency  of  selection  of  any  kind,  will,  after 
reading  the  above  discussion,  dispute  that  these  musical  instru- 
ments have  been  acquired  through  sexual  selection.  In  four 
other  Orders  the  members  of  one  sex,  or  more  commonly  of 
both  sexes,  are  provided  with  organs  for  producing  various 
sounds,  which  apparently  serve  merely  as  call-notes.  When 
both  sexes  are  thus  provided,  the  individuals  which  were  able 
to  make  the  loudest  or  most  continuous  noise  would  gain 
partners  before  those  which  were  less  noisy,  so  that  their  organs 
have  probably  been  gained  through  sexual  selection.  It  is 
instructive  to  reflect  on  the  wonderful  diversity  of  the  means 
for  producing  sound,  possessed  by  the  males  alone,  or  by  both 
sexes,  in  no  less  than  six  Orders.  We  thus  learn  how  effectual 
sexual  selection  has  been  in  leading  to  modifications  which 
sometimes,  as  with  the  Homoptera,  relate  to  important  parts  of 
the  organisation. 

From  the  reasons  assigned  in  the  last  chapter,  it  is  probable 
that  the  great  horns  possessed  by  the  males  of  many  Lamel- 
licorn,  and  some  other  beetles,  have  been  acquired  as  ornaments. 
From  the  small  size  of  insects,  we  are  apt  to  undervalue  their 
appearance.  If  we  could  imagine  a  male  Chalcosoma  (fig.  16), 
with  its  polished  bronzed  coat  of  mail,  and  its  vast  complex 


328  The  Descent  of  Man.  Paet  II. 

horns,  magnified  to  the  size  of  a  horse,  or  even  of  a  dog,  it  would 
be  one  of  the  most  imposing  animals  in  the  world. 

The  colouring  of  insects  is  a  complex  and  obscure  subject. 
When  the  male  differs  slightly  from  the  female,  and  neither  are 
briUiantly-coloured,  it  is  probable  that  the  sexes  have  varied 
in  a  slightly  different  manner,  and  that  the  variations  have  been 
transmitted  by  each  sex  to  the  same,  without  any  benefit  or 
evil  thus  accruing.  When  the  male  is  brilliantly-coloured  and 
difi"ers  conspicuously  from  the  female,  as  with  some  dragon-flies 
and  many  butterflies,  it  is  probable  that  he  owes  his  colours  to 
sexual  selection ;  whilst  the  female  has  retained  a  primordial  or 
very  ancient  type  of  colouring,  slightly  modified  by  the  agencies 
before  explained.  But  in  some  cases  the  female  has  a^Dparently 
been  made  obscure  by  variations  transmitted  to  her  alone, 
as  a  means  of  direct  protection ;  and  it  is  almost  certain  that 
she  has  sometimes  been  made  brilliant,  so  as  to  imitate  other 
protected  species  inhabiting  the  same  district.  When  the  sexes 
resemble  each  other  and  both  are  obscurely  coloured,  there  is 
no  doubt  that  they  have  been  in  a  multitude  of  cases  so  coloured 
for  the  sake  of  protection.  So  it  is  in  some  instances  when  both 
are  brightly-coloured,  for  they  thus  imitate  protected  species,  or 
resemble  surrounding  objects  such  as  flowers;  or  they  give 
notice  to  their  enemies  that  they  are  unpalatable.  In  other 
cases  in  which  the  sexes  resemble  each  other  and  are  both 
brilliant,  especially  when  the  colours  are  arranged  for  display, 
we  may  conclude  that  they  have  been  gained  by  the  male  sex  as 
an  attraction,  and  have  been  transferred  to  the  female.  We  are 
more  especially  led  to  this  conclusion  whenever  the  same  typo 
of  coloration*  prevails  throughout  a  whole  group,  and  we  find 
that  the  males  of  some  species  difier  widely  in  colour  from 
the  females,  whilst  others  differ  slightly  or  not  at  all,  with 
intermediate  gradations  connecting  these  extreme  states. 

In  the  same  manner  as  bright  colours  have  often  been 
partially  transferred  from  the  males  to  the  females,  so  it  has 
been  with  the  extraordinary  horns  of  many  Lamellicorn  and 
some  other  beetles.  So  again,  the  sound-producing  organs 
proper  to  the  males  of  the  Homoptera  and  Orthoptera  have 
generally  been  transferred  in  a  rudimentary,  or  even  in  a  nearly 
perfect  condition,  to  the  females;  yet  not  sufficiently  perfect  to 
be  of  any  use.  It  is  also  an  interesting  fact,  as  bearing  on 
sexual  selection,  that  the  stridulating  organs  of  certain  male 
Orthoptera  are  not  fully  developed  until  the  last  moult;  and  that 
the  colours  of  certain  male  dragon-flies  are  not  fully  develoi^ed 
until  some  little  time  after  their  emergence  from  the  pupal 
state,  and  when  they  are  ready  to  breed. 


Chap.  XI.  Summary  on  Insects.  329 

Sexual  selection  implies  that  the  more  attractive  individuals 
are  preferred  by  the  opposite  sex ;  and  as  with  insects,  when 
the  sexes  differ,  it  is  the  male  which,  with  some  rare  exceptions, 
is  the  more  ornamented,  and  departs  more  from  the  type  to 
which  the  species  belongs ; — and  as  it  is  the  male  which  searches 
eagerly  for  the  female,  we  must  suppose  that  the  females 
habitually  or  occasionally  prefer  the  more  beautiful  males,  and 
that  these  have  thus  acquired  their  beauty.  That  the  females 
in  most  or  all  the  orders  would  have  the  power  of  rejecting 
any  particular  male,  is  probable  from  the  many  singular  con- 
trivances  possessed  by  the  males,  such  as  great  jaws,  adhesive 
cushions,  spines,  elongated  legs,  &c.,  for  seizing  the  female ;  for 
these  contrivances  shew  that  there  is  some  difficulty  in  the  act, 
so  that  her  concurrence  would  seem  necessary.  Judging  from 
what  we  know  of  the  perceptive  powers  and  affections  of 
various  insects,  there  is  no  antecedent  improbability  in  sexual 
selection  having  come  largely  into  play ;  but  we  have  as  yet  no 
direct  evidence  on  this  head,  and  some  facts  are  opposed  to  the 
belief.  Nevertheless,  when  we  see  many  males  pursuing  the 
same  female,  we  can  hardly  believe  that  the  pairing  is  left  to 
blind  chance — that  the  female  exerts  no  choice,  and  is  not 
influenced  by  the  gorgeous  colours  or  other  ornaments  with 
which  the  male  is  decorated. 

If  we  admit  that  the  females  of  the  Homoptera  and  Orthoptera 
appreciate  the  musical  tones  of  their  male  partners,  and  that  the 
various  instruments  have  been  perfected  through  sexual  se- 
lection, there  is  little  improbability  in  the  females  of  other 
insects  appreciating  beauty  in  form  or  colour,  and  consequently 
in  such  characters  having  been  thus  gained  by  the  males.  But 
from  the  circumstance  of  colour  being  so  variable,  and  from  its 
having  been  so  often  modified  for  the  sake  of  protection,  it  is 
difficult  to  decide  in  how  large  a  jn-oportion  of  cases  sexual 
selection  has  played  a  part.  This  is  more  especially  difficult  in 
those  Orders,  such  as  Orthoptera,  HymenoiDtera,  and  Coleop- 
tera,  in  which  the  two  sexes  rarely  differ  much  in  colour ;  for 
we  are  then  left  to  mere  analogy.  With  the  Coleoptera,  however, 
as  before  remarked,  it  is  in  the  great  Lamellicorn  group,  placed 
by  some  authors  at  the  head  of  the  Order,  and  in  which  we 
sometimes  see  a  mutual  attachment  between  the  sexes,  that 
we  find  the  males  of  some  species  possessing  weapons  for  sexual 
strife,  others  furnished  with  wonderful  horns,  many  with  stridu- 
lating  organs,  and  others  ornamented  with  splendid  metallic 
tints.  Hence  it  seems  probable  that  all  these  characters  have 
been  gained  through  the  same  means,  namely  sexual  selection. 
With  butterflies  we    have    the    best  evidence,  as  the  males 


330  The  Descent  of  Man.  Paet  II. 

sometimes  take  pains  to  display  their  beautiful  colours;  and  we 
cannot  believe  that  they  would  act  thus,  unless  the  display  was 
of  use  to  them  in  their  courtship. 

When  we  treat  of  Birds,  we  shall  see  that  they  present  in 
their  secondary  sexual  characters  the  closest  analogy  with 
insects.  Thus,  many  male  birds  are  highly  pugnacious,  and 
some  are  furnished  with  special  weapons  for  fighting  with  their 
rivals.  They  possess  organs  which  are  used  dui'ing  the  breeding- 
season  for  producing  vocal  and  instrumental  music.  They  are 
frequently  ornamented  with  combs,  horns,  wattles  and  plumes 
of  the  most  diversified  kinds,  and  are  decorated  with  beautiful 
colours,  all  evidently  for  the  sake  of  display.  "We  shall  find 
that,  as  with  insects,  both  sexes  in  certain  groups  are  equally 
beautiful,  and  are  equally  provided  with  ornaments  which  are 
usually  confined  to  the  male  sex.  In  other  groups  both  sexes 
are  equally  plain-coloured  and  unornamented.  Lastly,  in  some 
few  anomalous  cases,  the  females  are  more  beautiful  than  the 
males.  We  shall  often  find,  in  the  same  group  of  birds,  every 
gradation  from  no  diff"erence  between  the  sexes,  to  an  extreme 
difference.  We  shall  see  that  female  birds,  like  female  insects, 
often  possess  more  or  less  plain  traces  or  rudiments  of  characters 
which  properly  belong  to  the  males  and  are  of  use  only  to  them. 
The  analogy,  indeed,  in  all  these  respects  between  birds  and 
insects  is  curiously  close.  W^hatever  explanation  applies  to  the 
one  class  probably  applies  to  the  other ;  and  this  explanation, 
as  we  shall  hereafter  attempt  to  shew  in  fui'ther  detail,  is  sexual 
selection. 


CHAPTEK  XII. 


Secondary  Sexual  Characters  of  Fishes,  Amphibians, 
AND  Reptiles. 

Fishes:  Courtship  and  battles  of  the  males — Larger  size  of  the  females 
— Males,  bright  colours  and  ornamental  appendages;  other  strange 
characters — Colours  and  appendages  acquired  by  the  males  during  the 
breeding-season  alone — Fishes  with  both  sexes  brilliantly  coloured 
— Protective  colours — The  less  conspicuous  colours  of  the  female  cannot 
be  accounted  for  on  the  principle  of  protection — Male  tishes  building 
nests,  and  taking  charge  of  the  ova  and  young.  Ajniphibians  :  Dif- 
ferences in  structure  and  colour  between  the  sexes — Vocal  organs. 
Reptiles  :  Chelonians — Crocodiles — Snakes,  colours  in  some  cases  pro- 
tective— Lizards,  battles  of — Ornamental  appendages — Strange  dif- 
ferences in  structure  between  the  sexes — Colours — Sexual  differences 
almost  as  great  as  with  birds. 

We  have  now  arrived  at  the  great  sub-kingdom  of  the  Vertebrata, 
and  will  commence  with  the  lowest  class,  that  of  Fishes.     The 


Chap.  XII.  Fishes.  33 1 

males  of  Plagiostomous  fishes  (sharks,  rays)  and  of  Chimaeroid 
fislies  are  provided  with  claspers  which  serve  to  retain  the 
female,  like  the  various  structures  possessed  by  many  of  the 
lower  animals.  Besides  the  claspers,  the  males  of  many  rays 
have  clusters  of  strong  sharp  spines  on  their  heads,  and  several 
rows  along  "  the  upper  outer  surface  of  their  pectoral  fins." 
These  are  present  in  the  males  of  some  species,  which  have 
other  parts  of  their  bodies  smooth.  They  are  only  temporarily 
developed  during  the  breeding-season ;  and  Dr.  GUnther  suspects 
that  they  are  brought  into  action  as  prehensile  organs  by  the 
doubling  inwards  and  downwards  of  the  two  sides  of  the  body. 
It  is  a  remarkable  fact  that  the  females  and  not  the  males  of 
some  species,  as  of  Ilaia  davata,  have  their  backs  studded  with 
large  hook-formed  spines.^ 

The  males  alone  of  the  capehn  {Mallotus  villosus,  one  of 
Salmonidse),  are  provided  with  a  ridge  of  closely-set,  brush-like 
scales,  by  the  aid  of  which  two  males,  one  on  each  side,  hold  the 
female,  whilst  she  runs  with  great  swiftness  on  the  sandy  beach, 
and  there  deposits  her  si3awn.^  The  widely  distinct  Monaccmthus 
scopas  presents  a  somewhat  analogous  structure.  The  male,  as 
Dr.  GiJnther  informs  me,  has  a  cluster  of  stiff,  straight  spines, 
like  those  of  a  comb,  on  the  sides  of  the  tail ;  and  these  in  a 
specimen  six  inches  long  were  nearly  one  and  a  half  inches  in 
length ;  the  female  has  in  the  same  place  a  cluster  of  bristles, 
which  may  be  compared  with  those  of  a  tooth-brush.  In 
another  species,  M.  peronii,  the  male  has  a  brush  like  that 
possessed  by  the  female  of  the  last  species,  whilst  the  sides  of 
the  tail  in  the  female  are  smooth.  In  some  other  species  of  the 
same  genus  the  tail  can  be  perceived  to  be  a  little  roughened  in 
the  male  and  perfectly  smooth  in  the  female;  and  lastly  in 
others,  both  sexes  have  smooth  sides. 

The  males  of  many  fish  fight  for  the  possession  of  the  females. 
Thus  the  male  stickleback  (Gasterosfeus  leiurus)  has  been  de- 
scribed as  "  mad  with  delight,"  when  the  female  comes  out  of  her 
hiding-place  and  surveys  the  nest  which  he  has  made  for  her. 
*'  He  darts  round  her  in  every  direction,  then  to  his  accumulated 
*'  materials  for  the  nest,  then  back  again  in  an  instant ;  and  as 
"  she  does  not  advance  he  endeavours  to  push  her  with  his  snout, 
"  and  then  tries  to  pull  her  by  the  tail  and  side- spine  to  the  nest."^ 

»  Yarrell's     'Hist,     of      British  1871,  p.  119. 

Fishes,'  vol.  ii.  1836,  pp.  417,  425,  3  See    Mr.     R.    Waringtoii's     in- 

436.     Dr.  Giinther  informs  me  that  terestiug   articles    in    '  Annals   and 

the  spines  in  R.  clacata  are  peculiar  Mag.  of  Nat.  Hist.'  Oct.   1852  and 

to  the  female.  Nov.  1855. 

-  *  The  American  Naturalist,' April 


332  The  Descent  of  Man.  Part  II. 

The  males  are  said  to  be  polygamists  ;^  they  are  extraordinarily 
bold  and  pugnacious,  whilst  "  the  females  are  quite  pacific." 
Their  battles  are  at  times  desperate ;  ''for  these  puny  com- 
"  batants  fasten  tight  on  each  other  for  several  seconds,  tumbling 
"  over  and  over  again,  until  their  strength  appears  completely 
"  exhausted."  With  the  rough-tailed  stickleback  (G^.  trachurus) 
the  males  whilst  fighting  swim  round  and  round  each  other, 
biting  and  endeavouring  to  pierce  each  other  with  their  raised 
lateral  spines.  The  same  writer  adds,^  "  the  bite  of  these  little 
"  furies  is  very  severe.  They  also  use  their  lateral  spines  with 
"  such  fatal  efiect,  that  I  have  seen  one  during  a  battle  absolutely 
"  rip  his  opponent  quite  open,  so  that  he  sank  to  the  bottom  and 
"  died."  When  a  fish  is  conquered,  "  his  gallant  bearing  forsakes 
"  him ;  his  gay  colours  fade  away ;  and  he  hides  his  disgrace 
"among  his  peaceable  companions,  but  is  for  some  time  the 
"  constant  object  of  his  conqueror's  persecution." 

The  male  salmon  is  as  pugnacious  as  the  little  stickleback ; 
and  so  is  the  male  trout,  as  I  hear  from  Dr.  Giinther.  Mr.  Shaw 
saw  a  violent  contest  between  two  male  salmon  which  lasted 
the  whole  day ;  and  Mr.  R.  Buist,  Sui3erintendent  of  Fisheries, 
informs  me  that  he  has  often  watched  from  the  bridge  at  Perth 
the  males  driving  away  their  rivals,  whilst  the  females  were 
spawning.  The  males  "  are  constantly  fighting  and  tearing  each 
"  other  on  the  spawning-beds,  and  many  so  injure  each  other  as 
"  to  cause  the  death  of  numbers,  many  being  seen  swimming  near 
"  the  banks  of  the  river  in  a  state  of  exhaustion,  and  apparently 
"  in  a  dying  state."^  Mr.  Buist  informs  me,  that  in  June  1868, 
the  keeper  of  the  Stormontfield  breeding-ponds  visited  the 
northern  Tyne  and  found  about  300  dead  salmon,  all  of  which 
with  one  exception  wei-e  males ;  and  he  was  convinced  that  they 
had  lost  their  lives  by  fighting. 

The  most  curious  point  about  the  male  salmon  is  that  during 
the  breeding-season,  besides  a  slight  change  in  colour,  "  the 
"  lower  jaw  elongates,  and  a  cartilaginous  projection  turns 
"  upwards  from  the  point,  which,  when  the  jaws  are  closed, 
"  occupies  a  deep  cavity  between  the  intermaxillary  bones  of  the 
upper  jaw."  '^  (Figs.  27  and  28.)  In  our  salmon  this  change  of 
structure  lasts  only  during  the  breeding-season;    but  in  the 

■*  Xoel  Humphreys,  'River    Gar-  experienced  observer  (Scrope's  *  Days 

dens,'  1857.  of  Salmon  Fishing/  p.  60)  remarks 

^  Loudon's  '  Mag.  of  Nat.  History,'  that  lilie  the  stag,  the  male  would, 

vol.  iii.  1830,  p.  331.  if  he  could,   keep   all   other  males 

«  'The  Field,'  June   29th,  18G7.  away. 
For     Mr.     Shaw's     statement,     see  ^  Yarrell,     '  History    of    British 

'Edinburgh  Review,'  1843.  Another  Fishes,'  vol.  ii.  1836,  p.  10. 


Ceap.  XIL 


Fishes. 


333 


Salmo  hjcabdon  of  N.-W.  America  the  change,  as  Mr.  J.  K.  Lord^ 
believes,  is  permanent,  and  best  marked  in  the  older  males  which 
have  previously  ascended  the  rivers.  In  these  old  males  the 
jaw  becomes  developed  into  an  immense  hook-like  projection,  and 


Fig.  27.     Head  of  male  common  salmon  (Salmo  salar)  during  the  breeding-season. 

TThis  drawing,  as  well  as  all  the  others  in  the  present  chaptnr,  have  been  executed 
by  the  well-known  artist,  Mr.  G.  Ford,  from  specimens  in  the  British  Museum,  under 
the  kind  superintendence  of  Dr.  Gtinther.J 

the  teeth  grow  into  regular  fangs,  often  more  than  half  an  inch 
in  length.  With  the  European  salmon,  according  to  Uv.  Lloyd,^ 
the   temporary  hook-like  structure   serves  to  strengthen  and 

*  'The  Naturalist  in  Vancouver's  ^  'Scandinavian  Adventures,' vol 

Island,'  vol.  i.  1866,  p.  54.  i.  1854,  pp.  100,  104. 


334 


The  Descent  of  Man, 


PartIL 


protect  the  jaws,  when  one  male  charges  another  with  wonderful 
violence  ;  but  the  greatly  developed  teeth  of  the  male  American 
salmon  may  be  compared  with  the  tusks  of  many  male  mammals, 
and  they  indicate  an  oifensive  rather  than  a  jDrotectivc  purpose. 


Fig  28 


Head  of  female  salmon. 


The  salmon  is  not  the  only  fish  in  which  the  teeth  differ  in 
the  two  sexes ;  as  this  is  the  case  with  many  rays.  In  the 
thornback  (^Eam  davatci)  the  adult  male  has  sharp,  pointed 
teeth,  directed  backwards,  whilst  those  of  the  female  are  broad 
and  flat,  and  form  a  pavement ;  so  that  tbese  teeth  differ  in  the 
two  sexes  of  the  same  species  more  than  is  usual  in  distinct 
genera  of  the  same  family.  The  teeth  of  the  male  become  sharp 
only  when  he  is  adult :  whilst  young  they  are  broad  and  fiat 


Cu AP.  X 1 1.  Fishes.  335 

like  those  of  the  female.  As  so  frequently  occurs  with  secondary 
sexual  characters,  both  sexes  of  some  species  of  rays  (for  instance 
R.  hatis),  when  adult,  possess  sharjD  pointed  teeth ;  and  here  a 
character,  proper  to  and  primarily  gained  by  the  male,  appears 
to  have  been  transmitted  to  the  offspring  of  both  sexes.  The 
teeth  are  likewise  pointed  in  both  sexes  of  11.  maculata,  but  only 
when  quite  adult ;  the  males  acquiring  them  at  an  earlier  age 
than  the  females.  We  shall  hereafter  meet  with  analogous 
cases  in  certain  birds,  in  which  the  male  acquires  the  plumage 
common  to  both  sexes  when  adult,  at  a  somewhat  earlier  age  than 
does  the  female.  With  other  species  of  rays  the  males  even  when 
old  never  possess  sharp  teeth,  and  consequently  the  adults  of  both 
sexes  are  provided  with  broad,  flat  teeth  like  those  of  the  young, 
and  like  those  of  the  mature  females  of  the  above-mentioned 
species.^^  As  the  rays  are  bold,  strong  and  voracious  fish,  wo 
may  suspect  that  the  males  require  their  sharp  teeth  for  fighting 
with  their  rivals ;  but  as  they  possess  many  parts  modified  and 
adapted  for  the  prehension  of  the  female,  it  is  possible  that  their 
teeth  may  be  used  for  this  purpose. 

In  regard  to  size,  M.  Carbonnier  "  maintains  that  the  female  of 
almost  all  fishes  is  larger  than  the  male ;  and  Dr.  Giinther  does 
not  know  of  a  single  instance  in  which  the  male  is  actually 
larger  than  the  female.  With  some  Cyprinodonts  the  male  is 
not  even  half  as  large.  As  in  many  kinds  of  fishes  the  males 
habitually  fight  together,  it  is  surprising  that  they  have  not 
generally  become  larger  and  stronger  than  the  females  through 
the  effects  of  sexual  selection.  The  males  suffer  from  their 
small  size,  for  according  to  M.  Carbonnier,  they  are  liable  to  be 
devoured  by  the  females  of  their  own  species  when  carnivorous, 
and  no  doubt  by  other  species.  Increased  size  must  be  in  some 
manner  of  more  importance  to  the  females,  than  strength  and 
size  are  to  the  males  for  fighting  with  other  males;  and  this 
perhaps  is  to  allow  of  the  production  of  a  vast  number  of  ova. 

In  many  species  the  male  alone  is  ornamented  with  bright 
colours;  or  these  are  much  brighter  in  the  male  than  the 
female.  The  male,  also,  is  sometimes  provided  with  appendages 
which  appear  to  be  of  no  more  use  to  him  for  the  ordinary 
purposes  of  life,  than  are  the  tail  feathers  to  the  peacock.  I  am 
indebted  for  most  of  the  following  facts  to  the  kindness  of  Dr. 
Giinther.  There  is  reason  to  suspect  that  many  tropical  fishes 
differ  sexually  in  colour  and  structure ;  and  there  are  some 
striking  cases  with  our  British  fishes.     The  male  Calliojiymns  lyra 

^•^  See    Yarrell's    account    of  the      cellent  figure,  and  p.  422,  432. 
rays  in  his '  Hist,  of  British  Fishes,'  ^^  As    quoted    in   'The    Fanner, 

vol.    ii.   1836,  p.  410,  with  an  ex-      1868,  p.  369. 


336 


The  Descent  of  Man. 


Part  II. 


has  been  called  the  gemmeoiis  dragonet  "  from  its  brilliant  gem- 
"  like  colours/'  When  fresh  caught  from  the  sea  the  body  is 
yellow  of  various  shades,  striped  and  spotted  with  vivid  blue  on 
the  head ;  the  dorsal  fins  are  pale  brown  with  dark  longitudinal 
bands;  the  ventral,  caudal,  and  anal  fins  being  bluish-black. 
The  female,  or  sordid  dragonet,  was  considered  by  Linnseus,  and 
by  many  subsequent  naturalists,  as  a  distinct  species ;  it  is  of  a 
dingy  reddish-brown,  with  the  dorsal  fin  brown  and  the  other 


Fig.  29.    Callionymus  lyra.    Upper  figure,  male ;  lower  figure,  female. 
N.B.  The  lower  figure  is  more  reduced  than  the  upper. 

fins  white.  The  sexes  differ  also  in  the  proportional  size  of  the 
head  and  mouth,  and  in  the  position  of  the  eyes  ;^^  but  the 
most  striking  difference  is  the  extraordinary  elongation  in  the 
male  (fig.  29)  of  the  dorsal  fin.  I\Ir.  W.  SaviUe  Kent  remarks 
that  this  "  singular  appendage  appears  from  my  observations 
"  of  the  species  in  confinement,  to  be  subservient  to  the  same 
"  end  as  the  wattles,  crests,  and  other  abnormal  adjuncts  of 
"  the  male  in  gallinaceous  birds,  for  the  jDurpose  of  fascinating 

'-  I  have  drawn  up  this  description  from  Yarrell's  '  British  Fishes,'voI.  1. 
1836,  pp.  261  and  2(36. 


Chap.  XII. 


Fishes. 


337 


**  their  mates."  ^^  The  young  males  resemble  the  adult  females 
in  structure  and  colour.  Throughout  the  genus  Callionymus/* 
the  male  is  generally  much  more  brightly  spotted  than  the 
female,  and  in  several  species,  not  only  the  dorsal,  but  the  anal 
fin  is  much  elongated  in  the  males. 

The  male  of  the  Coitm  scorpius,  or  sea-scorpion,  is  slenderer 
and  smaller  than  the  female.  There  is  also  a  great  difference 
in  colour  between  them.  It  is  difficult,  as  Mr.  Lloyd  ^^  remarks, 
"  for  any  one,  who  has  not  seen  this  fish  during  the  spawning- 
"  season,  when  its  hues  are  brightest,  to  conceive  the  admixture 
"  of  brilliaut  colours  with  which  it,  in  other  respects  so  ill- 
*'  favoured,  is  at  that  time  adorned."  Both  sexes  of  the  Lahrus 
mixtiis,  although  very  diiferent  in  colour,  are  beautiful;  the 
male  being  oraoge  with  bright  blue  stripes,  and  the  female 
bright  red  with  some  black  spots  on  the  back. 


Fig.  30.    Xiphophorus  Hellerii.    Upper  figure,  male ;  lower  figure,  female. 

In  the  very  distinci;  family  of  the  Cyprinodoutidse— inhabitants 
of  the  fresh  waters  of  foreign  lands— the  sexes  sometimes  differ 
much  in  various  characters.  In  the  male  of  the  MolUeuesia 
petenensis,'^^  the  dorsal  fin  is  greatly  developed  and  is  marked 

'6  With  respect  to  this  and  the 
following;  species  I  am  indebted  to 
Dr.  Giinther    for    information:  see 


'3  'Nature,'  July  1873,  p.  264. 

**  '  Catalogue  of  Acanth.  Fishes 
in  the  British  Museum,'  by  Dr. 
Giinther,  1861,  pp.  138-151. 

^^  *  Game  Birds  of  Sweden,'  &c.. 
1867,  p.  466. 


also  his  paper  on  the  'Fishes  of 
Central  America,'  in  'Transact. 
Zoolog.  Soc'  vol.  y\.  1868,  p.  485. 


338  The  Descent  of  Man.  -        Part  IT. 

with  a  row  of  large,  round,  ocellated,  bright-coloured  spots; 
whilst  the  same  fin  in  the  female  is  smaller,  of  a  different  shape, 
and  marked  only  with  irregularly  curved  brown  spots.  In  the 
male  the  basal  margin  of  the  anal  fin  is  also  a  little  produced 
and  dark  coloured.  In  the  male  of  an  allied  form,  the  Xipho- 
phorns  Helhrli  (fig.  30),  the  inferior  margin  of  the  caudal  fin  is 
developed  into  a  long  filament,  which,  as  I  hear  from  Dr.  Giinther, 
is  striped  with  bright  colours.  This  filament  does  not  contain 
any  muscles,  and  apparently  cannot  be  of  any  direct  use  to  the 
fish.  As  in  the  case  of  the  Callionymus,  the  males  whilst  young 
resemble  the  adult  females  in  colour  and  structure.  Sexual 
differences  such  as  these  may  be  strictly  compared  with  those 
which  are  so  frequent  with  gallinaceous  birds.^''' 

In  a  siluroid  fish,  inhabiting  the  fresh  waters  of  South  America, 
the  Plecostomus  harbatus^^  (fig.  31),  the  male  has  its  mouth  and 
inter-operculum  fringed  with  a  beard  of  stiff  hairs,  of  which  the 
female  shows  hardly  a  trace.  These  hairs  are  of  the  nature  of 
scales.  In  another  species  of  the  same  genus,  soft  flexible  ten- 
tacles project  from  the  front  part  of  the  head  of  the  male,  which 
are  absent  in  the  female.  These  tentacles  are  prolongations  of 
the  true  skin,  and  therefore  are  not  homologous  with  the  stiff 
hairs  of  the  former  species ;  but  it  can  hardly  be  doubted  that 
both  serve  the  same  purpose.  What  this  purpose  may  be,  it  is 
difficult  to  conjecture ;  ornament  does  not  here  seem  probable, 
but  we  can  hardly  suppose  that  stiff  hairs  and  flexible  filaments 
can  be  useful  in  any  ordinary  way  to  the  males  alone.  In  that 
strange  monster,  the  Chimcera  monsfrom,  the  male  has  a  hook- 
shaped  bone  on  the  top  of  the  head,  directed  forwards,  with  its 
end  rounded  and  covered  with  sharp  spines ;  in  the  female  "  this 
"  crown  is  altogether  absent,"  but  what  its  use  may  be  to  the 
male  is  utterly  unknown.^'' 

The  structures  as  yet  referred  to  are  permanent  in  the  male 
after  he  has  arrived  at  maturity  ;  but  with  some  Blennies,  and  in 
another  aUied  genus,^°  a  crest  is  developed  on  the  head  of  the 
male  only  during  the  breeding-season,  and  the  body  at  the  same 
time  becomes  more  brightly-coloured.  There  can  be  little  doubt 
that  this  crest  serves  as  a  temporary  sexunl  ornament,  for  the 
female  does  not  exhibit  a  trace  of  it.  In  other  species  of  the 
same  genus  both  sexes  possess  a  crest,  and  in  at  least  one  s^Decies 

'^  Dr.    Giinther    makes    this    re-  Water,'  July  1368,  p.  377,  with  a 

mark;  *  Catalogue  of  Fishes  in  the  figure.      Many  other  cases  could  be 

British  Museum,'  vol.  iii.  1861,  p.  added  of  structures  peculiar  to  the 

141.  male,    of   which    the    uses    are   not 

^*  See  Dr.  Giinther  on  this  genus,  known, 

m 'Proc.  Zoolog.  Soc' 1868,  p.  232.  =»  Dr.     Giinther,     '  Catalogue    of 

l^  F.    Buckland,    in     *  Land    and  Fishes,'  vol.  iii.  pp.  221  and  240. 


Chap.  XII. 


Fishes. 


339 


Fig.  31.  Plecostomus  barbatus.    L  i-p-n  i.b"' 


..3 ;  lower  figure,  female. 


340  The  Descent  of  Man.  Part  II. 

neither  sex  is  thus  provided.  In  many  of  the  Chromidse,  for 
instance  in  Geophagus  and  especially  in  Cichla,  the  males,  as  I 
hear  from  Professor  Agassiz/^  have  a  conspicuous  protuberance 
on  the  forehead,  which  is  wholly  wanting  in  the  females  and  in 
the  young  males.  Professor  Agassiz  adds,  "  I  have  often 
"  observed  these  fishes  at  the  time  of  spawning  when  the  ;pro- 
"  tuberance  is  largest,  and  at  other  seasons  when  it  is  totally 
*'  wanting,  and  the  two  sexes  shew  no  difference  whatever  in  the 
"  outline  of  the  profile  of  the  head.-  I  never  could  ascertain  that 
"  it  subserves  any  special  function,  and  the  Indians  on  the 
"  Amazon  know  nothing  about  its  use."  These  protuberances 
resemble,  in  their  periodical  appearance,  the  fleshy  caruncles  on 
the  heads  of  certain  birds ;  but  whether  they  serve  as  ornaments 
must  remain  at  present  doubtful. 

I  hear  from  Professor  Agassiz  and  Dr.  Giinther,  that  the  males 
of  those  fishes,  which  differ  permanently  in  colour  from  the 
females,  often  become  more  brilliant  during  the  breeding-season. 
This  is  likewise  the  case  with  a  multitude  of  fishes,  the  sexes  of 
which  are  identical  in  colour  at  all  other  seasons  of  the  year, 
The  tench,  roach,  and  perch  may  be  given  as  instances.  The 
male  salmon  at  this  season  is  "marked  on  the  cheeks  with 
*'■  orange-coloured  stripes,  which  give  it  the  ajDpearance  of  a 
**  Labrus,  and  the  body  partakes  of  a  golden  orange  tinge.  The 
"  females  are  dark  in  colour,  and  are  commonly  called  black- 
'*'fish."2^  An  analogous  and  even  greater  change  takes  place 
with  the  Salmo  eriox  or  bull  trout ;  the  males  of  the  char 
(iS.  umhla)  are  likewise  at  this  season  rather  brighter  in  colour 
than  the  females.^^  The  colours  of  the  pike  (Esox  reticulatus)  of 
the  United  States,  especially  of  the  male,  become,  during  the 
breeding-season,  exceedingly  intense,  brilliant,  and  iridescent.'^* 
Another  striking  instance  out  of  many  is  afforded  by  the  male 
stickleback  (Gasferosteus  leiurus),  which  is  described  by  Mr. 
Warington,^^  as  being  then  "beautiful  beyond  description." 
The  back  and  eyes  of  the  female  are  simply  brown,  and  the  belly 
white.  The  eyes  of  the  male,  on  the  other  hand,  are  "  of  the 
"  most  splendid  green,  having  a  metallic  lustre  like  the  green 
"  feathers  of  some  humming-birds.  The  throat  and  belly  are 
"  of  a  bright  crimson,  the  back  of  an  ashy-green,  and  the  whole 
"  fish  appears  as  though  it  were  somewhat  translucent  and 

2'  See  also  *  A  Journey  in  Brazil,'  Mag.  of  Nat.  History,'  vol.  vi.  1841, 

by  Prof,  and  Mrs.  Agassiz,  1868,  p.  p.  440. 
220.  24  'The  American  Agriculturalist,' 

"  Yarrell,  *  British   Fishes,'  vol.  1868,  p.  100. 
ii.  1836,  pp.  10,  12,  35.  25  .  Annals  and  Mag.  of  Nat.  Hist.' 

2^  W.  Thompson,  in  'Annals  and  Oct.  1852. 


Chap.  XII.  Fishes.  341 

"  glowed  with  an  internal  incandescence."  After  the  breeding- 
season  these  colours  all  change,  the  throat  and  belly  become  of  a 
paler  red,  the  back  more  green,  and  the  glowing  tints  subside. 

With  respect  to  the  courtship  of  fishes,  other  cases  have  been 
observed  since  the  first  edition  of  this  book  appeared,  besides  that 
already  given  of  the  stickleback.  Mr.  W.  S.  Kent  says  that  the 
male  of  the  Lalwus  mixtns,  which,  as  we  have  seen,  differs  in 
colour  from  the  female,  makes  "  a  deep  hollow  in  the  sand  of  the 
"  tank,  and  then  endeavours  in  the  most  persuasive  manner  to  in- 
"  duce  a  female  of  the  same  species  to  share  it  with  him,  swim- 
"  ming  backwards  and  forwards  between  her  and  the  completed 
''  nest,  and  x^lainly  exhibiting  the  greatest  anxiety  for  her  to  follow." 
The  males  of  Cantharus  lineatus  become,  during  the  breeding- 
season,  of  deep  leaden-black ;  they  then  retire  from  the  shoal,  and 
excavate  a  hollow  as  a  nest.  "Each  male  now  mounts  vigilant 
"  guard  over  his  respective  hollow,  and  vigorously  attacks  and 
*•  drives  away  any  other  fish  of  the  same  sex.  Towards  his  com- 
"  panions  of  the  opposite  sex  his  conduct  is  far  different ;  many  of 
"  the  latter  are  now  distended  with  spawn,  and  these  he  endeavours 
"  by  all  the  meaus  in  his  power  to  lure  singly  to  his  prepared 
"  hollow,  and  there  to  deposit  the  myriad  ova  with  which  they  are 
"  laden,  which  he  then  protects  and  guards  with  the  greatest  care.^" 

A  more  striking  case  of  courtship,  as  well  as  of  display,  by  the 
males  of  a  Chinese  Macropus  has  been  given  by  M.  Carbonnier, 
who  carefully  observed  these  fishes  under  confinement.-^  The 
males  are  most  beautifully  coloured,  more  so  than  the  females. 
During  the  breeding-season  they  contend  for  the  possession  of 
the  females;  and,  in  the  act  of  courtship,  expand  their  fins, 
which  are  spotted  and  ornamented  with  brightly  coloured  rays, 
in  the  same  manner,  according  to  M.  Carbonnier,  as  the  peacock. 
They  then  also  bound  about  the  females  with  much  vivacity,  and 
appear  by  "  I'etalage  de  leurs  vives  couleurs  chercher  a  attirer 
"  I'attention  des  femelles,  lesquelles  ne  paraissaient  indifferentes 
"  a  ce  manege,  elles  nageaient  avec  une  molle  lenteur  vers  les 
"  males  et  semblaient  se  comjDlaire  dans  leur  voisinage."  After 
the  male  has  won  his  bride,  he  makes  a  little  disc  of  froth  by 
blowing  air  and  mucus  out  of  his  mouth.  He  then  collects  the 
fertilised  ova,  drojDped  by  the  female,  in  his  mouth ;  and  this 
caused  M.  Carbonnier  much  alarm,  as  he  thought  that  they  were 
going  to  be  devoured.  But  the  male  soon  deposits  them  in  the 
disc  of  froth,  afterwards  guarding  them,  repairing  the  froth,  and 
taking  care  of  the  young  when  hatched.  I  mention  these  par- 
ticulars because,  as  we  shall  presently  see,  there  are  fishes,  the 

26  < Nature,'  May,  1873,  p.  25.  27  .Bull,  de  la  Soc.  d'Acclimat.' 

Paris,  July  1869,  and  Jan.  1870. 


342  The  Descent  of  Man.  Part  II. 

males  of  whicli  hatch  their  eggs  in  their  mouths  ;  and  those  who 
do  not  believe  in  the  principle  of  gradual  evolution  might  ask  how 
could  such  a  habit  have  originated ;  but  the  diflficulty  is  much 
diminished  when  we  know  that  there  are  fishes  which  thus 
collect  and  carry  the  eggs;  for  if  delayed  by  any  cause  in 
depositing  them,  the  habit  of  hatching  them  in  their  mouths 
might  have  been  acquired. 

To  return  to  our  more  immediate  subject.  The  case  stands 
thus :  female  fishes,  as  far  as  I  can  learn,  never  willingly  spawn 
except  in  the  presence  of  the  males ;  and  the  males  never  fertilise 
the  ova  except  in  the  presence  of  the  females.  The  males  fight 
for  the  possession  of  the  females.  In  many  species,  the  males 
whilst  young  resemble  the  females  in  colour ;  but  when  adult 
become  much  more  brilliant,  and  retain  their  colours  throughout 
life.  In  other  species  the  males  become  brighter  than  the  females 
and  otherwise  more  highly  ornamented,  only  during  the  season 
of  love.  The  males  sedulously  court  the  females,  and  in  one 
case,  as  we  have  seen,  take  pains  in  displaying  their  beauty 
before  them.  Can  it  be  believed  that  thej  would  thus  act  to  no 
purpose  during  their  courtship  ?  And  this  would  be  the  case, 
unless  the  females  exert  some  choice  and  select  those  males 
which  please  or  excite  them  most.  If  the  female  exerts  such 
choice,  all  the  above  facts  on  the  ornamentation  of  the  males 
become  at  once  intelligible  by  the  aid  of  sexual  selection. 

We  have  next  to  enquire  whether  this  view  of  the  bright 
colours  of  certain  male  fishes  having  been  acquired  through 
sexual  selection  can,  through  the  law  of  the  equal  transmission  of 
characters  to  both  sexes,  be  extended  to  those  groups  in  which  the 
males  and  females  are  brilliant  in  the  same,  or  nearly  the  same 
degree  and  manner.  In  such  a  genus  as  Labrus,  which  includes 
some  of  the  most  splendid  fishes  in  the  world — for  instance,  the 
Peacock  Labrus  {L.  -pavo),  described,-^  with  pardonable  exaggera- 
tion, as  formed  of  polished  scales  of  gold,  encrusting  lajDis-lazuli, 
rubies,  sapphires,  emeralds,  and  amethysts — we  may,  with  much 
probability,  accept  this  belief;  for  we  have  seen  that  the  sexes  in 
at  least  one  species  of  the  genus  differ  greatly  in  colour.  With 
some  fishes,  as  with  many  of  the  lowest  animals,  splendid  colours 
may  be  the  direct  result  of  the  nature  of  their  tissues  and  of  the 
surrounding  conditions,  without  the  aid  of  selection  of  any  kind. 
The  gold-fish  {Cyprinus  auratus),  judging  from  the  analogy  of 
the  golden  variety  of  the  common  carp,  is  perhaps  a  case  in  point, 
as  it  may  owe  its  splendid  colours  to  a  single  abrupt  variation, 
due  to  the  conditions  to  which  this  fish  has  been  subjected  under 

28  Bory  de  Saint  Vincent,    in  'Diet.  Class.  d'Hist.  Nat.'  torn.  ix.  1826 
p.  151. 


Chap.  XII.  Fishes.  343 


confinement.  It  is,  however,  more  probable  that  these  colours 
have  been  intensified  through  artificial  selection,  as  this  species 
has  been  carefully  bred  in  China  from  a  remote  period.^^  Under 
natural  conditions  it  does  not  seem  probable  that  beings  so 
highly  organised  as  fishes,  and  which  live  under  such  complex 
relations,  should  become  brilliantly  coloured  without  suffering 
some  evil  or  receiving  some  benefit  from  so  great  a  change,  and 
consequently  without  the  intervention  of  natural  selection. 

What,  then,  are  we  to  conclude  in  regard  to  the  many  fishes, 
both  sexes  of  which  are  splendidly  coloured?  Mr.  Wallace ^'^ 
believes  that  the  species  which  frequent  reefs,  where  corals  and 
other  brightly-coloured  organisms  abound,  are  brightly  coloured 
in  order  to  escape  detection  by  their  enemies ;  but  according  to 
my  recollection  they  were  thus  rendered  highly  conspicuous. 
In  the  fresh-waters  of  the  tropics  there  are  no  brilliantly- 
coloured  corals  or  other  organisms  for  the  fishes  to  resemble ; 
yet  many  species  in  the  Amazons  are  beautifully  coloured,  and 
many  of  the  carnivorous  Cyprinidse  in  India  are  ornamented 
with  "bright  longitudinal  lines  of  various  tints." ^^  Mr.  M'Clel- 
land,  in  describing  these  fishes,  goes  so  far  as  to  suppose  that 
"  the  peculiar  brilliancy  of  their  colours  "  serves  as  "  a  better 
"  mark  for  king-fishers,  terns,  and  other  birds  which  are 
"  destined  to  keep  the  number  of  these  fishes  in  check ; "  but  at 
the  present  day  few  naturalists  will  admit  that  any  animal  has 
been  made  conspicuous  as  an  aid  to  its  own  destruction.  It  is 
possible  that  certain  fishes  may  have  been  rendered  conspicuous 
in  order  to  warn  birds  and  beasts  of  prey  that  they  were 
unpalatable,  as  explained  when  treating  of  caterpillars;  but  it 
is  not,  I  believe,  known  that  any  fish,  at  least  any  fresh-water 
fish,  is  rejected  from  being  distasteful  to  fish-devouring  animals. 
On  the  whole,  the  most  probable  view  in  regard  to  the  fishes,  of 
which  both  sexes  are  brilliantly  coloured,  is  that  their  colours 
were  acquired  by  the  males  as  a  sexual  ornament,  and  were 
transferred  equally,  or  nearly  so,  to  the  other  sex. 

"^  Owing  to  some  remai*ks  on  this  has  been  "  produced  at  Hangchow  a 

subject,  made  in  my  work  '  On  the  "  variety  called  the  fire-fish,  from  its 

Variation  of  Animals  under  Domesti-  "intensely  red    colour.     It   is    uni- 

cation,'  Mr.  W.  F.  Mayers  ('  Chinese  "  versally  admired,  and  there  is  not 

Notes  and   Queries,'  Aug.   1868,  p.  "  a  household  where  it   is  not  cul- 

123)     has     searched     the     ancient  "  tivated,  w  rivalry  as  to  its  colour, 

(.•hinese     encyclopedias.     He     finds  "  and  as  a  source  of  profit." 

that    gold-fish  were  first  reared  iu  ^^  *  Westminster     Review,'     July 

confinement   during  the   Sung   Dy-  1867,  p.  7. 

nasty,  which  commenced  A.D.  960.  ^i  <  Judian  Cyprinidaj,'  by  Mr.   J, 

In    the     year     1129     these     fishes  M'Clelland,     'Asiatic      Researches, 

abounded.     In   another   place  it   is  vol.  xix.  part  ii.  1839,  p.  230. 
said  that  since  the  year  1548  there 


344  The  Descent  of  Ma7L  Part  II. 


We  have  now  to  consider  wliether,  when  the  male  differs  in  a 
marked  manner  from  the  female  in  colour  or  in  other  orna- 
ments, he  alone  has  been  modified,  the  variations  being  inherited 
by  his  male  offspring  alone;  or  whether  the  female  has  been 
specially  modified  and  rendered  inconspicuous  for  the  sake  of 
jn-otection,  such  modifications  being  inherited  only  by  the 
females.  It  is  impossible  to  doubt  that  colour  has  been  gained 
by  many  fishes  as  a  protection :  no  one  can  examine  the  speckled 
upper  surface  of  a  flounder,  and  overlook  its  resemblance  to  the 
sandy  bed  of  the  sea  on  which  it  lives.  Certain  fishes,  moreover, 
can  through  the  action  of  the  nervous  system,  change  their 
colours  in  adaptation  to  surrounding  objects,  and  that  within  a 
short  time.^^  One  of  the  most  striking  instances  ever  recorded 
of  an  animal  being  protected  by  its  colour  (as  far  as  it  can  be 
judged  of  in  preserved  specimens),  as  well  as  by  its  form,  is  that 
given  by  Dr.  Giinther^^  of  a  pipe-fish,  which,  with  its  reddish 
streaming  filaments,  is  hardly  distinguishable  from  the  sea-weed 
to  which  it  clings  with  its  prehensile  tail.  But  the  question  now 
imder  consideration  is  whether  the  females  alone  have  been 
modified  for  this  object.  We  can  see  that  one  sex  will  not  be 
modified  through  natural  selection  for  the  sake  of  protection 
more  than  the  other,  supposing  both  to  vary,  unless  one  sex  is 
exposed  for  a  longer  period  to  danger,  or  has  less  power  of 
escaping  from  such  danger  than  the  other;  and  it  does  not 
appear  that  with  fishes  the  sexes  differ  in  these  respects.  As 
far  as  there  is  any  difference,  the  males,  from  being  generally 
smaller  and  from  wandering  more  about,  are  exposed  to  greater 
danger  than  the  females;  and  yet,  when  the  sexes  differ,  the 
males  are  almost  always  the  more  conspicuously  coloured. 
The  ova  are  fertilised  immediately  after  being  deposited ;  and 
when  this  process  lasts  for  several  days,  as  in  the  case  of 
the  salmon,^*  the  female,  during  the  whole  time,  is  attended  by 
the  male.  After  the  ova  are  fertilised  they  are,  in  most  cases, 
left  unprotected  by  both  parents,  so  that  the  males  and  females, 
as  far  as  oviposition  is  concerned,  are  equally  exposed  to  danger, 
and  both  are  equally  important  for  the  production  of  fertile  ova ; 
consequently  the  more  or  less  brightly-coloured  individuals  of 
either  sex  would  be  equally  liable  to  be  destroyed  or  preserved, 
and  both  would  have  an  equal  influence  on  the  colours  of  their 
offspring. 

Certain  fishes,  belonging  to  several  families,  make  nests,  and 
some  of  them  take  care  of  their  young  when  hatched.     Both 

32  G.  Pouchet,  L'Institut.  Nov.  1,      327,  pi.  xiv.  anci  xv. 

1871,  p.  134.  ='•'  Yarrell,  '  British   Fishes,'  rcl. 

33  'Proc.    Zoolog.   Soc'  1865,  p.      ii.  p.  11. 


Chap.  XII.  Fishes.  345 


sexes  of  the  bright  coloured  Crenilabrus  massa  and  mdopa  work 
together  in  building  their  nests  with  sea-weed,  shells,  &c.^^ 
But  the  males  of  certain  fishes  do  all  the  work,  and  afterwards 
take  exclusive  charge  of  the  young.  This  is  the  case  with  the  dull- 
coloured  gobies,^''  in  which  the  sexes  are  not  known  to  differ  in 
colour,  and  likewise  with  the  sticklebacks  (Gasterosteus),  in  which 
the  males  become  brilliantly  coloured  during  the  spawning  season. 
The  male  of  the  smooth- tailed  stickleback  ((?.  leinrus)  j^erforms 
the  duties  of  a  nurse  with  exemplary  care  and  vigilance  during 
a  long  time,  and  is  continually  employed  in  gently  leading  back 
the  young  to  the  nest,  when  they  stray  too  far.  He  courageously 
drives  away  all  enemies,  including  the  females  of  his  own  species. 
It  would  indeed  be  no  small  relief  to  the  male,  if  the  female,  after 
depositing  her  eggs,  were  immediately  devoured  by  some  enemy, 
for  he  is  forced  incessantly  to  drive  her  from  the  nest.^^ 

The  males  of  certain  other  fishes  inhabiting  South  America 
and  Ceylon,  belonging  to  two  distinct  Orders,  have  the  extra- 
ordinary habit  of  hatching  within  their  mouths  or  branchial 
cavities,  the  eggs  laid  by  the  females.^®  I  am  informed  by 
Professor  Agassiz  that  the  males  of  the  Amazoniaa  species 
which  follow  this  habit,  "  not  only  are  generally  brighter  than 
"  the  females,  but  the  difference  is  greater  at  the  spawning-season 
"  than  at  any  other  time."  The  species  of  Geophagus  act  in  the 
same  manner ;  and  in  this  genus,  a  conspicuous  protuberance 
becomes  developed  on  the  forehead  of  the  males  during  the 
breeding-season.  With  the  various  species  of  Chromids,  as 
Professor  Agassiz  likewise  informs  me,  sexual  differences  in 
colour  may  be  observed,  "  whether  they  lay  their  eggs  in  the 
"  water  among  aquatic  plants,  or  deposit  them  in  holes,  leaving 
"  them  to  come  out  witiiout  further  care,  or  build  shallow  nests 
"  in  the  river  mud,  over  which  they  sit,  as  our  Promotis  does. 
"  It  ought  also  to  be  observed  that  these  sitters  are  among  the 
"  brightest  species  in  their  respective  families ;  for  instance, 
"  Hygrogonus  is  bright  green,  with  large  black  ocelli,  encircled 
"  with  the  most  brilliant  red."  Whether  with  all  the  species  of 
Chromids  it  is  the  male  alone  which  sits  on  the  eggs  is  not 
known.    It  is,  however,  manifest  that  the  fact  of  the  eggs  being 

^^  According  to  the  observatioas  nals  and  Mag.  of  Nat.  Hist.'  Novem- 

of  M.  Gerbe  ;  see  Giinther's  '  Record  ber  1855. 

of    Zoolog.    Literature,'     1^65,    p.  '*  Prof.  Wyman,  in  '  Proc*.  Boston 

194-.  Soc.  of  Nat.  Hist.'  Sept.    15,  1857. 

2^  Cuvier,   '  Rfegne   Animal,'   vol.  Also   Prof.  Turner,   in   '  Journal  of 

ii.  1829,  p.  242.  Anatomy  and   Phys.'  Nov.   1,   186(3, 

"  See     Mr.     Waringtou's     most  p.  78.     Dr.    Giinther    has    likewise 

interesting  description  of  the  habits  described  other  cases, 
of  the  Gasterosteus  Iciurus,   in  'An- 

16 


34^  The  Descent  of  Man.  Part  II. 

protected  or  unprotected  by  the  parents,  has  had  little  or  no 
influence  on  the  differences  in  colour  between  the  sexes.  It  is 
further  manifest,  in  all  the  cases  in  which  the  males  take 
exclusive  charge  of  the  nests  and  young,  that  the  destruction 
of  the  blighter-coloured  males  would  be  far  more  influential  on 
the  character  of  the  race,  than  the  destruction  of  the  brighter- 
coloured  females ;  for  the  death  of  the  male  during  the  period  of 
incubation  or  nursing  would  entail  the  death  of  the  young,  so 
that  they  could  not  inherit  his  peculiarities ;  yet,  in  many  of 
these  very  cases  the  males  are  more  conspicuously  coloured  than 
the  females. 

In  most  of  the  Lophobranchii  (Pipe-fish,  Hippocampi,  &c.) 
the  males  have  either  marsupial  sacks  or  hemispherical  de- 
pressions on  the  abdomen,  in  which  the  ova  laid  by  the  female 
are  hatched.  The  males  also  shew  great  attachment  to  their 
young.^^  The  sexes  do  not  commonly  differ  much  in  colour; 
but  Dr.  Gunther  believes  that  the  male  Hippocampi  are  rather 
brighter  than  the  females.  The  genus  Solenostoma,  however, 
offers  a  curious  exceptional  case,'*''  for  the  female  is  much  more 
vividly-coloured  and  spotted  than  the  male,  and  she  alone  has  a 
marsupial  sack  and  hatches  the  eggs;  so  that  the  female  of 
Solenostoma  differs  from  all  the  other  Lophobranchii  in  this 
latter  respect,  and  from  almost  all  other  fishes,  in  being  more 
brightly-coloured  than  the  male.  It  is  improbable  that  this 
remarkable  double  inversion  of  character  in  the  female  should 
be  an  accidental  coincidence.  As  the  males  of  several  fishes, 
which  take  exclusive  charge  of  the  eggs  and  young,  are  more 
brightly  coloured  than  the  females,  and  as  here  the  female  Sole- 
nostoma takes  the  same  charge  and  is  brighter  than  the  male,  it 
might  be  argued  that  the  conspicuous  colours  of  that  sex  which 
is  the  more  important  of  the  two  for  the  welfare  of  the  offspring, 
must  be  in  some  manner  protective.  But  from  the  large  number 
of  fishes,  of  which  the  males  are^  either  permanently  or  period- 
ically brighter  than  the  females,  but  whose  life  is  not  at  all 
more  important  for  the  welfare  of  the  species  than  that  of  the 
female,  this  view  can  hardly  be  maintained.  When  we  treat 
of  birds  we  shall  meet  with  analogous  cases,  where  there  has 
been  a  complete  inversion  of  the  usual  attributes  of  the  two 
sexes,  and  we  shall  then  give  what  appears  to  be  the  probable 
explanation,  namely,  that  the  males  have  selected  the  more 
attractive  females,  instead  of  the  latter  having  selected,  in 

39    Yarrell,     '  Hist,     of     British  Fishes  of  Zanzibar,'  by  Col.  Playfair, 

Fishes,'  vol.  ii.  183G,  pp.  329,  338.  1866,  p.  137,  has   re-examined  the 

*"  Dr.  Gunther,  since  publishing  specimens,   and    has   given   me    the 

an  account  of  this  species  in  '  The  above  information. 


Chap.  XII.  Fishes.  347 


accordance  with  the  usual  rule  throughout  the  animal  kingdom, 
the  more  attractive  males. 

On  the  whole  we  may  conclude,  that  with  most  fishes,  in 
which  the  sexes  differ  in  colour  or  in  other  ornamental  charac- 
ters, the  males  originally  varied,  with  their  variations  trans- 
mitted to  the  same  sex,  and  accumulated  through  sexual 
selection  by  attracting  or  exciting  the  females.  In  many  cases, 
however,  such  characters  have  been  transferred,  either  partially 
or  completely,  to  the  females.  In  other  cases,  again,  both  sexes 
have  been  coloured  alike  for  the  sake  of  protection;  but  in 
no  instance  does  it  appear  that  the  female  alone  has  had  her 
colours  or  other  characters  specially  modified  for  this  latter 
purpose. 

The  last  point  which  need  be  noticed  is  that  fishes  are  known 
to  make  various  noises,  some  of  which  are  described  as  being 
musical.  Dr.  Dnfosse,  who  has  especially  attended  to  this 
subject,  says  that  the  sounds  are  voluntarily  produced  in  several 
ways  by  different  fishes :  by  the  friction  of  the  pharyngeal  bones 
— by  the  vibration  of  certain  muscles  attached  to  the  swim- 
bladder,  which  serves  as  a  resounding  board — and  by  the  vibra- 
tion of  the  intrinsic  muscles  of  the  swim-bladder.  By  this  latter 
means  the  Trigla  produces  pure  and  long-drawn  sounds  which 
range  over  nearly  an  octave.  But  the  most  interesting  case  for 
us  is  that  of  two  species  of  Ophidium,  in  which  the  males  alone 
are  provided  with  a  sound-producing  apparatus,  consisting  of 
small  movable  bones,  with  proper  muscles,  in  connection  with 
the  swim-bladder,*^  The  drumming  of  the  Umbrinas  in  the 
European  seas  is  said  to  be  audible  from  a  depth  of  twenty 
fathoms ;  and  the  fishermen  of  Rochelle  assert  "  that  the  males 
"  alone  make  the  noise  during  the  spawning-time ;  and  that  it 
"  is  possible  by  imitating  it,  to  take  them  without  bait."*^  From 
this  statement,  and  more  especially  from  the  case  of  Ophidium, 
it  is  almost  certain  that  in  this,  the  lowest  class  of  the  Verte- 
brata,  as  with  so  many  insects  and  spiders,  sound-producing 
instruments  have,  at  least  in  some  cases,  been  developed  through 
sexual  selection,  as  a  means  for  bringing  the  sexes  together. 

*'  '  Comptes  Rendus.'     Tom.  xlvi.  the  Dutch  translation  of  this  w  ork 

1858,  p.  353.     Tom.  xlvii.   1858,  p.  (vol.  ii.,  p.  36),  gives  some  further 

916.     Tom.  liv.  1862,  p.  393.     The  particulars  on  the  sounds  made  by 

noise  made  by  the  Umbrinas  {Sciccna  lishes. 

ciqvMa),  is  said  by  some  authors  to  *^  The     Rev.     C.     Kingslcy,     in 

be  more  like  that  of  a  flute  or  organ,  <  Nature,'  May  1870,  p.  40. 
than  drumming :   Dr.  Zouteveen,  in 


343 


The  Descent  of  Man. 


Part  II. 


Amphibians. 

Urodela. — I  will  begin  with  the  tailed  amphibians.  The  sexes 
of  salamanders  or  newts  often  diifer  much  both  in  colour  and 
structure.  In  some  species  prehensile  claws  are  develoiDed  on 
the  fore-legs  of  the  males  during  the  breeding-season :  and  at 
this  season  in  the  male  Triton  palmipes  the  hind-feet  are  pro- 
vided with  a  swimming-web,  which  is  almost  completely 
absorbed  during  the  winter;   so  that  their  feet  then  resemble 


Fig.  32.    Triton  cristatus  (half  natural  size,  from  Bell's  '  British  Reptiles  '). 
Upper  figure,  male  during  the  breeding-season ;  lower  figure,  female. 


those  of  the  female.^^  This  structure  no  doubt  aids  the  male 
in  his  eager  search  and  pursuit  of  the  female.  Whilst  courting 
her  he  rapidly  vibrates  the  end  of  his  tail.  With  our  common 
newts  (Trito7i  pur/ctntus  and  cristntus)  a  deep,  much  indented 
crest  is  developed  along  the  back  and  tail  of  the  male  during  the 
breeding-season,  which  disappears  during  the  winter.  Mr.  St. 
George  Mivart  informs  me  that  it  is  not  furnished  with  muscles, 
and  therefore  cannot  be  used  for  locomotion.  As  during  the 
season  of  courtship  it  becomes  edged  with  bright  colours,  there 
can  hardly  be  a  doubt  that  it  is  a  masculine  ornament.  In 
many  species  the  body  presents  strongly  contrasted,  though 
lurid  tints,  and  these  become  more  vivid  during  the  breeding- 
season.  The  male,  for  instance,  of  our  common  little  newt 
{Triton  punctatus)  is  "brownish-grey  above,  passing  into  yellow 

"  Bell,  'History  of  British  Reptiles,'  2nd  edit.  1849,  pp.  156-159. 


Chap.  XII.  Amphibians.  349 

"  beneath,  wliicli  in  the  spring  becomes  a  rich  bright  orange, 
"  marked  everywhere  with  round  dark  si3ots."  The  edge  of  the 
crest  also  is  then  tipped  with  bright  red  or  violet.  The  female 
is  usually  of  a  yellowish-brown  colour  with  scattered  brown 
dots,  and  the  lower  surface  is  often  quite  plain.*^  The  young 
are  obscurely  tinted.  The  ova  are  fertilised  during  the  act  of 
deposition,  and  are  not  subsequently  tended  by  either  parent. 
We  may  therefore  conclude  that  the  males  have  acquired  their 
strongly-marked  colours  and  ornamental  apjiendages  through 
sexual  selection ;  these  being  transmitted  either  to  the  male 
offspring  alone,  or  to  both  sexes. 

Anura  or  But  rack  la. — With  many  frogs  and  toads  the  colours 
evidently  serve  as  a  protection,  such  as  the  bright  green  tints 
of  tree-frogs  and  the  obscure  mottled  shades  of  many  terrestrial 
species.  The  most  conspicuously-coloured  toad  which  I  ever 
saw,  the  rhri/niscus  nigricans,*^  had  the  whole  ui^per  surface  of 
the  body  as  black  as  ink,  with  the  soles  of  the  feet  and  j^arts  of . 
the  abdomen  spotted  with  the  brightest  vermilion.  It  crawled 
about  the  bare  sandy  or  open  grassy  plains  of  La  Plata  under  a 
scorching  sun,  and  could  not  fail  to  catch  the  eye  of  every  pass- 
ing creature.  These  colours  are  probably  beneficial  by  making 
this  animal  known  to  all  birds  of  prey  as  a  nauseous  mouthful. 

In  Nicaragua  there  is  a  little  frog  "  dressed  in  a  bright  livery 
"  of  red  and  blue  "  which  does  not  conceal  itself  like  most  other 
species,  but  hops  about  during  the  daytime,  and  Mr.  Belt  says""* 
that  as  soon  as  he  saw  its  happy  sense  of  security,  he  felt  sure 
that  it  was  uneatable. "  After  several  trials  he  succeeded  in 
tempting  a  young  duck  to  snatch  up  a  young  one,  but  it  was 
instantly  rejected ;  and  the  duck  "  went  about  jerking  its  head, 
"  as  if  trying  to  throw  off  some  unpleasant  taste." 

With  respect  to  sexual  differences  of  colour.  Dr.  Giinther 
does  not  know  of  any  striking  instance  either  with  frogs  or 
toads ;  yet  he  can  often  distinguish  the  male  from  the  female,  by 
the  tints  of  the  former  being  a  little  more  intense.  Nor  does 
he  know  of  any  striking  difference  in  external  structure  between 
the  sexes,  excepting  the  prominences  which  become  developed 
during  the  breeding-season  on  the  front-legs  of  the  male,  by 
which  he  is  enabled  to  hold  the  female.'*^    It  is  surprising  that 

*^  Bell,  *  History  of  British  Rep-  sikrmmensis   (Dr.   Anderson,    *  Proc. 

tiles,'  2nd  edit.  1849,  pp.  146,  151.  Zoolog.  Soc.,'  1871,  p.  204)  has  two 

*^  'Zoology  of  the  Voyage  of  the  plate-like  callosities  on  the  thorax 

"  Beagle,"  '  1843.     Bell,  ibid.  p.  49.  and  certain  rugosities  on  the  fingers, 

***  '  The  Naturalist  in  Nicaragua,'  which  perhaps  subserve  the  same  end 

1874,  p.  321.  as  the  above-mentioned  prominences. 

*''  The   male  alone   of  the  Bufo 


350  The  Descent  of  Man.  Part  II. 

these  animals  have  not  acquired  more  strongly-marked  sexual 
characters ;  for  though  cold-blooded  their  passions  are  strong. 
Dr.  GUnther  informs  me  that  he  has  several  times  found  an 
unfortunate  female  toad  dead  and  smothered  from  having  been 
so  closely  embraced  by  three  or  four  males.  Frogs  have  been 
observed  by  Professor  Hoffman  in  Giessen  fighting  all  day  long 
during  the  breeding-season,  and  with  so  much  violence,  that  one 
had  its  body  ripped  open. 

Frogs  and  toads  offer  one  interesting  sexual  difference,  namely, 
in  the  musical  powers  possessed  by  the  males;  but  to  speak 
of  music,  when  applied  to  the  discordant  and  overwhelming 
sounds  emitted  by  male  bull-frogs  and  some  other  species,  seems, 
according  to  our  taste,  a  singularly  inappropriate  expression. 
Nevertheless,  certain  frogs  sing  in  a  decidedly  pleasing  manner. 
Near  Eio  Janeiro  I  used  often  to  sit  in  the  evening  to  listen  to  a 
number  of  little  Hylse,  perched  on  blades  of  grass  close  to  the 
water,  which  sent  forth  sweet  chirping  notes  in  harmony.  The 
various  sounds  are  emitted  chiefly  by  the  males  during  the 
breeding-season,  as  in  the  case  of  the  croaking  of  our  common 
frog.*^  In  accordance  with  this  fact  the  vocal  organs  of  the 
males  are  more  highly-develojDed  than  those  of  the  females.  In 
some  genera  the  males  alone  are  provided  with  sacs  which  open 
into  the  larynx.^^  For  instance,  in  the  edible  frog  {liana  escultnta) 
"  the  sacs  are  peculiar  to  the  males,  and  become,  when  filled 
"  with  air  in  the  act  of  croaking,  large  globular  bladders,  stand- 
"  ing  out  one  on  each  side  of  the  head,  near  the  corners  of  the 
"  mouth."  The  croak  of  the  male  is  thus  rendered  exceedingly 
powerful ;  whilst  that  of  the  female  is  only  a  slight  groaning 
noise.^"  In  the  several  genera  of  the  family  the  vocal  organs 
differ  considerably  in  structure,  and  their  development  in  all 
cases  may  be  attributed  to  sexual  selection. 

Eeptiles. 

Chelonia. — Tortoises  and  turtles  do  not  offer  well-marked 
sexual  differences.  In  some  species,  the  tail  of  the  male  is 
longer  than  that  of  the  female.  In  some,  the  plastron  or  lower 
surface  of  the  shell  of  the  male  is  slightly  concave  in  relation  to 
the  back  of  -  the  female.  The  male  of  the  mud-turtle  of  the 
United  States  (Chrysemys  picta)  has  claws  on  its  front-feet 
twice  as  long  as  those  of  the  female ;  and  these  are  used  when 

*8  Bell,      'History     of     British  "  J.  Bishop,  in  'Todd's  Cyclop. 

Reptiles,'  1849,  p.  93.  of  Anat.  and  Phvs.'  vol.  iv.  p.'l503 

*o  Bell,  ibid.  p.  112-114. 


Chap.  XII.  Reptiles.  351 

the  sexes  unite.^^  With  the  huge  tortoise  of  the  Galapagos 
Islands  {Tedudo  nigra)  the  males  are  said  to  grow  to  a  larger 
size  than  the  females:  during  the  pairing-season,  and  at  no 
other  time,  the  male  utters  a  hoarse  bellowing  noise,  which  can 
be  heard  at  the  distance  of  more  than  a  hundred  yards ;  the 
female,  on  the  other  hand,  never  uses  her  voice.^^ 

With  the  Testudo  degans  of  India,  it  is  said  "  that  the  combats 
"  of  the  males  may  be  heard  at  some  distance,  from  the  noise 
"  they  produce  in  butting  against  each  other."  ^^ 

Crocodilia. — The  sexes  apparently  do  not  differ  in  colour ;  nor 
do  I  know  that  the  males  fight  together,  though  this  is  pro- 
bable, for  some  kinds  make  a  prodigious  display  before  the 
females.  Bartram^*  describes  the  male  alligator  as  striving 
to  win  the  female  by  splashing  and  roaring  in  the  midst 
of  a  lagoon,  *'  swollen  to  an  extent  ready  to  burst,  with  its 
"  head  and  tail  lifted  up,  he  spins  or  twirls  round  on  the 
"  surface  of  the  water,  like  an  Indian  chief  rehearsing  his  feats 
of  war."  During  the  season  of  love,  a  musky  odour  is  emitted 
by  the  submaxillary  glands  of  the  crocodile,  and  pervades  their 
haunts.^^ 

Ophidia.—'Dr.  Giinther  informs  me  that  the  males  are  always 
smaller  than  the  females,  and  generally  have  longer  and  slenderer 
tails ;  but  he  knows  of  no  other  difference  in  external  structure. 
In  regard  to  colour,  he  can  almost  always  distinguish  the  male 
from  the  female  by  his  more  strongly-pronounced  tints ;  thus 
the  black  zigzag  band  on  the  back  of  the  male  English  viper  is 
more  distinctly  defined  than  in  the  female.  The  difference  is 
much  plainer  in  the  rattle-snakes  of  N.  America,  the  male  of 
which,  as  the  keeper  in  the  Zoological  Gardens  shewed  me,  can  at 
once  be  distinguished  from  the  female  by  having  more  lurid 
yellow  about  its  whole  body.  In  S.  Africa  the  Bucephalus 
mpensis  presents  an  analogous  difference,  for  the  female  "is 
"  never  so  fully  variegated  with  yellow  on  the  sides  as  the 
"  male."  ^^  The  male  of  the  Indian  Dipsas  ajnodon,  on  the 
other  hand,  is  blackish-brown,  with  the  belly  partly  black, 
whilst  the  female  is  reddish  or  yellowish-olive,  with  the  belly 
either  uniform  yellowish  or  marbled  with  black.  In  the  Tragops 
dispar  of  the  same  country,  the  male  is  bright  green,  and  the 

51  Mr.     C.     J.     Maynard,     'The  British  India,' 1864,  p.  7. 
American  Naturalist,'  Dec.  1869,  p.  ^*  '  Travels     through     Carolina, 

555.  &c.,  1791,  p.  128. 

*2  See  my  <  Journal  of  Researches  ^^  Owen,    *  Anatomy    of    Verte- 

during  the  Voyage  of  the  "  Beagle,"  '  bratcs,'  vol.  i.  1866,  p.  615. 
1845,  p.  384.  5«  Sir  Andrew  Smith,  '  Zoolo?.  of 

53  Dr.      Giinther,     'Reptiles     of  S.  Africa:  Reptilia,' 1849,  pi.  xt 


352  The  Descent  of  Man.  Part  II. 


female  bronze-coloured.^''  No  doubt  the  colours  of  some  snakes 
are  protective,  as  shewn  by  the  green  tints  of  tree-snakes,  and 
the  various  mottled  shades  of  the  species  which  live  in  sandy 
places ;  but  it  is  doubtful  w^iether  the  colours  of  many  kinds, 
for  instance  of  the  common  English  snake  and  viper,  serve  to 
conceal  them;  and  this  is  still  more  doubtful  with  the  many 
foreign  species  which  are  coloured  with  extreme  elegance.  The 
colours  of  certain  species  are  very  different  in  the  adult  and 
young  states.^^ 

During  the  breeding-season  the  anal  scent-glands  of  snakes  are 
in  active  function  f^  and  so  it  is  with  the  same  glands  in  lizards, 
and  as  we  have  seen  with  the  submaxillary  glands  of  crocodiles. 
As  the  males  of  most  animals  search  for  the  females,  these 
odoriferous  glands  probably  serve  to  excite  or  charm  the  female, 
rather  than  to  guide  her  to  the  spot  where  the  male  may  be 
found.  Male  snakes,  though  appearing  so  sluggish,  are  amorous  ; 
for  many  have  been  observed  crowding  round  the  same  female, 
and  even  round  her  dead  body.  They  are  not  known  to 
fight  together  from  rivalry.  Their  intellectual  powers  are 
higher  than  might  have  been  anticipated.  In  the  Zoological 
Gardens  they  soon  learn  not  to  strike  at  the  iron  bar  with  which 
their  cages  are  cleaned ;  and  Dr.  Keen  of  Philadelphia  informs 
me  that  some  snakes  which  he  kept,  learned  after  four  or  five 
times  to  avoid  a  noose,  with  which  they  were  at  first  easily 
caught.  An  excellent  observer  in  Ceylon,  Mr.  E.  Layard,  saw  '^^ 
a  cobra  thrust  its  head  through  a  narrow  hole  and  sw^allow  a 
toad.  "  With  this  encumbrance  he  could  not  withdraw  him- 
"  self;  finding  this,  he  reluctantly  disgorged  the  precious  mor- 
"  sel,  which  began  to  move  off;  this  was  too  much  for  snake 
"  philosophy  to  bear,  and  the  toad  was  again  seized,  and  again 
"  was  the  snake,  after  violent  efforts  to  escape,  compelled  to  part 
"  with  its  prey.  This  time,  however,  a  lesson  had  been  learnt, 
"  and  the  toad  w^as  seized  by  one  leg,  withdrawn,  and  then 
"  swallowed  in  triumph." 

The  keeper  in  the  Zoological  Gardens  is  positive  that  certain 
snakes,  for  instance  Crotalus  and  Python,  distinguish  him  from 
all  other  persons.  Cobras  kej)t  together  in  the  same  cage 
apparently  feel  some  attachment  towards  each  other.*^^ 

"  Dr.  A.   Giinther,   'Reptiles   of  brates,' vol.  i.  1866,  p.  615. 

British    India/  Ray  Soc.   1864,  pp.  «<>  '  Rambles  in  Cevlon.' in 'Anuals 

304,  308.  and  Mag.  of  Nat.  Hist.'  2nd  series, 

58    Dr.    Stoliczka,    '  Journal     of  vol.  ix.  1852,  p.  333. 

Asiatic  Soc.  of  Bengal,'  vol.  xxxix.  **!  Dr.     Giinther,     *  Reptiles      of 

1870,  pp.  205,  211.  British  India,'  1864,  p.  340. 

^^  Owen,     '  Anatomy    of    Verte- 


Chap.  XII.  Reptiles.  353 


It  does  not,  however,  follow  because  snakes  have  some 
reasoning:  power,  strong  passions  and  mutual  affection,  that  they 
should  likewise  be  endowed  with  sulhcient  taste  to  admire 
brilliant  colours  in  their  partners,  so  as  to  lead  to  the  adorn- 
ment of  the  species  through  sexual  selection.  Nevertheless,  it  is 
difficult  to  account  in  any  other  manner  for  the  extreme  beauty 
of  certain  species ;  for  instance,  of  the  coral-snakes  of  S.  America, 
which  are  of  a  rich  red  with  black  and  yellow  transverse  bands. 
I  well  remember  how  much  surprise  I  felt  at  the  beauty  of  the 
first  coral-snake  which  I  saw  gliding  across  a  path  in  Brazil. 
Snakes  coloured  in  this  peculiar  manner,  as  Mr.  Wallace  states 
on  the  authority  of  Dr.  Giinther,"^  are  found  nowhere  else 
in  the  world  except  in  S.  America,  and  here  no  less  than  four 
genera  occur.  One  of  these,  Elaps,  is  venomous ;  a  second  and 
widely-distinct  genus  is  doubtfully  venomous,  and  the  two 
others  are  quite  harmless.  The  species  belonging  to  these 
distinct  genera  inhabit  the  same  districts,  and  are  so  like  each 
other,  that  no  one  "but  a  naturalist  would  distinguish  the 
"  harmless  from  the  poisonous  kinds."  Hence,  as  Mr.  Wallace 
l>elieves,  the  innocuous  kinds  have  probably  acquired  their 
colours  as  a  protection,  on  the  principle  of  imitation ;  for  they 
would  naturally  be  thought  dangerous  by  their  enemies.  The 
cause,  however,  of  the  bright  colours  of  the  venomous  Elaps 
remains  to  be  explained,  and  this  may  j^erhaps  be  sexual 
selection. 

Snakes  produce  other  sounds  besides  hissing.  The  deadly 
Ediis  carinata  has  on  its  sides  some  oblique  rows  of  scales  of  a 
peculiar  structure  with  serrated  edges ;  and  when  this  snake  is 
excited,  these  scales  are  rubbed  against  each  other,  which  pro- 
duces "  a  curious  prolonged,  almost  hissing  sound."  ^^  With 
respect  to  the  rattling  of  the  rattle-snake,  we  have  at  last  some 
definite  information :  for  Professor  Aughey  states,^^  that  on  two 
occasions,  being  himself  unseen,  he  watched  from  a  little  distance, 
a  rattle-snake  coiled  up  with  head  erect,  which  continued  to 
rattle  at  short  intervals  for  half  an  hour :  and  at  last  he  saw 
another  snake  approach,  and  when  they  met  they  paired. 
Hence  he  is  satisfied  that  one  of  the  uses  of  the  rattle  is  to  bring 
the  sexes  together.  Unfortunately  he  did  not  ascertain  whether 
it  was  the  male  or  the  female  which  remained  stationary  and 
called  for  the  other.  But  it  by  no  means  follows  from  the 
above  foct  that  the  rattle  may  not  be  of  use  to  these  snakes  in 
other  ways,  as  a  warning  to  animals  which  would  otherwise 

«2  '  Westminster  Review,' July  1st,      Soc'  1871,  p.  196. 
1867,  p.  32.  "  i  'i'jjg     American     Naturalist,' 

'^^  Dr.    Anderson,    '  Proc.  Zoolog.      1873,  p.  85. 


354  The  Descent  of  Man.  Part  II. 

attack  them.  Nor  can  I  quite  disbelieve  the  several  accounts 
which  have  appeared  of  their  thus  paralysing  their  prey  with  fear. 
Some  other  snakes  also  make  a  distinct  noise  by  rapidly  vibrating 
their  tails  against  the  surrounding  stalks  of  plants ;  and  I  have  my- 
self heard  this  in  the  case  of  a  Trigonocephalus  in  S.  America. 

Zacertilia.— The  males  of  some,  probably  of  many  kinds  of 
lizards  fight  together  from  rivalry.  Thus  the  arboreal  Anolis 
cristatellus  of  S.  America  is  extremely  pugnacious  :  "  During  the 
"  spring  and  early  part  of  the  summer,  two  adult  males  rarely 
"  meet  without  a  contest.  On  first  seeing  one  another,  they  nod 
"  their  heads  up  and  down  three  or  four  times,  and  at  the  same 
"  time  expanding  the  frill  or  pouch  beneath  the  throat ;  their 
"  eyes  glisten  with  rage,  and  after  waving  their  tails  from 
"  side  to  side  for  a  few  seconds,  as  if  to  gather  energy,  they  dart 
"  at  each  other  furiously,  rolling  over  and  over,  and  holding 
"  firmly  with  their  teeth.  The  conflict  generally  ends  in  one  of 
"  the  combatants  losing  his  tail,  which  is  often  devoured  by  the 
"  victor."  The  male  of  this  species  is  considerably  larger  than 
the  female  f^  and  this,  as  far  as  Dr.  Giinther  has  been  able  to 
ascertain,  is  the  general  rule  with  lizards  of  all  kinds.  The 
males  alone  of  the  Cyrtodactylus  rahidus  of  the  Andaman  Islands 
possesses  pre-anal  pores ;  and  these  pores  judging  from  analogy 
probably  serve  to  emit  an  odour.*'^ 

The  sexes  often  differ  greatly  in  various  external  characters. 
The  male  of  the  above-mentioned  Anolis  is  furnished  with  a 
crest  which  runs  along  the  back  and  tail,  and  can  be  erected  at 
pleasure ;  but  of  this  crest  the  female  does  not  exhibit  a  trace. 
In  the  Indian  Cophotis  ceylanica,  the  female  has  a  dorsal  crest, 
though  much  less  developed  than  in  the  male ;  and  so  it  is,  as 
Dr.  Giinther  informs  me,  with  the  females  of  many  Iguanas, 
Chameleons,  and  other  lizards.  In  some  species,  however,  the 
crest  is  equally  developed  in  both  sexes,  as  in  the  Iguana  tubercu- 
lata.  In  the  genus  Sitana,  the  males  alone  are  furnished  with  a 
large  throat-pouch  (fig.  33),  which  can  be  folded  up  like  a  fan, 
and  is  coloured  blue,  black,  and  red  ;  but  these  splendid  colours 
are  exhibited  only  during  the  pairing-season.  The  female  does 
not  possess  even  a  rudiment  of  this  appendage.  In  the  Anolis 
cristatellus,  according  to  Mr.  Austen,  the  throat  pouch,  which  is 
bright  red  marbled  with  yellow,  is  present  in  the  female,  though 
in  a  rudimental  condition.  Again,  in  certain  other  lizards,  both 
sexes  are  equally  well  provided  with  throat  pouches.    Here  we 

"  Mr.  N.  L.  Austen    kept   these  ^^  Stoliczka,  '  Journal  of  Asiatic 

animals    alive    for    a     considerable  Soc.  of  Bengal,'  vol.  xxxiv.  1870,  p. 

time ;  see  '  Land  and  Water,'  July  166. 
1867,  p.  9. 


Chap.  XIT. 


Reptiles. 


355 


Fig.  S"?.  Sitana  minor.*  Male  with  tlie 
gular  pouch  expanded  (from  Giiuther's 
'  Reptiles  of  India  '). 


see  with  species  belonging  to  the  same  group,  as  in  so  many 
previous  cases,  the  same  character  either  confined  to  the  males, 
or  more  largely  developed  in  them  than  in  the  females,  or 
again  equally  developed  in  both  sexes.  The  little  lizards  of 
the  genus  Draco,  which  glide 
through  the  air  on  their  rib- 
supported  parachutes,  and 
which  in  the  beauty  of  their 
colours  baffle  description,  are 
furnished  with  skinny  appen- 
dages to  the  throat  "  like  the 
wattles  of  gallinaceous  birds." 
These  become  erected  when 
the  animal  is  excited.  They 
occur  in  both  sexes,  but  are 
best  developed  when  the  male 
arrives  at  maturity,  at  which 
age  the  middle  appendage  is 
sometimes  twice  as  long  as  the  head.  Most  of  the  species  like- 
wise have  a  low  crest  running  along  the  neck  ;  and  this  is  much 
more  developed  in  the  full-grown  males,  than  in  the  females  or 
young  males. *^^ 

A  Chinese  species  is  said  to  live 
in  pairs  during  the  spring  ;  "  and  if 
"  one  is  caught,  the  other  falls  from 
*'  the  tree  to  the  ground,  and  allows 
"  itself  to  be  captured  with  impu- 
**  nity," — I  presume  from  despair.®^ 

There  are  other  and  much  more 
remarkable  differences  between  the 
sexes  of  certain  lizards.  The  male 
of  Ceratopliora  aspera  bears  on  the 
extremity  of  his  snout  an  appendage 
half  as  long  as  the  head.  It  is 
cylindrical,  covered  with  scales, 
flexible,  and  apparently  capable  of 
erection:  in  the  female  it  is  quite 
rudimental.  In  a  second  species 
of  the  same  genus  a  terminal  scale 
forms  a  minute  horn  on  the  summit  of  the  flexible  appendage ; 
All   the   foregoing   statements      nificent   work   on  the  '  Reptiles   of 


Fig.  34.  Ceratophora  Stoddartii 
Upper  figure,  male ;  lower  figure, 
female. 


and  quotations,  in  regard  to  Cophotis, 
Sitana  and  Draco,  as  well  as  the 
following  facts  in  regard  to  Cerato- 
phora and  Chama)leon,  are  from  Dr. 
Giinther  himself,  or  from  his  mag- 


British   India, 
122,  130,  135. 

*^*  Mr.    Swinhoe 
Soc'  1870,  p.  240. 


Ray  Soc.   1864,    pp. 
Proc.    Zoo  log. 


,56 


The  Descent  of  Man. 


Part  li. 


and  in  a  tliird  species  (C.  Stoddartii,  fig.  34)  the  whole  appen- 
dage is  converted  into  a  horn,  which  is  usually  of  a  white 
colour,  but  assumes  a  purplish  tint  when  the  animal  is  excited. 
In  the  adult  male  of  this  latter  species  the  horn  is  half  an  inch 
in  length,  but  it  is  of  quite  minute  size  in  the  female  and  in  the 
yoang.  These  appendages,  as  Dr.  Giinther  has  remarked  to  me, 
may  be  compared  with  the  combs  of  gallinaceous  birds,  and 
apparently  serve  as  ornaments. 


Fig.  35.    Chama.'lcon  bifurcus.    Upper  figure,  male ;  lower  figure,  female. 

In  the  genus  Cliamaileon  we  come  to  the  acme  of  difference 
between  the  sexes.  The  upper  part  of  the  skull  of  the  male 
C.  hifarcus  (fig.  35),  an  inhabitant  of  Madagascar,  is  produced 
into  two  great,  solid,  bony  projections,  covered  with  scales  like 
the  rest  of  the  head;  and  of  this  wonderful  modification  of 
structure  the  female    exhibits    only   a  rudiment.      Again,  in 


Chap.  XII. 


Reptiles, 


357 


Charnodeon  Owenii  (fig.  3G),  from  the  West  Coast  of  Africa,  the 
male  bears  on  his  snout  and  forehead  three  curious  horns,  of 
which  the  female  has  not  a  trace.  These  horns  consist  of  an 
excrescence  of  bone  covered  with  a  smooth  sheath,  forming  part  of 
the  general  integu- 
ments of  the  body, 
so  that  they  are 
identical  in  struc- 
ture with  those  of  a 
bull,  goat,  or  other 
sheath  -  horned  ru- 
minant. Although 
the  three  horns 
differ  so  much  in 
appearance  from 
the  two  great  pro- 
longations of  the 
skull  in  C.  h  if  lire  us, 
we  can  hardly  doubt 
that  they  serve  the 
same  general  pur- 
pose in  the  economy 
of  these  two  ani- 
mals. The  first  con- 
jecture, which  will  occur  to  every  one,  is  that  they  are  u^ed 
by  the  males  for  fighting  together;  and  as  these  animals  are 
very  quarrelsome,*^*  this  is  probably  a  correct  view.  Mr.  T.  W. 
Wood  also  informs  me  that  he  once  watched  two  individuals  of 
0.  pumilus,  fighting  violently  on  the  branch  of  a  tree ;  they  flung 
their  heads  about  and  tried  to  bite  each  other ;  they  then  rested 
for  a  time,  and  afterwards  continued  their  battle. 

With  many  lizards,  the  sexes  dififer  slightly  in  colour,  the 
tints  and  stripes  of  the  males  being  brighter  and  more  distinctly 
defined,  than  in  the  females.  This,  for  instance,  is  the  case  with 
the  above  Cophotis  and  with  the  Acnntlwdactylus  caper, sis  of 
S.  Africa.  Iif  a  Cordylus  of  the  latter  country,  the  male  is 
either  much  redder  or  greener  than  the  female.  In  the  Indian 
Calotes  nigrilahris  there  is  a  still  greater  diiFerence ;  the  lips  also 
of  the  male  are  black,  whilst  those  of  the  female  are  green.  In 
our  common  little  viviparous  lizard  (Zoofoca  vivipara)  "the 
"  under  side  of  the  body  and  base  of  the  tail  in  the  male  are 
"  bright  orange,  spotted  with  black ;  in  the  female  these  parts 
"  are  pale-greyish-green  without  spots." '°    We  have  seen  that 

«"  Dr.    Bucholz,    '  Monatsberkht  '»  Bell,      'History      of     British 

K.  Preuss.  Akad.'  Jtm.  1874,  p.  78.      Reptik-s,'  2nd  edit.  1849,  p.  40. 


Fig  36.    Cl)ama?leon  Owenii.    Upper  figure,  male 
lower  figure,  female. 


358  The  Descent  of  Man.  Part  II. 

the  males  alone  of  Sitana  possess  a  throat-pouch ;  and  this  is 
splendidly  tinted  with  blue,  black,  and  red.  In  the  Proctotretus 
tenuis  of  Chile  the  male  alone  is  marked  with  spots  of  blue^ 
green,  and  coppery-red.'^^  In  many  cases  the  males  retain  the 
same  colours  throughout  the  year,  but  in  others  they  become 
much  brighter  during  the  breeding-season ;  I  may  give  as  an 
additional  instance  the  Calotes  maria,  which  at  this  season  has  a 
bright  red  head,  the  rest  of  the  body  being  green.''^ 

Both  sexes  of  many  species  are  beautifully  coloured  exactly 
alike ;  and  there  is  no  reason  to  suppose  that  such  colours  are 
protective.  No  doubt  with  the  bright  green  kinds  which  live 
in  the  midst  of  vegetation,  this  colour  serves  to  conceal  them ; 
and  in  N,  Patagonia  I  saw  a  lizard  (Proctotretus  multimaculatus) 
which,  when  frightened,  flattened  its  body,  closed  its  eyes,  and 
then  from  its  mottled  tints  was  hardly  distinguishable  from  the 
surrounding  sand.  But  the  bright  colours  with  which  so  many 
lizards  are  ornamented,  as  well  as  their  various  curious  appen- 
dages, were  probably  acquired  by  the  males  as  an  attraction, 
and  then  transmitted  either  to  their  male  offspring  alone,  or  to 
both  sexes.  Sexual  selection,  indeed,  seems  to  have  played 
almost  as  important  a  part  with  reptiles  as  with  birds ;  and  the 
less  conspicuous  colours  of  the  females  in  comparison  with  the 
males  cannot  be  accounted  for,  as  Mr.  Wallace  believes  to  be  the 
case  with  birds,  by  the  greater  exposure  of  the  females  to  danger 
during  incubation. 


CHAPTER  XIII. 
Secondary  Sexual  Chaeacters  of  Birds. 

Sexual  differences — Law  of  battle  —  Special  weapons — Vocal  organs  — 
Instrunneutal  music — Love-antics  and  dances — Decorations,  permanent 
and  seasonal — Double  and  single  annual  moults — Display  of  ornaments 
by  the  males. 

Secondary  sexual  characters  are  more  diversified  and  con- 
spicuous in  birds,  though  not  perhaps  entailing  riiore  important 
changes  of  structure,  than  in  any  other  class  of  animals.  I  shall, 
therefore,  treat  the  subject  at  considerable  length.  Male  birds 
sometimes,  though  rarely,  possess  special  weapons  for  fighting 

^^  For  Proctotretus  see  *  Zoology  the  Indian  Calotes,  see  '  Reptiles  of 

of  the    Voyage    of  the    "  Beagle :"  British  India,'  by   Dr.  Giinther,  p. 

Keptiles,'  by  Mr.    Bell,  p.   8.     For  143. 

the  Lizards  of  S.  Africa,  see  '  Zoology  ^^  Giinther  in  *  Proc.  Zoolog.  Soc' 

of    S.     Africa:    Reptiles,'    by     Sir  1870,    p.     778,    with     a     coloured 

Andrew  Smith,  pi.  25  and  39.     For  figure. 


Ch.u'.  Xlir.  Birds.  359 


with  each  other.  They  charm  the  female  by  vocal  or  instru- 
mental music  of  the  most  varied  kinds.  They  are  ornamented 
by  all  sorts  of  combs,  wattles,  protuberances,  horns,  air-distended 
sacks,  top-knots,  naked  shafts,  plumes  and  lengthened  feathers 
gi'acefully  springing  from  all  parts  of  the  body.  The  beak  and 
naked  skin  about  the  head,  and  the  feathers  are  often  gorgeously 
coloured.  The  males  sometimes  pay  their  court  by  dancing,  or 
by  fantastic  antics  performed  either  on  the  ground  or  in  the  air. 
In  one  instance,  at  least,  the  male  emits  a  musky  odour,  which 
we  may  suppose  serves  to  charm  or  excite  the  female ;  for  that 
excellent  observer,  Mr.  Eamsay,^  says  of  the  Australian  musk- 
duck  (^Biziura  Jobata)  that  "  the  smell  which  the  male  emits 
"  during  the  summer  months  is  confined  to  that  sex,  and  in 
"  some  individuals  is  retained  throughout  the  year ;  I  have 
"  never,  even  in  the  breeding-season,  shot  a  female  which  had 
"  any  femell  of  musk."  So  powerful  is  this  odour  during  the 
pairing-season,  that  it  can  be  detected  long  before  the  bird  can 
be  seen.2  On  the  whole,  birds  appear  to  be  the  most  aesthetic  of 
all  animals,  excepting  of  course  man,  and  they  have  nearly  the 
same  taste  for  the  beautiful  as  we  have.  This  is  shewn  by  our 
enjoyment  of  the  singing  of  birds,  and  by  our  women,  both 
civilised  and  savage,  decking  their  heads  with  borrowed  plumes, 
and  using  gems  which  are  hardly  more  brilliantly  coloured  than 
the  naked  skin  and  wattles  of  certain  birds.  In  man,  however, 
when  cultivated,  the  sense  of  beauty  is  manifestly  a  far  more 
complex  feeling,  and  is  associated  with  various  intellectual 
ideas. 

Before  treating  of  the  sexual  characters  with  which  we  are 
here  more  particularly  concerned,  I  may  just  allude  to  certain 
differences  between  the  sexes  which  apparently  depend  on 
differences  in  their  habits  of  life;  for  such  cases,  though 
common  in  the  lower,  are  rare  in  the  higher  classes.  Two 
humming-birds  belonging  to  the  genus  Eustephanus,  which 
inhabit  the  island  of  Juan  Fernandez,  were  long  thought  to  be 
specifically  distinct,  but  are  now  known,  as  Mr.  Gould  informs 
me,  to  be  the  male  and  female  of  the  same  species,  and  they 
diifer  slightly  in  the  form  of  the  beak.  In  another  genus  of 
humming-birds  (Grypus),  the  beak  of  the  male  is  serrated  along 
the  margin  and  hooked  at  the  extremity,  thus  differing  much 
from  that  of  the  female.  In  the  Neomorpha  of  New  Zealand, 
there  is,  as  we  have  seen,  a  still  wider  difference  in  the  form  of 
the  beak  in  relation  to  the  manner  of  feeding  of  the  two  sexes. 
Something  of  the  same  kind  has  been  observed  with  the  gold- 

•  '  Ibis,'  vol.  iii.  (new  series)  1867,  ^  Gould,  '  Handbook  to  the  Birds 

p.  414.  of  Australia,'  1805,  vol.  ii.  p.  383. 


360  The  Descent  of  Ma?t.  Yawt  11 

finch  {Cardudis  eUgans),  for  I  am  assured  by  Mr.  J.  Jenrier  Weir 
tliat  the  birdcatchers  can  distingnish  the  males  by  their  slightly 
longer  beaks.  The  flocks  of  males  are  often  found  feeding  on 
the  seeds  of  the  teazle  (Dipsacus),  which  they  can  reach  with 
their  elongated  beaks,  whilst  the  females  more  commonly  feed 
on  the  seeds  of  the  betony  or  Scrophularia.  With  a  sKght 
difference  of  this  kind  as  a  foundation,  we  can  see  how  the  beaks 
of  the  two  sexes  might  be  made  to  differ  greatly  through  natural 
selection.  In  some  of  the  above  cases,  however,  it  is  possible 
that  the  beaks  of  the  males  may  have  been  first  modified  in 
relation  to  their  contests  with  other  males;  and  that  this 
afterwards  led  to  slightly  changed  habits  of  life. 

Laiv  of  Battle. — Almost  all  male  birds  are  extremely  pug- 
nacious, using  their  beaks,  wings,  and  legs  for  fighting  together. 
We  see  this  every  spring  with  our  robins  and  sparrows.  The 
smallest  of  all  birds,  namely  the  humming-bird,  is  one  ♦of  the 
most  quarrelsome.  Mr.  Gosse^  describes  a  battle  in  which  a 
pair  seized  hold  of  each  other's  beaks,  and  whirled  round  and 
round,  till  they  almost  fell  to  the  gi-ound ;  and  M.  Montes  de 
Oca,  in  speaking  of  another  genus  of  humming-bird,  says  that 
two  males  rarely  meet  without  a  fierce  aerial  encounter :  when 
kept  in  cages  "  their  fighting  has  mostly  ended  in  the  splitting  of 
"  the  tongue  of  one  of  the  two,  which  then  surely  dies  from 
"  being  unable  to  feed."^  With  W^aders,  the  males  of  the 
common  water-hen  {GalUnula  cMoropus)  "when  jDairing,  fight 
"  violently  for  the  females :  they  stand  nearly  upright  in  the 
"  water  and  strike  with  their  feet."  Two  were  seen  to  be  thus 
engaged  for  half  an  hour,  until  one  got  hold  of  the  head  of  the 
other,  which  would  have  been  killed,  had  not  the  observer 
interfered;  the  female  all  the  time  looking  on  as  a  quiet  spec- 
tator.^ Mr.  Blyth  informs  me  that  the  males  of  an  allied  bird 
(Gallicrex  cristatus)  are  a  third  larger  than  the  females,  and  are 
so  pugnacious  during  the  breeding- season,  that  they  are  kept  by 
the  natives  of  Eastern  Bengal  for  the  sake  of  fighting.  Various 
other  birds  are  kept  in  India  for  the  same  purpose,  for  instance, 
the  bulbuls  (Pycnonotus  haemorrhous)  which  "  fight  with  great 
"  spirit."*^ 

The  polygamous  ruff  (Machttes  puynax,  fig.  37)  is  notorious 
for  his  extreme  pugnacity  ;  and  in  the  spring,  the  males,  which 
are  considerably  larger  than  the  females,  congregate  day  after 


3  Quoted  by  Mr.  Gould,  'Intro- 

Ireland :    Birds,'   a'oI.    ii.    1850,  p. 

duction   to   the    Trochilida?,'    1861, 

327. 

p.  29. 

«  Jerdon,  'Birds  of  India,'  1863, 

*  Gould,  ibid.  p.  52. 

vol.  ii.  p.  96. 

5  W.  Thompson,   'Nat.    Hist,    of 

Chap.  XIII. 


Law  of  Battle. 


361 


.day  at  a  particular  spot,  wlierc  tlie  females  propose  to  lay  their 
eggs.  The  fowlers  discover  these  spots  by  the  turf  being 
trampled  somewhat  bare.  Here  they  fight  very  much  like  game- 
cocks, seizing  each  other  with  their  beaks  and  striking  with 
their  wings.  The  great  ruff  of  feathers  round  the  neck  is  then 
erected,  and  according  to  Col.  Montagu  "  sweeps  the  ground  as 
"  a  shield  to  defend  the  more  tender  parts;"  and  this  is  the  only 


362  The  Descent  of  Man,  Pakt  II. 

instance  known  to  me  in  the  case  of  birds,  of  any  structure 
serving  as  a  shield.  The  ruff  of  feathers,  however,  from  its 
varied  and  rich  colours  probably  serves  in  chief  part  as  an  orna- 
ment. Like  most  pugnacious  birds,  they  seem  always  ready  to 
fight,  and  when  closely  confined  often  kill  each  other ;  but 
Montagu  observed  that  their  pugnacity  becomes  greater  during 
the  spring,  when  the  long  feathers  on  their  necks  are  fully 
developed ;  and  at  this  period  the  least  movement  by  any  one 
bird  provokes  a  genei-al  battle.^  Of  the  pugnacity  of  web-footed 
birds,  two  instances  will  suffice :  in  Guiana  "  bloody  fights  occur 
"  during  the  breeding-season  between  the  males  of  the  wild 
"  musk-duck  (CaiVi'na  moschata') ;  and  where  these  fights  have 
**^  occurred  the  river  is  covered  for  some  distance  with  feathers."  * 
Birds  which  seem  ill-adapted  for  fighting  engage  in  fierce  con- 
flicts ;  thus  the  stronger  males  of  the  pelican  drive  away  the 
weaker  ones,  snapping  with  their  huge  beaks  and  giving  heavy 
blows  with  their  wings.  Male  snipe  fight  together,  "  tugging 
"  and  i^ushing  each  other  with  their  bills  in  the  most  curious 
"  manner  imaginable."  Some  few  birds  are  believed  never  to 
fight ;  this  is  the  case,  according  to  Audubon,  with  one  of  the 
woodpeckers  of  the  United  States  (Picks  awra^ws), although  "the 
"  hens  are  followed  by  even  half  a  dozen  of  their  gay  suitors."  ^ 

The  males  of  many  birds  are  larger  than  the  females,  and  this 
no  doubt  is  the  result  of  the  advantage  gained  by  the  larger  and 
stronger  males  over  their  rivals  during  many  generations.  The 
difference  in  size  between  the  two  sexes  is  carried  to  an  extreme 
point  in  several  Australian  species ;  thus  the  male  musk-duck 
(Biziura)  and  the  male  Ciudoramphus  cruralis  (allied  to  our 
pipits)  are  by  measurement  actually  twice  as  large  as  their 
respective  females.^''  With  many  other  birds  the  females  are 
larger  than  the  males ;  and  as  formerly  remarked,  the  explana- 
tion often  given,  namely,  that  the  females  have  most  of  the  work 
in  feeding  their  young,  will  not  suffice.  In  some  few  cases,  as 
we  shall  hereafter  see,  the  females  apparently  have  acquired 
their  greater  size  and  strength  for  the  sake  of  conquering  other 
females  and  obtaining  possession  of  the  males. 

The  males  of  many  gallinaceous  birds,  especially  of  the  poly- 
gamous kinds,  are  furnished  with  special  weapons  for  fighting 
with  their  rivals,  namely  spurs,  which  can  be  used  with  fearful 

^    Macgillivrar,       *  Hist.       Brit.  i.  p.  191.     For  pelicans  and  snipes, 

Birds,'  vol.  iv.  1852,  pp.  177-181.  see  vol.  iii.  pp.  138,  477. 

•  Sir  R.  Schomburgk,  in  '  Journal  '<>  Gould,  *  Handbook  of  Birds  of 
of  R.  Geograph.  Soc' vol.  xiii,  1843,  Australia,'  vol.  i.  p.  395;  vol,  ii.  p. 
p.  31.  383 

•  'Ornithological  Biography,' vol. 


Chap.  XII r.  Law  of  Battle.  363 

effect.  It  has  been  recorded  by  a  trustworthy  writer "  that  in 
Derbyshire  a  kite  struck  at  a  game-hen  accompanied  by  her 
chickens,  when  the  cock  rushed  to  the  rescue,  and  drove  his 
spur  right  through  the  eye  and  skull  of  the  aggressor.  The 
spur  was  with  difficulty  drawn  from  the  skull,  and  as  the  kite 
though  dead  retained  his  grasp,  the  two  birds  were  firmly 
locked  together;  but  the  cock  when  disentangled  was  very 
little  injured.  The  invincible  courage  of  the  game-cock  is 
notorious :  a  gentleman  who  long  ago  witnessed  the  brutal 
scene,  told  me  that  a  bird  had  both  its  legs  broken  by  some 
accident  in  the  cockpit,  and  the  owner  laid  a  wager  that  if  the 
legs  could  be  spliced  so  that  the  bird  could  stand  upright,  he 
would  continue  fighting.  This  was  effected  on  the  spot,  and  the 
bird  fought  with  undaunted  courage  until  he  received  his  death- 
stroke.  In  Ceylon  a  clo'sely  allied,  wild  species,  the  Gallus 
Stanley i,  is  known  to  fight  desperately  "in  defence  of  his 
"  seraglio,"  so  that  one  of  the  combatants  is  frequently  found 
dead.^^  An  Indian  partridge  ( Ortygornis  gularis),  the  male  of 
which  is  furnished  with  strong  and  sharp  spurs,  is  so  quarrel- 
some, "  that  the  scars  of  former  fights  disfigure  the  breast  of 
"  almost  every  bird  you  kill."  ^^ 

The  males  of  almost  all  gallinaceous  birds,  even  those  which 
are  not  furnished  with  spurs,  engage  during  the  breeding-season 
in  fierce  conflicts.  The  Capercailzie  and  Black-cock  (Tetrao 
urogallus  and  T.  tetrix),  which  are  both  polygamists,  have  regular 
appointed  places,  where  during  many  weeks  they  congregate  in 
numbers  to  fight  together  and  to  display  their  charms  before  the 
females.  Dr.  W.  Kovalevsky  informs  me  that  in  Eussia  he  has 
seen  the  snow  all  bloody  on  the  arenas  where  the  capercailzie 
have  fought ;  and  the  black-cocks  "  make  the  feathers  fly  in  every 
"  direction,"  when  several  "  engage  in  a  battle  royal."  The 
elder  Brehm  gives  a  curious  account  of  the  Balz,  as  the  love- 
dances  and  love-songs  of  the  Black-cock  are  called  in  Germaoy. 
The  bird  utters  almost  continuously  the  strangest  noises :  *'  he 
"  holds  his  tail  up  and  spreads  it  out  like  a  fan,  he  lifts  up.  his 
"  head  and  neck  with  all  the  feathers  erect,  and  stretches  his 
"  wings  from  the  body.  Then  he  takes  a  few  jumps  in  different 
"  directions,  sometimes  in  a  circle,  and  presses  the  under  part  of 
"  his  beak  so  hard  against  the  ground  that  the  chin  feathers  are 
"  rubbed  off.  During  these  movements  he  beats  his  wings  and 
"  turns  round  and  round.  The  more  ardent  he  grows  the  more 
"  lively  he  becomes,  until  at  last  the  bird  appears  like  a  frantic 

'>  Mr.   Hewitt    in    the   'Poultry      Nat.  Hist.' vol.  xiv.  1854,  p.  63. 
Book  by  Tegetmeier,'  1866,  p.  137.  i^  Jerdon,  'Birds  of  India,'  vol. 

-2  Layard,  'Annals  and  Mag.  of      iii.  p.  574. 


364  The  Descent  of  Man.  Part  II. 

"  creature."  At  snch  times  the  black-cocks  are  so  absorbed  that 
they  become  almost  blind  and  dea ',  but  less  so  than  the  caper- 
cailzie :  hence  bird  after  bird  may  be  shot  on  the  same  spot,  or 
even  caught  by  the  hand.  After  performing  these  antics  the 
males  begin  to  fight :  and  the  same  black-cock,  in  order  to  prove 
his  strength  over  several  antagonists,  will  visit  in  the  course  of 
one  morning  several  Balz-jDlaces,  which  remain  the  same  during 
successive  years.^^ 

The  peacock  with  his  long  train  appears  more  Mke  a  dandy 
than  a  warrior,  but  he  sometimes  engages  in  fierce  contests  :  the 
Kev.  W.  Darwin  Fox  informs  me  that  at  some  little  distance 
from  Chester  two  peacocks  became  so  excited  whilst  fighting, 
that  they  flew  over  the  wiiole  city,  still  engaged,  until  they 
alighted  on  the  top  of  St.  John's  tower. 

The  spur,  in  those  gallinaceous  bird's  which  are  thus  provided, 
is  generally  single ;  but  Polyplectron  (see  fig.  51,  p.  397)  has  two 
or  more  on  each  leg;  and  one  of  the  Blood- pheasants  {Ithaginis 
cruentus)  has  been  seen  with  five  spurs.  The  spurs  are  generally 
confined  to  the  male,  being  represented  by  mere  knobs  or  rudi- 
ments in  the  female ;  but  the  females  of  the  Java  peacock  {Pavo 
muticus)  and,  as  I  am  informed  by  IVIr.  Blyth,  of  the  small  fire- 
backed  pheasant  (EujjJocamus  erythroptliahnus)  jDossess  s^Durs. 
In  Galloperdix  it  is  usual  for  the  males  to  have  two  spurs,  and 
for  the  females  to  have  only  one  on  each  leg.^^  Hence  spurs  may 
be  considered  as  a  masculine  structure,  which  has  been  occasion- 
ally more  or  less  transferred  to  the  females.  Like  most  other 
secondary  sexual  characters,  the  spurs  are  highly  variable,  both 
in  number  and  development,  in  the  same  species. 

Various  birds  have  spurs  on  their  wings.  But  the  Egyptian 
goose  {ChenaJoj)ex  cegyptiacus)  has  only  "  bare  obtuse  knobs,"  and 
these  probably  shew  us  the  first  steps  by  which  true  spurs  have 
been  developed  in  other  species.  In  the  spur-winged  goose, 
Pledropterus  gamhensis,  the  males  have  much  larger  spurs  than 
the  females;  and  they  use  them,  as  I  am  informed  by  Mr. 
Bartlett,  in  fighting  together,  so  that,  in  this  case,  the  wing-spui-s 
serve  as  sexual  weapons ;  but  according  to  Livingstone,  they  are 
chiefly  used  in  the  defence  of  the  young.  The  Palamedea 
(fig.  88)  is  armed  with  a  pair  of  spurs  on  each  wing ;  and  these 
are  such  formidable  weapons,  that  a  single  blow  has  been  known 
to  drive  a  dog  howling  away.  But  it  does  not  appear  that  the 
spurs  in  this  case,  or  in  that  of  some  of  the  spur-winged  rails, 

1*    Brehm,    '  Illust.    Thierleben,'  Sweden,'  &c.,  1867,  p.  79. 
1867,  B.  iv.  s.  351.     Some  of  the  ^*    Jerdon,  'Birds  of  India:    ou 

foregoing  statements  are  taken  from  Ithaginis,  vol.  iii.  p.  523  ;  on  Gallo- 

L.    Lloyd,    'The    Game    Birds    of  perdix,  p.  541. 


Chap.  XIII. 


Law  of  Battle. 


3^5 


are  larger  in  the  male  than  in  the  female.^'^    In  certain  plovers, 
however,  the  wing-spurs  must  be  considered  as  a  sexual  cha- 


Fig.  38.    Palamedea  cornuta  (from  Brehm),  shewing  the  double  wing-spurs,  and  the 
filament  on  the  head. 


i«  For   the   Egyptian    goose,    see      p.  639.    For  Plectropterus,  <  Living- 
Macgillivray, '  British  Birds,'  vol.  iv.      stone's  Travels,'  p.  254.      For  Pala- 


^66  The  Descent  of  Man.  Part  II. 

racter.  Thus  in  the  male  of  our  common  peewit  (  Vanellus  cris- 
tatus)  the  tubercle  on  the  shoulder  of  the  wing  becomes  more 
prominent  during  the  breeding-season,  and  the  males  fight 
together.  In  some  species  of  Lobivanellus  a  similar  tubercle 
becomes  developed  during  the  breeding-season  "  into  a  short 
"■  horny  spur."  In  the  Australian  L.  lobatus  both  sexes  have  spurs, 
but  these  are  much  larger  in  the  males  than  in  the  females.  In 
an  allied  bird,  the  Hoplopterusarmatus,  the  spurs  do  not  increase 
in  size  during  the  breeding-season ;  but  these  birds  have  been 
seen  in  Egypt  to  fight  together,  in  the  same  manner  as  our 
peewits,  by  turning  suddenly  in  the  air  and  striking  sideways 
at  each  other,  sometimes  with  fatal  results.  Thus  also  they 
drive  away  other  enemies.^^ 

The  season  of  love  is  that  of  battle ;  but  the  males  of  some 
birds,  as  of  the  game-fowl  and  ruff,  and  even  the  young  males  of 
the  wild  turkey  and  grouse,^*  are  ready  to  fight  whenever  they 
meet.  The  presence  of  the  female  is  the  teterrima  belli  causa. 
The  Bengali  baboos  make  the  pretty  little  males  of  the  amadavat 
( tLstrtlda  amandava)  fight  together  by  placing  three  small  cages 
in  a  row,  with  a  female  in  the  middle ;  after  a  little  time  the  two 
males  are  turned  loose,  and  immediately  a  desperate  battle  en- 
sues.^^  When  many  males  congregate  at  the  same  appointed 
spot  and  fight  together,  as  in  the  case  of  grouse  and  various  other 
birds,  they  are  generally  attended  by  the  females,^"  which  after- 
wards pair  with  the  victorious  combatants.  But  in  some  cases  the 
pairing  precedes  instead  of  succeeding  the  combat :  thus  accord- 
ing to  Audubon,^^  several  males  of  the  Virginian  goat-sucker 
{Caprimulgus  virginiamis)  "court,  in  a  highly  entertaining 
"  manner  the  female,  and  no  sooner  has  she  made  her  choice, 
"  than  her  approved  gives  chase  to  all  intruders,  and  drives 


medea,  Brehm's  '  Thievleben,'  B.  iv.  2<*    Richardson  on  Tetrao  umhellus 

s.  740.     See  also  on  this  bird  Azara,  'Fauna   Bor,  Amer. :  Birds,'   1831, 

'Voyages    dans   TAme'rique  me'rid.'  p.  343.     L.  Lloyd, 'Game  Birds  of 

torn.  iv.  1809,  pp.  179,  253.  Sweden,'    1867,  pp.   22,  79,  on  the 

^'^  See,  on  our  peewit,  Mr.  R.  Carr  capercailzie  and  black-cock.  Brehm, 

in    '  Land    and    Water,'    Aug.    8th,  however,  asserts  ('  Thierleben,'  &c., 

1868,  p.  46.      In  regard    to  Lobi-  B.  iv.  s.  352)  that  in  Germany  the 

vanellus,    see    Jerdon's    '  Birds    of  grey-hens   do  not    generally  attend 

India,'  vol.  iii.  p.  647,  and  Gould's  the  Balzen  of  the  black-cocks,  but 

*  Handbook    of  Birds  of  Australia,'  this  is  an  exception  to  the  common 

vol.  ii.  p.  220.    For  the  Holopterus,  rule;    possibly    the     hens   may    lie 

see  Mr.  Allen  in  the  '  Ibis,'  vol.  v.  hidden  in  the    surrounding  bushes, 

1863,  p.  156.  as  is  known  to  be  the  case  with  the 

**  Audubon, '  Ornith.  Biography,'  grey-hens  in  Scandinavia,  and  with 

vol.  ii.  p.  492 ;  vol.  i.  pp.  4-13.  other  species  in  N.  America. 

1*  Mr.  Blyth,  '  Land  and  Water,  "^^  '  Ornithological       Biography, 

1867,  p.  212.  vol.  ii.  p.  275. 


Chap.  XIII.  Law  of  Battle.  367 

*'  them  beyond  his  dominions."  Generally  the  males  try  to  drive 
away  oi*  kill  their  rivals  before  they  pair.  It  does  not,  however, 
appear  that  the  females  invariably  prefer  the  victorious  males. 
I  have  indeed  been  assured  by  Dr.  W.  Kovalevsky  that  the 
female  capercailzie  sometimes  steals  away  with  a  young  male 
who  has  not  dared  to  enter  the  arena  with  the  older  cocks,  in 
the  same  manner  as  occasionally  happens  with  the  does  of  the 
red-deer  in  Scotland.  When  two  males  contend  in  presence  of 
a  single  female,  the  victor,  no  doubt,  commonly  gains  his  desire ; 
but  some  of  these  battles  are  caused  by  wandering  males  trying 
to  distract  the  peace  of  an  already  mated  pair.^ 

Even  with  the  most  pugnacious  species  it  is  probable  that  the 
pairing  does  not  depend  exclusively  on  the  mere  strength  and 
courage  of  the  male  ;  for  such  males  are  generally  decorated  with 
various  ornaments,  which  often  become  more  brilliant  during  the 
breeding-season,  and  which  are  sedulously  displayed  before  the 
females.  The  males  also  endeavour  to  charm  or  excite  their 
mates  by  love-notes^  songs,  and  antics ;  and  the  courtship  is,  in 
many  instances,  a  prolonged  affair.  Hence  it  is  not  probable 
that  the  females  are  indifferent  to  the  charms  of  the  opposite 
sex,  or  that  they  are  invariably  compelled  to  yield  to  the  victorious 
males.  It  is  more  probable  that  the  females  are  excited,  either 
before  or  after  the  conflict,  by  certain  males,  and  thus  un- 
consciously prefer  them.  In  the  case  of  Tetrao  umhellus,  a  good 
observer  ^^  goes  so  far  as  to  believe  that  the  battles  of  the  males 
"  are  all  a  sham,  performed  to  show  themselves  to  the  greatest 
*'  advantage  before  the  admiring  females  who  assemble  around ;  for 
*'  I  have  never  been  able  to  find  a  maimed  hero,  and  seldom  more 
"  than  a  broken  feather."  I  shall  have  to  recur  to  this  subject, 
but  I  may  here  add  that  with  the  Tetrao  cupido  of  the  United 
States,  about  a  score  of  males  assemble  at  a  particular  spot,  and 
strutting  about,  make  the  whole  air  resound  with  their  extra- 
ordinary noises.  At  the  first  answer  from  a  female  the  males 
begin  to  fight  furiously,  and  the  weaker  give  way;  but  then, 
according  to  Audubon,  both  the  victors  and  vanquished  search 
for  the  female,  so  that  the  females  must  either  then  exert  a 
choice,  or  the  battle  must  be  renewed.  So,  again,  with  one  of 
the  field-starlings  of  the  United  States  (Stumella  ludoviciana) 
the  males  engage  in  fierce  conflicts,  "  but  at  the  sight  of  a  female 
"  they  all  fly  after  her,  as  if  mad."  ^^ 

"  Brehm,    'Thierleben,'   &e.,    B.  24  Audubon's    '  Ornitholog.     Bio- 

iv.  1867,  p.  990.  Audubon, '  Ornith.  graphy  ;'  on   Tetrao  cupido,  vol.  ii. 

Biography,'  vol.  ii.  p.  492.  p.  492 ;  on  the  Sturnus,  vol.  ii.  p. 

23  'Land  and  Water,'  July  25th,  219. 
1868,  p.  14. 


368  The  Descent  of  Man.  Part  II. 

Vocal  and  instrumental  music. — With  birds  the  voice  serves  to 
express  various  emotions,  such  as  distress,  fear,  anger,  triumph, 
or  mere  happiness.  It  is  apparently  sometimes  used  to  excite 
terror,  as  in  the  case  of  the  hissing  noise  made  by  some  nestling- 
birds.  Audubon  ^°  relates  that  a  mght-heron  (Ardea  nycticorax, 
Linn.)  which  he  kept  tame,  used  to  hide  itself  when  a  cat 
approached,  and  then  "  suddenly  start  up  uttering  one  of  the 
"  most  frightful  cries,  apparently  enjoying  the  cat's  alarm  and 
"  flight."  The  common  domestic  cock  clucks  to  the  hen,  and 
the  hen  to  her  chickens,  when  a  dainty  morsel  is  found.  The 
hen,  when  she  has  laid  an  egg,  "  repeats  the  same  note  very  often, 
"  and  concludes  with  the  sixth  above,  which  she  holds  for  a 
longer  time ;  "  ^^  and  thus  she  expresses  her  joy.  Some  social 
birds  apparently  call  to  each  other  for  aid ;  and  as  they  flit  from 
tree  to  tree,  the  flock  is  kept  together  by  chirp  answering  chirp. 
During  the  nocturnal  migrations  of  geese  and  other  water-fowl, 
sonorous  clangs  from  the  van  may  be  heard  in  the  darkness 
overhead,  answered  by  clangs  in  the  rear.  Certain  cries  serve 
as  danger  signals,  which,  as  the  sportsman  knows  to  his  cost, 
are  understood  by  the  same  S];)ecies  and  by  others.  The 
domestic  cock  crows,  and  the  humming-bird  chii'ps,  in  triumph 
over  a  defeated  rival.  The  true  song,  however,  of  most  birds 
and  various  strange  cries  are  chiefly  uttered  during  the  breed- 
ing-season, and  serve  as  a  charm,  or  merely  as  a  call-note,  to  the 
other  sex. 

Naturalists  are  much  divided  with  respect  to  the  object  of  the 
singing  of  birds.  Tew  more  careful  observers  ever  lived  than 
Montagu,  and  he  maintained  that  the  "  males  of  song-birds  and 
"  of  many  others  do  not  in  general  search  for  the  female,  but, 
"  on  the  contrary,  their  business  in  the  spring  is  to  perch  on  some 
'•  conspicuous  spot,  breathing  out  their  full  and  amorous  notes, 
"  which,  by  instinct,  the  female  knows,  and  repairs  to  the  spot  to 
**  choose  her  mate."  ^^  Mr.  Jenner  Weir  informs  me  that  this 
is  certainly  the  case  with  the  nightingale.  Bechstein,  who  kept 
birds  during  his  whole  life,  asserts,  "that  the  female  canary 
"  always  chooses  the  best  singer,  and  that  in  a  state  of  nature 
"  the  female  finch  selects  that  male  out  of  a  hundred  whose 
"  notes  please  her  most."  ^^  There  can  be  no  doubt  that  birds 
closely  attend  to  each  other's  song.    Mr.  Weir  has  told  me  ot 

25  '  Ornithological  Biograph.'  vol.  vogel,'    1840,   s.  4.     Mr.    Harrisou 

V.  p.  601.  Weir    likewise   writes  to   me: — "I 

2®  The  Hon.  Daines    Barrington,  "  am  informed  that  the  best  singing 

'  Philosoph,  Transact.'  1773,  p.  252.  "  males  generally  get  a  mate  first, 

2^  '  Ornithological       Dictionary,'  "  when   they  are   bred  in  the  same 

1833,  p.  475.  "room." 

2*  '  Naturgeschichte  der   Stuben- 


Chap.  XIII.  Vocal  Music.  369 

the  case  of  a  bullfinch  which  had  been  iaught  to  pipe  a  German 
waltz,  and  who  was  so  good  a  performer  that  he  cost  ten 
guineas ;  when  this  bird  was  first  introduced  into  a  room  where 
other  birds  were  kept  and  he  began  to  sing,  all  the  others,  con- 
sisting of  about  twenty  linnets  and  canaries,  ranged  themselves 
on  the  nearest  side  of  their  cages,  and  listened  with  the  greatest 
interest  to  the  new  performer.  Many  naturalists  believe  that 
the  singing  of  birds  is  almost  exclusively  "the  effect  of  rivalry 
"  and  emulation,"  and  not  for  the  sake  of  charming  their  mates. 
This  was  the  opinion  of  Daines  Barrington  and  White  of 
Selborne,  who  both  especially  attended  to  this  subjcct.^^  Bar- 
rington, however,  admits  that  "  superiority  in  song  gives  to 
"  birds  an  amazing  ascendancy  over  others,  as  is  well  known  to 
"  bird-catchers." 

It  is  certain  that  there  is  an  intense  degree  of  rivalry  between 
the  males  in  their  singing.  Bird-fanciers  match  their  birds  to 
see  which  will  sing  longest ;  and  I  was  told  by  Mr.  Yarrel  that 
a  first-rate  bird  will  sometimes  sing  till  he  drops  down  almost 
dead,  or  according  to  Bechstein,^"  quite  dead  from  rupturing  a 
vessel  in  the  lungs.  Whatever  the  cause  maybe,  male  birds,  as 
I  hear  from  Mr.  Weir,  often  die  suddenly  during  the  season  of 
song.  That  the  habit  of  singing  is  sometimes  quite  ind-'pendent 
of  love  is  clear,  for  a  sterile,  hybrid  canary-bird  has  been  de- 
scribed ^^  as  singing  whilst  viewing  itself  in  a  mirror,  and  then 
dashing  at  its  own  image;  it  likewise  attacked  with  fury  a 
female  canary,  when  put  into  the  same  cage.  The  jealousy 
excited  by  the  act  of  singing  is  constantly  taken  advantage  of  by 
bird-catchers;  a  male,  in  good  song,  is  hidden  and  protected, 
whilst  a  stuffed  bird,  surrounded  by  limed  twigs,  is  exposed  to 
view.  In  this  manner,  as  Mr.  Weir  informs  me,  a  man  has  in  the 
course  of  a  single  day  caught  fifty,  and  in  one  instance  seventy, 
male  chaffinches.  The  power  and  inclination  to  sing  differ  so 
gTeatly  with  birds  that  although  the  price  of  an  ordinary  male 
chaffinch  is  only  sixpence,  Mr.  Weir  saw  one  bird  for  which  the 
bird-catcher  asked  three  pounds;  the  test  of  a  really  good 
singer  being  that  it  will  continue  to  sing  whilst  the  cage  is 
swung  round  the  owner's  head. 

That  male  birds  should  sing  from  emulation  as  well  as  for 
charming  the  female,  is  not  at  all  incompatible;  and  it  might 
have  been  expected  that  these  two  habits  would  have  concurred, 
like  those  of  display  and  pugnacity.     Some  authors,  however, 

"  'Philosophical      Transactions,'  ^^  '  Naturgesch.  der  Stubenvogel,' 

1773,     p.    263.     White's    'Natural  1840,  s.  252. 

History  of  Selborne,'  1825,  vol.  i.  p.  »'   Mr.  Bold,  '  Zoologist,' 1843-44, 

246.  p.  659. 

17 


370  The  Descent  of  Man.  Part  II. 

argue  that  the  song  of  the -male  cannot  serve  to  charm  the  female, 
because  the  females  of  some  few  species,  such  as  of  the  canary, 
robin,  lark,  and  bnllfinch,  especially  when  in  a  state  of  widow- 
hood, as  Bechstein  remarks,  pour  forth  fairly  melodious  strains. 
In  some  of  these  cases  the  habit  of  singing  may  be  in  part 
attributed  to  the  females  having  been  highly  fed  and  confined,^'* 
for  this  disturbs  all  the  usual  functions  connected  with  tlie  repro- 
duction of  the  siDecies.  Many  instances  have  already  been  given 
of  the  partial  transference  of  secondary  masculine  characters  to 
the  female,  so  that  it  is  not  at  all  surprising  that  the  females  of 
some  species  should  possess  the  power  of  song.  It  has  also  been 
argued,  that  the  song  of  the  male  cannot  serve  as  a  charm,  be- 
cause the  males  of  certain  species,  for  instance  of  the  robin,  sing 
during  the  autumn.^^  But  nothing  is  more  common  than  for 
animals  to  take  pleasure  in  practising  whatever  instinct  they 
follow  at  other  times  for  some  real  good.  How  often  do  we  see 
birds  which  fly  easily,  gliding  and  sailing  through  the  air  ob- 
viously for  pleasure  ?  The  cat  plays  with  the  captured  mouse, 
and  the  cormorant  with  the  captured  fish.  The  weaver-bird 
(Ploceus),  when  confined  in  a  cage,  amuses  itself  by  neatly 
weaving  blades  of  grass  between  the  wires  of  its  cage.  Birds 
which  habitually  fight  during  the  breeding-season  are  generally 
ready  to  fight  at  all  times;  and  the  males  of  the  capercailzie 
sometimes  hold  their  Bahen  or  eks  at  the  usual  place  of  as- 
semblage during  the  autumn.^^  Hence  it  is  not  at  all  surprising 
that  male  birds  should  continue  singing  for  their  own  amuse- 
ment after  the  season  for  courtship  is  over. 

As  shewn  in  a  previous  chapter,  singing  is  to  a  certain  extent 
an  art,  and  is  much  imj^roved  by  practice.  Birds  can,  be  taught 
various  tunes,  and  even  the  unmelodious  sparrow  has  learnt  to 
sing  like  a  linnet.  They  acquire  the  song  of  their  foster 
parents,^^  and  sometimes  that  of  their  neighbours.^'^  All  the 
common  songsters  belong  to  the  Order  of  Insessores,  and  their 
vocal  organs  are  much  more  complex  than  those  of  most  other 
birds ;  j  et  it  is  a  singular  fact  that  some  of  the  Insessores,  such 
as  ravens,  cj  ows,  and  magpies,  possess  the  proper  apparatus,^'' 

32  D.  Barrington,  'Phil.  Transact.'      Beohstein,  ibid,  s,  5. 

1773,  p.  262.     Bechstein,  'Stuben-  3«  Dureau    de    la     Malle    gives  a 

vogel,' 1840,  s.  4.  curious  instance  ('Annales  des  Sc. 

33  This  is  likewise  the  case  with  Nat.'  3rd  series,  Zoolog.  torn.  x.  p. 
the  water-ouzel,  see  Mr.  Hepburn  118)  of  some  wild  blackbirds  in  his 
in  the  'Zoologist,*  1845-1846,  p.  garden  in  Paris,  which  naturally 
1068.  learnt  a  rej-^ublican  air  from  a  caged 

3^  L.    Lloyd,     'Game     Birds     of      bird. 
Sweden,'  1867,  p.  25.  :"  Bishop,  in  '  Todd's  Cyclop,  of 

3^  Barrington,      ibid.      p.      294,      Aa<t.  and  Phys.'  vol.  iv.  p.  1496. 


CiiAP.  XIII.  Vocar Music,  37' 


tliough  they  never  sing,  and  do  not  naturally  modulate  their 
voices  to  any  great  extent.  Hunter  asserts  ^^  that  with  the  true 
songsters  the  muscles  of  the  larynx  are  stronger  in  the  males 
than  in  the  females ;  but  with  this  slight  exception  there  is  no 
difference  in  the  vocal  organs  of  the  two  sexes,  although  the 
males  of  most  species  sing  so  much  better  and  more  continuously 
than  the  females. 

It  is  remarkable  that  only  small  birds  properly  sing.  The 
Australian  genus  Menura,  however,  must  be  excepted ;  for  the 
Meniira  Alhrti,  which  is  about  the  size  of  a  half-grown  turkey, 
not  only  mocks  other  birds,  but  "  its  own  whistle  is  exceedingly 
"  beautiful  and  varied."  The  males  congregate  and  form  "  cor- 
rohonjing  places,"  where  they  sing,  raising  and  spreading  their 
tails  like  peacocks,  and  drooping  their  wings.=^^  It  is  also  remark- 
able that  birds  which  sing  well  are  rarely  decorated  with  brilliant 
colours  or  other  ornaments.  Of  our  British  birds,  excepting 
the  bullfinch  and  goldfinch,  the  best  songsters  are  plain-coloured. 
The  kingfisher,  bee-eater,  roller,  hoopoe,  woodpeckers,  &c.,  utter 
harsh  cries ;  and  the  brilliant  birds  of  the  tropics  are  hardly  ever 
songsters.-'^ '  Hence  bright  colours  and  the  power  of  song  seem 
to  replace  each  other.  We  can  perceive  that  if  the  plumage  did 
not  vary  in  brightness,  or  if  bright  colours  were  dangerous  to  the 
species,  other  means  would  be  employed  to  charm  the  females ; 
and  melody  of  voice  offers  one  such  means. 

In  some  birds  the  vocal  organs  differ  greatly  in  the  two  sexes. 
In  the  Tetrao  cupido  (fig.  39)  the  male  has  two  bare,  orange- 
coloured  sacks,  one  on  each  side  of  the  neck ;  and  these  are 
largely  inflated  when  the  male,  during  the  breeding-season, 
makes  his  curious  hollow  sound,  audible  at  a  great  distance. 
Audubon  proved  that  the  sound  was  intimately  connected  with 
this  apparatus  (which  reminds  us  of  the  air-sacks  on  each  side  ot 
the  mouth  of  certain  male  frogs),  for  he  found  that  the  sound  was 
much  diminished  when  one  of  the  sacks  of  a  tame  bird  was 
pricked,  and  when  both  were  pricked  it  was  altogether  stopped. 
The  female  has  "  a  somewhat  similar,  though  smaller  naked  space 
"  of  skin  on  the  neck ;.  but  this  is  not  capable  of  inflation."  *^    The 

38  As  stated  by  Barrington  in  in  Canada,'  by  Major  W-  Ross  King, 
'Philosoph.  Transact.'  1773,  p.  262.  1866     pp.    144-146      Mr.    T     \\ ., 

39  Gould,  '  Handbook  to  the  Birds  Wood     gives     in     the     'Student 
of  Australia,'  vol.  i.  1865,  pp.  308-  (April,   1870    p.   lib)  an  excellen 
310      See  als^   Mr.  T.  W.  Wood  in  account  ot  the  attitude  and  habits 
the  "'Student,'  Auril  1870,  p.  125.  of  this   bird  during    its    courtship. 

^0  See  remarks  to  this  eflect  in  He  states  that  the  ear-tufts  or  neck- 
Gould's  '  Introduction  to  the  Trochi-  plumes  are  erected,  so^  that  they 
lidic'  1861,  p.  22.  meet  over  the  crown  ot    the  heaU. 

^'' '  The  Sportsman  and  Naturalist  See  his  drawing,  fig.  39. 


ZT^ 


The  Descent  of  Man. 


Part  II. 


male  of  another  kind  of  grouse  {Tetrao   urophasianus),   whilst 

conrtmg  the  female,  has  his  "  bare  yellow  cesophagns  inflated  to 

a  prodigious  size,  fully  half  as  large  as  the  body;"  and  he 

then  utters  various  grating,  deep,  hollow  tones.    With  his  neck- 


feathers  erect,  his  wmgs  lowered,  and  buzzing  on- the  gound, 
and  his  long  pointed  tail  spread  out  like  a  fan,  he  displays  a 
variety  of  grotesque  attitudes.  The  oesophagus  of  the  female  is 
not  m  any  way  remarkable.^^ 


"   Richardson, 'Fauna  Bor.  American 
bid.  vol.  iv.  p.  507. 


Birds,'  1831,  p.  359.    Audubon, 


Chap.  Xlll. 


Vocal  Music. 


373 


It  seems  now  well  made  out  that  the  great  throat  pouch  of 
the  European  male  bustard  ( Otis  tarda),  and  of  at  least  four 
other  species,  does  not,  as  was  formerly  supposed,  serve  to  hold 
water,  but  is  connected  with  the  utterance  during  the  breeding- 


Fig.  40.    The  Umbrella-bird  or  Cephalopterus  ornatus  (male,  from  Brehm). 

season  of  a  peculiar  sound  resembling  "ock."^^  A  crow-like 
bird  inhabiting  South  America  (Cephalopterus  ornatus,  fig.  40)  is 
called  the  umbrella-bird,  from  its  immense  top-knot,  formed  of 


*^  The  following  papers  have 
been  lately  written  on  this  subject : 
Prof.  A.  Newton,  in  the  '  Ibis,' 
1862,  p.  107  ;  Dr.  Cullen,  ibid.  1865, 
p.  14-5 ;  Mr.  Flower,  iu  '  Proc.  Zool. 
Soc'  1865,  p.  747 ;  and  Dr.  Murie, 
in 'Proc.  Zool.  Soc'  1868,  p.  471. 


In  this  latter  paper  an  excellent 
figure  is  given  of  the  male  Austra- 
lian Bustard  in  full  display  with 
the  sack  distended.  It  is  a  singular 
fact  that  the  sack  is  not  developed  m 
all  the  males  of  the  same  species. 


374  'The  Descent  of  Man.  Paut  IT. 

bare  white  quills  surmounted  by  dark-blue  plumes,  which  it  can 
elevate  into  a  great  dome  no  less  than  five  inches  in  diameter, 
covering  the  whole  head.  This  bird  has  on  its  neck  a  long,  thin, 
cylindrical  fleshy  appendage,  which  is  thickly  clothed  with  scale- 
like blue  feathers.  It  probably  serves  in  part  as  an  ornament, 
but  likewise  as  a  resounding  apparatus;  for  Mr.  Bates  found 
that  it  is  connected  "  with  an  unusual  development  of  the 
"  trachea  and  vocal  organs."  It  is  dilated  when  the  bird  utters  its 
singularly  deep,  loud  and  long  sustained  fluty  note.  The  head- 
crest  and  neck-appendage  are  rudimentary  in  the  female.^* 

The  vocal  organs  of  various  web-footed  and  wading  birds  are 
extraordinarily  complex,  and  difier  to  a  certain  extent  in  the  two 
sexes.  In  some  cases  the  trachea  is  convoluted,  hke  a  French 
horn,  and  is  deeply  embedded  in  the  sternum.  In  the  wild 
swan  {('yjuui^  ferns)  it  is  more  deeply  embedded  in  the  adult  male, 
than  in  the  adult  female  or  young  male.  In  the  male  Merganser 
the  enlarged  portion  of  the  trachea  is  furnished  with  an  additional 
pau'  of  muscles."*^  In  one  of  the  ducks,  however,  namely  Anas 
punctata,  the  bony  enlargement  is  only  a  Httle  more  developed 
in  the  male  than  in  the  female.^*'  But  the  meaning  of  these 
differences  in  the  trachea  of  the  two  sexes  of  the  Anatidae  is  not 
understood;  for  the  male  is  not  always  the  more  vociferous; 
thus  with  the  common  duck,  the  male  hisses,  whilst  the  female 
utters  a  loud  quack.*'  In  both  sexes  of  one  of  the  cranes  {(ires 
virgo)  the  trachea  penetrates  the  sternum,  but  presents  "  certain 
"  sexual  modifications."  In  the  male  of  the  black  stork  there  is 
also  a  well-marked  sexual  difference  in  the  length  and  curvature 
of  the  bronchi.'***  Highly  important  structures  have,  therefore,  in 
these  cases  been  modified  according  to  sex. 

It  is  often  difficult  to  conjecture  whether  the  many  strange 
cries  and  notes  uttered  by  male  birds  during  the  breeding- 
season,  serve  as  a  charm  or  merely  as  a  call  to  the  female.  The 
soft  cooing  of  the  turtle-dove  and  of  many  pigeons,  it  may  be 
presumed,  pleases  the  female.     When  the  female  of  the  wild 

■''*  Bates,  '  The  Naturalist  on  the  of  eight,  and  yet  this  bird  (Jerdon, 
Amazons,'  1863,  vol.  ii.  p.  284;  'Birds  of  India,'  vol.  iii.  p.  763)  is 
Wallace,  in  '  Proc.  Zool.  Soc'  1850,  mute ;  but  Mr.  Blyth  informs  me 
p.  206.  A  new  species,  with  a  still  that  the  convolutions  are  not  con- 
larger  neck-appeudage  (C.  pendu-  stantly  present,  so  that  perhaps 
li(jer),  has  lately  been  discovered,  they  are  now  tending  towards 
see  '  Ibis,'  vol.  i.  p.  457.  abortion. 

*'  Bishop,   in  Todd's  '  Cyclop,  of  *^  '  Elements  of  Comp.  Anat.'  by 

Anat.  and  Phys.'  vol.  iv.  p.  1499.  R.  Wagner,  Eng.  translat.   1845,  p. 

*^  Prof.    Newton,  'Proc.   Zoolog.  111.     With  respect  to  the  swan,  as 

Soc'  1871,  p.  651.  given     above,     Yarrell's    'Hist,    of 

"  The    spoonbill    (Platalea)   has  British  Birds,'  2nd  edit.  1845,  vol. 

its  trachea  convoluted  into  a  figure  iii.  p.  193. 


CfiAP.  XIII.  Iiistrinnental  Music.  375 

turkey  utters  her  call  in  the  morning,  the  male  answers  by  a  note 
which  differs  from  the  gobbling  noise  made,  when  with  erected 
feathers,  rustling  wings  and  distended  wattles,  he  puffs  and 
struts  before  her.*^  The  spe^  of  the  black-cock  certainly  serves 
as  a  call  to  the  female,  for  it  has  been  known  to  bring  four  or 
five  females  from  a  distance  to  a  male  under  confinement;  but 
as  the  black-cock  continues  his  spel  for  hours  during  successive 
days,  and  in  the  case  of  the  capercailzie  "  with  an  agony  of 
"  passion,"  we  are  led  to  suppose  that  the  females  which  are 
present  are  thus  charmed.^"  The  voice  of  the  common  rook  is 
known  to  alter  during  the  breeding-season,  and  is  therefore  in 
some  way  sexual, '^^  But  what  shall  we  say  about  the  harsh 
screams  of,  for  instance,  some  kinds  of  macaws;  have  these 
birds  as  bad  taste  for  musical  sounds  as  they  apparently  have 
for  colour,  judging  by  the  inharmonious  contrast  of  their  bright 
yellow  and  blue  plumage  ?  It  is  indeed  possible  that  without 
any  advantage  being  thus  gained,  the  loud  voices  of  many  male 
birds  may  be  the  result  of  the  inherited  effects  of  the  continued 
use  of  their  vocal  organs,  when  excited  by  the  strong  passions 
of  love,  jealousy  and  rage;  but  to  this  point  we  shall  recur 
when  we  treat  of  quadrupeds. 

We  have  as  yet  spoken  only  of  the  voice,  but  the  males  of 
various  birds  practise,  during  their  courtship,  what  may  be  called 
instrumental  music.  Peacocks  and  Birds  of  Paradise  rattle  their 
quills  together.  Turkey-cocks  scrape  their  wings  against  the 
ground,  and  some  kinds  of  grouse  thus  produce  a  buzzing  sound. 
Another  North  American  grouse,  the  Tetrao  umhell/is,  y^hen  with 
his  tail  erect,  his  ruffs  displayed,  "  he  shows  off  his  finery  to  the 
"  females,  who  lie  hid  in  the  neighbourhood,"  drums  by  rapidly 
striking  his  wings  together  above  his  back,  according  to  Mr.  E. 
Haymond,  and  not,  as  Audubon  thought,  by  striking  them 
against  his  sides.  The  sound  thus  produced  is  compared  by 
some  to  distant  thunder,  and  by  others  to  the  quick  roll  of  a 
drum.  The  female  never  drums, ''  but  flies  directly  to  the  place 
"  where  the  male  is  thus  engaged."  The  male  of  the  Kalij- 
pheasant,  in  the  Himalayas,  "  often  makes  a  singular  drumming 
"  noise  with  his  wings,  not  unlike  the  sound  produced  by  shaking 
"  a  stiff  piece  of  cloth."  On  the  west  coast  of  Africa  the  little 
black-weavers  (Ploceus  ?)  congregate  in  a  small  party  on  the 
bushes  round  a  small  open  space,  and  sing  and  glide  through 

<9  C.  L.  Bonaparte,  quoted  in  the  Sweden,'  &c.,  1867,  pp.  22,  81. 
*  Naturalist     Library:    Birds,'    vol.  *'  Jenner,    *  Piiilosoph.    Transac- 

xiv.  p.  126.  tions,'  1824,  p.  20. 

*"  L.  Llovd,  '  The  Game  Birds  of 


376  The  Descent  of  Man.  Part  II 

the  air  with  quivering  wings,  "  which  make  a  rapid  whirring 
"  sound  hke  a  child's  rattle."  One  bird  after  another  thus  performs 
for  hours  together,  but  only  during  the  courting-season.  At  this 
season  and  at  no  other  time,  the  males  of  certain  night-jars  (Capri- 
mulgus)  make  a  strange  booming  noise  with  their  wings.  The 
various  species  of  wood-peckers  strike  a  sonorous  branch  with 
their  beaks,  with  so  rapid  a  vibratory  movement  that  "  the  head 
"  appears  to  be  in  two  places  at  once."  The  sound  thus  ]3ro- 
duced  is  audible  at  a  considerable  distance,  but  cannot  be 
described ;  and  I  feel  sure  that  its  source  would  never  be  con- 
jectured by  any  one  hearing  it  for  the  first  time.  As  this  jarring 
sound  is  made  chiefly  during  the  breeding-season,  it  has  been 
considered  as  a  love-song ;  but  it  is  perhaps  more  strictly  a 
love-call.  The  female,  when  driven  from  her  nest,  has  been 
observed  thus  to  call  her  mate,  who  answered  in  the  same 
manner  and  soon  appeared.  Lastly,  the  male  Hoopoe  iUpiq^a 
epops)  combines  vocal  and  instrumental  music ;  for  during  the 
breeding-season  this  bird,  as  Mr.  Swinhoe  observed,  first  draws 
in  air,  and  then  taps  the  end  of  its  beak  perpendicularly  down 
against  a  stone  or  the  trunk  of  a  tree,  "  when  the  breath  being 
"  forced  down  the  tubular  bill  produces  the  correct  sound."  If 
the  beak  is  not  thus  struck  against  some  object,  the  sound  is 
quite  different.  Air  is  at  the  same  time  swallowed,  and  the 
oesojihagus  thus  becomes  much  swollen ;  and  this  probably  acts 
as  a  resonator,  not  only  with  the  hoojDoe,  but  with  pigeons  and 
other  birds.^^ 

In  the  foregoing  cases  sounds  are  made  by  the  aid  of  structures 
already  present  and  otherwise  necessary ;  but  in  the  following 
cases  certain  feathers  have  been  specially  modified  for  the  express 
purpose  of  producing  sounds.  The  drumming,  bleating,  neigh- 
ing, or  thundering  noise  (as  expressed  by  different  observers) 
made  by  the  common  snipe  {Scolopax  gallinago)  must  have  sur- 
prised every  one  who  has  ever  heard  it.  This  bird,  during  the 
pairing-season,  flies  to  "  perhaps  a  thousand  feet  in  height,"  ana 

^-    For   the  foregoing    facts    see,  Weavers,  'Livingstone's  Expeditioj? 

on     Birds      of    Paradise,     Brehm,  to  the  Zambesi,'  1865,  p.  425.     0» 

'  Thierleben,'  Band  iii.  s.  325.     On  Woodpeckers,    Macgillivray,    '  Hist 

Grouse,    Kichardson,    '  Fauna     Bor.  of  British  Birds,'  vol.  iii.  1840,  pp 

Americ. :  Birds,'  pp.  343  and  359  ;  84,  88,  89,  and  95.     On  the  Hoopoe 

Major  W.  Ross  King,  '  The  Sports-  Mr.  Swinhoe,  in  '  Proc.  Zoolog.  Soc' 

man  in  Canada,'  1866,  p.  156;  Mr.  June  23,    1863    and    1871,  p.  348. 

Hayraond,    in     Prof.     Cox's    '  Geol.  On    the    Night-jar,   Audubon,   ibid. 

Survey  of  Indiana,'  p.  227;  Audu-  vol.  ii.  p.  255,  and  'American  Natu- 

bon,    '  American    Ornitholog.    Bio-  ralist,'  1873,  p.  672.     The  English 

graph.'   vol.     i.     p.     216.     On   the  Night-jar    likewise    makes    in    the 

Kalij-pheasant,     Jerdon,    'Birds   of  spring    a    curious   noise   during  its 

India,'    vol.    iii.     p.    533.     On    the  rapid  flight. 


CHAr.  XIIJ. 


Iiistniinentat  Music. 


m 


after  zig-zagging  about  for  a  time  clcsconds  to  the  earth  in  a 
curved  line,  with  outspread  tail  and  quivering  pinions,  and 
surprising  velocity.  The  sound  is  emitted  only  during  this 
rapid  descent.  No  one  was  able  to  explain  the  cause,  until 
M.  Meves  observed  that  on  each  side  of  the  tail  the  outer  feathers 
are  peculiarly  formed  (fig.  41),  having  a  stitf  sabre-shaped  shaft 


Fig.  41.    Outer  tail-feather  of  Scolopax  gallinago  (from  '  Proc.  Zool.  See'  1858). 


with  the  oblique  barbs  of  unusual  length,  the  outer  webs  being 
strongly  bound  together.  He  found  that  by  blowing  on  these 
feathers,  or  by  fastening  them  to  a  long  thin  stick  and  waving 
them  rapidly  through  the  air,  he  could  reproduce  the  drumming 
noise  made  by  the  living  bird.  Both  sexes  are  furnished  with 
these  feathers,  but  they  are  generally  larger  in  the  male  than 
in  the  female,  and  emit  a 
deeper  note.  In  some 
species,  as  in  ^S'.  frenata 
(fig.  42),  four  feathers, 
and  in  S.  javensis  (fig.  43), 
no  less  than  eight  on  each 
side  of  the  tail  are  greatly 
modified.  Different  tones 
are  emitted  by  the  feathers 
of   the    different    species 


Fig.  42.     Outer  tail-feather  of  Scolopax  frenata. 


Fig.  43.    Outer  tail-feather  of  Scolopax  javensis. 


when  waved  through  the  air ;  and  the  Scolopax  Wilsonii  of  the 
United  States  makes  a  switching  noise  whilst  descending  rapidly 
to  the  earth.^^ 

In  the  male  of  the  Chamcepetes  unicolor  (a  large  gallinaceous 
bird  of  America)  the  first  primary  wing-feather  is  arched  towards 
the  tip  and  is  much  more  attenuated  than  in  the  female.  In  an 
allied  bird,  the  Penelope  nUjra,  Mr.  Salvin  observed  a  male,  which, 
whilst  it  flew  downwards  "  with  outstretched  wings,  gave  forth 
"  a  kind  of  crashing  rushing  noise,"  like  the  falling  of  a  tree.^* 


^*  See  M.  Meves'  interesting 
paper  in  '  Proc.  Zool.  Soc'  1858,  p. 
199.  For  the  habits  of  the  snipe, 
Macgillivray,  '  Hist.  British  Birds,' 
vol.  iv.  p.  371.  For  the  American 
snipe,  Capt.  Blakiston,  'J bis,'  vol.  v. 
1863,  p.  131. 


"  Mr.  Salvin,  in  'Proc.  Zool. 
Soc'  1867,  p.  160.  I  am  much  in- 
debted to  this  distinguished  orni- 
thologist for  sketches  of  the  feathers 
of  the  Chamaipetes,  and  for  other 
information. 


;78 


The  Descent  of  Man. 


Part  II. 


^^^^i^^^ 


Fig.  44.  Primary  wing-featber  of  a 
Humming-bird,  the  Selasphoriis  pla- 
tycercus  (from  a  sketcli  by  i\Ir.  Sal- 
vin).  Upper  figure,  that  of  male; 
lower  figure,  corresponding  feather 
of  female. 


The  male  alone  of  one  of  the  Indian  bustards  (Sypheotides 
aiLritus)  has  its  primary  wing-feathers  greatly  acuminated;  and 
the  male  of  an  allied  species  is  known  to  make  a  humming  noise 

whilst  courting  the  female. ^^  In 
a  widely  different  group  of  birds, 
namely  Humming-birds,  the  males 
alone  of  certain  kinds  haye  either 
the  shafts  of  their  priir.ary  wing- 
feathers  broadly  dilated,  or  the 
webs  abruptly  excised  towards 
the  extremity.  The  male,  for  in- 
stance, of  SelasjjJwrm  jilatycercus, 
when  adult,  has  the  first  primary 
wing-feather  (fig.  44),  thus  excised. 
Whilst  flying  from  flower  to  flower 
he  makes  "  a  shrill,  almost  whist- 
ling noise  ;"^^  but  it  did  not  appear  to  Mr.  Salvin  that  the  noise 
was  intentionally  made. 

Lastly,  in  several  species  of  a  sub-genus  of  Pipra  or  Manakin, 
the  males,  as  described  by  Mr.  Sclater,  have  their  secondary  wing- 
feathers  modified  in  a  still  more  remarkable  manner.  In  the 
brilliantly-coloured  P.  delkiom  the  first  three  secondaries  are 
thick-stemmed  and  curved  towards  the  body ;  in  the  fourth  and 
fifth  (fig.  45,  «)  the  change  is  greater;  and  in  the  sixth  and 
seventh  {h,  c)  the  shaft  "  is  thickened  to  an  extraordinary  degree, 
"  forming  a  solid  horny  lump."  The  barbs  also  are  greatly 
changed  in  shape,  in  comparison  with  the  corresponding  feathers 
(d,  e,  f)  in  the  female.  Even  the  bones  of  the  wing,  which 
support  these  singular  feathers  in  the  male,  are  said  by  Mr. 
Fraser  to  be  much  thickened.  These  little  birds  make  an 
extraordinary  noise,  the  first  "  sharp  note  being  not  unlike  the 
"  crack  of  a  whip."  ^^ 

The  diversity  of  the  sounds,  both  vocal  and  instrumental, 
made  by  the  males  of  many  birds  during  the  breeding-season, 
and  the  diversity  of  the  means  for  producing  such  sounds, 
are  highly  remarkable.  We  thus  gain  a  high  idea  of  their 
importance  for  sexual  purposes,  and  are  reminded  of  the  con- 
clusion arrived  at  as  to  insects.  It  is  not  difficult  to  imagine 
the  steps  by  which  the  notes  of  a  bird,  primar'ly  used  as  a 
mere  call  or  for  some  other  purpose,  might  have  been  improved 


"  Jerdon,  '  Birds  of  India,'  vol. 
iii.  pp.  618,  621. 

^^  Gould,  '  Introduction  to  the 
Trochilida?,'  18G1,  p.  49.  Salvin, 
'Proc.  Zoolog.  Soc'  1867,  p.  160. 


"  Sclater,  in  '  Proc.  Zool.  Soc* 
1860,  p.  90,  and  in  'Ibis,'  vol.  iv. 
1862,  p.  175.  Also  Salvin,  in 
'Ibis,'  1860,  p.  37. 


Chap.  XIII.     Vocal  a? id  Ifistnunental  Music. 


379 


into  a  melodious  love  song.  In  the  case  of  the  modified 
feathers,  by  which  the  drumming,  whistling,  or  roaring  noises 
are  produced,  we  know  that  some  birds  during  their  courtship 
flutter,  shake,  or  rattle  their  unmodiiied  feathers  together ;  and 


Fig.  45.  Secondary  wing-feathers  of  Pipra  deliciosa  (from  Mr.  Sclater,  in  •  Proc.  Zool. 
Sec'  1860).  The  three  upper  feathers,  a,  &,  c,  from  the  male;  the  three  lower  corre- 
sponding feathers,  d,  e,/,  from  tlie  female. 

a  and  d,  fifth  secondary  wing-feather  of  male  and  femalp,  upper  surface. 

6  and  e,  sixth  secondary,  upper  surface,    c  and/,  seventh  secondary,  lower  surface. 

if  the  females  were  led  to  select  the  best  performers,  the  males 
which  possessed  the  strongest  or  thickest,  or  most  attenuated 
feathers,  situated  on  any  part  of  the  body,  would  be  the 
most  successful;  and  thus  by  slow  degrees  the  feathers  might  be 
modified  to  almost  any  extent.  The  females,  of  course,  would 
not  notice  each  slight  successive  alteration  in  shape,  but  only 


380     .  The  Descent  of  Man.  Part  II. 

the  sounds  tlius  produced.  It  is  a  curious  fact  that  in  the  same 
class  of  animals,  sounds  so  different  as  the  drumming  of  the 
snii^e's  tail,  the  tapping  of  the  woodpecker's  beak,  the  harsh 
trumpet-like  cry  of  certain  water-fowl,  the  cooing  of  the  turtle- 
dove, and  the  song  of  the  nightingale,  should  all  be  pleasing  to 
the  females  of  the  several  species.  But  we  must  not  judge  of  the 
tastes  of  distinct  species  by  a  uniform  standard ;  nor  must  we 
judge  by  the  standard  of  man's  taste.  Even  with  man,  we 
should  remember  what  discordant  noises,  the  beating  of  tom- 
toms and  the  shrill  notes  of  reeds,  please  the  ears  of  savages. 
Sir  S.  Baker  remarks,^^  that  "  as  the  stomach  of  the  Arab  prefers 
"  the  raw  meat  and  reeking  liver  taken  hot  from  the  animal,  so 
"  does  his  ear  prefer  his  equally  coarse  and  discordant  music  to 
"  all  other." 

Love- Antics  and  Dances. — The  curious  love  gestures  of  some 
birds  have  already  been  incidentally  noticed  ;  so  that  little  need 
here  be  added.  In  Northern  America,  large  numbers  of  a  grouse, 
the  Tetrao  pliasianellas,  meet  every  morning  during  the  breeding- 
season  on  a  selected  level  spot,  and  here  they  run  round  and 
round  in  a  circle  of  about  fifteen  or  twenty  feet  in  diameter,  sc 
that  the  ground  is  worn  quite  bare,  like  a  fairy-ring.  In  these 
Partridge-dances,  as  they  are  called  by  the  hunters,  the  birds 
assume  the  strangest  attitudes,  and  run  round,  some  to  the  left 
and  some  to  the  right.  Audubon  describes  the  males  of  a  heron 
(Ardea  herodias)  as  walking  about  on  their  long  legs  with  great 
dignity  before  the  females,  bidding  defiance-  to  their  rivals.  With 
one  of  the  disgusting  carrion-vultures  (Cathartes  jota)  the  same 
naturalist  states  that  "  the  gesticulations  and  parade  of  the 
"  males  at  the  beginning  of  the  love-season  are  extremely 
"  ludicrous."  Certain  birds  perform  their  love  antics  on  the  wing, 
as  we  have  seen  with  the  black  African  weaver,  instead  of  on  the 
ground.  During  the  spring  our  little  white-throat  {Sylvia 
cinerea^  often  rises  a  few  feet  or  yards  in  the  air  above  some 
bush,  and  "flatters  with  a  fitful  and  fantastic  motion,  singing  all 
"  the  while,  and  then  drops  to  its  perch."  The  great  English 
bustard  throws  himself  into  indescribably  odd  attitudes  whilst 
courting  the  female,  as  has  been  figured  by  Wolf.  An  allied 
Indian  bustard  (O^is  feew^raZensis)  at  such  times  "rises  perpen- 
"  dicularly  into  the  air  with  a  hurried  flapping  of  his  wings, 
"  raising  his  crest  and  puffing  out  the  feathers  of  his  neck  and 
"  breast,  and  then  drops  to  the  ground ;"  he  repeats  this 
manoeuvre  several  times,  at  the  same  time  humming  in  a 
peculiar  tone.  Such  females  as  happen  to  be  near  "  obey  this 
^s  *  The  Nile  Tributaries  of  Abyssinia,'  1867,  p.  203. 


Chap.  XIII.        Love  Antics  and  Decollation.  381 

'*  saltatory  summons,"  and  when  tlicy  approach  ho  trails  his 
wmgs  and  spreads  his  tail  like  a  turkey-cock.''-' 

But  the  most  curious  case  is  afforded  by  three  allied  genera  of 
Australian  birds,  the  famous  ^ower-birds, — no  doubt  the  co- 
descendants  of  some  ancient  species  which  first  acquired  the 
strange  instinct  of  constructing  bowers  for  performing  their 
Jove-antics.  The  bowers  (fig.  46),  which,  as  we  shall  hereafter 
see,  are  decorated  with  feathers,  shells,  bones,  and  leaves,  are 
built  on  the  ground  foi'  the  sole  purpose  of  courtship,  for  their 
nests  are  formed  in  trees.  Both  sexes  assist  in  the  erection  of 
the  bowers,  but  the  male  is  the  principal  workman.  So  strong 
is  this  instinct  that  it  is  practised  under  confinement,  and  Mr. 
Strange  has  described  *^^  the  habits  of  some  Satin  Bower-birds 
which  he  kept  in  an  aviary  in  New  South  Wales.  "  At  times 
"  the  male  will  chase  the  female  all  over  the  aviary,  then  go  to 
"  the  bower,  pick  up  a  gay  feather  or  a  large  leaf,  utter  a  curious 
"  kind  of  note,  set  all  his  feathers  erect,  run  round  the  bower  and 
"  become  so  excited  that  his  eyes  appear  ready  to  start  from  his 
"  head ;  he  continues  opening  first  one  wing  then  the  other, 
"  uttering  a  low,  whistling  note,  and,  like  the  domestic  cock, 
"  seems  to  be  picking  up  something  from  the  ground  until  at 
"  last  the  female  goes  gently  towards  him."  Captain  Stokes  has 
described  the  habits  and  "  play-houses  "  of  another  species,  the 
Great  Bower-bird,  which  was  seen  "  amusing  itself  by  flying 
"  backwards  and  forwards,  taking  a  shell  alternately  from  each 
"  side,  and  carrying  it  through  the  archway  in  its  mouth." 
These  curious  structures,  formed  solely  as  halls  of  assemblage, 
where  both  sexes  amuse  themselves  and  pay  their  court,  must 
cost  the  birds  much  labour.  The  bower,  for  instance,  of  the 
Fawn-breasted  species,  is  nearly  four  feet  in  length,  eighteen 
inches  in  height,  and  is  raised  on  a  thick  platform  of  sticks. 

Decoration. — I  will  first  discuss  the  cases  in  which  the  males 
are  ornamented  either  exclusively  or  in  a  much  higher  degree 
than  the  females,  and  in  a  succeeding  chapter  those  in  which 
both  sexes  are  equally  ornamented,  and  finally  the  rare  cases  in 
which  the  female  is  somewhat  more  brightly-coloured  than  the 

^^    For    Tetrao  phasianellus,    see  the  Indian  Bustard,  Jerdon,  *  Birds 

Richardson,  'Fauna,  Bor.  America,'  of  India,'  vol.  iii.  p.  618. 
p.  361,  and  for  further  particulars  ^'^  Gould,  'Handbook  to  the  Birds 

Capt.  Blakiston, 'Ibis,'1863,  p.  125.  of  Australia,'  vol.    i.   pp.  444,449, 

For   the  Cathartes  and  Ardea,  Au-  455.     The     bower     of     the     Satin 

dubon,  '  Ornith.  Biography,' vol.   ii.  Bower-bird    may    be     seen    in    the 

p.  51,  and  vol.   iii.  p.  89.     On  the  Zoological     Society's    Gardens,    Ke- 

White-throat,   Macgillivray,    '  Hist.  gent's  Park. 
British  Birds,'  vol    ii.  p.  354.     On 


2>^2 


The  Descent  of  Man. 


Part  II. 


"  Seo  remarks   to  this  efTect 
the    'Feelin?     of 


Beauty 


on      Animals/    by  Mr.   J.  Shaw,  in  the 
among      '  Athena>um,'  Nov.  24th,  1866,  p.681. 


Chap.  XIII.  Decoration.  383 

at  the  commencement  of  this  chapter,  arc  wonderfully  diver- 
sified. The  plumes  on  the  front  or  back  of  the  head  consist  of 
variously-shaped  feathers,  sometimes  capable  of  erection  or 
expansion,  by  which  their  beautiful  colours  are  fully  displayed. 
Elegant  ear-tufts  (see  fig.  89,  ante)  are  occasionally  present. 
The  head  is  sometimes  covered  with  velvety  down,  as  with  the 
pheasant;  or  is  naked  and  vividly  coloured.  The  throat,  also,  is 
sometimes  ornamented  with  a  beard,  wattles,  or  caruncles.  Such 
appendages  are  generally  brightly-coloured,  and  no  doubt  serve 
as  ornaments,  though  not  always  ornamental  in  our  eyes;  for 
whilst  the  male  is  in  the  act  of  courting  the  female,  they  often 
swell  and  assume  vivid  tints,  as  in  the  male  turkey.  At  such 
times  the  fleshy  appendages  about  the  head  of  the  male  Tragopan 
pheasant  {Ceriorrcis  Temminckii)  swell  into  a  large  lappet  on  the 
throat  and  into  two  horns,  one  on  each  side  of  the  splendid 
top-knot ;  and  these  are  then  coloured  of  the  most  intense  blue 
which  I  have  ever  beheld.^^  The  African  hornbill  (Bucorax 
abyssinirus)  inflates  the  scarlet  bladder-like  wattle  on  its  neck, 
and  with  its  wings  drooping  and  tail  expanded  "  makes  quite  a 
"  grand  appearance." ^^  ii^ven  the  iris  of  the  eye  is  sometimes 
more  brightly-coloured  in  the  male  than  in  the  female;  and  this 
is  frequently  the  case  with  the  beak,  for  instance,  in  our  common 
blackbird.  In  Buceros  corrugatus,  the  whole  beak  and  immense 
casque  are  coloured  more  conspicuously  in  the  male  than  in  the 
female ;  and  "  the  oblique  grooves  upon  the  sides  of  the  lower 
"  mandible  are  peculiar  to  the  male  sex.'"'^ 

The  head,  again,  often  supports  fleshy  appendages,  filaments, 
and  solid  protuberances.  These,  if  not  common  to  both  sexes, 
are  always  confined  to  the  males.  The  solid  protuberances  have 
been  described  in  detail  by  Dr.  W.  Marshall,*^^  who  shews  that 
they  are  formed  either  of  cancellated  bone  coated  with  skin,  or 
of  dermal  and  other  tissues.  With  mammals  true  horns  are 
always  supported  on  the  frontal  bones,  but  with  birds  various 
bones  have  been  modified  for  this  purpose ;  and  in  species  of  the 
same  group  the  protuberances  may  have  cores  of  bone,  or  be 
quite  destitute  of  them,  with  intermediate  gradations  connecting 
these  two  extremes.  Hence,  as  Dr.  Marshall  justly  remarks, 
variations  of  the  most  different  kinds  have  served  for  the 
development  through  sexual  selection  of  these  ornamental 
appendages.     Elongated  feathers  or  plumes  spring  from  almost 

^2  See  Dr.  Murie's  account  with  ^^  'Land  and    Water,'    1868,    p. 

coloured    figures   in  '  Proc.    Zoolog.  217, 
Soc'  1872,  p.  730.  ^^  'Ueberdie  Schiidelhocker,' &c., 

63  Mr.    Monteiro,    'Ibis,'  vol.  iv.  '  Niederlandischeil  Archiv  fiir    Zoo- 

1862,  p.  339.  logic,'  B.  I.  Heft.  2,  1872. 


384  TJie  Descent  of  Man.  Part  II. 

every  part  of  the  body.  The  feathers  on  the  throat  and  breast 
are  sometimes  developed  into  beautiful  ruffs  and  collars.  The 
tail-feathers  are  frequently  increased  in  length ;  as  we  see  in  the 
tail-coverts  of  the  peacock,  and  in  the  tail  itself  of  the  Argus 
pheasant.  With  the  peacock  even  the  bones  of  the  tail  have 
been  modified  to  support  the  heavy  tail-coverts.*'^  The  body  of 
the  Argus  is  not  larger  than  that  of  a  fowl ;  yet  the  length  from 
the  end  of  the  beak  to  the  extremity  of  the  tail  is  no  less  than 
five  feet  three  inches/^  and  that  of  the  beautifully  ocellated 
secondary  wing-feathers  nearly  three  feet.  In  a  small  African 
night-jar  (^Cosmetornis  vexillarius)  one  of  the  primary  wing- 
feathers,  during  the  breeding -season,  attains  a  length  of  twenty- 
six  inches,  whilst  the  bird  itself  is  only  ten  inches  in  length.  In 
another  closely-allied  genus  of  night-jars,  the  shafts  of  the  elon- 
gated wing-feathers  are  naked,  except  at  the  extremity,  where 
there  is  a  disc.''^  Again,  in  another  genus  of  night-jars,  the 
tail-feathers  are  even  still  more  prodigiously  developed.  In 
general  the  feathers  of  the  tail  are  more  often  elongated  than 
those  of  the  wings,  as  any  great  elongation  of  the  latter  impedes 
flight.  AYe  thus  see  that  in  closely-allied  birds  ornaments  of 
the  same  kind  have  been  gained  by  the  males  through  the 
development  of  widely  different  feathers. 

It  is  a  curious  fact  that  the  feathers  of  species  belonging  to 
very  distinct  groups  have  been  modified  in  almost  exactly  the 
same  peculiar  manner.  Thus  the  wing-feathers  in  one  of  the 
above-mentioned  night-jars  are  bare  along  the  shaft,  and  ter- 
minate in  a  disc  ;  or  are,  as  they  are  sometimes  called,  spoon  or 
racket-shaped.  Feathers  of  this  kind  occur  in  the  tail  of  a 
motmot  (Eiwiomota  sujyerciliarifi),  of  a  king-fisher,  finch,  hum- 
ming-bird, i^arrot,  several  Indian  drongos  (Dicrurus  and  Edolius, 
in  one  of  which  the  disc  stands  vertically),  and  in  the  tail  of 
certain  birds  of  paradise.  In  these  latter  birds,  similar  feathers, 
beautifully  ocellated,  ornament  the  head,  as  is  likewise  the  case 
with  some  gallinaceous  birds.  In  an  Indian  bustard  {Sypheotides 
auritus)  the  feathers  forming  the  ear-tufts,  which  are  about  four 
inches  in  length,  also  terminate  in  discs.*'^  It  is  a  most  singular 
fact  that  the  motmots,  as  Mr.  Salvin  has  clearly  shewn,''^"  give  to 
their  tail  feathers  the  racket-shape  by  biting  off  the  barbs, 
and,  further,  that  this  continued  mutilation  has  produced  a 
certain  amount  of  inherited  effect. 

66  Dr.  W.  Marshall,  <Uber  den  Vog-  1864-,  p.  114.  Livingstone,  'Expedi- 
elschwanz,'  ibid.  B.  I.  Heft  2,  1872.       tion  to  the  Zambesi,'  1865,  p.  60. 

67  Jardine's 'Naturalist  Library:  69  Jgrdon,  'Birds  of  India,'  vol, 
Birds,'  vol.  xiv.  p.  166.                             iii.  p.  620. 

««  Sclater,  in  the  'Ibis,'  vol.  vi.  7»  '  Proc.Zoolog.Soc' 1873,p.  429. 


Chap.  XIII.  Decoration.  385 

Again,  the  barbs  of  the  feathers  hi  various  widely-distiuct 
birds  are  filamentous  or  phimose,  as  with  some  herons,  ibises, 
birds  of  paradise,,  and  Gallinacea3.  In  other  cases  the  barbs 
disappear,  leaving  the  shafts  bare  from  end  to  end ;  and  these  in 
the  tail  of  the  Faradisea  apodu  attain  a  length  of  thirty-four 
inches  :'^^  in  /'.  Papuana  (fig.  47)  they  are  much  shorter  and 
thin.  Smaller  feathers  when  thus  denuded  appear  like  bristles, 
as  on  the  breast  of  the  turkey-cock.  As  any  fleeting  fashion  in 
dress  comes  to  be  admired  by  man,  so  with  birds  a  change  of 
almost  any  kind  in  the  structure  or  colouring  of  the  feathers  in 
the  male  appears  to  have  been  admired  by  the  female.  The 
fact  of  the  feathers  in  widely  distinct  groups,  having  been 
modified  in  an  analogous  manner,  no  doubt  depends  primarily 
on  all  the  feathers  having  nearly  the  same  structure  and  manner 
of  development,  and  consequently  tending  to  vary  in  the  same 
manner.  We  often  see  a  tendency  to  analogous  variability  in 
the  plumage  of  our  domestic  breeds  belonging  to  distinct 
species.  Thus  top-knots  have  appeared  in  several  sjjecies.  In 
an  extinct  variety  of  the  turkey,  the  top-knot  consisted  of  bare 
quills  surmovmted  with  plumes  of  down,  so  that  they  somewhat 
resembled  the  racket-shaped  feathers  above  described.  In 
certain  breeds  of  the  pigeon  and  fowl  the  feathers  are  plumose, 
with  some  tendency  in  the  shafts  to  be  naked.  In  the  Sebas- 
topol  goose  the  scapular  feathers  are  greatly  eloD gated,  curled, 
or  even  spirally  twisted,  with  the  margins  plumose.'^ 

In  regard  to  colour  hardly  anything  need  here  be  said,  for 
every  one  knows  how  splendid  are  the  tints  of  many  birds,  and 
how  harmoniously  they  are  combined.  The  colours  are  often 
metallic  and  iridescent.  Circular  spots  are  sometimes  sur- 
rounded by  one  or  more  differently  shaded  zones,  and  are  thus 
converted  into  ocelli.  Nor  need  much  be  said  on  the  wonderful 
difference  between  the  sexes  of  many  birds.  The  common 
peacock  offers  a  striking  instance.  Female  birds  of  paradise 
are  obscurely  coloured  and  destitute  of  all  ornaments,  whilst  the 
males  are  probably  the  most  highly  decorated  of  all  birds,  and 
in  so  many  different  ways,  that  they  must  be  seen  to  be  appre- 
ciated. The  elongated  and  golden-orange  plumes  which  sj^ring 
from  beneath  the  wings  of  the  Paradisea  cqwda,  when  vertically 
erected  and  made  to  vibrate,  are  described  as  forming  a  sort  of 
halo,  in  the  centre  of  which  the  head  "  looks  like  a  little  emerald 
"  sun  with  its  rays  formed  by  the  two  plumes." '^^    In  another 

"  Wcallace,  in  '  Annals  and  Mag.  "  See  my  work  on  '  The  Variation 

of  Nat.  Hist.'  vol.  XX.  1857,  p.  416  ;  of  Animals  and  Plants  under  Duines- 

and    in    his     '  Malay   Archipelago,'  tication,'  vol.  i.  pp.  289,  293. 

vol.  ii.  1869,  p.  390.  "  Quoted  from  M.  de  Lafresnaye, 


3S6 


The  Descent  of  Man. 


Paet  U. 


most  beautiful  species  the  head  is  bald,  "and  of  a  rich  cobalt 
"  blue,  crossed  by  several  lines  of  black  yelyety  feathers." ^^ 


in  'Annals  and  Mag.  of  Nat.  Hist.' 
vol.  xiii.  1854,  p.  157  :  see  also  Mr. 
Wallace's  much  fuller  account  in 
vol.   XX.    1857,    p.    412,  and  in  his 


'  Malav  Archipelas^o,' 

^*  Wallace,    'The    Malar   Archi- 
pelago,' vol.  ii.  1869,  p.  405. 


Chap.  XIII. 


Decoration. 


3S7 


Male  humming-birds  (figs.  48  and  49)  almost  vie  with  birds  of 
paradise  in  their  beauty,  as  every  one  will  admit  who  has  seen 
Mr.  Gould's  splendid  volumes,  or  his  rich  collection.  It  is  very 
remarkable  in  how  many  difterent  ways  these  birds  are  orna- 
mented. Almost  every  part  of  their  plumage  lias  been  taken 
advantage  of,  and  modified;   and  the  modifications  have  been 


Fig.  48. 


Lophornis  ornatus,  male  and  female  (from  Brchm). 


^.fciNS:,z. 


388 


The  Descent  of  Man. 


Part  II. 


carried,  as  Mr.  Gould  shewed  me,  to  a  wonderful  extreme  in 
some  species  belonging  to  nearly  every  sub-group.  Such  cases 
are  curiously  hke  those  which  we  see  in  our  fancy  breeds, 
reared  by  man  for  the  sake  of  ornament:  certain  individuals 
originally  varied  in  one  character,  and  other  individuals  of  the 


Fig.  49. 


Spathura  underwoodi,  male  and  female  (from  Brchm). 


Chap.  XTII.  Decoration.  389 

same  species  in  other  characters ;  and  these  have  been  seized  on 
by  man  and  much  augmented— as  shewn  by  the  tail  of  the  fan- 
tail-pigeon, the  hood  of  the  jacobin,  the  beak  and  wattle  of  the 
carrier,  and  so  forth.  The  sole  difference  between  these  cases  is 
that  in  the  one,  the  result  is  due  to  man's  selection,  whilst  in 
the  other,  as  with  humming-birds,  birds  of  paradise,  &c.,  it  is 
due  to  the  selection  by  the  females  of  the  more  beautiful  males. 

I  will  mention  only  one  other  bird,  remarkable  from  the  ex- 
treme contrast  in  colour  between  the  sexes,  namely  the  famous 
bell-bird  {Ghasmorhynchus  niveiis)  of  S.  America,  the  note  of 
which  can  be  distinguished  at  the  distance  of  nearly  three  miles, 
and  astonishes  every  one  when  first  hearing  it.  The  male  is 
pure  white,  whilst  the  female  is  dusky-green ;  and  white  is  a 
very  rare  colour  in  terrestrial  species  of  moderate  size  and  in- 
offensive habits.  The  male,  also,  as  described  .by  Waterton, 
has  a  spiral  tube,  nearly  three  inches  in  length,  which  rises  from 
the  base  of  the  beak.  It  is  jet-black,  dotted  over  with  minute 
downy  feathers.  This  tube  can  be  inflated  with  air,  through  a 
communication  with  the  palate;  and  when  not  inflated  hangs 
down  on  one  side.  The  genus  consists  of  four  species,  the  maloe 
of  which  are  very  distinct,  whilst  the  females,  as  described  by 
Mr.  Sclater  in  a  very  interesting  paper,  closely  resemble  each 
other,  thus  offering  an  excellent  instance  of  the  common  rule 
that  within  the  same  group  the  males  differ  much  more  from 
each  other  than  do  the  females.  In  a  second  species  {C.  nudi- 
collis)  the  male  is  likewise  snow-white,  with  the  exception  of  a 
large  space  of  naked  skin  on  the  throat  and  round  the  eyes, 
wliich  during  the  breeding- season  is  of  a  fine  green  colour.  In 
a  third  species  {C.  tricarunculatus)  the  head  and  neck  alone  of 
the  male  are  white,  the  rest  of  the  body  being  chesnut-brown, 
and  the  male  of  this  species  is  provided  with  three  filamentous 
projections  half  as  long  as  the  body— one  rising  from  the  base 
of  the  beak,  and  the  two  others  from  the  corners  of  the  mouth.'^^ 

The  colom-ed  plumage  and  certain  other  ornaments  of  the 
adult  males  are  either  retained  for  life,  or  are  periodically  re- 
newed during  the  summer  and  breeding-season.  At  this  same 
season  the  beak  and  naked  skin  about  the  head  frequently 
change  colour,  as  with  some  herons,  ibises,  gulls,  one  of  the 
bell-birds  just  noticed,  &c.  In  the  white  ibis,  the  cheeks,  the 
inflatable  skin  of  the  throat,  and  the  basal  portion  of  the  beak 
then  become  crimson.^*^     In  one  of  the  rails,  Gallicrex  cristatus,  a 

"  Mr.  Sclater,  'Intellectual  Ob-  plate,  in  the  '  Ibis,'  1865,  p.  90. 

server,'    Jan.     1867.      '  Waterton's  ^«  'Land     and   Water,'    1867,    p. 

Wanderings,'  p.  118.     See  also  Mr.  394. 
Salvia's  interesting   paper,  with    a 


390  The  Descent  of  Man.  Part  II. 

large  red  caruncle  is  developed  during  this  period  on  the  head 
of  the  male.  So  it  is  with  a  thin  horny  crest  on  the  beak  of 
one  of  the  pelicans,  P.  erythrorhynchus ;  for  after  the  breeding- 
season,  these  horny  crests  are  shed,  like  horns  from  the  heads 
of  stags,  and  the  shore  of  an  island  in  a  lake  in  Nevada  was 
found  covered  with  these  curious  exuvise.'' 

Changes  of  colour  in  the  plumage  according  to  the  season 
depend,  firstly  on  a  double  annual  moult,  secondly  on  an  actual 
change  of  colour  in  the  feathers  themselves,  and  thirdly  on  theii* 
dull-coloured  margins  being  periodically  shed,  or  on  these  three 
processes  more  or  less  combined.  The  shedding  of  the  de- 
ciduary  margins  may  be  compared  with  the  shedding  of  their 
down  by  very  young  birds ;  for  the  down  in  most  cases  arises 
from  the  summits  of  the  fii'st  true  feathers.'* 

With  respect  to  the  birds  which  annually  undergo  a  double 
moult,  there  are,  firstly,  some  kinds,  for  instance  snipes,  swallow- 
plovers  (Glareolse),  and  curlews,  in  which  the  two  sexes  resemble 
each  other,  and  do  not  change  colour  at  any  season.  I  do  not 
know  whether  the  winter  plumage  is  thicker  and  warmer  than 
the  summer  plumage, 'but  warmth  seems  the  most  probable  end 
attained  of  a  double  moult,  where  there  is  no  change  of  colour. 
Secondly,  there  are  birds,  for  instance,  certain  species  of  Totanus 
and  other  Grail atores,  the  sexes  of  which  resemble  each  other, 
but  in  which  the  summer  and  winter  plumage  differ  slightly  in 
colour.  The  difference,  however,  in  these  cases  is  so  small  that 
it  can  hardly  be  an  advantage  to  them ;  and  it  may,  perhaps, 
be  attributed  to  the  direct  action  of  the  different  conditions  to 
which  the  birds  are  exposed  during  the  two  seasons.  Thirdly, 
there  are  many  other  birds  the  sexes  of  which  are  alike,  but 
which  are  widely  different  in  their  summer  and  winter  plumage. 
Fourthly,  there  are  birds,  the  sexes  of  which  differ  from  each 
other  in  colour  ;  but  the  females,  though  moulting  twice,  retain 
the  same  colours  throughout  the  year,  whilst  the  males  undergo  a 
change  of  colour,  sometimes  a  gi^eat  one,  as  with  certain  bustards. 
Fifthly  and  lastly,  there  are  birds  the  sexes  of  which  differ  from 
each  other  in  both  their  summer  and  winter  plumage ;  but  the 
male  undergoes  a  greater  amount  of  change  at  each  recurrent 
season  than  the  female— of  which  the  ruff  {Machetes  pagnax) 
offers  a  good  instance. 

With  respect  to  the  cause  or  purpose  of  the  differences  in 
colour  between  the  summer  aud  winter  plumage,  this  may  in 
some  instances,  as  with   the    ptarmigan,'^    serve  during  both 

"  Mr.  D.  G.  Elliot,  in  '  Proc.  edited  by  P  L.  Sclater.  Ray  Soc. 
Zool.  Soc'  1869,  p.  589.  1867,  p   14. 

^^  Nitzsch's        '  Pterylography,'  "^  The    brown    mottled     summer 


Chap.  XIIL  Double  Animal  Motilt.  391 

seasons  as  a  protection.  When  the  difference  between  the  two 
plumages  is  slight,  it  may  perhaps  be  attributed,  as  already 
remarked,  to  the  direct  action  of  the  conditions  of  life.  But 
with  many  birds  there  can  hardly  be  a  doubt  that  the  summer 
plumage  is  ornamental,  even  when  both  sexes  are  alike.  Wo 
may  conclude  that  this  is  the  case  with  many  herons,  egrets, 
&c.,  for  they  acquire  their  beautiful  plumes  only  during  the 
breeding-season.  Moreover,  such  jDlumes,  top-knots,  &c.,  though 
possessed  by  both  sexes,  are  occasionally  a  little  more  developed 
in  the  male  than  in  the  female ;  and  they  resemble  the  plumes 
and  ornaments  possessed  by  the  males  alone  of  other  birds.  It 
is  also  known  that  confinement,  by  affecting  the  reproductive 
system  of  male  birds,  frequently  checks  the  development  of  their 
secondary  sexual  characters,  but  has  no  immediate  influence 
on  any  other  characters ;  and  I  am  informed  by  Mr.  Bartlett 
that  eight  or  nine  specimens  of  the  Knot  {Trinya  canutm)  re- 
tained their  unadorned  winter  plumage  in  the  Zoological  Gardens 
throughout  the  year,  from  which  fact  we  may  infer  that  the 
summer  jDlumage  though  common  to  both  sexes  partakes  of  the 
nature  of  the  exclusively  masculine  plumage  of  many  other 
birds.^° 

From  the  foregoing  facts,  more  especially  from  neither  sex  of 
certain  birds  changing  colour  during  either  annual  moult,  or 
changing  so  slightly  that  the  change  can  hardly  be  of  any  service 
to  them,  and  from  the  females  of  other  species  moulting  twice 
yet  retaining  the  same  colours  throughout  the  year,  we  may 
conclude  that  the  habit  of  annually  moulting  twice  has  not 
been  acquired  in  order  that  the  male  should  assume  an  orna- 
mental character  during  the  breeding- season ;  but  that  the 
double  moult,  having  been  originally  acquired  for  some  distinct 
purpose,  has  subsequently  been  taken  advantage  of  in  certain 
cases  for  gaining  a  nuptial  plumage. 

It  appears  at  first  sight  a  surprising  circumstance  that  some 
closely-allied  species  should  regularly  undergo  a  double  annual 


plumage  of  the  ptarmigan  is  of  as  statements     on     moulting,    see,    on 

much  importance    to  it,   as  a  pro-  snipes,     &c.,     Macgillivray,    '  Hist, 

tection,  as    the   white   winter   plu-  Brit.    Birds,'    vol.    iv.    p.    371;    on 

mage ;  for    in    Scandinavia,    during  Glareolse,     curlews,    and     bustards, 

the    spring,     when    the    snow    has  Jerdon,  '  Birds  of  India,'  vol,  iii.  pp. 

disappeared,  this  bird  is  known  to  615,  630,  683;  on  Totanus,  ibid.  p. 

suffer    greatly  from   birds  of  prey,  700  ;  on  the  plumes  of  herons,  ibid, 

before  it  has   acquired  its  summer  p.  738,   and    Macgillivray,   vol.   iv. 

dress  :  see  Wilhelm  von  Wright,  in  pp.  435  and  444,  and  Mr.   Stafford 

Lloyd,    'Game     Birds    of    Sweden,'  Allen,  in  the  '  Ibis,' vol.  v.  1863,  p. 

1867,  p.  125.  33. 
^^  In    regard     to    the     previous 


392  The  Descent  of  Man.  Part  II. 

moult,  and  others  only  a  single  one.  The  ptarmigan,  for 
instance,  moults  twice  or  even  thrice  in  the  year,  and  the  black- 
cock only  once  :  some  of  the  splendidly  coloured  honey-suckers 
(Nectarinise)  of  India  and  some  sub-genera  of  obscurely  coloured 
pipits  (Anthus)  have  a  double,  whilst  others  have  only  a  single 
annual  moult.^^  But  the  gradations  in  the  manner  of  moulting, 
which  are  known  to  occur  with  various  birds,  shew  us  how 
species,  or  whole  groups,  might  have  originally  acquii'ed  their 
double  annual  moult,  or  having  once  gained  the  habit,  have 
again  lost  it.  With  certain  bustards  and  plovers  the  vernal 
moult  is  far  from  complete,  some  feathers  being  renewed,  and 
some  changed  in  colour.  There  is  also  reason  to  believe  that 
with  certain  bustards  and  rail-like  birds,  which  properly  under- 
go a  double  moult,  some  of  the  older  males  retain  their 
nuptial  plumage  throughout  the  year.  A  few  highly  modified 
feathers  may  merely  be  added  during  the  spring  to  the  plumage, 
as  occurs  with  the  disc-formed  tail-feathers  of  certain  drongos 
(^BhringcL)  in  India,  and  with  the  elongated  feathers  on  the 
back,  neck,  and  crest  of  certain  herons.  By  such  steps  as  these, 
the  vernal  moult  might  be  rendered  more  and  more  complete, 
until  a  perfect  double  moult  was  acquired.  Some  of  the  birds 
of  paradise  retain  their  nuptial  feathers  throughout  the  year, 
and  thus  have  only  a  single  moult ;  others  cast  them  directly 
after  the  breeding-season,  and  thus  have  a  double  moult ;  and 
others  again  cast  them  at  this  season  during  the  first  year,  but 
not  afterwards ;  so  that  these  latter  species  are  intermediate  in 
their  manner  of  moulting.  There  is  also  a  great  difference  with 
many  birds  in  the  length  of  time  during  which  the  two  annual 
plumages  are  retained ;  so  that  the  one  might  come  to  be  re- 
tained for  the  whole  year,  and  the  other  completely  lost.  Thus 
in  the  spring  Machetes  pugnax  retains  his  ruff  for  barely  two 
months.  In  Natal  the  male  widow-bird  (Chera  progne)  acquires 
his  fine  plumage  and  long  tail-feathers  in  December  or  January, 
and  loses  them  in  March;  so  that  they  are  retained  only  for 
about  three  months.  Most  species,  which  undergo  a  double 
moult,  keep  their  ornamental  feathers  for  about  six  months. 
The  male,  however,  of  the  wild  Gcdlus  hankiva  retains  his  neck- 
hackles  for  nine  or  ten  months ;  and  when  these  are  cast  off,  the 
underlying  black  feathers  on  the  neck  are  fully  exposed  to  view. 
But  with  the  domesticated  descendant  of  tliis  species,  the  neck- 
hackles  of  the  male  are  immediately  replaced  by  new  ones  ;  so 

81  On  the  moulting  of  the  ptar-  359,  365,  369.     On  the  moulting  of 

migan,  sje  Gould's  'Birds  of  Great  Anthus,  see  Blyth,  in  'Ibis,'  1867, 

Britain.'     On     the     honey-suckers,  p.  32, 
Jerdon,  '  Birds  of  India,'  vol.  i.  pp. 


Chap.  .XTll.  Dotible  Aimtial  Moidt.  393 


that  we  here  see,  as  to  part  of  the  phimage,  a  double  moult 
changed  under  domestication  into  a  single  moult.^^ 

The  common  drake  (^Anas  loschas)  after  the  breeding- season 
is  well  known  to  lose  his  male  plumage  for  a  period  of  three 
months,  during  which  time  he  assumes  that  of  the  female.  The 
male  pintail-duck  (Ands  acuta)  loses  his  plumage  for  the  shorter 
period  of  six  weeks  or  two  months ;  and  Montagu  remarks  that 
"  this  double  moult  within  so  short  a  time  is  a  most  extra- 
"  ordinary  circumstance,  that  seems  to  bid  defiance  to  all  human 
"  reasoning."  But  the  believer  in  the  gradual  modification  of 
species  will  be  far  from  feeling  surprise  at  finding  gradations  of 
all  kinds.  If  the  male  pintail  were  to  acquire  his  new  plumage 
within  a  still  shorter  period,  the  new  male  feathers  would 
almost  necessarily  be  mingled  with  the  old,  and  both  with  some 
proper  to  the  female ;  and  this  apparently  is  the  case  with  the 
male  of  a  not  distantly-allied  bird,  namely  the  Merganser  serrator, 
for  the  males  are  said  to  "  undergo  a  change  of  plimiage,  which 
"  assimilates  them  in  some  measure  to  the  female."  By  a  little 
further  acceleration  in  the  process,  the  double  moult  would  be 
completely  lost.^^ 

Some  male  birds,  as  before  stated,  become  more  brightly 
coloured  in  the  spring,  not  by  a  vernal  moult,  but  either  by  an 
actual  change  of  colour  in  the  feathers,  or  by  their  obscurely- 
coloured  deciduary  margins  being  shed.  Changes  of  colour  thus 
caused  may  last  for  a  longer  or  shorter  time.  In  the  Pdecaims 
onocrotaliis  a  beautiful  rosy  tint,  with  lemon-coloured  marks  on 
the  breast,  overspreads  the  whole  plumage  in  the  spring ;  but 
these  tints,  as  Mr.  Sclater  states, "  do  not  last  long,  disappearing 
"  generally  in  about  six  weeks  or  two  months  after  they  have 
"  been  attained."  Certain  finches  shed  the  margins  of  their 
feathers  in  the  spring,  and  then  become  brighter  coloured,  while 
other  finches  undergo  no  such  change.  Thus  the  Frhigilla  tristis 
of  the  United  States  (as  well  as  many  other  American  species) 

^2  For  the   foregoing    statements  of  the  Herodias  bub'ilcus,  Mr.  S.  S. 

in    regard  to    partial    moults,    and  Allen,  in  'Ibis,'   1863,    p.  33.     Ou 

on  old  males  retaining  their  nuptial  Gallus    banhiva,  Blyth,  in   '  Annals 

plumage,    see    Jerdou,   on   bustards  and  Mag.  of  Nat.  Hist.' vol.  i.  1848, 

and  plovers,  in  '  Birds  of  India,' vol.  p.  455;  see,' also,   on  this   subject, 

iii.    pp.   617,    637,  709,    711.     Also  my    'Variation    of    Animals    under 

Blyth  in  '  Land  and  Water,'  1867,  Domestication,'  vol.  i.  p.  236. 

p.    84.     On  the  moulting  of  Para-  ^^  See  Macgillivray, '  Hist.  British 

disea,  see  an  interesting  article  by  Birds '  (vol.  v.  pp.  34,  70,  and  223), 

Dr.  W.  Marshall, 'Archives  Neerlan-  on    the    moulting    of   the   AnatidjB, 

daises,'  torn.  vi.  1871.  Ou  the  Vidua,  with  quotations  from  Watorton  and 

'Ibis,'    vol.    iii.    1861,   p.    133.     On  Montagu.      Also    Yarrell,    'Hist,   of 

the     Drongo-shrikes,    Jerdon,    ibid.  British  Birds,'  vol.  iii.  p.  243. 
vol.  i.  p.  435.     On  the  vernal  moult 
18 


394  •  The  Descent  of  Man.  Part  II. 

exhibits  its  bright  colours  only  when  the  winter  is  past,  whilst 
our  goldfinch,  which  exactly  represents  this  bird  in  habits,  and 
our  siskin,  which  represents  it  still  more  closely  in  structure, 
undergo  no  such  annual  change.  But  a  difference  of  this  kind 
in  the  plumage  of  allied  species  is  not  surprising,  for  with  the 
common  linnet,  which  belongs  to  the  same  family,  the  crimson 
forehead  and  breast  are  displayed  only  during  the  summer  in 
England,  whilst  in  Madeira  these  colours  are  retained  through- 
out the  year.^^ 

Di^ilay  hy  Male  Birds  of  their  Flumage. — Ornaments  of  all 
kinds,  whether  permanently  or  temporarily  gained,  are  sedulously 
displayed  by  the  males,  and  apparently  serve  to  excite,  attract, 
or  fascinate  the  females.  But  the  males  will  sometimes  display 
their  ornaments,  when  not  in  the  presence  of  the  females,  as 
occasionally  occurs  with  grouse  at  their  balz-places,  and  as  may 
be  noticed  with  the  peacock ;  this  latter  bird,  however,  evidently 
wishes  for  a  spectator  of  some  kind,  and,  as  I  have  often  seen, 
will  shew  off  his  finery  before  poultry,  or  even  pigs.^^  All  natu- 
ralists who  have  closely  attended  to  the  habits  of  birds,  whether 
in  a  state  of  nature  or  under  confinement,  are  unanimously  of 
opinion  that  the  males  take  delight  in  displaying  their  beauty. 
Audubon  frequently  speaks  of  the  male  as  endeavouring  in 
various  ways  to  charm  the  female.  Mr.  Gould,  after  describing 
some  peculiarities  in  a  male  humming-bird,  says  he  has  no  doubt 
that  it  has  the  power  ofdisplaying  them  to  the  greatest  advantage 
before  the  female.  Dr.  Jerdon  ^^  insists  that  the  beautiful  plum- 
age of  the  male  serves  "  to  fascinate  and  attract  the  female." 
Mr.  Bartlett,  at  the  Zoological  Gardens,  expressed  himself  to  me 
in  the  strongest  terms  to  the  same  effect. 

It  must  be  a  grand  sight  in  the  forests  of  India  "  to  come 
*' suddenly  on  twenty  or  thirty  pea  fowl,  the  males  displaying 
"  their  gorgeous  trains,  and  strutting  about  in  all  the  pomp  ot 
'  pride  before  the  gratified  females."  The  wild  turkey-cock 
erects  his  glittering  plumage,  expands  his  finely-zoned  tail  and 
barred  wing-feathers,  and  altogether,  with  his  crimson  and 
blue  wattles,  makes  a  superb,  though  to  our  eyes,  grotesque 

*^  On  the  pelican,  see  Sclater,  in  1863,  p.  230. 
'Proc,    Zool.     Soc'    1868,     p.    265.  *^  See  also  '  Ornamental  Poultry,' 

On   the    American   finches,  see  Au-  by  Rev.  E.  S.  Dixon,  1848,  p.  8. 
dubon,  '  Ornith.  Biography,'  vol.  i.  ^^  '  Birds  of  India,' introduct.  vol. 

pp.  174,  221,  and  Jerdon,  'Birds  of  i.  p.  xxiv. ;  on  the   peacock,  vol.  iii. 

India,'    vol.    ii.     p.     383.     On    the  p.  507.     See   Gould's  '  Introduction 

FringiUa  canna'nna  of  Madeira,  Mr.  to  the  Trochilidae,'  1861,  pp.  15  and 

E.  Vernon  Harcourt,  'Ibis,'  vol.  v.,  111. 


Chap.  XIII. 


Display  by  the  Male. 


395 


appearance.  Similar  facts  have  already  been  given  with  respect 
to  grouse  of  Yarious  kinds.  Turning  to  another  Order,  The 
male  Rupicola  crocea  (fig.  50)  is  one  of  the  most  beautiful  birds 
in  the  world,  being  of  a  splendid  orange,  with  some  of  the 
feathers  curiously  truncated  and  plumose.  The  female  is  brown- 
ish-green, shaded  with  red,  and  has  a  much  smaller  crest.  Sir 
R.  Schomburgk  has  described  their  courtship ;  he  found  one 


Fig.  50. 


Kupicola  crocea,  male  (T.  W.  Wood). 


of  their  meeting-places  where  ten  males  and  two  females  were 
present.  The  space  w^as  from  four  to  five  feet  in  diameter,  and 
appeared  to  have  been  cleared  of  every  blade  of  grass  and 
smoothed  as  if  by  human  hands.  A  male  "  was  capering,  to  the 
"  apparent  dehght  of  several  others.  Now  spreading  its  wings, 
"  throwing  up  its  head,  or  opening  its  tail  like  a  fan ;  now 
'-  strutting   about  with  a  hopping  gait  until    tired,  when  it 


39^  The  Descent  of  Man.  Part  II. 

"  gabbled  some  kind  of  note,  and  was  relieved  by  another.  Thus 
•'  three  of  them  successively  took  the  field,  and  then,  with  self- 
"  approbation,  withdrew  to  rest."  The  Indians,  in  order  to 
obtain  their  skins,  wait  at  one  of  the  meeting-places  till  the 
birds  are  eagerly  engaged  in  dancing,  and  then  are  able  to  kill 
with  their  poisoned  arrows  four  or  five  males,  one  after  the 
other.*^  With  birds  of  paradise  a  dozen  or  more  full-plumaged 
males  congregate  in  a  tree  to  hold  a  dancing-party,  as  it  is 
called  by  the  natives :  and  here  they  fly  about,  raise  their  wings, 
elevate  their  exquisite  plumes,  and  make  them  vibrate,  and 
the  whole  tree  seems,  as  Mr.  Wallace  remarks,  to  be  filled  with 
waving  plumes.  When  thus  engaged,  they  become  so  absorbed 
that  a  skilful  archer  may  shoot  nearly  the  whole  party.  These 
birds,  when  kept  in  confinement  in  the  Malay  Archipelago,  are 
said  to  take  much  care  in  keeping  their  feathers  clean;  often 
spreading  them  out,  examining  them,  and  removing  every  speck 
of  dirt.  One  observer,  who  kei)t  several  pairs  alive,  did  not  doubt 
that  the  display  of  the  male  was  intended  to  please  the  female.^^ 
The  Gold  and  Amherst  pheasants  during  their  courtship  not 
only  expand  and  raise  their  splendid  frills,  but  twist  them,  as  I 
have  myself  seen,  obliquely  towards  the  female  on  whichever  side 
she  may  be  standing,  obviously  in  order  that  a  large  surface  may 
be  displayed  before  her.^^^  They  likewise  turn  their  beautiful 
tails  and  tail-coverts  a  little  towards  the  same  side.  Mr.  Bartlett 
has  observed  a  male  Polyplectron  (fig.  51)  in  the  act  of  court- 
ship, and  has  shewn  me  a  specimen  stufi"ed  in  the  attitude 
then  assumed.  The  tail  and  wing-feathers  of  this  bird  are 
ornamented  with  beautiful  ocelli,  like  those  on  the  peacock's 
train.  Now  when  the  peacock  displays  himself,  he  expands  and 
erects  his  tail  transversely  to  his  body,  for  he  stands  in  front 
of  the  female,  and  has  to  shew  off,  at  the  same  time,  his  rich 
blue  throat  and  breast.  But  the  breast  of  the  Polyplectron  is 
obscurely  coloured,  and  the  ocelli  are  not  confined  to  the  tail- 
feathers.  Consequently  the  Polyplectron  does  not  stand  in  front 
of  the  female ;  but  he  erects  and  expands  his  tail-feathers  a  little 
obliquely,  lowering  the  expanded  wing  on  the  same  side,  and 
raising  that  on  the  opposite  side.  In  tliis  attitude  the  ocelli 
over  the  whole  body  are  exposed  at  the  same  time  before  the 

8'  *  Journal  of  R.  Geograph.  Soc'  ^^  Mr.    T.    W.    Wood    has    given 

vol.  X.  1840,  p.  236.  ('  The  Student,'  April  1870,  p.  115) 

**  '  Annals  and  Mag.  of  Nat.  Hist.'  a   full  account  of  this    manner    of 

vol.  xiii.  1854,  p.  157  ;  also  Wallace,  display,  by  the  Gold  pheasant  and 

ibid.  vol.  XX.  1857,  p.  412,  and 'The  by    the     Japanese     pheasant,     Ph. 

Malay  Archipehigo,'  vol.  ii.  1869,  p.  versicolor ;  and  he  calls  it  the  lateral 

252.     Also   Dr.   Bennett,  as  quoted  or  one-sided  display, 
by  Brehm,  'Thierleben,'  B.  iii.  s.  326. 


Chap.  XIII. 


Display  by  the  Male. 


397 


eyes  of  the  admiring  female  in  one  grand  bespangled  expanse. 
To  whichever  side  she  may  turn,  the  expanded  wings  and  the 
obliquely-held  tail  are  turned  towards  her.  The  male  Tragopan 
pheasant  acts  in  nearly  the  same  manner,  for  he  raises  the 
feathers  of  the  body,  though  not  the  wing  itself,  on  the  side 


B'ig.  51. 


Polyplectiou  chinquis,  male  (T.  W.  Wood). 


39^  The  Descent  of  Man.  Fart  II. 

which  is  opposite  to  the  female,  and  wliich  would  otherwise  be 
concealed,  so  that  nearly  all  the  beautifully  sj)otted  feathers  are 
exhibited  at  the  same  time. 

The  Argus  pheasant  affords  a  much  more  remarkable  case.  The 
immensely  developed  secondary  wing-feathers  are  confined  to  the 
male ;  and  each  is  ornamented  with  a  row  of  from  twenty  to 
twenty-three  ocelli,  above  an  inch  in  diameter.  These  feathers 
are  also  elegantly  marked  with  oblique  stripes  and  rows  of  spots 
of  a  dark  colour,  like  those  on  the  skin  of  a  tiger  and  leopard 
combined.  These  beautiful  ornaments  are  hidden  until  the  male 
shews  himself  off  before  the  female.  He  then  erects  his  tail,  and 
expands  his  wing-feathers  into  a  great,  almost  upright,  circular 
fan  or  shield,  which  is  carried  in  front  of  the  body.  The  neck 
and  head  are  held  on  one  side,  so  that  they  are  concealed  by  the 
fan ;  but  the  bird  in  order  to  see  the  female,  before  whom  he  is 
displaying  himself,  sometimes  pushes  his  head  between  two  of 
the  long  wing- feathers  (as  Mr.  Bartlett  has  seen),  and  then  pre- 
sents a  grotesque  appearance.  This  must  be  a  frequent  habit 
with  the  bird  in  a  state  of  nature,  for  Mr.  Bartlett  and  his  son  on 
examining  some  perfect  skins  sent  from  the  East,  found  a  place 
between  two  of  the  feathers,  which  was  much  frayed,  as  if  the 
head  had  here  frequently  been  pushed  through.  Mr.  Wood 
thinks  that  the  male  can  also  peep  at  the  female  on  one  side, 
beyond  the  margin  of  the  fan. 

The  ocelli  on  the  wing-feathers  are  wonderful  objects;  for  they 
are  so  shaded  that,  as  the  Duke  of  Argyll  remarks,^*^  they  stand 
out  hke  balls  lying  loosely  within  sockets.  When  I  looked  at 
the  specimen  in  the  British  Museum,  which  is  mounted  with  the 
wings  expanded  and  trailing  downwards,  I  was  however  greatly 
disappointed,  for  the  ocelli  appeared  flat,  or  even  concave.  But 
Mr.  Gould  soon  made  the  case  clear  to  me,  for  he  held  the 
feathers  erect,  in  the  position  in  wliich  they  w^ould  naturally  be 
displayed,  and  now,  from  ttie  hght  shining  on  them  from  above, 
each  ocellus  at  once  resembled  the  ornament  called  a  ball  and 
socket.  These  feathers  have  been  shewn  to  several  artists,  and 
all  have  expressed  their  admiration  at  the  perfect  shading.  It 
may  well  be  asked,  could  such  artistically  shaded  ornaments 
have  been  formed  by  means  of  sexual  selection  ?  But  it  will  be 
convenient  to  defer  giving  an  answer  to  this  question,  until  wo 
treat  in  the  next  chapter  of  the  principle  of  gradation. 

The  foregoing  remarks  relate  to  the  secondary  wing-feathers, 

but  the  primary  wing-feathers,  which  in  most  gaUinaceous  birds 

are  uniformly  coloured,  are  in  the  Argus    pheasant  equally 

wonderful.     They  are  of  a  soft  brown  tint  with  numerous  dark 

50  'The  Reign  of  Law,'  18G7,  p.  203. 


Chap.  XIII. 


Display  by  the  Male. 


399 


spots  each  of  which  consists  of  two  or  tliree  black  dots  with  a 
surrounding  dark  zone.  But  the  chief  ornament  is  a  space 
paraUel  to  the  dark-blue  shaft,  which  in  outline  forms  a  perfect 
second  feather  lying  within  the  true  feather.  This  inner  part  is 
coloured  of  a  lighter  chesnut.  and  is  thickly  dotted  with  minute 


Fis  52     Side  view  of  male  Argus  pheasant,  whilst  displaying  before  the  female 
nhsprvpd  and  sketched  from  nature  by  Mr.  T.  \V.  Wood. 


Observed  and  sketched  from 


400  The  Descent  of  Man.  Part  11. 

white  points.  I  have  shewn  this  feather  to  several  persons,  and 
many  have  admired  it  even  more  than  the  ball  and  socket 
feathers,  and  have  declared  that  it  was  more  like  a  work  of  art 
than  of  nature.  Now  these  feathers  are  quite  hidden  on  all 
ordinary  occasions,  but  are  fully  displayed,  together  with  the 
long  secondary  feathers,  when  they  are  all  expanded  together  so 
as  to  form  the  great  fan  or  shield. 

The  case  of  the  male  Argus  jDlieasant  is  eminently  interesting, 
because  it  affords  good  evidence  that  the  most  refined  beauty  may 
serve  as  a  sexual  charm,  and  for  no  other  purpose.  We  must  con- 
clude that  this  is  the  case,  as  the  secondary  and  primary  wing- 
feathers  are  not  at  all  displayed,  and  the  ball  and  socket  orna- 
ments are  not  exhibited  in  full  perfection,  until  the  male  assumes 
the  attitude  of  coui'tship.  The  Argus  pheasant  does  not  possess 
brilliant  colours,  so  that  his  success  in  love  appears  to  depend  on 
the  great  size  of  his  plumes,  and  on  the  elaboration  of  the  most 
elegant  patterns.  Many  wilf  declare  that  it  is  utterly  incredible 
that  a  female  bird  should  be  able  to  appreciate  fine  shading  and 
exquisite  patterns.  It  is  undoubtedly  a  marvellous  fact  that  she 
should  possess  this  almost  human  degree  of  taste.  He  who 
thinks  that  he  can  safely  gauge  the  discrimination  and  taste  o  t* 
the  lower  animals  may  deny  that  the  female  Argus  pheasant  can 
appreciate  such  refined  beauty ;  but  he  will  then  be  compelled 
to  admit  that  the  extraordinary  attitudes  assumed  by  the  male 
during  the  act  of  courtship,  by  which  the  wonderful  beauty  of 
his  plumage  is  fully  displayed,  are  purjDoseless ;  and  this  is  a 
conclusion  which  I  for  one  will  never  admit. 

Although  so  many  pheasants  and  allied  gaUinaceous  birds 
carefully  display  their  plumage  before  the  females,  it  is  remark- 
able, as  Mr.  Bartlett  informs  me,  that  tliis  is  not  the  case  with  the 
dull-coloured  Eared  and  Cheer  pheasants  {CrossoptiJon  auritum 
and  Phasiarms  tcallichii) ;  so  that  these  birds  seem  conscious 
that  they  have  little  beauty  to  display.  Mr.  Bartlett  has  never 
seen  the  males  of  either  of  these  species  fighting  together,  though 
he  has  not  had  such  good  opportunities  for  observing  the  Cheer 
as  the  Eared  pheasant.  Mr.  Jenner  Weir,  also,  finds  that  all 
male  birds  with  rich  or  strongly-characterised  plumage  are 
more  quarrelsome  than  the  dull-coloured  species  belonging  to 
the  same  groups.  The  goldfinch,  for  instance,  is  far  moi;p 
pugnacious  than  the  linnet,  and  the  blackbird  than  the  thrush. 
Those  birds  Avhich  undergo  a  seasonal  change  of  jilumage  like- 
wise becomes  much  more  pugnacious  at  the  period  when  they 
are  most  gaily  ornamented.  No  doubt  the  males  of  some 
obscui'ely-coloured  birds  fight  desperately  together,  but  it 
appears  that  when  sexual  selection  has  been  highly  influential. 


Chap.  XIII  Display  by  the  Male.  401 

and  has  given  bright  colours  to  the  males  of  any  species,  it  has 
also  very  often  given  a  strong  tendency  to  pngnacity,  \Ve  shall 
meet  with  nearly  analogous  cases  when  we  treat  of  mammals. 
On  the  other  hand,  with  birds  the  power  of  song  and  brilliant 
colours  have  rarely  been  both  acquired  by  the  males  of  the  same 
species ;  but  in  this  case,  the  advantage  gained  would  have  been 
the  same,  namely,  success  in  charming  the  female.  Nevertheless 
it  must  be  owned  that  the  males  of  several  brilliantly  coloured 
birds  have  had  their  feathers  specially  modified  for  the  sake  of 
producing  instrumental  music,  though  the  beauty  of  this  cannot 
be  compared,  at  least  according  to  our  taste,  M'ith  that  of  the 
vocal  music  of  many  songsters. 

We  will  now  turn  to  male  birds  which  are  not  ornaniented  in 
any  high  degree,  but  which  nevertheless  display  during  their 
courtship  whatever  attractions  they  may  possess.  These  cases 
are  in  some  respects  more  curious  than  the  foregoing,  and  have 
been  but  little  noticed.  I  owe  the  following  facts  to  Mr.  Weir, 
wdio  has  long  kept  confined  birds  of  many  kinds,  including  all  tbe 
British  Fringillida)  and  Emberizida3.  The  facts  have  been  selected 
from  a  large  body  of  valuable  notes  kindly  sent  me  by  him.  The 
bullfinch  makes  his  advances  in  front  of  the  female,  and  then 
puffs  out  his  breast,  so  that  many  more  of  the  crimson  feathers 
are  seen  at  once  than  otherwise  would  be  the  case.  At  the 
same  time  lie  twists  and  bows  his  black  tail  from  side  to  side  in 
a  ludicrous  manner.  The  male  chaflSnch  also  stands  in  front  of 
the  female,  thus  shewing  his  red  breast  and  "  blue  bell,"  as  the 
fanciers  call  his  head  ;  the  ^A•ings  at  the  same  time  being  slightly 
expanded,  with  the  pure  white  bands  on  the  shoulders  thus 
rendered  conspicuous.  The  common  linnet  distends  his  rosy 
breast,  slightly  expands  his  brown  wings  and  tail,  so  as  to  make 
the  best  of  them  by  exhibiting  their  white  edgings.  We  must, 
however,  be  cautious  in  concluding  that  the  wings  are  spread 
out  solely  for  display,  as  some  birds  do  so  whose  wings  are 
not  beautiful.  This  is  the  case  with  the  domestic  cock,  but  it 
is  always  the  wing  on  the  side  opposite  to  the  female  which  is 
expanded,  and  at  the  same  time  scraped  on  the  ground.  The 
male  goldfinch  behaves  diiferently  from  all  other  finches :  his 
wings  are  beautiful,  the  shoulders  being  black,  with  the  dark- 
tipped  wing-feathers  spotted  with  white  and  edged  with  golden 
yellow.  When  he  courts  the  female,  he  sways  his  body  from 
side  to  side,  and  quickly  turns  his  slightly  expanded  wings  first 
to  one  side,  then  to  the  other,  with  a  golden  flashing  effect. 
Mr.  Weir  informs  me  that  no  other  British  finch  turns  thus 
from  side  to  side  during  his  courtship,  not  even  the  closely- 
allied  male  siskin,  for  he  would  not  thus  add  to  his  beauty. 


402  TJie  Descent  of  Man.  Part  II. 

Most  of  the  British  Buntings  are  plain  coloured  birds  ;  but  in 
the  spring  the  feathers  on  the  head  of  the  male  reed-bnnting 
{Emheriza  schmiicvlus)  acquire  a  fine  black  colour  by  the 
abrasion  of  the  dusky  tips;  and  these  are  erected  during  the 
act  of  courtship.  Mr.  Weir  has  kef)t  two  species  of  Amadina 
from  Australia :  the  A.  caftan ot is  is  a  very  small  and  chastely 
coloured  finch,  with  a  dark  tail,  white  rump,  and  jet-black  upper 
tail-coverts,  each  of  the  latter  being  marked  with  three  large  con- 
spicuous oval  spots  of  w^hite.^^  This  species,  wdien  courting  the 
female,  slightly  spreads  out  and  vibrates  these  parti-coloured 
tail-coverts  in  a  very  i^eculiar  manner.  The  male  Amadina 
Lnthami  behaves  very  differently,  exhibiting  before  the  female 
his  brilliantly  spotted  breast,  scarlet  rump,  and  scarlet  upper 
tail-coverts.  I  may  here  add  from  Dr.  Jerdon  that  the  Indian 
bulbul  (Fi/cnonotus  Iwemorrhous)  has  its  under  tail-coverts  of  a 
crimson  colour,  and  these,  it  might  be  thought,  could  never  be 
"Well  exhibited ;  but  the  bird  "  w  hen  excited  often  spreads  them 
"  out  laterally,  so  that  they  can  be  seen  even  from  above."  ^- 
The  crimson  under  tail-coverts  of  some  other  birds,  as  with  one 
of  the  woodpeckers,  Ficus  major,  can  be  seen  without  any  such 
display.  The  common  pigeon  has  iridescent  feathers  on  the 
breast,  and  every  one  must  have  seen  how  the  male  inflates 
his  breast  whilst  courting  the  female,  thus  shewing  them  off 
to  the  best  advantage.  One  of  the  beautiful  bronze -winged 
pigeons  of  Australia  {Ocyphaps  JophoUs)  behaves,  as  described  to 
me  by  Mr.  Weir,  very  differently :  the  male,  whilst  standing 
before  the  female,  lowers  his  head  almost  to  the  ground,  spreads 
out  and  raises  his  tail,  and  half  expands  his  wings.  He  then 
alternately  and  slowly  raises  and  depresses  his  body,  so  that  the 
iridescent  metallic  feathers  are  all  seen  at  once,  and  glitter  in 
the  sun. 

Sufficient  facts  have  now  been  given  to  shew"  with  what  care 
male  birds  disj^lay  their  various  charms,  and  this  they  do  with 
the  utmost  skill.  Whilst  preening  their  feathers,  they  haver 
frequent  opportunities  for  admiring  themselves,  and  of  studying 
how  best  to  exhibit  their  beauty.  But  as  all  the  males  of  the 
same  species  display  themselves  in  exactly  the  same  manner,  it 
apjDcars  that  actions,  at  first  perhaps  intentional,  have  become 
instinctive.  If  so,  w^e  ought  not  to  accuse  birds  of  conscious 
vanity ;  yet  when  we  see  a  peacock  strutting  about,  with  ex- 
panded and  quivering  tail-feathers,  he  seems  the  very  emblem 
of  pride  and  vanity. 

'^  -Foi    the    description   of    these      p.  417. 
birds,  see  Gould's  'Handbook  to  the  ^^  'jJii^js  of  India,'  vol.  ii.  p.  96. 

Birds   of    Australia,'   vol.    i.    1865, 


Chap.  XIII.  Display  by  the  Male.  403 

The  various  ornaments  possessed  by  the  males  are  certainly 
of  the  highest  importance  to  them,  for  in  some  cases  they  have 
been  acquired  at  the  expense  of  greatly  impeded  powers  of 
flight  or  of  running.  The  African  night-jar  (Cosmetornis),  which 
during  the  pairing-season  has  one  of  its  primary  wing-feathers 
developed  into  a  streamer  of  very  great  length,  is  thereby  much 
retarded  in  its  flight,  although  at  other  times  remarkable  for  its 
swiftness.  The  "  unwieldy  size  "  of  the  secondary  wing-feathers 
of  the  male  Argus  pheasant  are  said  "  almost  entirely  to  deprive 
"  the  bird  of  flight."  The  fine  plumes  of  male  birds  of  jDaradise 
trouble  them  during  a  high  wind.  The  extremely  long  tail- 
feathers  of  the  male  widow-birds  (Vidua)  of  Southern  Africa 
render  "  their  flight  heavy ; "  but  as  soon  as  these  are  cast  off 
they  fly  as  well  as  the  females.  As  birds  always  breed  when 
food  is  abundant,  the  males  probably  do  not  suffer  much  in- 
convenience in  searching  for  food  from  their  impeded  powers 
of  movement ;  but  there  can  hardly  be  a  doubt  that  they  must 
be  much  more  liable  to  be  struck  down  by  birds  of  prey.  Nor 
can  we  doubt  that  the  long  train  of  the  peacock  and  the  long 
tail  and  wing-feathers  of  the  Argus  pheasant  must  render  them 
an  easier  prey  to  any  prowling  tiger-cat,  than  would  otherwise 
be  the  case.  Even  the  bright  colours  of  many  male  birds  cannot 
fail  to  make  them  conspicuous  to  their  enemies  of  all  kinds. 
Hence,  as  Mr.  Gould  has  remarked,  it  probably  is  that  such 
birds  are  generally  of  a  shy  disposition,  as  if  conscious  that  their 
beauty  was  a  source  of  danger,  and  are  much  more  difficult 
to  discover  or  approach,  than  the  sombre  coloured  and  compa- 
ratively tame  females,  or  than  the  young  and  as  yet  unadorned 
males.^^ 

It  is  a  more  curious  fact  that  the  males  of  some  birds  which 
are  provided  with  special  weapons  for  battle,  and  which  in  a 
state  of  nature  are  so  pugnacious  that  they  often  kill  each  other, 
suffer  from  possessing  certain  ornaments.  Cock-fighters  trim 
the  hackles  and  cut  off  the  combs  and  gills  of  their  cocks ;  and 
the  birds  are  then  said  to  be  dubbed.  An  undubbed  bird,  as  Mr. 
Tegetmeier  insists,  "  is  at  a  fearful  disadvantage ;  the  comb  and 
'*  gills  offer  an  easy  hold  to  his  adversary's  beak,  and  as  a  cock 
"  always  strikes  where  he  holds,  when  once  he  has  seized  his  foe, 

"'  On  the  Cosmetornis,  see  Living-  widow-bird,    Barrow's   *  Travels    in 

stone's  '  Expedition  to  the  Zambesi,'  Africa,'   vol.  i.  p.    243,  and  '  Ibis,' 

1865,  p.  66.  On  the  Argus  pheasant,  vol,  iii.  1861,  p.   l.')3.     Mr.   Gould, 

Jardine's  'Nat.  Hist.  Lib.:    Birds,'  on  the  shyness  of  male  birds, '  Hand- 

vol.    xiv.     p.     167.     On     Birds     of  book  to  Birds  of  Australia,'  vol.  i. 

Paradise,  Lesson,  quoted  by  Brehm,  1865,  pp.  210,  457. 
*  Thierleben,'  B.  iii.  s.  325.     On  the 


404  The  Descent  of  Man.  Part  II. 

"  he  has  him  entirely  in  his  power.  Even  supposing  that  the 
"  bird  is  not  killed,  the  loss  of  blood  suffered  by  an  nndnbhed 
"  cock  is  much  greater  than  that  sustained  by  one  that  has  been 
"  trimmed."  ^^  Young  turkey-cocks  in  iighting  always  seize  hold 
of  each  other's  wattles ;  and  I  jDresume  that  the  old  birds  fight  in 
the  same  manner.  It  may  perhaps  be  objected  that  the  comb 
and  wattles  are  not  ornamental,  and  cannot  be  of  service  to  the 
birds  in  this  way ;  but  even  to  our  eyes,  the  beauty  of  the  glossy 
black  Spanish  cock  is  much  enhanced  by  his  white  face  and 
crimson  comb ;  and  no  one  who  has  ever  seen  the  sj^lendid  blue 
wattles  of  the  male  Tragopan  pheasant,  distended  in  courtship, 
can  for  a  moment  doubt  that  beauty  is  the  object  gained.  From 
the  foregoing  facts  we  clearly  see  that  the  plumes  and  other 
ornaments  of  the  males  must  be  of  the  highest  imjDortance  to 
them ;  and  we  further  see  that  beauty  is  even  sometimes  more 
important  than  success  in  battle. 


CHAPTEK   XIV. 

Birds— con  ^mttec?. 


Choice  exerted  by  the  female — Length  of  courtship — Unpaired  birds — 
Mental  qualities  and  taste  for  the  beautiful — Preference  or  antipathy 
shewn  by  the  female  for  particular  males — Variability  of  birds — Varia- 
tions sometimes  abrupt — Laws  of  variation — Formation  of  ocelli — 
Gradations  of  character — Case  of  Peacock,  Argus  pheasant,  and  Urosticte. 

When  the  sexes  differ  in  beauty,  or  in  the  power  of  singing,  or 
in  producing  what  I  have  called  instrumental  music,  it  is  almost 
invariably  the  male  who  surpasses  the  female.  These  qualities, 
as  we  have  just  seen,  are  evidently  of  high  importance  to  the 
male.  When  they  are  gained  for  only  a  part  of  the  year  it  is 
always  before  the  breeding-season.  It  is  the  male  alone  who 
elaborately  displays  his  varied  attractions,  and  often  performs 
strange  antics  on  the  ground  or  in  the  aii*,  in  the  presence  of  the 
female.  Each  male  drives  away,  or  if  he  can,  kills  his  rivals. 
Hence  we  may  conclude,  that  it  is  the  object  of  the  male  to  induce 
the  female  to  pair  with  him,  and  for  this  purpose  he  tries  to  ex- 
cite or  charm  her  in  various  ways ;  and  tliis  is  the  opinion  of  all 
those  who  have  carefully  studied  the  habits  of  living  birds.  But 
there  remains  a  question  which  has  an  all-important  bearing  on 
sexual  selection,  namely,  does  every  male  of  the  same  species 
excite  and  attract  the  female  equally?  Or  does  she  exert  a  choice, 
and  prefer  certain  males  ?  This  latter  question  can  be  answered 
in  the  affirmative  by  much  dii*ect  and  indirect  evidence.  It  is  far 
»^  Tegetmeier,  '  The  Poultry  Book,'  1866,  p.  139. 


CiiAP.  XIV.  Length  of  Coiirtship,  405 

more  cliflBcult  to  decide  what  qualities  determine  the  clioice  of  the 
females ;  but  here  again  we  have  some  direct  and  indirect  evi- 
dence that  it  is  to  a  large  extent  the  external  attractions  of  the 
male ;  though  no  doubt  his  vigour,  courage,  and  other  mental 
qualities  come  into  play.  \Ye  will  begin  with  the  indirect  evidence. 
Length  of  Courtship. — The  lengthened  period  during  which 
both  sexes  of  certain  birds  meet  day  after  day  at  an  appointed 
place  probably  dej^ends  partly  on  the  courtship  being  a  pro- 
longed affair,  and  partly  on  reiteration  in  the  act  of  pairing. 
Thus  in  Germany  and  Scandinavia  the  balzeng  or  leks  of  the 
black-cocks  last  from  the  middle  of  Blarch,  all  through  April 
into  May,  As  many  as  forty  or  fifty,  or  even  more  birds  con- 
gregate at  the  leks;  and  the  same  place  is  often  frequented 
diQ-ing  successive  years.  The  lek  of  the  capercailzie  lasts  from 
the  end  of  March  to  the  middle  or  even  end  of  May.  In  North 
America  "  the  jDartridge  dances "  of  the  Tttrao  phasianellus 
"  last  for  a  month  or  more."  Other  kinds  of  grouse,  both  in 
North  America  and  Eastern  Siberia,^  follow  nearly  the  same 
habits.  The  fowlers  discover  the  hillocks  where  the  ruffs  con- 
gregate by  the  grass  being  trampled  bare,  and  this  shews  that 
the  same  spot  is  long  frequented.  The  Indians  of  Guiana  are 
well  acquainted  with  the  cleared  arenas,  where  they  exi:)ect  to 
find  the  beautiful  cocks  of  the  Eock ;  and  the  natives  of  New 
Guinea  know  the  trees  where  from  ten  to  twenty  male  birds  of 
paradise  in  full  plumage  congregate.  In  this  latter  case  it  is 
not  exjDressly  stated  that  the  females  meet  on  the  same  trees,  but 
the  hunters,  if  not  sj^ecially  asked,  would  i^robably  not  mention 
their  jDresence,  as  their  skins  are  valueless.  Small  parties  of  an 
African  weaver  (Ploceus)  congregate,  during  the  breeding-season, 
and  perform  for  hours  their  graceful  evolutions.  Large  numbers 
of  the  Solitary  snipe  {Scolojjax  major)  assemble  during  dusk  in 
a  morass  ;  and  the  same  place  is  frequented  for  the  same  purpose 
during  successive  years ;  here  they  may  be  seen  running  about 
"  like  so  many  large  rats,"  puffing  out  theii'  feathers,  flapping 
theii'  wings,  and  uttering  the  strangest  cries.^ 

^  Nordman  describes  ('  Bull.  Soc.  of  the  above  named  grouse  see 
Imp.  des  Nat.  Moscou,'  1861,  torn.  Brehm,  '  Thievleben,'  B.  iv.  s.  350; 
xxxiv.  p.  264)  the  balzen  of  Tetrao  also  L.  Lloyd,  *  Game  Birds  of 
urogalloides  in  Amur  Land.  He  Sweden,' 1867,  pp.  19, 78.  Richard- 
estimated  the  number  of  birds  son,  '  Fauna  Bor.  Americana,'  Birds, 
assembled  at  above  a  hundred,  not  p.  362.  Keferences  in  rec^ard  to  the 
counting  the  females,  which  lie  hid  assemblages  of  other  birds  have 
in  the  surrounding  bushes.  The  already  been  given.  On  Paradisea 
noises  uttered  ditler  from  those  of  see  Wallace,  in  *  Annals  and  Mag. 
T.  urofiallus.  of  Nat.  Hist.'  vol.  xx.  1857,  p.  412. 

2  With  respect  to  the  assemblages  On  the  sni])e,  Lloyd,  ibid.  p.  221. 


406  The  Descent  of  Man.  Part  II. 

Some  of  the  above  birds, — the  black-cock,  capercailzie, 
pheasant-grouse,  ruff.  Solitary  snipe,  and  perhaps  others, — are,  as 
is  believed,  polygamists.  With  such  birds  it  might  have  been 
thought  that  the  stronger  males  would  simply  have  driven  away 
the  weaker,  and  then  at  once  have  taken  possession  of  as  many 
females  as  possible ;  but  if  it  be  indispensable  for  the  male  to 
excite  or  please  the  female,  we  can  understand  the  length  of  the 
courtship  and  the  congregation  of  so  many  individuals  of  both 
sexes  at  the  same  spot.  Certain  strictly  monogamous  species 
likewise  hold  nuptial  assemblages ;  this  seems  to  be  the  case  in 
Scandinavia  with  one  of  the  ptarmigans,  and  their  leks  last  from 
the  middle  of  March  to  the  middle  of  May.  In  Australia  the 
lyre-bird  {Menura  superba)  forms  "small  round  hillocks,"  and 
the  M.  Alberti  scratches  for  itself  shallow  holes,  or,  as  they  are 
called  by  the  natives,  conohorying  places^  where  it  is  believed 
both  sexes  assemble.  The  meetings  of  the  M.  superba  are  some- 
times very  large ;  and  an  account  has  lately  been  published  ^  by 
a  traveller,  who  heard  in  a  valley  beneath  him,  tliickly  covered 
with  scrub,  "a  din  which  completely  astonished"  him;  on 
crawling  onwards  he  beheld  to  his  amazement  about  one  hun- 
dred and  fifty  of  the  magnificent  lyre-cocks,  "  ranged  in  order  of 
"  battle,  and  fighting  with  indescribable  fury."  The  bowers  of 
the  Bower-birds  are  the  resort  of  both  sexes  during  the  breeding- 
season  ;  and  "  here  the  males  meet  and  contend  with  each  other 
"  for  the  favours  of  the  female,  and  here  the  latter  assemble  and 
"  coquet  with  the  males."  With  two  of  the  genera,  the  same 
bower  is  resorted  to  during  many  years.* 

The  common  magpie  (Corvus  pica,  Linn,),  as  I  have  been  in- 
formed by  the  Eev.  W.  Darwin  Fox,  used  to  assemble  from  all 
parts  of  Delamere  Forest,  in  order  to  celebrate  the  "  great  mag- 
pie marriage."  Some  years  ago  these  birds  abounded  in  extra- 
ordinary numbers,  so  that  a  gamekeeper  killed  in  one  morning 
nineteen  males,  and  another  killed  by  a  single  shot  seven  birds 
at  roost  together.  They  then  had  the  habit  of  assembling  very 
early  in  the  spring  at  particular  spots,  where  they  could  be  seen 
in  flocks,  chattering,  sometimes  fighting,  bustling  and  flying 
about  the  trees.  The  whole  affair  was  evidently  considered  by 
the  birds  as  one  of  the  highest  importance.  Shortly  after  the 
meeting  they  all  separated,  and  were  then  observed  by  Mr.  Fox 
and  others  to  be  paired  for  the  season.  In  any  district  in  which 
a  species  does  not  exist  in  large  numbers,  gi-eat  assemblages 
cannot,  of  course,  be  held,  and   the   same   species  may  have 

.  3  Quoted  by  Mr.  T.  W.  Wood  in      of  Australia,'  vol.  1.  pp.   300,  308, 

the  '  Student,'  April,  1870,  p.  125.        448,  451.     On  the  ptarmigan,  above 

-  Gould,  '  Handbook  to  the  Birds      alluded  to,  see  Lloyd,  ibid.  p.  129. 


Chap.  XIV.  Unpaired  Birds.  407 

different  habits  in  different  countries.  For  instance,  I  have 
heard  of  only  one  instance,  from  Mr.  Wedderburn,  of  a  regular 
assemblage  of  black  game  in  Scotland,  yet  these  assemblages 
are  so  well  kno\Yn  in  Germany  and  Scandinavia  that  they  have 
received  special  names. 

Uiqxiircd  Birds. — From  the  facts  now  given,  we  may  conclude 
that  the  courtship  of  birds,  belonging  to  widely  different  groups, 
is  often  a  prolonged,  delicate,  and  troublesome  affiiir.  There  is 
even  reason  to  suspect,  improbable  as  this  will  at  first  appear, 
that  some  males  and  females  of  the  same  species,  inhabiting  the 
same  district,  do  not  always  please  each  other,  and  consequently 
do  not  pair.  Many  accounts  have  been  published  of  either  the 
male  or  female  of  a  pair  having  been  shot,  and  quickly  replaced 
by  another.  This  has  been  observed  more  frequently  with  the 
magpie  than  with  any  other  bird,  owing  perhaps  to  its  conspic- 
uous appearance  and  nest.  The  illustrious  Jenner  states  that 
in  Wiltshire  one  of  a  pair  was  daily  shot  no  less  than  seven 
times  successively,  "but  all  to  no  purpose,  for  the  remaining 
"  magpie  soon  found  another  mate " ;  and  the  last  pair  reared 
their  young.  A  new  partner  is  generally  found  on  the  succeed- 
.ing  day  ;  but  Mr.  Thompson  gives  the  case  of  one  being  replaced 
on  the  evening  of  the  same  day.  Even  after  the  eggs  are  hatched, 
if  one  of  the  old  birds  is  destroyed  a  mate  will  often  be  found ;  this 
occurred  after  an  interval  of  two  days,  in  a  case  recently  observed 
by  one  of  Sir  J.  Lubbock's  keepers.^  The  first  and  most  obvious 
conjecture  is  that  male  magpies  must  be  much  more  numerous 
than  females ;  and  that  in  the  above  cases,  as  well  as  in  many 
others  which  could  be  given,  the  males  alone  had  been  killed. 
This  apparently  holds  good  in  some  instances,  for  the  game- 
keepers in  Delamere  Forest  assured  Mr.  Fox  that  the  magpies 
and  carrion-crows  which  they  formerly  killed  in  succession  in 
large  numbers  near  their  nests,  were  all  males ;  and  they  accounted 
for  this  fact  by  the  males  being  easily  killed  whilst  bringing  food 
to  the  sitting  females.  Macgillivray,  however,  gives,  on  the 
authority  of  an  excellent  observer,  an  instance  of  three  magpies 
successively  killed  on  the  same  nest,  which  were  all  females ;  and 
another  case  of  six  magpies  successively  killed  whilst  sitting  on 
the  same  eggs,  which  renders  it  probable  that  most  of  them  were 
females  ;  though,  as  I  hear  from  Mr.  Fox,  the  male  will  sit  on  the 
eggs  when  the  female  is  killed. 

Sir  J.  Lubbock's  gamekeeper  has  repeatedly  shot,  but  how 
often  he  could  not  say,  one  of  a  pair  of  jays  (Garrulus  glandarius), 

5  On  magpies,  Jenner,  in  '  Phil.  p.  570.  Thompson,  in  *  Annals  and 
Transact.'  1824,  p.  21.  Macgil-  Mag.  of  Nat.  Hist.'  vol.  viii.  1842, 
livray,  'Hist.   British   Birds,'  vol.  i.      p.  404. 


4o8  The  Descent  of  Maji.  Part  II. 

and  has  never  failed  shortly  afterwards  to  find  the  survivor 
re-matched.  Mr.  Fox,  Mr.  F.  Bond,  and  others  have  shot  one  of 
a  pair  of  carrion-crows  (^(.'orvns  corone),  but  the  nest  was  soon 
again  tenanted  by  a  pair.  These  bh-ds  are  rather  common;  but 
the  peregrine-falcon  (Fako  peregrinus)  is  rare, yet  Mr,  Thompson 
states  that  in  Ireland  "  if  either  an  old  male  or  female  be  killed 
"  in  the  breeding- season  (not  an  uncommon  circumstance), 
"  another  mate  is  found  within  a  very  few  days,  so  that  the 
"  eyries,  notwithstanding  such  casualties,  are  sure  to  turn  out 
"  their  complement  of  young."  Mr.  Jenner  Weir  has  known  the 
same  thing  with  the  peregrine-falcons  at  Beachy  Head.  The 
same  observer  informs  me  that  three  kestrels  ( /-a/co  tinminculus), 
all  males,  were  killed  one  after  the  other  whilst  attending  the 
same  nest ;  two  of  these  were  in  mature  plumage,  but  the  third 
was  in  the  plumage  of  the  previous  year.  Even  with  the  rare 
golden  eagle  {Aqmla  chrysaetos),  Mr.  Birkbeck  was  assured  by  a 
trustworthy  gamekeeper  in  Scotland,  that  if  one  is  killed,  another 
is  soon  found.  So  with  the  white  owl  (Strix  flammed),  "  the 
"  survivor  readily  found  a  mate,  and  the  mischief  went  on." 

White  of  Selborne,  who  gives  the  case  of  the  owl,  adds  that 
he  knew  a  man,  who  from  believing  that  partridges  w^hen  paired, 
were  disturbed  by  the  males  fighting,  used  to  shoot  them ;  and 
though  he  had  widowed  the  same  female  several  times,  she 
always  soon  found  a  fresh  partner.  This  same  naturalist  ordered 
the  spaiTows,  which  deprived  the  house-martins  of  their  nests, 
to  be  shot ;  but  the  one  w^hich  was  left,  "  be  it  cock  or  hen, 
"  presently  procured  a  mate,  and  so  for  several  times  following." 
I  could  add  analogous  cases  relating  to  the  chaffinch,  nightingale, 
and  redstart.  With  respect  to  the  latter  bu'd  {Plapnkura  rutl- 
cilla\  a  writer  expresses  much  surprise  how  the  sitting  female 
could  so  soon  have  given  effectual  notice  that  she  was  a  widow, 
for  the  species  was  not  common  in  the  neighbourhood.  Mr. 
Jenner  Weir  has  mentioned  to  me  a  nearly  similar  case;  at 
Blackheath  he  never  sees  or  hears  the  note  of  the  wild  bullfinch, 
yet  when  one  of  his  caged  males  has  died,  a  wild  one  in  the  course 
of  a  few  days  has  generally  come  and  perched  near  the  widowed 
female,  whose  call-note  is  not  loud.  I  will  give  only  one  other  fact, 
on  the  authority  of  this  same  observer ;  one  of  a  pair  of  starlings 
(^Sturnus  vulgaris)  was  shot  in  the  morning;  by  noon  a  new 
mate  was  found  ;  this  was  again  shot,  but  before  night  the  pair 
was  complete  ;  so  that  the  disconsolate  widow  or  widower  was 
thrice  consoled  during  the  same  day.  Mr.  Engleheart  also 
informs  me  that  he  used  during  several  years  to  shoot  one  of  a 
pair  of  starlings  which  built  in  a  hole  in  a  house  at  Blackheath  ; 
but    the   loss  was  always  immediately  repaired.     During  one 


(/'hap.  XI  \^  Unpaired  Birds.  409 

season  lie  kept  an  account,  and  found  that  lie  liad  sliot  tliirty-five 
birds  from  the  same  nest;  these  consisted  of  both  males  and 
females,  but  in  what  jn-oportion  he  could  not  say  :  nevertheless, 
after  all  this  destruction,  a  brood  was  reared.*^ 

These  facts  well  deserve  attention.  How  is  it  that  there  arc 
birds  enough,  ready  to  replace  immediately  a  lost  mate  of  either 
sex  ?  Magpies,  jays,  carrion-crows,  partridges,  and  some  other 
birds,  are  always  seen  during  the  spring  in  pairs,  and  never  by 
themselves ;  and  these  offer  at  first  sight  the  most  perplexing 
cases.  But  birds  of  the  same  sex,  although  of  course  not  truly 
paired,  sometimes  live  in  pairs  or  in  small  parties,  as  is  known 
to  be  the  case  with  pigeons  and  partridges.  Birds  also  sometimes 
live  in  triplets,  as  has  been  observed  with  starlings,  carrion-crows, 
parrots,  and  partridges.  With  partridges  two  females  have  been 
known  to  live  with  one  male,  and  two  males  with  one  female. 
In  all  such  cases  it  is  probable  that  the  union  would  be  easily 
broken  ;  and  one  of  the  three  would  readily  pair  with  a  widow  or 
widower.  The  males  of  certain  birds  may  occasionally  be  heard 
pouring  forth  their  love-song  long  after  the  proper  time,  shew- 
ing that  they  have  either  lost  or  never  gained  a  mate.  Death 
from  accident  or  disease  of  one  of  a  pair,  would  leave  the  other 
free  and  single ;  and  there  is  reason  to  believe  that  female  birds 
during  the  breeding-season  are  especially  liable  to  premature 
death.  Again,  birds  which  have  had  their  nests  destroyed,  or 
barren  pairs,  or  retarded  individuals,  would  easily  be  induced  to 
desert  their  mates,  and  would  probably  be  glad  to  take  what 
share  they  could  of  the  pleasures  and  duties  of  rearing  offspring 
although  not  their  own.'^  Such  contingencies  as  these  probably 
explain  most  of  the  foregoing  cases.^    Kevertheless,  it  is  a  strange 

^  On  the    peregrine     falcon    see  1824.     In  regard  to  birds  living  in 

Tiiompson,  '  Nat.   Hist,  of  Ireland,  triplets,  I  owe  to  Mr.  Jenner  Weir 

Birds,'  vol.  i.  1849,  p.  39.     On  owls,  the  cases  of  the  starlings  and  parrots, 

sparrows,  and  partridges,  see  White,  and  to  Mr.  Fox,  of  partridges ;  on 

'Nat.    Hist,    of  Selborne,'    edit,   of  carrion-crows,  see  the  '  Field,'  1868, 

1825,  vol.  i.  p.  139.     On  the  Phoeni-  p.    415.     On     various    male    birds 

cura,    see  Loudon's  '  Mag.    of  Nat.  singing  after  the  proper  period,  see 

Hist.'  vol.  vii.  1834,  p.  245.  Brehm  Rev.    L.    Jenyns,    'Observations    in 

('Thierleben,'    B.    iv.    s.    991)  also  Natural  History,' 1846,  p.  87. 
alludes  to  cases  of  birds  thrice  mated  ^  The  following    case    has     been 

during  the  same  day.  given  ('  The  Times,'  Aug.  Gth,  1868) 

'  See  White  ('Nat.  Hist,  of  Sel-  by  the   Rev.   F.  0.   Morris,   on  the 

borne,'  1825,  a'oI.  i.  p.   140)  on  the  authority  of  the  Hon.  and  Rev.  0. 

existence,    early    in    the    season,    of  W.      Forester.     "  The     gameiceeper 

small  coveys  of  male  partridges,  of  "  here  found  a  hawk's  nest  this  year, 

which  fact  I  have   heard  other  in-  "  with  five  young  ones   on   it.     He 

stances.    See  Jenner,  on  the  retarded  "  took    four   and    killed   them,   but 

state  of  the    generative    organs    in  "  left  one  with  its  wings  clipped  as 

certain    birds,  iu   '  Pliil.    Transact.'  *'  a  decoy  to  destroy  the  old  ones  by. 


4IO  The  Descent  of  Man.  Part  II. 

fact  that  within  the  same  district,  during  the  height  of  the 
breeding-season,  there  should  be  so  many  males  and  females 
always  ready  to  repair  the  loss  of  a  mated  bird.  \Yhy  do  not 
such  spare  birds  immediately  pair  together  ?  Have  we  not  some 
reason  to  suspect,  and  the  suspicion  has  occurred  to  Mr.  Jenner 
Weir,  that  as  the  courtship  of  birds  appears  to  be  in  many  cases 
prolonged  and  tedious,  so  it  occasionally  happens  that  certain 
males  and  females  do  not  succeed  during  the  proper  season,  in 
exciting  each  other's  loTe,  and  consequently  do  not  pair?  This 
suspicion  will  appear  somewhat  less  imj^robable  after  we  have 
seen  what  strong  antipathies  and  preferences  female  birds 
occasionally  evince  towards  particular  males. 

Mental  Qualities  of  Birds,  and  their  Taste  for  the  Beautifid. — 
Before  we  further  discuss  the  question  whether  the  females 
select  the  more  attractive  males  or  accept  the  first  whom  they 
may  encounter,  it  will  be  advisable  briefly  to  consider  the 
mental  powers  of  birds.  Their  reason  is  generally,  and  perhaps 
justly,  ranked,  as  low  ;  yet  some  facts  could  be  given  ^  leading  to 
an  opposite  conclusion.  Low  powers  of  reasoning,  however,  are 
compatible,  as  we  see  with  mankind,  with  strong  aifections, 
acute  perception,  and  a  taste  for  the  beautiful ;  and  it  is  with 
these  latter  qualities  that  we  are  here  concerned.  It  has  often 
been  said  that  parrots  become  so  deeply  attached  to  each  other 
that  when  one  dies  the  other  pines  for  a  long  time;  but  Mr. 
Jenner  Weir  thinks  that  with  most  birds  the  strength  of  their 
affection  has  been  much  exaggerated.  Nevertheless  when  one  of  a 
pair  in  a  state  of  nature  hsis  been  shot,  the  survivor  has  been  heard 
for  days  afterwards  uttering  a  plaintive  call ;  and  Mr.  St.  John 
gives  various  facts  proving  the  attachment  of  mated  birds.^° 


"  They  were  both  shot  next  day,  ia  1870,  p.  278.    Speaking  of  Japanese 

"  the  act  of  feeding  the  young  one,  nut-hatches  in  confinement  he  says  : 

"and   the    keeper    thought    it    was  "  Instead  of  the  more  yielding  fruit 

"  done  with.     The  next  day  he  came  "  of  the  yew,  which   is    the    usual 

"  again  and  found  two  other  chari-  "  food  of  the  nut-hatch  of  Japan,  at 

"  table  hawks,  who  had  come  with  "  one  time  I  substituted  hard  hazel- 

"  an  adopted  feeling  to  succour  the  "  nuts.     As    the     bird    was    unable 

"  orphan.     These  two  he  killed,  and  "to  crack  them,  he  placed  them  one 

"then  left  the  nest.     On  returning  "  by  one  in  his  water-glass,  evidently 

"  afterwards    he    found    two     more  "  with  the  notion  that  they  would 

"  charitable  individuals  on  the  same  ''  in  time  become  softer — an  interest- 

"  errand  of  mercy.     One  of  these  he  "  ing  proof  of  intelligence   on   the 

"killed;    the    other    he    also    shot,  "  part  of  these  birds." 
"but     could     not    find.     Ko    more  '"  *A  Tour    in    Sutherlandshire,' 

"came  on  the  like  fruitless  errand."  vol.    i.    1849,  p.   185.     Dr.    Buller 

^  I  am  indebted  to  Prof  Newton  says  ('Birds  of  New  Zealand,'  1872, 

for  the  following  passage  from  Mr.  p.  56)  that  a  male  King  Lory  was 

Adam's   '  Travels   of  a  Naturalist,'  killed ;  and  the  female  "  fretted  and 


j^Chap.  XIV.  Mental  Qualities.  41 1 

Mr.  Bennett  relates  ^^  that  in  China  after  a  drake  of  the  beautiful 
mandarin  Teal  had  been  stolen,  the  duck  remained  disconsolate, 
though  sedulously  courted  by  another  mandarin  drake,  who  dis- 
played before  her  all  his  charms.  After  an  interval  of  three 
M'eeks  the  stolen  drake  was  recovered,  and  instantly  the  pair 
recognised  each  other  with  extreme  joy.  On  the  other  hand 
starlings,  as  we  have  seen,  may  be  consoled  thrice  in  the  same 
day  for  the  loss  of  their  mates.  Pigeons  have  such  excellent 
local  memories,  that  they  have  been  known  to  return  to  their 
former  homes  after  an  interval  of  nine  months,  yet,  as  I  hear 
from  Mr,  Harrison  Weir,  if  a  pair  which  naturally  would  remain 
mated  for  life  be  separated  for  a  few  weeks  during  the  winter, 
and  afterwards  matched  with  other  birds,  the  two,  when  brought 
together  again,  rarely,  if  ever,  recognise  each  other. 

Birds  sometimes  exhibit  benevolent  feelings-,  they  will  feed 
the  deserted  young  ones  even  of  distinct  species,  but  this  perhaps 
ought  to  be  considered  as  a  mistaken  instinct.  They  will  feed, 
as  shewn  in  an  earlier  part  of  this  work,  adult  birds  of  their  own 
species  which  have  become  blind.  Mr.  Buxton  gives  a  curious 
account  of  a  parrot  which  took  care  of  a  frost-bitten  and  crippled 
bird  of  a  distinct  species,  cleansed  her  feathers,  and  defended  her 
from  the  attacks  of  the  other  parrots  which  roamed  freely  about 
his  garden.  It  is  a  still  more  curious  fact  that  these  birds 
apparently  evince  some  sympathy  for  the  pleasures  of  their 
fellows.  When  a  pair  of  cockatoos  made  a  nest  in  an  acacia 
tree,  "  it  was  ridiculous  to  see  the  extravagant  interest  taken  in 
"  the  matter  by  the  others  of  the  same  species."  These  parrots, 
also,  evinced  unbounded  curiosity,  and  clearly  had  "  the  idea  of 
"  property  and  possession,"  ^^  They  have  good  memories,  for  in 
the  Zoological  Gardens  they  have  plainly  recognised  their  former 
masters  after  an  interval  of  some  months. 

Birds  possess  acute  powers  of  observation.  Every  mated  bird, 
of  course,  recognises  its  fellow.  Audubon  states  that  a  certain 
number  of  mocking-thrushes  {Mimus  iwlyglottus)  remain  all  the 
year  round  in  Louisiana,  whilst  others  migrate  to  the  Eastern 
States ;  these  latter,  on  their  return,  are  instantly  recognised, 
and  always  attacked,  by  their  southern  brethren.  Birds  under 
confinement  distinguish  different  persons,  as  is  proved  by  the 
strong  and  permanent  antipathy  or  affection  which  they  shew, 
without  any  apparent  cause,  towards  certain    individuals.     I 


"moped,  refused  her  food,  and  died  '^  'Acclimatization    of    Parrots,' 

"of  a  broken  heart,"  by    C,    Buxton,    M.P.    'Annals    and 

"   'Wanderings     in    New    South  Mag.  of  Nat,  Hist.'  Nov,   1868,  p. 

Wales,'  vol,  ii.  18;U,  p,  62,  381, 


412  The  Descent  of  Man.  Part  11. 

have  heard  of  numerous  instances  with  jays,  partridges,  canaries, 
and  especially  bullfinches.  Mr.  Hussey  has  described  in  how- 
extraordinary  a  manner  a  tamed  partridge  recog-nised  everybody ; 
and  its  likes  and  dislikes  were  very  strong.  This  bird  seemed 
"  fond  of  gay  colours,  and  no  new  gown  or  cap  could  be  jDut  on 
"  without  catching  his  attention,"  ^^  Mr.  Hewitt  has  described 
the  habits  of  some  ducks  (recently  descended  from  wild  birds), 
which,  at  the  approach  of  a  strange  dog  or  cat,  would  rush 
headlong  into  the  water,  and  exhaust  themselves  in  their  attempts 
to  escape ;  but  they  knew  Mr.  Hewitt's  own  dogs  and  cats  so  well, 
that  they  would  lie  down  and  bask  in  the  sun  close  to  them. 
They  always  moved  away  from  a  strange  man,  and  so  they 
would  from  the  lady  who  attended  them.,  if  she  made  any  great 
change  in  her  dress.  Audubon  relates  that  he  reared  and  tamed 
a  wild  turkey  which  always  ran  away  from  any  strange  dog; 
this  bird  escaped  into  the  woods,  and  some  days  afterwards 
Audubon  saw,  as  he  thought,  a  wild  turkey,  and  made  his  dog 
chase  it ;  but  to  his  astonishment,  the  bird  did  not  run  away, 
and  the  dog,  when  he  came  up,  did  not  attack  the  bird,  for  they 
mutually  recognised  each  other  as  old  friends." 

Mr.  Jenner  Weir  is  convinced  that  birds  pay  particular  atten- 
tion to  the  colours  of  other  birds,  sometimes  out  of  jealousy,  and 
sometimes  as  a  sign  of  kinship.  Thus  he  turned  a  reed-bunting 
{Emheriza  schmiiculus),  which  had  acquired  its  black  head-dress, 
into  his  aviary,  and  the  new-comer  was  not  noticed  by  any  bird, 
except  by  a  bullfinch,  which  is  likewise  black-headed.  This 
bullfinch  was  a  very  quiet  bird,  and  had  never  before  quarrelled 
with  any  of  its  comrades,  including  another  reed-bunting,  which 
had  not  as  yet  become  black-headed  :  but  the  reed-bunting  with 
a  black  head  was  so  unmercifully  treated,  that  it  had  to  be 
removed.  Spizacyanea,  during  the  breeding-season,  is  of  a  bright 
blue  colour ;  and  though  generally  peaceable,  it  attacked  S.  ciris, 
which  has  only  the  head  blue,  and  completely  scalped  the 
unfortunate  bird.  Mr.  Weir  was  also  obliged  to  turn  out  a 
robin,  as  it  fiercely  attacked  all  the  birds  in  his  aviary  with  any 
red  in  their  plumage,  but  no  other  kinds ;  it  actually  killed  a 
red-breasted  crossbill,  and  nearly  killed  a  goldfinch.  On  the 
other  hand,  he  has  observed  that  some  birds,  when  first  intro- 
duced, fly  towards  the  species  which  resemble  them  most  in 
colour,  and  settle  by  their  sides. 

As  male  birds  display  their  fine  plumage  and  other  ornaments 

"  'The  Zoologist,'  1847-1848,  p.  p.  39.  Audubon  on  the  wild  turkey, 

1602.  'Ornith.  Biography,'  vol.    i.  p.   14. 

^*  Hewitt  on  wild  ducks,  'Journal  On  the  mocking-thrush,  ibid.  vol.  i, 

of    Horticulture,'   Jan.     13,    1863,  p.  110. 


Chap.  XIV.  Taste  for  tJie  Beautiful.  413 

with  so  much  care  before  the  females,  it  is  obviously  probable 
that  these  appreciate  the  beauty  of  their  suitors.  It  is,  however, 
difficult  to  obtain  direct  evidence  of  their  capacity  to  appreciate 
beauty.  When  birds  gaze  at  themselves  in  a  looking-glass  (of 
which  many  instances  have  been  recorded)  we  cannot  feel  sure 
that  it  is  not  from  jealousy  of  a  supposed  rival,  though  this  is  not 
the  conclusion  of  some  observers.  In  other  cases  it  is  difficult  to 
distinguish  between  mere  curiosity  and  admiration.  It  is  per- 
haps the  former  feeling  which,  as  stated  by  Lord  Lilford,'''  attracts 
the  ruflf  towards  any  bright  object,  so  that,  in  the  Ionian  Islands, 
"  it  will  dart  down  to  a  bright-coloured  handkerchief,  regardless 
"  of  repeated  shots."  The  common  lark  is  drawn  down  from  the 
sky,  and  is  caught  in  large  numbers,  by  a  small  mirror  made  to 
move  and  glitter  in  the  sun.  Is  it  admiration  or  curiosity  which 
leads  the  magpie,  raven,  and  some  other  birds  to  steal  and 
secrete  bright  objects,  such  as  silver  articles  or  jewels  ? 

]\Ir.  Gould  states  that  certain  humming-birds  decorate  the 
outsides  of  their  nests  "  with  the  utmost  taste  ;  they  instinctively 
"  fasten  thereon  iDcautiful  pieces  of  flat  lichen,  the  larger  pieces 
"  in  the  middle,  and  the  smaller  on  the  part  attached  to  the 
"  branch.  Now  and  then  a  pretty  feather  is  intertwined  or 
"  fixstened  to  the  outer  sides,  the  stem  being  always  so  placed, 
"  that  the  feather  stands  out  beyond  the  surface."  The  best 
evidence,  however,  of  a  taste  for  the  beautiful  is  afforded  by  the 
three  genera  of  Australian  bower-birds  already  mentioned.  Their 
bowers  (see  tig.  46,  p.  382),  where  the  sexes  congregate  and  play 
strange  antics,  are  variously  constructed,  but  what  most  concerns 
us  is,  that  they  are  decorated  by  the  several  species  in  a  different 
manner.  The  Satin  bower-bird  collects  gaily-coloured  articles, 
such  as  the  blue  tail-feathers  of  parrakeets,  bleached  bones  and 
shells,  which  it  sticks  between  the  twigs,  or  arranges  at  the 
entrance.  Mr.  Gould  found  in  one  bower  a  neatly- worked  stone 
tomahawk  and  a  slip  of  blue  cotton,  evidently  procured  from  a 
native  encampment.  These  objects  are  continually  re-arranged, 
and  carried  about  by  the  birds  wiiilst  at  play.  The  bower  of 
the  Spotted  bower-bird  "  is  beautifully  lined  with  tall  grasses,  so 
"  disposed  that  the  heads  nearly  meet,  and  the  decorations  are 
"  very  profuse."  Eound  stones  are  used  to  keep  the  grass-stems 
in  their  proper  places,  and  to  make  divergent  paths  leading  to 
the  bower.  The  stones  and  shells  are  often  brought  from  a 
great  distance.  The  Regent  bird,  as  described  by  Mr.  Eamsay, 
ornaments  its  short  bower  with  bleached  land-shells  belonging  to 
five  or  six  species,  and  with  "  berries  of  various  colours,  blue,  red, 
'*  and  black,  whicn  give  it  when  fresh,  a  very  pretty  appearance. 
"  The  'Ibis,'  vol.  ii.  1860,  p.  844. 


414  The  Descent  of  Ma7i.  Part  II. 

"  Besides  these  there  were  several  newly-picked  leaves  and 
"  young  shoots  of  a  pinkish  colour,  the  whole  shewing  a  de- 
"  cided  taste  for  the  beautiful."  Well  may  Mr.  Gould  say,  that 
"  these  highly  decorated  halls  of  assembly  must  be  regarded  as 
"  the  most  wonderful  instances  of  bird-architecture  yet  dis- 
"  covered ;"  and  the  taste^  as  we  see,  of  the  several  species 
certainly  differs.^^ 

Preftrence  for  jMrticulur  Males  hy  the  Females. — Having  made 
these  preliminary  remarks  on  the  discrimination  and  taste  of 
birds,  I  will  give  all  the  facts  known  to  me,  which  bear  on  the 
preference  shewn  by  the  female  for  particular  males.  It  is 
certain  that  distinct  species  of  birds  occasionally  pair  in  a  state 
of  nature  and  produce  hybrids.  Many  instances  could  be 
given:  thus  Macgillivray  relates  how  a  male  blackbird  and 
female  thrush  "fell  in  love  with  each  other,"  and  produced 
oflfepring.^^  Several  years  ago  eighteen  cases  had  been  recorded 
of  the  occurrence  in  Great  Britain  of  hybrids  between  the  black 
grouse  and  pheasant ;  ^^  but  most  of  these  cases  may  perhaps  be 
accounted  for  by  solitary  birds  not  finding  one  of  their  own 
species  to  pair  with.  With  other  birds,  as  Mr.  Jenner  Weii-  has 
reason  to  believe,  hybrids  are  sometimes  the  result  of  the  casual 
intercourse  of  birds  building  in  close  proximity.  But  these 
remarks  do  not  apply  to  the  many  recorded  instances  of  tamed 
or  domestic  birds,  belonging  to  distinct  species,  which  have 
become  absolutely  fascinated  with  each  other,  although  living 
with  their  own  species.  Thus  Waterton^"  states  that  out  of  a 
flock  of  twenty-three  Canada  geese,  a  female  paired  with  a 
solitary  Bernicle  gander,  although  so  different  in  aj^pearance  and 
size;  and  they  produced  hybrid  offspring.  A  male  widgeon 
(Mareca  ^yenelope),  living  with  females  of  the  same  species,  has 
been  known  to  pair  with  a  pintail  duck,  Querqucdula  acuta. 
Lloyd  describes  the  remarkable  attachment  between  a  shield- 
drake  {TadoriM  vulpanser)  and  a  common  duck.     Many  ad- 

'^  Oa  the  oi'namented  nests  of  For  the  following  statements,  see  on 
humming-birds,  Gould,  '  Introduc-  the  widgeon,  Loudon's  '  Mag.  of  Nat. 
tion  to  the  Trochilidse,'  1861,  p.  19.  Hist.'  vol.  ix.  p.  616;  L.  Lloyd,  'Scan- 
On  the  bower-birds,  Gould,  'Hand-  diuavian  Adventures,'  vol.  i.  1854, 
book  .to  the  Birds  of  Australia,'  p.  452.  Dixon,  'Ornamental  and 
1865,  vol,  i.  pp.  444—461.  Ramsay,  Domestic  Poultry,'  p.  137  ;  Hewitt," 
in  the  'Ibis,'  1867,  p.  456.  in   'Journal  of  Horticulture,'  Jan. 

1'  '  Hist,  of  British  Birds,' vol.  ii.  13,   1863,  p.  40;   Bechstein,  '  Stu- 

p.  92.  benvogel,'    1840,    s.     230.     Mr.    J. 

'*  'Zoologist,'       1853-1854,      p.  Jenner   Weir   has    lately   given  me 

3946.  an  analogous  case  with  ducks  of  two 

^*  Waterton,     'Essays     on     Nat.  species. 
Hist.'  2nd    series,  pp.  42  and    117. 


Chap.  XIV.  Preference  by  tJie  Female.  415 

ditional  instances  could  be  given ;  and  the  Kev.  E.  S.  Dixon 
remarks  that  "  those  who  have  kept  many  different  species  of 
"  geese  together,  well  know  what  unaccountable  attachments 
"  they  are  frequently  forming,  and  that  they  are  quite  as  likely 
"  to  pair  and  rear  young  with  individuals  of  a  race  (species) 
"  apparently  the  most  alien  to  themselves,  as  with  their  own 
"  stock." 

The  Rev.  "W.  D.  Fox  informs  me  that  he  possessed  at  the  same 
time  a  pair  of  Chinese  geese  {^Anser  cygnoides),  and  a  common 
gander  with  three  geese.  The  two  lots  kept  quite  separate, 
until  the  Chinese  gander  seduced  one  of  the  common  geese  to 
live  with  him.  Moreover,  of  the  young  birds  hatched  from  the 
eggs  of  the  common  geese,  only  four  were  pure,  the  other 
eighteen  proving  hybrids ;  so  that  the  Chinese  gander  seems  to 
have  had  prepotent  charms  over  the  common  gander.  I  will 
give  only  one  other  case ;  Mr.  Hewitt  states  that  a  wild  duck, 
reared  in  captivity,  "  after  breeding  a  couple  of  seasons  with  her 
"  own  mallard,  at  once  shook  him  off  on  my  placing  a  male 
"  Pintail  on  the  water.  It  was  evidently  a  case  of  love  at  first 
"  sight,  for  she  swam  about  the  new-comer  caressingly,  though 
"  he  appeared  evidently  alarmed  and  averse  to  her  overtures  of 
"  affection.  From  that  hour  she  forgot  her  old  partner.  Winter 
"  passed  by,  and  the  next  spring  the  Pintail  seemed  to  have 
"  become  a  convert  to  her  blandishments,  for  they  nested  and 
"  produced  seven  or  eight  young  ones." 

What  the  cliarm  may  have  been  in  these  several  eases,  beyond 
mere  novelty,  we  cannot  even  conjecture.  Colour,  however, 
sometimes  comes  into  play ;  for  in  order  to  raise  hybrids  from 
the  siskin  (FringiUa  spinas)  and  the  canary,  it  is  much  the  best 
plan,  according  to  Bcchstein,  to  place  birds  of  the  same  tint 
together.  Mr.  Jenner  Weir  turned  a  female  canary  into  his 
aviary,  where  there  were  male  linnets,  goldfinches,  siskins, 
greenfinches,  chaffinches,  and  other  birds,  in  order  to  sec  which 
she  would  choose;  but  there  never  was  any  doubt,  and  the 
greenfinch  carried  the  day.  They  paired  and  produced  hybrid 
offspring. 

The  fact  of  the  female  preferring  to  pair  with  one  male  rather 
than  with  another  of  the  same  species,  is  not  so  likely  to  excite 
attention,  as  when  this  occurs,  as  we  have  just  seen,  between 
distinct  species.  The  former  cases  can  best  be  observed  with 
domesticated  or  confined  birds ;  but  these  are  often  pampered 
by  high  feeding,  and  sometimes  have  their  instincts  vitiated  to 
an  extreme  degree.  Of  this  latter  fact  I  could  give  sufficient 
proofs  with  pigeons,  and  especially  with  fowls,  but  they  cannot 
be  here  related.     Vitiated  instincts  may  also  account  for  some 


41 6  The  Descent  of  Man. .  Part  IT. 

of  tlic  hybrid  unions  above  mentioned ;  but  in  many  of  these 
cases  the  birds  were  allowed  to  range  freely  over  large  ponds, 
and  there  is  no  reason  to  suppose  that  they  were  unnaturally 
stimulated  by  high  feeding. 

With  respect  to  birds  in  a  state  of  nature,  the  first  and  most 
obvious  supposition  which  will  occur  to  every  one  is  that  the 
female  at  the  proper  season  accepts  the  first  male  whom  she 
may  encounter ;  but  she  has  at  least  the  opportunity  for  exerting 
a  choice,  as  she  is  almost  invariably  pursued  by  many  males. 
Audubon—  and  we  must  remember  that  he  spent  a  long  life  in 
prowling  about  the  forests  of  the  United  States  and  observing 
the  birds — does  not  doubt  that  the  female  deliberately  chooses 
her  mate ;  thus,  speaking  of  a  woodpecker,  he  says  the  hen  is 
followed  by  half-a-dozen  gay  suitors,  who  continue  performing 
strange  antics,  "  until  a  marked  preference  is  shewn  for  one.'' 
The  female  of  the  red-winged  starling  {Agelxus  phceniceus)  is 
likewise  pursued  by  several  males,  "  until,  becoming  fatigued, 
"  she  aligiits,  receives  their  addresses,  and  soon  makes  a  choice," 
He  describes  also  how  several  male  night-jars  repeatedly  plunge 
ttu-ough  the  air  with  astonishing  rapidity,  suddenly  turning,  and 
thus  making  a  singular  noise ;  "  but  no  sooner  has  the  female 
"  made  her  choice,  than  the  other  males  are  driven  away." 
With  one  of  the  vultures  (Calhartes  aura)  of  the  United  States, 
parties  of  eight,  ten,  or  more  males  and  females  assemble  on 
fallen  logs,  "  exhibiting  the  strongest  desire  to  please  mutually," 
and  after  many  caresses,  each  male  leads  off  his  partner  on  the 
wing.  Audubon  likewise  carefully  observed  the  wild  flocks  of 
Canada  geese  (Anser  canadensis),  and  gives  a  graphic  description 
of  their  love-antics;  he  says  that  the  birds  which  had  been 
previously  mated  "renewed  their  courtship  as  early  as  the 
"  month  of  January,  while  the  others  would  be  contending  or 
"  coquetting  for  hours  every  day,  until  all  seemed  satisfied  with 
"  the  choice  they  had  made,  after  which,  although  they  re- 
"■  mained  together,  any  j^erson  could  easily  perceive  that  they 
"  were  careful  to  keep  in  pairs.  I  have  observed  also  that  the 
"  older  the  birds,  the  shorter  were  the  preliminaries  of  their 
"  courtship.  The  bachelors  and  old  maids,  whether  in  regret, 
"  or  not  caring  to  be  disturbed  by  the  bustle,  quietly  moved 
"  aside  and  lay  down  at  some  distance  from  the  rest."^'  Many 
similar  statements  with  respect  to  other  birds  could  be  cited 
from  this  same  observer. 

Turning  now  to  domesticated  and  confined  birds,  I  will 
commence  by  giving  what  little  I  have  learnt  respecting  the 

20  Audubon  '  Oi-nitholog.  Biography,'  vol.  i,  pp.  191,349;  vol.  ii.  pp.  42, 
275  ;  vol.  iii.  p.  2. 


Chap.  XIV.  Preference  by  the  Female.  417 

courtship  of  fowls.  I  have  received  long  letters  on  this  subject 
from  Messrs.  Hewitt  and  Tegetmeier,  and  almost  an  essay  from 
the  late  Mr.  Brent.  It  will  be  admitted  by  every  one  that  these 
gentlemen,  so  well  known  from  their  published  works,  are 
careful  and  experienced  observers.  They  do  not  believe  that 
the  females  prefer  certain  males  on  account  of  the  beauty  of 
their  plumage ;  but  some  allowance  must  be  made  for  the  arti- 
ficial state  under  which  these  birds  have  long  been  kept.  ]\Ir. 
Tegetmeier  is  convinced  that  a  game-cock,  though  disfigured  by 
being  dubbed  and  with  his  hackles  trimmed,  would  be  accepted 
as  readily  as  a  male  retaining  all  his  natural  ornaments.  Mr. 
Brent,  however,  admits  that  the  beauty  of  the  male  probably 
aids  in  exciting  the  female ;  and  her  acquiescence  is  necessary. 
i\Ir.  Hewitt  is  convinced  that  the  union  is  by  no  means  left  to 
mere  chance,  for  the  female  almost  invariably  prefers  the  most 
vigorous,  defiant,  and  mettlesome  male ;  hence  it  is  almost 
useless,  as  he  remarks,  "  to  attempt  true  breeding  if  a  game-cock 
"  in  good  health  and  condition  runs  the  locality,  for  almost 
"  every  hen  on  leaving  the  roosting-plaee  will  resort  to  the 
"  game-cock,  even  though  that  bird  may  not  actually  drive 
"  a^vny  the  male  of  her  own  variety."  Under  ordinary  circum- 
stances the  males  and  females  of  the  fowl  seem  to  come  to  a 
mutual  understanding  by  means  of  certain  gestures,  described 
to  me  by  Mr.  Brent.  But  hens  will  often  avoid  the  officious 
attentions  of  young  males.  Old  hens,  and  hens  of  a  pugnacious 
disposition,  as  the  same  writer  informs  me,  dislike  strange 
males,  and  will  not  yield  until  well  beaten  into  compliance. 
Ferguson,  however,  describes  how  a  quarrelsome  hen  was 
subdued  by  the  gentle  courtship  of  a  Shanghai  cock.-' 

There  is  reason  to  believe  that  pigeons  of  both  sexes  prefer 
pairing  with  birds  of  the  same  breed;  and  dovecot-j^igeons 
dislike  all  the  highly  improved  breeds.^^  Mr.  Harrison  Weir 
has  lately  heard  from  a  trustworthy  observer,  who  keeps  blue 
pigeons,  that  these  drive  away  all  other  coloured  varieties,  such 
as  white,  red,  and  yellow ;  and  from  another  observer,  that  a 
female  dun  carrier  could  not,  after  repeated  trials,  be  matched 
with  a  black  male,  but  immediately  paired  with  a  dun.  Again. 
Mr.  Tegetmeier  had  a  female  blue  turbit  that  obstinately 
refused  to  pair  with  two  males  of  the  same  breed,  which  were 
successively  shut  up  with  her  for  weeks ;  but  on  being  let  out 
she  would  have  immediately  accepted  the  first  blue  dragon  that 
offered.     As  she  was  a  valuable  bird,  she  was  then  shut  up  for 

'*  '  Rare  aud  Prize  Poultry,' 185-4,  Plants  under  Domestication,' vol.  ii 
p.  ^7.  p.  103. 

'^  '  The  Variation  of  Animals  and 

19 


41 8  The  Descent  of  Man.  Tart  IT. 


many  weeks  with  a  siker  (i.e.,  very  pale  blue)  male,  and  at  last 
mated  with  him.  Nevertheless,  as  a  general  rule,  colour  appears 
to  have  little  influence  on  the  pairing  of  pigeons.  Mr.  Tegct- 
meier,  at  my  request,  stained  some  of  his  birds  wdth  magenta, 
but  they  were  not  much  noticed  by  the  others. 

"Female  pigeons  occasionally  feel  a  strong  antipathy  towards 
certain  males,  without  any  assignable  cause.  Thus  MM.  Boitard 
and  Corbie,  whose  experience  extended  over  forty-five  years, 
state :  "  Quand  une  femelle  eprouve  de  Fantipathie  pour  un 
"  male  avec  lequel  on  veut  I'accoupler,  malgre  tons  les  feux 
"  de  I'amour,  malgre  Palpiste  et  le  chenevis  dont  on  la  nourrit 
"  pour  augmenter  son  ardeur,  malgre  un  emprisonnement  de 
"  six  mois  et  meme  d'un  an,  elle  refuse  constamment  ses 
"caresses;  les  avances  empressees,  les  agaceries,  les  tournoie- 
"  mens,  les  tendres  roucoulemens,  rien  ne  pent  lui  plaire  ni 
*'  I'emouvoir ;  gonflee,  boudeuse,  blottie  dans  un  coin  de  sa 
"  prison,  elle  n'en  sort  que  pour  boire  et  manger,  ou  pour  re- 
"  pousser  avec  une  espece  de  rage  des  caresses  de  venues  trop  pres- 
'•'  santes."^^  On  the  other  hand,  Mr.  Harrison  Weir  has  himself 
observed,  and  has  heard  from  several  breeders,  that  a  female 
pigeon  will  occasionally  take  a  strong  fancy  for  a  particular  male, 
and  will  desert  her  own  mate  for  him.  Some  females,  accord- 
ing to  another  experienced  observer,  Eiedel,^*  are  of  a  profligate 
disposition,  and  prefer  almost  any  stranger  to  their  own  mate. 
Some  amorous  males,  called  by  our  English  fanciers  "  gay 
birds,"  are  so  successful  in  their  gallantries,  that,  as  Mr.  H. 
"Weir  informs  me,  they  must  be  shut  up  on  account  of  the 
mischief  which  they  cause. 

Wild  turkeys  in  the  United  States,  according  to  Audubon, 
"  sometimes  pay  their  addresses  to  the  domesticated  females, 
"'and  are  generally  received  by  them  with  great  pleasure."  So  that 
these  females  apparently  prefer  the  wild  to  their  own  males  ^ 

Here  is  a  more  curious  case.  Sir  E.  Heron  during  many  years 
kept  an  account  of  the  habits  of  the  peafowl,  which  he  bred  in 
large  numbers.  He  states  that  "  the  hens  have  frequently  great 
"  preference  to  a  particular  peacock.  They  were  all  so  fond 
"  of  an  old  pied  cock,  that  one  year,  when  he  was  confined 
"  though  still  in  view,  they  were  constantly  assembled  close  to 
"  the  trellice-walls  of  his  prison,  and  would  not  suffer  a  japanned 

23  Boitard      and      Corbie,     'Les  ^4  <  dj^ -paubenzucht,' 1824,  s.  80. 

Pigeons,  &G.,' 1824,  p.  12.     Prosper  ^5  <  Ornithological        Biography,' 

Lucas    ('Traite     de    I'Hered.     Nat.'  vol.  i.  p.  13.    See  to  the  same  eflect, 

toin.  ii.   1850,  p.  296)  has  himself  Dr.  Bryant,  in  '  Allen's  Mammals  arj 

observed   nearly  similar  facts  with  Birds  of  Florida,'  p.  344. 
pigeons. 


Chap.  XIV,  Preference  by  the  Female.  419 

"  peacock  to  touch  them.  On  his  being  let  out  in  the  autumn, 
*'  the  oldest  of  the  hens  instantly  courted  him,  and  was  success- 
"  ful  in  her  courtship.  The  next  year  he  was  shut  up  in  a  stable, 
"  and  then  the  hens  all  courted  his  rival."  ^*^  This  rival  was  a 
japanned  or  black-winged  peacock,  to  our  eyes  a  more  beautilul 
bird  than  the  common  kind. 

Lichtenstein,  who  was  a  good  observer  and  had  excellent 
opportunities  of  observation  at  the  Cape  of  Good  Hope,  assured 
Hudolphi  that  the  female  widow-bird  {CJura  'pro(/nc)  disowns  the 
male,  when  robbed  of  the  long  tail-feathers  with  whicli  he  is 
ornamented  during  the  breeding-season.  I  presume  that  this 
observation  must  have  been  made  on  birds  under  confinement.^' 
Here  is  an  analogous  case ;  Dr.  Jaeger,^^  director  of  the  Zoological 
Gardens,  of  Vienna,  states  that  a  male  silver-pheasant,  who  had 
been  triumphant  over  all  other  males  and  was  the  accepted 
lover  of  the  females,  had  his  ornamental  plumage  spoiled.  He 
was  then  immediately  superseded  by  a  rival,  who  got  the  upper 
hand  and  afterwards  led  the  flock. 

It  is  a  remarkable  fact,  as  shewing  how  important  colour  is  in 
the  courtship  of  birds,  that  Mr.  Boardman,  a  well-known  col- 
lector and  observer  of  birds  for  many  years  in  the  Northern 
United  States,  has  never  in  his  large  experience  seen  an  albino 
paired  with  another  bird;  yet  he  has  had  opportunities  of 
observing  many  albinos  belonging  to  several  species.^^  It  can 
hardly  be  maintained  that  albinos  in  a  state  of  nature  are 
incapable  of  breeding,  as  they  can  be  raised  with  the  greatest 
facility  under  confinement.  It  appears,  therefore,  that  we  must 
attribute  the  fact  that  they  do  not  pair,  to  their  rejection  by 
their  normally  coloured  comrades. 

Female  birds  not  only  exert  a  choice,  but  in  some  few  cases 
they  court  the  male,  or  even  fight  together  for  his  possession. 
Sir  E.  Heron  states  that  with  peafowl,  the  first  advances  are 
always  made  by  the  female ;  something  of  the  same  kind  takes 
place,  according  to  Audubon,  with  the  older  females  of  the  wild 
turkey.  With  the  capercailzie,  the  females  flit  round  the  male 
whilst  he  is  parading  at  one  of  the  places  of  assemblage,  and 
solicit  his  attention.^"    We  have  seen  that  a  tame  wild-duck 

2«  '  Proc.  Zool.  Soc'  1835,  p.  54.  ihre    Stellung    zu     Moral    und    Re- 

The  japanned  peacock  is  considered  ligiou,'  1869,  s.  59. 
by  Mr.  Sclater  as  a  distinct  species,  '*  This  statement  is  given  by  Mr 

and    has    been    named    Fato    nigri-  A.  Leith  Adams,  in  his  '  Field  and 

pennis;   but  the  evidence  seems    to  Forest    Rambles,'    1873,  p.    76,  and 

me  to  shew  that  it  is  only  a  variety.  accords  with  his  own  experience. 

"  Rudolphi,  'Beytriige    zur  An-  ^^  In   regard  to  peafowl,  see  Sir 

thropologie,'  1812,  s.  184-.  R.  Heron,  'Proc.  Zoolog.  Soc'  J 835, 

^^  '  Die  Darwin'sche  Thoorie,  und  p.  51-,  and   the    Rev.    E.   S.   Dixon, 


420  The  Descent  of  Man.  Part  II. 

seduced  an  unwilling  pintail  drake  after  a  long  courtship,  Mr. 
Bartlett  believes  that  the  Lophophorus,  like  many  other  gallina- 
ceous birds,  is  naturally  polygamous,  but  two  females  cannot  be 
X)laced  in  the  same  cage  with  a  male,  as  they  fight  so  much 
together.  The  following  instance  of  rivalry  is  more  surprising 
as  it  relates  to  bullfinches,  which  usually  pair  for  life.  Mr. 
Jenner  Weir  introduced  a  dull-coloured  and  ugly  female  into  his 
aviary,  and  she  immediately  attacked  another  mated  female  so 
unmercifully,  that  the  latter  had  to  be  separated.  The  new 
female  did  all  the  courtship,  and  was  at  last  successful,  for  she 
paired  with  the  male ;  but  after  a  time  she  met  with  a  just  re- 
tribution, for,  ceasing  to  be  pugnacious,  she  was  replaced  by  the 
old  female,  and  the  male  then  deserted  his  new  and  returned  to 
his  old  love. 

In  all  ordinary  cases  the  male  is  so  eager  that  he  will  accept 
any  female,  and  does  not,  as  far  as  we  can  judge,  prefer  one  to 
the  other  ;  but,  as  we  shall  hereafter  see,  exceptions  to  this  rule 
apparently  occur  in  some  few  groups.  With  domesticated  birds, 
I  have  heard  of  only  one  case  of  males  shewing  any  preference 
for  certain  females,  namely,  that  of  the  domestic  cock,  who, 
according  to  the  high  authority  of  Mr.  Hewitt,  prefers  the 
younger  to  the  older  hens.  On  the  other  hand,  in  effecting 
hybrid  unions  between  the  male  jtheasant  and  common  hens, 
Mr.  Hewitt  is  convinced  that  the  pheasant  invariably  prefers  the 
older  birds.  He  does  not  appear  to  be  in  the  least  influenced  by 
their  colour,  but  "  is  most  capricious  in  his  attachments :"  ^^ 
from  some  inexplicable  cause  he  shews  the  most  determined 
aversion  to  certain  hens,  which  no  care  on  the  part  of  the 
breeder  can  overcome.  Mr.  Hewitt  Informs  me  that  some  hens 
are  quite  unattractive  even  to  the  males  of  their  own  species,  so 
that  they  may  be  kept  with  several  cocks  during  a  whole  season, 
and  not  one  egg  out  of  forty  or  fifty  will  prove  fertile.  On  the 
other  hand,  with  the  Long-tailed  duck  {Harelda  glacialis),  "  it 
has  been  remarked,"  says  M.  Ekstrom,  "  that  certain  females 
*'  are  much  more  courted  than  the  rest.  Frequently,  indeed, 
"  one  sees  an  individual  surrounded  by  six  or  eight  amorous 
"  males."  Whether  this  statement  is  credible,  I  know  not ;  but 
the  native  sportsmen  shoot  these  females  in  order  to  stuff  them 
as  decoys.^ 

With  respect  to  female  birds  feeling  a  preference  for  particular 


'Ornamental   Poultry,'   18-t8,  p.  8.  "  Mr.  Hewitt,  quoted  in  '  Teget- 

For  the  turkey,  Audubon,  ibid.  p.  4,  meier's  Poultry  Book,'  1866,  p.  165. 

For  the  capercailzie,  Lloyd,  '  Game  ^2  Quoted  in  Lloyd's '  Game  Birds 

birds  of  Sweden,'  1867,  p.  23.  of  Sweden,'  p.  345. 


Chap.  XIV.  Preference  by  the  Female.     ■  42 1 

males,  we  must  bear  in  mind  that  we  can  judge  of  choice  being 
exerted,  only  by  analogy.  If  an  inhabitant  of  another  planet 
were  to  behold  a  number  of  young  rustics  at  a  fair  courting  a 
pretty  girl,  and  quarrelling  about  her  like  birds  at  one  of  their 
places  of  assemblage,  he  would,  by  the  eagerness  of  ihe  wooers 
to  please  her  and  to  display  their  finery,  infer  that  she  liad  the 
power  of  choice.  Now  with  birds,  the  evidence  stands  thus; 
they  have  acute  powers  of  observation,  and  they  seem  to  have 
some  taste  for  the  beautiful  both  in  colour  and  sound.  It  is 
certain  that  the  females  occasionally  exhibit,  from  unknown 
causes,  the  strongest  antipathies  and  ])references  for  particular 
males.  When  the  sexes  differ  in  colour  or  in  other  ornaments 
the  males  with  rare  exceptions  are  the  more  decorated,  either 
permanently  or  temporarily  during  the  breeding-season.  They 
sedulously  display  their  various  ornaments,  exe;.'t  their  voices, 
and  perform  strange  antics  in  the  presence  of  the  females.  Even 
well-armed  males,  who,  it  might  be  thought,  would  altogether 
depend  for  success  on  the  law  of  battle,  are  in  most  cases  highly 
ornamented;  and  their  ornaments  have  been  acquired  at  the 
expense  of  some  loss  of  power.  In  other  cases  ornaments  have 
been  acquired,  at  the  cost  of  increased  risk  from  birds  and 
beasts  of  prey.  With  various  species  many  individuals  of  both 
sexes  congregate  at  the  same  spot,  and  their  courtship  is  a  pro- 
longed affair.  There  is  even  reason  to  suspect  that  the  males 
and  females  within  the  same  district  do  not  always  succeed  in 
pleasing  each  other  and  pairing. 

What  then  are  we  to  conclude  from  these  facts  and  considera- 
tions ?  Does  the  male  parade  his  charms  with  so  much  pomp 
and  rivalry  for  no  purpose  ?  Are  we  not  justified  in  believing 
that  the  female  exerts  a  choice,  and  that  she  receives  the 
addresses  of  the  male  who  pleases  her  most  ?  It  is  not  probable 
that  she  consciously  deliberates ;  but  she  is  most  excited  or 
attracted  by  the  most  beautiful,  or  melodious,  or  gallant  males. 
Nor  need  it  be  supposed  that  the  female  studies  each  stripe  or  spot 
of  colour  ;  that  the  peahen,  for  instance,  admires  each  detail  in  the 
gorgeous  train  of  the  peacock— she  is  probably  struck  only  by 
the  general  effect.  Nevertheless,  after  hearing  how  carefully  the 
male  Argus  pheasant  displays  his  elegant  primary  wing-feathers, 
and  erects  his  ocellated  plumes  in  the  right  position  for  their 
full  effect ;  or  again,  how  the  male  goldfinch  alternately  displays 
his  gold-bespangled  wings,  we  ought  not  to  feel  too  sure  that  the 
female  does  not  attend  to  each  detail  of  beauty.  We  can  judge, 
as  already  remarked,  of  choice  being  exerted,  only  from  analogy ; 
and  the  mental  powers  of  birds  do  not  differ  fundamentally  from 
ours.     From  these  various  considerations  we  may  conclude  that 


422  The  Descent  of  Man.  Part  II. 

the  pairing  of  birds  is  not  left  to  chance ;  but  that  those  males, 
which  are  best  able  by  their  various  charms  to  please  or  excite 
the  female,  are  under  ordinary  circumstances  accepted.  If  this 
be  admitted,  there  is  not  much  difficulty  in  understanding  how- 
male  birds  have  gradually  acquired  their  ornamental  characters. 
All  animals  present  individual  differences,  and  as  man  can 
modify  his  domesticated  birds  by  selecting  the  individuals 
which  appear  to  him  the  most  beautiful,  so  the  habitual  or  even 
occasional  preference  by  the  female  of  the  more  attractive  males 
would  almost  certainly  lead  to  their  modification ;  and  such 
modifications  might  in  the  course  of  time  be  augmented  to 
almost  any  extent,  compatible  with  the  existence  of  the  species. 
Variability  of  Birds,  and  especially  of  their  Secondary  Sexual 
Characters. — Variability  and  inheritance  are  the  foundations  for 
the  work  of  selection.  That  domesticated  birds  have  varied 
greatly,  their  variations  being  inherited,  is  certain.  That  birds 
in  a  state  of  nature  have  been  modified  into  distinct  races  is 
now  universally  admitted.-^^  Variations  may  be  divided  into 
two  classes ;  those  which  appear  to  our  ignorance  to  arise  spon- 
taneously, and  those  which  are  directly  related  to  the  surrounding 
conditions,  so  that  all  or  nearly  all  the  individuals  of  the  same 
species  are  similarly  modified.  Cases  of  the  latter  kind  have 
recently  been  observed  with  care  by  Mr.  J.  A.  Allen,^*  who  shews 

'^  According  to  Dr.  Blasius  ('Ibis,'  difficult  to  account  for  the  dull  or 

vol.  ii.  1860,  p.  297),  there  are  425  dark  tints  of  almost  all  the  species 

indubitable  species  of  birds  which  inhabiting     certain     countries,     for 

breed  in  Europe,  besides  sixty  forms,  instance,  the  Galapagos  Islands  under 

which   are   frequently  regarded    as  the    equator,    the    wide    temperate 

distinct     species.     Of    the     latter,  phiins  of  Patagonia,  and,  as    it  ap- 

Blasius    thinks    that    only    ten    are  pears,  Egypt    (see    Mr.  Hartshorne 

really  doubtful,  and  that  the  other  in  the  'American  Naturalist,'  1873, 

fifty  ought  to  be  united  with  their  p.  747).     These  countries  are  open, 

nearest  allies ;  but  this  shews  that  and  afford  little    shelter  to    birds; 

there  must  be  a  considerable  amount  but  it  seems  doubtful  whether  the 

of  variation  with  some  of  our  Euro-  absence  of  brightly  coloured  species 

pean  birds.     It  is  also  an  unsettled  can  be  explained  on  the  principle  of 

point     with     naturalists,     whether  protection,  for  on  the  Pampas,  which 

several  North  American  birds  ought  are    equally  open,   though    covered 

to  be  ranked  as  specifically  distinct  by  green  grass,  and  where  the  birds 

from    the    corresponding    European  would  be  equally  exposed  to  danger, 

species.    So  again  many  North  Ame-  many    brilliant    and    conspicuously 

rican  forms  which  until  lately  were  coloured     species    are    common.     I 

named  as  distinct  species,  are  now  have  sometimes  speculated  whether 

considered  to  be  local  races.  the   prevailing    dull    tints    of    the 

^*  '  Mammals  and   Birds   of   East  scenery  in  the  above  named  countries 

Florida,'  also  an  '  Ornithological  Re-  may  not  have  affected  the  apprecia- 

connaissance  of   Kansas,    &c.     Not-  tion  of  bright  colours  by  the  birds 

withstanding    the    influence  of   cli-  inhabiting  them, 
mate  on  the  colours  of  birds,  it  is 


Chap.  XIV.  Variability,  423 


that  in  the  United  States  many  species  of  birds  gradually  become 
more  strongly  coloured  in  proceeding  southward,  and  more 
lightly  coloured  in  proceeding  westward  to  the  arid  plains  of  the 
interior.  Both  sexes  seem  generally  to  be  affected  in  a  like 
manner,  but  sometimes  one  sex  more  than  the  other.  This 
result  is  not  incompatible  with  the  belief  that  the  colours  of 
birds  are  mainly  due  to  the  accumulation  of  successive  yaria- 
tious  through  sexual  selection;  for  even  after  the  sexes  have 
been  greatly  differentiated,  climate  might  produce  an  equal 
effect  on  both  sexes,  or  a  greater  effect  on  one  sex  than  on  the 
other,  owing  to  some  constitutional  difference. 

Individual  differences  between  the  members  of  the  same 
species  are  admitted  by  every  one  to  occur  under  a  state  of 
nature.  Sudden  and  strongly  marked  variations  are  rare  ;  it  is 
also  doubtful  whether  if  beneficial  they  would  often  be  preserved 
through  selection  and  transmitted  to  succeeding  generations.^^ 
Nevertheless,  it  may  be  worth  while  to  give  the  few  cases  which 
I  have  been  able  to  collect,  relating  chiefly  to  colour, — simple 
albinism  and  melanism  being  excluded.  Mr.  Gould  is  well 
known  to  admit  the  existence  of  few  varieties,  for  he  esteems 
very  slight  differences  as  specific;  yet  he  states ^"^  that  near 
Bogota  certain  humming-birds  belonging  to  the  genus  Cynanthus 
are  divided  into  two  or  three  races  or  varieties,  which  differ 
from  each  other  in  the  colouring  of  the  tail  — *'  some  having  the 
"  whole  of  the  feathers  blue,  while  others  have  the  eight  central 
"  ones  tipped  with  beautiful  green."  It  does  not  appear  that 
intermediate  gradations  have  been  observed  in  this  or  the 
following  cases.  In  the  males  alone  of  one  of  the  Australian 
parrakeets  "  the  thighs  in  some  are  scarlet,  in  others  grass- 
"  green."  In  another  parrakeet  of  the  same  country  "  some 
"  individuals  have  the  band  across  the  wing-coverts  bright- 

35  '  Ori2;in  of  Species,'  fifth  edit.  valued  individuals  of  each  breed, 
1869,  p.  104.  I  had  always  per-  without  any  intention  on  his  part 
ccived,  that  rare  and  strongly-  to  modify  the  characters  of  the 
marked  deviations  of  structure,  de-  breed.  But  until  I  read  an  able 
serving  to  be  called  monstrosities,  article  in  the  '  North  British  Re- 
could  seldom  be  preserved  through  view  '  (March  1867,  p.  289,  et  seq.), 
natural  selection,  and  that  the  pre-  which  has  been  of  more  use  to  me 
servation  of  even  highly-beneficial  than  any  other  Review,  I  did  not 
variations  would  depend  to  a  certain  see  how  great  the  chances  were 
extent  on  chance.  I  had  also  fully  against  the  preservation  of  varia- 
appreciated  the  importance  of  mere  tions,  whether  slight  or  strongly 
individual  differences,  and  this  led  pronounced,  occurring  only  in  single 
me  to  insist  so  strongly  on  the  im-  individuals. 

portance  of  that  unconscious  form  ^^  '  lutroduct.  to  the  Trochilida,' 

of  selection  by  man,  which  follows  p.  102. 
from  the    preservation  of  the  most 


424  The  Descent  of  Man.  Part  II. 

"  yellow,  while  in  others  the  same  part  is  tinged  with  red."^^ 
In  the  United  States  some  few  of  the  males  of  the  Scarlet  Tanager 
{Tanngra  ri<6ra)  have  "a  beautiful  transverse  band  of  glowing 
"  red  on  the  smaller  wing-coverts ; ''  ^^  but  this  variation  seems 
to  be  somewhat  rare,  so  that  its  preservation  through  sexual 
selection  would  follow  only  under  unusually  favourable  circum- 
stances. In  Bengal  the  Honey  buzzard  (Pernis  cristata)  has 
either  a  small  rudimental  ciest  on  its  head,  or  none  at  all :  so 
slight  a  difference,  however,  would  not  have  been  worth  notice, 
had  not  this  same  species  possessed  in  Southern  India  "  a  well- 
"  marked  occipital  crest  formed  of  several  graduated  feathers."  ^^ 

The  following  case  is  in  some  respects  more  interesting.  A 
pied  variety  of  the  raven,  with  the  head,  breast,  abdomen,  and 
parts  of  the  wings  and  tail-feathers  white,  is  confined  to  the 
Feroe  Islands.  It  is  not  very  rare  there,  for  Graba  saw  during 
his  visit  from  eight  to  ten  living  specimens.  Although  the 
characters  of  this  variety  are  not  quite  constant,  yet  it  has  been 
named  by  several  distinguished  ornithologists  as  a  distinct 
species.  The  fact  of  the  pied  birds  being  pursued  and  persecuted 
with  much  clamour  by  the  other  ravens  of  the  island  was  the 
chief  cause  which  led  Briinnich  to  conclude  that  they  were 
specifically  distinct ;  but  this  is  now  known  to  be  an  error.^" 
This  case  seems  analogous  to  that  lately  given  of  albino  birds 
not  pairing  from  being  rejected  by  their  comrades. 

In  various  parts  of  the  northern  seas  a  remarkable  variety  of 
the  common  Guillemot  (Uria  troile)  is  found;  and  in  Feroe,  one 
out  of  every  five  birds,  according  to  Graba's  estimation,  presents 
this  variation.  It  is  characterised  *^  by  a  pure  white  ring  round 
the  eye,  witli  a  curved  narrow  white  line,  an  inch  and  a  half  in 
length,  extending  back  from  the  ring.  This  conspicuous  cha- 
racter has  caused  the  bird  to  be  ranked  by  several  ornithologists 
as  a  distinct  species  under  the  name  of  U.  lacrymans,  but  it  is 
now  known  to  be  merely  a  variety.  It  often  pairs  with  the 
common  kind,  yet  intermediate  gradations  have  never  been  seen ; 
nor  is  this  surprising,  for  variations  which  appear  suddenly,  are 
often,  as  I  have  elsewhere  shewn,*^  transmitted  either  unaltered 
or  not  at  all.  We  thus  see  that  two  distinct  forms  of  the  same 
species  may  co-exist  in  the  same  district,  and  we  cannot  doubt 

"  Gould,  '  Handbook  to  Birds  of  Faro,'  1830,  s.  51-54.  Macgillivray, 

Australia,'  vol.  ii.  pp.  32  and  68.  '  Hist.    British    Birds,'   vol.   iii.    p. 

38  Audubon,     '  Ornitholog.     Bio-  745.     '  Ibis,'  vol.  v.  1863,  p.  469. 
graphy,'  1838,  vol.  iv.  p.  389.  *^  Graba,    ibid.    s.    54.      Macgil- 

3**  Jerdon,  '  Birds  of  India,'  vol.  1.  livray,  ibid.  vol.  v.  p.  327. 
p.    108 ;  and   Mr.  Blyth,  in  '  Land  ^^  <  Variation     of    Animals     and 

and  Water,'  1868,  p.  381.  Plants  under  Domestication,'  vol.  ii. 

*^  Graba,  '  Tagebuch,  Reise  nach  p.  92. 


Chap.  XIV.  Variability.  425 

that  if  the  one  had  possessed  any  advantage  over  the  other,  it 
would  soon  have  been  multiplied  to  the  exclusion  of  the  latter. 
]f,  for  instance,  the  male  pied  ravens,  instead  of  being  persecuted 
by  their  comrades,  had  been  highly  attractive  (like  the  above 
pied  peacock)  to  the  black  female  ravens,  their  numbers  would 
have  rapidly  increased.  And  this  would  have  been  a  case  of 
sexual  selection. 

With  respect  to  the  slight  individual  differences  which  are 
common,  in  a  greater  or  less  degree,  to  all  the  members  of  the 
same  species,  we  have  every  reason  to  believe  that  they  are  by 
far  the  most  important  for  the  work  of  selection.  Secondary 
sexual  characters  are  eminently  liable  to  vary,  both  with  animals 
in  a  state  of  nature  and  under  domestication."*^  There  is  also 
reason  to  believe,  as  we  have  seen  in  our  eighth  chapter,  that 
variations  are  more  apt  to  occur  in  the  male  than  in  the  female 
sex.  All  these  contingencies  are  highly  favourable  for  sexual 
selection.  Whether  characters  thus  acquired  are  transmitted 
to  one  sex  or  to  both  sexes,  depends,  as  we  shall  see  in  the 
following  chapter,  on  the  form  of  inheritance  which  prevails. 

It  is  sometimes  diificult  to  form  an  opinion  whether  certain 
slight  differences  between  the  sexes  of  birds  are  simply  the 
result  of  variability  with  sexually-limited  inheritance,  without 
the  aid  of  sexual  selection,  or  whether  they  have  been  augmented 
through  this  latter  process.  I  do  not  here  refer  to  the  many 
instances  where  the  male  displays  splendid  colours  or  other 
ornaments,  of  which  the  female  pai'takes  to  a  slight  degree  ;  for 
these  are  almost  certainly  due  to  characters  primarily  acquired  by 
the  male  having  been  more  or  less  transferred  to  the  female.  But 
what  are  we  to  conclude  with  respect  to  certain  birds  in  which, 
for  instance,  the  eyes  differ  slightly  in  colour  in  the  two  sexes  V^ 
In  some  cases  the  eyes  differ  conspicuously;  thus  with  the 
storks  of  the  genus  Xenorhynchus,  those  of  the  male  are  blackish- 
hazel,  whilst  those  of  the  females  are  gamboge-yellow;  with 
many  hornbills  (Buceros),  as  I  hear  from  Mr.  Blyth/^  the  males 
have  intense  crimson  eyes,  and  those  of  the  females  are  white. 
In  the  Buceros  hicornis,  the  hind  margin  of  the  casque  and  a 
stripe  on  the  crest  of  the  beak  are  black  in  the  male,  but  not  so 
in  the  female.  Are  we  to  suppose  that  these  black  marks  and 
the  crimson  colour  of  the  eyes  have  been  preserved  or  augmented 
through  sexual  selection  in  the  males  ?     This  is  very  doubtful ; 

*^  On  these  points  see  also  '  Varia-  of  a  Podica  and  Gallicrex  in  '  Ibis,' 

tion  of  Animals  and  Plants   under  vol.   ii.    1860,  p.    206 ;  and  vol.  v. 

Domestication,'  vol.  i.  p.  253 ;  vol.  ii.  1863,  p.  426. 
pp.  73,  75.  *^  See    also    Jerdon,     '  Birds    of 

**  See,  for  instance,  on  the  irides  India,'  vol.  i.  pp.  243-245. 


426  The  Descent  of  Man.  Taut  II. 


for  Mr.  Bartlett  shewed  me  in  the  Zoological  Gardens  that  the 
inside  of  the  mouth  of  this  Buceros  is  black  in  the  male  and 
flesh-coloured  in  the  female;  and  their  external  appearance  or 
beauty  would  not  be  thus  affected.  I  observed  in  Chili  ^^  that 
the  iris  in  the  condor,  when  about  a  year  old,  is  dark-brown,  but 
changes  at  maturity  into  yellowish-brown  in  the  male,  and  into 
bright  red  in  the  female.  The  male  has  also  a  small,  longitu- 
dinal, leaden-coloured,  fleshy  crest  or  comb.  The  comb  of  many 
gallinaceous  birds  is  highly  ornamental,  and  assumes  rivid 
colours  during  the  act  of  courtship ;  but  what  are  we  to  think 
of  the  dull-coloured  comb  of  the  condor,  which  does  not  appear 
to  us  in  the  least  ornamental  ?  The  same  question  may  be  asked 
in  regard  to  various  other  characters,  such  as  the  knob  on  the 
base  of  the  beak  of  the  Chinese  goose  (^An^er  cygnoides),  which  is 
much  larger  in  the  male  than  in  the  female.  No  certain  answer 
can  be  given  to  these  questions ;  but  we  ought  to  be  cautious  in 
assuming  that  knobs  and  various  fleshy  appendages  cannot  be 
attractive  to  the  female,  when  we  remember  that  with  savage 
races  of  man  various  hideous  deformities — deep  scars  on  the 
face  with  the  flesh  raised  into  protuberances,  the  septum  of 
the  nose  pierced  by  sticks  or  bones,  holes  in  the  ears  and  lips 
stretched  widely  open— are  all  admired  as  ornamental. 

Whether  or  not  unimportant  differences  between  the  sexes, 
such  as  those  just  specified,  have  been  preserved  through  sexual 
selection,  these  difierences,  as  well  as  all  others,  must  primarily 
depend  on  the  laws  of  variation.  On  the  principle  of  correlated 
development,  the  plumage  often  varies  on  different  parts  of  the 
body,  or  over  the  whole  body,  in  the  same  manner.  AVe  see  this 
well  illustrated  in  certain  breeds  of  the  fowl.  In  all  the  breeds 
the  feathers  on  the  neck  and  loins  of  the  males  are  elongated, 
and  are  called  hackles ;  now  when  both  sexes  acquire  a  top-knot, 
which  is  a  new  character  in  the  genus,  the  feathers  on  the  head 
of  the  male  become  hackle-shaped,  evidently  on  the  principle  of 
correlation ;  whilst  those  on  the  head  of  the  female  are  of  the 
ordinary  shape.  The  colour  also  of  the  hackles  forming  the 
top-knot  of  the  male,  is  often  correlated  with  that  of  the  hackles 
on  the  neck  and  loins,  as  may  be  seen  by  comparing  these 
feathers  in  the  Golden  and  Silver-spangled  Polish,  the  Houdans, 
and  Creve-coeur  breeds.  In  some  natural  species  we  may 
observe  exactly  the  same  correlation  in  the  colours  of  these  same 
feathers,  as  in  the  males  of  the  splendid  Gold  and  Amherst 
pheasants. 

The  structure  of  each  individual  feather  generally  causes  any 

<«  'Zoology  of  the  Voyage  of  H.M.S.  Beagle,'  1841,  p.  6. 


Chap.  XIV.  •  Ocelli.  427 


change  in  its  colouring  to  be  symmetrical ;  we  see  tliis  in  the 
various  laced,  spangled,  and  pencilled  breeds  of  the  fowl ;  and 
on  the  principle  of  correlation  the  feathers  over  the  whole  body 
are  often  coloured  in  the  same  manner.  We  are  thus  enabled 
without  much  trouble  to  rear  breeds  with  their  plumage  marked 
almost  as  symmetrically  as  in  natural  species.  In  laced  and 
spangled  fowls  the  coloured  margins  of  the  feathers  are  abruptly 
defined ;  but  in  a  mongrel  raised  by  me  from  a  black  Spanish 
cock  glossed  with  green,  and  a  white  game-hen,  all  the  feathers 
were  greenish-black,  excepting  towards  their  extremities,  which 
were  yellowish- white;  but  between  the  white  extremities  and 
the  black  bases,  there  was  on  each  feather  a  symmetrical,  curved 
zone  of  dark-brown.  In  some  instances  the  shaft  of  the  feather 
determines  the  distribution  of  the  tints;  thus  with  the  body- 
feathers  of  a  mongrel  from  the  same  black  Spanish  cock  and  a 
silver-spangled  Polish  hen,  the  shaft,  together  with  a  narrow 
space  on  each  side,  was  greenish-black,  and  this  was  surrounded 
by  a  regular  zone  of  dark-brown,  edged  with  brownish-white. 
In  these  cases  we  have  feathers  symmetrically  shaded,  like  those 
which  give  so  much  elegance  to  the  plumage  of  many  natural 
species.  I  have  also  noticed  a  variety  of  the  common  pigeon 
with  the  wing-bars  symmetrically  zoned  with  three  bright 
shades,  instead  of  being  simply  black  on  a  slaty-blue  ground,  as 
in  the  parent- species. 

In  many  groups  of  birds  the  plumage  is  differently  coloured 
in  the  several  si^ecies,  yet  certain  spots,  marks,  or  stripes  are 
retained  by  all.  Analogous  cases  occur  with  the  breeds  of  the 
pigeon,  which  usually  retain  the  two  wing-bars,  though  they 
may  be  coloured  red,  yellow,  white,  black,  or  blue,  the  rest  of  the 
plumage  being  of  some  wholly  different  tint.  Here  is  a  more 
curious  case,  in  which  certain  marks  are  retained,  though 
coloured  in  a  manner  almost  exactly  the  opposite  of  what  is 
natural ;  the  aboriginal  pigeon  has  a  blue  tail,  with  the  terminal 
halves  of  the  outer  webs  of  the  two  outer  tail  feathers  white ;  now 
there  is  a  sub-variety  having  a  white  instead  of  a  blue  tail,  with 
precisely  that  part  black  which  is  white  in  the  parent -species.'''^ 

Formation  and  Variability  of  the  Ocelli  or  eye-like  Spots  oti  the 
Plumage  of  Birds. — As  no  ornaments  are  more  beautiful  than  the 
ocelli  on  the  feathers  of  various  birds,  on  the  hairy  coats  of  some 
mammals,  on  the  scales  of  reptiles  and  fishes,  on  the  skin  of 
amphibians,  on  the  wings  of  many  Lepidoptera  and  other  insects, 
they  deserve  to  be  especially  noticed.  An  ocellus  consists  of  a 
spot  within  a  ring  of  another  colour,  like  the  pupil  within  the 

*^  Bechsteia, 'Naluvgeschichte  Deutschlands,'  B.  iv,  1795,  s.  31,  on  a 
sub-variety  of  the  Monck  pigeou. 


428'  The  Descent  of  Man.  Part  II. 


iris,  but  the  central  spot  is  often  surrounded  by  additional  con- 
centric zones.  The  ocelli  on  the  tail-coverts  of  the  peacock  offer 
a  familiar  example,  as  well  as  those  on  the  wings  of  the  peacock- 
butterfly  (Vanessa).  Mr.  Trimen  has  given  me  a  description  of 
a  S.  African  moth  {Gyananis  isis),  allied  to  our  Emperor  moth, 
in  which  a  magnificent  ocellus  occupies  nearly  the  whole  surface 
of  each  hinder  wing ;  it  consists  of  a  black  centre,  including  a 
semi-transparent  crescent-shaped  mark,  surrounded  by  successive, 
ochre-yellow,  black,  ochre-yellow,  pink,  white,  pink,  brown,  and 
whitish  zones.  Although  we  do  not  know  the  steps  by  which 
these  wonderfully  beautiful  and  complex  ornaments  have  been 
developed,  the  process  has  probably  been  a  simple  one,  at  least 
with  insects  ;  for,  as  Mr.  Trimen  writes  to  me,  "  no  characters  of 
"  mere  marking  or  coloration  are  so  unstable  in  the  Lepidoptera 
"  as  the  ocelli,  both  in  number  and  size."  Mr.  Wallace,  who 
first  called  my  attention  to  this  subject,  shewed  me  a  series  of 
specimens  of  our  common  meadow-brown  butterfly  (Hipparchia 
j<mira)  exhibiting  numerous  gradations  from  a  simple  minute 
black  spot  to  an  elegantly-shaded  ocellus.  In  a  S.  African 
butterfly  {Cullo  leda,  Linn.),  belonging  to  the  same  family,  the 
ocelli  are  even  still  more  varia,ble.  In  some  si)ecimens  (A,  fig.  53) 
large  spaces  on  the  upper  surface  of  the  wings  are  coloured 
black,  and  include  irregular  white  marks ;  and  from  this  state 
a  complete  gradation  can  be  traced  into  a  tolerably  perfect 
ocellus  (A^),  and  this  results  from  the  contraction  of  the  irregular 
blotches  of  colour.  In  another  series  of  specimens  a  gradation 
can  be  followed  from  excessively  minute  white  dots,  surrounded 
by  a  scarcely  visible  black  line  (B),  into  perfectly  symmetrical 
and  large  ocelli  (B').**  In  cases  like  these,  the  development 
of  a  perfect  ocellus  does  not  require  a  long  course  of  variation 
and  selection. 

With  birds  and  many  other  animals,  it  seems  to  follow  from 
the  comparison  of  allied  species  that  circular  spots  are  often 
generated  by  the  breaking  up  and  contraction  of  stripes.  In  the 
Tragopan  pheasant  faint  white  lines  in  the  female  represent  the 
beautiful  white  spots  in  the  male ;  *^  and  something  of  the  same 
kind  may  be  observed  in  the  two  sexes  of  the  Argus  pheasant. 
However  this  may  be,  appearances  strongly  favour  the  belief  that 
on  the  one  hand,  a  dark  spot  is  often  formed  by  the  colouring 

^*  This    woodcut     has    been    en-  wings     of    this    butterfly,    in    his 

graved   from   a    beautiful  drawing,  '  Rhopalocera  Africae  Australis,'  p, 

most  kindly  made  for    me  by  Mr.  186. 

Trimen;  see  also  his  description  of  *'  Jerdon,  'Birds  of  India,'  vol.- 

the  wonderful  amount  of  variation  iii.  p.  517. 
iu  the  coloration  and  shape  of  the 


Chap.  XIV. 


Ocelli. 


429 


matter  being  drawn  towards  a  central  point  from  a  surrounding 
zone,  which  latter  is  thus  rendered  lighter ;  and,  on  the  other 
hand,  that  a  white  spot  is  often  formed  by  the  colour  being  driven 
away  from  a  central  point,  so  that  it  accumulates  in  a  surround- 
ing darker  zone.  In  either  case  an  ocellus  is  the  result.  The 
colouring  matter  seems  to  be  a  nearly  constant  quantity,  but  is 
redistributed,  either  centripetally  or  centrifugally.    The  feathers 


Fig.  5.3.    Cyllo  leda,  Linn.,  from  a  drawing  by  Mr.  Triinen,  shewing  the  extreme 
range  of  variation  in  the  ocelli. 


A.  Specimen,  from    Mauritius,  upper 

surface  of  fore-wing. 
Ai.  Specimen,  from  Natal,  ditto. 


B.  Specimen,  from  Java,  upper  surface 

of  hind-wing. 
&.  Specimen,  from  Mauritius,  ditto. 


of  the  common  guinea-fowl  offer  a  good  instance  of  white  spots 
surrounded  by  darker  zones ;  and  wherever  the  Avhite  spots  are 
large  and  stand  near  each  other,  the  surrounding  dark  zones 
become  confluent.  In  the  same  wing-feather  of  the  Argus 
pheasant  dark  spots  may  be  seen  surrounded  by  a  pale  zone, 
and  white  spots  by  a  dark  zone.  Thus  the  formation  of  an 
ocellus  in  its  most  elementary  state  appears  to  be  a  simple  affair. 
By  what  further  steps  the  more  complex  ocelli,  which  are  sur- 
rounded by  many  successive  zones  of  colour,  have  been  generated, 
I  will  not  pretend  to  say.  But  the  zoned  feathers  of  the  mongrels 
from  differently  coloured  fowls,  and  the  extraordinary  variability 
of  the  ocelli  on  many  Lepidoptera,  lead  us  to  conclude  that  their 
formation  is  not  a  complex  process,  but  depends  on  some  slight 
and  graduated  change  in  the  nature  of  the  adjoining  tissues. 


430  The  Descent  of  Man.  Part  II. 

Gradation  of  Secondary  Sexual  Characters. — Cases  of  gradation 
are  important,  as  shewing  us  that  highly  complex  ornaments 
may  be  acquired  by  small  successive  steps.  In  order  to  discover 
the  actual  steps  by  which  the  male  of  any  existing  bird  has 
acquired  his  magnificent  colours  or  other  ornaments,  we  ought 
to  behold  the  long  line  of  his  extinct  progenitors ;  but  this  is 
obviously  impossible.  We  may,  however,  generally  gain  a  clue 
by  comparing  all  the  species  of  the  same  group,  if  it  be  a  large 
one;  for  some  of  them  will  probably  retain,  at  least  partially, 
traces  of  their  former  characters.  Instead  of  entering  on 
tedious  details  respecting  various  groups,  in  which  striking 
instances  of  gradation  could  be  given,  it  seems  the  best  plan  to 
take  one  or  two  strongly  marked  cases,  for  instance  that  of  the 
peacock,  in  order  to  see  if  light  can  be  thrown  on  the  steps  by 
which  this  bird  has  become  so  splendidly  decorated.  The 
peacock  is  chiefly  remarkable  from  the  extraordinary  length  of  his 
tail-coverts ;  the  tail  itself  not  being  much  elongated.  The  barbs 
along  nearly  the  whole  length  of  these  feathers  stand  separate  or 
are  decomposed ;  but  this  is  the  case  with  the  feathers  of  many 
species,  and  with  some  varieties  of  the  domestic  fowl  and  pigeon. 
The  barbs  coalesce  towards  the  extremity  of  the  shaft  forming 
the  oval  disc  or  ocellus,  which  is  certainly  one  of  the  most 
beautiful  objects  in  the  world.  It  consists  of  an  iridescent,  in- 
tensely blue,  indented  centre,  surrounded  by  a  rich  green  zone, 
this  by  a  broad  coppery-brown  zone,  and  this  by  five  other  narrow 
zones  of  slightly  different  iridescent  shades.  A  trifling  character 
in  the  disc  deserves  notice;  the  barbs,  for  a  space  along  one  of 
the  concentric  zones  are  more  or  less  destitute  of  their  barbules, 
so  that  a  part  of  the  disc  is  surrounded  by  an  almost  transparent 
zone,  which  gives  it  a  highly  finished  aspect.  But  I  have  else- 
where described  ^^  an  exactly  analogous  variation  in  the  hackles 
of  a  sub-variety  of  the  game-cock,  in  which  the  tips,  having  a 
metallic  lustre,  "are  separated  from  the  lower  part  of  the 
"  feather  by  a  symmetrically  shaped  transparent  zone,  composed 
"  of  the  naked  portions  of  the  barbs."  The  lower  margin  or 
base  of  the  dark-blue  centre  of  the  ocellus  is  deeply  indented 
on  the  line  of  the  shaft.  The  surrounding  zones  likewise  shew 
traces,  as  may  be  seen  in  the  drawing  (fig.  54),  of  indentations, 
or  rather  breaks.  These  indentations  are  common  to  the  Indian 
and  Javan  peacocks  {Pavo  cristntns  and  P.  muticvs),  and  they 
seemed  to  deserve  particular  attention,  as  probably  connected 
with  the  development  of  the  ocellus  ;  but  for  a  long  time  I  could 
not  conjecture  their  meaning. 

so  i  Variation  of  Animals  and  Plants  under  Domestication,'  vol.  i.  p.  254 


Chap.  XIV. 


Gradation  of  Characters. 


431 


If  we  admit  the  principle  of  gradual  evolution,  there  must 
formerly  have  existed  many  species  which  presented  every  suc- 
cessive step  between  the  wonderfully  elongated  tail-coverts  of 
the  peacock  and  the  short  tail-coverts  of  all  ordinary  birds ;  and 
again  between  the  magnificent  ocelli  of  the  former,  and  the 
simpler  ocelli  or  mere  coloured  spots  on  other  birds ;  and  so  with 
all  the  other  characters  of  the  peacock.  Let  us  look  to  the  allied 
GallinacesB  for  any  still-existing  gradations.    The  species  and 


Fig.  54.  Feather  of  Peacock,  about  two-thirds  of  natural  size,  drawn  hy  Mr.  Ford. 
The  transparent  zone  is  represented  by  the  outermost  white  zone,  confimd  to  the 
upper  end  of  the  disc. 

sub-species  of  Polyplectron  inhabit  countries  adjacent  to  the 
native  land  of  the  peacock ;  and  they  so  far  resemble  this  bird 
that  they  are  sometimes  called  peacock-pheasants.  I  am  also  in- 
formed by  Mr.  Bartlett  that  they  resemble  the  peacock  in  their 
voice  and  in  some  of  their  habits.  During  the  spring  the  males, 
as  previously  described,  strut  about  before  the  comparatively 
plain-coloured  females,  expanding  and  erecting  tlieir  tail  and 


432  The  Descent  of  Man.  Part  II. 

wing-feathers,  which  are  ornamented  with  numerous  ocelli.  I 
request  the  reader  to  turn  back  to  the  drawing  (fig.  51,  p.  397) 
of  a  Polyplectron.  In  P.  riapohonis  the  ocelli  are  confined  to 
the  tail  and  the  back  is  of  a  rich  metallic  blue ;  in  which  re- 
spects this  species  approaches  the  Java  peacock.  P.  hardicickii 
possesses  a  peculiar  top-knot,  which  is  also  somewhat  like 
that  of  the  Java  peacock.  In  all  the  species  the  ocelli  on  the 
wings  and  tail  are  either  circular  or  oval,  and  consist  of  a 
beautiful,  iridescent,  greenish-blue  or  greenish-purple  disc,  with 
a  black  border.  This  border  in  P.  chinqiiis  shades  into  brown, 
edged  with  cream-colour,  so  that  the  ocellus  is  here  surrounded 
with  variously  shaded,  though  not  bright,  concentric  zones.  The 
unusual  length  of  the  tail-coverts  is  another  remarkable  character 
in  Polyplectron ;  for  in  some  of  the  species  they  are  half,  and  in 
others  two-thirds  as  long  as  the  true  tail-feathers.  The  tail- 
coverts  are  ocellated  as  in  the  peacock.  Thus  the  several  species 
of  Polyplectron  manifestly  make  a  graduated  approach  to  the 
peacock  in  the  length  of  their  tail-coverts,  in  the  zoning  of  the 
ocelli,  and  in  some  other  characters. 

Notwithstanding  this  approach,  the  first  species  of  Polyplectron 
which  I  examined  almost  made  me  give  up  the  search ;  for  I 
found  not  only  that  the  true  tail-feathers,  which  in  the  peacock 
are  quite  plain,  were  ornamented  with  ocelli,  but  that  the  ocelli 
on  all  the  feathers  differed  fundamentally  from  those  of  the 
peacock,  in  there  being  two  on  the  same  feather  (fig.  55),  one  on 
each  side  of  the  shaft.  Hence  I  concluded  that  the  early  pro- 
genitors of  the  peacock  could  not  have  resembled  a  Polyplectron. 
But  on  continuing  my  search,  I  observed  that  in  some  of  the 
species  the  two  ocelli  stood  very  near  each  other ;  that  in  the 
tail-feathers  of  P.  hardwickii  they  touched  each  other;  and. 
finally,  that  on  the  tail-coverts  of  this  same  species  as  well  as  of 
P.  malaccense  (fig.  56)  they  were  actually  confluent.  As  the 
central  part  alone  is  confluent,  an  indentation  is  left  at  both  the 
upper  and  lower  ends  ;  and  the  surrounding  coloured  zones  are 
likewise  indented.  A  single  ocellus  is  thus  formed  on  each  tail- 
covert,  though  still  plainly  betraying  its  double  origin.  These 
confluent  ocelli  differ  from  the  single  ocelli  of  the  peacock  in 
having  an  indentation  at  both  ends,  instead  of  only  at  the  lower 
or  basal  end.  The  explanation,  however,  of  this  difference  is 
not  difiicult ;  in  some  species  of  Polyplectron  the  two  oval 
ocelli  on  the  same  feather  stand  parallel  to  each  other ;  in  other 
species  (as  in  P.  cMiiquu)  they  converge  towards  one  end;  now 
the  partial  confluence  of  two  convergent  ocelli  would  manifestly 
leave  a  much  deeper  indentation  at  the  divergent  than  at  the 
convergent  end.     It  is  also  manifest  that  if  the  convergence  were 


Chap.  XIV.     Birds — Gradation  of  Characters. 


433 


strongly  pronounced  and  the  confluence  complete,  the  indentation 
at  the  convergent  end  would  tend  to  disappear. 

The  tail-feathers  in  both  species 
of  peacock  are  entirely  destitute 
of  ocelli,  and  this  apparently  is 
related  to  their  being  covered  up 
and  concealed  by  the  long  tail- 
coverts,  lu  this  respect  they 
differ  remarkably  from  the  tail- 
feathers  of  Polyplectron,  which 
in  most  of  the  species  are  or- 
namented with  larger  ocelli  than 
those  on  the  tail-coverts.  Hence 
I  was  led  carefully  to  examine  the 
tail-feathers  of  the  several  species, 
in  order  to  discover  whether  their 
ocelli  shewed  any  tendency  to  dis- 
appear; and  to  my  great  satis- 
faction, this  appeared  to  be  so. 
The  central  tail-feathers  of  P. 
7iapoleonis  have  the  two  ocelli  on 
each  side  of  the  shaft  perfectly 
developed ;  but  the  inner  ocellus 
becomes  less  and  less  conspicuous 
on  the  more  exterior  tail-feathers, 
until  a  mere  shadow  or  rudiment 
is  left  on  the  inner  side  of  the 
outermost  feather.  Again,  in  P. 
malaccense,  the  ocelli  on  the  tail- 
coverts  are,  as  we  have  seen, 
confluent ;  and  these  feathers  are 
of  unusual  length,  being  two- 
thirds  of  the  length  of  the  tail- 
feathers,  so  that  in  both  these 
respects  they  approach  the  tail- 
coverts  of  the  peacock.  Now  in 
P.  malaccense  the  two  central  tail- 
feathers  alone  are  ornamented, 
each  with  two  brightly-coloured 
ocelli,  the  inner  ocellus  having 
completely  disappeared  from  all 
the  other  tail-feathers.  Conse- 
quently the  tail-coverts  and  tail-feathers  of  this  species  of  Poly- 
plectron make  a  near  approach  in  structure  and  ornamentation 
to  the  corresponding  feathery  of  the  peacock. 


Fig.  55.  Part  of  a  tail-covert  of  Poly- 
plectron chinquis,  with  the  two  ocelli 
of  nat.  size. 


mm 


FiR.  56.  Part  of  a  tail-covcrt  of  Poly- 
plectron  malaccense,  with  the  two 
ocelli,  partially  confluent,  of  nat.  size. 


434  ^^^^  Descent  of  Man,  Fart  II. 


As  far,  then,  as  gradation  throws  light  on  the  steps  by  which 
the  magnificent  train  of  the  peacock  has  been  acquired,  hardly 
anything  more  is  needed.  If  we  picture  to  ourselves  a  progeni- 
tor of  the  peacock  in  an  almost  exactly  intermediate  condition 
between  the  existing  peacock,  with  his  enormously  elongated 
tail-coverts,  ornamented  with  single  ocelli,  and  an  ordinary 
gallinaceous  bird  with  short  tail-coverts,  merely  spotted  with 
some  colour,  we  shall  see  a  bird  allied  to  Polyplectron — that  is, 
with  tail-coverts,  capable  of  erection  and  expansion,  ornamented 
with  two  partially  confluent  ocelli,  and  long  enough  almost  to 
conceal  the  tail-feathers,  the  latter  having  already  partially  lost 
their  ocellL  The  indentation  of  the  central  disc  and  of  the 
surrounding  zones  of  the  ocellus,  in  both  species  of  peacock, 
speaks  plainly  in  favour  of  this  view,  and  is  otherwise  in- 
explicable. The  males  of  Polyplectron  are  no  doubt  beautiful 
birds,  but  their  beauty,  when  viewed  from  a  little  distance, 
cannot  be  compared  with  that  of  the  peacock.  Many  female 
progenitors  of  the  peacock  must,  during  a  long  line  of  descent, 
have  appreciated  this  superiority ;  for  they  have  unconsciously, 
by  the  continued  preference  of  the  most  beautiful  males,  rendered 
the  peacock  the  most  splendid  of  hving  birds. 

Argus  pheasant. — Another  excel Itnt  case  for  investigation  is 
offered  by  the  ocelli  on  the  wing-feathers  of  the  Argus  pheasant, 
which  are  shaded  in  so  wonderful  a  manner  as  to  resemble  balls 
lying  loose  within  sockets,  and  consequently  differ  from  ordinary 
ocelli.  No  one,  I  presume,  will  attribute  the  shading,  which  has 
excited  the  admiration  of  many  experienced  artists,  to  chance 
— to  the  fortuitous  concourse  of  atoms  of  colouring  matter. 
That  these  ornaments  should  have  been  formed  through  the 
selection  of  many  successive  variations,  not  one  of  which  was 
originally  intended  to  produce  the  ball-and-socket  effect,  seems 
as  incredible,  as  that  one  of  Piaphael's  Madonnas  should  have 
been  formed  by  the  selection  of  chance  daubs  of  paint  made  by  a 
long  succession  of  young  artists,  not  one  of  whom  intended  at 
first  to  draw  the  human  figure.  In  order  to  discover  how  the 
ocelli  have  been  developed,  we  cannot  look  to  a  long  line  of 
progenitors,  nor  to  many  closely-allied  forms,  for  such  do  not 
now  exist.  But  fortunately  the  several  feathers  on  the  wing 
suffice  to  give  us  a  clue  to  the  problem,  and  they  prove  to  de- 
monstration that  a  gradation  is  at  least  possible  from  a  mere 
sjDot  to  a  finished  ball-and-socket  ocellus. 

The  wing- feathers,  bearing  the  oceUi,  are  covered  with  dark 
stripes  (fig.  67)  or  with  rows  of  dark  spots  (fig.  59),  each  stripe 
or  row  of  spots  running  obliquely  down  the  outer  side  of  the 


Chap.  XIV.     Birds  —  Gradation  of  Characters.  435 


shaft  to  one  of  the  ocelli.     The  s]Dots  are  generally  elongated  in  a 

line  transverse  to  the  row  in  which  they  stand.     They  often 

become  conflueut,  either  in  the  line  of  the  row— and  then  they 

form  a  longitudinal  stripe— or   transversely,  that  is,  with  the 

spots  in  the   adjoining 

rows,   and    then    they 

form  trano  verse  stripes. 

A      spot       sometimes 

breaks  up  into  smaller 

spots,  which  still  stand 

in  their  proper  places. 

It  will  be  convenient 
first  to  describe  a  per- 
fect ball  -  and  -  socket 
ocellus.  This  consists 
of  an  intensely  black 
circular  ring,  surround- 
ing a  space  shaded  so 
as  exactly  to  resemble 
a  ball.  The  figure  here 
given  has  been  ad- 
mirably drawn  by  Mr. 
Ford  and  well  engraved, 
but  a  woodcut  cannot 
exhibit  the  exquisite 
shading  of  the  original. 
The  ring  is  almost 
always  slightly  broken 
or  interrupted  (see  fig. 
57)  at  a  point  in  the 
upper  half,  a  little  to 
the  right  of,  and  above 
the  white  shade  on  the 
enclosed  ball ;  it  is  also 
sometimes  broken  to- 
wards the  base  on  the 
right     hand.        These 

little  breaks  have  an  important  meaning.  The  ring  is  always 
much  thickened,  with  the  edges  ill-defined  towards  the  left- 
hand  upper  corner,  the  feather  being  held  erect,  in  the  posi- 
tion in  which  it  is  here  drawn.  Beneath  this  thickened  part 
there  is  on  the  surface  of  the  ball  an  oblique  almost  pure-white 
mark,  which  shades  off  downwards  into  a  pale-leaden  hue,  and 
this  into  yellowish  and  brown  tints,  which  insensibly  become 


Fie;.  57.     Part  of  secondary  wing-feather  of  Argus 
pheasant,   shewing  two   perfect  ocelli,  a   and  \j 
A,  B,  C,  D,  &c.,  are  dark  stripes  running  obliquely 
down,  each  to  an  ocellus. 
[Much  of  the  web  on  both  sides,  especially  to  the 

left  of  the  shaft,  has  been  cut  off.] 


43^ 


The  Descent  of  Man. 


Part  II. 


darker  and  darker  towards  the  lower  part  of  the  ball.  It  is  this 
shading  which  gives  so  admirably  the  effect  of  light  shining  on 
a  convex  surface.  If  one  of  the  balls  be  examined,  it  will  be  seen 
that  the  lower  part  is  of  a  brown  tint  and  is  indistinctly  sepa- 
rated by  a  curved  oblique  line  from  the  upper  part,  which  is 
yellower  and  more  leaden;  this  curved  oblique  line  runs  at  right 
angles  to  the  longer  axis  of  the  white  patch  of  light,  and  indeed 
of  all  the  shading ;  but  this  difference  in  colour,  which  cannot 
of  course  be  shewn  in  the  woodcut,  does  not  in  the  least  interfere 
with  the  perfect  shading  of  the  ball.  It  should  be  particularly 
observed  that  each  ocellus  stands  in  obvious  connection  either 
with  a  dark  stripe,  or  with  a  longitudinal  row  of  dark  spots,  for 
both  occur  indifferently  on  the  same  feather.  Thus  in  tig.  57  stripe 
A  runs  to  ocellus  a ;  B  runs  to  ocellus  h  ;  stripe  C  is  broken  in 
the  upper  part,  and  runs  down  to  the  next  succeeding  ocellus, 

not  represented  in  the  wood- 
cut ;  D  to  the  next  lower  one, 
and  zo  w'ith  the  strijDes  E 
and  F.  Lastly,  the  several 
ocelli  are  separated  from  each 
other  by  a  pale  surface  bear- 
ing irregular  black  marks. 

I  will  next  describe  the 
other  extreme  of  the  series, 
namely,  the  first  trace  of 
an  ocellus.  The  short  se- 
condary wing -feather  (fig. 
58),  nearest  to  the  body, 
is  marked  like  the  other 
feathers,  with  oblique,  lon- 
gitudinal, rather  irregular, 
rows  of  very  dark  spots. 
The  basal  spot,  or  that  near- 
est the  shaft,  in  the  five  lower 
rows  (excluding  the  lowest 
one)  is  a  little  larger  than 
the  other  spots  of  the  same 
row,  and  a  little  more  elon- 
gated in  a  transverse  direc- 
tion. It  differs  also  from  the  other  spots  by  being  bordered 
on  its  upper  side  with  some  dull  fulvous  shading.  But  this 
spot  is  not  in  any  way  more  remarkable  than  those  on  the 
plumage  of  many  birds,  and  might  easily  be  overlooked.  The 
next  higher  spot  does  not  differ  at  all  from  the  upper  ones  in 


Fig.  58.     Basal  part  of  the  secondary-wing 
feather,  nearest  to  the  body. 


Chap.  XIV.     Birds — Gradation  of  Characters, 


437 


the  same  row.  The  larger  basal  spots  occupy  exactly  the  same 
relative  position  on  these  feathers,  as  do  the  perfect  ocelli  on 
the  longer  wing-feathers. 

By  looking  to  the  next  two  or  three  succeeding  wing-feathers, 
an  absolutely  insensible  gradation  can  be  traced  from  one  of  the 
last-described  basal  spots,  together  with  the  next  higher  one  in 
the  same  row,  to  a  curious  ornament,  which  cannot  be  called  an 
ocellus,  and  which  I  will  name,  from  the  want  of  a  better  term, 
an  "  elliptic  ornament."  These  are  shewn  in  the  accompanying 
figure  (tig.  59).     We  here  see  several  oblique  rows.  A,  B,  C,  D, 


Fig.  59.  Portion  of  one  of  the  secondary  wing- feathers  near  to  the  body,  shewing  the 
so-called  elliptic  ornaments.  The  right-hand  figure  is  given  merely  as  a  diagram 
for  the  sake  of  the  letters  of  reference. 


A,  H,  C,  I),  &c.  Eows  of  spots  running 
down  to  and  forming  the  elliptic 
ornaments. 

&.  Lowest  spot  or  mark  in  row  B. 


C.  The  next  succeeding  spot  or  mark  in 

the  same  row. 
d.  Apparently  a  brokon  prolongation  of 

the  spot  c  in  the  same  row  B 


&c.  (see  the  lettered  diagram  on  the  right  hand),  of  dark  spots 
of  the  usual  character.  Each  row  of  spots  runs  down  to  and  is 
connected  with  one  of  the  elliptic  ornaments,  in  exactly  the 
same  manner  as  each  stripe  in  tig.  57  runs  down  to,  and  is 
connected  with,  one  of  the  ball-and-socket  ocelli.  Looking  to 
any  one  row,  for  instance,  B,  in  fig.  59,  the  lowest  mark  (/>)  is 
thicker  and  considerably  longer  than  tbe  up]ior  spots,  and  has 
its  left  extremity  pointed  and  curved  upward.s.    This  black  marlc 


438  The  Descent  of  Man.  Part  II. 

is  abruptly  bordered  on  its  upper  side  by  a  rather  broad  space  of 
richly  shaded  tints,  beginning  with  a  narrow  brown  zone,  which 
passes  into  orange,  and  this  into  a  pale  leaden  tint,  with  the  end 
towards  the  shaft  much  paler.  These  shaded  tints  together  fill  up 
the  whole  inner  space  of  the  elliptic  ornament.  The  mark  (6) 
corresponds  in  every  respect  with  the  basal  shaded  spot  of  the 
simple  feather  described  in  the  last  paragraph  (fig.  58),  but  is 
more  highly  developed  and  more  brightly  coloured.  Above  and  to 
the  right  of  this  spot  (6  fig.  59),  with  its  bright  shading,  there  is 
a  long  narrow,  black  mark  (c),  belonging  to  the  same  row,  and 
which  is  arched  a  little  downwards  so  as  to  face  (&).  This  mark 
is  sometimes  broken  into  two  portions.  It  is  also  narrowly  edged 
on  the  lower  side  with  a  fulvous  tint.  To  the  left  of  and  above 
c,  in  the  same  oblique  direction,  but  always  more  or  less  distinct 
from  it,  there  is  another  black  mark  {<i).  This  mark  is  generally 
sub-triangular  and  irregular  in  shape,  but  in  the  one  lettered  in 
the  diagram  it  is  unusually  narrow,  elongated,  and  regular.  It 
apparently  consists  of  a  lateral  and  broken  jDrolongation  of  the 
mark  (c),  together  with  its  confluence  with  a  broken  and  prolonged 
part  of  the  next  spot  above  ;  but  I  do  not  feel  sure  of  this.  These 
three  marks,  6,  c-,  and  c/,  with  the  intervening  bright  shades,  form 
together  the  so-called  elliptic  ornament.  These  ornaments  placed 
parallel  to  the  shaft,  manifestly  correspond  in  position  with  the 
ball-and-socket  ocelli.  Their  extremely  elegant  appearance  can- 
not be  appreciated  in  the  drawing,  as  the  orange  and  leaden  tints, 
contrasting  so  well  with  the  black  marks,  cannot  be  shewn. 

Between  one  of  the  elliptic  ornaments  and  a  perfect  ball-and- 
socket  ocellus,  the  gradation  is  so  perfect  that  it  is  scarcely 
possible  to  decide  when  the  latter  term  ought  to  be  used.  The 
passage  from  the  one  into  the  other  is  effected  by  the  elongation 
and  greater  curvature  in  opposite  directions  of  the  lower  black 
mark  (&  fig.  59),  and  more  especially  of  the  upf)er  one  (c\ 
together  with  the  contraction  of  the  elongated  sub-triangular  or 
narrow  mark  (c/),  so  that  at  last  these  three  marks  become  con- 
fluent, forming  an  irregular  elliptic  ring.  This  ring  is  gradually 
rendered  more  and  more  circular  and  regular,  increasing  at  the 
same  time  in  diameter.  I  have  here  given  a  drawing  (fig.  60) 
of  the  natural  siz^  of  an  ocellus  not  as  yet  quite  perfect.  The  lower 
part  of  the  black  ring  is  much  more  curved  than  is  the  lower 
mark  in  the  elliptic  ornament  (h  fig.  59).  The  upper  part  of 
the  ring  consists  of  two  or  three  separate  portions ;  and  there  is 
only  a  trace  of  the  thickening  of  the  portion  which  forms  the 
black  mark  above  the  white  shade.  This  white  shade  itself  is 
not  as  yet  much  concentrated ;  and  beneath  it  the  surface  is 
brighter  coloured  than  in  a  perfect  ball-and-socket  ocellus.     Even 


Chap.  XIV.    Birds — Gradation  of  Characters. 


439 


in  the  most  perfect  ocelli,  traces  of  the  junction  of  three  or  four 
elongated  black  marks,  by  which  the  ring  has  been  formed,  may 
often  be  detected.  The  irregular  sub-triangular  or  narrow  mark 
{d  fig.  59),  manifestly  forms,  by  its 
contraction  and  equalisation,  the 
thickened  portion  of  the  ring  above 
the  white  shade  on  a  perfect  ball- 
and-socket  ocellus.  The  lower  part 
of  the  ring  is  invariably  a  little 
thicker  than  the  other  parts  (see 
fig.  67),  and  this  follows  from  the 
lower  black  mark  of  the  elliptic 
ornament  {h  fig.  59)  having  origi- 
nally been  thicker  than  tlie  upper 
mark  (c).  Every  step  can  be  fol- 
lowed in  the  process  of  confluence 
and  modification ;  and  the  black 
ring  which  surrounds  the  ball  of 
the  ocellus  is  unquestionably  formed 
by  the  union  and  modification  of 
the  three  black  marks,  h,  c,  d,  of 
the  elliptic  ornament.  The  irre- 
gular zigzag  black  marks  between 
the  successive  ocelli  (see  again  fig. 
57)  are  plainly  due  to  the  breaking 
up  of  the  somewhat  more  regular  but  similar  marks  between 
the  elliptic  ornaments. 

The  successive  steps  in  the  shading  of  the  ball-and-socket 
ocelli  can  be  followed  out  with  equal  clearness.  The  brown, 
orange,  and  pale  leaden  narrow  zones,  which  border  the  lower 
black  mark  of  the  elliptic  ornament,  can  be  seen  gradually  to 
become  more  and  more  softened  and  shaded  into  each  other, 
with  the  upper  lighter  part  towards  the  left-hand  corner  ren- 
dered still  lighter,  so  as  to  become  almost  white,  and  at  the  same 
time  more  contracted.  But  even  in  the  most  perfect  ball-and- 
socket  ocelli  a  shght  difference  in  the  tints,  though  not  in  the 
shading,  between  the  upper  and  lower  parts  of  the  ball  can  be 
perceived,  as  before  noticed ;  and  the  line  of  separation  is  oblique, 
in  the  same  direction  as  the  bright-coloured  shades  of  the 
elliptic  ornaments.  Thus  almost  every  minute  detail  in  the 
shape  and  colouring  of  the  ball-and-socket  ocelli  can  be  shewn  to 
follow  from  gradual  changes  in  the  elliptic  ornaments ;  and  the 
development  of  the  latter  can  be  traced  by  equally  small  steps 
from  the  union  of  two  almost  simple  spots,  the  lower  one  (fig.  58) 
having  some  dull  fulvous  shading  on  its  upper  side. 


Fig.  60.  An  ocellus  in  an  inter- 
mediate condition  between  tbe 
elliptic  ornament  and  the  perfect 
ball-and-socket  ocellus. 


440 


The  Desce?tt  of  Man. 


Past  II 


The  extremities  of  tlie  longer  secondary  feathers  which  bear 
the  perfect  ball-and-socket  ocelli,  are  peculiarly  ornamented 
(fig.  61).  The  oblique  longitudinal  striiDes  suddenly  cease 
upwards  and  become  confused ;  and  above  this  limit  the  whole 

upper  end  of  the  feather  ('/)  is 
covered  with  white  dots,  sur- 
rounded by  little  black  rings, 
standing  on  a  dark  ground. 
The  oblique  stripe  belonging  to 
the  npjDermost  ocellus  {h)  is 
barely  represented  by  a  very 
short  iiTegular  black  mark  ^^  ith 
the  usual,  curved,  tran verse  base. 
As  tills  stripe  is  thus  abruptly 
cut  off,  we  can  perhaps  under- 
stand from  what  has  gone  before, 
how  it  is  that  the  upper  thickened 
part  of  the  ring  is  here  absent ;  for, 
as  before  stated,  this  thickened 
part  apparently  stands  in  some 
relation  with  a  broken  prolonga- 
tion from  the  next  higher  spot. 
From  the  absence  of  the  upper 
and  thickened  part  of  the  ring, 
the  uppermost  ocellus,  though 
perfect  in  all  other  respects, 
appears  as  if  its  top  had  been 
obliquely  sliced  off.  It  would, 
I  think,  perplex  any  one,  who 
believes  that  the  plumage  of 
the  Argus-pheasant  was  created 
as  we  now  see  it,  to  account  for 
the  imperfect  condition  of  the 
uppermost  ocellus.  I  should  add 
that  on  tlie  secondary  wing- 
feather  farthest  from  the  body, 
all  the  ocelli  are  smaller  and 
less  perfect  than  on  the  other 
feathers,  and  have  the  upper 
part  of  the  ring  deficient,  as  in  the  case  just  mentioned.  The 
imperfection  here  seems  to  be  connected  with  the  fact  that  the 
spots  on  this  feather  shew  less  tendency  than  usual  to  become 
confluent  into  stripes  ;  they  are  on  the  contrary,  often  broken  up 
into  smaller  spots,  so  that  two  or  three  rows  run  down  to  the 
same  ocellus. 


Fi.;-.  61.  Portion  near  summit  of  one  of 
the  secondary  wing-feathers,  bearing 
perfect  ball-and-socket  ocelli. 

a.  Ornamented  npper  part. 

h.  13  i>permost,  imperfect  ball-and-socket 
ocellus.  (The  shading  above  the 
white  mark  on  the  summit  of  the 
ocellus  is  here  a  little  too  dark.) 

c.  Perfect  ocellus. 


CHAr.  XIV.    Birds — Gradation  of  Characters.  441 

There  still  remains  another  very  curious  point,  first  observed 
by  Mr,  T.  W.  Wood;'^^  which  deserves  attention.  In  a  photograph, 
given  me  by  Mr.  Ward,  of  a  specimen  mounted  as  in  the  act  of 
display,  it  may  be  seen  that  on  the  feathers  which  are  held 
perpendicularly,  the  white  marks  on  the  ocelli,  representing 
light  reflected  from  a  convex  surface,  are  at  the  upper  or 
further  end,  that  is,  are  directed  upwards  ;  and  the  bird  whilst 
displaying  himself  on  the  ground  would  naturally  be  illuminated 
from  above.  But  here  comes  the  curious  point,  the  outer 
feathers  are  held  almost  horizontally,  and  their  ocelli  ought  like- 
wise to  appear  as  if  illuminated  from  above,  and  consequently 
the  white  marks  ought  to  be  placed  on  the  upper  sides  of  the 
ocelli ;  and  wonderful  as  is  the  fact  they  are  thus  placed  !  Hence 
the  ocelli  on  the  several  feathers,  though  occupying  very  different 
positions  with  respect  to  the  light,  all  appear  as  if  illumJnated 
from  above,  just  as  an  artist  would  have  shaded  them.  Never- 
theless they  are  not  illuminated  from  strictly  the  same  point 
as  they  ought  to  be ;  for  the  white  marks  on  the  ocelli  of  the 
feathers  which  are  held  almost  horizontally,  are  placed  rather 
too  much  towards  the  further  end ;  that  is  they  are  not  suffi- 
ciently lateral.  We  have,  however,  no  right  to  expect  absolute 
perfection  in  a  part  rendered  ornamental  through  sexual  selec- 
tion, any  more  than  we  have  in  a  part  modified  through  natural 
selection  for  real  use ;  for  instance  in  that  wondrous  organ  the 
human  eye.  And  we  know  what  Helmholtz,  the  highest  authority 
in  Europe  on  the  subject,  has  said  about  the  human  eye;  that 
if  an  optician  had  sold  him  an  instrument  so  carelessly  made,  he 
would  have  thought  himself  fully  justified  in  returning  it.^^ 

We  have  now  seen  that  a  perfect  series  can  be  followed,  from 
simple  spots  to  the  wonderful  ball-and-socket  ornaments.  Mr. 
Gould,  who  kindly  gave  me  some  of  these  feathers,  fully  agrees 
with  me  in  the  completeness  of  the  gradation.  It  is  obvious 
that  the  stages  in  development  exhibited  by  the  feathers  on  the 
same  bird,  do  not  at  all  necessarily  shew  us  the  steps  passed 
through  by  the  extinct  pj'ogenitors  of  the  species;  but  they 
probably  give  us  the  clue  to  the  actual  steps,  and  they  at  least 
prove  to  demonstration  that  a  gradation  is  possible.  Bearing  in 
mind  how  carefully  the  male  Argus  pheasant  displays  his  plumes 
before  the  female,  as  well  as  the  many  facts  rendering  it  probable 
that  female  birds  prefer  the  more  attractive  males,  no  one  who 
admits  the  agency  of  sexual  selection  in  any  case,  will  deny  that 
a  simple  dark  spot  with  some  fulvous  shading  might  be  converted, 
through  the  approximation  and  modification  of  two  adjoining 

*»  The  'Field,'  May  28,  1870.  Subjects,'  Eng.  trans,  1873,  pp.  219, 

Popular  Lectures  on  Scientific      227,  269,  390. 
20 


442  The  Descent  of  Man.  Part  II. 

spots,  together  with  some  slight  increase  of  colour,  into  one  of  the 
so-called  elliptic  ornaments.  These  latter  ornaments  have  been 
shewn  to  many  persons,  and  all  have  admitted  that  they  are 
beautiful,  some  thinking  them  even  more  so  than  the  ball-and- 
socket  ocelli.  As  the  secondary  plumes  became  lengthened 
through  sexual  selection,  and  as  the  elhptic  ornaments  increased 
in  diameter,  their  colours  apparently  became  less  bright ;  and 
then  the  ornamentation  of  the  plumes  had  to  be  gained  by  an 
improvement  in  the  pattern  and  shading ;  and  this  process  was 
carried  on  until  the  wonderful  ball-and-socket  ocelli  were  finally  , 
developed.  Thus  we  can  understand— and  in  no  other  way  as 
it  seems  to  me — the  present  condition  and  origin  of  the  orna- 
ments on  the  wing-feathers  of  the  Argus  pheasant. 

From  the  light  afforded  by  the  principle  of  gradation — from 
what  we  know  of  the  laws  of  variation — from  the  changes  which 
have  taken  place  in  many  of  our  domesticated  birds — and,  lastly, 
from  the  character  (as  we  shall  hereafter  see  more  clearly)  of  the 
immature  plumage  of  young  birds — we  can  sometimes  indicate 
with  a  certain  amount  of  confidence,  the  probable  steps  by  which 
the  males  have  acquired  their  brilliant  plumage  and  various 
ornaments ;  yet  in  many  cases  we  are  involved  in  complete 
darkness.  Mr.  Gould  several  years  ago  pointed  out  to  me  a 
humming-bird,  the  Urosticte  henjamini,  remarkable  for  the  curious 
difierences  between  the  sexes.  The  male,  besides  a  splendid 
gorget,  has  greenish-black  tail-feathers,  with  the  four  central 
ones  tipped  with  white ;  in  the  female,  as  with  most  of  the  allied 
species,  the  three  outtr  tail-feathers  on  each  side  are  tipped  with 
white,  so  that  the  male  has  the  four  central,  whilst  the  female 
has  the  six  exterior  feathers  ornamented  with  white  tips.  What 
makes  the  case  more  curious  is  that,  although  the  coloui-ing  of 
the  tail  difi"ers  remarkably  in  both  sexes  of  many  kinds  of 
humming-birds,  Mr.  Gould  does  not  know  a  single  species,  be- 
sides the  Urosticte,  in  which  the  male  has  the  four  central 
feathers  tipped  with  w'hite. 

The  Duke  of  Argyll,  in  commenting  on  this  case,"^  passes  over 
sexual  selection,  and  asks,  "  What  explanation  does  the  law  of 
"  natural  selection  give  of  such  specific  varieties  as  these  ?"  He 
answers  ''  none  whatever ;"  and  I  quite  agree  with  him.  But 
can  this  be  so  confidently  said  of  sexual  selection  ?  Seeing  in 
how  many  ways  the  tail-feathers  of  humming-birds  differ,  why 
should  not  the  four  central  leathers  have  varied  in  this  one 
species  alone,  so  as  to  have  acquired  white  tips  ?  The  variations 
may  have  been  gTadual,  or  somewhat  abrupt  as  in  the  case 
43  '  The  Reign  of  Law,'  1867,  p.  247. 


Chap.  XIV,    Birds — Gradation  of  Characters.  443 

recently  given  of  the  humming-birds  near  Bogota,  in  which 
certain  individuals  alone  have  the  "  central  tail-feathers  tipped 
'*  with  beautiful  green."  In  the  female  of  the  Urosticto  I 
noticed  extremely  minute  or  rudimental  white  tips  to  the  two 
outer  of  the  four  central  black  tail-feathers ;  so  that  here  wo 
have  an  indication  of  change  of  some  kind  in  the  plumage  of 
this  species.  If  we  grant  the  possibility  of  the  central  tail- 
feathers  of  the  male  varying  in  wliiteness,  there  is  nothing  strange 
in  such  variations  having  been  sexually  selected.  The  white 
tips,  together  with  the  small  white  ear-tufts,  certainly  add,  as 
the  Duke  of  Argyll  admits,  to  the  beauty  of  the  male ;  and 
whiteness  is  apparently  appreciated  by  other  birds,  as  may  be 
inferred  from  such  cases  as  the  snow-white  male  of  the  Bell-bird. 
The  statement  made  by  Sir  E,  Heron  should  not  be  forgotten, 
namely,  that  his  peahens,  when  debarred  from  access  to  the  pied 
peacock,  would  not  unite  with  any  other  male,  and  during  that 
season  produced  no  offspring.  Nor  is  it  strange  that  variations 
in  the  tail-feathers  of  the  Urosticte  should  have  been  specially 
selected  for  the  sake  of  ornament,  for  the  next  succeeding  genus 
in  the  family  takes  its  name  of  Metallura  from  the  splendour  of 
these  feathers.  We  have,  moreover,  good  evidence  that  humming- 
birds take  especial  pains  in  displaying  their  tail-feathers ;  Mr. 
Belt,^*  after  describing  the  beauty  of  the  Florisuga  mellivora,  says, 
•'*  I  have  seen  the  female  sitting  on  a  branch,  and  two  males 
'^  displaying  their  charms  in  front  of  her.  One  would  shoot  up 
"  like  a  rocket,  then  suddenly  expanding  the  snow-white  tail,  like 
"  an  inverted  parachute,  slowly  descend  in  front  of  her,  turning 

"  round  gradually  to  shew  off  back  and  front The 

"  expanded  white  tail  covered  more  space  than  all  the  rest  of  the 
"  bird,  and  was  evidently  the  grand  feature  in  the  performance. 
"  Whilst  one  male  was  descending,  the  other  would  shoot  up 
"  and  come  slowly  down  expanded.  The  entertainment  would 
"  end  in  a  fight  between  the  two  performers ;  but  whether  the 
"  most  beautiful  or  the  most  pugnacious  was  the  accepted 
"  suitor,  I  know  not."  Mr.  Gould,  after  describing  the  peculiar 
plumage  of  the  Urosticte,  adds,  "  that  ornament  and  variety  is 
"  the  sole  object,  I  have  myself  but  little  doubt."  ^^  If  this  be 
admitted,  we  can  perceive  that  the  males  which  during  former 
times  were  decked  in  the  most  elegant  and  novel  manner  would 
have  gained  an  advantage,  not  in  the  ordinary  struggle  for  life, 
but  in  rivalry  with  other  males,  and  would  have  left  a  larger 
number  of  offspring  to  inherit  their  newly-acquired  beauty. 

^*  'The  Naturalist  in  Nicaragua,'  ^*  'Introduction    to    th«    Trochi- 

1874,  p.  112  lida,'  18G1,  p.  110. 


444  The  Descent  of  Mmi.  Part  II. 


CHAPTEK   XV. 

BIRDS — continued. 

Discussion  as  to  why  the  males  alone  of  some  species,  and  both  sexes  of 
others,  are  brightly  coloured — Oq  sexually-limited  inheritance,  as  applied 
to  various  structures  and  to  brightly-coloured  plumage — Nidification  in 
relation  to  colour — Loss  of  nuptial  plumage  during  the  winter. 

We  have  in  this  chapter  to  consider,  why  the  females  of  many 
birds  have  not  acquired  the  same  ornaments  as  the  male ;  and 
why,  on  the  other  hand,  both  sexes  of  many  other  birds  are 
equally,  or  almost  equally,  ornamented?  In  the  following 
chapter  we  shall  consider  the  few  cases  in  which  the  female  is 
more  conspicuously  coloured  than  the  male. 

In  my  'Origin  of  Species'^  I  briefly  suggested  that  the  long 
tail  of  the  peacock  would  be  inconvenient,  and  the  conspicuous 
black  colour  of  the  male  capercailzie  dangerous,  to  the  female 
during  the  period  of  incubation ;  and  consequently  that  the 
transmission  of  these  characters  from  the  male  to  the  female 
offspring  had  been  checked  through  natural  selection.  I  still 
think  that  this  may  have  occurred  in  some  few  instances :  but 
after  mature  reflection  on  all  the  facts  which  I  have  been  able  to 
collect,  I  am  now  inclined  to  believe  that  when  the  sexes  differ, 
the  successive  variations  have  generally  been  from  the  first 
limited  in  their  transmission  to  the  same  sex  in  which  they  first 
arose.  Since  my  remarks  appeared,  the  subject  of  sexual 
coloration  has  been  discussed  in  some  very  interesting  papers 
by  Mr.  Wallace,^  who  believes  that  in  almost  all  cases  the 
successive  variations  tended  at  first  to  be  transmitted  equally 
to  both  sexes;  but  that  the  female  was  saved,  through  natural 
selection,  from  acquiring  the  conspicuous  colours  of  the  male, 
owing  to  the  danger  which  she  would  thus  have  incurred  during 
incubation. 

This  view  necessitates  a  tedious  discussion  on  a  difficult 
point,  namely,  whether  the  transmission  of  a  character,  which  is 
at  first  inherited  by  both  sexes,  can  be  subsequently  limited  in 
its  transmission  to  one  sex  alone  by  means  of  natural  selection. 
We  must  bear  in  mind,  as  shewn  in  the  preliminary  chapter  on 
sexual  selection,  that  characters  which  are  limited  in  their 
development  to  one  sex  are  always  latent  in  the  other.     An 

1  Fourth  edition,  1866,  p.  241.  1867.     'Journal   of  Tiavel,'  vol.  i. 

*' Westminster    Review/    July,      1868,  p.  73. 


Chap.  XV.    Birds — Sexually  Limited  InJieritance.      445 


imaginary  illustration  will  best  aid  us  in  seeing  the  difficulty  of 
the  case :  we  may  suppose  that  a  fancier  wished  to  make  a 
breed  of  pigeons,  in  which  the  males  alone  should  be  coloured 
of  a  pale  blue,  whilst  the  females  retained  their  former  slaty 
tint.  As  with  pigeons  characters  of  all  kinds  are  usually  trans- 
mitted to  both  sexes  equally,  the  fancier  would  have  to  try  to 
convert  this  latter  form  of  inheritance  into  sexually-limited 
transmission.  All  that  he  could  do  would  be  to  persevere  in 
selecting  every  male  pigeon  which  was  in  the  least  degree  of  a 
paler  blue;  and  the  natural  result  of  this  process,  if  steadily 
carried  on  for  a  long  time,  and  if  the  pale  variations  were 
strongly  inherited  or  often  recurred,  would  be  to  make  his 
whole  stock  of  a  lighter  blue.  But  our  fancier  would  be  com- 
pelled to  match,  generation  after  generation,  his  pale  blue  males 
with  slaty  females,  for  he  wishes  to  keep  the  latter  of  this 
colour.  The  result  would  generally  be  the  production  either  of 
a  mongrel  piebald  lot,  or  more  probably  the  speedy  and 
complete  loss  of  the  pale-blue  tint;  for  the  primordial  slaty 
colour  would  be  transmitted  with  prepotent  force.  Supposing, 
however,  that  some  pale-blue  males  and  slaty  females  were 
produced  during  each  successive  generation,  and  were  always 
crossed  together ;  then  the  slaty  females  would  have,  if  I  may 
use  the  exi^ression,  much  blue  blood  in  their  veins,  for  their 
fathers,  grandfathers,  &c.,  will  all  have  been  blue  birds.  Under 
these  circumstances  it  is  conceivable  (though  I  know  of  no 
distinct  facts  rendering  it  probable)  that  the  slaty  females  might 
acquire  so  strong  a  latent  tendency  to  pale-blueness,  that  they 
would  not  destroy  this  colour  in  their  male  offspring,  their 
female  offsj)ring  still  inheriting  the  slaty  tint.  If  so  the  desired 
end  of  making  a  breed  with  the  two  sexes  permanently  different 
in  colour  might  be  gained. 

The  extreme  importance,  or  rather  necessity  in  the  above  case 
of  the  desired  character,  namely,  pale-blueness,  being  present 
though  in  a  latent  state  in  the  female,  so  that  the  male  offspring 
should  not  be  deteriorated,  will  be  best  ai^preciated  as  follows : 
the  male  of  Soemmerring's  pheasant  has  a  tail  thirty-seven 
inches  in  length,  whilst  that  of  the  female  is  only  eight  inches ; 
the  tail  of  the  male  common  pheasant  is  about  twenty  inches, 
and  that  of  the  female  twelve  inches  long.  Now  if  the  female 
Soemmerring  pheasant  with  her  short  tail  were  crossed  with  the 
male  common  pheasant,  there  can  be  no  doubt  that  the  male 
hybrid  offspring  would  have  a  much  lon<jer  tail  than  that  of  the 
pure  offspring  of  the  co.mmon  pheasant.  On  the  other  hand,  if 
the  female  common  pheasant,  with  a  tail  much  longer  than  that 
of  the  female  Soemmerring  pheasant,  were  crossed  with  the  male 


44^  The  Descent  of  Man.  Part  II. 

of  the  latter,  the  male  hybrid  o£Fspring  would  have  a  much 
shorter  tail  than  that  of  the  pure  offspring  of  Scemmerring's 
pheasant.  2 

Our  fancier,  in  order  to  make  his  new  breed  with  the  males 
of  a  pale-blue  tint,  and  the  females  unchanged,  would  have  to 
continue  selecting  the  males  during  many  generations;  and 
each  stage  of  paleness  would  have  to  be  fixed  in  the  males, 
and  rendered  latent  in  the  females.  The  task  would  be  an 
extremely  difficult  one,  and  has  never  been  tried,  but  might 
possibly  be  successfully  carried  out.  The  chief  obstacle  would 
be  the  early  and  complete  loss  of  the  pale-blue  tint,  from  the 
necessity  of  reiterated  crosses  with  the  slaty  female,  the  latter 
not  having  at  first  any  latent  tendency  to  produce  pale-blue 
offspring. 

On  the  other  hand,  if  one  or  two  males  were  to  vary  ever  so 
slightly  in  paleness,  and  the  variations  were  from  the  first 
limited  in  their  transmission  to  the  male  sex,  the  task  of  making 
a  new  breed  of  the  desired  kind  would  be  easy,  for  such  males 
would  simply  have  to  be  selected  and  matched  with  ordinary 
females.  An  analogous  case  has  actually  occurred,  for  there  are 
breeds  of  the  pigeon  in  Belgium  ^  in  which  the  males  alone  are 
marked  with  black  striae.  So  again  Mr.  Tegetmeier  has  recently 
shewn  ^  that  dragons  not  rarely  produce  silver-coloured  birds, 
which  are  almost  always  hens;  and  he  himself  has  bred  ten 
such  females.  It  is  on  the  other  hand  a  very  unusual  event 
when  a  silver  male  is  produced;  so  that  nothing  would  be 
easier,  if  desired,  than  to  make  a  breed  of  dragons  with  blue 
males  and  silver  females.  This  tendency  is  indeed  so  strong 
that  when  Mr.  Tegetmeier  at  last  got  a  silver  male  and  matched 
him  with  one  of  the  silver  females,  he  expected  to  get  a  breed 
with  both  sexes  thus  coloured;  he  was  however  disappointed, 
for  the  young  male  reverted  to  the  blue  colour  of  his  grand- 
father, the  young  female  alone  being  silver.  No  doubt  with 
patience  this  tendency  to  reversion  in  the  males,  reared  from  an 
occasional  silver  male  matched  with  a  silver  hen,  might  be 
eliminated,  and  then  both  sexes  would  be  coloured  alike ;  and 
this  very  process  has  been  followed  with  success  by  Mr. 
Esquilant  in  the  case  of  silver  turbits. 

With  fowls,  variations  of  colour,  limited  in  their  transmis- 
sion to  the  male  sex,  habitually  occur.     When  this  form  of 

2  Temminck  says  that  the  tail  of  For     the     common     pheasant,     see 

the  female  Phasianus  Sccramerringii  Macgillivray,    '  Hist.    Brit.     Birds,' 

is   only    six   inches  long,  'Planches  vol.  i.  pp.  118-121. 

coloriees,'  vol.  v.  1838,  pp.  487  and  *  Dr.  Chapuis,  '  Le  Pigeon  Voya- 

488 :  the  measurements  above  given  gear  Beige,'  1865,  p.  87. 

were  irade  for  me  by  Mr.  ScLater.  *  The  'Field,'  Sept.  1872. 


Chap.  XV.    Birds — Sexually  Limited  Inheritance,      447 


inheritance  prevails,  it  might  well  happen  that  some  of  the 
successive  variations  would  be  transferred  to  the  female,  who 
would  then  slightly  resemble  the  male,  as  actually  occurs  in 
some  breeds.  Or  again,  the  greater  number,  but  not  all,  of  the 
successive  steps  might  be  transferred  to  both  sexes,  and  the 
female  would  then  closely  resemble  the  male.  There  can  hardly 
be  a  doubt  that  this  is  the  cause  of  the  male  pouter  pigeon 
having  a  somewhat  larger  crop,  and  of  the  male  carrier  pigeon 
having  somewhat  larger  wattles,  than  their  respective  females ; 
for  fanciers  have  not  selected  one  sex  more  than  the  other,  and 
have  had  no  wish  that  these  characters  should  be  more  strongly 
displayed  in  the  male  than  in  the"  female,  yet  this  is  the  case 
with  both  breeds. 

The  same  process  would  have  to  be  followed,  and  the  same 
difficulties  encountered,  if  it  were  desired  to  make  a  breed  with 
the  females  alone  of  some  new  colour. 

Lastly,  our  fancier  might  wish  to  make  a  breed  with  the  two 
sexes  differing  from  each  other,  and  both  from  the  parent- 
species.  Here  the  difficulty  would  be  extreme,  unless  the  suc- 
cessive variations  were  from  the  first  sexually  limited  on  both 
sides,  and  then  there  would  be  no  difficulty.  We  see  this  with 
the  fowl ;  thus  the  two  sexes  of  the  pencilled  Hamburghs  differ 
greatly  from  each  other,  and  from  the  two  sexes  of  the  abori- 
ginal Gallus  hankiva ;  and  both  are  now  kept  constant  to  their 
standard  of  excellence  by  continued  selection,  which  would  be 
impossible  unless  the  distinctive  characters  of  both  were  limited 
in  their  transmission.  The  Spanish  fowl  offers  a  more  curious 
case ;  the  male  has  an  immense  comb,  but  some  of  the  successive 
variations,  by  the  accumulation  of  which  it  was  acquired,  appear 
to  have  been  transferred  to  the  female ;  for  she  has  a  comb  many 
times  larger  than  that  of  the  females  of  the  parent-species.  But 
the  comb  of  the  female  differs  in  one  respect  from  that  of  the 
male,  for  it  is  apt  to  lop  over ;  and  within  a  recent  period  it  has 
been  ordered  by  the  fancy  that  this  should  always  be  the  case, 
and  success  has  quickly  followed  the  order.  Now  the  lopping  of 
the  comb  must  be  sexually  limited  in  its  transmission,  otherwise 
it  would  prevent  the  comb  of  the  male  from  being  perfectly 
upright,  which  would  be  abhorrent  to.  every  fancier.  On  the 
other  hand,  the  uprightness  of  the  comb  in  the  male  must 
likewise  be  a  sexually-limited  character,  otherwise  it  would 
prevent  the  comb  of  the  female  from  lopping  over. 

From  the  foregoing  illustrations,  we  see  that  even  with  almost 
unlimited  time  at  command,  it  would  be  an  extremely  difficult 
and  complex,  perhaps  an  impossible  process,  to  change  one  form 
of  transmission  into  the  other  through  selection.     Therefore, 


448  The  Descent  of  Man.  Part  II. 

without  distinct  evidence  in  each  case,  I  am  unwiUing  to  admit 
that  this  has  been  effected  in  natural  species.  On  the  other 
hand,  by  means  of  successive  variations,  which  were  from  the 
first  sexually  limited  in  their  transmission,  there  would  not  be 
the  least  difficulty  in  rendering  a  male  bird  widely  different  in 
colour  or  in  any  other  character  from  the  female ; "  the  latter 
being  left  unaltered,  or  shghtly  altered,  or  specially  modified  for 
the  sake  of  protection. 

As  bright  colours  are  of  service  to  the  males  in  their  rivalry 
with  other  males,  such  colours  would  be  selected,  whether  or 
not  they  were  transmitted  exclusively  to  the  same  sex.  Conse- 
quently the  females  might  be  expected  often  to  partake  of  the 
brightness  of  the  males  to  a  greater  or  less  degree;  and  this 
occurs  with  a  host  of  species.  If  all  the  successive  variations 
were  transmitted  equally  to  both  sexes,  the  females  would  be 
indistinguishable  from  the  males ;  and  this  likewise  occurs  with 
many  birds.  If,  however,  dull  colours  were  of  high  importance 
for  the  safety  of  the  female  during  incubation,  as  with  many 
ground  birds,  the  females  which  varied  in  brightness,  or  which 
received  through  inheritance  from  the  males  any  marked  ac- 
cession of  brightness,  would  sooner  or  later  be  destroyed.  But 
the  tendency  in  the  males  to  continue  for  an  indefinite  period 
transmitting  to  their  female  offspring  their  own  brightness, 
would  have  to  be  eliminated  by  a  change  in  the  form  of  in- 
lieritance ;  and  this,  as  shewn  by  our  jDrevious  illustration, 
would  be  extremely  difficult.  The  more  probable  result  of  the 
long-continued  destruction  of  the  more  brightly-coloured  females, 
supposing  the  equal  form  of  transmission  to  jDrevail,  would  be 
the  lessening  or  annihilation  of  the  bright  colours  of  the  males, 
owing  to  their  continual  crossing  with  the  duller  females.  It 
would  be  tedious  to  follow  out  all  the  other  possible  results; 
but  I  may  remind  the  reader  that  if  sexually-limited  variations 
in  brightness  occurred  in  the  females,  even  if  they  were  not  in 
the  least  injurious  to  them  and  consequently  were  not  elimi- 
nated, yet  they  would  not  be  favoured  or  selected,  for  the  male 
usually  accepts  any  female,  and  does  not  select  the  more  at- 
tractive individuals ;  consequently  these  variations  would  be 
liable  to  be  lost,  and  would  have  little  influence  on  the  character 
of  the  race ;  and  this  will  aid  in  accounting  for  the  females  being 
commonly  duller-coloured  than  the  males. 

In  the  eighth  chapter  instances  were  given,  to  which  many 
might  here  be  added,  of  variations  occurring  at  various  ages,  and 
inherited  at  the  corresponding  age.  It  was  also  shewn  that  va- 
riations which  occur  late  in  life  are  commonly  transmitted  to  the 
same  sex  in  which  they  first  appear ;  whilst  variations  occurring 


Chap.  XV.       Birds — Developmejit  of  Spurs.  449 

early  in  life  are  apt  to  be  transmitted  to  both  sexes ;  not  that  all 
the  cases  of  sexually-limited  transmission  can  thns  be  accounted 
for.  It  was  further  shewn  that  if  a  male  bird  varied  by  be- 
coming brighter  whilst  young,  such  variations  would  be  of  no 
service  until  the  age  for  reproduction  had  arrived,  and  there 
was  competition  between  rival  males.  But  in  the  case  of  birds 
living  on  the  ground  and  commonly  in  need  of  the  protection  of 
dull  colours,  bright  tints  would  *  be  far  more  dangerous  to  the 
youDg  and  inexperienced,  than  to  the  adult  males.  Conse- 
quently the  males  which  varied  in  brightness  whilst  young 
would  suffer  much  destruction  and  .be  eliminated  through 
natural  selection ;  on  the  other  hand,  the  males  which  varied  in 
this  manner  when  nearly  mature,  notwithstanding  that  they 
were  exposed  to  some  additional  danger,  might  survive,  and 
from  being  favoured  through  sexual  selection,  would  procreate 
their  kind.  As  a  relation  often  exists  between  the  period  of 
variation  and  the  form  of  transmission,  if  the  bright-coloured 
young  males  were  destroyed  and  the  mature  ones  were  suc- 
cessful in  their  courtship,  the  males  alone  would  acquire  bril- 
liant colours  and  would  transmit  them  exclusively  to  their 
male  offspring.  But  I  by  no  means  wish  to  maintain  that  the 
influence  of  age  on  the  form  of  transmission,  is  the  sole  cause  of 
the  great  difference  in  brilliancy  between  the  sexes  of  many 
birds. 

When  the  sexes  of  birds  differ  in  colour,  it  is  interesting  to 
determine  whether  the  males  alone  have  been  modified  by  sexual 
selection,  the  females  having  been  left  unchanged,  or  only  partially 
and  indirectly  thus  changed ;  or  whether  the  females  have  been 
specially  modified  through  natural  selection  for  the  sake  of  pro- 
tection. I  will  therefore  discuss  this  question  at  some  length, 
even  more  fully  than  its  intrinsic  importance  deserves ;  for  various 
curious  collateral  points  may  thus  be  conveniently  considered. 

Before  we  enter  on  the  subject  of  colour,  more  especially  in 
reference  to  Mr.  Wallace's  conclusions,  it  may  be  useful  to 
discuss  some  other  sexual  differences  under  a  similar  point  of 
view.  A  breed  of  fowls  formerly  existed  in  Germany*^  in  which 
the  hens  were  furnished  with  spurs ;  they  were  good  layers,  but 
they  so  greatly  disturbed  their  nests  with  their  spurs  that  they 
could  not  be  allowed  to  sit  on  their  own  eggs.  Hence  at  one  time 
it  appeared  to  me  probable  that  with  the  females  of  the  wild 
GiillinacesB  the  development  of  spurs  had  been  checked  through 
natural  selection,  from  the  injury  thus  caused  to  their  nests. 
This  seemed  all  the  more  probable,  as  wing-spurs,  which  would 
not  be  injurious  during  incubation,  are  often  as  well-developed 
6  Bechstein,   '  Natur^esch.  Deutschlands,'  1793,  B.  iii.  s.  339. 


45 o  The  Descent  of  Man.  Part  II. 


in  tlie  female  as  in  the  male ;  though  in  not  a  few  cases  they  are 
rather  larger  in  the  male.  When  the  male  is  furnished  with 
leg-spurs  the  female  almost  always  exhibits  rudiments  of  them, 
— the  rudiment  sometimes  consisting  of  a  mere  scale,  as  in 
G alius.  Hence  it  might  be  argued  that  the  females  had  abori- 
ginally been  furnished  with  well-developed  spurs,  but  that  these 
had  subsequently  been  lost  through  disuse  or  natural  selection. 
But  if  this  view  be  admitted,  it  would  have  to  be  extended  to 
innumerable  other  cases;  and  it  implies  that  the  female  pro- 
genitors of  the  existing  spur-bearing  sjDecies  were  once  encum- 
bered with  an  injurious  appendage. 

In  some  few  genera  and  sjDecies,  as  in  Galloperdix,  Acomus, 
and  the  Javan  peacock  {Pavo  muticas),  the  females,  as  well  as 
the  males,  possess  well-developed  leg- spurs.  Are  we  to  infer  from 
this  fact,  that  they  construct  a  different  sort  of  nest  from  that 
made  by  their  nearest  allies,  and  not  liable  to  be  injured  by  their 
spurs  ;  so  that  the  spurs  have  not  been  removed.  Or  are  we  to 
suppose  that  the  females  of  these  several  species  especially  require 
spurs  for  their  defence  ?  It  is  a  more  probable  conclusion  that 
both  the  presence  and  absence  of  spurs  in  the  females  result 
from  different  laws  of  inheritance  having  prevailed,  indepen- 
dently of  natural  selection.  With  the  many  females  in  which 
spurs  appear  as  rudiments,  we  may  conclude  that  some  few  of 
the  successive  variations,  through  which  they  were  developed  in 
the  males,  occurred  very  early  in  life,  and  were  consequently 
transferred  to  the  females.  In  the  other  and  much  rarer  cases, 
in  which  the  females  possess  fully  developed  spurs,  we  may 
conclude  that  all  the  successive  variations  were  transferred  to 
them ;  and  that  they  gradually  acquired  and  inherited  the  habit 
of  not  disturbing  their  nests. 

The  vocal  organs  and  the  feathers  variously-modified  for  pro- 
ducing sound,  as  well  as  the  proper  instincts  for  using  them, 
often  differ  in  the  two  sexes,  but  are  sometimes  the  same  in  both. 
Can  such  differences  be  accounted  for  by  the  males  having 
acquired  these  organs  and  instincts,  whilst  the  females  have 
been  saved  from  inheriting  them,  on  account  of  the  danger  to 
which  they  would  have  been  exposed  by  attracting  the  attention 
of  birds  or  beasts  of  prey  ?  This  does  not  seem  to  me  probable, 
when  we  think  of  the  multitude  of  birds  which  with  impunity 
gladden  the  country  with  their  voices  during  the  spring.'^    It  is 

"^  Daines     Barrington,     however,  cubation.     He  adds,  that  a  similar 

thought  it  probable  ('Phil.  Transact.'  view  may  possibly  account  for  the 

1773,  p.  164)  that  few  female  birds  inferiority    of    the    female    to     the 

sing,  because  the  talent  would  have  male  in  plumage, 
been  dauo-erous  to  them  during  in- 


CiiAP.  XV.       Bii'ds — Length  of  Female  s  Tail.  45 1 


a  safer  conclusion  that,  as  vocal  and  instrumental  organs  are  of 
special  service  only  to  the  males  during  their  courtship,  these 
organs  were  developed  through  sexual  selection  and  their  con- 
stant use  in  that  sex  alone— the  successive  variations  and  the 
effects  of  use  having  been  from  the  first  more  or  less  limited  in 
transmission  to  the  male  offspring. 

Many  analogous  cases  could  be  adduced  ;  those  for  instance  of 
the  plumes  on  the  head  being  generally  longer  in  the  male  than 
in  the  female,  sometimes  of  equal  length  in  both  sexes,  and 
occasionally  absent  in  the  female, — these  several  cases  occuring 
in  the  same  group  of  birds.  It  would  be  difficult  to  account  for 
such  a  difference  between  the  sexes  by  the  female  having  been 
benefited  by  possessing  a  slightly  shorter  crest  than  the  male, 
and  its  consequent  diminution  or  comj)lete  suppression  through 
natural  selection.  But  I  will  take  a  more  favourable  case, 
namely  the  length  of  the  tail.  The  long  train  of  the  peacock 
would  have  been  not  only  inconvenient  but  dangerous  to  the 
peahen  during  the  period  of  incubation  and  whilst  accompany- 
ing her  young.  Hence  there  is  not  the  least  a  'priori  improba- 
bility in  the  development  of  her  tail  having  been  checked 
through  natural  selection.  But  the  females  of  various  phea- 
sants, which  ai3parently  are  exposed  on  their  open  nests  to  as 
much  danger  as  the  peahen,  have  tails  of  considerable  length. 
The  females  as  well  as  the  males  of  the  Menura  superha  have 
long  tails,  and  they  build  a  domed  nest,  which  is  a  great 
anomaly  in  so  large  a  bird.  Naturalists  have  wondered  how  the 
female  Menura  could  manage  her  tail  during  incubation  ;  but  it 
is  now  known  ^  that  she  "  enters  the  nest  head  first,  and  then 
"  turns  round  with  her  tail  sometimes  over  her  back,  but  more 
"  often  bent  round  by  her  side.  Thus  in  time  the  tail  becomes 
"  quite  askew,  and  is  a  tolerable  guide  to  the  length  of  time  the 
"  bird  has  been  sitting."  Both  sexes  of  an  Australian  kingfisher 
{Tanysiptera  sylvia)  have  the  middle  tail-feathers  greatly  length- 
ened, and  the  female  makes  her  nest  in  a  hole ;  and  as  I  am 
informed  by  Mr.  E.  B.  Sharpe  these  feathers  become  much 
crumpled  during  incubation. 

In  these  two  latter  cases  the  great  length  of  the  tail-feathers 
must  be  in  some  degree  inconvenient  to  the  female  ;  and  as  in  both 
species  the  tail-feathers  of  the  female  are  somewhat  shorter  than 
those  of  the  male,  it  might  be  argued  that  their  full  development 
had  been  prevented  through  natural  selection.  But  if  the 
development  of  the  tail  of  the  peahen  had  been  checked  only 
when  it  became  inconveniently  or  dangerously  great,  she  would 
have  retained  a  much  longer  tail  than  she  actually  possesses ; 
»  Mr.  Ramsay,  in  '  Proc.  Zoolog.  Soc'  1868,  p.  50. 


452  The  Descent  of  Man.  Part  11. 

for  her  tail  is  not  nearly  so  long,  relatively  to  tlie  size  of  her 
body,  as  that  of  many  female  pheasants,  nor  longer  than  that 
of  the  female  turkey.  It  must  also  be  borne  in  mind,  that  in 
accordance  with  this  view  as  soon  as  the  tail  of  the  peahen 
became  dangerously  loflg,  and  its  development  was  consequently 
checked,  she  would  have  continually  reacted  on  her  male  pro- 
geny, and  thus  have  prevented  the  peacock  from  acquiring  his 
present  magnificent  train.  We  may  therefore  infer  that  the 
length  of  the  tail  in  the  peacock  and  its  shortness  in  the  pea- 
hen are  the  result  of  the  requisite  variations  in  the  male  having 
been  from  the  first  transmitted  to  the  male  offspring  alone. 

We  are  led  to  a  nearly  similar  conclusion  with  respect  to  the 
length  of  the  tail  in  the  various  species  of  pheasants.  In  the  Eared 
pheasant  {Croswptilon  auritum)  the  tail  is  of  equal  length  in  both 
sexes,  namely,  sixteen  or  seventeen  inches ;  in  the  common  phea- 
sant it  is  about  twenty  inches  long  in  the  male  and  twelve  in  the 
female ;  in  Soemmerring's  pheasant,  thirty-seven  inches  in  the 
male  and  only  eight  in  the  female ;  and  lastly  in  Eeeve's  phea- 
sant it  is  sometimes  actually  seventy-two  inches  long  in  the  male 
and  sixteen  in  the  female.  Thus  in  the  several  species,  the  tail  of 
the  female  differs  much  in  length,  irrespectively  of  that  of  the 
male ;  and  this  can  be  accounted  for,  as  it  seems  to  me,  with 
much  more  probability,  by  the  laws  of  inheritance, — that  is  by 
the  successive  variations  having  been  from  the  first  more  or  less 
closely  limited  in  their  transmission  to  the  male  sex,  than  by  the 
agency  of  natural  selection,  resulting  from  the  length  of  tail  being 
more  or  less  injurious  to  the  females  of  these  several  allied  species. 

We  may  now  consider  Mr.  Wallace's  arguments  in  regard  to 
the  sexual  coloration  of  birds.  He  believes  that  the  bright  tints 
originally  acquired  through  sexual  selection  by  the  males,  would 
in  all,  or  almost  all  cases,  have  been  transmitted  to  the  females, 
unless  the  transference  had  been  checked  through  natural  selec- 
tion. I  may  here  remind  the  reader  that  various  facts  opposed 
to  this  view  have  already  been  given  under  reptiles,  amphibians, 
fishes,  and  lepidoptera.  Mr.  Wallace  rests  his  belief  chiefly, 
but  not  exclusively,  as  we  shall  see  in  the  next  chapter, 
on  the  following  statement,^  that  when  both  sexes  are  coloured 
in  a  very  conspicuous  manner,  the  nest  is  of  such  a  nature 
as  to  conceal  the  sitting  bird;  but  when  there  is  a  marked 
contrast  of  colour  between  the  sexes,  the  male  being  gay  and 
the  female  dull-coloured,  the  nest  is  ojien  and  exposes  the 
sitting  bird  to  view.  This  coincidence,  as  far  as  it  goes,  cer- 
tainly seems  to  favour  the  belief  that  the  females  which  sit  on 
8  'Journal  of  Travel,'  edited  by  A.  Murray,  vol.  i.  1868,  p.  78. 


Chap*  XV.      Birds — Colour  and  Nidijicaiion,  453 

open  nests  have  been  specially  modified  for  the  sake  of  protec- 
tion ;  but  we  shall  presently  see  that  there  is  anotlier  and  more 
probable  explanation,  namely,  that  conspicuous  females  have 
acquired  the  instinct  of  building  domed  nests  oftener  than  dull- 
coloured  birds.  Mr.  Wallace  admits  that  there  are,  as  might 
have  been  expected,  some  exceptions  to  his  two  rules,  but  it  is  a 
question  whether  the  exceptions  are  not  so  numerous  as  seriously 
to  invalidate  them. 

There  is  in  the  first  place  much  truth  in  the  Duke  of  Argyll's 
remark  ^^  that  a  large  domed  nest  is  more  conspicuous  to  an 
enemy,  especially  to  all  tree-haunting  carnivorous  animals,  than 
a  smaller  open  nest.  Nor  must  we  forget  that  with  many  birds 
which  build  open  nests,  the  male  sits  on  the  eggs  and  aids  the 
female  in  feeding  the  young:  this  is  the  case,  for  instance,  with 
Pyranga  cvstivt,'^^  one  of  the  most  splendid  birds  in  the  United 
States,  the  male  being  vermilion,  and  the  female  light  brownish- 
green.  Now  if  brilliant  colours  had  been  extremely  dangerous 
to  birds  whilst  sitting  on  their  open  nests,  the  males  in  these 
cases  would  have  suffered  greatly.  It  might,  however,  be  of 
such  paramount  importance  to  the  male  to  be  brilliantly  coloured, 
in  order  to  beat  his  rivals,  that  this  may  have  more  than  com- 
pensated some  additional  danger. 

Mr.  Wallace  admits  that  with  the  King-crows  (Dicrurus), 
Orioles,  and  Pittidse,  the  females  are  conspicuously,  coloured, 
yet  build  open  nests ;  but  he  urges  that  the  birds  of  the  first 
group  are  highly  pugnacious  and  could  defend  themselves ;  that 
those  of  the  second  group  take  extreme  care  in  concealing  their 
open  nests,'but  this  does  not  invariably  hold  good ;  ^^  and  that 
with  the  birds  of  the  third  group  the  females  are  brightly 
coloured  chiefly  on  the  under  surface.  Besides  these  cases, 
pigeons  which  are  sometimes  brightly,  and  almost  always  con- 
spicuously coloured,  and  which  are  notoriously  liable  to  the 
attacks  of  birds  of  prey,  offer  a  serious  exception  to  the  rule,  for 
they  almost  always  build  open  and  exposed  nests.  In  another 
large  family,  that  of  the  humming-birds,  all  the  species  build 
open  nests,  yet  with  some  of  the  most  gorgeous  species  the  sexes 
are  alike ;  and  in  the  majority,  the  females,  though  less  brilliant 
than  the  males,  are  brightly  coloured.  Nor  can  it  be  maintained 
that  all  female  humming-birds,  which  are  brightly  coloured, 
escape  detection  by  their  tints  being  green,  for  some  display  on 
their  upper  surfaces  red,  blue,  and  other  colours.^^ 

'»  'Journal   of  Travel,' edited  bv  '^  jenlon,   'Birds   of  India,'  vol. 

A.  Murray,  vol.  i.  1868,  p.  281.       '  ii.    p.    108.     Gould's   'Handbook    of 

'•   Audubon,  '  Ornitnological  Bio-  the  Birds  of  Australia,'  vol.  i.  p.  463. 

graphy,'  vol.  i.  p.  233.  i3  y^^  instance,  the  female  Eupe- 


454  '^^^^  Descent  of  Man.  Tart  II. 

In  regard  to  birds  which  build  in  holes  or  construct  domed 
nests,  other  advantages,  as  Mr.  Wallace  remarks,  besides  con- 
cealment are  gained,  such  as  shelter  from  the  rain,  greater 
warmth,  and  in  hot  countries  protection  from  the  sun;^*  so  that 
it  is  no  valid  objection  to  his  view  that  many  birds  having  both 
sexes  obscurely  coloured  build  concealed  nests.^^  The  female 
Horn-bill  {Buceros),  for  instance,  of  India  and  Africa  is  protected 
during  incubation  with  extraordinary  care,  for  she  plasters  up 
with  her  own  excrement  the  orifice  of  the  hole  in  which  she  sits 
on  her  eggs,  leaving  only  a  small  orifice  through  which  the  male 
feeds  her ;  she  is  thus  kept  a  close  prisoner  during  the  whole 
period  of  incubation ;  '^^  yet  female  horn-bills  are  not  more  con- 
spicuously coloured  than  many  other  birds  of  equal  size  which 
build  open  nests.  It  is  a  more  serious  objection  to  Mr,  Wallace's 
view,  as  is  admitted  by  him,  that  in  some  few  groups  the  males 
are  brilliantly  coloured  and  the  females  obscure,  and  yet  the 
latter  hatch  their  eggs  in  domed  nests.  This  is  the  case  with  the 
Grallinae  of  Australia,  the  Superb  Warblers  (Maluridse)  of  the 
same  country,  the  Sun-birds  (Nectariniae),  and  with  several  of 
the  Australian  Honey-suckers  or  Meliphagidse.^'^ 

If  we  look  to  the  birds  of  England  we  shall  see  that  there  is  no 
close  and  general  relation  between  the  colours  of  the  female  and 
the  nature  of  the  nest  which  is  constructed.  About  forty  of  our 
British  birds  (excluding  those  of  large  size  which  could  defend 
themselves)  build  in  holes  in  banks,  rocks,  or  trees,  or  construct 
domed  nests.  If  we  take  the  colours  of  the  female  goldfinch, 
bullfinch,  or  blackbird,  as  a  standard  of  the  degree  of  con- 
spicuousness,  which  is  not  highly  dangerous  to  the  sitting 
female,  then  out  of  the  above  forty  birds,  the  females  of  only 
twelve  can  be  considered  as  conspicuous  to  a  dangerous  degree. 


tomena  macroura  has  the  head  and  very  hot  weather,  when  the  sun 
tail  dark  blue  with  reddish  loins;  was  shining  brightly,  as  if  their 
the  female  Lampornis  porphyrurus  eggs  would  be  thus  injured,  than 
is  blackish-green  on  the  upper  duringcool,  cloudy,  or  rainy  weather, 
surface,  with  the  lores  and  sides  of  *^  I  may  specify,  as  instances  of 
the  throat  crimson;  the  female  dull-coloured  birds  building  con- 
Eulampiis  jugularis  has  the  top  of  cealed  nests,  the  species  belonging 
the  head  and  back  green,  but  the  to  eight  Australian  genera,  de- 
loins  and  the  tail  are  crimson,  scribed  in  Gould's  '  Handbook  of  the 
Many  other  instances  of  highly  Birds  of  Australia,'  vol.  i.  pp.  340, 
conspicuous  females  could  be  given.  362,  365,  383,  387,  389,  391,  414. 
See  Mr.  Gould's  magnificent  work  "^  Mr.  C.  Home,  '  Proc.  Zoolog. 
on  this  family.  Soo.'  1869,  p.  243. 

**  Mr.  Salvin    noticed  in  Guate-  '^  On  the  nidification  and  colours 

mala   ('Ibis,'    1864,    p.     375)   that  of  these  latter  species,  see  Gould's 

humming-birds    were     much    more  '  Handbook,' &c.,  vol.  i.  pp.  504,  527. 
unwilliuo:  to  leave  their  nests  during 


Chap.  XV.      Birds — Colour  and  Nidification.  455 


the  remaining  twenty-eight  being  inconspicuous."  Nor  is  tlierc 
any  close  relation  within  the  same  genus  between  a  well-pro- 
nounced difference  in  colour  between  the  sexes,  and  the  nature 
ol' the  nest  constructed.  Thus  the  male  house  sparrow  (Passf^r 
domesticus)  differs  much  from  the  female,  the  male  tree-sparrow 
{P.  montanns)  hardly  at  all,  and  yet  both  build  well-concealed 
nests.  The  two  sexes  of  the  common  fly-catcher  (Musicajm 
grisold)  can  hardly  be  distinguished,  whilst  the  sexes  of  the 
pied  fly-catcher  (ilf.  lactuosa)  differ  considerably,  and  both 
species  build  in  holes  or  conceal  their  nests.  The  female  black- 
bird (Tardus  mtrida)  differs  much,  the  female  ring-ouzel  \T. 
torquatus)  differs  less,  and  the  female  common  thrush  {T.  musicus) 
hardly  at  all  from  their  respective  males ;  yet  all  build  open  nests. 
On  the  other  hand,  the  not  very  distantly-allied  water-ouzel 
{Cindus  aquaticm)  builds  a  domed  nest,  and  the  sexes  differ 
about  as  much  as  in  the  ring-ouzel.  The  black  and  red  grouse 
{Tetrao  tdrix  and  T.  scoticus)  build  open  nests  in  equally  well- 
concealed  spots,  but  in  the  one  species  the  sexes  differ  greatly, 
and  in  the  other  very  little. 

Notwithstanding  the  foregoing  objections,  I  cannot  doubt, 
after  reading  Mr.  Wallace's  excellent  essay,  that  looking  to  the 
birds  of  the  world,  a  large  majority  of  the  species  in  which  the 
females  are  conspicuously  coloured  (and  in  this  case  the  males 
with  rare  excej^tions  are  equally  conspicuous),  build  concealed 
nests  for  the  sake  of  protection.  Mr.  Wallace  enumerates^''  a 
long  series  of  groups  in  which  this  rule  holds  good ;  but  it  will 
suffice  here  to  give,  as  instances,  the  more  familiar  groups  of 
kingfishers,  toucans,  trogons,  puff- birds  (Capitonidae),  plantain- 
eaters  (Musophaga^),  woodpeckers,  and  parrots.  Mr.  Wallace 
believes  that  in  these  groups,  as  the  males  gradually  acquired 
through  sexual  selection  their  brilliant  colours,  these  were 
transferred  to  the  females  and  were  not  eliminated  by  natural 
selection,  owing  to  the  protection  which  they  already  enjoyed 

'*  I  have  consulted,  on  this  sub-  boarula   (?) ;    Erithacus   (?)  ;     Fru- 

ject,  Macgillivray's  '  British   Birds,'  ticola,  2  sp. ;  Saxicola  ;  Ruticilla,  2 

and  though   doubts  may  be   enter-  sp. ;    Sylvia,    3   sp. ;  Parus,   3  sp. ; 

tained    in  some  cases  in  regard   to  Mecistura ;     Anorthura ;      Certhia ; 

the   degree   of  concealment  of  the  Sitta ;     Yunx ;    Muscicapa,    2    sp. ; 

nest,    and    to    the    degree    of    con-  Hirundo,  3  sp. ;  and  Cypselus.     The 

spicuousness  of  the  female,  yet  the  females    of    the  following  12  birds 

following  birds,  which  all  lay  their  may  be    considered    as   conspicuous, 

e^gs  in  holes  or  in  domed  nests,  can  according     to     the     same  standard, 

hardly  be  considered,  by  the  above  viz.,   Pastor,   Motacilla  alba,  Parus 

standard,    as     conspicuous:     Passer,  major  and  P.  cseruleus,  Upupa,  Picus, 

2  species;  Sturnus,  of  which  the  4  sp.,  Coracias,  Alcedo,  and  Merops. 
female  is  considerably  less  brilliant  '^  '  Journal  of  Travel,'  edited  by 

than  the  male;  Cinclus  ;  Motacilla  A.  Murray,  vol.  i.  p.  78. 


456  The  Descent  of  Man.  Part  II. 

from  their  manner  of  nidification.  According  to  this  view,  their 
present  manner  of  nesting  was  acquired  before  their  present 
colours.  But  it  seems  to  me  much  more  probable  that  in  most 
cases,  as  the  females  were  gradually  rendered  more  and  more 
brilliant  from  partaking  of  the  colours  of  the  male,  they  were 
gradually  led  to  change  their  instincts  (supposing  that  they 
originally  built  open  nests),  and  to  seek  protection  by  building 
domed  or  concealed  nests.  No  one  who  studies,  or  mstance, 
Audubon's  account  of  the  differences  in  the  nests  of  the  same 
species  in  the  Northern  and  Southern  United  States,""  will  feel 
any  great  ditficulty  in  admitting  that  birds,  either  by  a  change 
(in  the  strict  sense  of  the  word)  of  their  habits,  or  through  the 
natural  selection  of  so-called  spontaneous  variations  of  instinct, 
might  readily  be  led  to  modify  their  manner  of  nesting. 

This  way  of  viewing  the  relation,  as  far  as  it  holds  good, 
between  the  bright  colours  of  female  birds  and  their  manner  of 
nesting,  receives  some  support  from  certain  cases  occurring  in 
the  Sahara  Desert.  Here,  as  in  most  other  deserts,  various  birds, 
and  many  other  animals,  have  had  their  colours  adapted  in  a 
wonderful  manner  to  the  tints  of  the  surrounding  surface. 
Nevertheless  there  are,  as  I  am  informed  by  the  Eev.  Mr. 
Tristram,  some  curious  exceptions  to  the  rule  ;  thus  the  male  of 
the  Monticola  cyanea  is  conspicuous  from  his  bright  blue  colour, 
and  the  female  almost  equally  conspicuous  from  her  mottled 
brown  and  white  plumage;  both  sexes  of  two  species  of  Dro- 
molsea  are  of  a  lustrous  black ;  so  that  these  three  species  are  far 
from  receiving  protection  from  their  colours,  yet  they  are  able  to 
survive,  for  they  have  acquired  the  habit  of  taking  refuge  from 
danger  in  holes  or  crevices  in  the  rocks. 

With  respect  to  the  above  groups  in  which  the  females  are 
conspicuously  coloured  and  build  concealed  nests,  it  is  not 
necessary  to  suppose  that  each  separate  species  had  its  nidifying 
instinct  specially  modified ;  but  only  that  the  early  progenitors 
of  each  group  were  gradually  led  to  build  domed  or  concealed 
nests,  and  afterwards  transmitted  this  instinct,  together  with 
their  bright  colours,  to  their  modified  descendants.  As  far  as  it 
can  be  trusted,  the  conclusion  is  interesting,  that  sexual  selection, 
together  with  equal  or  nearly  equal  inheritance  by  both  sexes, 
have  indirectly  determined  the  manner  of  nidification  of  whole 
groups  of  birds. 

According  to  Mr.  "Wallace,  even  in  the  groups  in  which  the. 
females,  from  being  protected  in  domed  nests  during  incubation, 

2"  See  many  statements  in  the  the  nests  of  Italian  birds  by  Eugenio 
'Ornithological  Biography.'  See,  Bettoni,  in  the  '  Atti  delhi  Societa 
also,  some  curious    observations  on      Italiana,'  vol.  xi.  1869,  p.  487. 


Chap.  XV.      Birds — Colour  and  Nidification.  457 

have  not  had  their  bright  colours  eliminated  through  natural 
selection,  the  males  often  differ  in  a  slight,  and  occasionally  in  a 
considerable  degree,  from  the  females.  This  is  a  significant  fact, 
for  such  differences  in  colour  must  be  accounted  for  by  some  of 
the  variations  in  the  males  having  been  from  the  first  limited  in 
transmission  to  the  same  sex ;  as  it  can  hardly  be  maintained  that 
these  differences,  esiDccially  when  very  slight,  serve  as  a  i^rotec- 
tion  to  the  female.  Thus  all  the  species  in  the  splendid  group 
of  the  Trogons  build  in  holes  ;  and  Mr.  Gould  gives  figures  ^^  of 
both  sexes  of  twenty-fiye  species,  in  all  of  which,  with  one  partial 
exception,  the  sexes  differ  sometimes  slightly,  sometimes  con- 
spciuously,  in  colour, — the  males  being  always  finer  than  the 
females,  though  the  latter  are  likewise  beautiful.  All  the  species 
of  kingfishers  build  in  holes,  and  with  most  of  the  species  the 
sexes  are  eciually  brilliant,  and  thus  far  Mr.  Wallace's  rule  holds 
good ;  but  in  some  of  the  Australian  species  the  colours  of  the 
females  are  rather  less  vivid  than  those  of  the  male ;  and  in  one 
splendidly-coloured  species,  the  sexes  differ  so  much  that  they 
were  at  first  thought  to  be  specifically  distinct.'-^  Mr.  E.  B.  Sharpe, 
who  has  especially  studied  this  group,  has  shewn  me  some 
American  species  (Ceryle)  in  which  the  breast  of  the  male  is 
belted  with  black.  Again,  in  Carcineutes,  the  difference  between 
the  sexes  is  conspicuous  :  in  the  male  the  upper  surface  is  dull- 
blue  banded  with  black,  the  lower  surface  being  partly  fawn- 
coloured,  and  there  is  much  red  about  the  head ;  in  the  female 
the  upi^er  surface  is  reddish-brown  banded  with  black,  and  the 
lower  surface  white  with  black  markings.  It  is  an  interesting 
fact,  as  shewing  how  the  same  peculiar  style  of  sexual  colouring 
often  characterises  allied  forms,  that  in  three  species  of  Dacelo 
the  male  differs  from  the  female  only  in  the  tail  being  dull-blue 
banded  with  black,  whilst  that  of  the  female  is  brown  with 
blackish  bars  ;  so  that  here  the  tail  differs  in  colour  in  the  two 
sexes  in  exactly  the  same  manner  as  the  whole  up^Dcr  surface  in 
the  two  sexes  of  Carcineutes. 

With  parrots,  which  likewise  build  in  holes,  we  find  analogous 
cases  :  in  most  of  the  species  both  sexes  are  brilliantly  coloured 
and  indistinguishable,  but  in  not  a  few  species  the  males  are 
coloured  rather  more  vividly  than  the  females,  or  even  very 
differently  from  them.  Thus,  besides  other  strongly-marked 
differences,  the  whole  under  surface  of  the  male  King  Lory 
(Aprosmictus  saipnlilus')  is  scarlet,  whilst  the  throat  and  chest  of 
the  female  is  green  tinged  with  red:  in  i\\Q  Euplie  ma  splcndida 

2'  See  his  *  Monograph  of  the  '  Handbook  to  the  Birds  of  Aus- 
Trogonidae,' first  edition.  tralia,' vol.  i.  p.   133 ;  see,  also,  pp 

2'^  Namely  CyanalcyoB.     Gould's      130,  136. 


458  TJie  Descent  of  Man.  Part  II. 

there  is  a  similar  difference,  tlie  face  and  wing-coverts  moreover 
of  the  female  being  of  a  paler  blue  than  in  the  male.^^  In  the 
family  of  the  tits  {Parinct),  which  build  concealed  nests,  the 
female  of  mir  common  blue  tomtit  {Par  us  cceruleus)  is  "much 
"  less  brightly  coloured  "  than  the  male ;  and  in  the  magnificent 
Sultan  yellow  tit  of  India  the  difference  is  greater.^* 

Again  in  the  great  group  of  the  woodpeckers,^-^  the  sexes  are 
generally  nearly  alike,  but  in  the  Megapicus  validas  all  those 
parts  of  the  head,  neck,  and  breast,  which  are  crimson  in  the 
male  are  pale  brown  in  the  female.  As  in  several  woodpeckers 
the  head  of  the  male  is  bright  crimson,  whilst  that  of  the  female 
is  plain,  it  occurred  to  me  that  this  colour  might  possibly  make 
the  female  dangerously  conspicuous,  whenever  she  put  her  head 
out  of  the  hole  containing  her  nest,  and  consequently  that  this 
colour,  in  accordance  with  Mr.  ^Yallace's  belief,  had  been  elimi- 
nated. This  view  is  strengthened  by  what  Malherbe  states  with 
respect  to  Indopicus  carlotta ;  namely,  that  the  young  females, 
like  the  young  males,  have  some  crimson  about  their  heads, 
but  that  this  colour  disaj^pears  in  the  adult  female,  whilst  it  is 
intensified  in  the  adult  male.  Nevertheless  the  following  con- 
siderations render  this  view  extremely  doubtful :  the  male  takes 
a  fair  share  in  incubation,-^  and  would  be  thus  almost  equally 
exposed  to  danger ;  both  sexes  of  many  species  have  their  heads 
of  an  equally  bright  crimson ;  in  otlier  species  the  difference 
between  the  sexes  in  the  amount  of  scarlet  is  so  slight  that  it 
can  hardly  make  any  appreciable  difference  in  the  danger 
incurred ;  and  lastly,  the  colouring  of  the  head  in  the  two  sexes 
often  differs  slightly  in  other  ways. 

The  cases,  as  yet  given,  of  slight  and  graduated  differences  in 
colour  between  the  males  and  females  in  the  groups,  in  which  as 
a  general  rule  the  sexes  resemble  each  other,  all  relate  to  species 
which  build  domed  or  concealed  nests.  But  similar  gradations 
may  likewise  be  observed  in  groups  in  which  the  sexes  as  a 
general  rule  resemble  each  other,  but  which  build  open  nests. 
As  I  have  before  instanced  the  Australian  parrots,  so  I  may  here- 
instance,  without  giving  any  details,  the  Australian  pigeons.^ 
It  deserves  especial  notice  that  in  all  these  cases  the  slight 

-'  Every  gradation  of  difference  taken  from  M.  Malherbe's  mag- 
between  the  sexes  may  be  followed  nificent  '  Monographic  des  Picideies,' 
in   the    parrots    of  Australia.     See  1861. 

Gould's  'Handbook,'  &c.,  vol.  ii.  pp.  ^e  Audubon's  '  Ornithological  Bio- 

14—102.  graphy,'  A'ol.  ii.  p.  75 ;  see  also  the 

'•'*  Macgillivray's  'British  Birds,'  'Ibis,'" vol.  i.  p.  268. 

vol.   li.  p.  433.     Jerdon,   '  Birds  of  ^'  Gould's     '  Handbook     to      the 

India,'  vol.  ii.  p.  282.  Birds  of  Australia,'  vol.  ii.  pp.  109- 

2^  All    the    following    facts     are  149. 


Chap.  XV.      Birds — Colon j'  aiid  Nidificatioit.  459 


differences  iu  plumage  between  the  sexes  are  of  the  same  general 
nature  as  the  occasionally  greater  differences.  A  good  illustra- 
tion of,  this  fact  has  already  been  afforded  by  those  kingfishers  in 
which  either  the  tail  alone  or  the  whole  upper  surface  of  the 
plumage  differs  in  the  same  manner  in  the  two  sexes.  Similar 
cases  may  be  observed  with  parrots  and  pigeons.  The  differ- 
ences in  colour  between  the  sexes  of  the  same  species  are,  also, 
of  the  same  general  nature  as  the  differences  in  colour  between 
the  distinct  species  of  the  same  group.  For  when  in  a  group  in 
which  the  sexes  are  usually  alike,  the  male  differs  considerably 
from  the  female,  he  is  not  coloured  in  a  quite  new  style.  Hence 
we  may  infer  that  within  the  same  group  the  special  colours  of 
both  sexes  when  they  are  alike,  and  the  colours  of  the  male,  when 
he  differs  slightly  or  even  considerably  from  the  female,  have 
been  in  most  cases  determined  by  the  same  general  cause ;  this 
being  sexual  selection. 

It  is  not  probaMe,  as  has  already  been  remarked,  that  differ- 
ences in  colour  between  the  sexes,  when  very  slight,  can  be  of 
service  to  the  female  as  a  protection.  Assuming,  however,  that 
they  are  of  service,  they  might  be  thought  to  be  cases  of 
transition ;  but  we  have  no  reason  to  believe  that  many  species 
at  any  one  time  are  undergoing  change.  Therefore  we  can 
hardly  admit  that  the  numerous  females  which  differ  very 
slightly  in  colour  from  their  males  are  now  all  commencing  to 
become  obscure  for  the  sake  of  protection.  Even  if  we  consider 
somewhat  more  marked  sexual  differences,  is  it  probable,  for 
instance,  that  the  head  of  the  female  chaffinch, — the  crimson  on 
the  breast  of  the  female  bullfinch, — the  green  of  the  female 
greenfinch, — the  crest  of  the  female  golden-crested  wren,  have 
all  been  rendered  less  bright  by  the  slow  process  of  selection  for 
the  sake  of  protection  ?  I  cannot  think  so ;  and  still  less  with  the 
slight  differences  between  the  sexes  of  those  birds  which  build 
concealed  nests.  On  the  other  hand,  the  differences  in  colour  be- 
tween the  sexes,  whether  great  or  small,  may  to  a  large  extent  be 
explained  on  the  principle  of  the  successive  variations,  acquired 
by  the  males  through  sexual  selection,  having  been  from  the 
first  more  or  less  limited  in  their  transmission  to  the  females. 
That  the  degree  of  limitation  should  differ  in  different  species  of 
the  same  group  will  not  surprise  any  one  who  has  studied  the 
laws  of  inheritance,  for  they  are  so  complex  that  they  appear  to 
us  in  our  ignorance  to  be  capricious  in  their  action.^** 

As  far  as  I  can  discover  there  are  few  large  groups  of  birds  in 
which  all  the  species  have   both  sexes  alike  and   brilliantly 

**■  See  remarks  to  this  effect  in  my  work  on  'Variation  under  Domesti- 
cation,' vol.  ii.  chap.  xii. 


460  TJie  Descent  of  Man.  Part  IT. 

coloured,  but  I  hear  from  Mr.  Sclater,  that  this  appears  to  be  the 
case  with  the  Musophagse  or  plantain-eaters.  Nor  do  I  believe 
that  any  large  group  exists  in  which  the  sexes  of  all  the  species 
are  widely  dissimilar  in  colour :  Mr.  Wallace  informs  me  that 
the  chatterers  of  S.  America  {Cotingidce)  offer  one  of  the  best 
instances ;  but  with  some  of  the  species,  in  which  the  male  has  a 
splendid  red  breast,  the  female  exhibits  some  red  on  her  breast ; 
and  the  females  of  other  species  shew  traces  of  the  green  and 
other  colours  of  the  males.  Nevertheless  we  have  a  near 
approach  to  close  sexual  similarity  or  dissimilarity  throughout 
several  groups :  and  this,  from  what  has  just  been  said  of  the 
fluctuating  nature  of  inheritance,  is  a  somewhat  surprising 
circumstance.  But  that  the  same  laws  should  largely  prevail 
with  allied  animals  is  not  surprising.  The  domestic  fowl  has 
produced  a  great  number  of  breeds  and  sub-breeds,  and  in  these 
the  sexes  generally  differ  in  plumage ;  so  that  it  has  been 
noticed  as  an  unusual  circumstance  when  in  certain  sub-breeds 
they  resemble  each  other.  On  the  other  hand,  the  domestic 
pigeon  has  likewise  produced  a  vast  number  of  distinct  breeds 
and  sub-breeds,  and  in  these,  with  rare  exceptions,  the  two  sexes 
are  identically  alike.  Therefore  if  other  species  of  Gallus  and 
Columba  were  domesticated  and  varied,  it  would  not  be  rash  to 
predict  that  similar  rules  of  sexual  similarity  and  dissimilarity, 
depending  on  the  form  of  transmission,  would  hold  good  in  both 
cases.  In  like  manner  the  same  form  of  transmission  has  gene- 
rally prevailed  imder  nature  throughout  the  same  groups, 
although  marked  exceptions  to  this  rule  occur.  Thus  within 
the  same  family  or  even  genus,  the  sexes  may  be  identically  alike, 
or  very  different  in  colour.  Instances  have  already  been  given 
in  the  same  genus,  as  with  sparrows,  fly-catchers,  thrushes  and 
grouse.  In  the  family  of  j^heasants  the  sexes  of  almost  all  the 
species  are  wonderfully  dissimilar,  but  are  quite  alike  in  the 
eared  pheasant  or  Crossopttlon  auritum.  In  two  species  of 
Chloephaga,  a  genus  of  geese,  the  male  cannot  be  distinguished 
from  the  females,  except  by  size ;  whilst  in  two  others,  the  sexes 
are  so  unlike  that  they  might  easily  be  mistaken  for  distinct 
.'■  pecies.-'* 

The  laws  of  inheritance  can  alone  account  for  the  following 
cases,  in  which  the  female  acquires,  late  in  life,  certain  characters 
proper  to  the  male,  and  ultimately  comes  to  resemble  him  more 
or  less  completely.  Here  protection  can  hardly  have  come  into 
play.  Mr.  Blyth  informs  me  that  the  females  of  Oriolus  meJano- 
ctphalus  and  of  some  allied  species,  when  sufficiently  mature  to 
breed,  differ  considerably  in  plumage  from  the  adult  males ;  but 
29  The  '  Ibis,'  vol.  vi.  1864,  p.  122. 


Chap.  XV,      Birds — Colour  and  Nidificatiou.  461 

after  the  second  or  third  moults  they  differ  only  in  their  beaks 
having  a  slight  greenish  tinge.  In  the  dwarf  bitterns  (Ardetta), 
according  to  the  same  authority,  "  the  male  acquires  his  final 
"  livery  at  the  first  moult,  the  female  not  before  the  third  or 
"  fourth  moult ;  in  the  meanwhile  she  presents  an  intermediate 
"  garb,  which  is  ultimately  exchanged  for  the  same  livery  as 
"  that  of  the  male."  So  again  the  female  Fulco  pei-egrihus  ac- 
quires her  blue  plumage  more  slowly  than  the  male.  Mr. 
Swinhoe  states  that  with  one  of  the  Drongo  shrikes  (Dicrurus 
macroccrcus)  the  male  whilst  almost  a  nestling,  moults  his  soft 
brown  plumage  and  becomes  of  a  uniform  glossy  greenish-black  ; 
but  the  female  retains  for  a  long  time  the  white  stria3  and  spots 
on  the  axillary  feathers ;  and  does  not  completely  assume  the 
uniform  black  colour  of  the  male  for  three  years.  The  same 
excellent  observer  remarks  that  in  tile  spring  of  the  second  year 
the  female  spoonbill  (Platalea)  of  China  resembles  the  male  of 
the  first  year,  and  that  apparently  it  is  not  until  the  third  spring 
that  she* acquires  the  same  adult  plumage  as  that  possessed  by 
the  male  at  a  much  earlier  age.  The  female  Bomhycilla  carolin- 
ensis  differs  very  little  from  the  male,  but  the  appendages,  which 
like  beads  of  red  sealing-wax  ornament  the  wing-feathers,^'^  are 
not  developed  in  her  so  early  in  life  as  in  the  male.  In  the  male 
of  an  Indian  parrakeet  (Palxomis  j'av m  ''ms)  the  upper  mandible 
is  coral-red  from  his  earliest  youth,  but  in  the  female,  as  Mr. 
Blyth  has  observed  with  caged  and  wild  birds,  it  is  at  first  black 
and  does  not  become  red  until  the  bird  is  at  least  a  year  old,  at 
which  age  the  sexes  resemble  each  other  in  all  respects.  Both 
sexes  of  the  wild  turkey  are  ultimately  furnished  with  a  tuft  of 
bristles  on  the  breast,  but  in  two-year-old  birds  the  tuft  is  about 
four  inches  long  in  the  male  and  hardly  apparent  in  the  female  ; 
when,  however,  the  latter  has  reached  her  fourth  year,  it  is  from 
four  to  five  inches  in  length. ^^ 

These  cases  must  not  be  confounded  with  those  where  diseased 
or  old  females  abnormally  assume  masculine  characters,  nor  with 

30  When  the  male  courts   the  fe-  '  Ibis,'   vol.    vi.   1864,   p.   366.     On 

male,  these  ornaments  are  vibrated,  'the   I3ombycilla,   Autlubon's    'Orni- 

aud  "  are  shewn  off"  to  great  advan-  tholog.   Biography,'  vol.   i.    p.   229. 

"tage,"  on  the  outstretched  wings:  On  the  Palaeornis,  see,  also,  Jcrdon, 

A.  Leith  Adams,  '  Field  and  Forest  '  Birds    of    India,'    vol.    i.    p.    263. 

Kambles,'  1873,  p.  153.  On  the  wild  turkey,  Audubon,  ibid. 

3'  On     Ardetta,     Translation     of  vol.  i,  p.  15  ;  but  I  hear  from  Judge 

Cuvier's   '  PJegne    Animal,'    by    Mr.  Caton    that    in    Ilhnois  the    female 

Blyth,    footnote,    p.    159.     On    the  very  rarely  acquires  a  tuft.    Analo- 

Peregrine     Falcon,     Mr.    Blyth,    in  gous     cases    with     the    females     of 

Charlesworth's  '  Mag.  of  Nat.  Hist.'  Petrocossvphus    are    given    by    Mr. 

vol.  i.  1837,  p.   304.     On  Dicrurus,  Pv.   B.  Sharpe,   '  Proc.   Zoolog.  Soc. 

*  Ibis,'  1863,  p.  44.     On  the  Platalea,  1872,  p.  496. 


462  The  Descejit  of  Man.  Paet  II. 


those  where  fertile  females,  whilst  young,  acquire  the  characters 
of  the  male,  through  variation  or  some  unknown  cause.^^  But 
all  these  cases  have  so  much  in  common  that  they  depend, 
according  to  the  hypothesis  of  pangenesis,  on  gemmules  derived 
from  each  part  of  the  male  being  present,  though  latent,  in  the 
female ;  their  development  following  on  some  slight  change  in 
the  elective  affinities  of  her  constituent  tissues. 

A  few  words  must  be  added  on  changes  of  plumage  in  relation 
to  the  season  of  the  year.  From  reasons  formerly  assigned  there 
can  be  little  doubt  that  the  elegant  plumes,  long  pendant 
feathers,  crests,  &c.,  of  egrets,  herons,  and  many  other  birds, 
which  are  developed  and  retained  only  during  the  summer, 
serve  for  ornamental  and  n^iptial  purposes,  though  common  to 
both  sexes.  The  female  is  thus  rendered  more  conspicuous 
during  the  period  of  incubation  than  during  the  winter ;  but 
such  birds  as  herons  and  egrets  would  be  able  to  defend  them- 
selves. As,  however,  plumes  would  probably  be  inconvenient 
and  certainly  of  no  use  during  the  winter,  it  is  possible  that  the 
habit  of  moulting  twice  in  the  year  may  have  been  gradually 
acquired  through  natural  selection  for  the  sake  of  casting  off 
inconvenient  ornaments  during  the  winter.  But  this  view 
cannot  be  extended  to  the  many  waders,  whose  summer  and 
winter  plumages  differ  very  little  in  colour.  With  defenceless 
species,  in  which  both  sexes,  or  the  males  alone,  become  extremely 
conspicuous  during  the  breeding-season, — or  when  the  males 
acquire  at  this  season  such  long  wing  or  tail-feathers  as  to  impede 
their  flight,  as  with  Cosmetornis  and  Vidua, — it  certainly  at  first 
appears  highly  probable  that  the  second  moult  has  been  gained 
for  the  special  purpose  of  throwing  off  these  ornaments.  We 
must,  however,  remember  that  many  birds,  such  as  some  of  the 
Birds  of  Paradise,  the  Argus  pheasant  and  peacock,  do  not  cast 
their  plumes  during  the  winter ;  and  it  can  hardly  be  maintained 
that  the  constitution  of  these  birds,  at  least  of  the  Gallinaceae, 
renders  a  double  moult  impossible,  for  the  ptarmigan  moults 
thrice  in  the  year.^^  Hence  it  must  be  considered  as  doubtful 
whether  the  many  species  which  moult  their  ornamental  plumes 
or  lose  their  bright  colours  during  the  winter,  have  acquired  this 
habit  on  account  of  the  inconvenience  or  danger  which  they 
would  otherwise  have  suffered. 

^"^  Of  these  latter  cases  Mr.  Blyth  also  recorded  a  similar  case  ('  Ornith. 

lias  recorded  (Translation  of  Cuvier's  Biog.'  vol.  v.  p.  519)  with    Tyranga 

'  Regne    Animal,'    p.    158)   various  ccstiva. 

instances    with    Lanius,     Ruticilla,  ^^  See    Gould's    '  Birds    of  Great 

Linaria,   and    Anas.     Audubon   has  Britain.' 


CuAP.  XVI.     Birds — InJieritance  Limited  by  Age.       463 

I  conclude,  therefore,  that  the  liabit  of  moulting  twice  in  the 
year  was  in  most  or  all  cases  first  acquired  for  some  distinct 
purpose,  perhaps  for  gaining  a  warmer  winter  covering ;  and  that 
variations  in  the  j^lumage  occurring  during  the  summer  were 
accumulated  through  sexual  selection,  and  transmitted  to  the  oif- 
spring  at  the  same  season  of  the  year  ;  that  such  variations  were 
inherited  either  by  both  sexes  or  by  the  males  alone,  according  to 
the  form  of  inheritance  which  prevailed.  This  appears  more 
probable  than  that  the  species  in  all  cases  originally  tended  to 
retain  their  ornamental  plumage  during  the  winter,  but  were 
saved  from  this  through  natural  selection,  resulting  from  the 
inconvenience  or  danger  thus  caused. 

I  have  endeavoured  in  this  chapter  to  shew  that  the  arguments 
are  not  trustworthy  in  favour  of  the  view  that  weapons,  bright 
colours,  and  various  ornaments,  are  now  confined  to  the  males 
owing  to  the  conversion,  by  natural  selection,  of  the  equal  trans- 
mission of  characters  to  both  sexes,  into  transmission  to  the  male 
sex  alone.  It  is  also  doubtful  whether  the  colours  of  many 
female  birds  are  due  to  the  preservation,  for  the  sake  of  protec- 
tion, of  variations  which  were  from  the  first  limited  in  their 
transmission  to  the  female  sex.  But  it  will  be  convenient  to 
defer  any  further  discussion  on  this  subject  until  I  treat,  in  the 
following  chapter,  of  the  differences  in  plumage  between  the 
young  and  old. 


CHAPTEE   XVI. 

BiEDS — concluded. 

The  immature  phimage  ia  relation  to  the  character  of  the  plumage  in 
both  sexes  when  adult — Six  classes  of  cases — Sexual  differences  between 
the  males  of  closely-allied  or  representative  species — The  female  as- 
suming the  characters  of  the  male — Plumage  of  the  young  in  relation 
to  the  summer  and  winter  plumage  of  the  adults — On  the  increase  of 
beauty  in  the  birds  of  the  world — Protective  colouring — Conspicuously- 
coloured  birds — Novelty  appreciated — Summary  of  the  four  chapters  on 
Birds. 

We  must  now  consider  the  transmission  of  characters,  as  limited 
by  age,  in  reference  to  sexual  selection.  The  truth  and  im- 
portance of  the  principle  of  inheritance  at  corresponding  ages 
need  not  here  be  discussed,  as  enough  has  already  been  said  on 
the  subject.  Before  giving  the  several  rather  complex  rules  or 
classes  of  cases,  under  which  the  differences  in  plumage  between 
the  young  and  the  old,  as  far  as  known  to  me,  may  be  included, 
it  will  be  well  to  make  a  few  preliminary  remarks. 


464  The  Descent  of  Man.  Part  II. 

With  animals  of  all  kinds  when  the  adults  differ  in  colour 
from  the  young,  and  the  colours  of  the  latter  are  not,  as  far  as 
we  can  see,  of  any  special  service,  they  may  generally  be 
attributed,  like  various  embryological  structures,  to  the  re- 
tention of  a  former  character.  But  this  view  can  be  maintained 
with  confidence,  only  when  the  young  of  several  species  resemble 
each  other  closely,  and  likewise  resemble  other  adult  species 
belonging  to  the  same  group ;  for  the  latter  are  the  living  proofs 
that  such  a  state  of  things  was  formerly  possible.  Young  lions 
and  pumas  are  marked  with  feeble  stripes  or  rows  of  spots,  and 
as  many  allied  species  both  young  and  old  are  similarly  marked, 
no  believer  in  evolution  will  doubt  that  the  progenitor  of  the 
lion  and  puma  was  a  striped  animal,  and  that  the  young  have 
retained  vestiges  of  the  stripes,  like  the  kittens  of  black  cats, 
which  are  not  in  the  least  striped  when  grown  up.  Many 
species  of  deer,  which  when  mature  are  not  spotted,  are  whilst 
young  covered  with  white  spots,  as  are  likewise  some  few  species 
in  the  adult  state.  So  again  the  young  in  the  whole  family  of 
pigs  ( Suid^e),  and  in  certain  rather  distantly  allied  animals,  such 
as  the  tapir,  are  marked  with  dark  longitudinal  stripes;  but 
here  we  have  a  character  apparently  derived  from  an  extinct 
progenitor,  and  now  preserved  by  the  young  alone.  In  all  such 
cases  the  old  have  had  their  colours  changed  in  the  course  of 
time,  whilst  the  young  have  remained  but  little  altered,  and  this 
has  been  effected  through  the  principle  of  inheritance  at  corre- 
siDonding  ages. 

This  same  principle  applies  to  many  birds  belonging  to 
various  groups,  in  which  the  young  closely  resemble  each 
other,  and  differ  much  from  their  respective  adult  parents.  The 
young  of  almost  all  the  Gallinacese,  and  of  some  distantly  allied 
birds  such  as  ostriches,  are  covered  with  longitudinally  striped 
down ;  but  this  character  points  back  to  a  state  of  things  so 
remote  that  it  hardly  concerns  us.  Young  cross-bills  (Loxia) 
have  at  first  straight  beaks  like  those  of  other  finches,  and  in 
their  immature  striated  plumage  they  resemble  the  mature 
redpole  and  female  siskin,  as  well  as  the  young  of  the  goldfinch, 
greenfinch,  and  some  other  allied  species.  The  young  of  many 
kinds  of  buntings  (Emberiza)  resemble  one  another,  and  like- 
wise the  adult  state  of  the  common  bunting,  E.  miliaria.  In 
almost  the  whole  large  group  of  thrushes  the  young  have  their 
breasts  spotted— a  character  which  is  retained  throughout  life 
by  many  species,  but  is  quite  lost  by  others,  as  by  the  Turdus 
migratorius.  So  again  with  many  thrushes,  the  feathers  on  the 
back  are  mottled  before  they  are  moulted  for  the  first  time,  and 
this  character  is  retained  for  life  by  certain  eastern  species. 


CuAP.  XVI.     Birds — Inheritance  Limited  by  Age.      465 

The  young  of  many  species  of  shrikes  (Lanius),  of  some  wood- 
peckers, and  of  an  Indian  pigeon  {Chakophaps  indicus),  are 
transversely  striped  on  the  under  surface;  and  certain  allied 
species  or  whole  genera  are  similarly  marked  when  adult.  In 
some  closely-allied  and  resplendent  Indian  cuckoos  (Chryso- 
Goccyx),  the  mature  species  differ  considerably  from  one  another 
in  colour,  but  the  young  cannot  be  distinguished.  The  young 
of  an  Indian  goose  (Sa7'kidiornis  melanonotus)  closely  resemble 
in  plumage  an  allied  genus,  Dendrocygna,  when  mature.* 
Similar  facts  will  hereafter  be  given  in  regard  to  certain  herons. 
Young  black  grouse  {Tetrao  tetria)  resemble  the  young  as  well 
as  the  old  of  certain  other  species,  for  instance  the  red  grouse 
or  T.  scoticus.  Finally,  as  Mr.  Blyth,  who  has  attended  closely 
to  this  subject,  has  well  remarked,  the  natural  affinities  of 
many  species  are  best  exhibited  in  their  immature  plumage ; 
and  as  the  true  affinities  of  all  organic  beings  depend  on  their 
descent  from  a  common  progenitor,  this  remark  strongly  con- 
firms the  belief  that  the  immature  plumage  approximately 
shews  us  the  former  or  ancestral  condition  of  the  species. 

Although  many  young  birds,  belonging  to  various  families, 
thus  give  us  a  glimpse  of  the  plumage  of  their  remote  pro- 
genitors, yet  there  are  many  other  birds,  both  dull-coloured  and 
bright-coloured,  in  which  the  young  closely  resemble  their 
parents.  In  such  cases  the  young  of  the  different  species  cannot 
resemble  each  other  more  closely  than  do  the  parents ;  nor  can 
they  strikingly  resemble  allied  forms  when  adult.  They  give  us 
but  little  insight  into  the  plumage  of  their  progenitors,  excepting 
in  so  far  that,  when  the  young  and  the  old  are  coloured  in  the 
same  general  manner  throughout  a  whole  group  of  species,  it  is 
probable  that  their  progenitors  were  similarly  coloured. 

We  may  now  consider  the  classes  of  cases,  under  which  the 
differences  and  resemblances  between  the  plumage  of  the  young 
and  the  old,  in  both  sexes  or  in  one  sex  alone,  may  be  grouped. 
Rules  of  this  kind  were  first  enounced  by  Cuvier ;  but  with  the 
progress  of  knowledge  they  require  some  modification  and  am- 
plification. This  I  have  attempted  to  do,  as  far  as  the  extreme 
complexity  of  the  subject  permits,  from  information  derived 
from  various  sources ;  but  a  full  essay  on  this  subject  by  some 
competent  ornithologist  is  much  needed.     In  order  to  ascertain 

'  In  regard  to  thrushes,  shrikes,  Oa    thrushes,    see    also    Audubon, 

and  woodpeckers,  see  Mr,  Blyth,  in  'Ornith.  Biography,'  vol.  ii.  p.  196. 

Charlesworth's  '  Mag.  of  Nat.  Hist,'  On  Chrysococcyx  and   Chalcophaps, 

vol.  i.   1837,  p.  304  ;  also  footnote  Blyth,  as  quoted  in  Jordon's  '  Birds 

to  his  translation  of  Cuvier's  '  Regne  of  India,' vol,  iii,  p.  485.     On  Sar- 

Animal,'   p.    159.     I   give   the  case  kidiornis,  Blyth,  in  '  Ibis,'  1867,  p. 

of  Loxia  on  Mr  Blyth's  information.  175. 
21 


466  The  Descent  of  Man.  Part  II. 

to  what  extent  each  rule  prevails,  I  have  tabulated  the  facts 
given  in  four  great  works,  namely,  by  Macgillivray  on  the  birds 
of  Britain,  Audubon  on  those  of  North  America,  Jerdon  on 
those  of  India,  and  Gould  on  those  of  Australia.  I  may  here 
premise,  first,  that  the  several  cases  or  rules  graduate  into  each 
other ;  and,  secondly,  that  when  the  young  are  said  to  resemble 
their  parents,  it  is  not  meant  that  they  are  identically  alike, 
for  their  colours  are  almost  always  less  vivid,  and  the  feathers 
are  softer  and  often  of  a  different  shape. 


RULES  OR  CLASSES  OF  CASES. 

I.  When  the  adult  male  is  more  beautiful  or  conspicuous 
than  the  adult  female,  the  young  of  both  sexes  in  their  first 
plumage  closely  resemble  the  adult  female,  as  with  the  common 
fowl  and  peacock ;  or,  as  occasionally  occurs,  they  resemble  her 
much  more  closely  than  they  do  the  adult  male. 

II.  When  the  adult  female  is  more  conspicuous  than  the 
adult  male,  as  sometimes  though  rarely  occurs,  the  young  of 
both  sexes  in  their  first  plumage  resemble  the  adult  male. 

III.  When  the  adult  male  resembles  the  adult  female,  the 
young  of  both  sexes  have  a  peculiar  first  plumage  of  their  own, 
as  with  the  robin. 

IV.  When  the  adult  male  resembles  the  adult  female,  the 
young  of  both  sexes  in  their  first  plumage  resemble  the  adults, 
as  with  the  kingfisher,  many  parrots,  crows,  hedge-warblers. 

V.  When  the  adults  of  both  sexes  have  a  distinct  winter  and 
summer  plumage,  whether  or  not  the  male  differs  from  the 
female,  the  young  resemble  the  adults  of  both  sexes  in  their 
winter  dress,  or  much  more  rarely  in  their  summer  dress,  or 
they  resemble  the  females  alone.  Or  the  young  may  have  an 
intermediate  character ;  or  again  they  may  differ  greatly  from 
the  adults  in  both  their  seasonal  plumages. 

VI.  In  some  few  cases  the  young  in  their  first  plumage  differ 
from  each  other  according  to  sex ;  the  young  males  resembling 
more  or  less  closely  the  adult  males,  and  the  young  females 
more  or  less  closely  the  adult  females. 

Class  I.— In  this  class,  the  young  of  both  sexes  more  or  less 
closely  resemble  the  adult  female,  whilst  the  adult  male  differs 
from  the  adult  female,  often  in  the  most  conspicuous  manner. 
Innumerable  instances  in  all  Orders  could  be  given;  it  will 
suffice  to  call  to  mind  the  common  i^heasant,  duck,  and  house- 
sparrow.  The  cases  under  this  class  graduate  into  others. 
Thus  the  two  sexes  when  adult  may  differ  so  slightly,  and  the 
young  so  slightly  from  the  adults,  that  it  is  doubtful  whether 


Chap.  XVI.     Birds —  Young  like  A  diilt  Females.         467 

such  cases  ought  to  come  under  the  present,  or  under  the  third 
or  fourth  classes.  So  again  the  young  of  the  two  sexes,  instead  of 
being  quite  alike,  may  differ  in  a  slight  degree  from  each  other, 
as  in  our  sixth  class.  These  transitional  cases,  however,  are 
few,  or  at  least  are  not  strongly  pronounced,  in  comparison  with 
those  which  come  strictly  under  the  present  class. 

The  force  of  the  present  law  is  well  shewn  in  those  groups, 
in  which,  as  a  general  rule,  the  two  sexes  and  the  ybung  are  all 
alike ;  for  when  in  these  groups  the  male  does  differ  from  the 
female,  as  with  certain  parrots,  kingfishers,  pigeons,  &c.,  the 
young  of  both  sexes  resemble  the  adult  female.^  We  see  the 
same  fact  exhibited  still  more  clearly  in  certain  anomalous  cases ; 
thus  the  male  of  lltUothrix  auriculata  (one  of  the  humming- 
birds) differs  conspicuously  from  the  female  in  having  a  s])lendid 
gorget  and  fine  ear-tufts,  but  the  female  is  remarkable  from 
having  a  much  longer  tail  than  that  of  the  male ;  now  the  young 
of  both  sexes  resemble  (with  the  exception  of  the  breast  being 
spotted  with  bronze)  the  adult  female  in  all  other  respects, 
including  the  length  of  her  tail,  so  that  the  tail  of  the  male 
actually  becomes  shorter  as  he  reaches  maturity,  which  is  a  most 
unusual  circumstance.^  Again,  the  plumage  of  the  male  goos- 
ander (Mergtfs  merganser)  is  more  conspicuously  coloured  than 
that  of  the  female,  with  the  scapular  and  secondary  wing-feathers 
much  longer;  but  differently  from  what  occurs,  as  far  as  I  know, 
in  any  other  bird,  the  crest  of  the  adult  male,  though  broader 
than  that  of  the  female,  is  considerably  shorter,  being  only  a 
little  above  an  inch  in  length ;  the  crest  of  the  female  being  two 
and  a  half  inches  long.  Now  the  young  of  both  sexes  entirely 
resemble  the  adult  female,  so  that  their  crests  are  actually  of 
greater  length,  though  narrower,  than  in  the  adult  male.* 

When  the  young  and  the  females  closely  resemble  each  other 
and  both  differ  from  the  males,  the  most  obvious  conclusion  is 

"^  See,  for   instance,   Mr.   Gould's  and  of  the  King  Lory,  with  which 

account  ('  Handbook  to  the  Birds  of  the  same  rule  prevails.     Also  Jerdon 

Australia,'  vol.  i.  p.  133)  of  Cyanal-  ('Birds  of  India,'  vol.  i.  p.  260)  on 

cyon  (one   of    the    Kingfishers)    iu  the   Palxornis   rosa,  in   which    the 

which,  however,  the    young   male,  young  are  more  like  the  female  than 

though  resembling  the  adult  female,  the  male.     See  Audubon  ('  Oruith. 

is  less  brilliantly  coloured.     In  some  Biograph.'   vol.    ii.   p.   475)  on    the 

species   of    Dacelo   the    males  have  two  sexes  and  the  young  of  Columba 

blue   tails,   and  the  females,  brown  passerina. 

ones ;  and  Mr.  R.  B.  Sharpe  informs  '  I  owe  this   information  to   Mr. 

me  that  the  tail  of  the  young  male  Gould,   who  shewed   me   the   spec'- 

of  i).  gandichaudi  is  at  ^rst  brown.  mens;  see  also   liis  'Introduction  to 

Mr.  Gould  has  described  (ibid.  vol.  the  Trochilida},'  1861,  p.  120. 
ii.  pp.  14,  20,  37)  the  sexes  and  the  *  Macgillivray, 'Hist.  Brit.  Birds, 

young   of  certaia   black    Cockatoos  vol.  v.  pp.  207-214. 


468  The  Descent  of  Man.  Part  IT. 

that  the  males  alone  have  been  modified.  Even  in  the  ano- 
malous cases  of  the  Heliothrix  and  Mergus,  it  is  probable  that 
originally  both  adult  sexes  were  furnished — the  one  species 
with  a  much  elongated  tail,  and  the  other  with  a  much  elon- 
gated crest — these  characters  having  since  been  partially  lost  by 
the  adult  males  from  some  unexplained  cause,  and  transmitted 
in  their  diminished  state  to  their  male  offspring  alone,  when 
arrived  at  the  corresponding  age  of  maturity.  The  belief  that 
in  the  present  class  the  male  alone  has  been  modified,  as  far  as 
the  differences  between  the  male  and  the  female  together  with 
her  young  are  concerned,  is  strongly  supported  by  some  re- 
markable facts  recorded  by  Mr.  Blyth,^  with  respect  to  closely- 
allied  species  which  represent  each  other  in  distinct  countries. 
For  with  several  of  these  representative  species  the  adult  males 
have  undergone  a  certain  amount  of  change  and  can  be  dis- 
tinguished ;  the  females  and  the  young  from  the  distinct  countries 
being  indistinguishable,  and  therefore  absolutely  unchanged. 
This  is  the  case  with  certain  Indian  chats  (Thamnobia),  with 
certain  honey-suckers  (Nectarinia),  shrikes  (Tephrodornis),  cer- 
tain kingfishers  (Tanysiptera),  Kallij  pheasants  (Gallophasis), 
and  tree-partridges  (Arboricola). 

In  some  analogous  cases,  namely  with  birds  having  a  different 
summer  and  winter  plumage,  but  with  the  two  sexes  nearly  alike, 
certain  closely-allied  species  can  easily  be  distinguished  in  their 
summer  or  nuptial  plumage,  yet  are  indistinguishable  in  their 
winter  as  well  as  in  their  immature  plumage.  This  is  the  case 
with  some  of  the  closely-allied  Indian  wag-tails  or  Motacillse. 
Mr.  Swinhoe^  informs  me  that  three  species  of  Ardeola,  a  genus 
of  herons,  which  represent  one  another  on  separate  continents, 
are  "most  strikingly  different"  when  ornamented  with  their 
summer  plumes,  but  are  hardly,  if  at  all,  distinguishable  during 
the  winter.  The  young  also  of  these  three  species  in  their 
immature  plumage  closely  resemble  the  adults  in  their  winter 
dress.  This  case  is  all  the  more  interesting,  because  with  two 
other  species  of  Ardeola  both  sexes  retain,  during  the  winter 
and  summer,  nearly  the  same  plumage  as  that  possessed  by  the 
three  first  species  during  the  winter  and  in  their  immature 
state;  and  this  plumage,  which  is  common  to  several  distinct 

^  See  his  admirable  paper  in  the  several    distinct    races,    solely     by 

'  Journal    of    the    Asiatic    Soc.    of  comparing  the  adult  males. 

Bengal,'  vol.  xix.   1850,  p.  223  ;  see  ^  See  also  Mr.  Swinhoe,  in  '  Ibis,' 

also  Jerdon,  '  Birds  of  hidia,'  vol.  i.  July  1863,  p.  131  ;  and  a  previous 

introduction,  p.  xxix.     In  regard  to  paper,  with  an  extract  from  a  note 

Tanysiptera,  Prof.  Schlegel  told  Mr.  by  Mr.  Blyth,  in  'Ibis,'  Jan.  1801, 

Blyth    that    he    could    distinguish  p.  25. 


Chap.  XVI.     Birds —  Young  like  A  diilt  Females.         469 

species  at  different  ages  and  seasons,  probably  shews  us  how 
the  progenitors  of  the  genus  were  coloured.  In  all  these  cases, 
the  nuptial  plumage  which  we  may  assume  was  originally 
acquired  by  the  adult  males  during  the  breeding-season,  and 
transmitted  to  the  adults  of  both  sexes  at  the  corresponding 
season,  has  been  modified,  whilst  the  winter  and  immature 
plumages  have  been  left  imchanged. 

The  question  naturally  arises,  how  is  it  that  in  these  latter 
cases  the  winter  plumage  of  both  sexes,  and  in  the  former  cases 
the  plumage  of  the  adult  females,  as  well  as  the  immature 
plumage  of  the  young,  have  not  been  at  all  affected?  The 
species  which  represent  each  other  in  distinct  countries  will 
almost  always  have  been  exposed  to  somewhat  different  con- 
ditions, but  we  can  hardly  attribute  to  this  action  the  modi- 
fication of  the  plumage  in  the  males  alone,  seeing  that  the 
females  and  the  young,  though  similarly  exposed,  have  not  been 
affected.  Hardly  any  fact  shews  us  more  clearly  how  subor- 
dinate in  importance  is  the  direct  action  of  the  conditions  of 
life,  in  comparison  with  the  accumulation  through  selection  of 
indefinite  variations,  than  the  surprising  difference  between  the 
sexes  of  many  birds ;  for  both  will  have  consumed  the  same  food, 
and  have  been  exposed  to  the  same  climate.  Nevertheless  we 
are  not  precluded  from  believing  that  in  the  course  of  time 
new  conditions  may  produce  some  direct  effect  either  on  both 
sexes,  or  from  their  constitutional  differences  chiefly  on  one  sex. 
We  see  only  that  this  is  subordinate  in  importance  to  the 
accumulated  results  of  selection.  Judging,  however,  from  a 
wide-spread  analogy,  when  a  species  migrates  into  a  new 
country  (and  this  must  precede  the  formation  of  representative 
species),  the  changed  conditions  to  which  they  will  almost 
always  have  been  exposed  will  cause  them  to  undergo  a  certain 
amount  of  fluctuating  variability.  In  this  case  sexual  selection, 
which  depends  on  an  element  liable  to  change— the  taste  or 
admiration  of  the  female — will  have  had  new  shades  of  colour 
or  other  differences  to  act  on  and  accumulate ;  and  as  sexual 
selection  is  always  at  work,  it  would  (from  what  we  know 
of  the  results  on  domestic  animals  of  man's  unintentional 
selection),  be  surprising  if  animals  inhabiting  separate  districts, 
which  can  never  cross  and  thus  blend  their  newly-acquired 
characters,  were  not,  after  a  sufficient  lapse  of  time,  differently 
modified.  These  remarks  likewise  apply  to  the  nuptial  or 
summer  plumage,  whether  confined  to  the  males  or  common  to 
both  sexes. 

Although  the  females  of  the  above  closely-allied  or  repre- 
sentative species,  together  with  their  young,  differ  hardly  at  all 


470  The  Descent  of  Man,  Part  II. 


from  one  another,  so  that  the  males  alone  can  be  distinguished, 
yet  the  females  of  most  species  within  the  same  genus  obviously 
differ  from  each  other.  The  differences,  however,  are  rarely 
as  great  as  between  the  males.  We  see  this  clearly  in  the 
whole  family  of  the  Gallinacese:  the  females,  for  instance,  of 
the  common  and  Japan  jDheasant,  and  especially  of  the  Gold  and 
Amherst  pheasant— of  the  silver  pheasant  and  the  wild  fowl 
— resemble  one  another  very  closely  in  colour,  whilst  the  males 
differ  to  an  extraordinary  degree.  So  it  is  with  the  females  of 
most  of  the  Cotingidse,  Fringillidse,  and  many  other  families. 
There  can  indeed  be  no  doubt  that,  as  a  general  rule,  the  females 
have  been  less  modified  than  the  males.  Some  few  birds, 
however,  offer  a  singular  and  inexplicable  exception ;  thus  the 
females  of  Faradisea  apoda  and  P.  papuana  differ  from  each 
other  more  than  do  their  respective  males  ;^  the  female  of  the 
latter  species  having  the  under  surface  pure  white,  whilst  the 
female  P.  apoda  is  deep  brown  beneath.  So,  again,  as  I  hear 
from  Professor  Newton,  the  males  of  two  species  of  Oxynotus 
(shrikes),  which  represent  each  other  in  the  islands  of  Mauritius 
and  Bourbon,^  differ  but  little  in  colour,  whilst  the  females  differ 
much.  In  the  Bourbon  species  the  female  appears  to  have 
partially  retained  an  immature  condition  of  plumage,  for  at 
first  sight  she  "  might  be  taken  for  the  young  of  the  Mauritian 
"  species."  These  differences  may  be  compared  with  those 
inexplicable  ones,  which  occur  independently  of  man's  selection 
in  certain  sub-breeds  of  the  game-fowl,  in  which  the  females  are 
very  different,  whilst  the  males  can  hardly  be  distinguished.^ 

As  I  account  so  largely  by  sexual  selection  for  the  differences 
between  the  males  of  allied  species,  how  can  the  differences 
between  the  females  be  accounted  for  in  all  ordinary  cases  ?  We 
need  not  here  consider  the  species  which  belong  to  distinct 
genera;  for  with  these,  adaptation  to  different  habits  of  life,  and 
other  agencies,  will  have  come  into  play.  In  regard  to  the 
differences  between  the  females  within  the  same  genus,  it  appears 
to  me  almost  certain,  after  looking  through  various  large  groups, 
that  the  chief  agent  has  been  the  greater  or  less  transference  to 
the  female  of  the  characters  acquired  by  the  males  through 
sexual  selection.  In  the  several  British  finches,  the  two  sexes 
differ  either  very  slightly  or  considerably  ;  and  if  we  compare 
the  females  of  the  greenfinch,  chaffinch,  goldfinch,  bullfinch, 
crossbill,  sparrow,  &c.,  we  shall  see  that  they  differ  from  one 

^Wallace,    'The     Malay    Archi-  'Ibis,' 1866,  p.  275. 

pelago,'  vol.  ii.  1869,  p.  394.  ^  '  Variation    of     Animals,    &c., 

®  These  species  are  described,  with  under  Domesticatign,'  vol.  i.  p.  251. 
coloured  figures,  by  M.  F.  Pollen,  in 


Chap.  XVJ.     Birds —  Young  like  A  dult  Females.         47 1 

another  chiefly  in  the  points  in  which  they  partially  resemble 
their  respective  males  ;  and  the  colours  of  the  males  may  safely 
be  attributed  to  sexual  selection.  "With  many  gallinaceous 
species  the  sexes  differ  to  an  extreme  degree,  as  with  the  peacock, 
pheasant,  and  fowl,  whilst  with  other  species  there  has  been  a 
partial  or  even  complete  transference  of  character  from  the  male 
to  the  female.  The  females  of  the  several  species  of  Polyplectron 
exhibit  in  a  dim  condition,  and  chiefly  on  the  tail,  the  splendid 
oc-elli  of  their  males.  The  female  partridge  differs  from  the 
male  only  in  the  red  mark  on  her  breast  being  smaller ;  and  the 
female  wild  turkey  only  in  her  colours  being  much  duller.  In 
the  guinea-fowl  the  two  sexes  are  indistinguishable.  There  is  no 
improbability  in  the  plain,  though  peculiarly  spotted  plumage 
of  this  latter  bird  having  been  acquired  through  sexual  selection 
by  the  males,  and  then  transmitted  to  both  sexes ;  for  it  is  not 
essentially  different  from  the  much  more  beautifully  spotted 
plumage,  characteristic  of  the  males  alone  of  the  Tragopan 
pheasants. 

It  should  be  observed  that,  in  some  instances,  the  transference 
of  characters  from  the  male  to  the  female  has  been  effected 
apparently  at  a  remote  period,  the  male  having  subsequently 
undergone  great  changes,  without  transferring  to  the  female  any 
of  his  later-gained  characters.  For  instance,  the  female  and  the 
young  of  the  black-grouse  (  Tdmo  tetriv)  resemble  pretty  closely 
both  sexes  and  the  young  of  the  red-grouse  (T.  scoticus)  ;  and  we 
may  consequently  infer  that  the  black-grouse  is  descended  from 
some  ancient  species,  of  which  both  sexes  were  coloured  in 
nearly  the  same  manner  as  the  red-grouse.  As  both  sexes  of 
this  latter  species  are  more  distinctly  barred  during  the  breeding 
season  than  at  any  other  time,  and  as  the  male  differs  slightly 
from  the  female  in  his  more  strongly-pronounced  red  and  brown 
tints,^"  we  may  conclude  that  his  plumage  has  been  influenced 
by  sexual  selection,  at  least  to  a  certain  extent.  If  so,  we  may 
further  infer  that  the  nearly  similar  plumage  of  the  female  black- 
grouse  was  similarly  produced  at  some  former  period.  But 
since  this  period  the  male  black-grouse  has  acquired  his  fine 
black  plumage,  with  his  forked  and  outwardly- curled  tail- 
feathers;  but  of  these  characters  there  has  hardly  been  any 
transference  to  the  female,  excepting  that  she  shews  in  her  tail 
a  trace  of  the  curved  fork. 

We  may  therefore  conclude  that  the  females  of  distinct  though 

allied  species  have  often  had  their  plumage  rendered  more  or 

less  different  by  the  transference  in  various  degrees,  of  characters 

acquired  by  the  males  through  sexual   selection,  both  during 

10  Macgillivray,  'Hist.  British  Birds,'  vol.  i.  pp.  172-174. 


472  The  Descent  of  Alan.         "         Part  II. 

former  and  recent  times.  But  it  deserves  especial  attention  that 
brilliant  colours  have  been  transferred  much  more  rarely  than 
other  tints.  For  instance,  the  male  of  the  red-throated  blue- 
breast  (6'?/a?<ecM?a  siiecica)  has  a  rich  blue  breast,  including  a  sub- 
triangular  red  mark ;  now  marks  of  nearly  the  same  shape  have 
been  transferred  to  the  female,  but  the  central  space  is  fulvous 
instead  of  red,  and  is  surrounded  by  mottled  instead  of  blue 
feathers.  The  Gallinaceae  offer  many  analogous  cases ;  for  none  of 
the  species,  such  as  partridges,  quails,  guinea-fowls,  &c.,  in  which 
the  colours  of  the  plumage  have  been  largely  transferred  from 
the  male  to  the  female,  are  brilliantly  coloured.  This  is  well 
exemplified  with  the  pheasants,  in  which  the  male  is  generally 
so  much  more  brilliant  than  the  female ;  but  with  the  Eared 
and  Cheer  pheasants  (^(Jrossoptilon  auritum  and  Fhasianus  wal- 
lichii)  the  sexes  closely  resemble  each  other  and  their  colours  are 
dull.  We  may  go  so  far  as  to  believe  that  if  any  part  of  the 
plumage  in  the  males  of  these  two  pheasants  had  been  brilliantly 
coloured,  it  would  not  have  been  transferred  to  the  females. 
These  facts  strongly  support  Mr.  Wallace's  view  that  with  birds 
which  are  exposed  to  much  danger  during  incubation,  the 
transference  of  bright  colours  from  the  male  to  the  female  has 
been  checked  through  natural  selection.  We  must  not,  however, 
forget  that  another  exjDlanation,  before  given,  is  possible  ;  namely, 
that  the  males  which  varied  and  became  bright,  whilst  they  were 
young  and  inexperienced,  would  have  been  exposed  to  much 
danger,  and  would  generally  have  been  destroyed ;  the  older  and 
more  cautious  males,  on  the  other  hand,  if  they  varied  in  a  like 
manner,  would  not  only  have  been  able  to  survive,  but  would 
have  been  favoured  in  their  rivalry  with  other  males.  Now 
variations  occurring  late  in  life  tend  to  be  transmitted  exclusively 
to  the  same  sex,  so  that  in  this  case  extremely  bright  tints  would 
not  have  been  transmitted  to  the  females.  On  the  other  hand, 
ornaments  of  a  less  conspicuous  kind,  such  as  those  possessed  by 
the  Eared  and  Cheer  pheasants,  would  not  have  been  dangerous, 
and  if  they  appeared  during  early  youth,  would  generally  have 
been  transmitted  to  both  sexes. 

In  addition  to  the  effects  of  the  partial  transference  of  charac- 
ters from  the  males  to  the  females,  some  of  the  differences 
between  the  females  of  closely  allied  species  may  be  attributed  to 
the  direct  or  definite  action  of  the  conditions  of  life.^^  With  the 
males,  any  such  action  would  generally  have  been  masked  by  the 
brilliant  colours  gained  through  sexual  selection;  but  not  so 
with  the  females.     Each  of  the  endless  diversities  in  plumage, 

"  See,  on  this  subject,  chap,  xxiii.  in  the  '  Variation  of  Auinnals  and 
Plants  under  Domestication.' 


Cuy\p.  XVI.     Birds —  Yotmg  like  Adult  Females.         473 


which  we  see  in  our  domesticated  birds  is,  of  course,  the  result 
of  some  definite  cause;  and  under  natural  and  more  uniform 
conditions,  some  one  tint,  assuming  that  it  was  in  no  way 
injurious,  would  almost  certainly  sooner  or  later  prevail.  The 
free  intercrossing  of  the  many  individuals  belonging  to  the  same 
species  would  ultimately  tend  to  make  any  change  of  colour, 
thus  induced,  uniform  in  character. 

Ko  one  doubts  that  both  sexes  of  many  birds  have  had  their 
colours  adapted  for  the  sake  of  protection ;  and  it  is  possible 
that  the  females  alone  of  some  species  may  have  been  modified 
for  this  end.  Although  it  would  be  a  difficult,  perhaps  an 
impossible  process,  as  shewn  in  the  last  chapter,  to  convert  one 
form  of  transmission  into  another  through  selection,  there  would 
not  be  the  least  difficulty  in  adapting  the  colours  of  the  female, 
independently  of  those  of  the  male,  to  surrounding  objects, 
through  the  accumulation  of  variations  which  were  from  the 
first  limited  in  their  transmission  to  the  female  sex.  If  the 
variations  were  not  thus  limited,  the  bright  tints  of  the  male 
would  be  deteriorated  or  destroyed.  Whether  the  females  alone 
of  many  species  have  been  thus  specially  modified,  is  at  present 
very  doubtful.  I  wish  I  could  follow  Mr.  Wallace  to  the  full 
extent;  for  the  admission  would  remove  some  difficulties.  Any 
variations  which  were  of  no  service  to  the  female  as  a  protection 
would  be  at  once  obliterated,  instead  of  being  lost  simply  by  not 
being  selected,  or  from  free  intercrossing,  or  from  being  elimin- 
ated when  transferred  to  the  male  and  in  any  way  injurious  to 
him.  Thus  the  plumage  of  the  female  would  be  kept  constant 
in  character.  It  would  also  be  a  relief  if  we  could  admit  that 
the  obscure  tints  of  both  sexes  of  many  birds  had  been  acquired 
and  preserved  for  the  sake  of  protection, — for  example,  of  the 
hedge-warbler  or  kitty- wren  {Accentor  modular  is  and  Troglodytes 
vulgaris),  with  respect  to  which  we  have  no  sufficient  evidence 
of  the  action  of  sexual  selection.  We  ought,  however,  to  be 
cautious  in  concluding  that  colours  which  appear  to  us  dull,  are 
not  attractive  to  the  females  of  certain  species  ;  we  should  bear 
in  mind  such  cases  as  that  of  the  common  house-sparrow, 
in  which  the  male  differs  much  from  the  female,  but  does  not 
exhibit  any  bright  tints.  No  one  probably  will  dispute  that  many 
gallinaceous  birds  which  live  on  the  open  ground,  have  acquired 
their  present  colours,  at  least  in  part,  for  the  sake  of  protection. 
We  know  how  well  they  are  thus  concealed;  we  know  that 
ptarmigans,  whilst  changing  from  their  winter  to  their  summer 
jjlumage,  both  of  which  are  protective,  suffer  greatly  from  birds 
of  prey.  But  can  we  believe  that  the  very  slight  difi'erences  in 
tints  and  markings  between,  for  instance,  the  female  black-grouse 


474  The  Descent  of  Man.  Part  II. 

and  red-grouse  serve  as  a  iDrotection  ?  Are  partridges,  as  they 
are  now  coloured,  better  protected  than  if  they  had  resembled 
quails  ?  Do  the  slight  di£ferences  between  the  females  of  the 
common  pheasant,  the  Japan  and  gold  pheasants,  serve  as  a 
protection,  or  might  not  their  plumages  have  been  interchanged 
with  impunity  ?  From  what  Mr.  Wallace  has  observed  of  the 
habits  of  certain  gallinaceous  birds  in  the  East,  he  thinks  that 
such  slight  differences  are  beneficial.  For  myself,  I  will  only 
say  that  I  am  not  convinced. 

Formerly  when  I  was  inclined  to  lay  much  stress  on  protection 
as  accounting  for  the  duller  colours  of  female  birds,  it  occurred 
to  me  that  possibly  both  sexes  and  the  young  might  aboriginally 
have  been  equally  bright  coloured ;  but  that  subsequently,  the 
females  from  the  danger  incurred  during  incubation,  and  the 
young  from  being  inexperienced,  had  been  rendered  dull  as  a 
protection.  But  this  view  is  not  supported  by  any  evidence, 
and  is  not  probable ;  for  we  thus  in  imagination  expose  during 
past  times  the  females  and  the  youDg  to  danger,  from  which  it 
has  subsequently  been  necessary  to  shield  their  modified 
descendants.  We  have,  also,  to  reduce,  through  a  gradual 
process  of  selection,  the  females  and  the  young  to  almost  exactly 
the  same  tints  and  markings,  and  to  transmit  them  to  the 
corresponding  sex  and  period  of  life.  On  the  supposition  that 
the  females  and  the  young  have  partaken  during  each  stage  of 
the  process  of  modification  of  a  tendency  to  be  as  brightly 
coloured  as  the  males,  it  is  also  a  somewhat  strange  fact  that  the 
females  have  never  been  rendered  dull-coloured  without  the 
young  participating  in  the  same  change;  for  there  are  no  in- 
stances, as  far  as  I  can  discover,  of  species  with  the  females  dull 
and  the  young  bright  coloured.  A  partial  exception,  however, 
is  offered  by  the  young  of  certain  woodpeckers,  for  they  have 
"  the  whole  upper  part  of  the  head  tinged  with  red,"  which 
afterwards  either  decreases  into  a  mere  circular  red  line  in  the 
adults  of  both  sexes,  or  quite  disappears  in  the  adult  females.^^ 

Finally,  with  respect  to  our  present  class  of  cases,  the  most 
probable  view  appears  to  be  that  successive  variations  in 
brightness  or  in  other  ornamental  characters,  occurring  in  the 
males  at  a  rather  late  period  of  life  have  alone  been  preserved ; 
and  that  most  or  all  of  these  variations,  owing  to  the  late  period 
of  life  at  which  they  appeared,  have  been  from  the  first  trans- 
mitted only  to  the  adult  male  offspring.  Any  variations  in 
brightness  occurring  in  the  females  or  in  the  young,  would  have 

^2  Audubon,  *  Oruith.  Biography,'      also  the  case  before  giveu  of  Indo- 
vol.  i.  p.  193.     Macgillivray,  'Hist,      incus  carlotta. 
Brit.     Birds '   vol.   iii.    p.    85.     See 


Cjiap.  XVI.    Birds —  Young  like  Adult  Males.  475 

been  of  no  service  to  tliem,  and  would  not  have  been  selected ; 
and  moreover,  if  dangerous,  would  have  been  eliminated.  Thus 
the  females  and  the  young  will  either  have  been  left  unmodified, 
or  (as  is  much  more  common)  will  have  been  partially  modified 
by  receiving  through  transference  from  the  males  some  of  his 
successive  variations.  Both  sexes  have  perhaps  been  directly 
acted  on  by  the  conditions  of  life  to  which  they  have  long 
been  exposed :  but  the  females  from  not  being  otherwise  much 
modified,  will  best  exhibit  any  such  effects.  These  changes 
and  all  others  will  have  been  kept  uniform  by  the  free  inter- 
crosdng  of  many  individuals.  In  some  cases,  especially  with 
ground  birds,  the  females  and  the  young  may  possibly  have  been 
modified,  independently  of  the  males,  for  the  sake  of  protection, 
80  as  to  have  acquired  the  same  dull  coloured  plumage. 

Class  II.  Whtn  the  adult  female  is  more  conspicuous  titan  the 
adult  male,  the  young  of  both  sexes  in  their  first  plumage  resemble 
the  adult  male. — This  class  is  exactly  the  reverse  of  the  last,  for 
the  females  are  here  brighter  coloured  or  more  conspicuous  than 
the  males ;  and  the  young,  as  far  as  they  are  known,  resemble 
the  adult  males  instead  of  the  adult  females.  But  the  difference 
between  the  sexes  is  never  nearly  so  great  as  with  many  birds  in 
the  first  class,  and  the  cases  are  comparatively  rare.  Mr. 
Wallace,  who  first  called  attention  to  the  singular  relation  which 
exists  between  the  less  bright  colours  of  the  males  and  their 
performing  the  duties  of  incubation,  lays  great  stress  on  this 
point,^^  as  a  crucial  test  that  obscure  colours  have  been  acquired 
for  the  sake  of  protection  during  the  period  of  nesting.  A 
different  view  seems  to  me  more  probable.  As  the  cases  are 
curious  and  not  numerous,  I  will  briefly  give  all  that  I  have 
been  able  to  find. 

In  one  section  of  the  genus  Turnix,  quail- like  birds,  the  female 
is  invariably  larger  than  the  male  (being  nearly  twice  as  large 
in  one  of  the  Australian  species),  and  this  is  an  unusual  circum- 
stance with  the  Gallinacese.  In  most  of  the  species  the  female  is 
more  distinctly  coloured  and  brighter  than  the  male,^^  but  in 
some  few  species  the  sexes  are  alike.  In  Turnix  taigoor  of  India 
the  male  "  wants  the  black  on  the  throat  and  neck,  and  the 
"  whole  tone  of  the  plumage  is  lighter  and  less  pronounced  than 
"  that  of  the  female."  The  female  appears  to  be  noisier,  and  is 
certainly  much  more  pugnacious  than  the  male;    so  that  the 

>3  <  Westminster     Review,'   July,  178,   180,    186,    and     188.     In   the 

1867,  and  A.  Murray,  '  Journal  of  British    Museum    specimens  of   the 

Travel,'  1868,  p.  83.  Australian    Plain-wanderer    (^Fedio- 

'■*  For  the  Australian  species,  see  notnus     torquatus)    may     be     seen, 

Gould's  '  Handbook,'  &c.,  vol.  ii.  pp.  shewinjj;  similar  sexual  dift'erenccs. 


4/6  TJie  Descent  of  Man.  Part  II. 

females  and  not  the  males  are  often  kept  by  the  natives  for 
fighting,  like  game-cocks.  As  male  birds  are  exposed  by  the 
English  bird-catchers  for  a  decoy  near  a  trap,  in  order  to  catch 
other  males  by  exciting  their  rivalry,  so  the  females  of  this 
Turnix  are  employed  in  India.  When  thus  exposed  the  females 
soon  begin  their  "  loud  purring  call,  which  can  be  heard  a  long 
"  way  off,  and  any  females  within  ear-shot  run  rapidly  to  the 
"  spot,  and  commence  fighting  with  the  caged  bird."  In  this  way 
from  twelve  to  twenty  birds,  all  breeding  •  females,  may  be 
caught  in  the  course  of  a  single  day.  The  natives  assert  that 
the  females  after  laying  their  eggs  associate  in  flocks,  and  leave 
the  males  to  sit  on  them.  There  is  no  reason  to  doubt  the 
truth  of  this  assertion,  which  is  supported  by  some  observations 
made  in  China  by  Mr.  Swinhoe.^^  Mr.  Blyth  believes,  that  the 
young  of  both  sexes  resemble  the  adult  male. 

The  females  of  the  three  species  of  Painted  Snipes  (Ehynchssa, 
fig.  62)  "  are  not  only  larger  but  much  more  richly  coloured  than 
"  the  males."  ^^  With  all  other  birds  in  which  the  trachea  differs 
in  structure  in  the  two  sexes  it  is  more  developed  and  complex 
in  the  male  than  in  the  female;  but  in  the  Rhynchxa  australis 
it  is  simple  in  the  male,  whilst  in  the  female  it  makes  four 
distinct  convolutions  before  entering  the  lungs.  ^^  The  female 
therefore  of  this  species  has  acquired  an  eminently  masculine 
character.  Mr.  Blyth  ascertained,  by  examining  many  speci- 
mens, that  the  trachea  is  not  convoluted  in  either  sex  of  B. 
hengalensis,  which  species  resembles  R.  ausfrah's  so  closely,  that 
it  can  hardly  be  distinguished  except  by  its  shorter  toes.  This 
fact  is  another  striking  instance  of  the  law  that  secondary 
sexual  characters  are  often  widely  different  in  closely-allied 
forms,  though  it  is  a  very  rare  circumstance  when  such  differ- 
ences relate  to  the  female  sex.  The  young  of  both  sexes  of  B. 
hengalensis  in  their  first  plumage  are  said  to  resemble  the 
mature  male.^®  There  is  also  reason  to  believe  that  the  male 
undertakes  the  duty  of  incubation,  for  Mr.  Swinhoe  ^^  found  the 
females  before  the  close  of  the  summer  associated  in  flocks,  as 
occurs  with  the  females  of  the  Turnix. 

The  females  of  Phahirojy/is  fulicarlus  and  P.  hyperhoreus  are 
larger,  and  in  their  summer  plumage  "  more  gaily  attired  than 
"  the  males."  But  the  difference  in  colour  between  the  sexes  is 
far  from  conspicuous.     According  to  Professor  Steenstrup,  the 

"  Jerdon,    'Birds  of  India,'   vol.  '"  Gould's     'Handbook     to     the 

iii.  p.  596.     Mr.    Swinhoe,  in  'Ibis,'  Birds  of  Australia,'  vol.  ii.  p.  275. 
1865,  p.  542  ;  1866,  pp.  131,  405.  '»  '  The  Indian  Field,'  Sept.  1858, 

'^  Jerdon,   '  Birds  of  India,'  vol.  p.  3. 
iii.  p.  677.  i»  '  Ibis,'  1866,  p.  298. 


Chap.  XVI.     Birds—  Young  like  Adult  Males,  477 


male  alone  of  P.ftdkanus  imdertakes  the  duty  of  incubation; 
this  is  hkewise  shewn  by  the  state  of  his  breast-feathers  during 
the  breeding-season.  The  female  of  the  dotterel  plover  {Eudro- 
mias'thorinellus)  is  larger  than  the  male,  and  has  the  red  and 


Fig.  62. 


Rhynchasa  capensis  (from  Brehm). 


black  tints  on  the  lower  surface,  the  white  crescent  on  the 
breast,  and  the  stripes  over  the  eyes,  more  strongly  pronounced. 
The  male  also  takes  at  least  a  share  in  hatching  the  eggs;  but 
the  female  likewise  attends  to  the  young.-^    I  have  not  been 

the  above-named  species  take  either 
the  whole  or  a  large  share  of 
the  duties  of  incubation,  and  that 
they  "shew  mucli  greater  devotion 
"  towards  their  young,  when  in 
"danger,  than  do  the  females."     So 


2<^  For  these  several  statements, 
see  Mr.  Gould's  '  Birds  of  Great 
Britain.'  Prof.  Newton  informs  me 
that  he  has  long  ))een  convinced, 
from  his  own  observations  and  from 
those  of  others,  that  the  males  of 


47  S  The  Descent  of  Man.  Part  II. 

able  to  discover  whether  with  these  species  the  young  resemble 
the  adult  males  more  closely  than  the  adult  females;  for  the 
comparison  is  somewhat  difficult  to  make  on  account  of  the  double 
moult. 

Turning  now  to  the  Ostrich  order :  the  male  of  the  common 
cassowary  {Camarim  guleatm)  would  be  thought  by  any  one 
to  be  the  female,  from  his  smaller  size  and  from  the  appendages 
and  naked  skin  about  his  head  being  much  less  brightly  coloured  ; 
and  I  am  informed  by  Mr.  Bartlett  that  in  the  Zoological 
Gardens,  it  is  certainly  the  male  alone  who  sits  on  the  eggs  and 
takes  care  of  the  young.^^  The  female  is  said  by  Mr.  T.  W. 
Wood  '^  to  exhibit  during  the  breeding  season  a  most  pugnacious 
disposition;  and  her  wattles  then  become  enlarged  and  more 
brilliantly  coloured.  So  again  the  female  of  one  of  the  emus 
(^DromKiis  irroratus)  is  considerably  larger  than  the  male,  and 
she  possesses  a  slight  top-knot,  but  is  otherwise  indistinguishable 
in  plumage.  She  appears,  however,  "  to  have  greater  power, 
"  when  angry  or  otherwise  excited,  of  erecting,  like  a  turkey- 
"  cock,  the  feathers  of  her  neck  and  breast.  She  is  usually  the 
"  more  courageous  and  pugilistic.  She  makes  a  deep  hollow 
"  guttural  boom  especially  at  night,  sounding  like  a  small  gong. 
*'  The  male  has  a  slenderer  frame  and  is  more  docile,  with  no 
"  voice  beyond  a  suppressed  hiss  when  angry,  or  a  croak."  He 
not  only  performs  the  whole  duty  of  incubation,  but  has  to 
defend  the  young  from  their  mother ;  "  for  as  soon  as  she 
''  catches  sight  of  her  progeny  she  becomes  violently  agitated, 
"  and  notwithstanding  the  resistance  of  the  father  appears  to 
"  use  her  utmost  endeavours  to  destroy  them.  For  months 
"  afterwards  it  is  unsafe  to  put  the  parents  together,  violent 
"  quarrels  being  the  inevitable  result,  in  which  the  female  gene- 
*'  rally  comes  off  conqueror."  ^^.  So  that  with  this  emu  we  have 
a  complete  reversal  not  only  of  the  parental  and  incubating 
instincts,  but  of  the  usual  moral  qualities  of  the  two  sexes ;  the 
females  being  savage,  quarrelsome,  and  noisy,  the  males  gentle 
and  good.  The  case  is  very  different  with  the  African  ostrich, 
for  the  male  is  somewhat  larger  than  the  female  and  has  finer 


It  IS,  as  he  informs  me,  with  Limosa  assertion,   as    Mr.  Bartlett   thinks, 

lapponica     and     some     few     other  may  be  accounted  for  by  the  female 

Waders,    in    which   the  females  are  visiting  the  nest  to  lay  her  eggs, 

larger  and  have  more  strongly  con-  _"  'The  Student,'  April,  1870,  p. 

trasted  colours  than  the  males.  124. 

2^  The  natives  of  Ceram  (Wallace,  ^'  See  the  excellent  account  of 
'  Malay  Archipelago,'  vol.  ii.  p.  150)  the  habits  of  this  bird  under  confine- 
assert  that  the  male  and  female  sit  ment,  by  Mr.  A.  W.  Bennett,  in  '  Land 
alternately   on    the  eggs  ;  but    this  and  Water,'  May,  18G8,  p.  233. 


CiiAp.  XV  I.     Birds —  Young  like  A  dult  Males.  479 

plumes  with  more  strongly  contrasted  colours;  ncvorthelc^ss  he 
undertakes  the  whole  duty  of  incubation.^^ 

I  will  specify  the  few  other  cases  known  to  me,  in  which  the 
female  is  more  conspicuously  coloured  than  the  male,  although 
nothing  is  known  about  the  manner  of  incubation.  With  the 
carrion-hawk  of  the  Falkland  Islands  {Milvago  leucums)  I  was 
much  surprised  to  find  by  dissection  that  the  individuals,  which 
had  all  their  tints  strongly  pronounced,  with  the  cere  and  legs 
orange-coloured,  were  the  adult  females;  whilst  those  with 
duller  plumage  and  grey  legs  were  the  males  or  the  young.  In 
an  Australian  tree-creeper  (Climacteris  ert/throjis)  the  female 
differs  from  the  male  in  "  being  adorned  with  beautiful,  ra- 
"  dialed,  rufous  markings  on  the  throat,  the  male  having  this 
"  part  quite  plain."  Lastly,  in  an  Australian  night-jar  "  the 
"  female  always  exceeds  the  male  in  size  and  in  the  brilliance 
"  of  her  tints ;  the  males,  on  the  other  hand,  have  two  white 
"  spots  on  the  primaries  more  conspicuous  than  in  the  female."  ^^ 

We  thus  see  that  the  cases  in  which  female  birds  are  more 
conspicuously  coloured  than  the  males,  with  the  young  in  their 
immature  plumage  resembling  the  adult  males  instead  of  the 
adult  females,  as  in  the  previous  class,  are  not  numerous,  though 
they  are  distributed  in  various  Orders.  The  amount  of  differ- 
ence, also,  between  the  sexes  is  incomparably  less  than  that 
which  frequently  occurs  in  the  last  class ;  so  that  the  cause  of 
the  difference,  whatever  it  may  have  been  has  here  acted  on  the 
females  either  less  energetically  or  less  persistently  than  on  the 

2-»  Mr.  Sclater,  on  the  incubation  than  that  of  the  male ;  the  head  of 

of  the  Struthiones, '  Proc.  Zool.  Soc.,'  the  male  is  of  a  rich  dark  bronzed 

June   9,    1863.     So   it   is  with   the  colour,  and  his  back  is  clothed  with 

i?Am  c^ani^mu  :  Captain  Musters  says  finely  pencilled    slate-coloured   fea- 

('  At   home  with  the  Patagonians,'  thers,  so  that  altogether  he  may  be 

1871,    p.    128),  that  the    male    is  considered  as  the  more  beautiful  of 

larger,  stronger    and    swifter  than  the   two.     He   is   larger  and    more 

the  female,  and  of  slightly  darker  pugnacious    than    the    female,   and 

colours  ;  yet  he  takes  sole  charge  of  does  not  sit  on  the  eggs.     So  that 

the  eggs  and  of  the  young,  just  as  in    all    these    respects   this    species 

does  the  male  of  the  common  species  comes  imder  our  first  class  of  cases ; 

of  Rhea.  but  Mr.  Sclater  ('  Proc.  Zool.  See' 

2^  For  the  Milvago,  see  '  Zoology  1866,  p.   150)  was  much  surprised 

of    the  Voyage     of    the     Beagle, '  to  observe  that  the  young  of  both 

Birds,  1841,  p.  16.    For  the  Climac-  sexes,  when  about  three  months  old, 

teris  and  night-jar  (Eurostopodus),  resembled  in  their  dark  heads   and 

see  Gould's  '  Handbook  to  the  Birds  necks   the   adult   males,   instead    of 

of  Australia,'  vol.  i.  pp.  602  and  97.  the  adult  females;  so  that  it  would 

The    New  Zealand    shieldrake   (7a-  appear  in  this  case  that  the  females 

dorna  variegata)  offers   a  quite  ano-  have  been  modified,  whilst  the  males 

nialous  case  ;  the  head  of  the  female  and    the    young    have    retained    a 

is  pure  white,  and  her  back  is  redder  former  state  of  plumage. 


480  The  Descent  of  Man,  Part  II. 

males  in  the  last  class.  Mr.  Wallace  believes  that  the  males 
have  had  their  colours  rendered  less  conspicuous  for  the  sake  of 
protection  during  the  period  of  incubation ;  but  the  dilference 
between  the  sexes  in  hardly  any  of  the  foregoing  cases  appears 
sufBciently  great  for  this  view  to  be  safely  accepted.  In  some 
of  the  cases,  the  brighter  tints  of  the  female  are  almost  confined 
to  the  lower  surface,  and  the  males,  if  thus  coloured,  would  not 
have  been  exposed  to  danger  whilst  sitting  on  the  eggs.  It 
should  also  be  borne  in  mind  that  the  males  are  not  only  in  a 
slight  degree  less  conspicuously  coloured  than  the  females,  but  are 
smaller  and  weaker.  They  have,  moreover,  not  only  acquired 
the  maternal  instinct  of  incubation,  but  are  less  pugnacious  and 
vociferous  than  the  females,  and  in  one  instance  have  simpler 
vocal  organs.  Thus  an  almost  complete  transposition  of  the 
instincts,  habits,  disposition,  colour,  size,  and  of  some  points  of 
structure,  has  been  effected  between  the  two  sexes. 

Now  if  we  might  assume  that  the  males  in  the  present  class 
have  lost  some  of  that  ardour  which  is  usual  to  their  sex,  so  that 
they  no  longer  search  eagerly  for  the  females ;  or,  if  we  might 
assume  that  the  females  have  become  much  more  numerous 
than  the  males — and  in  the  case  of  one  Indian  Turnix  the  females 
are  said  to  be  "  much  more  commonly  met  with  than  the 
"  males."  ^*^ — then  it  is  not  improbable  that  the  females  w^ould 
have  been  led  to  court  the  males,  instead  of  being  courted  by 
them.  This  indeed  is  the  case  to  a  certain  extent  with  some 
birds,  as  we  have  seen  with  the  peahen,  wild  turkey,  and  certain 
kinds  of  grouse.  Taking  as  our  guide  the  habits  of  most  male 
birds,  the  greater  size  and  strength  as  well  as  the  extraordinary 
pugnacity  of  the  females  of  the  Turnix  and  emu,  must  mean 
that  they  endeavour  to  drive  away  rival  females,  in  order  to  gain 
possession  of  the  male ;  and  on  this  view  all  the  facts  become 
clear;  for  the  males  would  probably  be  most  charmed  or  ex- 
cited by  the  females  which  were  the  most  attractive  to  them  by 
their  bright  colours,  other  ornaments,  or  vocal  powers.  Sexual 
selection  would  then  do  its  work,  steadily  adding  to  the  attrac- 
tions of  the  females ;  the  males  and  the  young  being  left  not  at 
all,  or  but  little  modified. 

Class  III.  When  the  adult  male  resembles  the  adult  female,  the 
young  of  hoth  sexes  have  a  peculiar  first  plumage  of  thtir  own. — In 
this  class  the  sexes  when  adult  resemble  each  other,  and  dififer 
from  the  young.  This  occurs  with  many  birds  of  many  kinds. 
The  male  robin  can  hardly  be  distinguished  from  the  female, 
but  the  young  are  widely  different,  with  their  mottled  dusky- 
olive  and  brown  plumage.  The  male  and  female  of  the  splendid 
2*^  Jerdon,  '  Birds  of  India,'  vol.  iii.  p.  598. 


Chap.  XVI.      Birds — Young  like  both  Adults.  481 

scarlet  ibis  are  alike,  whilst  the  young  are  brown ;  and  the 
scarlet-colour,  though  common  to  both  sexes,  is  apparently  a 
sexual  character,  for  it  is  not  well  developed  in  either  sex  under 
confinement ;  and  a  loss  of  colour  often  occurs  with  brilliant 
males  when  they  are  confined.  With  many  species  of  herons 
the  young  differ  greatly  from  the  adults;  and  the  summer 
plumage  of  the  latter,  though  common  to  both  sexes,  clearly 
nas  a  nuptial  character.  Young  swans  are  slate-coloured, 
whilst  the  matui-e  birds  are  pure  white;  but  it  would  be 
superfluous  to  give  additional  instances.  These  differences 
between  the  young  and  the  old  apparently  depend,  as  in  the  last 
two  classes,  on  the  young  having  retained  a  former  or  ancient 
state  of  plumage,  whilst  the  old  of  both  sexes  have  acquired  a  new 
one.  When  the  adults  are  bright  coloured,  we  may  conclude 
from  the  remarks  just  made  in  relation  to  the  scarlet  ibis  and  to 
many  herons,  and  from  the  analogy  of  the  species  in  the  first  class, 
that  such  colours  have  been  acquired  through  sexual  selection  by 
the  nearly  mature  males  ;  but  that,  differently  from  what  occurs 
in  the  first  two  classes,  the  transmission,  though  limited  to  the 
same  age,  has  not  been  limited  to  the  same  sex.  Consequently,  the 
sexes  when  mature  resemble  each  other  and  differ  from  the  young. 
Class  IV.  Whtn  the  adult  male  resembles  the  adult  female^  the 
youmj  of  both  sexes  in  their  first  jl  (image  resemble  the  adults. — In  this 
class  the  young  and  the  adults  of  both  sexes,  whether  brilliantly 
or'  obscurely  coloured,  resemble  each  other.  Such  cases  are,  I 
think,  more  common  than  those  in  the  last  class.  We  have  in 
England  instances  in  the  kingfisher,  some  woodjDeckers,  the  jay, 
magpie,  crow,  and  many  small  dull-coloured  birds,  such  as  the 
hedge-warbler  or  kitty-wren.  But  the  similarity  in  plumage 
between  the  young  and  the  old  is  never  complete,  and  graduates 
away  into  dissimilarity.  Thus  the  young  of  some  members  of 
the  kingfisher  family  are  not  only  less  vividly  coloured  than  the 
adults,  l)ut  many  of  the  feathers  on  the  lower  surface  are  edged 
with  brown,^^ — a  vestige  probably  of  a  former  state  of  the 
]:»lumage.  Frequently  in  the  same  group  of  birds,  even  within 
the  same  genus,  for  instance  in  an  Australian  genus  of  parro- 
keets  (Platycercus),  the  young  of  some  species  closely  resemble, 
whilst  the  young  of  other  species  differ  considerably,  from  their 
parents  of  both  sexes,  which  are  alike.^^  Both  sexes  and  the 
young  of  the  common  jay  are  closely  similar ;  but  in  the  Canada 
jay  {Ferisoreus  canadensis)  the  young  differ  so  much  from  their 
parents  that  they  were  formerly  described  as  distinct  species.^^ 

"  Jerdon,  '  Birds  of  India,'  vol.  i.  ^s  Qould,  Ibid.  vol.  ii.  pp.  37,  46. 

pp.  222,  228.  Gould's  '  Handbook  to  56. 

the  Birds  of  Australia,'  vol.  i.  pp.  ^9  Audubon,  '  Ornith.  Bio£raphy,' 

124,  130.  vol.  ii.  p.  5o. 


482  The  Descent  of  Man.  Part  II. 


I  may  remark  before  proceeding  that,  under  the  present  and 
next  two  classes  of  cases,  the  facts  are  so  complex  and  the  con- 
clusions so  doubtful,  that  any  one  who  feels  no  especial  interest 
in  the  subject  had  better  pass  them  over. 

The  brilliant  or  conspicuous  colours  wliich  climacterifce  many  birds 
in  the  present  class,  cai  rarely  or  never  be  of  service  to  them  as  a  pro- 
tection ;  so  that  they  have  probably  been  gained  by  the  males  through 
st-xual  seiecUon,  and  then  transferred  to  the  females  and  the  young. 
It  is,  however,  possible  that  the  males  may  have  selected  the  more 
attractive  females ;  and  if  these  transmitteil  their  characters  to  their 
offspring  of  both  sexes,  the  same  results  would  follow  as  fiora  the 
selectioa  of  the  more  attractive  maU  s  by  the  females.  But  there  is 
evidence  that  this  contingency  has  rarely,  if  ever,  occurred  in  any  of 
those  groups  of  birds  in  which  the  sexes  are  generally  alike ;  for,  if 
even  a  few  of  the  successive  variations  had  failed  to  be  transmitted  to 
both  sexes,  the  females  would  have  slightly  exceeded  the  males  in 
beauty.  Exactly  the  reverse  occurs  under  nature ;  for,  in  almost  every 
large  group  in  which  the  sexes  generally  resemble  each  other,  the 
males  of  some  few  species  are  in  a  slight  degree  more  brightly  coloured 
than  the  females.  It  is  again  possible  that  the  females  may  have 
selected  the  more  beautiful  males,  these  males  having  reciprocally 
selected  the  more  beautiful  females;  but  it  is  doubtful  whether  this 
double  process  of  selection  would  be  likely  to  occur,  owing  to  the 
greater  eagerness  of  one  sex  than  the  other,  and  whether  it  would  be 
more  efficient  than  selection  on  one  side  alone.  It  is,  therefore,  the 
most  probable  view  that  sexual  selection  has  acted,  in  the  present  class, 
as  far  as  ornamental  characters  are  concerned,  in  accordance  with  the 
geneial  rule  throughout  the  animal  kingdom,  that  is,  on  the  males  ; 
and  that  these  have  transmitted  their  gradually-acquired  colours, 
either  equally  or  almost  equally,  to  their  ottspring  of  both  sexes. 

Another  point  is  more  doubtful,  namely,  whether  the  successive 
variations  first  appeared  in  the  males  after  they  had  become  nearly 
mature,  or  whilst  quite  young.  In  either  case  sexual  selection  must 
have  acted  on  the  male  when  he  had  to  compete  with  rivals  for  the 
possession  of  the  female  ;  and  in  both  cases  the  characters  thus  acquired 
have  been  transmitted  to  both  sexes  and  all  ages.  But  these  characters, 
if  acquired  by  the  males  when  adult,  may  have  been  transmitted  at 
first  to  the  adults  alone,  and  at  some  subsequent  period  transferred  to 
the  young.  For  it  is  known  that,  when  the  law  of  inheritance  at 
corresponding  ages  fails,  the  ofispring  often  inherit  characters  at  an 
earlier  age  than  that  at  which  they  first  appeared  in  their  parents.^" 
Cases  apparently  of  this  kind  have  been  observed  with  birds  in  a  state 
of  nature.  For  instance  Mr.  Blyth  has  seen  specimens  of  Lanius  rufus 
and  of  Colymhus  glacialis  which  had  assumed  whilst  young,  in  a  quite 
anomalous  manner,  the  adult  plumage  of  their  parents.^^  Again,  the 
young  of  the  common  swan  {Cygnus  olor)  do  not  cast  off  their  dark 
feathers  and  become  white  until  eighteen  months  or  two  years  old  ;  but 
Dr.  F.  Forel  has  described  the  case  of  three  vigorous  young  birds,  out 
of  a  brood  of  four,  which  were  born  pure  white.     These  young  birds 


^0  'Variation    of     Animals    and  "  Charlesworths' *  Mag.  of  Nat. 

Plants  under  Domestication,'  vol.  ii.      Hi  '-,'  vol.  i,  1837,  pp.  305,  306, 
p.  79 


Chap.  XVI.      Birds—  Voting  like  both  Adults.  483 

were  not  albinoes,  as  shewn  by  the  colour  of  their  beaks  and  legs,  wliich 
nearly  retembled  the  same  parts  in  the  adults.^'^ 

It  may  be  worth  while  to  illustrate  the  above  three  modes  by  which, 
in  the  present  class,  the  two  sexes  and  the  young  may  have  come  to 
res(  mble  each  other,  by  the  curious  case  of  the  genus  I'asser.^^  In  the 
house-sparrow  (P.  rfome.vY/cH6)  the  male  differs  much  from  the  female 
and  from  the  young.  'Jlie  young  and  the  females  are  alike,  and 
resemble  to  a  large  extent  both  sexes  and  the  young  of  the  siJanow  of 
Palestine  (P.  hrachydactylus),  as  well  as  of  some  allied  species.  Wo 
may  therefore  assume  that  the  female  and  young  of  the  house-sparrow 
approximately  shew  us  the  jdumage  of  the  progenitor  of  the  genus. 
Now  with  the  tree-sparrow  (P.  montanus)  both  sexes  and  the  young 
closely  resemble  the  male  of  the  house-sparrow ;  so  that  they  have  ail 
been  moditied  in  the  same  manner,  and  all  depart  from  the  typical 
colouring  of  their  early  progenitor.  This  may  have  been  effected  by 
a  male  ancestor  of  the  tree-sparrow  having  varied,  firstly,  when  nearly 
mature  ;  or,  secondly,  whilst  quite  young,  and  by  having  in  either  case 
transmitted  his  modified  plumage  to  the  females  and  the  young;  or, 
thirdly,  he  may  have  varied  when  adult  and  transmitted  his  plumage 
to  both  adult  sexes,  and,  owing  to  the  failure  of  the  law  ot  inheritance 
at  corresponding  ages,  at  some  subsequent  period  to  his  young. 

It  is  impossible  to  decide  which  of  these  three  modes  has  generally 
prevailed  through  >ut  the  present  class  of  cases.  That  the  males  varied 
whilst  young,  and  transmitted  their  variations  to  tiieiroftspring  of  both 
sexes,  is  the  most  probable.  I  may  here  add  that  I  have,  with  little 
success,  endeavoured,  by  consulting  various  works,  to  decide  how  far 
the  period  of  variation  in  birds  has  generally  determined  the  transmis- 
sion of  characters  to  one  sex  or  to  both.  The  two  rules,  often  referred 
to  (namely,  that  variations  occurring  late  in  life  are  transmitted  to  one 
and  the  same  sex,  whilst  those  which  occur  early  in  life  are  transmitted 
to  both  sexes),  apparently  hold  good  in  the  first,^^  second,  and  fourth 
classes  of  cases ;  but  they  fail  in  the  third,  often  in  the  fifth,^^  and  in 
the  sixth  small  class.  They  apply,  however,  as  far  as  I  can  judge,  to 
a  considerable  majority  of  the  species ;  and  we  must  not  forget  the 
striking  generalisation  by  Dr.  W.  Marshall  with  respect  to  the  protu- 
berances on  the  heads  of  bu'ds.    Whether  or  not  the  two  rules  generally 


^2  '  Bulletin  de  la  Soc.  Vaudoise  Audubon,  '  Ornith.   Biography,'  vol. 

des  So.  Nat.'  vol.  x.   1869,  p.   132.  i.  p.  233,  280,378.)    The  Harlequin 

The    young    of   the    Polish     swan  duck  takes  three  years  (ibid.  vol.  iii. 

Cygnus  immutahilis  of  Yarrell,  are  p.   614).     The  male    of    the    Gold 

always  white ;  but  this  species,  as  pheasant,     as    I     hear    from      Mr. 

Mr.  Sclater  informs  me,  is  believed  Jenner  Weir,  can   be   distinguished 

to  be  nothing  more  than  a  variety  from  the  female  when  about  three 

of     the    domestic    swan     (^Cygnus  months  old,  but  he  does  not  acquire 

olor).  his  full  splendour  until  the  end   of 

I  am  indebted  to    Mr.    Blyth  the    September    m    the    following 


for   information    in    reaiard  to  this 


year. 


genus.     The    sparrow  of   Palestine  ^5  ^hus  the  76ts  tantalus  and  Grus 

belongs  to  the  sub-genus  Petronia.  americanus    take    four    years,    the 

2*  For     instance,    the     males    of  Flamingo    several    years,   and    the 

Tanngra  a'stiva  and  Fringilla  cyanea  Ardea  Ivdovkana  two  years,  before 

require   three    years,  the    mala   of  they  acquire  their  perfect  plumage. 

Fringilla  ciris  four   years,  to   com-  See  Audubon,  ibid.  vol.  i.  p.  121  j 

piste  their  beautiful  plumage.    (See  vol.  iii.  pp.  133,  139,  211. 


484  The  Descent  of  Man.  Part  II 

hold  g:ood,  we  may  conclude  from  the  facts  given  in  the  eighth  chapter, 
that  the  period  of  variation  is  one  important  element  in  determining  the 
form  of  transmission. 

With  birds  it  is  difficult  to  decide  by  what  standard  we  ought  to  jud^e 
of  the  earliness  or  lateness  of  the  period  of  variation,  whether  by  the 
age  in  reference  to  the  duration  of  life,  or  to  the  power  of  reproduction, 
or  to  tlie  number  of  moults  through  which  the  species  passes.  Tlie 
moulting  of  birds,  even  within  the  same  family,  sometimes  dilf  rs 
much  without  any  assignable  cause.  Some  birels  moult  so  early,  that 
nearly  all  the  body  featht-rs  are  cast  oif  before  the  first  wing-featht  rs 
are  fully  grown;  and  we  cannut  believe  that  this  was  the  primordial 
state  of  things.  When  the  period  of  moulting  has  been  accelerated, 
the  age  at  which  the  colours  of  the  a  Uilt  plumage  are  first  developed 
will  falsely  appear  to  us  to  be  earlier  tlian  it  really  is.  This  may  be 
illustrated  by  the  practice  folli>we  I  by  some  bird-fauciers,  who  pull 
out  a  few  feathers  from  the  breast  of  nestling  bullfinches,  and  from  the 
head  or  neck  of  young  gold-pheasants,  in  order  to  ascertain  their  sex ; 
for  in  the  male;",  these  feathers  are  immediately  replaced  by  coloured 
ones.'^  The  actual  duration  of  life  is  known  in  but  few  birds,  so  that 
we  can  hardly  judge  by  this  standard,  And,  with  reference  to  the 
period  at  which  the  power  of  reproduction  is  gained,  it  is  a  remark- 
able fact  that  various  birds  occasionally  breed  whiLt  retaining  their 
immature  plumage.^^ 

Tlie-fact  of  birds  breeding  in  their  immature  plumage  seems  opposed 
to  the  belief  that  sexual  selection  has  played  as  important  a  part,  as  1 
believe  it  Ins,  in  giving  oruiimental  colours,  plumes,  &c.,  to  the  males, 
and,  by  means  of  equal  transmission,  to  the  females  of  many  species. 
The  objection  would  be  a  valid  one,  if  the  younger  and  less  ornamented 
males  were  as  successful  in  winning  females  and  propagating  their 
kind,  as  the  (jlder  and  more  beautiful  males.  But  we  have  no  reason 
to  suppose  that  this  is  the  case.  Audubon  speaks  of  the  breeding  of 
the  immature  males  of  Ihis  tantalus  as  a  rare  event,  as  does  Mr. 
Swinhoe,  in  regard  to  the  immature  males  of  Oriolus.^®  If  the  young 
of  any  species  in  their  immature  plumage  were  more  successful  in 
winning  partners  than  the  adults,  the  adult  plumage  would  probably 


^^  Mr.  Blyth,  in   Charlesworth's  58):    but   Mr.   Blyth    informs    me 

*  Mag.  of  Nat.  Hist.'  vol.  i.  1837,  p.  that  certain  herons  apparently  are 

300.     Mr.    Bartlett    has    informed  dimorphic,  for  white  and  coloured 

me  in  regard  to  gold-pheasants.  individuals  of  the  same  age  may  be 

^'  I  have    noticed  the  following  observed.     The      Harlequin      duck 

cases    in    Audubon's   'Ornith.    Bio-  (J»as  A/sfrjomca,  Linn.)  takes  three 

graphy.'     The  redstart  of  America,  years  to  acquire  its   full  plumage, 

(3Iuscapica  ruticilla,  vol.  i.  p.  203).  though    many  birds    breed     in  the 

The  Ibis  tantalus  takes  four  years  to  second  year  (vol.  iii.  p.  614).     The 

come  to   full   maturity,  but  some-  White-headed  Eagle  {Falco  leucoce- 

times  breeds  in  the  second  year  (vol.  phalus,  vol.  iii.  p.  210)  is  likewise 

iii.  p.  133).     The   Grus  americanus  known  to  breed   in    its    immature 

takes   the    same    time,   but    breeds  state.     Some  species  of  Oriolus  (ac- 

before    acquiring  its   full   plumage  cording    to     Mr.    Blyth     and     Mr. 

(vol.  iii.  p.  211).      The    adults    of  Swinhoe,  in    'Ibis,'    July   1863,    p. 

Ardea    cxrulea    are    blue,    and   the  68)    likewise     breed    before     they 

young  white;  and   white,  mottled,  attain  their  full  plumage, 

and  mature  blue  birds    may  all  be  ^^  See  the  last  foot-note. 
seen  breeding  together  (vol.  iv.  p. 


Chap.  XYI.      Birds —  Voting  like  both  Adults.  485 

soon  be  lost,  as  the  males  would  prevail,  wliich  retained  their  immaturo 
dress  for  the  longest  period,  and  thus  the  character  of  the  species 
would  ultimately  be  modified.^"  If,  on  the  other  hand,  the  young  never 
succeeded  in  obtaining  a  female,  the  habit  of  early  reproduction  would 
perhaps  be  sooner  or  later  eliminated,  from  being  superfluous  and 
entailing  waste  of  power. 

The  plumage  of  certain  birds  goes  on  increasing  in  beauty  during 
many  years  after  they  are  fully  mature ;  this  is  the  case  with  the  train 
of  the  peacock,  with  some  of  the  birds  of  paradise,  and  with  the  crest 
and  plumes  of  certain  herons,  for  instance,  the  Ardea  liulovicana.*'^ 
But  it  is  doubtful  wliether  the  continued  development  of  such  feathers 
is  the  result  of  the  selection  of  successive  beneficial  variations  (though 
this  is  the  most  probable  view  with  birds  of  paradise)  or  merely  of  con- 
tinuous growth.  Most  fishes  continue  increasing  in  size,  as  long  as  they 
are  in  good  health  and  have  plenty  of  food;  and  a  somewhat  similar 
law  may  pi  evnil  with  the  plumes  of  birds. 

Class  V.  When  the  adults  of  both  sexes  have  a  distinct  winter  and 
summer  plumage,  lohether  or  not  the  male  differs  from  the  female,  the 
young  resemble  the  adidts  of  both  sexes  in  their  lointer  dress,  or  much 
more  rarely  in  their  summer  dress,  or  they  resemble  the  females  alone. 
Or  the  young  may  have  an  intermediate  character ;  or,  again,  they  may 
differ  greatly  from  the  adults  in  both  their  seasonal  plumages. — The  cases 
in  this  class  are  singularly  complex;  nor  is  this  surprising,  as  they 
depend  on  inheritance,  limited  in  a  greater  or  less  degree  in  three 
ditt'erent  ways,  namely,  by  sex,  age,  and  the  season  of  the  year.  In 
some  eases  the  individuals  of  the  same  species  pass  through  at  least 
five  distinct  states  of  plumage.  With  the  species,  in  which  the  male 
differs  from  the  female  during  the  summer  season  alone,  or,  which  is 
rarer,  during  both  si  asons,''^  the  young  generally  resemble  the  females, 
— ^as  with  the  so-called  goldfinch  of  North  America,  and  apparently 
with  the  splendid  Maluri  of  Australia.*^  With  those  species,  the  sexes 
of  which  are  alike  during  both  the  summer  and  winter,  the  young  may 


'*  Other    animals,   belonging    to  ^o  Jerdon,  'Birds   of  India,'  vol. 

quite   distinct    classes,    are     either  iii.    p.    507,    on    the    peacock.     Dr. 

habitually    or   occasionally  capable  Marshall,  thinks  that  the  older  and 

of  breeding  before  they  have  fully  more    brilliant    males    of    birds    of 

acquired     their    adult     characters,  paradise,   have  an    advantage    over 

This    is    the    case    with  the   young  the    younger  males;  see  'Archives 

males   of  the  salmon.     Several  am-  Neerlandaises,'  torn.   vi.   1871:. —  On 

phibians  have  been  known  to  breed  Ardea,   Audubon,  ibid.    vol.    iii.    p. 

whilst  retaining  their  larval  struc-  139. 

ture.      Fritz     Muller     has     shewn  *^  For  illustrative  cases  see  vol. 

('  Facts  and  Arguments  for  Darwin,'  iv.  of  Macgillivray's   '  Hist.     Brit. 

Eng.   trans.    1869,  p.   79)  that  the  Birds  ;' on  Tringa,  &c.,  pp.  229,  271  ; 

males  of  several    amphipod  crusta-  on    the    Machetes,  p.   172 ;  on    the 

ceans  become  sexually  mature  whilst  C/iaradrius  hiaticula,  p.  118  ;  on  the 

young ;  and  I  infer  that  this    is   a  Charadrius  pluvialis,  p.  94. 

case  of  premature  breeding,  because  ^"^  For  the  goldfinch  of  N.America, 

they  have  not  as  yet  acquired  their  Fringilla  tristis,  Linn.,  see  Audubon, 

fully-developed  claspers.     All  such  '  Ornith.    Biography,' vol.  i.  p.  172. 

facts     are     highly     interesting,     as  For  the   Maluri,  Gould's  '  Handbook 

bearing    on    one    means    by    which  to  the  Birds  of  Australia,*  vol.  i.  p. 

species  may  undergo  great  raodifi-  318. 
cations  of  character. 


4^6  The  Descent  of  Man.  Part  II. 

resemble  the  adults,  firsth%  in  their  winter  dsess ;  secondly,  and  this  is 
of  much  rarer  occurrence,  in  their  summer  dre^s;  thirdly,  they  may  be 
iutermediate  between  these  two  states ;  and,  fourthly,  they  may  dift'er 
greatly  from  the  adults  at  all  seasons.  We  have  an  instance  of  the 
first  of  these  four  cases  in  one  of  the  egrets  of  India  {Buphus  coro- 
mandus),  in  which  the  young  and  the  adults  of  both  sexes  are  white 
duringthe  winter,  the  adults  becoming  golden-buff  during  the  summer. 
With  the  gaper  {Anastomus  oscitans)  of  India  we  have  a  similar  case, 
but  the  colours  are  rever&ed :  for  the  young  and  the  adults  of  both 
sexes  are  grey  and  black  during  the  winter,  the  adults  becoming  white 
during  the  summer.*^  As  an  instance  of  the  second  case,  the  young  of 
the  razor-bill  {Alca  torda,  Linu.),  in  an  early  state  of  plumage,  are 
coloured  like  the  adults  during  the  summer;  and  the  young  of  the 
white-crowned  sparrow  of  North  America  (Fringilla  leucnphrys),  as  soon 
!is  fledged,  have  elegant  white  stripes  on  their  heads,  which  are  lost  by 
the  young  and  the  old  during  the  winter.^*  With  respect  to  the  third 
case,  namely,  that  of  the  young  having  an  intermediate  character  between 
the  summer  and  winter  adult  plumages,  Yarrell"  insists  that  this 
occurs  with  many  waders.  Lastly,  in  regiird  to  the  young  differing 
greatly  from  both  sexes  in  their  adult  summer  and  winter  plumages, 
this  occurs  with  some  herons  and  egrets  of  North  America  and  India, — 
the  young  alone  being  white. 

I  will  make  only  a  few  remarks  on  these  complicated  cases.  When 
the  young  resemble  the  females  in  their  summer  dress,  or  the  adults  of 
b.ith  sexes  in  their  winter  dress,  the  cases  differ  from  those  given  under 
Classes  I.  and  III.  only  in  the  characters  originally  acquired  by  tUe 
males  during  the  breeding-season,  having  been  limited  in  their  trans- 
misiion  to  the  corresponding  season.  When  the  adults  have  a  distinct 
summer  and  winter  plumage,  and  the  young  differ  from  both,  the  ca-e 
is  more  difficult  to  understand.  We  may  admit  as  probable  that  the 
young  have  retained  an  ancient  state  of  plumai^e  ;  we  can  account  by 
sexual  selection  for  the  summer  or  nuptial  plumage  of  the  adults,  but 
how  are  we  to  account  for  their  distinct  winter  plumage?  If  we  couid 
admit  that  this  plumage  serves  in  all  cases  as  a  protection,  its  ac- 
quirement would  be  a  simple  affair ;  but  there  seems  no  good  reason 
for  this  admission.  It  may  be  suggested  that  the  widely  different 
conditions  of  life  during  the  winter  and  summer  have  acted  in  a  direct 
manner  on  the  plumage;  this  may  have  had  some  effect,  but  I  have 
not  much  confidence  in  so  great  a  difference  as  we  sometimes  see 
between  the  two  plumages,  having  been  thus  caused.  A  more  probable 
explanation  is,  that  au  ancient  style  of  plumage,  partially  modified 
tliroujih  the  transference  of  some  characters  from  the  summer  plumage, 
has  been  retained  by  the  adults  during  the  winter.  Finally,  all  the 
cases  in  our  present  class  apparently  depend  on  characters  acquired  by 
the  adult  males,  having  been  variously  limited  in  theii*  transmission 
according  to  age,  season,  and  sex;  but  it  would  not  be  worth  while  to 
attempt  to  follow  out  these  complex  relations. 


*^  I  am   indebted   to    Mr.   Blyth  On   the    Fringilla    leucophryS,    Au- 

for  information  as  to  the  Buphus ;  dubon,  ibid.  vol.  ii.  p.  89.     I  shall 

see    also    Jerdon,   '  Birds  of  India,'  have  hereafter  to  refer  to  the  young 

vol.  ill.  p.  749,     On  the  Anastomus,  of  certain  herons  and  egrets  being 

see  Blyth,  in  'Ibis,'  1867,  p.  173.  white. 

*■*  On  the  Alca,  see  Macgillivrav,  *'"  '  Historv  of  British  Birds,'  vol 

'  Hist.  Brit.  Birds,'  vol.  v.  p.  347.  i.  1839,  p.  159. 


Chap,  XVI.     Birds —  Young  like  both  Adults,  487 

Class  VC.  The  young  in  their  first  plumage  differ  from  each  other 
according  to  sex  ;  the  young  males  resembling  more  or  less  closely  the  adult 
males,  and  the  young  females  more  or  less  closely  the  adult  females. — 
The  cases  in  the  present  class,  though  occurring  in  various  groups,  are 
not  numerous;  yet  it  seems  the  most  natural  thing  that  the  young 
should  at  first  somewhat  resemble  the  adults  of  the  same  sex,  and 
gradually  become  more  and  more  like  tliem.  The  adult  male  blackcap 
{Sylvia  atricapilla)  has  a  black  head,  tliat  of  the  female  being  reddisli- 
brown ;  and  I  am  informed  by  Mr.  Blyth,  that  the  young  of  b^tli  sexes 
can  be  distinguished  by  this  character  even  as  nestlings.  In  the  family 
of  thrushes  an  unusual  number  of  similar  cases  have  been  noticed  ;  thus, 
the  male  blackbird  {Turdus  merula)  can  be  distinguished  in  the  nest 
from  the  female.  Tlie  two  sexes  of  tlic  mocking  bird  (^Turdus  polyglottus, 
Linn.)  difler  very  little  from  each  other,  yet  the  males  can  easily  be. 
distinguished  at  a  very  eaily  age  from  the  females  by  shewing  more 
pure  white.''^  The  males  of  a  forest-thrush  and  of  a  rock-thrush 
(^Orocetes  erythrogastra  and  Petrocincla  cyanea)  have  much  of  their 
plumage  of  a  fine  blue,  whilst  the  females  are  brown  ;  and  the  nestling 
males  of  both  species  have  their  main  wing  and  tail-feathers  edged  with 
blue,  whilst  those  of  the  female  are  edged  with  brown.''^  In  the  young 
blackbird  the  wing  feathers  assume  their  mature  character  and  become 
black  after  the  others  ;  on  the  otlier  hand,  in  the  two  species  just  named 
the  wing- feathers  become  blue  before  tlie  others.  The  most  probable 
view  with  reference  to  the  cases  in  the  present  class  is  that  the  males, 
differently  from  what  occurs  in  Class  T;,  have  transmitted  their  colours 
to  their  male  offspring  at  an  earlier  age  than  that  at  which  they  were 
first  acquired ;  for,  if  the  males  had  varied  whilst  quite  young,  their 
characters  would  probably  have  been  transmitted  to  both  sexes.** 

In  Aithurus  polytmus,  a  humming-bird,  the  male  is  splenditUy  coloured 
black  and  green,  and  two  of  the  tail-feathers  are  immensely  lengthened  ; 
the  female  has  an  ordinary  tail  and  inconspicuous  colours;  how  the 
young  males,  instead  of  resembling  the  adult  female,  in  accordance 
with  the  common  rule,  begin  from  the  first  to  assume  the  colours  proper 
to  their  sex,  and  their  tail-feathers  soon  become  elongated.  I  owe  this 
information  to  Mr.  (tIouIiI,  who  has  given  me  the  following  more  striking 
and  as  yet  unpublished  case.  Two  liumming-birds  belonging  to  the 
genus  Eustephanus,  both  beautifully  coloured,  inhabit  the  small  island 
of  Juan  Fernandez,  and  have  always  been  ranked  as  specifically  dis- 
tinct. But  it  has  lately  been  ascertained  that  tlie  one,  which  is  of  a 
rich  chesnut  brown  colour  with  a  golden-red  head,  is  the  male,  whiUt 
the  other,  which  is  elegantly  variegated  with  green  and  white  with  a 
metallic-green  head  is  the  female.      Now  the   young  from  the  first 


*^  Audubon, '  Ornith.  Biography,'  '  Ornith.  Biography,'  vol.  iv.  p.  392), 

vol.  i.  p.  113.  and  so  it  is  with  the  nestlings  of  a 

•"  Mr.  C.  A.  Wright,    in  'Ibis,'  blue  nu.th.atch,  Bendrophila  frontalis 

vol.  vi.  1864,  p.  65.     Jerdon, '  Birds  of  India  (Jerdon,  'Birds  of  India,' 

of  India,' vol.  i.  p.  515.     See  also  on  vol.    i.    p.    389).     Mr,    Blyth     also 

the    blackbird,    Blyth     in   Charles-  informs   me   that   the   sexes   of  the 

worth's  '  Mag.  of  Nat.  History,'  vol.  stonechat,     Saxicola     rubicola,     are 

i.  1837,  p.  113.  distinguishable  at  a  very  early  age. 

■**  The  following  additional  cases  Mr.  Salvin  gives  ('Proc.  Zoolog.  Soc' 

maybe  mentioned;  the  young  males  1870,   p.   206),  the  case  of  a  hum- 

of  Tanagra rubra cixnhe distingnished  ming-bird,  like  the  following  one  of 

from   the  young  females  (Audubon,  Eustephanus. 


The  Descent  of  Man.  Part  IX 

somewhat  resemble  the  adults  of  the  corresponding  sex.,  the  resemblance 
gradually  becoming  more  and  more  complete. 

In  considering  this  last  case,  if  as  before  we  fake  tlie  plumage  of  the 
young  as  our  guide,  it  w  oidd  appear  that  both  sexes  have  been  rendered 
beautiful  indepenilently ;  and  not  that  one  sex  has  partially  transferred 
its  beauty  to  the  other.  The  male  apparently  has  acquired  liis  bright 
colours  through  sexual  selection  in  the  same  manner  as,  for  instance, 
the  peacock  or  pheasant  in  our  lirst  class  of  cases ;  and  the  female  in 
the  same  manner  as  the  female  Ehyncha>a  or  Turnix  in  our  second 
class  of  cases.  But  there  is  mucli  difficulty  in  understanding  how  this 
could  have  been  effected  at  the  same  time  with  the  two  sexes  of  the 
same  species.  Mr.  Salvin  states,  as  we  have  seen  in  the  eighth  chapter, 
that  with  certain  humming-birds  the  males  greatly  exceed  the  females 
in  number,  whilst  with  other  species  inhabiting  the  same  country  the 
females  greatly  exceed  the  males.  If,  then,  we  might  assume  that 
during  some  former  lengthened  period  the  males  of  the  Juan  Fernandez 
species  had  greatly  exceeded  the  females  in  number,  but  that  during 
another  lengthened  period  the  females  had  far  exceeded  the  males,  we 
could  understand  liow  the  males  at  one  time,  and  the  females  at 
another,  mi-ht  have  been  rendered  beautiful  by  tlie  selection  of  the 
brighter-coloured  individuals  of  either  sex ;  both  sexes  transmitting 
.  their  chaiacters  to  their  young  at  a  rather  earlier  age  than  usual. 
Whether  this  is  the  true  explanation  I  will  not  pretend  to  say ;  but  the 
case  is  too  remarkable  to  be  passed  over  without  notice. 

We  have  now  seen  in  all  six  classes,  that  an  intimate  relation 
exists  between  the  plumage  of  the  young  and  the  adults,  either 
of  one  sex  or  both.  These  relations  are  fairly  well  explained 
on  the  principle  that  one  sex — this  being  in  the  great  majority 
of  cases  the  male — first  acquired  through  variation  and  sexual 
selection  bright  colours  or  other  ornaments,  and  transmitted 
them  in  various  ways,  in  accordance  with  the  recognised  laws  of 
inheritance.  Why  variations  have  occurred  at  different  periods 
of  life,  even  sometimes  with  species  of  the  same  group,  we  do  not 
know,  but  with  respect  to  the  form  of  transmission,  one  important 
determining  cause  seems  to  be  the  age  at  which  the  variations 
first  appear. 

From  the  principle  of  inheritance  at  corresponding  ages,  and 
from  any  variations  in  colour  which  occurred  in  the  males  at  an 
early  age  not  being  then  selected— on  the  contrary  being  often 
eliminated  as  dangerous — whilst  similar  variations  occurring 
at  or  near  the  period  of  reproduction  have  been  preserved,  it 
follows  that  the  plumage  of  the  young  will  often  have  been  left 
unmodified,  or  but  little  modified.  We  thus  get  some  insight 
into  the  colouring  of  the  progenitors  of  our  existing  species.  In 
a  vast  number  of  species  in  five  out  of  our  six  classes  of  cases, 
the  adults  of  one  sex  or  of  both  are  bright  coloured,  at  least 
during  the  breeding-season,  whilst  the  young  are  invariably  less 
brightly  coloured  than  the  adults,  or  are  quite  dull  coloured ; 


Chap.  XVI.      Birds — Colour  and  Protectioji.  489 

for  no  instance  is  known,  as  far  as  I  can  discover,  of  ilie  young 
of  dull-coloured  species  displaying  bright  colours,  or  of  tlie  young 
of  bright-coloured  species  being  more  "brilliant  than  tlieir  parents. 
In  the  fourth  class,  however,  in  which  the  young  and  the  old 
resemble  each  other,  there  are  many  sj^ccius  (though  by  no 
means  all),  of  which  the  young  are  bright-coloured,  and  as  these 
form  whole  groups,  we  may  infer  that  their  early  progenitors 
were  hkewise  bright.  With  this  exception,  if  we  look  to  the 
birds  of  the  world,  it  appears  that  their  beauty  has  been  much 
increased  since  that  period,  of  which  their  immature  plumage 
gives  us  a  partial  record. 

On  the  Colour  of  the  Plumage  in  nlation  to  P7'oiection. — It  will 
have  been  seen  that  I  cannot  follow  Mr.  Wallace  in  the  belief 
that  dull  colours,  when  confined  to  the  females,  have  been  in 
most  cases  specially  gained  for  the  sake  of  protection.  There 
can,  however,  be  no  doubt,  as  formerly  remarked,  that  both 
sexes  of  many  birds  have  had  their  colours  modified,  so  as  to 
escape  the  notice  of  their  enemies ;  or  in  some  instances,  so  as  to 
approach  their  prey  unobserved,  just  as  owls  have  had  their 
plumage  rendered  soft,  that  their  flight  may  not  be  over- 
heard. Mr.  Wallace  remarks  ^^  that  "  it  is  only  in  the  tropics, 
"  among  forests  which  never  lose  their  foliage,  that  we  find 
"  whole  groups  of  birds,  whose  chief  colour  is  green."  It  will  be 
admitted  by  every  one,  who  has  ever  tried,  how  difficult  it  is  to 
distinguish  parrots  in  a  leaf-covered  tree.  Nevertheless,  we 
must  remember  that  many  parrots  are  ornamented  with  crimson, 
blue,  and  orange  tints,  which  can  hardly  be  protective.  Wood- 
peckers are  eminently  arboreal,  but  besides  green  species,  there 
are  many  black,  and  black-and-white  kinds — all  the  species 
being  apparently  exposed  to  nearly  the  same  dangers.  It  is 
therefore  probable  that  with  tree-haunting  birds,  strongly-pro- 
nounced colours  have  been  acquired  through  sexual  selection, 
but  that  a  green  tint  has  been  acquired  oftener  than  any  other, 
from  the  additional  advantage  of  protection. 

In  regard  to  birds  which  live  on  the  ground,  every  one  admits 
that  they  are  coloured  so  as  to  imitate  the  surrounding  surface, 
How  difficult  it  is  to  see  a  partridge,  snipe,  woodcock,  certain 
plovers,  larks,  and  night-jars  when  crouched  on  ground.  Animals 
inhabiting  deserts  oifer  the  most  striking  cases,  for  the  bare 
surface  afibrds  no  concealment,  and  nearly  all  the  smaller  quad- 
rupeds, reptiles,  and  birds  depend  for  safety  on  their  colours. 
Mr.  Tristram  has  remarked  in  regard  to  the  inhabitants  of  the 
Sahara,  that  all  are  protected  by  their  "isabelline  or  sand- 
"  '  Westminster  Review,'  July,  1867,  p.  5. 


490  ^/^^  Descent  of  Man.  Part  IT. 

"  colour."  ^0  Calling  to  my  recollection  the  desert-birds  of  South 
America,  as  well  as  most  of  the  ground-birds  of  Great  Britain, 
it  appeared  to  me  that  both  sexes  in  such  cases  are  generally 
coloured  nearly  alike.  Accordingly,  I  ai)plied  to  Mr.  Tristram 
with  respect  to  the  birds  of  the  Sahara,  and  he  has  kindly 
given  me  the  following  information.  There  are  twenty-six 
species  belonging  to  fifteen  genera,  which  manifestly  have  their 
plumage  coloured  in  a  protective  manner ;  and  this  colouring  is 
all  the  more  striking,  as  with  most  of  these  birds  it  differs  from 
that  of  their  congeners.  Both  sexes  of  thirteen  out  of  the 
twenty-six  species  are  coloured  in  the  same  manner  ;  but  these 
belong  to  genera  in  which  tliis  rule  commonly  prevails,  so  that 
they  tell  us  nothing  about  the  protective  colours  being  the  same 
in  both  sexes  of  desert-birds.  Of  the  other  thirteen  species,  three 
belong  to  genera  in  which  the  sexes  usually  differ  from  each 
other,  yet  here  they  have  the  sexes  alike.  In  the  remaining  ten 
species,  the  male  differs  from  the  female ;  but  the  difference  is 
confined  chiefly  to  the  under  surface  of  the  jDlumage,  w^hich  is 
concealed  when  the  bird  crouches  on  the  ground ;  the  bead  and 
back  being  of  the  same  sand-coloured  hue  in  the  two  sexes.  So 
that  in  these  ten  species  the  upper  surfaces  of  both  sexes  have 
been  acted  on  and  rendered  alike,  through  natural  selection,  for 
the  sake  of  protection ;  whilst  the  lower  surfaces  of  the  males 
alone  have  been  diversified,  through  sexual  selection,  for  the 
sake  of  ornament.  Here,  as  both  sexes  are  equally  well  pro- 
tected, we  clearly  see  that  the  females  have  not  been  prevented 
by  natural  selection  from  inheriting  the  colours  of  their  male 
parents;  so  that  we  must  look  to  the  law  of  sexually-limited 
transmission. 

In  all  parts  of  the  world  both  sexes  of  many  soft-billed  birds, 
especially  those  which  frequent  reeds  or  sedges,  are  obscurely 
coloured.  Ko  doubt  if  their  colours  had  been  brilliant,  they 
would  have  been  much  more  conspicuous  to  their  enemies ;  but 
whether  their  dull  tints  have  been  specially  gained  for  the  sake 
of  protection  seems,  as  far  as  I  can  judge,  rather  doubtful.  It 
is  still  more  doubtful  whether  such  dull  tints  can  have  been 
gained  for  the  sake  of  ornament.  We  must,  however,  bear  in  mind 
that  male  birds,  though  dull-coloured,  often  differ  much  from 
their  females  (as  with  the  common  sparrow),  and  this  leads  to 
the  belief  that  such  colours  have  been  gained  through  sexual 
selection,  from  being  attractive.  Many  of  the  soft-billed  birds 
are  songsters ;  and  a  discussion  in  a  former  chapter  should  not 

^^  '  Ibis,'  1859,  vol.  i.  p.  429,  et      his  experience  of  the    Sahara,   this 
seq.     Dr.   Kohlfs,  however,  remarks      statemeut  is  too  strong. 
to  me  in  a  letter  that,  according  to 


Chap.  XYI.      Birds — Conspicuous  Colours.  491 

be  forgotten,  in  which  it  was  shewn  that  the  best  songsters  are 
rarely  oraamented  with  bright  tints.  It  would  appear  that  female 
birds,  as  a  general  rule,  have  selected  their  mates  either  for  their 
sweet  voices  or  gay  colours,  but  not  for  both  charms  combined. 
Some  species,  which  are  manifestly  coloured  for  the  sake  of 
protection,  such  as  the  jack-snipe,  woodcock,  and  night-jar,  are 
likewise  marked  and  shaded,  according  to  our  standard  of  taste, 
with  extreme  eleganee.  In  such  cases  we  may  conclude  that 
both  natural  and  sexual  selection  have  acted  conjointly  for 
protection  and  ornament.  Whether  any  bird  exists  which  does, 
not  possess  some  special  attraction,  by  which  to  charm  the 
opposite  sex,  may  be  doubted.  When  both  ?exes  are  so  obscurely 
coloured  that  it  would  be  rash  to  assume  the  agency  of  sexual 
selection,  and  when  no  direct  evidence  can  be  advanced  shewing 
that  such  colours  serve  as  a  protection,  it  is  best  to  own  complete 
ignorance  of  the  cause,  or,  which  comes  to  nearly  the  same  thing, 
to  attribute  the  result  to  the  direct  action  of  the  conditions  of  life. 

Both  sexes  of  many  birds  are  conspicuously,  though  not 
brilliantly  coloured,  such  as  the  numerous  black,  W'hite,  or 
piebald  species ;  and  these  colours  are  probably  the  result  of 
sexual  selection.  W^ith  the  common  blackbird,  capercailzie, 
blackcock,  black  scoter-duck  (Oidemia),  and  even  with  one  of 
the  bh'ds  of  paradise  (^Lophorina  a^ra),  the  males  alone  are  black, 
whilst  the  females  are  brown  or  mottled ;  and  there  can  hardly 
be  a  doubt  that  blackness  in  these  cases  has  been  a  sexually 
selected  character.  Therefore  it  is  in  some  degree  probable  that 
the  complete  or  partial  blackness  of  both  sexes  in  such  birds  as 
crows,  certain  cockatoos,  storks,  and  swans,  and  many  marine 
birds,  is  likewise  the  result  of  sexual  selection,  accompanied  by 
equal  transmission  to  both  sexes  ;  for  blackness  can  hardly  serve 
in  any  case  as  a  protection.  With  several  birds,  in  which  the 
male  alone  is  black,  and  in  others  in  which  both  sexes  are 
black,  the  beak  or  skin  about  the  head  is  brightly  coloured,  and 
the  contrast  thus  afforded  adds  much  to  their  beauty ;  we  see 
this  in  the  bright  yellow  beak  of  the  male  blackbird,  in  the 
crimson  skin  over  the  eyes  of  the  black-cock  and  capercailzie,  in 
the  brightly  and  variously  coloured  beak  of  the  scoter-drake 
(Oidemia),  in  the  red  beak  of  the  chough  {Corvus()racidus,  Linn.), 
of  the  black  swan,  and  the  black  stork.  This  leads  me  to  remark 
that  it  is  not  incredible  that  toucans  may  owe  the  enormous 
size  of  their  beaks  to  sexual  selection,  for  the  sake  of  displaying 
the  diversified  and  vivid  stripes  of  colour,  with  wiiich  these 
organs  are  ornamented. ^^    The  naked  skin,  also,  at  the  base  of  the 

^^  No  satisfactory  explanation  has  size,  and  still  less  of  the  bright 
ever  been  oflered   of   the    immense      colours,  cf  the  toucan's  beak.     Mr. 


492  The  Descent  of  Man.  Part  II. 

beak  and  round  the  eyes  is  likewise  often  brilliantly  coloured ; 
and  Mr.  Gould,  in  speaking  of  one  species/-  says  that  the  colours 
of  the  beak  "  are  doubtless  in  the  finest  and  most  brilliant  state 
"  during  the  time  of  pairing."  There  is  no  greater  improbability 
that  toucans  should  be  encumbered  with  immense  beaks,  thougli 
rendered  as  light  as  possible  by  their  cancellated  structure,  for 
the  display  of  fine  colours,  (an  object  falsely  appearing  to  us 
unimportant),  than  that  the  male  Argus  pheasant  and  some  other 
birds  should  be  encumbered  with  plumes  so  loug  as  to  impede 
their  flight. 

In  the  same  manner,  as  the  males  alone  of  various  species  are 
black,  the  females  being  dull-coloured ;  so  in  a  few  cases  the 
males  alone  are  either  wholly  or  partially  white,  as  with  the 
several  bell-birds  of  South  America  (Chasmorhynchus),  the 
Antarctic  goose  (Betmida  antarctica),  the  silver-pheasant,  &c., 
whilst  the  females  are  brown  or  obscurely  mottled.  Therefore, 
on  the  same  principle  as  before,  it  is  probable  that  both  sexes  of 
many  birds,  such  as  white  cockatoos,  several  egrets  with  their 
beautiful  plumes,  certain  ibises,  gulls,  terns,  &c.,  have  acquired 
their  more  or  less  completely  white  plumage  through  sexual 
selection.  In  some  of  these  cases  the  plumage  becomes  white 
only  at  maturity.  This  is  the  case  with  certain  gannets,  tropic- 
birds,  &c.,  and  with  the  snow-goose  (^;?se/-  hyperhoreus).  As  the 
latter  breeds  on  the  "  barren  grounds,"  when  not  covered  with 
snow,  and  as  it  migrates  southward  during  the  winter,  there  is 
no  reason  to  suppose  that  its  snow-white  adult  plumage  serves 
as  a  protection.  In  the  Anastomus  oscitans,  we  have  still  better 
evidence  that  the  white  plumage  is  a  nuptial  character,  for  it  is 
developed  only  during  the  summer  ;  the  young  in  their  imma- 
ture state,  and  the  adults  in  their  winter  dress,  being  grey  and 
black.  With  many  kinds  of  gulls  (Larus),  the  head  and  neck 
become  pure  white  dui'ing  the  summer,  being  grey  or  mottled 
during  the  winter  and  in  the  young  state.     On  the  other  hand. 


Bates     ('The     Naturalist     on     the  by  its  breadth,  depth,   as    well    as 

Amazons,'    vol.    ii.     1863,    p.    341)  length,  is    not    intelligible    on   the 

states  that  they  use  their  beaks  for  view,  that    it   serves  merely  as  an 

reaching  fruit  at  the   extreme  tips  organ     of     prehension.     Mr.      Belt 

of  the    branches ;  and  likewise,    as  believes  ('  The  Naturalist    in  Nica- 

stated    by   other    authors,    for    ex-  ragua,'  p.  197),  that  the  principal 

tract ing  eggs  and  young  birds  from  use    of  the    beak    is    as   a   defence 

the  nests  of  other  birds.     But,   as  against    enemies,  especially   to    the 

Air.   Bates  admits,  the    beak   "  can  female  whilst  nesting  in  a  hole  in  a 

"  scarcely  be  considered  a  very  per-  tree. 

"fectly-formed  instrument  for  the  ^^  Ramphastos  carinatus,  Gould's 

"  end  to  which  it  is  applied."     The  '  Monogx-aph  of  Eamphasstidae.' 
great   bulk  of  the   beak,  as  shewn 


Chap.  XVI.       Birds— ConspictioJts  Colours.  493 


with  the  smaller  gulls,  or  sea-mews  (Gavia),and  with  some  terns 
(Sterna)  exactly  the  reverse  occurs;  for  the  heads  of  the  young 
birds  during  the  first  year,  and  of  the  adults  during  the  waiter, 
are  either  pure  white,  or  much  paler  coloured  than  during  the 
breeding-season.  These  latter  cases  offer  another  instance  of  the 
capricious  manner  in  which  sexual  selection  appears  often  to 
hive  acted.^"^ 

That  aquatic  birds  have  acquired  a  white  plumage  so  much 
oftenerthan  terrestrial  birds,  probably  depends  on  their  large 
size  and  strong  powers  of  flight,  so  that  they  can  easily  defend 
themselves  or  escape  from  birds  of  prey,  to  which  moreover  they 
are  not  much  exposed.     Consequently,  sexual  selection  has  not 
here  been  interfered  with  or  guided  for  the  sake  ot  protection. 
No  doubt  with  birds  which  roam  over  the  open  ocean,  the  males 
and  females  could  find  each  other  much  more  easily,  when  made 
conspicuous  either  by  being  perfectly  white  or  intensely  black; 
so  that  these  colours  may  possibly  serve  the  same  end  as  the 
call-notes  of  many  land-birds.^*    A  white  or  black  bird  when  it 
discovers  and  flies  down  to  a  carcase  floating  on  the  sea  or  cast 
up  on  the  beach,  will  be  seen  from  a  great  distance,  and  will 
guide  other  birds  of  the  same  and  other  species,  to  the  prey  ;  but 
as  this  would  be  a  disadvantage  to  the  first  finders,  the  indi- 
viduals which  were  the  whitest  or  blackest  would  not  thus 
procure  more  food  than  the  less  strongly  coloured  individuals 
Hence  conspicuous  colours  cannot  have  been  gradually  acquired 
for  this  purpose  through  natural  selection. 

As  sexual  selection  depends  on  so  fluctuating  an  element  as 
taste  we  can  understand  how  it  is  that,  within  the  same  group 
of  biVds  having  nearly  the  same  habits,  there  should  exist  white 
or  nearly  white,  as  well  as  black,  or  nearly  black  species,-lor 
instance,  both  white  and  black  cockatoos,  storks,  ibises,  swans, 
terns  and  petrels.  Piebald  birds  likewise  sometimes  occur  m 
the  same  groups  together  with  black  and  white  species;  for 
instance,  the  black-necked  swan,  certain  terns,  and  the  common 
ma-pie.  That  a  strong  contrast  in  colour  is  agreeable  to  birds, 
we^'may  conclude  by  looking  through  any  large  collection,  for 
the  sexes  often  differ  from  each  other  in  the  male  having  the  pale 

"  On  Larus,  Gavia,  and  Sterna,  high  in  the  air,  like  marine   birds 

see  Maccrillivray,  '  Hist.  Brit.  Birds,'  over  the  ocean,  three  or  tour  species 

vol     V    P    515,  584,  626.     On  the  are  almost  wholly  or  largely  wnite, 

Anser  "hvperboreus,  Audubon,  '  Or-  and  that  many  others  are  black      bo 

ni  h     Biography,'   vol.    iv.   p.    562.  that  here  again  conspicuous  colours 

On   the  AaitoLs,  Mr.  Bl^-th,    in  may  possibly  aid  the  sexes  in  ^jding 

abis,'1867,  p.  173.  each    other     during    the     bieedmg- 

»■»  *It  may   be  noticed  that   with  season, 
vultures,  which  roam  far  and  wide 


494  ^^^^  Descent  of  Man.  Part  IT. 

parts  of  a  purer  white,  and  the  variously  coloured  dark  parts  of 
still  darker  tints  than  the  female. 

It  would  even  appear  that  mere  novelty,  or  slight  changes  for 
the  sake  of  change,  have  sometimes  acted  on  female  birds  as  a 
charm,  like  changes  of  fashion  with  us.  Thus  the  males  of 
some  parrots  can  hardly  be  said  to  be  more  beautiful  than  the 
females,  at  least  according  to  our  taste,  but  they  diifer  in  such 
points,  as  in  having  a  rose-coloured  collar  instead  of  "  a  bright 
"  emeraldine  narrow  green  collar;"  or  in  the  male  having  a  black 
collar  instead  of  "  a  yellow  demi-collar  in  front,"  with  a  pale 
roseate  instead  of  a  plum-blue  head.^^  As  so  many  male  birds 
have  elongated  tail-feathers  or  elongated  crests  for  their  chief 
ornament,  the  shortened  tail,  formerly  described  in  the  male  of 
a  humming-bird,  and  the  shortened  crest  of  the  male  goosander, 
seem  like  one  of  the  many  changes  of  fashion  wliich  we  admire 
in  our  own  dresses. 

Some  members  of  the  heron  family  offer  a  still  more  curious 
case  of  novelty  in  colouring  having,  as  it  appears,  been  appre- 
ciated for  the  sake  of  novelty.  The  young  of  the  Ardea  asha  are 
white,  the  adults  being  dark  slate-coloured;  and  not  only  the 
young,  but  the  adults  in  their  winter  plumage,  of  the  allied 
li'uphus  coromanclus  are  white,  this  colour  changing  into  a 
rich  golden-buff  during  the  breeding-season.  It  is  incredible 
that  the  young  of  these  two  species,  as  well  as  of  some  other 
members  of  the  same  family, ^*^  should  for  any  special  purpose, 
have  been  rendered  pure  white  and  thus  made  conspicuous  to 
their  enemies ;  or  that  the  adults  of  one  of  these  two  species 
should  have  been  specially  rendered  white  during  the  winter  in 
a  country  which  is  never  covered  with  snow.  On  the  other  hand 
we  have  good  reason  to  beheve  thai  whiteness  has  been  gained  by 
many  birds  as  a  sexual  ornament.  ^Ye  may  therefore  conclude 
that  some  early  progenitor  of  the  Ardea  asha  and  the  Buphus 
acquired  a  white  plumage  for  nuptial  purposes,  and  transmitted 
this  colour  to  their  young;  so  that  the  young,  and  the  old 
became  white  like  certain  existing  egrets ;  and  that  the  w^hiteness 
was  afterw^ards  retained  by  the  young,  whilst  it  was  exchanged 
by  the  adults  for  more  strongly- pronounced  tints.  But  if  w^e 
could  look  still  further  back  to  the  still  earlier  progenitors  of 
these  two  species,  we  should  j^robably  see  the  adults  dark- 

^^    See     Jerdon     on     the     genus  specific  names.     Audubon  ('  Ornith. 

Palaeornis,  'Birds  of  India,'   vol.   i.  Biography,'  vol.  iii.  p.  416  ;  a'oI.  iv. 

p.  258-260.  p.  58)  seems  rather  pleased  at  the 

*s  The  young  of  ^rc?ca  rufescens  thought  that  this  remarkable  change 

and  A.  c(x,rulea  of  the  U.  States  are  of  plumage  will  greatly  "disconcert 

likewise    white,    the     adults    being  "  the  systematists." 
coloured   in  accordance  with    their 


Chap.  XVI.  Birds — Novelty  admired.  495 


coloured.  I  infer  that  this  would  be  the  case,  from  the  analogy 
of  many  other  birds,  which  are  dark  whilst  young,  and  when 
adult  are  white ;  and  more  especially  from  the  case  of  the  Ardta 
gnlaris,  the  colours  of  which  are  the  reverse  of  those  of  A.  asha, 
for  the  young  are  dark-coloured  and  the  adults  white,  the  young 
having  retained  a  former  state  of  plumage.  It  appears  there- 
fore that,  during  a  long  line  of  descent,  the  adult  progenitors 
of  the  Ardea  ashu,  the  Buphus,  and  of  some  allies,  have  under- 
gone the  following  changes  of  colour :  first,  a  dark  shade ; 
secondly,  pure  white ;  and  thirdly,  owing  to  another  change  of 
fashion  (if  I  may  so  express  myself),  their  present  slaty,  reddish, 
or  golden-buff  tints.  These  successive  changes  are  intelligible 
only  on  tlie  principle  of  novelty  having  been  admired  by  birds 
for  its  own  sake. 

Several  writers  have  objected  to  the  whole  theory  of  sexual 
selection,  by  assuming  that  with  animals  and  savages  the 
taste  of  the  female  for  certain  colours  or  other  ornaments 
would  not  remain  constant  for  many  generations;  that  first 
one  colour  and  then  another  would  be  admired,  and  con- 
sequently that  no  permanent  effect  could  be  produced.  We  may 
admit  that  taste  is  fluctuating,  but  it  is  not  quite  arbitrary.  It 
depends  much  on  habit,  as  we  see  in  mankind ;  and  we  may 
infer  that  this  would  hold  good  with  birds  and  other  animals. 
Even  in  our  own  dress,  the  general  character  lasts  long,  and  the 
changes  are  to  a  certain  extent  graduated.  Abundant  evidence 
will  be  given  in  two  places  in  a  future  chapter,  that  savages 
of  many  races  have  admired  for  many  generations  the  same 
cicatrices  on  the  skin,  the  same  hideously  perforated  lips, 
nostrils,  or  ears,  distorted  heads,  &c. ;  and  these  deformities 
present  some  analogy  to  the  natural  ornaments  of  various 
animals.  Nevertheless,  with  savages  such  fashions  do  not  endure 
for  ever,  as  we  may  infer  from  the  differences  in  this  respect 
between  aUied  tribes  on  the  same  continent.  So  again  the  raisers 
of  fancy  animals  certainly  have  admired  for  many  generations 
and  still  admire  the  same  breeds ;  they  earnestly  desire  slight 
changes,  which  are  considered  as  improvements,  but  any  great  or 
sudden'  change  is  looked  at  as  the  greatest  blemish.  "With  birds 
in  a  state  of  nature  we  have  no  reason  to  suppose  that  they 
would  admire  an  entirely  new  style  of  coloration,  even  if  great 
and  sudden  variations  often  occurred,  which  is  far  from  being 
the  case.  We  know  that  dovecot  pigeons  do  not  willingly 
associate  with  the  variously  coloured  fancy  breeds ;  that  albino 
i)irds  do  not  commonly  get  partners  in  marriage ;  and  that  the 
black  ravens  of  the  Fcroe  Islands  chase  away  their  piebald 
brethren.     But    this  dislike  of  a    sudden  change  would  not 


49^  The  Descent  of  Man.  Part  II. 


preclude  their  appreciating  slight  chaDges,  any  more  than  it 
does  in  the  case  of  man.  Hence  with  respect  to  taste,  which 
depends  on  many  elements,  but  partly  on  habit  and  partly 
on  a  love  of  novelty,  there  seems  no  improbability  in  animals 
admiring  for  a  very  long  period  the  same  general  style  of  ornamen- 
tation or  other  attractions,  and  yet  appreciating  slight  changes 
in  colours,  form,  or  sound. 

Summary  of  the  Four  Chapters  on  Birds. — Most  male  birds  are 
highly  pugnacious  during  the  breeding-season,  and  some  possess 
weapons  adapted  for  fighting  with  their  rivals.  But  the  most 
pugnacious  and  the  best  armed  males  rarely  or  never  depend  for 
success  solely  on  their  power  to  drive  away  or  kill  their  rivals,  but 
have  special  means  for  charming  the  female.  With  some  it  is  the 
power  of  song,  or  of  giving  forth  strange  cries,  or  instrumental 
music,  and  the  males  in  consequence  differ  from  the  females  in 
their  vocal  organs,  or  in  the  structure  of  certain  feathers.  From 
the  curiously  diversified  means  for  x^roducing  various  sounds,  we 
gain  a  high  idea  of  the  importance  of  this  means  of  courtship. 
Many  birds  endeavour  to  charm  the  females  by  love-dances  or 
antics,  performed  on  the  ground  or  in  the  air,  and  sometimes  at 
prepared  places.  But  ornaments  of  many  kinds,  the  most 
brilliant  tints,  combs  and  wattles,  beautiful  plumes,  elongated 
feathers,  top-knots,  and  so  forth,  are  by  far  the  commonest  means. 
In  some  cases  mere  novelty  appears  to  have  acted  as  a  charm. 
The  ornaments  of  the  males  must  be  highly  important  to  them, 
for  they  have  been  acquired  in  not  a  few  cases  at  the  cost  of 
increased  danger  from  enemies,  and  even  at  some  loss  of  power 
in  fighting  with  their  rivals.  The  males  of  very  many  species 
do  not  assume  their  ornamental  dress  until  they  arrive  at 
maturity,  or  they  assume  it  only  during  the  breeding-&eason,  or 
the  tints  then  become  more  vivid.  Certain  ornamental 
appendages  become  enlarged,  turgid,  and  brightly  coloured 
during  the  act  of  courtship.  The  males  display  their  charms 
with  elaborate  care  and  to  the  best  efect ;  and  this  is  done  in 
the  presence  of  the  females.  The  courtship  is  sometimes  a 
prolonged  affair,  and  many  males  and  females  congregate  at  an 
appointed  place.  To  suppose  that  the  females  do  not  appreciate 
the  beauty  of  the  males,  is  to  admit  that  their  splendid  decorations, 
all  their  pomp  and  display,  are  useless ;  and  this  is  incredible. 
Birds  have  fine  powers  of  discrimination,  and  in  some  few 
instances  it  can  be  shewn  that  they  have  a  taste  for  the  beautiful. 
The  females,  moreover,  are  known  occasionally  to  exhibit  a 
marked  preference  or  antipathy  for  certain  individual  males. 

If  it  be  admitted  that  the  females  prefer,  or  are  unconsciously 


Chap.  XVI.  Birds — Sicinmary.  497 

excited  by  the  more  beautiful  males,  then  the  males  would 
slowly  but  surely  be  rendered  more  and  more  attractive  through 
sexual  selection.  That  it  is  this  sex  which  has  been  chiefly 
modified,  we  may  infer  from  the  fact  that,  in  almost  every  genus 
where  the  sexes  diifer,  the  males  differ  much  more  from  one 
another  than  do  the  females ;  this  is  well  shewn  in  certain 
closely-allied  representative  species,  in  which  the  females  can 
hardly  be  distinguished,  whilst  the  males  are  quite  distinct. 
Birds  in  a  state  of  nature  offer  individual  differences  which 
would  amply  suffice  for  the  work  of  sexual  selection;  but  we 
have  seen  that  they  occasionally  present  more  strongly-marked 
variations  which  recur  so  frequently  that  they  would  imme- 
diately be  fixed,  if  they  served  to  allure  the  female.  The  laws 
of  variation  must  determine  the  nature  of  the  initial  changes, 
and  will  have  largely  infl^uenced  the  final  result.  The  grada- 
tions, which  may  be  observed  between  the  males  of  allied 
species,  indicate  the  nature  of  the  steps  through  which  they 
have  passed.  They  explain  also  in  the  most  interesting  manner 
how  certain  characters  have  originated,  such  as  the  indented 
ocelli  on  the  tail-feathers  of  the  peacock,  and  the  ball  and 
socket  ocelli  on  the  wing-feathers  of  the  Argus  pheasant.  It  is 
evident  that  the  briUiant  colours,  top-knots,  fine  plumes,  &c.,  of 
many  male  birds  cannot  have  been  acquired  as  a  protection; 
indeed,  they  sometimes  lead  to  danger.  That  they  are  not  due 
to  the  direct  and  definite  action  of  the  conditions  of  life,  we  may 
feel  assured,  because  the  females  have  been  exposed  to  the  same 
conditions,  and  yet  often  differ  from  the  males  to  an  extreme 
degree.  Although  it  is  probable  that  changed  conditions  acting 
during  a  lengthened  period  have  in  some  cases  produced  a 
definite  effect  on  both  sexes,  or  sometimes  on  one  sex  alone,  the 
more  important  result  will  have  been  an  increased  tendency  to 
vary  or  to  present  more  strongly  marked  individual  differences ; 
and  such  differences  will  have  afforded  an  excellent  ground- 
work for  the  action  of  sexual  selection. 

The  laws  of  inheritance,  irrespectively  of  selection,  appear  to 
have  determined  whether  the  characters  acquired  by  the  males 
for  the  sake  of  ornament,  for  producing  various  sounds,  and  for 
fighting  together,  have  been  transmitted  to  the  males  alone  or  to 
both  sexes,  either  permanently,  or  periodically  during  certain 
seasons  of  the  year.  Why  various  characters  should  have  been 
transmitted  sometimes  in  one  way  and  sometimes  in  another,  is 
not  in  most  cases  known ;  but  the  period  of  variability  seems  often 
to  have  been  the  determining  cause.  When  the  two  sexes  have 
inherited  all  characters  in  common  they  necessarily  resemble 
each  other ;  but  as  the  successive  variations  may  be  differently 


498  The  Descent  of  Man.  Part  II. 

transmitted,  every  possible  gradation  may  be  found,  even  witliin 
the  same  genus,  from  the  closest  similarity  to  the  widest  dis- 
similarity between  the  sexes.  With  many  closely-allied  sjDecies, 
following  nearly  the  same  habits  of  life,  the  males  have  come  to 
differ  from  each  other  chiefly  through  the  action  of  sexual 
selection;  w^hilst  the  females  have  come  to  differ  chiefly  from 
partaking  more  or  less  of  the  characters  thus  acquired  by  the 
males.  The  effects,  moreover,  of  the  definite  action  of  the 
conditions  of  life,  will  not  have  been  masked  in  the  females,  as 
in  the  males,  by  the  accumulation  through  sexual  selection  of 
strongly-pronounced  colours  and  other  ornaments.  Ihe  indi- 
viduals of  both  sexes,  however  affected,  will  have  been  kept  at 
each  successive  period  nearly  uniform  by  the  free  intercrossing 
of  many  individuals. 

With  species,  in  which  the  sexes  differ  in  colour,  it  is  possible 
or  i^robable  that  some  of  the  successive  variations  often  tended 
to  be  transmitted  equally  to  both  sexes;  but  that  wiien  this 
occurred  the  females  w^ere  prevented  from  acquiring  the  bright 
colours  of  the  males,  by  the  destructicm  which  they  suffered 
during  incubation.  There  is  no  evidence  that  it  is  possible  by 
natural  selection  to  convert  one  form  of  transmission  into 
another.  But  there  would  not  be  the  least  difficulty  in  rendering 
a  female  dull-coloured,  the  male  being  still  kept  bright-coloured, 
by  the  selection  of  successive  variations,  which  were  from  the 
first  limited  in  their  transmission  to  the  same  sex.  Whether 
the  females  of  many  species  have  actually  been  thus  modified, 
must  at  present  remain  doubtful.  When,  through  the  law  of 
the  equal  transmission  of  characters  to  both  sexes,  the  females 
were  rendered  as  conspicuously  coloured  as  the  males,  their 
instincts  appear  often  to  have  been  modified  so  that  they  were 
led  to  build  domed  or  concealed  nests. 

In  one  small  and  curious  class  of  cases  the  characters  and 
habits  of  the  two  sexes  have  been  completely  transposed,  for  the 
females  are  larger,  stronger,  more  vociferous  and  brighter 
coloured  than  the  males.  They  have,  also,  become  so  quarrel- 
some that  they  often  fight  together  for  the  possession  of  the 
males,  like  the  males  of  other  pugnacious  species  for  the 
possession  of  the  females.  If,  as  seems  probable,  such  females 
habitually  drive  away  their  rivals,  and  by  the  display  of  their 
bright  colours  or  other  charms  endeavour  to  attract  the  males, 
we  can  understand  how  it  is  that  they  have  gradually  been 
rendered,  by  sexual  selection  and  sexually- limited  transmission, 
more  beautiful  than  the  males — the  latter  being  left  unmodified 
or  only  slightly  modified. 

Whenever  the  \2,\i  of  inheritance  at  corresponding  ages  prevails, 


Chap.  XVI.  Birds — Sjiinmary.  499 


but  not  that  of  sexually-limited  transmission,  then  if  the 
parents  vary  late  in  life— and  we  know  that  this  constantly 
occurs  with  our  poultry,  and  occasionally  with  other  birds — 
the  young  will  be  left  unaffected,  whilst  the  adults  of  both  sexes 
will  be  modified.  If  both  these  laws  of  inheritance  prevail  and 
either  sex  varies  late  in  life,  that  sex  alone  will  be  modified,  the 
other  sex  and  the  young  being  unaffected.  When  variations  in 
brightness  or  in  other  conspicuous  characters  occur  early  in  life, 
as  no  doubt  often  happens,  they  will  not  be  acted  on  through 
sexual  selection  until  the  period  of  reproduction  arrives ;  con- 
sequently if  dangerous  to  the  young,  they  will  be  eliminated 
through  natural  selection.  Thus  we  can  understand  how  it  is 
that  variations  arising  late  in  life  have  so  often  been  pre- 
served for  the  ornamentation  of  the  males;  the  females  and 
the  young  being  left  almost  unaifected,  and  therefore  like  each 
other.  With  species  having  a  distinct  summer  and  winter 
plumage,  the  males  of  which  either  resemble  or  differ  from  the 
females  during  both  seasons  or  during  the  summer  alone,  the 
degrees  and  kinds  of  resemblance  between  the  young  and  the 
old  are  exceedingly  complex ;  and  this  complexity  apparently 
depends  on  characters,  first  acquired  by  the  males,  being 
transmitted  in  various  ways  and  degrees,  as  limited  by  age,  sex, 
and  season. 

As  the  young  of  so  many  species  have  been  but  little  modified 
in  colour  and  in  other  ornaments,  we  are  enabled  to  form  some 
judgment  with  respect  to  the  plumage  of  their  early  progenitors; 
and  we  may  infer  that  the  beauty  of  our  existing  species,  if  we 
look  to  the  whole  class,  has  been  largely  increased  since  that 
period,  of  which  the  immature  plumage  gives  us  an  indirect 
record.  Many  birds,  especially  those  which  live  much  on  the 
ground,  have  undoubtedly  been  obscurely  coloured  for  the  sake 
of  protection.  In  some  instances  the  upper  exposed  surface  of 
the  plumage  has  been  thus  coloured  in  both  sexes,  whilst  the 
lower  surface  in  the  males  alone  has  been  variously  ornamented 
through  sexual  selection.  Finally,  from  the  facts  given  in 
these  four  chapters,  we  may  conclude  that  weaj3ons  for  battle, 
organs  for  producing  sound,  ornaments  of  many  kinds,  bright 
and  conspicuous  colours,  have  generally  been  acquired  by  the 
males  through  variation  and  sexual  selection,  and  have  been 
transmitted  in  various  ways  according  to  the  several  laws  of 
inheritance —the  females  and  the  young  being  left  comparatively 
but  little  modified." 

*^  I  am  greatly  iudebted  to  the  birds,  and  the  two  following  ones 
kindness  of  Mr.  Sclater  for  having  on  mammals.  In  this  way  I  have 
looked  over  these  four  chapters  on      been  saved   from    making    mistakes 


500  The  Descent  of  Man.  Pakt  IJ. 


CHAPTER  XVII. 
Secondaiiy  Sexual  Characteks  of  Mammals. 

The  law  of  battle — Special  weapons,  confined  to  the  males — Cause  of 
absence  of  weapons  in  the  female — Weapons  common  to  both  sexes,  yet 
primarily  required  by  the  male — Other  uses  of  such  weapons — Their 
high  importance — Greater  size  of  the  male — Means  of  defence — On  the 
preference  shewn  by  either  sex  in  the  pairing  of  quadrupeds. 

With  mammals  the  male  appears  to  win  tlie  female  iniTcli  more 
tlirough  the  law  of  battle  than  through  the  display  of  his 
charms.  The  most  timid  animals,  not  provided  with  any  special 
weapons  for  fighting,  engage  in  desperate  conflicts  during  the 
season  of  love.  Two  male  hares  have  been  seen  to  fight  together 
until  one  was  killed;  male  moles  often  fight,  and  sometimes 
with  fatal  results ;  male  squirrels  engage  in  frequent  contests, 
"  and  often  wound  each  other  severely ;"  as  do  male  beavers,  so 
that  "hardly  a  skin  is  without  scars." ^  I  observed  the  same 
fact  with  the  hides  of  the  guanacoes  in  Patagonia ;  and  on  one 
occasion  several  were  so  absorbed  in  fighting  that  they  fearlessly 
rushed  close  by  me.  Livingstone  speaks  of  the  males  of  the 
many  animals  in  Southern  Africa  as  almost  invariably  shewing 
the  scars  received  in  former  contests. 

The  law  of  battle  prevails  with  aquatic  as  with  terrestrial 
mammals.  It  is  notorious  how  desperately  male  seals  fight, 
both  with  their  teeth  and  claws,  during  the  breeding-season; 
and  their  hides  are  likewise  often  covered  with  scars.  Male 
sperm-whales  are  very  jealous  at  this  season;  and  in  their 
battles  "  they  often  lock  their  jaws  together,  and  turn  on  their 
"  sides  and  twist  about ;"  so  that  their  lower  jaws  often  become 
distorted.^ 


about  the  names  of  the  species,  and  and    Bachman,    'Viviparous    Quad- 

from    stating    anything    as    a    fact  rupeds  of  N.  America,' 1846,  p.  269. 

which  is  known  to  this  distinguished  On  beavers,   Mr,    A.    H.   Green,   in 

naturalist  to  be  erroneous.     But  of  'Journal  of  Lin.  Soc.  Zoolog.' vol.  x. 

course  he  is  not  at  all  answerable  1869,  p.  362. 

for  the  accuracy  of  the  statements  ^  On  the  battles  of  seals,  see  Capt. 

quoted  by  me  from  various  autho-  C.  Abbott  in  '  Proc.  Zool.  Soc' 1868, 

rities,  p.    191;  also    Mr.    R.  Brown,  ibid. 

^  See  Waterton's  account  of  two  1868,  p.  436;  also  L.  Lloyd,  'Game 

hares  fighting,    '  Zoologist,'  vol.    i.  Birds    of  Sweden,'    1867,    p.    412 ; 

1843,     p.     211.     On     moles,     Bell,  also  Pennant.     On  the  sperm-whale, 

'Hist,   of  British   Quadrupeds,'   1st  see  Mr.  J.   H.  Thompson,  in  '  Proc. 

edit.  p.  100.   On  squirrels,  Audubon  Zool.  Soc'  1867,  p.  246. 


Chap.  XVII.        Mammals — Laiv  of  Battle.  50 1 

All  male  animals  which  are  furnished  with  special  weapons 
for  fighting,  are  well  known  to  engage  in  fierce  battles.  The 
courage  and  the  desperate  conflicts  of  stags  have  often  been 
described ;  their  skeletons  have  been  found  in  various  parts  of 
the  world,  with  the  horns  inextricably  locked  together,  shewing 
how  miserably  the  victor  and  vanq^uishcd  had  perished.^  No 
animal  in  the  world  is  so  dangerous  as  an  elephant  in  must. 
Lord  Tankerville  has  given  me  a  graphic  description  of  the 
battles  between  the  wild  bulls  in  Chillingham  Park,  the  descen- 
dants, degenerated  in  size  but  not  in  courage,  of  the  gigantic 
Bos  i^rimigenius.  In  1861  several  contended  for  mastery;  and  it 
was  observed  that  two  of  the  younger  bulls  attacked  in  concert 
the  old  leader  of  the  herd,  overthrew  and  disabled  him,  so  that 
he  was  believed  by  the  keepers  to  be  lying  mortally  wounded  in 
a  neighbouring  wood.  But  a  few  days  afterwards  one  of  the 
young  bulls  approached  the  wood  alone ;  and  then  the  "  monarch 
"  of  the  chase,"  who  had  been  lashing  himself  up  for  vengeance, 
came  out  and,  in  a  short  time,  killed  his  antagonist.  He  then 
quietly  joined  the  herd,  and  long  held  undisputed  sway.  Admiral 
Sir  J.  B.  Sulivan  informs  me  that,  when  he  lived  in  the  Falk- 
land Islands,  he  imported  a  young  English  stallion,  which 
frequented  the  hills  near  Port  William  with  eight  mares.  On 
these  hills  there  were  two  wild  stallions,  each  with  a  small  troop 
of  mares ;  "  and  it  is  certain  that  these  stallions  would  never 
"  have  approached  each  other  without  fighting.  Both  had  tried 
"  singly  to  fight  the  English  horse  and  drive  away  his  mares, 
"  but  had  failed.  One  day  they  came  in  together  and  attacked 
"  him.  This  was  seen  by  the  capitan  who  had  charge  of  the 
"  horses,  and  who,  on  riding  to  the  spot,  found  one  of  the  two 
"  stallions  engaged  with  the  English  horse,  whilst  the  other  was 
"  driving  away  the  mares,  and  had  already  separated  four  from 
"  the  rest.  The  capitan  settled  the  matter  by  driving  the  whole 
"  party  into  the  corral,  for  the  wild  stallions  would  not  leave 
"  the  mares." 

Male  animals  which  are  provided  with  efficient  cutting  or 
tearing  teeth  for  the  ordinary  purposes  of  life,  such  as  the 
carnivora,  insectivora,  and  rodents,  are  seldom  furnished  with 
weapons  especially  adapted  for  fighting  w^ith  their  rivals.  The 
case  is  very  difi"erent  with  the  males  of  many  other  animals. 
We  see  this  in  the  horns   of  stags  and  of  certain  kinds  of 

3  See  Scrope  ('  Art  of  Deer-stalk-  wapiti,  moose,   and    rein-deer  have 

ing,'   p.    17)  on  the  locking  of  the  been    found    thus    locked    together, 

horns    with     the     Cervus     elaphus.  Sir  A.  Smith  found  at  the  Cape   of 

Richardson,  in  '  Fauna  Bor.  Ameri-  Good     Hope    the    skeletons    of  two 

cana/  1829,  p.    252,  says  that  the  gnus  in  the  same  condition. 


i;02  The  Descent  of  Man.  Part  II. 

antelopes  in  wliicli  the  females  are  hornless.  "With  many  animals 
the  canine  teeth  in  the  upper  or  lower  jaw,  or  in  both,  are  much 
larger  in  the  males  than  in  the  females,  or  are  absent  in  the 
latter,  with  the  exception  sometimes  of  a  hidden  rudiment. 
Certain  antelopes,  the  musk-deer,  camel,  horse,  boar,  various 
apes,  seals,  and  the  walrus,  offer  instances.  In  the  females  of 
the  walrus  the  tusks  are  sometimes  quite  absent.*  In  the  male 
elephant  of  India  and  in  the  male  dugong^  the  upper  incisors 
form  offensive  weapons.  In  the  male  narwhal  the  left  canine 
alone  is  developed  into  the  well-known,  spirally-twisted,  so- 
called  horn,  which  is  sometimes  from  nine  to  ten  feet  in  length. 
It  is  believed  that  the  males  use  these  horns  for  fighting  to- 
gether ;  for  "  an  unbroken  one  can  rarely  be  got.  and  occasionally 
"  one  may  be  found  with  the  point  of  another  jammed  into  the 
"  broken  place."  ^  The  tooth  on  the  opposite  side  of  the  head  in 
the  male  consists  of  a  rudiment  about  ten  inches  in  length, 
which  is  embedded  in  the  jaw ;  but  sometimes,  though  rarely, 
both  are  equally  developed  on  the  two  sides.  In  the  female  both 
are  always  rudimentary.  The  male  cachalot  has  a  larger  head 
than  that  of  the  female,  and  it  no  doubt  aids  him  in  his 
aquatic  battles.  Lastly,  the  adult  male  ornithorhynchus  is  pro- 
vided with  a  remarkable  apparatus,  namely  a  spur  on  the  foreleg, 
closely  resembling  the  poison-fang  of  a  venomous  snake ;  but  ac- 
cording to  Harting,  the  secretion  from  the  gland  is  not  poisonous ; 
and  on  the  leg  of  the  female  there  is  a  hollow,  apparently  for  the 
reception  of  the  spur.'' 

When  the  males  are  provided  with  weapons  which  in  the 
females  are  absent,  there  can  hardly  be  a  doubt  that  these  serve 
for  fighting  with  other  males;  and  that  they  were  acquired 
through  sexual  selection,  and  were  transmitted  to  the  male  sex 
alone.  It  is  not  probable,  at  least  in  most  cases,  that  the  females 
have  been  prevented  from  acquiring  such  weapons,  on  account 
of  their  being  useless,  superfluous,  or  in  some  way  injurious. 
On  the  contrary,  as  they  are  often  used  by  the  males  for  various 

*  Mr.  Lament  ('Seasons  with  the  ^  Mr.  R.  Brown,  in  '  Proc.  Zool. 
Sea-Horses,'  1861,  p,  143)  says  that  Soc*  1869,  p.  553.  See  Prof.  Turner, 
a  good  tusk  of  the  male  wah-us  in  Joui'nal  of  '  Anat.  and  Phys.' 1872, 
weighs  4  pounds,  and  is  longer  than  p.  76,  on  the  homological  nature  of 
that  of  the  female,  which  weighs  these  tusks.  Also  Mr.  J.  W.  Clarke 
about  3  pounds.  The  males  are  on  two  tusks  being  developed  in  the 
described  as  fighting  ferociously.  males,  in  '  Proc.  Zoolog.  Soc'  1871, 
On  the   occasional   absence    of    the  p.  42. 

tusks   in   the    female,    see    Mr.    R.  ^    Owen    on    the    cachalot     and 

Brown,   '  Proc.  Zool.  Soc'  1868,  p.  Ornithorhynchus,  ibid.  vol.  iii.   pp. 

429.  638,  641.     Harting  is  quoted  by  Dr. 

*  Owen, 'Anatomy  of  Vertebrates,'  Zouteveen  in  the  Dutch  translat.  ol 
vol.  iii.  p.  283.  this  work,  vol.  ii.  p.  292. 


Chap.  XVI I.        Mammals —Lazv  of  Battle,  503 

purposes,  more  especially  as  a  defence  against  their  enemies,  it 
is  a  surprising  fact  that  they  are  so  poorly  developed,  or  quite 
absent,  in  the  females  of  so  many  animals.  With  female  deer 
the  development  during  each  recurrent  season  of  great  branching 
horns,  and  with  female  elephants  the  development  of  immense 
tusks,  would  be  a  great  waste  of  vital  power,  supposing  that 
they  were  of  no  use  to  the  females.  Consequently,  they  would 
have  tended  to  be  eliminated  in  the  female  through  natural 
selection ;  that  is,  if  the  successive  variations  were  limited  in 
their  transmission  to  the  female  sex,  for  otherwise  the  weapons 
of  the  males  would  have  been  injuriously  affected,  and  this 
would  have  been  a  greater  evil.  On  the  whole,  and  from  the 
consideration  of  the  following  facts,  it  seems  probable  that  when 
the  various  weapons  differ  in  the  two  sexes,  this  has  generally 
depended  on  the  kind  of  transmission  which  has  prevailed. 

As  the  reindeer  is  the  one  species  in  the  whole  family  of  Deer, 
in  which  the  female  is  furnished  with  horns,  though  they  are 
somewhat  smaller,  thinner,  and  less  branched  than  in  the  male, 
it  might  naturally  be  thought  that,  at  least  in  this  case,  they 
must  be  of  some  special  service  to  her.  The  female  retains  her 
horns  from  the  time  when  they  are  fully  developed,  namely,  in 
September,  throughout  the  winter  until  April  or  May,  when  she 
brings  forth  her  young.  Mr.  Crotch  made  particular  enquiries 
for  me  in  Norway,  and  it  appears  that  the  females  at  this  season 
conceal  themselves  for  about  a  fortnight  in  order  to  bring  forth 
their  young,  and  then  reappear,  generally  hornless.  In  Nova 
Scotia,  however,  as  I  hear  from  Mr.  H.  Eeeks,  the  female  some- 
times retains  her  horns  longer.  The  male  on  the  other  hand 
casts  his  horns  much  earlier,  towards  the  end  of  November.  As 
both  sexes  have  the  same  requirements  and  follow  the  same 
habits  of  life,  and  as  the  male  is  destitute  of  horns  during  the 
winter,  it  is  improbable  that  they  can  be  of  any  special  service 
to  the  female  during  this  season,  which  includes  the  larger  part 
of  the  time  during  which  she  is  horned.  Nor  is  it  probable 
that  she  can  have  inherited  horns  from  some  ancient  pro- 
genitor of  the  family  of  deer,  for,  from  the  fact  of  the  females  of 
so  many  species  in  all  quarters  of  the  globe  not  having  horns, 
we  may  conclude  that  this  was  the  primordial  character  of  the 
group.^ 

The  horns  of  the  reindeer  are  developed  at  a  most  unusually 

*  On  the  structure  and  shedding  regard  to  the  American  variety  or 

of  the  horns  of  the  reindeer,  Hott-  species ;  also   Major  W.  Ross  King, 

berg,   '  Amoenitates   Acad.'  vol.  iv.  '  The  Sportsman  in   Canada,'  1866, 

1788,     p.     149.      See     Richardson,  p.  80. 
*  Fauna  Bor.  Americano,'  p.  241,  in 


504  The  Descent  of  Man,  Part  II. 

early  age;  but  what  the  cause  of  this  may  be  is  not  known. 
The  eifect  has  apparently  been  the  transference  of  the  horns  to 
both  sexes.  We  should  bear  in  mind  that  horns  are  always 
transmitted  through  the  female,  and  that  she  has  a  latent 
capacity  for  their  development,  as  we  see  in  old  or  diseased 
females.^  Moreover  the  females  of  some  other  species  of  deer 
exhibit,  either  normally  or  occasionally,  rudiments  of  horns ;  thus 
the  female  of  Cervulus  moschatus  has  "bristly  tufts,  ending  in  a 
"  knob,  instead  of  a  horn ;"  and  "  in  most  specimens  of  the 
"  female  wapiti  {Cervus  canadensis)  there  is  a  sharp  bony  pro- 
"  tuberance  in  the  place  of  the  horn."^°  From  these  several 
considerations  we  may  conclude  that  the  possession  of  fairly 
well-developed  horns  by  the  female  reindeer,  is  due  to  the  males 
having  first  acquired  them  as  weapons  for  fighting  with  other 
males;  and  secondarily  to  their  development  from  some  un- 
known cause  at  an  unusually  early  age  in  the  males,  and  their 
consequent  transference  to  both  sexes. 

Turning  to  the  sheath-horned  ruminants:  with  antelopes  a 
graduated  series  can  be  formed,  beginning  with  species,  the 
females  of  which  are  completely  destitute  of  horns — passing  on 
to  those  which  have  horns  so  small  as  to  be  almost  rudimentary, 
(as  with  the  Anfilocapra  americana,  in  which  species  they  are 
present  in  only  one  out  of  four  or  five  females  ^^) — to  those  which 
have  fairly  developed  horns,  but  manifestly  smaller  and  thinner 
than  in  the  male  and  sometimes  of  a  different  shape,^^ — and 
ending  with  those  in  which  both  sexes  have  horns  of  equal  size. 
As  with  the  reindeer,  so  with  antelopes  there  exists,  as  pre- 
viously shewn,  a  relation  between  the  jDeriod  of  the  development 
of  the  horns  and  their  transmission  to  one  or  both  sexes ;  it  is 
therefore  probable  that  their  presence  or  absence  in  the  females 
of  some  species,  and  their  more  or  less  perfect  condition  in  the 
females  of  other  species,  depends,  not  on  their  being  of  any 
special  use,  but  simply  in  inheritance.     It  accords  with  this 

^  Isidore     GeofTroy    St.     Hilaire,  *  Catalogue   of    ^Mammalia    in    the 

'Essais  de  Zoolog.  Generale,'  1841,  British    Museum,'   part.   iii.  p.  220. 

p.  513.     Other  masculine  characters,  On  the  Cervus  canadensis  or  wapiti 

besides    the    horns,    are    sometimes  see  Hon.  J.  D.  Caton,  '  Ottawa  Acad, 

similarly  transferred  to  the  female  ;  of  Nat.  Sciences,'  May,  1868,  p.  9. 
thus  Mr.  Boner,  in  speaking  of  an  '^   I  am  indebted  to  Dr.  Canfield, 

old     female      chamois      ('  Chamois  for    this    information,    see    also   his 

Hunting  in  the   Mountains   of  Ba-  paper  in  '  Proc.  Zoolog.  Soc.,'  1866, 

varia,'  1860,  2nd  edit.  p.  363),  says,  p.  105. 

"  not  only  was  the  head  very  male-  '-  For    instance  the  horns  of  the 

"  looking,  but  along  the  back  there  female  Ant,  euchore  resemble  those 

"was  a  ridge  of  long  hair,  usually  of  a  distinct  species,   viz.   the  Ant. 

"  to  be  found  only  in  bucks."  dorcas   var.    Corine,  see  Desmarest, 

'*  On    the    Cervulus,    Dr.    Gray,  '  Mammalogie,'  p.  455. 


Chap.  XVII.       Mammals — Law  of  Battle.  505 


view  that  even  in  the  same  restricted  genus  both  sexes  of  some 
species,  and  the  males  alone  of  others,  are  thus  provided.  It  is 
also  a  remarkable  fact  that,  although  the  females  of  Antilope 
hezoartica  are  normally  destitute  of  horns,  Mr.  Blyth  has  seen  no 
less  than  three  females  thus  furnished ;  and  there  was  no  reason 
to  suppose  that  they  were  old  or  diseased. 

In  all  the  wild  species  of  goats  and  sheep  the  horns  are 
larger  in  the  male  than  in  the  female,  and  arc  sometimes 
quite  absent  in  the  latter.^^  In  several  domestic  breeds  of 
these  two  animals,  the  males  alone  are  furnished  Avith  horns; 
and  in  some  breeds,  for  instance,  in  the  sheep  of  North  Wales, 
though  both  sexes  are  properly  horned,  the  ewes  are  very 
liable  to  be  hornless.  I  have  been  informed  by  a  trustworthy 
witness,  who  purposely  inspected  a  flock  of  these  same  sheep 
during  the  lambing  season,  that  the  horns  at  birth  are  generally 
more  fully  developed  in  the  male  than  the  female.  Mr.  J.  Peel 
crossed  his  Lonk  sheep,  both  sexes  of  which  always  bear  horns, 
with  hornless  Leicesters  and  hornless  Shropshire  Downs;  and 
the  result  was  that  the  male  offspring  had  their  horns  con- 
siderably reduced,  whilst  the  females  were  wholly  destitute  of 
them.  These  several  facts  indicate  that,  with  sheep,  the  horns 
are  a  much  less  firmly  fixed  character  in  the  females  than  in 
the  males ;  and  this  leads  us  to  look  at  the  horns  as  properly 
of  masculine  origin. 

With  the  adult  musk-ox  {Ovibos  moscJiatns)  the  horns  of  the 
male  are  larger  than  those  of  the  female,  and  in  the  latter  the 
bases  do  not  touch. ^*  In  regard  to  ordinary  cattle  Mr.  Blyth 
remarks :  "  In  most  of  the  wild  bovine  animals  the  horns  are 
"  both  longer  and  thicker  in  the  bull  than  in  the  cow,  and  in 
"  the  cow-banteng  {Bos  sondaicus)  the  horns  are  remarkably 
"  small,  and  inclined  much  backwards.  In  the  domestic  races 
"  of  cattle,  both  of  the  humped  and  humpless  types,  the  horns 
"  are  short  and  thick  in  the  bull,  longer  and  more  slender  in  the 
"  cow  and  ox ;  and  in  the  Indian  buffalo,  they  are  shorter  and 
"  thicker  in  the  bull,  longer  and  more  slender  in  the  cow.  In 
"  the  wild  gaour  (Z>,  r/aurus)  the  horns  are  mostly  both  longer 
"  and  thicker  in  the  bull  than  in  the  cow."^^  Dr.  Forsyth  Major 
also  informs  me  that  a  fossil  skull,  believed  to  be  that  of  the 
female  Bos  etruscus,  has  been  found  in  the  Val  d'Arno,  which  is 
wholly  without  horns.  In  the  liliinoceros  simus,  as  I  may  add, 
the  horns  of  the  female  are  generally  longer  but  less  powerful 
than  in  the  male ;  and  in  some  other  species  of  rhinoceros  they 

"  Gray,  *  Catalogue  Mamm.  Brit,      ricana,'  p.  278. 
Mus."  })art  iii.  1852,  p.  160.  i^  '  Land    and    Water,'    1867,    p. 

'^  Richardson,  '  Fauna  Bor.  Ame-      346. 


5o6  The  Descent  of  Man.  Part  11. 

are  said  to  be  shorter  in  the  female.^*'  From  these  various  facts 
we  may  infer  as  probable  that  horns  of  all  kinds,  even  when  they 
are  equally  developed  in  the  two  sexes,  were  primarily  acquired 
by  the  male  in  order  to  conquer  other  males,  and  have  been 
transferred  more  or  less  completely  to  the  female. 

The  effects  of  castration  deserve  notice,  as  throwing  light  on 
this  same  point.  Stags  after  the  oi^eration  never  renew  their 
horns.  The  male  reindeer,  however,  must  be  excepted,  as  after 
castration  he  does  renew  them.  This  fact,  as  well  as  the  pos- 
session of  horns  by  both  sexes,  seems  at  first  to  prove  that  the 
horns  in  this  species  do  not  coDstitute  a  sexual  character  ;^^  but 
as  they  are  developed  at  a  very  early  age,  before  the  sexes 
differ  in  constitution,  it  is  not  surprising  that  they  should  be 
unaffected  by  castration,  even  if  they  were  aboriginally  acquired 
by  the  male.  With  sheep  botli  sexes  properly  bear  horns ;  and 
I  am  informed  that  with  Welch  sheep  the  horns  of  the  males  are 
considerably  reduced  by  castration;  but  the  degree  depends 
much  on  the  age  at  which  the  operation  is  performed,  as  is  like- 
wise the  case  with  other  animals.  Merino  rams  have  large  horns, 
whilst  the  ewes  "'  generally  speaking  are  without  horns ;"  and  in 
this  breed,  castration  seems  to  produce  a  somewhat  greater 
effect,  so  that  if  performed  at  an  early  age  the  horns  "  remain 
"  almost  undeveloped."^*  On  the  Guinea  coast  there  is  a  breed 
in  which  the  females  never  bear  horns,  and,  as  Mr.  Win  wood 
Reade  informs  me,  the  rams  after  castration  are  quite  desti- 
tute of  them.  With  cattle,  the  horns  of  the  males  are  much 
altered  by  castration;  for  instead  of  being  short  and  thick, 
they  become  longer  than  those  of  the  cow,  but  otherwise  re- 
semble them.  The  Ant  Hope  hezoartica  offers  a  somewhat  ana- 
logous case :  the  males  have  long  straight  spiral  horns,  nearly 
])arallel  to  each  other,  and  directed  backwards ;  the  females 
occasionally  bear  horns,  but  these  when  present  are  of  a  very 
different  shape,  for  they  are  not  spiral,  and  spreading  widely, 
bend  round  with  the  points  forwards.  Now  it  is  a  remarkable  fact 
that,  in  the  castrated  male,  as  Mr.  BIyth  informs  me,  the  horns 
are  of  the  same  peculiar  shape  as  in  the  female,  but  longer  and 
thicker.  If  we  may  judge  from  analogy,  the  female  probably 
shews  us,  in  these  two  cases  of  cattle  and  the  antelope,  the  former 

•^  Sir    Andrew   Smith,    '  Zoology  quiries  for  me   iu    Saxony   on    this 

of  S.  Africa,'  pi.  xix.     Owen,  'Ana-  subject.     H.  von  Nathusius  (' Vieh- 

tomy  of  Vertebrates,'  vol.  iii.  p.  624.  zuclit,'   1872,  p.   64)  says  that  the 

^^  This     is     the     conclusion     of  horns  of  sheep  castrated  at  an  early 

Seidlitz,  '  Die   Darwinsche  Theorie,'  period,  either  altogether  disappear 

1871,  p.  47.  or  remain  as  mere  rudiments;   but 

^^  I  am  much    obliged   to    Prof.  I  do   not   know  whether   he    refers 

Victor  Carus,  for  having  made  en-  to  merinos  or  to  ordinary  breeds. 


Chap.  XVII.       Mammals — Lazv  of  Battle.  507 

condition  of  the  liorns  in  some  early  progenitor  of  each  species. 
But  why  castration  should  lead  to  the  reappearance  of  an  early 
condition  of  the  horns  cannot  be  explained  with  any  certainty. 
Nevertheless,  it  seems  probable,  that  in  nearly  the  same  manner 
as  the  constitutional  disturbance  in  the  offspring,  caused  by  a 
cross  between  two  distinct  species  or  races,  often  leads  to  the 
reappearance  of  long-lost  characters  ;^^  so  here,  the  disturbance 
in  the  constitution  of  the  individual,  resulting  from  castration, 
produces  the  same  effect. 

The  tusks  of  the  elephant,  in  the  different  species  or  races, 
differ  according  to  sex,  nearly  as  do  the  horns  of  ruminants.  In 
India  and  Malacca  the  males  alone  are  provided  with  well- 
developed  tusks.  The  elephant  of  Ceylon  is  considered  by  most 
naturalists  as  a  distinct  race,  but  by  some  as  a  distinct  species, 
and  here  "  not  one  in  a  hundred  is  found  with  tusks,  the  few 
"  that  possess  them  being  exclusively  males."-*'  The  African 
elepliant  is  undoubtedly  distinct,  and  the  female  has  large  well- 
developed  tusks,  though  not  so  large  as  those  of  the  male. 

These  differences  in  the  tusks  of  the  several  races  and  species 
of  elephants — the  great  variability  of  the  horns  of  deer,  as 
notably  in  the  wild  reindeer — the  occasional  presence  of  liorns 
in  the  female  Antilope  hezcartica,  and  their  frequent  absence  in 
the  female  of  Antilocapra  amerlcana — the  presence  of  two  tusks 
in  some  few  male  narwhals — the  complete  absence  of  tusks  in  some 
female  walruses— are  all  instances  of  the  extreme  variability  of 
secondary  sexual  characters,  and  of  their  liability  to  differ  in 
closely-allied  forms. 

Although  tusks  and  horns  appear  in  all  cases  to  have  been 
primarily  developed  as  sexual  weapons,  they  often  serve  other 
purposes.  The  elephant  uses  his  tusks  in  attacking  the  tiger ; 
according  to  Bruce,  he  scores  the  trunks  of  trees  until  they  can 
be  thrown  down  easily,  and  he  lijvcwise  thus  extracts  the  fari- 
naceous cores  of  palms ;  in  Africa  he  often  uses  one  tusk,  always 
the  same,  to  probe  the  ground  and  thus  ascertain  whether  it  will 
bear  liis  weight.  Tlie  common  bull  defends  the  herd  with  his 
horns;  and  the  elk  in  Sweden  has  been  known,  according  to 
Lloyd,  to  strike  a  wolf  dead  with  a  single  blow  of  his  great 
liorns.  Many  similar  facts  could  be  given.  One  of  the  most 
curious  secondary  uses  to  which  the  horns  of  an  animal  may 
be  occasionally  put,  is  that  observed  by  Captain  Hutton-"  with 

'9  I    have    given   various  experi-  ^o  gij.  j    Emerson  Tenncnl,  '  Cey- 

ments   and   otiier   evidence  proving  ion,'    1859,    vol.    ii.    p.    274.       For 

tliat  this  is  the  case,  in  my  '  Varia-  Malacca,  '  Journal  of  Indian  Archi- 

tion  of  Animals   and   Plants   under  pelago,'  vol.  iv.  p.  357. 
Domestication,'  vol.  ii.,  18(>8,  pp.  39  21  '  Calcutta  Journal  of  Nat.  Hist.' 

-47.  vol.  ii.  1843,  p.  52(5. 


5o8  The  Descent  of  Man.  Part  II. 

the  wild  goat  {Capra  segagrus)  of  the  Himalayas  and,  as  it  is  also 
said,  with  the  ibex,  namely,  that  when  the  male  accidentally  falls 
from  a  height  he  bends  inwards  his  head,  and  by  alighting  on 
his  massive  horns  breaks  the  shock.  The  female  cannot  thus 
use  her  horns,  which  are  smaller,  but  from  her  more  quiet 
disposition  she  does  not  need  this  strange  kind  of  shield  so 
much. 

Each  male  animal  uses  his  weapons  in  his  own  peculiar 
fashion.  The  common  ram  makes  a  charge  and  butts  with 
such  force  with  the  bases  of  his  horns,  that  I  have  seen  a  power- 
ful man  knocked  over  like  a  child.  Goats  and  certain  species  of 
sheep,  for  instance  the  Ovis  cydocero^  of  Afghanistan,^  rear  on 
their  hind  legs,  and  then  rot  only  butt,  but "  make  a  cut  down 
"  and  a  jerk  up,  with  the  ribbed  front  of  their  scimitar-shaped 
"  horn,  as  with  a  sabre.  When  the  0.  cydoceros  attacked  a  large 
*'  domestic  ram,  who  was  a  noted  bruiser,  he  conquered  him  by 
"  the  sheer  novelty  of  his  mode  of  fighting,  always  closing  at 
'•'  once  with  his  adversary,  and  catching  him  across  the  face  and 
"  nose  ^ith  a  sharp  drawing  jerk  of  the  head,  and  then  bounding 
"  out  of  the  way  before  the  blow  could  be  returned."  In 
Pembrokeshire  a  male  goat,  the  master  of  a  flock  which  during 
several  generations  had  run  wild,  was  known  to  have  killed  several 
males  in  single  combat ;  this  goat  possessed  enormous  horns, 
measuring  thirty-nine  inches  in  a  straight  line  from  tip  to 
tip.  The  common  bull,  as  every  one  knows,  gores  and  tosses  his 
opponent ;  but  the  Italian  buffalo  is  said  never  to  use  his  horns, 
he  gives  a  tremendous  blow  with  his  convex  forehead,  and  then 
tramples  on  his  fallen  enemy  with  his  knees— an  instinct  which 
the  common  bull  does  not  possess.^^  Hence  a  dog  who  pins  a 
buffalo  by  the  nose  is  immediately  crushed.  We  must,  however, 
remember  that  the  Italian  buffalo  has  been  long  domesticated, 
and  it  is  by  no  means  certain  that  the  wild  parent-form  had 
similar  horns.  Mr.  Bartlett  informs  me  that  when  a  female 
Cape  buffalo  (Buhalus  cafer)  was  turned  into  an  enclosure 
with  a  bull  of  the  same  species,  she  attacked  him,  and  he  in 
return  pushed  her  nbout  with  great  violence.  But  it  was 
manifest  to  Mr.  Bartlett  that,  had  not  the  bull  shewn  dignified 
forbearance,  he  could  easily  have  killed  her  by  a  single  lateral 
thrust  with  his  immense  horns.  The  giraffe  uses  his  short 
hair-covered  horns,  which  are  rather  longer  in  the  male  than 
in  the  female,  in  a  curious  manner ;  for,  with  his  long  neck,  he 
swings  his  head  to  either  side,  almost  upside  down,  with  such 

"    Mr.     Blyth,    in    '  Land    and  goats  see  the  '  Field,'  1869,  p.  150. 
Water,'    March,    1867,   p.    134,  on  23  j^j^  £_  ]^j_  Bailly,   '  Sur  I'usago 

the  authority  of  Capt.  Hutton  and  des   Comes,'    &c.,    'Annal.    des   Sc. 

others.   For  the  wild  Pembrokeshire  Nat.'  torn.  ii.  1824,  p.  369. 


Chap.  XVI  F.        Mammals — Law  of  Battle. 


509 


force,  tliat  I  have  seen  ca  hard  plank  deei^ly  indented  by  a  single 
blow. 

With  antelopes  it  is  sometimes  diflBcult  to  imagine  how  they 
can  possibly  use  their  curiously-shaped  horns  ;  thus  the  spring- 
boc  (^Ant.  euchore)  has  rather  short  upright  horns,  with  the 
sharp  points  bent  inwards  almost  at  right  angles,  so  as  to  face 
each  other ;  ]\Ir.  Bartlett  does  not  know  how  they  are  used,  but 
suggests  that  they  would  inflict  a  fearful  wound  down  each  side 
of  the  face  of  an  antagonist.    The  slightly-curved  horns  of  the 


Fig.  63.    Oryx  leucoryx,  male  (from  the  Knowsley  Menagerie). 


Oryx  leucoryx  (fig.  63)  are  directed  backwards,  and  are  of  such 
length  that  their  points  reach  beyond  the  middle  of  the  back, 
over  which  they  extend  in  almost  parallel  lines.  Thus  they 
seem  singularly  ill-fitted  for  fighting ;  but  Mr.  Bartlett  informs 
me  that  when  two  of  these  animals  prepare  for  battle,  they  kneel 
down,  with  their  heads  between  their  fore  legs,  and  in  this 
attitude  the  horns  stand  nearly  parallel  and  close  to  the  ground, 
with  the  points  directed  forwards  and  a  little  upwards.  The  com- 
batants then  gradually  approach  each  other,  and  each  endeavours 
to  get  the  upturned  points  under  the  body  of  the  other;  if  one 
succeeds  in  doing  this,  he  suddenly  springs  up,  throwing  up  his 
head  at  the  same  time,  and  can  thus  wound  or  perhaps  even 
transfix  his  antagonist.  Both  animals  always  kneel  down,  so  as 
to  guard  as  far  as  possible  against  this  manoeuvre.  It  has  l>een 
recorded  that  one  of  these  antelopes  has  used  his  horns  with 
effect  even  against  a  lion ;  yet  from  being  forced  to  place  his 
head  between  the  fore-legs  in  order  to  bring  the  points  of  the 


5 10  The  Descent  of  Man.  Part  II. 


horns  forward,  he  would  generally  be  under  a  great  disadvantage 
when  attacked  by  any  other  animal.  It  is,  therefore,  not 
probable  that  the  horns  have  been  modified  into  their  present 
great  length  and  pecuhar  position,  as  a  protection  against  beasts 
of  prey.  We  can  however  see  that,  as  soon  as  some  ancient 
male  progenitor  of  the  Oryx  acquired  moderately  long  horns, 
directed  a  little  backwards,  he  would  be  compelled,  in  his  battles 
with  rival  males,  to  bend  his  head  somewhat  inwards  or  down- 
wards, as  is  now  done  by  certain  stags ;  and  it  is  not  improbable 
that  he  might  have  acquired  the  habit  of  at  first  occasionally  and 
afterwards  of  regularly  kneeling  down.  In  this  case  it  is  almost 
certain  that  the  males  which  possessed  the  longest  horns  would 
have  had  a  great  advantage  over  others  with  shorter  horns  ;  and 
then  the  horns  would  gradually  have  been  rendered  longer  and 
longer,  through  sexual  selection,  until  they  acquired  their  present 
extraordinary  length  and  position. 

With  stags  of  many  kinds  the  branches  of  the  horns  ofi'er  a 
curious  case  of  diflBculty;  for  certainly  a  single  straight  point 
would  inflict  a  much  more  serious  wound  than  several  diverging 
ones.  In  Sir  Philip  Egerton's  museum  there  is  a  horn  of  the 
red-deer  {Ctrvus  elaphns),  thirty  inches  in  length,  with  "not 
"fewer  than  fifteen  snags  or  branches;"  and  at  Moritzburg 
there  is  still  preserved  a  pair  of  antlers  of  a  red-deer,  shot  in 
1699  by  Frederick  I.,  one  of  which  bears  the  astonishing  number 
of  thirty-three  branches  and  the  other  twenty-seven,  making 
altogether  sixty  branches.  Eichardson  figures  a  jDair  of  antlers 
of  the  wild  reindeer  with  twenty-nine  points.^'*  From  the 
manner  in  which  the  horns  are  branched,  and  more  especially 
fi'om  deer  being  known  occasionally  to  fight  together  by  kicking 
with  their  fore-feet,^^  M.  Bailly  actually  comes  to  the  conclusion 
that  their  horns  are  more  injurious  than  useful  to  them  ?  But 
this  author  overlooks  the  pitched  battles  between  rival  males. 
As  I  felt  much  perplexed  about  the  use  or  advantage  of  the 
branches,  I  applied  to  Mr.  McNeill  of  Colonsay,  who  has  long 
and  carefully  observed  the  habits  of  red-deer,  and  he  informs 
me  that  he  has  never  seen  some  of  the  branches  brought  into 
use,  but  that  the  brow  antlers,  from  inclining  downwards,  are  a 
great  protection  to  the  forehead,  and  their  points  are  likewise 

2*  On     the    horns     of    red-deer,  Acnd,  of  Nat.  Science,'  May,  1868, 

Owen,    '  British    Fossil    Mammals,'  p.  9),  says  that  the  American  deer 

1846,   p.   478;    Eichardson  on    the  right     with    their    fore-feet,    after 

horns  of  the  reindeer,  '  Fauna  Bor.  "  the   question    of    superiority   has 

Americana,'   1829,    p.    240.     I    am  "  been  once  settled  and  acivnowledged 

indebted  to  Prof.  Victor  Carus,  for  "  in  the  herd."     Bailly, 'Sur  I'usage 

the  Moritzburg  case.  des  Cornes,'  '  Annales  des  So.  Nat. 

"  Hon.   J.  ^D.    Caton    ('Ottawa  tom.  ii.  1824,  p.  371. 


Chap.  XVII.      Mammals — Lazv  of  Battle.  5 1 1 

used  in  attack.  Sir  Philip  Egerton  also  informs  me  both  as  to 
red-deer  and  fallow-deer  that,  in  lighting,  they  suddenly  dash 
together,  and  getting  their  horns  fixed  against  each  other's 
bodies,  a  desperate  struggle  ensues.  When  one  is  at  last  forced 
to  yield  and  turn  round,  the  victor  endeavours  to  plunge  his 
brow  antlers  into  his  defeated  foe.  It  thiis  a])pears  that  the 
upper  branches  are  used  chiefly  or  exclusively  for  pushing  and 
fencing.  Nevertheless  in  some  species  the  iipper  branches  are 
used  as  weapons  of  offence;  when  a  man  was  attacked  by  a 
wapiti  deer  {Cei-vus  cimadeiisis)  in  Judge  Caton's  park  in 
Ottawa,  and  several  men  tried  to  rescue  him,  the  stag  "  never 
"  raised  his  head  from  the  ground ;  in  fact,  he  kept  his  face  almost 
"  flat  on  the  ground,  with  his  nose  nearly  between  his  fore  feet, 
"  except  when  he  rolled  his  head  to  one  side  to  take  a  new 
"  observation  preparatory  to  a  plunge."  In  this  position  the 
ends  of  the  horns  were  directed  against  his  adversaries.  "  In 
"  rolling  his  head  he  necessarily  raised  it  somewhat,  because  his 
"  antlers  were  so  long  that  he  could  not  roll  his  liead  without 
"  raising  them  on  one  side,  while,  on  the  other  side,  they  touched 
"  the  ground."  The  stag  by  this  procedure  gradually  drove  the 
party  of  rescuers  backwards,  to  a  distance  of  150  or  200  feet ; 
and  the  attacked  man  was  killed.^^  . 

Although  the  horns  of  stags  are  efficient  weapons,  there  can, 
I  think  be  no  doubt  that  a  single  point  would  have  been  much 
more  dangerous  than  a  branched  antler  ;  and  Judge  Caton,  who 
has  had  large  experience  with  deer,  fully  concurs  in  this  conclu- 
sion. Nor  do  the  branching  horns,  though  highly  important  as 
a  means  of  defence  against  rival  stags,  appear  perfectly  well 
adapted  for  this  purpose,  as  they  are  liable  to  become  interlocked. 
The  suspicion  has  therefore  crossed  my  mind  that  they  may 
serve  in  part  as  ornaments.  That  the  branched  antlers  of  stags 
as  well  as  the  elegant  lyrated  horns  of  certain  antelopes,  with 
their  graceful  double  curvature,  (fig.  G4),  are  ornamental  in  our 
eyes,  no  one  will  dispute.  If,  then,  the  horns,  like  the  splendid 
accoutrements  of  the  knights  of  old,  add  to  the  noble  appearance 
of  stags  and  antelopes,  they  may  have  been  modified  i3artly  for 
this  purpose,  though  mainly  for  actual  service  in  battle ;  but  I 
have  no  evidence  in  favour  of  this  belief. 

An  interesting  case  has  lately  been  published,  from  which  it 
appears  that  the  horns  of  a  deer  in  one  district  in  the  United 
States  are  now  being  modified  through  sexual  and  natural  selection. 
A  writer  in  an  excellent  American  Journal  "'^  says,  that  he  has 

26  See  a  most  intoresting  a/^ODUut  ^7  i  "d^q     Americaa     Naturalist,' 

in   the    Appendix    to    Hon.    J.    D.      Dec.  18G9,  p.  552. 
Caton's  paper,  as  above  quoted. 


512 


The  Descent  of  Man. 


Part  IL 


liunted  for  the  last  twenty-one  years  in  the  Adirondacks,  where 
the  Ccrvus  virginianus  abounds.  About  fourteen  years  ago  he 
first  heard  of  spike-horn  bucks.  These  became  from  year  to  year 
more  common ;  about  five  years  ago  he  shot  one,  and  afterwards 
another,  and  now  they  are  frequently  killed.  "  The  spike-horn 
"  differs  greatly  from  the  common  antler  of  the  C.  virginianus. 


Fig.  64.    Strepsiceros  Kudu  (from  Sir  Andrew  Smith's  '  Zoology  of  South  Africa'). 

it  consists  of  a  single  spike,  more  slender  than  the  antler,  and 
scarcely  half  so  long,  projecting  forward  from  the  brow,  and 
terminating  in  a  very  sharp  point.     It  gives  a  considerable 


Cha p.  XYII.       Mam mals — L  aw  of  Battle.  5 1 3 

"  advantage  to  its  possessor  over  the  common  buck.  Besides 
"  enabling  him  to  run  more  swiftly  through  the  thick  woods  and 
"  underbrush  (every  hunter  knows  that  does  and  yearling 
"  bucks  run  much  more  rapidly  than  the  large  bucks  when 
"  armed  with  their  cumbrous  antlers),  the  spike-horn  is  a  more 
"  effective  weapon  than  the  common  antler.  With  this  advantage 
"  the  spike-horn  bucks  are  gaining  upon  the  common  bucks,  and 
"  may,  in  time,  entirely  supersede  them  in  the  Adirondacks. 
"  Undoubtedly,,  the  first  spike-horn  buck  was  merely  an  acci- 
"  dental  freak  of  nature.  But  his  spike-horns  gave  him  an 
"  advantage,  and  enabled  him  to  propagate  his  peculiarity.  His 
"  descendants  having  a  like  advantage,  have  propagated  the 
"  peculiarity  in  a  constantly  increasing  ratio,  till  they  are 
"  slowly  crowding  the  antlered  deer  from  the  region  they 
"  inhabit."  A  critic  has  well  objected  to  this  account  by  asking, 
why,  if  the  simple  horns  are  now  so  advantageous,  were  the 
branched  antlers  of  the  parent-form  ever  developed  ?  To  this  1 
can  only  answer  by  remarking,  that  a  new  mode  of  attack  with 
new  weapons  might  be  a  great  advantage,  as  shewn  by  the  ca§e 
of  the  Ovis  cycloceros,\\]io  thus  conquered  a  domestic  ram  famous 
for  his  fighting  power.  Though  the  branched  antlers  of  a  stag 
are  well  adapted  for  fighting  with  his  rivals,  and  though  it 
might  be  an  advantage  to  the  prong-horned  variety  slowly  to 
acquire  long  and  branched  horns,  if  he-  had  to  fight  only 
with  others  of  the  same  kind,  yet  it  by  no  means  follows  that 
branched  horns  would  be  the  best  fitted  for  conquering  a  foe 
differently  armed.  In  the  foregoing  case  of  the  Oryx  Itucoryx,  it 
is  almost  certain  that  the  victory  would  rest  with  an  antelope 
having  short  horns,  and  who  therefore  did  not  need  to  kneel 
down,  though  an  oryx  might  protit  by  having  still  longer  horns, 
if  he  fought  only  with  his  proper  rivals. 

Male  quadrupeds,  which  are  furnished  with  tusks,  use  them  in 
various  ways,  as  in  the  case  of  horns.  The  boar  strikes  laterally 
and  upwards;  the  musk-deer  downwards  with  serious  effect.^^ 
The  walrus,  though  having  so  short  a  neck  and  so  unwieldy  a 
body,  "can  strike  either  upwards,  or  downwards,  or  sideways,  with 
"  equal  dexterity."'^  I  was  informed  by  the  late  Dr.  Falconer, 
that  the  Indian  elephant  fights  in  a  different  manner  according 
to  the  position  and  curvature  of  his  tusks.  When  they  are 
directed  forwards  and  upwards  he  is  able  to  Ping  a  tiger  to  a 
great  distance — it  .is  said  to  even  thirty  feet;  when  they  are 
short  and  turned  downwards  he  endeavours  suddenly  to  pin  the 

2S  Pallas,   'Spicilegia    Zoologica,'  -"  Lamont, '  Seasons  with  the  Sea- 

fasc.  xiii.  1779,  p.  18.  Horses,'  18(31,  p.  141. 

23 


514  The  Descent  of  Man.  Part  IT. 


tiger  to  the  ground  and,  in  consequence,  is  dangerous  to  the 
rider,  who  is  liable  to  be  jerked  off  the  howdah.^" 

Very  few  male  quadrupeds  possess  weapons  of  two  distinct 
kinds  specially  adapted  for  fighting  with  rival  males.  The  male 
muntjac-deer  (Cervulus),  however,  offers  an  exception,  as  he  is 
provided  with  horns  and  exserted  canine  teeth.  But  we  may 
infer  from  what  follows  that  one  form  of  weapon  has  often 
been  replaced  in  the  course  of  ages  by  another.  With  ruminants 
the  development  of  horns  generally  stands  in  an  inverse  relation 
with  that  of  even  moderately  developed  canine  teeth.  Thus 
camels,  guanacoes,  chevrotains,  and  musk-deer,  are  hornless, 
and  they  have  efficient  canines;  these  teeth  being  ''always  of 
"  smaller  size  in  the  females  than  in  the  males."  The  CamelidsB 
have,  in  addition  to  their  true  canines,  a  pair  of  canine-shaped 
incisors  in  their  upper  jaws.^^  Male  deer  and  antelopes,  on  the 
other  hand,  possess  horns,  and  they  rarely  have  canine  teeth ; 
and  these,  when  present,  are  always  of  small  size,  so  that  it  is 
doubtful  whether  they  are  of  any  service  in  their  battles.  In 
Antilope  monfana  they  exist  only  as  rudiments  in  the  young 
male,  disappearing  as  he  grows  old  ;  and  they  are  absent  in  the 
female  at  all  ages ;  but  the  females  of  certain  other  antelopes 
and  of  certain  deer  have  been  known  occasionally  to  exhibit 
rudiments  of  these  teeth.^^  Stallions  have  small  canine  teeth, 
which  are  either  quite  absent  or  rudimentary  in  the  mare;  but 
they  do  not  appear  to  be  used  in  fighting,  for  stallions  bite 
with  their  incisors,  and  do  not  open  their  mouths  wide  like 
camels  and  guanacoes.  Whenever  the  adult  male  possesses 
canines,  now  inefficient,  whilst  the  female  has  either  none  or 
mere  rudiments,  we  may  conclude  that  the  early  male  pro- 
genitor of  the  species  was  provided  with  efficient  canines,  which 
have  been  partially  transferred  to  the  females.  The  reduction  of 
these  teeth  in  the  males  seems  to  have  followed  from  some 
change  in  their  manner  of  fighting,  often  (but  not  in  the  horse) 
caused  by  the  development  of  new  weapons. 

Tusks  and  horns  are  manifestly  of  high  importance  to  their 

^<*  See    also    Corse     ('  PhilosojDh.  American  deer.     See   also   Falconer 

Transact.'     1799,    p.   212)    on   the  (' Palseont.    Memoirs    and     Notes,' 

manner  in  which  the   short-tusked  vol.  i.   1868,  p.  576)  on  canines  in 

Mookuah  variety  attacks  other  ele-  an  adult  female  deer.     In  old  males 

phauts.  of  the  musk-deer  the  canines  (Pallas, 

31  Owen,  'Anatomy  of  Verte-  '  Spic.  Zoolog:'  fasc.  xiii.  1779,  p. 
orates,'  vol.  iii.  p.  349.  18)  sometimes   grow  to  the   length 

32  See  Ruppell  (in  'Proc.  Zoolog.  of  three  inches,  whilst  in  old  females 
Soc'  Jan.  12,  1836,  p.  3)  on  the  a  rudiment  projects  scarcely  half 
canines  in  deer  and  antelopes,  with  an  inch  above  the  gums. 

a  note  by  Mr.  Martin  on  a  female 


Chap.  XVII.    Mammals — Greater  Size  of  Male.         515 


possessors,  for  their  development  consumes  much  organised 
matter.  A  single  tusk  of  the  Asiatic  elephant — one  of  the 
extinct  woolly  species — and  of  the  African  elephant,  have  been 
known  to  weigh  respectively  150,  160,  and  180  pounds ;  and 
even  greater  weights  have  been  given  by  some  authors.^^  With 
deer,  in  which  the  horns  are  periodically  renewed,  the  drain  on 
the  constitution  must  be  greater ;  the  horns,  for  instance,  of  the 
moose  weigh  from  fifty  to  sixty  pounds,  and  those  of  the 
extinct  Irish  elk  from  sixty  to  seventy  pounds — the  skull  of  the 
latter  weighing  on  an  average  only  five  pounds  and  a  quarter. 
Although  the  horns  are  not  periodically  renewed  in  sheep,  yet 
their  development,  in  the  opinion  of  many  agriculturists,  entails 
a  sensible  loss  to  the  breeder.  Stags,  moreover,  in  escaping 
from  beasts  of  prey  ai-e  loaded  with  an  additional  weight  for  the 
race,  and  are  greatly  retarded  in  passing  through  a  woody 
country.  The  moose,  for  instance,  with  horns  extending  five 
and  a  half  feet  from  tip  to  tip,  although  so  skilful  in  their  use 
that  he  will  not  touch  or  break  a  twig  when  walking  quietly, 
cannot  act  so  dexterously  whilst  rushing  away  from  a  pack 
of  wolves.  "During  his  progress  he  holds  his  nose  up,  so 
"  as  to  lay  the  horns  horizontally  back ;  and  in  this  attitude 
"  cannot  see  the  ground  distinctly.''^*  The  tips  of  the  horns  of 
the  great  Irish  elk  were-  actually  eight  feet  apart !  Whilst  the 
horns  are  covered  with  velvet,  which  lasts  with  the  red-deer  for 
about  twelve  weeks,  they  are  extremely  sensitive  to  a  blow ;  so 
that  in  Germany  the  stags  at  this  time  somewhat  change  their 
habits,  and  avoiding  dense  forests,  frequent  }'oung  woods  and 
low  thickets.^^  These  facts  remind  us  that  male  birds  have 
acquired  ornamental  plumes  at  the  cost  of  retarded  flight,  and 
other  ornaments  at  the  cost  of  some  loss  of  power  in  their  battles 
with  rival  males. 

With  mammals,  when,  as  is  often  the  case,  the  sexes  differ  in 
size,  the  males  are  almost  always  larger  and  stronger.  I  am 
informed  by  Mr.  Gould,  that  this  holds  good  in  a  marked 
manner  with  the  marsupials  of  Australia,  the  males  of  which 
appear  to  continue  growing  until  an  unusually  late  age.  But 
the  most  extraordinary  case  is  that  of  one  of  the  seals  {ddJor- 
hinus  ursinus),  a  full-grown  female  weighing  less  than  one-sixth 
of  a  full-grown  male.^''      Dr.  Gill  remarks  that  it  is  with  the 

'3  Emerson     Tennent,     'Ceylon,  143.      See     also     Owen,     *  British 

1859,  vol,  ii.  p.  275;  Owen,  '  Bri-  Fossil  Mammals,'  on  the  Irish  elk, 

tish  Fossil  Mammals,'  1846,  p.  245.  pp.  447,  455. 

3«  Richardson,  '  Fauna  Bor.  Ame-  "  '  Forest  Creatures,' by  C.  Bouor, 

ricana,'on  the  moose,  Alces  jyalmata,  1861,  p.  60. 

pp.  236,  237  ;  on  the  expanse  of  the  ^^  See  the  very  interesting  paper 

horns,  'Land  and  Water,'   18r39,  p.  by  Mr.  J.   A.   Alien  in  'Bull.   Mus, 


Si6  The  Descent  of  Man.  Part  II. 

polygamous  seals,  the  males  of  which  are  well  known  to  fight 
savagely  together,  that  the  sexes  differ  much  in  size ;  the  mono- 
gamous species  differing  but  little.  Whales  also  afford  evidence 
of  the  relation  existing  between  the  pugnacity  of  the  males  aud 
their  large  size  compared  with  that  of  the  female ;  the  males  of 
the  right -whales  do  not  fight  together,  and  they  are  not  larger, 
but  rather  smaller,  than  their  females;  on  the  other  hand,  male 
sperm-whales,  fight  much  together,  and  their  bodies  are  "  often 
"  found  scarred  with  the  imprint  of  their  rival's  teeth,"  and 
they  are  double  the  size  of  the  females.  The  greater  strength  of 
the  male,  as  Hunter  long  ago  remarked,^"  is  invariably  displayed 
in  those  parts  of  the  body  which  are  brought  into  action  in 
fighting  with  rival  males — for  instance,  in  the  massive  neck  of 
the  bull.  Male  quadrupeds  are  also  more  courageous  and 
pugnacious  than  the  females.  There  can  be  little  doubt  that 
these  characters  have  been  gained,  partly  through  sexual  selec- 
tion, owing  to  a  long  series  of  victories  by  the  stronger  and 
more  courageous  males  over  the  weaker,  and  partly  through  the 
inherited  effects  of  use.  It  is  probable  that  the  successive 
variations  in  strength,  size,  and  courage,  whether  due  to  mere 
variability  or  to  the  effects  of  use,  by  the  accumulation  of  which 
male  quadrupeds  have  acquired  these  characteristic  qualities, 
occurred  rather  late  in  life,  and  were  'consequently  to  a  large 
extent  limited  in  tiieir  transmission  to  the  same  sex. 

Ei-om  these  considerations  I  was  anxious  to  obtain  information 
as  to  the  Scotch  deerhound,  the  sexes  of  which  differ  more  in 
size  than  those  of  any  other  breed  (though  bloodhounds  differ 
considerably),  or  than  in  any  wild  canine  species  known  to  me. 
Accordingly,  I  applied  to  Mr.  Cupples,  well-known  for  his 
success  with  this  breed,  who  has  weighed  and  measured  many 
of  his  own  dogs,  and  who  has  with  great  kindness  collected  for 
me  the  following  facts  from  various  sources.  Fine  male  dogs, 
measured  at  the  shoulder,  range  from  28  inches,  which  is  low, 
to  83  or  even  34  inches  in  height ;  and  in  weight  from  80 
pounds,  which  is  light,  to  120  pounds,  or  even  more.  The 
females  range  in  height  from  23  to  27,  or  even  to  28  inches ;  and 
in  weight  from  £0  to  70,  or  even  80  pounds.^^     Mr.  Cupples 


Comp.  Zoolog.  of  Cambridge,  United  ^^  'Animal  Economy,'  p.  45. 
States,'  vol.  ii.  No.  1,  p.  82.  The  3*  See  also  Richardson's  '  Manual 
weights  were  ascertained  by  a  care-  on  the  Dog,'  p.  59.  Much  valuable 
ful  observer,  Capt.  Bryant.  Dr.  information  on  the  Scottish  deer- 
Gill  in  'The  American  Naturalist,'  hound  is  given  by  Mr.  McNeill, 
Jan.  1871,  Prof.  Shaler  on  the  who  tirst  called  attention  to  the 
relative  size  of  the  sexes  of  whales,  inequality  in  size  between  the, sexes, 
'American  Naturalist,'  Jan.  1873.  in  Scrope's  'Art  of  Deer  Stalking.' 


Chap.  XVII.    Mammals — Greater  Size  of  Male.  517 

concludes  that  from  95  to  100  pounds  for  the  male,  and  70  for 
the  female,  would  be  a  safe  average;  but  there  is  reason  to 
believe  that  formerly  both  sexes  attained  a  greater  weight. 
Mr.  Cupples  has  weighed  puppies  when  a  fortnight  old  ;  in  one 
litter  the  average  weight  of  four  males  exceeded  that  of  two 
females  by  six  and  a  half  ounces ;  in  another  litter  the  average 
weight  of  four  males  exceeded  that  of  one  female  by  less  than 
one  ounce;  the  same  males  when  three  weeks  old,  exceeded 
the  female  by  seven  and  a  half  ounces,  and  at  the  age  of  six 
weeks  by  nearly  fourteen  ounces.  Mr.  Wright  of  Yeldersley 
House,  in  a  letter  to  Mr.  Cupples,  says  :  "  I  have  taken  notes  on 
"  the  sizes  and  weights  of  puppies  of  many  litters,  and  as  far  as 
"  my  experience  goes,  dog-pui3pies  as  a  rule  differ  very  little 
"•  from  bitches  till  they  arrive  at  about  five  or  six  months  old ; 
"  and  then  the  dogs  begin  to  increase,  gaining  upon  the  bitches 
"  both  in  weight  and  size.  At  birth,  and  for  several  weeks 
"  afterwards,  a  bitch-puppy  will  occasionally  be  larger  than  any 
*'  of  the  dogs,  but  they  are  invariably  beaten  by  them  later." 
Mr.  McNeill,  of  Colonsay,  concludes  that  "the  males  do  not 
•'  attain  their  full  growth  till  over  two  years  old,  though  the 
"  females  attain  it  sooner."  According  to  Mr.  Cupples'  expe- 
rience, male  dogs  go  on  growing  in  stature  till  they  are  from 
twelve  to  eighteen  months  old,  and  in  w^eight  till  from  eighteen 
to  tw^enty-four  months  old ;  whilst  the  females  cease  increasing 
in  stature  at  the  age  of  from  nine  to  fourteen  or  fifteen  months, 
and  in  weight  at  the  age  of  from  twelve  to  fifteen  months. 
From  these  various  statements  it  is  clear  that  the  full  difference 
in  size  between  the  male  and  female  Scotch  deerhound  is  not 
acquired  until  rather  late  in  life.  The  males  almost  exclusively 
are  used  for  coursing,  for,  as  Mr.  McNeill  informs  me,  the 
females  have  net  sufficient  strength  and  weight  to  pull  down  a 
full-grown  deer.  From  the  names  used  in  old  legends,  it  appears, 
as  I  hear  from  Mr.  Cupples,  that,  at  a  very  ancient  period,  the 
males  w^ere  the  most  celebrated,  the  females  being  mentioned 
only  as  the  mothers  of  famous  dogs.  Hence,  during  many 
generations,  it  is  the  male  w^hich  has  been  chiefly  tested  for 
strength,  size,  speed,  and  courage,  and  the  best  will  have  been 
bred  from.  As,  however,  the  males  do  not  attain  their  full 
dimensions  until  rather  late  in  life,  they  will  have  tended,  in 
accordance  with  the  law  often  indicated,  to  transmit  their 
characters  to  their  male  offspring  alone ;  and  thus  the  great 
inequality  in  size  between  the  sexes  of  the  Scotch  deer-hound 
may  probably  be  accounted  for. 

I  hope  that  Mr.  Cupples  will  keep  account  and  history  of  this  f\imous 
to  his  intention  of  publishing  a  full      breed. 


^x8  The  Descent  of  Man.  Part  II. 

The  males  of  some  few  quadrupeds  possess  organs  or  parts 
developed  solely  as  a  means  of  defence  against  the  attacks  of 
other  males.  Some  kinds  of  deer  use,  as  we  have  seen,  the 
upper  branches  of  their  horns  chiefly  or  exclusively  for  de- 
fending themselves ;  and  the  Oryx  antelope,  as  I  am  informed 
by  Mr.  Bartlett,  fences  most  skilfully  with  his  long,  gently 
curved  horns;  but  these  are  likewise  used  as  organs  of  offence. 
The  same  observer  remarks  that  rhinoceroses  in  fighting,  parry 
each  other's  sidelong  blows  with  their  horns,  which  clatttr 
loudly  together,  as  do  the  tusks  of  boars.  Although  wild  boars 
fight  desperately,  they  seldom,  according  to  Brehm,  receive  fatal 
wounds,  as  the  blows  fall  on  each  other's  tusks,  or  on  the  layer 
of  gristly  skin  covering  the  shoulder,   called   by  the  German 

hunters,  the  shield;  and 
here  we  have  a  part  spe- 
cially modified  for  defence. 
With  boars  in  the  prime 
of  life  (see  fig.  65)  the 
tusks  in  the  lower  jaw  are 
used  for  fighting,  but  they 
become  in  old  age,  as 
Brehm  states,  so  much 
curved  inwards  and  up- 
wards over  the  snout, 
that  they  can  no  longer 
be  med  in  this  way 
They  may,  however,  still 

Fig.  65.     Head  of  Common  -wild  lior^r,  in  prime  , 

of  life  cfiom  Brehm).  scrve,    and     even    more 

effectively,  as  a  means 
of  defence.  In  compensation  for  the  loss  of  the  lower  tu.^ks 
as  weapons  of  offence,  those  in  the  upper  jaw,  which  always 
project  a  little  laterally,  increase  in  old  age  so  much  in  length 
and  curve  so  much  upwards,  that  they  can  be  used  for  attack. 
Nevertheless,  an  old  boar  is  not  so  dangerous  to  man  as  one  at 
the  age  of  six  or  seven  years."^ 

In  the  full-grown  male  Baburisa  pig  of  Celebes  (fig.  66),  the 
lower  tusks  are  formidable  weapons,  like  those  of  the  European 
boar  in  the  prime  of  life,  whilst  the  upper  tusks  are  so  long 
and  have  their  points  so  much  curled  inwards,  sometimes  even 
touching  the  forehead,  that  they  are  utterly  useless  as  weapons 
of  attack.  They  more  nearly  resemble  horns  than  teeth,  and 
are  so  manifestly  useless  as  teeth,  that  the  animal  was  formerly 
supposed  to  rest  his  head  by  hooking  them  on  to  a  branch! 
Their  convex  surfaces,  however,  if  the  head  were  held  a  little 
39  Brehm,  '  Thierleben,'  B.  ii.  s.  729-732. 


Chap.  XVII.        Mammals — Means  of  Defence.  519 

laterally,  would  serve  as  an  excellent  guard ;  and  hence,  perhaps, 
it  is  that  in  old  animals  they  "  are  generally  broken  off,  as  if  by 
"  fighting."^"  Here,  then,  we  have  the  curious  case  of  the 
upper  tusks  of  the  Babirusa  regularly  assuming  during  the 
prime  of  life,  a  structure  which  apparently  renders  them  fitted 
only  for  defence ;  whilst  in  the  European  boar  the  lower  tusks 
assume  in  a  less  degree  and  only  during  old  age,  nearly  the 
same  form,  and  then  serve  in  like  manner  solely  for  defence. 


tj6.    6kull  of  the  Babirusa  Pig  (from  Wallace's  '  Malay  Archipelago  ')• 


In  the  wart-hog  {Phacochoerus  cethiopicus,  fig.  67)  the  tusks  in 
the  upper  jaw  of  the  male  curve  upwards  during  the  prime  of 
life,  and  from  being  pointed  serve  as  formidable  weapons.  The 
tusks  in  the  lower  jaw  are  sharper  than  those  in  the  upper,  but 
from  their  shortness  it  seems  hardly  possible  that  they  can  be 
used  as  weapons  of  attack.  They  must,  however,  greatly 
strengthen  those  in  the  upper  jaw,  from  being  ground  so  as  to 
fit  closely  against  their  bases.     Neither  the  upper  nor  the  lower 

*"  See  Mr.  WalLace's  interesting  account  of  this  animal,  '  The  Malay 
Archipelago,'  1869,  vol.  i.  p.  435. 


520  The  Descejit  of  Man.  Part  II. 

tusks  appear  to  have  been  specially  modified  to  act  as  guards, 
though  DO  doubt  they  are  to  a  certain  extent  used  for  this  jDur- 
pose.  But  the  wart-hog  is  not  destitute  of  other  special  means 
of  protection,  for  it  has,  on  each  side  of  the  face,  beneath  the 
eyes,  a  rather  stiff,  yet  flexible,  cartilaginous,  oblong  pad 
(fig.  67),  which  projects  two  or  three  inches  outwards;  and 
it  appeared  to  Mr.  Bartlett  and  myself,  when  viewing  the  living 
animal,  that  these  pads,  when  struck  from  beneath  by  the  tusks 
of  an  opponent,  would  be  turned  upwards,  and  would  thus 
admirably  protect  the  somewhat  prominent  eyes.  I  may  add, 
on  the  authority  of  Mr.  Bartlett,  that  these  boars  when  figbting 
stand  directly  face  to  face. 


Fig.  67.  Head  of  female  ^Ethiopian  wart-hog,  from  'Proc.  Zool.  Soc'  1869,  shewing 
the  same  characters  as  the  male,  though  on  a  reduced  scale. 

N.B.  When  the  engraving  was  first  made,  1  was  under  the  impression  that  it  repre- 
sented the  male. 

Lastly,  the  African  river-hog  {Potomoclwerus  penicillatus)  has  a 
hard  cartilaginous  knob  on  each  side  of  the  face  beneath  the 
eyes,  which  answers  to  the  flexible  pad  of  the  wart-hog ;  it  has 
also  two  bony  prominences  on  the  upper  jaw  above  the  nostrils. 
A  boar  of  this  species  in  the  Zoological  Gardens  recently  broke 
into  the  cage  of  the  wart-hog.  They  fought  all  night  long,  and 
were  found  in  the  morning  much  exhausted,  but  not  seriously 
wounded.  It  is  a  significant  fact,  as  shewing  the  purpose  of  the 
above-described  projections  and  excrescences,  that  these  were 
covered  with  blood,  and  were  scored  and  abraded  in  an  extra- 
ordinary manner. 

Although  the  males  of  so  many  members  of  the  pig  family  are 


Chap.  XVII.     Mammals — Means  of  Defence.  521 

provided  with  weapons,  and  as  we  have  just  seen  with  means  of 
defence,  these  weapons  seem  to  have  been  acquired  witliin  a 
ratlier  late  geological  period.  Dr.  Forsyth  Major  specifies'^ 
several  miocene  species,  in  none  of  which  do  the  tusks  appear 
to  have  been  largely  developed  in  the  males  ;  and  Prof.  Eutimeyer 
was  formerly  struck  with  this  same  fact. 

The  mane  of  the  lion  forms  a  good  defence  against  the  attacks 
of  rival  lions,  the  one  danger  to  which  he  is  liable;  for  the 
males,  as  Sir  A.  Smith  informs  me,  engage  in  terrible  battles, 
and  a  young  lion  dares  not  approach  an  old  one.  In  1857  a 
tiger  at  Bromwich  broke  into  the  cage  of  a  lion  and  a  fearful 
scene  ensued :  "-the  lion's  mane  saved  his  neck  and  head  from 
"  being  much  injured,  but  the  tiger  at  last  succeeded  in  ripping 
"  up  his  belly,  and  in  a  few  minutes  he  was  dead."  ^^  The  broad 
ruff  round  the  throat  and  chin  of  the  Canadian  lynx  (^Felis 
canadensis)  is  much  longer  in  the  male  than  in  the  female ;  but 
whether  it  serves  as  a  defence  I  do  not  know.  Male  seals  are 
well  known  to  fight  desperately  together,  and  the  males  of 
certain  kinds  (Oiarta  jubata)*^  have  great  manes,  whilst  the 
females  have  small  ones  or  none.  The  male  baboon  of  the  Cape 
of  Good  Hope  (Cynocephalus  'porcarius)  has  a  much  longer  mane 
and  larger  canine  teeth  than  the  female ;  and  the  mane  probably 
serves  as  a  protection,  for,  on  asking  the  keepers  in  the  Zoo- 
logical Gardens,  without  giving  them  any  clue  to  my  object, 
whether  any  of  the  monkeys  especially  attacked  each  other  by 
the  nape  of  the  neck,  I  was  answered  that  this  was  not  the  case, 
except  with  the  above  baboon.  In  the  Hamadryas  baboon, 
Ehrenberg  compares  the  mane  of  the  adult  male  to  that  of  a 
young  lion,  whilst  in  the  young  of  both  sexes  and  in  the  female 
the  mane  is  almost  absent. 

It  appeared  to  me  probable  that  the  immense  woolly  mane  of 
the  male  American  bison,  which  reaches  almost  to  the  ground, 
and  is  much  more  developed  in  the  males  than  in  the  females, 
served  as  a  protection  to  them  in  their  terrible  battles ;  but  an 
experienced  hunter  told  Judge  Caton  that  he  had  never  observed 
anything  which  favoured  this  belief.  The  stallion  has  a  thicker 
and  fuller  mane  than  the  mare;  and  I  have  made  particular 
inquiries  of  two  great  trainers  and  breeders,  who  have  had  charge 
of  many  entire  horses,  and  am  assured  that  they  "  invariably 

■•i  '  Atti  della  Soc.  Italiana  di  Sc.  *^  Dr.  Murie,  on   Otaria,   '  Proc. 

Nat.'  1873,  vol.  xv.  fasc.  iv.  Zoolog.   Soc'   1869,  p.  109.     Mr.  J. 

^2  'The  Times,'  Nov.   10th,  1857.  A.  Allen,  in  the  paper  above  quoted 

In   regard  to  the  Canada  lynx,  see  (p.   75),   doubts  whether   the   hair, 

Audubon    and     Bachman,      '  Quad-  which  is  longer  on  the  nock   in  the 

rupeds  of    N.    America,'    1846,    p.  male  than  in  the  female,  deserves  to 


ioi 


be  called  a  mane. 


522  The  Descent  of  Man.  Part  II. 

"  endeaYour  to  seize  one  another  by  the  neck."  It  does  not, 
however,  follow  from  the  foregoing  statements,  that  when 
the  hair  on  the  neck  serves  as  a  defence,  that  it  was  originally 
developed  for  this  purpose,  though  this  is  probable  in  some  cases, 
as  in  that  of  the  lion.  I  am  informed  by  Mr.  McNeill  that  the 
long  hairs  on  the  throat  of  the  stag  (Cer<;^^s  dwphmC)  serve  as  a 
great  protection  to  him  when  hunted,  for  the  dogs  generally 
endeavour  to  seize  him  by  the  throat ;  but  it  is  not  probable  that 
these  hairs  were  specially  developed  for  this  purpose ;  otherwise 
the  young  and  the  females  would  have  been  equally  protected. 

Choice  in  Pairing  hy  either  Sex  of  Quadrupeds. — Before  de- 
scribing in  the  next  chapter,  the  diiferences  between  the  sexes  in 
voice,  odours  emitted,  and  ornaments,  it  will  be  convenient  here 
to  consider  whether  the  sexes  exert  any  choice  in  their  unions. 
Does  the  female  prefer  any  particular  male,  either  before  or 
after  the  males  may  have  fought  together  for  supremar^y;  or 
does  the  male,  when  not  a  polygamist,  select  any  particular 
female?.  The  general  impression  amongst  breeders  seems  to  be 
that  the  male  accepts  any  female ;  and  this  owing  to  his  eager- 
ness, is,  in  most  cases,  probably  the  truth.  Whether  the  female 
as  a  general  rule  indifferently  accepts  any  male  is  much  more 
doubtful.  In  the  fourteenth  chapter,  on  Birds,  a  considerable 
body  of  direct  and  indirect  evidence  was  advanced,  shewing 
that  the  female  selects  her  partner ;  and  it  would  be  a  strange 
anomaly  if  female  quadrupeds,  which  stand  higher  in  the  scale 
and  have  higher  mental  powers,  did  not  generally,  or  at  least 
often,  exert  some  choice.  The  female  could  in  most  cases 
escape,  if  wooed  by  a  male  that  did  not  please  or  excite  her ;  and 
when  pursued  by  several  males,  as  commonly  occurs,  she  would 
often  have  the  opportunity,  whilst  they  were  fighting  together, 
of  escaping  with  some  one  male,  or  at  least  of  temporarily  pairing 
with  him.  This  latter  contingency  has  often  been  observed  in 
Scotland  with  female  red-deer,  as  I  am  informed  by  Sir  Philip 
Egerton  and  others.^ 

It  is  scarcely  possible  that  much  should  be  known  about 
female  quadrupeds  in  a  state  of  nature  making  any  choice  in 
their  marriage  unions.  The  following  curious  details  on  the 
courtship   of  one  of  the  eared   seals   {CaUorhinus  ursinus)   are 

**  Mr.  Boner,  in  his  excellent  "  sanctuary  of  his  harem,  and  caii- 
description  of  the  habits  of  the  red-  "  ries  oft"  trophy  after  trophy."  Ex- 
deer  in  Germany  (' Forest  Creatures,'  actly  the  same  thing  occurs  with 
1861,  p.  81)  says,  "  while  the  stag  seals,  see  Mr.  J.  A.  Allen,  ibid.  p. 
"is  defending  his  rights  against  one  100. 
" intruder,     another     invades     the 


Chap.  XVIL     Mammals — Choice  in  Pairing.  523 

given  ^^  on  the  authority  of  Capt.  Bryant,  who  had  ample 
opportunities  for  observation.  He  says,  "  Many  of  the  females 
"  on  their  arrival  at  the  island  where  they  breed  appear  desirous 
"■  of  returning  to  some  particular  male,  and  frequently  climb  the 
"  outlying  rocks  to  overlook  the  rookeries,  calling  out  and 
"  listening  as  if  for  a  familiar  voice.  Then  changing  to  another 
"  place  they  do  the  same  again  ....  As  soon  as  a  female  reaches 
"  the  shore,  the  nearest  male  goes  down  to  meet  her,  making 
"  meanwhile  a  noise  like  the  clucking  of  a  hen  to  her  chickens. 
"  He  bows  to  her  and  coaxes  her  until  he  gets  between  her  and 
"  the  water  so  that  she  cannot  escape  him.  Then  his  manner 
"  changes,  and  with  a  harsh  growl  he  drives  her  to  a  place  in 
"  his  harem.  This  continues  until  the  lower  row  of  harems  is 
"  nearly  full.  Then  the  males  higher  ui)  select  the  time  when 
"  their  more  fortunate  neighbours  are  off  their  guard  to  steal 
"  their  wives.  This  they  do  by  taking  them  in  their  mouths 
"  and  lifting  them  over  the  heads  of  the  other  females,  and 
''  carefully  placing  them  in  their  own  harem,  carrying  them  as 
"  cats  do  their  kittens.  Those  still  higher  up  pursue  the  same 
''  method  until  the  whole  space  is  occupied.  Frequently  a 
"  struggle  ensues  between  two  males  for  the  possession  of  the 
"  same  female,  and  both  seizing  her  at  once  pull  her  in  two  or 
"  terribly  lacerate  her  with  their  teeth.  When  the  space  is  all 
*'  filled,  the  old  male  walks  around  complacently  reviewing  his 
"  family,  scolding  those  who  crowd  or  disturb  the  others,  and 
"  fiercely  driving  off  all  intruders.  This  surveillance  always 
"  keeps  him  actively  occupied." 

As  so  little  is  known  about  the  courtship  of  animals  in  a  state 
of  nature,  I  have  endeavoured  to  discover  how  far  our  domesti- 
cated quadrupeds  evince  any  choice  in  their  unions.  Dogs 
offer  the  best  opportunity  for  observation,  as  they  are  carefully 
attended  to  and  well  understood.  Many  breeders  have  expressed 
a  strong  opinion  on  this  head.  Thus,  Mr.  Mayhew  remarks, 
"  The  females  are  able  to  bestow  their  affections;  and  tender 
"  recollections  are  as  potent  over  them  as  they  are  known  to  be 
"  in  other  cases,  where  higher  animals  are  concerned.  Bitches 
"  are  not  always  prudent  in  their  loves,  but  are  apt  to  fling 
"  themselves  away  on  curs  of  low  degree.  If  reared  with  a 
"  companion  of  vulgar  appearance,  there  often  springs  up 
*'  between  the  pair  a  devotion  which  no  time  can  afterwards 
*'  subdue.  The  passion,  for  such  it  really  is,  becomes  of  a  more 
"  than  romantic  endurance."  Mr.  Mayhew,  who  attended 
chiefly  to  the  smaller  breeds,  is  convinced  that  the  females  are 

*^  Mr.  J.  A.  Allen  in  '  Bull,  Mus.  Comp.  Zoolog.  of  Cambridge,  United 
States,'  vol.  ii.  No.  1,  p.  99. 


524  The  Descent  of  Man.  Paet  II. 

strongly  attracted  by  males  of  a  large  size.  ^^  The  well-known 
veterinary  Blaine  states'*^  that  his  own  female  pug  became  so 
attached  to  a  spaniel,  and  a  female  setter  to  a  cur,  that  in 
neither  case  would  they  pair  with  a  dog  of  their  own  breed 
until  several  weeks  had  elapsed.  Two  similiar  and  trustworthy 
accounts  have  been  given  me  in  regard  to  a  female  retriever 
and  a  spaniel,  both  of  which  became  enamoured  with  terrier- 
dogs. 

Mr.  Cupples  informs  me  that  he  can  personally  vouch  for  the 
accuracy  of  the  following  more  remarkable  case,  in  which  a 
valuable  and  wonderfully-intelligent  female  terrier  loved  a 
retriever  belonging  to  a  neighbour  to  such  a  degree,  that  she 
had  often  to  be  dragged  away  from  him.  After  their  permanent 
separation,  although  repeatedly  shewing  milk  in  her  teats,  she 
would  never  acknowledge  the  courtship  of  any  other  dog,  and  to 
the  regret  of  her  owner  never  bore  puppies.  Mr.  Cupples  also 
states,  that  in  1868,  a  female  deerhound  in  his  kennel  thrice  pro- 
duced puppies,  and  on  each  occasion  shewed  a  marked  preference 
for  one  of  the  largest  and  handsomest,  but  not  the  most  eager, 
of  four  deerhounds  living  with  her,  all  in  the  prime  of  life. 
Mr.  Cupples  has  observed  that  the  female  generally  favours  a  dog 
whom  she  has  associated  with  and  knows;  her  shyness  and 
timidity  at  first  incline  her  against  a  strange  dog.  The  male,  on 
the  contrary,  seems  rather  inclined  towards  strange  females.  It 
appears  to  be  rare  when  the  male  refuses  any  particular  female, 
but  Mr.  Wright,  of  Yeldersley  House,  a  great  breeder  of  dogs, 
informs  me  that  he  has  known  some  instances  ;  he  cites  the  case  of 
one  of  his  own  deerhounds,  who  would  not  take  any  notice  of  a 
particular  female  mastiff,  so  that  another  deerhound  had  to  be 
employed.  It  would  be  superfluous  to  give,  as  I  could,  other 
instances,  and  I  will  only  add  that  Mr.  Barr,  who  has  carefully 
bred  many  bloodhounds,  states  that  in  almost  every  instance 
particular  individuals  of  opposite  sexes  shew  a  decided  prefer- 
ence for  each  other.  Finally,  Mr.  Cupples,  after  attending  to 
this  subject  for  another  year,  has  written  to  me,  "  I  have  had 
"  full  confirmation  of  my  former  statement,  that  dogs  in  breeding 
*'  form  decided  preferences  for  each  other,  being  often  influenced 
"  by  size,  bright  colour,  and  individual  characters,  as  well  as  by 
"  the  degree  of  their  previous  familiarity." 

In  regard  to  horses,  Mr.  Blenkiron,  the  greatest  breeder  of 
race-horses  in  the  world,  informs  me  that  stallions  are  so 
frequently  capricious  in  their  choice,  rejecting  one  mare  and 

**  'Dogs:  their  Management,' by  ^^  Quoted  by  Alex.  Walker  'On 

E.  Mayhew,   M.R.C.V.S.,  2nd   edit.'     Intermarriage,'   1838,   p.   276;    see 
1864,  pp.  187-192.  also  p.  244. 


Chap.  XVI II.     Ma^nmals — Choice  in  Pairing.  525 

without  cany  apparent  cause  taking  to  another,  that  various  artifices 
have  to  be  habitually  used.  The  famous  Monarque,  for  instance, 
would  never  consciously  look  at  the  dam  of  Gladiateur,  and  a 
trick  had  to  be  practised.  We  can  partly  see  the  reason  why 
valuable  race-horse  stallions,  which  are  in  such  demand  as  to  be 
exhausted,  should  be  so  particular  in  their  choice.  Mr.  Blenkiron 
has  never  known  a  mare  reject  a  horse ;  but  this  has  occurred 
in  Mr.  Wright's  stable,  so  that  the  mare  had  to  be  cheated. 
Prosper  Lucas  *^  quotes  various  statements  from  French  autho- 
rities, and  remarks,  "  On  voit  des  etalons  qui  s  eprennent  d'une 
"jument,  et  negligent  toutes  les  autres."  He  gives,  on  the 
authority  of  Baiilen,  similar  facts  in  regard  to  bulls;  and  Mr. 
H.  Eeeks  assures  me  that  a  famous  short-horn  bull  belonging  to 
his  father  "  invariably  refused  to  be  matched  with  a  black  cow." 
Hoffberg,  in  describing  the  domesticated  reindeer  of  Lapland 
says,  "  Foeminae  majores  et  fortiores  mares  prs6  cseteris  admittunt, 
"  ad  eos  confugiunt,  a  junioribus  agitatse,  qui  hos  in  fugam 
"  conjiciunt."^^  A  clergyman,  who  has  bred  many  pigs,  asserts 
that  sows  often  reject  one  boar  and  immediately  accept  another. 
From  these  facts  there  can  be  no  doubt  that,  with  most  of  our 
domesticated  quadrupeds,  strong  individual  antipathies  and  pre- 
ferences are  frequently  exhibited,  and  much  more  commonly  by 
the  female  than  by  the  male.  This  being  the  case,  it  is  impro- 
bable that  the  unions  of  quadrupeds  in  a  state  of  nature  should 
be  left  to  mere  chance.  It  is  much  more  probable  that  the 
females  are  allured  or  excited  by  particular  males,  who  possess 
certain  characters  in  a  higher  degree  than  other  males;  but 
what  these  characters  are,  we  can  seldom  or  never  discover  with 
certainty. 


CHAPTER  XVIII. 

Secondary  Sexual  Characters  of  Mammals — continued. 

Voice — Remarkable  sexual  peculiarities  in  seals — Odour — Development 
of  the  hair — Colour  of  the  hair  and  skin — Anomalous  case  of  the  temale 
being  more  ornamented  than  the  male — Colour  and  ornaments  due  to 
sexual  selection — Colour  acquired  for  the  sake  of  protection — Colour, 
though  common  to  both  sexes,  often  due  to  sexual  selection — On  the 
disappearance  of  spots  and  stripes  in  adult  quadrupeds — On  the  colours 
and  ornaments  of  the  Quadrumana — Summary. 

Quadrupeds  use  their  voices  for  various  purposes,  as  a  signal  of 
danger,  as  a  call  from  one  member  of  a  troop  to  another,  or  from 
the  mother  to  her  lost  offspring,  or  from  the  latter  for  protection 

^*  '  Traite  de  I'Hered.  Nat.'  torn.  ■•»  '  Amoenitates    Acad.'    vol.    iv. 

ii.  1850,  p.  296.  1788,  p.  160. 


526  TJie  Descent  of  Man.  Tart  II. 

to  their  motlier;  but  such  uses  need  not  here  be  considered. 
We  are  concerned  only  with  the  difference  between  the  voices  of 
the  sexes,  for  instance  between  that  of  the  lion  and  lioness,  or  of 
the  bull  and  cow.  Almost  all  male  animals  use  their  voices 
much  more  diu'ing  the  rutting-season  than  at  any  other  time  ; 
and  some,  as  the  giraffe  and  porcupine,^  are  said  to  be  completely 
mute  excepting  at  this  season.  As  the  throats  {i.e.  the  larynx 
and  thyroid  bodies  -)  of  stags  jDeriodically  become  enlarged  at  the 
beginning  of  the  breeding-season,  it  might  be  thought  that  their 
powerful  voices  must  be  somehow  of  high  importance  to  them  ; 
but  this  is  very  doubtful.  From  information  given  to  me  by 
two  experienced  observers,  Mr.  McNeill  and  Sir  P.  Egerton,  it 
seems  that  young  stags  under  three  years  old  do  not  roar 
or  bellow ;  and  that  the  old  ones  begin  bellowing  at  the  com- 
mencement of  the  breeding-season,  at  first  only  occasionally  and 
moderately,  whilst  they  restlessly  wander  about  in  search  of  the 
females.  Their  battles  are  prefaced  by  loud  and  prolonged 
bellowing,  but  during  the  actual  conflict  they  are  silent.  Animals 
of  all  kinds  which  habitually  use  their  voices  utter  various 
noises  under  any  strong  emotion,  as  when  enraged  and  pre- 
paring to  fight;  but  this  may  merely  be  the  result  ofnervons 
excitement,  which  leads  to  the  spasmodic  contraction  of  almost 
all  the  muscles  of  the  body,  as  when  a  man  grinds  liis  teeth  and 
clenches  his  fists  in  rage  or  agony.  No  doubt  stags  challenge 
each  other  to  mortal  combat  by  bellowing ;  but  those  with  the 
more  powerful  voices,  unless  at  the  same  time  the  stronger, 
better-armed,  and  more  courageous,  would  not  gain  any  advan- 
tage over  their  rivals. 

It  is  possible  that  the  roaring  of  the  lion  may  be  of  some 
service  to  him  by  striking  terror  into  his  adversary ;  for  when 
enraged  he  likewise  erects  his  mane  and  thus  instinctively  tries 
to  make  himself  appear  as  terrible  as  possible.  But  it  can 
hardly  be  supposed  that  the  bellowing  of  the  stag,  even  if  it  be 
of  service  to  him  in  this  way,  can  have  been  important  enough 
to  have  led  to  the  periodical  enlargement  of  the  throat.  Some 
writers  suggest  that  the  bellowing  serves  as  a  call  to  the  female ; 
but  the  experienced  observers  above  quoted  inform  me  that 
female  deer  do  not  search  for  the  male,  though  the  males  search 
eagerly  for  the  females,  as  indeed  might  be  expected  from  what 
we  know  of  the  habits  of  other  male  quadrupeds.  The  voice  of 
the  female,  on  the  other  hand,  quickly  brings  to  her  one  or  more 
stags,^  as  is  well  known  to  the  hunters  who  in  wild  countries 

*  Owen,     '  Anatomy    of     Verte-  3  See,    for    instance,    Major   W. 

brates,'  vol.  iii.  p.  585.  Ross    King    ('  The     Sportsman    in 

2  Ibid.  p.  595.  Canada'  1866,  p.  53,  131)  on  the 


(JuAP.  XV 1 1 1 .      Maminals —  Vocal  Organs.  5 27 

imitate  her  cry.  If  we  could  believe  that  the  male  had  the 
power  to  excite  or  allure  the  female  by  his  voice,  the  periodical 
enlargement  of  his  vocal  organs  wol^ld  be  intelligible  on  the 
principle  of  sexual  selection,  together  wdth  inheritance  limited  to 
the  same  sex  and  season ;  but  we  have  no  evidence  in  favour  ot 
this  view.  As  the  case  stands,  the  loud  voice  of  the  stag  daring 
the  breeding-season  does  not  seem  to  be  of  any  special  service  to 
him,  either  during  his  courtship  or  battles,  or  in  any  other  way. 
But  may  we  not  believe  that  the  frequent  use  of  the  voice,  undci* 
the  strong  excitement  of  love,  jealousy,  and  rage,  continued 
during  many  generations,  may  at  last  have  produced  an  in- 
herited effect  on  the  vocal  organs  of  the  stag,  as  well  as  of  other 
male  animals?  This  appears  to  me,  in  our  present  state  ot 
knowledge,  the  most  jorobable  view. 

The  voice  of  the  adult  male  gorilla  is  tremendous,  and  he  is 
furnished  with  a  laryngeal  sack,  as  is  the  adult  male  orang.* 
The  gibbons  rank  among  the  noisiest  of  monkeys,  and  the 
Sumatra  species  (^Hylobates  syndaciylus)  is  also  furnished  Avith  an 
air  sack ;  but  Mr.  Blyth,  who  has  had  opportunities  for  observa- 
tion, does  not  believe  that  the  male  is  noisier  than  the  female. 
Hence,  these  latter  monkeys  probably  use  their  voices  as  a 
mutual  call ;  and  this  is  certainly  the  case  with  some  quadrupeds, 
for  instance  the  beaver,^  Another  gibbon,  the  fJ.  agilis,  is  re- 
markable, from  having  the  power  of  giving  a  complete  and 
correct  octave  of  musical  notes,"  which  we  may  reasonably  sus- 
pect serves  as  a  sexual  charm ;  but  I  shall  have  to  recur  to  this 
subject  in  the  next  chapter.  The  vocal  organs  of  the  American 
Mycetes  caraya  are  one-third  larger  in  the  male  than  in  the 
female,  and  are  wonderfully  powerful.  These  monkeys  in  warm 
weather  make  the  forests  resound  at  morning  and  evening  with 
their  overwhelming  voices.  The  males  begin  the  dreadful  con- 
cert, and  often  continue  it  during  many  hours,  the  females, 
sometimes  joining  in  with  their  less  powerful  voices.  An 
excellent  observer,  Eengger,^  could  not  perceive  that  they  were 
excited  to  begin  by  any  special  cause  ;  he  thinks  that,  like  many 
birds,  they  delight  in  their  own  music,  and  try  to  excel  each 
other.  Whether  most  of  the  foregoing  monkeys  have  acquired 
their  powerful  voices  in  order  to  beat  their  rivals  and  charm  the 
females — or  whether  the  vocal  organs  have  been  strengthened 


habits  of  the  moose  and  wild  rein-  ®  C.   L.   Martin,   '  General    Intro- 
deer,  duction  to  the  Nat.  Hist,  of  Mamm. 

■*  Owen,    '  Anatomy    of     Verte-  Animals,'  1841,  p.  431. 
brates.' vol.  iii.  p.  600.  '   '  Naturgeschiclite    der     Sauge- 

*  Mr.  Green,  in  'Journal  of  Linn.  thiere  von  Paraguay,'  1830,  s.   15, 

Soc'  vol.  X.  Zoology,  1869,  p.  362.  21. 


528  The  Descent  of  Man.  Part  II. 

and  enlarged  through  the  inherited  effects  of  long-continued 
use  without  any  particular  good  being  thus  gained — I  will  not 
pretend  to  say ;  but  the  former  Tiew,  at  least  in  the  case  of  the 
Hylohates  agilis,  seems  the  most  j^robable. 

I  may  here  mention  two  very  curious  sexual  peculiarities 
occurring  in  seals,  because  they  have  been  supposed  by  some 
writers  to  affect  the  voice.  The  nose  of  the  male  sea-elephant 
(^Macrorhmus  prohoscideus)  becomes  greatly  elongated  during  the 
breeding-season,  and  can  then  be  erected.  In  this  state  it  is 
sometimes  a  foot  in  length.  The  female  is  not  thus  provided  at 
any  period  of  life.  The  male  makes  a  wild,  hoarse,  gurgling 
noise,  which  is  audible  at  a  great  distance  and  is  believed  to  be 
strengthened  by  the  proboscis ;  the  voice  of  the  female  being 
different.  Lesson  compares  the  erection  of  the  proboscis,  with 
the  swelling  of  the  wattles  of  male  galhnaceous  birds  whilst 
courting  the  females.  In  another  allied  kind  of  seal,  the  bladder- 
nose  {Ci/stophora  cristata),  the  head  is  covered  by  a  great  hood 
or  bladder.  This  is  supported  by  the  septum  of  the  nose,  which 
is  produced  far  backwards  and  rises  into  an  internal  crest  seven 
inches  in  height.  The  hood  is  clothed  with  short  hair,  and  is 
muscular ;  it  can  be  inflated  until  it  more  than  equals  the 
whole  head  in  size !  The  males  when  rutting,  fight  furiously  on 
the  ice,  and  their  roaring  "  is  said  to  be  sometimes  so  loud  as  to 
"  be  heard  four  miles  off.'  When  attacked  they  likewise  roar  or 
bellow ;  and  whenever  irritated  the  bladder  is  inflated  and 
quivers.  Some  naturalists  believe  that  the  voice  is  thus 
strengthened,  but  various  other  uses  have  been  assigned  to 
this  extraordinary  structure.  Mr.  E.  Brown  thinks  that  it 
serves  as  a  protection  against  accidents  of  all  kinds ;  but  this  is 
not  probable,  for,  as  I  am  assui'ed  by  INIr.  Lament  who  killed 
600  of  these  animals,  the  hood  is  rudimentary  in  the  females, 
and  it  is  not  developed  in  the  males  during  youth.^ 

Odour. — With  some  animals,  as  with  the  notorious  skunk  of 
America,  the  overwhelming  odour  which  they  emit  appears  to 
serve  exclusively  as  a  defence.  With  shrew-mice  (Sorex)  both 
sexes  possess  abdominal  scent-glands,  and  there  can  be  little 
doubt,  from  the  rejection  of  their  bodies  by  birds  and  beasts  of 
prey,  that  the  odour  is  protective;  nevertheless,  the  glands 
become  enlarged  in  the  males  during  the  breeding-season.     In 

^  On    the    sea-elephant,    see    an  1824,    p.     94.     Pennant    has    also 

article    by  Lesson,  in  'Diet..  Class.  collected     information     from     the 

Hist.  Nat.'  torn.  xiii.  p.  418.     For  sealers  on  this  animal.     The  fullest 

the    Cystophora    or    Stemmatopus,  account  is  given  by  Mr.  Brown,  in 

see  Dr.  Dekay,  '  Annals  of  Lyceum  '  Proc.  Zoolog.  Soc'  1868,  p.  435. 
of   Nat.    Hist.    New   York,'   vol.   i. 


Chap.  XVIII.     Mammals — Odours  emitted.  529 

many  other  quadrupeds  the  glands  are  of  the  same  size  in  both 
sexes,®  but  their  uses  are  not  known.  In  otlier  species  the 
glands  are  confined  to  the  males,  or  are  more  developed  than  in 
the  females  ;  and  they  almost  always  become  more  active  during 
the  rutting-season.  At  this  period  the  glands  on  the  sides  of 
the  face  of  the  male  elephant  enlarge,  and  emit  a  secretion 
having  a  strong  musky  odour.  The  males,  and  rarely  the 
females,  of  many  kinds  of  bats  have  glands  and  protrudable 
sacks  situated  in  various  parts ;  and  it  is  believed  that  these  are 
odoriferous. 

The  rank  effluvium  of  the  male  goat  is  well  known,  and  that 
of  certain  male  deer  is  wonderfully  strong  and  persistent.  On 
the  banks  of  the  Plata  I  perceived  the  air  tainted  with  the 
odour  of  the  male  Cervus  camjyestris,  at  half  a  mile  to  leeward  of 
a  herd ;  and  a  silk  handkerchief,  in  which  I  carried  home  a  skin, 
though  often  used  and  washed,  retained,  when  first  unfolded, 
traces  of  the  odour  for  one  year  and  seven  months.  This 
animal  does  not  emit  its  strong  odour  until  more  than  a  year 
old,  and  if  castrated  whilst  young  never  emits  iV^  Besides 
the  general  odour,  permeating  the  whole  body  of  certain  rumi- 
nants (for  instance.  Bos  moschatus)  in  the  breeding-season,  many 
deer,  antelopes,  sheep,  and  goats,  possess  odoriferous  glands  in 
various  situations,  more  especially  on  their  faces.  The  so-called 
tear-sacks,  or  suborbital  pits,  come  under  this  head.  These 
glands  secrete  a  semi-fluid  fetid  matter  which  is  sometimes  so 
copious  as  to  stain  the  whole  face,  as  I  have  myself  seen  in  an 
antelope.  They  are  "  usually  larger  in  the  male  than  in  the 
"  female,  and  their  development  is  checked  by  castration."" 
According  to  Desmarest  they  are  altogether  absent  in  the  female 
of  Ant  Hope  suhgutturosa.  Hence,  there  can  be  no  doubt  that 
they  stand  in  close  relation  with  the  reproductive  functions. 
They  are  also  sometimes  present,  and  sometimes  absent,  in 
nearly-allied  forms.  In  the  adult  male  musk-deer  (Mo.schus 
moschiferus),  a  naked  space  round  the  tail  is  bedewed  with  an 

^  As  with  the  castoreum   of  the  '°  Rengger,  '  Naturgeschichte  der  " 

beaver,   see    Mr.    L.     H.     Morgan's  Saiigethiere  von    Paraguay,'    1830, 

most  interesting  work,  'The  A  me-  s.   355.     This   observer    also    gives 

rican  BeaA'er,'  1868,  p.  300.     Pallas  some  curious  particulars  in  regard 

('  Spic.  Zoolog.'  fasc.  viii.   1779,  p.  to  the  odour. 

23)  has  well  discussed  the  odor! fer-  '^  Owen,  'Anatomy  of  Verte- 
ous  glands  of  mammals.  Owen  brates,'  vol.  iii.  p.  632.  See,  also 
('  Anat.  of  Vertebrates,'  vol.  iii,  p.  Dr.  Murie's  observations  on  these 
634)  also  gives  an  account  of  these  glands  in  the  '  Proc.  Zoolog.  Soc' 
glands,  including  those  of  the  ele-  1870,  p.  340.  '  Desmarest,  On  the 
phant,  and  (p.  763)  those  of  shrew-  Antilope  suhgutturosa,  '  Mamma- 
mice.  On  Bats,  Mr.  Dobson  in  logie,'  1820,  p.  455. 
'  Proc.  Zoolog.  Soc'  1873,  p.  241. 


5  30  .     The  Descettt  of  Man.  Part  II. 

odoriferous  fluid,  whilst  in  tlie  adult  female,  and  in  the  male 
until  two  years  old,  this  space  is  covered  with  hair  and  is  not 
odoriferous.  The  proper  musk-sack  of  this  deer  is  from  its 
position  necessarily  confined  to  the  male,  and  forms  an  additional 
scent-organ.  It  is  a  singular  fact  that  the  matter  secreted  by 
this  latter  gland  does  not,  according  to  Pallas,  change  in  con- 
sistence, or  increase  in  quantity,  during  the  rutting-season ; 
nevertheless  this  naturalist  admits  that  its  presence  is  in  some 
way  connected  with  the  act  of  reproduction.  He  gives,  however, 
only  a  conjectural  and  unsatisfactory  exjDlanation  of  its  use.^^ 

In  most  cases,  when  only  the  male  emits  a  strong  odour 
during  the  breeding-season,  it  probably  serves  to  excite  or  allure 
the  female.  We  must  not  judge  on  this  head  by  our  own  taste, 
for  it  is  well  kno^^ni  that  rats  are  enticed  by  certain  essential 
oils,  and  cats  by  valerian,  substances  far  from  agreeable  to  us ; 
and  that  dogs,  though  they  will  not  eat  carrion,  sniff  and  roll  on 
it.  From  the  reasons  given  when  discussing  the  voice  of  the 
stag,  we  may  reject  the  idea  that  the  odour  serves  to  bring  the 
iemales  from  a  distance  to  the  males.  Active  and  long-continued 
use  cannot  here  have  come  into  play,  as  in  the  case  of  the  vocal 
organs.  The  odour  emitted  must  be  of  considerable  importance 
to  the  male,  inasmuch  as  large  and  complex  glands,  fui'nished 
with  muscles  for  everting  the  sack,  and  for  closing  or  opening 
the  orifice,  have  in  some  cases  been  developed.  The  develop- 
ment of  these  organs  is  intelligible  through  sexual  selection,  if 
the  most  odoriferous  males  are  the  most  successful  in  winning 
the  females,  and  in  leaving  offspring  to  inherit  their  gradually- 
perfected  glands  and  odours. 

Development  of  the  Hair. — We  have  seen  that  male  quadrupeds 
often  have  the  hair  on  their  necks  and  shoulders  much  more 
developed  than  the  females;  and  many  additional  instances 
could  be  given.  Tliis  sometimes  serves  as  a  defence  to  the  male 
during  his  battles  ;  but  whether  the  hair  in  most  cases  has  been 
specially  developed  for  this  purp>OJO,  is  very  doubtful.  We  may 
feel  almost  certain  that  this  is  not  the  case,  when  only  a  thin 
and  narrow  crest  runs  along  the  back  ;  for  a  crest  of  this  kind 
would  afford  scarcely  any  protection,  and  the  ridge  of  the  back 
is  not  a  place  likely  to  be  injured;  nevertheless  such  crests  are 
sometimes  confined  to  the  males,  or  are  much  more  developed 
in  them  than  in  the  females.  Two  antelopes,  the  Tragekqjhus 
sc7nptus'^^  (see  fig.  70,  p.  544)  and  Fortax  pictu,  may  be  given  as 

^2  Pallas,  *  Spicilegia  Zoolog.' fasc.  '^  Dr.  Gray,  'Gleanings  from  the 

xiii.  1799,  p.  24  ;  Desmoulins,  '  Diet.      Menagerie  at  Knowsley,'  pi.  28. 
Class.  d'Hist.  Nat.'  torn.  iii.  p.  586. 


Cii  AP.  X  \"  1 1  i .     Mam  vials — Developmen  t  of  Hair.        5  3  i 

instances,  AVhen  stags,  and  the  males  of  the  wild  goat,  are 
enraged  or  terrified,  these  crests  stand  erect;'*  but  it  cannot  be 
supposed  that  they  have  been  developed  merely  for  the  sake  of 
exciting  fear  in  their  enemies.  One  of  the  above-named  an- 
telopes, the  Portax  incta,  has  a  large  well-defined  brush  of  black 
hair  on  the  throat,  and  this  is  much  larger  in  the  male  than  in 
the  female.  In  the  Ammotragus  trugelap/nis  of  North  Africa,  a 
member  of  the  sheep-family,  the  fore-legs  are  almost  concealed 
by  an  extraordinary  growth  of  hair,  which  depends  from  the 
neck  and  upper  halves  of  the  legs ;  but  Mr.  Bartlett  does  not 
believe  that  this  mantle  is  of  the  least  use  to  the  male,  in  whom 
it  is  much  more  developed  than  in  the  female. 

Male  quadrupeds  of  many  kinds  differ  from  the  females  in 
having  more  hair,  or  hair  of  a  diff"erent  character,  on  certain 
parts  of  their  faces.  '\  hus  the  bull  alone  has  curled  hair  on  the 
forehead.'^  In  three  closely-allied  sub-genera  of  the  goat  family, 
only  the  males  possess  beards,  sometimes  of  large  size ;  in  two 
other  sub-genera  both  sexes  have  a  beard,  but  it  disappears  in 
some  of  the  domestic  breeds  of  the  common  goat ;  and  neither 
sex  of  the  Hemitragus  has  a  beard.  In  the  ibex  the  beard  is  not 
developed  during  the  summer,  and  it  is  so  small  at  other  times 
that  it  may  be  called  rudimentary.'*^  With  some  monkeys  the 
beard  is  confined  to  the  male,  as  in  the  orang ;  or  is  much  larger 
in  the  male  than  in  the  female,  as  in  the  Mya  tes  caruya  and 
Fitlucia  satanas  (fig.  68).  So  it  is  with  the  whiskers  of  some 
species  of  Macacus,'^  and,  as  we  have  seen,  with  the  manes  of 
some  species  of  baboons.  But  with  most  kinds  of  monkeys  the 
various  tufts  of  hair  about  the  face  and  head  are  alike  in  both  sexes. 

The  males  of  various  members  of  the  ox  family  (Bovidse),  and 
of  certain  antelopes,  are  furnished  with  a  dewlap,  or  great  fold  ol 
skin  on  the  neck,  which  is  much  less  developed  in  the  female. 

Now,  what  must  we  conclude  with  resptct  to  such  sexual 
differences  as  these?  No  one  will  pretend  that  the  beards  of 
certain  male  goats,  or  the  dewlap  of  the  bull,  or  the  crests  ol 
hair  along  the  backs  of  certain  male  antelopes,  are  of  any  use 
to  them  in  their  ordinary  habits.  It  is  possible  that  the  im- 
mense beard  of  the  male  Pithecia,  and  the  large  beard  of  the 
male  orang,  may  protect  their  throats  when  fighting;  for 
the  keepers  in  the  Zoological   Gardens  inform  me  that  many 

1*  Judge    Caton   on    the  Wapiti,      p.  236 
'  Transact.     Ottawa       Acad.      Nat.  ^'^  See  Dr.  Gray's  '  Cat.  of  Mam- 

Sciences,'   18G8,  pp.  36,  40;  Blytii,      malia  in  British  Museum,'  part  iii. 
'  Land  and  Water,'  on   Capra  xga-      1852,  p.  144. 
grus,  1867,  p.  37.  ^'   Rengger,  '  Saugethiere,'  &c.  s. 

1^  '  Hunter's  Essays  and  Observa-  14;  Desmarest,  'Mamnialogie,'  p. 
lions,'  edited  by  Owen,  1861,  vol.  i.      lo'd. 


532 


The  Descent  of  Man. 


Part  11. 


monkeys  attack  each  other  by  the*throat ;  but  it  is  not  probable 
that  the  beard  has  been  developed  for  a  distinct  purpose  from 
that  served  by  the  whiskers,  moustache,  and  other  tufts  of  hair 
on  the  face ;  and  no  one  will  suppose  that  these  are  useful  as  a 
protection.  Must  we  attribute  all  these  appendages  of  hair  or 
skin  to  mere  purposeless  variability  in  the  male  ?  It  cannot  be 
denied  that  this  is  possible ;  for  in  many  domesticated  quad- 
rupeds, certain  characters,  apparently  not  derived  through  re- 
version from  any  wild  parent-form,  are  confined  to  the  males. 


^<^ 


^^^-^§:S 


Fig.  68. 


Pithecia  satanas,  male  (from  Brehni). 


or  are  more  developed  in  them  than  in  the  females— for  instance, 
the  hump  on  the  male  zebu-cattle  of  India,  the  tail  of  fat-tailed 
rams,  the  arched  outline  of  the  forehead  in  the  males  of  several 
breeds  of  sheep,  and,  lastly,  the  mane,  the  long  hairs  on  the 
hind-legs,  and  the  dewlap  of  the  male  of  the  Berbura  goat.^^  The 
mane,  which  occurs  only  in  the  rams  of  an  African  breed  of 


"^  See    the     chapters     on    these      '  Variation  of  Animals  tinder  Domes- 
ceveral    animals   in    vol.    i.    of  my      tication ; '  also  vol.  ii.  p.  73 ;  also 


Cha^p.  XVin.     Mammals — Ornamejital  Colours.  533 

sheep,  is  a  true  secondary  sexual  character,  for,  as  I  hear  from 
Mr.  Winwood  Reade,  it  is  not  developed  if  the  animal  be 
castrated.  Although  we  ought  to  be  extremely  cautious,  as 
shewn  in  my  work  on  'Variation  under  Domestication,'  in 
concluding  that  any  character,  even  with  animals  kept  by 
semi-civilised  people,  has  not  been  subjected  to  selection  by 
man,  and  thus  augmented,  yet  in  the  cases  just  specified  this  is 
improbable;  more  especially  as  the  characters  are  confined  to 
the  males,  or  are  more  strongly  developed  in  them  than  in  the 
females.  If  it  were  positively  known  that  the  above  African 
ram  is  a  descendant  of  the  same  iDrimitive  stock  as  the  other 
breeds  of  sheep,  and  if  the  Berbura  male-goat  with  his  mane, 
dewlap,  &c.,  is  descended  from  the  same  stock  as  other  goats, 
then,  assuming  that  selection  has  not  been  applied  to  these 
characters,  they  must  be  due  to  simple  variability,  together 
with  sexually-limited  inheritance. 

Hence  it  appears  reasonable  to  extend  this  same  view  to  all 
analogous  cases  with  animals  in  a  state  of  nature.  Nevertheless 
1  cannot  persuade  myself  that  it  generally  holds  good,  as  in  the 
case  of  the  extraordinary  development  of  hair  on  the  throat  and 
fore-legs  of  the  male  Ammotragus,  or  in  that  of  the  immense 
beard  of  the  male  Pithecia.  Such  study  as  I  have  been  able  to 
give  to  nature  makes  me  believe  that  parts  or  organs  which  are 
highly  developed,  were  acquired  at  some  period  for  a  special 
purpose.  With  those  antelopes  in  which  the  adult  male  is  more 
strongly-coloured  than  the  female,  and  with  those  monkeys  in 
which  the  hair  on  the  face  is  elegantly  arranged  and  coloured 
in  a  diversified  manner,  it  seems  probable  that  the  crests  and 
tufts  of  hair  were  gained  as  ornaments ;  and  this  I  know  is  the 
opinion  of  some  naturalists.  If  this  be  correct,  there  can  be 
little  doubt  that  they  were  gained  or  at  least  modified  through 
sexual  selection ;  but  how  far  the  same  view  may  be  extended 
to  other  mammals  is  doubtful. 

Colour  of  the  Tlalr  and  of  the  Naked  Skin. — I  will  first  give 
briefly  all  the  cases  known  to  me,  of  male  quadrupeds  differing 
in  colour  from  the  females.  With  Marsupials,  as  I  am.  informed 
by  Mr.  Gould,  the  sexes  rarely  differ  in  this  respect ;  but  the 
great  red  kangaroo  offers  a  striking  exception,  "  delicate  blue 
"  being  the  prevailing  tint  in  those  parts  of  the  female,  which 
"  in  the  male  are  red."^®    In  the  Didelphis  opossum  of  Cayenne 


chap.  XX.  on  the  practice  of  selection      logue,'  ibid.  p.  157. 

by  semi-civilised    people.     For  the  '^  Csj^Arani^/"  rw/w  ■,  Gould,  *  Mam- 

Berbura  goat,  see  Dr.  Gray,  '  Cata-      mals    of   Australia,'    1863,    vol.    ii. 


534  The  Desce7tt  of  Man.  Part  II. 

the  female  is  said  to  be  a  little  more  red  than  the  male.  Of  the 
Rodents  Dr.  Gray  remarks  :  "  African  squirrels,  especially  th()so 
"  found  in  the  tropical  regions,  have  the  fur  much  brighter 
"  and  more  vivid  at  some  seasons  of  the  year  than  at  others, 
"  and  the  fur  of  the  male  is  generally  brighter  than  that  of 
"  the  female." ^°  Dr.  Gray  informs  me  that  he  specified  the 
African  squirrels,  because,  from  their  unusually  bright  colours, 
they  best  exhibit  this  difference.  The  female  of  the  ]\lm  mmut/is 
of  Russia  is  of  a  paler  and  dirtier  tint  than  the  male.  In  a  large 
number  of  bats  the  fur  of  the  male  is  lighter  than  in  the  female.^^ 
Mr.  Dobson  also  remarks,  with  respect  to  these  animals :  "  Dif- 
"  ferences,  depending  partly  or  entirely  on  the  possession  by  the 
"  male  of  fur  of  a  much  more  brilliant  hue,  or  distinguished  by 
"  different  markings  or  by  the  greater  length  of  certain  portions, 
"  are  met  only,  to  any  appreciable  extent,  in  the  frugivorous  bats 
"  in  which  the  sense  of  sight  is  well  developed."  This  last 
remark  deserves  attention,  as  bearing  on  the  question  whether 
bright  colours  are  serviceable  to  male  animals  from  being  orna- 
mental. In  one  genus  of  sloths,  it  is  now  established,  as  Dr.  Gray 
states,  "  that  the  males  are  ornamented  differently  from  the 
"  females— that  is  to  say,  that  they  have  a  patch  of  soft  short 
"  hair  between  the  shoulders,  which  is  generally  of  a  more  or  less 
"  orange  colour,  and  in  one  species  pure  white.  The  females  on 
"  the  contrary,  are  destitute  of  this  mark." 

The  terrestrial  Carnivora  and  Insectivora  rarely  exhibit  sexual 
differences  of  any  kind,  including  colour.  The  ocelot  (Felis 
pardalis),  however,  is  exceptional,  for  the  colours  of  the  female, 
compared  with  those  of  the  male,  are  "  moins  apparentes,  le 
"  fauve  etant  plus  terne,  le  blanc  moins  pur,  les  raies  ay  ant 
"  moins  de  largeur  et  les  taches  moins  de  diametre."^^  The 
sexes  of  the  allied  Fells  mitis  also  differ,  but  in  a  less  degree ; 
the  general  hues  of  the  female  being  rather  paler  than  in  the 
male,  with  the  spots  less  black.  The  marine  Carnivora  or  seals, 
on  the  other  hand,  sometimes  differ  considerably  in  colour,  and 
they  present,  as  we  have  already  seen,  other  remarkable  sexual 
differences.  Thus  the  male  of  the  Otaria  nigrescens  of  the 
southern  hemisphere  is  of  a  rich  brown  shade  above ;  whilst  the 


Oa      the      Didelphis,       Desmarest,  United  States,'  1869,  p.  207.     Mr. 

'  Mammalogie,'  p.  256.  Dobson  on  sexual  characters  in  the 

2°  '  Annals  and  Mag.  of  Nat.  Hist.'  Chiroptera,    '  Proc.     Zoolog.     Soc' 

Nov.   1867,   p.   325.     On   the    3Ius  1873,  p.  241.     Dr.  Gray  on  Sloths, 

minutus,  Desmarest,  'Mammalogie,'  ibid.  1871,  p.  436. 

p.  304.  "])ggjj^^j.gg|.^  ;]^jj^j^j^j^]^jgjg^.  jg2()^ 

2'  J.  A.  Allen,   in    '  Bulletin    of  p.   220.     On   Felis    mitis,   Rengger, 

Mus.  Comp.  Zoolog.   of  Cambridge,  ibid.  s.  194. 


Chap.  XVIIT.     J\Ta7>nnals — Ornajfiental  Colours.  535 

female,  who  acquires  her  adult  tints  earlier  in  life  than  the  male, 
is  dark-grey  above,  the  young  of  both  sexes  being  of  a  deep 
chocolate  colour.  The  male  of  the  northern  Fhoca  groenhmdica 
is  tawny  grey,  with  a  curious  saddle-shaped  dark  mark  on  the 
back;  the  female  is  much  smaller,  and  has  a  very  different 
appearance,  being  "  dull  white  or  yellowish  straw-colour,  with  a 
"  tawny  hue  on  the  back ;"  the  young  at  first  are  pure  white, 
and- can  "hardly  be  distinguished  among  the  icy  hummocks  and 
"  snow,  their  colour  thus  acting  as  a  protection."  ^^ 

With  Ruminants  sexual  differences  of  colour  occur  more 
commonly  than  in  any  other  order.  A  difference  of  this  kind  is 
general  in  the  Strepsicerene  antelopes;  thus  the  male  nilghau 
(Port  IX  picta)  is  bluish-grey  and  much  darker  than  the  female, 
with  the  square  white  patch  on  the  throat,  the  white  marks  on 
the  fetlocks,  and  the  black  spots  on  the  ears  all  much  more 
distinct.  We  have  seen  that  in  this  species  the  crests  and 
tufts  of  hair  are  likewise  more  developed  in  the  male  than  in  the 
hornless  female.  I  am  informed  by  Mr.  Blyth  that  the  male, 
without  shedding  his  hair,  periodically  becomes  darker  during 
the  breeding-season.  Young  males  cannot  be  distinguished  from 
young  females  until  about  twelve  months  old ;  and  if  the  male  is 
emasculated  before  this  period,  he  never,  according  to  the  same 
authority,  changes  colour.  The  importance  of  this  latter  fact,  as 
evidence  that  the  colouring  of  the  Portax  is  of  sexual  origin 
becomes  obvious,  when  we  hear"-*  that  neither  the  red  summer- 
coat  nor  the  blue  winter-coat  of  the  Virginian  deer  is  at  all 
affected  by  emasculation.  With  most  or  all  of  the  highly-orna- 
mented species  of  Tragelaphus  the  males  are  darker  than  the 
hornless  females,  and  their  crests  of  hair  are  more  fully  developed. 
In  the  male  of  that  magnificent  antelope,  the  Derbyan  eland,  the 
body  is  redder,  the  whole  neck  much  blacker,  and  the  white  band 
which  separates  these  colours,  broader,  than  in  the  female.  In 
the  Cape  eland  also,  the  male  is  slightly  darker  than  the  female.^'^ 

In  the  Indian  black-buck  {A.  hezoartica),  which  belongs  to 
another  tribe  of  antelopes,  the  male  is  very  dark,  almost  black ; 
whilst  the  hornless  female  is  fawn-coloured.     We  meet  in  this 

2^  Dr.  Murie  on  the  Otaria, '  Proc.  142;    also    Dr.     Gray,     'Gleanings 

Zool.   Soc'   1869,   p.    108.     Mr.   R.  from   the    Menagerie  of  Knowsley,' 

Brown,  on  the  P.  groenlandica,  ibid.  in  which  there  is  a  splendid  drawing 

18G8,    p.    417.     See    also    on    the  of  the  Oreas  derbianus :  see  the  text 

colours  of  seals,  Desmarest,  ibid.  p.  on    Tragelaphus.     For     the      Cape- 

243,  249.  eland    {C)reas    cannci),    see    Andrew 

2*  Judge  Caton,  in  '  Trans.  Ottawa  Smith,  '  Zoology  of  S.  Africa,'  })1.  41 

Acad,  of  Nat.  Sciences,'  18G8,  p.  4.  and  42.     There   are    also   many   of 

^^  Dr.  Gray,  '  Cat.  of  Mamm.  in  these    antelopes    in    the     Zoological 

Brit.  Mas.'  part  iii.  1852,  pp.   134-  Gardens. 


536  The  Descent  of  Man.  Past  II. 

species,  as  Mr.  Blyth  informs  me,  with  an  exactly  similar  series 
of  facts,  as  in  the  Portax  pic.ta,  namely,  in  the  male  periodically 
changing  colour  during  the  breeding-season,  in  the  effects  of 
emasculation  on  this  change,  and  in  the  young  of  both  sexes 
being  indistinguishable  from  each  other.  In  the  Antilope  niger 
the  male  is  black,  the  female,  as  well  as  the  young  of  both 
sexes,  being  brown;  in  A.  smg-sing  the  male  is  much  brighter 
coloured  than  the  hornless  female,  and  his  chest  and  belly  are 
blacker;  in  the  male  A.  cnama,  the  marks  and  lines  which  occur 
on  various  parts  of  the  body  are  black,  instead  of  brown  as  in 
the  female;  in  the  brindled  gnu  {A.  gorgou)  "the  colours  of  the 
*'  male  are  nearly  the  same  as  those  of  the  female,  only  deeper 
"  and  of  a  brighter  hue."  ^'  Other  analogous  cases  could  be  added. 
The  Banteng  bull  {Bos  sondaims)  of  the  Malayan  Archipelago 
is  almost  black,  with  white  legs  and  buttocks ;  the  cow  is  of  a 
bright  dun,  as  are  the  young  males  until  about  the  age  of  three 
years,  when  they  rapidly  change  colour.  The  emasculated  bull 
reverts  to  the  colour  of  the  female.  The  female  Kemas  goat  is 
paler,  and  both  it  and  the  female  Capra  agagrus  are  said  to  be 
more  uniformly  tinted  than  their  males.  Deer  rarely  present 
any  sexual  cUfferences  in  colour.  Judge  Caton,  however, 
informs  me  that  in  the  males  of  the  wapiti  deer  (Cervns 
canadensis)  the  neck,  belly,  and  legs  are  much  darker  than  in 
the  female ;  but  during  the  winter  the  darker  tints  gradually 
fade  away  and  disappear.  I  may  here  mention  that  Judge 
Caton  has  in  his  park  thi*ee  races  of  the  Virginian  deer,  which 
diifer  slightly  in  colour,  but  the  differences  are  almost  ex- 
clusively confined  to  the  blue  winter  or  breeding  coat ;  so  that 
this  case  may  be  compared  with  those  given  in  a  previous 
chapter  of  closely-allied  or  representative  species  of  birds,  which 
differ  from  each  other  only  in  their  breeding  plumage.-'^  The 
females  of  Ceruus  pahidosus  of  S.  America,  as  well  as  the  young 
of  both  sexes,  do  not  possess  the  black  stripes  on  the  nose  and 
the  blackish-brown  line  on  the  breast,  which  are  characteristic  of 
the  adult  maleg.^^    Lastly,  as  I  am  informed  by  Mr.  Blyth,  the 

^^  On  the  Ant.  niger,  see  '  Proc.  ^'   '  Ottawa  Academy  of  Sciences,' 

Zool.  Soc'  1850,  p.  133.     With  re-  May,  21,  1868,  pp.  3,  5. 
spect  to  an  allied  species,  in  which  **  S.    ]\ruller,    on    the    Banteng, 

there  is  an  equal   sexual  ditference  '  Zoog.   Indischen  Archipel.'    1839- 

in  colour,  see    Sir    S.   Baker,  'The  1844,   tab.   35;  see  also  Raffles,  as 

Albert    Nyanza,'   1866,    vol.    ii.    p.  quoted   by  Mr.  Blyth,  in  'Laud  and 

3'i7.     For    the  A.   sing-sing,   Gray,  Water,'    1867,   p.   476.     On    goats, 

'Cat.   B.  Mus.'  p.  100.     Desmarest,  Dr.  Gray,  'Cat.  Brit.  Mus.'  p.   146  ; 

'  Mammalogie,'  p.  468,  on    the    A.  Desmarest,  '  Mammalogie,'    p.  482. 

caama.     Andrew    Smith,    '  Zoology  On  the   Ccrvus  paludosus,  Rengger, 

of  S.  Africa,'  on  the  Gnu.  ibid.  s.  345. 


Chap.  XVIII.     Mammals — Ornamental  Colours.         537 

mature  male  of  the  beautifully  coloured  and  spotted  axis  deer  is 
considerably  darker  than  the  female ;  and  this  hue  the  castrated 
male  never  acquires. 

The  last  Order  which  we  need  consider  is  that  of  the  Primates. 
The  male  of  the  Lemur  macaco  is  generally  coal-black,  whilst  the 
female  is  brown.^^  Of  the  Quadrumana  of  the  New  World,  the 
females  and  young  of  Mycetes  caraya  are  greyish-yellow  and  like 
each  other ;  in  the  second  year  the  young  male  becomes  reddish- 
brown  ;  in  the  third,  black,  excepting  the  stomach,  which,  how- 
ever, becomes  quite  black  in  the  fourth  or  fifth  year.  There  is 
also  a  strongly-marked  difference  in  colour  between  the  sexes  of 
Mycttts  senicidus  and  Cebus  capucinus  ;  the  young  of  the  former, 
and  I  believe  of  the  latter  species,  resembling  the  females. 
With  Pithecia  Itucocepliala  the  young  likewise  resemble  the 
females,  which  are  brownish-black  above  and  light  rusty-red 
beneath,  the  adult  males  being  black.  The  ruff  of  hair  round 
the  face  of  Ateles  manjinatus  is  tinted  yellow  in  the  male  and 
white  in  the  female.  Turning  to  the  Old  World,  the  males  of 
HyJohates  hoolock  are  always  black,  with  the  exception  of  a  white 
band  over  the  brows ;  the  females  vary  from  whity-brown  to  a 
dark  tint  mixed  with  black,  but  are  never  wholly  black.^''  In 
the  beautiful  Cercopithecns  diana,  the  head  of  the  adult  male  is 
of  an  intense  black,  whilst  that  of  the  female  is  dark  grey ;  in 
the  former  the  fur  between  the  thighs  is  of  an  elegant  fawn- 
colour,  in  the  latter  it  is  paler.  In  the  beautiful  and  curious 
moustache  monkey  (Cercopithecus  cephus)  the  only  difference 
between  the  sexes  is  that  the  tail  of  the  male  is  chesnut  and  that 
of  the  female  grey;  but  Mr.  Bartlett  informs  me  that  all  the 
hues  becomes  more  pronounced  in  the  male  when  adult,  whilst 
in  the  female  they  remain  as  they  were  during  youth  Ac- 
cording to  the  coloured  figures  given  by  Solomon  Miillcr,  the 
male  of  Semnopithecufi  chrysomelas  is  nearly  black,  the  female 
being  pale  brown.  In  the  Cercopithecus  cynosurns  and  griseo- 
viridis  one  part  of  the  body,  which  is  confined  to  the  male  sex,  is 
of  the  most  brilliant  blue  or  green,  and  contrasts  strikingly 
with  the  naked  skin  on  the  hinder  part  of  the  body,  which  is 
vivid  red. 

Lastly,  in  the  baboon  family,  the  adult  male  of  Cynocephalus 
hamadryas  difi"ers  from  the  female  not  only  by  his  immense 

29  Sclater,  *  Proc.  Zool.  Soc'  1866,  14  ;  and  Brehm,  '  Illustrirtes  Thier- 

p.  1.     The  same  fact  has  also  been  leben,'  B.  i.  s.   96,  107.     On  Ateles, 

fully  ascertained  by  MM.  Pollen  and  Desmarest,    '  Mamnialogic,'    p.    75. 

van  Dam.     See,  also,   Dr.   Gray   in  On    Hylobates,    Blyth,    'Land   and 

'  Annals  and  Mag.  of  Nat.  Hist.'  May  Water,'  1867,  p.  135.     On  the  Sem- 

1871,  p.  340.  nopithecus,    S.    Miiller,   'Zoog.   In- 

'*  On  Mycetes,   Rengger,  ibid,  s,  dischen  Archipel.*  tab.  x. 
24 


538 


The  Descent  of  Mmt. 


Part  II. 


mane,  but  slightly  in  the  colour  of  the  hair  and  of  the  naked 
callosities.  In  the  di'ill  (C.  leucophctus)  the  females  and  young 
are  much  paler-coloured,  with  less  green,  than  the  adult  males. 
No  other  member  in  the  whole  class  of  mammals  is  coloured  in 
so  extraordinary  a  manner  as  the  adult  male  mandrill  (C. 
mormon).  The  face  at  this  age  becomes  of  a  fine  blue,  with  the 
ridge  and  tip  of  the  nose  of  the  most  brilliant  red.  According  to 
some  authors,  the  face  is  also  marked  with  whitish  stripes,  and 


Fig.  C9.    Iload  of  male  :\randrill  (from  Gcrvais,  '  Hist.  Xat.  des  ilammiferes')- 


is  shaded  in  parts  with  black,  but  the  colours  appear  to  be 
variable.  On  the  forehead  there  is  a  crest  of  hair,  and  on  the 
chin  a  yellow  beard.  "  Toutes  Ics  parties  superieures  de  leurs 
"  cuisses  ct  le  grand  espace  nu  de  leurs  fesses  sont  cgalcment 
"  colores  du  rouge  le  plus  yif,  avcc  un  melange  de  bleu  qui  ne 


Chap.  XVIIT.     Mammals — Ornamental  Colours.         539 

"  manque  reellement  pas  d'elegancc."^^  When  the  animal  is 
excited  all  the  naked  parts  become  much  more  vividly  tinted. 
Several  authors  have  used  the  strongest  expressions  in  de- 
scribing these  resplendent  colours,  which  they  compare  with 
those  of  the  most  brilliant  birds.  Another  remarkable  pecu- 
liarity is  that  when  the  great  canine  teeth  are  fully  developed, 
immense  protuberances  of  bone  are  formed  on  each  cheek,  which 
are  deeply  furrowed  longitudinally,  and  the  naked  skin  over 
them  is  brilliantly-coloured,  as  just  described.  (Fig.  69.)  In 
the  adult  females  and  in  the  young  of  both  sexes  these  protuber- 
ances are  scarcely  perceptible ;  and  the  naked  parts  are  much 
less  bright  coloured,  the  face  being  almost  black,  tinged  with 
blue.  In  the  adult  female,  however,  the  nose  at  certain  regular 
intervals  of  time  becomes  tinted  with  red. 

In  all  the  cases  hitherto  given  the  male  is  more  strongly  or 
brighter  coloured  than  the  female,  and  differs  from  the  young  of 
both  sexes.  But  as  with  some  few  birds  it  is  the  female  which  is 
brighter  coloured  than  the  male,  so  with  the  Rhesus  monkey 
{Macaciis  rhesus)  the  female  has  a  large  surface  of  naked  skin 
round  the  tail,  of  a  brilliant  carmine  red,  which,  as  I  was 
assured  by  the  keepers  in  the  Zoological  Gardens,  periodically 
becomes  even  yet  more  vivid,  and  her  face  also  is  pale  red.  On 
the  other  hand,  in  the  adult  male  and  in  the  young  of  both  sexes, 
(as  I  saw  in  the  Gardens)  neither  the  naked  skin  at  the  posterior 
end  of  the  body,  nor  the  face,  shew  a  trace  of  red.  It  appears, 
however,  from  some  published  accounts,  that  the  male  does  occa- 
sionally, or  during  certain  seasons,  exhibit  some  traces  of  the  red. 
Although  he  is  thus  less  ornamented  than  the  female,  yet  in  the 
larger  size  of  his  body,  larger  canine  teeth,  more  developed 
whiskers,  more  prominent  superciliary  ridges,  he  follows  the 
common  rule  of  the  male  excelling  the  female. 

I  have  now  given  all  the  cases  known  to  me  of  a  difference  in 
colour  between  the  sexes  of  mammals.  Some  of  these  may  be 
the  result  of  variations  confined  to  one  sex  and  transmitted  to 
the  same  sex,  without  any  good  being  gained,  and  therefore 
without  the  aid  of  selection.  We  have  instances  of  this  with 
our  domesticated  animals,  as  in  the  males  of  certain  cats  being 
rusty-red,  whilst  the  females  are  tortoise-shell  coloured.  Analo- 
gous cases  occur  in  nature :  Mr.  Bartlett  has  seen  many  black 
varieties  of  the  jaguar,  leopard,  vulpine  phalanger,  and  wombat ; 

^'  Gervais,  '  Hist,  Nat.  des  Mam-  Desmarest,  *  IMainmalogie,'  p.  70. 
miferes,'  1854,  p.  103.  Figures  are  Geoflroy  St.-Hiiaire  aud  F.  Cuvier, 
given  of  the  skull  of  the  male.    Also      *  Hist.  Nat.  des  Mamm.'  1824,  torn.  i. 


540  TJie  Descent  of  Man.  Part  II. 

and  he  is  certain  that  all,  or  nearly  all  these  animals,  were  males. 
On  the  other  hand,  with  wolves,  foxes,  and  apparently  American 
squirrels,  both  sexes  are  occasionally  born  black.  Hence  it  is 
quite  possible  that  with  some  mamma's  a  difference  in  colour 
between  the  sexes,  especially  when  this  is  congenital,  may  simply 
be  the  result,  without  the  aid  of  selection,  of  the  occurrence  of 
one  or  more  variations,  which  from  the  first  were  sexually  limited 
in  their  transmission.  Nevertheless  it  is  improbable  that  the 
diversified,  vivid,  and  contrasted  colours  of  certain  quadrupeds, 
for  instance,  of  the  above  monkeys  and  antelopes  can  thus  be 
accounted  for.  We  should  bear  in  mind  that  these  colours  do 
not  appear  in  the  male  at  birth,  but  only  at  or  near  maturity ; 
and  that  unlike  ordinary  variations,  they  are  lost  if  the  male  be 
emasculated.  It  is  on  the  whole  probable  that  the  strongly- 
marked  colours  and  other  ornamental  characters  of  male 
quadrupeds  are  beneficial  to  them  in  their  rivalry  with  other 
males,  and  have  consequently  been  acquired  through  sexual 
selection.  This  view  is  strengthened  by  the  differences  in 
colour  between  the  sexes  occurring  almost  exclusively,  as  may 
be  collected  from  the  previous  details,  in  those  groups  and  sub- 
groups of  mammals,  which  present  other  and  strongly-marked 
secondary  sexual  characters ;  these  being  likewise  due  to  sexual 
selection. 

Quadrupeds  manifestly  take  notice  of  colour.  Sir  S.  Baker- 
repeatedly  observed  that  the  African  elephant  and  rhinoceros 
attacked  white  or  grey  horses  with  special  fury.  I  have  else- 
where shewn  '^'^  that  half- wild  horses  apparently  prefer  to  pair 
with  those  of  the  same  colour,  and  that  herds  of  fallow-deer  of 
different  colours,  though  living  together,  have  long  kept  distinct. 
It  is  a  more  significant  fact  that  a  female  zebra  would  not 
admit  the  addresses  of  a  male  ass  until  he  was  painted  so  as  to 
resemble  a  zebra,  and  then,  as  John  Hunter  remarks,  "  she 
"  received  him  very  readily.  In  this  curious  fact,  we  have 
"  instinct  excited  by  mere  colour,  which  had  so  strong  an  eifect 
"  as  to  get  the  better  of  everything  else.  But  the  male  did  not 
"  require  this,  the  female  being  an  animal  somewhat  similar  to 
"  himself,  was  sufficient  to  rouse  him."^^ 

In  an  earlier  chai)ter  we  have  seen  that  the  mental  powers  of 
the  higher  animals  do  not  differ  in  kind,  though  greatly  in 
degree,  from  the  corresponding  powers  of  man,  especially  of  the 
lower  and  barbarous  races ;  and  it  would  appear  that  even  their 
taste  for  the  beautiful  is  not  widely  different  from  that  of  the 

"  'The  Variation  of  Animals  and  '*  'Essays  and  Observations  by 

Plants  under  Domesticatiou,'  1868,      J.   Hunter,*  edited    by  Owen,  1801, 
vol.  ii.  pp.  102,  103.  vol.  i.  p.  194. 


Chap,  XVII  r.         Mammals  —  Ornaine?its.  541 

QuadriiiUcana.  As  the  negro  of  Africa  raises  the  flesh  on  his 
ftice  into  parallel  ridges  *'  or  cicatrices,  high  above  the  natural 
"  surface,  which  unsightly  deformities,  are  considered  great 
"personal  attractions;"^^ — as  negroes  and  savages  in  many 
parts  of  the  world  paint  their  faces  with  red,  blue,  white,  or 
black  bars, — so  the  male  mandrill  of  Africa  appears  to  have 
acquired  his  deeply-furrowed  and  gaudily-coloured  face  from 
having  been  thus  rendered  attractive  to  the  female.  No  doubt 
it  is  to  us  a  most  grotesque  notion  that  the  posterior  end  of  the 
body  should  be  coloured  for  the  sake  of  ornament  even  more 
brilliantly  than  the  face ;  but  this  is  not  more  strange  than  that 
the  tails  of  many  birds  should  be  especially  decorated. 

With  mammals  we  do  not  at  present  possess  any  evidence 
that  the  males  take,  pains  to  display  their  charms  before  the 
female  ;  and  the  elaborate  manner  in  wiiicli  this  is  performed  by 
male  birds  and  other  animals,  is  the  strongest  argument  in 
favour  of  the  belief  that  the  females  admire,  or  are  excited  by, 
the  ornaments  and  colours  displayed  before  them.  There  is, 
however,  a  striking  parallelism  between  mammals  and  birds  in 
all  their  secondary  sexual  characters,  namely  in  their  weapons 
for  fighting  with  rival  males,  in  their  ornamental  appendages, 
and  in  their  colours.  In  both  classes,  when  the  male  differs 
from  the  female,  the  young  of  both  sexes  almost  always  resemble 
each  other,  and  in  a  large  majority  of  cases  resemble  the  adult 
female.  In  both  classes  the  male  assumes  the  characters  proper 
to  his  sex  shortly  before  the  age  of  reproduction ;  and  if  emas- 
culated at  an  early  period,  loses  them.  In  both  classes  the 
change  of  colour  is  sometimes  seasonal,  and  the  tints  of  the 
naked  parts  sometimes  become  more  vivid  during  the  act  of 
courtship.  In  both  classes  the  male  is  almost  always  more 
vividly  or  strongly  coloured  than  the  female,  and  is  ornamented 
with  larger  crests  of  hair  or  feathers,  or  other  such  appendages. 
In  a  few  exceptional  cases  the  female  in  both  classes  is  more 
highly  ornamented  than  the  male.  With  many  mammals,  and 
at  least  in  the  case  of  one  bird,  the  male  is  more  odoriferous 
than  the  female.  In  both  classes  the  voice  of  the  male  is  more 
powerful  than  that  of  the  female.  Considering  this  jDarallelism 
there  can  be  little  doubt  that  the  same  cause,  whatever  it  may 
}.>e,  has  acted  on  mammals  and  birds ;  and  the  result,  as  far  as 
ornamental  characters  are  concerned,  may  be  attributed,  as  it 
appears  to  me,  to  the  long-continued  preference  of  the  indi- 
viduals of  one  sex  for  certain  individuals  of  the  opposite  sex, 
combined  with  their  success  in  leaving  a  larger  immber  of 
offspring  to  inherit  their  superior  attractions. 

3'  Sir  S.  Baker,  'The  Nile  Tributaries  of  Abyssinia,'  1867. 


542  The  Descent  of  Man.  Part  II. 

Equal  transmission  of  ornamental  characters  to  both  sexes. — With 
many  birds,  ornaments,  which  analogy  leads  us  to  believe  were 
primarily  acquired  by  the  males,  have  been  transmitted  equally, 
or  almost  equally,  to  both  sexes ;  and  we  may  now  enquire  how 
far  this  view  ajDplies  to  mammals.  With  a  considerable  number 
of  species,  especially  of  the  smaller  kinds,  both  sexes  have  been 
coloured,  independently  of  sexual  selection,  for  the  sake  of  pro- 
tection ;  but  not,  as  far  as  I  can  judge,  in  so  many  cases,  nor  in 
so  striking  a  manner,  as  in  most  of  the  lower  classes.  Audubon 
remarks  that  he  often  mistook  the  musk-rat,^^  whilst  sitting  on 
the  banks  of  a  muddy  stream,  for  a  clod  of  earth,  so  complete 
was  the  resemblance.  .  The  hare  on  her  form  is  a  familiar  instance 
of  concealment  through  colour;  yet  this  principle  partly  fails 
in  a  closely-allied  sjDecies,  the  rabbit,  for  when  running  to  its 
burrow,  it  is  made  conspicuous  to  the  sportsman,  and  no  doubt 
to  all  beasts  of  prey,  by  its  upturned  white  tail.  No  one  doubts 
that  the  quadrupeds  inhabiting  sncw-clad  regions  have  been 
rendered  white  to  protect  them  from  their  enemies,  or  to  favour 
their  stealing  on  their  prey.  In  regions  where  snow  never 
lies  for  long,  a  white  coat  would  be  injurious;  consequently, 
species  of  this  colour  are  extremely  rare  in  the  hotter  parts  of 
the  world.  It  deserves  notice  that  many  quadrupeds  inhabiting 
moderately  cold  regions,  although  they  do  not  assume  a  white 
winter  dress,  become  paler  during  this  season ;  and  this  appa- 
rently is  the  direct  result  of  the  conditions  to  which  they  have 
long  been  exposed.  Pallas  ^^  states  that  in  Siberia  a  change  of 
tliis  nature  occurs  with  the  wolf,  two  species  of  Mustela,  the 
domestic  horse,  the  Equus  liemionus,  the  domestic  cow,  two 
species  of  antelopes,  the  musk-deer,  the  roe,  elk,  and  rein- 
deer. The  roe,  for  instance,  has  a  red  summer  and  a  greyish- 
white  winter  coat;  and  the  latter  may  perhaps  serve  as  a 
protection  to  the  animal  whilst  wandering  through  the  leafless 
thickets,  sprinkled  -^ith  snow  and  hoar-frost.  If  the  above- 
named  animals  were  gradually  to  extend  their  range  into  regions 
perpetually  covered  with  snow,  their  pale  winter-coats  would 
})r()bably  be  rendered  through  natural  selection,  whiter  and 
whiter,  until  they  became  as  white  as  snow. 

Mr.  Eceks  has  given  me  a  curious  instance  of  an  animal  profit- 
ing  by  being  peculiarly  coloured.  He  raised  from  fifty  to  sixty 
white  and  brown  piebald  rabbits  in  a  large  walled  orchard; 
and  he  had  at  the  same  time  some  similarly  coloured  cats  in  his 

^*   Fil>er  zibcthicus,  Aiulubon   and  Glirium  ordine,'   1778,  p.  7.     What 

Bachin.in,  'The    Quatlrupoils  of    N.  1  have  called  the  roe  is  the  Caprco/us 

America,*  1846,  p.  109.  sibiricus  subccaudatus  of  Pallas. 

'"  'Novae  species  Quadrupedum  e 


Chap.  XVI II.     JMamnials — Spots  and  Stripes.  543 

house.  Such  cats,  as  I  have  often  noticed,  are  very  conspicnons 
during  daj^ ;  but  as  tliey  used  to  lie  in  watch  during  tlie  dusk 
at  the  mouths  of  the  burrows,  the  rabbits  apparently  did  not 
distinguish  them  from  their  parti-coloured  brethren.  The  result 
was  that,  within  eighteen  months,  every  one  of  these  parti- 
colourted  rabbits  was  destroyed ;  and  there  was  evidence  that 
this  was  effected  by  the  cats.  Colour  seems  to  be 'advantageous 
to  another  animal,  the  skunk,  in  a  manner  of  which  we  have  had 
many  instances  in  other  classes.  No  animal  will  voluntarily 
attack  one  of  these  creatures  on  account  of  the  dreadful  odour 
which  it  emits  when  irritated ;  but  during  the  dusk  it  would  not 
easily  be  recognised  and  might  be  attacked  by  a  beast  of  prey. 
Hence  it  is,  as  Mr.  Belt  believes,^^  that  the  skunk  is  provided  with 
a  great  white  bushy  tail,  which  serves  as  a  conspicuous  warning. 
Although  we  must  admit  that  many  quadrupeds  have  received 
their  present  tints  either  as  a  protection,  or  as  an  aid  in  j^rocuring 
prey,  yet  with  a  host  of  species,  the  colours  are  far  too  con- 
spicuous and  too  singularly  arranged  to  allow  us  to  suppose  that 
they  serve  for  these  purposes.  We  may  take  as  an  illustration 
certain  antelopes;  when  we  see  the  square  white  patch  on  the 
throat,  the  white  marks  on  the  fetlocks,  and  the  round  black 
spots  on  the  ears,  all  more  distinct  in  the  male  of  tlie  VorUvx  i^k'n 
than  in  the  female  ; — when  we  see  that  the  colours  arc  more 
vivid,  that  the  narrow  white  lines  on  the  flank  and  the  broad 
white  bar  on  the  shoulder  are  more  distinct  in  the  male  Ovms 
derhyanus  than  in  the  female  ; — when  we  see  a  similar  difference 
between  the  sexes  of  the  curiously-ornamented  Tragelaphus 
scriptns  (fig.  70), — we  cannot  believe  that  differences  of  this  kind 
are  of  any  service  to  either  sex  in  their  daily  habits  of  life.  It 
seems  a  much  more  probable  conclusion  that  the  various  marks 
were  first  acquired  by  the  males  and  their  colours  intensified 
through  sexual  selection,  and  then  partially  transferred  to  the 
females.  If  this  view  be  admitted,  there  can  be  little  doubt  that 
the  equally  singular  colours  and  marks  of  many  other  antelopes, 
though  common  to  both  sexes,  have  been  gained  and  transmitted 
in  a  like  manner.  Both  sexes,  for  instance,  of  the  koodoo 
{Stnpdaros  kmhi)  (fig.  64)  have  narrow  white  vertical  lines  on 
their  hind  flanks,  and  an  elegant  angular  white  mark  on  their 
foreheads.  Both  sexes  in  the  genus  Damalis  are  very  oddly 
coloured ;  in  1).  pyganja  the  back  and  neck  are  purplish-reel, 
shading  on  the  flanks  into  black  ;  and  these  colours  are  abruptly 
separated  from  the  white  belly  and  from  a  large  white  space  on 
the  buttocks;  the  head  is  still  more  oddly  coloured,  a  large 
oblong  white  mask,  narrowly-edged  with  black,  covers  the  face 
3^  'The  Naturalist  in  Nicaragua,'  p.  249. 


544 


The  Descent  of  Man. 


Part  II. 


up  to  the  eyes  (fig*  71);  there  are  three  white  stripes  on  the 
forehead,  and  the  ears  are  marked  with  white.  The  fawns  of 
this  species  are  of  a  uniform  pale  yellowish-brown.  In  DamaJis 
albifrons  the  colouring  of  the  head  dilfers  from  that  in  the  last 
species  in  a  single  white  stripe  replacing  the  three  stripes,  and 


lig.  70.        Tragelaphus  scrijitus,  male  (from  the  Knowsley  Menagerie). 

in  the  ears  being  almost  wholly  white.^^  After  having  studied 
to  the  best  of  my  ability  the  sexual  differences  of  animals  be- 
longing to  all  classes,  I  cannot  avoid  the  conclusion  that  the 
curiously-arranged  colours  of  many  antelopes,  though  common  to 
both  sexes,  are  the  result  of  sexual  selection  primarily  a})plicd  to 
the  male. 

"  See  the  fine  plates  in  A.  Smith's      *  Gleanings  from  the 
'  Zoology  of  S.  Africa,'  and  Dr.  (iray's      Kuowsley.' 


Monajrerie  of 


Chap.  XVllI.     Mammals—Spots  and  Stripes,  545 

The  same  conclusion  may  perhaps  be  extended  to  the  tiger,  one 
of  the  most  beautiful  animals  in  the  world,  the  sexes  of  which 
cannot  be  disthiguislied  by  colour,  even  by  the  dealers  m  wild 
beasts  Mr  Wallace  believes^'-'  that  the  striped  coat  ot  the  tiger 
-  so  assimilates  with  the  vertical  stems  of  the  bamboo,  as  to 
"  assist  greatly  in  concealing  him  from  his  approaching  prey. 
But  this  view  does  not  appear  to  me  satisfactory.  We  have 
some  slight  evidence  that  his  beauty  may  be  due   to  sexual 


Fig.  71.  Damalis  pygarga,  male  (from  the  Knowsley  Menagerie). 

selection,  for  in  two  species  of  Felis  the  analosrous  marks  and 
colours  are  rather  brighter  in  the  male  than  in  the  female.  The 
zebra  is  conspicuously  striped,  and  stripes  cannot  afford  any 
protection  on  the  open  plains  of  South  Africa.  BurchelP"  in  de- 
scribing a  herd  says,  "  their  sleek  ribs  glistened  in  the  sun,  and 
"  the  brightness  and  regularity  of  their  striped  coats  presented  a 
"  picture  of  extraordinary  beauty,  in  which  probably  they  are 

39  <■  Westminster  Review,'  July  1,  "^  'Travels  iu  South  Africa,'  1824, 

18G7,  p.  5.  vol.  ii.  p.  315. 


546  The  Descent  of  Man.  Part  ir. 

"  not  sui'i^assed  by  any  other  quadruped."  But  as  throughout 
the  wliole  group  of  the  Equidie  the  sexes  are  identical  in  colour, 
we  have  here  no  evidence  of  sexual  selection.  Nevertheless  he 
who  attributes  the  white  and  dark  vertical  stripes  on  tlie  flanks 
of  various  antelopes  to  this  process,  will  probably  extend  the 
same  view  to  the  Eoyal  Tiger  and  beautiful  Zebra. 

We  have  seen  in  a  former  chapter  that  when  young  animals 
belonging  to  any  class  follow  nearly  the  same  habits  of  life  as 
their  parents,  and  yet  are  coloured  in  a  different  manner,  it  may 
be  inferred  that  they  have  retained  the  colouring  of  some 
ancient  and  extinct  progenitor.  In  the  family  of  pigs,  and  in 
the  tapirs,  the  young  are  marked  with  longitudinal  stripes, 
and  thus  differ  from  all  the  existing  adult  species  in  these  two 
groups.  With  many  kinds  of  deer  the  young  are  marked  with 
elegant  white  spots,  of  which  their  parents  exhibit  not  a  trace. 
A  graduated  series  can  be  followed  from  the  axis  deer,  both 
sexes  of  wliich  at  all  ages  and  daring  all  seasons  are  beautifully 
spotted  (the  male  being  rather  more  strongly  coloured  than  the 
female),  to  species  in  which  neither  the  old  nor  the  young  arc 
spotted.  I  will  specify  some  of  the  steps  in  this  series.  The 
Mantchurian  deer  (Ceruus  manfchnricus)  is  spotted  during  the 
whole  year,  but,  as  I  have  seen  in  the  Zoological  Gardens,  the 
spots  are  much  plainer  during  the  summer,  when  the  general 
colour  of  the  coat  is  lighter,  than  during  the  winter,  when  the 
general  colour  is  darker  and  the  horns  are  fully  developed.  In 
the  hog-deer  {Uy<laphus  porcinas)  i\\Q  six>ts  are  extremely  con- 
spicuous during  the  summer  when  the  coat  is  reddish-brown, 
l>ut  quite  disappear  during  the  winter  when  the  coat  is  brown.^' 
In  both  these  species  the  young  are  spotted.  In  the  Virginian 
deer  the  young  are  likewise  spotted,  and  about  five  ]Der  cent,  of 
the  adult  animals  living  in  Judge  Caton's  park,  as  I  am  in- 
formed by  him,  temporarily  exhibit  at  the  period  when  the  red 
summer  coat  is  being  replaced  by  the  bluish  winter  coat,  a  row 
of  spots  on  each  flank,  which  are  always  the  same  in  number, 
though  very  variable  in  distinctness.  From  this  condition  there 
is  but  a  very  small  step  to  the  complete  absence  of  spots  in  the 
adults  at  all  seasons  ;  and,  lastly,  to  their  absence  at  all  ages  and 
seasons,  as  occurs  with  certain  species.  Fi-om  the  existence  of 
this  perfect  series,  and  more  especially  from  the  fawns  of  so  many 
species  being  s]X)tted,  we  may  conclude  that  the  now  living 
members  of  the  deer  family  are  the  descendants  of  some  ancient 

^'  Dr.  Gray,  '  Gleanings  from  the  deer    of    Ceylon,    says    it    is  more 

Mcn:i£;erie     of     Knowsley,'     p.     (54.  brightly   spotted    with   white    than 

Mr.  lilvtli,  ill  s])eakiug  ('Land  and  the  common  hog-deer,  at  the  seosou 

Watt-r,'    l«ijlt,   p.    4'JJ   of  the    hog-  wii-ju  it  renews  its  hums. 


Chap.  XVIII.     Mammals — Spots  and  Stripes.  547 

species  which,  like  tlic  axis  deer,  was  spotted  at  all  ages  and 
seasons.  A  still. more  ancient  progenitor  probably  somewhat 
resembled  the  Hyomoschus  aquaticus—fov  this  animal  is  spotted, 
and  the  hornless  males  have  large  exserted  canine  teeth,  of 
which  some  few  true  deer  still  retain  rudiments.  Hyomoschus, 
also,  offers  one  of  those  interesting  cases  of  a  form  linking  to- 
gether two  groups,  for  it  is  intermediate  in  certain  osteological 
characters  between  the  pachyderms  and  ruminants,  which  were 
formerly  thought  to  be  quite  distinct."*^ 

A  curious  difficulty  here  arises.  If  we  admit  that  coloured  spots 
and  stripes  were  first  acquired  as  ornaments,  how  comes  it  that 
so  many  existing  deer,  the  descendants  of  an  aboriginally  spotted 
animal,  and  all  the  species  of  pigs  and  tapirs,  the  descendants  of 
an  aboriginally  striped  animal,  have  lost  in  their  adult  state  their 
former  ornaments  ?  I  cannot  satisfactorily  answer  this  question. 
AVe  may  feel  almost  sure  that  the  spots  and  stripes  disappeared 
at  or  near  maturity  in  the  progenitors  of  our  existing  species,  so 
that  they  w^ere  still  retained  by  the  young ;  and,  owing  to  the  law 
of  inheritance  at  corresponding  ages,  were  transmitted  to  the 
young  of  all  succeeding  generations.  It  may  have  been  a  great 
advantage  to  the  lion  and  puma,  from  the  open  nature  of  their 
usual  haunts,  to  have  lost  their  stripes,  and  to  have  been  thus 
rendered  less  conspicuous  to  their  prey ;  and  if  the  successive 
variations,  by  which  this  end  was  gained,  occurred  rather  late  in 
life,  the  young  would  have  retained  their  striiDcs,  as  is  now  the 
case.  As  to  deer,  pigs,  and  tapirs,  Fritz  Miiller  has  suggested 
to  me  that  these  animals,  by  the  removal  of  their  spots  or  stripes 
through  natural  selection,  would  have  been  less  easily  seen  by 
their  enemies  ;  and  that  they  would  have  especially  required  this 
protection,  as  soon  as  the  carnivora  increased  in  size  and  number 
during  the  tertiary  periods.  This  may  be  the  true  explanation, 
but  it  is  rather  strange  that  the  young  should  not  have  been  thus 
protected,  and  still  more  so  that  the  adults  of  some  species  should 
have  retained  their  spots,  either  partially  or  completely,  during 
part  of  the  year.  We  know  that,  when  the  domestic  ass  varies 
and  becomes  reddish-brown,  grey,  or  black,  the  stripes  on  the 
shoulders  and  even  on  the  spine  frequently  disappear,  though  we 
cannot  explain  the  cause.  Very  few  horses,  except  dun-coloured 
kinds,  have  stripes  on  any  part  of  their  bodies,  yet  we  have  good 
reason  to  believe  that  the  aboriginal  horse  was  striped  on  the 
legs  and  spine*  and  probably  on  the  shoulders.'*^  Hence  the  dis- 
appearance of  the  spots  and  stripes  in  our  adult  existing  deer, 

*2  Falconer  and   Cautley,   *  Proc.  *'  'The  Variation  of  Animals  and 

Geoloe.  Soc'  1843;  and  Falconer's  Plants  under  Domestication,'  1868, 
*  Pal.  Jlemoirs,'  vol.  i.  p.  190.  vol.  i.  ]i.  Gl-64. 


548 


The  Descent  of  Man. 


Part  II. 


pigs,  and  tapirs,  may  bo  due  to  a  change  in  the  general  colour  of 
their  coats ;  but  whether  this  change  was  eiiected  through 
sexual  or  natural  selection,  or  was  due  to  the  direct  action  of  the 
conditions  of  life,  or  to  some  other  unknown  cause,  it  is  impossible 
to  decide.  An  observation  made  by  Mr.  Sclater  well  illustrates 
our  ignorance  of  the  laws  which  regulate  the  appearance  and 
disappearance  of  stripes;  the  species  of  Asinus  which  inhabit 
the  Asiatic  continent  are  destitute  of  stripes,  not  having  even 


Fifr.  72.  Tloatl  of  Senmopithecus  rubicundus.  This  and  the  following  figures  (from 
Prof.  (Jprvais)  are  given  to  shew  the  odd  arrangement  and  development  of  the  hair 
on  the  head. 


the  cross  shoulder-stripe,  whilst  those  which  inhabit  Africa  are 
conspicuously  striped,  with  the  ]iartial  exception  of  A.  ta."tuopus, 
■which  has  only  the  cross  shoulder-stripe  and  generally  some 
faint  bars  on  the  legs  ;  and  this  species  inhabits  the  almost  inter- 
noediate  region  of  Upper  l^^gypt  and  Abyssinia."** 

"  '  Proc.  Zool.  Soc'  1862,  ]).  \M.      L;indw.'     Bd.  xliii.  a.  222. 
Ser,  also,  Dr.  Iliutmanu,  '  Auu.  d. 


Chap.  XVI II.     Beauty  of  the  Qtcadriunana. 


549 


Quadrumana. — Before  we  conclude,  it  will  be  well  to  add  a 
few  remarks  on  the  ornaments  of  monkeys.  In  most  of  the 
species  the  sexes  resemble  each  other  in  colour,  but  in  some,  as  we 
have  seen,  the  males  differ  from  the  females,  especially  in  the 
colour  of  the  naked  parts  of  the  skin,  in  the  development  of  the 
beard,  whiskers,  and  mane.  Many  species  are  coloured  either  in 
so  extraordinary  or  so  beautiful  a  manner,  and  are  furnished 
with  such  curious  and  elegant  crests  of  hair,  that  we  can  hardly 
avoid  looking  at  these  characters  as  having  been  gained  for  the 
sake  of  ornament.     The  accompanying  figures  (tigs.  72  to  70) 


Fig  73     Head  of  Semnopitbecus  comatus  Fig.  74.    Head  of  Cebus  oapucinus. 


Fig.  75.    Head  of  Ateles  marginatus. 


Fig.  76.    Head  of  Cebus  vcllerosus. 


serve  to  shew  the  arrangement  of  the  hair  on  the  face  and  head 
in  several  species.  It  is  scarcely  conceivable  that  these  crests  of 
hair,  and  the  strongly  contrasted  colours  of  the  fur  and  skin,  can 
be  the  result  of  mere  variability  without  the  aid  of  selection ; 


550  The  Descent  of  M aft.  ] 'art  II. 

and  it  is  inconceivable  that  they  can  be  of  use  in  any  ordinary  way 
to  these  animals.  If  so,  they  have  probably  been  gained  through 
sexual  selection,  though  transmitted  equally,  or  almost  equally, 
to  both  sexes.  With  many  of  the  Quadrumana,  we  have  addi- 
tional evidence  of  the  action  of  sexual  selection  in  the  greater 
size  and  strength  of  the  males,  and  in  the  greater  develoiDment  of 
their  canine  teeth,  in  comparison  with  the  females, 

A  few  instances  will  suffice  of  the  strange  manner  in  which 
both  sexes  of  some  species  are  coloured,  and  of  the  beauty  of  others. 
The  face  of  the  Cercopithecus  pttaurhta  (fig.  77)  is  black,  the 
whiskers  and  beard  being  white,  with  a  defined,  round,  white 
spot  on  the  nose,  covered  with  short  white  hair,  which  gives  to 
the  animal  an  almost  ludicrous  aspect.  The  Semnopithecus 
frontatus  likewise  has  a  blackish  face  with  a  long  black  beard, 
and  a  large  naked  spot  on  the  forehead  of  a  bluish-white  colour. 
The  face  of  Macacus  lasiotus  is  dirty  flesh-coloured,  with  a 
defined  red  spot  on  each  cheek.  The  appearance  of  Ctrcocehus 
a'thiopjs  is  grotesque,  with  its  black  face,  white  whiskers  and 
collar,  chesnut  head,  and  a  large  naked  white  spot  over  each 
eyelid.  In  very  many  species,  the  beard,  whiskers,  and  crests  of 
hair  round  the  face  are  of  a  different  colour  from  the  rest  of  the 
head,  and  when  different,  are  always  of  a  lighter  tint,'*"  being 
often  pure  white,  sometimes  bright  yellow,  or  reddish.  The 
whole  face  of  the  South  American  Brachyurus  calvus  is  of  a 
"  glowing  scarlet  hue ; "  but  this  colour  does  not  appear  until 
the  animal  is  nearly  mature.^^  The  naked  skin  of  the  face  differs 
wonderfully  in  colour  in  the  various  species.  It  is  often  brown 
or  flesh-colour,  with  parts  perfectly  white,  and  often  as  black  as 
that  of  the  most  sooty  negro.  In  the  Brachyurus  the  scarlet  tint 
is  brighter  than  that  of  the  most  blushing  Caucasian  damsel.  It 
is  sometimes  more  distinctly  orange  than  in  any  Mongolian,  and 
in  several  species  it  is  blue,  passing  into  violet  or  grey.  In  all 
the  species  known  to  Mr.  Bartlett,  in  which  the  adults  of  both 
sexes  have  strongly-coloured  faces,  the  colours  are  dull  or  absent 
during  early  youth.  This  likewise  holds  good  with  the  mandrill 
atid  Khcsus,  in  which  the  face  and  the  posterior  parts  of  the  body 
are  brilliantly  coloured  in  one  sex  alone.  In  these  latter  cases 
we  have  reason  to  believe  that  the  colours  wore  acquired  through 
sexual  selection ;  and  we  are  naturally  led  to  extend  the  same 
view  to  the  foregoing  species,  though  both  sexes  when  adult 
have  their  faces  coloured  in  the  same  manner. 

*'  I    observed   this   fact    in    the  *  Hist.  Nat.  des  Mammif&res,'  torn. 

Zoological  Gardens;  and  many  cases  i.  1824. 

may  be  seen  in  the  coloured  plates  in  ■*"  Bates, 'The  Naturalist  on  the 

Geoin-oy  St.-llilaire  and  F.  Cuvier,  Amazons,'  18Gi),  vol.  ii.  p.  310. 


Chap.  XYIU.     Beauty  of  the  Qtiadriimaiia. 


551 


Although  many  kinds  of  monkeys  are  far  from  beautiful  accord- 
ing to  om-  taste,  other  species  are  universally  admired  for  tbeir 
elegant  appearance  and  bright  colours.  The  kemnopithecus 
ntmceusj  though  peculiarly  coloured,  is  described  as  extremely 


Fig.  11. 


Cercopithecus  petaurista  (from  Brehm). 


552  The  Descent  of  Man.  Part  II. 

pretty ;  tlie  orange-tinted  face  is  surrounded  by  long  whiskers 
of  glossy  whiteness,  with  a  line  of  chesnut-red  over  the  eye- 
brows ;  the  fur  on  the  back  is  of  a  delicate  grey,  with  a  square 
patch  on  the  loins,  the  tail  and  the  fore-arms  being  of  a  pure 
white ;  a  gorget  of  chesnut  surmounts  the  chest ;  the  thighs  are 
black,  with  the  legs  chesnut-red.  I  will  mention  only  two  other 
monkeys  for  their  beauty;  and  I  have  selected  these  as  pre- 
senting slight  sexual  differences  in  colour,  which  renders  it 
in  some  degree  probable  that  both  sexes  owe  their  elegant 
appearance  to  sexual  selection.  In  the  moustache-monkey 
{Cercopithecus  cephus)  the  general  colour  of  the  fur  is  mottled- 
greenish  with  the  throat  white  ;  in  the  male  the  end  of  the  tail 
is  chesnut,  but  the  face  is  the  most  ornamented  part,  the  skin 
being  chiefly  bluish-grey,  shading  into  a  blackish  tint  beneath 
the  eyes,  with  the  upper  lip  of  a  delicate  blue,  clothed  on  the 
lower  edge  with  a  thin  black  moustache;  the  whiskers  are 
o]-ange-colourcd,  with  the  upper  part  black,  forming  a  band 
which  extends  backwards  to  the  ears,  the  latter  being  clothed 
with  whitish  hairs.  In  the  Zoological  Society's  Gardens  I  have 
often  overheard  visitors  admiring  the  beauty  of  another  monkey, 
deservedly  called  CercopWiecus  diana  (fig.  78) ;  the  general 
colour  of  the  fur  is  grey ;  the  chest  and  inner  surface  of  the 
forelegs  are  white;  a  large  triangular  defined  space  on  the 
hinder  part  of  the  back  is  rich  chesnut ;  in  the  male  the  inner 
sides  of  the-  thighs  and  the  abdomen  are  delicate  fawn-coloured, 
and  the  top  of  the  head  is  black ;  the  face  and  ears  are  intensely 
black,  contrasting  finely  with  a  white  transverse  crest  over  the 
eye-brows  and  a  long  white  peaked  beard,  of  which  the  basal 
portion  is  black.^^ 

In  these  and  many  other  monkeys,  the  beauty  and  singular 
arrangement  of  their  colours,  and  still  more  the  diversified  and 
elegant  arrangement  of  the  crests  and  tufts  of  hair  on  their 
heads,  force  the  conviction  on  my  mind  that  these  characters 
have  been  acquired  through  sexual  selection  exclusively  as 
ornaments.  • 

Summary. — The  law  of  battle  for  the  possession  of  the  female 
appears  to  prevail  throughout  the  whole  great  class  of  mammals. 
Most  naturalists  will  admit  that  the  greater  size,  strength, 
courage,  and  pugnacity  of  the  male,  his  special  weapons  of 
offence,  as  well  as  his  special  means  of  defence,  have  been 

*''  I  have  seen  mo.st  of  the  above  ^Ir.  W.  C.  Martin's  'Nat.  Hist,  of 

monkeys  in  the  Zoological  Society's  ]\lamnialia,'   1841,  p.  4(j0  ;  see  also 

(Jardeus.     The    description    otl   the  pp.  475,  523. 
Hc'miwjjit/ic'cuf  7Wi)VLUs  is  taken  from 


Chap.  XVIII.         Mammals — Summary, 


553 


acquired  or  modified  through  that  form  of  selection  wliich  I 
have  called  sexual.  This  does  not  depend  on  any  superiority  in 
the  general  struggle  for  life,  but  on  certain  individuals  of  one 
sex,  generally  the  male,  being  successful  in  conquering  other 
males,  and  leaving  a  larger  number  of  offspring  to  inherit  their 
superiority  than  do  the  less  successful  males. 


Fig.  78. 


Cercopithecus  diana  (from  Brehm). 


There  is  another  and  more  peaceful  In'nd  of  contest,  m  which 
the  males  endeavour  to  excite  or  allure  the  females  by  various 
charms.  This  is  probably  carried  on  in  some  cases  by  the  i)ower- 
ful  odours  emitted  by  the  males  during  the  breeding-season ;  the 
odoriferous  glaivls  having  been  acquired  through  sexual 
selection.    Whether  the  same  view  can  be  extended  to  the  voice 


5  54  The  Descent  of  Man.  Part  IT, 


is  doubtful,  for  the  vocal  organs  of  the  males  must  have  been 
strengthened  by  use  during  maturity,  under  the  powerful  excite- 
ments of  love,  jealousy  or  rage,  and  will  consequently  have  been 
transmitted  to  the  same  sex.  Various  crests,  tufts,  and  mantles 
of  hair,  which  are  either  confined  to  the  male,  or  are  more 
developed  in  this  sex  than  in  the  female,  seem  in  most  cases  to  be 
merely  ornamental,  though  they  sometimes  serve  as  a  defence 
against  rival  males.  There  is  even  reason  to  suspect  that  the 
branching  horns  of  stags,  and  the  elegant  horns  of  certain  ante- 
lopes, though  properly  serving  as  weapons  of  offence  or  defence, 
have  been  partly  modified  for  ornament. 

When  the  male  diifers  in  colour  from  the  female,  he  generally 
exhibits  darker  and  more  strongly-contrasted  tints.  We  do  not 
in  this  class  meet  with  the  splendid  red,  blue,  yellow,  and  green 
tints,  so  common  with  male  birds  and  many  other  animals.  The 
naked  parts,  however,  of  certain  Quadrumana  must  be  excepted ; 
for  such  parts,  often  oddly  situated,  are  brilliantly  coloured 
in  some  species.  The  colours  of  the  male  in  other  cases  may  be 
due  to  simple  variation,  without  the  aid  of  selection.  But  when 
the  colours  are  diversified  and  strongly  pronounced,  when  they 
are  not  developed  until  near  maturity,  and  when  they  are  lost 
after  emasculation,  we  can  hardly  avoid  the  conclusion  that 
they  have  been  acquired  through  sexual  selection  for  the  sake  of 
ornament,  and  have  been  transmitted  exclusively,  or  almost 
exclusively,  to  the  same  sex.  When  both  sexes  are  coloured  in 
the  same  manner,  and  the  colours  are  conspicuous  or  curiously 
arranged,  without  being  of  the  least  apparent  use  as  a  protection, 
and  especially  when  they  are  associated  with  various  other  orna- 
mental appendages,  we  are  led  by  analogy  to  the  same  conclusion, 
namely,  that  they  have  been  acquired  through  sexual  selection, 
although  transmitted  to  both  sexes.  That  conspicuous  and 
diversified  colours,  whether  confined  to  the  males  or  common  to 
both  sexes,  are  as  a  general  rule  associated  in  the  same  groups 
and  sub-groups  with  other  secondary  sexual  characters  serving 
for  war  or  for  ornament,  will  be  found  to  hold  good,  if  we  look 
back  to  the  various  cases  given  in  this  and  the  last  chapter. 

The  law  of  the  equal  transmission  of  characters  to  both  sexes, 
as  far  as  colour  and  other  ornaments  are  concerned,  has  prevailed 
far  more  extensively  with  mammals  than  with  birds;  but 
weapons,  such  as  horns  and  tusks,  have  often  been  transmitted 
either  exclusively  or  much  more  perfectly  to  the  males  than  to  the 
females.  This  is  surprising,  for,  as  the  males  generally  use  their 
weapons  for  defence  against  enemies  of  all  kinds,  their  weapons 
would  have  been  of  service  to  the  females.  As  far  as  we  can  sec, 
their  al)sence  in  this  sex  can  be  accounted  for  only  by  iho 


Chap.  XVII r.  Mammals — Sitimnary,  555 


form  of  inheritance  which  has  prevailed.  Finally,  with  quad- 
rupeds the  contest  between  the  individuals  of  the  same  sex, 
whether  peaceful  or  bloody,  has,  with  the  rarest  exceptions, 
been  contined  to  the  males;  so  that  the  latter  have  been  modified 
through  sexual  selection,  far  more  commonly  than  the  females, 
either  for  fighting  with  each  other  or  for  alluring  the  opposite 
sex. 


(    556    ) 


Part  III. 
SEXUAL  SELECTION  IN  EELATION  TO  MAN, 


AND  CONCLUSION. 


CHAPTER  XIX. 
Secondary  Sexual  Characters  of  Man. 

Differences  between  man  and  woman — Causes  of  such  differences  and  of 
certain  characters  common  to  both  sexes — Law  of  battle — Differences  in 
mental  powers,  and  voice — On  the  influence  of  beauty  in  determining 
the  marriages  of  mankind — Attention  paid  by  savages  to  ornaments— 
Their  ideas  of  beauty  in  woman — The  tendency  to  exaggerate  each 
natui-al  ijeculiarity. 

With  mankind  th3  differences  between  the  sexes  are  greater 
than  in  most  of  the  Quadrumana,  but  not  so  great  as  in  some, 
for  instance,  the  mandrill.  Man  on  an  average  is  considerably 
taller,  heavier,  and  stronger  than  woman,  with  sqnarer  shoulders 
and  more  plainly-pronounced  muscles.  Owing  to  the  relation 
which  exists  between  muscular  development  and  the  projection  of 
the  brovvs,"^  the  superciliary  ridge  is  generally  more  marked  in 
man  than  in  woman.  His  body,  and  especially  his  face,  is  more 
hairy,  and  his  voice  has  a  different  and  more  powerful  tone.  In 
certain  races  the  women  are  said  to  differ  slightly  in  tint  from 
the  men.  For  instance,  Schweinfurth,  in  speaking  of  a  negress 
belonging  to  the  Monbuttoos,  who  inhabit  the  interior  of  Africa  a 
few  degrees  north  of  the  Equator,  says,  "  Like  all  lier  race,  she  had 
*•  a  skin  several  shades  lighter  than  her  husband's,  being  some- 
"  thing  of  the  colour  of  half-roasted  coffee."^  As  the  women 
labour  in  the  fields  and  arc  qnite  unclothed,  it  is  not  likely  that 
they  differ  in  colour  from  the  men  owing  to  less  exposnre  to  the 
weather.  European  women  arc  perhaps  the  brighter  coloured  of 
tlie  two  sexes,  as  may  be  seen  when  both  have  been  equally 
exposed. 

'  Schaaffhauson,     translation     in  *  'The  Haart  of  Africa,'  English 

'Anthropological  Review,' Oct.  18G8,      Transl.  1873,  vol.  i.  p.  544. 
pp.  419,  420,  427. 


Chap.  XIX.  Ma?i — Sexual  Differences.  557 


Man  is  more  courageous,  pugnacious  and  energetic  than 
woman,  and  has  a  more  inventive  genius.  His  brain  is  abso- 
lutely larger,  but  whether  or  not  proportionately  to  his  larger 
body,  has  not,  I  believe,  been  fully  ascertained.  In  woman  the 
face  is  rounder  ;  the  jaws  and  the  base  of  the  skull  smaller ;  the 
outlines  of  the  body  rounder,  in  parts  more  prominent ;  and  her 
pelvis  is  broader  than  in  man;^  but  this  latter  character  may 
perhaps  be  considered  rather  as  a  primary  than  a  secondary 
sexual  character.  She  comes  to  maturity  at  an  earlier  age  than 
man. 

As  with  animals  of  all  classes,  so  with  man,  the  distinctive 
characters  of  the  male  sex  are  not  fully  developed  until  he  is 
nearly  mature;  and  if  emasculated  they  never  appear.  The 
beard,  for  instance,  is  a  secondary  sexual  character,  and  mjile 
children  are  beardless,  though  at  an  early  age  they  have  abun- 
dant hair  on  the  head.  It  is  probably  due  to  the  rather  late 
apjjearance  in  life  of  the  successive  variations  whereby  man  has 
acquired  his  masculine  characters,  that  they  are  transmitted  to 
the  male  sex  alone.  Male  and  female  children  resemble  each 
other  closely,  like  the  young  of  so  many  other  animals  in 
which  the  adult  sexes  diifer  widely ;  they  likewise  resemble  the 
mature  female  much  more  closely  than  the  mature  male.  The 
female,  however,  ultimately  assumes  certain  distinctive  cha- 
racters, and  in  the  formation  of  her  skull,  is  said  to  be  inter- 
mediate between  the  child  and  the  man.  *  Again,  as  the  young  of 
closely  allied  though  distinct  species  do  not  differ  nearly  so  much 
from  each  other  as  do  the  adults,  so  it  is  with  the  children  of  the 
different  races  of  man.  Some  have  even  maintained  that  race- 
differences  cannot  be  detected  in  the  infantile  skuli.^  In  regard 
to  colour,  the  new-born  negro  child  is  reddish  nut-brown,  which 
soon  becomes  slaty- grey  ;  the  black  colour  being  fully  developed 
within  a  year  in  the  Soudan,  but  not  until  three  years  in  Egypt. 
The  eyes  of  the  negro  are  at  first  blue,  and  the  hair  chesnut- 
brown  rather  than  black,  being  curled  only  at  the  ends.  The 
children  of  the  Australians  immediately  after  birth  are  yellowish- 
brown,  and  become  dark  at  a  later  age.  Those  of  the  Guaranys 
of  Paraguay  are  whitish-yellow,  but  they  acquire  in  the  course 
of  a  few  weeks  the  yellowish-brown  tint  of  their  joarents. 
Similar  observations  have  been  made  in  other  parts  of  America.*^ 

^  Ecker,  translation  in  'Anthro-  355;  Vogt, '  Lectures  on  Man,' Eng. 

pological  Review,' Oct.  1868,  pp.  351-  translat.  p.  81. 

35'i.     The  comparison  of  the  form  ^  Schaaffhausen,     '  Anthropolog. 

of  the  skull  in  men  and  women  has  Review,'  ibid.  p.  429. 

been  followed  out  with  much  care  "  Pruner-Bey,   on    negro    infants 

by  Welcker.  as   quoted   by   Vogt,   '  Lectures    on 

^  Ecker  and  Welcker,  ibid.  p.  352,  Man,'   Eng.  translat.   18G4-,  p.  189; 


558  The  Desceitt  of  Man.  Part  III. 

I  have  specified  the  foregoing  diiferences  between  the  male  and 
female  sex  in  mankind,  because  they  are  curiously  like  those  of 
the  Quadrumana.  With  these  animals  the  female  is  mature  at 
an  earlier  age  than  the  male ;  at  least  this  is  certainly  the  case  in 
the  Ctbas  uzarcej  The  males  of  most  species  are  larger  and 
stronger  than  the  females,  of  which  fact  the  gorilla  affords  a  well- 
known  instance.  Even  in  so  trifling  a  character  as  the  greater 
prominence  of  the  superciliary  ridge,  the  males  of  certain 
monkeys  differ  from  the  females/  and  agree  in  this  respect  with 
mankind.  In  the  gorilla  and  certain  other  monkeys,  the  cranium 
of  the  adult  male  presents  a  strongly-marked  sagittal  crest,  which 
is  absent  in  the  female ;  and  Ecker  found  a  trace  of  a  similar 
difference  between  the  two  sexes  in  the  Australians.^  With 
monkeys  when  there  is  any  difference  in  the  voice,  that  of  the 
male  is  the  more  powerful.  We  have  seen  that  certain 
male  monkeys  have  a  well-developed  beard,  which  is  quite 
deficient,  or  much  less  developed  in  the  female.  No  instance 
is  known  of  the  beard,  whiskers,  or  moustache  beiug  larger 
in  the  female  than  in  the  male  monkey.  Even  in  the  colour 
of  the  beard  there  is  a  curious  parallelism  between  man  and 
the  Quadrumana,  for  with  man  when  the  beard  differs  in  colour 
from  the  hair  of  the  head,  as  is  commonly  the  case,  it  is,  I  believe, 
almost  always  Qf  a  lighter  tint,  being  often  reddish.  I  have 
repeatedly  observed  this  fact  in  England ;  but  two  gentlemen 
have  lately  written  to  me,  saying  that  they  form  an  exception 
to  the  rule.  One  of  these  gentlemen  accounts  for  the  fact 
by  the  wide  difference  in  colour  of  the  hair  on  the  paternal 
and  maternal  sides  of  his  family.  Both  had  been  long  aware  of 
this  peculiarity  (one  of  them  having  often  been  accused  of 
dyeing  his  beard),  and  had  been  thus  led  to  observe  other  men, 
and  were  convinced  that  the  exceptions  were  very  rare.  Dr. 
Hooker  attended  to  this  little  point  for  me  in  Eussia,  and  found 
no  exception  to  the  rule.  In  Calcutta,  Mr.  J.  Scott,  of  the 
Botanic  Gardens,  was  so  kind  as  to  observe  the  many  races  of  men 
to  be"  seen  there,  as  well  as  in  some  other  parts  of  India,  namely, 
two    races    in  Sikliim,   the    Bhoteas,  Hindoos,   Burmese,  and 


for  further  facts  on   negro  infants,  '   Rengger,      '  Sivugethiere,'      <S:c. 

as  quoted  from   Winterbottom   and  1830,  s.  49. 

Camper,  see  Lawrence,  'Lectures  on  ^  As,  in  Macacus  c>/nomoIgus  (Des- 

Physiology,' &c.   1822,  p.  451.     For  marest,  '  Mammalogie,'  p.  65),  and 

the    intants    of    the    Guaranys,   see  in    Ilylohates    agilis   (GeofiVoy    St.- 

Kengger,   '  Saugethiere,'   &c,    s.    3.  Hilaire  and  F.  Cuvier,  '  Hist.  Nat. 

See  also  Godron,  '  De  I'Esp^ce,'  tom.  des  Llamm.'  1824,  tom.  i.  p.  2). 

ii.    1859,  p.  253.     For  the  Austra-  ^  '  Anthropological  Review,'  Oct. 

Hans,  Waitz,  '  Introduct.  to  Anthro-  1868,  p.  353. 

pology,'  Eng.  translat.  1863,  p.  99. 


Chap.  XIX.         Man — Sexual  Differences.  559 


Chinese,  most  of  which  races  have  very  little  hair  on  the  face 
and  he  always  found  that  when  there  was  any  ditference  in 
colour  between  the  hair  of  the  head  and  the  beard,  the  latter  was 
invariably  lighter.  Now  with  monkeys,  as  has  already  been 
stated,  the  beard  frequently  differs  strikingly  in  colour  from  the 
hair  of  the  head,  and  in  such  cases  it  is  always  of  a  lighter  hue, 
being  often  pure  white,  sometimes  yellow  or  reddish. '° 

In  regard  to  the  general  hairiness  of  the  body,  the  women  in 
all  races  are  less  hairy  than  the  men ;  and  in  some  few 
Quadrumana  the  under  side  of  the  body  of  the  female  is  less 
hairy  than  that  of  the  male.  ^^  Lastly,  male  monkeys,  like  men, 
are  bolder  and  fiercer  than  the  females.  They  lead  the  troop, 
and  when  there  is  danger,  come  to  the  front.  We  thus  see  how 
close  is  the  parallelism  between  the  sexual  differences  of  man 
and  the  Quadrumana.  With  some  few  species,  however,  as 
with  certain  baboons,  the  orang  and  the  gorilla,  there  is  a  con- 
siderably greater  difference  between  the  sexes,  as  in  the  size  of 
the  canine  teeth,  in  the  development  and  colour  of  the  hair,  and 
especially  in  the  colour  of  the  naked  parts  of  the  skin,  than  in 
mankind. 

All  the  secondary  sexual  characters  of  man  are  highly  variable, 
even  within  the  limits  of  the  same  race  ;  and  they  differ  much  in 
the  several  races.  These  two  rules  hold  good  generally  through- 
out the  animal  kingdom.  In  the  excellent  observations  made  on 
board  the  Novara,  ^^  the  male  Australians  were  found  to  exceed 
the  females  by  only  65  millim.  in  height,  whilst  with  the  Javans 
the  average  excess  was  218  millim. ;  so  that  in  this  latter  race  the 
difference  in  height  between  the  sexes  is  more  than  thrice  as 
great  as  with  the  Australians.  Numerous  measurements  were 
carefully  made  of  the  stature,  the  circumference  of  the  neck  and 
chest,  the  length  of  the  back-bone  and  of  the  arms,  in  various 

'"  Mr.  Blyth  inlorms  me  that  he  thus  Mr.  D.  Forbes  has  never,  as  he 

has    only  seen    one  instance  of  the  informs  me,  seen  an  instance  with 

beard,  whiskers,  &c.,  in    a  monkey  the  Aymaras   and    Quichuas    of  S. 

becoming    white    with    old    age,   as  America. 

is    so  commonly  the  case  with   us.  "  This  is  the   case  Avith  the  fe- 

This,      however,     occurred     in    an  males  of  several  species  of  Hylobates, 

aged      Macacus     cynomolgus,    kept  see  GeoffVoy  St.-Hiliare  and  F.  Cu- 

in    confinement,    whose    moustaches  vier,  '  Hist.  Kat.  des  Mamm.'  tom.  i. 

were    "  remarkably    long    and    hu-  See,  also,    on    H.    lar.  '  Fenny  Cy- 

"  man-like."     Altogether    this    old  clopedia,'  vol.  ii.  pp.  149,  150. 

monkey  presented    a    ludicrous  re-  ^2  xhe  results   were    deduced  by 

semblance  to    one   of  the    reigning  Dr.    Weisbach    from    the    measure- 

monarchs  of  Europe,  after  whom  he  ments    made    by   Drs.   K.  Scherzer 

was     universally     nick-named.     In  and  Schwarz,  see  '  Reiseder  Novara: 

certain  races  of    man   the   hair   on  Anthropolog.    Theil,'  1867,  ss.  216, 

the  head  hardly  ever  becomes  grey;  231,  234,  236,  239,  269. 


560  The  Descent  of  Man.  Part  III. 

races ;  and  nearly  all  these  measurements  shew  that  the  males 
differ  much  more  from  one  another  than  do  the  females.  This 
fact  indicates  that,  as  far  as  these  characters  are  concerned,  it  is 
the  male  which  has  been  chiefly  modified,  since  the  several  i-aces 
diverged  from  their  common  stock. 

The  development  of  the  beard  and  the  hairiness  of  the  body 
differ  remarkably  in  the  men  of  distinct  races,  and  even  in 
different  tribes  or  families  of  the  same  race.  We  Europeans  see 
this  amongst  ourselves.  In  the  Island  of  St.  Kilda,  according  to 
Martin,^^  the  m.en  do  not  acquire  beards  until  the  age  of  thirty 
or  upwards,  and  even  then  the  beards  are  very  thin.  On 
the  EuropsBO- Asiatic  continent,  beards  prevail  until  we  pass 
beyond  India ;  though  with  the  natives  of  Ceylon  they  are  often 
absent,  as  was  noticed  in  ancient  times  by  Diodorus.^^  East- 
ward of  India  beards  disappear,  as  with  the  Siamese,  Malays, 
Kalmucks,  Chinese,  and  Japanese ;  nevertheless  the  Ainos,^^  who 
inhabit  the  northernmost  islands  of  the  Japan  Archipelago,  are 
the  hairiest  men  in  the  world.  With  negroes  the  beard  is 
scanty  or  wanting,  and  they  rarely  have  whiskers ;  in  both  sexes 
the  body  is  frequently  almost  destitute  of  fine  down.^*'  On  the 
other  hand,  the  Papuans  of  the  Malay  Archipelago,  who  are 
nearly  as  black  as  negroes,  possess  well-developed  beards.^^ 
In  the  Pacific  Ocean  the  inhabitants  of  the  Fiji  Archipelago  have 
large  bushy  beards,  whilst  those  of  the  not  distant  archi- 
pelagoes of  Tonga  and  Samoa  are  beardless;  but  these  men 
belong  to  distinct  races.  In  the  Ellice  group  all  the  inhabitants 
belong  to  the  same  race;  yet  on  one  island  alone,  namely 
Nunemaya,  "the  men  have  sj^lendid  beards;"  whilst  on  the 
other  islands  "  they  have,  as  a  rule,  a  dozen  straggling  hairs  for 
"a  beard." ^** 

Throughout  the  great  American  continent  the  men  may  be 
said  to  be  beardless ;  but  in  almost  all  the  tribes  a  few  short 
hairs  are  apt  to  appear  on  the  face,  especially  in  old  age.  With 
the  tribes  of  North  America,  Catlin  estimates  that  eighteen  out 
of  twenty  men  are  completely  destitute  by  nature  of  a  beard ; 

13  'Voyage   to    St.     Kilda'  (3rd  remarkable  that  in  the  United  States 

edit.  1753),  p.  37.  ('  Investigations     in     Military    and 

'*  Sir    J.    E.    Tennent,    '  Ceylon,'  Anthropological  Statistics  of  Ame- 

vol.  ii.  1859,  p.  107.  rican   Soldiers,'   1869,  p.  569)    the 

'*  Quatrefages,  '  Revue  des  Cours  pure  negroes  and  their  crossed  otF- 

Scientitiques,'    Aug.    29,    1868,    p.  spring  seem  to  have  bodies  almost 

630;     Vogt     'Lectures     on     Man,'  as  hairy  as  Europeans. 

Eug.  translat.  p.  127.  *^  Wallace,    'The    Malay    Arch.' 

'^  On  the  beards  of  negroes,  Vogt,  vol.  ii.  1869,  p.  178. 

'Lectures,'  &c.  p.  127;   Waitz,  'In-  '^  Dr.  J.  Barnard  Davis  On  Oce- 

troduct.    to    Anthropology,'    Engl.  anic  Races,  in 'Anthropolog.  Review,' 

translat.   1863,  vol.  i.  p.  96.     It^is  April,  1870,  pp.  185,  191. 


Chap.  XIX.  Man — Laiv  of  Battle.  561 

but  occasionally  there  may  be  seen  a  man,  who  has  neglected  to 
pluck  out  the  hairs  at  puberty,  with  a  soft  beard  an  inch  or  two 
iu  length.  The  Guaranys  of  Paraguay  differ  from  all  the 
sarrounding  tribes  in  having  a  small  beard,  and  even  some  hair 
on  the  body,  but  no  whiskers.^^  I  am  informed  by  Mr.  1). 
Forbes,  who  particularly  attended  to  this  point,  that  the  Aymaras 
and  Quichuas  of  the  Cordillera  are  remarkably  hairless,  yet  in 
old  age  a'  few  straggling  hairs  occasionally  appear  on  the  chin. 
The  men  of  these  two  tribes  have  very  little  hair  on  the  various 
parts  of  the  body  where  hair  grows  abundantly  in  Europeans, 
and  the  women  have  none  on  the  corresponding  parts.  The 
hair  on  the  head,  however,  attains  an  extraordinary  length  in 
both  sexes,  often  reaching  almost  to  the  ground ;  and  this  is 
likewise  the  case  with  some  of  the  N.  American  tribes.  In  the 
amount  of  hair,  and  iu  the  general  shape  of  the  body,  the 
sexes  of  the  American  aborigines  do  not  differ  so  much  from 
each  other,  as  in  most  other  races.^"  This  fact  is  analogous 
with  what  occurs  with  some  closely  allied  monkeys;  thus  the 
sexes  of  the  chimpanzee  are  not  as  diffei-ent  as  those  of  the  orang 
or  gorilla!^^ 

In  the  previous  chapters  we  have  seen  that  with  mammals, 
birds,  fishes,  insects,  &c.,  many  characters,  which  there  is  every 
reason  to  believe  were  primarily  gained  through  sexual  selection 
by  one  sex,  have  been  transferred  to  the  other.  As  this  same  form 
of  transmission  has  apparently  prevailed  much  with  mankind, 
it  will  save  useless  repetition  if  we  discuss  the  origin  of  cha- 
racters peculiar  to  the  male  sex  together  with  certain  other 
characters  common  to  both  sexes. 

Lain  of  Battle.— ^i'ih  savages,  for  instance  the  Australians,  the 
women  are  the  constant  cause  of  war  both  between  members  of 
the  same  tribe  and  between  distinct  tribes.  So  no  doubt  it  was 
in  ancient  times ;  "  nam  fuit  ante  Helenam  mulier  teterrima  belli 
"  causa."  With  some  of  the  North  American  Indians,  the  con- 
test is  reduced  to  a  system.     That  excellent  observer,  Hearne,-'- 

'^  Catlin,  'North    American    la-  21  Rutimeyer,    'Die  Grenzen   der 

(Hans,'  3rd  edit.  1842,  vol.  ii.  p.  227.  Thierwelt ;     eine    Betrachtung    zu 

On  the  Guaranys,  see  Azara,  '  Voy-  Dnrwin's  Lehre,'  1868,  s.  54. 
ages  dans  I'Ame'rique  Merid.'  torn.  -^  '  A    Journey    from    Prince    of 

ii.    1809,    p.     58;     also    Rengger,  Wales  Fort.' 8vo.  edit.  Dublin,  179i5, 

'  Saugethiere  von  Paraguay,'  s.  3.  p.  1(34.     Sir  J.  Lubboclc  ('  Origin  ot 

2»  Prof,  and  Mrs.  Agassiz  (' Jour-  Civilisation,'    1870,    p.    69)    gives 

uey  in  Brazil,'  p.  530)  remark  that  other  and  similar  cases    in    North 

the  sexes  of  the  American  Indians  America.     For   the    Guanas    of    S. 

differ  less  than  those  of  the  negroes  America  see  Azara,   '  Voyages,'  &c. 

f.nd  of  the  higher  races.     See  also  torn.  ii.  p.  94. 

25 


562  The  Descent  of  Man.  Part  111. 

says : — "  It  has  ever  been  the  custom  among  these  people  for 
"  the  men  to  wrestle  for  any  woman  to  whom  they  are  at- 
"  tached ;  and,  of  coui'se,  the  strongest  party  always  carries  oft 
"  the  prize.  A  weak  man,  unless  he  be  a  good  hunter,  and  well- 
"  beloved,  is  seldom  permitted  to  keep  a  wife  that  a  stronger 
"  man  thinks  worth  his  notice.  This  custom  prevails  through- 
"  out  all  the  tribes,  and  causes  a  great  spirit  of  emulation  among 
"  their  youth,  who  are  upon  all  occasions,  from  their  childhood, 
"  trying  their  strength  and  skill  in  wrestling.'^  With  the  Guanas 
of  South  America,  Azara  states  that  the  men  rarely  marry  till 
twenty  years  old  or  more,  as  before  that  age  they  cannot  conquer 
their  rivals. 

Other  similar  facts  could  be  given ;  but  even  if  we  had  no 
evidence  on  this  head,  we  might  feel  almost  sure,  from  the 
analogy  of  the  higher  Quadi'umana,'^^  that  the  law  of  battle  had 
prevailed  with  man  during  the  early  stages  of  his  development. 
The  occasional  appearance  at  the  i^resent  day  of  canine  teeth 
which  project  above  the  others,  with  traces  of  a  diastema  or 
oiDen  space  for  the  reception  of  the  opposite  canines,  is  in  all 
probability  a  case  of  reversion  to  a  former  state,  when  the 
progenitors  of  man  were  provided  with  these  weapons,  like  so 
many  existing  male  Quadrumana.  It  was  remarked  in  a  former 
chapter  that  as  man  gradually  became  erect,  and  continually 
used  his  hands  and  arms  for  fighting  with  sticks  and  stones,  as 
well  as  for  the  other  purposes  of  life,  he  would  have  used  his 
jaws  and  teeth  less  and  less.  The  jaws,  together  with  their 
muscles,  would  then  have  been  reduced  through  disuse,  as 
would  the  teeth  through  the  not  well  understood  principles  of 
correlation  and  economy  of  growth ;  for  we  everywhere  see 
that  parts,  which  are  no  longer  of  service,  are  reduced  in  size. 
By  such  steps  the  original  inequality  between  the  jaws  and  teeth 
in  the  two  sexes  of  mankind  would  ultimately  have  been  obhte- 
rated.  1'he  case  is  almost  parallel  with  that  of  many  male 
Ruminants,  in  which  the  canine  teeth  have  been  reduced  to 
mere  rudiments,  or  have  disappeared,  apparently  in  consequence 
of  the  development  of  horns.  As  the  p.iodigious  difference 
between  the  skulls  of  the  two  sexes  in  the  orang  and  gorilla 
stands  in  close  relation  with  the  development  of  the  immense 
canine  teeth  in  the  males,  we  may  infer  that  the  reduction  of 
the  jaws  and  teeth  in  the  early  male  progenitors  of  man  must 
have  led  to  a  most  striking  and  favourable  change  in  his 
appearance. 

^2  On  the  fighting  of   the    male      p.  423.     On  Prcib]itis    cnicUxs,  see 
goriUas,  see  Dv.  Savage,  in  'Boston      llie  'Indian  Field,'  1859,  p.  146. 
Journal  ol"  Nat.   Hist.'  vol.  v.  1847, 


Chap.  XIX.  Menial pozvcrs  of  Man  and  Woman,        563 

There  can  be  little  doubt  that  the  greater  size  and  strength  of 
man,  in  comparison  with  woman,  together  with  his  broader 
shoulders,  more  developed  muscles,  rugged  outline  of  body,  his 
greater  courage  and  pugnacity,  are  all  due  in  chief  part  to  in- 
heritance from  his  half-human  male  ancestors.  These  characters 
would,  however,  have  been  preserved  or  even  augmented  during 
the  long  ages  of  man's  savagery,  by  the  success  of  the  strongest 
and  boldest  men,  both  in  the  general  struggle  for  life  and  in 
their  contests  for  wives ;  a  success  which  would  have  ensured 
their  leaving  a  more  numerous  progeny  than  their  less  favoured 
brethren.  It  is  not  probable  that  the  greater  strength  of  man 
was  primarily  acquired  through  the  inherited  effects  of  his 
having  worked  harder  than  woman  for  his  own  subsistence 
and  that  of  his  family  ;  for  the  women  in  all  barbarous  nations 
are  compelled  to  work  at  least  as  hard  as  the  men.  With 
civilised  people  the  arbitrament  of  battle  for  the  possession  of 
the  women  has  long  ceased ;  on  the  other  hand,  the  men,  as  a 
general  rule,  have  to  work  harder  than  the  women  for  their 
joint  subsistence,  and  thus  their  greater  strength  will  have  been 
kept  up. 

Difference  in  the  Mental  Poicers  of  the  two  Sexes.- — With  respect 
to  differences  of  this  nature  between  man  and  woman,  it  is 
probable  that  sexual  selection  has  played  a  highly  important 
part.  I  am  aware  that  some  writers  doubt  whether  there  is  any 
such  inherent  difference ;  but  this  is  at  least  probable  from  the 
analogy  of  the  lower  animals  which  present  other  secondary 
sexual  characters.  No  one  disputes  that  the  bull  differs  in 
disposition  from  the  cow,  the  wild-boar  from  the  sow,  the 
stallion  from  the  mare,  and,  as  is  well  known  to  the  keepers  of 
menageries,  the  males  of  the  larger  apes  from  the  females. 
Woman  seems  to  differ  from  man  in  mental  disposition,  chiefly 
in  her  greater  tenderness  and  less  selfislmess;  and  this  holds 
good  even  with  savages,  as  shewn  by  a  well-known  passage  in 
Mungo  Park's  Travels,  and  by  statements  made  by  many  other 
travellers.  Woman,  owing  to  her  maternal  instincts,  displays 
these  qualities  towards  her  infants  in  an  eminent  degree  ;  there- 
fore it  is  likely  that  she  would  often  extend  them  towards  her 
fellow- creatures.  Man  is  the  rival  of  other  men ;  he  delights 
in  competition,  and  this  leads  to  ambition  which  passes  too 
easily  into  selfishness.  These  latter  qualities  seem  to  be  his 
natural  and  unfortunate  birthright.  It  is  generally  admitted 
that  with  woman  the  powers  of  intuition,  of  rapid  perception, 
and  perhaps  of  imitation,  are  more  strongly  marked  than  in 
man:   but  some,  at  least,  of  these  faculties  are  characteristi'' 


564  The  Descent  of  Man.  Part  III. 

of  the  lower  races,  and  therefore  of  a  past  and  lower  state  of 
civilisation. 

The  chief  distinction  in  the  intellectual  powers  of  the  two 
sexes  is  shewn  by  man's  attaining  to  a  higher  eminence,  in 
whatever  he  takes  np,  than  can  woman — whether  requiring 
deep  thought,  reason,  or  imagination,  or  merely  the  use  of  the 
senses  and  hands.  If  two  Ksts  were  made  of  the  most  eminent 
men  and  women  in  poetry,  painting,  sculpture,  music  (inclusive 
both  of  composition  and  performance),  history,  science,  and 
philosophy,  wath  half-a-dozen  names  under  each  subject,  the 
two  hsts  would  not  bear  comparison.  We  may  also  infer,  from 
the  law  of  the  deviation  from  averages,  so  well  illustrated  by 
Mr.  Galton,  in  his  work  on  *  Hereditary  Genius,'  that  if  men  are 
capable  of  a  decided  pre-eminence  over  women  in  many  subjects, 
the  average  of  mental  power  in  man  must  be  above  that  of  woman. 

Amongst  the  half-human  progenitors  of  man,  and  amongst 
savages,  there  have  been  struggles  between  the  males  during 
many  generations  for  the  possession  of  the  females.  But  mere 
bodily  strength  and  size  would  do  little  for  victory,  unless 
associated  with  courage,  perseverance,  and  determined  energy. 
With  social  animals,  the  young  males  have  to  pass  through 
many  a  contest  before  they  win  a  female,  and  the  older  males 
have  to  retain  their  females  by  renewed  battles.  They  have, 
also,  in  the  case  of  mankind,  to  defend  their  females,  as  well  as 
their  young,  from  enemies  of  all  kinds,  and  to  hunt  for  their  joint 
subsistence.  But  to  avoid  enemies  or  to  attack  them  with 
success,  to  capture  wild  animals,  and  to  fashion  weapons,  requires 
the  aid  of  the  higher  mental  faculties,  namely,  observation, 
reason,  invention,  or  imagination.  These  various  faculties  will 
thus  have  been  continually  put  to  the  test  and  selected  during 
manhood ;  they  will,  moreover,  have  been  strengthened  by  use 
during  this  same  period  of  life.  Consequently,  in  accordance 
with  the  principle  often  alluded  to,  we  might  expect  that  they 
would  at  least  tend  to  be  transmitted  chiefly  to  the  male  offspring 
at  the  corresponding  period  of  manhood. 

Now,  when  two  men  are  put  into  competition,  or  a  man  with 
a  woman,  both  possessed  of  every  mental  quality  in  equal  per- 
fection, save  that  one  has  higher  energy,  perseverance,  and 
courage,  the  latter  will  generally  become  more  eminent  in  every 
pursuit,  and  will  gain  the  ascendancy.^^  He  may  be  said 
to  possess  genius — for  genius  has  been  declared  by  a  great 

•^  J.  Stuart   Mill  remarks  ('  The  "  require  most  plodding,  and    long 

Subjection  of  Women,' 18G9,  J).  122),  "hammering    at   single   thoughts." 

"The  tilings   in    whicli   man    most  What   is  this  but  energy  and  pcr- 

"  excels    woman     are    those     which  severance? 


Chap.  XIX.  Mental  powers  of  Man  and  Woman.       565 

authority  to  be  x>atiencG ;  and  patience,  in  this  sense,  means 
unflinching,  undaunted  perseverance.  But  this  view  of  genius 
is  perhaps  deficient ;  for  without  the  higher  powers  of  the  imagi- 
nation and  reason,  no  eminent  success  can  be  gained  in  many 
subjects.  These  latter  faculties,  as  well  as  the  former,  will  have 
been  developed  in  man,  partly  through  sexual  selection, — that 
is,  through  the  contest  of  rival  males,  and  partly  through  natural 
selection, — that  is,  from  success  in  the  general  struggle  for  life ; 
and  as  in  both  cases  the  struggle  will  have  been  during  maturity, 
the  characters  gained  will  have  been  transmitted  more  fully  to 
the  wale  than  to  the  female  offspring.  It  accords  in  a  striking 
manner  with  this  view  of  the  modification  and  re-inforcement  of 
many  of  our  mental  faculties  by  sexual  selection,  that,  firstly, 
they  notoriously  undergo  a  considerable  change  at  puberty,-'^ 
and,  secondly,  that  eunuchs  remain  throughout  life  inferior  in 
these  same  qualities.  Thus  man  has  ultimately  become  superior 
to  woman.  It  is,  indeed,  fortunate  that  the  law  of  the  equal 
transmission  of  characters  to  both  sexes  prevails  with  mammals ; 
otherwise  it  is  probable  that  man  would  have  become  as  superior 
in  mental  endowment  to  woman,  as  the  peacock  is  in  ornamental 
plumage  to  the  peahen. 

It  must  be  borne  in  mind  that  the  tendency  in  characters 
acquired  by  either  sex  late  in  life,  to  be  transmitted  to  the  same 
sex  at  the  same  age,  and  of  early  acquired  characters  to  be 
transmitted  to  both  sexes,  are  rules  which,  though  general,  do 
not  always  hold.  If  they  always  held  good,  we  might  conclude 
(but  I  here  exceed  my  proper  bounds)  that  the  inherited  elfects 
of  the  early  education  of  boys  and  girls  would  be  transmitted 
equally  to  both  sexes;  so  that  the  present  inequality  in  mental 
power  between  the  sexes  would  not  be  effaced  by  a  similar  course 
of  early  training ;  nor  can  it  have  been  caused  by  then-  dissimilar 
early  training.  In  order  that  woman  should  reach  the  same 
standard  as  man,  she  ought,  when  nearly  adult,  to  be  trained  to 
energy  and  perseverance,  and  to  have  her  reason  and  imagination 
exercised  to  the  highest  point;  and  then  she  would  probably 
transmit  these  qualities  chiefly  to  her  adult  daughters.  All  women, 
however,  could  not  be  thus  raised,  unless  during  many  genera- 
tions those  who  excelled  in  the  above  robust  virtues  were 
married,  and  produced  offspring  in  larger  numbers  than  other 
women.  As  before  remarked  of  bodily  strength,  although  men 
do  not  now  fight  for  their  wives,  and  this  form  of  selection  has 
passed  away,  yet  during  manhood,  they  generally  undergo  a 
severe  struggle  in  order  to  maintain  themselves  and  their 
fiimilies ;  and  this  will  tend  to  keep  up  or  even  increase  theii 

^5  JMaudsley,  '  Miiid  auJ  I'.ody,'  p.  31. 


566  The  Descent  of  Man.  Part  III. 

mental  powers,  and,  as  a  consequence,  the  present  inequality 
between  the  sexes.^*^ 

Voice  and  Musical  Powers. — In  some  species  of  Quaclrumana 
there  is  a  great  difference  between  the  adult  sexes,  in  the  ix)wer 
of  their  voices  and  in  the  development  of  the  vocal  organs ;  and 
man  api^ears  to  have  inherited  this  difference  from  his  early 
iprogenitors.  His  vocal  cords  are  about  one-third  longer  than  in 
woman,  or  than  in  boys ;  and  emasculation  produces  the  same 
effect  on  him  as  on  the  lower  animals,  for  it  "  arrests  that  pro- 
"  minent  growth  of  the  thyroid,  &c.,  which  accompaniea  the 
"  elongation  of  the  cords."  -^  With  resj^ect  to  the  cause  of  this 
difference  between  the  sexes,  I  have  nothing  to  add  to  the 
remarks  in  the  last  chapter  on  the  probable  effects  of  the  long- 
continued  use  of  the  vocal  organs  by  the  male  under  the  ex- 
citement of  love,  rage  and  jealousy.  According  to  Sir  Duncan 
Gibb,-**  the  voice  and  the  form  of  the  larynx  differ  in  the  different 
races  of  mankind ;  but  with  the  Tartars,  Chinese,  &c.,  the  voice 
of  the  male  is  f^aid  not  to  differ  so  much  from  that  of  the  female, 
as  in  most  other  races. 

The  capacity  and  love  for  singing  or  music,  though  not  a 
sexual  character  in  man,  must  not  here  be  passed  over.  Although 
the  sounds  emitted  by  animals  of  all  kinds  serve  many  purposes, 
a  strong  case  can  be  made  out,  that  the  vocal  organs  were 
primarily  used  and  perfected  in  relation  to  the  propagation  of  the 
species.  Insects  and  some  few  spiders  are  the  lowest  animals 
wliich  volmitarily  j^roduce  any  sound;  and  this  is  generally 
effected  by  the  aid  of  beautifully  constructed  stridulating  organs, 
which  are  often  confined  to  the  males.  The  sounds  thus  pro- 
duced consist,  I  believe  in  all  cases,  of  the  same  note,  repeated 
rhythmically ;  ^  and  this  is  sometimes  ^Dleasing  even  to  the 
ears  of  man.  The  chief  and,  in  some  cases,  exclusive  purpose 
appears  to  be  either  to  call  or  charm  the  opposite  sex. 

The  sounds  produced  by  fishes  are  said  in  some  cases  to  be 
made  only  by  the  males  during  the  breeding-season.    All  the 

^*  An  observation  by  Vogt  bears  "  skulls."     But  Vogt  admits  ('  Lec- 

on  this  subject :    he  says,  "It  is   a  tures  on  Man,*  Eng.  translat.  18G4, 

*'  remarkable  circumstance,  that  the  p.   81)   that  more  observations  are 

"  dilferencc    between  the    sexes,  as  requisite  on  this  point, 
"regards   the     cranial     cavity,    in-  '•''  Owen,     'Anatomy    of    Verte- 

"  creases   with   the   deveh^pment   of  brates,' vol.  iii.  p.  G03. 
"  the  race,  so  that  the  male  Euro-  -**  '  Journal  of  the  Anthropolog, 

*'  pean  excels  much  more  the  female,  Soc'  April,  1869,  p.  Ivii.  and  Ixvi. 
"than     the     negro     the     negress.  -"  Dr.  Scudder,  'Notes   on    Stri- 

"  Welcker   confirms  this  statement  duhitiou,'  in   '  Proc.  lioston   Soc.  of 

"of    Ilusclike     fi-om    his    measure-  Nat.  Hist.' vol.  xi.  April,  18G8. 
"  ments     of     ncirro     and     German 


Chap.  XIX .  Man  —Musical  Powers.  5  67 


air-breathing  Vortcbrata  necessarily  possess  an  apparatus  for 
inhaling  and  expelling  air,  with  a  pipe  capable  of  being  closed 
at  one  end.  Hence  when  the  primeval  members  of  this  class 
were  strongly  excited  and  their  muscles  violently  contracted, 
purposeless  sounds  would  almost  certainly  have  been  produced ; 
and  these,  if  they  proved  in  any  way  serviceable,  might  readily 
have  been  modified  or  intensified  by  the  preservation  of  properly 
adapted  variations.  The  lowest  Vertebrates  which  breathe  air 
are  Amphibians;  and  of  these,  frogs  and  toads  possess  vocal 
organs,  wiiich  are  incessantly  used  during  the  breeding- season, 
and  which  are  often  more  highly  developed  in  the  male  than  in 
the  female.  The  male  alone  of  the  tortoise  utters  a  noise,  and 
this  only  during  the  season  of  love.  Male  alligators  roar  or 
bellow  during  the  same  season.  Every  one  knows  how  much 
birds  use  their  vocal  organs  as  a  means  of  courtship ;  and  some 
species  likewise  perform  what  may  be  called  instrumental  music. 
In  the  class  of  Mammals,  with  which  we  are  here  more  par- 
ticularly concerned,  the  males  of  almost  all  the  species  use  their 
voices  during  the  breeding-season  much  more  than  at  any  other 
time;  and  some  are  absolutely  mute  excepting  at  this  season. 
With  other  species  both  sexes,  or  only  the  females,  use  their 
voices  as  a  love-call.  Considering  these  facts,  and  that  the  vocal 
organs  of  some  quadrupeds  are  much  more  largely  developed  in 
the  male  than  in  the  female,  either  permanently  or  temporarily 
during  the  breeding-season ;  and  considering  that  in  most  of  the 
lower  classes  the  sounds  produced  by  the  males,  serve  not  only 
to  call  but  to  excite  or  allure  the  female,  it  is  a  surprising  fact 
that  wo  have  not  as  yet  any  good  evidence  that  these  organs  are 
used  by  male  mammals  to  charm  the  females.  The  American 
Mycdes  caraya  perhaps  forms  an  exception,  as  does  the  Hylohatcs 
agiUs,  an  ape  allied  to  man.  This  gibbon  has  an  extremely  loud 
but  musical  voice.  Mr.  Waterhouse  states,^"  "  It  appeared  to 
"  me  that  in  ascending  and  descending  the  scale,  the  intervals 
"  were  always  exactly  half-tones ;  and  I  am  sure  that  the  highest 
"  note  was  the  exact  octave  to  the  lowest.  The  quality  of  the 
"  notes  is  very  musical ;  and  I  do  not  doubt  that  a  good  violinist 
"  would  be  able  to  give  a  correct  idea  of  the  gibbon's  compo- 
"  sition,  excepting  as  regards  its  loudness."  Mr.  Waterhouse 
then  gives  the  notes.  Professor  Owen,  who  is  a  musician,  con- 
firms the  foregoing  statement,  and  remarks,  though  erroneously, 
that  this  gibbon  "alone  of  brute  mammals  may  be  said  to 
"  sing."    It  appears  to  be  much  excited  after  its  performance. 

30  Given    in    W.    C.    L.    Martin's      Owen,    '  Anatomy    of   Vertebrates,' 
'Oeneral  lutroduct.  to  Nat.  Hist,  of     vol.  iii.  p.  000. 
Mamm.    AnimaLs,'     18-H,    p.   4;]2 ; 


568  The  Descent  of  Man.  Part  III. 

Unfortunately,  its  habits  liave  never  been  closely  observed  in  a 
state  of  nature;  but  from  the  analogy  of  other  animals,  it  is 
probable  that  it  uses  its  musical  powers  more  especially  during 
the  season  of  courtship. 

This  gibbon  is  not  the  only  species  in  the  genus  which  sings, 
for  my  son,  Francis  Darwin,  attentively  listened  in  the  Zoo- 
logical Gardens  to  H.  leuciscus  whilst  singing  a  cadence  of  three 
notes,  in  true  musical  intervals  and  with  a  clear  musical  tone. 
It  is  a  more  surprising  fact  that  certain  rodents  utter  musical 
sounds.  Singing  mice  have  often  been  mentioned  and  exhibited, 
but  imposture  has  commonly  been  suspected.  We  have,  how- 
ever, at  last  a  clear  account  by  a  well-knowTi  observer,  the  Eev. 
S.  Lockwood,^^  of  the  musical  powers  of  an  American  species, 
the  Hesperomys  cognatus,  belonging  to  a  genus  distinct  from  that 
of  the  English  mouse.  This  little  animal  was  kept  in  confine- 
ment, and  the  performance  was  repeatedly  heard.  In  one  of  the 
two  chief  songs,  "  the  last  bar  would  frequently  be  prolonged  to 
*'  two  or  three ;  and  she  would  sometimes  change  from  C  sharp 
**  and  D,  to  C  natural  and  D,  then  warble  on  these  two  notes 
"  aw^hile,  and  wind  up  with  a  quick  chirp  on  C  sharp  and  D. 
"  The  distinctness  between  the  semitones  was  very  marked,  and 
"  easily  appreciable  to  a  good  ear."  Mr.  Lockwood  gives  both 
songs  in  musical  notation;  and  adds  that  though  this  little 
mouse  "  had  no  ear  for  time,  yet  she  would  keep  to  the  key  of  B 
"  (two  flats)  and  strictly  in  a  major  key."  ..."  Her  soft  clear 
"  voice  falls  an  octave  with  all  the  precision  possible ;  then  at  the 
"  wind  up,  it  rises  again  into  a  very  quick  trill  on  C  sharp 
"  and  D." 

A  critic  has  asked  how  the  cars  of  man,  and  he  ought  to  have 
added  of  other  animals,  could  have  been  adapted  by  selection  so 
as  to  distinguish  musical  notes.  But  this  question  shews  some 
confusion  on  the  subject ;  a  noise  is  the  sensation  resulting  from 
the  co-existence  of  several  aerial  "simple  vibrations"  of  various 
periods,  each  of  w'hich  intermits  so  frequently  that  its  separate 
existence  cannot  be  perceived.  It  is  only  in  the  want  of  con- 
tinuity of  such  vibrations,  and  in  their  want  of  harmony  inter  se, 
that  a  noise  diifers  from  a  musical  note.  Thus  an  ear  to  be 
capable  of  discriminating  noises — and  the  high  importance  of 
this  power  to  all  animals  is  admitted  by  every  one — must  be  sen- 
sitive to  musical  notes.  AYe  have  evidence  of  this  capacity  even 
low  down  in  the  animal  scale:  thus  Crustaceans  are  provided 
with  auditory  hairs  of  diflercnt  lengths,  which  have  been  seen  to 
vibrate  when  the  proper  musical  notes  are  struck.^^    As  stated 

=*'  The     'American      Naturalist,'  ^2  Hehnholtz,  'Tliebrie  Phys.  de 

1871,  p.  7G1.  la  Miisi(iue,'  1868,  p.  187. 


(Jhap.  XIX.  Man — Musical  Powers,  569 


in  a  previous  chapter,  similiar  observations  have  been  niado 
on  the  hairs  of  the  antennrc  of  gnats.  It  has  been  positively 
asserted  by  good  observers  that  spiders  are  attracted  by  music. 
It  is  also  well  known  that  some  dogs  howl  when  hearing  par- 
ticular toncs.^^  Seals  apparently  appreciate  music,  and  their 
fondness  for  it  "  was  well  known  to  the  ancients,  and  is  often 
"  taken  advantage  of  by  the  hunters  at  the  present  day."  ^^ 

Therefore,  as  far  as  the  mere  perception  of  musical  notes  is 
concerned,  there  seems  no  special  difficulty  in  the  case  of  man 
or  of  any  other  animal.  Helmholtz  has  explained  on  physio- 
logical principles  why  concords  are  agreeable,  and  discords  dis- 
agreeable to  the  human  car ;  but  we  are  Httle  concerned  with 
these,  as  music  in  harmony  is  a  late  invention.  We  are  more  con- 
cerned with  melody,  and  here  again,  according  to  Helmholtz,  it  is 
intelligible  why  the  notes  of  our  musical  scale  are  used.  The 
ear  analyses  all  sounds  into  their  component "  simple  vibrations," 
although  we  are  not  conscious  of  this  analysis.  In  a  musical 
note  the  lowest  in  pitch  of  these  is  generally  predominant,  and 
the  others  which  are  less  marked  are  the  octave,  the  twelfth, 
the  second  octave,  &c.,  all  harmonies  of  the  fundamental  pre- 
dominant note ;  any  two  notes  of  our  scale  have  many  of  these 
harmonic  over-tones  in  common.  It  seems  pretty  clear  then, 
that  if  an  animal  always  wished  to  sing  precisely  the  same  song, 
he  would  guide  himself  by  sounding  those  notes  in  succession, 
which  possess  many  over-tones  in  common — that  is,  he  would 
choose  for  his  song,  notes  which  belong  to  our  musical  scale. 

But  if  it  be  further  asked  why  musical  tones  in  a  certain 
order  and  rhythm  give  man  and  other  animals  pleasure,  we 
can  no  more  give  the  reason  than  for  the  pleasantness  of  certain 
tastes  and  smells  That  they  do  give  pleasure  of  some  kind  to 
animals,  we  may  infer  from  their  being  produced  during  the 
season  of  courtship  by  many  insects,  spiders,  fishes,  amphibians, 
and  birds ;  for  unless  the  females  were  able  to  appreciate  such 
sounds  and  were  excited  or  charmed  by  them,  the  persevering 
efforts  of  the  males,  and  the  complex  structures  often  possessed 
by  them  alone,  would  be  useless ;  and  this  it  is  impossible  to 
believe. 

Human  song  is  generally  admitted  to  be  the  basis  or  origin 
of  instrumental  music.  As  neither  the  enjoyment  nor  the 
capacity  of  producing  nuisical  notes  are  faculties  of  the  least 

"^  Several  accounts  have  been  another  instance  of  a  dog  always 
published  to  tliis  effect.  ]\lr.  Peacli  whining,  when  one  note  on  a  con- 
writes  to  me  that  he  has  repeatedly  certina,  which  was  out  of  tunc,  was 
found  that  an  old  dog  of  his  howls  played. 

when  B  flat  is  sounded  on  the  flute,  ^■*  Mr.  R.  Brown,  in  '  Proc.  Zool. 

and  to  no  other   note.     I  may  add  boc'  1868,  p.  41U. 


570  The  Descent  of  Man.  Part  111. 

use  to  man  in  reference  to  liis  daily  habits  of  life,  they  must  be 
ranked  amongst  the  most  mysterious  with  which  he  is  endowed. 
They  are  present,  though  in  a  very  rude  condition,  in  men  of  all 
races,  even  the  most  savage ;  but  so  different  is  the  taste  of  the 
several  races,  that  our  music  gives  no  pleasure  to  savages,  and 
their  music  is  to  us  in  most  cases  hideous  and  unmeaning.  Dr. 
Seemann,  in  some  interesting  remarks  on  this  subject,^^  "  doubts 
*'  whether  even  amongst  the  nations  of  Western  Europe,  in- 
"  timately  connected  as  they  are  by  close  and  frequent  inter- 
"  course,  the  music  of  the  one  is  interpreted  in  the  same  sense  by 
"  the  others.  By  travelling  eastwards  we  find  that  there  is  cer- 
"  tainly  a  different  language  of  music.  Songs  of  joy  and  dancc- 
"  accompaniments  are  no  longer,  as  with  us,  in  the  major  keys, 
"  but  always  in  the  minor."  Whether  or  not  the  half-human 
progenitors  of  man  possessed,  like  the  singing  gibbons,  tlie 
capacity  of  producing,  and  therefore  no  doubt  of  appreciating, 
musical  notes,  we  know  that  man  possessed  these  faculties  at  a 
very  remote  period.  M.  Lartet  has  described  two  flutes,  made 
out  of  the  bones  and  horns  of  the  reindeer,  found  in  caves 
together  v/ith  flint  tools  and  the  remains  of  extinct  animals. 
The  arts  of  singing  and  of  dancing  are  also  very  ancient,  and 
are  now  practised  by  all  or  nearly  all  the  lowest  races  of  man. 
Poetry,  wliicli  may  be  considered  as  the  offspring  of  song,  is  like- 
wise so  ancient,  that  many  persons  have  felt  astonished  that  it 
should  have  arisen  during  the  earliest  ages  of  which  we  have 
any  record. 

We  see  thiit  the  musical  faculties,  whicli  are  not  wholly  deficient 
in  any  race,  are  capable  of  prompt  and  high  development,  for  Hot- 
tentots and  Negroes  have  become  excellent  musicians,  although  in 
their  native  countries  they  rarely  practise  anything  that  we  should 
consider  music.  Schweinfurth,  however,  was  pleased  with  some 
of  the  simple  melodies  which  he  heard  in  the  interior  of  Africa. 
But  there  is  nothing  anomalous  in  the  musical  faculties  lying 
dormant  in  man :  some  species  of  birds  which  never  naturally 
sing,  can  without  much  difficulty  be  taught  to  do  so;  thus  a 
house-sparrow  has  learnt  the  song  of  a  linnet.  As  these  two 
species  are  closely  allied,  and  belong  to  the  ordci-  of  Insessores, 
which  includes  nearly  all  the  singing-birds  in  the  world,  it  is 
possible  that  a  progenitor  of  the  sparrow  may  have  been  a 
songster.  It  is  more  remarkable  that  parrots,  belonging  to 
a  group  distinct  from  the  Insessores,  and  having  differently 

35  'Journal  of  Anthropolog.  Soc'  cond  eiKtion,   1869,  which    confaiu 

Oct.    1870,    p.   civ.     See    also    the  an  admirable  account  of  the  liabits 

several  later  cha))tor.s  in   Sir  John  of  savages. 
Lubbock's    '  Prehistoric  Times,'  t;e- 


Chap.  XIX.  Man— Musical  Powers.  571 

constructed  vocal  organs,  can  be  taught  not  only  to  speak,  but  to 
pipe  or  whistle  tunes  invented  by  man,  so  tliat  they  must  have 
some  musical  capacity.  Nevertheless  it  would  be  very  rash  to 
assume  that  parrots  are  descended  from  some  ancient  form  whicli 
was  a  songster.  Many  cases  could  be  advanced  of  organs 
and  instincts  originally  adapted  for  one  purpose,  having  been 
utilised  for  some  distinct  purpose.^*'  Hence  the  capacity  for 
high  musical  development,  which  the  savage  races  of  man 
possess,  may  be  due  cither  to  the  practice  by  our  semi-human 
progenitors  of  some  rude  form  of  music,  or  simply  to  their 
having  acquired  the  proper  vocal  organs  for  a  different  purpose. 
But  in  this  latter  case  we  must  assume,  as  in  the  above  instance 
of  parrots,  and  as  seems  to  occur  with  many  animals,  that  they 
already  possessed  some  sense  of  melody. 

Music  arouses  in  us  various  emotions,  but  not  the  more 
terrible  ones  of  horror,  fear,  rage,  &c.  It  awakens  the  gentler 
feelings  of  tenderness  and  love,  which  readily  pass  into  devotion. 
In  the  Chinese  annals  it  is  said,  "Music  hath  the  power  of 
"  making  heaven  descend  upon  earth."  It  likewise  stirs  up  in  us 
the  sense  of  triumph  and  the  glorious  ardour  for  war.  "^lese 
powerful  and  mingled  feelings  may  well  give  rise  to  the  sense 
of  subHmity.  We  can  concentrate,  as  Dr.  Seemann  observes, 
greater  intensity  of  feeling  in  a  single  musical  note  than  in 
pages  of  writing.  It  is  probable  that  nearly  the  same  emotions, 
but  much  weaker  and  far  less  complex,  are  felt  by  birds  when 
the  male  pours  forth  his  full  volume  of  song,  in  rivalry  with 
other  males,  to  captivate  the  female.  Love  is  still  the  com- 
monest theme  of  our  songs.  As  Herbert  Spencer  remarks, 
"  music  arouses  dormant  sentiments  of  which  we  had  not 
"  conceived  the  possibility,  and  do  not  know  the  meaning ;  or, 
"  as  Eichter  says,  tells  us  of  things  we  have  not  seen  and*  shall 
"  not  see."  Conversely,  when  vivid  emotions  are  felt  and  ex- 
pressed by  the  orator,  or  even  in  common  speech,  musical  cadences 
and  rhythm  are  instinctively  used.  The  negro  in  Africa  when 
excited  often  bursts  forth  in  song;  "  another  will  reply  in  song, 
"  while  the  company,  as  if  touched  by  a  musical  wave,  murmur 

^®  Since  this  chapter  was  printed,  "  resulting    advantages    as  well    as 

1   have    seen    a  valuable  article  by  "limiting    disadvantages,  actual  or 

Mr.     Ghauncey     Wright     ('  North  "  possible,  which    the    principle    of 

Anier.    Review,'     Oct.     1870,    page  "  utility  may  not  have  comprehended 

293),  who,  in  discussing  the  above  "  in  its  action."  As  I  have  attempted 

subject,  remarks,  "  There   are  many  to  shew  in  an  curly  chapter  of  this 

''consequences  of  the  ultimate  laws  work,  this  principle  has  an  impor- 

"  or  uniformities  of  nature,  through  taut  bearing  on  the  acquisition  by 

"  which  the  acquisitiou  of  one  use-  man  of  some  of  his  mental  charac- 

"  ful  power  will  bring  with  it  niauy  tcristics. 


572  The  Descent  of  Man.  Part  ill. 

"  a  chorus  in  perfect  uiiison."^^  Even  monkeys  express  strong 
feelings  in  different  tones— anger  and  impatience  by  low, — fear 
and  pain  by  high  notes.^^  The  sensations  and  ideas  thus 
excited  in  us  by  music,  or  expressed  by  the  cadences  of  oratory, 
appear  from  their  vagueness,  yet  depth,  like  mental  reversions 
to  the  emotions  and  thoughts  of  a  long-past  age. 

All  these  facts  with  respect  to  music  and  impassioned  speech 
become  intelligible  to  a  certain  extent,  if  we  may  assume  that 
musical  tones  and  rhythm  were  used  by  our  half-human  an- 
cestorS;  during  the  season  of  courtship,  when  animals  of  all  kinds 
are  excited  not  only  by  love,  but  by  the  strong  passions  of 
jealousy,  rivalry,  and  triumph.  From  the  deeply-laid  principle  of 
inherited  associations,  musical  tones  in  this  case  would  be  likely 
to  call  up  vaguely  and  indefinitely  the  strong  emotions  of  a  long- 
past  age.  As  we  have  every  reason  to  suppose  that  articulate 
speech  is  one  of  the  latest,  as  it  certainly  is  the  highest,  of  the 
arts  acquired  by  man,  and  as  the  instinctive  power  of  producing 
musical  notes  and  rhythms  is  developed  low  down  in  the  animal 
series,  it  would  be  altogether  opposed  to  the  principle  of  evolution, 
if  we  were  to  admit  that  man's  musical  capacity  has  been  deve- 
lopefl  from  the  tones  used  in  impassioned  speech.  We  must 
suppose  that  the  rhythms  and  cadences  of  oratory  are  derived 
from  previously  developed  musical  powers.^^  \ye  can  thus 
understand  how  it  is  that  music,  dancing,  song,  and  poetry  are 
such  very  ancient  arts.  We  may  go  even  further  than  this, 
and,  as  remarked  in  a  former  chapter,  believe  that  musical  sounds 
afforded  one  of  the  bases  for  the  development  of  language.^" 

2'  Winwood  Reade,  'The  Martyr-  the     opposite     sex.     Thus    musical 

dom    of   Man,'    1872,    p.   441,    and  tones  became  firmly  associated  with 

*  African  Sketch  Book,'  1873,  vol.  ii.  some  ol"  the  strongest   passions  an 

p.  313*  animal  is  capable  of  feeling,  and  are 

'^  Rengger,     '  Siiugethiere     von  consequently  used  instinctively,  or 

Paraguay,'  s.  49.  through    association,   when   strong 

^^  See  the  A-ery  interesting  dis-  emotions  are  expressed  in  speech, 
cussion  on  the  '  Origin  and  Function  Mr.  Spencer  does  not  offer  any 
of  Music,'  by  Mr.  Herbert  Spencer,  satisfactory  explanation,  nor  can  I, 
in  his  collected  '  Essays,'  1858,  p.  why  high  or  deep  notes  should  be 
359.  Mr.  Spencer  comes  to  an  ex-  expressive,  both  with  man  and  the 
actly  opposite  conclusion  to  that  at  lower  animals,  of  certain  emotions, 
which  I  have  arrived.  He  con-  Mr.  Spencer  gives  also  an  in- 
cludes, as  did  Diderot  formerly,  that  teresting  discussion  on  the  relations 
the  cadences  used  in  emotional  between  poetry,  recitative,  and  song, 
speech  afford  the  foundation  from  *"  I  find  in  Lord  Monboddo's 
which  music  has  been  developed;  *  Origin  of  Language,' vol.  i.  (1774), 
whilst  I  conclude  that  musical  notes  p.  469,  that  Dr.  Blacklock  likewise 
and  rhj-thm  were  first  acquired  by  thought  "that  the  first  language 
the  male  or  female  progenitors  of  "among  men  was  music,  and  that 
mankind  for  the  sake  of  charming  "  before  our  ideas  were  expressed  by 


Ch/.p.  XIX.         Man — Love  of  Ornaiiiciit.  573 

As  the  males  of  several  quadrumanous  animals  have  their 
vocal  organs  much  more  develoiDcd  than  in  the  females,  and  as  a 
gibbon,  one  of  the  anthropomorphous  apes,  pours  forth  a  whole 
octave  of  musical  notes  and  may  be  said  to  sing,  it  appears  pro- 
bable that  the  progenitors  of  man,  cither  the  males  or  females  or 
both  sexes,  before  acquiring  the  power  of  expressing  their  mutual 
love  in  articulate  language,  endeavoured  to  charm  each  other 
with  musical  notes  and  rliythm.  So  little  is  known  about  the 
use  of  the  voice  by  the  Quadrumana  during  the  season  of  love, 
that  we  have  no  means  of  judging  whether  the  habit  of  singing 
was  fii'st  acquired  by  our  male  or  female  ancestors.  Women 
are  generally  thought  to  possess  sweeter  voices  than  men,  and  as 
far  as  this  serves  as  any  guide,  we  may  infer  that  they  first 
acquired  musical  powers  in  order  to  attract  the  other  scx.''^  But 
if  so,  this  must  have  occurred  long  ago,  before  our  ancestors  had 
become  sufficiently  human  to  treat  and  value  their  women  merely 
as  useful  slaves.  The  impassioned  orator,  bard,  or  musician, 
when  with  his  varied  tones  and  cadences  he  excites  the  strongest 
emotions  in  his  hearers,  little  suspects  that  he  uses  the  same 
means  by  which  his  half-human  ancestors  long  ago  aroused  each 
other's  ardent  passions,  during  their  courtship  and  rivah-y. 

T]\e  Influence  of  Beauty  in  determining  the  Marriages  of  Man- 
kiwi. — In  civilised  life  man  is  largely,  but  by  no  means  exclu- 
sively, influenced  in  the  choice  of  his  wife  by  external  appearance ; 
but  we  are  chiefly  concerned  with  primeval  times,  and  our  only 
means  of  forming  a  judgment  on  this  subject  is  to  study  the 
habits  of  existing  semi-civilised  and  savage  nations.  If  it  can  be 
shewn  that  the  men  of  different  races  prefer  women  having 
various  characteristics,  or  conversely  with  the  women,  we  have 
then  to  enquire  whether  such  choice,  continued  during  many 
generations,  would  produce  any  sensible  effect  on  the  race, 
either  on  one  sex  or  both  according  to  the  form  of  inheritance 
which  has  prevailed. 

It  will  be  well  first  to  shew  in  some  detail  that  savages  pay  the 
greatest  attention  to  their  i3ersonal  apj^earance.^^   That  they  have 

"articulate  sounds,  they  were  com-  selves,  is  given  by  the  Italian  tra- 

"  municated  by  tones,  A^aried  accord-  Atelier,  Prof.  Mantegazza,  '  Kio  de  la 

"  ing  to  difierent  degrees  of  gravity  Phita,  Viaggi    e    Studi,'    18G7,    pp. 

"  and  acuteuess."  525-545  ;  all    the  following    state- 

■*'  See    an    interesting    discussion  ments,  when  other    references   are 

on  this  subject  by  Hiickei,  'Gene-  not  given,  are  taken  from  this  work, 

relle  Morph,' B.  ii.  1866,  s.  246.  See,     also,    Waltz,     '  Introduct.     to 

^2  A  full  and  excellent  account  of  Anthropolog.'    Kng.    transl.    vol.    i. 

the  manner  in  which  savages   in  all  1863,  p.  275,  ct  passim.     Lawrence 

parts  of  the  world  ornament  them-  also  gives  very  full    details  in  his 


574  The  Descent  of  Man.  Part  III. 

a  passion  for  ornament  is  notorious ;  and  an  English  philosopher 
goes  so  far  as  to  maintain,  that  clothes  were  first  made  for 
ornament  and  not  for  warmth.  As  Professor  Waitz  remarks, 
"  however  poor  and  miserable  man  is,  he  finds  a  pleasui-e  in 
*'  adorning  himself."  The  extravagance  of  the  naked  Indians  of 
South  America  in  decorating  themselves  is  shewn  "  by  a  man  of 
"  large  stature  gaining  with  difficulty  enough  by  the  labour  of  a 
"  fortnight  to  procure  in  exchange  the  cliica  necessary  to  jDaint 
"  himself  red."  ^^  The  ancient  barbarians  of  Europe  during  the 
Eeindeer  period  brought  to  their  caves  any  brilliant  or  singular 
objects  which  they  happened  to  find.  Savages  at  the  present 
day  everywhere  deck  themselves  with  plumes,  necklaces,  armlets, 
ear-rings,  &c.  They  i3aint  themselves  in  the  most  diversified 
manner.  "  If  painted  nations,"  as  Humboldt  observes,  "  had 
"  been  examined  with  the  same  attention  as  clothed  nations,  it 
"  would  have  been  perceived  that  the  most  fertile  imagination 
"  and  the  most  mutable  caprice  have  created  the  fashions  of 
"  painting,  as  well  as  those  of  garments." 

In  one  part  of  Africa  the  eyelids  are  coloured  black ;  in  another 
the  nails  are  coloured  yellow  or  purple.  In  many  places  the 
hair  is  dyed  of  various  tints.  In  different  countries  the  teeth  are 
stained  black,  red,  blue,  &c.,  and  in  the  Malay  Archipelago  it  is 
thought  shameful  to  have  white  teeth  ''like  those  of  a  dog."  Not 
one  great  country  can  be  named,  from  the  Polar  regions  in  the 
north  to  New  Zealand  in  the  south,  in  which  the  aborigines  do 
not  tati.oo  themselves.  This  practice  was  followed  by  the  Jews 
of  old,  and  by  the  ancient  Britons.  In  Africa  some  of  the 
natives  tattoo  themselves,  but  it  is  a  much  more  common 
practice  to  raise  protuberances  by  rubbing  salt  into  incisions 
made  in  various  parts  of  the  body ;  and  these  are  considered  by 
the  inhabitants  of  Kordofau  and  Darfur  "  to  be  great  piersonal 
"  attractions."  In  the  Arab  countries  no  beauty  can  be  perfect 
until  the  checl?s  "  or  temples  have  been  gashed."  ^^  In  South 
America,  as  Humboldt  remarks,  "  a  mother  would  be  accused  of 
"  culpable  indifference  towards  her  children,  if  she  did  not 
"  employ  artificial  means  to  shape  the  calf  of  the  leg  after  the 
"  fashion  of  the  country."    In  the  Old  and  New  Worlds  the 


'Lectures    on     Physiology,'     1822.  *^  Humboldt, 'Personal  Narrative,' 

Since  this  cliapter  was  written  Sir  Eng.  Trauslat.  vol.   iv.  p.  515;  on 

J.  Lubbock  has  published  his  '  Origin  the  imagination  shewn  in  painting 

of  Civilisation,'  1870,  in  which  theie  the  body,  p.  522  ;  on  modifying  the 

is  an    interesting    cliapter    on    the  form  of  the  calf  of  the  leg,  p.  466. 
I)resent  subject,  and  from  which  (j)]).  ■*•*  '  The  Nile  Tributaries,'  1867  ; 

42,  48)    I    have    taken   some    facts  '  The  Albert  N'yanza,'  1866,  vol.  i. 

about   savages   dyeing     their    teeth  p.  218. 
and  hair,  and  piercing  their  teeth. 


CiiAp.  XIX.  Man — Love  of  Ornament.  575 

shape  of  the  skull  was  formerly  modified  during  infancy  in  the 
most  extraordinary  manner,  as  is  still  the  case  in  many  places, 
and  such  deformities  are  considered  ornamental.  For  instance, 
the  savages  of  Colombia^^  deem  a  much  flattened  head  "an 
"  essential  point  of  beauty." 

The  hair  is  treated  with  especial  care  in  various  countries  ;  it 
is  allowed  to  grow  to  full  length,  so  as  to  reach  to  the  ground,  or 
is  combed  into  *'  a  compact  frizzled  mop,  which  is  the  Papuan's 
"pride  and  glory."'***  In  Northern  Africa  "a  man  requires  a 
"  period  of  from  eight  to  ten  years  to  perfect  his  coiffure."  With 
other  nations  the  head  is  shaved,  and  in  parts  of  South  America 
and  Africa  even  the  eyebrows  and  eyelashes  are  eradicated.  The 
natives  of  the  Upper  Nile  knock  out  the  four  front  teeth,  saying 
that  they  do  not  wish  to  resemble  brutes.  Further  south,  the 
Batokas  knock  out  only  the  two  upper  incisors,  which,  as 
Livingstone*^  remarks,  gives  the  face  a  hideous  appearance, 
owing  to  the  prominence  of  the  lower  jaw;  but  these  people  think 
the  presence  of  the  incisors  most  unsightly,  and  on  beholding 
some  Europeans,  cried  out,  "  Look  at  the  great  teeth ! "  The 
chief  Sebituani  tried  in  vain  to  alter  this  fashion.  In  various 
parts  of  Africa  and  in  the  Malay  Archipelago  the  natives  file  the 
incisors  into  points  like  those  of  a  saw,  or  pierce  them  with 
holes,  into  which  they  insert  studs. 

As  the  face  with  us  is  chiefly  admired  for  its  beauty,  so  with 
savages  it  is  the  chief  seat  of  mutilation.  In  all  quarters  of  the 
world  the  septum,  and  more  rarely  the  wings  of  the  nose  are 
pierced ;  rings,  sticks,  feathers,  and  othei  ornaments  being  in- 
serted into  the  holes.  The  ears  are  everywhere  pierced  and 
similarly  ornamented,  and  with  the  Botocudos  and  Lenguas  of 
South  America  the  hole  is  gradually  so  much  enlarged  that  the 
lower  edge  touches  the  shoulder.  In  North  and  South  America 
and  in  Africa  either  the  upper  or  lower  lip  is  pierced ;  and  with 
the  Botocudos  the  hole  in  the  lower  lip  is  so  large  that  a  disc  of 
wood,  four  inches  in  diameter,  is  placed  in  it.  Mantegazza  gives 
a  curious  account  of  the  shame  felt  by  a  South  American  native, 
and  of  the  ridicule  which  he  excited,  when  he  sold  his  tcmheta, — 
the  large  coloured  piece  of  wood  which  is  passed  through  the 
hole.  In  Central  Africa  the  women  perforate  the  lower  lip  and 
wear  a  crystal,  which,  from  the  movement  of  the  tongue,  has  "  a 
"  wriggling  motion,  indescribably  ludicrous  during  conversation," 

**  Quoted     by    Prichard,    '  Phys.  445.     On  the  coiffure  of  the  Afii- 

Ilist.  of  Mankind,'  4th  edit.  vol.  i.  cans,    Sir    S.    Baker,    'The    Albert 

1851,  p.  321.  N'yanza,'  vol.  i.  p.  210. 

<«  On     the     Papuans,     WaUace,  "' Travels,' p.  533. 
*  The  Malay  Arcliipolago,'  v(-l.  ii.  p. 


5/6  TJie  Descent  of  Man.  Part  111. 


(( ( 


The  wife  of  the  chief  of  Latooka  told  Sir  S.  Baker  "*^  that  Lady 
Baker  "  would  be  much  improved  if  she  would  extract  her  four 
"  front  teeth  from  the  lower  jaw,  and  wear  the  long  pointed 
*'  polished  crystal  in  her  under  lip."  Further  south  with  the 
Makalolo,  the  upper  lip  is  perforated,  and  a  large  metal  and 
bamboo  ring,  called  a  pehle,  is  worn  in  the  hole.  "  This  caused 
"  the  lip  in  one  case  to  project  two  inches  beyond  the  tip  of  the 
"  nose;  and  when  the  lady  smiled  the  contraction  of  the  muscles 
"  elevated  it  over  the  eyes.  '  Why  do  the  women  wear  these 
"  '  things^? '  the  venerable  chief,  Chinsurdi,  was  asked.  Evidently 
"  surprised  at  such  a  stuj^id  question,  he  replied, '  For  beauty  ! 
"  '  They  are  the  only  beautiful  things  women  have  ;  men  have 
beards,  women  have  none.  What  kind  of  a  person  would  she 
be  without  the  pelele  ?  She  would  not  be  a  woman  at  all 
** '  with  a  mouth  like  a  man,  but  no  beard.'  "^^ 

Hardly  any  part  of  the  body,  which  can  be  unnaturally 
modified,  has  escaped.  The  amount  of  suffering  thus  caused 
must  have  been  extreme,  for  many  of  the  operations  require 
several  years  for  theii'  completion,  so  that  the  idea  of  their 
necessity  must  be  imperative.  The  motives  are  various;  the 
men  paint  their  bodies  to  make  themselves  appear  terrible  in 
battle;  certain  mutilations  are  connected  with  religious  rites, 
or  they  mark  the  age  of  puberty,  or  the  rank  of  the  man,  or  they 
serve  to  distinguish  the  tribes.  Amongst  savages  the  same 
fashions  prevail  for  long  periods,^°  and  thus  mutilations,  from 
whatever  cause  first  made,  soon  come  to  be  valued  as  distinctive 
marks.  But  self-adornment,  vanity,  and  the  admiration  of 
others,  seem  to  be  the  commonest  motives.  In  regard  to  tattoo- 
ing, I  was  told  by  the  missionaries  in  New  Zealand,  that  when 
they  tried  to  persuade  some  girls  to  give  up  the  practice,  they 
answered, ''  We  must  just  have  a  few  lines  on  our  lips  ;  else  when 
"  we  grow  old  we  shall  be  so  very  ugly."  With  the  men  of  New 
Zealand,  a  most  capable  judge  ^^  says,  "  to  have  fine  tattooed  faces 
"  was  the  great  ambition  of  the  young,  both  to  render  themselves 
"  attractive  to  the  ladies,  and  conspicuous  in  war."  A  star 
tattooed  on  the  forehead  and  a  spot  on  the  chin  are  thought  by 
the  women  in  one  part  of  Africa  to  be  irresistible  attractions.^- 

••*  'The    Albert   K'yanza,'    ISGG,  "fashion    for    dr'^ssing    the    hair." 

vol.  i.  p.  217.  See  Agassiz    ('Journey    in    Brazil,' 

^^  Livingstone,  'British  Associa-  18G8,  p.  318)  on  the    invariability 

tion,'    18GU  ;    report    given    in    the  of  the  tattooing  of  the  Amazonian 

'  Athena'iim,'  July  7,  ISGO,  p.  29.  Indians. 

*"  Sir  S.   Bahor  (ibid.  vol.   i.  p.  ^'   Kev.  R.  Taylor,  '  New  Zealand 

210)   speaking    of    the     natives    of  and  its  Inhabitants,'  1855,  p.  152. 
Central    Africa   says,  "every   tribe  ^-  Mantegazza,  '  Viaggi  c  Studi,' 

"  has   a    dibtinct     and     unchanging  p.  542. 


Chap.  XIX.  Man — Beauty.  577 

In  most,  but  not  all  parts  of  the  world,  tlio  men  arc  more 
ornamented  than  the  women,  and  often  in  a  different  manner ; 
sometimes,  though  rarely,  the  women  are  hardly  at  all  orna- 
mented. As  the  ^romen  are  made  by  savages  to  perform  the 
greatest  share  of  the  work,  and  as  they  are  not  allowed  to  eat 
the  best  kinds  of  food,  so  it  accords  with  the  characteristic 
selfishness  of  man  that  they  should  not  be  allowed  to  obtain,  or 
use  the  finest  ornaments.  Lastly,  it  is  a  remarkable  fact,  as 
proved  by  the  foregoing  quotations,  that  the  same  fashions  in 
modifying  the  shape  of  the  head,  in  ornamenting  the  hair,  in 
painting,  tattooing,  in  perforating  the  nose,  lips,  or  ears,  in 
removing  or  filing  the  teeth,  &c.,  now  prevail,  and  have  long 
prevailed,  in  the  most  distant  quarters  of  the  world.  It  is 
extremely  improbable  that  these  practices,  followed  by  so  many 
distinct  nations,  should  be  due  to  tradition  from  auy  common 
source.  They  indicate  the  close  similarity  of  the  mind  of  man, 
to  whatever  race  he  may  belong,  just  as  do  the  almost  iiniversal 
habits  of  dancing,  masquerading,  and  making  rude  pictures. 

Having  made  these  preliminary  remarks  on  the  admiration 
felt  by  savages  for  various  ornaments,  and  for  deformities  most 
unsightly  in  our  eyes,  let  us  see  how  far  the  men  are  attracted 
by  the  appearance  of  their  women,  and  what  are  theii'  ideas  of 
beauty.  I  have  heard  it  maintained  that  savages  are  quite 
indifferent  about  the  beauty  of  their  women,  valuing  them  solely 
as  slaves  ;  it  may  therefore  be  well  to  observe  tlw,t  this  conclusion 
does  not  at  all  agree  with  the  care  which  the  women  take  in 
ornamenting  themselves,  or  with  their  vanity.  BurchelP^  gives 
an  amusing  account  of  a  Bush-w^oman  who  used  as  much 
grease,  red  ochre,  and  shining  powder  "  as  would  have  ruined 
"  any  but  a  very  rich  husband."  She  displayed  also  "  much 
"  vanity  and  too  evident  a  consciousness  of  her  superiority."  Mr. 
Winwood  Eeade  informs  me  that  the  negroes  of  the  West  Coast 
often  discuss  the  beauty  of  their  women.  Some  competent 
observers  have  attributed  the  fearfully  common  practice  of 
infanticide  partly  to  the  desire  felt  by  the  women  to  retain  their 
good  looks.^*  In  several  regions  the  women  wear  charms  and 
use  love-philters  to  gain  the  affections  of  the  men;  and  Mr. 
Brown  enumerates  four  plants  used  for  this  purpose  by  the 
women  of  North-Western  America/^ 

"  'Travels    in  S.   Africa,'   1824,  'Voyages,'  &c.  torn.  ii.  p.  IIG. 

vol.  i.  p.  41-1:,  *^  Oa  the  vegetable    productions 

^*  See,    for    references,    Gerlaud  used  by  tlie  North-Western  Anieri- 

•  Ueber  das  Aussterbeuder  Naturvol-  can  hidiaus,  'Pharmaceutical  Jour- 

ker,'  18G8,  s.  51,  o.'},  hh  ;  also  Azara,  nal,'  vol.  x. 


578  The  Descent  of  Man.  Part  111. 

Hearne/^  an  excellent  observer,  who  lived  many  years  with  the 
American  Indians,  says,  in  speaking  of  the  women,  "Ask  a 
"  Northern  Indian  what  is  beauty,  and  he  will  answer,  a  broad 
"  flat  face,  small  eyes,  high  cheek-bones,  t!iree  or  four  broad 
"  black  lines  across  each  cheek,  a  low  forehead,  a  large  broad 
"  cliin,  a  clumsy  hook  nose,  a  tawny  hide,  and  breasts  hanging 
"  down  to  the  belt."  Pallas,  who  visited  the  northern  parts  of 
the  Chinese  empire,  says  *'  those  women  are  preferred  who  have 
"  the  Mandschu  form ;  that  is  to  say,  a  broad  face,  high  cheek- 
"  bones,  very  broad  noses,  and  enormous  ears ; "  ^'^  and  Vogt 
remarks  that  the  obliquity  of  the  eye,  which  is  proper  to  the 
Chinese  and  Japanese,  is  exaggerated  in  their  pictures  for  the 
purpose,  as  it  "  seems,  of  exhibiting  its  beauty,  as  contrasted 
"  with  the  eye  of  the  red-haired  barbarians."  It  is  well  known, 
as  Hue  repeatedly  remarks,  that  the  Chinese  of  the  interior 
think  Europeans  hideous,  with  their  white  skins  and  prominent 
noses.  The  nose  is  far  from  being  too  prominent,  according  to  our 
ideas,  in  the  natives  of  Ceylon ;  yet  "  the  Chinese  in  the  seventh 
"  century,  accustomed  to  the  flat  features  of  the  Mongol  races, 
"■  were  surprised  at  the  prominent  noses  of  the  Cingalese ;  and 
*'  Thsang  described  them  as  having  '  the  beak  of  a  bird,  with  the 
"  '  body  of  a  man.' " 

Finlayson,  after  minutely  describing  the  people  of  Cochin 
China,  says  that  their  rounded  heads  and  faces  are  their  chief 
characteristics ;  and,  he  adds,  "  the  roundness  of  the  whole 
"  countenance  is  more  striking  in  the  women,  who  are  reckoned 
"  beautiful  in  proportion  as  they  display  this  form  of  face."  The 
Siamese  have  small  noses  with  divergent  nostrils,  a  wide  mouth, 
rather  thick  lips,  a  remarkably  large  face,  with  very  high  and 
broad  cheek-bones.  It  is,  therefore,  not  wonderful  that "  beauty, 
"  according  to  our  notion  is  a  stranger  to  them.  Yet  they  con- 
"  sider  their  own  females  to  be  much  more  beautiful  than  those 
"  of  Europe."^^ 

It  is  well  known  that  with  many  Hottentot  women  the 
posterior  part  of  the  body  projects  in  a  wonderful  manner ;  they 
are  steatopygous ;  and  Sir  Andrew  Smith  is  certain  that  this 
peculiarity  is  greatly  admired  by  the  men.^^    He  once  saw  a 

^^  '  A    Journey    from     Prince    of  furd  and  Finlayson,  *  Phys.  Hist,  of 

Wales  Fort,'  8vo.  edit.  179G,  p.  89.  Mankind,'  vol.  iv.  pp.  534,  535. 

*^  Quoted     by    Prichard,    'Phys.  ^^  Idem  illustrissimus  viator  dixit 

Hist,  of  Mankind,'  3rd  edit.  vol.  iv.  mihi    proecinctorium    A'el    tabulam 

1844,  p.  519;  Vogt,    'Lectures   on  foemina?,  quod  nobis  teterrimum  est, 

Man,'    Eng.    translat.    p.    129.     On  quondam    permagno     aestimari    ab 

the  opinion  of   the  Chinese  on   the  hominibus  in  hac  gonte.     Nunc  res 

Cingalese, E.Tennent, 'Ceylon,' 1859,  mutata  est,  et  censent  talem    con- 

vol.  ii.  p.  107.  formationem  miaime  optandam  esse. 

*^  Prichard   as  taken  from  Crp- 


Chap.  XIX.  Alan — Beauty.  579 

woman  who  was  considered  a  beauty,  and  she  was  so  immensely 
developed  behind,  that  when  seated  on  level  ground  she  could 
not  rise,  and  had  to  push  herself  along  until  she  came  to  a  slope. 
Some  of  the  women  in  various  negro  tribes  have  the  same  pecu- 
liarity ;  and,  according  to  Burton,  the  Somal  men  "  are  said  to 
"  choose  their  wives  by  ranging  them  in  a  line,  and  by  picking 
"  her  out  who  projects  farthest  a  tergo.  Nothing  can  be  more 
"  hateful  to  a  negro  than  the  opposite  form."  ^ 

With  respect  to  colour,  the  negroes  rallied  Mungo  Park  on  the 
whiteness  of  his  skin  and  the  prominence  of  his  nose,  both  of 
which  they  considered  as  "  unsightly  and  unnatural  conforma- 
"  tions."  He  in  return  praised  the  glossy  jet  of  their  skins  and  the 
lovely  depression  of  their  noses ;  this  they  said  was,  '*  honey- 
"  mouth,"  nevertheless  they  gave  him  food.  The  African  Moors, 
also,  "  knitted  their  brows  and  seemed  to  shudder  "  at  the  white- 
ness of  his  skin.  On  the  eastern  coast,  the  negro  boys  when  they 
saw  Burton,  cried  out  "  Look  at  the  white  man ;  does  he  not  look 
*'  like  a  white  ape  ?  "  On  the  western  coast,  as  Mr.  Win  wood 
Reade  informs  me,  the  negroes  admire  a  very  black  skin  more 
than  one  of  a  lighter  tint.  But  their  horror  of  whiteness  may  be 
attributed,  according  to  this  same  traveller,  partly  to  the  belief 
held  by  most  negroes  that  demons  and  spirits  are  white,  and 
partly  to  their  thinking  it  a  sign  of  ill-health. 

The  Banyai  of  the  more  southern  part  of  the  continent  arc 
negroes,  but "  a  great  many  of  them  are  of  a  light  coffee-and-milk 
"  colour,  and,  indeed,  this  colour  is  considered  handsome 
"throughout  the  whole  country;"  so  that  here  we  have  a 
different  standard  of  taste.  With  the  Kafirs,  who  differ  much 
from  negroes,  "  the  skin,  except  among  the  tribes  near  Delagoa 
"  Bay,  is  not  usually  black,  the  prevailing  colour  being  a  mixture 
''of  black  and  red,  the  most  common  shade  being  chocolate. 
"  Dark  complexions,  as  being  most  common  are  naturally  held  in 
"  the  highest  esteem.  To  be  told  that  he  is  light-coloured,  or 
"  like  a  white  man,  would  be  deemed  a  very  poor  compliment  bv 
"  a  Kafir.  I  have  heard  of  one  imfortunate  man  who  was  so  very 
"  fair  that  no  girl  would  marry  him."  One  of  the  titles  of  the 
Zulu  king  is  "  You  who  are  black."  ^^  Mr.  Galton,  in  speaking 
to  me  about  the  natives  of  S.  Africa,  remarked  that  their  ideas  of 

«»  '  The  Anthropological  Review,'  Burton's    statement    is    quoted    by 

November,  1864,  p.  237.     For  ad-  Schaaffhausen,  '  Archiv  fiir  Anthro- 

ditional  references,  see  Waitz,  '  In-  polog.' 18GG,  s.  1G;5,    On  the  Banyai, 

troduct.     to    Anthropology,'    Eng.  Livingstone,    'Travels,'  p.  G4.     On 

translat.  18G3,  vol.  i.  p.  105.  the    Kafirs,    the    Rev.    J.    Shooter, 

«i  Mungo     Park's     'Travels     in  'The  Kafirs  of  Natal  and  the  Zulu 

Africa,'   4to.    1816,    pp.    53,     131.  Country,'  1857,  p.  1. 


580  The  Descent  of  Man.  Tart  III. 

beauty  seem  very  different  from  ours ;.  for  in  one  tribe  two  slim, 
slight,  and  pretty  girls  were  not  admired  by  the  natives. 

Turning  to  other  quarters  of  the  world ;  in  Java,  a  yellow,  not 
a  white  girl,  is  considered,  according  to  Madame  Pfeiffer,  a 
beauty.  A  man  of  Cochin  China  "  spoke  with  contempt  of  the 
"  wife  of  the  English  Ambassador,  that  she  had  wliite  teeth  like 
"  a  dog,  and  a  rosy  colour  like  that  of  potato-flowers."  We 
have  seen  that  the  Chinese  dislike  our  white  skin,  and  that  the 
N.  Americans  admire  "  a  tawny  hide."  In  S.  America,  the  Yura- 
caras,  who  inhabit  the  wooded,  damp  sIojdcs  of  the  eastern 
Cordillera,  are  remarkably  pale-coloured,  as  their  name  in  their 
own  language  expresses ;  nevertheless  they  consider  European 
women  as  very  inferior  to  their  own.*^^ 

In  several  of  the  tribes  of  North  America  the  hair  on  the  head 
grows  to  a  wonderful  length ;  and  Cathn  gives  a  curious  proof 
how  much  this  is  esteemed,  for  the  chief  of  the  Crows  was 
elected  to  this  oflice  from  having  the  longest  hair  of  any  man  in 
the  tribe,  namely  ten  feet  and  seven  inches.  The  Aymaras  and 
Quichuas  of  S.  America,  likewise  have  very  long  hair ;  and  this, 
as  Mr.  D.  Eorbes  informs  me,  is  so  much  valued  as  a  beauty, 
that  cutting  it  off  was  the  severest  punishment  which  he  could 
inflict  on  them.  In  both  the  Northern  and  Southern  halves  of 
the  continent  the  natives  sometimes  increase  the  ai)parent  length 
of  their  hair  by  weaving  into  it  fibrous  substances.  Although 
the  hair  on  the  head  is  thus  cherished,  that  on  the  face  is  con- 
sidered, by  the  North  American  Indians  "•  as  very  vulgar,"  and 
every  hair  is  carefully  eradicated.  This  practice  jDrevails 
throughout  the  American  continent  from  Yancouvers  Island  in 
the  north  to  Tierra  del  Fuego  in  the  south.  When  York 
Minster,  a  Fuegian  on  board  the  'Beagle,'  was  taken  back  to 
his  country,  the  natives  told  him  he  ought  to  i)ull  out  the  few 
short  hau's  on  his  face.  They  also  threatened  a  young  missionary, 
who  was  left  for  a  time  with  them,  to  strip  him  naked,  and 
pluck  the  hairs  from  his  face  and  body,  yet  he  was  far  from  being 
a  hairy  man.  This  fashion  is  carried  so  far  that  the  Indians  of 
Paraguay  eradicate  their  eyebrows  and  eyelashes,  saying  that 
ihey  do  not  wish  to  be  like  horses.''^ 

«2  For    the    Javaus   and    Cochin-  G.  Catlin,  3rd  edit.  1842,  vol.  i.  p. 

Chinese,  see  Waltz,    'Introduct.  to  49;  vol.  ii.  p.  227.     On  the  natives 

Anthropology,'  Eng.  transUit.  vol.  i.  of    Vancouver's  Island,  see   Sproat, 

p.    oO.j.     On    the    Yura-caras,    A.  '  Scenes  and  Studies  of  Savage  Life,* 

(I'Orligny,  as   quoted    in    Prichard,  18()8,    p.    25.     On    the    Indians    of 

'  I'hys.  Hist,  of  Mankind,'  vol.  v.  3rd  I'araguay,  Azara,  '  Voyages,'  torn.  ii. 

edit.  p.  470.  p.  1U5. 

^^  'North  American  Indians,'  by 


CiiAi'.  XIX.  Man— Beauty.  581 

It  is  remarkable  that  throughout  the  world  the  races  which 
are  almost  completely  destitute  of  a  beard,  dislike  hairs  on  the 
face  and  body,  and  take  pains  to  eradicate  them.  The  Kalmucks 
are  beardless,  and  they  are  well  known,  like  the  Americans,  to 
pluck  out  all  straggling  hairs ;  and  so  it  is  with  the  Polynesians, 
some  of  the  Malays,  and  the  Siamese.  Mr.  Veitch  states  that  the 
Japanese  ladies  **  all  objected  to  our  whiskers,  considering  them 
"  very  ugly,  and  told  us  to  cut  them  off,  and  be  like  Japanese 
"  men."  The  New  Zealandors  have  short,  curled  beards ;  yet 
they  formerly  plucked  out  the  hairs  on  the  face.  They  had  a 
saying  that  "  there  is  no  woman  for  a  hairy  man  ;"  but  it  would 
appear  that  the  fashion  has  changed  in  New  Zealand,  perhaps 
owing  to  the  presence  of  Europeans,  and  I  am  assured  that 
beards  are  iiow  admired  by  the  Maorics.*'* 

On  the  other  hand,  bearded  races  admire  and  greatly  valne 
their  beards ;  among  the  Anglo-Saxons  every  part  of  the  body 
had  a  recognised  value  ;  "  the  loss  of  the  beard  being  estimated 
"  at  twenty  shillings,  while  the  breaking  of  a  thigh  was  fixed  at 
"  only  twelve."  "^^  In  the  East  men  swear  solemnly  by  their 
beards.  We  have  seen  that  Chinsurdi,  the  chief  of  the  Makalolo 
in  Africa,  thought  that  beards  were  a  great  ornament.  In  the 
Pacific  the  Fijian's  beard  is  '*  profuse  and  bushy,  and  is  his 
"  greatest  pride ; "  whilst  the  inhabitants  of  the  adjacent  archi- 
pelagoes of  Tonga  and  Samoa  are."  beardless,  and  abhor  a  rough 
"  chin."  In  one  island  alone  of  the  Ellice  group  "  the  men  are 
"  heavily  bearded,  and  not  a  little  proud  thereof."  ^^ 

We  thus  see  how  widely  the  different  races  of  man  differ  in 
their  taste  for  the  beautiful.  In  every  nation  sufficiently  ad- 
vanced to  have  made  effigies  of  their  gods  or  of  their  deified 
rulers,  the  sculptors  no  doubt  have  endeavoured  to  express  their 
highest  ideal  of  beauty  and  grandeur."  Under  this  point  of  view 
it  is  well  to  compare  in  our  mind  the  Jupiter  or  Apollo  of  the 
Greeks  with  the  Egyptian  or  Assyrian  statues ;  and  these  with 
the  hideous  bas-reliefs  on  the  ruined  buildings  of  Central 
America. 

I  have  met  with  very  few  statements  ojiposed  to  this  conclusion. 

"  On  the  Siamese,  Prichard,  ibid.  tion,'  1870,  p.  321. 
vol.   iv.  p.  533.     On  the  Japanese,  ""  Dr.  Barnard  Davis  quotes  Mr. 

Veitch    in    '  Gardeners'    Chronicle,'  Prichard   and  others  for  these  facts 

1860,  p.   1104.     On   the  Kew  Zea-  in    regard    to    the     Polynesians,    in 

landers.     Mantegazza,     '  Viaggi     e  '  Anthropological     Ileviow,'     April, 

Studi,'  18G7,  p.  5213.     For  the  other  1870,  p.  185,  191. 
nations  mentioned,  see  references  in  ^^  Ch.  Conite  has  remarks  to  tliis 

Lawrence,  'Lectures  on  Physiology,'  effect  in  his  '  Trait«  de  Legislation,' 

&c.  1822,  p.  272.  3rd  edit.  1837,  p.  13G. 

**  Lubbock,   'Origin  of  Civilisa- 


5  82  The  Descent  of  Man.  Part  III. 

Mr.  WinwoocI  Eeade,  however,  -who  has  had  ample  oppor- 
tunities for  observation,  not  only  with  the  negroes  of  the  West 
Coast  of  Africa,  but  with  those  of  the  interior  who  have  never 
associated  with  Europeans,  is  convinced  that  their  ideas  of 
beauty  are  on  the  luliole  the  same  as  ours ;  and  Dr.  Eohlfs  writes 
to  me  to  the  same  effect  with  respect  to  Bomu  and  the  countries 
inhabited  by  the  Pullo  tribes.  Mr.  Pieade  found  that  he  agreed 
with  the  negroes  in  their  estimation  of  the  beauty  of  the  native 
girls ;  and  that  iheir  appreciation  of  the  beauty  of  European 
women  corresponded  with  ours.  They  admire  long  hair,  and 
use  artificial  mtans  to  make  it  appeal'  abundant ;  they  admire 
also  a  beard,  though  themselves  very  scantily  provided.  Mr. 
Eeade  feels  doubtful  what  kind  of  nose  is  most  appreciated  :  a 
girl  has  been  heard  to  say,"  I  do  not  want  to  man-y  him,  he  has 
"  got  no  nose ;"  and  this  shews  that  a  very  flat  nose  is  not  admired. 
We  should,  however,  bear  in  mind  that  the  depressed,  broad 
noses  and  projecting  jaws  of  the  negroes  of  the  West  Coast  are 
exceptional  types  with  the  inhabitants  of  Africa.  Notwithstand- 
ing the  foregoing  statements,  Mr.  Eeade  admits  that  negToes 
"  do  not  like  the  colour  of  our  skin ;  they  look  on  blue  eyes  with 
"  aversion,  and  they  think  our  noses  too  long  and  our  lips  too 
'•'  thin."  He  docs  not  think  it  probable  that  negroes  would  ever 
prefer  the  most  beautiful  European  woman,  on  the  mere  grounds 
of  physical  admiration,  to  a  good-looking  negress.*^^ 

The  general  truth  of  the  principle,  long  ago  insisted  on  by 
Humboldt,*^^  that  man  admires  and  often  tries  to  exaggerate 
whatever  characters  nature  may  have  given  him,  is  shewn  in 
many  ways.  The  practice  of  beardless  races  extirpating  every 
trace  of  a  beard,  and  often  all  the  hairs  on  the  body,  affords  one 
illustration.  The  skull  has  been  greatly  modified  during  ancient 
and  modern  times  by  many  nations;  and  there  can  be  little 
doubt  that  this  has  been  practised,  especially  in  N.  and  S. 
America,  in  order  to  exaggerate  some  natural  and  admired 

"^  The    'African    Sketch    Book,'  consider    us  as  superior  beings.     I 

vol.  ii.   187o,    pp.    253,    394,    5'21.  should  add  that  a  most  experienced 

The    Fuegians,   as   I  have  been    in-  ob&erver,     Capt.     Burton,     believes 

formed  by   a  missionary  who  long  that   a   woman  whom  we    consider 

resided   with  them,  consider  Euro-  beautiful  is  admired  throughout  the 

pean  women  as  extremely  beautiful ;  world,     'Anthropological     Review,' 

but  from  what  we  have  seen  of  the  March,  1864,  p.  245. 

judgment  of  the  other  aborigines  of  ^"  '  Personal      Narrative,'      Eng. 

America,  I  cannot   but  think    that  translat.  vol.  iv.  p.   518,  and  else- 

this  must  be  a  mistake,  unless  in-  where.    Mantegazza,  in  his  '  Viaggi 

deed  the  statement  refers  to  the  few  e  Studi,'  1867,  strongly  insists   on 

Fuegians  who   have  lived  for  some  this  same  principle, 
time  with  Europeans,  and  who  must 


CnAi'.  XIX.  Man^ Beauty.    .  583 

peculiarity.  Many  American  Indians  arc  known  to  admire  a  head 
so  extremely  flattened  as  to  appear  to  us  idiotic.  The  natives  on 
the  north-western  coast  compress  the  head  into  a  pointed  cone  ; 
and  it  is  their  constant  practice  to  gather  the  hair  into  a  knot  on 
the  top  of  the  head,  for  the  sake,  as  Dr.  Wilson  remarks,  "  of 
*'  increasing  the  apparent  elevation  of  the  favourite  conoid  form." 
The  inhabitants  of  Arakhan  "  admire  a  broad,  smooth  forehead, 
"  and  in  order  to  produce  it,  they  fasten  a  plate  of  lead  on  the 
"  heads  of  the  new-born  children."  On  the  other  hand,  "  a  broad, 
*'  well-rounded  occiput  is  considered  a  great  beauty  "  by  the 
natives  of  the  Fiji  islands.'^" 

As  with  the  skull,  so  with  the  nose ;  the  ancient  Huns  during 
the  age  of  Attila  were  accustomed  to  flatten  the  noses  of  their 
infants  with  bandages,  "  for  the  sake  of  exaggerating  a  natural 
"  conformation.'*'  With  the  Tahitians,  to  be  called  long-nose  is 
considered  as  an  insult,  and  they  compress  the  noses  and  fore- 
heads of  their  children  for  the  sake  of  beauty.  The  same  holds 
with  the  Malays  of  Sumatra,  the  Hottentots,  certain  Negroes, 
and  the  natives  of  Brazil.'^^  The  Chinese  have  by  nature  un- 
usually small  feet ;  "'^  and  it  is  well  known  that  the  women  of 
the  upiDcr  classes  distort  their  feet  to  make  them  still  smaller. 
Lastly,  Humboldt  thinks  that  the  American  Indians  prefer 
colouring  their  bodies  with  red  paint  in  order  to  exaggerate 
their  natural  tint ;  and  until  recently  European  women  added 
to  their  naturally  bright  colours  by  rouge  and  white  cosmetics  ; 
but  it  may  be  doubted  wiiether  barbarous  nations  have  generally 
had  any  such  intention  in  painting  themselves. 

In  the  fashions  of  our  own  dress  we  see  exactly  the  same 
principle  and  the  same  desire  to  carry  every  point  to  an  extreme ; 
we  exhibit,  also,  the  same  spirit  of  emulation.  But  the  fashions 
of  savages  are  far  more  permanent  than  ours ;  and  whenever 
their  bodies  are  artificially  modified,  this  is  necessarily  the  case. 
The  Arab  W'Omen  of  the  Upper  Nile  occupy  about  three  days  in 
dressing  their  hair ;  they  never  imitate  other  tribes, "  but  simply 

■o  On  the  skulls  of  the  American  "  On    the    Huns,    Gotlron,    <  De 

tribes,  see  Nott  and  Gliddon,' Types  I'Espece,'   torn.    ii.     1859,    p.    300. 

of  Mankind,' 1854,  p.  440;  Prichard,  On  the  Tahitians,  Waitz,  'Anthro- 

'  Phys.   Hist,    of    Mankind,'    vol.    i.  polog.'  Eng.  translat.  vol.  i.  p.  305. 

3rd  edit.  p.  321 ;  on  the  natives  of  Marsden,  quoted  by  Prichard, '  Phys. 

Arakhan,     ibid.    vol.    iv.     p.     537.  Hist,  of  Mankind,'  3rd  edit.  vol.  v. 

Wilson,        '  Pliysical       Ethnology,'  p.     67.     Lawrence,    '  Lectures     on 

Smithsonian    histitution,     1863,  p.  Physiology,'  p.  337. 
288;    on  the  Pijians,   p.   290.     Sir  "  This   fact  was    ascertained    in 

J.    Lubbock    ('Prehistoric    Times,'  the 'Reise  der  Novara:  Anthropolog. 

2nd  edit.    1869,  p.   506)  gives  an  Thiel,' Dr.  Weisbach,  1867,  s.  265. 
tscellent  resume  on  this  subject. 


584  The  Descent  of  Man,  Part  TIL 

"  vie  with  eacli  other  in  the  superlativeness  of  their  own  style." 
Dr.  "Wilson,  in  si3€aking  of  the  compressed  skulls  of  various 
American  races,  adds,  "such  usages  are  among  the  least  eradi- 
"  cable,  and  long  survive  the  shock  of  revolutions  that  change 
"  dynasties  and  efface  more  important  national  peculiarities."^^ 
The  same  principle  comes  into  play  in  the  art  of  breeding; 
and  we  can  thus  understand,  as  I  have  elsewhere  explained,^* 
the  wonderful  development  of  the  many  races  of  animals  and 
plants,  which  have  been  kept  merely  for  ornament.  Fanciers 
always  wish  each  character  to  be  somewhat  increased ;  they  do 
not  admire  a  medium  standard ;  they  certainly  do  not  desire 
any  great  and  abrupt  change  in  the  character  of  their  breeds ; 
tliey  admire  solely  what  they  are  accustomed  to,  but  they 
ardently  desire  to  see  each  characteristic  feature  a  little  more 
developed. 

The  senses  of  man  and  of  the  lower  animals  seem  to  be  so 
constituted  that  brilliant  colours  and  certain  forms,  as  well  as 
harmonious  and  rhythmical  sounds,  %\nq  pleasure  and  are  called 
beautiful;  but  why  this  should  be  so,  we  know  not.  It  is 
certainly  not  true  that  there  is  in  the  mind  of  man  any  universal 
standard  of  beauty  with  respect  to  the  human  body.  It  is, 
however,  possible  that  certain  tastes  may  in  the  course  of  time 
become  inherited,  though  there  is  no  evidence  in  favour  of  this 
belief;  and  if  so,  each  race  would  possess  its  own  innate  ideal 
standard  of  beauty.  It  has  been  argued  ^^  that  ugliness  consists 
in  an  approach  to  the  structure  of  the  lower  animals,  and  no 
doubt  this  is  partly  true  with  the  more  civilised  nations,  in 
which  intellect  is  highly  appreciated  ;  but  this  explanation  will 
hardly  apply  to  all  forms  of  ugliness.  The  men  of  each  race 
prefer  what  they  are  accustomed  to;  they  cannot  endure  any 
great  change ;  but  they  like  variety,  and  admire  each  cha- 
racteristic carried  to  a  moderate  extreme.'^''  Men  accustomed  to 
a  nearly  oval  face,  to  straight  and  regular  features,  and  to  bright 
colours,  admire,  as  we  Europeans  know,  these  points  when 
strongly  developed.  On  the  other  hand,  men  accustomed  to 
a  broad  face,  with  high  cheek-bones,  a  depressed  nose,  and  a 
black  skin,  admire  these  peculiarities  when  strongly  marked. 

"  'Smithsonian Institution,' 18G3,  Anthropologie,'  1866,  s.  164. 
J).   289.     On  the  fashions   of  Arab  '«  Jlr.  Bain  has  collected  ('Men- 
women,    Sir    S.    Baker,    'The    Kile  tal  and  Moral    Science,'    18G8,   pp. 
Tributaries,'  1867,  p.  121.  304-314)  about  a    dozen    more  or 

"■•  '  The  Variation  of  Animals  and  less  different  theories  of  the  idea  of 

Plants  under  Domestication,'  vol.  i.  beauty ;  but    none    are    quite    the 

p.  214  ;  vol.  ii.  p.  240.  same  as  that  here  given. 

"  Schaaffhausen,     'Archiv      fiir 


Chap.  XX.  Man — lexical  Selection.  5^5 

No  doubt  cliaracters  of  all  kinds  may  be  too  much  developed 
for  beauty.  Hence  a  perfect  beauty,  which  implies  many 
characters  modified  in  a  particular  manner,  will  be  in  every 
race  a  prodigy.  As  the  great  anatomist  Bichat  long  ago  said, 
if  every  one  were  cast  in  the  same  mould,  there  would  be  no 
such  tiling  as  beauty.  If  all  our  women  were  to  become  as 
beautiful  as  the  Venus  de'  Medici,  we  should  for  a  time  be 
charmed  ;  but  we  should  soon  wish  for  variety  ;  and  as  soon  as 
we  had  obtained  variety,  we  should  wish  to  see  certain  cha- 
racters a  little  exaggerated  beyond  the  then  existing  common 
standard. 


CHAPTEE  XX. 
Secondary  Sexual  Characters  of  IIk-^— continued. 

On  the  effects  of  the  continued  selection  of  women  according  to  a  different 
standard  of  beauty  in  each  race — On  the  causes  which  interfere  with 
sexual  selection  in  civilised  and  savage  nations — Conditions  favourable 
to  sexual  selection  during  primeval  times — On  the  manner  of  action 
of  sexual  selection  with  mankind — On  the  women  in  savage  tribes  having 
some  power  to  choose  their  husbands — Absence  of  hair  on  the  body,  and 
development  of  the  beard — Colour  of  the  skiu — Summary. 

We  have  seen  in  the  last  chapter  that  with  all  barbarous  races 
ornaments,  dress,  and  external  appearance  are  highly  valued ; 
and  that  the  men  judge  of  the  beauty  of  their  women  by  widely 
different  standards.  We  must  next  inquire  whether  this  pre- 
ference and  the  consequent  selection  during  many  generations  of 
those  women,  which  appear  to  the  men  of  each  race  the  most 
attractive,  has  altered  the  character  either  of  the  females  alone, 
or  of  both  sexes.  With  mammals  the  general  rule  appears 
to  be  that  characters  of  all  kinds  are  inherited  equally  by  the 
males  and  females ;  we  might  therefore  expect  that  with  man- 
kind any  characters  gained  by  the  females  or  by  the  males 
through  sexual  selection,  would  commonly  be  transferred  to  the 
offspring  of  both  sexes.  If  any  change  has  thus  been  effected,  it 
is  almost  certain  that  the  different  races  would  be  differently 
modified,  as  each  has  its  own  standard  of  beauty. 

With  mankind,  especially  with  savages,  many  causes  interfoi(_' 
with  the  action  of  sexual  selection  as  far  as  the  bodily  frame  is 
concerned.  Civilised  men  are  largely  attracted  by  the  mental 
charms  of  women,  by  their  wealth,  and  especially  by  their  social 
position ;  for  men  rarely  marry  into  a  much  lower  rank.  The 
men  who  succeed  in  obtaining  the  more  beautilul  women,  will 
not  have  a  better  chance  of  leaving  a  long  line  of  descendants 
2() 


586  TJie  Descent  of  Man.  Part  III. 

than  other  men  with  plainer  wives,  save  the  few  who  bequeath 
their  fortunes  according  to  primogeniture.  With  respect  to  the 
opposite  form  of  selection,  namely  of  the  more  attractiye  men  by 
the  women,  although  in  civilised  nations  women  have  free  or 
almost  free  choice,  which  is  not  the  case  with  barbarous  races, 
yet  their  choice  is  largely  influenced  by  the  social  position  and 
wealth  of  the  men ;  and  the  success  of  the  latter  in  life  depends 
much  on  their  intellectual  powers  and  energy,  or  on  the  fruits  of 
these  same  powers  in  their  forefathers.  iS^o  excuse  is  needed  for 
treating  this  subject  in  some  detail ;  for,  as  the  German  philo- 
sopher Schopenhauer  remarks,  "the  final  aim  of  all  love 
"  intrigues,  be  they  comic  or  tragic,  is  really  of  more  importance 
"  than  all  other  ends  in  human  life.  What  it  all  turns  upon  is 
"  nothing  less  than  the  composition  of  the  next  generation.  .  .  . 
"  It  is  not  the  weal  or  woe  of  any  one  individual,  but  that  of  the 
"  human  race  to  come,  which  is  here  at  stake."  ^ 

There  is,  however,  reason  to  believe  that  in  certain  civilised 
and  semi-civilised  nations  sexual  selection  has  effected  some- 
thing in  modifying  the  bodily  frame  of  some  of  the  members. 
Many  persons  are  convinced,  as  it  appears  to  me  with  justice, 
that  our  aristocracy,  including  under  this  term  all  wealthy 
families  in  which  primogeniture  has  long  prevailed,  from  having 
chosen  during  many  generations  from  all  classes  the  more  beau- 
tiful women  as  their  wives,  have  become  handsomer,  according 
to  the  European  standard,  than  the  middle  classes ;  yet  the 
middle  classes  are  placed  under  equally  favoi\rable  conditions  of 
life  for  the  perfect  development  of  the  body.  Cook  remarks  that 
the  superiority  in  personal  appearance  "  which  is  observable  in 
"  the  erees  or  nobles  in  all  the  other  islands  (of  the  Pacific)  is 
"  found  in  the  Sandwich  islands ; "  but  this  may  be  chiefly  due 
to  their  better  food  and  manner  of  life. 

The  old  traveller  Chardin,  in  describing  the  Persians,  says 
their  ''  blood  is  now  highly  refined  by  frequent  intermixtures 
"  with  the  Georgians  and  Circassians,  two  nations  which  surpass 
"  all  the  world  in  personal  beauty.  There  is  hardly  a  man  of 
"  rank  in  Persia  who  is  not  born  of  a  Georgian  or  Circassian 
J'  mother."  He  adds  that  they  inherit  their  beauty,  "not  from 
*'  their  ancestors,  for  without  the  above  mixture,  the  men  of 
*'  rank  in  Persia,  who  are  descendants  of  the  Tartars,  would  be 
*'  extremely  ugly."  -    Here  is  a  more  curious  case ;  the  priestesses 

*  'Schopenhauer  and  Darwinism,'  &c.  1822,  p.   393),   who  attributes 

in  'Journal  of  Anthropology,'  Jan.  the  beauty  of  the  upper  classes  io 

1871,  p.  323.  England  to   the  men    having    long 

"^  These  quotations  are  taken  from  elected  the  more  beautiful  women. 
Lawrence  ('  Lectures  on  Physiology,' 


CuAp.  XX.      Man  — Checks  to  Sexual  Selection.  5  8/ 

who  attended  the  temple  of  Venus  Erycina  at  SanGiuliano  in 
Sicily,  were  selected  for  their  beauty  out  of  the  whole  of  Greece 
they  were  not  vestal  virgins,  and  Quatrefages/  who  states  the 
foregoing  fact,  says  that  the  women  of  San-Giuliano  are  now 
famous  as  the  most  beautiful  in  the  island,  and  are  sought  by 
artists  as  models.  But  it  is  obvious  that  the  evidence  in  all  the 
above  cases  is  doubtful. 

The  following  case,  though  relating  to  savages,  is  well  worth 
giving  from  its  curiosity.  Mr.  AVinwood  Eeade  informs  me  that 
the  Jollofs,  a  tribe  of  negroes  on  the  west  coast  of  Africa,  "  are 
"  remarkable  for  their  uniformly  fine  appearance."  A  friend  of 
his  asked  one  of  these  men,  *'  How  is  it  that  every  one  whom  I 
*'  meet  is  so  iine-looking,  not  only  your  men,  but  your  women  ?" 
The  Jollof  answered,  "  It  is  very  easily  explained :  it  has  always 
"  been  our  custom  to  pick  out  our  worse-looking  slaves  and  to 
"  sell  them."  It  need  hardly  be  added  that  with  all  savages, 
female  slaves  serve  as  concubines.  That  this  negro  should  have 
attributed,  whether  rightly  or  wrongly,  the  fine  appearance  of 
his  tribe  to  the  long-continued  elimination  of  the  ugly  women  is 
not  so  surprising  as  it  may  at  first  appear  ;  for  I  have  elsewhere 
shewn  *  that  negroes  fully  appreciate  the  importance  of  selection 
in  the  breeding  of  their  domestic  animals,  and  I  could  give  from 
Mr.  Eeade  additional  evidence  on  this  head. 

The  Causes  ivhich  prevent  or  check  the  Action  of  Sexual  Selection 
with  Savages. — The  chief  causes  are,  first,  so-called  communal 
marriages  or  promiscuous  intercourse ;  secondly,  the  consequences 
of  female  infanticide ;  thirdly,  early  betrothals ;  and  lastly,  the 
low  estimation  in  which  women  are  held,  as  mere  slaves.  These 
four  points  must  be  considered  in  some  detail. 

It  is  obvious  that  as  long  as  the  pairing  of  man,  or  of  any 
other  animal,  is  left  to  mere  chance,  with  no  choice  exerted  by 
either  sex,  there  can  be  no  sexual  selection ;  and  no  effect  will  be 
produced  on  the  offspring  by  certain  individuals  having  had  an 
advantage  over  others  in  their  couj-tship.  Now  it  is  asserted 
that  there  exist  at  the  present  day  tribes  which  practise  what 
Sir.  J.  Lubbock  by  courtesy  calls  communal  marriages ;  that  is, 
all  the  men  and  women  in  the  tribe  are  husbands  and  wives  to 
one  another.  The  licentiousness  of  many  savages  is  no  doubt 
astonishing,  but  it  seems  to  me  that  more  evidence  is  requisite, 
before  we  fully  admit  that  th&ir  intercourse  is  in  any  case  pro- 
miscuous.   Nevertheless  all  those  who  have  most  closely  studied 

*  *  Aathropologie,'  'Revue  des  Plants  under  Domestication,'  vol.  i. 
Cours    Scientifiques,'  Oct.   1868,  p.      p.  207. 

721.  *  Sir  J.  Lubbock,  'The  Origin  of 

*  'The  Variation  of  Animals  and      CiTilisation,'  1870,  chap.  iii.  especi- 


588  The  Descent  of  Man.  Part  IIL 

tlie  subject;^  and  whose  judgment  is  worth  much  more  than  mine, 
believe  that  communal  marriage  (this  expression  being  variously 
guarded)  was  the  original  and  universal  form  throughout  the 
world,  including  therein  the  intermarriage  of  brothers  and  sisters. 
The  late  Sir.  A.  Smith,  who  had  travelled  widely  in  S.  Africa,  and 
knew  much  about  the  habits  of  savages  there  and  elsewhere,  ex- 
pressed to  me  the  strongest  opinion  that  no  race  exists  in  which 
woman  is  considered  as  the  property  of  the  community.  I  believe 
that  his  judgment  w\as  largely  determined  by  what  is  implied  by . 
the  term  marriage.  Throughout  the  following  discussion  I  use 
the  term  in  the  same  sense  as  when  naturalists  speak  of  animals 
as  monogamous,  meaning  thereby  that  the  male  is  accepted  by  or 
chooses  a  single  female,  and  lives  wath  her  either  during  the 
breeding-season  or  for  the  wdiole  year,  keeping  possession  of 
her  by  the  law  of  might ;  or,  as  when  they  speak  of  a  polygamous 
species,  meaning  that  the  male  lives  with  several  females.  This 
kind  of  marriage  is  all  that  concerns  us  here,  as  it  suflQces  for 
the  work  of  sexual  selection.  But  I  know  that  some  of  the 
writers  above  referred  to,  imply  by  the  term  marriage,  a  recog- 
nised right,  protected  by  the  tribe. 

The  indii-ect  evidence  in  favour  of  the  belief  of  the  former 
prevalence  of  communal  marriages  is  strong,  and  rests  chiefly  on 
the  terms  of  relationshij)  which  are  employed  between  the 
members  of  the  same  tribe,  implying  a  connection  with  the  tribe, 
and  not  with  either  parent.  But  the  subject  is  too  large  and 
comj^lex  for  even  an  abstract  to  be  here  given,  and  I  will  confine 
myself  to  a  few  remarks.  It  is  evident  in  the  case  of  such 
marriages,  or  where  the  marriage  tie  is  very  loose,  that  the 
relationship  of  the  child  to  its  father  cannot  be  known.  But  it 
seems  almost  incredible  that  the  relationship  of  the  child  to  its 
mother  should  ever  be  completely  ignored,  especially  as  the  women 
in  most  savage  tribes  nurse  their  infants  for  a  long  time. 
Accordingly,  in  many  cases  the  hues  of  descent  are  traced 
through  the  mother  alone,  to  the  exclusion  of  the  father.  But 
in  other  cases  the  terms  employed  exi^ress  a  connection  with  the 

ally  pp.  60-67.     Mr.   M'Lennan,  in  memoir  on  the  classificatory  system 

his    extremely    rahiable    work    on  of  relationship    ('  Proc.     American 

'  Primitive  Marriage,'  1865,  p.  163,  Acad,   of    Sciences,'  vol.    vii.    Feb. 

speaks    of   the    union  of   the  sexes  1868,  p.  475),  concludes  that  poly- 

"  in    the    earliest    times    as    loose,  gamy  and  all   forms     of    marriage 

"  transitory,    and    in    some    degree  during  primeval  times  were  essen- 

"  promiscuous."  Mr.  M'Lennan  and  tially   unknown.     It    appears    also, 

Sir  J.  Lubbock  have  collected  much  from  Sir  J.    Lubbock's  work,   that 

evidence  on  the  extreme  licentious-  Bachofen     likewise     believes    that 

ness  of  savages  at  the  present  time.  communal     intercourse     originally 

Mr.  L.  H.  Morgan,  in  his  intcjesting  prevailed. 


Chap.  XX.      Ma7i — Checks  to  Sexuhl  Selection.  589 

tribe  alone,  to  the  exclusion  even  of  the  mother.  It  seems 
possible  that  the  connection  between  the  related  members  of  the 
same  barbarous  tribe,  exposed  to  all  sorts  of  danger,  might  be  so 
much  more  important,  owing  to  the  need  of  mutual  protection 
and  aid,. than  that  between  the  mother  and  her  child,  as  to  lead 
to  the  sole  use  of  terms  expressive  of  the  former  relationships  ; 
but  Mr.  Morgan  is  convinced  that  this  view  is  by  no  means 
sufficient. 

The  terms  of  relationship  used  in  different  parts  of  the  world 
may  be  divided,  according  to  the  author  just  quoted,  into  two 
great  classes,  the  classificatory  and  descriptive,— the  latter  being 
employed  by  us.  It  is  the  classificatory  system  which  so  strongly 
leads  to  the  belief,  that  communal  and  other  extremely  loose 
forms  of  marriage  were  originally  universal.  But  as  far  as  I  can 
see,  there  is  no  necessity  on  this  ground  for  believing  in  abso- 
lutely jDromiscuous  intercourse  ;  and  I  am  glad  to  find  that  this 
is  Sir.  J.  Lubbock's  view.  Men  and  women,  like  many  of  the 
lower  animals,  might  formerly  have  entered  into  strict  though 
temporary  unions  for  each  birth^and  in  this  case  nearly  as  much 
confusion  would  have  arisen  in  the  terms  of  relationship,  as  in 
the  case  of  promiscuous  intercourse.  As  far  as  sexual  selection 
is  concerned,  all  that  is  required  is  that  choice  should  be  exerted 
before  the  parents  unite,  and  it  signifies  little  whether  the  unions 
last  for  Kfe  or  only  for  a  season. 

Besides  the  evidence  derived  from  the  terms  of  relationship, 
other  lines  of  reasoning  indicate  the  former  wide  prevalence  of 
communal  marriage.  Sir.  J.  Lubbock  accounts^  for  the  strange 
and  widely-extended  habit  of  exogamy — that  is,  the  men  of  one 
tribe  taking  wives  from  a  distinct  tribe, — by  communism  having 
been  the  original  form  of  intercourse;  so  that  a  man  never 
obtained  a  wife  for  himself  unless  he  captured  her  from  a 
neighbouring  and  hostile  tribe,  and  then  she  would  naturally 
have  become  his  sole  and  valuable  property.  Thus  the  practice 
of  captui'ing  wives  might  have  arisen ;  and  from  the  honour  so 
gained  it  might  ultimately  have  become  the  universal  habit. 
According  to  Sir  J.  Lubbock,"  we  can  also  thus  understand  "  the 
"  necessity  of  expiation  for  marriage  as  an  infringement  of  tribal 
"  rites,  since,  according  to  old  ideas,  a  man  had  no  right  to 
"  appropriate  to  himself  that  which  belonged  to  the  whole 
"  tribe."  Sir  J.  Lubbock  farther  gives  a  curious  body  of 
facts  shewing  that  in  old  times  high  honour  was  bestowed  on 
women  who  were  utterly  licentious;  and  this,  as  he  explains,  is 

•  Address  to  British  Association  dition  of  the  Lower  Races  of  Man,  ' 
*  On  the  Social  and  Religious  Cou-      1870,  p.  20. 


590  The  Descent  of  Man.  Part  III. 

intelligible,  if  we  admit  that  promiscuous  intercourse  was  the 
aboriginal,  and  therefore  long  revered  custom  of  the  tribe  J 

Although  the  manner  of  development  of  the  maiTiage-tie  is  an 
obscure  subject,  as  we  may  infer  from  the  divergent  opinions  on 
several  points  between  the  tliree  authors  who  have  studied  it 
most  closely,  namely,  Mr.  Morgan,  Mr,  M'Lennan,  and  Sir  J, 
Lubbock,  yet  from  the  foregoing  and  several  other  lines  of 
evidence  it  seems  i^robable  ^  that  the  habit  of  marriage,  in  any 
strict  sense  of  the  word,  has  been  gradually  developed  ;  and  that 
almost  promiscuous  or  very  loose  intercourse  was  once  ex- 
tremely common  thi-oughout  the  world.  Nevertheless  from  the 
strength  of  the  feeling  of  jealousy  all  through  the  animal 
kingdom,  as  well  as  from  the  analogy  of  the  lower  animals,  more 
particularly  of  those  which  come  nearest  to  man,  I  cannot 
believe  that  absolutely  promiscuous  intercourse  prevailed  in 
times  past,  shortly  before  man  attained  to  his  present  rank  in 
the  zoological  scale.  Man,  as  I  have  attempted  to  shew,  is 
certainly  descended  from  some  ajDC-like  creature.  With  the 
existing  Quadrumana,  as  far  as  their  habits  are  known,  the 
males  of  some  species  are  monogamous,  but  live  during  only  a 
part  of  the  year  with  the  females;  of  this  the  orang  seems  to 
afford  an  instance.  Several  kinds,  for  example  some  of  the 
Indian  and  American  monkeys,  are  strictly  monogamous,  and 
associate  all  the  year  round  with  their  wives.  Others  are  poly- 
gamous, for  example  the  gorilla  and  several  American  sj^ecies, 
and  each  family  lives  separate.  Even  when-  this  occurs,  the 
families  inhabiting  the  same  district  are  probably  somewhat 
social :  the  chimpanzee,  for  instance,  is  occasionally  met  with  in 
large  bands.  Again,  other  species  are  polygamous,  but  several 
males,  each  with  his  own  females,  live  associated  in  a  body,  as  with 
several  species  of  baboons.^  We  may  indeed  conclude  from  what 
we  know  of  the  jealousy  of  all  male  quadrupeds,  armed,  as  many 
of  them  are,  with  special  weapons  for  battling  with  their  rivals, 
that  promiscuous  intercoui'se  in  a  state  of  nature  is  extremely 

'  'Origin  of  Civilisation,'  1870,  classificatory  system  of  relationship 
p.  8G.  In  the  several  works  ahove  can  be  otherwise  explained, 
quoted,  there  will  be  found  copious  ^  Brehm  ('Illust.  Theirleben,'  B. 
evidence  on  relationship  throucrh  i.  p.  77)  says  Cynocephalus  hama- 
the  females  alone,  or  with  the  tribe  dryas  lives  in  great  troops  contain- 
alone,  ing  twice  as  many  adult  females  as 

'  Mr.  C.  Staniland  Wake  argues  adult  males.     See  Renggor  on  Ame- 

strongly    ('Anthropologia,'    ]\Iarch,  rican  polygamous  species,  and  0\ven 

1874,  ]).  197)  against  the  views  held  ('  Anat.  of  Vertebrates,'  vol.  iii.  p. 

by  these  three  writers  on  the  former  746)     on    American     monogamous 

prevalence    of  almost    promiscuous  species.     Other  references  might  be 

intercourse;  and  he  thinks  that  the  added 


Cr.Ar.  XX.     Ma?i — Checks  to  Sexual  Selection.  591 

improbable.  The  pairing  may  not  last  for  life,  but  only  for  each 
birth;  yet  if  the  males  which  are  the  strongest  and  best  able  to 
defend  or  otherwise  assist  their  females  and  young,  were  to 
select  the  more  attractive  females,  this  would  suffice  for  sexual 
selection. 

Therefore,  looking  far  enough  back  in  the  stream  of  time,  and 
judging  from  the  social  habits  of  man  as  he  now  exists,  the  most 
probable  view  is  that  he  aboriginally  lived  in  small  communities, 
each  with  a  single  wife,  or  if  powerful  with  several,  whom  he 
jealously  guarded  against  all  other  men.  Or  he  may  not  have 
been  a  social  animal,  and  yet  have  lived  with  several  wives,  like 
the  gorilla ;  for  all  the  natives  "  agree  that  but  one  adult  male 
"  is  seen  in  a  band ;  when  the  young  male  grows  up,  a  contest 
"  takes  place  for  mastery,  and  the  strongest,  by  killing  and 
"  driving  out  the  others,  establishes  himself  as  the  head  of  the 
"  community."  ^'^  The  younger  males,  being  thus  expelled  and 
wandering  about,  would,  when  at  last  successful  in  finding  a 
partner,  prevent  too  close  interbreeding  within  the  limits  of  the 
same  family. 

Although  savages  are  now  extremely  licentious,  and  although 
communal  marriages  may  formerly  have  largely  prevailed,  yet 
many  tribes  practise  some  form  of  marriage,  but  of  a  far  more  lax 
nature  than  that  of  civilised  nations.  Polygamy,  as  just  stated, 
is  almost  universally  followed  by  the  leading  men  in  every  tribe. 
Nevertheless  there  are  tribes,  standing  almost  at  the  bottom  of 
the  scale,  which  are  strictly  monogamous.  This  is  the  case  with 
the  Veddahs  of  Ceylon  :  they  have  a  saying,  according  to  Sir  J. 
Lubbock,^^  "  that  death  alone  can  separate  husband  and  wife." 
An  inteHigent  Kandyan  chief,  of  course  a  polygamist,  "  was 
"  perfectly  scandalised  at  the  utter  barbarism  of  living  with 
"  only  one  wife,  and  never  parting  until  separated  by  death." 
It  was,  he  said,  '*  Just  like  the  Wanderoo  monkeys."  Whether 
savages  who  now  enter  into  some  form  of  marriage,  either  poly- 
gamous or  monogamous,  have  retained  this  habit  from  j^rimeval 
times,  or  whether  they  have  returned  to  some  form  of  marriage, 
after  passing  through  a  stage  of  promiscuous  intercourse,  I  will 
not  pretend  to  conjecture. 

Lifanticid€.—T\i\s,  practice  is  now  very  common  throughout 
the  world,  and  there  is  reason  to  believe  that  it  prevailed  much 
more  extensively  during    former  times.^^    Barbarians    find    it 

>o  Dr.  Savacre,  in  '  Boston  Journal  ^-    Mr.      M'Lenuan,     'Primitive 

of  Nat.    Hist.''  vol.  v.   1845-47,  p.  Marriage,'  1865.     See  especially  on 

423.  exogamy  and   infanticiJe,   pp.    loO, 

"  'Prehistoric    Times,'   1839,   p.  lo8,  165. 
424. 


592  TJie  Descent  of  Man.  Part  III. 

difficult  to  support  themselves  and  their  children,  and  it  is  a 
simple  plan  to  kill  their  infants.  In  South  America  some  tribes, 
according  to  Azara,  formerly  destroyed  so  many  infants  of  both 
sexes,  that  they  were  on  the  point  of  extinction.  In  the  Poly- 
nesian Islands  women  have  been  known  to  kill  from  four  or  five, 
to  even  ten  of  their  children ;  and  Ellis  could  not  find  a  single 
woman  who  had  not  killed  at  least  one.  Wherever  infanticide 
prevails  the  struggle  for  existence  will  be  in  so  far  less  severe, 
and  all  the  members  of  the  tribe  will  have  an  almost  equally  good 
chance  of  rearing  their  few  surviving  children.  In  most  cases  a 
larger  number  of  female  than  of  male  infants  are  destroyed,  for 
it  is  obvious  that  the  latter  are  of  more  value  to  the  tribe,  as 
they  will,  when  grown  up,  aid  in  defending  it,  and  can  support 
themselves.  But  the  trouble  experienced  by  the  women  in  rearing 
children,  their  consequent  loss  of  beauty,  the  higher  estima- 
tion set  on  them  when  few  and  their  happier  fate,  are  assigned 
by  the  women  themselves,  and  by  various  observers,  as  additional 
motives  for  infanticide.  In  Austraha,  where  female  infanticide 
is  still  common,  Sir  G.  Grey  estimated  the  proportion  of  native 
women  to  men  as  one  to  three ;  but  others  say  as  two  to  three. 
In  a  village  on  the  eastern  frontier  of  India,  Colonel  MacCulloch 
found  not  a  single  female  child.^^ 

"When,  owing  to  female  infanticide,  the  women  of  a  tribe  were 
few,  the  habit  of  capturing  wives  from  neighbouring  tribes  "^ould 
naturally  arise.  Sir  J.  Lubbock,  however,  as  we  have  seen, 
attributes  the  practice  in  chief  part,  to  the  former  existence  of 
communal  marriage,  and  to  the  men  having  consequently 
captured  women  from  other  tribes  to  hold  as  theu'  sole  jDroperty. 
Additional  causes  might  be  assigned,  such  as  the  communities 
being  very  small,  in  which  case,  marriageable  women  would 
often  be  deficient.  That  the  habit  was  most  extensively  practised 
during  former  times,  even  by  the  ancestors  of  civilised  nations, 
is  clearly  shewn  by  the  i^reservation  of  many  curious  customs 
and  ceremonies,  of  which  Mr.  M'Lennan  has  given  an  interesting 
account.  In  our  own  marriages  the  "  best  man  "  seems  origin- 
ally to  have  been  the  chief  abettor  of  the  bridegroom  in  the  act 
of  capture.  Now  as  long  as  men  habitually  procured  their  wives 
through  violence  and  craft,  they  would  have  been  glad  to  seize  on 
any  woman,  and  would  not  have  selected  the  more  attractive  ones. 
But  as  soon  as  the  practice  of  procuring  wives  from  a  distinct 
tribe  was  effected  through  barter,  as  now  occurs  in  many  places, 

■3  Dr.  Gerland  (*  Ueber  das  Aus-  54-.     Azara  ('Voyages,'  &c.  torn.  ii. 

sterbea  der  Naturvolker,'  1868)  has  pp.  94,  116)  enters  in  detail  on  the 

collected  much    information  on  in-  motives.     See  also  M'Lennan  (ibid, 

fanticide,  see    especially  s.   27,  51,  p.  139)  for  cases  in  India. 


Chap.  XX.      Man — Checks  to  Sexual  Se lection.  593 

the  more  attractive  women  would  generally  have  been  purchased. 
The  incessant  crossing,  however,  between  tribe  and  tribe,  which 
necessarily  follows  from  any  form  of  this  habit,  would  tend  to 
keep  all  the  people  inhabiting  the  same  country  nearly  uniform 
in  character ;  and  this  would  interfere  with  the  power  of  sexual 
selection  in  differentiating  the  tribes. 

The  scarcity  of  women,  consequent  on  female  infanticide,  leads, 
also,  to  another  practice,  that  of  polyandry,  still  common  in 
several  j)arts  of  the  world,  and  which  formerly,  as  Mr.  M'Lennan 
believes,  prevailed  almost  universally  ;  but  this  latter  conclusion 
is  doubted  by  Mr.  Morgan  and  Sir.  J.  Lubbock.^*  '  Whenever  two 
or  more  men  are  compelled  to  marry  one  woman,  it  is  certain 
that  all  the  women  of  the  tribe  will  get  married,  and  there  will 
be  no  selection  by  the  men  of  the  more  attractive  women.  But 
under  these  circumstances  the  women  no  doubt  will  have  the 
power  of  choice,  and  will  prefer  the  more  attractive  men.  Azara, 
for  instance,  describes  how  carefully  a  Guana  woman  bargains 
for  all  sorts  of  privileges,  before  accepting  some  one  or  more 
husbands ;  and  the  men  in  consequence  take  unusual  care  of 
their  personal  appearance.  So  amongst  the  Todas  of  India,  who 
practise  polyandry,  the  girls  can  accept  or  refuse  any  man.^^ 
A  very  ugly  man  in  these  cases  would  perhaps  altogether  fail 
in  getting  a  wife,  or  get  one  later  in  hfe ;  but  the  handsomer 
men,  although  more  successful  in  obtaining  wives,  would  not,  as 
far  as  we  can  see,  leave  more  offspring  to  inherit  their  beauty 
than  the  less  handsome  husbands  of  the  same  women. 

Early  Betrothals  and  Slavery  of  Women. — With  many  savages 
it  is  the  custom  to  betroth  the  females  whilst  mere  infants ;  and 
this  would  effectually  prevent  preference  being  exerted  on  either 
side  according  to  personal  aiDpearance.  But  it  would  not  prevent 
the  more  attractive  women  from  being  afterwards  stolen  or  taken 
by  force  from  their  husbands  by  the  more  powerful  men ;  and 
this  often  happens  in  Australia,  America,  and  elsewhere.  The 
same  consequences  with  reference  to  sexual  selection  would  to  a 
certain  extent  follow,  when  women  are  valued  almost  solely  as 
slaves  or  beasts  of  burden,  as  is  the  case  with  many  savages. 
The  men,  however,  at  all  times  would  prefer  the  handsomest 
slaves  according  to  their  standard  of  beauty. 

We  thus  see 'that  several  customs  prevail  with  savages  which 
must  greatly  interfere  with,  or  completely  stop,  the  action  of 
sexual  selection.     On  the  other  hand,  the  conditions  of  life  to 

^*  *  Primitive  Marriage,'  p.  208  ;  polyandry. 
Sir  J.  Lubbock,  '  Origiu  of  Civilisa-  '^  Azara,  '  Voyages,'  kc.  torn.  ii. 

tion,'  p.  100.     See  also  Mr.  Morgan,  pp.      92-95,       Colonel       Marshall, 

loc.  cit.,  oh  the  former  i)re valence  of  'Amongst  the  Todas,   p.  212. 


594  ^^^^  Descent  of  Man.  Part  III. 

which  savages  are  exposed,  and  some  of  their  habits,  are  favour- 
able to  natural  selection ;  and  this  comes  into  play  at  the  same 
time  with  sexual  selection.  Savages  are  known  to  suffer  severely 
from  recurrent  famines;  they  do  not  increase  their  food  by 
artificial  means ;  they  rarely  refrain  from  marriage/^  and  generally 
marry  whilst  young.  Consequently  they  must  be  subjected  to 
occasional  hard  struggles  for  existence,  and  the  favoured  indi- 
viduals will  alone  survive. 

At  a  very  early  period,  before  man  attained  to  his  present 
rank  in  the  scale,  many  of  his  conditions  would  be  different  from 
what  now  obtains  amongst  savages.  Judging  from  the  analogy 
of  the  lower  animals  he  would  then  either  live  with  a  single 
female,  or  be  a  polygamist.  The  most  powerful  and  able  males 
would  succeed  best  in  obtaining  attractive  females.  They  would 
also  succeed  best  in  the  general  struggle  for  life,-  and  in  defend- 
ing their  females,  as  well  as  their  offspring,  from  enemies  of  all 
kinds.  At  this  early  period  the  ancestors  of  man  would  not  be 
sufficiently  advanced  in  intellect  to  look  forward  to  distant 
contingencies ;  they  would  not  foresee  that  the  rearing  of  all 
their  children,  especially  their  female  children,  would  make  the 
struggle  for  life  severer  for  the  tribe.  They  would  be  governed 
more  by  their  instincts  and  less  by  their  reason,  than  are  savages 
at  the  present  day.  They  would  not  at  that  period  have 
partially  lost  one  of  the  strongest  of  all  instincts,  common  to  all 
the  lower  animals,  namely  the  love  of  their  young  offspring  ;  and 
consequently  they  would  not  have  practised  female  infanticide. 
Women  would  not  have  been  thus  rendered  scarce,  and  poly- 
andry would  not  have  been  practised ;  for  hardly  any  other  cause, 
except  the  scarcity  of  women  seems  sufficient  to  break  down  the 
natural  and  widely  prevalent  feeling  of  jealousy,  and  the  desire  of 
each  male  to  possess  a  female  for  himself.  Polyandry  would  be  a 
natural  stepping-stone  to  communal  marriages  or  almost  pro- 
miscuous intercourse  ;  though  the  best  authorities  believe  that 
this  latter  habit  preceded  polyandry.  During  primordial  times 
there  would  be  no  early  betrothals,  for  this  implies  foresight.  Nor 
would  women  be  valued  merely  as  useful  slaves  or  beasts  of 
burthen.  Both  sexes,  if  the  females  as  well  as  the  males  were  per- 
mitted to  exert  any  choice,  would  choose  their  partners  not  for 
mental  charms,  or  property,  or  social  position,  btit  almost  solely 
from  external  appearance.     All  the  adults  would  marry  or  pair, 

^^  Burchell  says  ('Travels  in  S.  Azara  ('Voyages    dans    I'Amerique 

Africa,'  vol.   ii.    1824,  p.  58),  that  Merid.'  torn.  ii.  1809,  p.  21)  makes 

among  the  wild  nations  of  Southern  precisely  the  same  remark  in  regard 

Africa,  neither  men  nor  women  ever  to  the  wild  Indians  of  South  Ame- 

pass  their  lives  in  a  state  of  celibacy.  rica. 


Chap.  XX.       Man — Mode  of  Sexual  Selection.  595 

1 

and  all  the  ofifspring,  as  far  as  that  was  possible,  would  be  reared ; 
so  that  the  struggle  for  existence  would  be  periodically  ex- 
cessively severe.  Thus  during  these  times  all  the  conditions  for 
sexual  selection  would  have  been  more  favourable  than  at  a 
later  period,  when  man  had  advanced  in  his  intellectual  powers 
but  had  retrograded  in  his  instincts.  Therefore,  whatever 
influence  sexual  selection  may  have  had  in  producing  the 
differences  between  the  races  of  man,  and  between  man  and 
the  higher  Quadrumana,  this  influence  would  have  been  more 
powerful  at  a  remote  period  than  at  the  present  day,  though 
probably  not  yet  wholly  lost. 

Tlxe  Manner  of  Action  of  Sexual  Selection  v  ith  Manldnd. — With 
primeval  men  under  the  favourable  conditions  just  stated,  and 
with  those  savages  who  at  the  present  time  enter  into  any 
marriage  tie,  sexual  selection  has  probably  acted  in  the  following 
manfter,  subject  to  greater  or  less  interference  from  female  in- 
fanticide, early  betrothals,  &c.  The  strongest  and  most  vigorous 
men,— those  who  could  best  defend  and  hunt  for  their  famihes, 
who  were  provided  with  the  best  weapons  and  possessed  the 
most  property,  such  as  a  large  number  of  dogs  or  other 
animals,— would  succeed  in  rearing  a  greater  average  number  of 
offspring  than  the  weaker  and  poorer  members  of  the  same 
tribes.  There  can,  also,  be  no  doubt  that  such  men  would 
generally  be  able  to  select  the  more  attractive  women.  At 
present  the  chiefs  of  nearly  every  tribe  throughout  the  world 
succeed  in  obtaining  more  than  one  wife.  I  hear  from  Mr. 
Mantell,  that  until  recently,  almost  every  girl  in  New  Zealand, 
who  was  pretty,  or  promised  to  be  pretty,  was  tapii  to  some 
chief.  With  the  Kafirs,  as  Mi\  C.  Hamilton  states,^^  "  the 
"  chiefs  generally  have  the  pick  of  the  women  for  many  miles 
"  round,  and  are  most  persevering  in  establishing  or  confii-ming 
"  their  privilege."  We  have  seen  that  each  race  has  its  own 
style  of  beauty,  and  we  know  that  it  is  natural  to  man  to  admire 
each  characteristic  point  in  his  domestic  animals,  dress,  orna- 
ments, and  personal  appearance,  when  carried  a  little  beyond  the 
average.  If  then  the  several  foregoing  propositions  be  admitted, 
and  I  cannot  see  that  they  are  doufetful,  it  would  be  an  inex- 
plicable circumstance,  if  the  selection  of  the  more  attractive  women 
by  the  more  powerful  men  of  each  tribe,  who  would  rear  on  an 
average  a  greater  number  of  children,  did  not  after  the  lapse 
of  many  generations  somewhat  modify  the  character  of  the  tribe. 

When  a  foreign  breed  of  our  domestic  animals  is  introduced 
into  a  new  country,  or  when  a  native  breed  is  long  and  carefully 

"  'Anthropological  Review,' Jan.  1870,  p.  xyi. 


59^  The  Descent  of  Man.  Part  ]  II. 

attended  to,  either  for  use  or  ornament,  it  is  found  after  several 
generations  to  have  undergone  a  greater  or  less  amount  of  change, 
whenever  the  means  of  comparison  exist.  This  follows  from 
unconscious  selection  during  a  long  series  of  generations — that  is, 
the  preservation  of  the  most  approved  individuals — without  any 
wish  or  expectation  of  such  a  result  on  the  part  of  the  breeder. 
So  again,  if  during  many  years  two  careful  breeders  rear  animals 
of  the  same  family,  and  do  not  compare  them  together  or  with  a 
common  standard,  the  animals  are  found  to  have  become,  to  the 
surprise  of  their  owners,  shghtly  different.  ^^  Each  breeder  has 
impressed,  as  Von  Nathusius  well  expresses  it,  the  character  of 
his  own  mind — his  own  taste  and  judgment — on  his  animals. 
What  reason,  then,  can  be  assigned  why  similar  results  should 
not  follow  from  the  long-continued  selection  of  the  most  admired 
women  by  those  men  of  each  tribe,  who  were  able  to  rear  the 
greatest  number  of  children?  This  would  be  unconscious 
selection,  for  an  effect  would  be  produced,  independently  of  any 
wish  or  expectation  on  the  part  of  the  men  who  preferred  certain 
women  to  others. 

Let  us  suppose  the  members  of  a  tribe,  practising  some  form 
of  marriage,  to  spread  over  an  unoccupied  continent ;  they  would 
soon  split  up  into  distinct  hordes,  separated  from  each  other  by 
various  barriers,  and  still  more  effectually  by  the  incessant  wars 
between  all  barbarous  nations.  The  hordes  would  thus  be 
exposed  to  slightly  different  conditions  and  habits  of  life,  and 
would  sooner  or  later  come  to  differ  in  some  small  degree.  As 
soon  as  this  occurred,  each  isolated  tribe  would  form  for  itself  a 
slightly  different  standard  of  beauty ;  ^^  and  then  unconscious 
selection  would  come  into  action  through  the  more  powerful 
and. leading  men  preferring  certain  women  to  others.  Thus 
the  differences  between  the  tribes,  at  first  very  slight,  would 
gradually  and  inevitably  be  more  or  less  increased. 

With  animals  in  a  state  of  nature,  many  characters  proper  to 
the  males,  such  as  size,  strength,  special  weapons,  courage  and 
pugnacity,  have  been  acquired  through  the  law  of  battle.  The 
semi-human  progenitors  of  man,  like  their  allies  the  Quadrumana, 
will  almost  certainly  havS  been  thus  modified ;  and,  as  savages 
still  fight  for  the  possession  of  their  women,  a  similar  process  of 

'*  'The  Variation  of  Animals  and  French    artists,    that    the    idea    of 

Plants  under  Domestication,'  vol.  ii.  beauty   is  not  absolutely  the  same 

pp.  210-217.  even    throughout    Europe:  see  the 

'^  An    ingenious   writer    argues,  '  Lives   of  Haydn   and   Mozart,'  by 

from  a  comparison  of  the  pictures  Bombet     (otherwise      M.     Beyle), 

of   Raphael,    Rubens,    and    modern  English  translat.  p.  278. 


CiiAi'.  XX.      Man — Mode  of  Sexual  Selection.  597 

selection  has  probably  gone  on  in  a  greater  or  less  degree  to  the 
present  day.  Other  characters  proper  to  the  males  of  the  lower 
animals,  such  as  bright  colours  and  various  ornaments,  have  been 
acquired  by  the  more  attractive  males  having  been  preferred  by 
the  females.  There  are,  however,  exceptional  cases  in  which  the 
males  are  the  selecters,  instead  of  having  been  the  selected.  We 
recognise  such  cases  by  the  females  being  more  highly  orna- 
mented than  the  males, — their  ornamental  characters  having 
been  transmitted  exclusively  or  chiefly  to  their  female  offspring. 
One  such  case  has  been  described  in  the  order  to  which  man 
belongs,  that  of  the  Khesus  monkey. 

Man  is  more  powerful  in  body  and  mind  than  woman,  and  in 
the  savage  state  he  keeps  her  in  a  far  more  abject  state  of  bondage, 
than  does  the  male  of  any  other  animal ;  therefore  it  is  not  sur- 
prising that  he  should  have  gained  the  power  of  selection. 
Women  are  everywhere  conscious  of  the  value  of  their  o"uti  beauty ; 
and  when  they  have  the  means,  they  take  more  delight  in  deco- 
rating themselves  with  all  sorts  of  ornaments  than  do  men.  They 
borrow  the  plumes  of  male  birds,  with  which  nature  has  decked 
this  sex  in  order  to  charm  the  females.  As  women  have  long  been 
selected  for  beauty,  it  is  not  surprising  that  some  of  their  succes- 
sive variations  should  have  been  transmitted  exclusively  to  the 
same  sex;  consequently  that  they  should  have  transmitted 
beauty  in  a  somewhat  higher  degree  to  their  female  than  to  their 
male  offspring,  and  thus  have  become  more  beautiful,  according 
to  general  opinion,  than  men.  Women  however,  certainly 
transmit  most  of  their  characters,  including  some  beauty,  to 
their  offspring  of  both  sexes  ;  so  that  the  continued  preference  by 
the  men  of  each  race  for  the  more  attractive  women,  according  to 
their  standard  of  taste,  will  have  tended  to  modify  in  the  same 
manner  all  the  individuals  of  both  sexes  belonging  to  the  race. 

With  respect  to  the  other  form  of  sexual  selection  (which  with 
the  lower  animals  is  much  the  more  common),  namely,  when  the 
females  are  the  selecters,  and  accept  only  those  males  which 
excite  or  charm  them  most,  we  have  reason  to  believe  that  it 
formerly  acted  on  our  progenitors.  Man  in  all  probability  owes 
his  beard,  and  perhaps  some  other  characters,  to  inheritance 
from  an  ancient  progenitor  who  thus  gained  his  ornaments.  But 
this  form  of  selection  may  have  occasionally  acted  during  later 
times ;  for  in  utterly  barbarous  tribes  the  w^omen  have  more 
power  in  choosing,  rejecting,  and  tempting  their  lovers,  or  of 
afterwards  changing  their  husbands,  than  might  have  been 
expected.  As  this  is  a  point  of  some  importance,  I  will  give  in 
detail  such  evidence  as  I  have  been  able  to  collect. 

Hearne  describes  how  a  woman  in  one  of  the  tribes  of  Arctic. 


598  The  Descent  of  Man.  Part  III. 

America  repeatedly  ran  away  from  her  husband  and  joined  her 
lover  ;  and  with  the  Charruas  of  S.  America,  according  to  Azara, 
divorce  is  quite  optional.  Amongst  the  Abipones,  a  man  on 
choosing  a  wife,  bargains  with  the  parents  about  the  price.  But 
"it  frequently  happens  that  the  girl  rescinds  what  has  been 
"  agreed  upon  between  the  parents  and  the  bridegroom, 
"  obstinately  rejecting  the  very  mention  of  marriage."  She  often 
runs  away,  hides  herself,  and  thus  eludes  the  bridegroom. 
Captain  Musters  who  lived  with  the  Patagonians,  says  that  their 
marriages  are  always  settled  by  inclination;  "if  the  parents 
"  make  a  match  contrary  to  the  daughter's  will,  she  refuses  and 
"  is  never  compelled  to  comply."  In  Tierra  del  Fuego  a  young 
man  first  obtains  the  consent  of  the  parents  by  doing  them  some 
service,  and  then  he  attempts  to  carry  oif  the  girl ;  "  but  if  she  is 
"  unwilling,  she  hides  herself  in  the  woods  until  her  admirer  is 
"  heartily  tired  of  looking  for  her,  and  gives  up  the  pui'suit ;  but 
"  this  seldom  happens.''  In  the  Fiji  Islands  the  man  seizes  on 
the  woman  whom  he  wishes  for  his  wife  by  actual  or  jDretended 
force ;  but  "  on  reaching  the  home  of  her  abductor,  should  she  not 
"  approve  of  the  match,  she  runs  to  some  one  who  can  protect 
"  her ;  if,  however,  she  is  satisfied,  the  matter  is  settled  forthwith." 
With  the  Kalmucks  there  is  a  regular  race  between  the  bride  and 
bridegroom,  the  former  having  a  fair  start ;  and  Clarke  "  was 
"  assured  that  no  instance  occurs  of  a  girl  being  caught,  unless 
"  she  has  a  partiality  to  the  pursuer."  Amongst  the  wild  tribes 
of  the  Malay  Archipelago  there  is  also  a  racing  match;  and  it 
appears  from  M.  Bourien's  account,  as  Sir  J.  Lubbock  remarks, 
that  "  the  race  '  is  not  to  the  swift,  nor  the  battle  to  the  strong,' 
"but  to  the  young  man  who  has  the  good  fortune  to  please 
"  his  intended  bride."  A  similar  custom,  with  the  same  result, 
prevails  with  the  Koraks  of  North-Eastem  Asia. 

Turning  to  Africa :  the  Kafirs  buy  their  wives,  and  girls  are 
severely  beaten  by  their  fathers  if  they  will  not  accept  a  chosen 
husband  ;  but  it  is  manifest  from  many  facts  given  by  the  Kev. 
Mr.  Shooter,  that  they  have  considerable  power  of  choice.  Thus 
very  ugly,  though  rich  men,  have  been  known  to  fail  in  getting 
wives.  The  girls,  before  consenting  to  be  betrothed,  compel  the 
men  to  sliew  themselves  off  first  in  front  and  then  behind,  and 
"  exhibit  their  paces."  They  have  been  known  to  propose  to  a 
man,  and  they  not  rarely  run  away  with  a  favoured  lover.  So 
again,  Mr.  Leslie,  who  was  intimately  acquainted  with  the  Kafirs, 
says,  "  it  is  a  mistake  to  imagine  that  a  girl  is  sold  by  her  father 
"  in  the  same  manner,  and  with  the  same  authority,  with  which 
"  he  would  dispose  of  a  cow."  Amongst  the  degraded  Bush- 
men of  S.  Africa,  *'  when  a  girl  has  grown  up  to  womanhood 


Chap.  XX.       Ma?i — Mode  of  Sexual  Selection.  599 

"  without  having  been  betrothed,  which,  however,  does  not  often 
"  happen,  hor  lover  must  gain  her  approbation,  as  well  as  that  of 
"the  parents." -°  Mr,  Win  wood  Eeade  made  inquiries  for  me 
with  respect  to  the  negroes  of  Western  Africa,  and  he  informs 
me  that  "the  women,  at  least  among  the  more  intelligent  Pagan 
"  tribes,  have  no  difficulty  in  getting  the  husbands  whoiu  they 
"  may  desire,  although  it  is  considered  unwomanly  to  ask  a 
"  man  to  marry  them.  They  are  quite  capable  of  falling  in  love, 
"  and  of  forming  tender,  passionate,  and  faithful  attachments." 
Additional  cases  could  be  given. 

We  thus  see  that  with  savages  the  women  are  not  in  quite  so 
abject  a  state  in  relation  to  marriage,  as  has  often  lx?en  supposed. 
They  can  tempt  the  men  whom  they  pi'cfer,  and  can  sometimes 
reject  those  whom  they  dislike,  either  before  or  after  marriage. 
Preference  on  the  part  of  the  women,  steadily  acting  in  any  one 
direction,  would  ultimately  affect  the  character  of  the  tribe  ;  for 
the  women  would  generally  choose  not  merely  the  handsomest 
men,  according  to  their  standard  of  taste,  but  those  who  were  at 
the  same  time  best  able  to  defend  and  support  them.  Such  well- 
endowed  pairs  would  commonly  rear  a  larger  number  of  offspring 
than  the  less  favoured.  The  same  result  would  obviously  follow 
in  a  still  more  marked  manner,  if  there  was  selection  on  both  sides ; 
that  is  if  the  more  attractive,  and  at  the  same  time  more  powerful 
men  were  to  prefer,  and  were  preferred  by,  the  more  attractive 
women.  And  this  double  form  of  selection  seems  actually  to 
have  occurred,  especially  during  the  earlier  jDcriods  of  our  long 
history. 

We  will  now  examine  a  little  more  closely  some  of  the  charac- 
ters which  distinguish  the  several  races  of  man  from  one  another 
and  from  the  lower  animals,  namely,  the  greater  or  less  deficiency 
of  hair  on  the  body,  and  the  colour  of  the  skin.  We  need  say 
nothing  about  the  great  diversity  in  the  shape  of  the  features  and 
of  the  skull  between  the  different  races,  as  we  have  seen  in  the 
last  chapter  how  different  is  the  standard  of  beauty  in  these 

2"  Azara,  *  Voyages,' &c.  torn.  ii.  mitive  Marriage,' 1865,  p.  32.     On 

p.   23.     DobrJzhofi'er,  'An  Account  the  Malays,   Lubbock,   ibid.  p.   76. 

of  the  Abipones,'   vol.  ii.    1822,  p.  The  Rev.  J.  Shooter,  'On  the  Karirs  . 

207.     Capt.  Musters,  in  'Pvoc.    K.  of  Natal,'  1857,  pp.  52-60.     Mr.  D. 

Geograph.    Soc.,'    vol.     xv.    p.    47.  Leslie, 'Kafir  Character  and  Customs,' 

Williams  on  the  Fiji  Islanders,  as  1871,    p.  4.       On    the    Bush-men, 

quoted    by    Lubbock,     'Origin     of  Burchell,    'Travels    In    S.    Africa,' 

Civilisation,'   1870,  p.  79.     On  the  vol.  ii.  1824,  p.  59.     On  the  Koraks 

Fuegians,  King  and  FitzRoy,  '  Voy-  by   McKcnnan,   as    quoted    by    Mr. 

ages  of  the  Adventure  and  Beagle,'  Wake,  in  '  Anthropologia,' Oct.  1873, 

vol.  ii.  1839,  p.   182.     On  the  Kal-  p.  75. 
mucks,  quoted  by  M'Lennan,  '  Pri- 


6oo  The  Descent  of  Man.  Part  HI. 

respects.  These  characters  will  therefore  probably  have  been 
acted  on  through  sexual  selection ;  but  we  have  no  means  of 
judging  whether  they  have  been  acted  on  chiefly  from  the  male 
or  female  side.  The  musical  faculties  of  man  have  likewise  been 
already  discussed. 

Absetice  of  Hair  on  the  Body,  and  its  Development  on  the  Face 
and  Head.— From  the  presence  of  the  woolly  hair  or  lanugo  on 
the  human  foetus,  and  of  rudimentary  hairs  scattered  over  the 
body  during  maturity,  we  may  infer  that  man  is  descended  from 
some  animal  which  was  born  hairy  and  remained  so  during  life. 
The  loss  of  hair  is  an  inconvenience  and  probably  an  injury  to  man, 
even  in  a  hot  climate,  for  he  is  thus  exposed  to  the  scorching 
of  the  sun,  and  to  sudden  chills,  especially  during  wet  weather. 
As  Mr.  Wallace  remarks,  the  natives  in  all  countries  are  glad  to 
protect  their  naked  backs  and  shoulders  with  some  slight  covering. 
No  one  supposes  that  the  nakedness  of  the  skin  is  any  direct 
advantage  to  man ;  his  body  therefore  cannot  have  been  divested 
of  hair  through  natural  selection.  ^^  Nor,  as  shewn  in  a  former 
chapter,  have  we  any  evidence  that  this  can  be  due  to  the 
direct  action  of  climate,  or  that  it  is  the  result  of  correlated 
development. 

The  absence  of  hair  on  the  body  is  to  a  certain  extent  a 
secondary  sexual  character ;  for  in  all  parts  of  the  world  women 
are  less  hairy  than  men.  Therefore  we  may  reasonably  suspect 
that  this  character  has  been  gained  through  sexual  selection. 
We  know  that  the  faces  of  several  species  of  monkeys,  and  large 
surfaces  at  the  posterior  end  of  the  body  of  other  species,  have 
been  denuded  of  hair ;  and  this  we  may  safely  attribute  to  sexual 
selection,  for  these  surfaces  are  not  only  vividly  coloured,  but  some- 
times, as  with  the  male  mandrill  and  female  rhesus,  much  more 
vividly  in  the  one  sex  than  in  the  other,  especially  during  the 
breeding-season.  I  am  informed  by  Mr.  Bartlett  that,  as  these 
animals  gradually  reach  maturity,  the  naked  surfaces  grow 
larger  compared  with  the  size  of  their  bodies.  The  hair,  how- 
ever, appears  to  have  been  removed,  not  for  the  sake  of  nudity, 
but  that  the  colour  of  the  skin  may  be  more  fully  displayed.    So 

^^  'Contributions  to   the   Theory  A'iew  ('Transactions  of  Devonshire 

of  Natural   Selection,'  1870,  p.  346.  Assoc,  for  Science,'  1870)  remarks, 

Mr.  Wallace  believes  (p.  350)  "  that  that   had    Mr.   Wallace  ''  employed 

"  some  intelligent  power  has  guided  "his  usual  ingenuity  on  the  ques- 

"  or  determined  the  development  of  "  tion  of    man's    hairless    skin,   he 

"  man  ;  "  and  he  considers  the  hair-  "  might  have  seen  the  possibility  of 

less  condition  of  the  skin  as  coming  "  its  selection  through  its  superior 

under  this   head.     The   Rev.   T.   K.  "  beauty  or  the  health  attaching  to 

Stebbing,   in  commenting    on    this  "  superior  cleanliness." 


Chai'.  XX.  HI  an — Absence  of  Hair.  6oi 

again  with  many  birds,  it  appears  as  if  the  head  and  neck  had 
been  divested  of  feathers  through  sexual  selection,  to  exliibit  the 
brightly-coloured  skin. 

As  the  body  in  woman  is  less  hairy  than  in  man,  and  as  this 
character  is  common  to  all  races,  we  may  conclude  that  it  was 
our  female  semi-human  ancestors  who  were  first  divested  of  hair, 
and  that  this  occurred  at  an  extremely  remote  i^eriod  before  the 
several  races  had  diverged  from  a  common  stock.  Whilst  our 
female  ancestors  were  gradually  acquiring  this  new  character  of 
nudity,  they  must  have  transmitted  it  almost  equally  to  their 
offspring  of  both  sexes  whilst  young ;  so  that  its  transmission, 
as  with  the  ornaments  of  many  mammals  and  birds,  has  not  been 
limited  either  by  sex  or  age.  There  is  nothing  surprising  in  a 
partial  loss  of  hair  having  been  esteemed  as  an  ornament  by  our 
ape-like  progenitors,  for  we  have  seen  that  innumerable  strange 
characters  have  been  thus  esteemed  by  animals  of  all  kinds,  and 
have  consequently  been  gained  through  sexual  selection.  Nor 
is  it  surprising  that  a  slightly  injurious  character  should  have 
been  thus  acquired  ;  for  we  know  that  this  is  the  case  with  the 
plumes  of  certain  birds,  and  with  the  horns  of  certain  stags. 

The  females  of  some  of  the  anthropoid  apes,  as  stated  in  a 
former  chapter,  are  somewhat  less  hairy  on  the  under  surface 
than  the  males ;  and  here  we  have  what  might  have  afforded  a 
commencement  for  the  process  of  denudation.  With  respect  to 
the  completion  of  the  process  through  sexual  selection,  it  is  well 
to  bear  in  mind  the  New  Zealand  proverb,  "  There  is  no  woman 
"  for  a  hairy  man."  All  who  have  seen  photographs  of  the 
Siamese  hairy  family  will  admit  how  ludicrously  hideous  is  the 
opposite  extreme  of  excessive  hairiness.  And  the  king  of  Siam 
had  to  bribe  a  man  to  marry  the  first  haary  woman  in  the 
family;  and  she  transmitted  this  character  to  her  young  off- 
spring of  both  sexes.  ^^ 

Some  races  are  much  more  hairy  than  others,  especially  the 
males ;  but  it  must  not  be  assumed  that  the  more  hairy  races, 
such  as  the  European,  have  retained  their  primordial  condition 
more  completely  than  the  naked  races,  such  as  the  Kalmucks 
or  Americans.  It  is  more  probable  that  the  hairiness  of  the 
former  is  due  to  partial  reversion;  for  characters  which  have 
been  at  some  former  period  long  inherited,  are  always  apt  to 
return.  We  have  seen  that  idiots  are  often  very  hairy,  and  they 
are  apt  to  revert  in  other  characters  to  a  lower  animal  type.  It 
does  not  appear  that  a  cold  climate  has  been  influential  in 
leading  to  this  kind  of  reversion ;  excepting  perhai^s  with  the 

^  'The  Variation  of  Animals  and  Plants  under  Domestication,' vol  ii 
1868,  p.  327. 


6o2  The  Descent  of  Man.  Tart  III. 

negroes,  who  have  been  reared  during  several  generations  in 
the  United  States/^  and  possibly  with  the  Ainos,  who  inhabit 
the  northern  islands  of  the  Japan  archipelago.  But  the  laws  of 
inheritance  are  so  complex  than  we  can  seldom  understand  their 
action.  If  the  greater  hairiness  of  certain  races  be  the  result  of 
reversion,  unchecked  by  any  form  of  selection,  its  extreme 
variability,  even  within  the  limits  of  the  same  race,  ceases  to  be 
remarkable.^* 

With  respect  to  the  beard  in  man,  if  we  turn  to  our  best  guide, 
the  Quadrumana,  we  find  beards  equally  developed  in  both 
sexes  of  many  species,  but  in  some,  either  confined  to  the  males, 
or  more  developed  in  them  than  in  the  females.  From  this  fact 
and  from  the  curious  arrangement,  as  well  as  the  bright  colours 
of  the  hair  about  the  heads  of  many  monkeys,  it  is  highly 
probable,  as  before  explained,  that  the  males  first  acquired  their 
beards  through  sexual  selection  as  an  ornament,  transmitting 
them  in  most  cases,  equally  or  nearly  so,  to  their  offspring  of 
both  sexes.  We  know  from  Eschricht^^  that  with  mankind,  the 
female  as  well  as  the  male  foetus  is  furnished  with  much  hair  on 
the  face,  especially  round  the  mouth;  and  this  indicates  that 
we  are  descended  from  progenitors,  of  whom  both  sexes  were 
bearded.  It  appears  therefore  at  first  sight  probable  that  man 
has  retained  his  beard  from  a  very  early  period,  whilst  woman 
lost  her  beard  at  the  same  time  that  her  body  became  almost 
completely  divested  of  hair.  Even  the  colour  of  our  beards  seems 
to  have  been  inherited  from  an  ape-like  progenitor ;  for  when 

2'  '  Investigations    into    Military  races    of  man  would  be   eminently 

and    Anthropological    Statistics    of  liable   to   revert   to  the  primordial 

American  Soldiers,'  by  B.  A.  Gould,  hairy  character  of  their  early  ape- 

1869  ;  p.  568  : — Observations  were  like  progenitors, 

carefully  made   on  the  hairiness   of  "^^  Hardly  any  view  advanced  in 

2129  black    and  coloured    soldiers,  this  work  has    met  with  so  much 

whilst  they  were  bathing  ;  and  by  disfavour  (see  for  instance,  Spengel, 

looking  to  the  published  table,  "  it  '  Die  Fortschritte  des  Darwinismus,' 

"  is  manifest  at  a  glance  that  there  1874,  p.  80)  as  the  above  explana- 

"  is  but  little,  if  any,  difference  be-  tion  of  the  loss  of  hair  in  mankind 

"  tween   the   white   and   the   black  through  sexual  selection;  but  none 

"  races  in  this  respect."     It  is,  how-  of  the  opposed  arguments  seem  to 

ever,  certain  that  negroes  in  their  na-  me  of  much  weight,  in  comparison 

tive  and  much  hotter  land  of  Africa,  with    the   facts    shewing    that    the 

have  remarkably  smooth  bodies.     It  nudity  of  the  skin  is  to  a  certain 

should    be    particularly    observed,  extent  a  secondary  sexual  character 

that    both    pure    blacks    and    mu-  in  man  and  in  some  of  the  Quad- 

lattoes  were  included  in  the  above  rumana. 

enumeration ;  and  this  is  an  unfor-  -^    '  Ueber      die     Richtung     der 

tunate  circumstance,  as  in  accordance  Haare    am    Menschlicheu    Korper,* 

with  a  principle,  the  truth  of  which  in  dialler's  'Archiv  fur  Anat.  und 

I    have    elsewhere    proved,   crossed  Phys.'  1837,  s.  40. 


Chap.  XX.  Beards,  603 


there  is  any  difference  in  tint  between  the  hair  of  the  head  and 
the  beard,  the  latter  is  lighter  coloured  in  all  monkeys  and  in 
man.  In  those  Quadrumana  in  which  the  male  has  a  larger 
beard  than  that  of  the  female,  it  is  fully  developed  only  at 
maturity,  just  as  with  mankind ;  and  it  is  possible  that  only  the 
later  stages  of  develoijment  have  been  retained  by  man.  In 
opposition  to  this  view  of  the  retention  of  the  beard  from  an 
early  period,  is  the  fact  of  its  great  variability  in  different  races, 
and  even  within  the  same  race;  for  this  indicates  reversion, — 
long  lost  characters  being  very  apt  to  vary  on  re-appearance. 

Nor  must  we  overlook  the  part  which  sexual  selection  may 
have  played  in  later  times  ;  for  we  know  that  with  savages,  the 
men  of  the  beardless  races  take  infinite  pains  in  eradicating  every 
hair  from  their  faces  as  something  odious,  whilst  the  men  of 
the  bearded  races  feel  the  greatest  jnide  in  their  beards.  Tlie 
women,  no  doubt,  participate  in  these  feelings,  and  if  so  sexual 
selection  can  hardly  have  failed  to  have  effected  something  in  the 
course  of  later  times.  It  is  also  possible  that  the  long-continued 
habit  of  eradicating  the  hair  may  have  produced  an  inherited 
effect.  Dr.  Brown-Sequard  has  shewn  that  if  certain  animals  are 
operated  on  in  a  particular  manner,  their  offspring  are  affected. 
Further  evidence  could  be  given  of  the  inheritance  of  the  effects 
of  mutilations;  but  a  fact  lately  ascertained  by  Mr.  Salvin^"^ 
has  a  more  direct  bearing  on  the  present  question ;  for  he  has 
shewn  that  the  motmots,  which  are  known  habitually  to  bite  off 
the  barbs  of  the  two  central  tail-feathers,  have  the  barbs  of  these 
feathers  naturally  somewhat  reduced.^''^  Nevertheless  with  man- 
kind, the  habit  of  eradicating  the  beard  and  the  hairs  on  the 
body  would  probably  not  have  arisen  until  these  had  already 
become  by  some  means  reduced. 

It  is  difficult  to  form  any  judgment  as  to  how  the  hair  on  the 
head  became  developed  to  its  present  great  length  in  many  races. 
Eschricht-*  states  that  in  the  human  foetus  the  hair  on  the  face 
during  the  fifth  month  is  longer  than  that  on  the  head ;  and 
this  indicates  that  our  semi-human  progenitors  were  not 
furnished  with  long  tresses,  which  must  therefore  have  been 
a  late  acquisition.  This  is  likewise  indicated  by  the  extra- 
ordinary difference  in  the  length  of  the  hair  in  the  different 
races ;  in  the  negro  the  hair  forms  a  mere  curly  mat ;  with  us 

^^  'Oa  the  tail-feathers  of  Morao-  Some      distinguished     ethnologists, 

tus'   '  Proc.  Zoolog.  Soc.,'   1873,  p.  amongst  others  M.  Gosse  of  Geneva, 

429.  believe  that  artificial  modifications 

2^   ^Mr.     Sproat     has    suggested  of  the  skull  tend  to  be  inherited. 
('Scenes    and    Studies    of     Savage  28  '  Ugber  die  Richtung,'  ibid.  ». 

Life,'  1868,  p.  25)  this  same  view,  40. 


6o4  The  Descent  of  Man.  Pabt  III. 

it  is  of  great  length,  and  with  the  American  natives  it  not  rarely 
reaches  to  the  ground.  Some  species  of  Semnopithecus  have 
their  heads  covered  with  moderately  long  hair,  and  this  probably 
serves  as  an  ornament  and  was  acquired  through  sexual  selection. 
The  same  view  may  perhaps  be  extended  to  mankind,  for  we 
know  that  long  tresses  are  now  and  were  formerly  much  admired, 
as  may  be  observed  in  the  works  of  almost  every  poet ;  St.  Paul 
says,  "  if  a  woman  have  long  hair,  it  is  a  glory  to  her ;"  and  we 
have  seen  that  in  North  America  a  chief  was  elected  solely  from 
the  length  of  his  hair. 

Colour  of  the  Skin. — The  best  kind  of  evidence  that  in  man  the 
colour  of  the  skin  has  been  modified  through  sexual  selection  is 
scanty ;  for  in  most  races  the  sexes  do  not  differ  in  this  respect, 
and  only  slightly,  as  we  have  seen,  in  others.  We  know,  however, 
from  the  many  facts  already  given  that  the  colour  of  the  skin  is 
regarded  by  the  men  of  all  races  as  a  highly  important  element 
in  their  beauty ;  so  that  it  is  a  character  which  would  be  likely 
to  have  been  modified  through  selection,  as  has  occurred  in 
innumerable  instances  with  the  lower  animals.  It  seems  at  first 
sight  a  monstrous  supposition  that  the  jet-blackness  of  the  negro 
should  have  been  gained  through  sexual  selection ;  but  this  view 
is  supported  by  various  analogies,  and  we  know  that  negroes 
admire  their  own  colour.  "With  mammals,  when  the  sexes 
differ  in  colour,  the  male  is  often  black  or  much  darker  than 
the  female ;  and  it  depends  merely  on  the  form  of  inheritance 
Avhether  this  or  any  other  tint  is  transmitted  to  both  sexes  or  to 
one  alone.  The  resemblance  to  a  negro  in  minature  of  Pithecia 
satunas  with  his  jet  black  skin,  white  rolling  eyeballs,  and  hair 
parted  on  the  top  of  the  head,  is  almost  ludicrous. 

The  colour  of  the  face  differs  much  more  widely  in  the  various 
kinds  of  monkeys  than  it  does  in  the  races  of  man  ;  and  we  have 
some  reason  to  believe  that  the  red,  blue,  orange,  almost  white 
and  black  tints  of  their  skin,  even  when  common  to  both  sexes, 
as  well  as  the  bright  colours  of  their  fur,  and  the  ornamental 
tufts  about  the  head,  have  all  been  acquired  through  sexual 
selection.  As  the  order  of  development  during  growth,  generally 
indicates  the  order  in  which  the  characters  of  a  species  have 
been  developed  and  modified  during  j^revious  generations ;  and 
as  the  newly-born  infants  of  the  various  races  of  man  do  not 
differ  nearly  as  much  in  colour  as  do  the  adults,  although  their 
bodies  are  as  completely  destitute  of  hair,  we  have  some  slight 
evidence  that  the  tints  of  the  different  races  were  acquired  at  a 
period  subsequent  to  the  removal  of  the  hair,  which  must  have 
occurred  at  a  very  early  period  in  the  history  of  man. 


Ckap.  XX.  Sunnnary  on  Man.  605 

Summary, — Wc  EQcay  conclude  that  the  greater  size,  strength, 
courage,  pugnacity,  and  energy  of  man,  in  comparison  with 
woman,  were  acquired  during  primeval  times,  and  have  subse- 
quently been  augmented,  chiefly  through  the  contests  of  rival 
males  for  the  possession  of  the  females.  The  greater  intellectual 
vigour  and  power  of  invention  in  man  is  probably  due  to 
natural  selection,  combined  with  the  inherited  effects  of  habit, 
for  the  most  able  men  will  have  succeeded  best  in  defending  and 
providing  for  themselves  and  for  their  wives  and  offspring.  As 
far  as  the  extreme  intricacy  of  the  subject  permits  us  to  judge, 
it  appears  that  our  male  ape-like  progenitors  acquired  their 
beards  as  an  ornament  to  charm  or  excite  the  opjjosite  sex,  and 
transmitted  them  only  to  their  male  offspring.  The  females 
apparently  first  had  their  bodies  denuded  of  hair,  also  as  a 
sexual  ornament;  but  they  transmitted  this  character  almost 
equally  to  both  sexes.  It  is  not  improbable  that  the  females 
were  modified  in  other  respects  for  the  same  purpose  and  by  the 
same  means ;  so  that  women  have  acquired  sweeter  voices  and 
become  more  beautiful  than  men. 

It  deserves  attention  that  with  mankind  the  conditions  were 
in  many  respects  much  more  favourable  for  sexual  selection, 
during  a  very  early  period,  when  man  had  only  just  attained 
to  the  rank  of  manhood,  than  during  later  times.  For  he  would 
then,  as  we  may  safely  conclude,  have  been  guided  more  by  his 
instinctive  passions,  and  less  by  foresight  or  reason.  He  would 
have  jealously  guarded  his  wife  or  wives.  He  would  not  have 
practised  infanticide ;  nor  valued  his  wives  merely  as  useful 
slaves  ;  nor  have  been  betrothed  to  them  during  infancy.  Hence 
we  may  infer  that  the  races  of  men  were  differentiated,  as  far 
as  sexual  selection  is  concerned,  in  chief  part  at  a  very  remote 
epoch ;  and  this  conclusion  throws  light  on  the  remarkable  fact 
that  at  the  most  ancient  period,  of  which  we  have  as  yet  any 
record,  the  races  of  man  had  already  come  to  differ  nearly  or 
quite  as  much  as  they  do  at  the  present  day. 

The  views  here  advanced,  on  the  part  which  sexual  selection 
has  played  in  the  history  of  man,  want  scientific  precision.  He 
who  does  not  admit  this  agency  in  the  case  of  the  lower  animals, 
will  disregard  all  that  I  have  written  in  the  later  chapters 
on  man.  We  cannot  positively  say  that  this  character,  but  not 
that,  has  been  thus  modified  ;  it  has,  however,  been  shewn  that 
the  races  of  man  difibr  from  each  other  and  from  their  nearest 
allies,  in  certain  characters  which  are  of  no  service  to  them  in 
their  daily  habits  of  life,  and  which  it  is  extremely  probable 
would  have  been  modified  through  sexual  selection.  We  have 
seen  that  with  the  lowest  savages  the  people  of  each  tribe  admire 


6o6  The  Descent  of  Man.  Part  III. 

their  own  cliaracteristic  qualities, — the  shape  of  the  head  and 
face,  the  squareness  of  the  cheek-bones,  the  prominence  or 
depression  of  the  nose,  the  colour  of  the  skin,  the  length  of  the 
hair  on  the  head,  the  absence  of  hair  on  the  face  and  body,  or 
the  presence  of  a  great  beard,  and  so  forth.  Hence  these  and 
other  such  points  could  hardly  fail  to  be  slowly  and  gradually 
exaggerated,  from  the  more  powerful  and  able  men  in  each  tribe, 
who  would  succeed  in  rearing  the  largest  number  of  offspring, 
having  selected  during  many  generations  for  their  wives  the 
most  strongly  characterised  and  therefore  most  attractive 
women.  For  my  own  part  I  conclude  that  of  all  the  causes 
which  have  led  to  the  differences  in  external  appearance  between 
the  races  of  man,  and  to  a  certain  extent  between  man  and  the 
lower  animals,  sexual  selection  has  been  the  most  efficient. 


CHAPTEE  XXL 

General  Summary  and  Conclusion. 

Main  conclusion  that  man  is  descended  from  some  lower  form — Manner  of 
development — Genealogy  of  man — Intellectual  and  moral  faculties — ■ 
Sexual  selection — Concluding  remarks. 

A  BRIEF  summary  will  be  sufficient  to  recall  to  the  reader's  mind 
the  more  salient  points  in  this  work.  Many  of  the  views  which 
have  been  advanced  are  highly  speculative,  and  some  no  doubt 
will  prove  erroneous ;  but  I  have  in  every  case  given  the  reasons 
which  have  led  me  to  one  view  rather  than  to  another.  It 
seemed  worth  while  to  try  how  far  the  principle  of  evolution 
would  throw  light  on  some  of  the  more  complex  problems  in  the 
natural  history  of  man.  False  facts  are  highly  injurious  to  the 
progress  of  science,  for  they  often  endure  long ;  but  false  views, 
if  supported  by  some  evidence,  do  little  harm,  for  every  one 
takes  a  salutary  pleasure  in  proving  their  falseness ;  and  when 
this  is  done,  one  path  towards  error  is  closed  and  the  road  to 
truth  is  often  at  the  same  time  opened. 

The  main  conclusion  here  arrived  at,  and  now  held  by  many 
naturahsts  who  are  well  competent  to  form  a  sound  judgment, 
is  that  man  is  descended  from  some  less  highly  organised  form. 
The  grounds  upon  which  this  conclusion  rests  will  never  be 
shaken,  for  the  close  similarity  between  man  and  the  lower 
animals  in  embryonic  development,  as  well  as  in  innumerable 
points  of  structure  and  constitution,  both  of  high  and  of  the 
most  trifling  importance, — the  rudiments  which  he  retains,  and 
the  abnormal  reversions  to  which  he  is  occasionally  liable, — 


Chap.  XXI.  General  Suniniary.  607 

are  facts  which  cannot  be  disputed.  They  have  long  been 
known,  but  until  recently  they  told  ns  nothing  with  respect  to 
the  origin  of  man.  Now  when  viewed  by  the  light  of  onr  know- 
ledge of  the  whole  organic  world,  their  meaning  is  unmistakable. 
The  great  principle  of  evolution  stands  up  clear  and  firm,  when 
these  groups  of  facts  are  considered  in  connection  with  others, 
such  as  the  mutual  affinities  of  the  members  of  the  same  group, 
their  geographical  distribution  in  past  and  present  times,  and 
their  geological  succession.  It  is  incredible  that  all  these  facts 
should  speak  falsely.  He  who  is  not  content  to  loolc,  like  a 
savage,  at  the  phenomena  of  nature  as  disconnected,  cannot  any 
longer  believe  that  man  is  the  work  of  a  separate  act  of  creation. 
He  will  be  forced  to  admit  that  the  close  resemblance  of  the 
embryo  of  man  to  that,  for  instance,  of  a  dog — the  construction 
of  his  skull,  limbs  and  whole  frame  on  the  same  plan  with  that  of 
other  mammals,  independently  of  the  uses  to  which  the  parts 
may  be  put— the  occasional  re-appearance  of  various  structures, 
for  instance  of  several  muscles,  which  man  does  not  normally 
possess,  but  which  are  common  to  the  Quadrumana — and  a 
crowd  of  analogous  facts— all  point  in  the  plainest  manner  to  the 
conclusion  that  man  is  the  co- descendant  with  other  mammals 
of  a  common  progenitor. 

We  have  seen  that  man  incessantly  presents  individual  differ- 
ences in  all  parts  of  his  body  and  in  his  mental  faculties.  These 
differences  or  variations  seem  to  be  induced  by  the  same  general 
causes,  and  to  obey  the  same  laws  as  with  the  the  lower  animals. 
In  both  cases  similar  laws  of  inheritance  prevail.  Man  tends  to 
increase  at  a  greater  rate  than  his  means  of  subsistence;  con- 
sequently he  is  occasionally  subjected  to  a  severe  struggle  for 
existence,  and  natural  selection  will  have  effected  whatever  lies 
within  its  scope.  A  succession  of  strongly-marked  variations  of 
a  similar  nature  is  by  no  means  requisite ;  slight  fluctuating 
differences  in  the  individual  suffice  for  the  work  of  natural  selec- 
tion ;  not  that  we  have  any  reason  to  suppose  that  in  the  same 
species,  all  parts  of  the  organisation  tend  to  vary  to  the  same 
degree.  We  may  feel  assured  that  the  inherited  effects  of  the 
long-continued  use  or  disuse  of  parts  will  have  done  much  in 
the  same  direction  with  natural  selection.  Modifications  formerly 
of  importance,  though  no  longer  of  any  special  use,  are  long- 
inherited.  When  one  part  is  modified,  other  parts  change 
through  the  principle  of  correlation,  of  which  we  have  instances 
in  many  curious  cases  of  correlated  monstrosities.  Something 
may  be  attributed  to  the  direct  and  definite  action  of  the 
sm-rounding  conditions  of  life,  such  as  abundant  food,  heat  or 
moisture;  and   lastly,  many  characters  of  slight  physiological 


6o8  The  Descent  of  Man.  Part  III. 

importance,  some  indeed  of  considerable  importance,  have  been 
gained  through  sexnal  selection. 

No  doubt  man,  as  well  as  every  other  animal,  presents 
structures,  which  seem  to  our  limited  knowledge,  not  to  be  now 
of  any  service  to  him,  nor  to  have  been  so  formerly,  either  for  the 
general  conditions  of  life,  or  in  the  relations  of  one  sex  to  the 
other.  Such  structures  cannot  be  accounted  for  by  any  form  of 
selection,  or  by  the  inherited  effects  of  the  use  and  disuse  of 
parts.  We  know,  however,  that  many  strange  and  strongly- 
marked  peculiarites  of  structure  occasionally  appear  in  our 
domesticated  productions,  and  if  their  unknown  causes  were  to 
act  more  uniformly,  they  would  probably  become  common  to  all 
the  individuals  of  the  species.  We  may  hope  hereafter  to 
understand  something  about  the  causes  of  such  occasional  modi- 
fications, especially  through  the  study  of  monstrosities :  hence 
the  labours  of  experimentalists,  such  as  those  of  M.  Camille 
Dareste,  are  full  of  promise  for  the  future.  In  general  we  can 
only  say  that  the  cause  of  each  slight  variation  and  of  each 
monstrosity  lies  much  more  in  the  constitution  of  the  organism, 
than  in  the  nature  of  the  surrounding  conditions ;  though  new 
and  changed  conditions  certainly  play  an  important  part  in 
exciting  organic  changes  of  many  kinds. 

Through  the  means  just  specified,  aided  perhaps  by  others 
as  yet  undiscovered,  man  has  been  raised  to  his  present  state. 
But  since  he  attained  to  the  rank  of  manhood,  he  has  diverged 
into  distinct  races,  or  as  they  may  be  more  fitly  called,  sub- 
sjDecies.  Some  of  these,  such  as  the  Negro  and  European,  are 
so  distinct  that,  if  specimens  had  been  brought  to  a  naturalist 
without  any  further  information,  they  would  undoubtedly  have 
been  considered  by  him  as  good  and  true  species.  Nevertheless 
all  the  races  agree  in  so  many  unimportant  details  of  structure 
and  in  so  many  mental  peculiarities,  that  these  can  be  accounted 
for  only  by  inheritance  from  a  common  progenitor ;  and  a  pro- 
genitor thus  characterised  would  probably  deserve  to  rank  as  man. 

It  must  not  be  supposed  that  the  divergence  of  each  race  from 
the  other  races,  and  of  all  from  a  common  stock,  can  be  traced 
back  to  any  one  pair  of  progenitors.  On  the  contrary,  at  every 
stage  in  the  process  of  modification,  all  the  individuals  which 
were  in  any  way  better  fitted  for  their  conditions  of  life,  though 
in  different  degrees,  would  have  survived  in  greater  numbers 
than  the  less  well-fitted.  The  process  would  have  been  like  that 
followed  by  man,  when  he  does  not  intentionally  select  particular 
individuals,  but  breeds  from  all  the  superior  individuals,  and 
neglects  the  inferior.  He  thus  slowly  but  surely  modifies  his 
stock,  and  unconsciouslv  forms  a  new  strain.    So  with  respect 


Chap.  XXI.  General  Summary.  609 


to  modifications  acquired  independently  of  selection,  and  due 
to  variations  arising  from  the  nature  of  the  organism  and  the 
action  of  the  surrounding  conditions,  or  from  changed  habits  of 
life,  no  single  pair  will  have  been  modified  much  more  than  the 
other  pairs  inhabiting  the  same  country,  for  all  -will  have  been 
continually  blended  through  free  intercrossing. 

By  considering  the  embryological  structure  of  man, — the 
homologies  which  he  presents  with  the  lower  animals,— the 
rudiments  which  he  retains,— and  the  reversions  to  which  he  is 
liable,  we  can  partly  recall  in  imagination  the  former  condition 
of  our  early  progenitors ;  and  can  approximately  place  them  in 
their  proper  place  in  the  zoological  series.  We  thus  learn  that 
man  is  descended  from  a  hairy,  tailed  quadruped,  probably 
arboreal  in  its  habits,  and  an  inhabitant  of  the  Old  World. 
This  creature,  if  its  whole  structure  had  been  examined  by  a 
naturalist,  would  have  been  classed  amongst  the  Quadrumana, 
as  surely  as  the  still  more  ancient  progenitor  of  the  Old  and 
New  World  monkeys.  The  Quadrumana  and  all  the  higher 
mammals  are  probably  derived  from  an  ancient  marsupial 
animal,  and  this  through  a  long  line  of  diversified  forms,  from 
some  amphibian-like  creature,  and  this  again  from  some  fish-like 
animal.  In  the  dim  obscurity  of  the  past  we  can  see  that  the 
early  progenitor  of  all  the  Vertebrata  must  have  been  an  aquatic 
animal,  provided  with  branchiae,  with  the  two  sexes  united  in 
the  same  individual,  and  with  the  most  important  organs  of  the 
body  (such  as  the  brain  and  heart)  imperfectly  or  not  at  all 
developed.  This  animal  seems  to  have  been  more  like  the  larvae 
of  the  existing  marine  Ascidians  than  any  other  known  form. 

The  high  standard  of  our  intellectual  powers  and  moral  dis- 
position is  the  gTcatest  difficulty  which  presents  itself,  after  we 
have  been  driven  to  this  conclusion  on  the  origin  of  man.  But 
every  one  who  admits  the  principle  of  evolution,  must  see  that 
the  mental  powers  of  the  higher  animals,  which  are  the  same  in 
kind  with  those  of  man,  though  so  different  in  degree,  are 
capable  of  advancement.  Thus  the  interval  between  the  mental 
powers  of  one  of  the  higher  apes  and  of  a  fish,  or  between  those 
of  an  ant  and  scale-insect,  is  immense ;  yet  their  development 
does  not  offer  any  special  difiiculty ;  for  with  our  domesticated 
animals,  the  mental  faculties  are  certainly  variable,  and  the 
variations  are  inherited.  No  one  doubts  that  they  'are  of  the 
utmost  importance  to  animals  in  a  state  of  nature.  Therefore 
the  conditions  are  favourable  for  their  development  through 
natural  selection.  The  same  conclusion  may  be  extended  to  man ; 
the  intellect  must  have  been  all-important  to  him,  even  at  a  very 
27 


6 10  The  Descent  of  Man.  Pakt  III. 

remote  period,  as  enabling  liim  to  invent  and  use  language,  to 
make  weapons,  tools,  trajDS,  &c.,  whereby  with  the  aid  of  his 
social  habits,  he  long  ago  became  the  most  dominant  of  all  living 
creatures, 

A  great  stride  in  the  development  of  the  intellect  will  have 
followed,  as  soon  as  the  half-art  and  half-instinct  of  language 
came  into  use ;  for  the  continued  use  of  language  will  have 
reacted  on  the  brain  and  produced  an  inherited  effect ;  and  this 
again  will  have  reacted  on  the  improvement  of  language.  As  Mr. 
Chauncey  Wright^  has  well  remarked,  the  largeness  of  the  brain 
in  man  relatively  to  his  body,  compared  with  the  lower  animals, 
may  be  attributed  in  chief  part  to  the  early  use  of  some  simple 
form  of  language, — that  wonderful  engine  which  affixes  signs  to 
all  sorts  of  objects  and  qualities,  and  excites  trains  of  thought 
which  would  never  arise  from  the  mere  impression  of  the 
senses,  or  if  they  did  arise  could  not  be  followed  out.  The 
higher  intellectual  powers  of  man,  such  as  those  of  ratiocination, 
abstraction,  self-consciousness,  &c.,  probably  follow  from  the  con- 
tinued improvement  and  exercise  of  the  other  mental  faculties. 

The  development  of  the  moral  qualities  is  a  more  interesting 
problem.  The  foundation  lies  in  the  social  instincts,  including 
under  this  term  the  family  ties.  These  instincts  are  highly 
complex,  and  in  the  case  of  the  lower  animals  give  special 
tendencies  towards  certain  definite  actions;  but  the  more  im- 
portant elements  are  love,  and  the  distinct  emotion  of  sympathy. 
Animals  endowed  with  the  social  instincts  take  pleasure  in  one 
another's  company,  warn  one  another  of  danger,  defend  and  aid 
one  another  in  many  ways.  These  instincts  do  not  extend  to  all 
the  individuals  of  the  species,  but  only  to  those  of  the  same 
community.  As  they  are  highly  beneficial  to  the  species,  they 
have  in  all  probability  been  acquired  through  natural  selection. 

A  moral  being  is  one  who  is  capable  of  reflecting  on  his 
past  actions  and  their  motives — of  approving  of  some  and 
disapproving  of  others ;  and  the  fact  that  man  is  the  one  being 
who  certainly  deserves  this  designation,  is  the  greatest  of  all 
distinctions  between  him  and  the  lower  animals.  But  in  the 
fourth  chapter  I  have  endeavoured  to  shew  that  the  moral  sense 
follows,  firstly,  from  the  enduring  and  ever-present  nature  of  the 
social  instincts ;  secondly,  from  man's  appreciation  of  the  appro- 
bation and  disapprobation  of  his  fellows ;  and  thirdly,  from  the 
high  activity  of  his  mental  faculties,  with  past  impressions  ex- 
tremely vivid ;  and  in  these  latter  respects  he  difiers  from  the 
lower  animals.    Owing  to  this  condition  of  mind,  man  cannot 

*  *  On  the  Limits  of  Natural  Selection,*  in  j;he  *  North  American 
Review,'  Oct.  1870,  p.  295. 


CiiAP.  XXr.  General  Suifiinary.  6ii 

avoid  looking  both  backwards  and  forwards,  and  comparing 
past  impressions.  Hence  after  some  temporary  desire  or  passion 
has  mastered  his  social  instincts,  he  reflects  and  compares  the 
now  weakened  impression  of  such  past  impulses  with  tlie  cvcr- 
preseut  social  instincts ;  and  he  then  feels  that  sense  of  dissatis- 
fiiction  which  all  unsatisfied  instincts  leave  behind  them,  he 
therefore  resolves  to  act  differently  for  the  future, — and  this  is 
conscience.  Any  instinct,  permanently  stronger  or  more  endui-ing 
than  another,  gives  rise  to  a  feeling  which  we  express  by  saying 
that  it  ought  to  be  obeyed^  A  pointer  dog,  if  able  to  reflect  on 
his  past  conduct,  would  say  to  himself,  I  ought  (as  indeed  we 
say  of  him)  to  have  pointed  at  that  hare  and  not  have  yielded 
to  the  passing  temptation  of  hunting  it. 

Social  animals  are  impelled  partly  by  a  wish  to  aid  the  members 
of  their  community  in  a  general  manner,  but  more  commonly  to 
perform  certain  definite  actions.  Man  is  impelled  by  the  same 
general  wish  to  aid  his  fellows ;  but  has  few  or  no  special  instincts. 
He  differs  also  from  the  lower  animals  in  the  power  of  expressing 
his  desires  by  words,  which  thus  become  a  guide  to  the  aid  required 
and  bestowed.  The  motive  to  give  aid  is  likewise  much  modified  in 
man  :  it  no  longer  consists  solely  of  a  blind  instinctive  impulse, 
but  is  much  influenced  by  the  praise  or  blame  of  his  fellows. 
The  appreciation  and  the  bestowal  of  praise  and  blame  both 
rest  on  sympathy ;  and  this  emotion,  as  we  have  seen,  is  one  of 
the  most  important  elements  of  the  social  instincts.  Sympathy, 
though  gained  as  an  instinct,  is  also  much  strengthened  by 
exercise  or  habit.  As  all  men  desire  their  own  happiness, 
praise  or  blame  is  bestowed  on  actions  and  motives,  according  as 
they  lead  to  this  end ;  and  as  happiness  is  an  essential  part  of 
the  general  good,  the  greatest-happiness  principle  indirectly 
serves  as  a  nearly  safe  standard  of  right  and  wrong.  As  the 
reasoning  powers  advance  and  experience  is  gained,  the  remoter 
effects  of  certain  lines  of  conduct  on  the  character  of  the  indi- 
vidual, and  on  the  general  good,  are  perceived ;  and  then  the  self- 
regarding  virtues  come  within  the  scope  of  public  opinion,  and 
receive  praise,  and  their  opposites  blame.  But  with  the  less 
civilised  nations  reason  often  errs,  and  many  bad  customs  and 
base  superstitions  come  within  the  same  scope,  and  are  then 
esteemed  as  high  virtues,  and  their  breach »as  heavy  crimes. 

The  moral  faculties  are  generally  and  justly  esteemed  as  of 
higher  value  than  the  intellectual  powers.  But  we  should  bear 
in  mind  that  the  activity  of  the  mind  in  vividly  recalling  past 
impressions  is  one  of  the  fundamental  though  secondary  bases 
of  conscience.  This  affords  the  strongest  argument  for  educating 
and  stimulating  in  all  possible  ways  the  intellectual  faculties  of 


6 1 2  The  Descent  of  Man.  Pa rt  1 II. 

every  liuman  being.  No  doubt  a  man  with  a  torpid  mind,  if  his 
social  affections  and  sympathies  are  well  developed,  will  be  led 
to  good  actions,  and  may  have  a  fairly  sensitive  conscience.  But 
whatever  renders  the  imagination  more  vivid  and  strengthens 
the  habit  of  recalling  and  comparing  past  impressions,  will 
make  the  conscience  more  sensitive,  and  may  even  somewhat 
compensate  for  weak  social  affections  and  sympathies. 

The  moral  nature  of  man  has  reached  its  present  standard, 
partly  through  the  advancement  of  his  reasoning  powers  and 
consequently  of  a  just  public  opinion,  but  especially  from 
his  sympathies  having  been  rendered  more  tender  and  widely 
diffused  through  the  effects  of  habit,  example,  instruction,  and 
reflection.  It  is  not  imiorobable  that  after  long  practice  virtuous 
tendencies  may  be  inherited.  With  the  more  civilised  races,  the 
conviction  of  the  existence  of  an  all-seeing  Deity  has  had  a 
potent  influence  on  the  advance  of  morality.  Ultimately  man 
does  not  accept  the  praise  or  blame  of  his  fellows  as  his  sole 
guide,  though  few  escape  this  influence,  but  his  habitual 
convictions,  controlled  by  reason,  afford  him  the  safest  rule. 
His  conscience  then  becomes  the  supreme  judge  and  monitor. 
Nevertheless  the  first  foundation  or  origin  of  the  moral  sense 
lies  in  the  social  instincts,  including  sympathy;  and  these 
instincts  no  doubt  were  primarily  gained,  as  in  the  case  of  the 
lower  animals,  through  natural  selection. 

The  belief  in  God  has  often  been  advanced  as  not  only  the 
greatest,  but  the  most  complete  of  all  the  distinctions  between 
man  and  the  lower  animals.  It  is  however  impossible,  as  we 
have  seen,  to  maintain  that  this  belief  is  innate  or  instinctive  in 
man.  On  the  other  hand  a  belief  in  all-pervading  spiritual 
agencies  seems  to  be  universal ;  and  apparently  follows  from  a 
considerable  advance  in  man's  reason,  and  from  a  still  greater 
advance  in  his  faculties  of  imagination,  curiosity  and  wonder.  I 
am  aware  that  the  assumed  instinctive  belief  in  God  has  been 
used  by  many  persons  as  an  argument  for  His  existence.  But 
this  is  a  rash  argument,  as  w^e  should  thus  be  compelled  to 
believe  in  the  existence  of  many  cruel  and  malignant  spirits,  only 
a  little  more  powerful  than  man ;  for  the  belief  in  them  is  far 
more  general  than  in  'a  beneficent  Deity.  The  idea  of  a  universal 
and  beneficent  Creator  does  not  seem  to  arise  in  the  mind  of  man, 
until  he  has  been  elevated  by  long-contin^^ed  culture. 

He  who  believes  in  the  advancement  of  man  from  some  low 
organised  form,  will  naturally  ask  how  does  this  bear  on  the  belief 
in  the  immortality  of  the  soul.  The  barbarous  races  of  man,  as 
Sir  J.  Lubbock  has  shewn,  possess  no  clear  belief  of  this  kind  ; 


Chap.  XXI.  General  Siunviary,  613 


but  arguments  derived  from  the  i)rimeval  beliefs  of  savages  aro, 
as  we  have  just  seen,  of  little  or  no  avail.  Few  persons  feel  any 
anxiety  from  the  impossibility  of  determining  at  what  precise 
period  in  the  development  of  the  individual,  from  the  first  trace 
of  a  minute  germinal  vesicle,  man  becomes  an  immortal  being ; 
and  there  is  no  greater  cause  for  anxiety  because  the  period 
cannot  possibly  be  determined  in  the  gradually  ascending  organic 
scale.  - 

I  am  aware  that  the  conclusions  arrived  at  in  this  work 
will  be  denounced  by  some  as  highly  irreligious ;  but  he  who 
denounces  them  is  bound  to  shew  why  it  is  more  irreligious  to 
explain  the  origin  of  man  as  a  distinct  species  by  descent  from 
some  lower  form,  through  the  laws  of  variation  and  natural 
selection,  than  to  explain  the  birth  of  the  individual  through 
the  laws  of  ordinary  reproduction.  The  birth  both  of  the 
species  and  of  the  individual  are  equally  parts  of  that  grand 
sequence  of  events,  which  our  minds  refuse  to  accept  as  the 
result  of  blind  chance.  The  understanding  revolts  at  such  a 
conclusion,  whether  or  not  we  are  able  to  believe  that  every  slight 
variation  of  structure, — the  union  of  each  pair  in  marriage, — 
the  dissemination  of  each  seed, — and  other  such  events,  have  all 
been  ordained  for  some  special  purpose. 

Sexual  selection  has  been  treated  at  great  length  in  this 
work;  for,  as  I  have  attempted  to  shew,  it  has  played  an 
important  part  in  the  history  of  the  organic  world.  I  am 
aware  that  much  remains  doubtful,  but  I  have  endeavoured 
to  give  a  fair  view  of  the  whole  case.  In  the  lower  divisions 
of  the  animal  kingdom,  sexual  selection  seems  to  have  done 
nothing :  such  animals  are  often  affixed  for  life  to  the  same 
spot,  or  have  the  sexes  combined  in  the  same  individual,  or 
what  is  still  more  important,  their  perceptive  and  intellectual 
faculties  are  not  suflQciently  advanced  to  allow  of  the  feelings  of 
love  and  jealousy,  or  of  the  exertion  of  choice.  AYhen,  however, 
we  come  to  the  Arthropoda  and  Yertebrata,  even  to  the  lowest 
classes  in  these  two  great  Sub-Kingdoms,  sexual  selection  has 
efiected  much. 

In  the  several  great  classes  of  the  animal  kicgdom, — in 
mammals,  birds,  reptiles,  fishes,  insects,  and  even  crustaceans, 
— the  differences  between  the  sexes  follow  nearly  the  same 
rules.  The  males  are  almost  always  the  wooers;  and  they 
alone  are  armed  with  special  weapons  for  fighting  with  their 
rivals.     They  are  generally  stronger  and  larger  than  the  females, 

'  The  Rev.  J.  A.  Picton  gives  a  discussion  to  this  effect  in  his  *Ne\v 
Theories  and  the  Old  Faith,'  1870. 


6 14  ■    TJie  Descent  of  Man.  Part  III. 

and  are  endowed  with  the  requisite  qualities  of  courage  and 
pugnacity.  They  are  provided,  either  exclusively  or  in  a  much 
liigher  degree  than  the  females,  with  organs  for  vocal  or  instru- 
mental music,  and  with  odoriferous  glands.  They  are  ornamented 
with  infinitely  diversified  appendages,  and  with  the  most 
brilliant  or  cousidIcuous  colours,  often  arranged  in  elegant 
patterns,  whilst  the  females  are  unadorned.  When  the  sexes 
differ  in  more  important  structures,  it  is  the  male  which  is 
provided  with  special  sense-organs  for  discovering  the  female, 
with  locomotive  organs  for  reaching  her,  and  often  with 
prehensile  organs  for  holding  her.  These  various  structures  for 
charming  or  securing  the  female  are  often  develoi^ed  in  the  male 
during  only  part  of  the  year,  namely  the  breeding-season.  They 
have  in  many  cases  been  more  or  less  transferred  to  the  females ; 
and  in  the  latter  case  they  often  appear  in  her  as  mere  rudiments. 
They  are  lost  or  never  gained  by  the  males  after  emasculation. 
Generally  they  are  not  developed  in  the  male  during  early 
youth,  but  appear  a  short  time  before  the  age  for  reproduction. 
Hence  in  most  cases  the  young  of  both  sexes  resemble  each  other ; 
and  the  female  somewhat  resembles  her  young  offspring  through- 
out life.  In  almost  every  great  class  a  few  anomalous  cases 
occur,  where  there  has  been  an  almost  complete  transposition  of 
the  characters  proper  to  the  two  sexes ;  the  females  assuming 
characters  which  properly  belong  to  the  males.  This  surprisiDg 
uniformity  in  the  laws  regulating  the  differences  between  the 
sexes  in  so  many  and  such  widely  separated  classes,  is  intelligible 
if  we  admit  the  action  of  one  common  cause,  namely  sexual 
selection. 

Sexual  selection  depends  on  the  success  of  certain  individuals 
over  others  of  the  same  sex,  in  relation  to  the  propagation  of  the 
species ;  whilst  natural  selection  depends  on  the  success  of  both 
sexes,  at  all  ages,  in  relation  to  the  general  conditions  of  life. 
The  sexual  struggle  is  of  two  kinds ;  in  the  one  it  is  between  the 
individuals  of  the  same  sex,  generally  the  males,  in  order  to  drive 
away  or  kill  their  rivals,  the  females  remaining  passive ;  whilst  in 
the  other,  the  struggle  is  likewise  between  the  individuals  of  the 
same  sex,  in  order  to  excite  or  charm  those  of  the  opposite 
sex,  generally  the  females,  which  no  longer  remain  passive,  but 
select  the  more  agi-eeable  piartners.  This  latter  kind  of  selection 
is  closely  analogous  to  that  which  man  unintentionally,  yet 
effectually,  brings  to  bear  on  his  domesticated  productions, 
when  he  preserves  during  a  long  period  the  most  pleasing  or 
useful  individuals,  without  any  wish  to  modify  the  breed. 

The  laws  of  inheritance  determine  whether  characters  gained 
through  sexual  selection  by  either  sex  shall  be  transmitted  to  the 


Chap.  XXI.  General  Siumnary,  615 

same  sex,  or  to  both ;  as  well  as  tlic  age  at  which  they  shall  be 
developed.  It  appears  that  variatious  arising  late  m  life  are 
commonly  transmitted  to  one  and  the  same  sex.  Variability  is 
the  necessary  basis  for  the  action  of  selection,  and  is  wholly 
independent  of  it.  It  follows  from  this,  that  variations  of  the 
same  general  nature  have  often  been  taken  advantage  of  and 
accumulated  through  sexual  selection  in  relation  to  the  propaga- 
tion of  the  species,  as  well  as  through  natural  selection  in 
relation  to  the  general  purposes  of  life.  Hence  secondary 
sexual  characters,  when  equally  transmitted  to  both  sexes  can  be 
distinguished  from  ordinary  specific  characters  only  by  the  light 
of  analogy.  The  modifications  acquired  through  sexual  selection 
are  often  so  strongly  pronounced  that  the  two  sexes  have 
frequently  been  ranked  as  distinct  species,  or  even  as  distinct 
genera.  Such  strongly-marked  differences  must  be  m  some 
manner  highly  important;  and  we  know  that  they  have  been 
acquired  in  some  instances  at  the  cost  not  only  of  inconvenience, 
but  of  exposure  to  actual  danger. 

The  belief  in  the  power  of  sexual  selection  rests  chiefly  on 
the  following  considerations.  Certain  characters  are  confined 
to  one  sex;  and  this  alone  renders  it  probable  that  in  most 
cases  they  are  connected  with  the  act  of  reproduction.  In 
innumerable  instances  these  characters  are  fully  developed  only 
at  maturity,  and  often  during  only  a  part  of  the  year,  which 
is  always  the  breeding-season.  The  males  (passing  over  a  few 
exceptional  cases)  are  the  more  active  in  courtship ;  they  are  the 
better  armed,  and  are  rendered  the  more  attractive  m  various  ways. 
It  is  to  be  especially  observed  that  the  males  display  their 
attractions  with  elaborate  care  in  the  presence  of  the  females; 
and  that  they  rarely  or  never  display  them  excepting  during 
the  season  of  love.  It  is  incredible  that  all  this  should  be 
purposeless.  Lastly  we  have  distinct  evidence  with  some  quad- 
rupeds and  birds,  that  the  individuals  of  one  sex  are  capable  of 
feeling  a  strong  antipathy  or  preference  for  certain  individuals 
of  the  other  sex. 

Bearing  in  mind  these  facts,  and  the  marked  results  of  mans 
unconscious  selection,  when  applied  to  domesticated  animals  and 
cultivated  plants,  it  seems  to  me  almost  certain  that  if  the 
individuals  of  one  sex  were  during  a  long  series  of  generations  to 
prefer  pairing  with  certain  individuals  of  the  other  sex,  charac- 
terised in  some  peculiar  manner,  the  offspring  would  slowly  but 
surely  become  modified  in  this  same  manner.  I  have  not 
attempted  to  conceal  that,  excepting  when  the  males  are  more 
numerous  than  the  females,  or  when  polygamy  prevails,  it  is 
doubtful  how  the  more  attractive  males  succeed  in  leaving  a 


6i6  The  Descent  of  Man.  Part  IIT. 

larger  number  of  offspring  to  inherit  their  superiority  in  orna- 
ments or  other  charms  than  the  less  attractive  males ;  but  I 
have  shewn  that  this  would  probably  follow  from  the  females, — 
especially  the  more  vigorous  ones,  which  would  be  the  first  to 
breed, — preferring  not  only  the  more  attractive  but  at  the  same 
time  the  more  vigorous  and  victorious  males. 

Although  we  have  some  positive  evidence  that  birds  appre- 
ciate bright  and  beautiful  objects,  as  with  the  bower-birds  of 
Australia,  and  although  they  certainly  appreciate  the  power  of 
song,  yet  I  fully  admit  that  it  is  astonishing  that  the  females 
of  many  birds  and  some  mammals  should  be  endowed  with 
sufficient  taste  to  appreciate  ornaments,  which  we  have  reason  to 
attribute  to  sexual  selection ;  and  this  is  even  more  astonishing 
in  the  case  of  reptiles,  iish,  and  insects.  But  we  really  know 
little  about  the  minds  of  the  lower  animals.  It  cannot  be 
supposed,  for  instance,  that  male  birds  of  paradise  or  pea<30cks 
should  take  such  pains  in  erecting,  spreading,  and  vibrating 
their  beautiful  plumes  before  the  females  for  no  purpose.  We 
should  remember  the  fact  given  on  excellent  authority  in  a 
former  chapter,  that  several  i3eahens,  when  debarred  from  an 
admired  male,  remained  widows  during  a  whole  season  rather 
than  pair  with  another  bird. 

Nevertheless  I  know  of  no  fact  in  natuj'al  history  more  wonder- 
ful than  that  the  female  Argus  pheasant  should  ai^preciate  the 
exquisite  shading  of  the  ball-and-socket  ornaments  and  the 
elegant  patterns  on  the  wing-feathers  of  the  male.  He  who 
thinks  that  the  male  was  created  as  he  now  exists  must  admit 
that  the  great  j^lumes,  which  prevent  the  wings  from  being  used 
for  flight,  and  which  are  displayed  during  courtship  and  at  no 
other  time  in  a  manner  quite  peculiar  to  this  one  species,  were 
given  to  him  as  an  ornament.  If  so,  he  must  likewise  admit  that 
the  female  was  created  and  endowed  with  the  capacity  of  appre- 
ciating such  ornaments.  I  differ  only  in  the  conviction  that  the 
male  Argus  pheasant  acquired  his  beauty  gradually,  through  the 
preference  of  the  females  during  many  generations  for  the  more 
highly  ornamented  males  ;  the  aesthetic  capacity  of  the  females 
having  been  advanced  through  exercise  or  habit,  just  as  our  own 
taste  is  gradually  improved.  In  the  male  through  the  fortunate 
chance  of  a  few  feathers  being  left  unchanged,  we  can  distinctly 
trace  how  simple  spots  with  a  little  fulvous  shading  on  one  side 
may  have  been  developed  by  small  steps  into  the  wonderful 
ball-and-socket  ornaments;  and  it  is  probable  that  they  were 
actually  thus  developed. 

Everyone  who  admits  the  principle  of  evolution,  and  yet 
feels  great  difficulty  in  admitting  that  female  mammals,  birds, 


Chap.  XXI.  General  Summary,  617 

reptiles,  and  fish,  could  have  acquired  the  high  taste  implied  by 
the  beauty  of  the  males,  and  which  generally  coincides  with  our 
own  standard,  should  reflect  that  the  nerve- cells  of  the  brain  in 
the  highest  as  well  as  in  the  lowest  members  of  the  Vertebrate 
series,  are  derived  from  those  of  the  common  progenitor  of  this 
great  Kingdom.  For  we  can  thus  see  how"  it  has  come  to  pass 
that  certain  mental  faculties,  in  various  and  widely  distinct 
groups  of  animals,  have  been  developed  in  nearly  the  same 
manner  and  to  nearly  the  same  degree. 

The  reader  who  has  taken  the  trouble  to  go  through  the 
several  chapters  devoted  to  sexual  selection,  will  be  able  to 
judge  how  far  the  conclusions  at  which  I  have  arrived  are 
supported  by  sufficient  evidence.  If  he  accepts  these  conclusions 
he  may,  I  think,  safely  extend  them  to  mankind ;  but  it  would 
be  superfluous  here  to  repeat  what  I  have  so  lately  said  on  the 
manner  in  which  sexual  selection  apparently  has  acted  on  man, 
both  on  the  male  and  female  side,  causing  the  two  sexes  to  differ 
in  body  and  mind,  and  the  several  races  to  differ  from  each 
other  in  various  characters,  as  well  as  from  their  ancient  and 
lowly-organised  progenitors. 

He  w^ho  admits  the  principle  of  sexual  selection  wall  be  led 
to  the  remarkable  conclusion  that  the  nervous  system  not  only 
regulates  most  of  the  existing  functions  of  the  body,  but  has 
indirectly  influenced  the  progressive  development  of  various 
bodily  structures  and  of  certain  mental  qualities.  Courage, 
pugnacity,  perseverance,  strength  and  size  of  body,  weapons  of 
all  kinds,  musical  organs,  both  vocal  and  instrumental,  bright 
colours  and  ornamental  appendages,  have  all  been  indirectly 
gained  by  the  one  sex  or  the  other,  through  the  exertion  of 
choice,  the  influence  of  love  and  jealousy,  and  the  ai^preciation 
of  the  beautiful  in  sound,  colour  or  form  ;  and  these  powers  of 
the  mind  manifestly  depend  on  the  development  of  the  brain. 

Man  scans  with  scrupulous  care  the  character  and  pedigree 
of  his  horses,  cattle,  and  dogs  before  he  matches  them ;  but 
when  he  comes  to  his  own  marriage  he  rarely,  or  never,  takes 
any  such  care.  He  is  impelled  by  nearly  the  same  motives  as 
the  lower  animals,  when  they  are  left  to  their  own  free  choice, 
though  he  is  in  so  far  superior  to  them  that  he  highly  values 
mental  charms  and  virtues.  On  the  other  hand  he  is  strongly 
attracted  by  mere  wealth  or  rank.  Yet  he  might  by  selection  do 
something  not  only  for  the  bodily  constitution  and  frame  of  his 
offspring,  but  for  their  intellectual  and  moral  qualities.  Both 
sexes  ought  to  refrain  from  marriage  if  they  are  in  any  marked 
degree  inferior  in  body  or  mind ;  but  such  hopes  are  Utopian  and 


6 1 8  The  Descent  of  Man.  Part  III . 

will  never  be  even  partially  realised  until  the  laws  of  inheritance 
are  thoroughly  known.  Everyone  does  good  service,  who  aids 
towards  this  end.  "When  the  principles  of  breeding  and  in- 
-heritance  are  better  understood,  we  shall  not  hear  ignorant 
members  of  our  legislature  rejecting  with  scorn  a  plan  for 
ascertaining  whether  or  not  consanguineons  marriages  are 
injurious  to  man. 

The  advancement  of  the  welfare  of  mankind  is  a  most  intricate 
problem :  all  ought  to  refrain  from  marriage  who  cannot  avoid 
abject  poverty  for  their  children ;  for  poverty  is  not  only  a  great 
svil,  but  tends  to  its  own  increase  by  leading  to  recklessness  in 
marriage.  On  the  other  hand,  as  Mr.  Galton  has  remarked,  if 
the  prudent  avoid  marriage,  whilst  the  reckless  marry,  the 
inferior  members  tend  to  supplant  the  better  members  of 
society.  Man,  like  every  other  animal,  has  no  doubt  advanced 
to  his  present  high  condition  through  a  struggle  for  existence 
consequent  on  his  rapid  multiplication ;  and  if  he  is  to  advance 
still  higher,  it  is  to  be  feared  that  he  must  remain  subject  to  a 
severe  struggle.  Otherwise  he  would  sink  into  indolence,  and 
the  more  gifted  men  would  not  be  more  successful  in  the  battle 
of  life  than  the  less  gifted.  Hence  our  natural  rate  of  increase, 
though  leading  to  many  and  obvious  evils,  must  not  be  greatly 
diminished  by  any  means.  There  s];iould  be  open  competition 
for  all  men ;  and  the  most  able  should  not  be  prevented  by  laws 
or  customs  from  succeeding  best  and  rearing  the  largest  number 
of  offspring.  Important  as  the  struggle  for  existence  has  been 
and  even  still  is,  yet  as  far  as  the  highest  part  of  man's  nature 
is  concerned  there  are  other  agencies  more  important.  For  the 
moral  quantities  are  advanced,  either  directly  or  indirectly, 
much  more  through  the  effects  of  habit,  the  reasoning  powers, 
instruction,  religion,  &c.,  than  through  natural  selection; 
though  to  this  latter  agency  may  be  safely  attributed  the 
social  instincts,  which  afforded  the  basis  for  the  development  of 
the  moral  sense. 

The  main  conclusion  arrived  at  in  this  work,  namely  that 
man  is  descended  from  some  lowly  organised  form,  will,  I  regi'et 
to  think,  be  highly  distasteful  to  many.  But  there  can  hardly 
be  a  doubt  that  we  are  descended  from  barbarians.  The  aston- 
ishment which  I  felt  on  first  seeing  a  pai'ty  of  Feugians  on  a 
wild  and  broken  shore  will  never  be  forgotten  by  me,  for  the 
reflection  at  once  rushed  into  my  mind — such  were  our  ancestors. 
These  men  were  absolutely  naked  and  bedaubed  with  paint, 
theu-  loDg  hair  was  tangled,  their  mouths  frothed  with  excite- 
ment, and  their  expression  was  wild,  startled,  and  distrustful. 


CiiAi'.  XXI.  General  Suuwiary.  619 

They  possessed  hardly  any  arts,  and  like  wild  animals  lived  on 
what  they  could  catch  ;  they  had  no  government,  and  were 
merciless  to  every  one  not  of  their  own  small  tribe.  lie  who  has 
seen  a  savage  in  his  native  land  will  not  feel  much  shame,  if 
force(f  to  acknowledge  that  the  blood  of  some  more  humble 
creature  flows  in  his  veins.  For  my  own  part  I  would  as  soon 
be  descended  from  that  heroic  little  monkey,  who  braved  his 
dreaded  enemy  in  order  to  save  the  life  of  his  keepci',  or  from 
that  old  baboon,  who  descending  from  the  mountains,  carried 
away  in  triumph  his  young  comrade  from  a  crowd  of  astonished 
dogs — as  from  a  savage  who  delights  to  torture  his  enemies, 
offers  up  bloody  sacrifices,  practises  infanticide  without  r«morsc, 
treats  his  wives  like  slaves,  knows  no  decency,  and  is  haunted 
by  the  grossest  superstitions. 

Man  may  be  excused  for  feeling  some  j^ride  at  having  risen, 
though  not  through  his  own  exertions,  to  the  very  summit  of  the 
organic  scale  ;  and  the  fact  of  his  having  thus  risen,  instead  of 
having  been  aboriginally  placed  there,  may  give  him  hope  for  a 
still  higher  destiny  in  the  distant  future.  But  we  are  not  here 
concerned  with  hopes  or  fears,  only  with  the  truth  as  far  as  our 
reason  permits  us  to  discover  it ;  and  I  have  given  the  evidence 
to  the  bestof  my  ability.  We  must,  however,  acknowledge,  as  it 
seems  to  me,  that  man  with  all  his  noble  qualities,  with  sympathy 
which  feels  for  the  most  debased,  with  benevolence  which  extends 
not  only  to  other  men  but  to  the  humblest  living  creature,  with 
his  god-like  intellect  which  has  penetrated  into  the  movemcats 
and  constitution  of  the  solar  system — with  all  these  exalted 
powers — Man  still  bears  in  his  bodily  frame  the  indehble  stamp 
of  his  lowly  orjgiii. 


INDEX. 


AGRION. 


Abbot,  C,  on  the  battles-  of  seals, 
500. 

Abductor  of  the  fifth  metatarsal, 
presence  of,  in  man,  42. 

Abercrombie,  Dr.,  on  disease  of  the 
brain  affecting  speech,  88. 

Abipones,  marriage  customs  of  the, 
598. 

Abou-Simbel,  caves  of,  168. 

Abortion,  prevalence  of  the  practice 
of,  46. 

Abstraction,  power  of,  in  animals,  83. 

Acalles,  stridulation  of,  306. 

Acanthodactylus  capensis,  sexual  dif- 
ferences of  colour  in,  357. 

Accentor  modularis,  473. 

Acclimatisation,  difference  of,  in  dif- 
ferent races  of  men,  167. 

Achetidae,  stridulation  of  the,  282, 
283,  285  ;  rudimentary  stridulating 
organs  in  female,  288. 

Acilius  sulcatus,  elytra  of  the  female, 
276. 

Acomiis,  development  of  spurs  in  the 
female  ol',  450. 

Acridiidse  stridulation  of  the,  282, 
286  ;  rudimentary  stridulating  or- 
gans in  female,  288. 

Acromio-basilar  muscle,  and  quad- 
rupedal gait,  42. 

Acting,  178. 

Actinice.  bright  colours  of,  260. 

Adams,  Mr.,  migration  of  birds,  108  ; 
intelligence  of  nut-hatch,  418;  on 
the  Bomb'/cilia  carolinensis,  461. 

Admiral  butterfly,  312. 

Adoption  of  the  young  of  other  ani- 
mals by  female  monkeys,  70. 

Advancement  in  the  organic  scale. 
Von  Baer's  definition  cf,  164. 

Acby,  on    the   ditibreuce  between    the 


skulls  of  man  and  the  quadrumana, 
149. 

^Esthetic  faculty,  not  highly  deve- 
loped in  savages,  93. 

Affection,  maternal,  70  ;  manifestation 
of,  by  animals,  70 ;  parental  and 
filial,  partly  the  result  of  natural 
selection,  105 ;  mutual,  of  birds, 
410 ;  shewn  by  birds  in  confine- 
ment, for  certain  persons,  411. 

Africa,  probably  the  birthplace  of 
man,  155 ;  South,  crossed  popula- 
tion of,  174 ;  South,  retention  of 
colour  by  the  Dutch  in,  193  ;  South, 
proportion  of  the  sexes  in  the  but- 
terflies of,  250  ;  tattooing  practised 
in,  574 ;  Northern,  coiffure  of 
natives  of,  575. 

Agassiz,  L.,  on  conscience  in  dogs, 
103;  on  the  coincidence  of  the 
races  of  man  with  zoological  pro- 
vinces, 169  ;  on  the  number  of 
species  of  man,  174;  on  the  court- 
ship of  the  land-snails,  262  ;  on  the 
brightness  of  the  colours  of  male 
fishes  during  the  breeding  season, 
340  ;  on  the  frontal  protuberance 
of  the  males  of  Gcophagxis  and 
Cichia,  340,  345 ;  male  fishes 
hatching  ova  in  their  mouths,  345  ; 
sexual  ditlerences  in  colour  of  chro- 
mids,  345 ;  on  the  slight  sexual 
differences  of  the  South  Americans, 
561 ;  on  the  tattooing  of  the  Ama- 
zonian Indians,  576. 

Age,  in  relation  to  the  transmission 
of  characters  in  birds,  463 ;  va- 
riation in  accordance  with,  in  birds, 
484. 

Agelceiis  pliccniceus,  225,  416. 

Ageronia  feronia,  noise  produced  by, 
307. 

Agrion,  dimorphism  in.  290. 


INDEX. 


621 


Agrion  Eamlmrii,  sexes  of,  290. 

AgrionidcE,  ditTerence  in  the  sexes  of, 
290. 

Agrotis  exclainationis,  316. 

Ague,  tertian,  dog  suffering  from,  8. 

A'ithurus  polytmus,  young  of,  487. 

Ainos,  hairiness  of  the,  5tJ0. 

Albino  birds,  419. 

Alca  torda,  young  of,  48G. 

Alces  palmata,  515. 

Alder  and  Hancock,  M^f.,  oa  the 
nudi-branch  moUusca,  204. 

Allen,  J.  A,,  vigour  of  birds  earliest 
hatched,  212,  213;  effect  of  dif- 
ference of  temperature,  light,  &;c., 
on  birds,  225 ;  colours  of  birds, 
422 ;  on  the  relative  size  of  the 
sexes  of  Callorhinus  ursiniis,  515; 
on  the  mane  of  Otaria  jubata,  521  ; 
on  the  pairing  of  seals,  523 ;  on 
sexual  differences  in  the  colour  of 
bats,  534. 

S.,   on    the    habits   of  Hoplo- 

pterus,  366  ;  on  the  plumes  of 
herons,  391  ;  on  the  vernal  moult 
of  Herodias  bubulcus,  393. 

Alligator,  courtship  of  the  male,  221, 
351 ;  roaring  of  the  male,  567. 

Amadavat,  pugnacity  of  male,  366. 

Amadina  Lathami,  display  of  plumage 
by  the  male,  402. 

castanotls,   display   of  plumage 

by  the  male,  402. 

Amazons,  butterflies  of  the.  250  ; 
fishes  of  the,  343. 

America,  variation  in  the  skulls  of 
aborigines  of,  26  ;  wide  range  of 
aborigines  of,  169 ;  lice  of  the 
natives  of,  170;  general  beardless- 
ness  of  the  natives  of,  560. 

,    North,    butterflies    of,     250; 

Indians  of,  women  a  cause  of  strife 
among  the,  561  ;  Indians  of,  their 
notions  of  female  beauty,  577, 
580. 

,  South,  character  of  the  natives 

of,  168  ;  population  of  parts  of, 
173  ;  piles  of  stones  in,  179  ;  ex- 
tinction of  the  fossil  horse  of,  191 ; 
desert-birds  of,  490  ;  slight  sexual 
difference  of  the  aborigines  of,  561  ; 
prevalence  of  infanticiile  in,  592. 

American  languages,  often  highly 
artificial,  91. 

Americans,    wide  geographical   range 


of,  29;  native,  variability  of,  174; 

and    negroes,    ditTerence    of,    197  ; 

aversion  of,  to  hair  on  the  face,  580. 
Amrnophila,  on  the  jaws  of,  275. 
Ammoiragus  tragelaphus^  hairy   foie- 

legs  of,  531,  533. 
Amphibia,   affinity  of,   to    the   ganoid 

fishes,    159 ;    vocal   organs  of  the, 

566. 
Amphibians,     165,     348  ;      breeding 

whilst  immature,  485 
Amphioxus^  159. 
Amphipoda,    males    sexually    mature 

wliile  young,  485. 
Amunoph    III.,    negro    character    of 

features  of,  168. 
Anal  appendages  of  insects,  275. 
Analogous  variation  in  the  plumage  of 

birds,  385. 
Anas,  462. 

• acuta,  male  plumage  of,  393. 

boschas,  male  plumage  of,  393. 

histrionica,  484. 

punctata,  374, 

Anastomus  oscitans,  sexes   and  young 

of,  486  ;  white  nuptial  plumage  of, 

492. 
Anatida?,  voices  of,  374. 
Anax  Junius,  differences   in  the  sexes 

of,  290. 
Andaman    islanders,    susceptible     to 

change  of  climate,  188. 
Anderson,  Dr.,  on  the  tail  of  Macacus 

brunncus,  59 ;   the   Bufo  sikimtnen- 

sis,  349;  sounds  of  Lchi's  cirinata, 

353. 
Andrwna  fnha,  292. 
Anglo-Saxons,  estimation  of  the  beard 

among  the,  581. 
Animals,  domesticated,  more    fertile 

than  wild,  45 ;    cruelty  of  savages 

to,    118;    characters    common    to 

man  and,  146 ;  domestic,  change  of 

l)reeds  of,  596. 
Annelida,  264;  colours  of,  265. 
Anobium  tessellatum,  sounds  produced 

by,  306. 
Anolis  cristatellus,  ntale,  crest  of,  354  ; 

pugnacity  of  the  male,  354  ;  throat- 
pouch  of,  354. 
Anser  canadensis,  416. 
C'/gnoides,  415  ;  knob  at  the  base 

of  the  beak  of,  426. 

hyperboreus,  whiteness  of,  492. 

Antelope,  prong-horned,  horns  of,  234. 


622 


ANTELOPES. 


INDEX. 


Antelopes,  gonerally  polygamous.  217  ; 
horns  of,  234,  504 ;  canine  teeth  of 
some  male,  502  ;  use  of  horns  of 
509  ;  dorsal  crests  in,  530 ;  dew- 
laps of,  531 ;  winter  change  of  two 
species  of,  543  ;  peculiar  markings 
of,  543. 

Antennje,  furnished  with  cushions  in 
the  male  of  Penthe,  276. 

Anthidium  manicatum,  large  male  of, 
279. 

Anthocharis  cardamines,  308,  312; 
sexual  difference  of  colour  in,  322. 

genutia,  312. 

sara,  312. 

Anthophora  acervorum,  large  male  of, 
279. 

retusa,  diff'erence  of  the  sexes  in, 

292. 

Anthropida?,  152. 

Anthus,  moulting  of,  392. 

Antics  of  birds,  380. 

Antigua,  Dr.  Nicholson's  observations 
on  yellow  fever  in,  195. 

Antilocapra  amoHcana,  horns  of,   234, 

504,  507. 

Antilope  hezom^tica,  horned  females  of, 

505,  506,  507  ;  sexual  difference  in 
the  colour  of,  535. 

Dorcas  and  euclxorc,  504. 

euchore,  horns  of,  509. 

moivtana,  rudimentary  canines  in 

the  young  male  of,  514. 

nigcr,  sing-sing,  caama,  and  gor- 

gon,  sexual  differences  in  the  colours 
of,  536. 

oreas,  horns  of,  234. 

sa«(/a,  polygamous  habits  of,  217. 

strepsiceros^  horns  of,  234. 

• subgutturosa,  absence  of  sub- 
orbital pits  in,  529. 

Antipathy,  shewn  by  birds  in  confine- 
ment, to  certain  persons,  411. 

Ants,  147;  large  size  of  the  cerebral 
ganglia  in,  54 ;  soldier-,  large  jaws 
of,  63  ;  playing  together,  69  ; 
memory  in,  74  ;  intcommunication 
of,  by  means  of  the  antennae,  89  ; 
habits  of,  147 :  difference  of  the 
sexes  in,  292  ;  recognition  of  each 
other  by,  after  separation,  292. 

White,  habits  of,  291. 

Auura,  349. 

Apat  inia  muliebris,  male  unknown, 
254. 


Apathus.  difference  of  the  sexes  in, 
292. 

Apatura  Iris,  307,  308. 

Apes,  difference  of  the  young,  from 
the  a  iu,t,  8  ;  semi-erect  attitude  of 
some,  5'-'  ;  mastoid  processes  of,  53  ; 
influences  of  the  jaw-muscles  on 
the  physiognomy  of,  54 ;  female, 
destitute  of  large  canines,  63 ; 
building  platforms,  82 ;  imitative 
faculties  of,  129 ;  anthropomor- 
phous, 153  ;  probable  speedy  ex- 
termination of  the,  156  ;  Gratiolet 
on  the  evolution  of,  177  ;  canine 
teeth  of  male,  502  ;  females  of 
some,  less  hairy  beneath  than  the 
males,  601. 

long-armed,  their  mode  of  pro- 
gression, 52. 

Aphasia,  Dr.  Batemau  on,  88. 

Apis  mdlifica,  large  male  of,  279. 

Apollo,  Greek  statues  of,  581. 

Apoplexy  in  Cebus  Azarse,  7. 

Appendages,  anal,  of  insects,  276. 

Approbation,  influence  of  the  love  of, 
109,  116,  131. 

Aprosmictus  scapuhtus,  4:bl . 

Apus,  proportion  of  sexes,  255. 

Aquatic  birds,  frequency  of  white 
plumage  in,  495. 

Aquila  chrysaetos,  408. 

Arab  women,  elaborate  and  peculiar 
coiffure  of,  583. 

Arabs,  fertility  of  crosses  with  other 
races,  171;  gashing  of  cheeks  and 
temples  among  the,  574. 

Arachnida,  272. 

Arakhan,  artificial  widening  of  the 
forenead  by  the  natives  of,  583. 

ArboiHcola,  young  of,  468. 

Archeopteryx,  158. 

Arctiidfe,  coloration  of  the,  314. 

Ardea  asha,  rufescens,  and  cxruler, 
change  of  colour  in,  494,  495. 

cset^ulea,  breeding   in  immature 

plumage,  484. 

gularis,  change   of  plumage  in, 

495. 

herodias,    love-gestures    of   the 

male,  380. 

ludoticiana,  age  of  mature  plu- 
mage in,  483 ;  continued  growth 
of  crest  and  plumes  in  the  male  of, 
485. 

nycticorax,  cries  of,  368. 


INDEX. 


AUSTRALIA. 


623 


Ardeola,  young  of,  468. 

Ardctta,  changes  of  plumage  in,  461. 

Argeateuil,  22. 

Argus  pheasant,  384-,  403,  462  ;  dis- 
play of  plumage  by  the  male,  398  ; 
ocellated  spots  of  the,  428 ;  grada- 
tion of  characters  in  the,  434. 

Argyll,  Duke  of,  on  the  physical 
weakness  of  man,  63  ;  the  fashion- 
ing of  implements  peculiar  to  man, 
82  ;  on  the  contest  in  man  between 
right  and  wrong,  12o  ;  on  the  pri- 
mitive civilisation  of  man,  143  ;  on 
the  plumage  of  the  male  Argus 
pheasant,  398 ;  on  Urosticte  Ben- 
yamm/,  442,  443 ;  on  the  nests  of 
birds,  453. 

Argijnnis,  colouring  of  the  lower  sur- 
face of,  314. 

Aricoris  epitus,  sexual  differences  in 
the  wings  of,  277. 

Aristocracy,  increased  beauty  of  the, 
586. 

Arms,  proportions  of,  in  soldiers  and 
sailors,  32  ;  direction  of  the  hair  on 
the,  151. 

and  hands,  free  use  of,  indi- 
rectly correlated  with  diminution 
of  canines,  53. 

Arrest  of  development,  35,  36. 

Arrow-heads,  stone,  general  resem- 
blance of,  179. 

Arrows,  use  of,  179. 

Arteries,  variations  in  the  course  of 
the,  26. 

Artery,  effect  of  tying,  upon  the 
lateral  channels,  32. 

Arthropoda,  265. 

Arts  practised  by  savages,  179. 

Ascension,  coloured  incrustation  on 
the  rocks  of,  263. 

Ascidia,  affinity  of  the  lancelet  to, 
159;  tadpole-like  hvrva;  of,  159. 

Ascidians,  262 ;  bright  colours  of 
some,  260. 

Asinus,  Asiatic  and  African  species  of, 
548. 

tcsniopus,  548. 

Ass,  colour-variations  of  the,  547. 

Ateles,  effects  of  brandy  on  an,  7 ; 
absence  of  the  thumb  in,  51. 

beehebuth,  ears  of,  15. 

marginatus,   colour   of  the  ruff 

of,  537 ;  hair  on  the  head  of,  549. 

Atcuchus,  striduhition  of,  306, 


Atcuch'is,  ciciitricosus,  habits  of,  300. 

Athalia,  proportions  of  the  sexes  in, 
254. 

Atropus  pidsatorivs,  291. 

Attention,  manifestj.tions  of,  in  ani- 
mals, 73. 

Audouin,  V.,  on  a  hymenopterous  pa- 
rasite with  a  sedentary  male,  221. 

Audubon,  J.  J.,  on  the  pinioned  goose, 
105;  on  the  speculum  oi  Mergus 
cucullatus,  236  ;  on  the  pugnacity 
of  male  birds,  362,  366  ;  on  Tetrao 
cupido,  367  ;  on  Ardea  n;/ct'Corar, 
368  ;  on  Sturnella  ludoviciana,  368  ; 
on  the  vocal  organs  of  Tetrao 
cnpido,  371;  on  the  drumming  of 
the  male  Tetrao  umbellus,  375 ;  on 
sounds  produced  by  the  nightjar, 
376 ;  on  Ardea  herodias  and  Cat.'t- 
artes  jota,  380 ;  on  Jlirmis  pol;/- 
glottics,  393 ;  on  display  in  male 
birds,  394  ;  on  the  spring  change  of 
colour  in  some  finches,  394  ;  recog- 
nition of  a  dog  by  a  turkey,  412 ; 
selection  of  mate  by  femile  birds, 
416;  on  the  turkey,  411,  412,  419; 
on  variation  in  the  male  scarlet 
tanager,  424 ;  on  the  musk-rat, 
5-f2 ;  on  the  habits  of  Pijranga 
wstiva,  453  ;  on  local  differences  in 
the  nests  of  the  same  species  of 
birdi,  456  ;  on  the  habits  of  wood- 
peckers, 458  ;  on  Bombj/cida  caro- 
linensis,  461 ;  on  young  females  of 
Tijranga  wstlva  acquiring  male  cha- 
racters, 462 ;  on  the  immature 
])lumage  of  thrushes,  464;  on  the 
immature  plumage  of  birds,  465  et 
se I. ;  on  birds  breeding  in  immature 
plumage,  484;  on  the  growth  of 
the  crest  and  plumes  in  the  male 
A  7-dea  ludoviciana,  485 ;  on  the 
change  of  colour  in  some  species  of 
Ardea,  494 

Audubon  and  Bachman,  M.iVf.,  on 
squirrels  fighting,  500 ;  on  tlie 
Canadian  lynx,  521. 

Aughey,  Prof.,  on  rattlesnakes,  353. 

Austen,  N.L.,  on  Anolis  cristatellus, 
354. 

Australia,  not  the  birthplace  of  man, 
155  ;  half-castes  killed  by  the 
natives  of,  170;  lice  of  the  natives 
of,  170;  prevalence  of  female  in- 
lauticide  in,  592. 


624 


AUSTRALIA. 


INDEX. 


BARRINGTON. 


Australia,  South,  variation  in  the 
skulls  of  aborigines  of,  26. 

Australians,  colour  of  newborn  chil- 
dren of,  557  ;  relative  height  of  the 
sexes  of,  559  ;  women  a  cause  of 
war  among  the,  561. 

Axis  deer,  sexual  difference  in  the 
colour  of  the,  537. 

Aymaras,  measurements  of  the,  34 ; 
no  grey  hair  among  the,  559 ; 
hairlesbness  of  the  face  in  the,  561  ; 
long  hair  of  the,  580. 

Azara,  on  the  proportion  of  men  and 
women  among  the  Guaranys,  244 ; 
on  Falamedea  cornuta,  366  ;  on  the 
beards  of  the  Guaranys,  561  ;  on 
strife  for  women  among  the  Guanas, 
561  ;  on  infanticide,  577,  592 ;  on 
the  eradication  of  the  eyebrows  and 
eyelashes  by  the  Indians  of  Para- 
guay, 580 ;  on  polyandry  among 
the  Guanas,  593  ;  celibacy  unknown 
among  the  savages  of  South  Ame- 
rica, 594 ;  on  the  freedom  of 
divorce  among  the  Charruas,  598. 


B. 

Babbage,  C,  on  the  greater  proportion 
of  illegitimate  female  births,  244. 

Babirusa,  tusks  of  the,  518. 

Baboon,  revenge  in  a,  69 ;  rage  ex- 
cited in,  by  reading,  71  ;  manifes- 
tation of  memory  by  a,  74 ;  em- 
ploying a  mat  for  shelter  against 
the  sun,  82  ;  protected  from  punish- 
ment by  its  companions,  103. 

■ ,  Cape,  mane  of  the  male,  521  ; 

Hamadryas,  mane  of  the  male, 
521. 

Baboons,  effects  of  intoxicating  li- 
quors on,  7  ;  ears  of,  15  ;  diversity 
of  the  mental  faculties  in,  27;  bands 
of,  50;  habits  of,  51;  variability 
of  the  tail  in,  58  ;  manifestation  of 
maiernal  affection  by,  70 ;  using 
stones  and  sticks  as  weapons,  81  ; 
co-operation  of,  101  ;  silence  of,  on 
plundering  expeditions,  104;  ap- 
parent polygamy  of,  217  ;  poly- 
gamous and  social  habits  of,  590. 

Bachman,  Dr.,  on  the  fertility  of  mu- 
lattoes,  171. 

Baer,  K.  E.  von,  on  embryonic  deve- 
lopment, 9.  I 


Bagehot,  W.,  on  the  social  virtues 
among  primitive  men,  117  ;  slavery 
formerly  beneficial,  117 ;  on  the 
value  of  obedience,  130  ;  on  human 
progress,  132  ;  on  the  persistence 
of  savage  tribes  in  classical  times, 
183. 

Bailly,  E.  M.,  on  the  mode  of  fighting 
of  the  Italian  buffalo,  508  ;  on  the 
fighting  of  stags,  510. 

Bain,  A.,  on  the  sense  of  duty,  98  ; 
aid  springing  from  sympathy,  103  ; 
on  the  basis  of  sympathy,  106  ;  on 
love  of  approbation,  kc,  109  ;  on 
the  idea  of  beauty,  584. 

Baird,  W.,  on  a  difference  in  colour 
between  the  males  and  females  of 
some  Entozoa,  260. 

Baker,  Mr.,  observation  on  the  pro- 
portion of  the  sexes  in  pheasant- 
chicks,  247. 

,  Sir  S.,  on  the  fondness  of  the 

Arabs  for  discordant  music,  380  ;  on 
sexual  difference  in  the  colours  of 
an  antelope,  536  ;  on  the  elephant 
and  rhinoceros  attacking  white  or 
grey  horses,  540  ;  on  the  disfigure- 
ments practised  by  the  negroes, 
541 ;  on  the  gashing  of  the  cheeks 
and  temples  practised  in  Arab 
countries,  574  ;  on  the  coiffure  of 
the  North  Africans,  575  ;  on  the 
perforation  of  the  lower  lip  by  the 
women  of  Latooka,  575  ;  on  the 
distinctive  characters  of  the  coiffure 
of  central  African  tribes,  576  ;  on 
the  coiffure  of  Arab  women,  584. 

"  Balz  "  of  the  Black-cock,  363,  405. 

Bantam,  Sebright,  211,  238. 

Banteng,  horns  of,  505  ;  sexual  dif- 
ferences in  the  colours  of  the,  536.^ 

Banyal,  colour  of  the,  579. 

Barbarism,  primitive,  of  civilised 
nations,  143. 

Barbs,  filamentous,  of  the  feathers, 
in  certain  birds,  385,  430. 

Barrage,  F.,  on  the  Simian  resem- 
blances of  man,  3. 

Barr,  Mr.,  on  sexual  preference  in 
dogs,  524. 

Barrington,  Daines,  on  the  language 
of  birds,  86 ;  on  the  clucking  of 
the  hen,  368  ;  on  the  object  of  the 
song  of  birds,  369  ;  on  the  singing 
of    female    birds,     370 ;    on    birds 


IIAUROW. 


INDEX. 


625 


acquiring  the  songs  of  other  birds, 
370  ;  on  the  muscles  of  the  larynx 
in  song  birds,  371 ;  on  the  want  of 
the  power  of  song  by  female  birds, 
450. 
Barrow,  on  the  widow-bird,  403. 
Bartels,  Dr.,  supernumerary  mammae 
in  men,  37. 

Bartlett,  A.  D.,  period  of  hatching  of 
birds'  eggs,  165  ;  on  the  tragopau, 
220 ;  on  the  development  of  the 
spurs  in  Crossoptilon  auritum,  236  ; 
on  the  fighting  of  the  males  of 
Flectropterus  gambensis,  364 ;  on 
the  knot,  391  ;  on  display  in  male 
birds,  394  ;  on  the  display  of  plu- 
mage by  the  male  J'oli/plectron, 
396  ;  on  Crossoptilon  auritum  and 
Phasianus  Wallichii,  400 ;  on  the 
habits  of  Lophophorus,  420  ;  on  the 
colour  of  the  mouth  in  Buceros 
bicornis,  426  ;  on  the  incubation  of 
the  cassowary,  478  ;  on  the  Cape 
Buffalo,  508  :  on  the  use  of  the 
horns  of  antelopes,  509 ;  on  the 
fighting  of  male  wart-hogs,  520  ; 
on  Ammotragus  tragelaphus,  531  ; 
on  the  colours  of  Cercopit hocus 
cephus,  537  ;  on  the  colours  of  the 
faces  of  monkeys,  550 ;  on  the 
naked  surfaces  of  monkeys,  600. 

Bartram,  on  the  courtship  of  the 
male  alligator,  351. 

Basque  language,  highly  artificial,  91. 

Bate,  C.  S.,  on  the  superior  activity 
of  male  Crustacea,  221  ;  on  the 
proportions  of  the  sexes  in  crabs, 
255 ;  on  the  chelae  of  Crustacea, 
266  ;  on  the  relative  size  of  the 
sexes  in  Crustacea,  268 ;  on  the 
colours  of  Crustacea,  270. 

Bateman,  Dr.,  tendency  to  imitation 
in  certain  diseased  states,  72  ;  on 
Aphasia,  88. 

Bates,  H.  W.,  on  variation  in  the 
form  of  the  head  of  Amazonian 
Indians,  28  ;  on  the  proportion  of 
the  sexes  among  Amazonian  but- 
terflies, 250  ;  on  sexual  differences 
in  the  wings  of  butterflies,  277  ; 
on  the  field  -  cricket,  283 ;  on 
Pijrodes  pAilcherrimus,  294  ;  on  the 
horns  of  Lamellicorn  beetles,  295, 
297 ;  on  the  colours  of  Epicalice, 
&c.,    309 ;    on   the    coloration    of 


tropical  butterflies,  311  ;  on  the 
variability  of  Pajyilio  Sesostris  and 
Childrence,  320 ;  on  male  and  fe- 
male butterflies  inhabiting  ditTerent 
stations,  321  ;  on  mimicry,  323 ; 
on  the  caterpillar  of  a  Sphinx,  325  ; 
on  the  vocal  organs  of  the  um- 
brella-bird, 374 ;  on  the  toucans, 
492  ;  on  Brachyurus  calvus,  550. 

Batokas,  knocking  out  two  upper  in- 
cisors, 575. 

Batrachia,  349  ;  eagerness  of  male, 
221. 

Bats,  scent-glands,  529:  sexual  dif- 
fei-ences  in  the  colour  of,  534  •,  fur 
of  male  frugivorous,  534. 

Battle,  law  of,  144;  among  beetles, 
299 ;  among  birds,  360  ;  among 
mammals,  500  et  seq. ;  in  man, 
561. 

Beak,  sexual  difference  in  the  forms  of 
the,  359  ;  in  the  colour  of  the,  383. 

Beaks,  of  birds,  bright  colours  of,  491. 

Beard,  development  of,  in  man,  557  ; 
analogy  of  the,  in  man  and  the 
quadrumana,  558  ;  variation  of  the 
development  of  the,  in  different 
races  of  men,  559  ;  estimation  of, 
among  bearded  nations,  581 ;  pro- 
bable origin  of  the,  602. 

,  in  monkeys,  150  ;  of  mammals, 

531. 

Beautiful,  taste  for  the,  in  birds,  410  ; 
in  the  quadrumana,  540. 

Beauty,  sense  of,  in  animals,  92  ;  ap- 
preciation of,  by  birds,  413  ;  in- 
fluence of,  573,  576  ;  variability  of 
the  standard  of,  596. 

Beavan,  Lieut.,  on  the  development 
of  the  horns  in  Cervus  Eldi,  234. 

Beaver,  instinct  and  intelligence  of 
the,  67,  68;  voice  of  the,  527; 
castoreum  of  the,  529. 

Beavers,  battles  of  male,  500. 

Bechstein,  on  female  birds  choosing 
the  best  singers  among  the  males, 
368  ;  on  rivalry  in  song-birds,  369  ; 
on  the  singing  of  female  birds,  37u  ; 
on  birds  acquiring  the  songs  of 
other  birds,  370 ;  on  pairing  the 
canary  and  siskin,  415;  on  a  .sub- 
variety  of  the  monk  pigeon,  427 ; 
on  spurred  hens,  449. 

Beddoe,  Dr.,  on  causes  of  difference  m 
stature,  31. 


626 


EKIO-EATKR. 


INDEX. 


Bee-eater,  371. 

Bees,  99 ;  pollen-baskets  and  stings 
of,  63 ;  destruction  of  drones  and 
queens  by,  106  ;  female,  secondary 
sexual  chavacters  of,  208 ;  propor- 
tion of  sexes,  254;  difference  of 
the  sexes  in  colour  and  sexual 
selection,  292. 

Beetle,  luminous  larva  of  a,  277. 

Beetles,  294- ;  size  of  the  cerebral 
ganglia  in,  51-;  dilatation  of  the 
fore  tarsi  in  male,  275  ;  blind,  29-1 ; 
stridulation  of,  301. 

Belgium,  ancient  inhabitants  of,  182. 

Bell,  Sir  C,  on  emotional  inuscles  in 
man,  3;  "snarling  muscles,"  41; 
on  the  hand,  51. 

,  T.,  on  the  numerical  proportion 

of  the  sexes  in  moles,  247  ;  on  the 
newts,  348  ;  on  the  croaking  of  the 
frog,  350 ;  on  the  difference  in  the 
coloration  of  the  sexes  in  Zootoca 
vivipara,  357 ;  on  moles  fighting, 
500. 

Bell-bird,  sexual  difference  in  the 
colour  of  the,  389. 

Bell-birds,  colours  of,  492. 

Belt,  Mr.,  on  the  nakedness  of  tropical 
mankind,  57  ;  on  a  spider-moukey 
and  eagle,  102  ;  habits  of  ants,  147  ; 
Lampyrida>  distasteful  to  mammals, 
277  ;  mimicry  of  Zejjtalides,  325 ; 
colours  of  Nicaraguan  frogs,  349  ; 
display  of  humming-birds,  443  ;  on 
the  toucans,  492 ;  protectiA'e  co- 
louring of  skunk,  543. 

Benevolence,  manifested  by  birds, 
411. 

Bennett,  A.  W.,  attachment  of  mated 
birds,  411:  on  the  habits  of  D,o- 
mccus  irroraiu-^,  478. 

,     Dr.,    on    birds     of    paradise, 

396. 

Berbers,  fertility  of  crosses  with 
other  races,  171. 

iJcniicla  antarctica  colours  of,  492. 

Beruicle  gander  pairing  with  a  Canada 
goose,  414. 

liort,  M.,  crustaceans  distinguish 
colours,  271. 

Bertillon,  M.,  arrested  development 
anil  polydactylism,  37. 

Bettoui,  E.,  on  local  differences  in 
the  nests  of  Italian  birds,  456. 

B3yle,  AI.,  set'  Bombet. 


Bhoteas,  colour  of  the  beard  in,  558, 

Bhringa,  disciform  tail-feathers  of, 
392. 

Bianconi,  Prof.,  on  structures  as  ex- 
plained through  mechanical  prm- 
ciples,  24. 

Bibio,  sexual  differences  in  the  genus, 
280. 

Bichat,  on  beauty,  585. 

Bickes,  proportion  of  sexes  in  man, 
243. 

Bile,  coloured,  in  many  animals,  261. 

Bimana,  149. 

Birds,  imitations  of  the  songs  of  other 
birds  by,  73  ;  dreaming,  74  ;  killed 
by  telegi-aph  wires,  80 ;  language 
of,  86  ;  sense  of  beauty  in,  92 ; 
pleasure  of,  in  incubation,  105 ; 
male,  incubation  by,  1 63  ;  and  rep- 
tiles, alliance  of,  165  ;  sexual  dif- 
ferences in  the  beak  of  some,  208  ; 
migratory,  arrival  of  the  male  be- 
fore the  female,  212 ;  appareiit 
relation  between  polygamy  and 
marked  sexual  differences  in,  220 ; 
monogamous,  becoming  polygamous 
iinder  -domestication,  220  ;  eager- 
ness of  male  in  pursuit  of  the  fe- 
male, 221  ;  wild,  numerical  propor- 
tion of  the  sexes  in,  247  ;  secondary 
sexual  characters  of,  358  ;  differ- 
ence of  size  in  the  sexes  of,  362; 
fights  of  male,  witnessed  by  females, 
367  ;  display  of  male,  to  ca})tivate 
the  females,  367  ;  close  attention 
of,  to  the  songs  of  others,  368  ;  ac- 
quiring the  song  of  their  foster- 
parents,  370  ;  brilliant,  rarely  good 
songsters,  371;  love -antics  and 
dances  of,  380 ;  coloration  of,  385 
et  seq. ;  moulting  of,  390  ct  seq.  ; 
unpaired,  407  ;  male,  singing  out 
of  season,  409  ;  mutual  affection  of, 
410 ;  in  confinement,  distinguish 
persons,  411;  hybrid,  production 
of,  414;  Albino,  419;  European, 
number  of  species  of,  422 ;  varia- 
bility of,  422  ;  geographical  distri- 
bution of  colouring,  422  ;  gradation 
of  secondary  sexual  characters  in, 
430;  obscurely  coloured,  building 
concealed  nests,  454  ;  young  female, 
acquiring  male  characters,  462; 
breeding  in  immature  plumage, 
484;    moulting  of,   484;    aquatic, 


INDEX. 


627 


frequency  of  white  plumage  in, 
493 ;  vocal  courtship  of,  567 ; 
naked  skin  of  the  head  and  neck  in, 
601. 

Birgus  latro,  habits  of,  270. 

Birkbeck,  Mr.,  on  the  finding  of  new 
mates  by  golden  eagles,  408. 

Birthplace  of  man,  155. 

Births,  numerical  proportions  of  the 
sexes  in,  in  animals  and  man,  215, 
216;  male  and  female,  numerical 
proportion  of,  in  England,  242. 

Bischoff,  Prof,  on  the  agreement  be- 
tween the  brains  of  man  and  of  the 
orang,  6  ;  figure  of  the  embryo  of 
the  dog,  10;  on  the  convolutions 
of  the  brain  in  the  human  foetus, 
11  ;  on  the  difference  between  the 
skulls  of  man  and  the  quadrumana, 
149  ;  resemblance  between  the  ape's 
and  man's,  200. 

Bishop^  J.,  on  the  vocal  organs  of 
frogs,  350  ;  on  the  vocal  organs  of 
corvine  birds,  370  ;  on  the  trachea 
of  the  Merganser,  374. 

Bison,  American,  co-operation  of,  101; 
mane  of  the  male,  521. 

Bitterns,  dwarf,  coloration  of  the 
sexes  of,  461. 

Biziura  lobata,  musky  odour  of  the 
male,  359  ;  large  size  of  male,  362. 

Blackbird,  sexual  differences  in  the, 
219 ;  proportion  of  the  sexes  in 
the,  348  ;  acquisition  of  a  song  by, 
370 ;  colour  of  the  beak  in  the 
sexes  of  the,  383,  491 ;  pairing 
with  a  thrush,  414 ;  colours  and 
nidification  of  the,  455 ;  young  of 
the,  487  ;  sexual  difference  in  colo- 
ration of  the,  491. 

Black-buck,  Indian,  sexual  difference 
in  the  colour  of  the,  535. 

Blackcap,  arrival  of  the  male,  before 
the  female,  212;  young  of  the, 
487. 

Black-cock,  polygamous,  219;  pro- 
portion of  the  sexes  in  the,  248 ; 
pugnacity  and  love-dance  of  the, 
363  ;  call  of  the,  375  ;  moulting  of 
the,  392;  duration  of  the  court- 
ship of  tlie,  405;  and  pheasant, 
hybrids  of,  414 ;  sexual  difference 
in  coloration  of  the,  491;  crimson 
eye-cere  of  the,  491. 

Blacklock,  Dr.,  on  music,  572. 


Black-grouse,  characters  of  young, 
465,  471. 

Blackwall,  J.,  on  the  speaking  of  the 
m.agpie,  90;  on  the  desertion  of 
their  young  by  swallows,  108  ;  on 
the  superior  activity  of  male  spiders, 
221  ;  on  the  proportion  of  the 
sexes  in  spiders,  254;  on  sexual 
variation  of  colour  in  spiders,  272  ; 
on  male  spiders,  272.    • 

Bladder-nose  Seal,  hood  of  the,  528. 

Blaine,  on  the  aflections  of  dogs, 
523. 

Blair,  Dr.,  on  the  relative  liability  of 
Europeans  to  yellow  fever,  194. 

Blake,  C.  C,  on  the  jaw  from  La 
Naulette,  40. 

Blakiston,  Capt.,  on  the  American 
snipe,  377  ;  on  the  dances  of  Tetrao 
phasianellus,  381. 

Blasius,  Dr.,  on  the  species  of  Euro- 
pean birds,  422. 

Blcdius  taurus,  hornlike  processes  of 
male,  299. 

Bleeding,  tendency  to  profuse,  237. 

Blenkiron,  Mr.,  on  sexual  preference 
in  horses,  524. 

Blennies,  crest  developed  on  the  head 
of  male,  during  the  breeding  sea- 
son, 338. 

Bletkisa  multipiinctata,  stridulation 
of,  302. 

Bloch,  on  the  proportions  of  the  sexes 
in  fishes,  249. 

Blood,  arterial,  red  colour  of,  261. 

pheasant,  number  of  spurs  in, 

364. 

Bluobreast,  red-throated,  sexual  dif- 
ferences of  the,  472. 

Blumeubach,    on    ]\Ian,    28 ;    on   the 
large  size  of  the  nasal  cavities  in  . 
American    aborigines,  34;    on    the 
position  of  man,  149  ;  on  the  num- 
ber of  species  of  man,  174. 

Blyth,  E.,  on  the  structure  of  the 
hand  in  the  species  of  Hylobates, 
51  ;  observations  on  Indian  crows, 
102;  on  the  development  of  the 
horns  in  the  Koodoo  and  Eland  an- 
telopes, 234;  on  the  pugnacity  of  the 
males  of  Gallinula  cristata,  360  ;  on 
the  presence  of  spurs  in  the  female 
Exqjlocamus  erijthrophthalmus,  364 ; 
on  the  pugnacity  of  the  amadavat, 
366  ;  on  the  spoonbill,  374  ;  on  the 


62S 


BOAR. 


INDEX. 


BOURBOX. 


moulting  of  Anthus,  392 ;  on  the 
moulting  of  bustards,  plovers,  and 
Oallus  bunkiva,  392 ;  on  the  Indian 
honey-buzzard,  424  ;  on  sexual  dif- 
ferences in  the  colour  of  the  eyes 
of  hornbills,  425  ;  on  Oriolus  me- 
lanocephalus,  460 ;  on  PalcBornis 
javanicus,  461;  on  the  genus  Ar- 
detta,  461  ;  on  the  peregrine  folcon, 
461  ;  on  young  female  birds  acquir- 
ing male  characters,  461 ;  on  the 
immature  plumage  of  birds,  465 ; 
on  representative  species  of  birds, 
468  ;  on  the  young  of  Turnix,  476 ; 
on  anomalous  young  of  Lanius 
rufus  and  Colymbus  glaoialis,  482  ; 
on  the  sexes  and  young  of  the  spar- 
rows, 483  ;  on  dimorphism  in  some 
herons,  484 ;  on  the  ascertainment 
of  the  sex  of  nestling  bullfinches 
by  pulling  out  breast-feathers,  484  ; 
on  orioles  breeding  in  immature 
plumage,  484 ;  on  the  sexes  and 
young  of  Buphus  and  Anastomus, 
486  ;  on  the  young  of  the  black- 
cap and  blackbird,  487 ;  on  the 
young  of  the  stonechat,  487;  on 
the  white  plumage  of  Anastomus, 
493 ;  on  the  horns  of  Bovine  ani- 
mals, 505  ;  on  the  horns  oi  Antiloje 
bezoartica,  bOl ;  on  the  mode  of 
fighting  of  Otis  cycloceros,  508 ; 
jjn  the  voice  of  the  Gibbons,  527  ; 
on  the  crest  of  the  male  wild  goat, 
531 ;  on  the  colours  of  Portax 
pkta,  535  ;  on  the  colours  of  Anti- 
lope  bezoartica,  536  ;  on  the  colour 
of  the  Axis  deer,  536  ;  on  sexual 
difference  of  colour  in  Jli/lobates 
hoolock,  537  ;  on  the  hog-deer,  546  ; 
on  the  beard  and  whiskers  in  a 
monkey  becoming  white  with  age, 
550. 

Boar,  wild,  polygamous  in  India,  217  ; 
use  of  the  tu.sks  by  the,  513  ;  fight- 
ing of,  518. 

Boai'dman,  Mr.,  Albino  birds  in  U.  S., 
419. 

Boitard  and  Corbit?,  MM.,  on  the 
transmission  of  sexual  peculiarities 
in  pigeons,  230 ;  on  the  antipathy 
shewn  by  some  female  pigeons  to 
certain  males,  418. 

Bold,  Mr.,  on  the  singing  of  a  sterile 
hybrid  canary,  369. 


Bombet,    on    the    variability    of   the 

standard  of  beauty  in  Europe,  596. 
Bombus,    difference    of  the  sexes   in, 

292. 
Bombycidae,  coloration  of,  313  ;  pair- 
ing of  the,  318  ;  colours  of,  318. 
Bombycilla   carolmensis,    red  append- 
ages of,  461. 
Bombyx  cynthia,  278 ;  proportion  of 

the  sexes  in,  250,  253  ;  pairing  of, 

318. 
mori,   difference  of  size    of  the 

male  and  female  cocoons  of,  278  ; 

pairing  of,  318. 
Pernyi,  proportion  of  sexes  of, 

253. 
Yamamai,  278  ;  M.  Personnat  on, 

251  ;  proportion  of  sexes  of,  253. 
Bonaparte,  C.  L.,  on  the  call-notes  of 

the  wild  turkey,  375. 
Bond,  F.,  on  the  finding  of  new  mates 

by  crows,  408. 
Bone,  implements  of,  skill  displayed 

in  making,  49. 
Boner,  C,    on    the    transfer  of  male 

characters  to  an  old  female  chamois, 

504;  on  the  habits  of  stags,  515  ; 

on  the  pairing  of  red  deer,  522. 
Bones,    increase    of,    in    length    and 

thickness,  when  carrying  a  greater 

weight,  32. 
Bonizzi,   P.,  difference    of   colour    in 

sexes  of  pigeons,  230. 
Bonnet  monkej^,  151. 
Bonwick,   J.,    extinction    of    Tasma- 

nians,  183,  184. 
Boomerang,  145. 
Boreus  hyemalis,  scarcity  of  the  male, 

254. 
Bory  St.  Vincent,  on  the  number  of 

species  of  man,  174;  on  the  colours 

of  Labrus  pavo,  342. 
Bos  etruscus,  505. 

gaurus,  horns  of,  505. 

moschatus,  529. 

primigenius,  501. 

sondaicus,  horns  of,  505  ;  colours 

of,  536. 
Botocudos,  144;  mode  of  life  of,  197  ; 

disfigurement  of  the  ears  and  lower 

lip  of  the,  575. 
Boucher  de  Perthes,  J.  C.  de,  ou  the 

antiquity  of  man,  2. 
Bourbon,  proportion  of  the  sexes  in  a 

species  of  Papilio  from,  250. 


INDEX. 


BROCA. 


620 


Bourien,  on  the  marriage-customs  of 
the  savages  of  the  Malay  Archi- 
pelago, 598. 

Bovidae,  dewlaps  of,  531. 

Bower-birds,  406  ;  habits  of  the,  381 ; 
ornamented  playing-places  of,  92, 
413. 

Bows,  use  of,  179. 

Brachycephalic  structure,  possible 
explanation  of,  56. 

Brachyura,  268. 

Brachyurus  calvus,  scarlet  face  of, 
550. 

Bradley,  Mr.,  abductor  ossis  metatarsi 
quinti  in  man,  42. 

Brain,  of  man,  agreement  of  the,  with 
that  of  lower  animals,  6  ;  convolu- 
tions of,  in  the  human  foetus,  1 1 ; 
influence  of  development  of  mental 
faculties  upon  the  size  of  the,  54 ; 
influence  of  the  development  of,  on 
the  spinal  column  and  skull,  55  ; 
larger  in  some  existing  mammals 
than  in  their  tertiary  prototypes, 
81  ;  relation  of  the  development  of 
the,  to  the  progress  of  language, 

87  ;  disease  of  the,  affecting  speech, 

88  ;  difference  in  the  convolutions 
of,  in  different  races  of  men,  167  ; 
supplement  on,  by  Prof.  Huxley, 
199  ;  development  of  the  gyri  and 
sulci,  204. 

Brakenridge,  Dr.,  on  the  influence  of 

climate,  32. 
Brandt,  A.,  on  hairy  men,  1 9. 
Braubach,  Prof.,  on  the  quasi-religious 

feeling  of  a  dog  towards  his  master, 

96  ;    on  the  self-restraint  of  dogs, 

103. 
Brauer,  F.,  on  dimorphism  in  Neuro- 

themis,  291. 
Brazil,  skulls  found  in  caves  of,  168  ; 

population  of,  173  ;  compression  of 

the  nose  by  the  natives  of,  583. 
Break  between  man   and   the   apes, 

156. 
Bream,  proportion  of  the  sexes  in  the, 

249. 
Breeding,  age  of,  in  birds,  484. 
season,  sexual  characters  making 

their  appearance   in   the,  in  birds, 

390. 
Brehm,  on  the  effects  of  intoxicating 

liquors    on    monkeys,    7 ;    on    the 

recognition  of  women  by  male  Cyno- 


ccphali,  8 ;  on  the  diversity  of  the 
mental  foculties  of  monkeys,  27 ; 
on  the  habits  of  baboons,  51  ;  on 
revenge  taken  by  monkeys,  69;  on 
manifestations  of  maternal  affection 
by  monkeys  and  baboons,  70  ;  on 
the  instinctive  dread  of  monkeys 
for  serpents,  71 ;  on  the  use  of 
stones  as  missiles  by  baboons,  81  ; 
on  a  baboon  using  a  mat  for  shelter 
from  the  sun,  82  ;  on  the  signal- 
cries  of  monkeys,  87  ;  on  sentinels 
posted  by  monkeys,  101;  on  co-ope- 
ration of  animals,  101 ;  on  an  eagle 
attacking  a  young  Cercopithecus, 
101 ;  on  baboons  in  confinement  pro- 
tecting one  of  their  number  from 
punishment,  103 ;  on  the  habits 
of  baboons  when  plundering,  104 ; 
on  polygamy  in  Cynoceplialus  and 
Cebus,  217;  on  the  numerical  pro- 
portion of  the  sexes  in  birds,  247  ;  on 
the  love-dance  of  the  black-cock, 
363  ;  on  Palamedea  cornuta,  366  ; 
on  the  habits  of  the  Black-grouse, 
366 ;  on  sounds  produced  by  birds 
of  paradise,  376  ;  on  assemblages  of 
grouse,  405  ;  on  the  finding  of  new 
mates  by  birds,  409 ;  on  the  fight- 
ing of  wild  boars,  518 ;  on  the 
habits  of  Cynoceplialus  hamadryas, 
590. 

Brent,  Mi*.,  on  the  courtship  of  fowls, 
417. 

Breslau,  numerical  proportion  of  male 
and  female  births  in,  243. 

Bridgman,  Laura,  88. 

Brimstone  butterfly,  312  ;  sexual  dif- 
ference of  colour  in  the,  322. 

British,  ancient,  tattooing  practised 
by,  574, 

Broca,  Prof.,  on  the  occurrence  of  the 
supra-condyloid  foramen  in  the 
human  humerus,  22 ;  anthropo- 
morphous apes  more  bipedal  than 
quadrupedal,  53;  on  the  capacity  of 
Parisian  skulls  at  different  periods, 
55;  comparison  of  modern  and 
mediaeval  skulls,  55 ;  on  tails  of 
quadrupeds,  58  ;  on  the  influence  of 
natural  selection,  61 ;  on  hybridity 
in  man,  170 ;  on  human  remains 
from  Les  Eyzies,  181  ;  on  the  cause 
of  the  difference  between  Europeans 
and  Hindoos,  192. 


630 


INDEX. 


BURCHELL. 


Brodie,  Sir  B.,  on  the  origin  of  the 
moral  sense  in  man,  98. 

Bronn,  H.  G.,  on  the  copulation  of 
insects  of  distinct  species,  275. 

Bronze  period,  men  of,  in  Europe, 
128. 

Brown,  R.,  sentinels  of  seals  generally 
females,  100;  on  the  battles  of 
seals,  500 ;  on  the  narwhal,  502 ; 
on  the  occasional  absence  of  the 
tusks  in  the  female  walrus,  502  ; 
on  the  bladder-nose  seal,  528 ;  on 
the  colours  of  the  sexes  in  Phoca 
Groenlandica,  535 ;  on  the  appre. 
ciation  of  music  by  seals,  569  ;  on 
plants  used  as  love-philters,  by 
North  American  women,  577. 

Brown,  Dr.  Crichton,  injury  to  infants 
during  parturition,  244. 

Brown-Sequard,  Dr.,  on  the  inheri- 
tance of  the  effects  of  operations  by 
guinea-pigs,  60,  603. 

Bruce,  on  the  use  of  the  -elephant's 
tusks,  507. 

Brulerie,  P.  de  la,  on  the  habits  of 
Ateuchus  cicatricosus,  300  ;  on  the 
stridulation  of  Ateuchus,  306. 

Briinnich,  on  the  pied  ravens  of  the 
Feroe  islands,  424. 

Bryant,  Dr.,  preference  of  tame  pigeon 
for  wild  mate,  418. 

,    Capt,    on    the    courtship    of 

Callorhinus  xusinus,  522, 

Bubas  bison,  thoracic  projection  of, 
298. 

Bucephalus  capensis,  difference  of  the 
sexes  of,  in  colour,  351. 

Buceros,  nidification  and  incubation  of, 
454. 

• bicornis,  sexual  differences  in  the 

colouring  of  the  casque,  beak,  and 
mouth  in,  425, 

' corrugatu!^,  sexual  dilTerences  in 

the  beak  of,  383. 

Biichner,  L.,  on  the  origin  of  man, 
3  ;  on  the  use  of  the  human  foot  as 
a  prehensile  organ,  52  ;  on  the  mode 
of  i)rogression  of  the  apes,  52  ;  on 
want  of  self-consciousness,  &c,,  in 
savages,  83. 

Bucholz.,  Dr.,  quarrels  of  chameleons, 
357. 

Buckhand,  F.,  on  the  numerical  pro- 
portion of  the  sexes  in  rats,  247; 
on    the   proportion  of  the  sexes  in 


strosa,  338. 
Buckland,  W.,  on  the  complexity  of 

crinoids,  91. 
Buckler,  W.,   proportion  of  sexes  of 

Lepidoptera  reared  by,  253. 
Buckinghamshire,  numerical  propor- 
tion of  male  and  female  births  in,  242. 
Bucoi'ax  abyssinicus,  inflation  of  the 

neck-wattle    of    the    male    during 

courtship,  383. 
Budytes  Bait,  212. 
Buffalo,  Cape,  508. 

,  Indian,  horns  of  the,  505. 

,  Italian,  mode  of  fighting  of  the, 

508. 
Buffon,  on  the  number  of  species  of 

man,  174. 
Bufo  sikimmensis,  349. 
Bugs,  281. 
Buist,  R.,  on  the  proportion   of  the 

sexes  in  salmon,  249;  on  the  pug- 
nacity of  the  male  salmon,  332. 
Bulbul,  puo;nacity  of  the  male,  360 ; 

display  of  under  tail-coverts  by  the 

male,  402. 
Bull,  mode  of  fighting  of  the,  508  ; 

curled  frontal  hair  of  the,  531. 
Buller,  Dr.,  on  the   Huia,   208  ;    the 

attachment  of  birds,  410. 
Bullfinch,  sexual  differences    in  the, 

219;  piping,  369;    female,  singing 

of  the,  370  ;  courtship  of  the,  401 ; 

widowed,  finding  a  new  mate,  408  ; 

attacking     a     reed-bunting,    412; 

nestling,  sex  ascertained  by  pulling 

out  breast-feathers,  484, 
Bullfinches     distinguishing    persons, 

412  ;  rivalry  of  female,  42U. 
Bulls,  two   young,  attacking  an  old 

one,  101  ;  wild,  battles  of,  501. 
Bull-trout,  male,  colouring  of,  during 

the  breeding  season,  340. 
Bunting,  reed,  head   feathers  of  the 

male,  402  ;  attacked  by  a  bullfinch, 

412. 
Buntings,  characters  of  young,  464. 
Buphus  coromandus,  sexes  and  young 

of,  486 ;  change  of  colour  in,  494, 

495. 
Burchell,  Dr.,  on  the  zebra,  545  ;  on 

the  extravagance  of  a  bush  woman 

in  adorning  herself,  577  ;  celibacy 

unknown    among    the    savages   of 

South  Africa,  594;   on   the   naar- 


BURKE. 


INDEX. 


CANINE. 


031 


riage-customs  of  the  Bushwomen, 
598. 

Burke,  on  the  number  of  species  of 
man,  174. 

Burmese,  colour  of  the  beard  in,  358. 

Burton,  Capt.,  on  negro  ideas  of 
female  beauty,  579  ;  on  a  universal 
ideal  of  beauty,  582. 

Bushmen,  64. 

Bushwoman,  extravagant  ornamenta- 
tion of  a,  577. 

Bushwomen,  hair  of,  107  ;  marriage- 
customs  of,  598. 

Bustard,  throat-pouch  of  the  male, 
373  ;  humming  noise  produced  by  a 
male,  377  ;  Indian,  ear-tufts  of,  384. 

Bustards,  occurrence  of  sexual  dif- 
ferences and  of  polygamy  among 
the,  219  ;  love-gestures  of  the  male, 
380;  double  moult  in,  390,  392. 

Butler,  A.  G.,  on  sexual  differences  in 
the  wings  of  Aricoris  epitus,  277  ; 
courtship  of  butterflies,  307 ;  on 
the  colouring  of  the  sexes  in 
species  of  Thecia,  310 ;  on  the  re- 
semblance of  fyhias  glaucippe  to  a 
leaf,  313  ;  on  the  rejection  of  cer- 
tain moths  and  caterpillars  by 
lizards  and  frogs,  326. 

Butterfly,  noise  produced  by  a,  307 ; 
Emperor,  307,  308  ;  meadow  brown, 
instability  of  the  ocellated  spots  of, 
428. 

Butterflies,  proportion  of  the  sexes  in, 
250 ;  forelegs  atrophied  in  some 
male,  277  ;  sexual  difference  in  the 
neuration  of  the  wings  of,  277 ; 
pugnacity  of  male,  307  ;  protective 
resemblances  of  the  lower  surface 
of,  311  ;  display  of  the  wings  by, 
314;  white,  alighting  upon  bits  of 
paper,  317  ;  attracted  by  a  dead 
.specimen  of  the  same  species,  317  ; 
courtship  of,  317  ;  male  and  female, 
inhabiting  different  stations,  321. 

Buxton,  C,  observations  on   macaws, 
102  ;  on  an  instance  of  benevolence 
in  a  parrot,  411. 
Buzzard,  Indian  honey-,  variation  in 
the  crest  of,  424. 


Cabbage  butterflies,  312. 
Cachalot,  larsre  head  of  the  mak 


502. 


Cadences,  musical,  perception  of,  by 
animals,  569. 

Ca3cum,  20;  large,  in  the  early  pro- 
genitors of  man,  160. 

Cairina  moschata,  pugnacity  of  the 
male,  362. 

Californian  Indians,  decrease  of,  258. 

CalUanassa,  chelas  of,  figured,  267. 

Callidnjas,  colours  of  sexes,  318. 

Callionymus  lyra,  characters  of  the 
male,  335. 

Callorhinus  ursinus,  relative  size  of 
the  sexes  of,  515 ;  courtship  of, 
522. 

Calotes  maria,  358. 

nigrilabris,  sexual   difference   in 

the  colour  of,  357. 

Cambridge,  O.  Pickard,  on  the  sexes 
of  spiders,  255  ;  on  the  size  of  male 
Nephila,  273. 

Camel,  canine  teeth  of  male,  502, 
514. 

Campbell,  J.,  on  the  Indian  elephant, 
218;  on  the  proportion  of  male 
and  female  births  in  the  harems  of 
Siam,  245. 

Campi/lopterus  Iiemileuciirus,  248. 

Canaries  distinguishing  persons,  412 

Canary,  polygamy  of  the,  220  ;  change 
of  plumage  in,  after  moulting,  238  ; 
female,  selecting  the  best  singing 
male,  268 ;  sterile  hybrid,  singing 
of  a,  369 ;  female,  singing  of  the, 
370 ;  selecting  a  greenfinch,  415  ; 
and  siskin,  pairing  of,  415. 

Canestrini,  G.,  on  rudimentary  cha- 
racters and  the  origin  of  man,  3  ; 
on  rudimentary  characters,  11;  on 
the  movement  of  the  ear  in  man, 
14 ;  on  the  variability  of  the  ver- 
miform appendage  in  man,  21;  on 
the  abnormal  division  of  the  malar 
bone  in  man,  39 ;  on  abnormal  con- 
ditions of  the  human  uterus,  39 ; 
on  the  persistence  of  the  frontal 
suture  in  man,  39  ;  on  the  propor- 
tion of  the  sexes  in  silk-moths,  250, 
251  ;  secondary  sexual  characters 
of  spiders,  272. 

Cancer  pagu>-us,  266. 

Canfield,  Dr.,  on  the  horns  of  the 
Antilocapra  234. 

Canine  teeth  in  man,  46  ;  diminution 
of,  in  man,  53 ;  diminution  of,  in 
horses,    53 ;    disappearance    of,    in 


632 


CANINES. 


INDEX. 


male  ruminants,  53 ;  Urge,  in  the 
eai'ly  progenitors  of  man,  IGO. 

Canines,  and  horns,  inverse  develop- 
ment of,  614. 

Canoes,  use  of,  48,  180. 

Cantharis,  difference  of  colour  in  the 
sexes  of  a  species  of,  294. 

Cantharus  lineatus,  341. 

Capercailzie,  polygamous,  219  ;  pro- 
portion of  the  sexes  in  the,  248  ; 
pugnacity  of  the  male,  363  ;  pairing 
of  the,  367  ;  autumn  meetings  of 
the,  370 ;  call  of  the,  375 ;  dura- 
tion of  the  courtship  of,  405  ;  beha- 
viour of  the  female,  419;  inconve- 
nience of  black  colour  to  the  female, 
444  ;  sexual  difference  in  the  colora- 
tion of  the,  491;  crimson  eye-cere 
of  the  male,  491. 

Capitonidse,  colours  and  nidification  of 
the,  455. 

Capra  cegagrus,  508 ;  crest  of  the 
male,  531  ;  sexual  difference  in  the 
colour  of,  536. 

Capreolus  Sibirions  suhecaudatus,  542. 

Caprice,  common  to  man  and  animals, 
93. 

Caprimulgus,  noise  made  by  the  males 
of  some  species  of,  with  their 
wings,  376. 

virginianus,  pairing  of,  366. 

Carabidse,  302. 

Carbonnier,  on  the  natural  history  of 
the  pike,  249  ;  on  the  relative  size 
of  the  sexes  in  fishes,  335 ;  court- 
ship of  Chinese  Macropus,  341. 

Carcineutes,  sexual  difference  of 
colour  in,  457. 

Cafcinus  mcenas,  268,  269. 

Cardinalis  virginianus,  225. 

Carduelis  elegans,  sexual  differences  of 
the  beak  in,  360. 

Carnivora,  marine,  polygamous  habits 
of,  218;  sexual  differences  in  the 
colours  of,  534. 

Carp,  numerical  proportion  of  the 
sexes  in  tlie,  249. 

Carr,  K.,  on  the  peewit,  366. 

Carrier  pigeon,  late  development  of 
the  wattle  in  the,  238. 

Carrion  beetles,  stridulation  of,  302. 

Cams,  Prof.  V.,  on  the  development 
of  the  horns  in  merino  sheep,  235. 

Cassowary,  sexes  and  incubation  of 
the,  478. 


Castina,  mode  of  holding  wings,  315. 

Castoreum,  529. 

Casuarius  galeatus,  478. 

Cat,  convoluted  body  in  the  extremity 
of  the  tail  of  a,  23  ;  sick,  sympathy 
of  a  dog  with  a,  103. 

Cataract  in  Cebiis  Azarce,  7. 

Catarrh,  liability  of  Cebus  Azarce  to,  7. 

Catarrhine  monkeys,  153. 

Caterpillars,  bright  colours  of,  325. 

Cathartes  aura,  416. 

jota,  love-gestures  of  the  male, 

380. 

Catlin,  G.,  correlation  of  colour  and 
texture  of  hair  in  the  Mandans, 
197;  on  the  development  of  the 
beard  among  North  American 
Indians,  660 ;  on  the  great  length 
of  the  hair  in  some  North  American 
tribes,  680. 

Caton,  J.  D.,  on  the  development  of 
the  horns  in  Cervus  virginianus  and 
strongyloceros,  234  ;  on  the  presence 
of  traces  of  horns  in  the  female 
wapiti,  504 ;  on  the  fighting  of 
deer,  610  ;  on  the  crest  of  the  male 
wapiti,  531  ;  on  the  colours  of  the 
Virginian  deer,  535  ;  on  sexual  dif- 
ferences of  colour  in  the  wapiti, 
636  ;  on  the  spots  of  the  Virginian 
deer,  546. 

Cats,  dreaming,  74;  tortoise-shell,  230, 
232,  237  ;  enticed  by  valerian,  530  ; 
colours  of,  543, 

Cattle,  rapid  increase  of,  in  South 
America,  47  ;  domestic,  lighter  in 
winter  in  Siberia,  229 ;  horns  of,  234, 
506  ;  domestic,  sexual  differences  of, 
late  developed,  238  ;  numerical  pro- 
portion of  the  sexes  in,  246. 

Caudal  vertebrae,  number  of,  in  ma- 
caques and  baboons,  68 ;  basal,  of 
monkeys,  imbedded  in  the  body,  59. 

Cavolini,  observations  on  ^erranus, 
162. 

Cebus,  maternal  affection  in  a,  70; 
gradation  of  species  of,  175. 

—  ApcUa,  205. 

Azara;,  liability  of,  to  the  same 

diseases  as  man,  7  ;  distinct  sounds 
produced  by,  84 ;  early  maturity 
of  the  female,  558. 

capucimis,    polygamous,    217; 

sexual  differences  of  colour  in,  537  ; 
hair  on  the  head  of,  549. 


INDEX. 


CIIELONIA. 


633 


Ccbus  vcllerosus,  hair  on  the  head  of, 

549. 
Cecidomyidae,  proportions  of  the  sexes 

in,  354. 
Celibacy,  unknown  among  the  savages 

of  South  Africa  and  South  America, 

594. 
Centipedes,  274. 
Cephalopoda,    absence    of    secondary 

sexual  characters  in,  263. 
Cephalopterus  ornatus,  373. 

• penduliger,  374. 

Cerambyx  hews,  stridulant  organ  of, 

303. 
Ceratodus,  paddle  of,  37. 
Ceratophora  aspera,  nasal  appendages 

of,  355. 

Stoddartii,  nasal  horn  of,  355. 

Cerceris,  habits  of,  291. 

Cercocebus  cethiojJS,  whiskers,  &c.,  of, 

550. 
Cercopithecus,    young,    seized    by    an 

eagle    and  rescued    by    the   troop, 

101 ;  definition  of  species  of,  175. 
Cephas,  sexual  difterence  of  colour 

in,  537,  552. 
— —    cynosurus      and     griseoviridls, 

colour  of  the  scrotum  in,  537. 
■ Diana,    sexual     differences    of 

colour  in,  537,  552,  553. 
griseo-viridis,  101. 

petaurista,   whiskers,    &c.,    of, 

550. 

Ceres,  of  birds,  bright  colours  of,  491. 

Ceriornis  Temminckii,  swelling  of  the 
wattles  of  the  male  during  court- 
ship, 383. 

Cervulus,  weapons  of,  514. 

moschatas,    rudimentary  horns 

of  the  female,  504. 

Cervus  alces,  234. 

. campestris,  odour  of,  529. 

canadensis,   traces   of  horns    in 

the  female,  504  ;  attacking  a  man, 

511 ;  sexual  difference  in  the  colour 

of,  536. 
. elaphus,  battles  of  male,   501  ; 

horns   of,  with   numerous   points, 

510. 

Eldi,  234. 

mantchuricus,  546. 

paludosus,  colours  of,  536. 

strongyloceros,  234. 

virginianus,   234  ;    horns   of,   in 

course  of  modification,  511. 
28 


Ceryle,  male  black-belted  in  some 
species  of,  457. 

Cetacea,  nakedness  of,  56. 

Ceylon,  frequent  absence  of  beard  in 
the  natives  of,  560. 

Chaffinch,  proportion  of  the  sexes  in 
the,  248  ;  courtship  of  the,  401. 

Chaffinches,  369 ;  new  mates  found 
by,  408. 

Chalcophaps  indicus,  characters  of 
young,  465. 

Chalcosorna  atlas,  sexual  differences 
of,  295. 

Ghamceleo,  sexual  differences  in  the 
genus,  356. 

hifurcus,  356,  357. 

Owenii,  357. 

pumilus,  357. 

Cliamcepetes  unicolor,  modified  wing- 
feather  in  the  male,  377. 

Chameleons,  354. 

Chamois,  danger-signals  of,  100; 
transfer  of  male  characters  to  an 
old  female,  504. 

Champneys,  Mr.,  acromio-basilar 
muscle  and  quadrupedal  gait,  42. 

Chapuis,  Dr.,  on  the  transmission  of 
sexual  peculiarities  in  pigeons,  230; 
on  streaked  Belgian  pigeons,  238, 
446. 

Char,  male,  colouring  of,  during  the 
breeding  season,  340. 

Characters,  male,  developed  in  fe- 
males, 227 ;  secondary  sexual, 
transmitted  through  both  sexes, 
227;  natural,  artificial,  exaggera- 
tion of,  by  man,  582. 

Charadrius  hiaticula  and  pJuvialis, 
sexes  and  young  of,  485. 

Chardin  on  the  Persians,  586. 

Charms,  worn  by  women,  577. 

Charruas,  freedom  of  divorce  among 
the,  598. 

Chasmorhynchws,  difference  of  coloui 
in  the  sexes  of,  389  ;  colours  of,  492. 

• niveus,  389. 

tricarunculatus,  389. 

Chastity,  early  estimation  of,  1 1 9. 

Chatterers,  sexual  differences  in,  219. 

Cheever,  Kev.  H.  T.,  census  of  the 
Sandwich  Islands,  257. 

Cheiroptera,  absence  of  secondary 
sexual  characte]-s  in,  218. 

Chelge  of  Crustacea,  266,  271. 

Chelonia,  sexual  dirfirences  in   350. 


634 


CHENALOPEX. 


INDEX. 


cociiix-ciiijrA, 


Chenalopex  cejyptiacus,  wing-kuobs  of, 
364. 

Cheraprogne,  392,  419. 

Chest,  proportions  of,  in  soldiers  and 
sailors,  32 ;  large,  of  the  Quechua 
and  Aymara  Indians,  34. 

Chevrotains,  canine  teeth  of,  514. 

Ckiasognathus,  stridulation  of,  306. 

Grantii,  mandibles  of,  300. 

Children,  legitimate  and  illegitimate, 
proportion  of  the  sexes  in,  243. 

Chiloe  lice  of  the  natives  of,  170 ; 
population  of,  173. 

C/iimcera  monstrosa,  bony  process  on 
the  head  of  the  male,  338. 

Chimasroid  fishes,  prehensile  organs  of 
male,  331. 

Chimpanzee,  561;  ears  of  the,  14; 
representatives  of  the  eyebrows  in 
the,  19;  hands  of  the,  50;  absence 
of  mastoid  processes  in  the,  53 ; 
platforms  built  by  the,  66 ;  crack- 
ing nuts  with  a  stone,  81 ;  direc- 
tion of  the  hair  on  the  arms  of  the, 
151  ;  supposed  evolution  of  the, 
177  ;  polygamous  and  social  habits 
of  the,  590. 

China,  North,  idea  of  female  beauty 
in,  578. 

,  Southern,  inhabitants  of,  197 

Chinese,  use  of  flint  tools  by  the,  145  ; 
difficulty  'of  distinguishing  the 
races  of  the,  167  ;  colour  of  the 
beard  in,  558 ;  general  beardless- 
ness  of  the,  560 ;  opinions  of  the, 
on  the  appearance  of  Europeans  and 
Cingalese,  578  ;  compression  of  the 
feet  of,  583. 

Chinsurdi,  his  opinion  of  beards,  576, 
581. 

Chlamydera  maculata,  382. 

Chloeon,  pedunculated  eyes  of  the 
male  of,  274. 

C/ilocphcuja,  coloration  of  the  sexes  in, 

460. 
C/ilorocccIus  Tanana  (figured),  285. 

Chorda  Dorsalis,  161. 

Chough,  red  beak  of  the,  491. 

ChromidiB,    frontal    protuberance    in 
male,    340 ;    sexual   difl'ereuces   iu 
colour  of,  345. 
Chrysemys  picta,   long  claws  of  the 

male,  350. 
C/iri/sococcyx,  characters  of  young  of, 
465. 


Chrysomelidae,  stridulation  of,  302. 

Cicada  pruinosa,  282, 

septendecim,  282, 

Cicadas,  songs  of  the,  281  ;  rudimen- 
tary sound-organs  in  females  of,  288. 

Cicatrix  of  a  burn,  causing  modifica- 
tion of  the  facial  bones,  55. 

Cichla,  frontal  protuberance  of  male 
340. 

Cimeti&re  du  Sud,  Paris,  22. 

Cincloramphus  amralis,  large  size  of 
male,  362. 

Cinclus  aqiiatkus,  455. 

Cingalese,  Chinese  opinion  of  the  ap- 
pearance of  the,  578, 

Cirripedes,  complemental  males  of, 
208. 

Civilisation,  effects  of,  upon  natural 
selection,  133;  influence  of,  in  the 
competition  of  nations,  183. 

Clanging  of  geese,  &c.,  368. 

Claparede,  E.,  on  natural  selection 
applied  to  man,  49. 

Clarke,  on  the  marriage-customs  of 
the  Kalmucks,  598. 

Classification,  148. 

Claus,  C,  on  the  sexes  of  Saphirina, 
271. 

Cleft-palate,  inherited,  35. 

Clinutcteris  eryth7'ops,  sexes  of,  479. 

Climate,  31  ;  cool,  favourable  to  hu- 
man progress,  133;  power  of  sup- 
porting extremes  of,  by  man,  182  ; 
want  of  connexion  of,  with  colour, 
192. 

Cloaca,  existence  of  a,  in  the  early 
progenitors  of  man,  161. 

Cloacal  passage  existing  in  the  hu- 
man embryo,  9. 

Clubs,  used  as  weapons  before  disper- 
sion of  mankind,  180. 

Clucking  of  fowls,  368. 

Chithra  A-punctata,  stridulation  of, 
302. 

Coan,  Mr.,  Sandwich-islanders,  187. 

Cobbe,  Miss,  on  morality  in  hypothe 
tical  bee-community,  99. 

Cobra,  ingenuity  of  a,  352. 

Coccus,  147. 

Coccyx,  22,  23  ;  in  the  human  em- 
bryo, 9;  convoluted  body  at  the 
extremity  of  the,  23  ;  imbedded  iu 
the  body,  59. 

Cociiin-China,  notions  of  beauty  of 
the  inhabitants  of,  578,  580. 


INDEX. 


COSMETORNIb. 


635 


Cock,  blind,  fed  by  its  companions, 
103;  game,  killing  a  kite,  303; 
comb  and  wattles  of  the,  403  ;  pre- 
ference shewn  by  the,  for  young 
hens,  420  ;  game,  transparent  zone 
in  the  hackles  of  a,  430. 

Cock  of  the  rock,  405. 

Cockatoos,  491,  492,  493;  nestling, 
411  ;  black,  immature  plumage  of, 
467. 

Ccelenterata,  absence  of  secondary 
sexual  characters  in,  260. 

Coffee,  fondness  of  monkeys  for,  7. 

Cold,  supposed  effects  of,  32  ;  power 
of  supporting,  by  man,  182. 

Coleoptera,  294 ;  stridulation  of,  284  ; 
stridulant  organs  of,  discussed,  303. 

Colias  edusa  and  hyale,  319. 

Collingwood,  C,  ou  the  pugnacity  of 
the  butterflies  of  Borneo,  307  ;  on 
butterflies  being  attracted  by  a 
dead  specimen  of  the  same  species, 
317. 

Colobus,  absence  of  the  thumb,  51. 

Colombia,  flattened  heads  of  savages 
of,  575. 

Colonists,  success  of  the  English  as, 
142. 

Coloration,  protective,  in  birds,  489. 

Colour,  supposed  to  be  dependent  on 
light  and  heat,  32;  correlation  of, 
with  immunity  from  certain  poisons 
and  parasites,  193  ;  purpose  of,  in 
lepidoptera,  316 ;  relation  of,  to 
sexual  functions,  .in  fishes,  343  ; 
difference  of,  in  the  sexes  of  snakes, 
351 ;  sexual  differences  of,  in 
lizards,  357;  influence  of,  in  the 
pairing  of  birds  of  different  species, 
415 ;  relation  of,  to  uidification, 
453,  456 ;  sexual  dilTerences  of,  in 
mammals,  533,  540  ;  recognition  of, 
by  quadrupeds,  540  ;  of  cliildren, 
iu  different  races  of  man,  557  ;  of 
the  skin  in  man,  604. 

Colours,  admired  alike  bv  man  and 
animals,  93  ;  bright,  due  to  sexual 
selection,  261  ;  bright,  among  tlie 
lower  animals,  261,  262;  bright, 
protective  to  butterflies  and  moths, 
313;  bright,  in  male  fishes,  335, 
340  ;  transmission  of,  in  birds,  448. 

Colquhoun,  example  of  reasoning  in 
a  retriever,  78. 

Columba  passcrina,  young  of,  467. 


Colymbus  glacialis,  anomalous  young 

of,  482. 
Comb,  development  of,  in  fowis,  239. 
Combs  and  wattles  in  male  birds,  403. 
Community,  preservation  of  variations 

useful  to  the,  by  natural  selection, 

62. 
Composite,  gz'adation  of  species  among 

the,  175. 
Comte,  C,  on  the  expression  of  the 

ideal  of  beauty  by  sculpture,  581. 
Conditions  of  life,  action  of  changed, 

upon   man,    30 ;    influence    of,    on 

plumage  of  birds,  472. 
Condor,  eyes  and  comb  of  the,  472. 
Conjugations,  origin  of,  91. 
Conscience,  114,  126;  absence  of,  in 

some  criminals,  116. 
Constitution,  difference  of,  in  different 

races  of  men,  168. 
Consumption,  liability  of  Ceb'is  Azarce 

to,    7  ;    connexion    between    com- 
plexion and,  194. 
Convergence  of  characters,  177. 
Cooing  of  pigeons  and  doves,  374. 
Cook,   Capt.,    on    the    nobles    of  the 

Sandwich  Islands,  586. 
Cope,  E.  D.,  on  the  Diuosauria,  158. 
Cophotis  ceylnnica,  sexual  differences 

of,  354,  357. 
Cupris,  295. 

Isidis,     sexual     differences     of, 

296. 

lunaris,  stridulation  of,  303. 

Corals,  bright  colours  of,  260. 
Coral-snakes,  353. 

Cordylus,  sexual  difference  of  colour 
in  a  species  of,  357. 

Corfu,  habits  of  the  Chaffinch  in,  248. 

Cornelius,  on  the  proportions  of  the 
sexes  in  Lucanus  Ccrvus,  253. 

Corpora  Wolffiana,  161 ;  agreement 
of,  with  the  kidneys  of  fishes,  11. 

Correlated  variation,  43. 

Correlation,  influence  of,  in  the  pro- 
duction of  races,  197. 

Corse,  on  the  mode  of  fighting  of  the 
elephant,  514. 

Corvus  corone,  408. 

graculus,  red  beak  of,  491. 

pica,      nuptial      assembly     of, 

406. 
Corydalis  comutus,  large  jaws  of  the 

male,  275. 
Cosmetornis,  462. 


636 


COSMETORNIS. 


INDEX. 


CYANECULA. 


Cosmetomis  vexillarius,  elongation  of 
wiug-feathers  in,  384,  403. 

Cotingidse,  sexual  differences  in,  219; 
coloration  of  the  sexes  of,  460  ;  re- 
semblance o*f  the  females  of  distinct 
species  of,  470. 

Coitus  scorpius,  sexual  differences  in, 
337. 

Coulter,  Dr.,  on  the  Californian  In- 
dians, 258, 

Counting,  origin  of,  144 ;  limited 
power  of,  in  primeval  man,  180. 

Courage,  variability  of,  in  the  same 
species,  69  ;  universal  high  appre- 
ciation of,  118;  importance  of, 
130  ;  characteristic  of  men,  564. 

Courtship,  greater  eagerness  of  males 
in,  221  ;  of  fishes,  331,  341  ;  of 
birds,  367,  405. 

Cow,  winter  change  of  colour,  542. 

Crab,  devil,  269. 

,  shore,  habits  of,  268. 

Crahro  cribrarius,  dilated  tibise  of  the 
male,  276. 

Crabs,  proportions  of  the  sexes  in, 
255. 

Cranz,  on  the  inheritance  of  dexterity 
in  seal  catching,  33. 

Crawfurd,  on  the  number  of  species 
of  man,  174. 

Crenilabrus  massa  and  C.  melops,  nests 
built  by,  345. 

Crest,  origin  of,  in  Polish  fowls,  231. 

Crests,  of  birds,  difference  of,  in  the 
sexes,  467  ;  dorsal  hairy,  of  mam- 
mals, 530. 

Cricket,  field-,  stridulation  of  the, 
283  ;  pugnacity  of  male,  289. 

,     house-,     stridulation    of    the 

283,  284. 

Crickets,  sexual  differences  in,  289. 

Criocerida",  striduhition  of  the,  302. 

Crinoids,  complexity  of,  91. 

Cloaking  of  frogs,  350. 

Crocodiles,  musky  odour  of,  during 
the  breeding  season,  351. 

Crocodilia,  351. 

Crossbills,  characters  of  young,  464. 

Crosses  in  man,  173. 

Crossing  of  races,  etfects  of  the,  192. 

Crossoptilon  auritum,  400,  452,  472  ; 
adornment  of  both  sexes  of,  235  ; 
sexes  alike  in,  460. 

Crotch,  G.  R.,  on  the  stridulation  of 
beetles,   302,  304  ;  on  the  stridu- 


lation of  Heliopathes,  305 ;  on  the 
stridulation  of  Acalles,  306 ;  habit 
of  female  deer  at  breeding  time,  503. 

Crow  Indians,  long  hair  of  the,  580. 

,  young  of  the,  481. 

Crows,  491  ;  vocal  organs  of  the,  370  ; 
living  in  triplets,  409. 

,  carrion,  new  mates  found  bv, 

407. 

,  Indian,  feeding  their  blind  com- 
panions, 103. 

Cruelty  of  savages  to  animals,  118. 

Crustacea,  parasitic,  loss  of  limbs  by 
female,  208 ;  prehensile  feet  and 
antennae  of,  209  ;  male,  more  active 
than  female,  221 ;  parthenogenesis 
in,  255 ;  secondary  sexual  charac- 
ters of,  265 ;  amphipod,  males 
sexually  mature  while  young,  485  ; 
auditory  hairs  of,  568. 

Crystal  worn  in  the  lower  lip  by  some 
Central  African  women,  575. 

Cuckoo  fowls,  238. 

Culicidae,  208,  280  ;  attracted  by  each 
other's  humming,  280. 

Cullen,  Dr.,  on  the  throat-pouch  of 
the  male  bustard,  373. 

Cultivation  of  plants,  probable  origin 
of,  133. 

Cupples,  Mr.,  on  the  numerical  pro- 
portion of  the  sexes  in  dogs,  sheep, 
and  cattle,  246 ;  on  the  Scotch 
deerhound,  516 ;  ou  sexual  pre- 
ference in  dogs,  524. 

Curculionida?,  sexual  difference  in 
length  of  snout  in  some,  208 ; 
hornlike  processes  in  male,  299 ; 
musical,  301,  302. 

Curiosity,  manifestations  of,  by  ani- 
mals, 71. 

Curlews,  double  moult  in,  390. 

Cursores,  comparative  absence  of 
sexual  differences  among  the,  219. 

Curtis,  J.,  on  the  proportion  of  the 
sexes  in  At/ialia,  254. 

Cuvier,  F.,  on  the  recognition  of 
women  by  male  quadrumana,  8. 

,  G.,  on  the  number  of  caudal 

vertebras  in  the  mandrill,  58 ;  on 
instinct  and  intelligence,  67  ;  views 
of,  as  to  the  position  of  man,  149; 
on  the  position  of  the  seals,  150 ; 
on  Jlcctocoty'ey  263. 

Cyanecula  suecicay  sexual  differences 
of,  472. 


CYANALCYON. 


INDEX. 


di::ndropiiila. 


^17 


CyavMlcyon,     sexunl      diftercnce      in 

Dances  of  birds,  380. 

colours    of,    457 ;    immature    plu- 

Dancing, universality  of,  178. 

mage  of,  407. 

Daniell,   Dr.,  his  experience   of  resi- 

Cychrus, sounds  produced  by,  304. 

dence  in  West  Africa,  195. 

Ct/cnia  mcndica,  sexual   ditl'erence  of, 

Darfur,  protuberances  artificially  pro- 

in colour,  316. 

duced  by  natives  of,  574, 

Cygnus  fe>~us,  trachea  of,  374. 

Darwin,    F.,    on    the    stridulation    of 

olor,  white  young  of,  482. 

Dermestes  murinua,  302. 

Cyllo  Leda,  instability  of  the  ocellatcd 

Dasychira  pudibunda,  sexual  difierence 

spots  of,  428. 

of  colour  in,  316. 

Cynantlms,    variation   in   the    genus, 

Davis,  A.  H.,  on  the  pugnacity  of  the 

423. 

male  stag-beetle,  300. 

Cvnipida?,  proportions  of  the  sexes  in, 

,  J.  B.,  on   the    capacity   of  the 

'254. 

skull  in  various  races  of  men,  54  ; 

Cynocephalus,  difference  of  the  young 

on   the   beards  of  the  Polynesians, 

from  the  adult,  8 ;    male,  recogni- 

560. 

tion   of  women  by,  8  ;   polygamous 

Death-rate   higher  in  towns   than  in 

habits  of  species  of,  217. 

rural  districts,  139. 

chacina.  70. 

Death-tick,  306. 

gelada,  81. 

De  Candolle,  Alj^h.,  on  a  case  of  in- 

 hamadryas,  81,  590;  sexual  dif- 

herited power  of  moving  the  scalp, 

ference  of  colour  in,  537. 

13. 

leucophus,  colours  of  the  sexes 

Declensions,  origin  of,  91. 

of,  538. 

Decoration  in  birds,  381. 

mormon,   colours    of  the    male, 

Decticus,  285. 

538,  540,  550. 

Deer,  233  ;  development  of  the  horns 

porcarius,   mane    of    the    male, 

in,  233  ;  spots  of  young,  464,  546  ; 

521. 

horns  of,  503,  506  ;    use   of  horns 

Cypridina,  proportions  of  the  sexes  in. 

of,  510,  518  ;  horns  of  a,  in  course 

255. 

of  modification,  511;    size   of  the 

Cyprinidae,  proportion  of  the  sexes  in 

horns  of,  515  ;  female,  pairing  with 

the,  249. 

one  male,  whilst  others  are  fighting 

,  Indian,  343. 

for    her,  522  ;    male,  attracted  by 

Cyprinodontidae,  sexual  differences  in 

the  voice  of  the  female,  527  ;  male, 

the,  335,  337. 

odour  emitted  by,  529. 

Cyprinus  auratus,  342. 

,  Axis,  sexual   difference    in    the 

Cypris,  relations  of  the  sexes  in,  255. 

colour  of  the,  537. 

Cyrtodactylus  rubidus,  354. 

,  fallow,  diiiereut  coloured  herds 

Cystophora  cristata,  hood  of,  528. 

of,  540. 

,  Jklantchurian,  546. 

D. 

,  Virginian,  546 ;  colour  of  the, 

not    affected     by    castration,    535 ; 

Daceh,  sexual  difference  of  colour  in. 

colours  of,  536. 

457. 

Deerhound,  Scotch,  greater  size  of  the 

Gaudichaudi,    young    male    of, 

male,  237,  516. 

467. 

Defensive  organs  of  mammals,  518. 

Dal-ripa,  a  kind  of  ptarmigan,  248. 

De  Geer,  C,  on  a  female   spider  de- 

Damalis albifrons,  peculiar  markings 

stroying  a  male,  273. 

of,  544. 

Dekay,  Dr.,  on  the  bladder-nose  seal, 

pygo-fgfi,  peculiar  markings  of. 

528. 

543. 

Delorenzi,  G.,  division  of  malar  bone. 

Dampness  of  climate,  supposed  influ- 

39. 

ence  of,  on  the  colour  of  the  skin. 

Demerara,  yellow  fever  in,  194. 

32,  103. 

Dendrocygna,  465. 

Panaidae,  308. 

DendrophUa  frontalis,  young  of,  487. 

6sS 


IXDEX. 


DOGS. 


Denison,  Sir  W.,  manner  of  ridding 
themselves  of  vermin  among  the 
Australians,  57  ;  extinction  of  Tas- 
manians,  184-. 

Denny,  H.,  on  the  lice  of  domestic 
animals,  169. 

Dermestes  murinus,  stridulation  of, 
302. 

Descent  traced  through  the  mother 
alone,  588. 

Deserts,  pi-otective  colouring  of  ani- 
mals inhabiting,  489. 

Desmnrest,  on  the  absence  of  sub- 
orbital pits  in  Antilope  subguttu- 
rosa,  529 ;  on  the  whiskers  of 
Macacus,  531  ;  on  the  colour  of 
the  opossum,  534- ;  on  the  colours 
of  the  sexes  of  Mxis  minutus,  534  ; 
on  the  colouring  of  the  ocelot,  534  ; 
on  the  colours  of  seals,  535  ;  on 
Antilope  caama,  536 ;  on  the 
colours  of  goats,  536 ;  on  sexual 
difference  of  colour  in  Ateles  mar- 
gtnatus,  537  ;  on  the  mandrill, 
539  ;  on  Macacus  cynomolgus,  558. 

Desmoulins,  on  the  number  of  species 
of  man,  174  ;  on  the  musk-deer, 
530. 

Desor,  on  the  imitation  of  man  by 
monkeys,  72. 

Despine,  P.,  on  criminals  destitute  of 
conscience,  116. 

Development,  embryonic,  of  man,  9, 
11  ;  correlated,  426. 

Devil,  not  believed  in  by  the  Fue- 
gians,  95. 

Devil-crab,  269. 

Devonian,  fossil  insect  from  the,  289. 

Dewlaps,  of  cattle  and  antelopes,  531. 

Dladema,  sexual  differences  of  colour- 
ing in  the  species  of,  309. 

Diamond-beetles,  bright  colours  of, 
294. 

Diastema,  occurrence  of,  in  man,  35. 

Diastvlidaj,  proportion  of  the  sexes  in, 
255. 

Dicrurus,  racket-shaped  feathers  in, 
384  ;  nidification  of,  453. 

macrocercus,  change  of  plumage 

in,  461. 

Didelphis  opossum,  sexual  difference 
in  the  colour  of,  533. 

Differences,  comparative,  between  dif- 
ferent species  of  birds  of  the  same 
sex,  470. 


Digits,  supernumerary,  more  frequent 
in  men  than  in  women,  223  ;  super- 
numerary, inheritance  of,  232 ; 
supernumerary,  earlv  devel  pment 
of,  237. 
Dimorphism  in  females  of  water- 
beetles,  276  ;  in  JS'eurothemis  and 
Agrion,  291. 
Diodorus,  on  the  absence  of  beard  ir» 

the  natives  of  Ceylon,  560. 
Dipelicus    Cantori,   sexnal  differences 

of,  296. 
Diplopoda,    prehensile    limbs    of    the 

male,  274. 
Dipsas   cynodon,  sexual    difference   in 

the  colour  of,  351. 
Diptera,  280. 
Disease,  generated  by  the  contact  of 

distinct  peoples,  183. 
Diseases    common    to    man    and    the 
lower    animals,    7 ;     difference    of 
liability    to,   in   different  races    of 
men,    167 ;    new,   effects   of,  upon 
savages,     182;     sexually    limited, 
237. 
Display,     coloration    of    Lepidoptera 
for,    314 ;    of    plumage    by    male 
birds,  394,  402. 
Distribution,  wide,  of  man,  48  ;    geo- 
graphical,   as  evidence    of  specific 
distinctness  in  man,  169. 
Disuse,  effects  of,  in  producing  rudi- 
mentary organs,   12;    and    use    of 
parts,  effects  of,  32  ;  of  parts,  in- 
fluence of,  on  the  races  of  men,  197. 
Divorce,  freedom  of,  among  the  Char- 

ruas,  598. 
Dixon,  E.  S.,  on  the  pairing  of  different 
species  of  geese,  415  ;  on  the  court- 
ship of  peafowl,  419. 
Dobrizhoffer,  on  the  marriage-customs 

of  the  Abipones,  599. 
Dobson,  Dr.,  on  the  Cheiroptera,  218  ; 
scent-glands  of  bats,  529 ;  fru- 
givorous  bats,  534. 
Dogs,  suffering  from  Tertian  ague,  8  ; 
memory  of,  74 ;  dreaming,  74 ; 
diverging  when  drawing  sledges 
over  thin  ice,  75;  exercise  of  reason- 
ing faculties  by,  78  ;  domestic,  pro- 
gress of,  in  moral  qualities,  80; 
distinct  tones  uttered  by,  81; 
parallelism  between  his  affection  for 
his  master  and  religious  feeling,  96  ; 
sociability  of  the,  100;  sympathy  of, 


DOLlCHOCErHALlC. 


INDEX. 


639 


with  a  sick  cat,  103  ;  sympathy  of, 
with  his  master,  103 ;  their  posses- 
sion of  conscience,  103  ;  possible  use 
of  the  hair  on  the  fore-legs  of  the, 
151 ;  races  of  the,  176;  numerical 
proportion  of  male  and  female  births 
in,  246 ;  sexual  affection  between  in- 
dividuals of,  524;  howling  at  certain 
notes,  569  ;  rolling  in  carrion,  530. 

Dolichocephalic  structure,  possible 
cause  of,  56. 

Dolphins,  nakedness  of,  56. 

Domestic  animals,  races  of,  176  ; 
change  of  breeds  of,  596. 

Domestication,  influence  of,  in  re- 
moving the  sterility  of  hybrids,  172 

D'Orbigny,  A.,  on  the  influence  of 
dampness  and  dryness  on  the  colour 
of  the  skin,  193  ;  on  the  Yuracaras, 
582. 

Dotterel,  477. 

Doubleday,  E.,  on  sexual  differences 
in  the  wings  of  butterflies,  277. 

,   H.  on   the  proportion    of   the 

sexes  in  the  smaller  moths,  251 ; 
males  of  Lasiocampa  quercus  and 
on  the  attraction  of  the  Sa- 
turnia  carpini  by  the  female  252 ; 
on  the  proportion  of  the  sexes  in 
the  Lepidoptera,  252  ;  on  the  tick- 
ing of  Anobium  tesselatum,  306 ; 
on  the  structure  of  Ageronia 
feronia,  307 ;  on  white  butterflies 
alighting  upon  paper,  317. 

Douglas,  J.  W.,  on  the  sexual  differ- 
ences of  the  Hemipjtera,  281 ;  on  the 
colours  of  British  Homoptera,  282. 

Down,  of  birds,  390. 

Draco,  gular  appendages  of,  355. 

Dragonet,  Gemmeoiis,  336. 

Dragon-flies,  caudal  appendages  of 
male,  276  ;  relative  size  of  the 
sexes  of,  279 ;  difference  in  the 
sexes  of,  290  ;  want  of  pugnacity 
by  the  male,  291. 

Drake,  breeding  plumage  of  the,  393. 

Dreams,  74 ;  a  possible  source  of  the 
belief  in  spiritual  agencies,  94. 

Drill,  sexual  difference  of  colour  in 
the,  538. 

DromcBUS,  irroratus,  478. 

Dromolaea,  Saharan  species  of,  456. 

Drongo  shrike,  461. 

Drongos,  racket-shaped  feathers  in 
the  tails  of,  384,  392. 


Dryness,  of  climate,  supposed  influence 
of,  on  the  colour  of  the  skin,  193. 

Drijopitheciis,  155. 

Duck,  harlequin,  age  of  mature  plu- 
mage in  the,  483 ;  breeding  in  im- 
mature plumage  484. 

,  long-tailed,  preference  of  male, 

for  certain  females,  420. 

■ ,  pinta*,  pairing  with  a  wigcon, 

414. 

,  voice  of  the,  374  ;  pairing  with 

a  shield-drake,  414 ;  immature 
plumage  of  the,  466. 

,  wild,  sexual  difierences  in  the 

219  ;  speculum  and  male  charac- 
ters of,  236  ;  pairing  with  a  pintail 
drake,  415. 

Ducks,  wild,  becoming  polygamous 
under  partial  domestication,  219  ; 
dogs  and  cats  recognised  by,  412. 

Dufosse,  Dr.,  sounds  produced  by  fish, 
347. 

Dugong,  nakedness  of,  56 ;  tusks  of, 
502. 

Dujardin,  on  the  relative  size  of  the 
cerebral  ganglia  in  insects,  54. 

Duncan,  Dr.,  on  the  fertility  of  early 
marriages,  138  ;  comparative  health 
of  married  and  single,  140. 

Dupont,  M.,  on  the  occurrence  of  the 
supra-condyloid  foramen  in  the 
humerus  of  man,  22. 

Durand,  J.  P.,  on  causes  of  variation, 
30. 

Dureau  de  la  Malle,  on  the  songs  of 
birds,  86  ;  on  the  acquisition  of  an 
air  by  blackbirds,  370. 

Dutch,  retention  of  their  colour  by 
the,  in  South  Africa,  193. 

Duty,  sense  of,  97. 

Duvaucel,  female  Hylobatcs  washing 
her  young,  70, 

Dyaks,  pride  of,  in  mere  homicide,  1 1 7. 

Dynastes,  large  size  of  males  of,  279. 

Dynastini,  stridulation  of,  303. 

Dytiscus,  dimorphism  of  females  of, 
276  ;  grooved  elvtra  of  the  female, 
276. 


E. 

Eagle,  young  Cercopithecus  rescued 
from,  by  the  troop,  101. 

,  white-headed,  breeding  in  im- 
mature plumage,  484. 


640 


INDEX. 


KNGLEHEAR'] 


Eagles,  golden,  new  mates  found  by, 
408. 

Ear,  motion  of  the,  13 ;  external 
shell  of  the,  useless  in  man,  14 ; 
rudimentary  point  ofthe,  in  man,  15. 

Ears,  more  variable  in  men  than 
women,  224 ;  piercing  and  orna- 
mentation of  the,  575. 

Earwigs,  parental  feeling  4b,  106. 

Echidna,  156. 

Echini,  bright  colours  of  some,  260. 

Echinodermata,  absence  of  secondary 
sexual  characters  in,  260. 

Echis  carinata,  353. 

Ecker,  figure  of  the  human  embryo, 
10 ;  on  the  development  of  the 
gyri  and  sulci  of  the  brain,  204 ; 
on  the  sexual  differences  in  the 
pelvis  in  man,  557  ;  on  the  pre- 
sence of  a  sagittal  crest  in  Austra- 
lians, 558. 

Edentata,  former  wide  range  of,  in 
America,  169  ;  absence  of  secondary 
sexual  characters  in,  218. 

Edolius,  racket-shaped  feathers  in, 
384. 

Edwards.  Mr.,  on  the  proportion  of 
the  sexes  in  North  American  species 
o(Fapilio,  250. 

Eels,  hermaphroditism  of,  162. 

Egerton,  Sir  P.,  on  the  use  of  the 
antlers  of  deer,  510  ;  on  the  pairing 
of  red  deer,  522  ;  on  the  bellowing 
of  stags,  526. 

Eggs,  hatched  by  male  fishes,  345. 

Egret,  Indian,  sexes  and  young  of,  486. 

Egrets,  breeding  plumage  of,  391  ; 
white,  492. 

Ehrenberg,  on  the  mane  of  the  male 
Hamadryas  baboon,  521. 

Ekstrom,  M.,  on  Harelda  glacialis, 
420. 

Elachista  rufocinerea,  habits  of  male, 
252. 

Eland,  development  of  the  horns  of 
the,  234. 

Elands,  sexual  differences  of  colour  in 
535 

Elaphomijia,  sexual  differences  in,  280. 

Elaphrus  ulijinosus,  stridulation  of, 
302. 

Elaps,  353. 

Elatcridae,  proportions  of  the  sexes  in, 
253. 

Elaters,  luminous,  278. 


Elephant,  156 ;  rate  of  increase  of 
the,  47  ;  nakedness  of  the,  57  ;  In- 
dian, forbearance  to  his  keeper, 
104 ;  polygamous  habits  of  the, 
218  ;  pugnacity  of  the  male,  501 ; 
tusks  of,  502,  503,  507,  515  ; 
Indian,  mode  of  fighting  of  the, 
513 ;  male,  odour  emitted  by  the, 
529 ;  attacking  white  or  grey 
horses,  540. 

Elevation  of  abode,  modifying  in- 
fluence of,  35. 

Elimination  of  inferior  individuals, 
137. 

Elk,  507;  winter  change  of  the,  542. 

,  Irish,  horns  ofthe,  515. 

El  lice  Islands,  beards  of  the  natives, 
560,  581. 

Elliot,  R.,  on  the  numerical  pro- 
portion of  the  sexes  in  young  rats, 
247  ;  on  the  proportion  of  the 
sexes  in  sheep,  246. 

,  D.  G.,    on    Felecanus   erythro- 

rhynchus,  390. 

,    Sir    W.,    on    the    polygamous 

habits  of  the  Indian  wild  boar, 
218. 

Ellis,  on  the  prevalence  of  infanticide 
in  Polynesia,  592. 

Elphinstone,  Mr.,  on  local  differences 
of  stature  among  the  Hindoos,  31 ; 
on  the  difficulty  of  distinguishing 
the  native  races  of  India,  167. 

Elytra,  of  the  females  of  Dytiscus^ 
Acilius,  Hydroporus,  276. 

Emberiza,  characters  of  young,  464. 

miliaria,  464. 

schccniclus,  41 2  ;    head-feathers 

of  the  male,  402. 

Embryo  of  man,  9,  10  ;  ofthe  dog,  10. 

Embryos  of  mammals,  resemblance  of 
the,  25. 

Emigration,  137. 

Emotions  experienced  by  the  lower 
animals  in  common  with  man,  69 ; 
manifested  by  animals,  71. 

Emperor  moth,  315. 

Emulation  of  singing-birds,  369. 

Emu,  sexes  and  incubation  of,  478. 

Endurance,  estimation  of,  118. 

Energy,  a  characteristic  of  men,  565. 

England,  numerical  propcrtion  of 
male  and  female  births  in,  242. 

Engleheart,  Mr.,  on  the  finding  of 
new  mates  by  starlings,  408. 


INDEX. 


FACULTIES. 


641 


English,  success  of,  as  colonists,  142. 

Engravers,  short-sighted,  33. 

Entomostraca,  268. 

Entozoa,  ditference  of  colour  between 
the  males  and  females  of  some,  260. 

Envy,  persistence  of,  112. 

Eocene  period,  possible  divergence  of 
man  during  the,  156. 

Eolidae,  colours  of,  produced  by  the 
biliary  glands,  261. 

Epeira  nigra,  small  size  of  the  male 
of,  273 

Ephemeras,  274. 

Ephemerida;,  290. 

Ephippiger  vitiuin,  stridulatmg  organs 
of,  284,  288. 

Epicalia,  sexual  differences  of  colour- 
ing in  the  species  of,  309. 

Equus  hemionus,  winter  change  of, 
542. 

Erateina,  coloration  of,  315. 

Ercolani,  Prof,  hermaphroditism  in 
eels,  162. 

Erect  attitude  of  man,  51,  52. 

Eristalis,  courting  of,  280. 

Eschricht,  on  the  development  of  hair 
in  man,  18;  on  a  lanuginous 
moustache  in  a  female  foetus,  19; 
on  the  want  of  definition  between 
the  scalp  and  the  forehead  in  some 
children,  151  ;  on  the  arrangement 
of  the  hair  in  the  human  foetus,  152; 
on  the  hairmess  of  the  face  in  the 
human  foetus  of  both  sexes,  602,  603. 

Esmeralda,  difference  of  colour  m  the 
sexes  of,  294. 

Esox  lucius,  249. 

reticulatus,  340. 

Esquimaux,  64,  133  ;  their  belief  in 
the  inheritance  of  dexterity  in 
seal-catching,  33  ;  mode  of  life  of, 
197. 

Estrelda  amandava,  pugnacity  of  the 
male,  366. 

Eubagis,  sexual  differences  of  colour- 
ing in  the  species  of,  309. 

Euchirus  longimanus,  sound  produced 
by,  304. 

Eudromias  morinellus,  477. 

Eulamjns  jugularis,  colours  of  the 
female,  454. 

Euler,  on  the  rate  of  increase  in  the 
United  States,  44. 

Eumomota  superciliaris,  racket-shaped 
feathers  in  the  tail  of,  384. 


Eup)ctomena  macroura,  colours  of  the 
female,  453. 

Euphema  splendida,  457. 

Euplocamus  erytkrophthahnus,  posses- 
sion of  spurs  by  the  female,  364. 

Europe,  ancient  inhabitants  of,  181. 

Europeans,  difference  of,  from  Hin- 
doos, 192 ;  hairiness  of,  probably 
due  to  reversion,  601. 

Eurostopodus,  sexes  of,  479. 

Eurygnathus,  different  proportions  of 
the  head  in  the  sexes  of,  276. 

Eustephanus,  sexual  differences  of  spe- 
cies of,  359  ;  young  of,  487. 

Exaggeration  of  natural  characters  by 
man,  582. 

Exogamy,  588,  591. 

Expression,  resemblances  in,  between 
man  and  the  apes,  150. 

Extinction  of  races,  causes  of,  181. 

Eye,  destruction  of  the,  32  ;  change 
of  position  in,  55 ;  obliquity  of, 
regarded  as  a  beauty  by  the  Chinese 
and  Japanese,  578. 

Eyebrows,  elevation  of,  13  ;  develop- 
ment of  long  hairs  in,  19 ;  in  mon- 
keys, 151  ;  eradicated  in  parts  of 
South  America  and  Africa,  575  ; 
eradication  of,  by  the  Indians  of 
Paraguay,  580. 

Eyelashes,  eradication  of,  by  the  In- 
dians of  Paraguay,  580. 

Eyelids,  coloured  black,  in  part  of 
Africa,  574. 

Eyes,  pillared,  of  the  male  of  Chloeon, 
274 ;  diflerence  in  the  colour  of, 
in  the  sexes  of  birds,  425. 

Eyton,  T.  C,  observations  on  the  de- 
velopment of  the  horns  in  the 
fallow-deer,  234. 

Eyzies,  Les,  human  remains  from, 
181. 


F. 

Fabre,  M.,  on  the  habits  of  Cerceris, 

291. 
Facial  bones,  causes  of   modification 

of  the,  55. 
Faculties,   diversity  of,   in   the  same 

race  of  men,  26  ;  inheritance  of,  27 ; 

diversity  of,  in  animals  of  the  same 

species,  28;    mental,   variation  of, 

in  the  same   species,  66  ;  of  birds, 

410. 


642 


INDEX. 


Fakirs,  Indian,  tortui'es  undergone  by, 
118. 

Falco  leucocephalus,  484. 

peregrinus,  408,  461. 

tinnunculus,  408. 

Falcon,  peregrine,  new  mate  found 
by,  408. 

Falconer,  H.,  on  the  mode  of  Kghting 
of  the  Indian  elephant,  513 ;  on 
canines  in  a  female  deer,  514;  on 
H'^omoschus  aquaticics,  547. 

Falkland  Islands,  horses  of,  181. 

Fallow-deer,  different  coloured  herds 
of,  540. 

Famines,  frequency  of,  among  savages, 
46. 

Farr,  Dr.,  on  the  structure  of  the 
uterus,  38  ;  on  the  effects  of  pro- 
fligacy, 137 ;  on  the  influence  of 
marriage  on  mortality,  139,  140. 

Farrar,  F.  W.,  on  the  origin  of  lan- 
guage, 87  ;  on  the  crossing  or  blend- 
ing of  languages,  91  ;  on  the 
absence  of  the  idea  of  God  in  cer- 
tain races  of  men,  93 ;  on  early 
marriages  of  the  poor,  138  ;  on  the 
middle  ages,  141. 

Fashions,  long  prevalence  of,  among 
savages,  576,  584. 

Faye,  Prof,  on  the  numerical  propor- 
tion of  male  and  female  births  in 
Norway  and  Russia,  243  ;  on  the 
greater  mortality  of  male  children 
at  and  before  birth,  243. 

Feathers,  modified,  producing  sounds, 
377  et  seq.,  450 ;  elongated,  in 
male  birds,  383,  403  ;  racket- 
shaped,  384 ;  barbless  and  with 
filamentous  barbs  in  certain  birds, 
385  ;  shedding  of  margins  of,  393. 

Feeding,  high,  probable  influence  of, 
in  the  pairing  of  birds  of  dilferent 
species,  415. 

Feet,  thickening  of  the  skin  on  the 
soles  of  the,  33 ;  modification  of, 
in  man,  52. 

Felis  canadensis,  throat-ruff  of,  521. 

pardalis   and    F.    mitis,    sexual 

differences  in  the  colouring  of,  534. 

Female,  behaviour  of  the,  during 
courtship,  222. 

birds,  differences  of,  470. 

Femalej,  presence  of  rudimentary 
male  organs  in,  162 ;  preference 
of,  for  certain  males,  214  ;  pursuit 


of,  by  males,  221  ;  occurrence  of 
secondary  sexual  characters  in, 
225  ;  development  of  male  charac- 
ters by,  227. 

Females  and  males,  comparative  num- 
bers of,  213,  215;  comparative  mor- 
tality of,  while  young,  216. 

Femur  and  tibia,  proportions  of,  in  the 
Aymara  Indians,  34. 

Fenton,  Mr.,  decrease  of  Maories, 
184  ;  infanticide  amongst  the 
Maories,  256. 

Ferguson,  Mr.,  on  the  courtship  of 
fowls,  417. 

Fertility  lessened  under  changed  con- 
ditions, 188. 

Fertilisation,  phenomena  of,  in  plants, 
222  ;  in  the  lower  animals,  222. 

Fevers,  immunity  of  Negroes  and 
Mulattoes  from,  193. 

Fiber  zihethicus,  protective  colouring 
of  it,  542. 

Fick,  H.,  effect  of  conscription  for 
military  service,  134. 

Fidelity  of  savages  to  one  another, 
118;  importance  of,  124. 

Field-slaves,  difference  of,  from  house- 
slaves,  196. 

Fijians,  burying  their  old  and  sick 
parents  alive,  102  ;  estimation  of 
the  beard  among  the,  581 ;  admi- 
ration of,  for  a  broad  occiput, 
683. 

Fiji  Archipelago,  population  of  the, 
173. 

Islands,  beards  of  the   natives, 

560,  581 ;  mariiage-customs  of  the, 
598. 

Filial  affection,  partly  the  result  of 
natural  selection,  105. 

Filum  termiuale,  23. 

Finch,  racket-shaped  feathers  in  the 
tail  of  a,  384. 

Finches,  spring  change  of  colour  in. 
393  ;  British,  females  of  the,  46o. 

Fingers,  partially  coherent,  in  species 
of  Jlylobates,  51. 

Finlayson,  on  the  Cochin  Chinese, 
578. 

Fire,  use  of,  49,  145,  180. 

Fischer,  on  the  pugnacity  of  the  m;!k' 
of  Lethrus  ccphalotes,  300. 

Fish,  eagerness  of  male,  221;  propor- 
tion of  the  sexes  in,  249;  sounds 
produced  by,  347. 


INDEX. 


FRINGILLID.E. 


643 


Fishes,  kidneys  of,  represented  by- 
Corpora  Wolffiana  in  the  human 
embryo,  11;  male,  hatching  ova 
in  their  mouths,  163  ;  receptacles 
for  ova  possessed  by,  208  ;  relative 
size  of  the  sexes  in,  335  ;  fresh- 
water, of  the  tropics,  343  ;  protec- 
tive resemblances  in,  344 ;  change 
of  colour  in,  344 ;  nest-building, 
346 ;  spawning  of,  345 ;  sounds 
produced  by,  347,  5GG  ;  continued 
growth  of,  485. 

Flamingo,  age  of  mature  plumage, 
483. 

Flexor  poll  ids  longus,  similar  variation 
of,  in  man,  42. 

Flint  tools,  145. 

Flints,  ditficulty  of  chij^ping  into 
form,  49. 

Floresuga  melUvora,  443. 

Florida,  Quiscalus  major  in,  248. 

Flounder,  coloration  of  the,  344. 

Flower,  W.  H.,  on  the  abductor  of 
the  fifth  metatarsal  in  apes,  42 ; 
on  the  position  of  the  Seals,  150  ; 
on  the  Pithecia  monachus,  201  ; 
on  the  throat-pouch  of  the  male 
bustard,  373. 

Fly-catchers,  colours  and  nidification 
of,  455. 

Foetus,  human,  woolly  covering  of 
the,  19;  arrangement  of  the  hair 
on,  152. 

Food,  influence  of,  upon  stature,  31. 

Foot,  prehensile  power  of  the,  re- 
tained in  some  savages,  52 ;  pre- 
hensile, in  the  early  progenitors  of 
man,  160. 

Foramen,  supra  -  condyloid,  excep- 
tional occurrence  of  in  the  humerus 
of  man,  21,  43  ;  in  the  early  pro- 
genitors of  man,  160. 

Forbes,  D.,  on  the  Aymara  Indians, 
34 ;  on  local  variation  of  colour 
in  the  Quichuas,  196  ;  on  the  hair- 
lessness  of  the  Aymaras  and  Qui- 
chuas, 561  ;  on  the  long  hair  of 
the  Aymaras  and  Quichuas,  559, 
580. 

Forel,  F.,  on  white  young  swans, 
282. 

Formica  riifa,  size  of  the  cerebral 
ganglia  in,  64. 

Fossils,  absence  of,  connecting  man 
with  the  apes,  166. 


Fowl,  occurrence  of  spurs  in  the 
female,  227  ;  game,  early  pugnacity 
of,  239  ;  Polish,  early  development 
of  cranial  peculiarities  of,  239 ; 
variations  in  plumage  of,  385  ;  ex- 
amples of  correlated  development 
in  the,  426  ;  domestic,  breeds  and 
subbreeds  of,  460. 

Fowls,  spangled  Hamburgh,  229,  238  ; 
inheritance  of  changes  of  plumage 
by,  229 ;  sexual  peculiarities  in, 
transmitted  only  to  the  same  sex, 
230 ;  loss  of  secondary  sexual  cha- 
racters by  male,  231 ;  Polish,  origin 
of  the  crest  in,  231  ;  period  of  in- 
heritance of  characters  by,  238  ; 
cuckoo-,  238  ;  development  of  the 
comb  in,  239 ;  numerical  propor- 
tion of  the  sexes  in,  247  ;  courtship 
of,  417  ;  mongrel,  between  a  black 
Spanish  cock  and  different  hens, 
427;  pencilled  Hamburgh,  differ- 
ence of  the  sexes  in,  447  ;  Spanish, 
sexual  differences  of  the  comb  in, 
447  ;  spurred,  in  both  sexes,  449. 

Fox,  W.  D.,  on  some  half-tamed  wild 
ducks  becoming  polygamous,  and 
on  polygamy  in  the  guinea-fowl  and 
canary-bird,  220  ;  on  the  proportion 
of  the  sexes  in  cattle,  247 ;  on  the 
pugnacity  of  the  peacock,  364 ;  on 
a  nuptial  assembly  of  magj^ies, 
406  ;  on  the  finding  of  new  mates 
by  crows,  407  ;  on  partridges  living 
in  triplets,  409  ;  on  the  pairing  of  a 
goose  with  a  Chinese  gander,  415. 

Foxes,  wariness  of  young,  in  hunting 
districts,  80  ;  black,  540. 

Fraser,  C,  on  the  different  colours  of 
the  sexes  in  a  species  of  Squilla, 
271. 

,  G.,  colours  of  Thecia,  312. 

Frere,  Hookham,  quoting  Theognis  on 
selection  in  mankind,  29. 

Fringilla  cannabina,  394. 

ciris,  age  of  mature  plumage  in, 

483. 

cyanea,  age  of  mature  plumage 

in,  483. 

leucophrys,  young  of,  486. 

spinus,  416. 

.  tristis,  change  of  colour   in,  in 

spring,  393  ;  young  of,  485. 

Fringillida,  rssemblance  of  the  fe- 
males of  distinct  species  of,  470. 


644 


FROGS. 


INDEX. 


GELASIMUS. 


Frogs,  349 ;  male,  temporary  recep- 
tacles for  ova  possessed  by,  208 ; 
ready  to  breed  before  the  females, 
212;  fighting  of,  349  ;  vocal  organs 
of,  350. 

Frontal  bone,  persistence  of  the  suture 
in,  39. 

Fruits,  poisonous,  avoided  by  animals, 
66. 

Fuegians,  133,  143  ;  difference  of  sta- 
ture among  the,  31 ;  power  of  sight 
in  the,  33  ;  skill  of,  in  stone-throw- 
ing, 49 ;  resistance  of  the,  to  their 
severe  climate,  63,  182;  mental 
capacity  of  the,  65  ;  quasi-religious 
sentiments  of  the,  95  ;  resemblance 
of,  in  mental  characters,  to  Euro- 
peans, 178 ;  mode  of  life  of  the, 
197 ;  aversion  of,  to  hair  on  the 
face,  580  ;  said  to  admire  European 
women,  582. 

Fulgoridae,  songs  of  the,  281. 

Fur,  whiteness  of,  in  arctic  animals, 
in  winter,  229. 

Fur-bearing  animals,  acquired  saga- 
city of,  80. 


G. 


the 


Gallicrex,    sexual    difference    in 
colour  of  the  irides  in,  425. 

cristatus,    pugnacity    of    male, 

360 ;  red  caruncle  occurring  in 
the  male  during  the  breeding- 
season,  389. 

Gallinaceaj,  frequency  of  polygamous 
habits  and  of  sexual  ditierences  in 
the,  219;  love-gestures  of,  380; 
decomposed  feathers  in,  385  ;  stripes 
of  young,  464  ;  comparative  sexual 
differences  between  the  species  of, 
470,  471  ;  plumage  of,  472. 

Gallinaceous  birds,  weapons  of  the 
male,  362  ;  racket-shaped  feathers 
on  the  heads  of,  384. 

Gallinula  chloropus,  pugnacity  of  the 
male,  360. 

cristata,  pugnacity  of  the  male, 

360. 

GalloperdiXy  spurs  of,  364  ;  develop- 
ment of  spurs  in  the  female,  450. 

Gallophasis,  young  of,  468. 

Gallns  hankiva,  447  ;  neck-hackles  of, 
392. 

Stanleyi,  pugnacity  of  the  male, 

.  363. 


Galls,  60. 

Galton,  Mr,,  on  hereditary  genius,  28  ; 
gregariousness  and  independence 
in  animals,  104 ;  on  the  struggle 
between  the  social  and  personal 
impulses,  125 ;  on  the  effects  of 
natural  selection  on  civilised  na- 
tions, 133  ;  on  the  sterility  of  sole 
daughters,  135 ;  on  the  degree  of 
fertility  of  people  of  genius,  136  ; 
on  the  early  marriages  of  the  poor, 
138  ;  on  the  ancient  Greeks,  140  ; 
on  the  Middle  Ages,  141  ;  on  the 
progress  of  the  United  States,  142  ; 
on  South  African  notions  of  beautv, 
579. 

Gammarus,  use  of  the  chelae  of,  268. 

marinus,  270. 

Gannets,  white  only  when  mature, 
492. 

Ganoid  fishes,  159,  165. 

Gaour,  horns  of  the,  505. 

Gap  between  man  and  the  apes,  156. 

Gaper,  sexes  and  young  of,  486. 

Gardner,  on  an  example  of  rationality 
in  a  Gelasimus,  270. 

Garrulus  glandarius,  407. 

Gartner,  on  sterility  of  hybrid  plants, 
172. 

Gasteropoda,  272 ;  pulmoniferous, 
courtship  of,  262. 

Gasterosteus,  220 ;  nidification  of, 
345. 

leiurus,  331,  340,  345. 

trac/iurus,  332. 

Gastrojjhora,  wings  of,  brightly  co- 
loured beneath,  315. 

Gauchos,  want  of  humanitv  among 
the,  123. 

Gaudry,  M,,  on  a  fossil  monkey,  154. 

Gavia,  seasonal  change  of  plumage  in, 
493. 

Geese,  clanging  noise  made  by,  368 
pairing  of  different  species  of,  415; 
Canada,  selection  of  mates  by,  416. 

Gegenbaur,  C,  on  the  number  of 
digits  in  the  Ichthyopterygia,  37  ; 
on  the  hermaphroditism  of  the 
remote  progenitors  of  the  verte- 
brata,  161  ;  two  types  of  nipple  in 
mammals,  162. 
Gelasirnus,  proportions  of  the  sexes 
in  a  species  of,  254  ;  use  of  the 
enlarged  chelas  of  the  male,  268  ; 
pugnacity  of  males  of,  269;  rational 


GE1IMULE3. 


INDEX. 


GONKI'TKRVX. 


645 


actions  of  a,  270;  difference  of 
colom*  in  the  sexes  of  a  species  of, 
271. 

Gemmules,  dormant  in  one  sex,  231. 

Genius,  28  ;  hereditary,  b64. 

-,  fertility  of  men  and  women  of, 

136. 

Geoffroy-Saint-Hilaire,  Isid.,  on  the 
recognition  of  women  by  male 
quadrumana,  8 ;  on  monstrosities, 
30  ;  coincidences  of  arrested  devel- 
opment with  polydactylism,  37  ; 
on  animal-like  anomalies  in  the 
human  structure,  40  ;  on  the  cor- 
relation of  monstrosities,  44;  on 
the  distribution  of  hair  in  man  and 
monkeys,  57  ;  on  the  caudal  ver- 
tebrae of  monkeys,  58 ;  on  cor- 
related variability,  60 ;  on  the 
classification  of  man,  147  ;  on  the 
long  hair  on  the  heads  of  species  of 
Semnopithecus,  151 ;  on  the  hair  in 
monkeys,  152 ;  on  the  develop- 
ment of  horns  in  female  deer,  504 ; 
and  F.  Cuvier,  on  the  mandrill, 
539  ;  on  Hylobates,  558,  559. 

Geographical  distribution,  as  evidence 
of  specific  distinctions  in  man,  169. 

Geometrae,  brightly  coloured  beneath, 
315. 

Geophagus,  frontal  protuberance  of 
male,  340,  345 ;  eggs  hatched  by 
the  male,  in  the  mouth  or  bran- 
chial cavity,  345. 

Georgia,  change  of  colour  in  Germans 
settled  in,  196. 

Geotrupes,  stridulation  of,  303,  304. 

Gerbe,  M.,  on  the  nest-building  of 
Crenilabrus  massa  and  C.  mclops, 
345. 

Gerland,  Dr.,  on  the  prevalence  of 
infanticide,  117,  577,  592  ;  on  the 
extinction  of  races,  182. 

Gervais,  P.,  on  the  hairiness  of  the 
gorilla,  57  ;  on  the  mandrill,  538. 

Gesture-languag£,  178. 

Ghost-moth,  sexual  difference  of 
colour  in  the,  316. 

Gibbs,  Sir  D.,  on  differences  of  the 
voice  in  different  races  of  men,  566. 

Gibbon,  Hoolock,  nose  of,  150. 

Gibbons,  voice  of,  527. 

Gill,  Dr.,  male  seals  larger  than 
females,  219;  sexual  differences  in 
locals,  515. 


Giraffe,  its  mode  of  using  the  horns, 
508 ;  mute,  except  in  the  rutting 
season,  526. 

Girard,  M.,  disputes  descent  of  verti- 
brates  from  Ascidians,  160  ;  colour 
of  sponges  and  Ascidians,  261  : 
musky  odour  of  Sphinx,  308. 

Girau'd-Teulon,  on  the  cause  of  short 
sight,  34. 

Glanders,  communicable  to  man  from 
the  lower  animals,  7. 

Glands,  odoriferous,  in  mammals,  529, 
530. 

Glareola,  double  moult  in,  390. 

Glomeris  limbata,  difference  of  colour 
in  the  sexes  of,  274. 

Glow-worm,  female,  apterous,  208 ; 
luminosity  of  the,  277. 

Gnats,  dances  of,  280  ;  auditory  powers 
of,  569. 

Gnu,  sexual  differences  in  the  colour 
of  the,  536. 

Goat,  male,  wild,  falling  on  his  horns, 
508  ;  male,  odour  emitted  by,  529  ; 
male,  wild,  crest  of  the,  531  ; 
Berbura,  mane,  dewlap,  &c.,  of  the 
male,  532  ;  Kemns,  sexual  differ- 
ence in  the  colour  oif  the,  536. 

Goats,  sexual  diffei'ences  in  the  horns  of, 
230  ;  horns  of,  235,  505 ;  mode  of 
fighting  of,  508 ;  domestic,  sexual 
differences  of,  late  developed,  237  ; 
beards  of,  531. 

Goatsucker,  Virginian,  pairing  of  the, 
366. 

Gobies,  nidification  of  345. 

God,  want  of  the  idea  of,  in  some  races 
of  men,  93. 

Godron,  M.,  on  variability,  29  ;  on 
difference  of  stature,  31 ;  on  the 
want  of  connexion  between  climate 
and  the  colour  of  the  skin,  192  ;  on 
the  odour  of  the  skin,  198;  on  the 
colour  of  infants,  558. 

Goldfinch,  371,  394;  proportion  of 
the  sexes  in  the,  248 ;  sexual  dif- 
ferences of  the  beak  in  the,  359  ; 
courtship  of  the,  401. 

■ ,  North  American,  young  of,  485 

Gold-fish,  342. 

Gomphus,  proportions  of  the  sexes  in, 
254;  difference  in  the  sexes  of, 
290. 

Gonepteryx  lihamni,  312  ;  sexual  dif- 
ference of  colour  in,  322. 


646 


INDEX. 


GRAY. 


Goodsir,  Prof.,  on  the  affiaity  of  the 
lancelet  to  the  ascidians,  159. 

Goosander,  young  of,  467. 

Goose,  Antarctic,  colours  of  the,  492. 

,  Canada,    pairing    with    a   Ber- 

nicle  gander,  414. 

,  Chinese,    knob  on   the   beak  of 

the,  426. 

,  Egyptian,  364. 

,  Sevastopol,  plumage  of,  385. 

,  Snow-,  whiteness  of  the,  492. 

,  Spur-winged,  364. 

Gorilla,  561  ;  semi-erect  attitude  of 
the,  52 ;  mastoid  processes  of  the, 
53 ;  direction  of  the  hair  on  the 
arms  of  the,  151 ;  manner  of  sit- 
ting, 151 ;  supposed  to  be  a  kind  of 
mandrill,  177  ;  polygamy  of  the  217, 
590,  691 ;  voice  of  the,  527  ;  cra- 
nium of,  558 ;  fighting  of  male, 
562. 

Gosse,  P.  H.,  on  the  pugnacity  of  the 
male  Humming-bird,  360. 

,  M.,  on  the  inheritance  of  artifi- 
cial modifications  of  the  skull,  603. 

Gould,  B.  A.,  on  variation  in  the 
length  of  the  legs  in  man,  26  ; 
measurements  of  American  soldiers, 
30,  32  ;  on  the  proportions  of  the 
body  and  capacity  of  the  lungs  in 
ditferent  races  of  men,  167  ;  on  the 
the  inferior  vitality  of  mulattoes, 
171. 

,  J.,  on  migration  of  swifts,  108  ; 

on  the  arrival  of  male  snipes  before 
the  females,  212;  on  the  numerical 
proportion  of  the  sexes  in  birds, 
247  ;  on  Xeomorpha  Grypus,  359  ; 
on  the  species  of  Eustephanus,  359  ; 
on  the  Australian  musk-duck,  359  ; 
on  the  relative  size  of  the  sexes  in 
Briziura  lohata  and  Cinchramphus 
cruralis,  362  ;  on  Lobivanelhts  loba- 
tus,  366 ;  on  the  habits  of  Menura 
Alberti,  371  ;  on  the  rarity  of  song 
in  brilliant  birds,  371;  on  Selas- 
phorus  platycercus,  378 ;  on  the 
Bower-birds,  381,  406;  on  the 
ornamental  plumage  of  the  Hum- 
ming-birds, 387  ;  on  the  moulting 
of  the  ptarmigan,  392 ;  on  the  dis- 
play of  plumage  by  the  male  Hum- 
ming-birds, 394 ;  on  the  shyness  of 
adorned  male  birds,  403 ;  on  the 
decoratiou  of  the  bowers  of  Bowor- 


birds,  413;  on  the  decoration  of 
their  nests  by  Humming-birds,  413  ; 
on  variation  in  the  genus  Cynan- 
thus,  423 ;  on  the  colour  of  the 
thighs  in  a  male  parakeet,  424  ;  on 
Urosticte  Benjamini,  442,  443 ;  on 
the  nidification  of  the  Orioles,  454  • 
on  obscurely-coloured  birds  build 
ing  concealed  nests,  454;  ontrogons 
and  kingfishers,  456  ;  on  Austra- 
lian parrots,  458  ;  on  Australian 
pigeons,  458  ;  on  the  moulting  of 
the  ptarmigan,  462  ;  on  the  imma- 
ture plumage  of  birds,  466  etseq.; 
on  the  Australian  species  of  Turnix, 
473;  on  the  young  oi  Aithums pohj' 
tmus,  487  ;  on  the  colours  of  the  bills 
of  toucans,  491;  on  the  relative 
size  of  the  sexes  in  the  marsupials 
of  Australia,  515;  on  the  colours 
of  the  Marsupials,  533. 

Goureaux,  on  the  stridulation  of  Mu- 
tilla  europce-T,  293. 

Gout,  sexually  transmitted,  237. 

Graba,  on  the  Pied  Ravens  of  the 
Feroe  Islands,  424;  variety  of  the 
Guillemot,  424. 

Gradation  of  secondaiy  sexual  cha- 
racters in  birds,  430. 

Grallatorfts,  absence  of  secondary 
sexual  characters  in,  219;  double 
moult  in  some,  390. 

Grallina,  nidification  of,  454. 

Grasshoppers,  stridulation  of  the,  286. 

Gratiolet,  Prof,  on  the  anthropo- 
morphous apes,  154 ;  on  the  evo- 
lution of  the  anthropomorphous 
apes,  177  ;  on  the  dilference  in  the 
development  of  the  brains  of  apes 
and  of  man,  203. 

Gray,  Asa,  on  the  gradation  of  species 
among  the  Compositse,  175. 

,  J.  E.,  on  the   caudal  vertebrae 

of  monkeys,  58 ;  on  the  presence 
of  rudiments  of  horns  in  the  female 
of  Cervulus  moschcdus,  504;  on  the 
horns  of  goats  and  sheep,  505  ;  on 
the  beard  of  the  ibex,  531 ;  on  the 
Berbura  goat,  533  ;  on  sexual  dif- 
ferences in  the  coloration  of  Rodents, 
534  ;  ornaments  of  male  sloth,  534; 
on  the  colours  of  the  Elands,  535 ; 
on  the  Sing-sing  antelope,  536 ; 
on  the  colours  of  goats,  536  ;  00  the 
hog-deer,  546. 


INDEX. 


647 


*' Greatest  happiness  principle,"  120, 
121. 

Greeks,  ancient,  140. 

Green,  A.  H.,  on  beavers  fighting, 
500  ;  on  the  voice  of  the  beaver, 
527. 

Greenfinch,  selected  by  a  female 
canary,  415. 

Greg,  W.  R.,  on  the  effects  of  natural 
selection  on  civilised  nations,  133 ; 
ou  the  early  marriages  of  the 
poor,  138  ;  on  the  Ancient  Greeks, 
141. 

Grenadiers,  Prussian,  29. 

Grey,  Sir  G.,  on  female  infanticide  in 
Australia,  592. 

Greyhounds,  numerical  proportion  of 
the  sexes  in,  215,  216  ;  numerical 
proportion  of  male  and  female 
births  in,  246,  258. 

Grouse,  red,  monogamous,  219;  pug- 
nacity of  young  male,  366  ;  pro- 
ducing a  sound  by  scraping  their 
wings  upon  the  ground,  374;  dura- 
tion of  courtship  of,  405  ;  colours 
and  nidification  of,  455. 

Gruber,  Dr.,  on  the  occurrence  of 
the  supra-condyloid  foramen  in  the 
humerus  of  man,  21 ;  on  division 
of  malar  bone,  39  ;  stridulation  of 
locust,  284. 

Griis  americanus,  age  of  mature  plu- 
mage in,  483 ;  breeding  in  im- 
mature plumage,  484. 

vii'go,  trachea  of,  374. 

GnjUus  campestris,  284;  pugnacity 
of  male,  289. 

domesticus,  284. 

Grjjp  IS,  sexual  differences  in  the  beak 
in,  359. 

Guanacoes,  battles  of,  500 ;  canine 
teeth  of,  514. 

Guanas,  strife  for  women  among  the, 
562  ;  polyandry  among  the,  593. 

Guanche  skeletons,  occurrence  of  the 
supra-condyloid  foramen  in  the 
humerus  of,  22. 

Guaranys,  proportion  of  men  and 
women  among,  244  ;  colour  of  new- 
born children  of  the,  557  ;  beards 
of  the,  561. 

Guene'e,  A.,  on  the  sexes  of  Ihjp'vy- 
thra,  251 

Guilding,  L.,  oa  the  stridulation  of 
the  Locustidac,  283 


Guillemot,  variety  of  the,  424. 

Guinea,  sheep  of,  with  males  only 
horned,  234. 

Guinea-fowl,  monogamous,  219  ;  occa- 
sional polygamy  of  the,  220 ; 
markings  of  the,  429. 

Guinea-pigs,  inheritance  of  the  effects 
of  operations  by,  603. 

Gulls,  seasonal  change  of  plumage  in, 
492  ;  white,  492. 

Giinther,  Dr.,  on  paddle  of  Ceratodus, 
37  ;  on  hermaphroditism  in  Serra- 
nus,  162;  on  male  fishes  hatching 
ova  in  their  mouths,  163,  345 ;  on 
mistaking  infertile  female  fishes  for 
males,  249 ;  on  the  prehensile 
organs  of  male  Plagiostomous  fishes, 
331  ;  spines  and  brushes  on  fishes, 
331  ;  on  the  pugnacity  of  the  male 
salmon  and  trout,  332  ;  on  the 
relative  size  of  the  sexes  in  fishes, 

335  ;  on  sexual  differences  in  fishes, 

336  et  seq. ;  on  the  genus  Callio- 
nymus,  337  ;  on  a  protective  re- 
semblance in  a  pipe-fish,  344;  on 
the  genus  Solenostoma,  346  ;  on  the 
coloration  of  frogs  and  toads,  349 ; 
combats  of  Testudo  elegans,  351  ; 
on  the  sexual  differences  in  the 
Ophidia,  351 ;  on  differences  of  the 
sexes  of  lizards,  354  ct  seq, 

Qynanisa    Isis,    ocellated    spots     of, 

428. 
Gypsies,    uniformity     of,    in    various 

parts  of  the  world,  193. 


Habits,  bad,  facilitated  by  familiarity. 
123  ;  variability  of  the  force  of, 
125. 

Hackel,  E.,  on  the  origin  of  man,  3 ; 
on  rudimentary  characters,  11;  on 
death  caused  by  inflammation  of 
the  vermiform  appendage,  21  ;  on 
the  canine  teeth  in  man,  40;  on 
the  steps  by  which  man  became  a 
biped,  52  ;  on  man  as  a  member  of 
the  Catarrhine  group,  155  ;  on  the 
position  of  the  Lemurida;,  157;  on 
the  genealogy  of  the  Mammalia, 
158 ;  on  the  lancelet,  159  ;  on  the 
transparency  of  pelagic  animals, 
261  ;  on  the  musical  powers  of 
women,  573. 


648 


INDEX 


HEMITRAGCS. 


Hagen,    H.,    and     Walsh,    B.  D.,    on 

American  Neuroptera,  254. 
Hair,  development  of,  in  man,  18 ; 
character  of,  supposed  to  be  deter- 
mined by  light  and  heat,  32  ;  distri- 
bution of,  in  man,  57,  600  ;  possibly- 
removed  for  ornamental  purposes, 
58 ;  arrangement  and  direction  of, 
151 ;  of  the  early  progenitors  of 
man,  160  ;  different  texture  of,  in 
distinct  races,  167  ;  and  skin,  cor- 
relation of  colour  of,  197  ;  develop- 
ment of,  in  mammals,  530  ;  manage- 
ment of,  among  different  peoples, 
575 ;  great  length  of,  in  some 
Korth  American  tribes,  580 ;  elon- 
gation of  the,  on  the  human  head, 
603. 

Hairiness,  difference  of,  in  the  sexes 
in  man,  559  ;  variation  of,  in  races 
of  men,  559. 

Hairs  and  excretory  pores,  numerical 
relation  of,  in  sheep,  198. 

Hairy  family,  Siamese,  601. 

Halbertsma,  Prof.,  hermaphroditism 
in  Serranus,  162. 

Haraadryas  baboon,  turning  over 
stones,  101  ;  mane  of  the  male,  521. 

Hamilton,  C,  on  the  cruelty  of  the 
Kaffirs  to  animals,  118;  on  the 
engrossment  of  the  women  by  the 
Kaffir  chiefs,  595. 

Hammering,  difficulty  of,  49. 

Hancock,  A.,  on  the  colours  of  the 
nudibranch  Mollusca,  261,  264. 

Hands,  larger  at  birth,  in  the  chil- 
dren of  labourers,  33  ;  structure  of, 
in  the  quadrumana,  50  ;  and  arms, 
freedom  of,  indirectly  correlated 
with  diminution  of  canines,  53. 

Handwriting,  inherited,  88. 

Handvside,  Dr.,  supernumerary  mam- 
mae in  men,  37. 

Harcourt,  E.  Vernon,  on  Frinjilla 
cannabina  394. 

Hxreldi  glicialis,  420. 

Hare,  protective  colouring  of  the, 
542. 

Hares,  battles  of  male,  500. 

Harlan,  Dr.,  on  the  difference  be- 
tween field-  and  house-slaves,  196. 

Harris,  J.  M.,  on  the  relation  of  com- 
plexion to  climate,  195. 

• ,  T.  W.,  on  the  Katy-did  locust, 

283  ;    on    the   stridulution    of  the 


grasshoppers,  286 ;  on  (Ecanthus 
nivalis,  289 ;  on  the  colouring  of 
Lepidoptera,  314;  on  the  colouring 
of  Saturnia  lo,  316. 

HartingjSpur  of  the  Ornithorhvnchus, 
502. 

Hartman,  Dr.,  on  the  singing  of  Ci.ada 
septendeciin,  282. 

Hatred,  persistence  of,  112. 

Haughton,  S.,  on  a  variation  of  the 
flexor  p-jllicis  longus  in  man,  42. 

Hawks,  feeding  orphan  nestling,  409. 

Hayes,  Dr.,  on  the  diverging  of  sledge- 
dogs  on  thin  ice,  75. 

Haymond,  R.,  on  the  drumming  of  the 
male  Tetrao  umbellus,  375  ;  on  the 
drumming  of  birds,  376. 

Head,  altered  position  of,  to  suit  the 
erect  attitude  of  man,  55;  hairiness 
of,  in  man,  57  ;  processes  of,  in 
male  beetles,  295  ;  artificial  alte- 
rations of  the  form  of  the,  583. 

Hearne,  on  strife  for  women  among 
the  North  American  Indians,  361 ; 
on  the  North  American  Indians' 
notion  of  female  beauty,  578  ;  re- 
peated elopements  of  a  Noi'th  Ame- 
rican woman,  597. 

Heart,  in  the  human  embryo,  9. 

Heat,  supposed  effects  of,  32. 

Hectocotyle,  263. 

Hedge  warbler,  473 ;  voung  of  the, 
481. 

Heel,  small  projection  of,  in  the 
Aymara  Indians,  35. 

Hegt,  M.,  on  the  development  of  the 
spurs  in  peacocks,  236. 

Heliconidae,  308 ;  mimicry  of,  by 
other  butterflies,  323. 

Heliopatlies,  stridulation  peculiar  to 
the  male,  305. 

Heliothrix  auriculjta,  young  of,  467, 
468. 

Helix  pomatia,  example  of  individual 
attachment  in,  263. 

Hellins,  J.,  proportions  of  sexes  of 
Lepidoptera  reared  by.  253. 

Helmholtz,  on  pleasure  derived  from 
harmonies,  92 ;  on  the  vibration  of 
the  auditory  hairs  of  Crustacea, 
568 ;  the  physiology  of  harmony, 
659. 

Hemiptcra,  281. 

Hanitragus,  beardless  in  both  sexes, 
531. 


IIEMSBACII. 


INDEX. 


649 


Hemsbach,  IM.  von,  on  medial  mamma 
in  man,  37. 

Hepburn,  Mr.,  on  the  autumn  song  of 
the  water-ouzel,  370. 

Hepialus  humnU,  sexual  diflerence  of 
colour  in  the,  316. 

Herbs,  poisonous,  avoided  by  animals, 
66.- 

Hermaphroditism  of  embryos,  161. 

Ilerodias  hubulcus,  vernal  moult  of, 
393. 

Heron,  Sir  R.,  on  the  habits  of  pea- 
fowl, 418,  419,  443. 

love-gestures  of  a,  380 

Herons,  decomposed  feathers  in,  385  ; 
breeding  plumage  of,  391,  392 ; 
young  of  the,  481  ;  sometimes 
dimorphic,  484  ;  continued  growth 
of  crest  and  plumes  in  the  males  of 
some,  485 ;  change  of  colour  in 
some,  494. 

Hesperomys  cognatus,  568. 

Hetcerina,  proportion  of  the  sexes  in, 
254 ;  difference  in  the  sexes  of, 
290. 

JTeterocerus,  stridulation  of,  3C2. 

Hewitt,  Mr.,  on  a  game-cock  killing 
a  kite,  363 ;  on  the  recognition  of 
dogs  and  cats  by  ducks,  412  ;  on 
the  pairing  of  a  wild  duck  with  a 
pintail  drake,  415;  on  the  court- 
ship of  fowls,  417  ;  on  the  coupling 
of  pheasants  with  common  hens, 
420. 

Hilgendorf,  sounds  produced  by  crus- 
taceans, 274. 

Hindoo,  his  horror  of  breaking  his 
caste,  122,  124. 

Hindoos,  local  difference  of  stature 
among,  31  ;  difference  of,  from 
Europeans,  192 ;  colour  of  the 
beard  in,  558. 

IlipparcJda  Janira,  319;  instability 
of  the  ocellated  spots  of,  428, 

Hippocampus,  development  of,  163 ; 
marsupial  recejitacles  of  the  male, 
346. 

minor,  202. 

Hippopotamus,  nakedness  of,  56. 

Hips,  proportions  of,  in  soldiers  and 
sailoi's,  32. 

Hodgson,  S.,  on  the  sense  of  duty,  97. 

Hoffberg,  on  the  horns  of  the  rein- 
deer, 503 ;  on  sexual  preferences 
shown  by  reindeer,  525. 


Hoffman,  Prof.,  protective  colours, 
281 ;  fighting  of  frogs,  350. 

Hog-deer,  546 

Hog,  wart-,  519  ;  river-,  520. 

Holland,  Sir  H.,  on  the  effects  of  new 
diseases,  182. 

Homologous  structures,  correlated 
variation  of,  43. 

Homoptera,  281 ;  stridulation  of  the, 
and  Orthoptera,  discussed,  288. 

Honduras,  Quiscalus  major  in,  248. 

Honey-buzzard  of  India,  variation  in 
the  crest  of,  424. 

Honey-suckers,  moulting  of  the,  392  ; 
Australian,  nidification  of,  454. 

Honour,  law  of,  121. 

Hooker,  Dr.,  forbearance  of  elephant 
to  his  keeper,  104  ;  on  the  colour 
of  the  beard  in  man,  558. 

Hookham,  Mr.,  on  mental  concepts  in 
animals,  83. 

Hoolock  Gibbon,  nose  of,  150. 

Hoopoe,  371 ;  sounds  produced  by  the 
male,  376. 

Hoplopterus  armatus,  wing-spurs  of, 
366. 

Hornbill,  African,  inflation  of  the 
neck-wattle  of  the  male  during 
courtship,  383. 

Hornbills,  sexual  difference  in  the 
colour  of  the  eyes  in,  425 ;  nidifi- 
cation and  incubation  of,  454. 

Home,  C,  on  the  rejection  of  a 
brightly-coloured  locust  by  lizards 
and  birds,  289. 

Horns,  sexual  differences  of,  in  sheep 
and  goats,  230  ;  loss  of,  in  female 
merino  sheep,  231  ;  development 
of,  in  deer,  233;  development  of,' 
in  antelopes,  234;  from  the  head 
and  thorax,  in  male  beetles,  297  ; 
of  deer,  503,  506,  515;  and  canine 
teeth,  inverse  development  of, 
514. 

Horse,  fossil,  extinction  of  the,  in 
South  America,  191;  polygamous, 
217;  canine  teeth  of  male,  502; 
winter  change  of  colour,  542. 

Horses,  rapid  increase  of,  in  South 
America,  47  ;  diminution  of  canine 
teeth  in,  53;  dreaming,  74;  of 
the  Falkland  Islands  and  Pam- 
pas, 181  ;  numerical  proportion  of 
the  sexes  in,  215,  216;  lighter  in 
winter    in    Siberia,    229 ;     sexual 


6so 


HOTTENTOT. 


INDEX. 


HVDROPORUS. 


preferences  in,   524;    pairing  pre- 
ferently  with   thitse    of   the    same 
colour,  540 ;   numerical  proportion 
of  male  and  female  births  in,  245 ; 
formerly  striped,  547. 
Hottentot  women,  peculiarities  of,  174. 
Hottentots,  lice  of,  170 ;  readily  be- 
come   musicians,    570 ;    notions  of 
female    beauty  of  the,  578 ;  com- 
pression of  nose  by,  583. 
Hough,    Dr.    S.,    men's    temperature 
more  variable  than  women's,  224  ; 
proportion  of  sexes  in  man,  243. 
House-slaves,  difference  of,  from  iield- 

slaves,  196. 
Houzeau,  on  the  baying  of  the  dog, 
75  ;  on  reason  in  dogs,  76  ;  birds 
killed  by  telegraph  wires,  80  ;  on 
the  cries  of  domestic  fowls  and 
parrots,  85,  87  ;  animals  feel  no 
pity,  102 ;  suicide  in  the  Aleutian 
islands,  117. 
Ho  worth,  H.  H.,  extinction  of  savages, 

183. 
Huber,  P.,  on  ants  playing  together, 
69 ;  on  memory  in  ants,  74 ;  on  the 
intercommunication  of  ants,  89  ;  on 
the    recognition  of  each   other  by 
ants  after  separation,  292. 
Hue,  on  Chinese  opinions   of  the  ap- 
pearance of  Europeans,  578. 
Huia,  the,  of  New  Zealand,  208. 
Human,  man  classed  alone  in  a,  king- 
dom, 147. 

sacrifices,  96. 

Humanity,  unknown  among  some 
savages,  118  ;  deficiency  of,  among 
savages,  123. 
'Humboldt,  A.  von,  on  the  rationality 
of  mules,  78  ;  on  a  parrot  preserv- 
ing the  language  of  a  lost  tribe, 
181 ;  on  the  cosmetic  arts  of 
savag(7s,  574;  on  the  exaggeration 
of  natural  characters  by  man,  582  ; 
on  the  red  painting  of  American 
Indians,  583. 
Hume,    D.,   on  sympathetic   feelings, 

109. 
Humming-bird,  racket-shaped  feathers 
in  the  tail  of  a,  384;  display  of 
})lumage  by  the  male,  394. 
Hiiinining-birds,  ornament  their  nests, 
92,  413;  polygamous,  219;  pro- 
portion of  the  sexes  in,  248,  488  ; 
6€xual    differences   in,   359,    442 


»   I 


pugnacity  of  Piale,  360 ;  modified 
primaries  of  male,  378;  coloration 
of  the  sexes  of,  387 ;  display  by,  443 : 
nidification  of  the,  453 ;  colours  of 
female,  453  ;  young  of,  487. 
Humphrevs,  H.  N.,  on  the  habits  of 

the  stickle-back,  220,  332. 
Hunger,  instinct  of,  112. 
Huns,  ancient,  flattening  of  the  nose 

by  the,  583. 
Hunter,  J.,  on  the  number  of  species 
of  man,  174  ;  on  secondary  sexual 
characters,    207  ;    on    the    general 
behaviour  of  female  animals  during 
courtship,  222  ;  on  the  muscles  of 
the  larynx  in  song-birds,  371;  on 
the  curled  frontal  hair  of  the  bull, 
531  ;  on  the  rejection  of  an  ass  by 
a  female  zebra,  540. 
Hunter,  W.  W.,   on  the  recent  rapid 
increase  of  the  Santali,  45;  on  the 
Sautali,  192. 
Huss,  Dr.  Max,  on  mammary  glands, 

162. 
Hussey,   Mr.,  on  a  partridge    distin- 
guishing persons,  412. 
Hutchinson,  Col.,  example  of  reason- 
ing in  a  retriever,  78. 
Hutton,  Capt.,  on  the  male  wild  goat 

falling  on  his  horns,  507. 
Huxley,  T.  H.,  on  the  structural 
agreement  of  man  with  the  apes, 
2 ;  on  the  agreement  of  the  brain 
in  man  with  that  of  lower  animals, 
6 ;  on  the  adult  age  of  the  orang, 
8  ;  on  the  embryonic  development 
of  man,  9 ;  on  the  origin  of  man, 
3,  11  ;  on  variation  in  the  skulls  of 
the  natives  of  Australia,  26 ;  on 
the  abductor  of  the  fifth  meta- 
tarsal in  apes,  42  ;  on  the  nature 
of  the  reasoning  power,  77  ;  on  the 
position  of  man,  150;  on  the  sub- 
orders of  primates,  152 ;  on  the 
Lemuridce,  157  ;  on  the  Dinosauria, 
158  ;  on  the  amphibian  affinities  of 
the  Ichthyosaurians,  159  ;  on  vari- 
ability of  the  skull  in  certain  races 
of  man,  174;  on  the  races  of  man, 
176;  supplement  on  the  brain,  199. 
Hybrid  birds,  production  of,  414. 
Hydrophobia    communicable  between 

man  and  the  lower  animals,  7. 
Ilydroporus,   dimorphism  of  females 
of,  276. 


HYKLAPllUS. 


INDEX. 


INSECTIVORA. 


651 


Hyelaphus  poreinus,  546. 

Hygrogonus,  345. 

Bi/la,  singing  species  of,  350. 

I/ylobatas,  absence  of  the  thumb  in,  51 ; 
upi-ight  progression  of  some  species 
of,  52 ;  maternal  aflection  in  a, 
70 ;  direction  of  the  hair  on  the 
arms  of  species  of,  151  ;  females  of, 
less  hairy  below  than  males,  558. 

■ agilis,  51  ;  hair  on  the  arms  of, 

151  ;  musical  voice  of  the,  527  ; 
superciliary  ridge  of,  558  ;  voice  of, 
5(J7. 

hooloch,     sexual     difference    of 

colour  in,  537. 

lar,  51  ;  hair   oa   the   arms  of, 

151. 

leuciscns,  51  ;  song  of,  568. 

syndactylus,  51 ;    laryngeal    sac 

of,  527. 

Hylophila  prasinana,  308. 

Hymenoptera,  291  ;  large  size  of  the 
cerebral  ganglia  in,  54 ;  classifica- 
tion of,  148  ;  sexual  differences  in 
the  wings  of,  277  ;  aculeate,  rela- 
tive size  of  the  sexes  of,  279. 

Hymenopteron,  parasitic,  with  a 
sedentary  male,  221. 

Hyomuschus  aquati-us,  547. 

Hyperytkra,  proportion  of  the  sexes 
in,  251. 

Hypogymna  dispar,  sexual  difference 
of  colour  in,  316. 

Hypopyra,  coloration  of,  315. 


Ibex,  male,  falling  on  his  horns,  508 ; 
beard  of  the,  531. 

Ibis,  white,  change  of  colour  of 
naked  skin  in,  during  the  breeding 
season,  389  ;  scarlet,  young  of  the, 
481. 

tantalus,  age  of  mature  plumage 

in,  483 ;  breeding  in  immature 
plumage,  484. 

Ibises,  decomposed  feathers  in,  385 ; 
white,  492  ;  and  black,  493. 

Ichneumouidaj,  difference  of  the  sexes 
in,  292. 

Ic/ithyopterygia,  37. 

Ichthyosaurians,  159. 

Idiots,. microcephalous,  their  charac- 
ters and  habits,  35  ;  hairiness  and 
animal  nature  of  their  actions,  36  : 


microcephalous,  imitative  faculties 
of,  87. 

Iguana  tuberculata,  354. 

Iguanas,  354. 

Illegitimate  and  legitimate  children, 
proportion  of  the  sexes  in,  244. 

Imagination,  existence  of,  in  animals, 
74. 
^Imitation,  68 ;  of  man  by  monkeys, 
72  ;  tendency  to,  in  monkeys,  micro- 
cephalous idiots  and  savages,  87  ; 
inlluence  of,  120. 

Immature  plumage  of  birds,  463,  466. 

Implacentata,  157. 

Implements,  employed  by  monkeys, 
81;  fashioning  of,  peculiar  to 
man,  82. 

Impregnation,  period  of,  influence  of, 
u]ion  sex,  245. 

Improvement,  progressive,  man  alone 
supposed  to  be  capable  of,  79. 

Incisor  teeth,  knocked  out  or  filed  by 
some  savages,  575. 

Increase,  rate  of,  44 ;  necessity  of 
checks  in,  47. 

Indecency,  hatred  of,  a  modern  virtue, 
119. 

India,  difficulty  of  distinguishing  the 
native  races  of,  167 ;  Cyprinidte 
of,  343 ;  colour  of  the  beard  in 
races  of  men  of,  558. 

Indian,  North  American,  honoured 
for  scalping  a  man  of  another  tribe, 
117. 

Individuality,  in  animals,  83. 

Indopicus  carlotta,  colours  of  the 
sexes  of,  458. 

Infanticide,  prevalence  of,  46,  117, 
256  ;  supposed  cause  of,  577  ;  pre- 
valence and  causes  of,  591  et  seq. 

Inferiority,  supposed  physical,  of 
man,  64. 

Inflammation  of  the  bowels,  occur- 
rence of,  in  Cehus  Azara:,  7. 

Inheritance,  27 ;  of  long  and  short 
sight,  33  ;  of  effects  of  use  of  vocal 
and  mental  organs,  88  ;  of  moral 
tendencies,  123,  126;  laws  of, 
227;  sexual,  232;  sexually  limi- 
ted,  444. 

Inquisition,  influence  of  the,  141. 

Insanity,  hereditary,  28. 

Insect,  fossil,  from  the  Devonian,  289. 

Insectivora,  534;  absence  of  secondary 
sexual  characters  in,  218. 


652 


INDEX. 


JKRDON. 


Insects,  relative  size  of  the  cerebral 
gauglia  in,  54;  male,  appearance 
of,  before  the  females,  212  ;  pursuit 
of  female,  by  the  males,  221 ;  period 
of  development  of  sexual  characters 
in,  236 ;  secondary  sexual  cha- 
racters of,  274;  stridulation,  566. 

Insessores,  vocal  organs  of,  370, 

instep,  depth  of,  in  soldiers  and 
sailors,  32. 

Instinct  and  intelligence,  67. 

,    migratory,     vanquishing    the 

maternal,  107,  113. 

Instinctive  actions,  the  result  of  in- 
heritance, 105. 

impulses,  difference  of  the  force 

of,  110,  111;  and  moral  impulses, 
alliance  of,  110. 

Instincts,  66 ;  complex  origin  of, 
through  natural  selection,  67 ; 
possible  origin  of  some,  67 ;  ac- 
quired, of  domestic  animals,  104  ; 
variability  of  the  force  of,  107  ; 
difference  of  force  between  the 
social  and  other.  111,  126  ;  utilised 
for  new  purposes,  571. 

Instrumental  music  of  birds,  375,  378. 

Intellect,  influence  of,  in  natural 
selection  in  civilised  society,  136. 

Intellectual  faculties,  their  influence 
on  natural  selection  in  man,  127  ; 
probably  perfected  through  natural 
selection,  128. 

Intelligence,  Mr.  H.  Spencer  on  the 
dawn  of,  67. 

Intemperance,  no  reproach  among 
savages,  119;  its  destructiveness, 
137. 

Intoxication  in  monkeys,  7. 

Iphias  glaucippc,  313. 

Iris,  sexual  difference  in  the  coiour  of 
the,  in  birds,  383,  425. 

Ischio-pubic  muscle,  41. 

Wiajmis  crucntu<,  number  of  spurs 
in,  364. 

lult'S,  tarsal  suckers  of  the  males  of, 
274 

J. 

Jackals  learning  from  dogs  to  bark, 

73. 
Jack-snipe,  coloration  of  the,  491 
Jacquinot,  on  the  number  of  species 

of  man,  174. 
Jaeger,  Dr.,  l(;ngth  of  bones  increased 


from  carrying  weights,  32;  on  the 
difficulty  of  approaching  herds  of 
wild  animals,  100 ;  male  Silver- 
pheasant,  rejected  when  his  plu- 
mage was  spoilt,  419. 

Jaguars,  black,  539. 

Janson,  E.  W.,  on  the  proportions  of 
the  sexes  in  Tomicus  villusus,  253  ; 
on  stridulant  beetles,  302. 

Japan,  encouragement  of  licentious- 
ness in,  46. 

Japanese,  general  beardlessness  of  the, 
•  560;  aversion  of  the,  to  whiskers, 
581. 

Jai'dine,  Sir  W.,  on  the  Argus  phea- 
sant, 384,  403. 

Jarrold,  Dr.,  on  modifications  of  the 
skull  induced  by  unnatural  position, 
56. 

Jarves,  Mr.,  on  inf^mticide  in  the 
Sandwich  Islands,  257. 

Javans,  relative  height  of  the  sexes 
of,  559  ;  notions  of  female  beaut v, 
580. 

Jaw,  influence  of  the  muscles  of  the, 
upon  the  physiognomy  of  the  apes, 
54. 

Jaws,  smaller  proportionately  to  the 
extremities,  33 ;  influence  of  food 
upon  the  size  of,  33 ;  diminution 
of,  in  man,  53 ;  in  man,  reduced 
by  correlation,  562. 

Jay,  young  of  the,  481  ;  Canada, 
young  of  the,  481. 

Jays,  new  mates  found  by,  407  ;  dis- 
tinguishing persons,  412. 

Jeffreys,  J.  Gwyn,  on  the  form  of  the 
shell  in  the  sexes  of  the  Gastero- 
poda, 262  ;  on  the  influence  of  light 
upon  the  colours  of  shells,  263. 

Jelly-fish,  bright  colours  of  some, 
260. 

Jenner,  Dr.,  on  the  voice  of  the  rook, 
375;  on  the  finding  of  new  mates  by 
magpies,  407;  on  retardation  of  the 
generative  functions  in  birds,  409. 

Jenyns,  L.,  on  the  desertion  of  their 
young  by  swallows,  108  ;  on  male 
birds  singing  after  the  proper 
season,  409. 

Jerdon,  Dr.,  on  birds  dreaming,  74 ; 
on  the  pugnacity  of  the  male  bul- 
bul,  360  ;  on  the  pugnacity  of  the 
male  Ortygornis  gularis,  363 ;  on 
the  spurs  of  GalloperdUj  364:',    on 


JEV0N3. 


INDEX. 


KIRBY. 


653 


the  habits  of  Lobivanellus,  366  ;  on 
the  spoonbill,  374;  on  the  drum- 
mino-  of  the   Kalij-pheasant,  376 ; 
on  fndian  bustards,  378;  on   Otis 
bengalensis,  381  ;    on  the  ear-tufts 
of  Sypheotidcs  auritus,  384  ;  on  the 
double  moults  of  certain  birds,  391 ; 
on    the    moulting    of   the    honey- 
Suckers',  392  ;  on  the  moulting  of 
bustards,  plovers,  and  drongos,  393  ; 
on  the  spring  change  of  colour  in 
some   finches,    393;    on  display  in 
male  birds,  394 ;  on  the  display  ot 
the  under-tail  coverts  by  the  male 
bulbul,  402  ;  on  the   Indian  honey- 
buzzard,  424  ;  on  sexual  differences 
in  the  colour  of  the  eyes  of  horn- 
bills,  425  ;  on  the  markings  of  the 
Tragopan   pheasant,   428;    on    the 
niditication  of  the  Orioles,  453  ;  on 
the    nidification   of  the    hornbills, 
454  ;  on  the  Sultan  yellow-tit,  458  ; 
on  Falceomisjavanicus,  461  ;  on  the 
immature   plumage    of  birds,  465 
et  seq.  ;  on  representative  species  ot 
birds,  468  ;  on  the  habits  of  Turnup 
476;  on  the  continued  incre^ase  of 
beauty    of  the    peacock,    485;    on 
coloration  in  the  genus  Palxornis, 
494. 
Jevons,  W.  S.,  on  the  migrations  ot 

man,  47. 
Jews,  ancient  use  of  flmt  tools  by  the, 
145;    uniformity    of,     in    various 
parts  of  the  world,  193  ;  numerical 
proportion    of    male    and    female 
births    among   the,  243;    ancient, 
tattooing  practised  by,  574. 
Johnstone,  Lieut.,  on  the  Indian  ele- 
phant, 218. 
JoUofs,  tine  appearance  of  the,  587. 
Jones,  Albert,  proportion  of  sexes  of 

Lepidoptera,  reared  by,  253. 
Juan    Fernandez,   humming-birds   of, 

487.  •     ^      , 

Junonia,  sexual  differences  of  colour- 
ing in  species  of,  310. 
Jupiter,    comparison    with    Assyrian 
effigies,  581. 


Kaffir    skull,  occurrence  of  the  dia- 
stema in  a,  40. 
Kaffirs,  their  cruelty  to  animals,  118  ; 


lice  of  the,  170;  colour  ot  the, 
579 ;  engrossment  of  the  hand- 
somest women  by  the  chiefs  of  the, 
595  ;  marriage-customs  of  the,  598. 
Kalij-pheasant,     drumming     of     the 

male,  375  ;  young  of,  468. 
Kallima,  resemblance  of,  to  a  withered 

leaf,  311. 
Kalmucks,  general   beardlessness    of, 
560;  aversion  of,  to   hairs  on  the 
face,    581  ;     marriage-customs    of 
the,  598. 
Kangaroo,  great  red,  sexual  difference 

in  the  colour  of,  533. 
Kant,  Imm.,  on   duty,   97  ;    on  self- 
restraint,  110;  on  the  number  of 
species  of  man,  174. 
Katy-did,  stridulation  of  the,  283. 
Keen,  Dr.,  on  the   mental  powers  of 

snakes,  352. 
Keller,    Dr.,    on    the     difficulty    of 

fashioning  stone  implements,  49. 
Kent,  W.  S.,  elongation  of  dorsal  fin 
of  Callionymus    li/ra,  336 ;    court- 
ship of  Lubrus  mivtus,  341 ;  colours 
and  courtship  of  Cantharus  lineatus^ 
341. 
Kestrels,  new  mates  found  by,  408. 
Kidney,   one,  doing  double  work  in 

disease,  32. 
King,  W.  R.,  on  the  vocal  organs  of 
Tetrao  cupido,  371  ;  on  the  drum- 
ming of  grouse,  376  ;  on  the  rein- 
deer,  503;    on   the   attraction    of 
male   deer   by   the   voice    of    the 
female,  526. 
King  and  Fitzroy,  on  the  marriage- 
customs  of  the  Fuegians,  599. 
King-crows,  nidification  of,  453. 
Kingfisher,  371;  racket-shaped  feathers 

in  the  tail  of  a,  384. 
Kingfishers,  colours   and  nidification 
of  the,  455,  457,   459;  immature 
plumage  of  the,  467,  468  ;  young 
of  the,  481. 
King  Lory,  457  ;  immature  plumage 

of  the,  467. 
Kingsley,  C,  on  the  sounds  produced 

by  Uinhrina,  347. 
Kirby  and  Spence,  on  sexual  differ- 
ences in  the  length  of  the  snout  in 
Curculionidae,  208;  on  the  court- 
ship of  insects,  221 ;  on  the  elytra 
of  Djti.^cus,  276  ;  on  peculiarities 
in   the  legs  of  male  insects,  276  ; 


654 


INDEX. 


on  the  relative  size  of  the  sexes  in 
insects,  279 ;  on  the  Fulgoridae, 
281  ;  on  the  habits  of  Termites^ 
291 ;  on  difference  of  colour  in  the 
sexes  of  beetles,  294  ;  on  the  horns 
of  the  male  lamellicorn  beetles, 
297  ;  on  hornlike  processes  in  male 
Curculionidse,  299  ;  on  the  pug- 
nacity of  the  male  stag-beetle,  299. 

Kite,  killed  by  a  game-cock,  363, 

Knot,  retention  of  winter  plumage  by 
the,  391 

Knox,  R.,  on  the  semilunar  fold,  17  ; 
on  the  occurrence  of  the  supra- 
eondyloid  foramen  in  the  humerus 
of  man,  21 ;  on  the  features  of  the 
young  Memnon,  168. 

Koala,  length  of  the  coecum  in,  20. 

Kohus  elli20siprymnus,  proportion  of 
the  sexes  in,  24:7. 

Kolreuter,  on  the  sterility  of  hybrid 
plants,  172, 

Koodoo,  development  of  the  horns  of 
the,  234  ;  markings  of  the,  543. 

Koppen,  F,  T,,  on  the  migratory  lo- 
cust, 283. 

Kordofan,  protuberances  artificially 
produced  by  natives  of,  574. 

Koraks,  marriage  customs  of,  598, 

Korte,  on  the  proportion  of  sexes  in 
locusts,  254  ;  Piussian  locusts,  283. 

Kovalevsky,  A.,  on  the  affinity  of 
the  Ascidia  to  the  Vertebrata,  159, 
160. 

,  W,,  on    the   pugnacity  of  the 

male  capercailzie,  363 ;  on  the 
pairing  of  the  capercailzie,  367. 

Krause,  on  a  convoluted  body  at  the 
extremity  of  the  tail  in  a  Macacus 
and  a  cat,  23. 

Kupffer,  Prof,  on  the  affinity  of  the 
Ascidia  to  the  Vertebrata,  160. 


Ld'ndocera  Pancinii,  prehensile  organs 
of  the  male,  266 

Labrus,  splendid  colours  of  the  species 
of,  342. 

inutus,    sexual     differences    in, 

337,  341. 

pavo,  342. 

Lacertilia,  sexual  differences  of,  354. 

Lifresuaye,  M.  de,  on  birds  of  para- 
disc,  385. 


Lamarck,  on  the  origin  of  man,  3. 

Lamellibranchiata,  262. 

Lamellicorn  beetles,  horn-like  pro- 
cesses from  the  head  and  thorax  of, 
295,  298  ;  influence  of  sexual  selec- 
tion on,  301. 

Lamellicornia,  stridulation  of,  303. 

Lament,  Mr.,  on  the  tusks  of  the 
walrus,  502 ;  on  the  use  of  its 
tusks  by  the  walrus,  513  ;  on  the 
bladder-nose  seal,  528. 

Lampornis  porphyrurus,  colours  of 
the  female,  454. 

Lampyridae,  distasteful  to  mammals, 
277. 

Lancelet,  159,  165. 

Landois,  H.,  gnats  attracted  by  sound, 
280  ;  on  the  production  of  sound 
by  the  Cicadse,  281  ;  on  the  stridu- 
lating  organ  of  the  crickets,  284  ; 
on  Decticus,  285 ;  on  the  stridula- 
ting  organs  of  the  Acridiidae,  286  ; 
stridulating  apparatus  in  Orthop- 
tera,  288  ;  sounds  produced  by 
Atropus,  291 ;  on  the  stridulation 
of  Necrophorns,  302  ;  on  the  stridu- 
lant  organ  of  Cerambyx  heros,  303  ; 
on  the  stridulant  organ  of  Geo- 
tinipes,  303  ;  on  the  stridulating 
organs  in  the  Coleoptera,  304 ;  on 
the  ticking  of  Andbium,  306. 

Landor,  Dr.,  on  remorse  for  not  obey 
ing  tribal  custom,  114. 

Language  an  art,  86 ;  articulate,  origin 
of,  86  ;  relation  of  the  progress  of, 
to  the  development  of  the  brain, 
87  ;  effects  of  inheritance  in  pro- 
duction of,  88  ;  complex  structure 
of,  among  barbarous  natrons,  91; 
natural  selection  in,  91  ;  gesture, 
178  ;  primeval,  180;  of  a  lost  tribe 
preserved  by  a  parrot,  181. 

Languages,  presence  of  rudmients  m, 
90 ;  classification  of,  90  ;  variability 
of,  90  ;  crossing  or  blending  of,  90  ; 
complexity  of,  no  test  of  perfection 
or  proof  of  special  creation,  92  ;  re- 
semblance of,  evidence  of  commu- 
nity of  origin,  148. 

and  species,  identity  of  evidence 

of  their  gradual  development,  90. 

Lanius,  462  ;  characters  of  voung, 
464. 

rufiis,     anomalous    rouug    of, 

482. 


LANKE3TER. 


INDEX. 


LIBKLLULID/E. 


655 


Lankester,  E.  R.,  on  comparative  lon- 
gevity, 13o,  136  ;  on  the  destruc- 
tive effects  of  intemperance,  137. 

Lanugo,  of  the  human  foetus,  19, 
600. 

Lapponian  language,  highly  artificial, 
91. 

Lark,  proportion  of  the  sexes  in  the, 
248  ;  female,  singing  of  the,  370. 

Larks,  attracted  by  a  mirror,  413. 

Lartet,  E.,  comparison  of  cranial  ca- 
pacities of  skulls  of  recent  and 
tertiary  mammals,  55  ;  on  the  size 
of  the  brain  in  mammals,  81;  on 
Dryopithecus,  155  ;  on  pre-historic 
flutes,  570. 

Larus,  seasonal  change  of  plumage  in, 
492. 

Larva,  luminous,  of  a  Brazilian  beetle, 
278. 

Larynx,  muscles  of  the,  in  song-birds, 
371. 

Lasiocampa  quercus,  attraction  of 
males  by  the  female,  252  ;  sexual 
difference  of  colour  in,  316. 

Latham,  R.  G.,  on  the  migrations  of 
man,  48. 

Latooka,  perforation  of  the  lower  lip 
by  the  women  of,  576. 

I.aunllard,  on  the  abnormal  division 
of  the  malar  bone  in  man,  39. 

Lawrence,  W.,  on  the  superiority  of 
savages  to  Europeans  in  power  of 
sight,  33  ;  on  the  colour  of  negro 
infants,  558  ;  on  the  fondness  of 
savages  for  ornaments,  578  ;  on 
beardless  races,  581  ;  on  the  beauty 
of  the  English  aristocracy,  586. 

Lavard,  E.  L.,  on  an  instance  of  i"a- 
tionality  in  a  cobra,  352;  on  the 
pugnacity  of  Gallus  Stanleyi,  363. 

Laycock,  Dr.,  on  vital  periodicity,  8  ; 
theioid  nature  of  idiots,  36. 

Leaves,  autumn,  tints  useless,  262. 

Lecky,  Mr.,  on  the  sense  of  duty,  97  ; 
on'  suicide,  117  ;  on  the  practice  of 
celibacy,  119 ;  his  view  of  the 
crimes  of  savages,  119  ;  on  the 
gradual  rise  of  morality,  125. 

Leconte,  J.  L.,  on  the  stridulant  organ 
in  the  Coprini  and  Dynastini,  303. 

Lee,  H.,  on  the  numerical  proportion 
of  the  sexes  in  the  trout,  249. 

Leg,  calf  of  the,  artificially  modified, 
574. 


Legitimate  and  illegitimate  children, 
proportion  of  the  sexes  in,  244. 

Legs,  variation  of  the  length  of  the,  in 
man,  26 ;  proportions  of,  in  sol- 
diers and  sailors,  32  ;  front,  atro- 
phied in  some  male  butterflies,  277  ; 
peculiarities  of,  in  male  insects,  277. 

Leguay,  on  the  occurrence  of  the 
supra-condyloid  foramen  in  the 
humerus  of  man,  22. 

"  Lek  "  of  the  black-cock  and  caper- 
cailzie, 405. 

Lemoine,  Albert,  on  the  origin  of  lan- 
guage, 87. 

Lcimir  macaco,  sexual  difierence  of 
colour  in,  537, 

Lemurida^,  152;  ears  of  the,  15; 
variability  of  the  muscles  in  the, 
41 ;  position  and  derivation  of  the, 
157;  their  origin,  165. 

Lemurs,  uterus  in  the,  38. 

Lenguas,  disfigurement  of  the  ears  of 
the,  575. 

Leopards,  black.  539. 

Lepidoptera,  307  ;  numerical  propor- 
tions of  the  sexes  in  the,  250 ; 
colouring  of,  308  ;  ocellated  spots 
of,  427. 

Lepidosircn,  159,  165. 

J>eptalides,  mimicry  of,  325. 

Leptorhynchiis  angustatus,  pugnacity 
of  male,  299. 

Leptura  testacea,  difference  of  colour 
in  the  sexes  of,  294. 

Leroy,  on  the  wariness  of  young  foxes 
in  hunting-districts,  80;  on  the 
desertion  of  their  young  by  swal- 
lows, 108. 

Leslie,  D.,  marriage  customs  of  Kaffirs, 
598. 

Lesse,  valley  of  the,  22. 

Lesson,  on  the  birds  of  paradise,  219, 
403  ;  on  the  sea-elephant,  528. 

Lessona,  M.,  observations  on  Scrran  'S, 
162. 

Lethrns  ccphalotes,  pugnacity  of  the 
males  of,  297,  300. 

Leuckart,  R.,  on  the  vesicula  prosta- 
tica,  24  ;•  on  the  influence  of  the 
age  of  parents  on  the  sex  of  off- 
spring, 245. 

Levator  clavicula:  muscle,  42. 

Libelliila  dcpressa,  colour  of  the  male, 
290. 

Libellulidae,  relative  size  of  the  sexes 


656 


LICE. 


INDEX. 


LUBBOCK. 


of,  279  ;  difierence  in  the   sexes  of, 
290. 

Lice  of  domestic  animals  and  man, 
169. 

Licentiousness,  a  check  upon  popuLi- 
lion,  46 ;  prevalence  of,  among 
savages,  119. 

Lichtenstein,  on  Chera  progne,  4J  9. 

Life,  inheritance  at  corresponding 
periods  of,  228,  232. 

Light,  effects  on  complexion,  32 ;  in- 
fluence of,  upon  the  colours  of 
shells,  263. 

Lilford,  Lord,  the  ruff  attracted  by 
bright  objects,  413. 

Limosa  lapponka,  478. 

Linaria,  462. 

montuna,  248. 

Lindsay,  Dr.  W.  L.,  diseases  commu- 
nicated from  animals  to  man,  7  ; 
madness  in  animals,  79  ;  the  dog 
considers  his  master  his  God,  96. 

Linnaeus,  views  of,  as  to  the  position 
of  man,  149. 

Linnet,  numerical  proportion  of  the 
sexes  in  the,  248 ;  crimson  fore- 
head and  breast  of  the,  394 ;  court- 
ship of  the,  401. 

Lion,  polygamous,  218  ;  mane  of  the, 
defensive,  521  ;  roaring  of  the,  526. 

Lions,  stripes  of  young,  464. 

Lips,  piercing  of  the,  by  savages,  575. 

Lithohius,  prehensile  appendages  of 
the  female,  274. 

Lithosia,  coloration  in,  314. 

L'ittorina  llttorea,  262. 

Livingstone,  Dr.,  manner  of  sitting  of 
gorilla,  151  ;  on  the  influence  of 
dampness  and  dryness  on  the  colour 
of  the  skin,  193  ;  on  the  liability 
of  negroes  to  tropical  fevers  after 
residence  in  a  cold  climate,  194; 
on  the  spur-winged  goose,  364;  on 
weaver-birds,  376  ;  on  an  African 
night-jar,  384,  403  ;  on  the  battle- 
scars  of  South  African  male  mam- 
mals, 500  ;  on  the  removal  of  the 
upper  incisors  by  the  Batokas,  575; 
on  the  perforation  of  the  uj)per  lip 
by  the  Makalolo,  576  ;  on  the 
Banyai,  579. 

Livonia,  numerical  proportion  of  male 
and  female  births  in,  215,  243. 

Lizards,  relative  size  of  the  sexes 
of,  354  ;  gular  pouches  of,  354. 


Lloyd,  L.,  on  the  polygamy  of  the 
capercailzie  and  bustard,  219  ;  on 
the  numerical  proportion  of  the 
sexes  m  the  capercailzie  and  black- 
cock, 248  ;  on  the  salmon,  333  ; 
on  the  colours  of  the  sea-scorpion, 
337 ;  on  the  pugnacity  of  male 
grouse,  364;  on  the  capercailzie 
and  black-cock,  366,  370 ;  on  th^ 
call  of  the  capercailzie,  375  ;  on 
assemblages  of  grouse  and  snipes, 
405 ;  on  the  pairing  of  a  shield- 
drake  with  a  common  duck,  414 ; 
on  the  battles  of  seals,  500 ;  on  the 
elk,  507. 

Lohivanellus,  wing-spurs  in,  366. 

Local  influences,  effect  of,  upon  sta- 
ture, 31. 

Lock  wood,  Mr.,  on  the  development 
of  Hippocampus,  163. 

,  Rev.  S.,  musical  mouse,  568. 

Locust,  bright-coloured,  rejected  by 
lizards  and  birds,  289. 

,   migratorv,  283 ;  selection    by 

female,  283. 

Locusts,  proportion  of  sexes  in,  254 ; 
stridulation  of,  284. 

Locustidae,  stridulation  of  the,  282, 
284  ;  descent  of  the,  285. 

Longicorn  beetles,  difference  of  the 
sexes  of,  in  colour,  294  ;  stridula- 
tion of,  303. 

Lonsdale,  Mr.,  on  an  examjde  of  per 
sonal  attachment  in  Helix  poiU'jtia, 
263. 

Lophobranchii,  marsupial  receptacles 
of  the  males,  346. 

Lophophorus,  habits  of,  420. 

Lophorina  atra,  sexual  ditference  in 
coloration  of,  491. 

Lophornis  ornatiis,  387 

Lord,  J.  K.,  on  Salmo  lycaodon,  333. 

Lorv,  Kino;,  457  ;  immature  plumage 
of  the,  467. 

Love-antics  and  dances  of  birds,  380. 

Lowne,  B.  T.,  on  Musca  vomitoria, 
54,  280. 

Loxia,  characters  of  young  of,  464. 

Lubbock,  Sir  .J.,  on  the  antiquity  of 
man,  2  ;  on  the  origin  of  man,  3 ; 
on  the  mental  capacity  of  savages, 
65 ;  on  the  origin  of  implements, 
82  ;  on  the  simplification  of  lan- 
guages, 92  ;  on  the  absence  of  the 
idea  of  God  among  certain  races  of 


lucakid;e. 


INDEX.. 


MACaiLLIVRAV. 


657 


men,  94;  on  the  origin  of  the 
belief  in  spiritual  agencies,  95; 
on  superstitions,  96  ;  on  the  sense 
of  duty,  97  ;  on  the  practice  of 
burying  the  old  and  sick  among 
the  Fijians,  102  ;  on  the  im- 
morality of  savages,  119;  on  JMr. 
Wallace's  claim  to  the  origination 
of  the  idea  of  natural  selection, 
49 ;  on  the  absence  of  remorse 
among  savages,  131  ;  on  the  former 
barbarism  of  civilised  nations,  143  ; 
on  improvements  in  the  arts  among 
savages,  144  ;  on  resemblances  of 
the  mental  characters  in  different 
races  of  men,  178 ;  on  the  arts 
practised  by  savages,  179 ;  on  the 
power  of  counting  in  primeval  man, 
180;  on  the  prehensile  organs  of 
the  m»le  Labidocera  Darwinii,  266  ; 
on  Chloeon,  274 ;  on  Smynthurus 
luteus,  279  ;  finding  of  new  mates 
by  jays,  407  ;  on  strife  for  women 
among  the  North  American  Indians, 
561  ;  on  music,  570;  on  the  orna- 
mental practices  of  savages,  574 ; 
on  the  estimation  of  the  beard 
among  the  Anglo-Saxons,  581  ; 
on  artificial  deformation  of  the 
skull,  583  ;  on  "  communal  mai-- 
riages,"  587,  588 ;  on  exogamy, 
589,  592;  on  the  Veddahs,  591; 
on  polyandry,  593. 

Lucanidse,  variability  of  the  mandibles 
in  the  male,  300. 

Lucanus,  large  size  of  males  of,  278. 

cervus,    numerical     proportion 

of  sexes  of,  253  ;  weapons  of  the 
male,  299. 

elaphus,  use  of   mandibles  of, 

300  ;  large  jaws  of  male,  275. 

Lucas,  Prosper,  on  pigeons,  418;  on 
sexual  preference  in  horses  and 
bulls,  525. 

Lunar  periods,  8,  164. 

Lund,  Dr.,  on  skulls  found  in  Bra- 
zilian caves,  168 

Lungs,  enlargement  of,  in  the  Qui- 
chua  and  Aymara  Indians,  34; 
a  modified  swim-bladder,  161  ; 
different  capacity  of  in  races  of 
man,  167. 

Luminosity  in  insects,  277. 

Luschka,  Prof.,  on  the  termination  of 
the  coccyx,  23. 

2^ 


Luxury,  expectation  of  life  unin- 
fluenced by,  136. 

Lycccna,  sexual  differences  of  colom*- 
ing  in  species  of,  310. 

Lyell,  Sir  C,  on  the  antiquity  of 
man,  2 ;  on  the  origin  of  man, 
3 ;  on  the  parallelism  of  the  de- 
velopment of  species  and  languages, 
90 ;  on  the  extinction  of  lan- 
guages, 90 ;  on  the  Inquisition, 
141  ;  on  the  fossil  remains  of  ver- 
tebrata,  157 ;  on  the  fertility  of 
mulattoes,  171. 

Lynx,  Canadian,  throat-ruff  of  the, 
521. 

Lyre-bird,  assemblies  of,  406. 


Macacus,  ears  of,  15;  convoluted 
body  in  the  extremity  of  the  tail 
of,  23  ;  A'ariability  of  the  tail  in 
species  of,  58  ;  whiskers  of  species 
of,  531, 

hrunneus,  59. 

cynomol'jus,   superciliary  ridge 

of,  558  ;  beard  and  whiskers  of, 
becoming  white  with  age,  559. 

ecaudatus,  60. 

lasiutus,  facial  spots  of,  550.  • 

radiatics,  151. 

rhesus,  sexual  difference    in  the 

colour  of,  539,  550. 

Macalister,  Prof.,  on  variations  of 
the  palmaris  accessorius  muscle, 
27  ;  on  muscular  abnormalities  in 
man,  42,  43  ;  on  the  greater  varia- 
bility of  the  muscles  in  men  than 
in  women,  223. 

Macaws,  Mr.  Buxton's  observations 
on,  102;  screams  of,  375. 

McCanu,  J.,  on  mental  individuality, 
84. 

McClelland,  J.,  on  the  Indian  Cypri- 
nidiE,  343. 

Macculloch,  Col.,  on  an  Indian  vil- 
lage without  any  female  children, 
592. 

,  Dr.,  on  tertian  ague  in  a  dog,  8. 

Macgillivray,  W.,  on  the  vocal  organs 
of  birds,  90 ;  on  the  Egyptian 
goose,  365  ;  on  the  habits  of  wood- 
peckers, 376  ;  on  the  habits  of  the 
snipe,  377 ;  on  the  whitethroat, 
381 ;  on  the  moulting  of  the  snipes ; 


658 


INDEX. 


MAMMALIA. 


391 ;  on  the  moulting  of  the 
Anatidae,  393  ;  on  the  finding  of 
new  mates  by  magpies,  407  ;  on 
the  pairing  of  a  blackbird  and 
thrush,  414  ;  on  pied  ravens,  424  ; 
on  the  guillemots,  424 ;  on  the 
colours  of  the  tits,  458  ;  on  the 
immature  plumage  of  birds,  466 
et  seq. 

Machetes,  sexes  and  young  of,  485  ; 

^ugnax,  supposed  to  be  polyga- 
mous, 219  ;  numerical  proportion 
of  the  sexes  in,  248  ;  pugnacity  of 
the  male,  360 ;  double  moult  in,  390. 

Mcintosh,  Dr.  colours  of  the  Nemer- 
tians,  265. 

McKennan,  marriage  customs  of 
Koraks,  598. 

Mackintosh,  on  the  moral  sense,  97. 

MacLachlan,  R.,  on  Apatania  mulie- 
hris  and  Boreus  hy emails,  254 ; 
on  the  anal  appendages  of  male 
insects,  275  ;  on  the  pairing  of 
dragon-flies,  279  ;  on  dragon-flies, 
290,  291  ;  on  dimorphism  in 
Agrion,  291  ;  on  the  want  of  pug- 
nacity in  male  dragon-flies,  291  ; 
colour  of  ghost-moth  in  the  Shet- 
land Islands,  316. 

McLennan,  Mr.,  on  infanticide,  46, 
591 ;  on  the  origin  of  the  belief  in 
spiritual  agencies,  94;  on  the  pre- 
valence of  licentiousness  among 
savages,  119,  588;  on  the  primi- 
tive barbarism  of  civilised  nations, 
143  ;  on  traces  of  the  custom  of 
the  forcible  capture  of  wives,  144, 
592  ;  on  polyandry,  593. 

Macnamara,  Mr.,  susceptibility  of 
Andaman  islanders  and  Nepalese 
to  change,  188. 

McNeill,  Mr.,  on  the  use  of  the  ant- 
lers of  deer,  510  ;  on  the  Scotch 
deerhound,  516;  on  the  long  haii's 
on  the  throat  of  the  stag,  522 ;  on 
the  bellowing  of  stags,  526. 

Macropus,  courtship  of,  341. 

Macrorhinus  proboscideus,  structure 
of  the  nose  of,  528. 

Magpie,  power  of  speech  of,  90 ; 
nuptial  assemblies  of,  406  ;  new 
mates  found  by,  407  ;  stealing 
bright  objects,  4i;? ;  young  of  the, 
481 ;  coloration  of  the,  493. 

Magpies,  vocal  organs  of  the,  370. 


Maillard,  M.,  on  the  proportion  of 
the  sexes  in  a  species  of  Papiiio 
from  Bourbon,  250. 

Maine,  Sir  Henry,  on  the  absorption 
of  one  tribe  by  another,  128;  a 
desire  for  miprovement  not  general, 
132. 

Major,  Dr.  C.  Forsyth,  on  fossil 
Italian  apes,  155;  skull  of  Bos 
'  etruscus,  505 ;  tusks  of  miocene 
pigs,  521. 

Makalolo,  perforation  of  the  upper 
lip  by  the,  576. 

Malar  bone,  abnormal  division  of,  in 
man,  39. 

Malay,  Archipelago,  marriage-cus- 
toms of  the  savages  of  the,  598. 

Malays,  line  of  separation  between 
the  Papuans  and  the,  169  ;  general 
beardlessness  of  the,  560  f  staining 
of  the  teeth  among,  574 ;  aversion 
of  some,  to  hairs  on  the  face, 
581. 

and  Papuans,  contrasted  cha- 
racters of,  168. 

Male  animals,  struggles  of,  for  the 
possession  of  the  females,  212,  213  ; 
eagerness  of,  in  courtship,  221, 
222  ;  generally  more  modified  than 
female,  221,  223  ;  differ  in  the,  same 
way  from  females  and  young,  232. 

characters,  developed  in  females, 

227  ;  transfer  of,  to  female  birds, 
471. 

,  sedentary,  of  a  hymenopterous 

parasite,  221, 

Malefactors,  137. 

Males,  presence  of  rudimentary  female 
organs  in,  162, 

and  females,  comparative  num- 
bers of,  213,  215;  comparative 
mortality  of,  while  young,  216. 

Malherbe,  on  the  woodpeckers,  458. 

Mallotus  peronii,  331. 

■ villosuSy  331. 

Malthus,  T,,  on  the  rate  of  increase 
of  population,  44,  45.  46, 

Maluridic,  niditication  of  the,  454-. 

Malurus,  young  of,  485. 

Mamma2,  208 ;  rudimentary,  in  male 
mammals,  11,  23,  161,  162,163: 
supernumerary,  in  women,  36 ;  of 
male  human  subject,  37. 

Mammalia,  Prof.  Owen's  classification 
of,  148;  genealogy  of  the,  158. 


MAMMALS. 


INDEX. 


MAYHEW. 


659 


Mammals,  recent  and  tertiary,  com- 
parison of  cranial  capacity  of,  55 ; 
nipples  of,  162;  pursuit  of  female, 
by  the  males,  221 ;  secondary  sexual 
characters  of,  500 ;  weapons  of, 
501 ;  i-elative  size  of  the  sexes  of, 
515;  parallelism  of,  with  birds  in 
secondary  sexual  characters,  541 ; 
voices  of,  used  especially  during  the 
breeding  season,  567. 

Man,  variability  of,  26  ;  erroneously 
regarded  as  more  domesticated  than 
other  animals,  28 ;  migrations  of, 
47 ;  wide  distribution  of,  48  ;  causes 
of  the  nakedness  of,  57  ;  supposed 
physical  inferiority  of,  64  ;  a  mem- 
ber of  the  Catarrhine  group,  155  ; 
early  progenitors  of,  160;  transition 
from  ape  indefinite,  180;  numerical 
proportions  of  the  sexes  in,  215; 
diO'erence  between  the  sexes,  223 ; 
proportion  of  sexes  amongst  the 
illegitimate,  244 j  different  com- 
plexion of  male  and  female  negroes, 
556  ;  secondary  sexual  characters  of, 
556  ;  primeval  condition  of,  594. 

Mandans,  correlation  of  colour  and 
texture  of  hair  in  the,  197. 

Mandible,  left,  enlarged  in  the  male 
of  Tapliroderes  distortus,  21 Q. 

Mandibles,  use  of  the,  in  Arnmophila, 
275  ;  large,  of  Corydalis  cornutus, 
275;  large,  of  male  Lucanus  ela- 
phus,  275. 

Mandrill,  number  of  caudal  vertebrae 
in  the,  58  ;  colours  of  the  male, 
538,  541,  550. 

Mantegazza,  Prof.,  on  last  molar 
teeth  of  man,  20  ;  bright  colours 
in  male  animals,  224  ;  on  the  orna- 
ments of  savages  573  et  seq.  ;  on 
the  beardlessness  of  the  New  Zea- 
landers,  581 ;  on  the  exaggeration  of 
natural  characters  by  man,  582. 

Mantell,  W.,  on  the  engrossment  of 
pretty  girls  by  the  IS'ew  Zealand 
chiefs,  595. 

Mantis,  pugnacity  of  species  of,  289. 

Maories,  mortality  of,  184;  infanti- 
cide and  proportion  of  sexes,  256  ; 
distaste  for  hairiness  amongst  men, 
581. 

Marcus  Aurelius,  on  the  origin  of  the 
moral  sense,  98  ;  on  the  influence 
of  habitual  thou:j,hts,  123. 


Mareca  pcnclope,  414. 

Marks,  retained  throughout  groups 
of  birds,  427. 

Marriage,  restraints  upon,  among  sa- 
vages, 46  ;  influence  of,  upon  morals, 
119;  influence  of,  on  mortality, 
139  ;  development  of,  590. 

Marriages,  early,  138,  139  ;  com- 
munal, 587,  589. 

IMarshall,  Dr.  \V.,  protuberances  on 
birds'  heads,  235,  383 ;  on  the 
moulting  of  birds,  393  ;  advantage 
to  older  birds  of  paradise,  485. 

,    Col.,    interbreeding     amongst 

Todas,  189;  infanticide  and  pro- 
portion of  sexes  with  Todas,  255  ; 
choice  of  husbands  amongst  Todas, 
593. 

,  Mr.,  on  the  brain  of  a  Bush- 
woman,  167. 

Marsupials,  157  ;  development  of  the 
nictitating  membrane  in,  17;  uterus 
of,  39 ;  possession  of  nipples  by, 
162;  their  origin  from  Monotre- 
mata,  165 ;  abdominal  sacks  of, 
208  ;  relative  size  of  the  sexes  of, 
515  ;  colours  of,  533. 

Marsupium,  rudimentary,  in  male 
marsupials,  161. 

Martin,  W.  C.  L.,  on  alarm  manifested 
by  an  orang  at  the  sight  of  a  turtle, 
72  ;  on  the  hair  in  Ilylobatcs,  152  ; 
on  a  female  American  deer,  514  ; 
on  the  voice  of  Hylobates  agiiis, 
527  ;  on  Semnopithecus  nemccus,  552. 

.  on  the  beards  of  the  inhabitants 

of  St.  Kilda,  560. 

Martins  deserting  their  young,  108. 

,  C,  on  death  caused  by  inflam- 
mation uf  the  vermiform  appen- 
dage, 21. 

Mastoid  processes  in  man  and  apes,  53. 

Maudsley,  Dr.,  on  the  influence  of  the 
sense  of  smoU  in  man,  18;  on 
idiots  smelling  their  food,  36  ;  on 
Laura  Bridgman,  88  ;  on  the  de- 
velopment of  the  A'ocal  organs,  89  ; 
moral  sense  failing  in  iucij>ient 
madness,  124 ;  change  of  mental 
faculties  at  puberty  in  man,  565. 

Mayers,  W.  F.,  on  the  domestication 
of  the  goldfish  in  China,  343. 

Mayhew,  E.,  on  the  affection  between 
individuals  of  different  sexes  in  the 
dog,  523. 


66o 


JIAYNARD. 


INDEX. 


SIOCtGKIDGE. 


Maynard,  C.  J.,  on  the  sexes  of  Chrij- 

semi/s  picta,  351. 
Meckel,  on  correlated  variation  of  the 

muscles  of  the  arm  and  leg,  44, 
Medicines,    ciFect    produced    by,    the 

same  in  man  and  in  monkeys,  7. 
Medusce,  bright  colours  of  some,  260. 
Megalithic  structures,  prevalence  of, 

179. 
Megapicus   validus,   sexual  difference 

of  colour  in,  458. 
Megasoma,    large   size    of    males    of, 

279. 
Meigs,  Dr.   A.,   on   variation    in    the 

skulls  of  the   natives   of  Amei-ica, 

26. 
Meinecke,  on  the   numerical  propor- 
tion  of   the    sexes    in   butterflies, 

250. 
Melanesians,  decrease  of,  185. 
Meldola,   Mr.,  colours  and   marriage 

flight  of  Colias  and  Fieris,  319. 
Meliphagidse,  Australian,  nidification 

of,  454. 
Melita,   secondary  sexual    characters 

of,  268. 
Meloe,    difference    of   colour    in   the 

sexes  of  a  species  of,  294. 
Memory,    manifestations   of,    in    ani- 
mals, 74. 
Memnon,  young,  168. 
Mental    characters,    difference    of,   in 

different  races  of  men,  167. 
faculties,   diversity   of,    in    the 

same  race  of  men,  26  ;  inheritance 

of,  27  ;   variation   of,  in  the  same 

species,  27,  QQ ;    similarity  of  the 

in  different  races  of  man,  178  ;  of 

birds,  410. 
■ •  powers,    difference    of,    in    the 

two  sexes  in  man,  563. 
Menura  Alberti,  406  ;  song  of,  371. 
superba,  406  ;   long  tails  of  both 

sexes  of,  451. 
Merqanser,     trachea    of    the     male, 

374. 
■  serrator,  male  plumage  of,  393. 

Mergus  cucullatus,  speculum  of,  236. 

merganser,  young  of,  467. 

Mctal/ura,   splendid    tail-feathers   of, 

443. 
Methoca  ichncumoniJes,  large  male  of, 

279. 
Meves,   M.,  on  the  drumming  of  the 

snipe,  377. 


Jlexicans,  civilisation  of  the,  not 
foreign,  145. 

Meyer,  on  a  convoluted  body  at  tht 
extremity  of  the  tail  in  a  Macacus 
and  a  cat,  23. 

,  Dr.    A.,   on  the   copulation   of 

Phryganidce  of  distinct  species,  275. 

,    Prof.    L.,    on    development    of 

helix  of  ear,  15,  16 ;  men's  ears 
more  variable  than  women's,  224  ; 
antennae  serving  as  ears,  280. 

Migrations  of  man,  effects  of,  47. 

Migratory  instinct  of  birds,  105 ; 
vanquishing  the  maternal,  107, 
113. 

Mill,  J.  S.,  on  the  origin  of  the  moral 
sense,  98  ;  on  the  "  greatest  hap- 
piness principle,"  120  ;  on  the  dif- 
ference of  the  mental  powers  in 
the  sexes  of  man,  564. 

Millipedes,  274. 

Milne-Edwards,  H.,  on  the  u'se  of  the 
enlarged  chelae  of  the  male  Gelasi- 
musy  267. 

Ililvago  leucurus,  sexes  and  young  of, 
479. 

Mimicry,  323. 

Mimus  polyglottus,  411. 

Mind,  difference  of,  in  man  and  the 
highest  animals,  126  ;  similarity 
of  the,  in  different  races,  178. 

Minnow,  proportion  of  the  sexes  in 
the,  249. 

Mirror,  larks  attracted  by,  413. 

Mitchell,  Dr.,  interbreeding  in  the 
Hebrides,  189. 

Mitford,  selection  of  children  in 
Sparta,  29. 

Mivart,  St.  George,  on  the  reduction 
of  organs,  12  ;  on  the  ears  of  the 
lemuroidea,  15 ;  on  variability  of 
the  muscles  in  lemuroidea,  41,  48  ; 
on  the  caudal  vertebrae  of  monkeys, 
58 ;  on  the  classiflcation  of  the 
primates,  153  ;  on  the  orang  and 
on  man,  154  ;  on  differences  in  the 
lemuroidea,  155  ;  on  the  crest  of 
the  male  newt,  348. 

JMobius,  Prof.,  on  reasoning  powers 
in  a  pike,  75. 

Mocking-thrush,  partial  migration  of, 
411 ;  young  of  the,  487. 

Jlodiflcations,  unserviceable,  02. 

Moggridge,  J.  T.,  on  habits  of  spiders, 
69;  on  habits  of  ants,  147. 


MOLES. 


INDEX. 


MOTACILLS:. 


66i 


Moles,  numerical  proportion  of  the 
sexes  in,  247  ;  b;ittles  of  male,  500. 

Alolliencsia  pctencnsis,  sexual  ditier- 
ence  in,  3o7. 

Mollusca,  beautiful  colours  and  shapes 
of,  2(J3 ;  absence  of  secondary 
sexual  characters  in  the,  2(J2. 

Molluscoida,  159,  2G2. 

Monacanthus  scopas  and  M.  Feronii, 
331. 

Mongolians,  perfection  of  the  senses 
in,  3-1-. 

Monkey,  protecting  his  keeper  from 
a  baboon,  103,  110;  bonnet-,  151; 
rhesus-,  sexual  difference  in  colour 
of  the,  539,  550 ;  moustache-, 
colours  of  the,  537. 

Monkeys,  liability  of,  to  the  same 
diseases  as  man,  7  ;  male,  recog- 
nition of  women  by,  8 ;  diversity 
of  the  mental  faculties  in,  27  ; 
breaktng  hard  fruits  with  stones, 
50;  hands  of  the,  50,  51;  basal 
caudal  vertebrae  of,  imbedded  in  the 
body,  59 ;  revenge  taken  by,  ti9 ; 
maternal  affection  in,  70 ;  varia- 
bility of  the  fiiculty  of  attention 
in,  74  ;  American,  manifestation  of 
reason  in,  77 ;  using  stones  and 
sticks,  81 ;  imitative  faculties  of, 
87 ;  signal-cries  of,  87  ;  mutual 
kindnesses  of,  101;  sentinels  posted 
by,  101  ;  human  characters  of, 
150;  American,  direction  of  the 
hair  on  the  arms  of  some,  151  ; 
gradation  of  species  of,  175;  beards 
of,  531  ;  ornamental  characters  of, 
549  ;  analogy  of  sexual  differences 
of,  with  those  of  man,  558 ;  dif- 
ferent degrees  of  difference  in  the 
sexes  of,  561;  expression  of  emo- 
tions by,  572  ;  generally  mono- 
gamous habits  of,  590  ;  polygamous 
habits  of  some,  590 ;  naked  surfaces 
of,  600. 

Monogamy,  not  primitive,  144. 

Monogenists,  176. 

Mononijchus  pseudacori,  stridulation 
of,  305. 

Monotremata,  157  ;  development  of 
the  nicitating  membrane  in,  17; 
lactiferous  glands  of,  162  ;  connect- 
ing mammals  with  reptiles,  165. 

Monstrosities,  analogous,  in  man  and 
lower  animals,  30  ;  caused  by  arrest 


of  development,  35  ;  correlation  of, 
44;  transmission  of,  173. 

Montagu,  G.,  on  the  habits  of  the 
bhick  and  red  grouse,  219;  on  the 
pugnacity  of  the  ruft",  361  ;  on  the 
singing  of  birds,  368;  on  tlie 
double  moult  of  the  male  pintail, 
393. 

]\Ionteiro,  Mr.,  on  Bucorax  abyssi- 
nicus,  383. 

Montes  de  Oca,  M.,  on  the  pugnacity 
of  male  Humming-birds,  360. 

Montlcola  cyanca,  456. 

Monuments,  as  traces  of  •extinct 
tribes,  181. 

Moose,  battles  of,  501  ;  horns  of  the, 
an  incumbrance,  515, 

Moral  and  instinctive  impulses,  alli- 
ance of,  111. 

faculties,     their    influence    on 

natural  selection  in  man,  127. 

rules,   distinction    between  the 

higher  and  lower,  122. 

sense,   so-called,  derived    from 

the  social  instipcts,  120,  121;  origin 
of  the,  124. 

tendencies,  inheritance  of,  124. 

Morality,  supposed  to  be  founded  in 
selfishness,  120  ;  test  of,  the  general 
welfare  of  the  community,  121; 
gradual  rise  of,  125;  influence  of  a 
high  standard  of,  132. 

Morgan,  L.  H.,  on  the  beaver,  67  ;  on 
the  reasoning  powers  of  the  beaver, 
75 ;  on  the  forcible  capture  of 
wives,  144;  on  the  castoreum  of 
the  beaver,  529  ;  marriage  unknown 
in  primeval  times,  588;  on  poly- 
andry, 593. 

Morlcy.  J.,  on  the  appreciation  of 
})raise  and  fear  of  blame,  146. 

Morris,  F.  0.,  on  hawks  feeding  an 
orphan  nestling,  409. 

Morse,  Dr.,  colours  of  mollusca,  264. 

Moi-selli,  E.,  division  of  the  malar 
bone,  39. 

Mortality,  comparative,  of  females 
and  males,  216,  243. 

Morton,  on  the  number  of  species  of 
man,  174. 

Moschkau,  Dr.  A.,  on  a  speaking 
starling,  85. 

Moschus  moschiferus,  odoriferous  or- 
gans of,  529. 

Motacillcc,  Indian,  young  of,  468. 


662 


MOTHS. 


INDEX. 


MUSIC. 


Moths,  313;  absence  of  mouth  in 
some  males,  208 ;  apterous  female, 
208  ;  male,  prehensile  use  of  the 
tarsi  by,  209 ;  male,  attracted  by 
females,  252;  coloration  of,  315; 
sexual  differences  of  colour  in,  316. 

Motmot,  inheritance  of  mutilation  of 
tail  feathers,  60,  603;  racket- 
shaped  feathers  in  the  tail  of  a, 
384. 

Moult,  double,  463;  double  annual, 
in  birds,  390. 

Moulting  of  birds,  484. 

Moults,»partial,  392, 

Mouse,  song  of,  568. 

Moustache-monkey,  colours  of  the, 
537,  552. 

Moustaches,  in  monkeys,  150. 

Mud-turtle,  long  claws  of  the  male, 
350. 

Mulattoes,  persistent  fertility  of, 
171;  immunity  of,  from  yellow 
fever,  193. 

Mule,  sterility  and  strong  vitality  of 
the,  171. 

Mules,  rational,  78. 

Muller,  Ferd.,  on  the  Mexicans  and 
Peruvians,  145. 

,  Fritz,  on  astomatous  males  of 

Tanais,  208  ;  on  the  disappearance 
of  spots  and  stripes  in  adult  mam- 
mals, 547  ;  on  the  proportions  of 
the  sexes  in  some  Crustacea,  255 ; 
on  secondary  sexual  characters  in 
various  Crustaceans,  265  et  seq. ; 
musical  contest  bet-.veen  male  Ci- 
cada:, 282 ;  mode  of  holding  wings 
inCastnia,  315  ;  on  birds  shewing  a 
preference  for  certain  colours,  317  ; 
on  the  sexual  maturity  of  young 
amphipod  Crustacea,  485. 

— — ,  Hermann,  emergence  of  bees 
from  pupa,  214;  pollen-gathering 
of  bees,  228  ;  proportion  of  sexes 
m  bees,  254  ;  courting  of  Uristalis, 
280 ;  colour  and  sexual  selection 
with  bees,  292. 

,  J.,  on  the  nictitating  membrane 

and  semilunar  fold,  17. 

■ ,  Max,  on  the  origin  of  language, 

87  ;  language  implies  power  of 
general  conception,  88 ;  struggle 
for  life  among  the  words,  &c.,  of 
languages,  91. 

,  S.,  on  the  banteug,   536 ;    on 


the  colours  of  SemnopiiJiecus  chry- 
somelas,  537. 

Muntjac-deer,  weapons  of  the,  514. 

Murie,  J.,  on  the  reduction  of  organs, 
12  ;  on  the  ears  of  the  Lemuroidea, 
15  ;  on  variability  of  the  muscles 
in  the  Lemuroidea,  41,  48  ;  basal 
caudal  vertebras  of  Maca:us  hrun- 
neus  imbedded  in  the  body,  69 ; 
on  the  manner  of  sitting  in  short- 
tailed  apes,  59 ;  on  diti'erences  in 
the  Lemuroidea,  155;  on  the 
throat-pouch  of  the  male  bustard, 
373;  on  the  mane  of  Otaria  juhata, 
521 ;  on  the  sub-orbital  pits  of 
Ruminants,  529  ;  on  the  colours  of 
the  sexes  in  Otaria  nigresceiis,  535. 

Murray,  A.,  on  the  Pediculi  of  dif- 
ferent races  of  men,  169. 

T.  A.,  on  the  fertility  of  Austra- 
lian women  with  white  men,  170. 

Mus  coninja,  80. 

minutw,  sexual  difference  in  the 

colour  of,  534. 

Musca  vomitoria,  54. 

Muscicapa  grisola,  455. 

luctuosa,  455. 

ruticilla,  breeding  in  immature 

plumage,  484. 

Muscle,  ischio-pubic,  41. 

Muscles,  rudimentary,  occurrence  of, 
in  man,  12  ;  variability  of  the,  26  ; 
effects  of  use  and  disuse  upon,  32 ; 
animal-like  abnormalities  of,  in 
man,  41  ;  correlated  variation  of, 
in  the  arm  and  leg,  44  ;  variability 
of,  in  the  hands  and  feet,  48 ;  of 
the  jaws,  influence  of,  on  the  phy- 
siognomy of  the  Apes,  54;  habitual 
spasms  of,  causing  modifications  of 
the  facial  bones,  55  ;  of  the  early 
progenitors  of  man,  160 ;  greater 
variability  of  the,  in  men  than  in 
women,  223. 

Musculns  sternalis.  Prof.  Turner  on 
the,  13. 

Music,  178  ;  of  birds,  368  ;  discord- 
ant, love  of  savages  for,  380  ;  rea- 
son of  power  of  perception  of  notes 
in  animals,  568  ;  power  of  distin- 
guishing notes,  569  ;  its  connection 
with  primeval  speech,  670 ;  differ- 
ent appreciation  of,  by  different 
peoples,  670;  origin  of,  569,  573 
effects  of,  671. 


MUSICAL. 


INDEX. 


Ni:W  ZEALAND. 


663 


Musical   cadences,   perception   of,  by 

animals,  569  ;  powers  of  man,  566 

et  seq. 
Musk-deer,  canine  teeth  of  male,  502, 

513,   514- ;  male,   odiferous   organs 

of  the,  529  ;  winter  change  of  the, 

542. 
Musk-duck,    Australian,    359 ;    large 

size  of  male,  362  ;  of  Guiana,  pug- 
nacity of  the  male,  362. 
Musk-ox,  horns  of,  505. 
Musk-rat,   protective   resemblance  of 

the,  to  a  clod  of  earth,  542. 
Musophagcc,   colours  and  nidification 

of  the,  455  ;  both  sexes  of,  equally 

brilliant,  460. 
Mussels  opened  by  monkeys,  50. 
Mustela,  winter  change  of  two  species 

of,  542. 
Musters,    Capt.,    on    Jihea   Barwinii, 

479  ;    marriages    amongst  Patago- 

nians,  598. 
Mutilations,    healing    of,   8;    inheri- 
tance of,  60. 
Mutilia  europcea,  stridulation  of,  292. 
Mutillidae,  absence  of  ocelli  in  female, 

275. 
Mycetes    caraya,    polygamous,    217  ; 

vocal  organs  of,  527  ;  beard  of,  531 ; 

sexual    difterences    of    colour    in, 

537  ;  voice  of,  567. 
■ seniculus,    sexual    differences  of 

colour  in,  537. 
Myriapoda,  274. 

N. 

Nageli,  on  the  influence  of  natural 
selection  on  plants,  61  ;  on  the 
gradation  of  species  of  plants,  175. 

Nails,  coloured  yellow  or  purple  in 
part  of  Africa,  574. 

Narwhal,  tusks  of  the,  502,  507. 

Nasal  cavities,  large  size  of,  in  Ame- 
rican aborigines,  34. 

Nascent  organs,  12, 

Nathusius,  H.  von,  on  the  improved 
breeds  of  pigs,  177 ;  male  domes- 
ticated animals  more  variable  than 
females,  223;  horns  of  castrated 
sheep,  506 ;  on  the  breeding  of 
domestic  animals,  596. 

Natural  selection,  its  effects  on  the 
early  progenitors  of  man,  47  ;  in- 
fluence of,  on  man,  60,  62  ;  limita- 


tion of  the  principle,  61  ;  influence 
of,  on  social  animals,  62  ;  Mr,  Wal- 
lace on  the  limitation  of,  by  the 
influence  of  the  mental  faculties  in 
man,  127  ;  influence  of,  in  the  pro- 
gress of  the  United  States,  142  ;  in 
relation  to  sex,  259. 
Natural  and  sexual  selection  con- 
trasted, 226. 
Naulette,  jaw  fi-om,  large  size  of  the 

canines  in,  40. 
Neanderthal   skull,  capacity    of  the, 

55. 
Neck,  proportion  of,  in  soldiers  and 

sailors,  32. 
Kecrophorus,    stridulation    of,     302, 

304. 
Kectarinia,  young  of,  468. 
Kedannice,    moulting    of    the,    392 ; 

nidification  of,  454. 
Negro,  resemblance  of  a,  to  Europeans, 

in  mental  characters,  178. 
Negro-women,  their  kindness  to  Mungo 

Park,  118. 
Negroes,  Caucasian  features  in,  167  ; 
character  of,  168  ;  lice  of,  170  ;  fer- 
tility of,  when  crossed  with  other 
races,  171;  blackness  of,  170,  173; 
variability  of,  174,175;  immunity 
of,  from  yellow  fever,193;  diflerence 
of,  from  Americans,  197  ;  disfigure- 
ments of  the,  541  ;  colour  of  new- 
born children  of,  557  ;  comparative 
beardlessness  of,   560  ;  readily  be- 
come musicians,  570 ;  appreciation 
of  beauty  of  their  women  by,  577, 
579 ;  idea   of  beauty  among,  582  ; 
compression   of  the   nose   by  some, 
583. 
Nemertians,  colours  of,  264. 
Neolithic  period,  145. 
Neomorpha,   sexual  difTerence  of  the 

beak  in,  359. 
Nephila,  size  of  male,  273. 
Nests,   made  by  fishes,  345 ;  decora- 
tion of,  by  Humming-birds,  413. 
Neumeister,  on  a  change  of  colour  in 
pigeons    after   several    moultings, 
238. 
Neuration,  difference  of,  in  the  two 
sexes   of  some   butterflies  and  hy- 
menoptera,  277. 
Neuroptera,  254,  290. 
Neurothemis,  dimorphism  in,  291. 
New    Zetland,    expectation     bjr    the 


664 


INDEX. 


natives  of,  of  their  extinction,  191; 
practice  of  tattooing  in,  576  ;  aver- 
sion of  natives  of,  to  hairs  on  the 
face,  581 ;  pretty  girls  engrossed 
by  the  chiefs  in,  595. 
Newton,  A.,  on  the  throat-pouch  of 
the  male  bustard,  373;  on  the  dif- 
ference between  the  females  of  two 
species  of  Oxynotus,  470 ;  on  the 
habits  of  the  Phalarope,  dotterel, 
and  godwit,  477. 
Newts,  348. 

Nicholson,  Dr.,  on  the  non-immunity 
of  dark    Europeans    from    yellow 
fever,  195. 
Nictitating  membrane,  17,  101, 
Nidification,  of  fishes,  344  ;  relation 
of,  to  colour,  453,  456  ;  of  British 
birds,  454. 
Night-heron,  cries  of  the,  368. 
Nightingale,     arrival     of    the     male 
before  the  female,  212;  object  of  j 
the  song  of  the,  368. 
Nightingales,    new    mates  found   by 

408. 
Nightjai",  selection  of  a  mate  by  the 
female,  416;    Australian,  sexes  of 
479  ;  coloration  of  the,  491. 
Nightjars,  noise   made  by  some  male, 
with  their  wings,  376  ;  elongated 
feathers  in,  384,  403. 
Nilghau,  sexual  differences   of  colour 

in  the,  535. 
Nilsson,  Prof.,  on  the  resemblance  of 
stone    arrow-heads    from    various 
places,  179  ;  on  the  development  of 
the  horns  in  the  reindeer,  234. 
Nipples,  absence  of,  in  Monotremata, 

162. 
Nitsche,  Dr.,  ear  of  foetal  orang,  17. 
Nitzsch,  C.  L.,  on  the  down  of  birds, 

390. 
Noctuai,    brightly-coloured    beneath, 

315. 
Noctuidoe,  coloration  of,  313. 
Nomadic      habits,     unfavourable     to 

human  progress,  133. 
Nordmann,  A.,  on  Tetrao  urogalloidcs, 

405. 
Norfolk  island,  half-breeds  on,  190. 
Norway,  numerical  proportion  of  male 

and  female  births  in,  243. 
No.se,  resemblance  of,  in  man  and  the 
apes,  153;  piercing  and  ornamen- 
tation of  the,  575  ;    very  flat,  not 


admired  in  negroes,  582 ;  flattening 
of  the,  583. 

Nott  and  Gliddon,  on  the  features  of 
Kameses  II.,  168;  on  the  features 
of  Amunoph  111.,  168  ;  on  skulls 
from  Brazilian  caves,  168 ;  on  the 
immunity  of  negroes  and  mulattoes 
from  yellow  fever,  193  ;  on  the  de- 
formation of  the  skull  among  Ame- 
rican tribes,  583. 

No  vara,  voyage  of  the,  suicide  m  New 
Zealand,  117. 

Nudibranch  MoUusca,  bright  colours 
of,  264. 

Numerals,  Roman,  144;  origin  of, 
264. 

Nunemaya,  natives  of,  bearded,  349, 
560. 

Nut-hatch  of  Japan,  intelligence  of, 
410. 


Obedience,  value  of,  130. 

Observation,  powers  of,  possessed  bv 
birds,  411. 

Occupations,  sometimes  a  cause  of 
diminished  stature,  31 ;  effect  of, 
upon  the  proportions  of  the  bodv, 
31. 

Ocelli,  absence  of,  in  female  Mutil- 
Mx,  274. 

of  birds,  formation  and  vari- 
ability of  the,  427. 

Ocelot,  sexual  differences  in  the  colour- 
ing of  the,  534. 

Oci/phaps  lophotes,  402. 

Odonata,  254.  / 

Odonestis  potatoria,  sexual  difference 
of  colour  in,  316. 

Odour,  correlation  of,  with  colour  of 
skin,  197  ;  emitted  by  snakes  in 
the  breeding-season,  352  ;  of  mam- 
mals, 528. 

(Ecanthus  nivalis,  difference  of  colour 
in  the  sexes  of,  289. 

pellucidus,  289. 

Ogle,  Dr.  W.,  relation  between  colour 

and  power  of  smell,  18. 
Oidemia,  491. 

Olivier,  on  sounds  produced  by 
Pimclia  striata,  306. 

Omahplia    brunnea,   stridulation    of, 

303. 
Onitis  furcifcr,  processes  of  anterior 


ONTHOPHAGUS. 


INDEX 


66s 


femora  of  the  male,  and  on  the 
head  and  thorax  of  the  female, 
297. 

Onthophagus,  295. 

rangifer,   sexual  differences    of, 

296  ;  variation  in  the  horns  of  the 
male,  297. 

Ophidia,  sexual  differences  of,  351. 

Ophidium,  347. 

Opossum,  wide  range  of,  in  America, 
169. 

Optic  nerve,  atrophy  of  the,  caused 
oy  destruction  of  the  eye,  32. 

Orang-Outan,  561  ;  Bischoff  on  the 
agreement  of  the  brain  of  the, 
with  that  of  man,  6 ;  adult  age 
of  the,  8  ;  ears  of  the,  14 ;  ver- 
miform appendage  of,  21 ;  hands 
of  the,  50  ;  absence  of  mastoid  pro- 
cesses In  the,  53  ;  platforms  built 
by  the,  Q^  ;  alarmed  at  the  sight  of 
a  turtle,  72  ;  using  a  stick  as  a 
lever,  81 ;  using  missiles,  81 ;  using 
the  leaves  of  the  Pandanus  as  a 
night  covering,  82 ;  direction  of 
the  hair  on  the  arms  of  the,  151  ; 
its  aberrant  characters,  154 ;  sup- 
posed evolution  of  the,  177  ;  voice 
of  the,  527 ;  monogamous  habits 
of  the,  590 ;  male,  beard  of  the, 
531. 

Oranges,  treatment  of,  by  monkeys, 
50. 

Orange-tip  butterfly,  308,  312,  313. 

Orchestia  Darwinii^  dimorphism  of 
males  of,  268. 

Tucuratinga,  limbs  of,  267,  271. 

Ordeal,  trial  by,  96. 

Oreas  canna,  colours  of,  535, 

Derhianus,  colours  of,  535,  543. 

Organs,  prehensile,  209  ;  utilised   for 

new  purposes,  571. 

Organic  scale,  von  Baer's  definition  of 
jirogress  in,  164. 

Orioles,  nidification  of,  453. 

Oriolus,  species  of,  breeding  in  im- 
mature plumage,  484. 

melanocephalus,  coloration  of  the 

sexes  in,  460. 

Ornaments,     prevalence    of    similar, 

.  179;  of  male  birds,  367  ;  fondness 
of  savages  for,  574. 

Ornamental  characters,  equal  trans- 
mission of,  to  both  sexes,  in  mam- 
mals, 541;  of  monkeys,  549. 


Ornithoptera  crcesus,  250. 

Ornithorhynchus,  156 ;  reptilian  ten- 
dency of,  159 ;  spur  of  the  male, 
502. 

Orocetes  erythrognstra,  voung  of,  487. 

Orrony,  Grotto  of,  22.  ' 

Orsodacna  atra,  difference  of  colour  in 
the  sexes  of,  294. 

Orthoptera,  282 ;  metamorphosis  of, 
237 ;  stridulating  apparatus  of, 
283,  288;  colours  of,  289;  rudi- 
mentary stridulating  organs  in 
female,  288;  stridulation  of  the, 
and  Homoptera,  discussed,  288. 

Ortygornis  gularis,  pugnacity  of  the 
male,  363. 

Oryctcs,  stridulation  of,  303  ;  sexual 
differences  in  the  stridulant  organs 
of,  305. 

Oryx  leucoryx,  use  of  the  horns  of, 
509,  518. 

Osphranter  rufus,  sexual  difference  in 
the  colour  of,  533. 

Ostrich,  African,  sexes  and  incuba- 
tion of  the,  478. 

Ostriches,  stripes  of  young,  464. 

Otaria  jubata,  mane  of  the  male, 
521. 

nigrescens,     difference     in     the 

coloration  of  the  sexes  of,  534. 

Otis  bengalensis,  love-antics  of  the 
male,  380. 

tarda,  throat-pouch  of  the  male, 

373  ;  polygamous,  219. 

Ouzel,  ring-,  colours  and  nidification 
of  the,  455. 

,  water-,  colours  and  nidifica- 
tion of  the,  455. 

Ovibos  mosc'tatus,  horns  of,  505. 

Ovipositor  of  insects,  208. 

Oris  cycloceros,  mode  of  fighting  of, 
508,  513. 

Ovule  of  man,  9. 

Owen,  Prof.,  on  the  Corpora  Wolf- 
fiana,  11 ;  on  the  great  toe  in  man, 
11  ;  en  the  nictitating  membrane 
and  semilunar  fold,  17;  on  the 
development  of  the  posterior  mo- 
lars in  different  races  of  man,  20 ; 
on  the  length  of  the  ca?cum  in  the 
Koala,  20 ;  on  the  coccygeal  ver- 
tebra?, 23  ;  on  rudimentary  struc- 
tures belonging  to  the  reproductive 
system,  24 ;  on  abnormal  condi- 
tions   of   the    human    uterus,   38 , 


666 


INDEX. 


PARAGUAY. 


on  the  number  of  digits  in  the 
Ichthyopteiygia,  37  ;  on  the  canine 
teeth  in  man,  40 ;  on  the  walking 
of  the  chimpanzee  and  orang,  50 ; 
on  the  mastoid  processes  in.  the 
higner  apes,  53 ;  on  the  hairiness 
of  elephants  in  elevated  districts, 
57 ;  on  the  caudal  vertebras  of 
monkeys,  58  ;  classification  of  mam- 
malia, 148 ;  on  the  hair  in  mon- 
keys, 152 ;  on  the  piscine  affinities 
of  the  Ichthyosaurians,  159 ;  on 
pologamy  and  monogamy  among 
the  antelopes,  217;  on  the  horns 
of  Antilocapra  americana,  234  ;  on 
the  musky  odour  of  crocodiles 
during  the  breeding  season,  351  ; 
on  the  scent-glands  of  snakes,  352  ; 
on  the  Dugong,  Cachalot  and  Cr- 
nithorhynchus,  502  ;  on  the  antlers 
of  the  red  deer,  510  ;  on  the  den- 
tition of  the  Camelidas,  514 ;  on  the 
horns  of  the  Irish  elk,  515  ;  on 
the  voice  in  the  giraffe,  porcupine, 
and  stag,  526  ;  on  the  laryngeal 
sac  of  the  gorilla  and  orang,  527  ; 
on  the  odoriferous  glands  of  mam- 
mals, 528,  529;  on  the  effects  of 
emasculation  on  the  vocal  organs  of 
men,  566  ;  on  the  voice  of  Hylo- 
bates  agilis,  567  ;  on  American 
monogamous  monkeys,  567. 

Owls,   white,   new    mates  found   by, 
408. 

Oxynotus,  difference  of  the  females  of 
two  species  of,  470. 


P. 


Pachydermata,  218. 

Pachytylus  migratorius,  283. 

Paget,  on  the  abnormal  development 
of  hairs  in  man,  19  ;  on  the  thick- 
ness of  the  skin  on  the  soles  of  the 
feet  of  infants,  33. 

Painting,  pleasure  of  savages  in,  178. 

Palccmnn,  chelae  of  a  si)ecies  of,  267. 

Pal(vornis,  sexual  diflerences  of  colour 
in,  494. 

—— javanicua,  colour  of  beak  of, 
461. 

rosa,  young  of,  467. 

Palamedea  cornuta,  spurs  on  the 
wings,  364. 


Paleolithic  period,  145. 

Palestine,  habits  of  the  chaffinch  in, 

248. 
Pallas,  on  the  perfection  of  the  senses 
in  the  Mongolians,  34 ;  on  the 
want  of  connexion  between  cli- 
mate and  the  colour  of  the  skin, 
192  ;  on  the  polygamous  habits  of 
Antilope  Saiga,  217  ;  on  the  lighter 
colour  of  horses  and  cattle  in 
winter  in  Siberia,  229  ;  on  the 
tusks  of  the  musk-deer,  513,  514; 
on  the  odoriferous  glands  of  mam- 
mals, 529 ;  on  the  odoriferous 
glands  of  the  musk-deer,  530  ;  on 
winter  changes  of  colour  in  mam- 
mals, 542  ;  on  the  ideal  of  female 
beauty  in  North  China,  578. 
Palmaris  accessorius,  muscle  variations 

of  the,  27. 
Pampas,  horses  of  the,  181. 
Pangenesis,  hypothesis  of,  228,  231. 
Panniculus  carnosus,  13. 
Pansch,  on  the  brain  of  a  foetal  Cebus 

apella,  205. 
Papilio,  proportion  of  the  sexes  in 
North  American  species  of,  250 ; 
sexual  differences  of  colouring  in 
species  of,  309  ;  coloration  of  the 
wings  in  species  of,  312. 

ascanius,  309. 

Sesostris   and    Childrcna:,  varia- 
bility of,  319. 

Turnus,  250. 

Papilionidse,  variability  in  the,  319. 
Papuans,  line  of  separation  between 
the,  and   the  Malays,  169  ;  beards 
of  the,  560  ;  hair  of,  575. 
and  Malays,  contrast  in  charac- 
ters of,  168. 
Paradise,  Birds  of,  405,  462  ;  supposed 
by  Lesson   to  be  polygamous,  219 
rattling    of  their  quills  by,    375 
racket -shaped    feathers    in,    384 
sexual  differences  in  colour  of,  385 
decomposed  feathers  in,  385,  403 
display  of  plumage    by   the   male, 
395. 
Paradisea  apala,  barbless  feathers  in 
the  tail  of,  385  ;  plumage  of,  385; 
and  P.  papiiana,  385  ;  divergence 
of  the  females  of,  470  ;  increase   of 
beauty  with  age,  485. 
Paraguay,   Indians  of,  eradication  of 
eyebrows  and  eyelashes  by,  580. 


PARAKEET. 


INDEX. 


PERCH. 


667 


Parakeet,  Australian,  variation  in  the 
colour  of  the  thighs  of  a  male, 
423. 

Parallelism  of  development  of  species 
and  languages,  90. 

Parasites  on  man  and  animals,  7,  8 ; 
as  evidence  of  specific  identity  or 
distinctness,  169  ;  immunity  from, 
correlated  with  colour,  193. 

Parental  feeling  in  earwigs,  star- 
fishes, and  spiders,  106  ;  afl'ection, 
partly  a  result  of  natural  selection, 
105. 

Parents,  age  of,  influence  upon  sex  of 
offspring,  245. 

Parinaj,  sexual  difference  of  colour  in, 
458. 

Park,  Mungo,  negro-women  teaching 
their  children  to  love  the  truth, 
118;  his  treatment  by  the  negro- 
women,  118,  562  ;  ou  negi'o  opinions 
of  the  appearance  of  white  men, 
579. 

Parker,  Mr.,  no  bird  or  reptile  in  line 
of  mammalian  descent,  158. 

Parrot,  racket-shaped  feathers  in  the 
tail  of  a,  384 ;  instance  of  benevo- 
lence in  a,  411. 

Parrots,  change  of  colour  in,  60 ; 
imitative  faculties  of,  73 ;  living 
in  triplets,  409  ;  affection  of,  410  ; 
colours  and  nidification  of  the,  455, 
457,  458 ;  immature  plumage  of 
the,  467 ;  colours  of,  489 ;  sexual 
differences  of  colour  in,  494  ;  musi- 
cal powers  of,  570. 

Parthenogenesis  in  the  Tenthredinje, 
254  ;  in  Cynipida?,  254  ;  in  Crusta- 
cea, 255, 

I'artridge,  monogamous,  219  ;  propor- 
tion of  the  sexes  in  the,  248  ;  fe- 
male, 471. 

" dances,"  380,  405. 

Partridges,  living  in  triplets,  409  ; 
spring  coveys  of  male,  409  ;  distin- 
guishing persons,  412. 

Parus  ojeruleus,  458. 

Passer,  sexes  and  young  of,  483. 

hrachydactylus,  483. 

domesticus,  455,  483. 

montanus,  455,  483. 

Patagonians,  self-sacrifice  by.  111; 
marriages  of,  598. 

Patterson,  Mr.,  on  the  Agrionidae, 
250. 


Patteson,  Bishop,  decrease  of  Melaue- 

sians,  185. 
Paulistas  of  Brazil,  173. 
Pavo  cristatus,  236,  430. 
miidcus,  236,  430 ;  possession  of 

spurs  by  the  female,  364,  450. 

nigripennis,  419. 

Payaguas  Indians,  thin  legs  and  thick 

arms  of  the,  32. 
Payan,  Mr.,  on  the  proportion  of  the 

sexes  in  sheep,  246. 
Peacock,    polygamous,    219;    sexual 

characters   of,   236 ;    i)Ugnacity   of 

the,  364  ;  rattling  of  the  quills  by, 

375  ;  elongated  tail-coverts  of  the, 

384,  402  ;  love  of  display  of  the, 

394, 43 1 ;  ocellated  spots  of  the,  430 ; 

inconvenience  of  long  tail  of  the,  to 

the  female,  444,  451,  452;  continued 

increase  of  beauty  of  the,  485. 

butterfly,  312. 

Peafowl,  preference   of  females  for  .1 

particular  male,  418  ;  first  advances 

made  by  the  female,  419. 
Pediculi  of  domestic  animals  and  man, 

169. 
Pedigree  of  man,  165. 
Pedionomus  torquatus,  sexes  of,  473. 
Peel,  J.,  on  horned  sheep,  505. 
Peewit,  wing-tubercles  of  the   male, 

366. 
Pelagic  animals,  transparency  of,  261. 
Pelecanus erythrorh}jnchus,  horny  crest 

on  the  beak  of  the  male,  during  the 

breeding  season,  390. 
onocrotalus,  spring  plumage   of, 

393. 
Pelelcf!,  an  African  ornament,  576. 
Pelican,  blind,  fed  by  his  companions, 

102  ;  young,  guided   by   old   birds, 

102  ;  pugnacity  of  the  male,  362. 
Pelicans,  fishing  in  concert,  101. 
Pelobius   Hcnnanni,    stridulation    of, 

303,  304. 
Pelvis,  alteration  of,  to  suit  the  erect 

attitude  of  man,  53  ;  differences  of 

the,  in  the  sexes  in  man,  557. 
Penelope  nigra,  sound  produced  by  the 

male,  377. 
Pennant,  on  the  battles  of  seals,  500; 

on  the  bladder-nose  seal,  528. 
Penthe,  antennal  cushions  of  the  male, 

276. 
Perch,    brightness    of    male,    during 

breeding  season,  340. 


662> 


PEREGRINE. 


INDEX. 


Peregrine  falcon,  new  mate  found  by, 

408. 
Period  of  variability,  relation   of,  to 

sexual  selection,  240. 
Periodicity,  vital.  Dr.  Laycock,  on,  8. 
Periods,  lunar,  followed  by  functions 

in  man  and  animals,  8,  164, 
of  life,    inheritance    at    corre- 

spondmg,  228,  232, 
Perisoreus  canadensis,  young  of,  481. 
Feritrichia,  difference  of  colour  in  the 

sexes  of  a  species  of,  294. 
Periwinkle,  262. 
Pernis  cristata,  424. 
Perrier,  M.,  on  sexual  selection,  210  ; 

on  bees,  292. 
Perseverance,  a  characteristic  of  man, 

564. 
Persians,  said  to  be  improved  by  in- 
termixture with  Georgians  and  Cir- 
cassians, 586. 
Personnat,  M.,  on  Bomhyx   Yamamai, 

251. 
Peruvians,   civilisation    of    the,    not 

foreign,  145. 
Petrels,  colours  of,  493. 
Petrocinda  cyanea,  young  of,  487. 
Peti'ocossyphus,  461. 
Pctronia,  483. 
Pfeiffer  Ida,  on  Javan  ideas  of  beautv, 

580. 
Phacochcerus    oethiopicus,    tusks    and 

pads  of,  519. 
Phalanger,  Vulpine,  black  varieties  of 

the,  539. 
Phalaropus  fulicarius,  476. 

hyperboreus,  476. 

Phanxus,  298. 

carnifex,  variation  of  the  horns 

of  the  male,  297. 

faunus,   sexual    differences    of, 

296. 

lancifer,  295. 

Phaseolarctus  cincreus,  taste  for  rum 

and  tobacco,  7. 
Phasgonura   viridissiina,    stridulation 

of,"  284,  285. 
Phasianus  Scemmerringii,  446. 
versicolor,  396. 

Wallichii,  400,  472. 

Pheasant,  polygamous,  219  ;  and  black 

grouse,  hybrids  of,  414;  production 
of  hybrids  with  the  common  fowl, 
420  immature  plumage  of  the, 
460. 


Pheasant,  Amherst,  display  of,  396. 

,  Argus,    334,    462  ;     display    of 

plumage  by  the  male,  398  ;  ocel- 
lated  spots  of  the  428,  434  ;  grada- 
tion of  characters  in  the,  434. 

,  Blood-,  364. 

,  Cheer,  400,  472. 

,  Eared,  235,  400,  472  ;    length 

of  the  tail  in  the,  452  ;  sexes  alike 
in  the,  460. 

,    Golden,    display    of    plumage 

by  the  male,  396  ;  age  of  mature 
plumage  in  the,  483  ;  sex  of  young, 
ascertained  by  pulling  out  head- 
feathers,  484. 

,  Kalij,  drumming  of  the  male, 

375. 

,  Pieeve's,  length  of  the  tail  in, 

452. 

,  Silver,  triumphant  male,  de- 
posed on  account  of  spoiled  plu- 
mage, 419  ;  sexual  coloration  of  the, 
492. 

,  Scemmerring's,  445,452. 

,  Tragopan,     383 ;      display     of 

plumage  by  the  male,  397  ;  mark- 
ings of  the  sexes  of  the,  428. 

Pheasants,  period  of  acquisition  of  male 
characters  in  the  family  of  the, 
235 ;  proportion  of  sexes  in  chicks 
of,  247  ;  length  of  the  tail  in,  445, 
451,  452. 

Philters,  worn  by  Avomen,  577. 

Phoca  grccnlandica,  sexual  difference 
in  the  coloration  of,  535. 

Phccnicura  ruticilla,  408. 

Phosphorescence  of  insects,  277. 

Phryganidai,  copulation  of  distinct 
species  of,  275. 

Phryniscus  nigricans,  349. 

Physical  inferioi'ity,  supposed,  of  man, 
64. 

Pickering,  on  the  number  of 
of  man,  174. 

Picton,  J.  A.,on  the  soul  of  man,  613. 

Picus  auratus,  362. 

major,  402. 

Pieris,  312  ,319. 

Pigeon,  female,  deserting  a  weakened 
mate,  214 ;  carrier,  late  develop- 
ment of  the  wattle  in,  238 : 
pouter,  late  development  of  the 
crop  in,  238;  domestic,  breeds  and 
sub-breeds  of,  460. 

Pigeons,  nestling,  fed  by  the  secretion 


jpecies 


INDEX. 


I'OLVGAMY. 


669 


of  the  crop  of  both  parents,  163 ; 
changes  of  plumage  in,  229  ;  trans- 
mission of  sexual  peculiarities  in, 
230 ;  Belgian,  with  black-streaked 
males,  231,  238,  446  ;  changing 
colour  after  several  moultings,  238  ; 
numerical  proportion  of  the  sexes 
in,  247  ;  cooing  of,  374  ;  variations 
in  plumage  of,  385 ;  display  of 
plumage  by  male,  402 ;  local  me- 
mory of,  411  ;  antipathy  of  female, 
to  certain  males,  418  ;  pairing  of, 
418;  profligate  male  and  female, 
418 ;  wing-bars  and  tail-feathers 
of,  427  ;  suppositious  breed  of,  445  ; 
pouter  and  carrier,  peculiarities  of, 
predominant  in  males,  447 ;  nidi- 
lication  of,  453 ;  Australian,  458  j 
immature  plumage  of  the,  467. 

Pigs,  origin  of  the  improved  breeds 
of,  177  ;  numerical  proportion  of 
the  sexes  in,  247  ;  stripes  of  young, 
464,  546  ;  tusks  of  «iiocene,  521  ; 
sexual  preference  shewn  by,  525. 

Pike,  American,  brilliant  colours  of 
the  male,  during  the  breeding 
season,  340. 

,  reasoning  powers  of,  75  ;  male, 

devoured  by  females,  249. 

Pike,  L.  0.,  on  the  psychical  elements 
of  religion,  95. 

Pimelia  striata,  sounds  produced  by 
the  female,  306. 

Pinel,  hairiness  in  idiots,  36. 

Pintail,  drake,  plumage  of,  393 ; 
pairing  with  a  wild  duck,  415. 

■ Duck,  pairing  with  a  wigeon, 

414. 

Pipe-fish,  filamentous,  344 ;  mar- 
supial receptacles  of  the  male,  346. 

Pipits,  moulting  of  the,  392. 

Pipra,  modified  secondary  wing- 
feathers  of  male,  378. 

deliciosa,  378,  379. 

Pirates  stridulus,  stridulation  of,  281. 

Pitcairn  island,  half-breeds  on,  190. 

Pithccia  leucocephala,  sexual  dilfer- 
ences  of  colour  in,  537. 

Satanas,   beard    of,   531,   532 ; 

resemblance  of,  to  a  negro,  604. 

Pits,  suborbital,  of  Ruminants,  529. 

Pittidae,  nidification  of,  453. 

Placentata,  157. 

Plagiostomous  fishes,  331. 

Plain-wanderer,  Australian,  473. 


Planaria;^  bright  colours  of  some, 
260. 

Plantain-eaters,  colours  and  nidi- 
fication of  the,  455  ;  both  sexes  of, 
equally  brilliant,  460. 

Plants,  cultivated,  more  fertile  than 
wild,  45  ;  Niigeli,  on  natural  selec- 
tion in,  61 ;  male  flowers  of,  mature 
before  the  female,  212  ;  phenomena 
•of  fertilisation  in  222. 

Platalca,  374;  change  of  plumage  in, 
461. 

Platijblemnus,  289. 

Platycercus,  young  of,  481. 

Platyphyllum  concamun,  283,  286. 

Platyrrhine  monkeys,  153. 

Platijsma  myoides,  13. 

Plecostomus,  head-tentacles  of  the 
males  of  a  species  of,  338. 

barbatus,  peculiar  beard  of  the 

male,  338. 

Plectropterus  gambensis,  spurred  wings 
of,  364. 

Ploceus,  370,  375,  405. 

Plovers,  Aving-spurs  of,  365  ;  double 
moult  in,  392. 

Plumage,  changes  of,  inheritance  of, 
by  fowls,  22J ;  tendency  to  ana- 
logous variation  in,  385 ;  display 
of,  by  male  birds,  394,  402  ;  changes 
of,  in  relation  to  season,  462  ;  im-r 
mature,  of  birds,  464,  465  ;  colour 
of,  in  relation  to  protection,  489. 

Plumes  on  the  head  in  birds,  differ- 
ence of,  in  the  sexes,  451. 

Pneumora,  structure  of,  286. 

Pudica,  sexual  difference  in  the  colour 
of  the  irides  of,  425. 

Poeppig,  on  the  contact  of  civilised 
and  savage  races,  183. 

Poison,  avoidance  of,  by  animals,  80. 

Poisonous  fruits  and  herbs  avoided  by 
animals,  ijG, 

Poisons,  immunity  from,  correlated 
with  colour,  193. 

Polish  fowls,  origin  of  the  crest  in, 
231. 

Pollen  and  van  Dam,  on  the  colours 
of,  Lemur  macaco,  537. 

Polyandry,  593 ;  in  certain  Cypri- 
nidaj,  249  ;  among  the  Llateridae, 
253. 

Polydactylism  in  man,  37. 

Polygamy,  iuHuence  of,  upon  sexual 
selection,    216;    superinduced    by 


6/0 


POLYGENISTS. 


INDEX. 


PUFF-BIROS. 


domestication,  220 ;  supposed  in- 
crease of  female  births  by,  245  ;  in 
the  stickleback,  332. 

Polygenists,  176. 

Polynesia,  prevalence  of  infanticide 
in,  592. 

Polynesians,  wide  geographical  range 
of  29  ;  difference  of  stature  among 
the,  31 ;  crosses  of,  173;  variability 
of,  174 ;  heterogeneity  of  the,  192  ; 
aversion  of,  to  hairs  on  the  face, 
581. 

Polyplectron,  number  of  spurs  in,  364; 
display  of  plumage  by  the  male, 
396 ;  gradation  of  characters  in, 
431 ;  female  of,  471. 

chinquis,  397,  432. 

Hard'.cickii,  432. 

malaccense.  432,  433. 

Napoleonis,  432,  433. 

Polyzoa,  262. 

Pontoporeia  affinis,  266. 

Porcupine,  mute,  except  in  the  rut- 
ting season,  526. 

Pores,  excretory,  numerical  relation 
of,  to  the  hairs  in  sheep,  198. 

Pnrpitce,  bright  colours  of  some,  260. 

Portax  picta,  dorsal  crest  and  throat- 
tuft  of,  530  ;  sexual  differences  of 
colour  in,  535,  543. 

Portunus  piiber,  pugnacity  of,  269. 

Potamochcerus  penicillatus,  tusks  and 
facial  knobs  of  the,  520. 

Pouchet,  G.,  the  relation  of  instinct 
to  intelligence,  67 ;  on  the  in- 
stincts of  ants,  147  ;  on  the  caves 
of  Abou-Simbel,  168 ;  on  the  im- 
munity of  negroes  from  yellow 
fever,  193;  change  of  colour  in 
fishes,  344. 

Pouter  pigeon,  late  development  of 
the  large  crop  in,  238. 

Powell,  Dr.,  on  stridulation,  281. 

Power,  Dr.,  on  the  different  colours 
of  the  sexes  in  a  species  of  Squilla, 
271. 

Powys,  Mr.,  on  the  habits  of  the 
chaffinch  in  Corfu,  248. 

Pre-eminence  of  man,  48 

Preference  for  males  by  female  birds, 
414,  420;  shewn  by  mammals,  in 
pairing,  522. 

Prehensile  organs,  209. 

Prcshi/tis  eniellus,  fighting  of  the 
male,  562. 


Preyer,  Dr.,  on  function  of  shell  of 
ear,  14;  on  supernumerary  mam- 
mas in  women,  37. 
Prichard,  on  the  difference  of  stature 
among  the  Polynesians,  31  ;  on  the 
connection  between  the  breadth  of 
the  skull  in  the  Mongolians  and  the 
perfection  of  their  senses,  34 ;  on 
the  capacity  of  British  skulls  of 
different  ages,  55  ;  on  the  flattened 
heads  of  the  Colombian  savages, 
575  ;  on  Siamese  notions  of  beauty, 
578  ;  on  the  beardlessness  of  the 
Siamese,  581  ;  on  the  deformation 
of  the  head  among  American  tribes 
and  the  natives  of  Arakhan,  583. 
Primary  sexual  organs,  207. 
Primates,  149,  205  ;  sexual  differences 

of  colour  in,  537. 
Primogeniture,  evils  of,  135. 
Prionidse,    difference  of  the    sexes  in 

colour,  294. 
Proctotretus  mttltimaculatus,  358. 

tenuis,  sexual  difference  in    the 

colour  of,  358. 
Profligacy,  137. 

Progenitors,  early,  of  man,  160. 
Progress,  not  the  normal  rule  in  hu- 
man society,  133  ;  elements  of,  141. 
Prong-horn  antelope,  horns  of,  234. 
Proportions,  difference  of,  in  distinct 

races,  167. 
Protective    colouring    in    butterflies, 
312;  in  lizards,  358;  in  birds,  473, 
489  ;  in  mammals,  542. 

nature  of  the  dull  colouring  of 

female  Lepidoptera,  321,  322,  324. 

resemblances  in  fishes,  344. 

Protozoa,  absence  of  secondary  sexual 

characters  in,  260. 
Prunei--Bey,  on  the  occurrence  of  the 
supra-condyloid     foramen     in    the 
humerus  of  man,  22  ;  on  the  colour 
of  negro  infants,  S57. 
Prussia,  numerical  proportion  of  male 

and  female  births  in,  243. 
Psocus,  proportions   of  the   sexes   in, 

254. 
Ptarmigan,  monogamous,  219 ;  sum- 
mer  and  winter  plumage    of   the,  . 
390,  392  ;    nuptial  assemblages  of, 
406  ;  triple  moult  of  the,  462 ;  pro- 
tective coloration  of,  473. 
Puff-birds,  colours  and  nidification  of 
the.  455. 


PUONACITV. 


INDEX. 


671 


Pugnacity  of  fine-plumaged  male 
birds,  400. 

Pumas,  stripes  of  young,  464. 

Puppies  learning  from  cats  to  clean 
their  faces,  73. 

Pycnonotus  hccmorrhous,  pugnacity  of 
the  male,  360 ;  display  of  under 
tail  coverts  by  the  male,  402. 

Pyranga  ccstiva,  male  aiding  in  incu- 
bation, 453. 

Pijrodes,  difference  of  the  sexes  in 
colour,  294. 

Q. 

Quadrumana,  hands  of,  50  ;  differences 
between  man  and  the,  149  ;  sexual 
differences  of  colour  in,  537  ;  orna- 
mental characters  of,  549  ;  analogy 
of  sexual  differences  of,  with  those 
of  man,  558 ;  fighting  of  males  for 
the  females,  562 ;  monogamous 
habits  of,  590  ;  beards  of  the,  602. 
Quain,   R.,   on  the    variation    of  the 

muscles  in  man,  26. 
Quatrefages,  A.  de,  on  the  occur- 
rence of  a  rudimentary  tail  in  man, 
22  ;  on  variability,  30 ;  on  the 
moral  sense  as  a  distinction  between 
man  and  animals,  97  ;  civilised  men 
stronger  than  savages,  136  ;  on  the 
fertility  of  Australian  women  with 
white  men,  170;  on  the  Paulistas 
of  Brazil,  173  ;  on  the  evolution  of 
the  breeds  of  cattle,  177 ;  on  the 
Jews,  193  ;  on  the  liability  t.f 
negroes  to  tropical  fevers  after 
residence  in  a  cold  climate,  194; 
on  the  difference  between  field- 
and  house-slaves,  196;  on  the  in- 
fluence of  climate  on  colour,  196  ; 
colours  of  annelids,  265;  on  the 
Ainos,  560;  on  the  women  of 
San  Giuliano,  586. 
Querquedula  acuta,  414. 
Qu<^telet,  proportion  of  sexes  in  man, 
343;  relative  size  in  man  and 
woman,  244. 
Quichua  Indians,  34;  local  variation 
of  colour  in  the,  196;  no  grey 
hair  among  the,  559  ;  hairlessness 
of  the,  561  ;  long  hair  of  the,  580. 
Quiscalus  major,  225  ;  proportions  of 
the  sexes  of,  in  Florida  and  Hondu- 
ras, 248. 


B. 

Rabbit,  white  tail  of  the,  542. 
Rabbits,  domestic,  elongation   of  the 
skull  in,   56  ;    modification  of  the 
skull,    in    by   the    lopping   of   the 
ear,    56 ;    danger-signals    of,    100  ; 
numerical  proportion  of  the  sexes 
in,  247. 
Races,  distinctive  characters  of,  167, 
168  ;  or  species  of  man,  168;  crossed, 
fertility    or    sterility    of,  170;     of 
man,    variability    of  the,    174;  of 
man,    resemblance    of,    in   mental 
characters,     178 ;      formation    01, 
181;  of  man,   extinction  of,   181; 
effects   of  the  crossing  of,  192  ;  of 
man,    formation    of  the,   192;    of 
man,  children  of  the,  557  ;  beard- 
less, aversion  of,  to  hairs   on   the 
foce,  580. 
Raflies,  Sir  S.,  on  the  banteng,  536. 
Rafts,  use  of,  48,  180. 
Rage,  manifested  by  animals,  69, 
liaia  batis,  teeth  of,  335. 

clavata,  female    spined  on  the 

back,  331 ;  sexual  difference  in  the 
teeth  of,  334. 

maculata,  teeth  of,  335. 

Rails,  spur-winged,  364. 
Ram,  mode   of  fighting  of  the  508  ; 
African,    mane    of  an,    532  ;     fat- 
tailed,  532. 
Rameses  II.,  features  of,  168. 
liamsay,     Mr.,     on     the     Australian 
musk-duck,   359;    on    the   regent- 
bird,  413  ;    on    the    incubation    of 
Menura  superba,  451. 
Ihvia  csculcnta,  vocal  sacs  of,  350. 
Rat,  common,   general  dispersion   of, 
a  consequence  of  superior  cunning, 
80 ;  supplantation   of   the    native, 
in   New  Zealand,  by  the   European 
rat,  191  ;  common,  said  to  be  poly- 
gamous,   218;     numerical   propor- 
tion of  the  sexes  in,  247. 
Rats,  enticed  by  essential  oils,  530. 
Rationality  of  birds,  410. 
Rattle-snakes,  difference  of  the  sexes 
in  the,  351  ;  rattles  as  a  call,  353, 
Raven,    vocal    organs    of    the    370  ; 
stealing  bright  objects,  413;  pied, 
of  the  Feroe  Islands,  424. 
Rays,  prehensile  organs  of  male,  331. 


6/: 


RAZOR-BILL. 


INDEX. 


RICHARDSON. 


Razor-bill,  young  of  the,  486. 

Reade,  Winwood,  suicide  among 
savages  in  Africa,  117  ;  mulattoes 
not  prolific,  171  ;  effect  of  castra- 
tion of  horned  sheep,  506  ;  on  the 
Guinea  sheep,  235  ;  on  the  occur- 
rence of  a  mane  in  an  African 
ram,  533 ;  on  the  negroes'  appre- 
ciation of  the  beauty  of  their 
women,  577  ;  on  the  admiration  of 
negroes  for  a  black  skin,  579;  on 
the  idea  of  beauty  among  negroes, 
582 ;  on  the  Jollofs,  587  ;  on  the 
marriage-customs  of  the  negroes, 
599. 

Reason,  in  animals,  75. 

Redstart,  American,  breeding  in  im- 
mature plumage,  484-. 

Redstarts,  new  mates  found  by,  408. 

Reduvidae,  stridulation  of,  28i. 

Reed-bunting,  head-feathers  of  the 
male,  402  ;  attacked  by  a  bull-finch, 
412. 

Reefs,  fishes  frequenting,  343. 

Reeks,  H.,  retention  of  horns  by 
breeding  deer,  503  ;  cow  rejected 
by  a  bull,  525 ;  destruction  of  pie- 
bald rabbits  by  cats,  542. 

Regeneration,  partial,  of  lost  parts  in 
man,  8. 

Regent-bird,  413. 

Reindeer,  horns  of  the,  233  ;  battles 
of,  501  ;  horns  of  the  female,  503; 
antlers  of,  with  numerous  points, 
510;  winter  change  of  the,  542; 
sexual  preferences  shewn  by,  552. 

Relationship,  terras  of,  590. 

Religion,  defieiency  of,  among  cer- 
tain races,  93  ;  psychical  elements 
of,  95. 

Remorse,  114;  deficiency  of,  among 
savages,  131. 

Rengger,  on  the  diseases  of  Cchus 
Azarce,  7  ;  on  the  diversity  of  the 
mental  faculties  of  monkeys,  27 ; 
on  the  Payaguas  Indians,  32  ;  on 
the  inferiority  of  Europeans  to 
savages  in  their  senses,  33';  re- 
venge taken  by  monkeys,  69 ;  on 
maternal  affection  in  a  Cebus,  70 ; 
on  the  reasoning  powers  of  Ameri- 
can monkeys,  77 ;  on  the  use  of 
stones  by  monkeys  for  cracking 
hard  nuts,  81 ;  on  the  sounds  ut- 
tered by  Cebus  A  zara,  84 ;  on  the 


signal-cries  of  monkeys,  87  ;  on  the 
pol3'^gamous  habits  of  Mycetes  ca- 
raya,  217  ;  on  the  voice  of  the  how- 
ling monkeys,  527  ;  on  the  odour 
of  Cervus  campestris,  529 ;  on  the 
beards  of  Mycetes  caraya  and 
Fithecia  Satanas,  531 ;  on  the 
colours  of  Felis  mitis,  534 ;  on  the 
colours  of  Cervus  paludosus,  536  ; 
on  sexual  differences  of  colour  m 
Jlycetes,  537  ;  on  the  colour  of  the 
infant  Guaranys,  558  ;  on  the  early 
maturity  of  the  female  of  Cebus 
azarce,  558  ;  on  the  beards  of  tne 
Guaranys,  561 ;  on  the  emotional 
notes  employed  by  monkeys,  572; 
on  American  polygamous  monkeys, 
590. 

Representative  species,  of  birds,  468. 

Reproduction,  unity  of  phenomena 
of,  throughout  the  mammalia,  8  ; 
period  of,  in  birds,  484. 

Reproductive  system,  rudimentary 
structures  ki  the,  23  ;  accessory 
parts  of,  161. 

Reptiles,  350. 

and  birds,  alliance  of,  165. 

Resemblances,  small,  between  man 
and  the  apes,  150. 

Retrievers,  exercise  of  reasoning 
faculties  by,  78. 

Revenge,  manifested  by  animals,  69. 

Reversion,  36  ;  perhaps  the  cause  o'' 
some  bad  dispositions,  137. 

Jihagium,  diflerence  of  colour  in  the 
sexes  of  a  species  of,  294. 

Rhamphastos  carinatus,  492. 

Fhea  darwmii,  479. 

Rhinoceros,  nakedness  of,  57 ;  horns 
of,  505  ;  horns  of,  used  defensively, 
518;  attacking  white  or  grey 
horses,  540. 

Fhynchcea,  sexes  and  young  of,  47G. 

austral  is,  476. 

bengalensis,  476. 

capensis,  477. 

Rhythm,  perception  of,  bv  animals, 
569. 

Richard,  M.,  on  rudimentary  muscles 
in  man,  12. 

Richardson,  Sir  J.,  on  the  pairing  of 
Tetrao  umbellus,  366 ;  on  Tetrao 
urophasianus,  372 ;  on  the  drum- 
ming of  grouse,  376  ;  on  the  dances 
of    Tetrao  phasiaiwltuSf    381  ;     on 


RICHARDSON. 


INDEX. 


^7Z 


assemblages  of  grouse,  405  ;  on  the 
battles  of  male  deer,  501 ;  on  the 
reindeer,  503  ;  on  the  horns  of  the 
musk-ox,  505;  on  antlers  of  the 
reindeer  with  numerous  points, 
510  ;   on  the  moose,  515. 

Richardson,  on  the  Scotch  deer- 
hound,  516. 

Richter,  Jean  Paul,  on  imagination,  74. 

Riedel,  on  profligate  female  pigeons, 
218. 

Riley,  Mr.,  on  mimicry  in  butterflies, 
324 ;  birds'  disgust  at  taste  of 
certain  caterpillars,  326. 

King-ouzel,  colours  and  nidification 
of  the,  455. 

Ripa,  Father,  on  the  difficulty  of  dis- 
tinguishing the  races  of  the  Chinese, 
167. 

Rivalry,  in  singing,  between  male 
birds,  369. 

River-hog,  African,  tusks  and  knobs 
of  the,  520. 

Rivers,  analogy  of,  to  islands,  159. 

Roach,  brightness  of  male  during 
breeding-season,  340. 

Robbery,  of  strangers,  considered 
honourable,  117, 

Robertson,  Mr.,  remarks  on  the  de- 
velopment of  the  horns  in  the  roe- 
buck and  red-deer,  234. 

Robin,  pugnacity  of  the  male,  360  ; 
autumn  song  of  the,  370  ;  female 
singing  of  the,  370 ;  attacking 
other  birds  with  red  in  their  plu- 
mage, 412;   young  of  the,  480. 

Robinet,  on  the  difference  of  size  of 
the  male  and  female  cocoons  of  the 
silk-moth,  278. 

Rodents,  uterus  in  the,  38 ;  absence 
of  secondary  sexual  characters  in, 
218;  sexual  differences  in  the 
colours  of,  534. 

Roe,  winter  change  of  the,  542. 

Rohlfs,  Dr.,  Caucasian  features  in 
negro,  167  ;  fertility  of  mixed 
races  in  Sahara,  171;  colours  of 
birds  in  Sahara,  490  ;  ideas  of  beauty 
amongst  the  Bornuans,  582. 

Rolle,  F.,  on  the  origin  of  man,  3  ; 
on  a  change  in  German  fiimilies 
settled  in  Georgia,  196. 

Roller,  harsh  cry  of,  371. 

Romans,  ancient,  gladiatorial  exhi- 
bitions of  the,  123. 


Ronjou,  M.  A.,  coincidence  of  arrested 

development    with    polydactylism 

37. 
Rook,  voice  of  the,  375. 
Rossler,   Dr.,  on  the  resemblance    of 

the  lower  surface   of  butterflies  to 

the  bark  of  trees,  311. 
Rostrum,    sexual     difference    in    the 

length  of,  in  some  weevils,  208. 
Royer,    Madlle.,     mammals      giving 

suck,  163. 
Rudim«ntary  organs,  11  ;    origin  of, 

24. 
Rudiments,  presence  of,  in  languages, 

90. 
Rudolphi,  on   the  want   of  connexion 

between  climate  and  the  colour  of 

the  skin,  192. 
Ruff,    supposed     to   be    polygamous, 

219;    proportion    of  the    sexes   in 

the,  248;  pugnacity  of  the,    361; 

double  moult  in,  390,392;  duration 

of  dances  of,  405  ;  attraction  of  the, 

to  bright  objects,  413. 
Ruminants,    male,   disappearance     of 

canine  teeth  in,  53,  562  ;  generally 

polygamous,   217  ;    suborbital  pits 

of,  529  ;  sexual  differences  of  colour 

in,  535. 
Rupicola  crocea,  display   of  plumage 

by  the  male,  395. 
Rtippell,  on  canine  teeth    in  deer  and 

antelopes,  514. 
Russia,  numerical  proportion  of  male 

and  female  births  in,  215,  243. 
Ruticilla,  462. 

RUtimeyer,  Prof.,  on  •  the   physiogno- 
my of  the  apes,  54  ;    on  the  sexual 

differences  of  monkeys,  561. 
Rutlandshire,    numerical    proportion 

of  male  and  female  births  in,  242. 


S. 

Sachs,  Prof.,  on  the  behaviour  of  the 
male  and  female  elements  in  fertili- 
sation, 222. 

Sacrifices,  human,  144. 

Sagittal  crest  in  male  apes  and 
Australians,  558. 

Sahara,  fertility  of  mixed  races  in, 
171  ;  birds  of  the,  456  ;  animal  m- 
habitants  of  the,  489. 

Sailors,  growth  of,  delayed  by  con- 
ditions of  life,  31 ;  long-sighted,  33. 


6/4 


INDEX. 


SCIIOMBURGE. 


Sailors  and  soldiers,  diflference  in  the 
proportions  of,  32. 

St.  John,  Mr.,  on  the  attachment  of 
mated  birds,  410. 

St.  Kilda,  beards  of  the  inhabitants 
of,  560. 

Salmo  eriox,  and  8.  umhla,  colouring 
of  the  male,  during  the  breeding 
season,  340. 

lyr_aodon,  333. 

salar^  333. 

Salmon  leaping  out  of  fresh  water, 
107  ;  male,  ready  to  breed  before 
the  female,  212;  proportion  of 
the  sexes  in,  249  ;  male,  pugna- 
city of  the,  332  ;  male,  characters 
of,  during  the  breeding  season,  332, 
340  ;  spawning  of  the,  344  ;  breed- 
ing of  immature  male,  485. 

Salvin,  0.,  inheritance  of  mutilated 
feathers,  60,  384,  603;  on  the 
Humming-birds,  219,  454;  on  the 
numerical  proportion  of  the  sexes 
in  Humming-birds,  248,  487  ;  on 
Ckamoepetes  and  Penelope,  377 ; 
on  Selasphorus  platycercus,  378 ; 
Pipra  deliciosa,  378  ;  on  C/iasmo- 
rhynchus,  389. 

Samoa  Islands,  beardlessness  of  the 
natives  of,  560,  581. 

Sand-skipper,  270. 

Sandwich  Islands,  variation  in  the 
skulls  of  the  natives  of  the,  26; 
decrease  of  native  population,  186  ; 
papulation  of,  257  ;  superiority  of 
the  nobles  in  the,  587. 

Islanders,  lice  of,  170. 

San-Giuliano,  women  of,  586. 

Santali,  recent  rapid  increase  of  the, 
45;  Mr.  Hunter  on  the,  192. 

Saphirina,  characters  of  the  males  of, 
271. 

Sarhidiornis  melanonotus,  characters 
of  the  young,  465. 

Sars,  0.,  on  Pontoporcia  affinis, 
266. 

Sacuniia  carpini,  attraction  of  males 
by  the  female,  252. 

/o,  difference  of  coloration  in  the 

sexes  of,  316. 

Saturniidce,  coloration  of  the,  314, 
315. 

Savage,  Dr.,  on  the  fighting  of  the 
male  gorillas,  562  ;  on  the  habits 
of  the  gorilla,  591. 


Savage  and  Wyman  on  the  polyga- 
mous habits  of  the  gorilla,  217. 

Savages,  uniformity  of,  exaggerated, 
28  ;  long-sighted,  33  ;  rate  of  in- 
crease among,  usually  small,  45  ; 
retention  of  the  prehensile  power 
of  the  feet  by,  52  ;  imitative  facul- 
ties of,  87 ;  129 ;  causes  of  low 
morality  of,  119;  tribes  of,  sup- 
planting one  another,  128 ;  im- 
provements in  the  arts  among, 
144 ;  arts  of,  179 ;  fondness  of, 
for  rough  music,  380 ;  attention 
paid  by,  to  personal  appearance, 
574  ;  relation  of  the  sexes  among, 
591. 

Saviotti,  Dr.,  division  of  malar  bone, 
39. 

Saw-fly,  pugnacity  of  a  male,  291. 

Saw-flies,  proportions  of  the  sexes  in, 
254. 

Saxicola  ruhicola,  young  of,  487. 

Scalp,  motion  of  the,  13. 

Scent-glands  in  snakes,  352. 

Schaaffhausen,  Prof.,  on  the  develop- 
ment of  the  posterior  molars  in 
different  races  of  man,  20  ;  on  the 
jaw  from  La  Naulette,  40  ;  on  the 
correlation  between  muscularity 
and  prominent  supra-orbital  ridges, 
44 ;  on  the  mastoid  processes  of 
man,  53  ;  on  modifications  of  the 
cranial  bones,  56  ;  on  human  sacri- 
fices, 144  ;  on  the  probable  speedy 
extermination  of  the  anthropomor- 
phous apes,  156  ;  on  the  ancient 
inhabitants  of  Europe,  181 ;  on 
the  effects  of  use  and  disuse  of 
parts,  197  ;  on  the  superciliary 
ridge  in  man,  556  ;  on  the  ab- 
sence of  race-differences  in  the  in- 
fant skull  in  man,  557  ;  on  ugliness, 
584. 

Schaum,  H.,  on  the  elytra  of  Dytiscus 
and  Hydroporus,  276. 

Schelver,  on  dragon-flies,  290. 

Schiodte,  on  the  stridulation  of  Hete- 
rocerus,  302. 

Schlegel,  F.  von,  on  the  complexity  of 
the  languages  of  uncivilised  peo- 
ples, 91. 

,  Prof.,  on  Tanysiptfra,  468. 

Schleicher,  Prof.,  on  the  origin  of 
language,  87. 

Schomburgk,  Sir  R.,  on  the  pugnacity 


SCHOOLCUAl'T. 


INDEX. 


SEMNOPITHECUS. 


675 


of  the  male  musk-duck  of  Guiana, 
362  ;  on  the  courtship  of  Kupicola 
crocca,  395. 
Schoolcraft,  Mr.,  on  the  difficulty  of 

fashioning  stone  implements,  49. 
Schweinfurth,  complexion  of  negroes, 

556. 
Sciccna  aquila,  347. 
Sclater,  P.  L.,  on  modified  secondary 
wing-feathers  in  the  males  of  Fipra, 
378  ;     on    elongated    feathers    in 
nightjars,  384;   on   the  species  of 
Ciiasmorhynchus,  385  ;  on  the  plu- 
mage   of     Pelecanus     onocroiahis, 
393  ;  on  the  plantain-eaters,  460  ; 
on  the  sexes  and  young  of  Tadorna 
variegata,  479  ;  on  the   colours  of 
Lemur  macaco,  537  ;  on  the  stripes 
in  asses,  548. 
Scolecida,  absence  of  secondary  sexual 

characters  in,  260. 
Scolopax    frcnata,    tail-feathers     of, 
377. 

gallinago,  drumming  of,  376. 

javensis,  tail-feathers  of,  377. 

major,  assemblies  of,  405. 

Wilsonii,    sound    produced    by, 

377. 
Scolytus,  stridulation  of,  302. 
Scoter-duck,  black,  sexual  difference 
in   coloration  of  the,  491 ;  bright 
beak  of  male,  491. 
Scott,   Dr.,  on  idiots  smelling  their 
food,  36. 

,  J.,  on  the  colour  of  the  beard 

in  man,  558. 
Scrope,  on  the  pugnacity  of  the  male 
salmon,    332 ;    on    the    battles    of 
stags,  501. 
Scudder,    S.     H.,    imitation    of    the 
stridulation    of     the    Orthoptera, 
283 ;    on    the  stridulation  of  the 
Acridiidae,  286 ;  on  a  Devonian  in- 
sect, 289  ;  on  stridulation,  566. 
Sculpture,  expression  of  the  ideal  of 

beauty  by,  581. 
Sea-anemonies,  bright  colours  of,  260. 
Sea-bear,  polygamous,  218. 
Sea-elephant,  male,  structure  of  the 
nose  of  the,  527  ;  polygamous,  218. 
Sea-lion,  polygamous,  218. 
Seal,  bladder-nose,  528. 
Seals,    their    sentinels    generally    fe- 
males, 100;  evidence  furnished  by, 
on  classification,  150 ;  polygamous 


habits  of,  218;  battles  of  male, 
500 ;  canine  teeth  of  male,  502 ; 
sexual  dilferences,  515;  pairing  of, 
522  ;  sexual  peculiarities  of,  528  ; 
in  the  coloration  of,  534 ;  apprecia- 
tion of  music  by,  569. 
Sea-scorpion,    sexual    differences    in, 

337. 
Season,  changes  of  colour  in  birds,  in 
accordance  with  the,  390 ;  changes 
of  plumage  of  birds  in  relation  to, 
462. 
Seasons,  inheritance  at  corresponding, 

230. 
Sebituani,    African    chief,    trying   to 

alter  a  fashion,  575. 
Sebright  Bantam,  238. 
Secondary    sexual    characters,    207 ; 
relations    of    polygamy    to,    217  ; 
transmitted   through    both    sexes, 
226  ;  gradation  of,  in  birds,  430. 
Sedgwick,  W.,  on  hereditary  tendency 

to  produce  twin«,  45. 
Seemann,  Dr.,  on  the  different  appre- 
ciation of  music  by  different  peoples, 
570;  on  the  effects  of  music,  571. 
Seidlitz,  on  horns  of  reindeer,  506. 
Selaspliorus    platycercus,     acuminate 

first  primary  of  the  male,  378. 
Selby,  P.   J.,    on   the   habits    of  the 

black  and  red  grouse,  219. 
Selection,  double,  225. 

of  male  by  female  birds,  404, 

421. 
,  methodical,  of  Prussian  grena- 
diers, 29. 

,  sexual,  explanation  of,  209,  213, 

220  ;    influence   of,  on  the  colour- 
ing of  Lepidoptera,  321. 

,  sexual  and  natural,  contrasted, 

226. 
Self-command,    habit    of,    inherited, 

115  ;  estimation  of,  118. 
Self-consciousness,  in  animals,  83. 
Self-preservation,  instinct  of,  112. 
Self-sacrifice,  by  savages,   111;   esti- 
mation of,  118. 
Semilunar  fold,  17. 
Seinnopithecus,  154 ;  long  hair  on  the 
heads  of  species  of,  150,  604. 

chrysomelas,    sexual    differences 

of  colour  in,  537. 

comatus,  ornamental  hair  on  the 

head  of,  549. 
frontatus,  beard,  &c.,  of,  550. 


ey^ 


SEMNOPITHECUS. 


INDEX. 


Semnopithecus  nasica,  nose  of,  150. 
nemceus,  colouring  of,  551. 

rubicundus,  ornamental  l;air  ou 

the  head  of,  548. 

Senses,    inferiority    of  Europeans    to 

savages  in  the,  33. 
Sentinels,  among  animals,  100,  107. 
Serpents,    instinctively    dreaded     by 

apes  and  monkeys,  67,  71. 
Ferraniis,  hermaphroditism  in,  161. 
Sex,  inheritance  limited  by,  230. 
Sexes,  relative  proportions  of,  in  man, 

242,  558  ;  probable  relation  of  the, 

in  primeval  man,  591. 
Sexual    characters,    secondary,    207  ; 

relations    of    polygamy    to,    217  ; 

transmitted    through    both    sexes, 

226  ;  gradation  of,  in  birds,  430. 

and  natural  selection,  con- 
trasted, 226. 

characters,  effects  of  the  loss  of 

231  ;  limitation  of,  231. 

differences  in  man,  8. 

selection,    explanation    of,    209, 

213,  220;  influence  of,  on  the 
colouring  of  Lepidoptera,  319  ;  ob- 
jections to,  495  ;  action  of,  in  man- 
kind, 595. 

similarity,  255. 

Shaler,  Prof.,  sizes  of  sexes  in  whales, 
516. 

Sharks,  prehensile  organs  of  male, 
331. 

Sharpe,  Dr.,  Europeans  in  the  tropics, 
195. 

,  R.   B.,  on    Tanysiptera  sylvia, 

451  ;  on  Ceryle,  457  ;  on  the  young 
male  of  Dacelo  Gaudichaudi,  467. 

Shaw,  Mr.,  on  the  pugnacity  of  the 
male  salmon,  332. 

,  J.,  on  the  decorations  of  birds, 

382. 

Sheep,  danger-signals  of,  100  ;  sexual 
ditfsrences  in  the  horns  of,  230  ; 
horns  of,  235,  505  ;  domestic,  sexual 
differences  of,  late  developed,  237  ; 
numerical  proportion  of  the  sexes 
in,  246 ;  inheritance  of  horns  by 
one  sex,  505  ;  effect  of  castration, 
506;  mode  of  fighting  of,  508; 
arched  foreheads  of  some,  532. 

Merino,  loss  of  horns  in  females 

of,  231  ;  horns  of,  235. 

Shells,  difference  in  form  of,  in  male 
and     female     Gasteropoda,     262 ; 


beautiful  colours  and  shapes  of, 
264. 

Shield-drake,  pairing  with  a  common 
duck,  414;  New  Zealand,  sexes  and 
young  of,  479. 

Shooter,  J.,  on  the  Kaffirs,  579 ;  on 
the  marriage-customs  of  the  Kaffirs, 
598. 

Shrew-mice,  odour  of,  528. 

Shrike,  Drongo,  461. 

Shrikes,  characters  of  young,  465. 

Shuckard,  W.  E.,  on  sexual  differ- 
ences in  the  wings  of  Hymenoptera, 
277. 

Shyness  of  adorned  male  birds,  403. 

Siagonium,  proportions  of  the  sexes 
in,  253 ;  dimorphism  in  males  of, 
299. 

Siam,  proportion  of  male  and  female 
births  in,  245. 

Siamese,  general  beardlessness  of  the. 
560  ;  notions  of  beauty  of  the,  578  ; 
hairy  family  of,  601. 

Sidgwick,  H.,  on  morality  in  hypo- 
thetical bee  community,  99 ;  our 
actions  not  entirely  directed  by 
pain  and  pleasure,  120. 

Siebold,  C.  T.  von,  on  the  proportion 
of  sexes  in  the  Apus,  255 ;  on  the 
auditory  apparatus  of  the  stridii- 
lant  Orthoptera,  283. 

Sight,  inheritance  of  long  and  short, 
33. 

Signal-cries  of  monkeys,  87. 

Silk-moth,  proportion  of  the  sexes 
in,  250,  251;  Ailanthus,  Prof. 
Canestrini,  on  the  destruction  of 
its  larvae  by  wasps,  251 ;  differ- 
ence of  size  of  the  male  and  fe- 
male cocoons  of  the,  278 ;  pairing 
of  the,  318. 

Simiada;,  152  ;  their  origin  and  divi- 
sions, 165. 

Similarity,  sexual,  225. 

Singing  of  the  Cicad*  and  Fulgorida), 
281  ;  of  tree-frogs,  350  ;  of  birds, 
object  of  the,  368. 

Sirenia,  nakedness  of,  56. 

Sirex  juvencus,  292. 

Siricidae,  difference  of  the  sexes  in, 
292. 

Siskin,  394;  pairing  with  a  canary, 
415. 

Sitana,  throat-pouch  of  the  males  of, 
355,  308. 


INDEX. 


SPARROWS. 


^■77 


Size,  relative,  of  the  sexes  of  insects, 
278. 

Skin,  movement  of  the,  13 ;  ncaked- 
ness  of,  in  man,  56  ;  colour  of  the, 
192. 

and  hair,   correlation  of  colour 

of,  197. 

Skull,  variation  of,  in  man,  26  ; 
cubic  contents  of,  no  absolute  test 
of  intellect,  54 ;  Neanderthal,  ca- 
pacity of  the,  55  ;  causes  of  modi- 
fication of  the,  55 ;  ditference  of, 
in  form  and  capacity,  in  difterent 
races  of  men,  168;  variability  of 
the  shape  of  the,  174;  differences 
of,  in  the  sexes  in  man,  557  ;  arti- 
ficial modification  of  the  shape  of, 
575. 

Skunk,  odour  emitted  by  the,  528 ; 
white  tail  of,  protective,  543. 

Slavery,  prevalence  of,  117;  of  wo- 
men, 593. 

Slaves,  difference  between  field  and 
house-slaves,  196, 

Sloth,  ornaments  of  male,  534. 

Smell,  sense  of,  in  man  and  animals, 
17. 

Smith,  Adam,  on  the  basis  of  sym- 
pathy, 106. 

,  Sir  A.,   on  the  recognition    of 

women  by  male  CynocepUa'A,  8  ;  on 
revenge  by  a  baboon,  69  ;  on  an 
instance  of  memory  in  a  baboon, 
74  ;  on  the  retention  of  their  colour 
by  the  Dutch  in  South  Africa,  193  ; 
on  the  polygamy  of  the  South 
African  antelopes,  217;  on  the 
polygamy  of  the  lion,  218;  on  the 
proportion  of  the  sexes  in  Kohus 
cUipsiprymnus,  247  ;  on  Bucephalus 
capensis,  351  ;  on  South  African 
lizards,  358 ;  on  fighting  gnus, 
501  ;  on  the  horns  of  rhinoceroses, 
506  ;  on  the  fighting  of  lions,  521 ; 
on  the  colours  of  the  Cape  Eland, 
535;  on  the  colours  of  the  gnu, 
536;  on  Hottentot  notions  of 
beauty,  578;  disbelief  in  commu- 
nistic marriages,  588. 

• ,  F.,  on  the  Cynipida?  and  Ten- 

••thredinidae,  254;  on  the  relative 
size  of  the  sexes  of  Aculeate  Hymen- 
optera,  279 ;  on  the  ditlereuce  be- 
tween the  sexes  of  ants  and  bees, 
292;  on  the  stridulation  of   Trox 


sabulosus,  303;  on  the  stridulation 
of  Mononychus  pseudacori,  305. 

Smy7ithurus  luteus,  courtship  of,  279. 

Snakes,  sexual  dilierences  of,  351  ; 
mental  powers  of,  352  ;  male,  ar- 
dency of,  352. 

*'  Snarling  muscles,"  41. 

Snipe,  drumming  of  the,  376  ;  colo- 
ration of  the,  491. 

,  painted,    sexes    and  young    of, 

476. 

,  solitary,  assemblies  of,  405. 

Snipes,  arrival  of  male  before  the 
female,  212;  pugnacity  of  male, 
362;  double  moult  in,  390. 

Snow-goose,  whiteness  of  the,  492. 

Social  animals,  affection  of,  for  each 
othei-,  102 ;  defence  of,  by  the 
males,  107. 

Sociability,  the  sense  of  duty  con- 
nected with,  98 ;  impulse  to,  in 
animals,  105;  manifestations  uf, 
in  man,  108 ;  instinct  of,  in  ani- 
mals, 109. 

Sociality,  probable,  of  primeval  men, 
62 ;  influence  of,  on  the  develop- 
ment of  the  intellectual  faculties, 
128  ;  origin  of,  in  man,  129. 

Soldiers,  American,  measurements  of, 
29. 

and   sailors,    difference    in    the 

proportions  of,  32. 

Solenostoma,  bright  colours  and  mar- 
supial sack  of  the  females  of, 
346. 

Song  of  male  birds  appreciated  by 
their  females,  92  ;  want  of,  in 
brilliant  plumaged  birds,  401  ;  of 
birds,  450. 

Sorex,  odour  of,  528. 

Sounds  admired  alike  by  man  and 
animals,  92 ;  produced  by  fishes, 
347 ;  produced  by  male  frogs  and 
toads,  350  ;  instrumentally  pro- 
duced by  birds,  376  et  seq. 

Spain,  decadence  of,  141. 

Sparassus  smaragdulus,  difference  of 
colour  in  the  sexes  of,  272. 

Sparrow,  pugnacity  of  the  male,  360 ; 
acquisition  of  the  Linnet's  song  by 
a,  370  ;  coloration  of  the,  473  ;  im- 
mature plumage  of  the,  466. 

,   white-crowned,  young  of  the, 

486. 

Sparrows,  house-  and  tree-,  455. 


678 


IN^DEX. 


Sparrows,  new  mates  found  by,  408. 

,    sexes    and    young     of,    483 ; 

learning  to  sing,  570. 

Spatliura  Underwoodi,  388, 

Spawning  of  fishes,  341,  344, 

Spear,  used  before  dispersion  of  man, 
180. 

Species,  causes  of  the  advancement, 
of,  137  ;  distinctive  characters  of, 
166;  or  races  of  man,  168;  steri- 
lity and  fertility  of,  when  crossed, 
170;  supposed,  of  man,  174;  gra- 
dation of,  175  ;  difficulty  of  defin- 
ing, 175 ;  representative,  of  birds, 
468;  of  birds,  comparative  differ- 
ences between  the  sexes  of  distinct, 
469. 

Spectrum  femoratum,  difference  of 
colour  in  the  sexes  of,  289, 

Speech,  connection  between  the  brain 
and  the  faculty  of,  88  ;  connection 
of  intonation  with  music,  570.  . 

«  Spel  "  of  the  black-cock,  375. 

Sj)encer,  Herbert,  on  the  influence  of 
food  on  the  size  of  the  jaws,  33 ; 
on  the  dawn  of  intelligence,  67 ; 
on  the  origin  of  the  belief  in  spirit- 
ual agencies,  94 ;  on  the  origin  of 
the  moral  sense,  123 ;  on  music, 
571,  572. 

Spengel,  disagrees  with  explanation  of 
man's  hairlessness,  602. 

Sperm-whales,  battles  of  male,  500. 

Sphingidse,  coloration  of  the,  314. 

Sphinx,  Humming-bird,  317. 

,    Mr.    Bates  on   the  caterpillar 

of  a,  325. 

moth,  musky  odour  of,  308. 

Spiders,  272 ;  parental  feeling  in, 
106  ;  male,  more  active  than  female, 
221 ;  proportion  of  the  sexes  in,  254 ; 
secondary  sexual  characters  of,  272  ; 
courtship  of  male,  273  ;  attracted 
by  music,  273  ;  male,  small  size  of, 
273. 

Spilosoma  menthastri,  rejected  by 
turkeys,  316, 

Spine,  alteration  of,  to  suit  the  erect 
attitude  of  man,  53, 

Spirits,  fondness  of  monkeys  for,  7. 

Spiritual  agencies,  belief  in,  almost 
universal,  94, 

Spiza  cyanea  and  civis,  412, 

Spoonbill,  374;  Chinese,  change  of 
plumage  in,  461. 


Spots,  retained  throughout  groups  of 
birds,  427  ;  disappearance  of,  in 
adult  mammals,  546. 

Sprengel,  C.  K.,  on  the  sexuality  of 
plants,  212, 

Spring-boc,  horns  of  the,  509, 

Sproat,  Mr.,  on  the  extinction  of 
savages  in  Vancouver  Island,  183; 
on  the  eradication  of  facial  hair  by 
the  natives  of  Vancouver  Island, 
580 ;  on  the  eradication  of  the 
beard  by  the  Indians  of  Vancouver 
Island,  603, 

Spurs,  occurrence  of,  in  female  fowls, 
227,  231  :  development  of,  in 
various  species  of  Phasianidoe,  235  ; 
of  Gallinaceous  birds,  362,  364; 
development  of,  in  female  Gal- 
linaceas,  449, 

Sijuilla,  different  colours  of  the  sexes 
of  a  species  of,  271. 

Squirrels,  battles  of  male,  500 ;  Af- 
rican, sexual  differences  in  the 
colouring  of,  534 ;  black,  540. 

Stag,  long  hairs  of  the  throat  of,  521 ; 
horns  of  the,  227,  229  ;  battles  of, 
501 ;  horns  of  the,  with  numerous 
branches,  510 ;  bellowing  of  the, 
526  ;  crest  of  the,  531. 

beetle,  numerical  proportion  of 

sexes  of,  253  ;  large  size  of  male, 
278  ;  weapons  of  the  male,  299, 

Stainton,  H.  T.,  on  the  numerical 
proportion  of  the  sexes  in  the 
smaller  moths,  251 ;  habits  of  Ela- 
chista  rufocinerea,  252  ;  on  the 
coloration  of  moths,  315  ;  on  the 
rejection  of  Spilosoma  menthastri, 
by  turkeys,  316;  on  the  sexes  of 
Agrotis  exclamationis,  316, 

Staley,  Bishop,  mortality  of  infant 
Maories,  187. 

Stallion,  mane  of  the,  521. 

Stallions,  two,  attacking  a  third,  101  ; 
fighting,  501 ;  small  cauiue  teeth 
of,  514, 

Stansbury,  Capt,,  observations  on 
pelicans,  102, 

Staphylinida?,  hornlike  processes  in 
male,  299. 

Starfishes,  parental  feeling  in,  106  ; 
bright  colours  of  some,  260. 

Stark,  Dr.,  on  the  death-rate  in  towns 
and  rural  districts,  138;  on  the 
influence  of  marriage  on  mortality. 


INDEX. 


SUPRA-COND^LOID. 


679 


139  ;  on  the  higher  mortality  of 
males  in  Scotland,  243. 

Starling,  American  field-,  pugnacity  of 
male,  367. 

,  red-winged,  selection  of  a  mate 

by  the  female,  416. 

Starlings,  three,  frequenting  the  same 
nest,  219,  409;  new  mates  found 
by,  408. 

Statues,  Greek,  Egyptian,  Assyrian, 
&c.,  contrasted,  581. 

Stature,  dependence  of,  upon  local 
influences,  31. 

Staudiuger,  Dr.,  on  breeding  Lepid- 
optera,  251 ;  his  list  of  Lepid- 
optera,  252. 

Staunton,  Sir  G.,  hatred  of  indecency 
a  modern  virtue,  119. 

Stealing  of  bright  objects  by  birds, 
413. 

Stebbing,  T.  R.,  on  the  nakedness  of 
the  human  body,  600. 

Stemmatopus,  528. 

Stendhal,  see  Bombet. 

Stenohothrus  prutonim,  stridulation, 
286. 

Stephen,  Mr.  L.,  on  the  difference  in 
the  minds  of  men  and  animals,  78  ; 
on  general  concepts  in  animals, 
89 ;  distinction  between  material 
and  formal  morality,  111. 

Sterility,  general,  of  sole  daughters, 
135 ;  when  crossed,  a  distinctive 
character  of  species,  166 ;  under 
changed  conditions,  189,  191. 

Sterna,  seasonal  change  of  plumage  in, 
493. 

Stickle-back,  polygamous,  220  ;  male, 
courtship  of  the,  331  ;  male,  bril- 
liant colouring  of,  during  the  breed- 
ing season,  340  ;  nidification  of  the, 
345. 

Sticks  used  as  implements  and  wea- 
pons by  monkeys,  81. 

Sting  in  bees,  208. 

Stokes,  Capt.,  on  the  habits  of  the 
great  bower-bird,  381. 

Stoliczka,  Dr.,  on  colours  in  snakes, 
352. 

Stonechat,  young  of  the,  487. 

Stone  implements,  difficulty  of  making, 
49  ;  as  traces  of  extinct  tribes,  181. 

Stones,  used  by  monkeys  for  breaking 
■  hard  fruits  and  as  missiles,  50 ; 
piles  of,  179. 


Stork,  black,  sexual  differences  in  the 
bronchi  of  the,  374  ;  red  beak  of 
the,  491. 

Storks,  491,  493 ;  sexual  difference 
in  the  colour  of  the  eyes  of,  425. 

Strange,  Mr.,  on  the  satin  bower- 
bird,  381. 

Stretch,  Mr.,  on  the  numerical  pro- 
portion in  the  sexes  of  chickens, 
247.  ■ 

Strepsiceros  kudu,  horns  of,  512 ;  mark- 
ings of,  543. 

Stridulation,  by  males  of  Theridion, 
274  ;  of  the  Orthoptera  and  Homop- 
tera  discussed,  289 ;  of  beetles, 
301. 

Stripes,  retained  throughout  groups 
of  birds,  427 ;  disappearance  of,  in 
adult  mammals,  546. 

Strix  flammea,  408. 

Structure,  existence  of  unserviceable 
modifications  of,  61. 

Struggle  for  existence,  in  man,  142, 
146. 

Struthers,  Dr.,  on  the  occurrence  of 
the  supra-condyloid  forame%in  the 
humerus  of  man,  21. 

Sturnella  ludoviciana,  pugnacity  of  the 
male,  367. 

Sturnus  vulgaris,  408. 

Sub-species,  175. 

Suffering,  in  strangers,  indifference  of 
savages  to,  117. 

Suicide,  137  ;  formerly  not  regarded 
as  a  crime,  117  ;  rarely  practised 
among  the  lowest  savages,  117. 

SuidcTe,  stripes  of  young,  464. 

Sulivan,  Sir  B.  J.,  on  speaking  of 
parrots,  85  ;  on  two  stallions  at- 
tacking a  third,  501. 

Sumatra,  compression  of  the  nose  by 
the  Malays  of,  583. 

Sumner,  Archb.,  man  alone  capable  of 
progressive  improvement,  79. 

Sun-birds,  nidification  of,  454. 

Superstitions,  144  ;  prevalence  of, 
122. 

Superstitious  customs,  96. 

Superciliary  ridge  in  man,  556,  558. 

Supernumerary  digits,  more  frequent 
in  men  than  in  women,  223  ;  in- 
heritance of,  232  ;  early  develop- 
ment of,  237. 

Supra-condyloid  foramen  in  the  early 
progenitors  of  man,  160. 


68o 


SUSPICION. 


INDEX. 


Suspicion,  prevalence  of,  among  ani- 
mals, 69. 

Swallow-tail  butterfly,  312. 

Swallows  deserting  their  youns:,  108, 
113. 

Swan,  black,  wild,  trachea  of  the, 
374 ;  white  young  of,  482  ;  red  beak 
of  the,  491  ;  black-necked,  493. 

Swans,  491,  493;  young,  481. 

Swaysland,  Mi.,  on  the  arrival  of 
migratoi-y  birds,  212. 

Swifts,  migration  of,  108. 

Swinhoe,  R.,  on  the  common  rat  in 
Formosa  and  China,  80  ;  behaviour 
of  lizards  when  caught,  355  ;  on 
the  sounds  produced  by  the  male 
hoopoe,  376 ;  on  Dicrurus  macro- 
cercus  and  the  spoonbill,  461  ;  on 
the  young  of  Ardeola,  468 ;  on  the 
habits  of  Turnix,  476 ;  on  the 
habits  of  Rhynchoca  bengalensis, 
476  ;  on  Orioles  breeding  in  imma- 
ture plumage,  484. 

Sylvia  atricapilla,  young  of,  487. 

cinerea,  aerial  love-dance  of  the 

male,  380 

Sympathy,  134  ;  among  animals,  102  ; 
its  supposed  basis,  106. 

Sympathies,  gradual  widening  of, 
123. 

Syngnathous  fishes,  abdominal  pouch 
in  male,  163. 

Sypheotides  auritus,  acuminated  pri- 
maries of  the  male,  378  ;  ear-tufts 
of,  384 


Tabanidje,  habits  of,  208. 

Tadorna  variegata,  sexes  and  young 
of,  479. 

vulpanser,  414. 

Tahitians,  145  ;  compression  of  the 
nose  by  the,  583. 

Tail,  rudimentary,  occurrence  of,  in 
man,  22  ;  convoluted  body  in  the 
extremity  of  the,  23  ;  absence  of, 
in  man  and  the  higher  apes,  58  ; 
variability  of,  in  species  of  Macacus 
and  in  baboons,  58  ;  presence  of,  in 
the  early  progenitors  of  man,  160; 
length  of,  in  pheasants,  445,  451, 
452  ;  diflerence  of  length  of  the, 
in  the  two  sexes  of  birds,  451. 

Tait,  Lawson,  on   the   eflects   of  na- 


tural selection  on  civilised  nations, 
133. 

Tanager,  scarlet,  variation  in  the 
male,  424. 

Tanagra  oestiva,  age  of  mature  plu- 
mage in,  483. 

rM6ra424;  young  of,  487. 

Tanais,  absence  of  mouth  in  the  males 
of  some  species  of,  208  ;  relations 
of  the  sexes  in,  255 ;  dimorphic 
males  of  a  species  of,  265. 

Tankerville,  Earl,  on  the  battles  of 
wild  bulls,  501. 

Tanysiptera,Yaces  of,  determined  from 
adult  males,  468. 

Sylvia,  long  tail-feathers  of,  451. 

Taphroderes  distortus,  enlarged  left 
mandible  of  the  male,  277. 

Tapirs,  longitudinal  stripes  of  young, 
464,  546. 

Tarsi,  dilatation  of  front,  in  male 
beetles,  275. 

Tarsius,  156. 

Tasmania,  half-castes  killed  by  the 
natives  of,  170. 

Tasmanians,  extinction  of,  184. 

Taste,  in  the  Quadrumana,  541. 

Tattooing,  178  ;  tmiversality  of,  574. 

Taylor,  G.,  on  Quiscalus  major,  248. 

Tea,  fondness  of  monkeys  for,  7. 

Tear-sacks,  of  Ruminants,  529. 

Teebay,  Mr.,  on  changes  of  plumage 
in  spangled  Hamburgh  fowls,  229. 

Teeth,  rudimentary  incisor,  in  Rumi- 
nants, 11  ;  posterior  molar,  in  man, 
20  ;  wisdom,  20  ;  diversity  of,  26  ; 
canine,  in  the  early  progenitors  ot 
man,  160 ;  canine,  of  male  mam- 
mals, 502 ;  in  man,  reduced  by 
correlation,  562  ;  staining  of  the, 
574;  front,  knocked  out  or  filed 
by  some  savages,  575. 

Tegetmeier,  Mr.,  on  the  transmission 
of  colours  in  pigeons  by  one  sex 
alone,  231;  numerical  proportion 
of  male  and  female  births  in  dogs, 
246 ;  on  the  abundance  of  male 
pigeons,  247  ;  on  the  wattles  of 
game-cocks,  403  ;  on  the  courtship 
of  fowls,  417;  on  the  loves  of  pigeons, 
418;  on  dyed  pigeons,  418;  blue 
dragon  pigeons,  446. 

Tembeta,  S.  American  ornament,  575. 

Temper,  in  dogs  and  horses,  inherited, 
69. 


TENCH. 


INDEX. 


TOADS. 


68 1 


Tench,  proportions  of  the  sexes  in  the 

249;    brightness    of  male,    during 

breeding  season,  340. 
Tenebrionidse,  stridulation  of,  302. 
Tennent,  Sir  J.  E.,  on   the  tusks  of 

the   Ceylon    Elephant,   507,    515; 

on  the  frequent  absence  of  beard  in 

the  natives  of  Ceylon,  560  ;  on  the 

Chinese   opinion  of  the    aspect  of 

the  Cingalese,  578. 
Tennyson,    A.,     on    the    control    of 

thought,  123. 
Tenthredinidpe,    proportions    of    the 

sexes  in,    254;    fighting    habits  of 

male,  291 ;  difference  of  the  sexes 

in,  292. 
Tephrodoniis,  young  of,  468. 
Terai,  in  India,  182. 
Termites,  habits  of,  291. 
Terns,  white,  492 ;  and  black,  493. 
,  seasonal  change  of  plumage  in, 

492. 
.  Terror,  common  action  of,  upon  the 

lower  animals  and  man,  69. 
Testudo  elcgans,  351. 

nigra,  351. 

Tetrao  cupido,  battles  of,  367  ;  sexual 

difference  in  the  vocal  organs   of, 

371. 
phasianellus,    dances    of,    380 ; 

duration  of  dances  of,  405. 

scoticus  455,  465,  471. 

tetrix,  455,  465, 471 ;  pugnacity 

of  the  male,  363. 
umbeUics,  pairing  of,  367  ;  battles 

of,  367  ;  drumming    of  the    male, 

375. 

urogalloides,  dances  of,  405. 

urogallus,  pugnacity  of  the  male, 

363. 

urophasianus,   inflation   of  the 

oesophagus  in  the  male,  372. 

Thamnobia,  young  of,  468. 
Thaumalea  picta,  display  of  plunjage 

by  the  male,  396. 
Theda,  sexual  differences  of  colouring 

in  species  of,  310. 

ruhi,   protective    colouring   of, 

312. 

Inecophora  fovea,  308. 
Theognis,  selection  in  mankind,  29. 
Theridion,  stridulation   of  males  of, 
273. 

lineatum,  272. 

Thomisus   citreas,   and   T.  floricokns, 

30 


difference  of  colour  in  the  sexes  of, 

272. 
Thompson,  J.  H.,  on  the    battles    of 

sperm-whales,  500. 
,  W.,  on   the    colouring   of  the 

male   char    during    the    breeding 

season,  340  ;   on  the  pugnacity  of 

the   males  of   GaUinula   chloropus, 

360  ;  on  the  finding  of  new  mates 

by  magpies,  407  ;  on  the  finding  of 

new  mates   by    Peregrine    falcons, 

408. 
Thorax,  j.rocesses  of,  in  male  beetles, 

295. 
Thorell,  T.,  on  the  proportion  of  the 

sexes  in  spiders,  255. 
Thornback,  difference  in  the  teeth  of 

the  two  sexes  of  the,  334. 
Thoughts,  control  of,  123. 
Thrush,    pairing   with    a   blackbird, 

414 ;    colours   and   nidification   of 

the,  455. 
Thrushes,  characters  of  young,  455, 

464. 
Thug,  remorse  of  a,  117. 
Thumb,  absence  of,  in  Ateles  and  Hy- 

lobates,  51. 
Thury,  M.,  on  the  numerical  propor- 
tion  of  male   and    female    births 

among  the  Jews,  243. 
T/iylacinus,  possession  of  the  marsupial 

sack  by  the  male,  161 
Thysanura,  279. 
Tibia,  dilated,  of  the  male  Crabo  cri- 

brarius,  276. 
and  femur,   proportions  of,   in 

the  Aymara  Indians,  34. 
Tierra   del  Fuego,  marriage-customs 

of,  598. 
Tiger,  colours  and  markings  of  the, 

545. 
Tigers,  depopulation  of  districts  by,  in 

India,  46. 
Tillus  elongatus,  difference  of  colour 

in  the  sexes  of,  294. 
Timidity,  variability  of,  in  the  same 

species,  69. 
Tineina,  proportion  of  the  sexes  in, 

250. 
Tipula,  pugnacity  of  male,  280. 
Tits,   sexual   difference  of  colour  in, 

458. 
Toads,  349 ;  male,  treatment  of  ova 

by  some,  163  ;  male,  ready  to  breed 

before  the  female,  212. 


682 


INDEX. 


TUEDDS. 


Todas,  iufanticide  and  proportion  of 
sexes,  255 ;  practise  polyandry, 
593 ;  choice  of  husbands  amongst, 
593. 

Toe,  great,  condition  of,  in  the  human 
embryo,  11. 

Tomicus  villosus  proportion  of  the 
sexes  in,  253. 

Tomtit,  blue,  sexual  difference  of  co- 
lour in  the,  458. 

Tonga  Islands,  beardlessness  of  the 
natives  of,  560,  581. 

Tooke,  Home,  on  language,  86. 

Tools,  flint,  145 ;  used  by  monkeys, 
81 ;  use  of,  48. 

Topknots  in  birds,  384. 

Tortoise,  voice  of  the  male,  567, 

Tortures,  submitted  to  by  American 
savages,  118. 

Totanus,  double  moult  in,  390. 

Toucans,  colours  and  nidification  of 
the,  455;  beaks  and  ceres  of  the, 
491. 

Towns,  residence  in,  a  cause  of  dimi- 
nished stature,  31. 

Toynbee,  J.,  on  the  external  shell  of 
the  car  in  man,  14. 

Trachea,  convoluted  and  imbedded  in 
the  sternum,  in  some  birds,  374 ; 
structure  of  the,  in  Rhynchcea,  476. 

Trades,  affecting  the  form  of  the 
skull,  56. 

Tragelaphus,  sexual  differences  of 
colour  in,  535. 

scriptus,  dorsal   crest   of,  530 ; 

markings  of,  543,  544. 

Tragopan,  220;  swelling  of  the  wat- 
tles of  the  male,  during  courtship, 
383;  display  of  plumage  by  the 
male,  397  ;  markings  of  the  sexes 
of  the,  428. 

Tracjops  dispar,  sexual  difference  in 
the  colour  of,  351. 

Training,  effect  of,  on  the  mental 
difference  between  the  sexes  of 
man,  565. 

Transfer  of  male  characters  to  female 
birds,  470. 

Transmission,  equal,  of  ornamental 
characters,  to  both  sexes  in  mam- 
mals, 542. 

Traps,  avoidance  of,  by  animals,  80 ; 
use  of,  48. 

Treachery,  to  comrades  avoidance  of, 
by  savages,  111 


Tremex  columhce,  292. 

Tribes,  extinct,  128;  extinction  of, 
182. 

Trichius,  difference  of  colour  in  the 
sexes  of  a  species  of,  294. 

Trigla,  347. 

Trimen,  R.,  on  the  proportion  of  the 
sexes  in  South  African  butterflies, 
250  ;  on  the  attraction  of  males  by 
the  female  of  Lasiocampa  quercus, 
252  ;  on  Pneumora,  288 ;  on  dif- 
ference of  colour  in  the  sexes  of 
beetles,  294;  on  moths  brilliantly 
coloured  beneath,  315  ;  on  mimicrv 
in  butterflies,  325;  on  Gynanis'a 
Isls,  and  on  the  ocellated  spots  of 
Lepidoptera,  428;  on  CyUo  Ledt, 
429. 

Tringa,  sexes  and  young  of,  485. 

cornuta,  391. 

Triphceni,  coloration  of  the  species 
of,  313. 

Tristram,  H.  B.,  on  unhealthy  dis- 
tricts in  North  Africa,  1 94 ;  on 
the  habits  of  the  chaffinch  in  Pales- 
tine, 248 ;  on  the  birds  of  the 
Sahai'a,  456 ;  on  the  animals  in- 
habiting the  Sahara,  489. 

Triton  cristatns,  348. 

palmipes,  348. 

p-mctatus,  348. 

Troglodyte  skulls,  greater  than  those 
of  modern  Frenchmen,  55. 

Troglodytes  vulgaris,  473. 

Trogons,  colours  and  nidification  of 
the,  455,  456. 

Tropic-birds,  white  only  when  mature, 
492. 

Tropics,  freshwater  fishes  of  the, 
343. 

Trout,  proportion  of  the  sexes  in, 
249  ;  male,  pugnacity  of  the,  332. 

Trox  sabulosus,  stridulation  of,  303. 

Truth,  not  rare  between  members  of 
the  same  tribe,  118;  more  highly 
appreciated  by  certain  tribes,  122. 

Tulloch,  Major,  on  the  immunity  of 
the  negro  from  certain  fevers,  193. 

Tumbler,  almond,  change  of  plumage 
in  the,  238. 

Tardus  merula,  455  ;  young  of,  487, 

migratorius,  464. 

musicns,  455. 

polyglottus,  young  of,  487t 

torquutus,  455 


TURKEY. 


INDEX. 


6^^ 


Turkey,  wild,  pugnacity  of  young 
male,  366  ;  wild,  notes  of  the,  375  ; 
swelling  of  the  wattles  of  the  male, 
383 ;  variety  of,  with  a  top-knot, 
384;  recognition  of  a  dog  by  a,  412  ; 
male,  wild,  acceptable  to  domesti- 
cated females,  418  ;  wild,  first  ad- 
vances made  by  older  females,  419  ; 
wild,  breast-tuft  of  bristles  of  the, 
461. 

Turkey-cock,  scraping  of  the  wings 
of,  upon  the  ground,  375  ;  wild, 
display  of  plumage  by,  394;  fight- 
ing habits  of,  404. 

Turner,  Prof.  W.,  on  muscular  fasci- 
culi in  man  referable  to  the  pan- 
niculus  carnosus,  13 ;  on  the  occur- 
rence of  the  supra-condyloid  foramen 
in  the  hnman  humerus,  21 ;  on 
muscles  attached  to  the  coccyx  in 
man,  22  ;  on  the  filum  terminale  in 
man,  23  ;  on  the  variability  of  the 
muscles,  26  ;  on  abnormal  conditions 
of  the  human  uterus,  39  ;  on  the 
development  of  the  mammary 
glands,  162 ;  on  male  fishes  hatch- 
ing ova  in  their  mouths,  163,  345; 
on  the  external  perpendicular  fissure 
of  the  brain,  200  ;  on  the  bridging 
convolutions  in  the  brain  of  a  chim- 
panzee, 201. 

lurnix,  sexes  of  some  species  of,  475  ; 
480. 

Turtle-dove,  cooing  of  the,  374. 

Tuttle,  H.,  on  the  number  of  species 
of  man,  174. 

Tylor,  E.  B.,  on  emotional  cries,  ges- 
tures, &o.,  of  man,  85;  on  the 
origin  of  the  belief  in  spiritual 
agencies,  94  ;  remorse  for  violation 
of  tribal  usage  in  marrying,  115; 
on  the  primitive  barbarism  of 
civilised  nations^  143  ;  on  the  origin 
of  counting,  144 ;  inventions  of 
savages,  145  ;  on  resemblances,  of 
the  mental  characters  indifferent 
races  of  man,  178. 

Type  of  structure,  prevalence  of,  164. 

Typhceus,  stridulatiug  organs  of,  301  ; 
stridulation  of,  303. 

Tyranga  cestiva,  462. 

Twins,  tendency  to  produce,  here- 
ditary, 45. 

Twite,  proportion  of  the  sexes  in  the, 
248. 


U. 

Ugliness,  said  to  consist  in  an  approach 
to  the  lower  animals,  584 

Umbrella-bird,  373, 

Umhrina,  sounds  produced  by,  347. 

United  States,  rate  of  increase  in,  44  ; 
influence  of  natural  selection  on 
the  progress  of,  142 ;  change  under- 
gone by  Europeans  in  the,  196. 

Upupa  epops,  sounds  produced  by  the 
male,  376, 

Uraniidfe,  coloration  of  the,  314. 

[fria  troilc,  variety  of  (=  U,  lacry- 
7nnns),  424. 

Urodela,  348. 

Urosticte  Benjammi,  sexual  differences 
in,  442. 

Use  and  disuse  of  parts,  effects  of,  32  , 
influence  of,  on  the  races  of  man, 
197. 

Uterus,  reversion  in  the,  38  ;  more  or 
less  divided,  in  the  human  subject, 
38,  43  ;  double,  in  the  early  pro- 
genitors of  man,  161. 

V. 

Vaccination,  influence  of,  134. 

Vancouver  Island,  Mr.  Sproat  on  the 
savages  of,  183  ;  natives  of,  eradi- 
cation of  facial  hair  by  the,  580. 

Vanellus  cristatus,  wing  tubercles  of 
the  male,  366. 

VanessoB,  308  ;  resemblance  of  lower 
surface  of,  to  bark  of  trees,  311. 

Variability,  causes  of,  28  ;  in  man, 
analogous  to  that  in  the  lower 
animals,  29 ;  of  the  races  of  man, 
174;  greater  in  men  than  in 
women,  223;  period  of,  relation  of 
the,  to  sexual  selection,  240 ;  of 
birds,  422  ;  of  secondary  sexual 
characters  in  man,  559. 

Variation,  laws  of,  29 ;  correlated, 
43;  in  man,  146;  analogous,  152- 
analogous,  in  plumage  of  birds,  385 

Variations,  spontaneous,  44. 

Varieties,  absence  of,  between  two 
species,  evidence  of  their  distinct- 
ness, 167. 

Variety,  an  object  in  nature,  493. 

Variola,  communicable  between  man 
and  the  lower  animals,  7. 


634 


VAUREAL. 


INDEX. 


WALLACE. 


Vaureal,  human  bones  from,  22. 

Veddahs,  monogamous  habits  of,  591. 

Veitch,  Mr.,  on  the  aversion  of 
Japanese  ladies  to  whiskers,  581. 

Vengeance,  instinct  of,  113. 

Venus  Erycina,  priestesses  of,  587. 

Vermes,  264. 

Vermiform  appendage,  21. 

Verreaux,  M.,  on  the  attraction  of 
numerous  males  by  the  female  of 
an  Australian  Bomhyx,  252. 

Vertebrae,  caudal,  number  of,  in 
macaques  and  baboons,  58 ;  of 
monkeys,  partly  imbedded  in  the 
body,  59. 

Vertebvata,  330 ;  common  origin  of 
the,  158 ;  most  ancient  progenitors 
of,  161 ;  origin  of  the  voice  in  air- 
breathing,  567. 

Vesicula  prostatica,  the  homologue  of 
the  uterus,  24-,  161. 

Vibrissse,  represented  by  long  hairs 
in  the  eyebrows,  19. 

Vidua,  403,  462. 

axillaris,  219. 

Villerme,  M.,  on  the  influence  of 
plenty  upon  stature,  31. 

Vinson,  Aug.,  courtship  of  male 
spider,  273  ;  on  the  male  of  Epeira 
nigra,  273. 

Viper,  difference  of  the  sexes  in  the, 
351. 

Virey,  on  the  number  of  species  of 
man,  174. 

Virtues,  originally  social  only,  116  : 
gradual  appreciation  of,  131. 

A'iscera,  variability  of,  in  man,  27. 

Vlacovich,  Prof.,  on  the  ischio-pubic 
muscle,  41. 

Vocal  music  of  birds,  368. 

organs  of  man,  87  ;  of  birds,  90, 

450 ;  of  frogs,  350 ;  of  the  Inses- 
sores,  370;  difference  of,  in  the 
sexes  of  birds,  371  primarily  used 
in  relation  to  the  propagation  of 
the  species,  566. 

Vogt,  Karl,  on  the  origin  of  species,  1  ; 
on  the  origin  of  man,  3 ;  on  the 
semilunar  fold  in  man,  17  ;  on  mi- 
crocephalous idiots,  35  ;  on  the  imi- 
tative faculties  of  microcephalous 
idiots,  87;  on  skulls  from  Brazilian 
caves,  168  ;  on  the  evolution  of  the 
races  of  man,  177  ;  on  the  formation 
of  the  skull  in  women,  557  :  on  the 


Ainos  and  negroes,  560 ;  on  the  in- 
creased cranial  difference  of  the 
sexes  in  man  with  race  development, 
566  ;  on  the  obliquity  of  the  eye  in 
the  Chinese  and  Japanese,  578. 

Voice  in  mammals,  525 ;  in  monkeys 
and  man,  558  ;  in  man,  566  ;  origin 
of,  in  air-breathing  vertebrates,  567. 

Von  Baer,  definition  of  advancement 
in  the  organic  scale,  164. 

Vulpian,  Prof.,  on  the  resemblance 
between  the  brains  of  man  and  of 
the  higher  apes,  6. 

Vultures,  selection  of  a  mate  by  the 
female,  416  ;  colours  of,  493. 


W. 

Waders,  young  of,  486. 

Wagner,  E.,  on  the  occurrence  of  the 
diastema  in  a  Kaffir  skull,  40;  on 
the  bronchi  of  the  black  stork,  374, 

Wagtail,  Pvay's,  arrival  of  the  male 
before  the  female,  212. 

Wagtails,  Indian,  young  of,  468. 

Waist,  proportions  of,  in  soldiej-s  and 
sailors,  32. 

Waitz,  Prof.,  on  the  number  of  spe- 
cies of  man,  174 ;  on  the  liability 
of  negroes  to  tropical  fevers  after 
residence  in  a  cold  climate,  194; 
on  the  colour  of  Australian  infants, 
558 ;  on  the  beardlessness  of 
negroes,  560 ;  on  the  fondness  of 
mankind  for  ornaments,  573 ;  on 
negro  ideas  of  female  beauty,  579  ; 
on  Javan  and  Cochin  Chinese  ideas 
of  beauty,  580. 

Walckenaer  and  Gervais,  spider  at- 
tracted by  music,  273  ;  on  the  Myri- 
apoda,  274. 

Waldeyer,M  ,  on  the  hermaphroditism 
of  the  vertebrate  embryo,  161. 

Wales,  North,  numerical  proportion  ot 
male  and  female  births  in,  242. 

Walker,  Alex.,  on  the  large  size  of  the 
hands  of  labourers'  children,  33. 

,  F.,  on  sexual  differences  in 

the  diptera,  280. 

Wallace,  Dr.  A.,  on  the  prehensile 
use  of  the  tarsi  in  male  moths,  209  ; 
on  the  rearing  of  the  Ailanthus 
silkmoth,  251 ;  on  breeding  Lepi- 
doptera,  251 ;  proportion  of  sexes  of 
Bombyx  cynthia,  B.  yamamai,  and 


INDEX. 


WATERIIOUSE. 


685 


B.  Pernyi  reared  by,  253 ;  on  the 
development  of  Bomhyx  cynthia 
and  B.  yamamai,  278  ;  on  the  pair- 
ing oi  Bomhyx  cynthia,  318. 
Wallace,  A.  R.,  on  the  origin  of 
man,  3  ;  on  the  power  of  imitation 
in  man,  68 ;  on  the  use  of  missiles 
by  the  orang,  82 ;  on  the  varying 
appreciation  of  truth  among  diffei'- 
ent  tribes,  122  ;  on  the  limits  of 
natural  selection  in  man,  49,  127  ; 
on  the  occurrence  of  remorse  among 
savages,  131 ;  on  the  effects  of 
natural  selection  on  civilised  na- 
tions, 133 ;  on  the  use  of  the  con- 
vergence of  the  hair  at  the  elbow  in 
the  orang,  151 ;  on  the  contrast  in 
the  characters  of  the  Malays  and 
Papuans,  168 ;  on  the  line  of  sepa- 
ration between  the  Papuans  and 
Malays,  169;  on  the  birds  of  para- 
dise, 219 ;  on  the  sexes  of  Ornithop- 
tera  Crcesus,  250;  on  protective  re- 
semblances, 261  ;  on  the  relative  sizes 
of  the  sexes  of  insects,  278  ;  on  Ula- 
phomyia,  280 ;  on  the  pugnacity  of 
the  males  of  Leptorhjnckus  angus- 
tatus,  299  ;  on  sounds  produced  by 
Euchirus  longimanus,  304;  on  the 
colours  of  Diadeina,  309 ;  on  Kal- 
lima,  311 ;  on  the  protective  colour- 
ing of  moths,  313  ;  on  bright  colora- 
ration  as  protective  in  butterflies, 
314;  on  variability  in  the  Papi- 
lionidae,  320 ;  on  male  and  female 
butterflies  inhabiting  different  sta- 
tions, 321  ;  on  the  protective  nature 
of  the  dull  colouring  of  female  but- 
terflies, 321,  322,  325  ;  on  mimicry 
in  butterflies,  324;  on  the  bright 
colours  of  caterpillars,  325 ;  on 
brightly-coloured  fishes  frequenting 
reefs,  343;  on  the  coral  snakes,  353  ; 
on  Paradisea  apoda,  385;  on  the 
display  of  plumage  by  male  birds 
of  paradise,  395 ;  on  assemblies  of 
birds  of  paradise,  405 ;  on  the  in- 
stability of  the  ocellated  spots  in 
Hipparchia  Janira,  428;  on  sexually 
limited  inheritance,  444 ;  on  the 
sexual  coloration  of  birds,  452, 472, 
473,  475,  480;  on  the  relation  be- 
tween the  colours  and  nidification 
of  birds,  452,  455 ;  on  the  colora- 
tion of  the  Cotingidae,  460 ;  on  the 


females  of  Paradisea  apoda  and 
papuana,  470 ;  on  the  incubation 
of  the  cassowary,  478  ;  on  protective 
coloration  in  birds,  489 ;  on  the 
Babirusa,  519;  on  the  markings  of 
the  tiger,  545  ;  on  the  beards  of  the 
Papuans,  560;  on  the  hair  of  the 
Papuans,  575 ;  on  the  distribution 
of  hair  on  the  human  body,  600. 

Walrus,  development  of  the  nicti- 
tating membrane  in  the,  17  ;  tusks 
of  the,  502,  507  ;  use  of  the  tusks 
by  the,  513. 

Walsh,  B.  D.,  on  the  proportion  of 
the  sexes  in  Papilio  Tumus,  250; 
on  the  Cynipidaj  and  Cecidomyidce, 
254 ;  on  the  jaws  of  Ammophila, 
275;  on  Corydalis  cornutus,  275; 
on  the  prehensile  organs  of  male 
insects,  275 ;  on  the  antennae  of 
Penthe,  275 ;  on  the  caudal  appen- 
dages of  dragon-flies,  276  ;  on  Platy- 
phyllum  cone  (vum,  286  ;  on  the  sexes 
of  the  Ephemeridse,  289 ;  on  the 
difference  of  colour  in  the  sexes  of 
Spectrum  femoratum,  289 ;  on  sexes 
of  dragon-flies,  290 ;  on  the  differ- 
ence of  the  sexes  in  the  Ichneu- 
monidce,  292  ;  on  the  sexes  of  Orso- 
dacna  at'Xi,  294;  on  the  variation 
of  the  horns  of  the  male  Phanceus 
carnifex,  297 ;  on  the  coloration  of 
the  species  of  Anthocharis,  312. 

Wapiti,  battles  of,  501 ;  traces  of 
horns  in  the  female,  504;  attacking 
a  man,  51 1  ;  crest  of  the  male,  531 ; 
sexual  difference  in  the  colour  of 
the,  536. 

Warbler,  hedge-,  473  ;  young  of  the, 
481. 

Warblers,  superb,  nidification  of,  454. 

Wariness,  acquired  by  animals,  80. 

Warington,  R.,  on  the  habits  of  the 
stickle-backs,  331,  345 ;  on  the 
brilliant  colours  of  the  male 
stickle-back  during  the  breeding 
season,  340. 

Wart-hog,  tusks  and  pads  of  the,  519. 

Watchmakers,  short-sighted,  33. 

Waterhen,  360. 

Waterhouse,  C.  0.,  on  blind  beetles, 
294 ;  on  difference  of  colour  in 
the  sexes  of  beetles,  294. 

,  G.  R.,  on  the  voice  of  Hylo\>o.tcs 

agilis,  567. 


em 


WATER-OUZEL. 


INDEX. 


\\ESTWO0D. 


Water-ouzel,  455 ;  autumn  song  of 
the,  370. 

Watcrton,  C,  on  the  Bell-bird,  389  ; 
on  the  pairing  of  a  Canada  goose 
with  a  Bernicle  gander,  414 ;  on 
hares  fighting,  500. 

Wattles,  disadvantageous  to  male 
birds  in  fighting,  404. 

Weale,  J.  Mansel,  on  a  South  African 
caterpillar,  325. 

Wealth,  influence  of,  134. 

Weapons,  used  by  man,  48  ;  employed 
by  monkeys,  81 ;  offensive,  of  males, 
210;  of  mammals,  501  et  seq. 

Weaver-bird,  370. 

Weaver-birds,  rattling  of  the  wings 
of,  375;  assemblies  of,  405. 

Webb,  Dr.,  on  the  wisdom  teeth,  20. 

Wedderburn,  Mr,,  assembly  of  black 
game,  407. 

Wedgwood,  Hensleigh,  on  the  origin 
of  language,  87. 

Weevils,  sexual  difference  in  length  of 
snout  in  some,  208. 

Weir,  Harrison,  on  the  numerical 
projxirtion  of  the  sexes  in  pigs  and 
rabbits,  247  ;  on  the  sexes  of  young 
pigeons,  247  ;  on  the  songs  of  birds, 
368  ;  on  pigeons,  411 ;  on  the  dis- 
like of  blue  pigeons  to  other  co- 
loured varieties,  417  ;  on  the  de- 
sertion of  their  mates  by  female 
pigeons,  418. 

■ ,  J.  Jenner,   on  the  nightingale 

and  blackcap,  212  ;  on  the  relative 
sexual  maturity  of  male  birds,  213  ; 
on  female  pigeons  deserting  a  feeble 
mate,  214;  on  three  starlings  fre- 
quenting the  same  nest,  219  ;  on 
the  proportion  of  the  sexes  in  Ma- 
c/ietes  pugnax  and  other  birds,  247, 
248 ;  on  the  coloration  of  the  Tri- 
pkoence,  313;  on  the  rejection  of 
certain  caterpillars  by  birds,  326  ; 
on  sexual  diflerences  of  the  beak  in 
the  goldfinch,  360 ;  on  a  ])iping 
bullfinch,  369  ;  on  the  object  of  the 
nightingale's  song,  368;  on  song- 
birds, 369  ;  on  the  pugnacity  .  of 
male  fine-plumaged  birds,  400  ;  on 
the  courtship  of  birds,  401  ;  on  the 
finding  of  new  mates  by  Peregrine- 
falcons  and  Kestrels,  408;  on  the 
bullfinch  and  starling,  408  ;  on  the 
cause  of  birds  remaining  unpaired. 


409  ;  on  starlings  and  i>arrots  living 
in  triplets,  409  ;  on  recognition  of 
colour  by  birds,  411  ;  on  hybrid 
birds,  414 ;  on  the  selection  of  a 
greenfinch  by  a  female  canary,  415  ; 
on  a  case  of  rivalry  of  female  bull- 
finches, 420  ;  on  the  maturity  of 
the  golden-pheasant,  483. 

Weisbach,  Dr.,  measurement  of  men 
of  different  races,  167  ;  on  the 
greater  variability  of  men  than  of 
women,  223 ;  on  the  relative  pro- 
portions of  the  body  in  the  sexes  of 
different  races  of  man,  559, 

Weismann,  Prof.,  colours  of  Lyccencc, 
312. 

Welcker,  M.,  on  brachycephaly  and 
dolichocephaly,  56  ;  on  sexual  dif- 
ferences in  the  skull  in  man,  557. 

Wells,  Dr.,  on  the  immunity  of  co- 
loured races  from  certain  poisons, 
193. 

Westring,  on  the  stridulation  of  males 
of  IVieridion,  273 ;  on  the  stridu- 
lation of  Heduviiis  perso7iatus,  281 ; 
on  the  stridulation  of  beetles,  302  ; 
on  the  stridulation  of  Omaloplia 
brunnea,  303  ;  on  the  stridulating 
organs  of  the  Coieoptera,  304 ; 
on  sounds  produced  by  Cychnts, 
304. 

Westropp,  H.  M.,  on  reason  in  a  bear, 
76 ;  on  the  prevalence  of  certain 
forms  of  ornamentation,  179. 

Westwood,  J.  0.,  on  the  classification 
of  the  Hymenoptera,  148  ;  on  the 
Culicidse  and  Tabanidae,  208  ;  on  a 
Hymenopterous  parasite  with  a 
sedentary  male,  221;  on  the  pro- 
portions of  the  sexes  in  Lttcanus 
cervus  and  Siajonium,  253 ;  on  the 
absence  of  ocelli  in  female  mu- 
tillidae,  274 ;  on  the  jaws  of  .4m- 
mophila,  275;  on  the  copulation  of 
insects  of  distinct  species,  275  ;  on 
the  male  of  Crabw  cribraritts,  276  ; 
on  the  pugnacity  of  male  'Jipuloe, 
280  ;  on  the  stridulation  of  Pirates 
striduhts,  281 ;  on  the  Cicada;,  281 ; 
on  the  stridulating  organs  of  the 
crickets,  284 ;  on  IJp/tippiger  litium, 
284,  288 ;  on  Pneumora,  287  ;  on 
the  pugnacity  of  the  Mantides,  289  ; 
on  Platyblemnus,  289  ;  on  difference 
in  the  sexes  of  the  Agrionidoe,  290; 


INDEX. 


WOODPECKERS. 


6^7 


ou  the  pugnacity  of  the  males  of  a 
species  of  Tenthredinfe,  291  ;  on 
the  pugnacity  of  the  male  stag- 
beetle,  299  ;  on  Bledius  taurus  and 
Siagonium,  299  ;  .  on  lamellicorn 
beetles,  301 ;  on  the  coloration  of 
Lithosia,  314. 

Whale,  Sperm-,  battles  of  male,  500. 

Whales,  nakedness  of,  56. 

Whately,  Archb.,  language  not  pe- 
culiar to  man,  84  ;  on  the  primitive 
civilisation  of  man,  143. 

Whewell,  Prof.,  on  maternal  affection, 
70. 

Whiskers,  in  monkeys,  150. 

White,  F.  B.,  noise  produced  by  Hylo- 
phila,  308. 

,  Gilbert,  on  the  proportion  of 
the  sexes  in  the  partridge,  248  ;  on 
the  house-cricket,  283  ;  on  the  ob- 
ject of  the  song  of  birds,  369 ;  on 
the  finding  of  new  mates  by  white 
owls,  408  ;  on  spring  coveys  of  male 
partridges,  409. 

Whiteness,  a  sexual  ornament  in 
some  birds,  494;  of  mammals  in- 
habiting snowy  countries,  542. 

White-throat,  aerial  love-dance  of  the 
male,  380. 

Whitney,  Prof.,  on  the  development 
of  language,  86 ;  language  not  in- 
dispensable for  thought,  88. 

Widow-bird,  polygamous,  219  ;  breed- 
ing plumage  of  the  male,  392, 
403  ;  female,  rejecting  the  un- 
adorned male,  419. 

Widows  and  widowers,  mortality  of, 
140. 

Wigeon,  pairing  with  a  pintail  duck, 
414. 

Wilckens,  Dr.,  on  the  modification  of 
domestic  animals  in  mountainous 
regions,  35 ;  on  a  numerical  re- 
lation between  the  hairs  and  excre- 
tory pores  in  sheep,  198. 

Wilder,  Dr.  Burt,  on  the  greater  fre- 
quency of  supernumerary  digits  in 
men  than  in  women,  223. 

Williams,  on  the  marriage-customs  ot 
the  Fijians,  599. 

Wilson,  Dr.,  on  the  conical  heads  of 
the  natives  of  North-Western  Ame- 
rica, 583 ;  on  the  Fijians,  583  ;  on 
the  persistence  of  the  fashion  of 
compressing  the  skull,  584. 


Wing-spurs,  449. 

Wings,  dilleronces  of,  in  the  two  sexes 
of  butterflies  and  Hymenoptera, 
277  ;  plav  of.  in  the  courtship  oi 
birds,  40''l. 

Winter,  change  of  colour  of  mam- 
mals in,  542. 

Witchcraft,  96. 

Wives,  traces  of  the  forcible  capture 
of,  144. 

Wolf,  winter  change  of  the,  542. 

Wolff,  on  the  variability  of  the  vis- 
cera in  man,  27. 

Wollaston,  T.  V.,  on  Eurygnathus, 
277  ;  on  musical  Curculionidge,  301  ; 
on  the  stridulation  of  Acalles,  306, 

Wolves  learning  to  bark  from  dogs, 
73  ;  hunting  in  packs,  101. 

,  black,  540. 

Wombat,  black  varieties  of  the,  539. 

Women  distinguished  from  men  by 
male  monkeys,  8  ;  preponderance  of, 
in  numbers,  244 ;  selection  of,  for 
beauty,  397  ;  etlects  of  selection 
of,  in  accordance  with  different 
standards  of  beauty,  584 ;  practice 
of  capturing,  589,  592 ;  early  be- 
trothals and  slavery  of,  593 ;  free- 
dom of  selection  by,  in  savage 
tribes,  598. 

Wonder,  manifestations  of,  by  ani- 
mals, 71. 

Wonfor,  Mr.,  on  sexual  peculiarities 
in  the  wings  of  butterflies,  277. 

Wood,  J.,  on  muscular  variations  m 
man,  26,  41,  43 ;  on  the  greater 
variability  of  the  muscles  in  men 
than  in  women,  223. 

,  T.  W.,  on  the  colouring  of  the 

orange-tip  butterfly,  313  ;  on  the 
habits  of  the  Saturniidoe,  315; 
quarrels  of  chamfeleons,  357 ;  on 
the  habits  of  Menura  Alberti,  371 ; 
on  Tetrao  cupido,  371 ;  on  the  dis- 
play of  plumage  by  male  pheasants, 
396  ;  on  the  ocellated  spots  of  the 
Argus  pheasant,  441 ;  on  the  habits 
of  the  female  cassowary,  478. 

Woodcock,  coloration  of  the,  491. 

Woodpecker,  selection  of  a  mate  by 
the  female,  416. 

Woodpeckers,  371  ;  tapping  of,  376; 
colours  and  nidification  of  the,  455, 
458,  489  ;  characters  of  voung,  465, 
474,  481. 


688 


INDEX. 


zyGvENida:. 


Wcolner,  Mr.,  observations  on  the 
ear  in  man,  15. 

Wormald,  Mr.,  on  the  coloration  of 
Hiipopyra,  315. 

Wounds,  healing  of,  8. 

Wren,  473;  young  of  the,  481. 

Wright,  C.  A.,  on  the  young  of  Oro- 
ceies  and  Peti'ocincla^  487. 

,   Chauncey,    great   brain-power 

requisite  for  language,  48 ;  on  cor- 
relative acquisition,  571;  on  the 
enlargement  of  the  brain  in  man, 
610, 

,  Mr.,  on  the  Scotch  deerhound, 

517  ;  on  sexual  preference  in  dogs, 
524;  on  the  rejection  of  a  horse 
by  a  mare,  525. 

,  W.  von,  on  the  protective  phi- 
mage  of  the  Ptarmigan,  391. 

Writing,  144. 

Wyman,  Prof.,  on  the  prolongation  of 
the  coccyx  in  the  human  embryo, 
11;  on  the  condition  of  the  great 
toe  in  the  human  embryo,  11 ;  on 
the  occurrence  of  the  supra-condy- 
loid  foramen  in  the  humerus  of 
man,  22  ;  on  variation  in  the  skulls 
of  the  natives  of  the  Sandwich 
Islands,  26  ;  on  the  hatching  of  the 
eggs  in  the  mouths  and  branchial 
cavities  of  male  fishes,  163,  345. 


X. 

Xenarchus,  on  the  Cicadse,  281. 

Xenophon,  selection  in  mankind  advo- 
cated by,  29. 

XenoHiynchus,  sexual  difference  in  the 
colour  of  the  eyes  in,  425. 

Xip'iophorus  Hellerii,  peculiar  anal  fin 
of  the  male,  337,  338. 

Xylocopa,  difference  of  the  sexes  in, 
292. 


Yarrell,  W.,  on  the  habits  of  the 
Cyprinidae,  250;  on  Raia  clavata, 
331 ;  on  the  characters  of  the  male 
salmon  during  the  breeding  season, 
332,  342  ;  on  the  characters  of  the 
rays,  334;  on  the  gemmeous  drago- 
net,  336  ;  on  the  spawning  of  the 
salmon,  344;  on  the  incubation  of 
the  Lophobranchii,  346  ;  on  rivalry 
in  song-birds,  369 ;  on  the  trachea 
of  the  swan,  374  ;  on  the  moulting 
of  the  AnatidfE,  393  ;  on  the  young 
of  the  waders,  486. 

Yellow  fever,  immunity  of  negroes 
and  mulattoes  from,  193. 

Youatt,  Mr.,  on  the  development  of 
the  horns  in  cattle,  235. 

Yura-caras,  their  notions  of  beauty, 
580. 


Z. 

Zebra,  rejection  of  an  ass  by  a  female, 

540  ;  stripes  of  the,  645 
Zebus,  humps  of,  532. 
Zigzags,  prevalence  of,  as  ornaments, 

179. 
Zincke,  Mr.,  on  European  emigration 

to  America,  142. 
Zootoca  vivipara,  sexual  difference  in 

the  colour  of,  357. 
Zouteveen,   Dr.,    polydactylism,    37 ; 

proportion  of  sexes  at  Cape  of  Good 

Hope,    243 ;    spiders  attracted   by 

music,  273  ;  on  sounds  produced  bv 

fish,  347. 
Zygainidtc,  coloration  of  the,  314. 


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849  &  S51   Broadway,  N.  Y. 


THE    WORKS    OF 


Prof.  JOHN  TYNDALL,  LLD.,  F.R.S. 


V. 

HOURS  OF   EXERCISE    IN   THE   ALPS. 

One  vol.,  i2mo.     With  Illustrations.     Cloth,  $2.00. 

"  The  present  volume  is  for  the  most  part  a  record  of  bodily  action,  written  partly  to 
preserve  to  myself  the  memory  of  strong  and  joyous  hours,  and  partly  for  the  pleasure  of 
those  who  find  exhilaration  in  descriptions  associated  with  mountain-life." — From  Author's 
Preface. 

VI. 
FARADAY    AS    A    DISCOVERER. 

One  vol.,  i2mo.     Cloth,  $1.00. 

"  It  has  been  thought  tlesirable  to  give  you  and  the  world  some  image  of  Michael 

Faraday  as  a  scientific  investigator  and  discoverer I  have 

returned  from  my  task  with  such  results  as  I  could  gather,  and  also  with  the  wish  that 
these  results  were  more  worthy  than  they  are  of  the  greatness  of  my  theme." — The 
Author. 

VII. 

FORMS  OF  WATER,  IN   CLOUDS,   RAIN,   RIVERS,   ICE, 
AND  GLACIERS. 

This  is  the  first  volume  of  the  International  Scientific  Series,  and  is  a  valu- 
able  and  interesting  work.     One  vol.,  i2mo.     Cloth,  $1.50. 

VIII. 

CONTRIBUTIONS   TO   MOLECULAR   PHYSICS   IN   THE 
DOMAIN    OF   RADIANT    HEAT. 

A  Series  of  Memoirs  published  in  the  **  Philosophical  Transactions  "  and 
*'  Philosophical  Magazine."     With  Additions. 

D.  APPLETON  Si  CO.,  Publishers, 

B49  &  551   Broadway,   N.  Y. 


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