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LIBRARY 

OF   THK 

UNIVERSITY  OF  CALIFORNIA. 


GIKT  OK 


Accession  86461>        class 


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The  Contemporary  Science  Series. 

Edited  by  Havelock  Ellis. 


I.  THE  EVOLUTION  OF  SEX.     By  Prof.  PATRICK  GEDDES 

and  J.  A.  THOMSON.    With  90  Illustrations.    Third  Edition.     35.  6d. 

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TUNZELMANN.     With  88  Illustrations.     33.  6d. 

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TAYLOR.     Illustrated.     Second  Edition.     35.  6d. 

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account  to  which  he  can  turn  his  exceptionally  wide  and  varied  information.  .  .  . 
Masterly  and  exhaustive." — Pall  Mall  Gazette. 

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GAZZA.     Illustrated.     Second  Edition.     35.  6d. 

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conquest,  encroachment,  and  the  heavy  hand  of  Roman  law." — Scottish  Leader. 

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Revised  and  Enlarged.     33.  6d. 

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XIII.  THE   EVOLUTION   OF   MARRIAGE.     By  Professor 
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SIMS  WOODHEAD.     Illustrated.     Third  Edition.     35.  6d. 
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35.  6d. 

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and  the  educational  question  any  one  has  carried  forward  the  new  doctrine  so  boldly 
to  its  extreme  logical  consequence."—  Professor  SULLY  in  Mind. 

XVI.  THE  MAN  OF  GENIUS.     By  Prof.  LOMBROSO.     Illus- 
trated.   35.  6d. 

"  By  far  the  most  comprehensive  and  fascinating  collection  of  facts  and  generalisa- 
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Science. 

XVII.  THE   HISTORY  OF  THE    EUROPEAN   FAUNA. 
By  R.  F.  SCHARFF,  B.Sc.,  PH.D.,  F.Z.S.     6s. 

XVIII.  PROPERTY:  ITS  ORIGIN  AND  DEVELOPMENT. 

By  CH.  LETOURNEAU,  General  Secretary  to  the  Anthropological 
Society,  Paris,  and  Professor  in  the  School  of  Anthropology,  Paris. 
33.  6d. 

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interpreted  his  facts  with  considerable  judgment  and  learning."—  Westminster 
Jteciew. 


XIX.  VOLCANOES,     PAST    AND    PRESENT.      By    Prof. 
EDWARD  HULL,  LL.D.,  F.R.S.     33.  6d. 

"  A  very  readable  account  of  the  phenomena  of  volcanoes  and  earthquakes." — Nature. 

XX.  PUBLIC   HEALTH.      By   Dr.    J.    F.    J,   SYKES.      With 

numerous  Illustrations.     35.  6d. 

"Not  by  any  means  a  mere  compilation  or  a  dry  record  of  details  and  statistics, 
but  it  takes  up  essential  points  in  evolution,  environment,  prophylaxis,  and  sanitation 
bearing  upon  the  preservation  of  public  health." — Lancet. 

XXI.  MODERN   METEOROLOGY.     AN   ACCOUNT   OF  THE 
GROWTH    AND    PRESENT    CONDITION    OF    SOME    BRANCHES    OF 
METEOROLOGICAL  SCIENCE.     By  FRANK  WALDO,  PH.D.,   Member 
of  the   German  and   Austrian    Meteorological    Societies,    etc. ;    late 
Junior   Professor,    Signal    Service,    U.S.A.      With    112   Illustrations. 
33.  6d. 

"  The  present  volume  is  the  best  on  the  subject  for  general  use  that  we  have  seen." 
— Daily  Telegraph  (London). 

XXII.  THE  GERM-PLASM :  A  THEORY  OF  HEREDITY. 

By  AUGUST  WEISMANN,  Professor  in  the  University  of  Freiburg-in- 

Breisgau.     With  24  Illustrations.     6s. 

"There  has  been  no  work  published  since  Darwin's  own  books  which  has  so 
thoroughly  handled  the  matter  treated  by  him,  or  has  done  so  much  to  place  in  order 
and  clearness  the  immense  complexity  of  the  factors  of  heredity,  or,  lastly,  has 
brought  to  li^ht  so  many  new  facts  and  considerations  bearing  on  the  subject."— 
British  Medical  Journal. 

XXIII.  INDUSTRIES    OF    ANIMALS.     By    F.    HOUSSAY. 

With  numerous  Illustrations.     35.  6d. 

"  His  accuracy  is  undoubted,  yet  his  facts  out-marvel  all  romance.  These  facts  are 
here  made  use  of  as  materials  wherewith  to  form  the  mighty  fabric  of  evolution." 
— Manchester  Guardian. 

XXIV.  MAN  AND  WOMAN.     By  HAVELOCK  ELLIS.     Illus- 
trated.    Second  Edition.     6s. 

"Mr.  Havelock  Ellis  belongs,  in  some  measure,  to  the  continental  school  of  anthro- 
pologists ;  but  while  equally  methodical  in  the  collection  of  facts,  he  is  far  more 
cautious  in  the  invention  of  theories,  and  he  has  the  further  distinction  of  being  not 
only  able  to  think,  but  able  to  write.  His  book  is  a  sane  and  impartial  considera- 
tion, from  a  psychological  and  anthropological  point  of  view,  of  a  subject  which  is 
certainly  of  primary  interest." — Athenaeum. 

XXV.  THE   EVOLUTION   OF    MODERN    CAPITALISM. 

By  JOHN  A.  HOBSON.  M.A.     35.  6d. 

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conscientious  as  it  is  acute,  a  keen  sense  of  the  importance  of  certain  points  as  to 
which  economists  of  all  schools  have  hitherto  been  confused  and  careless,  and  an 
impartiality  generally  so  great  as  to  give  no  indication  of  his  [Mr.  Hobson's]  per- 
sonal sympathies."— Pall  Mall  Gazette. 

XXVI.  APPARITIONS  AND  THOUGHT-TRANSFERENCE. 

By  FRANK  PODMORE,  M.A.     35.  6d. 

"A  very  sober  and  interesting  little  book.  .  .  .  That  thought-transference  is  a 
veal  thing,  though  not  perhaps  a  ver>  common  thing,  he  certainly  shows." — Spectator. 

XXVII.  AN    INTRODUCTION    TO   COMPARATIVE 
PSYCHOLOGY.     By  Professor  C.  LLOYD  MORGAN,  F.R.S.     With 
Diagrams.     6s. 

as  it  takes  shape  in  a  mind 
Tell  written,  extremely  entertain- 


XXVIII.  THE   ORIGINS   OF   INVENTION:  A  STUDY  OF 
INDUSTRY    AMONG    PRIMITIVE    PEOPLES.      By    OTIS    T.    MASON, 
Curator  of  the  Department  of  Ethnology  in  the  United  States  National 
Museum.     35.  6d. 

"A  valuable  history  of  the  development  of  the  inventive  faculty."— Nature. 

XXIX.  THE   GROWTH   OF   THE   BRAIN:    A   STUDY  OF 
THE  NERVOUS  SYSTEM  IN  RELATION  TO  EDUCATION.     By  HENRY 
HERBERT  DONALDSON,  Professor  of  Neurology  in  the  University  of 
Chicago.     33.  6d. 

"  We  can  say  with  confidence  that  Professor  Donaldson  has  executed  his  work  with 
much  care,  judgment,  and  discrimination."—  The  Lancet. 

XXX.  EVOLUTION    IN    ART:    As    ILLUSTRATED    BY   THE 
LIFE-HISTORIES  OF  DESIGNS.     By  Professor  ALFRED  C.  HADDON, 
F.R.S.     With  130  Illustrations.     6s. 

"It  is  impossible  to  speak  too  highly  of  this  most  unassuming  and  invaluable 
book."— Journal  Anthropological  Institute. 

XXXI.  THE  PSYCHOLOGY  OF   THE  EMOTIONS.     By 

TH.  RIBOT,  Professor  at  the  College  of  France,  Editor  of  the  Revue 
Philosophique.     6s. 

"Charmingly  written,  and  full  of  lucid  explanation  and  brilliant  comparison.  A 
masterly  exposition."— British  Medical  Journal. 

XXXII.  HALLUCINATIONS  AND  ILLUSIONS :  A  STUDY 
OF  THE  FALLACIES  OF  PERCEPTION.     By  EDMUND  PARISH.     6s. 

"  The  most  comprehensive  and  most  scientific  work  on  false  perception  that  has  up 
till  now  been  written  in  any  language." — Journal  of  Mental  Science. 

XXXIII.  THE  NEW  PSYCHOLOGY.     By  E.  W.  SCRIPTURE, 

Ph.D.  (Leipzig).     With  124  Illustrations.     6s. 

"  We  have  at  present  no  work  in  English  which  gives  in  so  compact  a  form  so 
comprehensive  a  view  of  the  subject." — Liverpool  Post. 

XXXIV.  SLEEP  :  ITS  PHYSIOLOGY,  PATHOLOGY,  HYGIENE,  AND 
PSYCHOLOGY.     By  MARIE  DE  MANACEINE  (St.  Petersburg).     Illus- 
trated.    35.  6d. 

"The  book  is  a  complete  and  wonderfully  interesting  exposition,  and  as  such 
ought  to  receive  a  hearty  welcome." — Scotsman. 

XXXV.  THE    NATURAL    HISTORY    OF    DIGESTION. 
By  A.   LOCKHART  GILLESPIE,    M.D.,   F.R.C.P.   ED.,   F.R.S.   ED. 
With  a  large  number  of  Illustrations  and  Diagrams.     6s. 

"Dr.  Gillespie's  work  is  one  that  has  been  greatly  needed.  No  comprehensive 
collation  of  this  kind  exists  in  recent  English  literature.  All  the  important  woi  k 
that  has  appeared  within  the  past  few  years  is  discussed  so  far  as  the  limits  of  the 
book  allow  of  discussion,  and  extremely  little  of  value  has  been  omitted.  Not  least 
interesting  are  the  accounts  of  the  author's  own  original  work." — American  Journal 
of  the  Medical  Sciences. 

XXXVI.  DEGENERACY:  ITS  CAUSES,  SIGNS,  AND  RESULTS. 
By  Professor    EUGENE  S.    TALBOT,    M.D.,    Chicago.     With   Illus- 
trations.    6s. 

"  The  author  is  bold,  original,  and  suggestive,  and  his  work  is  a  contribution  of 
real  and  indeed  great  value,  more  so  on  the  whole  than  anything  that  has  yet 
appeared  in  this  country."— American  Journal  of  Psychology. 

XXXVII.  THE  RACES  OF  MAN.     By  J.  DENIKER.     6s. 

XXXVIII.  THE     PSYCHOLOGY     OF     RELIGION.      AN 
EMPIRICAL  STUDY  OF  THE  GROWTH  OF   RELIGIOUS  CONSCIOUS- 
NESS.    By  EDWIN  DILLER  STARBUCK,   PH.D.,   Assistant  Professor 
of  Education,  Leland  Stanford  Junior  University.     6s. 


THE  CONTEMPORAR  V  SCIENCE  SERIES. 


EDITED  BY  HAVELOCK  ELLIS. 


THE  HISTORY  OF  THE  EUROPEAN  FAUNA 


THE   HISTORY  OF  THE 
EUROPEAN    FAUNA 


BY 


R.  F.tSCHARFF,  B.Sc.,  PH.D.,  F.Z.S. 

Keeper  of  the  Natural  History  Collections,  Science  and  Art  Museum  t 

Dublin ;  Member  of  the  Royal  Irish  Academy ;  Corresponding 

Member  of  the  Senckenbergische  Naturforschende 

Gcsellschaft. 


WITH    ILLUSTRATIONS. 


LONDON 
WALTER    SCOTT,    LIMITED 

PATERNOSTER  SQUARE 
1899 


CONTENTS. 


CHAPTER  I. 

PAGE 

INTRODUCTION      ..................  1-36 

CHAPTER    II. 
PRELIMINARY  CONSIDERATIONS  .........  37~88 

CHAPTER  III. 

THE   FAUNA  OF   BRITAIN  ...  .........         89-13! 

CHAPTER  IV. 

THE  ARCTIC   FAUNA          .............  ..      132-188 

CHAPTER  V. 

THE  SIBERIAN   MIGRATION          ............      189-244 

CHAPTER  VI. 

THE  ORIENTAL  MIGRATION        ...          .  ........      245-286 

CHAPTER  VII. 

THE  LUSITANIAN   FAUNA  ............      287-308 

CHAPTER  VIII. 

THE  ALPINE  FAUNA         ...          •  ............ 


BIBLIOGRAPHY  ...............      35^354 

INDEX  ..................      355-364 

86461 


PREFACE. 


OUR  knowledge  of  the  present  and  past  fauna  of 
Europe  is  as  yet  insufficient  to  indicate  with  precision 
the  original  homes  of  its  component  elements,  but  I 
hope  that  the  lines  of  research  laid  down  here,  and 
the  method  of  treatment  adopted,  will  aid  zoologists 
and  geologists  in  collecting  materials  for  a  more  com- 
prehensive study  of  the  history  of  our  animals.  I 
trust  also  that  a  fresh  impulse  will  be  given  by  the 
publication  of  this  book  to  the  study  of  the  Geo- 
graphical Distribution  of  Species.  Collectors  of 
Beetles,  Butterflies,  Shells,  and  Fossils  may  derive 
some  useful  hints  by  its  perusal  and  thus  direct  their 
studies,  so  as  to  add,  by  accuracy  in  observation, 
to  our  knowledge  of  the  former  geographical  re- 
volutions which  have  moulded  our  islands  and  con- 
tinents. To  geographers,  a  survey  of  some  of  the 
more  important  changes  in  the  distribution  of  land 
and  water  in  past  times — based  upon  the  composi- 
tion of  our  fauna — will  be  interesting.  The  subject, 
however,  is  a  complex  one.  I  have  ventured  to 
indicate  a  suitable  method  of  treatment,  and  as  such 
this  attempt  to  elucidate  the  history  of  the  European 
fauna  should  be  received. 


VI  PREFACE. 

This  work  was  written  as  the  outcome  of  a  paper 
published  in  the  Proceedings  of  the  Royal  Irish 
Academy  (3rd  series,  vol.  iv.,  1897),  "On  the  Origin 
of  the  European  Fauna."  A  summary  of  that  paper 
appeared  in  Nature  (vol.  Ivi.,  1897),  and  fuller  extracts 
of  more  important  parts,  with  some  criticisms,  in  the 
Geological  Magazine  (N.S.,  sec.  iv.,  vol.  iv.,  1897).  I 
freely  acknowledge  the  value  of  these  criticisms, 
which  have  largely  assisted  me  to  amplify  and  to 
improve  upon  the  ideas  laid  down  in  the  paper. 

I  have  found  that  it  greatly  facilitates  comprehension 
of  the  arguments  used,  to  give  a  few  maps  indicating 
in  a  general  way  the  extent  of  former  seas  and 
continents.  I  may  in  this  way,  as  Mr.  Kendall 
has  pointed  out,  have  submerged  many  square 
miles  of  land  which  had  never  been  covered  by  the 
sea, — at  least  not  within  recent  geological  times, — 
but  the  maps  were  intended  as  illustrations  of  my 
views  in  a  broad  spirit  only. 

Some  zoologists  may  be  surprised  that,  in  some 
cases,  I  have  not  followed  the  latest  views  in  revised 
nomenclature.  I  felt  that  in  a  work  of  this  kind 
it  was  of  supreme  importance  to  employ  names  still 
current  in  our  leading  text-books,  such  as  Lepus 
variabilis  for  the  Mountain  Hare,  instead  of  Lefius 
timidus.  After  each  chapter  I  have  endeavoured  to 
give  a  short  summary  of  contents,  while  a  biblio- 
graphy of  the  principal  works  and  papers  consulted 
will  be  found  at  the  end.  I  should  also  acknowledge 
the  aid  which  I  have  received  from  such  excellent 


PREFACE.  Vli 

works  of  reference  as  the  British  Museum  Catalogues 
of  Birds,  by  Dr.  Bowdler  Sharpe,  and  those  of  Reptiles, 
Amphibia,  and  Fishes,  by  Dr.  Giinther  and  Dr. 
Boulenger.  The  valuable  works  en  Mammalia  by 
Sir  W.  Flower,  Mr.  Lydekker,  Mr.  Grev£,  and  Dr. 
Trouessart,  were  indispensable  to  me. 

To  Sir  William  Flower,  Mr.  Lydekker,  Professor 
Sars,  and  Professor  Smitt,  I  am  especially  indebted 
for  allowing  me  to  reproduce  drawings  from  their 
works,  and  to  my  friend  Mr.  Welch  for  some  beauti- 
ful photographs.  The  Council  of  the  Royal  Irish 
Academy  also  kindly  gave  me  permission  to  reprint 
the  maps  used  in  illustration  of  my  paper.  Professor 
Haddon  first  suggested  my  writing  this  book,  and 
gave  me  many  useful  hints ;  and  great  assistance  was 
rendered  me  by  my  colleague,  Mr.  G.  H.  Carpenter, 
in  revising  the  proofs.  To  both  of  these  kind  friends 
I  desire  to  acknowledge  my  deep  sense  of  gratitude. 

R.  F.  SCHARFF. 


THE   HISTORY   OF  THE 
EUROPEAN    FAUNA. 


CHAPTER   I. 

INTRODUCTION. 

/  EVERY  student  of  natural  history,  whether  he  be 
'  interested  in  birds,  butterflies,  or  shells,  contributes 
his  share  of  facts  which  help  to  show  how  the  fauna 
of  his  country  has  originated.  The  capture  of  a 
Swallow-tail  or  of  a  Marbled  White  Butterfly  in  Eng- 
land at  once  furnishes  material  for  reflection  as  to  the 
reason  of  its  absence  from  Scotland  and  Ireland. 
Why  should  the  Nightingale  allow  its  beautiful  song 
to  be  heard  in  England,  and  never  stray  across 
the  Channel  to  the  sister  isle  or  cross  the  borders  of 
North  Britain?  Lovers  of  bird-life  and  sportsmen, 
who  have  observed  the  habits  of  the  Ptarmigan  in  the 
wild  mountain  recesses  of  Scotland,  are  aware  that 
nowhere  else  in  the  British  Islands  do  we  meet  with 
this  interesting  member  of  the  grouse  family,  and 
many  no  doubt  have  allowed  their  minds  to  dwell 
upon  the  causes  of  its  singularly  local  distribution. 


2  HISTORY  OF   THE   EUROPEAN    FAUNA. 

All  these  animals  have  a  wide  range  in  other  parts 
of  the  world.  In  past  times,  before  man  began  to 
make  observations  on  the  geographical  distribution  of 
birds  and  butterflies,  or  even  before  the  appearance  of 
man  in  Northern  Europe,  they  may  have  lived  all 
over  the  British  Islands.  For  some  reason  or  other 
they  are  perhaps  dying  out  or  withdrawing  towards 
their  original  home,  which  may  either  be  northward, 
or  to  the  east  or  south.  If  we  had  some  clue  as  to 
their  former  history  from  fossil  evidence — or,  in  other 
words,  if  their  remains  had  been  preserved  to  us  in 
geological  deposits, — we  should  have  less  difficulty  in 
deciding  this  problem.  But  butterflies  are  scarcely 
ever  preserved  in  a  fossil  state,  and  birds  very  rarely. 
We  know  little  or  nothing,  therefore,  of  their  past 
history  from  direct  evidence,  and  are  obliged  to  trust 
to  indirect  methods  of  research  which  will  be  indi- 
cated later  on. 

X^Mammals  and  Snails  tell  us  their  story  more  plainly. 
*  The  bones  of  the  former  and  the  shells  of  snails  are 
easily  preserved,  and  thus  furnish  us  with  the  neces- 
sary data  as  to  their  past  history,  for  we  find  them 
abundantly  in  most  of  the  recent  geological  deposits. 
Among  the  mammals  of  the  British  Islands  there 
are  some  instances  of  distribution  which  much 
resemble  those  I  have  quoted.  Thus  the  Arctic  Hare 
(Lepus  variabilis]  is  in  the  British  Islands  confined  to 
Ireland  and  to  the  mountains  of  Scotland  ;  and  if  it 
were  not  for  the  fact  that  its  bones  have  been  dis- 
covered in  a  cave  in  the  south-west  of  England,  we 


INTRODUCTION.  3 

should  perhaps  never  have  known  that,  formerly,  it 
must  have  inhabited  that  country  as  well.  Of  other 
mammals  we  possess  fossil  and  also  historical  evi- 
dence of  their  having  once  lived  in  these  islands. 
Such  are  the  Wolf  and  the  Wild  Boar,  both  of  which 
were  abundant  in  Great  Britain  and  Ireland.  The 
latter  is  a  distinctly  southern  species.  We  assume 
this,  because  its  remains  have  never  been  found  in 
high  northern  latitudes ;  nor  does  it  now  occur  in 
Northern  Europe  or  Northern  Asia,  whilst  all  its 
nearest  relatives  live  in  sub-tropical  or  tropical 
climates.  The  Arctic  Hare,  on  the  contrary,  has 
probably  come  to  us  from  the  north.  Its  remains 
are  unknown  even  in  Southern  Europe,  and  the 
more  we  approach  the  Arctic  Regions,  the  more 
abundant  it  becomes.  Thus  we  have  here  two 
instances  of  British  mammals,  one  of  which,  the 
Wild  Boar,  has  died  out — as  it  were  in  a  southerly 
direction  ;  whilst  the  other,  the  Arctic  Hare,  is  ap- 
parently retreating  towards  the  north. 

There  are  also  some  British  mammals  of  which  we 
have  no  fossil  history,  at  least  of  which  no  remains 
have  as  yet  been  found  in  these  islands.  Such  a 
one  is  the  Harvest  Mouse  (Mus  minutus).  It  has 
a  somewhat  restricted  range  in  England,  and  only 
just  crosses  the  Scottish  border  in  the  east.  From 
the  rest  of  Scotland  and  from  the  whole  of  Ireland 
it  is  absent.  To  judge  from  this  distribution,  in 
connection  with  the  fact  of  its  being  unknown  as  a 
British  fossil  species,  it  is  probably  a  late  immigrant 


4  HISTORY   OF   THE    EUROPEAN    FAUNA 

to  England,  and  has  not  had  time  to  spread,  through- 
out Scotland  at  any  rate.  But  it  is  also  absent 
from  Scandinavia,  from  the  Spanish  peninsula,  from 
almost  the  whole  of  Italy  and  the  Alps,  as  also  from 
the  Mediterranean  Islands,  whilst  the  little  mouse 
occurs  abundantly  right  across  Siberia.  We  shall 
learn  more  about  centres  of  dispersion  later  on ; 
meanwhile  I  should  mention  that  such  a  distribution 
indicates  that  the  Harvest  Mouse  has  most  likely 
originated  in  the  east,  and  has  spread  from  there 
westward  in  recent  geological  times. 

Conchologists  have  long  ago  been  acquainted  with 
the  fact  that  many  molluscs,  for  example  the  so-called 
"Stone-cutter"  Snail  (Helix lapicida)  and  the  "Cheese 
Snail "  (Helix  obvoluta),  have  a  very  restricted  range 
in  the  British  Islands.  Both  are  entirely  absent 
from  Scotland  and  Ireland,  the  Cheese  Snail  being 
confined  to  South-eastern  England.  The  Stone- 
cutter has  rather  a  wider  range,  is  even  known  from 
a  Welsh  locality,  and  is  met  with  as  far  north  as 
Yorkshire.  Their  distribution  would  indicate,  there- 
fore, that  while  both  are  recent  immigrants,  the 
Cheese  Snail  is  probably  the  last  comer.  This 
supposition  is  in  so  far  supported  by  fossil  evidence, 
as  the  latter  is  unknown  in  the  fossil  state,  whilst  the 
Stone-cutter  has  been  described  by  Messrs.  Kennard 
and  Woodward  (p.  243) a  as  occurring  in  the  cave 
deposit  known  as  the  Ichtham  fissure,  and  also  from 

1  The  numbers  in  brackets  throughout  this  work  refer  to  the  page- 
number  in  the  Bibliography  at  the  end. 


INTRODUCTION.  5 

several  English  pleistocene  and  holocene  deposits. 
The  Stone-cutter  can  scarcely  be  looked  upon  as  a  very 
recent  immigrant  in  the  light  of  this  evidence,  though 
we  have  no  proof  of  its  having  ever  had  a  much 
wider  range  in  the  British  Islands  than  it  has  to-day. 

Among  the  lichens,  which  so  abundantly  cover  the 
rocks  and  trees  in  South-western  Ireland,  and  which 
impart  such  a  characteristic  feature  to  the  scenery, 
we  find  a  beautifully  spotted  slug  (Geomalacus  macu- 
losus}.1  It  is  a  stranger  to  the  rest  of  the  British 
Islands,  and  indeed  occurs  nowhere  else  in  Northern 
Europe.  We  have  to  travel  as  far  as  Northern 
Portugal  before  we  again  meet  with  it,  and  it  is 
there  also  that  its  nearest  relations  live. 

Many  more  similar  examples  might  be  quoted,  but 
enough,  I  think,  has  been  said  to  show  that  the 
British  fauna  is  made  up  of  several  elements  whose 
original  homes  may  lie  widely  apart  and  in  different 
directions.  We  have  fossil  evidence  that  some  of 
the  northern  species,  and  also  a  few  of  the  southern 
ones,  have  become  extinct  within  comparatively 
recent  times ;  others  are  apparently  on  the  verge  of 
extinction,  whilst  many  not  only  maintain  their 
position  in  the  constant  struggle  for  existence,  but 
are  even  extending  their  range. 
/X^The  problem  of  tracing  the  origin  of  the  British 
fauna,  or  at  least  that  of  some  of  the  more 
characteristic  members  of  every. section  or  element, 

1  A  map  giving  its  exact  distribution  in  Ireland  will  be  found  on 
p.  300,  and  a  figure  of  the  slug  on  p.  298. 


6  HISTORY  OF   THE   EUROPEAN    FAUNA. 

appears  at  first  a  somewhat  difficult  task.     Indeed, 
the    means    of   dispersal    of   the    various    groups  of 
animals   are  so  different  that  it   occurred   to   me  it 
might    be   better    to   deal    with   the    mammals,    the 
birds,  the  reptiles,  and  so  forth,  all  separately.     This 
idea  I  have  attempted  to  follow  to  some  extent,  with 
most  satisfactory  results.      The  British  fauna  of  the 
present  day  is  no  doubt  complex,  but   no   more   so 
than  the  fauna  of  the  most  recent  of  our  geological 
deposits — the    Pleistocene.      However,   when    we   go 
back  still  further   and    look    at    the   earlier  Tertiary 
remains,  we  find  the  fauna  becoming  less  complex. 
Northern     species    disappear,    and    the    strata     are 
entirely  filled  with  the  remains  of  southern  animals 
and  plants.     Geologists  indeed  are  quite  unanimous 
in  their  belief,  that  the  fauna  of  the  British  Islands 
during  the  earlier  epochs  of  the  Tertiary  Era  was  a 
southern  one  ;   that  it  then   gradually   became  more 
temperate,  until  at  last,  in  more  recent  times,  decidedly 
northern   forms  invaded   the  country.      These   seem 
to  have  driven    out — to  some   extent    at    least — the 
southern     species;    but    more     recently    again,    the 
southerners,    reinforced    by   an    eastern     contingent, 
appear  to  have  gained   territory  and  are  advancing 
into  the  area  held  by  the  northerners.     The  eastern 
invasion  does  not  seem  to  have  affected  Ireland  at 
all,  and  we  find  the  country  there  divided  between  the 
southern  and  northern  animals.     We  can  thus  roughly 
construct  a   map  as   I   have   done  here,  showing,  by 
means  of  horizontal   and  sloping  lines,   the   principal 


INTRODUCTION. 


areas  inhabited  at  the  present  time  by  the  species  of 
northern,  southern,  and  eastern  origin  (Fig.  i). 


FIG.  i. — Map  of  the  British  Islands,  indicating  approximately  the 
areas  inhabited  by  the  northern,  southern,  and  eastern  animals. 
The  horizontal  lines  represent  the  areas  of  northern  species,  the 
sloping  lines  those  of  southern  and  eastern  ones. 

In  the  problems  which  are  being  discussed  in  this 
work  I  have  often  found  it  of  advantage,  in  order  to 


8  HISTORY  OF   THE   EUROPEAN    FAUNA. 

facilitate  the  comprehension  of  the  arguments  used,  to 
give  maps.  Some  of  these  represent  the  geographical 
conditions  at  the  particular  epoch  referred  to  in  the 
text,  but  they  merely  claim  to  give  a  general  idea. 
There  was  never  any  intention  to  make  them  corre- 
spond with  all  the  data  of  which  we  have  geological 
evidence.  They  are  what  I  might  call  "diagrammatic." 
In  comparing  them  with  reconstructions  of  former 
physical  geography  such  as  have  been  attempted  from 
time  to  time,  I  hope  geologists  will  therefore  deal 
leniently  with  the  faults  I  may  have  committed,  and 
remember  that  the  maps  are  "  impressions,"  or  "  dia- 
grams," and  not  faithful  representations  of  all  the 
geographical  revolutions  witnessed  by  some  of  our 
remote  forefathers  at  any  particular  period. 
r.  The  knowledge  we  gain  from  a  study  of  the  British 
Tertiary  deposits  enables  us  to  affirm  positively  that 
both  the  eastern  and  the  northern  species  arrived  in 
these  islands  comparatively  recently,  but  that  the 
southern  forms  must  have  migrated  northward  from 
the  Continent  long  ages  ago.  Since  the  northern 
and  the  eastern  migrations — that  is  to  say,  those 
coming  from  the  north  and  east — were  the  last  to 
arrive  in  Northern  Europe,  the  remains  of  the  animals 
contained  in  the  most  recent  deposits  of  that  portion 
of  our  continent  will  furnish  us  with  a  clue  as  to  the 
extent  of  the  area  inhabited  by  them.  This  is  not 
all,  however.  It  is  also  possible  to  discover  from  these 
remains  the  direction  which  the  animals  that  they 
belonged  to  came  from.  As  we  shall  learn  later  on, 


INTRODUCTION.  9 

a  migration  on  a  vast  scale  entered  Europe  during 
the  Pleistocene  epoch — the  most  recent  of  the  geo- 
logical epochs,  during  which  great  extensions  of 
glaciers  occurred  in  the  mountainous  regions  of 
Europe.  The  latter  period  is  known  to  us  as  the  Ice 
Age  or  Glacial  period.  This  will  be  described  more 
fully  in  Chapter  II.,  meanwhile  I  may  mention  that 
we  presume  that  this  migration  came  from  the  east, 
because  no  remains  of  the  members  of  that  particular 
fauna  are  known  from  Spain,  Southern  Italy,  Scandi- 
navia, Ireland,  or  from  the  Balkan  peninsula.  The 
number  of  species  evidently  belonging  to  this  same 
migration,  moreover,  become  fewer  as  we  proceed 
westward,  and  a  large  proportion  of  them  still  inhabit 
Northern  Asia,  though  most  of  them  are  now  extinct 
in  Europe.  After  having  thoroughly  studied  such  a 
recent  geological  migration,  we  learn  to  understand 
others  better,  though  the  more  ancient  they  are,  the 
fewer  are  the  traces  and  the  more  difficult  are  they  to 
follow. 

Then  again  we  have  to  take  into  consideration  the 
fact,  that  whilst  mammals,  particularly  the  larger 
herbivores,  are  forced  to  migrate  frequently  owing 
to  scarcity  of  food  or  temporary  changes  of  climate, 
many  of  the  invertebrates  remain  practically  unaffected 
by  either.  Most  of  our  land  mollusca,  for  instance, 
are  satisfied  with  meagre  provender,  and  stand  ex- 
tremes of  climate  well,  as  long  as  there  is  sufficient 
moisture.  As  a  result  of  their  peculiar  disposition, 
many  of  them,  no  doubt,  have  survived  through 


10  HISTORY  OF  THE   EUROPEAN   FAUNA, 

several  geological  epochs,  and  have  witnessed  vast 
geographical  revolutions  in  their  immediate  surround- 
ings, whilst  mammals  are  comparatively  short-lived. 
Being  driven  from  one  country  to  another,  and  ex- 
posed to  innumerable  enemies,  new  types  appear  and 
old  ones  rapidly  vanish  ;  in  fact,  there  are  almost 
constant  changes  in  the  mammalian  fauna  as  we 
pass  from  one  epoch  to  another. 

I  have  until  now  referred  more  particularly  to  the 
British  fauna  and  the  North  European  in  general, 
because  the  history  of  our  own  animals  interests 
us  all  more  than  those  of  any  other  European  area. 
It  is,  moreover,  preferable  to  commence  our  inves- 
tigations into  the  origin  of  the  European  fauna 
by  the  study  of  a  small  district.  This  should,  if 
possible,  be  an  island.  If  we  took  a  slice  of  the 
continent  like  France  or  Germany,  we  should  find 
the  problem  more  complex.  Instead  of  choosing 
the  British  Islands,  we  might,  however,  take  an  island 
like  Corsica  or  Sardinia.  In  either  of  these  we  should 
discover  peculiarities  in  the  composition  of  their  fauna 
precisely  similar  to  those  which  I  have  indicated  to 
be  present  in  the  British  fauna.  We  should  find 
probably  a  more  striking  endemic1  element,  which 

1  The  term  endemic  will  be  employed  throughout  this  work  as  applied 
to  species  peculiar  to  a  country  and  not  found  elsewhere.  Autoch- 
thonous will  be  used  in  speaking  of  a  species  which  has  originated  in  a 
country  to  which,  however,  it  is  not  peculiar ;  e.g. ,  the  Chamois  is  an 
autochthonous  Alpine  species,  but  occurs  also  in  the  Pyrenees  and 
Caucasus.  An  indigenous  species  is  one  native  to  a  country,  as 
opposed  to  the  term  "  introduced,"  and  is  applicable  to  all  species 
v\hich  have  reached  it  by  ordinary  migration. 


INTRODUCTION.  II 

with  us  is  so  meagre  that  it  can  almost  be  left 
unnoticed ;  the  main  features,  however,  remain 
nearly  the  same.  The  fauna  of  both  of  these 
islands  is  composed  of  a  strong  southern  element, 
of  an  eastern  and  a  northern  one,  and  in  addition 
we  have  here  species  whose  ancestors  lived  in 
Western  Europe. 

Before  investigating  more  minutely  the  problems 
suggested  by  the  composition  of  the  faunas  of  these 
insular  and  also  of  some  continental  areas,  it  is 
necessary  that  we  should  thoroughly  understand 
all  about  the  migrations  of  animals.  One  of  the 
principal  objects  of  this  work  is  to  show  how  the 
autochthonous  animals  of  Europe,  /.&,  those  which 
have  originated  there,  may  be  distinguished  from  the 
immigrants,  and  to  trace  the  latter  to  the  home 
of  their  ancestors.  But  in  doing  so,  it  is  necessary 
to  refer  to  the  many  important  geographical  changes 
which  have  occurred  in  Europe  during  the  latest 
geological  epochs.  The  study  of  the  geographical 
distribution  of  the  European  fauna,  as  expounded  in 
this  work,  will  in  many  instances  confirm  the  theories 
as  to  geographical  changes  based  upon  geological 
foundations.  But  in  every  case  the  views  herein 
advocated  are  founded  upon  the  geographical  distri- 
bution of  living  and  extinct  organisms  alone. 

A  terrestrial  mammal  like  the  deer  can,  under 
ordinary  circumstances,  only  reach  one  part  of  a 
country  from  another  by  walking  or  running  to  it; 
but  a  beetle,  such  as  the  cockchafer,  has  two  different 


12  HISTORY   OF   THE   EUROPEAN    FAUNA. 

modes  of  progression.  It  may  walk  or  fly.  In  both, 
however,  there  is  a  third  mode  of  transport — an  in- 
voluntary one.  The  deer  may  be  suddenly  seized  by 
a  flood  whilst  crossing  a  river,  and  carried  far  away 
without  necessarily  coming  to  grief.  The  beetle  in 
a  similar  manner  could  be  transported  to  a  distant 
country,  or  it  might  be  caught  in  a  whirlwind  and 
blown  hundreds  of  miles  off. 

We  may  thus  distinguish  between  the  natural  or 
active  and  the  accidental  or  passive  means  of  distribu- 
tion of  animals.  The  active  mode  of  dispersal  again 
may  be  only  migratory,  as  in  most  animals,  or  periodic 
and  migratory,  as  in  some  birds  and  fishes.  It  is  of 
course  the  tendency  of  every  species  to  spread  in  all 
directions  from  its  original  home,  provided  it  does  not 
encounter  obstacles,  such  as  want  of  food,  unsuita- 
bility  of  climate  or  soil,  or  barriers  such  as  mountains, 
rivers,  or  the  sea.  Birds  might  be  thought  to  be  little 
interfered  with  by  any  of  these  barriers,  but,  as  Dr. 
Wallace  has  shown,  they  are  almost  as  much  affected 
by  them  in  their  distribution  as  mammals  are. 

This  then  is  the  ordinary  migratory  distribution. 
Periodic  distribution  obtains  with  migratory  birds 
and  fishes.  The  annual  flight  of  swallows  to  their 
northern  summer  residence  comes  under  the  heading 
of  periodic  migration  or  distribution,  but  apart  from 
this,  the  swallow  must  seek  to  extend  its  range  by 
the  ordinary  method,  like  every  other  animal. 
Similarly,  the  herring  migrates  periodically  into 
shallow  water  to  spawn,  only  to  return  again  to  its 


INTRODUCTION.  13 

deeper  home,  where,  as  its  numbers  increase,  there 
must  be  a  tendency  to  spread.  We  have  in  these 
cases,  therefore,  both  a  periodic  and  an  ordinary 
movement  of  migration. 

Now,  in  studying  the  composition  of  a  fauna,  and 
especially  its  origin,  it  is  of  the  utmost  importance  to 
be  able  to  determine  approximately  the  percentage  of 
accidental  arrivals  and  of  the  ordinary  migrants — 
that  is  to  say,  of  those  which  have  reached  the 
country  owing  to  accidental  distribution,  and  of 
animals  which  have  adopted  the  usual  course  of 
migration.  It  is  of  all  the  more  import  to  review 
this  subject  of  accidental,  or,  as  Darwin  called  it,  "  the 
occasional  means"  of  distribution,  as  both  he  and 
Dr.  Wallace  have,  I  venture  to  think,  somewhat  over- 
estimated its  significance.  No  one  doubts  that  acci- 
dental transportal  takes  place,  but  the  question  is 
whether  the  accidentally  transported  animals  arrive 
living  and  reach  a  spot  where  suitable  food  is  pro- 
curable, and  whether  they  are  able  to  propagate  their 
own  species  in  the  new  locality.  For  it  must  be  clear 
to  anybody  that  the  accidental  transportal  of  a  beetle 
or  of  a  snail  to  a  new  country  cannot  affect  its  fauna 
or  add  one  permanent  member  to  it  unless  all  these 
conditions  are  fulfilled.  As  a  matter  of  fact,  only 
exceedingly  few  instances  are  on  record  of  man 
having  witnessed,  for  example,  the  accidental 
transportal  across  the  sea  to  an  island  of  a  live 
animal. 

To  mention  an  example,  Colonel  Feilden  informs 


14  HISTORY   OF  THE   EUROPEAN    FAUNA. 

us  (Zoologist,  1888)  that,  when  living  on  the  island 
of  Barbadoes,  an  alligator  arrived  one  day  on  the 
shore,  and  at  the  same  time  a  tree  measuring  40  feet 
in  length,  which  was  recognised  as  a  Demerara  species, 
was  likewise  stranded.  He  thinks  that  there  can  be 
no  doubt  that  the  alligator,  which  was  alive  when  it 
reached  Barbadoes,  was  transported  by  the  tree,  thus 
covering  a  distance  of  250  miles  from  the  nearest 
land.  Numerous  observations  on  the  accidental 
transportal  of  seeds  and  tree-trunks  from  one  island 
to  another,  and  from  a  continent  to  an  island,  have 
been  recorded,  and  even  on  our  own  shores  we  may 
witness  the  occasional  arrival  of  such  vegetable  pro- 
ducts from  a  far  distant  land.  On  the  west  coast  of 
Ireland  it  not  unfrequently  happens  that  large  West 
Indian  beans  are  stranded,  and  in  this  as  well  as  in 
many  other  similar  cases  the  seeds  have  often  proved 
none  the  worse  for  their  prolonged  immersion  in  sea- 
water.  That  locusts  are  sometimes  blown  to  great 
distances  from  the  land  is  not  so  surprising,  since 
their  power  of  steering  through  the  air  is  very  limited. 
Darwin  mentions  (p.  327)  having  caught  one  370 
miles  from  the  coast  of  Africa,  and  that  swarms  of 
them  sometimes  visited  Madeira.  Sir  Charles  Lyell 
relates  that  green  rafts  composed  of  canes  and  brush- 
wood are  occasionally  carried  down  the  Parana  River 
in  South  America  by  inundations,  bearing  on  them 
the  tiger,  cayman,  squirrels,  and  other  quadrupeds. 

But  though   actual  observations  of  such  abnormal 
instances  of  the  dispersal  of  animals  are  few,  many 


INTRODUCTION.  1 5 

experiments  have  been  made  to  demonstrate  the  pos- 
sibility of  a  passive  transportal  of  species  over  wide 
distances.  It  was  especially  Darwin  who  gave  a  great 
stimulus  by  setting  the  example  to  those  interested 
in  natural  history  in  the  conduct  of  such  researches. 
He  was  struck  by  the  fact  that,  though  land- 
shells  and  their  eggs  are  easily  killed  by  sea-water, 
almost  all  oceanic  islands,  even  the  smallest  and  most 
isolated,  are  inhabited  by  them,  and  felt  that  there 
must  be  some  unknown  but  occasionally  efficient 
means  for  their  transportal  (p.  353).  To  quote  his 
words:  "It  occurred  to  me  that  land-shells,  when 
hibernating  and  having  a  membranous  diaphragm 
over  the  mouth  of  the  shell,  might  be  floated  in  chinks 
of  drifted  timber  across  moderately  wide  arms  of  the 
sea.  And  I  find  that  several  species  in  this  state 
withstand  uninjured  an  immersion  in  sea-water  during 
seven  days:  one  shell,  the  HelLr pomatia,  after  having 
been  thus  treated  and  again  hibernating,  was  put  into 
sea-water  for  twenty  days,  and  perfectly  recovered. 
During  this  length  of  time  the  shell  might  have  been 
carried  by  a  marine  current  of  average  swiftness  to  a 
distance  of  660  geographical  miles.  As  this  Helix 
has  a  thick  calcareous  operculum,  I  removed  it,  and 
when  it  had  formed  a  new  membranous  one,  I  again 
immersed  it  for  fourteen  days  in  sea-water,  and  again 
it  recovered  and  crawled  away.  Baron  Aucapitaine 
has  since  tried  similar  experiments  :  he  placed  one 
hundred  land-shells,  belonging  to  ten  species,  in  a 
box  pierced  with  holes,  and  immersed  it  for  a 


16  HISTORY   OF    THE    EUROPEAN    FAUNA. 

fortnight  in  the  sea.  Out  of  the  hundred  shells, 
twenty-seven  recovered.  The  presence  of  an  oper- 
culum  seems  to  have  been  of  importance,  as  out  of 
twelve  specimens  of  Cyclostoma  elegans  which  it  thus 
furnished,  eleven  revived.  It  is  remarkable,  seeing 
how  well  the  Helix pomatia  resisted  with  me  the  salt- 
water, that  not  one  of  fifty-four  specimens  belonging 
to  four  other  species  of  Helix  tried  by  Aucapitaine, 
recovered.  It  is,  however,  not  at  all  probable  that 
land-shells  have  often  been  thus  transported ;  the  feet 
of  birds  offer  a  more  probable  method." 

We  have  here  positive  evidence  that  such  shells  as 
Helix pomatia  and  Cyclostoma  elegans  might  easily  be 
transported  to  an  island  from  the  mainland.  The 
former  occurs  in  France,  Holland,  and  England,  and 
the  latter  all  along  western  continental  Europe  and 
England.  And  yet  neither  of  these  species  inhabits  the 
Canary  Islands,  Madeira,  or  Ireland,  none  of  which  are 
at  too  great  a  distance  from  Europe  to  be  within  easy 
reach  for  a  floating  object.  The  fact  that  Cyclostoma 
elegans  does  not  live  in  Ireland  is  of  particular  interest 
in  connection  with  the  floating-theory  just  quoted,  as 
on  all  sides  of  Ireland  dead  specimens  have  been  picked 
up  on  the  shore,  showing  that  marine  currents  carry 
specimens  and  have  thus  transported  them  for 
countless  centuries.  Nevertheless  the  species  has  not 
established  itself  in  Ireland.  If  such  a  fate  meets  a 
land-shell  of  the  type  of  Cyclostoma  elegans,  it  may  be 
asked,  with  some  justification,  what  chance  slugs  or 
the  smaller  non-opcrculated  species  would  have  to 


INTRODUCTION.  17 

reach  an  island  like  Ireland  alive  from  the  main- 
land, and  to  colonise  it  successfully. 

Both  slugs  and  their  eggs  are  killed  by  a  short 
immersion  in  sea-water,  as  I  have  proved  experi- 
mentally. I  have  also  subjected  slugs,  in  the  act  of 
crawling  on  twigs,  to  an  artificial  spray  of  sea-water. 
This  seemed  to  irritate  their  tender  skins  to  such  an 
extent  that  they  curled  themselves  up,  released  their 
hold  on  the  twig  and  let  themselves  drop  to  the 
ground.  If  we  supposed,  therefore,  that  a  slug  had 
successfully  reached  the  sea,  transported  on  a  tree- 
trunk,  the  moisture  would  tend  to  lure  it  forth  from 
its  hiding-place  under  the  bark,  whilst  the  mere  spray 
would  prove  fatal  to  its  existence.  Those  species  of 
snails  and  slugs  which  lead  an  underground  existence, 
rarely  venturing  above  ground,  such  as  Testacella  and 
Coecilianella,  would  have  even  less  chance  of  being 
accidentally  carried  to  some  distant  island. 

The  suggestion  advanced  by  Darwin  (p.  353),  that 
young  snails  just  hatched  might  sometimes  adhere  to 
the  feet  of  birds  roosting  on  the  ground  and  thus  be 
transported,  appears  to  me  so  extremely  improbable 
as  to  be  scarcely  worth  serious  consideration.  Indeed, 
as  Darwin  himself  acknowledged  later  on,  it  does  not 
help  us  very  much  to  suggest  possible  modes  of 
transport.  What  we  require  is  direct  evidence.  How 
far  we  are,  however,  from  obtaining  it,  may  be  inferred 
from  Mr.  Kew's  remark  (p.  119),  that  "  we  have 
little  or  no  actual  evidence  of  precise  modes  of 
dispersal  even  for  short  distances  on  land." 


1 8  HISTORY   OF   THE    EUROPEAN    FAUNA. 

"  'Hv> 

A  very  curious  statement  was  made  by  a  well- 
known  French  conchologist,  the  late  M.  Bourguignat, 
with  regard  to  introductions  of  mollusca.  Whether 
he  had  any  actual  facts  collected  in  support  of  it,  I 
cannot  say,  but  he  maintained  that  species  accidentally 
transported,  with  the  exception  of  those  under  mari- 
time influence,  can  only  be  acclimatised  from  north  to 
south,  and  not  from  south  to  north — from  east  to 
west,  but  not  from  west  to  east  (p.  353). 

The  whole  theory  of  the  accidental  or  abnormal 
dispersal  of  mollusca  appears  to  have  been  originated 
by  Darwin,  in  order  to  account  for  their  presence  on  so- 
called  Oceanic  islands.  His  views  v/ere  strongly  sup- 
ported by  Wallace,  who  defines  these  islands  (p.  243) 
as  those  which  are  of  volcanic  or  coralline  formation 
usually  far  from  continents,  entirely  without  indi- 
genous land  mammals  or  amphibians,  but  with  a  fair 
number  of  birds  and  insects,  and  usually  with  some 
reptiles. 

I  do  not  wish  it  to  be  understood  that  I  am  in  any 
way  undervaluing  the  great  works  of  these  dis- 
tinguished naturalists.  Darwin's  views  have  had 
more  influence  in  advancing  Zoology  than  those  of  any 
man,  and  his  fame  is  unassailable.  Nevertheless,  I 
feel  that  his  theories  regarding  the  origin  of  the  faunas 
of  oceanic  islands  require  revision. 

The  formerly  prevalent  belief  of  the  permanence  of 
ocean  basins  has  been  shaken  by  the  utterances  of 
some  of  the  greatest  geologists  of  our  day,  whilst 
many  positively  assert  that  what  is  now  deep  sea 


INTRODUCTION.  19 

of  more  than  1000  fathoms  was  dry  land  within 
comparatively  recent  geological  epochs.  Thus  the 
Azores  are  classed  by  Darwin  and  Wallace  among 
the  oceanic  islands — that  is  to  say,  among  such 
as  have  received  their  fauna  and  flora  by  flotsam 
and  jetsam.  But  Professor  Neumayr  believes,  on 
geological  grounds,  that  the  Old  and  New  Worlds 
were  connected  by  a  land-bridge  during  Tertiary 
times  right  across  the  Atlantic,  and  that  the 
Canary  Islands,  Madeira,  and  Azores  (p.  547) 
are  the  last  remnants  of  this  continent.  This 
meets  with  the  entire  approbation  of  Dr.  von 
Ihering,  who  has  recently  re-investigated  the  sub- 
ject from  a  faunistic  point  of  view  (p.  135).  Take 
another  instance  of  one  of  Wallace's  most  typical 
oceanic  islands,  the  Galapagos  Group.  Their  fauna 
and  flora  have  recently  been  most  thoroughly 
re-explored  by  an  American  expedition,  the  result 
of  which,  according  to  Dr  Baur,  goes  to  show 
that  these  islands  must  have  formed  part  of  the 
mainland  of  South  America  at  no  distant  date.  The 
fauna  and  flora  are  therefore  to  be  regarded  as  having 
reached  them  in  the  normal  mode,  viz.,  by  migration 
on  land.  According  to  Mr.  Beddard  (p.  138),  it  is 
difficult  to  see  how  earthworms  could  be  transported 
across  the  sea.  Floating  tree-trunks  have  been  ob- 
served far  out  at  sea,  but  unless  the  water  remained 
absolutely  calm  during  the  long  period  necessary  for 
the  drifting  by  currents  so  that  no  splashing  occurred, 
the  worms  would  probably  be  killed.  Yet  earthworms 


20  HISTORY  OF  THE   EUROPEAN   FAUNA. 

do  occur  on  oceanic  islands.  It  is  indeed  quite 
possible  that  our  views  with  regard  to  the  origin  of 
the  remainder  of  the  Pacific  Islands  may  change  very 
materially,  and  once  more  revert  to  what  Dr.  Gould 
expressed  nearly  fifty  years  ago  in  the  following 
words  :  "  From  a  consideration  of  the  land-shells  on 
the  Pacific  Islands,  it  seems  possible  to  draw  some 
fair  inferences  as  to  the  relations  of  the  lands  which 
once  occupied  the  area  of  the  Pacific  Ocean,  and 
whose  mountain  peaks  evidently  now  indicate  or 
constitute  the  islands  with  which  it  is  now  studded." 
Indeed  Dr.  von  Ihering  goes  so  far  as  to  positively 
state  that  in  his  opinion  the  Polynesian  Islands  are 
not  volcanic  eruptions  of  the  sea  floor,  which  being 
without  life  were  successively  peopled  from  Australia 
and  the  neighbouring  islands,  but  the  remains  of 
a  great  Pacific  continent,  which  was  in  early 
mesozoic  times  connected  with  other  continental  land 
masses  (a,  p.  425). 

Before  coming  to  a  decision  on  the  part  played  by 
flotsam  and  jetsam  in  the  constitution  of  an  island 
fauna,  those  who  have  studied  the  problem  on  the 
spot  should,  however,  have  a  voice  in  the  matter. 
And  though,  from  my  experience  in  northern  latitudes, 
I  feel  sure  that  island  faunas  there  are  but  slightly 
affected  by  occasional  dispersal  of  species,  Mr. 
Hedley,  who  has  made  the  fauna  of  the  Pacific 
Islands  his  special  study,  assures  me  that  drift 
migration  plays  an  important  rdle  in  that  region. 
I  hope  we  may  soon  have  a  more  detailed  account 


INTRODUCTION.  21 

of  his  particular  observation  bearing  on  this  interest- 
ing subject. 

On  the  other  hand,  Mr.  Simpson,  who  has  gained 
considerable  experience  of  oceanic  dispersal  in  the 
West  Indian  region,  though  he  acknowledges  having 
often  noticed  bamboo  rafts,  which  would  be  suitable  in 
the  transportal  of  invertebrates,  nevertheless  does  not 
attach  much  importance  to  this  means  of  distribution. 
"  The  fact,"  he  remarks,  "  that  the  operculates  (oper- 
culate  land-shells)  form  so  large  a  proportion  of  the 
Antillean  land-snail  fauna,  that  a  majority  of  the 
genera  are  found  on  two  or  more  of  the  islands  and 
the  mainland,  while  nearly  every  species  is  absolutely 
restricted  to  a  single  island,  appears  to  me  to  be 
very  strong  testimony  in  favour  of  a  former  general 
land  connection"  (p.  428). 

Amphibians  are  affected  in  the  same  manner  by 
sea-water  as  slugs  are.  The  accidental  transportal  of 
an  amphibian  from  the  mainland  to  an  island  is  there- 
fore almost  inconceivable.  And  the  presence  of  frogs, 
toads,  and  newts  in  the  British  Islands,  in  Corsica 
and  Sardinia,  indicates,  if  nothing  else  did,  that  all 
these  islands  were  at  no  distant  date  united  with  the 
continent  of  Europe. 

As  regards  the  terrestrial  reptiles,  the  case  is  some- 
what different.  Many  of  them  readily  take  to  the 
sea,  and,  as  probably  all  snakes  and  some  Hzards  are 
able  to  swim,  it  is  possible  that  sometimes,  though 
very  rarely,  they  might  reach  islands  if  not  too  far 
from  a  continent.  Instances  of  accidental  transportal 


22  HISTORY  OF   THE   EUROPEAN    FAUNA. 

of  land-reptiles  to  islands  have  actually  been  observed. 
But  the  fact  of  the  occurrence  of  such  instances  by  no 
means  proves  that  reptiles  thus  conveyed  are  able  to 
establish  themselves  permanently  in  their  new  home. 
Sir  Charles  Lyell  records  in  his  Principles  of  Geology 
that  a  large  boa-constrictor  was  once  seen  floating  to 
the  island  of  St.  Vincent,  twisted  round  the  trunk  of 
a  tree.  It  appeared  so  little  injured  by  its  long 
voyage  from  South  America,  that  it  captured  some 
sheep  before  it  was  killed. 

Mammals  might  be  accidentally  conveyed  to  islands 
on  such  rafts  as  have  been  described  by  Sir  Charles 
Lyell,  and  there  are  instances  on  record  of  their 
having  crossed  short  distances  of  sea  by  swimming. 
Elephants  and  also  deer  and  pigs  are  good  swimmers, 
the  former  having  been  known  to  swim  for  six  hours 
at  a  stretch.  "But,"  remarks  Mr.  Lydekker  (p.  13), 
"it  may  be  assumed  that  about  twenty  miles  is  the 
utmost  limit  which  mammals  are  likely  to  cross  by 
swimming,  even  when  favoured  by  currents.  Such 
passages  as  these  must,  however,  be  of  very  rare 
occurrence,  for  a  terrestrial  mammal  is  not  likely 
to  take  it  into  its  head  to  swim  straight  out  to  sea 
in  an  unknown  direction.  Moreover,  supposing  a 
mammal,  near  to  a  particular  island,  to  have  arrived 
there  by  swimming,  unless  it  happen  to  be  a  pregnant 
female,  or  unless  another  individual  of  the  same 
species  but  of  the  opposite  sex  should  arrive  soon 
after  (a  most  unlikely  event),  it  would  in  due  course 
die  without  being  able  to  propagate  its  kind." 


INTRODUCTION.  23 

All  zoologists,  indeed,  are  quite  in  accord  with  Dr. 
Wallace's  view  as  expressed  in  Island  Life  (p.  74). 
"  Whenever  we  find  that  a  considerable  number  of 
the  mammals  of  two  countries  exhibit  distinct  marks 
of  relationship,  we  may  be  sure  that  an  actual  land- 
connection,  or  at  all  events  an  approach  to  within 
a  very  few  miles  of  each  other,  has  at  one  time 
existed."  As  all  the  European  islands  come  under 
this  category,  their  mammals  exhibiting  distinct 
relationship  with  those  on  the  European  continent, 
they  all  have  been  connected  with  it  formerly. 

Perhaps  the  most  powerful  of  all  agents  in  the 
transportal  of  species  by  accidental  means  is  man. 
But  his  actions  may  be  accidental  as  well  as  in- 
tentional. We  have  therefore  to  distinguish  between 
the  animals  disseminated  all  over  the  world  by  pure 
chance,  and  those  which  have  been  introduced  into 
new  countries  purposely.  Invertebrates,  such  as 
snails,  centipedes,  woodlice,  beetles,  and  cockroaches, 
are  constantly  being  unintentionally  carried  with 
vegetables,  fruit,  trees,  and  with  timber  from  one 
country  to  another.  Earthworms  are  sometimes 
transported  in  the  balls  of  earth  in  which  the  roots 
of  trees  are  enveloped.  As  regards  molluscs,  Mr. 
Kew  believes  (p.  178)  that  during  the  last  three 
centuries  at  least,  human  agency  has  influenced 
their  disposal  more  than  all  other  causes  taken 
together.  A  large  number  of  species  of  invertebrates 
in  America  are  said  to  owe  their  existence  in  that 
country  to  accidental  introduction  by  man.  In 


24  HISTORY   OF   THE   EUROPEAN    FAUNA. 

most  cases,  however,  no  particular  reason  can  be 
assigned  why  they  should  have  been  thus  introduced, 
and  as  a  matter  of  fact  there  are  always  individual 
differences  of  opinion  as  to  the  precise  number  of 
such.  Certain  it  is,  that  though  the  number  of 
supposed  introductions  from  Europe  to  America 
is  very  large,  those  which  have  been  carried  from 
America  to  Europe  is  exceedingly  small.  In  fact, 
I  remember  only  two  instances  of  accidental  animal 
importations  from  America  which  have  firmly 
established  themselves  in  Europe,  viz.,  a  small 
fresh-water  mollusc,  Planorbis  dilatatus^  and  the 
much-dreaded  vine-pest,  Phylloxera  vastatrix. 

As  a  rule  the  animals  die  out  very  shortly  after 
their  arrival  on  foreign  soil.  Many  instances, 
nevertheless,  are  on  record,  especially  in  the  case 
of  molluscs,  where  snails  thus  transported  have  not 
only  survived  but  are  apparently  in  a  flourishing 
condition  and  spreading.  Helix  aspersa,  for  example, 
our  large  garden  snail,  has  been  naturalised  in  many 
foreign  countries  by  French  and  Portuguese  sailors, 
who  had  taken  them  on  board  their  ships  as  food. 

It  certainly  cannot  be  denied  that  a  number  of 
species  among  almost  all  groups  of  invertebrates  have 
been  unintentionally  conveyed  by  man  from  Europe 
into  foreign  countries.  It  has  been  proposed  by  Dr. 
von  Ihering  to  apply  the  term  "cenocosmic"  to  those 
species  which  have  become  spread  all  over  the  world 
through  artificial  means,  and  thus  to  distinguish 
them  from  cosmopolitan  ones  which  have  attained  a 


INTRODUCTION.  25 

similar  range  naturally.  The  latter  he  calls  "  palin- 
cosmic "  species  (a,  p.  422).  Many  so-called  ceno- 
cosrnic  ants  are  believed  by  Dr.  von  Ihering  to  be 
palincosmic.  We  are  altogether  too  apt  to  regard 
cosmopolitan  as  synonymous  with  introduced,  and  we 
should  hesitate  before  concluding  that  because  one  of 
.our  common  European  species  occurs  in  Australia  or 
South  America,  it  must  have  been  transported  there 
recently  by  human  agency.  Some  of  our  widely- 
distributed  forms  are  probably  of  very  great  antiquity, 
and  may  have  spread  to  distant  lands  in  early  Ter- 
tiary times,  when  a  different  state  of  the  geographical 
conditions  enabled  them  to  do  so. 

I  cannot  quote  a  more  appropriate  instance 
than  the  molluscan  fauna  of  Madeira.  No  less 
than  thirteen  of  the  Madeiran  snails  are  looked 
upon  as  having  been  introduced  from  Europe  by 
human  agency,  on  the  sole  evidence  that  these 
happen  to  be  common  European  species.  Yet  the 
correctness  of  this  supposition  must  be  questioned 
in  face  of  the  interesting  observation  made  by 
Darwin  (p.  357),  "that  Madeira  and  the  ad- 
joining islet  of  Porto  Santo  possess  many  distinct 
but  representative  species  of  land-shells,  some  of 
which  live  in  crevices  of  stone ;  and  although  large 
quantities  of  stone  are  annually  transported  from 
Porto  Santo  to  Madeira,  yet  this  latter  island  has  not 
become  colonised  by  the  Porto  Santo  species.  Never- 
theless, both  islands  have  been  colonised  by  European 
land -shells,  which  no  doubt  had  some  advantage 


26  HISTORY   OF   THE  EUROPEAN    FAUNA, 

over  the  indigenous  species."  Darwin,  therefore, 
meets  the  evident  anomaly  by  suggesting  that  the 
European  species  are  supposed  to  possess  some 
advantages  as  colonisers.  But  the  true  explanation 
appears  to  me  to  lie  in  the  supposition  that  the 
European  land-shells  found  in  the  Madeiran  Islands 
are  all,  or  for  the  greater  part,  ancient  forms  which 
survived  both  there  and  on  the  continent,  whilst  the 
remainder  of  the  forms  inhabiting  these  islands  are 
either  such  as  are  now  extinct  in  Europe,  or  have 
become  modified  since  their  arrival  there  from  the 
continent  at  a  time  when  extensive  land-connections 
allowed  a  free  migration  by  land. 

The  theory  of  accidental  introductions  is  an  ex- 
tremely popular  one.  It  allows  free  scope  to  a  host 
of  speculations,  and  once  the  idea  has  taken  firm 
root  that  a  certain  species  is  introduced,  especially 
among  the  class  of  naturalists  who  by  way  of  ex- 
periment are  wont  to  create  new  centres  of  dispersion 
in  their  own  neighbourhood,  evidence  to  the  contrary 
must  be  of  the  most  convincing  nature  to  shake  the 
popular  belief.  Thus,  it  is  almost  regarded  as  an 
established  fact  by  conchologists  and  others,  that  the 
fresh-water  mussel  (Dreyssensiapolymorpka)  was  intro- 
duced into  England  at  the  beginning  of  this  century. 
Though  it  has  been  proved  that  this  species  is  quite 
unable  to  live  in  pure  sea-water,  yet  the  view  that  it 
has  been  carried  from  the  Black  Sea  ports  to  this 
country  attached  to  the  bottom  of  ships  is  maintained 
by  many,  whilst  others  incline  to  the  theory  that  the 


INTRODUCTION.  2/ 

shell  came  with  timber.  But  Dreyssensia  polymorpha 
was  by  no  means  always  confined  to  the  Caspian  and 
Black  Sea  areas;  it  occurs  abundantly  in  the  lower 
continental  boulder-clay  (see  p.  230),  and  no  doubt  it 
had  at  one  time  a  much  wider  geographical  distribution. 
Itp  appears  to  me  possible,  that  it  was  able  to  maintain 
itself  in  certain  fresh-water  lakes  and  slow-flowing 
rivers  in  Northern  Europe,  from  which  it  might  have 
spread  since  the  introduction  of  canals  into  Europe  at 
the  beginning  of  the  century.  As  the  larva  of  this 
fresh-water  mussel  is  free-swimming,  its  propagation 
is  much  favoured  by  canals.  Quickly-flowing  rivers 
are  fatal  to  its  existence,  since  the  delicate  larvae  are 
swept  out  to  sea  and  perish.  Such  an  hypothesis  as 
this  is  strengthened  by  the  fact  of  its  recent  discovery 
in  a  sandy  layer  fifteen  feet  below  the  present  surface 
under  the  streets  of  London  in  a  deposit  which  prob- 
ably, as  Mr.  Woodward  remarks  (p.  8),  was  accumu- 
lated in  the  early  days  of  the  city's  existence.  In 
spite  of  Mr.  Woodward's  interesting  find,  and  Dr. 
Jeffreys'  opinion,  who  always  maintained  that  this 
shell  was  indigenous  to  England,  popular  belief  still 
clings  tenaciously  to  the  introduction  theory. 

Among  man's  intentional  introductions  into  a  new 
country,  no  instance  is  better  known  than  that  of  the 
rabbit  to  Australia.  Rabbits  are  entirely  confined  to 
Europe.  In  their  transplantation  to  Australia  they 
were  carried  to  a  country  with  a  different  climate  and 
among  new  surroundings.  Yet  the  rabbits  flourished, 
and  within  comparatively  few  years  increased  to  such 


28  HISTORY  OF   THE   EUROPEAN    FAUNA. 

an  extent  as  to  become  a  burden  and  pest  to  the 
country.  It  may  be  remembered  though,  that,  owing 
to  the  complete  absence  of  small  carnivores,  which 
act  with  us  as  a  check  upon  the  too  rapid  increase  of 
this  rodent,  the  speed  with  which  it  established  itself 
in  the  new  surroundings  is  not  so  very  surprising. 

Many  of  the  English  settlers  in  the  New  World  felt 
that  America  lacked  the  presence  of  our  familiar 
birds.  The  homely  sparrow  was  therefore  brought 
over,  with  the  result  that  the  Agricultural  Department 
of  the  United  States  is  now  devising  means  for  its 
destruction,  so  rapid  has  been  its  increase. 

Similarly,  the  inhabitants  of  Jamaica,  annoyed  by 
the  great  profusion  of  rats  in  their  island,  sent  over  to 
India  for  a  number  of  mongoose.  These  have  deci- 
mated the  rats  since  their  arrival,  but  they  have 
multiplied  to  such  an  extent  as  to  be  a  serious 
menace  to  the  native  fauna. 

To  give  an  instance  nearer  home,  the  Capercaillie 
(Tetrao  urogallus)  was  successfully  introduced  into 
Scotland  in  1837.  From  its  different  centres  of 
distribution  it  is  spreading  in  all  directions  where 
sufficient  cover  is  obtainable.  But  this  case  differs 
from  the  others  very  materially,  in  so  far  as  this  bird 
was  formerly  a  native  of  Scotland,  and  only  became 
extinct  during  the  last  century. 

However,  although  there  are  many  examples  of  un- 
doubtedly successful  introductions  by  human  agency, 
quite  as  many,  or  perhaps  more,  unsuccessful  ones 
might  be  quoted.  In  fact,  it  is  by  no  means  easy  to 


INTRODUCTION.  29 

establish  a  species  in  any  new  locality.  Frequently 
it  happens  that  the  species  seems  to  be  on  the  increase 
at  first,  but  then  there  is  a  decline,  and  after  a  few 
years  the  new  plantation  has  entirely  vanished.  In 
other  cases,  the  species  disappears  immediately  after 
the  introduction  takes  place,  or  lingers  on  for  many 
years  if  it  receives  special  and  uninterrupted  pro- 
tection. 

It  may  not  be  generally  known  that  the  English 
Hare  {Lepus  Europeans)  is  not  found  in  Ireland,  where 
the  Mountain  Hare  (Lepus  variabilis)  alone  occurs. 
Attempts  to  acclimatise  the  English  species  have 
been  made  in  a  number  of  places  in  Ireland,  but 
many  of  them  have  been  failures,  and  not  one  of 
them  has  been  a  signal  success.1  Similarly,  the 
endeavour  to  introduce  the  French  or  Red-legged 
Partridge  (Caccabis  rufd)  into  Ireland  has  met  with  a 
like  result.  According  to  Dr.  Day,  it  was  tried  during 
the  summer  of  1869  to  naturalise  the  Sterlet  (Acipenser 
ruthenus)  from  Russian  waters  into  the  Duke  of 
Sutherland's  River  Fleet  by  importing  artificially  im- 
pregnated ova.  From  one  hundred  and  fifty  to  two 
hundred  lively  young  sterlets  are  said  to  have  been 
turned  out,  but  nevertheless  the  experiment  met  with 
no  success.  Several  fortunately  abortive  efforts  were 
also  made  in  British  rivers  to  establish  Silurus  glanis, 
a  hideous  monster  of  a  fish,  and  quite  unpalatable. 

1  I  might  refer  any  one  more  specially  interested  in  these  intro- 
ductions to  an  article  on  this  subject  in  the  Irish  Naturalist  of  March 
1898,  by  Mr.  Barrett- Hamilton. 


30  HISTORY   OF   THE    EUROPEAN    FAUNA. 

The  Natterjack  Toad  (Bufo  calamitd}  has  a  very 
local  distribution  in  the  British  Islands.  In  Ireland 
it  is  found  only  along  the  coast  of  Dingle  Bay  in 
County  Kerry,  where  it  is  known  among  the  peasantry 
as  the  Black  Frog.  There  is  no  doubt  about  its 
being  indigenous  there,  and  though  it  has  not  spread 
beyond  the  very  limited  area  of  its  habitat,  the  Irish 
climate  cannot  be  said  to  be  unsuited  to  its  existence. 
Yet  it  seems  to  be  extremely  difficult  to  acclimatise 
it  elsewhere,  for  though  no  less  than  sixty  specimens 
were  turned  out  in  Phcenix  Park,  Dublin,  about 
forty  years  ago,  none  of  them  were  ever  seen 
afterwards.  They  were  placed  in  the  vicinity  of  one 
of  the  lakes,  so  as  to  give  them  ample  scope  for 
breeding  and  developing  the  young,  and  in  sur- 
roundings which  were  considered  eminently  suitable 
at  the  time. 

It  has  occasionally  happened,  too,  that  animals  are 
introduced  by  kindly-disposed  persons  with  the  view 
of  adding  a  species  to  their  fauna,  in  complete 
ignorance  of  their  previous  existence  in  the  country 
where  they  wished  to  naturalise  them.  Thus  we  are 
told  that  in  the  year  1699  one  of  the  Fellows  of 
Trinity  College,  Dublin,  procured  Frog's  spawn  from 
England  in  order  to  add  that  amphibian  to  the  Irish 
fauna.  It  was  placed  in  a  ditch  in  the  College  Park, 
whence  the  species  is  supposed  to  have  gradually 
spread  all  over  the  island.  This  story  is  quoted  by 
many  writers  as  the  true  history  of  the  Frog  in 
Ireland,  and  is  given  as  an  example  of  the  rapidity 


INTRODUCTION. 

with  which  animals  spread.  Unfortunately  the  would- 
be  introducer  seemed  unaware  that,  according  to 
Stuart's  History  of  Armagh,  the  first  Frog  which  was 
ever  seen  in  Ireland  made  its  appearance  in  a  pasture 
field  near  Waterford  about  the  year  1630,  that  is  to 
say,  seventy  years  before  its  introduction  in  Dublin.1 
But  even  Stuart  was  mistaken  in  supposing  that  no 
Frog  had  ever  been  seen  in  Ireland  before,  since 
Giraldus  Cambrensis,  in  his  Topography  of  Ireland, 
mentions  that  a  Frog  was  found  in  a  meadow  near 
Waterford  in  the  year  1187. 

Certain  British  species  of  vertebrates  are  generally 
looked  upon  as  introduced  species,  though  we  cannot 
trace  any  record  of  their  first  establishment,  and  it  is 
quite  possible  that,  though  there  was  local  extinction 
and  subsequent  local  re-introduction,  they  are  truly 
indigenous  and  may  never  have  become  totally 
extinct.  Such  are,  for  instance,  the  Rabbit  (Leptts 
cuniculus)  and  the  Pheasant  (fhasianus  colchicus}. 
The  latter  certainly  had  become  naturalised  in 
England  before  the  Norman  invasion. 

But  cases  of  introduction  such  as  those  above  re- 
ferred to  are  by  no  means  confined  to  the  vertebrates, 
similar  instances  among  invertebrates  being  numer- 
ous enough.  I  am  sure  every  naturalist  is  personally 
acquainted  with  a  good  number,  and  it  is  hardly 
necessary  that  I  should  quote  in  any  detail  after 

1  I  should  recommend  those  who  are  particularly  interested  in  the 
full  history  of  the  Irish  frog  to  read  the  notes  on  this  subject  contained 
in  vol.  ii.  of  the  Irish  Naturalist. 


32  HISTORY  OF   THE    EUROPEAN   FAUNA. 

what  has  been  said  on  the  subject  generally.  The  two 
species  of  snails,  Helix pomatia  and  Cyclostoma  elegans^ 
both  of  which  occur  in  England,  and  which  I  had  occa- 
sion to  mention  among  those  experimented  on  by 
Darwin,  were  turned  out  in  several  suitable  localities  in 
Ireland  by  Thompson,  but  failed  to  establish  them- 
selves. The  former,  according  to  Mr.  Kew,  was  also 
introduced  into  Scotland  and  Norway,  whilst  fifty  or 
sixty  specimens  were  brought  to  Petersfield  in  Eng- 
land, but  none  of  these  trials  at  acclimatisation 
were  successful.  As  among  vertebrates,  a  large  num- 
ber of  the  so-called  successful  introductions  rest  upon 
insufficient  evidence. 

When  we  once  more  carefully  review  the  evidence 
as  to  the  undoubted  difficulty  attendant  on  intentional 
introduction  of  animals  by  human  agency,  placed  as 
they  often  were  in  most  suitable  localities,  we  must 
feel  that  accidental  introduction  cannot  play  an  im- 
portant r61e  in  the  making  of  the  fauna  of  any  country. 
Especially  is  this  the  case  with  an  island  fauna. 
Vertebrates  are  almost  altogether  excluded,  and  in- 
vertebrates must  arrive  singly  as  a  rule,  often  stranded 
on  an  inhospitable  and  unsuitable  shore.  Their 
chances  of  surviving  a  passage  by  sea,  of  finding 
suitable  food  and  shelter  and  a  mate  in  order  to  pro- 
create their  species,  appear  to  me  infinitesimally  small. 
Yet  there  may  be  some  such  cases.  However,  I  quite 
agree  with  Mr.  Andrew  Murray — a  high  authority 
on  geographical  distribution — that  "colonisation  or 
'occasional  dispersal  is  insufficient  to  account  for  the 


INTRODUCTION.  33 

character  of  the  faunas  and  floras  of  oceanic  islands; 
and  I  believe  that  the  normal  mode  in  which  islands 
have  been  peopled,  has  been  by  direct  continuity  with 
the  land  at  some  former  period,  or  by  contiguity  so 
close  as  to  be  equivalent  to  junction  "  (p.  15).  "  That 
a  slight  intermixture,"  he  continues,  "due  to  Mr. 
Darwin's  colonisation,  occurs  in  many  (probably  in 
all)  I  am  ready  to  admit ;  and  from  instances  to  be 
afterwards  noticed,  I  am  disposed  to  reckon  the  pro- 
portions of  such  intermixtures  in  the  flora,  in  the 
most  favourable  circumstances,  at  not  more  than  two 
per  cent.  In  the  fauna  I  think  it  must  be  much  less." 

Mr.  Murray's  views,  though  they  relate  only  to 
oceanic  islands,  are  likewise  applicable  to  continental 
islands  such  as  our  own.  I  think  we  might  take  the 
admixture  in  the  British  fauna  due  to  occasional, 
including  human  introduction,  as  amounting  to  five 
per  cent.  It  is  better  to  take  a  high  estimate,  so  as  to 
include  all  the  species  about  whose  native  land  there 
might  be  some  reasonable  doubt.  Now  of  what 
importance,  after  all,  is  this  five  per  cent?  The 
remaining  ninety-five  per  cent,  of  the  species  of 
animals  belonging  to  the  British  fauna  undoubtedly 
migrated  to  these  islands  in  the  normal  way  by  land. 

It  is  of  great  importance,  in  dealing  with  the 
question  of  the  origin  of  the  British  fauna,  to 
thoroughly  grasp  this  conclusion — that  ninety-five 
per  cent,  of  the  animals  have  reached  us  by  land. 
We  can  afford  in  fact  to  ignore  the  five  per  cent, 
altogether.  It  is  an  insignificant  factor.  As  regards 

3 


34  HISTORY   OF   THE    EUROPEAN    FAUNA. 

the  botanical  aspect  of  the  question,  botanists  are 
quite  in  accord  with  the  zoologists,  and  entirely 
share  their  views  in  the  belief  of  a  former  land- 
continuity  between  the  British  Islands  and  the 
Continent.  "  It  cannot  be  denied,"  says  Professor 
Blytt  (p.  32),  "that  a  plant  of  one  or  another 
species  may,  in  an  exceptional  case,  migrate,  without 
human  assistance,  all  at  once,  across  large  tracts 
of  land  and  sea,  and  that  such  migration,  if  operating 
during  geological  periods,  might  introduce  a  number 
of  species  even  into  distant  oceanic  islands ;  but 
when  the  question  is  of  whole  communities  of  plants, 
such  as  the  above  enumerated  elements  in  our  flora, 
then  such  an  accidental  and  sudden  transport  across 
large  tracts  can  only  be  conceived  to  be  at  all 
probable  in  the  case  of  Arctic  plants  carried  by 
drifting  ice  to  a  bare  country  without  native  flora  ; 
as  to  the  other  species,  we  must  imagine  that  the 
migration  during  the  gradual  change  of  climate  has 
proceeded  sloivly  and  step  by  step  across  connected 
tracts  of  cotmtty.  In  that  manner  we  may  assume 
that  our  country  has  in  the  course  of  time  obtained 
its  present  covering  of  plants.  Each  of  the  above- 
named  elements  in  our  flora  has  doubtless  its 
corresponding  element  in  our  fauna.  The  fauna 
and  flora  of  a  region  stand  in  relation  of  com- 
plicated dependence  to  each  other.  The  animals 
live  on  the  plants.  The  fecundation  of  the  plants 
takes  place  in  a  great  degree  by  means  of  insects ; 
their  seeds  are  often  scattered  by  resident  birds  and 


INTRODUCTION.  35 

quadrupeds.  Everything  indicates  that  conveyance 
1o  small  distances  is  the  rule,  and  that  sudden  and 
long  migration  is  the  exception." 

The  conviction  which  has  been  gained  by  zoologists 
and  botanists,  that  the  British  Islands  once  formed 
part  of  the  Continent,  is  based  on  the  present  British 
fauna  and  flora.  The  remains,  however,  of  animals 
which  used  formerly  to  live  in  these  countries,  such  as 
the  Mammoth,  the  Irish  Elk,  the  Cave  Bear,  and  many 
others,  tell  us  the  same  tale.  They  could  not  have 
peopled  England  by  swimming  across  the  Channel, 
or  even  by  walking  across  solid  ice,  as  has  once  been 
suggested.  Nothing  but  a  land-connection  induced 
them  to  explore  this  country  more  closely,  and 
finally  to  decide  on  settling  there. 

The  origin  of  the  British  fauna  will  be  discussed 
more  in  detail  in  the  third  chapter.  The  methods 
of  investigation  adopted,  along  with  a  general  scheme 
of  this  book,  will  be  found  in  the  next. 

The  manner  in  which  the  origin  of  the  fauna  of  any 
particular  continental  area  can  be  traced  is  very 
similar  to  that  adopted  in  the  case  of  an  island. 
Portions  of  the  continent  of  Europe  can  be  shown 
to  have  been  islands  in  former  times.  Thus  the 
Crimea,  now  a  peninsula  united  to  the  mainland 
by  the  narrow  isthmus  of  Perekop,  must  have  been 
an  island  in  comparatively  recent  times.  The  absence 
of  a  number  of  striking  and  familiar  South  Russian 
species  of  mammals  and  reptiles  proves  this  to  have 
been  the  case.  It  was  probably  long  after  the  appear- 


36  HISTORY   OF   THE    EUROPEAN    FAUNA. 

ance  of  man,  though  before  historic  times,  that  these 
changes  took  place. 

We  shall  learn  in  the  subsequent  chapters,  that 
by  a  careful  study  of  the  fauna  and  flora  the 
fact  can  be  established,  not  only  of  the  former 
connection  of  an  island  with  a  continent,  but  also 
whether  such  union  existed  (geologically  speaking) 
within  recent  or  more  remote  times.  The  better 
the  fauna  is  known,  both  recent  and  fossil,  the  more 
precisely  can  the  period  of  connection  be  indicated, 
and  its  duration  determined. 


CHAPTER  II. 

PRELIMINARY  CONSIDERATIONS. 

I  INTEND  to  give  in  this  chapter  a  general  outline 
of  the  subject  which  will  be  discussed  in  the 
subsequent  ones.  This  will  include  a  brief  history 
of  the  great  events,  in  recent  geological  times,  which 
have  modified  the  evolution  of  the  European  fauna 
by  the  influence  which  they  have  exerted  on  the 
course  of  the  successive  streams  of  migration. 

The  composition  of  the  European  fauna  is  the 
first  item  which  will  have  to  be  taken  into  con- 
sideration. But  not  only  must  the  existing  species 
of  animals  be  dealt  with  :  the  extinct  ones,  too, 
at  least  those  which  have  lived  in  Europe  during 
late  Tertiary  times,  will  be  useful  for  our  inquiries. 
A  knowledge  of  the  past  faunas  is  a  most  important 
factor  in  tracing  the  original  home  of  the  European 
animals. 

Where  a  species  first  originated,  whether  this  was 
in  one  or  several  places,  or,  in  other  words,  where 
it  first  had  its  home,  cannot  be  determined  with 
absolute  certainty  in  the  present  state  of  our 
knowledge,  but  as  a  rule  it  can  be  indicated 
approximately  with  a  fair  amount  of  precision.  In 

37 


38  HISTORY   OF   THE    EUROPEAN    FAUNA. 

a  few  instances,  species  may  possibly  have  had  a 
dual  origin.  The  majority  of  naturalists  doubt 
that  there  are  any  such,  but  it  seems  to  me 
that  almost  the  same  forces  may  have  acted  in 
different  localities  on  certain  forms  so  as  to  produce, 
in  very  exceptional  circumstances,  similar  species. 
The  vast  majority  of  animals,  however,  have  no 
doubt  originated  in  one  locality;  or,  we  might  say, 
almost  all  species  have  but  one  home. 

We  may  assume  that  every  animal  gradually 
extends  its  range  by  migration,  as  the  result  of  the 
natural  increase  of  the  species  necessitating  a  search 
for  fresh  feeding  grounds.  Every  species  thus  tends 
to  slowly  take  possession  of  all  the  habitable  parts 
of  the  globe  to  which  it  has  access.  They  would 
all  naturally  spread  from  their  original  homes  in 
every  direction,  unless  prevented  by  an  impassable 
barrier.  We  have  already  learned  that  to  all  land 
animals,  the  sea  acts  as  such  a  barrier.  Mountains 
and  rivers  act  also  in  a  similar  way,  but  not  to  the 
same  extent.  It  is  not  difficult  to  understand  also 
that  a  forest  may  be  a  formidable  barrier  to  a  typical 
inhabitant  of  the  open  country  and  vice  versa,  whilst 
a  desert  is  impassable  to  almost  all  terrestrial  organ- 
isms. Some  species  are  scarcely  affected  by  climate, 
and  flourish  equally  well  in  the  tropics  and  in 
temperate  or  cold  countries  ;  the  majority,  however, 
are  greatly  influenced  by  it.  "  No  more  striking 
illustration,"  remarks  Merriam  (p.  38),  "could  be 
desired  of  the  potency  of  climate  compared  with 


1'RELiMiNARY   CONSIDERATIONS.  30 

the  inefficiency  of  physical  barriers,  than  is  presented 
by  the  almost  total  dissimilarity  of  the  North 
American  Tropical  and  Sonoran  Regions,  though 
in  direct  contact,  contrasted  with  the  great  similarity 
of  the  Boreal  Regions  of  North  America  and  Eurasia, 
now  separated  by  broad  oceans,  though  formerly 
united,  doubtless,  in  the  region  of  Behring  Sea." 

To  return  to  the  composition  of  the  European 
fauna,  we  now  know  positively  that  a  number  of  the 
mammals  and  birds  inhabiting  Central  and  Eastern 
Europe  are  of  Siberian  origin.  How  they  came,  and 
when,  will  form  the  subject  for  discussion  in  Chapter 
V.  At  present  it  will  suffice  to  mention  that  in  the 
superficial  deposits  belonging  to  the  Pleistocene 
series  of  the  North  European  plain  have  been  dis- 
covered the  remains  of  many  typical  members  of  the 
Siberian  Steppe-fauna.  Some  of  these,  such  as  the 
Saiga- Antelope  (Saiga  tartarica),  Fig.  2,  still  inhabit 
portions  of  Eastern  Europe,  whilst  others  have  re- 
treated to  their  native  land.  But  it  might  be  asked, 
how  is  it  known  that  these  species  did  not  originate 
in  Europe,  and  thence  migrate  to  Siberia?  Because 
if  they  had  originated  on  our  continent,  they  would 
have  spread  there.  They  would  have  invaded 
Northern  and  Southern  Europe,  and  they  would 
probably  have  left  some  remains  in  Spain,  Italy,  or 
Greece.  They  would  also  have  left  some  of  their 
relations  in  Europe ;  but  all  their  nearest  allies, 
too,  are  Asiatic.  Moreover, — and  this  completes,  I 
think,  the  proof  of  their  Siberian  origin, — the  Pleisto- 


46 


HISTORY   OF   THE   EUROPEAN    FAUNA. 


PRELIMINARY   CONSIDERATIONS.  4! 

cene  remains  of  these  animals  in  Europe  become 
less  abundant,  and  the  number  of  species  likewise 
decreases,  as  we  proceed  from  east  to  west.  With 
these  remains  of  Steppe  animals  are  generally  asso- 
ciated those  of  others,  which  we  must  also  look  upon 
as  Siberian  emigrants,  such  as  the  Pikas  or  tailless 
Hares  belonging  to  the  genus  Lagomys,  the  pouched 
Marmots  (Spennophilus),  and  others.  Some  of  them, 
as  I  have  mentioned,  still  inhabit  Central  and  Eastern 
Europe,  whilst  others  have  a  wider  distribution  on  our 
continent. 

This  migration  must  have  been  an  unusually  large 
one.  It  has  been  suggested  that  the  Glacial  period 
had  some  connection  with  it,  and  there  can  be  little 
doubt,  as  we  shall  see  later  on,  that  a  change  of 
climate  probably  brought  about  this  great  Siberian 
invasion  of  Europe.  But  other  causes  might  tend 
in  the  same  direction,  such  as  want  of  sufficient 
food  after  a  few  years  of  great  increase  of  any  parti- 
cular species.  It  is  not  known  to  what  we  owe 
the  periodic  visits  of  the  Central  Asiatic  Sandgrouse 
(Syrrhaptes  paradoxus\  Fig.  3,  but  certain  it  is  that 
immense  flocks  of  these  birds  invade  Europe  from 
time  to  time  at  the  present  day,  just  as  those 
mammals  may  have  done  in  past  ages. 

The  Siberian  migrations  will  be  spoken  of  in  the 
subsequent  pages,  as  the  Siberian  element  of  the 
European  fauna.  These  migrations,  however,  are  not 
the  only  ones  which  reached  Europe  from  Asia. 
The  sixth  chapter  deals  with  migrations  which  have 


42  HISTORY   OF   THE    EUROPEAN    FAUNA 

influenced  our  fauna  far  more  than  the  Siberian.  The 
latter  did  not  last  long,  nor  did  they  affect  the  whole 
of  Europe.  But  what  I  may  call  the  Oriental  migra- 
tions spread  to  every  corner  of  Europe  and  certainly 
lasted  throughout  the  whole  of  the  Tertiary  Era.  The 
Oriental  element  came  probably  from  Central  and 


FIG.  3.  — Central  Asiatic  Sandgrouse  (Syrrhaftes  faradoxus}. 

Southern  Asia,  and  in  its  march  to  Northern  Europe 
it  was  joined  by  local  European  migrations.  For  on 
our  continent,  too,  animals  originated  and  spread  in 
all  directions  from  their  centres  of  dispersal.  A 
separate  chapter  has  been  given  to  the  Alpine  fauna, 
and  another  to  that  of  South-western  Europe,  which 
will  be  known  by  the  name  of  the  Litsitanian  element. 


PRELIMINARY   CONSIDERATIONS.  43 

Finally,  animals  have  also  reached  us  from  the  north, 
and  in  the  fourth  chapter  the  history  of  that  remark- 
able migration  will  be  fully  discussed  under  the  title 
of  the  Arctic  element  of  the  European  fauna. 

It  is  generally  believed  that  Africa  played  an  im- 
portant role  in  the  peopling  of  our  continent,  but 
this  is  quite  a  mistake.  The  eminent  Swiss  palae- 
ontologist Riitimeyer  was  quite  right  in  saying  (p. 
42)  that  it  is  much  more  probable  that  Morocco, 
Algeria,  and  Tunis  were  stocked  with  animals  by  way 
of  Gibraltar,  and  perhaps  also  by  Sicily  and  Malta, 
from  Europe,  than  the  South  of  Europe  from  Africa. 

I  have  already  referred  to  what  are  known  as 
"centres  of  dispersion"  of  animals,  but  before  con- 
tinuing to  explain  the  general  outline  of  this  book, 
it  will  be  necessary  to  make  a  few  additional  remarks 
on  the  subject. 

Since  every  animal  naturally  tends  to  spread  in 
every  direction  from  its  original  home — that  is  to 
say,  from  the  place  of  its  origin — the  latter  should 
correspond  with  the  centre  of  its  range.  And  in 
any  particular  group  of  animals  the  maximum 
number  of  species  should  be  formed  in  the  area  or 
zone  which  is  the  centre  of  its  distribution.  In  the 
great  majority  of  instances  this  is  probably  the  case, 
in  the  higher  animals  perhaps  less  so  than  in  the  lower; 
still  the  rule  must  hold  good  that  the  original  home 
of  a  species  is  generally  indicated  by  the  centre  of 
its  geographical  distribution. 

Take  for  example  our  familiar  Badger  (Meles  taxus). 


44  HISTORY   OF   THE   EUROPEAN   FAUNA. 

It  inhabits  Europe  and  Northern  Asia.  It  is  absent 
apparently  from  many  parts  of  Central  Asia,  but  it 
appears  again  farther  south  in  Palestine,  Syria,  Persia, 
Turkestan,  and  Tibet.  West  Central  Asia  would  be 
about  the  centre  of  its  range.  That  this  corresponds 
to  its  place  of  origin  is  indicated  by  the  fact  that  the 
only  three  other  Badgers  known — viz.,  M.  anakuma, 
M.  leucuruS)  and  M.  albogularis — are  confined  to  Asia. 
If  we  examine  the  fossil  history  of  the  genus,  we  find 
that  the  two  most  ancient  instances  of  the  existence 
of  Badgers  have  been  discovered  in  Persia,  where  M. 
Polaki  and  M.  maraghanus  occur  in  miocene  deposits. 
The  latter  had  migrated  as  far  west  as  Greece  in  mio- 
cene times  ;  no  other  trace  of  the  Badger,  however,  is 
known  from  Europe  until  we  come  to  the  pleistocene 
beds.  There  are  a  good  many  cases  known  among 
mammals  where  the  centre  of  dispersion  would  indicate 
to  us  a  similar  origin.  On  the  other  hand,  there  may 
be  no  fossil  evidence  of  the  occurrence  of  a  species, 
or  of  its  ancestors,  in  Asia,  whilst  such  has  been 
discovered  in  Europe.  I  think,  however,  that  the 
present  range  of  a  species  forms  a  safer  criterion  for 
the  determination  of  its  original  home,  as  the  Asiatic 
continent  is  still  practically  unworked  from  a  palaeon- 
tological  point  of  view.  In  a  letter  which  I  received 
from  Professor  Charles  Deperet,  he  advocates  the 
view  that  the  wild  Boar  (Sus  scrofa)  is  probably  of 
European,  and  not,  as  I  maintained  (c,  p.  455),  of 
Asiatic  origin  ;  because  there  seemed  to  be  a  direct 
descent  from  Hyotherium  of  the  middle  miccene  of 


PRELIMINARY  CONSIDERATIONS.  45 

Europe,  through  the  upper  miocene  Pig  of  the  Mount 
Leberon  (Sus  major)  and  of  Eppelsheim  (Sus 
antiquus\  and  the  pliocene  Pigs  of  Montpellier  (Sus 
provincialis)  and  of  the  Auvergne  (Sus  arvernensis}. 
No  doubt  this  appears  rather  a  strong  case  in  favour 
of  the  European  origin  of  the  wild  Boar,  but  although 
the  Tertiary  strata  of  Asia,  as  I  remarked,  are  as  yet 
little  known,  a  number  of  fossil  pigs  are  known  from 
India,  Persia,  and  China,  the  oldest  being  the  upper 
miocene  Persian  Pig  (Sus  maraghanus).  Pigs  are 
therefore  as  old  in  Asia  as  in  Europe,  and  as  a  direct 
intercourse  between  the  two  continents  probably  never 
ceased  since  miocene  times,  it  is  not  surprising  that 
this  genus  should  occur  in  both.  Even  if  the  genus  had 
its  origin  in  Europe,  it  is  quite  possible  that  in  later 
Tertiary  times,  the  active  centre  of  origin  was  shifted 
to  the  neighbouring  continent,  and  that  henceforth 
many  new  species  issued  forth  from  Asia,  some  of 
which  may  subsequently  have  been  modified  on 
reaching  our  continent.  The  wild  Boar  (Sus  scrofa\ 
however,  to  judge  from  its  general  range,  I  must  look 
upon  as  merely  an  immigrant  in  Europe.  I  have  no 
doubt  that  it  originated  somewhere  in  Asia,  probably 
in  the  south. 

The  view  I  take  of  the  origin  of  our  European 
Boar  is  also  supported  by  Dr.  Forsyth  Major's  recent 
researches.  He  was  led  to  a  re-investigation  of  the 
history  of  the  Pig  while  examining  a  la'-ge  number 
of  fossil  skulls  in  the  Museum  at  Florence,  and  came 
to  the  conclusion  that  only  three  or  four  species  of 


46  HISTORY   OF    THE    EUROPEAN    FAUNA. 

recent  wild  pigs  can  be  clearly  distinguished  (b,  p.  298). 
One  of  these,  viz.,  Sus  vittatus^  he  thinks,  is  trace- 
able in  slight  modifications  frcm  Sardinia  to  New 
Guinea  and  from  Japan  to  South  Africa.  The  centre 
of  distribution  of  this  species  lies  in  Southern  Asia. 
Of  the  three  remaining  species,  two,  viz.,  Sus  ver- 
rucosus  and  *S.  barbarus,  are  entirely  confined  to  the 
great  islands  which  form  part  of  the  Malay  Archi- 
pelago. Finally,  Sus  scrofa,  our  Central  European 
wild  Boar,  is  so  closely  related  to  5.  vittatus  that  the 
Sardinian  Boar  might  be  looked  upon  as  a  variety  of 
either  the  one  or  the  other.  At  any  rate,  Dr.  Major 
recognises  clearly  in  Sus  vittatiis  the  representative 
of  the  ancestral  stock  of  which  Sus  scrofa  is  a  some- 
what modified  offshoot. 

The  fauna  of  Europe  consists,  as  I  have  mentioned, 
to  a  large  extent  of  immigrants  from  the  neighbouring 
continents.  This  is  especially  noticeable  among  the 
higher  animals.  When  we  come  to  the  lower,  such  as 
the  amphibia,  we  find  a  larger  percentage,  and  among 
the  land  mollusca  the  great  majority,  to  be  of  Euro- 
pean origin.  The  foreigners  are,  as  we  learned,  called 
Orientals,  Siberians,  and  Arctics.  For  the  sake  of 
convenience,  only  two  of  the  great  European  centres 
of  origin  have  a  chapter  devoted  to  themselves, 
namely,  the  Alpine  and  the  Lusitanian  centres. 
There  is  another,  -  however,  of  almost  equal  im- 
portance which  lies  in  the  east. 

In  the  British  Islands  there  is  only  an  exceedingly 
small  and  insignificant  group  of  species  which  are 


PRELIMINARY   CONSIDERATIONS.  47 

peculiar,  and  which  we  may  consider  to  have  had 
their  origin  there.  Almost  the  whole  of  the  British 
fauna  is  composed  of  streams  of  migrants  which  came 
from  the  north,  south,  and  east,  though  many  of 
these  immigrant  species  have  since  their  arrival  been 
more  or  less  distinctly  modified  into  varieties  or  local 
races. 

The  eminent  French  conchologist  Bourguignat  (a,  p. 
352)  was  of  opinion  that,  as  far  as  terrestrial  mollusca 
were  concerned,  there  are  in  Europe  three  principal 
centres  of  creation  or  dispersion — all  situated  in 
mountainous  countries  and  not  in  the  plains.  He 
distinguished  the  Spanish,  Alpine,  and  Tauric  centres, 
and  believed  that  almost  all  species  known  from 
Europe  had  originated  in  one  of  these  three,  and  that 
each  of  them  possessed  quite  a  distinct  type  of  its 
own.  This  theory  seems  to  agree  very  well  with  the 
facts  of  distribution.  Let  us  take,  for  instance,  the 
genus  Clausilid)  a  pretty  turret-shaped  snail,  which 
abounds  on  old  ruined  walls.  Only  two  species,  viz., 
CL  laminata  and  CL  bidentata,  are  met  with  in  Ireland. 
In  England  we  find  the  same  species  with  the  addition 
of  two  others,  CL  biplicata  and  Cl.  Rolphii.  Crossing 
over  the  Channel  to  Belgium,  these  four  species  occur 
again,  and  also  several  others  not  known  in  England. 
In  Germany  the  list  of  Clausilice  mounts  up  to  twenty- 
five  species,  including  all  those  found  in  the  British 
Islands.  As  we  proceed  eastward  the  number  of 
species  of  this  genus  increases  steadily,  and  when  we 
reach  the  Caucasus  or  the  Balkan  Peninsula  the  con- 


48  HISTORY    OF   THE   EUROPEAN    FAUNA. 

chologist  is  able  to  make  a  collection  of  several 
hundred  different  kinds,  whilst  farther  east  again  they 
diminish.  This  clearly  indicates  there  is  in  South- 
Eastern  Europe  a  powerful  centre  of  creation  of 
Clausilice,  from  which  the  species  have  spread  all 
over  Europe.  But  it  is  by  no  means  certain  that  this 
centre  was  always  in  our  continent,  for  in  South- 
Eastern  Asia  and  the  Malay  Archipelago  Clausilice 
increase  once  more.  It  is  interesting  to  note,  however, 
that  almost  all  these  eastern  forms  belong  to  the  sub- 
genus  Phcedusa  (vide  Boettger),  which  had  only  been 
known  as  a  fossil  genus  from  a  few  species  in  the 
Eocene  and  Oligocene  of  Southern  Europe.  The 
first  centre  of  origin,  therefore,  may  possibly  have 
been  in  Southern  Asia,  and  in  these  early  Tertiary 
limes  a  second  centre  may  have  become  established 
in  Southern  Europe  from  which  the  sub-genus  Gar- 
nieria  went  eastward,  Macroptychia  southward,  and 
Nenia  westward  across  the  Atlantis  to  South  America. 
Only  a  few  remnants  of  these  primitive  Clausilicz 
are  now  left  in  Europe,  such  as  the  interesting  Cl. 
(Laminifera]  Fault. 

As  an  example  of  a  genus  which  has  its  centre  of 
distribution  in  South-Western  Europe  we  might  take 
that  to  which  our  common  brown  garden  slug  belongs, 
viz.,  Arion.  Dr.  Simroth,  who  was  the  first  to  point 
out  that  the  species  of  Arion  had  spread  over  our  con- 
tinent from  South-Western  Europe  (p.  5),  is  inclined 
to  the  belief  that  the  Arionidce  had  originated  on  the 
old  land-bridge  between  Europe  and  North  America. 


PRELIMINARY  CONSIDERATIONS.  49 

which  is  generally  known  by  the  name  of  "  Atlantis." 
From  this  a  branch  went  westward  to  the  New 
World  and  another  eastward  as  far  as  Southern  Asia, 
but  Arion  and  a  number  of  other  genera  are  more  or 
less  confined  to  South-Western  Europe.  Only  a  few 
species  of  Anon  have  a  wide  range  in  Europe,  one  of 
them,  A.  subfuscus,  crossing  the  borders  of  our  con- 
tinent into  Siberia.  In  the  British  Islands  and  in 
Western  Germany,  which  are  about  equi-distant  from 
the  supposed  creative  centre  of  the  genus,  there  are 
found  five  species.  In  France  six  or  seven  species 
are  met  with,  and  in  Spain  and  Portugal  about  ten. 
Towards  the  east,  Arions  diminish  in  number.  This 
genus,  therefore,  forms  part  of  a  migration  which  I 
have  designated  as  "  Lusitanian "  from  Lusitania, 
the  name  applied  by  the  Romans  to  what  we 
now  call  Portugal.  Another  genus  of  slugs,  Geo- 
malacuS)  is  interesting  from  the  fact  that  one  species 
occurs  in  the  British  Islands,  being  otherwise  con- 
fined to  the  Lusitanian  province.  Parmacella,  a  slug- 
like  animal  bearing  a  tiny  shell  at  the  extremity  of  its 
tail,  has  probably  likewise  had  its  origin  in  this  part  of 
Europe.  All  this,  however,  will  be  more  fully  referred 
to  in  the  seventh  chapter,  which  deals  with  the 
Lusitanian  fauna. 

As  regards  the  Alpine  centre  of  origin,  Dr.  Kobelt 
considers  three  groups  of  mollusca  as  especially 
characteristic  of  the  Alps,  viz.,  the  sub -genus  Carnpylaea 
of  the  great  and  widely-spread  genus  Helix,  and  the 
genera  Pomatias  and  Zonites.  The  latter,  which  is  not 

4 


5O  HISTORY  OF   THE    EUROPEAN    FAUNA. 

to  be  confounded  with  our  British  Hyalinia  (formerly 
united  with  Zonites\  does  not  extend  very  far  south 
or  north  of  the  Alps.  There  may  be  others  too,  which 
owe  their  origin  to  these  mountains,  but  most  of 
the  terrestrial  mollusca  are  exceedingly  ancient,  and 
many  genera  have  existed  long  before  the  Alps  had 
made  their  appearance  above  the  surface  of  the  early 
Tertiary  seas.  It  should  be  remembered  that  Hya- 
linia and  Pupa,  both  British  genera,  are  known  from 
carboniferous  deposits  in  forms  which  closely  approach 
those  living  at  the  present  day,  and  in  these  and  a 
great  number  of  other  instances,  it  is  quite  impossible 
to  determine  the  original  home  of  the  genus. 

This  little  digression  on  centres  of  dispersion  will 
help  us  to  understand  in  what  manner  the  indigenous 
element  of  the  European  fauna  joined  in  with  the 
alien  members  as  they  arrived  in  our  continent.  The 
species  confined  to  South-Eastern  England  need  not 
necessarily  have  come  to  us  from  Eastern  Europe  or 
Siberia.  Alpine  species  spread  northward  probably 
at  the  same  time  as  the  Siberian  animals  went  west- 
ward. An  Alpine  form  may  therefore  have  joined  a 
batch  of  the  latter  and  entered  Eng-land  with  them. 
Even  a  Lusitanian  animal  may  have  mingled  with 
these  migrants,  so  that  all  three  elements  may  occur 
together  in  one  locality. 

But  these  are  exceptions.  The  migrations  have,  as 
a  rule,  not  joined  to  any  great  extent;  indeed,  all 
those  naturalists  who  have  carefully  examined  the 
problem  of  the  origin  of  the  European  fauna,  have  felt 


PRELIMINARY  CONSIDERATIONS.  51 

that  it  was  composed  of  elements  which  arrived  at 
different  times. 

The  great  Russian  naturalist,  the  late  Professor 
Brandt,  distinguished  five  phases  in  the  history  of  the 
Eurasian  mammalian  fauna  (pp.  249-254).  During  the 
first  phase — an  uncertain  period  of  long  duration — the 
mammals  held  intact  their  position  in  the  northern 
half  of  Asia.  The  Mammoth,  the  Hairy  Rhinoceros, 
Bison,  Musk  Ox,  Wild  Sheep,  Reindeer,  and  perhaps 
Tigers,  Hyaenas,  etc.,  lived  then,  with  numerous 
peculiar  Rodents,  under  such  climatic  conditions, 
according  to  Brandt,  that  they  were  able  to  extend 
their  range  along  with  tree  vegetation  to  the  extreme 
north  of  the  Asiatic  continent  This,  he  thinks,  seems 
to  have  been  the  case  especially  with  the  Reindeer, 
Mammoth,  Rhinoceros,  and  Musk  Ox.  The  second 
phase  was  characterised  by  the  dispersion  of  the 
Northern  Asiatic  mammalian  fauna  towards  Central, 
Southern,  and  Western  Europe,  and  this  period  lasted 
until  the  complete  extermination  of  the  Mammoth. 
The  third  phase  dates  from  the  time  when  the  Mam- 
moth and  the  Hairy  Rhinoceros  had  become  extinct, 
whilst  the  fourth  commenced  with  the  disappearance 
of  the  Reindeer  in  Europe,  and  terminated  when  the 
Wild  Ox  in  the  feral  state  had  become  unknown. 
Finally,  the  last  phase  constitutes  the  present  time. 
Lartet  held  similar  views,  and  also  believed  that 
Europe  was  peopled  by  successive  migrations  from 
Asia. 

Botanists   have    worked    at    the    problem    of    the 


52  HISTORY   OF  THE   EUROPEAN    FAUNA. 

European  flora  much  more  systematically,  and  our 
knowledge  of  the  origin  of  that  flora  has  been  greatly 
increased  within  the  last  twenty  years,  chiefly  by 
the  researches  of  Professor  Engler.  More  recently, 
detailed  studies  have  been  made  in  Scandinavia  by 
Professor  Blytt,  in  the  Alps  by  Dr.  Christ  and  Mr. 
Ball,  in  Germany  by  Professor  Drude,  Dr.  Schulz, 
and  many  others.  Dr.  Schulz  (p.  i)  is  of  opinion 
that  the  great  majority  of  the  European  plants  have 
either  migrated  to  or  have  originated  in  our  conti- 
nent since  the  beginning  of  the  Pliocene  epoch,  and 
that  the  original  home  of  the  immigrants  must  be 
looked  for  in  Asia  and  in  Arctic  America.  From  the 
latter  an  almost  uninterrupted  migration  must  have 
taken  place  during  the  greater  part  of  Tertiary  times 
up  to  the  commencement  of  the  Pliocene  epoch,  partly 
over  a  direct  land-connection  with  Europe  by  way  of 
Greenland,  Iceland,  and  the  Faroes,  and  also  vid 
Spitsbergen,  Franz  Josef  Land  and  Novaya  Zemlya, 
and  partly  by  an  indirect  one  across  the  Behring 
Straits  between  Alaska  and  Kamtchatka. 

A  good  deal  of  work  still  requires  to  be  done  before 
zoologists  have  acquired  the  same  intimacy  with  the 
European  fauna  as  botanists  have  with  the  flora. 
However,  the  view  that  our  animals  all  come  from 
Asia,  as  was  long  ago  believed,  has  been  abandoned 
for  some  time.  The  first  to  bring  under  the  notice 
of  naturalists  the  hypothesis,  that  there  must  have 
been  two  distinct  migrations  of  northern  animals  to 
Central  Europe — one  from  the  north,  and  another 


PRELIMINARY   CONSIDERATIONS.  53 

from  the  cast — was  the  late  Mr.  Bogdanov.  The 
Arctic  species,  of  which  remains  have  been  discovered 
in  the  Pyrenees — namely,  the  Reindeer,  Arctic  Hare, 
Willow  Grouse,  etc.,  he  thought  had  nothing  to  do 
with  those  which  invaded  Europe  from  Siberia  during 
the  Glacial  period.  He  maintained  that  the  former 
had  quite  a  distinct  origin,  and  came  from  Scandi- 
navia (p.  26). 

As  I  shall  deal  with  this  problem  more  fully  in  a 
subsequent  chapter,  I  need  only  mention  that  I  fully 
agree  with  the  view  expressed  by  Mr.  Bogdanov  that 
two  distinct  migrations  of  northern  species  to  Central 
Europe  can  be  traced. 

No  one,  I  think,  has  done  more  in  fostering  a  care- 
ful study  of  the  migrations  of  animals  than  our 
distinguished  geologist  Professor  Boyd  Dawkins.  He 
did  not  follow  Bogdanov  in  distinguishing  two  Arctic 
migrations;  however,  he  did  more  in  constructing  a 
very  ingenious  chart  (a,  p.  ill)  representing  the  geo- 
graphy of  Europe  during  the  last  and  most  recent 
geological  epoch — the  Pleistocene — and  indicating  on 
it  the  probable  extent,  during  that  time,  of  an  eastern 
and  a  southern  migration  of  mammals.  The  map  is 
very  instructive,  and  is  the  first  ever  published  giving 
a  clear  idea  of  a  southern  and  an  eastern  migration 
to  Europe.  He  believed  that  the  migration  of  the 
southern  mammals  northward,  took  place  conjunctly 
with  the  westward  movement  of  the  eastern  species. 
Having  once  reached  Europe,  the  southern  species 
are  supposed  to  have  passed  northward  in  summer 


54  HISTORY  OF  THE   EUROPEAN    FAUNA. 

time,  whilst  the  eastern  forms  (he  calls  them  northern) 
would  swing  southwards.  The  two  migrations  would 
thus  occupy,  at  different  times  of  the  year,  the 
same  tract  of  ground  (a,  p.  1 13).  From  the  mingling 
of  the  remains  of  the  Hyaena  with  those  of  the 
Reindeer  and  Hippopotamus  in  the  Kirkdale  Cavern, 
he  infers  that  the  former  preyed  upon  the  Reindecr 
at  one  time  of  the  year,  and  on  the  Hippopotamus  at 
another.  He  argues  that  in  such  a  manner  might 
be  explained  the  curious  mixture  of  northern  and 
southern  types  which  we  find  in  the  British  pleistocene 
and  in  cave  deposits. 

Besides  mammals,  the  only  European  animals 
which  have  received  some  attention  with  a  view  to  a 
study  of  their  origin,  are  the  Butterflies  and  the  Land- 
Snails.  The  entomologists  who  have  taken  up  the 
problem  have  in  so  far  scarcely  produced  satisfactory 
results,  as  they  all  seemed  to  be  bound  down  to  the 
hypothesis  that  practically  all  the  butterflies  had 
been  destroyed  in  Europe  during  the  Glacial  period. 
Hofman,  in  his  interesting  little  work,  comes  to  the 
conclusion  (p.  50),  that  only  in  Greece  and  Spain 
could  a  small  remnant  of  the  butterflies  have  survived 
the  extreme  rigours  of  climate.  Greece  was  at  that 
time  connected  with  Asia  Minor,  and  Spain  with 
North  Africa  ;  and  the  author  supposes  that  the  semi- 
alien  fauna  inhabiting  these  tracts  was  mainly  re- 
sponsible for  the  re-stocking  of  Southern  Europe,  but 
that  the  main  mass  of  our  butterflies  are  post-glacial 
Siberian  immigrants. 


PRELIMINARY   CONSIDERATIONS.  55 

The  work  published  by  Messrs.  Speycr  deals  only 
with  the  origin  of  the  Central  European  Butterflies. 
The  period  during  which  our  European  species 
originated  is  not  specified,  but  the  authors  believe 
that  they  had  their  home  either  in  Southern  Russia 
or  Central  Asia.  The  fact  that  the  number  of 
butterflies  decreases  very  considerably  as  we  pro- 
ceed north-westward  in  Europe  appears  to  them 
to  substantiate  these  views.  The  apparent  dislike 
evinced  by  butterflies  to  the  damp  Atlantic  Coast 
climate,  they  think,  clearly  indicates  that  they  had 
originated  in  a  dry  and  more  continental  climate. 
The  history  of  the  North  European  Butterflies 
and  Moths  has  been  carefully  described  by  Mr. 
Petersen.  He  adopts  Hofman's  theory  as  to  the 
almost  total  extinction  of  the  Lepidoptera  in 
Europe  during  the  Glacial  period.  The  chief  immi- 
gration to  Europe  after  that  period  is,  he  thinks, 
Siberian. 

At  first  there  appeared  species  which  belonged 
to  a  cold  climate,  and  which  now  live  in  ele- 
vated regions  ;  then  came  forms  suited  to  a  milder 
climate,  which  established  themselves  on  the  north- 
easterly slopes  of  the  Alps.  The  most  recent  addition 
which  our  continent  has  received  from  Siberia  is, 
according  to  Mr.  Petersen,  the  present  Scandinavian 
fauna.  Scandinavia  has  obtained  a  larger  number  of 
species  than  the  European  plain,  because  to  this  last 
migration  were  added  such  as  prefer  a  northern  or 
Alpine  climate. 


56  HISTORY  OF   THE   EUROPEAN    FAUNA. 

As  a  contribution  to  the  history  and  composition  of 
the  European  fauna,  by  far  the  most  important  work 
ever  published  is  that  of  Dr.  Kobelt,  the  eminent 
German  conchologist.  Whilst  the  researches  into  the 
origin  of  the  Lepidoptera,  above  described,  have  been 
marred  by  the  prevalent  prejudice  as  to  the  dele- 
terious effects  of  a  glacial  climate  on  the  butterflies, 
the  present  author  boldly  works  out  the  problem  on 
independent  lines.  He  shuns  theories  and  specula- 
tions almost  altogether.  His  great  work,  as  yet 
practically  unknown,  the  result  of  a  lifetime  of  the 
most  painstaking  labour,  ranks  among  the  most  im- 
portant contributions  to  zoogeography.  I  shall  have 
frequent  occasion  to  refer  to  it  throughout  these  pages. 
Meanwhile  some  of  his  more  remarkable  conclusions 
may  be  mentioned.  "  Comparing  all  classes  of 
animals  as  to  their  zoogeographical  importance,  the 
highest  rank  must  undoubtedly  be  accorded  to  the 
land-snails  "  (i.,  p.  7).  "  The  Pleistocene,  and  with  it 
the  land  and  fresh-water  molluscan  fauna  of  the 
present  day  has  been  gradually  evolved  from  the 
Tertiary  one,  and  its  roots  can  be  traced  through  the 
Cretaceous  to  the  Jurassic  epoch.  During  the  whole 
of  that  time  no  sudden  appearance  of  a  new  fauna 
can  be  demonstrated.  Quite  slowly,  step  by  step,  the 
Cretaceous  is  succeeded  by  the  Tertiary  fauna,  and 
one  after  the  other  of  the  characteristic  palaearctic 
genera  appear — first  the  fresh-water,  then  the  land 
forms"  (p.  141).  "The  division  of  the  North  Alpine 
from  the  South  Alpine  fauna  must  be  older  than  the 


PRELIMINARY   CONSIDERATIONS.  57 

Glacial  period  ;  and  the  present  Central  European 
fauna  had  already  become  developed  from  the  Plio- 
cene in  all  its  details  of  form  and  distribution  before 
the  commencement  of  the  Ice  Age"  (p.  162).  "We 
must  draw  the  conclusion  from  the  preceding  remarks, 
that  the  present  (palaearctic)  molluscan  fauna  in  its 
distribution  is  older  than  the  Glacial  period,  and  that 
the  latter  produced  merely  a  retreat  of  the  fauna 
from  the  most  inhospitable  regions  of  Europe  with 
a  subsequent  re-immigration,  but  did  not  cause  its 
destruction"  (i.,  p.  169). 

A  few  attempts  have  also  been  made  by  naturalists 
to  trace  the  origin  of  the  fauna  of  some  smaller 
European  areas.  Thus  Riitimeyer,  in  dealing  with 
the  mammalian  fauna  of  Switzerland,  remarks  (p.  31) 
"that  it  seems  certain  that,  in  spite  of  many  local 
disturbances,  the  continuity  of  generations  was  never 
interrupted  throughout  the  whole  of  the  Tertiary 
period  until  the  present  day." 

An  even  more  interesting  memoir  is  that  of  Mr. 
Koppen  on  the  origin  of  the  Crimean  fauna.  It  is 
only  recently,  according  to  this  author,  that  this  pen- 
insula has  become  connected  with  Southern  Russia. 
And  it  is  for  this  reason  that  the  Squirrel  and  a  number 
of  other  animals,  and  also  plants,  present  in  Russia, 
are  absent  from  the  Crimea.  Originally  the  latter  prob- 
ably formed  a  westward  continuation  of  the  Caucasus, 
and  at  that  time  it  was  surrounded  by  the  sea  on 
all  other  sides.  "  Much  later,"  he  continues,  "  after 
and  in  consequence  of  a  local  subsidence,  the  country 


$8  HISTORY  OF   THE   EUROPEAN   FAUNA. 

between  the  Caucasus  and  the  Crimea  became 
interrupted.  The  latter  existed  for  a  long  time  as 
an  island,  and  only  much  later,  in  recent  geological 
times,  did  it  become  united  with  Southern  Russia' 
by  means  of  the  isthmus  of  Perekop." 

There  is,  on  the  whole,  a  great  diversity  of  opinion 
as  to  how  the  European  fauna  has  originated  ;  how- 
ever, except  in  Dr.  Kobelt's  work,  no  attempt  has 
hitherto  been  made  to  collect  together  all  the  available 
information,  and  to  include  in  the  inquiry  more  than 
one  class  of  animals.  The  little  work  which  I  venture 
to  bring  before  the  public  will  not  by  any  means  ex- 
haust the  subject,  nor  is  our  knowledge  of  the  Euro- 
pean fauna  sufficient  to  give  more  than  a  mere  sketch 
of  many  of  the  animal  groups  mentioned.  As  we 
have  learned  in  the  introduction,  different  classes  of 
animals  are  not  all  of  equal  importance  in  indicating 
the  changes  which  have  taken  place  in  the  distribution 
of  land  and  water.  While  Dr.  Kobelt  is  of  opinion 
that  the  land-snails  are  by  far  the  most  important 
in  such  an  inquiry,  Mr.  Lydekker  believes  that 
mammals  afford  the  safest  and  truest  indications 
of  such  changes.  Mr.  Beddard  puts  in  a  claim  for 
earthworms,  as  even  a  narrow  strait  of  sea-water 
forms  an  insuperable  barrier  to  their  dispersion. 
Dr.  Wallace  agrees  with  Mr.  Lydekker,  and  goes 
so  far  as  to  say  (p.  74)  that  "  whenever  we  find 
that  a  considerable  number  of  the  mammals  of  two 
countries  exhibit  distinct  marks  of  relationship,  we 
may  be  sure  that  an  actual  land-connection,  or  at 


PRELIMINARY   CONSIDERATIONS.  59 

all  events  an  approach  to  within  a  very  few  miles 
of  each  other,  has  at  one  time  existed."  Besides 
the  groups  referred  to,  I  claim  that  particular 
attention  should  be  devoted  to  Amphibia,  which, 
contrary  to  Wallace,  I  hold  do  not  possess  special 
facilities  for  dispersal ;  and  also  to  spiders  and  to 
all  wingless  animals  leading  a  subterranean  life,  such 
as  some  of  the  wood-lice,  planarian  worms  and 
apterous  beetles. 

A  thorough  knowledge  of  the  changes  in  the  dis- 
tribution of  land  and  water  is  desirable  in  order  to 
appreciate  the  extent  and  variations  of  former  mi- 
grations. A  study  of  the  British  fauna,  for  example, 
teaches  us  that  the  British  Islands  were  once  con- 
nected with  one  another  and  with  the  continent  of 
Europe  between  England  and  France.  It  was 
Professor  James  Geikie,  I  believe,  who  first  pointed 
out,  many  years  ago,  that  the  area  now  covered  by 
the  Irish  Sea  was  formerly  in  all  probability  a  fresh- 
water lake.  This  had  its  outlet  at  the  southern  ex- 
tremity in  the  form  of  a  stream  into  which  most  likely 
flowed  the  smaller  rivers  from  the  south-east  of 
Ireland,  and  which  was  joined  from  the  east  by  the 
Severn,  and  finally  debouched  into  the  Atlantic 
(Fig.  4).  The  range  in  the  British  Islands  of  those 
species  which  have  migrated  to  them  from  the  south, 
indicates  that  whilst  the  Atlantic  Ocean  had  gradually 
crept  up  and  flooded  the  area  between  Ireland  and 
Wales,  and  had  turned  the  fresh-water  lake  into  a  bay, 
communication  between  Scotland  and  Ireland  was 


6o 


HISTORY   OF   THE   EUROPEAN    FAUNA. 


still  possible.     The  occurrence  of  many  Scandinavian 
species  in  Scotland  which  are  absent  on  the  continent 


^r 


FIG.  4.  —  Map  of  the  British  Islands  and  surrounding  area  at  a  time 
when  the  earlier  members  of  the  southern  migration  reached 
England.  (Only  some  of  the  rivers  have  been  indicated.  The 
shaded  parts  represent  water,  the  light  land.) 

of  Europe,   indicates    that    these  two  countries   also 
were  united  formerly.     Most  geologists  hold  that  such 


PRELIMINARY  CONSIDERATIONS.  6 1 

a  connection,  if  it  existed,  must  have  broken  down 
in  Pliocene  times.  Professor  Judd,  however,  has  ex- 
pressed his  belief  (p.  1008)  that  it  still  existed  until 
after  the  appearance  of  man  in  Northern  Europe,  and 
that  our  forefathers  might  have  been  able  to  walk 
dry  foot  from  Scotland  to  Norway. 

I  shall  also  show  on  distributional  evidence,  in  the 
fourth  chapter,  that  until  recent  geological  times 
Scandinavia  was  continued  northward,  by  way  of 
Bear  Island,  with  Spitsbergen  and  probably  Franz 
Josef  Land,  which  islands  again  were  joined  with 
North  Greenland  and  Arctic  North  America,  and 
that  the  polar  fauna  and  flora  were  able  to  spread  on 
this  land-connection  to  both  America  and  Europe. 

That  Gibraltar  was  connected  with  Morocco,  and 
Sicily  with  Southern  Italy  and  Greece  on  the  one 
hand,  and  with  Tunis  on  the  other,  is  more  generally 
recognised;  whilst  Professor  Suess  has  shown  (vol.  i., 
p.  442),  on  purely  geological  grounds,  that  the 
Egean  Sea  was  dry  land  up  till  quite  recently — 
certainly,  he  thinks,  till  after  the  appearance  of  man. 
This  supposition  enables  us  to  understand,  as  will  be 
more  fully  discussed  in  the  sixth  chapter,  how  the 
Oriental  fauna  entered  Europe.  Such  minor  zoo- 
geographical  problems  as  the  occurrence  of  the  Wild 
Goat  of  Asia  Minor  {Capra  cegagtus)  on  the  islands  of 
Crete  and  on  some  of  the  Cyclades  now  almost 
explain  themselves.  The  Sea  of  Marmora  is  prob- 
ably a  modern  formation,  so  that  Asia  Minor  ex- 
tended not  long  ago  beyond  the  Turkish  capital,  but 


62  HISTORY  OF   THE   EUROPEAN    FAUNA. 

Dr.  Kobelt  believes  that  an  arm  of  the  Black  Sea 
communicated  up  till  recent  times  along  the  lower 
course  of  the  Maritza  with  the  Gulf  of  Saros. 
It  can  be  shown  also  that  Sardinia  and  Corsica 
formed  part  of  the  continent  of  Europe,  and  that 
their  present  fauna  and  flora  reached  them  by  migra- 
tion on  land. 

The  Russian  naturalists,  Brandt  and  Koppen,  be- 
lieved that  at  no  very  distant  date  a  sea  extended  right 
across  Eastern  Russia  from  the  Caspian  to  the  Arctic 
Ocean,  whilst  Professor  Boyd  Dawkins  expressed 
himself  in  very  similar  language  as  follows  (c,  p.  35) : 
"  Before  the  lowering  of  the  temperature  in  Central 
Europe  the  sea  had  already  rolled  through  the  low 
country  of  Russia,  from  the  Caspian  to  the  White 
Sea  and  the  Baltic,  and  formed  a  barrier  to  western 
migration  to  the  Arctic  mammals  of  Asia."  These 
naturalists  based  their  opinions  on  distributional 
evidence,  but  additional  facts  will  be  brought 
forward  in  the  fifth  chapter  to  substantiate  these 
views. 

These  are  some  of  the  more  important  geographical 
events  which  will  be  dealt  with  in  detail  in  the  subse- 
quent chapters  in  connection  with  the  history  of  the 
migrations  of  the  European  fauna. 

A  separate  chapter  has  been  devoted  to  the  British 
fauna  and  its  origin,  since  it  plays  a  very  important 
part  in  the  evolution  of  that  of  our  continent.  So 
essential  is  a  thorough  knowledge  of  this  fauna,  that 
I  think  it  would  be  difficult  to  understand,  without 


PRELIMINARY  CONSIDERATIONS.  63 

it,  the  main  features  of  the  great  migrations  ;  and  I 
have  before  now  expressed  the  opinion  that  the 
British  fauna  forms  the  key  to  the  solution  of  the 
problem  of  the  origin  of  European  animals.  WTe 
know  that  our  British  species  came  to  us  by  land 
— at  least  the  bulk  of  them.  But  we  want  to 
know  what  direction  they  came  from,  and  at  what 
time  they  arrived.  When  Ireland  became  discon- 
nected from  Great  Britain,  and  the  latter  from 
Scandinavia  and  France,  is  another  interesting 
problem.  Professor  Boyd  Dawkins  has  indicated 
to  us  a  method  of  the  special  line  of  research  to 
meet  such  inquiries.  "  The  absence,"  he  says  (by 
p.  xxix),  "  of  the  beaver  and  the  dormouse  from  Ire- 
land must  be  due  to  the  existence  of  some  barrier 
to  their  westward  migration  from  the  adjacent  main- 
land, and  the  fact  that  the  Alpine  hare  is  indi- 
genous, while  the  common  hare  is  absent,  implies 
that,  so  far  as  relates  to  the  former  animal,  the  barrier 
did  not  exist." 

Many  members  of  the  great  Siberian  invasion 
reached  England,  but  Ireland  remained  entirely 
free  from  these  migrants.  The  assumption  there- 
fore seems  not  unreasonable,  that  the  latter  country 
at  the  time  of  their  arrival  was  no  longer  joined 
to  England.  The  great  bulk  of  the  Irish  fauna 
is  composed  of  Lusitanian,  Alpine,  and  Oriental 
immigrants,  and  there  is  besides  a  distinctly  Arctic  or 
North  American  element.  All  these,  of  course,  must 
have  established  themselves  in  Ireland  before  the 

^^SJB*AJT^. 
IWIVETR*SITY 


64  HISTORY   OF   THE   EUROPEAN    FAUNA. 

Siberian  fauna  set  foot  in  England,  since  it  has  been 
shown  that  a  continuous  land-surface  was  necessary 
for  their  migration.  Owing  to  the  perfect  preserva- 
tion of  the  remains  of  the  Siberian  migrants  in  recent 
continental  deposits,  the  history  of  that  migration 
can  be  clearly  followed,  and  it  is  possible  even  to 
determine  the  date  of  its  arrival  in  England — in 
geological  language  at  any  rate.  The  time  of  the 
colonisation  of  Ireland  can  be  thus  approximately 
fixed  as  having  taken  place  at  a  period  prior  to  the 
arrival  of  the  Siberian  migrants  in  England. 

All  those  who  have  seriously  studied  the  problems 
presented  by  our  British  fauna — notably  the  late 
Professor  Forbes,  and  more  recently  Mr.  Carpenter 
and  myself — are  agreed  that  the  Lusitanian  element 
is  the  oldest,  and  that  the  newest  is  that  which  has 
come  to  us  from  the  east. 

The  sequence  of  events  in  the  British  Islands  was 
probably  as  follows  : — The  first  comers  were  the 
members  of  that  fauna  which  issued  from  South- 
western Europe;  then  came  the  Alpine,  and  at  the 
same  time  probably  the  Arctic  and  the  Oriental; 
and  finally  the  Eastern  or  Siberian.  The  migrations 
of  all  but  the  last  continued,  uninterruptedly,  for  very 
long  periods. 

The  study  of  these  migrations  has  convinced  me 
that,  though  climate  was  a  powerful  factor  in  the 
evolution  or  history  of  the  European  fauna,  the 
geographical  changes  which  took  place  on  our  con- 
tinent in  later  Tertiary  times  exerted  a  yet  stronger 


PRELIMINARY   CONSIDERATIONS.  65 

influence.  The  principal  climatic  disturbance  is 
generally  supposed  to  have  been  the  so-called  "  Ice 
Age."  So  firmly  rooted  is  the  conviction,  among 
naturalists  of  the  present  day,  of  the  enormous 
destruction  which  this  period  produced  on  our 
European  fauna,  so  that  all  animal  life  practically 
disappeared  from  large  areas  of  our  continent,  that 
it  is  desirable  that  we  should  now  shortly  review 
the  history  of  that  remarkable  period  in  order  to 
ascertain  in  how  far  these  views  are  corroborated 
by  facts.  Frequent  reference,  moreover,  will  be 
made  throughout  this  work  to  the  theories  con- 
nected with  the  Glacial  period. 

It  has  been  stated  by  an  eminent  geologist  that 
during  part  of  the  Glacial  period  the  climate  was  such 
that  neither  plants  nor  animals  could  have  existed  in 
the  British  Islands.  If  that  had  been  so,  it  is  evident 
that  very  few  organisms  could  have  even  survived  in 
France,  though  a  number  of  Arctic  species  might 
have  dragged  on  an  existence  in  Southern  Europe. 
At  any  rate,  on  the  return  of  more  genial  conditions, 
the  Arctic  species  would  undoubtedly  have  been  the 
first  to  gain  admission  to  the  British  Islands,  to 
re-people  the  arid  wastes.  Our  supposition  that  the 
Lusitanian  element  in  the  British  fauna  is  the  oldest 
would  therefore  be  wrong.  From  early  Tertiary 
times  onward,  the  climate  of  Europe,  which  was 
then  semi-tropical,  gradually  became  more  and  more 
temperate;  until  finally  the  Ice  Age  or  Glacial 
period  arrived,  during  which,  according  to  Professor 

5 


66  HISTORY   OF   THE   EUROPEAN   FAUNA. 

J.  Geikie — one  of  our  highest  authorities  on  this 
subject — a  great  part  of  Northern  Europe  became 
practically  uninhabitable  owing  to  the  severity  of 
the  climate. 

To  enable  us  to  judge  better  of  the  true  value  of 
the  many  hypotheses  which  have  been  advanced 
to  account  for  this  supposed  extraordinary  fall  of 
temperature  during  the  "Ice  Age,"  we  must  compare 
the  views  of  other  authorities  with  the  one  just 
quoted.  I  do  not  propose  to  discuss  the  causes  which 
have  led  to  the  production  of  the  Glacial  period — 
those  interested  in  these  questions  should  consult 
the  writings  of  Dr.  Croll,  Professor  J.  Geikie,  Pro- 
fessor Bonney,  Mr.  Falsan,  and  others — but  merely  to 
give  the  climatic  aspects  from  a  physical,  zoological, 
and  botanical  point  of  view. 

According  to  Professor  Penck  (a,  p.  12),  the  nature 
of  the  glacial  climate  can  be  determined  by  comparing 
the  snow-line  of  the  Glacial  period  with  that  of  the 
present  day.  The  position  of  the  snow-line  is  de- 
pendent on  two  climatic  factors — viz.,  precipitation 
and  temperature.  We  know  the  height  at  which 
snow  must  have  lain  permanently  during  the  Glacial 
period,  or  during  the  maximum  phase  of  glaciation. 
If  the  Ice  Age  had  been  produced  solely  by  an 
increase  of  snowfall,  as  has  been  suggested,  Professor 
Penck  tells  us  that  then  it  must  have  snowed  three  or 
four  times  as  much  as  it  does  now.  But  he  does  not 
adopt  the  view  that  the  Ice  Age  is  due  to  an  increase 
of  snowfall  alone.  His  calculations,  based  upon  the 


PRELIMINARY   CONSIDERATIONS.  67 

height  of  the  snow-line,  tend  to  show  that  a  general 
decrease  of  temperature  to  the  extent  of  from  4-5 
degrees  Centigrade  (all  other  atmospheric  conditions 
remaining  the  same  as  now)  would  be  sufficient  to 
give  us  back  the  Glacial  period. 

Professor  Neumayr  (p.  619)  adopted  a  similar  prin- 
ciple in  determining  the  temperature  which  prevailed 
in  Europe  during  the  Glacial  period.  Snow  now  lies 
in  the  Pyrenees  1000  metres  higher 'than  it  did  then, 
1,200  metres  higher  in  the  Alps,  and  800  metres 
higher  in  the  Tatra  mountains.  Since  the  tempera- 
ture in  Central  Europe  decreases  by  half  a  degree 
Centigrade  for  every  100  metres  of  elevation,  it  follows 
that  if  the  glacial  phenomena  had  only  been  brought 
about  by  a  decrease  of  temperature  without  an  in- 
crease of  moisture,  we  should  have  had  a  reduction 
of  temperature  during  the  Glacial  period  of  six 
degrees  Centigrade  in  the  Pyrenees,  of  seven  degrees 
in  the  Alps,  and  of  four  in  the  Tatra  mountains. 
The  general  lowering  of  the  temperature  of  Europe, 
says  Professor  Neumayr,  could  not  have  amounted  to 
more  than  six  degrees  Centigrade.  Moreover,  he  is 
of  opinion  that  the  very  low  snow-line  in  the  British 
Islands  proves  that  even  during  the  Ice  Age  a  com- 
paratively mild  climate  prevailed  there,  and  that 
the  climatic  conditions  generally,  in  the  different 
parts  of  Europe,  were  relatively  about  the  same  as 
they  are  now. 

Professor  J.  Geikie  does  not  give  us  his  views  as  to 
the  temperature  of  the  Glacial  period,  but  he  main- 


68  HISTORY   OF   THE   EUROPEAN   FAUNA. 

tains  that  a  lowering  of  the  temperature  is  evinced 
not  only  by  the  widespread  phenomena  of  glaciation, 
but  by  the  former  presence  in  our  temperate  latitudes 
of  a  northern  fauna  and  flora. 

Mr.  Charles  Martins,  who  based  his  calculations 
on  the  temperature  during  the  Glacial  period  on 
the  glaciers  of  Chamounix,  concluded  that  it  only 
needed  a  lowering  of  the  temperature  to  the  extent 
of  four  degrees  Centigrade  to  bring  the  glaciers  down 
to  the  plain  of  Geneva,  and  in  fact  give  us  back  the 
Glacial  period.  It  need  not  surprise  us,  therefore, 
that  the  French  geologist,  Mr.  Falsan,  the  author 
of  La periode  glaciere,  is  of  opinion  (p.  230)  that  the 
mean  annual  temperature  of  France  during  the 
Glacial  period  was  approximately  from  6-9  degrees 
Centigrade,  perhaps  more.  Close  to  the  immense 
glaciers  of  the  Rhone,  it  might  have  been  about 
six  degrees.  This  is  the  actual  mean  annual  tempera- 
ture of  the  South-west  of  Sweden  and  Norway,  or 
the  North  of  Scotland. 

Although  all  these  investigations  tend  to  show 
that  the  climate  of  Europe  during  the  Glacial  period 
was  by  no  means  so  severe  as  we  are  often  led  to 
believe,  yet  there  exists  also  a  school  of  geologists 
who  maintain  there  was  actually  a  higher  temperature 
than  at  present.  The  inconsistency  of  mentioning 
heat  in  connection  with  ice  and  snow  is  more 
apparent,  however,  than  real,  for  we  must  remember 
Tyndall's  original  remark  on  this  subject.  It  is  the 
snow,  he  says,  which  feeds  the  glaciers.  But  the 


PRELIMINARY  CONSIDERATIONS.  69 

snow  comes  from  the  clouds,  and  these  again 
originate  from  the  vapours  which  the  sun  causes 
to  be  absorbed  from  the  ocean.  Without  the  sun's 
heat,  we  should  have  no  water  vapour  in  the 
atmosphere ;  without  vapour,  no  clouds ;  without 
clouds,  no  snow ;  without  snow,  no  glaciers.  The 
ice  of  glaciers,  therefore,  owes  its  origin  indirectly 
to  the  sun's  heat.  It  has  been  supposed  that  if  the 
sun's  heat  diminished,  larger  glaciers  would  form 
than  those  existing  to-day,  but  the  diminution  of  the 
solar  heat  would  infallibly  reduce  the  amount  of 
water  vapour  in  the  air,  and  would  thus  stop  the 
very  source  of  glaciers. 

Mr.  Falsan  even  admits  that  without  a  change  of 
the  mean  annual  temperature  (p.  201)  of  Europe, 
the  central  portions  of  our  continent  might  at  this 
period  have  enjoyed  an  insular  climate.  This  more 
equable  and  humid  climate  could,  within  certain 
limits,  favour  the  development  of  the  ancient  glaciers 
by  increasing  the  snowfall  and  slackening  the  summer 
rate  of  melting. 

It  seems  evident  then,  according  to  these  views, 
that  with  a  comparatively  slight  change  of  the 
atmospheric  conditions  in  the  British  Islands,  we 
might  have  glaciers  back  again  on  all  our  highest 
mountain  ranges  in  England,  Scotland,  and  Ireland. 
But  a  widespread  belief  seems  to  prevail  that  the 
presence  of  glaciers  implies  a  very  low  temperature. 
Snow  and  ice,  however,  are  formed  as  soon  as  the 
temperature  falls  below  freezing  point;  it  does  not 


7O  HISTORY   OF   THE   EUROPEAN   FAUNA. 

matter  whether  there  be  I  or  20  degrees  of  cold. 
Winters  with  a  few  degrees  of  frost  will  be  just  as 
favourable  for  the  growth  of  glaciers  as  winters  with 
the  most  severe  cold. 

Let  us  now  see  what  the  fauna  and  flora,  as  far 
as  we  know  it,  tell  us  of  the  climate  of  the  Glacial 
period.  At  the  very  outset  of  our  inquiry  we  are 
confronted  with  one  very  serious  difficulty  in  the 
problem,  and  that  is  the  supposed  occurrence  of  inter- 
glacial  mild  phases  alternating  with  colder  ones  during 
the  Ice  Age.  At  first,  when  traces  of  a  temperate  flora 
and  fauna  were  discovered  intercalated  between  two 
layers  of  boulder  clay,  their  presence  was  explained 
by  the  supposition  of  a  mild  inter-glacial  period. 
The  famous  Forest-bed  on  the  east  coast  of  England 
was  also  pronounced  to  be  an  inter-glacial  deposit, 
though  not  coming  precisely  under  this  definition. 
In  a  few  places  one  such  bed  was  found,  in  some  two 
or  more,  and  in  others  none  at  all.  Professor  James 
Geikie  discovered  the  evidences  of  no  less  than  five 
of  such  inter-glacial  epochs  (p.  612)  in  Europe. 
Lest  a  reader  of  that  author's  remarkable  work  on 
the  Ice  Age  might  carry  with  him  the  idea  that  his 
hypotheses  had  met  with  general  acceptance,  a  few 
quotations  from  almost  equally  high  authorities  on 
glacial  matters  will  be  useful.  "That  the  glaciers," 
remarks  Professor  Bonney  (p.  245),  "  were  liable  to 
important  oscillations  seems  to  be  proved,  but  whether 
the  evidence  suffices  to  establish  inter-glacial  epochs, 
in  the  usual  sense  of  the  words,  is  more  doubtful. 


tRELIMiNARY   CONSIDERATIONS.  ?I 

When  the  snow-fields,  as  in  the  Alps,  were  much 
more  extensive  than  they  are  at  present,  the  glaciers 
which  radiated  from  them  would  be  more  sensitive 
to  minor  climatal  change.  Even  now  they  oscillate 
considerably.  But  during  a  Glacial  epoch,  an  inch, 
either  more  or  less,  of  precipitation  might  mean  a 
considerable  advance  or  retreat  of  the  ice  in  the  low- 
lands." French  geologists  look  with  even  less  favour 
on  Professor  Geikie's  theories.  Mr.  Falsan  (p.  212) 
says  that  he  agrees  with  Messrs.  Favre,  de  Saporta, 
Lory,  de  Mortillet,  Desor,  de  Lapparent,  Lortet, 
Chantre,  Benoit,  Fontannes,  Deperet,  and  many 
other  geologists,  that  there  was  only  a  single  Glacial 
period,  which,  according  to  each  particular  region, 
might  be  divided  into  several  phases,  or  into  their 
equivalents — viz.,  one  or  more  extensions  of  the 
ancient  glaciers.  But,  on  the  whole,  the  view  that 
there  was  at  least  one  inter-glacial  phase  in  the 
Glacial  period  meets  with  more  general  acceptance 
among  geologists,  I  think,  though  the  other  opinion 
agrees  much  better  with  the  nature  of  the  fauna 
and  flora  as  it  has  been  revealed  to  us  from  the 
pleistocene  deposits. 

The  occurrence  of  the  remains  of  such  arctic  species 
of  mammals  as  the  Musk-Ox,  Arctic  Fox,  Glutton, 
Lemming,  and  many  others  in  these  deposits,  is 
frequently  held  up  to  us  by  geologists  as  a  proof 
of  the  prevalence  of  an  arctic  climate  while  these 
beds  were  laid  down.  And  indeed  this  appears 
at  first  a  most  satisfactory  explanation  of  the 


^2  HISTORY   OF   THE   EUROPEAN   FAUNA. 

phenomenon.  But  we  must  not  judge  the  climate 
of  Europe  by  their  presence  alone.  As  I  shall 
explain  more  fully  in  Chapter  V.,  these  species 
invaded  Europe  owing  to  two  circumstances.  Firstly, 
because  the  climate  of  Siberia  was  becoming  colder, 
necessitating  a  southward  movement,  with  a  con- 
sequent over-population  in  a  reduced  area;  secondly, 
because  a  new  short  route  to  Europe  had  been 
opened  up  for  them  about  the  same  time  (see 
p.  221).  An  invasion  of  Europe  therefore  took  place 
from  east  to  west.  Similar  invasions  occur  even  at 
the  present  day,  though  not  caused  by  a  change  in 
our  climate,  for  every  now  and  then  immense  flocks 
of  the  Siberian  Sandgrouse  emigrate  to  our  continent. 
The  mammalian  migrants  referred  to  are  not  to  be 
looked  upon  as  constituting  the  whole  of  our  fauna 
at  that  time.  Europe  had  a  fauna  of  its  own,  and 
these  invaders  merely  mingled  with  our  animals. 
There  was,  no  doubt,  a  keen  struggle  for  existence, 
as  the  result  of  which  the  weaker  in  many  cases 
succumbed.  The  hypothesis,  however,  that  these 
Siberian  migrants  occupied  an  empty  continent, 
forsaken  by  its  pre-glacial  inhabitants,  is  not  sup- 
ported by  any  facts. 

All  those  who  have  investigated  the  pleistocene 
fauna  have  been  struck  by  the  extraordinary  mixture 
of  northern  and  southern  types  of  animals.  Professor 
Dawkins  attempted  to  explain  these  facts  by  the  sup- 
position (p.  113)  that  "in  the  summer  time  the  southern 
species  would  pass  northwards,  and  in  the  winter 


PRELIMINARY  CONSIDERATIONS.  73 

time  the  northern  would  sway  southwards,  and  thus 
occupy  at  different  times  of  the  year  the  same 
tract  of  ground,  as  is  now  the  case  with  the  elks 
and  reindeer."  "  In  some  of  the  caverns,"  he 
continues  (p.  114),  "such  as  that  of  Kirkdale,  the 
hyaena  preyed  upon  the  reindeer  at  one  time  of  the 
year,  and  the  hippopotamus  at  another." 

A  similar  mingling  of  northern  and  southern 
faunas  has  also  been  observed  in  France.  Mr. 
Falsan  tells  us  (p.  236),  that  the  remains  of  the 
mammals  gathered  and  determined  by  Lartet  and 
Gaudry  belong  partly  to  species  which  have  been 
wrongly  regarded  as  indications  of  a  severe  climate, 
and  partly  to  such  as  are  accustomed  to  a  relatively 
mild  temperature.  In  several  localities  in  France, 
viz.,  at  Levallois,  St.  Acheul,  and  Arcy,  the  remains  of 
the  Hippopotamus  have  occurred  together  with  those 
of  the  Reindeer;  whilst,  according  to  Sir  H.  Howorth, 
the  Lion  has  been  found  together  with  northern  Voles 
at  Bicetre,  near  Paris.  It  is  stated  by  the  same 
authority  (p.  115)  that  much  the  same  conditions 
exist  in  Germany.  "The  lion  and  the  spotted  hyaena, 
the  mammoth  and  rhinoceros,  were  found  with  the 
marmot,  the  suslik,  the  lemming,  the  pica,  and  the 
reindeer."  At  another  locality  near  Thiede,  remains 
of  the  Mammoth,  woolly  Rhinoceros,  Horse,  Ox, 
Reindeer,  Arctic  Fox,  Lemming,  and  Pica  are  met 
with  in  the  same  deposit.  In  quoting  the  presence 
of  these  northern  animals  in  Europe  as  evidence  of  an 
arctic  climate,  we  commit  a  fatal  mistake.  Indeed, 


74  HISTORY   OF   THE   EUROPEAN    FAUNA. 

breeders  of  animals  and  those  acquainted  with  zoo- 
logical gardens  know  perfectly  well  that  it  is  much 
easier  to  keep  a  northern  species  in  a  southern 
climate,  than  a  southern  species  in  a  northern  one. 
If  in  a  Central  European  deposit  occur  a  mixture  of 
northern  and  southern  forms  of  animals,  the  presence 
of  the  latter  is  more  remarkable  than  that  of  the 
former.  Logically,  we  should  look  upon  the  occurrence 
of  southern  species  in  the  north,  therefore,  as  support- 
ing the  view  that  a  mild  climate  had  induced  them 
to  travel  northward.  The  only  indication,  indeed, 
of  the  presence  of  a  Monkey  in  the  British  Isles 
in  former  times  comes  to  us  from  the  very  same 
strata  which  have  also  yielded  the  remains  of  the 
Siberian  mammals. 

Before  I  conclude  the  consideration  of  the  pleisto- 
cene fauna,  it  may  be  of  interest  to  hear  what 
Mr.  Lydekker,  one  of  our  highest  authorities 
on  fossil  mammals,  has  to  say  on  this  subject. 
"  The  most  remarkable  feature  connected  with 
this  fauna  is  the  apparently  contradictory  evidence 
which  it  affords  as  to  the  nature  of  the  climate 
then  prevalent.  The  Glutton,  Reindeer,  Arctic 
Fox,  and  Musk-Ox  are  strongly  indicative  of 
a  more  or  less  arctic  climate;  many  of  the  Voles 
{Micr<otus\  Picas  (Lagomys),  and  Susliks  {Sper- 
mophilus\  together  with  the  Saiga  Antelope, 
appear  to  point  equally  strongly  to  the  prevalence 
of  a  Steppe-like  condition  ;  while  the  Hippopotamus 
and  Spotted  Hyaena  seem  as  much  in  favour  of  a 


PRELIMINARY  CONSIDERATIONS.  75 

sub-tropical  state  of  things.  Many  attempts  have 
been  made  to  reconcile  these  apparently  contra- 
dictory circumstances ;  one  of  the  older  views  being 
that  while  the  tropical  types  of  animals  lived  during 
a  warm  interlude,  they  migrated  southwards  with 
the  incoming  of  colder  conditions  to  the  arctic  type 
of  fauna.  Since,  however,  it  has  now  been  ascertained 
that  the  remains  of  both  tropical  and  arctic  forms 
have  been  found  lying  side  by  side  in  the  same  bed, 
it  is  perfectly  certain  that  such  an  explanation  will 
not  meet  the  exigencies  of  the  case  "  (p.  300). 

In  Germany  the  remains  of  the  Siberian  mammals 
occur  to  a  large  extent  in  a  pleistocene  deposit  known 
as  "  loess,"  and  the  theory  has  of  late  years  gained 
ground  that  the  latter  is  the  fine  dust-like  sand 
accumulated  during  an  intensely  arctic  dry  climate. 
That  many  of  the  mammals  discovered  in  the  "  loess" 
now  inhabit  the  dry  steppes  of  Eastern  Europe  and 
North-Western  Asia  seems  to  lend  support  to  this 
supposition;  but  besides  the  mammals  there  are  also 
land  and  freshwater  shells  in  this  deposit.  The  mol- 
luscan  fauna  certainly  indicates  no  steppe-character, 
according  to  Dr.  Kobelt  (b,  i.  p.  166). 

The  attempt  to  utilise  the  Siberian  migrants  to 
Europe  as  indicators  of  a  severe  climate  there,  cer- 
tainly fails  to  establish  conviction.  But  it  may  be 
asked,  surely  the  remains  of  the  Alpine  and  Arctic 
plants  which  have  been  found  in  pleistocene  deposits 
must  decide  this  question  in  favour  of  one  or  the 
other  hypothesis?  Let  us  test  it. 


76  HISTORY  OF  THE   EUROPEAN   FAUNA. 

Plants  being  more  directly  affected  than  animals 
by  changes  of  temperature  and  rainfall,  remarks 
Mr.  Clement  Reid  (p.  185),  give  evidence  of  the 
highest  value  when  we  inquire  into  former  climatic 
conditions.  The  severity  of  the  climate  during  the 
Glacial  period  is  often  assumed  from  the  occurrence 
in  pleistocene  strata  of  such  plants  as  Dryas  octopetala^ 
some  species  of  willow,  the  dwarf  birch,  and  others, 
which  are  now  found  in  high  latitudes  and  in  the  Alps, 
but  are,  as  a  rule,  absent  from  the  plain  of  Northern 
Europe.  Professor  J.  Geikie  goes  so  far  as  to  state 
(p.  398)  that  it  was  unlikely  that  southern  England 
during  the  climax  of  the  glacial  cold  had  much  if  any 
vegetation  to  boast  of,  and  continues,  "  It  is  certain, 
however,  that  it  was  clothed  and  peopled  by  an  Arctic 
flora  and  fauna  when  the  climatic  conditions  were 
somewhat  less  severe,  relics  of  that  flora  having  been 
detected  at  Bovey  Tracey."  He  believes,  therefore, 
that  an  Arctic  flora  took  possession  of  England  as 
soon  as  the  climate  enabled  it  to  live  in  the  country. 
Arctic  plants,  according  to  this  explanation  of  the 
sequence  of  events,  were  the  first  immigrants  to 
reconquer  the  dreary,  plantless  wastes  and  make 
them  habitable  for  mammals. 

Fortunately  these  views  do  not  at  all  agree  with 
those  of  many  of  our  leading  European  botanists  and 
others  entitled  to  have  a  voice  in  the  matter.  Pro- 
fessor Warming  is  of  opinion  that  the  main  mass  of 
the  present  flora  of  Greenland  survived  the  Glacial 
period  in  that  country  (p.  403);  whilst  Professor 


PRELIMINARY  CONSIDERATIONS.  77 

Drude  has  shown  (p.  288)  that  all  plant  life  could 
not  possibly  have  been  destroyed  in  northern 
countries.  He  maintains  that  the  greater  part  of 
the  Arctic  floral  elements  which  unite  Greenland 
and  Scandinavia  must  have  survived  the  Glacial 
period  in  these  countries  in  sheltered  localities. 
Indeed,  he  justly  remarks,  where  at  the  present 
moment  do  we  find  such  plantless  wastes  ?  Green- 
land, Franz- Josef  Land,  and  Grinnell  Land,  situated 
in  high  Arctic  latitudes,  all  have  a  flora  composed 
of  flowering  plants  and  cryptograms.  "  I  cannot 
understand,"  he  continues  (p.  286),  "  why  a  flora, 
possibly  mixed  with  northern  forms  but  in  the  main 
points  agreeing  with  our  present  floral  elements, 
should  not  have  persisted  throughout  the  Ice  Age 
even  in  the  heart  of  Germany."  "To  my  mind," 
says  Col.  Feilden,  the  well-known  Arctic  traveller 
(b,  p.  51),  "it  seems  indisputable  that  several  plants 
now  confined  to  the  polar  area  must  have  originated 
there,  and  have  outlived  the  period  of  greatest  ice- 
development  in  that  region."  The  theory  in  favour  of 
a  survival  of  the  pre-glacial  flora  has  been  especially 
strengthened  by  the  late  Mr.  Ball  (than  whom 
probably  no  botanist  possessed  a  better  knowledge 
of  Alpine  plants),  who  was  strongly  in  favour 
of  this  view  as  far  as  the  Alps  are  concerned. 
"Is  it  credible,"  he  says  (p.  576),  "that  in  the 
short  interval  since  the  close  of  the  Glacial  period 
hundreds  of  very  distinct  species  and  several  genera 
have  been  developed  on  the  Alps,  and,  what  is  no  less 


78  HISTORY  OF  THE    EUROPEAN   FAUNA. 

hard  to  conceive,  that  several  of  these  non-Arctic 
species  and  genera  should  still  more  recently  have 
been  distributed  at  wide  intervals  throughout  a  dis- 
continuous mountain  chain  some  1,500  miles  in  length, 
from  the  Pyrenees  to  the  Eastern  Carpathians  ?  "  Mr. 
Ball's  remarks,  indeed,  just  touch  upon  a  very  important 
characteristic  of  all  the  so-called  Alpine  plants.  In 
Europe  they  chiefly  occur  in  Scandinavia  and  the 
central  and  southern  mountain  ranges,  whilst  they 
are  mostly  absent  from  the  intervening  lowlands. 
Again,  we  find  a  large  number  of  species  in  the 
mountains  of  Central  Asia  and  in  some  of  the  North 
American  mountains.  Almost  all  species  of  Alpine 
plants,  in  fact,  are  examples  of  discontinuous  distribu- 
tion; and  this,  as  every  naturalist  knows,  is  always,  in 
both  animals  and  plants,  a  proof  of  antiquity. 

The  glacial  or  Alpine  flora  is  very  old,  and  must 
have  originated  long  before  the  Ice  Age.  But  it  might 
be  urged,  why  should  these  plants  be  now  almost  con- 
fined to  the  Arctic  regions  and  the  higher  mountain 
ranges,  where  the  temperature  undoubtedly  is  very 
low,  if  they  had  originated  during  a  pre-glacial 
period  probably  much  milder  than  the  present? 
The  answer  can  be  given  by  those  who  have  made 
Alpine  plants  their  special  study,  and  who  have 
attempted  to  grow  them  by  administering  to  them 
a  temperature  and  such  climatic  conditions  as  to 
be  most  conducive  to  good  health.  We  should  all 
expect  these  plants  to  be  very  robust,  and  especially 
to  be  able  to  stand  extremely  low  temperatures.  But, 


PRELIMINARY   CONSIDERATIONS.  79 

strange  to  say,  the  very  opposite  is  the  case.  Pro- 
fessor Blytt  tells  us  (p.  19)  that  "Arctic  and  Alpine 
species  in  the  Christiania  Botanic  Gardens  endure  the 
strongest  summer  heat  without  injury,  while  they  are 
often  destroyed  when  not  sufficiently  covered  during 
winter."  The  English  climate  then,  one  would 
think,  ought  to  suit  these  plants,  since  the  winters 
are  not  too  cold;  but  we  find  that  at  Kew  Gardens 
the  large  collection  of  Alpine  plants  have  to  be 
wintered  in  frames  under  glass  in  order  to  keep 
them  in  good  health ;  and  Professor  Dyer,  the 
Director  of  the  Gardens,  thinks  they  are  mostly 
intolerant  of  very  low  temperatures  (compare  also 
pp.  161-164). 

Such  being  the  constitution  of  Alpine  plants,  how 
could  they  possibly  have  originated  during  the  Glacial 
period  and  wandered  from  the  mountains  into  the 
plains,  across  numbers  of  formidable  barriers,  often 
exposed  to  icy  winds,  for  thousands  of  miles?  As  a 
matter  of  fact,  Alpine  plants  have  survived  in  the 
high  North  and  in  the  Alps  because  they  are  there 
permanently  protected  during  winter  by  a  covering  of 
snow  from  very  low  temperatures,  and  they  are  at  the 
same  time  prevented  from  drying  up.  If  they  are 
given  sufficient  moisture  and  a  constant,  mild  tempera- 
ture they  seem  to  do  very  well.  Such  conditions  are 
afforded  them  in  many  parts  of  the  British  Islands,  and 
we  find  indeed  the  Mountain  Avens  (Dryas  octopetala), 
one  of  the  most  typically  Arctic  plants,  growing  wild 
in  profusion  on  the  coast  of  Galway,  in  Ireland,  at  sea- 


8O  HISTORY  OF  THE   EUROPEAN    FAUNA. 

level.  The  winter  temperature  of  that  part  of  Ireland 
resembles  that  of  southern  Europe,  being  no  less  than 
12°  Fahr.  above  freezing  point  This  fact  appears 
to  strengthen  the  view  not  only  that  the  Alpine 
flora  is  of  pre-glacial  origin,  but  that  the  climate  of 
Europe  during  the  Glacial  period  was  mild. 

Having  now  shortly  reviewed  the  state  of  our 
knowledge  with  regard  to  the  former  presence  in 
our  temperate  latitudes  of  Arctic  animals  and  plants, 
it  still  remains  for  me  to  give  a  succinct  statement  of 
the  light  thrown  by  this  fauna  and  flora  on  the  wide- 
spread phenomena  of  glaciation.  It  is  necessary  to 
do  so,  because,  though  the  greater  development  of 
glaciers  on  the  mountains  of  Europe  in  former  times 
does  not  presuppose  the  prevalence  of  an  Arctic 
climate,  the  survival  through  the  Ice  Age  of  a  fauna 
and  flora  could  not  possibly  have  taken  place  in 
northern  Europe  if  the  theories  of  glaciation  now  so 
much  in  vogue  are  really  true.  Professor  Geikie 
reminds  us,  in  speaking  of  his  native  country  (p. 
67),  that  "we  must  believe  that  all  the  hills 
and  valleys  were  once  swathed  in  snow  and 
ice;  that  the  whole  of  Scotland  was  at  some 
distant  date  buried  underneath  one  immense  mer 
de  glace,  through  which  peered  only  the  higher 
mountain  tops."  That  under  such  conditions  no 
fauna  or  flora  to  speak  of  could  have  survived  in 
Scotland  is  evident.  Then  again  he  argues  (p.  426) 
that  because  in  the  great  plain  of  Europe  we  meet 
occasionally  with  striated  rock-surfaces  and  roches 


PRELIMINARY   CONSIDERATIONS.  8 1 

moutonnfos  very  similar  to  those  produced  by  the 
glaciers  of  Switzerland,  it  must  have  been  traversed 
by  "inland  ice"  flowing  from  Scandinavia  and  the 
Baltic  southward.  The  boulder  clay  of  Germany  is 
supposed  to  have  accumulated  underneath  this 
vast  "  mer  de  glace"  as  he  calls  it.  There  is 
no  question  here  of  a  simple  local  development 
of  glaciers,  such  as  could  have  existed  under  a 
mild  and  moist  climate;  practically  all  the  plants 
and  animals  would  have  been  annihilated  in  northern 
Europe  under  such  conditions,  as  there  were  no  areas 
free  from  ice.  A  more  vivid  idea  of  the  state  of 
Europe  during  the  epoch  of  maximum  glaciation 
will  be  obtained  by  looking  at  Professor  Geikie's 
map  (p.  437).  The  whole  of  Scandinavia,  Iceland, 
Scotland,  Ireland,  and  Switzerland  is  there  repre- 
sented as  having  been  completely  enveloped  in  ice, 
and  also  the  greater  part  of  Russia,  Germany,  and 
England.  In  speaking  of  Scandinavia  (p.  424)  he 
remarks  that  "the  whole  country  has  been  moulded 
and  rubbed  and  polished  by  one  immense  sheet 
of  ice,  which  in  its  deeper  portions  could  hardly 
have  been  less  than  5000  feet  or  even  6000 
feet  thick."  The  greater  portion  of  the  area  in- 
dicated as  having  been  underneath  a  sheet  of  ice  is 
thickly  covered  with  superficial  accumulations  of 
gravel,  sand,  and  clay.  The  latter  is  generally 
spoken  of  as  "  boulder  clay,"  and,  with  the  associated 
sand  and  gravel,  it  may  be  observed  equally  well  in 

Russia  or  Germany,  in   England  or  Ireland.      As  a 

6 


82  HISTORY   OF   THE    EUROPEAN    FAUNA. 

rule  these  stony  clays  thicken  out  as  they  are  traced 
from  the  high-lying  tracts  to  the  low  grounds;  and 
especially  near  the  mountains  the  rock-surfaces  are 
often  polished  and  striated.  "  For  many  years  it  was 
believed,"  continues  Professor  Geikie  (p.  432),  "that 
all  those  superficial  deposits  were  of  iceberg  origin. 
The  low  grounds  of  Northern  Europe  were  supposed 
to  have  been  submerged  at  a  time  when  numerous 
icebergs,  detached  from  glaciers  in  Scandinavia  and 
Finland,  sailed  across  the  drowned  countries,  dropping 
rock-rubbish  on  the  way.  Such  was  thought  to  have 
been  the  origin  of  the  erratics,  stony  clay,  and  other 
superficial  accumulations,  and  hence  they  came  to  be 
known  as  the  'great  northern  drift  formation."'  "  But," 
he  adds  (p.  433),  "  when  the  phenomena  came  to 
be  studied  in  greater  detail  and  over  a  wider  area, 
this  explanation  did  not  prove  satisfactory.  The 
facts  described  in  the  preceding  paragraphs — the 
occurrence  of  striated  surfaces  and  roches  mou- 
tonnees,  the  disturbed  appearances  associated  with 
the  till,  and  the  not  infrequent  presence  of  giants' 
kettles — convinced  geologists  that  all  the  vast 
regions  over  which  boulder-clay  is  distributed  were 
formerly  occupied  by  the  'inland  ice'  of  Scan- 
dinavia." 

I  think  Professor  Geikie  over-estimates  the  value  of 
the  evidences  which  appear  to  be  in  favour  of  his 
theory.  His  treatise  on  the  Ice  Age  leaves  one 
under  the  impression  that  the  older  view  of  the 
marine  origin  of  the  boulder-clay  is  not  only 


PRELIMINARY  CONSIDERATIONS.  83 

done  with  for  good  and  all,  but  that  no  geologists 
nowadays  believe  in  it.  If  a  more  careful  study  of 
the  glacial  phenomena  has  led  most  geologists  to 
abandon  what  I  might  call  the  "marine  view"  in 
favour  of  the  terrestrial  one,  a  more  careful  study 
of  the  fauna  and  flora  will,  I  venture  to  think,  have 
the  opposite  effect.  However,  it  appears  that  even 
from  a  purely  geological  point  of  view  more  can 
be  said  in  favour  of  the  old  theory  than  Pro- 
fessor Geikie  and  his  school  are  ready  to  admit. 
Thus  we  are  told  by  Professor  Bonney  (p.  280), 
in  referring  to  the  boulder-clay,  that  "the  singular 
mixture  and  apparent  crossing  of  the  paths  of 
boulders  are  less  difficult  to  explain  on  the  hypo- 
thesis of  distribution  by  floating  ice  than  on  that  of 
transport  by  land-ice,  because,  in  the  former  case, 
though  the  drift  of  winds  and  currents  would  be 
generally  in  one  direction,  both  might  be  varied  at 
particular  seasons.  So  far  as  concerns  the  distribu- 
tion and  thickness  of  the  glacial  deposits,  there  is  not 
much  to  choose  between  either  hypothesis;  but  on 
that  of  land-ice  it  is  extremely  difficult  to  explain 
the  intercalation  of  perfectly  stratified  sands  and 
gravel  and  of  boulder-clay,  as  well  as  the  not  in- 
frequent signs  of  bedding  in  the  latter."  "  Anything," 
writes  Professor  Cole  (p.  239),  "that  keeps  open  the 
position  maintained  by  Lyell  and  others,  that 
extensive  glaciation  is  compatible  with  mild  and 
sheltered  nooks  and  corners,  and  that  much  of 
the  distribution  of  boulder-clay  was  performed  in 


84  HISTORY  OF  THE   EUROPEAN    FAUNA. 

seas  and  not  on  land,  may  be  welcomed  by 
rationalists,  at  any  rate  until  further  research  has 
been  carried  on  among  the  Arctic  glaciers.  At 
present  every  year  brings  evidence  of  modern 
marine  boulder-clays  in  high  latitudes,  and  removes 
us  farther  and  farther  from  belief  in  a  moraine 
profonde"  That  foraminifera  are  occasionally  found 
in  boulder-clay  has  been  known  for  a  long  time, 
but  it  is  only  within  recent  years  that  these  marine 
organisms  have  been  shown  to  occur  in  so  many 
localities,  that  Mr.  Wright,  who  examined  a  large 
number  of  samples,  says  (p.  269),  "  I  am  forced 
to  the  conclusion  that  the  Scottish  as  well  as 
the  Irish  boulder-clay  is  a  true  marine  sedimentary 
deposit " 

In  the  fourth  and  fifth  chapters  I  shall  return  to 
this  subject  again,  and  mention  a  number  of  facts 
of  distribution  which  appear  to  me  much  easier  of 
explanation  by  means  of  the  marine  than  by  the  land- 
ice  theory.  But  I  do  not  propose  to  go  into  further 
geological  details  in  this  volume,  as  I  think  I  have 
clearly  conveyed  my  position  in  this  controversy. 

Before  concluding  this  short  review  of  the  glacial 
problem,  so  far  as  it  affects  the  origin  of  the  European 
fauna,  I  should  like  to  refer  to  the  opinion  of  one 
who  has  devoted  years  to  the  study  of  the  glacial 
phenomena  in  the  Arctic  Regions,  viz.,  Col. 
Feilden.  "To  a  certain  extent,"  he  says  (a,  p.  57), 
"all  boulder  clays  at  home  arc  fragmentary  when 
compared  with  the  boulder-bearing  beds  of  Kolguev, 


PRELIMINARY   CONSIDERATIONS.  85 

which  we  may  safely  assume  are  50  miles  in  length 
by  40  in  width,  with  a  thickness  of  not  less  than  250 
feet,  probably  far  more,  all  lying  in  one  undisturbed 
mass.  It  is  suggestive  that  all  the  glacial  deposits 
which  I  have  met  with  in  Arctic  and  Polar  lands, 
with  the  exception  of  the  terminal  moraines  now 
forming  above  sea-level  in  areas  so  widely  separated 
as  Smith's  Sound,  Grinnell  Land,  North  Greenland, 
Spitsbergen,  Novaya  Zemlya,  and  Arctic  Norway, 
should  be  glacio-marine  beds.  Throughout  this 
broad  expanse  of  the  Arctic  Regions  I  have  come 
across  no  beds  that  could  be  satisfactorily  assigned 
to  the  direct  action  of  land-ice;  that  is  to  say,  beds 
formed  in  situ  by  the  grinding  force  and  pressure  of 
an  ice-sheet.  On  the  contrary,  so  far  as  I  can  judge, 
the  glacial  beds  which  I  have  traced  over  the  exten- 
sive area  mentioned  above  have  all  been  deposited 
subaqueously  and  re-elevated." 

One  of  the  strongest  arguments  that  can  be  used 
against  the  view  of  the  marine  origin  of  the  glacial 
phenomena  in  Northern  Europe  seems  to  me  the  fact 
that  we  find  polished  rock-surfaces  far  removed  from 
the  source  of  glaciers,  and  so  exactly  resembling  those 
produced  at  the  present  day  by  our  Alpine  glaciers  as 
to  appear  identical  to  the  experienced  eye.  Most  of 
such  striated  and  polished  rocks  occurring  in  the 
higher  mountain  ranges  of  Scandinavia,  and  also  of 
the  British  Islands,  have  no  doubt  been  actually  pro- 
duced by  glaciers,  whilst  those  in  the  plain,  some- 
times hundreds  of  miles  away  from  the  mountains, 


86  HISTORY   OF  THE   EUROPEAN   FAUNA. 

must  have  originated  in  a  similar  manner;  that  is  to 
say,  by  a  heavy  mass  of  material  containing  stones 
being  slowly  dragged  over  the  rock-surfaces.  The 
weight  which  causes  the  stones  to  polish  the  latter  is 
generally  ice,  but  it  is  quite  conceivable  that  any  other 
substance,  especially  if  it  is  in  a  semi-solid  state,  must 
act  and  operate  in  much  the  same  way.  All  polished 
rock-surfaces  are  carved  by  glaciers,  because  we  can 
see  them  done  by  glaciers  every  day,  is  the  argument 
commonly  used  nowadays.  It  was  not  so  formerly. 
But  Mr.  Mallet  and  his  views  are  almost  forgotten 
now;  his  name  does  not  even  appear  in  our  great 
modern  works  on  the  Ice  Age.  His  argument  was 
that  as  the  land  rose  out  of  the  glacial  sea,  the  mud 
which  had  accumulated  round  the  shore  slipped 
downward  in  a  direction  determined  by  the  contour 
of  the  surrounding  valleys  and  mountains.  The 
moment  the  land  rose  above  water-level,  the  large 
mass  of  gravel  and  mud  lying  upon  it  slipped  down- 
ward. During  a  steady  rising  of  the  land  there  would 
therefore  be  produced  a  continuous  sliding  down  of 
this  mud-glacier,  which  would  groove  and  polish  the 
rock  underneath  it,  in  the  same  manner  as  the  ice- 
glaciers  do  in  the  Alps  (p.  47).  Professors  Sedgwick 
and  Haughton  became  strong  adherents  of  Mr. 
Mallet's  theory  at  the  time,  but  it  seems  later  on 
to  have  fallen  into  disfavour  with  geologists,  who 
may  not  even  be  thankful  to  have  it  brought  to 
light  again. 


PRELIMINARY   CONSIDERATIONS.  8/ 


SUMMARY  OF  CHAPTER  II. 

I  have  endeavoured  to  show  in  this  chapter  how  we  can  deter- 
mine approximately  the  original  home  of  an  animal.  By  this 
means  we  are  able  to  study  the  component  elements  of  the 
European  fauna,  which  is  found  to  consist  to  a  large  extent  of 
migrants  from  the  neighbouring  continents.  There  is  a  Siberian, 
an  Oriental,  and  an  Arctic  element  in  it.  The  remainder  of  the 
fauna  is  derived  from  local  centres  of  dispersal.  "What  was 
formerly  believed  to  have  been  one  great  northern  migration 
now  resolves  itself,  on  closer  study,  into  two  very  distinct 
ones— the  Siberian  and  the  Arctic.  The  mammals  have 
received  most  attention  hitherto,  because  their  remains  are  so 
frequently  met  with,  thus  enabling  us  more  easily  to  investigate 
their  past  history;  but  butterflies  and  snails  have  not  been 
neglected,  and  at  least  one  very  remarkable  work  on  the  latter 
has  been  published  dealing  with  their  origin  in  Europe  and 
in  the  remainder  of  the  Palsearctic  region. 

The  former  distribution  of  land  and  water  is  intimately  con* 
nected  with  the  origin  of  the  European  fauna,  and  the  changes 
which  have  taken  place  in  this  respect  may  be  best  traced  by 
the  present  distribution  of  mammals,  snails,  and  earthworms. 
In  this  manner  the  British  Islands  may  be  shown  to  have  been 
connected  with  one  another  and  with  the  Continent;  Spain 
with  Morocco  across  the  Straits  of  Gibraltar;  Greece  with  Asia 
Minor,  and  so  forth. 

The  British  fauna  has  played  such  an  important  part  in  the 
evolution  of  the  European  fauna,  that  it  forms  the  key  to  the 
solution  of  the  wider  problem.  In  it  five  elements  are 
recognisable,  of  which  the  Lusitanian  element  is  the  oldest, 
and  the  Siberian  the  most  recent.  It  has  been  deemed 
advisable  to  conclude  this  chapter  with  a  short  review  of  the 


88  HISTORY   OF   THE   EUROPEAN    FAUNA. 

history  of  the  Glacial  period  in  its  climatic  effects  on  the 
animals  and  plants  of  Europe.  A  number  of  writers  are 
quoted  who  have  conducted  special  researches  in  determining 
the  temperature  of  our  continent  at  the  time.  The  fauna  of 
Europe  is  frequently  described  as  having  been  of  an  Arctic 
nature,  but  as  a  matter  of  fact  there  existed  during  the  Ice 
Age  a  striking  and  most  remarkable  mingling  of  a  northern 
and  a  southern  fauna.  The  presence  of  Siberian  mammals  in 
Europe  is  said  to  have  been  due  to  the  prevalence  of  a  dry 
steppe  climate,  but  this  view  is  not  supported  by  other  evidence. 
The  Alpine  flora  in  a  wide  sense  is  probably  pre-glacial  in 
origin,  and  appears  to  have  survived  the  Ice  Age  where  it 
is  now  known  to  exist.  A  few  words  on  the  phenomena  of 
glaciation  are  added  before  bringing  the  chapter  to  a  close. 


CHAPTER   III. 

THE   FAUNA  OF   BRITAIN. 

THE  British  Islands  are,  as  I  have  remarked,  very 
suitable  as  a  starting-point  for  our  investigations. 
Their  fauna  and  flora  are  fairly  well  known,  and  the 
distribution  of  the  large  animals  at  any  rate,  which 
are  of  course  of  much  importance  in  these  researches, 
has  been  as  much  studied  as  that  of  any  other  area 
in  Europe.  We  possess  in  England  an  abundance  of 
the  remains  of  past  animal  life,  and  a  combination  of 
the  data  furnished  by  both  of  these  important  factors 
will  enable  us  to  draw  up  a  history  of  the  origin  of 
the  present  British  fauna. 

In  the  first  chapter  I  indicated  that  in  the  fauna  of 
the  British  Islands  three  divisions  or  elements  are 
recognisable — a  northern,  a  southern,  and  an  eastern. 
These  elements  correspond  to  migrations  which  can 
be  proved  to  have  arrived  in  this  country  at  different 
periods  in  past  times.  When  we  investigate  these 
migrations  more  closely,  the  eastern  is  found  to  be 
composed  partly  of  European  and  partly  of  Siberian 
species.  The  southern  is  made  up  of  European 
and  of  Central  and  Southern  Asiatic  species.  To 
make  matters  still  more  complex,  the  southern  and 

89 


90  HISTORY   OF   THE   EUROPEAN    FAUNA. 

eastern  migrations  insensibly  merge  into  one  another, 
so  that  it  is  often  very  difficult  to  determine  to  which 
of  them  an  animal  may  belong.  The  European 
species  spread  principally  from  three  centres  over 
Europe — viz.,  from  the  Lusitanian,  Alpine,  and  the 
Balkan  centres.  The  southern  element  of  the  British 
fauna  is  therefore  composed  of  animals  which  have 
originated  in  these  three  centres,  and  in  Central  and 
Southern  Asia.  The  Balkan  species  have  been 
included  with  those  coming  from  the  latter  centre 
under  the  term  "Oriental"  migration.  The  sixth 
chapter  is  devoted  to  it,  whilst  the  Lusitanian  and 
Alpine  migrations  have  each  a  chapter  to  them- 
selves. 

The  Arctic  Hare  is,  as  I  have  already  mentioned, 
one  of  the  mammals  of  the  northern  element  of  the 
British  fauna.  It  is  now  confined  to  the  mountains  of 
Scotland  and  the  plain  and  mountains  of  Ireland. 
But  in  former  times  it  had  a  wider  range  in  the 
British  Islands.  The  Stoat  is  another  distinctly 
northern  mammal.  It  occurs  with  us,  as  Messrs. 
Thomas  and  Barrett-Hamilton  have  pointed  out, 
in  two  distinct  varieties  or  species,  the  one  being 
confined  to  Great  Britain,  the  other  to  Ireland.  As 
I  shall  explain  more  fully  later  on  (p.  135),  I  have 
reasons  to  believe  that  the  Irish  Stoat  came  from  the 
Arctic  Regions  as  a  northern  migrant,  but  that  the 
English  Stoat,  on  the  other  hand,  reached  England 
with  the  Siberian  fauna  from  the  east.  A  third 
northern  animal,  now  extinct  in  the  British  Islands, 


THE   FAUNA   OF   BRITAIN.  91 

is  the  Reindeer.  It  is  supposed  to  have  died  out  in 
these  countries  not  very  many  centuries  ago,  and 
records  have  been  handed  down  to  us  that  it  still 
inhabited  Scotland  as  late  as  the  thirteenth  century. 
Like  the  Stoat,  it  occurred  in  two  well-known  varie- 
ties, distinguished  from  one  another  by  the  shape 
and  form  of  the  antlers.  In  the  English  pleistocene 
deposits  the  remains  of  both  kinds  are  met  with 
mingled  together,  whilst  in  Ireland  only  one  of  them 
has  been  found.  The  explanation  of  this  case  is 
similar  to  that  of  the  two  stoats.  One  of  the  varieties, 
which  we  may  call  the  northern  one,  came  to  us  from 
the  Arctic  Regions;  the  second  wandered  to  the 
British  Islands  at  a  later  period,  when  Ireland  had 
probably  become  separated  from  England.  It  was 
therefore  unable  to  penetrate  so  far  west. 

One  of  the  most  familiar  examples  of  a  northern 
British  bird  is  the  Red  Grouse  (Lagopus  scoticus).  By 
most  authorities  it  is  looked  upon  as  a  species  distinct 
from  the  Scandinavian  Willow  Grouse  (Lagopus 
albus\  but  except  in  colour  it  is  undistinguishabie 
from  it,  and  the  eggs  are  identical.  The  whole  genus 
Lagopus  is  a  distinctly  Arctic  one,  and  there  can  be 
no  doubt  that  the  British  Grouse  belongs  to  the 
northern  migration,  just  like  the  Arctic  Hare.  The 
Ptarmigan  (Lagopus  mutus)  and  the  Snow  Bunting 
are  also  migrants  from  the  north.  Though  as  resident 
British  birds  they  are  quite  confined  to  Scotland,  the 
remains  of  the  former  have  been  found  in  a  cave  in 
the  south  of  Ireland,  showing  that  its  range  in  the 


92  HISTORY   OF   THE    EUROPEAN    FAUNA. 

British  Islands  was  formerly  more  extensive.  Another 
bird  which  probably  came  to  our  shores  with  this 
same  migration,  though  it  is  now  unfortunately  ex- 
tinct, is  the  Great  Auk  (Alca  impennis),  of  which 
some  specimens  have  luckily  been  preserved  in  our 
museums.  From  the  occurrence  of  its  remains  in 
kitchen-middens  and  other  recent  deposits,  the  Great 
Auk  is  known  to  have  inhabited  the  coasts  of  Scotland, 
Ireland,  and  Scandinavia,  as  well  as  those  of  New- 
foundland. Mr.  Ussher  recently  found  the  bones  of 
this  bird  near  Waterford,  which,  I  believe,  is  the  most 
southern  locality  known.  The  manner  of  their  occur- 
rence leaves  no  doubt  that  the  bird  had  been  used  as 
food  by  the  early  races  of  man.  In  all  probability  it 
originated  in  the  Arctic  Regions,  and  subsequently 
spread  south  on  either  side  of  the  Atlantic.  We 
need  not  here  refer  to  the  many  winter  visitants, 
— northern  birds  which  appear  regularly,  or  at  more 
or  less  long  intervals,  in  these  islands, — although  in 
most  of  the  ornithological  works  they  are  included 
under  the  term  "British  Birds." 

All  the  British  reptiles  and  amphibia  appear  to 
have  reached  us  from  the  south  or  east,  but  among 
the  fishes  there  are  a  good  many  northern  forms. 
The  whole  salmon  family  —  the  Salmonidcs  —  are 
typical  northern  immigrants.  The  Stickleback  (Gas- 
terosteus  aculeatus\  too,  has  undoubtedly  come  to  us 
from  the  north.  The  genus  Cottus,  like  Gasterosteus> 
is  certainly  Arctic  in  origin.  Originally  freshwater 
forms,  many  species  are  now  found  between  tide- 


THE    FAUNA   OF   BRITAIN.  93 

marks,  and  of  these  a  few  have  migrated  southward 
along  the  coasts  of  the  great  continents.  Thus  we 
meet  with  various  species  of  Coitus  as  far  south  as 
California  and  Japan,  on  the  American  and  Asiatic 
coasts  of  the  Pacific  respectively.  In  Europe,  two 
species,  viz.,  C.  scorpio  and  C.  bubalis,  range  as  far 
south  as  the  French  coast.  Our  freshwater  Cottns, 
the  Miller's  thumb  (Cottus  gobio),  has  migrated  to 
us  from  the  north  with  the  Arctic  species.  All  the 
freshwater  forms,  indeed,  of  this  genus  are  typically 
Arctic. 

A  large  number  of  land  and  freshwater  invertebrates 
too  have  no  doubt  reached  us  from  the  north.  Some 
of  them  may  have  originated  in  Scandinavia  or  within 
the  Arctic  Circle,  but  others  probably  came  still 
farther,  either  from  America  or  even  from  Asia,  and 
used  the  Arctic  land-connection  via  Greenland  in  their 
migration  to  Europe.  As  I  shall  give  a  number  of 
additional  instances  of  such  migrants  in  the  succeed- 
ing chapters,  I  need  not,  perhaps,  dwell  upon  them 
now  any  longer,  except  to  mention  a  few  of  the 
more  typical  ones.  Vertigo  alpestris,  a  minute  snail 
with  an  amber-coloured  shell,  and  our  freshwater 
pearl-mussel,  Unto  (Margaritana)  margaritifery  belong 
to  this  migration.  Then  among  butterflies  we  may 
cite  the  Marsh-ringlet  (Coenonympha  t)phon\  and 
among  beetles,  Pelophila  borealis  and  BletJiisa  multi- 
punctata.  There  are  a  number  of  northern  spiders, 
among  which  a  few  certainly  indicate  an  Arctic 
origin,  or  at  any  rate,  that  they  have  wandered  to 


94  HISTORY   OF   THE   EUROPEAN    FAUNA. 

Europe  across  Greenland  and  the  old  Arctic  land- 
connections.  Bathyphantes  nigrinus^  Linyphia  insignis, 
and  Drapetisca  socialis,  for  instance,  are  three  British 
species  whose  range  indicates  a  northern  origin,  and 
which  also  occur,  according  to  Mr.  Carpenter,  in 
North  America.  Mr.  Carpenter  also  tells  me  that  the 
Collembolan,  Isotoma  littoralis,  is  a  typical  northern 
migrant  He  has  recently  discovered  it  in  the  west 
of  Ireland,  its  only  station  in  the  British  Islands. 

Among  the  Crustacea,  the  genus  Apus  forms  an 
exceedingly  interesting  illustration  of  the  northern 
migration,  Apus  glacialis  having  been  discovered  in 
a  Scottish  pleistocene  freshwater  deposit,  whilst  it  is 
now  almost  confined  to  the  Arctic  regions. 

To  the  same  group  of  animals  also  belong  the  three 
remarkable  species  of  freshwater  sponges,  Ephydatia 
crateriformis,  Heteromeyenia  Ryderi,  and  Tubella  pen- 
sylvanicay  which  Dr.  Hanitsch  has  described  from  some 
lakes  in  Western  Ireland.  None  of  these  are  known 
from  Great  Britain  or  from  the  continent  of  Europe. 
A  few  North  American  plants  grow  wild  in  the  same 
district.  That  any  of  these  should  owe  their  existence 
in  Ireland  to  accidental  introduction  appears  to  me 
exceedingly  improbable.  In  a  former  contribu- 
tion to  this  subject  (a,  p.  475)  I  assumed  that 
these  American  plants  and  animals  had  migrated 
to  Europe  at  the  same  time  as  the  other  northern 
forms  referred  to.  My  friend  Mr.  Carpenter,  how- 
ever, takes  exception  to  this  (p.  383),  and  I  quite 
recognise  the  force  of  his  argument.  "Their  very 


THE   FAUNA   OF   BRITAIN.  95 

restricted  and  discontinuous  ranges,"  he  says,  "  along 
the  extreme  western  margin  of  Europe  mark  them 
as  decidedly  older  than  those  northern  animals  and 
plants  which  have  a  circumpolar  distribution."  We 
have  indeed  quite  similar  examples  in  the  Oriental 
migration,  of  which  part  is  very  ancient,  surviving 
here  and  there  and  exhibiting  discontinuous  distri- 
bution. We  may  therefore  look  upon  these  American 
immigrants  as  among  the  oldest  members  of  that 
northern  stock  which  have  survived  in  our  islands 
— probably  a  mere  remnant  of  a  once  luxuriant  flora 
and  fauna. 

In  order  to  show  the  importance  of  the  Eastern  or 
Siberian  element  in  the  English,  or,  we  might  say 
with  Dr.  Sclater,  the  Anglo-Scotian  mammalian 
fauna,  I  herewith  give  a  list  of  the  species  of 
mammals  which  probably  migrated  to  Great  Britain 
from  Siberia.  I  have  marked  with  an  asterisk  those 
which  still  exist  in  this  country  (not  in  Ireland),  or 
have  become  extinct  within  historic  times. 


Canis  lagopus.  *  Mus  minutus. 

Gulo  luscus.  *  Arvicola  agrestis. 

*  Mustela  erminea.  *        „       amphibius. 

*  „        putorius.  „       arvalis. 

*  „        vulgaris.  *         „       glareolus. 

*  Sorex  vulgaris.  ,,       gregalis. 
Lagomys  pusillus.                                    ,,       ratticeps. 

*  Castor  fiber.  Equus  caballus. 
Spermophilus  Eversmanni.  Saiga  tartarica. 

„  erythrogenoides.         Ovibos  moschatus. 


96  HISTORY   OF   THE  EUROPEAN    FAUNA. 

Cricetus  songarus.  Alces  latifrons. 

Myodes  lemmus.  „      machlis. 

Cunictilus  torquatus.  Rangifer  tarandus. 


We  have  evidence  that  most  of  these  twenty-six 
species  of  mammals  came  from  Eastern  Europe,  but 
there  is  no  reason  to  suppose  that  they  originated  there. 
On  the  contrary,  it  is  highly  probable,  as  I  said  be- 
fore, that  their  native  home  is  Siberia,  and  that  they 
entered  Europe  to  the  north  of  the  Caspian.  Along 
with  these,  vast  numbers  of  other  forms  of  life,  and 
also  plants,  swarmed  into  our  continent,  and  as  we 
advance  eastward  from  England  we  meet  with  them 
in  increasing  numbers  to  the  present  day.  But  not 
only  on  the  Continent  do  we  find  these  survivals  of 
the  great  Siberian  migration,  which  has  been  so  ably 
described  by  Professor  Nehring;  no  less  than  nine 
species  still  inhabit  Great  Britain  (if  we  include  the 
recently  extinct  Beaver).  On  the  other  hand,  not 
more  than  three  have  been  found  fossil  in  Ireland, 
and  of  these  only  one  still  survives.  This  very  signifi- 
cant fact  will  be  referred  to  again  more  fully  on  p.  153. 
Meanwhile  it  should  be  remembered  that  these 
three  species,  viz.,  Mustela  erminea,  Equus  caballus, 
and  Rangifer  tarandus,  occur  in  Ireland  in  varieties 
distinct  from  those  found  in  Central  Europe.  It  is 
upon  this,  and  many  other  circumstances,  that 
I  founded  my  belief  that  Ireland  was  already 
separated  from  England  at  the  time  of  the  arrival 
of  the  Siberian  emigrants  in  the  latter  country.  As 


THE   FAUNA  OF   BRITAIN.  97 

we  shall  see,  the  Irish  Stoat,  Horse,  and  Reindeer 
probably  came  by  a  different  route  from  that  taken 
by  the  English  representatives  of  the  same  species. 

Very  few  of  the  lower  animals  of  Siberian  origin 
have  reached  the  British  Islands.  Most  of  those 
which  were  formerly  thought  to  be  Siberian  are  either 
of  East  European  or  of  Central  and  South  Asiatic 
origin,  though  they  probably  joined  the  Siberian 
migration  on  their  way  to  England.  The  Arctic 
migration  brought  a  greater  variety  of  species  to 
this  country  than  the  Siberian,  but  neither  the  one 
nor  the  other  has  contributed  more  than  a  small  per- 
centage to  the  British  fauna.  The  bulk  of  that  fauna 
is  derived  from  the  various  European  centres  of  dis- 
persal, and  especially  from  Central  and  Southern  Asia. 

Those  animals  which  have  their  home  in  the  latter 
area,  I  have  named  Orientals,  though  it  must  be  re- 
membered that  they  need  not  necessarily  have  come 
from  what  is  known  among  zoologists  as  the  "  Oriental 
Region."  The  terms  "  Oriental  animals  "  and  "  Ori- 
ental migration  "  are  used  here  in  a  wider  sense,  and 
include  even  those  species  which  reached  Central  and 
Northern  Europe  from  South-Eastern  Europe.  It  is 
astonishing,  what  a  vast  number  of  both  vertebrate 
and  invertebrate  animals  can  be  traced  back  to  this 
Oriental  migration.  Great  tracts  of  Europe  were 
repeatedly  submerged  beneath  the  sea  during  Tertiary 
times,  and  on  their  re-appearance  were  formed  into 
green  fields  and  pastures  new  for  the  rich  Asiatic 
fauna,  which  was  ever  ready  to  flood  the  neighbouring 

7 


98  HISTORY  OF   THE   EUROPEAN   FAUNA, 

continent.     This  went  on,  and  not  for  a  comparatively 
short  space  of  time,  as  in  the  case  of  the  Siberian 
invasion ;    the     immeasurable    ages    which    passed, 
whilst  several  of  the  Tertiary  epochs  dawned  upon 
Europe,   witnessed    an    almost   constant    stream    of 
Asiatic    immigrants    pouring   in  upon    us.      Europe 
returned  her  own    products   in    exchange,   but  they 
must  have  been  scanty  in  comparison  to  the  enor- 
mous   number  of    species  which  have    undoubtedly 
originated    in    Central    and    Southern    Asia.     Very 
many  of  the  widely  distributed  forms  in  the  British 
Islands  are  of  Oriental   origin.      Among  these   are 
also   the    cosmopolitan    species,   such   as    the    Barn 
Owl  (Strix  flammed]  and  the  Painted  Lady  Butter- 
fly   ( Vanessa    cardui}.      A    great    number    of    our 
British  Mammals,  Birds,  Butterflies,  and  Beetles  have 
come   to   us  with   the    Oriental   migration.     But,  as 
I  shall  explain  in  the  special  chapter  devoted  to  it, 
the  earlier  migrants  from  the  south-east  found  their 
northward   progress   barred    by   a   great   sea    which 
stretched  through  Central  Europe  from  west  to  east. 
The  Mediterranean  was  then  divided  into  two  smaller 
basins.     On  their  arrival  in  Greece,  which  was  then 
connected    with    Asia    Minor    and    Southern    Italy, 
the  Oriental  migrants  seem  to  have  turned  westward, 
skirting  the  shores  of  the  Mediterranean.     When  they 
finally  reached  Spain,  many  then  changed  their  course 
northward  (see  Fig.   5,  p.  117)  and  wandered  to  the 
British   Islands   with   the   Lusitanian   animals   which 
came  from  South- Western  Europe. 


THE   FAUNA  OF   BRITAIN.  99 

Dr.  Wallace  makes  mention  of  a  fairly  large  num- 
ber of  species  and  varieties  of  Lepidoptera,  Coleoptera, 
and  land  and  freshwater  Mollusca,  supposed  to  be 
peculiar  to  the  British  Islands.  Even  if  these  were  all 
found  to  be  of  British  origin,  most  of  their  nearest 
relatives  are  continental  species.  Many,  however,  must 
be  looked  upon  as  mere  races  or  sub-species  of 
familiar  continental  forms.  But  others,  such  as  Geo- 
•jualacus  maculosus  and  Asiminea  Grayana,  are  by  no 
means  confined  to  the  British  Islands.  Some  of 
the  so-called  varieties  enumerated  by  Dr.  Wallace 
are  merely  slight  individual  variations  in  form  and 
colour,  which,  only  by  the  extraordinary  tendency  of 
the  variety-monger  to  advertise  himself,  have  received 
a  distinct  Latin  denomination.  The  number  of  the 
remaining  species,  after  weeding  out  the  unworthy 
ones,  will  be  found  to  be  insignificant. 

Similarly,  the  list  of  seventy -five  species  and 
varieties  of  flowering  plants  included  by  Dr.  Wallace 
among  the  forms  peculiar  to  the  British  Islands 
(p.  360)  is  reduced  by  Sir  Joseph  Hooker  to  twenty. 
The  remainder  are  to  be  considered  as  varietal  forms 
of  a  very  trifling  departure  from  the  type,  or  as 
hybrids. 

Just  as  we  distinguish  in  the  British  Islands  the 
parts  inhabited  by  Englishmen,  Scotchmen,  and  Irish- 
men, so  we  can  recognise  three  divisions  in  the  animal 
world,  and  these  roughly  correspond  to  the  boundaries 
of  England,  Scotland,  and  Ireland.  Most  of  the 
eastern  species  inhabit  England,  most  of  the  northern 


IOO         HISTORY  OF  THE   EUROPEAN    FAUNA. 

ones  are  confined  to  Scotland,  whilst  Ireland  is  occu- 
pied chiefly  by  southern  animals.  This,  however,  is 
only  a  very  rough-and-ready  method  of  sub-dividing 
the  British  Islands  into  their  component  parts 
according  to  the  origin  of  their  faunas.  On  closer 
study  such  a  division  is  found  to  be  unsatisfactory. 
The  eastern  species  do  not  really  stop  at  the  Scottish 
frontier,  they  range  far  into  Scotland.  Nor  are  the 
northern  forms  confined  to  the  latter  country.  Many 
of  them  range  into  Ireland,  and  also  into  England.  I 
have  constructed  a  map  of  the  British  Islands  showing 
approximately  the  boundaries  of  the  northern,  eastern, 
and  southern  species  (p.  7),  but  even  this  may  not 
altogether  meet  with  the  views  of  an  ornithologist 
or  conchologist.  For  every  group  of  animals  the 
boundaries  would  probably  require  to  be  marked 
differently.  There  is  also  a  good  deal  of  overlapping, 
so  that  the  attempt  to  define  the  limits  of  the  various 
elements  meets  with  great  difficulties.  But  the  map 
represents,  I  think,  fairly  well  the  general  impression 
one  receives  as  to  the  disposition  of  its  component 
elements,  after  a  careful  study  of  the  British  fauna 
as  a  whole. 

The  distribution  of  the  British  plants  has  been 
worked  out  much  more  thoroughly  than  that  of  the 
animals.  It  need  not  surprise  us,  therefore,  that  the 
first  attempt  to  separate  the  British  Islands  into 
natural  divisions  was  made  by  a  botanist — the  late 
Mr.  Watson.  As  he  himself  pointed  out,  in  making 
these  divisions  he  did  not  take  into  consideration  the 


THE  FAUNA  OF   BRITAIN.  IOI 

origin  of  the  British  species.  They  represent  merely 
groups  of  assemblages  of  plants  of  different  types 
of  vegetation.  Edward  Forbes,  on  the  other  hand, 
founded  his  districts  on  the  origin  of  plants.  His 
work  is  not  only  the  first  of  the  kind,  but  it  is  a 
classical  essay,  and  remains  one  of  the  most  remark- 
able contributions  to  the  literature  on  the  geographical 
distribution  of  living  organisms  known  to  science. 
The  vegetation  of  the  British  Islands,  he  informs  us 
(p.  4),  presents  a  union  of  five  well-marked  floras, 
four  of  which  are  restricted  to  definite  provinces, 
whilst  the  fifth,  besides  exclusively  claiming  a  great 
part  of  the  area,  overspreads  and  commingles  with 
all  the  others.  These  are — 


I.  Mountainous  districts  of  South-)  T 

west  and  West  of  Ireland    .     .  [Lusitaman  type. 

II.  South-west     of    England,     and),-  ,,. 

South-east  of  Ireland      .     ,    .) Galilean  type. 

III.  South-east  of  England. 

IV.  Mountains  of  Scotland,  Cumber-)  c       j- 

land,  and  Wales     .     .          .     .j Scandinavian  type. 

V.  General  Flora Germanic  type. 

Professor  Forbes  points  out,  in  connection  with  the 
plants  of  the  Germanic  type,  that  the  fauna  accom- 
panying this  flora  presents  the  same  peculiarities  and 
diminishes  westward  and  to  the  north.  This  type 
includes,  therefore,  almost  all  the  species  which  can 
be  shown  to  have  come  to  us  directly  from  the  east, 
few  if  any  of  which  have  penetrated  to  Ireland. 

On   a   previous    occasion,   the    same    author    had 


102          HISTORY   OF   THE   EUROPEAN    FAUNA. 

divided  the  British  Islands  into  ten  districts,  accord- 
ing to  the  distribution  of  their  molluscan  fauna. 
These  are — 

I.  The  Channel  Isles. 
II.  South-east  of  England  (including  Cambridgeshire). 

III.  South-west  of  England. 

IV.  North-east  of  England. 

V.  North-west  of  England  (including  Isle  of  Man). 
VI.  North  of  Ireland. 
VII.  South  of  Ireland. 
VIII.  South  of  Scotland. 
IX.  North  of  Scotland. 
X.  Shetland  Isles. 

In  a  short  paper  on  this  subject  (b,  p.  5),  I  have 
shown  that  some  of  these  districts  are  founded  on 
erroneous  data,  whilst,  with  the  knowledge  now  at 
our  disposal,  others  can  no  longer  be  maintained  as 
distinct.  I  thought  then  that  the  molluscan  fauna 
warranted  a  division  of  the  British  Islands  into  the 
following  two  provinces  : — 

I.  England  and  Wales  (except  the  South-west). 
II.  South-west  of  England  and  Wales  and  the  whole  of 
Ireland  and  Scotland. 

The  second  district  contains  some  species  of  mol- 
luscs which  are  almost  entirely  absent  from  the  first, 
such  as  Geomalacus  maculosus,  Testacella  Maugei^ 
Helix  piscina,  Helix  revelata,  Helix  acuta,  and  Pupa 
ringens.  These  are  all  of  Lusitanian  origin,  and  do 
not  occur  in  Central  Europe.  Scotland  alone  cannot 


THE   FAUNA  OF   BRITAIN.  10$ 

be  classed  as  a  separate  province,  since  it  does  not 
contain  a  single  species  peculiar  to  itself.  But,  along 
with  Ireland  and  the  South-west  of  England  and 
Wales,  it  is  distinguished  from  the  remainder  of 
these  countries  by  the  almost  total  absence  of  what 
have  been  called  Germanic  types. 

A  French  conchologist,  the  late  Dr.  Fischer,  dealt 
with  the  British  molluscan  fauna  in  a  somewhat 
similar  spirit  (p.  57).  He  divided  the  British  area 
into  two  districts,  but  these  differ  from  mine  in  so 
far  as  the  South-west  of  England  and  Wales  and  the 
West  of  Ireland  form  one  ;  the  remainder  of  England 
and  Ireland  as  well  as  the  whole  of  Scotland  the 
other.  His  classification  is  of  particular  interest, 
since  the  first  district  represents  part  of  a  larger 
Atlantic  province,  the  second  a  portion  of  the  Ger- 
manic province  of  the  European  sub-region.  The 
latter  he  looks  upon  as  one  of  the  sub-regions  of 
the  great  Palsearctic  Region.  Attention  is  thus 
drawn  to  the  intimate  relationship  existing  between 
the  western  parts  of  the  British  Islands  and  the 
Spanish  peninsula  on  the  one  hand,  and  between  the 
eastern  portions  and  Central  Europe  on  the  other. 

Mr.  Jordan's  North-Sea-and-Baltic  district  includes 
Scotland  and  the  North  of  Ireland,  whilst  England 
joined  with  the  West  and  South  of  Ireland  forms  part 
of  his  Celtic  province.  Both  of  these  districts  or  pro- 
vinces belong  to  Mr.  Jordan's  greater  Germanic 
Region  (p.  302). 

In  the  collection  illustrating  the  geographical  dis- 


104         HISTORY   OF   THE    EUROPEAN    FAUNA. 

tribution  of  animals  in  the  Dublin  Museum,  the 
British  species  have  been  grouped  into  three  divisions. 
One  contains  those  with  a  wide  range  over  the  British 
Islands,  another  the  characteristic  forms  of  the  south- 
east and  lowland  districts  of  Great  Britain,  and  the 
third  the  Irish  and  the  western  and  highland  Anglo- 
Scotian  species.  Mr.  Carpenter  has  named  the  last 
two  divisions  the  "Teutonic"  and  the  "Celtic"  More 
recently,  he  has  recognised  that  this  last  division 
contains  two  distinct  groups  ;  one  including  animals 
of  northern,  the  other  those  of  southern  origin. 
He  acknowledges  indeed,  just  as  I  do,  three  distinct 
faunas  in  the  British  Islands,  with  the  addition  of 
the  group  of  generally  distributed  species  of  un- 
determined origin. 

Many  other  naturalists  have  worked  in  the  direc- 
tion I  have  indicated — namely,  in  grouping  the 
British  animals  into  several  distinct  assemblages, 
without,  however,  taking  their  foreign  range  into 
consideration,  or  their  origin.  I  have  already  referred 
to  the  useful  work  done  by  botanists,  who  have  been 
the  pioneers  in  the  science  of  the  geographical  dis- 
tribution of  living  organisms.  Among  the  British 
naturalists  who  have  applied  the  principles  of  Watson 
to  zoology,  A.  G.  More  deserves  to  be  specially  men- 
tioned. He  was  the  first  to  make  a  serious  study  of 
the  British  fauna  on  the  lines  laid  down  by  that  dis- 
tinguished botanist.  In  conjunction  with  E.  Boyd, 
he  published  a  valuable  essay  on  the  "  Distribution  of 
Butterflies  in  Great  Britain,"  and  later  on  the  birds 


THE   FAUNA  OF   BRITAIN.  IO5 

were  similarly  dealt  with.  All  the  more  important 
groups  of  animals  are  now  being  studied  with  a  view 
to  determining  their  exact  range  in  these  islands. 
Mr.  Harvie-Brown,  Mr.  J.  W.  Taylor,  Mr.  Eagle  Clarke, 
Mr.  Miller  Christy,  Mr.  Ussher,  Mr.  Harrington,  and 
a  number  of  others  have  considerably  advanced  our 
knowledge  in  this  direction  in  recent  years. 

Any  such  contributions  are  to  be  welcomed  as 
furnishing  us  with  the  necessary  data  to  solve  the 
problem  of  the  origin  of  the  British  fauna.  Mean- 
while we  know  enough  to  enable  us  to  assert 
positively  that  the  latter  has  reached  us  by  land- 
connections  from  various  parts  of  Europe  (cf.  p.  35). 
This  statement  of  course  refers  to  the  bulk  jgf  the 
Britishfauna^__The  small  proportion  of  indigenous 
species,  or  such  as  have  been  introduced  accidentally, 
may  be  left  out  of  consideration  when  dealing  with 
the  great  mass  of  animals  which  have  evidently 
migrated  to  the  British  Islands  on  land  now  sunk 
beneath  the  sea  (see  Fig.  4,  p.  60).  Opinions  of 
zoologists,  botanists,  and  geologists  are  practically 
unanimous  on  this  subject ;  yet  there  are  two  other 
theories,  which  have  from  time  to  time  been  advanced 
to  arrniini-fnrJJTej^i^itL^if  thgJRrifrfch  fauna.  Only 

the  first  of  these,  however,  can  claim  the  serious 
attention  of  those  interested  in  the  problem.  Its 
chief  contention  lies  in  the  oft-asserted  dictum  of 
the  "imperfection  of  geological  record"  It  has  been 
suggested,  in  fact,  that  the  British  fauna,  instead 
of  having  migrated  to  our  islands,  might  have 


IO6         HISTORY  OF   THE   EUROPEAN    FAUNA. 

orjgimited  there,  but  that,  owing  to  the  fragmentary 
nature  of  our  Tertiary  deposits,  all  trace  of  their 
early  history  had  disappeared.  "The  origin  of 
European  species,"  remarks  Professor  Cole  (p.  238), 
"within  the  area  of  the  British  Isles,  and  their 
migration  outwards  when  local  conditions  became 
less  favourable  for  their  multiplication,  are  pos- 
sibilities that  seem  too  often  disregarded.  Yet 
the  geologist  must  see  in  the  western  borderland 
of  modern  Europe  a  diminished  continent  from 
which  land-animals  must  have  again  and  again 
moved  eastward."  "  Hence  geologists  may  fairly  be 
unwilling  to  look  on  our  isles  as  barren  lands  waiting 
to  be  peopled  in  pliocene  or  later  times.  Far  rather 
has  the  breaking  up  of  a  broad  land-area  along  the 
present  continental  edge  sent  our  land-fauna  to  the 
new  steppes  that  opened  eastward,  leaving  us  a 
mere  diminished  remnant  to  struggle  with  the  glacial 
period." 

There  are  in  Professor  Cole's  views  many  points 
with  which  I  readily  agree.  In  the  first  place,  he 
acknowledges  that  migration  has  taken  place  on 
land,  so  that  we  have  our  land-connection  between 
Great  Britain  and  the  Continent  whatever  theory  we 
accept  as  to  the  direction  taken  by  the  migrants. 
That  the  western  borderland  of  Europe  has  given 
rise  to  many  important  assemblages  of  animals  in 
past  times,  seems  to  me  also  exceedingly  probable, 
nor  do  I  look  upon  the  British  Islands  as  "  barren 
lands  waiting  to  be  peopled  in  pliocene  or  later 


THE   FAUNA   OF   BRITAIN.  IO/ 

times."  On  the  contrary,  I  believe  an  almost  un- 
interrupted stream  of  migrants  poured  into  the 
British  Isles  before  pliocene  times  from  the  south. 
But  what  I  thoroughly  disagree  with,  is  the  remark 
that  our  British  land-fauna  has  been  sent  to  the  new 
steppes  that  opened  eastward.  These  are  the  more 
or  less  arid  portions  of  Eastern  Europe.  Professor 
Cole  no  doubt  has  in  mind  those  species  of  mammals 
which  I  have  included  in  what  I  called  the  Siberian 
migration,  and  of  which  we  have  fossil  evidence 
in  the  late  Tertiary  deposits  of  Europe.  It  would 
be  impossible  here  to  discuss  this  subject  fully, 
especially  as  I  have  done  so  in  the  subsequent 
chapters;  but,  even  if  we  had  no  geological  record 
whatsoever,  the  present  range  of  the  species  in 
question  and  their  nearest  relatives  must  convince 
us  that  they  could  not  have  originated  in  Western 
Europe.  However,  on  the  strength  of  the  geological 
evidence,  Professor  Nehring — the  only  one  who  has 
made  this  fauna  his  special  study — remarks  (p.  228), 
that  there  seems  scarcely  any  doubt  that  this  steppe- 
fauna  just  referred  to  had  come  to  us  from  the  east. 
Professors  Boyd  Dawkins,  Brandt,  and  Lartet  held 
similar  views. 

The  theory  that  an  ice-sheet  stretched  across  a 
narrow  sea  might  be  the  means  of  aiding  a  fauna 
across  from  the  mainland  to  an  island,  is  particularly 
inapplicable  to  the  British  Islands.  Neither  Mr. 
Kinahan  nor  Mr.  Lamplugh,  the  two  supporters  of 
this  view,  have,  however,  taken  the  trouble  to  apply 


108         HISTORY   OF   THE   EUROPEAN   FAUNA. 

it  to  more  than  one  species  of  the  British  fauna.  An 
ice-bridge,  thinks  Mr.  Kinahan,  "  could  easily  have 
connected  Scotland  and  Ireland,  thus  giving  a  land 
causeway  for  migration"  (p.  3).  Mr.  Lamplugh 
throws  more  light  on  this  interesting  speculation  by 
giving  us  the  name  of  an  animal  which  he  believes 
crossed  a  narrow  sea  on  a  bridge  of  ice.  This  animal 
unfortunately  happens  to  be  one  whose  remains  have 
never  been  found  in  high  northern  latitudes,  viz.,  the 
Irish  elk  (Cervus  giganteus).  And  because  he  is  of 
opinion  that  this  species  of  extinct  deer  found  its 
way  to  the  Isle  of  Man  from  the  mainland  on  a 
waning  ice-sheet,  he  sees  no  reason  why  certain 
elements  of  the  Irish  fauna  should  not  have  been 
similarly  introduced. 

It  seems  of  no  advantage  to  begin  the  discussion 
on  the  origin  of  the  British  fauna  by  assuming  the 
former  existence  of  ice-bridges,  and  the  possibility  of 
a  migration  across  them  of  some  of  its  members.  If 
a  glacier  connected  Scotland  and  Ireland,  the  climate 
of  both  countries  (since  they  were  highlands  and 
acted  as  the  feeders  of  the  ice-sheet)  must  have 
been  uncomfortable  to  the  majority  of  the  British 
species.  What  were  the  inducements  that  could 
have  prompted  those  which  had  braved  the  dis- 
comforts of  Scotland  to  emigrate  to  Ireland  at 
such  a  time?  What  light  does  it  throw  on  the 
origin  of  the  Irish  fauna  as  a  whole,  to  advance  the 
extremely  improbable  hypothesis  that  certain  ele- 
ments of  it  may  have  reached  Ireland  by  an  ice* 


THE   FAUNA  OF   BRITAIN.  IOQ 

bridge?  If  any  species  came  to  that  country  in  such 
an  unusual  manner,  surely  they  must  have  been  Arctic 
or  northern  forms.  But  what  about  the  southern 
species,  which  form  the  bulk  of  the  Irish  fauna  and 
also  the  flora?  Even  the  Arctic  element  of  the 
British  fauna,  which  probably  includes,  besides  the 
Reindeer,  many  hundreds  of  species,  could  not,  I  think, 
have  migrated  to  these  islands  on  an  ice-bridge.  In- 
deed, I  agree  with  most  of  the  writers  who  have  dealt 
with  the  subject,  in  asserting  that  the  northern  as  well 
as  all  the  other  elements  of  our  fauna  utilised  for  their 
migration  the  old  land-bridges  which  connected  these 
islands  with  one  another  and  with  the  Continent. 

There  is  a  greater  diversity  of  opinion  as  to  the  age 
during  which  the  British  fauna  arrived  in  these  islands. 
This  is  naturally  a  much  more  complicated  problem, 
but  it  is  one  which  I  am  convinced  will  ultimately  be 
solved  mainly  by  means  of  a  study  of  the  geographical 
distribution  of  animals  and  plants.  I  ft  we  can  settle 
the  relative  ages  of  the  various  migrations,  wejhereby 
supply  afTTTTTpoTtant  fink  in  our  attempt  to  reconstruct 
the  past  geographicallealures  of  the  BritTsrTTslands. 
The  range  ofthe~7Brttish  species  will  give  us  ^rr 
idea  of  the  nature  of  the  land-connections  and  their 
gradual  changes  in  course  of  time.  Geological  data 
are  exceedingly  valuable  in  these  inquiries,  but  it  is 
a  fatal  mistake  to  build  our  geographical  theories 
and  the  origin  of  the  British  fauna  as  a  whole 
entirely  on  the  assumptions  of  a  certain  school  of 
geologists.  Unfortunately,  Dr.  White's  very  interest- 


1 10         HISTORY  OF  THE   EUROPEAN    FAUNA. 

ing  remarks  on  the  British  fauna  for  this  reason  lose 
much  of  the  value  which  they  might  otherwise 
possess. 

In  his  remarkable  essay  the  late  Edward  Forbes 
affirms  that  the  flora  peculiar  to  the  west  of 
Ireland,  of  which  the  strawberry  tree  (Arbutus  unedo) 
is  the  most  striking  example,  and  which  exhibits  such 
strong  southern  affinities,  is  not  only  much  the  most 
ancient  of  our  island  floras,  but  that  it  is  actually  of 
miocene  age.  It  migrated  to  Ireland  from  Spain  at 
a  very  remote  period,  during  which  he  supposed  that 
a  direct  land-connection  existed  between  the  two 
countries.  The  destruction  of  this  old  land-bridge, 
he  thinks,  must  have  taken  place  before  the  com- 
mencement of  the  Glacial  period.  Climatal  changes 
during  that  time  destroyed  the  mass  of  the  southern 
flora  which  had  thus  reached  Ireland,  the  survivors 
being  species  such  as  were  most  hardy  (saxifrages, 
heaths,  etc.),  which  he  considers  to  be  the  only  relics 
of  this  most  ancient  portion  of  our  flora. 

The  northern  or  Arctic  fauna  and  flora,  according 
to  the  same  author,  established  themselves  in  the 
British  Isles  during  the  Glacial  period — at  a  time 
when  these  were  groups  of  islands  in  the  midst  of 
an  ice-bound  sea.  Finally,  the  great  mass  of  our 
animals  and  plants  migrated  from  the  Continent  to 
England  after  the  Glacial  period.  "  The  migration  of 
the  species,"  he  says,  "  less  speedy  of  diffusion,  which 
are  now  peculiar  to  England  was  arrested  by  the 
breaking  up  of  the  land-connection  between  England 


THE  FAUNA  OF  BRITAIN.  Ill 

and  Ireland,  and  thence  the  famous  deficiencies  of 
the  sister  isle,  as,  for  instance,  its  freedom  from 
reptiles"  (p.  10).  He  is  also  of  opinion,  that  the 
separation  between  England  and  the  Continent  took 
plade  at  a  later  date  than  that  between  England 
and  Ireland. 

According  to  Dr.  A.  R.Wallace  (p.  338),  we  possessed 
just  before  and  during  the  Glacial  period  "a  fauna 
almost  or  quite  identical  with  that  of  adjacent  parts 
of  the  Continent,  and  equally  rich  in  species."  But 
the  submersion,  he  thinks,  which  is  supposed  to  have 
occurred  during  the  latter  part  of  the  Glacial  period, 
destroyed  the  greater  part  of  the  life  of  our  country. 
When  England  again  became  continental,  continues 
Dr.  Wallace,  this  fauna  was  succeeded  by  an  assem- 
blage of  animals  from  Central  Europe.  "  But  sufficient 
time  does  not  seem  to  have  elapsed  for  the  migra- 
tion to  have  been  completed  before  subsidence  again 
occurred,  cutting  off  the  further  influx  of  purely 
terrestrial  animals,  and  leaving  us  without  the  number 
of  species  which  our  favourable  climate  and  varied 
surface  entitle  us  to."  The  comparative  zoological 
poverty  of  Ireland  he  attributes  to  the  fact  that  "  the 
depth  of  the  Irish  Sea  being  somewhat  greater  than 
that  of  the  German  Ocean,  the  connecting  land 
would  there  probably  be  of  small  extent  and  of  less 
duration,  thus  offering  an  additional  barrier  to 
migration." 

Dr.  Wallace's  explanation  of  the  origin  of  the 
British  fauna  is  disappointing  after  Forbes's  careful 


112         HISTORY  OF   THE   EUROPEAN   FAUNA. 

study  and  critical  inquiry  into  its  component  ele- 
ments. So  great  an  authority  on  geographical 
distribution  might  have  given  us  more  lucid  state- 
ments of  his  views  on  a  variety  of  topics  connected 
with  this  subject. 

In  speaking  of  the  fauna  of  Ireland,  Professor 
Leith  Adams,  Professor  Dawkins,  and  Mr.  Alston  are 
evidently  only  thinking  of  the  mammals,  which  form 
but  a  very  small  proportion  of  it.  The  first-men- 
tioned palaeontologist  held  that  there  was  a  land- 
communication  between  Scotland  and  Ireland  at  the 
close  of  the  Glacial  period,  by  which  the  greater 
portion  of  the  mammals  that  had  found  their  way  to 
the  former  country  crossed  to  the  latter  (p.  100). 
And,  he  continues,  the  severance  between  the  two 
countries  must  have  taken  place  before  the  slow- 
travelling  Mole,  the  Beaver,  the  forest-haunting  Elk 
and  the  Roebuck  had  time  to  arrive. 

Much  in  the  same  spirit  are  Mr.  Alston's  remarks 
on  this  subject  (p.  5).  "  The  absence  from  the  known 
fossil  fauna  of  Scotland  and  Ireland  of  most  of  the 
characteristic  post-pliocene  English  animals,  shows 
that  the  northward  migration  of  these  forms  was  slow, 
gradually  advancing  as  the  glacial  conditions  of 
the  northern  parts  of  our  islands  decreased  in  in- 
tensity. Thus  it  is  not  difficult  to  suppose  that  the 
Hedgehog,  Ermine,  Badger,  Squirrel,  and  Mountain 
Hare  may  have  found  their  way  through  southern 
Scotland  into  Ireland  long  before  they  were  able  to 
penetrate  into  the  still  sub-arctic  regions  of  the  High- 


XJI^ 
^  n«- 

THE   FAUNA  OF   BRITAIN. 

lands.  Subsequently,  when  the  improvement  of  the 
climate  had  continued,  the  Shrews  and  Voles  may 
well  have  found  their  way  northward  along  the  com- 
paratively  genial  coasts,  before  the  larger  beasts  of 
prey  could  find  a  sufficient  stock  of  game." 

That  the  Bear,  Wolf,  Stag,  Horse,  Mammoth,  and 
Reindeer  lived  in  Ireland  before  the  Glacial  period 
is  considered  highly  probable  by  Professor  Boyd 
Dawkins  (a,  p.  152). 

Only  the  Butterflies  are  dealt  with  in  Dr.  Buchanan 
White's  clever  little  essay  on  distribution.  And, 
as  I  remarked  before,  his  conclusions  are  some- 
what marred  by  the  unwarrantable  assumption  that 
our  islands  at  no  distant  date  were  totally  destitute 
of  all  plant-life,  and  were  therefore  uninhabitable  by 
animals.  But  his  paper  differs  in  so  far  from  most  of 
the  others,  that  he  has  made  a  thorough  study  of  the 
one  group  he  deals  with.  In  some  respects  it  may  serve 
as  a  model  to  future  students  in  its  general  treatment 
of  the  problem  he  has  set  himself  to  work  out  He 
adopts  the  principle,  even  for  butterflies,  that  though 
it  is  possible  for  them  to  be  blown  over  from  the 
Continent,  they  have  probably  migrated  with  the  rest 
of  our  indigenous  fauna  and  flora  across  the  dry  bed 
of  the  German  Ocean.  His  conclusions  are  that 
Britain  derived  its  butterfly  fauna  from  continental 
Europe  in  post-glacial  times,  that  the  Arctic  and 
Alpine  species  were  the  first  arrivals,  and  that  one 
part  of  the  Irish  species  reached  Ireland  by  way  of 
Scotland,  another  from  the  south.  He  assumes,  of 


I  14         HISTORY  OF  THE   EUROPEAN   FAUNA. 

course,  that  Great  Britain  and  Ireland  were  connected 
at  that  time. 

Within  the  last  few  years  the  spell  which  has  bound 
naturalists  to  accept  the  theory  of  a  total  destruction 
of  life  during  the  Glacial  period  is  happily  vanishing, 
and  more  enlightened  views  are  gaining  ground. 
The  Lusitanian  species  of  plants  in  the  west  of  Ire- 
land, which  had  already  furnished  Forbes  with  an 
argument  in  favour  of  survival,  are  also  regarded  by 
Mr.  Bulman  as  the  remnants  of  a  pre-glacial  flora 
which  was  exterminated  everywhere  else  by  the 
cold  (p.  265).  This  view  of  the  survival  of  a  pre- 
glacial  fauna  and  flora  has  since  been  accepted  by 
Mr.  Carpenter,  whilst  I  also  have  endeavoured  to 
bring  fresh  evidence  into  the  field  in  its  favour. 
We  both  agree  with  Edward  Forbes  in  considering 
the  Lusitanian  element  as  the  oldest  section  of  our 
fauna  and  flora,  and  that  it  came  long  before  the 
Glacial  period.  But  we  differ  somewhat  from  him, 
in  so  far  as  we  do  not  limit  that  element  to  Ireland. 
It  seems  also  to  be  represented  in  South-western 
England  and  Wales,  though  it  is  there  less  con- 
spicuous. 

This  decision  as  to  the  relative  age  of  the  British 
South-western  fauna  has  not  been  arrived  at  from  any 
geological  considerations.  The  conviction  that  it 
must  be  older  than  the  other  sections  has  been  gained 
solely  from  a  study  of  the  geographical  distribution 
of  the  species  belonging  to  that  fauna.  Many  of 
them  exhibit  what  is  known  as  "discontinuous  distri- 


THE   FAUNA  OF   BRITAIN.  115 

bution,"  which  zoologists  are  agreed  to  regard  as  a 
sign  of  antiquity.  Thus  Geomalacus  maculosus,  the 
Kerry  Slug,  is  in  the  British  Islands  confined  to  South- 
western Ireland  (see  Fig.  19,  p.  300),  and  on  the  Con- 
tinent it  is  unknown  north  of  North-western  Spain. 
The  Millepede,  Polydesmus  gallicus,  has  a  wider  range 
in  Ireland,  and  is  also  known  from  France  and  the 
Azores.  Two  Earthworms  of  the  Spanish  and 
Mediterranean  region,  viz.,  Allolobophora  veneta  and 
Georgii^  have  been  discovered  in  Ireland,  but  are 
apparently  unknown  in  England  or  France ;  whilst 
the  Weevil,  OtiorrJiyncJius  auropunctatus,  does  not 
occur  north  of  the  Auvergne  Mountains  in  France 
except  in  Ireland.  A  very  large  number  of  instances 
might  be  mentioned  of  species  found  in  South- 
western Europe,  France,  the  South-west  of  England 
and  Ireland.  Enough,  however,  has  been  said  to 
show  the  nature  of  the  fauna,  and  there  is,  as 
Forbes  has  pointed  out,  a  corresponding  flora. 

A  great  number  of  the  species  belonging  to  the 
South-western  British  element  seem  to  have  origin- 
ated in  South-western  Europe,  or  at  any  rate  to  have 
spread  over  our  continent  from  that  part.  Their 
home  lay  therefore  probably  in  a  warm,  damp 
climate,  and  it  seems  a  reasonable  inference  to 
suppose  that  they  spread  north  at  a  time  when 
the  temperature  over  the  British  Islands  was  much 
higher  than  what  it  is  now.  Any  one  familiar  with 
our  Bristle  fern,  or  Killarney  fern,  as  it  is  called  in 
Ireland  (Trichomanes  radicans},  will  readily  admit  that 


Il6         HISTORY  OF   THE   EUROPEAN   FAUNA. 

it  must  have  come  to  us  at  such  an  epoch.  It  at  once 
suggests  some  shady  waterfall  in  a  tropical  forest,  and 
indeed  the  home  of  the  genus  is  South  America.  It 
is  one  of  those  plants  which  have  evidently  migrated 
to  us  from  South-western  Europe,  'a  mere  remnant 
of  a  once  luxuriant  flora. 

Sir  Archibald  Geikie  tells  us  (p.  837),  and  in  the 
main  every  one  agrees  with  him,  that  at  the  beginning 
of  the  Tertiary  era  in  which  we  now  live,  the  climate 
was  of  a  tropical  and  subtropical  character  in  Europe. 
Gradually  it  became  more  temperate,  and  eventually 
it  passed  into  a  phase  of  extreme  cold,  but  since  that 
time  the  cold  has  again  gradually  diminished.  It  is 
quite  evident,  therefore,  that  from  a  purely  geological 
point  of  view  our  south-western  flora  must  have 
migrated  northward  before  the  cold  came  on,  and 
survived  in  sheltered  localities  under  the  influence 
of  the  mild  coast  climate.  Some,  however,  suppose 
that  there  occurred  a  phase  of  extreme  mildness  im- 
mediately after  the  Glacial  period,  and  that  it  was 
during  that  time  that  the  Lusitanian  fauna  and  flora 
became  established  in  the  British  Islands.  To  this 
Professor  James  Geikie  replies  (£,  p.  169),  "there  are  few 
points  we  can  be  more  sure  of  than  this,  that  since 
the  close  of  the  Glacial  epoch — since  the  deposition  of 
the  clays  with  Arctic  shells  and  the  Saxicava  sands 
• — there  have  been  no  great  oscillations,  but  only  a 
gradual  amelioration  of  climate.  It  is  quite  impossible 
to  believe  that  any  warm  period  could  have  intervened 
between  the  last  Arctic  and  the  present  temperate 


THE  FAUNA  OF   BRITAIN.  117 

conditions  without  leaving  some  notable  evidence  in 


the  superficial  deposits  of  Scotland,  Scandinavia,  and 
North  America."     Thus  it  appears  that  on  the  whole 


Il8         HISTORY  OF   THE   EUROPEAN    FAUNA. 

the  assumption  that  the  Lusitanian  fauna  and  flora 
are  very  ancient  and  pre-glacial  is  also  supported 
on  geological  evidence. 

The  course  of  events  in  the  origin  of  the  British 
fauna  might  have  been  therefore  somewhat  as 
follows : — In  early  Tertiary  times,  when  the  climate 
all  over  Western  Europe  was  moist  and  semi-tropical, 
a  migration  proceeded  northward  from  the  south- 
western corner  of  Europe.  This  was  strengthened 
by  Oriental  migrants  which  had  moved  westward 
along  the  Mediterranean  basin  (Fig.  5,  No.  i). 
Owing  to  geographical  changes  supervening,  the 
Alpine  fauna  (No.  2)  was  then  enabled  to  colonise 
the  British  Islands,  and  subsequently  another  migra- 
tion had  begun  to  come  in  from  the  south-east 
(No.  3).  The  climate  had  meanwhile  gradually 
become  more  temperate  and  drier,  About  the  same 
time,  or  even  earlier,  an  Arctic  migration  commenced 
to  pass  southward  (No.  4),  and  finally  the  Siberian 
animals  (No.  5)  poured  into  our  continent.  The 
arrows  in  the  map  indicate  the  directions  followed  by 
the  different  migrants  as  they  travelled  to  the  British 
Islands.  The  arrows  are  not  meant  to  represent  the 
whole  nor  the  full  extent  of  the  migrations  from 
any  particular  centre,  but  only  in  so  far  as  they 
affect  our  islands.  Moreover,  it  would  be  im- 
possible to  indicate  on  one  map  the  geographical 
conditions  which  obtained  during  the  several  migra- 
tions. It  must  be  remembered  that  during  the  time 
which  elapsed  while  they  passed  into  the  British 


THE  FAUNA  OF   BRITAIN.  1 19 

Islands,  these  were  joined  in  the  north  to  Scandinavia 
and  in  the  south  to  Belgium  and  France.  The  various 
phases  of  geographical  evolution  of  Europe  will  be 
studied  in  the  subsequent  chapters,  and  maps  will 
then  be  given  to  show  as  far  as  possible  in  a 
general  way  the  leading  characteristics  of  these 
great  changes. 

I  have  now  given  some  reasons  for  the  belief  that 
several  different  migrations  of  animals  entered  the 
British  Islands  in  later  Tertiary  times.  I  have  also 
shown  why  some  of  them  must  be  looked  upon  as 
being  older  than  others,  and  in  so  far  we  have  come 
to  a  decision  as  to  their  relative  ages.  It  still  remains 
for  us,  however,  to  examine  how  their  geological  ages 
can  be  approximately  determined.  We  require  for 
this  purpose  palasontological  aid. 

In  the  fifth  chapter  will  be  found  the  history  of 
the  Siberian  migration.  And  since  we  possess  most 
valuable  records  of  it  in  the  numerous  fossil  remains 
discovered  in  Central  and  Western  Europe,  we  are 
able  to  trace  their  progress  from  the  east  to  the 
west  in  a  very  complete  and  satisfactory  manner.  In 
England  their  first  appearance  dates  from  the  Forest- 
Bed,  for  here  we  find  remains  of  the  Glutton  (Gulo 
luscus\  Musk-Ox  (Ovibos  moschatus\  and  others  (see 
p.  204).  It  seems  reasonable  to  suppose,  therefore, 
that  the  first  entry  of  these  Siberian  mammals  into 
Europe  took  place  at  or  just  before  the  Forest-Bed 
period.  But  Professor  Nehring  tells  us  in  his  remark- 
able work  on  the  Tundra  and  Steppes  (p.  222),  that  in 


120         HISTORY   OF   THE   EUROPEAN    FAUNA. 

Germany  the  remains  of  the  same  mammals  occur  in 
deposits  which  are  certainly  more  recent  than  the 
lower  continental  boulder  clay;  and  he  is  inclined  to 
the  belief  that  they  migrated  into  Europe  during  the 
inter-glacial  phase  which  is  supposed  to  have  separated 
the  earlier  from  the  later  stage  of  the  Glacial  period. 
It  is  evident  that  in  this  case  the  inter-glacial  period 
in  Germany  would  have  corresponded  to,  and  be  con- 
temporaneous with,  our  Forest-Bed  period.  The 
deposits  immediately  preceding  the  Forest-Bed  would 
also  be  contemporaneous  with  the  lower  continental 
boulder  clay.  Although  this  may  seem  rather  a 
startling  statement  to  make,  from  the  evidence 
which  will  be  brought  forward  in  the  fourth  and 
fifth  chapters  I  am  inclined  to  the  belief  that  such 
is  probably  the  case. 

Having  once  arrived  at  a  determination  of  the 
exact  geological  period  during  which  the  Siberian 
mammals  invaded  our  continent,  and  having  also 
previously  determined  the  relative  ages  of  the  various 
other  migrations,  we  have  advanced  another  step  in 
the  direction  we  are  aiming  at  Let  us  suppose  that 
the  Siberian  migration  actually  reached  the  British 
Islands  during  the  Forest-Bed  period.  Since  the 
Siberian  migration  is  the  most  recent  of  those  which 
entered  the  British  Islands,  the  others  must  have 
commenced  their  march  before  the  Forest-Bed 
period.  Now  it  was  Professor  Boyd  Dawkins  who 
first  indicated  to  us,  as  I  have  remarked  before, 
the  method  of  research  to  be  adopted  in  an  attempt 


THE  FAUNA  OF   BRITAIN.  121 

to  determine  the  geological  age  of  the  different 
migrations  in  so  far  as  they  affected  the  British 
Islands.  I  may  be  excused,  therefore,  for  again 
quoting  the  following  important  passage  in  one  of 
his  works.  "  The  absence,"  he  says  (£,  p.  xxix),  "  of 
the  beaver  and  the  dormouse  from  Ireland  must  be 
due  to  the  existence  of  some  barrier  to  their  westward 
migration  from  the  adjacent  mainland,  and  the  fact 
that  the  Alpine  hare  is  indigenous,  while  the  common 
hare  is  absent,  implies  that,  so  far  as  relates  to  the 
former  animal,  the  barrier  did  not  exist."  The  Beaver, 
Dormouse,  and  Common  Hare  are  either  Siberians  or 
later  migrants  from  elsewhere,  and  there  can  be  no 
doubt  that  at  the  Forest-Bed  period  Ireland  was 
already,  or  was  just  being,  separated  from  England. 
All  the  southern  species,  that  is  to  say  all  the 
Lusitanian,  Alpine,  and  Oriental  forms  occurring  in 
Ireland,  must  therefore  be  older  than  that  period.  I 
have  advocated  similar  views  in  a  former  essay  on 
this  subject.  Mr.  Carpenter  recently  advanced  some 
interesting  and  valuable  criticisms  on  these  views, 
which  we  may  examine  a  little  more  closely  (p.  385). 
"  While,  then,"  he  remarks,  "  I  find  myself  in  almost 
complete  agreement  with  Dr.  Scharff  with  regard  to 
the  older  sections  of  our  fauna,  I  think  that  those 
widespread  species  which  survived  the  Glacial  period 
must  have  been  confined  to  the  more  southern  parts 
of  our  area,  and  have  only  subsequently  spread 
northwards  and  westwards  to  Scotland  and  Ireland." 
He  suggests,  in  fact,  that  the  widespread  British 


122         HISTORY   OF   THE   EUROPEAN   FAUNA. 

species  belong  to  a  younger  or  newer  section  of  our 
fauna  than  the  local  ones.  In  many  cases  this  may 
be  quite  true,  but  we  possess  also  a  large  number  of 
common  and  widely-spread  forms  which  bear  the 
impress  of  antiquity  upon  them.  We  have  the  most 
positive  proof  of  the  antiquity  of  the  very  common 
small  circular  Snail  (Helix:  rotundata},  since  it  was 
found  in  miocene  freshwater  deposits  near  Bor- 
deaux. Many  other  examples  might  be  mentioned 
to  show  that,  though  discontinuous  range  is  generally 
a  proof  of  antiquity,  continuous  range  is  not  always  a 
sign  of  the  opposite.  Some  species,  in  fact,  appear  to 
be  short-lived  and  disinclined  to  spread,  whilst  others 
multiply  rapidly  even  under  a  change  of  temperature 
and  climate,  and  are  to  be  found  almost  everywhere. 
But  even  if  we  supposed,  with  Mr.  Carpenter,  that 
these  widely-ranging  species  must  have  been  confined 
during  the  Glacial  period  to  the  more  southern  parts 
of  England,  the  idea  that  they  afterwards  made  their 
way  northwards  along  the  eastern  shore  of  the  Irish 
Sea  and  then  passed  into  Ireland,  does  not  appeal  to 
me.  Southern  England  was  occupied  at  that  very 
same  time  by  an  assemblage  of  Siberian  mammals. 
Mr.  Carpenter  thinks  these  might  have  been  kept 
out  of  Ireland  by  an  arm  of  the  sea  until  the  land- 
connection  with  North-western  England  had  broken 
down.  But  if  an  arm  of  the  sea  could  keep  out  the 
Siberian  mammals  it  would  also  keep  out  the  widely- 
spread  British  species  of  the  general  fauna.  On  the 
other  hand,  I  quite  admit  that  my  view  of  the  survival 


THE   FAUNA  OF   BRITAIN.  123 

in  Ireland  of  the  pre-glacial  fauna  is  somewhat 
difficult  to  accept,  considering  that  we  have  such 
undoubted  evidence  of  a  very  extensive  submergence. 
The  case  of  Isle  of  Man,  quoted  by  Mr.  Carpenter, 
can  be  met,  I  think,  by  the  supposition  that  it  was 
connected  with  Cumberland  until  quite  recently,  and 
quite  independently  of  any  connection  between  Eng- 
land and  Ireland;  that  the  Isle  of  Man,  in  fact, 
was  always  a  cape  or  peninsula  of  the  mainland,  and 
only  recently  became  separated  by  local  subsidences 
or  by  the  action  of  the  sea. 

Part  of  the  history  of  the  British  fauna  will  be 
referred  to  again  in  the  next  chapter,  which  deals 
with  the  Arctic  migration.  We  need  not  therefore 
dwell  any  longer  on  this  subject  here.  There  is  one 
matter,  however,  which  is  of  importance  in  connection 
with  the  geographical  conditions  of  the  British  Islands 
at  the  time  when  the  greater  portion  of  our  fauna 
arrived  from  abroad. 

On  page  60  will  be  found  a  map  indicating  the 
physical  geography  of  that  part  of  the  ancient  con- 
tinent on  which  what  are  now  the  British  Islands  were 
situated.  Only  one  large  river  has  been  marked  on 
that  map,  namely,  that  flowing  out  of  a  lake  which 
occupied  part  of  the  Irish  Sea.  Another  probably  dis- 
charged its  waters  into  the  Atlantic  midway  between 
France  and  England,  whilst  the  Thames  may  have 
been  a  tributary  of  the  Rhine,  as  it  emptied  itself 
into  the  sea  near  our  south-east  coast.  I  have  shown 
in  a  previous  essay  that  the  former  presence  of  a  fresh- 


124         HISTORY   OF   THE   EUROPEAN    FAUNA. 

water  lake  between  England  and  Ireland  is  indicated 
by  the  distribution  of  the  Charrs  and  also  by  the 
various  species  of  British  Coregonus.  There  are 
three  British  species  of  Coregonus,  viz.,  C.  clupeoides, 
C.  vandesius,  and  C.  pollan.  These  are  confined  to 
the  lakes  of  North  Wales,  North-western  England, 
South-western  Scotland,  and  Ireland.  All  but  the 
latter  communicate  at  present  directly  with  the  Irish 
Sea.  The  lakes  of  the  latter  country,  however,  must 
have  done  so  at  a  time  when  the  west  of  Ireland 
stood  at  a  higher  level  than  it  does  now.  The 
ancestors  of  the  three  Coregonus  species,  and  also 
those  of  the  Charrs,  then  lived  in  the  large  freshwater 
lake  indicated  on  the  map  (p.  60),  and  when  the  sea 
gradually  crept  up  the  river  valley  and  finally  con- 
verted the  lake  into  a  gulf,  the  freshwater  fish  took 
refuge^  in  the  rivers  which  supplied  it  with  water. 
^NQW  as  for  the  continuous  sea-shore  between  the 
coast  of  Brittany  and  the  south-west  of  Ireland, 
zoological  distribution  again  aids  us  in  proving  that 
such  must  have  actually  existed  at  no  very  distant 
geological  date.  Most  of  our  common  shore  forms 
of  life  migrate  along  the  coast  exactly  as  land 
animals  do — step  by  step.  Their  eggs  are  care- 
fully attached  to  fixed  objects,  so  as  not  to  be 
carried  away  by  the  waves,  whilst  the  young  often 
remain  and  grow  old  in  some  particular  little  pool, 
rarely  venturing  farther  than  a  few  yards  from  the  spot 
where  they  first  saw  the  light  of  day.  A  number  of 
such  shore  forms  are  found  on  the  west  coast  of  France, 


THE   FAUNA   OF   BRITAIN.  125 

the  same  species  recurring  again  on  the  south-west 
coasts  of  England  and  Ireland,  thus  clearly  indicating 
a  former  continuity  of  coast-line  between  these  points, 
now  separated  by  deep  sea.  A  very  familiar  example 
to  British  zoologists  is  the  purple  rock-boring  Sea- 
urchin  {Strongylocentrotus  lividus),  but  there  are  a 
great  many  others,  such  as  the  semi-marine  Beetles 
Octhebiiis  Lejolisii  and  dEpophilus  Bonnairei^  the 
Crustaceans  Achceus  Cranchii,  Inachus  leptochirus, 
Gonoplax  angulata,  T/iia  assidua,  Callianassa  sub- 
terraneay  the  Fishes  Blennius  galerita  and  Lepado- 
gaster  Decandollii^  and  the  Molluscs  Otina  otis, 
Donax  politus,  and  Amphidesma  castaneum. 

Before  concluding  this  chapter,  a  few  words  as  to 
my  views  on  the  conditions  prevailing  during  the 
Glacial  period  will  not  be  out  of  place.  They  do  not 
differ  very  much  from  those  held  formerly  by  most 
geologists;  and  even  at  present  there  are,  as  I  have 
mentioned  before,  a  few  upholders  of  those  older 
views. 

The  sea,  I  think,  must  have  gradually  crept  across 
England  from  the  east  during,  or  shortly  after,  the 
Forest-Bed  period,  so  as  to  separate  the  south  from 
the  north,  whilst  Ireland  and  Scotland  were  then  still 
connected  with  one  another.  At  a  later  stage,  the  sea 
also  partially  invaded  Ireland,  and  this  condition  is 
very  roughly  represented  on  the  accompanying  map. 
Mr.  Kendall  kindly  drew  my  attention  to  the  fact 
that  several  notable  areas  on  which  shelly  drift  has 
been  observed  are  here  placed  upon  the  land;  but 


126         HISTORY  OF  THE  EUROPEAN   FAUNA. 

it  must  be  remembered  that  one  stage  only  can  be 
shown  on  the  map,  and  that  the  sea  covered  more 
ground  a  little  later.  Many  of  the  smaller  islands  in 
the  glacial  sea,  too,  are  not  shown.  The  map,  in  fact, 


FlG.  6. — Map  of  the  British  Islands,  showing  approximately  in  what 
manner  the  sea  may  have  invaded  the  country  from  the  east  during, 
or  shortly  after,  the  Forest-Bed  period.  The  darkly  shaded 
parts  indicate  the  areas  covered  by  water,  and  the  lightly  shaded 
and  white  portions  what  was  land  at  that  time. 


THE   FAUNA  OF   BRITAIN.  \2J 

is  merely  meant  to  give  a  general  idea  of  the  manner 
in  which  the  great  northern  sea  moved  westward  and 
slowly  covered  a  large  portion  of  the  British  Islands. 
These  peculiar  geographical  conditions  explain,  I 
think,  better  than  anything,  the  absence  from  parts 
of  the  Midlands  and  the  north  of  England  of 
such  a  number  of  terrestrial  invertebrates  which 
are  otherwise  widely  distributed  over  the  British 
Islands.  In  spite  of  the  fact  that  a  large  portion 
of  the  British  Islands  became  submerged,  we 
possessed  at  that  time  an  extensive  area  which 
has  since  been  claimed  by  the  sea,  so  that  there 
was  ample  room  for  the  present  fauna  to  survive 
the  Glacial  period.  The  climate  during  this  period 
was  probably  much  the  same  as  it  is  at  present, 
though  moister,  with  cooler  summers  and  milder 
winters. 

It  may  be  asked  what  proof  we  have  of  such  an 
extensive  submergence  of  England  and  Ireland.  My 
own  views  are  principally  based  on  the  general 
distribution  of  the  fauna  in  the  British  Islands,  and 
the  belief  that  nothing  but  a  mild  climate  during 
the  Glacial  period  could  have  brought  it  about.  On 
purely  geological  grounds,  however,  some  geologists, 
notably  Mr.  Mellard  Reade,  have  come  to  a  similar 
conclusion.  "  The  whole  of  Lancashire  and  Cheshire," 
he  remarks  (a,  p.  542),  "  from  sea-level  up  to  about 
400  feet,  and  in  places  600  feet,  is  covered  by  a 
continuous  mantle  of  boulder-clay  and  sands." 
"  These  clays,  as  a  rule,  contain  distributed  through 


128         HISTORY  OF  THE   EUROPEAN   FAUNA. 

them,  in  a  greater  or  less  degree,  fragments  of  shells 
and  some  perfect  ones.  I  myself  have  recorded 
forty-four  species."  Again  he  continues  (pp.  545 
and  546) :  "  A  large  part  of  Ayrshire  is  covered  with 
similar  shelly  boulder-clays  from  sea-level  to  1061 
feet  at  Dippal.  These  Ayrshire  high-level  shells 
have,  in  the  majority  of  cases,  been  taken,  not  from 
sand  and  gravel  beds,  but  from  boulder-clay,  and  in 
that  respect  they  are  most  important  and  unique. 
In  Moel  Try  fan  the  shells  are  found  in  sands  and 
gravels  at  982  feet;  on  the  range  of  hills  from  Miaera 
to  Llangollen  from  1000-1200  feet;  also  in  sands  and 
gravels  at  Gloppa,  near  Oswestry,  at  1100-1200  feet; 
and  near  Macclesfield  at  a  level  of  about  1200  feet. 
In  Ireland  marine  shells  can  be  traced  almost  from 
sea-level  to  a  height  of  over  1000  feet." 

"Again,"  continues  the  same  author,  "if  we  look 
broadly  at  the  distribution  of  these  shelly  deposits, 
we  find  that  they  occur  all  round  our  maritime  coasts 
in  Lancashire,  Cheshire,  and  Wales,  in  Cumberland 
and  Westmoreland,  Wigtonshire  and  Ayrshire,  and 
along  the  eastern  coast  of  Ireland.  The  same  is 
to  be  said  of  the  eastern  coasts  of  England  and 
Scotland." 

That  a  very  considerable  change  of  sea-level  has 
taken  place  in  some  parts  of  the  British  Islands  would 
appear  to  a  zoologist  the  most  logical  conclusion  after 
an  examination  of  these  "  high-level  shelly  sands  and 
gravels,"  but  the  shells  contained  in  them  are  now 
generally  supposed  to  have  been  carried  there  frozen 


THE   FAUNA   OF   BRITAIN.  1 29 

in  the  sole  of  a  glacier  or  pushed  up  in  front  of  it 
The  older  view,  however,  which  agrees  so  much  better 
with  the  facts  of  distribution,  fortunately  has  not 
disappeared  among  geologists.  "When  we  call  up," 
says  Mr.  Mellard  Reade  (3,  p.  435),  "before  our 
mental  vision  the  simple  and  well-known  facts  of 
nature  which  suffice  to  explain  the  marine  drifts  on 
the  theory  of  submergence,  it  seems  unnecessary 
to  resort  to  the  ingenious  and  artificial  system  of 
physics  elaborated  to  explain  the  phenomena  of 
land-ice." 

"  When  we  have  more  knowledge  of  the  glaciers 
of  the  Arctic  Regions,  and  facts,  in  place  of 
ingenious  suppositions,  to  base  our  reasoning  upon, 
we  may  possibly  have  to  revise  all  our  glacial  con- 
ceptions. In  the  meantime,  the  submergence  theory 
of  the  origin  of  high-level  shelly  gravels  and  sands 
seems  to  me  by  far  the  simpler  of  the  two  theories, 
and  the  most  consistent  with  the  facts  and  phenomena 
which  the  labours  of  a  succession  of  enthusiastic  geo- 
logists have  made  us  acquainted  with." 

Among  those  geologists,  and  they  form  the  majority, 
who  hold  that  Ireland  was  covered  by  land-ice,  there  is 
a  great  diversity  of  opinion  as  to  its  extent.  Messrs. 
Close,  Kinahan,  J.  Geikie,  and  others  believe  that 
the  ice  covered  practically  everything,  whilst  others 
who  claim  to  have  examined  the  ground  with  equal 
care,  such  as  Professor  Carvill  Lewis,  were  led  to 
believe  that  the  south  of  Ireland,  with  the  exception 
of  a  few  local  glaciers,  was  free  from  ice.  The  glacial 

9 


130          HISTORY   OF   THE    EUROPEAN    FAUNA. 

phenomena  of  the  country  can  therefore  be  inter- 
preted in  different  ways,  even  by  those  who  are 
convinced  that  they  are  due  to  land-ice  and  not  to 
icebergs  or  mud-glaciers. 


SUMMARY  OF  CHAPTER  III. 

The  history  of  the  British  fauna  is  not  only  of  interest  to  us 
from  a  sentimental  point  of  view,  it  is  a  convenient  starting- 
point  in  the  study  of  the  larger  European  problem.  The  fauna, 
broadly  speaking,  is  composed  of  three  foreign  elements,  viz.,  the 
northern,  eastern,  and  southern,  to  which  may  be  added  a  small 
endemic  one.  Examples  are  given  of  the  more  noteworthy  forms 
belonging  to  each  of  these.  This  leads  us  to  the  subject  of  the 
natural  divisions  of  the  British  Islands  according  to  their  animal 
inhabitants.  Zoologists  attempted  at  first  to  subdivide  these 
countries,  on  the  lines  laid  down  by  botanists,  into  a  large 
number  of  provinces.  Forbes  proposed  ten  such  divisions  for 
mollusca,  and  subsequently  five,  which  were  ultimately  reduced 
by  others  to  two  or  three. 

The  opinions  of  biologists  are  almost  unanimous  in  attributing 
the  bulk  of  the  British  fauna  and  flora  to  migrations  by  land 
from  the  Continent,  but  two  other  theories,  viz.,  those  of  Pro- 
fessor Cole  and  Messrs.  Kinahan  and  Lamplugh,  are  also 
referred  to.  The  first  believes  in  a  possible  migration  eastward 
from  Western  Europe,  and  the  latter  support  the  view  of  the 
former  existence  of  ice-bridges  to  assist  the  fauna  in  their 
migrations. 

An  endeavour  is  next  made  to  determine  at  what  geological 
periods  the  various  migrations  entered  the  British  Islands. 
There  is  considerable  difference  of  opinion  on  this  subject. 
Some  believe  that  the  British  fauna  is  altogether  post-glacial ; 
a  few  think  that  it  is  partly  so  and  the  remainder  glacial ; 


THE   FAUNA   OF   BRITAIN.  131 

others  again  hold  that  a  portion  is  pre-glacial  and  the  rest 
glacial  and  post-glacial.  Those  who  have  studied  the  subject 
most  closely  feel  convinced  that  the  south-western  or  Lusitanian 
fauna,  and  also  the  flora,  must  have  arrived  before  the  Glacial 
period  and  survived  the  latter  in  these  Islands.  It  seems 
reasonable  to  suppose,  therefore,  that  the  climate  cannot 
have  been  very  severe  during  the  so-called  Ice-Age.  This 
Lusitanian  fauna  must  be  looked  upon  as  the  oldest  portion  of  the 
British  fauna.  The  Alpine  and  Oriental  migrations  arrived  next. 
After  these  came  the  Arctic,  and  finally  the  Eastern  or  Siberian. 
As  the  fossil  evidence  is  most  complete  with  regard  to  the  last, 
we  are  able  to  determine  with  precision  not  only  the  direc- 
tion whence  this  migration  came,  but  approximately  its  geo- 
logical age.  It  arrived  in  Germany  from  the  east  after  the 
deposition  of  the  lower  boulder-clay.  Since  the  boulder-clay 
is  looked  upon  as  a  glacial  deposit,  the  Siberian  migration 
reached  Central  Europe  after  the  first  portion  of  the  Glacial 
period  had  passed.  In  England  it  makes  its  first  appearance 
in  the  Forest-Bed,  which  would  therefore  correspond  to  the 
"  Loess  "  formation  of  Central  Europe.  All  the  other  migrations 
are  older  than  the  Siberian.  They  must  therefore  have  come 
to  Great  Britain  during  the  earlier  part  of  the  Glacial  period 
or  before  it. 

The  chapter  concludes  with  a  short  statement  on  the  physical 
geography  of  the  British  Islands  during  the  time  when  these 
migrations  entered  them.  That  there  existed  a  continuous 
coast-line  between  France  and  Ireland  is  proved  by  the 
occurrence  of  a  considerable  number  of  identical  shore 
species,  whilst  the  former  existence  of  a  freshwater  lake  on 
the  site  of  the  present  Irish  Sea  is  indicated  by  the  dis- 
tribution of  some  freshwater  fishes. 


CHAPTER  IV. 

THE  ARCTIC   FAUNA. 

THE  lands  lying  within  the  Polar  Circle  are  inhabited 
by  an  assemblage  of  animals  and  plants,  many  of 
which  are  peculiar  to  those  regions.  They  are  mostly 
adapted  to  the  abnormal  conditions  of  life  prevailing 
in  the  high  latitudes  of  our  globe — the  long,  dark 
winters,  and  the  short  summers  of  one  long  day. 
Though  the  numbers  of  species  and  of  individuals  arc 
few,  there  is  a  keen  struggle  for  existence  in  those 
regions.  The  prevailing  colour  of  the  ground  is  white, 
and  since  a  resemblance  in  the  colour  of  an  animal 
to  the  ground  it  lives  on  acts  as  a  protection  to  weak 
ones,  and  also  enables  Carnivores  to  approach  their 
prey  with  greater  facility,  it  is  not  surprising  that  we 
should  find  the  majority  of  polar  animals  coloured 
white.  As  I  remarked,  the  polar  area  contains  a  very 
distinct  set  of  species;  most  of  them,  however,  range 
beyond  the  confines  of  the  Arctic  Circle.  It  is  there- 
fore scarcely  justifiable  to  raise  this  Arctic  area  into  a 
distinct  zoological  region  equivalent  to  the  great  zoo- 
geographic  regions,  which  have  been  established  by 
Sclater  and  Wallace,  though  we  might,  with  Dr. 
Brauer,  look  upon  it  as  a  sub-region. 

132 


THE  ARCTIC  FAUNA.  133 

There  are  six  typical  Polar  Land-mammals,  one  of 
which,  the  Polar  Bear,  is  semi-aquatic.  The  Reindeer 
(Rangifer  tarandus)  occurs  upon  almost  all  the  polar 
lands,  and  it  has  often  been  a  source  of  speculation 
in  what  manner  it  has  reached  such  remote  islands  as 
Spitsbergen  and  Novaya  Zemlya — the  former  of  the 
two  being  so  remote  from  a  continent.  There  is  no 
doubt  that  Reindeer  are  great  wanderers,  owing  to 
the  difficulty  of  rinding  sufficient  food-supply  for  the 
large  herds  in  which  they  are  accustomed  to  travel ; 
and  for  this  reason  they  can  cross,  and  have  been 
known  to  cross,  distances  of  from  ten  to  twenty  miles 
on  ice.  The  Behring  Straits,  when  frozen  over  in 
winter,  is  frequently  traversed  by  them.  But  I  quite 
agree  with  Dr.  Brauer  (p.  260)  that  it  is  impossible  to 
account  for  their  presence  in  Spitsbergen  by  an  im- 
migration from  either  Novaya  Zemlya,  Greenland,  or 
Scandinavia,  under  the  present  geographical  con- 
ditions. The  seas  between  the  former  island  and 
the  other  land-masses  referred  to  are  rarely  entirely 
frozen  over.  Even  if  this  should  occur,  the  dis- 
tances between  Spitsbergen  and  Greenland,  Novaya 
Zemlya,  or  Scandinavia  are  so  great,  that  a  migration 
across  ice  is  quite  excluded  from  the  range  of  pos- 
sibilities, since  Reindeer  could  not  subsist  without 
food  during  the  time  it  would  take  to  travel  from 
one  to  the  other.  The  manner  in  which  it  did  reach 
Spitsbergen  and  Greenland  will  be  discussed  more 
fully  below,  and  I  will  therefore  proceed  to  mention 
the  other  Arctic  mammals. 


134         HISTORY   OF   THE   EUROPEAN    FAUNA. 

One  of  the  most  important  and  most  typical  species 
is  the  Polar  Bear  (Ursus  maritimus\  the  greater  part 
of  whose  life  is  spent  on  the  ice  and  in  the  sea.  The 
fact  that  its  favourite  nourishment  consists  of  seals 
proves  its  excellent  and  keen  faculties  of  sight  and 
hearing,  and  its  facility  in  swimming.  But  it  is  not  a 


FIG.  7.  —The  Musk-Ox  ( Ovibos  moschatus).   (From  Flower  &  Lydekker's 
Mammals ,  p.  358.     London:  Adam  &  Chas.  Black.) 

dainty  feeder,  and  lives  upon  almost  all  animals  which 
come  within  its  reach  ;  birds,  land-mammals,  or  fish 
are  not  despised  in  times  of  scarcity.  Its  fur  through- 
out the  year  is  coloured  white,  though  in  old  bears  it 
assumes  a  more  yellowish  hue. 

Another  large  mammal,  perhaps  less  well  known, 
is  the  Musk-Ox   {Ovibos  moschatus,   Fig.   7),   which 


THE   ARCTIC   FAUNA.  135 

resembles  in  size  the  smaller  varieties  of  Oxen,  but  in 
structure  and  habits  is  closely  allied  to  the  Sheep.  As 
is  implied  by  the  specific  name,  it  exhales  a  musky 
odour ;  this  does  not,  however,  appear  to  be  due  to 
the  secretion  of  a  special  gland,  as  is  the  case  in  other 
animals  with  a  similar  smell.  The  skin  is  covered 
with  long  brown  thickly-matted  hair,  interspersed 
with  white.  It  is  confined  to  the  most  northerly 
parts  of  North  America  and  the  American  Arctic 
islands,  and  to  North  Greenland.  Though  not  now 
living  in  the  Old  World,  it  seems  formerly  to  have 
been  abundant  in  Siberia,  and,  as  we  shall  learn  later 
on,  it  was  one  of  the  species  which  took  part  in  the 
great  Siberian  invasion  of  Europe.  Its  remains  have 
been  found  not  only  in  Germany  and  France,  but  also 
in  the  south  of  England. 

The  Polar  Fox  (Cants  lagopus]  occurs  throughout 
the  Polar  Regions,  and  on  islands  where  even  the 
Reindeer  and  the  Musk-Ox  are  unknown.  Beyond 
the  Polar  Circle,  its  range  extends  into  Northern 
Asia,  to  the  extreme  north  of  North  America,  and  the 
mountains  of  Scandinavia.  Like  its  congeners,  it 
had  in  pleistocene  times  a  more  southerly  extension, 
and  fossil  remains  have  been  met  with  in  various 
parts  of  continental  Europe  and  in  England. 

The  Stoat  (Mttstela  erminea\  which  is  known  and 
much  valued  in  commerce  under  the  name  of  Ermine, 
was  formerly  believed  to  occur  only  in  Arctic  America 
and  the  northern  parts  of  the  Old  World,  but  in  more 
recent  years  it  has  been  discovered  in  a  number  of 


136         HISTORY   OF   THE   EUROPEAN    FAUNA. 

the  northern  islands,  such  as  Saghalien,  in  the  islands 
of  the  Behring  Straits,  the  Aleutian  islands,  and  also 
in  Greenland  and  Spitsbergen.  In  Europe,  it  is  found 
as  far  south  as  the  Arctic  Hare,  or  perhaps  even 
farther,  and  it  flourishes  in  the  Alps  up  to  a  height 
of  9000  feet.  It  offers  a  parallel  to  the  Arctic  Hare 
in  the  fact  that  in  some  countries,  such  as  Ireland,  it 
only  rarely  turns  white  in  winter.  The  Irish  form  of 
the  Stoat  differs  so  much  from  the  English,  that 
Messrs.  Thomas  and  Barrett-Hamilton  are  of  opinion 
that  it  is  specifically  distinct,  as  I  mentioned  in 
speaking  of  the  divisions  of  the  British  fauna 
(p.  90). 

The  Arctic  Hare  (Lepus  variabilis]  is  almost  the 
only  one  of  the  typical  Arctic  mammals  which  still 
inhabits  the  British  Islands,  and  for  that  reason  it  is 
to  most  of  us  more  familiar  than  any  of  the  preceding 
species.  Hares  have  been  described  from  Green- 
land by  the  name  of  Lepus  glacialis,  from  the 
European  Alps  as  Lepus  alpinus,  and  under  other 
names  from  Arctic  North  America;  but  though  slight 
differences  in  the  fur  and  even  in  the  skull  can  be 
pointed  out,  there  is  no  doubt  that  all  these  are  only 
varieties  or  races  of  what,  in  the  British  Islands,  is 
known  as  the  Irish  or  the  Scotch  Mountain  Hare, 
Lepus  variabilis.  In  the  Arctic  Regions  this  Hare 
remains  white  throughout  the  year,  but  in  Scan- 
dinavia and  some  other  parts  its  fur  becomes  brown 
in  the  summer,  and  in  Ireland  it  frequently  remains 
entirely  brown  during  the  whole  year,  and  never,  or 


THE   ARCTIC   FAUNA. 


137 


only  in  very  rare  cases,  becomes  entirely  white  in 
winter.  Besides  Scandinavia,  Scotland,  and  Ireland, 
it  is  found  in  Northern  Russia,  and  also  in  the 
Pyrenees,  the  Alps,  and  the  Caucasus.  In  Asia  it 
occurs  not  only  on  the  mainland  of  Siberia,  but  it  has 


FIG.  8. — Map  of  the  northern  hemisphere,  to  show  the  geographical 
distribution  of  the  Arctic  Hare  (Leptis  variabilis]  indicated  in 
black. 

been  obtained  on  the  Akita  Mountains  in  Japan  and 
on  the  Mioko  San  Mountain,  and  also  on  the  island 
of  Saghalien.  It  had  in  former  times  a  more  exten- 
sive range,  and  its  remains  have  been  discovered  in 


138         HISTORY   OF  THE   EUROPEAN    FAUNA. 

England  and  in  a  number  of  places  on  the  continent 
of  Europe.  The  peculiarity  of  its  range,  which  will 
be  explained  more  fully  directly,  lies  in  the  fact  of 
the  occurrence  of  isolated  colonies  in  the  mountains 
of  Europe,  in  Ireland  and  Scotland,  and  in  the 
mountains  of  Japan  (Fig.  8).  From  a  distributional 
point  of  view,  it  is  one  of  the  most  interesting  species 
of  mammals,  and  its  history  throws  a  flood  of  light 
on  the  geographical  changes  which  have  occurred  in 
former  times. 

One  more  species  must  be  mentioned,  and  that  is 
the  Banded  Lemming  {Cuniculus  torquatus},  which 
occurs  chiefly  in  Arctic  America,  Northern  Siberia, 
and  Greenland.  Though  frequently  mistaken  for  the 
Scandinavian  Lemming,  there  is  a  striking  difference 
in  the  character  of  the  teeth,  which  has  induced 
zoologists  to  put  them  into  distinct  genera.  The 
Arctic  Lemming,  moreover,  is  distinguished  from 
the  Scandinavian  by  the  absence  of  external  ears,  the 
densely  furred  feet,  and  by  the  great  length  of  the  two 
middle  claws  in  the  fore-feet.  There  are  two  species 
of  the  true  Lemming,  namely,  the  one  just  referred  to, 
Myodus  leimnuS)  and  Myodus  obensis.  These  may  be 
looked  upon  as  more  or  less  Arctic  species,  since  they 
occur  within  the  Polar  Circle,  but  they  are  not  so 
exclusively  confined  to  that  region  as  the  Banded 
Lemming  {Cuniculus  torquatus}.  The  remains  of 
both  Cuniculus  torquatus  and  of  Myodus  leinmus  have 
been  found  in  British  pleistocene  deposits. 

Until  recently  no  Lemming  remains  had  been  found 


THE   ARCTIC   FAUNA.  139 

to  the  south  of  France,  but  Mr.  Barrett-Hamilton 
announced  to  us  a  short  time  since  that  Dr.  Gadow 
had  discovered  some  skeletons  with  their  skins  still 
preserved  in  a  cave  in  Northern  Portugal.  These 
were  found  to  belong  to  the  Scandinavian  Lemming 
(M.  lemmus)y  and  the  author  incidentally  expressed  the 
opinion  that  there  was  some  possibility  of  this  species 
still  inhabiting  the  mountains  of  Spain. 

The  Lemming  multiplies  with  great  rapidity  under 
favourable  conditions.  In  speaking  of  his  experiences 
in  Siberia  Dr.  Brehm  says  (p.  79):  "All  the  young  of 
the  first  litter  of  the  various  Lemming  females  thrive, 
and  six  weeks  later  at  the  most  these  also  multiply. 
Meanwhile  the  parents  have  brought  forth  a  second 
and  a  third  litter,  and  these  in  their  turn  bring  forth 
young.  Within  three  months  the  heights  and  low 
grounds  of  the  tundra  teem  with  lemmings,  just  as 
our  fields  do  with  mice  under  similar  circumstances. 
Whichever  way  we  turn  we  see  the  busy  little  crea- 
tures, dozens  at  a  single  glance,  thousands  in  the  course 
of  an  hour.  But  the  countless  and  still  increasing 
numbers  prove  their  own  destruction.  Soon  the  lean 
tundra  ceases  to  afford  employment  enough  for  their 
greedy  teeth.  Famine  threatens,  perhaps  actually  sets 
in.  The  anxious  animals  crowd  together  and  begin 
their  march,  hundreds  join  with  hundreds,  thousands 
with  other  thousands,  the  troops  become  swarms,  the 
swarms  armies.  They  travel  in  a  definite  direction, 
at  first  following  old  tracks,  but  soon  striking  out 
new  ones;  in  unending  files — defying  all  computation 


140         HISTORY  OF  THE   EUROPEAN   FAUNA. 

— they  hasten  onwards;  over  the  cliffs  they  plunge 
into  the  water.  Thousands  fall  victims  to  want  and 
hunger;  the  army  behind  streams  on  over  their 
corpses;  hundreds  of  thousands  are  drowned  in  the 
water  or  are  shattered  at  the  foot  of  the  cliffs;  the 
remainder  speed  on;  other  hundreds  and  thousands 
fall  victims  to  the  voracity  of  Arctic  and  red  foxes, 
wolves  and  gluttons,  rough-legged  buzzards  and 
ravens,  owls  and  skuas  which  have  followed  them; 
the  survivors  pay  no  heed.  Where  these  go,  how  they 
end,  none  can  say;  but  certain  it  is,  that  the  tundra 
behind  them  is  as  if  dead,  that  a  number  of  years 
pass  ere  the  few  who  have  remained  behind  and  have 
managed  to  survive  slowly  multiply  and  visibly  re- 
people  their  native  fields."  This  eloquent  passage 
reminds  us  of  the  manner  in  which  migrations  of 
all  kinds  of  animals  have  taken  place  in  former  times, 
and  are  still  taking  place.  It  is  principally  want 
of  food  which  compels  them  to  search  for  new 
homes. 

On  page  91  I  have  referred  to  some  birds  which 
have  come  to  us  from  the  north.  One  of  these,  the 
Snow  Bunting  (Plectrophenax  nivalis],  is  a  typically 
Arctic  species.  In  summer  it  is  widely  distributed, 
and  is  found  in  Spitsbergen,  Novaya  Zemlya,  Siberia, 
and  the  Arctic  Regions  generally.  In  winter  it 
migrates  down  into  North  America,  into  Japan, 
Northern  China,  Turkestan,  Southern  Russia,  and 
occasionally  even  across  Europe  into  North  Africa. 
Very  characteristic  Arctic  birds  are  the  Eider  Ducks 


THE   ARCTIC   FAUNA. 


141 


belonging  to  the  genus  Somateria.  Three  species 
have  visited  the  British  Islands.  The  common  Eider 
Duck  (S.  mollissima},  which  is  of  such  high  com- 
mercial value,  is  abundant  in  Norway  and  northward, 


FIG.  9.— The  Great  Auk  (Aka  imfcniris). 

throughout  the  Polar  Regions.  The  appearance  of 
the  King  Eider  (S.  spectabilis)  on  our  coasts  is  an 
extremely  rare  occurrence,  and  even  in  Norway  it  is 
only  known  as  a  visitor,  but  on  Novaya  Zemlya  and 
along  the  Arctic  shores  of  Siberia,  in  Greenland  and 


142          HISTORY   OF   THE    EUROPEAN    FAUNA. 

Arctic  North  America,  it  is  known  to  breed.  The 
third  species,  Steller's  Eider  (S.  Stelleri),  seems  to  be 
still  rarer,  and  only  in  the  Aleutian  islands  and  in  the 
north  of  Alaska  can  it  be  said  to  be  at  all  abundant. 
It  is  probable  that  the  famous  Great  Auk  (Alca 
impennis,  Fig.  9)  also  was  a  typical  Arctic  species. 
Its  range  extended  to  both  sides  of  the  Atlantic.  In 
Newfoundland  and  on  the  coast  of  Iceland  it  is  known 
to  have  been  met  with  in  considerable  numbers 
within  historic  times;  and  no  doubt,  like  all  Arctic 
species,  it  extended  farther  southwards  at  a  more 
remote  period. 

The  members  of  the  genus  Lagopus,  including  the 
various  species  of  Grouse,  are  likewise  of  northern 
origin.  The  British  Red  Grouse  (L.  scoticiis),  which 
may  be  looked  upon  as  a  form  of  the  Scandinavian 
Willow  Grouse  (L.  albus)  (compare  p  91),  constitutes 
in  some  respects  a  curious  case  of  parallelism  with 
the  Arctic  Hare,  since  the  latter,  in  its  more  southern 
station,  generally  retains  the  summer  fur  throughout 
the  year.  The  allied  Ptarmigan  (L.  mutus]  inhabits 
Scandinavia,  the  Ural  Mountains,  and  some  of  the 
Asiatic  mountain  ranges.  It  is  also  found  in  the 
European  Alps  and  in  the  Pyrenees.  The  North 
European  range  of  the  Ptarmigan  suggests  that 
we  are  dealing  with  an  ancient  species  which  came 
south  from  the  Arctic  Regions  at  about  the  same 
time  as  the  Arctic  Hare;  but  it  is  more  probable, 
as  I  have  shown  in  a  subsequent  chapter  (p.  334), 
that  this  species  has  entered  Europe  more  recently 


THE   ARCTIC   FAUNA.  143 

with  the  Siberian  migrants  from  Central  Asia,  where 
indeed  the  genus  had  its  original  home.  The  Black 
Cock  (Tetrao  tetrix)  and  the  Capercaillie  (Tetrao 
urogallus)  have  also  come  to  us  from  the  east, 
and  have  even  penetrated  into  Ireland.  They  are 
therefore  some  of  the  few  instances  of  members 
of  the  Siberian  invasion  having  become  temporarily 
established  there. 

Reptiles  and  amphibia  are  altogether  unknown  in 
the  Polar  Regions,  but  a  large  number  of  fish,  chiefly 
marine,  have  taken  their  origin  there.  The  Salmon 
family  are  of  Arctic  origin,  as  also  are  the  Stickle- 
backs and  the  Perches,  many  of  the  Cod  family,  the 
Herrings,  and  several  of  the  Flat  fish. 

It  would  lead  me  too  far  to  refer  to  the  invertebrate 
fauna  of  the  Polar  Regions,  but  a  few  remarks  on  the 
Arctic  plants  may  not  be  out  of  place. 

The  principal  Arctic  genera  are  Satix,  Ranunculus, 
Draba,  Pedicularis,  Potentilla,  Saxifraga,  Carex, 
Juncus,  Luzula,  Eriophorum,  and  others. 

Among  the  most  characteristic  Arctic  plants  may 
be  mentioned  Dry  as  octopetala,  to  which  I  have  already 
referred  as  occurring  in  the  west  of  Ireland;  Saxifraga 
oppositifolia,  another  British  species,  occurs  in  the 
higher  mountains  of  Scotland,  Ireland,  and  Wales; 
Braya  alpina,  Papaver  nudicaule.  Lychnis  apetala, 
Diapensia  lapponica,  and  Lobelia  Dortmanna,  which  is 
found  in  the  lakes  of  Scotland  and  Ireland.  The 
dwarf  birch  (Betula  nand)  also,  which  still  occurs 
in  Scotland  and  the  North  of  England,  and  which 


144         HISTORY   OF   THE   EUROPEAN    FAUNA. 

had  formerly  a  wider  range  in  the  British  Islands, 
should  be  included  among  these;  but  there  are  other 
plants  probably  of  Arctic  origin,  though  not  now 
occurring  in  the  Arctic  Regions,  and  to  these  may 
be  classed  the  so-called  American  species  of  plants 
which  are  found  on  the  northern  and  western  coasts 
of  Ireland,  in  the  Hebrides,  in  Scotland,  and  in 
North  America.  These  are  no  doubt  the  relics  of  an 
Arctic  flora  which  flourished  in  high  latitudes  in  past 
times  when  the  climate  there  was  more  temperate 
A  list  of  these  species  will  be  found  on  page  166. 

As  none  of  them  occur  in  Siberia,  they  must 
either  have  found  their  way  to  North  America  and 
to  Europe  from  the  Arctic  Regions,  or  have  travelled 
from  North  America  across  the  latter  to  Europe. 
In  any  case  a  former  land-connection  between  the 
two  continents  must  have  existed.  This  becomes 
the  more  evident  when  we  examine  the  remarkable 
results  obtained  by  the  late  Professor  Heer,  who  first 
described  the  Tertiary  plant-beds  in  North  Greenland. 
No  less  than  282  species  of  plants  have  been  described 
by  this  eminent  botanist  from  these  deposits.  A  large 
number  of  the  plants  found  were  trees  belonging  to 
the  genus  Sequoia^  Thujopsis^  and  Salisburia,  besides 
beeches,  oaks,  planes,  poplars,  limes,  and  magnolias. 
That  they  grew  on  the  spot  is  proved  by  the  fruits, 
which  have  been  obtained  from  these  beds  in  various 
stages  of  growth. 

From  a  similar  deposit  in  Spitsbergen  a  large 
number  of  fossil  plants  have  also  been  brought  to 


THE   ARCTIC   FAUNA.  145 

light,  many  of  which  are  identical  with  those  found 
in  Greenland ;  and  some  of  the  Greenland  forms 
(such  as  Taxodium  distichum  and  Sequoia  Langsdorfii) 
have  been  found  too  in  Alaska,  showing  that  there 
was  probably  a  continuity  of  land  between  Spits- 
bergen and  North  America  by  way  of  Greenland. 
Two  species  of  Sequoias,  namely,  S.  sempervirens 
and  5.  gigantea,  the  well-known  Californian  giant 
trees,  are  very  closely  allied  to  the  Greenland  forms 
discovered  by  Professor  Heer. 

Heer  assigned  the  Arctic  plant-bearing  beds  to  the 
Miocene  epoch,  but  doubts  have  been  recently  thrown 
upon  this  opinion  by  Mr.  Starkie  Gardner,  who 
brought  forward  arguments  in  support  of  his  theory 
of  their  being  of  the  Eocene  age.  Professor  Heer, 
however,  was  able  to  meet  these  criticisms,  and  he  is 
ably  supported  in  his  views  by  Professor  Engler  and 
other  eminent  continental  botanists. 

It  is  evident  that  under  the  present  conditions  of 
temperature  none  of  those  plants  could  have  flourished 
in  Greenland.  The  climate  must  have  been  much 
milder  than  it  is  at  present.  Professor  Heer  estimated 
from  the  general  aspect  of  the  fossil  flora  that  the 
mean  annual  temperature  of  North  Greenland  was 
at  least  nine  degrees  centigrade,  and  that  the  mean 
winter  temperature  was  not  below  zero. 

It  will  hardly  be  necessary  for  me  to  review  here 
the  various  theories  which  have  been  advanced  by 
geologists  and  botanists  to  account  for  this  remark- 
ably high  temperature  in  such  northern  latitudes 

JO 


146         HISTORY  OF   THE   EUROPEAN    FAUNA. 

Any  one  who  has  read  the  writings  of  the  late  Dr. 
Croll  cannot  help  being  struck  by  the  facts  he  adduces 
to  show  the  importance  of  ocean  currents  in  relation 
to  the  distribution  of  heat  over  the  globe,  and  it  seems 
to  me  that  the  view  which  attributes  the  mild  climate 
prevailing  in  former  times  in  Greenland  to  warm 
ocean  currents  reaching  the  Polar  Circle  is  the 
one  least  open  to  serious  objections.  If  we  suppose 
that  the  North  Atlantic  Ocean  was  bridged  by  a 
land-connection  between  Scandinavia  and  Greenland 
by  way  of  Spitsbergen,  and  between  Greenland  and 
North  America,  the  Polar  Ocean  would  be  practically 
a  closed  sea.  If,  then,  a  wide  passage  existed  some- 
where about  Behring  Straits  to  allow  a  warm  current 
to  enter  and  circulate  within  the  Arctic  Seas,  we 
should  have  the  southern  shores  of  Greenland  washed 
by  the  warm  Atlantic  current  and  the  northern  shores 
by  a  warm  Pacific  current,  which  combination  would 
undoubtedly  produce  the  effect  of  raising  the  tem- 
perature throughout  the  Polar  Regions  very  con- 
siderably; and  especially  would  that  be  the  case 
with  regard  to  Greenland  and  the  neighbouring 
islands. 

It  might  be  urged  that  the  constant  darkness 
during  winter  must  have  had  an  injurious  action 
upon  the  flora,  but  it  is  found  that  in  countries 
such  as  Northern  Russia,  where  southern  plants  are 
housed  during  winter  in  greenhouses,  the  light  being 
almost  entirely  excluded  by  a  covering  of  straw,  no 
serious  damage  is  done  thereby  to  the  plants. 


,THE   ARCTIC  FAUNA.  147 

It  seems  probable  that  a  similar  gradual  refrigera- 
tion 'of  climate  in  northern  latitudes  has  taken  place 
after  Miocene  times  as  has  been  proved  to  have 
occurred  in  Europe. 

Some  years  ago  Dr.  Haacke  propounded  the  hypo- 
thesis that  the  centre  of  creation  of  all  the  larger 
groups  of  animals  was  situated  in  the  region  of  the 
North  Pole,  and  that  the  newly  originated  groups  must 
always  push  the  older  ones  farther  and  farther  south 
into  the  most  remote  corners  of  the  earth.  As 
instances  of  the  correctness  of  his  view  he  quotes 
the  fact  that  the  more  ancient  mammals,  such  as 
Monotremes,  Marsupials,  Lemurs,  Edentates,  and 
Insectivores,  all  inhabit  the  more  southerly  parts 
of  the  world.  The  Apteryx,  Moa,  Rhea,  and  the 
Ostrich,  as  well  as  ^Epyornis,  which  is  only  recently 
extinct,  are  found  in  the  same  regions.  But  we  have 
no  palseontological  evidence  in  favour  of  these  ex- 
travagant views.  Fossil  Edentates  and  Marsupials 
are  almost  entirely  confined  to  the  Southern  Hemi- 
sphere, and  the  supposition  that  because  these 
primitive  mammals  inhabit  the  extreme  south  of  our 
great  continental  land-masses,  they  therefore  came 
from  the  north,  cannot  be  said  to  be  an  argument. 
Nevertheless,  I  am  quite  with  Dr.  Haacke  in  consider- 
ing that  the  North  Pole,  or,  we  might  say,  the  lands 
within  the  Arctic  Circle,  have  been  the  place  of 
origin  of  some  of  our  European  mammals,  and  there 
can  be  no  doubt  that  certain  species  in  other  groups, 
among  invertebrates  and  also  plants,  have  originated 


148         HISTORY   OF   THE    EUROPEAN   FAUNA. 

in  the  Polar  Regions.  The  facts  of  geographical 
distribution  teach  us  that  in  these  regions  there  has 
been  a  centre  of  origin  within  comparatively  recent 
geological  times.  I  have  on  a  previous  occasion 
drawn  attention  to  the  range  of  the  Reindeer:  that 
it  lives  almost  throughout  the  Polar  lands,  and  that  it 
spreads  into  North  America,  Northern  Europe,  and 
Northern  Asia.  We  have,  again,  fossil  proof  that  its 
range  extended  down  to  the  Pyrenees  in  Europe  in 
pleistocene  times.  But  there  is  not  a  scrap  of 
evidence  that  it  ever  during  any  time  occurred 
farther  south,  either  in  Europe,  Asia,  or  North 
America.  Its  original  home  must  therefore  have 
been  in  the  Polar  Regions,  for  if  it  had  originated 
either  in  Central  Europe,  Asia,  or  America,  there  is  no 
reason  why  it  should  not,  in  the  natural  course  of 
events,  have  extended  its  range  to  the  south  as  well 
as  to  the  north. 

The  Arctic  Hare  presents  us  with  a  very  similar 
case  of  distribution.  Like  the  Reindeer,  it  inhabits,  as 
we  have  learned,  the  Polar  Regions  and  the  northerly 
parts  of  the  Old  World  and  the  New;  but  while  we 
have  only  fossil  evidence  of  the  former,  more  southerly, 
extension  of  the  range  of  the  Reindeer,  the  Arctic  Hare 
furnishes  us  with  a  still  stronger  proof  of  its  past 
southward  range  in  the  survival  of  small  isolated 
colonies  in  some  of  the  southern  mountain  ranges  of 
Europe  and  Asia.  It  is  generally  believed  that  the 
occurrence  of  the  Arctic  Hare  in  these  southern  moun- 
tains is  a  standing  testimony  to  the  severity  of  the 


THE   ARCTIC   FAUNA.  149 

climate  at  the  time  when  it  commenced  its  southerly 
increase  of  range,  but  I  have  already  shown  that  the 
climate  of  Europe  at  that  time  was  not  necessarily 
colder  than  it  is  at  present,  but  that  it  may  have  been 
somewhat  milder  (p.  80).  I  think  that  a  vast  increase 
of  ice  in  the  Polar  Regions  has  taken  place  only  at.  a 
comparatively  recent  date,  and  that  both  the  Reindeer 
and  the  Arctic  Hare  originated  there  during  a  much 
more  temperate  climate  than  obtains  at  present. 
A  great  sensation  was  produced  among  European 
zoologists  and  anthropologists  when  the  discovery 
was  first  announced  that  the  remains  of  the  Reindeer 
had  been  found  in  the  Pyrenees,  and  it  naturally  gave 
rise  to  many  speculations  as  to  the  nature  of  the 
climate  at  the  time  when  its  range  extended  so  far 
south.1  The  greater  number  of  our  best  authorities 
are  still  of  opinion  that  the  existence  of  the  Reindeer 
in  Southern  Europe  points  to  the  prevalence  of  an 
arctic  climate  in  that  region.  It  is  generally  over- 
looked, however,  that  the  Reindeer-remains  occur  in 
company  with  many  typically  southern  animals,  which, 

1  A  very  interesting  piece  of  information  has  been  given  us,  recently, 
by  Mr.  Barrett- Hamilton  on  the  Arctic  Fox  of  Spitsbergen.  In  com- 
paring the  skulls  of  Spitsbergen  Foxes  with  those  of  Europe,  he  found 
that  the  former  are  much  smaller,  and  represent  a  distinct  race  or  sub- 
species. This  small  race  he  believes  to  be  confined  to  Greenland, 
Iceland,  Spitsbergen,  and  Novaya  Zemlya,  whilst  the  larger  one  occurs 
in  Europe,  Asia,  and  on  the  Commander  Islands.  This  fact  favours 
the  view  which  I  have  advocated  in  Chapter  V.,  that  the  Arctic  Fox  in 
Europe  is  a  Siberian  migrant,  and  did  not  come  from  the  north  with 
the  Reindeer  and  Arctic  Hare. 


ISO         HISTORY  OF   THE   EUROPEAN    FAUNA. 

if  they  had  been  found  alone,  would  have  been  held 
to  be  a  certain  indication  of  a  warm  climate.  The 
French  geologist  Professor  Lartet,  indeed,  was  of 
opinion  that  the  temperature  during  the  time  when 
the  Reindeer  lived  in  the  Pyrenees  must  have  been 
rather  milder  than  it  is  at  present  (compare  pp.  71-75). 
Similarly,  Mr.  Harle  argues,  that  the  extremely 
cold  climate  probably  did  not  extend  to  South- 
western France,  since  that  area  only  received  occa- 
sional visits  from  some  of  the  representatives  of  the 
Arctic  fauna. 

Long  ago  North  American  zoologists  recognised 
the  existence  in  their  country  of  two  well-marked 
races  of  the  Reindeer  (Caribou) — a  smaller  one  with 
rounded  antlers  (Fig.  10),  and  a  larger  one  in  which 
the  antlers  are  more  or  less  flattened  out  (Fig.  n). 
Two  somewhat  similar  races  can  also  be  traced  in  the 
fossil  remains  of  the  Reindeer  in  Europe.  It  was,  I 
think,  Gervais  who  first  pointed  out  that  the  Reindeer 
remains  from  the  north  of  France  differed  from  those 
found  in  the  south;  and  Lartet  referred  to  the  fact 
that  the  southern  remains  were  more  like  what,,  in 
America,  is  called  the  Barren-ground.  Caribou,  while 
those  from  Central  European  deposits  all  belonged  to 
the  Siberian  variety,  which  is  more  like  the  Wood- 
land Caribou  ot  North  America.  In  Ireland,  Pro- 
fessor Leith  Adams  also  drew  attention  to  the 
curious  fact  that  all  the  Irish  Reindeer  remains 
resemble  the  Norwegian  variety  rather  than  the 
Siberian;  and  Mr.  Murray  was  so  much  struck  by 


THE   ARCTIC   FAUNA. 


FlG.  10  — Head  of  a  Barren-ground  Reindeer  in  the  Dublin  Museum 
(photographed  by  Mr.  McGoogan). 


I  §2         HISTORY   OF   THE    EUROPEAN    FAUNA. 


FIG.  II.—  Head   of  a   Woodland    Reindeer   in   the   Dublin    Museum 
(photographed  by  Mr.  McGoogan). 


THE  ARCTIC  FAUNA.  153 

the  close  resemblance  between  the  Spitsbergen  and 
Greenland  forms  with  the  Barren-ground  Caribou,  that 
he  based  some  speculations  on  a  former  land-connec- 
tion between  these  countries  on  this  circumstance. 

We  have,  therefore,  records  of  the  present  or  the 
former  existence  of  a  Reindeer  resembling  the  North 
American  Barren-ground  form  in  Greenland,  Spits- 
bergen, Scandinavia,  Ireland,  and  the  South  of 
France.  In  England  the  remains  of  the  two  forms 
occur  mixed,  but  I  do  not  know  in  how  far  either  the 
one  or  the  other  predominates.  The  Barren-ground 
Reindeer  is  in  Europe  altogether  confined  to  the 
west;  the  most  easterly  locality  that  I  am  acquainted 
with  being  Rixdorf,  near  Berlin.  The  majority  of 
the  European  remains  of  the  Reindeer  seem  to 
belong  to  the  Siberian  or  Woodland  variety,  and  it 
would  appear  as  if  some  intercrossing  between  the 
two  forms  had  occurred  in  Lapland,  since  it  is  stated 
that  in  that  country  the  Reindeer  is  somewhat  inter- 
mediate between  the  two.  All  the  Asiatic  remains 
also  resemble  the  Woodland  variety. 

As  far  as  I  know,  no  explanation  has  been 
attempted  to  account  for  this  peculiar  range  in 
Europe  of  the  two  forms  of  Reindeer.  But  if  we  look 
more  closely  into  the  mode  of  occurrence  of  the  Rein- 
deer remains,  we  find  that  the  Barren-ground  form, 
seems  to  have  existed  in  Western  Europe  long  before 
the  other  variety  made  its  appearance  there.  It  was 
pointed  out  by  Struckmann  that  the  Reindeer  in 
Southern  Europe  occurs  in  older  deposits  than  in 


154         HISTORY   OF   THE    EUROPEAN   FAUNA. 

the  north.  In  speaking  of  the  northern  ones,  he  had  of 
course  chiefly  the  German  deposits  in  view.  It  is  in 
one  of  the  oldest  pleistocene  deposits  in  Germany 
that  the  isolated  instance,  referred  to  above,  of  the 
occurrence  of  the  Barren-ground  Reindeer,  near  Berlin, 
has  been  noted. 

There  is  still  a  further  point  which  illustrates  the 
supposition  that  the  Barren-ground  Reindeer  was  a 
more  ancient  inhabitant  of  Europe  than  the  Wood- 
land one.  The  latter  in  all  Central  European  stations 
(in  fact  almost  wherever  it  occurs  fossil)  is  asso- 
ciated with  the  remains  of  the  typical  inhabitants 
of  Siberia,  such  as  the  Glutton,  Sousliks,  Lemmings, 
and  others;  but  in  the  deposits  in  which  the  Barren- 
ground  Reindeer  have  been  found  in  South-western 
France,  no  other  Arctic  mammal  finds  a  place. 
Again,  in  Irish  deposits  none  of  the  Siberian 
migrants  are  found.  The  only  explanation  of  this 
remarkable  fact  is  that  the  two  varieties  of  the 
Reindeer  have  come  to  Europe  by  different  routes. 
We  have  learned  already  from  the  observations  of 
Mr.  Murray  that  there  are  evidences  of  the  existence 
of  a  former  land-connection  between  North  America, 
Greenland,  and  Spitsbergen.  Professor  Petersen 
tells  us  that,  according  to  recent  surveys,  a  high 
submarine  plateau  with  a  sharp  fall  of  1000  fathoms 
towards  the  Atlantic  Ocean  begins  from  Northern 
Norway  and  is  continued  as  far  as  Spitsbergen. 
Several  islands,  such  as  Bear  Island,  King  Charles 
Land,  and  others,  arise  from  this  plateau,  and  these 


THE   ARCTIC  FAUNA.  155 

must  be  looked  upon  as  the  remains  of  a  sunken 
land  (Fig.  12). 

From  Arctic  America,  thinks  Professor  Schulz 
(p.  i),  we  probably  have  had  an  uninterrupted 
migration  during  the  greater  part  of  later  Tertiary 
times  up  to  the  commencement  of  the  Pliocene  epoch 
— partly  over  a  direct  land-connection  between  Green- 
land, Iceland,  and  the  Faroes,  and  also  between  Arctic 
America,  Spitsbergen,  Franz  Josef  Land,  etc.  There 
was  also  a  connection  between  Asia  and  Alaska. 

The  distribution  of  the  Barren-ground  Reindeer  in 
Europe  seems  to  warrant  the  belief  that,  at  the  time 
it  began  its  southward  wanderings  from  the  Polar 
area,  Northern  Norway  must  have  been  connected 
with  Greenland  in  the  manner  just  indicated,  but, 
as  I  shall  explain  later  on,  Russian  Lapland  and 
part  of  Northern  Russia,  or  the  land  between  the 
White  Sea  and  the  Baltic,  must  at  that  time  have 
been  submerged  by  the  sea.  The  greater  part  of 
Denmark  and  the  lowlands  of  Sweden  were  likewise 
submerged,  but  Scandinavia  extended  south  as  far  as 
Scotland,  while  Scotland  was  connected  with  Ireland, 
and  the  latter  with  England  and  France.  The  Rein- 
deer migrating  south  into  Scandinavia  could  only 
reach  the  continent  of  Europe  by  way  of  the  British 
Islands.  It  appeared  there  in  the  west  and  gradually 
extended  its  range  east,  where,  as  I  mentioned  above, 
it  has  occurred  in  a  few  isolated  localities. 

The  advent  of  the  Woodland  form  of  the  Reindeer 
in  Europe  took  place  at  a  much  later  stage.  It  came, 


156         HISTORY  OF  THE   EUROPEAN   FAUNA. 

as  I  indicated,  with  the  hordes  of  Siberian  migrants 
which  invaded  Europe  during  what  is  known  as  the 
Inter-glacial  phase  of  the  Glacial  period.  Scan- 


FiG.  12.- — Map  of  Europe,  indicating  the  parts  which  were  probably 
submerged  (shaded)  at  the  commencement  of  the  Glacial  period. 
The  light  portions  represent,  approximately,  the  extent  of  the 
land  at  that  time. 

dinavia,  not  being  then  directly  connected  with  con- 
tinental Europe,  was  not  accessible  to  it;  neither 
was  Ireland,  which  had  by  that  time  become  dis- 


THE  ARCTIC   FAUNA.  157 

connected  from  Great  Britain.  None  of  the  Siberian 
migrants  seem  to  have  been  able  to  cross  the  River 
Garonne,  and  \ve  therefore  find  neither  the  Woodland 
Reindeer  nor  any  of  the  typical  Siberian  species 
represented  in  the  Pyrenean  deposits. 

The  Woodland  Reindeer  persisted  in  continental 
Europe  until  comparatively  recent  times,  and  it  has 
since  made  its  way  into  Scandinavia  across  Northern 
Russia,  and  probably  mingled  with  the  older  stock  of 
the  Barren -ground  form.  In  the  same  way,  it  may 
have  come  about  that  in  the  English  pleistocene 
deposits  the  remains  of  the  two  races  occur. 

In  a  recent  contribution  to  our  knowledge  of  the 
deer  tribe  (c,  p.  88),  Mr.  Lydekker  suggests  that  the 
former  division  of  the  Reindeer  races  into  the  two 
forms  of  Woodland  and  Barren-ground  Caribou,  no 
longer  holds  good.  He  now  recognises  no  less  than 
six  races,  as  follows  :— 

1.  Rangifer  tarandus  typicus. 

2.  „  „  spitzbergensis. 

3.  „  „  caribou. 

4.  „  „  terrae-novse. 

5.  „  „  grcenlandicus. 

6.  „  „  arcticus. 

I  hardly  think  these  can  be  considered  of  equal 
value;  indeed,  though  there  may  be  differences 
between  R.  groenlandicus,  typicus,  arcticus,  and  spilz- 
bergensis,  the  antlers  exhibit  a  certain  much  closer 
relationship  among  one  another  than  to  R.  terrcz- 
novce  and  caribou.  But  the  whole  subject  is  by  no 


158         HISTORY  OF  THE   EUROPEAN   FAUNA. 

means  as  well  known  as  could  be  wished,  and  a  very 
careful  comparative  study  of  recent  and  fossil  remains 
of  the  Reindeer  from  various  parts  of  the  Old  and 
New  Worlds  is  much  needed  to  put  our  views  on  a 
firmer  basis. 

The  presence  of  the  Arctic  Hare  in  Ireland  and 
the  absence  of  the  common  European  Hare  (Lepus 
europceus)  can  be  explained  in  a  somewhat  similar 
manner.  The  Arctic  Hare  is  the  older  of  the  two 
species — corresponding  with  the  Barren-ground  Rein- 
deer— and  the  European  Hare  the  newer  one, 
associating,  like  the  Woodland  Reindeer,  in  its 
westward  migration  with  Siberian  animals,  though 
probably  of  Oriental  origin. 

Let  us  once  more  refer  back  again  to  the  map  on  page 
137  indicating  the  geographical  distribution  of  the 
Arctic  Hare.  Its  discontinuous  range  and  its  isolated 
position  in  the  Alps,  Pyrenees,  and  the  Japanese 
mountains,  all  tend  to  show  that  it  is  an  ancient 
species.  Moreover,  its  presence  in  Ireland  in  the 
plain  as  well  as  in  the  mountains,  clearly  points  to  the 
fact  that,  in  the  British  Islands  at  any  rate,  the  Arctic 
Hare  was  the  first  comer,  and  that  subsequently  the 
European  Hare  invaded  these  countries.  It  probably 
found  Ireland  then  no  longer  accessible,  having  since 
become  separated  from  England.  Again  and  again 
do  we  find  the  statement  repeated,  that  the  presence 
of  the  Arctic  Hare  in  Europe  is  a  clear  proof  of  the 
former  prevalence  in  our  continent  of  an  Arctic  climate. 
But  if  so,  why  should  this  Hare  at  present  live  and 


THE   ARCTIC  FAUNA.  159 

thrive  in  Ireland,  which  has  a  particularly  mild  climate 
in  winter,  and  be  absent  from  so  many  continental 
stations  where  the  temperature  more  resembles  that 
of  its  native  home?  If  we  suppose  that  the  European 
Hare  migrated  to  Europe  from  the  east,  after  the 
Arctic  Hare  had  become  established  in  Western 
Europe,  and  drove  the  latter  into  the  mountains 
or  northward  whenever  the  two  came  into  contact, 
we  should  have,  it  seems  to  me,  a  better  ex- 
planation of  the  range  presented  by  the  two  species. 
I  was  formerly  of  opinion  that  the  European  Hare 
had  come  with  the  Siberian  animals  from  Siberia, 
but  it  appears  to  me  more  likely  now,  that  it  reached 
our  continent  with  the  Oriental  migrants,  and  only 
then  joined  the  Siberians  in  Eastern  Europe. 

The  evidence  in  favour  of  a  former  land-connection 
between  Scandinavia  and  Greenland,  rests  on  many 
other  facts  besides  those  already  brought  forward. 
That  some  form  of  land-connection  formerly  existed 
between  Europe  and  Greenland  is  now  indeed  almost 
universally  accepted.  That  it  was  situated  more  to 
the  south  between  Scotland  and  Greenland  is  a  sup- 
position which  has  been  actively  supported  by  many 
leading  authorities,  but  it  seems  to  me  that  if  such  a 
land-bridge  existed,  it  must  have  been  in  very  early 
Tertiary  times,  whilst  the  northern  one,  such  as  I 
have  indicated,  may  have  originated  later  and  per- 
sisted until  a  recent  geological  date. 

The  distribution  of  few  groups  of  animals  is  now 
better  known  than  that  of  the  larger  butterflies  and 


l6o         HISTORY  OF  THE   EUROPEAN    FAUNA. 

moths  (Macro-lepidoptera}\  even  those  of  Siberia  have 
been  fairly  well  investigated.  The  interesting  facts  ob- 
tainable from  their  distribution  are  therefore  of  special 
value.  No  less  than  243  species  of  Lepidoptera  are 
mentioned  by  Moschler  as  being  common  to  North 
America  and  Europe.  It  is  extremely  probable  that 
a  fair  number  of  these  have  either  migrated  direct 
from  America  to  Europe  or  vice  'versa,  though  many 
may  be  of  Asiatic  origin,  and  have  wandered  east 
and  west  from  their  original  home.  The  following 
twelve  species  are  mentioned  by  Petersen  (p.  38) 
as  occurring  in  Arctic  Europe  and  also  in  Arctic 
North  America,  but  not  in  Asia : — Colzas  nastes, 
Colias  hecla,  Syrichthus  centaurece,  Pachnobia  carnea, 
Plusia  parilis,  Anarta  Richardsoni,  Anarta  Schon- 
herri,  Anarta  lapponica,  Anarta  Zetterstedti^  Cidaria 
frigidaria,  Cidaria  polata,  Eupithecia  hyperboreata; 
and  these,  as  he  remarks,  point  to  the  possibility 
of  a  former  direct  land-connection  between  Europe 
and  North  America. 

Mr.  Petersen  believes  that  the  chief  immigration 
into  the  Arctic  area  of  Europe  is  post-glacial  and 
took  place  from  Siberia,  since  the  majority  of  the 
species  are  still  to  be  found  in  that  country  at  the 
present  day  (p.  57).  He  also  draws  particular  atten- 
tion to  a  fact, — which  I  shall  discuss  more  fully  in  the 
next  chapter, — namely,  that  the  most  characteristically 
Arctic  forms  of  Northern  Europe,  which  also  partly 
occur  in  the  Alps,  are  entirely  absent  from  the 
Caucasus. 


THE   ARCTIC    FAUNA.  l6l 

Adopting  the  glacial  views  of  some  of  our  leading 
geologists,  Petersen  comes  to  the  logical  conclusion 
that  Central  Europe  could  not  have  possessed 
any  butterflies  during  the  height  of  the  Glacial 
period,  but  since  all  evidences  seem  to  point  to  the 
chief  migration  from  Siberia  having  taken  place  after 
the  Glacial  period,  he  concludes  that  they  must  have 
survived  the  severe  cold  of  that  time  in  Central  Asia. 
He  leaves  us,  however,  to  imagine  .under  what  possible 
geographical  conditions  the  climate  in  Europe  could 
be  too  severe  for  a  lepidopterous  fauna,  while  at  the 
same  time  Central  Asia  could  maintain  an  abundant 
one. 

In  a  suggestive  note  on  the  origin  of  European 
and  North  American  Ants,  Professor  Emery  states 
(p.  399)  that  a  great  number  of  North  American 
ants  are  specifically  identical  with  European  ones; 
whilst  Dr.  Hamilton  tells  us  (p.  89),  as  an  instance, 
that  specimens  of  the  beetle  Loricera  ccemJescens 
from  Lake  Superior  and  from  Scotland  do  not  seem 
to  vary  to  the  extent  of  a  hair  on  the  antennae. 
He  enumerates  487  species  of  Coleoptera  as  being 
common  to  North  America,  Northern  Asia,  and 
Europe,  many  of  which  no  doubt  have  migrated  by 
the  Americo-European  land-connection. 

Arctic  Scandinavia  or  Lapland,  according  to  Sir 
Joseph  Hooker,  contains  three-fourths  of  the  entire 
number  of  species  of  plants  known  from  the  whole 
circumpolar  area.  His  view,  that  the  Greenland 
flora  is  almost  exclusively  Lapponian, — having  only 

II 


162         HISTORY   OF  THE   EUROPEAN    FAUNA. 

an  extremely  slight  admixture  of  American  or 
Asiatic  types, — again  points  to  a  former  more 
intimate  connection  between  North  America  and 
Arctic  Europe,  and  indeed  he  remarks  (p.  252), 
"  It  is  inconceivable  to  me  that  so  many  Scan- 
dinavian plants  should,  under  existing  conditions 
of  sea,  land,  and  temperature,  have  not  only  found 
their  way  to  Greenland  by  migration  across  the 
Atlantic,  but  should  have  stopped  short  on  its 
western  coast  and  not  crossed  to  America." 

Hooker's  view,  that  the  Scandinavian  flora  is  of 
great  antiquity,  that,  at  the  advent  of  the  Glacial 
period,  it  was  everywhere  driven  southwards,  and 
that  during  the  succeeding  warm  epoch  the  sur- 
viving species  returned  north,  has  been  adopted  by 
the  great  majority  of  naturalists. 

The  natural  corollary  of  this  'theory  is  that  there 
must  have  been,  between  the  beginning  of  the  Glacial 
period  and  the  present  time,  either  two  independent 
land-connections  between  the  Polar  Regions  and 
Northern  Europe  at  different  epochs  to  enable 
animals  and  plants  to  travel  southwards  and  once 
more  to  regain  their  former  northern  home,  or, 
that  during  the  whole  of  the  Glacial  period  the 
Polar  Regions  were  uninterruptedly  connected  with 
Northern  Europe,  until  the  fauna  and  flora  had  once 
more  reached  their  northern  goal,  after  the  Polar 
lands  had  been  desolated  by  the  supposed  rigours  of 
that  period. 

In  following  the  history  of  the  Arctic  migration  to 


THE  ARCTIC  FAUNA.  163 

Europe,  it  is  of  great  importance  to  determine  the 
nature  and  the  time  of  duration  of  these  land- 
connections.  The  Greenland  flora  is  a  very  in- 
structive one  in  helping  us  to  understand  many 
of  the  problems  connected  with  the  origin  of  the 
European  plants  and  animals.  To  judge  from  the 
remarks  of  Professor  James  Geikie  and  Mr.  Clement 
Reid,  no  flowering  plants  could  have  existed  in 
the  British  Islands  during  the  height  of  the  Glacial 
period,  and  one  would  suppose  that  the  cold  in  Green- 
land at  that  time  must  have  been  far  more  intense 
than  in  England.  If  no  flowering  plants  could  exist 
in  the  latter  country,  then  very  surely  none  could  in 
Greenland,  where  the  climate  was  of  necessity  by  far 
more  rigorous.  It  will  be  a  surprise,  therefore,  to 
those  who  are  acquainted  only  with  Professor  Geikie's 
views  of  the  nature  of  the  Glacial  period,  that  two  of 
the  most  eminent  Swedish  botanists,  who  have  made 
a  special  study  of  the  flora  of  Greenland,  have  come 
to  the  conclusion  that  a  survival  of  flowering  plants 
has  taken  place  in  Greenland  itself  from  pre-glacial 
times.  According  to  Professor  Nathorst  (p.  200), 
only  a  few  plants  could  have  survived  the  Glacial 
period  in  Greenland.  The  species  now  peculiar  to 
that  country  may  perhaps,  he  thinks,  be  the  remnants 
of  those  which  existed  in  pre-glacial  times.  Mr. 
Warming,  on  the  other  hand,  is  of  opinion  that  the 
main  mass  of  Greenland's  present  flora  survived  the 
Glacial  period  there  (p.  403),  and  that  the  remainder 
was  carried  from  Europe  and  North  America  by 


164         HISTORY  OF  THE   EUROPEAN    FAUNA. 

occasional  means  of  distribution  of  the  nature  in- 
dicated by  Darwin. 

Very  similar  views  on  the  origin  of  the  present 
Polar  flora  are  expressed  by  Colonel  Feilden,  who 
says,  "  To  my  mind  it  seems  indisputable  that  several 
plants  now  confined  to  the  Polar  area  must  have 
originated  there  and  have  outlived  the  period  of 
greatest  ice-development  in  that  region"  (£,  p.  50). 
No  land-connection  at  all  need  be  supposed  to  have 
existed  in  recent  geological  times,  that  is  to  say, 
during  the  Glacial  period  or  after,  if  Mr.  Warming's 
and  Colonel  Feilden's  views  be  adopted.  A  pre-glacial 
connection  would  be  sufficient  to  explain  the  general 
features  of  distribution.  An  admission  is  thus  ob- 
tained from  these  two  independent  authorities  that 
the  climate  during  the  Glacial  period  must  have 
been  vastly  less  severe  in  the  Polar  Regions  than 
is  generally  conceded.  I  am  of  opinion  that  not 
only  the  whole  of  the  present  flora,  but  also  the 
fauna  of  Greenland  survived  the  Glacial  period  in 
that  country. 

If  we  suppose  that  an  extensive  centre  of  origin 
existed  in  the  Polar  area,  or  we  may  say  in  Green- 
land, both  animals  and  plants  would  have  been  able 
to  spread  from  it  into  Northern  Europe  and  North 
America  by  means  of  the  land-connections  which  are 
generally  supposed  to  have  existed  in  pliocene  times, 
that  is  to  say,  just  before  the  commencement  of  the 
Glacial  period.  There  must  have  been  at  this  time 
a  connection  too  between  Scotland  and  Scandinavia, 


THE   ARCTIC   FAUNA.  465 

which  will  be  dealt  with  more  fully  presently.  The 
important  point  is  to  consider  what  light  the  Green- 
land flora  arid  fauna  will  throw  upon  the  problem 
of  the  continuity  of  the  aforesaid  land-connec- 
tion during  the  Glacial  period.  We  have  seen 
that  the  Barren-ground  Reindeer,  a  typically  Polar 
species,  penetrated  as  far  south  as  the  Pyrenees,  the 
Arctic  Hare  went  as  far,  while  a  number  of  other 
species  of  Polar  animals  and  also  of  plants  occur 
in  the  Alps.  Of  these  it  remains  to  be  seen  how 
many  have  come  direct  by  way  of  Northern  Europe 
or  from  the  Polar  Regions  by  way  of  Asia.  At  any 
rate,  as  the  origin  of  the  Alpine  animals  and  plants 
will  be  discussed  in  another  chapter,  there  is  no  need 
to  dwell  on  this  subject  at  present. 

From  the  nature  of  the  distribution  in  Ireland  of 
Arctic  plants  and  animals,  which  occur  mostly  on  the 
north  and  west  coasts,  it  would  seem  that  a  stream  of 
migration  entered  from  Scotland,  and  I  have  no  doubt 
that  that  same  migration  came  into  Scotland  directly 
from  Scandinavia  by  a  route  over  which  now  roll  the 
waves  of  the  North  Sea.  There  is,  moreover,  as  I 
already  mentioned  on  p.  94,  a  very  interesting  so-called 
American  element  in  the  north-western  European 
flora,  that  is  to  say,  plants  now  found  in  North-west 
Europe  and  North  America  without  occurring  in 
Greenland  or  any  of  the  islands  which  might  have 
formed  the  former  highway  between  the  Old  World 
and  the  New.  These  are  probably  some  of  the  more 
ancient  Polar  plants  which  have  become  extinct  in 


166          HISTORY  OF   THE   EUROPEAN   FAUNA. 

the  Arctic  Regions  and  survive  in  isolated  patches  in 
favourable  localities.  We  find  seven  species  of  these 
American  plants  in  Ireland,  almost  entirely  confined 
to  the  north  and  west  coasts.  These  are  Spiranthes 
Romanzoviana,  Sisyrinchium  anceps,  Naias  flexilis, 
Eriocaulon  septangular e^Juncus  tennis •,  and  Polygonum 
sagittifoliuui.  To  them  must  be  added  another  plant 
recently  discovered  by  the  Rev.  Mr.  Marshall  in  the 
south  of  Ireland,  namely  Sisyrinchium  calif  or nicum. 
As  I  have  mentioned  in  former  writings,  there  are 
three  species  of  North  American  freshwater-sponges 
in  Ireland  which  have  not  hitherto  been  discovered 
elsewhere  in  Europe  or  in  Asia.  These,  namely 
Ephydatia  crater  if ormis,  Heteromeyenia  Ryderi,  and 
Tubella  pennsylvanica,  all  occur  in  some  of  the  lakes 
near  the  western  coast  of  Ireland. 

There  are  in  all  groups  of  animals  instances  of  species 
which  are  confined  to  Europe  and  North  America,  while 
unknown  from  the  Asiatic  continent,  but  none,  as  far 
as  is  known,  have  such  a  very  discontinuous  range  as 
that  of  the  animals  and  plants  just  referred  to.  In 
some  cases  the  species  still  occur  in  Greenland,  and 
in  this  way  make  it  still  clearer  that  their  migration 
in  former  times  took  place  from  one  continent  to 
the  other  by  way  of  that  country.  As  an  interesting 
instance  of  such  distribution  may  be  mentioned  the 
Common  Stickleback  (Gasterostens  aculeatus],  which 
is  found  in  Greenland,  North  America,  and  Europe, 
but  is  quite  absent  from  Asia.  Then  again,  the  Nine- 
spined  Stickleback  (Gasterosteus  pungitius]  is  confined 


THE  ARCTIC  FAUNA.  167 

to  Western  Europe  and  North  America,  though  an 
allied  species,  Gasterostcus  sinensis,  lives  in  China  and 
has  probably  penetrated  there  from  the  New  World 
across  the  old  Behring  Straits  land-connection. 

The  Coleoptera  Diachila  arctica,  Elaphrus  lapponicus, 
and  Blethisa  multipunctata  are  good  instances  of 
species  which  have  come  to  us  from  North  America 
by  way  of  Greenland.  I  have  already  referred  to 
the  Lepidoptera,  but  might  add  that  eleven  species 
of  Anarta  occur  in  Scandinavia,  eight  of  which 
reappear  again  in  Labrador,  none  of  them,  however, 
being  met  with  in  Siberia.  Then  again,  take  the  in- 
teresting Crustacean  Lepidurus  (Apus]  glacialis.  It 
is  found  in  Greenland,  Spitsbergen,  Lapland,  and 
Norway;  and  formerly,  as  we  know  from  fossil 
evidence,  it  ranged  into  Scotland.  Another  Phyllo- 
pod,  viz.,  Branchinecta  palndosa,  inhabits  Greenland, 
Lapland,  and  Norway.  Mr.  Kennard  suggests  that 
the  freshwater  Snail  Planorbis  glaber  might  also 
belong  to  the  same  migration.  And  there  are  no 
doubt  large  numbers  of  others. 

Professor  Emery  mentions  that  Northern  Europe 
possesses  one  peculiar  genus  of  Ant,  viz.,  Anergates. 
This  is  closely  allied  to  Epoccus,  another  genus  con- 
fined to  North  America.  It  seems  probable,  there- 
fore, that  both  of  these  have  sprung  from  an  Arctic 
genus  which  sent  two  branches  southward  into  the 
two  continents  without  there  being  any  migration 
through  Asia. 

The  general  range  of  the  Arctic  plants  and  animals 


1 68         HISTORY   OF   THE   EUROPEAN    FAUNA. 

gives  no  reason  to  suppose  that  the  Greenland  fauna 
and  flora  of  the  present  day  were  exterminated  by  the 
Glacial  period  and  then  reintroduced  into  that  country. 
Nor  have  we  any  evidence  that  such  a  fauna  and  flora 
migrated  across  the  British  Islands  northward.  The 
Greenland  animals  and  plants  too  are  altogether  much 
more  like  the  Lapland  ones  than  those  of  Scotland. 
It  will  also  become  evident  to  the  reader  of  this  work 
that  no  very  extensive  migrations  could  have  taken 
place  during  the  post-glacial  period,  and  that  almost 
everything  points  to  a  survival  of  both  fauna  and 
flora  in  northern  latitudes  throughout  the  Glacial 
period. 

If  we  take  into  consideration  the  palaeontological 
evidence  of  the  two  races  of  Reindeer  in  Europe,  one  of 
which  came  to  us  from  the  north,  and  that  the  Arctic 
Hare  and  one  of  the  races  of  the  Stoat  entered  our 
continent  from  the  same  direction — when  we,  more- 
over, carefully  review  the  numerous  other  instances 
quoted  of  plants  and  animals  which  could  only  have 
reached  us  from  the  north,  the  irresistible  conclusion  is 
forced  upon  us  that  a  land-connection  existed  at  no 
very  distant  period  between  Northern  Europe  and  the 
Arctic  Regions  of  North  America.  This  is  not  a  new 
hypothesis.  Many  geologists  are  of  opinion  that  a 
land-passage  did  exist  within  comparatively  recent 
times,  uniting  Europe,  Greenland,  and  North  America. 
But  the  position  of  this  old  land-bridge,  as  I  have 
mentioned,  has  been  generally  placed  somewhat 
farther  south  than  I  should  feel  inclined  to  put  it. 


THE  ARCTIC   FAUNA.  169 

The  fact  that  very  extensive  glaciers  formerly 
covered  the  mountains  of  Scandinavia  on  the  eastern 
side,  whilst  they  scarcely  reached  the  sea  on  the  west 
(Feilden,  a,  p.  721),  seems  to  favour  the  view  of  a 
warm  current  having  washed  the  western  shores.  As 
I  shall  attempt  to  show  later  on  (p.  179),  the  Arctic 
Ocean  extended  across  Northern  Russia  at  that  time 
from  the  White  Sea  to  the  Baltic — that  is  to  say,  to 
the  eastern  shores  of  Scandinavia,  which  country  was 
then  joined  to  the  north  of  Scotland.  The  predis- 
posing agents  to  a  copious  snowfall  existed  in 
Scandinavia,  viz.,  an  excessive  evaporation  of  the 
warm  Atlantic  waters  and  unusual  precipitation  in 
the  form  of  snow  owing  to  the  cold  given  off  by 
the  Arctic  waters  on  the  east  side  of  the  mountains. 
It  is  therefore  probable  that  the  land -connection 
which  united  Europe  and  North  America  was  farther 
north  than  has  been  supposed. 

If  we  sail  straight  across  from  Northern  Scandinavia 
to  Greenland,  we  traverse  an  exceedingly  deep  marine 
basin ;  but  if  we  examine  the  sub-marine  bank  which 
runs  all  along  the  coast  of  the  former  country  from 
south  to  north,  we  find  that  it  does  not  end  when  the 
extreme  north  of  the  land  is  reached.  The  bank 
extends  much  farther  north,  and  is  continued  as  far 
as  Spitsbergen.  As  I  have  said  before,  the  latter, 
as  well  as  Bear  Island,  must  be  looked  upon  as  the 
remains  of  a  large  mass  of  sunken  land — the  ancient 
Scandinavia  stretching  far  into  the  Arctic  Circle.  Pro- 
fessor Nathorst  speaks  of  Spitsbergen  as  a  northern 


170         HISTORY  OF   THE   EUROPEAN    FAUNA. 

continuation  of  Europe,  not  only  geographically,  but 
also  botanically  and  geologically.  However,  this 
northern  land  must  have  stretched  even  farther — not 


FIG.  13  — Map  of  Europe,  indicating  approximately  the  distribution  of 
land  and  water  during  the  earlier  stages  of  the  Glacial  period — 
shortly  after  the  period  represented  in  Fig.  12,  p.  156.  The  darkly 
shaded  parts  indicate  the  areas  covered  by  water,  and  the  white 
portions  what  was  land  at  the  time. 

perhaps  farther  north,  but  farther  west.  Here  lay  the 
old  land-connection  between  Scandinavia,  Greenland, 
and  North  America  (Fig.  13).  One  of  the  highest 


THE  ARCTIC   FAUNA.  171 

authorities  on  the  geographical  distribution  of  plants, 
Professor  Engler,  maintains  that  the  arguments  in 
favour  of  this  Arctic  connection  of  America  with 
Europe  are  more  weighty  than  those  for  a  land-bridge 
between  Greenland,  Iceland,  the  Faroes,  and  Great 
Britain.  Moreover,  he  is  of  opinion  that  a  certain 
number  of  species  of  plants  belonging  to  the  Alpine 
flora  of  Arctic  Siberia  have  travelled  from  Scandinavia 
via  Greenland  and  North  America  to  Eastern  Asia, 
and  not  direct  from  Scandinavia  to  Siberia  (p.  143). 

That  this  ancient  Arctic  land-connection  existed 
almost  throughout  the  Glacial  period  appears  to  me 
probable.  It  has  often  been  suggested  that  such  a 
land-barrier  was  one  of  the  principal  causes  of  the 
production  of  the  glacial  phenomena  in  Europe, 
and  as  such  it  must  have  existed  intact  certainly 
during  the  earlier  stages  of  the  Glacial  period. 
The  barrier  must  then  have  gradually  subsided  in 
one  or  two  places ;  and  once  a  breach  was  formed, 
the  complete  union  between  the  Atlantic  and 
the  Arctic  Oceans  could  not  have  been  long 
delayed. 

The  terrestrial  fauna  and  flora,  as  we  have  seen, 
lend  strong  support  to  the  view  of  the  former 
connection  between  Scandinavia  and  Greenland,  but 
many  other  facts  point  in  the  same  direction.  It  was 
Edward  Forbes  who  first  drew  attention  to  the  pres- 
ence of  a  number  of  species  of  littoral  molluscs  on 
the  coast  of  Finmark  which  also  occur  on  the  coast  of 
Greenland,  and  he  expressed  the  firm  conviction  that 


172         HISTORY  OF   THE   EUROPEAN    FAUNA. 

they  indicated  by  their  existence  on  both  sides  of  the 
Atlantic  some  ancient  continuity  of  the  coast-line. 
He  held  that  the  line  of  migration  of  these  mollusca 
was  probably  from  west  to  east,  and  that  it  must 
have  taken  place  during  physical  conditions  entirely 
different  from  those  prevailing  at  present  If  Forbes's 
view  is  correct,  a  current  must  have  existed  from  the 
north  coast  of  North  America  along  the  northern 
shore  of  the  ancient  land  which  stretched  east  as 
far  as  Europe.  We  have  also  some  palaeontological 
evidence  bearing  on  the  existence  of  such  a  current 

(P-  173). 

As  we  shall  learn  presently,  the   early  stages  of 

the  Glacial  period  were  accompanied  by  a  marine 
transgression  over  Northern  Russia  and  Germany — 
an  overflow,  as  it  were,  of  the  waters  of  the  Arctic 
Ocean  covering  a  great  part  of  Northern  Europe, 
with  the  exception  of  Norway.  One  continuous 
ocean  ultimately  extended  from  the  east  coast 
of  England  across  Holland,  Northern  Germany,  and 
Russia  to  the  White  Sea  (Fig.  12,  p.  156).  The 
south  of  England  being  at  that  time  joined  to 
France,  and  Scotland  to  Scandinavia,  there  was  no 
direct  communication  between  this  large  North 
European  Sea  and  the  Atlantic.  The  glaciers 
which  took  their  origin  in  the  Scandinavian  Moun- 
tains discharged  icebergs  into  this  sea,  and  many  of 
them  no  doubt  were  stranded  on  the  east  coast  of 
England.  The  boulders  of  Scandinavian  origin  which 
have  been  discovered  in  recent  geological  deposits  on 


THE  ARCTIC  FAUNA.  173 

that  coast  have  generally  been  traced  to  the  action  of 
land-ice,  but  the  supposition  that  they  have  been 
carried  by  icebergs — the  older  theory — appears  to 
me  the  more  probable  one.  Such  boulders  begin 
to  make  their  first  appearance  in  the  Red  Crag,  a 
deposit  which  is  now  looked  upon  as  belonging  to 
the  newer  pliocene  series.  But  whether  we  call  it 
pliocene  or  pleistocene  really  matters  little.  The 
important  fact  is,  that  glacial  phenomena,  consisting 
of  the  appearance  of  boulders  foreign  to  the  country 
together  with  an  invasion  of  Arctic  shells,  are  now 
ushered  in  upon  a  coast  which  shortly  before  teemed 
with  the  southern  life  of  a  Mediterranean  character. 
Among  the  new  arrivals  in  these  English  crags  there 
are  no  less  than  eighteen  species  of  North  American 
marine  mollusca.  Since  the  German  Ocean  had  then 
no  direct  communication  with  the  Atlantic,  these 
mollusca  could  only  have  come  from  the  White 
Sea,  and  Forbcs's  Arctic  current  would  offer  an 
explanation  of  the  manner  in  which  they  were 
enabled  to  migrate  there  from  their  original 
home. 

It  might  be  urged  that  we  have  no  grounds 
for  the  supposition  that  the  German  Ocean  was 
practically  a  closed  basin;  and  that  these  American 
species  probably  inhabited  at  that  time  the  whole  of 
the  North  Atlantic  Ocean.  But  if  such  had  been  the 
case,  we  ought  to  have  evidence  of  the  occurrence  of 
some  of  these  species  in  the  newer  Tertiary  deposits 
along  the  west  coasts  of  the  British  Islands.  Such 


174         HISTORY  OF  THE   EUROPEAN   FAUNA. 

beds  exist;  there  is,  however,  not  a  trace  in  any 
of  them  of  any  American  mollusca.  In  examin- 
ing the  marine  deposits  of  St.  Erth,  on  the  coast  of 
Cornwall,  which  are  believed  to  be  of  about  the  same 
age  as  the  newer  crags,  Messrs.  Kendall  and  Bell 
were  much  struck  by  the  absence  of  the  species 
characteristic  of  the  latter.  The  St.  Erth  fauna  led 
them  to  believe  that  the  Arctic  Ocean  could  not 
then  have  opened  into  the  Atlantic,  but  that  a  land- 
communication  had  existed  between  Europe  and 
North  America,  so  as  to  form  a  barrier  of  separation 
between  the  two  oceans.  This  again  perfectly  har- 
monises with  the  views  I  have  expressed,  and 
supports  them. 

Let  us  now  look  a  little  more  closely  at  the 
history  and  the  fauna  of  the  Baltic  and  the  adjoining 
lakes,  in  order  to  gain  additional  information  as  to  the 
geographical  changes  which  have  had  such  lasting 
influence  on  the  peninsula  of  Scandinavia,  The 
Baltic  is  a  shallow  sea  covering  an  area  of  184,496 
square  miles,  and  its  waters  are  decidedly  brackish. 
The  fauna  is  a  poor  one,  being  too  salt  for  the  purely 
freshwater  species  and  not  salt  enough  for  the  typical 
marine  forms.  The  absence  of  some  animals  which 
we  should  expect  to  find  there  is  one  of  the  remark- 
able features  about  the  Baltic,  but,  on  the  other  hand, 
some  species  occur  which  are  altogether  strangers  to 
the  fauna.  And  these,  moreover,  are  confined  to  the 
extreme  northern  end  of  the  sea.  I  need  only  refer 
to  the  Arctic  Seal  (P/wca  annelata),  which  is  confined 


THE   ARCTIC   FAUNA.  175 

to  the  Gulf  of  Bothnia,  and  to  the  four-horned  sting- 
fish  (Cottus  quadricornis,  Fig.  14,  p.  178),  neither  of 
which  occur  on  the  west  coast  of  Scandinavia.  But 
there  are  others  which  point  in  an  equally  unmistakable 
manner  to  the  former  existence  of  a  marine  connection 
between  the  Baltic  and  the  southward  prolongation  of 
the  Arctic  Ocean — known  as  the  White  Sea.  It  is 
generally  admitted  now  that  such  a  union  between 
these  two  seas,  viz.,  the  Baltic  and  the  White  Sea, 
occurred  in  recent  geological  times,  but  opinions 
differ  as  to  the  duration  of  this  connection.  I 
adhere  to  the  view  expressed  by  Murchison  and 
others,  that  the  boulder-clay  is  a  marine  deposit.  I 
am  also  convinced  that  the  Arctic  Ocean,  as  I  have 
already  mentioned,  transgressed  over  the  lowlands 
of  Northern  Russia  at  about  the  time  when  the 
newer  crags  were  being  deposited  on  the  east  coast 
of  England;  that  the  same  large  sea  also  covered 
Northern  Germany,  Denmark,  Holland,  and  the  low- 
lands of  Sweden,  and  laid  down  the  lower  continental 
boulder-clay  which  is  spread  over  such  vast  tracts  of 
land  in  those  countries.  I  shall  have  occasion  to  refer 
to  this  again  more  fully  in  the  next  chapter;  mean- 
while, it  should  be  remembered  that  this  stage  was 
followed  by  a  partial  retreat  of  the  northern  sea, 
though  Scandinavia  did  not  become  joined  to  the 
Continent.  The  date  of  this  retreat  of  the  sea,  repre- 
sented in  Fig.  13,  corresponds  probably  to  what  is 
know  as  the  inter-glacial  phase  of  the  Glacial  period, 
and  I  think  it  must  have  been  during  this  time  that 


1/6         HISTORY  OF  THE   EUROPEAN    FAUNA. 

the  Forest-Bed   on  the   coast   of  Norfolk   was   laid 
down.1 

None  of  the  Siberian  mammals  apparently  entered 
Scandinavia  at  the  time  when  they  invaded  Central 
Europe  and  penetrated  as  far  west  as  England  and 
Western  France.  Nor  did  the  great  Oriental  mammals, 
like  the  Mammoth  and  others,  reach  Scandinavia;  and 
Professor  Pohlig  argued,  on  the  strength  of  these 
facts,  that  the  latter  country  was  either  for  a 
very  short  time  only  free  from  ice,  or  that  it  had 
defective  land-communication  with  the  Continent 
during  inter -glacial  times.  This  seems  to  me 
scarcely  to  explain  the  facts  of  distribution  and 
account  satisfactorily  for  the  absentees.  Nor  does  it, 
of  course,  harmonise  with  the  views  that  I  have 
announced  above.  Professor  Engler's  remark  (p.  131), 
that  Scandinavia  probably  projected  above  the  glacial 
sea  as  an  island,  is  more  in  accordance  with  these 
views,  though  the  term  island  is  scarcely  applicable 
to  that  country,  since  it  was  always,  as  I  said,  in- 
directly joined  to  the  Continent  (vide  Fig.  13,  p.  170). 
The  fauna  of  Scandinavia,  both  fossil  and  recent,  points 
to  a  direct  isolation  of  that  country  from  the  continent 
of  Europe  during  a  considerable  period. 

Another  proof  that  Northern  Russia  and  the  low- 
lands of  Sweden  were  covered  by  the  sea  comes  to  us 
from  a  study  of  the  fauna  of  the  relict  lakes — the 
"  Reliktenseen  "  of  Leuckart.  This  name  was  first 
applied  by  Leuckart  to  lakes  containing  marine 

1  I  have  already  expressed  this  view  on  p.  120. 


THE   ARCTIC   FAUNA.  1 77 

organisms,  which  are  supposed  to  have  been 
flooded  by,  or  to  have  been  in  close  communication 
with  the  sea  at  some  former  period,  like  the  lakes 
Ladoga  and  Onega  in  Russia.  His  views  have  been 
worked  out  subsequently  in  greater  detail  by  Loven 
and  O.  Peschel,  who  gave  them  their  strong  adherence. 
Many  leading  zoologists,  such  as  Professor  Sars  and 
others,  have  since  adopted  them,  and  though  dis- 
credited by  Professor  Credner,  the  theory  still 
offers  the  best  explanation  for  the  origin  of  marine 
animals  in  freshwater  lakes. 

Professor  Credner's  contention,  that  marine  mollusca 
are  always  absent  from  these  relict  lakes,  seems  at  first 
sight  a  stumbling-block  to  the  theory.  But  the  ex- 
planation is  really  simple  enough.  It  is  to  Dr.  Sollas 
that  we  owe  a  very  ingenious  explanation  of  the  origin 
of  freshwater  faunas.  He  showed  that  all  freshwater 
organisms  in  their  early  stages  of  development  are  pro- 
vided either  with  some  process  enabling  them  to  attach 
themselves  to  a  foreign  object,  or  that  they  pass  this 
period  within  the  body  of  the  parent.  This  is  a 
provision  of  nature  to  prevent  freshwater  organisms 
from  being  floated  out  to  sea,  where  they  would 
perish,  until  they  reach  maturity  and  can  cope  with 
floods  and  currents.  Had  Professor  Credner  been 
aware  of  Dr.  Sollas's  views,  no  doubt  he  would  have 
modified  his  criticisms,  for,  as  most  marine  mollusca 
have  free- swimming  larvae,  they  would  have  little 
chance  of  becoming  permanent  residents  of  lakes. 
During  their  larval  stage,  marine  molluscs  are  quite  a 

12 


178         HISTORY  OF   THE   EUROPEAN    FAUNA. 

prey  to  the  currents  of  the  sea.  They  have  practically 
no  swimming  organs,  and  only  move  by  lashing  to  and 
fro  the  tender  cilia  with  which  they  are  provided. 

This  disposes,  therefore,  of  Professor  Credner's 
main  criticisms.  As  for  the  fauna  of  the  relict 
lakes,  we  are  now  only  concerned  with  those  of 
Northern  Russia,  Finland,  and  Sweden.  In  the 
lakes  Wetter  and  Wener  in  the  latter  country 
occurs  the  four-horned  sting-fish  (Coitus  quadricornisy 


FlG.  14.— The  Four-horned  Sting- fish  (Coitus  qitadricornis],  reduced 
from  Professor  Smitt's  figure  in  the  Fishes  of  Scandinavia. 

Fig.  14),  which,  as  we  have  learned,  also  inhabits  the 
northern  part  of  the  Baltic,  and,  as  was  suggested, 
migrated  there  at  a  time  when  the  latter  was 
connected  with  the  White  Sea.  The  principal  food 
of  this  little  fish  consists  in  a  marine  Crustacean 
called  Idotea  entomon,  an  animal  allied  to  our 
common  woodlouse.  This  is  a  typical  marine 
species,  but  it  occurs  also  in  the  relict  lakes  of 
the  countries  mentioned  above,  as  well  as  in  the 
Baltic  and  the  Caspian.  Perhaps  the  best  known 


THE  ARCTIC  FAUNA.  179 

form  with  a  similar  range  is  the  Schizopod  crus- 
tacean Mysis  relzcta1  (Fig.  15),  which  is  clearly  a  des- 
cendant of  the  Arctic  marine  Mysis  oculata>  of  which 
it  was  formerly  considered  a  mere  variety.  The  two 
Amphipods  Gammaracanthus  relictus  and  Pontoporeia 
ajjinis,  and  the  Copepod  Limnocalanus  macrurus,  are 
three  additional  well-known  Arctic  crustaceans  whose 
range  differs  but  little  from  those  above-mentioned.2 


FIG.   15. — Mysis  relicta,  a  small  shrimp-like  Crustacean,  after  Sars 
(enlarged). 

These  facts  all  go  to  prove  that  the  sea  formerly 
covered  the  lowlands  of  Sweden,  Finland,  and 
Northern  Russia.  The  fauna  of  Scandinavia,  as 
we  have  seen,  indicates  that  during  the  greater  part 
of  the  Glacial  period  the  country  was  not  directly 
connected  with  continental  Europe  as  it  is  now.  It 
seems  that  the  barrier  of  separation  probably  con- 

1  The  occurrence  of  this  species  in  Lough  Neagh  in  Ireland,  pointing 
to  a  connection  between  the  Irish  Sea  and  the  Baltic,  will  be  referred  to 
later  on;  as  also  that  of  two  allied  forms  in  the  Caspian  Sea. 

-  For  additional  species  with  a  similar  range,  vide  Nordquist. 


I  SO         HISTORY   OF   THE    EUROPEAN   FAUNA. 

sisted  of  a  broad  expanse  of  ocean  on  which  floated 
numerous  icebergs,  which  originated  from  the  Scan- 
dinavian glaciers  as  they  reached  the  sea.  This  was 
a  cold  sea,  whilst  Western  Scandinavia  was  washed 
by  the  Gulf  Stream  (vide  Fig.  12,  p.  156).  We 
might  look  upon  the  boulder-clay  which  covers  such 
vast  tracts  of  country  in  Northern  Germany,  Russia, 
and  Holland  as  deposits  formed  by  this  sea  rather 
than  the  ground-moraine  of  a  huge  Scandinavian 
glacier.  I  shall  refer  to  this  subject  again  in  the 
next  chapter ;  meanwhile  it  may  be  remembered 
that  the  boulder-clay  of  Northern  Europe  exactly 
resembles  in  all  important  particulars  the  similar 
accumulations  met  with  in  the  British  Islands. 
They  resemble  one  another  also  in  the  occasional 
occurrence  of  sea-shells,  the  frequent  appearance  of 
bedded  deposits,  and  the  often  inexplicable  course 
taken  by  boulders  from  their  source  of  origin.  There 
occurs  often  a  singular  mixture  and  an  apparent 
crossing  of  the  paths  of  boulders  in  the  boulder- 
clay.  Professor  Bonney  remarks  (p.  280)  that  these 
are  less  difficult  to  explain  on  the  hypothesis  of 
distribution  by  floating  ice  than  on  that  of  transport 
by  land-ice,  because,  in  the  former  case,  though  the 
drift  of  winds  and  currents  would  be  generally  in  one 
direction,  both  might  be  varied  at  particular  seasons. 
So  far  as  concerns  the  distribution  and  thickness  of 
the  glacial  deposits,  he  says  there  is  not  much 
to  choose  between  either  hypothesis;  but  on  that 
of  land-ice  it  is  extremely  difficult  to  explain  the 


THE  ARCTIC   FAUNA.  l8l 

intercalation  of  perfectly  stratified  sands  and  gravels 
and  of  boulder-clay,  as  well  as  the  not  infrequent 
signs  of  bedding  in  the  latter.  Two  divisions  are 
generally  recognisable  in  the  continental  boulder-clay 
• — a  lower  and  an  upper.  An  inter-glacial  phase 
characterised  by  a  less  severe  climate  is  assumed 
to  have  intervened  between  the  deposition  of  the 
two.  In  Russia  no  such  division  can  as  a  rule  be 
made  out,  and  sea-shells  are  either  entirely  absent  or 
extremely  scarce.  It  has  been  pointed  out  by  Pro- 
fessor J.  Geikie  that  the  erratics — a  name  applied  to 
boulders  in  boulder-clay — in  the  upper  division  have 
travelled  in  a  different  direction  from  those  contained 
in  the  lower.  Taking  for  granted  that  the  boulder- 
clay  is  a  marine  deposit,  this  phenomenon  seems 
to  indicate  that  the  current  which  prevahed  during 
the  early  part  of  the  Glacial  period  in  this  North 
European  ocean  was  different  from  the  prevailing 
current  during  the  latter  part.  I  have  attempted 
to  explain  this  circumstance  by  the  supposition 
that  during  the  early  part  of  the  Glacial  period 
the  Northern  Sea  had  a  connection  with  the  Ponto- 
Caspian  Sea — a  sea  formed  by  the  junction  of  the 
Black  Sea  and  the  Caspian  (Fig.  12,  p.  156).  There 
is  geological  evidence,  as  will  be  explained  in  the 
following  chapter,  that  the  area  of  these  two  seas  was 
considerably  larger  in  glacial  times  than  it  is  now, 
and  that  they  were  joined  across  the  valley  of  the 
Manytch.  After  the  inter-glacial  phase  of  the  Glacial 
period,  the  North  European  Ocean  became  connected 


1 82         HISTORY  OF   THE   EUROPEAN    FAUNA. 

with  the  Atlantic  Ocean  across  the  north  of  England 
(Fig.  6,  p.  126),  the  junction  between  the  former 
and  the  Ponto-Caspian  having  meanwhile  become 
dry  land  (Fig.  13,  p.  170).  A  fresh  current,  now 
flowing  westward,  was  set  up  in  the  North  Euro- 
pean Ocean,  which  accounts  for  the  fact  just  cited 
that  the  erratics  in  the  upper  continental  boulder- 
clay  have  travelled  in  a  different  direction  from 
those  in  the  lower.  The  boulder-clay  laid  down  by 
the  sea  on  the  midland  and  northern  counties  of 
England,  just  as  was  the  case  with  the  similar  deposit 
on  the  Continent,  is  generally  accredited  to  the 
action  of  land-ice.  It  is  by  most  geologists  looked 
upon  as  the  ground-moraine,  partly  of  the  huge 
Scandinavian  glacier  which  is  supposed  to  have 
impinged  upon  the  English  coast,  partly  of  local 
British  glaciers. 

But  renewed  geological  investigations  on  this  point 
throw  doubts  upon  these  theories.  Thus  Mr.  Harmer 
remarks  in  a  recent  contribution  to  glacial  literature 
(p.  775),  that  "it  is  difficult  to  see  how  the  Baltic 
glacier  could  have  reached  East  Anglia,  though  ice- 
floes with  Scandinavian  boulders  might  easily  have 
done  so,  while  had  the  Norwegian  ice  filled  the  North 
Sea  and  overflowed  the  county  of  Norfolk,  some  evi- 
dence of  its  presence  ought  to  be  found  in  the  glacial 
beds  of  Holland." 

All  the  phenomena  of  distribution  of  the  British 
fauna  and  flora  are,  as  we  have  seen,  much  more  easily 
explained  by  the  supposition  of  a  damp,  temperate 


THE   ARCTIC   FAUNA.  I&3 

climate,  such  as  might  have  been  produced  by  the 
proximity  of  a  cold  sea  on  one  side  and  of  a  warm 
one  at  the  other,  than  by  invokirg  an  arctic  climate 
with  enormous  glaciers.  Most  of  the  living  animals 
and  plants  would  have  been  exterminated  under  the 
latter  conditions.  Palaeontological  evidence  in  Great 
Britain  clearly  indicates  that  southern  species  migrated 
first  to  these  islands,  that  Arctic  species  were  then 
driven  south  from  their  native  lands, — probably  owing 
to  insufficient  food-supply  and  climatic  changes  in 
the  north, — that  finally  eastern  species  invaded  the 
country — all  this  without  the  annual  temperature 
of  Europe  being  apparently  much  affected.  For  we 
find  in  the  British  pleistocene  deposits — and  Mr. 
Lydekker  draws  particular  attention  to  this  remark- 
able fact — a  curious  intermingling  of  southern  and 
northern  mammals,  which  undoubtedly  lived  side  by 
side.  Everybody  knows  that  northern  and  Arctic 
species  can  live  perfectly  well  in  a  temperate  climate, 
but  that  it  is  almost  impossible  to  acclimatise 
southern  animals  in  an  Arctic  or  even  temperate 
one.  We  have  in  this  circumstance  almost  a  proof, 
therefore,  that  the  climate  cannot  have  been  very  cold. 
Though  a  cold  sea  bathed  the  shores  of  Eastern 
England,  and  even  eventually  invaded  a  portion  of 
Northern  England,  the  warm  ocean  on  the  west 
must  have  effectually  prevented  any  great  lowering 
of  temperature. 

At  the  time  when  the  North  European  Sea  flooded 
a  portion  of  England,  Scandinavia  was  still  connected 


184         HISTORY  OF   THE   EUROPEAN   FAUNA. 

with  Scotland,  and  the   latter  with  Ireland  (Fig.   6, 
p.  126).    There  is  no  doubt  that  the  food-supply  in  the 
Arctic   Regions  was  decreasing  with  an  increase  of 
snowfall  and  with  the  gradual  lowering  of  the  land, 
which  reduced  also  the  habitable  area.    Arctic  species 
therefore  were  driven  south  in  search  of  fresh  pastures. 
But  it  need  not  be  supposed  that  anything  like  a  vast 
destruction  of  the  fauna  of  the  Arctic  Regions  took 
place.     Only  fewer  mammals  were  able  to  find  food 
in   a  given   space  than  heretofore.     This  southward 
migration    may   have    commenced,    in    the    case   of 
plants  and  the  invertebrates,  at  a  much  earlier  time, 
— during   the   Miocene   or  Pliocene  Epochs, — but  it 
is  doubtful  whether  the  mammals  and   birds  which 
we    find    in    our    pleistocene    and    recent    deposits 
began  to  travel   south  much  before  the  commence- 
ment of  the  Glacial  period.      The  beginning  of  the 
Glacial  period  in  England,  I  think,  is  indicated  by 
the  deposition  of  the  Red  Crag,  though  the  latter  is 
generally   regarded    as    belonging    to    the    pliocene 
series.      Much    of    the    northward    migration    from 
the   British   Islands  of  Lusitanian    and  other  forms 
had  then  ceased,  but  we  have  in  Scandinavia,  just 
as  in  these  islands,  a  southern  relict  fauna  and  flora, 
plants  and  animals  which  had  wandered  across  what 
is  now  the  German  Ocean   from  Scotland  to  Scan- 
dinavia,  and    have   never   become    extinct    in    that 
country  to  the  present  day.     I   need  only  mention 
the  Red  Deer,  the  Badger,  and  Slugs  of  the  genus 
A  rion. 


THE  ARCTIC   FAUNA.  185 

Professor  Blytt  directs  attention  to  some  such 
southern  relict  species  of  plants  now  only  found  in  the 
extreme  south-west  of  Scandinavia,  such  as  Asplenium 
niarinuni)  Hymenophyllum  Wilsoni,  Carex  binervis^ 
Scilla  verna,  Erica  cinerea,  Conopodimn  denudalum, 
Meum  athamanticum,  and  Rosa  involuta  (p.  28). 

The  Arctic  fauna  and  flora  in  Scandinavia — that 
is  to  say,  the  descendants  of  those  species  which 
migrated  direct  from  Greenland  and  Spitsbergen, 
as  we  have  seen,  are  numerous.  They  of  course  per- 
sisted throughout  the  Glacial  period  in  the  country, 
and  are  now  in  many  localities  being  exterminated 
partly  by  change  of  climate,  partly  by  a  keen  com- 
petition with  more  vigorous  rivals  which  have  come 
to  Scandinavia  from  the  east.  It  is  a  curious  circum- 
stance, as  pointed  out  by  Professor  Blytt,  that  the 
Arctic  plants  in  the  Botanic  Gardens  at  Christiania 
are  able  to  stand  almost  any  amount  of  sunshine, 
but  are  very  liable  to  be  injured  by  the  frost,  and  have 
to  be  covered  in  the  winter.  A  similar  observation 
has  been  made  in  the  case  of  the  Alpine  plants  at 
Kew  Gardens,  which  have  to  be  wintered  in  frames, 
though  their  homes  are  either  in  the  high  Alps — among 
the  everlasting  snows — or  in  the  intensely  cold  climate 
of  Greenland.  Many  of  the  Scandinavian  plants  ex- 
hibit instances  of  discontinuous  distribution,  thus  show- 
ing their  ancient  origin ;  and  there  is  altogether  nothing 
in  the  fauna  and  flora  of  that  country  which  might 
lead  us  to  believe  that  these  were  exterminated 
during  the  Glacial  period  and  reintroduced  subse- 


1 86         HISTORY   OF  THE   EUROPEAN    FAUNA. 

qucntly.  The  climate  during  that  period  in  Scan- 
dinavia was  probably  more  equable  and  moister, — 
with  a  greater  snowfall  in  winter  and  with  less 
sun  to  melt  the  snow  during  summer, — so  that  the 
development  of  glaciers  took  more  formidable  dimen- 
sions, chiefly  on  the  east  side.  The  lowlands  of 
Sweden  were  covered  by  the  sea,  whilst  many  of 
the  valleys  were  choked  with  ^  great  glaciers,  which 
cast  off  portions  of  ice  as  they  reached  the  sea,  just 
as  the  Greenland  and  other  northern  glaciers  do  {vide 
p.  237).  A  country  which  at  the  present  day  probably 
somewhat  resembles  the  former  Scandinavia  climati- 
cally is  Tierra  del  Fuego,  in  the  extreme  south  of 
South  America.  Though  there  is  an  abundant 
snowfall,  so  that  glaciers  reach  the  sea  in  many 
parts  of  the  country,  the  flora  has  been  described  by 
travellers  as  luxuriant;  and  it  appears  that  the  fauna 
also  is  richer  than  might  be  expected  from  the  cheer- 
less climate. 

Towards  the  latter  part  of  the  Glacial  period  the 
land-connection  between  Scandinavia,  Spitsbergen, 
and  Greenland  broke  down,  and  the  waters  of  the 
Arctic  and  Atlantic  Oceans  joined.  Whether  it  was 
at  this  time  or  later  that  the  other  land-connection 
between  Scandinavia  and  Scotland  collapsed  is 
difficult  to  determine;  but  it  is  certain,  I  think, 
that  Scotland  was  still  united  with  Ireland  even  after 
these  two  great  land-bridges  ceased  to  exist. 


•y 

THE   ARCTIC   FAUNAV.       O*J£^     1 87 


SUMMARY  OF  CHAPTER  IV. 

The  fauna  of  the  Arctic  Regions  is  much  poorer  than  that  of  the 
other  regions  which  are  dealt  with  in  this  work.  In  some  groups, 
such  as  Reptiles  and  Amphibia,  there  are  no  representatives  at  all, 
but  no  doubt  a  larger  number  of  species  existed  there  in  earlier 
Tertiary  times.  At  least  we  have  fossil  evidence  that  during 
the  Miocene  Epoch  plants  of  many  families  flourished  in  Green- 
land of  which  no  vestige  is  now  left  in  the  Polar  area.  Climatic 
conditions  must  therefore  have  changed,  as  in  Europe.  A 
gradual  refrigeration  took  place,  owing  probably  to  the  slow 
withdrawal  of  the  current  which  supplied  the  Arctic  Sea  with 
warmth.  Greenland  and  Europe  were  then  connected,  and 
the  Arctic  Ocean  was  separated  from  the  Atlantic.  This  land- 
connection  is  supposed  to  have  lain  far  north  between  Scan- 
dinavia, Spitsbergen,  and  Greenland,  and  must  have  persisted 
until  towards  the  end  of  the  Glacial  period. 

As  the  temperature  decreased  and  the  land-area  available  in 
the  north  diminished,  the  surplus  population,  consisting  of 
animals  and  plants,  and  possibly  also  of  human  beings,  moved 
southward.  We  have  traces  in  Europe,  and  especially  in  the 
British  Islands,  of  a  very  early  migration  from  the  north  in  the 
so-called  American  plants  and  in  the  freshwater  sponges.  The 
geographical  distribution  of  some  of  the  Arctic  species  of  mam- 
mals is  referred  to  in  greater  detail,  to  show  how  the  relative 
age  of  their  entry  into  Europe  can  be  determined.  Two  forms 
of  Reindeer,  resembling  the  Barren-ground  and  Woodland 
varieties,  have  been  met  with  in  European  deposits,  but  only 
the  former  occurs  in  Ireland  and  the  south  of  France,  whilst 
eastward  the  other  becomes  more  common,  and  finally  is  the 
only  one  found.  It  is  believed  that  the  Barren-ground  is  the 
older  form  as  far  as  Europe  is  concerned,  and  that  it  came  to 
us  with  the  Arctic  migration,  and  that  the  other  Reindeer 
reached  Europe  much  later  from  Siberia,  when  Ireland  had 
already  become  detached  from  England.  The  range  of  the 


1 88         HISTORY  OF  THE   EUROPEAN   FAUNA. 

Arctic  Hare  is  equally  instructive.  It  must  have  been  a  native 
of  Europe  since  early  glacial  or  pre-glacial  times— before  the 
common  English  Hare  had  made  its  appearance  in  Central 
Europe.  Along  with  other  Arctic  forms,  it  entered  Northern 
Europe  directly  from  the  Arctic  Regions,  by  means  of  the  former 
land-connection  which  joined,  as  I  remarked,  Lapland  with 
Spitsbergen,  Greenland,  and  North  America.  There  need  not 
have  been  a  post-glacial  connection  between  Europe  and 
Greenland ;  the  present  flora  of  that  country  may  have  survived 
the  Glacial  period  in  the  Arctic  Regions,  as  has  been  main- 
tained by  some  botanists  and  other  authorities.  Professor 
Forbes  argued  from  the  occurrence  of  the  same  species  of 
shore  mollusca  on  the  coast  of  Finmark  and  Greenland  that 
these  two  countries  were  not  long  ago  joined,  so  that  a  slow 
migration  from  west  to  east  along  an  ancient  coast-line  could 
have  taken  place.  That  such  a  migration  actually  occurred  is 
further  made  probable,  judging  from  the  presence  of  American 
mollusca  in  the  Crag  deposits  on  the  east  coast  of  England. 
These  came  into  the  North  Sea  in  the  first  place  direct  from  the 
Arctic  Ocean  at  a  time  when  the  two  oceans  freely  communi- 
cated with  one  another  across  the  lowlands  of  Northern  Russia, 
Northern  Germany,  and  Holland.  Arctic  shells  are  also  found 
below  the  boulder-clay  on  the  Baltic  coast,  and  a  free  com- 
munication such  as  indicated  is  generally  held  to  have  taken 
place  at  no  very  distant  date.  The  so-called  "relict  species" 
— marine  animals  left  in  freshwater  lakes  in  districts  formerly 
covered  by  this  sea— lend  some  support  to  this  view.  But  the 
view  that  the  continental  boulder-clay  is  a  marine  deposit  is  not 
now  held  except  by  a  few,  though  I  here  bring  it  forward  again, 
as  it  seems  to  me  to  fit  in  so  much  better  with  the  known  facts 
of  distribution.  The  sea  just  referred  to  probably  existed 
throughout  the  greater  part  of  the  Glacial  period;  and  icebergs, 
which  originated  from  the  Scandinavian  glaciers,  would  have 
brought  detritus  and  boulders  to  the  lowlands.  Scandinavia 
was  then  connected  with  Scotland,  and  England  with  France. 


CHAPTER  V. 

THE   SIBERIAN    MIGRATION. 

IN  dealing  with  the  British  fauna  in  particular,  I 
have  drawn  attention  to  the  fact  that  it  is  chiefly  in 
the  south  of  England  that  we  find  fossil  remains 
of  eastern  species  of  mammals  in  recent  geological 
deposits.  We  can  actually  trace  the  remains  of 
these  species  and  their  course  of  migration  across 
part  of  the  Continent  towards  Eastern  Europe,  and 
as  none  of  their  bones  have  been  discovered  in  the 
southern  or  northern  parts  of  our  Continent,  it  must 
be  assumed  that  their  home  lay  in  Siberia,  where 
many  still  exist  to  the  present  day,  and  where 
closely  allied  forms  also  are  found.  Some  of  these 
Siberian  migrants  have  remained  in  England  and 
on  the  Continent  to  the  present  day.  Many  have 
become  extinct.  But  the  animals  forming  this 
eastern  migration  did  not  all  originate  in  Siberia, 
though  I  have  sometimes  spoken  of  them  collectively 
as  Siberian  migrants.  There  must  have  been  other 
centres  of  dispersion  of  species  in  Europe.  We  know 
that  a  very  active  centre  of  development — at  any  rate 
for  land-mollusca — lay  in  South-eastern  Europe,  either 
in  the  Caucasus  or  in  the  Balkan  peninsula,  or  more 


HISTORY   OF   THE   EUROPEAN    FAUNA. 

probably  in  both.  The  Alps  no  doubt  produced  a 
number  of  species  which  have  spread  north  and 
south,  and  may  in  their  wanderings  have  joined  the 
Siberian  migrants  in  their  western  course,  and  thus 
have  reached  the  British  Islands.  Nevertheless,  the 
majority  of  the  mammals  belonging  to  the  eastern 
clement  of  the  British  fauna  (vide  p.  95)  have  un- 
doubtedly originated  in  Siberia.  The  Polecat  (Mus- 
tela  putorius}  and  the  Harvest  Mouse  (Mus  minutus], 
for  instance,  are  members  of  that  eastern  migration. 
Both  occur  throughout  Central  Europe  and  a  large 
portion  of  Siberia,  but  are  absent  from  the  extreme 
north  and  south  of  Europe  and  also  from  all  the 
Mediterranean  Islands.  A  Siberian  species,  which 
has  never  penetrated  so  far  west  as  the  British 
Islands,  nor  even  so  far  north  as  Scandinavia  or 
south  to  Italy,  is  what  is  known  in  Germany  as  the 
"Hamster"  (Cricetus  frumentarius],  a  little  Rodent 
which  spends  the  winter  asleep  in  its  burrows,  and 
surrounds  itself  with  a  great  accumulation  of  food- 
material  carried  there  during  autumn.  The  common 
English  Hare,  which  I  formerly  regarded  as  an 
instance  of  a  Siberian  mammal,  must  now  find  a 
place  among  the  Oriental  migrants.  Its  history  is 
very  instructive,  and  I  shall  have  an  opportunity  later 
on  to  refer  to  it  again.  Meanwhile,  it  may  be  men- 
tioned that  though  this  Hare  inhabits  Europe  in  two 
varieties  or  races,  one  of  which,  Lepus  mediterra- 
neus,  is  confined  to  Southern  Europe,  the  latter  owes 
its  origin  to  an  earlier  migration  from  Asia. 


THE   SIBERIAN    MIGRATION.  IQI 

When  we  come  to  consider  the  eastern  birds,  we 
have  to  distinguish  between  resident  species  and 
migratory  ones.  The  Black-throated  Thrush  (Turdus 
atrigularis],  which  has  been  twice  obtained  in  the 
British  Islands,  is  a  mere  straggler  to  Europe,  and 
is  not  known  to  breed  there  at  all.  Better  known 
birds,  perhaps,  are  the  Golden  Thrush  (  Turdus  varius\ 
which  has  even  occurred  as  far  west  as  Ireland,  the 
Rock-Thrush  (Monticola  saxatilis)  and  the  Scarlet 
Grosbeak  {Carpodacus  erythrinus],  which  breed  in 
Eastern  Europe,  but  are  known  only  as  occasional 
visitors  in  the  west. 

To  judge  by  their  distribution,  the  Bullfinches 
(Pyrrhula)  are  of  Asiatic  origin,  for  seven  species  out 
of  ten  are  confined  to  that  continent.  Our  common 
Bullfinch  (P.  europed]  probably  came  with  the  Oriental 
migrants,  or  perhaps  its  ancestors  did.  But  the  larger 
Northern  or  Russian  Bullfinch  (P.  major]  has  no  doubt 
entered  our  Continent  directly  from  the  east.  We  have 
in  many  groups  similar  instances  of  closely  allied  species 
er  varieties,  one  of  which,  originating  at  a  somewhat 
later  stage  than  the  other,  took  a  different  route  of 
migration  from  that  followed  by  its  near  relative. 

The  Pine -Grosbeak  (Pimcola  enudeator)  is  only 
known  to  British  ornithologists  as  an  exceedingly 
rare  visitor.  Its  real  home  lies  in  the  northern  parts 
of  Europe,  Asia,  and  North  America,  and  it  is  one 
of  the  most  typical  of  the  Siberian  migrants. 

But  there  are  a  number  of  other  species  of  birds, 
which,  though  probably  not  of  Siberian  origin, 


IQ2          HISTORY  OF  THE   EUROPEAN   FAUNA. 

only  migrated  westward  recently,  and  have  either 
not  yet  reached  the  British  Islands,  or  which  lead  one 
to  suppose,  from  their  British  range,  that  they  are 
eastern  forms. 

Such,  for  instance,  is  the  Nightingale  (Danlias 
luscinia\  which  is  probably  of  Oriental  origin,  but 
only  visits  England  regularly  in  spring.  There  is  no 
authenticated  record  of  its  ever  having  migrated 
either  to  Scotland  or  Ireland. 

The  Bearded  Titmouse  (Panurus  biarmicus)  is  one 
of  the  eastern  birds  still  resident  in  England,  though 
unfortunately  it  seems  tcr  be  on  the  verge  of  extinc- 
tion. It  is  unknown  in  Scotland  and  Ireland. 
Another  resident  eastern  species  is  the  Nuthatch 
(Sitta  ca>sia\  but  neither  of  these  is  probably  of 
Siberian  origin. 

The  majority  of  the  European  Reptiles  are  probably 
of  eastern  origin.  Among  our  British  species,  the 
Common  Viper  (Pelias  berus\  for  example,  is  a 
typically  eastern  form.  It  is  almost  unknown  in 
Southern  Europe  proper — that  is  to  say,  in  Italy, 
the  Balkan  peninsula,  and  the  Mediterranean  Islands, 
but  its  range  extends  in  the  west  as  far  as  Spain,  and 
in  the  east  right  across  the  Asiatic  continent  to 
Japan.  It  is  well  known  that  the  Viper  occurs  in 
Scotland,  and  that  neither  it  nor  any  other  snake  is 
found  in  Ireland.  There  is  a  legend,  indeed,  that 
snakes  did  once  exist  in  Ireland  and  were  banished 
from  the  island  by  St.  Patrick,  but  unfortunately 
we  have  no  historical  evidence  that  such  an 


THE   SIBERIAN   MIGRATION.  IQ3 

interesting  event  actually  took  place.  The  Sand- 
Lizard  (Lacerta  agilis\  another  British  species,  may 
be  looked  upon  as  an  eastern  form.  It  is  quite  absent 
from  Italy,  the  Balkan  peninsula,  and  the  Medi- 
terranean Islands,  but  extends  throughout  Central 
Europe  to  the  east. 

Among  the  species  of  eastern  Reptiles  which  have 
a  mere  local  range  in  Europe  might  be  mentioned 
the  two  Lizards,  Phrynocephalus  auritus  and  Agarna 
sanguinolenta.  They  belong  to  the  family  Iguanidcs^ 
which  includes  some  very  large  species.  Both  of 
them  are  Asiatic  forms,  which  have  only  just  pene- 
trated across  the  eastern  steppes  into  Europe,  where 
they  inhabit  the  arid  regions  between  the  Caspian 
and  the  River  Don  in  Southern  Russia. 

The  species  of  Mammals  living  in  Europe  at  the 
present  day  have,  with  few  exceptions,  migrated 
to  our  continent  from  other  parts  of  the  world. 
With  regard  to  the  Birds,  it  is  possible  that  a 
somewhat  larger  number  proportionally  may  be 
of  European  origin.  Still,  the  great  majority  are, 
I  think,  to  be  regarded  as  immigrants.  The  autoch- 
thones are  about  equal  to  the  immigrant  reptiles, 
but  many  of  the  European  Amphibians  and  the 
majority  of  the  Fishes  have  probably  originated  on 
our  continent.  Some  of  the  European  Amphibia — 
especially  among  the  tailless  forms — appear  to  be 
immigrants  from  Asia.  Thus  the  distribution  of 
Rana  arvalis  in  Europe  is  remarkably  like  that  of 
a  Siberian  migrant.  This  frog  occurs  in  Siberia, 

13 


194         HISTORY   OF   THE   EUROPEAN    FAUNA. 

ranging  southward  as  far  as  Persia  and  parts  of 
Asia  Minor.  Crossing  the  European  border,  we 
find  it  in  Russia,  Upper  Hungary,  North  and  Central 
Germany, — being  rarer  in  the  south, — Denmark,  and 
Scandinavia.  According  to  Bedriaga,  it  crosses  the 
Rhine  only  in  Alsace,  but  occurs  no  farther  west.  It 
only  just  enters  Holland.  If  we  suppose  the  species 
to  have  originated  in  Central  Europe,  we  should 
expect  to  find  it  in  Switzerland,  France,  and  perhaps 
England.  If  it  had  its  ancestral  home  in  Eastern 
Europe,  we  might  expect  it  to  occur  on  the  Balkan 
peninsula.  It  seems  to  me  more  probable,  therefore, 
that  Rana  arvalis  came  with  the  Siberian  migration. 
This  need  not  cause  surprise,  as  the  genus  Rana  is 
certainly  not  European.  Out  of  about  no  species, 
only  four  are  peculiar  to  Europe,  the  rest  are  scattered 
over  all  parts  of  the  globe.  Moreover,  the  fact  that 
these  four  species  are  confined  to  Southern  Europe 
would  seem  to  indicate  that  the  first  species  entered 
from  the  south,  and  there  either  became  modified  or 
spread  over  nearly  the  whole  continent,  as  did,  for 
instance,  Rana  esculenta  and  R.  temporaries  Neither 
of  these  is  by  any  means  confined  to  Europe.  R. 
esculenta  ranges  right  across  the  Asiatic  continent  to 
Japan,  and  also  enters  North  Africa,  while  the  other 
has  a  wide  distribution  in  northern  and  temperate 
Asia. 

The  various  groups  of  Vertebrates  are  not  dependent 
on  each  other  in  their  migrations.  Mammals  and  Birds 
extend  their  range  with  so  much  greater  facility  than 


THE  SIBERIAN    MIGRATION.  195 

Reptiles  and  Amphibians,  that  the  surplus  population 
of  our  neighbouring  continents  readily  poured  into 
Europe  when — owing  to  changes  of  climate  perhaps — 
they  forsook  their  original  homes. 

We  observe  much  the  same  differences  of  origin  in 
the  various  groups  of  European  Invertebrates.  The 
Central  European  Molluscan  fauna,  remarks  Dr. 
Kobelt,  had  already  developed  from  the  pliocene 
— in  almost  all  its  details,  as  regards  formation  of 
species  and  distribution — when  the  Ice-Age  com- 
menced (3,  i.  p.  162).  Certain  very  interesting  disloca- 
tions, however,  in  the  range  of  land  mollusca  can  be 
proved  to  have  taken  place  about  that  time.  Thus, 
as  Dr.  Kobelt  has  pointed  out,  the  genus  Zonites, 
which  is  now  almost  confined  to  the  south-east  of 
Europe,  occurs  in  inter-glacial  deposits  in  the  valley 
of  the  Neckar,  and  even  as  far  west  as  the  Seine.  If 
we  might  judge  from  this  single  instance,  a  molluscan 
migration  from  the  east  to  the  west  seems  to  have 
occurred  either  in  early  or  pre-glacial  times.  That 
Helix  pomatia  has  migrated  only  comparatively 
recently  from  the  East  to  Western  Europe  is 
rendered  probable  by  its  general  range  in  northern 
and  western  Europe,  but  I  cannot  agree  with  Dr. 
Kobelt  in  the  belief  that  Helix  aspersa  is  of  an 
equally  recent  origin  in  the  North.  No  matter 
whether  it  has  been  found  fossil  or  no,  its  range  in 
the  British  Islands  points  to  its  having  penetrated  to 
Ireland  when  the  latter  was  still  connected  with  the 
Continent  by  way  of  England.  Its  migration  from 


196         HISTORY  OF  THE   EUROPEAN   FAUNA. 

the  Mediterranean  dates  therefore  from  early  pleisto- 
cene or  late  pliocene  times. 

In  referring  to  the  sixty-five  species  of  Land  and 
Freshwater  Mollusca  which  have  been  described  from 
the  continental  "Loess,"  Dr.  Kobelt  states  (p.  166) 
that  this  fauna  has  certainly  not  a  steppe-character. 
It  does  not  therefore  strengthen  Professor  Nehring's 
view  that  Europe  during  the  deposition  of  the  loess 
had  a  climate  comparable  to  that  of  the  Siberian 
steppes.  The  Glacial  period  had  hardly  any  effect 
on  the  molluscan  fauna  of  Europe.  Dr.  Kobelt 
believes  in  a  certain  movement  of  that  fauna  from 
the  least  favourable  areas,  with  a  subsequent 
re-immigration;  but  even  that  could  not  have 
taken  place  on  a  large  scale.  Nothing  like  a 
destruction  of  the  fauna  occurred,  as  far  as  we 
know  from  fossil  evidence. 

Not  a  single  species  of  land  or  freshwater  mollusc 
can  be  quoted  as  having  migrated  to  Europe  from 
Siberia  in  recent  geological  times.  The  molluscan 
fauna  of  the  latter  country  is  so  closely  connected 
with  that  of  Europe,  that  it  is  quite  impossible  to 
elevate  it  to  the  rank  of  a  sub-region  of  the  Holarctic 
Region.  Dr.  Kobelt  insists  that  Siberia  cannot  even 
claim  to  be  placed  into  a  distinct  province.  Accord- 
ing to  the  same  authority,  we  find  no  species  in  the 
whole  Siberian  molluscan  fauna  which  we  might 
regard  as  having  immigrated  since  the  close  of  the 
Glacial  period.  Even  to  attempt  the  location  of 
the  original  homes  of  many  of  the  species  which 


THE  SIBERIAN    MIGRATION.  197 

Siberia  has  in  common  with  Europe,  seems  hopeless. 
Such  forms  as  Arion  hortensis,  which  has  been 
obtained  in  Siberia,  and  which,  as  we  have  seen,  must 
have  originated  in  Western  Europe,  migrated  in 
pliocene  or  miocene  times,  possibly  along  the 
shores  of  the  Mediterranean  and  across  Asia  Minor. 
We  have  evidence,  therefore,  of  an  eastward  migration 
among  the  land  and  freshwater  mollusca  in  later 
Tertiary  times,  but  not  of  a  westward  one  from 
Siberia. 

A  very  different  view  is  presented  to  us  by  the 
coleopterous  fauna  of  Europe.  Many  of  our  Euro- 
pean Beetles  are  Siberian  migrants.  Let  us  take, 
for  instance,  the  Tiger  Beetles  (Cicindelidce).  There 
are  over  forty  species  of  the  genus  Cicindela  in 
Europe,  five  of  which  reach  the  British  Islands. 
This  seems  a  large  number;  but  there  are  altogether 
no  less  than  6co  species  of  the  genus  scattered  over 
the  greater  part  of  the  world,  many  of  them  being 
Asiatic.  The  genus  is  certainly  not  of  European 
origin,  for  not  only  are  most  of  the  European  species 
confined  to  the  Caucasus  and  the  south-east  generally, 
but  no  Cicindelidce  whatsoever  occur,  for  example,  in 
Madeira  or  the  Canaries,  where  we  should  expect 
some  to  have  persisted  if  the  genus  had  originated 
on  our  continent.  Moreover,  of  the  five  tribes  into 
which  the  large  family  of  Cicindelidce  can  be  sub- 
divided, only  two  range  to  Europe,  and  one  of 
them  is  represented  by  only  a  single  species  on  our 
continent. 


198         HISTORY  OF   THE   EUROPEAN    FAUNA. 

Some  of  the  Cicindclas  may  have  come  with  the 
Oriental  migration.  I  think  this  was  the  case  with 
the  only  Irish  species  of  the  genus,  C.  campestris.  It 
occurs  all  over  continental  Europe  and  Northern  Asia, 
and  varieties  of  the  species  are  known  from  Corsica, 
Sicily,  Crete,  the  Cyclades,  Sardinia,  Asia  Minor, 
Greece,  and  Spain.  Five  species  of  Cicindela^  as  I 
said,  are  known  from  England,  of  which  C.  silvatica 
and  C.  maritima  are  certainly  Siberian  migrants,  and 
perhaps  C.  hybrida  too.  Neither  of  the  two  first 
species  is  found  in  Southern  Europe  or  in  Spain, 
where  we  should  expect  them  to  occur  had  they 
originated  on  our  continent.  C-  silvatica  and 
maritima  have  no  doubt  entered  Europe  from 
Siberia  in  recent  geological  times,  probably  soon 
after  a  way  was  opened  up  across  the  Tchornosjem 
district  of  Southern  Russia — that  is  to  say,  in 
inter-glacial  times.  The  former  spread  along  the 
Central  European  plain  as  far  west  as  the  south-east 
of  England  when  Great  Britain  still  formed  part  of 
France.  C.  maritimay  which  preferred  the  proximity 
of  the  sea,  migrated  along  the  shores  of  the  Caspian 
and  then  across  Russia  to  the  shores  of  the  Baltic 
and  North  Sea,  and  has  penetrated  a  little  farther 
north  and  west  in  England  than  its  near  relative. 
C.  litterata  has  a  very  similar  distribution  and  origin, 
but  instead  of  wandering  so  far  west  as  the  British 
Islands,  it  seems  to  have  preferred  extending  its 
range  southward,  and  has  just  reached  Northern 
Italy. 


THE  SIBERIAN    MIGRATION.  1 99 

The  closely  allied  Ground-beetles  (Carabida)  furnish 
us  with  equally  interesting  and  instructive  proofs  of  a 
migration  from  Asia.  Over  300  species  of  Carabus 
are  known  to  science.  The  number  of  species 
inhabiting  Asia  and  Europe  are  about  equal.  But 
the  genus  does  not  extend  its  range  to  Southern 
Asia  or  to  South  America  or  Australia.  Very 
few  species  enter  Africa,  and  only  nine  North 
America,  of  which  three  also  occur  in  Siberia.  The 
genus  is  unknown  in  Madeira,  and  only  represented 
by  three  species  in  the  Canary  Islands.  To  judge 
from  its  distribution,  it  has  probably  originated  in 
Western  Asia.  Probably  some  Carabi  of  European 
origin  have  spread  into  Asia,  but  the  Asiatic — or 
we  might  say  the  Siberian — origin  and  subsequent 
migration  westward  of  a  number  of  well-known  forms 
appears  to  me  evident.  Such  forms  as  C.  clatkratus, 
C.  granulatus,  and  C.  cancellatus  are  no  doubt  of 
European  origin,  and  have  only  in  recent  geological 
times  extended  their  range  across  Northern  Asia, 
whilst  C.  marginalist  coming  fiom  Siberia,  can  hardly 
be  said  to  have  invaded  Europe,  since  it  has  never 
been  met  with  farther  west  than  the  eastern  provinces 
of  Prussia. 

Among  the  Carabidtz  there  are  altogether  very 
many  examples  pointing  to  a  migration  from  Asia  to 
Europe,  but  I  do  not  wish  here  to  give  a  list  of  all 
such  cases,  and  only  refer  to  a  few  of  the  more, 
remarkable  ones.  One  of  the  European  species  of 
Demetrias  (D.  unipunctatus),  known  to  English  ento- 


200         HISTORY  OF   THE   EUROPEAN   FAUNA. 

mologists  as  a  south-eastern  form,  seems  to  have 
arrived  with  the  Siberian  migration,  whilst  the  closely 
allied  D.  atricapillus,  which  has  been  able  to  reach 
Ireland,  has  a  wider  range  and  came  earlier  with  the 
Orientals. 

Messrs.  Speyer  state  (p.  68)  that  almost  all  those 
species  of  Central  European  Butterflies  whose 
northern  limit  is  deflected  southward  as  we  approach 
the  west  coast  of  Europe,  inhabit  also  the  Volga 
country  and  the  adjoining  parts  of  Asia.  Many  of 
them  are  much  commoner  there  than  in  Central 
Europe,  and  it  appears  probable  to  the  authors  of 
the  Geographical  Distribution  of  Butterflies  that 
these  species  came  from  the  east.  Asia  and  Central 
Europe  have,  according  to  Messrs.  Speyer,  no  fewer 
than  156  species  in  common.  Mr.  Petersen  esti- 
mates that  no  less  than  91  per  cent,  of  the  Arctic- 
European  Butterflies  also  occur  in  Siberia.  He 
made  a  special  study  of  the  Arctic  Macro-lepi- 
doptera,  and  came  to  the  conclusion  that  Central 
Asia,  not  having  been  glaciated  in  the  Ice-Age, 
offered  a  possibility  of  existence  to  both  animals 
and  plants.  Here,  he  thinks,  was  the  principal  centre 
to  which  Europe  owed  its  re-population  in  post-glacial 
times.  Mr.  Petersen  is  of  opinion  (p.  40)  that  the 
Arctic-European  Lepidoptera  are  composed  of  two 
elements  —  the  pliocene  relics  which  persisted  in 
Europe  during  the  Glacial  period,  and  the  new 
immigrants  from  Siberia. 

No  doubt  Siberia  supplied  Europe  with  a  number 


THE   SIBERIAN    MIGRATION.  2OI 

of  species  of  Butterflies  and  Moths  in  recent  geo- 
logical times,  but  we  need  not  necessarily  suppose 
that  these  arrived  only  after  the  Glacial  period.  Even 
the  most  extreme  glacialists  admit  that  large  areas 
on  our  continent  were  free  from  ice  at  the  height  of 
the  Ice-Age,  Siberia  had  therefore  no  particular 
advantage  over  Europe  in  giving  an  asylum  to 
Butterflies  and  Moths  which  were  escaping  from  the 
rigours  of  a  supposed  arctic  climate.  But  we  have 
already  learned  (p.  80)  that  the  climate  during  the 
Glacial  period  probably  differed  but  little  from  that 
which  we  enjoy  at  the  present  day,  and  we  may 
assume,  therefore,  that  the  Lepidoptera  of  Siberia 
migrated  during  that  time  or  even  earlier  to 
Europe. 

Let  us  for  a  moment  reconsider  some  instances  of 
mammalian  migration  from  Siberia,  with  a  view  to 
studying  more  closely  the  nature  of  these  great 
events.  I  mentioned  the  fact  that  some  of  the 
Siberian  migrants  have  remained  in  England,  that 
more  have  settled  down  permanently  on  our  con- 
tinent, but  that  many  others  have  either  become 
entirely  extinct  or  do  not  live  any  longer  in 
Europe. 

Of  the  mammals  which  made  their  appearance  in 
Great  Britain  in  recent  geological  times,  t.e.y  during 
and  since  the  deposition  of  the  Forest-Bed  for  example, 
the  following  species  probably  came  direct  from 
Siberia  across  the  plains  of  Europe,  as  already  men- 
tioned (p.  95): — 


2O2         HISTORY  OF   THE   EUROPEAN   FAUNA. 

Canis  lagopus.  *  Mus  minutus. 

Gulo  luscus.  *  Arvicola  agrestis. 

*  Mustela  erminea.  *        „       amphibius. 

*  „        putorius.  „       arvalis. 

*  „        vulgaris.  *        „       glareolus. 

*  Sorex  vulgaris.  „      gregalis. 
Lagomys  pusillus.                                   „       ratticeps. 

*  Castor  fiber.  Equus  caballus. 
Spermophilus  Eversmanni.  Saiga  tartarica. 

„           erythrogenoides.  Ovibos  moschatus. 

Cricetus  songarus.  Alces  latifrons. 

Myodes  lemmus.  ,,      machlis. 

Cuniculus  torquatus.  Rangifer  tarandus. 

*  Those  marked  with  an  asterisk  still  inhabit  Great  Britain,  or  did  so 
within  historic  times. 

Of  the  arrival  of  many  of  these  in  Europe  we  have 
geological  proof,  as  they  have  left  their  bones  in 
recent  pleistocene  deposits,  and  are  unknown  from 
older  European  strata.  The  remote  ancestors  of 
others,  such  as  Sorex  and  Lagomys,  no  doubt  lived  in 
Europe,  but  the  recent  species  probably  had  their 
original  homes  in  Asia.  It  is  evident  that  in  recent 
geological  times  there  existed  no  active  centre  of 
origin  for  mammals  in  Europe,  and  that  our  continent 
was  largely  dependent  on  the  neighbouring  one  for 
the  supply  of  its  mammalian  fauna.  A  shifting  of  the 
centre  of  development  from  Europe  to  Asia  appears 
to  have  taken  place  occasionally,  as  already  men- 
tioned (p.  45).  Mr.  Lydekker  has  drawn  attention 
to  the  fact  that  though  the  remote  ancestors  of  the 
Elephantidcz  resided  in  Europe,  neither  the  latter 


THE  SIBERIAN    MIGRATION.  203 

continent  nor  North  America  was  the  home  of  the 
direct  ancestor  of  any  of  the  true  Elephants. 
Similarly,  though  we  have  had  our  Sorex  in  Europe 
from  the  Upper  Eocene  and  Lagomys  from  the 
Middle  Miocene,  the  geographical  distribution  of 
Sorex  vidgaris  and  Lagomys  pusillus  does  not  support 
the  view  that  they  are  of  European  origin  and  have 
migrated  to  Asia.  Their  absence  from  most  of 
the  European  islands  indicates  either  an  extremely 
recent  origin  or  a  recent  immigration  from  Asia, 
and  the  latter  view  seems  to  me  much  the  more 
probable. 

No  less  than  twenty-six  species  of  the  Siberian 
mammals  penetrated  as  far  west  as  the  British  Islands, 
and  nine  of  these  still  inhabit  Great  Britain.  Some 
of  the  remaining  seventeen  species  probably  lived  only 
for  a  very  short  time  in  England,  and  the  rest 
gradually  became  extinct  one  by  one.  This  process 
of  extinction  of  the  aliens  still  continues.  The  Beaver 
(Castor fiber)  has  died  out  within  recent  historic  times. 
We  possess  legends  and  uncertain  historic  records 
pointing  to  the  existence  of  the  Reindeer  in  Scotland 
as  recently  as  about  seven  centuries  ago.  But  much 
the  same  state  of  things  has  happened  on  the  Con- 
tinent. The  Glutton  (Gulo  hiscus),  which  still  lived 
in  Northern  Germany  last  century,  has  now  entirely 
vanished  from  that  country,  as  also  the  Reindeer. 
The  Lemmings  have  found  an  asylum  in  Scandinavia. 
The  Musk-Ox  (Ovibos  moschatus)  has  disappeared 
not  only  from  Europe  but  also  from  Asia,  and  is  now 


204         HISTORY   OF   THE   EUROPEAN    FAUNA. 

confined  to  Arctic  America  and  Greenland.  The 
Horse  no  longer  occurs  in  Europe  in  the  wild  state, 
and  the  Saiga  Antelope  (Saiga  tartaricd)  has  retreated 
to  the  Steppes  of  Eastern  Europe  and  Western 
Siberia. 

As  we  proceed  more  and  more  eastward  across 
Central  Europe,  we  find  that  a  larger  and  larger 
percentage  of  the  Siberian  migrants  have  adopted 
the  new  country  as  their  permanent  home,  though  in 
France  and  Germany,  as  well  as  in  Austria,  we  have 
evidence  that  a  great  number  of  Siberian  species,  which 
formerly  lived  there,  have  either  become  entirely 
extinct,  or  have  retreated  towards  the  land  of  their 
origin.  There  is  a  prevalent  belief  that  these  migrants 
have  taken  refuge  on  the  higher  European  mountain 
ranges,  but  this  idea  is  altogether  erroneous,  as  will 
be  shown  in  the  chapter  dealing  with  the  origin  of  the 
Alpine  fauna. 

One  of  the  Jerboas  (Alactaga  jaculus)  occurs  fossil 
as  far  west  as  Western  Germany,  but  it  is  now  con- 
fined to  Russia  and  Western  Siberia.  The  Bobak 
marmot  (A rctoniys  bobak\  which  has  a  similar  range 
now,  probably  inhabited  France  in  former  times. 
A  Siberian  species  which  has  retreated  but  little  is 
the  Hamster  (Cricetus  vulgaris).  Its  fossil  remains 
have  been  found  in  Central  France,  but  it  does  not 
now  occur  west  of  the  Vosges  Mountains. 

It  appears,  therefore,  as  if  a  wave  of  migration  had 
swept  over  Central  Europe  from  east  to  west,  that 
those  species  which  were  able  to  adapt  themselves  to 


THE   SIBERIAN   MIGRATION.  20$ 

the  new  surroundings  had  remained,  and  as  if  the  rest 
had  died  out  or  were  gradually  retreating  to  the 
east. 

Ornithologists  are  well  acquainted  with  the  fact 
that  in  some  years  there  is  an  unusually  large  exodus 
from  Eastern  Europe  and  Siberia  of  birds;  and  that 
species  like  the  Waxwing  (Ampelis  garrulus)  then 
appear  in  great  numbers.  But  the  appearance  of  this 
bird  in  Western  Europe  is  not  looked  upon  as  so 
remarkable  as  that  of  Pallas's  Sandgrouse  (Syrrhaptes 
paradoxus,  Fig.  3,  p.  42),  a  typical  inhabitant  and  resi- 
dent of  the  Arctic  Steppes.  The  last  great  irruption 
took  place  in  1888,  and  many  birds  reached  even  the 
extreme  west  of  Ireland  in  May  and  June  of  that 
year.  A  few  weeks  before,  it  had  been  announced  to 
the  German  papers  that  large  flocks  of  this  peculiar 
pigeon-like  bird  had  arrived  in  the  eastern  provinces; 
and  though  the  vast  majority  vanished  as  quickly  as 
they  had  come,  a  certain  number  remained  for  a  year 
or  so  in  the  newly  visited  countries,  and  some  even 
bred  in  England. 

Twenty-five  years  before,  in  1863,  a  similar  migra- 
tion had  occurred,  though  not  perhaps  on  quite  such 
a  vast  scale,  and  a  few  small  flocks  had  made  their 
appearance  in  Western  Europe  on  several  occasions 
between  these  dates. 

It  may  not  be  generally  known  that  no  other  bird 
has  been  honoured  by  our  Government  in  a  like 
manner,  for  it  is  the  only  animal  for  whose  protec- 
tion a  separate  Act  of  Parliament  has  been  passed. 


2O6         HISTORY  OF   THE   EUROPEAN    FAUNA. 

In  spite  of  this  unusual  precaution,  the  species  has 
not  survived  to  add  another  member  to  the  resident 
British  fauna.  The  wave  of  migration  from  the  east 
has  come  and  vanished  again  just  like  so  many  others 
with  which  history  is  familiar. 

These  migrations  from  the  east  occurring  at  the 
present  day  give  us  some  idea  of  those  of  which  we 
have  fossil  evidence,  and  which  all  had  their  origin 
in  Central  and  Northern  Asia.  Almost  all  the  species 
of  mammals  to  which  I  have  referred  as  being  of 
Siberian  origin  have  been  found  in  the  fossil  state 
in  comparatively  recent  geological  deposits  within  a 
certain  very  limited  area.  None  of  the  typical  species 
have  ever  been  found  in  Southern  Europe  proper, 
including  the  Mediterranean  islands.  It  must  be 
remembered  that  though  the  Reindeer  is  a  Siberian 
migrant,  the  form  of  the  Reindeer  which  was  found 
in  the  Pyrenees  belonged  to  a  distinct  variety — in 
fact,  to  a  much  earlier  migration  which  issued  from 
the  Arctic  European  Regions,  and  to  which  I  have 
referred  in  detail  (pp.  150-158).  Curiously  enough, 
no  deposits  of  these  typical  Siberian  mammals 
have  ever  been  obtained  in  Scandinavia — only  in 
Russia,  Austria,  Switzerland  (the  lowlands),  Germany, 
Belgium,  France,  and  England.  To  facilitate  a  study 
of  the  extent  of  these  migrations,  I  have  constructed 
a  map  on  which  the  probable  course  taken  across 
Central  Europe  is  roughly  indicated  by  dots 
(Fig.  16). 

In  the  migrations  of  to-day  we  perceive  the  same 


THE   SIBERIAN    MIGRATION. 


207 


tendency  as  in  the  older  ones  of  which  we  have  fossil 
evidence,  viz.,  generally  a  spreading  of  species  on  a 
large  scale  over  new  territory,  and  then  a  gradual 


FIG.  1 6. —Map  of  Europe.  The  doited  portions  represent,  approxi- 
mately, the  course  of  migration  of  the  Siberian  mammals.  The 
principal  mountain  ranges  are  roughly  indicated  in  black. 

shrinkage  towards  their  original  home,  with  an  occa- 
sional survival  of  small  colonies  in  the  invaded  part. 
It  must  not  be  supposed  that  this  observation  applies 


208         HISTORY   OF   THE   EUROPEAN    FAUNA. 

alone  to  the  Siberian  migration.  In  the  case  of  the 
Arctic  one,  precisely  the  same  thing  has  happened, 
and  we  shall  see  that  the  Southern  (migration  from 
the  south)  agrees  in  this  respect  with  the  others. 

As  for  the  immediate  cause  of  these  migrations,  it  is 
to  be  looked  for  either  in  the  scarcity  of  food  dependent 
upon  a  temporary  or  permanent  change  of  climate,  or 
in  an  excessive  increase  in  numbers  of  a  particular 
species.  I  do  not  propose  to  trace  back  migrations 
beyond  the  Pliocene  Epoch,  or  indeed  much  beyond 
the  beginning  of  the  Glacial  period,  which  is  regarded 
as  a  phase  of  the  most  recent  geological  epoch,  viz., 
the  Pleistocene.  During  the  period  in  question,  we 
have  indirect  evidence  of  one  vast  migration  from 
Siberia  into  Europe  across  the  lowlands  lying  to  the 
north  of  the  Caspian  and  to  the  south  of  the  Ural 
Mountains.  There  is  a  general  consensus  of  opinion 
that  this  migration  took  place  in  Pleistocene  times. 
Professor  Nehring  thinks  that  there  can  be  no  doubt 
(p.  222)  that  the  Siberian  migrants  arrived  in 
Northern  Germany  after  the  first  stage  or  division 
of  the  Glacial  period,  and  lived  there  probably 
during  the  inter-glacial  phase  which  occurred  be- 
tween the  first  and  second  stages — if  indeed  we  look 
upon  this  period  as  being  divisible  into  two  distinct 
stages. 

Judging  from  the  evidence  of  distribution  of 
mammals  in  pleistocene  Europe,  Professor  Boyd 
Dawkins  came  to  the  conclusion  (p.  113)  that  the 
climate  of  our  continent  "was  severe  in  the  north 


THE  SIBERIAN    MIGRATION.  209 

and  warm  in  the  south,  while  in  the  middle  zone, 
comprising  France,  Germany,  and  the  greater  part 
of  Britain,  the  winters  were  cold  and  the  summers 
warm,  as  in  Middle  Asia  and  North  America."  "  In 
the  summer  time  the  southern  species  would  pass 
northwards,  and  in  the  winter  time  the  northern 
would  swing  southwards,  and  thus  occupy  at  different 
times  of  the  year  the  same  tract  of  ground,  as  is  now 
the  case  with  the  Elks  and  Reindeer."  Very  different 
are  the  views  of  Professor  Nehring  on  this  subject. 
According  to  him,  the  climate  in  Germany  must 
have  been  extremely  cold  and  damp,  resembling 
that  of  Greenland,  though  perhaps  not  quite  so 
arctic.  Professor  Nehring  does  not  at  all  believe 
that  southern  and  northern  species  of  mammals  could 
have  lived  in  Central  or  Northern  Europe  at  the 
same  time;  though  of  this  we  have  undoubted  geo- 
logical evidence  (pp.  72-75).  He  thinks  that  the 
supposed  commingling  of  southern  and  northern 
types,  which  has  actually  been  shown  by  Professor 
Dawkins  to  occur,  is  either  due  to  careless  observa- 
tion or  to  the  fact  that  some  of  the  species  need  not 
necessarily  have  lived  where  their  bones  were  found 

(P-  133). 

The  most  reliable  conclusions  as  regards   former 

conditions  of  vegetation  and  climate  can  be  drawn, 
according  to  Professor  Nehring,  from  the  smaller 
burrowing  mammals,  such  as  the  marmots,  sousliks, 
etc.  He  is  of  opinion  that  a  great  portion  of 
Northern  Europe,  where  their  remains  have  been 


210         HISTORY  OF   THE   EUROPEAN    FAUNA. 

discovered,  must  have  possessed  tundras  and  steppes, 
as  we  find  them  nowadays  in  Siberia,  and  a  climate 
similar  to  that  of  Northern  Asia.  It  is  presumed 
that  the  climate,  after  the  maximum  cold  of  the  first 
stage  of  the  Ice- Age,  ameliorated  so  far  as  to  permit 
these  mammals  to  exist  in  Europe. 

The  natural  question,  however,  which  is  forced 
upon  us  in  reading  Professor  Nehring's  interesting 
and  suggestive  work  is,  where  did  all  these  steppe 
animals  live  during  the  earlier  part  of  the  Ice- Age? 
No  traces  of  their  remains  have  been  discovered  in 
Southern  Europe,  and  it  can  therefore  certainly  be 
affirmed  that  they  could  not  have  lived  there.  If 
Central  and  Northern  Europe  were  uninhabitable 
for  mammals,  Central  and  Northern  Asia  must  have 
been  even  more  so,  and  we  have  to  fall  back  upon  the 
Oriental  Region  as  a  possible  home  of  these  species 
during  the  assumed  maximum  cold  of  the  Glacial 
period.  In  invading  Europe  from  the  Oriental 
Region  these  Siberian  mammals  would  have  taken 
the  shorter  route  by  Asia  Minor  and  Greece,  which 
was  open  to  them.  This  they  certainly  did  not  do, 
which  proves  that  they  came  directly  from  Siberia  to 
Europe  without  retreating  first  to  Southern  Asia. 

But  it  seems  to  me  that  there  is  no  necessity  for 
assuming  such  drastic  changes  of  climate  to  have 
taken  place  at  all  (compare  pp.  75-80).  We  really 
have  no  idea  under  what  precise  climatic  conditions 
the  Siberian  mammals  lived  in  their  original  home. 
The  only  thing  we  can  be  certain  of  is  that  the 


THE   SIBERIAN   MIGRATION.  211 

smaller  burrowing  mammals  would  not  have  chosen 
a  wood  to  live  in,  if  they  could  possibly  help  it. 
Prairies,  or  sand-dunes  with  short  grass  or  shrubs, 
such  as  abound  in  Europe  near  the  sea-coast,  would 
suit  these  species  perfectly.  If  we  suppose  Northern 
Germany  to  have  been  covered  by  sea  (p.  156)  during 
part  of  the  Pleistocene  Epoch,  forests  would  probably 
not  have  grown  there  for  a  very  considerable  time 
afterwards,  owing  to  the  excessive  salinity  of  the  soil, 
but  a  tract  of  sandy  country  would  have  been  left 
on  the  retreat  of  the  sea.  Possibly  a  slight  change 
of  climate  in  the  original  home  of  these  steppe-species 
may  have  reduced  their  habitable  area,  and  thus 
caused  their  migration  into  Europe. 

But  this  migration  problem  cannot  be  solved 
without  tracing  the  mammals  to  their  place  of 
origin  and  investigating  their  early  history.  This  I 
shall  attempt  to  do  presently;  meanwhile,  it  would 
be  interesting  to  note  whether  other  groups  of 
animals  support  Professor  Nehring's  steppe-theory. 

Among  groups  other  than  mammals,  the  most 
important,  for  the  purpose  of  drawing  conclusions 
as  to  former  physical  conditions  and  climate,  are  the 
mollusca.  Their  remains  have  been  well  preserved, 
and  are  easily  identified.  Though  Professor  Nehring 
argues  that  the  molluscs  found  along  with  the  small 
mammals  harmonise  perfectly  with  the  assumption 
of  a  steppe-climate  (p.  212),  I  cannot  at  all  agree 
with  him.  He  enumerates  the  following  sixteen 
species  as  having  been  discovered  by  him  : — 


212         HISTORY   OF   THE   EUROPEAN   FAUNA. 

1.  Pupa  muscorum.  9.  Helix  pulchella. 

2.  Chondrula  tridens.  10.  Do.    hortensis. 

3.  Cionella  lubrica.  u.  Do.    obvoluta. 

4.  Patula  ruderata.  12.  Hyalinia  radiatula. 

5.  Do.    rotundata.  13.  Succinea  oblonga. 

6.  Helix  striata.  14.  Limnaea  peregra. 

7.  Do.    hispidia.  15.  Clausilia  sp. 

8.  Do.    tenuilabris.  16.  Pisidium  pusillum. 

Only  two  of  these  can  be  looked  upon  as  typically 
northern  species,  viz.,  Patula  ruderata  and  Helix 
tenuilabris^  though  both  of  them  are  still  found 
living  locally  in  Germany.  Some  of  the  others 
are  decidedly  southern  species,  like  Chondrula  tridens, 
Helix  obvoluta,  H.  rottmdata,  and  H.  striata.  All  the 
rest  live  and  flourish,  for  example,  in  Ireland  at  the 
present  day,  where,  as  we  all  know,  anything  but  a 
dry  steppe-climate  prevails. 

Dr.  Kobelt  quite  agrees  with  me  in  thinking  that 
the  remains  of  the  mollusca  found  along  with  the 
so-called  "steppe-mammals"  afford  no  proof  of  a 
steppe-character  of  the  country  at  the  time  when 
they  were  alive  (p.  166).  Nor  do  the  mollusca  which 
have  been  found  in  England  in  the  Forest-Bed  and 
the  succeeding  pleistocene  strata  support  such  a 
view.  The  Forest-Bed,  generally  regarded  as  belong- 
ing to  the  Upper  Pliocene,  I  believe  to  be  an  inter- 
glacial  pleistocene  deposit — contemporaneous  with  the 
loess  formation  in  Germany.  Of  fifty-nine  species 
of  land  and  freshwater  mollusca  which  have  been 
discovered  in  this  bed,  forty-eight  species,  according 


THE   SIBERIAN    MIGRATION. 

to  Mr.  Clement  Reid  (p.  186),  are  at  present  living 
in  Norfolk,  six  are  extinct,  two  are  continental 
forms  living  in  the  same  latitudes  as  Norfolk,  and  the 
other  three  are  all  southern  forms.  Not  a  single 
species  has  a  particularly  northern  range.  Of  the 
land  and  freshwater  mollusca  of  the  South  of  Eng- 
land in  the  succeeding  pleistocene  deposits,  six 
species  are  now  no  longer  living  in  the  British 
Islands,  but  only  one  (Helix  ruderata)  can  be  looked 
upon  as  an  Arctic  or  Alpine  form.  After  this  short 
digression  on  the  mollusca,  I  will  briefly  recapitulate 
what  is  known  about  the  early  history  of  the  Siberian 
mammals,  which  will  assist  us  in  tracing  the  cause  of 
their  migration  to  Europe. 

We  have  in  Siberia  problems  quite  as  difficult  of 
solution  as  the  European  ones.  Volumes  have  been 
written  to  explain  the  former  presence  of  Arctic 
mammals  like  the  Reindeer  in  Southern  Europe,  and 
the  most  extraordinary  demands  on  the  credulity  of 
the  public  have  been  made  by  some  geologists  in 
their  attempts  to  account  for  this  comparatively  simple 
problem.  In  Northern  Asia  a  somewhat  similar 
phenomenon,  but  much  more  difficult  of  explanation, 
has  taken  place.  Mammals  have  been  found  fossil  in 
recent  geological  deposits  in  localities  where  they  do 
not  now  occur,  and  apparently  the  Siberian  and  the 
European  deposits  are  of  about  the  same  age.  Now, 
however,  comes  the  extraordinary  difference.  In 
Europe  the  Arctic  mammals  went  southward,  but 
in  Siberia  the  Southern  ones  went  northward.  Not 


214         HISTORY   OF   THE   EUROPEAN    FAUNA. 

only  do  we  find  the  Saiga-Antelope,  Tiger,  Wild 
Horse,  European  Bison,  Mammoth,  and  Rhinoceros 
in  the  extreme  north  of  the  mainland  of  Siberia;  their 
remains  have  even  been  obtained  in  the  New  Siberian 
Islands.  As  these  islands  are  situated  in  the  same 
latitude  as  the  northern  part  of  Novaya  Zemlya, — 
indeed,  not  far  south  of  the  latitude  of  Spitsbergen, 
— the  fact  of  such  huge  mammals  having  been  able 
to  find  subsistence  there  at  apparently  quite  a  recent 
geological  period  seems  an  astounding  fact.  It  may 
be  urged  that  their  bones  might  have  been  carried  so 
far  north  by  ice,  or  by  some  other  equally  powerful 
agency.  But  Tcherski  and  all  other  palaeontologists 
who  have  examined  these  northern  deposits  are 
unanimous  in  the  belief  that  these  herbivores  and 
carnivores  lived  and  died  where  their  remains  are 
now  found.  "It  is  evident,"  says  Tcherski  (p.  451), 
"that  these  large  animals  could  only  have  lived  in 
those  extremely  northern  latitudes  under  correspond- 
ingly favourable  conditions  of  the  vegetation,  viz., 
during  the  existence  of  forests,  meadows,  and 
steppes."  He  also  is  of  opinion  that  the  moist  climate 
which  evidently  prevailed  in  Europe  during  Post- 
tertiary  (Pleistocene)  times  must  have  modified  the 
Siberian  climate  in  so  far  as  to  render  it  milder. 
The  existence  of  the  Aralo-Caspian  basin  (Fig.  12, 
p.  156)  must  also  have  tended  in  the  same  direction. 
It  appears  then  that,  at  the  time  when  plants  and 
animals  are  believed  to  have  retired  southward  in 
Europe  before  the  supposed  advancing  Scandinavian 


THE  SIBERIAN    MIGRATION.  215 

ice-sheet,  no  agency  existed  in  North  Siberia  which 
was  able  to  suppress  and  to  annihilate  the  forest  and 
meadow  vegetation,  and  drive  away  the  fauna  con- 
nected with  it.  We  know,  continues  Tcherski,  that 
such  genera  as  Bison,  Colus  (Saiga\  Rhinoceros, 
Elephas,  and  Equus  are  met  with  in  all  horizons 
of  the  diluvium  of  West  Siberia.  He  therefore 
comes  to  the  conclusion  (p.  474),  that  these  and 
other  facts  imply  that  the  retreat  of  the  North 
Asiatic  fauna  commenced  about  the  end  of  the 
Tertiary  Era  (Pliocene),  and  that  it  was  continued 
very  slowly  throughout  the  Post-tertiary  (Pleistocene) 
Epoch,  without  any  visible  changes  in  its  southward 
direction,  even  during  the  time  of  the  most  important 
glacial  developments  in  Northern  Europe.  Only  after 
the  conditions  disappeared  which  had  produced  the 
augmentation  of  an  atmospheric  moisture,  did  the 
climate  of  North  Siberia  become  deadly  to  a 
temperate  fauna  and  flora.  Tundras  then  spread 
over  the  meadow-lands  and  remnants  of  forests, 
whilst  arctic  animals  replaced  the  large  ungulates 
and  carnivores  which  had  wandered  far  away  from 
their  native  southern  home. 

This  is  Tcherski's  explanation  of  the  extraordinary 
events  which  he  has  chronicled,  after  years  of  the 
most  arduous  labour  and  under  conditions  of  peculiar 
hardship.  And  though  his  work  cannot  be  over- 
estimated, and  his  opinions  should  receive  the  most 
careful  consideration,  yet  I  fear  the  explanation  will 
not  be  looked  upon  as  entirely  satisfactory.  Every  one 


2l6         HISTORY   OF   THE   EUROPEAN    FAUNA. 

will  agree  with  him  that  the  climate  of  Siberia  must 
have  been  greatly  moister  in  pliocene  and  pleistocene 
times  than  it  is  now.  The  Aralo- Caspian  covered 
a  vast  area  of  South-western  Siberia.  Freshwater 
basins  existed  along  the  east  of  the  Ural  Mountains, 
while  Central  Asia  was  studded  over  with  a  number  of 
large  lakes,  which  have  now  almost  entirely  vanished. 
But  that  the  generally  assumed  refrigeration  of 
Europe  must  have  had  a  chilling  effect  on  the 
Siberian  atmosphere  seems  to  me  evident.  That  the 
whole  of  Northern  Europe  should  have  been  made 
uninhabitable  owing  to  the  advance  of  the  Scandi- 
navian ice-sheet,  while  North  Siberia  at  the  same 
time  supported  forests,  meadows,  and  a  temperate 
fauna,  is  incredible.  At  the  approach  of  winter,  at 
any  rate,  the  animals  would  have  been  driven  south- 
ward for  thousands  of  miles  to  seek  shelter  from  the 
snows  and  cold  and  to  obtain  nourishment,  and  it 
would  scarcely  have  been  possible  for  them  to 
undertake  such  vast  migrations  at  every  season. 
Professor  James  Geikie's  suggestion  (p.  706),  that 
the  Mammoth  and  Woolly  Rhinoceros  could  have 
survived  the  Pleistocene  Epoch  in  Southern  Siberia, 
does  not  appear  to  solve  the  problem,  as  that  part 
of  Asia  must  have  participated  in  the  great  cold 
which  is  said  to  have  prevailed  all  over  Europe. 

Let  us  now  concede,  for  the  sake  of  argument, 
that  the  current  views  regarding  the  pleistocene 
climate  of  Europe  are  correct.  We  are  told  by  Pro- 
fessor Geikie  that  the  climate  of  Scotland  during 


THE   SIBERIAN    MIGRATION.  217 

part  of  the  Pleistocene  Epoch  was  so  cold,  that  the 
whole  country  was  buried  underneath  one  immense 
mer  de  glace,  through  which  peered  only  the  higher 
mountain-tops  (p.  67).  If  this  was  the  state  of  climate 
in  close  proximity  to  the  Atlantic,  it  must  probably 
have  been  still  more  severe  on  the  European  con- 
tinent. Now  at  the  present  time  Siberia  has  the 
reputation  of  being  the  coldest  country  in  the  world, 
and  the  mercury  of  the  thermometer  is  said  to  remain 
frozen  for  weeks  during  winter,  even  in  the  south. 

With  the  prevailing  dampness  in  pleistocene  times 
the  snowfall  throughout  Siberia  would  have  been 
much  heavier  than  at  present,  though  it  would  have 
modified  the  temperature  to  some  extent.  Under 
such  circumstances  Southern  Siberia  could  not  have 
been  a  desirable  place  of  residence  for  large  mammals. 
It  would  have  been  necessary  for  the  Mammoth  and 
the  other  species  referred  to,  to  wander  farther  into 
the  extreme  south  of  Asia  or  Europe  to  find  a 
suitable  refuge  during  the  arctic  conditions  which 
are  supposed  to  have  prevailed  in  Northern  Europe. 
To  quote  Professor  J.  Geikie's  own  words  (p.  706): 
"  They  (Mammoth,  etc.)  would  seem  to  have  lived  in 
Southern  Siberia  throughout  the  whole  Pleistocene 
period,  from  which  region  doubtless  they  originally  in- 
vaded our  Continent.  But  with  the  approach  of  our 
genial  forest-epoch  (penultimate  inter-glacial  stage) 
they  gradually  vanished  from  Europe,  to  linger  for  a 
long  time  in  Siberia  before  they  finally  died  out." 
It  is  suggested,  therefore,  by  the  author  that  the 


218         HISTORY  OF   THE   EUROPEAN    FAUNA. 

Mammoth  and  the  other  mammals  whose  remains 
have  been  discovered  on  the  New  Siberian  Islands 
found  their  way  there  during  one  of  the  late  inter- 
glacial  stages  of  the  Ice-Age.  But  there  is  no 
astonishment  expressed  by  Professor  Geikie  at  the 
extraordinary  change  of  climate  which  must  have 
occurred  in  Siberia  to  allow  of  such  migrations.  I 
can  find  no  very  definite  statement  in  this  author's 
work  as  to  the  nature  of  the  climate  in  Europe  during 
those  inter-glacial  phases,  but  he  remarks  (p.  129) 
"  that  the  evidence  of  the  Scottish  inter-glacial  beds, 
so  far  as  it  went,  did  not  entitle  us  to  infer  that 
during  their  accumulation  local  glaciers  may  not 
have  existed  in  the  Highland  valleys."  There  is  no 
evidence,  in  other  words,  of  the  existence  in  Europe 
of  a  milder  climate  than  that  prevailing  at  present. 
Still  less  can  there  be  any  ground  for  the  supposition 
that  the  climate  of  the  whole  of  Siberia  ameliorated 
to  such  an  extent  that  forests  and  meadows  could 
develop  as  far  north  as  the  New  Siberian  Islands ; 
for  if  the  temperature  in  Europe  was  then  about 
the  same  as  now,  that  of  Siberia  could  not  have 
been  vastly  higher  than  it  is  at  present. 

It  is  highly  improbable,  therefore,  that  a  sufficiently 
mild  climate  prevailed  in  the  extreme  north  of 
Siberia  during  the  so-called  later  inter-glacial  periods 
to  induce  the  mammals  to  which  I  have  referred  to 
seek  fresh  pastures  there, 

The  late  Professor  Brandt,  one  of  the  highest 
zoological  authorities  in  Russia,  was  of  opinion  that 


THE  SIBERIAN   MIGRATION.  2 19 

at  the  commencement  of  the  Glacial  period  the  great 
mammals  of  Northern  Siberia  either  perished  or 
migrated  southward.  From  there  they  gradually 
penetrated  into  European  Russia.  He  believed  that 
before  the  Glacial  period  a  connection  qxisted 
between  the  Aralo-Caspian  Sea  and  the  Arctic 
Ocean,  carrying  warm  water  northward.  The  gradual 
disappearance  of  this  marine  channel  caused  a 
decrease  of  warmth  in  Northern  Asia,  so  that  large 
accumulations  of  frozen  soil  and  ice  were  formed, 
which  still  more  depressed  the  temperature.  This, 
he  suggested,  probably  took  place  at  the  time  when 
the  Glacial  period  commenced  in  North-western 
Europe. 

It  has  been  urged  against  these  views  of  Tcherski 
and  Brandt,  that  the  bone  beds  in  the  Liakov  Islands 
(New  Siberian  Islands)  rest  partly  upon  a  solid  layer 
of  ice  of  nearly  seventy  feet  thick.  This  mass  of  ice, 
it  was  thought,  must  have  accumulated  during  the 
Glacial  period.  As  the  bones  rest  upon  it,  the 
mammals  could  only  have  lived  in  those  islands  in 
more  recent  times,  after  the  Ice- Age  had  passed  away. 
Nothing,  apparently,  can  be  clearer,  and  yet  in  the 
face  of  this  seeming  proof  one  feels,  as  I  have  men- 
tioned before,  that  if  such  an  extraordinary  revolution 
of  climate  as  is  implied  by  this  admission  had  taken 
place,  we  should  be  able  to  perceive  the  traces 
throughout  the  northern  hemisphere.  In  this  di- 
lemma, a  suggestion  made  by  Dr.  Bunge,  who  visited 
the  New  Siberian  Islands  recently  at  the  instance  of 


22O         HISTORY   OF  THE   EUROPEAN   FAUNA. 

the  Imperial  Academy  of  St.  Petersburg,  helps 
us  out  of  the  difficulty.  He  found  that,  as  a 
rule,  these  so-called  fossil  glaciers  contain  seams 
of  mud  and  sand,  and  he  argued  that  the  ice  had 
formed,  and  is  still  forming  at  the  present  day,  in 
fissures  of  the  earth.  I  entirely  concur  with  this 
view.  Neither  palaeontology  nor  the  geographical 
distribution  of  animals  lend  any  support  to  the  other 
theory,  and  I  think  we  may  conclude  that  Brandt's 
view  in  the  main  is  probably  the  correct  explanation 
of  the  phenomena  which  we  have  discussed.  Some 
important  facts  of  distribution  are  more  easily  explic- 
able on  this  assumption.  Why,  for  instance,  should 
the  Siberian  fauna  of  pliocene  times  have  remained 
in  Siberia  and  not  have  migrated  to  Europe  at  that 
time?  The  pliocene  mammals  of  Siberia  are  mostly 
of  southern  origin.  Their  range  increased  enormously 
during  the  epoch  throughout  Northern  Asia.  We 
should  expect  them,  therefore,  to  have  crossed  the 
Caspian  plains,  or  even  the  low-lying  Ural  Mountains, 
to  pour  into  the  neighbouring  continent.  But  Pro- 
fessor Brandt  explained  how  they  were  prevented 
from  spreading  west.  An  arm  of  the  sea  stretched 
from  the  Aralo-Caspian  to  the  Arctic  Ocean,  thus 
raising  an  effectual  barrier  between  the  two  con- 
tinents.  There  is  some  evidence  for  the  belief,  as  we 
shall  learn  presently,  that  this  marine  barrier  existed 
also  during  the  early  part  of  the  pleistocene  epoch, 
After  having  greatly  expanded  during  pliocene  times, 
the  fauna  of  Siberia  gradually  withdrew  from  the 


THE  SIBERIAN    MIGRATION.  221 

northern  regions  during  the  earlier  portion  of  the 
succeeding  epoch.  It  was  only  after  the  marine 
connection  above  referred  to  ceased  to  exist,  or 
became  disconnected,  that  an  entry  into  Europe 
was  possible. 

A  fauna,  to  some  extent  composed  of  species  now 
inhabiting  the  steppes  of  Eastern  Europe  and  Siberia, 
poured  into  the  neighbouring  continent.  On  p.  95  I 
have  given  a  list  of  those  which  reached  as  far  west 
as  the  British  Islands,  but,  as  I  mentioned,  many 
other  species  came  from  the  east  about  this  time. 
With  regard  to  the  early  history  of  the  Siberian 
mammals,  I  favour  a  view  somewhat  between  that 
of  Tcherski  and  that  of  Brandt.  The  outpouring  of 
the  fauna  into  Europe  seems  to  me  to  indicate  that 
there  was  a  sudden  change  of  climate  in  Siberia. 
This  was  produced,  perhaps,  by  the  rupture  of  the 
marine  connection  between  the  Arctic  Ocean  and 
the  Aralo-Caspian.  Such  an  event  would  not  only 
have  caused  the  sudden  shrinkage  of  the  area  avail- 
able for  food-supply  by  lowering  the  temperature  in 
Siberia,  it  would  have  acted  also  as  a  means  in 
assisting  the  fauna  to  enter  a  new  continent  where 
an  inconsiderable  number  of  mammals,  already  estab- 
lished, were  mostly  dispossessed  of  their  homes  by  the 
advancing  eastern  host. 

Brandt's  theory,  however,  of  a  marine  connection 
between  the  Arctic  Ocean  and  the  Aralo-Caspian  is 
by  no  means  generally  accepted.  That  the  Caspian 
Sea  was  at  that  time  greatly  larger  than  it  is  at 


222          HISTORY   OF   THE   EUROPEAN    FAUNA. 

present,  and  joined  to  the  Sea  of  Aral  and  the  Black 
Sea,  is  acknowledged  by  everybody.  That  the 
deposits  laid  down  by  this  huge  inland  sea  reach 
as  far  north  as  the  shores  of  the  river  Kama,  in 
Central  Russia,  is  also  well  known  to  geologists. 
But  what  comes  rather  as  a  surprise,  is  that  Professor 
Karpinski,  whom  we  must  take  as  one  of  the  highest 
authorities  on  the  geology  of  Russia,  asserts  that  this 
Aralo-Caspian  Sea  was  probably  joined  by  a  system 
of  lakes  or  channels  to  the  Arctic  Ocean  (p.  183).  He 
was  by  no  means  the  first,  though,  to  put  forward 
such  a  theory.  We  have  already  learned  that  Professor 
Brandt  held  a  somewhat  similar  view,  though  he 
believed  in  something  more  than  a  connection  by 
mere  channels,  and  Mr.  Koppen,  and  also  the  Russian 
traveller  Mr.  Kessler,  agreed  with  him.  So  much 
was  Professor  Boyd  Dawkins  impressed  with  their 
arguments  at  the  time,  that  he  wrote  (c,  p.  148) : 
"  Before  the  lowering  of  the  temperature  in  Central 
Europe,  the  sea  had  already  rolled  through  the  low 
country  of  Russia,  from  the  Caspian  to  the  White 
Sea  and  the  Baltic,  and  formed  a  barrier  to  western 
migration  to  the  Arctic  mammals  of  Asia" 

In  one  particular  Professor  Dawkins's  views  differ 
from  those  of  almost  all  the  previous  writers.  His 
connection  between  the  Caspian  and  the  Arctic 
Ocean  is  placed  to  the  west  of  the  Ural  Mountains, 
while  it  had  always  been  assumed  by  the  Russian 
writers  to  have  lain  on  the  eastern  or  Asiatic  side  of 
that  mountain  range.  Thus,  when  Tcherski  in  recent 


THE  SIBERIAN    MIGRATION.  223 

years  announced  that  the  tract  on  this  eastern  side  of 
the  mountains  was  covered  by  freshwater  deposits, 
his  discovery  seemed  once  for  all  to  settle  the  problem 
of  the  arctic  marine  connection  in  the  negative.  As 
Professor  Dawkins's  theory  has,  however,  received 
much  additional  affirmative  evidence  by  current 
faunal  researches,  a  connection  between  the  Caspian 
(or  Aralo-Caspian)  and  the  Arctic  Ocean  (White  Sea) 
may  have  actually  existed  within  recent  geological 
times. 

What  relict  lakes  are,  has  already  been  explained 
(p.  176),  and  their  fauna  will  again  be  referred  to  in 
a  subsequent  chapter.  I  might  perhaps  be  allowed 
to  repeat  that  such  lakes  are  supposed  to  have  been 
flooded  by,  or  to  have  been  in  close  connection  with, 
the  sea  at  some  former  period.  Many  of  the  Swedish 
lakes  are  spoken  of  as  relict  lakes  (Reliktenseen), 
because  they  contain  a  number  of  marine  species  of 
animals  which  have  now  become  adapted  to  live  in 
fresh  water,  but  all  of  whose  nearest  relatives  inhabit 
the  sea.  One  of  these,  the  schizopod  crustacean  Mysis 
relicta, — a  shrimp-like  creature, — which  was  formerly 
believed  to  inhabit  also  the  Caspian,  is  of  particular 
interest.  More  recently,  the  occurrence  of  this  Mysis 
in  the  Caspian  was  denied,  but  though  this  denial  has 
been  confirmed  by  Professor  Sars  in  his  memoir  on 
the  crustaceans  of  the  great  Russian  inland  sea,  he 
has  been  enabled  to  add  two  new  species  of  Mysis 
to  the  list  of  those  already  known  to  science.  These 
are  M.  caspia  and  M.  micropthalma,  and  both  are 


224         HISTORY  OF  THE   EUROPEAN   FAUNA. 

closely  related  to  the  arctic  marine  Mysis  oculata. 
According  to  Professor  Sars,  the  genus  Mysis  as  a 
whole  may  be  regarded  as  arctic  in  character.  The 
occurrence  of  these  two  species,  therefore,  in  his 
opinion,  points  to  a  recent  connection  of  the  Caspian 
with  the  Glacial  Sea. 

A  large  number  of  other  crustaceans  have  been 
described  by  the  same  author  from  the  Caspian.  Of 
the  order  Cumacea,  which  is  exclusively  marine, 
ten  species  are  mentioned,  but  none  of  these  seems 
to  range  beyond  the  Caspian.  Among  the  smaller 
species  of  crustaceans,  a  minute  pelagic  copepod 
(Limnocalanus  grimaldii)  also  inhabits  the  Baltic 
and  the  Arctic  Ocean.  The  marine  isopod  Idctea 
entomon,  related  to  the  common  wood-louse,  has  a 
similar  distribution. 

Genuine  Arctic  species  of  Fishes  do  not  seem  to 
occur  in  the  Caspian,  though  some,  viz.,  Clupea 
caspiay  Atherina  pontica,  Clupionella  Grimmi,  and 
Syngnathus  bucctilentus,  are  almost  certainly  the 
descendants  of  marine  forms. 

The  Seal  of  the  Caspian  (Phoca  caspica)  is  closely 
allied  to  the  Arctic  Seal,  and  its  presence  alone  in 
that  sea  indicates  that  at  no  very  distant  date — at 
any  rate  since  pliocene  times — a  closer  connection 
with  the  Arctic  Ocean  existed  than  at  present. 

I  am  sure  it  will  be  readily  granted  that  there  is 
zoological  evidence  for  the  belief  of  such  a  connection 
or  union  between  the  two  great  seas.  However,  it 
may  be  urged  that  owing  to  the  presence  of  an  ice- 


THE   SIBERIAN    MIGRATION. 


225 


sheet  in  Northern  Europe  during  the  Glacial  period, 
such  a  connection  must  either  have  been  pre-glacial 
or  have  existed  after  that  period.  But  the  connec- 
tion must  have  occurred  at  a  time  when  the  Caspian 


FIG.  17. — Map  of  European  Russia  (after  Karpinski).  The  faintly 
dotted  parts  indicate  the  areas  covered  by  boulder-clay,  the 
strongly  dotted  ones  those  exhibiting  Aralo-Caspian  and  other 
post-pliocene  deposits. 

extended  far  to  the  north — when  indeed  the  so-called 
post-tertiary  Caspian  deposits  were  laid  down  (Fig. 
17).  Since  the  boulder-ciay  which  covers  the  plain 
of  Northern  Russia  is  assumed  to  be  the  ground- 

15 


226          HISTORY   OF   THE    EUROPEAN    FAUNA. 

moraine  of  the  great  northern  ice-sheet,  we  might 
expect  to  find  that  the  Caspian  deposits  were  not 
contemporaneous  with  it.  Curiously  enough,  it  has 
been  shown  by  Mr.  Sjogren  that  all  observations 
have  pointed  to  the  fact  that  these  two  deposits  do 
not  overlie  one  another,  but  occur  side  by  side, 
and  are  therefore  contemporaneous.  This  seems  to 
warrant  our  belief,  that  while  the  boulder-clay  was 
being  laid  down  in  Northern  Europe,  the  Aralo- 
Caspian  Sea  had  some  communication  with  the 
White  Sea. 

The  boulder-clay  of  Northern  Continental  Europe, 
as  already  stated,  is  now  generally  recognised  to 
be  the  product  of  a  huge  ice-sheet  which  invaded 
the  lowlands  of  Continental  Europe  from  the  Scan- 
dinavian mountains.  Though  Alpine  glaciers  at  the 
present  day  produce  little  or  no  ground  moraine, 
these  ancient  larger  ice-sheets,  or  "mers-de-glace,"  are 
believed  to  have  deposited  immense  layers  of  mud 
containing  scratched  and  polished  stones.  Many  of 
the  latter  have  been  carried  great  distances  from 
their  source  of  origin.  The  Scandinavian  ice-sheet 
is  supposed  to  have  advanced  as  far  south  as  the 
line  indicated  on  the  map,  after  which  it  gradually 
retreated.  On  this  point,  however,  as  in  almost  every 
detail  connected  with  the  Glacial  period,  geologists 
are  at  variance.  Professor  James  Geikie  maintains, 
that  there  were  no  less  than  four  Glacial  periods, 
separated  from  one  another  by  milder  inter-glacial 
phases.  On  the  Continent  the  view  of  two  Glacial 


THE   SIBERIAN    MIGRATION.  227 

and  one  inter-glacial  period  is,  I  think,  more  gene- 
rally adopted.  Professor  Geikie's  four  periods  seem 
to  me  to  have  originated  i-n  a  desire  to  correlate 
the  British  pleistocene  deposits  with  the  continental 
ones,  and  at  the  same  time  to  retain  the  old  view  of 
the  inter-glacial  position  of  the  Forest-Bed.  The 
two  theories  agree  in  so  far  as  that  in  both  the 
glacial  conditions  culminate  in  a  maximum  glacia- 
tion,  followed  by  a  more  temperate  phase  of  climate, 
with  consequent  retreat  of  the  ice-sheets,  and  finally 
by  a  renewed  advance  of  the  glaciers. 

We  are  told  that  there  is  not  the  slightest  doubt 
about  it  that  a  marked  but  gradual  decrease  of 
temperature  took  place  all  over  Europe  either 
during  the  beginning  of  the  Pleistocene  or  towards 
the  end  of  the  Pliocene  Epoch. 

We  might  reasonably  suppose,  then,  that  a  similar 
climatic  effect  was  produced  in  Siberia,  in  con- 
sequence of  which  the  fauna  would  have  been 
obliged  to  retreat  from  the  extreme  northern  lati- 
tudes southward.  No  doubt  great  efforts  would 
have  been  made  by  the  members  of  the  Siberian 
fauna — at  any  rate  by  those  possessing  strong 
power  of  locomotion — to  extend  their  range  in 
other  directions.  But  we  have  no  evidence  that 
a  migration  from  Siberia  came  to  Eastern  Europe 
at  that  time.  It  seems,  therefore,  as  if  the  barrier 
referred  to  by  Brandt,  Koppen,  Boyd  Dawkins, 
and  others  (p.  222),  had  existed  at  this  time.  This 
would  have  effectually  prevented  an  overflow  of 


228         HISTORY  OF   THE   EUROPEAN    FAUNA. 

the  fauna  from  Siberia.  Only  in  deposits  later 
than  the  lower  continental  boulder-clay  do  we  find 
traces  of  a  Siberian  migration.  The  time  of  maxi- 
mum glaciation  had  then  passed  away ;  the  great 
glacier  which  was  believed  to  have  invaded  the 
lowlands  of  Northern  Europe  had  again  retreated, 
before  the  Siberian  mammals  made  their  appearance 
in  Germany. 

It  has  been  stated  above  (p.  226)  that  while  the 
Russian  boulder-clay  was  being  laid  down,  the  Aralo- 
Caspian  probably  had  some  communication  with  the 
White  Sea. 

But  how  can  this  view  be  reconciled  with  the 
existence  of  a  huge  mer  de  glace  in  the  northern 
plains  of  Russia?  The  existence  of  the  ice-sheet 
has  been  conjured  up  in  order  to  explain  the 
presence  of  the  boulder-clay.  But  not  long  ago  a 
very  different  interpretation  of  the  origin  of  this  clay 
was  given;  and  one,  I  may  say,  which  explains  the 
history  of  the  Siberian  and  the  European  fauna  in  a 
more  satisfactory  manner  than  is  done  by  the  ice- 
sheet  hypothesis.  It  is  that  the  boulder-clay  is  not 
the  product  of  land-ice,  but  has  been  deposited  by  a 
sea  with  floating  icebergs.  Thus  the  latter  hypothesis 
does  not  deny  the  existence  of  glaciers,  but  allows 
.the  mud  to  be  deposited  on  the  floor  of  a  turbid 
sea,  instead  of  beneath  an  immense  mer  de  glace.  I 
need  hardly  mention  that  this  view,  which  was 
formerly  universally  accepted  by  geologists,  is  now 
scouted  by  almost  every  authority,  both  British  and 


THE   SIBERIAN    MIGRATION.  229 

Continental.  I  should  scarcely  venture  the  attempt 
to  revive  old  memories  and  stir  up  again  long 
forgotten  controversies,  were  it  not  for  the  fact  that 
many  new  points  have  arisen  in  the  course  of  the 
above  inquiries,  which  appear  to  me  so  very  difficult 
to  explain  by  the  land-ice  hypothesis,  while  they  are 
comparatively  easy  to  understand  when  we  adopt 
the  old  theory  of  the  marine  origin  of  the  boulder- 
clay.  But  a  few  geologists  even  at  the  present  day, 
while  believing  in  the  land-ice  theory,  recognise  that 
the  marine  hypothesis  should  have  some  considera- 
tion shown  to  it.  I  need  only  remind  glacialists  of  the 
work  recently  published  by  Professor  Bonney.  "  The 
singular  mixture,"  he  remarks  (p.  280),  "and  apparent 
crossing  of  the  paths  of  boulders,  as  already  stated, 
are  less  difficult  to  explain  on  the  hypothesis  of 
distribution  by  floating  ice  than  on  that  of  transport 
by  land-ice,  because,  in  the  former  case,  though  the 
drift  of  winds  and  currents  would  be  generally  in  one 
direction,  both  might  be  varied  at  particular  seasons. 
So  far  as  concerns  the  distribution  and  thickness  of 
the  glacial  deposits,  there  is  not  much  to  choose 
between  either  hypothesis;  but  on  that  of  land-ice 
it  is  extremely  difficult  to  explain  the  intercalation  of 
perfectly  stratified  sands  and  gravels  and  of  boulder- 
clay,  as  well  as  the  not  infrequent  signs  of  bedding 
in  the  latter." 

Now  with  regard  to  the  land -ice  theory,  several 
serious  difficulties  present  themselves  in  connection 
with  the  origin  of  the  European  fauna.  In  the  first 


230         HISTORY  OF  THE  EUROPEAN    FAUNA. 

place,  as  the  climate  renders  Northern  Siberia 
almost  uninhabitable  for  mammals  at  the  present 
day,  how  much  more  severe  must  it  have  been 
during  the  time  of  the  maximum  glaciation  in 
Europe.  As  the  then  existing  fauna  was  not 
driven  into  Europe,  where  could  it  possibly  have 
survived  ?  Secondly,  how  can  we  reconcile  the 
contemporaneous  existence  of  a  great  inland  sea 
(the  Aralo-Caspian)  containing  survivals  of  mild 
Sarmatic  times  with  an  immense  glacier  almost 
touching  it  on  its  northern  shores?  How  did 
one  of  the  most  characteristic  species  of  that  sea, 
Dreyssensia  polymorpha,  come  to  make  its  appearance 
in  the  lower  boulder-clay  of  Prussia  and  then  dis- 
appear in  the  upper?  And  finally,  how  are  we  to 
explain  the  sudden  appearance  of  a  Siberian  fauna 
after  the  deposition  of  the  lower  boulder-clay,  except 
by  the  removal  of  a  barrier  which  had  prevented 
their  egress  from  Siberia? 

If  we  assume  that  the  continental  boulder-clay  of 
Russia  has  been  formed  in  the  manner  so  ably 
explained  by  Murchison,  de  Verneuil,  and  von  Key- 
serling,  viz.,  by  a  sea  with  floating  icebergs,  the 
temperature  of  Siberia  might  have  been  higher  than 
at  present,  and  have  supported  a  fauna  in  more 
northern  latitudes. 

The  contemporaneousness  of  the  deposits  of  this 
sea  with  those  of  the  Aralo-Caspian  is  also  rendered 
more  intelligible.  If  we  suppose,  moreover,  the  con- 
nection between  the  Aralo-Caspian  and  the  White 


THE  SIBERIAN   MIGRATION.  231 

Sea  (Fig.  12,  p.  156)  to  have  existed  at  this  time,  we 
possess  an  explanation  of  the  method  of  migration  of 
the  Arctic  marine  species  into  the  Southern  and  of 
the  Caspian  species  (Dreyssensia)  into  the  Northern 
Sea. 

An  inter-glacial  phase  is  believed  to  have  super- 
vened after  the  deposition  of  the  lower  boulder-clay, 
and  it  is  during  this  period  that  the  Siberian  species 
first  appeared  in  Central  Europe.  If  we  assume  then 
that  the  retreat  of  the  Northern  Sea  (Fig.  13,  p.  170) 
opened  up  a  passage  for  the  Siberian  fauna,  we  have  in 
this  very  fact  also  an  explanation  of  the  extraordinarily 
large  exodus  of  Asiatic  "mammals,  because  the  great 
reduction  of  the  marine  area  in  Northern  Europe 
would  have  had  an  important  influence  in  lowering 
the  temperature  in  Asia.  Only  a  sudden  change  of 
climate  in  Siberia  could  have  brought  about  the 
migration  of  the  vast  hordes  of  Asiatic  mammals 
whose  remains  we  find  in  Central  and  Western 
Europe  in  deposits  of  that  period. 

Throughout  this  work  we  are  made  acquainted 
with  facts  which  bear  out  the  view  that  the  climate 
during  the  greater  part  of  the  Glacial  period  was 
mild  rather  than  intensely  arctic  in  Europe.  That 
a  huge  ice-sheet  could  have  covered  Northern  Europe 
under  such  conditions  appears  to  me  very  doubt- 
ful. No  one  can  deny,  however,  that  glaciers  must 
have  existed  during  the  Glacial  period  in  all  the 
mountainous  regions  of  Central  and  Northern 
Europe,  though  their  existence  is  not  incompatible 


232         HISTORY  OF  THE   EUROPEAN   FAUNA. 

with  a  mild  climate.  Tree-ferns  and  other  tropical 
vegetation  grow  at  the  foot  of  glaciers  in  New 
Zealand.  We  need  not  even  go  so  far  afield,  for  in 
Switzerland  grapes  ripen  and  an  abundant  fauna  and 
flora  thrive  in  close  proximity  to  some  of  the  well- 
known  glaciers. 

One  matter  of  importance  still  remains  to  be  con- 
sidered before  concluding  this  chapter,  viz.,  the 
fauna  contained  in  the  English  geological  deposit 
known  as  the  "  Forest-Bed."  This  interesting 
deposit  is  exposed  at  the  base  of  a  range  of  cliffs 
on  the  coast  of  Norfolk.  It  is  composed  of  beds 
of  estuarine  and  marine  origin.  The  tree-stumps 
formerly  believed  to  be  the  remains  of  trees  in  situ 
have,  after  more  careful  examination,  proved  to  be 
in  all  cases  drifted  specimens.  A  portion  of  the 
"Forest-Bed"  no  doubt  was  laid  down  in  close  proxi- 
mity to  a  large  river,  and  subject  to  being  periodi- 
cally flooded  by  it.  It  is  not  absolutely  certain,  there- 
fore, that  all  the  mammals  whose  remains  occur  in 
this  deposit  lived  in  England  or  whether  only  on  the 
banks  of  the  river  farther  south.  Nevertheless,  we 
may  take  for  granted  that  some  of  them  did. 
England  was  at  the  time  connected  with  France 
and  Belgium,  and  for  our  purpose  it  matters  little 
whether  they  had  crossed  the  Channel  or  inhabited 
those  parts  of  the  Continent  through  which  the  great 
river  flowed  which  sent  its  alluvial  detritus  as  far  as 
the  plains  of  Norfolk.  All  we  have  to  remember 
is  that  certain  mammals,  which  appear  to  have 


THE   SIBERIAN    MIGRATION.  233 

originated  in  Siberia,  and  of  which  we  have  some 
evidence  that  they  crossed  Central  Europe  in  their 
westward  course,  had  now  reached  the  great  river 
just  alluded  to,  which  some  geologists  believe  to  have 
been  the  Rhine. 

I  have  had  occasion  to  refer  to  a  number  of  British 
mammals  (p.  202) — some  of  which  are  now  extinct — 
which  I  believe  to  have  migrated  across  the  plains  of 
continental  Europe  direct  from  Siberia.  There  were 
twenty-six  species  of  these  Siberian  mammals;  and  no 
less  than  ten  of  these  occur  in  the  Forest-Bed.  None 
appear  in  any  older  British  deposit.  It  is  perfectly 
clear,  therefore,  that  the  Forest-Bed  must  have  been 
laid  down  after  their  immigration  into  Europe.  They 
probably  wandered  to  Western  Europe  very  soon 
after  crossing  the  eastern  boundaries  of  our  conti- 
nent ;  the  deposits  in  which  they  are  found  are  there- 
fore contemporaneous.  But  we  have  learned  above 
(p.  208),  that  the  beds  in  Eastern  Europe  in  which  the 
Siberian  mammal-remains  are  found  are  more  recent 
than  the  lower  boulder-clay.  As  already  stated,  the 
Forest-Bed  must  also  be  more  recent  than  the  lower 
continental  boulder-clay,  and  should  be  included  in 
the  pleistocene  series. 

That  the  Forest-Bed  is  an  inter-glacial  deposit  has 
been  urged  long  ago  by  various  writers.  Professor 
Geikie  regards  it  as  stratigraphically  contempor- 
aneous with  the  peat  and  freshwater  beds  below  the 
lower  diluvium  of  Western  and  Middle  Germany,  and 
as  having  been  laid  down  during  the  first  Inter-glacial 


234         HISTORY  OF   THE   EUROPEAN   FAUNA. 

Epoch  of  the  great  Ice- Age.  The  fact  that  no 
boulder-clay  underlies  the  Forest-Bed  seems  rather 
a  strong  argument  against  the  view  of  its  being  an 
inter-glacial  deposit.  It  lies  directly  on  what  is 
known  as  the  Newer  Pliocene  Crags.  If  the  Forest- 
Bed  is  included  in  the  pleistocene  series,  as  I  sug- 
gested it  should,  the  crags,  or  a  portion  of  them, 
would  therefore  be  equivalent  as  regards  time  of 
deposition  to  the  lower  continental  boulder-clay. 
And  again,  if  the  lower  continental  boulder-clay  is 
contemporaneous  with  the  Newer  Crags,  the  latter 
should  also  be  classed  with  the  pleistocene  strata.  I 
can  scarcely  hope  that  geologists  will  be  ready  to 
admit  such  a  sweeping  change  of  nomenclature 
without  a  protest.  I  venture,  therefore,  to  explain 
more  fully  my  reasons  for  adhering  to  these  un- 
orthodox views. 

Let  us  look  once  more  at  the  map  which  I  con- 
structed (Fig.  12,  p.  156)  to  elucidate  the  migration 
of  the  Arctic  terrestrial  species  to  the  British  Islands. 
It  will  be  noticed  that  one  continuous  ocean  extends 
from  the  east  coast  of  England  across  Holland, 
Northern  Germany,  and  Russia  to  the  White  Sea. 
At  the  same  time  Greenland  and  Northern  Scandi- 
navia, Scotland  and  Southern  Scandinavia,  are  united 
by  a  narrow  strip  of  land,  and  so  are  England  and 
France.  The  waters  of  the  Atlantic  and  this  North 
European  Sea  do  not  therefore  intermingle  at  any 
point,  the  two  seas  being  absolutely  independent  of 
one  another. 


THE  SIBERIAN   MIGRATION.  235 

Such  I  assume  to  have  been  the  geographical  con- 
dition of  Northern  Europe  during  the  deposition  of 
the  Red  Crag.  Arctic  mollusca  were  then  brought 
to  the  east  coast  of  England,  and  boulders  were 
scattered  through  the  beds  laid  down  on  that  coast 
by  icebergs  which  had  been  cast  off  by  Scandinavian 
glaciers  on  reaching  the  sea.  Bedded  clays  which 
have  yielded  arctic  shells  He  beneath  the  lower  con- 
tinental boulder-clay  on  the  Baltic  coast-lands  and  on 
the  coast  of  the  White  Sea.  According  to  Professor 
Geikie,  marine  clays  on  the  same  geological  horizon 
reach  an  elevation  of  some  230  feet.  "  It  would  seem, 
then,"  he  says,  "that  before  the  deposition  of  the 
lower  boulder-clay  of  those  regions  the  Baltic  Sea 
had  open  communication  with  the  German  Ocean " 
(p.  442).  All  these  clays  are  evidently  deposits  of  the 
same  sea.  But  apart  from  the  fact  that  the  Red  Crag 
and  these  Baltic  deposits  are  the  oldest  of  the  upper 
Tertiary  beds  containing  arctic  shells,  there  is  no 
evidence  that  they  are  contemporaneous. 

Overlying  the  same  Baltic  deposits  comes  the  lower 
boulder-clay,  reaching  a  thickness  of  several  hundred 
feet  in  some  parts  of  Germany.  It  presents,  like  the 
upper  clay,  frequent  interstratification  with  well- 
bedded  deposits  of  sand  and  gravel.  The  scarcity  of 
marine  mollusca,  the  occurrence  of  striated  surfaces, 
and  the  occasional  presence  of  so-called  giants' 
kettles,  appear  to  favour  the  view,  which  at  present 
is  generally  adopted  by  both  British  and  Continental 
geologists,  that  the  boulder-clay  owes  its  origin  to 


236         HISTORY  OF   THE   EUROPEAN    FAUNA. 

land-ice.  I  have  stated  on  several  occasions  that  the 
view  of  the  marine  origin  of  the  boulder-clay  agrees 
best  with  the  known  facts  of  distribution,  and  with 
the  history  of  the  European  fauna  (pp.  80-86,  and 
p.  129).  It  may  be  urged  that  if  the  lower  boulder- 
clay  were  contemporaneous  with  the  British  Crags 
which  succeeded  the  Red  Crag,  how  can  we  explain 
the  fact  that  these  crags  contain  plenty  of  shells, 
while  in  the  lower  continental  boulder-clay  there 
are  scarcely  any  ? 

But  as  yet  our  knowledge  of  the  conditions  of  life 
of  the  marine  mollusca  and  of  their  distribution  is 
extremely  scanty.  We  are  apt  to  imagine  that  the 
bottom  of  the  sea  is  covered  by  a  more  or  less 
uniform  thick  layer  of  shells;  but  whenever  a  careful 
survey  of  the  nature  of  the  deposits  now  forming 
there  has  been  made,  such  is  by  no  means  found  to 
be  the  case.  Some  of  the  best  results  obtained  by 
that  useful  body,  the  Liverpool  Marine  Biological 
Committee,  have  been  precisely  in  this  direction. 
A  most  interesting  account  has  been  published  by 
Professor  Herd  man  and  Mr.  Lomas  on  the  floor 
deposits  of  the  Irish  Sea,  in  which  the  authors  state 
(p.  217),  that  "a  place  may  be  swarming  with  life 
and  yet  leave  no  trace  of  anything  capable  of  being 
preserved  in  the  fossil  state,  whereas  in  other  places, 
barren  of  living  things,  banks  of  drifted  and  dead 
shells  may  be  found,  and  remain  as  a  permanent 
deposit  on  the  ocean  floor." 

Owing   to   the    fact   of  the   peculiar   geographical 


THE   SIBERIAN    MIGRATION.  237 

position  of  Scandinavia  at  this  time — an  isthmus  of 
land  with  a  high  mountain  range  lying  between  the 
warm  Atlantic  and  the  cold  Arctic  Sea — the  snowfall 
must  have  been  excessive,  and  large  glaciers  were 
evidently  forming.  These  produced  icebergs  as  soon 
as  the  lower  parts  had  advanced  to  the  Baltic  coast- 
land  and  deposited  their  detritus  in  the  sea.  Immense 
masses  of  mud  and  stones  were  thus  cast  to  the 
bottom  of  the  sea,  and  under  these  circumstances  no 
delicate  mollusca  or  other  marine  life  probably  could 
have  developed  within  a  considerable  distance  from 
the  shore.  To  judge  from  the  direction  pursued  by 
the  majority  of  the  boulders  from  their  source  of 
origin,  the  prevailing  current  during  the  deposition  of 
the  lower  boulder-clay  was  from  north-west  to  south- 
east. It  is  possible  that  little  marine  life,  except 
free-swimming  forms,  would  have  been  able  to  live 
within  the  Russian  area  of  this  sea.  But  the  free- 
swimming  larvae  of  molluscs  and  other  surface 
species  were  not  prevented  from  passing  from  the 
White  Sea  south-westward,  and  in  sheltered  localities 
where  little  or  no  mud  deposition  was  going  on, 
these  no  doubt  might  have  developed  into  adults 
on  the  sea-floor.  It  is  quite  conceivable,  therefore, 
that  in  one  portion  of  the  North  European  Sea,  which 
was  fully  exposed  to  the  destructive  influences  of  the 
iceberg  action,  the  fauna  was  scanty  or  totally  absent, 
while  in  another  part  there  lived  a  fairly  abundant 
one.  The  unfossiliferous  state  of  the  lower  continental 
boulder-clay  does  not,  therefore,  offer  any  serious 


238         HISTORY    OF   THE   EUROPEAN    FAUNA. 

difficulty  to  the  supposition  that  some  of  the  so- 
called  Newer  Pliocene  Crags  of  the  east  coast  of 
England  were  laid  down  at  the  same  time  by  the 
same  sea. 

This  would  also  explain  how  the  Arctic  species 
come  to  inhabit  the  Caspian,  as  the  old  Aralo-Caspian 
Sea  could  have  had  some  communication  (Fig.  12, 
p.  156)  with  the  North  European  Sea.  And  this  again 
offers  an  explanation  of  the  otherwise  mysterious 
occurrence  of  the  Caspian  Dreyssensia  polymorpJia 
in  the  lower  continental  boulder-clay. 

The  climatic  reasons  for  the  supposition  that  the 
boulder-clay  is  a  marine  deposit  have  already  been 
given  (p.  66).  However,  it  may  be  asked  what  about 
the  glacial  flora  which  has  been  proved  to  have  existed 
all  over  the  plains  of  Northern  Europe  ? — what  about 
the  relics  of  this  same  flora  which  still  linger  on  in  a 
few  localities  to  the  great  delight  of  the  systematic 
botanist  ?  They  have  been  spoken  of  as  indications  of 
a  former  Arctic  climate  in  Europe.  The  presence  of 
an  Arctic  species  such  as  Dryas  octopetala  in  any  of 
the  pleistocene  deposits  is  often  looked  upon  as  an 
absolute  proof  of  a  very  severe  climate  having  pre- 
vailed at  the  time  they  were  laid  down.  Professor 
Geikie  tells  us  that  the  South  of  England  was 
clothed  with  an  Arctic  flora,  when  the  climate  be- 
came somewhat  less  severe  than  it  had  been  during 
the  climax  of  the  glacial  cold  (p.  398).  Relics  of 
such  a  flora  have  been  detected  at  Bovey  Tracey,  in 
Devonshire,  the  Arctic  plants  found  comprising 


THE  SIBERIAN    MIGRATION.  239 

Betnla  nana  and  B.  alba,  Salix  cinerea  and  Arctosta- 
phylos  uva-ursi. 

Now  three  of  these  four  species  of  plants  are  still 
natives  in  the  British  Islands,  and  all  are  forms 
which  probably  came  to  us  with  the  Arctic  migration 
which  I  described  in  Chapter  IV.  They  travelled 
south  with  the  reindeer,  or  before  it,  and  may  have 
covered  large  tracts  of  country  at  the  time.  With  the 
increased  struggle  for  existence  on  the  arrival  of  the 
Siberian  and  Oriental  migrants,  they  have  probably 
been  evicted  by  these  more  powerful  rivals.  A 
discovery  of  their  remains  does  not  necessarily 
indicate  that  a  great  change  of  climate  has  taken 
place  since  they  lived  in  the  country.  And  certainly 
these  Arctic  plants  cannot  be  taken  as  indicating  a 
low  temperature,  for  it  has  been  shown  that  Alpine 
plants  are  mostly  intolerant  of  very  low  temperatures. 
"  Arctic  and  Alpine  species  in  the  Botanical  Gardens 
at  Christiania,"  says  Professor  Blytt  (p.  19),  "endure 
the  strongest  summer  heat  without  injury,  while  they 
arc  often  destroyed  when  not  sufficiently  covered 
during  the  winter."  Similar  observations  have  been 
made  in  other  countries.  For  this  reason  they  have 
to  be  generally  wintered  in  frames  in  the  Botanic 
Gardens  at  Kew  and  Dublin,  and  are  thus  exposed  to 
higher  temperatures  than  at  present  obtain  in  the 
British  Islands.  This  fact  suggests  that  the  Alpine 
and  Arctic  plants  really  did  not  originate  in  countries 
with  cold  temperatures.  They  probably  made  their 
first  appearance  long  before  the  Glacial  period — 


240         HISTORY   OF   THE    EUROPEAN    FAUNA. 

perhaps  in  early  Tertiary  times — chiefly  in  the  Arctic 
Regions,  which  at  that  time  had  a  mild  climate. 
They  have  since  become  adapted  to  live  in  cold 
countries  where  they  flourish,  provided  they  receive 
sufficient  moisture  in  the  summer,  and  are  protected 
from  severe  frost  in  the  winter  by  a  covering  of  snow. 

When  we  carefully  examine  the  present  range 
of  Arctic  plants  in  the  British  Islands,  a  curious 
fact  presents  itself  which  no  doubt  has  frequently 
been  noted  by  botanists,  viz.,  that  some  of  the  most 
characteristically  Arctic  species,  and  some  which 
are  often  quoted  by  glacialists  in  support  of  their 
theories,  flourish  at  the  present  moment  in  very 
mild  situations.  I  have  already  referred  to  the  fact 
that  the  Mountain  Avens  (Dryas  octopetala)  abounds 
in  the  west  of  Ireland  (County  Galway)  down  to  sea- 
level.  Now  it  is  well  known  that  the  mean  winter 
temperature  of  that  part  of  Ireland  resembles  that 
of  Southern  Europe,  being  no  less  than  12°  F. 
(  =  7°  Cent.)  above  freezing  point.  The  plant,  of 
course,  is  here  a  native,  and  not  introduced.  This 
instance  shows  clearly,  that  as  long  as  more  vigorous 
competitors  are  absent,  and  as  long  as  it  is  not 
exposed  to  severe  frost  or  undue  dryness,  this  and 
allied  species  do  just  as  well  in  a  mild  climate  as  in 
their  native  Arctic  home. 

In  his  interesting  essay  on  the  distribution  of  the 
Arctic  plants  in  Europe  during  the  Glacial  period, 
Professor  Nathorst  adduces  the  fact  that  all  the 
localities  but  one,  in  which  remains  of  such  plants 


THE  SIBERIAN   MIGRATION.  241 

have  been  discovered,  lie  either  within  or  close  to 
the  limits  of  the  maximum  extension  of  the  supposed 
northern  ice-sheet,  or  within  those  of  the  former 
Alpine  glaciers.  Whether  we  look  upon  the  boulder- 
clay  as  a  marine  or  a  terrestrial  product,  it  is  quite 
conceivable  that,  in  many  instances,  the  remains  of 
the  Arctic  plants  may  have  been  carried  by  ice  to 
great  distances  from  where  they  grew.  The  prob- 
ability, however,  is  in  favour  of  most  of  them 
having  lived  where  their  remains  are  now  found. 
Now,  it  is  a  remarkable  fact,  that  the  single  instance 
in  Europe  of  a  deposit  of  Arctic  plants  having  been 
found  far  removed  from  the  maximum  extension 
of  the  northern  ice-sheet  is  the  one  quoted  above, 
viz.,  at  Bovey  Tracey,  in  Devonshire.  Even  up 
to  recent  times  Arctic  plants  may  have  persisted  at 
Bovey  Tracey  just  as  they  do  in  Gal  way  under  the 
influence  of  a  mild  coast  climate.  Similar  circum- 
stances may  have  led  to  their  survival  along  the 
shores  of  the  sea  which  deposited  the  North 
European  boulder-clay,  while  they  moved  north- 
ward from  the  Alps  along  with  the  glaciers,  which 
always  supplied  them  with  an  abundance  of  moisture. 
Alpine  plants  probably  became  exterminated  in  the 
plain  of  Central  Europe  at  a  much  earlier  period. 


242         HISTORY  OF   THE   EUROPEAN   FAUNA. 


SUMMARY  OF  CHAPTER  V. 

What  has  been  spoken  of  in  the  earlier  parts  of  this  book  as  the 
eastern  migration,  refers  in  a  general  way  to  the  animals  which 
have  come  to  England  from  the  east.  But  these  are  by  no  means 
natives  of  one  country  alone.  We  can  trace  a  number  of  the 
British  mammals  to  a  Siberian  origin,  and  also  some  birds; 
among  many  of  the  lower  vertebrates  and  invertebrates,  how- 
ever, there  are  few  species  which  have  reached  us  from  Siberia. 
They  may  have  had  their  original  homes  in  the  Alps,  in  Eastern 
Europe,  or  in  Central  and  Southern  Asia,  and  have  joined  in 
their  westward  course  the  later,  more  quickly  travelling 
mammals.  Many  instances  are  given  from  all  the  more  im- 
portant groups  of  animals  to  show  how  we  may  proceed  in 
approximately  identifying  the  home  of  a  species. 

The  periodical  invasion  into  our  continent  of  Pallas's  Sand- 
grouse  and  other  birds,  suggests  an  explanation  as  to  the  cause 
of  the  great  westward  migration  in  former  times  of  the  Siberian 
mammals.  Since  a  considerable  amount  of  fossil  evidence  is 
available  to  show  the  path  of  migration  pursued  by  these 
mammals,  other  important  problems,  such  as  the  time  of  their 
arrival  in  Europe  and  the  geographical  conditions  surrounding 
them,  may  perhaps  be  approximately  ascertained,  and  thus 
throw  much  light  on  the  general  features  of  the  European 
fauna.  It  has  been  proved  by  Professor  Nehring  that  the 
Siberian  mammals  arrived  in  Eastern  Europe  after  the  deposi- 
tion of  the  lower  continental  boulder-clay.  He  believes 
that  the  climate  of  Germany  at  that  time  had  ameliorated  so 
far,  after  the  maximum  cold  of  the  Glacial  period,  that  steppes 
with  a  Siberian  fauna  could  exist.  Other  groups,  such  as  the 
Mollusca,  however,  do  not  support  Professor  Nehring's  theory, 
and  in  order  to  arrive  at  an  independent  solution  of  this  and  the 
other  problems  referred  to,  a  short  history  is  given  of  the  Siberian 


THE   SIBERIAN   MIGRATION.  243 

fauna.  Recent  geological  ages  have  witnessed  the  arrival  in 
Southern  Europe  of  mammals  now  almost  confined  to  the  arctic 
and  subarctic  regions.  In  Siberia,  on  the  other  hand,  many 
southern  species  penetrated,  apparently  about  the  same  time, 
to  the  extreme  northern  limits  of  that  country.  The  greatest 
authority  on  the  Siberian  fossil  fauna,  Tcherski,  believes  that 
this  took  place  in  pliocene  times,  the  gradual  retreat  occupy- 
ing the  whole  of  the  Glacial  period.  If  this  were  correct,  the 
retreat  from  the  Arctic  Regions  would  have  occurred  at 
the  same  time  when,  according  to  our  European  authorities, 
Professors  Nehring  and  Geikie,  the  much  more  southern 
parts  of  our  continent  were  already  uninhabitable.  But 
Siberia  could  not  have  supported  the  large  mammals  at  all 
at  a  time  when  Europe  was  uninhabitable,  as  it  would  be 
difficult  to  conceive  under  what  geographical  conditions  the 
climate  of  the  latter  was  arctic  and  that  of  the  former  temperate. 
If  the  whole  fauna  was  driven  into  Southern  Asia,  how  is  it  that 
the  Siberian  invasion  of  Europe  occurred  immediately  after  the 
deposition  of  the  lower  boulder-clay,  that  is  to  say,  after  the 
earlier  part  of  the  Glacial  period?  The  difficulty  can  be  met  by 
the  supposition  that  both  Europe  and  Siberia  had  a  temperate 
climate  at  that  time.  This  view  is  supported  by  certain 
evidences,  fully  described,  of  a  connection  between  the  Caspian 
and  the  White  Sea,  which  would  have  had  the  effect  of  influ- 
encing the  climate.  The  Siberian  fauna  would  thus  have  been 
prevented  from  spreading  westward  in  Pliocene  and  early 
Glacial  times.  But  on  the  disappearance  of  the  marine  con- 
nection, a  way  would  have  been  opened  into  our  continent, 
which  again  had  an  effect  on  the  climate.  The  latter  would 
have  become  sensibly  colder  and  thus  have  reduced  the 
habitable  area  of  the  Siberian  fauna. 

Such  geographical  conditions  would  have  been  incompatible 
with  a  great  northern  mer  de  glace,  and  the  boulder-clay  in 
Northern  Europe  could  not  have  represented  a  ground  moraine 
but  is  a  marine  deposit.  The  sea  is  supposed  to  have  covered 


244         HISTORY  OF   THE   EUROPEAN   FAUNA. 

the  Northern  Russian  and  German  plains,  and  into  it  icebergs 
discharged  the  detritus  which  had  accumulated  on  them  when 
they  were  still  Scandinavian  glaciers. 

As  regards  the  time  of  the  arrival  of  the  Siberian  migrants  in 
Europe,  the  English  Forest-Bed  gives  us  an  additional  clue  to 
its  determination.  Since  Siberian  migrants  are  unknown  from 
earlier  deposits  than  this,  it  is  reasonable  to  suppose  that  they 
arrived  in  England  about  the  time  when  it  was  laid  down.  But 
since  they  appear  in  Germany  in  the  inter-glacial  beds  subse- 
quent to  the  deposition  of  the  lower  boulder-clay,  the  former  are 
probably  contemporaneous  with  the  Forest-Bed.  Some  of  the 
deposits  generally  regarded  as  upper  pliocene  by  British  geolo- 
gists would  therefore  have  to  be  classed  with  the  lower 
continental  boulder-clay  as  lower  pleistocene.  In  connection 
with  this  theory  some  interesting  faunistic  data  are  given 
which  seem  to  support  it. 

In  conclusion,  the  former  presence  of  Arctic  plants  in  Central 
Europe  and  their  bearing  on  the  climatic  problems  are 
discussed. 


CHAPTER  VI. 

THE  ORIENTAL   MIGRATION. 

THE  Oriental  migration  is  closely  related  to  the 
Siberian.  Both  have  originated  within  the  Asiatic 
continent,  and  in  many  respects  a  strict  line  cannot 
be  drawn  between  them.  There  can  be  no  doubt 
that  some  of  the  species  which  we  regard  as  Siberian 
migrants  had  their  original  home  in  more  southern 
latitudes,  and  thus  may  have  formed  part  of  the 
older  Oriental  migration.  The  home  of  that  migra- 
tion I  take  to  be  Central  and  Southern  Asia,  that  is 
to  say,  everything  south  of  the  Altai  Mountains  and 
the  Caucasus.  Its  members  have  reached  Europe 
across  an  old  land-connection  which  united  Turkey, 
Greece,  and  Syria,  while  the  Siberian  animals  in- 
vaded our  continent  to  the  north  of  the  Caspian  and 
Caucasus. 

The  Siberian  immigrants  into  Europe  on  the  whole 
are  not  very  numerous,  but  it  is  different  with  those 
from  the  more  southern  parts  of  the  Asiatic  con- 
tinent. The  members  of  the  Oriental  migration 
form  a  very  large  percentage  of  the  European  fauna. 
No  other  migration  has  affected  our  continent  so 
powerfully,  because  it  continued  uninterruptedly  for 

245 


246         HISTORY  OF   THE   EUROPEAN    FAUNA. 

a  very  long  time.  Hence  its  results  can  be  traced 
from  one  corner  of  Europe  to  the  other.  We  have 
seen  that  the  Siberian  migration  only  commenced 
after  the  first  portion  of  the  Glacial  period  had  passed 
away.  The  Oriental,  however,  persisted  throughout, 
or  at  any  rate  for  the  greater  part  of  that  period.  It 
commenced  ages  before  it,  in  miocene  times,  or  even 
earlier.  And  as  the  ^Egean  Sea,  which  broke  up  the 
highway  of  the  Oriental  migrants,  is  only  of  recent 
formation,  there  was  a  steady  westward  march  for  a 
very  considerable  time.  No  doubt  the  migration  was 
also  favoured  by  the  fact  that  scarcely  any  formidable 
barriers  had  to  be  crossed. 

Many  instances  might  be  quoted  of  the  same 
species  forming  part  of  the  Oriental  and  also  of  the 
Siberian  migration,  but  as  a  rule  the  Siberian  mi- 
grant belongs  to  a  distinct  variety,  or  has  such  well- 
marked  racial  characters  as  to  be  at  once  detected 
from  its  more  southern  relative.  Among  the  ex- 
amples of  Oriental  migrants  which  I  have  occasion 
to  bring  forward,  such  instances  will  be  specially 
dealt  with. 

In  its  wild  state  the  Red  Deer  (Cervus  elaphus)  is 
almost  extinct  in  the  British  Islands,  though  it  still 
occurs  in  the  moorlands  of  Devonshire  and  Somerset- 
shire in  England,  in  the  south-west  of  Ireland,  and 
in  some  localities  in  Scotland.  Fifty  years  ago  it  was 
also  found  wild  in  several  other  of  the  Irish  western 
counties ;  and  in  the  seventeenth  century  it  was 
common  in  most  of  the  mountainous  districts  of 


THE   ORIENTAL   MIGRATION.  247 

Ireland.  Its  remains  have  been  found  fossil  in  the 
marls  and  caves  of  Ireland,  and  in  the  Forest-Bed, 
as  well  as  in  a  large  number  of  caves  in  England. 
The  history  of  the  Red  Deer  in  other  countries 
is  very  similar.  In  Scandinavia  it  flourished  as 
far  north  as  the  sixty-eighth  degree  of  latitude, 
whereas  it  is  now  quite  extinct  on  the  main- 
land, though  still  lingering  on  in  some  of  the 
western  islands.  Denmark  and  Switzerland  know 
it  no  more,  and  it  is  almost  extinct  in  Belgium. 
Nearly  throughout  Europe  where  it  occurs,  its 
numbers  are  diminishing,  greatly  owing,  perhaps,  to 
the  relentless  persecution  by  man,  but  its  gradual 
disappearance  must  likewise  be  partly  due  to  other 
causes.  Formerly  it  inhabited  every  country  of 
Europe  and  all  the  larger  islands.  It  still  exists 
in  Corsica  and  Sardinia,  and  at  an  earlier  period 
it  was  also  met  with  on  the  island  of  Malta. 
The  Red  Deer  found  in  Corsica  and  Sardinia  is 
smaller  than  that  inhabiting  Central  Europe,  and  is 
by  some  authorities  regarded  as  a  distinct  species, 
which  has  been  named  Cervus  corsicanus.  But 
Sir  Victor  Brooke  has  pointed  out  that  the  antlers 
of  some  of  the  Scotch  Deer  agree  in  every  point 
with  those  of  the  Sardinian  species.  Indeed,  the 
West  European  Red  Deer  altogether  is  a  small- 
antlered  form,  compared  with  the  Eastern  one.  This 
character,  however,  is  only  a  racial  one,  and  not  of 
specific  value.  In  the  pleistocene  deposits  of  Eastern 
and  Central  Europe,  a  very  large-antlered  race  has 


248         HISTORY  OF   THE   EUROPEAN   FAUNA. 

been  discovered,  and  identified  by  Professor  Nehring 
with  Ceruus  canadensis — the  Canadian  Red  Deer. 
Tcherski,  the  Siberian  traveller,  believed  that  Cervus 
canadensis  was  identical  with,  or  a  variety  of,  the 
Asiatic  species  of  Deer,  Cervus  eustephanus,  Cervus 
xanthopygus,  and  Cervus  maral.  Some  authorities 
— and  to  these-belong  Mr.  Lydekker — think  that  we 
ought  perhaps  to  regard  the  whole  number  of  Red 
Deer-like  forms  as  local  varieties  of  one  widely- 
spread  species.  Besides  the  deer  already  referred  to, 
the  following  belong  to  this  same  group: — Cervus 
cashmirianuS)  Cervus  affinis,  Cervus  Roosvelti,  from 
North  America,  and  the  North  African  Cervus  bar- 
barus. 

The  question  now  is,  where  have  these  varieties 
originated  ?  Or,  if  we  go  to  the  root  of  the  matter, 
where  is  the  original  home  of  their  ancestors?  Con- 
sidering that  so  many  Cervidcz  have  been  found  in 
French  and  English  pliocene  deposits,  and  that 
remains  of  the  Red  Deer  occur  not  only  in  the 
English  Forest-Bed,  but  have  been  found  associated 
with  those  of  the  Pigmy  Hippopotamus  in  Malta, 
it  would  only  be  reasonable  to  suppose  that  the 
genus  Cervus  had  originated  in  Europe.  It  might 
also  be  argued  with  equal  force  that  the  Red  Deer 
had  its  birthplace  in  our  continent.  But  when  we 
carefully  study  its  present  range  this  verdict  cannot 
be  accepted.  The  view  of  the  Asiatic  origin  of  the 
Red  Deer,  so  ably  maintained  by  Koppen,  cor- 
responds far  better  with  its  present  distribution, 


THE   ORIENTAL  MIGRATION.  249 

especially  if  we  look  upon  the  Asiatic,  North 
American,  and  North  African  forms  as  varieties  of 
the  same  species. 

If  the  Red  Deer  were  of  European  origin,  it  must 
have  come  into  existence  at  a  time  when  Malta  was 
part  of  the  mainland,  when  North  Africa  and  the 
British  Islands  were  connected  with  the  continent  of 
Europe,  and  of  course  before  the  deposition  of  the 
Forest-Bed.  Such  land-connections  existed  probably 
during  the  Pliocene  Epoch.  Migrants  would  have 
wandered  from  Europe  into  Asia.  These  would 
have  developed  into  larger  races,  which  again 
furnished  emigrants  for  North  America.  The  latter 
crossed  by  the  old  land-connection  which  once  joined 
America  and  Asia  at  Behring's  Straits.  During 
pleistocene  times  the  large  Siberian  race  would  now 
have  re-migrated  to  the  home  of  its  ancestors  in 
Europe,  for  we  find  the  remains  only  in  Central 
and  Eastern  Europe,  indicating  that  an  invasion  of 
the  Red  Deer  from  Asia  must  then  have  taken 
place. 

Against  this  view  of  the  European  origin  of  the 
Red  Deer,  it  may  be  urged  that  deer  are  known  from 
Indian  as  well  as  from  European  pliocene  deposits, 
and  that  a  migration  could  have  taken  place  from 
the  Oriental  Region  to  Europe  just  as  easily  as 
from  the  latter  to  Asia.  The  majority  of  the  species 
of  the  genus  Ceruus  (in  a  wide  sense),  moreover, 
are  Asiatic,  ranging  to  Borneo,  Sumatra,  and  the 
Philippine  Islands,  all  of  which  islands  have  been 


250         HISTORY  OF  THE   EUROPEAN    FAUNA. 

separated  from  the  mainland  for  a  considerable  time. 
Finally,  the  original  home  of  a  species,  as  we  have 
learned,  generally  corresponds  with  the  centre  of  its 
geographical  range,  and  this  lies  in  the  case  of  the 
Red  Deer  in  Central  Asia. 

One  of  the  highest  authorities  on  the  deer  family, 
Sir  Victor  Brooke,  also  was  of  opinion  that  the 
Cervida  originated  in  Asia,  and  from  there  spread 
east  and  west  Of  the  two  divisions  into  which  true 
deer  are  divided,  viz.,  the  Plesiometacarpalia  and 
the  Telemetacarpalia,  the  former  is  almost  confined  to 
the  Old  and  the  latter  to  the  New  World.  The  only 
North  American  species  belonging  to  the  first 
division  is  the  Canadian  Red  Deer,  which  fact 
clearly  indicates  its  recent  immigration  to  that 
continent. 

There  were  probably  two  distinct  migrations  of 
the  Red  Deer  into  Europe.  An  older  one  coming 
from  Asia  Minor  into  Greece,  which  stocked  Sardinia, 
Corsica,  Malta,  and  North  Africa  in  the  first  place, 
when  these  were  still  connected  with  one  another. 
This  same  migration  likewise  affected  western  con- 
tinental Europe,  the  Irish  Red  Deer  being  probably 
the  descendant  of  this  very  ancient  stock.  The 
latter  entered  the  island  when  it  was  still  part  of  the 
Continent.  The  later  migration  of  a  larger  form  came 
from  Siberia  and  spread  mainly  over  Eastern  and 
Central  Europe,  but  it  appears  that  it  also  reached 
England,  although  there  is  no  evidence  of  any  of 
these  Siberian  deer  having  ever  inhabited  Ireland. 


THE  ORIENTAL  MIGRATION.  251 

The  range  of  this  deer,  therefore,  to  some  extent 
corresponds  to  that  of  another  described  on  p.  153. 
We  found  then  that  two  races  of  Reindeer  had 
migrated  to  the  British  Islands — one  from  the  Arctic 
Regions,  and  the  other  from  Siberia,  but  that  only 
the  former  had  reached  Ireland. 

The  so-called  Irish  Elk  (Cervus  giganteus)  has 
been  referred  to  the  Oriental  migration,  but,  as 
stated  below,  it  has  some  claims  to  be  regarded 
as  a  European.  Unfortunately  it  is  now  extinct; 
it  seems  not  unlikely,  however,  that  it  inhabited 
Ireland  when  man  had  already  made  his  appear- 
ance on  the  island.  Although  its  remains  are  found 
in  such  extraordinary  abundance  in  Ireland,  it 
certainly  did  not  originate  there.  It  lived  also 
in  England  and  Scotland,  and  in  the  Isle  of  Man, 
in  France,  Denmark,  Germany,  Austria,  North  Italy, 
and  Russia.  Its  remains  have  been  discovered 
even  in  Siberia.  It  must  either  have  originated 
in  Europe  and  then  migrated  to  Asia,  or  have  had  its 
birthplace  in  Asia  and  wandered  to  Europe.  There 
is  nothing  to  lead  any  one  to  assert  positively  that 
either  of  these  two  continents  was  the  one  in  which 
the  original  home  of  the  Irish  Elk  was  situated,  and 
we  can  only  be  guided  in  this  case  by  the  history  of 
its  nearest  relatives.  These  are  the  Fallow  Ueer 
(Cervus  dama).  There  are  two  very  closely  allied 
species,  the  Persian  and  the  European,  but  several 
others  have  been  discovered  in  the  Forest-Bed  and 
the  pliocene  deposits  of  the  Auvergne.  As  no 


252         HISTORY  OF   THE   EUROPEAN    FAUNA. 

remains  of  the  Fallow  Deer  are  known  from  Asia, 
it  seems  probable  that  it  and  also  the  Irish  Elk 
originated  in  Southern  Europe,  and  only  invaded 
Asia  in  early  pleistocene  times. 

The  Mammoth  (Elephas  primigenius)  is  a  familiar 
example  among  a  large  number  of  mammals  which 
have  come  to  us  about  the  same  time  from  Asia  by 
the  Asia  Minor  route.  It  had  a  much  wider  range 
than  the'  Irish  Elk,  since  its  remains  have  been  dis- 
covered in  a  large  number  of  European  localities  as 
far  west  as  Ireland,  also  in  Siberia,  and  even  North 
America.  Though  we  have  had  Proboscidea  in  Europe 
from  the  Middle  Miocene  onwards,  Mr.  Lydekker 
(d,  p.  viii.)  holds  that  "  our  comparatively  full  know- 
ledge of  Lower  Miocene  and  Upper  Eocene  mam- 
malian faunas  of  the  greater  part  of  Europe  and 
North  America,  renders  it  almost  certain  that  neither 
of  those  regions  was  the  home  of  the  direct  ancestors 
of  the  Elephantidce ;  and  we  must  therefore  look 
forward  to  the  discovery  of  marnmaliferous  Lower 
Miocene  or  Upper  Eocene  strata  in  some  other 
region  of  the  (probably  old)  world  which  may  yield 
these  missing  forms." 

The  genus  Elephas  makes  its  first  appearance  in 
the  Upper  Miocene  of  India.  Our  European  E. 
antiquus  is,  according  to  Professor  Zittel,  probably 
identical  with  E.  armeniacus  of  Asia  Minor,  while 
E.  meridionalis  agrees  in  all  essential  characters 
with  the  Indian  E.  hysudricus.  The  Indian  and 
European  species  of  fossil  elephants  altogether  are 


THE  ORIENTAL  MIGRATION.  253 

very  closely  related,  and  the  supposition  that  they 
all  have  had  their  original  home  in  the  Oriental  Region 
offers,  I  think,  no  serious  obstacle.  The  view  of  the 
European  origin  of  the  mammoth  especially  is  open 
to  very  serious  objections.  It  does  not  occur  in  any 
European  pliocene  deposits,  and  could  not  therefore 
have  originated  in  our  Continent  until  pleistocene 
times.  That  it  should  then  have  commenced  its 
travels  through  Europe  and  Siberia  to  the  New 
Siberian  Islands  and  North  America  seems  almost 
an  impossibility.  But  if  we  suppose  the  mammoth 
to  have  had  its  home  in  India  in  pliocene  times,  it 
could  then  easily  have  migrated  to  all  the  parts  of 
the  world  where  its  remains  have  been  discovered. 

Of  the  Asiatic  mammals  still  living,  some  have 
only  just  crossed  the  borders  of  Europe  and  then 
died  out  again.  Similar  cases  have  been  referred  to 
in  discussing  the  Siberian  migration.  Thus  remains 
of  the  camel  have  been  found  in  Roumania  and  in 
Southern  Russia  in  pleistocene  deposits.  Others  have 
lingered  on  to  the  present  day.  Crocidura  etrusca, 
for  instance,  still  lives  in  Southern  France,  Italy, 
Sicily,  and  North-western  Africa.  All  its  nearest 
relations  are  typically  Oriental  species.  In  spite  of 
the  fact  that  a  Crocidura  is  known  from  French  and 
German  miocene  deposits,  the  general  range  of  the 
genus  suggests  an  Oriental  origin.  In  early  Tertiary 
times  a  section  spread  into  African  territory  and 
another  eastward  as  far  as  the  island  of  Timor.  This 
may  possibly  have  happened  in  miocene  times,  when 


254         HISTORY   OF   THE   EUROPEAN   FAUNA. 

a  few  species  likewise  found  their  way  into  Europe. 
Many  other  mammals  have  wandered  still  farther 
west,  and  now  form  an  important  percentage  of  the 
European  fauna. 

Of  Birds,  too,  a  large  number  might  be  mentioned 
which  had  their  home  in  Asia  and  have  found  their 
way  to  Europe  with  the  Oriental  migrants.  A  few 
instances  have  already  been  alluded  to,  and  somq 
additional  ones  may  be  specified  at  random,  without 
attempting  to  give  a  complete  list. 

Some  of  the  Wagtails  (Motacilla),  as  I  men- 
tioned in  the  last  chapter,  have  certainly  come  to 
us  with  the  Siberian  migration;  but  others  seem 
to  be  Oriental,  such  as  Motacilla  melanope,  which  is 
resident  in  Southern  Europe  and  migratory  in  the 
North.  M.  campestris — the  Yellow  Wagtail — has 
a  most  peculiar  discontinuous  range.  One  colony 
breeds  in  the  British  Isles  and  Western  Europe 
generally,  where  it  is  known  as  a  summer  visitor, 
retiring  to  WTest  Africa  during  winter;  another  is 
found  from  South-east  Russia  to  Turkestan  in 
summer,  and  winters  in  Southern  Africa.  This  fact 
may  possibly  be  due  to  two  distinct  migrations  from 
Asia  having  taken  place:  an  earlier  one  from  the 
South-east — that  is  to  say,  an  Oriental  one — and 
a  Siberian  one  more  recently.  In  this  case  the 
members  of  the  two  migrations  have  not  become 
sufficiently  differentiated  to  be  regarded  as  distinct 
varieties.  Though  most  of  the  Wagtails  have  a 
somewhat  northern  range,  none  (except  perhaps  M. 


THE   ORIENTAL  MIGRATION.  255 

borealis)  are  truly  Arctic;  and  indeed,  as  almost  all  of 
them  pass  the  winter  in  southern  latitudes,  it  may 
be  assumed  that  they  are  of  southern  and  not  of 
northern  origin. 

The  Dippers  (Cinclus)  are  practically  unknown  in 
the  Central  European  plain,  but  they  occur  in 
Western  Europe  as  far  north  as  Scandinavia,  also 
in  the  Alps,  Carpathians,  and  Southern  Europe,  in- 
cluding Sicily  and  Sardinia.  Some  authorities  dis- 
tinguish three  species,  others  only  one.  As  a  matter 
of  fact,  the  difference  between  the  three  forms  is  very 
slight,  and  their  nests  and  eggs  are  undistinguishable. 
Eight  other  species  have  been  recognised,  and  all 
these  are  either  Asiatic  or  American.  As  one  of  the 
American  forms  is  peculiar  to  Peru  and  another  to 
Ecuador  and  Columbia,  and  since  the  genus  as  a 
whole  is  a  mountain-genus,  it  probably  is  an  ancient 
one.  Its  European  range  alone,  however,  implies  that 
it  has  inhabited  our  continent  for  a  considerable 
time  and  is  no  new-comer.  We  may  look  upon  it  as 
of  Asiatic  origin.  The  ancestors  have  spread  east 
and  west,  the  European  species  having  arrived  with 
the  earlier  Oriental  migrants,  and  wandered  along 
the  Mediterranean  at  a  time  when  the  geographical 
conditions  of  that  sea  were  vastly  different  from  what 
they  are  to-day. 

Not  quite  so  ancient  as  the  Dippers,  but  like- 
wise Asiatic  in  their  origin,  are  the  Bullfinches 
(Pyrrhida).  The  closely  allied  Pine-Grosbeak  {Pint- 
cola  enucleator]  has  already  been  referred  to  (p.  191) 


256         HISTORY  OF   THE   EUROPEAN   FAUNA. 

as  a  member  of  the  Siberian  migration.  The 
distribution  of  the  European  Bullfinch  (P.  europcea) 
is  very  interesting,  as  it  occurs  in  two  distinct 
forms,  by  some  authorities  regarded  as  races,  by 
others  as  species.  In  all  probability  these  two 
races  owe  their  origin  to  two  different  migrations 
from  the  same  ancestral  stock.  We  may  suppose 
that  P.  europcza  came  to  Europe  along  with  the 
Oriental  migration,  spreading  chiefly  over  the  south 
and  west,  while  another  branch  developed  in  Siberia 
into  the  larger  and  more  brilliant  race  (P.  major), 
which  subsequently  entered  the  neighbouring  con- 
tinent with  the  Siberian  fauna.  The  latter  race 
inhabits,  according  to  Mr  Saunders,  Northern  and 
Eastern  Europe,  and  also  Siberia.  All  the  other 
species — there  are  eight  more — except  one,  are  found 
in  Asia.  This  one  species,  which  inhabits  the  Azores, 
appears  to  be  more  closely  related  to  one  of  the 
Siberian  bullfinches  than  to  the  European.  It  stands 
isolated,  and  is  an  extraordinary  instance  of  discon- 
tinuous distribution,  as  no  Bullfinch  inhabits  either 
Madeira  or  the  Canary  Islands.  We  must  assume 
that  the  form  connecting  it  with  the  Asiatic  prob- 
ably lived  in  Southern  Europe,  and  has  become 
extinct. 

One  of  the  most  typically  Oriental  genera  of  birds 
is  Phaslanus,  to  which  our  Common  Pheasant  belongs. 
Out  of  twenty  species,  nineteen  are  found  exclusively 
in  Asia,  most  of  them  being  confined  to  the  central 
plateaux  of  that  continent.  Only  one  species  passes 


THE   ORIENTAL   MIGRATION.  257 

the  confines  of  Asia  into  Greece,  Turkey,  and 
Southern  Russia.  This  is  Phasianus  colchicus.  For- 
merly, however,  the  Pheasant  appears  to  have  had  a 
wider  range  in  Europe,  for  three  species  are  known 
fossil  from  France.  Altogether,  it  is  not  quite  certain 
whether  the  Pheasant  is  not  really  an  indigenous  bird 
in  the  British  Islands,  having  survived  from  pre- 
glacial  times.  It  is  believed  that  the  Romans  brought 
it  to  England,  but  there  is  no  record  of  an  introduc- 
tion at  that  time. 

Among  the  older  Oriental  bird  migrants  might  be 
mentioned  the  Fire-crested  WrQr\(Regutusignicapiltus)y 
which  has  even  occasionally  visited  England.  It  be- 
comes commoner  as  we  go  south-eastward.  In  Asia 
Minor  it  is  more  abundant  than  the  Gold-crest ;  and 
throughout  the  year  it  is  resident  in  Southern  Europe, 
where  it  occurs  in  Turkey,  Greece,  Italy,  Spain, 
Sardinia,  and  Malta.  On  the  opposite  shore,  in 
North-west  Africa,  it  again  makes  its  appearance, 
and  its  range  extends  westward  to  the  Canaries  (R. 
teneriffa)  and  Madeira  (R.  maderensis). 

The  genus  to  which  our  common  Goldfinch  belongs, 
viz.,  Carduelis,  is  also  probably  of  Oriental  origin,  and 
may  be  looked  upon  as  one  of  the  earlier  migrants. 
That  species  (C.  elegans)  breeds  throughout  Europe, 
except  in  the  extreme  north,  but  it  is  especially 
abundant  in  Southern  Europe  and  North-west  Africa. 
It  is  also  resident  in  Madeira  and  the  Canaries. 
Eastward  its  range  extends  to  Persia.  A  larger 
race  (C.  major)  inhabits  Western  Siberia  and  crosses 

17 


258         HISTORY   OF   THE   EUROPEAN    FAUNA. 

the  European  border  into  Russia.  It  interbreeds 
in  Siberia  with  C.  caniceps^  an  East  Siberian 
form. 

A  few  instances  of  Reptiles  and  Amphibia  with  a 
similar  range  will  show  that  the  Oriental  migration 
was  not  confined  to  the  higher  vertebrates. 

Two  species  of  the  genus  Eremias  (fodarcis]  occur 
in  South-eastern  Europe.  This  is  a  genus  of  Lizards 
with  rather  a  wide  distribution,  ranging  from  Central 
Asia  to  South  Africa  southward  and  China  eastward. 
Altogether  there  are  twenty-four  species,  two  of  which 
just  enter  Europe;  and  of  the  rest  half  are  Asiatic 
and  half  African.  Even  if  the  genus  were  of  African 
origin,  it  is  extremely  unlikely  that  the  Asiatic 
species  came  by  way  of  Europe.  We  may  assume, 
therefore,  with  a  fair  degree  of  probability  that  the 
two  European  species  wandered  westward  along  with 
the  Oriental  migrants. 

The  genus  Ablepharns  belongs  to  a  family  of 
Lizards  in  which  the  legs  are  either  very  fully 
developed,  or  quite  absent  as  in  the  Slow-worm 
(Anguis  fragilis).  It  is  an  ancient  genus,  having  a 
wide  range  from  Central  Asia  to  Australia  on  the  one 
hand,  and  to  South  Africa  on  the  other.  One  species 
of  this  Scink-like  Lizard,  viz.,  Ablepharus  pannonicus, 
enters  Europe  in  the  south-east,  inhabiting  Greece  as 
far  north  as  Southern  Hungary.  In  Asia  it  is  found 
in  Syria  and  North  Arabia.  This  clearly  signifies 
that  the  Lizard  is  an  Oriental  migrant. 

Among    the    Snakes    which    participated    in    the 


THE  ORIENTAL  MIGRATION.  259 

Oriental  migration  might  be  mentioned  Eryx  jaculus> 
whose  home  is  probably  in  Western  Asia.  It  is 
known  in  Europe  from  the  Greek  islands  of  Tinos 
and  Naxos,  from  Turkey  and  Southern  Russia. 
Another,  a  peculiar  worm-like  form,  lives  under- 
ground in  damp  earth  and  under  stones — Typhlops 
lumbricalis.  This  species  inhabits  the  mainland  of 
Greece  as  well  as  the  Greek  islands,  and  Asia  Minor 
as  far  as  the  Caucasus. 

A  most  interesting  case  of  distribution  is  that  of 
the  pretty  little  Toad  so  well  known  on  the  Continent 
under  the  name  of  "  fire-toad  "  (Bombinator  igneus). 
Though  some  authorities,  such  as  Boulenger,  recog- 
nise only  one  form  of  Bombinator^  others  are  of 
opinion  that  two  well-marked  varieties  exist  in 
Europe.  These  are  looked  upon  by  Dr.  von  Bedriaga 
as  good  species,  but  he  acknowledges  that  they  are 
rather  critical  and  difficult  to  identify.  No  other 
species  of  Bombinator  occur  in  Europe.  Bombinator 
pachypus,  the  western  race, — or  if  we  choose  to  call  it 
species, — occurs  in  France,  Germany,  Switzerland, 
Austria,  Sicily,  and  Greece.  B.  igneus — the  eastern 
race — is  found  in  Southern  Sweden,  Denmark,  Ger- 
many, Austria,  and  Russia.  The  latter  has  therefore 
a  more  northerly  and  easterly  range.  The  species  is 
not  known  from  Siberia,  but  makes  its  appearance 
again  in  China  in  a  form  which,  according  to  Dr.  von 

1  Since  writing  the  above  account,  Mr.  Boulenger,  in  his  new  work 
on  the  Batrachia  of  Europe,  has  accepted  the  specific  distinctions 
between  the  two  fire-toads. 


26O         HISTORY   OF   THE   EUROPEAN    FAUNA. 

Bedriaga,  docs  not  quite  agree  with  either  of  the  two 
European  races. 

Now  if  we  supposed  Bombinator  to  have  originated 
in  Europe,  its  absence  from  the  British  Islands,  most 
of  the  Mediterranean  islands,  and  the  greater  part 
of  Scandinavia  would  not  be  easy  of  explanation, 
while  as  an  Asiatic  migrant  the  European  range  is 
more  readily  understood.  Its  apparent  absence  from 
Western  Asia  might  quite  likely  be  due  to  the  fact 
that  the  zoology  of  that  part  of  the  Continent  is  only 
now  being  investigated.  The  latter  has,  moreover, 
undergone  great  physical  changes  in  recent  geological 
times.  The  supposition  that  one  migration  of  Bom- 
binator from  the  south-east  has  taken  place,  and 
then  another  from  the  east,  seems  to  explain  this 
case  of  distribution,  as  other  similar  ones,  in  a  most 
satisfactory  manner. 

The  Tree-Frog  (Hyla  arborea)  must  be  an  ancient 
species,  but  it  is  not  of  European  origin.  Few  genera 
of  Amphibia  have  a  wider  distribution  than  Hyla. 
There  are  only  three  species  in  Asia,  Europe,  and 
Africa,  the  remaining  129  being  confined  to  America 
and  Australia.  Two  of  the  three  Old  World  Tree- 
frogs  are  so  closely  allied  that  until  recently  they 
were  regarded  as  mere  varieties  of  one  another. 
These  are  Hyla  arborea  and  //.  chinensis.  The 
former  is  found  in  Asia  Minor,  Persia,  China  and 
Japan,  and  in  most  of  the  Mediterranean  islands  and 
Southern  Europe  generally.  It  does  not  occur  in  the 
British  Islands,  Norway,  or  North  Russia,  but  in 


THE  ORIENTAL  MIGRATION. 

South  Sweden,  Germany,  France,  and  Spain.  It  is 
also  known  from  North  Africa  and  from  Madeira,  the 
Canaries,  and  the  Salvages.  The  occurrence  of  the 
Tree-Frog  an  so  many  of  the  Mediterranean  islands 
is  of  particular  interest,  especially  as  four  well-marked 
varieties  have  been  distinguished  by  our  leading  her- 
petologists,  so  that  the  more  minute  features  of  the 
various  forms  can  be  traced  from  island  to  island, 
adding  one  more  proof — if  proof  were  needed — of 
their  former  continuity.  Of  course,  that  Hyla  arborea 
must  be  considered  an  Oriental  migrant  seems  so 
evident  that  it  scarcely  needs  further  comment. 

A  number  of  mollusca  might  be  mentioned  whose 
range  indicates  that  they  have  migrated  to  Europe 
from  Asia  Minor.  Buliminus  pupa  is  one  of  these. 
It  is  known  from  Asia  Minor,  Greece,  South  Italy, 
Sicily,  and  Algeria.  Buliminus  detritus  is  perhaps 
better  known,  being  common  in  some  parts  of 
Germany.  From  there  its  range  spreads  east  as 
far  as  Asia  Minor.  Many  closely  allied  species 
inhabit  Western  Asia,  to  which  they  are  confined, 
while  others  enter  on  European  territory  in  some 
of  the  Greek  islands.  B.  fasciolatus  occurs  on  the 
islands  of  Crete,  Rhodes,  Cyprus,  and  in  Greece 
and  Syria.  Most  of  the  species  of  Buliminus  have 
a  very  restricted  range,  but  Buliminus  obscurus  is 
found  almost  all  over  Europe,  from  Ireland  in  the 
west  to  the  Crimea  and  Transcaucasia  in  the  east. 

Whether  the  sub-genus  Pomatia  of  the  genus 
Helix — to  which  the  so-called  Roman  Snail  belongs — 


262         HISTORY  OF  THE   EUROPEAN   FAUNA. 

is  of  Asiatic  origin,  or  whether  some  of  the  species 
have  migrated  from  Europe  to  Asia,  I  am  not  pre- 
pared to  say;  but  there  can  be  no  doubt  that  Helix 
pomatia  has  reached  Western  Europe  from  the  east. 

On  the  whole,  the  number  of  mollusca  which  we 
might  point  to  as  having  migrated  to  Europe  is  not 
large,  the  great  majority  being  indigenous  to  our 
continent.  However,  some  of  the  other  groups  of 
invertebrates  differ  very  materially  in  that  respect 
from  the  mollusca.  I  cannot  leave  the  consideration 
of  the  mollusca  without  referring  to  the  fact  that 
there  appears  to  be  a  very  important  centre  of 
distribution  in  South-eastern  Europe.  It  is  from 
this  centre  that  many  species  have  spread  north 
and  south,  east  and  west  Take,  for  example,  the 
genus  Clausilia,  a  small  land-shell  shaped  like  a 
pointed  round  tower,  and  abundant  on  old  walls 
and  tree  trunks.  In  England  we  have  four  species 
of  Clausilia>  in  Ireland  only  two.  In  the  greater 
part  of  Spain  only  our  common  Cl.  bidentata  occurs. 
As  we  go  east  the  number  of  species  rapidly 
increases.  A  maximum  is  reached  in  South-eastern 
Europe,  where  hundreds  of  different  kinds  are  found. 
Towards  Northern  Europe  a  similar  decrease  of 
species  takes  place.  So  far  the  history  of  the 
Clausilice  seems  perfectly  simple.  An  active  centre 
of  origin  appears  to  exist  in  South-eastern  Europe, 
from  which  the  species  radiate  out  in  all  directions. 
But  when  we  come  to  look  more  closely  into  the 
extra-European  distribution  of  the  genus,  and 


THE   ORIENTAL   MIGRATION.  263 

especially  when  we  examine  its  past  history,  we 
find  that  its  origin  is  extremely  complex,  and  dates 
back  to  a  much  more  remote  period  than  would 
have  been  imagined,  had  we  merely  taken  into 
account  its  present  range  in  our  own  continent. 
Professor  Boettger,  who  is  the  highest  authority  on 
Clausilia,  tells  us  that  the  genus  is  known  from 
the  earliest  deposits  of  the  Tertiary  Era.  About 
700  species  are  now  known,  and  these  have 
been  sub-divided  by  Professor  Boettger  and  others 
into  a  number  of  sub-genera.  Some  of  these  are 
extinct,  but  the  great  majority  are  still  living. 
The  sub-genus  Phcedusa  occurs  in  the  eocene 
and  oligocene  of  Southern  Europe,  but  it  is  extinct 
as  far  as  our  continent  is  concerned.  Close  upon 
a  hundred  species,  however,  still  inhabit  India, 
the  Malayan  Islands,  China,  Ceylon,  and  Japan. 
Then  again,  the  sub-genus  Laminifera  occurs  in  the 
oligocene  and  miocene  of  Central  Europe,  and 
survives  in  a  single  species,  CL  Pauli>  in  South- 
western France.  The  groups  Garnieria  of  China, 
Macroptyckia  of  East  Africa,  Boettgeria  of  Madeira, 
and  Nenia  of  South  America,  have  no  fossil  repre- 
sentatives. We  have  here  some  very  remarkable 
cases  of  discontinuous  distribution  which  testify  to 
the  antiquity  of  the  genus,  and  this  is  certainly 
confirmed  by  the  fossil  evidence.  However,  it  is 
hardly  likely  that  the  headquarters,  as  it  were,  of 
Clausilia  have  always  been  in  South-eastern  Europe. 
Most  of  that  part  of  the  Continent  has  been  sub- 


264         HISTORY  OF  THE   EUROPEAN    FAUNA. 

merged  since  eocene  times  more  than  once.  The 
peculiar  distribution  of  the  genus  might  be  explained, 
I  think,  if  we  supposed  the  original  home  of  Clausilia 
to  have  been  in  Southern  Asia,  that  from  this  centre 
Southern  Europe  was  colonised,  where  a  new  centre 
developed  in  oligocene  and  miocene  times,  sending 
colonies  off  to  Madeira  and  across  the  old  land- 
connection  which  united  Northern  Africa  and  South 
America  about  that  time.  The  most  active  centre 
of  development  then  gradually  shifted  eastward 
again,  while  the  older  centres  were  perhaps  sub- 
merged during  the  physical  changes  in  the  distri- 
bution of  land  and  water. 

I  should  have  mentioned  that  the  species 
wandering  westward  and  northward  from  this 
South-European  centre  of  distribution,  would 
naturally  have  joined  the  migrants  which  came 
from  beyond  the  borders  of  our  continent.  They 
might  thus  appear  to  be  true  Oriental  migrants, 
and  on  a  previous  occasion  I  grouped  all  these 
together  under  the  term  of  "Southern  Fauna,"  as 
I  assumed  the  observer  to  be  stationed  in  the 
British  Islands.  All  new-comers  from  the  south- 
east, south,  or  south-west  of  Europe  would  be  to 
him  southerners  quite  irrespective  of  their  original 
home,  which  might  be  in  Southern  Europe,  Asia, 
or  Africa. 

The  Swallow-tail  is  well  known  to  all  collectors 
of  Butterflies  in  England,  though  it  has  of  late 
years  become  very  rare  and  is  now  confined  to  a 


THE  ORIENTAL  MIGRATION.  265 

few  localities  in  the  east  of  England.  The  members 
of  the  family  Papilionidce,  to  which  it  belongs,  are 
mostly  large  and  striking  species,  and  their  distri- 
bution is  therefore  more  accurately  known  than 
that  of  the  smaller  and  less  conspicuous  butterflies. 
Only  four  different  kinds  of  Swallow-tail  Butterflies 
inhabit  Europe,  but  in  Southern  Asia  and  the  Malay 
peninsula  they  attain  their  maximum  as  regards 
numbers;  and  there  we  find  a  great  many  species  of 
this  genus  Papilio.  Of  the  four  European  species  only 
one,  viz.,  Papilio  hospiton,  is  peculiar  to  Europe ;  all 
the  others  range  into  Asia.  It  would  seem,  therefore, 
as  if  this  genus  was  an  Asiatic  one  and  had  migrated 
to  Europe,  and  that  the  route  taken  was  the  one  from 
Asia  Minor  across  to  Greece.  We  have  a  similar 
case  in  the  closely  allied  genus  Tliais  two  of  the 
three  European  species  living  also  in  Asia  Minor. 
Thais  cerisyi  inhabits  some  of  the  Greek  islands,  as 
well  as  the  mainland  of  Turkey  and  Greece. 

Another  genus  of  the  great  family  Papilionidce 
with  which  most  lepidopterists  are  well  acquainted 
is  Parnassius.  What  butterfly- hunter  has  been  in 
Switzerland  without  hearing  of,  or  seeing,  the  famous 
Parnassius  Apollo  ?  We  have  four  European  species  of 
Parnassius t  only  one  of  which  is  peculiar  to  our  con- 
tinent, but  the  locality  where  it  occurs,  the  Caucasus, 
is  on  the  borders  of  Asia.  Almost  all  the  other 
species  are  Asiatic,  none  however  range  to  the  south. 
Its  headquarters,  and  I  think  its  original  home,  are 
the  mountains  of  Central  Asia.  From  there  it 


266          HISTORY  OF   THE    EUROPEAN   FAUNA. 

has  spread — some  species  to  the  Himalayas,  and 
a  few  to  Europe  and  North  America.  But  these 
migrations  are  not  of  very  recent  date.  Parnassius 
no  doubt  arrived  accompanied  by  a  large  number 
of  other  Central  Asiatic  mountain  insects  and 
plants.  I  shall  refer  to  the  latter  again  when 
dealing  with  the  origin  of  the  Alpine  fauna,  but 
meanwhile  it  might  be  mentioned  that  the  famous 
Swiss  "Edelweiss"  (Leontopodium  alpinuni),  which 
we  are  accustomed  to  regard  as  a  typical  Alpine 
plant,  is  certainly  of  Asiatic  origin.  In  some  parts 
of  Southern  Siberia  it  is  one  of  the  common  meadow- 
flowers,  and  ranges  from  there  south  into  Kashmere, 
but  not  northward.  Like  the  Apollo,  it  does  not 
occur  in  Scandinavia  or  Northern  Siberia.  Both 
plant  and  insect  evidently  migrated  from  Central 
Asia,  directly  westward  along  the  southern  border  of 
the  sea,  which  extended  from  that  region  as  far  as 
the  European  Alps  in  early  Tertiary  times.  At  that 
time  the  Caucasus  was  possibly  still  connected  with 
the  Balkan  Mountains,  across  what  is  now  the  Black 
Sea,  and  that  may  have  been  the  highway  on  which 
they  travelled  west. 

Some  of  the  Clouded-Yellows — butterflies  apper- 
taining to  the  genus  Colias — formed  part  of  the 
Oriental  migration.  The  genus  is  undoubtedly  of 
Asiatic  origin,  and  while  many  of  the  species  have 
turned  northward,  ranging  across  Siberia  and  North 
America,  others  have  taken  a  southern  and  westward 
turn  and  thus  reached  Europe.  We  have  two 


THE  ORIENTAL  MIGRATION.  267 

Clouded-Yellows   in    Western    Europe,  and  both  of 
them  must  have  come  with  this  migration. 

A  very  good  example  of  an  Oriental  migrant  is 
Danais  clirysippus,  a  magnificent  butterfly  found  in 
Greece  and  Southern  Italy.  In  Asia  it  is  known  from 
Syria,  Persia,  and  from  the  whole  of  the  southern 
portion  of  the  Continent.  The  genus  Danais  (in 
its  wide  sense)  is  a  large  one,  and  principally 
occurs  in  the  warmer  regions  of  Asia.  Three 
species  are  found  in  North  America  and  only  one  in 
Europe. 

Among  the  beetles  belonging  to  this  migration, 
there  is  one  of  very  considerable  interest  from  a  dis- 
tributional point  of  view,  for  all  the  species  of  the 
genus — even  the  whole  family  to  which  the  genus 
belongs — are  what  is  known  by  zoologists  as  "  Com- 
mensalists."  These  are  animals  habitually  associating 
and  living  in  close  connection  with  others  with  which 
they  are  not  tied  by  any  family  relations  or  kinship. 
Such  a  state  of  close  and  permanent  friendship  is 
called  "  commensalism."  Now  it  appears  as  if  the 
members  of  this  family  of  beetles  (Clavigeridce)  had  of 
their  own  free  will  formed  such  a  close  connection 
with  colonies  of  ants — sometimes  with  one  species, 
sometimes  another.  They  are  the  permanent  guests 
of  the  ants,  and  in  return  they  secrete  a  fluid  which  is 
apparently  highly  prized  by  them.  All  of  the  Clavi- 
gers  are  provided  with  peculiar  club-shaped  antennae, 
with  which  they  ungraciously  beat  their  hosts,  when 
they  are  in  want  of  food.  According  to  some  authori- 


268         HISTORY  OF  THE   EUROPEAN   FAUNA. 

ties,  they  even  occasionally  gnaw  at  the  pupae  and 
larvae  of  the  ant  with  which  they  live. 

Such  beetles  naturally  can  only  have  extremely 
limited  means  of  distribution,  and  they  are  com- 
parable in  that  respect  with  the  woodlice  of  the  genus 
Platyarthrus,  to  which  I  have  already  had  occasion 
to  refer.  All  the  species  of  Claviger  are  confined  to 
Europe,  chiefly  to  the  south,  but  one  species,  CL 
testaceus,  has  wandered  farther  north  and  occurs  in 
the  nest  of  the  ant  Lasius  flavus  in  the  south  of 
England,  Ireland,  and  Scotland.  Though  none  of  the 
Clavigers  can  be  claimed  as  Oriental  migrants,  the 
centre  of  distribution  of  the  genera  belonging  to  the 
Clavigerida  is  in  Southern  Asia,  and  it  is  probable 
that  the  ancestors  of  the  European  Clavigers  have 
spread  westward  from  that  region  to  Europe,  eastward 
to  Australia  and  Japan,  and  southward  to  Madagascar 
and  South  Africa.  The  genus  Hopatroides,  belonging 
to  the  same  family  as  the  so-called  Spanish-fly  (Tene- 
brionidcz})  has  twelve  species  in  Western  Asia  and 
Greece.  One  only,  H.  thoracicus — an  instance  of  dis- 
continuous distribution — occurs  in  Andalusia.  Amphi- 
coma  is  represented  in  Western  Asia  and  the  Balkan 
peninsula  by  fifteen  species,  while  three  others  are 
met  with  in  North-west  Africa  and  Southern  Spain. 

A  genus  of  Dragon  -  fly,  Onychogomphusy  has  in 
Europe  a  somewhat  similar  distribution  to  Claviger, 
but  it  has  besides  a  very  extensive  foreign  range. 
There  are  altogether  thirty-five  species ;  of  these  ten 
are  Holarctic,  twelve  Oriental,  five  Mascarene,  and 


THE   ORIENTAL   MIGRATION.  269 

eight  Ethiopian.  The  centre  of  distribution  is  there- 
fore in  the  Oriental  region,  and  we  may  assume  that 
in  all  probability  the  genus  has  originated  there, 
the  European  species  having  travelled  west  with 
the  Oriental  migration  at  an  early  date  of  the 
Tertiary  Era. 

Ryothemis,  another  genus  of  Dragon  -  flies,  has 
originated  perhaps  somewhat  farther  east  than  the 
last,,  for  no  less  than  thirteen  species  are  found 
in  Australia,  a  like  number  in  India,  five  in  Mada- 
gascar and  Africa,  and  five  in  the  Holarctic  region. 
Both  of  these  genera  are  entirely  absent  from 
America,  and  they  have  possibly  travelled  to  Europe 
together. 

Among  the  European  OrtJwptera — the  group  to 
which  our  Earwigs  and  Grasshoppers  belong — there 
are  also  a  good  many  instances  of  Oriental  migrants. 
One  of  the  most  striking  of  these  is  the  curious 
"praying  insect"  (Mantis  religiosd].  It  occurs  all 
over  Southern  Europe,  and  ranges  as  far  north  as  the 
north  of  France.  It  is  also  found  in  Southern  Ger- 
many and  in  Austria,  and  has  a  vast  extra-European 
range.  There  are  even  records  of  its  occurrence  from 
all  parts  of  Southern  Asia  and  Java  and  a  great  part 
of  Africa.  That  it  belongs  to  an  extreme'y  ancient 
genus  is  testified  by  the  fact  of  its  presence  in 
Mauritius,  Japan,  Australia,  New  Zealand,  South 
America,  and  Madagascar.  The  genus  Bacilhis — to 
which  the  typical  Stick-insects  belong — has  a  some- 
what similar  geographical  distribution.  But  no  less 


2/0         HISTORY   OF   THE   EUROPEAN   FAUNA. 

than  four  species  of  Bacillus  are  known  from  Europe, 
according  to  our  great  authority  Mr.  Brunner  von 
Wattenwyl — all  from  the  south ;  and  some  of  these 
also  range  into  North  Africa.  There  are  thirty-two 
other  species  distributed  over  Southern  Asia,  Africa, 
Australia,  New  Zealand,  and  the  Sandwich  Islands. 

Volumes,  indeed,  might  be  filled  with  lists  of 
species  and  genera  of  terrestrial  invertebrates  of 
Oriental  origin,  but  I  will  not  weary  the  reader 
with  further  enumeration  of  such  instances.  Just 
two  more,  however,  before  concluding,  as  I  have 
not  alluded  to  the  large  group  of  the  Arachnida. 

Two  peculiar  spider-like  genera,  viz.,  Galeodes  and 
Rhax>  are  found  in  Southern  Europe.  Both  occur 
also  in  North  Africa,  and  in  Western  and  a  portion  of 
Southern  Asia.  As  the  whole  family  altogether  has 
an  Asiatic  character,  I  cannot  agree  with  Mr.  Pocock, 
who  considers  them  of  European  origin  and  believes 
that  they  are  migrating  eastward. 

But  not  only  terrestrial  forms  migrated  to  Europe 
from  Western  and  Southern  Asia.  Freshwater 
species  also  took  part  in  this  great  Oriental  migra- 
tion. 1  need  only  refer  to  the  freshwater  Crab 
(Thelphusa  fluviatilis),  with  which  Southern  Euro- 
peans are  familiar.  It  is  the  sole  representative  of  a 
large  genus  which  ranges  east  as  far  as  Australia  and 
southward  to  Madagascar  and  the  Cape  of  Good 
Hope.  The  European  species  is  found  in  Turkey, 
Cyprus,  Greece,  Southern  Italy,  Sicily,  North  Africa, 
Southern  Spain,  Syria,  and  Persia. 


THE   ORIENTAL  MIGRATION.  271 

There  is  a  corresponding  flora  with  a  range  exactly 
similar  to  that  of  some  of  the  animals  quoted.  Thus 
the  Balkan  Rhododendron  (Rhododendron  ponticmri) 
is  again  met  with  in  the  western  Mediterranean 
region  in  Southern  Spain.  The  Cedar  occurs  in 
local  varieties  in  the  Himalayan  Mountains,  in  the 
Lebanon,  and  the  Atlas  Mountains.  Both  of  these 
are  instances  of  discontinuous  distribution,  a  proof  of 
their  antiquity;  but  a  large  number  of  plants  have  a 
continuous  range  between  Asia  Minor  and  Spain. 

On  looking  through  these  few  instances  of  what  have 
been  called  Oriental  migrants,  one  cannot  help  being 
struck  by  the  fact  that  the  species  after  their  entry 
into  Europe  evidently  did  not  all  follow  the  same  path 
during  their  westward  advance.  We  have  seen  that 
a  good  many  seem  to  have  travelled  either  due  west 
or  north-west  on  entering  our  continent  from  Asia 
Minor.  They  may  now  perhaps  be  found  in  Greece, 
Southern  Italy,  Algiers,  and  Spain,  also  probably  on 
some  of  the  intervening  islands  in  the  Greek  Archipe- 
lago, in  Sicily,  Sardinia,  and  Corsica,  or  they  may  have 
travelled  north-east  and  occur  in  the  Alps.  This  distri- 
bution indicates  undoubtedly,  as  I  have  already  set  forth 
in  another  memoir  (c,  p.  459),  that  land  extended  from 
Asia  Minor  across  Greece  to  Southern  Italy,  that  the 
latter  again  was  disconnected  with  Central  Italy,  but 
united  with  Sicily,  Sardinia,  and  Tunis,  and  that  the 
Straits  of  Gibraltar  did  not  exist  at  the  time  when 
these  species  migrated  westward.  Some  species 
are  only  to  be  found  as  far  west  as  Southern  Italy, 


2/2         HISTORY   OF  THE   EUROPEAN    FAUNA. 

while  others  occur  in  Central  and  Northern  Europe, 
scarcely  in  the  South,  and  not  at  all  in  the 
larger  Mediterranean  islands  or  in  North  Africa. 
This  appears  to  me  to  indicate  that  the  late  comers 
from  the  east  found  that  geographical  changes  had 
taken  place  in  Southern  Europe  which  prevented 
them  from  following  the  same  track  as  the  older 
immigrants.  They  were  now  obliged  to  turn  directly 
northward  and  then  westward.  It  may  be  asked, 
why  should  not  the  earlier  migrants  have  taken  the 
same  route?  This  question  will  be  answered  imme- 
diately. Meanwhile  it  should  be  clearly  understood 
that  there  probably  was  an  older  and  a  newer  migra- 
tion from  the  east.  The  Oriental  genera — from 
whose  general  range  we  know  that  they  must  be 
very  ancient  indeed,  such  as  Mantis  and  Bacillus — 
are  almost  invariably  confined  to  Southern  Europe. 
There  they  are  frequently  found  on  some  of  the 
Mediterranean  islands.  The  earlier  migrants  there- 
fore went  westward  and  the  later  ones  north- 
ward. 

Let  us  now  inquire  a  little  into  the  reasons  why 
such  different  courses  were  pursued  by  the  migrants 
— why  the  Oriental  migration  divided  into  two 
streams,  an  older  and  a  newer. 

During  early  Tertiary  times,  and  probably  through- 
out the  Miocene  and  Pliocene  Epochs,  the  ^Egean 
Sea  did  not  exist.  From  the  island  of  Crete  to  the 
Peloponnesus,  and  from  Asia  Minor  to  Thessaly 
and  Macedonia,  stretched  a  vast  and  fertile  plain 


THE   ORIENTAL   MIGRATION.  273 

dotted  over  with  numerous  freshwater  lakes.  Grad- 
ually the  sea  encroached  upon  this  land  from  the 
south,  owing  chiefly  to  extensive  subsidences  having 
taken  place.  Only  very  recently,  says  Professor 
Suess,  did  the  whole  of  the  ^Egean  continent  subside 
(i.,  p.  437).  Huge  cliffs  of  levantine  freshwater 
deposits  now  mark  the  new  coast-line,  and  the  Medi-. 
terranean  advances  steadily  towards  the  Black  Sea 
and  the  Sea  of  Asov.  A  new  order  of  things  is  now 
established,  continues  the  famous  author  of  Das 
Antlitz  der  Erde;  where  there  were  high  mountains 
we  now  behold  a  deep  sea,  in  some  places  many 
thousand  feet  deep.  All  this  took  place  quite 
recently, — geologically  speaking, — certainly  in  post- 
glacial times;  and  man  may  even  have  witnessed 
these  imposing  events.  Most  geologists  admit  the 
correctness  of  these  views.  They  are,  moreover, 
built  upon  such  solid  geological  evidence,  that  even 
if  the  science  of  zoogeography  had  not  yet  taught  us 
anything,  naturalists  would  not  hesitate  in  accepting 
them. 

Animals  and  plants  were  free  to  migrate  from 
Central  and  Southern  Asia  to  Greece  by  land  for 
untold  ages.  The  vast  accumulation  of  mammalian 
bones  which  have  been  discovered  at  Pikermi,  and  so 
ably  described  by  Gaudry,  are  probably  to  a  large 
extent  the  remains  of  Asiatic  immigrants  to  Europe. 
Many  of  these  resemble  forms  still  living  in  South 
Africa,  which  implies  that  a  highway  existed  also  at 
that  time  between  Asia  and  Africa.  Among  these  is 

18 


2/4         HISTORY   OF   THE    EUROPEAN   FAUNA. 

a   giraffe   and    antelopes    closely   allied    to   African 
species,  and  other  most  interesting  mammals. 

In  still  earlier  European  deposits — the  Miocene — 
we  find  the  ancestors  of  modem  Elephants,  which 
are  probably  of  Asiatic  origin.  The  remains  of 
several  kinds  of  monkeys  occur,  whose  nearest 
relations  are  now  confined  to  Southern  Asia. 
Altogether  the  fauna  bears  a  strong  Asiatic  facies. 
Many  of  our  European  terrestrial  invertebrates 
probably  arrived  about  this  time  from  Asia.  The 
struggle  for  existence  being  keener  and  the  facility 
for  migration  much  greater  in  the  higher  vertebrates, 
they — or  at  any  rate  the  mammalian  faunas — were 
subjected  to  more  rapid  changes  than  the  inverte- 
brates. I  have  repeatedly  expressed  my  belief  that 
a  great  number  of  our  familiar  insects  and  mollusca 
inhabited  Europe  long  before  our  present  mammals 
came  into  existence.1 

Let  us  now  follow  one  of  the  miocene  Oriental 
migrants  starting  from  Central  Asia  on  its  way  to 
Europe.  Very  soon  after  leaving  its  home,  it  must 
have  encountered  a  sea  which  extended  at  that  time 
from  the  Eastern  Mediterranean  to  the  borders  of 
Afghanistan.  In  following  a  westward  course,  the 
emigrant  was  compelled  to  keep  along  the  northern 
shore  of  it.  We  do  not  know  the  state  of  the 
physical  geography  of  the  region  between  the  Black 

1  In  some  cases  the  accuracy  of  this  view  is  proved  by  fossil  evidence, 
Helix  rottindata,  a  common  and  widely  spread  British  species,  having 
been  found  in  miocene  strata  near  Bordeaux. 


THE   ORIENTAL   MIGRATION.  275 

Sea  and  the  Tianshan  Mountains,  but  it  seems 
certain  that  a  considerable  extent  of  dry  land 
enabled  a  wanderer  from  Central  or  Southern  Asia 
to  reach  the  Balkan  peninsula  by  skirting  the 
northern  shore  of  that  large  miocene  sea.  No 
miocene  deposits  occur  north  of  Teheran  or  of  the 
Upper  Euphrates,  nor  are  they  known  from  the 
islands  of  the  ^Egean  Sea  or  the  lands  surrounding  it. 
From  the  Balkan  peninsula  it  was  possible  for  our 
migrant  to  reach  the  European  Alps,  which  were 
then  slowly  rising  as  a  peninsula  out  of  the  western 
portion  of  the  great  miocene  sea.  What  are  now 
the  Alps  was  then  hilly  ground,  which  was  being 
raised  from  the  bottom  of  the  sea.  It  was  no  doubt 
connected  with  the  Balkan  peninsula,  so  that  an 
intercourse  of  species  could  take  place  between  this 
newly-formed  peninsula  and  Central  Asia.  I  say 
peninsula,  because  the  miocene  sea  almost  completely 
surrounded  it.  From  the  Western  Mediterranean  a 
wide  gulf  extended  up  the  Rhone  valley  into  that  of 
the  Rhine  as  far  north  as  Maintz.  Then  skirting  along 
the  northern  outliers  of  the  Tyrol,  the  gulf  can  be 
followed  as  far  east  as  Transylvania.  It  is  quite 
probable  that  it  extended  much  farther  east  still,  but 
there  is  as  yet  no  geological  evidence  forthcoming. 
At  any  rate,  our  Asiatic  migrant  turning  northward 
from  the  Balkan  peninsula  found  its  farther  progress 
barred  once  more  by  an  arm  of  the  same  sea  which  in 
its  earlier  peregrinations  had  stopped  it  from  going 
south  (cf.  Suess,  i.,  p.  406). 


2/6         HISTORY   OF   THE   EUROPEAN    FAUNA. 

In  later  miocene  times  the  sea  does  not  seem  to 
have  surrounded  the  Alps  to  the  same  extent  as  it 
did  before,  but  it  certainly  extended  from  the  Eastern 
Alps  to  the  shores  of  the  Sea  of  Asov,  so  that  the 
direct  northward  passage  was  still  more  or  less  barred 
to  the  Oriental  immigrants.  At  the  same  time  Alpine 
species  were  now  able  to  emigrate  to  the  North 
European  provinces.  During  the  last  stages  of  this 
epoch,  the  same  sea  increased  its  area  very  consider- 
ably in  an  eastward  direction.  One  continuous 
expanse  of  water  now  stretched  from  the  Alps  as 
far  as  the  Sea  of  Aral  in  Central  Asia,  perhaps  even 
farther. 

During  pliocene  times  especially,  the  northern  parts 
of  the  Balkan  peninsula  were  occupied  by  a  series  of 
freshwater  lakes,  while  Greece  was  joined  to  Southern 
Italy,  Sicily,  and  Tunis.  Central  and  Northern  Italy 
were  represented  by  a  long  narrow  peninsula  con- 
nected in  the  north  with  the  Alps.  Corsica  and 
Sardinia  were  joined  to  Sicily,  and  the  Straits  of 
Gibraltar  did  not  exist.  When  I  first  published  my 
views  regarding  these  geographical  conditions  of  the 
Mediterranean  area,  Professor  Deperet  was  good 
enough  to  send  me  his  criticisms  from  a  purely 
geological  standpoint.  He  is  of  opinion  that  though 
Sicily  and  Sardinia  might  at  this  time  have  still  been 
connected  with  Tunis,  the  Straits  of  Messina  must 
already  have  been  formed — in  other  words,  Southern 
Italy  and  Sicily  could  no  longer  have  been  connected 
with  one  another.  This  opinion  is  based  upon  the 


THE  ORIENTAL   MIGRATION.  277 

fact  that  in  the  upper  strata  of  the  enormously  thick 
Sicilian  pliocene  deposits  are  found  a  number  of 
arctic  or  subarctic  species  of  mollusca  which  are 
entirely  foreign  to  the  Mediterranean  fauna.  It  is 
generally  supposed  that  these  reached  the  Mediter- 
ranean area  by  the  newly  opened  Straits  of  Gibraltar 
in  later  pliocene  times,  and  that  the  lower  Sicilian 
deposits  must  therefore  have  been  laid  down  earlier. 
So  far  the  deductions  are  perfectly  correct,  if  we  assume 
the  northern  mollusca  to  have  arrived  in  the  Atlantic 
at  the  time  stated.  However,  they  must  have  reached 
the  Atlantic  much  later — not  till  pleistocene  times — 
if  we  adopt  the  above-stated  suggestions  as  to  the 
age  of  the  Forest-Bed  (cf.  p.  125).  Moreover,  the 
great  similarity  between  the  faunas  of  Southern 
Spain  and  North-western  Africa  indicate  that  the 
formation  of  the  Straits  of  Gibraltar  is  of  very 
recent  date.  The  northern  mollusca,  of  course, 
could  not  have  reached  Sicily  till  later.  To  suppose 
that  the  Sicilian  deposits  have  been  uplifted  7000  feet 
since  then  is  no  doubt  contrary  to  all  our  geological 
teaching,  but  we  must  remember  that  this  is  altogether 
an  exceptional  case.  The  area  in  question  has  prob- 
ably ever  since  been  in  the  immediate  neighbourhood 
of  an  active  volcano,  and  the  rate  of  the  uplift  has 
therefore  been  immeasurably  greater  than  at  other 
localities  with  which  this  one  might  be  compared.  The 
disconnection  between  Tunis,  Sicily,  and  Southern 
Italy  was  evidently  produced  by  a  subsidence  of  the 
tract  of  land  uniting  these  countries.  If  we  suppose 


2/8         HISTORY   OF   THE   EUROPEAN   FAUNA. 

that  this  happened  in  early  pliocene  times,  we  have 
either  to  take  for  granted  that  the  terrestrial  fauna 
and  flora  of  these  countries  are  of  miocene  origin,  or 
that  they  were  joined  again  during  the  Pleistocene 
Epoch.  The  range  of  a  very  large  number  of  animals 
and  plants  is  such  as  can  only  be  explained  by 
assuming  that  Tunis,  Sicily,  Sardinia,  Corsica,  and 
Southern  Italy  were  connected  with  one  another.  Of 
such  extensive  land-connections  subsequent  to  the 
arrival  of  the  northern  marine  mollusca  we  possess, 
however,  no  geological  evidence  whatsoever;  and  it  is 
extremely  improbable  that  the  land-areas  which  had 
sunk  were  once  more  raised  before  again  subsiding. 
The  many  animals  whose  presence  in  the  Mediter- 
ranean Region  bears  witness  to  these  ancient  land- 
connections  could  not  have  arrived  there  in  miocene 
times — in  fact,  they  could  hardly  have  lived  there 
before  the  end  of  the  Pliocene  Epoch.  On  the  other 
hand,  it  seems  difficult  to  believe,  once  the  Straits  of 
Gibraltar  were  open  and  the  waters  of  the  Atlantic 
able  to  enter  the  Mediterranean,  that  the  sunken 
parts  between  Sicily,  Italy,  and  Tunis  could  have 
been  raised  without  affecting  the  entire  area  of  that 
sea.  Nor  is  it  likely  that  the  junction  between  these 
countries  could  have  then  been  brought  about  by  a 
general  lowering  of  the  Mediterranean  waters.  As  it 
may  be  asked  what  evidences  we  possess  at  all  for 
the  supposition  of  such  land-connections  as  I  have 
indicated,  also  that  Southern  Italy  and  Greece 
were  connected,  a  few  of  the  more  salient  instances 


THE   ORIENTAL   MIGRATION.  279 

of  distribution  bearing  on  this  problem  may  be  of 
interest. 

I  have  already  referred  to  the  occurrence  of  the 
remains  of  a  small  race  of  Red  Deer  in  the  caves 
of  Malta,  similar  to  those  still  living  in  North- 
west Africa,  Corsica,  and  Sardinia.  The  Black- 
mouthed  Weasel  (Mustela  boccamela)  inhabits  Persia, 
Asia  Minor,  Greece,  South  Italy,  Sicily,  and  Sardinia, 
while  Mustela  africana  is  found  in  Malta  and  Algiers. 
The  European  Porcupine  inhabits  Asia  Minor,  the 
island  of  Rhodos,  Greece,  Southern  Italy,  Sicily, 
North  Africa,  and  Spain.  Then  we  have  the  Wild 
Sheep  of  Asia  Minor,  Cyprus,  Sardinia,  and  Corsica, 
all  of  which  are  closely  allied.  The  small  shrew-like 
Crocidura  etrusca  occurs  in  South  France,  Italy, 
Sicily,  and  North  Africa.  Many  other  mammalia 
might  be  quoted,  but  these  are  sufficient  for  our 
purpose. 

There  are  a  good  many  reptiles  and  amphibians 
with  a  similar  distribution.  The  European  Chamaeleon 
(Chamczleon  vulgaris)  has  been  found  in  South  Spain, 
North  Africa,  and  Sicily.  The  Snake  Periops  hippo- 
crepis  is  confined  to  Spain,  Sardinia,  and  Greece. 
The  worm-like  Lizard  Blanus  cinereus  inhabits  some 
of  the  Greek  islands,  North  Africa,  and  Spain. 
Another  Lizard  belonging  to  the  Scincid&  has  also 
been  found  in  some  of  the  Greek  islands,  Sicily, 
Sardinia,  Southern  Spain,  and  the  Canary  Islands. 
Discoglosstis  pictus — a  toad" — occurs  in  Spain,  North- 
west Africa,  Malta,  Sicily,  Sardinia,  and  Corsica. 


280         HISTORY  OF   THE    EUROPEAN    FAUNA. 

A  variety  of  the  Tree  Frog  (Hyla  arborea  Savignyi} 
is  found  in  Europe  only  in  Corsica,  Sardinia,  and  the 
Greek  Archipelago. 

Eight  species  of  Reptiles  and  Amphibia — some  of 
which  I  have  just  referred  to — are  enumerated  by  Dr. 
Forsyth  Major  as  occurring  eastward  and  westward 
of  the  Italian  peninsula  (and  almost  all  also  in  North 
Africa)  without  being  known  on  the  mainland  of  Italy. 
And  in  order  to  show  that  Sardinia  and  Corsica  are 
more  closely  related  to  North  Africa  than  to  Italy, 
he  indicates  the  general  range  of  the  Reptiles  and 
Amphibians  found  in  these  islands.  Of  the  twenty- 
one  species,  only  twelve  inhabit  Italy,  but  at  least 
sixteen  North  Africa  and  seventeen  Spain.  Indeed, 
he  shows  that  Corsica,  Sardinia,  Sicily,  and  North- 
west Africa  form  a  zoogeographical  province,  from 
which  Italy,  with  the  exception  of  a  few  localities 
on  its  west  coast,  is  excluded.  It  is  a  remarkable 
fact  that  there  are  a  few  localities  on  the  west 
coast  of  Italy  which  in  their  fauna  and  flora 
exhibit  closer  relationship  with  Corsica  and  Sar- 
dinia than  with  the  mainland.  Thus  Dr.  Major 
pointed  out  that  the  Catena  Mettalifera,  the  Monte 
Argentario,  and  Monte  Circeo  all  belong  to  what 
we  may  call  the  former  Tyrrhenian  continent. 
They  are  to  be  regarded  as  its  eastern  limits,  which 
remained  standing,  while  the  central  portion — now 
occupied  by  the  Tyrrhenian  Sea — subsided,  and  is 
at  present  covered  by  deep  sea.  Subsequently  these 
remnants-  of  the  old  continent  became  joined  with 


THE   ORIENTAL   MIGRATION.  28 1 

the  newly-formed  Italian  peninsula,  but  the  plants 
and  animals  belonging  to  the  older  flora  and  fauna 
were  mostly  destroyed  by  newer  and  more  vigorous 
immigrants.  A  few  of  the  more  hardy  ones  survived, 
and  are  a  standing  testimony  of  the  geographical 
revolutions  of  that  part  of  Southern  Europe. 

That  the  Mediterranean  area  has  undergone  such 
profound  geographical  changes  as  I  have  endeavoured 
to  indicate  is  no  new  theory.  Many  zoologists  who 
have  investigated  the  fauna  of  that  region,  and  have 
attempted  to  explain  the  faunistic  relations,  had  to 
acknowledge  that  the  migrations  must  have  taken 
place  under  geographical  conditions  entirely  different 
from  those  obtaining  at  present.  Riitimeyer  long 
ago  remarked  that  it  seemed  to  him  much  more 
probable  that  Morocco,  Algeria,  and  Tunis  were 
peopled  by  way  of  Gibraltar,  and  perhaps  also  by 
Sicily  and  Malta  from  Europe,  than  Southern 
Europe  from  Africa.  After  careful  conchological 
researches  in  the  Western  Mediterranean  region, 
Dr.  Kobelt  came  to  the  conclusion  that  formerly 
Southern  Spain  and  Morocco  must  have  been 
united  by  a  broad  land-connection.  Sicily  and 
Algeria  do  not  apparently  show  any  very  intimate 
relationship  conchologically,  but  farther  west — in  the 
mountains  of  Tetuan — Dr.  Kobelt  discovered  a  colony 
of  Sicilian  forms.1 

1  There  are  a  great  many  instances  of  discontinuous  distribution 
among  Oriental  Invertebrates.  Thus  the  Freshwater  Crab  ( Telphitsa 
faiviatilis]  occurs  in  Southern  Italy,  Greece,  Turkey,  Cyprus,  and 


282          HISTORY  OF   THE   EUROPEAN   FAUNA. 

"  The  close  relationship,"  remarks  Dr.  Major  (a, 
p.  106),  "shown  in  the  fauna  of  Corsica  and  Sardinia 
to  Africa,  permits  the  supposition  that  the  connec- 
tion with  these  islands  had  persisted  to  a  much  more 
recent  date  than  that  with  Europe." 

Many  other  authors  have  pointed  out  the  close 
similarity  existing  between  the  faunas  of  Southern 
Europe  and  North  Africa.  We  need  only  refer  to 
the  writings  of  Professor  Suess,  Milne-Edwards,  and 
Boyd  Dawkins.  Mr.  Blanchard  went  even  so  far  as 
to  say,  "  a  comparer  les  plantes  et  les  animaux  de  la 
Sicile  et  de  la  Tunesie,  on  se  croirait  sur  le  meme 
terrain"  (p.  1047). 

No  less  than  113  species  of  phanerogamic  plants 
are  enumerated  by  Professor  Engler  (p.  53)  as  occur- 
ring in  the  Mediterranean  coast  region  east  and  west 
of  Italy  without  being  found  in  that  peninsula,  or  at 
least  only  in  the  extreme  south  of  it.  But  he  tells  us 
that  these  species  represent  only  a  portion  of  such 
plants,  which  are  extremely  numerous. 

In  taking  a  general  survey  of  these  plants,  Pro- 
fessor Engler  is  of  opinion  that  their  range  implies 
that  a  large  number  of  the  Mediterranean  species 
have  migrated  along  a  line  which  can  be  drawn 
between  North  Africa,  Sicily,  Greece,  Crete,  and  Asia 
Minor,  and  that  from  this  line  the  distribution  started 
northward  again. 

Asia  Minor.  Another  crustacean— a  Freshwater  Crayfish — (Hemicari- 
dina  Desmaresti]  inhabits  Spain,  Corsica,  Sardinia,  Sicily,  and  Asia 
Minor. 


THE  ORIENTAL  MIGRATION.  283 

Many  of  these  plants  then,  and  also  some  of  the 
animals  I  have  referred  to,  formed  part  of  the  older 
stream  of  migration  which  entered  Europe  from  Asia 
Minor  (vide  Fig.  5,  p.  117).  There  were  only  two 
courses  open  to  them  as  they  arrived  on  our  continent 
during  earlier  Tertiary  times.  They  could  either  go 
straight  west  towards  Greece,  or  in  a  more  northward 
direction  to  the  newly-formed  Alps.  As  the  latter 
were  raised,  some  of  the  immigrants  were  modified 
so  as  to  adapt  themselves  to  the  new  surroundings. 
Others  became  extinct ;  but  a  great  many  have  per- 
sisted in  the  Alps  to  the  present  day  and  exhibit 
discontinuous  distribution,  having  meanwhile  dis- 
appeared in  the  intermediate  tract  between  the  latter 
and  their  original  home  in  Asia.  The  lowlands  of 
Eastern  and  Central  Europe  were  either  occupied  by 
the  sea  or  by  large  freshwater  lakes,  so  as  effectually 
to  prevent  a  direct  migration  northward. 

When  the  newer  migrants  arrived  from  Asia  not 
only  had  the  Alps  risen  to  a  lofty,  mountain  chain 
acting  as  an  effectual  barrier,  but  Southern  Italy  and 
Greece  had  become  disconnected.  Some  time  after, 
Sicily  and  Southern  Italy  also  became  separated. 
Meanwhile  the  stream  of  migrants  which  consisted 
less  and  less  of  typically  southern  forms,  emigrants 
from  Central  Asia  and  even  Southern  Siberia, 
mingled  with  the  southern  forms  on  their  way  to 
Europe,  and  these  now  poured  across  the  newly 
opened  plain  of  Central  and  Northern  Europe.  But 
it  was  not  until  some  time  after  this  that  the  Mediter- 


284         HISTORY   OF   THE   EUROPEAN    FAUNA. 

ranean  Sea  broke  across  the  ^Egean  region,  and  that 
the  Northern  Sea  retired  from  the  plains  of  Eastern 
Russia  to  admit  the  typical  Siberian  fauna  and  flora 
into  our  continent  (vide  pp.  189-241). 

I  cannot  close  this  chapter  without  referring  to  the 
active  distributional  centre — or  I  might  say,  centre  of 
origin — of  species  situated  in  South-eastern  Europe. 
No  group  of  animals  is  more  instructive  in  eluci- 
dating the  paths  of  migration  from  this  centre  than 
the  terrestrial  mollusca.  Wherever  the  original  home 
of  the  genus  Clausilia  may  have  been  in  early  Tertiary 
times,  it  is  certain  that  the  most  active  centre  of 
origin  is  now,  and  has  been  for  a  considerable 
time  past,  in  South-eastern  Europe.  One  of  the 
earliest  migrants  from  that  modern  centre  of  this 
interesting  genus  is  Clausilia  bidentata,  which  is  the 
only  species  found  in  Southern  Spain,  and  one  of 
the  two  met  with  in  Ireland,  and  which  has  been 
observed  in  high  altitudes  in  the  Alps  and  in 
Scandinavia.  As  we  go  eastward  from  Western 
Europe  the  number  of  species  of  Clausilia^  as  we 
have  seen,  increases  until  we  reach  a  maximum  in 
the  Balkan  peninsula  and  the  region  of  the  Caucasus. 
Limax,  Agriolimax,  and  Amalia,  three  genera  of 
slugs,  likewise  appear  to  have  originated  in  the  same 
region  and  spread  over  Europe  from  there.  Some 
species  like  Limax  maximus  and  L.  marginatus  are 
very  ancient,  and  probably  commenced  their  wander- 
ings in  early  Tertiary  times.  In  this  manner  many 
animals  of  European  origin  have  joined  the  Oriental 


THE  ORIENTAL   MIGRATION.  285 

migrants  in  their  westward  and  also  in  their  later 
northward  travels.  In  a  similar  way  species  of  plants 
and  animals  of  Alpine  origin  might  have  joined  these 
migrants  in  their  northward  course,  and  it  is  only 
when  we  come  to  carefully  analyse  the  constituent 
parts  of  all  these  members  which  have  come  to  us  in 
England  from  the  south,  that  we  realise  the  com- 
plexity of  their  origin.  Finally,  even  the  Siberian 
migrants  mingled  with  the  later  Oriental  ones,  and 
in  some  cases  the  decision  as  to  whether  a  certain 
species  belongs  to  the  former  or  to  the  latter  migration 
becomes  a  matter  of  great  difficulty. 


SUMMARY   OF   CHAPTER  VI. 

LIKE  the  last  chapter,  this  deals  with  the  Asiatic  migrants. 
But  while  the  former  described  the  history  of  the  northern 
invasion,  those  animals  which  entered  Europe  from  the  south- 
east are  here  more  particularly  referred  to.  They  originated  in 
Central,  Southern,  and  Western  Asia.  It  is  not  easy  to  dis- 
criminate in  all  cases  between  this  Oriental  migration  and  the 
Siberian.  To  a  certain  extent,  even  an  entry  of  Northern 
Asiatic  species  has  taken  place  by  the  southern  route,  and 
•vice  versa.  On  the  other  hand,  southern  species  might  have 
come  to  Europe  by  the  southern  route— that  is  to  say,  to 
the  south  of  the  Caspian — and  also  by  the  northern,  which 
lay  to  the  north  of  that  great  inland  sea.  The  Red  Deer  is  a 
good  example.  It  arrived  on  our  continent  by  both  routes. 
However,  there  is  a  racial  difference  in  the  members  of  the  two 
migrations.  The  small  race  now  found  in  Corsica,  Sardinia, 
North-west  Africa,  and  Western  Europe,  is  probably  the  older 


286         HISTORY   OF   THE   EUROPEAN   FAUNA. 

of  the  two,  while  the  larger  one — resembling  the  American 
Wapiti  Deer — arrived  very  much  later  from  Siberia. 

The  Mammoth,  Wild  Boar,  Badger,  the  Dippers  and  Phea- 
sants, are  all  Oriental  species  which  have  come  to  us  from  the 
south-east ;  but  there  are  also  Reptiles  and  Amphibians,  and  a 
host  of  Invertebrates.  Not  all  the  animals,  for  instance,  which 
have  reached  us  in  England  from  the  south-east  are  of  Asiatic 
origin.  There  is  an  active  centre  of  distribution  in  South- 
eastern Europe  itself,  from  which  species  radiate  out  in  all 
directions.  This  fact  is  well  illustrated  by  the  genus  Clausilia. 
Species  from  this  centre,  and  also  from  the  Alps,  joined  the 
Oriental  stream  in  their  northward  course. 

In  reviewing  a  number  of  instances  of  Oriental  species  in 
Europe,  one  is  struck  by  the  peculiarity  of  their  having  ap- 
parently followed  two  distinct  routes.  All  entered  from  Asia 
Minor,  which  is  proved  to  have  been  connected  with  Greece 
until  recent  geological  times.  From  here  some  seem  to  have 
proceeded  straight  west,  others  northward.  Further  study 
reveals  the  fact  that  the  first  route  was  followed  by  a  much 
older  set  of  migrants  at  a  time  when  the  Mediterranean  area 
was  greatly  different  from  what  it  is  at  the  present  day. 
Greece  was  then  joined  to  Southern  Italy,  Sicily,  and  Tunis. 
The  latter  was  also  connected  with  Sardinia  and  Corsica,  and 
the  Straits  of  Gibraltar  did  not  exist.  Under  such  geographical 
conditions  a  direct  migration  on  land  from  Southern  Greece  to 
Spain  was  not  only  possible,  but  was  actually  undertaken  by 
a  very  large  number  of  Oriental  species. 


CHAPTER  VII. 

THE   LUSITANIAN   FAUNA. 

UNDER  the  Roman  Emperor  Augustus,  the  Spanish 
peninsula  was  divided  into  three  provinces,  one  of 
which — namely  Lusitania — occupied  a  large  portion  of 
the  present  area  of  Portugal.  The  term  "  Lusitanian  " 
is  therefore  almost  synonymous  with  Portuguese,  but 
it  has  frequently  been  applied  by  zoologists  and 
botanists  in  a  much  wider  sense,  so  as  to  vaguely 
include  the  extreme  south-west  of  Europe  without 
any  definite  limits.  Neither  do  I  propose  to  restrict 
the  term  to  everything  found  within  the  borders  of 
Portugal  For  the  sake  of  convenience,  we  may 
designate  as  Lusitanian  forms  those  animals  and 
plants  which  have  migrated  to  Central,  Southern,  or 
Northern  Europe  from  South-western  Europe.  They 
may  really  be  North-west  African  species,  or  they 
may  have  originated  on  land  which  lay  to  the  west  of 
Portugal,  and  which  is  now  mostly  buried  beneath  a 
deep  sea.  Nevertheless,  we  have  received  them  from 
the  extreme  south-western  portion  of  our  continent 
— they  have  come  to  greater  Europe  from  that 
direction. 

In  discussing  the  component  elements  of  the  British 
287 


288         HISTORY   OF   THE   EUROPEAN    FAUNA. 

fauna  and  flora  in  the  third  chapter,  I  have  already 
referred  to  the  distinguishing  characters  of  the  Lusi- 
tanian  migrants  and  to  their  distribution.  I  need 
only  repeat,  therefore,  that  these  are  now  principally 
confined  to  the  south-western  portions  of  the  British 
Islands.  The  late  Edward  Forbes  was  the  first  to 
trace  the  Lusitanian  flora  to  its  native  home.  In  his 
classical  memoir  on  the  geological  relations  of  the 
existing  fauna  and  flora  of  the  British  Isles,  he  laid 
the  foundations  of  a  new  method  of  research.  We  are 
as  yet  only  beginning  to  realise  the  far-reaching 
conclusions  obtainable  by  a  careful  study  of  the 
geographical  distribution  of  animals  and  plants, 
though  the  lines  of  investigation  were  indicated 
by  him  more  than  fifty  years  ago.  Forbes  was 
of  opinion  that  the  Lusitanian  element  in  the  British 
flora  was  of  miocene  age,  and  that  it  survived  the 
Glacial  period  on  a  now  sunken  land  to  the  south- 
west of  Ireland.  Mr.  Carpenter  and  myself  agree 
in  so  far  that  we  are  both  inclined  to  look  upon 
this  Lusitanian  flora  and  the  accompanying  fauna 
in  Ireland  as  of  pre-glacial  origin.  But  I  am 
not  quite  satisfied  that  the  Lusitanian  migration 
ceased  to  come  north  then.  It  may  have  received 
a  temporary  check ;  but  the  presence,  for  instance, 
of  the  Dartford  Warbler  {Melizophilus  undatus)  in 
the  south-east  of  England  would  seem  to  indicate 
that  its  northward  migration  took  place  in  very 
recent  times.  It  is  possible  also  that  the  very 
restricted  occurrence  of  the  Dartford  Warbler  may 


THE   LUSITANIAN    FAUNA.  289 

imply  that  it  is  gradually  withdrawing  towards  its 
centre  of  origin  from  a  former  wider  range.  Such 
an  eventuality,  as  we  have  seen,  has  actually  taken 
place  in  a  great  number  of  instances. 

It  is  not  only  in  the  British  Islands  that  we 
perceive  the  influence  of  the  Lusitanian  element. 
Scandinavia,  Russia— indeed  almost  every  part  of 
Europe — can  boast  of  some  migrants  which  have 
originated  in  South-western  Europe  or  on  the 
mysterious  lands  which  lay  beyond  it.  As  a  rule, 
however,  we  notice  a  marked  decrease  of  Lusitanian 
species  as  we  travel  eastward  from  Western  Europe. 
Nevertheless,  certain  forms  have  travelled  far  beyond 
the  confines  of  our  continent,  and  we  certainly  meet 
with  them  in  Asia  and  Northern  Africa. 

It  is  remarkable  that  we  are  apt  to  mistake  some- 
times for  Lusitanian  migrants  species  which  are  of 
Oriental  origin.  In  a  previous  paper  I  classed  such 
animals  which  had  apparently  originated  in  South- 
western Europe,  but  had  really  come  from  Asia  by  a 
circuitous  southern  route,  with  the  Lusitanians.  How- 
ever, there  is  really  no  reason  why  the  two  should 
not  be  kept  apart,  provided  we  can  discriminate 
between  the  pseudo-Lusitanians  and  the  true  ones. 
I  have  already  indicated  in  the  last  chapter  how 
these  pseudo-Lusitanian  migrants  originated. 

Supposing  an  Oriental  species  had  left  Asia  for 
Europe  in  miocene  times,  it  would  on  its  arrival  in 
Greece  have  had  to  decide  between  two  courses.  It 
could  either  advance  into  the  newly-formed  Alpine 

19 


290         HISTORY   OF   THE   EUROPEAN    FAUNA. 

peninsula  and  there  remain,  or  at  once  push  on  west- 
ward into  Southern  Italy,  Sicily,  and  Tunis,  by  means 
of  the  old  land-connections,  and  thence  into  Southern 
Spain.  The  Atlantic  communicated  at  that  time  with 
the  Mediterranean  across  the  valley  of  the  Guadal- 
quivir ;  but  that  connection  ceased  to  exist  towards 
the  end  of  the  Miocene  Epoch,  when  the  Oriental 
migrants  were  free  to  ramble  through  Spain  and  the 
whole  of  the  North  European  plain.  I  have  indicated 
on  a  previous  occasion  (a,  p.  484)  that  the  earliest 
members  of  the  Red  Deer  migration,  which  have  left 
their  traces  in  the  caves  of  Malta,  and  whose  descend- 
ants still  live  in  Corsica,  Sardinia,  and  North  Africa, 
may  have  found  their  way  to  Northern  Europe  in 
this  manner.  Many  other  Asiatic  mammals  probably 
reached  the  British  Islands  in  a  similar  way. 

I  cannot  call  to  mind  any  large  species  of 
mammal  which  we  might  reasonably  suppose  to 
have  originated  in  South-western  Europe.  Even 
among  the  smaller  ones,  few  give  us  any  definite 
clue  in  this  respect.  For  instance,  the  present  range 
of  the  genus  Myogale — a  small  Insectivore  belonging 
to  the  Mole  family  (Talpidce) — teaches  us  nothing. 
The  two  living  species  show  discontinuous  distribu- 
tion, and  are  almost  confined  to  Europe.  Myogale 
occurs  fossil  in  French  miocene  deposits,  but  is 
unknown  beyond  the  confines  of  our  continent.  It  is 
therefore  probably  of  West  European  origin.  The 
gap  between  the  South  Russian  M.  moschata  and  the 
Spanish  M.pyrenaica  is  bridged  over  in  so  far  as  we 


THE   LUSITANIAN   FAUNA.  2QI 

know  from  fossil  evidence  that  the  former  had  a 
much  wider  range  in  pleistocene  times,  being  then 
found  in  England,  Belgium,  and  Germany.  Ta!pa> 
too, —  to  which  genus  our  common  Mole  belongs, — 
seems  to  be  a  West  European  genus,  since  it  occurs 
in  French  miocene  deposits.  However,  it  would  be 
difficult  to  name  many  more  recent  genera  which  could 
be  included  in  the  area  which  I  propose  to  investigate 
in  this  chapter.  The  genus  Lepus  is  probably  not  of 
Lusitanian  origin,  but  the  sub-genus  Oryctolagus — to 
which  our  common  Rabbit  belongs — has  no  doubt  had 
its  original  home  in  that  region.  Only  two  species  of 
Lepus  {Oryctolagus)  are  known,  one  of  which — Lepus 
lacostei — has  been  met  with  in  French  pliocene 
deposits.  The  other  is  the  Rabbit  (L.  cuniculus). 
Though  generally  considered  to  have  been  intro- 
duced into  the  British  Islands,  no  reason  can  be 
brought  forward  in  favour  of  such  a  supposition, 
especially  as  it  is  known  to  have  spread  into 
Germany  in  pleistocene  times  from  South-western 
Europe.  It  occurs  in  France,  the  Spanish  peninsula, 
North-western  Africa,  and  on  some  of  the  Mediter- 
ranean islands.  Its  nearest  living  relatives,  as  we 
should  almost  expect,  are  found  in  South  America. 

Of  the  Lusitanian  Birds  I  have  already  mentioned 
the  so-called  Dartford  Warbler  {Melizophilus  undatus\ 
which  ranges  from  the  south  of  England  to  the 
extreme  south-west  of  Europe.  A  second  species 
occurs  on  the  Balearic  Islands  and  on  Corsica, 
Sardinia,  and  Sicily.  The  Andalusian  Bush-quail 


HISTORY  OF  THE   EUROPEAN   FAUNA. 

(Turnix  sylvatica)  is  probably  of  North  African 
origin,  and  has  subsequently  spread  into  Southern 
Spain  and  Portugal,  and  eastward  as  far  as  Sicily. 
It  is  an  instance  of  a  migrant  utilising  the  old 
Mediterranean  land-connections  in  the  opposite  direc- 
tion from  that  described  in  the  last  chapter. 

Two  of  our  British  Wagtails  are  very  closely 
related,  so  much  so  that  it  requires  a  very  critical  eye 
to  distinguish  them  even  at  close  range.  They  also 
frequently  interbreed.  In  their  distribution,  however, 
there  is  a  considerable  difference  between  the  White 
Wagtail  (Motacilla  alba}  and  the  Pied  Wagtail 
(M.  lugubris).  While  the  former  ranges  almost 
all  over  Europe  and  Asia,  the  latter  is  a  local 
form  resident  in  the  British  Islands,  Southern 
Scandinavia,  and  France,  and  a  winter  visitor  to 
Spain  and  North-west  Africa.  The  genus  Motacilla 
is  probably  Oriental  in  its  origin,  but  it  seems  as  if 
the  Pied  Wagtail  was  a  Lusitanian  species  which 
had  gradually  spread  northward,  only  to  return  to 
South-western  Europe  in  severe  weather  for  shelter. 

The  Bearded  Titmouse  (Panurus  biarmicus] — the 
only  representative  of  the  family  Panuridce — may 
possibly  be  a  Lusitanian  bird.  The  fact  of  its 
being  absent  from  Scandinavia  and  Northern  Russia 
is  suggestive  of  a  southern  origin.  It  is  doubtful 
whether  the  bird  occurs  on  the  south  side  of  the 
Mediterranean,  but  it  is  common  in  the  south  of 
France  and  Spain,  and  has  also  been  observed  in 
Sicily,  Greece,  and  Asia  Minor.  In  Central  Europe 


THE   LUSITANIAN   FAUNA.  293 

it  is  found  sparingly,  and  eastward  its  range  extends 
as  far  as  Turkestan. 

The  genus  FringilUi>  which  belongs  to  the  great 
family  of  the  Finches,  appears  to  be  not  only  of 
European  origin,  but,  if  the  range  of  the  species 
counts  for  anything,  I  should  feel  inclined  to  locate 
their  home  in  the  south-west.  Altogether,  five 
species  are  known.  One  of  them,  viz.,  Fringilla 
leydea,  is  confined  to  the  Island  of  Teneriffe ;  another, 
F.  madeirensis,  is  found  in  Madeira,  the  Canaries, 
and  the  Azores ;  a  third,  F.  spodiogenys,  inhabits 
North-west  Africa.  The  two  remaining  species 
have  a  much  wider  range.  F.  ccclebs — the  common 
Chaffinch — occurs  in  Europe,  while  its  range  extends 
eastward  to  Western  Siberia,  Persia,  and  Turkestan. 
The  other — F.  montifringilla,  known  as  the  Bramb- 
ling — is  more  common  in  Northern  Europe,  and 
generally  frequents  the  more  northern  latitudes  of 
Asia  as  far  as  Japan. 

It  might  be  urged  that  the  peculiar  little  blue 
Magpie  of  Spain — Cyanopolius  Cooki — should  find  a 
place  among  the  Lusitanian  species,  since  there  is 
no  bird  like  it  anywhere  else  in  Europe.  But  in 
Eastern  Siberia  there  lives  a  bird  so  closely  allied 
as  to  be  barely  distinguishable  from  it.  Neverthe- 
less, since  there  are  some  distinguishing  characters,  it 
has  received  a  distinct  name — C.  cyanus.  This  is  a 
most  interesting  and  remarkable  case  of  discontinuous 
distribution,  which  may  perhaps  be  explained  by  the 
supposition  that  the  genus  is  of  Oriental  origin,  and 


294         HISTORY  OF  THE   EUROPEAN    FAUNA. 

has  died  out  at  its  former  headquarters  in  Southern 
Asia  and  all  along  the  line  of  migration,  except  at 
the  extreme  limits  of  the  range  in  both  directions — 
east  and  west. 

As  we  go  down  in  the  scale  of  life — among  the  lower 
vertebrates  and  invertebrates — we  meet  with  a  greater 
number  of  prominent  members  of  the  Lusitanian 
migration.  The  Bullfinch,  Dipper,  and  Chough, 
which  might  be  thought  to  be  of  Lusitanian  origin, 
are,  as  I  have  shown  in  the  last  chapter,  Asiatic. 

The  European  snakes  seem  to  be  all  of  eastern 
origin,  unless  TropidonotiLS  viperinus  might  be  claimed 
as  a  Lusitanian  form.  Of  very  great  interest  from 
a  zoogeographical  point  of  view  is  our  only  European 
member  of  the  South  American  and  African  family 
Amphisb&nida.  This  species — Blanus  cinereus — is 
of  the  size  and  shape  of  an  ordinary  earth-worm, 
from  which,  however,  it  may  be  distinguished  by  its 
snake-like  wriggling  motions.  It  lives  under  stones 
in  Spain  and  Portugal,  North-west  Africa,  and 
Greece.  It  has,  therefore,  a  somewhat  similar  dis- 
tribution to  that  of  many  of  the  animals  and 
plants  referred  to  in  the  last  chapter.  But  here 
we  have  an  animal  which  has  evidently  utilised  the 
old  Mediterranean  route  described  on  p.  271,  from 
west  to  east.  Two  other  species  of  Blanus  inhabit 
Asia  Minor  and  Syria,  but  most  of  its  nearest  relations 
either  live  in  South  America  or  tropical  Africa.  In 
migrating  to  North  and  West  Africa,  its  ancestors 
probably  made  use  of  the  land-bridge  which  spanned 


THE   LUSITANIAN    FAUNA. 

the  Atlantic  in  early  Tertiary  times.  Another 
Lusitanian  Lizard — belonging  not  to  an  aberrant 
group,  but  to  the  typical  Lacertidae — is  Psammo- 
dromus  hispanicus.  It  is  rather  variable  in  colour — 
generally  of  a  brown  or  green — and  grows  to  a  length 
of  about  four  or  five  inches.  It  occurs  throughout 
the  Spanish  peninsula  and  also  in  Southern  France. 
One  of  the  handsomest  European  Lizards,  which 
reaches  almost  a  foot  in  length, — of  an  olive  colour 
with  greenish  or  mother-of-pearl  reflection,  and  with 
two  yellow  stripes  along  each  side  of  the  body,— is 
an  allied  species  (P.  algirus).  From  the  Spanish 
peninsula  it  passes  into  Southern  France  and  North 
Africa.  Two  other  species  of  the  genus  are  confined 
to  North-west  Africa. 

It  is  quite  possible  that  the  genus  Pelobates  is  of 
south-western  origin.  Of  the  two  known  species  of 
this  genus  of  Toads,  one  is  found  in  the  Central 
European  plain  and  the  other  on  the  Spanish  penin- 
sula and  in  France.  The  closely  allied  Pelodytes 
punctatus,  too,  is  confined  to  this  south-western 
district,  and  their  nearest  relations  are  found  in 
Mexico.  Similarly,  the  genus  to  which  the  Midwife 
Toad  (Alytes  obstetricans)  belongs  may  have  its 
original  home  in  that  part  of  Europe.  Of  the  two 
species,  one  is  confined  to  France,  Switzerland, 
Belgium,  and  Western  Germany,  and  the  other,  viz., 
Alytes  cisternasii,  to  Spain.  Discoglossus  pictus — a 
well-known  and  conspicuous  Toad  in  Southern 
Europe — inhabits  Spain,  Algiers,  and  Tunis,  the 


$96         HISTORY   OF   THE   EUROPEAN    FAUNA. 

islands  of  Malta,  Sicily,  Sardinia,  and  Corsica.  From 
the  general  range  of  the  family  Discoglossidtv,  as 
given  in  Mr.  Boulenger's  excellent  catalogue,  it 
appears  that  nowhere  in  the  vast  space  between 
China  and  New  Zealand  has  any  member  of  the 
family  been  discovered.  The  peculiar  genus  of 
Salamander — CJiioglossa — is  quite  confined  to  the 
Spanish  peninsula. 

The  Butterflies  Nemeobius  htcina  and  Charaxes 
jasius  may  also  have  had  their  home  in  that  south- 
western district.  To  this  migration  also  seems  to 
belong  the  genus  Gonepteryx,  which  has  so  peculiar  a 
range  in  the  British  Islands.  The  only  British  species, 
known  as  the  Brimstone  Butterfly  (Goneptcryxrhanmi\ 
occurs  in  the  south  of  England  and  in  the  south  and 
west  of  Ireland.  It  is  met  with  over  the  greater  part 
of  Europe,  and  its  range  extends  into  Asia  Minor  and 
Northern  India,  and  then  it  reappears  again  in  dis- 
tinct varieties  in  Japan  and  the  Amur  district.  Three 
other  species  of  Gonepteryx  are  known  from  Tibet 
and  India,  and  one  (G.  cleopatrd]  from  Southern 
Europe  and  Northern  Africa.  All  the  remaining 
species  inhabit  the  west,  viz.,  Brazil,  Mexico,  and 
Venezuela.  That  the  genus  has  migrated  from 
America  eastward  to  Europe  appears  to  be  more 
probable  than  a  migration  in  the  opposite  direction. 
At  any  rate,  that  an  exchange  of  species  between 
the  south-western  portion  of  the  Holafctic  Region 
and  the  Neotropical  area  took  place  is  indicated  by 
the  fact,  not  only  that  a  variety  of  G.  cleopatra  has 


THE   LUSITANIAN    FAUNA.  2Q7 

been  found  in  Madeira,  but  also  that  the  Canary 
Islands  possess  a  distinct  form  of  Goneptetyx,  viz., 
G.  cleobule. 

Dr.  Kobelt  has  given  us  such  an  exhaustive  memoir 
on  the  characteristic  Mollusca  of  the  different 
zoogeographical  provinces  of  Europe,  that  we  are 
particularly  well  informed  as  regards  that  group  of 
Invertebrates.  He  tells  us  that  the  group  Torquilla 
of  the  genus  Pupa — which  is  a  small  chrysalis-like 
snail — is  especially  characteristic  of  the  Pyrenees, 
Spain,  and  Portugal.  In  a  certain  measure  they 
replace  there  the  Clausilice  which,  as  we  have  seen  in 
the  last  chapter,  have  come  from  the  east  and  are 
almost  entirely  absent  in  the  south-west  of  Europe. 
Of  about  seventy  species  of  Torquilla,  the  larger 
number  are  confined  to  this  district,  and  some,  which 
like  Pupa  (Torquilla}  granum,  range  eastward,  have 
travelled  along  the  old  Mediterranean  highway,  via 
Algiers,  Sicily  and  Greece,  to  Asia  Minor.  They 
are  still  found  along  the  whole  of  this  route. 

Sifnilarly,  we  are  told  by  the  same  author,  that 
Gonostoma — a  group  of  the  large  genus  Helix — has  a 
number  of  species  in  the  same  south-western  district, 
while  only  one,  viz.,  Helix  obvoluta,  occurs  in  England 
and  Germany,  and  two  in  the  Alps.  Southward  we 
again  find  many  representatives  crossing  over  to 
North  Africa,  among  which  Helix  lenticula  has  a 
similar  range  to  Piipa  granum,  which  I  have  just 
referred  to.  The  Alpine  sub-genus  Cainpylcea  is 
quite  absent  in  the  Lusitanian  district. 


298         HISTORY  OF   THE   EUROPEAN    FAUNA. 

Among  our  own  British  testaceous  Land  Mollusca, 
several  Helices,  viz.,  Helix  pisana,  ericetorum,  virgata, 
acuta,  fusca,  rotundata,  aculeata,  and  probably  many 
others,  have  come  to  us  from  the  south-west.  The 
species  of  Hyalinia  are  undoubtedly  of  very  remote 
origin,  and  it  would  be  futile  at  the  present  state  of 
our  knowledge  to  speculate  as  to  their  home.  Some 
of  our  species  may  possibly  be  of  British  origin. 
Balea  perversa  is  probably  a  south-western  species, 
and  certainly  Pupa  anglica,  which  is  quite  confined  to 
Western  Europe. 


•••••••••••••MMraraEMW^ 

FIG.  1 8. — The  Spotted  Slug  (Geomalacus  maculosus}. 

Much  more  characteristic  of  South-western  Europe, 
however,  than  these  land-shells  are  some  of  the  slugs. 
The  peculiar  genus  Geomalacus  is  almost  entirely 
confined  to  Portugal.  One  species,  which  I  have 
had  several  occasions  to  refer  to  in  illustration  of  the 
term  "discontinuous  distribution,"  ranges  far  beyond 
the  confines  of  that  country.  This  is  Geomalacus 
maculosus  (Fig.  18),  first  discovered  in  the  south- 
west of  Ireland,  and  more  recently  also  in  Portugal. 
Although  careful  search  has  been  made  for  it  in 


THE   LUSITANIAN    FAUNA.  299 

other  parts  of  the  British  Islands,  this  slug  has  only 
been  found  in  the  portion  of  Ireland  just  indicated. 
Within  the  last  few  years  I  have  taken  it,  up  to  a 
height  of  over  a  thousand  feet,  on  the  promontory 
north  of  the  Kenmare  River,  also  from  sea-level  up 
to  a  considerable  height  near  Glengarirf,  and  more 
recently  Messrs.  Praeger  and  Welch  discovered  it  in 
abundance  near  the  town  of  Kenmare.  But  beyond 
this  rather  circumscribed  area  in  the  counties  of  Cork 
and  Kerry  it  does  not  occur  (vide  Fig.  19).  Several 
Portuguese  species  of  this  interesting  genus  have 
since  been  added  to  science  by  Dr.  Simroth  and 
others.  Dr.  Simroth,  too,  has  promulgated  the  view 
that  the  genus  Arion — to  which  our  common  brown 
garden  slug  belongs — is  of  Lusitanian  origin.  Indeed, 
the  number  of  species  of  Arion  diminishes  as  we 
leave  that  province,  though  one  extends  beyond  the 
borders  of  Europe  into  Siberia.  The  same  number  of 
species,  viz.  five,  occur  in  Germany  and  in  England. 
Testacella — a  slug-like  mollusc — which  lives  under- 
ground on  earthworms,  and  of  which  genus  three 
species,  viz.  T.  mangei,  T.  haliotidea,  7\  scutulum, 
are  known  to  inhabit  the  British  Islands,  is  another 
Lusitanian  animal.  All  the  species  are  confined  to 
\Vestern  Europe  and  North  Africa  ;  they  do  not  even 
reach  Germany  or  Switzerland. 

I  have  had  occasion  to  mention  once  before  an  ex- 
tremely interesting  genus  of  blind  Woocllouse,  viz., 
Platyarthnis.  Like  Testacella,  it  lives  underground, 
and  also  resembles  it  in  its  general  range.  Its  distribu- 


300         HISTORY  OF  THE   EUROPEAN   FAUNA. 


tion  is  therefore  of  particular  interest.  It  is  difficult  to 
conceive  that  Platyarthrus,  from  its  peculiar  mode  of 
life,  could  have  crossed  any  formidable  barrier,  such 


FIG.  19. — Map  of  the  British  Islands  on  which  the  geographical  distri- 
bution of  Geomalaats  maculosus  is  indicated  in  black. 

as  even  a  narrow  straits  of  sea.  Its  occurrence  in 
Spain  and  North  Africa  indicates,  therefore,  that  the 
Straits  of  Gibraltar  did  not  exist  at  the  time  when 


THE   LUSITANIAN   FAUNA.  3OI 

it  undertook  the  migration  southward,  just  as  the 
English  Channel  and  the  Irish  Sea  could  not  have 
been  there  when  it  wandered  to  England  and  Ireland. 
The  species  which  occurs  in  the  south  of  England 
has  a  wide  range  in  Ireland,  and  reaches  in  Scotland 
its  most  northern  European  limit  of  distribution. 
Platyarthrus  is  only  one  of  the  Lusitanian  genera  of 
woodlice.  In  Ireland — chiefly  on  the  west  coast — we 
also  find  a  brilliantly  coloured  Woodlouse,  which  is 
absent  from  Great  Britain, viz.  Metoponorthuscingendus. 
It  reappears  again  on  the  Continent  in  the  south  of 
France.  Its  range  is  therefore  suggestive  of  a  Lusi- 
tanian origin  ;  and  indeed,  when  we  examine  the 
general  distribution  of  the  genus  Metoponorthus,  we 
find  that  out  of  the  forty-four  known  species,  fully 
one-half  arc  confined  to  Western  Europe  and  North 
Africa. 

My  friend  and  colleague,  Mr.  Carpenter,  informs 
me  that  among  the  Irish  Spiders  he  is  acquainted 
with,  the  following  are  to  be  looked  upon  as  Lusi- 
tanian species  : — 

Dysdera  crocota.  Agroeca  celans. 
Oonops  pulcher.  do.     gracilipes. 

Tegenaria  hibernica.  Teutana  grossa. 

Theridion  aulicum.  Cnephalocotes  curtus. 

Lasseola  inornata.  Porrhomma  myops. 

Of  the  Coleoptera,  the  genera  Trichis,  Glycia,  and 
Singilis,  all  belonging  to  the  Running  Beetles  (Cara- 
bidce],  are  almost  confined  to  the  Spanish  penin- 
sula. 


3O2         HISTORY  OF   THE   EUROPEAN   FAUNA. 

The  beetles  Rhopalomesites  Tardyi,  Eurynebria 
complanata,  and  Otiorrhynchus  auropimctatus  also 
belong  to  this  fauna,  as  also  the  Earthworms  Allolo- 
bophora  veneta  and  A.  Georgii,  and  the  Millipede 
Polydesmus  gallicus. 

It  will  be  evident  to  every  one  from  these  few 
instances  of  Lusitanian  species,  that  somewhere  in 
South-western  Europe  and.  North-western  Africa, 
and  also,  perhaps,  in  a  larger  now  submerged  western 
land-area,  there  existed  an  active  centre  of  develop- 
ment, from  which  animals  spread  in  all  directions. 

If  the  presence  of  Platyarthrus  in  North-west  Africa 
proves  that  the  Straits  of  Gibraltar  had  come  into 
existence  after  its  southward  migration,  it  also 
suggests  that  the  ancestral  home  of  this  woodlouse 
was  in  the  Spanish  peninsula.  Whether  this  sup- 
position is  correct  or  not,  does  not  affect  the  Straits 
of  Gibraltar  problem,  for  in  a  migration  northward 
into  Spain  from  Morocco  a  land-connection  would  be 
equally  necessary.  Almost  every  group  of  vertebrates 
and  invertebrates  furnishes  instances  of  species  which 
must  have  crossed  the  Straits  on  dry  land.  Many 
naturalists  have  come  to  this  conclusion,  and  have 
clearly  expressed  their  views  on  the  subject  At 
the  commencement  of  the  present  period,  says  Mr. 
Bourguignat  (p.  354),  the  north  of  Africa  was  a 
peninsula  of  Spain,  the  Straits  of  Gibraltar  did  not 
exist,  and  the  Mediterranean  communicated  by  the 
Sahara  with  the  Atlantic. 

The  faunas  of  North-west   Africa  and  the  south- 


THE   LUSITANIAN   FAUNA.  303 

western  portion  of  our  continent  are  so  closely 
related,  that  an  uninterrupted  intercourse  by  land 
must  have  existed  for  a  very  long  period.  The 
Mediterranean,  however,  throughout  the  Tertiary 
period — at  any  rate  since  miocene  times — must  have 
had  almost  constant  communication  with  the  Atlantic. 
According  to  Professor  Suess,  this  was  the  case.  The 
Atlantic  was  joined  with  the  Mediterranean  across 
the  valley  of  the  Guadalquivir  during  the  Miocene 
Epoch,  so  that  Andalusia  must  have  belonged  to 
North  Africa  in  those  days.  The  Straits  of  Gibraltar 
are  supposed  to  have  been  formed  in  the  next  epoch. 
I  have  already  expressed  my  disagreement  with  that 
theory  from  a  zoogeographical  point  of  view.  The 
old  Guadalquivir  connection  probably  persisted  much 
longer, — though  interrupted  by  temporary  periods  of 
a  partial  retreat — so  as  to  uncover  sufficient  land  to 
allow  of  an  interchange  during  miocene  as  well  as 
pliocene  times  between  the  European  and  North 
African  faunas.  It  is  in  this  way,  perhaps,  that  some 
of  the  members  of  the  Alpine  fauna  have  reached 
Spain  by  way  of  Corsica,  Sardinia,  and  North-western 
Africa,  and  vice  versd.  The  Balearic  Islands  were 
then  connected  with  Spain;  and  we  find  there  many 
curious  survivals  which  have  long  ago  become  extinct 
on  the  mainland 

That  the  Straits  of  Gibraltar  are  only  of  recent 
formation  has  been  suggested  on  zoogeographical 
evidence  by  Bourguignat,  Simroth,  Kobelt,  and  many 
others.  Dr.  Kobelt  believes  that  the  former  land- 


304         HISTORY  OF  THE   EUROPEAN    FAUNA. 

connection  between  the  south  of  Spain  and  Morocco 
was  much  wider  than  is  generally  assumed,  and  that 
the  coast-line  stretched  from  Oran  in  Algeria  straight 
across  to  Cartagena  in  Spain  (^,  ii.,  p.  228). 

My  allusions  to  the  lands  lying  beyond  the  Lusi- 
tanian  province,  refer  chiefly  to  the  Canary  Islands 
and  Madeira.  Whatever  doubts  Dr.  Wallace  had 
on  the  subject  of  their  former  connection  with 
Morocco,  it  cannot  be  denied  that  they  used  to  be 
of  much  larger  extent,  especially  in  miocene  and 
pliocene  times.  It  seems  extremely  probable  that 
these  islands  formed  part  of  the  mainland  of  North 
Africa  until  comparatively  recently,  and  that  they 
are  the  last  traces  of  a  sunken  continent  which  united 
Africa  and  South  America.  A  discussion  of  this 
problem,  however,  must  be  deferred,  as  it  is  a  com- 
plicated one,  and  one  which  would  lead  me  alto- 
gether outside  the  scope  of  this  little  volume.  I 
hope  I  shall  have  an  opportunity  to  publish  my 
views  on  this  subject  before  long,  meanwhile  the 
reader  must  content  himself  with  this  mere  state- 
ment. 

During  the  greater  portion  of  the  Miocene,  and  I 
think  for  part  of  the  Pliocene  Epoch  too,  the  advance 
of  the  Lusitanian  species  eastward  was  barred  on 
the  continent  of  Europe  by  an  arm  of  the  sea  which 
stretched  northward  along  the  Rhone  valley  from 
the  Mediterranean.  The  Lusitanian  forms  which 
originated  in  Southern  Spain  were  able  to  travel  east 
during  these  times  by  way  of  North-west  Africa, 


THE  LUSITANIAN   FAUNA.  3°5 

Sicily,  Southern  Italy,  and  Greece;  and  it  is  possible 


FIG.  20.— The  Strawberry-tree  (Arbutus  unedo)  in  its  native  habitat  in 
the  south-west  of  Ireland.     (From  a  photograph  by  Robert  Welch. ) 

that  some  may  have  reached  the  Alps  in  this  manner, 

and    Eastern    Europe    generally.      That    the    Lusi- 

20 


306 


HISTORY  OF   THE   EUROPEAN    FAUNA. 


tanian  centre  was  never  a  very  active  one  compared 
with,  for  instance,  the  Oriental  is  indicated  by  many 
distributional  facts.  It  is  difficult  to  understand, 
however,  why  the  Oriental  species,  on  the  whole, 
have  migrated  so  far  west,  while  few  Lusitanians  have 
gone  very  far  east.  This  seems  to  have  been  noted 


FlG.  21. — The  Irish  Spurge  {Euphorbia  hiberna]  in  its  native  habitat 
in  the  south  of  Ireland.     (From  a  photograph  by  Robert  Welch.) 

particularly  in  the  case  of  the  flora.  Mr.  Bonnet  drew 
attention  to  the  fact  that  in  Tunis  there  are  none  of  the 
absolutely  characteristic  plants  of  Morocco  and  Spain, 
while  the  Oriental  flora  is  represented  by  a  good 
many  species.  Lusitanian  species  have  spread  chiefly 
southward  into  North  Africa,  and  northward  into 
France,  the  British  Islands,  and  even  Scandinavia. 


THE   LUSlTANlAN    FAUNA.  30? 

As  I  have  mentioned  in  the  third  chapter,  there  are  a 
good  many  species  of  Lusitanian  origin  in  the  British 
Islands.  However,  we  have  only  a  mere  remnant 
of  what  we  ought  to  have,  had  the  climate  been  less 
trying.  It  is  probable,  too,  that  the  submergence 
destroyed  a  good  many  plants  and  the  insects  depen- 
dent on  them.  That  the  Lusitanian  fauna  is  very 
ancient  in  the  British  Islands  is  proved  by  the  fact 
of  the  discontinuous  distribution  of  so  many  species. 
A  greater  number  survived  in  Ireland  than  in 
England. 

Altogether — and  this  was  strongly  urged  by 
Edward  Forbes — the  Lusitanian  element  is  the 
oldest  of  the  components  of  our  fauna,  and  it  must 
have  poured  into  the  British  Islands  for  many  geo- 
logical periods  almost  without  cessation.  The  same 
author,  in  his  classic  essay,  refers  especially  to  the 
Lusitanian  flora,  two  prominent  members  of  which 
are  the  British  plants,  Arbutus  unedo  (Fig.  20,  p.  305) 
and  EupJiorbia  hiberna  (Fig.  21,  p.  306).  The  former 
has  a  wide  range  in  the  Mediterranean  region,  and 
occurs  in  the  British  Islands  only  in  the  south-west 
of  Ireland.  The  Spurge,  on  the  other  hand,  is  also 
found  in  the  south-west  of  England,  besides  Ireland 
and  Southern  Europe, 


308         HISTORY   OF   THE   EUROPEAN    FAUNA. 


SUMMARY  OF  CHAPTER  VII. 

The  term  "  Lusitanian "  is  in  this  chapter  employed  in  the 
wide  sense,  as  indicating  the  South-west  of  Europe  and  North- 
western Africa.  From  this  centre,  and  probably  also  from  a 
now  sunken  land  which  lay  to  the  west  of  it,  issued  a  fauna  and 
flora  of  which  we  have  abundant  evidence  in  our  own  islands, 
especially  in  Ireland.  Edward  Forbes  held  that  the  Lusitanian 
element  of  the  British  flora  was  of  miocene  age,  and  that  it 
survived  the  Glacial  period  in  this  country. 

At  the  time  when  the  Straits  of  Gibraltar  did  not  exist,  and 
when  there  was  free  land  communication  between  Asia  Minor, 
Greece,  and  Tunis,  many  Oriental  species  migrated  westward 
by  this  ancient  Mediterranean  route  as  far  as  Spain.  They 
would  then  have  invaded  the  more  central  parts  of  Europe  from 
the  south-west,  without  however  being  of  Lusitanian  origin.  Of 
the  true  Lusitanian  mammals  a  typical  example  is  the  Rabbit. 
Then  we  have  a  few  birds  and  several  interesting  reptiles  and 
amphibians.  The  genus  to  which  the  Brimstone  Butterfly 
belongs  is  also  of  south-western  origin.  A  number  of  Mollusca 
are  mentioned  which  from  their  range  likewise  indicate  a  Lusi- 
tanian origin.  Most  of  our  British  Slugs  and  many  of  our 
larger  Snails  belong  to  this  group. 

All  these  are  merely  a  small  remnant  of  what  we  received 
from  South-western  Europe  during  the  Miocene  and  Pliocene 
Epochs.  But  they  spread  into  many  parts  of  Europe,  and  a 
few  even  crossed  into  Asia.  The  antiquity  of  the  Lusitanian 
element  in  our  fauna  is  especially  indicated  by  the  frequent 
recurrence  of  "discontinuous  distribution"  among  the  species 
belonging  to  that  section. 


CHAPTER  VIII. 

THE    ALPINE    FAUNA. 

WE  are  told  by  Sir  Archibald  Geikie  (p.  851)  that 
"  from  the  Pyrenees  eastwards,  through  the  Alps  and 
Apennines  into  Greece,  and  the  southern  side  of 
the  Mediterranean  basin,  through  the  Carpathian 
Mountains  and  the  Balkan  into  Asia  Minor,  and 
thence  through  Persia  and  the  heart  of  Asia  to 
the  shores  of  China  and  Japan,  a  series  of  massive 
limestones  has  been  traced,  which,  from  the  abun- 
dance of  their  characteristic  foraminifera,  have  been 
called  the  Nummulitic  Limestone.  Unlike  the  thin, 
soft,  modern-looking,  undisturbed  beds  of  the  Anglo- 
Parisian  area,  these  limestones  attain  a  depth  of 
sometimes  several  thousand  feet  of  hard,  compact, 
sometimes  crystalline  rock,  passing  even  into  marble, 
and  they  have  been  folded  and  fractured  on  such  a 
colossal  scale  that  their  strata  have  been  heaved  up 
into  lofty  mountain  crests  sometimes  10,000,  and 
in  the  Himalaya  range  more  than  16,000  feet  above 
the  sea."  "  Nowhere  in  Europe,"  continues  the 
same  author  (p.  860),  "do  oligocene  strata  play 
so  important  a  part  in  the  scenery  of  the  land,  or 
present  on  the  whole  so  interesting  and  full  a  picture 

309 


3IO         HISTORY  OF  THE   EUROPEAN    FAUNA. 

of  the  state  of  Europe  when  they  were  deposited,  as 
in  Switzerland.  Rising  into  massive  mountains,  as 
in  the  well-known  Rigi  and  Rossberg,  they  attain  a 
thickness  of  more  than  6000  feet."  "By  far  the  larger 
portion  of  these  strata  is  of  lacustrine  origin.  They 
must  have  been  formed  in  a  large  lake,  the  area  of 
which  probably  underwent  gradual  subsidence  during 
the  period  of  deposition,  until  in  Miocene  times  the 
sea  once  more  overflowed  the  area." 

From  these  remarks  by  our  most  eminent  British 
geologist,  we  gather  that  in  early  Tertiary  times 
much  of  the  present  area  of  Switzerland  was  either  a 
sea  or  a  large  freshwater  lake.  The  Alps  were  then 
appearing  in  this  sea,  probably  as  a  chain  of  islands, 
and  in  the  beginning  of  the  Miocene  Epoch  one  large 
elongated  island  had  made  its  appearance — the  future 
European  Alps.  I  have  already  mentioned  that  the 
Miocene  Sea  skirted  the  Alps  from  the  Mediterranean 
up  the  valley  of  the  Rhone  and  along  its  northern  and 
eastern  margin.  Miocene  marine  deposits  are  also 
known  from  the  Southern  Alps  and  the  east  side  of 
the  Apennines,  from  Corsica,  Sardinia,  and  Malta. 
No  trace,  however,  of  them  has  been  noticed  any- 
where along  the  ^Egean  Sea  or  on  the  Balkan 
peninsula.  The  Alps  were  therefore  connected  to  the 
east  with  the  outliers  of  the  Balkan  Mountains,  and 
in  this  way  with  Asia,  from  which  they  received  so 
large  a  proportion  of  their  fauna  and  flora.  In 
pliocene  times  the  sea  still  washed  the  southern  shore 
of  the  Alps,  but  to  the  north  dry  land  gradually 


THE   ALPINE   FAUNA.  311 

supplemented  the  sea,  and  the  Alpine  fauna  and 
flora  were  able  to  pour  into  the  plain.  It  was  then 
that  the  Arctic  species — which  we  have  learned  had 
migrated  into  Northern  Europe  from  the  north — found 
their  way  to  the  Alps.  In  a  similar  way  Lusitanian 
forms — in  fact,  species  from  almost  all  parts  of 
Europe — were  now  free  to  wander  to  the  newly  opened 
up  peninsula  which  had  become  part  of  the  main- 
land of  Europe.  The  typical  Siberian  species  had 
not  entered  our  continent  at  that  time,  it  was  not  till 
much  later — not  until  the  middle  of  the  Pleistocene 
Epoch — that  they  made  their  appearance  at  the  foot 
of  the  Alps,  but,  as  we  shall  see  later  on,  it  is  doubt- 
ful whether  many  of  these  species  ever  reached  the 
mountains. 

The  fauna  of  the  Alps,  and  also  the  flora,  is 
therefore  made  up  of  a  number  of  component  ele- 
ments. In  the  first  place  we  have  the  Oriental  element 
— the  migrants  from  Central  and  Southern  Asia. 
When  the  nature  and  origin  of  the  Oriental  fauna  in 
Europe  was  discussed,  reference  was  made  to  the  fact 
(p.  272)  that  we  can  distinguish  an  older  from  a  newer 
Oriental  migration.  Both  of  these  have  entered  the 
Alps.  As  we  might  anticipate,  many  of  the  older 
Oriental  migrants  have  developed  into  new  species, 
laying  the  foundation  of  an  indigenous  Alpine 
element.  From  the  fact  that  they  set  foot  on  the 
Alpine  peninsula,  it  might  be  expected  that  there 
could  have  existed  no  mountains  to  speak  of.  The 
climate  was  mild  and  damp.  Now  as  the  country 


312          HISTORY  OF   THE   EUROPEAN    FAUNA. 

rose,  and  a  formidable  mountain  range  took  the  place 
of  a  hilly  island,  the  whole  fauna  was  lifted  up 
and  transferred  to  entirely  different  conditions.  A 
modification  of  their  structure  to  suit  the  new  sur- 
roundings was  therefore  to  be  anticipated,  and  that  is 
exactly  what  occurred,  though  not  in  all  cases. 

Take,  for  example,  the  goats  which  are  of  Asiatic 
origin.  Every  one  has  heard  of  the  "  Steinbock," — the 
Alpine  mountain  goat  (Capra  ibex) — though  very  few 
have  seen  it  in  its  native  haunts,  where  it  is  now  on 
the  verge  of  extinction.  A  closely  allied  species 
(Capra  sibirica)  inhabits  the  Altai'  and  Himalayan 
Mountains;  a  third  species  (Capra  sinaitica)  lives  in 
Palestine,  and  has  entered  Egypt  by  way  of  the 
Sinaitic  peninsula.  Another  (C.  cegagrus)  occurs  in 
Asia  Minor,  Persia,  the  island  of  Crete,  and  some  of 
the  Cyclades.  This  exemplifies  what  I  remarked  in  the 
last  chapter  about  ihe  former  land-connection  between 
Greece  and  the  Asiatic  continent.  Finally,  we  have 
the  Pyrenean  Goat  (Capra  pyrenaica\  which  is  found 
in  the  Pyrenees,  the  higher  ranges  of  Central  Spain, 
in  Andalusia,  and  Portugal,  thus  indicating  that  it 
probably  reached  the  Spanish  peninsula  from  the 
south  by  means  of  the  old  Sicilo-Algerian  highway, 
especially  as  remains  of  the  species  occur  in  the 
cave  deposits  of  Gibraltar.  The  ancestors  of  the 
goat-like  Antelope — known  as  the  Chamois  (Rupicapra 
tragus] — no  doubt  also  came  from  Asia.  The  genus  is 
not  represented  there,  but  Nemorhcedus  and  Budorcas 
are  allied  Asiatic  genera,  while  the  Rocky  Mountain 


THE   ALPINE   FAUNA.  313 

Goat  (Haploceros  montanus)  also  has  certain  affinities 
with  the  Chamois.  Besides  the  Alps,  the  latter 
occurs  in  the  Caucasus  and  the  Pyrenees.  The 
Alpine  Marmot  (Arctomys  mar  motto)  is  sometimes 
quoted  as  owing  its  origin  to  the  Siberian  pleistocene 
migration,  but  it  does  not  occur  in  Siberia  now,  nor 
is  there  any  palaeontological  evidence  that  it  was  ever 
found  there.  The  genus  Arctomys  is  an  ancient 
Asiatic  genus,  to  judge  from  its  general  range. 
Only  two  species  occur  in  Europe,  one  of  which, 
the  true  Siberian  Marmot  (A.  bobac\  just  enters  our 
continent  in  the  east — or  rather,  it  is  one  of  those 
species  which  came  to  us  in  pleistocene  times  and  are 
now  gradually  retreating  towards  their  native  land. 
The  genus,  however,  is  probably  not  of  Siberian 
origin.  No  less  than  seven  other  species  occur  in 
Asia,  six  of  which  are  confined  to  Central  Asia  and 
the  Himalayan  Mountains,  while  four  have  wandered 
to  North  America.  The  sequence  of  events,  therefore, 
was  that  the  ancestor  of  Arctomys  marmotta  probably 
came  to  the  Alps  direct  from  Central  Asia  by  way 
of  Asia  Minor  in  miocene  or  pliocene  times.  It  has 
since  become  modified  into  a  distinct  species,  and  has 
spread  to  the  European  plain,  where  it  occurs  fossil 
in  pleistocene  strata,  and  to  the  Carpathian  Moun- 
tains and  the  Pyrenees. 

The  great  majority  of  species  of  the  large  genus 
Microtus  (Arvicola)  are  Asiatic,  and  there  can  be  little 
doubt  that  it  has  originated  in  that  continent.  There 
is  one  species  of  Vole  {Microtus  nivalis)  which  occurs 


3 14         HISTORY   OF  THE   EUROPEAN    FAUNA. 

in  the  high  Alps,  and  which  has  been  supposed  to  be  a 
typical  Alpine  form.  It  is  known,  however,  to  occur 
also  in  North  Italy  and  in  Bohemia,  while  Microtus 
leucurus  of  the  Pyrenees  is  identical  with  this 
species.  But  its  range  is  by  no  means  confined  to 
Europe,  for  it  has  also  been  discovered  in  Syria 
and  Palestine,  while  a  closely  allied  form  exists  in 
the  Himalayan  Mountains.  This  shows  clearly  that 
the  species  has  migrated  to  the  Alps  from  Asia 
Minor.  That  this  migration  may  have  taken  place 
at  an  early  period — at  a  time  when  Sardinia  and 
Corsica  were  still  connected  with  Southern  Europe — 
is  indicated  by  the  occurrence  of  an  extinct  Vole 
{Microtus  brecciensis)  in  Sardinian  and  Corsican 
pleistocene  (?)  deposits. 

All  the  Alpine  species  mentioned  except  the 
Chamois  can  be  easily  traced  to  their  former  Asiatic 
home.  But  even  it  has  its  nearest  relations  in  Asia. 
I  might  also  refer  to  another  Vole  (Evotomys  Nageri) 
which  is  practically  confined  to  the  Alps  and 
Northern  Italy,  and  which  has  probably  originated 
there,  though  most  of  its  nearest  relations  are  either 
Asiatic  or  North  American  species. 

But  besides  these  Asiatic  immigrants  and  their 
modified  descendants  we  have  a  small  truly  native 
Alpine  mammalian  fauna.  Sqrexalpinus — the  Alpine 
Shrew — occurs  only  in  the  Alps,  the  Harz  Mountains, 
Pyrenees,  and  Carpathians.  The  genus  has  been 
found  in  European  eocene  strata, — in  vastly  older 
deposits  in  our  own  continent  than  elsewhere, — so 


THE   ALPINE   FAUNA.  315 

that  it  is  extremely  probable  that  it  has  originated 
there.  It  may  then  have  developed  a  new  centre  of 
distribution  in  the  newly-formed  Alps,  where  both 
Sorex  alpinus  and  5.  minutus  (pygmceus)  have  their 
home.  From  there  they  again  spread — perhaps 
already  in  miocene  times — to  Asia  and  North 
America,  where  a  large  number  of  new  species 
originated.  It  seems  to  me  even  probable  that  one 
of  these  Asiatic  species  of  Sorex,  viz.  S.  araneus 
(vulgar is}>  subsequently  migrated  towards  the  old 
home  of  its  forefathers,  since  we  find  it  more  or  less 
confined  to  Central  and  Northern  Asia  and  Northern 
Europe. 

Though  the  origin  of  the  Alpine  Hare  has  already 
been  referred  to  and  fully  discussed  in  a  previous 
chapter  (p.  148),  the  conclusions  arrived  at  may  be 
once  more  repeated.  The  Alpine  Hare  (Lepus 
variabilis]  is  of  Arctic  origin.  It  spread  southward 
into  Europe,  North  America,  and  Asia  in  early  glacial 
times,  and  reached  our  continent  from  Spitsbergen  by 
means  of  a  direct  land-connection  with  Lapland. 
The  Scandinavian  peninsula  was  then  separated  from 
Russia,  but  connected  with  Scotland  and  Ireland 
(Fig.  13,  p.  170).  Since  England  was  then  united  to 
France,  the  Alpine  Hare  was  able  to  invade  western 
continental  Europe  and  all  the  mountain  ranges. 
Its  range  is  very  discontinuous,  small  colonies  being 
scattered  all  over  the  mountainous  parts  of  the 
Northern  Hemisphere,  while  the  European  Hare — 
a  closely  allied  species — occurs  in  the  plain,  and  now 


316         HISTORY  OF   THE   EUROPEAN    FAUNA. 

occupies  to  some  extent  the  former  haunts  of  the 
Alpine  Hare  (cf.  Fig.  8,  p.  137).  Might  not  the 
European  Hare,  as  suggested,  possess  some  advan- 
tages which  enabled  it  to  drive  the  other  into  more 
inaccessible  parts,  thus  producing  the  peculiarity 
of  range?  The  present  distribution  of  the  Alpine 
and  the  European  Hare  (L.  Europceus)  appears  to  me 
to  strongly  support  such  an  assumption.  It  is  not 
the  cold  which  has  driven  the  Alpine  Hare  to  the 
Alps;  and  its  presence  there  is  not,  as  is  often  sup- 
posed, a  "standing  testimony  of  a  former  arctic 
climate"  in  Europe,  but  merely  the  necessary  con- 
sequence of  the  weaker  species  being  thrust  into 
less  accessible  regions  by  a  stronger  rival. 

Muscardinus  avellanarhis, — the  common  Dormouse, 
— though  by  no  means  confined  to  the  Alps,  has  prob- 
ably originated  there.  It  is  found  up  to  a  height 
of  nearly  5000  feet  in  these  mountains,  and  is  spread 
over  Europe  at  nearly  equal  distances  from  the  Alps 
in  all  directions.  Being  absent  from  Ireland,  Scot- 
land, Norway,  and  Northern  Russia,  it  seems  as 
if  it  had  only  diffused  northward  in  more  recent 
times. 

The  closely  allied  genus  Myoxus  is  likewise  of 
European  extraction,  some  species  being  known  from 
French  eocene  deposits. 

There  are  only  a  few  typically  Alpine  Birds.  One  of 
these  is  the  Alpine  Accentor  (Accentor  collaris],  which 
on  rare  occasions  visits  England,  and  Northern 
Europe  generally.  It  is,  however,  by  no  means 


THE   ALPINE   FAUNA.  317 

peculiar  to  the  European  Alps ;  a  variety  of  this 
species  occurs  in  Central  Asia,  Eastern  Siberia,  and 
Japan.  The  only  other  Accentor  inhabiting  our 
continent  is  the  Hedge  Accentor  (A.  modularis), 
which  is  resident  over  the  greater  part  of  it,  and 
also  in  North  Africa  and  the  Mediterranean  Islands. 
It  also  extends  its  range  across  the  ^gean  Sea  to 
Asia  Minor,  so  that  really  not  a  single  Accentor  is 
peculiar  to  Europe. 

Both  the  European  species  are  evidently  old  forms, 
and  the  genus,  as  might  be  expected,  is  certainly 
Asiatic.  No  less  than  ten  other  species  of  Accentor 
are  known,  all  of  which  are  confined  to  Central  Asia 
and  the  Himalayan  Mountains,  and  are  therefore  all 
Holarctic.  I  may  mention  that  much  difference  of 
opinion  still  exists  as  to  the  true  zoological  position 
of  this  anomalous  genus.  It  has  been  located  in 
several  different  families  by  various  ornithologists, 
but  has  not  yet  found  a  permanent  resting-place. 
Another  bird  generally  considered  to  be  peculiar  to 
Switzerland  is  the  Alpine  Chough  {Pyrrhocorax 
alpinus),  but  its  range  extends  across  Asia  Minor 
to  the  Himalayas.  Whether  the  European  Chough 
should  not  form  a  distinct  genus  is  a  matter  of 
opinion.  Some  of  our  leading  ornithologists,  like 
Dr.  B.  Sharpe,  are  inclined  to  separate  it  from 
Pyrrhocorax  ;  however,  there  is  no  doubt  that  it  is 
closely  related  to  the  Alpine  Chough,  whatever  view 
we  may  take  of  the  generic  distinctness.  It  inhabits 
principally  Western  and  Southern  Europe,  also 


318          HISTORY   OF   THE    EUROPEAN    FAUNA. 

North  Africa ;  and  its  range  extends  eastward  to 
the  Himalayas,  China,  and  Eastern  Siberia.  If  any 
doubt  still  existed  as  to  the  Asiatic  origin  of  the 
Choughs,  it  may  be  noted  that  the  only  two  other 
closely  allied  genera,  viz.,  Corcorax  and  Podoces,  live 
in  Australia  and  Central  Asia  respectively. 

There  are  two  other  birds  to  which  I  should  like 
to  refer.  These  are  the  Rock  Sparrow  and  the  Alpine 
Snow  Finch.  The  first  of  these  (Petronza  stultd)  is 
by  no  means  peculiar  to  the  Alps.  It  is  the  only 
species  of  the  genus  inhabiting  Europe;  and  besides 
the  Alps  it  occurs  in  Southern  Europe  generally, 
and  ranges  as  far  west  as  the  Canaries  and  Madeira. 
Eastward  it  is  not  found  beyond  Central  Asia.  Of 
the  remaining  five  species  of  Petronia,  two  occur  in 
Asia  (including  India)  and  three  in  Africa.  Whether 
the  genus  is  African  or  Asiatic  is  immaterial  for  our 
purpose,  since,  in  any  case,  the  only  European  species 
came  to  us  from  the  east  with  the  Oriental  migration. 
The  distribution  of  the  Alpine  Snow  Finch  (Monti- 
fringilla  nivalis]  is  very  similar  to  that  of  the  birds  we 
have  just  been  considering.  It  inhabits  the  Alps  up 
to  a  great  height,  but  occurs  also  on  the  Pyrenees  and 
other  South  European  mountain  ranges  as  far  east  as 
Palestine,  where  again  it  is  found  in  the  Lebanon. 
The  genus  Montifringilla  has  seventeen  other  species. 
Twelve  of  these  live  in  Central  Asia  and  Japan, 
extending  as  far  north  as  Kamtchatka,  while  five 
inhabit  Western  North  America  right  down  to 
Mexico.  There  is  every  probability  that  in  this  case 


THE  ALPINE   FAUNA.  319 

also  we  have  to  deal  with  an  Asiatic  genus  which 
spread  eastward  to  America,  and  westward  to  Europe. 
As  regards  the  Reptiles,  there  are  no  peculiar 
Alpine  forms,  but  among  the  Amphibia  some  species 
deserve  to  be  mentioned.  Up  to  an  elevation  of 
10,000  feet  we  find  in  the  Alps  the  Black  Salamander 
(Salamandra  atra}\  and  it  is  apparently  quite  peculiar 
to  them,  never  having  been  observed  in  the  plains. 
The  handsome  black  and  yellow  Salamander  (Sala- 
mandra  mac^llosa} — so  well  known  as  a  terrarium 
specimen — likewise  occurs  in  the  Alps,  and  it  has 
besides  a  fairly  wide  distribution  in  Europe.  It  is 
known  from  Southern  Germany,  the  Pyrenees,  Spain, 
Portugal,  Sardinia,  Corsica,  Greece,  Syria,  and  Algiers. 
A  third  species  (S.  caucasicd)  inhabits  the  Caucasus. 
The  evidence  of  distribution  here  points  emphatically 
to  an  Alpine  origin  of  the  genus  Salamandra.  We 
cannot  tell  where  the  ancestors  of  Salamandra  may 
have  come  from,  but  several  other  genera  of  Sala- 
mandridcz  are  certainly  Asiatic.  Our  common  Newt 
(Molge  vnlgaris]  belongs  to  a  genus  with  nineteen 
species,  several  of  which  are  peculiar  to  Europe.  The 
general  range  of  the  genus,  however,  extends  to  North 
America,  and  it  is  more  probable  therefore  that  it 
originated  in  Asia.  If  so,  it  certainly  must  have 
passed  into  Europe  at  a  very  early  date.  Let  us 
assume  the  first  Molges  to  have  traversed  the  ^Egean 
Sea  on  terra  firma  to  Greece  in  miocene  times,  they 
might  thus  have  been  able  to  travel  straight  on  to 
the  old  Tyrrhenian  continent  of  which  Corsica  and 


320         HISTORY   OF   THE   EUROPEAN    FAUNA. 

Sardinia  now  form  the  remains,  and  also  on  to  North- 
west Africa.  Indeed,  we  find  high  up  in  the  Corsican 
mountains  an  interesting  large  brownish-grey  Newt 
{Molge  montana),  and  another  in  Sardinia  (Molge 
Rusconii).  Again,  in  Algiers  there  are  two  species, 
viz.,  Molge  Poireti  and  M.  Hagenmillleri,  while  the 
Moroccan  M.  Waltlii  passes  into  the  south  of  Spain. 
Here  Molge  boscce,  M.  aspera,  and  M.  mannorata 
originated,  the  latter  passing  into  France. 

Another  branch  of  the  Molge  tribe  turned  north- 
ward from  Greece  towards  the  newly  forming  Alps; 
and  there  originated  Molge  alpestris  and  M.  palmata, 
which  more  recently  have  spread  into  England  (one  at 
least),  Germany,  France,  Austria,  and  Southern  Italy. 
Molge  vulgaris  is  an  Asiatic  species  which  wandered 
northward  after  entering  Europe,  covering  a  large 
area,  but  never  reached  the  extreme  south  or  south- 
west. M.  cristata — the  large  Water  Newt — has  a 
similar  but  not  quite  so  extended  a  range,  while 
M.  vittata  never  managed  to  cross  the  borders  of 
Asia  Minor.  Some  of  the  other  species  occur  in 
China,  Japan,  and  North  America. 

None  of  the  tailless  Batrachians — the  Frogs  and 
Toads — are  peculiar  to  the  Alps,  but  one,  viz.  Rana 
temper aria>  ascends  to  the  height  of  no  less  than 
10,000  feet.  It  is  our  common  British  Frog.  No 
other  Frog  probably  ranges  so  far  north  or  to  such 
heights. 

Let  us  now  inquire  what  the  invertebrate  fauna  of 
the  Alps  teaches  us.  We  are  told  by  Dr.  Kobelt, 


THE  ALPINE  FAUNA.  32! 

the  great  authority  on  European  land  shells,  that  a 
uniformity  of  character  marks  the  Alpine  Molluscan 
fauna  (bt  i.,  p.  251).  One  of  the  characteristic  genera 
Campylaea — often  looked  upon  as  a  sub-genus  of 
Helix — is  a  group  containing  somewhat  flattened 
conspicuous  snails  of  large  size.  These  are  found 
everywhere  in  the  Alps,  and  wherever  they  occur 
beyond  the  confines  of  these  mountains,  remarks  Dr. 
Kobelt,  their  origin  from  the  main  stock  is  easily 
traced.  They  have  been  gathered  in  the  Apen- 
nines in  Sicily,  and  even  beyond  the  Mediterranean  in 
Algeria.  On  the  Balkan  peninsula  they  occur  right 
down  to  the  most  southern  point  of  Greece,  but 
are  not  met  with  either  in  Crete  or  Asia  Minor. 
One  species  has  been  found  sub-fossil  in  Thuringia 
in  Northern  Germany. 

Another  truly  Alpine  genus,  says  Dr.  Kobelt,  is 
the  operculate  Pomatias,  which  in  its  geographical 
distribution  offers  some  interesting  modifications  from 
that  of  Campylaea.  Less  limited  to  high  elevations, 
it  has  spread  over  a  greater  part  of  the  plains.  This 
has  happened  especially  in  France,  while  in  Germany 
one  species  advances  almost  as  far  north  as  Heidel- 
berg. In  other  directions  also  the  genus  has  travelled 
beyond  the  limits  of  range  of  Campylaea.  Pomatias 
occurs  in  the  Pyrenees  and  Northern  Spain,  in 
Sardinia  and  Crete,  and  may,  according  to  the  same 
author,  be  expected  in  Asia  Minor,  although  no 
species  has  as  yet  been  met  with  there.  In  Greece, 

again,  it  has  been  observed,  and  numerous  species 

21 


322          HISTORY   OF   THE   EUROPEAN    FAUNA. 

inhabit  Tunis  and  Algeria.  Dr.  Kobelt  connects  the 
wider  range  of  Pomatias  with  the  geological  history 
of  the  genus  (b,  i.,  p.  253).  He  tells  us  that  species  of 
Pomatias  have  been  found  in  eocene  deposits  differ- 
ing but  little  from  our  present  forms,  while  undoubted 
Campylaece  are  not  met  with  till  we  reach  the  upper 
Miocene. 

Zonites  is,  according  to  Dr.  Kobelt,  a  third  Alpine 
genus,  whose  range  scarcely  differs  from  the  other 
two  (b,  i.,  p.  254).  The  centre  of  distribution  lies  at 
present  in  one  of  the  branches  of  the  most  southern 
Alpine  chain  which  help  to  form  a  large  portion  of 
the  Balkan  peninsula.  The  bulk  of  the  species 
inhabit  that  peninsula,  the  Greek  Islands  (except 
Crete)  and  Asia  Minor.  Neither  in  the  Tyrol  nor 
in  Switzerland  do  we  find  any  Zonites^  and  the  few 
species  that  do  occur  in  the  south-eastern  Alps 
only  just  cross  the  outliers  of  these  mountains. 
Between  the  south-western  Alps  and  the  Rhone  we 
again  find  a  Zonites — a  remarkable  case  of  discon- 
tinuous distribution,  since  the  nearest  other  habitat 
of  the  genus  is  Monte  Gargano  in  South-eastern 
Italy,  which  is  known  to  harbour  a  good  many 
interesting  geographical  puzzles. 

We  still  have  a  good  deal  to  learn  as  regards  the 
molluscan  fauna  of  Sicily,  Sardinia,  and  Corsica. 
These  islands  have  scarcely  been  more  than  skimmed 
by  conchologists,  and  Zonites  may  inhabit  one  or  all 
of  these,  which  might  indicate  to  us  the  manner  in 
which  this  genus  travelled  from  Southern  Italy  to 


THE   ALPINE   FAUNA.  323 

Provence  in  the  south  of  France.  The  distribution 
of  Zonites  certainly  dees  not  seem  to  imply  an 
Alpine  origin,  because  it  is  almost  completely  absent 
from  the  Alps  proper.  But  I  do  not  think  my  views 
differ  materially  from  those  of  Dr.  Kobelt,  since  the 
Alps,  in  the  wide  sense,  include  the  mountains  of  the 
Balkan  peninsula,  where  I  should  feel  inclined  to 
locate  the  ancestral  home  of  the  genus. 

The  small  operculate  genus  Acme  is  a  similar 
case.  Dr.  Kobelt  places  the  centre  of  distribution 
on  the  southern  slope  of  the  Alps,  but  scarcely 
any  of  the  species  inhabit  the  Alps  proper.  Some 
occur  in  France,  others  in  North  Africa,  Sicily, 
Southern  Italy,  and  the  Caucasus.  It  is  evidently 
a  very  ancient  genus.  The  species  live  in  moss  or 
underground,  and  are  not  likely  to  be  transported 
across  the  sea  by  accidental  or  occasional  means  of 
distribution. 

Still  another  genus,  which  resembles  Acme  in  its 
geographical  distribution,  is  Daudebardia — a  small 
slug-like  mollusc  with  a  tiny  shell.  It  does  not, 
however,  range  nearly  so  far  north  or  west  as  Acme> 
for  it  occurs  neither  in  the  British  Islands  nor  in 
Spain  or  the  Pyrenees. 

I  shall  not  be  able  to  refer  to  more  than  a  few  of 
the  most  typical  Alpine  species  of  Lepidoptera,  but 
they  may  be  taken  as  fair  examples  of  the  geo- 
graphical distribution  of  the  rest  of  the  group. 

Even  those  visitors  to  Switzerland  who  do  not 
claim  to  be  naturalists  have  heard  of  the  remarkably 


324         HISTORY  OF   THE   EUROPEAN    FAUNA. 

handsome  and  stately  Butterfly  known  as  Apollo. 
To  the  ardent  entomologist,  the  first  sight  of  this 
typical  Alpine  species  is  a  never-to-be-forgotten  de- 
light, and  he  generally  brings  home  with  him  a  rich 
harvest  of  specimens.  The  more  experienced  Butter- 
fly hunter  knows  that  there  are  no  less  than  three 
different  kinds  of  Apollo — or,  as  we  should  say  more 
correctly,  of  Parnassius — in  Switzerland.  There  is 
first  the  common  Apollo  (Parnassius  Apollo],  then 
the  rarer  and  more  local  P  delius>  which  inhabits 
more  elevated  regions,  and  finally  the  still  scarcer  P. 
mnemosyne>  which  is  only  known  from  the  highest 
mountain  ranges.  It  may  be  a  surprise  to  those  who 
have  accustomed  themselves  to  connect  Apollo  with 
the  Alps,  and  who  think  the  two  belong  together  and 
cannot  do  without  one  another,  to  hear  that  it  is 
by  no  means  confined  to  them.  It  is  also  found  in 
Scandinavia,  France,  Spain,  Russia,  and  in  Siberia. 
Parnassius  delius  is  confined  to  the  European  Alps 
and  the  mountains  of  Central  Asia,  while  P.  mnemo- 
syne  is  known  from  the  Pyrenees,  Sweden,  Hungary, 
Sicily,  Russia,  and  Western  Asia.  One  other  Par- 
nassius inhabits  Europe,  viz.,  P.  Nordmanni  of  the 
Caucasus,  but  all  the  remaining  species  of  the  genus 
— and  there  arc  nearly  thirty  more — are  confined  to 
Central  Asia.  A  few,  as  we  have  seen,  have  reached 
Europe,  some  have  travelled  to  the  Himalayan 
Mountains,  and  others  to  Western  North  America. 
The  centre  of  distribution  is  certainly  in  Central 
Asia,  and  we  have  no  reason  to  suppose  that  the 


THE  ALPINE  FAUNA.  325 

original  home  in  this  case  does  not  agree  with  that 
centre. 

Melitcea,  a  genus  to  which  some  of  our  British 
Fritillaries  belong,  has  also  some  typically  Alpine 
members.  Two  of  these,  viz.  M.  cynthia  and  M. 
asteria,  are  peculiar  to  the  Alps,  the  latter  being 
only  found  at  considerable  elevations.  Most  of  the 
remaining  fourteen  European  species  are  also  found 
in  Central  Asia.  Thus  the  isolated  M.  mattirna, 
which  in  Europe  is  confined  to  Lapland,  is  also 
known  from  the  Altai'  Mountains,  which  again  are 
near  the  centre  of  distribution,  since  some  species 
of  Melitcea  range  across  the  Northern  Pacific  to 
Western  North  America. 

The  small  British  Mountain  Ringlet,  and  also  the 
Scotch  Argus,  belong  to  a  genus  of  butterflies  which 
is  very  characteristic  of  the  European  Alps.  But 
owing  to  its  enormous  geographical  distribution,  its 
probable  home  is  somewhat  difficult  to  ascertain. 
Nevertheless  it  is  a  noteworthy  genus,  especially  so 
from  the  fact  that  the  two  British  species  Erebia 
epiphron  and  E.  czthiops  are  taken  at  first  sight 
for  true  Arctic  migrants.  As  neither  of  them,  how- 
ever, occurs  in  Scandinavia,  Greenland,  or  Arctic 
America,  this  supposition  must  be  abandoned.  They 
must  be  looked  upon  as  species  which  once  had  a 
wider  range  in  the  southern  parts  of  the  British 
Islands,  and  which  have  survived  in  a  few  isolated 
localities,  where  they  are  apparently  on  the  verge  of 
extinction. 


326         HISTORY  OF  THE  EUROPEAN   FAUNA. 

About  sixty  species  of  Erebia  are  known  to 
science,  half  of  which  are  found  in  Europe,  the 
remainder  in  Siberia,  the  Himalayas,  Arctic  America, 
Chili,  Patagonia,  South  Africa,  and  Madagascar. 
Though  a  few  do  range  into  these  outlying  regions 
of  the  earth,  Central  Asia  seems  to  lie  near  the  centre 
of  distribution  of  the  genus,  and  the  probability  is 
that  it  also  was  its  original  home.  Most  of  the 
European  species  are  high  Alpine  forms — E.  glacialis 
being  met  with  at  a  height  of  10,000  feet — and 
these  are  generally  quite  peculiar  to  the  Alps, 
showing  that  their  ancestors  came  from  Asia  at  an 
early  date,  probably  by  way  of  Asia  Minor  and 
Greece.  A  few,  as  for  instance  E.  lappona^  range 
right  across  to  the  Altai'  Mountains  from  the  Alps, 
and  at  least  one — E.  melas — is  found  in  Greece. 
Erebia  migrations  seem  therefore  to  have  taken 
place  by  the  Southern  or  Oriental  route  at  different 
geological  periods.  But  some  of  the  European 
species  which  are  more  or  less  confined  to  the  plain, 
and  are  either  absent  from  Switzerland  or  do  not 
reach  the  higher  elevations,  appear  to  me  to  have 
come  by  the  more  direct  northern  or  Siberian  high- 
way, at  a  still  more  recent  period.  These  are  Erebia 
cethiops,  medusa,  ligea,  and  ambla. 

Only  one  species  of  the  well-known  Polar  genus 
(Eneis,  viz.  (E.  aello,  occurs  in  the  Alps.  It  has 
always  been  taken  at  very  high  elevations  near  the 
verge  of  the  snow-line  on  the  most  lofty  parts 
of  the  Simplon  Pass,  and  other  similar  situations. 


THE  ALPINE  FAUNA.  327 

Altogether  about  a  dozen  species  of  this  genus  of 
butterfly  are  known,  most  of  which  are  confined  to 
the  polar  regions  of  the  Old  World  and  the  New, 
though  some  have  found  their  way  to  the  extreme 
south  end  of  South  America,  in  what  manner  is  still 
a  mystery.  Like  the  preceding  genera,  this  also 
appears  to  have  emerged  from  Central  Asia.  The 
genus,  too,  is  closely  allied  to  the  last,  and  though  its 
range  is  not  quite  so  extensive,  it  resembles  it  in 
many  respects.  The  Alpine  species  of  CEneis  came 
to  Europe  by  the  Oriental  route.  But  the  Lapland 
species — at  any  rate  CE.  jutta  and  (E.  bore — have 
taken  a  somewhat  circuitous  route  to  reach  our 
continent.  They  first  migrated  from  Asia  to  North 
America,  and  then  by  the  old  land-connections  by 
way  of  Greenland  to  Lapland.  It  is  noteworthy  that 
Professor  Engler  felt  convinced  (cf.  p.  171;  that  the 
occurrence  of  many  of  the  Arctic  plants  in  North 
Scandinavia  and  Siberia  could  be  best  explained  by 
the  assumption  of  such  a  migration  from  Asia  via 
North  America  to  Europe  rather  than  by  the  shorter 
route. 

There  are  far  more  Alpine  beetles  than  butterflies, 
but  their  geographical  distribution  is  less  well  known, 
and  it  is  therefore  not  at  all  safe  to  base  important 
conclusions  as  to  the  origin  of  a  fauna  on  that  group 
alone ;  however,  as  far  as  my  limited  knowledge  of 
the  Coleoptera  of  the  Alps  goes,  their  general  range 
seems  to  agree  perfectly  with  other  orders  of  insects. 
Many  can  also  be  traced  to  an  Asiatic  home,  and 


328         HISTORY  OF   THE   EUROPEAN   FAUNA. 

the  route  they  came  by  is  the  Oriental  and  not  what 
I  have  called  the  Siberian. 

Take,  for  instance,  the  genus  Nebria,  of  which  we 
have  one  species  in  England — a  black  insect  with  a 
bright  reddish-yellow  border  and  long  light  legs — 
known  as  N.  livida.  There  are  about  eighty  Euro- 
pean species,  most  of  which  are  confined  to  the  Alps, 
the  Caucasus,  the  Pyrenees,  Spain,  and  Greece.  The 
genus,  however,  ranges  all  over  the  Holarctic  Region, 
that  is  to  say  roughly,  over  Europe,  Central  and 
Northern  Asia,  and  North  America.  The  centre  of 
distribution  lies  in  Central  Asia.  If  the  genus  had 
poured  into  Europe  by  the  northern  or  Siberian 
route,  we  should  probably  now  find  many  species  in 
Northern  Russia,  Germany,  and  France ;  but  this  is 
not  the  case,  and  we  may  therefore  assume  with  some 
justification  that  the  Southern  or  Oriental  route  was 
the  only  one  available  at  the  time  when  the  bulk  of 
the  species  of  Nebria  wandered  to  Europe.  Many 
of  the  Nebrias  occur  in  Switzerland  and  in  the  Alps, 
generally  on  the  margins  of  the  snow-fields  and 
glaciers,  like  N.  Germari  and  Brunii.  Others,  for 
example,  N.  atrata,  ascend  to  the  highest  limit  of 
animal  life,  having  been  observed  at  a  height  of  over 
10,000  feet. 

Of  the  remaining  orders  of  insects  we  know  as  yet 
very  little.  Central  Asia  and  even  Siberia  are  only 
beginning  to  be  explored,  and  their  invertebrate 
fauna — except  Lepidoptera  and  Coleoptera — is  practi- 
cally unknown.  However,  I  cannot  conclude  this 


THE  ALPINE  FAUNA.  329 

short  summary  of  some  of  the  more  characteristic 
Alpine  animals  without  referring  to  the  Grasshoppers 
which  are  so  conspicuous  in  the  mountains.  The 
mountain  air  simply  rings  during  a  bright  summer's 
day  with  the  loud  and  cheerful  song  of  millions 
of  these  insects.  It  is  one  of  the  most  vivid  impres- 
sions a  tourist  brings  back  from  Switzerland — this 
constant  shrill  sound  issuing  from  an  apparently 
invisible  source. 

Among  these  Grasshoppers  there  are  some  highly 
characteristic  Alpine  genera.  Pezotettix — formerly 
known  as  Podisma — is  one  of  these.  P.  alpinus  is 
almost  confined  to  the  high  Alps ;  with  P.  mendax 
it  occurs  in  lower  levels  chiefly  towards  the  south- 
east, that  is  to  say,  in  the  direction  of  Hungary, 
Servia,  and  Dalmatia.  P.  frigidus  occurs  not  only 
in  the  high  Alps,  but  also  in  Lapland.  P.  ScJimidti 
and  P.  salamandra  are  found  in  Carinthia,  Servia, 
and  Transylvania;  and  one  species  also  inhabits 
the  Pyrenees  and  another  the  Italian  Mountains. 
Finally,  the  only  English  species  of  Pezotettix^ 
viz.  P.  pedestris,  has  been  taken  in  Sweden,  Den- 
mark, and  then  again  in  Austria,  Hungary,  Servia, 
etc.,  as  far  east  as  the  Volga,  and  also  on  the  high 
Alps,  in  Sardinia  and  the  Abruzzi  Mountains  in 
Italy. 

Very  little,  as  I  remarked,  is  known  of  the  Asiatic 

range  of  this  genus,  but  either  the  same  or  a  closely 

-  allied   one  has  many   representatives   in    North   and 

South    America.      Whether    Pezotettix    is    therefore 


330        HISTORY  OF  THE   EUROPEAN   FAUNA. 

of  Asiatic  origin  we  cannot  positively  affirm,  but 
whatever  view  we  take,  the  general  range  of  the 
European  species  indicates  that  the  migration  took 
place  from  the  Alps  in  a  south-easterly  direction,  or 
to  them  in  a  north-westerly  one.  That  is  to  say  the 
Oriental  route,  and  not  the  Siberian,  was  utilised  by 
the  migrants. 

Fortunately,  we  know  a  little  more  about  another 
Grasshopper  genus,  called  Chrysochraon.  There  are 
only  two  species,  one  of  which,  Chr.  dispar,  has  been 
found  from  Northern  France  to  the  mountains  of 
Servia,  but  not  in  the  Alps.  The  other,  Chr. 
brachypterus,  has  a  somewhat  similar  range  in  the 
plain;  but,  moreover,  it  inhabits  the  Alps  up  to  a 
considerable  height.  It  is  interesting  to  note  that 
both  these  Grasshoppers  again  turn  up  on  the  Amur 
in  Eastern  Siberia. 

In  conclusion,  I  might  mention  one  more  Grass- 
hopper, viz.  Tettix,  because  it  includes  a  species-—  T. 
bipunctatus — which,  though  well  known  in  the  plain  of 
Middle  and  North  Europe,  ascends  the  Alps  to  a 
height  of  nearly  10,000  feet.  It  is  one  of  the  few 
instances  I  know  of  an  animal  occurring  in  the 
same  form  in  such  an  enormous  range  of  altitude 
— from  sea-level  to  the  highest  regions  where  animal 
life  is  known  to  exist.  It  is  also  known  from  Asia 
Minor  and  Siberia.  T.  subulatus  has  a  similar  dis- 
tribution, but  is  more  common  in  Southern  Europe 
than  the  other.  T.  fuliginosus  occurs  in  Lapland 
and  Siberia,  T.  meridionalis  and  T.  depressus  all 


THE  ALPINE  FAUNA.  331 

along  the  shores  of  the  Mediterranean.  There  can  be 
no  doubt  that  here  also  we  can  trace  migration  to  or 
from  Siberia,  and  again,  as  on  previous  occasions,  by 
the  Oriental  route. 

We  now  possess  a  fair  general  idea  of  the  fauna  of 
the  Alps.  We  have  learned  that  a  good  many  of 
the  animals  are  indigenous,  and  that  others  have 
migrated  to  the  Alps  by  various  routes.  The  majority 
of  these  have  come  from  Central  and  Southern  Asia 
with  what  has  been  described  as  the  Oriental  migra- 
tion. A  much  smaller  number  have  reached  the 
Alps  from  the  north  and  the  west,  but  none  of  the 
latter  are  among  the  high  Alpine  forms.  What  will 
be  the  most  surprising  revelation  is  that  the  eastern 
species,  which  arrived  in  Europe  with  the  Siberian 
migration,  are  practically  absent  from  the  Alps 
proper.  No  doubt  some  of  them  still  survive  in  the 
lowlands  of  Switzerland  and  the  Tyrol,  but  none  of 
the  true  Alpine  fauna  owes  its  origin  to  the  Siberian 
migration.  If  we  compare  the  Alpine  mammals  with 
the  Siberian  forms  which  reached  England  (vide 
p.  202),  we  at  once  perceive  the  difference.  We 
should  expect  to  find  in  the  Alps — if  not  the  Rein- 
deer and  the  Glutton — the  Arctic  Fox,  the  little 
Pica,  the  Lemmings,  and  the  pouched  Marmots.  It 
might  be  urged  that  some  of  the  smaller  Siberian 
carnivores  and  rodents  do  inhabit  the  Alps.  So  they 
do.  The  Stoat  and  Weasel  have  found  such  a  con- 
genial home  in  Europe,  both  in  the  plain  and 
mountains,  that  they  have  spread  rapidly  to  the 


332        HISTORY  OF  THE   EUROPEAN   FAUNA. 

latter,  and  no  doubt  reached  within  a  comparatively 
short  time  the  great  heights  at  which  they  now 
occur  in  the  Alps.  But  the  Voles  (Arvicola)  have 
scarcely  spread  beyond  the  region  of  fields  and 
cultivated  ground.  A  height  of  5000  feet  at  the 
most  marks  their  maximum  altitude  in  the  Alps. 

The  fauna  which  reached  the  Alps  in  miocene  and 
pliocene  times,  as  well  as  the  indigenous  element, 
must  have  survived  the  Glacial  period  in  their 
mountain  home.  Though  I  think  that  the  con- 
ditions of  the  climate  at  that  time  and  the  size  of 
the  Scandinavian  glaciers  have  been  greatly  ex- 
aggerated, there  can  be  no  doubt  at  all  about  the 
enormous  size  of  many  of  the  Alpine  glaciers  at  this 
period.  The  climate  was  probably  much  moister  but 
not  colder  than  what  it  is  now,  possibly  warmer. 
The  snowfall  was  therefore  greater,  so  that  glaciers 
filled  many  of  the  lower  valleys  of  Switzerland  which 
are  now  quite  free  from  ice,  and  even  invaded  the 
plain.  But  there  is  no  reason  whatsoever  why  the 
Alps  should  not  even  then  have  supported  a  luxuriant 
fauna  and  flora  as  they  do  now.  Possibly  many  of 
the  miocene  plants  and  animals  became  extinct  then, 
but  extinction  of  species  occurs  at  the  present  day. 
We  hear  complaints  that  the  Chamois  and  the 
Steinbock  have  nearly  vanished ;  we  know  that  the 
Marmot  is  now  much  scarcer  than  it  used  to  be, 
and  that  the  Edelweiss  and  many  other  plants  are 
more  and  more  difficult  to  find,  and  seem  rapidly  to 
disappear.  No  doubt  all  this  is  in  a  great  measure 


THE   ALPINE   FAUNA.  333 

due  to  the  influence  of  man,  but  not  altogether. 
There  is  a  constant  struggle  for  existence  going 
on  among  the  animals  and  plants  themselves — the 
stronger  and  fitter  species  driving  the  less  fit  and 
weaker  into  a  corner,  where  they  finally  succumb. 
This  happens  now  just  as  it  did  in  pliocene  and 
pleistocene  times,  and  need  not  imply  change  of 
climate. 

As  soon  as  the  Miocene  sea  to  the  north  of  the 
mountains  had  retreated,  a  portion  of  the  Alpine 
fauna  poured  into  the  plain,  and  many  species 
have  found  their  way  to  the  British  Islands,  a  few 
to  Scandinavia  and  Russia.  Westward  too,  the  sea 
soon  after  retired  and  opened  a  way  for  those 
Alpine  species  which  were  vigorous  enough  to  extend 
their  range  in  that  direction.  South-eastward,  of 
course,  a  highway  had  long  ago  been  open,  and 
Alpine  forms  which  were  able  to  migrate  towards 
the  incoming  Oriental  stream,  had  no  difficulty  in 
doing  so.  When  they  arrived  in  Greece,  some  turned 
westward  again  and  populated  Sicily,  Southern  Italy, 
Sardinia,  Corsica,  and  Northern  Africa,  while  others 
crossed  over  to  Asia  Minor,  which  was  then  con- 
nected with  Greece,  and  wandered  towards  the 
Central  Asiatic  or  the  Himalayan  Mountains. 

But,  as  I  remarked,  few  of  the  typical  Alpine 
species  reached  Scandinavia  and  Lapland.  I  have 
already  referred  to  the  similarity  between  the  North- 
ern Scandinavian  and  the  Alpine  faunas  in  a  pre- 
vious chapter,  and  I  have  shown  that  this  resemblance 


334         HISTORY   OF   THE   EUROPEAN   FAUNA. 

cannot  altogether  be  explained  by  the  supposition  of 
an  interchange  in  the  faunas  of  the  two  regions.  That 
this  has  taken  place  to  some  extent  is  probable,  but 
the  resemblance  appears  more  especially  due  to  the 
fact  that  the  Alps  and  Scandinavia  have  been  peopled 
from  the  same  centres  of  distribution. 

In  order  to  make  this  matter  quite  clear,  I  will  give 
a  familiar  example  as  an  instance  of  the  manner  in 
which  the  present  distribution  can  be  explained  with- 
out taking  recourse  to  direct  migration  from  the  Alps 
to  Scandinavia  or  vice  versa.  The  example  I  will 
take  is  that  of  a  family  of  birds,  not  only  of  extreme 
interest  from  the  fact  of  its  northern  range,  but  also 
from  the  pleasure  it  gives  to  those  addicted  to  sport. 
This  is  the  grouse  family,  the  Tetraonida. 

Let  us  commence  with  our  British  Grouse  (Lagopus 
scoticus\  which  is  peculiar  to  the  British  Islands. 
In  Norway  we  find  a  Grouse  (L.  albus)  which  differs 
in  habit,  and  in  the  fact  of  its  turning  white  in 
winter ;  otherwise  it  is  so  closely  allied  to  our 
Grouse  that  many  ornithologists  do  not  separate 
them  specifically.  No  doubt  the  British  Grouse  is  a 
descendant  of  this  Scandinavian  Willow-grouse.  The 
latter  is  known  also  to  inhabit  Greenland  and  Arctic 
North  America,  and  it  is  even  found  beyond  Behring 
Straits  in  Northern  Siberia.  En  route  between 
Scandinavia  and  Asia,  travelling  in  a  westward 
direction,  we  meet  with  two  other  very  local  species 
of  Grouse,  which  may  be  looked  upon  as  offshoots 
of  L.  rnpestris — viz.,  L.  hyperboreus  of  Spitsbergen, 


THE   ALPINE   FAUNA.  335 

and  leucurus  of  Western  North  America.  In  Asia  we 
then  again  find  two  kinds  of  Grouse,  very  closely 
related,  and  by  some  indeed  regarded  as  belonging 
to  the  same  species.  These  are  L.  rupestris  and 
L.  mutus.  Mr.  Ogilvie-Grant  tells  us  of  the  former 
(p.  49),  that  it  is  merely  a  more  northern  rufous  form 
of  L.  mutus,  and  .that  it  goes  through  similar  changes 
of  plumage.  In  summer  the  males  are  readily  dis- 
tinguishable, but  in  winter  it  is  impossible  to  tell 
one  from  the  other.  "  L.  rupestris  taken  as  a  whole," 
says  Mr.  Ogilvie-Grant,  "appears  to  us  barely  specifi- 
cally distinct  from  L.  mutus"  L.  rupestris  occurs 
not  only  in  Northern  Asia,  but  crosses  the  Behring 
Straits  to  Arctic  America,  being  still  found  on  the 
Aleutian  Islands,  which  represent  the  last  remains 
of  the  former  land-bridge  between  Asia  and  North 
America,  then  eastward  to  Greenland  and  Iceland. 
However,  while  this  form  does  not  cross  the  confines 
of  Asia  in  a  westerly  direction,  its  near  relative 
L.  mutus — better  known  as  the  Ptarmigan — does; 
and  may  perhaps  have  entered  Europe  as  a  Siberian 
and  also  as  an  Arctic  migrant  It  is  still  found  in 
the  Ural  Mountains,  in  Finland,  and  the  highlands 
of  Scandinavia.  It  is  gradually  being  driven  out  of 
the  Alpine  lowlands,  while  it  has  long  ago  dis- 
appeared from  Germany,  France,  and  Austria — in 
fact,  from  all  the  lowlands  of  Europe.  It  has  also 
been  met  with  in  the  Pyrenees  and  in  some  of  the 
Spanish  mountains.  Similarly,  the  bird  has  become 
extinct  in  England  and  Ireland,  while  it  is  becoming 


336        HISTORY   OF   THE   EUROPEAN    FAUNA. 

more  and  more  scarce  in  Scotland.  The  centre  of 
distribution  of  the  genus  lies  in  Arctic  America,  and 
from  there  the  genus  has  spread  to  Europe  and 
Asia.  L.  albus  and  L.  mutus  appear  in  our  continent 
chiefly  as  Arctic  migrants. 

The  Black  Grouse  (Lyrurus  tetrix)  belongs  to  a 
closely  allied  genus,  which  has  only  two  species. 
One  of  these  is  very  local  in  distribution,  being  con- 
fined to  the  Caucasus,  but  the  smallness  of  range  is 
to  some  extent  compensated  for  by  the  peculiarity 
of  its  name,  which  is  L.  mlokosiewiczi.  The  Black 
Grouse,  on  the  contrary,  is  widely  distributed.  It 
inhabits  Northern  Asia  from  the  Pacific  to  the  Ural 
Mountains,  and  extends  as  far  south  as  Pekin  and 
the  Tian  Shan  range.  In  Europe  it  is  found  from 
the  extreme  east  to  the  Pyrenees,  the  Apennines  on 
the  south,  and  to  Great  Britain  and  Scandinavia 
in  the  north.  It  is  important  to  note  its  absence 
from  Spain,  the  Mediterranean  islands,  and  Ireland ; 
and  we  have  learned  that  it  is  one  of  those  Siberian 
migrants  which  have  succeeded  in  establishing  them- 
selves in  the  Alps. 

The  Capercaillie  (Telrao  urogaUus) — another  great 
favourite  with  sportsmen — is  now  generally  separated 
generically  from  the  Black  Grouse,  though  they  are 
of  course  near  relations.  Its  range  greatly  resembles 
that  of  the  Black  Grouse,  except  that  it  does  not  go 
quite  as  far  east  in  Siberia,  not  having  been  met 
with  beyond  Lake  Baikal.  From  there  it  is  found 
westward  as  far  as  the  Pyrenees.  It  occurs  also  in 


THE   ALPINE   FAUNA.  337 

the  Carpathians  and  the  Alps.  In  England,  where  it 
used  to  be  known  by  the  name  Cock  of  the  Wood,  it 
became  extinct  at  some  remote  period  in  history, 
while  it  lingered  on  in  Scotland  and  Ireland  until 
the  end  of  the  last  century.  In  Scotland  it  has  been 
reintroduced  into  several  counties,  and  being  pro- 
tected, it  appears  to  spread  from  these  artificial 
centres  of  distribution. 

Like  the  Black  Cock,  the  Capercaillie  is  a  Siberian 
migrant,  and  it  is  one  of  the  few  Siberian  species 
which  have  reached  Ireland,  as  I  have  had  occasion 
to  mention  in  dealing  with  the  origin  of  the  British 
fauna.  Two  other  species  of  Capercaillie  and  an 
allied  genus  (Falcipennis]  are  met  with  in  the  extreme 
north-east  of  Siberia,  and  six  other  genera,  all  be- 
longing to  the  grouse  family,  are  confined  to 
North  America.  We  have  therefore  a  very  intimate 
relationship  between  the  grouse  of  Asia  and  those  of 
North  America,  some  species  even  ranging  right 
across  the  two  continents. 

The  last  genus  of  this  very  interesting  family  is 
Tetrastes.  This  grouse  is  not  familiar  to  British 
ornithologists,  since  it  is  entirely  absent  from  the 
British  Islands.  But  sportsmen  who  have  tramped 
over  Scandinavia  know  it  well  by  the  name  of 
Hazel  Grouse.  It  is  ashy  grey  in  colour,  barred  and 
vermiculated  with  black.  The  Common  Hazel 
Grouse  (Tetrastes  bonasia)  is  found  from  Northern 
Spain  in  the  west  right  through  the  mountainous 

parts  of  Central  and  Northern  Europe  and  Northern 

22 


338         HISTORY  OF  THE   EUROPEAN    FAUNA. 

Asia  to  Kamtchatka  and  the  Russian  convict  island 
of  Saghalien  in  the  Pacific.  Besides  the  Common 
Hazel  Grouse,  two  other  species  are  known,  one  from 
Eastern  Russia  and  the  other  from  China. 

Having  now  shortly  reviewed  the  whole  grouse 
family,  we  have  seen  that,  although  some  species 
live  within  the  Polar  Circle,  the  majority  are 
more  or  less  confined  to  the  more  temperate 
or  rather  the  less  arctic  parts  of  the  Northern 
Hemisphere.  They  are  quite  absent  from  Southern 
Asia  and  even  the  southern  parts  of  North  America, 
and  almost  so  from  the  Mediterranean  basin.  The 
whole  range  of  the  family  is  therefore  suggestive  of 
a  northern  origin,  and  this  view  agrees  perfectly  with 
all  the  details  of  distribution.  The  centre  of  dis- 
tribution lies  in  Northern  Asia,  or  in  Arctic  North 
America.  From  there  the  great  genus  Lagopus 
spread  east  and  west,  reaching  Europe  by  these 
vastly  divergent  routes  at  a  time  when  the  physical 
geography  was  very  different  from  what  it  is  to- 
day. Several  of  the  species  common  to  the  Alps 
and  Scandinavia  have  migrated  from  Siberia  direct 
to  Eastern  Europe.  But  we  can  now  imagine  how 
from  a  similar  centre  in  Asia — perhaps  at  a  rather 
more  remote  time — a  species  spread  eastward  across 
North  America  and  Greenland  to  Scandinavia,  and 
westward  along  the  mountain  ranges  of  the  Tian 
Shan  and  the  mountains  of  Asia  Minor  to  Greece, 
and  finally  to  the  Alps.  We  should  then  have  the 
same  species  in  the  Alps  and  in  Scandinavia,  not  far 


THE  ALPINE   FAUNA.  339 

removed  from  one  another;  but  how  different  were 
their  paths  of  migration !  This,  however,  is  not  an 
imaginary  instance.  Such  a  migration  must  have 
actually  taken  place  in  a  good  number  of  instances 
among  the  terrestrial  invertebrates  and  also  among 
plants. 

The  view  still  current  among  many  zoologists  and 
botanists,  that  animals  and  plants  were  driven  down 
into  the  plain  from  the  mountains  of  Europe  during 
the  height  of  the  Glacial  period  and  there  lived 
together  till  the  return  of  a  more  genial  temperature, 
when  they  retreated  to  their  mountain  homes,  is  a 
very  plausible  one.  During  their  sojourn  in  the 
plain,  the  plants  and  animals — say  from  Scandi- 
navia— intermingled  with  those  from  the  Alps  ;  and 
when  the  time  of  separation  came,  many  Alpine 
forms  retired  northward  with  the  Scandinavians, 
while  many  Scandinavians  would  go  with  the 
Alpines  to  their  home.  In  this  way  the  similarity 
between  the  Alpine  and  Scandinavian  faunas  and 
floras  is  assumed  to  have  been  brought  about. 
These  theories,  first  promulgated  by  Edward  Forbes, 
were  hailed  with  general  satisfaction  by  the  scientific 
world.  Even  Darwin  says  of  them  (p.  331),  that 
grounded  as  they  are  on  the  perfectly  well- 
ascertained  occurrence  of  a  former  Glacial  period, 
they  seemed  to  him  to  explain  in  a  satisfactory 
manner  the  present  distribution  of  the  Alpine  and 
Arctic  productions  of  Europe.  To  the  present  day 
this  view  meets  with  much  favour  among-  naturalists. 


340        HISTORY  OF   THE   EUROPEAN    FAUNA. 

It  is  somewhat  similar  to  one  which  has  recently 
been  strongly  supported  by  Professor  Nehring  and 
accepted  by  Professor  Th.  Studer  and  many  others. 
They  have  never  made  it  quite  clear  whether  the 
pre-glacial  fauna  and  flora  are  supposed  to  have 
been  absolutely  destroyed  by  the  glacial  climate, 
or  whether  part  of  them  have  been  able  to  take 
refuge  somewhere  in  the  south;  but  the  great  mass 
of  our  Alpine  plants  and  animals  are  believed  to 
have  been  derived  from  the  Siberian  invasion,  which 
I  have  fully  described  in  the  fifth  chapter.  This 
invasion  spread  over  the  European  plain,  and  when 
the  climate  ameliorated,  both  animals  and  plants 
migrated  north  and  south  to  the  mountains.  This 
view  agrees  with  the  earlier  theory,  except  that  the 
adaptation  to  Alpine  conditions  would,  according  to 
the  former,  have  taken  place  since  the  close  of  the 
Glacial  period,  during  which  time  no  such  modifica- 
tion or  change  of  species  seems  to  have  been  pro- 
duced in  other  parts  of  the  world.  The  characteristic 
fauna  of  the  Alps,  as  has  been  gathered  from 
the  preceding  pages,  is  mainly  of  Central  Asiatic 
rather  than  of  Siberian  origin.  Migration  to  the 
Alps  took  place  by  the  Oriental  route  long  before 
the  Siberian  invasion.  Some  of  the  species  of  the 
latter  have  penetrated  to  the  Alps,  but  these  Siberian 
species  have  not  given  to  the  fauna  of  the  highest 
European  mountain  range  the  striking  character  with 
which  we  all  associate  it. 

Before  concluding  this  chapter,  a  few  remarks  on 


THE   ALPINE   FAUNA.  341 

the  botanical  aspect  of  the  Alpine  problem  might  not 
be  out  of  place.  It  will  enable  us  to  judge  which  of 
the  views  indicated  is  the  more  probable,  and  will 
add  to  the  interest  which  may  have  been  aroused  by 
the  perusal  of  this  sketch  of  the  fauna  of  the  Alps. 
Very  much  the  same  train  of  argument  was  applied 
as  to  the  course  of  events  in  the  formation  of  the 
Alpine  flora  as  in  the  case  of  the  fauna.  The  plants 
were  all  supposed  to  have  been  killed  or  driven  away 
by  the  arctic  temperature  of  the  Glacial  period,  and 
their  place  taken  by  new  migrants  from  the  north  or 
east  when  the  climate  ameliorated. 

Professor  Engler,  one  of  the  highest  living 
authorities  on  the  geographical  distribution  of  plants, 
is  of  opinion  (p.  102)  that  a  large  number  of  the 
indigenous  Alpine  species  did  not  originate  till  after 
the  close  of  the  Glacial  period,  because  so  many  of 
them  are  absent  from  the  Sierra  Nevada  in  Spain, 
where  the  condition  for  their  well-being  exists, 
while  many  have  evidently  spread  from  the 
Alps  to  the  Carpathian  Mountains  and  to  the 
Pyrenees.  He  does  not  believe  that  a  glacial  flora 
could 'have  existed  in  the  plain  between  the  Sierra 
Nevada  and  the  Pyrenees  during  the  Glacial  period 
(p.  109).  In  speaking  of  the  Caucasus,  Professor 
Engler  informs  us  (p.  117)  that  a  good  many  species 
which  do  not  occur  in  the  Alps  reached  these 
mountains  from  Siberia.  Apart  from  the  northern 
glacial  plants,  the  Caucasus  has  only  few  species  in 
common  with  the  Alps,  more  with  the  Balkan  moun- 


342        HISTORY  OF    THE   EUROPEAN   FAUNA. 

tains  and  Northern  Persia.  Turning  to  Afghanistan, 
our  author  mentions' (p.  121)  a  few  Alpine  plants  as 
occurring  in  that  country,  and  likewise  in  the 
Caucasus  and  the  Himalayas.  He  considers  it 
probable  that  the  route  of  migration  of  some  glacial 
plants  from  the  east  to  the  west,  and  vice  versa,  lay 
across  the  Afghan  mountains.  Many  of  our  Alpine 
plants  occur  in  the  Siberian  mountains,  but  in  the  Altai' 
and  Eastern  Siberia  generally  a  considerable  number 
of  these  are  by  no  means  confined  to  the  mountains 
(p.  125).  They  are  also  met  with  in  the  lower  regions, 
and  the  rare  Alpine  Edelweiss  (Leontopodium  alpinuvi} 
frequently  covers  wide  tracts  in  the  plain,  and  is 
passed  by  almost  unnoticed  by  the  Siberian  botanist. 
Special  attention  is  drawn  by  Professor  Engler  to 
the  fact  (p.  130)  that  several  of  the  Siberian  plants 
inhabit  the  Alps  and  the  Caucasus,  but  are  not 
found  in  Scandinavia.  And  from  this  he  deduces  the 
conclusion  that  part  of  the  Siberian  flora  migrated 
in  a  south-westerly  direction  towards  the  Caucasus 
and  the  mountains  of  the  Mediterranean  area,  exactly 
in  the  manner  indicated  in  respect  to  the  fauna  of  the 
Alps.  We  learned  that  the  migration  to  the  Alps 
from  Central  and  perhaps  also  parts  of  Northern 
Asia  took  a  south-westerly  course  first,  and  was 
then  followed  by  one  in  an  easterly  direction.  I 
called  the  former  the  Oriental  migration  and  the 
latter  the  Siberian.  Later  on  Professor  Engler 
states  (p.  142)  that  the  main  mass  of  the  Siberian 
forms  of  plants  certainly  wandered  westward  to 


THE   ALPINE   FAUNA.  343 

the  south  of  the  Ural.  This  is  proved  by  the 
numerous  glacial  plants  found  in  the  Caucasus,  while 
the  glacial  flora  of  the  Ural  Mountains  is  poor. 
Finally,  he  expresses  the  opinion  that  the  probability 
of  most  of  the  Alpine  plants  occurring  in  Arctic 
Siberia,  having  wandered  from  the  Alps,  by  way 
of  Scandinavia,  Greenland,  and  North  America,  to 
North-eastern  Siberia,  is  greater  than  the  direct 
migration  from  Europe  to  Siberia  (p.  143). 

Another  continental  writer  on  the  Alpine  flora  who 
deserves  special  mention  is  Dr.  Christ.  His  observa- 
tion that  Alpine  plants  by  no  means  suffer  from  a 
high  temperature  (p.  309),  but  solely  from  a  drying 
up  of  the  soil,  seems  to  me  to  point  to  the  correctness 
of  the  view  I  have  expressed  on  several  occasions, 
that  these  plants  have  originated  long  before  the 
Glacial  period  at  a  time  when  the  climate  was 
warmer  and  moister  than  it  is  now.  It  seems  quite 
natural  to  Dr.  Christ  that  the  Arcto-Alpine  flora 
should  have  originated  in  Asia,  but  he  excepts  thirty 
species  which  are  absent  from  Northern  Asia,  though 
occurring  in  America  (p.  327).  These  he  thinks  have 
penetrated  direct  from  America  to  the  Alps  by  way 
of  Scandinavia,  since  no  less  than  twenty-three  still 
occur  in  the  latter  country.  In  the  human  population 
of  the  Alps,  he  continues  (p.  336),  one  can  distinguish 
an  indigenous  Celtic  race,  a  Germanic  colder  and 
more  apathetic  race,  and  a  more  lively  Roman  one. 
The  flora  is  composed  of  quite  a  similar  mixture. 
We  find  also  an  indigenous  element — an  Arctic  and 


344        HISTORY  OF  THE  EUROPEAN   FAUNA. 

a  Mediterranean  one.  The  last  element  is  a  survival 
of  the  Tertiary  flora  of  the  Central  European  plateau 
(p.  532)-  The  plants  were  driven  down  to  the  shores 
of  the  Mediterranean,  and  it  is  only  after  the  retreat 
of  the  glaciers  that  a  few  of  them  have  been  able  to 
regain  their  ancient  territory.  The  incoming  Asiatic 
and  North  American  flora  likewise  retired  at  the  end 
of  the  Glacial  period  to  the  Alps  and  the  Arctic 
countries,  and  left  isolated  traces  of  its  former 
abundance  on  the  North  European  plain.  The 
bulk  of  the  Arctic  or  Alpine  flora  is  held  to  be 
of  Asiatic  origin.  Since  Siberia  shows  little  trace 
of  having  been  glaciated,  owing  to  the  dryness  of  the 
climate,  a  rich  flora  was  able  to  develop  there,  which 
spread  into  Europe  as  soon  as  the  vanishing  glaciers 
made  room  for  it. 

These  are  the  views  of  Professor  Engler  and  Dr. 
Christ.  They  agree  in  so  far  as  both  of  them  main- 
tain that  the  bulk  of  the  Alpine  flora  is  post-glacial 
— that  is  to  say,  that  it  has  developed  quite 
recently,  or  migrated  to  the  Alps  after  the  glaciers 
had  retreated  from  the  plain  to  the  mountain 
recesses.  It  is  assumed  by  Dr.  Christ  that  while 
Europe  was  practically  uninhabitable,  a  rich  flora 
survived  in  Northern  Asia,  because  the  climate  there 
was  too  dry  for  the  development  of  glaciers.  Due 
consideration  in  this  interesting  speculation,  however, 
is  not  given  to  the  fact  which  he  himself  emphasised, 
that  Alpine  plants  are  particularly  prone  to  suffer 
from  a  dry  climate.  Even  a  moderately  dry  cold 


THE    ALPINE   FAUNA.  345 

kills  most  of  them.  How  can  we  then  reconcile  this 
fact  with  the  theory  of  their  origin  in  a  dry  and 
intensely  cold  climate?  I  quite  agree  with  the  views 
as  to  the  Asiatic  origin  of  the  bulk  of  the  Alpine  flora, 
while  the  dry  state  of  the  Siberian  climate  is  certainly 
indicated  by  the  extremely  feeble  development  of  the 
glaciers  during  a  .large  part  of  the  Glacial  period. 
We  know,  however,  that  in  Pliocene  and  even  in 
early  Glacial  times  the  atmospheric  conditions  must 
have  been  very  different  in  Siberia.  A  great  slice  of 
Central  Asia  was  under  water,  and  numerous  fresh- 
water lakes  covered  the  lowlands  in  the  north,  so 
that  the  climate  must  have  been  damp  though  not 
cold  enough  for  the  formation  of  extensive  glaciers. 
Everything,  in  fact,  seems  to  indicate  that  the  migra- 
tion of  the  Asiatic  Alpine  flora  took  place  at  a  very 
early  date — probably  long  before  the  Glacial  period — 
either  by  the  Oriental  or  by  the  Arctic  route  via 
North  America,  Greenland,  and  Scandinavia.  But 
would  this  not  necessitate  a  survival  of  the  Alpine 
plants  in  the  Alps  themselves?  That  is  the  view  which 
has  already  been  expressed  with  regard  to  the  fauna, 
and  the  flora  probably  followed  a  very  similar  course. 
This  is  by  no  means  a  novel  theory,  however,  and 
though  unfortunately  an  untimely  death  has  removed 
one  of  our  very  best  authorities  on  the  Alpine  flora 
before  he  had  completed  his  life's  work,  we  have 
some  indications  in  the  earlier  writings  of  John  Ball 
that  his  opinions  on  the  origin  of  that  flora  did  not 
coincide  with  those  held  by  the  leading  continental 


346        HISTORY   OF   THE   EUROPEAN    FAUNA. 

authors.  To  quote  the  words  of  this  distinguished 
botanist  (p.  576):  "Is  it  credible  that  in  the  short 
interval  since  the  close  of  the  Glacial  period  hundreds 
of  very  distinct  species  and  several  genera  have  been 
developed  in  the  Alps,  and — what  is  no  less  hard  to 
conceive — that  several  of  these  non-Arctic  species 
and  genera  should  still  more  recently  have  been 
distributed  at  wide  intervals  throughout  a  discon- 
tinuous chain  some  1500  miles  in  length,  from  the 
Pyrenees  to  the  Eastern  Carpathians?  Nor  would 
the  difficulties  cease  there.  You  would  have  left  un- 
explained the  fact  that  many  of  the  non- Arctic  types 
which  are  present  in  the  Alps  are  represented  in  the 
mountains  of  distant  regions,  not  by  the  same, 
but  by  allied  species,  which  must  have  descended 
from  a  common  ancestor;  that  one  species  of 
Wulfenia>  for  example,  inhabits  one  small  corner  of 
the  Alps,  that  another  is  found  in  Northern  Syria, 
while  a  third  allied  species  has  its  home  in  the 
Himalaya."  Mr.  Ball  is  of  opinion  (p.  584)  that 
the  effects  of  the  Glacial  period  have  been  greatly 
overrated.  "  Even  during  the  period  of  maximum 
cold  the  highest  ridges  of  the  Alps  were  not  com- 
pletely covered  with  snow  and  ice;  for  we  still  see 
by  the  appearance  of  the  surface,  the  limit  above 
which  the  ancient  ice  did  not  reach,  and  in  the  middle 
zone  the  slopes  that  rose  above  the  ancient  glaciers 
had  a  summer  climate  not  very  different  from  that 
which  now  prevails.  In  my  opinion  the  effect  of  the 
Glacial  period  on  the  growth  of  plants  in  the  Alps 


THE  ALPINE   FAUNA.  347 

was  to  lower  the  vertical  height  of  the  zones  of 
vegetation  by  from  one  to  two  thousand  feet."  He 
acknowledges  that  there  was  probably  a  moderate 
diminution  of  the  mean  temperature  of  Europe  with 
an  increased  snowfall,  so  as  to  cause  a  great  extension 
of  glaciers  on  all  the  mountains  of  Northern  Europe. 
"But  that  the  clim.ate  of  Middle  Europe  was  such  that 
the  plants  of  the  high  Alps  could  spread  across  the 
plains  seems  to  me  an  improbable  supposition" 
(p.  584). 

On  the  Continent,  also,  some  botanists  seem  to 
feel  that  Forbes's  theories  of  the  origin  of  the 
Alpine  flora,  which  were  at  first  hailed  with  such 
delight,  and  accepted  by  almost  every  naturalist  as 
the  final  verdict,  must  be  modified  in  -  the  light 
of  recent  researches.  Professor  Krasan  believes  that 
many  plants  which  now  live  in  the  high  Alps 
flourished  in  pliocene  times  at  sea-level  (p.  37). 
"  Especially  the  evergreen  species  exhibit  the  im- 
pression of  an  originally  mild  climate — of  a  climate 
without  winter  frosts — for  otherwise  the  plants 
would  have  developed  into  species  with  deciduous 
leaves."  To  the  favourable  conditions,  consisting  in 
periodic  snowfalls  and  high  summer  temperature, 
must  be  attributed  the  fact  that  in  the  highlands  so 
many  more  species  from  Tertiary  times  have  survived 
than  in  the  plains.  The  temperature  was  probably 
much  higher  during  the  Glacial  period  than  is 
generally  believed.  The  climate  was  more  moist, 
thus  contributing  to  an  abundant  snowfall,  while  the 


348        HISTORY   OF   THE   EUROPEAN    FAUNA. 

survivors  of  ancient  Tertiary  times  were  able  to 
repeople  the  parts  which  were  temporarily  devastated 
by  the  advancing  glaciers. 

In  so  short  a  chapter  it  is  impossible  to  deal  with 
the  Alpine  fauna  in  a  manner  more  deserving  of  this 
theme.  I  have  merely  sought  to  give  a  sketch  of 
the  general  outlines  of  the  subject  and  to  suggest 
another  possible  mode  of  origin  of  Alpine  animals 
than  that  currently  believed  in  by  naturalists.  It  is 
to  be  hoped  these  suggestions  will  be  useful  to  those 
intending  to  reinvestigate  the  problems  raised  in  this 
chapter.  When  our  knowledge  of  the  fauna  of  Asia 
is  more  complete,  it  will  be  possible  to  give  a  more 
thorough  and  in  many  respects  a  more  satisfactory 
history  of  the  European  fauna  than  at  present. 


SUMMARY  OF  CHAPTER  VIII. 

In  early  Tertiary  times  the  area  now  covered  by  the  European 
Alps  was  covered  by  the  sea.  Islands  slowly  rose  above  the 
surface  of  the  waters,  which  finally  coalesced  to  form  a  peninsula 
connected  with  the  mainland  in  the  east.  Animals  now  began 
to  invade  the  new  territory  which  continued  to  rise,  while  the 
sea  retired  farther  and  farther  to  the  north  and  south.  During 
the  Pliocene  Epoch  the  sea  ceased  to  wash  the  northern  shores 
of  the  Alps,  and  both  emigration  and  immigration  became 
possible  in  that  direction,  and  also  from  and  to  the  west. 

The  Alpine  fauna  and  also  the  flora  are  made  up  of  a  number 
of  elements,  the  eastern  one  being  the  oldest.  The  latter  is 
represented  in  the  Alps  by  the  older  and  newer  Oriental  migra- 
tion. The  general  range  of  the  Alpine  Steinbock,  Chamois, 
Marmot,  Vole,  Shrew,  and  Hare  are  specially  referred  to.  The 


THE   ALPINE   FAUNA.  349 

Alpine  birds  are  few  in  number,  and  all  of  them  are  readily 
traceable  to  an  Asiatic  ancestry.  Among  the  Amphibia,  the 
Salamanders  are  considered  of  Alpine  origin. 

Dr.  Kobelt  tells  us  that  a  uniformity  of  character  marks  the 
Alpine  molluscan  fauna.  Campylaea, — often  considered  a  sub- 
genus  of  Heli.\\ — Pomatias,  Zonites,  are  looked  upon  as  truly 
Alpine  genera.  For  very  long  periods  the  Alps  seem  to  have 
received  no  addition  to  their  molluscan  fauna  from  other  areas. 
The  case  is  very  different  with  the  Lcpidoptera^  some  of  the 
most  striking  species  being  evidently  Asiatic  immigrants. 
Some  examples  of  Coleoptera  and  Orthoptera  are  mentioned, 
and  their  origin  discussed. 

We  find  as  the  result  of  these  considerations  that  the 
majority  of  the  Alpine  species  are  either  indigenous  or  have 
come  from  Asia  with  the  Oriental  migration.  None  of  the 
northern  or  western  immigrants  appear  to  be  among  the 
characteristic  Alpine  species,  and  it  seems  that  the  Siberian 
migrants  have  not  retired  to  the  Alps,  as  some  naturalists  have 
been  led  to  suppose.  It  is  evident  that  the  fauna  must  have 
survived  the  Glacial  period  on  the  Alps,  though  according 
to  geological  evidence  glaciers  of  enormous  size  originated 
on  these  mountains. 

The  identity  of  many  Alpine  species  with  Scandinavian  ones 
appears  at  first  sight  due  to  a  direct  migration  from  the  Alps 
to  Scandinavia  or  vice  versd.  Perhaps  such  a  migration  has 
taken  place  to  some  extent,  but  it  is  probable  that  from  a  Central 
Asiatic  centre  some  species  spread  across  Arctic  America  into 
Northern  Europe,  and  also  westward  to  the  Alps.  The  Grouse 
family  forms  an  interesting  example. 

There  are  two  older  theories  which  explain  the  similarity 
between  the  Scandinavian  and  Alpine  faunas.  Forbes's  view, 
which  gained  most  adherents  among  naturalists,  was  that  the 
Scandinavian  and  Alpine  animals  were  driven  into  the  plain  by 
the  cold  during  the  Glacial  period,  and  when  they  ultimately 
regained  their  homes,  some  individuals  of  the  northern  species 


350          HISTORY  OF   THE    EUROPEAN    FAUNA. 

moved  southward,  and  a  few  of  the  southern  ones  northward. 
By  the  more  recent  theory  of  Nehring,  the  Siberian  animals 
which  invaded  our  continent  from  the  east,  and  then  spread 
northward  to  Scandinavia  and  southward  to  the  Alps,  formed 
the  nucleus  of  the  faunas  of  these  two  areas.  The  objections 
to  both  of  these  views  are  fully  set  forth  in  this  chapter. 

A  few  remarks  on  the  botanical  aspect  of  the  Alpine  problem 
conclude  the  chapter.  The  origin  of  the  flora  has  been  ex- 
plained in  a  very  similar  manner  to  that  of  the  fauna.  But 
already  Ball  and  Krasan  have  raised  their  voices  against  the 
current  theories,  as  the  facts  of  distribution  appear  to  them 
more  satisfactorily  explained  on  lines  more  consonant  with 
those  which  I  have  used  in  discussing  the  origin  of  the  Alpine 
fauna.  One  of  the  most  important  conclusions  obtained  by  this 
study  of  the  flora  in  conjunction  with  the  fauna,  is  that  I  have 
emphasised  in  most  of  the  preceding  chapters — viz.,  that  the 
Glacial  period  in  Europe  was  not  a  time  of  extreme  cold, 
and  that  its  destructive  effect  on  the  animals  and  plants  was  by 
no  means  such  as  is  currently  believed. 


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Nathorst,  A.  G.— Kritische  Bemer- 
kungen  liber  die  Geschichte  d. 
Vegetation  Gronlands,  "  Botan- 
ische  Jahrbucher,"  vol.  xiv., 
1891. 

Nehring,  A. — "  Tundren  und  Step- 
pen  der  Jetzt  undVorzeit,"  1890. 

Neumayr,  M. —  "Erdgeschichte," 
vol.  ii.,  1887. 

Ogilvie,  W.  R.  Grant.— "Game 
Birds,"  1895. 

Penck,  A. — Die  grosse  Eiszeit, 
"HimmelundErde." 

Petersen,  W. — Die  Lepidopteren- 
fauna  d.  arktisch.  Gebiete  von 
Europa,  "  Beitrage  z.  Kenntniss 
d.  Russ.  Reichs,"  vol.  iv.,  1888. 

Pettersen,  K.— Arktis,  "Arch.  f. 
Math,  og  Naturvid.,"  1882. 

Pohlig,  H. — Dentition  und  Kranio- 
logie  des  Elephas  antiquus  (part 
2),  "Nova  Acta  Acad.  Carol. - 
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Reade,  Mellard— a.  The  Present 
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"Geological  Magazine,"  vol.  iii., 
1896.  b.  High  Level  Shelly 
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Reid,  C. — The  Pliocene  Deposits 
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Riitimeyer,  L.  — "Die  Herkunft 
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Sars,  G.  O. — Crustacea  Caspica, 
"Bull.  Acad.  Imp.  d.  Sciences 

23 


354 


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04. 

Scharff,  R.  F. — a.  On  the  Origin 
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deposit,  "Trans.  Geol.  Soc. 
Glasgow,"  vol.  x.,  1896. 


INDEX. 


Abkpharus  pannonicu s,  258 
Accentor  collaHs,  316 

modidaris,  317 

Accidental  distribution,  12,  26 
Acipenser  ruthenus,  29 
Acme,  323 

Adams,  Leith,  112,  150 
/Egean    continent,    subsidence    of, 

272-273 

Agriolimax,  284 
AlactagajacuhiS)  204 
Aha  impennis,  92,  142 
Alligator,    accidental    dispersal  of, 

14 
AHolobophora  Georgit,  115,  302 

veneta,  115,  302 
Alpine  Accentor,  316 

Chough,  317 

fauna,  survival  of,  during  Glacial 
period,  332 

flora,  341-348 

flora,  age  of,  78,  79 

flora,  suitable  conditions  for,  78- 

79 

Hare,  as  a  test  of  climate,  316 
Snow-finch.  318 
Alps,  component  elements  of  fauna, 

311-312 

Alston,  E.  R.,  312 
Alytes  cisternasii,  295 

obstetricans,  295 
Amalia,  284 

America,  introductions  from,  24 
American    beetles    in    the   British 

Islands,  167 
plants,  origin  of,   in  the   British 

Islands,  144 
plants  in  Ireland,  1 66 
sponges  in  Ireland,  166 
Anipelis  garruhtst  205 


Amphibia  of  Europe,  193-194 

dispersal  of,  21 
Aniphicoma,  268 
Andalusian  Bush-quail,  291-292 
Anergates,  167 
Anglo-Scotian  fauna,  95 
Animals  as  tests  of  climate,  71-75 
Antelope,  39 

Ants,  European  origin  of,  161 
Apollo,  325 

Apus  glacialiS)  94,  167 
Aralo- Caspian    Sea    and    Arctic 
Ocean,     connection    between, 
219-231 

Arbutus  unedo,  IIO,  307 
Arctic  animals  in  Caspian  Sea,  238 

flora  in  Europe,  238-241 

Hare,  origin  of,  148-149,  158-159 

Hare,  range  of,  136-138 

Lepidoptera,  200 

plant-beds  at  Bovey  Tracy,  238 

plants,  143-144 

plants,  delicacy  of,  185 

mollusca  in  Red  Crag,  235-236 

Seal,  174 

sub-region,  132 
Arctotnys  bobak,  204,  313 

mannoi 7a,  313 
Arion,  48,  299 

snbf^tsc^ls,  49 

Arionidie,  origin  of,  48,  299 
Asiminea  Grayana,  99 
Aucapitaine,     Baron,     experiments 

by,  15 

Autochthonous  species,  10 
Azores,  remains  of  a  continent,  19 

Bacillus,  269-270 
Badger,  distribution  of,  43 
Balea  perversa,  298 


INDEX. 


Ball,  J.,  77,  345-347 

Baltic  Sea,  fauna  of,  174 

Baltic  and  White  Seas,  connection 

between,  175 

Banded  Lemming,  range  of,  138 
Barn  Owl,  98 
Barrett- Hamilton,    G.   E.   H.,  29, 

90,  139,  149 

Barriers  to  animal  migration,  38 
Barrington,  R.  M.,  105 
Bathyphantes  nigrinus,  94 
Baur,  G.,  19 
Bearded  Titmouse,  292 
Beaver,  203 

Beaver  absent  from  Ireland,  121 
Beddard,  F.  E.,  19,  58 
Bedriaga,  J.  von,  259-260 
Beetles  common  to  North  America 

and  Europe,  161 

American,  in  British  Islands,  167 
Bell  (vide  Kendall  and  Bell) 
Black  Cock,  143 
Black  Grouse,  336 
Blanchard,  E.,  282 
B latins  cinereus,  279,  294 
Blelhisa  multipunctata,  93 
Blytt,  A.,  34,  52,  79,  185,  239 
Boar,  distribution  of,  44 
Bobak  Marmot,  204 
Boettger,  O. ,  48,  263 
Bogdanov,  M.  M.,  53 
Bombinatof,  259-260 
igneus,  259 
pachypus,  259 
Bonnet,  306 
Bonney,  T.  G.,  66,   70,  83,    180, 

229 

Boulenger,  G.  A.,  259 
Boulder-clay,  foraminifera  in,  84 
origin  of,  81,  175,  180-181,  229- 

230 
of  Continental  Europe,  180,  226- 

231 

Bourguignat,  J.  R.,  18,  47,  302 
Bovey  Tracy,  arctic  plant  beds  at, 

238 

Boyd,  E.,  104 
Branchinecta  palludosa,  167 
Brandt,   J.    F.,   51,    62,    107,   218, 

222 

Brauer,  A.,  132-133 


Brehm,  A.  E.,  139 

Bristle-fern,  115 

British  flora  during  Glacial  period, 

163 

British  Islands,  Lusitanian  flora  in, 
288,  306-307 

submergence  of,  127 
Brooke,  Sir  V.,  247,  250 
Brunner  von  Wattenwyl,  270 
Btifo  calamita,  30 
Buliminus  detritus ,  261 

fasciolatus,  261 

obscurus,  261 

pupa,  261 

Bullfinch,  origin  of,  191,  255-256 
Bulman,  G.  W.,  114 
Bunge,  D.,  219 

Bush-quail,  Andalusian,  291-292 
Butterflies,    Arctic   distribution   of, 
159-161 

origin  of  North  European,  55 

Caccabis  rufa,  29 

Campylaea,  sub-genus  of  Helix,  49, 

321 

Canary  Islands,  origin  of,  19 
Cam's  lagopus,  135 
Capercaillie,  28,  143,  336-337 
Capra  ibex,  312 

tzgagrus,  61,  312 

pyrenaica,  312 

sibirica,  312 

sinaitica,  312 
Carabus,  199 
Carduelis,  257 

caniceps,  258 

elegans,  257 

major t  257 
Carpenter,  G.   H.,  64,   94-95,   104, 

114,  121-123,  288>  301 
Caspian  Fishes,  224 

post-pliocene  deposits;  222,  226 

Sea,  Arctic  animals  in,  238 

seal,  224 
Castor  fiber,  203 
Cave  deposits,  mixture  of  Northern 

and  Southern  animals  in,  54 
Cenocosmic  species,  24 
Centre  of  distribution,  12,  43,  47 

origin,  shifting  of,  202 
Cervida,  origin  of,  248 


INDEX. 


357 


Cervus  dama,  251 

elaphus,  246-250 

giganiciis,  108,  251 
Chaincelcon  vulgaris,  279 
Chamois,  312 
Charrs,  origin  of,  124 
Chioglossa,  296 
Chough,  Alpine,  317 
Christ  H.,  52,  343'344 
Christy,  Miller,  105 
Chrysochraon,  330 
Cicindela,  197-198 
Cinclus,  255 
Clarke,  Eagle,  105 
Clausilia,  47,  262-264,  284 

bidentata,  47,  262,  284 

biplicala,  47 

faminafa,  47 

Pauli,  48,  263 

Rolphii,  47 
Claviger,  267-268 

tes/acetts,  268 

Climate   in  Glacial    period,  65  8 1, 
127,  149,  182-183 

in  Pleistocene  Europe,  208-209 
Close,  Maxwell,  129 
Cod-fish,  origin  of,  143 
Coecilianella,  17 
Coenonyjiipha  typhon,  93 
Cole,  G.  J.,  83,  106-107 
Coleoptera   of  Europe,    origin   of, 

197 

common   to   Europe   and   North 

America,  161 
Colias,  266 
Coregonus,  124 

clubeoides,  124 

pollan,  124 

vandesiiiS)  124 
Cosmopolitan  species,  24 
Coitus,  92-93 

bubalis,  93 

gobio,  93 

quadricornis,  175 

scorpio,  93 

Credner,  R.,  177-178 
Cricetus  frunifntariuS)  190 

vttlgaris,  204 

Crimea  formerly  an  island,  35 
Crimean  fauna,  57 
Croll,  J.,  66 


Crocidura,  253 

etnisca,  253,  279 
Ctitiiciilus  tonjuatus^  138 
Current,  Arctic,  172-173 
Cyanopolius  Coo/a,  293 

cyamts,  293 
Cyclostoma  elegans,  16 

Danais  chrysipftts,  267 
Dartford  Warbler,  288 
Darwin,  C,  13-15,  17-19,  25,  32- 

33,  339 

Daudebardia,  323 
Daulias  luscinea^  192 
Dawkins,  Boyd,   53,  62-63,  72-73, 

107,    II2-II3,     120,     208,     222- 

223, 282 

Day,  F.,  29 
Deperet,  C.,  44,  276 
Dippers,  origin  of,  255 
Discoglossus pictus,  279,  295-296 
Dispersal  of  Amphibia,  59 

of  beetles  [apterous],  59 

of  British  butterflies,  113 

of  earthworms,  58 

of  planarian  worms,  59 

of  spiders,  59 

of  wood-lice,  59 
Distribution,  centres  of,  12,  43,  47 

discontinuous,  114 
Dormouse,  316 

absent  from  Ireland,  121 
Drapetisca  socialis,  94 
Dreyssensia  polymorpha,  26,    230- 

231 

Drift,  a  marine  deposit,  129 
Drude,  O.,  52,  77 
Dryas  octopetela,  79,  238 
Dual  origin,  possibility  of,  38 
Dyer,  Th.,  79 

Earth-worms,    distribution    of,    19, 

23,.  58 

Edelweiss,  266,  342 
Egean    Continent,    subsidence    of, 

272-273 

Eider-duck,  range  of,  141 
ElepkttmtidaSi  origin  of,  202,  252- 

253 

Elephas  primigemus,  214,  252 
Emery,  C.,  161,  167 

23* 


358 


INDEX. 


Endemic  species,  10 

Engler,  A.,  52,  145,  171,  176,  282, 

341-342 

English  Hare,  29 
Ephydatia  craierifonnis,  94 
Epoccus,  167 
Erebia,  325-326 

athiops,  325-326 

epiphron,  325 

glacialis,  326 

lappona,  326 
Eremias,  258 
Erratics,  181-182 
Eryx  jacu/us,  259 
Euphorbia  hiberna,  307 
European  beetles,  origin  of,  197 

butterflies,  origin  of,  200 

land   and    fresh-water    mollusca, 
origin  of,  196 

mammals,  origin  of,  106,  193 
Etirynebria  coniplanata,  302 
Evotomys  Nageri,  314 
Extension  of  range,  mode  of,  38 

Fallow  Deer,  251 

Falsan,  A.,  66,  68-69,  7*»  73 

Feilden,    H.    W.,    13,    77,    84-85, 

164 
Finmark,    Greenland   mollusca    on 

coast  of,  171 

Fire-toads,  origin  of,  259 
Fischer,  P,,  103 
Fishes,  Caspian,  224 
"  Flotsam  and  jetsam  "  theory,  20 
Foraminifera  in  boulder-clay,  84 
Forbes,  E.,  64,  101,  no.  114-115, 

171,  288,  339 

Forest-Bed  an  inter-glacial  deposit, 
70 

corresponding       to      continental 
inter-glacial  deposits,   120 

fauna  of,  232-234 

mollusca,  212-213 
Fossil  glaciers,  220 
Fresh-water  faunas,  origin  of,  177 
Fringilla,  293 

ccelebs,  293 

madeirensis,  293 

•niontifringilla,  293 

spodiogenys,  293 

teydea,  293 


Frog,  introduction  of,  into  Ireland, 


Gadow,  H.,  139 
Galapagos  Islands,  19 
Gale  odes,  270 

Gammaracanthus  relictus,  179 
Gardner,  J.  S.,  145 
Garnieria,  sub-genus  of  Clausilia,  48 
Gasterosteus  aculeatus,  92 
Gaudry,  A.,  73,  273 
Geikie,  Sir  A.,  116,  309 
Geikie,  J.,   59,   66-67,    70,    75-76, 
80-83,  116,  129,  163,  181,  216- 
217,  226-227,  233,  235,  238 
Geographical   changes,   importance 

of,  64 
Geomalacus   maculostis,   5>   49?   99? 

102,  115,  298-299 
Gervais,  E.,  150 
Glacial  climate  in  France,  150 

period,    climate   of,    65-81,    127, 
149,  182-183 

period  in  Scandinavia,  176 

period,  survival  of  animals   and 

plants  during,  65 
Glaciation  of  Ireland,  129 
Glutton,  119,  203 
Goldfinch,  origin  of,  257 
Gonepteryx,  296-297 

ckobule,  297 

cleopatra,  296 

rhamni,  296 
Gould,  J.  E.,  20 
Great  Auk,  range  of,  92,  142 
Greenland  flora,  161-162 

flora,  survival  of,  163-164 

miocene  temperature  in,  146 

mollusca  on  coast  of  Finmark,  1  7  1 

Tertiary  plants,  144-145 
Grouse,  91,  334 

black,  336 
Gu/o  hiscus,  119,  203 

Ilaacke,  W.,  147 
Hamilton,  John,  161 
Hamster,  190,  204 
Hanitsch,  R.,  94 
Hare,  2,  29,  90 

Arctic,  2,  90-91 

English,  29 


INDEX. 


359 


Hare,  Alpine,  as  a  test  of  climate, 

316 

Harle,  E.,  150 
Harmer,  F.  W.,  182 
Harvest- mouse,  3,  190 
Harvie-Brown,  J.  A  ,  105 
Plaughton,  W./86 
Hazel-grouse,  337 
Hedge  Accentor,  317 
Hedley,  C,  20 
Heer,  O.,  144-145 
Helix,  4 

actifa,  102,  298 

aculeata,  298 

aspersa,  24,  195 

er ice  tor  urn,  298 

fusca,  298 

lapicida.)  4 

obvolula>  4,  297 

pi sana,  102,  298 

pomatia,  15,  32,  195,  262 

revelata,  102 

rotundata,  122,  274,  298 

ruder  at  a,  212-213 
Herdman,  W.  A.,  and  Lomas,  J., 

236 

Herring,  origin  of,  143 
Heteroineyenia  Ryderi,  94 
Hofman,  E.,  54 
Hooker,  Sir  J.,  99,  161 
Hopatroides  thoracictis,  268 
Howorth,  Sir  H.,  73 
Hyalinia,  antiquity  of,  50 
Hyla,  260 

arborea,  260-261,  280 

chinensisy  260 

Ice-Age,  climate  of,  65.  81 
Ice-bridge,  migration  on,  108 
Idotea  entomon,  178,  224 
Ihering,  H.  von,  19,  20,  24-25 
Indigenous  species,  10 
Inter-glacial  deposit,  the  Forest-Bed 
an,  70 

periods,  climate  of,  181,  218 
Introduced  species,  10,  23,  27-29 
Introduction  by  man,  32-33 
Introductions  from  America,  24 
Ireland,  glaciation  of,  129 
Irish  and  Scotch  Hare,  136 

Elk,  108,  251-252 


Irish  fauna,  composition  of,  63 

Stoat,  136 

Isle  of  Man,  fauna  of,  123 
hotoma  lit  tor  a  Us,  94 

Jeffreys,;.  G.,  27 
Jerboa,  204 
Jordan,  H.,  103 
Judd,  J.  W.,  61 

Karpinski,  H.,  222 

Kendall,  P.  J.,  125 

Kendall,  P.  J.,  and  Bell,  A.,  174 

Kennard,  A.  S.,  167 

Kennard,    A     S.,  and  Woodward, 

B.  B.,  4 

Kessler,  H.,  222 
Kew,  II.  W.,  17,  23,  32 
Killarney  fern,  115 
Kinahan,  G.  H.,  107-108,  129 
Kirkdale  Cavern,  remains  in,  54 
Kobelt,    W.,   49,    56,   58,  62,  75, 

195-196,  212,    281,    297,   303, 

321-323 

Koppen,  F.  T.,  51,  62,  222,  248 
Krasan,  F.,  347 

Lagomys,  origin  of,  41,  203 
Lagopus,  334 

albus,  91,  142,  334 
hyperboreus,  334 
leucttnts,  335 
mutus,  91,  142,  334 
1-upestris,  334 
scoticus,  334 
LaDiinifera,)  sub-genus  of  Clatisilia^ 

48 

Lamplugh,  G.  W.,  107-108 
Land-connection  between  America 

and  Northern  Europe,  61 
British   Islands  and  France,  59- 

60 
British  Islands  and  Scandinavia, 

61 

Europe  and  North  Africa,  61 
Europe  and  North  America,  6 1, 

168-172 

Greece  and  Asia  Minor,  61 
Greenland  and  Europe,  155,  159 
Ireland  and  Spain,  no 
Land  Mollusca,  migrations  of,  9 


360 


INDEX. 


Land  shells,  West  Indian,  21 
Lartet,  E.,  51,  73,  107,  150 
Lemming,  range  of,  138-140 
Leon'.opodium  alpinu/n,  266,  342 
Lepus,  27,  29,  31 
)  136 

iS)  27,  31,  291 
us,  29,  316 

glacialis,  136 

lacostei,  291 

variabilts,  2,  29,  136,  315 
Lepidoptera,  range  of,  159-160 

Arctic,  200 

surviving  Glacial  period,  54 
Leuckart,  R.,  176 
Lewis,  Carvill,  129 
Liwax,  284 

marginal-its ;  284 

maximus,  284 

Limnocalanus  macriirusy  ]  79 
Linyphia  insignis,  94 
"  Loess"  fauna,  75,  196 
Lomas  (vide  Herdman  and  Lomas) 
Loven,  S.,  177 

Lusitanian  flora  in  British  Islands, 
288,  306-307 

spiders,  301 
Lydekker,   R.,  14,  22,  32,  58,  157, 

183,  202,  248,  252 

Lyell,  Sir  C.,  14,  22 
Lyrtirtis  tetrix^  336 

Macro ptyckia,  sub-genus  of  Clatt- 
silia,  48 

Madeira,  19 

molluscan  fauna  of,  25 
remains  of  a  continent,  19 

Major,  Forsyth,  45-46,  280,  282 

Mallet,  R.,  86 

Mammoth  in  Siberia,  214-217 
range  of,  252-253 

Mammals,  dispersal  of,  9,  21 

Mantis  religiosa,  269 

Maps,  general  plan  of,  8 

Margaritana,  93 

Marine  connection  between  Caspian 
Sea  and  Arctic  Ocean,  62,  219- 
231 

mollusca,  distribution  of,  236 
origin  of  boulder-clay,  82-86,  228- 


Marine  shells  above  sea-level,  127- 
128 

transgression  in  Northern  Russia, 

172 
Marmot,  Alpine,  313 

Bobak,  204 
Marsh-ringlet,  93 
Marshall,  Rev.,  id6 
Martins,  C,  68 
Mediterranean    land-  connections, 

276-282 
Meles,  43 

albogularis,  44 

anakunia,  44 

t'eucurus,  44 

maraghanus,  44 

polaki,  44 

taxus,  43 
Melitcea  aster ia,  325 

cynthia,  325 

mattirna,  325 
Melizoph  :lus  undatus,  288 
Merriam,  C.  H.,  38 
Metoponorthus  cingendus,  301 
Microtus,  313 

brecciensis,  314 

leucurus,  314 

nivalis,  313 
Mid- wife  Toad,  295 
Migrations,  8-9 

cause  of,  208 

of  British  shore-forms,  124 
Migration  from  Asia  to  Europe  by 

North  America,  327 
Migration  on  ice-bridge,  108 

waves  of,  204-206 
Miller's  thumb,  93 
Milne-Edwards,  A.,  282 
Mingling  of  Southern  and  Northern 

Mammals,  72-75,  209 
Miocene  geography,  274-276 

temperature  in  Greenland,  146 
Mi o gale  mo  s  chat  a  ^  290 

pyrenaica,  290 
Molge  alpestris,  320 

cristata,  320 

montana,  320 

palmata^  320 

rusconii,  320 

vulgaris,  319 
Mollusca  in  Loess,  196 


INDEX. 


361 


Mollusca,  distribution  of  marine,  236 
Molluscan  fauna,  divisions  of  British, 
102 

Madeira,  25 

Porto  Santo,  25 
Molluscs,  dispersal  of,  17,  24 
Mongoose,  28 
Montifringilla  nivalis,  318 
More,  A.  G.,  104 
Moschler,  E.,  160 
Motacillct)  254,  292 

alba,  292 

boreal  is,  255 

campestris,  254 

lugubris,  292 

melanope,  254 
Mountain  Avens,  79 
Mountain-ringlet,  325 
Mouse,  distribution  of  Harvest,  3 
Mud-glaciers,  86 
Murchison,  R.,  175,  230 
Murray,  Andrew,  32-33,  150,  154 
Muscardinus  avellanarius,  316 
Musk-Ox,  range  of,  119,   134-135, 

203. 

Mus  minutusy  3,  95,  190 
Miistela  africana,  279 

boccamela,  279 

erminea,  135- 136 

putorius,  190 
Myodus  lemmuS)  138 

obensis,  138 
Myoxus,  316 
Mysis  caspica,  223 

necropthalma,  223 

relicta,  179,  223 

Nathorst,   A.    G.,    163,    169,    240- 

241 

Natterjack  Toad,  30 
Nebria  atrata,  328 

livida,  328 
Nehring,    A.,    96,    107,    119,    196, 

208-211,  340 

Nenia,  sub-genus  of  Claiisilia,  48 
Neumayr,  M.,  19,  67 
Newt,  319 
New    Siberian    Islands,    origin    of 

bone-beds,  219 
extinct  fauna  of,  218 
Nightingale,  192 


Nordquist,  179 

North  American  marine   mollusca 

in  crag  deposits,  173 
North  European  Sea,  172 

Ocean  basins,  permanence  of,  18 
Oceanic  Islands,  18 
Oeneis,  326-327 

aello,  326 

Onychogomphus,  268 
Oriental  migration,   old   and  new, 

272 

Oriental  plants,  282-283 
Origin,  centre  of,  shifting,  202 
Otiorrhynchus  auropunctatus,  115, 

302 
Ovibos  moschatus,  119,  134-135,  203 

Pacific  Continent,  20 
Painted-lady  Butterfly,  98 
Palincosmic  species,  25 
Pallas's  Sandgrouse,  205 
Panurus  biarmicus,  292 
Fapilio  hospifon,  265 
Parmacella,  49 
Parnassitis,  265-266,  324 

apollo,  265,  324 

delitts,  324 

mnemosyne,  324 

Nordmanni,  324 
Partridge,  29 
Pelias  beruS)  192 
Pelobates,  295 
Pelodytes  punclattts,  295 
Pelophila  boreatis,  93 
Penck,  A.,  66 
Perches,  origin  of,  143 
Periops  hippocrepis,  279 
Peschel,  O.,  177 
Petersen,  VV.,  55,  154,  160,  200 
Petronia  stulta,  318 
Pezotettix,  329-330 

alpinus,  329 

pedestris,  329 
Phasianus,  256-257 

cokhicus,  31,  257 
Pheasant,  origin  of,  256-257 
Phoca  annelata,  174 

caspica,  224 


Phcedusa,    sub-genus   of   Clausili 


362 


INDEX. 


Phylloxera  vastalrix,  24 
Pigs,  origin  of,  44-46 
Pika,  41 

Pine-Grosbeak,  191 
Pinicola  enucleator,  191 
Planorbis  dilatetus,  24 

glaber,  167 
Plants,  American,  in  Ireland,  166 

as  tests  of  climate,  75-80 

migration  of,  34 
Platyarthrus,  299-302 
Plectrophenax  nivalis,  140 
Pleistocene   climate   of  Asia,    210, 
214-215 

Europe,  climate  in,  208-209 

fauna,     northern    and     southern 
animals  in,  72-75 

mollusca,  211-213 
Pliocene  deposits  of  Sicily,  277 

geography,  276-277 
Pohlig,  H.,  176 
Polar  Bear,  134 

Fox,  range  of,  135,  149 

origin  of  animals,  147 
Pole-cat,  190 

Polydesnius  gallicus,  115,  302 
Polynesian  Islands,  20 
Pomalias,  49,  321-322 
Pontoporeia  affiu/s,  179 
Porto  Santo,  molluscan  fauna  of,  25 
Pouched  Marmot,  41 
Praeger,  R.  L  ,  298 
Praying  insect,  269 
Proboscidea,  origin  of,  252 
Psanirnodromus  algirus,  295 

hispanicuS)  295 
Ptarmigan,  91,  142 
Ptipa,  antiquity  of,  50 
Pupa,  297-298 

anglica,  298 
granum,  297 

ringens,  102 

Pyrrhocorax  alpimis,  317 
Pyrrhula,  191,  255 

e^^rop<za,  191,  256 

major,  191,  256 

Rabbit,  introduction  of,  27,  31,  291 
Rana  temporaria,  194,  320 
Range,  extension  of,  38 
Rangifer  tarandus,  133,  150-158 


Reade,  Mellard,  127-129 

Red  Crag,  Arctic  mollusca  in,  235- 

236 

Red  Deer,  246-250 
Red  Grouse,  91,  142 
Red-legged  Partridge,  29 
Reguhis  ignicapilhis,  257 
maderensis,  257 
teneriffiz,  257 
Reid,  Clement,  76,  163 
Reindeer,  range  and  varieties  of,  91, 

133,  149-158 

Relict  lakes,  176-179,  223 
Reptiles,  dispersal  of,  21 
Reptiles  of  Europe,  192-193 
Rhax,  270 

Rhinoceros,  distribution  of,  214-216 
Rhododendron  ponticum^  271 
Rhopalomesites  Tardyi,  302 
Rock  Sparrow,  318 
Route    of    migration    of    Siberian 

mammals,  210 
Rupicapra  tragus,  312 
Russia,     marine    transgression    in, 

172 

submergence  of  North,  155 
RiUimeyer,  L.,  43,  57,  281 
Ryothemis,  269 

Saiga  Antelope,  39,  204 
Saiga  tartarica,  39,  204 
Salamandra  atra,  319 
caitcasica,  319 
maculosa,  319 

Salnionidce,  origin  of,  92,  143 
Sandgrouse,   migration  of  Pallas's, 

41,  205 

Sars,  O.,  177,  223-224 
Saunders,  H.,  256 
Seal,  Arctic,  174 
Sea-urchin,  125 
Sedgwick,  A.,  86 
Scandinavia,    absence    of    Oriental 

and   Siberian   mammals   from, 

176,  206 

during  Glacial  period,  176 
Scandinavian    boulders,    origin    of, 

172-173. 

butterflies  in  America,  167 
flora,  antiquity  of,  162,  185 
glaciers,  172 


INDEX. 


363 


Scandinavian  lepidopterous  fauna,  55 

Schulz,  A.,  52,  155 

Sclater,  P.  L.,  95 

Scotch  Argus,  325 

Scotch  and  Irish  Hare,  136 

Siberia,  climate  of,  during  Glacial 

period,  217 

Siberian   birds   in   British    Islands, 
191-192 

mammals    absent    from    Scandi- 
navia, 176,  206 

mammals  absent  from   Southern 
Europe,  206 

mammals  in  Great  Britain,  95-96, 
190,  202-204 

migration,  date  of,  208 

steppe-fauna,  39 
Sicily,  Pliocene  deposits  in,  277 
Silurus  glanis,  29 
Simpson,  C.  T.,  21 
Simroth,  H. ,  48,  299 
Sjogren,  H.,  226 
Slugs,  dispersal  of,  16-17 
Snails,  dispersal  of,  17 
Snow-Bunting,  range  of,  91,  140 
Snow-Finch,  318 
Sollas,  W.,  177 
Somateria  mollissima,  141 
Sorex^  202,  3 14-3 1 5 

alpinus,  3 14-3 1 5 

araneus,  315 

minulus,  315 
Southern  mammals  in  Arctic  Siberia, 

213-216 

Sparrow,  introduction  of,  28 
Spcmtop1tilust  41 
Speyer,  A.  and  A.,  55,  200 
Spiders,  Lusitanian,  301 
Sponges,  American,  in  Ireland,  166 
St.  Erth  fauna,  174 
Steinbock,  312 
Steppe-fauna,  Siberian,  39 
Sterlet,  introduction  of,  29 
Stick  insect,  269 
Stickleback,  origin  of,  92,  143,  166- 

167 

Stoat,  range  of,  90-91,  135-136 
Strails   of  Gibraltar,   age   of,   277- 

278,  281 

Strix  Jlammea>  98 
Stronglyocentrotus  lividus,  125 


Struckmann,  C. ,  153 
Studer,  Th.,  340 

Sub-marine  plateau  between   Nor- 
way and  Spitsbergen,  154 
Suess,  E.,  61,  273,  282,  303 
Stts  scrofa,  44-46 
Swallow-tail  Butterfly,  264-265 
Syrrhaptes  paradoxus,  41,  205 

Tailless  Hare,  41 
Talpa,  291 
Taylor,  J.  W.,  105 
Tcherski,  J.  D.,  214-215,  222 
Tertiary  plants  in  Greenland,  144- 

145 
Test  ace  I  la,  17,  299 

haliotidea,  299 

maugeii  102,  299 

scutulum.)  299 
Tetrao,  28 

urogallus,  28,  143,  336-337 
Tetrastes  bonasia,  337 
Tettix,  330 

bipunctatus,  330 
Thais  cerisyi)  265 
Thelphusa  fluviatilis,  270,  281 
Thomas,  O.,  90 
Thompson,  W.,  32 
Time     of    disconnection     between 
Great  Britain  and  Ireland,  63 
Titmouse,  bearded,  292 
Toad,  Natter-jack,  30 
Tree-frog,  260,  280 
Trichomanes  radicans,  115 
Tropidonotus  viperimts,  294 
Tubclfa  pensylvam'ca,  94 
Tundras  in  Northern  Europe,  209- 

210 

Turnix  sylvatica,  291-292 
Tyndall,  J.,  68 
Typhlops  lumbricalis,  259 
Tyrrhenian  Continent,  280 

Unio(Maigaritana}  margaritifer^ 
Ursus  maritinwS)  134 
Ussher,  R.  J.,  92,  105 

Vanessa  cardui^  98 

Vertigo  alpestris,  93 
Viper,  range  of,  192 
Vole,  range  of,  313-314 


3^4 


INDEX. 


Wag-tails,  origin  and  range  of,  254, 

292 
Wallace,  A.  R.,   12-13,   18-19,  23, 

58,  99,  in,  304 
Warming,  E.,  76,  163 
Watson,  H.  C.,  100 
Wax-bill,  migrations  of,  205 
Welch,  R.,  298 
West-Indian  land-shells,  21 


White,  Buchanan,  109,  113 
Willow-grouse,  91,  142 
Woodward,    B.    B.    (vide   Kennard 

and  Woodward) 
Wright,  J.,  84 


Zittel,  252 

Zonites,  49,  195,  322-323 


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This  volume  contains  about  400  drawings  of  birds 
by  John  Duncan,  naturalist  and  artist.  The  drawings 
are  accompanied  by  a  concise  description  of  each  bird. 


Mr.  Chas.  Dixon,  in  the  Introduction  to  this  work, 
says: — "Just  a  century  ago,  British  naturalists  and  the 
British  public  were  captivated  by  the  first  volume  of 
Bewick's  classic  History  of  British  Birds — a  work,  com- 
pleted by  the  issue  of  the  second  volume  seven  years  later, 
that  gave  more  or  less  accurate  pictures  of  all  the  species, 
and  one  that  has  retained  a  high  place  in  the  affections  of 
ornithologists  and  artistic  people  generally  down  to  the 
present  time.  To  a  certain  extent,  Mr.  Duncan's  volume 
very  closely  resembles  the  immortal  engraver  on  wood; 
and  in  some  respects,  as  those  who  have  the  pleasure  of 
inspecting  his  remarkable  series  of  drawings  will  find  for 
themselves,  it  is  superior.  The  present  book,  so  far  as 
British  birds  are  concerned,  is  practically  unique.  For  the 
first  time,  an  up-to-date  manual,  containing  an  accurate 
and  artistic  picture  of  every  species,  is  placed  within  the 
reach  of  the  multitude  by  reason  of  its  very  modest  price." 


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The  Natural  History  of  Digestion. 

BY    A.    LOCKHART    GILLESPIE, 

M.D.,  F.R.C.P.  ED.,  F.R.S.  ED. 

Dr.  Gillespie,  who  has  long  been  known  as  an  original  investigator  in 
this  department  of  physiology,  has  in  the  present  volume  attempted  to 
bring  together  all  the  facts  and  recent  discoveries  bearing  on  this  subject 
of  great  scientific  and  practical  importance.  Dealing  with  the  subject  in 
much  detail,  as  well  as  broadly  and  comprehensively,  the  book  appeals 
both  to  medical  specialist  and  general  reader.  The  author  shows  that 
digestion  is  a  process  which  occurs  throughout  animated  nature,  and 
beginning  with  digestion  in  plants,  and  describing  many  original  and 
other  experiments  with  carnivorous  plants,  he  passes  on  to  digestion  in 
the  lower  animals,  and  then  deals  fully  with  the  many  complicated  prob- 
lems offered  by  digestion  in  the  higher  animals  and  man.  The  practical 
questions  of  diet  are  examined  in  their  various  aspects,  and  the  influence 
of  alcohol,  tea,  and  other  stimulants  discussed. 


Crown  8vo,  Cloth,  Price  6s.     With  Illustrations. 

Degeneracy : 

ITS    CAUSES,    SIGNS,    AND    RESULTS. 

BY  PROFESSOR  EUGENE  S.  TALBOT,  M.D.,  Chicago. 

This  volume  presents,  in  a  simple  and  comprehensive  way,  the  basis  of 
fact  on  which  the  speculations  of  Nordau  and  others  have  been  founded. 
It  is  the  first  book  written  in  English  by  a  competent  authority  dealing 
broadly  with  this  subject.  The  author  deals  with  more  especial  fulness 
with  the  signs  of  degeneracy  in  the  head  and  face — ears,  jaws,  teeth,  etc. 
— as  those  which  he  has  chiefly  studied,  and  which  are  of  most  general 
interest ;  but  he  also  discusses  degeneracy  in  the  body  generally,  as  well 
as  its  mental  forms.  The  chief  causes  tending  to  produce  degeneracy  in 
modern  life  are  discussed — heredity,  climate,  foods,  alcohol,  education, 
etc. — and  the  methods  of  combating  them  considered.  The  book  is  fully 
illustrated,  chiefly  from  original  photographs. 

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Great  Writers 

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Museum,  London. 

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LIFE  OF  LONGFELLOW.    By  Professor  ERIC  S.  ROBERTSON. 
LIFE  OF  COLERIDGE.    By  HALL  CAINE. 
LIFE  OF  DICKENS.     By  FRANK  T.  MARZIALS. 
LIFE  OF  DANTE  GABRIEL  ROSSETTI.     By  J.  KNIGHT. 
LIFE  OF  SAMUEL  JOHNSON.     By  Colonel  F.  GRANT. 
LIFE  OF  DARWIN.     By  G.  T.  BETTANY. 
LIFE  OF  CHARLOTTE  BRONTfi.     By  A.  BIRRELU 
LIFE  OF  THOMAS  CARLYLE.     By  R.  GARNETT,  LL.D. 
LIFE  OF  ADAM  SMITH.    By  R.  B.  HALDANE,  M.P. 
LIFE  OF  KEATS.     By  W.  M.  ROSSETTI. 
LIFE  OF  SHELLEY.    By  WILLIAM  SHARP. 
LIFE  OF  SMOLLETT.    By  DAVID  HANNAY. 
LIFE  OF  GOLDSMITH.    By  AUSTIN  DOBSON. 
LIFE  OF  SCOTT.    By  Professor  YONGE. 
LIFE  OF  BURNS.     By  Professor  BLACKIE. 
LIFE  OF  VICTOR  HUGO.     By  FRANK  T.  MARZIALS. 
LIFE  OF  EMERSON.    By  RICHARD  GARNETT,  LL.D. 
LIFE  OF  GOETHE.    By  JAMES  SIME. 
LIFE  OF  CONGREVE.    By  EDMUND  GOSSE. 
LIFE  OF  BUNYAN.     By  Canon  VENABLES. 
LIFE  OF  CRABBE.     By  T.  E.  KEBBEL. 
LIFE  OF  HEINE.    By  WILLIAM  SHARP. 
LIFE  OF  MILL.     By  W.  L.  COURTNEY. 
LIFE  OF  SCHILLER.     By  HENRY  W.  NEVINSON. 
LIFE  OF  CAPTAIN  MARRYAT.     By  DAVID  H  ANN  AT. 
LIFE  OF  LESSING.     By  T.  W.  ROLLESTON. 
LIFE  OF  MILTON.    By  R.  GARNETT,  LL.D. 
LIFE  OF  BALZAC.     By  FREDERICK  WEDMORE. 
LIFE  OF  GEORGE  ELIOT.     By  OSCAR  BROWNING. 
LIFE  OF  JANE  AUSTEN.     By  GOLDWIN  SMITH. 
LIFE  OF  BROWNING.     By  WILLIAM  SHARP. 
LIFE  OF  BYRON.     By  Hon.  RODEN  NOEL. 
LIFE  OF  HAWTHORNE.     By  MONCURE  D.  CONWAY. 
LIFE  OF  SCHOPENHAUER.    By  Professor  WALLACE. 
LIFE  OF  SHERIDAN.    By  LLOYD  SANDERS. 
LIFE  OF  THACKERAY.     By  HERMAN  MERIVALE  and  FRANK  T. 

MARZIALS. 

LIFE  OF  CERVANTES.    By  H.  E.  WATTS. 
LIFE  OF  VOLTAIRE.     By  FRANCIS  ESPINASSE. 
LIFE  OF  LEIGH  HUNT.     By  COSMO  MONKHOUSE. 
LIFE  OF  WHITTIER.     By  W.  J    LINTON. 
LIFE  OF  RENAN.    By  FRANCIS  ESPINASSE. 
LIFE  OF  THOREAU.    By  H.  S.  SALT. 


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Ibsen's  Prose  Dramas 

^EDITED  BY  WILLIAM  ARCHER 

Complete  in  Five  Vols.     Crown  Svo,  Cloth,  Price  3*.  6d.  each. 

Set  of  Five  Vols.,  in  Case,  17 s.  6d.  ;  in  Half 'Morocco \ 

in  Case,  325.  6d. 

'  We  stem  at  last  to  be  shown  men  and  women  as  they  are  ;  and  at  first 
it  is  more  than  we  can  endure.  .  .  .  All  Ibsen* s  characters  speak  and  act 
as  if  they  were  hypnotised,  and  under  their  creator's  imperious  demand 
to  reveal  themselves.  There  never  -was  such  a  mirror  held  ttp  to  nature 
before;  it  is  too  terrible.  .  .  .  Yet  we  must  return  to  Ibsen,  with  his 
remorseless  surgery,  his  remorseless  electric-light,  until  we,  too,  have 
frown  strong  and  learned  to  face  the  naked— if  necessary,  the  flayed  and 
bleeding — reality' — SPEAKER  (London). 

VOL.  I.  'A  DOLL'S  HOUSE,'  'THE  LEAGUE 
OF  YOUTH,'  and  'THE  PILLARS  OF  SOCIETY.' 
With  Portrait  of  the  Author,  and  Biographical  Introduc- 
tion by  WILLIAM  ARCHER. 

VOL.  II.  'GHOSTS,'  'AN  ENEMY  OF  THE 
PEOPLE,'  and  'THE  WILD  DUCK.'  With  an  Intro- 
ductory  Note. 

VOL.  III.  'LADY  INGER  OF  OSTRAT,'   'THE 
•     VIKINGS    AT    HELGELAND,'   'THE    PRETEND- 
ERS.'   With  an  Introductory  Note  and  Portrait  of  Ibsen. 

VOL.  IV.  '  EMPEROR  AND  GALILEAN.'  With 
an  Introductory  Note  by  WILLIAM  ARCHER. 

VOL.  V.  '  ROSMERSHOLM,'  'THE  LADY 
FROM  THE  SEA,'  'HEDDA  GABLER.'  Translated 
by  WILLIAM  ARCHER.  With  an  Introductory  Note. 

The  sequence  of  the  plays  in  eachvolume  is  chronological ;  the  complete 
«et  of  volumes  comprising  the  dramas  presents  them  in  chronological  order. 


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Library  of  Humour 

Cloth  Elegant,  Large  Crown  8vo,  Price  y.  6d.  per  Vol. 

'  The  books  are  delightful  in  every  way,  and  are  notable  for  the  high 
ttandard  of  taste  and  the  excellent  judgment  that  characterise  their 
editing,  as  well  as  for  the  brilliancy  of  the  literature  that  they  contain.' 
—BOSTON  (U.S. A)  GAZETTE. 


VOLUMES  ALREADY  ISSUED. 

THE  HUMOUR  OF  FRANCE.  Translated,  with  an 
Introduction  and  Notes,  by  ELIZABETH  LEE.  With 
numerous  Illustrations  by  PAUL  FRENZENY. 

THE  HUMOUR  OF  GERMANY.  Translated,  with 
an  Introduction  and  Notes,  by  HANS  MULLER-CASENOV. 
With  numerous  Illustrations  by  C.  E.  BROCK. 

THE  HUMOUR  OF  ITALY.  Translated,  with  an 
Introduction  and  Notes,  by  A.  WERNER.  With  50  Illus- 
trations and  a  Frontispiece  by  ARTURO  FALDI. 

THE  HUMOUR  OF  AMERICA.  Selected  with  a 
copious  Biographical  Index  of  American  Humorists,  by 
JAMES  BARR. 

THE  HUMOUR  OF  HOLLAND.  Translated,  with 
an  Introduction  and  Notes,  by  A.  WERN-ER.  With 
numerous  Illustrations  by  DUDLEY  HARDY. 

THE  HUMOUR  OF  IRELAND.  Selected  by  D.  J. 
O'DONOGHUE.  With  numerous  Illustrations  by  OLIVER 
PAQUE. 

THE  HUMOUR  OF  SPAIN.  Translated,  with  an 
Introduction  and  Notes,  by  SUSETTE  M.  TAYLOR.  With 
numerous  Illustrations  by  H.  R.  MILLAR. 

THE  HUMOUR  OF  RUSSIA.  Translated,  with 
Notes,  by  E.  L.  BOOLE,  and  an  Introduction  by  STEPNIAK. 
With  50  Illustrations  by  PAUL  FRENZENY. 

THE  HUMOUR  OF  JAPAN.  Translated,  with  an 
Introduction  by  A.  M.  With  Illustrations  by  GEORGE 
BIGOT  (from  drawings  made  in  Japan).  [In  preparation* 


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The  contents  of  these  little  handbooks  are  so  arranged  as  to 
permit  direct  and  immediate  reference.  All  dialogues  or  enquiries  not 
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EVERY-DAY  HELP  SERIES 

OF  USEFUL  HAND-BOOKS.    Price  6d.  each, 

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Contributors— J.  LANGDON  DOWN,  M.D.,  F.R.C.P.;  HENRY 
POWER,  M.B.,  F.R.C.S. ;  J.  MORTIMER -GRANVILLE,  M.D.; 
J.  CRICHTON  BROWNE,  M.D.,  LL.D.;  ROBERT  FARQUHARSON, 
M.D.  Edin.;  W.  S.  GREENFIELD,  M.D.,  F.R.C.P.;  and  others. 

The  Secret  of  a  Clear  Head. 

Common  Mind  Troubles. 

The  Secret  of  a  Good  Memory. 

The  Heart  and  its  Function. 

Personal  Appearances  in  Health  and  Disease. 

The  House  and  its  Surroundings. 

Alcohol:  Its  Use  and  Abuse. 

Exercise  and  Training. 

Baths  and  Bathing. 

Health  in  Schools. 

The  Skin  and  its  Troubles. 

How  to  make  the  Best  of  Life. 

Nerves  and  Nerve-Troubles. 

The  Sight,  and  How  to  Preserve  it. 

Premature  Death:  Its  Promotion  and  Prevention. 

Change,  as  a  Mental   Restorative. 

Youth:  Its  Care  and  Culture. 

The  Gentle  Art  of  Nursing  the  Sick. 

The  Care  of  Infants  and  Young  Children. 

Invalid  Feeding,  with  Hints  on  Diet. 

Every-day  Ailments,  and  How  to  Treat  Them. 

Thrifty  Housekeeping. 
Home  Cooking. 
Flowers  and  Flower  Culture. 
HOW  to  do  Business.     A  Guide  to  Success  in  Life. 
How  to  Behave.     Manual  of  Etiquette  and  Personal  Habits. 
How  to  Write.     A  Manual  of  Composition  and  Letter  Writing. 
How  to  Debate.     With  Hints  on  Public  Speaking. 
Don't :   Directions  for  avoiding  Common  Errors  of  Speech. 
The  Parental  Don't :  Warnings  to  Parents. 
Why  Smoke  and  Drink.    By  James  Parton. 
Elocution.    By  T.  R.  W.  Pearson,  M.A.,  of  St.  Catharine's  College, 
Cambridge,  and  F.  W.  Waithman,  Lecturers  on  Elocution. 


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VOLUMES  ALREADY   ISSUED— 


1  ROMANCE  OF  KING  ARTHUR. 

2  THOREAU'S  WALDEN. 

3  TKOREAU'S  "  WEEK." 

4  THOREAU'S  ESSAYS. 

5  ENGLISH  OPIUM-EATER. 

6  LANDOR'S  CONVERSATIONS. 

7  PLUTARCH'S  LIVES. 

8  RELIGIO  MEDICI,  &c. 

9  SHELLEY'S  LETTERS. 

10  PROSE  WRITINGS  OF  SWIFT. 

11  MY  STUDY  WINDOWS. 

12  THE  ENGLISH  POETS. 

13  THE  DIGLOW  PAPERS. 

14  GREAT  ENGLISH  PAINTERS. 

15  LORD  BYRON'S  LETTERS. 

16  ESSAYS  BY  LEIGH  HUNT. 

17  LONGFELLOW'S  PROSE. 

18  GREAT  MUSICAL  COMPOSERS. 

19  MARCUS  AURELIUS. 

20  TEACHING  OF  EPICTETUS. 

21  SENECA'S  MORALS. 

22  SPECIMEN  DAYS  IN  AMERICA, 

23  DEMOCRATIC  VISTAS. 

24  WHITE'S  SELBORNE. 

25  DEFOE'S  SINGLETON. 

26  MAZZINI'S  ESSAYS. 

27  PROSE  WRITINGS  OF  HEINE. 
23  REYNOLDS'  DISCOURSES. 

29  PAPERS    OF    STEELE    AND 

ADDISON. 

30  BURNS'S  LETTERS. 

31  VOLSUNGA  SAGA. 

32  SARTOR  RESARTUS. 

33  WRITINGS  OF  EMERSON. 

34  LIFE  OF  LORD  HERBERT. 

35  ENGLISH  PROSE. 

36  IBSEN'S  PILLARS  OF  SOCIETY. 

37  IRISH  FAIRY  AND  FOLK  TALES. 
33  ESSAYS  OF  DR.  JOHNSON. 

39  ESSAYS  OF  WILLIAM  HAZLITT. 

40  LANDOR'S  PENTAMERON,  &c. 

41  POE'S  TALES  AND  ESSAYS. 

42  VICAR  OF  WAKEFIELD. 

43  POLITICAL  ORATIONS. 

44  AUTOCRAT    OF    THE    BREAK- 

FAST-TABLE. 

45  POET  AT  THE  BREAKFAST-TABLE 

46  PROFESSOR    AT    THE    BREAK- 

FAST-TABLE. 

47  CHESTERFIELD'S  LETTERS. 
43  STORIES  FROM  CARLETON. 

49  JANE  EYRE. 

50  ELIZABETHAN  ENGLAND. 

51  WRITINGS  OF  THOMAS  DAVIS 

52  S PENCE'S  ANECDOTES. 


53  MORE'S  UTOPIA. 

54  SADI'S  GULISTAN. 

55  ENGLISH  FAIRY  TALES. 

56  NORTHERN  STUDIES. 

57  FAMOUS  REVIEWS. 

58  ARISTOTLE'S  ETHICS. 

59  PERICLES  AND  ASPASIA. 

60  ANNALS  OF  TACITUS. 

61  ESSAYS  OF  ELLA, 

62  BALZAC. 

63  DE  MUSSET'S  COMEDIES. 

64  CORAL  REEFS. 

65  SHERIDAN'S  PLAYS. 

66  OUR  VILLAGE. 

67  MASTER  HUMPHREY'S  CLOCK 

68  TALES  FROM  WONDERLAND. 

69  JERROLD'S  ESSAYS. 

70  THE  RIGHTS  OF  WOMAN. 

71  "THE  ATHENIAN  ORACLE." 

72  ESSAYS  OF  SAINTE-BEUVE. 

73  SELECTIONS  FROM  PLATO. 

74  HEINE'S  TRAVEL  SKETCHES. 

75  MAID  OF  ORLEANS. 

76  SYDNEY  SMITH. 

77  THE  NEW  SPIRIT. 

78  MALORY'S      BOOK     OF     MAR- 

VELLOUS ADVENTURES. 

79  HELPS'  ESSAYS  &  APHORISMS. 

80  ESSAYS  OF  MONTAIGNE. 

81  THACKERAY'S  BARRY  LYNDON. 

82  SCHILLER'S  WILLIAM  TELL. 

83  CARLYLE'S  GERMAN  ESSAYS. 

84  LAMB'S  ESSAYS. 

85  WORDSWORTH'S  PROSE. 

86  LEOPARDI'S  DIALOGUES. 

87  THE  INSPECTOR-GENERAL. 

88  BACON'S  ESSAYS. 

89  PROSE  OF  MILTON. 

90  PLATO'S  REPUBLIC. 

91  PASSAGES  FROM  FROISSART. 

92  PROSE  OF  COLERIDGE. 

93  HEINE  IN  ART  AND  LETTERS. 

94  ESSAYS  OF  DE  QUINCEY. 

95  VASARI'S  LIVES    OF   ITALIAN 

PAINTERS. 

96  LESSING'S  LAOCOON. 

97  PLAYS  OF  MAETERLINCK. 

98  WALTON'S  COMPLETE  ANGLER. 

99  LESSING'S  NATHAN  THE  WISE. 

100  STUDIES  BY  REN  AN. 

101  MAXIMS  OF  GOETHE. 

102  SCHOPENHAUER. 

103  RENAN'S  LIFE  OF  JESUS. 

104  CONFESSIONS OPST. AUGUSTINE 

105  PRINCIPLES    OF    SUCCESS    IN 

LITERATURE. 


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SLEEP:  ITS  PHYSIOLOGY,  PATHOLOGY,  HYGIENE,  AND 
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THE  NATURAL  HISTORY  OF  DIGESTION.  By  A.  LOCK- 
HART  GILLESPIE,  M.D.,  F.R.C.P.  ED.,  F.R.S.  ED.  6s. 

DEGENERACY :  ITS  CAUSES,  SIGNS,  AND  RESULTS.  By  Prof. 
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London :  WALTER  SCOTT,  LIMITED,  Paternoster  Square. 


THE  CANTERBURY  POETS. 

EDITED  BY  WILLIAM  SHARP.    Cloth,  Cut  and  Uncut  Edges,  is. ;  Red  Roan, 

Gilt  Edges,  2s.  6d. ;  Pad.  Morocco,  Gilt  Edges,  53. 
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1  CHRISTIAN  YEAR 

2  COLERIDGE 

3  LONGFELLOW 

4  CAMPBELL 

5  SHELLEY 

6  WORDSWORTH 

7  BLAKE 

8  WHITTIER 

9  POE 

10  CHATTERTON 

11  BURNS.     Songs 

12  BURNS.     Poems 

13  MARLOWE 

14  KEATS 

15  HERBERT 

16  HUGO 

17  COWPER 

18  SHAKESPEARE'S  POEMS,  etc. 

19  EMERSON 

20  SONNETS  OF  THIS  CENTURY 

21  WHITMAN 

22  SCOTT.     Lady  of  the  Lake,  etc. 

23  SCOTT.     Marmion,  etc. 

24  PRAED 

25  HOGG 

26  GOLDSMITH 

27  LOVE  LETTERS,  etc. 

28  SPENSER 

29  CHILDREN  OF  THE  POETS 

30  JONSON 

31  BYRON.     Miscellaneous. 

32  BYRON.     Don  Juan. 

33  THE  SONNETS  OF  EUROPE 

34  RAMSAY 

35  DOBELL 

36  POPE 

37  HEINE 

38  BEAUMONT  &  FLETCHER 

39  BOWLES,  LAMB,  etc. 

40  SEA  MUSIC 

41  EARLY  ENGLISH  POETRY 

42  HERRICK 

43  BALLADES  AND  RONDEAUS 

44  IRISH  MINSTRELSY 

45  MILTON'S  PARADISE  LOST 

46  JACOBITE  BALLADS 

47  DAYS  OF  THE  YEAR 

48  AUSTRALIAN  BALLADS 

49  MOORE 

50  BORDER  BALLADS 


51  SONG-TIDE 

52  ODES  OF  HORACE 

53  OSSIAN 

54  FAIRY  MUSIC 

55  SOUTHEY 

56  CHAUCER 

57  GOLDEN  TREASURY 

58  POEMS  OF  WILD  LIFE 

59  PARADISE  REGAINED 

60  CRABBE 

61  DORA  GREENWELL 

62  FAUST 

63  AMERICAN  SONNETS 

64  LANDOR'S  POEMS 

65  GREEK  ANTHOLOGY 

66  HUNT  AND  HOOD 

67  HUMOROUS  POEMS 

68  LYTTON'S  PLAYS 

69  GREAT  ODES 

70  MEREDITH'S  POEMS 

71  IMITATION  OF  CHRIST 

72  UNCLE  TOBY  BIRTHDAY  BK 

73  PAINTER-POETS 

74  WOMEN  POETS 

75  LOVE  LYRICS 

76  AMERICAN  HUMOROUS  VERSE. 

77  MINOR  SCOTCH  LYRICS 

78  CAVALIER  LYRISTS 

79  GERMAN  BALLADS 

80  SONGS  OF  BERANGER 

81  RODEN  NOEL'S  POEMS 

82  SONGS  OF  FREEDOM 

83  CANADIAN  POEMS 

84  CONTEMPORARY  SCOTTISH  VERSB 

85  POEMS  OF  NATURE. 

86  CRADLE  SONGS. 

87  BALLADS  OF  SPORT. 

88  MATTHEW  ARNOLD. 

89  CLOUGH'S  BOTHIE. 

90  BROWNING'S  POEMS. 

Pippa  Passes,  etc.    Vol.  1. 

91  BROWNING'S  POEMS. 

A  Blot  in  the  'Scutcheon,  etc.    Vol.  2. 

92  BROWNING'S  POEMS. 

Dramatic  Lyrics.    Vol.  3. 

93  MACKAY'S  LOVER'S  MISSAL 

94  HENRY  KIRKE  WHITE 

95  LYRA  NICOTIANA 

96  AURORA  LEIGH. 


London:  WALTER  SCOTT,  LIMITED,  Paternoster  Square. 


The    Canterbury    Poets. 

IMPORTANT  ADDITIONS. 
WORKS    BY   ROBERT    BROWNING. 


VOL.  I. 

Pippa  Passes,  and  other  Poetic  Dramas, 

by   Robert   Browning.     With   an    Introductory   Note 
by  Frank  Rinder. 

VOL.  II. 

A   Blot    in    the    'Scutcheon,   and    other 

Poetic  Dramas,  by  Robert  Browning.     With  an  Intro- 
ductory Note  by  Frank  Rinder. 

VOL.  III. 

Dramatic    Romances    and    Lyrics ;    and 

Bordello,  by  Robert  Browning.     To  which  is  prefixed 
an  Appreciation  of  Browning  by  Miss  E.  DIXON. 

BINDINGS. 

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LONDON:  WALTER  SCOTT,  LTD.,  Paternoster  Square. 


The   World's    Great    Novels. 

Large  Crown  &vo,  Illustrated,  $s.  6d.  each. 
Uniform  with  the  New  Edition  of  "Anna  Karenina  " 

A  series  of  acknowledged  masterpieces  by  the  most  eminent  writers 
of  fiction. 


THE     COUNT     OF     MONTE -CRISTO.       By 

ALEXANDRE  DUMAS.  With  Sixteen  Full-page  Illustrations 
drawn  by  FRANK  T.  MERRILL,  and  over  noo  pages  of  letter- 
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THE  THREE  MUSKETEERS.    By  ALEXANDRE 

DUMAS.  With  Twelve  Full-page  Illustrations  by  T.  EYRE 
MACKLIN,  a  Photogravure  Frontispiece  Portrait  of  the  Author, 
and  over  600  pages  of  letterpress,  printed  from  large  clear  type. 

TWENTY    YEARS    AFTER.     By  ALEXANDRE 

DUMAS.  With  Sixteen  Full-page  Illustrations  by  FRANK  T. 
MERRILL,  and  800  pages  of  letterpress,  set  from  new  type. 

LES  MIS&RABLES.     By  VICTOR  HUGO.     With 

Eleven  Full-page  Illustrations,  and  1384  pages  of  letterpress. 

NOTRE     DAME.       By    VICTOR    HUGO.       With 

numerous  Illustrations. 

JANE  EYRE.     By  CHARLOTTE  BRONTE.     With 

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Tolstoy's  Great  Masterpiece.     New  Edition  of  Anna  Karenina. 

ANNA    KARENINA:     A    NOVEL.     By    COUNT 

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"  Other  novels  one  can  afford  to  leave  unread,  but  Anna  Karenina 
never;  it  stands  eternally  one  of  the  peaks  of  all  fiction." — Review 
of  Reviews. 

LONDON:  WALTER  SCOTT,  LTD.,  PATERNOSTER  SQUARE. 


L 


AN    INITIAL 

W.U.  BE  ASSESS^ 
THIS  BOOK  ON  TH1  .  DATE 


PENAUTV 
FOURTH 


LD  21-50m-8