Skip to main content

Full text of "The natural history of Juan Fernandez and Easter Island"

See other formats 


THE 


Natural  History 


Juan  Fernandez 


AND 


Easter  Island 


EDITED  BY 
CARL  SKOTTSBERG 


THE  NATURAL  HISTORY 
OF  JUAN  FERNANDEZ 
AND  EASTER  ISLAND 

EDITED   BY  DR.    CARLSKOTTSBERG 


VOL.  I 
GEOGRAPHY,  GEOLOGY, 
ORIGIN  OF  ISLAND  LIFE 


WITH   14  PLATES 


AWE 


UPPSALA   1920-1956 

ALMQVIST&WIKSELLS  BOKTRYCKERI  AB 


707782 


Table  of  Contents. 

1.  Skottsberg,   Carl.  Notes  on  a  visit  to  Easter  Island 3 

2.  Hagerman,  T.   H.   Beitrage  zur  Geologic  der  Juan  Fernandez-Inseln      ....  21 

3.  QuENSEL,   P.  Additional    Comments    on    the    Geology   of  the  Juan  Fernandez 

Islands     37 

4.  Skottsberg,   C.   A  Geographical  Sketch  of  the  Juan  Fernandez  Islands   ...  89 
5. Derivation  of  the  Flora  and  Fauna  of  Juan  Fernandez  and  Easter  Island  193 


The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.     Vol.  I. 


"\iica4a  "Ho  Orwo 


EASTER   ISLAND 
after  Ihe  map  of  fhe  Qrilean  H\'dr.  OHice 

-t:aooooo 

Ueiifhi  1/1  miners  abox-e  sea   leve? 
.  Obsrrvatiart  spaf 


310 


/^i&oVo^uH 


I.    Notes  on  a  visit  to  Easter  Island. 

♦ 

■■.    '  By 
CARL  SKOTTSBERG. 

With  14  plates,  i  map,  and  3  text  figures. 

While  working  on  the  Juan  Fernandez  Islands,  our  party  obtained  per- 
mission to  accompany  the  Chilean  corvette  » General  Baquedano»  on  her  cruise 
to  Easter  Island,  in  191 7.  A  short  and  very  popular  account  of  our  visit 
appears  in  my  book  »Till  Robinson-on  och  varldens  ande»  (1918).  Although 
the  purpose  of  our  survey  was  purely  biological,  no  scientist  visiting  the  famous 
island  can  help  taking  a  vivid  interest  in  the  archaeological  remains,  and  we 
occasionally  made  a  (ew  observations.  However,  I  have  refrained  from  writing 
anything  on  the  subject,  as  I  had  to  wait  for  the  publication  of  the  results 
obtained  by  Mr.  and  Mrs.  SCORESBY  RouTLEDGE  of  the  British  »Mana»  Ex- 
pedition. Last  year  Mrs.  RoUTLEDGE  published  a  most  interesting  account  of 
their  work  on  the  island  (The  Mystery  of  Easter  Island),  where  the  ancient 
monuments  of  all  kinds  are  amply  described  and  illustrated.  A  second  volume 
will  follow,  containing  the  detailed  descriptions  of  the  prehistoric  remains. 

I  willingly  admit  that  this  little  paper  will  appear  rather  unnecessary 
since  the  British  Expedition  has  explored  the  place  with  such  a  wonderful 
accuracy.  But  it  is  Mrs.  RoUTLEDGE's  excellent  narrative  which  has  induced 
me  to  collect  a  few  notes  and  to  use  them  as  a  basis  for  a  discussion  of  some 
interesting  points.  I  have  also  found  it  worth  while  to  add  a  number  of  my 
photographs,  which  may  be  of  some  value. 

The  »General  Baquedano»  sailed  from  Iquique  on  May  27th,  1917.  She 
carried  a  Government  commission  presided  over  by  Bishop  Rafael  Edwards, 
a  prominent  Chilean  ecclesiastic,  who  went  to  continue  his  studies  on  native 
conditions  and  to  distribute  a  large  amount  of  materials,  clothing  etc.  among 
the  members  of  the  little  island  colony.  Capitan  de  fragata  J.  T.  Merino  was 
in  command  of  the  vessel,  and  he  as  well  as  the  bishop  and  the  officers  of  the 
ship  did  all  in  their  power  to  assist  us  in  our  undertaking. 

After  a  rather  uneventful  cruise  our  vessel  anchored  in  La  Perouse  Bay 
on  June  15th,  and  the  same  day  we  made  our  first  excursion  along  the  north 
coast.  We  were  bound  for  Hanga  Roa  (Cook's  Bay)  but  were  detained  in  La 
Perouse    on    account    of  adverse  winds;  finally  we  resolved  to  cross  the  island 


,  t ARL    SKOTTSBERO 


(,n  liorseback,  and  arrived  at  Matavcri,  the  scat  of  tlie  farm  house,  on  the  19th. 
hi  the  meantime  u  e  liad  made  some  excursions  in  the  northeastern  part,  where 
Mt  Katiki  was  ascended.  At  Matavcri  we  were  cordially  received  by  Mr. 
ri:k(  V  i:i)MlNhS,  the  mana-er,  and  were  invited  to  to  take  up  our  quarters  in 
his  house.  ( juite  naturally,  the  natives  were  in  a  state  of  great  excitement 
ovc-r  the  arrival  of  the  vessel  with  their  much  beloved  bishop,  the  missionaries 
(two  Capuchin  l^rethrcn)  and  the  many  useful  articles  reported  to  be  onboard; 
and  conse(iuentl\-  the\-  wore  rather  unwilling  to  render  us  any  assistance.  It 
was  only  through  the  kind  intervention  of  the  bishop  that  we  were  able  to  get 
an  important  })ait  of  our  scientific  baggage,  which  had  been  left  on  the  beach 
at  La  Perouse,  transported  to  Mataveri.  From  our  headquarters  the  district 
round  Hanga  Koa  and  Ilanga  Piko  etc.  was  visited  and  several  trips  under- 
taken to  Kano  Kao  and  Orongo,  the  famous  stone  village.  Further,  our  work 
was  exlendend  to  Kano  Aroi  and  Mt.  Terevaka,  the  highest  mountain,  and 
also  to  tlie  south  coast  and  to  Rano  Raraku,  the  image  mountain. 

We  had  expected  to  remain  at  least  three  weeks  on  the  island,  and  greatly 
regretted  that  we  were  ordered  onboard  already  after  a  fortnight's  stay,  espe- 
cially as  my  ca{)acit\'  for  hard  work  had  become  reduced  on  account  of  illness. 
The   >Ha«iucdano»   left  ICaster  Island  on  July   1st. 


(IKNKK.M.    NOTF.S   ON    TIIK    GEOGKArilV    OF    EASTER    ISLAND 

Tlic  topographical  features  of  the  island  are  fairly  well  illustrated  on  the 
accom[)anying  map,  the  result  of  Chilean  Navy  surveys  of  later  years.  The 
position  of  the  observation  spot  in  Ilanga  Roa  is  given  as  Lat.  27°  08'  06"  S., 
Long.  10/  25'  54"  W.  Mrs.  Roitledge's  map  is  based  on  U.  S.  Hydro- 
graphic  ()ft'ice  chart  no.  i  1 19,  from  which  it  differs  in  the  position  of  some  of 
the  mountains  and  in  the  geographical  names.  There  are  certain  discrepancies 
between  this  map  and  the  Chilean  one.  and  the  difference  between  the  latter 
and  the  U  S.  chart  are  still  more  considerable,  especially  in  the  configuration 
of  tlic  northwestern  part  of  the  island.  There  has  been  some  confusion  in  the 
placing  of  the  names,  but  I  take  it  for  granted  that  all  the  names  used  by 
Mrs.   R.  art-  properly  s|)elt  and  righth'  placed. 

1  lie  island  is  known  to  be  wholly  volcanic.  There  are  no  signs  of  recent 
action,  save  for  a  couple  of  tcj)id  sj^rings  below  high  water  mark  reported  to, 
but  never  seen  l)\  us.  It  is  rather  curious  that  both  THOMSON  (Smiths.  Inst. 
Ann.  kcp.  iSS.^.  Washington  1S91)  and  Agassiz  (Mem.  Mus.  Comp.  Zool. 
(  ambridgc  33,  i<i(>9)  should  discuss  volcanic  eruptions  and  great  earthquakes 
as  a  possible  reason  for  the  destruction  of  the  megalithic  monuments  and  for 
the  (lisappearani  e  of  the  greater  part  of  the  population,  as  all  signs  of  recent 
catastrophes  are  entirely  absent.  On  the  other  hand,  not  a  few  craters  are 
well  preserved;  someiinies  thev  are  arranged  on  distinct  lines  suggesting  lines 
of  less  resistance  in  the  older,  more  or  less  horisontal  basaltic  beds,  which  form 
the  bulk  of  the  island.  1  lie  tufas  and  ashes  of  the  numerous  cones  present  a 
great  variation  ot  colour  contributing  to  lessen  the  monotony  of  the  scenery. 
The  attention  of  the  visitor  is  especially  drawn  to  the  three  great  A"^;/^.    Rano 


NOTES    ON    A    VISIT    TO    EASTER    ISLAND  5 

is  the  native  name  for  a  mountain  which  contains  water.  R.  Kao  forms  the 
broad  southwestern  promontory.  The  greatest  height  of  the  rim  was  found  by 
the  writer  to  be  316  m.,  as  the  result  of  three  aneroid  observations  (differences 
in  temperature  duly  considered)  at  different  occasions.  Agassiz  has  i  327  feet 
or  403  m.;  the  Chilean  map,  400  m.  The  crater  lake  measures,  according  to 
Cooke,  2085  feet  across  (634  m.);  this  figure  may  be  the  result  of  a  careful 
observation,  but  seems  too  small.  Its  surface  was  found  by  me  to  be  120  m. 
above  sea  level;  THOMSON  and  CooKE  (Smiths.  Inst.  Ann.  Rep.  1897,  Wash- 
ington 1899),  say,  respectively,  that  it  is  700  and  600  feet  below  the  crater 
rim,  thus  according  to  their  figures  for  the  latter  corresponding  to  a  height  of 
190  or  160  m.;  taking  316  m.  as  the  starting  point  we  get  106  or  134  m.  I 
am  sorry  that  we  were  unable  to  examine  the  thickness  of  the  peat  that 
covers  the  sheet  of  water  save  for  some  irregular  pools,  which  do  not  appear 
to  have  decreased  much  in  size  since  the  photographs  of  the  Albatross  Ex- 
pedition were  taken.  No  reliable  figure  for  the  depth  of  the  lake  has  been 
obtained;  according  to  CoOKE,  Mr.  Salmon  tried  to  sound,  but  at  a  depth 
of  300  feet  the  line  broke  without  having  reached  the  bottom.  I  need  not  tell 
that  according  to  the  belief  of  the  islanders,  the  pool  in  Rano  Kao  belongs  to 
the  category  of  famous  lakes  without  a  bottom. 

The  lake  is  partly  surrounded  by  stands  of  a  very  robust  buUrush,  an 
endemic  variety  of  the  widespread  Scirpus  riparius,  called  paschalis  by  Dr. 
KUKENTHAL. 

The  country  NE  of  Rano  Kao  is  hilly,  one  of  the  cones  being  known  as 
Punapau  (Plate  i),  the  seat  of  the  hat  quarry.  The  northwestern  corner  of  the 
island  is  occupied  by  the  highest  mountain,  the  Terevaka  (Plate  2),  non  seldom 
veiled  by  a  bank  of  clouds.  This  name  is  not  mentioned  in  Mrs.  RouTLEDGE's 
book,  where  the  entire  high  land  in  question  is  called  Rano  Aroi.  But  the 
latter  name  only  applies  to  the  crater  on  the  southeast  slope  of  the  mountain. 
On  the  Chilean  map  appears  Cerro  Terevaka,  separated  from  Rano  Aroi  (or 
Roi)  by  a  shallow  depression,  and  both  names  were  recognized  by  the  native 
Juan  Tepano,  who  accompanied  us  to  this  place.  The  top  of  Terevaka  did 
not  present  any  marked  signs  of  being  a  crater;  the  height  was  found  to  be 
530  m.,  which  I  believe  is  nearer  to  the  truth  that  the  figure  770  put  down  on 
the  Chilean  map.  I  had  expected  to  find  some  notable  difference  in  the  flora 
of  the  highland,  but  was  rather  disappointed.  The  cryptogams,  mosses  and 
lichens,  however,  played  a  much  greater  part  here  than  in  the  lowlands,  where 
they  are  of  a  very  slight  importance.  Rano  Aroi  is  a  very  modest  and  shallow 
rano  and  cannot  at  all  be  compared  with  the  grand  R.  Kao.  The  height  was 
found  to  be  425  m.  The  lake  is  overgrown  with  vegetation.  There  is  a  gap 
in  the  east  wall  through  which,  after  prolonged  rains,  the  water  flows  down 
to  another  pool,  which  empties  itself  into  a  long,  narrow  fissure,  crossed  by 
the  track  from  La  Perouse  Bay  to  Hanga  Roa.  This  fissure  does  not  seem  to 
have  been  eroded  by  water  but  suggests  a  volcanic  origin. 

The  land  along  the  north  coast,  from  the  hills  backing  Anakena  Cove  to 
Katiki,  is  a  rather  flat  basaltic  plain,  with  occasional  outcrops  of  hard  rock 
and  strewn  with  innumerable  sharp-edged  stones,  partly  hidden  by  the  coarse 
grass  and   making  walking  disagreeable,  more  so  for  a  person  in  a  state  of  ill- 


TAkL    SKOTISUKKG 


health.  The  slope  of  Katiki  (another  name  not  found  on  Mrs.  RouTLEDGE's 
map,  but  frecpienlly  used)  is  comparatively  gentle.  The  top  was  found  to  be 
412  m.  higli  (300  m.  on  the  Chilean  ma[)  must  be  wrong);  it  presents  a  rather 
striking  appearance,  forming  a  shallow  circular  basin,  perfectly  dry  and  with 
a  flat  bottom  5-6  m.  below  the  rim,  which  is  75—80  m.  across.  On  the  north 
slope  is  a  succession  of  three  cones,  of  which  the  northernmost  is  gradually 
eaten  away  through  tlie  action  of  the  sea.  The  one  nearest  Katiki,  Vaintu 
Rova,  is  of  a  light  yellowish  colour;  the  liight  is  310  m.  On  the  south  slope 
wc  came  across  a  deep  fissure,  containing  rain  water  and  surrounded  by  a  fine 
growth  of  ferns.  The  natives,  of  course,  all  know  this  rare  watering-place,  and 
I  guess  this  is  the  well  spoken  of  by  Captain  CooK  in  his  second  voyage: 
•  Towards  the  eastern  end  of  the  island  they  met  with  a  well  whose  water  was 
|)erfectly  fresh,  being  considerably  above  the  level  of  the  sea;  but  it  was  dirty, 
(Mving  to  the  filthiness  or  cleanliness  (call  it  what  you  will)  of  the  natives,  who 
never  go  to  drink  v.ithout  washing  themselves  all  over  as  soon  as  they  have 
ilone  .  .  .  (the  edition  in  Iweryman's  Library,  p.  163).  North  of  Vaintu  Rova 
stands  the  somewhat  lower  Tea-tea,  the  » white  mountain*. 

SW  of  Katiki  the  famous  image  mountain,  Rano  Raraku,  is  situated,  so 
ably  described  and  illustrated  by  Mrs.  ROUTLEDGE,  who  gained  an  intimate 
knowledge  of  this  unique  place.  It  is  shown  on  Plate  5.  Between  this  rano 
and  the  hills  east  of  llanga  Roa  there  is  an  extensive  plain,  only  broken  by 
a  few  higher  eminences. 

The  visitor,  even  if  he  be  not  a  geographer,  cannot  fail  to  notice  the 
absence  of  every  trace  of  valley  or  ravine  caused  by  the  action  of  running 
water.  It  is  almost  with  surprise  that,  one  learns  the  figure  for  rainfall,  i  218 
mm.,  the  average  of  8  years'  observations.  This  is,  indeed,  no  small  amount, 
surpassing  that  of  Juan  h\'rnandez,  where  erosion  has  modelled  the  entire  island 
into  a  system  of  deej)  valleys  and  sharp  ridges,  l^ut  in  Easter  Island  there  is 
no  stream,  no  brook;  only  in  the  crater  lakes  water  is  always  found.  The 
great  .scarcity  of  water  makes  the  high  development  of  the  ancient  culture 
(juitc  astonishing.  The  climate  is  warmer  in  Easter  Island  than  in  Juan  Fer- 
nandez, the  evaporation  undoubtedly  much  greater,  the  winds  at  least  equally 
tre<iucnt.  Anybody  will  note  the  rapid  disappearance  of  the  water;  after  a 
licavy  raiti,  the  ground  may  become  soaking  wet;  nevertheless,  some  hours 
later,  it  is  perfectly  dry,  the  result  of  the  combined  forces  of  the  burning  sun, 
the  strong  winds,  and  the  extreme  j)orosity  of  the  soil.  Occasionally,  water  is 
encountered  by  digging  deep  holes;  but  to  dig  through  the  hard  rocks  must 
have  l)ccn  too  difficult  a  matter  for  the  natives.  Subterraneous  streams  are 
reported,  and  arc,  of  course,  to  be  expected;  and  running  water  has  played  an 
important   })att  in  the  formation  of  the  numerous  caves  round  the  coast. 

1  iic  vegetation  is  extremely  poor;  if  one  comes  from  Juan  Fernandez, 
the  contrast  is  very  striking.  The  island  is  destitute  not  only  of  wood,  but  of 
trees,  cxci-pt  for  a  few  specimens  in  the  crater  of  Rano  Kao,  where  the  last 
stunted  liwarfs  of  the  famous  toromiro  (SopJiora  foroiiiiro)  still  linger,  in  com- 
pany with  mahute  ' Ihvnssnurt/a  ptipyrifcni),  hau-hau  (or  jau-jau,  Spanish  j), 
called    //////;/A/A/.-  l,v  l-t  I  MIS    I  believe   that  it  is    T.  scmitriloba\  and  \:\  [Cor- 


NOTES   ON    A    VISIT   TO    EASTER    ISLAND  7 

dyline  iennhialis  f:a).^  From  the  earliest  descriptions  is  seen  that  the  island 
was  never  wooded;  planted  Eucalypts,  Melia  azedarach  and  other  subtropical 
trees  do  pretty  well,  however. 

In  his  »Informe»  (Memorias  del  Ministerio  de  Relaciones  esteriores,  culto 
i  colonizacion,  Santiago  1892),  P.  P.  TORO  says:  »En  otro  tiempo  formo  (i.  e. 
the  toromiro)  sin  duda  bosques  pues  en  diversas  partes  de  la  isla  se  ven  todavia 
innumerables  i  tupidos  troncos  secos  de  dos  a  tres  metros  de  alto.  Parece  in- 
dudable  que  esos  bosques  naturales  han  desaparecido,  secandose  la  mayor  parte 
de  los  arboles  a  consecuencia  principalmente  de  la  introduccion  de  animales 
vacunos  i  ovejunos  que  han  quebrado  las  plantas  o  les  han  comido  la  corteza.» 

Thomson  and  especially  Cooke  also  speak  of  the  numerous  groups  of 
trees  of  small  dimensions:  »In  other  parts  of  the  island  may  be  seen,  in  places 
in  considerable  numbers,  a  hardwood  tree,  more  properly  bush  or  brush,  called 
by  the  natives  toromiro,  all  or  nearly  all  dead  and  decaying  by  reason  of 
being  stripped  of  their  bark  by  the  flocks  of  sheep  which  roam  at  will  all  over 
the  island*. 

When  Cook  visited  the  place  in  1774,  he  certainly  did  not  come  across 
anything  like  a  forest,  for  he  expresses  himself  thus:  »the  country  appeared 
quite  barren  and  without  wood».  Both  ROGGEVEEN  and  FoRSTER  (A  voyage 
round  the  world.  London  1777,  Vol.  I)  assert  that  there  was  nothing  like  a 
forest  on  the  island.  FoRSTER  gives  a  good  description  of  the  general  appear- 
ance of  the  place;  he  mentions  about  ten  species  of  plants,  among  them  paper 
mulberry,  Hibiscus  populneus  and  Mimosa:  whether  Hibiscus  still  occurs^  I 
cannot  tell,  but  the  Mimosa  is  certainly  Sophora;  this  was  the  largest  tree, 
and  a  very  small  one:  »there  was  n6t  a  tree  upon  the  island,  which  exceeded 
the  height  of  ten  feet».  In  some  places,  on  the  hills,  Sophora  seems  to  have 
formed  small  shrubberies.  FoRSTER  did  not  estimate  the  flora  to  comprise 
more  than  twenty  species,  including  the  cultivated  plants.  On  board  the  »Ba- 
quedano»,  Bishop  EDWARDS  showed  me  a  letter  from  Brother  EUGENIO  Eyraud, 
the  missionary  of  the  island,  to  his  Superior  General  in  Valparaiso,  dated 
December,  1864.  He  writes:  »la  vegetacion  toda  de  yerbas  y  arbustos,  fal- 
tando  los  arboles  y  plantas  elevadas».  It  is  possible  that  Brother  EUGENiO 
includes  the  toromiro  within  his  »arbustos».  They  cannot  have  been  of  any 
considerable  size,  or  he  would  not  have  pointed  out  that  trees  were  missing. 
A  toromiro  of  3  m.  decidedly  has  the  look  of  a  small  tree,  not  of  a  shrub, 
especially  in  a  place  where  there  are  no  larger  plants  to  compare  it  with. 
TORO  must  be  mistaken  when  he  believes  that  there  had  grown  forests  on  the 
island  not  long  before  his  arrival.  Mr.  CoOKE  does  not  tell  if  he  saw  the 
numerous  trunks  himself  or  if  he  was  only  told  of  their  existence. 

^  In  Anuario  Hidrogr.  de  la  Mar,  de  Chile  (30)  1916,  p.  55  C.  De  la  Maza  mentions  a 
small  indigenous  tree  which  he  calls  »tumahiti».  »La  madera  del  tumahiti  es  bastante  dura  y 
mas  o  menos  flexible.  Lo  utilizan  los  canacas  para  construir  yugos,  arados  etc.  Es  el  unico 
arbol  que  crece  libremente  en  toda  la  isla.»  The  fruit  is  described  as  oval-shaped,  yellow, 
I  cm.  long,  with  a  stone  within,  and  of  bitter  taste.  There  is,  however,  no  tree  growing  freely 
over  the  island.  The  description  of  the  fruit  suggests  Melia,  but  this  is  called  miotaijl  (vide 
F.  FuENTES,  Resefia  botanica  de  la  Isla  de  Pascua.  Inst.  Centr.  Meteor,  y  geofi's.  de  Chile. 
No.  4.  Santiago  191 3).  Tumahiti  sounds  like  a  corruption  for  te  mahute,  the  paper  mulberry, 
which  certainly  cannot  supply  materials  for  the  implements  mentioned. 


S  CARI.    SKOIISBEKG 

The  dwarf  trees  now  cxistiiiL;  are,  as  has  already  been  stated,  almost 
wholly  confined  to  the  crater  of  Rano  Kao  and  are  on  the  verge  of  extinction. 
They  ^row  not  far  from  the  lake,  where  the  steep  slope  is  covered  with  very 
larj^e  blocks,  which  prevent  the  sheep  from  reaching  them.  Mr.  EDMUNDS  told 
me  about  some  trees  which  grow  along  the  steep  bluff  of  the  eastern  headland; 
unfortunatls',  we  were  prevented  from  visiting  the  place,  what  I  hope  some 
future  naturalist  will  do. 

The  greatest  part  of  the  island  is  covered  with  grass,  widespread  species, 
either  Polynesian  or  introduced  from  the  old  world  via  Chile,  Tahiti  or  other 
places.  Occasionally  ferns  arc  found,  also  outside  \.\\c  x^no,  Microlcpia  strigosa 
being  the  most  common.  We  discovered  two  species  of  Ophioglossum,  to  which 
the  natives  attribute  medicinal  (jualities.  Among  the  cryptogams  are  some 
endemic  species.  A  detailed  account  of  the  Flora  will  appear  in  the  volume 
dedicated  to  the  Botany  of  the  expedition. 

The  terrestrial  Fauna  is  very  poor;  no  indigenous  mammals  or  land  birds 
are  known.  There  are  two  species  of  lizards,  both  of  wide  range.  The  insect 
fauna  is  remarkably  poor;  some  species  have  been  introduced  by  man,  e.  g. 
cockroaches  and  flies,  which  have  increased  enormously. 


ARCIIAKOI.OGICAL  REMAINS:   THE   AHU 

Tliese  structures,  the  burial-places  —  but  not  the  only  ones  —  of  the  is- 
landers, liave  been  called  » terraces*  by  most  authors.  Such  a  word  tells  us 
very  little,  while  the  word  ahu  is  a  proper  technical  term,  strongly  and  justly 
recommended  by  Mrs.  RoUTLEDCH-:  for  regular  use. 

There  are  several  kinds  of  ahu.  The  most  striking  is  the  image-ahu, 
which  carried  the  now  fallen  statues  of  stone.  We  know  that  all  the  images 
were  purposely  u{)set  as  a  result  of  internal  warfare.  During  our  staty  in  La 
Tcrouse  Hay  we  devoted  some  time  to  the  inspection  of  the  ahu.  One  not  far 
from  the  landing-{)lace,  close  to  the  beach,  was  said  to  be  one  of  the  best 
preserved,  although  not  one  of  the  largest  and  having  supported  one  statue 
only.  'Ihis  ahu  was  measured  and  described  (see  Plates  3-4  and  text  fig.  l). 
The  central  part  is  23,;  m.  long  and  protrudes  4  m.  in  front  of  the  wings;  as 
it  stands  on  somewhat  higher  ground,  it  rises  above  the  wings,  in  spite  of  the 
front  wall  being  rather  low  or,  about  1,35  m.  The  front  wall  has  a  foundation 
of  small  stones  and  flat  slabs,  followed  by  large,  very  well  wrought  blocks; 
one  of  these  was  2,3  m.  long,  o,y  high,  and  0,45  broad,  another  2,45  long  and 
I  in  high.  They  are  closely  (itted.  The  space  behind  this  wall  was  filled 
with  bouUkrs,  the  surface  paved  with  larger,  flat  stones,  making  a  level 
platform.  ^ 

I  he  wings  are  larger  than  the  centre,  the  cast  31,3,  the  west  32  m.,  giving 
as  total  length  of  the  ahu  86,7  m.  Their  front  wall  is  as  high  as  or  even 
higher  (about  2  m)  than  that  f)f  the  centre  and  is  similar  in  construction, 
without  being  so  neatly  finished.  The  central  part  of  the  ahu,  behind  the  plat- 
form, had  been  disturbed,  a  stone  wall  of  7,6  m.  .stretching  obliquely  along  the 
fallen    image.      Ihe    aim    sl()i)cs    gently    inwards.     This   .slope,  which  could  be 


NOTES    ON    A    VJSIT    TO    EASTER    ISLAND 


traced  inland  about  12  m.  (measured  from  the  front  wall),  is  divided  by  a  wall 
(not  visible  above  the  surface),  as  described  and  figured  by  Mrs.  R.  The  sea- 
ward part  has  a  surface  of  boulders  almost  free  from  vegetation,  while  behind 
the  dividing-line  it  looks  like  an  old  pavement  with  grass  between  the  stones. 
According  to  Mrs.  R.,  the  vaults  for  bones  are  found  in  this  part.  In  the  case 
in  question  an  open  vault  was  to  be  seen  in  the  seaward  part  of  the  west 
wing;  it  measured  1,9  X  0,7  m.  with  a  depth  of  i  m.  Two  slabs  had  covered 
it.  We  do  not  know  if  the  ahu  were  built  with  many  vaults,  or  if  accomoda- 
tions for  the  bones  of  the  deceased  were  arranged  on  each  occasion. 


I    '    '    ■    '   (K    M    1/   1(1   I   U  '!'<(/   ■. 


Fig.  I.     Diagrammatic  sketch  of  an  ahu  west  of  the  landing-place  in  La  Perouse  Bay. 
Scale  I  :  750.    s  fallen  image,  /i  hat,  7/1  stone- wall,  ^  vault. 

In  the  centre  of  the  front  platform  one  large  image  or  moai  had  stood 
(Plate  4).  It  had  been  brought  down  by  undermining  the  foundation  stone. 
This  moai  was  the  only  one  carefully  measured: 


Total  length   10,27  ni. 
Length  of  body  6,55  m. 

»  »    head  2,52  m. 

»  »    neck  1,2  m. 

Width  of  body  at  base  2,y  m. 
Thickness  of  body  at  base   1,6  m. 


Width  across  shoulders  3,2  m. 

»  »        head  2,6  m. 

»  »        neck  1,9  m. 

Circumference  round  shoulders  7,9  m. 

»  »       neck  5,2  m. 

Length  of  ear  2,4  m. 


Close  by,  the  hat  or  crown  rested  flat  upon  the  ground;  its  height  was 
1,85,  its  circumference  7,  the  greatest  and  smallest  diameters  2,5  and  1,85  m., 
respectively.  It  is  of  the  usual  red  stone  from  Punapau,  but  not  finished.  The 
finished  hat  had  a  knob  on  top  and  an  oval  depression  below;  this  one  showed 
neither  on  the  exposed  side.  According  to  Mrs.  R.  the  hats  were  finished  after 
having  arrived  at  the  ahu;  I  suppose  this  one  never  adorned  the  image.  Could 
it  not  be  possible  that  the  stone  wall  spoken  of  above  was  part  of  a  construc- 
tion on  which  the  hat  was  to  be  rolled  up  to  the  top  of  the  image.? 

Naturally,  I  have  tried  to  identify  the  ahu  described  here  with  one  of 
those  mentioned  by  Mrs.  R.  Its  position  at  once  suggested  the  Faro,  of  which 
a  sketch  is  communicated  by  Mrs.  R.  As  far  as  I  can  see,  this  must  be  the 
same  (if  so,  it  will  undoubtedly  be  recognized  by  Mrs.  R.  from  my  photographs), 
but  there  are  some  differences  that  I  am  unable  to  account  for.  In  the  sketch, 
there  is  no  trace  of  the  stone  wall  alongside  the  image,  the  two  parts  of  this 
latter  are  far  too  much  apart,  and  the  hat  seems  to  rest  on  its  cylindrical 
side.     Then,    there  is    the    size    of  the    image:    Mrs.    R.    gives    32  feet,  adding 


CAKL    SKOTTSHKRG 


that  It  is  the  hir^cst  one  ever  found  on  an  ahu  and  the  last  one  to  be  upset. 
Are  there  two  exactly  siniihir  ahu  close  to  each  other,  each  with  an  unusually 
lari^e  imager  I  think  not.  The  measurements  were  taken  by  my  wife  and 
myself  with  a  tape  25   m.  loni;  and  the  figures  committed  to  paper  on  the  spot. 

It  couUl  po>sibly  also  be  the  same  ahu  as  no.  34  Punahoa  of  THOMSON 
1.  c,  p.  505.  lie  also  <^ives  the  total  length  of  the  single  moai  as  32  feet. 
Ilmvever,  the  entire  structure  is  said  to  have  a  length  of  175  feet  and  a  width 
«)f  S  feet,  which  figures  must  be  entirely  incorrect  if  ahu  Pare  is  meant. 

\W  what  kind  of  apparatus  or  devices  the  statues  were  transported  from 
Kano  Raraku  to  the  coast,  in  some  cases  to  rather  inaccessible  places,  remains 
a  m\ster\'.  The  natives  possessed  strong  cordage,  and  Mrs.  R.  has  made  out 
that  long  lines  were  used  occasionally,  but  veritable  hawsers  would  have  been 
needed  to  drag  the  statues  along  over  the  ground  in  the  manner  imagined  by 
Thomson.  We  have  seen  that  there  is  evidence  against  the  island  ever  having 
produced  good  sized  timber  suitable  for  rollers.  THOMSON  thinks  that,  after  a 
smooth  road  had  been  constructed,  »the  images  were  dragged  by  means  of 
ropes  made  of  indigenous  hemp»;  »seaweed  and  grass  made  excellent  lubri- 
cants*. He  could  »clearly  see  how  it  was  accomplished  with  a  large  force  of 
able-bodietl  men»  (p.  49S).  I  must  confess  that  I  find  it  less  easy  to  understand 
how  the  work  was  done,  for  the  least  obstacle  would  become  a  serious  one; 
and  the  roads  must  have  been  made  as  smooth  as  a  fioor  in  order  to  serve 
the  puri)ose,  the  images  being  rather  fragile.  Mrs.  R.  has  traced  the  few  high- 
wa\-s  leading  from  Rano  Raraku  to  the  coast;  but  if  really  the  images  were 
draggetl  up  to  the  numerous  ahu  all  round  the  island,  these  roads  cannot  have 
sufficed,  but  an  elaborate  network  of  very  smooth  paths  was  required,  of  which 
all  traces  would  have  disappeared.  It  is  true  that  seaweeds  are  plentiful,  but 
there  is  no  species  of  any  considerable  size  and  I  fail  to  see  how  the  quantities 
re(|nired  could  have  been  brought  together.  It  is  astonishing  that  no  tradition 
on  the  means  of  transport  survives.  According  to  Mrs.  R.  the  natives  in- 
variabl\-  offered  one  exi)lanation:  that  the  images  were  transported  by  the  aid 
of    supernatural   forces. 

<  )n  p.  4X6  Thonlsox  discusses  the  possibilit)^  of  a  transport  by  sea.  Near 
a  grouj)  of  ahu  he  discovered  a  fine  landing-place  made  by  art,  »admirably 
adapted  to  the  landing  of  heavy  weights*.  hVom  old  drawings  we  know  what 
the  aboriginal  canoes  were  like  —  not  a  single  one,  as  far  as  I  know,  has  been 
preserved  to  our  <la\s  but  they  were  not  strong  enough  to  support  any  very 
heavy  weight  (  )tie  might  suggest  that  large  rafts  were  built  but,  on  the  other 
hand,  there  are  several   ahu  which  are  unaccessible  from  the  shore. 

.Still,  there  is  another  nietod  to  be  reckoned  with,  although  further  specu- 
lation on  this  matter  may  appear  j)retty  useless.  Some  sort  of  a  sledge-like 
apparatus  could  h.i\c  been  cntistructed  without  the  need  of  timber  of  any  con- 
sitleral)le  si/.e.  .\  sledge  would  slide  (luite  well  over  the  grass,  provided  that 
the  road  was  cleared  from  stones.  A  great  number  of  people  could  be  simul- 
taneously engaged  in  j)n!liiig,  while,  if  rollers  were  used,  the  image  must  have 
been  more  diffK-ult  t(»  handle.  (  )nce  arrived  at  the  ahu,  a  sloping  causeway 
could  have  been  built,  from  which  the  image  was  lowered  down  in  position, 
or,  the  same  method  could  have  been  used  that  was  aj^plied  when  raising  the 


NOTES    ON    A    VISIT    TO    EASTER    ISLAND  II 

images  on  Rano  Raraku.  Still,  this  was  easier,  as  the  statue  was  steadied  by 
the  hole  in  the  ground.  For  details,  see  Mrs.  R.,  p.  189.  Also  compare  what 
is  said  above  on  ahu  Paro:  if  a  slope  was  built  for  the  image,  the  same  one 
might  have  been  used  for  bringing  the  crown  to  the  top,  and  perhaps  the  wall 
mentioned  above  will  have  to  be  explained  in  another  and  more  natural  manner 
than  the  one  indicated.  Anyhow,  to  'erect  the  statues  on  the  platform  in  the 
precise  position  required,  turning  their  backs  to  the  sea,  must  have  demanded 
not  only  great  skill  but  also  perseverance,  a  quality  not  very  characteristic  of 
the  present  native  population. 

It  is  the  merit  of  Mr.  and  Mrs.  R.  to  have  made  out  the  various  types  of 
ahu  and  to  have  pointed  out  that  not  a  few  have  been  rebuilt.  Some  structures 
of  this  kind  were  noted  by  us.  A  »poe-poe»,  not  far  from  the  western  slope 
of  Mt.  Katiki,  was  even  sketched  (another  by  Mrs.  R.,  fig.  95);  a  stone  pillar 
stands  on  the  surface,  which  is  covered  by  grass. 


THE   IMAGE   MOUNTAIN 

Rano  Raraku  has  been  called  the  most  interesting  spot  in  the  island. 
Truly,  the  sight  of  this  wonderful  mountain  with  its  quarries  and  statues  is  one 
not  likely  te  be  forgotten.  The  place  has  been  admirably  well  surveyed  by 
Mr.  and  Mrs.  R.,  making  it  quite  unnecessary  for  the  present  writer  to  say 
anything  on  the  subject,  but  he  thinks  that  the  reproduction  of  some  of  his 
photographs  will  be  found  pardonable.  Plate  5  gives  a  total  view  of  the 
mountain  from  the  SW,  plate  6  is  a  familiar  sight  from  the  outer  slope.  On 
plate  7  is  shown  one  of  the  very  largest  statues  in  the  quarries,  no.  41  in  Mrs. 
R.'s  diagram,  also  figured  by  her,  Fig.  49,  and  by  Agassiz,  Plate  39.  Neither 
of  these  shows  more  than  the  left  part  of  the  image.  It  must  be  about  16  m. 
long,  while  the  largest  statue  is  just  over  20.^  Mrs.  R.  finds  it  difficult  to 
believe  that  the  latter  was  ever  made  to  be  launched;  the  same,  then,  can  be 
said  of  no.  41.  It  is  hard  to  see  where  the  limit  for  the  capacity  of  the  an- 
cient islanders  should  be  drawn.  Surely  there  are  images  in  the  quarries  that 
are  little  more  than  rock:carvings,  but  I -am  not  prepared  to  include  the  two 
just  mentioned  under  such   a  heading. 

The  two  prostrate  statues  on  Plate  8  are  nos.  64 — 65  in  Mrs.  R.'s  dia- 
gram; they  show  the  narrowed  base.  Plate  9  is  also  from  the  inside  of  the 
crater,  a  quarry  high  up  in  the  gap.  It  represents  the  heads  of  two  images 
(possibly  nos.  15 — 16  of  the  diagram),  one  but  roughly  modelled,  one  finished. 
The  first  has  a  large  »wart»  on  its  cheek;  perhaps  this  applies  to  the  case 
mentioned  by  Mrs.  R.  on  p.  181,  where  it  is  stated  that  the  unexpected  occur- 
rence of  large  and  hard  nodules  in  the  rock  could  cause  the  whole  work  to 
be  abandoned. 

It  is  the  great  triumph  of  Mrs.  R.  to  have  unveiled  the  mystery  of  the 
scattered  statues  outside  the  mountain,  which  were  formerly  believed  to  have 
been    dropped    on   their  way  to  the  coast  and  left  lying.     We  know  now  that 

^  Geiseler  (resp.  Weisser)  describes  a  statue  of  23  m.  length  (Die  Oster-Insel.  Berlin 
1883,  p.  9). 


12  CAKI,    SK.OTTSEF.RG 

neither  the  iinaLTcs  stamlini;  on  the  mountain  nor  those  found  scattered  over 
the  ishuui  were  ever  intended  for  the  ahu,  but  that  the  latter  lined  the  roads 
leadin<;  from  the  mountain  to  the  coast.  I  cannot  add  anything  to  the  ex- 
phmation  of  the  standing  statues.  If  they  were  put  up  to  celebrate  »bird- 
nien».  it  seems  cjuaint  that  not  tiic  names  of  these  heroes  but  those  of  the 
workers  should  have  become  attached  to  them,  but  such  is  the  tradition,  as 
told  by  Mrs.   R. 

I'he  rutle  stone  implements  (toki)  used  by  the  sculptors  are  often  found. 
In  Ilanga  Koa  we  came  across  a  large  and  very  well  wrought  stone  adze  of  a 
ratiier  international  type,  but  not  found  in  any  of  the  accounts  on  the  island. 
It  had  been  picked  up  on  tlie  seashore  at  low  water  and  is  quite  incrustated 
witii  the  siiells  of  animals.  It  measures  20  cm.  (Plate  14,  fig.  i).  A  few  stone 
chisels  were  also  obtained;  two  are  figured  on  Plate   14,  fig.  2,  3. 


KKM.MNS   OV    llorSK.S    ANP    PLANTATIONS 

I''oundations  of  old  houses  are  seen  in  many  places,  and  several  were 
noteil  on  our  excursion  to  Alt.  Katiki.  Of  one  a  sketch  was  made,  also  showing 
the  paved  area  in  front,  but  without  foundation-stones  for  a  porch.  None  of 
these  dwelling-houses  are  left.  Between  Mataveri  and  Hanga  Roa  are  a  couple 
of  grass  huts  (Plate  10)  which  give  us  a  faint  idea  of  what  the  old  houses 
were  like.  The\-  are  small,  lack  the  stone  foundation,  and  have  the  entrance 
at  one  end. 

There  is  another  kind  of  structure  in  the  shape  of  low,  very  strongly 
built  towers  of  stone  which  cannot  fail  to  arouse  the  curiosity  of  the  visitor. 
The  present  pef)ple  do  not  seem  to  be  sure  as  to  their  former  use.  A  fine 
tower  at  the  landing-place  in  Hanga  Ho  Orno  (La  Perouse)  is  shown  on  Plate 
II,  another  is  figured  and  described  by  Mrs.  R.  (p.  218,  fig  87),  a  third  one, 
in  a  ruined  state,  by  Thomson  (p.  484).  The  first-mentioned  has  a  height  of 
3,.»  m.  and  is  6,;  7  m.  across  at  the  base.  The  only  entrance  is  0,65  m.  high 
and  0,9  wide.  The  roof  is  vaulted  inside;  outside,  the  wall  ends  in  a  girdle  of 
stones.  The  size  of  the  stones  bears  witness  of  the  prehistoric  era  and  of  the 
makers  of  the  j^reat  ahu.  Mrs.  R.,  on  the  authority  of  some  resident,  explains 
these  structures  as  look-out  towers  whence  watchers  on  land  communicated  the 
wlicreabouts  of  the  fish  to  those  at  sea;  these  contained  a  small  chamber 
below  which  was  us(-d  as  a  .sleeping  aj)artmcnt»   (p.  218). 

It  sounds  strange  that  these  solid  towers  should  have  had  no  other  pur- 
pose; tlian  to  serve  as  lookout  stations  for  fishermen.  In  order  to  keep  a  good 
look  out,  presumably  to  follow  the  movements  of  shoals  —  it  has  not  been 
proved  that  there  is  any  fish  here  of  the  social  type,  and  Mrs.  R.  states  that 
»tish  are  not  picntifuh  —  an  observer  must  seek  an  eminence  dominating  a 
considerable  space  of  water.  TiioMsox  tells  us  that  from  the  towers  the  move- 
ments of  the  turtles  wer(>  watched.  The  ob.server  must  keep  outside  the 
t(nver.  or  on  the  top  of  it,  not  a  very  comfortable  place.  Really,  the  tower 
itselt  would  have  been  little  more  than  a  refuge  in  bad  weather  and  during 
the    night,    but    for    such    a    jnupose    a    much    simpler  structure  would  indeed 


NOTES    ON    A    VISIT    TO    EASTER    ISLAND 


13 


suffice.  Thus  the  importance  of  the  tower  with  its  lower  apartment  appears 
to  stand  in  no  reasonable  proportion  to  the  vast  amount  of  labour  required  to 
build  it.  We  should  perhaps  remember  that  the  permanent  dwelling-houses 
were  much  more  fragile.  The  tower  suggests  some  kind  of  fortress,  with  a 
chamber  for  stores  or  treasures:  it  would  be  easy  to  defend  the  entrance. 
Speaking  of  the  narrow  entrances  to  the  Orongo  houses,  Thomson  remarks 
(p.  483):  »The  low  contracted  entrances  were  used  here  as  well  as  elsewhere 
for  defence.  Factional  fights  were  common,  and  it  was  necessary  that  every 
house  should  be  guarded  against  surprise  and  easily  defended ».  He  adds: 
» Another  reason  might  be  found  for  making  the  openings  as  small  as  possible, 
in  the  absence  of  doors  to  shut  out  the  storms».  But,  at  least  at  Orongo, 
there  were  plenty  of  slabs  suitable  for  doors  if  wanted. 


2.     Diagrams    of  old    plantations;  a  seen  from  above;  b  three  types  of  shelter,  in  section. 
I  =  bananas,  2  =  Melia  azedarach,  3  =  Andropogon  halepensis.    Scale  i  :  230. 


On  the  other  hand,  an  enemy  could  pull  down  the  roof  over  the  occupants. 
Also,  we  should  expect  to  find  a  communication  between  the  tower  and  the 
underground  chamber,  which  we  have  not  found.  So,  after  all,  this  theory  may 
not  hold  good.  Perhaps  prisoners  were  locked  up  in  the  towers,  where  they 
could  be  easily  guarded  till  the  hour  arrived  to  put  them  to  death.  But  it  is 
also  necessary  to  consider  whether  these  buildings  may  not  have  had  some 
relation  to  unkown  rites.  Did  they  have  a  ritual  purpose,  we  do  not  need  to 
wonder  about  their  elaborate  structure,  as  natives  may  invest  any  amount  of 
labour  in  connection  with  religious  or  other  ritual  buildings. 

There  is  a  description  and  figure  of  a  similar  building  in  La  Pi^ROUSE's 
Voyage,  reproduced  by  Stolfe  in  Ymer,  1883.  He  states  that  they  are  only 
found  on  the  top  of  Rano  Raraku,  which  is,  of  course,  a  mistake.  They  were 
oval  in  shape;  close  by  was  the  underground  chamber  with  its  separate  en- 
trance.    In  several  respects,  especially  concerning  the  ahu,  there  are  great  dis- 


14 


CAF^L    SKOTTSBERG 


crepancics  bctw  ccn  tlic  statements  and  illustrations  of  the  old  navigators  and 
the  results  obtained  by  modern  explorers.  A  critical  examination  of  the  old 
stories  would  be  welcome,  and  it  is  to  be  expected  that  Mrs.  RoUTLEDGE  will 
undertake  to  scrutinize  the  entire  literature.  There  are,  e.  g.,  in  La  PIiROUSE's 
and  I'lNAKl's  narratives  designs  of  ahu  which  do  not  at  all  correspond  to 
modern  descriptions  or  photographs. 

In  the  vicinit)-  ot  Ilanga  IIo  Orno  we  saw  many  remains  of  native  planta- 
tions. 'ihe\-  are  of  several  types.  One,  seen  in  fig.  2  a,  is  probably  of  a 
later    date,    as    the    material    has    been    taken    from    an  ahu,  the  front  wall  of 


l-ur    V    n.  Tsv.)   .iMrd-mcii)>  on  rock  at  Oron^o  (hciolu  ol  the  rock  1,6—1,7111.)  b.   Incised  marks 

on  door  poost  at  Orongo. 


which  fornix  the  back  wall  of  the  garden.  Circular  miniature  gardens  are 
represented  in  fig.  2  /'.  The  need  of  shelter  and  moisture  is  well  unterstood. 
Mclia  was  said   to  be  u.oun  for  the  sake  of  the  timber.    Probably  it  is  of  recent 


intii)(iuction. 


•>K'»N(;o    AND   WW.   lURI)   cui/r 


Mrs.  koriiKixiK  (i,-votcd  nuich  time  to  the  survev  of  the  Orongo  village, 
an(l  as  a  detailed  plan  was  made  and  every  hou.se  measured  and  described,  I 
shall   content  mysell    with   a   few  short  remarks. 

The  last  house  (if  I  remember  right)  towards  the  gap  of  the  crater  rim. 
close     to    the    sculptured    n.cks,    had    one  door-.post  with  incised  carvings  left. 


NOTES    ON    A    VISIT    TO    EASTER    ISLAND 


15 


vide  fig.  3  b.  This  house  was  pointed  out  to  us  as  the  house  of  Ariki.  The 
ariki  was  the  chief  of  the  Miru  clan,  the  authority  on  the  script  (i.  e.  the 
»ariki-mau»,  vide  R.  p.  241;  all  Miru  were  also  called  ariki).  Now,  the  same 
design  was  found  on  a  skull  in  the  possession  of  the  schoolmaster,  Mr.  I.  VlVES, 
and  this  skull  was  attributed  to  an  ariki.  Unfortunately,  the  owner  did  not 
want  to  part  with  his  treasure.  The  design  is  unlike  the  one  figured  by  Mrs. 
R.  (fig.  96)  of  another  Miru  skull. 

The  Bird  Cult  i?  described,  with  full  details,  by  Mrs.  R.  Special  attention 
is  paid  to  the  rock  carvings.  I  sketched  a  couple  of  the  »bird-men»  (fig.  3  a). 
Their  meaning  is  not  known  with  certainty.  Mrs.  R.  believes  them  either  to 
represent  one  of  the  egg-gods  (they  were  spoken  of  as  »Make-make»)  or  made 
to  immortalize  the  bird-men,  the  winners  of  the  egg-race;  she  finds  the  latter 
explanation  more  probable.  I  have  not  been  able  to  form  an  independent 
opinion.  The  same  carvings  are  seen  on  a  flat  stone  opposite  Orongo,  marking 
the  place  where  the  path  descends  into  the  crater  of  Rano  Kao. 

All  that  is  left  of  prehistoric  remains,  at  least  of  the  large  ones,  will 
remain  on  the  island.  Shortly  before  our  visit  a  law  was  passed  prohibiting  the 
removal  of  statues  etc.,  so  that  we  had  to  abandon  our  idea  of  bringing  home 
a  small  image  presented  to  us  by  one  of  the  residents.  The  »Mana»  was  just 
in  time  to  rescue  the  small  but  unique  statue  from  Motu  Nui. 


•     .         •  WOOD   CARVINGS 

The  famous  wooden  statues  as  well  as  other  pieces  of  carving  are  gone 
from  the  island  for  ever.  What  is  ofi"ered  to  passing  visitors  is  not  worth 
mentioning.  The  art  is  gone.  One  old  moai-miro,  in  a  very  much  decayed 
state,  had  been  discovered  in  a  cave  after  the  departure  of  the  »Mana».  It  was 
presented  to  Bishop  EDWARDS.  In  1908,  while  staying  at  Valparaiso,  a  Swedish 
captain,  Mr.  G.  Karstrom,  who  had  been  shipwrecked  on  Easter  Island  many 
years  before,  presented  me  with  two  beautiful  wooden  images,  one  of  which 
is  in  the  Etnographical  Museum  in  Stockholm;  the  other  is  owned  by  a  private 
person. 


HOUSEHOLD   GOODS,   WEAPONS,    ETC. 

Very  little  of  this  kind  is  now  to  be  encountered.  Sticks  used  for  net- 
knitting  are  available,  and  so  are  baskets  or  rather  bags  made  of  bullrushes 
(figured  by  THOMSON  on  Plate  51).  There  are  still  some  people  skilled  in  the 
preparation  of  tapa  cloth  from  the  mahute  and  of  strings  from  the  hau-hau, 
and  we  had  samples  made  for  the  collection.  Curiously  enough,  Geiseler 
does  not  mention  the  latter  plant,  but  states  that  all  the  cordage,  fishing-nets 
etc.  were  made  from  the  bullrushes. 

It  is  generally  stated  that  the  islanders  never  possessed  any  earthen-ware. 
Contrary    to  this,  RUTLAND  (Transactions  New  Zeal.   Inst.  29,   1896)  says  that 


,6  CARL  sKonsr^KKG 

the  earliest  liiscoverers  liad  seen  Mude  carthen-\vare»  on  the  island,  a  statement 
due  to  some  misinterpretation.  Of  stone  implements,  besides  the  toki,  and  the 
stone  adze  and  chisels,  we  t;x>t  one  lishiiook,  very  neatly  wrought  but  unfor- 
tunately not  complete,  as  the  point  is  missing  (Plate  14,  fig.  4)-  There  is  a 
drawing  of  one  of  these  hooks  in  Thomson's  report  (Plate  58).  Another  curious 
article  is  tlie  si)hcrical  stone  ball,  Fig.  5  on  Plate  14.  It  shows  two  holes 
which  communicate  so  that  a  string  can  be  passed  through,  and  may  have 
been  worn  as  an  ornament.  No  explanation  was  offered.  Perhaps  it  is  a 
fetish  stonc».  llioM.soN  has  described  and  figured  many  such  stones,  but 
none  of  them    i)resent  any  likeness  to  this  one. 

The  object  on  Plate  14,  Fig.  6  is  not,  as  might  be  suspected,  a  broken 
spear-liead  or  ))iataa,  but  has  been  given  its  present  shape  on  purpose.  It  fits 
well  into  the  hand  and  may  have  been  used  as  a  knife  or  scrape.  But  if  it  was 
used  with  a  handle,  my  explanation  may  not  be  satisfactory.  Spear-heads  are 
ct)mmonl\-  found  in  the  soil  and  also  manufactured  to  satisfy  the  demand  of 
visitors.  Two,  of  an  ordinar\'  type  and  apparently  old,  are  seen  on  Plate  14, 
h'igs.  7,  S.  According  to  THOMSON  there  were  at  least  nine  kinds,  all  with 
dilVerent  names,  a  statement  well  needing  the  corroboration  of  Mrs.  R. 


ORKIIN    OF    THE    PEOPLE 

The  histor\-  of  Faster  Island  is  full  of  mystery,  but  I  think  that  Mrs.  R. 
has  come  pretty  near  the  solution  of  some  of  the  problems.  She  has  drawn 
some  imj)ortant  conclusions  from  the  legendary  traditions  still  alive.  A  tale 
of  two  ditTerent  races  and  two  successive  colonizations  runs  through  the  old 
legends.  Tlie  anthropological  evidence  seems  to  be  in  favour  of  a  double  origin, 
Melancsian  and  I'oKnesian.  The  comparative  studies  of  the  Bird  Cult  in  the 
.Solomon  Islands  and  I^aster  Island  (by  PI.  Balfolr,  vide  Mrs.  R.)  seem  nothing 
less  than  convincing.  The  bird  rei)resented  in  the  numerous  carvings,  paintings 
etc.  of  ICaster  Island  is  not  the  holy  bird  of  this  place,  but  the  frigate  bird, 
u<»rshi|)|)cil  in  the  Solomon  Islands.  'Phe  bird  figures  were  called  penguins  by 
Lkhm.wn  (l-.ssai  dune  bibliograi)hie,  Anthropos,  1907),  which  undoubtedly 
must  be  a  mistake,  especiall)'  as  penguins  hardly  ever  visit  these  waters. 

it  wi-  sum  up  the  results  obtained,  there  is  evidence  that  the  Easter  Is- 
landers ate  ot  a  twofold  origin  and  that,  after  the  Melanesian  immigration,  a 
P(»l\-nesian  immigration  followed.  l"he  population  now  tends  to  assume  a 
multicoloured  aspect;  there  has  been  a  late  influence  from  Tahiti  (so  we  were 
told)  and  various  white  men  ha\'e  contributed  towards  the  »amelioration»  of  the 
race.  The  xouul;  girl  figured  on  Plates  12-13  ^^•'■'^  ^^^^  to  be  of  » pure  Easter 
Island  race  ,  but  whether  rejjrcsenting  a  IVlelanesian  or  Polynesian  type,  I  am 
unai)le   to  trli. 

A  critical  examination  of  the  language  would  be  of  interest.  Many  words 
are  the  same  as  in  the  Maori  or  other  Polynesian  tongues,  such  as  viaunga 
(mountain),  inaJiul,  (paper  mulberry),  //  (Cord^line),  ciimara  (sweet  potato)  a.  o. 
A    large    vocai)ular\-,  collected  by   Padre   Roi  SSEI,,  was  publi.shed  in  Santiago, 


NOTES    ON    A    VISIT   TO    EASTER    ISLAND 


17 


191 7,  but  it  miist  be  used  with  much  criticism,  as  it  contains  many  Tahitian 
words  and  also  corrupted  EngHsh,  French  or  Spanish;  good  examples  are  anio 
(agneau)  and  rnutone  =  sheep,  himene  (hymn)  =  to  sing,'  teperanate  =  serpent, 
tokini  =  stockings,  tiaporo  (diablo)  =  devil,  viretute  =  virtue,  given  without 
reservation.  A  closer  look  reveals  that  the  material  is  not  at  all  so  rich  as  the 
number  of  words  would  indicate,  for  the  author  has  invented  hundreds  of  ex- 
pressions for  ideas  wholly  unfamiliar  to  the  aboriginal  soul,  by  combining  the 
words  and  extending  their  meaning  in  a  most  improper  manner,  e.  g.  expres- 
sions for  cabin,  desert,  doctrine,  palace,  river,  saint,  W.  C.  etc.  etc.  to  quote 
a  few  obvious  examples,  of  which  scores  could  be  given.  This  is,  I  believe, 
a  common  missionary  method  to  enrich  the  language  with  ideas  and  expressions 
necessary  for  the  translation  of  reHgious  and  other  books,  but  otherwise  never 
used  by  the  natives. 

Concerning  the  name  of  the  sweet  potato,  see  below.  It  has  been  ad- 
vanced as  indicating  an  American  influence  previous  to  the  Columbian  era. 
Rutland  (1.  c.)  thinks  that  the  ancient  monuments  bear  witness  of  a  constant 
communication  between  the  island  and  Peru  and  Mexico:  »from  hence  architects 
of  Easter  Island  may  have   been  derived ». 

CULTIVATED   PLANTS 

If  we  knew  the  history  of  the  cultivated  plants,  many  a  mystery  related 
to  the  history  of  mankind  would  be  solved.  But,  unfortunately,  discussion 
often  begins  with  the  original  home  of  the  wild  parents  of  these  plants,  and 
there  it  also  ends. 

The  first  record  of  domesticated  plants  in  Easter  Island  is  that  of  RoG- 
GEVEEN,  the  discoverer  of  the  island  or,  at  least,  the  first  white  man  to  set 
his  foot  upon  it.  He  makes  the  following  statement  on  p.  120  (De  Reis  van 
Jacob  Roggeveen.  Worken  uitgeven  door  de  Linschoten-Vereenigung  4. 
191 1):  »en  toegebragt  worden  alles  wat  sy  hadden,  bestaende  en  boomvrugten, 
aardgewasch  en  hoenderen»,  that  is,  tree-fruits,  soil-fruits  (rootcrops)  and  hens; 
and,  farther  down:  »want  na  verloop  van  een  kleynen  tijd  bragten  sy  eene 
menigte  van  suykerriet,  hoenderen,  ubaswortelen  en  bananas»,  that  is  sugar- 
cane, bananas  and  ubas-roots.  But  what  is  ubas?  Most  likely  the  same  word 
as  the  Malesian  ubi  (uwi,  huvvi),  yams  (Dioscorea  alata),  now  called  ufi  in  the 
island.  All  these  plants  are  of  Old  World  origin  and  have  spread  from  the 
Indo-malayan  region  over  the  Pacific.  According  to  Friderici,  the  same  word, 
in  a  corrupted  form,  is  current  in  South  America:  »Dieses  Wort  schlagt  eine 
Briicke  iiber  den  grossen  Ozean:  es  gehort  als  op  unter  der  Bezeichnung  'siisse 
Kartoffel'  zum  Sprachschatz  der  Chimu,  des  kiistenbewohnenden  Kulturvolkes 
westlichen  Siidamerikas*  (Wiss.  Ergebn.  seiner  amtl.  Forschungsreise  nach  dem 
Bismarck  Archipel  im  Jahre   1908).    But  the  bridge  in  question  seems  to  be  weak. 

The  word  cumara  is  used  for  sweet  potato  [Ipomaea  batatas  or  Batatas 
edulis)  from  New  Zealand  through  Polynesia  to  Easter  Island.  According  to 
Cheeseman  (Manual  of  the  New  Zealand  Plora)  the  Maori  introduced  the  plant 
from  Polynesia  when  they  colonized  the  country  (supposingly  1350  — 1400),  and 
it    was    described  by  SOLANDER   as  Convolvulus  chrysorhizus,  now  reduced  to 

2  —  20199.     The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.     Vol.  I. 


I  8  CARL    SKOT'ISBI-RG 

a  synonym  of  I.  batatas.  This  plant  is  universally  considered  to  be  of  Central 
American  origin,  although  wild  plants  are  not  found  nowadays,  nor  is  the  more 
precise  locality  known  where  they  grew.  We  must  consider  whether  the  same 
species  was  not  originally  a  native  both  in  America  and  in  the  Polynesian  legion 
or,  whether  the  cultivated  forms  were  not  derived  from  more  than  one  wild 
species,  so  that  it  is  unnecessary  to  suppose  that  the  sweet  potato  was  intro- 
duced to  Polynesia  from  the  American  coast.  Yams  is  obtained  from  forms  of 
several  wild  species  characteristic  of  different  continents. 

If  this  theory  holds  good,  we  should  expect  to  find  different  names  for 
the  sweet  potato  on  the  two  sides  of  the  Pacific.  But  according  to  R.  Lenz 
(Diccionario  etimolojico.  Santiago  1910),  the  word  cumara  is  found  in  the 
Quichua  language;  it  is  not  indicated  as  the  principal  name  of  the  sweet  potato, 
which  is  apichii,  but  nevertheless  used,  according  to  this  author,  for  a  »clase 
parecida*  of  the  camote,  thus  for  some  form  of  the  same  plant.  From  this 
fact  some  people  would  conclude  that,  as  the  plant  is  American  and  called 
cumara  by  the  Quichua,  it  was  introduced  to  Polynesia  under  the  same  name 
long  before  the  Columbian  era.  It  is  useless  to  discuss  this  matter  any  further 
till  we  know  more  of  the  history  of  the  camote  and  also,  whether  the  word 
cumara  in  Quichua  really  applies  to  the  true  sweet  potato  and,  if  such  be  the 
case,  belongs  to  the  original  Quichua  language  or  has  been  introduced  through 
the  ICuropeans.  If  old  communications  existed  between  America  and  Polynesia, 
many  other  proofs  must  be  found.  Much  has  been  written  about  old  land- 
bridges  across  the  ocean,  considered  by  some  naturalists  to  be  indispensable 
for  the  explanation  of  the  distribution  of  animals  and  plants.  But  generally 
their  existence  was  supposed  to  have  ceased  long  before  the  age  of  Man.  Only 
H.XLI.IKR  (Uber  friihere  LandbriJcken,  Pflanzen-  und  Volkerwanderungen  zwi- 
schen  Australasien  und  Amerika.  Mededeel.'s  Rijks  Herb.  Leiden  13,  191 2) 
gives  them  a  longevity  sufficent  to  let  people  march  across.  I  am  afraid  that 
such  bridges  rest  on  a  very  unstable  foundation. 

To  return  to  the  sweet  potato,  we  have  seen  that  it  is  not  mentioned  by 
Ro<;gevkkn  as  existing  in  Easter  Island  in  1722.  CoOK  and  FoRSTER  found 
it  in  cultivation.  At  that  time  also  Broussonetia,  Thespesia  (also  Triumfetta?) 
and  toromiro  were  cultivated  in  addition  to  taro,  bananas  and  sugar-cane. 
.According  to  tradition  all  of  them  were  brought  by  Hotu  Matua's  party,  the 
first  settlers.  The  barahii  mentioned  by  F.  ViDAL  GoRMAZ,  Jeografia  nautica, 
p.  177  (Anuario  Midrogr.  de  la  Marina  de  Chile,  7)  is,  to  judge  from  the  de- 
scripton,  the  same  as  the  hau-hau.  The  calabash  mentioned  by  THOMSON, 
p.  535  is  Lagenaria  vulgaris.  At  present,  the  following  food-plants  are  cultiv- 
ated: sugar,  wheat,  Indian  corn,  taro,  pineapple,  yams,  bananas,  white  mul- 
berry, figs,  maniok,  oranges,  lemons,  grapes,  peaches,  quince,  plums,  beans, 
sweet  potatoes,  tomatoes,  melons,  artichokes  and  lettuce,  but  several  of  these 
only  on  a  very  small  scale  and  exclusively  in  the  garden  of  Mataveri.  Some 
tobacco  is  also  grown.  I  do  not  know  what  Thomson  means  by  the  »two 
varieties  of  indigenous  hemp»,  as  there  is  no  plant  of  this  kind  either  in  a 
cultivated  or  abandoned  state.  The  cordage  has  always  been  prepared  from 
the  hau-hau.  as  far  as  I  have  been  able  to  ascertain.  Nor  does  Mrs.  R.  refer 
to  any   such   plant,  nor  to  the  hau  hau. 


NOTES    ON    A    VISIT    TO    EASTER    ISLAND  1 9 

THE   FUTURE  OF   THE   ISLAND 

The  power  of  resistance  of  the  Easter  Island  people  was  definitely  crushed 
through  the  Peruvian  slave  raids,  and  through  missionaries  and  farmers  they 
lost  the  strength  which  lies  in  the  possession  of  an  aboriginal  culture.  Their 
removal  to  Hanga  Roa,  where  a  village  was  formed,  was  very  unlucky,  as  it 
meant  giving  up  many  small  plantations  and  induced  the  people  to  lead  a 
parasitic  life,  expecting  everything  from  their  new  rulers.  Although  they  have 
left  so  many  wonderful  monuments  to  bear  witness  of  earlier  busy  days  and 
of  a  people  of  warriors,  they  are  now,  with  few  exceptions,  lazy  beggars.  In 
part  this  may  be  due  to  their  pronounced  feelings  of  animosity  against  the 
intruders,  as  they  regard  themselves  as  the  true  possessors  of  the  island.  It 
appears  that  ever  since  the  establishment  of  a  farming  company  the  state  of 
affairs  has  never  been  lucky,  and  Mrs.  R.  has  an  interesting  tale  to  tell  of  an 
anxious  time.  I  do  not  at  all  believe  that  the  present  manager  is  to  blame, 
for  we  got  the  impression  that  he  is  as  well  liked  as  any  white  man  in  his 
position  can  expect  to  be.  In  Chile,  nobody  seems  to  have  taken  much  notice 
of  the  distant  colony  till  Bishop  Edw.ards  entered  the  field.  During  his  first 
visit,  in  1916,  he  informed  himself  of  the  state  of  things,  and  he  returned  in 
1917  invested  with  powers  to  put  everything  right  if  he  could.  Among  other 
things  he  wanted  to  take  up  war  against  the  leprosy.^  Not  quite  5  %  of  the 
population  suffer  from  this  disease;  they  are  confined  to  a  colony  some  distance 
from  Hanga  Roa.  Apparently  it  is  not  very  contagious,  for  the  isolation  is 
not  quite  effective.  The  surgeon  of  the  »Baquedano»,  Dr.  G.  LONGO,  examined 
almost  every  soul,  but  only  one  or  two  new  cases  were  discovered.  As  acco- 
modations for  the  most  advanced  cases  had  been  wanting,  the  vessel  this  time 
brought  materials  for  the  construction  of  a  small  hospital  which  was  to  be 
erected  by  the  new  »subdelegado»  or  governor.  Captain  MERINO  carried  in- 
structions to  examine  the  claims  against  the  company,  and  a  meeting  was  held 
where  the  natives  put  forth  their  demands.  I  understood  that  the  Company 
was  said  to  have  taken  possession  of  more  land  than  it  was  entitled  to  and 
that  the  natives  wanted  it  to  be  restored.  Officers  went  round  with  natives 
who  indicated  the  seats  of  their  former  homes  and  fields,  and  parts  of  the  land 
were  measured.  The  scheme  was,  I  think,  that  certain  parts  should  be  restored 
to  the  old  owners,  that  the  village  should  be  abandoned,  and  that  the  natives 
should  move  into  »the  camp»  in  order  to  become  selfsustaining.  A  certain 
amount  of  native  labour  should  be  granted  to  the  manager  at  a  fixed  rate  of 
pay.  I  have  had  ho  chance  to  learn  how  far  the  realisation  of  this  humanitary 
scheme  has  advanded;  nor  would  I  venture  to  foretell  if  it  is  likely  to  meet 
with  success. 

EASTER   ISLAND   AS   A    FIELD   OF   PSEUDOGEOGRAPHICAL  SPECULATION 

Finally,  I  shall    make  a  few  remarks  in  addition  to  what  Mrs.  R.  tells  us 
(p.  290)  of  the  theosophists'  views  of  Easter  Island,  which  are  based  on  errors 

^  The    surgeon  of  the  »Mohican»,  Dr.  Cooke,  does  not  mention  this  malady  as  existing 
in  the  island  in  the  year   1886.     It  was  imported  from  Tahiti. 


2  0  CARL    SKOTTSBEKG 

re^^ardini,^  the  cxistino^  monuments.  Last  year  a  small  book  appeared,  entitled 
»lJet  sunkne  kontinent  (Atlantis)»,  b\-  a  Xoruegian,  C.  SuND  (Copenhagen  I919), 
where  also  the  supposed  Pacific  continent  is  spoken  of.  As  might  be  expected, 
I'Lastcr  Island  forms  an  important  item.  With  my  permission,  two  of  my  photo- 
<,'raphs  were  reproduced.  No  doubt  Mr.  SUND  regards  himself  as  excused  for 
his  mistakes,  for  he  has  quoted  various  obscure  authors;  but  it  must  be  regretted 
that  lie  should  not  happen  to  draw  from  a  single  reliable  source,  not  even 
from  m\-  popular  dcscrif)ti()n,  which  was  known  to  him.  Mr.  SUND  tells  us  of 
the  Iv^^vptian  influence  in  Easter  Island,  of  the  enormous  foundation  walls  and 
ruins  of  temples;  almost  ever\-  mountain  had  sculptured  designs  of  goods,  fishes 
and  p\'ramids,  the  cave  paintings  w:ere  in  the  l^oltec  or  Egyptian  style,  etc. 
There  are  300  tablets  with  script  on  the  island  (if  it  were  but  true!),  waiting 
to  be  deciphered.  On  the  mountain  terraces  are  fortresses  with  walls  up  to 
80  feet  high.  The  pyramid  is  the  architectonical  principle,  built  as  the  Egj^ptiari 
one,  even  with  the  same  kind  of  cement.  All  materials,  bricks,  glass,  porcelain, 
ever>thing  was  known  in  I£aster  Island;  religion,  symbols  and  habits  were  the 
same  as  in  I\gypt,  only,  the  culture  of  the  island  was  older.  There  are  fan- 
tastic groups  of  statues  roundabout,  gods  of  hard  store  with  faces  up  to  25 
feet  high,  in  the  highlands  there  are  images  on  high  stone  pillars  or  staircase- 
like foundations,  and  with  square  hats  of  stone,  most  of  them  covered  with 
script  in  a  probably  forgotten  language.  Round  them  are  the  remains  of  large 
walls  and  buildings,  so  they  probably-  stood  in  vast  temple-yards.  And  so 
forth.  No  wonder  that  Mr.  SUNI)  draws  the  most  surprising  conclusions.  Now, 
this  must  not  be  taken  too  seriously  and  will  do  no  harm  in  scientific  circles. 
The  general  reader,  however,  will  get  a  rather  curious  idea  of  Easter  Island. 
I  dare  say  the  place  is  remarkable  enough  in  itself  and  need  not  be  glorified 
bv  such  fantastic  inventions. 


iMually,  I  wish  to  express  my  sincere  gratitude  for  kind  assistance  to  the 
Commander  and  Officers  of  the  »General  Baquedano»,  to  Bishop  RafaEL  Ed- 
wards, Mr.  I'KK(  V  I-:i)MiN])s,  Mr.  I.  ViVES  and  Baron  Erland  NoRDENSKlOLD. 


Explanation  of  Plate  14. 

1.  StiMic  ;u1/.:,  no.    U).    i.    307;   iu)t   quite   V-'- 

2.  .Stone  ciiiscl,  no.    10.   1.   320;  alino.st  ^i. 

V  ■  >^         no.    K).    I.    :;:i  ;  ,^       a,. 

4.     I'lsh  hook  of  stone,  no.   iq.   i     :,2y,  almost  "/;. 

V  Stone   1^11.  no.    10.    I.   500,  not   qu;?e   nat.  si/e. 

(\     Knite  or  scrape-  no.   19.   i.   31^,  not  ciuite  nat    si/e. 
7.     Speai-Iie.id,  no.    ly.    i.   31  -,.   •'  „ 

■'^-  "  .    UO.     lU.     I.     :;i  J  .    -j.:. 

'I'lie  orii,'inaK  in  the   .Museum.   Ciothenbiii". 


Nat.  Hist.  Juan  Fernandez  and  Easter  hi.    Vol.  I. 


Plate  i 


bo 


c 


o 

bC 
C 


.\'^//.    I  list.    '7iui)i   I'iDhVhIiz  tvni  luutir  Isl.     ]\>l.    I. 


Platk   2. 


<A 

^ 

o 

c3 

S 

^ 

c 

a 

8 

o 

^ 

(U 

^ 

c 

(L> 

c 

(U 

C/2 

(U 

hd" 

t 

c 

^ 

r^ 

2 

> 

o 

p^ 

■•n 

<\i 

c3 

fe 

biO 

(-] 

C 

■»-> 

g3 

O 

ffi 

C 

Nat.   Hist.  Juan  Fernandez  and  Easter  IsL    Vol.  I. 


Plate  3. 


(U 

TD 

bJO 

n 

c^ 

rri 

p 

c; 

c 

(U 

•-^ 

J3   ^ 


.\(//.    Hi  si.    '7im>/   l-ii)uvi(ii\:   cDui  l-.astd  I  si.     I'd.   I. 


Plate  4. 


m 


0 

c3 

03 

G 

4> 

•'"' 

^ 

c 

^ 

^ 

1^ 

^ 

'2, 

be 

*-i 

_•'-' 

o; 

'  > 

-C 

"^ 

cu 

0 

3 

H 

"cJ: 

13 

>-( 

OJ 

c3 

bJD 

> 

c3 

-^ 

_s 

-1^ 

(U 

rZZ 

rO 

Vi-( 

OJ 

0 

Ji 

^ 

0 

— < 

oj 

.s 

^ 

(U 

c/J 

^ 

03 

c 
0 

c3 

m 

a 

trt 

rt 

t-c 

C/2 

*>: 

Nat,  Hist.  Juan  Fernandez  and  Easter  Isl.    Vol.  I. 


Plate   5. 


.\(//.    llist.    '"fUiVi   l-t))iiuhii::   auJ  l.astiy   I  si.     \'ol.  I. 


Plate  6. 


/ 


m 


Nat.  Hist.  Jtian  Fcrnartdez  and  Easter  Isl.    Vol.  I. 


Plate  7, 


\<i/.    Hist.    '7ua)i   l\))iiviiii::  a)id  luistcr  Isl.     ]'ol.   I. 


Plate  8. 


Nat  Hist.  Jtian  Fernandez  and  Easter  hi.    Vol.  I. 


Plate  9, 


3    o 

0) 


5       rt 


la 


\(!/.   Hist,    'lua)!  l-\>)uvuii::   a)id  f.astir  I  si.    I'o/.  I. 


Plate   io. 


Nat.  Hist.  Juan  Fernandez  and  Easter  Is/.    ]^o/,  I. 


Plate   ii, 


Xd/.   Hist.    Juan  Foiitvidcz  ajid  liastcr  Is/.     J 'of.   I. 


Plate    12. 


Nat.  Hist.  Juan  Fcj'nandez  and  Easter  Isl.    Vol.  I. 


Plate   13. 


Photo  by  K.  B'dckstrbm 


Same  girl  as  in  Plate   12. 


.\(i/.    I  list.    '7iui//   J-\)iiii)!(/i-:  ivid  Juis/ir  Is/.     I'ol.    /. 


Pl.ATK    14. 


2.    Beitrage  zur  Geologic  der  Juan  Fernandez-Inseln. 

Von 

TOR  H.  HAGERMAN. 

Mit  12  Textfiguren. 

Die  Schwedische  Pazifik-Expedition  1916 — 1917  unter  der  Leitung  von 
Professor  C.  Skottsberg  brachte  unter  anderem  eine  Gesteinsammlung  von 
den  Juan  Fernandez-  und  Oster-Inseln  zuriick,  die  dem  Mineralogischen  Institut 
der  Hochschule  zu  Stockholm  zur  Bearbeitung  iibergeben  wurde.  Bei  der  von 
mir  vorgenommenen  Untersuchung  von  etwa  50  Handstiicken  von  den  Juan 
Fernandez-Inseln  stand  mir  ausserdem  Quensel's  Material  von  einer  friiheren 
Beschreibung^  desselben   geologischen  Gebietes  zur  Verfiigung. 

Die  Juan  Fernandez-Inseln  liegen  zwischen  33  und  34°  S.  Br.,  660  km  W. 
von  Valparaiso.  Die  Inselgruppe  besteht  aus  zwei  grosseren  Inseln,  Masatierra, 
Flacheninhalt  ca.  95  qkm  und  W.  von  derselben  Masafuera,  85  qkm.  Nahe  der 
erstgenannten  liegt  eine  kleine  Insel  St.  Clara,   5   qkm. 

Der  Gebirgsgrund  dieser  Inseln  besteht  ausschliesslich  aus  Effusivgesteinen. 
Eine  exakte  Altersbestimmung  derselben  kann  kaum  gemacht  werden,  da  keine 
Sedimentgesteine  vorhanden  sind.  QUENSEL^  nimmt  an  dass  die  vulkanischen 
Gesteine  kaum  alter  als  jungtertiar  sein  diirften,  und  es  ist  seitdem  nichts  be- 
kannt  geworden,  was  fiir  eine  veranderte  Auffassung  sprache. 

Die  Inselgruppe  ist  einer  kraftigen  Erosion  ausgesetzt  gewesen,  sodass  die 
urspriinglichen  Vulkankegel  nicht  mehr  zu  erkennen  sind.  Besonders  auf  Masa- 
fuera, wo  die  Wasserscheide  weit  nach  W.  verschoben  liegt,  hat  sich  eine  aus- 
gesprochene  Canon-Landschaft  gebildet.  Die  wilden  Terrainformen  sind  deutlich 
aus  den  zahlreichen  Photographien  zu  erkennen,  von  denen  viele  in  Skottsberg's 
Reisebeschreibung^,  wie  auch  in  Vol.  I  und  II  dieses  Werkes  veroffentlicht 
worden  sind. 

Masatierra. 

Die  Untersuchung  des  Materials  von  Masatierra  hat  erneut  bestatigt,  was 
Quensel  bereits  hervorhebt,  namlich,  dass  die  Gesteine  untereinander  chemisch 
und  mineralogisch  nahe  verwandt  sind,  und  sich  im  wesentlichen  nur  strukturell 
voneinander   unterscheiden.    Sie    konnen    als    verschiedene  Erstarrungsformen  ein 


1  P.  D.    Quensel,    Die    Geologie   der  Juan-Fernandez-Inseln.    Bull.  Geol.  Ups.,  Vol.  XI, 
p.  253-290. 

2  L.  c.  p.  256. 

3  C.  Skottsberg,  Till  Robinsonon  och  varldens  ande.  Stockholm  1918. 

3—248.      The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.      Vol.   I. 


2  2  TOR    H.     HAGERMAN 

unci  (lesselbcn  Magmas  betrachtet  werden;  spaterhin  sind  allenthalben  stellen- 
wcisc  sckundare  Veranderungen  infolge  hydrotliermaler  Prozesse  entstanden,  die 
abweichende  Ausbildunoen   hervorgerufen   haben. 

Die  Gesteine  sind  im  primaren  Zustande  durchwegs  mehr  oder  weniger  olivin- 
reiclie  Hasalte  mit  der  Zusammensetzung:  Olivin,  Pyroxen,  Plagioklas,  Magnetit, 
Ilnienit  und  oft  ein   wenig  Glasbasis. 

Charakteristisch  fur  alle  diese  verschicdenen  Teilen  der  Insel  entnommenen 
Proben  ist  besonders  die  Zusammensetzung  des  Pyroxens.  Dieser  besteht  aus 
einem  Titanaugit,  sofort  erkennbar  an  seiner  schwach  rotvioletten  Farbe  und 
starken  Dispersion  der  optischen  Aciisen.  13och  ist  zu  bemerken,  dass  derselbe 
nicht  zu  den  extremsten  Typen  gehort. 

Audi  die  Felds{nite  weisen  eine  konstante  Zusammensetzung  auf.  Sie  sind 
fast  aile  zwillingsgebildet  nach  dem  Albit-  oder  Karlsbader-Gesetz.  In  Schnitten 
senkrecht  zu  M  zeigen  die  Albitlamellen  eine  max.-Auslosung  von  32 — 33°,  in 
einigen  iMnzclfallen  diese  W'erte  mit  hochst  2°  variierend.  Der  Feldspat  ist  also 
ein   Labrador  von  eincr  Durchschnittzusammensetzung  Ab42An58. 

Da  die  Hasalte  also  mineralogiscli  einander  nahe  vervvandt  sind,  wurden  sie 
liauptsaclilich   nach  der  Struktur  in  folgende  Typen  eingeteilt: 

basaltische  Laven,  (teils  dichte,  teils  grobkornig  doleritische,  teils  schlackige) 
Tuftc,   hydrothermale   Um\vandlungS[)rodukte. 

Basaltische  Laven. 

Diese  (iesteine  bilden,  wie  friihere  Verfasser  bereits  betont  haben,  den  Haupt- 
bestandteil  der  Insel.  Auf  Grund  des  vorliegenden  Materials  konnte  man  den 
X'erlauf  der  verschicdenen  Lavastrome  und  deren  Neigungsverhaltnisse  nicht  be- 
stinimen   und  so  auch  keine  Klarheit  iiber  die  P>uptionsstellen  erlangen. 

Die  Fundstiitten  der  dem  Verfasser  zur  Untersuchung  vorliegenden  Hand- 
stucke  sind  ziemlich  gleichmassig  iiber  die  Insel  verteilt.  Von  den  zu  den  Laven 
gchorenden  (iesteinsproben  sind  ungefahr  ^/^  sehr  porose  und  schlackige  Typen, 
wahrend  ^3  (6  Stck.)  dichte,  dabei  gleichmassigere  und  feinkornige  Gesteine 
darstellen. 

Die  Struktur  dieser  Gesteine  ist  im  allgemeinen  hypokristallin  porphyrisch. 
In  einigen  I-'allen,  besonders  bci  den  dichten  Typen,  kommt  es  vor,  dass  Glas- 
basis ganz   fehlt. 

Mit  Bczug  auf  die  mincralogische  Zusammensetzung  der  basaltischen  Laven 
sind  folgende  Mineralien  beobachtet  worden :  Olivin  nebst  dessen  Umwandlungs- 
produkten,  die  vorgenannten  Pyroxene  und  Plagioklase  sowie  ILrzmineralien  (Mag- 
netit und   Ilmenit). 

Die  I'Linsprenglinge  sind  Olivine  und  ihre  Umwandlungsprodukte  sowie 
i'eldspat.  Die  Augitkorner  konnen  sich  zuweilen  der  Grosse  der  Einsprenglinge 
naiiern. 

Die  Olivineinsprenglinge  erreichen  ihre  grosste  durchschnittliche  Ausdehnung, 
ca.  2  mm,  in  Proben,  die  von  dem  nordlichen  Ufer  der  Padrebucht  stammen. 
Gerade  diese  ICinsprenglinge  zeigen  meistens  in  einer  scharf  begrenzten  Zone 
die  von  Oll.NSF.i.'  fruher  beschriebene  Iddingsitumvvandlung.  Aus  Dunnschlififen 
der    genannten    (icsteine  ist  klar  ersichtlich,   wie  die  Umwandlung  von  den   Ran- 

'  L.  c.  p.  260. 


BEITRAGE    ZUR    GEOLOGIE    DER   JUAN    FERNANDEZ- INSELN  23 

dern  nach  der  Mitte  zu  ausgegangen  ist  und  sich  0,02 — 0,03  mm  in  den  Olivin 
hineinerstreckt  hat.  An  manchen  Stellen  ist  die  Umwandlung  langs  der  Spalt- 
risse  vor  sich  gegangen,  wahrend  anderweitig  die  Durchgange  merkwiirdigerweise 
vollkommen  unveranderte  Teile  der  OHvinkerne  durchqueren.  (Vergl.  Fig.  i.) 
In  der  Gesamterscheinung  des  obengenannten  Praparates  mochte  der  Verfasser 
die  Iddingsitbildung  als  das  Resultat  einer  von  aussen  kommenden  chemischen 
Beeinflussung  ansehen.  WASHINGTON^  verweist  die  Iddingsitbildung  bis  auf  mag- 
matischen  Ursprung  zurijck.  Man  braucht  vielleicht  nicht  so  weit  zu  gehen,  da 
dieselbe  ebensogut  einer  hydrothermalen  Umwandlungsperiode  zugeschrieben  wer- 
den  kann. 


Fig.   I.     An    den    Randern   iddingsitumwandelte    Olivine.    Olivinbasalt  von   der    Padrebucht. — 

Vergr.  56  x  .    Photo  Hj.  Olsson. 

In  einem  anderen  Gestein  von  der  Padrebucht,  das  etvvas  feinkorniger  ent- 
wickelt  ist,  treten  auch  Feldspateinsprenghnge  von  0,5 — i  mm  Lange  auf,  neben 
Resten  von  Olivinen.  Der  Olivinumwandlungsprozess  hat  hier  ein  anderes  Pro- 
dukt  hervorgebracht,  namUch  gewohnHchen  Serpentin.  Dies  ist  auch  bei  den  der 
Grundmasse  angehorenden  OHvinkornern  der  Fall.  Dieselben  sind  vollkommen 
als  Serpentin  ausgeflossen.  Auch  der  Augit  scheint  an  den  Randern  etwas  ange- 
grififen  zu  sein.  Beinahe  identisch  entwickelt  sind  zwei  andere  feinkornige  Basalte, 
der  eine  von  der  Mitte  des  siidlichen  Ufers,  am  Fuss  des  Yunque,  der  andere  von 
einem  Gebirgsriicken  (385  m  ii.  M.)  SW.  von  Tres  Puntas,  W.  von  der  Villagra- 
Bucht,  herstammend.  Iddingsit-  und  Serpentinumwandlungen  der  Olivine  sind, 
wie  ich  besonders  hervorheben  mochte,  niemals  in  ein  und  demselben  Gestein 
gleichzeitig  angetroffen  worden. 

Die  beiden  erwahnten  Gesteine  von  der  Padrebucht  und  an  beiden  Seiten 
von  Villagra  zeigen  eine  eigentiimliche  primare  Struktur  der  gleichkornig  ent- 
wickelten   Grundmasse,  indem  die  Plagioklase  mit  einer  mittleren   Ausdehnung 


1  Italian  petrogr.  sketches.   Journ.  Geology.    4  (1896),  p.  835 — 836. 


24 


TOR    H.     HAGERMAN 


von  0,2  mm  sich  vielfach  radialstrahlig  mit  den  Augitindividuen  geordnet  haben. 
1st  man  erst  einmal  auf  diese  spharolitahnlichen  Bildungen  aufmerksam  gewor- 
den,  so  findet  man  sie  haufig  hauptsachlich  in  den  feinkornigsten  Proben  dieser 
Gesteine  ausgebildet.  Besonders  schone  Beispiele  hiervon  zeigt  ein  Handstiick, 
das  der  »ryramides  einem  Ikrggipfel  ungefahr  in  der  Mitte  der  Insel  unvveit 
des  SKIA'IKK-Denkmals,  entnommen  wurde.  (Fig.  2.) 

lune    sehr   ahnliche    Krscheinung    ist    von   Reiter^  beschrieben  worden.    Er 
sclimolz    45%    Albit,    45%   Augit    und    10%   Magnetit   zusammen.    »Der    Schliff 


V    /r 


Pyroxen      (-^^">PU<tioklAS       ^  M  Ag  Weht 


Fig.  2.     Phigioklas-AugitSpharoIit.  I5asalt  von  der  '>Pyramide».  —  Vergr.  250  x 

Zeichnung  vom  Vcrf. 


eincr  durcli  7  Stundcn  abgckiihlten  Schmelze  zeigt  eine  sphariodale  Anordnung 
der  Kristalle,  in  dem  niagnetitreiche  Kerne  von  Glaspartien  mit  einzelnen  aus- 
gcschiedenen  Augit-  und  Plagioklasleisten  und  Kristalliten  umgeben  sind.  Die 
W'iederhokmg  des  X'crsuchcs  bei  30-stundiger  Abkiihlung  ergab  eine  Schmelze 
mit  kornig-pori)hyrisclicr  Struktur.  Zweifelsohne  ist  dies  audi  in  dem  vorliegenden 
l^'alle  zutrcffeiid,  indcm  die  s{)harolitrulirenden  Laven  einer  raschen  und  ungestorten 
Abkuhlung  ausgcset/.t  gewesen   sein   durften. 

In  Anschluss  an  dicse  Spharolite  seien  hier  die  in  Pig.  3  abgebildeten  kreuz- 
formig  licgenden  Olivinkristalle  erwahnt.  Der  DiinnschlifT  entstammt  einem  etwas 
grobkornigen    Gestein    von    Bahia  Cumberland.    Wie  aus  der  Figur  deutlich   her- 

^   U.  H.  Rkii  F.R,  Kxporimentellc  Studien  an  Silikatschmelzen.  Neues  Jahrbuch.  Beil.  Bd.  22 
1906  ,  p.   197. 


BEITRAGE    ZUR    GEOLOGIE    DER  JUAN    FERNANDEZ-INSELN 


25 


vorgeht,  handelt  es  sich  urn  eine  skelettartige  Ausbildung  der  Kristallindividuen. 
Eine  gesetzmassige  Verwachsung  der  verschiedenen  Individuen  habe  ich  nicht 
nachweisen  konnen. 


Im  Vaqueriatal  tritt,  wie  aus  der  untenstehenden  Photographic  (Fig.  4)  er- 
sichtlich,  ein  fast  horizontal  liegendes  Gestein  auf.  Nach  ihrer  grobkristallinischen 
Struktur  zu  urteilen,  sind  diese  Basalte 
moglicherweise  als  intrusiv  aufzufassen> 
Das  nur  an  zwei  Seiten  zugeschlagene 
Handstiick  ist  vorziiglich  durch  Schrump- 
fung  unter  rechtem  Winkel  zerkluftet. 
(Fig.  5.)  U.  d.  M.  zeigt  dasselbe  ein  un- 
verandertes  hochkristallinisches  Aussehen. 
Reichlich  albitlamellierte  Feldspatleisten, 
durchschnittlich  ca.  2  mm  lang,  bedin- 
gen  mit  Olivin-  und  Titanaugitkristallen 
eine  ophitische  Struktur.  Ausser  diesen 
Mineralien  habe  ich  nebst  Magnetit  hier 
und  da  ein  Biotitkorn  gefunden.  Moglicher- 
weise erstreckt  sich  dieser  Basalt  bis  zur 
Cumberland  Bay,  wo  eine  ahnliche  Aus- 
bildung von  QUENSEL^  beschrieben  wurde. 
Auch  das  Material  Skottsberg's  enthalt 
eine  ahnliche  Probe  von  dort,  in  losem 
Block  gefunden.  Der  Mineralbestand  der 
beiden  letztgenannten  Handstiicke  ist  der- 
selbe  wie  jener  der  Vaqueriaprobe  nur 
mit  dem  Unterschied,  dass  kein  Biotit  vor- 
handen  ist.  Auf  Grund  seiner  grobkristallinischen  Struktur  muss  das  obenerwahnte 
Gestein  zu  den  doleritischen  Basalten  gerechnet  werden. 

Feinkorniger,  aber  im  iibrigen  dem  vorgenannten  Gestein  vollig  gleich,  ist 
der  bei  »Tres  Puntas»  genommene  Basalt.  Die  Handstiicke  bestehen  aus  langen, 
schmalen,  dreiseitigen  Prismen. 


Fig.  3  a. 


Zentrisch   angeordnete    Olivine. 
Vergr.  56  x  .     Verf.  phot. 


Unter  den  schlackigen  Laven  weisen  einige  eine  auffallende  Analogic  zu 
rezenter  Oberflachenbildung  auf.  Besonders  ist  dies  der  Fall  bei  einem  sehr  po- 
rosen  glasreichen  Gestein  vom  Ufer  s.  von  Yunque.  Der  vorerwahnte,  auf  dem 
Gipfel  der  »Pyramide»  befindliche  dichte  Basalt  hat  ein  schlackiges  und  glasiges 
Lavabett  als  Unterlage. 

In  einigen  anderen  der  schlackigen  Gesteine  sind  die  Locher  mehr  oder  we- 
niger  mit  Opal,  Chlorit,  Serpentin  und  Calcit  ausgefiillt. 


1  Vergl.  jedoch  Quensel  p.  263—264. 

2  L.  c.  p.  263. 


26 


TOR    H.    HAGKRMAN 


Tuffe. 

Unter  dem  mitgebracliten  Material  befinden  sich  zwei  Proben  von  ausge- 
sprochenen  Tufien.  Der  eine,  ein  poroses,  dichtes  Gestein,  stammt  von  El  Puente, 
dem  Istmus  zwischen  der  Padrebucht  und  Carbajal  und  ist  ein  Palagonittuff  mit 
einigen  sporadischen  Augit-  und  Magnetitkornern.  Der  andere  Tuff  stammt  von 
dem  nordlichen  Ufer  der  Padrebucht,  von  wo  einige  umgevvandelte  Olivinba- 
salte  (s.  S.  23)  herriihren.  Ausser  Augit  und  Magnetit  enthalt  derselbe  einige 
grossere  vollkommen  reine  Olivinkorner  in  einem  teilvveise  kryptokristallinisch  aus- 
sehenden    Glase.    Stellt    man    diese    verschiedenen    Bildungen    aus   der   Nahe   der 


Fi^.  3  b.     Detail  von   Fig.  3  a.—  Verg^r.   170  x.  Verf.  phot. 


Padrebucht  zusammen,  so  gelangt  man  zu  der  Auffassung,  dass  dieses  Gebiet  frische 
Spuren  vulkanischer  Tatigkcit  aufweist.  Vergleicht  man  die  obengenannten  Tuffe 
mit  den  von  OlKNSKl.'  bescliriebenen  roten  Tuffen  von  der  Cumberland  Bay,  so 
scheinen  die  letztgenannten  nicht  so  empfindlich  gegen  Verwitterung  zu  sein, 
wie  besonders  die  Palagonittuffe. 

Hydrothermale  Bildungen. 

K\n  aragonithaltiges  (iestein  vom  Ufer  gleich  siidlich  vom  Yunquegipfel  diirfte 
als  liydrothermal  umgewandelter  Basalt  angesehen  vverden.  (Fig.  6.)  Das  Hand- 
stuck  ist  ein  von  weissen  Streifen  durchzogenes  scharfgriines  Gestein,  das  u.  d.  M. 

1  L.  c.  p.  266. 


BEITRAGE    ZUR    GEOLOGIE    DER   JUAN    FERNANDEZ-INSELN 


27 


grosse  Augitkristalle  in  einer  vollig  zerflossenen  Serpentinmasse  zeigt.  Das  Pra- 
parat  ist  von  Aragonitbandern  durchzogen.  Dieses  Gestein  muss  als  ein  stark 
umgebildeter  Olivinfels  bezeichnet  werden.  QUENSEL^  hat  ganz  frische  Gesteine 
von  letztgenanntem  Typus  angetroffen  und  beschrieben.  Vielleicht  kann  die  Ara- 
gonitbildung  hier  eine  Andeutung  geben,  auf  welche  Weise  der  Olivin  chemisch 
umgewandelt  worden  ist. 

Auf  derselben  Stelle  wurde  auch  eine  reine  Kalksinterbildung  gefunden,  was 
darauf  hinweist,  dass  diese  Gegend  in  spaterer  Zeit  postvulkanischen  Prozessen 
hydrothermaler  Natur  ausgesetzt  gewesen  ist. 

In  Fig.  7  ist  eine  eigentumliche  Bildung  dargestellt,  wie  sie  auf  dem  offenen 


Fig.  4.    Das  Vaqueriatal.  Wasserfall  iiber  den  saulenformig  abgesonderten  doleritischen  Basalt. 

Plateau  bei  Puente  vorkommt.  Es  sind  lange,  in  einem  »Sandfeld»  aufrechtste- 
hende  rohrahnliche  Bildungen,  Aragonit,  Pyroxen,  Magnetit  sowie  etwas  Olivin 
in  einem  Zement  von  Karbonat  enthalten.  Die  wahrscheinlichste  Deutung  dieser 
Phanomene  ist  wohl,  dass  mit  Calciumkarbonat  gesattigte  thermale  Gewasser  iiber 
eine  Vegetationsdecke  geflossen  sind,  wobei  Wurzeln  etc.  mit  einer  Kruste  von 
oben  angegebener  Zusammensetzung  iiberzogen  wurden. 


Santa  Clara. 

An  dem  Siidende  von  Masatierra  liegt  die  kleine  Insel  Santa  Clara.  Von 
Santa  Clara  selbst  ist  keine  Probe  mitgebracht,  dagegen  von  der  kleinen  Insel 
Morro  de  los  alelies,  die  bei  tiefstem  Wasserstand  mit  der  Hauptinsel  zusammen- 


1  L.  c.  p.  265. 


28 


TOR    H.    HAGERMAN 


hangt.  Sowohl  in  Diinnschlifif  wie  in  Handstiick 
zeigt  dieses  Gestein,  das  als  fast  vertikale  Gange 
auftritt,  eine  voUige  tjbereinstimmung  mit  einem 
der  dichten  Basalte  von  Masatierra  vom  Gipfel  des 
Cerro  Negro  SO  von  Yunque,  190  m  ii.  d.  M.  Ur- 
spriingliche  Plagioklaseinsprenglinge  von  bis  zu  2 
mm  Lange  sind  oft  so  stark  kaolinisiert,  dass  beim 
Schleifen  nur  die  Hohlraume  iibrig  geblieben  sind 
und  dem  Gestein  ein  falsches,  schlackiges  Aus- 
sehen  verleihen.  Hierbei  ist  interessant,  dass  der 
Feldspat  der  Grundmasse  sich  frisch  beibehalten 
hat.  Die  Olivine  sind  natiirlich  vollig  in  Serpentin 
umgewandelt.  Das  ganze  Praparat  ist  von  Ilme- 
nitskeletten  durchwachsen. 

Masafuera. 


I'ig.  5.     Handstiick  voin  Vaqueria 
K'ang.    Verf.  phot. 


Etwa  180  km  vvestlich  von  Masatierra  erhebt 
sich  die  Insel  Masafuera.  Abweichend  von  Masa- 
tierra in  Bezug  auf  die  einheitliche  Mineralzusammensetzung  der  Gesteine  Uefert 
Masafuera  Beispiele  petrographisch  weit  verschiedener  Typen.  Basalte  mit  den 
dazugehorenden  Gangformen  von  ungefahr  gleichem  Mineralbestand  wie  die  auf 
Masatierra  vorkommenden  gibt  es  zvvar  auch  hier,  ausserdem  finden  sich  aber 
auch  an  Erzmineralien  stark  iibersattigte  Basalte,  sowie  den  Trachytandesiten  sich 
nahernde  Gesteine.  Am  interessantesten  ist  jedoch  das  Vorkommen  von  reinen 
Alkaligesteinen,  wie  z.  B.  die  von  QUENSEL  angefiihrten  Natrontrachyte. 


Natrontrachyt. 

Leider  ist  das  einzige  mitgebrachte  Handstiick  dieses  Gesteins  von  einem 
losen  Block  am  Fuss  der  Steilwand  von  Tierras  Blancas  abgeschlagen.  Skotts- 
HER(;  hat  indessen  miindlich  berichtet,  dass  zahlreiche  Blocke  desselben  Gesteines 
in  den  Talusbildungen  von  Tierras  Blancas  vorkommen,  und  dass,  soweit  er  ver- 
stehcn  koniitc,  dasselbe  hellgraue  Gestein  den  ganzen  oberen  Teil  der  Steilwand 
bildct;  seiner  Kartenskizze  nach  zu  urteilen  tritt  dasselbe  bereits  400  m  ii.  d.  M. 
auf.  Dies  ist  von  Bedeutung  fiir  das  Feststellen  der  Eruptionsfolge,  die  spater  kurz 
erwalint  wcrden  soil. 

Das  (iestein  besteht  aus  gleichmassigen  Kornern  und  ist  sehr  reich  an  Feld- 
spat. Die  I^'eldsj)atleisten  erreichen  eine  Lange  von  0,3 — 0,4  mm.  Sie  sind  gut 
parallclorientiert  und  verleihen  dem  Gestein  eine  trachytoidale  Struktur.  Da  Albit- 
zwillinge  nicht  vorhanden  sind  und  der  Feldspat  durch  die  Anlagerung  der  diinnen 
Individucn  unscharfe  Bcgrenzungen  zeigt,  konnte  eine  genaue  Bestimmung  des- 
selben nicht  ausgefuhrt  vverden.  Die  Lichtbrechung  halt  sich  im  allgemeinen  etwas 
liber  Kollolith  (n  1,535),  stellenweise  ist  das  Relief  jedoch  ganz  verschwunden. 
Urn  eine  nalierc  Kcnntnis  von  den  Feldspaten  zu  bekommen,  ist  eine  Alkalibestim- 
mung  des  Gesteins  ausgefuhrt  worden.  Diese  ergab  3,45  %  KgO  und  7,34  %  NagO. 
Dies  wiirdc  einem  Gehalt  von  20,44  %  Ortoklas  und  62,27  %  Albit  im  Gestein  ent- 
sprechen.  Auf  (irund  der  Lichtbrechungsverhaltnisse  diirfte  jedenfalls  neben  einem 
Kali-Xatronfeidspat  auch   cin  saurer  Plagioklas  der  Oligoklasreihe  vorhanden  sein. 


BEITRAGE    ZUR    GEOLOGIE    DER  JUAN    FERNANDEZ-INSELN 


29 


Eine  geringere  Menge  Pyroxen  tritt  ebenfalls  auf.  Die  durchschnittliche 
Ausdehnung  desselben  ist  0,1  mm,  die  Farbe  ist  gelbbraun,  die  kristallographische 
Ausbildung  schlecht  entwickelt.  Eine  Ausloschung  von  c:x  =  40°  (ungef.)  deutet 
auf  Augit.  Schliesslich  war  auch  Magnetit  vorhanden,  der  ofters  fliessende  Be- 
grenzung  der  graubraunen  Glasbasis  gegeniiber  zeigt. 


..—  Aragonitband,  von 
einem  Augitindivi- 
duum  iiberquert 


Fig.  6.     Hydrothermal  umgewandelter  Olivinfels  von  dem  Yunque. 
—  Vergr.  12,5  x,     Verf.  phot. 


J 

9 

m 

i^fe' 

^■^M 

---^r 

'^:f^ 

y  •   « 

■    .       '     z           ,             . 

.    .    .    * 

Fig.  7.     Sinterbildung  von  Puente.    Massstab  in  Cm.     Verf.  phot. 


Der  von  QuENSEL^  beschriebene  Natrontrachyt  ist  ungefahr  1200  m  ii.  d,  M. 
gefunden  worden  und  zeigt  bei  einem  Vergleich  einige  Abweichungen.  Das  Hand- 
stuck  scheint  im  Gegensatz  zu  dem  obenbeschriebenen  etwas  verkieselt  zu  sein. 
U.  d.  M.  bemerkt  man  sofort,  dass  der  Pyroxen  hier  abweichend  von  dem  vorer- 
wahnten  farblos  ist.  QUENSEL  bezeichnet  denselben  als  Diopsid.  Ferner  ist  die 
farbige  Glasbasis  nicht  vorhanden. 

1  L.  c.  p.  283. 


30 


TOR    H.    HAGERMAN 


Andesit. 


Von  (lem  Berggipfel  »Las  Torres >,  1 370  m  ii.  d.  M.,  und  in  losen  Blocken 
von  dem  Ik'rgrucken  XO  von  der  genannten  Stelle  1 200  m  ii.  d.  M.  sind  Hand- 
stiicke  von  einem  feldspatreichcn  aschgrauen  Gestein  mitgebracht  worden.  Obwohl 
etwas  olivinreicher,  stelien  dieselben  den  von  Olensel^  beschriebenen,  einem 
1 100  m  ii.  d.  M.  liegenden  Niveau  entnommenen  Trachytandesiten  sehr  nahe.  Als 
Kinsprenglinge  koniinen  Feldspat  und  Olivin  vor,  u.  d.  M.  zeigt  der  erstgenannte 
eine  Zusammensetzung  von  AbggAnga  und  erreicht  eine  Korngrosse  von  3 — 4  mm. 
Die  Olivine  sind  etwas  kleiner.  Sie  werden  im  allgemeinen  nur  ca.  i  mm  in 
Diameter  und  sind  wenig  verandert,  nur  an  den  Randern  zeigt  sich  eine  schwach 
gelbe  Farbe,  wo  die  Umwandlung  begonnen  hat.  In  der  Grundmasse  dominiert 
der  Plagioklas  mit  einer  Ausdehnung  von  ca.  0,1  mm.  Die  Pyroxene  und  Magnetite 
sind  noch  kleiner,  im  allgemeinen  nur  0,05  mm.  Die  Magnetite  sind  vollig  idio- 
morph.  Die  Klassifizicrung  hiehergehorender  Gesteine  ist  etwas  unsicher.  Die 
trachytoidale  Struktur  konnte  auf  einen  gevvissen,  nicht  wahrnehmbaren  Alkali- 
gelialt  in  der  (irundmasse  deuten.  Geniigende  Griinde,  sie  als  Trachytandesite  zu 
bezeiclinen,  liegen  jedoch  nicht  vor. 

Sehr  iiiteressant  ist  ein  bei  Correspondencia  (1420  m  ii.  d.  M.)  genommenes 
(iestein.  Mikroskopisch  zeigt  dieses  Handstiick  ein  aschgraues,  porphyrisches 
Aussehcn,  doch  sind  einige  Partien  bedeutend  dunkler.  Als  Kinsprenglinge  kom- 
men  1^'eldspat  und  Olivin  vor.  U.  d.  M.  zeigt  der  erstgenannte  eine  Zusammen- 
setzung Ab42An58  und  erreicht  eine  Korngrosse  von  durchschnittlich  2  mm  bei 
einer  niax.-Lange  von   5   mm. 

Der  Olivin  ist  vollig  frisch,  die  Kinsprenglinge  treten  aber  in  zwei  verschie- 
dencn  ICntwicklungen  auf.  Dies  steht  im  Zusammenhang  damit,  dass  das  Gestein, 
wie  bercits  erwahnt,  nicht  vollig  homogen  ist.  Die  dunkleren  Partien  erweisen 
sich  bei  mikroskopischer  Untersuchung  als  bedeutend  magnetit-  und  ilmenitreicher 
als  der  iibrige  Teil  des  Gesteins. 

In  den  dunkleren  Schlieren  finden  sich  nun  Olivine  mit  Magnetiteinschliissen 
vollgesteckt.  Iksonders  an  den  Randern  ist  der  Magnetit  so  reichlich  vorhanden, 
dass  die  Olivinkorner  vollig  opak  sind.  Sowohl  aus  diesem  Grunde  als  auch  in- 
folge  der  abgerundeten  Form  der  Mineralkcirner  scheint  es,  als  ob  diese  Olivin- 
korner einer  kraftigen  Resorption  ausgesetzt  gewesen  waren.  Naheliegend  ist  nun, 
dass  diese  dunkleren  Schlieren  mit  ihrem  grosseren  Kisengehalt  Riickstande  auf- 
geloster  Hruchstucke  sind,  welche  urspriinglich  zu  dem  Typus  gehorten,  die  einem 
schlackigen  l^asalt  vom  (jipfel  des  Inocentas  entsprechen,  der  spater  beschrieben 
werden  soil. 

In  einer  luitfernung  von  kaum  i^/g  mm  von  einem  der  erwahnten  Olivin- 
korner treten  Individuen  des  anderen  Typus  auf.  Diese  sind  ganz  einschlussfrei, 
vollig  idiomorph  mit  scharfen  Hegrenzungsflachen,  erreichen  einen  Durchschnitt  bis 
zu  4  mm  und  ents{)rechen  vermutlich  der  intratellurischen  Olivingeneration  des 
Ilauptgesteins. 

Der  Feldspat  in  der  Grundmasse  der  helleren  Schlieren  tritt  in  Stengein  von 
ca.  0,14  mm  Lange  auf  und  verleiht  durch  seine  Parallelorientierung  dem  Gestein 
eine  Fluidalstruktur.  Die  Zusammensetzung  desselben  ist  Ab46An54.  Im  iibrigen 
enthalt  die  Grundmasse  Magnetit  und  Pyroxen. 

Die    (irundmasse    der    dunkleren  Partien  unterscheidet  sich  von  der  obenge- 

^  L.  c.  p.  282. 


BEITRAGE    ZUR    GEOLOGIE    DER   JUAN    FERNANDEZ-INSELN  3 1 

nannten  durch  ihren  Gehalt  an  Ilmenit,  leicht  erkennbar  an  seinem  tafelformigen 
Habitus.  Ausserdem  kommen  hier  feine  Nadeln  vor,  welche  aus  einem  ziemlich 
stark  lichtbrechenden  Mineral  bestehen.  Dasselbe  ist  pleochroitisch  von  braun- 
gelber  bis  gelbgruner  Farbe  und  weist  parallele  Ausloschung  auf.  Wegen  der 
kleinen  Dimensionen  der  Korner  konnte  eine  sichere  Bestimmung  derselben  nicht 
ausgefuhrt  werden.  Mit  grosster  Wahrscheinlichkeit  liegt  hier  nur  eine  feinblattrige 
Ausbildung  von  Ilmenit  vor. 

Basaltische  Laven. 

Auf  dem  Uferplateau  an  der  Ostseite  der  Insel  bei  dem  Casastal  steht  ein 
feinkorniger  Basalt  mit  porphyrischen  Feldspat-  und  Olivineinsprenglingen  an. 
Die  Feldspate  erreichen  eine  Lange  von  0,9  mm  und  erweisen  sich  als  Plagio- 
klase  mit  einer  Zusammensetzung  von  Ab4oAn6o  und  stimmen  also  mit  dem  Feldspat 
der  Masatierra-Basalten  iiberein.  Die  von  QuENSEL^  erwahnten,  stark  basischen 
Feldspatkerne  habe  ich  nicht  angetroffen.  Die  Olivinkorner  erreichen  in  diesem 
Praparat  eine  Grosse  von  0,3 — 0,4  mm  und  sind  etwas  iddingsitumgewandelt.  Von 
den  Mineralien  in  der  Grundmasse  werden  die  Plagioklase  am  grossten,  0,08  mm. 
Der  Pyroxen  ist  dagegen  so  klein,  dass  eine  nahere  Bestimmung  sich  nicht  aus- 
fiihren  Hess.  Er  erscheint  in  kleinen,  viereckigen,  farblosen  Kornern,  meistens 
zusammen  mit  dem  Magnetit. 

Diesem  Gestein  sehr  nahe  verwandt  ist  dasjenige,  welches  am  Ufer  des  Mono- 
Tales  ansteht.  Makroskopisch  sind  die  beiden  Gesteine  einander  sehr  ahnlich. 
U.  d.  M.  tritt  jedoch  ein  Unterschied  auf,  und  zwar  indem  die  Grundmasse  des 
Monobasaltes  hier  bedeutend  mehr  Olivin  enthalt,  weshalb  man  dieses  Gestein 
auch  wegen  der  zahlreicheren  Olivineinsprenglinge  als  einen  Olivinbasalt  bezeichnen 
muss,  wahrend  sich  das  erstgenannte  den  Feldspatbasalten  nahert. 

In  Quebrada  del  Ovalo,  ungefahr  150  m  ii.  d.  M.,  steht  eine  saulenformige 
Basaltkuppe  an,  die  dem  Tal  seinen  Namen  gegeben  hat.  Sie  ist  oben  horizon- 
talzerkliiftet,  wahrend  weiter  unten  eine  prismatische  Vertikalzerkliiftung  ansetzt. 
Vorausgesetzt,  dass  die  Kuppe  aus  ein  und  demselben  Gesteine  besteht,  kann 
dies  als  ein  gutes  Beispiel  der  von  Iddings^  dargetanen  Veranderung  in  der 
Richtung  fiir  den  kleinsten  Schrumpfwiderstand  innerhalb  des  erstarrten  Gesteins 
angesehen  werden. 

Die  Handstiicke  sind  den  unteren  Teilen  der  20 — 30  m  hohen  Saule  ent- 
nommen  und  bestehen  aus  einem  porphyrischen  Olivinbasalt.  Die  Einsprenglinge 
sind  Olivine  von  ca.  0,6  mm  Durchmesser.  Dieselben  sind  an  den  Randern  dunkel- 
rot,  kaum  durchleuchtend,  was  wahrscheinlich  durch  einen  Gehalt  an  freiem  FcgOg 
verursacht  wird.  Untenstehende  Photographic  (Fig.  9)  stellt  einen  dieser  iibrigens 
sehr  sparlich  vorkommenden  Einsprenglinge  dar. 

Die  Grundmasse  ist  der  des  vorstehend  erwahnten  Olivinbasalts  voUig  gleich, 
nur  etwas  grober.  Der  Plagioklas,  ein  Labrador,  erreicht  eine  Lange  von  ca. 
0,12  mm,  das  ganze  Praparat  ist  parallelorientiert.  Besonders  hervorgehoben  sei, 
dass  das  Olivin  in  der  Grundmasse  nicht  rotpigmentiert  ist.  Im  iibrigen  findet 
sich  Magnetit  und  der  farblose  Pyroxen. 


1  S.  p.  276. 

2  J.  P.  Iddings,  The  columnar  structure  in  the  igneous  rocks  of  Orange  Mountain,  N.J. 
Amer.  Journ.  31  (1886),  p.  321. 


32 


TOR    H.    HAGERMAN 


Der  hochste  Berg  auf  Masafuera  ist  der  Inocentes.  Von  dem  Gipfel,  ca.  1 500 
m  ii.  d.  M.,  wurde  ein  rotlicher,  sehr  schlackiger  porphyrischer  Basalt  mitgebracht, 
der  sich  besonders  durch  seinen  hohen  Gehalt  an  Erzmineralien  auszeichnet. 

Unter  dem  Mikroskop  erwies  sich  die  Grundmasse  als  hyalopilitisch.  Die 
Einsprenglinge  bestehen  hauptsachlich  aus  grossen,  im  ersten  Augenblick  voll- 
standig  opaken  Kornern  von  ca.  3  mm  Durchmesser.  Bei  genauerer  Untersuchung 
erweisen  sich  jedoch  mehrere  als  stellenweise  durchsichtig.    Diese  Teile  besitzen 


Fig.  8  a.     Das  heterogene  Ciestein  von  Correspondencia.  Dunklere  Partien  oben  und  unten 
sichtbar.  —  Vergr.    10  x.    Photo.  E.  Dahlstrom. 


die  hohen  Interferenzfarben  des  OHvins,  dazu  weisen  die  Korner  auch  im  iibrigen 
den  Habitus  des  Oiivins  auf.  Bei  Beobachtung  in  konvergentem  polarisiertem 
Licht  bei  einem  Achsenaustritt  wurde  keine  Krummung  des  Achsenbalkens  beob- 
achtet,  vveshalb  der  Achsenvvinkel  nahezu  90°  sein  durfte.  Dies  deutet  auf  cine 
fur  liasahe  normale  Zusammensetzung  von  OHvin  mit  verhaltnismassig  niedrigem 
Eisengehalt. 

Kand.  S.  L.\M)E1<(;kkn  fuhrte  eine  Eisen-  und  Titanbestimmung  der  Ge- 
steinsarten  aus,  vvobci  folgende  Werte  erhalten  wurden:  18,82%  FeaOg  und  3,59  % 
TiOg.  Dies  entspricht  einem  Gehalt  von  6,81%  Ilmenit  und  1 5,44  %  Magnetit. 
Diese  Zahlen  sind  natiirlich  zu  hoch,  da  Fe  auch  in  den  geringen  Mengen  Olivin, 
Pyroxen    und    Glasbasis    enlhalten    ist,   die  sich   in  dem  Gestein  befinden.    Es  ist 


BEITRAGE    ZUR    GEOLOGIE    DER    JUAN    FERNANDEZ-INSELN 


33 


jedoch  besonders  interessant,  dass,  obgleich  das  Magma  eisenubersattigt  war,  das 
Eisen  die  Konstitution  des  Olivins  nicht  nennenswert  beeinflusste. 

Von  solchen  magnetitijbersattigten  Olivinen  spricht  Doss^  in  einer  Beschrei- 
bung  von  Basalten  aus  Syrien:  »In  diesen  Gesteinen  beherbergt  der  porphyrische 
Olivin  eine  derartige  Menge  von  Magnetitkornern,  dass  dieselben  meist  die  Halfte, 
zuweilen  ungefahr  Vs  des  ganzen  Kristalldurchschnittes  einzunehmen  scheinen».  An 
einer  spateren  Stelle  schreibt  er:   »Das  Extrem  hiervon  tritt  dann  ein,  wenn  der 


6V\ 


Fig.  8  b.     Detail  von  Fig.  8  a  (bei  — ).  —  Vergr.  210  x,    Zu  beachten  ist  die  parallele 
Anordnung  des  Magnetits.    Photo.  E.  Dahlstrom. 


Olivinkrystall  einen  breiten,  vollig  opaken,  schwarzen  Saum  von  Magneteismen 
besitzt».  Das  Phanomen  stimmt  mit  dem  vorliegenden  voUkommen  uberein,  nur 
dass  bier  der  entgegengesetzte  Fall  vorliegt,  indem  das  ganze  Korn  voUstandig 
opak  sein  kann,  auch  in  sehr  diinnen  Praparaten. 

Reiter2  hat  sich  mit  der  Zusammenschmelzung  von  Olivin  und  Magnetit 
beschaftigt  und  sagt:  »Bei  Abkuhlung  tritt  eine  gewisse  Ubersattigung  ein.  Vom 
Magnetit  scheidet  sich  ein  Teil  ab,  dann  wird  der  ubersattigte  Olivin  ausge- 
schieden  .  .  . 


1  B.  Doss,  Die  basaltischen  Laven  und  Tuffe  der  Provins  Hauran  und  vom  Diret  et-TuluI 
in  Syrien.    T.  M.  P.  M.    Bd.  7  (1886),  p.  483—484. 

2  L.  c.  p.  232. 


34 


lOR    H.    HAGKRMAN 


Wir  erhalten  hiemit  also  Magnetite  in  zonarer  Anordnung  in  den  nachher 
entstehentlen  Olivinen  ein^esclilossen;  ...» 

Die  umstehende  Tliotographie  (Fig.  lo)  zeigt,  wie  der  Magnetit  parallel  an- 
geordnet  ist,  sodass  die  durchsichtigen   Partien  in   Streifen  auftreten. 

Unter  den  I^insprenglingen  komnit,  wenn  auch  selten,  hier  und  da  ein  Feld- 
si)atindividiiiim  \  or.  dessen  Zusamniensetzung  ungefahr  Ab4oAn6o  ist;  es  handelt 
sich  also  um  einen  Labrador,  unbedeutend  abweichend  von  dem  Feldspate  der 
ubrigen   (iesteinsarten. 

Die  basischen  Tlagioklase  der  Grundmasse  zeigten  zum  Teil  unscharfe  Be- 
grenzungen.  durcli  die  Anlagerung  der  diinneren  Individuen  verursacht. 

Die  Pyroxene  sind  bedeutend  kleiner  und  erreichen  einen  Durchschnitt  von 
ca.  0,02—0.03  mm,  wenn  auch  einzelne  stengelige  Individuen  etwas  grosser  werden 


-0*^3^ 


r/iA 


Fi 


•'K- 


9.     Kotj)igmentierte  Olivineinsprenglinge.    Basalt  von  El  Ovalo.  —  Vergr.  56 
Photo.   Hi.  Olsson. 


konncn.  \'on  diesen  kommen  zvvei  verschiedene  Ausbildungen  vor.  Der  eine  (nor- 
male)  ist  mit  den  in  den  beschriebenen  basaltischen  Gesteinen  auftretenden  Py- 
roxen  indentiscli  und  vermutlich  als  Diopsid  zu  betrachten,  da  er  beinahe  farblos 
und  kaiun  pleochroitisch  bei  einer  Ausloschung  von  ca.  45°  ist.  Der  andere  Typus 
ist  braimgelb  und  tritt  besonders  uin  die  Locher  herum,  niemals  zusammen  mit 
dem  erstgenannten  Tyjius  atif.  stellt  aber  wahrscheinlich  nur  eine  Pigmentierung 
dcsselbeu  dar.  Auch  hierin  zeigt  das  Gestein  eine  grosse  Analogic  mit  den  vor- 
erwahnten.  von  Dos.s  beschriebenen  I^asalten.  \\x  sagt  auf  Seite  481:  »Hier  be- 
sitzen  die  in  der  Xahe  der  von  Kalkspat  ausgefiillten  Hohlraume  gelegenen  Augite 
eine   goldgelbc   I'\-irbc;  .  .  .    . 

Im  iibrigcn  bcsteht  die  (irimdmasse  aus  kleinen,  idiomorphen  Magnetitkornern, 
Ilmenit  luul  Glasbasis.  Der  ganze  (iebirgskamm  soil  aus  diesem  ausgesprochen 
basischen   Gestein   l;cstehen. 

I^ereits  mikroskopisch  zeigt  ein  anderes  Handstuck,  das  vom  Strandabhange 
nahe    des    Casatales    abgeschlagen   ist,  eine  auffallende  Ahnlichkeit  mit  dem  Ino- 


BEITRAGE    ZUR    GEOLOGIE    DER   JUAN    FERNANDEZ-INSELN  35 

centesgestein.  (Laut  Angabe  von  Skottsberg  soil  das  ganze  ostliche  Ufer  aus 
diesem  Gesteinstypus  bestehen,  abwechselnd  mit  den  obenerwahnten  Feldspat- 
basalten.) 

Auch  hier  treten  magnetitfuhrende  Olivine  auf,  wenn  auch  sparlicher.  Die 
Feldspate  sind  dieselben  und  besteht  der  einzige  Unterschied  darin,  dass  das 
Gestein  glasreicher  ist,  weshalb  der  in  dem  letztgenannten  Gestein  vorkommende 
Pyroxen  nicht  mit  Bestimmtheit  hat  wahrgenommen  werden  konnen.  Nimmt  man 
an,  dass  dieses  Gestein  einer  rascheren  Abkiihlung  ausgesetzt  gewesen  ist  als 
das  vorgenannte,  kann  man  dasselbe  diesem  gleichstellen  und  moglicherweise 
eine  Andeutung  iiber  eine  nicht  unbedeutende  Ausbreitung  der  fraglichen  stark 
basischen  Gesteinsart  finden. 


Fig.   10.     Magnetitiibersattigte  Olivinsprenglinge  im  Basalt  von  Inocentes.    Verf.  phot. 

Umstehende  Tabelle  ist  eine  Zusammenstellung  iiber  die  Veranderlichkeit 
der  efifusiven  Masafuera-Gesteine  im  Verhaltnis  zum  Niveau  der  Fundstatten.  Die 
Gesteine  sind  nach  abnehmender  Basisitat  geordnet  und  dabei  das  Material  Skotts- 
BERG's  wie  auch  dasjenige  QuENSEL's  beriicksichtigt. 

Zwar  fallen  die  Lavabetten  etwas  nach  NNO  ab,  doch  diirfte  mit  Riicksicht 
auf  den  geringen  Umfang  der  Insel  die  umstehende  Tabelle  ein  gutes  Bild  der 
Eruptionsfolge  gevvahren,  wobei  die  altesten  Gesteine  beim  Meeresniveau,  also 
links  beginnen. 

Vor  allem  verstosst  der  stark  basische  Basalt  aus  der  Hohe  von  1500  m  ii. 
d.  M.  gegen  die  unter  Zugrundlegung  der  Eruptionsfolge  vorgebrachte  Theorie, 
dass  die  Gesteine  hier  gravitativ  differentiert  sein  sollen.  Die  Gesetzmassigkeit, 
die  QUENSEL  bei  diesen  Gesteinen  gefunden  zu  haben  glaubte  —  mit  den  sauren 
und  alkalireichen  als  den  jiingsten  —  scheint  mir  schwerlich  mit  diesen  Beob- 
achtungen  in  Ubereinstimmung  gebracht  werden  zu  konnen. 


36 


TOR    H.    HAGERMAN 


1  Natrontrachvte  .  .  . 

! 

i   !  o 

!          i          1 

1 

X 

i 
1 



1 

1 

1     1 

O 

O 

1 

Basanit 

1 

X 

j^^^^l^^FeUispat      ^q  i         '\         1 

Ohvin-          i  oo 

O 

1          i 

Er/iibersatti^ae  Ha 
salte 

0 

i          1 

O 

13       14  1500 

m  ii.  d.  M. 


Die  \'erteilung  der  Effusivgesteine  der  InseJ  Masafuera. 
O  bezeichnet  Das  Material  Skottsberg's. 
X  »  »  ,  QUENSEL's. 


Ks  ist  selbstverstandlich  denkbar,  dass  die  in  den  tieferen  Teilen  des  Zufluss- 
kanales  versunkcnen  schweren  Magmas  zuletzt  herausgestossen  wurden,  doch  spricht 
gcgcn  diese  Annahme  das  bereits  in  einer  Hohe  von  400  m  ii.  d.  M.  auftretende 
alkalisciie  Ciestein. 

Scliliesslich  ist  aus  der  Tabelle  zu  ersehen,  dass  aus  einer  Hohe  von  500 — 
1000  111  Liberhaupt  kein  Material  untersucht  ist.  Die  Masafuera  Gesteine  sind 
jcdenfalls  in  Minblick  aiif  den  Mineralbestand  so  verschiedenartig,  dass  eine  nahere 
L'ntcrsuchung  der  noch  unbekannten  Hohenlagen  wichtige  Aufschliisse  ergeben 
durfte. 

/'»'j-/  ^7-intcd  February  2^th,  ig2^. 
Reprinted  ivithout  change  April  13th,  igjj4. 


3-  Additional  Comments  on  the  Geology  of  the 
Juan  Fernandez  Islands. 


By 

PERCY  OUENSEL. 


Contents. 

Page 

Introduction      ^^ 

Main  Geological  Features 40 

Petrology  of  the  Volcanic  Formations 44 

Masatierra 4^ 

Masafuera 56 

Regional  Relations. 

Tectonic  Connections -74 

Petrographic  Connections 77 

General  Conclusions      79 

Acknowledgements 82 

Tables  of  Analyses 83 

Bibliography 86 


Introduction. 

The  Juan  Fernandez  Islands  consist  of  Masatierra,  Masafuera  and  Santa 
Clara.  In  many  publications  the  name  Juan  Fernandez  has  been  used  to  denote 
Masatierra  only,  the  other  islands  then  being  indicated  by  the  names  as  above. 

The  largest"  island,  Masatierra,  situated  660  km  from  and  nearly  due  west 
of  Valparaiso,  measures  95  square  km.  Masafuera,  170  km  further  westward,  is 
64  square  km.  The  small  island  Santa  Clara,  close  to  Masatierra,  is  only  5 
square  km  in  area. 

During  more  or  less  casual  visits  to  Juan  Fernandez  stray  observations  have 
been  recorded  on  geological  features  of  the  islands.  In  all  cases  they  refer  to 
Masatierra.  It  may  be  of  interest  in  this  connection  to  give  a  summary  recount 
thereof. 

The  first  samples  of  volcanic  rocks  of  Masatierra  were,  as  far  as  known, 
collected  by  Lord  CocilCRANE  in  January  1823.  He  was  returning  to  Europe 
after  5  years  service  as  admiral  in  Chilean  service  during  the  war  of  independ- 
ence. The  ship  at  his  disposal,  'Colonel  Allen',  touched  at  Masatierra  for  two 
days.  Mrs  Maria  Graham,  a  passenger  on  board,  has  in  her  diary  given  the 
following  details,  recorded  by  Thomas  Sutcliffe  in  his  book  Cmsoniana:  "Lord 

3  —  516795     The  Nat.  Hist,  of  Juan  Fernandez  and  Easter   Isl.    Vol.  I. 


3S  IMKlV    (JIKNSKL 

Cochcrane  brought  from  the  siiniiiiit  (1.500  feet)  a  piece  of  black  porous  lava; 
and  uiuler  it  lie  foiiiul  some  dark  iiardened  clay  full  of  cells,  the  inside  of  which 
appear  sli^litly  \  itritk-d.  The  ishuul  seems  chiefly  composed  of  this  porous  lava; 
the  strata  of  which,  beini;  crossed  at  ri^ht  an^^les  by  a  very  compact  black  lava, 
dij)  on  tile  eastern  side  of  the  island  about  22'  and  on  the  west  side  l6°,  pointing 
to   the   centre   of  tiie   island   as   an   apex"    (l,    j).   198). 

In  1S30  C\  Hi:kii:r<»  j^ublished  some  observations  under  the  title  'Notice 
sur  rilistoire  naturelle  de  file  juan  hV-rnandez'.  With  regard  to  geological  ques- 
tions he  sa\s:  "je  pense  ciu'un  geologue  n'y  trouverait  cjue  du  basalte  dans  les 
etats,  lueiue  dans  cehii  de  la  j)lus  parfaite  decomposition;  plusieurs  blocs  sont 
parsemes  dune  cristallisation  j)articulicre,  a  laquelle  on  donne,  je  croix,  le  nom 
d()li\ine  ...  11  n'\-  a  pas  de  trace  de  volcan ;  les  pierres  qu'on  prend  pour 
de  la  hue,  et  dont  (iuel(|ues-unes  ressemblent  assez  aux  scories  on  de  la  pierre 
j)once.  ne  sont,  a  mon  avis,  (jue  du  basalte  decompose;  on  trouve  aussi  cette 
rociie  sous  forme  sj)hericiue.  et  composee  de  couches  concentriques  ..."  (2, 
p.  34;;   compare   in   the   latter  respect   h^ig.  10  on   page   52   of  this  paper). 

.\.  (Ai.ix  i.Kic;!!,  who  accomj)anied  Captain  P.  Parker  Kixc;  on  the  sur- 
\f\ing  \()\ages  of  II.  M.S.  Adventure  and  Beagle  on  their  first  expedition 
1S26  1S30,  read  before  the  (Geological  Society  of  London  on  Jan.  5th,  1831,  a 
statement  on  'The  geology  of  the  island  of  Juan  Fernandez'.  In  the  Proceedings 
of  that  N'ear  the  following  account  of  Caldcleugh's  discourse  is  given:  "The 
author  could  discover  no  trace  of  a  volcano,  said  to  exist  here  by  former  visitors; 
all  the  rocks,  according  to  him,  consist  of  basaltic  greenstone  and  trap  of  various 
mineralogical  structure,  both  amorphous  and  vesicular,  together  with  trappean 
concretions,  no  other  contained  minerals  being  observable  except  olivine  and 
chaux  carbonatee  metastaticpie.  It  is  further  mentioned  that  the  basalt  in  parts 
is  almost  colunuiar,  and  in  others  has  a  [)eaked  and  serrated  outline,  the  mass 
being,  here  and  there,  traversed  by  dykes.  Owing  to  the  peculiar  character  of 
this  basalt,  and  especially  from  the  great  quantity  of  olivine,  the  author  compares 
its  age  with  that  of  the  basalt  of  Bohemia,  the  Rhine,  the  Vivarrais  and  Beaulieu 
in  Provence  '  (3,  p.  256,  also  i)uhlishe(l  in  the  Phil.  Mag.  and  Annals  of  Philosophy, 
\'ol.  l.\,    I  S3  I,   }).  220). 

Captain  King  recaj)itulates  Caldcleugh's  narrative,  as  given  above,  with  the 
addition:  "In  captain  llAi.i.s  interesting  journal,  there  is  a  list  of  geological  and 
mmeralogical  specimens,  of  which  one  from  Alasafuera  is  named  vesicular  lava" 
I4.    p.  304).    '1  he   uhimate   destiny   of  these  specimens   is   unknown. 

Members  of  tlie  Dumont  dl'rville  expedition,  when  visiting  Masatierra  in 
1S3S.  collected  and  specified  several  different  saiu})les  of  the  lavas  from  the  island 
'-^-  P-  ■'4'-  1  he  material  tor  the  new  analyses  of  basalts  from  Masatierra,  which 
L\(K<iix  recentl\-  caused  to  be  made  and  which  will  be  referred  to  later  on,  are 
evidentlx-  trom  this  collection,  as  Lacroix  says  they  were  made  from  specimens 
collected  by  the  Dumont  dCrMlle  exj)edition.  G.  (iUANCiK  records  some  observa- 
tions as  tollows:  "Toutes  les  roclies  aj)j)artiennent  a  diverses  varictes  de  trapp 
et  de  diorite  basaltique  amorphes  et  vesiculaires,  on  ne  trouve  dans  ces  roches 
volcanicjues  aucun  autres  mineraux   (jue  de  I'olivine  et  de  la  chaux  metastatique. 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  39 

La  roche  basaltique  s'y  presente  le  plus  souvent  en  couches  superposees,  quel- 
quefois  en  escarpements  interrompus  et  fractionncs  et  sont  traverses  par  des  dikes 
d'ejections  plus  modernes.  Le  basalt  forme  des  pics  eleves  dans  I'lle  et  sur  quelques 
points  prend  une  disposition  prismatique  fort  remarquable''  (6,  p.  39). 

In  1896  L.  H.  Plate  published  a  paper  'Zur  Kenntnis  der  Insel  Juan  Fernan- 
dez'. Plate  was  a  zoologist  and  the  paper  deals  with  the  zoology  of  Masatierra, 
but  some  introductory  remarks  refer  to  geological  observations  from  which  the 
following  instances  may  be  quoted:  "Das  Gestein  der  Insel  ist  ausschliesslich 
vulkanischer  Natur  und  besteht  aus  schwarzer,  basaltischer  Lava,  der  an  einzelnen 
Stellen  weissliche  oder  rothliche  Tuffe  eingelagert  sind  .  .  .  Die  Insel  fallt  fast  iiber- 
all  mit  senkrechten  Wanden,  deren  Hohe  zwischen  ico  und  300  m  schwankt,  gegen 
das  Meer  zu  ab  und  nur  in  den  Hafen  der  Ansiedelung  (Bahia  Cumberland),  dem 
Puerto  Ingles,  dem  Puerto  Frances  und  der  Bahia  de  la  Vaqueria  erstrecken  sich 
die  Thaler  bis  an  die  Ki.iste,  so  dass  man  in  diesen  Stellen  ohne  Miihe  vom 
Meer  in  das  Innere  der  Insel  vordringen  kann  .  .  .  Die  Lavawande  sind  deutlich 
geschichtet  ,  .  ,  Betrachtet  man  nun  vom  Meer  aus  eine  (solche)  Lavawand,  so 
erkennt  man  sofort  dass  sie  geschichtet  ist;  denn  sie  wird  in  ganzer  Ausdehnung 
von  zahlreichen  horizontalen  Linien  durchsetzt  .  .  .  Diese  horizontalen  Linien  sind 
wohl  der  Ausdruck  des  successiven  Aufbaues  der  Insel.  Aus  einem  submarinen 
Krater  ergossen  sich  Lavastrome  und  breiteten  sich  auf  dem  Grund  der  Oceans 
aus.  Die  PLruptionen  wiederholten  sich  haufig,  und  so  floss  eine  Lavaschicht  iiber 
die  andere,  um  spater  zum  Teil  iiber  die  Oberflache  des  Meeres  gehoben  zu 
werden  .  .  .  Ohne  Zweifel  war  die  Insel  in  friiheren  Erdperioden  sehr  viel  grosser. 
Eine  Untersuchung  des  Meeresbodens  zwischen  Masatierra  und  Masafuera  wird  viel- 
leicht  spater  den  Beweis  bringen,  dass  die  beiden  Inseln,  welche  jetzt  92  Seemeilen 
von  einander  liegen,  urspriinglich  nur  eine  einzige  bildeten  oder  doch  wenigstens 
die  hochsten  Punkte  desselben  submarinen  Plateaus  darstellen  und  daher  gleich- 
zeitig  entstanden  sein  miissen"   (7,  p.  221). 

The  observations,  hitherto  recorded,  have  hardly  more  than  historical  interest. 
As  little  attention  had  been  paid  to  the  geology  of  the  Juan  Fernandez  Islands 
before  the  last  decade  of  the  past  century,  these  casual  comments  have,  however^ 
here  been  included  to  evade  oblivion. 

The  first  observations  of  any  importance  regarding  the  rocks  of  Masatierra 
are  based  on  specimens,  collected  by  members  of  the  Challenger  Expedition  in 
1876.  A.  RenarD  has  given  a  description  thereof  in  a  paper  titled  'Rocks  of 
the  Island  of  Juan  Fernandez'.  (This  name  here  signifies  Masatierra.)  He  says 
that  "the  rocks  which  have  been  submitted  to  examination  all  belong  to  the 
basalt  type,  and  it  seems  probable  that  the  whole  island  is  made  up  of  those 
rocks  that  we  are  about  to  describe.  The  rocks,  which  form  the  central  mass  of 
the  island,  appear  in  the  specimens  as  dolerites  or  as  common  basalts  . .  .  Among 
the  specimens  collected  on  the  coast  of  Juan  Fernandez  it  is  necessary  to  mention 
a  greyish  very  scoriaceous  rock  from  which  stand  out  large  crystals  of  plagioclase 
of  a  waxy  and  milky  appearance.  This  rock  is  a  dolerite  with  large  vesicules.  Under 


40 


I'KRrV    (UF.NSKL 


the  microscope  tlie  fiiiulaineiUal  mass,  in  wiiich  the  plagioclase  crystals  are  embedded^ 
has  a  dolerite  structure.  Tlie  felspar  crwstals  show  large  extinction  angles  (38° — 41°) 
which  ma\-  be  compared  with  tiiose  of  b\townite.  The  sections  of  the  mineral 
are  cracked  and  pervaded  with  zeolite  matter,  which  forms  an  irregular  network. 
This  matter  which  looks  slightlx'  gre\-,  when  seen  by  ordinary  light,  remains 
obscured  l)etween  crossed  nicols  .  .  .  The  olivine,  of  which  large  sections  are  seen, 
is  uniform!)-  changed  into  a  red  hematite;  these  sections,  however,  still  show 
extinctions  like  those  of  the  unaltered  olivine"  (8,  [).  176).  The  characteristic 
change  of  olivine  to  iddingsite  in  the  basalts,  which  will  be  treated  below  in 
some  detail,  is  here  evidentl\-  noted  b)-  Renard,  though  by  him  named  hematite. 
I"'or  the  rest  his  description  mostly  refers  to  the  colour  and  texture  of  differ- 
ent samples  of  tlolerite  and   basalt. 

In  1886  L.  l).\K.\i'SK\  published  a  short  report  on  detached  rounded  lumps 
of  magnesite  from  Hahia  del  Padre  in  Masatierra,  locally  known  as  "piedras  de 
campana'  but  named  "(dockenstein"  by  the  author  (9).  R.  PoilLMANN  later  (1893) 
described  this  singular  formation  in  rather  more  detail  and  discussed  its  origin  (10). 

An  account  of  some  geological  and  petrographical  observations  by  J.  ScHULZE 
and  K.  I'(  »lll.M.\NN,  participants  in  Fedkrico  Joiiow's  expedition  to  the  islands 
in  1891,  has  been  published  by  the  latter  as  an  introduction  to  Johow's  monograph: 
'Ivstudios  sobre  la  Flora  de  las  islas  de  Juan   Fernandez'   (11,   p.  i). 

As  a  member  of  the  Swedish  Magellanian  Expedition  of  1907 — 1909  I  had 
the  opportunity,  together  with  Professor  Skott.SBERG,  of  visiting  the  islands  in 
1908  and  later  described  the  rock  specimens  then  collected  in  a  paper:  'Die 
(ieologie  der  Juan  I^'ernandezinseln'  (12).  In  1916 — 17  Skottsberg  again  visited  the 
islands  and  collected  specimens  from  some  new  localities.  These  were  subsequently 
described  by  '1\  H.\(;krmax  under  the  title  'Beitrage  zur  Geologie  der  Juan 
I'ernande/.  Inseln'   ( I  3). 

Since  Skottsberg's  visit  in  1917,  no  further  exploration  in  the  field  has  been 
|)ublishe(l  concerning  the  geology  of  the  islands  in  question.  And,  with  the  exception 
ot  tour  recent  analyses  of  lavas  from  Masatierra  from  collections  made  by  members 
of  Dumont  d  Frvillcs  A'oyage  au  I'ole  Sud  et  dans  rOceanie'  in  1838,  commented 
on  1)\-  L.\(k(»i\  (14,  p.  64),  no  further  observations  have,  as  far  as  known,  been 
j)ublishcd  on  the  gecjlogy  or  petrography  of  the  Juan  Fernandez  Islands.  On  the 
other  hand  (|uestions  relating  to  their  lithological  connection  with  other  intra- 
|)acific  islands  and  their  geophysical  position  in  relation  to  eastern-pacific  volcanic 
centres   has   been    tlie  subject   of  repeated   discussions   during   later  years. 

Main  Geological  Features. 

1  he  two  islands  Masatierra  and  Masafuera  j)resent  very  different  aspects  with 
regard  to  their  bulk  conti-uration.  Masatierra  exhibits  a  rugged  appearance  of 
isolated  jagged  cliffs  lI"io.  21.  The  highest  peak,  VA  Vunque,  is  927  m.  On  the 
clear-cut  sliore  boundc-d  escarpments  one  can  distinguish  many  hundred  lava 
beds  overlying  one  another.  In  thickness  they  can  vary  from  .some  few  metres 
up   to   20  m   or   more. 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  4! 


Fig.  I.  Perspective  view  of  Masafuera  from  the  East.  (Reproduced  from  F.  Johow,  Estudios  sobre 
la  flora  de  las  Islas  de  Juan  Fernandez.   Santiago  de  Chile   1896.; 


Masafuera  presents  notable  dissimilarities  to  Masatierra.  Its  dome-shaped, 
sharply  bounded  outline  with  an  approximately  oval  circumference  and  its  impos- 
ing height  give  an  impression  of  a  volcanic  configuration  deviating  from  that 
of  Masatierra.  On  the  west  side  of  the  island  the  cliffs  fall  all  but  perpendicularly 
for  over  i  ooo  m.  The  shore  line  below  is  a  stony  and  sandy  reach,  in  part 
named  Loberia  vieja  on  account  of  the  number  of  fur  seals  found  there  at  certain 
seasons.  The  eastern  coastline  has  a  very  different  appearance.  As  the  highest 
ridge  lies  towards  the  west,  the  fall  on  this  side  is  not  so  precipitous.  Instead 
it  is  traversed  by  numerous  narrow  and  steep  sub-parallel  erosion  valleys  (que- 
bradas).  Attempts  to  ascend  the  higher  parts  of  the  island  can  only  be  made  from 
the  eastern  side  by  means  of  these  quebradas,  a  climb  which  is  anything  but 
easy  going  (Fig.  3).    The  highest  peak  is   i  500  m. 

The  sketch  in  Fig.  i  gives  an  excellent  conception  of  the  singular  configura- 
tion of  Masafuera's  eastern  escarpments. 

The  Juan  Fernandez  Islands  are  exclusively  formed  of  volcanic  material,  in 
many  respects  of  much  the  same  nature  as  in  other  volcanic  islands  of  the 
eastern  Pacific,  All  indications  tend  to  show  that  they  are  of  late  origin.  Both 
VON  Wolf  (15,  1929,  p.  771)  and  J.  BrCggen  (16,  p,  59)  consider  them  to  be 
late  tertiary  or  pleistocene  in  age.  Lacroix  says  regarding  all  the  non-coral- 
line islands  of  the  southern  central  Pacific:  "leur  age  precis  est  indetermine, 
mais  il  est  certainement  tertiaire,  pleistocene  ou  meme,  dans  certains  cas,  recent" 

(14,  P-  55)- 

No  signs  of  recent  activity  have  been  found  on  the  Juan  Fernandez  Islands. 
That  the  immediate  environs  have  recently  been  subjected  to  sub-marine  volcanic 
activity  is,  however,  evident  from  a  narrative,  published  in  the  Report  of  the 
Challenger  F:xpedition  as  follows:  "In  1835  Masatierra  appears  to  have  been 
governed  by  a  Mr  SUTCLIFFE,  an  Englishman  in  Chilean  service.  He  was  present 
when  the  earth-quake  took  place  on  the  20th  of  February  of  that  year,  of  which 
he  gives  the  following  account:  At  11.30  a.m.  the  sea  rose  over  the  mole  and 
afterwards  retired,  leaving  the  greater  part  of  Cumberland  Bay  dry,  so  much  so 
that  old  anchors  on  the  bottom  became  clearly  visible.  The  earth  then  began  to 
shake  violently,  and  a  tremendous  explosion  was  heard,  the  sea  still  receding 
in  immense  rollers,  which  afterwards  returned,  violently  rising  to  such  a  height 
that  the  settlement  was  literally  covered  and  washed  away,  when  the  sea  again 
receded.  The  phenomenon  occurr-ed  four  times,  causing  much  destruction,  uproot- 
ing trees  and  drowning  cattle.   Shortly  after  the  explosion,  a  large  column,  some- 


42 


I'F.KCN     (JUKNSEL 


I■"iJ,^  2.  'I'lu'    south    sIoijc   of  the  high  ridge  along  the  western  lialf  of  Masatierra  as  seen  from 

Portozuelo.    Photo  C.  Skottsberg. 


what  rcscinblin<;  a  water-spotit,  was  seen  ascending  from  the  sea  off  point  l^acalab, 
which  proved  to  be  smoke,  but  at  7  |).m.  volcanic  flames  were  visible  through 
the  smoke,  which  lasted  till  2  a.m.  on  the  2ist.  The  depth  of  the  water  on  the 
spot,  where  the  eruj)ti()n  took  j)lace,  was  from  50  to  80  fathoms;  no  alteration 
in  the  dej)th  was  detected  after  the  eruption  had  subsided"  (17,  p.  818).  Sutclifife 
has  published  an  account  of  the  'earthquake'  in  a  separate  publication  (18)  and 
reproduced   a  sketch   of  the  sidj-marine  eruj)tion   (19,   p.  387). 

It  is  obvious  that  these  narratives  must  refer  to  a  sub-marine  volcanic  explo- 
sion. Cii  ARMS  1)  \i<\\  IN  also  mentions  the  phenomenon  in  his  'Geological  Observa- 
tions on  the  X'olcanic  Ishmds'  (20,  p.  149).  Renard  gives  the  f)osition  of  the 
e.\|)losion  as  i  I-jioHsh  mile  from  the  island  and  remarks  "that  the  close  prox- 
imit>-  of  a   \olcanic  centre  seems  therefore  to  be  implied"   (8,   p.  176). 

1  he  sid) marine  eruption  nuist  be  taken  as  conclusive  evidence  that  the 
iuHuediate  nei^hl)f)iMhood  of  the  Juan  I'Y-rnandez  Islands  has  been  the  seat  of 
volcamc  action  within  the  last  11;  years.  A  point  of  further  interest  is  that  the 
e.xplosion  was  simultaneous  with  violent  earthcjuakes  on  the  Chilean  coast,  as 
Darwin    already   obserxcd   (see   j).  75). 

Hruggen  refers  to  some  finther  observations  of  sub-marine  eruptions  in 
the  vicinity  oi'  the  Juan  l-'ernandez  Islands,  recorded  by  Fk.  Goll  in  his  paper 
'J)ie  lu-dbeben  Chiles'  (Munchener  Geogr.  Studien  1904,  Nr.  14).  The  following 
denotements  by  GoM,  are  taken   from   Hriiggen   (16,   p.  332): 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE  JUAN    FERNANDEZ    ISLANDS  43 


Fig,  3.  Quebrada  del  Varadero  on  the  east  coast  of  Masafuera.  Photo  C.  Skottsberg. 


44  PERrv   (jrHNSKi, 

"'Se«4un  CioW  se  prodiijo",  cl  12  de  febrero  de  1839  una  erupcion  submarina 
\-  marenioto  a  unas  120  kni  al  este  de  la  Isla  de  ^las  a  Tierra.  rLl  mismo  autor 
cita  la  obscrvacion  si<;iiicnte  hecha  en  un  punto  iin  poco  mas  austral:  'l^^n  octubre 
de  1867,  se  sintio  un  temblor  submarino  en  34  55'  S  y  7/°  38'  W  (unas  lOO 
millas  SIC  de  Juan  I-'ernande/.V  despues  el  buque  navego  durante  dos  boras  por 
agua  tie  color  bianco  lechoso.  habiendo  mucho  pescado  muerto  en  la  superficie,' 
Se  trata  probablemente  del  mismo  fencHiieno  que  describe  JoSK  M.  Po.MAR  en 
la  forma  sigiiiente,  aun(]ue  dice  (jue  el  [)unto  se  halla  a  100  millas  al  S\V  en 
\  ez  de  Si'',  de  Juan  I-'ernantlez:  'ICn  1867,  el  capitan  SlMPSON  de  la  barca  britanica 
C'oronella  navegaba  en  el  I'acitico  con  mucha  calma  y  vientos  contrarios,  con 
e.\cej)ci()n  de  un  hierte  \  iento  acompaiiado  de  siete  temblores  que  se  produjeron 
como  a  100  millas  al  SW  de  la  Isla  de  Juan  Fernandez;  durante  dos  horas  navego 
|)or  agua  tan  blanca  como  leche;  sondeo,  pero  no  toco  fondo  en  100  pies  de 
|)rorim(latl,  \  io  muclios  pescados  muertos  y  una  gran  cantidad  de  pajaros  por 
todas  j)artes.  Agregaba  el  capitan  Simpson  que  si  hubiera  estado  10  millas  mas 
adelante,  el  cluxiiie  hubiera  sido  peor  y  hubiera  causado  averias  al  buque.'" 

Concerning  the  earthquake  of  Vallenar  in  the  province  of  Atacama  on  the 
loth  of  Nov.  1922  Ikuggen  cites  I^AII.EV  WlLUS'  observations  on  the  contem- 
poraneous volcanic  activity  on  San  Felix. 

I^ruggen  concludes  his  opinion  on  the  submarine  explosions,  given  above, 
as  follows:  "A  la  teoria  del  origin  de  los  tsunamis  por  erupciones  submarinas  podria 
objetarse  (jue  serian  erupciones  muy  excepcionales,  ya  que  consisterian  en  una  sola 
o  nuiy  pocas  explosiones  que  causan  las  pocas  olas  sismicas,  apagandose  luego 
la  actividad.  V.n  realidad  se  tratara  solamente  de  las  primeras  explosiones  que 
abren  la  chimenea  para  la  salida  de  la  lava  o  de  los  gases  y  que  tienen  la  fuerza 
suhciente  para  causar  el  tsunami.  Tambien  en  otro  sentido,  las  erupciones  sub- 
marinas se  distinguen  de  las  de  los  volcanes  de  los  Andes,  que  solo  excepcional- 
mente  entran  en  actividad  durante  los  terremotos.  Los  volcanos  submarines 
parecen  estar  en  relacicm  mas  estrecha  con  los  focos  sismicos  de  los  grandes 
terremotos  chilenos"  (16,   p.  332). 

Petrology  of  the  Volcanic  Formations. 

The  following  descrij)tion  of  the  rocks  of  Masatierra  and  Masafuera  is  based 
on  specimens  collected  by  myself  in  1908  and  by  Skottsbkrc;  in  191 7.  In  many 
cases  the  two  collections  supplement  one  another  and  help  to  elucidate  to  a 
certain   degree  the  distribution   of  the  somewhat  varying  types  of  the  lava  flows. 

1  he  specimens  from  Masatierra  in  both  collections  give  conclusive  evidence 
that  this  inland  in  the  main  is  formed  of  a  rather  uniform  series  of  basaltic  lava 
beds,  only  diverging  in  respect  of  coarser  or  finer  grain  or  of  a  higher  or  lower 
content  ot  oJiMiie.  The  specimens  from  Masafuera  on  the  other  hand  indicate 
more    obx  lou-,    dissimilarities  in   the  com[)osition   of  the  rocks  at  different  levels. 

In  the  following  the  principal  petrological  features  of  the  two  islands  will 
first  be  treated.  I'lHlcr  a  later  heading  references  will  be  made  to  resemblances 
in   various  respects  to  other  volcanic   islands  of  the  eastern  Pacific. 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  45. 

Masatierra. 

(Lat.  33°  37i'  S..  Long.  78°  50'  W.) 

The  two  collections  of  specimens  from  Masatierra  by  Skottsberg  and  my- 
self have  partly  been  taken  from  different  localities.  The  central  parts  of  the 
island  are  however  sparsely  represented.  Somewhat  more  complete  series  of  rock 
samples  originate  from  the  heights  around  Bahia  Cumberland  (Ensenada  de  San 
Juan  Bautista),  Puerto  Frances  and  Portezuelo  as  well  as  from  Bahia  del  Padre. 
From  the  adjacent  small  island  Santa  Clara  there  is  only  one  specimen.  It  is 
not  easy  to  single  out  the  local  distribution  of  the  different  lava  beds  on  account 
of  the  inconnected  localities  from  which  specimens  have  been  collected.  Some 
characteristic  features  may  however  be  found  which  indicate  that  certain  types 
of  lavas  are  restricted  to  localised  areas.  To  some  extent  one  may  then  draw 
conclusions  regarding  the  sequence  of  the  volcanic  eruptions. 

The  predominating  rocks  of  Masatierra  are  olivine  basalts,  differing  only  in 
their  content  of  olivine.  Lava  beds  from  around  Puerto  Frances  are  exceptionally 
rich  in  this  mineral,  the  content  of  which  can  reach  40  vol.%  of  the  rock  (Fig.  6).  Such 
rocks,  with  an  extreme  content  of  olivine,  I  named  picrite  basalts  in  my  earlier 
paper  (12,  p.  265).  Lacroix  originally  named  such  basalts  'picrite  feldspatique' 
but  later  discarded  this  name,  substituting  for  it  the  name  oceanite',  under  which 
name  he  includes  the  basalts  of  the  Juan  Fernandez  Islands  with  an  exceptionally 
high  content  of  olivine  (14,  p.  65).  Since  the  name  picrite  basalt,  as  originally 
defined  in  my  former  paper,  has  later  been  adopted ^  I  will  retain  this  name  for 
the  rocks  in  question,  with  the  name  oceanite  as  synonym. 

From  the  extremely  olivine-rich  basalts  around  Puerto  Frances  there  is  every 
transition  over  intermediate  types  to  feldspar  basalts  without  any  olivine  at  all 
or  with  only  a  very  insignificant  amount  thereof.  Such  rocks,  however,  have  a 
relatively  limited  distribution  on  Masatierra.  In  general  one  may  say  that  types 
with  a  very  high  content  of  olivine  are  restricted  to  lower  elevations,  whereas 
higher  up  more  normal  olivine  basalts  predominate. 

A  second  feature  of  dissimilarity  in  the  basalts  is  found  in  their  texture. 
Many  of  the  lava  beds  show  a  coarse-grained  ophitic  texture  and  may  be  classed 
as  dolerites,  and  have  been  so  named  by  Hagerman  and  myself  in  our  previous 
papers.  Such  rocks  are  usual  around  Puerto  Frances  and  Bahia  Cumberland  as 
well  as  at  Vaqueria  and  Tres  Puntas.  Specimens  from  all  these  localities  are 
in  outward  appearance  very  much  alike.  In  general  these  rocks,  in  contrast  to 
other  lavas,  are  singularly  fresh.  Only  the  olivine  often  shows  a  dark  brown 
rim,  indicating  an  incipient  alteration  to  iddingsite  {c(.   12,  p.  260). 

The  doleritic  basalts  seem  only  to  occur  at  lower  horizons,  where  the  lava 
beds  generally  attain  their  greatest  thickness.  All  the  specimens  thereof  at  hand 
are  from  between  sea  level  and  200  m.  A  sample  in  Skottsberg's  collection  from 
Bahia  Cumberland  (Fig.  7)  and  another  from  Vaqueria  are  both  from  1 50  m  above  sea 


'  The  name  oceanite  was  first  proposed  by  Lacroix  in  1923  (in  Mineralogie  de  Madagascar, 

Vol.  Ill,    p.  49).    The    name    is  given  "k  cause  de  leur  abondance  dans  les  iles  du  Pac-hque". 

^  Holmes,  Q.  J.  Geol.  Soc,   172,   1916,  p,  231;     Washington,  Am.  J.  of  Sc.  V,  1923.  p.  471- 


rKKCV     (llKN'SEl, 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  47 


Fig.  5.  Basaltic  lava  beds.  Western  head  of  Bahia  Cumberland,  Masatierra.  Photo  C.  Skottsberg. 


level.  The  two  speciinens  are  in  texture  and  mineral  composition  identical.  A 
third  sample  from  Tres  Puntas  is  from  200  m.  Specimens  in  my  earlier  collection 
were   taken    from   the  lower  lava  beds  in  the  quebradas  around  Puerto  Frances. 

As  no  scoriaceous  or  slaggy  development  is  to  be  seen  in  the  upper  or  lower 
parts  of  the  doleritic  beds,  which  have  a  very  coarse-grained  texture,  they  may 
well  represent  intrusions  between  previously  consolidated  lava  flows,  in  accordance 
with  what  Daly  has  assumed  to  be  the  case  with  similar  doleritic  rocks  in  Hawaii 
(R.  Daly,  Differentiation  in  Hawaii.  Journ.  of  Geology,  Vol.  9,    191 1,  p.  291). 

The  doleritic  basalts  and  the  picrite  basalts  have  much  the  same  mineralogical 
composition.  The  only  essential  difference  is  the  higher  content  of  olivine  in  the 
picrite  basalts.  The  other  rock-forming  constituents  in  both  rocks  are  labradorite, 
a  pleochroic  Ti-augite  and  magnetite. 

In  the  picrite  basalts  (oceanites)  from  around  Puerto  Frances  as  well  as  in 
the  olivine  basalts  in  general  the  phenocrysts  of  olivine  are  often  more  or  less 
altered  to  iddingsite.  In  some  cases  only  insignificant  rests  of  olivine  are  left; 
the  iddingsite  pseudomorphs,  however,  still  retain  the  crystal  habit  of  the  olivine. 
To  the  petrogenetic  problem  of  iddingsite  as  representing  a  deuteric  mineral  I 
will  return  later  in  connection  with  equivalent  alterations  in  certain  lava  beds  of 
Masafuera,  where  the  'iddingsitisation'  has  gone  further  and  there  gives  the  rocks 
a  very  characteristic  aspect  (see  p.  60). 

In  the  picrite  basalts  from  Puerto  Frances  inclusions  of  dunite  occur.  The 
large  olivine  crystals  of  this  rock  are  singularly  fresh,   without  an}^  signs  of  even 


48 


PF.RCV    QU  F.N  SEL 


^ 


M^^m^'- 


^;^ 


v!i>;?r: 


> 


Fij,''.  6.   Ficritc  basalt    oceanite).    x  9.  Puerto  Frances,  Masatierra. 


ig.  7-    I  ><'l('riti<  basalt,    x  ^y.   Bahia  Cumberland,  Masatierra. 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEIZ    ISLANDS  49 

a  periferic  alteration  to  iddingsite  which  is  otherwise  usual  in  the  olivine  of  the 
surrounding  basaltic  lavas. 

At  higher  levels  the  basalts  have  a  more  normal  character.  Large  phenocrysts 
of  olivine,  or  olivine  and  feldspar,  lie  in  a  more  or  less  glassy  groundmass  of 
augite,  olivine  and  labradorite  with  abundant  small  grains  of  magnetite,  llmenite 
in  tabular  form  or  in  skeleton  cry.stals  is  now  often  present.  Most  of  these  lavas 
are  vesicular,  scoriaceous  or  slaggy.  The  cavities  are  in  many  cases  rimmed  or 
filled  with  opal. 

The  occurrence  of  basalts  of  this  type  is  widespread  up  to  the  highest  parts 
of  the  island,  and  they  are  without  doubt  the  dominant  lavas  of  Masatierra. 

Somewhat  divergent  lava  beds  seem,  however,  to  predominate  at  intermediate 
horizons.  All  the  specimens  at  hand  of  this  type  are  holocrystalline  rocks  of  an 
ash-grey  colour,  aphanitic  and  aphyric  in  texture  (Fig.  8).  They  generally  show  a 
characteristic  light  zone  of  weathering  and  a  tendency  to  develope  a  columnar  struc- 
ture (Fig.  9).  The  fine-grained  mineral  assemblage  consists  of  augite,  labradorite  and 
very  abundant  magnetite  in  small  euhedral  crystals,  evenly  dispersed  throughout 
the  rock.  llmenite  is  also  generally  present  in  skeleton  crystals.  Stray  small 
grains  of  olivine  may  in  some  cases  be  observed;  in  other  specimens  olivine  is 
completely  absent.  In  vesicular  lavas  of  this  type  the  cavities  are  again  more  or 
less  filled  with  opal  (Fig.  8  b). 

These  lava  beds,  which  represent  the  only  specific  feldspar  basalts  of  Masa- 
tierra, are  found  at  heights  between  400  and  500  m  (Cordon  Chifladores  400  m, 
Portezuelo  500  m).  At  an  elevation  below  100  m  at  Punta  Larga  in  the  more 
western  part  of  the  island  the  same  type  has  been  found,  but  then  in  the  form  of 
a    dike,    which    may    signify  a  channel  for  the  analogous  lavas  at  higher  levels. 

The  very  characteristic  aspect  of  these  rocks  seems  to  indicate  that  they 
represent  a  definite  epoch  of  intrusion,  intermediate  between  the  doleritic  basalts 
and  picrite  basalts  of  the  lower  parts  and  the  scoriaceous  olivine  basalts  of  the 
higher  horizons.  An  analysis  has  been  made  of  a  very  similar  rock  from  Masafuera, 
which  confirms  its  classification  there  as  a  feldspar  basalt  (see  p.  66). 

Lacroix's  four  new  analyses  of  basaltic  lavas  from  Masatierra  indicate  that 
the  analysed  rocks  are  similar  in  composition.  It  is  regrettable  that  the  specimens 
all  originate  from  Bahia  Cumberland.  Probably  the  members  of  the  d'Urville 
expedition  only  brought  back  samples  from  that  locality  and  these  were  there- 
fore the  only  specimens  available  in  Paris  for  Lacroix's  analyses.  On  the  other 
hand  the  insignificant  dip  of  the  lava  flows  on  Masatierra  (i5' — 20°)  may  infer 
that  the  analysed  rocks  can  be  taken  as  representative  in  chemical  composition 
for  the  basal  basalts  of  the  island.  Microscopic  determinations  of  corresponding 
specimens  from  the  other  localities  point  in  the  same  direction. 

With  the  exception  of  the  ultra-femic  picrite  basalts  from  Puerto  Frances 
and  the  inclusions  therein  of  dunitic  rocks,  and  the  light  grey  feldspar  basalts 
of  intermediate  horizons,  we  may  conclude  that  the  dominant  rocks  of  Masatierra 
consist  of  rather  normal  olivine  basalts  deviating  principally  in  texture  and  in  a 
varying  content  of  olivine. 


so 


rKKcV    (UENSKL 


I'"ij(.  H  a.   X'esirular    aphyric   feldspar    basalt    cokminar  structure  . 

(see  Fig-.  9  . 


30.    Portezuelo,   Masatierra 


Fig.  <S  1).  Same  spe(  inien  as  Fig.  8  a,  niagiiified    x  65.    Vesicle  in  upper  right  corner  filled  with 

o})al.    Portezuelo,   Masatierra, 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE  JUAN    FERNANDEZ    ISLANDS  5  I 


Fig  9    Northern    wall    of   the    Portezuelo    Pass,   showing  columnar  structure  of  feldspar  basalt 

CFig.  8  a).  Photo  C.  Skottsberg. 


PERCY    QUKNSEL 


li-. 


uial  weathering  of  basalt.    Bahia  Cumberland,   Masaticrra.  Photo  C.  Skotisbci^;. 


Oiiantities  of  pyroclastic  material  are  found  interbedded  between  the  lava 
Hows  all  over  the  island.  Pohlmann  has  described  such  products  from  Bahia 
del  Padre  as  follows:  "Uberall  sieht  man  Decken  von  basaltischem,  meist  saulen- 
formi^  abj^esondertcm  P'.ruptiv^estein  mit  Schichten  von  vulkanischem  Auswurfs- 
niaterial   wie  Asche,   Lapilli,  l^omben  u.  s.  w.  wechsellagernd. 

Mine  solchc  aus  Lapilli  und  l^omben  zusammengesetzte  Schicht  von  mehreren 
Mctern  Maclitij^keit  an  der  rechten  Seite  der  Bucht  in  der  Nahe  der  sog.  Kapelle 
<;elc^fen,  liefcrt  das  Material  der  sog.  'Glockensteine',  Der  Vorgang  ihrer  l^ildung 
ist,  kurz  gcsagt,  folgendcr:  zuniichst  entstehen  traubige  und  nierformige  Con- 
crctioncn  zwischcn  den  losen  zusammengefiigten  vulkanischen  Massen;  dieseweissen 
Knollen  (JMg.  i  2)  gelangen  beini  Abstiirzen  der  Schichten  an  den  Strand  und  erhalten 
als  Kollstcinc  (lurch  die  Thiitigkeit  des  Wassers  ihre  gerundete  P'orm"  (lO,  p.  321). 

I  ha\e  prc\i<)us]y  noted  that  the  lava  beds  of  the  eastern  parts  of  the  island 
are  also  often   interbedded   with   agglomeratic  layers  (12,   p.  257). 

Some  stra>'  occurrences  of  superficial  tuffs  seem  still  to  have  evaded  destruction 
by  erosion.  Wlu-n  Kcnard  says  that  "amongst  the  specimens  collected  at  Juan 
I'"cnian(iez  (Masaticrra)  !)>•  the  Ciiallenger  l^x|)edition  in  1 875,  we  have  not  found 
any  specimen  which  iniglit  belong  to  any  recent  eruption,  no  tuffs,  no  volcanic 
ash  are  to  be  found  and  e\cr\thing  seems  to  prove  that  they  have  been  washed 
away  by  the  waves  and  the  atmosj)heric  agencies"  (8,  j).  176),  this  last  conclusion 
seems  (juestionable.    1  lagerman   refers  to  two  specimens  in  Skottsberg's  collections 


ADDITIONAL    COMMENTS    ON    THE    OEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  53 

as  representing  recent  tuffacious  material.  The  one  he  describes  as  "ein  poroses, 
dichtes  Gestein  von  El  Puente,  dem  Istmus  zwischen  der  Padrebucht  und  Carbajal 
und  ist  ein  Palagonittuff  mit  einigen  sporadischen  Augit-  und  Magnetitkornern. 
Der  andere  Tuff  stammt  von  dem  nordlichen  Ufer  der  Padrebucht,  von  wo  einige 
umgewandelte  Olivinbasalte  herriihren"  (13,  p.  26).  As  these  specimens  have 
been  collected  without  any  observations  regarding  their  petrological  position,  one 
cannot  draw  any  conclusive  evidence  as  to  their  age,  but  the  specimens  from  both 
the    localities  have  every  appearance  of  being  recent  pyroclastic  sediments. 

I  have  previously  described  rocks  from  Bahia  Cumberland,  filling  out  the 
greater  part  of  the  bay,  which  I  assumed  to  be  of  recent  tuffaccous  origin.  The 
description  was  given  as  follows:  "In  den  zentralen  Teilen  von  Cumberland  Hay 
liegen  noch  ziemlich  machtige,  meist  lebhaft  rot  gefarbte  Tufflager,  die  sehr  stark 
umgewandelt  sind.  Bruchstucke  von  Olivinkristallen,  lapilliartige  Lavabruchstucke, 
P>zkorner  und  Glas  liegen  in  einer  Grundmasse,  die  aus  einer  weichen,  mit  Mes.ser 
schneidbaren,  roten  lateritahnlichen  Lehmsubstanz  besteht,  die  durch  Verwitterung 
aus  dem  ursprlinglichen  Tuffmaterial  hervorgegangen  sein  durfte.  Uberall  in  den 
Tuffsedimenten  ist  eine  deutliche  Lagerung  sichtbar"  (12,  p.  266).  It  is  over  40 
years  since  I  visited  the  locality  and  naturally  I  cannot  now  rely  on  any  personal 
recollection.  The  inundation  in  connection  with  the  volcanic  disturbance  of  1835, 
referred  to  above  (p.  41),  may  have  wrought  such  havoc,  that  superficial  deposits 
could  have  been  re-formed.  But  the  composition  of  the  formation,  as  well  as  my 
notes  from  the  field,  offer  indications  that  the  tuffaceous  material  of  Bahia  Cumber- 
land also  represents  pyroclastic  sediments  of  recent  volcanic  origin. 

Even  if  only  trivial  remains  of  tuffaceous  formations,  indicative  of  late  volcanic 
activity  on  the  island,  are  left,  the  submarine  explosion  of  1835  confirms  without 
doubt  that  the  area  in  the  immediate  vicinity  has  at  that  time  been  subjected 
to  disturbances  of  volcanic  nature. 

Some  rocks  from  around  Bahia  del  Padre  deserve  special  notice.  Schulze 
and  Pohlmann  have  observed  the  deviating  nature  of  the  rock  assemblage  and 
the  latter  has  commented  thereon  as  follows:  "De  suma  importancia  para  esplicar 
la  formacion  jeolojica  de  Masatierra  es  la  entrada  a  Bahia  del  Padre,  situada  en 
la  parte  suroeste  de  la  isla.  Aqui  se  observa  debajo  de  las  capas  basalticas  ya 
descritas  un  grand  macizo  de  roca  compacta  verdosa,  que  es  andesitica.  Segun 
la  opinion  de  Schulze'  que,  a  me  parecer,  es  correcta,  esta  roca  verdosa  es  la 
mas  antigua  de  la  isla.  P^n  ningun  otro  punto,  ni  en  Masatierra,  ni  en  Masafuera 
se  ha  observado  una  configuracion  jeolojica  analoga  a  la  mencionada"  (11,  p.  4). 

The  rocks  referred  to  by  Pohlmann  certainly  show  a  divergent  aspect,  l^ut 
I  do  not  think  they  can  be  taken  as  represehting  rocks  belonging  to  a  more 
ancient  formation  than  the  lavas  around,  nor  that  they  should  be  classified  as 
andesites,  as  Pohlmann  has  assumed.  I  have  as  the  result  of  microscopical 
examination  of  the  rocks  come  to  the  conclusion  that  the  deviating  character  is 
the  result  of  post-volcanic  alterations  through  thermal  processes.  In  my  earlier 
paper  I  have  given  in  some  detail  the  reasons  for  this  conclusion  (12,  p.  266),  and 

'  Ur.  Schulze  died  before  publishing  his  observations. 

4  —  516795     The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.    Vol.  I. 


54 


\'\\<C\     (UKNSKI. 


I      \'<.|. 


llumciatc.    I'.aliia   CumixTland,   Masalicrra.     Photo  C.  Skottsbcrg. 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  55 


Fig.  12.  Lumps  of  magnesite  ;the  so-called  Glockenstein .  Nat.  size.  Rahia  del  Padre,  Masatierra. 
Reproduced  from   P.  Quensel,  12,  p.  269. 


Fig.  13.    Scoriaceous    lava    with    newtonite    and    bole.    Nat.    size.  Bahia  del  Padre,  Masatierra. 
Reproduced  from  P.  Quensel,  12,  p.  269. 


will  in  this  connection  only  summarize  the  facts.  The  rocks  in  question  are  very 
fine-grained  and  of  a  greenish  colour.  The  abundant  cleavage  plains  are  coated 
with  epidote  and  calcite.  Pyrite  is  finely  dispersed  throughout  the  rock.  Under 
the  microscope  one  can  see  that  original  phenocrysts  of  both  olivine  and  augite 
are  more  or  less  completely  changed  to  serpentine  and  chlorite.  The  matrix  con- 
sists of  olivine,  augite,  plagioclase,  magnetite  and  ilmenite  —  i.e.,  the  usual 
mineral    assemblage   of  the  basaltic  lavas.    The  greater  part  of  the  silicates  are. 


I'KRiV     (JlKNSKL 


li()uc\cr,  \cr\-  nuich  altered  to  serpentine,  chlorite  and  prehnite.  Secondary  albite 
and  some  scapolite  is  also  j)resent  toj^ether  with  abnndant  pyrite.  Small  cavities 
are  tilled  with  epidote  ami  calcite.  in  the  slaj^i^y  lavas  in  the  immediate  vicinity 
lai\L;er  ca\  ities  are  tilled  with  a  j)iirel\-  white  cla>--like  substance  which  was  found 
to  be  newtonite.  In  intimate  association  with  this  mineral  is  nearly  always  found 
a  vellowi^ii  waw  substance  which  shows  ever\'  resemblance  to  what  mineralogi- 
call\-  ma\-  be  si^nitied  as  bole  (I''i<;.  13).  It  is  very  brittle  with  a  conchoidal 
fraclme.  in  water  it  readil\'  disintegrates  into  small  angular  fragments.  Tliis  bole 
mineral  is  also  t'ound  tilling  cracks,  or  occurs  in  smaller  masses  in  the  cavities 
of  the   basalts  around   the   baw 

in  t!ie  agglomeritic  lavas  of  the  vicinit)-  large  cavities  are  filled  with  hard 
compact  magnesite.  As  earl\-  as  1886  Darapsky  described  this  mineral  by  the 
r.ame  "( ilockenstein"  and  gave  an  analysis  thereof,  which  shows  it  to  be  an 
exceptionall)-  pure  magnesite  (9,  p.  I  13).  Without  doubt  the  magnesite  is  primarily 
derued  tVom  olivine,  the  decomposition  being  caused  by  the  same  processes  as 
ha\e  changed   the   basaltic  lavas  nearby. 

The  whole  asj^ect  of  the  rocks  from  l^ahia  del  Padre,  with  magnesite,  calcite, 
serpentine,  chlorite,  sca{)olite  and  pyrite  as  secondary  minerals,  seems  without 
doubt  to  indicate  that  the  lava  beds  in  question  have  been  subjected  to  alterations 
in  connection  with  thermal  processes  during  some  intermediate  phase  of  volcanic 
activity  on  the  island.  There  seems  no  reason  to  classify  them  as  andesites  of 
an   older   formation,  as  is  done  by  Schulze  and  Pohlmann. 

In  a  s[)ecimen  collected  by  Skottsberg  from  the  shore  south  of  El  Yunque, 
Ilagerman  also  tound  evidence  of  a  far-reaching  decomposition.  Under  the  heading 
"!  Ixclrothermale  I^ildungen"  he  gives  the  following  description:  "Das  Handstiick 
ist  ein  von  weissen  .Streifen  durchzogenes  scharfgriines  Gestein,  das  u.  d.  M. 
grosse  Augitkristalle  in  einer  vollig  zeri^ossenen  Serpentinmasse  zeigt.  Das  Pra- 
))arat  ist  von  Aragonitbandern  durchzogen.  Dieses  Gestein  muss  als  ein  stark 
umgebildetes  (  )livinfels  bezeichnct  werdcn"  (13,  p.  27).  The  large  olivine  crystals  of 
the  duniti'.  tound  as  inclusions  in  the  picrite  basalt  at  Puerto  Prances,  show,  on  the 
contrarx-,  no  signs  of  secondar\-  alteration  (see  p.  47).  There  is  therefore  no  doubt 
that  the  higiily  decomposed  '( )hvinfels'  described  by  Hagerman  has  succumbed 
to  a  later  decomposition  of  much  the  same  nature  as  has  been  active  around 
I)ahia    (K-1    I'adre. 


Masafuera. 

I. at.   ^y   52'  S..    Lon^.  <So''  54'  W. : 

1  he  la\as  ot  Masafuera  piesent  a  more  varying  aspect  than  those  of  Masa- 
tierra  and  contam  .sc\cral  t>pes  of  petrographic  interest.  They  have,  however, 
hitherto  only  been  sinumanl\-  described  in  the  j)a|)ers  by  myself  (l  2,  p.  274)  and 
b\'    I  lagerman   1  1  3.   p.  j8i. 

I  he  rocks  which  predominate  at  lower  levels  are  mostly  vesicular  to  slaggy 
basalts.  The\-  are  well  represented  around  the  Ouebrada  de  las  Casas.  At  higher 
levels  the   basalts  consist  of  more  compact  lava  beds.    In  contrast  to  the  basalts 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  57 


QwebradxvNegrcb 
Q  Saitdahto 
Q  SandaJLo 

QuiebraSa.  PcUo 


Chx>%<ta 
QiLeibrajdcLiiano 


Ra3a  de-la,  CoLonicL 
Casae  o  Banuedxxnjo 
Ctuca. 

Q.BUndaSo 


Q  Cdbreros 


P*«-Negra. 


Cables  10 

(§065  f*')  ■ 


3  SeaMiles 


Fig.  14.   Masafuera.   From   the  Chilean  Gov.  chart  of  1921.   Scale   1:70000. 


S8 


)r  F.N  SKI. 


Fig.  15.  \- 


.t    !li 


nicrior  of  Quel^rada  do  la  Loberia  vieja  towards  the  shore. 
Photo  C.  SkottsberL--. 


of  Masatierra,  olivine  is  more  scarce  in  the  dominant  "basalts  of  Masafuera,  but 
is  sin(^tilarly  rich  in  the  basaltic  dikes  which  in  hundredfold  vertically  traverse 
the  island. 

The  rocks  |)re\ailin<^  at  heights  up  to  c.  i  ooo  m  may  for  the  greater  part 
be  classihed  as  dark  vesicular  feldspar  basalts,  with  large  phenocrysts  of  labradorite 
(Ab  45,  An  55).  Olivine  is  scarce  as  phenocrysts  but  occurs  in  varying  amount 
in  the  groundmass  together  with  augite,  labradorite  and  magnetite  in  a  dark, 
glassy    matrix   (V\i^.  16). 

In  many  sj)ecimcns  of  the  vesicular  lavas  the  vesicles  are  filled  or  lined  with 
zeolite  minerals.  In  some  samples  the  \esicles  present,  from  the  outward  rim, 
fust  a  coating  ot  glass,  followed  by  chalcedon  and  chabasite  and  a  central 
replenishment  of  well   develoj)ed   natrolite  spherolites. 

At  an  elevatif)n  of  about  i  ooo  m  a  lava  bed  of  a  \ery  ditTerent  aspect  is 
met  witii.  In  contrast  to  the  dark  basalts  of  lower  horizons,  the  rock  now  in 
(jiiestion  is  light  gre\-  in  colour.  Large  j)henocrysts  of  olivine  (up  to  5  mm  in 
diameter)  and  labrad'jrite  are  tmiformly  distributed  in  a  very  fine-grained  ground- 
mass,  consistmg  of  augite,  slender  laths  of  labradorite,  abundant  small  grains  of 
magnetite  and   a  small   amount  of  a   nearly  colourless  glass  (Fig.   17). 

In  outward  appearance,-  this  lava  has  a  very  singular  asj:)ect.  The  surface 
teels  rough  and  grain\-  which,  together  with  the  light  grey  colour,  at  first  gives 
the  impression   that   the  rock   would   ha\e  a  trachytic  or  trachy-andesitic  compo- 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  59 


Fig.  16.  Vesicular  feldspar  basalt,   x  8.  Normal  development  under  i  000  m  elevation.  Masafuera. 


sition.  The  analysis  (Table  I,  No.  7),  however,  shows  that  the  lava  must  be 
assigned  to  the  basaltic  group,  though  in  many  respects  it  obviously  differs  from 
all  the  dark  basaltic  rocks  of  lower  elevations  on  the  island.  With  some  hesitation, 
in  my  former  publication  I  referred  the  rock  to  the  basanite  group,  assuming 
that  a  content  of  virtual  nepheline  might  be  concealed  in  the  groundmass,  which 
was  readily  affected  when  treated  with  HCl.  This  circumstance  Lacroix  assumes 
to  be  characteristic  for  the  basanitoid  lavas  of  the  Pacific,  where  he  says  the 
nepheline  often  "n'est  pas  exprimee;  elle  est  restee  a  I'etat  potential  dans  la  verre. 
La  poudre  des  ces  roches,  traitee  par  HCl  s'attaque  plus  au  moins  facilement" 
(C.  R.  169,  1919,  p.  402).  Bartii  has,  in  his  paper  'Pacificite,  an  anemousite 
basalt',  suggested  that  the  virtual  nepheline  may  be  concealed  in  the  plagioclase 
in  the  form  of  carnegieite.  For  such  basalts  he  proposes  the  name  pacificite,  or 
if  olivine  is  present,  olivine  pacificite  and  on  account  of  my  former  description, 
refers  the  lava  in  question  from  Masafuera  to  the  pacificite  group  (21,  p.  60; 
22,   p.  380,   510). 

In  publishing  the  description  of  the  rocks  from  the  Juan  Fernandez  Islands 
in  my  earlier  publication  the  analyses  were  not  recalculated  according  to  modern 
methods.  This  has  been  done  now  and  the  results  recorded  in  the  appendixed 
table  of  analyses.  With  regard  to  the  rock,  then  tentatively  named  basanilc,  the 
calculated  norm  shows  no  nepheline  and  an  excess  of  1.99  %  of  aluminium.  Ihis, 
together  with  a  content  of  3.36  %  H2O,  gives  the  impression  tiiat  the  rock  is  de- 
composed. On  this  account  Lacroix,  in  reproducing  my  anal  jos  ■::  ilie  Juan 
Fernandez    rocks,    omitted  this  analysis,  "car  les  resultats  ne  curresi-   nclcr.t  pas 


6o 


rF.RCV    ()L-F.NSKL 


17.   (  )li\  inc  h; 


lit.   Nat.  size.  Elevation  c.  i  000  in.   Masafuera.  The  olivine  plienocrysts  more 
or  less  completeh-   altered   to   deuteric  iddingsite. 


:i  la  composition  mineralooique  clccrite;  le  calcul  fait  voir  qu'il  s'a^^it  d'une  roche 
altcrce"     ( 14,  |).  65). 

A  reiiewetl  scriitin}-  of  the  rock  has  offered  new  aspects  regardin<^  its  petro- 
looic  and  j^etro^raphic  position.  The  essential  point  has  been  that  the  large  olivine 
phenocrwsts  are  altered  to  such  a  great  extent  to  iddingsite.  We  have  every  inter- 
mediate phase  frotii  a  more  or  less  broad  rim  of  iddingsite  around  a  kernel  of  per- 
fectly fresh  ()li\  ine  to  c()mj)lete  pseudomorphs  of  iddingsite,  retaining  the  crystal 
habit  of  the  olivine.  The  circimistancc  that  the  kernels  of  olivine  are  absolutely  un- 
altered, leads  to  the  conclusion  that  the  iddingsite  is  not  a  product  of  normal 
weathering  but  a  deuteric  mineral,  derived  during  a  final  stage  of  cooling  of  the 
la\a   in    which    it  occiu's. 

l)iM'ing  later  }'ears  sexeral  papers  have  been  published,  in  which  the  origin 
ol  iddingsite  has  been  discussed  in  detail.  The  conceptions,  there  j:)ut  forth,  to 
all  intent  conform  with  the  characteristic  features  of  the  lava  from  iMasafuera. 
As  the  petrogcnesis  of  the  rock  in  (juestion  is  of  a  certain  interest,  some  signi- 
ficant (|uotatioiis  may  be  given,  relating  to  the  formation  of  iddingsite  from 
ditk-rrnt    localities. 

I\i»ss  and  .Shannon  simimarize  their  conclusions  as  follows:  "Iddingsite  is 
not  confmed  to  weathered  surfaces;  its  development  shows  no  proximity  to  joint 
cracks  and  e\  idences  of  weathering  in  associated  minerals  is  entirely  absent.  Thus 
it  is  concluded  that  iddingsite  is  not  a  product  of  ordinary  weathering  but 
a  deuteric  mineral,  tiiat  is  to  say  the  result  of  metasomatic  {)rocesses  associated 
with  the  later  stages  (jf  a  cooling  magma."  They  also  emphasize  that  the 
magma  must  ha\e  come  to  rest  before  iddingsite  formed,  for  though  it  is  a 
very  brittle  mineral,  it  is  ne\er  fractured  or  distorted  by  flow  (C  S.  National 
Museum,    l*roc.  6-,    1925,   Art.  7,   p.  i<S). 

Al  Korssi.Ai  comes  to  the  conclusion  that  "iddingsite  is  the  result  of  oxida- 
tion  [)rocesses  that  acted  rapidly  on   the  olivine  during  the  liberation  of  copious 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  6 1 

active  volatile  phases  at  the  time  of  eruption"  (Linnean  Soc.  of  X.  S.  W.,  Proc.  51, 
1926,  p.  617). 

Finally  Edwards  in  his  paper  'The  formation  of  iddin^site'  says:  "It  is 
concluded,  therefore,  that  iddingsite  forms  during  or  after  extrusion,  according 
to  the  temperature  of  the  magma  at  the  time  of  extrusion,  and  that,  if  the  magma 
has  cooled  sufficiently  before  extrusion,  not  enough  iron  oxides  are  left  in  the 
residual  volatiles  for  iddingsite  to  be  formed"    (Am.  Min.   23,    1938,  j).  281). 

The  formula  for  iddingsite  is  given  by  Ross  and  Shannon  as  MgO  •  FegOa  • 
3  Si02-4  H2O  with  magnesia  replaced  in  part  by  CaO,  approximately  in  the  ratio 
I  :  4.  The  calculated  composition  according  to  this  formula  includes  7.90  %  HaO"*" 
and  7.90  %   HaO"   (ibid.  p.  17). 

All  the  authors,  now  quoted,  come  to  the  same  conclusion  that  the  forma- 
tion of  iddingsite  cannot  be  regarded  as  the  result  of  normal  weathering  processes 
but  that  the  mineral  must  be  of  deuteric  origin,  formed  during  the  later  stages 
of  a  cooling  magma.  The  development  of  abundant  iddingsite  in  the  lava  flow 
from  Masafuera  leads  to  the  same  conclusion.  I  may  also  remark  that  most  of 
the  photomicrographs,  representing  different  stages  of  alteration  from  olivine  to 
iddingsite,  published  in  PI.  I — II  in  Ross  and  Shannon's  paper,  correspond  in 
nearly  every  detail  with  equivalent  alterations  in  the  lava  from  Masafuera  (Fig. 
18).  As  iddingsite  according  to  the  formula  of  Ross  and  Shannon  contains 
15.80%  H2O,  we  may  conclude  that  an  appreciable  amount  of  the  3.46%  H2O, 
according  to  the  analysis  of  the  lava  in  question  (Table  I,  No.  7),  enters  the 
iddingsite  molecule. 

Another  singular  alteration  in  the  iddingsite  basalt  in  question  is  also  worthy 
of  notice.  The  slender  labradorite  laths  in  the  groundmass  as  well  as  some 
phenocrysts  of  the  same  mineral  are  found  to  be  more  or  less  completely 
altered  to  an  isotropic  substance,  though  retaining  their  crystal  habit  perfectly 
intact.  I  had  observed  this  circumstance  in  my  former  publication  on  the  Geology 
of  the  Juan  Fernandez  Islands  (12,  p.  278)  and  then  came  to  the  conclusion  that 
the  invading  material  was  glass,  formed  through  fusion  of  the  feldspars  in  the 
volcanic  vent   (Fig.  19). 

Dr  Otto  Mellis  has  kindly  undertaken  a  renewed  investigation  of  this 
isotropic  alteration  product.  At  first  he  also  came  to  the  conclusion  that  it  con- 
sisted of  glass,  formed  by  fusion  of  the  feldspars  during  a  period  of  high-temperature 
autometamorphism.  A  more  detailed  study,  however,  led  to  different  conclusions. 
Dr  Mellis  gives  the  result  as  follows:  "Repeated  determinations  showed  that  the 
refractive  index  of  the  isotropic  material  lies  between  1.49  and  1.50.  These  values 
must  exclude  the  presence  of  glass,  formed  by  fusion  of  a  labradoritic  feldspar. 
The  index  of  refraction  should  according  to  Franco  and  Schairer  in  such  a  case 
lie  between  1.53  and  1.54  (Journ.  of  Geology,  Vol.  59,  1951,  p.  266).  A  further 
evidence  in  this  respect  is  that,  when  treated  with  HCl,  the  isotropic  matter 
readily  gelatinized,  which  is  not  in  accordance  with  what  one  can  assume  to  be 
the  case  with  glass,  formed  by  fusion  of  a  basic  plagioclase.  Opal  must  for  the 
same  reason  be  excluded  as  a  possible  component.  Continued  investigations 
resulted    in    the   conclusion  that  the  alteration  of  the  feldspars  ir   the  iddingsite 


VFAiCX    ()rKNSKI. 


l•'i;^^  i8.  hklinj^site  forming   sharp  outer  borders  around  unaltered  oli\inc.     <   loo.  From  specimen 
Fig.  17.   Elevation    i  000  m.   Masafuera. 


aV,  ^*,.. -< 


fy— 


r 


^^f^  JSr^^. 


y 


Fij^.  19.   I'henocryst   of  labradorite,  traversed  In'  a  zeolitic  matter,    x   170.   From  specimen 

I-"i<^.   17.   Masafuera. 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  65 

basalt  must  be  attributed  to  an  invasion  of  zeolitic  matter.  As  there  seems  to 
be  a  slight  difference  in  refractive  indices  in  the  alteration  product,  this  is  prob- 
ably a  mixture  of  different  zeolites  or  related  material.  It  has,  however,  not 
been  possible  to  determine  the  definite  nature  of  the  zeolites.  The  fact  that  they 
show  no  sign  of  birefringence  indicates  that  they  must  belong  to  the  isotropic 
group  or  to  those  of  very  near  zero  birefringence." 

It  is  of  interest  to  note  that  Renard  evidently  has  observed  a  very  similar 
circumstance  in  a  lava  flow  from  Masatierra,  when  he  says  that  sections  of  a 
basic  feldspar  from  a  greyish  very  scoriaceous  lava  under  the  microscope  are 
seen  to  be  "cracked  and  pervaded  with  zeolitic  matter,  which  forms  an  irregular 
network.  This  matter,  which  looks  slightly  grey,  when  seen  in  ordinary  light, 
remains  obscured  between  crossed  nicols."  This  description  might  as  well  refer 
to  the  zeolitisation  of  the  lava  bed  from  Masafuera,  especially  as  Renard  in  the 
same  lava  mentions  olivine,  uniformly  changed  to  iddingsite  (Renard  says  hematite, 
8,  p.    176). 

It  therefore  seems  probable  that  lava  beds,  in  which  the  content  of  olivine  has 
been  more  or  less  changed  to  deuteric  iddingsite,  also  have  succumbed  to  a  high 
degree  of  zeolitisation.  The  fact  that  the  large  phenocrysts  of  olivine,  only 
rimmed  with  iddingsite,  show  no  signs  of  alteration,  whereas  the  feldspars  in  the 
same  section  may  be  completely  altered  to  an  isotropic  substance  of  zeolitic 
composition,  must  indicate  that  also  the  zeolitisation  of  the  rock  has  taken  place 
at  an  early  stage  and  cannot  be  attributed  to  periods  of  normal  w^eathering  or 
later  thermal  activity. 

In  consequence  of  the  changes  in  chemical  composition  which  must  have 
taken  place  in  connection  wTth  the  formation  of  iddingsite  and  the  zeolitisation 
of  the  lava,  it  is  no  longer  possible  to  establish  the  primary  composition  of  the 
rock  in  question.  Though  the  whole  aspect  of  the  lava  bed  so  obviously  differs 
from  the  normal  olivine  basalts  on  Masafuera,  there  is  evidently  not  reason  enough 
with  any  degree  of  certitude  to  classify  the  rock  as  an  olivine  basanitoid.  The 
presence  of  alkaline  rocks  on  Masafuera  must  for  the  present,  therefore,  be 
restricted  to  the  occurrence  of  the  soda-trachytic  lava  flows,  now  to  be  described. 

At  an  elevation  between  i  ooo  and  i  lOO  m  there  occur  rocks  of  quite  another 
type.  In  outward  appearance  they  are  dense,  aphanitic  and  aphyric  in  texture 
and  of  a  light  yellowish  green  colour.  The  dominant  minerals  are  an  acid  plagio- 
clase  and,  subordinate,  orthoclase.  Olivine,  a  light  green  diopsidic  pyroxene,  and 
ore  minerals  are  sparingly  found  in  small  individuals  between  the  feldspar  laths, 
which  have  a  well-defined  trachytoid  orientation  (Fig.  20).  An  analysis  of  this 
rock  was  made  for  my  former  description  and  is  reproduced  in  the  annexed  table 
of  the  analyses  of  the  Juan  Fernandez  rocks  (Table  I,  No.  8).  I  named  the  rock 
soda-trachyte. 

On  my  survey  I  had  assumed  that  rocks  of  this  composition  only  existed 
on  and  formed  the  highest  parts  of  the  island,  as  at  that  time  no  higluM'  elevations 
were  reached.  New  specimens  in  Skottsberg's  later  collection  di^::o\  e  my  earlier 
supposition  in  two  essential  respects.    In  the  first  place,  rock.-,  of  mi  ch  the  same 


^'4 


I'KIUN     ()rKNSEL 


Fig.  20.  Sotla   trachyte.     :■:  50.  Elevation  c.   I  050  m.  Masafuera. 


Iiytc    boulder.     •    30.    Xic.  +.  Elevation  about  400  m. 
Ticrras   lUancas,  Masafuera. 

character  as  the  .soihi  trachyte  have  now  been  found  at  lower  levels.  Secondly, 
Skottsbet-o  reached  the  .siniiniit  of  the  i.sland  and  was  there  able  to  establish  that 
basaltic  lavas  ()cciij)y  tlie  whole   highest  i)art  of  the  island. 

The    new    find    of    a  rock,   closely  related  to  the  soda-trachyte  from  higher 
levels,  was  a  loose  boulder  at  the  foot  of  Ticrras  Blancas.   According  to  Skotts- 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE  JUAN    FERNANDEZ    ISLANDS  65 


F\g.  22.  Vesicular  phyric  feldspar  basalt  with  stray  phenocrysts  of  olivine, 
cimen  from   i  420  m  elevation.  Masafuera. 


30,  Analysed  spe- 


ij.  Aphyric  feldspar  basalt. 

El  Ovalo,  Masafuera. 


66  \'V\iC\     (JUKNSKL 

berg's  intorniation,  boulders  of  tliis  character  were  ver}'  numerous  in  the  talus 
under  a  steep  escar|)nient  and  can  here  hardlx'  ha\e  originated  from  higher  eleva- 
tions tlian   about  400  ni. 

In  outwarti  a[)|)earance  this  rock  shows  an  aspect  different  to  the  compact 
soda  tiachxte  i)et\\een  I  000  and  i  loom,  which  on  account  of  its  dense  texture 
at  first  sii;ht  i^ives  the  impression  of  a  (piart/itic  rock.  The  new  specimen  has 
on  the  other  hand  a  roui^h.  loose  grained  structure.  Minute  inter^jjranular  cavities 
in  the  rock  are  seen  under  the  microscope  to  be  tilled  with  limonitic  matter.  But 
allowing  for  tiiis  structural  dissimilarity  the  rocks  from  the  talus  below  Tierras 
Hlancas  seem  to  l)e  closel}'  related  to  the  soda-traclu'te  as  described  above  (Fig.  21). 
Taking  into  consideration  the  seemingl}-  horizontal  j)()sition  of  the  lava  beds  of 
-Masafuera.  the  conclusion  must  be  that  la\as  of  soda-trachytic  composition  have 
been  emittetl  at  different  times  and  that  they  have  alternated  with  lavas  of  more 
normal   basaltic  comj)()siti()n   (see  j).  71). 

.S|)ecimens  of  the  rocks  from  between  i  100  and  i  400  m  j)resent  again  a 
different  asj^ect.  I'^ive  sam|)les  from  I  lOO,  I  200,  I  300  and  I  420  m  all  have 
\er\-  much  the  same  a})pearance.  The}'  are  all  phyric  lavas  of  an  ash-grey  colour, 
i-'eldspar  and,  more  rarel}-,  olivine  occur  as  ])henocrysts  in  a  groundmass  of 
basic  plagioclase,  augite  and  abundant  magnetite.  In  a  somewhat  vesicular  specimen 
from   I  100  m  olivine  is  also  present  as  sporadic  grains  in  the  groundmass. 

The  groundmass  has  in  most  of  the  specimens  a  trachytic  texture;  the  narrow 
laths  of  feldsjjar  circuiting  the  phenocrysts  in  a  more  or  less  well-defined 
manner    (I'^ig.  22). 

In  m\-  previous  paj)er  I  named  the  rocks  of  this  nature  trachyandesite. 
Ilagerman  sa\-s,  referring  to  much  the  same  specimens  in  Skottsberg's  collection: 
"Die  Klassifi/ierung  hiehergehdriger  Gesteine  ist  etwas  unsicher.  Geniigende 
Grimde  sie  als  Trach\-andesite  zu  bezeichnen,  liegen  jedoch  nicht  vor"  (13,  p.  30). 
I  iagerman   names  the  rock  andesite  in   his  j)aper. 

Tliese  rocks  evidently  have  a  rather  wide  distribution  on  ^lasafuera,  j^robably 
lorming  the  whole  comi)lex  of  la\a  beds  between  i  100  and  i  420  m.  To  certify 
their  j)etrographic  ])osition  an  analysis  has  now  been  made  of  a  typical  specimen 
(  Tabk-  I,  No.  5).  The  calculated  norm  and  Xiggli  values  definitely  classify  the 
rock  as  a  tel(lsj)ar  basalt,  in  many  respects  of  similar  composition  as  some  of 
tile  analxsed  basalts  from  Masatierra  (Table  I,  \o.  i — 4).  A  difference  of  interest 
is  the  still  lower  content  of  KoO;  the  Xiggli  value  X'  is  now  0.08  against  O.I  5 — 0.20 
in    the  basalts    fiom    Masatierra. 

I  hese  phyric  feldspar  basalts  from  ^lasafuera  in  general  aspect  also  very 
much  resemble  the  apii\ric  fel(lsj)ar  basalts  from  between  400 — 600  m  elevation 
on  Masatierra.  (  )ccasionall\-  at  lower  levels  they  also  on  Masafuera  may  be  aphyric 
in  texture  il'ig.  23).  dhey  seem  also  in  several  respects  to  correspond  to  certain 
ph\ric  and  aph\ric  basalts  from  the  I  lawaiian  Islands,  as  described  by  Washinotox 
(Petrology  of  the  I  lawaiian  Islands.  I.  Kohala  and  Mauna  Kea.  Am.  J.  of  Sc.  V, 
'9-3'  P-  4''^"*-  ^  '>  this  (juestion  I  will  return  under  a  concluding  heading  dealing 
with  the  chemical  and  petrological  connections  between  the  rock  of  the  Juan 
I-'ernandez   Islands  and   other   volcanic   islands  of  the  eastern   Pacific. 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  67 


Fig. 


•4.  Olivine  basalt,  supersaturated  with  iron  oxide,   x  35.  Vesicular  flow-breccia.  Los  Inocentes 
^elevation  c.  i  500  m),   Masafuera. 


Fig.  25.  Olivine    phenocryst    from   specimen  Fig.  24.     x   no.    Specks    and 
olivine  as  rests  in  a  pseudomorph  after  olivine.    Los  Inocentes. 


cak^  <:i  tUKiiLcred 


6S 


PKKCV    ()1-i:nski. 


I"ij4.  26.   Disintegrated    phenocryst    of  olivine,  caused  by  high  temperature  oxidation.    The  rim 

around  the  olivine  crystal  is  hematite.  Within  the  olivine  the  light  grains  are  mineral  components, 

formed  by  exsolution.     <   150.  Los  Inocentes,  Masafuera.  Photo  P.  Ramdohr. 


Pig.  27.   I'art  of  disintegrated  olivine  |)heno(  ryst,  enlarged    x  600.  Against  the  dark  background 

of    olivine    the    exsolution    proiUn  t  is  seen   in   the  form  of  small  composite  grains  consisting  of 

hematite    light    and  a  darker  undefined  component.  Larger  lij^ht  grains  see  PI.  II,  P^ig.  I. 

Photo   P.  Ramdohr. 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE  JUAN    FERNANDEZ    ISLANDS  69 


Fig.  28.  High-temperature  exsolution  of  spinel,  enclosed  in  olivine  phenocryst  (PI.  I,  Fig.  2).  Light 
lamellae,    according  to    Ramdohr    probably   magnetite  in  groundmass  of  excess  spinel,     x  600. 

Photo  P.  Ramdohr. 


Fig.  29.    Magnetite  in  groundmass  of  the  lava  bed  as  seen  in  PI.  I,  Fig.  i    pa:t-a!iy  c!):inge(l  to 
hematite  (martite).    x  300.   Photo  P.  Ramdohr, 

5  —  516795     The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.    Vol.  I. 


■JO  I'KKCN     (JIKN'SKI. 

An  observation  of  importance  with  rej^ard  to  the  distribution  of  different 
lava  beds  on  Masafuera  is  tlie  find  of  a  deviatini^  type  of  olivine  basalt  from  Los 
Inocentes.  Accordin*;  to  Skottsber^'s  observations  such  la\as  probably  occupy 
the  hii^diest  part  of  the  island,  representing^  elevations  above  I  420  m.  The  few 
samples  brouj^ht  back  are  hi<4hl\'  scoriaceous  flow  breccias  containing  a  high 
content  of  iron  oxides  (h'ig.  24).  Hagerman  has  in  the  preceding  publication 
of  this  series  given  a  descrij)tion  of  these  lavas  which  he  characterizes  as  slaggy 
ohvine    basalts    with    large    olivine    phenocrysts,    supersaturated    with    magnetite 

(•3.   V-  ^S'^- 

'l"he  phenocrysts  of  olivine  are  under  the  microscope  found  to  be  almost 
opacjiie,  due  to  the  {precipitation  of  new-formed  ore  minerals.  A  varying  amount 
of  residual  olivine  is,  however,  nearly  always  to  be  observed  in  the  form  of  specks 
or  streaks  (l^'ig.  25).  \o  signs  of  alteration  are  to  be  observed  in  this  olivine. 
Optical  determinations  indicate  that  only  a  low  content  of  about  8%  FeO  is 
present   in   the  molecule. 

The  groundmass  consists  of  slender  laths  of  labradorite,  small  grains  of  augite, 
magnetite,  ilmenite  and  pseudobrookite  in  a  dark  brown  glass  matrix. 

To  determine  the  mineral  composition  of  the  pseudomorphs  after  olivine 
Professor  S.  Gavklin  and  Dr  Uv'i"KNlM)GAARl)'r  kindly  undertook  to  examine  some 
polished  sections  of  the  rock.  Professor  P.  Ramdoiir  (Heidelberg)  contempora- 
neously supervised  a  section  for  the  same  reason.  It  thereby  became  apparent 
that  the  seemingly  opacpie  constituent  was  not  magnetite  and  that  the  mineral 
assemblage  of  the  pseudomorphs  was  of  a  complicated  nature.  Professor  Ramdohr 
has  taken  four  photomicrographs  thereof  and  kindly  put  them  at  my  disposal. 
They  are  reproduced  in  Fig.  26 — 29  with  Ramdohr's  explanatory  notes.  In  Fig.  26 
the  essential  components  can  be  observed.  A  rim  of  hematite  is  seen  to  encircle 
an  i(liom()r|)hic  crystal  of  olivine  with  specks  of  disintegrated  minerals.  In  the 
enlarged  microphotograph  Fig.  27  these  minerals  are  seen  in  the  form  of  small 
lighter  grains  uniformly  distributed  against  the  dark  background  of  olivine.  One 
can  now  observe  that  the  grains  consist  of  two  constituents.  The  one  component 
is  hematite.  Repeated  attempts  have  been  made  to  determine  the  second  com- 
ponent both  in  polished  sections  and  with  X-ray  powder  photographs.  No  con- 
clusive evidence  regarding  the  true  nature  of  this  mineral  has,  however,  been 
attained. 

The  singular  alteration  of  the  olivine  phenocrysts  must  in  all  probability  be 
connected  with  the  saiue  processes  as  have  controlled  the  formation  of  the  deuteric 
iddingsite,  though  in  the  samj)lcs  at  hand  this  mineral  is  not  extant.  P:dwards 
seems  to  have  described  a  very  similar  formation  in  the  iddingsite-bearing  basalts 
from  two  X'ictorian  localities  in  Australia.  After  concluding  that  the  iddingsite 
must  have  been  formed  before  the  ultimate  consolidation  of  the  lava  flow,  Fdwards 
says:  "In  some  instances,  however,  the  action  has  gone  further,  and  a  rim  of  iron 
oxid  is  formed  on  the  outer  margin  of  the  iddingsite.  Fventually  all  the  original 
olivine  vanishes,  and  the  iddingsite,  which  had  formed  a  rim  about  it,  is  completely 
rej)laced  by  magnetite  ...  It  is  essential  for  the  formation  of  iddingsite  that  the 
magma  should  not  only  be  rich  in  water  vapour,  but  that  it  should  have  differentiated 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE  JUAN    FERNANDEZ    ISLANDS  7  I 

in  such  a  manner  as  to  give  rise  to  an  iron-rich  final  fluid"  (Am.  Min.  23,  1938, 
p.  280).  A  photomicrograph  in  the  text  shows,  according  to  Edwards,  "a  pseudo- 
morph  of  iron  ore  after  iddingsite,  itself  a  pseudomorph  after  olivine". 

These  suppositions  seem  also  referable  to  the  very  similar  formation  on 
Masafuera.  The  lava  bed  in  question  contains  33.86  %  FeO  according  to  deter- 
mination by  Landergren  (13,  p.  33).  This  is  more  than  double  the  content  of 
FeO  in  any  other  basalts  of  the  Juan  Fernandez  Islands.  There  seems  little  doubt 
that  the  pseudomorphs  in  question  have  formed  under  much  the  same  conditions 
as  advanced  by  Edwards.  This  is  furthermore  supported  by  the  nearly  identical 
appearance  of  the  photomicrographs  of  the  pseudomorphs  from  the  Australian 
localities  and  from  Masafuera,  in  both  cases  in  connection  with  lavas  containing 
iddingsite.  Edwards  assumes  that,  via  an  intermediate  stage  of  iddingsite,  the  ulti- 
mately formed  component  is  magnetite.  As  narrated  above,  this  is  not  the  case 
in  the  lava  from  Masafuera,  where  the  pseudomorphs  are  found  to  be  of  a  more 
complicated    composition,    formed   without    iddingsite    as   an  intermediate  phase. 

All  the  observations  given  above  seem  to  indicate  that  the  iddingsite-bearing 
basalt  from  elevations  about  i  000  m  as  well  as  the  flow  breccias  from  the  highest 
elevations  on  Masafuera  have  been  subjected  to  an  automorphic  re-mineralisation, 
prior  to  the  final  consolidation  of  the  magma.  This  would  suggest  that  the  volcanic 
eruptions  have  been  interposed  by  periods,  during  which  the  lava  in  a  molten 
state  has  temporally  stagnated  in  the  volcanic  vent  under  conditions  which  in 
connection  with  active  volatile  phases  have  led  to  the  formation  of  such  deuteric 
minerals  as  iddingsite  and  to  the  partial  high  temperature  exsolution  of  the  oli- 
vine phenocrysts. 

The  conclusions  which  may  be  drawn  regarding  the  distribution  of  the 
different  lavas  of  Masafuera  would  be  now  that  rocks  of  more  normal  basaltic 
composition,  principally  feldspar  basalts,  occupy  the  lower  and  intermediate  eleva- 
tions of  the  island,  whereas  the  highest  parts  consist  of  an  olivine  basalt,  super- 
saturated with  iron  oxides.  At  some  intermediate  elevations  lavas  of  a  more 
alkaline  character  are  found  in  the  form  of  at  least  two  beds  of  soda-trachyte, 
interposed  between  dominant  flows  of  basaltic  lavas. 

Horizontal  distribution  of  different  types  of  lava  on  Masafuera. 


in  m    X  100 


o      I       2      3      4      5      6      7      8      91011121314   1500 


Olivine  basalt  supersaturated 
with  magnetite 

Light  grey    phyric  feldspar 
basalt 

Soda-trachyte 

Iddingsite-bearing  phyric  oli- 
vine basalt 

Dark     phyric     and    aphyric 
feldspar-  and  olivine  basalt 

5*— 516795 


X 

X 

X 

X 

X 

X 

X 

X 

v 

X 

I'F.KfV    (JUFNSF.L 


Fi^^  30.   Picrite  basalt  (masafuerite;.   Dike   rock.  Nat.  size.   Loberia  vieja,   Masafuera. 

}  Ia<^erman  has  recorded  in  his  paper  in  tabular  form  his  conception  of  the 
horizontal  distribution  of  the  different  lavas  of  Masafuera.  This  is  reproduced  on  p. 
71    with  sotne  slight  corrections  on  the  base  of  renewed  examination. 

My  former  supposition  that  the  basaltic  lavas  only  occur  up  to  an  elevation 
of  about  I  000  m,  from  there  on  being  succeeded  by  more  alkaline  rocks  of 
soda-trachytic  composition,  is  no  longer  in  agreement  with  more  recent  observations, 
based  on  Skottsberg's  new  collections.  Any  thought  of  gravitative  differentiation 
to  explain  the  sequence  of  the  volcanic  rocks,  which  I  tentatively  proposed  in 
my  former  publication,  must  in  the  light  of  later  observations  be  discarded  now. 


The  numerous  basaltic  dikes,  traversing  the  whole  island  in  a  VV^est-East 
direction,  are  worth  special  notice  as  representing  rocks  exceptionally  rich  in 
olivine.  In  this  res})ect  they  exceed  the  most  olivine-rich  picrite  basalts  from 
Puerto  I'Vances  on  Masatierra.  l^OWKN  has  commented  on  these  rocks  as  follows: 
"( )ne  otiier  rock  may  be  mentioned  in  this  connection.  It  is  a  picrite  basalt  from 
Juan  PY'rnandc/.,  a  dike,  not  a  lava,  but  quenched  so  as  to  reveal  the  fact  of  its 
origin.  In  it  is  shown  the  highest  amount  of  normative  olivine  (53  %)  of  any  rock 
termed  basalt  by  the  author  describing  it.  Great  crystals  of  olivine  lie  in 
an  aphanitic  ground  composed  mainly  of  plagioclase  and  augite  (Fig.  30 — 32). 
Some  of  the  olivmc  basalts  of  this  island  group  are,  locally  at  least,  about 
as  rich  in  olivine  as  this  dike,  but  they  have  not  been  analyzed.  Their  high 
olivine  content  is  invariably  due  to  an  increased  amount  of  phenocrysts  of 
olivine  about  i  cm  in  diameter.  Plainly  these  crystals  were  not  in  solution 
in    the    dike    or    flow    material    at    the    time    of   its  intrusion  or  extrusion.    This 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  73 


Fig.  31.   Masafuerite  dike,    x  9.   Loberia  vieja,  Masafuera. 


Fig.  32.  Masafuerite  dike,    x  9.  Analogous  to  sample  Fig.  31   but  with  rims  of  iddingsitc 
all  phenocrysts  of  olivine.   Loberia  vieja,  Masafuera. 


74 


PERCY    (UKNSEL 


fact  does  not  prove  that  they  were  not  in  solution  in  that  material  at  an  earlier 
time.  Hut  if  one  finds  the  condition  siiown  by  these  basalts  to  be  invariably 
true  of  all  rocks  rich  in  olivine,  which  have  suffered  quenching,  one  must 
conclude  that  large  amounts  of  olivine  occur  in  solution  in  magmatic  liquids. 
A  survey  of  igneous  provinces  leaves  no  question  that  such  rocks  do  have  this 
character,  that  is,  thc\'  always  contain  either  all  of  their  olivine  or  all  in  excess 
of  a  (juite  small  amount  (apparently  some  12 — 15  %)  as  relatively  large  pheno- 
crysts.  rhe\-,  therefore,  force  acceptance  of  the  stated  conclusion"  (23,  p.  163). 
The  (juotation  above  refers  to  purely  theoretical  questions  but  indicates  the  extreme 
j)osition  these  rocks  hold  in  petrographic  classification  with  regard  to  the  abnor- 
mally high  content  of  olivine  in  basaltic  lavas.  JoilANNSEN  has  named  these  rocks 
masafuerite,  with  the  following  argument:  "The  picrite  basalt  from  Masafuera  of 
the  islands  of  the  Juan  Fernandez  group  is  a  most  extraordinary  rock  .  .  .  While 
this  particular  rock  occurs  as  a  dike,  on  the  adjacent  island  of  Masatierra,  for  example 
at  Puerto  h'rances,  there  is  a  similar  rock  in  the  form  of  a  lava  flow  with  large 
olivine  crystals  in  a  groundmass  containing  more  or  less  the  same  mineral.  I  am 
placing  the  rock  among  the  hypabyssals  on  the  basis  of  the  occurrence  on  Masa- 
fuera. To  all  olivine-melabasalt  dikes  which  contain  more  olivine  than  any  other 
mineral  and  in  addition  carry  basic  plagioclase  and  augite,  I  should  like  to  apply 
the   name  masafuerite"   (24,  p.  334). 

We  already  have  the  name  picrite  basalt  for  the  lava  flows  of  much  the  same 
composition  on  Masatierra,  with  Lacroix's  name  oceanite  as  synonym,  given  with 
the  following  definition:  "les  roches  basaltiques  porphyriques  a  olivine  sont  parfois 
extraordinairement  riches  en  peridot;  dans  I'echantillon  analyse,  tons  les  grains 
de  ce  mineral  se  touchent,  ils  sont  reunis  par  une  petite  quantite  de  plagues  de 
labrador,  englobant  des  microlites  d'augite  et  des  lames  d'ilmenite"  (14,  p.  44). 
Johann.sen  has  restricted  the  name  masafuerite  to  aschistic  dike  rocks  of  a 
picrite  basaltic  magma. 

These  dikes  may  represent,  at  least  in  part,  transmission  channels  for  the 
upj)er  basalt  beds,  supersaturated  with  iron  oxides.  A  significant  feature  in  this 
respect  is  that  they  contain  numerous  'schlieren'  of  darker  colour,  due  to  abundant 
minute  grains  of  magnetite.  These  streaks  may  indicate  relics  from  a  magmatic 
flow,  subse(juently  consf)lidated  in  the  shape  of  the  olivine  basalt,  supersaturated 
in  iron  oxides,  which  now  forms  the  highest  parts  of  the  island.  At  a  later  period 
the  ciiannels  may  then  have  been  filled  with  the  melanocratic  magma,  which  now 
characterizes  them  as  such  singular  rocks.  Sequent  intrusions  of  this  nature  might 
lead  to  phases  of  crystalHsation  in  accordance  with  l^owen's  conception  of  these 
dikes  as  (jucnched  rocks,  referred  to  above. 


Regional  Relations. 
Tectonic  Connections. 

In   the   introductory  lines   I   already  noted  that  during  recent  years  as  well  as 
in   older  reports  speculations  have  been   offered  regarding  connections  in  one  or 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  75 

Other  respect  between  the  Juan  Fernandez  Islands  and  the  volcanic  islands  of  the 
eastern  Pacific.    Some  quotations  may  be  given. 

Charles  Darwin  already  expressed  views  regarding  geotectonic  connections 
between  the  Juan  Fernandez  Islands  and,  on  the  one  side,  the  South  American 
continent,  on  the  other,  the  Galapagos  Islands.  In  his  'Observations  on  the 
volcanic  islands  and  parts  of  South  America'  he  says:  "Some  authors  have 
remarked  that  volcanic  islands  occur  scattered,  though  at  very  unequal  distances, 
along  the  shores  of  the  great  continents,  as  if  in  some  measure  connected  with 
them.  In  the  case  of  Juan  Fernandez,  situated  330  miles  from  the  coast  of  Chile, 
there  was  undoubtedly  a  connection  between  the  volcanic  forces  acting  under 
this  island  and  under  the  continent  as  was  shown  during  the  earthquake  of  1835, 
The  islands,  moreover,  of  some  of  the  small  volcanic  groups,  which  border  the 
continents,  are  placed  in  lines,  related  to  those  along  which  the  adjoining  shores 
of  the  continent  trend;  I  may  instance  the  lines  of  intersection  at  the  Gala- 
pagos"   (20,  p.  144). 

In  his  paper  'Constitution  lithologique  des  lies  volcaniques  de  la  Polynesie 
Australe'  Lacroix  gives  expression  to  much  the  same  trend  of  thought  when 
he  writes:  "Particulierement  interessantes  sont  les  iles  volcaniques  qui  se  trouvent 
a  une  plus  ou  moins  distance  de  I'Amerique  du  Sud,  les  iles  Juan  Fernandez, 
San  Felix  et  San  Ambrosio,  et  enfin  Galapagos,  puis  au  large  de  I'Amerique 
centrale,  I'ile  Clipperton.  Bien  que  la  connaissance  de  la  lithologie  de  ces  iles 
soit  loin  d'etre  completement  eclaircie,  on  peut  a  present  assurer  que  leur  laves 
different  de  celles  des  Cordilleres  des  Andes,  c'est-a-dire  de  la  serie  circumpacifique 
et  montrer  qu'elles  se  rattachent  a  la  serie  intrapacifique"   (14,  p.  64). 

In  'La  Face  de  la  Terre'  Emmanuel  de  Margerie  refers  to  the  same 
subject  as  follows:  "Le  Relay  de  la  marine  des  Etats-Unis  a  signale,  au  large 
de  Valparaiso  5.651  metres.  A  I'ouest  de  ces  fosses  sont  situees  les  deux  iles 
volcaniques  anciennes  de  San  Felix  et  San  Ambrosio;  au  Sud  de  ces  iles  le 
croiseur  Chilien  Presidente  Pinto  a  trouve,  sur  une  etendue  de  760  km,  des 
profondeurs  si  faibles  qu'il  est  probable  qu'une  crete  sous-marine,  orientee  a 
peu    pres    N — S,   s'allonge    dans    la    direction    de  I'ile  Juan  Fernandez"  (25,  III, 

P-I359)- 

In  his  'Description  and  Geology'  of  San  Felix  and  San  Ambrosio,  Bailev 
Willis  writes:  "San  Felix  and  San  Ambrosio  are  volcanic  islands  in  the  South 
Pacific  Ocean,  San  Felix  being  situated  in  latitude  26°:  5'  south  and  longitude  80°;' 
west  of  Greenwich  and  San  Ambrosio  lying  about  16  km  to  the  east-south-east. 
They  are  about  500  miles  west  of  Chanaral  on  the  east  coast  of  Chile,  and  the 
same  distance  due  north  of  the  group  of  Juan  Fernandez  and  Mas-a-fuera.  The 
South  Pacific  charts  show  several  rocks  or  islets  and  some  whose  existence  is 
recorded  as  doubtful,  which,  with  the  above-named  islands,  form  an  archipelago 
strewn  on  a  narrow  submarine  ridge  that  extends  along  the  meridian  ol  80 
degrees  west  from  about  36  degrees  south  to  26  degrees  south,  the  ridge  being 
defined  by  the  2.000-meter  contour  line.  Knowing  that  all  these  islands  and 
islets  are  peaks  of  volcanoes,  we  may  suspect  that  there  are  more  of  them  than 
we    can    see;    but  this  must  remain  an  unverified  guess  until  detailed  soundings 


PEKCN     OlENSEI. 


can  be  made.  The  deptli  of  the  ocean  in  this  region,  which  lies  west  of  the 
Ricliartls  Deej),  varies  from  4.000  to  5.000  meters.  The  islands,  therefore,  repre- 
sent tlie  summits  of  volcanoes  probably  sixteen  to  eighteen  thousand  feet  or 
more  in  height  —  that  is  to  say,  they  compare  with  the  volcanoes  of  the  Andes, 
which  are  situated  on  the  other  side  of  the  deep"   (26,  p.  365). 

In  an  interesting  paf)er  on  the  geology  of  Galapagos,  Cocos  and  Easter 
Island,  L.  J.  CliriUi  has  j)ublished  some  noteworthy  remarks  on  the  regional 
relations  of  the  \olcanic  islands  of  the  eastern  central  Pacific.  He  writes  as 
follows:  "L'nder  the  east  central  Pacific  there  lies  a  vast  area,  the  Albatross 
Plateau,  under  depths  of  less  than  2.000  fathoms,  though  on  all  sides  the  depths 
exceed  this  figure.  Xo  islands  rise  from  the  central  part  of  this  plateau,  but  at 
each  end  is  an  archipelago  that  appears  to  have  been  built  up  on  a  set  of 
intersecting  fissures,  the  Martjuesas  at  the  western  end,  and  the  Galapagos  at 
the  eastern.  On  or  near  its  southern  margin  too  there  are  several  volcanic  islands, 
including  the  Mangareva  (Gambier)  Archipelago,  Pitcairn,  Piaster,  Sala-y-Gomez 
and   the  Juan   I^'ernandez  Islands. 

"It  is  suggested  that  the  plateau  constitutes  a  resistant  block  which  has 
withstood  lateral  pressure  that  has  been  brought  to  bear  on  it  from  all  sides, 
that  around  its  margin  it  has  become  cracked  and  fissured,  and  that  on  the 
fissures  volcanic  islands  have  been  erected.  That  these  islands  owe  their  origin  to  a 
common  cause  is  suggesed  by  the  similarity  of  their  structure  and  geological 
history,  so  far  it  is  known.  .  .  .  Petrographically,  too,  these  islands  resemble  each 
other  and  differ  from  most  of  the  other  Pacific  islands.  The  most  striking  charac- 
teristics of  their  rocks  are  the  almost  complete  absence  of  nepheline-bearing 
types  and  the  presence  of  virtual  free  silicia  in   many. 

"Cocos,  St.  Vd'ix  and  St.  Ambrose  islands  are  constituted  in  part  of  nephe- 
line  rocks,  and  for  this  reason  they  are  regarded  as  lying,  not  on  the  resistant 
block,  but  beyond  its  eastern  margin.  Petrographically  they  resemble  the  Society 
Islands  and   Austral   Islands  which  lie  to  the  west  of  the  plateau. 

"It  is  thought  that  beyond  the  margins  of  the  block  the  crust  is  more 
pliable  and  has  yielded  to  pressure,  with  the  formation  of  anticlines  and  synclines. 
\'olcanoes  that  have  produced  nepheline-bearing  rocks  have  been  erected  on  the 
anticlines.  The  folds  have  tended  in  the  western  area  to  migrate  from  southwest 
to  northeast  with  a  wave-like  motion  j)roved  by  the  history  of  their  coral  reefs. 
I  here  is  not  sufficient  evidence,  however,  to  determine  whether  the  folds  which 
l)r()bably  underlie  Cocos,  St.  Felix  and  St.  Ambrose  islands  have  suffered  a 
similar   movement  '   (27,   |).  43). 

Ji  .\\  Hiu(;(;k\  has  recently  in  his  book  'Fundamentos  de  la  Geologia  de 
Chile'  discussed  the  geotectonic  position  of  the  Juan  P'ernandez  Islands.  He 
writes:  "klsta  zona  (la  regicui  situada  al  este  de  Llico,  en  Arauco)  de  dislocaciones 
tan  extranas  a  la  structura  de  la  Cordillera  de  los  Andes,  coincide  con  la  region 
donde  una  ancha  loma  submarina  se  desprende  del  continente.  F:ncima  de  la 
loma  se  levantan  las  islas  Juan  I^'ernandez  y  mas  al  norte  las  de  San  Ambrosio 
y  San   Felix. 

Parece  que  se  trata  de  una  antigua  Cordillera  que  se  separo  del  actual  conti- 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  77 

nente  entre  Magellanes  y  Arauco  y  que  se  hundio  posteriormente.  A  este  cordillera 
o  simplemente  zona  continental,  que  llamaremos  'Tierra  de  Juan  Fernandez',  se 
debera  el  termino  de  las  capas  de  la  Quiriquina  en  el  sur  de  Arauco  y  tambien 
el  hecho  que  sedimentos  marinos  de  Eoceno  no  se  conozcan  mas  a  sur. 

Pero  en  el  Oligoceno,  cuando  la  costa  del  piso  de  Navidad  se  extendio 
hasta  la  region  de  Ypiin  (45°  L.  S.),  se  habia  hundido  una  gran  parte  de  la 
Tierra  de  Juan  Fernandez,  conservandose  probablemente  cierta  extencion  en  la 
vecindad  de  las  islas  volcanicas,  cuyas  rocas  se  formaron  solo  mas  tarde  en 
erupciones  posteriores.  A  juzgar  por  el  grado  de  denudacion  y  en  vista  de  las 
actividades  volcanicas  recientes  en  Mas  a  Tierra  y  San  Felix,  la  parte  volcanica 
de  estas  islas  se  habra  formado  en  el  Terciario  superior,  probablemente  en  el 
Plioceno,  cuando  existia  todavia  un  resto  de  la  antigua  Tierra  de  Juan  Fernandez, 
de  la  cual  immigro  la  flora  del  Eoceno.  Cuando  mas  tarde  se  hundio  tambien 
este  resto,  sobresalian  solamente  las  partes  volcanicas,  constituyendo  las  islas  de 
Juan  Fernandez,  que  Servian  de  refugio  para  la  flora"  (16,  p.  59). 

The  references  now  given  suffice  to  show  how  the  position  of  the  Juan 
Fernandez  Islands  has  from  different  points  of  view  been  geotectonically  connected 
with  other  volcanic  islands  or  groups  of  islands  of  the  eastern  Pacific. 


Petrographic  Connections. 

In  many  papers  of  recent  years,  petrological  and  petrographical  connections 
between  the  rocks  of  the  Juan  Fernandez  Islands  and  those  of  other  volcanic 
islands  of  the  Pacific  Ocean  have  also  been  the  subject  of  discussion.  The  types 
of  lava  which  in  this  respect  have  been  of  special  interest  are  the  extremely 
melanocratic  picrite  basalts  (occanites  of  Lacroix)  and  their  occasional  combi- 
nations with  more  alkaline  rocks.  A  short  summary  of  the  literature  on  this 
subject  may  be  given  first. 

We  may  conveniently  begin  with  the  islands  of  San  Felix  and  San  Ambro- 
sio,  which  geographically  lie  nearest.  The  distance  is  760  km  due  north  of  Juan 
Fernandez.  H.  S.  Washington  has  given  a  petrographical  description  of  the 
rocks.  I  may  quote  some  lines  from  his  general  conclusions:  "It  would  appear 
from  the  specimens  brought  back  by  Willis  that  the  lavas  of  San  Felix  volcano 
are,  so  far  as  known,  only  of  two  kinds  —  a  decidedly  sodic  trachyte  and  a 
somewhat  variable  nepheline  basanite,  which  seems  to  be  highly  vitreous.  There 
is  little  doubt  that  the  yellow  tuff  is  derived  from  a  nephelite  basanite  magma 
closely  similar  to  that  of  the  flows.  The  prominent  characteristic  of  these  two 
types  of  lava  is  their  high  content  in  alkalies,  especially  in  soda,  while  high 
titanium  and  phosphorus  appear  to  be  other  constant  characters  of  minor  but 
still  considerable  interest.  This  conclusion  as  to  the  generally  highly  sodic 
character  of  the  San  Felix  lavas  is  subject  to  the  limitations  imposed  by  the 
absence  of  specimens  from  the  lower  flows  and  from  various  parts  of  the  island. 
Such  basaltic  lavas,  especially  if  highly  vitreous,  may  appear  megascopically  to 
be   very  uniform  and  yet  modally  and  chemically  very  diverse.    It  is,  therelore. 


7  8  I'KRCV    (jrKNSRl. 

possible  tluit  earlier,  lowermost  flows  are  less  sodic  and  more  typically  basaltic 
than  the  upper,  which  were  the  ones  examined.  "  Washington  continues:  "In 
this  {:)redominantly  highly  sodic  character  of  the  lavas  San  Felix  appears  to 
differ  widel\'  from  other  Pacific  islands.  At  IMasafuera,  it  is  true,  both  soda 
trach\-te  (and  nephcline  basanite)',  closely  like  those  of  San  Felix,  occur,  but 
these  are  accompanied  b\'  basalt  and  [)icrite  basalt,  whereas  at  the  neighbouring 
juan  b'ernandez  (Masatierra)  the  lavas  appear  to  be,  to  judge  from  Quensel's 
description,  only  olivine  basalt  with  neither  trachyte  nor  basanite.  Trachyte,  also 
highl\'  sodic,  occurs  at  several  other  Pacific  volcanic  islands,  as  do  also  nephelite 
basanite  and  similar  rocks  high  in  soda;  but  at  all  of  them  the  predominant 
lavas  are  more  or  less  normal  basalts  or  andesites;  so  that  the  general  magmatic 
character  is  basaltic  —  that  is  to  say,  sodi-calcic,  somewhat  modified  by  distinctly 
sodic  facies"  (26,  p.  382). 

Referring  to  the  trachyte  of  San  Felix  Washington  says  that  "in  thin  section 
the  rock  shows  a  somewhat  peculiar  texture,  which  resembles  that  of  the  trachyte 
of  Masafuera  described  by  Quensel,  that  of  Puu  Anahulu  on  the  island  of  Hawaii 
and  of  the  trachyte  of  Lahaina  on  Maui.  The  texture  seems  to  be  rather  usual 
in  the  trachytes  of  the  Intra-Pacific  volcanic  islands.  Ill-defined  laths  of  alkali 
feldspar  make  up  most  of  the  rock.  Most  of  these  are  arranged  irregularly,  but 
here  and  there  flow  texture  is  evident"   (26,   p.  375). 

In  several  places  in  his  paper  'La  constitution  lithologique  des  iles  vol- 
canitjues  de  la  Polynesie  Australe'  Lacroix  compares  the  petrographic  character 
ot  the  lavas  of  the  Juan  Fernandez  Islands  with  those  of  other  volcanic  islands 
of  the  Pacific.  Concerning  the  basalts  of  the  Galapagos  Islands  he  observes  that 
they  "offrent  I'analogie  la  plus  grande  avec  les  basaltes  de  Masatierra  et  avec 
ceux  des  iles  (iambier,  c'est-a-dire  avec  les  plus  calciques  des  basaltes  du 
I'acificjue  et  les  plus  pauvres  en  potasse"  (14,  p.  68).  And  further  on  in  the  same 
paper:  "Les  iles  Juan  P'ernandez  se  groupent  au  voisinage  des  iles  Marquises  et 
1  on  a  vu  (ju'il  faut  comparer  ce  que  Ton  sait  des  roches  des  iles  Galapagos  aux 
donnces  concernant  ceux  des  iles  Gambier"  (14,  p.  jy]. 

'1  hese  conclusions  of  Lacroix  refer  to  the  normal  basalts  of  Masatierra  and 
are  founded  on  the  four  new  analyses  of  such  basalts.  The  similarity  in  respect 
ot  the  Gambier  Islands,  however,  goes  a  step  further,  as  Lacroix  describes  from 
tiiere,  associated  with  more  normal  basalts,  typical  oceanites  (picrite  basalts), 
which,   as  we  have  seen,  also  occur  on   Masatierra. 

A  third  author  who  has  brought  the  Juan  Fernandez  rocks  under  discussion 
with  reterence  to  chemical  similarities  with  other  volcanic  islands  of  the  Pacific  is 
Conrad  Hi  kki.  Under  the  title  'Chemismus  und  provinziale  Verhaltnisse  der 
jungcruptivcn  (iesteine  des  pazifischen  Oceans  und  seine  Umrandung'  he  coor- 
dinates under  the  heading  "Typus  Hawaii"  (in  contrast  to  "Typus  Tahiti"),  the 
rocks  of  Hawaii,  the  Leeward  group,  Juan  Fernandez  and  Samoa,  remarking  that 
the  Juan  I-'ernandez,  San  P'elix  and  San  Ambrosio  lavas  are  good  representatives 
of  the  group,  the  basanite  from  Masafuera,  however,  showing  a  small  deficiency 

'    This  name  now  discarded    see  p.  63). 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE  JUAN    FERNANDEZ    ISLANDS  79 

in  al  —  alk  (28,  p.  177).  —  This  deficiency  is  now  explained  by  the  fact  that 
the  analysed  specimen  has  attained  an  abnormal  chemical  composition  through 
the  formation  of  deuteric  minerals  (see  p.  60). 

A  second  part  of  Chubb's  paper  on  the  geology  of  Galapagos,  Cocos  and 
Easter  islands  contains  the  "Petrology  of  the  Galapagos  Islands"  by  C.  Richard- 
son. Under  a  concluding  heading  he  says:  "The  Juan  Fernandez  Islands  are  the 
only  islands  on  which  both  types  of  basalt  (porphyric  with  dominant  phenocrysts 
of  olivine  or  with  basic  plagioclase)  are  found  in  addition  to  soda  trachyte  similar 
to  that  occurring  in  the  Galapagos  Archipelago.  Although  oceanites  (and  basanitic 
lavas)  are  also  present,  the  Juan  Fernandez  Islands  are  petrologically  closer  to 
the  Galapagos  than  any  other  islands"  (27,  p.  64). 


General  Conclusions. 

A  characteristic  feature  of  the  volcanic  rocks  of  the  Pacific  is  the  universal 
predominance  of  sodium  over  potassium.  Lacroix  says  that  in  this  respect 
"toutes  les  roches  etudiees  presentent  la  commune  caracteristique  d'etre  plus  riche 
en  soude  qu'en  potasse"  (14,  p.  55).  BuRRi  comes  to  the  same  conclusion,  based 
on  recalculations  of  all  available  analyses.  He  states  that  the  Niggli  value  k  is 
always  under  0.4  and  for  the  most  typical  regions  of  volcanic  rocks  of  the  intra- 
Pacific  Ocean  under  0.25   (28,  p.  173). 

The  value  k  in  the  four  new  analyses  from  Masatierra  in  no  case  exceeds 
0.20  (average  0.17).  And  the  light  grey  basalt  from  Masafuera  has  a  still  lower 
content  of  K2O  (Niggli  value  k  0.08).  Therefore,  with  regard  to  low  percentage 
of  potassium  the  basaltic  lavas  of  both  Masatierra  and  Masafuera  must  be  con- 
sidered in  this  respect  as  representative  for  the  basalts  of  the  volcanic  islands 
of  the  Pacific. 

In  other  respects  the  petrographic  relationship  between  Juan  Fernandez 
and  other  intra-pacific  islands  has  been  interpreted  somewhat  differently.  The 
reason  is,  however,  easy  to  explain.  In  some  cases  only  the  basaltic  rocks 
of  Masatierra  have  been  taken  into  consideration,  in  other  cases  special  notice 
has  been  given  to  the  soda-trachytic  lavas  of  Masafuera  as  indicating  a  casual 
presence  of  more  alkaline  rocks.  Since  no  rocks  of  alkaline  character  occur  on 
Masatierra,  this  island  has  petrographically  been  connected  most  closely  with  the 
Gambler  and  Marquesas  Islands  as,  according  to  Lacroix,  representing  "les  plus 
calciques  des  basaltes  du  Pacifique  et  les  plus  pauvres  en  potasse"  (14,  p.  68  and  ']']\ 

In  chemical  composition  the  basaltic  lavas  of  the  Juan  Fernandez  Islands  also 
show  similarities  with  some  of  the  basalts  from  the  Hawaiian  Islands.  Several 
analyses  of  phyric  and  aphyric  feldspar  basalts  from  Kohala  and  Maunakea  as 
well  as  from  Kilauea,  published  by  Washington  are  very  similar  to  those  of 
the  basalts  from  Juan  Fernandez  (Am.  J.  of  Sc.  5,  1923,  p.  482—87  and  6, 
p.  341).  It  can  be  suggestive  that  together  with  both  ancient  and  recent  lavas 
of  Kilauea,  Washington  also  describes  chrysophyric  picrite  basalts  of  much  the 
same  character  as  those  from  Masatierra  and  Masafuera.  Another  similarity  can 
be    given.     WHITMAN    CROSS    has    described,    together  with  the  normal  basalts. 


8o  PKRCV    QUENSEL 

soda-trachyte  from  Maui  and  Analuilu,  on  the  first  island  in  connection  with  a 
'picritic  basalt'  [V.  S.  (ieol.  Survey,  Prof.  Paper  88,  191  5,  p.  26 — 28).  This  corre- 
sponds exactly  to  the  rock  assenibhif^e  of  Masafuera. 

Petro(^ra{)hic  description  of  the  Hawaiian  basalts  from  the  mentioned  localities 
is,  also  in  other  respects,  found  to  agree  with  both  megascopic  and  microscopic 
features  of  the  basalts  of  Juan  Fernandez.  Washington  describes  an  aphyric  basalt 
from  Kohala  as  follows:  "The  type  is  a  light  grey,  almost  aphanitic  lava,  except 
that  some  rare,  very  small  feldspar  [)henocr)'sts  may  be  present,  and  a  few  pheno- 
crysts  of  olivine  are  seen  in  most  specimens  .  .  .  The  texture  is  rough  and  trachytic, 
so  that  the  rock  would  probably  be  considered  an  andesite  or  trachyte  in  the 
field.  Most  specimens  are  dense  and  very  fine-grained  or  aphanitic,  but  vesicular 
forms  may  occur"  (1.  c,  p.  485).  This  description  might  as  well  refer  to  the  light 
grey  basalts  of  Masafuera  at  elevations  between  i  lOO  and  i  400  m,  which  I  also, 
before  an  analysis  was  made,  tentatively  denoted  as  a  trachy-andesite  (12,  p.  282). 

Although  certain  lavas  of  Hawaii  evidently  present  similarities  with  the  basalts 
from  Juan  I'ernandez,  the  general  assemblage  has,  however,  a  different  character. 
According  to  Washington  "olivine-free  labradorite  basalts  constitutes  the  most 
abundant  type,  followed  in  abundance  by  andesine  basalt  and  then  by  oligoclase 
andesite'  (Am.  J.  of  Sc.  6,  1923,  p.  355).  The  high  percentage  of  andesine  basalt 
and  andesite  denote  a  magmatic  sequence  differing  from  that  of  the  non-alkaline 
lavas  of  more  southern  latitudes  of  the  Pacific.  It  may,  therefore,  be  advisible  for 
the  present  to  com[)ly  with  Lacroix  when  he  says:  "Les  roches  de  cette  ile 
(Hawaii)  constitueraient  une  division  speciale,  ayant  une  originalite  propre" 
(14,   p.  76). 

If  we  take  into  consideration  the  assemblage  of  olivine  and  feldspar  basalts 
and  soda-trachytic  lavas  on  Masafuera  as  a  characteristic  feature  for  this  island 
it  seems  evident  that,  as  Richardson  already  has  assumed,  the  rocks  of  the 
(kila|)agos  Archii)elago  display  the  closest  similarities.  According  to  the  analyses, 
j)ublished  by  Richardson,  both  the  basalts  and  the  soda-trachyte  are  in  chemical 
composition  very  similar  to  equivalent  rocks  of  Masafuera.  Also  soda-trachyte  is 
f)f  the  same  scarce  occurrence  in  the  Galapagos  Islands  as  on  Masafuera,  the 
only  sample  being  collected  by  Darwin  on  the  Beagle  voyage  of  1835.  Richardson 
says:  "Juan  T'ernandez  are  the  only  islands  on  which  both  types  of  basalt  are 
found  in  addition  to  soda  trachyte  similar  to  that  occurring  in  the  Galapagos 
archipelago  '  1 27,  j).  64).  The  low  content  of  potassium  is  in  common  for  the 
basalts   from   both   island   grouj)s. 

On  the  other  hand  we  must  evidently  exclude  any  petrographical  relationship 
between  Juan  I^'ernandez  and  San  P'elix — San  Ambrosio  where  the  lavas  have,  as 
far  as  is  known,  a  more  j)ronounced  alkaline  composition,  classified  by  Washington 
as  soda-trachytes  and  nc))hcline  basanites  (1.  c,  p.  382).  Richardson  says:  "The 
Juan  I<ernandcz  are  i)etrologically  closer  to  the  Galapagos  than  are  any  other 
islands  .  .  .  Hoth  are  situated  comparatively  near  the  American  coast  of  the  Pacific, 
but  their  similarity  is  not  shared  by  San  VeWx  and  San  Ambrosio  islands,  or 
any  other  islands  on  that  side  of  the  Pacific"  (27,  p.  64).  Lacroix  comes  to  the 
same  conclusion.    He  finds  the  closest  connection  with  San  Felix — San  Ambrosio 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  8 1 

to  be  the  highly  alkahne  rocks  of  Tahiti  in  the  mid-Pacific  (14,  p.  ']']).  Though 
lying  geographically  nearest  Juan  Fernandez,  the  rocks  of  San  Felix — San  Ambrosio 
evidently  represent  lavas  of  a  more  alkaline  composition,  with  the  exception  of 
Cocos  Island,  not  otherwise  met  with  among  the  volcanic  islands  of  the  eastern 
Pacific. 

We  may  conclude,  therefore,  that  the  lavas  of  the  Juan  P^rnandez  Islands 
in  chemical  composition,  and  to  a  certain  degree  also  with  regard  to  their  general 
aspect  and  fluctuations  in  petrographic  character,  must  be  regarded  as  most  closely 
associated  with  the  rocks  of  Galapagos  and  in  some  respects  also  with  some 
rock  assemblages  of  the  Hawaiian  Islands,  and  in  the  more  central  parts  of  the 
Pacific  Ocean,  with  the  Gambler  and  Marquesas  island  groups. 

These  connections  are  purely  based  on  petrographical  and  chemical  similari- 
ties. It  is  of  interest  to  find  how  these  connections  coincide  with  Chubb's  more 
theoretical  ideas  regarding  the  regional  distribution  of  the  volcanic  islands  of  the 
Pacific.  I  refer  to  the  quotation  above  (p.  ']6)  and  will  here  only  recapitulate  the 
following:  "No  islands  rise  from  the  central  part  of  this  plateau  (the  Albatross 
plateau),  but  at  each  end  is  an  archipelago  that  appears  to  have  been  built  up 
on  a  set  of  intersecting  fissures,  the  Marquesas  at  the  western  end,  and  the  Gala- 
pagos at  the  eastern.  On  or  near  its  southern  margin  too  there  are  several  vol- 
canic islands,  including  the  Mangareva  (Gambier)  archipelago,  Pitcairn,  Easter, 
Sala  y  Gomez  and  the  Juan  Fernandez  islands." 

An  approximately  symmetrical  distribution  of  alkaline  and  non-alkaline  rocks 
of  the  volcanic  islands  in  the  Pacific,  which  are  situated  between  Lat.  0°  and 
35°  S.  may,  therefore,  be  assumed  to  encircle  the  Albatross  plateau,  itself  so 
obviously  unendowed  with  any  islands.  The  outward  lying  connections  would 
then  include  San  Felix-Ambrosio  and  Cocos  on  an  eastern  marginal  zone  and 
the  Austral  and  Society  Islands  on  the  western  margin.  The  rocks  of  these 
islands  all  have  a  pronounced  alkaline  character  and  have  been  referred  by  Lacroix 
to  "la  serie  nephelinique"  of  the  Pacific  islands.  Inward  zones  of  connection  would 
include  Juan  Fernandez — Galapagos  and  the  Gambier  (Mangareva)  and  Marquesas 
island  groups,  characterized  by  absence  of  nepheline-bearirig  types  and  the  presence 
of  virtual  free  silica  in  many.  Lacroix  unites  them  as  belonging  to  "la  serie 
sans   nepheline"    (14,  p.  59). 

A  plausible  explanation  of  these  circumstances  might  be  that  tectonic  dis- 
turbances at 'different  intervals  have  located  cracks  and  fissures  around  a  resistant 
block,  represented  by  the  Albatross  plateau.  Volcanic  eruptions  of  different 
magmatic  composition  have  then  been  localised  to  different  areas,  the  alkaline 
lavas  being  restricted  to  peripherical  dislocations  in  contrast  to  the  non-alkaline 
lavas,  located  along  inner  lines  of  connection.  Easter  Island  and  Sala  y  Gomez, 
composed  of  lavas  of  a  deviating  type  and  composition,  may  have  been  orientated 
by  intersecting  fissures  along  other  trends. 

The  climax  of  volcanic  activity  on  all  these  islands  belongs  to  past  periods 
of  probably  late  tertiary  or  pleistocene  age.  However,  several  of  the  islands 
bordering  the  American  continent  still  manifest  obvious  indications  of  vulcanic 
nature.    On   San  Felix  volcanic  gases  were  issuing  from  a  crevice  on  the  southern 


82  I'KRCv  qiens?:l 

rim  in  May  1923.  l^aile\'  Willis  sa\'s:  "In  this  sense  and  to  this  extent  we 
may  consider  San  VcWx  an  active  volcano"  (26,  p.  370).  The  Galapagos  Islands 
are  still  the  seat  of  volcanic  eruptions.  As  late  as  1925  lava  flows  reached  the 
sea,  {)()urin^  over  the  1 00  foot  cliffs  (27,  p.  9).  The  Juan  Fernandez  Islands 
on  the  other  hand  show  no  signs  of  recent  volcanic  activity  but,  as  recorded 
above,  one,  and  possibly,  several  sub-marine  explosions  have  taken  place  in  their 
immediate  vicinity  during  the  past  century. 

The  formation  of  fissures  in  locating  the  position  of  the  volcanic  islands 
bordering  the  South  American  continent  may  be  conceived  to  be  connected  with 
dislocations  of  the  oceanic  sub-stratum  as  an  after-sway  of  displacements  during 
the  formation  of  the  continental  mountain  ranges.  Time  connections  between 
violent  eartlujuakes  on  the  continent  and  volcanic  activity  on  some  of  the  adjacent 
islands  are  significant  in  this  respect. 

However,  much  of  what  has  been  said  regarding  connections  of  the  volcanic 
islands  of  the  Pacific,  whether  founded  on  petrological  or  petrographical  similarities 
or  on  geotectonic  orientation,  must  still  be  considered  as  conjectural.  For  the 
present,  we  must  agree  with  Daly  that  "a  glance  at  the  larger  aspects  of  Pacific 
petrolog\-  shows  how  pitifully  slight  is  our  knowledge  of  the  island  petrography. 
Now  is  not  the  time  for  settled  convictions.  Now  is  the  time  for  concerted, 
persistant  effort,  leading  to  a  thorough  exploration  of  the  Pacific  archipelagos, 
under  the  auspices  of  a  single  institution  with  a  staff  of  cooperating  observers" 
(R.  Daly,  Petr()gra{)hy  of  the  Pacific  Islands.  Bull.  Geol.  Soc.  of  America,  27,  1916, 
P-  331)- 

Acknowledgements . 

The  writer  is  indebted  to  Professor  P.  Ramdoiir,  Heidelberg,  and  to  Pro- 
fessor S.  (kWKI.ix  and  Dr  W.  Uytenp.ogaardt  for  friendly  cooperation  in  deter- 
mining the  ore  minerals.  Professor  Ramdohr  has  taken  some  photomicrographs 
of  the  same  and  kindly  put  them  at  my  disposal   for  reproduction. 

Dr  ().  Mki.i.is  has  helpfully  co-operated  in  taking  most  of  the  other  photo- 
micrographs. The  landsca[)e  j)hotographs  have  kindly  been  put  at  my  disposal  by 
Professor  C".  Skoi  1  si!KR(;.  They  are  taken  by  him  on  his  visits  to  the  Juan 
i'ernandez   Islands  in    1908   and    1917. 

A  grant  from  the  foundation  Lars  Mikrtas  Mixnk  is  gratefully  acknowledged. 

.Mineralogical  Department,  University  of  Stockholm, 
March    1952. 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE  JUAN    FERNANDEZ    ISLANDS  83 

Table  I. 
Analyses  of  the  rocks  from  the  Juan  Fernandez  Islands. 


SiOa  .  .  . 
TiOj  ... 
Al,03  .  .  . 
Fe^O,  .. 
FeO  ... 
MnO  .  .  . 
CaO  .  .  . 
BaO.... 
MgO  .  .  . 
Na^O  .  .  . 
K^O  .  .  .  . 
P.O.  ... 
Cr,03 . . . 

F 

CI    

S 


Masatierra  (I— IV) 

Masafuera 

(V— VIII) 

I 

II 

III 

IV 

V 

VI 

VII 

VIII 

46.08 

46.10 

46.50 

47.62 

46.51 

43-37 

43-47 

63.43 

348 

3.80 

3.04 

3-42 

3-25 

1.03 

2.68 

0.28 

15.98 

16.54 

13-34 

16.24 

16.68 

8.48 

17.30 

18.64 

2.75 

5.48 

5-34. 

474 

4.30 

2.91 

6.87 

2.78 

8.83 

7.25 

6.71 

7-14 

8.17 

11.00 

7.09 

1.02 

0.23 

0.24 

O.II 

0.18 

0.18 

0.13 

0.07 

0.09 

10.54 

10.64 

10.04 

10.60 

8.85 

O.OI 

6.40 

5-03 

6.09 

1.68 

6.53 

4.58 

8.89 

4.41 

25-93 

8.60 

1.38 

3-61 

2.94 

2.32 

2.60 

2.80 

1-33 

2-53 

6.77 

1.36 

0.88 

0.63 

0.68 

0.36 

0.58 

0.74 

3.82 

0.32 

0.30 

0.29 

0.36 

0.46 

0.19 

0.27 
0.13 

0.18 

0.15 
0.08 

0.08 

0.18 

0.04 

— 

— 

— 

— 

- 

trace 

0.12 

0.01 

0.34 

0.96 

1.63 

1.24 

1.09 

— 

— 

— 

0.05 

0.37 

1.39 

1.61 

1.06 

0.19 

3-36 

0.24 

100.10 

100.08 

100.23 

99.84 

100.35 

100.25 

99.48 

100.35 

I.  Olivine  basalt  (dolerite).    Masatierra,    Bahia  Cumberland.  Raoult  anal. 
II.  Olivine  basalt.    Masatierra,  Bahia  Cumberland.  Raoult  anal. 

III.  Olivine  basalt.    Masatierra,  Bahia  Cumberland.  Raoult  anal. 

IV.  Aphyric  basalt.    Masatierra,  Bahia  Cumberland.  Raoult  anal. 

V.  Feldspar  phyric  basalt.    Elevation   i  420  m.  Masafuera.    A.-M.  Bystrom  anal. 
VI.  Picrite  basalt  (Masafuerite,  Oceanite)  dike  rock.    Masafuera.    N.  Sahlbom  anal. 
VII.  Iddingsite-basalt.    Elevation   i  000  m.    Masafuera.  N.  Sahlbom  anal. 
VIII.  Soda  trachyte.    Elevation   i  200  m.    Masafuera.    N.  Sahlbom  anal. 

Analyses  I — IV  have  been  made  for  A.  Lacroix  from  specimens  collected  at  Bahia  Cumberland 
by  members  of  Dumont  d'Urville's  expedition  in  1854,  These  analyses  were  first  published  in 
Lacroix'  paper:  Constitution  lithologique  des  lies  volcaniques  de  la  Polyne'sie  Australe  in  1927. 

Analysis  No.  V  is  new,  representing  a  sample  of  the  abundant  light  grey  feldspar  basalts  at 
higher  elevations  on  Masafuera. 

Analyses  VI — VlII  are  produced  from  my  earlier  paper  on  the  geology  of  the  Juan  Fernandez 
Islands  (No.  12). 


84 


PER IV    ()UEXSEL 


Analyses    calculated  as  water  free. 


■ii(\,  . 

VVJ    >3 

FrO  , 
Mn(  >  . 
CaO  , 
I'.aO.  . 
M.^O 
Nii,,(  ). 
K,()  . 
IV>.   . 

I'  .  .  .  . 

a  .. . 


I 

11 

III 

1\- 

y 

VI 

VII 

VIII 

4r..2i 

4().()H 

47-S3 

48.60 

47.36 

43-34 

45-27 

63.36 

349 

3.83 

312 

3-49 

3-31 

103 

2.79 

0.28 

If). 03 

i^'.75 

1372 

16.57 

16.99 

8.48 

18.02 

18.62 

27^> 

5-55 

5-49 

4.84 

4.38 

2.91 

7-15 

2.78 

8..S6 

7-34 

6.90 

7.29 

8.32 

10.99 

7.38 

1.02 

0.23 

024 

0.1 1 

0.18 

0.18 

0.13 

0.07 

0.09 

10.53 

10.78 

10.33 

10.82 

9.01 
0.01 
6.62 

5-03 

6.34 

1.68 

0.55 

4M 

9.15 

4.50 

25.91 

8.96 

1.38 

3.62 

2.98 

2.39 

2.65 

2.85 

1.33 

2.64 

6.76 

1.36 

0.89 

0.65 

0.69 

0-37 

0.58 

0.77 

3-81 

0,32 

0.30 

0.30 

0.37 

0.47 

0.19 

0.28 

0.18 

— 

1 

— 

■ — 

— 

0.14 

— 

— 

1 



0.15 
0.08 

0.08 

0.19 

0.04 

1  100.00 

1  100.00 

100.00 

100.00 

100.00 

1 00.00 

100.00 

100.00 

Molecular  proportions  (x  100) 


Sio, 

!•>.<  \ 
!-•<•( ) 
.MnO 
(■;!(  ). 

r.ao 
.Mk<> 

Na/) 

IV  \ 
CrV'a 
I"  ... 
CI    .  . 


Ill 


76.94 
4-36 

15.72 
i-73 

12.33 
0.32 

18.84 

16.24 
5.84 
1.44 
0.22 


[V 


77.72 
4.81 

16.43 
3-47 

10.22 

0.34 
19.22 

11.51 
4.81 
0.94 
0.21 


79.64 

3-9' 
13.72 

3.44 

9.60 

0.15 

18.42 

22.69 

3.85 
0.69 
0.21 


80.92 

4.36 

16.25 

3.03 
10.15 

0.25 
19.29 

1 1. 16 
4.27 
0.73 
0.26 


VI 


VII 


78.85 
4.13 

16.67 
2.74 

11.58 
0.25 

16.06 
0.01 

16.17 
4.60 
0.39 
0.33 

0.79 
0.23 


72.16 
1.29 
8.32 
1.82 

15.30 
0.18 

8.97 

64.26 
2.14 
0.62 
0.13 


0.23 


75-37 
3-48 

17.68 
4.48 

10.27 
o.io 

11.30 

22.22 
4.26 
0.82 
0.28 
0.09 

0.54 


105.49 

0-35 
18.27 

1-74 
1.42 

0.13 
2.99 

3.42 

10.90 

4.04 

0.13 


0.1 1 


ADDITIONAL    COMMENTS    ON    THE    GEOLOGY    OF    THE   JUAN    FERNANDEZ    ISLANDS  85 

Norms. 


Q 

C  . 

or 

ab 

an 

ne 

di 

hy 

ol 

ap 

il   . 

ml 

hm 

cm 

sal. 

fern 


II 


8.08 
18.14 
23.48 

6.76 
21.78 

10.46 
0.74 
6.62 
4.01 


56.39 
4361 


0.14 

5-23 
25.22 
29.71 

17-39 
6.28 

0.71 
7.30 
8.03 


60.30 
3971 


III 


0.40 

3.84 
20.19 
24.81 

19.32 
16.84 

0.71 
5-93 
7-97 


49.24 

50.77 


IV 


V 


VI 


VII 


4.25 

4.06 

22.39 
31.29 

16.00 

7.50 

0.87 
6.62 

7.02 


61.99 
38.01 


0.66 
2.17 

24.12 

32.49 

7-43 
19.20 

I.I  I 
6.27 
6.34 


59-44 
40.35 


3-45 
1 1.22 

15-47 


6.65 
3-51 
53-07 
0.44 
1.96 
421 


30.14 
69.84 


1-99 
4.56 

22.33 
29.62 


23.40 
1.58 
0.67 
5.28 

10.37 

0.20 
58.50 
41.50 


Quantitative  system. 


I. 

Ill 

3 

4 

Camptonose 

or 

:ab 

an 

II. 

III 

4 

4 

dosodic 

or 

ab 

an 

III. 

III 

4 

4 

dosodic 

or 

ab 

an 

IV. 

III 

4 

4 

dosodic 

or 

ab 

an 

V. 

III 

4 

5 

persodic 

or 

ab 

an 

VI. 

IV 

I 

4 

doferrous 

or 

ab 

an 

VII. 

III 

4 

4 

dosodic 

or 

ab 

an 

VIII. 

I 

2 

4 

Laurvikose 

or 

ab 

an 

16.14:36.55  :  47.31 
8.69:41.92:49.39 
7.86  :  41.34:  50.80 
7.03:38.78:54.19 
3.69:41.03:  55.28 

11.44:37.23:  51.33 
8.07:39.51  :  52.42 

25.91  :  65.88:    8.21 


Niggli   values. 


4-49 

0.79 

22.48 

57.15 
7.12 


3-43 

0.45 

0.53 
2.78 
0.86 

91-93 
8.05 


si  .  . 
qz  . 
al  .. 
f m  . 
c  ,  . 
alk 
c/fm 
ti  . . 

P  •- 
k  . 
mg. 
o. . . 
w     . 


103.7 

-35-5 
21.2 
43-6 
25.4 
9.8 
0.58 
5.88 
0.30 
0.20 
0.50 

O.II 

0.21 


1 10.4 

-22.4 

23-3 
41.2 

27.3 
8.2 
0.66 
6.83 
0.30 
0.16 
0.40 
0.24 
0.40 


III 


105. 1 

-18.9 

17.8 

51-9 

24.3 

6.0 

0.47 
5.16 
0.28 
0.15 
0.58 
0.17 
0.41 


IV 


118. 7 
-10.5 

23.9 
40.5 
28.3 
l-7> 
0.70 
6.40 
0.38 
0.15 
0.40 
0.22 
0.37 


110.7 

23-4 
47.0 
22.6 
7.0 
0.48 
5.80 
0.66 
0.08 
0.48 
0.16 
0.32 


VI 


VII 


69.8 

41.0 

8.0 
80.6 

8.7 

2.7 

O.II 

1.25 
0.13 
0.22 
0.77 
0.04 
0.19 


99-7 
27.1 
23-4 
550 
14.9 
6.7 
0.27 
4.60 
0.26 
0.16 

0.53 
0.22 
0.46 


VIII 


236.3 
+   2.3 
40.9 
18.9 
6.7 
33-5 
0.35 
0.78 
0.29 
0.27 
0.40 
0.41 
0.69 


86  PKRCV    QU KNSEL 


Bibliography. 

1.  'V\i.   Sr  1(1  iFKK.   Criisoiiiana.   Manchester    1843. 

2.  C.    Hkrtkko.   Notice  siir  I'Histoire  naturelle  de  Tile  Juan  Fernandez.  Annales  des 

Sciences  Naturelles,    Paris    ICS30,   I'ome  XXI,   p.  345. 

3.  A.   Cai.dc  i.FiuJH.   On  the  (leology  of  the  Island  of  Juan  Fernandez.  Geol.  Soc.  of 

London,    Proceedings,   Vol.    i,  1826 — 1833,  p.  256.   (Also  published  in  Phil. 
.Mag.   and   Annals   of  Philosophy,   Vol.   IX,    1831,  p.  220.) 

4.  P.    Pakkkr   Kin(;.   Narrative  of  the  surveying  voyages  of  H.  M.  S.   Adventure  and 

Beagle.   Vol.  i.    Proceedings  of  the  first  ex])edition  1826 — 1830,  p.  304. 

5.  J.    DiMoNi    dTrviiik.   \'oyage  au  Pole  Sud  et  dans  I'Oceanie.  Histoire  du  Voyage, 

III,    1842.  }) .  114. 

6.  I.   (1ran(.k.    (leologie,    Mineralogie    et  (leographie  physique  du  Voyage.   Dumont 

d'Urville,    Voyage    au    Pole  Sud  et  dans  I'Oceanie,    2^  Partie,    1854,  p.  39. 

7.  L.    Pi  A  IK.    Zur    Kenntnis    der    Insel    Juan    Fernandez.  Verh.  der  Gesellschaft  fiir 

Frdkunde  /.u  Berlin.  Band  XXIII,  1896,  p.  221. 
<S.  A.  RiNARD.  l\ej)ort  on  the  rock  specimens  collected  on  the  Oceanic  Islands 
during  the  voyage  of  H.  M.  S.  Challenger.  Rocks  of  Juan  Fernandez.  Report 
of  the  Challenger  Fxpedition,  Vol.  II,  1889,  No.  15.  (Also  published  in 
I'rench  under  the  title  'Notice  sur  les  roches  de  Pile  de  Juan  Fernandez'. 
Bull.  Acad.  Belgique,  tome  10,  1885,  p.  569.) 
9.     L.    Daraj'skv.    Uber  den  C.lockenstein  von  Juan  Fernandez.  Verh.  des  deutschen 

wissenschaftlichen  Vereins  zu  Santiago,    1886,   Heft  3,   p.  113. 
o.     R.    PoHi.MANN.    Das  Vorkommen  und  Bildung  des  sog.   Glockensteins  (Magnesit). 
Ibid.   Band   II.    1893,   Heft   5—6. 

I. Noticias  preliniinares  sobre  las  condiciones  jeograficas  i  jeolojicas  del  Archi- 

pielago.    Publ.    in    V.   Johow:   Fstudios  sobre  la   Flora  de  las  islas  de  Juan 
I-'ernande/.   Santiago   de   Chile    1896,   p.  i. 
2.     P.   (^)rKNSKi..    Die  (ieologie  der  Juan  Peinandezinseln.   Bull.  Ceol.  Inst,   of  Upsala, 
\  ol.    XI,    I  9  I  2,   J).    252. 
r.    Ha(;i:r.man.     lieitriige    zur    Geologic    der    Juan    Fernandezinseln.    The  Natural 
History  of  Juan  P'ernandez  and  Faster  Island.   Vol.   I,    1924. 
14.     A.    La(  Roix.    La    constitution    lithologique    des    iles    volcaniques  de  la  Polynesie 

Australe.   Mem.  Acad,   des  Sciences.   Paris.   Tome    59,    1927. 
I  5.     !■.  \()N  Wo  IK.  X'ulkanisnnis.  Stuttgart.  'Peil  I,  19  i  3,  p.  290;  'Peil  II,  1929,  p.  77  i,  805. 
i<>-     ]■    Bkm(;(;kn.    Inindainentos  de  la  (ieologia  de  Chile.   Santiago  de  Chile  1950,  p.  59, 

17.    'P.  II.    Ti/Akii,    H.  N.  MosKiKv,  J.  G.  Puchanan  and  J.  Murray.  Narrative  of  the 

cruise   of  II.  M.S.   Challenger.    Vol.    I,    1885,   p".  818. 
>8.     T.   SiK  i.iKiK.     The    eartluiuake    of    Juan    Fernandez    as    it    occurred    in  the  year 
I  S3  5.    London    i.S3(). 

Sixteen    years   in   Chile   and    Peru    from    1822  to  1839.   London  i84i,p.  387 
(with   a  sketch   ot   the  sub-marine  explosion). 
i'w.    Darwin.    ( ;colo,L;i(  al   observations  on   the  volcanic  islands  of  South   America, 

visited   during   the  voyage  of  H.  M.  S.   P)eagle.   London    1876,   ]).  144. 
I'.  Bar  in.    Pa(  ificite,   an   anemousite  basalt.  Journ.  Washington   Acad,   of  Sc.  Vol. 
-X  X,    I  ()3o,    p.  60. 
-  -  Mineral()gi(  al     Petrf)^ra])hy    of   the   Pacific   lavas.     Am.  J.   of  Sc.   Vol.   XXI, 
1  93  I ,    ]).  3 So,    510. 
L.    BowKN.   The   evolution   of  igneous  rocks.   Princeton    1928,   p.  163. 


ADDITIONAL    COMMENTS    ON     THE    GEOLOGY    OF     IHE   JUAN    FERNANDEZ    ISLANDS  87 

24.  A.  JoHANNSEN.   Petrography,   111,  j).  334. 

25.  E.   DE  Margerie.   La   face  de  la  'Jerre.   Ill,   ]).  1359. 

26.  Bailey  Willis  and   H.  S.   Washington.  San   Felix  and  San   Ambrosio,  their  geo- 

logy and  petrography.   Bull.  Geol.  Soc.  of  America.   Vol.  35,    1924,  p.  365. 

27.  L.  Chubb  and  C.  Richardson.   Geology  of  Galaprgos,  Cocos  and  Kaster  Islands. 

Bernice   P.   Bishop  Museum,   Honolulu.   Bull.    100,    1933,   p.  43,  47,  64. 

28.  C.   BuRRi.    Chemismus    und   provinciale    Verhiiltnisse    der  jungeruptiven  (iesteine 

des    pacifischen    Oceans    und   scine.'r   I'mrandung.    Schw.  Min.-petrogr.   Mitt. 
Band  6,    1926,   p.  177. 

Unfortunately  an  interesting  paper  by  Gorden  A.  Macdonald  on  the  'Hawaiian 
Petrographic  Province',  published  in  the  Bull.  (ieol.  Soc.  of  America  (60:2,  1949, 
p.  1588),  in  which  comparisons  with  Juan  Fernandez  and  other  Central  Pacific  Islands 
are   discussed,   has  evaded  my  attention  until  this   paj)er  was  already  in  print. 


4-  A  Geographical  Sketch  of  the  Juan  Fernandez  Islands. 

By 

C.  SKOTTSBERG. 

The  Juan  Fernandez  Islands  were  discovered  on  the  22nd  November,  1574, 
by  the  Spanish  navigator  JUAN  FERNANDEZ  who  called  them  Las  Islas  de  Santa 
Cecilia.  They  consist  of  two  islands,  distant  from  each  other,  Masatierra  with  its 
satellite  Santa  Clara,  and  Masafuera.  Masatierra  lies  360  miles  W  of  Valparaiso, 
Masafuera  92  miles  W  of  Masatierra.  According  to  the  charts  the  position  of  the 
light  in  Cumberland  Bay  on  Masatierra  is  33°37'i5"  S.  and  78°49'5o"  W.,  and 
of  the  summit  of  Masafuera,  33^46'  S.  and  8o°46'  W. 

The  islands  are  of  volcanic  origin  and  considered  to  be  late  Tertiary.  They 
show  no  signs  of  recent  activity,  but  a  submarine  eruption  near  Pta  Bacalao  in 
Masatierra  is  reported  by  Sutclifife  to  have  occurred  in  1835,  and  another  E  of 
this  island  by  Goll  in  1839  (Bruggen  pp.  326,  332).  Sutclifife  (i,  Plate  p.  387) 
published  a  drawing  of  the  eruption;  the  landscape  is  a  pure  flight  of  fancy. 
Some  visitors  have  wanted  to  recognize  a  number  of  extinct  craters.  To  this  ques- 
tion I  shall  return  later.  When  Ulloa  thought  that  he  saw  flames  bursting  from 
the  summit  of  Mt.  Yunque,  he  certainly  made  a  mistake. 

No  geographer  has,  as  far  as  I  know,  visited  the  islands,  but  many  notes  on 
their  configuration  and  topography  are  found  in  the  narratives  of  early  naviga- 
tors as  well  as  in  the  official  reports  to  the  Oficina  Hidrografica  in  Valparaiso 
by  the  Commanders  of  surveying  ships.  Certain  observations  on  the  former  dis- 
tribution of  the  forests  were  referred  to  in  an  earlier  paper  (Skottsberg  3).  Many 
popular  descriptions  of  the  nature  and  life  on  Masatierra  have  appeared  (see  Bib- 
liography), some  also  paying  attention  to  Masafuera.  The  latest,  by  JORGE  Guz- 
man Parada,  contains  much  useful  material  and  will  often  be  referred  to  here. 

Comments  on  some  earlier  descriptions  and  maps  of  Masatierra. 

The  most  interesting  account  of  this  island  from  the  i8th  century  is  found 
in  Walter's  narrative  of  Captain  (later  Lord)  Anson's  voyage.  The  illustrations 
are,  even  if  not  quite  so  accurate  as  the  author  thinks,  vastly  superior  to  the 
contemporaneous  ones  in  Ulloa's  work.  Plate  XIV  is  a  prospect  from  ¥.,  including 
Santa  Clara  (called.  Goat  I.),  the  rock  El  Verdugo  (Monkey  Key)  and  part  of 
the  north  coast  of  Masatierra,  seen  under  almost  right  angle  and  with  the  con- 
spicuous  mountains  in  correct  position.  Plate  XV  is  a  map,  not  bad  in  its  main 

6-537351   The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.   Vol.  i. 


90 


C.  SKOTTSBERG 


features;  of  the  mountains  only  Mt.  Vunque  appears.  The  names  on  the  map  are 
Monkey  Key,  luist  Bay  (Pto'  Frances),  the  Spout  (a  cascade  not  far  from  Pta 
Bacalao).  West  Hay  (Pto  Ingles)  and  Sugarloaf  Bay  (Vaqueria).  Woods  cover  the 
east  half;  the  treeless  west  half  erroneously  includes  the  still  well  wooded  Villa- 
gra  valleys.  Plate  W'l  is  a  Special  of  Cumberland  Bay,  of  which  PI.  XVII  gives 
a  good  view,  and  PI.  Will  shows  the  Commodore's  camp  in  the  valley  later 
named  in  commemoration  of  his  visit.  Masatierra  was  the  rendezvous  of  Anson's 
scjuadron  and  brought  salvation  to  the  remnants  of  the  crews,  of  which  the  greater 
[)art  had  fallen  a  victim  to  scorbut.  The  winter  months  of  1741  were  spent  here 
and   the  ships    refitted. 

Am'omo  1)E  Ui.i.o.v's  narrative  is  accompanied  by  a  panorama  of  the  south 
side  of  Masatierra  showing  Mt.  Yunque,  Mt.  Piramide,  Co  Negro  and  Damajuana, 
but  other  details  cannot  be  identified.  The  map  (Plate  IV)  is  a  rough  sketch. 
Three  bays  have  names,  Puerto  del  Ingles,  Englishman's  harbour,  very  likely 
named  to  commemorate  Selkirk  as  the  cave  called  "Robinson's  grotto"  is  found 
here,  Puerto  Grande  de  Juan  P'ernandez  (Cumberland  Bay)  and  Puerto  de  Juan 
P'ernandez  (Pto  Frances).  Some  other  (nameless)  coves  are  indicated,  e.g.  Pangal. 
Three  rivers  empty  in  the  harbours. 

I  do  not  know  the  circumstances  under  which  the  survey  by  F^RANCISCO 
Am.\I)()R  de  Amava  was  made.  It  resulted  in  a  map  published  in  1795  which 
has  formed  the  basis  of  the  charts  still  in  use,  but  it  may  not  have  been  known 
to  Tho.mas  Sutcijffe,  whose  book  "Crusoniana"  (1843)  is  accompanied  by  a 
map  with  more  details  than  the  older  ones;  with  regard  to  the  coast  line  it 
is  inferior  to  Anson's.  Sutcliffe  was  Governor  of  the  islands  in  the  1830's,  There 
are  many  names,  but  as  I  have  not  seen  Amaya's  original  map  I  do  not  know 
which  are  new.  Cumberland  Bay  is  called  Port  of  Juan  Fernandez;  W^est  Bay, 
Ulloa's  Puerto  del  Ingles,  Selkirk  Bay,  and  East  Bay  F'rench  Bay.  Sugar-loaf  Bay 
(V^aqueria)  is  called  Sandal  Bay,  an  interesting  name;  perhaps  most  of  the  sandal- 
wood was  obtained  here  in  Sutcliffe's  time.  West  of  this  place  we  find  Desola- 
tion Bay,  a  well  chosen  name;  now  called  Bahia  Juanango.  Herradura,  undoubt- 
edly an  old  S{)anish  name,  is  known  now  as  Bahia  del  Padre;  La  Punta  is  Pta 
de  la  Isla.  The  east  ca[)e,  now  Pta  or  Cabo  Hueso  de  Ballena,  is  called  Pta  de 
Juanango.  On  the  south  coast  we  find  Caravajal  (Carvajal),  Loberia,  Villagra, 
Chamelo  and  Monkey  I.  These  names  are,  however,  misplaced.  Sutcliffe's  Carva- 
jal is  Bahia  Tierra  Blanca,  a  name  placed  by  him  inland  at  the  foot  of  the  hills 
(where  it  belongs),  the  two  bights  on  both  sides  of  "Loberia",  B.  Chupones  and 
B.  X'lllagra,  are  nameless;  the  former  is  also  called  Tierras  Amarillas  on  some 
charts,  a  name  used  by  Sutcliffe  for  a  tract  of  land  back  of  his  Tierras  Blancas. 
X'illagra  is  located  east  instead  of  west  of  Mt.  Yunque,  and  Chamelo  used  for 
the  coast  now  called  Playa  Larga.  The  interior  shows  some  topographical  features; 
a  mountain  range  can  be  followed  from  east  cape  to  beyond  the  misplaced  Yunque, 
and  north   of  this  is  a  short  row  of  hills,  corresponding  to  Cordon  Central,  which 

'  Abbrcridtions.  I'>.  r.ahia  bay,  C.  -  Cordon  (range,  ridge),  Co  =  Cerro  (mountain),  L.  = 
Loberia  scaling  grounds,  M.-Morro  small  islet,  rock),  Pta  =  Punta  (point,  cape),  Pto  =  Puerto 
(port,  harbour,  ().  =Quebrada  j-iarrow  valley,  gorge),  V.  =  Valle  (valley). 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ   ISLANDS  9I 

separates  "Anson's  vale"  from  "Lonsdale"  (now  Valle  Colonial).  A  name  not 
found  on  any  other  map  is  "Kay's  town",  the  settlement  in  Cumberland  Bay. 
This  name  was  given  by  Sutcliffe  in  commemoration  of  one  JOMN  Kay  who, 
through  his  technical  skill,  greatly  furthered  the  textile  industry  in  England  and 
whose  biography  appears  in  "Crusoniana",  The  Salsipuedes  ridge  and  the  ridge 
between  Pto  Ingles  and  Vaqueria  are  marked,  while  the  conspicuous  crest  unit- 
ing Yunque  and  Salsipuedes  has  disappeared  altogether.  The  topography  of  the 
western  section  is  poor,  only  Tres  Puntas  placed  in  correct  position.  The  name 
"Puente"  is  misplaced,  but  certainly  refers  to  the  elevated  isthmus  between  Car- 
vajal  and  Herradura. 

Some  later  surveys  and  maps. 

From  time  to  time  the  Chilean  Hydrographic  Office  despatched  a  vessel  to 
the  islands  as  part  of  the  work  on  a  "derrotero"  for  the  entire  coast  of  the  Re- 
public. The  reports  were  published  in  the  Anuario  Hidrografico  de  la  Marina  de 
Chile.  Lopez  (1876)  mainly  repeats  older  statements  with  regard  to  distances, 
size  of  the  islands,  altitudes  etc.,  ViEL  (1878)  concentrated  his  attention  on 
the  possibilities  of  making  Masatierra  productive,  Vidal  Gormaz  (1881)  little 
more  than  copied  Lopez.  The  chart  was  not  much  improved.  More  information 
on  the  nature  of  the  coast,  the  serviceableness  of  the  harbours  and  anchorages, 
landmarks  etc.  are  found  in  the  compiled  "Instrucciones  nauticas"  of  1896.  For 
Cumberland  Bay  the  original  Spanish  name  Bahia  San  Juan  Bautista  is  used, 
and  some  other  early  names  are  preferred,  Bahia  del  Este,  B.  del  Oeste,  Pan 
de  Azucar  (Sugar-loaf,  also  Cerro  Alto)  and  B.  Pan  de  Azucar  (Vaqueria),  etc. 
Gunther's  report  of  1920  has  little  to  add  to  the  Instrucciones.  A  new  chart 
had  now  been  published  and  is  reproduced  in  a  very  small  scale.  The  distances 
between  certain  points  indicated  by  Giinther  agree  rather  well,  with  regard  to 
the  east  section  of  Masatierra,  with  those  on  my  map,  while  considerable  dif- 
ference is  noted  in  the  length  of  the  long,  narrow  western  section,  12.96  km  ac- 
cording to  Giinther,  10.25  on  my  map,  so  that  the  total  length  between  Pta  de 
la  Isla  and  Pta  Hueso  Ballena  becomes  22.2  and   18.5  km,  respectively. 

From  American,  French  and  English  sources  the  well-known  editor  of  geo- 
graphical and  nautical  works  L.  Friederichsen  of  Hamburg  compiled  a  new  map 
to  accompany  Ermel's  popular  account  of  his  visit  to  Masatierra  (1889).  The 
central  portion  is  much  disfigured,  but  the  general  trend  of  the  mountain  ranges 
more  or  less  correct,  the  details,  however,  erroneous  in  many  cases.  Most  of  the 
names  used  are  Spanish.  Some  are  still  in  use  on  the  British  and  Cliilean  charts, 
where,  however,  Punta  is  used  for  Cabo:  C.  del  Padre,  C.  Tunquillar  (Tinquillar), 
C.  Lemos,  Morro  Juanango,  C.  de  los  Negros  (now  also  called  Pta  Suroeste), 
B.  de  la  Vaqueria,  C.  Salinas,  Sal  si  puedes,  C.  San  Carlos,  C.  Loberia,  C.  Bac- 
alao,  C.  Pescadores,  C.  Frances,  Corrales  de  Molina  (a  series  of  hanging  gorges 
E  of  Mt.  Yunque),  Morro  Vifiillo,  Bahia  Chupones,  C.  O'Higgins.  Some  of  the 
names  on  Friederichsen's  map  are  now  forgotten:  Bahia  de  la  Fe  (  =  B.  Juanango), 
El    Palillo    (west  head  of  Pangal),  C.  Madurgo  (W  of  the  east    cape,  here  called 


92  C.   SKOTTSBERG 

C.  Giiasabullena,  a  corruption  of  Hueso  de  Ballena),  Morro  Caletas  (  =  EI 
Verciii<ijo),  C.  Chupones  (now  Pta  Larga),  Bahia  Aguabuena  [now  Tierra  Blanca, 
but  modern  charts  liave  Pta  Aguabuena  between  T.  Blanca  and  Carvajal  (Coq- 
bajal  of  Friederichsen)j.  The  to{)ography  is  much  clearer  and  more  correct  than 
in  any  of  the  earlier  maps.  With  the  exception  of  Cerro  Alto  and  Yunque  no 
names  of  mountains  have  been  put  in. 

R.  PoKlll.MANNs  sliort  description  of  the  islands,  with  special  reference  to  the 
geology,  serves  as  an  introduction  to  Johow's  well-known  work  on  the  natural 
histor\-.  Johow  s  map  of  Masatierra,  based  on  "recientes  trabajos  recopilados  per 
la  ()hcina  Ilidrogratica  en  1895",  gives  a  very  unsatisfactory  idea  of  the  topo- 
graph)-. 

Amador  dc  Amaya's  map  of  1795,  with  additions  and  corrections  by  the 
British  (no.  1383)  and  Chilean  navies,  remained  the  basis  of  all  charts  until  191 7, 
when  I  handed  over  my  notes  and  sketches  to  the  Oficina  Hidrografica.  From 
1918  on  several  editions  have  appeared.  Pta  Suroeste  replaces  Friederichsen's 
Pta  de  los  Xegros,  but  the  latter  should  be  preferred  because  not  this  point  but 
Pta  de  la  Isla  is  the  south-west  point  of  Masatierra.  For  Monkey  Key  El  Verdugo 
is  sanctioned,  Cabo  Chamelo  is  replaced  by  Los  Chamelos,  referring  to  the  rocks 
outside,  Cabo  Viudo  by  C.  Norte,  with  the  rock  in  front  called  Morro  Viudo. 
K  of  Co  Tres  Puntas  Co  Chumacera  appears.  More  important  is  that,  for  the 
first  time,  the  valleys  between  Pto  Frances  and  Pta  Pescadores  have  been  ind- 
icated and  named.  On  the  Special  of  Cumberland  Bay  is  the  new  name  Cordon 
de  las  Cabras  for  the  ridge  generally  called  C.  Central.  The  British  chart  "with 
corrections  from  the  Chilean  Gov.  chart  of  1921"  reproduced  by  Quensel  (2  p.  46) 
shows  the  topography  more  distinctly  than  the  former  except  of  the  east  sec- 
tion, where  all  the  improvements  have  been  omitted.  A  new  name  is  Pta  Mere- 
daxia  for  Pta  del  Padre.  The  latest  edition,  revised  up  to  March  1953,  is  iden- 
tical, but  for  the  topography  a  different  technique  has  been  used. 

The  names  used  by  Guzman  in  his  text  do  not  always  agree  with  those  on 
the  map.  Me  has  taken  up  Herradura  for  Bahia  del  Padre,  which  is  all  right,  but 
when  he  called  Bahia  Juanango  "l^^nsenada  Pan  de  Azucar,  cuyo  nombre  lo  debe 
a  su  islote  Juanango"  —  the  conical  Morro  —  he  made  a  mistake,  because  the 
name  Pan  de  Azucar  belongs  to  Vaqueria  and  refers  to  Co  Alto. 

During  our  expedition  I  tried  to  sketch  the  distribution  of  the  forest,  using 
the  chart  as  a  basis.  The  position  of  the  boundaries  was  determined  with  help 
of  simultaneous  aneroid  and  temperature  readings;  the  same  observations  were 
made  at  sea  level  before  and  after  every  excursion  and  the  elevations  calculated 
from  tables  I  had  received  from  the  late  IVofessor  AxEL  Hamberg.  This  method 
does  not,  of  course,  give  exact  results,  but  it  gives  more  reliable  figures  than 
the  altimeter.  Our  large  series  of  photographs  has  been  a  great  help.  Neverthe- 
less the  need  of  a  map.  based  on  a  real  survey,  was  deeply  felt,  and  when,  in 
195  I,  I  was  going  to  |)ut  my  notes  in  shape  for  publication,  I  approached  the 
Chilean  government  through  the  Swedish  Legation  in  Santiago  and  asked  for 
assistance  from  the  Chilean  Air  l-'orce.  This  was  most  generously  granted.  During 
a  flight  on  April  8,  1952,  Masatierra  was  photographed;  unfortunately  it  was  rather 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  93 

late  in  the  season  and  much  of  the  island  was  hidden  by  clouds.'  Though  some 
corrections  could  be  made  along  the  south  coast  and  around  Cumberland  Bay, 
the  result  was  not  what  I  had  hoped  for.  A  second  attempt  was  planned,  but  had 
to  be  given  up,  and  the  work  was  discontinued.  Fortunately  the  Swedish  engineer 
Mr.  Bertil  Frodin,  then  a  resident  of  Santiago  and  a  most  helpful  channel  during 
my  negotiations  with  the  authorities,  had  been  invited  to  join  the  first  flight, 
and  his  series  of  Kodachrome  pictures,  most  of  them  taken  from  the  plane,  was 
graciously  put  at  my  disposal.  They  have  proved  to  be  of  very  great  help;  the 
reproductions  here  will,  I  dare  say,  testify  to  their  high  value.  A  comparison 
with  our  photographs  allowed  me  to  identify  practically  every  single  forest  patch 
shown,  and  their  size  and  shape  was  almost  the  same  in  1952  as  in  1917.  With  the 
aidf  of  all  this  material  a  new  sketch  map,  reproduced  here  in  reduced  size,  was  drawn 
{fig.    i).  I  want  to  emphasize  that  this  map  is  a  sketch  only. 

Main  geographical  features. 

Masatierra  can  be  inscribed  in  an  obtuse-angled,  isosceles  triangle  with  the 
hypotenuse  (the  distance  from  Pta  Isla  to  Pta  Hueso  Ballena)  12.5  nautical  miles 
(23  km)  long  on  the  sea  chart  and  the  greatest  width  (from  Pta  Salinas  to  Los 
Chamelos)  4.2  miles  (7.8  km);  circumference  34  miles  (=53  km),  area  93  sq.  km. 
These  are  the  figures  generally  quoted,  but  others  are  also  found:  length  15.5, 
width  3.75  miles  (Lopez),  22  and  8  km  (Ermel),  25  and  9  km  (Branchi),  etc.  The 
figures  obtained  from  my  map  are:  length  (  =  hypotenuse)  18.5  km,  width  7  km, 
area  75.2  sq.  km. 

Masatierra  is  a  deeply  eroded  and  very  rugged  mountain  range  (fig.  2)  rising 
abruptly  from  a  submarine  ridge  running  S — N  and  bordered  by  deep  water;  the 
bathymetrical  conditions  will  not  be  discussed  here.  There  is  hardly  any  level 
land  on  the  island  worth  speaking  of.  Where  the  soil  is  not  covered  by  forest, 
as  on  the  barren  seaward  slopes  of  the  valleys,  on  the  coast  escarpments  and 
on  the  precipitous  ridges  rising  high  above  the  continuous  forest  cover,  hundreds 
of  lava  beds  overlying  one  another  can  be  distinguished,  varying  in  thickness 
from  a  few  m  (in  cases  less  than  one)  up  to  20  or  more  (Quensel  2  p.  40).  The 
location  of  the  main  summit  ridge  and,  as  a  consequence,  the  trend  of  the  val- 
leys, depends  on  the  dip  of  the  lava  beds.  From  the  east  highland  to  Mt.  Yunque 
and  from  Pta  San  Carlos  to  Co  Alto,  the  tilt  is  N  to  NE,  above  Pto  Frances 
14 — 18°  (fig.  13),  at  Centinela  and  Pangal  about  20°,  between  San  Carlos  and  Pto 
Ingles   12 — 13°  (fig.  3),  at  Co  Alto  20°  or  a  little  more.  From  the  east  end  to  in- 

'  One  of  Mr.  Frodin's  photographs  of  Masatierra  seen  from  the  air  in  2  000  m  altitude 
was  reproduced  in  the  daily  paper  "Dagens  Nyheter".  The  explanation  says:  ".  .  .  covered  with 
white  clouds  that  later  lifted,  enabling  us  to  map  the  islands  accurately".  And  in  the  text  we 
read:  "We  flew  to  and  fro  over  Masatierra  and  took  series  of  photographs  which  will  be  put 
together  to  form  maps  in  scale  i:  15000.  We  had  the  good  luck  to  get  the  summits  quite  free 
from  clouds  .  .  ."  This  story  is  confirmed  by  Mr.  Frodin's  kodachromes  which  show  the  central 
and  northern  parts  of  Masatierra  very  clear.  On  the  aerial  map  in  i  :  28  500,  submitted  to 
me  by  the  Chilean  Air  Force,  the  island  is,  however,  more  or  less  covered  with  clouds,  and 
not  one  of  the  conspicuous  mountains  could  be  identified  with  certainty. 


94 


C.  SKOTTSHERG 


5^ 


O 

a 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


95 


Fig.  2.  Eastern  half  of  Masatierra,  seen  from  the  air.  —  Photo  B.  Frodin  8/4   1952. 


Fig.  3.  End  of  Cordon  Salsipuedes  close  to  Pta  San  Carlos,  showing-  the  dip  of  the  lava  beds. 
At  the  foot  of  the  ridge  the  Cemetery.  —  Photo  C.  Skottsberg  ^/^   191 7. 


96  C.  SKOTTSBERG 

eluding  the  Vunque  massif,  the  crest  follows  the  south  coast,  an  imposing  bar- 
ranca several  hundred  m  high,  with  almost  vertical  gorges.  As  a  rule  the  saddles 
are  impassable  (figs.  11,  14,  31).  From  the  saddle  between  Yunque  and  Piramide 
(figs.  21,  22)  the  ridge  turns  NNW  and  in  a  shallow  curve  runs  right  across  the 
widest  part  of  the  island.  The  reason  is  that  the  beds,  at  least  the  middle  and 
upper  ones,  are  horizontal;  at  the  foot  of  Mt.  Yunque  a  very  insignificant  tilt  of 
5 — 9°  was  observed  in  one  place  (fig.  7).  The  central  part  of  the  island  receives 
the  greatest  precipitation,  and  erosion  has  worked  inland  from  two  opposite  direc- 
tions, but  the  Cumberland  valley  system  has  been  considerably  more  deepened 
than  the  Villagra  system.  The  ridge  is  600  to  700  m  high  in  this  section.  A 
narrow  pass,  Portezuelo  de  Villagra,  often  spoken  of  as  "Selkirk's  lookout",  forms 
the  only  practicable  passage  between  the  two  sides  of  the  island.  West  of  Vaqueria 
the  ridge  reaches  the  north  side  of  the  island,  turns  SW  and  follows  the  coast, 
rising  to  at  least  500  m  in  the  highest  peaks,  then  getting  lower  and  lower  and 
disap{)earing  as  we  approach  El  Puente,  flat  and  sandy  and  only  50  m  above 
sea  level.  The  small  peninsula  forming  the  extreme  west  of  Masatierra  is  crowned 
by  a  hill  at  least  twice  as  high.  At  Carvajal  the  beds  appear  to  be  horizontal 
(fig.  47),  but  K  of  the  isthmus  they  are  tilted  SE,  and  consequently  all  the  valleys 
trend  toward  the  south  coast.  The  dip  is  slight. 

The  change  in  position  of  the  backbone,  combined  with  its  decreasing  eleva- 
tion, has  a  profound  influence  not  only  on  the  morphology,  but,  as  a  consequence 
of  the  direction  of  the  prevailing  winds,  also  on  the  local  climate  and  thereby  on  the 
vegetation.  Climatic  dates  will  be  found  in  my  paper  on  the  vegetation  (3  pp.  812 — 
818);  the  common  wind  direction  is  SE  to  SW  (together  78%).  Along  the  east 
and  central  section  the  air  currents  are  suddenly  forced  up  over  crests  500 — 
900  m  high,  cooled  and  condensed,  and  rain  drenches  the  ridges  (fig.  4).  This  is  the 
forest  country,  where  the  deep  valleys  are  covered  with  verdure.  The  region 
around  Mt.  Yunque  may  be  shrouded  in  mist  while  all  the  country  west  enjoys 
sunshine.  Yery  often  the  lower  cloud  limit  is  knife-sharp  (Skottsb.  3  fig.  2  on 
p.  808).  V'lg.  4  shows  clouds  also  over  the  West  and  on  Santa  Clara.  But  W  of 
Cerro  Chumacera,  where  the  main  ridge  forms  the  upper  edge  of  the  long  north- 
ern escarpment,  the  air  does  not  hit  a  high,  precipitous  wall  but  rises  gradually, 
and  the  elevation  is  too  modest  to  allow  the  rain-bringing  clouds  to  gather  ex- 
cept now  and  then  during  the  winter  months.  This  is  the  barren,  treeless,  grass- 
covered  land. 


Geology  and  morphology. 

Xo  extensive  geological  survey  has  been  made  in  these  islands.  Our  know- 
ledge is  mainly  based  on  OlKNSEL's  .short  visit  in  1908,  when  he  studied  the 
stratigraphy  at  a  limited  number  of  places  and  later  gave  an  account  of  the 
geology,  petrography  and  mincralogical  compo.sition  of  the  rocks,  but  his  mater- 
ial was  too  small  to  allow  us  to  trace  the  different  kinds  of  strata  from  one  end 
of  the  island  to  the  other.  For  my  own  part  I  had  no  geological  training,  but 
during  our   19 16 — 17  campaign  I  collected  rock  specimens  in  many  places.  They 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


97 


Fig.  4.  Masatierra  and  Santa  Clara  (right,  behind  the  promontory)  seen  in  SW  from  3000  m  alt. 

—  Photo  B.  Frodin  8/^  1952. 


were  described  by  Hagerman.  The  joint  material  served  Ouensel  for  a  renewed 
study,  enlarged  to  a  discussion  of  the  geotectonic  connection  between  Juan  Fer- 
nandez and  other  volcanic  islands  of  the  East  Pacific. 

Even  a  casual  visitor  cannot  fail  to  observe  the  difference  in  appearance 
and  colour  between  the  lower  brownish,  yellowish  and  reddish  slopes  and  profiles 
and  the  higher,  light  to  dark  gray  ridges;  see  the  water  colour  sketch  in  Skottsb. 
2,  opposite  p.  52,  These  two  horizons  can  be  followed  from  the  east  end  to  Tres 
Puntas,  possibly  to  Cerro  Enrique,  but  no  samples  were  brought  from  the  ex- 
treme western  section  with  the  exception  of  a  few  from  Bahia  del  Padre.  The 
island  is  "in  the  main  formed  by  a  rather  uniform  series  of  basaltic  lava  beds, 
only  diverging  in  respect  of  coarser  and  finer  grain  or  of  a  higher  or  lower 
content  of  olivine"  (Quensel  2  p.  44).  Rocks  with  a  very  high  content  of  olivine 
(picrite  basalts)  seem  to  be  restricted  to  lower  elevations;  higher  up  more  nor- 
mal basalts,  less  rich  in  olivine,  predominate,  but  between  the  two  extremes  there 
is  every  transition.  Of  the  lower  lava  beds,  up  to  200  m  above  sea  level,  "many 
show  a  coarse-grained  ophitic  texture  and  may  be  classed  as  dolerites"  (I.e.  p.  45); 
these  lavas  have  been  traced  from  Pto  Frances  to  Tres  Puntas.  The  dolerites  are 
very  resistant  and  show,  at  least  where  observed  by  me,  a  columnar  structure. 
They  form  thresholds  in  some  of  the  valleys.  The  most  conspicuous  ones  were 
met  with  in  Vaqueria  (fig.  5)  —  a  piece  of  a  column  was  figured  by  Hagerman 
p.  28  —  and  on  the  south  side  of  the  island  below  Chumacera  and  Tres  Puntas. 
At  Chumacera  the  bed  is  about  3  m  thick.  The  pillars  appear  as  long  and  narrow. 


98 


C.  SKOTTSBERG 


Fig.   5.    \aqueria    valley,    Masatierra,    showing    bed    of   doleritic  basalt,  —  After  a  photograph 

bought  in  Valparaiso. 


F'ig.  6.  Spheroidal  weathering  of  basalt,  Valle  Colonial,  Cu.nbLiicUid   Day,  —  Photo  C.  Skotts- 

berg  24   191 7. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


99 


Zz~       Hard  basalt 
',',','/,'      Less  hard  basalt 
?oV     Agglomerate  beds 
Dikes  black 


Fig.  7.  Diagrammatic  sketch  of  a  profile  of  the  seaward  base  of  Mt.  Yunque.  Height  c.  350—400  m. 

3-sided  prisms.  The  dolerites  have  a  very  fresh  appearance  and  are  supposed  to 
represent  intrusions  between  previously  consoHdated  flows  (Quensel  2  pp.  45,  47). 

The  lava  beds  at  lower  elevations  in  east  and  central  Masatierra  are  inter- 
bedded  with  agglomerate  layers,  formed  by  tuffaceous  material  as  explained  by 
Quensel  and  illustrated  by  his  fig.  1 1  (2  p.  54).  The  photograph  was  taken  near 
the  entrance  to  Anson's  valley  at  approximately  50  m  above  sea  level.  In  a 
clayey  ground  mass  of  a  deep  brick-red  colour  hard  blocks  of  various  shapes 
and  sizes  are  embedded,  showing  spheroidal  weathering;  I.e.  fig.  10  is  a  fine 
example  from  the  same  bed.  Another,  from  the  floor  of  Valle  Colonial  near  the 
trail  to  Portezuelo,  is  seen  in  my  fig.  6.  The  entire  exposed  surface  weathers  in 
this  fashion;  the  soft  ground  mass  is  washed  out  and  carried  into  the  sea  and 
the  hard  blocks  left  lying.  The  same  kind  of  agglomerate  is  found  also  in  other 
valleys.  Stratification  is  quite  distinct  in  the  profile  illustrated  by  Quensel.  His 
opinion  is  that  these  beds  are  pyroclastic  sediments  of  recent  volcanic  origin. 
According  to  my  notes  the  volcanic  agglomerate  was  observed  overlayered  by  hard, 
gray  basalt  in  Pto  Frances  and  at  the  seaward  base  of  Mt.  Yunque  (fig.  7).  In 
the  profoundly  eroded  Cumberland  valleys  all  the  upper  strata  have  been  removed. 

All  through  the  island  the  lower  horizons  are  traversed  by  vertical  dikes  of 
hard  lava  striking  approximately  N — S.  Nowhere  are  they  better  observed  than 
on  the  imposing  perpendicular  escarpment  between  Vaqueria  and  Juanango 
bay,  where  the  wall  is  ribbed  with  innumerable  dikes  which,  thanks  to  their  greater 
hardness,  project  above  the  rim  to  form  a  serrated  edge  (figs.  8,  9). 

The  geology  of  Bahia  del  Padre  was  considered  by  POEIILMANN  to  be  of 
particular  interest  and  importance.  The  lowermost  bed  at  the  entrance  to  the  bay 
was  identified  by  him  as  an  andesite  representing  a  much  older  formation  than 
the  overlaying  basalts  and  tuffs  and  exposed  only  in  this  place,  but  Quensel  came 
to  the  conclusion  that  we  have  no  reason  to  classify  these  beds  as  andesites  of 
an  older  formation  (i  p.  266,  2  p.  56);  they  "have  been  subjected  to  alterations 
in  connection  with  thermal  processes  during  some  intermediate  phase  of  volcanic 
activity". 


C.  SKOTTSBERG 


Fig.  8.    The    escarpment    between    Vaquen'a  and  Juanango,   Masatierra,  showing  the  numerous 
vertical  dikes.  —  Photo  C.  Skottsberg  26/^  i^qS. 


j^ 

"^ 

^ < 

. 

^ 

e:  ~::^ 



J 

I  H 



^ 

A 

== 

:=  = 



1— 

y 

^ 

^ 

^ 

— 

— 

— 

^ 

/ 

uL 

— 



— 



— 

(^ 

E 

E 

E 

— 



— 

— 



— 

— 



z 

I 

I 

_ 

n 

Z- 

_ 

~1 

— 

— 

- 

- 

^" 

- — 

■ 

— 



— 









~ 

„ 

— 

— 

— 

— 

— 

— 

— 

—  - 

— ■ 

== 

— 

.  [= 

— 

— -- 

— 

/->    / 

=~ 

y 

y 

y 

UJ- 

-L-J 

^ 

^r-' 

J .— 

^^ 



- 

Fig.  9.  Diagrammatic  sketch  of  part  of  the  escarpment  between  Vaquen'a  and  Juanango;  comp. 
hg.  8.   Dikes  black;  in  nature  they  do  not  run  quite  so  regularly. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


Fig.   JO.  El  Puente  seen  from  the  shore  in  Bahia  del  Padre,  Masatierra. 

15/j  1917. 


Photo  C.  Skottsberg 


Fig.  10  shows  the  profile  of  the  Puente.  The  foot  of  the  chff  is  more  or 
less  hidden  under  talus  material.  A  tufif  bed  overlayers  a  sequence  of  basalt 
and  agglomerate  beds;  one  of  these,  rather  distorted,  can  be  followed  righ  across 
the  slope  on  PI.  102:  i,  Skottsb.  3.  The  tuff  was  described  by  Hagerman  p.  26. 
It  resembles  a  coarse-grained  sandstone  and  disintegrates  easily.  The  Puente 
(figs.  45,  46)  and  the  adjacent  slopes  on  both  sides  are  covered  with  white,  mob- 
ile sand  where  small  dunes  and  ripplemarks  announce  wind  action,  and  the  wind 
carries  the  sand  out  into  the  bay.  On  the  sand,  standing  more  or  less  upright, 
are  numerous  peculiar  more  or  less  tube-shaped  concretions  (see  fig.  7  in  Hager- 
man's  paper).  Hagerman  p.  29  calls  them  sinter  concretions  formed  by  a  number 
of  minerals  in  a  cement  of  CaCOg:  "Die  wahrscheinliche  Deutung  dieser  Phano- 
mene  ist  wohl,  dass  mit  Calciumkarbonat  gesattigte  thermale  Gewasser  iiber  eine 
Vegetationsdecke  geflossen  sind,  wobei  Wurzeln  etc.  mit  einer  Kruste  von  oben 
angegebener  Zusammensetzung  iiberzogen  wurden."  This  should  bear  witness  of 
late  volcanic  activity  contemporaneous  with  the  existence  ot  a  more  humid  cli- 
mate than  the  present  one,  permitting  a  vegetation  cover  to  thrive.  Now  there 
are  neither  any  hot  springs  saturated  with  lime  nor  is  this  region  covered  with 
native  plants;  the  Puente  is  a  field  of  pure  sand  with  large  patclies  of  weeds 
along  the  edge  (fig.  46).  There  are  indications  that  the  west  part  subsided  in 
geologically  recent  time,  when  Masatierra  and  Santa  Clara  hung  together.  With 
greater  elevation  rains  were  more  frequent  and  where  the  land  is  now  barren,  it 
bore  shrubs  and  trees. 


I02  C.   SKOTTSHKRG 

More  normal  olivine  basalts,  less  rich  in  olivine  than  the  rocks  character- 
istic of  the  lower  horizons,  are  "widespread  up  to  the  highest  part  of  the  island" 
(Oiiensel  2  p.  49).  They  are  dark  gray  and  as  a  rule  vesicular,  scoriaceous  or 
slaggy,  but  they  are  hard,  more  resistant  to  denudation,  and  form  the  elevated 
crests  and  crags  all  along  the  ridges.  At  intermediate  horizons,  approximately 
between  400  and  500  m  (Cordon  Chifladores  400  m,  Portezuelo  500  m),  feldspar 
basalts  seem  to  predominate  (Ouensel  I.e.).  These  lavas  are  ash  gray,  aphanitic  and 
aphyric  in  texture  and  have  a  tendency  to  develop  a  columnar  structure.  Rocks 
of  the  same  type  were  found  near  Pta  Larga  at  less  than  100  m  in  the  form  of 
a  dike,  "which  may  signify  a  channel  for  the  analogous  lavas  at  higher  levels". 
These  beds  are  supposed  to  "represent  a  definite  epoch  of  intrusion,  intermediate 
between  the  doleritic  basalts  and  picrite  basalts  of  the  lower  parts  and  the  sco- 
riaceous olivine  basalts  of  the  higher  horizons".  Very  likely  the  thick  bed  seen 
on  PI.  97:2  in  Skottsb.  3,  about  450  m  above  sea  level,  belongs  to  this  type. 
I  admit  that  the  photograph  selected  by  Quensel  to  illustrate  this  formation  (fig. 
9,  p.  51)  has  the  same  outward  appearance,  but  the  altitude  is  approximately 
625  m,  and  as  this  place  is  out  of  reach  —  the  climber  cannot  depend  on  the 
shallow-rooted  shrubs  and  ferns  —  no  specimens  were  taken  there;  the  samples  I 
brought  came  from  500  to  575  m,  and  whether  or  not  the  beds  are  intermediate 
is  impossible  to  tell,  as  no  rocks  from  a  higher  elevation  than  575  m  have  been 
examined.  The  samples  from  this  level  are  vesicular  aphyric  feldspar  basalts.  We 
know  nothing  about  the  mineraiogical  composition  of  the  rocks  forming  the 
highest   summit.    Mt.    Yunque    rises    about    350  m  above  Portezuelo  de  Villagra. 


A  geographical  reconnaissance  of  Masatierra. 

Pta  Ilueso  de  Ballena,  where  we  shall  start  our  circuit  of  the  island,  plunges 
abruptly  into  the  sea,  forming  an  escarpment  of  perhaps  300  m.  A  dominant 
feature  of  the  coast  is  that  talus  deposits  are  insignificant  or  lacking,  so  that 
the  surf  is  able  to  undermine  the  wall  and  to  excavate  caves.  Only  in  the  coves 
where  a  valley  has  been  eroded  down  to  sea  level,  a  beach  is  found  which  leaves 
a  f)assage  along  the  foot  of  the  escarpment. 

Between  the  east  cape  and  the  Frances  valley  the  land  rises  to  500  m  or 
more.  Some  shallow  cjuebradas,  filled  with  forest,  descend  north  toward  the  sea 
but  do  not  reach   very  far  down  (fig.   11). 

Pto  I-nviccs  does  not  deserve  to  be  called  a  harbour;  it  offers  no  protection 
even  as  an  occasional  anchorage  (Instrucc.  naut.  p.  226).  It  is  a  small,  open  cove 
facing  N  and  IC,  but  with  winds  from  other  quarters  landing  is  easy.  The  beach 
consists  of  rounded  stones  and  coarse  shingles;  here  as  elsewhere  the  surf  re- 
moves all  minor  particles.  Tlie  lower  slopes  of  the  valley  are  very  barren,  the  soil 
is  exposed  or  covered  with  })atches  of  weeds,  and  the  marks  of  running  water 
and  the  tracks  of  cattle  are  everywhere  to  be  seen  (fig.  12).  Some  little  distance 
from  the  shore  and  about  50  m  above  sea  level  is  a  small  shack.  The  streambed 
occupies  the  entire  narrow  bottom  of  the  broadly  V-shaped  valley  to  which  sev- 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


03 


Fig.  II.  Most  easterly  part  of  Masatierra,  seen  from  the  air,  looking  east.  The  detached,  coni- 
cal rock  is  El  Verdugo.  Ridges  a  (centre)  and  b.  —  Photo  B.  Frodin  ^\^  1952. 


Fig.   12. 


Pto    Frances   seen   from   a  spur  about  350  m  above  the  sea. 

"/12   1916. 


Photo  C.  Skottsberg 


I04 


C.   SKOTTSBERG 


Fig.   13.   Rulge  a  overlooking  the  E  branch  of  Frances  valley.  ^  Photo  C.  Skottsberg  ^'\^  1917. 


\.   14.   Pto  Frances,  W  valley-branch.   Left,  c\  right,  d.  —  Photo  C.  Skottsberg  1^/4  iQi?- 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


105 


Fig.  15.  Prospect  of  the  E  part  of  Masatierra,  seen  toward  SE  from  an  airplane  at  low  eleva- 
tion,   showing    the    land    from    Pto  Frances  to  Quebr.  Pesca  de  los  Viejos;  between  them,  the 
small  and  steep  Q.  Lapiz.  The  dotted  line  follows  the  crest  of  Cordon  Chifladores.  From  left  to 
right  ridges  a,  b  and  c.  —  Photo  B.  Frodin  8/4  1952. 


t*     " 


'^■'"'^f  If^"- 


»^-.     ^ 


Fig.  16.    Continues    fig.    15,    to  Pta  Pescadores.  From  left  to  right  Quebr.  Lapiz,  Pesca  de  ios 
Viejos,   Laura  and  Piedra  Agujereada;  ridges  a,  b,  c,  d.  —  Photo  B.  Frodin  s/i  1952. 

7  -  537351   TA^  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.   Vol.  i. 


Io6  C.   SKOTTSBERG 

eral  quebradas  belong,  filled  with  inaqui-infested,  grazed  liima  forest'  lower  down 
but  higher  up  with  fine  primeval  stands.  Some  gullies  cut  deep  into  the  ridge 
and  end  in  an  impassable  saddle;  on  the  south  side  is  a  precipice.  Figs.  13  and 
14  show  two  of  these  gorges.  Fig.  15  is  a  general  view  of  the  Frances  system; 
the  letters  a,  b  and  c  denote  three  conspicuous  crests,  easy  to  identify  also  on 
tigs.  II,  13,  14  and  16.  The  stream  has  some  water  also  during  the  drier  sum- 
mer half  year. 

From  Pto  Frances  the  coast  runs  in  an  almost  straight  line  to  Pta  Pescadores 
(Fishermen's  point).  The  trail  crosses  three  valleys,  Pesca  de  los  Viejos,  Laura  and 
Piedra  Agujereada,  none  of  which  has  been  eroded  down  to  the  sea  (fig.  16),  but 
all  have  been  cut  back  deep  and  widened  to  form  a  basin  furrowed  by  numerous 
small  tributaries.  To  call  these  valleys  "apenas  unas  grietas",  as  Guzman  does 
(p.  26),  is  not  to  do  them  justice. 

Cordon  de  los  Cliijiadores  (Whistlers'  ridge,  fig.  16)  is,  as  most  of  the  ridges 
extending  X — S,  wide  near  the  sea  and  narrows  inland,  in  the  steeper  rise  up 
toward  the  crest  approaching  the  knife-edge  type.  As  everywhere  along  the  north 
side  of  the  island  the  country  near  the  coast  is  treeless.  Possibly  the  forest  never 
went  clear  down  to  the  cliff.  This  question  was  raised  by  Johow  and  commented 
on  by  me  (3  p.  800).  I  forgot  then  to  mention  Ermel's  theory  that,  as  the  islands 
are  so  much  younger  than  the  mainland,  a  vegetation  cover  has  not  yet  had  time 
to  spread  to  the  coast!  There  is  no  need  for  a  discussion.  On  C.  Chifladores  the 
first  forest  is  met  with  about  300  m  above  the  sea,  covering  the  ridge  another 
50  m  and  then  passing  into  the  usual  low  scrub. 

Near  the  seaward  slope  of  the  ridge  is  a  dry  crevice  where  a  few  stunted  lumas 
linger.   We  named  it   Q.  del  Lapiz  (because  I  lost  a  pencil  there). 

O.  de  la  Pesca  de  los  Viejos  (Old  folks'  fishing  place,  fig.  17)  had  very 
little  water  in  the  stream  in  December  and  not  much  more  in  April.  Along  the 
outer  slopes  the  inclination  is  gentle;  the  middle  section  is  steeper,  up  to  35°. 
Scattered  trees  are  remnants  of  the  once  closed  forest.  The  interior  is  well  wooded 
u[)  to  the  main  ridge.   Not  much  maqui  was  seen  in  this  valley. 

O.  de  Laura  (fig.  18).  Near  the  sea  the  valley  sides  slope  23 — 24°,  a  litde 
farther  inland  about  30°.  None  of  these  valleys  is  deeply  eroded.  There  was  some 
water  in  the  stream  in  December,  but  nothing  in  April  when  the  picture  published 
in  Skottsb.  3  fig.  32  on  p.  889  was  taken."  Fig.  18  shows  the  valley  in  August 
The  outer  section  is  desert-like,  the  soil  naked  or  covered  with  weeds,  but  a  few 
solitary  trees  may  be  seen.  The  slope  facing  \l  is  more  barren  than  the  opposite 
slope.  The  interior  is  filled  with  good,  thick-stemmed  forest. 

A  high,  well  wooded  crest,  rising  to  at  least  650  m,  forms  the  background 
of  O.  de  la  Piedra  Ai^ujereada  (fig.  19)  which  got  its  name  from  a  rock  pierced 

'  For  some  of  the  leading  species  the  local  names  are  used.  Canelo,  Drimys  conferii- 
folia\  L'houU,  Juf/fi/n  auslralis\  Luma,  Aof/iof>iyniti  fcrnandesiana  (Masatierra)  and  Myrceuge- 
nia  Schulzei  (Masafiiera;;  Maqui,  Aristotelin  ;;/«7^//'/  chilensis;  a  macal  is  a  maqui  grove);  Naran- 
jillo,  l-agani  mnyu  Masatierra  and /-'.  ^.r/^/vm  ;  Masafuera; ;  Pangue,  C/^'w^/t-r^ /^//^//rt  (Masatierra) 
and   (/'.   Masafuc?-nc  Masafuera  . 

'  The  text  is  incorrect.   Tlie  photograph  was  taken  by  the  author  17.4.  191 7. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN   FERNANDEZ  ISLANDS 


107 


Fig-.   17.  Quebr.  Pesca  de  los  Viejos,  looking  S;  ridge  d.  —  Photo  C.  Skottsberg  1^/4  191; 


'i'^i!i^''.S?f^.        iu-'it 


Fig.   18.  Quebr.  Laura,  looking  N.  —  Photo  K.  Backstrom  Aug.   1917. 


io8 


C.  SKOTTSBERG 


Fig.   19.  The  interior  of  Quebr.  Piedra  Agujereada.  —  Photo  K.  Backstrom  Aug.  191 7. 


by  a  hole  (aguja,  needle).  The  exterior  section  (see  Skottsb.  3  fig.  33  on  p.  890) 
has  the  same  character  as  in  the  valleys  mentioned,  while  the  interior  is  filled  with 
fine  forest  covering  the  ridges  on  both  sides  and  above  400  m  very  damp  and  rich 
in  tree-ferns.  The  vegetation  cover  acts  as  a  sponge  and  only  a  minor  part  of 
the  precipitation  will  feed  the  stream,  its  lower  course  being  dry  during  the  summer; 
in  winter  the  water  rushes  down  to  the  sea  as  a  cascade,  mentioned  by  Anson 
("The  Spout")  and  referred  to  in  Instrucc.  naut.  p.  227,  where  it  is  said  to  run  quite 
dry  at  the  end  of  the  winter. 

El  Rabanal  (rabano  =  Raphanus  sathnis,  formerly  abundant  here)  is  very  unlike 
the  other  valleys  with  its  wide,  almost  level  floor;  in  Johow's  time  it  was  densely 
wooded,  but  in  1905  it  was  ravaged  by  fire,  and  the  forest  never  came  back.  In 
March  191 7  the  dry  soil  was  covered  with  the  dead  stalks  and  innumerable  young 
rosettes  of  Silybum  inaria7m)n  (see  Skottsb.  3  fig.  34  on  p.  891).  PI.  86  (I.e.)  shows 
the  same  spot  in  August,  when  a  vigorous  new  growth  had  sprung  up.  The  shadow 
across  the  Silybctuui  indicates  the  streambed.  There  are  a  few  dying  lumas,  maqui 
is  plentiful  in  the  quebradas,  succeeded  toward  the  interior  by  degraded  luma-canelo 
forest.   Higher  up  are  better  stands  (I.e.   PI.  94:  i). 

Rising  at  an  angle  of  about  85°  Pta  Bacalao  forms  the  end  of  the  Centinela 
ridge.  It  got  its  name  because  the  bacalao  ("stock-fish"),  the  commonest  and  most 
valuable  fish  in  these  waters  (fig.    100),  is  very  abundant  here. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


09 


O 

^  o 


2  I 


c  o 

c  bo 

03  13 


C.   SKOTl'SHERG 


Fig.  2  1.  View   W  from  the  summit  of  Centinela  ridge,  c.  780  m,  toward  Mt.  Yunque  and  Mt. 

I'inimide    right',  in  front  of  Yunque  Mt.  Damajuana;  below  Piramide  the  crest  between  Quebr. 

Minero  and  Pangal.  —  Photo  C.  Skottsberg  ^^4  ^9^7- 


Cordoji  Centinela  (Sentinel  ridge).  The  broad  northernmost  part  of  this  ridge 
forms  a  small  meseta,  upon  which,  320  m  above  the  sea,  stands  the  wireless  station  ' 
(fig.  20).  The  ridge,  which  is  steep  on  both  sides  (45 — 55°),  is  very  barren  and  shows 
the  stratification  on  the  yellowish-gray  lava  beds  very  plainly.  A  zigzag  trail,  cut 
in  the  rock,  leads  from  Pangal  to  the  station,  where  one  has  a  splendid  view  of 
Cumberland  bay  and  the  mountains  behind  (fig.  27).  Following  the  ridge,  which 
gradually  gets  very  narrow,  we  have  the  rare  opportunity  to  walk  right  across  the 
island  to  the  to[)  of  the  main  ridge  and  to  look  down  on  the  south  coast,  almost  800  m 
below  our  feet  (fig.  21).  The  gradient  is  gentle  all  the  way  up,  from  5°  a  little  south 
of  the  station  to  about  20°  farther  up.  From  about  300  m  there  is  forest  on  both 
sides,  closing  over  the  ridge  a  little  higher  up.  A  very  dense  scrub  covers  the  crest 
(I.e.   PI.  90:  2). 

I^ta  Loheria  (lobo,  seal;  [)lace  where  sealing  was  practised  in  old  times)  and 
Pta  San  Carlos  (fig.  27)  are  the  headlands  of  Cumberland  bay.  The  distance 
between   them   is  about  2   km. 

Bahia  Cionherland  or  San  Juan  Ihmiista  is  the  only  harbour  in  the  islands 
where  large  ships  find  good  anchorage;  see  the  charts  and  descriptions  by  Walter, 
Sutclifie,   Lopez,   Vidal  Ciormaz,  Glinther  etc.  The  bay  is  open  toward  N  and  NE» 

'  Ace.  to  (iuzman  p.  16  not  used  now. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


Fig.  22.    Cumberland    Bay    from  Pta  Loberia  to  Cordon  Central,  seen  from  the  air  in   looo  m 

elevation.  From  left  to  right,  Pangal,  Cordon  Escarpado,  Quebr.  Minero-Damajuana,  La  Dama- 

juana,  Valle  Anson  with  El  Yunque.  —  Photo  B.  Frodin  ^/^  1952. 

but  winds  from  these  quarters  are  uncommon  also  during  the  winter.  The  sudden 
squalls  coming  down  through  the  valleys  with  great  force  should  be  looked  out  for. 
Four  valleys  end  in  the  bay,  from  E  to  W  Pangal,  Minero-Damajuana,  Anson 
and  Colonial.  The  scenery  round  the  bay  is  very  impressive,  a  semicircle  of  bold 
mountains,  green  gorges  and  gray  precipices  (figs.  22,  27).  Many  of  the  names  quoted 
by  Guzman  are  unknown  to  me,  for  inst.  Cordon  de  la  Falda  Larga  ("falda"  in  the 
sense  of  "long  skirt"),  perhaps  =  the  back  wall  of  Q.  del  Minero,  Picacho  con  la 
Piedra  con  Letras  (.^Cerro  Piramide),  Cerro  de  los  Munoces,  Picacho  de  la  Mona 
(=  she-monkey)  and  Cerro  el  Tope  ("top").  Pico  Central,  a  long  established  name, 
is  not  mentioned  by  Guzman. 

ii/  Pangal.  The  entrance  to  this  picturesque  gorge  is  crossed  by  the  trail  to 
Centinela,  but  it  is  also  easy  to  land  on  the  stony  beach.  The  main  branch  is  a 
blind  alley  and  the  only  true  canyon  on  Masatierra,  recalling  on  a  small  scale  the 
magnificent  gorges  of  Masafuera.  The  U-shaped  gorge  ends  in  a  high  wall,  luxuriant 
with  verdure,  through  which  a  small  waterfall  leaps  down.  The  altitude  of  the  val- 
ley bottom  at  the  foot  of  the  fall  is  only  about  200  m.  Pis.  'j'^,  87  and  98  (Skottsb.  3) 
give  a  good  idea  of  the  vegetation  with  its  stately  tree-ferns  and  giant  pangues 
{Gunnera  peltata)  which  gave  its  name  to  this  valley.  No  wonder  that  Walpole 
found  that  the  "rhubarb"  grew  so  luxuriantly  on  Masatierra,  that  it  was  too  coarse 
to  be  good!  Above  the  waterfall  the  valley  widens  and  is  filled  with  primeval  forest 
(I.e.  PI.  89:  i).  A  branch  with  a  patch  of  forest  comes  down  from  the  Centinela 
ridge  (fig.   23). 


C.   SKOTTSBERG 


Fig.  23.  Centinela  ridge  and  Pangal,  seen  from  the  slope  of  Salsipuedes  above  the  Cemetery. 

Photo  B.  Frodin  2»/i  1952- 


Cordon  Escarpado  (=  steep  ridge)  with  its  Picacho,  365  m  high  (fig.  22),  sepa- 
rates Pangal  from  the  next  valley. 

Q.  del  Minero  (Miner's  v.)  comes  down  to  the  water  rather  steeply  (figs. 
22,  24).  We  did  not  survey  this  valley,  but  to  judge  from  the  colours  on  the 
Kodachrome  film  only  the  higher  slopes  have  native  luma  forest,  the  lower  being 
covered  by  maqui.  There  were  no  plantations  here  in  1916 — 17.  A  steep  spur 
separates  Q.  Minero  from 

Q.  de  la  Daniajuana  (figs.  25,  26).  The  lower  slopes  have  been  cleared,  but 
between  1  50  and  200  m  elevation  a  dense  macal  fills  the  bottom,  followed  by  a 
mixed  luma-maqui  forest  and  finally  a  belt  of  native  wood. 

La  Damajiiana  (The  Demijohn,  figs.  25,  26)  is  a  very  characteristic  landmark. 
The  cone  crowns  a  short,  high  and  narrow  spur,  continued  toward  the  sea  by  a 
long  ridge,  which  is  barren  on  the  west  side.  This  ridge  can  be  followed  up  to 
the  base  of  the  cone,  alt.  c.  430  m,  and  from  there  round  the  bend  into  the  valley 
and  up  to  c.  530  m.  The  gradient  is  steep,  nowhere  under  35°.  There  are  ledges 
of  harder  rock,  on  both  sides  with  a  small  waterfall,  the  first  at  230  m  when  going 
up  the  valley.  The  sides  of  the  cone  are  precipitous,  the  height  2338  ft  (739  m) 
according  to  the  1^2nglish  chart,  712  m  on  the  latest  Chilean  map,  but  only  570  m 
according  to  Branchi;  this  figure  is  too  small.  As  far  as  I  know  this  mountain  has 
not  been  ascended.  An  attempt  should  be  made  from  the  south. 

\  ^alle  de  Anson  drains  the  loftiest  part  of  Masatierra  and  is  watered  by  two 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN   FERNANDEZ  ISLANDS 


»»3 


Fig.  24.  Quebr.  del  Minero.  —  Photo  C.  Skottsberg  2^/^^  ,916. 


permanent  streams.  Bounded  by  Damajuana  on  the  east  and  by  Cordon  Central  on 
the  west  it  is  dominated  by  the  square  Yunque  massif  (figs.  22,  31).  Due  to  the 
scant  water  supply  in  the  side  gullies  near  the  sea,  the  distal  part  of  the  valley  floor 
inclines  steeply,  whereas  the  middle  section  has  been  levelled,  being  almost  hori- 
zontal at  the  small  clearing  known  as  Plazoleta  (or  Plazuela)  del  Yimque  ("small 
square"),  situated  about  220  m  above  the  sea  (Skottsb.  3  PI.  89:  2),  but  from  here 
the  gradient  gets  very  steep.  It  is  evident  from  the  pictures  in  Walter's  narrative 
that  quite  some  clearing  was  done  by  Anson's  party,  very  likely  as  far  up  as  to 
Plazoleta,  and  there  is  little  native  forest  below  this  point.  In  1916 — 17  the  trail 
went  through  an  extensive  macal,  and  the  lower  slopes  near  the  sea  were  barren. 
Very  few  people  lived  in  this  valley  (fig.  27).'  The  forests  of  the  interior  were  in 
good  condition  and  the  upper  montane  type  luxuriated  below  the  saddle  between 
Damajuana  and  Yunque  from  400  to  600  m. 

El  Yunque  (The  Anvil)  presents,  from  all  directions,  the  same  venerable 
appearance  (figs.  21,  22,  25,  31,  51).  The  walls  are  everywhere  steep  with  gradients 

'  Plazoleta  is  where,  in  1930,  HuGO  Weber  went  to  live  all  alone  as  a  modern  Robinson, 
as  told  in  his  interesting  book.  He  built  a  hut,  cleared  the  ground,  made  a  garden  and  raised 
chickens.  He  married  in  1932,  built  a  more  substantial  home  and  extended  his  cultivations.  In 
1942  he  left  the  island  and  settled  on  the  mainland.  The  little  farm  is  still  his  property  but  without 
a  tenant,  and  I  am  told  by  Dr.  Guillermo  Kuschel  that  the  place  was  overgrown  with  Rubus 
ulmifoUus  when  he  visited  the  island  a  few  years  ago. 


1 14 


C.  SKOTTSBERG 


Fig. 


frc 


25.  Damajuana  and  Vunque  from  tlic   ucui  u>   I'aiigal,  looking  SW. 

"/4   1917. 


Photo  C.  Skottsberg 


55°  to  70°,  in  places  almost  perpendicular.  The  exact  height  is  not  known; 
the  fit^iires  vary  from  1700  ft  (537  m,  Walpole)  to  983  m  (Viel);  Branchi  has  805  m, 
tiie  English  chart  3005  ft  (913  m),  the  new  Chilean  915  m,  Instrucc.  naut.  927  m. 
The  first  ascent  was  made  in  1795,  a  second  shortly  after,  the  third  in  1923,  after 
which  the  attempt  has  been  repeated  with  success  a  few  times  (Guzman  p.  30). 
Only  the  route  from  the  IJamajuana-Yunque  saddle  seems  to  lead  to  the  summit. 
The  figure  838  m  (doubtless  too  high)  on  Friederichsen's  map  corresponds  to  a  small 
{/mnacle  set  on  the  saddle,  the  "Camote"  (=  sweet  potato,  bulb).  The  ascent,  which 
is  difficult  and  dangerous,  was  described  by  Tp:nz,  who  was  the  first  to  give  us 
any  information  on  the  topography  and  plant  life  of  the  summit;  see  Skottsb.  3 
pp.   S97— 898. 

Cordivi  Coitral  (figs.  22,  27,  31,  99).  From  the  broad  gable  of  Mt.  Piramide, 
F  of  its  centre,  this  sharp-edged  ridge  runs  down  to  the  coast,  separating  the  Anson 
and  Colonial  valleys  and  widening  to  a  fan-shaped,  barren  and  sandy  front,  slop- 
ing down  to  the  stony  beach.  Pico  Central,  c.  570  m,  marks  the  end  of  the  harder 
basalts  Ijcloiiging  to  the  upper  horizons;  I.e.  PI.   89:  2,  Johow  PI.  IV. 

\'allc  ('dlofiiai  (V.  del  l^olvorin  of  Guzman;  p.  =  powder-house),  the  seat  of 
the  village,  is  the  only  valley  on  Masatierra  that  approaches  maturity.  The  floor 
rises  gently  to  the  foot  of  Mt.  Piramide,  a  distance  of  about  2  km,  and  is  watered 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


115 


Fie    26     Mt     Damajuana    seen    from    Anson's   valley.    In  the  foreground  a  section  through  an 
^*       "  agglomerate  bed.  —  Photo  C.  Skottsberg  2^/12   1916. 


i6 


C.   SKOTTSBERG 


Fig.  27.  Valle  Colonial  seen  from  the  Centinela  ridge,  c.  350  m  above  sea  level.  Left,  Cordon 
Central;  above,  Portezuelo  de  Villagra  and  Cordon  Salsipuedes,  ending  in  the  vertical  escarpment 
of  Pta  San  Carlos,  behind  which,  on  the  west  side  of  Pto  Ingles,  towers  Cerro  Alto.  —  Photo 

C.  Skottsberg  "/4  19 17. 

by  two  permanent  streams,  which  receive  several  small  tributaries  from  the 
surrounding  gullies.  The  valley  bottom  was  cleared  of  its  native  forest  centuries 
ago;  already  80  years  ago  practically  nothing  was  left  below  250  m.  Macales  and 
maqui-luma  stands  fill  the  interior,  and  only  farther  up  the  gullies  we  find  better 
forest.  Material  washed  down  from  the  sides  have  contributed  to  build  the  wide, 
stony  and  sandy  beach,  the  largest  piece  of  level  ground  on  this  island  (figs.  3, 
20,  27).  I  cannot  remember  having  heard  of  any  names  for  the  streams.  Guzman 
mentions  3  streams  emptying  in  Cumberland  Bay,  Arroyo  del  Hospital,  A.  de  la 
Turbia  (turbio  -=  turbid)  and  A.  de  la  Reina;  the  first  is,  I  suppose,  where  Anson 
had   his  hospital. 

FA  Pirdmidc  (figs.  21,  29,  31),  separated  from  Mt.  Yunque  by  a  narrow  saddle 
(W  l^ortezuelo  del  Yunque),  towers  above  the  colony.  Johow's  PI.  V  dates  from  von 
Rodt's  reign  and  shows  the  scenery  when  no  village  existed.  The  figure  809  m 
on  Friederichsen's  map  stands  for  the  summit  of  Mt.  Piramide.  From  both  N  and 
S  the  ascent  is  rather  steep,  40 — 50°  the  first  stretch,  but  gets  more  gentle  higher 
up,  25—20";  toward  the  valleys  on  both  sides  the  slopes  are  precipitous  (Skottsb. 
3  PI.  9i)- 

Portezuelo  de  Villagra  (fig.  28).  No  visitor,  even  if  he  only  has  a  single  day 
at  his  disposal,  fails  to  visit  this  famous  spot  with  the  Selkirk  memorial  tablet.  The 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


Fig.  28.    Portezuelo    de    Villagra,    seen    from  above  the  settlement  in  Valle  Colonial. 

C.  Skottsberg  3«/ia  1916. 


Photo 


altitude  of  the  pass  is  generally  stated  to  be  550  m;  an  average  of  20  aneroid  read- 
ings gave  593  m,  which  I  believe  is  too  much;  about  575  ought  to  be  approximately 
correct.  The  north  "door-post"  rises  with  an  angle  of  80°  (Quensel  2  fig.  9  on 
p.  51),  the  south  is  formed  by  the  slope  of  Mt.  Piramide;  see  fig.  29,  a  transverse 
profile  with  a  gradient  on  either  side  of  about  70°.  The  trail  across  has,  in  some 
places,  been  worn  down  in  the  red  forest  soil  and  has  been  deeply  eroded  by 
running  water.  From  the  saddle  one  has  a  grand  view  of  the  south  side  of  Masa- 
tierra  from  Mt.  Yunque  to  Pta  O'Higgins  and  Santa  Clara  (figs.  49,  50). 


A  volcanic  island,  especially  if  considered  to  be  of  late  Tertiary  origin,  often 
has  craters  or  shows  other  signs  of  recent  activity,  and  it  is  not  surprising  that 
visitors  to  Masatierra  have  tried  to  locate  old  centres  of  eruption.  Walpole  writes 
p.  93:  "We  skirted  up  the  western  side,  which  shows  three  semicircular  craters, 
whose  sides  toward  the  sea  are  broken  down,  thus  forming  bays  within  their  basins. 
Of  these,  Cumberland  Bay  is  the  central  .  .  ."  The  other  two  were,  I  presume,  Pto 
Frances  and  Pto  Ingles  (if  not  Bahia  del  Padre).  And  Quensel  (i  p.  256),  referring 
to  Cumberland  Bay,  speaks  of  "die  lockeren  Tuffmassen,  die  noch  einen  alten 
Kraterboden  bedecken  .  .  ."  and  p.  257:  "Machtige,  oft  rot  gefarbte  Tuffablagerun- 
gen  fallen  den  Talboden,  was  darauf  hindeutet,  dass  es  sich  nicht  um  ein  P>osionstal, 


C.   SKOl  rSBKRCr 


Fi^.   2().    South    side    of    Portezuelo    with    tlic    .Scikhk    iiicinoriai 

I'hoto  (".   Skottsberj;  ^^j^^   1916. 


)ict,  seen  troni   the  pass. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  I  I  9 

sondern  urn  eine  teilweise  erhaltene  Kraterbildung  handelt."  Finally,  "Ob  das  west- 
lich  von  dem  Portezuelo  sich  offnende  cirkusformige  Tal  audi  eine  Kraterbildung 
darstellt,  lasst  sich  nicht  entscheiden.  In  einer  vorlaufigen  Mitteilung  habe  ich  diese 
Moglichkeit  angedeutet,  und  vieles  scheint  mir  darauf  hinzudeuten,  dass  urspriinglich 
ein  Doppelkrater  hier  gestanden  hat,  wobei  gerade  der  enge  Rucken  des  Portezuelo 
die  Scheidemauer  bildet."  This  was  written  after  our  visit  in  August  1908,  and  at 
that  time  I  shared  my  companion's  opinion  (i  p.  136).  During  my  later  expedition, 
when  I  came  to  know  most  of  the  island  better,  I  arrived  at  a  different  conclusion 
(see  e.g.  2  p.  57).  The  geological  structure  is  the  same  all  the  way  from  Pto  Frances 
to  Bahi'a  Juanango,  we  find  the  same  agglomerate  beds  more  or  less  well  exposed 
in  the  valley  bottoms,  but  nobody  would  be  inclined  to  deny  that  they  are  typical 
erosion  valleys  where  denudation,  thanks  to  the  scant  supply  of  running  water,  is 
very  slow;  in  many  cases  abrasion  has  worked  faster  than  erosion.  In  Pto  Frances 
the  stream  has  barely  managed  to  lower  its  bed  down  to  sea  level  near  the  coast. 
In  Pto  Ingles,  where  the  surrounding  ridges  are  very  high,  the  distal  part  of  the 
valley  has  been  widened  and  levelled,  and  quite  some  alluvial  soil  has  been  formed. 
The  same  procedure  is,  I  think,  responsible  for  the  formation  of  the  Cumberland 
valley  system;  there  is  nothing  that  speaks  in  favour  of  a  crater  theory  or  against 
its  origin  as  the  result  of  erosion.  The  streams  drain  the  wettest  part  of  the  island, 
where  erosion  is  greater  and  faster  and  accumulation  a  factor  of  some  consequence. 
As  I  said,  my  first  impression  was  that  the  Colonial  valley  was  the  old  crater  and 
its  wall  formed  by  Cordon  Central,  Mt.  Piramide  and  the  Salsipuedes  ridge,  and 
I  think  this  also  was  what  glimmered  in  Quensel's  mind.  However,  the  thick,  strati- 
fied deposits  of  brick-red,  deeply  weathered  tuff  with  "Bruchstiicke  von  Olivin- 
krystallen,  Lapilli-artige  Lavabruchstiicke,  Erzkorner  und  Glas"  also  occur  in 
Anson's  valley,  separated  from  Valle  Colonial,  the  supposed  crater,  by  the  over 
500  m  high  Cordon  Central  which  runs  down  to  the  shore  of  the  bay.  I  mentioned 
above  that  my  photograph  of  the  section  through  an  agglomerate  bed  published 
by  Quensel  was  taken  in  Anson's  valley.  Either  this  is  another  broken-down  crater, 
or  the  deposits  are  ejections  from  the  "Colonial"  volcano.  This  would  make  us 
postulate  that  the  Anson  valley  had  been  excavated  even  to  a  lower  level  than  the 
present  one  when  the  volcano  was  active,  and  this  seems  questionable.  In  his 
second  paper  Quensel  quotes  his  earlier  discussion  (p.  53)  and  adds:  "It  is  over 
40  years  since  I  visited  the  locality  and  naturally  I  cannot  now  rely  on  any  personal 
recollection",  and  "But  the  composition  of  the  formation,  as  well  as  my  notes  from 
the  field,  offer  indications  that  the  tuffaceous  material  of  Bahia  Cumberland  also 
represents  pyroclastic  sediments  of  recent  volcanic  origin".  He  refers  to  the  sub- 
marine eruption  1835  off  Pta  Bacalao.  I  willingly  admit  that  I  lack  the  necessary 
training  to  get  to  the  bottom  with  a  geological  problem,  but  my  observation  near  the 
foot  of  Mt.  Yunque  (see  above  p.  99)  suggests  that  the  agglomerate  bed  underlying 
the  basalt  is  of  the  same  nature  as  the  formation  in  Cumberland  Bay.  With  regard 
to  Villagra,  there  is  no  semicircular  valley  corresponding  to  the  opposite  one,  for 
only  on  the  north  and  east  sides  are  ridges  resembling  a  crater  wall.  Until  a  geo- 
logist-vulcanologist  has  had  an  occasion  to  study  the  island  it  is  better  to  leave  the 
question  of  the  nature  of  Cumberland  Bay  open. 


r.   SKOITSBERG 


F'ij^.  30.  Cordon  Salsipuedes  from  Cumberland  Bay,  with  the  still  wooded   quebradas  nos.  3  to 
5  (counted  from  Pta  San  Carlos);  beyond,  the  deep  Quebr.  Gutierrez.  —  Photo  B.  Frodin  2*/^  1952. 


Fig.  31.  Cumberland   I'.ay  in  SSK,  seen  from  the  air.  From  left  to  right  Quebr.  Minero-Dama- 

juana,   Damajuana,  Portezuelo  del  ^'unque,  Valle  Anson,  El  Yunque,  El  Piramide  with  Cordon 

and   Pico  Central,  .Salsipuedes  with  Pta  San  Carlos.  —  Photo  B.  Frodin  »U  1952. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


Fig.  32.  From  the  trail  across  the  Salsipuedes  highland,  looking  SE.  Photo  C.  Skottsberg  ^i/^  191 7. 


Cordon  Salsipuedes  forms  the  western  boundary  of  Cumberland  Bay  and  runs 
from  the  main  range  to  Pta  San  Carlos  (figs.  3,  27,  31),  along  the  east  side  of  the 
Salsipuedes  highland  between  the  bay  and  Pto  Ingles.  Whether  the  name,  which 
means  "get  out,  if  you  can",  was  originally  applied  to  the  coastal  escarpment,  380  m 
high  on  Friederichsen's  map,  I  cannot  tell.  A  well-worn  zigzag  trail,  also  used 
as  a  bridle-path,  unites  the  colony  with  Pto  Ingles  and  crosses  the  ridge  at  about 
410  m.  From  here  it  is  not  difficult  to  follow  the  ridge  up  to  over  700  m;  who 
attempts  to  continue  until  the  rib  abuts  on  the  backbone  takes  his  life  in  his  hands, 
for  the  ridge  is  a  knife-edge,  studded  wath  crags.  Already  at  about  600  m  it  is  only 
I — 2  m  wide,  and  the  rise,  quite  gentle  farther  down,  increases  to  25 — 30°.  As 
we  climb,  scrub,  brushwood  and  forest  succeed  each  other  on  the  slopes. 

The  declivity  facing  the  Colony  is  sculptured  by  a  series  of  shallow  gullies,  of 
which  the  two  nearest  to  the  coast  are  treeless,  whereas  the  others  are  wooded. 
A  comparison  between  fig.  30  from  1952  and  Skottsb.  3  PI.  90:  i  from  19 16  makes 
it  clear  that  the  forest  patches  have  undergone  no  perceptible  change  in  size.  The 
floor  of  the  fourth  quebrada  (counted  from  Pta  San  Carlos)  slopes  30°;  the  forest 
is  of  an  open,  degraded  luma-canelo  type  with  much  maqui.  There  is  no  water  in 
these  gullies.  The  gullies  of  the  main  ridge,  O.  Gutierrez  and  O.  del  Monte 
Maderugo,  were  thickly  wooded  in   19 1 7  (monte  maderugo  =  forest  full  of  timber). 

The  trail  to  Pto  Ingles  ascends  the  buttress  between  the  gullies  no.  2  and  3, 
descends  on  an  undulating  slope  (fig.  32),  crosses  Loma  de  los  Muiioces  and  descends 

8  —  537351   The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.  Vol.  I. 


122  C.  SKOTTSHERG 

rather  abruptly  into  V.  Ingles.  The  quebradas  coming  down  from  the  main  ridge 
are  filled  with  a  remarkable  Dicksonia  forest  near  the  range  —  note  Q.  Helechos  ("Fern 
gully")  —  followed  by  luma  groves  and  macal;  I  refer  to  my  description,  3  p.  909. 
Fog  is  a  very  important  climatic  factor  here. 

Pto  luf^U's  offers  no  protection  and  is  not  used  as  a  harbour  (fig.  33).  As  a  rule 
there  is  a  heavy  swell  and  the  boulders  on  the  beach  are  shifted  to  and  fro  making 
landing  uncomfortable,  perhaps  dangerous.  Close  to  the  east  headland  a  rock  pro- 
jects, separating  a  miniature  cove  from  the  main  bay.  The  beach  is  a  steep  wall 
of  boulders,  but  with  a  tolerably  calm  sea  landing  is  easy  at  the  foot  of  the  rock, 
which  is  {)ierced  by  a  tunnel,  "una  roca  agujereada  sobre  una  playa  de  piedras 
grandes",  as  this  place  is  described  in  Instrucc.  naut.  p.  229;  referring  to  Selkirk, 
Guzman  calls  the  tunnel  "la  portada  del  Solitario"  (p.  23).  Through  the  tunnel  or, 
at  high  tide,  across  the  rock,  we  gain  the  bay,  and  immediately  to  the  left,  about 
5  m  above  sea  level  (Branchi),  is  the  famous  "Robinson's  cave",  a  favourite  goal 
for  visiting  tourists  (fig.  34).  It  is  hardly  probable  that  the  cave,  described  in  some 
detail  by  Guzman  I.e.,  served  the  recluse  as  his  permanent  abode. 

Fig.  35  is  a  general  view  of  the  valley  seen  from  an  airplane,  but  this  picture 
does  not  show  the  extent  of  the  beach  flat  and  of  the  wide,  gently  sloping  valley 
floor.  The  stream  has  water  at  all  seasons.  A  ridge  extending  halfway  down  the 
valley  divides  it  in  two;  the  east  branch  comes  from  a  crescent-shaped  saddle,  equally 
conspicuous  from  both  sides  of  the  island  (figs.  35,  36)  and  reported  to  be  impassable. 
The  dividing  cordon  can  be  followed  along  the  crest  up  to  about  550  m,  where  it 
gets  so  narrow  that  further  advance  becomes  too  hazardous  (see  Skottsb.  3  PI. 
90:  2).  All  the  low  land  in  the  valley  (fig.  36)  has  been  cleared  by  fire  and  the  forest 
replaced  by  extensive  weed  fields,  but  in  the  branch  valleys  and  side  gullies  is  some 
good  forest,  where  a  few  chonta  palms  have  been  spared  (Skottsb.  3  PI.  88).  There 
is  much  naranjillo,  but  little  maqui.  The  continuous  cover  of  herbs  and  grasses 
testifies  that  erosion  is  slight,  and  so  is  the  inclination  of  the  cleared  valley  floor. 
It  would  [)erhaps  be  possible  to  reforest  this  valley  with  luma  and  other  native  trees. 
At  present  (or  at  least  in    1917)  it  is  grazed;  a  tropilla  is  seen  in  fig.  36. 

Ccrro  Alto  boldly  terminates  the  dividing  ridge  between  Pto  Ingles  and  Vaqueria 
(fi^^-  37>  38).  Possibly  this  ridge  gives  access  to  the  summit  which  is  about  600  m 
high;  an  older  figure  says  627  (Friederichsen,  Johow).  All  other  sides  are  almost 
or  cjuite  [)erpendicular.   Patches  of  forest  are  seen  on  the  flanks  of  the  cone. 

Bahia  dc  la  Wiqucria  serves,  as  the  name  suggests,  as  a  cattle  ranch.  The 
cove  is  useless  as  an  anchorage,  but  landing  is  easy  enough  with  a  calm  sea.  There 
is  no  level  beach;  the  stream,  which  is  permanent,  gropes  its  way  between  a  wall 
of  boulders  of  all  sizes.  Mg.  39  is  a  general  view  of  the  valley  seen  from  the  air. 
The  outer  [)art  is  grass  land  with  scattered  trees  on  the  slopes,  closing  to  form  groves 
higher  up,  the  interior  is  densely  wooded  (Skottsb.  3  fig.  35  on  p.  894).  The  ani- 
mals in  X'aquen'a  have  been  left  to  run  wild,  and  the  visitor  should  look  out  for 
the  bulls. 

As  far  as  I  could  sec,  the  geological  structure  is  the  same  as  in  Cumberland 
Hay,  with  the  same  red  volcanic  agglomerate,  and  it  was  during  our  visit  to  Vaqueria 
that  I   began   to  doubt  the  crater  character  of  the  former.   Halfway  up  the  valley 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


123 


liiilii 


Fig.  33.    Pto   Ingles  with  the  foot  of  Cerro  Alto  (note  tilt  of  the  strata).  Mrs.  S.  seated  on  an 
old  Spanish  gun.  —  Photo  C.  Skottsberg  20^1  1917. 


.V.  ... 


^r-yr         Vjs^-; 


Fig.  34.    Ihe  cave  ("Robinson's  grotto")  in  Pto  Ingles.  —  Photo  P.  Quensel  "/g  1908. 


SKOTTSBERG 


35.    The    surroundings    of   Pto   Ingles,   seen  from  the  air;  note  the  crescent-shaped  saddle 
o\erlooking  the  south  coast  of  the  island.  —  Photo  B.  Frodin  ^1^  1952. 


Fig.  36.  The  gently  sloping  floor  of  V'alle  Ingles  with  the  main  range  in  the  background;  right, 
the  dividing  ridge  where,  in   1908,  the  last  living  Satitalum  grew.  —  Photo  K.  Backstrom  191 7. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN   FERNANDEZ  ISLANDS 


25 


Fig.  37.  Cerro  Alto,  seen  from  Cordon  Salsipuedes,  The  small  eminence  above  the  escarpment 
is  the  same  shown  at  the  extreme  right  in  fig.  38.  —  Photo  C.  Skottsberg  ^'12  191 6. 


Fig.  38.  Cerro  Alto  seen  from  a  point  on  the  crest  of  the  central  ridge  in  Valle  Ingles,  c.  400  m 
above  sea  level,  looking  N.  —  Photo  C.  Skottsberg  ^^/^  1917. 


126 


SKOTTSHERG 


^  'ft-  39-     I  'i6     Vaquen'a    cove    and    valley,    seen    from  the    air.  Left,   Cerro  Alto,  right,    ridge 

between  Vaqueria  and  Juanango;  the  top  of  Cerro  Chumacera  visible  behind.  — 

Photo  B.  Frodin  8/4  1952. 


I   Co  Agiulo 


7  »  9 

:i  I'ir;'uni(ie;    3  Kl  Yunque;    4  Co  Chumacera;   5  Co  Tres  Puntas;  6  V.  Villagra; 
7  \illa  Alcmana;  8  M.  Juanango;  9  V.  Juanango. 

Fig.  40.   I5ahia  Juanango  with   Pta  Negros  and  Morro  Juanango,  seen  from  the  air  looking  SE. 

—  Photo  B.   Frodin  8/4   1952. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  I  27 

we  meet  the  conspicuous  dolerite  bed  mentioned  above  (p.  97  and  fig.  5).  A  pas- 
sage across  the  steep  rocky  and  scrub-covered  ridge  leads  us  down  into  the 
Juanango  valley. 

The  spectacular  escarpment  between  Vaqueria  and  Pta  Negros  was  described 
and  illustrated  above  (p.  99  and  fig.  8).  Near  the  entrance  to  Vaqueria  is  a  low  rock 
and  a  little  farther  west,  o^  Pta  Norte,  another  called  Morro  del  Viudo  (Widower's 
rock).  These  rocks  rest  on  a  submarine  abrasion  terrace,  clearly  distinguishable  at 
low  tide.  The  coast  wall  projects  west  in  a  long,  narrow  and  curved  spur,  not  unlike 
a  saw-blade,   Pta  de  los  Negros,  and  SE  of  this  lies  Morro  Juanango  (fig.  40). 

Bahia  del  Juanango  is  a  wide  bight,  protected  against  winds  from  N  and  E, 
but  otherwise  open.  Seen  from  some  distance  out  to  sea  it  looks  forbidding,  and 
the  name  "Desolation  Bay"  seems  well  chosen,  but  on  a  closer  view  two  green 
quebradas  come  in  sight.  Villa  Alemana  (Germantown),  accessible  with  difficulty, 
and  Q.  Juanango. 

Q.  del  Juanafigo.  The  beach  in  the  little  cove  is  of  the  same  kind  as  in 
Vaqueria,  a  low  escarpment  and  large,  angular  stone-blocks,  but  with  a  suitable 
wind  landing  is  easy.  My  diary  calls  these  blocks  "conglomerate",  without  much 
doubt  identical  with  the  agglomerate  found  in  Vaqueria.  The  valley  is  full  of  weeds 
near  the  sea,  but  as  it  isn't  grazed  native  grasses  are  abundant  farther  in.  At  about 
200  m  above  the  sea  are  the  first  forest  patches,  and  a  little  higher  up,  at  the  small 
waterfall,  is  good  forest.  The  threshold  is,  I  suppose,  formed  by  the  same  dolerite 
bed  as  in  Vaqueria,  but  I  did  not  bring  any  specimens. 

From  Juanango  to  Bahia  del  Padre  the  coast  escarpment  trends  SW  in  wide, 
slightly  concave  curves  to  Pta  Letnos  and  Pta  Tunquillar  and  thence  to  Pta 
Meredaxia,  the  east  head  of  B.  del  Padre.  Figs.  41  and  42  make  further  descrip- 
tions superfluous.  Two  conspicuous  mountains  tower  high  above  the  coast-line, 
Chumacera  and  Tres  Puntas,  overlooking  both  sides  of  the  island;  the  latter  with  its 
three  peaks  ought  to  be  a  fitting  goal  for  expert  climbers  (fig.  43).  On  the  slope 
of  Chumacera  are  found  the  most  westerly  forest  patches  on  Masatierra;  from  here 
the  country  is  treeless. 

Bahia  del  Padre  (Parson's  Bay)  got  its  name  from  the  configuration  of  the 
rock  W  of  the  entrance.  The  old  Spanish  name  B.  Herradura  (Horseshoe  B.)  is  still 
used  by  some  authors.  The  diameter  is  about  250  m.  It  is  a  convenient  harbour 
for  small  boats,  but  the  entrance,  guarded  by  rocks,  is  narrow  (fig.  44).  Landing 
on  the  beach  of  sand  and  shingles  (Skottsb.  3  PI.  102:  i)  is  comfortable  with  a 
moderate  swell,  but  may  be  difficult.  The  cove  is  a  natural  amphitheatre,  but  we 
do  not  find  very  much  of  the  "risuenas  representantes  del  mundp  vegetal"  praised 
by  Guzman  (p.  24),  except  a  patch  of  salt-meadow  [Salicornia)  along  the  beach, 
because  the  flora  is  poor  and  mainly  consists  of  weeds,  among  which  the  gilly- 
flowers are  conspicuous.  Nowhere  is  the  climate  drier.  El  Puente  was  described  and 
illustrated  above  (p.  loi,  fig.  10);  the  flat,  sandy  surface  is  seen  on  figs,  45  and  46, 
the  former  showing  the  wind-polished  tufT  beds. 

Looking  at  the  maps  and  photographs  one  is  struck  by  the  peculiar  appearance 
of  B.  Padre.  It  is  evident  that  it  is  no  valley,  no  result  of  erosion.  Branchi  wrote 
(p.  168):  "Un  crater  muy  pronunciado  puede  suponerse  en  la  Bahia  del  Padre",  and 


128 


C.   SKOTTSBERG 


Fig.  41, 


Jl 


Fig.  42. 

Hgs    4 1 --42.    North    coast    of  western   Masatierra  from   h.  Juanango  to  Pta  Lemos,  seen  from 
<Juebr.  Juanango.  Cerros  Cluimacera    note  patches  of  luma,  Tres  Puntas  and  PInrique.  —  Photo 

C.   Skottsberg  ^/^   1917. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN   FERNANDEZ   ISLANDS  1 29 


Fig.  43.  Cerro  Tres  Puntas.  —  Photo  Hans  Frey 


C.   SKOTTSHERG 


Fig.  44.  The  entrance  to  Bahia  del  Padre.  —  Photo  C.  Skottsberg  ^'^j^  191 7. 


I  am  tempted  to  endorse  his  opinion.  Both  Quensel  and  Hagerman  came  to  the 
conclusion  that  the  rocks  have  been  subject  to  post-volcanic  thermal  processes,  and 
Hagerman,  after  a  description  of  the  palagonite  tuff  from  Puente,  writes:  "Stellt 
man  diese  verschiedenen  Bildungen  aus  der  Nahe  der  Padrebucht  zusammen,  so 
gelangt  man  zu  der  Auffassung,  dass  dieses  Gebiet  frische  Spuren  vulkanischer 
Tatigkeit  aufvveist"  (p.   26). 

On  the  naked  sand  we  found  many  living  beetles  (no  reference  is  made  to  them 
in  vol.  Ill)  and  empty  shells  of  four  species  of  landshells,  Fernandezia  tryoniY''\%- 
bry,  Succinea  fernandi  Reeve  (also  in  sand  at  Tierra  Blanca),  S.  texia  Odhner  and 
S.  semiglobosa  Pfeiff.  (also  in  sand  on  Santa  Clara).  These  delicate  creatures  of  the 
humid  forests  are  entirely  unfamiliar  to  these  dry  and  barren  surroundings,  and  it 
is  difficult  to  account  for  the  presence  of  these  shells  here  as  well  as  at  Tierra 
Blanca  and  on  Santa  Clara.  Are  they  a  testimony  of  a  more  humid  climate  per- 
mitting some  kind  of  brushwood  to  exist,  a  period  during  which  the  concretions 
mentioned  above  (p.  loi)  were  formed?  Or  did  dwarf  trees  such  as  Dendroseris 
I'ltoralis,  Rea  pruijiata  and  Clieiwpodinm  Saiictae  Clarae,  on  which  landshells  lived, 
grow  here  in  historical  time  but  before  goats  were  introduced?  Certain  facts  do  speak 
in  favour  of  this  theory.  A  few  specimens  of  Kea  and  Dendroseris  still  occur  on  the 
south  coast  of  Masatierra,  particularly  on  M.  Vinillo  and  on  M.  Juanango,  where  the 
goats  cannot  get  them.  What  did  this  now  barren  country  look  like  when  the  islands 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


Fig.  45.  View  from  Puente,  looking  S.  Wind-eroded  tuff  beds  and  sand.  Behind,  Pta  O'Higgins 
with  its  morro,  in  the  background  Santa  Clara.  —  Photo  K.  Backstrom  "/^  191 7. 


Fig.  46.  Mobile  sand  on  Puente,  in  the  foreground  a  dense  growth  of  weeds,  mainly  Clienopo- 
dlum  multifidum.  C.  S.  as  measure.  —  Photo  K.  Backstrom  ^^/^  191 7. 


32 


C.   SKOTTSBERG 


were  discovered?  Not  as  to-day  I  am  sure,  but  more  or  less  like  Morro  del  Spartan 
at  Santa  Clara,  as  already  suggested  by  Johow  p.  261.  I  would  think  that  80,  in 
some  valleys  90  per  cent  of  the  soil  is  now  occupied  by  introduced  weeds.  We 
have  no  reason  to  believe  that  there  was  a  desert  when  the  immigration  of  aliens 
began. 

It  is  to  be  regretted  that  we  did  not  study  how  the  shells  found  did  occur, 
if  only  on  the  surface  or  also  deeper  down  embedded  in  the  sand.  And  w-e  have 
to  find  out  if  landshells  live  on   the  plants  mentioned  above. 


With  Pta  dc  la  Is/a  we  reach  the  end  of  the  north  coast.  Time  did  not 
permit  us  to  visit  the  small  peninsula  W  of  El  Puente,  and  we  shall  now  pro- 
ceed along  the  south  side  of  Masatierra. 

P/a  (y///[i^x///s,  watched  by  a  nameless  morro  (small  skerry),  and  the  inhosp- 
itable coast  cliffs  of  BaJiia  Carvajal  are  seen  in  fig.  47.  With  a  NW  wind 
boats  engaged  in  langost  fishing  at  Santa  Clara  find  shelter  here.  From  Pta 
Agiiabuoia  to  Co  Xegro  the  coast  shows  several  well-marked  bights. 

Haliia  Ticna  Blanca  or  Tierras  Blaiicas  has  its  name  from  the  white  sand 
above  the  bay.  Just  as  the  other  bays  on  this  coast  it  is  bordered  by  cliffs  and 
beaten  by  a  never-resting  surf.  To  land  anywhere  on  this  side  of  the  island  is 
possible  only  under  very  exceptional  conditions  (fig.  48). 

Pta  I.a>\i!^a  (Long  Pt.)  separates  Tierra  Blanca  from  the  next  bay;  in  the 
background  rises  a  rounded  hill  which  I  take  to  be  identical  with  Guzman's  Co  Enri- 
que. From  a  distance  it  seems  to  be  formed  by  basalts  of  the  higher  horizons,  but 
no  specimens  were  brought. 

Pallia  Chupones  derives  its  name  from  "chupon",  in  Chile  a  vernacular  name 
for  (ireii!;ia  spliacelata  Reg.  (Bromeliaceae),  which  has  edible  fruits  (chupar  =  suck), 
and  ap|)lic(l  by  the  fishermen  of  Masatierra  to  the  extremely  rare  Hespevogreigia 
as  well  as  to  Ocliagaria;  in  this  case  the  former,  an  inhabitant  of  the  wettest 
and  loftiest  ridges,  is  not  to  be  thought  of,  whereas  the  latter,  a  typical  xerophyte, 
very  likcl)-  occurs  here,  even  if  we  did  not  observe  it  W  of  Tres  Puntas,  where 
it  co\  crs  rock  faces  (see  Skottsb.  3  PI.  97).  The  slopes  round  the  bay  are  grass- 
land, in  tile  western  half  almost  pure  Aveiia  barbata,  otherwise  with  extensive 
|)atchcs  of  the  native  StipctiDii.  All  streambeds  in  this  western  section  are  dry 
most   of  the   year. 

Ijuna  Psiarpa(ta\  385  m  high  where  we  crossed  it  near  the  main  range, 
separates  B.  Ciiuj)ones  from  \'illagra  Bay,  this  taken  in  a  wide  sense.  Off  the 
j)<)int  lies  Mono  I'ihillo  (vifiilla  means  "small  hill  planted  with  vines",  but  the 
name  must  refer  to  something  quite  different  in  this  case);  the  gradient  of  the 
surface  shows  tiie  tilt  of  the  lava  beds  (fig.  49). 

Pallia    (tr     Mllagra    and    its    valleys,    the    bay    taken    in    a    wide  sense  and 

\\\  my  tield  notes  I  called  this  ridge  "Cordon  ?2scarpado",  and  this  name  also  appears 
on  tlic  Chilfan  chart,  and  on  my  map  I  had  used  the  same  name  for  the  ridge  between  Pangal 
and  <^).  Mincro,  and  as  it  was  published  in  this  sense  Skottsb.  3  pp.  890,  915  ,  I  have  renamed 
the  other  ridge  Loma   Kscarpada. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


^33 


Fig.  47.  View  from  Tuente  toward  liahia  Carvajal  and  Pta  O'Higgins.  Behind,  Santa  Clara. 

Photo  K.  Backstrom  ^^j-^  1917. 


Fig.  48.  Bahia  Tierra  Blanca  seen  from  Pta  Larga.  —  Photo  C.  Skottsberg  ^/^  191 7. 


34 


C.   SKO'lTSBERG 


V\^^.  49.    \'ie\v    from    Portezuelo  de  Villagra  looking  SW,  with  Santa  Clara  in  the  background. 

Morro    V'inillo    in    the  middle,  right  the  south  coast  of  Masatierra  from  Bahia  Villagra  to  Pta 

O'Higgins.  —  Photo  C.  Skottsberg  ^^l^^  1916. 


extending  from  M.  Vinillo  to  Los  Chamelos.  The  broad  slopes  E  of  Loma 
Escarpada,  locally  known  as  Los  bajos  de  Villagra,  are  strewn  with  lava  boulders 
and  furrowed  by  dry,  stony  streambeds.  In  the  background  two  very  conspicuous 
mountains,  already  seen  on  the  north  coast,  Co  Tres  Puntas  and  Co  Chumacera 
(Rowlock  Mt.,  with  a  deep  vertical  slit),  rise  to  a  height  of  perhaps  500  m 
(Branchi's  figure  650  is  certainly  too  high).  In  the  dry  streambed  below  Tres 
Tuntas  we  found  a  patch  of  pangue,  which  reaches  its  farthest  west  here.  Chu- 
macera looks  like  an  enormous  rock  slab  standing  on  end.  At  the  foot  water 
was  found  also  during  the  dry  season,  and  here  is  the  westernmost  luma  forest 
on  this  side,  and  below  a  threshold  (see  above  p.  97  and  Skottsb.  3  fig.  36  on 
|).  896)  with  a  small  waterfall  a  grove  of  BoeJimeria.  The  next  gully  has  forest 
down  to  about  300  m  above  the  sea.  Fig.  50  gives  a  good  idea  of  the  nature 
of  this  country. 

( )n  all  the  earlier  charts  and  maps  the  bay  is  presented  as  forming  a  regular 
ciu've.  but  the  aerial  survey  proved  that  this  is  not  the  case;  see  fig.  2.  The 
coast  cliffs  arc  lower  here  than  farther  west,  but  there  are  very  few  places  where 
it  is  |)()ssible  to  get  down  to  the  water.  It  can  be  done  not  far  from  Cerro 
Negro,   where  we   found   access  to  the   beach. 

X'illagra  is  watered  1)\'  three  permanent  streams  and  densely  wooded.  The 
scener)-  is  even   more  grand   than  on  the  Cumberland  side,  with  the  sequence  of 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


135 


Fig.  50.    View   from    Portezuelo   de   Villagra  lookiiig  W.  From  W  to  E  Cerros  Enrique,  Tres 
Puntas  and  Chumacera.  —  Photo  C.  Skottsberg  "/^  1917. 


summits  from  Co  Agudo  (sharp)  to  Mt.  Yunque,  more  imposing  perhaps  from  this 
side  than  from  any  other  (fig.  51).  A  peak  between  Agudo  and  Chumacera  is 
called  Oreja  del  Conejo  (Rabbit's  ear)  by  Guzman.  The  higher  slopes  are  pre- 
cipitous, with  gradients  of  60 — 70°  and  sculptured  with  numerous  hanging 
gullies,  carrying  water  after  every  rain,  when  many  little  cascades  tumble  down 
from  the  summit  of  Mt.  Yunque  (fig.  2)  between  the  carpets  of  ferns  and  pangue. 
The  Villagra  valleys  have  not  been  levelled  by  erosion  as  much  as  Valle  Colo- 
nial, V.  Ingles  or  V.  Anson;  the  inclination  in  the  middle  and  lower  sections  is 
20 — 30°.  The  forest  in  Q.  de  la  Choza  (responsible  for  this  name  is  a  small 
shack  below  the  lower  timber-line,  I.e.  PI.  85:  i)  and  on  the  slope  of  Mt.  Piramide 
(I.e.  PI.  91)  is  primeval;  it  extends  up  to  the  level  of  the  Villagra  pass  and  is 
very  wet  and  rich  in  species.  Some  maqui  is  seen  here  and  there  at  lower  eleva- 
tions. The  forest  comes  to  a  sudden  stop  200 — 250,  in  cases  300  m  above  the 
sea.  On  a  former  occasion  (3  p.  895,  PI.  85:  i)  I  have  discussed  the  nature  of 
this  timber-line.  Below  the  forest  degraded  grass-land  with  foreign  grasses  and 
herbs  dominates  over  the  natural  Siipeium.  Along  the  streams  a  fringe  of  pangue 
runs  down  toward  the  sea  (I.e.  PI.  85:  2).  Nobody  lives  in  Villagra,  but  it  is 
grazed  over  by  cattle  on  the  lower  slopes. 

Mt.  Yunque  has  already  been  described;  I  shall  add  here  what  Tenz  I.e. 
has  to  say  about  the  summit.  "Se  ve  arriba  una  altiplanicie  muy  extensa  y  suave- 
mente    inclinada  hacia  oeste  a  poca  profundidad,  de  forma  rectangular,  rodeada 


^36 


C.   SKOTTSHERG 


FiK-   5'-   1-1   N  uiKiiie  seen  from  the  \'illagra  slope  c.   175  m  above  sea  level.  —  Photo  C.  Skotts- 

berg  «/i   191 7. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


137 


Fig.  52.    South    coast  of  Masatierra  from  Cerro  Negro  to  beyond  Corrales  de  Molina.  After  a 

water-colour  sketch  by  the  author. 


de  cordones  en  los  cuales  sobresalen  varias  cimas.  Desde  ellas  nacen,  en  distintas 
direcciones,  quebradas  en  que  corren  cristalinas  vertientes."  The  buttress  project- 
ing S  is  prolonged  to  a  ridge  ending  in  a  low  cone,  Co  Negro,  190  m  high  (one 
reading  only;  I.e.  PL  92:  1). 

From  El  Yunqiie  to  EL  Verdugo  the  main  ridge  of  the  island  presents  the 
picture  of  a  sky-high  rock  wall  (figs.  2,  11,  52).  E  of  Mt  Yunque  several  nearly 
vertical,  trough-shaped  gorges  have  been  dug  out,  each  with  a  cascade  and  a 
patch  of  forest  and  known  as  Corrales  de  Molina  (corral  =  enclosure;  probably 
named  for  Padre  IGNACIO  Molina,  an  Italian-Chilean  naturalist  of  the  i8th 
century  and  author  of  a  Compendio  in  which  3  plant  species  from  Masatierra 
were  mentioned).  Goat-hunters  cross  the  ridge  here  and  descend  into  the  gullies, 
which  have  been  described  with  much  detail  by  Weber  who  went  there  several 
times.  Fig.  52  shows  that  it  is  no  easy  going.  Hence  follows  a  naked  vertical 
wall,  exposing  the  regularly  stratified  lava  (fig.  52,  right).  Along  the  shore  Playa 
Larga  extends,  marked  "Low  beach"  on  the  English  chart,  presumably  a  low 
abrasion  ledge.  The  country  farther  E  is  a  succession  of  clifi"  walls  and  gorges; 
see  figs.  2  and  11.  The  sinister  name  ir/  Fifr^//^^  (executioner,  hangman,  fig.  11) 
reminds  of  the  dangers  on  this  coast,  where  no  light  warns  the  sailor. 

9  ~  537351   The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.  Vol.  I. 


38 


C.   SKOTTSRERG 


^* 'h-  53-  'Santa  Clara  from  the  air,  looking"  SE.  —  Photo  B.  Frodin  ^/^   1952. 


Santa  Clara. 

Santa  Clara  or  Goat  Island  is  a  barren,  desolate  islet  separated  from  Masa- 
tierra  by  a  shallow,  about  1500  m  wide  strait,  dreaded  for  its  strong  currents 
and  turbulent  waters.  The  depth  is  19 — 20  fathoms  according  to  Anson's  map; 
I  sounded   20 — 45  m. 

The  older  literature  gives  little  information  about  this  island,  but  it  was 
described  in  some  detail  by  Guzman  pp.  48 — 53,  to  whom  the  reader  is  referred. 
Various  bays  and  morros  are  mentioned,  with  names  probably  given  by  the 
fishermen.  The  circumference  of  the  island  is  said  to  be  9  km,  the  area  500 
hectares.  Measurements  on  the  new  map  gave  a  length  of  3.5  km,  a  maximum 
width  of  1.25  km  and  an  area  of  2.5  sq.  km.  This  map  is  based  on  the  aerial 
survey  of  1952  and  on  Mr.  Frodin's  photographs  (fig.  53).  The  length  profile  is 
seen  in  its  full  extension  on  fig.  48.  A  comparison  shows  that  all  the  older  maps 
are  very  defective. 

The  island  is  everywhere  bordered  by  steep  cliff*  walls  (fig.  53).  Above  is 
a  sandy  table-land,  studded  with  hills,  of  which  the  easternmost,  incorrectly 
placed  on  other  maps,  is  about  375  m  high  (366,  Lopez).  Guzman  calls  it  Co 
Negros,  but  as  we  have  one  Co  Negro  on  the  south  coast  of  Masatierra,  I 
named  it  Cerro  Jolioic  in  commemoration  of  the  author  of  the  well-known  work 
on  the  flora  of  Juan  Fernandez.  Close  to  the  coast  are  several  morros.  The  west 
side  of  the  island  is  beaten  by  a  heavy  surf  and  inaccessible,  but  on  the  inner 
side,  behind  M.  del  Spartan,  landing  is  —  but  not  always  —  possible. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  I  39 

On  Anson's  map  of  1741  Santa  Clara  is  shown  as  covered  with  some  kind 
of  arboreous  vegetation.  Walter  does  not  tell  if  a  landing  was  made,  but  the 
name  Goat  I.  shows  that  there  were  goats  on  the  island,  and  Ulloa's  map  of 
1742  leaves  it  treeless.  Just  as  Johow  we  found  the  place  very  barren  and  the 
vegetation  mostly  formed  by  weeds  with  Ai/ejia  in  dominance,  all  very  dry 
during  the  summer.  When  Giinther  says  that  there  were  "algunos  arboles  en  la 
pendiente  del  este"  he  either  did  see  some  specimens  o^  Dendroscrjs  on  iha  c\\f(s 
above  the  water  or  refers  to  Morro  Spartan  (also  called  M.  de  los  Alelfes; 
fishermen  are  said  to  have  collected  seeds  of  Matthiola  in  B.  del  Padre  and  to 
have  scattered  them  on  Santa  Clara).  Relics  of  the  original  flora  found  a  refuge 
on  the  morro,  which  is  separated  from  the  island  by  a  channel  less  than  10  m 
wide  and  blocked  by  huge  boulders  which  are  exposed  at  low  tide.  The  current 
rushes  through  the  narrow  channel,  and  to  judge  from  the  vegetation  the  goats 
are  unable  to  cross.  I  believe  that  Johow  (p.  261)  was  right  in  assuming  that, 
before  the  introduction  of  this  pest,  Santa  Clara  must  have  looked  much  like 
M.  Spartan.  The  altitude  of  the  island  is  sufficient  to  catch  the  trade  clouds,  as 
shown  on  fig.  4,  and  we  are  told  that  rains  are  not  unfrequent  during  the  winter 
months  (Johow  p.  261),  giving  origin  to  an  abundant  vernal  flora  of  annuals. 
During  our  brief  visit  in  January  we  did  not  see  any  trace  of  water;  but  we 
read  in  Instrucc.  naut.  p.  230  that  a  stream  leaps  over  the  cliff  at  the  NW  point, 
a   statement    repeated    by    Guzman   who  calls  it   Chorro  de  Doiia  Maria  (p.   50). 

To  judge  from  our  passing  observations,  the  geology  is  the  same  as  of 
western  Masatierra,  uncounted,  light  yellowish  or  brownish  volcanic  beds  tra- 
versed by  numerous  vertical  dikes  of  a  hard  gray  basalt.  I  regret  having  neglected 
to  bring  samples  of  the  lava  beds  for  comparison  with  the  rocks  from  Puente, 
The  dike  rock  was  described  by  Hagerman  p.  28  and  found  to  be  identical  with 
the  basalt  from  the  top  of  Co  Negro.  The  sample  came  from  a  dike  on  M.  Spartan. 
We  landed  at  the  foot  of  this  dike  which  forms  a  flight  of  steps  leading  to  the 
table-land  of  the  morro;  see  Skottsb.  3  p.  924  and  PI.  103.  The  dikes  project 
as  flat  slabs  above  the  softer  beds. 


Masafuera,  former  surveys  and  maps. 

Until  our  visit  in  19 17  Masafuera  was  much  less  known  than  her  sister  island. 
It  had  been  inhabited  more  permanently  only  during  the  period  of  the  penal 
settlement  1909 — 19 13,  a  misfortune  that  befell  this  ocean  castle  a  second  time 
in   1927 — 1930.  Only  few  scientists  have  visited  Masafuera. 

Of  the  early  navigators  few  paid  a  visit  to  this  island.  One  of  Commodore 
Anson's  captains,  on  his  way  to  Cumberland  Bay,  happened  to  come  up  under 
Masafuera  and  reported  that  the  island  was  not,  as  former  navigators  had  imagined, 
a  barren  rock,  but  "almost  every  where  covered  with  trees  and  verdure,  and 
was  near  four  miles  in  length".  He  had  not  been  able  to  land,  but  added  that 
"it  appeared  to  him  far  from  impossible,  but  some  small  bay  might  be  found 
on  it,  which  might  ofl"er  sufficient  shelter  for  any  ship  desirous  of  refreshing 
ashore"   (Walter  p.    134).  As  four  ships  of  the  squadron  were  missing  the  Com- 


140 


C.  SKOTTSBERG 


MASAFUERA 


Vicente  Forrai 


Kodado  del  Sindilo 


Scale  1:100000 


GENERALSTABENS  LITOGRAFISKA  ANSTALT 
STOCKHOLM   1953 


Fig.   54.  Map  of  Masafuera.  After  Skottsberg. 


modore  sent  a  sloop  to  Masafuera  to  look  for  them  there,  and  when  she  returned 
after  a  fruitless  search,  during  which  the  island  was  circumnavigated,  a  report 
was  drawn  up  with  the  first  real  description  of  the  island.  Anchorage  had  been 
found  on  the  north  side  close  to  the  shore  but  protected  from  S  only  (I.e.  p.  156). 
PI.  X.XI  is  a  view  of  the  Xl^  side  of  "Masa-Fuero",  as  the  name  is  spelled.  It 
shows  the  table-land  uj)  to  the  highest  hills  well  forested,  and  we  have  little 
reason  to  doubt  that  such  was  the  situation  200  years  ago.  Four  places  along 
the  coast  are  marked  with  the  letters  a,  b,  b,  and  c\  a  seems  to  indicate  a  reef, 
b  and  b  arc  the  entrances  to  two  of  the  canyons.  I  believe  that  a  also  indicates 
(J.  Casas  and  that  b  are  (J.  Ovalo  and  (,).  Sanchez;  c  is  the  waterfall  coming 
down  from  ().  Larga.  PI.  XXII,  pretending  to  show  the  west  side,  is  less  easy 
to  read.  1  think  tliat  it  re[)resents  the  north-west  section  of  the  coast  from  Cabo 
Xorte  south  toward  Loberia  X'ieja,  but  nothing  like  the  perfectly  cylindric  tower 
on    the    extreme    right  in  the  picture  exists  —  the  only  thing  I  can  think  of  is 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  I  41 

a  very  irregular  rock  close  to  the  beach  in  Loberia.  The  promontory  in  the 
middle  is  Buque  Varado.  Groups  of  trees  are  seen  only  above  this  place  and 
above  Loberia  Nueva. 

Ulloa  found  the  island  very  inhospitable:  "La  Isla  de  afucra  de  Jiam  Fer- 
ftajtdea  es  toda  muy  alta,  y  tan  escarpada,  y  escabrosa  que  no  tiene  parage 
conmodo  para  desembarcar"  (p.  287).  On  a  former  occasion  (3  p.  796)  I  quoted 
the  narratives  of  Byron  and  Carteret.  The  island  was  well  wooded  in    1765. 

The  hydrographic  expeditions  despatched  by  the  Oficina  in  Valparaiso  paid 
little  attention  to  Masafuera.  When  Lopez  reports  that  the  island  was  "cubierta 
de  arbolado"  in  1875  (p.  6^)  he  must  have  included  the  Dkksonia  jungle  on 
the  slopes  of  Mt.  Inocentes.  Johow  (p.  96)  asserts  that  the  luma  was  common 
all  over  the  island  except  near  the.  coast  and  in  the  highland,  but  this  does  not 
necessarily  mean  that  it  formed  extensive  woods.  We  find  the  same  statement 
in  Instrucc.  naut.  of  1896.  It  is  not  known  when  the  destruction  of  the  forest 
started  in  earnest,  but  it  is  probable  that  the  exploitation  of  the  sandal-wood, 
which  led  to  its  complete  extermination,  had  serious  consequences  for  the  forest 
as  such.  The  names  of  two  valleys  and  a  place  on  the  south  coast  testify  that 
a  species  of  Sanialuni  grew  on  the  island.  The  possibility  that  forest  fires  have 
ravaged  the  woods  must  not  be  forgotten.  Foreign  grasses  spread  into  the  cleared 
spaces  and  prevented  the  germination  of  the  seeds  of  the  native  trees.  The 
direct  influence  of  the  goats,  introduced,  I  believe,  during  the  17th  century, 
remains  to  be  found  out.  They  greedily  devour  the  arboreous  Compositae  and 
the  endemic  herbs,  but  I  cannot  tell  if  they  eat  the  leaves  of  the  luma,  naranjillo 
and  canelo.  I  can  testify  from  my  own  experience  that  there  was  better  forest 
in  1908  than  in  191 7,  an  undisputable  consequence  of  the  activity  of  the  con- 
victs during  the  intermediate  period.  Nevertheless  Giinther  (1920)  repeats  the 
old  statement  that  Masafuera  was  covered  with  trees. 

In  1895  the  Oficina  Hidrografica  published  the  first  map  showing  the  prin- 
cipal topographical  features.  New  dates  had  been  provided  by  Johow.  During 
our  visit  in  191 7  we  soon  discovered  that  this  map  was  too  defective  to  be  used 
as  a  basis  for  a  map  of  the  vegetation,  and  we  did  what  we  could  to  correct 
it.  The  result  was  a  sketch  put  at  the  disposal  of  the  Oficina,  which  used  it 
for  a  new  chart.  When  the  flight  over  the  islands  was  undertaken  in  1952,  Masa- 
fuera was  covered  by  clouds.  A  new  map  is  urgently  needed. 


Main  geographical  features. 

In  shape  and  general  appearance  Masafuera  differs  profoundly  from  her 
sister  island.  The  island  (fig.  54)  is  a  solid  rectangular  block  with  slightly  rounded 
corners,  tilted  NE  so  that  the  escarpment  along  the  west  side  is  very  much 
higher  than  on  the  east  side,  where  it  is  quite  respectable  (figs.  55  —  57).  I  he 
table-land  is  traversed  by  numerous  parallel  deep  valleys  running  NK — L  (figs. 
57,  58),  the  high  west  wall  by  a  number  of  precipitous  gorges;  this  side  has 
much  the  same  appearance  as  the  south  side  of  Masatierra  E  of  Mt.  Yunque 
(fig.    52). 


42 


C.   SKOTTSBERG 


Fi) 


%-*j 


W.  Quebr.  de  las  Casas  at  the  extreme  left.  —  Photo 
Backstrom  Feb.   191 7. 


The  question  may  be  raised  if  not  the  eroding  forces  have  been  more  active 
in  bygone  times  than  they  appear  to  be  at  present  when  so  many  of  the  streams 
are  dry  or  carry  very  Httle  water  during  the  summer  half  year.  If  it  is  true,  and 
this  is  believed  by  many,  that  the  Ice  Age  was  characterized  by  high  precipita- 
tion values,  the  climate  of  Juan  Fernandez,  otherwise  not  much  influenced  by  the 
glaciation  in  the  southern  Andes,  must  have  been  more  rainy  than  now  and,  as 
a  consecjucnce,  the  eroding  forces  stronger.  The  end  of  the  Pleistocene  left  the 
islands  much   as  we  see  them  now. 

The  coast  lacks  bays,  there  is  no  harbour,  not  even  a  sheltered  cove  to 
accomodate  small  craft.  The  landing  places  from  where  the  interior  can  be  reached 
are  the  entrances  to  O.  Sanchez  and  O.  Casas,  but  only  under  favorable  con- 
ditions with  regard  to  wind  and  sea.  It  happens  that  an  expedition  has  to  return 
with  its  errand  unaccomplished.  Abrasion  has  created  a  terrace  along  the  west 
side,  continued  by  a  reef  studded  with  rocks  (fig.  59).  At  the  foot  of  the  coast 
cliffs  a  talus  has  accumulated,  sloping  down  to  a  beach  where  the  surf  washes 
out  the  finer  material  leaving  the  boulders.  It  is  possible  to  scramble  along  the 
shore  from  ().  Sanchez  south  round  the  island  and  from  there  north  along  the 
west  side  to  the  north  extremity  of  the  Loberia  Vieja  plain,  separated  from  the 
Loben'a  Vcntana  beach  by  an  impassable  obstacle.  Figs.  60  and  61  show  that 
it  is  no  easy  going,  and  with  a  heavy  swell  it  may  not  be  possible  to  pass  the 
southeast  corner  of  the  island.  The  opportunity  to  make  this  circuit  round  ^/^ 
of  the   shore    and    to    proceed    from    one    canyon    entrance    to    the    next  greatly 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ   ISLANDS  I  43 

simplifies    matters    for    the    surveyor    where,    in    contradistinction    to    Masatierra, 
boats  cannot  be  used  to  move  from  one  place  to  another. 

The  longitudinal  axis  is  directed  N  by  W— S  by  E.  The  statements  regarding 
the  size  of  the  island  vary  a  great  deal:  9  miles  (14.4  km)  by  4  miles  (6.4  km)  with 
a  circumference  of  about  46  km  and  an  area  of  85  sq.  km,  all  according  to  Lopez; 
11.7  by  7.4  km  (Johow's  map);  Branchi,  10  by  6  km,  area  84  sq.km;  Gunther, 
9.25  by  6  km,  and  Guzman,  17(1)  by  7  km,  a  circumference  of  55  km  and  an 
area  of  85  sq.  km.  The  figures  taken  from  my  map  are  10.3  by  6.2  km  and  the 
area   57.6  sq.  km. 

Geology  and  morphology. 

In  1908  Ouensel,  the  only  geologist  who  ever  visited  Masafuera,  made  a 
hasty  survey  of  the  rocks  along  the  coast  from  Q.  Casas  to  Loberia  Vieja  and 
north  of  Casas  from  the  shore  to  perhaps  1300  m  altitude.  His  field  notes  and 
specimens  served  him  for  a  description  of  the  principal  geological  features.  In 
structure  the  two  islands  differ  considerably.  Both  are  built  up  of  innumerable 
volcanic  beds  of  varying  thickness  and  tilted  about  20°,  both  exhibit  two  more 
or  less  distinct  horizons,  both  are  traversed  by  vertical  dikes,  but  here  the 
resemblance  ends.  On  Masatierra  the  lower  strata  consist  of  lavas  very  rich  in 
olivine,  interbedded  with  tuftaceous  deposits,  deeply  weathered  and  denuded, 
resulting  in  the  formation  of  V-shaped  valleys.  The  lower  horizon  of  Masafuera 
consists  of  hard,  dark  gray  to  black  vesicular  feldspar  basalts  (Quensel  2  p.  58),  less 
rich  in  olivine  and  very  resistant  to  the  denuding  forces.  These  rocks  prevail 
up  to  approximately  1000  m.  My  notes  from  191 7  contain  a  statement  that  W 
of  Vicente  Porras,  where  Playa  Larga  begins,  a  bed  of  "conglomerate"  was 
observed,  traversed  by  harder  dikes,  but  unfortunately  no  specimen  was  pre- 
served. In  a  paper  not  quoted  by  Quensel,  who  did  not  mention  the  occurrence 
of  agglomerate  beds  on  Masafuera,  R.  A.  Philippi  described  14  rock  samples  col- 
lected by  Germain,  most  of  them  "losen,  von  der  Hohe  heruntergerollten  Blocken 
entnommen".  I  doubt  that  anybody  will  be  able  to  identify  these  rocks  from 
Philippi's  unscientific  description,  but  two  of  the  samples  suggest  the  occurrence 
of  tufi"aceous  deposits:  they  consist  of  "Rapilli"  cemented  together. 

At  about  1000  m  a  lava  bed  of  a  different  aspect  was  met  with,  a  light  gray, 
"iddingsite-bearing  phyric  olivine  basalt"  (Quensel  2  pp.  58 — 63).  Between  1000 
and  1 100  m  a  rock  of  quite  another  type  occurred,  described  as  a  light  yellow- 
ish green  soda  trachyte.  Apparently  a  closely  related  type  is  found  also  at 
lower  levels;  boulders  of  this  rock  are  so  common  in  the  talus  along  the 
south  shore  that  the  place  was  called  Tierras  Blancas.  Quensel  arrived  at  the 
conclusion  that  "lavas  of  soda-trachytic  composition  have  been  emitted  at  different 
times  and  that  they  have  alternated  with  lavas  of  more  normal  basaltic  com- 
position"  (2  p.  66). 

The  upper  horizon,  from  about  iioo  to  1400  m,  is  formed  by  dense,  ash 
gray  feldspar  basalts,  which  probably  form  the  whole  upper  complex  of  lava 
beds    with    exception    of  the    summit    of  Mt.  Inocentes;   these  rocks  very  much 


44 


C.   SKOTTSBERG 


m-^^' 


V\^.  56.   East  coast  of  Masafuera  between  Pta  Negra  and  Playa  Ancha,  with  entrance  to  Q.  del 
Varadero.  —  Photo  C.  Skottsberg  21/^  19 17. 


Fig.   57.    Part    of  the  east  side  of  Masafuera  seen  from  the  air.  The  highland  is  covered  with 
thick  fog.  —  Photo  B.  Frodin  8/4  1952. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISI-ANDS 


45 


Fig.  58.   View   from    the   high   land   above    Las    Chozas  looking  toward  the  Ovalo,  Pasto  and 
Sandalo  valleys.  —  Photo  C.  Skottsberg  3/3  1917. 


^^*mA 


'"^f^mt^ 


Fig.   59.  The  shore  at  Loberia  Vieja  with  a  crag  on  the  reef.  —  Photo  K.  Backstrom  1V2  ^9^7- 


146 


C.   SKOTTSBERG 


Fig.  60.  At  the  landing-place  in   Las  Casas.  Our  party  is  about  to  embark,  assisted  by  the  crow 
of  the  schooner.  —  Photo  K.  Backstrom  1^/3   191 7. 


Fig.  61.  As  tig.  60,  showing  ijcach  ot  large  boulders.  —  Photo  K.   Backstrom  "/j   1917. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


f47 


Fig.  62.    Slope  of  Mt.  Inocentes  seen  from  Cordon  Atravesado,  c.  1350  m  ctouvc  sea  level.  — 

Photo  C,  Skottsberg  ^^/j  191 7. 


resemble  the  hard  basalts  between  400  and  600  m  on  Masatierra  (I.e.).  The  sum- 
mit of  Mt.  Inocentes  consists  of  a  still  different  rock,  a  slaggy  olivine  basalt 
supersaturated  with  iron  oxides  (I.e.  p.  70).  The  extension  and  thickness  of  this 
bed   is  unknown. 

As  on  Masatierra  numerous  vertical  dikes  traverse  Masafuera,  in  this  case 
in  W — E  direction.  The  rock  is  extraordinarily  rich  in  olivine  and  the  dikes 
may  represent,  at  least  in  part,  channels  for  the  beds  forming  the  summit  of 
the  island  (I.e.  p.   74). 

The  difference  in  geological  structure  between  the  two  islands  helps,  I  think, 
to  explain  the  profound  difference  in  morphology.  The  streams,  eroding  deeper 
and  deeper  into  the  land,  have  excavated  valleys  which,  in  their  distal  parts, 
are  V-shaped,  with  steep  sides,  which  rapidly  become  steeper  inland,  where  the 
perfect  U-shape  is  retained  in  the  canyon,  with  the  entire  narrow  width  of  the 
gently  sloping  bottom  occupied  by  the  streambed.  In  the  innermost  part,  where 
the  land  reaches  its  greatest  elevation  and  the  precipitation  its  highest  figures, 
lateral  erosion  has  widened  the  valley  and  a  fan-shaped  series  of  hanging  gorges 
has  been  formed.  The  canyon  ends  in  a  high,  almost  vertical  wall  with  a  water- 
fall. Figs.  63 — 65,  68 — 71,  73,  75  and  'j'j  serve  to  illustrate  the  valley  formation. 
In  the  northern,  drier  and  less  high  half  of  the  island  no  deep  canyons  were 
formed.    The  ridges  left  standing  between  the  valleys  are  very  unlike  the  ridges 


148 


C.   SKOTTSBERG 


^^^^^H 

1 

^ 

mki 

'^^i^^^^^^^^^^^^^^^i 

^^H 

-.^jE^^^^^^^ 

.SUhHwI^^I 

^^H 

^ 

1 

^^^.r^l^gi^.      '^^ 

1 

^ 

■ 

^^^^K^^jj^^^^^^l 

H 

^^^^^^HPf 

t 

H 

^^pn " 

•' 

^^9|^^^^^^| 

JM^H^^^^^^^^^^H 

H 

■ 

^"^^^9^^^! 

^^%w^^^^H 

g  rT^^B 

HB.- 

., 

^;f  f-rt,iv^.  ?  ~.iJ:-i<i^j^i^m^^M  .*    :f 

J 

CQ 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


149 


Fig.  64.  In  the  Casas  canyon  after  a  few  days  of  dry  weather.  The  rhubarb-like  plant  is  Gunnera 
Masafuerae.  —  Photo  K.  Backstrom  Feb.   19 17. 


Fig.  65.  Looking  down  into  the  interior  of  Casas  canyon  from  the  crest  of  Cordon  del  Barril. 

—  Photo  K.  Backstrom  1/3  19' 7- 


ISO 


C.   SKOTISBERG 


V        **-.        \  t " 

Fij(.  66.  The  grass  covered  table-iand  ijetween  the  Casas  and  Vacas  valleys,  looking  N.  The 
stone-building,  an  observation  post  overlooking  the  ocean,  dates  from  the  first  convict  settlement. 

—  Photo  C.  Skottsberg  ^^/j  191 7. 

in  Masatierra,  because  the  hard  basalt  layers  have  not  been  removed  by  denuda- 
tion, but  are  left  as  a  cover  right  down  to  the  sea,  forming  gently  sloping,  wide 
plains.  Farther  inland  the  ridge  gradually  narrows.  See  figs.  57  and  58.  Beds  of 
greater  resistance  form  ledges  along  the  valley  slopes  and  thresholds  across  the 
streambed. 


The  south  half  of  Masafuera. 

Shrouded  in  clouds  most  of  the  time,  Los  Inocefites  rises  above  the  west 
wall,  an  imposing  dome  forming  the  south  half  of  the  island.  Opinions  have 
differed  very  much  regarding  the  altitude:  2000  m  on  the  older  charts  as  well 
as  on  Johow's  map,  1836  (Lopez),  1850  (Viel ;  850  is  a  misprint),  2300  (v.  Rodt), 
1624  (Hranchi)  and  1840  (Guzman).  My  single  reading,  carefully  worked  out, 
gave  only  1500  m,  a  figure  possibly  too  low.  Toward  W,  N  (fig.  62)  and  NE 
the  summit  falls  precipitously;  it  is  connected  with  the  north  table-land  by  a 
ridge,  C.  Atravesado,  bordered  on  both  sides  by  a  precipice  and  so  narrow  that  the 
stretch  above  the  Vacas  valley,  where  a  pinnacle  rises  from  the  knife-edge,  is  not 
passed  by  the  goats.  y\bove  Casas  the  highest  point  is  c.  1370  m.  Where  Ermel 
got  the  idea  from  that  the  summit  is  covered  with  eternal  snow  (p.  113)  is  hard 
to  understand. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


15 


Fig.  67.  Cordon  del  Barril,  looking  W  toward  C.  Atravesado.  Note  goat  track  along  the  ridge. 

—  Photo  C.  Skottsberg  1/3  ^9^7- 


Q.  de  las  Casas,  once  the  headquarters  of  the  convict  settlement  and  the 
ordinary  landing-place,  is  called  O.  Baquedano  by  Guzman,  commemorating  the 
corvette  frequently  sent  to  the  outlying  islands  of  the  Republic.  From  the 
beach,  bordered  by  lofty  headlands  down  to  the  water,  we  enter  the  valley,  which 
is  about  200  m  wide  here,  with  slopes  receding  under  an  angle  of  40 — 45°  (fig, 
63).  It  gradually  narrows  so  that  the  streambed  offers  the  only  access  into  the 
canyon.  After  a  downpour  the  stream  overflows  and  the  valley  is  closed;  a  couple 
of  days  with  dry,  sunny  weather  and  the  bed,  filled  with  boulders  of  all  sizes, 
resting  on  the  bedrock,  lies  dry,  with  pools  of  crystal  clear  water  in  the  depres- 
sions and  back  of  the  thresholds  (fig.  64).  The  most  spectacular  part  of  the  gorge 
begins  about  i  km  from  the  entrance,  where  the  width  at  the  bottom  may  fall 
below  10  m,  while  the  walls  are  many  hundred  m  high  (fig.  65).  From  a  botan- 
ist's viewpoint  Casas  is  one  of  Nature's  conservatories;  see  Skottsb.  3  pp. 
639 — 640  and  Pi.  no.  Johow  called  this  place  "El  Pangal",  and  nowhere  is  the 
pangue  [Gunnera  Masafuerae)  more  luxuriant  (I.e.  PI.  iii).  About  2  km  from  the 


152 


C.   SKOTTSBERG 


FiJ,^  68.  Entrance  to  Quebr.  de  las  Vacas.  —  Photo  C.  Skottsberg  ^^j^  1917- 


entrance  and  210  m  above  the  sea  the  passage  is  obstructed  by  a  threshold  dam- 
ming up  a  deep  pool  which  cannot  be  passed  round,  and  into  this  pool  the 
water  comes  down  in  a  cascade.  Q.  Casas  drains  C.  Atravesado.  The  eroding 
power  of  the  current  is  great,  and  I  suppose  that  the  stream  is  more  or  less 
permanent  during  the  winter  half  year,  but  it  must  have  taken  a  very  long  time 
to  dig  a  gorge    1000  m  deep  through  the  hard  basalt. 

C.  del  luirril.  Between  Casas  and  Vacas  lies  a  sloping,  triangular  table- 
land, about  1.5  km  wide  along  the  top  of  the  coastal  escarpment,  which  is  125  m 
high  in  tliis  {)lace,  and  extending  west  c.  2  km,  where  the  cordon  proper  begins. 
The  ruin  seen  on  fig.  66  lies  190  m  above  the  sea;  the  tilt,  corresponding  to 
the  dip  of  the  lava  beds,  is  20°  E.  Three  small  gullies  break  the  monotony  of 
the  grass-land,  (.).  ("hica,  ().  Hlindado  and  O.  Cabreros.  Q.  Cliica  (-=  small)  is  very 
shallow  and  dry,  but  some  luma  trees  are  seen.  Q.  del  Bliiidado  (B.  means 
ironclad  cruiser,  but  may  have  a  very  different  significance  here)  is  much  larger 
and  tlie  water  comes  down  over  a  threshold  about  500  m  upstream  and  has  dug 
out  a  miniature  caiu'on.  Tiiere  is  good  forest  between  400  and  500  m  and  some 
groups  of  trees  a  little  farther  clown.  There  is  some  forest  also  in  Q.  de  los 
Cahreros  ((ioat-hunters'  valle)-).  Where  the  narrowing  rocky  Barril  ridge,  which 
has  its  name  from  a  barrel  sha|)ed  monolith,  begins,  the  elevation  is  730  m.  By 
and  by  the  crest  gets  very  narrow,  but  the  rise  is  gentle,  and  there  is  a  goat 
track    all    the    way  up  to  C.   Atravesado  (fig.  G'j).  The  ridge  slopes   50 — 60°  on 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  I  53 

both  sides,  and  we  look  down  into  the  Casas  and  Vacas  canyons  (figs.  65,  69). 
From  C.  Atravesado  the  road  N  to  Piano  de  la  Mona  lies  open. 

Q.  de  las  Vacas  (  =  cow;  this  place-name  is  old,  but  no  cattle  existed  on 
the  island  in  1908.  Reintroduced  in  1909,  they  had  been  killed  off  before  19 17 
and  were  introduced  again  in  1927).  Following  the  beach  south  from  Casas 
past  Pta  Langlois  we  reach  the  entrance  to  this  second  large  canyon,  which 
drains  the  highest  part  of  Mt.  Inocentes.  It  happened  that  when  Casas  was  dry, 
the  Vacas  stream  was  still  alive.  The  entrance  is  quite  picturesque  thanks  to 
the  deep  side-valley  separated  from  the  main  quebrada  by  a  high  spur  ending 
in  a  sharp  peak  (fig.  68).  The  interior  has  the  same  morphology  as  Casas;  fig. 
70  shows  the  drainage  basin  viewed  from  the  Barril  ridge,  fig.  71  a  typical,  low 
threshold  of  harder  basalt.  Some  witches'  cauldrons  were  observed  (fig.  72). 

Little  Vacas  is  accessible  to  a  point  about  i  km  from  the  sea,  where  a  very 
high  cliff  barrier  impedes  further  progress. 

The  easiest  way  to  reach  the  table-land  S  of  Vacas  is  to  climb  a  low  bar- 
ranca at  Play  a  Ancha,  a  level  stretch  of  stony  beach,  from  where  Q.  de  los 
Inocentes  with  its  untouched  forest  and  fern-groves  is  gained  without  difficulty. 
Crossing  this  valley  high  up  we  climbed  the  Inocentes- Vacas  ridge  which  leads 
to  the  top.  To  our  surprise  we  were  not  stopped  by  the  fern  forest  we  had  reason 
to  expect  judging  from  our  experience  farther  south;  the  stony  and  rocky  ridge 
was  covered  with  Alpine  heath  which  continued  right  to  the  Inocentes  summit. 
The  ridge  can  also  be  gained  from  Little  Vacas.  The  slope  is,  as  seen  on  fig.  68, 
steep,  but  it  is  terraced,  and  on  our  return  we  made  a  quick  descent  here. 

From  the  ridge  south  of  O.  Inocentes  a  good  view  is  obtained  across  the 
interior  of  the  narrow  Q.  Angosta  (fig.  73).  The  morphology  is  the  same  as  in 
Casas  and  Vacas,  but  the  valley  got  its  name  because  it  is  narrower  than  any 
other,  only  about  5  m  wide  at  the  bottom,  a  magnificent  blind  alley  where,  c.  i  km 
from  the  entrance,  we  are  confronted  with  a  precipice  and  waterfall  once  more. 
The  photograph  shows  the  columnar  structure  of  the  threshold.  At  the  foot  of  the 
waterfall  the  vegetation  is,  as  in  all  these  gorges,  luxuriant. 

Quebr.  del  Varadero  (varadero  =  ship-yard,  which  does  not  fit  here;  varar 
also  =  to  be  stopped,  and  a  strong  surf  may  prevent  passing  the  escarpment 
south  of  the  canyon  entrance).  Our  first  attempt  to  ascend  Mt.  Inocentes  was 
made  from  this  valley  at  the  place  seen  in  fig.  74,  where  we  gained  the  ridge 
at  400  m  above  see  level.  On  account  of  the  slippery  grass  the  climb  was  a  little 
hazardous.  To  begin  with,  the  ridge  itself  was  easy  until,  at  c.  740  m,  the  tree- 
ferns,  which  had  begun  to  appear  on  the  sides,  gathered  on  the  very  crest,  form- 
ing an  impenetrable  thicket.  Having  crawled  through  the  soaking  wet  barricades 
of  decaying  trunks  under  the  closed  roof  of  the  fronds  and  made  perhaps  500  m 
in  an  hour,  we  had  to  give  up.  The  altitude  was  c.  950  m  and  Dicksonia  con- 
tinued in  every  direction  as  far  as  we  could  see  from  a  solitary  canelo  rising 
above  the  fern  roof 

A  short  walk  into  the  Varadero  gorge  offers  a  strikingly  wild  scenery  (fig. 
75).  A  picture  of  the  short  and  narrow  side  valley  was  published  by  Quensel 
(2  fig.   3   on  p.  43).   Gradients  of  60—80°  are  the  rule  in  these  gorges. 

io~53735i   ^^^  ^<^i-  Hist,  of  Juan  Fernandez  and  Easter  I  si.  Vol.  I. 


54 


C.   SKOTTSBERG 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


55 


156 


C.   SKOTTSHERG 


Fig.  71.  A  low  threshold  in  the  Vacas  canyon.  —  Photo  C.  Skottsberg  ^^1^  191 7- 


^; 


I'ig.   72.  'rwo  witches'   cauldrons  in   the   \'acas  canyon.  —   I'hoto  C.   Skottsberg  "/j   1917- 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


57 


Fig.  73.  A  view  of  the  interior  of  Quebr.  Angosta,  seen  from  the  ridge  N  ol  lins  v.uicy,  550  m 
above  sea-level.  —  Photo  C.  Skottsberg  ^j^  191 7- 


T/ie  sotitJi  coast.  The  strip  of  beach  is  narrow  S  of  Varadero,  and  the  escarp- 
ment reaches  an  imposing  height  at  Pta  Negra  (fig.  56),  A  short  km  beyond 
this  point  a  gorge  with  a  cascade  is  passed,  Chorro  de  Dona  Maria,  whether  the 
same  lady  to  whom  the  chorro  on  Santa  Clara  was  dedicated  I  cannot  tell.  The 
talus  W  of  this  place  derives  its  name,  Tierras  Blancas,  from  the  light  colour  of 
the  deposits.  Along  Rodado  del  Sdndalo  (rodar,  to  make  a  turn)  the  land  rises  gently; 
the  shore  is  fringed  by  a  reef,  on  which  the  sea  breaks  with  a  thundering  roar 
(fig.  ']^.  At  Vicente  Porras  we  arrive  on  the  west  coast.  The  long  and  broad  abra- 
sion terrace.  Play  a  Larga,  was  referred  to  above  (p.  142);  a  level  plain  like  this  is  a 
rare  phenomenon  on  Juan  Fernandez.  At  Loberia  Vieja,  "the  old  sealing  grounds", 
where  some  fur-seals  were  still  to  be  seen,  the  width  is  i  1/2  cable  (277.5  "i)  ^^' 
cording  to  Giinther,  a  figure  in  good  accordance  with  our  estimation.  Several  hang- 
ing gorges  face  the  playa,  coming  down  from  Inocentes  and  C.  Atravesado,  and  two 
streams  find  their  way  across  the  plain  to  the  sea.  One  of  the  gorges,  described  as 
Q.  de  la  Loberia  Vieja  (Skottsb.  3  p.  941)  was  surveyed  by  us.  The  entrance  is 
barred  by  very  large  boulders,  and  one  has  to  climb  along  the  wall  to  get  into 
the  valley.  Quensel  published  a  photograph  of  the  entrance  (2  fig.  15  on  p.  58). 
The  short  canyon  ends  in  the  ever-present  cliff  wall  with  its  waterfall  (fig.  'j'i). 
The  altitude  of  the  valley  floor  is  only  c.  190  m. 

A   short    but    impassable    stretch    prevents    us    from   reaching  the  remaining 


158 


r.   SKOTTSHERG 


^= 


FiK.  74;  l^ntraiKc  to  Ouebr.  del  X'ar.ulcro,  from  where,  up  the  south  wall  (left)  an  ascent  was 
made.    The    knob   on  the  extreme  right  is  shown  in  fi(,^  75.  —  Photo  C.  Skottsberg  "/a   1917. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


159 


Fig.  75.  Looking  into  the  Varadero  canyon.  —  Photo  C.  Skottsberg  ^i/,^  1917. 


1  6o 


;kottsberg 


P^ig.  76.  Coast  of  Masafuera  between  Tierras  Blancas  and  Vicente  Porras. 

Strom  ^^/a   191 7. 


Photo  K.  Back- 


"loberias",  L.  \^entaua  (=^  window,  a  hole  in  the  rock),  L.  Nueva  and  L.  del  Buque 
I  'arado. 

Playa  del  Buque  Varado  ("the  beach  of  the  stranded  ship")  is  a  wide  tongue 
formed  by  the  talus  deposits;  probably  one  or  several  landslides  have  occurred 
here,  where  tlie  barranca  is  of  formidable  height  and  steepness  (fig.  78).  A  stream 
coming  from  the  gullies  above  has  cut  a  winding  bed  (fig.   82). 

I'jiseiiada  loltthi  is  a  shallow  open  cove  W  of  Cabo  Norte.  When  it  is  im- 
possible to  approach  the  east  coast  boats  anchor  here,  where  landing,  as  a  rule, 
meets  with  little  difficulty.  Consequently,  numerous  weeds  mix  with  the  native 
herbs  and  grasses  covering  the  slopes  at  the  foot  of  the  barranca.  Unfortunately 
Token  is  (juite  unfit  as  a  base  camp,  as  the  highland  cannot  be  reached,  nor 
is  there  a  j)assage  along  the  shore  to  O.   Sanchez. 


The  table-land  N  of  0-  de  las  Casas. 

The  Plauo  de  la  Mona  ("she-monkey's  plain"),  a  name  of  unknown  origin 
(there  are  no  monkeys  on  Juan  Fernandez),  and  the  rim  of  the  coast  escarpment 
can  be  reached  from  several  valleys.  The  usual  route  takes  us  up  a  zigzag  trail 
from  Casas,  across  a  dry  gully  and  through  the  large,  branched  and  well  for- 
ested O.  del  Mono  (another  cjueer  name)  to  the  abandoned  convict  settlement 
Las   Chozas,  situated  among  the  remnants  of  luma  forest   550 — 650  m  above  sea 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


6i 


Fig.  ^T.  Interior  of  the   Loberia  Vieja  canyon.  —  Photo  C.  Skottsberg  ^'/a  19^ 7- 


C.   SKOTTSHERG 


I•'i^^  7<S.    Pta    del    liii.in  and  Playa  del   Buque  Varado,  seen  from  the  sea  looking   S.  —  Photo 

C.  Skottsberg  ^^/g  191 7. 


level;  another  50 — 100  m  and  the  undulating  grass-,  fern-  and  pangue-covered 
Mona  plain,  crossed  by  many  little  gullies,  dry  during  the  summer,  lies  open  in 
every  direction  (fig.  79).  At  one  of  the  tributaries  to  O.  Pasto,  1130m  above 
the  sea,  a  pool  of  water  in  the  rocky  stream-bed  made  the  place  a  convenient 
camp  site  ("Campo  Correspondencia").  A  short  walk  brings  us  to  the  edge  of 
the  i)lain  and  to  Las  Torres  (Towers),  c.  1370  m,  and  Co  Correspondencia, 
c.  1420  m.  V\\r.  80  is  a  view  of  one  of  the  Loberia  gorges  seen  from  Corres- 
pondencia. ihe  Alpine  flora  is  well  represented  on  these  hills.  Near  the  top  of 
the  high  western  barranca  some  shallow  depressions  almost  lack  vegetation  (fig.  81). 
The  soil  is  cla\-ey  and  the  surface,  which  shows  distinct  signs  of  water  erosion, 
strewn  with  stones  and  cracked  in  places.  Some  small  boulders  rested  on  short 
jMllars  of  cla>'.  The  ground  is  perhaps  flooded  during  the  winter.  I  was  told  that 
{)atches  of  snow  have  been  observed  here,  but  this  needs  corroboration.  Wind 
erosion   might  be  res[)onsible  for  the  formation  of  the  small  "tables". 

Ai'ciiida  dc  las  Cahras,  "Goat  Avenue",  is  a  well-beaten  track  running  along 
the  very  edge  of  the  abyss  from  C.  Atravesado  to  Co  Verde.  From  about 
1100-125001  alt.  one  looks  down  on  Buque  Varado  (fig.  82)  and  Q.  del  hndn 
(Magnet  gorge,  fig.  83).  One  of  the  former  inmates  of  the  convict  settlement  told 
me  that  he  had  been  em{)loyed  cutting  a  trail  down  the  precipice  to  Buque 
Varado,    where    also    huts  were  built,  but  we  saw  no  signs  of  either.  There  are 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN   FERNANDEZ  ISLANDS 


163 


] 


Fig.  79.    View    across    Piano  de  la  Mona  toward  the  Larga  and  Sanchez  valleys,  elev.  940  m. 

—  Photo  C.  Skottsberg  25/2   ^9^7- 


Fig.   80.    Looking    down    into    one    of  the  upper  gorges  of  Quebr.  Loberia  from  an  altitude  of 
c.   1400  m.  —  Photo  C.  Skottsberg  "/a  ^9^7- 


i64 


C.   SKO'lTSHERG 


Fi.l,^  8i.    Small    depression    at    the    west    edge    of  the    Masafuera  table-land,  c.   1400  m;  naked 
soil,  on  the  stones  mosses  and  lichens.  Left,  a  patch  oS.  Lophosoria.  —  Photo  C.  Skottsberg  ^^/a  191 7- 


Lookins 


down  on   I'laya  del    liuque  Varado  from  c.   1 200  m. 
^3   I9I7. 


Photo  C.  Skottsberg 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


165 


Fig.    83.   Quebr.  del  Iman  seen  from  c.   iioo  m.  Left,  Buque  Varado.  —  Photo  C.   Skottsberg 

73  1917- 


Fig    84    Piano  de  la  Mona,  looking  N,  with  Cerro  Verde  in  the  background.  —  Photo  C.  Skotts- 
berg 73  191 7. 


66 


C.   SKOTTSHERG 


Fig.  85.    Looking   S  tf)\var(l   Loberia  Nueva  from  Avenida  de  las  Cabras  N  of  Buque  Varado. 

—   Photo  C.  Skottsberg  '/a   1917. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


167 


I  68  C.   SKOTTSBKRG 

only  scattered  patches  of  trees  on  this  side,  while  ferns,  especially  Lophosoria, 
pant^ue,  <^rass  etc.  form  a  more  or  less  continuous  cover.  A  picture  taken  from  a 
point  between  Buque  \'arado  and  Iman  gives  some  idea  of  the  scenery  (fig.  85). 
At  the  north  extremity  of  the  plain  stands  a  green,  rounded  hill,  Cerro  Verde 
(fig.  84),  mentioned  as  Pico  \orte  in  Instrucc.  naut.  p.  230  and  said  to  be  1 340  m 
high.   This  figure  is  too  high,   I  believe. 

From  Cabo  Norte  to  0-  de  las  Casas. 

The  coast  cliffs  from  Cabo  Xorte  to  the  Sanchez  shore  are  almost  perpen- 
dicular and  about  300  m  high.  Half  way  a  cascade  tumbles  down  to  the  beach 
in  two  lea[)s,  coming  from  the  gap  of  O.  Larga.  Just  as  the  other  valleys  on  the 
east  side  this  has  a  forest  belt  above  c.  300  m  and  grass-land  below.  Here  the 
trees  cover  also  part  of  the  ridge  between  Larga  and  Sanchez,  apparently  the 
largest  continuous  patch  of  luma  forest  on  Masafuera. 

0.  lie  Scvichez  is  very  long,  narrow  and  twice  furcate,  and  there  is  forest 
in  all  the  branches.  None  of  the  other  valleys  is  quite  destitute  of  trees.  The 
small  0.  Xegra  (also  called  O.  del  Plan  Negro)  does  not  reach  the  shore.  Q.  del 
Saudalito  and  O.  del  Sdndalo  do  not  seem  to  offer  anything  of  particular  interest. 
The  talus  slope  permits  a  fairly  comfortable  passage  as  far  south  as  to  the  large 
O.  del  Pasto.  "Pasto"  means  pasture,  and  there  is  better  grass-land  with  abun- 
dant native  grasses  along  this  part  of  the  coast  than  anywhere  else.  Pasto  is  a 
large  valley  with  its  sources  near  the  edge  of  table-land.  The  main  stream  runs 
over  a  high  threshold.  S  of  Pasto  a  land-slide  is  crossed  before  reaching  Q.  del 
0/mlo,  which  derives  its  name  from  a  conspicuous  hill  inside  the  entrance,  a  good 
exam[)le  of  columnar  structure.  The  passage  is  a  little  fatiguing  until  we  are  op- 
posite O.  del  Mono,  another  big  valley  emptying  its  waters  through  a  narrow 
crevice  (fig.  86).  From  here  going  is  easy  to  Casas,  where  we  have  completed 
our  circuit  of  the  island. 

Human  influence  on  Juan  Fernandez. 

The  discovery  of  the  two  virgin  and  uninhabited  islands  in  1574  was  soon 
followed  by  the  first  inroad  into  its  living  world.  If  we  are  to  believe  the  histo- 
rians cited  e.g.  by  Guzman,  I^Y'rnandez  returned  and  settled  on  Masatierra  where  he 
introduced  60  Indians  (Weber),  a  few  goats  and  pigs  and  devoted  himself  to 
agriculture,  fishing  and  sealing.  His  stay  is  said  to  have  been  short,  but  his  goats 
remained  and  multiplied.  After  P\M-nandez'  death  the  island  was  turned  over  to 
the  ( )r(lcr  of  Jesuits,  and,  in  the  beginning  of  the  17th  century,  the  first  fruit- 
trees  were  planted  and  vegetables  introduced  (Weber  p.  162).  During  the  "era 
ot  the  buccaneers"  the  islands  served  as  a  place  of  refreshment,  but  the  damage 
was,  I  (laresa\-,  confined  to  the  surroundings  of  the  harbours.  Their  first  visit  is 
said  to  lia\e  taken  [)lace  in  1680.  A  change  for  the  worse  came  when  L'Heremite 
reported  that  the  precious  sandal-wood  was  abundant.  I  refer  to  Johow's  instructive 
account  of  the  history  of  this  ill-fated  tree,  pp.  127 — 131.  L'Heremite's  visit  took 
place   in    1624.  and   already  at  the  middle  of  the  century  a  lucrative  traffic  was 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


69 


Fig.  87.    Ruins  of  the  old  Spanish  fort  in  Cumberland  Ray.  —  Photo  C.  Skottsberg  28/4  191 7- 


in  full  swing  between  the  islands  and  Perii.  Toward  the  middle  of  the  i8th  century 
there  cannot  have  been  very  much  left  on  either  island,  or  the  sandal-wood  should 
not  have  escaped  Anson  and  his  men  who  spent  months  there  and  must  have 
known  of  the  former  trade. 

The  hunt  for  sandal-wood  ended  in  the  extermination  of  this  species  and 
had,  I  presume,  serious  effects  on  the  forest  in  general,  with  the  final  result  that 
all  the  lower  slopes  became  cleared  of  trees,  leaving  the  field  open  for  an  in- 
vasion of  weeds;  Ulloa  for  instance  was  struck  by  the  abundance  of  Avena. 

Having  suffered  serious  losses  from  the  raids  of  the  martial  English  navi- 
gators the  Spaniards  took  possession  in  1750  and  made  up  their  mind  to  trans- 
form Masatierra  into  a  stronghold.  I  shall  not  dwell  upon  the  history,  enough  said 
that  in  1779  7  places  were  fortified.  Part  of  the  walls  in  Cumberland  Bay  are  still 
standing  (fig.  87),  and  old  guns  lie  about  in  other  places  (fig.  33).  Nor  have  I  any 
good  reason  to  retale  the  melancholy  history  of  the  time  when  Masatierra  served 
as  a  prison  for  banished  patriots  during  the  years  of  resurrection  and  later.  Its  role 
as  a  depository  for  undesirable  citizens  came  to  a  definite  close  in  1855.  Then 
came  the  tenants. 

A  source  of  destruction  of  the  forest,  not  at  all  negligible,  were  the  fre- 
quent visits  of  American  whalers  during  the  19th  century  who  called  to  supply 
themselves  with  water,  wood  and  goats'  meat.  The  large  herds  of  goats  had  been 
II  ""537351   T^h^  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.  Vol.  I. 


lyo  C.   SKOTTSBERG 

a  i^rcat  asset  durin*^  the  era  of  the  corsairs,  and  in  order  to  stop  this  traffic 
mastiffs  were  landed  as  early  as  in  1686  or  Sy  (Hurney  IV  p.  210).  The  result 
was  tiiat  the  t^oats  had  become  very  shy  at  the  time  of  Anson's  visit;  he  estim- 
ated them  to  be  about  200  only,  but  the  dogs  were  plentiful,  savage  and  dan- 
gerous. I  lis  men  saw  no  dogs  on  Masafuera,  whereas  goats  abounded,  but  later 
dogs  were  introduced  also  to  this  island  which  was  spoken  of  as  "Isla  de  Perros". 
There  were  some  dogs  left  when  Johow  went  to  Masafuera  in  1892,  but  none 
in  1908.  In  1830  they  were  exterminated  on  Masatierra.  Freed  from  their  worst 
enemies,  the  goats  increased  in  number;  in  1877  Viel  estimated  them  at  3000. 
The  new  lessee  von  Rodt  persecuted  them  for  the  sake  of  their  hide,  and  in 
1892  the  number  had  sunk  to  about  looo,  while  there  were  200  on  Santa  Clara 
and  4000  on  Masafuera;  if  these  figures  are  reliable  is  hard  to  tell.  We  came 
across  a  small  herd  on  the  south  side  of  Masatierra  in  1917,  and  once  in  a  while 
a  daring  hunter  brought  down  a  buck  from  the  inaccessible  crags  where  they  liked 
to  hide  themselves.  At  that  time  they  enjoyed  the  protection  of  the  Government  — 
the  descendants  from  Robinson  Crusoe's  goats  were  regarded  as  sacred.  They 
were  abundant  on  Masafuera,  and  even  if  their  number  was  reduced  during  the 
time  of  the  convict  station,  they  soon  recovered.  To  judge  from  what  Weber  tells 
us  the  island  swarmed  with  goats  after  the  departure  of  the  convicts  in  1930, 
and  they  went  right  down  to  the  beach.  Their  damage  to  the  indigenous  vegeta- 
tion (and  thereby  to  the  fauna)  cannot  be  estimated  nor  disputed.  Their  ravages 
had  left  their  marks  everywhere,  and  several  peculiar  endemic  plant  species 
were  on  the  verge  of  extinction   in    1917. 

Beside  goats  Juan  Fernandez  brought  pigs  which  ran  wild;  but  they  were 
killed  off  long  ago,  nor  are  there  any  wild  asses  left,  while  cattle  and  horses 
came  to  stay.  I  suppose  that  ever  since  the  first  Spanish  colony  200  years  ago 
domestic  animals  have  existed  on  Masatierra.  In  1813  there  were  1000  sheep,  100 
goats,  500  cows,  200  horses,  10  mules  and  40  pigs  (Guzman  p.  178),  but  a  decade 
later  very  few  seem  to  have  been  left.  Again  the  Government  stocked  the  island 
(Sutcliffe  2  p.  206),  and  in  1833  there  were  350  sheep,  120  tame  goats,  70  cows, 
15  horses,  8  mules  and  30  pigs  (Guzman  I.e.),  but  the  tenants  took  no  interest 
in  farming,  and  in  i860  Mackenna  registered  15  sheep,  98  cows,  15  horses  and 
15   mules  only.   In    1878  and    1892,  only  horses  remained. 

It  is  easy  to  understand  that  the  early  visitors  got  a  very  favourable  impres- 
sion of  Masatierra,  its  agreeable  climate,  volcanic  soil,  evergreen  forests,  luxuriant 
verdure  and  good  water,  and  concluded  that  the  island  was  fertile  if  only  its 
natural  resources  were  utilized.  Certainly  they  did  not  fail  to  observe  that  there 
was  very  little  level  and  moderately  sloping  land,  but  the  valleys  of  Cumberland 
Hay  and  Pto  Ingles  must  have  looked  inviting.  We  can  also  understand  that  in 
those  times  it  did  not  occur  to  anybody  that  the  living  world  was  unique,  but 
It  is  strange  that  educated  peoi)le  like  Governor  Sutcliffe  or  Mrs.  Graiiam  could 
dream  of  Masatierra  as  fit  not  only  to  (ecd  a  considerable  population  but  to  be 
able  to  export  its  products.  Mrs.  Graham,  cited  by  Sutcliffe  (2),  wrote  in  1824: 
"The  island  might  maintain  easily  2000  persons,  exchanging  the  surplus  of  beef, 
wines,  brandy,  for  bread  and  clothing;   and  its  wood  and  water  would  render  it 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  1  7  I 

invaluable  as  a  port."  And  Sutcliffe  (i  pp.  442—443)  cites  a  letter  written  to  him 
by  one  MANUEL  DE  Salas,  Sept.  i,  1835,  when  Sutcliffe  had  been  appointed 
governor: 

"The  clays  and  soils  of  various  colours  brought  to  my  memory  the  minerals  of 
Golconda  and  Brazil  .  .  .  their  (the  islands')  geographical  position  places  them  in  a  like 
situation  and  renders  probable  the  same  effects  and  equal  productions  .  .  .  One  occurs 
to  me  which  might  be  of  great  importance;  such  is  the  making  of  bricks,  of  which 
immense  quantities  are  brought  from  England  and  the  United  States,  and  for  which 
you  have  all  the  elements  in  abundance;  there  are,  the  firewood,  the  clays,  and,  above 
all,  the  idle  hands." 

Still  the  tenants,  who  succeeded  each  other,  did  not  coin  money.  Guzman 
made  an  estimation  of  the  arable  land,  iio  hectares  on  level  ground,  70  on  the 
lower  slopes;  in  addition  there  were  20  on  Santa  Clara  and  100  on  Masafuera, 
and,  if  the  forest  was  cleared,  grazing  land  of  much  greater  size  would  be  obtained. 
The  colonists  took  no  interest  in  cattle.  Lopez  argued  (1878)  that  if  the  tenant 
were  more  enterprising,  he  would  go  in  for  cattle  and  sheep,  of  which  the  island 
could  maintain  loooo  —  the  expectations  had  grown!  Viel's  report  (pp.  19 — 22), 
founded  on  his  observations  in  1877  —  a  few  days  I  presume  —  reflects  the 
ideas  of  his  time  regarding  the  future  of  Masatierra.  He  admits  that  its  fertility 
was  more  apparent  than  real  although  chonta  and  sandal-wood  grew  in  the  for- 
ests, both  highly  appreciated  for  their  valuable  wood  and  a  possible  source  of  a 
small  trade  —  it  was  50  years  since  anybody  saw  a  sandal  tree!  A  road  around 
the  island  would  become  a  great  asset,  because  various  industries  could  be  started 
in  different  places  .  .  .  Timber  was  plentiful,  but  the  contract  forbade  the  lessee 
to  use  it,  because  the  disappearance  of  the  forest  would  endanger  the  water 
supply.  Viel's  personal  opinion  was  that  this  danger  did  not  exist,  "the  island, 
situated  in  mid-ocean,  accumulates  the  clouds  and  condensation  produces  the 
rains  which  feed  the  streams  and  irrigate  the  land;  thus  there  is  no  reason  why 
the  lack  of  trees  would  have  a  contrary  effect".^  He  forgot  all  about  erosion.  He 
recommended  to  His  Excellency  to  authorize  the  lessee  to  utilize  the  forest;  be- 
sides, he  wrote,  "if  dead  trees  are  left  lying,  the  fresh  growth  will  be  poor  and 
raquitic  (!)";  one  cannot  help  wondering  why,  after  thousands  of  years,  when  the 
fallen  logs  were  left  to  decay,  there  was  still  dense,  healthy  forest.  Unfortunately, 
"the  quality  of  the  native  timber  was  not  good,  for  which  reason  it  should  be  better 
gradually  to  replace  the  indigenous  trees  by  better  ones,  which  later  would  become 
a  source  of  income  to  the  Republic.  Pines  should  be  planted  at  once  on  a  large 
scale;  it  would  be  wise  to  take  advantage  of  the  present  tenant  who  would  be 
glad  to  look  after  the  plantations,  provided  that  some  minor  behalves  which 
wouldn't  in  any  way  harm  the  State  or  render  his  contract  more  favourable  were 
conceded  him."^  It  was  necessary  to  supply  good  timber;  if  not,  the  whalers  would 
cease  to  come.  Fruit-growing  ought  to  be  improved,  wheat  grew  well,  potatoes 
could  be  exported  to  northern  Chile  with  great  gain.  In  this  connection  Viel  says 
that  zarzaparilla  —  here  =  Acaejta  argentea,  a  noxious  weed  —  was  common  and 


1  Translated  from  Spanish. 


72 


C.   SKOTTSBERG 


maqui  not  less  abundant;  it  is  not  clear  whether  or  not  he  regarded  them  as  a 
source  of  wealth. 

As  a  more  promising  industry  than  forestry  Vie!  recommends  raising  cattle. 
The  island  could  easily  maintain  looo,  during  the  Spanish  time  there  had  been 
800,  and  now  the  number  of  domesticated  animals  was  98  cows  and  50  horses, 
but  many  had  run  wild.  The  3000  wild  goats  ought  to  be  protected  by  a  closed 
season;  800  skins  had  been  exported  recently.  Apparently  the  fur  seals  did  not, 
in  his  opinion,  need  })rotection:  "the  output  was  mediocre,  a  consequence  of  the 
settlers  not  being  sufficiently  intelligent  and  industrious"^  —  he  forgot  that  the 
j)oor  result  of  sealing  was  a  consequence  of  ruthless  persecution  during  a  cent- 
ury. The  average  number  of  seals  killed  per  annum  was  700  —  one  would  call 
this  a  fair  number  considering  the  small  size  of  the  islands.  Fishing  was  neg- 
lected, onl\'  very  little  salted  or  dried  fish  had  been  exported.  There  was  plenty 
"langosta",  and  it  ought  to  be  made  into  preserves.  Viel  concludes  his  report 
with  the  following  words:  "El  estado  actual  es  bien  lamentable."  Had  the  Govern- 
ment listened  to  his  recommendations  the  situation  would  have  become  much 
more  lamentable. 

Of  this  official  report  to  the  Minister  of  Finance  and  to  the  Governor  of 
\'alparaiso  Fkllexberg  published  a  translation,  but  he  protests  against  Viel's  belief 
that  the  disappearance  of  the  forest  would  not  harm  the  water  supply.  As  an  appen- 
dix he  published  3  letters  from  the  new  tenant,  addressed  to  his  relatives  in  Bern. 

Ai.KRKi)  vox  Root,  of  Swiss  descent,  an  ex-officer  in  the  Austrian  army 
and  a  well-educated  gentleman,  settled  on  contract  on  Masatierra  as  tenant  of 
the  islands,  h'.ncouraged  by  the  many  favorable  reports  on  their  resources,  he  ex- 
pected to  make  his  fortune,  and  he  had  some  money  to  invest  in  the  enterprise. 
His  first  letter,  written  shortly  after  his  arrival  and  dated  June  5,  1877,  contains 
dates  on  the  size  and  |)osition  of  the  place;  he  tells  that  there  were  7000  wild 
goats  and  [)asture  for  1000  head  of  cattle  and  that  he  intended  to  start  "verschie- 
dene  Industrien  '.  I'A'idently  he  went  to  work  without  delay;  on  March  13,  1878, 
he  writes  that  he  had  timber  worth  3000  Dollars  ready  to  be  shipped  to  the  coast. 
He  reported  from  Masafuera  17.6.  1878  that  this  island  was  considerably  larger 
than  Masatierra  (!)  and  that  there  were  large  forests  and  the  most  beautiful  grass- 
lands where  it  would  be  easy  to  feed  20000  sheep.  He  had  killed  more  than 
700  fur  seals. 

I  he  castle  in  the  air  vanished,  von  Rodt  never  founded  any  new  industries 
on  Masatierra,  there  never  was  a  sheep-farm  on  Masafuera.  He  lost  his  money, 
but  he  remained  true  to  his  beloved  Masatierra,  where  it  is  easy  enough  to  make 
a  living,  but  perhaps  not  a  fortune.  The  von  Rodt  dynasty  was  still  going  strong 
when   we  \  isitcd  the  islands;  the  sons  of  Alfred  made  their  living  as  lobster ^  fish- 

^  'I'ranslated  from   .Spanish. 

2  i'rofrssor  Kari.  I..\N(;,  head  of  the  Dept.  of  Evcrtebratcs  in  the  Nat.  Hist.  Museum,  Stock- 
hohii,  on  my  ro(|ucst  kindly  made  a  dihgent  search  for  an  English  equivalent  to  the  Spanish 
word  langosta  as  name  of  the  large  Decapod  Jasus  Lalatidci  (formerly  known  as  Palinurtis 
ln>n/a/is  ,  which  from  a  scientific  viewpoint  is  no  real  lobster.  He  reports  that  Palinurus  vul- 
i^uiris  and  related  forms  are  called  "rock  lobster";  "spiny  lobster"  is  another  name.  Yox  the  sake 
of  brevity  "lobster"  or     langosta"  will  be  used  here. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  I  73 

ers,  cultivated  their  corn-fields  and  gardens  and  reared  a  new  crop  of  sturdy 
fishermen. 

The  damaging  influence  of  von  Rodt's  reign  on  the  native  flora  and  fauna 
must  not  be  overrated.  It  is  true  that  the  maqui  spread,  but  also  that  the  goats 
decreased  in  number.  The  population  remained  very  small,  and  the  cultivated 
soil  did  not  extend  much.  In  comparison  with  recent  disasters  the  encroach  during 
the  last  decades  of  the  19th  century  was  of  little  significance,  except,  perhaps,  with 
regard  to  the  chonta  palm,  one  of  Nature's  great  treasures.  And  BuRGER  tells 
us  (p.  Ill)  that  when  von  Rodt  failed  to  "develop"  the  islands,  he  became  a 
protector  of  their  nature. 

In  1877  the  population  of  Masatierra  counted  64  persons,  29  of  whom  were 
male,  children  not  included  (demographic  figures  taken  from  Guzman  p.  37),  in 
1878  74  (22),  in  1879  141  (51),  and  in  1880  147  (55)  —  the  rise  probably  due  to  the 
arrival  of  labourers  contracted  by  the  tenant.  Then  it  began  to  sink:  82  (24)  in 
1885,  61  (23)  in  1886.  In  1885  the  German  merchant  Alexander  Ermel  visited 
Masatierra  as  partaker  in  a  tourist  excursion.  As  many  before  and  after  him  he 
fell  for  the  beauty  of  the  scenery  and  he  returned  with  fantastic  ideas  of  the  value 
of  the  natural  resources,  which  nobody  had  understood  to  utilize  properly.^  Con- 
sequently he  did  not  hold  a  very  high  opinion  of  von  Rodt.  Even  barren  Santa 
Clara  became  fertile  in  Ermel's  eyes.  For  Masafuera  only  no  great  future  was  in 
store;  it  was  too  inaccessible.  "Daher  wird  Masafuera  fiir  die  Zukunft  allein  die 
traurige  Bestimmung  haben,  als  ein  in  Wolken  eingehiillter,  schauriger  Felsen 
dazustehen,  dessen  Nahe  die  Seefahrer  soviel  als  moglich  vermeiden  werden" 
(p.    115). 

In  spite  of  the  very  short  duration  of  his  visit  (3  days),  Ermel  judged  him- 
self competent  to  discuss  the  economic  importance  of  the  islands  for  Chile.  Masa- 
tierra might  well  serve  as  a  health-resort.  Its  principal  industry  would  be  fishing. 
Here  he  was,  as  the  future  showed,  right,  and  the  same  is  true  when  he  argues  that 
the  seals  needed  protection.  In  former  days  the  seals  had  been  extremely  abund- 
ant, not  only  the  fur  seal,  but  also  the  sea  elephant.  L'Heremite  saw  thousands 
of  sea  lions  and  seals  (Burney  III  p.  18).  Walter  reports  on  the  "sea  lions",  which 
occurred  in  large  herds,  but  PI.  XIX  shows  that  his  "sea  lions"  were  sea  elephants. 
UUoa  tells  us  (p.  287)  that  "the  beaches  and  rocks  were  everywhere  crowded 
with  seals  in  such  abundance  that  no  free  space  was  left  where  one  could  walk; 
they  did  not  leave  a  passage  between  them".^  Captain  ROGERS  (1709)  says 
that  a  compact  string  of  seals  lined  the  beach  of  Cumberland  Bay  (Guzman 
p.  215).  The  main  slaughter  of  the  fur  seal  commenced  in  1797;  it  is  reported  that 
in   1801   a  single  ship  carried  one  million  skins  to  the  London  market.  If  this  be 


1  These  illusions  have  been  very  tough.  In  an  American  newspaper  (Meridian  Star,  Miss.) 
the  following  paragraph  appeared  on  March  i,  1928:  "Crusoe  island  fruitful. — Juan  Fernandez 
Island,  on  which  Alexander  Selkirk,  the  reputed  original  of  Robinson  Crusoe,  lived  for  four 
years,  is  one  of  the  most  fruitful  spots  in  South  America,  according  to  a  recent  survey.  Every 
known  plant  seems  to  grow  there.  One  Frenchman  who  was  shipwrecked  there  40  years  ago 
refuses  to  leave."  No  comments  needed,  but  the  climax  is  priceless. 

2  Translated  from  Spanish.  ... 


174 


C.   SKOTTSHERG 


true,  it  is  a  wonder  that  sealing  could  be  practised  with  a  profit  over  a  period  of 
40  years.  The  seals  have  disappeared  from  Masatierra,  the  sea  elephant  is  extinct 
in  the  waters  of  juan  Fernandez.  As  I  told  above,  we  found  a  small  herd  of 
fur  seals  at  Loberia  Vieja.  Since  1891  it  is  protected  by  law,  to  what  effect  I  shall 
not  say. 

Hut  let  us  return  to  l^rmel.  l^ven  if  fishing  ranked  first,  the  wealth  of  the 
land  was  b\'  no  means  contemptible:  timber  and  fuel,  chonta,  sandalo,  charcoal, 
"womit  Herr  von  Rodt  in  den  ersten  Jahren  seiner  Pachtzeit  einen  lohnenden 
Anfang  gemacht  hatte";  all  kinds  of  fruit,  probably  also  grain  and  wine,  and 
breeding  cattle,  llamas  and  vicunas  should  be  tried.  He  also  underlined  Masatierra's 
strategic  {position. 

However,  nothing  happened  except  that  the  population  continued  to  dwindle, 
reaching  its  lowest  figure,  35  (12),  in  1893.  When  Johow  revisited  the  islands  in 
1895  it  had  grown  to  54.  In  1891 — 92  Johow  headed  an  official  commission  and 
he  outlined  a  program  for  the  future  management  of  the  islands.  This  document 
was  published  in  his  book,  pp.  267 — 274.  It  is  not  without  interest,  and  a  sum- 
mary will  be  given  here.  It  begins  by  stating  that  the  utilization  must  be  based 
on  the  principal  marine  products,  the  langosta  (jfasus  Lalandei\  confined  to 
Juan  I^'ernandez  and  the  Desventuradas  Islands  (San  Ambrosio  and  San  Felix),  and 
the  bacalao  {Polyprion  prog7tatiis),  also  absent  from  the  coast  of  the  mainland.  Even 
the  latter  would  become  a  valuable  article  of  export,  the  more  so  because  it  is 
one  of  the  w^orst  enemies  of  Jastis.  The  waters  abound  in  many  other  kinds  of 
savoury  fishes.  When,  however,  the  commission  recommends  to  repeal  the  law  of 
1 89 1,  prohibiting  the  destruction  of  the  fur  seals,  with  the  motive  that  seals  are 
the  most  dreaded  enemies  of  the  fishes,  no  responsible  authority  in  our  days 
would    agree.    Poachers    can    be  relied  upon  to  keep  the  number  of  seals  down. 

Johow  states  that  agriculture  will  never  become  profitable,  for  the  simple 
reason  that  there  is  very  little  arable  land  and  that  the  soil,  once  deprived  of 
its  natural  vegetation  cover,  will  be  subject  to  erosion.  Wheat  should  be  imported 
from  the  mainland  and  land  utilization  limited  to  cultivation  of  potatoes  and  veget- 
ables for  local  consumption.  Nothing  is  said  of  the  aspects  as  grazing  land. 
There  were  very  few  animals  in  Johow's  time.  After  these  introductory  remarks  the 
commission   proceeds  to  answer  a  number  of  questions. 

With  regard  to  the  administration,  some  kind  of  authority  should  be  estab- 
lished on  Masafuera.  A  small  steamer  would  become  necessary  to  serve  the  com- 
munications between  Masatierra  and  Valparaiso.  An  "inspector  de  colonizacion" 
should  be  appointed.  The  question  whether  it  would  be  necessary  to  prohibit 
the  cutting  of  chonta  and  sandalo  is  answered  in  the  affirmative.  Johow  estimated 
the  number  of  full-grown  chontas  at  100  to  150,  and  he  had  occasion  to  visit 
the  last  living  sandalo.  They  were  already  under  provisional  protection  and  it 
was  recommended  "no  solo  por  motivos  meramente  ideales  o  sea  cientificos,  sino 
tambien  por  consideraciones  de  caracter  comercial  i  economico"  that  the  law 
should  remain  in  force  and  violation  be  subject  to  severe  penalty:  "No  seria,  a 
nuestro  juicio,  exajerado  castigar  la  destruccion  de  un  solo  ejemplar  de  chonta 
o  de  sandalo  como  delito  de  robo  comun,  i   prohibir  tambien  su  corta  bajo  pre- 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  I  7  5 

testo  cientifico."  And  the  commission  made  public  that,  at  repeated  occasions, 
the  crew  —  they  forgot  to  add  "and  officers"  —  of  vessels  belonging  to  the  Armada 
Nacional  had  possessed  themselves  of  large  quantities  of  chonta  trunks  —  with 
the  assistance  of  the  colonists.  (Such  is  the  attitude  of  the  majority  of  people 
all  over  the  world  that  if  an  individual  stands  before  the  last  living  specimen  of 
a  plant  or  animal,  he  will  grab  it,  because  if  he  doesn't,  someone  else  will  have  it.) 

In  the  1890's  there  were  no  plantations  needing  protection,  but  in  Johow's 
opinion  the  existence  of  the  native  forest  was  jeopardized  by  the  ravages  of  a 
parasitic  fungus,  Antennaria  scoriadea  Berk.  (Limacinia  fernandeziana  Neger  ex 
Johow),  and  he  feared  that  it  would  "concluir  poco  a  poco  con  la  vejetacion  de 
muchas  quebradas  ".  The  fungus  is  still  plentiful,  but  its  killing  capacity  was,  as 
far  as  I  could  see,  greatly  overrated.  The  commission  recommended  to  make  pun- 
ishable to  set  the  woods  on  fire,  and  to  enjoin  the  inhabitants  and  visitors  in 
need  of  wood  to  use  only  the  fallen  trunks.  If  I  remember  right  there  later  was 
a  regulation  that  only  the  maqui  could  be  used  for  fuel.  In  the  report  nothing 
is  said  about  the  damnable  habit  of  visitors  to  rob  the  forest  of  the  stately  en- 
demic tree-ferns  which,  always  without  success,  they  tried  to  plant  in  their  gardens 
on  the  mainland.  Ermel  saw  the  lifeboats  of  his  ship  return  laden  with  young 
chontas  and  tree-ferns  and  looking  like  gardens  afloat.  I  have  witnessed  the  same 
traffic  myself,  but  I  do  not  know  if  it  still  flourishes  —  there  is  a  long  way  now 
to  the  young  chontas  and  to  the  tree-ferns. 

A  question  if  fishing  with  dynamite  ought  to  be  forbidden  was  answered  in 
the  affirmative,  but  not  because  it  would  reduce  the  fish  supply  very  much  — 
though  it  is  admitted  that  the  demand  for  fish  might  be  greater  in  the  future  — 
but  rather  because  "la  dinamita  .  .  .  manejada  por  personas  de  poca  o  ninguna 
ilustracion,  facilmente  orijina  funestos  incidentes  .  .  .".  With  regard  to  the  langosta 
no  scarcity  had  made  itself  felt,  but  it  was  recommended  to  leave  it  in  peace 
during  the  breeding  season. 

Among  the  remaining  questions,  of  which  many  were  referred  to  specialists 
and  not  answered  by  the  commission,  one  is  of  interest:  it  was  recommended  to 
exterminate  the  wild  dogs  and  the  native  buzzards  on  Masafuera.  The  motive  was 
to  take  better  care  of  the  goats,  still  considered  to  be  an  important  source  of 
meat;  the  meat  is,  I  admit,  very  palatable.  A  bounty  was  therefore  suggested. 
Nobody  will  grieve  the  loss  of  the  wild  dogs;  they  are  not  good  company,  but 
in  the  case  of  the  buzzard,  Buteo  erythrojioius  exsul,  the  care  for  the  goats  gets 
in  conflict  with  the  interests  of  science.  In  his  catalogue  of  the  insular  Ornis  Johow 
remarks  that  the  "aguiluchos  quizas  representan  una  variedad  endemica  si  no  una 
especie  distinta  de  la  del  continente"  —  nevertheless  he  would  not  hesitate  to 
exterminate  it.  Had  he  known  the  flora  of  Masafuera  better,  he  would  have  been 
less  kindly  disposed  toward  the  goats. 

Finally  the  commission  underlines  the  urgent  need  of  regular  communications 
with  the  mainland  with  rapid  steamers  permitting  the  products  of  the  fishing  in- 
dustry to  arrive  fresh  at  their  destination. 

Thus  the  wants  and  needs  were  stated,  but  little  was  accomplished.  The 
wild  dogs  on  Masafuera  were  exterminated,  but  fortunately  the  buzzard  escaped. 


76 


C.   SKOTTSllERC 


h  ig.  88.  The  boat  harbour  in  Cumberland  Bay.  —  Photo  C.  Skottsberg  ^/jg   1916. 


Fig.  89.  Lobster  fishers  working  for  Recart  y  Doniez  by  their  boats  during  the  closed  season. 
Cumberland  Bay.  —  Photo  K.  Backstrom  Dec.   19 16. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


77 


Fig.  90.   The   take   is   brought   into  the  Recart  y  Doniez  factory;  left,  Hve  lobsters;  right,  the 
metal  cooking  baskets.  —  Photo  C.  Skottsberg  »/i  191 7. 


P 


Fig.  91,  The  harbour  in  Cumberland  Bay,  showing  three  schooners  at  anchor,  a  motor  launch 
and    a    number    of  fish-chests    used    for   storing   the  take  until  shipped  alive  to  Valparaiso,  — 

Photo  C.  Skottsberg  3/i   1917. 

12  —537351   The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.  Vol.  I. 


C.   SKOTTSBERG 


P'ig.  92.  The  village  in  Cumberland  Bay,  seen  from  the  slope  of  Salsipuedes. 

'berg  21/^   1 91 7. 


Photo  C.  Skotts- 


Masafuera  got  no  supervisor,  the  management  of  the  forests  continued  as  before, 
the  clionta  continued  to  be  cut  unlawfully  —  a  common  occupation  during  the 
winter  months.  Weber  says  p.  116  —  nothing  was  done  for  the  fishing  industry, 
no  steamer  came  and  went  to  the  benefit  of  the  inhabitants.  Life  went  on  as  before. 
Let  us  return  to  the  langosta.  Ever  since  the  days  of  the  discovery  of  the 
islands  this  magnificent  crayfish  had  been  appreciated  as  very  good  eating,  and  at 
Anson's  time  the  beaches  literally  swarmed  with  large-sized  specimens.  No  in- 
strument was  needed  to  catch  any  amount  in  one  or  two  feet  of  water.  How 
long  this  happy  state  of  affairs  lasted  I  do  not  know,  nowadays  the  best  catch 
is  made  in  depths  from  40  to  80  meters.  In  spite  of  the  value  of  this  marine 
product,  which  from  time  to  time  was  shipped  to  the  mainland,  none  of  the 
tenants  had  the  means  to  organize  the  industry  on  a  larger  scale.  A  company 
was  formed  in  1893,  but  even  with  a  catch  of  35  to  40  thousand  annually  and 
exporting  some  40000  tins  of  preserved  tails,  the  business  did  not  pay,  and  a 
new  company  formed  in  1900  also  failed  (see  Guzman  p.  209).  In  19 14  Messrs. 
Recart  y  Doniez  started  their  establishment.  We  had  the  very  best  opportunity 
to  watch  the  fishing  from  the  catch  of  the  sharks  used  as  bait  to  handling  the 
langosts  in  the  factory  (figs.  88—91)  and  to  the  export  of  the  living  animals,  in 
the  com()any  of  which  we  made  our  return  voyage  to  Valparaiso  onboard  one  of 
the  schooners.  The  stern   is  built  as  a  well  with  free  circulation  of  the  water,  and 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


79 


-C^  ". 


?y^. 


Fig.  93.    Carica  papaya   in    the  garden  of  Mr.  Charpentier,  Valle  Anson. 

berg  i*/4  1917- 


Photo  C.  Skotts- 


if  the  passage  could  be  completed  in  2  V2  to  3  days  all  went  well,  but  in  too 
calm  or  too  bad  weather  the  motor  alone  did  not  help  very  much,  the  crossing 
was  slow  and  it  happened  that  very  few  of  the  lobsters  arrived  alive.  Two  minor 
companies  were  also  in  operation.  The  closed  season  was  strictly  observed.  On 
the  2d  of  January  at  6  a.m.  a  rifle  shot  signalled  the  open  season,  and  the 
launches  towed  a  string  of  boats,  each  party  rushing  to  gain  the  best  grounds 
which  it  monopolized  during  the  season.  Recart  y  Doniez  also  held  the  licence 
to  fish  at  Masafuera  and  sent  a  schooner  thither  several  times.  I  don't  remember 
having  heard  that  any  kind  of  fish  was  exported  at  that  time,  but  fish  was  the 
staple  food  on  the  island.  About  200  people  lived  there,  most  of  the  men  being 
employed  by  the  companies.  The  village  in  Cumberland  Bay  (fig.  92)  looked 
quite  inviting  with  its  neat,  if  not  always  too  well  kept  houses,  vegetable  gardens, 
fruit  trees  and  flower-beds,  set  among  exotic  trees  like  araucarias,  eucalypts, 
poplars,  pines,  Albizzia,  Eriobotrya  and  so  forth.  Johow  has  published  a  paper 
on  the  plants  cultivated  in  the  islands,  and  also  a  list  in  his  book  pp.  263 — 266. 
With  the  exception  of  the  fig  and  the  quince  most  of  the  fruit  trees  yielded  in- 
different fruits.  The  cliinate  is  of  the  Mediterranean  type,  but  at  the  same  time 
pronouncedly  oceanic,  and  the  lack  of  a  period  of  real  warm  weather  seemed  to 
be  responsible  for  the  failure  of  Citrus  fruits.  The  more  surprising  was  it  to  find 
the  papaya  in  cultivation  in  a  garden  belonging  to  a  colonist  of  French  descent. 


i8o 


C.   SKOTTSBERG 


Fig.  94.    Threshing   the  wheat  with  a  tropilla  of  horses,  Cumberland  Bay.  —  Photo  K.  Back- 

strom   1917. 


m 

mmmmmemwsm 

'■■--■^m-         ™ ■— ■ -— ,^^^i*  . .  1  _^ 

■-— p--- 

^^^— jMjJfc       ,  i.  "■■./*;*»■                         1 

Fig.  95.  Ch..;;  ...A...^.   ;.,c  wind.  Same  place  as  lig.  94.  -       Tholo  K.  Backslrui;.   uji] 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


8r 


Fig.  96.  A  rodeo  in  Cumberland  Bay,  —  Photo  K.  Backstrom  191 7. 


The  specimens  were  small,  and  so  were  the  fruits,  which  we  had  no  opportunity 
to  taste  (fig.  93).  This  seems  to  be  the  only  tropical  tree  thriving  in  this  warm- 
temperate  climate.  Several  ornamental  plants  have  become  naturalized  in  the  neigh- 
bourhood of  the  village,  e.g.  Zantedeschia  aethiopica,  Pelargonium  spp.,  Tropaeoluni 
ntajus,  Matthiola  incana  and  Lochnera  rosea. 

Wheat-fields  were  few  and  small  and  harvesting  methods  quite  primitive; 
see  figs.  94  and  95.  Most  of  the  flour  was  imported,  and  there  was  also  a  shortage 
of  potatoes.  Unfortunately  the  number  of  animals  had  increased  very  much,  much 
more  than  would  be  deemed  necessary,  but  milk-cows  were  few.  Imported  wine 
was  the  daily  beverage,  here  as  in  other  parts  of  Chile.  Herds  of  cattle  roamed 
through  all  the  valleys  from  Pto  Frances  to  Vaqueria  on  the  north  and  from  the 
foot  of  Mt.  Yunque  to  Puente  on  the  south  side  of  the  island.  I  regret  having 
neglected  to  ask  their  number;  every  animal  had  its  owner.  Fig.  96  shows  a 
rodeo  in  Cumberland  Bay.  Only  the  herd  in  Vaqueria  was  left  to  run  wild.  Horses 
were  seen  in  some  valleys,  but  I  cannot  remember  having  seen  any  slieep,  and 
we  were  never  offered  mutton. 

After  19 1 7  the  population  remained  about  the  same  for  years,  but  in  1930 
it  had  increased  to  298,  of  which  155  were  male,  including  the  boys.  Ten  years 
later  it  had  grown  to  434  (225). 

I  shall  leave  Masatierra  for  the  present  and  try  to  follow  the  fate  of  Masafuera. 


C.  SKOTTSP.KRG 


J 


Fig.  97.  View  from  the  south  slope  of  Casas  valley,  Masafuera,  showing  the  headquarters  of  the 
first  convict  settlement.  —  Photo  K.  Backstrom  Feb.   1917. 

The  first  house  was  built  in  1867  (Weber).  Goat-hunters  or  fishermen  used  to  visit 
the  island.  In  1908  we  found  some  people  staying  there,  but  there  was  no  perma- 
nent settlement.  The  next  year  saw  a  radical  change.  The  full  history  is  told  by 
Guzman  pp.  87 — loi.  A  short  summary  is  given  here.  Guzman  begins  with  the 
following  words:  "Tal  vez  la  descripcion  de  la  flora  natural  de  la  isla  de  Mas  Afuera 
que  hiciera  el  sabio  sueco  Skottsberg,  hizo  creer  a  un  Ministro  de  Justicia  del 
Presidente  Pedro  Montt,  que  dicha  isla  era  el  sitio  ideal  para  la  fundacion  de  un 
presidio  agricola."  Certainly  Guzman  is  joking,  the  more  so  as  I  tried  on  my 
return  to  start  a  campaign  in  favour  of  preserving  the  native  flora  and  fauna 
—  not  the  goats  however!  —  and  never  said  a  word  about  Masafuera  as  a  suitable 
place  for  agricultural  or  any  kind  of  commercial  experiments.  Two  months  later 
a  decree  was  signed,  transforming  the  island  into  a  settlement  of  criminals,  and 
in  April  1909  "Carcel  Pedro  Montt"  received  its  first  inmates  and  soon  after  a  second 
lot,  together  170.  For  the  State  it  was  no  cheap  affair  to  erect  substantial  headquar- 
ters at  the  entrance  to  Casas  valley,  to  build  a  jetty  in  the  turbulent  water  of  the  so- 
called  landing-place  and  a  schooner  to  serve  the  colony,  but  the  parents  of  the 
scheme  were  optimistic  and  thought  that  once  in  operation,  the  establishment 
would  soon  become  self-sup[)orting,  a  miracle  to  be  performed  by  cutting  trees 
and  cultivating  the  soil.  Goats  were  plentiful  and  easy  to  get  hold  of,  at  least  to 
begin  with.  "Tal  era  la  conviccion  que  tenian  los  creadores  del  presidio  en  la  fe- 
racidad  de  la  estcril  Mas  Afuera,  que  al  alcalde  se  le  designo  con  el  singular  titulo 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


183 


Fig.  98.  The  ruined  forest  and  village  of  Las  Chozas,  Masafuera.  —  Photo  K.  Backstrom  Feb.  191 7. 

de  Director  y  ]e^e  de  Cultivos",  Guzman  writes.  This  official  was  a  thrifty  fellow, 
but  he  could  not  call  up  arable  land  where  there  was  none.  Houses  were  built  in 
many  places,  vegetables  sown  and  potatoes  planted,  catde  introduced  and  killed 
surreptitiously  by  the  convicts,  but  much  food  had  to  be  imported,  and  when  the 
boarders  obtained  permission  to  send  for  their  wives  or  relatives  to  keep  them  com- 
pany the  population  rose  to  350  persons.  Good  luck  did  not  favour  this  humanitarian 
enterprise,  the  situation  on  the  island  became,  to  say  the  least,  unpleasant,  the 
schooner  was  shipwrecked  and  lost,  and  in  191 3  Masafuera  was  abandoned.  The 
buildings  in  Casas  were  in  tolerably  good  condition  in  191 7  (fig.  97),  but  the 
wooden  huts  more  or  less  fallen  to  pieces.  Ruined  forests,  abandoned  potato 
fields  and  a  host  of  new  weeds  told  the  story  (fig.  98).  Of  domestic  animals  only 
two  horses  could  be  discovered.  We  caught  them  because  we  could  use  them. 
But,  as  Guzman  says,  "los  juristas  no  podian  aceptar  que  una  Naturaleza 
tan  prodiga  en  helechos  y  en  plantas  sub-alpinas,  no  fuera  generosa  tambien  para 
con  las  legumbres,  hortalizas  y  frutales  .  .  .  ",  and  in  1927  "Prision  Carlos  Ibanez" 
was  established,  this  time  not  reserved  for  ordinary  criminals  — in  addition  150 
political  offenders  were  exiled  to  Masafuera.  More  trees  were  cut,  new  houses 
built,  new  seeds  sown,  but  the  harvest  was  no  richer  than  before,  and  in  1930 
the  colony  was  discontinued.  Guzman  seems  to  fear  a  repetition  of  this  sad  story, 
to  judge  from  his  concluding  remark  "...  el  hombre  con  sus  disposiciones  legales 


i84 


C.   SKOTTSBERG 

Table  I. 
Lobster  catcli    1940—46.  After  Guzman. 


i<>40, 

i«(4i 

1042, 

i')43 

1044 

K'45 


Gross  weight  in  kg 


Number 

Total 

Living 

Tinned 

log  250 

11^  531 

15  533 

96998 

103  6()3 

100  590 

14415 

86175 

134  5«') 

II-:  531^ 

20  903 

91  628 

(,2  287 

64  160 

14929 

49231 

^7  959 

28  6oo2 

II  947 

16633 

^'3516 

24  220 

13643 

10557 

3^>43i 

58  120 

— 

— 

y  con  su  persistencia,  indiferente  del  pasado,  espera  el  momento  propicio  para 
construir  un  nuevo  penal,  sobre  las  ruinas  del  septimo  presidio"  (there  had  been 
5  on  Masatierra). 

(3n  January  the  i6th,  1935,  President  Arturo  Alessandri  signed  a  decree 
making  the  Juan  Fernandez  Islands  (together  with  Easter  I.)  a  National  Park,  and 
two  (ierman  residents,  Weber  and  Carlos  Bock,  both  addicted  to  the  study  of  the 
fauna  and  flora,  were  appointed  honorary  forest  inspectors.  Bock  soon  died,  and 
Weber  left  the  islands  after  some  years.  The  regulations  were  strict;  had  they  been 
followed  the  goal  for  which  I  had  fighted  so  many  years,  supported  by  Chilean  col- 
leagues, would  have  been  attained.  The  rules  did  not  infringe  upon  the  reasonable 
rights  of  the  colonists,  nor  was  the  fishing  industry  affected.  In  order  to  collect 
specimens  of  the  indigenous  plants  and  animals,  also  for  scientific  purposes,  a  license 
was  required.  No  permanent  habitations  could  be  erected  on  Masafuera  and  Santa 
Clara.  It  is  evident  that  Weber,  in  spite  of  his  earnest  intentions,  did  not  quite 
understand  what  effective  conservation  involves  or  he  would  not  have  written  the 
following  words  (p.  138):  "Die  Tierwelt  von  Juan  Fernandez  war  leider  sehr 
sparlich,  es  soUte  mehr  Leben  in  die  schweigsamen  Walder  kommen"  —  it  did 
not  occur  to  him  that  the  introduction  of  foreign  animals,  even  a  few  birds,  would 
disturb  Nature's  balance  in  a  way  never  properly  to  be  foreseen. 

Unfortunately  a  law  has  little  effect  without  adequate  means  to  enforce  it. 
Without  money  and  men  with  authority  a  conservation  program  cannot  be  realized, 
and  in  the  case  before  us  there  was  neither.  Serious  inroad  upon  the  forests 
might,  however,  be  prevented,  and  with  the  declining  lobster  industry  and  the 
non-existent  possibilities  to  expand  agriculture  there  seemed  to  be  no  danger  that 
more  people  would  settle  on  Masatierra.  Ikit  what  happened?  Guzman's  interesting 
account  gives  the  answer,  I.e.  p.  37. 

'  This  figure,  which  is  identical  with  the  figure  for  1940,  must  be  incorrect.  In  all  other  cases 
save  one  1941  the  average  weight  of  the  langosts  slightly  exceeded  I  kg,  and  it  was  very  little 
below  that  weight  in    1941. 

*  On  p.  202  r,u7.iii;'m  cjuotes  a  very  different  figure:  47238  kg  obtained  during  the  period 
Jan. — Aug.  The  balance  cannot,  1  am  sure,  have  been  consumed  locally,  because  the  people 
fish  for  the  companies,  so  that  some  other  explanation  has  to  be  found. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


85 


^^S-  99-    The   central    part  of  Masatierra,  seen  from  the  air  and  showing  the  cultivations  and 

plantations  around  Cumberland  Bay:  from  left  to  right  V.  Colonial,  V.  Anson  and  Quebr.  del 

Minero.  —  Photo  B.  Frodin  ^/^  1952. 

In    1940   the   population  amounted  to  434  persons  (225  men  and  boys),  in 

1943  77^   (198  men;  the  number  of  women  was  123,  of  children  no  less  than  450); 

1944  shows  a  drop  to  653,  a  result,  one  would  think,  of  the  heavy  drop  in  the 
number  of  lobsters  caught.  In  1948  the  population  was  about  600.  The  total 
export  of  lobster,  living  and  tinned,  is  shown  in  Table  I  (after  Guzman). 

As  we  see,  the  proportion  of  preserved  lobsters  underwent  a  rapid  decrease 
during  these  years;  the  export  of  live  animals  was  undoubtedly  more  profitable. 
The  reasons  for  the  sudden  rise  in  total  output  in  1946  are  not  known  to  me, 
nor  if  it  indicated  a  reliable  increase  or  just  a  transient  improvement;  after  1946 
no  figures  have  been  available  to  me. 

The  growth  of  the  population  between  1940  and  1943  cannot  have  had 
anything  to  do  with  the  fishing  industry,  and  the  drop  from  1943  to  1944  does 
not  run  parallel  with  the  quantity  of  lobsters;  besides,  the  effect  would  not  show 
before   1945,  when  no  census  was  taken. 

More  people  meant  more  houses,  more  gardens,  more  timber  and  fuel  used, 
etc.  The  effect  is  shown  by  comparing  figs,  24,  27  and  92  with  22,  30  and  99. 
In  191 7  the  lower  slopes  of  Q.  del  Minero  were  barren,  in  1952  there  were  plant- 
ations; Anson's  valley,  where  few  people  lived  in  191 7,  bears  sign  of  much 
activity,  and  the  aspect  of  the  settlement  in  the  Colonial  valley  has  changed  a 
great  deal  from  19 17  to  1952.  Everybody  welcomes  that  the  devastated  slopes 
near  the  sea,  where  erosion  is  a  serious  problem,  have  been  planted,  nobody 
objects    to    the   growth    of  the    population    as  long  as  fishing  provides  a  decent 


l86  C.   SKOTTSHERG 

livelihood  for  the  majority.  This,  however,  is  not  the  case.  The  time  has  come 
when  the  old  dreams  to  utilize  the  land,  fostered  by  well-meaning  patriots,  are 
coming  true.  The  statistics  tell  us  that  Masatierra  is  being  transformed  into  a 
cattle  and  sheep  farm,  a  new  and  strange  type  of  National  Park.  In  1945  the 
island  had  5000  sheep.  600  head  of  cattle,  500  tame  (.')  goats  and  300  horses, 
and  even  if  the  horses  and  cows  mostly  graze  the  open  country  with  grasses  and 
herbs,  mainly  introduced  weeds,  they  do  not  despise  the  light  forest  which,  and 
this  was  easy  to  observe  already  in  191 7,  suffers.  The  habits  of  the  sheep  and 
goats  are  different;  wild  sheep  were  observed  by  Weber  in  the  thirties  (p.  116). 
There  are  no  fences  and  nothing  to  prevent  these  animals  to  run  wild  among  the 
crests,  where  it  is  difficult  to  follow  them.  No  palatable  plant  species  is  likely  to 
escape  these  gluttons  and  in  contradistinction  to  the  naturalist,  they  need  no  license. 
It  goes  without  saying  that  they  will  take  to  the  mountains  when,  at  the  end  of 
the  drier  season,  food  becomes  scarce.  Guzman  recommends  introducing  new  kinds 
of  grasses,  better  adapted  to  the  climate,  a  measure  probably  unheard  of  in  combina- 
tion with  the  management  of  a  national  park.  It  should  not  be  forgotten  that  one 
Mr.  Orro  Rir.CKiEi.,  "el  gran  amigo  de  las  ciencias  naturales"  as  Guzman  calls 
him,  celebrated  the  creation  of  the  national  park  by  introducing  6  pairs  of  rab- 
bits (!)  which,  as  everywhere  else,  will  take  what  the  sheep  leave;  nor  that  a  parti- 
cularly dangerous  weed,  the  zarzamora  [Rubus  ulmifolius),  a  wellknown  pest  on  the 
mainland,  was  introduced  on  purpose  to  be  used  as  living  fences.  The  success  was 
comj)lete.  The  thruslies  took  care  of  the  dispersal  of  the  fruits,  invasion  is  going 
on  everywhere,  and  I  have  just  been  told  that  the  entire  Plazoleta  del  Yunque  pre- 
sents the  picture  of  an  impenetrable  "living  fence".  From  the  same  source  I  have 
the  information  that  the  unlawful  cutting  of  the  Chonta  has  not  been  stopped  — 
I  doubt  that  anybody  tried  to  stop  it  —  but  is  going  on  just  as  before  and  with 
the  same  method:  before  a  tree  is  felled  a  cut  with  an  ax  is  made  near  the  base 
in  order  to  know  the  thickness  of  the  wood  cylinder;  if  it  is  too  thin  to  be  of 
value  the  tree  is  left  standing,  a  potential  victim  of  fungus  attacks. 

It  remains  to  be  mentioned  that  17  colonists  settled  on  Masafuera  some 
years  ago  (72  {)ersons  in  all).  Their  principal  occupation  is  said  to  be  lobster 
fishing. 

Basing  his  opinion  on  his  [)crsonal  knowledge  of  the  islands  and  on  a  wealth 
of  material  from  various  sources,  (Guzman  discusses  the  present  situation  and  the 
{)ossibilities  to  improve  it.  There  can  be  no  doubt  that  he  is  interested  in  the 
f)reservation  of  the  nature,  but  it  is  also  evident  that  he  greatly  underrates  the 
dangers  jeopardizing  the  survival  of  the  indigenous  flora  and  fauna.  The  pros  and 
cons  in  agriculture  and  silviculture  are  set  forth  in  detail.  In  his  appreciation  of 
the  fruit  r^roduccd  he  differs  cf)nsi(lerably  from  Johow  who  did  not  hold  a  very 
high  opinion  of  the  (juality,  and  our  impression  was  not  too  favourable.  Oranges 
and  lemons,  not  grown  in  our  time,  are  said  to  prosper  in  the  valleys  —  this  may 
be  true,  but  what  are  the  fruits  like.'  The  wheat,  of  poor  quality  according  to 
Johow,  is  excellent,  which  must  mean  that  new  strains  have  been  introduced,  A 
circumstance  in  favour  of  agriculture,  Guzman  says,  is  that  as  a  national  park  all 
the  land  belongs  to  the  State,  and  the  concessioned  farmer  needs  not  worry  about 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  I  87 

rents  or  amortizations.  On  the  other  hand,  Guzman  continues,  the  factors  render- 
ing agriculture  difficult  are  also  conspicuous.  The  men  working  for  the  companies 

—  there  are  three  of  them,  but  the  Recart  y  Doniez  Co.  was  taken  over  "en 
estado  ruinoso"  by  Cia  Santa  Sofia  —  earn  much  more  money  than  the  farm  hands, 
with  the  result  that  everybody  prefers  to  be  a  fisherman.  The  schooners  cannot 
accommodate  the  export  of  fruit  or  vegetables.  Good  soil  is  rare  and  shallow, 
because  the  islands  are  too  young  (!).  The  zarzamora  is  taking  possession  of  the 
best  agricultural  soil  and  is  becoming  a  serious  problem.  The  strong  winds  are 
another  inconvenience,   to  be  overcome  by  planting  hedges. 

The  most  serious  obstacle  is  the  very  limited  extension  of  arable  land,  referred 
to  above  p.  171;  300  hectares,  in  9  different  localities,  including  Masafuera  and 
Santa  Clara,  might  be  utilized,  with  an  annual  output  of  cereals  amounting  to 
2300  quintales.  If  800  are  reserved  for  local  consumption  and  300  for  seed-corn, 
1200  remain  for  export  (I.e.  p.  160)  to  compete  on  the  market  with  millions  of 
quintales  produced  on  the  mainland,  and  having  to  carry  the  cost  of  freight!  It 
it  disgusting  to  find  that  such  imaginary  hopes  have  been  carried  from  genera- 
tion to  generation.  How  much  of  the  land  is  level?  Almost  nothing.  And  what 
would  happen  if  the  slopes,  where  the  inclination  rarely  falls  below  15  to  20°, 
and  the  soil  is  very  thin,  are  plowed?  The  badly  eroded  areas  seen  everywhere 
give  the  answer.  When  Guzman  recommends  the  declivities  for  tree-planting  he 
gets  on  safer  ground.  And  he  believes  that  reforestation,  large  olive  plantations 
and  the  creation  of  orchards  and  artificial  meadows  would  transform  Masatierra 
into  an  "alegre  huerto  isleno,  deleite  de  los  turistas"  (I.e.  p.    164). 

Masatierra  is  already  blessed  with  sheep,  cattle,  goats  and  rabbits,  all  in  excess 

—  will  it  also  be  necessary  to  add  a  host  of  tourists?  Guzman  holds  a  very  high 
opinion  of  the  attractions  of  Juan  Fernandez.  He  quotes  "el  celebre  viajero  Carlos 
Rowsel"  (I  regret  never  having  heard  his  name  before)  who  once  said  that  if 
Masatierra  were  situated  60  instead  of  360  miles  from  Valparaiso  it  would  be  "la 
Reina  de  los  Balnearios"  in  the  Pacific  —  and  this  in  spite  of  the  fact  that  there 
is  not  the  slightest  trace  of  a  bathing  beach  on  the  island.  And  when  Guzman 
compares  Juan  Fernandez  to  Hawaii  and  finds  that  the  one  is  just  as  marvellous 
as  the  other,  he  has  lost  all  contact  with  reality.  I  should  add  that  I  happen  to 
be  very  well  acquainted  with  both,  and  that  nobody  could  appreciate  the  beauties  of 
Juan  Fernandez  more  than  I  do.  Call  it  a  miniature  Hawaii  if  you  like,  but  deprived 
of  the  tropical  luxuriance,  the  colourful  Royal  history  of  the  natives,  the  white, 
palm-shadowed  beaches  of  coral  sand,  the  gigantic  mountains  and  active  volcanoes, 
not  to  mention  the  comfort  and  luxury  offered  by  the  busy  cities,  the  large  hotels, 
beautiful  camp  sites,  excellent  motor  roads  and  easy  and  rapid  communications 
by  sea  and  air  with  the  outside  world!  The  nature  of  the  indigenous  living 
world  is,  in  principle,  of  the  same  type  in  the  two  cases,  but  incomparably  richer 
and  more  varied  in  Hawaii  —  there  is  room  for  half  Masatierra  in  the  caldera  of 
Haleakala.    To    the  average  tourist  Masatierra  remains  Robinson  Crusoe's  island 

—  this  is  its  main  attraction.  I  have  no  doubt  that  the  traffic  will  grow  and  the 
tourists  become  a  moderate  source  of  income  which  nobody  will  envy  the  kind  and 
hospitable  colonists,  but  I  am  afraid  that  Guzman  is  too  optimistic  when  he  thinks 


I  88  C.   SKOTTSBERG 

that  the  day  is  near  when  Juan  Fernandez  will  become  "el  balneario  habitual  de 
los  sonadores  del  mundo"  (p.  229)  and  when  large  passenger  planes  will  land  on 
the  big  aerodrome  on  Masatierra.  At  present  not  even  a  tiny  plane  is  able  to  land 
without  crashing,  but  there  are  some  places  where  an  autogiro  could  come  down 
safely.  The  roadstead  in  Cumberland  Bay  is  so  far  the  only  place  where  a  plane 
(a  Catalina)  has  alighted,  but  with  a  high  wind  and  a  heavy  swell  the  situation 
gets  unpleasant.  The  level  land  by  the  sea  in  front  of  the  colony  is  the  only 
place  where  a  small  airport  could  be  constructed.  Its  length  will  be  500  m  or,  if 
the  front  slope  of  Cordon  Central  with  the  old  prison  caves  be  blasted,  perhaps 
800  m,  but  the  buildings  belonging  to  the  fishing  companies  and  a  good  many 
other  houses  would  have  to  disappear.  On  the  table-land  of  Masafuera,  about 
1200  m  above  sea  level,  another  airport  could  be  built  at  enormous  cost,  another 
one,  probably  a  little  cheaper,  on  the  Loberia  plain.  Neither  would  serve  any 
sensible  purpose  if  not  a  strategic  one.  Let  us  limit  our  plans  of  making  Juan 
Fernandez  a  popular  goal  of  the  tourist  to  an  improvement  of  the  communications 
and  to  a  couple  of  modest  guest-houses  —  and  to  impress  on  the  visitor  that 
he  finds  himself  in  a  sanctuary  where  he  has  to  keep  his  hands  off.  An  unknown 
number  of  plants  and  animals  barely  manage  to  hold  their  own,  and  obviously 
many  are  on  the  verge  of  extinction,  pronouncedly  stenotopic  as  they  are.  The 
trail  to  Portezuelo  ought  to  be  improved  and  kept  in  repair,  for  everybody  will 
want  to  see  Selkirk's  Lookout,  to  read  the  memorial  tablet  and  behold  the  grand 
views.  And  there  is  no  point  within  easy  reach  where  the  endemic  flora  is  —  or 
was,  at  least,  in    191 7  —  better  displayed. 


If  we  want  to  preserve  a  unique  living  world  of  very  great  scientific  interest 
and  as  such  belonging  not  to  a  single  country  but  to  the  whole  world,  these  are 
the  rules: 

to  limit  plantations  and  fields  to  the  waste-land  on  the  north  side  of  Masa- 
tierra; 

to  encourage  gardening  for  local  consumption; 

to  declare  war  on  the  introduced  noxious  weeds,  goats  and  rabbits; 

greatly  to  reduce  the  number  of  domestic  animals  and  to  keep  them  out  of 
the  native  forest; 

to  reduce  the  number  of  wild  goats  on  Masafuera  and  keep  it  on  a  minimum 
or,  which   would  be  the  best,  to  exterminate  them; 

to  teach  the  inhabitants  not  to  disturb  Nature's  equilibrium; 

to  enforce  the  Law  of  Jan.  31,  1935,  by  appointing  a  sufficient  number  of 
salaried  supervisors  and  guardians. 

Human  influence  has  cut  its  mark  deep  and  it  has  changed  the  natural  scen- 
ery greatly  without  adding  to  its  beauty.  In  part  this  has  been  inevitable,  if  man 
was  to  live  on  the  islands,  but  there  has  been  and  is  too  much  senseless  destruc- 
tion. This  is  the  more  to  be  regretted  as  the  welfare  of  the  population  need  not 
at  all  depend  on  either  breeding  cattle  or  sheep-farming.  Once  the  lobster  meant 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS 


189 


Fig.  100.  Large  specimen  of  bacalao,  Cumberland  Bay.  The  gentleman  is  Don  Natalio  San- 
chez,   subdelegado    civil    and    the  highest  authority  in  the  islands  at  the  time  of  our  visit.  — 

Photo  K.  Backstrom  1917. 


90 


C.   SKOTTSHERG 


everjthint;.  To-day  we  can  speak  with  Guzman  of  the  "Crepusculo  de  la  langosta". 
The  situation  is  ahirniinir,  he  calls  for  measures  to  put  an  end  to  the  decline,  and 
he  j)roposes  certain  ways  and  regulations.  I  would  like  to  add  that  in  a  case  as 
serious  as  this  the  best  thing  to  do  is  to  {)roclaim  the  whole  year  a  closed  season 
tluring  a  sufhciently  long  period.  An  industry  based  on  the  enormous  supply 
of  fisii,  with  the  bacalao  (hg.  lOo)  heading  the  list,  would,  if  carefully  handled, 
become  a  profitable  enterprise.  A  ship  with  modern  equipment  would  bring  the 
frozen  fish  to  the  mainland  where  it  would  find  a  ready  market.  The  new  methods 
of  cold  storage  permit  deep-frozen  food,  fish  not  excepted,  to  keep  absolutely 
fresh  during  any  length  of  time.  Beside  the  bacalao,  the  most  abundant  of  all  and 
considered  to  be  the  most  delicious,  there  are  several  other  kinds  of  commercial 
value  unknown  on  the  coast  of  Chile.  1  shall  add  one  more  remark.  Cumberland 
l^a\-  ought  to  be  an  ideal  place  for  a  biological  station.  The  fauna  and  flora, 
terrestrial  as  well  as  marine,  ofi'ers  a  wide  field  for  scientific  research,  no  less 
important  from  a   {practical   viewpoint. 

The  present  management  of  these  precious  islands  is  not  in  good  accordance 
with  the  intrinsic  meaning  of  the  Law  and,  in  some  cases,  directly  violates  it. 
If  the  responsible  authorities  do  not  change  their  attitude,  Juan  Fernandez  will 
become  a  second  Saint   Helena  and  a  disgrace  to  an   enlightened  world. 

S[)eaking  as  on  behalf  of  the  Chilean  nation,  I  would  like  to  say,  with  a 
slight  alteration,   what  (iuzman  said  about  the  threatened  langosta  (p.  223): 

\o  permitamos  que  nuestras  islas  se  transformen  en  el  recuerdo  de  una 
riqueza  extinguida  en  forma  irreparable.  Asi  lo  exige  nuestro  prestigio  de  nacion 
civilizada,  asi  lo  esperan  de  nosotros  las  generaciones  futuras. 

December   1953. 

Bibliography. 

Andkkson,   Cj.  \V.    a    new,    authentic    and    complete    collection    of  voyages  round  the 

world.    I-ondon.    No  date.  (Byron's  and  Carteret's  visits  to  Juan  Fernandez.) 
Anson,   ().   A   voyage  round  the  world  in  the  years   1740 — 44.  Compiled  by  Richard 

Wai.tkr.    London    1749. 
Branchi,    K.  C.    La   Isla  de  Robinson.   Valparaiso    1922. 
Brugcen,   M.J.    Inmdamentos  de  la  geologia  de  Chile.   Santiago    1950. 
liiRNKv,  James.   A   (  hronological  history  of  the  discoveries  in  the  South  Sea  or  Pacific 

()(can.   London.   Vol.   Ill   (1813,   L'Heremite),   IV  (1816). 
P.lr(;er,   O.    Die   Rohinson-Inscl.    Leipzig    1909. 

Krmki,,    a.    Line   Rcise  nach   dcr   Rohinson-Crusoe-lnsel.   Hamburg    1889. 
I'RODIN,  B.    Kx|)eflition   na  svenskt  uppdrag   kartlade   sagoon   i   Stilla  havet.   (Expedition 

on    Swedish   initiative   map])cd   the   fairy   isle   in   the   Pacific   Ocean.)  Dagens  Ny- 

hetcr  2«  ^j    1(^-2. 
(Irak  AL\rin,   A.    l'rol)lenias    c(  onoinicos    de    Juan    Fernandez.   Departamento    General 

de    la    Pro(hu(i6n.    ()fi(  ina    de    Enlace    Agn'cola.    Memorandum    158.   Santiago, 

29   dc   Ahril   de    1944.   ('fypcwrittcn.) 
Cramam,    Maria.  Journal   of  a   residence   in   Chile  during  the  year  1822.  London  1824. 
(iCNitn-.R,    E.    Derrotero   de   la   costa   de   Chile.    V.   Anuario   Hidrografico   de   la  Marina 

de   Chile.    32.    1920. 
(jfZMAN    Parada,  J.   Cutiihres   oceanicas.   Santiago.   No   date. 


A  GEOGRAPHICAL  SKETCH  OF  THE  JUAN  FERNANDEZ  ISLANDS  I9I 

Hagerman,  T.  H.   Beitrage  zur  Geologie  der  Juan  Fernandez-lnseln.  Nat.  Hist,  of  Juan 

Fernandez  and  Easter  Island.  1:  i.    1920. 
Hawkesworth,  J.    An  account  of  the  voyages  undertaken   ...   for  making  discoveries 

in  the  Southern  Hemisphere.  I.  London  1783.  (Byron's  and  Carteret's  journeys.) 
Instrucciones    nauticas    de    la    costa    de  Chile.  Anuario   Hidrogrdfico  de  la  Marina  de 

Chile.  XX.    1896. 
JoHOw,  F.  Estudios  sobre  la  Flora  des  Islas  de  Juan  Fernandez.   Santiago    1896. 
Juan,   G.  y  Ulloa,  A.  de.  Relaci6n  historica  del  viage  a  la  America  meridional.  Ft.  2, 

vol.   III.   Madrid    1748. 
Lopez,  Juan    E.    Esploracion    de    las    islas  esporadicas  de  la  costa  de  Chile.   Anuario 

Hidrografico  de  la  Marina  de  Chile.  II.  Santiago    1876. 
Philippi,    R.    a.    Geognostische    Beschaffenheit  der  Insel  Masafuera.   Neues  Jahrb.   fiir 

Mineralogie  etc.   Stuttgart   1857. 
Philippson,    a.    Die    Erosion    des  fliessenden   VVassers  und  ihr  Einfluss  auf  die  Land- 

schaftstypen.  Geogr.  Bausteine  herausgeg.  von  Dr.  Hermann  Haack.  Gotha  1914. 
Poehlmann,   R.  Noticias  preliminares  sobre  las  condiciones  jeograficas  i  jeolOjicas  del 

Archipielago  (de  Juan  Fernandez).   In:  Johow,   Estudios. 
QuENSEL,  P.  (i)   Die  Geologie  der  Juan  Fernandez-lnseln.   Bull.  Geol.  Inst,  of  Upsala. 

XI.    1912. 
(2)  Additional  Comments  on  the  Geology  of  the  Juan  Fernandez  Islands.   Nat. 

Hist,  of  Juan  Fernandez  and  Easter  Island.   I:  3.    1952. 
Skottsberg,   C.  (i)  The  Wilds  of  Patagonia.   London    191 1. 

(2)  Till  Robinson-on  och  varldens  ande.   Stockholm  191 8. 

■ — ■ —  (3)  The  vegetation  of  the  Juan  Fernandez  Islands.  Nat.  Hist,  of  Juan  Fernandez 

and   Easter  Island.   II:  29.    1953. 
SuTCLiFFE,  Th.  (i)  Sixteen  years  in  Chile  and  Peru  from  1822  to  1839.  London  1841. 
(2)  Crusoniana  or  Truth  versus  Fiction,  elucidated  in  the  history  of  the  Island 

of  Juan  Fernandez.  Manchester   1843. 
Tenz,    O.    Una    ascensi6n  emocionante  a  la  cumbre  del  Monte  Yunque.  El  Mercurio 

V3   192  T.  Valparaiso. 
ViDAL  GoRMAz,  F.  Jeografia  nautica  de  la  Repiiblica  de  Chile.  Anuario   Hidrografico 

de  la  Marina  de  Chile.   VII.    1881. 
ViEL,    O.    Islas  de  Juan  Fernandez.  Anuario  Hidrografico  de  la  Marina  de  Chile.  IV. 

1878. 
Walter,  R.  See  Anson. 
Walpole,  F.  Four  years  in  the  Pacific  in  Her  Majesty's  ship  "Collingwood",  from  1844 

to    1848.  London    1850. 
Weber,   Hugo.   Signalmaat  Weber.  Zehn  Jahre  auf  der  Robinson-Insel.  Reutlingen  1940. 


Addition. 

This  paper  was  in  page  proof  when  my  friend  Mr.  G.  Looser  sent  me  a  report 
written  by  Dr.  Alberto  Graf  Marin,  Head  of  the  Oficina  de  Enlace  Agricola  in  Sant- 
iago, and  kindly  put  at  my  disposal  (see  Bibliography).  Dr.  Graf  Marin  visited  Masa- 
tierra  in  January  1944  in  order  to  inform  himself  of  the  economic  situation  of  the 
population,  452  persons  in  1943,  a  figure  very  different  from  the  one  quoted  by  Guzmiin 
for  the  same  year,  771;  both  cannot  be  correct.  Ciraf  Marin  judged  the  living  condi- 
tions to  be  comparatively  good,  even  if  much  remained  to  be  improved  in  the  way 
of  housing,  teaching,  medical  care,  entertainment,  travel  facilities,  etc.  It  goes  without 
saying  that  lobster  fishing  gave  the  main  income;  again,  there  is  a  discrepancy  be- 
tween his  figures  and  those  of  Guzman:  84  100  lobsters  shipped  to  the  mainland  in 
1943,  whereas  the  total  catch  was   62  287   according  to  Guzmdn.  The  average  annual 


IQ2  C.  SKOTTSBERG 

profit  derived  from  the  export  of  wool  was  about  ^/e  of  the  income  from  fishing,  the 
number  of  sheej)  3000.  Finally,  some  mutton  was  sold  to  Cia  Oto  Hnos,  apparently 
the  only  fishing  enterprise  in  operation  (comp.,  however,  p.  187).  With  regard  to  the 
lobster  (iraf  Marin  was  told  that  it  breeds  the  year  round  whence  follows  that  the 
fears  later  expressed  by  Guzman  would  seem  groundless  and  a  closed  season  —  after 
iq^i    16 — 309   —   unnecessary. 

Although  Ciraf  Marin  recognizes  the  fishing  industry  as  the  main  source  of  wealth 
he  revives  the  old  dream  of  Masatierra  as  a  future  rich  farming  country,  and  his  plans 
were  drawn  up  without  regard  to  the  fact  that  Masatierra  forms  part  of  a  National 
Tark,  protected  by  a  law  which,  nevertheless,  he  does  mention  in  passing.  He  recommends 
to  import  farmers  from  the  region  of  Concepcion,  where  the  climate  is  similar.  As 
the  soil,  save  for  the  houses  and  gardens,  is  fiscal  property,  anybody  is  entitled  to 
supply  himself  with  wood  in  the  forest  —  only  maqui  is  used  for  fuel,  however.  The 
forest  should  be  subject  to  economic  management  by  multiplying  the  few  good  timber- 
trees  and  by  introducing  new  ones;  the  Mediterranean  cork-oak  is  particularly  recom- 
mended beside  Acacia  tnelanoxylon  and  Pinus  radiata.  Of  the  endemic  trees  two  are 
(or  rather  were)  of  commercial  value,  the  chonta  palm  and  the  sdndalo;  the  former  — 
and  also  the  latter,  if  it  could  be  rediscovered  —  ought  to  be  propagated  in  nurseries 
and  planted  on  a  grand  scale.  He  tells  us  that  the  last  living  sandal  tree  seen  by 
lohow  in  1892  and  by  me  in  1908  was  ordered  by  the  Subdelegado  Vera  in  1918 
or  19  to  be  cut  down  —  but  in  1916  I  was  assured  by  the  same  man  who  brought 
me  to  the  tree  in  1908  that  it  had  died  and  the  wood  been  taken  care  of.  I  have 
no  reason  whatsoever  to  doubt  the  correctness  of  his  statement. 

With  regard  to  the  black  pest  Antennaria  (Limacinia)  Graf  Marin  shares  my 
opinion  that  Johow  exaggerated  its  dangerousness.  A  more  serious  enemy  of  the  luma 
is  an  insect,  Saissetia  oleae,  and  he  recommends  the  introduction,  successfully  arranged 
for  on  the  mainland  by  himself,   of  its  parasites. 

Even  if  various  fruits  can  be  produced  in  sufficient  quantity  to  be  exported  — 
the  water-supply  is  good  and  the  streams  might  be  canalized  and  used  for  irrigation  — 
raising  live  stock  should  constitute  the  main  occupation  of  the  farmers,  but  it  is 
jeopardized  by  the  spread  of  the  zarzaparilla  {Acaena  argentea)  and  of  the  zarzamora 
[Rubus  ulmifolius).  They  ought  to  be  exterminated,  which  is  just  as  desirable  from 
the  naturalist's  viewpoint,  but  much  easier  said  than  done.  In  order  to  improve  the 
pasture  alfalfa  and  new  and  more  nutritious  foreign  grasses  should  be  introduced.  Above 
all,  to  stimulate  general  utilization  of  the  land  and  to  prevent  that  a  single  thrifty 
person  take  possession  of  most  of  it,  the  island  ought  to  be  parcelled  and  the  size 
of  the  lots  fixed  with  regard  to  their  rentability.  The  total  value  of  Masatierra,  settle- 
ments not  included,  is  said  to  be  220000  pesos,  of  Masafuera  1 20  000  (considerably 
more  to-day  that  10  years  ago).  From  a  scientific  and  scenic  viewpoint  the  value  of 
these  islands  cannot  be  expressed  in  figures  —  it  is  inestimable.  If  the  plan  set  forth 
in  (iraf  Marin's  interesting  report  materializes,  Juan  Fernandez  will  stand  as  a  unique 
e\am|)lc  of  a   National   Park,  the  home  of  a  unique  fauna  and  flora,  offered  for  sale. 


5.  Derivation  of  the  Flora  and  Fauna  of  Juan  Fernandez 
and  Easter  Island. 

By 
C.  SKOTTSBERG. 

With   I   Map. 

Part  1. 
THE  JUAN  FERNANDEZ  ISLANDS. 

Chapter  I. 
Composition,  distribution  and  relationships  of  the  Flora. 

The  statements  made  below  are  based  on  the  results  laid  down  in  vol.  II  of 
this  work  and  in  a  number  of  papers  published  from  time  to  time  prior  to  the 
survey  in  the  field  undertaken  by  myself  and  collaborators  during  a  three  months' 
stay  in  the  islands,  Dec,  1954-March,  1955.  Later  on  some  minor  changes 
will  have  to  be  made  in  the  lists  of  the  Vascular  Plants,  but  as  they  will  not  alter 
the  conclusions  drawn  I  have  preferred  not  to  include  them  here,  more  so  as  the 
species  in  question,  some  of  them  at  least,  have  to  be  subjected  to  a  close  taxo- 
nomical  study.  With  regard  to  the  Cryptogams,  to  be  referred  to  a  number  of 
specialists,  the  revision  of  the  collections  will  take  some  time,  but  even  if  a  few 
new  species  will  be  described,  and  other  additions  made  to  the  lists,  the  pro- 
portions between  the  various  geographical  elements  will,  I  think,  remain  much  the 
same. 

I.  Angiospermae. 

In  the  islands  42  families  are  represented  of  which  one,  Lactoridaceae,  is 
endemic  and  monotypic.  Until  recently  Juan  Fernandez  (Masatierra)  was  the  only 
oceanic  island  possessing  an  endemic  family,  but  after  the  discovery  of  the  genus 
Degeneria,  which  forms  the  monotypical  family  Degeneriaceae,  it  shares  this 
honour  with  Fiji;  the  Fiji  group  is,  however,  not  as  "oceanic",  in  the  current 
sense  of  this  word,  as  Juan  Fernandez. 

The  largest  families  are  Compositae  with  28  (13  gen.),  Cyperaceae  with  14 
(7  gen.),  and  Gramineae  with  13  species  (10  gen.),  but  these  are  very  large  fami- 
lies almost  everywhere;  Campanulaceae  (2  gen.)  and  Juncaceae  have  6  (2  gen., 
but  some  species  of  Juncus  may  not  be  truly  indigenous),  Rubiaceae  5  (4  gen  ), 

13  "~  557857  The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.    Vol.  I 


194  ^-   SKOTTSBERG 

Umbelliferae  5  (3  gen.),  Chenopodiaceae  4  (2),  Myrtaceae  4  (4),  Piperaceae  4(1), 
Rosaceae  4  (3),  Solanaceae  4  (2),  and  Urticaceae  4  (3),  Caryophyllaceae  3  (2), 
Cruciferae  3  (i),  (iunncraceae  3(1),  Halorrhagidaceae  3  (i),  Berberidaceae  2  (i), 
Ikomeliaceae  2  (2),  Convolvulaceae  2  (2),  Labiatae  2  (i),  Leguminosae  2  (i), 
Plantaginaceae  2  (1),  Rutaceae  2  (i),  Scrophulariaceae  2  (2);  represented  by  i 
species  only:  Aizoaceae,  Horaginaceae,  Callitrichaceae,  Empetraccae,  Ericaceae, 
luipliorbiaceae,  Flacourtiaceae,  Iridaceae,  Lactoridaceae,  Loranthaceae,  Palinae, 
Ranunculaceae,  Rliamnaceae,  Santalaceae,  Saxifragaceae,  Verbenaceae,  Wintera- 
ceae,  17  families  or  40%  of  the  entire  number.  Some  of  the  families  are  alto- 
gether small,  but  others  are  large  and  widespread  also  in  Andean  America — 
as,  for  instance,  Umbelliferae-Hydrocotyloideae  [Azorella\  Caryophyllaceae,  Cruci- 
ferae, Labiatae,  Leguminosae  (Adesniia,  Cassm),  Plantaginaceae,  Scrophulariaceae 
[Calceolayia],  Horaginaceae  [Plagiobotrys  and  related  genera),  Euphorbiaceae,  Iri- 
daceae [SisyrijuJiiuui],  Ranunculaceae,  Rhamnaceae,  Verbenaceae.  The  absence 
of  such  families  as  Amaryllidaceae,  Asclepiadaceae,  Eagaceae,  Geraniaceae,  Lau- 
raccac,  Liliaceae,  Xolanaceae,  Onagraceae,  Orchidaceae,  Oxalidaceae,  Polygona- 
ceae,  Portulacaceae,  Valerianaceae  and  Violaceae  is  noteworthy,  being  more  or 
less  well,  some  of  them  very  well,  represented  on  the  Chilean  mainland. 

Of  the  42  families  Masatierra  possesses  39,  seven  of  these  also  found  on 
its  satellite  Santa  Clara,  and  Masafuera  30;  12  are  confined  to  Masatierra  (Plan- 
taginaceae also  on  Santa  Clara)  and  3  to  Masafuera;  27  are  common  to  Masa- 
tierra and  Masafuera.  The  families  are,  in  contradistinction  to  the  genera  and 
species,  evenly  distributed  over  the  group,  as  shown  by  the  list  of  Genera  and 
Species,  Table  I. 

The  number  of  genera  is  89;  of  these  71  are  found  on  Masatierra,  9  on 
Santa  Clara  and  54  on  Masafuera;  35  are  known  from  Masatierra  only,  including 
those  known  from  Santa  Clara,  all  also  found  on  Masatierra;  18  are  confined 
to  Masafuera,  36  shared  by  both  islands.  Expressed  in  percentage:  Masatierra 
39.3%,  Masafuera  20.2%,  Masatierra  +  Masafuera  40.5%.  Thus  less  than  one 
half  of  the  genera  are  common  to  both  islands.  The  floristic  difi"erence  between 
them  is  [)artly  explained  by  the  difi"erence  in  altitude  and  thereby  in  climate — 
Masafuera  has  an  ali)ine  and  subalpine  flora  for  which  there  is  no  room  on 
Masatierra,  partly  by  the  more  varied  topography  of  this  island,  which  has  a  richer 
flora.  '1  he  difference  becomes  particularly  obvious  when  endemism  is  considered. 

Of  the  89  genera,  17  (19  ?{,)  are  endemic;  of  these  12  (70.6  %)  are  confined 
to  .\L'isatierra,  i  (5.9%)  to  Masafuera  and  4  (23.5%)  found  on  both  islands;  see 
Table   II. 

Of  71  genera  known  from  Masatierra  16  (22.5  %)  are  endemic  to  the  islands, 
of  54  found  on  Masafuera,  5  (9.3  %);  of  35  genera  only  found  on  Masatierra,  12  are 
endemic  1 34. 3  "„),  the  corresponding  figures  for  Masafuera  are  18,  I  and  5.5  % 
and,  for  the  genera  occurring  on  both  islands,  36,  4  and  1 1. 1  %.  These  figures 
serve  to  illustrate  the  great  difference  in  the  distribution  of  the  endemic  genera 
commonly  looked  upon  as  ref)resenting  the  most  ancient  element  among  the 
Angiosperins  in  Juan    h'ernandez. 

Ot   the  total   number  of  species,    147,  99  are  found  on  Masatierra,  9  on  Santa 


DERIVATION  OF  THE  FLORA  AND  FAUNA 


195 


Table  I. 
Endemics  bold-faced,  endemic  genera  in  capital  letters. 


Abrotanella  crassipes  Skottsb 

Acaena  ovalifolia  Ruiz  et  Pav 

—  masafuerana  Bitter 

Agrostis  masafuerana  Pilger 

Apium  fernandezianum  Johow 

Azara  fernandeziana  Gay 

Berberis  corymbosa  Hook,  et  Arn 

—  masafuerana  Skottsb 

Boehmeria  excelsa  Wedd 

Callitriche  Lechleri  (Hegelm.)  Fassett 

Calystegia  tugurionim  R.  Br 

Cardamine  chenopodiifolia  Pars 

—  flaccida  Cham,  et  Schlechtd 

—  Kruesselii  Johow 

Carex  berteroniana  Steud 

—  Banksii  Boott 

CENTAURODENDRON  dracaenoides  Johow 

Centella  triflora  (Ruiz  et  Pav.)  Nannf 

Chaetotropis  chilensis  Kunth 

—  imberbis  (Phil.) 

Chenopodium  crusoeanum  Skottsb 

—  nesodendron  Skottsb 

—  Sanctae  Clarae  Johow 

Chusquea  fernandeziana  Phil 

Cladium  scirpoideum  (Steud.)  Benth.  et  Hook,  f, 

Colletia  spartioides  Bert 

Coprosma  Hookeri  (G.  Don)  W.  R.  B.  Oliver  .    . 

—  pyrifolia  (Hook,  et  Arn.)  Skottsb 

CUMINIA  eriantha  Benth 

—  fernandezia  Colla 

Cyperus  eragrostis  Lam 

—  reflexus  Vahl 

Danthonia  coUina  Phil 

DENDROSERIS  litoralis  Skottsb 

—  macrantha  (Bert.)  Skottsb 

—  macrophylla  D.  Don      

—  marginata  (Bert.)  Hook,  et  Arn 


t 

rt 
rt 

t 

(U 

u 

% 

nS 

cj 

S 

a 

(§ 

% 

^ 

S 

196 


C.   SKOTTSBERG 


j  Dichoiulra  repens  l-'orst 

!  Drim\  s  confertifolia  Phil 

Dysopsis  hirsuta  (Muell.  Arg.)  Skottsb.     .    .    . 

Kleocharis  fuscopurpurea  (Steiul.)  H.  Pfeiff. 

Empetrum  rubrum  Vahl 

Erigeron  fruticosus  DC 

—  Ingae  Skottsb 

—  Innocentium  Skottsb 

—  luteoviridis  Skottsb 

—  rupicola  Phil 

—  turricola  Skottsb 

Eryngium  bupleuroides  Hook,  et  Arn.      .    .    . 

—  inaccessum  Skottsb 

—  sarcophyllum  Hook,  et  Arn 

Escallonia  Callcottiae  Hook,  et  Am 

Euphrasia  formosissima  Skottsb 

Fagara  externa  Skottsb 

—  mayu  (Bert.)  li^ngl 

Galium  masafueranum  Skottsb 

Gnaj)halium  spiciforme  Sch.  Bip 

Gunncra  bracteata  Steud 

— •  Masafuerae  Skottsb 

—  peltata  Phil 

Halorrhagis  asperrima  Skottsb 

—  masafuerana  Skottsb 

—  masatierrana  Skottsb 

I  Hedyotis  thesiifolia  St.  Hil 

Hesperogreigia  Herteroi  Skottsb 

HESPEROSERIS  gigantea  (Johow)  Skottsb. 

JUANIA  australis  (Mart.)  Drude 

JuiK  us  capillac ens  I. am 

- —  dombev.'iuus  (iay 

—  imbri(atiis  Laliarpc       

—  l)lanif..lins  K.  I',r 

—  ])ro(cnis  I'..  Mvv 

Kock'ria  iiii(  ratlicra  (  Dcsv.)  Griseb 

LACTORIS  fernandeziana  Pliil 

Lagcnopliora  nari..1i    Iraiu  h 

Libcrtia  formosa  C.rali 


1 

0 

a 

2 

1 

_ 

+ 

— 

+ 

— 

+ 

- 

+ 

- 

+ 

- 

+ 

- 

+ 

- 

4- 

_ 

+ 

- 

+ 

- 

+ 

- 

+ 

- 

■+ 

- 

+ 

- 

+ 

- 

+ 

- 

+ 

- 

+ 

- 

+ 

- 

+ 

DERIVATION  OF  THE  FLORA  AND  FAUNA 


197 


Lobelia  alata  Labill 

Luzula  masafuerana  Skottsb 

Margyricarpus  digynus  (Bitter)  Skottsb 

MEGALACHNE  berteroniana  Steud 

—  masafuerana  (Skottsb.  et  Pilger)  Hubbard  ms.     .    .    . 

Mimulus  glabratus  H.  B.  K.  forma 

Myrceugenia  Schulzei  Johow 

Myrteola  nummularia  (Poir.)  Berg 

Nertera  granadensis  (L.  fil.)  Druce  var 

Nicotiana  cordifolia  Phil 

NOTHOMYRCIA  fernandeziana  (Hook,  et  Arn.)  Kausel 

OGHAGAVIA  elegans  Phil 

Oreobolus  obtusangulus  Gaud 

Parietaria  humifusa  Rich 

Paronychia  chilensis  DC 

Peperomia  berteroana  Miq 

—  fernandeziana  Miq 

—  margaritifera  Bert 

—  Skottsbergii  C.  DC 

Pernettya  rigida  DC 

PHOENIGOSERIS  berteriana  (Dene)  Skottsb 

—  pinnata  (Bert,  ex  Dene)  Skottsb 

—  regia  Skottsb 

Phrygilanthus  Berteroi  (Hook,  et  Arn.)  Reiche 

Piptochaetium  bicolor  (Vahl)  Presl 

Plantago  fernandezia  Bert 

—  truncata  Cham,  var 

PODOPHORUS  bromoides  Phil 

Ranunculus  caprarum  Skottsb 

REA  micrantha  Bert,  ex  Dene 

—  neriifolia  Bert,  ex  Dene 

—  pruinata  (Johow)  Skottsb 

Rhaphithamnus  venustus  (Phil.)  B.  L.  Robins 

RHETINODENDRON  Berterii  (Dene)  Hemsl 

ROBINSONIA  evenia  Phil 

—  gayana  Dene 

—  gracilis  Dene 

—  Masafuerae  Skottsb 

—  thurifera  Dene 


03 

0 

1 

03 

+ 

_ 

+ 

- 

- 

+ 

+ 

- 

- 

+ 

- 

+ 

- 

- 

+ 

- 

- 

+ 

- 

- 

+ 

- 

- 

+ 

- 

- 

+ 

- 

- 

+ 

+ 

- 

- 

+ 

- 

- 

- 

- 

+ 

+ 

- 

+ 

+ 

- 

- 

+ 

- 

+ 

+ 

- 

+ 

+ 

- 

- 

- 

- 

+ 

+ 

- 

+ 

+ 

- 

- 

+ 

-  . 

— 

- 

- 

+ 

+ 

- 

- 

+ 

- 

- 

+ 

- 

- 

+ 

+ 

- 

+ 

- 

- 

- 

- 

+ 

+ 

- 

- 

+ 

- 

+ 

+ 

- 

+ 

- 

+ 

+ 

- 

— 

+ 

- 

+ 

- 

- 

+ 

— 

— 

- 

- 

+ 

+ 

- 

- 

198 


C.  SKOTTSBERG 


Riibus  geoides  Sm j 

Salicornia  fruticosa  l\unth j 

i  Santalum  fernandezianum  F.  Phil j 

Scirpus  cernuus  \'ahl      

j  —  nodosus  Rottb ! 

I  SELKIRKIA  Berteroi  (CoUa)  Hemsl 

I  Solanuni  fernandezianum  Phil 

!  —  masafueranum  Bitter  et  Skottsb 

—  robinsonianum  Hitter 

;  Sophora  fernandeziana  (Phil.)  Skottsb 

I  —  masafuerana  (Phil.)  Skottsb 

I  Spergularia  confertiflora  Steud 

I  —  masafuerana  Skottsb 

i  Stipa  (Xassella)  laevissima  (Phil.)  Speg 

—  neesiana  Trin.  et  Rupr 

SYMPHYOGHAETA  macrocephala  (Dene)  Skottsb 

i 
Tetragonia  expansa  Murr 

:  Trisetum  chromostachyum  Desv 

;  Ugni  Selkirkii  (Hook,  et  Arn.)  Berg 

j  Uncinia  brevicaulis  Thouars 

i  —  costata  Kuekenth 

—  Douglasii  Boott 

—  phleoides  Pers 

—  tenuis  Poepp 

Urtica  fernandeziana  (Rich.)  Ross 

j  —  Masafuerae  Phil 

j  Wahlenbergia  Berteroi  Hook,  et  Arn 

j  —  fernandeziana  (A.  DC.  p.  p.)  Skottsb 

j  —  Grahamae  Hemsl 

—  Larrainii  (Colla)  Skottsb 

i  —  Masafuerae  (Phil.)  Skottsb 

I  YUNQUEA  Tensii  Skottsl) 


Clara,  and  74  on  ^Ma.safuera;  6<S  -  46.3  %  (68.7  %  of  99)  are  restricted  to  Masa- 
tierra,  i  -  0.7  %  to  Santa  Clara,  and  47-32  %  (63.5  %  of  74)  to  Masafuera, 
4  -  2.7  %  are  common  to  Ma.satierra  and  Santa  Clara  with  the  exclusion  of  Masa- 
fuera, 4  -  2.7  %  found  in  all  three  islands  and  23  =  15.6%  on  Masatierra  and 
Masafuera  excluding  those  also  occurring  on  Santa  Clara.  Thus  only  27  species 
-^  18.4  °o   are  reported  from  both   Masatierra  and   Masafuera,  a  surprisingly  small 


DERIVATION  OF  THE  FLORA  AND  FAUNA 


199 


Table  II. 
Distribution  of  the  endemic  genera. 


Centaurodendron 
Cuminia  .  .  .  . 
Dendroseris  .  . 
Hesperoseris     .    . 

Juania 

Lactoris  .  .  .  . 
Megalachne  .  .  . 
Nothomyrcia  .  . 
Ochagavia  .  .  , 
Phoenicoseris  .  . 
Podophorus      .    , 

Rea 

Rhetinodendron  , 
Robinsonia  .  .  . 
Selkirkia  .  .  .  . 
Symphyochaeta  , 
Yunquea  .    .    .    . 


I      + 


number,  for  even  if  we  pay  due  attention  to  the  physiographic  difference  between 
them,  it  tells  a  story  of  effective  isolation  with  little  possibility  for  an  exchange 
across  the  92  miles  of  water  separating  them. 

Of  the  total  number  of  species  (147),  loi  are  endemic  =  68.7  %;  of  the  99 
species  found  on  Masatierra,  66  (66.'j  %)  are  endemic  in  Juan  Fernandez;  the  cor- 
responding figures  for  Masafuera  are  74  and  47  (63.5  %),  and  for  Santa  Clara  9 
and  5  (55.5  %).  Thus  endemism  plays  about  the  same  role  in  all  cases,  and  this 
is  apparent  also  when  local  endemism  is  considered.  Of  the  68  species  restricted 
to  Masatierra  50  (73.5  %)  are  local  endemics,  of  the  47  species  restricted  to  Masa- 
fuera, 34  (72.3  %);  the  single  species  restricted  to  Santa  Clara  is  endemic.  Of  the 
27  species  common  to  Masafuera  and  Masatierra  13  (48%)  are  endemic  in  Juan 
Fernandez.  If,  to  the  68  species  restricted  to  Masatierra,  4  also  found  on  Santa 
Clara  are  added,  the  figures  are  72  and  73.6%.  Of  the  total  number  of  endemic 
species  (lOi),  50  are,  as  we  have  seen,  confined  to  Masatierra  (49.5  %),  3  (3  %) 
to  Masatierra  +  Santa  Clara,  i  (i  %)  to  Santa  Clara,  34  (33.7%)  to  Masafuera, 
12  (11.9%)  common  to  Masatierra  and  Masafuera,  and  i  (i  %)  found  on  all  three 
islands.  The  difference  between  Masatierra  and  Masafuera  stands  out  even  more 
clearly.  It  is  less  pronounced  when  we  come  to  the  46  non-endemic  species:  only 
on    Masatierra    18    (39.1  %),    on    Masatierra   and    Santa   Clara   i    (2.2  %),  only  on 


C.  SKOTTSBERG 


Masafuera  13  (28.3  %),  on  Masatierra  and  Masafuera  11  (23.9  %),  and  on  all  three 
islands  3  (6.5  %).  However,  several  of  the  species  restricted  to  Masatierra  are, 
perhaps,  not  truly  indigenous,  whereas  the  indigenous  character  of  the  species 
only  recorded  from  Masafuera  cannot  be  doubted;  11  of  them  are  mountain  plants. 
With  regard  to  the  former,  some  were  commented  upon  by  Joiiow.  In  his  table 
"Continjente  B,  I^^species  autoctonas,  pero  no  endemicas",  pp.  221-222,  he  men- 
tions several  species  regarded  as  doubtful  natives:  3  species  of  Gnaphaihwi,  Mimu- 
liis,  Solanum  furcatian  ( =  robinsonianum),  Tetragonia  cxpansa,  Spergularia  rubra 
(=  confertiflora),  Parietaria  )iuiuifusa\  Trisetum  c/iroifwstachyum  and  DantJioiiia 
colliiia.  With  the  exception  of  GnapJialiian  spp.  I  have  listed  them  as  native. 
There  seems  to  be  little  reason  to  regard  Tetrago7iia,  a  wide-spread  thalassocho- 
rous  plant,  as  anthropochorous,  and  the  Spergularia  is  known  only  from  Juan 
Fernandez  and  Desventuradas  Is.  Danthoiiia  gave  me  the  impression  of  being 
just  as  autochthonous  as  PiptocJiactiuin  and  the  two  species  of  Stipa,  and  I  find  no 
good  reason  to  exclude  either  Mimulus,  Parietaria  or  Trisetum  (this  never  seen 
by  us).  With  regard  to  Solanum  "'furcatimi'  (not  furcatum  of  Dunal)  I  share 
Joi row's  doubts.  It  was  described  by  BlTTER  as  vS*.  robi^isonianum,  a  Morella  micro- 
species  of  5.  nigrum  L.  coll.  and  found  on  all  three  islands.  This  assemblage 
as  represented  on  the  mainland  has  never  been  seriously  studied.  Possibly  vS".  robifi- 
sonianum  is  a  case  of  the  same  kind  as  the  dandelion  discovered  on  the  islands 
in  19 1 7  and  described  as  Taraxaaim  ferna7idezianum  Dahlst.,  a  microspecies  of 
the  Eurasiatic  Vulgaria.  There  cannot  be  the  slightest  doubt  that  it  is  an  alien 
introduced  from  Chile,  where  these  weeds  have  not  been  studied.  From  JOIlOW's 
list  I  have  excluded  Bahia,  Amblyopappus,  Erythraea,  Monocosmia  and  Phalaris. 
Further,  some  species  not  recorded  from  Juan  Fernandez  by  JOHOW  or  earlier 
authors,  but  for  the  time  being  listed  as  native  by  me,  are  under  strong  suspi- 
cion: J  uncus  capillaceus  and  pla7iifolius,  Ce7itella  and  Hedyotis,  perhaps  also  Paro- 
7iycJiia.  If,  in  the  future,  we  shall  be  able  to  purify  the  list,  the  percentage  of 
endemic  species  will  rise  to  72  %   or  even  more. 

The  genera  richest  in  species  are  Erigero7i  with  6,  yu7icus,  Robi7iso7iia,  U7ici- 
7iia  and  \Vakle7ibergia  with  5,  and  De7idroseris  and  Pepero77iia  with  4  species  each; 
eight  genera  have  3,  15  two,  and  59  only  one  species.  The  average  number  of 
species  to  a  genus  is   1.65. 


The  systematical  position  of  the  endemites  and  the  distribution  of  the  genera  and 

species  also  found  elsewhere. 

Gramineae. 

Stipa  L.   About  250  sp.  (Bkws),  widespread  both  hemisph.,  subtrop.-temp. 

7ieesia7ia  Trin.  et  Rupr.  Mcx.  to  Boliv.  and  Chile,  Braz.,  Argent.,  Urug.  Poly- 
morphous (^^p.  771). 

Iae?rissi7na  (Phil.)  Speg.  Peru,  Chile,  Argent.  With  a  number  of  S.  Amer.  sp. 
referred  to  a  separate  genus  [Xassella). 


DERIVATION  OF  THE  FLORA  AND  FAUNA 


Piptochaetium  Presl.  20;  N.  Amer.  to  extratrop.  S.  Amer. 

bicolor  (Vahl)  Desv.  Chile:  Valdivia;  Braz.,  Argent. 
Podophorus  bromoides  Phil.  In  22y  I  referred  to  Brachyely thrum  Beauv.  and  Aphan- 
elythrum  Hack,  as  the  nearest  relatives.  Mr.  C.  E.  Hubbard  kindly  informed  me 
that  it  resembles  the  former  in  a  number  of  important  features  and  that  he  regards 
both  genera  as  belonging  to  a  relatively  ancient  group  of  grasses.  The  distribution 
of  Brachyely  thrum  is  disjunct  after  a  well-known  pattern:  B.  erectum  (Schreb.) 
Beauv.,  N.E.  Amer.,  and  B.  japonicum  Hack,  ex  Honda,  China,  Korea,  Japan. 
PiLGER  (/p^)  brings  this  genus  to  Festuceae-Festucinae  next  to  Aphanely thrum, 
placing  Podophorus  with  the  Brominae. 
Chaetotropis  Kunth.   2  (?3). 

imberbis  (Phil.)  Near  the  following. 

chilensis  Kunth.  Peru,  Chile,  Braz.,  Argent.,  Urug. 
Agrostis  L.  About   125;  very  wide-spread  in  temp,  and  cold  climates. 

masafuerana  Pilger.  "Gehort  in  die  Verwandtschaft  von  A.  canina  L.  In  der 
Tracht  A.  magellanica  Lam.  sehr  ahnlich"  (PiLGER  ipy).  This  is  recorded  from 
Magell.,  Falkl.,  N.  Zeal.  (55).  F.  B.  H.  Brown  (j5.  I.  84)  says  that  his  A.  rapensis 
from  Rapa  Id.  is  "very  closely  allied"  to  magellajiica,  but  to  judge  from  his 
illustrations  they  seem  to  have  little  in  common. 
Trisetum  Pers.  About  65 ;  N.  and  S.  temp,  (also  S.  Braz.). 

chromostachyum  Desv.  Centr.-S.  Chile. 
Danthonia  DC.  About  100;  essentially  southern  (S.  Amer.,  S.  and  E.  Afr.,  Austral., 
N.  Zeal.),  but  extending  north  to  N.  Amer.,  S.  Eur.,  and  India. 

collina  Phil.  S.  Chile. 
Koeleria  Pers.   60    (Domin);  N.  temp.,  S.E.  Austral.,  Tasm.,  N.  Zeal.,  S.  Amer. 
And.  to  Patag.,  Falkl. 

micrathera  (Desv.)  Griseb.  S.  Chile. 
Megalachne   Steud.  2  very   distinct   sp.,  better oniana  Steud.  and  masafuerana 
(Skottsb.  et  Pilg.)  Hubbard  ms.  Reduced  to  Bromus  by  PiLGER  {iQj),  but  restored  to 
generic  rank  in  his  posthumous  paper  on  the  system  of  the  Gramineae  [igS). 
Chusquea  Kunth.  About   100  (.?)  sp.  Mex.  to  Argent,  and  S.  Chile. 

fernandeziana  Phil.  Related  to  Chilean  species. 

Cyperaceae. 

Cyperus  L.  A  world-wide,  essentially  tropical  genus  of  about  600  sp.  (KuKEN- 
THAL). 

er agrostis  Lam.  (vegetus  Willd.).  N.  and  Centr.  Amer.  to  Braz.,  Urug.,  Argent. 
and  S.  Chile;  Easter  I.  Often  introduced  and  perhaps  not  indigenous  in  Juan 
Fernandez. 

reflexus  Vahl.  Mex.  and  Tex.  to  Braz.,  Argent,  and  S.  Chile.  Indigenous.^ 
Scirpus  L.  About  400;  cosmopolitan. 

nodosus  Rottb.  Circumpolar,  S.  temp.  zone.  Recently  the  Chilean  plant  was 
segregated  as  .S.  molinianus  Beetle,  but  I  am  unable  to  recognize  this  as  specifi- 
cally distinct  [326). 


202  C.  SKOTTSBERG 

cermius    \'ahl.    Subcosmopolitan    and    of  variable  habit;  south  to  Fueg.  and 
Falkl. 
FJeocharis  R.   Br.   A  world-wide  genus  of  about   150  sp.  (Svenson). 

fiistopiirpurea   (Steud.)   H.  Pfeifif.    S.  Chile.   Listed  before  as  a  subspecies  or 
variety  of  //.  7>iaculosa  (Vahl)  R.  Br.  (W.  Ind.-Braz.). 

Oreoholus  R.  Br.  An  austral-circump.  genus  of  6  sp.,  extending  north  to  Malaysia 
and   Hawaii. 

obtusaugiilus  Gaud.  Andes  of  Colomb.  and  Ecuad.  to  Fueg.,  Falkl. 
Cladium  R.  Br.  48  (Ki  kkxtii.\l);   widespread,  trop.-subtrop.  with  preponderance 
S.I'^.  Asia-X.  Guin.-Austral.-X.  Zeal.,  north  to  Hawaii,  W.  Ind.,  Braz.  2  boreal  sp. 

scirpoideum  (Steud.)  Benth.  et  Hook.   f.  Nearly  related  to   C.  angtistifolium 
(Gaud.)  Benth.   et   Hook.   f.   (X.   Guin.,  Tahiti,   Hawaii). 

rnciuia  Pers.   32;   circump.-austr.-subantarct.   with  outposts  north   of  the  P^quator 
in   Centr.  Amer.  and  Philipp.  Is.;  greatest  concentration  of  species  in  N.  Zeal. 

brevicaulis   Thouars.   S.  Chile    to  Fueg.,    P'alkl.;    Trist.    da  C;   St.  Paul  and 
Amsterd.  Is. 

Douglasii  Boott. 

costata  Kuekenth.  Related  to  the  former. 

pJihoidcs  Pers.   Andes  from  Colomb.  to  S.  Chile  and  Patag. 

tenuis  Poepp.   Centr.  Amer..^;  Andes  of  Chile  to  Fueg. 
Carex  L.  About   1500;   worldwide,  but  comparatively  few  trop.  sp. 

Haiiksii  Boott.  Andes  of  Centr.  Chile  to  P'ueg.  Belongs  to  the  boreal  section 
I^Vigidae-P^uliginosae  of  KlKENTIlAL. 

berteroniana  Steud.  Sect.  Echinochlaenae  of  KCkenthal,  with  few  excep- 
tions (Austral.,  Tasm.,  Norfolk  I.  and  one  Chilean  sp.)  confined  to  N.  Zeal. 

Palmae. 
Juania  australis  (Mart.)  Drude.  Affinities  undoubtedly  trop.  Andean.  Included 
in  the  Iriarteae  by  Dkl  DE,  but  perhaps  nearer  to  the  Morenieae,  or  possibly 
regarded  as  type  of  a  separate  subtribe.  HUTCIIINSON  brought  Juaiiia  next  to 
Ctroxylo}!,  following  Bextiiam  and  HooKER,  Croizat  (7/)  remodelled  Morenieae, 
including  Jutuiia  and  the  Mascarene  IlyopJiorhe,  otherwise  linked  with  Chamae- 
doreae. 

Bromeliaceae. 
Ilcsperogreigia  Berteroi  Skottsb.  2  or  perhaps  more  sp.  Subtrop.  Andean,  related 
to  (ireigia  («S  sp.,  Costa  Rica,  Venez.-S.  Chile).  LOOSER  called  Hesperogreigia 
monotypical,  because  my  {)ai)er  (2^0)  had  escaped  him;  if  not,  he  would  not  have 
written  "^'Ouiza  dc  la  afinidad  de  Greigia.^"  [ijo.  291).  Inspired  by  L.  B.  SMITH 
he  sup{)ressed  I fispcrognigia  in  an  appendix  to  his  paper  (p.  299),  an  attitude 
I  cannot  take.   The  two  genera  differ  very  much.^ 

1  .Sepals  free.  Pericarp  thin.  .Seed  testa  thin,  with  a  thick  coat  of  mucilage,  forming  a  massive 
pulp.  Raphe  inconspicuous.  .Scales  with  irrej.;ular,  more  or  less  isodiametric  cells    .  Hesperogreigia 
Sepals  connate  into  a  tube.  Pericarp  fleshy.  Testa  very  hard,  without  mucilage.  Raphe  very  con- 
spicuous. Scales  with  long  and  narrow  cells  radiating  from  centre Greigia 


DERIVATION  OF  THE  FLORA  AND  FAUNA  203 

Ochagavia  elegans  Phil.  Very  near  the  Chilean  genus  Rliodostachys\  see  22g.  iio 
and  24g.  jy^. 

Juncaceae. 

Luzula  DC.  A  world-wide  essentially  temp,  and  mainly  boreal  genus  of  about 
80  sp.,  also  represented  on  trop.  mountains  and  in  the  S.  Braz.  highland;  south 
to  Fueg.,  Falkl.  and  N.  Zeal. 

masafuerana  Skottsb.  Nearly  related  to  some  Andean  sp.  (Mex.,  Boliv.- 
Fueg.)  neading  taxonomic  revision. 

Junciis  L.  Mainly  temp.-subtrop.;  about  225.  Well  represented  S.  hemisph.  (about 
50),  especially  Austral,  and  N.  Zeal.,  less  so  temp.  S.  Amer. 

imbricatus  Laharpe.  Ecuador-S.  Chile,  Braz.,  Argent.,  Urug. 

capillaceus   Lam.    Ecuador,    Centr.    Chile,   Braz.,  Argent.,  Urug.  Introduced? 

procerus  E.  Mey.  S.   Chile. 

dombeyanus  Gay.  Peru,  Chile,  Braz.,  Argent.,  Urug. 

plaiiifolius  R.  Br.  S.  Chile,  Austral.,  Tasm.,  N.  Zeal.  Introduced.^ 

Iddaceae. 

Libertia  Spreng.   5  sp.,  3   Chile,  2  N.  Zeal. 
formosa  Grah.  S.  Chile. 

Piperaceae. 

Peperomia  Ruiz  et  Pav.  Possibly  over   1500;   pantropical. 

berteroana  Miq.  Extremely  like  P.  tristanensis  Christoph.  from  Gough  I.; 
see  24^. 

margaritifera  Bert,  ex  Hook.  Possibly  related  to  P.  Reineckei  C.  DC.  (Samoa). 

Skottsbergii  C.  DC.  Allied  to  the  former. 

These  species  form  a  separate  subgenus  Tildenidium  Skottsb.  (^^7),  which 
seems  to  stand  closer  to  palaeotropical  than  to  neotropical  groups.  P.  berteroayia 
and  tristanensis  show  affinity  to  P.  urvilleana  A.  Rich.  (Austral.,  N.  Zeal.,  Lord 
Howe  I.,  Norfolk  I.)  and  P.    Vente7iati  Miq,  (Java). 

fernandeziana  Miq.  Central  (Frai  Jorge,  Talinay)  and  S.  Chile  (Valdivia).  Be- 
longs to  subgen.  Sphaerocarpidium. 

Urticaceae. 

Urtica  L.  About  40  sp.  widely  scattered  in  temp,  zones  on  both  hemisph.,  well 
represented  in  Amer.  (13,  Mex.— Fueg.). 

Masafuerae  Phil.  Related  to   [/.  echinata  Benth.  (Ecuador). 

fernandeziana  (Rich.)  Ross.  A  very  distinct  species  without  near  relatives 
(Weddell). 

Boehmeria  Jacq.  About   100,  trop.-subtrop. 

excelsa  (Bert,  ex  Steud.)  Wedd.  Seems  to  be  more  nearly  related  to  Pacific 
than  to  American  species.  Very  similar  to  B.  dealbata  Cheesem.  (Kermadec  Is.). 


204 


C.   SKOTTSBERG 


Parictaria  L.   Some  30  sp.;   in  all  parts  of  the  globe.  Several  medit. -orient.,  others 
N.   and   S.   Amer.,  one  [P.  debilis  Forst.)  supposed  so  be  widely  distributed. 

/luniifiisa  Rich.  Chile.  Until  now  listed  as  debilis  which,  in  Weddell's  mon- 
ograph, is  a  collective  species.  I  have  shown  [2^1)  that  Forster's  species  from 
Australia,  New  Zealand  and  Polynesia  has  little  to  do  with  the  forms  referred  to 
this  but  occurring  elsewhere  and  that  none  of  the  specimens  from  S.  Amer.  seen 
by  me  belong  to  true  debilis.  Pending  a  revision  of  the  genus,  Jinmifusa  is  a  cor- 
rect name  for  the  Chilean  plant. 

Loranthaceae. 

Phrygilantlius  hjchl.  About  30;   bicentric:   Mex.-Braz.   and  Chile,  Argent.,  Urug.; 
Austral.,  X.  Zeal.,  X.  (kiin.,  Philipp.  Is. 

Berteroi  (Ih^ok.  et  Arn.)  Reiche.  Related  to  Andean  sp. 

Santalaceae. 

Sautaluui  L.  18;   Austral. ~X.  Guin.  5,  Melan.  2,  Polyn.  2,  Hawaii  7,  Bonin  i,  and 
the  following.   See  2?^  and  270. 

fernandezianutn  ¥.  Phil.  P^xtinct.  Belongs  to  sect.  Polynesica  Skottsb.;  not, 
as  has  been  said  (///),  related  to  S.  freycinetiaimni  of  Hawaii  which  belongs  to  a 
different  section  {2'/6). 

Chenopodiaceae. 

Clie)iopodiu})i   L.   Over  250;   world-wide,  but  essentially  temp. 

Sanctae  Clarae  Johow,  crusoeanum  Skottsb.  and  nesodendron  Skottsb.  On 
my  request  Dr.  P.  Aei.LKX,  who  created  sect.  Skottsbergia  to  receive  the  three 
island  endemics,  sent  me  the  following  remarks:  "Ich  wiirde  sagen,  dass  die  drei 
Arten  zu  kciner  der  uns  heute  bekanntgewordenen  Arten  irgendwelche  verwandt- 
schaftliche  Zijge  und  l^eziehungen  aufvveisen.  P^s  sind  vollig  isolierte  Typen,  Re- 
Hkte  eines  im  Meere  versunkenen  Plorenreiches."  Evidently  Dr.  Aellen  does 
not  support  the  idea  that  they  are  related  lo  pajiiculatum  Hook.  (X.  Amer.,  Peru- 
Chile)  or  oahuejise  Meyen  (Plawaii),  which  latter  has  the  same  arboreous  habit. 
Salicorjiia   L.   Widely  distributed,   about  30  sp. 

fruticosa  L.  (peruviana  Kunth).  Taken  in  a  wide  sense  Mediterr.,  S.  Afr.,  W. 
Ind.,   Polyn.;  peruricvia,   \V.   coast  of  S.  Amer.   to   S.   Chile. 

Aizoaceae. 

Tetragojiia  P.   About  25   s[).   Strongly  represented  in  S.  Afr.,  several  sp.  in  Chile. 
expajisa  Murr.   Coasts  and   islands  of  the  Pacific;  Braz. 

Caryophyllaceae. 

Spergularia  Presl.   A   widely  scattered  genus  of  about  40  sp.;   numerous  in  Chile. 
eojifertijioya  Steud.   San   Ambrosio.   Related  to  species  from  Centr.   Chile. 
masafuerana  Skottsb.   Related  to  the  former  and  to  S.  media  (L.)  Presl. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  205 

Paronychia  L.  About  40,  scattered  in  temp,  and  subtrop.  regions. 
chilensis  DC.  Centr.-S.  Chile,  S.  Braz.,  Argent. 

Ranunculaceae. 

Ranunculus  L.  Probably  over  300;  world-wide,  particularly  boreal,  numerous  sp. 
N.  Zeal. 

caprarum  Skottsb.  Apparently  with  distinct  relations  in  New  Zealand,  not 
near  boreal  or  S.  American  species  (22^.  125;  jyj). 

Berberidaceae. 

Berberis  L.  About  540  sp.  distributed  over  the  N.  hemisph.  and  extending  south 
along  the  Andes  to  Fueg.;  also  S.  Braz. 

corymbosa  Hook,  et  Arn.  Referred  to  sect.  Corymbosae  Schneid.  (3,  trop. 
And.).  Not  close  to  Chilean  species. 

masafuerana  Skottsb.  Near  the  former. 

Winteraceae. 

Drimys  Forst.  40;  29  N.  Guin.,  6  Austral.,  i  Borneo-Philipp.  Is.,  3  Mex.-Fueg., 
and  the  following. 

confertifolia   Phil.    Near   D.    Winteri  Forst.  (Centr.  and  S.  Chile  to  Fueg.). 

Lactoridaceae. 

Lactoris  fernandeziana  Phil.  With  regard  to  the  position  of  the  family  system- 
atists  disagree,  some  bringing  it  to  Polycarpicae,  others  to  Piperales.  GUNDERSEN 
(/20)  asserts  that  it  belongs  to  the  latter,  but  it  differs  from  this  in  very  important 
characters. 

Cfuciferae. 

Cardamine  L.  About   130;    world-wide,    essentially  temperate,    many  Chilean  sp. 
chenopodiifolia  Pers.  Boliv.,  Braz.,  Argent.,  Urug. 
flaccida  Cham,  et  Schlechtd.  Centr.  and  S.  Amer.  to  S.  Chile. 
Kruesselii  Johow.  Related  to   C.  vulgaris  Phil.  (S.  Chile). 

Saxifragaceae. 

Escallonia  Mutis.  About  50,  Colomb.— Fueg.,  Braz.,  Argent.,  Urug. 

Callcottiae  Hook,  et  Arn.  Occupies  a  rather  independent  position  among 
the  Chilean  sp.   [ij^]- 

Rosaceae. 

Rubus  L.  A  very  large,  temp,  and  trop. -montane  genus,  richly  developed  in  the 
N.  hemisph. 

geoides  Sm.   S.  Chile  to  Fueg.,  Falkl. 
Margyricarpus  Ruiz  et  Pav.  4  or   5,  trop.  Andes  to  Patag. 


2o6  C.   SKOTTSBERG 

digynus  (Bitter)  Skottsb.  Near  J/,  pinnatus  (Lam.)  O.  K.  (Peru,  Chile,  S. 
Hraz.,   Argent.). 

Acaoia  L.  An  austral  genus  of  over  lOO  sp.,  the  majority  in  S.  Amer.;  12 
Austral. -X.  Zeal.,  single  sf).   Calif.,   Hawaii,  the  Cape,  etc. 

masafuerana  I^itter.   Near  A.  antarctica  Hook.   f.  (Magell.,  Fueg.). 

oralifolia  Ruiz  et  Pav.  subsp.  australis  l^itter.  S.  Chile  to  Fueg.,  Falkl.  Per- 
haps accidentally  introduced  to  Juan  P"ernandez. 

Leguminosae. 

Soplioni  L.  70-80,  Old  and  New  World;  sect.  Tetrapterae  ("Edwardsia")  austral- 
circump.;    17   closeh'  related  species. 

fernandeziana  (Phil.)  Skottsb.  and  masafuerana  (Phil.)  Skottsb.  Very  near 
the  Chilean  ''tetraptera^  [S.  iiiacuabiaua  Phil.),  not  identical  with  vS".  tetraptera 
Ait.   from  X.   Zeal. 

Rutaceae. 

lui^s;ara  L.  Over  200,  pantrop.;  a  large  palaeotrop.  group  well  represented  in  the 
S.  Pacific  (Austral.,  X.  Caled.,  Polyn.),  extending  north  to  Hawaii.  The  two  Juan 
I-'ernandez  species  F.  mayu  (Bert.,  Hook,  et  Arn.)  Engl,  and  externa  Skottsb., 
which  are  very  closely  allied,  form  the  sect.  Mayni  Engl.  Related  to  W.  Pacific  sp. 

Euphofbiaceae. 

Dysopsis  Baill.   3;    i    in  Pxuador  \D. paucidentata  (M.  Arg.)  Skottsb.],    i  in  S.  Chile 
\D.  gleclio})widcs  (Rich.)  M.  Arg.   p.  p.],  and  the  following. 
hirsiita  (M.  Arg.)  Skottsb. 

Callitrichaceae. 

Calliiriche  L.   Widely  distributed  in  the  N.   hemisph.,    austral-circump.,    south  to 
Austral,  and   S.   Chile  but  absent  from  the  southern  half  of  Afr. 
LechUri  (Hegelm.)  Fassett.   S.   Chile  to  Magell.  [24^.  781). 

Rhamnaceae. 

Colli tia  C'onim.    17,   Andean  and  extratrop.   S.   Amer. 

spartioides  Ik-rt.  ex  Colla.  A  well-marked  species,  related  to  Chilean  and 
other  S.   Amer.   forms. 

Flacourtiaceae. 

Az(ir(7   Ruiz  et   Pav.   20~2 1    sj).;    18-19  Chile,    i    Braz.,    I    Argent. 

fernandeziana    (iay.    Closely    related    to  A.  scrrata  Ruiz  et  Pav.  (S.   Chile). 

Myrtaceae. 

r^'-;//  'I'urcz.    15,   Mex.,  Centr.   Amer.,  Andes,  south   to  Chiloe,  Venez.,  Braz. 

Selkirkii  (Hook,  et  Arn.)  Berg.  Probably  related  to  species  in  Centr.  Amer. 
and  \'enez.,  perhaps  also  to  f\  Ccvidollci  (Barn.)  Berg  from  Chile;  not  very  close 
to    r.  Molijiae  Turcz.  (S.   Chile;   Kauskl  in  litt.). 


DERIVATION  OF  THE  FLORA  AND  FAUNA  207 

Myrteola  Berg.   12,  Colomb.-Chile. 

mimmularia  (Poir.)  Berg.  Chile,  Cord.  Linares  to  Fueg.,  F'alkl. 
Nothomyrcia  Kausel.  A  monotypical  genus,  related  to  Chilean  genera. 

fernandeziana  (Hook,  et  Arn.)  Kausel. 
Myrceugenia  Berg.  About  40;  trop.  Andes,  S.  Braz.  (many  sp.)  and  about  20  in 
Chile,  south  to  Chiloe. 

Schulzei  Johow.  Related  to  M.  planipes  (Hook,  et  Arn.)  Berg  (S.  Chile). 

Gunneraceae. 

Gunnera  L.  About  30  sp.  Subgen.  Pangue,  16  sp.  Costa  Rica,  Colomb.-S.  Chile, 
Magell.,  Braz.;  Hawaii  2  sp.,  but  some  more,  of  doubtful  taxonomic  status,  have 
been  described.  Other  subgenera  in  the  Andes  of  Colomb.  to  Chile,  south  to 
Fueg.  and  F'alkl.;  Urug.;  S.  and  Centr.  Afr.;  Malays.-Philipp.  Is.;  N.  Zeal.,  Tasm. 

peltata  Phil,  and  Masafuerae  Skottsb.  related  to  Andean  sp. 

bracteata  Steud.  Seems  to  come  closer  to  the  Hawaiian  G.  kaMaie7isis  Rock 
than  to  S.  Amer.  species. 

Halorrhagidaceae. 

Halorrhagis  Forst.  About  80,  the  great  majority  Austral.-N.  Zeal.,  single  sp. 
scattered  north  to  Indonesia,  Philipp.  Is.,  China  and  Japan,  and  east  to  Rapa 
and  J.  Fern. 

asperritna  Skottsb.,  masatierrana  Skottsb.  and  masafuerana  Skottsb.  belong 
to  Subsect.  Cercodia  and  are  closely  related  to  H.  erecia  (Murr.)  Schindl.  (N.  Zeal.) 
and  other  Austral,  and  N.  Zeal.  sp.  All  J.  Fern,  forms  were  formerly  incorrectly 
identified  with  erecia. 

Umbelliferae. 

Cejttella  L.  29;  19  Afr.,  Madag.,  i  China,  2  Austral.,  5  trop.  Amer.  and  the 
widely  dispersed   C.  asiatica  (L.)  Urb. 

triflora  (Ruiz  et  Pav.)  Nannfeldt.  Centr.  and  S.  Chile,  formerly  included  under 
asiatica.  Introduced  .r* 

Eryngium  L.  About  230,  in  all  parts  of  the  world,  but  with  two  centres:  W.  Eur.— 
Medit.  and  trop.-subtrop.  Amer.  Poorly  represented  in  N.  Amer.  and  Australas. 

bupleuroides  Hook,  et  Arn.,  inaccessum  Skottsb.  and  sarcophyllum  Hook, 
et  Arn.  form  a  special  sect.  Fruticosa,  but  differ  mainly  in  being  arborescent  from 
the  species  occurring  on  the  mainland. 
Apium  L.  About  30;  in  all  parts  of  the  world. 

fernandezianutn  Johow.  A  well-marked  species,  probably  not  nearly  related 
to  the  Chilean  species,  but  showing  some  affinity  to  A. prosiratiini  Labill.  (Austral.) 
or  australe  Thouars  (Tristan  da  C.) 

Ericaceae. 

Pernettya  Gaud.  About   12,  6  or  7  Mex.  and  Centr.  Amer.  and  along  the  Andes 
to  Fueg.  and  Falkl.;    i   Galapag.  Is.,  2  N.  Zeal.,  2  Tasm. 
rigida  (Bert.)  DC.  A  well-marked  sp.  [2_$2). 


2o8  C-   SKOTTSBERG 

Empetraceae. 

Kmpctruin  L.  A  bipolar  genus,  E.  ?iij{runi  I.,  north,  /:.  rubrum  Vahl  south,  but 
according  to  Gool)  the  latter  is  represented  in  the  Subarctic  by  special  forms. 
See  _v^.  781. 

rubriun  Vahl.  Andes  of  S.  Chile  and  Argent,  to  Fueg.  and  Falkl.;  Tristan  da  C. 

Convolvulaceae. 

Dichondra  Forst.   5   trop. -subtrop.  Amer.,   i   X.  Zeal.,   and  the  following. 

)cpe)is  Forst.  Widely  spread  over  both  hemispheres,  north  to  N.  Amer.  and 
China,  south  to  S.  Chile  and  N.  Zeal. 
Calystegia  R.   Hr.  10-20;  temp. -subtrop.  in  all  parts  of  the  world. 

tugiiriorum  R.  Br.   S.  Chile  (Hantelmanni  Phil.)  and  N.  Zeal. 

Boraginaceae. 

Selkirkia  Berteroi  (Colla)  Hemsl.  According  to  Johnston  (148)  very  near  Hackelia 
()j)iz,  a  genus  centering  in  western  N.  Amer.,  with  outposts  in  S.  Amer.  and 
I'Airas.;  //.   revoluta  (Ruiz  et  Pav.)  Johnst.  Peru  to  Boliv.   and  Argent. 

Verbenaceae. 

Khaphitliaiinius  Miers.  2,  one  in  Centr.  and  S.  Chile.  Near  CitJiarexylon  L.,  a  neo- 
trop.  genus  of  about  20  sp.,  Mex.  and  W.  Ind.  to  Boliv.  and  Braz. 
venustus  (Phil.)  B.  L.  Robins.  A  very  distinct  species. 

Labiatae. 

Cuminia  Colla.  2.  An  isolated  genus,  referred  to  Prasioideae  by  El'LlNG  (go),  a 
palae()tr()j)ical  group  best  developed  in  Hawaii,  but  whereas  Cumiuia  has  the  drupe 
of  this  tribe,  it  has  the  corolla  of  Stachyoidcac-Me7itJiiuae,  where  BRIQUET  placed 
it;  lUkciKKS  statement  that  the  flowers  are  "Lippenbliiten"  (./j.  23)  is  erroneous. 
fernandezia  Colla  and  eriantha  Benth.  Perhaps  united  by  intermediate  forms. 

Solanaceae. 

Sola)iu))i  L.  Prcjbably  over  looo;  in  all  parts  of  the  world,  richly  represented  in 
trop.    Amer,   man\-   in   Chile. 

fernandezianiitn   Piiil.    Related  to  .S".   tKhcrosuui   L.   coll. 

masafiieranutn   Hitter  et  .Skottsb.   A   \ery  well-marked   Morella. 

robinsonianutrj   Bitter.   .See  above  j).   200. 
XicotidJKJ  L.  60;  45  Anicr.,  C"alif.-Mex.,  And.  S.  vXmer.  (hxuad.-Peru-Chile,  ]3raz.- 
Argent.-Patag.),    15    Austral,   (i    \.   Caled.,  etc.). 

coydifolia  IMiil.  Belongs  to  the  Rustica  group,  confined  to  trop.  Amer.  and 
Australas.,  and  related  to  X.  Raijiioiidii  ^lacbr.  (Andes  of  S.  Peru),  solaJiifolia 
\\'al[).  (.\.  and  Centr.  Chile)  and  piviiculata  L.  (Perii,  \.  Chile).  Dr.  GooDSl'KKD 
ex[)resscd  his  opinion  (in  litt.)  that  the  Rustica  complex  originated  in  the  region 
now  occupied  by  western  ]^oli\  ia  and  l\>ru  and  extended  to  the  "J"^'^  Fernandez 
land",  becoming  isolated  during  the  final  uplift  of  the  Andes  or  possibly  even  earlier. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  209 

See  also  112.  F.  B.  H.  Brown  (jj.  III.  262)  suggested  an  affinity  between  cordifolia 
2ind  fatuhivensis  F.  B.  H.  Brown  (Marquesas),  but  they  belong  to  different  subgenera 
and  the  latter  is  referred  to  N.  fragrans  Hook,  from  N.  Caled.  as  a  variety  {112). 

Scfophulariaceae. 

Mimulus  L.  About  130;  widely  dispersed,  predominantly  western  N.  Amer.  (80 
Calif.),  south    to  S.  Chile;  2  Afr.,  Madag.,  4  E.Asia,  5  Austral.,  Tasm.,  N.  Zeal. 

glabratus  H.  B.  K.  N.  Amer.  to  Boliv.,  Argent,  and  Chile;  polymorphous,  the 
island  form  very  close  to  var.  parviflorus  (Lindl.)  Grant  {114). 
Euphrasia  L.  About  100;    bor.-circump.  with  isolated  populations  in  the  S.  Andes, 
south    to    Fueg.  and  Falkl.,  and  in  Austral.-N.  Zeal.,    the  gap  between  E.  Asia 
and  Australia  bridged  over  by  trop.  mountain  stations;  see  map  y8.22/\. 

formosissima  Skottsb.  The  fact  that  this  species  is  very  unlike  the  Trifidae 
of  Chile  makes  it  particularly  interesting.  In  22g.  169  I  emphasized  the  differ- 
ence between  formosissima  and  the  Australes  of  N.  Zealand  and  placed  it  nearer 
to  the  boreal  Semicalcaratae.  Wettstein,  in  his  contribution  to  my  paper  (I.e.  209), 
expressed  the  opinion  that  it  could  be  attached  to  a  Japanese  group  of  species, 
intermediate  between  Australes  and  Semicalcaratae.  The  question  was  taken  up 
by  Du  RiETZ  who  thinks,  with  good  reason  I  believe,  that  I  overrated  its  rela- 
tions to  boreal  species  (77.  533)  and  that,  in  most  respects,  it  is  more  nearly  related 
to  N.  Zealand  forms. 

Plantaginaceae. 

Plantago  L.  About  270;  world-wide,  essentially  temp. 

fernandezia  Bert.  The  nearest  relative  appears  to  be  P.  priiiceps  Cham,  et 
Schld.  of  Hawaii.  Both  belong  to  sect.  Palaeopsy Ilium,  scattered  over  the  S.  hemisph. 
and  extending  north  to  N.  Amer.,  Hawaii  and  S.  Eur.:  N.  Amer.  5,  S.  Amer.  i, 
S.  Eur.  2,  Afr.  5,  St.  Helena  i,  Madag.  i.  Lord  Howe  I.  i,  Auckl.  Is.  i,  Rapa  2, 
and  Hawaii  9. 

iruncata  Cham.  Centr.  and  S.  Chile.  Pilger  distinguished  the  island  form  as 
a  separate  subspecies  close  to  ssp.  firma  Pilger,  but  I  doubt  that  it  deserves  the 
rank  assigned  to  it,  and  it  is  even  possible  that  it  is  a  very  late  arrival  in  the 
islands. 

Rubiaceae. 

Hedyotis  L.  A  large  pantrop.  genus. 

thesiifolia  St.  Hil.  Trop.-subtrop.  S.  Amer.,  in  Chile  south  to  Chiloe.  Very 
likely  not  truly  indigenous  in  J.   Fern. 

Nertera  Banks  et  Sol.  10-12;  centering  in  N.  Zeal.  (4  endem.  sp.),  north  to  Malaya 
and  Hawaii;  Tristan  da  C;  S.  Amer.  from  Falkl.  and  Fueg.  to  Colomb.  and  Mex. 

granadensis  (L.  fil.)  Druce.   A  polymorphous  species,  reported  from  S.  Amer., 
Tristan  da  C,  Austral.,  Tasm.,  N.  Zeal.,  Java  and  Hawaii — see  24.4,  where  I  pointed 
out  that  it  is  heterogeneous  but  that  the  plant  from  Masafuera  seems  to  be  iden- 
tical with  the  form  common  in  Magell.  and  Falkl. 
Coprosma  Forst.  90;  a  western  Pacific  genus  centering  in  N.  Zeal.  (39)  and  distri- 

14  ~  557857   The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.   Vol.  I 


2IO  C.   SKOTTSBERG 

buted  from  Australia  to  Malays.,  Melan.  and  Polyn.  with  a  secondary  centre  in 
Hawaii  (i8);  entirely  absent  from  Amer. 

Hookeri  (G.  Don)  \V.  R.  B.  Oliver  {1^2).  Forms  a  separate  monotypical  section. 

pyrifolia  (Hook,  et  Arn.)  Skottsb.  Belongs  to  a  section  of  8  Polyn.  sp.  (4 
Tahiti,    i    Cook  Is.,    i    Rapa,    1    Pitcairn,    i  J.  Fern.). 

CaliiDH  L.  World-wide,  essentially  boreal.  Over  500  have  been  described,  almost 
^'4  of  these  Medit.-Orient.;  about  50  in  S.  Amer.,  mostly  along  the  Andes  and 
extending  south  to  Fueg.,  Falkl.   and  S.   Georgia. 

masaftierantim  Skottsb.  Related  to  C  criocarpuin  Bartl.,  DC.  and  tricJiocar- 
pu))i  DC.  (both  Coquimbo-Cord.  Linares),  the  fruit  as  in  masafueraimm  covered 
with  straight  bookless  hairs. 

Campanulaceae. 

Wahlciibergia  Schrad.  l^ssentially  S.  African;  about  230  sp.  (9  S.  Amer.,  10  S. 
Fur.-Oricnt..  2  or  3  St.  Helena,  20  trop.  Afr.,  6  Madag.-Mascar.,  150S.  Afr.,  9  As., 
I    \.  (iuin.,  8  Austral.,  8  N.  Zeal,   5  J.  Fern,). 

Larrainii  (Bert,  ex  Colla)  Skottsb.,  fern andeziana  A.  DC.  p.  p.  and  Grahamae 
Hemsl.  are  closely  related  to  each  other. 

Masafuerae  (Phil.)  Skottsb.  approaches  the  former,  but  has  the  large  tuber 
of  the  following. 

Berteroi  Hook,  et  Arn.  occupies  a  rather  independent  position. 

Most  African  species  are  annuals  and  quite  unlike  the  island  species,  and 
these  have  little  in  common  with  the  single  Chilean  or  the  other  American  forms, 
nor  with  those  from  Australia  or  N.  Zealand,  even  if  there  is  a  superficial  likeness 
between  /['.  Masafuerae  and  a  couple  of  perennial  S.  African  species  such  as 
//'.  lickloiiii  Buek  and  oxypJiylla  A.  DC.  On  the  other  hand,  the  resemblance 
between  the  J.  Fernandez  and  St.  Helena  species  is  quite  striking,  and  in  spite  of 
the  difference  in  the  number  of  carpels,  the  ovary  being  trimerous  in  the  former 
and  dimerous  in  the  latter,  the  possibility  of  a  common  origin  cannot  be  dismissed. 
The  number  of  carpels  in  the  genus  varies  between  2  and  5;  in  most  species 
tiiey  arc  3.  Hkmsi.ev  {i2y.  61)  regarded  the  St.  Helena  species  as  allied  to  African 
and  Juan   I^Y'rnandez  species. 

Lobelia  L.  350-370;  particularly  numerous  in  Amer.  and  Afr.,  less  so  in  As., 
Austral,  and  Oceania,  2   in  Juir. 

alata  Labill.   S.   Chile,  S.  Afr.,  Austral.  A  sea-side  sp. 

Compositae. 

Lagenop flora  Forst.  A  biccntric  austr.-subantarct.  genus  of  16  sp.,  most  numerous 
in   Austral. -N.   Zeal.;   Fiji,   Rapa,  extending  north  to  Philipp.  Is.   and  Hawaii. 

JIario/i  I^Vanch.  Andes  of  S.   Chile  to  Fueg. 
]-j-io;ero}i  L.    A  large    bor.-temp.    genus.    Ind.   Kew.  lists  about  700  sp.  as  valid, 
half  of  them   in   X.   Amer.  and  about   lOO  in   S.  Amer.,  where  the  genus  is  richly 
developed  along  the  Andes,  south  to  Patag.,  Fueg.  and  Falkl.  Not  few  are  known 
from    tro{).    mountains    in    the    W.   and  P2.   hemispheres;    very  few  reported  from 


DERIVATION  OF  THE  FLORA  AND  FAUNA  211 

Australia.  One  species  is  found  on  Rapa,  related  to  the  Juan  Fernandez  species 
according  to  Brown  [jj.  III.  338). 

fruticosus  DC.  and  luteoviridis  Skottsb.  are  related  to  each  other. 

Ingae  Skottsb.,  Innocentium  Skottsb.^  and  turricola  Skottsb.  form  another 
group. 

rupicola  Phil,  stands  apart  from  all  other  species. 

ViERllAPPER   (see   22p.  1 82)   suggested    that  the  island  species  are  allied  to 
Andine  species,  but  also  that  E.  fruticosus  comes  very  close  to  E.  lepidotus  Less. 
of  Hawaii;  this  is,  however,  now  referred  to  Tetramolopiu7n  by  SllERFF.  There  is 
no  Erigerori  in  Hawaii. 
Gnaphalium  L.   A    large  subcosmopol.,  essentially  temp,  genus  needing  revision. 

spicifonne  Sch.  Bip.;  comp.  22g.  187-188.  Patag.  Fueg.  The  assemblage  to 
which  the  alpine  species  of  Masafuera  belongs  is  in  a  state  of  taxonomic  confusion. 
The  identity  with  the  Magellanian  plant  may  be  doubted,  but  I  am  convinced 
that  the  island  form  cannot  be  referred  to  e.\t\\Qr  americanum  W\\\.,  ptirpureum  L., 
spicatuin  Lam.  or  mucronatum  Phil,  which,  together  with  consanguineuin  Gaud.,  are 
regarded  as  forms  of  a  single  polymorphous  taxon. 

Abrotaiiella  Cass.  An  austr.-subantarct.  genus  of  20  sp.,  the  majority  in  N.  Zeal. 
with  the  subantarct.  islands  (9)  and  W.  Patag.-Fueg.-Falkl.  (5);  of  the  remainder 
I   in  Austral.,  2  Tasm.,   i   N.  Guin.,   i   Rodriguez  I.,  and  the  following. 

crassipes  Skottsb.  Very  near  A.  Moseleyi  Skottsb.  nom.  (W.  Patag.);  see 
22g.  189-190. 

Robinsonia  DC.  Bentiiam  (20.460)  remarks  on  Robinsonia  2iW^  Rhetinodendroni 
"Although  their  connexion  with  Eusenecioneae  seems  greater  than  with  any  other 
subtribe  or  tribe,  yet  in  their  dioecious  capitula,  in  the  presence  of  small  free 
anthers  without  pollen  in  the  female  floret  and  some  other  points  they  approach 
the  subtribe  Petrobieae  of  Helianthoideae."  The  idea  was  rather  attractive,  be- 
cause the  Petrobieae  inhabit  St.  Helena  and  S.  America.  Another  suggestion  men- 
tioned the  Hawaiian  Dubautia  and  Raillardia,  but  they  are  not  dioecious.  Re- 
cently an  undisputable  relative  was  discovered  in  New  Guinea,  Brachionosiylunt 
Mattfeld  (j/c?.  27-28) : 

Von  Senecio,  dessen  pacifische  Arten  unserer  Pflanze  recht  nahe  kommen,  unter- 
scheidet  sich  diese  Gattiing  wesentlich  durch  die  Zweihausigkeit  .  .  .  und  waiter  in 
Zusammenhang  damit  durch  die  Form  der  Griffelschenkel  der  Scheibenbliiten,  die 
der  Fegehaare  ganz  entbehren  und  vorn  nicht  gestutzt  sondern  abgerundet  sind.  Diese 
Merkmale  hat  sie  aber  mit  den  auf  Juan  Fernandez  endemischen  Gattungen  Robinsonia 
und  Rhetinodendron  gemeinsam,  von  denen  sie  sich  iiberhaupt  durch  kein  generisches 
Merkmal  unterscheidet.  Aber  die  Ubereinstimmung  erstreckt  sich  sogar  auf  kleinere 
Merkmale,  wie  den  leicht  hinfalligen  Pappus,  die  kurzen,  aber  verhaltnismiissig  sehr 
breiten,  etvvas  vorspringend  gestreiften  Zungen,  die  Form  der  durch  ein  kleines  Be- 
cherchen  gekronten  Achaenen,  und  schliesslich  die  Wuchsform  und  Verzweigungsart : 
bei  beiden  schliesst  der  schopfig  beblatterte  Stengel  mit  dem  Bliitenstande  ab,  wahrend 
der  Fortsetzungsspross  aus  der  Achsel  eines  der  oberen  Laubblatter  unter  dem  Blii- 
tenstande entspringt  und  diesen  bald  zur  Seite  driingt.  ...  So  bleibt  als  einziger  Un- 
terschied  der  durch  die  sehr  verschiedene  Blattform  bedingte  habituelle  Findruck.  .  .  . 

1  Described  as  a  variety  of  the  former  and  now  raised  to  specific  rank. 


212  C.   SKOTTSBERG 

Die  Rliitcnkopfe  sind  bei  Robinsonia  erheblich  zahlreicher  und  kleiner,  kurz  und  breit 
glockenformig,   bei  unserer  Pflanze  aber  schmalglockig. 

Wiirden  diese  Sippen  demselben  pflanzengeographischen  Gebiete  angehoren,  so 
wiirde  man  sie  sicher  nicht  generisch  trennen  konnen.  Was  hier  aber  wesentlich  dazu 
zwingt,  ist,  dass  die  Zweihiiiisigkeit,  die  ja  hier  die  wesentliche  Ubereinstimmung  be- 
dingt,  bei  den  Kompositen  zu  den  verschiedensten  Malen  in  den  verschiedensten  und 
auch  in  denselben  (iru})pen  entvvickelt  wiirde  und  daher  keineswegs  als  Kriterium 
fur  eine  generische  Verwandtschaft  angesehen  werden  kann. 

To  this  I  shall  remark  that  nobody  would  think  of  uniting  Rohhisonia  and 
Rlicti}wdeudron,  because  they  differ  in  essential  floral  characters.  Brachionostylmn 
is,  to  judge  from  the  description  and  plate  XC:  B  (only  ?  known),  closely  related 
to  Robi)iso7iia  but  not  to  RJietinodendron.  The  mode  of  growth  is  not  quite  the 
same,  for  in  Robinsonia  (as  well  as  in  Rheti7iodendroii)  2  or  3  innovations  are 
developed,  and  none  of  them  continues  the  mother  axis.  Besides,  Brachionosiylimi 
has  petiolate,  penninerved  leaves  of  a  type  current  in  dicotyledons  and  widely 
different  from  the  sessile,  linear-lanceolate,  broadly  clasping  and  parallel-veined 
leaves  of  Robinsonia. 

gayana  Dene  and  thurifera  Dene  are  a  pair  of  related  species. 

evenia  Phil,  and  Masafuerae  Skottsb.  form  another  pair. 

gracilis  Dene  stands  more  isolated. 
Symphyochaeta  (DC.)  Skottsb.   See  2^g.  785. 

macrocephala  (Dene)  Skottsb. 
Rhetinodendron  Aleisn.  Berterii  (Dene)  Hemsl.   See  above. 

Centaurodendron    dracaenoides  Johow.  Not,  as  has  been  supposed,  just  an  ar- 
boreous  Centaurca,  but  differing  materially  in   flower  structure  [24J). 
Yunqiiea   Tenzii  Skottsb.   See  2jy.  163. 

Dendroseris  C.  Don.  An  isolated  genus,  perhaps  distantly  related  to  TJiavmo- 
sevis  Phil,  from  Desventuradas  Is.,  but  probably  not  to  Fiichia  Hook.  fil.  Ben- 
lllAM  stated  (20.480)  that  the  achenes  of  Dendroseris  differ  from  those  of  the 
Cichorioideae  in  general,  and  that  also  the  involucre  and  the  habit  are  different. 
With  regard  to  the  achenes  this  is  certainly  true  of  Dendroseris  s.  str.,  and  also 
of  PJioenicoscris,  whereas  they  are  of  a  rather  normal  type  in  Rea  and  Hespe- 
roscris.  All  are  devoid  of  paleae,  which  are  present  in  Tlianinoseris  and  Fitchia, 
a  distinctive  feature  pointed  out  by  1^p:ntiiam,  but  the  awned  achenes  of /7/f///^ 
differ  from  those  of  all  the  other  genera  mentioned.  In  spite  of  the  profound 
discrepances  in  sucli  im[)ortant  characteristics  as  inflorescence,  involucre,  recep- 
tacle and  stigma  Tlianinoseris  has  more  in  common  with  the  Dendroseris  as- 
semblage than   with   other  genera. 

macrophylla  \).  Don,  macrantha  (Bert,  ex  Dene)  Skottsb.,  margitiata  (Bert, 
ex   Dene)   I  look,   et  y\rn.  and  litoralis  Skottsb.,  all  lumped  together  by  JOIIOW,  are 
well-marked  s{)ecics;   comp.   22^1.201-204. 
Phoenicoseris   Skottsb.,  2./9.  787. 

pinnata  (Bert,  ex  Dene)  Skottsb.  and  fter^er/ana  (Dene)  Skottsb.  (Masatierra) 
are  closely  related  but  quite  distinct;  regia  Skottsb.  (Masafuera)  differs  from  both 
in  leaf  shape,  but  fertile  specimens  have  not  been  found. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  213 

Rea  Bert,  ex  Dene  p.  p.  See  24^.  788. 

neriifolia  Dene.  Little  is  known  of  this,  but  it  is  undoubtedly  a  very  good 
species;   micrantha  Bert,  ex  Dene  and  pruinata  (Johow)  Skottsb.  are  near  rela- 
tives but  good  species  (comp.  22g.  207). 
Hesperoseris  Skottsb.  gigantea  (Johow)  Skottsb.  See  24p.  788. 

Vicarious  species. 

VlERHAPPER  (2yd)  distinguished  between  true  and  false  vicarism.  True  vi- 
carists  have  arisen  from  a  common  initial  species  and  become  differentiated  either 
within  the  limits  of  the  area  this  once  occupied  or  after  penetration  into  a 
different  habitat,  followed  by  isolation,  whereas  in  the  case  of  pseudo-vicarism 
they  have  a  different  origin;  a  second  species  may  invade  the  area  of  the  first 
and  colonize  such  parts  of  the  area  as  are  unsuitable  to  the  latter.  Very  often 
the  term  "vicarious"  has  been  taken  in  a  much  wider  sense:  any  two  related 
species  replacing  each  other  in  separate  areas  were  called  vicarious,  and  phyto- 
geographers  used  the  term  to  designate  two  plant  species,  related  or  not,  that 
played  corresponding  roles  in  two  closely  allied  plant  communities;  in  this  case 
Nothomyrcia  fernandeziana  and  Myrceuge7iia  Schulzei,  which  form  the  bulk  ot 
the  forest  in  Masatierra  and  Masafuera,  respectively,  are  vicarists,  although  they 
belong  to  different  genera.  Species  not  fulfilling  the  conditions  claimed  by  VlER- 
HAPPER were  called  "substitute  species". 

WULFF  [2gi.  66-62)  devotes  considerable  space  to  a  discussion  of  vicarism. 
He  agrees  with  VlERHAPPER:  true  vicarism  is  a  result  of  one  taxon  breaking 
up  into  two,  adapted  to  different  habitat  conditions.  Cain  (42.  265)  expresses 
himself  in  slightly  different  words,  but  their  meaning  is  the  same:  "closely  re- 
lated allopatic  species  which  have  descended  from  a  common  ancestral  popula- 
tion and  attained  at  least  spatial  isolation."  He  summarizes  statements  made 
by  Drude,  Diels,  Wulff  and  Setchell,  who  called  species  vicarious  if  they 
were  only  slightly  discontinuous  morphologically  but  widely  so  geographically; 
certainly  anybody  would  call  them  vicarists  but  use  the  same  term  for  a  pair 
of  intimately  related  forms  of  which  one  inhabits  granite  and  the  other  limestone 
within  the  same  geographical  area. 

I  am  afraid  that,  in  most  cases,  we  know  very  little  or  nothing  at  all  of 
the  ancestry  of  species  we  are  used  to  call  vicarious,  let  it  be  that  we  have 
reason  to  assume  that  they  have  differentiated  out  of  a  common  population  some 
time  in  the  past.  This  is,  at  least,  the  situation  in  Juan  Fernandez.  Here  I 
should  perhaps  refer  to  what  Cain  (I.e.  276)  calls  polytopic  species,  i.e.  when 
the  same  taxon  occurs  in  two  or  more  discrete  areas,  disjunction  being  the 
result  either  of  dispersal  from  one  original  centre  or  of  the  breaking  up  of  an 
area  through  subsidence  or  formation  of  some  other  kind  of  barrier ;  upon  a  close 
investigation  it  has  been  shown  in  many  instances  that  slightly  different  forms 
within  the  original  population  happened  to  become  isolated  and  appear  as  ex- 
amples of  true  vicarism. 

Johow  {ijo.  233)  held  forth  that  6  species  endemic  in  Masatierra  correspond 
to  6  other  species  endemic  in  Masafuera,  but  he  did  not  use  the  term  vicarious. 


214  ^'  SKOTTSBERG 

These  were  (the  species  from  Masatierra  mentioned  first):  Dendroseris  niicrantha- 
:gigantca  (belong  to  different  genera),  WaJiloibergia  Bcrteroi-tuberosa  {=  Masa- 
fuerae,  not  very  closely  related),  JMyrceugcnia  fernandezia7ia-Sc)iulzei  {^\^^x^x\\. 
genera),  liryngium  buplcuyoides-sarcopJiyllujn  (only  distantly  related),  Cardamiue 
alsophila  ( =  Jlaccida) -  Kruessi'/ii{v\ot  nearly  related),  and  Urtica glo^neruliflora ( =  fer- 
7iandt'zia)ia)~Masafucrae  (belong  to  different  sections).  There  is  not  among  them 
a  single  example  of  either  vicarism  or  pseudo-vicarism  in  the  sense  of  ViERllArPER. 
In  1914  (-?-?/)  I  distinguished  4  pairs  of  vicarists  in  Juan  Fernandez: /^<f;/^/r^- 
stris  inicraiitha-gigajitea,  Myrceugenia  fcruandeziana-Schulzei,  Giinnera peltata- 
Masafuerac,  and  Pepcrouiia  luargaritifera-Skottsbergii;  the  flora  of  Masafuera  was 
at  that  time  little  known.  Of  these  pairs  the  last  two  still  hold  good,  the  species 
replace  each  other  from  a  taxonomical  as  well  as  from  a  sociological  viewpoint; 
nevertheless  I  am  not  prepared  to  argue  that  they  arose  out  of  one  initial  spe- 
cies: several  species  may  have  been  involved,  for  they  differ  in  a  number  of 
minor  characters  and  all  we  can  say  is  that  they  make  the  impression  of  coming 
from  the  same  stock.  On  the  basis  of  our  present  knowledge  of  their  taxonomy 
the  following  nine  pairs  are  distinguished;  the  species  replace  each  other,  call 
them  \icarious  or  substitute:  Berberis  corymbosa-masafjierana,  CJienop odium  crii- 
soeanum-nesodendron,  Dendroseris  macrayitha-macrophylla,  F agar  a  mayu— externa, 
Gunner  a  peltata- Masafuerae,  Halorrhagis  masatierrana-masafuerana,  Peperomia 
inargaritifera-Skottsbergii,  Robinsonia  eve^iia  —  Masafuerae,  and  Sophora  ferna7i- 
deziajia- masafuerana.  As  regards  Berberis,  Fagara,  Halorrhagis  and  Sophora, 
perhaps  also  Gumiera,  Peperomia  and  Robinsonia,  the  two  members  of  a  pair  are 
closely  related,  but  in  the  case  of  Halorrhagis  the  situation  is  complicated  be- 
cause there  is  a  third  species,  endemic  in  Masafuera,  H.  aspe^^rima;  masaftiera7ia 
seems,  however,  to  be  a  better  match  for  masatierrana  than  asperri7na.  As  we 
have  seen,  there  is  also  a  third  species  of  Che7iop odium,  endemic  to  Santa  Clara, 
very  likely  formerly  occurring  also  on  Masatierra,  but  extinct  there.  Of  the  3 
species  of  Dendroseris  inhabiting  Masatierra,  7narginaia  and  litoralis  form  a  pair 
of  one  inland  and  one  coast  species. 

An  unbalanced  flora. 

It  has  always  been  argued  that  an  island  flora,  where  the  number  of  species 
is  small  compared  with  the  number  of  genera  and  many  large  and  widely  distri- 
buted families,  well  developed  under  different  conditions,  poorly  represented  or 
altogether  absent,  furnishes  one  of  the  best  proofs  of  the  absolute  oceanity  of  its 
abode,  and  Juan  Fernandez  is  no  exception  from  this  rule.  I  have  mentioned  this 
above  (|).  194)  pointing  out  that  most  of  these  families  have  numerous  genera 
and  sj)ecies  on  the  opi)osite  mainland,  thus  Caryophyllaceae  20  genera,  Compo- 
.sitae  some  130,  Cruciferae  28,  Leguminosae  22,  Scrophulariaceae  18,  Umbelliferae 
about  30,  and  so  forth,  and  examples  of  important  families  in  Chile  not  found 
in  the  islands  were  also  given.  This  state  of  things  calls  for  an  explanation.  JOIIOW 
paid  much  attention  to  it;  as  he  believed  that  the  islands  had  been  isolated  from 
the  very  beginning  he  blamed  chance  in  all  cases  where,  according  to  the  cur- 
rent opinion,  the  diaspores  were  adapted  for  dispersal  across  a  wide  expanse  of 


DERIVATION  OF  THE  FLORA  AND  FAUNA  215 

water.  He  mentions  Compositae-Labiatiflorae,  Calyceraceae,  Valerianaceae,  Cacta- 
ceae,  Orchidaceae  and  Dioscoreaceae  as  good  examples.  In  other  cases  he  blames 
the  diaspores  as  in  Nolanaceae,  Leguminosae,  Violaceae,  Fagaceae,  Amaryllidaceae 
and  Liliaceae. 

Also  with  reference  to  life-forms  the  island  flora  makes  an  impression  of  being 
unbalanced,  a  fact  already  discussed  at  some  length  in  2^1.  825-830;  I  shall 
only  repeat  that  woody  species  are  in  overwhelming  majority  and  annuals  almost 
absent,  whereas  they  are  very  numerous,  also  proportionately,  on  the  mainland. 
Already  in  19 14  [22^]  I  argued  that  climatic  differences  alone  do  not  offer  an 
explanation  and  that  historical  causes  must  be  taken  into  account. 

Geographical  elements. 

In  his  survey  of  the  flora  p.  229-232  JOIIOW  discussed  its  composition  and 
distinguished  various  elements.  The  endemic  genera  and  a  few  peculiar  endemic 
species  of  non-endemic  genera  form  his  first  (and  most  ancient)  group;  the  second 
group  contains  the  remainder  of  endemic  species:  (a)  markedly  distinct,  (b)  nearly 
related  to  continental  species.  Both  groups  make  up  "Continjente  A".  The  third 
group,  "Continjente  B",  contains  the  species  found  elsewhere,  all  occurring  in  South 
America  except  Halorrhagis  ''alata\  America  is  claimed  as  the  source  of  the  en- 
demic element  as  well:  "tanto  las  especies  del  primero  como  las  del  segundo  grupo 
pertenecen,  con  la  unica  excepcion  del  Santalum,  a  jeneros  representados,  si  no 
en  Chile,  a  lo  menos  en  alguna  parte  de  la  costa  occidental  de  Sud-America" — 
this  did  not,  however,  apply  to  the  isolated  Compositae,  to  Lactoris  etc.,  which 
JOHOW  regarded  as  originated  from  ancestors  to  be  looked  for  in  the  Tertiary 
flora  of  Chile.  Halorrhagis,  Santalum  and  Coprosma,  mentioned  later,  were  sup- 
posed to  have  immigrated  from  western  Pacific. 

It  serves  no  purpose  to  go  into  detail;  when  JOHOW  wrote  his  book  our 
knowledge  of  the  flora  was  too  defective  to  allow  him  to  arrive  at  anything  like 
safe  conclusions.  The  same  may  be  said  of  my  1914  paper  (22^),  even  if  my  short 
visit  to  the  islands  revealed  the  existence  of  an  up  till  then  unknown  element. 
Four  main  groups  were  distinguished:  I.  Old  Pacific  (Engler's  Altoceanisches 
Element),  comprising  genera  or  species  supposed  to  have  a  long  history  behind 
them  within  the  precincts  of  the  Pacific  and  lacking  near  relatives;  subdivision 
A,  Endemic  genera,  with  11  sp.,  and  Non-endemic  genera  with  10  sp.;  subdivision 
B  with  allied  species  in  Hawaii,  Polynesia,  Australia  and  New  Zealand,  16  en- 
demic species — the  genus  Dendroseris  was  also  placed  here — and  i  non-endemic 
{Halorrhagis).  Group  II,  called  Neotropical,  contained  6  endemic  species;  Cuminia 
and  Juania  were  included  here.  Group  III,  called  Chilean,  was  the  largest  and  was 
divided  into  3  lots:  A,  very  distinct  species,  18  [Ochagavia  placed  here);  B,  less 
well-marked  species,  1 1 ;  C,  also  found  in  Chile,  28,  but  among  them  were  6  not 
now  regarded  as  native.  So  far  the  main  difference  between  this  arrangement  and 
JOllOW's  lies  in  the  greater  number  of  species  with  supposed  west  Pacific  connec- 
tions. Finally  Group  IV,  Subantarctic-Magellanian,  4  species,  was  added. 

The  study  of  our  1916-1917  collection  added  many  species  not  known  before 
and    gave    rise    to    a    fresh    analysis    of  the    vascular   plants   [2jg),    of  which    a 


2l6  C.   SKOTTSBERG 

short  summary  is  given  here  (translated  from  French).  The  ferns  are  not  included 
below. 

1.  Neotropical  and  Andean  element.  Species  either  found  in  Chile  (with  one  exception,  c) 

or  endemic  but  allied  to  Chilean  si)ecies;  47. 

a)  in  Central  or  S.  Chile,  20. 

b)  in  the  Magellan  region,  but  not  of  subantarctic  character,  3. 

c)  in  South  America,  but  not  in  Chile,  i. 

d)  endemic  species  of  non-endemic  genera,  21. 

e)  ,,  ,,       ,,  endemic  ,,      ,  2. 

2.  Element  consisting  of  wide-spread  species,  also  inhabiting  Chile,  5. 

3.  Neotropical  element,  not  represented  in  Chile,  7. 

a)  endemic  species  of  non-endemic  genera,  3. 

b)  species  belonging  to  endemic  genera,  4. 

4.  Magellanian  -  Old  Antarctic  element,  27. 

a)  subantarctic  species,  10. 

b)  austral  species,  5. 

c)  endemic  species  allied  to  subantarctic  species,  7. 

d)  endemic  s])ecies  not  allied  to  subantarctic  species,  5. 

5.  Pacific  element,  59. 

a)  isolated  endemic  species  of  wide  genera,  21. 

b)  endemic  species  belonging  to  genera  or  sections  of  west  to  central  Pacific  distribu- 
tion or,  if  belonging  to  wide  genera,  then  more  closely  related  to  Pacific  species,  i  7 . 

c)  species  of  isolated  endemic  genera  with  supposed  transpacific  relations,  18, 

d)  sj)ecies  of  endemic  genera  with  unknown  relations,  3. 

This  analysis  was  based  on  my  memoir  on  the  Phanerogams  published  in 
1922  (22^).  Since  that  time  several  pending  questions  have  been  restudied,  additional 
plant  material  has  come  to  hand,  and  recent  monographs  have  been  consulted. 
The  results  were  communicated  above,  and  we  shall  now  proceed  to  the  following 
arrangement  of  the  angiosperms. 

I.  Andine-Chilean  element. —  69  (46.9  %). 

a.  Endemic  species  (34).  In  two  cases,  OcJiagavia  and  NotJiomyrcia,  also  the 
genus  endemic,  but  nearly  related  to  Chilean  genera.  Chaetotropis  imberbis,  Chus- 
quea  fernandeziana,  Uncinia  Douglasii  and  costata,  Hesperogreigia  Berteroi,  Ocha- 
gavia  elegans,  Luzula  masafuerana,  Phrygilanthus  Berteroi,  Spergularia  masafue- 
rana,  Cardamine  Kruesselii,  P^scallonia  Callcottiae,  Margyricarpus  digynus,  Sophora 
fernandeziana  and  masafuerana,  Dysopsis  hirsuta,  Colletia  spartioides,  Azara  fer- 
nandeziana, Nothomyrcia  fernandeziana,  Myrceugenia  Schulzei,  Gunnera  peltata 
and  Masafuerae,  Apium  fernandezianum,  Pernettya  rigida,  Rhaphithamnus  venus- 
tus,  Solanum  fernandezianum,  masafueranum  and  robinsonianum,  Galium  masa- 
fueranum,  P>igeron  fruticosus,  luteoviridis,  Ingae,  Innocentium,  turricola  and 
rupicola. 

b.  Known  from  continental  Chile,  many  also  in  other  parts  of  S.  America  or 
of  still  wider  distribution  (33):  Stipa  neesiana  and  laevissima,  Piptochaetium  bicolor, 
Chaetotropis  chilensis,  Trisetum  chromostachyum,  Danthoniacollina,  Koeleria  micra- 
thera,  C"\-perus  eragrostis  and  reflexus,  Scirpus  cernuus  and  nodosus,  Eleocharis 
fuscopurpurea,  Uncinia  phleoides,  Juncus  imbricatus,  capillaceus,  procerus,  dom- 
beyanus  and  planifolius,  Libertia  formosa,  Salicornia  fruticosa,  Tetragonia  expansa. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  21  7 

Peperomia  fernandeziana,  Parietaria  humifusa,  Paronychia  chilensis,  Cardamine 
flaccida,  Callitriche  Lechleri,  Centella  triflora,  Dichondra  repens,  Calystegia  tugu- 
riorum,  Mimulus  glabratus,  Plantago  truncata,  Hedyotis  thesiifolia,  Lobelia  alata. 

c.  Also  known  from  San  Ambrosio:  Spergularia  confertiflora. 

d.  S.  American,  but  not  reported  from  Chile:  Cardamine  chenopodiifolia. 

II.  Subantarctic-Magellanian  element. — 15  (10.2  %). 

a.  Endemic  (4):  Agrostis  masafuerana,  Drimys  confertifolia,  Acaena  masafue- 
rana,  Abrotanella  crassipes. 

b.  Not  endemic,  in  several  cases  extending  north  along  the  Andes  (11):  Oreo- 
bolus  obtusangulus,  Uncinia  brevicauli?  and  tenuis,  Carex  Banksii,  Rubus  geoides, 
Acaena  ovalifolia,  Myrteola  nummularia,  Empetrum  rubrum,  Nertera  granadensis, 
Lagenophora  Harioti,  Gnaphalium  spiciforme. 

III.  Neotropical  element. — 19  (12.9  %). 

a.  Belonging  to  endemic  genera  without  or  with  only  distant  relations  to  the 
present  Andean  flora  but  presumably  of  neotropical  ancestry  (5):  Megalachne 
berteroniana  and  masafuerana,  Podophorus  bromoides  (only  tentatively  referred  here), 
Centaurodendron  dracaenoides,  Yunquea  Tenzii. 

b.  Belonging  to  endemic  genera  of  undoubted  neotropical  affinity  (2):  Juania 
australis,  Selkirkia  Berteroi. 

c.  Endemic  species  belonging  to  widespread  genera  and  presumably  of  neo- 
tropical ancestry  (7):  Urtica  fernandeziana,  Chenopodium  Sanctae  Clarae,  crusoe- 
anum    and    nesodendron,    Eryngium  bupleuroides,  inaccessum   and  sarcophyllum. 

d.  Endemic  species  of  unquestionably  neotropical  parentage  (5):  Urtica  Masa- 
fuerae,  Berberis  corymbosa  and  masafuerana,  Ugni  Selkirkii,  Nicotiana  cordifolia. 

IV.  Pacific  element. — 26  (17.7  %). 

Affinities  along  the  route  New  Zealand-Australia— Melanesia-Polynesia-Hawaii; 
all  endemic, 

a.  The  genus  endemic  (9):  Cuminia  fernandezia  and  eriantha,  Robinsonia 
gayana,  thurifera,  evenia,  Masafuerae  and  gracilis,  Symphyochaeta  macrocephala, 
Rhetinodendron  Berterii. 

b.  Only  the  species  endemic  (17):  Cladium  scirpoideum,  Carex  berteroniana, 
Peperomia  margaritifera  and  Skottsbergii,  Boehmeria  excelsa,  Santalum  fernande- 
zianum,  Ranunculus  caprarum,  Fagara  mayu  and  externa,  Gunnera  bracteata, 
Halorrhagis  asperrima,  masatierrana  and  masafuerana,  Euphrasia  formosissima, 
Plantago  fernandezia,   Coprosma  Hookeri  and  pyrifolia. 

V.  Atlantic  -  S.  African  element. — 6  (4.1  %).  Endemic. 

Peperomia  berteroana,  Wahlenbergia  Larrainii,  fernandeziana,  Grahamae,  Masa- 
fuerae and  Berteroi. 

VI.  Eu-Fernandezian  element.  —  i  2  (8.2  %). 

Isolated  endemics  of  unknown  parentage,  the  Cicoriaceous  genera  forming  a 
natural  group. 


2l8  C.  SKOTTSBERG 

Lactoris  fernandeziana,  Dendroseris  macrophylla,  macrantha,  marginata  and 
litoralis,  Phoenicoseris  pinnata,  berteriana  and  regia,  Rea  neriifolia,  micrantha  and 
pruinata,   Hesperoseris  gigantea. 

The  ditierences  between  this  arrangement  and  the  one  of  1934  are  consider- 
able, but  partly  at  least  more  apparent  than  real.  Group  i  of  1934  corresponds 
(if  we  exclude  (b),  which  from  a  purely  geographical  point  of  view  has  to  go  to 
the  present  group  II)  to  I,  but  Selkirkia  has  now  been  placed  in  III,  a  group 
correspontling  to  the  3d  element  of  1934.  Group  4  was  rather  heterogeneous  and 
included,  beside  Magellanian  species,  several  bicentric  ones  and  some  endemics 
of  austral-circumpolar  affinity  now  referred  to  I  a.  Group  5,  Pacific  element,  in- 
cluded the  new  groups  IV-VI  and  part  of  III. 

II.  Pteridophyta. 

Six  families  (taken  in  the  old  sense)  are  represented,  Ophioglossaceae  only 
on  Masatierra,  Lycopodiaceae  only  on  Masafuera.  The  number  of  genera  is  23; 
21  (91.3%)  are  found  on  Masatierra  and  the  same  number  on  Masafuera,  3  on 
Santa  Clara.  Two  genera  are  confined  to  Masatierra  (8.7%)  and  2  to  Masafuera, 
19  (82.6%)  shared  by  both  islands,  3  of  them  also  known  from  Santa  Clara.  A 
single  genus  [Thyrsoptcris]  is  endemic  in  Juan  P'ernandez  and  found  on  Masatierra 
and   Masafuera. 

Of  the  53  s{)ecies  listed  43  occur  on  Masatierra,  3  on  Santa  Clara  and  45 
on  Masafuera.  Only  8  species  are  restricted  to  Masatierra,  15.1%  (18.6%  of  the 
total  found  there)  and  10  to  Masafuera,  18.9%  (22.2%  of  the  total);  35  [66%) 
are  found  on  both. 

There  are  18  endemic  species  (34%),  of  which  16  inhabit  Masatierra  and  14 
Masafuera;  no  endemic  is  found  on  Santa  Clara.  Of  the  endemics  4  are  confined 
to  Masatierra  and  2  to  Masafuera,  12  having  been  recorded  from  the  two  islands; 
in  percentage:   22.2,    ii.i   and  (^^.'jX,  respectively. 

A  high  percentage  of  ferns  is  to  be  expected  in  "oceanic"  islands,  whether 
or  not  isolated  from  the  beginning,  and  Juan  Fernandez  is  no  exception  from  this 
rule,   for  of  the   200  vascular  plants  26.5  %   are  Pteridophytes. 

Of  the  35  species  also  found  elsewhere  4  are  confined  to  Masatierra,  8  to 
Masafuera  and  23  found  on  both  islands,  3  of  these  also  on  Santa  Clara;  in  per- 
cctUage  11.4,  22.9  and  65.7,  respectively.  The  difi"erences  in  percentage  between 
endemic  and  non-endemic  species  is  due  to  the  occurrence,  in  the  highland  of 
.Masafuera,  of  a  few  mountain  ferns  not  found  on  the  other  island.  But  whether 
our  analysis  bears  upon  families,  genera  or  species  the  distribution  over  the  archi- 
pelago is  \ery  much  more  even  than  the  dispersion  of  the  angiosperms,  and  this 
is  of  course  what  we  have  every  reason  to  expect.  Also,  the  number  of  species 
to  a  genus  is  greater,   2.3. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  219 

The   systematical   position   of  the   endemites   and  the  geographical  distribution 
of  the  genera  and  species  also  found  elsewhere. 

Hymenophyllaceae. 

Trichomanes  L.  coll.  The  island  species  belong  to  Vandenhoschia  Copel.,  a  pan- 
trop.  and  circum-austr.  genus  of  about  25  sp.,  extending  north  to  N,  Amer.,  Engl, 
and  Japan. 

philippianum  Sturm.  "One  of  the  most  distinct  species  of  Ti'ichomanes  . .  . 
its  cellular  structure  quite  unique"  (CllRlSTENSEN  ^^.3).  —  Also  in  S.  Chile?  (see 
I.e.  2). 

Ingae  C.   Chr.  Belongs  to  the  neotrop.  pyxidiferum  group. 

exsectum  Kze.  S.  Chile,  south  to  Chiloe.  Near  the  neotrop.  T.  tener'utn  Kze. 
Serpyllopsis  v.  d.  Bosch.  A  monotypical  genus  without  near  relatives  (Copeland 

^9'  37). 

caespiiosa  (Gaud.)  C.  Chr.  (var.  fernandeziana  C.  Chr.  et  Skottsb.,  slightly 
different  from  the  forms  described  from  the  mainland).  Falkl.  and  Fueg.  to  S.  Chile. 
Hymenoglossum  Presl.  Monotypical  and  without  near  affinity  to  any  known  genus 
(Copeland  I.e.). 

cruentum  (Cav.)  Presl.  S.  Chile,  Valdivia  to  49°  s.  lat. 
Hymenophyllum  Sm.  coll. 

a.  Mecodium  Presl.  About  100,  pantrop.  and  austral,  with  several  sp.  in  N. 
Zeal.;  north  to  Sakhalin. 

cuneaUwt  Kze.  S.  Chile,  Valdiv.-W.  Patag.;  Marquesas  Is.,  Rapa.  Near 
H.  polyanthes  Sw.,  a  variable  pantrop.  sp. 

caudiculatum  Mart.  Peru,  S.  Braz.,  S.  Chile,  Valdiv.-49°  s.  lat. 

fuciforme  Sw.  S.  Chile  to  W.  Patag.  A  very  distinct  sp.,  referred  to  Meco- 
dium with  considerable  doubt  (CoPELAND  ^<5.  95). 

b.  HyfnenophyllMH  s.  str.  About  25,  austr.-circump.  with  outlying  stations  in 
W.  Eur.  and  Japan;  H.  peltatum  (Poir.)  Desv.  widely  scattered  in  slightly  different 
forms. 

pectinatum  Cav.  S.  Chile,  Valdiv.-Fueg.  An  aberrant  species,  CoPELAND  6g.  34. 

falklandicmn  Bak.  W.  Patag.,  P'ueg.,  Falkl.,  S.Georgia.  Related  to /'^//^?///w 
which,  \{  falklandicum  really  belongs  to  Mecodium,  where  CoPELAND  placed  it 
-<5^.  94,  also  must  be  brought  here;  peltatum  is  known  from  extratropical  regions 
of  both  hemispheres  and  also  reported  from  Chile  (perhaps  =  var.  Mejiziesii  '(^x^^X) 
C.  Chr.  61. 4),  by  Ciiristensen  also  quoted  for  Juan  Fernandez — he  must  have 
forgotten  that  in  62.  1 1   all  material  from  there  was  referred  to  falklandicum. 

rugosum  C.  Chr.  et  Skottsb.  Related  to  //.  tunbridgerise  (L.)  Sm.,  W.  Ind., 
Venez.,  Chile,  Atl.  islands,  Mediterr.,  S.  Afr.,  Austral.,  Tasm.,  N.  Zeal. 

c.  Sphaerocioyiium  Copel.  A  pantrop.  and  austral  genus  of  about  50  sp.,  richest 
developed  in  trop.  Amer. 

ferrugineum  Colla.  S.  Chile,  Valdiv.-Fueg.  Related  to  the  pantropical  H.  ci- 
liatum  Sw.  and  belonging  to  a  small  austral-tricentric  group  of  closely  allied  spe- 
cies, H.  Frankliiiiae  Col.  (N.  Zeal,  probably  =  ferrugineum,  CllRlSTENSEN  61.  5), 


C.  SKOTTSBERG 


Table  III. 

List  of  species. 
Endemics  bold  faced,  endemic  genera  in  capital  letters. 


Adiantum  chilense  Kaiilf 

Arthropteris  altescandens  (Colla)  J.  Sm, 
Asplenium  dareoidcs  Desv 

—  obliquum  Forst.  var 

—  macrosorum  Bert,  ex  Colla      .    .    . 

—  stellatum  Colla 

Blcchiinm  auriculatum  Cav 

—  chilense  (Kaulf.)  IVIett 

—  cycadifolium  (Colla)  Sturm  .... 

—  longicauda  C.  Chr 

—  Schottii  (Colla)  C.  Chr 

—  valdiviense  C.  Chr 

Cystopteris  fragilis  (L.)  Bernh 

Dicksonia  berteriana  (Colla)  Hook.    .    . 

—  externa  Skottsb 

Dryopteris  inaequalifolia  (Colla)  C.  Chr. 
Elaphoglossum  Lindenii  (Bory)  Moore  . 
(ileichenia  cf.  litoralis  (Phil.)  C.  Chr.  .    . 

—  pedalis  (Kaulf.)  Spr 

—  quadripartita  (Poir.)  Moore     .... 
Histio[)teris  incisa  (Thunb.)  J.  Sm.      .     . 
Hymenoglossum  cruentum  (Cav.)  Presl 
Hymenophyllum  caudiculatum  Mart.      . 

—  cuneatum  Kze 

falklandicum  Bak 

—  ferrugineum  Colla 

—  fuci forme  S\v 

—  pectinatum  Cav 

—  plicatum  Kaulf 

-  rugosum  C.  Chi.  et  Skottsb.     .    .    . 

— ■  secundum  Hook,  et  Cirev 

—  tortuosum  Hook,  ct  Crev 

Hypolepis  ru.t^osuia  (Labill.)  J.  Sm.     .     . 
Lophosoria  (|uadripinnata  ((Imcl.)  C.  Chr. 

Lycopodium  ma.uellanicum  S\v 

— •  scariosum  l-'orst 

Ophioglossum  fernandezianum  C.  Chr. 

Pellaea  chilensis  I'ee 

Polypodium  magellanicum  (Desv.)       .     . 


Mt 


SC 


DERIVATION  OF  THE  FLORA  AND  FAUNA 


—  intermedium  Colla 

—  lanceolatum  L 

—  Masafuerae  Phil 

—  trichomanoides  Sw 

Polystichum  berterianum  (Colla)  C.  Chr, 

—  vestitum  (Forst.)  Presl 

Pteris  berteroana  Ag 

—  chilensis  Desv 

—  semiadnata  Phil 

Serpyllopsis  caespitosa  (Gaud.)  C.  Chr.  . 
THYRSOPTERIS  elegans  Kze  .  .  . 
Trichomanes  exsectum  Kze 

—  Ingae  C.  Chr 

—  philippianum  Sturm      


Mt 

sc 

Mf 

+ 

- 

+ 

+ 

- 

+ 

- 

- 

+ 

+ 

- 

- 

+ 

- 

+ 

+ 

- 

+ 

+ 

- 

+ 

+ 

- 

+ 

+ 

- 

+ 

+ 

- 

+ 

+ 

- 

+ 

+ 

- 

+ 

+ 

- 

- 

+ 

- 

- 

aeriigi7wsimi  (Thouars)  Carm.  (Tristan  da  C.  and  N.  Amsterd.  I.),  and  Marlothii 
Brause  (Cape). 

d.  Meringium  Copel.  About  60,  S.  Chile,  Afr.,  Ceylon,  Formosa,  N.  Guin., 
N.  Zeal.,  Fiji. 

secundum  Hook,  et  Grev.  S.  Chile,  Valdiv.-Fueg. 

plicatum  Kaulf.  See  2/i.g.'j6i.  S.  Chile,  Valdiv.-W.  Patag. 

tortuosum  Hook,  et  Grev.   S.   Chile,  Valdiv.-Fueg.,  Falkl. 


Cyatheaceae. 

Thyrsopteris  elegans  Kze.  Unanimously  regarded  as  a  very  ancient  type  (BowER, 
Seward,  Berry,  Winkler,  Copeland,  etc.).  Berry  (27.  88)  pointed  out  that 
Cyatheoides  thyrsopteroides  Berry  is  remarkably  similar  to  Thyrsopteris;  it  was, 
however,  found  sterile,  and  he  adds  that  most  of  the  fossils  formerly  referred  to 
Thyrsopteris  are  based  on  too  slender  evidence.  Seward  (2.?o.  221)  says:  "Thyr- 
sopteris is  very  closely  allied  to  certain  Jurassic  Ferns  from  the  Yorkshire  coast 
and  many  other  places:  geological  evidence  points  to  a  remote  antiquity,  and  its 
present  isolation  is  in  all  probability  the  last  phase  in  its  history  of  a  direct  deriva- 
tive of  a  widely  scattered  Jurassic  type."  CoPELAND  called  it  "a  relic  from  the 
time  when  Dicksonia  and  Cyathea  had  a  common  ancestor"  ((5^.  48)  and  he  thinks 
that  it  is  allied  to  Culcita  Presl.  He  brought  Thyrsopteris  to  his  large  and  possibly 
very  heterogeneous  family  Pteridaceae,  where  also  Dicksonia  is  placed. 

Winkler's  statement  that  Thyrsopteris  is  found  on  Masafuera  only  [28y.  472) 
is  erroneous. 
Lophosoria  Presl.  One  polymorphous  species. 

quadripinnata  (J.  F.  Gmel.)  C.  Chr.  Trop.  and  subtrop.  Amer.,  Mex.-Chile  and 
south  to  W.  Patag.,  49°  s.  lat.  The  taxonomical  status  of  the  various  forms  re- 
mains to  be  settled.   The  island  form  (or  forms,  for  there  seems  to  be  some  dif- 


222  C.   SKOTTSBERG 

fercnce  between  the  plants  of  Masatierra  and  Masafuera)  is  certainly  unlike  the 
form  inhabitint^   Chile;  see  62.  16. 

l)ickso7iia  Lllcrit.  24;  i  Mex.,  3  Centr.  Amer.  to  Colomb.,  i  Ecuad.-Braz.,  2  Peru; 
1  St.  Helena,  i  Malays.,  4  X.  Guin.,  2  Austral.-Tasm.,  3  N.  Zeal.,  3  X.  Caled., 
I    Mji-Sanioa. 

berteroana  Colla  and  externa  Skottsb.  are  very  near  each  other  and  closely 
related  not  to  the  Andean  group  but  to  I),  laiiaia  Col.  from  X.  Zeal.  CllRlST 
{■,-9.154)  further  cjuotes  as  relatives  1).  antarctica  Labill.  (Austral.),  Bluffiei  {)^zt) 
Moore  (Indones.,  Philij)p.),  gnvidis  Ros.  (X.  Guin.)  and  species  from  X.  Caledonia 
and  the  South   Sea  islands. 

Polypodiaceae. 
Cvstopii-ris  Bernh.   15;    i  boreal,    10  Ind.-China,  Japan,  X.  Zeal.,  2  Eur.,  i  S.  Braz. 

fragilis  (L.)  Bernh.  coll.  A  wide-spread,  polymorphous  sp.,  found  in  all  parts 
of  the  world  (83°  n.  lat.  to  S.  Georgia)  except  Australia.  The  form  occurring  on 
Masafuera  presumably  related  to  a  form  from  the  mainland  [62.  18). 
/>';-)Yy'/r;7>  Adans.  Sens  u  C.  Chr.  about  1 200  sp.;  s.  str.  Coi'KLANl)  about  150;  "an 
assemblage  of  genera  that  must  be  segregated  in  their  entirety  before  of  use  for 
a  study  of  distribution"   (Coi'KLAXD  6g). 

inaeqtialifolia  (Colla)  C.   Chr.  Belongs  to  a  neotrop.  group  and  comes  nearer 
to  tropical  forms  than  to  the  south  Andean  I),  spcctabilis  (Kaulf.)  C.  Chr.,  which 
extends  south  to   CUiile. 
PolysticJiuui  Roth.  About  225;   widely  spread,   numerous  in  E.  As. 

berterianum  (Colla)  C.  Chr.  Xear  P.  adiantiforrnc  Eorst.,  spread  through  the 
S.   temp,   zone,  north  to  W.  Ind. 

I'i'stituni  (k'orst.)  I'resl  coll.   S.  Amer.   (also  Chile),  Austral.,  X.  Zeal. 
Artliyoptcris  ].  Sm.  About  20;  j)alaeotrop.,  recorded  from  trop.  Afr.,  north  to  Arab.; 
Madag.,   Australas.,  north  to  l^hilipj).  Is.;   Xew  Caled.,  Fiji,  Samoa;   best  developed 
X.   (iuin.,  X.   Caled.  and  Madag. 

altescandens  (Colla)  J.  .Sm.   Closely  related  to  Pacific  forms  (62.21). 
Asplciiiii))!   L.  650-700;   cosmopolitan. 

obliquiiDi  Eorst.  var.  cJioidrophyllum  (Bert.)  Mett.  Typical  obliqutwi  reported 
from  S.   Chile,  Austral,   and  X.   Zeal. 

tnacrosorutn  Bert,   ex   Colla.  Distantly  related  to  neotrop.  sp.;  see  62.  23. 

stellatutn  Colla.   Belongs  to  the  pantrop.  ///;/;^/<r? /;/;;/  group;  see  ^2.  24. 

day f aides  Desv.  (magcllanicum  Kaulf.).  S.  Chile,  Eueg.,  Ealkl.  Closely  related 
to  A.  alrarczciise  Rudm.  Ikown  of  Diego  Alvarez  (Tristan  da  C);  see  61.  13. 
lUiilniuui   L.    180-200;   essentially  southern. 

auriculatuni  Cav.  Temj).  S.  Amer.,  common  in  Centr.  and  S.  Chile  and  related 
to   //.   anstralc   L.   (S.   Afr.). 

raldinioisc  C.  Chr.  Chile,  south  to  Chiloe.  Related  to  B.  lanccolaium  (R.  Br.) 
.Sturm   from   Austral,   and   \.  Zeal.   As  regards  the  nomenclature,  see  i6g.  54,  and 

Schottii  (Colla)  C.  Chr.  Stands  near  B.  aitcnuatiwi  (Willd.)  C.  Chr.  (S.  Afr., 
IC.   Austral.,  Polyn.)  and  nieridejise  (Kaulf.)  C.   Chr.  (trop.  Amer.);  see  62.  27. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  22$ 

chilense  (Kaulf.)  Mett.  Centr.  Chile  to  W.  Patag.;  Falkl.  Belongs  to  the  capense 
group  (S.  Afr.,  Indomal.),  see  62.  2^]  and  i6g.^']. 

cycadifolium  (Colla)  Sturm.  Almost  too  near  B.  viagellmiicum  (Desv.)  Mett. 
(S.  Chile  to  Fueg.  and  Falkl.);  both  related  to  B.  tabulare  (Thunb.)  Kuhn  (S.  and 
E.  Afr.,  Madag.,  Mascaren.). 

longicauda  C.  Chr.  Very  near  B.  Sprucei  C.  Chr.  from  trop.  S.  Amer.  (Ecuad., 
Boliv.,  Braz.),  see  2^0.  HiCKEN  records  it  from  Tucuman  in  Argent.  (133.  246), 
a  statement  overlooked  also  by  Christensen. 

Pellaea  Link.  About  80,  the  majority  in  S.  Amer.  (south  to  Chile),  S.  Afr.  and 
islands;  N.  Amer.  north  to  Canada;  N.  Zeal. 

chilensis  Fee.  Very  close  to  P.  nivea  (Poir.)  Prantl  (Ariz.-Chile),  see  62.  30. 
Hypolepis  Bernh.  A  pantrop.  genus  of  45-50  sp.,  half  of  them  trop.  Amer.,  several 
Afr.  and  surrounding  islands,  others  N.  Guin.-E.  As.  and  Austral. -Poly n. 

rugosida  (Labill.)  J.  Sm.  Centr.  and  S.  Chile;  the  typ.  sp.  Austral.,  N.  Zeal., 
var.    villosoviscida  (Thouars)  C.  Chr.  [61.  6)  on  Tristan  da  C;  Polypodiu7n  viscidum 
Roxb.  from  St.  Helena  very  likely  is  another  variety. 
Adiafitum  L.  200-225;   widely  distributed  but  most  numerous  in  S.  Amer. 

chilense  Kaulf.  Peru-Chile,  Patag.,  Falkl. 
Pteris  L.  270-280,  mainly  trop.  but  extending  north  to  S.  Eur.  and  south  to  S.  Afr., 
Tasm.  and  N.  Zeal. 

chilensis  Desv.  Near  P.  leptophylla  Sw.  (S.  Braz.). 

semiadnata  Phil.  S.  Chile,  Valdiv.-Huafo  I.  A  distinct  sp.,  possibly  related 
to  P.  pidchra  Schlechtd.  from  Mexico. 

berteroana  Ag.  Related  to  P.  comans  Forst.  (Austral.,  Tasm.,  N.  Zeal.)  and 
particularly  to  P.  endlicheriana  Ag.  (Norfolk  I.). 

Hisiiopteris  (Ag.)  J.  Sm.  An  assemblage  of  trop.  and  austr.-circump.  forms  which 
according  to  CllRlSTENSEN  cannot  claim  to  be  regarded  as  different  species 
[62.  36),  whereas  Copeland  [6g.  60)  speaks  of  local,  derived  species  (a  concentra- 
tion in  Indomal.-Polyn.)  differing  sufficiently  from  the  following  polymorphous  taxon. 

incisa  (Thunb.)  J.  Sm.  Chile,  S.  Afr.,  Tristan  da  C,  Austral,  Tasm.,  N.  Zeal., 
Polynes. 

Polypodiwn  L.  coll.  World-wide;  Ind.  Fil.  registers  more  than  iioo  sp.  The  genus 
is  now  generally  broken  up,  most  radically  by  CoPELAND,  who  segregates  numer- 
ous genera  partly  restored  from  synonymy. 

a.  Granimitis  Sw.  About  150;  an  essentially  southern  genus,  reaching  north 
to  West  Indies  and  Bonin  Is.  and  richest  developed  in  New  Guinea. 

magellanica  Desv.  S.  Chile,  Valdiv.-Fueg,,  And.  Patag.  (Rio  Negro);  Tristan 
da  C,  Marion  I.  In  62.  36  as  P.  Billardieri  (Willd.)  C.  Chr.  var.  niagellanician 
(Desv.)  C.  Chr.,  but  later  restored  to  specific  rank  by  CHRISTENSEN  61.  18.  Distri- 
bution of  Billardieri:  Austral.,  Tasm.,  N.  Zeal.,  Auckl.  and  Campb.  Is.,  Lord 
Howe  and  Norfolk  Is.,  St.  Paul  and  N.  Amsterd.  Is.,  Kerguel. 

b.  Synammia  Presl,  According  to  Copeland  6g.  184  only  one  species,  wS. 
Feuillei  (Bert.)  Copel.  [Polypodiwn  trilobmn  Cav.),  but  I  fail  to  see  how  we  could 
exclude  the  island  species. 


224 


C.  SKOTTSBERG 


intermedium  Colla  [translucens  (Kze)  Fee].  JOHOW  united  P.  irilobimi  Cav. 
and  califoyjiicuDi  Kaulf.  with  traiislucens,  and  they  seem  to  belong  to  the  same 
small  group,  to  which  also  P.  Espiiiosae  W'eatherby  (Chile,  Atacama)  must  be 
rcferretl. 

c.  PolvpodiioJi  s.  str.,  about  75   sp.,  mainly  N.   hemisph.   and  neotrop. 
Masafiierae  Phil.   S.   Peru  to  N.  Chile  (Antofagasta),  Argent.;   comp.  i^g.  14 

and  16S.  33.   \>ry  close  to  P.  pycnocarpiiui  C.   Chr.  (Mex.-N.   Chile). 

d.  Xiplwptcris  Kaulf.  About   50,  pantrop. 

tyichonianoides  Sw.  Trop.  Amer.   A   puzzling  record  (see  2^g.  766). 

e.  Plcopeliis  Humb.  et  Bonpl.  About  40,  pantrop.;  the  following  sp.  wide- 
spread. 

la}iceolatii>n  L.  Mex.  and  \\\  Ind.  to  subtrop.  S.  Amer.  (also  in  Chile),  St. 
Helena,  Tristan  da  C,  Afr.,  Madag.,  Indomal.,  Hawaii. 

lUapJioglossiiui  Schott.    A  pantrop.   genus  of  more  than  400  sp.,  very  numerous 
in   And.   S.   Amer.,  4  sp.  on  Tristan  da  C,  and  many  sp.  in  Polyn.  and  Hawaii. 

Li)idenii  (Bory)  Moore.   Mex.  to  Ecuad.   and  Braz. 

Gleicheniaceae. 
GleicJienia  Sm.  About   130  sp.,  the  majority  of  them  referred  to  5//f//<f?7/i- (Kaulf.) 
Ching,    to    which    also  the  species  in  J.  P'ern.   belong;    about  a  dozen  sp.  spread 
over    the   austral    zone;    5    Chile,  one  extending  to  Falkl.;    i    S.  Afr.,    i   Madag., 
Mascaren.  and  Seych.  Is.,    i   Tasm.,   i   Tasm.  and  N.  Zeal.,  2  N.  Zeal. 

quadripartita  (Poir.)  Moore.   S.   Chile  to  Fueg. 

pedalis  (Kaulf.)  Spreng.   S.   Chile,  south  to  Chonos  Is. 

cf.  litoralis  (Phil.)  C.  Chr.  Hardly  identical  with  this  little  known  species  and 
perhaps  only  a  form  of  G.  pedalis. 

Ophioglossaceae. 
OpJiioglossuDi  L.   28  sp.   recognized  by  Clausen  {64);  the  genus  "scattered  with 
remarkable  uniformity  over  the  habitable  globe"   (COPELAM)  6g.  12). 

fertiandezianum  C.  Chr.  Not  very  near  0.  ypaiicnse  Mart.  (Colomb.,  Braz.) 
as  Chris  ri-:NSEX  thought,  but  close  to  0.  scariosuni  Clausen  from  Peru,  Dept.  Junin. 

Lycopodiaceae. 
Lycopodiuin  L.   A  large  world-wide  genus,  badly  ill-treated  by  Nessel,   who  listed 
several   undistinguishable  species  in  J.  P^ern.;  see  2/i.g.  ^66. 

j/Kii^r/Iimiiinn   Sw.   .S.  Chile  to  P\ieg.,   P'alkl.,   S.  Georgia,  Marion  I.,  Kerguel. 

scariosiiiH  I^'orst.  Chile,  Valdiv. -Guaytecas  Is.  [L.  gayajiuni  Remy),  N.  Zeal.; 
Tr(){).   Andes,   Braz.  [L.  Jiissicui  Desv.).   See  I.e.   J^S'] . 

In  his  treatment  of  the  geographical  groups  distinguished  by  him,  JOIIOW 
included  the  ferns  —  no  1^'ern  Allies  were  at  that  time  known  from  Juan  Fernandez. 
All  were  classified  as  American  exce[)t  3,  Dicksonia  berteroafia,  said  to  be  similar 
to  a  species  from  T'iji,  Ptrris  bcrtcroaiia  (identified  with  comans),  and  '' Aspleniimi 
/oJigissiuiNiN  Illume",  the  s[)ecies  now  known  as  BlecJinimi  longicatida.  In  1914 
[22^)  I  went  a  step  further;  to  my  "Altpazifisches  P^lement"  5  species  were  referred, 
to  the  Trop.  American  5,  and  all  the  rest  to  the  Chilean.  In  1934  I  attempted 
a  more  detailed  subdivision   (279,  here  translated  from  P>ench). 


DERIVATION  OF  THE  FLORA  AND  FAUNA  225 

1.  Neotropical  and  Andean  element.  Species  either  found  in  Centr.  and  S.  Chile  or 
endemic  but  allied  to  Chilean  species;   28. 

a)  in  Chile,   22. 

b)  in  the  Magellan,  region,  but  not  of  subantarctic  character.   3. 
d)  endemic  species  of  non-endemic  genera,  3. 

2.  Element  consisting  of  wide-spread  species,  also  inhabiting  Chile,   2. 

3.  Neotropical  element,  not  represented  in   Chile,   7. 

a)  non-endemic  species,    i. 

b)  endemic  species  of  non-endemic  genera,   6. 

4.  Magellanian-Old  Antarctic  element,   9. 

a)  non-endemic  Magellanian  species,   2. 

b)  ,,  ,,         Old  Antarctic  but  not  Magellanian  species,  4. 

c)  endemic  species  allied  to  subantarctic  species,   2.  , 

d)  ,,  ,,        not  allied  to  subantarctic  species,    i. 

5.  Pacific  element,   5. 

a)  isolated  endemic  species  of  wide  genera,    i. 

b)  endemic  species  belonging  to  genera  or  sections  of  west  to  central  Pacific  distribu- 
tion,  3. 

d)  species  of  endemic  genus  with  unknown  relations,    i. 

A  modified  arrangement  is  attempted  below,  corresponding  to  the  analysis 
of  the  angiosperms. 

I.  Andine-Chilean  element. — 34  (64.1  %). 

a.  Endemic  species  (5,  9.4%):  Hymenophyllum  rugosum,  Polystichum  berteri- 
anum,  Blechnum  cycadifolium,  Pellaea  chilensis,  Polypodium  intermedium. 

d.  Known  from  continental  Chile,  many  also  in  other  parts  of  S.  America 
or  of  still  wider  distribution  (29,  54.7%):  Trichomanes  exsectum,  Hymenoglossum 
cruentum,  Hymenophyllum  cuneatum,  caudiculatum,  fuciforme,  pectinatum,  ferru- 
gineum,  secundum,  plicatum  and  tortuosum,  Lophosoria  quadripinnata,  Cystopteris 
fragilis,  Polystichum  vestitum,  Asplenium  obliquum,  Blechnum  auriculatum,  valdi- 
viense  and  chilense,  Hypolepis  rugosula,  Adiantum  chilense,  Pteris  chilensis  and 
semiadnata,  Histiopteris  incisa,  Polypodium  magellanicum,  Masafuerae  and  lanceo- 
latum,  Gleichenia  quadripartita,  pedalis  and  cf.  litoralis,  Lycopodium  scariosum. 

II.   Subantarctic-Magellanian  element. — 4  (75%). 
Not  endemic,  extending  north  but  not  beyond  Centr.  Chile:  Serpyllopsis  caes- 
pitosa,  Hymenophyllum  falklandicum,  Asplenium  dareoides,  Lycopodium  magel- 
lanicum. 

III.  Neotropical  element. — 9  (17  %). 

a.  Endemic  species  (7,  13.1  %):  Trichomanes  philippianum  and  Ingae,  Dryo- 
pteris  inaequalifolia,  Asplenium  macrosorum  and  stellatum,  Blechnum  longicauda, 
Ophioglossum   fernandezianum. 

b.  Not  endemic  (2,  3.8  %):  Polypodium  trichomanoides,  Elaphoglossum  Lindenii. 

IV.  West  Pacific  element. — 5  (94%). 

Affinities  in  Australia,  New  Zealand  and  Oceania,  not  in  America.  All  en- 
demic: Dicksonia  berteroana  and  externa,  Arthropteris  altescandens,  Blechnum 
Schottii,  Pteris  berteroana. 

15  ""  557857  The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.    Vol.  I 


226  C.   SKOTTSBERG 

V.   Eu-Fernandezian  element. —  i  (1.9%). 

Endemic  genus  witliout  affinities  among  the  living  ferns:  Thyrsopteris  elegans. 

As  could  be  expected,  the  Chilean  element  is  stronger  among  the  ferns  than 
among  the  angiosperms,  followed  in  great  distance  by  the  neotropical,  which  also 
is  comparatively  stronger,  whereas  the  Magellanian  and  the  Pacific  are  less  im- 
portant, especially  the  latter;  long-distance  advocates  w^ould  expect  the  opposite, 
because  ferns  ought  to  travel  across  the  ocean  much  more  readily  than  flowering 
plants. 

III.  Musci. 

The  following  list  is  based  on  Brotiierus'  paper  in  vol.  II  ( ?^)  and  on  his 
treatment  of  the  group  in  the  2nd  edition  of  Natiirl.  Pflanzenfam.  In  many  cases 
the  figure  for  the  number  of  species  (in  brackets)  is  too  low,  as  numerous  mosses 
have  been  described  in  later  years,  but  I  have  refrained  from  searching  the  litera- 
ture. Much  information  was  obtained  from  Herzog's  work  (i2g)  and  in  pt.  2  of 
Irmscher's  book  (i4j).  I  am  indebted  to  Dr.  Herman  Persson  for  kind  assistance. 

Mi  =-  Masatierra,  Mf  =  Masafuera. 

Ditrichaceae. 

l^Uiiridium  Brid.  (about  30).  Wide-spread,  mainly  temperate. 

Robinsomi  (Mont.)  Mitt.  Chile,  Urug.  Belongs  to  a  S.  Amer.  group. — Mt. 
DitricJiuni  Timm  (about   50).  In  all  parts  of  the  world. 

affine  (CM.)  Hampe.  S.  Chile  and  Patag.,  E.  Austral.,  N.  Zeal,  Auckl.  Is. 
Not  listed  in  210,  where  D.  elongatiim  (H.  f.  et  W.)  Mitt,  is  quoted  for  Chile,  Austral., 
Tasm.  and  N.  Zeal.— Mt,  Mf. 

lougisetum  (Hampe)  Jaeg.  S.   Chile  to  Fueg. — Mf. 
Ceratodoi  Brid.  (2,  one  trop.). 

puypurcus  (L.)  Brid.   Cosmopol. — Mt,  Mf. 

Pottiaceae. 

llymc}wst()))iu))i  R.   Br.  (50-60).  All  over  the  world. 

kuuzcanum  (CM.)  Broth.   S.   Chile.— Mt. 
(jyuiuostannufi   Hedw.   (10).   All  over  the  world. 

calcariUDi   Jkyol.  germ.   Eur.,  Afr.,  As.,  N.  Amer.,  S.  Amer.:  P3cuad.-Chile; 
Austral.,  Tasm.,  X.  Zeal.— Mt,  Mf. 
Tricliostomum  Hedw.  (81).  Cosmopolitan. 

hracliydoiitiuin  Bruch.  Eur.,  Caucas.,  N.  Afr.,  Macaron.,  Mascar.,  Japan,  N. 
Zeal.,  but  not  mentioned  in  210. — Mf. 

Dicranaceae. 

AmpJiidium  Xees  (10).  Very  widely  distributed. 

cyatJiicarpum  (Mont.)  Broth.  Ecuad.-Chile,  S.  Georgia,  Afr.,  E.  Austral.,  Tasm., 
N.  Zeal.— Mt,  :\If. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  227 

Dicranella  Schimp.  (6o).  All  continents,  numerous  S.  Amer.  and  As. 

costata  Broth. — Mt,  Mf. 
Oncophorus  Brid.  (8).  Southern  S.  Amer.,  Eur.,  E.  As.,  Ceylon. 

fuegiaims  Card.  Patag.-Fueg. — Mf. 
Dicranolonia  Ren.  [']6).  Almost  exclusively  S.  hemisph,,  austr.-subantarct.,  i  S.  Afr. 

fernandezianum  Broth.  Near  D.  Dusenii  Broth.  (S.  Chile). — Mt, 

capillifoliwn  Broth.  S.  Chile-Fueg. — Mt. 

capillifolioides  Broth.  Near  the  former. — Mf. 

Menziesii  (Tayl.)  Par.  S.  Chile,  E.  Austral.,  Tasm.,  Norfolk  I.,  N.  Zeal., 
Chatham  Is.,  Auckl.  Is. — Mt. 

Billardieri  (Schwaegr.)  Par.  Peru-Fueg.,  Falkl,  S.  Afr.,  Austral.,  Tasm.,  N. 
Zeal,  Auckl.  and  Campb.  Is. — Mt,  Mf. 

nigricaule  Angstr.  S.  Chile,  south  to  Fueg. — Mf. 
Campylopus  Brid.  (about  500).  All  over  the  world,  mainly  trop.-subtrop. 

mtroflexus  (Hedw.)  Mitt.  N.  Amer.,  S.  Amer.,  Ecuad.  to  Urug.  and  Fueg., 
islands  of  W.  and  E.  Afr.,  Ascens.,  St.  Helena,  Tristan  da  C,  Marion  I.,  Austral., 
Tasm.,  N.  Zeal.,  Auckl.,  Campb.  and  Antipodes  Is.,  Pacific  islands. — Mt. 

trimcatus  CM.  Chile.— Mt. 

polytricJwides  De  Not.  W.  and  S.  Eur.,  N.  Afr.,  Mascar. — Mt. 

aberrans  Broth.  A  very  peculiar  sp. — Mt,  Mf. 

areodiciyon  (C.  M.)  Mitt.  Centr.  and  S.  Amer.,  Venez.-Boliv. — Mf. 

subareodictyon  Broth.  Related  to  the  former. — Mf. 

blindioides  Broth. — Mf. 
Thysanomitrium  Schwaegr.  (31).  Trop.-subtrop.,  especially  Old  World,  extending 
south   to  the  austr.-subantarct.  zone. 

Richardi  Schwaegr.  Centr.  and  S.  Amer.  to  Chile. — Mt,  Mf. 

leptodiis  (Mitt.)  Broth.  \Campylopus  clavatus  (R.  Br.)  H.  f.  et  W.]  Ecuad.,  Chile, 
Austral,  Tasm.,  N.  Zeal,  Auckl.  and  Campb.  Is. — Mt,  Mf. 

Dicnemonaceae. 

Eucamptodon  Mont.  (7).  Chile  (i).  Austral  (i),  N.  Caled.  {4),  N.  Zeal  +  N.  Caled.(i). 
perichaetialis  Mont.  S.   Chile  to  Magell — Mf. 

Fissidentaceae. 

Fissidens  Hedw.  (over  700).  All  continents,  very  numerous  in  trop.  zones. 

fernandezianus  Broth. — Mt,  Mf. 

crassicuspes  Broth.  Related  to  F.crass/pes  Wi\s.  (Eur.  Mediterr.,  Madeira). — Mt. 

rigidulus  Hook.  f.  et  Wils.  Ecuad.-W.  Patag.,  Austral,  Tasm.,  N.  Zeal. — 
Mt,  Mf. 

leptochaete  Dus.  Chile.— Mt. 

maschalanthus  Mont.  Chile,  south  to  W.  Patag. — Mt,  Mf. 

pycnotylus  Broth.  Very  near  the  former. — Mt. 

asplenioides  (Sw.)  Hedw.  Centr.  Amer.  to  Peru,  S.  Chile  and  Braz.,  W.  Afr., 
Macaron.,  Tristan  da  C,  Indomal,  Queensl,  N.  Zeal. — Mt. 


228  C.  SKOTTSBERG 

LeptodoJitiuDi  Mampe  (80).  Most  numerous  in  Amer.,  40%   And. 

fernandezianum  Ikoth.  Related  to  L.  luteum  (Tayl.)  Mitt.  (trop.  And.). — Mf. 
Diiiymodon  Hedw.  (91).  Subcosmopol.,  mainly  temp.,  essentially  Amer.:  Mex,— 
Centr.   Amer. -Peru   and  Chile;    i    Antarct. 

calytnperidictyon  Broth. — ^Mt. 

linearis  l^roth. — -Mf. 
Tottula  Hetlw.  (220).  Subcosmopol.,  mainly  temp. 

scabriutrris  (C.  M.)  Mitt.   Chile.— Mt. 

flaij^cllayis  (Schimp.)  Mont.   Chile.— Mt. 
G}i)H)}iia  IChrh.  (230).   Subcosmopol.   but  rare  in  the  trop.  zones. 

phyllorhizans  Broth. — Mt. 
Kluicomitriu))i  Ikid.  (80).  As   Grimmia. 

subuiirritum  (CM.)  Par.  W.  Patag.-P\ieg.— Mf. 

sy)}iphyodo)itu})i  (C.  M.)  Jaeg.  S.  Chile  to  P\ieg.  and  Falkl.,  S.  Afr.,  Kerguel., 
Tasm.,   \.   Zeal.  Not  listed   for  N.  Zeal,   in  210. — Mt,  Mf. 

striatipiliDJi  Card.   S.  Chile  to  Fueg.,  S.  Georgia;   N.  Zeal.  [^216). — Mf. 

lajiugiiiosuni  (Hedw.)  Brid.   Cosmopol. — Mf. 

loriforme  Dus.   W.   Patag. — Mf. 

conrolutuni  Mont.   S.   Chile  to   W.   Patag.— Mf. 

Ptychomitfiaceae. 

Ptycho))utrin))i  (Bruch)  P'urnr.  (62).   Of  wide  distribution   in   temp,  zones. 
fernandezianum  (Mitt.)  Jaeg.^Mt,  Mf. 

Orthotrichaceae. 

Zyo^odoii  Hook.  f.  et  Tayl.  (about  100).  N.  and  Centr.  Amer.  (6),  trop.  S.  Amer. 
(40),  .S.  Chile  and  Argent.  (20),  \\\\x.  (4),  trop.  Afr.  (2),  trop.  As.  (6),  \l.  Austral.- 
X.   Zeal.  (10). 

i)itcn)ii'dius  B.  et  S.  S.  Amer.,  Afr.,  Monsoon  reg.,  Austral.,  Tasm.,  N.  Zeal. 
—  Mt.  [26^).  Z.  oboi'alis  Mitt.,  a  doubtful  species  (,?/?.  139).  is  supposed  to  be  iden- 
tical  with  i?ifrn>ifdii(s. 

Moiziesii  (Schwaegr.)  W.  Arn.   S.   Chile,  Austral.,  Tasm.,  N.   Zeal. — Mt. 
Sfoioi/iitriinn  (Mitt.)   l^roth.  (2).   Viiru,   Chile. 

pojtastichuni  (Mont.)  Ikoth.   Peri'i-W.   Patag.— Mf. 
riota  Mohr  (43).    X.   and   S.   temj).   zones,   ^3  Chile,   some  N.   Zeal. 

fernandeziana  Malta  (.?/?).  Xear  ('.  rufula  (Mitt.)  Jaeg.  (Chile-W.  Patag., 
Argent.,  X.  Zeal.).— Mf. 

Macromitriion  Brid.  (415).  Troix-subtro|).,  special  gr()uj)s  Chile-Patag.  and  P2.  Austral.- 
Tasm.-X.   Zeal. 

Iiyinoiostouiu))!   Mont.   S.   Chile-P'ueg. — Mt. 

saxatile  Mitt. — Mt,  Mf. 

fernandezianum  Broth.  Related  to  J/,  aspcruluni  Mitt.  (Tasm.,  N.  Zeal.;  not  in 
210).— Vii. 

Masafuerae  l^roth.   Xear  the  former. — Mf. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  229 

Funariaceae. 

Funaria  Schreb.  (200).  A  world-wide  genus. 

hygroinetrica  (L.)  Sibth.  Cosmopol. — Mt,  Mf. 

Bryaceae. 

Mielichhoferia  Hornsch.  (100).  Widely  distributed,  centering  in  the  Andes. 

longiseta  C.  M.  Ecuador. — Mt. 
Bryum  Dill,  (about  800).  All  over  the  world. 

Lechleri  C.  M.  Chile.— Mf. 

fernandezianum  Broth.  Related  to  B.   Cruegerz  Hampe  (nGotrop.). — Mt,  Mf. 

Leptostomaceae. 

Leptostomutn  R.  Br.  (i  i).  S.  Chile  (2),  Indomal.,  Austral.,  N.  Zeal.,  Norfolk  I.,  Campb.  I. 
Menziesii  (Hook.)  R.  Br.   S.  Chile  to   W.  Patag.  and  Fueg.— Mf. 

Eustichiaceae. 

Eustichia  (Brid.)  Mitt.  (8).  6  Mex.-S.  Chile,   i   S.  Afr.,   i   islands  of  E.  Afr. 
Poeppigii  (C.  M.)  Par.  S.  Chile-Magell.— Mt,  Mf. 

Rhizogoniaceae . 

Rhizogonium  Brid.  (27).  Austr.-circump.  (S.  Amer.,  S.  Afr.,  Austral.,  N.  Zeal.,  Polyn.), 
extending  into  the  N.  hemisph.;    i   pantrop. 

Nouae  Hollandiae  Brid.  v2iX. patagonicum  Card,  et  Broth.  W.  Patag.;  the  typical 
sp.  Austral.,  Tasm.,  N.  Zeal.— Mt. 

mnioides  (Hook.)  Schimp.  Colomb.-Fueg.,  E.  Austral.,  Tasm.,  N.  Zeal. — Mf. 

Bartramiaceae. 

Anacolia  Schimp.  (7).  W.  N.  Amer.-trop.  Andes  (5),   i   Medit.,   i   Ethiop. 

subsessilis  (Tayl.)  Broth.  Mex.-Ecuad.— Mt,  Mf. 
Bartramia  Hedw.  (iio).   Subcosmopol.,  a  few  sp.  subantarct.-circump. 

aristata  Schimp.   Chile.— Mt,  Mf. 

patens  Brid.  W.  Patag.-Fueg.,  Falkl.,  S.  Georgia,  Kerguel.,  E.  Austral.,  Tasm., 
N.  Zeal. — Mf.  Not  mentioned  in  210,  where  B.  halleriajta  Hedw.,  widely  distributed 
in  the  N.  hemisphere,  is  credited  to  Patag.,  Fueg.,  Austral.,  Tasm.  and  N.  Zeal. — Mf. 

fernandeziana  Card. — Mt. 
Philonotis  Brid.  (174).  Subcosmopol.,  two  monotyp.  sections  in  S.  Amer. 

krauseana  (C.  M.)  Jaeg.  Centr.  Chile  to  W.  Patag.— Mt,  Mf. 

glabrata  Broth.  Related  to  the  former. — Mt. 

scabrifolia  (Hook.  f.  et  Wils.)  Broth.  Circump.-subantarct.,  forming  a  separate 
section.  Ecuad.-Fueg.,  Falkl.,  S.  Georgia,  S.  Afr.,  Marion  I.,  Kerguel.,  Austral., 
Tasm.,  N.  Zeal,  Auckl.  and  Campb.  Is.— Mt,  Mf. 

vagans  (Hook.  f.  et  Wils.)  Mitt.  Forms  a  separate  section.  Chile-Fueg.,  S. 
Georgia. — Mf  (an  aberrant  form). 

Breutelia  Schimp.  (104).  Best  developed  trop.  mountains,  rare  N.  Amer.,  numerous 
austr.-subantarct. 

Masafuerae  Broth. — Mf. 


230 


C.   SKOTTSBERG 


Hedwigiaceae. 

Rhacocarpus  Lindb.  (24).  Centr.  Andes  and  Braz.  mountains  etc.;  almost  confined 
to  the  S.   hcmisph. 

Iliiuiboldtii  \\\(^oV)  Lindb.  Me.x.,  \V.  Ind.,  Colomb.-And.  Patag.,  Fueg.,  Falkl., 
Centr.   Afr..  Madag.,  Reunion,  Austral.,  Tasm.,  N.  Zeal.  A  polymorphous  sp. — Mf. 

Cryphaeaceae. 

Deudrocryf^haea  Par.  et  Schimp.  (4).  Austr.-subantarct.,  2  Chile,  I  Argent.-Patag., 
I    Tasm.-\.  Zeal. 

cuspidata  (Sull.)  Broth.   S.   Chile. — Mf. 
Cvptodon  I'ar.  et  Schimp.  (6).  2  Austral.-N.  Zeal,  1  X.  Caled.,  2  Fiji-Samoa-Tonga, 
I  J.   Fern. 

crassinervis    Broth.   Close  to   C.  (210  under   CrypJiaea)  dilatatus  (Hook.  f.  et 
Wils.)  Par.  (h:.  Austral.,  X.  Zeal.).— Mf. 

Lepyrodontaceae. 

Lcpyrodou  Hampe  (6-8).  i  bicentric  (Chile-Fueg.,  P3.  Austral.,  Tasm.,  N.  Zeal., 
Canipb.  I.),   2  or  3  trop.  Andes  to  Braz.  and  Argent.,   2  or  3   Chile,    I   N.  Zeal. 

pannilus  Mitt.   Centr.  and  S.   Chile.— Mt,  Mf. 

tojnffitosus  (Hook.)  Mitt.  Colomb.-Perii,  Braz.,  Argent.,  W.  Patag. — Mt. 

iuiplexus  (Kze)  Par.   Chile,  south  to  Fueg.- — Mt. 

Ptychomniaceae. 

PlycJioDuuH))!  1  look.  f.  et  Wils.  (9,  or  8  if  cyg)iisetu))i  =  acicularc).  Austral-subant- 
arct.,  e.xtending  north  to  Braz.  and  Hawaii;  Argent,  and  Chile  to  Pueg.,  \\.  Austral., 
Tasm.,   X.  Zeal,   and  subant.   islands.   Lord  Howe  L,  X.   Caled.,  Polyn. 

suhaciculare  Besch.   Chile,  south   to  I^\ieg. — Mt. 

falcatiilum  Broth.   Related  to  the  former. — Mt. 

ptychocaypian  (Schwacgr.)  Mitt.   Chile,  south  to  \V.  Patag.— Mf. 

Neckeraceae. 

WeyDioutliia  Ikoth.  (3-4).   Austr.-circump.,  Hawaii. 

iHol/is   (Hedw.)    Broth.    Centr.    and    S.   Ciiile  to  Magell.,  PI  Austral.,  Tasm., 
X.   Zeal.-  Mt,  Mf. 
Ltplodo}!  Mohr  (4I   Afr.   and   islands  (3),  and  the  following. 

Sniithii  (Dicks.)  Mohr.   Argent.,  Chile;   S.   ICur.,  Caucas.,  Afr.,  Macaron.;  Y.. 
Austral.,   X.  Zeal.^Mt,   Mf. 

Xcckcra  Hedw.  (127).  Widely  distributed  troi).-subtrop.,  extending  south  to  Chile 
and    X.   Zcal.-Auckl.   I. 

rotundata  Broth.   A  very  distinct  sp. — Mf. 
PoyothainjiiuDi   IHeisch.   (51).    i    W.   X.   Amer.,  41    Centr.  and  S.  Amer.,  extending 
south  to  Patag.,   7   trop.   Afr.,    i    Ceylon,    i    X.   Zeal. 

fasciculaiion  (Sw.)    Meisch.   W.   Ind.,   Colomb.-Peru,   Ikaz. — Mf. 

arbuscnlans  (C.   M.)   Broth.   Chile,   Patag. — Mf. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  23 1 

Thamnium  Bryol.  eur.  (33).  N.  Amer.,  W.  and  S.  Eur.,  Macaron.,  S.  Afr.,  N.  and 
E.  As.,  Indomal.,  Austral,  N.  Zeal.,  Melan.  The  Juan  Fernandez  species  belong 
to  a  southern  section. 

rigidum  (Mitt.)  Broth.— Mt. 

latinerve  (Mitt.)  Broth.— Mt. 

Caroli  Broth.  Related  to  rigidum. — Mt. 

Ingae  Broth.  Very  near  the  former. — Mt. 

crassinervium  (Mitt.)  Broth. — Mt. 

proboscideum  Broth.  Near  the  former. — Mt. 

assitnile  Broth.  Very  near  the  former. — Mt. 

confertum  (Mitt.)  Broth.— Mt. 
Piimatella  (C.  M.)  Fleisch.  (37).  Pantrop. 

tnacrosticta  Broth.  A  very  distinct  sp. — Mf. 

Hookeriaceae. 

Distichophyllum  Dozy  et  Molk.  (93).  Essentially  austral,  extending  south  to  Austral., 
Tasm.,  N.  Zeal;  well  represented  Indomal.  region. 

subelimbatum  Broth. — Mt. 

assitnile  Broth.  Near  the  former. — Mt. 

fernandezianum  Broth.  Related  to  D,  rotundifoliuin  (Hook.  f.  et  Wils.)  Broth. 
(Chile,  Patag.,  E.  Austral,  Tasm.,  N.  Zeal).— Mf. 

Pterygophyllum  Brid.  (32).  Mainly  S.  hemisph.,  6  S.  Chile-Fueg.,  25  Austral-Tasm.- 
N.  Zeal,  2  of  these  also  S.  Amer. 

anomalum  (Schwaegr.)  Mitt.  F'ueg. — Mt.  Perhaps  only  an  extreme  water  form 
of  the  following  (jjj.  85) 

obscurum  (Mont.)  Mitt.  S.   Chile  to  W.  Patag.,  Falkl,  Tasm.— Mf. 

tenuinerve  Broth.  Very  near  the  former. — Mt. 

denticulatum  (Hook.  f.  et  Wils.)  Mitt.  W.  Patag.-Fueg.,  Falkl,  E.  Austral, 
Tasm.,  N.  Zeal,  Auckl  and  Campbell  Is. — Mt.  Not  included  in  210,  where 
P.  dentatum  (Hook.  f.  et  Wils.)  Mitt,  is  supposed  to  occur  in  Chile  and  P'uegia. 
Eriopus  (Brid.)  C.  M.  (25).  Some  trop.  Andes,   i    S.  Afr.,  numerous  N.  Zeal. 

leptoloma  Broth.  Related  to  E.  apiculatus  (Hook.  f.  et  Wils.)  Mitt.  (S.  Chile- 
Fueg.,  Austral,  Tasm.,  N.  Zeal).— Mt,  Mf. 

grandiretis  Broth.  Near  the  former. — Mf. 
Lamprophyllum  Schimp.  Monotypical;  more  or  less  distantly  related  to  neotrop. 
genera. 

splendidissimum  (Mont.)  Schimp.  S.  Chile  and  W.  Patag. — Mf. 

Hypopterygiaceae. 

Lopidium  Hook.  f.  et  Wils.  (16).    14  paleotrop.,  2  S.  Amer.  (Braz.,  Chile). 

concinnum  (Hook.)  Fleisch.  Chile,  south  to  W.  Patag.,  Austral,  Tasm.,  N.  Zeal, 
Auckl.  Is.— Mt. 

Hypopterygium  Brid.  (61).  Widely  distributed  N.  and  S.  Amer.,  Afr.,  As.,  Indomal, 
one  small  group  austr.-bicentr. 


232  C.   SKOTTSBERG 

Thouvii  (Schwaegr.)  Mont.  S.  Chile-Fueg.  Belongs  to  a  group  of  4sp.  (3  Chile, 
I   N.  Zeal.).— Mf. 

Rhacopilaceae. 

Rhacopilum  I'alis.  (51).  Pantrop.,   mainly  southern,  south  to  X.  Zeal. 

fcDiaudeziiDiuin  Card.   S.   Chile.   Related  to  R.  toinentosum  (Sw.)  Brid.  (trop.- 
subtrop.  Amer.). — Mt,  Mf. 

Thuidiaceae. 

riiuidiion  Bryol.  eur.  (161).  All  over  the  world,  especially  in  humid  mountain  climates. 

Masafuerae   l^roth.    Related    to    7\  fuluastrum    (Mitt.)   Jaeg.   (Tristan  da  C, 
X.   Zeal,  but  not  included  in  210)  and  to  the  following  sp. — Mf. 

I'aldiviae  Broth.   S.   Chile.— Mt,  Mf. 

Amblystegiaceae. 

Sciayouiiuni  Mitt.  (22).   Mainly  austral,  the  majority  southern  S.  Amer.,  i  Cape,  2 
Ciiina,   2   1^.  Austral.,    i  X.  Zeal.;  the  single  X.  Amer.  sp.  forms  a  separate  section. 
Sect.  Aloma  6  (i    Boliv.,  2   Fueg.,  2  \l.  Austral.,   i   X.  Zeal.). 
pacJiyluDia  (Mont.)  Par.  S.  Chile-W.  Patag.— Mt,  Mf. 

Hypnaceae. 

Stcrcodoji  Mitt.  (7).   2   Mex.,    i    Chile,  4  mountains  of  Asia. 
LccJilcri  (C.   M.)  Mitt.   Centr.   Chile  to  W.  Patag.— Mt. 
Isoptervi^iuui  Mitt.  (170).   Widely  distributed,  preponderately  trop.-subtrop. 
fernandezianutn  Broth.  Related  to  /.  teuerum  (Sw.)  Mitt,  (neotrop.). — Mt. 

Sematophyllaceae. 

RhaphidostegiiiDi  (Bryol.  eur.)  De  Xot.  (106).  Subcosmopol.,  trop.  and  temp.  The 
C'hilcan  species  sometimes  regarded  as  belonging  to  a  separate  genus  RJiapJiido- 
rJiyiichiion. 

Masafuerae  Ikoth.   Similar  to  R.  cyparissioides  (Hornsch.)  l^esch.  (Braz.). — Mf. 

aberrans  Broth.  Similar  to  R.  callidion  (Mont.)  Jaeg.  (S.  Chile  to  W.  Patag.). — 
Mt,   Mf. 

cacspifosuiii  (.Sw.)  Jaeg.   \V.   Ind.,  trop.-subtrop.   S.  Amer. — Mt,  Mf. 

caespitosoides   Broth.   Related  to  the  former. — Mf. 

brachycladulum  J^roth. — Mt. 
Rij^odiion   Kzc.  (19).  Trop.-subtrop.   7\mer.,   2   Afr. 

toxcniuni  (Schwaegr.)  Schim[).  Trop.   S.  Amer.  to  W.  Patag. — Mt,  Mf. 

arboresicus  (C.   M.)  Broth.   S.   Chile-\V.   Patag.— Mt,  Mf. 

hylocoDiioidcs  Card,   et  Broth.  Patag. — Mf. 

robustutn  l^roth.      Mt. 

Looseri    Thcr.   Related  to  A',  gracile  Ren.  et  Card.  (Costa  Rica). — Mt. 

tcDnarix  C.   M.  (elegantulum   Card.).   S.   Chile-W.  Patag. — Mt. 
R/iyuc/iostf^i^iian   Ikyol.   eur.   (130).   Temp.-subtrop.,   almost  cosmopol. 

coiuplcvium  (Mitt.)  Jaeg.   Centr.   Chile. — Mt,   Mf. 

toiuifoliuiu  (Hedw.)  Jaeg.   Urug.,  S.   Chile,  E.  Austral.,  Tasm.,  X.  Zeal.— Mt. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  233 

Catagoniopsis  Broth.  Monotypical,  related  to   Catagonium  (S.  hemisph.) 
berteroava  (Mont.)  Broth.  Centr.  Chile. — Mt. 

Hypnodendraceae. 

Hypnodendron  Lindb.  (28).   Chile  (2),  IndomaL,  Austral.,  Polyn.,  Hawaii. 
microstictum  Mitt.  Chile. — Mf. 

Polytrichaceae. 

Oligotrichmn  Lam.  et  DC.  (13).  Widely  scattered:  2  W.  N.  Amer.,  4  Chile;  single 
sp.  Braz.,  trop.  Andes,  Arct.-alp.,  Eur.,  bor.-circump. 

caiialiculatum  (Hook.)  Mitt.  var.  The  typical  sp.  in  S.  Chile. — Mf. 
Psilopilum  Brid.  (17).  A  bipolar  genus,  tricentric  in  the  S.  hemisph. 

antarcticu7n  C.  M.  Boliv.,  Fueg.,  Falkl.,  S.  Georgia,  Kerguel. — Mf. 
Polytrichadelphus  (C.  M.)  Mitt.  (22).  Centering  in  the  Andes  (17),  i  W.  N.  Amer., 
I   Braz.,  2  Magell.,   i   subant.-bicentric. 

magellanicus  (L.)  Mitt.  Patag.,  Fueg.,  Falkl.,  E.  Austral.,  Tasm.,  N.  Zeal.,  Auckl. 
and  Campb.  Is. — Mf. 
Dendroligotrichum  (C.  M.)  Broth.  Monotypical. 

dendroides  (Brid.)  Broth.  Peru,  S.  Chile  south  to  Fueg.,  N.  Zeal. — Mf. 

The  geographical  distribution  of  the  mosses  is  less  well  known  than  of  the 
vascular  plants,  and  some  of  the  species  now  considered  to  be  endemic  in  Juan 
Fernandez  will  perhaps  be  discovered  on  the  mainland.  No  specialist  ever  visited 
the  islands,  where  most  likely  further  species,  endemic  or  known  from  elsewhere, 
will  be  found.  Species  now  only  recorded  from  Masatierra  may  be  found  onMasafuera, 
and  vice  versa. 

The  131  species  of  mosses  hitherto  recorded  from  Juan  Fernandez  belong  to  65 
genera  (2:  i),  all  known  from  elsewhere;  48  species  (36.6%)  are  endemic,  a  high 
figure  in  a  spore-bearing  group,  higher  than  for  the  Pteridophytes  (34%).  On  Masa- 
tierra 84^  were  collected,  50  of  these  (38.1  %)  not  found  on  Masafuera,  where  81 
were  recorded,  of  which  47  (35.9  %)  are  restricted  to  this  island;  34  (26  %)  are  known 
from  both  islands.  No  mosses  have  been  reported  from  Santa  Clara. 

The  mosses  are  less  evenly  distributed  than  the  ferns,  presenting  a  higher 
degree  of  local  endemism,  but  this  may  be  due  to  insufficient  knowledge  of  their 
distribution.  Endemics  are  more  local  than  non-endemics;  of  the  former  (48),  24 
(50%)  are  only  known  from  Masatierra,  and  16  (33.3  %)only  from  Masafuera,  8  (16.7  %) 
being  found  on  both  islands;  of  the  non-endemics  (83),  26  inhabit  Masatierra  (31.3  %), 
31  Masafuera  (37.4%)  and  26  (31.3  %)  both  islands.  Endemics  are  proportionately 
more  numerous  on  Masatierra,  where  32  species  or  38  %  are  endemic;  the  figures  for 
Masafuera  are  24  and  29.6.  It  is  remarkable  that  all  the  8  species  of  Thamnium 
are  endemic  on  Masatierra. 

For  two  reasons  it  is  a  difficult  task  to  segregate  with  sufficient  accuracy  the 

1  Brotherus  p.  420  records  Rhacomitrium  subnigritum  from  both  islands,  but  the  locality 
quoted  for  Masatierra  is  situated  on  Masafuera. 


234 


C.   SKOTTSBERG 


various  geographical  elements  of  which  the  flora  is  made  up:  many  of  the  non- 
endemic  species  have  a  more  or  less  wide  distribution,  and  the  accurate  systematic 
position  of  tlie  endemic  ones  is,  in  many  cases,  uncertain  or  quite  unknown;  where 
no  information  was  given  by  their  author  and  nobody  has  studied  them  after  him, 
I  have  not  ventured  to  find  a  place  for  them.  Thus  1 1  species  had  to  be  left  out, 
bringing  the  number  down  to  120,  on  which  the  percentages  have  been  calculated. 
Even  so  it  stands  to  reason  that  the  arrangement  below  cannot  be  definite  because 
many  doubtful  points  remain  to  be  cleared  by  the  bryologist. 

As  to  the  83  non-endemic  species  the  main  thing  is  whether,  regardless  of 
their  total  distribution,  they  have  been  found  in  S.  America  or  not;  in  consequence 
of  the  geographical  position  of  Juan  Fernandez  they  must  be  referred  to  one  of 
the  American  groups  I-III.  Xo  less  than  25  %  are  austral  or  subantarctic  and 
bicentric;   their  origin,  if  antarctic  or  not,  will  not  concern   us  here. 

The  geographical  elements. 

I.   Andine-Chilean  element. — 77    (64.2%). 

ci.  luidemic  species  (15):  Dicranoloma  fernandezianum  and  capillifolioides,  Fissi- 
dens  pycnotylus,  Didymodon  calymperidictyon  and  linearis,  Ulota  fernandeziana, 
Philonotis  glabrata,  Ptychomnium  falcatulum,  Distichophyllum  fernandezianum, 
Pterygo[)hyllum  tenuinerve,  Eriopus  leptoloma  and  grandiretis,  Thuidium  Masa- 
fuerae,   Rhaphidostegium  aberrans,  Rigodium  robustum. 

b.  Also  known  from  Chile,  rarely  extending  to  the  extreme  south  {57): 
Pleuridium  Robinsonii,  Ditrichum  affine  andlongisetum,  Hymenostomumkunzeanum, 
Amphidium  cyathicarpum,  Dicranoloma  capillifolium,  Menziesii  and  nigricaule, 
Campylopus  truncatus,  Thysanomitrium  Richardi  and  leptodus,  Eucamptodon 
perichaetialis,  Fissidens  rigidulus,  leptochaete,  maschalanthus  and  asplenioides, 
Tortula  scabrinervis  and  flagellaris,  Rhacomitrium  loriforme  and  con  volutum,  Zygodon 
intermedins  and  Menziesii,  Stenomitrium  pentastichum.  Macromitrium  hymeno- 
stomum,  Ikyum  Lechleri,  Leptostomum  Menziesii,  EustichiaPoeppigii,Rhizogonium 
nmioides,  Hartramia  aristata,  Philonotis  krauseana  and  vagans,  Rhacocarpus  Hum- 
boldtii,  Dcndrocryphaea  cuspidata,  Lepyrodon  parvulus  and  implexus,  Ptychomnium 
subacicularc  and  j)tyciu)carpum,  Weymouthia  mollis,  Leptodon  Smithii,  Porotham- 
nium  arbusculans,  Lamprophyllum  splendidissimum,  Lopidium  concinnum,  Hypo- 
|)tcrygium  Thouini,  Khacopilum  fernandezianum,  Thuidium  Valdiviae,  Sciaromium 
pachyloma,  Stereodon  Lechleri,  Rigodium  toxarium,  arborescens,  hylocomioides 
and  tamarix,  Rhynchostegium  complanum  and  tenuifolium,  Catagoniopsis  berte- 
roana,  Hypnodendron  microstictum,  Oligotrichum  canaliculatum,  Dendroligotrichum 
dendroides. 

c.  Cosmopolitan  (5):  Ceratodon  purpureus,  Gyinnostomum  calcareum,  Cam- 
pylof)us  introflexus,  Rhacomitrium  lanuginosum,  l^\inaria  hygrometrica. 

II.    Subantarctic-Magellanian   element. ^ — 13    (10.8%). 

All  non-endemic,  found  in  the  far  south,  many  going  north  to  the  latitude 
of  Valdivia  or  even  farther;  several  occur  on  the  P'alkland  Is.,  South  Georgia  or 
other  subantarctic  islands,  and  not  few  reappear  in  New  Zealand,  etc. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  235 

Oncophorus  fuegianus,  Dicranoloma  Billardieri,  Rhacomitrium  subnigritum, 
symphyodontum  and  striatipilum,  Rhizogonium  Novae  Hollandiae,  Bartramia  patens, 
Philonotis  scabrifolia,  Pterygophyllum  anomalum,  obscurum  and  denticulatum, 
Psilopilum  antarcticum,  Polytrichadelphus  magellanicus. 

III.  Neotropical  element. — 14   ( r  i .  7  %). 

Recorded  from  tropical  America  but  not  from  Chile,  or  related  to  tropical 
species. 

a.  Endemic  (8):  Campylopus  subareodictyon,  Leptodontium  fernandezianum, 
Bryum  fernandezianum,  Pinnatella  macrosticta,  Isopterygium  fernandezianum,  Rha- 
phidostegium  Masafuerae  and  caespitosoides,  Rigodium  Looseri. 

b.  Non-endemic  (6):  Campylopus  areodictyon,  Mielichhoferia  longiseta,  Anacolia 
subsessilis,  Lepyrodon  tomentosus,  Porothamnium  fasciculatum,  Rhaphidostegium 
caespitosum. 

IV.  West  Pacific  element.— 13   (10.8%). 

Endemic  species,  allied  to  S.W.  Pacific  species  (Australia,  Tasmania,  New  Zea- 
land etc.)  but,  as  far  as  known,  not  to  S.  American  species:  Macromitrium  fer- 
nandezianum and  Masafuerae,  Cyptodon  crassinervis,Distichophyllumsubelimbatum 
and  assimile,  Thamnium  rigidum,  latinerve,  Caroli,  Ingae,  crassinervium,  probosci- 
deum,  assimile  and  confertum. 

V.   Atlantic   element. — 3   (2.5  %). 

a.  Endemic:  Fissidens  crassicuspes. 

b.  Not  endemic:  Trichostomum  brachydontium,  Campylopus  polytrichoides. 

Endemic  species  of  unknown  position:  Dicranella  costata,  Campylopus  aberrans 
and  blindioides,  Fissidens  fernandezianus,  Grimmia  phyllorhizans,  Ptychomitrium 
fernandezianum,  Macromitrium  saxatile,  Bartramia  fernandeziana,  Breutelia  Masa- 
fuerae, Neckera  rotundata,  Rhaphidostegium  brachycladulum. 

The  dominance  of  a  South  American  element  is  self-evident;  groups  I  to  III 
make  up  86.7  %.  It  is  hard  to  draw  a  line  between  I  and  II;  the  Magellanian  species 
are  supposed  to  have  come  from  the  far  south,  and  theirs  is  a  more  southern  area, 
but  many  of  the  species  referred  to  I  ^  may  have  had  the  same  history  though, 
at  present,  they  do  not  reach  so  far  south. 

The  Atlantic  element  is  artificial.  Fissidens  crassicuspes  is,  if  Brotiierus  is  right, 
related  to  a  species  that  has  its  nearest  station  on  Madeira,  but  the  genus  is  a  very 
large  one,  and  another  connection  may  be  found.  Campylopus  polytrichoides  \'s>  diXv 
Atlantic  species  with  its  nearest  locality  on  Madeira;  its  presence  on  Masatierra 
is  indeed  surprising.  Trichostomum  brachydontium  is  scattered  over  half  the  globe, 
with  its  nearest  stations  in  Macaronesia,  but  it  extends  not  only  to  and  beyond 
the  Mediterranean  region,  but  turns  up  on  the  island  of  Reunion,  in  Japan  and 
on  New  Zealand.  Have  we  to  do  with  isolated  remnants  of  a  once  more  continuous 
area,  or  is  it  still  to  be  discovered  in  other  places.?  Is  it  a  bipolar  species.? 


236 


C.   SKOTTSBERG 


Of  the  species  left  aside  for  the  present,  Dicrmiella  costata  (many  in  S.  Amer.), 
PtvchomitriuDi  fevnaudeziamini  (other  sp.  in  the  Andes),  Macroniityiiim  saxaiile  (many 
in  S.  Amer.),  Bartrauiia  fernandcziima  and  Breutelia  Masafuerae  (many  austral  sp.) 
may  turn  out  to  belong  to  an  American  element.  The  occurrence  in  Masatierra 
of  8  endemic  species  of  Thatiniium  is  astonishing,  for  not  one  is  quoted  for  South 
America,  whereas  related  species  are  found  in  Australia-New  Zealand,  Oceania  and 
Malaysia;  for  this  reason  I  have  referred  the  species  in  Masatierra  to  the  Pacific 
element. 

IV.  Hepaticae. 

Xo  handbook  equal  to  Brotherl's'  comparatively  modern  account  of  the  mos- 
ses exists  of  the  Hepaticae.  ScillFKNERs  treatment  of  this  group  in  the  1st  edition  of 
Xatiirl.  Pflanzenfamilien  is  too  antiquated  to  be  of  much  use.  Much  important  infor- 
mation is,  however,  found  in  Hkrzog's  work  [i2g)  as  well  as  in  Domin's  paper  (7^), 
and  also  in  this  case  Dr.  Pkrsson  kindly  helped  me,  but  to  search  the  voluminous 
s[)ecial  literature  of  the  last  thirty  years  was  not  to  be  thought  of.  The  number  of 
species  given  is,  in  many  cases  at  least,  too  low,  but  I  don't  think  this  matters 
very  much. 

The  following  list  is  based  on  PlVAXs'  and  Herzog's  papers  (gj,  ijo)  with 
a  few  alterations  (/-?/).  Several  species  credited  to  Juan  Fernandez  by  Stephani 
( i^j-,^)  but  not  mentioned  by  Herzog  are  included  here;  some  of  them  are,  perhaps^ 
identical  with  other  species. 

Marchantiales. 

Plagiochasuia  Lehm.  et  Lindenb.  (about  20).  Mostly  trop.-subtrop. 

ntpcstrc  (Forst.)  St.  Cosmopol.,  also  Chile. — Mt,  Mf. 
Rehoulia  Raddi  (i). 

Iieinispliaerica  (L.)  Raddi.   Cosmopol.,  also  Chile. — Mt,  Mf. 
LiDiularia  (Mich.)  Adans.  (i). 

cruciata  (L.)  Dumort.  S.  Amer.,  also  Chile,  Medit.  Fur.,  Atl.  islands,  Afr.^ 
Austral.— Mt,   Mf. 

Maycliautia  L.   (about   50).   Subcosmopol.,  numerous  trop. 

polyifwypha   L.   Cosmopol.,  also  Chile.- — Mt,   Mf. 

hertcrpiDia  Lehni.  et  Lindenb.  Chile  south  to  I^'ueg.,  Falkl.,  St.  Helena,  S.  Afr.,, 
Marion   I.,   Kerguel.,  Austral.,  Tasm.,  N.  Zeal. — Mt,  Mf. 

foliacca  Mitt.   S.   Chile,  Tasm.,   N.   Zeal.— Mt,   Mf. 

Metzgeriales. 

Riccaydiii  )^.V .    (iray  (140-150,  but  much   higiier  figures  are  given).  N.   temp.  6^ 
trop.   Amer.  43,   Afr.    14,   trop.   As. -Oceania   53,   austr.-subantarct.   35. 

fucgiensis  Massal.   S.   Chile  to  Fueg. — Mf. 

brcniramosa  (St.)   I^vans.    Falkl. — Mt,   Mf. 

adglutinata  Fvans. — Mt,   Mf. 

jusularis  SchitTn.   St.   Paul  and   New  Amsterd.   Is. — Mt. 

pariabilis  Fvans.   S.   Chile. — Mt. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  237 

leptostachya  Evans.  Related  to  the  former. — Mt. 

nudimitra  (St.)  Evans.  S.  Chile  to  W.  Patag. — Mf. 
Metzgeria  Raddi  (about  50).  Well  developed  trop.,  some  widely  spread  N.  hemisph. 
in   oceanic  climates. 

decrescens  St.  S.  Chile  to  W.  Patag. — Mf. 

decipiens  (Massal.)  Schiffn.  et  Gottsche.  Centr.  Chile  to  W.  Patag.,  Austral., 
N.  Zeal.,  Antipodes  Is.— Mt,  Mf. 

multiformis  Evans.  Closely  related  to  the  former. — Mt,  Mf. 

violacea  (Ach.)  Dumort.  Centr.  and  S.  Chile,  Fueg.,  Argent.,  W.  and  S.  Afr.  N. 
Zeal.,  Antip.  Is.  Brought  to  M.  decipiens  as  a  variety  by  HoDGSON  [136.  278). — Mf. 
Hymenophytum  Dumort.  (5).  S.  Amer.,  Ind.,  Austral.,  Tasm.,  N.  Zeal.,  Melanes. 

flabellatum  (Labill.)  Dumort.  Colomb.,  Chile,  Austral.,  Tasm.,  N.  Zeal.,  N.  Caled., 
Fiji.— Mf. 
Symphyogyna  Nees  et  Mont.  (39).  Trop.  and  austral. 

circinata  Nees  et  Mont.  Centr.  and  S.  Chile. — Mt,  Mf. 

Hochstetteri  Nees  et  Mont.  Falkl— Mt,  Mf. 

hymenophylluni  (Hook.)  Nees  et  Mont.  Trop.  Amer..^  Chile,  Austral.,  Tasm., 
N.  Zeal. — Mt,  Mf.  Probably  identical  with  vS.  podophylla  (Thunb.)  Mont,  et  Nees 
from  Centr.  and  S.  Africa  (S.  Arnell,  ms.). 

Pallavicinia  S.  F.  Gray  (about  25).  Trop.-subtrop.,  Colomb.,  Indomal.;  Kerguel., 
N.  Zeal. 

xiphoides  (Tayl.)  St.  N.  Zeal.— Mf. 
Monocloa  Hook.  (2,    i   trop.  Amer.). 

Forsteri  Hook.  S.  Chile-W.  Patag.,  N.  Zeal.— Mt,  Mf. 
Androcryphaea  Nees  (i ;  Noteroclada  Tayl.). 

confluens  (Tayl.)  Nees.  Mex.;   Colomb.-Chile,  south  to  Fueg.,  Braz.,  Argent., 
Falkl.,  Kerguel.— Mf. 
Fossombronia  Raddi  (26).  Trop.-subtrop.,  extending  into  cooler  regions. 

fernandeziensis  St. — Mt. 

Anthocerotales. 

Anthoceros  L.  (about   140).  Mainly  trop. 

Skottsbergii  (St.  ex  p.)  Evans.— Mt,  Mf. 
Megaceros  Campb.  (about  40).  Mex.,  W.  Ind.,  S.  Amer.  south  to  Chile,  Reunion, 
Indomal.-N.  Zeal.,  Oceania. 

ftiegiensis  St.   W.  Patag.-Fueg.   Related  to  neotrop.  sp. — Mt,  Mf. 

Jungermanniales . 

Solenostoma  Mitt.  (30).  Subcosmop.,  but  now  partly  referred  to  Aplozia  Dum.,  partly 
to  Plectocolea. 

crassuhim  (Mont.)  St.   S.  Chile.— Mf. 

obtusiflorum  St. — Mt. 

rostratum  St.— Mf. 
Jmnesoniella  Spruce  (20).  Mainly  trop.-subtrop.  (i  Engl.)  but  also  important  in  the 
austral  zone. 


238  C.   SKOTTSBERG 

coloyata  (Lchni.)  Spruce.  Centr.  Amer.,  S.  Amer.  south  to  Fueg.,  Falkl.;  Tristan 
da  C,  S.  Afr.,  Marion  I..  Kerguel..  \\.  Austral.,  Tasm.,  N.  Zeal,  Auckl.,  Campb.  and 
Antijiodes  Is. — Mt,   Mf. 

)nalui}ia  St.   Fall<l. — Mt. 

oeuops  St.   S.  Chile,  south   to   Fue^^.,   S.  Georgia. — Mt. 

gyamiiflora  (Lehni.  et  (k)ttsche)  Spruce.  Colomb.-Chile,  south  to  Fucg.;  Tristan 
da  C,   Marion   I.,  Tasni.^ — Mt. 
Auastrop/iyliinn  Xees  (31).   Very  widely  distributed. 

leucocephaliini  (Tayl.)  Spruce.   X^enez.,  Peru,   Fueg. — Mf. 
AcrobolbHS  Xees  (11).   Essentially  austral:   2   S.  Amer.,    i    subantarct.,  i  Himal.,  7 
Austral.,   X.   Zeal.,   X.   Caled. 

cxcisus  (Mitt.)  Schiffn.   W.   Patag.,  Fueg.,   Kerguel.  — Mf. 
A)iasirepta  (Lindb.)  Schiftn.  (4).  2  southern  S.  Amer.,  i  Alaska-P^ur.-E.  As.-Hawaii, 
I    Himal. 

bifida  St.  S.   Chile:   Valdivia,  Magell.— Mt,  Mf. 
Plai^iochila  Dumort.  (about  1000).  Very  few  bor.  zone;  trop.,  particularly  neotrop., 
but   well   represented  southern   S.   Amer. 

i:;aya}ia   Gottsche.   S.  Chile.  —  Mt. 

fasiiata  St.   Chiloe.— Mt,  Mf. 

hyadesiaua  Bcsch.   et  Massal.   S.   Chile,  P\ieg.— Mt,  Mf. 

deform  if olia  St.   W.   Patag.— Mf. 

chilocusu  St.   S.   Chile  to  W.  Patag.— Mf. 

rcctani!;ulata  St.   W.  Patag. — Mf. 

rciiiotidfiis  St.   S.   Chile,   Magell.— Mf. 

fuscobriinnea  St.  Almost  too  closely  related  to  P.  riibescens  Lehm.  et  Lindenb 
(Syn.   l\  chilcusis  St.  according  to  Hkkzog  770).— Mt,  Mf. 

pudftcusis  St.   Chiloe. — ^It. 

hoDiomalla  St.   W.  Patag. — Mf. 

mcsiivia  Lindenb.   S.   Chile  south   to  Magell. — Mt. 

riparia  St.   W.   Patag.— Mt,  Mf. 

squarrosa  St.   W.   Patag.— Mt,  Mf. 

robusta  St.   W.   and   \\.   Patag.,  Fueg. — Mf. 

i'/ahj  Hook.  f.  et  Tayl.  Inieg.— Mf. 

Xo/arisii  Lehm.   Falkl.  — Mt,   Mf. 
'ryliouDitJius   Mitt.   (25).   .S.   hemisph.,   best  represented  Australia;    i    \V.   Ind. 

liwbalus  St.    W.    I'atag.-- -Mt. 

silvaticus  St.      MW 

densiretis   Her/..      Mf. 

bilobatus  St.  —  Mt. 
My/in  S.   V .  (ira>'  (40-50).  Tiie  majority  neotrop.  mountains  (Mex.,  W.  Ind.,  Centr. 
and   S.   Amer.   south   to  Magell.),  4  i\fr.,    i    .S.   Georgia. 

rrpe?is  (Mitt.)   I  lerz.    Magell.,   X.   Zeal.— Mf. 

t'uscovircus  (Tayl.)  St.   See   /  ,'0.714.   Centr.   and  S.  Chile  to  Fueg. — Mf. 

li^s^ulata  (St.)   Ilerz.   \V.   Patag.— Mt. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  239 

Lophocolea   Dumort.    (250-300).    World-wide;    main  concentrations  subantarct.  S. 
Amer.  (60),  S.  As.-Oceania  (35),  and  Austral.-N.  Zeal.  (70), 

rotu7idifolia  St.  W.  Patag.,  Fueg.,  Falkl. — Mf. 

fertiandeziaria  St.   Centr.  Chile  (Coquimbo). — Mt,  Mf. 

pallidevirens  (Tayl.)  St.  Centr.  (Coquimbo)  and  S.  Chile  to  W.  Patag.,  Fueg., 
Falkl.,  Marion  I.— Mt,  Mf. 

papulosa  St. — Mt. 

attenuata  St.   S.   Chile  to  W.  Patag.— Mt,  Mf. 

textilis  Tayl.  W.  Patag.-Fueg.,  Falkl.— Mf. 

diver genticiliata  St.  S.   Chile,  W.  Patag.,  Fueg.— Mt,  Mf. 

mtiricata  Nees.  W.  Ind.,  S.  Braz.,  Centr.  and  S.  Chile  to  W.  Patag.,  S.  Afr., 
Reunion,  Java,  N.   Guin.,  E.  Austral.,  N.  Zeal.— Mf. 

angulata  St. — Mt. 

cJiilensis  St.  Chile. — Mf. 

submuricata  Herz.  Near  muricata,  but  also  very  close  to  L.  fragrans  Tayl. 
(atl.-mediterr.,  130.  720). — Mt,  Mf. 

Ckiloscyphus  Corda  (150-200).  Trop.-subtrop.,  best  developed  S.  hemisph.  (about 
50  Austral.-Tasm.-N.  Zeal.,  8  subantarct.  Amer.). 

integrifolius  Lehm.  et  Lindenb.   Centr.   Chile  to  Fueg. — Mt,  Mf. 

lobatus  St.  W.  Patag.,  Fueg. — Mf. 
Saccogyna  Dumort.  (9—10).  Braz.   2,  Chile   i,  Eur.-Macaron.  i,  Indomal.-Polyn.  2, 
Tasm.-N.  Zeal.  3;   Hawaii.? 

sqiiarristipula  Herz.  S.  Chile  to  W.  Patag. — Mt. 
Marsupidium    Mitt,    (about   40.?).  Austr.-circump.,  subantarct.  Amer.,  Austral.-N. 
Zeal. 

piljferiim  St.  S.   Chile,  E.  Austral,  N.  Caled.— Mt,  Mf. 
Adelanthiis  Mitt.  (7).  Widely  scattered:  W.  Ind.,  Ecuad.,  Chile,  Eur.,  St.  Helena. 

sphalerus  Hook.  f.  et  Tayl.   W.  Patag.-Fueg. — Mt. 
Bazzania  S.   F.  Gray  (230).  World-wide. 

cerina  (St.)  Fulford.  S.  Chile,  W.  Patag.,  Falkl,  Tristan  d.  C— Mt,  Mf. 

peruviana  (Lehm.  et  Lindenb.)  Trev.  Peru,  Chile  to  W.  Patag. — Mt. 
Lepidozia   Dumort.   (about  200).   World-wide,  but  few  north  temp.,  most  numerous 
trop.-subtrop.,  many  austr. -subantarct.  (70-80). 

sejuncta  (Angstr.)  St.  Mex.,  W.  Ind.,  S.  Braz.,  Azor.,  W.  and  S.  Afr.,  Tasm. 
— Mf. 

bicuspidata  Massal  Centr.   Chile,  W.  Patag.,  Fueg. — Mt. 

pseudozoopsis  Herz.   S.  Chile  to  W.  Patag.  (7^7.51). — Mt. 

fernandeziensis  St.  S.  Chile  (/jo.  725).  Near  L.  plumulosa  and  Lindenb ergii  St. 
(N.  Zeal).— Mt. 

plumulosa    Lehm.  et  Lindenb.  S.   Chile  to  W.  Patag.  and  Fueg.,  Falkl,  N. 
Zeal,  Auckl  Is.,  Antipodes  Is. — Mt. 

fragillima  Herz.  Related  to  Chilean  species. — Mt. 

disticha  St.— Mt. 

Jacquemontii  St.  Centr.  Chile-Magell,  Fueg. — Mt,  Mf. 


240  C.  SKOTTSBERG 

Uerberta  S.   F.   Gray  (15).   Widely  scattered. 

riiuciuata  (Tayl.)  Herz.   S.   Chile  to  W.   Patag.— Mf. 
LepicoUil  Dumort.   (5).   Scattered  S.  hemisph. 

ochyoleucii  (Spreng.)  Spruce.   S.   Braz.,  S.   Chile  to  Fueg.,  Cape. — Mt,  Mf. 
Lepidolaeua  Dumort.  (12).   S.   temp,  and  cold  zones. 

mai:;ella)uca  (Lam.)  Schiffn.  S.  Chile  to  Fueg.,  E.  Austral.,  Tasm.,  N.  Zeal. 
— Mf. 

Trichocolfa    Dumort.    (32).    i    l^or.  zone,    19  trop.  Amer.,   7  trop.  As.-Oceania,   5 
Austral. -N.  Zeal. 

opposita  St.  Near    7\  australis  St.  (X.  Zeal.). — Mt. 

rettiallata  St.   S.   Chile  to  W.   and  E.  Patag.— Mt,  Mf. 
Scliistochila    Dumort.  (83).   P^ssentially  southern:   austral-subantarct.  (20),  Afr.  (7), 
Indomal.-Oceania  (35),  Austral. -N.  Zeal.  (21). 

berteroana  (Hook.)  St.   S.   Chile.— Mt,   Mf. 

Skottsbergii   St.    Related    to    5.  stratosa   (Mont.)    St.    (S.   Chile  to  Fueg.). — 
Mt,   Mf. 

pachyla  (Tayl.)  St.   W.  Patag.-MagelL— Mf. 

splacliuophylla  (Tayl.)  St.  W.  Patag.-Fueg.,  N.  Zeal. — Mf. 
Balantiopsis  (Xees)  St.  (17).    i   Braz.,   i    Oueensl.,  the  remainder  austr.-subantarct. 

caucellata  (Xees)  St.   S.   Chile  to  W.  Patag.— Mf. 

cJiilensis  St.  S.   Chile  to  W.  Patag. — Mf. 

purpiirata  Mitt.   S.   Chile.  Determination  uncertain. — Mf. 

hians  Ilerz. — Mt. 

lancifolia  St. — Mt. 
Radula  Dumort.  (220).   Mainly  trop.-subtrop.  Bor.  zone  (Eur.,  N.  Amer.,  7),  trop.- 
subtroj).  Amer.  (66),  Afr.  (37),  Indomal.-Oceania  (69),  Austral.-X.  Zeal.  {29),  sub- 
antarct.  (15). 

hastaia  St.   S.   Chile  to  luieg.- Mf. 

niicyoloha  St.   S.   Chile. — Mt,  Mf. 

Duscnii  St.   S.   Chile  to  \V.  Patag.— Mf. 

Mittvnii  St.   I^'alkl.— Mt,  Mf. 

Madothcca    Dumort.    (153).    Trop.-subtrop.    Amer.    (45),    trop.    As.-Oceania    [6"]), 
Austral. -\.   Zeal.   etc.   (12). 

chiloLsis    Lchm,    et    Lindenb.    var.  feriiandeziensis    Herz.    Centr.    Chile    (Co- 
(juimbo;   main  sp.   S.   Chile  to  W.   I'atag.). — Mt,  Mf. 

subsquarrosa  Xees  et  Mont.   S.   Chile  to   Fueg. — Mt. 
I'fulliuna   kaddi  (;oo-6oo).   World-wide,  but  mainly  trop.  and  southern:  X.  Amer. 
(35)- 

lickloui  Spr.  (crassa  Herz.).   Ccntr.   Chile  to  W.  Patag.;  W.  and  S.  Afr.— Mt. 

cluloisis  St.  S.   Chile  to  W.   Patag. — Mf. 

lobulaia   Hook.   f.   et   Wils.   W.   Patag.,   P\ieg. — Mf. 

})uio;cllanica    (Sprcng.)  Web.   et  Xees.   Centr.   and  S.   Chile  to  Fueg.,  Tristan 
da  C.   Tasm.,   Campb.   I. — Mt,   Mf. 

stipatiloba  St.   Centr.   Chile  to  W.   Patag.— Mt. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  24I 

Lopholejeunea   Spruce   (74).  Trop.  and  austral;  trop.  Amer.  (13),  trop.  Afr.  (17), 
trop.  As.-Oceania  (38),  Austral.-N.  Zeal.  (6). 
spinosa  St. — Mt,  Mf. 

Brachiolejeioiea  Spruce  (65).  Trop.  Amer.  (26),  Chile  (i),  Afr.  (10),  trop.  As.-Oce- 
ania (22),  Austral.-N.  Zeal.  (6). 

spruceaita  (Massal.)  St.  S.  Chile-Magell. — Mf. 
Harpalejeunea  Spruce  (57).  Trop.  Amer.  (36),  Chile  (2),  Afr.  (2),  As.-Oceania  (8), 
Austral.-N.  Zeal.  (9). 

oxyota  (Mont.)  St.  Centr.  (Coquimbo)  and  S.  Chile  to  W.  Patag,  Tristan  da  C. — Mf. 

setifera  (St.)  Herz.  Magell. — Mf. 
Strepsilejeunea   Spruce   (47).  Trop.  Amer.,  south  to  Chile  (19),  Afr.  (6),  As.  (7), 
Austral.-N.  Zeal.  (7),  subantarct.  (8). 

acuminata  (Lehm.  et  Lindenb.)  St.  S.  Chile. — Mt. 

squarrosula  Herz. — Mf. 

macroloba  Herz. — Mf. 
Siphonolejeunea  Herz.  (i). 

nudicalymia  Herz.  Centr.  Chile  (Coquimbo,  iji.  65). — Mt,  Mf. 
Lejeiinea  Lib.  (190).  Pantrop.,  scarce  toward  the  south. 

reticulata  Herz.  Related  to  Chilean  sp. — Mt,  Mf. 
Apha7iolejeunea  Evans  (13).  Trop.-subtrop. 

asperrima  St.  S.   Chile. — Mt. 

diaphana  Herz.  Centr.  Chile  (Coquimbo,  S.  Arnell  ms.). — Mt. 
Cololejeunea  Spruce  (about  80).  Mainly  trop. 

Skottsbergii  Herz.  Nearly  related  to  a  species  from  N.  Zealand. — Mt,  Mf. 
Colura  Dumort.  (30).  Trop.  to  subantarct.;  Amer.  (8),  Magell.  (2),  Eur.  (i),  Afr. 
(2),  Indomal.  (16). 

bulbosa  Herz.  W.  Patag.  (j?7.  66).— Mf. 

The  124  Hepaticae,  of  which  25  (20.2%)  are  endemic,  belong  to  47  genera 
(2.6:1);   27  are  thallose,  97  foliose.  None  have  been  reported  from  Santa  Clara. 

Of  the  27  thallose  species  21  have  been  recorded  from  Masatierra  and  23 
from  Masafuera;  4(15  %)  are  known  from  Masatierra  only,  7  (25  %)  restricted  to 
Masafuera,  16  common  to  the  two  islands.  Of  the  5  endemics  2  have  been  found 
on  Masatierra  only,  3  on  both  islands.  Endemism  is  stronger  on  Masatierra  (25  %) 
than  on  Masafuera  (13  %). 

Of  the  97  foliose  species  59  occur  on  Masatierra  and  6^]  on  Masafuera;  30 
(30.9  %)  are  restricted  to  Masatierra,  38  (39.2  %)  to  Masafuera,  29  (29.9  %)  are  found 
on  both  islands.  Endemic  species  20  (21.6%),  of  these  9  only  on  Masatierra,  5 
only  on  Masafuera  and  6  found  on  both  islands. 

Of  the  59  species  recorded  for  Masatierra  i  5  are  endemic  in  the  islands  (25.4  %); 
the  corresponding  figures  for  Masafuera  are  67,  1 1  and  16.4  %.  After  the  discovery 
of  Siphonolejeunea  on  the  mainland  of  Chile  there  is  no  endemic  genus  in  Juan 
Fernandez. 

Of  the  total  number  of  species,  124,  80  occur  on  Masatierra  and  90  on  Masa- 

i6  -  557857  The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.  Vol.  I 


242 


C.  SKOTTSBERG 


fuera;  of  these  20  {25  %)  and  14  (16.5  %),  respectively,  belong  to  the  endemic 
element;  34  species  are  restricted  to  Masatierra  (27.4%),  44  to  Masafuera  (35.5  %) 
and  46  (37.1  %)  shared  by  the  two  islands.  The  corresponding  figures  for  the 
endemic  species  are:  Masatierra  11  (44%),  Masafuera  5  (20%),  both  islands  9 
(36  %),  and  for  the  non-endemics  23  (23.2  %),  39  (39.4  %)  and  37  (37.4  %).  Of  the  99 
species  also  found  elsewhere  Masatierra  has  60  and  Masafuera  ']6.  This  island  is 
richer  in  non-endemic  species,  many  of  which   belong  to  higher  altitudes. 

Geographical  elements. 
I.  Andine-Chilean  element.  —  97  (78.3  %). 

a.  luulemic  (20):  Riccardia  adglutinata  and  leptostachya,  Metzgeria  multifor- 
mis, Solenostoma  obtusiflorum  and  rostratum,  Plagiochila  fuscobrunnea,  Tylunan- 
thus  silvaticus,  bilobatus  and  densiretis,  Lophocolea  papulosa,  angulata  and  sub- 
nuiricata,  Lepidozia  fragillima  and  disticha,  Schistochila  Skottsbergii,  Balantiopsis 
hians  and  lancifolia,  Strepsilejeunea  squarrosula  and  macroloba,  Lejeunea  reticulata. 

b.  Also  known  from  Chile  and  not  restricted  to  the  extreme  south  (73):  Mar- 
chantia  berteroana  and  foliacea,  Riccardia  fuegiensis,  variabilis  and  nudimitra, 
Metzgeria  decrescens,  decipiens  and  violacea,  Hymenophytum  flabellatum,  Sym- 
phyogyna  circinata  and  hymenophyllum,  Monoclea  Forsteri,  Androcryphaea  con- 
fluens,  Solenostoma  crassulum,  Jamesoniella  colorata  and  grandiflora,  Anastrophyl- 
lum  leucocephalum,  Anastrepta  bifida,  Plagiochila  gayana,  fasciata,  hyadesiana, 
deformifolia,  chilocnsis,  rectangulata,  remotidens,  pudetensis,  homomalla,  neesiana, 
riparia,  S([uarrosa  and  robusta,  Tylunanthus  limbatus,  Mylia  repens,  fuscovirens 
and  ligulata,  Lophocolea  fernandeziana,  attenuata,  divergenticiliata,  chilensis  and 
muricata,Chiloscyphusintegrifolius,Saccogynasquarristipula,Marsupidiumpiliferum, 
Bazzaniacerinaand  peruviana,  Lepidozia bicuspidata,  pseudozoopsis,  fernandeziensis, 
plumulosa  and  Jaccjuemontii,  Herberta  runcinata,  Lepicolea  ochroleuca,  Lepidolaena 
magellanica,Trichocolea  verticillata,  Schistochila  berteroana,  Balantiopsis  cancellata, 
chilensis  and  purj)urata,  Radula  hastata.  microloba  and  Dusenii,  Madotheca  chi- 
lensis and  subsfjuarrosa,  Frullania  hxklonii,  chilensis,  magellanica  and  stipatiloba, 
Brachiolejeunea  spruceana,  I  larpalejeunea  oxyota,  Strepsilejeunea  acuminata,  Sipho- 
nolejeunea  nudicalycina,  Aphanolcjeunea  asperrima,   Colura  bulbosa. 

c.  Cosiuopolitan  (4):  Plagiochasma  rupestre,  Reboulia  hemisphaerica,  Lunu- 
laria  cruciata,   Marchantia  polymorpha. 

II.  Subantarctic-Magellanian  element. — 18  (14.5  %). 

All  non-endemic:  Riccardia  breviramosa,  Symphyogyna  Hochstetteri,  Mega- 
ceros  fuegiensis,  Jamesoniella  maluina  and  oenops,  Acrobolbus  excisus,  Plagiochila 
data  and  Xotarisii,  Lophocolea  rotundifolia,  pallidevirens  and  textilis,  Chiloscyphus 
lobatus,  Adelanthus  sphaleriis,  Schistochila  pachyla  and  splachnophylla,  Radula 
Mittenii,   Frullania  lobulata,   I  larpalejeunea  setifera. 

III.  Neotropical  element. —  5  (4.0  %). 
a.   Fndemic  (4):   Fossombronia  fernandeziensis,   Anthoceros   Skottsbergii,  Lo- 
pholejeunea  spinosa,  Aphanolcjeunea  dia[)hana. 
h.  Not  endemic:   Lepidozia  sejuncta. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  243 

IV.  West  Pacific  element.— 3  (2.4  %). 

a.  Endemic  (2):  Trichocolea  opposita,  Cololejeunea   Skottsbergii. 

b.  Not  endemic:  Pallavicinia  xiphoides. 

V.  Known  from  St.  Paul  and  New  Amsterdam  Is. —  i  (0.8  %), 
Riccardia  insularis. 

The  American  element  is  in  absolute  dominance;  120  species,  96.8  %,  belong 
to  elements  I-III.  The  Pacific  group  is  very  small,  but  as  we  shall  find  later,  many 
species  are  austral  and  bicentric. 

Table  IV. 

Comparison  between  the  Angiosperms,  Pteridophytes  and  Bryophytes.  Figures  in  %. 

A,  Andine-Chilean;  M,  Magellanian;  N,  Neotropical;  P,  West  Pacific. 

A  M  N                P 

Angiosperms 46.9  10.2  12.9  17.7 

Pteridophytes 64.1  7.5  17.0                9.4 

Mosses 64.2  10.8  1 1.7  10.8 

Hepatics 78.3  14.5  4.0               2.4 

The  lower  percentage  of  Mosses  in  A,  as  compared  with  the  Hepatics,  may 
be  due  to  imperfect  knowledge  of  the  distributioh  and  to  the  fact  that  1 1  moss 
species  had  to  be  left  out  of  consideration. 

V.  Lichenes. 

What  I  have  said  above  when  dealing  with  the  Bryophytes  holds  good  of 
the  Lichens  in  a  still  higher  degree:  no  lichenologist  ever  visited  Juan  Fernandez, 
and  a  non-specialist  is  bound  to  pass  over  many  species;  collecting  crustaceous 
lichens  growing  on  hard  basalt  is  not  easy  and  generally  time-absorbing.  Our 
collection  is  listed  after  Zahlbruckner  [2(^6,  2^^),  where  also  the  species  found 
by  other  collectors,  but  not  found  by  us,  are  included.  Some  changes  had  to  be 
made;  for  instance,  Zahlbruckner  did  not  distinguish  between  Sticta  and  Pseu- 
docyphellaria\  on  my  request,  Dr.  R.  Santesson  kindly  revised  the  nomenclature. 
Some  determinations  in  these  and  other  genera  were  corrected  by  him  (see  vol. 
11.  886).  Later  he  went  over  the  proof-sheets  of  this  paper  and  added  further 
corrections.  I  thank  him  for  generous  assistance. 

It  is  to  be  regretted  that  ZAHLBRUCKNER  did  not  indicate  the  distribution 
of  the  non-endemic  species,  and  the  statements  in  his  Catalogus  are  too  general. 
I  have  tried  to  collect  further  data  from  a  number  of  papers  on  the  flora  of  Chile 
and  the  Subantarctic  and  Antarctic  regions  (7,  J2,  /j,  jj^,  iy6,  lyy,  182,  184,  2g^, 
J2y-j2p)  but  it  was  beyond  my  possibilities  to  search  the  entire  literature.  Dr. 
A.  H.  Magnusson,  who  kindly  took  the  trouble  to  go  over  my  list,  supplied  much 
useful  information, 

SC  =  Santa  Clara. 


244 


C.  SKOTTSBERG 


Verrucariaceae. 

I'errucaria  Schrad.  (about  2/0).   All   parts  of  the  world. 

microspora  Xyl.   X.   Amer.,  Chile,  l^^ur.,  Jap. — Mt. 
Microglaeua  Koerb.  (37).   luir.,  except   i    Hraz.,    i    Socotra,  and  the  following. 

fernandeziana  Zbr.^Mt,  SC. 

Dermatocarpaceae. 

X01  )>ia}tdi)ta  X\'l.   (i). 

piilchella  Xyl.   X.  and  S.   Amer.,  south  to  Fueg.,  Eur.,  Afr.,  St.  Paul's  I.,  As., 
Hawaii,  X.  Zeal.— Mt. 

Pyrenulaceae. 

Arthopyroiia  Mass.  (150).  Tro[).-subtrop. 

Ci>icho}iac  M.  Arg.   Widely  distributed  in  the  trop.  zone;  also  Hawaii. — Mt. 

adnexa  M.  Arg.  var.  leptosperma  Zbr.  The  typ.  sp.  Braz. — Mf. 

pliDiorbis  M.   Arg.  Troj).-subtrop.,  also  Hawaii. — Mt. 
Poriua  M.   Arg.   (235).  All  over  the  globe,  south   to  X.   Caled.   and  N.   Zeal. 

fernandeziana  Zbr.   Belongs  in  the  vicinity  of  P.  cJdorotica  (Ach.)  M.  Arg., 
a  cosniop.  sp. — Mt. 

rufocarpella  7An. — Mt. 

depressula  Zbr.  Possibly  related  to  P.  exscrta  M.   Arg.  (Braz.).— Mt,  Mf. 
Pyrcnula  Ach.  (170).   Widely  distributed  in  the  trop.  zone. 

aspisiea  Ach.  Trop.,  also   Hawaii. — Mt. 

juanunillaua  Trev.  Trop.;   Hawaii,   S.  Chile. — Mt. 

Kioithii  h'ee.  Trop. — Mt,  Mf. 

Astrotheliaceae. 

Pyroiastnoii   ICschw.   (18).   Trop. 
chiloise  Mont.   Chile. — Mt. 

Sphaerophoraceae. 

SpJiacrophonis  Pers.  (S).   Of  wide  distribution,  centering  S.  hemisph. 

)Hela)iociirpus  (Sw.)  DC.  Almost  cosmop.;  Chile,  south  to  I'^ueg.,  P^alkl.;  N.  Zeal., 
Auckl.   and  Campb.   Is. — Mt,  Mf. 

Arthoniaceae. 

Artliouia  Ach.   (about   370).  The  majority  trop.-subtrop. 

Cytisi  Massal.   var.   meridionalis  7An.  The  typ.   s[).   luir. — Mt. 

subnebulosa  Zbr.    Related  to  A.  rphelodes  Xyl.  (X.  Caled.)  and  scitula  Krmph. 
(I^raz.)— Mt. 

berberina  Zbr.   Related  to  .-/.   rar/a  (Ach.)  Xyl.  (trop.  Amer.). — Mt. 

coftipliDiaia   l*'ee.    I'antroj).,  also   Hawaii;   Chile.  —Mt. 

Graphidaceae. 

Grapliis  Adans.  (about   280).   The  majority  tr()|).-subtrop. 
intricata   Fee.   Widely   distributed   trop.   zone. — Mt. 
Dumastii  Spreng.   Trop. — Mt. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  245 

Phaeographina  M.  Arg.  (80).  Trop.-subtrop. 

scalpturata  M.  Arg.  Trop.-subtrop.  S.  Amer.,  N.  Zeal. — Mt. 

Chiodectonaceae. 

Enterostigma  M.  Arg.  (2).   1   trop.  Amer.,  and  the  following. 
Skottsbergii  Zbr. — Mf. 

Dirinaceae. 

Dirma  Fr.  (12).  Of  wide  distribution. 
limitata  Nyl.  Chile. — Mt. 

Lecanactidaceae. 

Schismatomma  Fw.  et  Koerb.  (80).  Mainly  warmer  countries. 
accede7is  (Nyl.)  Zbr.   Chile.— Mt. 

Chfysotrichaceae. 

Byssocaiilon  Mont.  (5).  Austral.-Oceania. 

nivetirn  Mont.   Subtrop.,  north  to  Japan,  south  to  Chile  and  N.  Zeal. — Mt. 

Thelotremaceae . 

Ocellularia  Spreng.  (over   100).  Mainly  warmer  regions. 

subdenticulata  Zbr. — Mf. 
Thelotrema  (Ach.)  M.  Arg.  (over   100).  Mainly  warmer  regions. 

lepadinum    Ach.    Widely   distributed;    also    Hawaii,    Chile,    south    to    Fueg., 
N.  Zeal.,  Auckl.  Is.— Mt,  Mf. 

Diploschistaceae. 

Diploschistes  Norm.  (30).  Cold-temp.,  trop. -alp. 

actinostomus  (Pers.)  Zbr.  N.  and  S.  temp.;   Hawaii. — Mt. 

sc7'uposus  (Schreb.)  Norm.  N.  and  S.  temp.;   S.  Chile,  south  to  Fuegia. — Mt. 

Gyalectaceae. 

Dimerella  Trev.  (39).  Mostly  trop. 

lutea  (Dicks.)  Trev.  Widely  distributed,  also  S.  Chile,  Patag.;  Haw.— Mt,  Mf. 
Gyalecta  Ach.  (60-70).  Mainly  colder  climates. 

jenejisis  (Batsch)  Zbr.  N.  Amer.,  Eur.;  N.  Zeal. — Mt. 
Pachyphiale  Loennr.  (4).   2   S.W.  Eur.,    i    bor.-temp.,  and  the  following. 

cornea  (With.)  Poetsch  et  Schiederm.  Eur.;   Chile. — Mt. 

Coenogoniaceae. 

Coenogonium    Ehrenb.   (30).   Centr.  and  trop.  S.  Amer.,  Afr.,  trop.  As.,  Austral. 

velutmum  Zbr.  S.  Chile  (comm.  by  R.  Santesson). — Mt. 
Racodium  Fr.  (i). 

rupestre  Pers.  N.  and  S.  Amer.  (Staten  I.,  comm.  by  R.  Santesson);  Eur. — Mt. 


246  C.   SKOTTSBERG 

Collemaceae. 

Lemniopsis   (\'ain.)    Zbr.    (4).    i    temp.    X.   Amer.  4-  Eur.,   2   S.W.   Eur.,    and    the 
following. 

polychidioides  Zbr. — Mt. 
/V/ys///(i  Massal.  (10).   W.   Ind.,  trop.   As.,  Japan,  Austral.,  N.   Caled. 

chilinse  Hue.   Chile. — Mt. 
Lepf(\i^iii»i  (Ach.)  S.   V.  (iray  (over   100).  All  over  the  world. 

niolucciviuui  (Pers.)  \'ain.   Widely  scattered;   Hawaii,  also  Chile. — Mt,  Mf. 

tronelloides  (L.  fil.)  S.   F.   (iray.  Trop. -tern]).,  in   Chile  south   to  W.  Patag., 
S.  (icorgia. — Mt. 

cvivifsiffis    (Ach.)    Koerb.    Tenijx-subtrop.,    e.g.   Hawaii,  S.  Amer. — Mt,  Mf. 

p/iyl/ocarpu/n  (Pers.)  Mont.   Trop.-subtrop.,   S.   Amer.,  Chile,  S.  Afr.,  Philipp., 
Australia. Mt. 

Moiziesii  (Sm.)  Mont.   Mount,  of  trop.  Amer.,  Hawaii,  Chile  south  to  Fueg., 
Ealkl.,  S.  (Georgia.— Mf. 

callitluDnniou  (Tayl.)  Xyl.  Trop.  Amer. — Mt. 

Pannariaceae. 

Pcinfiiliflla  M.  Arg.  (40).   Widely  scattered,  warm  and  cold  climates. 

}iii:^roci}icta  (Mont.)  M.  Arg.  Chile,  south  to  Fueg.,  W.  Afr.  islands,  Hawaii, 
Au.stral.,  X.  Zeal.— Mt,  Mf. 

symptychia  (Tuck.)  Zbr. — ^\t. 

pycuophora  (Xyl.)  R.   Sant.  var.  subdivisa  (Zbr.)  R.  Sant.  W.  Patag.  [32^].  The 
typ.  sp.  X.  Zeal. — Mt. 
I\vi}ia]ia  Del.  (about  60).   Widely  spread  over  the  w^orld. 

fia\i^i(usis  Zbr.   P\ieg. — Mt. 

hilaris  Zbr.      Mt. 

nihiiriuosa  Del.  .Scattered;  e.g.  Hawaii,  Chile,  Falkl.,  St.  Helena,  Campb.  I. — Mt. 

ruhijriuosa  var.  vulcanica  Zbr.  l^erhaps  specifically  distinct. ^ — Mt. 
Afiiss(i/o}ij^ia  Koerb.  (2,    i    Iv   As.). 

catuosii  (Dicks.)   Koerb.   Mount,   of  N.  Amer.  and  Eur.,  P'alkl,  X.  Zeal.— Mf. 
I\s(>ro)fia  X\-l.  (al)out  60).   Mainlx'  cold  and   temp.,  centering  in  N.   Zeal.  etc. 

vulcanicum  Zbr. — Mf. 

cephalodinutn  Zbr.      Mt. 

p/io/ido/iofi  (Mont.)   M.   Arg.   Chile,  south   to  Fueg. — Mt. 

spliiuctriuum  (Mont.)  Xyl.   S.   Amer.,  also  Chile,  south  to  P'ueg.;    X.  Zeal. — 
Mt,  Mf. 

dasycladuni  Zbr.  —  Mt. 

atigustisecttim  Zbr. — Mt. 

Stictaceae. 

Lobaria  Schreb.   (70).   Mainly  warmer  countries. 

croiulata  (Del.)  Trev.   Trop.  Amer.,  south  to  Chile,  Hawaii,  Austral-Oceania. 
—  Mt. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  247 

Pseudocyphellaria  Vain,  (with  Sticta  about  200).  Humid  trop.  to  temp,  climates. 

argyracea   (Del.)    Vain.    Chile,    S.    Afr.,    Madag.,    Mascar.,    Malays.,   Hawaii, 
N.  Zeal,  Polyn.— Mt,  Mf. 

intricata  (Del.)  Vain.  S.  Amer.;  Chile  south  to  Fueg.,  Falkl.;  Ireland,  Macaron., 
Mascar.,  Cape,  Tristan  da  C,  Hawaii. — Mt. 

fragillima  (Bab.).  S.  Amer.,  Austral.,  N.  Zeal.,  Auckl.  and  Campb.  Is. — Mt. 

subvariabilis  (Nyl.)  Vain.  Philipp.,  Austral.,  N.  Zeal. — Mt. 

chloroleuca  (Hook.  f.  et  Tayl.)  Du  Rietz.   S.  Chile  to  Fueg.,  N.  Zeal.— Mf. 

cinnamomea  (Rich.)  Vain.  S.  Chile  to  Fueg.,  Austral.,  Tasm.,  Philipp. — Mt,  Mf. 

berteroana  (Mont.). — Mt. 

hirsiita  (Mont.)  Malme.  S.  Amer.;   Chile  south  to  Fueg. — Mt. 

Guillemini  (Mont.).  S.  Chile-Fueg.— Mt. 

gilva  (Ach.)  Malme.  S.  Chile  to  Fueg.,  Falkl.,  S.  Afr.,  Australia.— Mt. 

mougeotiaita  (Del.)  Vain.  Warmer  countries;   Hawaii,  also  Chile. — Mt,  Mf. 

aurata  (Ach.)  Vain.  Trop.-subtrop.;  N.  Amer.,  Chile,  W.  Eur.,  St.  Helena, 
Hawaii,  Austral. — Mt,  Mf. 

nitida  (Tayl.)  Malme.  S.  Chile  to  W.  Patag.,  Fueg.— Mf. 

endochrysea  (Del.)  Vain.  S.  Chile  to  Fueg.,  Falkl.,  S.  Georgia;  Austral.,  N.  Zeal., 
Auckl.  and  Campb.  Is.— Mt,  Mf. 

Durvillei  (Del.)  Vain.  S.  Chile  to  Fueg.,  Falkl.,  N.  Zeal.— Mt. 

Jiavicans  (Hook.  f.  et  Tayl.)  Vain.  W.  Patag.-Fueg.;  Philipp.,  Hawaii,  Australia. 
— Mt. 

Freycmetii  {De\)  Malme.  S.  Chile  to  Fueg.,  Falkl.,  S.  Georgia,  Austral.,  Tasm., 
N.  Zeal.,  Campb.  and  Antipodes  Is. — Mt,  Mf. 

Richardi  (Mont.)  Raes.  W.  Patag.-Fueg.,  N.  Zeal,  Auckl.  Is.— Mt. 
Sticta  Schreb. 

Weigelii  (Ach.)  Vain.    Widespread,  also  Hawaii,  S.   Chile  to  Fueg.;  Austral, 
N.  Zeal— Mt. 

liyieariloba  (Mont.)  Nyl   S.  Amer.,  south  to  Magell— Mt. 

latifrons  A.  Rich.   Chile,  N.  Zeal— Mt. 

laciniata  (Huds.)  Zbr.  S.  Amer. — Mt. 
Nephroma  Ach.  (27).  N.  and  S.  temp. 

plumbeu7n  Mont.   Chile. — Mt. 

cellulosum  (Sm.)  Ach.  S.  Chile  to  Fueg.,  Tasm.,  N.  Zeal. — Mt. 

antarcticum  (Wulf.)  Nyl.  S.  Chile  to  Fueg.,  Tasm. — Mt,  Mf. 

australe  A.  Rich.  S.   Chile,  N.  Zeal— Mt. 
Peltigera  Pers.  (20).   World-wide. 

rufescens  (Neck.)  Humb.  Cosmop.;    Patag.,    Fuegia,  Falkl,  S.  Georgia. — Mt. 

polydactyla   (Neck.)    Hoffm.  Cosmop.;    Chile,  south  to  Fueg.,  Falkl,  Tristan 
da  C,  Marion  I.,  Kerguel,  Auckl.  and  Campb.  Is.— Mt,  Mf. 

Lecideaceae. 

Lecidea  (Ach.)  Th.  Fr.  (1000  or  more.?).  Preferably  cold  or  temp,  climates. 
avium  Zbr.  Related  to  L.  aeruginosa  Nyl.  (Chile). — Mt. 


248 


C.   SKOTTSBERG 


inactiva  Zbr. — Mt. 

cyanosarca  Zbr. — Mt,  Mt. 

Uucoplaca  M.   Ar^^  C'hile.  — Mt. 

leucozonata  7hx. — Mt. 

euttrolema  X>1.  Temp.   Kur.,  etc.  St.   Helena,  etc. — Mt,  Mf. 

laiYpea  Ach.  Temp,  widely  distributed. — Mt. 

viridiins  Lamy.   luir. — Mt. 

mutabilis  Fee.  \.   and  S.  Amer.,  also  Chile,  W.  Eur. — Mt. 

iiterica  Tayl.  S.  Amer.,  also  Chile. — Mf. 
Catillaria  (Ach.)  Th.   Fr.  (about   150).   Widespread  N.  and  S.  hemisph. 

i)iter)nixta  Arn.   Widespread,  south  to  N.  Zeal. — Mt. 

mclasttiria  (Nyl.)  Zbr.  S.  Amer.,  Chile  south  to  F^'ueg.,  N.  Zeal.— Mt,  Mf. 

eudochronia  (Fee)  Zbr.  N.   and  S.  Amer.^Mt. 

Umochlora  (Mont.)  Zbr.   Chile.— Mt,  Mf. 

theobromina  Zbr. — Mt. 
Mi\i^ii/os/>()rii  Mey.  et  Flot.   (about   50).   Warmer  regions. 

rersicolor  (I'"ee)  Zbr.  var.  tnicrocarpa  Zbr.  The  typ.  sp.  S.  Amer.,  N.  Zeal. — Mt. 
liiuidia  (De  Not.)  Zbr.  (at  least  200).   World-wide. 

ouiolcuca  Kickx.   Almost  cosmop.,  also  S.   Amer.,  N,  Zeal. — Mt,  Mf. 

arciutifia  (Ach.)  Arn.  var.  hyposcotina  Zbr.  The  typ.  sp.  F^ur. — Mt. 

df/iips(2?is  Zbr.   Hawaii. — Mt. 

subluifola  (Xyl.)  A.  Zbr.  l^raz.— Mf. 
T(>)U)iia  Til.   Vx.  (about  80).  Mainly  temp. 

hullata  (Mey.  et  Flot.)  Zbr.  Fxuador:  Chimborazo. — Mf. 
I.of'iuiiutn   Koerb.  (about   50).   Almost  cosmop. 

U'UioxautIiu))i  (.Spreng.)  Zbr.  var.  albidius  Zbr.  The  typ.  sp.  subtrop.,  south 
to  Austral,  and  X.  Zeal.;   Hawaii. — Mt. 

s/>.,  different  from  the  former,  described  as  JMyxodictyon  lopadioides  by  ZahL- 
r.RrCKNF.K   p.  3<S3.    Chile.— Mt. 
l\hizocaypo)i  DC',  (about  90).   Cosmopol.,  cold  to  temp,  climates. 

i::c()ij;)apliicu))i  (L.)  DC.   Cold  and  temp.,  both  hemisph.;  Patag.,  F\ieg.,  Falkl., 
Kerguel.,   W.   Antarct.     Mf. 

microspermum  7An.   .Similar  to  the  following  sp.— Mf. 

dbscuration  (Ach.)  Massal.   var.  deminututn  Zbr.  The  typ.  sp.  cold  and  temp, 
climates.      Mf. 

Phyllopsoraceae. 
Phyllopsora   ^F   Arg.   (25).   Trop.- subtrop. 

part'ifolia  (I'ers.)  AF   Arg.   Widesi)read,  also  Chile,  Hawaii  and  N.  Zeal. — Mt. 

Cladoniaceae. 

Baeoinyccs  Pers.   (34).    Majority   trop.;   7   X.   Zeal. 

chiUiisis  (Mont.)  Cromb.   Chile. — Mt,   Mf. 
C/adonid  (Hill)  Vain,  (about   280).   All  over  the  world. 

pycnodada  (Pers.)  Xyl.   Holiv.,   S.   Chile  to  Fueg.,  Falkl.;  Tristan  da  C.?—Mt, 


DERIVATION  OF  THE  FLORA  AND  FAUNA  249 

Mf.  To  this  belongs  C.  alpestris  of  Zahlbruckner  zgd.  370,  a  species  not  occur- 
ring in  the  south  hemisphere  (Santesson  J2y  and  in  letter). 

bacillaris  (S.  F.  Gray)  Nyl.  Very  widespread;   Chile,  Fueg.,  Falkl.,  Campb.  I. 
— Mt. 

didyma  Vain.  Mex.-Fueg.,  Austral.,  N.  Caled.,  Hawaii. — Mt. 

coccifera   (L.)    Willd.  Very  widespread;    Chile,  Patag.,  Fueg.,  Falkl.,  Tristan 
da  C,  W.  Antarct.— Mt,  Mf. 

<^gg^^g<^f<^  (Sw.)  Ach.  N.  and  S.  Arner.,  Chile  to  Fueg.,  Falkl.;   St.  Helena, 
S.  Afr.,  Madag.,  Auckl.  and  Campb.  Is.,  Macquarie  I.;  Asia;    Hawaii. — Mt,  Mf. 

fiircata  (Huds.)  Schrad.  Cosmopol.;    Chile  to  Fueg.,    Falkl.,    S.  Georgia,  W. 
Antarct.,  Antipodes  Is.;   Hawaii. — Mt. 

gracilis   (L.)  Willd.   Cosmopol.;    Chile  to  Fueg.,    Falkl.,  S.  Georgia,    Tristan 
da  C;   Hawaii;  Kerguel.,  W.  Antarct.— Mt,  Mf. 

pyxidata  (L.)  Fr.  Cosmopol.;   Chile  to  Fueg.,  Falkl.,  Tristan  da  C,  Kerguel., 
St.  Paul's  I.;  Hawaii.— Mf. 

finibriata  (L.)  Fr.   Cosmopol.;   Chile  to  Fueg.,  Falkl.,  Marion  I.,  Kerguel.,  W. 
Antarct.,  Auckl.  and  Campb.  Is.;   Hawaii. — Mt,  Mf. 

pityrea  (Flk.)  Fr.   Cosmopol.;  Falkl,  Tristan  da  C;  Hawaii. — Mt,  Mf. 
Stereocaulon  Schreb.  (about  90).  N.  and  S.  temp.,  trop.-subtrop.  mountains. 

patagoniciiDi  M.  Lamb.  S.  Chile  and  Patag.  to  Fueg.,  Falkl. — Mt,  Mf. 

ramtilosum  (Sw.)  Raensch.  N.  and  S.  Amer.,  Chile  south  to  Magell.,  N.  Zeal. 
— Mt,  Mf. 

implexum  Th.  Fr.  S.  Amer.,  south  to  Fueg.;  N.  Zeal. — Mt,  Mf  (f.  compactius 
(Zbr.)  M.  Lamb.). 

Acarosporaceae. 

Acarospora  Massal.  (over  200).   World-wide. 

smaragdula  (Wahlenb.)  Massal.  var.  N.  temp. — Mf. 
xanthophana  (Nyl.)  Jatta.  S.  Amer.  mountains. — SC. 

Pertusariaceae. 

Coccotrenia  M.  Arg.  (i). 

granulatiim  (Hook.  f.  et  Tayl.)  R.  Sant.  n.  comb.  (C.  curbitula  M.  Arg.,  Porina 
granulata  Hook.  f.  et  Tayl).  Chile:  Valdiv.-Fueg.,  Ceyl,  Jap.,  Philipp.,  N.  Zeal. 
—  Mt. 
Pertusaria  DC.  (about  200).  World-wide. 

leioplaca  (Ach.)  Schaer.  Cosmop. — Mf. 

polycarpa  Krph.  var.   monospora  Zbr,  The  typ,  sp.  Braz. — Mf. 

hadrocarpa  Zbr.  Similar  to  A.  cerebrinula  Zbr.  (Falkl). — Mt,  Mf. 

Skottsbergii  Zbr.  Related  sp.  in  Chile.— Mt,  SC. 
Mela7iaria  Erichs.  (15).  N.  and  S.  Amer.,  Eur.,  S.  Afr.,  As.,  N.  Zeal. 

melanospora  (Nyl)  Erichs.  S.  Amer.,  also  Chile. — Mt,  SC,  Mf. 

Lecanoraceae. 

Lecanora  Ach.  (over  200).  World-wide. 

masafuerensis  Zbr.   Close  to  L.  subimmersa  (Fee)  Vain.  (Braz.). — Mf. 


250  C.  SKOTTSBERG 

coarctata  (Sin.)  Ach.  Widespread  temp.;   Chile. — Mf. 

atra  (Huds.)  Acli.  Cosmop.;   Chile,  Falkl.;   Hawaii. — Mt,  Mf. 

Ingae  Zbr.— Mt,  Mf. 

albilli)U}  M.  Arg.   var.  validior  Zbr.  The  typ.  sp.  Fueg. — Mf. 

dis/^ersa  (Pers.)  Flk.  Widespread  N.  hemisph. — Mt,  Mf. 

polytropa    (Fhrh.)    Ach.    Cosmop.;    Chile,    Falkl.,  W.  Antarct.;  Hawaii. — Mf. 

thrysolcuca  (Sm.)  Ach.  W.  Arct.,  Fur.  mountains,  Antarct.,  etc. — Mt. 

saxkola  (Poll.)  Ach.  X.  Amer.,  p:ur.;   .^ Chile.— Mt. 
Placopsis  Nyl.  (31).   Widely  distrib.,  but  mainly  austral,  18  southern  S.  Amer. 

chilcua  M.  Lamb.   Chile.— Mf. 

fuscidula  M.   Lamb.  S.   Chile  to  P^ueg.,  Tristan  da  C. — Mt,  Mf. 

pareUina   (Xyl.)    M.    Lamb.    Boliv.    And.,   Chile  to  P'ueg.,  Java,  S,  Austral., 
X.  Zeal,   Plawaii.— Mt. 

gelida  (L.)  Ach.  Circumpol.  Arct.  and  temp,  oceanic;   Chile,  Tristan  da  C, 
Kerguel.,  Java,    X.    Zeal.,    Hawaii. — Mf  (var.    subreagens  M.   Lamb,  but  identity 
doubtful). 
Caudclariella  M.  Arg.  (27).  X.  and  S.  Amer.,  Eur. 

intellina  I\L  Arg.  Widespread;   Chile,  Falkl.,  Hawaii. — Mt,  Mf. 
Myxodiityon  Massal.  (3,    i   Australia). 

chrysostictum  (Tayl.)  Mass.  Chile,  X.  Zeal— Mt.  M.  lopadioides  Zbr.  2g6.  383 
is  a  species  of  Lopadiiim  (Santesson  in  letter). 

Parmeliaceae. 

J\iruie/ia  (Ach.)  De  Xot.  (about  400).   World-wide. 

laevigata   (Sm.)    Ach.    Widespread  temp,  and  trop.;  also  Hawaii  and  Chile. 
— Mt. 

lacrigatula  Xyl.   Ikaz.- — Mt. 

rei'oluta  P1k.   Widespread  temp. -trop.  Also  Hawaii. — Mf. 

cctrata    Ach.    W^y  widespread  temp,   and  trop.,  also  Hawaii. — Mt,   SC,  Mf. 

saxatilis  (L.)  Ach.   Widespread,  in  Chile  south  to  F'ueg.,   Falkl;   W.  Antarct. 
— Mt. 

couspeysa  (P'.hrh.)  Ach.  Cosmop.,  also  Hawaii,  P^alkl— Mt,  SC,  Mf. 

ahstrusa  \'ain.   Hraz. — Mf. 

polata  Ach.   Widespread;  also  Hawaii  and  S.   Chile.  —  Mt. 

)tilglicrr€)isis  Xyl   Trop.-subtrop. — Mf. 

pilosella  Hue.   X.   Amer.   to   Mex.,   PLur. — Mt,  Mf. 

piloselloides  Zbr. — Mt. 

cetrayioidt's  Del   Very  widespread.— Mf. 

uncrosdcta  M   Arg.  'Prop.   Amer. — Mt. 

caperata  (L.)  Ach.  Temp,  zones;   Chile,   Hawaii. — Mt,  Mf. 

sored ica  Xyl.   Calif.,   Mex. — Mf. 
Meiiegazzia  Mass.  (30).    P'ew  north  hemisj)h.,  majority  S.  Amer.  (11)  and  Austral- 
'Pasm.-X.  Zeal.  (14). 

sauguinascens  (Raes.)  R.   Sant.  [^^28.  i  i,  Parmelia  pertusa  (Schrad.)  Schaer.  in 
2g6.  389),  Valdiv.   to  Fueg.— Mt,  Mf. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  25 1 

Ratnalina  Ach.  (about   lOo).  World-wide. 

linearis  (Sw.)  Ach.  Warmer  regions;   Chile,  Fueg.,  Falkl.,  N.  Zeal. — Mt. 

us7tea  (L.)  Howe  Jr.  Trop.-temp.,  N.  Amer.,  Chile. — Mf. 
Usnea  Wigg.  (about   lOo).  Cosmop. 

dasypogoides  Nyl.  Rodriguez  I. — Mt,  Mf. 

florida  (L.)  Hoffm.  Very  widespread,  also  Chile. — Mf. 

subtorulosa  (Zbr.)  Motyka.  Piaster  I. — Mf. 

angulata  Ach.  N.  Amer.,  south  to  Mex.;   S.  Amer.  to  Chile. — Mf. 

Caloplacaceae. 

Blastenia  (Massal.)  Th.  Fr.  (about  60).  World-wide. 

fernandeziana  Zbr. — Mt,  SC. 

ferruginea  (Huds.)  Massal. — Mt.  Determination  probably  incorrect  ace.  to  Dr. 
Magnusson. 
Bombyliospora  De  Not.  (25).  Mostly  trop.-subtrop. 

dolichospora  (Nyl.)  Zbr.  Chile. — Mt. 
■Caloplaca  Th.  Fr.  (over   100).  World-wide. 

clandestina  Zbr. — Mf. 

Selkirkii  Zbr.— Mt,  Mf. 

rubina  Zbr.  Easter  I.— Mt,  SC,  Mf. 

isidioclada  Zbr. — Mf. 

subcerina  (Nyl.)  Zbr.  var.  aurantiaca  Zbr.  Trop. — Mf. 

elegans  (Link)  Th.  Fr.  var.  australis  Zbr.  Chile,  W.  Patag.  The  typ.  sp.  cold 
to  temp.  N.  and  S.  hemisph. — Mt,  SC,  Mf. 

orthoclada  Zbr.  In  the  vicinity  of  C  Feliponei  Zbr.  (Urug.). — Mf. 

Teloschistaceae. 

Teloschistes  Norm.  (12).  World-wide. 

flavicans   (Sw.)    M.    Arg.    Widespread    trop.-subtrop.,    in  S.  Amer.  south  to 
Fueg.,  Falkl.— Mt. 

Buelliaceae. 
Buellia  De  Not.  (about  200).  World-wide. 

concinna  (Stzbgr.)  Th.  Fr.  var.  oceanica  Zbr.  The  typ.  sp.  Eur. — SC. 

siphoniatula   Zbr.    Similar   to  B.  posthabita  (Nyl.)  Zbr.  (Colomb.)  and  falk- 
.landica  Darb.  (P^alkl.). — Mt. 

stellulata  (Tayl.)  Mudd.   Cosmop.   Chile;  Kerguel— Mt,  SC,  Mf. 

halophila  M.  Arg.  Australia.— Mt. 
•  halophiloides  Zbr.  Easter  \. — Mt. 

fei'nandeziana  Zbr.   Easter  I. — Mt. 

masafuerana  Zbr. — Mf. 

barrilensis  Zbr. — Mf. 

Physciaceae. 
-Pyxine  (Fr.)  Nyl.  (16).  Warmer  regions. 

curvatula  Zbr. — Mt. 


C.  SKOTTSBERG 


Pliyscia  (Schreb.)  X'aiii.  (100-150).  Cosmop.,  most  numerous  temp,  regions. 

picta  (Sw.)  Xyl.   Widespread  trop.-subtrop.  Also  Hawaii. — Mt. 
Anaptychia  Koerb.   (30).   Widespread,  mainly  warmer  regions. 

Inpolcuca  (Muhlb.)  Massal.  Widespread,  also  Hawaii. — Mt. 

pccti}iata  (Zbr.)  R.   Sant.   Patag.  (Nahuelhuapi),  Fueg. — Mt. 

Hymenolichenes. 

Cora  Fr.  (8).  Trop.-subtrop. 

pai'onia  (Sw.)  Fr.  IMex. -Chile,  south  to  Fueg.;   St.   Helena. — Mt,  Mf. 

The  list  includes  194  species;  103  (53  %)  of  these  are  restricted  to  Masatierra, 
including  3  also  found  on  Santa  Clara,  2  (i  %)  have  only  been  encountered  on 
this  islet,  39  (20  %)  only  on  Masafuera;  50  (25.7  %)  are  listed  for  both  islands, 
6  of  them  also  found  on  Santa  Clara.  The  number  of  endemic  species  is  36  (18.5  %), 
a  number  likely  to  be  reduced  when  the  lichen  flora  of  South  America  becomes 
better  known;  some  species  described  by  Zaiilbruckner  as  endemic  in  Juan 
Fernandez  have  later  been  found  on  the  mainland  or  on  Hawaii.  New  discoveries 
will,  on  the  other  hand,  be  made  in  the  islands. 

Masatierra  has  153  species,  of  which  26  (17%)  are  endemic  in  Juan  Fer- 
nandez; the  corresponding  figures  for  Santa  Clara  are  11  and  3  (27.3  %)  and  for 
Masafuera  89  and  15  (16.8%).  Masatierra  is  richer  in  lichen  species  than  Masa- 
fuera; this  may  have  something  to  do  with  the  greater  variety  of  substratum  of- 
fered by  the  numerous  species  of  trees  and  shrubs  inhabiting  only  Masatierra. 
Of  100  corticolous  species  62  are  restricted  to  Masatierra,  16  to  Masafuera  and 
22  occur  on  both  islands;  thus  84  species  have  been  collected  on  the  former  and 
only  38  on   the  latter. 

The  greater  wealth  of  Masatierra  is  also  shown  in  the  number  of  endemic  spe- 
cies found  only  on  Masatierra,  21  (of  36)  or  58.3%;  3  were  found  also  on  Santa 
C'lara.  The  figure  for  Masafuera  is  10  =  27.7%;  of  these  5  belong  to  the  highland 
above  1 000  m.  Only  5  species  (14%)  have  been  found  on  both  islands.  I  suppose 
that  the  superiority  of  Masatierra  depends  on  the  greater  variation  of  habitat. 
r\iture  research  will,  I  suppose,  yield  numerous  additional  species,  but  I  do  not 
expect  that  the  relation  between   the  islands  will  be  much  altered. 

It  is  difficult  to  arrive  at  a  geographical  classification  of  the  lichens  because 
in  too  many  cases  only  very  general  information  is  given:  "in  the  tropics",  "in 
warmer  regions",  "in  temperate  regions"  and  so  forth,  and  "cosmopolitan"  is  used 
too  generously.  Statements  suggesting  the  most  surprising  disjunctions  are  not 
uncommon;  some  are  probably  due  to  wrongly  named  specimens.  Unfortunately 
the  liclien  flora  of  Chile  is  not  very  well  known.  I  have  tried  to  find  out  if  a 
s[)ecies  called  cosmopolitan  has  been  recorded  for  Chile.  Many  world-wide  lichens 
have  been  found  in  Juan  l-'ernandcz;  if  also  found  in  Chile  they  were  referred 
to  the  Chilean  element. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  253 

I.  Andine-Chilean  element. — ^106  (62.4%). 

a.  Endemic  species  supposed  to  be  most  nearly  related  to  Chilean  species  (7): 
Psoroma  vulcanicum,  cephalodinum,  dasycladum  and  angustisectum,  Pseudocyphel- 
laria  berteroana,  Lecidea  avium,  Pertusaria  Skottsbergii. 

b.  Also  known  from  Chile  (88):  Verrucaria  microspora,  Normandina  pulchella, 
Pyrenula  mammillana,  Pyrenastrum  chilense,  Sphaerophorus  melanocarpus,  Arthonia 
complanata,  Dirina  limitata,  Schismatomma  accedens,  Byssocaulon  niveum,  Thelo- 
trema  lepadinum,  Diploschistes  scruposus,  Dimerella  lutea,  Pachyphiale  cornea, 
Coenogonium  velutinum,  Racodium  rupestre,  Physma  chilense,  Leptogium  molucca- 
num,  tremelloides,  cyanescens,  phyllocarpum  and  Menziesii,  Parmeliella  nigrocincta 
and  pycnophora  var.,  Pannaria  rubiginosa,  Psoroma  pholidotum  and  sphinctrinum, 
Lobaria  crenulata,  Pseudocyphellaria  argyracea,  intricata,  cinnamomea,  hirsuta, 
Guilleminii,  gilva,  mougeotiana  and  aurata,  Sticta  Weigelii,  lineariloba  and  lati- 
frons,  Nephroma  plumbeum  and  australe,  Peltigera  rufescens  and  polydactyla, 
Lecidea  leucoplaca,  mutabilis  and  icterica,  Catillaria  melastegia  and  leucochlora, 
Rhizocarpon  geographicum,  Phyllopsora  parvifolia,  Baeomyces  chilensis,  Cladonia 
pycnoclada,  bacillaris,  didyma,  coccifera,  aggregata,  furcata,  gracilis,  pyxidata, 
fimbriata  and  pityrea,  Stereocaulon  ramulosum  and  implexum,  Coccotrema  granu- 
latum,  Melanaria  melanospora,  Lecanora  coarctata,  atra  and  polytropa,  Placop- 
sis  chilena,  fuscidula,  parellina  and  gelida,  Candelariella  vitellina,  Myxodictyon 
chrysostictum,  Parmelia  saxatilis,  laevigata,  conspersa,  perlata  and  caperata,  Mene- 
gazzia  sanguinascens,  Ramalina  linearis  and  usnea,  Usnea  florida  and  angulata, 
Bombyliospora  dolichospora,  Caloplaca  elegans,  Buellia  stellulata,  Theloschistes 
flavicans,  Cora  pavonia. 

c.  Widespread  to  cosmopolitan  species  expected  to  occur  on  the  mainland 
of  Chile  (11):  Diploschistes  actinostomus,  Pseudocyphellaria  fragillima,  Sticta  laci- 
niata,  Catillaria  intermixta  and  endochroma,  Bacidia  endoleuca,  Pertusaria  leio- 
placa,  Parmelia  revoluta,  cetrata  and  cetrarioides,  Physcia  picta. 

II.   Subantarctic-Magellaniati  element. — 14  (8.2  %). 

a.  Endemic:  Pertusaria  hadrocarpa. 

b.  Also  in  W.  Patagonia,  Fuegia  etc.  (13):  Pannaria  fuegiensis,  Pseudocyphel- 
laria chloroleuca,  nitida,  endochrysea,  Durvillei,  flavicans,  Freycinetii  and  Richardi, 
Nephroma  cellulosum  and  antarcticum,  Stereocaulon  patagonicum,  Lecanora  albel- 
lina  (endem.  var.),  Anaptychia  pectinata. 

III.   Neotropical  element. — 26  (153%). 
Endemic,  or  found  or  expected  in  South  America,  but  not  recorded  for  Chile. 

a.  Endemic  (5):  Arthonia  subnebulosa  and  berberina,  Enterostigma  Skotts- 
bergii, Lecanora  masafuerensis,  Caloplaca  orthoclada. 

b.  Not  endemic,  some  perhaps  to  be  expected  in  Chile  (14):  Arthopyrenia 
Cinchonae,  adnexa  and  planorbis,  Phaeographina  scalpturata,  Leptogium  calli- 
thamnion,  Megalospora  versicolor  (endem.  var.),  Bacidia  subluteola,  Toninia  bullata, 
Acarospora  xanthophana,  Pertusaria  polycarpa  (endem.  var.),  Parmelia  laevigatula, 
abstrusa  and  microsticta,  Anaptychia  hypoleuca. 


254 


C.  SKOTTSBERG 


C-.  Widespread  tropical-subtropical  species,  probably  also  occurring  in  South 
America  (7):  Pyrenula  aspistea  and  Kunthii,  Graphis  intricata  and  Dumastii,  Lo- 
padiuni  leucoxanthum  (endem.  var.),  Parmelia  nilgherrensis,  Caloplaca  subcerina 
(endem.   var.). 

IV.  Pacific  element.  —  7  (4.1  %). 

Pscudoc\phellaria  subvariabilis,  l^acidia  delapsans,  Usnea  subtorulosa,  Calo- 
placa rubina,  Huellia  halophila,  fernandeziana  and  halophiloides. 

V.  Boreal  element. — 16  (9.4%). 

if.   luulemic:   Leniniopsis  polychidioides. 

/;.  Not  endemic  (15):  Arthonia  cytisi  (endem.  var.),  Gyalecta  jenensis,  Massa- 
longia  carnosa,  Lecidea  enteroleuca,  latypea  and  viridans,  l^acidia  arceutina 
(endem.  var.),  Riiizocarjxni  obscuratum  (endem.  var.),  Acarospora  smaragdula, 
Lecanora  dis{)ersa,  chrysoleuca  and  saxicola.  Parmelia  pilosella  and  soredica,  Buel- 
lia  concinna  (endem.  var.). 

VI.   Only  reported  from  Juan  Fernandez  and  Rodriguez  I. 

Usnea  dasy{)ogoides. 

The  following  22  endemic  species  had  to  be  left  out,  their  taxonomic  rela- 
tions being  unknown:  Microglaena  fernandeziana,  Porina  depressula,  fernandeziana 
and  rufocarpella,  Ocellularia  subdenticulata,  Parmeliella  symptychia,  Pannaria  hilaris, 
Lecidea  inactiva,  cyanosarca  and  leucozonata,  Catillaria  theobromina,  Rhizocarpon 
microspermum,  Lecanora  Ingae,  Parmelia  piloselloides,  Blastenia  fernandeziana, 
Caloi)laca  clandestina,  Selkirkii  and  isidioclada,  Buellia  siphoniatula,  masafuerana 
and  barrilensis,  I^yxine  curvatula — further,  Lopadium  sp,  and  the  dubious  Blastenia 
sp.   have  been   excluded.  The  {percentages  were  calculated  w'ith    170  as  a  total. 

VI.   Fungi. 

Our  knowledge  of  the  fungus  flora  is  very  limited,  and  time  did  not  permit 
us  to  pay  due  attention  to  this  group.  Of  Basidiouiycctcs  about  40  species  were 
identified  by  RoMi:[,i,  {2of),  including  those  enumerated  by  JoilOW  (several  spe- 
cies doubtful),  luidemic  species  few.  Keisslkr's  list  of  Ascomycetes  (759,  i6o\ 
with  additions  by  AkWiDssoN  (.?//),  short  as  it  is — only  31  species — gives  some 
idea  of  the  relation  between  the  geographical  elements.  Of  the  20  named  spe- 
cies, 9  were  known  before  from  S.  America  (mostly  Chile)  and  I  from  N.  Amer- 
ica; 5  are  wides[oread  and  5  endeiuic,  one  of  these  belonging  to  an  endemic 
genus.  Liimniiiia  scoriadca  (Berk.)  Keissl.  is  also  known  from  Chile,  Java  and  New 
Zealand.  Three  of  the  endemic  genera  of  Compositae  have  their  special  rusts, 
the  endemic  I'lupluasia  is  attacked  by  the  same  Urcdo  that  is  found  on  two  species 
of  sect.  Irifidac  in  Chile,  Azara  fcy)ia)ideziana  by  the  rust  known  from  A.  integrifolia 
in  Chile,  and  Rubus  geoides  is  accompanied  by  the  same  parasite  as  in  P^uegia  (j_5(?). 

Of  (lasteroDiycetes  only  2  named  s[)ecies  were  reported;  o^  i\\QSQ  lieodictyon 
jrraci/e  l^erk.  is  of  geographical  interest:  S.  America,  S.  Africa,  Australia  and  New 
Zealand,  the  second  species  was  known  before  from  N.  America  and  Samoa  [102). 


DERIVATION  OF  THE  FLORA  AND  FAUNA  255 

The  Myxophyta,   i8  species,  are  more  or.  less  cosmopolitan  [loi). 

I  have  tried  above  to  indicate  where  the  native  plants  have  their  nearest  sta- 
tions outside  the  islands  or,  if  endemic,  where  their  closest  relatives  occur.  Sta- 
tistics like  this  serve  to  assign  to  a  local  flora  its  position  within  a  certain  floristic 
region  and,  when  dealing  with  an  oceanic  island,  to  trace  the  sources  from  where 
its  living  world  is  likely  to  have  been  derived.  The  position  generally  assigned 
to  Juan  Fernandez  is  that  of  an  outpost  from  South  America.  In  Engler's  Syl- 
labus the  islands  form  the  "Gebiet  von  Juan  Fernandez  und  Masafuera"  under 
"Zentral-  und  siidamerikanisches  Florenreich";  it  is  characterized  thus:  "Gattungen 
vorzugsweise  verwandt  mit  denen  der  chilenischen  tjbergangsprovinz",  i.e.  Cen- 
tral Chile  and  the  Valdivian  forest  region,  Engler's  transitional  belt  between  the 
Andean  and  Magellanian  provinces.  Good  [log)  distinguished  a  "Region  of  Juan 
Fernandez"  under  his  "Neotropical  Kingdom",  giving  it  the  same  rank  as  the 
"Amazone  Region",  the  "Andean  Region",  etc.  We  have  seen  that  there  is  an 
unmistakable  floristic  agreement  between  Juan  Fernandez  and  South  Chile,  but 
also  that  it  is  far  from  complete,  and  both  Engler  and  GoOD  were  well  aware 
of  the  presence  of  elements  that  had  little  to  do  with  the  flora  of  the  mainland. 
They  were  barely  recognized  by  JoilOW  (see  above  p.  215)  who,  with  his  faith 
in  unlimited  transoceanic  dispersal,  paid  little  attention  to  them;  they  were  too 
few  to  disturb  the  Chilean  picture.  Not  until  the  flora  had  become  better  known 
did  its  strange  features  stand  out  in  a  clearer  light.  It  is  surprising  that  GuiLLAU- 
MIN  who  knew  and  quoted  the  synopsis  published  in  vol,  II  of  this  work  (2.?^) 
failed  to  recognize  them.  In  his  paper  on  the  floristic  divisions  of  the  Pacific  he 
states  that  Juan  Fernandez  lies  outside  Oceania  and  he  includes  it  in  his  discus- 
sion only  for  the  sake  of  comparison.  The  flora  is  characterized  as  follows  [118. 
931):  "Sur  142  Phanerogames  indigenes,  la  moitie  sont  endemiques  mais  appa- 
rentees  aux  especes  chiliennes;  les  autres  sont  cosmopolites  ou  Sudamericaines, 
surtout  chiliennes."  It  is  not  easy  to  understand  how  he  arrived  at  this  conclusion. 
On  the  other  hand,  some  authors  were  led  astray  by  the  difference  between  the 
islands  and  the  continent.  Ermei.,  who  got  his  impression  from  a  short  visit  to 
Masatierra,  wrote  (^7.48): 

.  .  .  muss  die  Zusammengehorigkeit  dieser  Insel  zum  amerikanischen  Festlande,  zu 
welcher  Ansicht  deren  geringe  geographische  Entfernung  von  selbst  hinleiten  konnte,  aufs 
nachdriicklichste  in  Abrede  gestellt  werden,  weil  die  beiderseitige  Flora  and  Fauna  zu 
grosse  Verschiedenheiten  aufweist,  wo  wir  an  geeigneter  Stella  die  notigen  Beweise  bei- 
bringen  werden.  Die  Gestaltung  der  Flora  ordnet  die  Inselgruppe  vielmehr  dem  aiistralischen 
Weltteile  zu,  von  dem  der  grosste  Teil  heruntergegangen  ist. 

It  is  hardly  necessary  to  mention  that  his  proofs  were  based  on  his  ignorance 
of  the  composition  of  the  island  flora  and  very  likely  also  of  the  floras  of  Australia 
and  Chile. 


256  C.  SKOTTSBERG 


Chapter  II, 


Sources  of  the  island  flora  as  judged  by  the  total  distribution  of  the 

geographical  elements  distinguished,  with  special  reference  to  the 

composition  of  the  Chilean  flora. 

I  have  attempted  above  to  describe  tlie  distribution  of  the  non-endemic  spe- 
cies, to  state  where  the  endemics  have  their  relatives,  if  any^  and  to  distinguish 
a  number  of  geograpliical  elements.  Now  1  shall  proceed  a  little  further  and  look 
at  the  matter  from  a  wider  horizon.  A  species  was  called  Chilean  because  it  is 
found  also  in  Chile  or  has  its  relatives  in  the  south  Andean  flora,  or  it  was  referred 
to  a  Magellanian  group  because  it  occurs  only  in  the  farthest  south  of  the  conti- 
nent, and  so  forth;  but  in  order  to  know  something  of  the  genesis  and  history 
of  each  group  we  cannot  stop  here.  We  shall  find  that  our  "Chilean  element", 
Chilean  from  our  insular  viewpoint,  consists  of  several  types,  each  with  its  own 
distribution  pattern.  To  speak  with  WULFF  (2^1.  203),  until  now  we  occupied  our- 
selves with  the  geograpJiic  elements,  now  we  shall  try  to  trace  the  ^r?/r//r  ones, 
"species  grouped  according  to  their  region  of  origin,  thus  reflecting  the  genesis 
of  a  given  flora".  He  very  properly  adds:  "To  determine  the  region  of  origin  of 
a  species" — and  indeed  also  of  a  genus  or  family — "is  often  a  very  difficult  mat- 
ter, requiring  a  monographic  study.  .  .  ."  With  regard  to  Juan  Fernandez,  the  ge- 
netic elements  are  congruent  to  Wulff's  "migration  elements". 

I.  Angiospermae. 

Of  the  two  species  of  Stipa,  necsiana  is  distributed  from  Mexico  through  the 
tro{)ical  Andes  to  Central  Chile  and  east  to  Brazil,  Argentina  and  Uruguay,  lae- 
nissinia  (Xassella)  a  typical  Andean  species;  the  former  is  neotropic,  the  latter 
Chilean,  extending  north  into  Peru  and  east  into  Argentina.  Almost  the  same 
area  is  occupied  by  riptocliaeiiuin  bicolor,  a  genus  limited  to  extratropical  South 
America. 

rodopJioiHs  is  a  unicjue  anomaly  without  known  neotropical  affinities,  as  it 
were,  a  far-travelled  member  of  an  Arcto-tertiary  flora,  if  its  affinity  with  Brachy- 
clythruDi  hits  the  mark;   in  Pil(;er's  opinion  it  sides  with  MegalacJine. 

The  sj)ccies  of  L'haetotropis  were  referred  to  the  neotropical  group — "tropical" 
not  to  be  taken  in  a  purely  climatic  sense,  because  a  species  included  under  this 
heading  may  just  as  well  be  subtro[)ical  and  even   extend  into  a  temperate  zone. 

Agrostis  inasafucyana  and  the  bicentric  A.  viagcllmiica  were  linked  together, 
but  f)nly  provisionally,  because  in  a  large  and  world-wide  genus  like  Agrostis  the 
relationships  cannot  be  safely  judged  without  a  thorough  taxonomic-genetic  study 
of  the  whole  genus.  Assuming  that  PllXiFR  was  right,  Antarctica  becomes  in- 
volved, and  the  two  species — and  probably  others  as  well — should  be  classified  as 
"Old  Antarctic  ',  or,  as  I  now  prefer  to  term  them,  Antarcto-tertiary  (correspond- 


DERIVATION  OF  THE  FLORA  AND  FAUNA  257 

ing  to  Arcto-tertiary).  An  Antarcto-tertiary  taxon  is  not  necessarily  of  Antarctic 
ancestry;  strictly  spoken  it  certainly  should  be,  but  there  are  numerous  cases 
where  the  point  of  origin  was  either  the  South  American  or  the  Australasian- 
New  Zealand  centre,  and  Antarctica  only  served  as  a  trans-continental  route  of 
migration. 

Trisetum  chromostachymn  is  Chilean,  but  the  genus  is  both  N.  and  S.  temper- 
ate, perhaps  originally  Arcto-tertiary?  Danthonia  collhta  and  Koelerm  micrathera 
also  are  Chilean,  but  the  distribution  pattern  of  the  genera  indicates  that  Antarc- 
tica eventually  was  involved  in  their  history. 

Megalackne,  temporarily  referred  to  Bromus  but  once  more  stated  to  be  an 
independent  genus,  is  a  relict  type  with  unknown  history,  eventually  a  remnant 
of  a  pre- Andean  flora  (comp.  above  p.   217). 

The  genus  Chusqiiea  belongs  to  the  neotropical  element  in  the  flora  of  Chile, 
and  the  same  is  true  of  the  species  of  Cyperus.  Scirpiis  nodosus  and  cernuus  are 
circumpolar  seaside  plants  and  most  likely  thalassochorous;  if  Antarctica  had  a 
share  in  their  earlier  history  is  impossible  to  say,  but  it  is  not  improbable.  Eleo- 
charis  is  another  large  world-wide  genus,  perhaps  too  wide-spread  to  allow  us  to 
locate  its  place  of  origin. 

Oreobolus.  The  Antarcto-tertiary  character  of  Oreobolus  can  hardly  be  dis- 
puted; it  is  often  referred  to  as  a  classical  example  of  an  Antarctic  genus.  Of  the 
6  species  recognized  by  KUKENTIIAL,  i  (with  2  varieties)  is  found  in  S.  and  E. 
Australia,  Tasmania  and  New  Zealand,  2  in  N.E.  New  Guinea,  one  of  them 
extending  to  Borneo,  i  in  N.  Sumatra  and  Malacca,  i  in  Hawaii,  and  O.  obtusangu 
lus  in  Chile  from  the  Cordilleras  of  Valdivia  to  Fuegia,  the  Falkland  Is.  and  Juan 
Fernandez.  [A  seventh  species,  0.  pfeifferianus  Barros,  was  identified  by  KUKEN- 
THAL  [16^.  vill)  with  pjunilio  var.  pectinaius\ 

Cladiiim  (16^.  Xll)  is  wide-ranging  but  it  is  not  cosmopolitan  in  spite  of  the 
large  areas  occupied  by  C.  7nariscus  L.  and  its  varieties,  among  v^\\\q\\  j amaiceiise 
is  circumpolar  and  distributed  also  south  of  the  equator.  The  main  distribution 
of  subg.  Machaerina  (inch  Vincentia),  where  C.  scirpoideum  of  Juan  Fernandez 
belongs,  is  palaeotropical  with  11  species  (Madagascar,  Mascarene  Is.,  Australia, 
Lord  Howe  I.,  New  Guinea,  Indonesia,  Oceania);  5  are  neotropical  (W.  Indies, 
Brazil).  Its  austral  character  is  clear  enough.  The  closest  relative  of  scirpoideum 
is  not  an  American  species  but  C.  angustifolium  (Gaud.)  Benth.  et  Hook.  fil.  (New 
Guinea,  Tahiti,  Hawaii).  Subg.  Baumea,  with  29  scattered  from  Australia,  where 
there  are  18  species,  over  the  Indomalayan  region  west  to  Ceylon,  north  to  Hong- 
kong and  Japan,  east  to  Melanesia  and  Hawaii,  has  15  in  Australia-Tasmania- 
New  Zealand  and  i  in  the  region  of  Madagascar  and  Mascarene  Is.  Together  the 
two  subgenera  cover  the  3  sectors,  the  African,  the  Australian-Malaysian  and  the 
American.  In  the  centre  of  this  vast  area  lies  Antarctica  or,  as  it  were,  Gond- 
wana  Land.  The  history  of  Cladium  may  well  lead  back  to  the  Mesozoic,  and 
it  seems  natural  to  refer  the  genus  to  the  Antarcto-tertiary  element.  The  same 
applies  to  Uncinia,  one  of  the  generally  recognized  Antarctic  genera,  represented 
in  Juan  Fernandez  by  5  species,  2  of  them  endemic.  Within  the  South  American 
sector  are   13  species  distributed  along  the  Andes  with  a  concentration  toward  the 

17  "557857  The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.  Vol.  I 


238  C.   SKOTTSBKRG 

south,  one  species  ^oin<^  nortli  as  tar  as  Mexico  and  the  West  Indies,  i  extending 
east  from  I^^ie^ia  to  the  Falkhmd  Is.  and  Tristan  da  Cunha.  On  the  opposite  side 
of  Antarctica  are  18  species.  14  of  tliese  indit^enous  in  New  Zealand,  where  8  are 
endemic,  the  remainder  scattered  alon^  tiie  route  Macquarie  I.-Auckland  and  Camp- 
bell Is.-Tasmania-Australia-New  Zealand-Lord  Howe  and  Norfolk  Is.;  one  New 
Zealand  species  reappears  in  Hawaii,  another  on  Kerguelen  and  New  Amster- 
dam I.,  and  Marion  1.  has  an  endemic  species.  Finally,  2  species  are  found  in  New 
(iuinea,  one  of  them  also  reported  from  Borneo,  and  the  Philippines  have  I  en- 
demic species,  l^oth  sections  of  sub^.  liuriicijiia  are  represented  in  America,  only 
Stenandrae  in  the  opj)osite  sector.  The  monotypical  subg.  Pseudocarcx  is  Magel- 
lanian. 

It  is  not  easy  to  find  one's  way  through  the  labyrinth  of  the  enormous  and 
still  (growing  ^cnus  Carcx,  world-wide  but  unbalanced  as  the  tropics  are  poor  in 
species  in  comparison  with  the  temperate  and  cold  zones.  Our  two  island  species, 
the  endemic  C.  hfrttrojiiivia  and  the  south  Andean  Baiiksii,  belong  to  different 
sections  and  different  geographic  groups,  the  former  to  sect.  Echinochlaenae:  of 
20  species  16  are  endemic  in  New  Zealand  (one  with  a  variety  on  Norfolk  I.), 
I  in  Australia,  1  in  Tasmania;  the  little  known  Chilean  C.  laniprocarpa  Phil,  and 
L  .  bfrti-yojiiaiia  are  far-flung  outposts,  but  it  lies  near  at  hand  to  assume  that  they 
or  their  ancestors  migratetl  across  Antarctica.  C.  Bavksii  belongs  to  PVigidae- 
I'"uli<4inosae,  a  boreal  group  centering  in  luu'asia  with  one  species  in  Pacific  North 
America,  but   I   cannot  tell  if  luviksii  comes  near  this  species. 

The  s\stematic  position  of  Juiuiia  austraiis  was  briefly  discussed  in  22g. 
109  and  above  p.  202,  but  whether  we  bring  it  to  Morenieae  or  Iriarteae  or  let  it 
form  a  separate  subtribe  it  remains  a  member  of  the  neotropical  element.  HUTCIIIN- 
S(>N  followed  Hi:n'1  HAM  and  HooKKR  in  placing  it  next  to  Ceroxylon,  but  this 
genus  is  |)ol\gam()-monoecious  and  the  stigma  becomes  basal  in  fruit.  To  me 
Ckni/A  1  s  opinion  lacks  foundation;  he  solves  what  he  calls  "a  hopeless  conflict 
among  taxonomists"  (7/.  85)  by  deriving  Juajiia  from  "a  massive  center  of  origin 
of  angiospermy  at   the  Mascarenes  "   (p.    103). 

Ochai:;ai'ia  and  I It'sf^crooyeiiria  belong  to  an  Andean-neotropical  assemblage 
of  genera,   with   close  relati\es  in   Chile. 

To  judge  from  Hr(  IIK\  \i  s  monogra[)h  of  the  Juncaceae  in  Pflanzenreich 
f.uzula  uiasafucnvia  nuist  be  referred  to  a  grouj)  of  Andean  species,  L.  racentosa 
Desv.  (.Mex.  S.  Chile),  cxcelsa  Huch.  (Boliv.),  ///^vyw]'/;// Buch.  (Argent.),  /.^'^^^//V 
Buch.  (C^hile),  and  cliiloisis  .\ees  et  Mey.  (Chile,  south  to  l^'uegia),  but  in  the  same 
grou})  we  find  A.  spicata  (L.)  DC.  (Arct.  circumj).  and  Alpine)  and  ahyssinica  Pari. 
(ICthiop.,  Brit.  \\.  Afr.),  and  the  |)()ssibilit}'  that  the  Andine  species  are  of  boreal 
origin  should  be  considered.  On  the  other  hand  much  speaks  in  favour  of  a  south- 
ern origin  of  Juncaceae:  the  subantarctic  genera  Marsippospennuui  dind  RostkovJay 
the  isolated  rnoinuiu  in  South  Africa,  endemic  Andine  genera  like  Oxychloc  and 
Patosia,  well  marked  endennc  species  of  I.uzula  in  New  Zealand  and  the  Magel- 
lanian  region,  and  the  subantarctic-bicentric  Jinicns  scluuchzcrioides  group.  It  is 
true  that  4  of  the  5  J// ;/r//.s- species  reported  from  Juan  Fernandez — some  of  them 
perhaps  not  native— inhabit  the  Andes,  3  going  east  to  Brazil,  Argentina  and  Uru- 


DERIVATION  OF  THE  FLORA  AND  FAUNA  25  9 

guay,  the  fifth,  J.  planifolius,  shared  by  Chile  and  AustraHa-Tasmania-New  Zea- 
land, but  it  is  also  true  that  even  if  we  look  for  the  origin  of  the  family  in 
Antarctica,  a  possibility  pointed  out  by  Weimarck  {281),  we  must  count  with 
secondary  centres  of  evolution  in  the  boreal  zone  where  the  overwhelming  major- 
ity of  sections  and  species  are  found.  Some  50  species  are  in  the  South  hemi- 
sphere with  main  centres  in  South   Africa  (20)  and  Australia  (17). 

Libertia  with  3  species  in  Chile  and  2  in  New  Guinea-Australia-New  Zealand 
tells  a  story  of  an  Antarctic  past,  and  the  group  Sisyrinchineae  has  a  stronghold 
in  the  South,  where  the  genera  concentrate;  Sisyrinchium  itself  centres  in  South 
America,  where  4  small  genera  are  endemic  [Symphyosteinon,  Chamaelum,  Sole- 
fiomelus  and  Tapeinia);  South  Africa  is  another  stronghold  [Aristea,  Wilsenia^ 
Bobartia,  Klattia  and  Cleanthe),  and  '^[Orthosanthus,  Diplarrhena  and  Paiersojiia) 
are  found  in  the  East  Australian  -  Indomalayan  region,  but  this  does  not  entitle  us 
to  derive  Iridaceae  from  the  far  south. 

Of  the  4  island  species  of  Peperomia  only  P.  fernandezia?ia  (Chile)  is  neo- 
tropical, while  the  other  species  appear  to  be  more  nearly  allied  to  palaeotropical 
ones  (Java,  Australia,  Oceania);  P.  berteroana  occupies  a  unique  geographical  posi- 
tion (p.  203).  The  genus  is  of  tropical  origin  and  centres  in  America,  but  if  we 
try  to  understand  the  history  of  the  endemic  species  of  Juan  Fernandez,  the  possi- 
bility of  Antarctica  as  a  migration  route  should  be  considered. 

Whereas  Urtica  masafuerajia^  one  of  the  few  indigenous  annuals,  is  related 
to  a  species  from  Ecuador,  U.  feriiandeziana  with  the  habit  of  a  miniature  tree 
{22g.  862)  appears  to  lack  near  relatives.  The  family  is,  I  suppose,  of  tropical 
origin,  but  the  actual  centre  of  Urtica  is  in  the  north  temperate  and  subtropical 
zones;  there  are  14  species  in  Eurasia,  including  the  Mediterranean  region,  and 
7  in  North  and  Central  America  with  a  secondary  centre  in  the  Andes  (Colomb.- 
Fueg.  6);  3  species  are  tropical  (Braz.-Urug.,  Ethiopia,  Java).  There  are  a  few 
species  in  the  south  hemisphere,  2  S.  Afr.,  i  Australia,  2  N.  Zeal,  and  i  Auckl.  Is. 
U.  fernandeziana  seems  to  represent  an  ancient  type. 

Boehmeria  excelsa  was  described  by  BuRGER  (^7)  as  an  elegant  shrub  which  came 
from  S.  America;  it  is  a  clumsy  tree  and  not  related  to  neotropical  species.  It 
comes  nearest  to  B.  dealbata  (Kermadec  Is.)  and  points  west,  not  east. 

Parietaria  hiimiftisa  (Chile);  see  above  p.  204.  Belongs  to  a  neotropical  group 
which  has  not  been  cleared  up  [241)  but  is  also  closely  related  to  the  Australian- 
Polynesian  debilis  of  FORSTER.  Weddell's  debilis  is  a  mixture  of  varieties  scat- 
tered over  the  globe  and  most  likely  consists  of  several  good  species. 

Phrygilanthus  Berteroi.  The  family  Loranthaceae  is  \xo\i\Q.di\,h\xX.Phrygilanthus 
is  of  Antarctic  origin.  Of  the  7  sections,  4  (about  20  species)  are  American  and 
range  from  Lower  California  to  S.  Chile,  i  with  2  species  belongs  to  Australia 
and  New  Zealand,  i  (i  species)  to  New  Zealand,  and  i  has  one  species  in  Aus- 
tralia, another  in  New  Guinea  and  a  third  in  the  Philippines — a  typically  austral- 
bicentric  genus. 

The  genus  Santalum  ranges  from  Australia  and  Melanesia  to  Micronesia  and 
Malaysia  and  east  to  Polynesia  and  Hawaii  with  a  distant  outlying  station  in 
Juan    Fernandez;    see   Tuyama's   map    (^70),  Of  the  4  sections  distinguished  by 


26o  C.  SKOTTSBERG 

TrVAMA  (who  divided  my  section  Eusanialum  in  2  and  referred  the  Hawaiian 
tLusantala  to  a  sejxirate  section  SolenaJithus),  Fjisantalinu  s.  str.  is  the  largest  with  9 
species,  SoloiaJithus  and  sect,  llawaiiensia  (together  8)  confined  to  Hawaii  (incl.  Lay- 
san),  and  sect.  Poly)icsica  (2,  with  varieties)  to  Polynesia;  in  this  section  the  extinct 
.V.  fi-riiajideziajnofi  occupies  a  rather  independent  position.  "Aber  woher  kam  der 
frenulartige  l'>eniit  Santaluni.-"  lU'RC.KR  [4.1.  22)  exclaims,  "zweifelsohne  fern  aus 
ostindischen  Meercn,  wo  die  W'iege  seines  altberiihmten  Geschlechtes  steht" — 
nothing  could  give  a  wronger  idea  of  the  history  of  the  sandalwoods,  because 
everything  points  toward  Antarctica  as  their  cradle.  There  are  related  endemic 
genera  in  Australia,  Mida  in  New  Zealand,  Jixocarpus  ranges  from  Australia  across 
Polynesia  to  Hawaii,  and  several  endemic  genera  are  at  home  in  temperate  S. 
America  [Arjojia,  lodiiia,  Ouidia,  QuincJianialiuui,  and  Xa7iodea  in  the  extreme 
south  and  in  the  Falkland  Is.);  one  is  tempted  to  regard  the  family  as  of  Antarctic 
ancestry,  but  with  Thesiiini  in  mind  it  might  be  safer  to  speak  of  a  special 
Antarcto-tertiary  centre. 

The  3  endemic  species  of  CheJiopodiuin  were  commented  upon  above  (p.  204). 
In  general  appearance  they  are  very  like  the  Hawaiian  oahuense  but  the  special- 
ist oj)ines  that  they  are  not  nearly  related  to  this  nor  to  other  species,  I  can  find 
no  better  place  for  them  than   in  a  neotropical  group. 

SalicoDiia  fruticosa,  taken  in  a  wide  sense,  is  a  thalassochorous  plant  scattered 
along  tropical  and  subtropical  coasts;  the  same  is  the  case  with  Tetragoiiia  ex- 
pausa  in  the  S.  hemisphere,  but  while  Saliconiia  is  a  world-wide  genus,  Tetra- 
i:;o)iia  has  a  stronghold  in   South  Africa  and  several  endemic  species  in  Chile. 

Sptrj^idar/a  is  a  wide-ranging,  but  mainly  boreal  genus  with  a  vigorous  branch 
in  Andean  America  but  absent  from  Australia,  New  Zealand  and  Oceania.  The 
2  island  species,  of  which  .V.  co7ifertiJiora  is  also  found  on  San  Ambrosio,  are 
closely  related  to  Chilean  species.  Paronychia  has  about  the  same  distribution 
j)attcrn  as  Spcyo^ularia,  but  is  poorly  represented   in   S.  America. 

RamDuulus  capraniui.  A  very  large  essentially  boreal-temperate  genus  with 
Avell  stocked  branches  in  S.  America  and  on  New  Zealand.  The  Masafueran  en- 
demic stands  a|)art  from  its  American  congeners  and  approaches  certain  New  Zea- 
land sj)ecies,  j)crhaps  also  the  Hawaiian  ones.  An  Antarctic  migration  route  seems 
probable. 

Fuyheris  coryinbosa  and  niasafuerana  belong  to  a  small  section  confined  to 
the  tropical  Andes  and  not  extending  to  Chile,  where  we  have  many  other  spe- 
cies. The  present  area  of  the  Berberidaceae  testifies  to  its  Arcto-tertiary  charac- 
ter; it  centres  in  \\.  and  S.  Asia  and  ranges  far  south  only  in  America,  where  3 
species  reach   I'\iegia. 

Of  the  six  genera  of  W'interaceae  (<\?),  Bubbia  has  2  species  in  Australia, 
I  on  Lord  Howe  I.,  <S  in  New  Caledonia  and  19  in  New  Guinea,  Belliohun  4 
in  New  Caledonia  and  4  in  the  Solomon  Is.,  Pseudowintcra  2  in  New  Zealand, 
Jixospoimiin  2  and  Zy^^^ogyjiu))!  6  in  New  Caledonia,  and  Drimys  6  in  Australia 
{i  also  in  Tasmania),  29  in  New  (Guinea  and  i  on  Borneo,  Celebes  and  in  the 
Philippines,  all  these  belonging  to  sect.  Tasma7iia\  the  other  section,  Eudrimys, 
is  American  with  4  species,  J),  confcrtifolia  endemic  in  Juan  Fernandez.  This  sec- 


DERIVATION  OF  THE  FLORA  AND  FAUNA  261 

tion  is  a  distant  branch  of  an  Australasian  family  and  ranges  from  the  uttermost 
south  along  the  mountains  to  Mexico  and  to  Roraima  in  Brazil.  That  Antarctica 
once  played  a  role  in  the  history  of  Drimys  is  proved  by  the  Tertiary  fossils 
discovered  in  West  Antarctica. 

Lactoris  is  generally  looked  upon  as  a  primitive  member  of  the  Ranales  and 
claimed  to  belong  to  the  most  primitive  element  in  the  island  flora.  It  has  no 
relatives  in  America.  It  is  no  typical  member  of  the  Magnoliales,  an  Arcto-tertiary 
order,  which  it  needs  a  Croizat  to  derive  from  the  Antarctic.  Geographically, 
Lactoris  is  a  parallel  to  Degeneria  of  Fiji,  but  the  affinities  of  the  latter  are  not 
questionable  and  they  have  little  in  common.  It  lies  near  at  hand  to  think  of  the 
small  family  Lardizabalaceae,  endemic  in  Chile.  Hutchinson  regards  Lactorida- 
ceae  as  "closely  related  to  the  Wiriteraceae,  of  which  it  is  probably  a  reduced 
derivative"  [140.  II.  85) — but  is  the  perfect  trimery  a  result  of  reduction.?  And  are  not 
the  never  quite  closed  carpels  an  indication  of  primitiveness?  I  daresay  most  sys- 
tematists  agree  that  the  Polycarpicae,  whether  regarded  as  07te  order  or  split  up,  are 
among  the  oldest  living  angiosperms.  The  distribution  of  Winteraceae  and  the 
occurrence  in  the  South  Hemisphere  of  small  isolated  families  as  Degeneriaceae, 
Lactoridaceae  and  Lardizabalaceae  suggests  that  the  Antarctic  continent  was  one 
of  the  centres  of  evolution. 

Cardamifte  is  a  world-wide,  mainly  temperate  and  essentially  boreal  genus, 
extending  into  the  tropics  and  south  to  Fuegia  and  New  Zealand.  The  3  island 
species,  one  of  them  endemic,   were  commented  on  above  (p.   205). 

Among  the  numerous  Chilean  species  of  Escallojiia,  E.  Callcotiiae  stands  a 
little  apart  from  the  rest  {is8)\  the  genus  is  spread  along  the  Andes  and  extends 
to  Brazil  and  Uruguay.  The  subfamily  Escallonioideae  is  austral-circumpolar:  Tri- 
beles  (i  S.  Chile-Fueg.),  Valdivia  (i  S.  Chile),  Forgesia  (i)  and  Berenice  (i), 
Reunion,  Anopterus  (2,  Tasm.,  E.  Austral.),  Cuttsia  (i  E.  Austral),  Argophylliim 
(10  E.  Austral.,  N.  Caled.),  Colmeiroa  (i  Lord  Howe  I.),  Carpodeius  (i  N.  Zeal., 
N.  Guin.),  Quijitinia  (15  Austral. -N.  Guin.-N.  Caled.,  Philipp.),  Potiingeria  (i  N.E. 
Ind.) — a  distribution  suggesting  an  Antarctic  origin. 

Rubus  geoides  has  a  single  near  relative,  R.  radicans,  in  S.  Chile.  They  differ 
very  much  from  the  numerous  north  temperate  and  tropical  montane  species  and 
form  their  own  section  or  subgenus,  and  their  resemblance  to  the  Tasmanian 
R.  gunnianus  Hook.  Icon.  Plant.  Ill  is  no  proof  of  affinity.  They  seem  to  represent 
an  isolated  offshoot  from  the  north  which  has  become  cut  off  from  its  source  of 
origin  and  found  its  way  into  the  subantarctic  zone. 

Acae7ta.  Nobody  if  not  Croizat  would  argue  that  Rosaceae  are  a  southern 
family,  but  this  cannot  prevent  us  from  assuming  that  the  Sanguisorba  assemblage 
of  genera  has  gone  through  part  of  its  evolution  at  least  in  the  far  south.  BiTTER 
{31)  distinguished  10  sections:  I,  13  species,  S.  Amer.;  II,  i.  The  Cape;  III,  2, 
I  S.  Amer.,  i  Hawaii;  IV,  i,  S.  Amer.;  V,  8,  6  S.  Amer.,  i  J.  Fern.,  i  Tasm.; 
VI,  I,  S.  Amer.;  VII,  28,  26  S.  Amer.,  i  Cahf.,  i  Tasm.;  VIII,  64,  58  S.  Amer., 
I  Tristan  da  C,  i  N.  Amsterd.  I.,  3  N.  Zeal.,  i  N.  Zeal.-Tasm.-Austral.-N.  Guin.; 
IX,  I,  N.  Zeal.;  and  X,  2,  N.  Zeal.  Two  Magellanian  species  occur  on  S.  Georgia 
and   I   on    Kerguelen.  The  circumpolar  distribution  shows  no  gaps.  About  70  % 


262  C.  SKOTTSBERG 

of  the  species  inhabit  extratropical  S.  America,  but  many  of  them  are  so  closely 
related  that  they  are  little  more  than  microspecies  and  in  not  few  cases  based 
on  one  or  two  specimens  from  a  single  locality,  and  the  number  of  separable 
taxa  will  perhaps  be  reduced  when  more  material  becomes  available.  I^e  this  as 
it  may,  7  sections  are  represented  in  America,  5  in  the  Australian-X.  Zealand 
area,  3  are  common  to  both,  one  of  them  ranging  to  the  African  sector,  where 
another  section  is  endemic.  As  regards  A.  i)iasafuera7ia  see  p.  206  above.  I  sup- 
j)ose  we  can  draw  no  other  conclusion  from  this  distribution  than  that  Acaeiia  is 
an  Antarcto-tertiary  genus,  2  sections  having  developed  numerous  species  in  the 
Andes  and  Patagonia. 

Mari:;yricaypus,  a  small  Andean  genus,  is  so  closely  allied  to  Acaciia  that  they 
have  j)roduced  a  bigeneric  hybrid  in  Juan  Fernandez.  Two  more  genera  are  found 
in  the  Andes,  TetvaglocJii}!  and  Polylepis.  In  S.  Africa  we  have  the  large  genus 
Cliffortia.  The  remaining  genera  Sangiijsorba,  Poteriuiu  and  Beiicomia  (Macarone- 
sia)  belong  to  the  X.   hemis{)here. 

Sophoni  sect.  Tetrapterae  is  austral-circumpolar:  New  Zealand  (3  species), 
Chatham  I.  (i).  Lord  Howe  I.  (i).  Austral  Is.  (i),  Rapa  (i),  Marquesas  (i),  Hawaii 
(i),  I'^aster  I.  (i),  Juan  Fernandez  (2),  Chile  (2;  S.  Diacrocarpa,  however,  rather  unlike 
all  the  others),  Diego  Alvarez  (i),  and  Reunion  (i).  With  the  exception  o'i macrocarpa 
and  the  Hawaiian  cJirysopJiylla  the  remaining  species  used  to  be  united  under 
tetraptera  Ait.,  otherwise  endemic  in  New  Zealand.  They  are  very  closely  related, 
but  distinct;  it  is  of  minor  importance  if  we  call  them  species  or  geographical 
subs{)ecies.  Unless  we  believe  that  kS.  tetraptera  was  carried  by  water  from  island 
to  island  and  was  transformed  into  a  new  species  wherever  it  landed,  we  must 
look  upon  Antarctica  as  a  one-time  centre  of  a  polymorphous  population,  which 
radiated  in  various  directions;  we  shall  not  discuss  here  how  this  may  have  hap- 
pened. We  have  not  to  do  with  litoral  but  with  inland  plants;  the  pods  are 
ada|:)tc(l  to  float,  assisted  by  the  four  narrow  wings,  Joiiow  says,  but  some  of  the 
forms  have  no  wings  at  all,  and  even  if  they  have,  the  pods  open  on  the  tree  and 
discharge  their  seeds. 

I'agara  inayu  and  exterjia  form  their  own  section.  When  HC'RGER  said  [4.1.  19) 
that  I'agara  had  migrated  to  Juan  Fernandez  from  the  primeval  forests  of  Peru 
and  Colombia  he  oxerlooked  that  the  affinity  is  with  palaeotropical  rather  than 
with  neotropical  si)ecies;  there  are  numerous  species  scattered  from  Australia  and 
Xew  Caledonia  to  Polynesia  and  Hawaii,  where  many  are  endemic.  Rutaceae  were 
})erhaps  represented  in  the  Antarctic  in  Tertiary  times,  and  we  have  too  look  for 
a   route  across  to  the  American  sector. 

The  family  liuphorbiaeeae  is  pantroj)ical,  let  alone  that  Euphorbia  has  at- 
tained a  world-wide  distribution  and  flourishes  also  in  temjierate  climates.  Dysopsis 
is  Andean,  Seidelia  (2)  and  Leidesia  (i)  South  African,  the  fourth  genus  of  the 
JSPercurialis  grouj),  Mercurialis  (<S),  ranges  from  Xorth  l^urope  to  the  Mediterranean 
and  is  found  in  \\.  Asia.  The  southern  genera  seem  to  be  more  closely  con- 
nected mutually  than  with  Mereiirialis.  The  disjunctions  are  interesting  and  difficult 
to  explain,  unless  we  can  find  good  reason  to  look  for  a  common  source  in  the 
Antarctic. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  263 

The  systematical  position  of  Callitriche  has  been  discussed  many  times,  but 
we  know  nothing  of  its  history  or  where  it  started.  The  genus  is  world-wide,  but 
many  species  are  not  wide-spread  and  some  are  quite  local,  among  them  C.  Lech- 
leri,  which  may  have  been  carried  to  Juan  Fernandez  from  the  mainland  by 
accident. 

The  tribe  Colletieae  of  the  otherwise  wide-ranging  Rhamnaceae  is  called 
austral-antarctic  by  Sussengut  (Xaturl.  Pflanzenfam.  2nd  ed.);  except  Adolphia 
(Mexico-U.S.A.)  the  genera  are  distributed  over  Andean  and  extratropical  S.  Amer- 
ica, centering  in  Chile,  and  Colletia  spartioides  finds  its  place  with  the  Andean 
element.  It  should  be  mentioned  that  Discaria  (ii)  extends  south  to  Fuegia  and 
reappears  in  Australia  (i)  and  New  Zealand  (i),  suggesting  transantarctic  migra- 
tion from  America. 

The  family  Flacourtiaceae  is  tropical;  Azara  is  neotropical  with  about  19 
species  in  Chile,  i  in  Brazil  and  i  in  Argentina.  The  tribe  to  which  it  belongs 
is  well  developed  in  the  south  hemisphere:  tropical  America  and  Africa,  Mada- 
gascar and  neighbouring  islands,  Asia,  New  Guinea  and  [Xylosma]  Oceania  to 
Hawaii,  but  Antarctica  may  not  at  all  be  involved  in  its  history. 

Myrtaceae.  A  very  large  world-wide  and  tropical-subtropical  family.  Of  the 
subfamily  Myrtoideae,  some  2400  species,  75  %  are  American,  the  remainder  scat- 
tered over  Asia,  Africa,  Australia  and  Oceania.  The  Leptospermoideae,  some  850, 
are  restricted  to  Australasia  with  the  single  exception  of  Tepualia,  monotypical 
and  endemic  in  the  Chilean  rain  forests,  south  to  West  Patagonia,  a  most  inter- 
esting case  of  disjunction.  Berry  [26]  regards  the  family  to  be  of  American  ori- 
gin and  to  have  attained  its  present  distribution  before  the  close  of  the  Creta- 
ceous. Basing  his  conclusions  on  fossil  evidence  he  thinks  that  during  the  cooling 
down  of  the  climate  during  late  Tertiary,  the  ancestral  stock  of  Myrtoideae  with- 
drew from  North  America  to  the  neotropical  zone;  the  Australian  Leptospermo- 
ideae represent  the  remnants  of  the  Cretaceous  radiation  during  which  numerous 
new  types  became  evolved.  Some  of  these  eventually  invaded  Antarctica  and 
Tepualia  survives  on  Chilean  soil. 

The  Juan  Fernandez  Myrtoideae  are  closely  linked  to  Andean  types.  For  Ugni 
Selkirkii  ^v\d  Myrteola  Tiuniinularja  see  p.  206.  NotJiomyrcia,  now  restricted  toMasa- 
tierra,  may  or  may  not  have  inhabited  a  larger  area.  Myrceugenia  has  about  20 
species  in  Chile. 

Gunneraceae  (often  placed  as  a  subfamily  under  Halorrhagidaceae)  is  a  classi- 
cal example  of  a  tricentric  Antarcto-tertiary  type.  Its  long  and  complicated  history 
is  reflected  in  its  taxonomic  differentiation;  6'/<';/;7^r<^  is  composed  of  6  subgenera. 
Pangue  is  the  largest  with  10  species  ranging  from  Costa  Rica  to  S.  Chile,  i  in 
Brazil,  3  in  Juan  Fernandez  and  2  in  Hawaii.  Perperisuvi  is  monotypical  with 
separate  varieties  in  S.  Africa,  British  E.  Africa  and  Madagascar,  Ostenia  an  aber- 
rant monotype  endemic  in  Uruguay.  Misandra  includes  3  species,  one  extending 
from  Colombia  to  Fuegia  and  Falkland,  one  restricted  to  the  S.  Chilean  Andes 
and  one  to  subantarctic  America;  Milligajiia  has  8  species  in  New  Zealand  and 
I  in  Tasmania.  The  monotypical  Pseudogmmera  inhabits  New  Guinea,  Java,  Su- 
matra and  the  Philippines.  No  subgenus  is  found  in  more  than  one  sector.  It  is 


264 


C.  SKOTTSBERG 


surprisinj^  that  the  geographically  isolated  Hawaiian  species  are  closely  related  to 
the  species  of  Juan  I^>rnandez.  We  know  other  examples  of  this  connection,  but 
to  construct  a  route  between  Hawaii  and  Juan  Fernandez  meets  with  serious  obsta- 
cles, and  Pa)igue  may  have  reached  Hawaii  along  a  quite  different  route.  The 
evolution   and  differentiation  of  the  subgenera  very  likely  took  place  in  Antarctica. 

Scilixni-KR  (Pflanzenreich)  expressed  his  opinion  of  the  history  of  Halor- 
rhagidaceac  thus:  "Aus  der  geographischen  Verbreitungder  ursprlinglichsten  Halor- 
rhagaceen,  namlich  der  Gattung  Halorrhagis,  ist  mit  Sicherheit  zu  folgern,  dass  die 
Familie  antarktischen  Ursprungs  ist."  It  is  not  easy  to  see  how  he  arrived  at  this 
conclusion,  though  I  think  it  is  correct,  because  at  that  time  the  endemic  species 
of  Juan  Fernandez  were  unknown;  they  passed  as  //.  erecta,  a  New  Zealand  en- 
demic, and  this,  incorrectly  attributed  also  to  Juan  Fernandez  and  Chile,  had  been 
carried  there  on  purpose:  "Der  Standort  auf  Juan  Fernandez  und  in  Chile  ist  kein 
urspriinglicher,  sondern  durch  die  auch  in  Neuseeland  erfolgende  Verwendung  der 
Pflanze  als  Futterptianze  erklart."  But  there  was  no  Ilaloi'rJiagis  on  the  mainland, 
and  the  species  indigenous  in  Juan  Fernandez  were  not  used  as  forage,  let  it  be 
that  the  introduced  animals  eat  them.  Halorrhagis,  eminently  Australian  (59  out 
of  80  species)  and  with  7  species  in  New  Zealand,  extends  north  to  Indomalaya, 
IVIicronesia,  S.F.  China  and  Japan  and  east  to  Rapa  (not  known  to  Schindler) 
and  Juan  h^rnandez — see  Ti'YAMa's  map  of  distribution  [2yi].  The  genus  "well 
illustrates  the  not  infrequent  extension  of  an  Australasian  group  far  north  of  the 
equator,  and  the  much  rarer  condition  of  occurrence  in  Juan  Fernandez  but  not 
in  continental  America"  {log.  108).  HalorrJiagis  shows  the  same  distribution  pat- 
tern as  ScDitaluDi. 

Coitclla  is  essentially  African,  see  above  p.  207;  the  distribution  is  tricentric 
with  some  remote  stations.  The  widespread  C.  asiatica  is  scattered  over  a  broad 
belt  but  not  reported  from  America.  Very  likely  the  Hydrocotyloideae,  a  sepa- 
rate family  according  to  some  authors,  are  of  Antarctic  origin,  but  C.  triflora 
may  well   have  reached  Juan   Fernandez  with  the  traffic  from  Chile. 

The  peculiar  endeinic  species  of  Eryjigiuui  differ  so  much  in  habit  from  all 
other  species  of  this  large  and  widespread  genus  that  they  have  been  referred 
to  a  se{)arate  genus,  a  rank  to  which  they  are  not  entitled,  liryvgiuni  concen- 
trates in  tiie  Mediterranean  region  and  in  tropical  South  America,  where  also  the 
island  species  Ijelong,  in  sj)ite  of  their  arborescent  habit;  to  quote  TURMKI.  [26g. 
130):  "Se  rattachant  a  la  Region  chilienne,  on  cite  les  especes  de  Juan  Fernandez 
li.  hupleuvoidcs,  sai'copJiylluiii  et  hiaccessuni  .  .  . ,  plantes,  du  moins  pour  les  deux 
premieres,  arbustives  s'o[)posant  radicalement  aux  autres  especes  des  territoires 
voisins"— but  viacccssuni  also  is  a  dwarf  tree,  more  so  I  would  say  than  sarco- 
pliyllufti.  I  doubt  that  they  descend  from  herbaceous  continental  forms;  they  be- 
long to  an   ancient  type  and   find   their  proper  place  with  the  neotropical  element. 

The  tribe  Apioideae-Amnfineae  is  very  widely  spread,  with  a  concentration 
in  the  \orth  hemisphere,  Apiuni  distributed  also  in  the  south  temperate  zone,  and 
A.  gyai'eoh)is  L.  is  frecjuently  regarded  as  bipolar  species.  I  don't  believe  that 
any  of  the  southern  forms  should  be  included;  they  are,  however,  in  need  of 
revision.  A.  fcDicDidcziajiuin  is  a  well-marked  species,  related,  but  not  very  closely, 


DERIVATION  OF  THE  FLORA  AND  FAUNA  265 

to  a  series  of  forms,  probably  good  species,  reported  from  subantarctic  America, 
Falkland,  Tristan  da  Cunha,  Australia,  etc.  and  possibly  descending  from  an 
old  Antarctic  stock. 

Pernettya:  13  species,  8  Mexico  and  Centr.  America  to  Chile,  south  to  Fuegia 
and  Falkland,  i  Galapagos  Is.,  2  in  Tasmania  and  2  in  New  Zealand;  P.  rigida 
is  linked  to  Andean  species,  but  very  distinct  [2^2).  The  genus  is  more  diversi- 
fied in  the  American  sector,  and  this  seems  to  be  where  it  originated,  having 
reached  New  Zealand  across  the  Antarctic,  if  not  with  Gaultheria,  well  developed 
in  New  Zealand,  descending  from  a  common  Antarcto-tertiary  stock. 

Empetrum  is  a  bipolar  genus,  the  family  most  likely  of  boreal  origin.  Con- 
cerning E.  rubrum  see  2^g.jd>\. 

Of  the  7  Dichondra  species  5  are  neotropical,  i  endemic  in  New  Zealand 
and  D.  repefis  (incl.  sericea)  spread  round  the  world.  It  is  common  on  the  coast 
of  Chile  and  possibly  adventitious  in  Juan  Fernandez.  The  occurrence  of  an  en- 
demic species  in  New  Zealand  suggests  that  Antarctica  witnessed  part  of  its 
history. 

Calystegia.  About  25  species  have  been  described,  scattered  over  the  globe, 
C.  sepiiwi  sensu  lat.  reported  from  America,  Eurasia,  N.  Africa,  Australia,  New 
Zealand,  Easter  I.,  etc.  and  evidently  very  easily  naturalized.  The  plant  found 
on  Masafuera  and  also  on  the  mainland  was  described  as  C.  Hantelman7ii  Phil, 
and  later  identified  with  tugurioriwi  from  New  Zealand;  see  2^g.  783.  If  this  is 
correct,  C.  tugiiriorurn  offers  one  of  the  very  few  cases  of  a  species  restricted  to 
Chile  and  New  Zealand,  but  even  if  they  are  kept  apart,  they  present  a  remark- 
able case  of  disjunction. 

Selkirkia  Berteroi,  the  only  representative  of  Boraginaceae,  so  richly  devel- 
oped on  the  mainland,  was  regarded  as  an  isolated,  Old  Pacific  type  [22"/.  31,.?^^. 
593)  until  Johnston  {148)  showed  that  it  comes  close  to  Hackelia  and  differs  from 
this  principally  by  its  arboreous  habit.  With  Urtica  fernandezimia,  the  species  of 
EryngiwH  etc.  I  refer  it  to  the  neotropical  element. 

Rhaphithannius  vc7iustiis  is  of  neotropical  ancestry  (see  above  p.  208);  the 
second  species  is  common  in  Centr.  and  S.  Chile. 

Briquet  placed  Cuminia  in  the  Stachyoideae-Menthinae  next  to  Oreosphacus 
Phil.,  a  shrub  of  the  high  Cordillera  in  the  boundary  region  between  Chile  and 
Argentina,  but  this  genus  has  a  schizocarp  of  four  nuts.  It  has  also  been  com- 
pared with  Bystropogon  L'Herit.  (Canary  Is.,  Colomb.-Peru,  different  sections).  Ep- 
LING  (using  the  fancy  name  JoJioivid)  referred  Cuminia  to  Prasioideae,  an  ancient 
group  showing  great  disjunctions:  Prasitmi  Mediterranean,  Stenogyne,  Phyllostegia 
and  Haplosiachys  in  Hawaii,  Bosirychmtthus  and  Gomphostemma  in  Asia;  Cuminia 
differs  in  the  shape  of  the  corolla  (see  above  p.  208),  but  even  so  it  seems  to  repre- 
sent a  palaeotropic  element  in  the  island  flora. 

Solanum  fernandezianum  is  a  distinct  species  of  indubitable  neotropical  and 
Andean  ancestry.   With  regard  to  ^.  robinsonia?ium ,  see  above  p.   200. 

Nicotiajta  cordifolia  has,  according  to  GOODSPEED  [112.  347),  its  closest  resem- 
blance, in  flower  structure  as  well  as  in  general  habit,  to  N.  Raimondii.  Crosses 


266  C.  SKOTTSBERG 

with  this  species  and  with  solanifolia  gave  evidence  of  fundamental  affinities  between 
the  island  endemic  and  species  of  the  mainland  to  the  north  (I.e.). 

Mi})iulus  is  essentiall)'  a  western  N.  American  genus  with  few  species  else- 
where; the  somewhat  polymorphous  M.  glabratus  ranges  from  N.  America  to 
Bolivia,   Chile  and  Argentina. 

The  different  opinions  on  the  systematic  \)os\t\on  o{  Euphrasia  forifios is siifi a 
have  already  been  referred  to  above.  Within  the  area  of  the  geographically  isolated 
Chilean-Magellanian  'Irifidac  occurs  the  semicalcarate  E.  pcrpitsillaV\\\\.  (S.  Chile). 
Both  would  indicate  a  road  from  the  Australian-New  Zealand  area  across  the  Ant- 
arctic to  America,  just  as  the  tropical  mountains  of  Malaysia  served  as  a  road 
between  Asia  and  Australia,  as  Dl'  RiETZ  thinks  (77.  536):  "The  Euphrasia  popu- 
lation of  Juan  r>rnandez  may  therefor  very  well  have  formed  the  northern  end 
of  a  population  so  far  south  that  the  lack  of  close  relationships  between  E.for- 
niosissiDia  and  the  species  of  Middle  Chile  is  fully  explained,  and  the  semical- 
carate anthers  of  the  more  southern  E.  perpusilla  may  be  the  last  South  American 
remnant  of  this  old  connection."  Another  question,  not  yet  answered,  is  this:  are 
the  very  well-marked  Trifidae,  reaching  from  subantarctic  America  to  the  Andes 
of  S.  Chile,  likewise  descendants  from  a  remote  southern  population  or  did  they 
originate  in   Chile? 

Plantago  fernaiuiezia  is  another  arborescent  member  of  a  herbaceous  world- 
wide genus  and  often  cited  as  an  example  of  a  mysterious  connection  between 
Hawaii  and  Juan  Fernandez,  but  if  we  look  at  the  total  distribution  of  sect.  Palaeo- 
psyliiuui  (see  above  p.  209)  its  character  of  an  austral  group  is  revealed,  even  if 
it  extends  north  to  X.  America  and  S.  Europe.  In  the  south  it  is  circumpolar, 
and  the  route  from  New  Zealand  (Auckland  Is.)  via  Rapa  to  Hawaii  can  be  traced 
— a  radiation  from  Antarctica  seems  not  unthinkable. 

Plantago  truucaia,  represented  in  Juan  Fernandez  by  an  endemic  (.')  variety, 
belongs  to  the  large  section  Xoiwrbis  and  needs  no  further  comments. 

The  neotropical  Ilcdyotis  ihesiifolia  is  a  recent  addition  to  the  flora  and  may 
have  been  accidentally  introduced. 

Xcrtcra  is  an  austral  circumpolar  genus  allied  to  Coprosvia  and  best  devel- 
oped in  Xcw  Zealand  where  5  species  occur,  4  of  them  endemic;  the  fifth  is 
X.  grauadfusis  (depressa),  claimed  to  be  very  widely  spread.  One  species  is  en- 
demic in  Tristan  da  Cunha.  Recent  observations  tend  to  show  that  granaderisis 
includes  taxonomically  distinct  forms;  the  Malaysian  plant  is  not  identical  with 
the  iXndean,  and  other  forms  will  perhaps  become  distinguished  after  a  critical 
revision.  He  this  as  it  may,  the  genus  is  Antarcto-tertiary  and,  if  we  link  Tristan 
da   Cunha  with   Africa,  tiicentric. 

CoprosDia  is  a  parallel  to  I lalurrJiagis  but  differs  in  being  present  in  Hawaii; 
another  difference  is  that  the  species  of  Juan  Fernandez  do  not  have  their  closest 
relatives  in  New  Zealand  or  Australia — lloolccri  forms  its  own  section,  pyrifolia 
is  of  Polynesian  affinity.  As  the  genus  is  absent  from  America  as  well  as  from 
Africa  it  should  lie  near  at  hand  to  refer  it  to  a  West  Pacific  element,  a  position 
favoured  by  the  relationships  o{  pyrifolia,  and  to  the  believers  in  transoceanic 
migration  combined  with  evolution  of  local  endemics  wherever  (T^/r^j-;;/^  happened 


DERIVATION  OF  THE  FLORA  AND  FAUNA  267 

to  land  this  is  the  only  course  to  take,  even  if  the  sudden  appearance  of  an  iso- 
lated type  like  Hookeri  becomes  somewhat  embarrassing.  With  an  extension  north 
and  east  of  an  Antarctic  borderland  Coprosnia  could  pass  as  of  Antarcto-tertiary 
origin. 

C.  pyrifolia  offers  a  good  example  of  an  incorrect  taxonomic  position  leading 
to  false  conclusions.  It  was  described  as  Psychotria  and  referred  to  a  neotropical 
group:  it  came,  as  Burger  writes  (^j.  20),  "von  den  Urwaldern  Perus  und  Co- 
lombias". 

Of  the  more  than  500  species  described  under  Galitun  about  400  are  distri- 
buted over  the  boreal  zone  and  some  50  or  60  are  known  from  South  America, 
extending  from  the  tropical  Andes  to  Fuegia,  the  Falkland  Is.  and  South  Georgia; 
about  30  are  African,  the  remaining  species  being  divided  between  India,  Malaysia, 
Australia,  New  Zealand,  etc.  Galiiun  is  a  boreal  genus  with  a  strong  represen- 
tation along  the  Andes,  a  not  uncommon  case.  G.  masafuera?inm  is  allied  to 
species  from  the  mountains  of  Central  Chile,  probably  also  to  species  found  farther 
north.  Nothing  indicates  that  Antarctica  ever  had  a  share  in  the  history  of  this 
genus. 

Whether  Campanulaceae-Lobelioideae  evolved  in  the  tropics  or  in  the  far  South 
will  not  be  discussed  here.  The  remarkable  concentration  of  arboreous  genera, 
two  of  them  large,  in  Hawaii,  has  given  rise  to  much  speculation,  and  the  occur- 
rence of  other  endemic  genera  in  Polynesia  as  well  as  the  Australian  affinity  of 
Brighamia  of  Hawaii  have  led  some  authors  to  look  for  the  origin  of  the  sub- 
family (or  family)  in  the  Antarctic.  In  our  special  case  we  can  leave  this  question 
aside  because  Lobelia  alata  is  a  widely  distributed  seaside  species,  tricentric  in 
the  South  Hemisphere. 

The  distribution  of  Wahlenbergia,  a  large  and  widespread  genus,  is  inter- 
esting. It  is  essentially  southern,  of  the  about  230  species  described  150  are  South 
African  and  20  tropical  African,  6  are  reported  from  Madagascar  and  the  Masca- 
rene  islands.  North  it  extends  to  the  Mediterranean,  the  Orient  and  Asia,  together 
some  25  species.  Scattered  species  are  known  from  New  Guinea  (i),  Australia  (7), 
New  Zealand  (7),  Lord  Howe  I.  (2)  and  St.  Helena  (3).  Species  are  few  in  America 
(North  Amer.  i,  South  Amer.  8);  a  single  species,  W.  gracilis,  ranges  over  the 
south  hemisphere. 

If  we  look  at  the  related  genera,  some  13  in  number,  the  dominant  position 
of  South  Africa  becomes  still  more  conspicuous:  7  are  exclusively  or  preponder- 
antly South  African,  i  is  from  tropical  Africa,  E.  Indies  and  Brazil,  4  Asiatic 
and  I  from  S.E.  Europe,  all  according  to  the  old  synopsis  in  Natiirl.  Pflanzenfam- 
Our  island  species  are,  as  we  have  seen,  not  matched  in  Africa,  nor  in  America 
or  other  parts  of  the  world  except  on  St.  Helena,  see  above  p.  210.  All  known 
facts  suggest  that  the  genus  had  a  long  and  complicated  history  in  the  far  south 
and  that  S.  Africa  is  a  secondary  centre  where  evolution  has  been  progressive 
and  prolific  in  species  formation.  It  is  important  to  remember  that  our  island  forms 
are  not  a  group  of  closely  related  taxa  but  that  W.  Masaftierae  and  particularly 
Berteroi  stand  apart  from  the  rest.  The  distribution  of  the  genus  can  hardly  be 
explained  without  admitting  Antarctica  into  the  picture. 


268  C.   SKOTTSBERG 

The  distribution  of  Lagowphora  clearly  testifies  to  its  Antarctic  origin:  3 
Magellanian  species,  one  of  them  also  on  Tristan  da  Cunha  and  one  on  Masafuera, 
7  in  New  Zealand,  2  in  Australia,  one  of  these  north  to  the  Philippines,  i  in  Fiji 
and  3  in  Hawaii,  the  four  Pacific  ones  connected  with  L.  Bil/ardieri  Cdiss.  (Austral.), 
whereas  the  Magellanian  species  point  toward  L.  puiuila  Cheesein.  2iX\d.  petiolata 
Hook.   fil.  (New  Zealand). 

In  Iiyii:;tyo}i  we  meet  again  with  a  large  boreal  genus  strongly  represented 
along  the  Andes  and  ranging  south  to  Patagonia,  P^iegia  and  P'alkland;  135  species 
are  reported  from  Kurasia,  inch  India,  345  from  North  America,  35  from  Centr. 
America  and  the  W.  Indies,  95  from  S.  America,  10  from  Africa,  8  from  Australia 
and  a  single  species  from  Oceania  (Rapa),  E.  rapensis  F.  B.  H.  Brown,  and  this  is 
compared  with  our  island  species.  To  judge  from  the  description  it  has  the  same 
habit  as  these,  a  shrub  about  3  dm  tall  with  the  leaves  in  terminal  rosettes.  In 
I  lawaii  the  genus  is  represented  by  l^ctra)nolopiu7)i,  which  is  very  close  to  Erigeroit\ 
\'lKKii.\ri'KR  [22g.  182)  suggested  that  the  Juan  P^rnandez  species  came  near  the 
Hawaiian  /:.  lepidotus  Less.,  which  is  now  referred  to  Tetraniolopmnt,  and  that  we 
ought  to  look  for  relatives  among  the  Andine  and  Mexican  forms.  Among  our 
island  species  li.  nipicola  differs  much  in  habit  from  the  rest,  but  all  belong  together 
and  most  likely  represent  a  special  branch  of  the  Andine  Erigeroii  flora  which,  in 
its  turn,  comes  from  a  boreal  stock. 

To  what  I  said  above  about  GnaphaEiuj>i  spicifornie  nothing  can  be  added 
at  present.  Chile  is  well  provided  with  poorly  limited  species  related  to  G.  pur- 
pur  ciiui  L.;  most  of  them  are  badly  known.  Their  boreal  parentage  can  hardly 
be  doubted. 

Ahrotajiclla  resembles  LagejiopJiora  in  its  distribution,  but  is  absent  from  Oce- 
ania; tiie  majority  inhabits  New  Zealand  with  its  subantarctic  islands  (9  sp.),  i  is 
found  in  X'ictoria,  2  in  Tasmania,  i  in  New  Guinea  and  5  in  S.  America  (W. 
Patagonia  to  h\iegia  and  P'alkland);  in  addition,  one  is  found  on  Rodriguez  I. 
Tiie  genus  is,  as  it  were,   tricentric  and  its  Antarcto-tertiary  character  indisputable. 

The  concentration  of  isolated  arborescent  Compositae  in  the  Pacific  was  em- 
|)hasized  by  Bkn'IIIAM;  tiie  main  grouj)S  of  the  family  are  represented  among  them^ 
and  the  accunuilation  of  endemic  genera  in  Hawaii  and  Juan  P^rnandez  has  led 
to  nuich  discussion.  Tiic  enormous  development  and  differentiation  in  the  family 
on  American  soil  is  an  undeniable  fact,  but  simply  to  derive  the  endemic  Pacific 
genera  from  America  as  did  (ill'l'V  [121)  does  not  seem  [)ossible.  Speculations  (comp. 
f.  inst.  Sl'/i  (111:1.1,  2r())  led  back  to  Antarctica,  but  not  until  BracJiioioslyluin  was 
discovered  in  .\ew  (iuinea  and  found  to  be  nearly  related  to  Robinsoiiia  o'i  ]\x^Vi 
l^>rnandez  were   we  able  to  stand   on   tolerably  firm   ground. 

With  regard  to  Geiitaurodoidroji,  to  which  J//';/*//^'^^'?  seems  to  be  related,  the 
situation  is  different.  Coitaurca  and  all  the  genera  of  Centaureinae  belong  to  the 
Old  World  with  the  exce})tion  of  a  single  species  in  N.  America  and  a  few  in 
the  Andes,  south  to  Centr.  Cliile.  Tlie  grou{)  ranges  over  luirope  with  a  strong- 
hold in  the  Mediterranean,  the  Orient,  P'.thiopia  and  through  Centr.  Asia  to  Japan. 
The  Chilean  species  of  Centaurca  belong  to  the  Plectocephali,  but  Ceiitauroden- 
droji  differs  not  only  from  this  section   but  from  all  in  sex  distribution  (24J).  It 


DERIVATION  OF  THE  FLORA  AND  FAUNA  269 

serves  no  purpose  to  say  that  Centaurodendron  descends  from  some  continental 
Centaurea  which  became  arboreous  under  insular  conditions,  because  the  character 
of  the  ray  floret  is  entirely  opposed  to  such  a  theory  and  Centaurodendron  seems 
to  be  a  more  ancient  type  than  Centaurea,  a  relic  from  an  epoch  previous  to  the 
final  uplift  of  the  Andes, 

The  four  Cichoriaceous  genera,  to  which  Z//<3'w;/d7^^;7V  of  the  Desventuradas 
Is.  shows  some  slight  affinity,  are  even  more  isolated.  The  only  other  genus  to 
which  they  bear  some  resemblance  is  the  Polynesian  Fitchia,  but  the  differences 
are  too  profound  to  allow  us  to  visualize  an  Antarctic-Pacific  ancestry  of  the  Den- 
droseris  group  and  it  remains  us  to  link  it  with  the  neotropical  element. 

Referring  to  the  synopsis  given  above  the  Phanerogams  are  rearranged  in 
the  following  way  according  to  their  supposed  source  of  origin. 

I.  Antarcto-tertiary  element. — 62  sp.  (42.2  %). 

1.  Distribution  pattern  austral-circumpolar,  bicentric  or  tricentric. — 33  sp. 

a.  Endemic  species  (17):  Cladium  scirpoideum,  Uncinia  Douglasii  and  costata, 
Carex  berteroniana,  Luzula  masafuerana,  Drimys  confertifolia,  Phrygilanthus  Ber- 
teroi,  Acaena  masafuerana,  Sophora  fernandeziana  and  masafuerana,  Gunnera  pel- 
tata,  Masafuerae  and  bracteata,  Apium  fernandezianum  (.^),  Pernettya  rigida,  Plan- 
tago  fernandezia,  Abrotanella  crassipes. 

b.  Species  also  found  on  the  continent  (16):  Danthonia  collina,  Koeleria  micra- 
thera  (.f"),  Oreobolus  obtusangulus,  Uncinia  brevicaulis,  phleoides  and  tenuis,  Juncus 
imbricatus,  capillaceus,  acutus,  dombeyanus  and  planifolius,  Libertia  formosa, 
Acaena  ovalifolia,  Centella  triflora,  Nertera  granadensis,  Lagenophora  Harioti. 

2.  Genera  only  found  in  S.  America,  but  supposed  to  be  of  Antarctic  origin. 
The  species  are  endemic:    Escallonia  Callcottiae,  Margyricarpus  digynus. — 2  sp. 

3.  Endemic  genera  or  species  as  far  as  known  without  continental  American 
affinities,  either  suggesting  an  ancient  Antarcto-Pacific  track  east  from  Australasia 
without  reaching  America,  or  having  arrived  along  the  road  over  the  Scotia  Arc 
without  leaving  any  traces  in  the  present  S.  American  flora. — 21   sp. 

a.  Belonging  to  endemic  genera  (7):  Robinsonia  gayana,  thurifera,  evenia,  Masa- 
fuerae and  gracilis,  Symphyochaeta  macrocephala,  Rhetinodendron  Berterii. 

b.  Endemic  species  of  genera  of  wider  distribution  (14):  Peperomia  berteroana, 
margaritifera  and  Skottsbergii,  Santalum  fernandezianum,  Boehmeria  excelsa,  Ra- 
nunculus caprarum,  Fagara  mayu  and  externa,  Halorrhagis  asperrima,  masatier- 
rana  and  masafuerana,  Euphrasia  formosissima,  Coprosma  Hookeri  and  pyrifolia. 

4.  Endemic  species  with  relatives  in  the  African  sector  only:  Wahlenbergia 
Larrainii,  fernandeziana,  Grahamae,  Masafuerae  and  Berteroi. — 5   sp. 

5.  Endemic  family,  possibly  of  old  Antarctic,  perhaps  pre-Tertiary  ancestry: 
Lactoris  fernandeziana. —  i   sp. 

II.  Neotropical- Andean  element. — 54  sp.  (36.7  %). 

I.  Endemic  genera  or  species  of  neotropical  parentage;  non-endemic  species 
South  American. — 31  sp. 

a.  Belonging  to  endemic  genera  (4):  Juania  australis,  Ochagavia  elegans,  Notho- 
myrcia  fernandeziana,  Selkirkia  Berteroi. 


270  C.   SKOTTSBKRG 

b.  luidcmic  species  of  non-endemic  genera,  some  of  wider  distribution  (14): 
Chaetotropis  imberbis.  Chusquea  fernandeziana,  Hesperogreigia  Berteroi,  Urtica 
Masafiierae,  Dysopsis  hirsuta.  Colletia  spartioidcs,  Azara  fernandeziana,  Ugni  Sel- 
kirkii,  Myrceugenia  Schulzei,  Rhaphithamnus  venustus,  Solanum  fernandezianum, 
robinsonianum  and  masafiieranum,   Xicotiana  cordifolia. 

c.  Sj)ecies  also  found  on  the  continent  (13):  Stipa  neesiana  and  laevissima, 
Piptochaetium  bicolor,  Chaetotropis  chilensis.  Cyperus  eragrostis  and  reflexus,  Eleo- 
charis  fuscopurj)urea,  Peperomia  fernandeziana,  Parietaria  humifusa,  Myrteola  num- 
mularia,   Minnilus  ghibratus,  Phuitago  truncata,   Hedyotis  thesiifolia. 

2.  luidemic  genera  or  endemic  sections  of  wide-ranging  genera  without  relatives 
in  the  present  continental  flora  but  supposed  to  descend  from  extinct  neotropical 
ancestors. — 23  sp. 

a.  Ik'longing  to  endemic  genera  (16):  Podophorus  bromoides,  Megalachne  ber- 
teroniana  and  masafuerana,  Centaurodendron  dracaenoides,  Yunquea  Tenzii,  Den- 
droseris  macrophylla,  macrantha,  marginata  and  litoralis,  Rea  neriifolia,  micrantha 
and   j)ruinata,   IMioenicoseris  j^nnata,  berteriana  and  regia,   Hesperoseris  gigantea. 

h.  Ik'longing  to  endemic  sections  (7):  Urtica  fernandeziana,  Chenopodium 
Sanctae  Clarae,  crusoeanum  and  nesodendron,  Eryngium  bupleuroides,  inaccessum 
and  sarc()j)h\'llum. 

III.   Arcto-tertiary  element. —  23  sp.  (15.6%). 

Cicncra    essentially    boreal   but  extending  south  along  the  Andes  or  reappearing  in  S. 

America. 

a.  luidemic  species  of  wide-ranging  genera  (13):  Agrostis  masafuerana,  Sper- 
gularia  confertiflora  (also  Desventuradas)  and  masafuerana,  Berberis  corymbosa 
and  masafuerana,  Cardamine  Kruesselii,  Galium  masafueranum,  Erigeron  fruticosus, 
luteoviridis,   Ingae,   Innocentium,  turricola  and  rupicola. 

b.  Species  also  fountl  on  the  continent  (lo):  Trisetum  chromostachyum,  Carex 
Banksii,  Paronychia  chilensis,  Cardamine  chenopodiifolia  and  flaccida,  Rubus  geo- 
ides,  Callitriche  Lechleri,  P^mpetrum  rubrum,  Calystegia  tuguriorum(.?),  Gnaphalium 
s{)ici  forme. 

IV.   Palaeotropic  element. —  2  sp.  (1.4%). 

Plndemic  genus:   Cuminia  fernandezia  and  eriantha. 

V.   Austral  element  of  wide-ranging  seaside  species. — 6  sj).  (4,1  %). 

Scirpus  nodosus  and  ccrnuus,  Salicornia  fruticosa  (peruviana),  Tetragonia  ex- 
[)ansa.  Dichondra  repens.    Lobelia  alata. 

I'Acn  if  Antarctica  is  recognized  as  an  important  source  of  evolution  of  both 
plants  and  animals  and  as  a  centre  from  where  large-scale  migration  took  place, 
a  percentage  of  42  may  seem  surprisingly  high,  and  I  admit  that  some  genera 
or  species  have  been  referred  to  group  I  with  considerable  hesitation.  This  does 
not,  with  very  few  cxceptif)ns,  aj)i)ly  to  I.  i,  22.4%  of  the  total  or  53.2%  of  I. 
Luzula  and  Jiiucus  were  [)laced  here  because  the  family  was  regarded  as  Antarctic, 
but  even   if  this  be  true  it  is  po.ssible  that  the  sections  including  our  species  are 


DERIVATION  OF  THE  FLORA  AND  FAUNA  27I 

of  boreal  origin  and  has  spread  south.  Pernettya  is  a  bicentric  genus,  but  it 
is  much  better  developed  in  America  and  may  have  originated  with  Gaultheria 
in  the  Antarctic.  The  same  should  apply  to  Nicotiana  according  to  Merrill 
(jo<5.  310)  who  thinks  that  its  actual  distribution  was  attained  in  the  Tertiary  pe- 
riod by  way  of  Antarctica;  however,  N.  cordifolia  cannot  be  removed  from  II.  i.  b 
and  find  a  better  place  with  I.  i.a.  On  the  other  hand,  a  truly  Antarctic  genus 
may  have  produced  numerous  species  in  one  sector  and  few  in  another,  or  the 
few  may  be  a  remnant  of  a  larger  population.  In  group  I.  3  Peperomm  berteroana 
calls  for  attention.  As  I  have  shown  (^^5)  it  is  so  close  to  P.  tristanejish  that 
their  common  origin  cannot  be  doubted  and  that  little  prevents  us  from  regarding 
them  as  forms  of  one  species,  very  distinct  but  clearly  related  to  the  other  two 
species  endemic  in  Juan  Fernandez,  and  they  point  west.  This  is  the  reason  why 
all  three  were  referred  to  I.  Wahlenbergia  (I.  4)  is  another  mysterious  case,  but  as 
it  is  neither  boreal  nor  palaeotropical  or  neotropical  but  South  African  (and  on  St. 
Helena),  only  a  far  southern  ancestry  remains  to  explain  the  disjunction. 

Of  the  endemic  genera  belonging  to  group  II.  i  only  Juaiiia  is  taxonomically 
isolated,  Rhodostachys  of  Chile  is  brought  to  Ochagavia  by  some  authors,  and 
Selkirkia  comes  very  near  Hackelia.  With  regard  to  the  endemic  Cichoriaceous 
genera  authors'  opinions  differ.  Bentiiam  looked  upon  the  arborescent  Pacific 
Compositae  as  relics  of  an  old  Polynesian  flora  but  did  not  refer  directly  to  Ant- 
arctica as  source;  Guppy  {^121]  quoting  Bentham  believed  that,  with  the  Hawaiian 
Lobelioideae,  they  belonged  to  an  ancient  flora  of  the  Pacific  which  had  origi- 
nated in  America  and  gained  dominance  during  what  he  termed  "Age  of  Lobe- 
liaceae"  and  "Age  of  Compositae",  respectively.  Antarctica  seems  not  to  have 
meant  anything  at  all  to  him,  but  as  I  have  discussed  his  ideas  in  some  detail 
in  an  earlier  paper  [2/f.8),  I  shall  not  enter  upon  this  subject  here. 

Calystegia  tuguriorum  is  doubtful  case,  but  few  people  will  be  inclined  to 
think  that  it  originated  independently  in  Chile  and  New  Zealand. 


In  a  paper  of  1928  [^218)  Setchell  discussed  what  he  called  the  two  prin- 
cipal elements  in  the  Pacific  flora,  the  Indomalayan  and  the  "Subantarctic".  With 
this  he  did  not  understand  what,  from  a  geographical  viewpoint,  I  call  subant- 
arctic, nor  the  species  of  old  Antarctic  genera  found  in  the  bogs  of  Hawaii,  the 
Pacific  flora  with  which  he  was  particularly  concerned,  but  the  group  he  later 
{2ig)  called  "the  Old  pacific  and  antarctic  element",  for  it  included  also  the  ar- 
boreous Lobeliaceae  and  Compositae,  which  he  believed  had  a  common  origin 
in  high  southern  latitudes.  They  had  migrated  north  along  different  lines,  the 
Compositae  taking  an  easterly  course  along  the  route  Juan  Fernandez-Hawaii,  the 
Lobeliaceae,  which  are  absent  from  Juan  Fernandez  but  have  left  traces  in  Poly- 
nesia, a  more  westerly;  a  third  line  is  called  the  Dammara  (Agathis)-Podocarpus 
line,  an  Australasian  line  running  over  Fiji.  What  interests  us  here  is  that  the 
isolated  genera  of  Compositae  were  regarded  as  Antarctic.  As  we  have  seen,  I 
did  not  venture  to  include  the  Cichoriaceous  genera,  but  I  cannot  assure  that 
Setchell  was  wrong. 


272 


C.   SKOTTSBKRG 


Before  quotin*]^  some  other  authors  who  have  paid  special  attention  to  the 
Antarctic  i)roblems  I  want  to  add  a  few  remarks  on  the  relations  between  the  far 
South  and  South  America-Juan  Fernandez.  The  Antarctic  flora  invaded  the  Amer- 
ican continent,  advancinj^  especially  along  the  rising  Andes  and  in  some  cases 
extending  north  of  the  Iu]uator.  It  must  have  been  a  climatically  diversified  flora, 
for  the  great  southern  continent  must  have  had  coast  and  inland,  lowland  and 
mountain  climates,  and  consequently  the  Antarctic  plants  in  S.  America  have 
varying  claims  on  moisture  and  heat.  The  less  exacting  plants  are  concentrated 
in  West  Patagonia,  r\iegia  and  the  Falkland  Is.,  rising  higher  and  higher  as  they 
advance  north  along  the  Cordillera,  whereas  such  as  require  more  favourable  tem- 
perature conditions  are  found  farther  north  in  S.  Chile  and  in  the  montane  region 
of  the  tropical  Andes.  I^^ather  Rambo,  a  wellknown  expert  on  the  flora  of  S.  Brazil, 
called  my  attention  (letter  of  Nov.  30,  1953)  to  an  Antarctic  element  in  the  high- 
lands of  Rio  Grande  do  Sul,  where  such  genera  as  Acae?ia,  Margyricarptis,  Es- 
callojiia,  GuuJicra,  /)r/;^n's and  PJirygilanthusdiX^  represented;  in  202.  30  he  mentions 
Ayaucaria  ajigusiifolia,  Podocarpus  Lmnberti,  Drimys  Wmteri,  Acae7ia  fuscesceiis, 
h'tichsia  regia,  Ciumicra  manicata,  and  Griselhija  riiscifolia,  elements  which,  in  his 
opinion,  constitute  the  last  remnants  of  the  old  flora  that  inhabited  the  southern 
lands  united  until  early  Tertiary  times. 

IliLL  (/iV)  was  cautious  in  his  judgment  of  the  importance  of  Antarctica  in 
the  history  of  the  plant  world.  He  quotes  Seward  who  had  pointed  out  that  a 
number  of  families  now  largely  in  the  South  Hemisphere  were  present  in  the  North 
Hemisphere  already  in  Mesozoic  times  and  that  this  would  lead  us  to  derive  them 
from  there,  which  most  likely  is  true  in  certain  cases.  He  also  refers  to  SMUTS, 
who  looked  at  the  "ancient  lands  of  the  Southern  Hemisphere"  as  the  cradle  of 
the  peculiar  S.  African  flora,  and  HiLL  formulates  the  following  question  (p.  1497): 

Did  the  angiosperms  originate  in  the  north  and  migrate  southwards  and  then,  having 
rea(  lied  the  south,  evolve  along  special  lines  in  lands  comparatively  isolated  from  the 
land  masses  in  the  north,  or  did  certain  grouj)S  first  ap])ear  in  the  South  Hemisphere 
in  an  ancient  Antarctic  Continent  and  become  dis})ersed  northward  into  our  j)resent- 
(lay   New   Zealand,   Australia,    South   America  and   South   Africa? 

With   regard  to  Juan    h'ernandez  he  finds  that 

the  present  flora  suggests  that  at  one  time  these  islands  formed  part  of  the  Antarctic 
continent  or  were  united  to  the  extreme  south  of  Chile,  a  view  which  is  shared  by 
Jhering  and  Joly,  who  believe  that  such  a  connection  may  have  existed  in  early  Ter- 
tiary times  and  that  Kerguelen  Island  was  prol)al)ly  part  of  a  large  land  mass  at  the 
same   period  .  .  . 

and  he  continues: 

It  seems  also  likely,  on  geological  evidence,  that  Antarctica  extended  in  the  Ter- 
tiary ei)0ch  towards  Tasmania  and  Australia,  and  so  to  Asia,  and  towards  New  Zea- 
land, and  the  distribution  of  certain  j)resent-day  ])lants  in  the  Australasian  region  lends 
consideral)]e  suj)port  to  such   an   extension   (p.  1479). 

Nevertheless,  the  problem  is  not  as  simple  as  it  might  seem  to  be: 


DERIVATION  OF  THE  FLORA  AND  FAUNA  273 

We  must  be  prepared  to  modify  our  views  extensively  in  those  cases  which  appear 
to  afford  fairly  certain  evidence  of  distribution  from  a  Southern  centre,  but  from  the 
evidence  available  it  seems  probable  that  such  genera  as  Calceolaria,  Jovellana,  Fuchsia, 
Ourisia,  Pelargonium,  Caltha,  Lilaeopsis,  Gunnera,  Hebe,  Fernettya,  Azorella,  Drapeies 
and  other  Thymelaeaceae,  Noihofagus,  Eucryphia,  Laurelia,  probably  the  Proteaceae, 
some  of  the  Ericaceae  .  .  .  may  have  originated  in  some  continental  area  in  the  South 
Hemisphere,  whence  their  descendants  spread  northwards  ... 

It  is  of  course  possible  that  in  early  times  many  of  the  plants  from  w^hich  these 
Southern  Genera  evolved  belonged  to  the  Northern  region  and  gradually  migrated  to 
the  south  and  having  reached  the  Southern  regions  they  there,  in  a  state  of  some  isola- 
tion, evolved  peculiar  types  which  in  their  turn  spread  again  northwards  .  .  .  without 
however  crossing  the  Equator  and  reaching  the  North  Hemisphere.  On  this  view  we 
may  perhaps  best  regard  many  of  the  present  day  genera  which  are  now  only  found 
in  the  South  Hemisphere  (p.  1480). 

He  adds, 

It  may  be  almost  impossible  in  these  latter  days  to  make  any  certain  pronounce- 
ment as  to  which  natural  families  or  genera,  if  any,  actually  originated  in  the  Southern 
Hemisphere,  w^hen  it  is  remembered  that  there  have  been  a  succession  of  alternations 
of  warm  temperate  and  cold  glacial  periods  at  various  geological  epochs  not  only  in 
the  Arctic  but  also  in  the  Antarctic  regions. 

He  thinks  that  the  cold  spells  gave  rise  to  an  extensive  migration  south  and 
north,  respectively,  but  he  did  not  recognize  the  glaciations  as  contemporary  in 
the  two  hemispheres. 

Of  other  modern  authors,  W.  H.  Camp  (4.J)  interests  us  here.  My  Antarcto-tertiary 
element  lists  Drimys  as  a  classical  example  of  an  Antarctic  genus;  in  fact,  the 
entire  family  Winteraceae  is  supposed  to  be  of  Antarctic  origin.  On  Camp's  map, 
however,  the  original  seat  of  the  family  is  placed  in  the  region  of  the  old  Bering 
Land  bridge,  from  where  tracks  lead  south  through  Asia  to  Australia-Tasmania 
and  through  western  N.  America  to  S.  America.  The  related  Magnoliaceae  are 
circumboreal  and  have  left  numerous  fossil  remains  in  the  north.  Another  map 
shows  Laurelia,  Eucryphia,  Luzuriaga  and  Jovellana,  all  as  a  rule  considered  to 
be  Antarctic;  here  Camp  has  two  alternatives,  one  track  leading  from  the  north 
as  in  Drimys,  another  from  the  south.  Other  southern  families  shown  on  his  maps 
are  Tetrachondraceae,  Eucryphiaceae,  Centrolepidaceae,  Epacridaceae  (Hawaii  in- 
advertently left  out),  Stylidiaceae,  Restionaceae,  Liliaceae-Milliganieae,  Philesiaceae, 
Halorrhagidaceae  and  Gunneraceae.  He  emphasizes  the  fact  that  of  the  natural 
families,  some  300  in  number,  103  have  a  fairly  restricted  range,  and  that  80  of 
these  are  on  the  South  Hemisphere.  I  shall  quote  him  in  full. 

In  the  majority  of  instances  an  analysis  of  a  group  indicates  that  the  primitive 
members  are  on  the  southern  part  of  the  group's  range.  If  we  were  to  follow  the  well- 
known  dicta  of  Matthews  we  could  conclude  that  these  primitive  forms  were  "driven" 
unto  the  southern  land  mass  extremities  by  the  more  highly  specialized,  better  adapted, 
and  more  recently  derived  groups  of  the  north.  The  natural  corollary  to  this,  therefore, 
is  that  we  should  find  the  majority  of  the  peculiar  specialized  and  more  recently  derived 
families  farthest  away  from  the  primitive  forms.  However,  as  can  quite  easily  be  seen 
in  the  angiosperms,  in  most  instances  this  is  not  the  case.  I  therefore  incline  to  the 
conclusion  that  these  southern  land  masses  are  not  only  the  original  homes  of  the  great 

18  -  557857  T^^e  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.      Vol.  I 


74 


C.   SKOTTSBERG 


majority  of  our  basic  angiosi)erm  groups,  but  that  also  it  has  been  on  these  same  southern 
land  masses  where  the  greater  part  of  their  primary  evolutionary  divergences  took  place 
(pp.  180-181), 

and  he  continues, 

It  seems  likely  that  the  angiosperms,  as  a  group,  arose  on  this  southern  land  mass 
contemporaneously  with  the  Paleozoic  of  the  northern  (Holarctic)  land  mass  and  that 
the  divergences  of  the  basic,  generalized  familial  groups  had  been  accomplished  in  this 
southern   hind   mass   certainly   by  the   mid-Mesozoic. 

This  would  make  the  Jurassic  the  great  period  of  ascendency  of  the  angiosperms. 

C'ami's  theory  does  not,  at  least  at  first  sight,  agree  too  well  with  his  opinion 
on  the  origin  of  the  Magnoliales,  always  regarded  as  a  primitive  group  though 
not  of  restricted  range;  the  Winteraceae  have  a  stronghold  in  New  Guinea-Australia- 
Xew  Zealand,  and  the  actual  distribution  of  this  family  testifies  to  a  southern  origin. 

Recently  SlKHBlNS  (^57)  took  up  the  question  of  the  history  of  the  Ranales. 
He  points  out  (p.   8)  that  this  order  » 

includes  a  high  j)roj)ortion  of  sj)ecies  which  on  the  basis  of  all  characteristics  must  be 
])laccd  not  only  in  monotypic  genera  but  even  in  monogeneric  or  digeneric  families. 
Thcv  are  obviously  relict  types  of  which  the  close  relatives  have  long  been  extinct.  .  .  . 
Finally,  distributional  studies  show  that  the  genera  and  species  are  at  present  strongly 
concentrated  in  eastern  Asia  and  Australasia,  and  at  least  one  family,  the  Winteraceae^ 
may  have  radiated  from  the  latter  center  (A.  C.  Smith  1945).  This  family  was  dispersed 
through   the   Antarctic  regions. 

We  cannot  deny^  the  possibility  that  New  Guinea  was  the  birthplace  of  types 
that  now  generally  pass  as  Antarctic,  a  theory  first  advanced  by  Miss  GiBBS  (lod) 
and  perhaps  strengthened  by  the  sensational  discovery  there  of  a  great  number  of 
Xothofagus  species,  but  it  is  quite  clear  that,  if  they  are  bicentric,  they  must  have 
migrated  across  what  is  now  the  Antarctic  continent  in  order  to  arrive  in  S.  America 
which  again  com[)els  us  to  assume  that  connections  were  established  on  both  sides, 
and  the  existence  of  fossil  plants  belonging  to  taxa  now  living  in  the  two  sectors 
furnishes  additional  evidence  that  Antarctica  took  an  important  part  in  their  history. 
Anyhow,  Xothofa^i^iis  survives  in  greater  variation  in  New  Guinea  than  either  in 
New  Zealand  or  in  Chile,  though  these  are  situated  nearer  to  the  Antarctic  continent. 

Before  Antarctica  was  recognized  as  a  possible  centre  of  evolution,  the  Holarctic 
region  was  claimed  as  the  great  and  only  cradle  of  temperate  plant  families  which 
s{)read  soutii  during  the  Tertiary  just  as,  in  pre-Tertiary  times,  tropical  families 
had  extended  to  the  j)rcsent  Arctic.  GoRDuX  (77?),  although  he  finds  undeniable 
proofs  of  an  old  Antarctic  radiating  centre,  carefully  scrutinizes  the  possibility  of  a 
northern  origin  of  temperate  types  now  found  in  the  south,  referring  to  Oliver's 
theory  that  :i  genus  like  Xothofagus  originated  in  \.  America  and  that  two  tracks 
lead  sf)uth,  one  to  subantarctic  America,  one  across  the  Ikring  land  to  Australia 
and  Xew  Zealand,  but  he  doesn't  think  it  probable  that  we  can  explain  the  dis- 
junctions of  numerous  taxa  in  the  south  in  this  way  and  that  the  chances  for  cool 
climate  types  to  cross  the  broad  and  well-stocked  tropical  belt  must  have  been  small. 

To  me  it  would  seem  more  acceptable  to  fall  back  on  an  ancient  pantropical 


DERIVATION  OF  THE  FLORA  AND  FAUNA  275 

flora,  spread  from  pole  to  pole  before  any  temperate  zones  had  become  sharply 
delimited  and  when  the  distribution  of  land  and  sea  was  quite  different  from  what 
it  is  now.  With  the  appearance  of  distinct  climatic  belts  a  sorting  out  of  meso- 
and  microthermic  groups  followed,  and  these  belts,  both  north  and  south,  started 
to  produce  their  own  particular  new  groups  which  dispersed  toward  the  equatorial 
zone  and  eventually  met  and  passed  it  where  mountain  ranges  offered  a  passage. 
This  is  in  agreement  with  Florin's  opinion  (pj)  that  the  temperate  floras,  angio- 
sperms  as  well  as  gymnosperms,  developed  independently  throughout  the  Tertiary 
period  in  the  north  and  south  hemispheres.  The  march  of  Antarcto-tertiary  types 
north  corresponds  to  the  march  of  Arcto-tertiary  types  south. 

Recently  Antarctica  and  the  far  south  in  general  as  the  birthplace  of  the  world's 
floras  has  found  an  eloquent  advocate  in  Ll^ON  Croizat  (yi),  who  throws  all  other 
speculative  authors  in  the  shade.  His  ideas  were  criticised  by  Sparre  (2j6),  but, 
as  Juan  Fernandez  is  regarded  by  Croizat  as  "one  of  the  most  interesting  domains 
of  general  phytogeography",  I  cannot  pass  him  in  silence. 

The  original  southern  plant  world,  Croizat  says,  spread  over  the  globe  in 
a  way  not  to  be  deducted  from  the  present  map;  we  have  to  go  back  to  the  map  as 
it  looked  during  the  Cretaceous,  but  he  does  not  tell  which  of  the  constructions 
he  prefers,  only  that  he  dismisses  Wegener's  theories;  that  everybody  who  devotes 
himself  to  the  study  of  plant  migrations  should  pay  attention  not  only  to  the  present 
configuration  of  the  continents  goes  without  saying.  His  own  ideas  of  Cretaceous 
geography  in  the  south   are  expressed  on  p.   252: 

It  is  most  probable,  sure  we  would  venture  to  say,  that  an  ancient  shore  connected 
every  land  in  the  deep  south  of  our  maps,  not  only,  but  that  the  lands,  now  vanished, 
but  once  extant  between  the  approach  of  Kerguelen  Islands  and  Magellania  are  the  ultimate 
hub  of  angiospermous  dispersal. 

Here  extends  the  Patagonian-Mascarene  baseline  with  its  two  triangles,  Natal-Ker- 
guelen-Tristan  da  Cunha  (the  Afro-antarctic  triangle)  and  Ceylon-Madagascar- 
Mascarenes,  the  "hub"  spoken  of  above.  Croizat  points  at  South  Georgia  as  a 
proof  of  the  existence  of  an  old  Antarctic  flora: 

The  theory  that  a  nebulous  'Glacial  Epoch'  killed  off  very  nearly  all  the  ancient 
vegetation  of  the  antarctic  islands  is  shown  to  be  false  by  the  comparatively  large  plant- 
world  still  endemic  in  South  Georgia,  indeed  a  sizeable  flora  if  we  consider  that  it  thrives 
in  what  is  now  hardly  better  than  a  glaciated  mountain  straight  out  of  the  ocean  (p.  255). 

Now,  Croizat  otherwise  deals  with  the  phanerogams  only;  there  is  not  a  single 
endemic  species  in  South  Georgia,  but  there  are  many  endemic  bryophytes  and 
lichens.  And  he  forgets  to  mention  that  the  "nebulous"  glacial  epoch  has  left  its 
very  distinct  marks  there,  because  we  know  that  the  lowland  was  ice-covered  where 
the  phanerogams  now  form  closed  communities  and  that  it  is  very  improbable  that 
the  higher  flora  survived.  It  is  surprising  that,  in  this  connection,  he  does  not  men- 
tion Kerguelen  where  endemic  genera  such  as  Pringlea  and  Lyallia  surely  survived. 
A  second  baseline  indicates  the  extension  of  an  Antarcto-Gondwana  in  the 
S.  Indian  Ocean,  a  third  lies  along  60°  S.  and  runs  from  south  and  east  of  Chatham 
Is.  to  contact  S.  America.  From  these  centres  the  angiosperms  started  to  disperse 


276 


C.  SKOTTSBERG 


during  late  Jurassic  and  very  early  Cretaceous  times  through  three  "gates"  in  three 
main  streams,  the  West  Polynesian,  the  Magellanian  and  the  African  gate.  Many 
of  the  genera  occurring  in  Juan  l>>rnandez,  endemic  or  not,  are  mentioned  by  Croizat  ; 
several  of  them  are,  also  in  my  opinion,  of  Antarctic  origin. 

Onoholus.  Instead  of  allowing  it  to  enter  S.  America  through  the  "Magellanian 
gate  '  Ckoiza  r  makes  a  circuit  along  his  Pacific  baseline  from  New  Zealand  to  F'uegia. 

Juauia.  I  ha\  e  already  discussed  the  taxonomic  position  of  this  genus  which 
Ckoi/ai'  derives  from  a  Mascarene  centre  in  company  with  related  Andine  genera. 
Ik  nillNSON  linked  it  with  Ceruxyloii  to  form  a  tribe  Ceroxyleae;  Croizat  remarks 
that 

this  mav  be  better  taxonomy,  we  do  not  know,  but  does  not  change  substances  in  the 
least,  'i'he  ingredients  that  make  up  the  Morenieae,  Iriarteae  and  Ceroxyleae  are  all  from 
the  same  kitchen.  In  one  case  Hyophorbe  speaks,  in  another  so  does  the  face  of  dispersal. 
Kvervthing  in  the  end  rhymes  to  the  very  same.  With  or  without  Hyophorbe,  seasoned 
with  this  or  that  sauce,  the  stew  does  not  change  .  .  .  classification  of  such  a  genus  as 
this  is   bound    forever  to   remain   the   j)laything  of  opinion   (pp.   456-457), 

a  statement  quite  characteristic  of  this  author's  method  of  approaching  a  phyto- 
geogra[)hical  problem. 

Pipcyo))iia  has  its  stronghold  in  tropical  America  where  a  number  of  subgenera 
are  found.  To  CROIZAT  the  genus  is  Pacific  and  Juan  F'ernandez  one  of  the  starting- 
points  of  its  dispersal  toward  the  heart  of  S.  America,  the  four  island  species  leading 
straight  to  north  Chile,  lie  based  this  opinion  on  YUNCKER  who  wants  to  bring 
them  to  subgen.  SphacrocarpidiuDi^  where,  however,  only  one  of  them  belongs,  while 
the  other  three,  as  was  brought  forth  above  (p.  259),  point  west,  one  of  them  inti- 
mately related  to  P.  ty'istanciisis,  which,  unexpectedly  it  is  true,  brings  the  Antarctic 
within  sight.  In  a  footnote  CkOlZA  T  admits  that  I  may  be  right  in  my  opinion  of 
the  taxonomic  {)osition  of  P.  bcrtcroana\  none  the  less  he  finds  the  contact  Juan 
hYM'nandez- Tristan  not  at  all  j)eculiar.  EDipetruni  is  another  example,  both  "a 
mere  asj)cct  f)f  the  standard  dispersal  between  Africa  and  South  America"  (p.  102). 
Hut  tiiis  "standard  dispersal",  as  discussed  by  Engler,  refers  to  a  number  of 
tr()j)ical  genera  and  families  bearing  witness  of  a  transatlantic  bridge.  In  Croizat's 
opinion   the  track   Mascarenes-Africa-America  was  a  very  important  route. 

I'alms  and  Pepcromias  are  pantropical  and  range  south  to  New  Zealand  and 
central  Chile,  and  there  are  man\'  endemic  palm  genera  on  the  Indian  and  Pacific 
islands  south  of  the  Ivpiator,  whence  follows  that  the  field  is  open  for  speculation 
and  that  the  possibilit)-  of  a  ])rimary  centre  in  the  south  may  be  discussed  without 
breaking  the  rules  of  the  game,  but  to  derive  exclusively  neotropical  families  from 
a  Mascarene  centre  cannot  fail  to  cause  astonishment.  Croizat,  dealing  with  Cac- 
taceae,  Bromeliaceae,  Kai)ateaceae  etc. — and  in  an  earlier  chapter  he  asserts  that 
Malesherbiaceae  also  had  their  starting  j)oint  in   Africa — says  that 

to  credit  grouj)s  of  the  kind  with  "American"  origin  steps  must  be  taken  as  a  prelim- 
inary which  no  one  trained  in  |)rccise  thinking  will  encourage.  .  .  .  Rhipsalis  is  doubtless 
a  (iondwanic  genus  because  it  is  rooted  within  the  triangle  Ceylon-Madagascar-Masca- 
renes.  It  well  may  have  reached  ]')razil  and  the  West  Indies  from  West  Africa.  .  .  .  the 
baseline  of  the  Cactaceae  scpiarely  rests  between  Patagonia  and  the  Mascarenes,  and  if 


DERIVATION  OF  THE  FLORA  AND  FAUNA  277 

Rhipsalis  is  Gondwanic  it  is  likely,  to  say  the  least,  that  the  archetypes  of  the  family 
itself  stem   from  the  Afroantarctic  triangle  (pp.   364,   365). 

Much  could  be  said  to  this,  but  I  shall  confine  myself  to  remarking  that  the 
claims  of  Rhipsalis  outside  America  to  true  citizenship  are  still  under  suspicion  and 
that  the  existence  of  this  specialized  type  of  angiosperms  in  the  Gondwana  flora 
needs  proofs  of  some  sort.  But  Croizat  does  not  stop  here.  On  p.  523  we  read: 

Considering  that  the  New  World  is  uniformly  at  the  receiving  end  of  angiospermous 
tracks  throughout  the  first  epoch  of  migration  we  are  drawn  to  conclude  that  none  of 
the  primary  angiosperms  ever  originated  in  the  Americas. 

But  who  will  tell  us  where  the  first  angiosperms  were  evolved?  We  have  to 
go  back  to  times  when  "the  Americas",  as  we  know  them,  did  not  exist  and  where, 
just  as  everywhere  else,  all  centres  of  origin  are  secondary.  I  cannot  see  that  his 
opinion  that  the  primary  centre  must  be  looked  for  in  an  "Afroantarctic  triangle" 
rests  on  a  stable  foundation,  but  everybody  will  agree  when  he  formulates  the 
following  recommendation  to  the  "phytogeographers  of  the  Academic  school": 
"Look,  and  keep  silent  awhile  ere  you  speak." 

Phrygilanthus  and  Santalum\  compare  what  will  be  said  below  about  Coprosma. 

Dysopsis.  Antarctic  according  to  Croizat  p.  51;  I  regarded  it  as  neotropical. 

"A  large  part  of  the  Ericaceae  originated  with  the  Empetraceae  in  the  Afro- 
antarctic Triangle",  Croizat  writes  p.  381.  That  the  Anlarctic  has  been  used  as 
a  migration  route  of  the  Gaultherioideae  seems  certain,  let  alone  where  the  family 
or  complex  of  families  had  its  beginning.  Croizat  finds  that  the  occurrence  of  a 
species  of  Pernettya  in  Juan  Fernandez  and  of  another  in  Galapagos  is 

one  of  the  most  interesting  aspects  of  the  Gaultherioideae.  ...  a  track  of  this  nature  is 
nothing  unusual,  of  course,  because  dispersals  of  this  extent  and  nature  are  commonplace 
between  the  vicinity  of  Juan  Fernandez  and  Hawaii,  not  to  speak  of  the  Galapagos  and 
the  Revilla  Gigedos  (pp.  167,  168). 

We  have  examples  of  a  floristic  "contact"  between  Juan  Fernandez  and  Hawaii, 
but  they  are  very  few  and  to  call  this  type  of  dispersal  commonplace  is  an  inadmis- 
sible exaggeration  of  facts.  The  American  area  oi  Penietiya  extends  from  Fuegia- 
Falkland  to  Mexico,  and  its  presence  on  Juan  Fernandez  and  Galapagos,  outlying 
stations  west  of  the  continent,  is  not  surprising;  we  need  not  construct  a  special  east 
Pacific  track  to  explain  them,  nor  assume  that  "the  South  Pacific  is  a  fundamental 
center  of  the  Ericaceae". 

I  have  mentioned  Empetrum  and  the  boreal  centre  of  Empetraceae,  also  that 
Croizat  wants  them  to  have  followed  the  same  stream  as  Ericaceae:  "they  begin 
their  visible  dispersal  only  westward  from  the  Cape  region"  (p.  349),  "with  the 
present  Atlantic  as  the  axis  of  their  distribution"  (p.  353).  Empetrum  used  to  pass 
as  a  classical  example  of  bipolarism,  but  to  those  who  still  believe  that  bipolarity 
exists  Croizot  says:  "bipolarism,  as  we  know,  is  a  bugaboo  of  academic  phyto- 
geography" — I  am  afraid  that  some  of  us  didn't  know  that. 

The  case  of  Cuminia  (for  which  Croizat  uses  the  fancy  name  Johowia),  whether 
or  not  to  be  linked  with  the  Old  World  and  Hawaiian  Prasioideae,  is  embarrassing: 


278  C.   SKOTTSBERG 

my  discussion  with  F.  H.  H.  l^Rowx  [jji]  made  Croizat  remark  that  "the  debate 
proved  futile,  as  could  be  anticipated".  To  attempt  a  solution  and  to  discuss  possible 
routes  of  mi<i;ration   is  called   "obvious  nonsense". 

Of  the  presumable  history  of  liupJirasia  enough  has  been  said  above.  In  Croi- 
ZAl's  opinion  there  cannot  be  the  cjuestion  of  a  north-south  track  from  Asia  across 
Malaysia  to  Australia  and  New  Zealand;  his  map  fig.  6  shows  the  centre  far  south 
in  the  Pacific,  from  where  tracks  lead  east  to  S.  America  and  west  to  New  Zea- 
land and  Australia  and  thence  from  there  to  Eurasia  and  along  the  Aleutian  arc  to 
N.   America. 

I'rom  the  lu'pothetical  "main  angiosi)ermous  center"  south  of  Madagascar, 
where  Plantao^o-Pa/afopsyl/iii/fi  was  born,  one  branch  leads  west  around  S.  America 
to  Juan  Fernandez,  another  east  to  Polynesia  and  from  there  to  Hawaii,  alterna- 
tively reached  directly  from  Juan  P^rnandez.  I  preferred  the  first  alternative,  but 
even  if  the  circumpolar  distribution  suggests  an  Antarctic  origin  I  can  see  no  good 
reason  to  place  the  starting  point  south  of  Madagascar. 

Copros))ia  is  one  of  the  standard  Antarctic  dispersal  types  but  lacking  in  Africa 
and  in  continental  America.  P>om  its  distribution  Croizat  draws  the  following 
conclusions. 

This  dispersal  shows  a)  An  Antarctic  range  in  the  southern  Pacific  which  fed  Co- 
prosma  to  all  lands  between  Juan  Fernandez  and  E.  Java,  b)  This  range  could  possibly 
reach  Hawaii  from  West  along  the  line  East  Malaysia-Hawaii,  c)  The  trend  of  this  type 
of  disj)ersal  is  all  westward  from  the  Americas  (Juan  Fernandez  to  New  Zealand,  Aus- 
tralia, New  (iuinea,  Malaysia)  ultimately  veering  again  eastward  to  reach  Hawaii  (p.  101). 

To  make  Juan  P^ernandez  the  source  of  Coprosnia  in  New  Zealand  is  utterly  im- 
possible and  the  statement  is  not  in  agreement  with  Croizat's  map,  fig.  29,  which 
shows  a  baseline  along  lat.  60°  between  the  longitudes  of  New  Zealand  and  Fuegia 
and  arrows  pointing  west  to  New  Zealand  and  northeast  to  Juan  F'ernandez. 

1  am  not  going  to  enter  upon  a  discussion  of  the  origin  of  a  world-wide  family 
like  Com{)ositae;  I  shall  only  quote  the  following  sentence:  "The  perfect  circum- 
[)olar  distribution  of  I'araxacuui  viagellanicuin  ...  in  itself  tells  us  where  is  to  be 
sought  the  ultimate  origin  of  the  Compositae"  (p.  63).  Even  if  it  is  true  that  this 
interesting  dandelion  also  inhabits  New  Zealand,  the  conclusions  would  seem  too 
far-reaching. 

I  shall  conclude  this  discussion  with  a  summary  of  a  recent  paper  by  AXELROD 
(/■r)  who  gives  us  a  palaeontologist's  view  on   angiosperm  evolution. 

The  cradle  of  angiosperms  stood  in  the  tro})ical  belt,  the  width  of  which  has 
undergone  considerable  change  during  the  geological  ages.  The  angiosperms  origi- 
nated and  evolved  in  uplands  during  Permian  to  Triassic  times  and  gradually 
descended  to  the  lowlands  to  replace  the  vanishing  floras  now  extinct  or  repre- 
sented by  such  remnants  as  the  Cycadophyta.  P\)ssil  records,  especially  of  pollen, 
make  it  probable  that  many  families  existed  in  early  Jurassic,  and  from  middle 
Cretaceous,  when  angiosperms  had  gained  dominance  over  gymnosperms  and  ferns, 
the  presence  of  numerous  families  belonging  to  very  different  orders  bear  witness 
of  a  long  history.   By  that  time  three  different  lowland  floras  had  developed,  the 


DERIVATION  OF  THE  FLORA  AND  FAUNA  279 

Tropic-,  Arcto-,  and  Antarcto-Cretaceous,  and  it  is  the  latter  that  interests  us  here. 
AXELROD  emphasizes  that  numerous  data  allow  us  to  conclude  that  many  families 
and  genera  generally  regarded  as  having  evolved  in  the  temperate  zones  originated 
in  the  tropics. 

The  data  seem  to  support  the  view  that  angiosperms  have  not  had  an  exclusively 
holarctic  source,  or  a  wholly  austral  center  of  origin.  They  were  being  assembled  in 
both  regions  by  the  gradual  adaptation  of  basic  tropical  groups  to  extratropical  conditions 
during  the  long  period  of  Permo-Triassic  down  to  the  Cretaceous.  On  this  basis  the 
Arcto-Cretaceous  and  Antarcto-Cretaceous  floras  of  higher  temperate  latitude  represent 
vegetation  types  whose  genera  were  derived  largely  by  the  long  and  continued  differen- 
tiation of  successively  derivative  members  of  original  tropical  and  border-tropical  angio- 
sperms. From  this  standpoint,  the  temperate  regions  to  the  north  (holarctic)  and  south 
(antarctic)  are  subordinate  or  secondary  centers  in  early  angiosperm  evolution. 

The  conclusions  we  can  draw  from  this  are  (i)  that  the  origin  and  primary  seat 
of  the  angiosperms  was  in  the  tropics  before  anything  like  the  present  map  existed; 
(2)  that  this  tropical  flora  gradually  gave  birth  to  a  temperate  flora  also  in  the 
south;  (3)  that  Antarctica  became  the  centre  of  a  varied  Antarcto-tertiary  flora 
which  spread  north  and  over  S.  America  reached  the  region  where  now  stands 
Juan  Fernandez. 

II.  Pteridophyta. 

Owing  to  their  great  age  and  perhaps  also  to  their  greater  faculty  of  dispersal 
many  of  the  genera  and  some  species  have  attained  a  very  extensive  range  making 
it  difficult  or  impossible  to  assign  them  to  one  of  the  elements  proposed  in  the  sub- 
division of  the  angiosperms.  Of  23  genera  found  in  Juan  Fernandez,  16  are  very 
wide-spread;  some  of  them  belong  to  the  largest  fern  genera,  and  even  if  we  avail 
ourselves  of  the  generic  concept  used  by  Copeland  (dp),  the  situation  remains 
practically  the  same.  To  Copeland  50-75  %  of  the  living  ferns  are  of  austral  or 
Antarctic  ancestry ;  we  shall  see  to  what  extent  this  is  true  of  the  Juan  Fernandez  flora. 

With  regard  to  the  Hymenophyllaceae  COPELAND  remarks  (dy.  174)  that  "no 
other  plant  family  of  its  size  and  diversity  is  quite  so  conspicuously  Antarctic  in 
origin  as  this  one". 

Trichommies-VandenboscJiia  is  pantropical;  as  2  of  the  3  insular  species  are 
decidedly  neotropical  in  their  affinities,  also  T.  phzlippianum  is  referred  to  the  same 
element. 

Serpyllopsis  is  subantarctic-American  and  best  attached  to  an  Antarcto-tertiary 
element. 

Hymenoglossum  should  perhaps  be  referred  to  the  same  element,  but  its  re- 
lations are  just  as  unknown  as  those  of  Serpyllopsis. 

Hymeiiophyllum-Mecodium  is  a  pantropical  group,  also  represented  in  New  Zea- 
land; cuneatum  and  caudiculatum  are  most  likely  of  neotropical  origin  and  this  may 
be  true  also  o{fuciforme.  H.  pectinatum  stands,  COPELANI)  says,  apart  from  the  other 
species,  but  I  can  find  no  good  reason  to  call  in  Antarctic,  while  the  distribution 
of  Sphaerocionium  ferrugineuni  and  its  near  relatives  testifies  to  their  Antarctic  origin 
in  spite  of  their  affinity  to  ciliatiim  (comp.   above  p.   219).  Hymenophylluni  ^.  str. 


28o  C.  SKOTTSBERG 

as  limited  by  Coi'Ki.AM)  1938  (66)  does  not  \x\q\w<\^  falklandicum  [Mecodium  accord- 
ing to  CcM'KLAXD  p.  94),  though  it  is  closely  akin  to  peitatiim,  and  H.  riigosiim 
"is  to  //.  tuuhridi::€US€  what  H .  falklandiciim  is  to  H.  peltatuin'  (62.  13).  According 
to  CliRisr  (-r^*.  146),  pi'ltattou  and  tiinhridgense  are  essentially  Antarctic-andine  but 
found  north  as  far  as  Mexico,  also  reported  from  S.  Africa  and  New  Zealand,  ex- 
tending in  insular  climates  to  western  L2urope.  The  conclusion  would  be  that  also 
falkUvidicuni  and  rNgosiiffi  should   be  brought  to  the  Antarctic  group. 

IlymoiopJivlluni-Mey'nigiuDi  secundum,  plicatiiui  and  tortuosum  are  subantarctic- 
American  in  their  present  distribution;  JMcrijigiuni  is  austral  and  extends  north  to 
tropical  Asia,   and   its  origin   most  likely  Antarctic. 

'Iliyrsoptcris.  Mesozoic  fossils  supposed  to  come  near  this  genus  have  been  re- 
ported from  the  boreal  zone;  besides,  from  the  Tertiary  in  Chile  (see  below  p.  399). 
llll.l.  [1J4.  1477)  calls  it  "a  remnant  of  a  once  widespread  group  which  migrated 
southwards,  possibly  in  Cretaceous  times,  from  the  Northern  Hemisphere".  With 
regard  to  its  present  unique  station  COPELANI)  [6y .  175)  writes: 

As  juan  l-'ernandez  can  hardly  have  evolved  so  peculiar  a  fern,  its  most  reasonable 
origin,  as  a  juan  Fernandez  fern,  is  Antarctica,  whether  directly  or  through  southern 
Chile.  Kuropean  fossils  have  been  referred  to  this  genus,  but  Juan  Fernandez  was  not 
colonized   from   FAirope. 

I.op/iosor/a  is  exclusively  neotropical  at  present,  but  if  we  are  to  believe,  as 
C<)i'i:i,AM)  thinks,  that  the  Cyatheaceae  derive  from  Antarctica,  it  is  unlikely  that 
Lopliosaria  would  be  an  exception. 

The  Antarctic  origin  of  Dicksonia  seems  very  clear  to  judge  from  its  actual 
distribution  pattern.  To  what  was  said  above  p.  222  I  shall  add  that  along  the 
track  New  Zealand-Australia-Xew  Caledonia-Fiji (-Samoa)-Malaysia  14  species  are 
found,  along  the  Andean  path  9,  and  that  the  species  of  Juan  F^ernandez  are  not 
linked  with   the  neotropical  but  with  southwest  Pacific  species. 

'Fhc    present   distribution   of  the  genus   Cystopteris  suggests  a  boreal  origin. 

In  its  wide  as  well  as  in  its  more  restricted  sense  Dryoptcris  is  world-wide, 
and  C()i'i;i  and's  remark  (^7.  181)  that  he  has  little  doubt  that  the  group  as  a 
whole  is  of  southern  origin  (see  also  6g.  122)  is  perhaps  little  more  than  a  guess. 
I  think  we  can  take  it  ior  granted  that  hundreds  of  species  have  evolved  in  the 
neotropical  region,  where  the  single  Juan  FY-rnandez  species  D.  inaequalij'olia  has 
its   relatives. 

rolystiiliuui .  Mistern  .Asia  is  the  centre  of  greatest  concentration  ;  the  section 
occurring  there  is,  in  C(  )ii;i,AM)'s  oj)inion,  the  least  primitive,  and  we  have  a  number 
ot  southern  hi-  or  tricentric  frjrms,  among  them  P.  vcsiiiuvi  coll.  The  endemic  P. 
bertiricDiuui  is  very  close  to  the  circumj)()lar  adicDitiforme.  To  quote  CoPKLAND:  "The 
case  for  ;ui  Antarctic  origin  of  Polysticlnou  is  so  clear  that  the  evidence  has  long 
been  familiar  even  t(j  those  hesitant  to  draw  the  obvious  conclusion"  [dj.  1 81); 
and,  "Tile  distribution  .  .  .   testifies  clearly  to   its  Antarctic  origin"   [6g.  109). 

Aythroptcris  is  the  only  fern  genus  found  in  Juan  Fernandez  which  is  absent 
from  continental  America,  but  otherwi.se  typically  austral.  Its  Antarctic  origin  is 


DERIVATION  OF  THE  FLORA  AND  FAUNA  28l 

beyond  doubt,  and  A.  altescajidens  has  its  nearest  allies  in  Oceania;  comp.  above 

p.    222. 

Aspleniwn.  CoPELAND  points  out  that  some  groups  at  least  are  southern  and 
that  perhaps  the  same  applies  to  the  whole  genus:  "it  migrated  in  its  present 
form  from  Antarctica"  {6g.  167).  Of  the  species  found  in  Juan  Fernandez,  obliquum 
is  austral-bicentric,  stellaimn  close  to  neotropical  species,  macrosorum  most  likely 
of  neotropical  affinity;  day^eoides  comes  very  near  alvarezerise  from  Tristan  da 
Cunha    and    is    not,    as    has  been  suggested,  related  to  the  boreal  ruta  muraria. 

Blechnum.  According  to  Copeland  (^9.157)  no  fern  genus  is  more  con- 
spicuously austral  in  its  present  distribution  and  thus  more  evidently  Antarctic 
in  origin.  Of  the  6  species  found  in  Juan  Fernandez  auriculatmn ,  chilense  and  cy- 
cadifolium  belong  to  an  Afro-american  group,  Schottii  and  perhaps  also  valdiviefise 
of  austral-circumpolar  type,  and  longicauda  intimately  related  to  the  neotropical 
Spriicei\  even  so  it  cannot  blur  the  picture  of  an  old  Antarctic  genus. 

Pellaea  is  an  austral-tricentric  genus  extending  north  to  Canada,  but  P.  chi- 
lensis,  endemic  in  Juan  Fernandez,  but  dangerously  close  to  a  widespread  American 
species,  belongs  to  a  group  that,  according  to  Copeland  [6^.  70),  is  best  allowed 
to  form  a  separate  genus;  I  have  brought  it  to  the  neotropical  element. 

Hypolepis  is  pantropical,  especially  neotropical,  but  numerous  species  are 
scattered  in  the  equatorial  and  austral  zones:  Africa  with  neighbouring  islands, 
Malaysia  and  north  to  Japan,  Australia-New  Zealand-Lord  Howe-Melanesia-Hawaii 
— when  Copeland  said  "without  surviving  Chilean  representatives"  he  'iox^otrugo- 
sula.  This  species  is  bicentric  or,  including  Polypodium  villosoviscidum  of  Tristan, 
tricentric;  there  is  another  variety  on  St.  Helena.  Thus  when  CoPELAND  6y.  lyy 
calls  Hypolepis  "an  old  antarctic  genus  now  at  home  chiefly  in  the  Tropics  but 
with  two  of  its  paths  still  occupied",  we  can  add  that  also  the  third  path  is  indicated. 

Adimituin  is  most  numerous  in  S.  America,  and  A.  chilense  and  related  species 
tell  us  nothing  of  the  earlier  history  of  the  genus;  CoPELAND  6g.  78:  "I  suppose 
that  at  least  the  most  of  the  extant  species  are  of  Antarctic  ancestry,  but  the 
genus  is  old  and  may  have  lived  elsewhere  without  interruption";  in  dy.  178  he 
says  that  "Antarctica  has  played  a  major  part  in  its  history",  but  that  the  genus 
"may  include  elements  surviving  from  other  floras  and  from  an  age  prior  to  the 
great  dispersal  from  the  south".   I  refer  A.  ckilefise  to  the  neotropical  group. 

Pteris.  The  great  wealth  of  species  is  in  the  tropics  and  only  few  extend  as 
far  south  as  to  south  Chile,  S.  Africa  and  New  Zealand.  P.  chileiisis  is  of  neo- 
tropical character,  perhaps  also  P.  semiadnata,  while  berteroana  belongs  to  a  group 
regarded  by  CoPELAND  as  Antarctic  to  judge  from  its  distribution  pattern  (see 
above  p.  223). 

Histiopteris  incisa  is  a  pantropical  and  circum-austral,  polymorphous  fern  with 
a  number  of  "local  derived  species"  (Copeland)  in  Indonesia  and  Polynesia.  The 
genus  is,  Copeland  says  [6g.6o),  "evidently  old  enough  to  be  a  migrant  from 
Antarctica". 

Polypodium  is  an  aggregation  of  unities  of  different  origin  and  history.  Gram- 
mitis  is,  in  Copeland's  opinion  (<57.  184),  "a  plain  Antarctic  case",  and  G.  Bil- 
lardieri  "surrounds  Antarctica  more  closely  and  completely  than  does  any  other 


282  C.   SKOTTSBERG 

fern  '  [dg.  21 1;  see  above  p.  223);  it  was  at  that  time  supposed  to  be  circum- 
polar,  but  is  replaced  in  S.  America  by  /\  DiagclUmicum.  As  mentioned  above, 
I  believe  that  if  a  t>enus  Syiiaimiiia  with  PolypodiiDii  Feuillei  2,s  the  typical  species 
is  acce{)ted,  the  endemic  P.  i>iier)iiedin)n  finds  its  place  with  this,  but  I  fail  to  see 
why  Coi'KLANi)  y6S)  regards  .V.  Ju'uil/ii  di^  "clearly  Antarctic",  whereas  the  nearly 
relatetl  CofiiophUhiuDi  is  called  "a  northern  genus".  Of  the  remaining  species 
P.  Masafiicrae  and  P.  (Xij)ho|)teris)  trichouajioidcs  are  neotropical  and  P.  (Pleo- 
peltis)  lajiceolatio)!  pantropical  but  not  reported  from  Australia  and  not  indicating 
an   Antarctic  origin. 

We  have  no  good  reason  to  look  for  an  Antarctic  ancestry  of  Elaphoglossmn, 
though  C'()ii;i,.\M)  thinks  that  it  "may  have  come  from  the  south"  {dj.  185),  and 
P.   Li)idi)iii  is  a  neotropical  species. 

The  species  of  Gleichoiia  occurring  in  Juan  Fernandez  belong  to  Sticherus, 
a  genus  segregated  by  some  modern  authors;  11  species  are  scattered  over  the 
austral  zone,  and  in  Cotkland's  opinion  Gleicheniaceae  are  "obviously  and  en- 
tirely of  Antarctic  ancestry"  (<5^.  26),  or  "entirely  Antarctic  at  some  stage  of  its 
histor}- '  [dy.  173);  how  this  should  be  explained  I  cannot  tell  unless  he  means 
that  the  family  characters  first  evolved  in  the  Antarctic,  secondary  centres  of 
evolution  having  become  established  in  the  tropics.  The  genetic  relations  between 
the  species  inhabiting  the  three  sectors  (see  above  p.  224)  should  be  studied. 
Meanwhile  I  shall  refer  the  5  species  occurring  in  southern  S.  America  to  the 
Antarcto-tertiary  element. 

The  Ophioglossaceac  are  an  ancient  family,  "scattered  with  remarkable  uni- 
formity over  the  habitable  globe"  (CoPELANl)  6g.  12).  In  dy.  167  he  paid  special 
attention  to  Hotrychiion  australe^.  Br.  which  has  the  peculiar  distribution  Argentina, 
Australia,  Tasmania  and  New  Zealand  and  "may  be  regarded  as  an  obvious  immi- 
grant from  the  south".  It  is  surprising  that  the  species  found  in  Patagonia  and 
I'\iegia  is  not  this,  but  the  northern  B.  lunaria  L.  We  have  no  reason  to  regard 
OphioglossiDn  fcy}ia)idi-zia)ium  as  coming  from  the  south. 

Lycopodiion  was  not  discussed  by  CoPELANi).  L.  magellaiiiaiin  is  subantarctic- 
circumpolar,  and  part  of  the  history  of  the  genus  may  have  been  enacted  in  Ant- 
arctica, the  more  so  as  L.  scariosuni^  which  belongs  to  another  section,  is  bicentric. 

With  reference  to  the  discussion  above  the  Pteridophytes  are  arranged  as 
follows. 

I.   Antarcto-tertiary  element. — 32  sp.  (60.4%). 

1.  Distribution  pattern  austral-circumpolar,  bicentric  or  tricentric. — 20  sp. 
a.  ICndemic  sj)ecies  (2):  Hymenophyllum  rugosum,  Polystichum  berterianum. 
h.    .Species    also    found    in    S.  America    (Chile),  many  of  them  with  a  wider 

distribution  (w)  in  the  S.  hemisphere  (18):  Hymenophyllum  secundum,  plicatum, 
tortuosum,  ferrugineum  and  falklandicum  (w),  Polystichum  vestitum  (w),  Asplenium 
obliquum  (w)  and  dareoides,  Hlechnum  valdiviense  and  chilense,  Hypolepis  rugosula 
(w),  Ilistiopteris  incisa  (w),  Polypodium  magellanicum  (w),  Gleichenia  pedalis, 
quadripartita    and  cf.  litoralis,   Lycopodium  magellanicum  (w)  and  scariosum  (w). 

2.  S[)ecies  belonging  to  genera  endemic  in   S.  America  but  supposed  to  be 


DERIVATION  OF  THE  FLORA  AND  FAUNA  283 

of  Antarctic  origin:  Serpyllopsis  caespitosa,  Hymenoglossum  cruentum(?),  Lopho- 
-soria  quadripinnata. — 3  sp. 

3.  Endemic  species  without  near  relatives  in  America,  suggesting  either  an 
ancient  southern  Pacific  west-east  path  or  an  Antarctic-Magellanian  track  now 
not  occupied:  Dicksonia  berteroana  and  externa,  Arthropteris  altescandens,  Blech- 
num  Schottii,  Pteris  berteroana. — 5  sp. 

4.  Species  closely  related  to  S.  African  ones. — 2  sp. 

a.  Endemic:  Blechnum  cycadifolium. 

b.  Also  on  the  mainland:  Blechnum  auriculatum. 

5.  Endemic  species  very  nearly  related  to  a  neotropical  species:  Blechnum 
longicauda. —  i   sp. 

6.  Endemic  genus  without  affinities  to  living  genera:  Thyrsopteris  elegans. 
—  I   sp. 

II.  Neotropical- Andean  element. — 20  sp.  (37.7  %). 

a.  Endemic  species  (8):  Trichomanes  Ingae  and  philippianum,  Dryopteris 
inaequalifolia,  Asplenium  stellatum  and  macrosorum,  Pellaea  chilensis,  Polypodium 
intermedium,  Ophioglossum  fernandezianum. 

b.  Also  on  the  mainland,  restricted  to  Chile  or  more  wide-ranging  (10):  Tricho- 
manes exsectum,  Hymenophyllum  cuneatum,  caudiculatum,  fuciforme  and  pecti- 
natum,  Adiantum  chilense,  Pteris  chilensis  and  semiadnata,  Polypodium  Masafuerae 
and  lanceolatum. 

c.  Non-endemic  neotropical  species  not  found  in  Chile  (2):  Polypodium  tricho- 
inanoides,  Elaphoglossum  Lindenii. 


III.  Arcto-teftiary  element. —  i  sp.  (1.9  %). 
Also  in  Chile:   Cystopteris  fragilis  var. 


The  Antarctic  element  is  considerably  larger  than  in  the  angiosperms,  60.4  % 
against  42.2,  the  neotropical  being  of  the  same  size,  37.7  and  36.7%.  On  the  other 
hand,  the  boreal  element  is  very  insignificant  (even  open  to  doubt).  No  living 
pteridologist  has  a  wider  general  knowledge  of  the  ferns  than  E.  B.  Copeland,  and 
«ven  if  his  new  system  does  not  appeal  to  everybody,  we  are  bound  to  pay  attention 
to  his  theories.  In  several  of  his  writing  he  points,  as  we  have  seen,  to  Antarctica 
as  the  main  source.   On  the  other  hand  he  admits  that 

ferns  existed  well  over  the  world  longer  ago  than  I  have  would  try  to  explain  their 
presence  as  immigrants  from  Antarctica,  and  must  be  supposed  to  have  maintained 
■continuous  existence  elsewhere.  This  being  so,  the  preponderance  of  ferns  of  apparent 
Antarctic  origin  in  the  world  to-day  is  surprising  indeed  ((5/.  188). 

This  origin  is  not  readily  revealed  in  the  large  and  world-wide  families  and 
genera;  the  best  proofs  are  furnished  by  small  families  restricted  to  high  southern 
latitudes  and  bi-  or  tricentric  in  distribution — most  of  these  families  have  been 
created  as  a  result  of  the  modern  splitting  process.  With  a  single  southern  genus 
in  a  large  tropical  family  we  cannot  feel  on  safe  ground,  and  if  in  a  large  tropi- 
cal   genus    only    a    solitary   species    is   southern,   we   are  not  inclined  to  classify 


284  C.  SKOTTSBERG 

the  genus  as  Antarctic  in  origin.  But,  CorELAND  remarks,  if  the  ferns  as  a  group 
were  of  tropical  origin  the\-  would  be  expected  to  be  far  more  abundant  north- 
ward, where  the  land  areas  increase,  than  southward  where  the  land  decreases 
— and  very  much  so  —  in  size,  and  this  would  be  still  more  so  if  they  were  of 
northern  origin.  The  conclusion  is  that  if  a  family  or  genus  is  mainly  southern 
to  da\',  this  fact  is  a  strong  indication  of  its  Antarctic  origin  and  that,  if  the  pre- 
sent range  is  wholl}'  southern,  the  evidence  becomes  almost  conclusive.  If  the 
more  primitive  families  or  genera  are  found  to  be  characteristically  southern,  a 
southern  seat  of  old  fern  evolution  is  almost  demonstrated  [68.  626).  But  when  it 
comes    to    the  large  families,  their  history  may  be  much   more  difficult  to  read: 

A  large  family,  even  if  of  southern  origin,  cannot  possibly  be  predominantly  southern 
in  i)resent  distribution,  because  tropical  and  northern  species  must  outnumber  the  whole 
tlora   of  Antarctic   America,   or  even   of  New   Zealand   {6y .  158). 

With  this  he  wants  so  sa}%  I  suppose,  that  a  vigorous  family  has  invaded  the 
tr()j)ics  and  also  extended  north  under  rapid  evolution  of  genera  and  species  con- 
cealing the  j)rimary  origin;  he  believes  this  evolution  to  have  been  so  rapid  that 
the  j)eriod  since  the  Miocene  has  been  sufficient  to  create  most  of  the  existing 
species  and  a  large  part  of  the  genera  and  for  their  spread  over  any  expanse  ot 
suitable  land  area  (I.e.). 

We  have  no  evidence  that  the  Tropics  as  a  whole  were  at  any  past  time  unfit  for  ferns; 
and  the  assumi)tion  that  this  has  been  the  chief  place  of  evolution  obviates  the  neces- 
sity of  assuming  and  explaining  migration  in  latitude.  It  is  only  w-hen  we  open  our 
eyes  to  anomalies  in  present  distribution  that  appeal  to  other  places  of  evolution  be- 
( omcs   necessary   [6y.  163). 

(ireat  geographical  disjunctions,  of  which  the  Juan  Fernandez  fern  flora  offers 
man\-  examples,  are  such  anomalies,  and  they  seem  to  show  that  Antarctica  has 
taken   a   very   important  part  in   fern  history. 

III.  Musci. 

In  his  analysis  of  the  Alagellanian  moss  flora  Cardot  (48)  distinguished  an 
Antarctic  element  and  he  believed  in  a  common  origin  of  the  floras  of  Magella- 
nia  and  New  Zealand  (j).  44).  Hkkzoc;  (i2g)  found  that  Juan  Fernandez  has  "eine 
last  \-ollstandig  austral  antarktische  Moosvegetation",  but  also  possesses  "ein  paar 
tropisch  annuitende  Artcn:  ThysaiioDiiiriuni  RicJiardi,  Porothanniium  fasciculaiuvi, 
Pi)niatilla  luaooslicta  und  ein  kliacopiliDiiy  The  monotypical  Chilean  genus  Lam- 
l^yof'hylluDi  was  also  mentioned  as  a  genus  of  tropical  ancestry.  A  circumpolar 
austral-antarctic  element  is  said  to  dominate  in  the  moss  flora  of  Patagonia  and 
l'\iegia;  characteristic  genera,  also  re[)resented  in  Juan  Fernandez,  are  Dkraiioloma, 
(  lota,  Dindyocryf^hacd,  I .cpyycuhi}!,  rtycliouinium,  Weyniouthia,  DisticJiophylluni^ 
r/in\^-(>/^/iyI/in//,  liriof^us,  J f]f(phn\i^ju»i,  Sciaro))iiiu)i,  perhaps  also  CatagoJiiopsis 
and  Psihpilii))!.  Ii<ms(  [ikr  (//,')  gives  many  examples  of  austral-bicentric  taxa: 
Lcpyrodo)itac(ai\  /\>Iy/r/i/i/i</i//^//!(s^  Wcymouthia,  PtcrygopJiylluni,  Sciaromiuin  sect. 
Aloma,  Hyp(ptn-yo;iu))i  sect.  Stoiobasis,  I lypnodcndron,  and  species  of  Disticko- 
phylluui^    J  ovinia,   Macyoviityiuui   and  MielichJioferia. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  285 

The  synopsis  pp.  226 — 233  led  to  a  classification  according  to  the  actual  geo- 
graphic distribution.  We  have  to  look  for  austral-antarctic  species  and  genera 
in  the  Andine-Chilean,  Subantarctic-Magellanian  and  Pacific  groups;  thereby  I 
have  tried  to  follow  the  same  principles  as  applied  before.  Many  of  the  endemic 
species  are  difficult  to  place  because  their  author  did  not  compare  them  with  other 
species  or  discuss  their  position;  1 1,  belonging  to  large  and  widespread  genera  were 
left  out  (see  list  p.  235).  The  8  species  of  Tham7imm,  all  endemic,  were  tentatively 
included  in  group  IV.  In  Natiirl.  Pflanzenfam.  2d  ed.  Brotherus  enumerates  34 
species,  but  none  at  all  are  quoted  for  S.  America.  Sect.  I  is  entirely  austral, 
with  a  single  species  in  tropical  Asia  (Sumatra,  Java,  Borneo,  Philippines);  2  are 
found  in  Australia  and  New  Zealand,  i  in  New  Caledonia  and  8  in  Juan  Fernandez. 
Of  sect.  II,  3  species  are  southern  (New  Hebrides,  E.  Australia-New  Zealand,  New 
Zealand-Java-Sumatra),  the  remainder  scattered  over  the  boreal  zone,  3  N. 
America,  3  mediterr.-macaronesian,  i  England,  i  W.  Europe  to  Japan,  2  Cauca- 
sus, 9  E.  Asia. 

The  total  number  of  species  included  in  the  following  synopsis  is   120. 

I.  Antarcto-tertiary  element. — 67  sp.  (55.8  %). 

1.  Bicentric  or  tricentric  (t)  distribution:  Ditrichum  affine,  Amphidium  cyathi- 
carpum,  Dicranoloma  Billardieri  (t)  and  Menziesii,  Campylopus  introflexus  (t), 
Thysanomitrium  leptodus,  Fissidens  rigidulus  and  asplenioides  (t),  Rhacomitrium 
symphyodontum  (t),  Zygodon  intermedins  and  Menziesii,  Rhizogonium  Novae  Hollan- 
diae  and  mnioides,  Bartramia  patens  (t),  Philonotis  scabrifolia  (t),  Rhacocarpus  Hum- 
boldtii  (t),  Weymouthia  mollis,  Leptodon  Smithii  (t),  Pterygophyllum  obscurum 
and  denticulatum,  Lopidium  concinnum,  Rhynchostegium  tenuifolium,  Polytricha- 
delphus  magellanicus,  Dendroligotrichum  dendroides. — 24  sp. 

2.  Endemic  or  also  found  in  Chile  and  restricted  to  the  S.  American  sector, 
in  either  case  with  Australian-Neozelandic  relations,  with  the  exception  of  Eusti- 
chia. — 30  sp. 

a.  Endemic  species  (9):  Dicranoloma  fernandezianum,  capillifolioides  and  nigri- 
caule,  Ptychomnium  falcatulum,  Distichophyllum  fernandezianum,  Pterygophyllum 
tenuinerve,  Eriopus  leptoloma  and  grandiretis,  Thuidium  Masafuerae. 

b.  Also  in  Chile  (21):  Ditrichum  longisetum,  Dicranoloma  capillifolium,  Eu- 
camptodon  perichaetialis,  Rhacomitrium  subnigritum  and  striatipilum,  Macromitrium 
hymenostomum,  Leptostomum  Menziesii,  Eustichia  Poeppigii  (other  species  in 
America  and  Africa),  Dendrocryphaea  cuspidata,  Lepyrodon  parvulus,  tomentosus 
and  implexus,  Ptychomnium  subaciculare  and  ptychocarpum,  Pterygophyllum  ano- 
malum,  Hypopterygium  Thouini,  Thuidium  Valdiviae,  Sciaromium  pachyloma, 
Catagoniopsis  berteroana,  Hypnodendron  microstictum,  Psilopilum  antarcticum. 

3.  Endemic  species  of  presumably  Australasian  and  Polynesian  affinity,  not 
or  more  distantly  related  to  American  species:  Macromitrium  fernandezianum  and 
Masafuerae,  Cyptodon  crassinervis,  Thamnium  rigidum,  latinerve,  Caroli,  Ingae, 
crassinervium,  proboscideum,  assimile  and  confertum,  Distichophyllum  subelimba- 
tum  and  assimile. — 13  sp. 


286  C.   SKOTTSBERG 

II.  Neotropical  element.  —  26  sp.  (21.7  %). 

a.  ICndemic  species  (11):  Campylopus  subareodictyon,  Leptodontium  fernan- 
dezianum,  Didymodon  calymperidictyon  and  linearis,  Bryum  fernandezianum,  Pin- 
natclla  niacrosticta,  Isopterygium  fernandezianum,  Rhaphidostegium  Masafuerae 
and  caespitosoides,   Rigodiuni  robustum  and   Looseri. 

Ik  Also  in  S.  America  (15):  Gymnostomum  calcareum  (widely  dispersed),  Cam- 
pyl()l)us  areodictyon,  Thysanomitrium  Richardi,  Mielichhoferia  longiseta,  Anacolia 
siibsessilis,  Porothamnium  fasciculatiim  and  arbusculans,  Lamprophyllum  splendi- 
dissimum,  Rhacopilum  fernandezianum,  Stereodon  Lechleri,  Rhaphidostegium 
caespitosum,   Rigodium  toxarium,  arborescens,  hylocomioides  and  tamarix. 

III.  Chilean  element. —  24  sp.  (20.0  %). 

(7.  luidemic  species  (4):  Fissidens  pycnotylus,  Ulota  fernandeziana,  Philonotis 
glabrata,  Rhaphidostegium  aberrans. 

/\  Also  in  Chile,  mostly  southern  (17):  Pleuridium  Robinsonii,  Hymenosto- 
mum  kunzcanum,  Oncophorus  fuegianus,  Campylopus  truncatus,  Fissidens  lepto- 
chaete  and  maschalanthus,  Tortula  scabrinervis  and  flagellaris,  Rhacomitrium  lori- 
forme  and  convolutum,  Stenomitrium  pentastichum,  Bryum  Lechleri,  Bartramia 
aristata,  Philonotis  krauseana  and  vagans,  Rhynchostegium  complanum,  Oligotrichum 
canaliculatum. 

(-.  Cosmopolitan  (3):  Ceratodon  purpureus,  Rhacomitrium  lanuginosum,  Funa- 
ria  hygrometrica. 

Xoie. — On  p.  234  Campylopus  introfJexus  was  placed  here;  it  was  transferred  to 
I:  I  on  account  of  its  circumpolar  distribution  in  the  far  south,  from  where  it  may  have 
invaded  the  tropics  and  migrated  north,  but  it  is  perhaps  just  as  probable  that  it  is 
an   old   j)antroj)i(al   species  which   has  spread  both   north   and  south. 

IV.  Atlantic-Mediterranean  element. — 3  sp.  (2.5  %). 
Comp.  ahove,  p.  235. 

a.   I'Jidcmic:   h'issidcns  crassicuspes. 

h.   \ot  endemic:   Campylo[His  polytrichoides,  Trichostomum  brachydontium. 

W'c  have  seen  above  that  Hkrzog  regarded  the  moss  flora  of  Juan  P^ernan- 
dcz  as  almost  entirely  austral-antarctic  and  that  the  tropical  element  was  small. 
Most  likely  he  added  the  species  forming  my  Chilean  element,  because  they  are 
concentrated  in  the  moist  southern  part  of  the  country  and  in  several  cases  range 
south  into  the  subantarctic  zone  which  is,  however,  no  proof  of  their  Antarctic 
ancestry.  A  species  like  l\)r()tJiai)niiui}i  arbusculans  is,  as  far  as  I  know,  restricted 
to  Chile  and  I'atagonia,  but  the  genus  is  essentially  tropical,  Stereodon  Lechlen 
extends  from  S.  Chile  to  W.  I'atagonia,  but  is  the  only  species  reported  south 
of  the  lujuator;  Rii^odiuDi  is  American  except  2  species  found  in  Africa  and 
essentially  tropical.  On  the  other  hand  it  is  difficult  to  draw  a  limit  between 
groups  II  and  III,  but  the  s[)ecies  with  a  southern  area  are  so  numerous  that  I 
felt  obliged  to  make  this  distinction.  The  Atlantic  element  will  perhaps  disappear 
when  the  distribution  becomes  better  known. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  287 

The  bi-  or  tricentric  species  found  in  Juan  Fernandez  represent  a  minor  frac- 
tion only  of  the  mosses  common  to  S.  America  and  New  Zealand.  Sainsbury's 
new  Flora  (210)  contains  nearly  60  species  with  this  type  of  distribution,  and  to 
these  some  occurring  in  Australia  and  Tasmania,  but  not  in  New  Zealand,  may 
be  added.  Ditrichum  affine,  Trichosiomum  brachydontium,  Rhacomiirium  symphyo- 
dontum,  Bartramia  patens,  and  Pterygophyllum  denticulatum,  listed  for  New  Zealand 
by  Brotherus  (j4),  are  not  recorded  by  Sainsbury. 

Consequently,  what  Miss  FULFORD  (see  below  p.  289)  says  about  the  Hepati- 
cae  is  true  also  of  the  mosses. 

IV.  Hepaticae. 

In  an  interesting  paper  on  the  distribution  of  the  Hepaticae  UOMlN  (yd)  lays 
stress  upon  that  they  are  just  as  specialized  geographically  as  the  flowering 
plants,  and 

einen  uralten,  heiitzutage  in  den  gemassigten  iind  kalteren  Gebieten  sozusagen  erstarr- 
ten,  sich  von  ihren  ganz  speziellen  Standorten  nicht  weiter  ausbreitenden,  wenig  an- 
passungsfahigen  Typus  darstellen,  welcher  auf  der  nordlichen  Hemisphare  seinen  Ent- 
wickelungsgang  in  weit  zuriickliegenden  Epochen  durchgemacht  iind  bereits  zur  Tertiar- 
zeit  in  den  Hauptziigen  beendet  hat  (p.  3). 

He  thinks  that  the  evolution  still  continues  in  the  tropics.  Numerous  genera  are 
widespread,  but  the  majority  of  genera  and  species  inhabit  the  tropical  and  south 
temperate  rain  forests  in  America,  Africa  (perhaps  not  so  rich.?),  Malaysia,  Austral- 
asia and  Oceania.  Austral-antarctic  genera  mentioned  by  DOMIN  are  Schistochila, 
Balantiopsis,  Adelanthus  (hardly  austral),  Lophocolea,  Chiloscyphus,  Trichocolea,  and 
many  genera  with  their  greatest  wealth  of  species  in  the  tropics  are  well  repre- 
sented in  the  temperate  and  cool  south,  such  as  Riccardia,  Symphyogyjia,  Pla- 
giochila,  Madotheca,  Lepidozia,  Radula,  Frullania,  while  few,  in  cases  very  few, 
species  are  found  in  the  boreal  zone.  From  Herzog's  handbook  the  following 
genera  may  also  be  quoted  as  tropical-austral,  or  essentially  or  exclusively  south- 
ern: HymenophytuDi,  Acrobolbus,  Tylunanthus,  Jainesoniella,  Saccogyna,  Lepicolea 
and  Marsupidiuni,  to  mention  only  genera  also  found  in  Juan  Fernandez.  Lepi- 
dolaeiia  is  restricted  to  the  south  temperate  and  cold  zones.  In  the  following  ar- 
rangement I  have  been  guided  by  these  authors. 

I.  Antarcto-tertiary  element. — 84  sp.  (67.7  %). 

1.  Bicentric  or  tricentric  (t)  species:  Marchantia  berteroana  (t)  and  foliosa, 
Riccardia  insularis,  Metzgeria  decipiens  and  violacea(t),  Hymenophytum  flabellatum, 
Symphyogyna  hymenophyllum,  Monoclea  Forsteri,  Pallavicinia  xiphoides,  Jame- 
soniella  colorata  (t)  and  grandiflora  (t),  Acrobolbus  excisus,  Mylia  repens,  Lophocolea 
pallidevirens  and  muricata  (t),  Marsupidium  piliferum,  Bazzania  cerina,  Lepidozia 
sejuncta  (t)  and  plumulosa,  Lepicolea  ochroleuca  (t),  Lepidolaena  magellanica, 
Schistochila  splachnophylla,   Frullania  magellanica  (t). — 23  sp. 

2.  Restricted  to  the  American  sector,  endemic  in  Juan  Fernandez  or  also 
found  on  the  mainland,  connected  with  species  in  the  Australia-New  Zealand  sec- 
tor.— 61   sp. 


288  C.   SKOTTSBERG 

a.  luidemic  (17):  Riccardia  adghitinata  and  leptostachya,  Metzgeria  multifor- 
mis, TylunanthussilvaticLis.  bilobatiis  and  densiretis,  Lophocolea  papulosa,  angulata 
and  submuricata,  Trichocolea  opposita,  Schistochila  Skottsbergii,  Balantiopsis  hians 
and  lancifolia,  Lopholejcunca  spinosa,  Stre{)silejeunea  squarrosula  and  macroloba, 
Cololejeiinea  Skottsbergii. 

/;.  Also  found  in  Chile  (44):  Riccardia  fuegiensis,  breviramosa,  variabilis  and 
nudimitra,  Metzgeria  decrescens,  Symphyogyna  circinata  and  Hochstetteri,  Jame- 
soniella  maluina  and  oenops,  Tylunanthus  limbatus,  Mylia  fuscovirens  and  ligulata, 
Lo[)hocolea  rotundifolia,  fernandeziensis,  chilensis,  attenuata,  textilis  and  divergenti- 
ciliata,  Chiloscyphus  integrifolius  and  lobatus,  Saccogyna  squarristipula,  Adelanthus 
sphalerus,  Lepidozia  bicuspidata,  pseudozoopsis,  fernandeziensis  and  Jacquemontii, 
Trichocolea  verticillata,  Schistochila  berteroana  and  pachyla,  Balantiopsis  cancellata, 
chilensis  and  purpurata,  Radula  hastata,  microloba,  Mittenii  and  Dusenii,  Madotheca 
chilensis  and  snbsquarrosa,  FruUania  Eckloni,  chilensis,  lobulata  and  stipatiloba, 
Hrachiolejeunea  spruceana,  Strepsilejeunea  acuminata. 

II.  Neotropical  element. — 13  sp.  (10.5  %). 

a.  I'jidemic  (3):  Fossombronia  fernandeziensis,  Anthoceros  Skottsbergii,  Le- 
jeunea  reticulata. 

Ik  Also  in  tropical  South  America,  extending  to  Chile,  or  southern,  but  pre- 
sumably of  tropical  origin  (10):  Megaceros  fuegiensis,  Androcryphaea  confluens, 
Anastrophyllum  leucocephalum,  Bazzania  peruviana,  Harpalejeunea  oxyota  and 
setifera,  Siphonolejeunea  nudicalycina,  Aphanolejeunea  asperrima  and  diaphana, 
Col  lira  bulbosa. 

III.  Chilean  element. —  27  sp.  (21.8  %). 

Chilean  species  without  tropical  connection,  nor  suggesting  Antarctic  rela- 
tions. 

a.  luulemic  (5):  Solenostoma  obtusiflorum  and  rostratum,  Plagiochila  fusco- 
brunnea,   Lejiidozia  fragilhma  and  disticha. 

d.  Also  in  Ciiile,  especially  in  the  south  (18):  Solenostoma  crassulum,  Ana- 
stre[)ta  bifida,  Plagiochila  gayana,  fasciata,  hyadesiana,  deformifolia,  chiloensis, 
rectangulata,  remotidens,  pudetensis,  homomalla,  neesiana,  riparia,  squarrosa, 
robusta,  data  and   Xotarisii,   I  Icrberta  runcinata. 

c.  Cosmop(j]itan  (4):  Plagiochasma  rupestre,  Reboulia  hemisphaerica,  Lunularia 
cruciata,   Marciiantia  p()l\-morj)ha. 

ICven  if  not  the  Juan  r'ernandez  Hepatics  are  almost  entirely  Antarctic  as 
Hkr/jx;  thinks,  the  Antarcto-tcrtiary  element  is  proportionally  larger  than  in  any 
other  grouj),  and  it  is  not  imj)ossible  that  genera  like  Megaceros  or  Harpalejeunea, 
the  island  species  of  wiiich  were  referred  to  the  neotropical  element,  are  Antarctic, 
and  that  this  is  true  also  of  JUizzaiiia,  represented  in  I  and  II,  but  on  the  other 
hand  we  must  admit  that  pantropical  genera  like  Riccardia,  Anthoceros,  Mega- 
ceros, Hazzaiiia,  Lepidozia,  Radula,  Madotheca,  FruUania  etc,  may  have  extended 
far  south  during  a  warmer  period  and  given  rise  to  important  groups  all  around 
the  circumpolar  belt,  and  that  we  have  to  show  that  such  disjunct  southern  groups 


DERIVATION  OF  THE  FLORA  AND  FAUNA  289 

are  interrelated  in  such  a  manner  that  we  can  derive  the  genera  from  a  common 
Antarctic  centre.  This  is  a  problem  that  only  an  expert  and  all-round  hepatologist 
can  solve.  In  our  island  case  the  result  may  be  that  the  Antarctic  element  will 
shrink  and  the  Neotropical  swell;  we  may  have  to  merge  the  Chilean  element  in 
the  Neotropical.  Disregarding  the  cosmopolitan  species,  anthropophilous  and  anthro- 
pochorous  and  very  likely  of  recent  introduction,  we  find  that  the  bulk  of  the 
Chilean  species  is  formed  by  the  14  species  of  Plagiochila\  we  need  a  critical 
revision  of  this  gigantic  assemblage  before  we  shall  know  anything  of  its  origin 
and  history.  Also  small  but  widely  scattered  genera  offer  difficulties:  ^/^^i-Zr^'//^, 
Adelanthus  and  Herberta,  for  instance. 

The  discontinuous  distribution  of  many  southern  genera  and  several  species 
was  commented  upon  by  Miss  Fulford  (/oj.  846): 

The  distribution  patterns  indicate  that  they  were  contemporaries  at  a  time  when  free 
migration  was  possible  between  South  America,  Africa,  Australia,  New  Zealand  and  the 
Sikkim  area  in  N.E.  India.  They  also  indicate  that  the  Antarctic  Continent  has  been  of 
great  importance  in  the  distribution  of  genera  and  identical  species  in  South  America,  the 
Antarctic  Islands  [rm^  Subantarctic],  Australia— New  Zealand  and  probably  Africa. 

Finally,  let  us  repeat  our  comparison  of  the  four  phyla  analyzed.  No  obviously 
Arcto-tertiary  (Boreal)  species  were  distinguished  among  the  Hepaticae.  The  Chilean 
element  accepted  in  the  mosses  and  hepatics  has  been  combined  with  the  Neotropical 
to  form  a  Neotropical-Chilean  group  (NC);  A,  Antarctic;  B,  Boreal. 

Table  V. 

A  NC  B 

Angiosperms 42.2  36.7  15.6 

Pteridophytes 60.4  37.7  1.9 

Mosses 55.8  41.7  2.5 

Hepatics 67.7  32.3  — 

V.  Lichenes. 

Unlike  the  Archegoniates  the  Lichens  offer  insuperable  difficulties  when  we 
try  to  trace  eventual  centres  of  origin  and  evolution.  Of  67  genera  represented  in 
Juan  Fernandez  no  less  than  some  55  have  a  wide,  in  cases  world-wide  distribu- 
tion; they  are  called  cosmopolitan,  subcosmopolitan,  tropical,  temperate  and  so 
forth,  and  with  regard  to  about  70  species  the  situation  is  the  same.  Only  in  a 
very  limited  number  of  instances  a  genus,  or  a  group  of  related  species  is,  to 
judge  from  its  present  distribution,  concentrated  to  a  limited  area,  tempting  us 
to  consider  it  a  centre  of  evolution.  On  the  other  hand  we  must  not  forget  that 
lichens  are  lichenized  fungi;  modern  lichenologists  do  not  regard  them  as  a  spe- 
cial phylum  but  range  them  with  the  different  fungus  orders  and  families  accord- 
ing to  the  taxonomic  position  of  the  fungus  component.  Nothing  prevents  us 
from  assuming  that  the  same  lichen  species  originated  in  different  places  widely 
apart  where  both  fungus  and  microscopic  alga  happened  to  be  present  at  the  time 
when    lichen-forming    fungi    existed   as  independent  organisms.  Nor  should  it  be 

19  ~  557857  The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.      Vol.  I 


290 


C.  SKOTTSBERG 


forgotten  that  numerous  green  and  bluegreen  algae  are  cosmopolitan  and  that  the 
fungus  spores  as  well  as  the  thallogenous  diaspore  (in  fact  more  important  for 
dispersal)  of  lichens  are  quite  resistant  and  adapted  to  long-distance  transporta- 
tion, also  that  they  have  a  greater  facility  to  get  established  where  they  happen 
to  land  than  the  spores  of  ferns  and  bryophytes. 

During  early  Tertiary  times — and  certainly  much  earlier — South  America  had 
a  rich  and  varied  lichen  flora  of  tropical  and  subtropical  character.  With  the  grad- 
ual climatic  diflerentiation  and  the  evolution  of  cold-resistant  forms  the  rising 
Andes  provided  stations  for  both  northern  and  southern  lichens;  migrations  in 
both  directions  helped  to  equalize  the  I^oreal-arctic  and  Austral-antarctic  floras  and 
furthered  the  origin  of  a  bipolar  element. 

With  our  present  insufficient  knowledge  of  the  distribution  of  the  lichens  all 
we  can  say  with  regard  to  a  majority  of  species  or  genera  is  that  they  are  tropi- 
cal-subtropical or  temperate  and  either  northern  or  southern;  it  is  rarely  pos- 
sible to  decide  in  favour  of  the  north  or  the  south  as  a  primary  source  of  the 
Andean  tenifierate  flora.  The  exceptions  from  this  rule  are  few;  for  instance,  gen- 
era like  Pachypliialc,  Lemmopsis  and  Massalongia  may  be  classified  as  northern, 
while  I\vssocaulo}i,  PscudocypJiellarici,  Xephronia  and  Stercocaiilon  are  southern  or, 
if  bipolar,  have  a  southern  centre  making  it  permissible  to  conclude  that  the 
Antarctic  continent  has  played  an  important  part  in  their  history  and  that  the 
bicentric  distribution  of  so  many  species,  also  in  the  larger  genera,  testifies  to  an 
Antarctic  origin  or  at  least  indicates  that  a  transantarctic  migration  route  once 
existed.  Thus  I  have  ventured  to  distinguish  an  Antarcto-tertiary  element  also  in 
the  C'hilean-Juan  I'Y'rnandez  lichen  flora.  Here  as  in  other  groups  the  possibility 
that  a  bicentric  species  may  have  reached  its  stations  from  the  north  must  be 
taken  into  account;  that  a  circumglobal  species  has  reached  southern  Chile  and 
.\ew  Zealand   is  no  {)roof  of  a  transantarctic  connection. 

Below  an  attempt  is  made  to  distribute  the  Juan  Fernandez  lichens  among 
geographical-genetic  groups.  Only  164  species  could  be  included.  No  place  at  all 
could  be  assigned  to  many  of  the  endemic  species — see  above  p.  254 — and  the  fol- 
lowing non-endemic  ones  v.ith  their  disjunct  areas  were  also  excluded:  Bacidia 
delapscDis,  I  sjica  dasypogoides  and  suhtorulosa,  Caloplaca  rubina,  Buelliti  htilophiloides 
'dndfcr)i(nidirjia?ia. 

The  arrangement  proposed  is  far  from  satisfactory  and  will,  I  am  sure,  be 
subjected  to  criticism.  With  our  very  imperfect  knowledge  of  the  real  distribution 
of  lichens  called  "wide-spread"  or  even  "cosmopolitan",  mistakes  in  assigning  a 
species  to  a  certain  element  are  unavoidable.  Thus,  many  were  referred  to  group 
I  with  great  hesitation  because  it  is  iinpossible  to  decide  if  they  have  reached  their 
austral-bicentric  stations  (south  Chile,  New  Zealand)  independently  from  the  north, 
or  it  Antarctic  routes  arc  involved.  The  species  lumped  under  IV  surely  represent 
several   different  distribution   patterns. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  291 

I.  Antarcto-tertiary  element. —  58  sp.  (35.4  %). 
b,  Austral-bicentric;  t,  Austral-tricentric  species. 

a.  Endemic  species  (5):  Psoroma  vulcanicum,  cephalodinum,  dasycladum  and 
angustisectum,  Pseudocyphellaria  berteroana. 

b.  Non-endemic  (53):  Sphaerophorus  melanocarpus,  Phaeographina  scalpturata 
(b),  Byssocaulon  niveum  (b),  Leptogium  phyllocarpum  (b),  Parmeliella  nigrocincta  (t) 
and  pycnophora,  Pannaria fuegiensis  and  rubiginosa (b), Massalongia  carnosa,  Psoroma 
pholidotum  and  sphinctrinum  (b),  Lobaria  crenulata  (b),  Pseudocyphellaria  argyracea 
(t),  intricata  (b),  fragillima  (b),  subvariabilis  (b),  chloroleuca  (b),  cinnamomea  (b),  hir- 
suta,  Guillemini,  gilva  (t),  mougeotiana,  aurata  (t),  nitida,  endochrysea  (b),  Durvillei 
(b),  flavicans  (b),  Freycinetii  (b)  and  Richardi  (b),  Sticta  Weigelii  (b),  lineariloba, 
latifrons  (b)  and  laciniata,  Nephroma  plumbeum,  cellulosum  (b),  antarcticum  (b)  and 
australe  (b),  Catillaria  melastegia  (b),  Megalospora  versicolor  (b),  Phyllopsora  parvifolia 
(b),  Cladonia  pycnoclada,  didyma  (b)  and  aggregata  (t),  Stereocaulon  patagonicum, 
ramulosum  (b)  and  implexum  (b),  Coccotrema  granulatum  (b),  Lecanora  albellina, 
Placopsis  chilena,  fuscidula  (b)  and  parellina  (b),  Myxodictyon  chrysostictum  (b), 
Buellia  halophila  (b). 

II.  Andean  tropical  to  temperate  element. — 54  sp.  (32.9  %). 

a.  Endemic  species  (9):  Arthonia  subnebulosa  and  berberina,  Enterostigma 
Skottsbergii,  Coenogonium  velutinum,  Lecidea  avium,  Pertusaria  hadrocarpa  and 
Skottsbergii,  Lecanora  masafuerensis,  Caloplaca  orthoclada. 

b.  Non-endemic  (45):  Arthopyrenia  cinchonae,  adnexa  and  planorbis,  Pyrenula 
aspistea,  mammillana  and  Kunthii^,  Pyrenastrum  chilense,  Arthonia  complanata, 
Graphis  intricata  and  Dumastii,  Dirinalimitata,  Schismatommaaccedens,  Thelotrema 
lepadinum,  Dimerella  lutea,  Physma  chilense,  Leptogium  moluccanum,  tremelloides, 
cyanescens,  Menziesii  and  callithamnion,  Lecidea  leucoplaca  and  icterica,  Catillaria 
endochroma  and  leucochlora,  Bacidia  endoleuca  and  subluteola,  Toninia  bullata, 
Lopadium  leucoxanthum,  Baeomyces  chilensis,  Acarospora  xanthophana,  Per- 
tusaria polycarpa,  Melanaria  melanospora,  Parmelia  laevigatula,  abstrusa,  nilgher- 
rensis,  soredica  and  microsticta,  Menegazzia  sanguinascens,  Usnea  angulata,  Bom- 
byliospora  dolichospora,  Caloplaca  subcerina,  Theloschistes  flavicans,  Physcia  picta, 
Anaptychia  pectinata,  Cora  pavonia. 

III.  Boreal  element. — 14  sp.  (8.5  %). 

a.  Endemic:  Lemmopsis  polychidioides. 

b.  Non-endemic  (13):  Verrucaria  microspora,  Arthonia  cytisi,Gyalecta  jenensis, 
Pachyphiale  cornea,  Racodium  rupestre,  Lecidea  enteroleuca  and  viridans,  Bacidia 
arceutina,  Acarospora  smaragdula,  Lecanora  dispersa  and  saxicola,  Parmelia  pilosella, 
Buellia  concinna. 

IV.  Pantemperate-Bipolar  to  Cosmopolitan  element. — 38  sp.  (23.2  %). 

Normandina  pulchella,  Diploschistes  actinostomus  and  scruposus,  Peltigera 
rufescens   and    polydactyla,   Lecidea  latypea  and  mutabilis,  Catillaria  intermixta. 


292  C.  SKOTTSBERG 

Rhizocarpon  <^eographicum  and  obscuratum,  Cladonia  bacillaris,  coccifera,  furcate, 
gracilis,  pyxidata,  rtmbriata  and  pityrea,  Pertusaria  leioplaca,  Lecanora  coarctata, 
atra,  polytropa  and  clirysoleuca,  Placopsis  gelida,  Candelariella  vitellina,  Parmelia 
laevigata,  revoluta,  cetrata,  saxatilis,  conspersa,  perlata,  cetrarioides  and  caperata, 
Ramalina  linearis  and  iisnea,  Usnea  florida,  Caloplaca  elegans,  Buellia  stellulata, 
Anaptychia   hypoleuca. 

Chapter  III. 
Composition,  distribution  and  relationships  of  the  Fauna. 

There  are  no  indigenous  reptiles,  amphibians,  freshwater  fishes  or  mammals 
on  the  islands;  of  the  introduced  mammals,  goats,  rats  and  mice  were  naturalized 
centuries  ago  and  during  the  last  20  years  also  rabbits  and  Nasna  rufa. 

Aves. 

Lr)\Ni!KR(;  j-;4  ;  names  in  brackets  used  by  Goodai.l-Johnson-Philippi  (//o). 

Indigenous  land-birds. 

Tunius  ifKii^f/iiuiicus  King  (T.  falklandii  magellanicus).  Both  islands.  North 
Chile  to  Patagonia  and  Fuegia;  according  to 57.  V.  224  not  known  to  be  migratory. 
"More  richly  coloured  with  buff  below  than  specimens  from  the  mainland  avail- 
able to  me",  L()XXP,KRG  wrote  I.e.  3,  a  colouring  characteristic  of  T.falklmtdi- 
cus  (Ouoy  et  (jain),  comp.  5/.  V.  pi.  XIII.  In  no  this  is  called  T.  f.  falklandii  diV\d 
the  island   form  is  referred  to  inagellanicus. 

A}iacretiS  fevjiajideziaiius  (Phil.)  (Spizitornis  f.).  Pandemic  on  Masatierra.  Anae- 
retcs  is  a  neotropical  genus  of  7  species  (Ecuador  to  N.  Argentina  and  Chile). 
A.  fer?iirndc.zi(Uiiis  is  related  to  a  Chilean  species. 

Aphyasiura  Diasafucrae  Phil,  et  Landb.  Endemic  on  Masafuera.  A  second 
species  ranges  from   Centr.   Chile  to  Patagonia  and  Fuegia. 

Ciuclodfs  oustalcti  Scott  ssp.  bacckstroe^nii  Lonnb.  Both  islands,  endemic  but 
very  near  the  typical  species  (Antofagasta-Chiloe).  A  genus  of  13  species  (Ecuad. 
to  Argent.,  Patag.,   I'alkl.   Is.). 

liiislcpluvnis  fiyjuvidcusis  (King  as  'J'rochiliis)  Gould  (Thaumaste  f.).  Pandemic, 
forming  an  endemic  genus  according  to  no.  300:  "Coloracion  totalmente  diferente 
de  la  de  cualcjuier  I'icaflor  cjue  habita  el  continente."  These  authors  leave  open 
the  (juestion  whether  or  not  /:'.  Icyboldi  Gould  of  Masafuera  is  distinct  but  list  it 
as  111.  f.  Icyboldi  and  the  form  from  Masatierra  as  lit.  f.fernandoisis.  L()NNBERG 
p.  7  gave  gofxl  reasons  for  considering  them  as  identical.  The  humming-bird 
seems  to  be  extinct  on   Masafuera. 

l'Mstcpha)ius  gaUritus  Mol.  (as  Trochilus;  Orthorhyncha  sephanoides  Lesson 
et  Garnot  i<S27  —  but  Molina's  name  must  be  about  50  years  older;  Sephanoides 
s.,  Hi)).  Masatierra  and  on  the  mainland  from  Centr.  Chile  to  P^uegia.  Said  to 
migrate  to  the  coast  and  spend  the  winter  there  (/7^),  a  statement  not  quoted 
by  GoODALl.. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  293 

Asio  flammeus  Pontoppidan  (A.  f.  suinda  Vieillot).  Masatierra.  The  typical  spe- 
cies almost  cosmopolitan;  suinda  ranges  over  South  America  from  Venezuela  to 
Fuegia. 

Cercneis  sparverius  (L.)  ssp.  fernandensis  Chapm.  (Falco  s.  f.).  Masatierra, 
endemic,  the  species  distributed  from  North  America  and  the  West  Indies  to  northern 
South  America,  another  ssp.  in  Chile.  A  genus  of  28  species  and  very  wide 
distribution,  but  not  recorded  for  Oceania. 

Buteo  erythronotus  King  ssp.  exsul  Salvin  (B.  polyosoma  e.).  Endemic  on  Masa- 
fuera,  an  accidental  visitor  to  Masatierra;  typical  erythronotus  ranges  from  Peru 
to  the  Magellan  Straits  and  Falkland.  On  Masafuera  the  principal  food  of  the 
buzzard  are  rats,  mice  and  young  goats'  kids,  all  introduced  by  man,  but  it  has 
been  observed  attacking  petrels  and  thrushes.  A  widespread  genus  of  33  species 
(Amer.,  Euras.,  Austral,  Ocean.). 

Breeding  sea-birds. 

Fregetta  grallaria  Vieillot.  Masatierra  and  Santa  Clara  and  also  Desventu- 
radas  (San  Ambrosio);  coasts  and  islands  of  the  Indian  and  Pacific  Oceans. 
4  species,  subtropical-tropical  seas. 

Puffinus  creatopus  Coues.  Masatierra  and  Santa  Clara;  California-S.  Chile, 
where  it  breeds  on  Mocha  I.  and  islands  in  the  vicinity  of  Chiloe.  Migrates  during 
the  winter  to  Peru  and  along  the  coast  north  to  Alaska,  returning  south  in  No- 
vember. The  genus  (28  sp.)  is  world-wide. 

Pterodroma  neglecta  Schleg.  Masatierra  and  Santa  Clara;  also  on  San  Am- 
brosio and  further  reported  from  Lord  Howe  and  Kermadec  Is.  Strolls  north  during 
the  winter.  A  genus  of  about  30  species  spread  over  the  south  hemisphere  and 
extending   north  to  the  north  Atlantic  and  to  Japan. 

Pterodroma  exterfia  Salvin  (P.  e.  externa).  Endemic  on  Masafuera,  migrates 
north  as  far  as  Costa  Rica.  Another  race  breeds  on  Tristan  da  Cunha,  a  third 
on  Kermadec  Is. 

Pterodroma  cooki  Gray  ssp.  defilippiana  Gigl.  et  Salvin.  Endemic  to  Santa 
Clara,  Masatierra  and  Desventuradas.  Typical  cooki  ow  New  Zealand,  P.  cooki  orien- 
talis  Murphy  on  the  coast  of  Peru  and  Chile,  but  breeding  places  unknown ; 
defilippiana  is  said  to  extend  its  flights  to  Peru. 

Pterodroma  cooki  ssp.  masafuerae  Lonnb.  (P.  leucoptera  Masafuerae).  Masafuera, 
endemic.  At  first  Lonnberg  felt  inclined  to  identify  the  bird  with  P.  c.  leucoptera 
"in  spite  of  the  zoogeographical  difficulties  for  such  a  theory"  (p.  15);  LoNNBERG's 
opinion  is  strengthened  by  the  fact  that,  whereas  defilippiana  and  leucoptera  are 
surface-breeding  like  neglecta,  externa  and  masafuerae  are  burrowing. 

Two  aliens  are  naturalized  in  the  islands,  a  melanistic  form  of  Columba  livia 
Briss.  in  the  i8th  century  and  now  common,  d^nd  Lop hortyx  calif ornicus  ShdiW  et 
Nodd^  introduced  191 2  or  191 3.  Several  landbirds  have  been  observed  as  acci- 
dental visitors,  Crymophilus  ftdicarius  and  Buteo  obsoletus  migrants  from  the  north 
2,wd  Belonopterus  chilensis,  Cathartes  sp.,  Cygnus  melanocoryphus,  Haematopus  ater 
and   Circus  maculosus  from  the  opposite  coast.  Petrels,  albatrosses,  Cape  pigeons 


294 


C.   SKOTTSBKRG 


and    penguins     arc     occasionally     seen     around     the    islands    but    do    not    breed 
there. 

Of  the  i;  s{)ecies  l)reedin<^  on  the  islands  3  are  endemic;  of  the  remaining 
12,  6  are  re|)resented  b\-  endemic  subs|)ecies  of  which,  however,  2  also  breed  on 
the  Desventuradas  Islands.  Including  these,  60  %  of  the  birds  are  endemic,  a  high 
figure  in  animals  as  mobile  as  birds  are.  Otherwise,  the  poverty  of  the  island  ornis 
is  noteworth}-,  as  also  the  fact  that  the  affinities  of  the  landbirds  are  all  with 
S.   America. 

With  regard  to  the  actual  distribution  we  can  distinguish  the  following  two 
groups. 

I.    South  American  (especially  Chilean)  group. — 10  sj). 

a.  luulcmic  (6):  Anaeretes  fernandczianus,  Aphrastura  masafuerae,  Euste- 
phanus  fernandcnsis,  Cinclodes  oustaleti  baeckstroemii,  Cercneis  sparverius  fer- 
nandcnsis,   lUitco  erythronotus  exsul. 

/>.  Not  endemic  (4):  Turdus  magellanicus,  luistephanus  galeritus,  Asio  flam- 
meus,   Pufhnus  creatopus. 

II.   South  Pacific  group.  —  5  sp. 

a.  l^ndemic  (3):  Pterodroma  externa  externa,  cooki  defilippiana  and  cooki 
masafuerae. 

i).  Xot  endemic  (2):   Fregetta  grallaria,  Pterodroma  neglecta. 

The  fust  grouj)  includes  of  more  tropical  birds  jhiaeretcs,  Cinclodes  and  Eu- 
stiphaiius,  and  of  more  temperate  Turdus,  Asio,  CevcJuieis,  Buteo  and  Puffiuus. 
Of  the  endemic  species  luistepliauus  fernandcnsis  is  the  most  notable,  in  certain 
characters  a  unicpie  t}'pe  in  the  family  Trochilidae.  The  second  group  is  of  par- 
ticular interest  as  including,  beside  the  widespread  frigate-bird,  four  species  of 
Plcrodroina  not  breeding  on  the  mainland,  where,  perhaps,  a  special  race  of 
/'.  cooki  breeds.  The  genus  is  essentially  austral-circumpolar,  as  it  were  tricentric, 
w  ith  Tristan  da  Ounha  representing  the  African  sector.  Cases  like  those  of /^.  neglecta 
and  externa  call  for  a  common  source  and  suggest  that  Pterodroma  belongs  to 
an  Antarcto  tertiary  element  which  inhabited  the  coasts  and  islands  of  Antarctica 
n   prjglacia!   timjs. 

Oligochaeta. 

Mich Al.l.sKx  (/<S7)  regards  all  the  earth-worms  of  Juan  Fernandez  as  adven- 
titious. The  single  strictly  South  American  Kerria  saltensis  was,  he  believes,  in- 
troduced from  Chile  with  the  human  traffic,  and  this  is  also  true  of  the  three  species 
of  Allolobof^hora,  introduced  to  Chile  from  luirope,  and  by  Friedericia  galha.  The 
occurrence  ot  I\uhydrihis  rcrrucosus  offers  more  interest.  It  was  known  from  Great 
Ikitain,  the  Hebrides,  S.W.  Africa  and  h'uegia,  everywhere  living  on  the  seashore; 
on   Masatierra  it  was  not  found  on   the  beach   but  inland  in  a  freshwater  stream. 

Hirudinea. 

The  leech  discovered  in  19 17  in  the  highland  of  Masafuera  was  described 
as  a  new  species  of  the  Australian  genus  Pliilaenion,   Ph.  skottsbergi  L.  Joh,  [147). 


DERIVATION  OF  THE  FLORA  AND  FAUNA  295 

Whether  or  not  the  two  species  described  from  Samoa  and  Madagascar,  respec- 
tively, belong  to  PJiilaeDion  remains  to  be  settled.  JOHANSSON  expressed  some 
doubts  with  regard  to  the  position  of  the  Juan  Fernandez  leech  and  after  his  death 
the  question  was  taken  up  by  NvBELlN  who  showed  that  it  should  form  a  separate 
genus,  Nesophilaemon  [i88).  With  its  allies  it  forms  a  small  austral  group,  pos- 
sibly of  old  Antarctic  ancestry.  It  must,  of  course,  have  a  host;  it  was  found  in 
the  extremely  wet  Dicksonia  forest,  where  Pterodroma  cooki  masafuerae  makes 
its  burrows,  the  only  possible  host  existing  here  (see  also  14.'/.  442).  Truly  no 
leech  has  been  collected  on  the  bird,  but  extremely  few  specimens  of  this  have 
been  taken  care  of  and  examined. 

Crustacea. 
Amphipoda  (5^). 

OrcJiestia  chileJisis  (chiliensis)  Milne-Edw.  Both  islands,  terrestrial  and  found 
from  near  the  shore  to  almost  600  m  altitude.  It  is  a  bicentric  species,  known 
also  from  Chile  and  New  Zealand. 

Isopoda  (277). 

Beside  3  cosmopolitan  species  2  endemic  ones  have  been  found,  both  be- 
longing to  widespread  genera,  Ligia  litiginosa  and  Philoscia  minjica,  the  latter 
referred  to  a  new  subgenus. 

Arachnoidea. 
Araneae  [22). 

I  am  indebted  to  Professor  Alb.  Tullgren,  who  supplied  much  information 
on  the  distribution  of  the  genera.  Berland  lists  24  species,  of  which  4  are  cos- 
mopolitan and  also  occur  in  Chile;  of  the  remaining  20  one,  belonging  to  the  very 
large  and  widely  spread  genus  Araneus  and  perhaps  new,  was  left  unnamed.  The 
other  19  are  enumerated  below.  Here  as  in  the  following  +  signifies  an  endemic 
species,  +  +  an  endemic  genus.  Mt  =  Masatierra,  SC  =  Santa  Clara,  Mf  =  Masafuera. 

Ariadna  maxima  Nicolet.  Chile. — Mt,  Mf.  The  genus  widely  distributed 
(N.  and  S.  Amer.,  Afr.,  E.  Ind.,  Australia). 

+  Theridion  Baeckstroemi  Berl. — Mt.  The  genus  is  cosmopolitan. 

Th.  gracile  Keyserl.   Chile. — Mf. 

+  Lephthyphantes  Fernandezi  Berl. — Mf.  A  cosmopolitan  genus,  but  only  a 
single  species  recorded  for  S.  America  (Patagonia). 

+  Macrargus pacificus  Berl. — Mt.  The  genus  is  known  from  northern  N.  Amer- 
ica and  Europe,  but  Tullgren  (in  litt.)  doubts  that  it  has  a  wide  distribution 
in  America;  thus,  its  appearance  in  Juan  Fernandez  is  rather  unexpected. 

+  M.  australis  Berl.— Mf. 

+  Leptorhopirum  (.^)  Platei  F.  Cambr. — Mt.  Doubtfully  referred  to  this  Eu- 
ropean genus,  from  which  it  differs  in  certain  characters:  "I'epigyne  est  d'un  type 
tout  a  fait  different"  (22.430). 

+  Tmeticus  Defoei  F.  Cambr. — Mt.  The  genus  used  to  be  quoted  from  N. 
America  and  Europe,  but  TULLGREN  informed  me  that  a  great  many  species  are 
now  referred  to  other  genera. 


296  C.   SKOTTSBERG 

Mtta  uigrohumeraiis  F.  Cambr.— Mt,  endemic?  (see  I.e.  430).  The  genus  is 
known   from  all  continents. 

^Selkirkiclla  alhoiruttata  Berl.  — Mt.  The  genus  is  related  to  the  neotropical 
G)iolus,  known  from  Perii,  l^razil  and  Chile,  and  appears  not  to  be  restricted  to 
Juan  Fernandez;  Hi:rl.\M)  has  seen  a  very  closely  related  species  from  Valdivia 
[22.A12). 

Miivs)naucheuius  scg)>untatus  Simon.  Patagonia  and  Fuegia. — Mt.  A  genus 
of  2  species,  the  second  one  from  the  Magellanian  region. 

^  Misumoiops  Sjoestedti  Berl. — Mt.  An  American,  especially  X.  American 
genus. 

+  (uiycjma  Skottsbergi  l^erl. — Mt.  A  S.  American,  especially  Chilean  genus 
of  numerous  species. 

(',.  niaculaiipes  Keyserl.   Chile. — Mt. 

rOxysoma  Delfini  Simon. — Mt.  A  S.  American  genus. 

^  Plulisca  or)iata  Berl. — Mt.  A  subantarctic-magellanian  genus  extending  north 
into  Chile. 

^/V/.  iugeus  Herl.— Mt. 

■r  Lycosa  I'CDiajidezi  F.   Cambr. — Mt.  The  genus  is  cosmopolitan. 

lii'oplirys  quilpiioisis  Simon.  Centr.  Chile. — Mt.  The  genus  is  known  from 
Centr.  and  S.   America,  ICuroj^e,   S.  Africa  and  Japan. 

Of  the  19  species  enumerated  13  or  perhaps  14  are  endemic  in  the  islands; 
15  (11  or  12  endemic)  are  restricted  to  Masatierra,  3  (2  endemic)  to  Masafuera 
and  a  single  Chilean  species  found  on  both  islands.  The  only  conclusion  we  can 
draw  from  these  figures  is  that  most  likely  only  a  minor  part  of  the  spiders  occur- 
ring on  the  islands  is  known.  It  is  quite  possible  that  there  is  a  marked  differ- 
ence between  the  two  islands,  but  it  is  not  probable  that  Masafuera  is  so  poor 
and  that  Santa  Clara  is  devoid  of  spiders.   Only  a  short  visit  was  paid  to  this  islet. 

The  fauna  makes  the  impression  of  being  almost  entirely  neotropical  or,  at 
least,  S.  American,  with  the  exception  of  the  two  species  of  Macrargus,  which 
are  of  boreal  parentage;  of  the  doubtful /.^//^///^//r//;;/ nothing  can  be  said.  The 
{)resence  of  a  southern,  eventually  Antarctic  element  is  indicated  by  MecysinaucJic- 
)iiiis,  possibly  also  by  riiilisca,  but  so  far  there  is  no  sign  of  a  bicentric  group. 
\\  hethcr  it  can   be  distinguished   in   subantarctic  America  I   cannot  tell. 

Acarina. 

TR.'u;.\Kl)n  [268)  enumerates  28  species,  of  which  2  are  cosmopolitan,  the  re- 
mainder endemic.  He  points  out  that  the  collection,  the  first  ever  made  in  Juan  Fer- 
nandez, undoubtedly  represents  onl}'  a  small  part  of  the  acarofauna;  this  is  evident 
already  from  the  fact  that  not  a  single  sj^ecies  came  from  Masafuera  or  Santa  Clara. 
Whether  the  cosmopolitan  sj)ecies  are  late  arrivals  or  not  is  impossible  to  tell,  but 
very  likely  they  are.  This  would  mean  that  the  entire  acarofauna  is  endemic,  and 
new  investigations  will  not  change  its  independent  character  unless  some  species 
are  discovered  on   the  mainland.   Of  the  23  indigenous  genera  only  one — probably 


DERIVATION  OF  THE  FLORA  AND  FAUNA 


297 


a  second  will  have  to  be  described — is  endemic  and  several  have  a  wide  distri- 
bution. 

The  scant  knowledge  of  this  neglected  group  in  these  regions,  particularly 
in  Oceania,  is  to  be  regretted;  it  certainly  does  not  yet  lend  itself  to  zoogeo- 
graphical  speculations.  Nevertheless  it  deserves  to  be  mentioned  that  Euiergvs 
similis  Trag.  belongs  to  a  genus  hitherto  recorded  only  from  New  Zealand  and 
that  Phyllhermannia  deiitata  Trag.  is  related  to  a  neozelandic  species;  the  genus 
is  also  found  elsewhere. 

Pseudoscorpionidea  (jp). 

+  +  Asterocherftes  vittatus  Beier. — Mt.  The  genus  has  its  greatest  resemblance 
to    Thalassochernes  Beier  from  New  Zealand. 
+  Chelanops  insularis  Beier. — Mt. 

+  Ch.  ktischeli  Beier. — ^Mt.  Related  to  a  Chilean  species. 
+  Geogarypus  bucculentus  Beier. — Mt. 
'V  Parachernes  kuscheli  Beier. — Mt,  Mf. 
+  +  Protoivirthius  fernandezianus  Beier.- — Mf. 
^P.  robustus  Beier. — Mt. 

Neotropical  elements  are  present,  but  species  with  their  relatives  in  the 
Australian-Polynesian  region  are  in  dominance  and  part  of  the  fauna  shows  not- 
able archaic  characters  (I.e.  205). 

Myriapoda. 

The  very  small  and  incomplete  collection — no  specimens  were  brought  from 
Masafuera  or  Santa  Clara — was  studied  by  Verhoeff  {2^4).  In  order  to  get  some 
information  on  the  distribution  of  the  genera  I  asked  Dr.  Otto  Sciiubart  of 
Pirassununga,  Brazil,  for  assistance,  and  he  most  liberally  put  his  wide  knowledge 
of  this  group  at  my  disposal  (letter,  Aug.  27,  1954).  Several  changes  had  to  be 
made  in  the  nomenclature;  the  names  used  by  Verhoeff,  if  different,  have  been 
put  in  brackets. 

Diplopoda. 

Brachyiulus  pusilhis  Leach  (Microbrachyiulus  litoralis  Verh.).  Indigenous  in 
western  Europe,  adventitious  in  N.  America  and  Argentina. 

Brachydesmus  supertis  Latzel.  A  European  species,  adventitious  in  N.  America 
and  Argentina. 

+  Aulacode sinus  insulanus  (Verh.)  Schubart  (Semnosoma,  Verh.).  Endemic.  A 
genus  of  16  species,  distributed  over  Chile  and  Argentina  and  belonging  to  the 
austral  family  Sphaerotrichopidae  (S.  Amer.,  S.  Afr.,  Madag.,  Nossi  Be,  Austral., 
Tasm.,  N.  Zeal.,  N.  Caled.,  Hawaii). 

Nopoiulus  veiiustus  Meinert  (pulchellus  Leach).  Widely  distributed  in  Europe, 
introduced  to  N.  America  and  Chile. 

Cylindroiulus  frisius  oceanicus  Verh.  Typical  frishis  {C.  O^veni  Bollman) 
introduced  to  N.  America,  Argentina  (also  in  forma  oceanicus),  S.  Africa  and  St. 
Paul's  I. 


298  C.   SKOTTSBERG 

Chilopoda. 

+  Xesoj^fop/ii/iis  laticpllis  (Attems)  Schubart  (Geophilus,  Verb.).  Endemic.  The 
^einis,  wbicb  has  not  been  reported  from  S.  America,  includes  after  the  latest  revi- 
sion by  Airr.MS  (as  subgenus  of  Geopliilus)  i  i  species  (i  SAV.  Austral.,  i  N.  Zeal, 
I    X.   Caled..    i    An  nam,   3  Jap.   and   2   Eur.). 

r  Xi'S(>o-i-()p/iiii(s   biU-ckstrocini  (Verb.)  Schubart  (Geophilus,  Verb.).   Endemic. 

Schizotoiia  alaccr  (Pocock)  Silvestri.  Chile,  south  to  Fuegia,  Argentina.  A 
genus  of  6  s[)ecies  (3  Chile  and  Argent,  to  Patag.,  Eueg.,  i  E.  Austral.,  1  N.  Zeal., 
I    Chatham  Is.). 

Litliobio)>wyplia  afyicaua  Porat  (Lamyctes  insignis  Pocock,  insignisbaeckstroemi 
X'erh.).  Widely  distributed  over  Africa;  also  Tristan  da  Cunha,  St.  Paul's  I., 
S.W.  Australia  and  Hawaii.  The  genus  very  wide-ranging  [N.  Amer.,  W.  Ind., 
S.  Amer.  (also  Chile),  Afr.,  \\.  Ind.,  Austral.,  Tasm.,  N.  Zeal.,  Chatham  Is.,  N. 
Caled.,  Kermadec  Is.,  Guam,  Hawaii]. 

If  we  exclude  the  4  species  regarded,  rightly  I  presume,  as  introduced  with 
the  human  traffic,  5  species  remain,  3  of  these  endemic  in  Juan  Eernandez.  This 
is  indeed  a  very  small  number,  but  in  spite  of  being  so  few,  they  tell  a  story  of 
an   austral-circumpolar,   presumably  Antarcto-tertiary  element. 

CoUembola. 

Of  the  8  species  distinguished  by  SciloTT  [216),  the  first  ever  collected  in 
Juan  PY'rnandez.  2  inhabit  Chile,  3  are  known  from  various  parts  of  the  world 
and  3  endemic.  As  long  as  so  little  is  known  about  the  distribution  of  this  group 
it  does  not  lend  itself  to  zoogeographical  speculations.  The  occurrence  of  widely 
dispersed  boreal  species  in  S.  America  and  other  parts  of  the  south  hemisphere 
(Australia,  New  Zealand  etc.)  is  noteworthy,  but  whether  their  wdde  range  is  due 
to  the  great  age  of  CoUembola  or  a  result  of  later  dispersal  is  unknown. 

Thysanura  [222,  2pj). 

■  Is(>U'pis))ia  luiucctois  Silvestri. — Mt,  Mf.  The  specific  epithet  refers  to  the 
intermediate  position  between  IsolcpisDia  and  ll€tcrolcpis7>ia\  the  species  is  com- 
pared with   forms  known   from   Africa  and  Australia. 

'  h  KuscJielocliilis  Ocliai:;aviac  W'ygodz. — Mt.  A  monotypical  endemic  genus 
related  to  Alloinacliilis  and  XesoniacJiilis  from  Australia,  but  not,  as  far  as  known, 
to  an  American   genus. 

Among  the  Invertel)rates  treated  above  the  endemic  leech  offers  great  in- 
terest. Of  Arachnoidea  the  Pseudoscorpionidea  include  a  remarkable  Antarcto- 
tertiary  element,  whereas  the  true  spiders,  strangely  enough,  are  quite  disap- 
pointing  in   this  respect,   even   more  so   than   the  centipedes. 

Insecta. 

In  order  to  get  an  idea  of  the  zoogeography  of  the  island  insects  I  asked  a 
number    of  specialists  for  information  on  the  general  distribution  of  genera  and 


DERIVATION  OF  THE  FLORA  AND  FAUNA  299 

Species.  For  their  readiness  to  supply  me  with  the  necessary  data  I  am  much 
obhged  to  Dr.  Olof  Ahlberg,  Stockholm  (Thysanoptera),  Dr.  KjELL  Ander, 
Linkoping  (Orthoptera),  Dr.  Per  Brinck,  Lund  (Coleoptera),  Mr.  Nils  Bruce, 
Gardby  (Coleoptera),  Dr.  Lars  Brundin,  Stockholm  (Coleoptera),  Mr.  Felix  Bryk, 
Stockholm  (Lepidoptera),  Dr.  W.  E.  China,  London  (Hemiptera),  Dr.  K.-H.  FoRSS- 
LUND,  Stockholm  (Trichoptera),  Dr.  G.  J.  Kerrich,  London  (Hymenoptera),  Dr. 
K.  Princis,  Lund  (Orthoptera),  Mr.  Bo  Tjeder,  Falun  (Neuroptera)  and  Dr. 
B.  P.  UVAR(3V,  London  (Orthoptera). 

JOHOW  (ijo)  enumerates  26  species  of  insects  from  Juan  Fernandez;  some  finds 
may,  I  presume,  have  escaped  his  notice,  but  probably  not  many,  and  it  is  evident 
that  the  entomofauna  was  very  little  known  at  that  time.  During  our  survey  19 16-17 
a  fair  number  of  insects  were  collected  and  many  novelties  were  described  in  vol. 
3  of  this  work,  but  the  collection  gave  the  impression  of  being  very  fragmentary. 
The  intense  collecting  undertaken  in  195  i  and  1952  by  the  Rev.  Dr.  GuiLLERMO 
KUSCHEL  revealed,  however,  the  existence  in  the  islands  of  a  surprisingly  rich 
and  varied  insect  world.  As  Dr.  ALEXANDER,  the  wellknown  specialist  on  Tipu- 
lidae,  expresses  himself  (^.35):  "Father  Kuschel's  collecting  has  completely 
revolutionized  our  knowledge  of  the  insect  fauna  of  the  islands  in  many  groups,  in- 
cluding the  crane-flies" — only  3  species  were  known,  the  number  now  amounts  to  37. 

Until  now  only  a  part  of  Dr.  Kusciiel's  large  material  has  been  worked  up 
by  specialists,  and  I  can  only  refer  to  what  has  been  published  (142,  208,  2g2, 
2gj,  jog,  314),  but  for  some  groups  we  now  have  sufficient  data  to  form  an 
opinion  of  the  zoogeographical  position  of  the  islands  as  far  as  the  insects  go.  At 
the  end  of  1954  Dr.  KusCHEL  joined  my  new  expedition  to  the  islands  and  brought 
back  a  third  very  large  collection.  When  all  his  material  has  been  studied,  the  insect 
fauna  of  Juan  Fernandez  will  be  better  known  than  that  of  most  isolated  islands.  At 
present  about  340  indigenous  species  have  been  recorded,  of  which  about  230(70  %) 
are  regarded  as  endemic.  Dr.  KuscHEL  (letter,  Oct.  16,  1955)  calculates  that  of 
a  total  of  about  600  species  collected  by  him,  about  360  still  await  publication- 
Among  them  are  25-30  flies,  probably  over  50  butterflies,  many  endemic,  at 
least  180  beetles  (more  than  120  weevils,  of  which  4  have  been  introduced 
accidentally,  the  remainder  being  endemic),  and  some  40  hymenopters. 

Orthoptera. 
Dermaptera  [22j). 

+  Euborellia  annulipes  (Lucas). — Mt,  SC,  Mf.  The  genus  S.  Amer.,  E.  Afr., 
Orient,  Ind.,  Ceylon,  Tasm. 

Anisolabis  Bormansi  Scudd.  Galapagos  Is.,  Easter  I.— Mt.  A  large  genus  of 
world-wide  distribution. 

Saltatoria  [j8,  22j). 

^Hoplospyriuin  Skottsbergi  Chopard.— Mt.  An  American  genus,  the  species 
related  to  species  from  N.  America  and  Chile. 

Trimeroiropis  ochraceipeimis  Blanch.   Chile.— Mt.  The  genus  is  American. 


300 


C.  SKOTTSBERG 


Corrodentia. 

Isoptera. 

-  Kiiloti-nfics  i^^racilii^^Jicitus  luiierson. — Mt.  The  only  Termite  known  from  Juan 
Fernandez.  "The  wing  venation  is  close  to  that  of  Kalotermes  broiini  Froggatt 
from  New  Zealand"   [8j.  393). 

Mallophaga  [266). 

Puffimis  creatopus  and  Ptcrodroma  neglecta  and  exterua  are  infested  with  the 
same  mallophagous  parasites  found  on  these  and  related  birds  in  other  regions.  A 
new   Halipcurus  is  mentioned  but  not  described. 

Thysanoptera  (7). 

Aeolothrips  fasciatus  L.  Boreal. — Mt. 

-X-  Physotlirips  Skottsbcrgi  Ahlb. — Mt.  The  genus  is  distributed  over  N.  Amer- 
ica,  luirope,  W.  Asia  and  Australia. 

-SencotJirips  vieptiis  Ahlb. — Mt.  Perhaps  nearest  to  a  Californian  species. 
The  genus  is  otherwise  confined  to  F^urope,  where  it  is  widely  spread. 

rinips  tabaci  Lindem.  X.  America. — Mt.  The  genus  known  from  N.  America, 
luirasia,  \.  Africa  and  Australia. 

The  two  non-endemic  species  may  have  been  introduced  accidentally.  Both 
were   found  in   the  spathe  of  Zantcdcschja  acthiopica,    cultivated  and  naturalized. 

Neuroptera  (zcVf,  g2,  I2j). 

+  +  Couclioptcrella  kusclieli  Handsch. — Mt. 

+  6'.   Diacidata   Handsch. — Mt. 

(layoinyia  falcata  (Blanch.).  Chile,  Argentina. ^ — Mt,  Mf.  A  small  S.  American 
genus. 

'  I Ic))U'yobiiis  Siocstedti  Xavas  (M.  fumosus  l^>sb. -Peters.,  H.  nigrinus  F"sb.- 
Peters.). — Mt.  An  almost  world-wide  genus,  absent  from  the  S.  hemisphere  except 
for  the  Andean   region  (Colomb.,  Perii,  l^oliv.,  Ikaz.,  Argent.,   Chile). 

-i  //.   Skottsbn-gi  Xav.-is.— Mt,   Mf. 

Trichoptera  [21^. 

+  Australoinyia  inastUierra  Schmid.  Mt.  The  genus  is  known  from  Chile^ 
Patagonia  and    I^'alkland   Is. 

■\-A.   luasafuera  .Schmid. — Mf. 

I'ergcr  Por/iri  Xav.   Centr.   Ciiile.  — Mt.   A  Chilean   genus. 

Lepidoptera  (tJ). 

A  great  number  of  genera  and  species  will  have  to  be  added  when  Dr. 
Klsciikl's  material   has  been   determined. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  3OI 

Tineomorpha. 

Gelechidae. 

+  +  Apotheioeca  synaphrista  Meyr. — Mt.  The  genus  is  closely  allied  to  the  large 
and  widespread   Gelechia  (N.  and  S.  Amer.,  Galap.,  Palearct.,  Macaron.,  S.  Afr., 
Australia). 
Oecophoridae. 

-\-Depressaria  relegata  Meyr. — Mt.  Near  a  species  from  the  Andes  of  Ecuador. 
The  genus  is  Holarctic  and  also  found  in  S.  Africa. 

Endrosis  lactella  Schifif. — Mf.  Widely  spread,  domestic. 
Tineidae. 

Monopis  crocicapitella  Clem. — Mt,  Mf.  In  most  parts  of  the  globe,  domestic. 

Pyralimorpha. 

Pyralidae. 

+  Crambus  fernandesellus  Hamps. — Mt.  A  world-wide  genus. 

Elasmopalpus  lignosellus  Zell.   Centr.   and  S.  Amer. — Mt. 

Ephestia  kuehniella  Zell.   Widespread,  domestic. 

+  "^ Fernandocrambiis  Baeckstj'oemi  Kwx'w . — Mt,  Mf.  The  genus  nearly  related 
to   Crambus. 

+  F.  brunneus  Auriv. — Mt. 

+  F.  fuscus  Auriv. — Mt. 

+  +yuania  anmdata  Auriv. — Mt.  Similar  in  some  ways  to  Ptochosiola  Meyr. 
(S.  Afr.,  Australia). 

Nomophila  7iociuella  Schiff.  Cosmopol. — Mt,  Mf;  adventitious. 

■\-Pionea  fiunipefinis  (Warren)  Hamps. — Mt.  A  world-wide  genus. 

Scoparia  Ragonoti  Butl. — Mt,  Mf.  Chile.  A  very  widespread  genus. 

Geometfina. 

Geometridae. 

^  Eupithecia  halosydne  Prout. — Mt.  A  widespread  genus,  but  not  found  in 
Australasia. 

+  E.  (.?)  znepta  Prout.— Mt. 

-\-E.  physocleora  Prout. — Mt. 

■\-Lobophora  insulai'is  Auriv. — Mt.  An  essentially  Palaearctic  genus. 
Tortricidae. 

+  Crocidosema  insulana  Auriv. — Mt.  A  S.  American  genus. 

■vEulia  griseiceps  Auriv. — Mt.  The  genus  Holarctic,  also  in  Hawaii;  few  else- 
where. 

■\-E.  Robinsoni  Auriv. — Mt. 

+  E.  striolana  Auriv. — Mt. 

Noctuina. 

Noctuidae. 

Copitarsia  turbata  Herr.-Sch.  Venezuela,  Colombia. — Mt.  The  genus  in  Mex., 
Centr.  and  S.  Amer.  (Argent.,  Chile). 

Feltia  malefida  Guen.  Amer.  (south  to  Chile);   Macaronesia. — Mt. 


30. 


C.   SKOTTSBERG 


-V  ~  Hoplotaysia  inai^iia  Auriv. — Mt.   Related  to   Copitarsia. 

Li'ucauia  inipuiicta  (iucii.   Chile.— ^It.  A  bipolar  genus  (Palaearct.,  N.  Zeal.). 

-^  Lvcopltotia  luuiks/roojii  Auriv. — Mt.  The  genus  widespread  (Arct.,  Amer., 
luir.,   S.   Afr.,   Madag.,   X.   Zeal.). 

L.   DitssiiDH  Gucn.   Chile,   Patag. — Mt. 

Kac/nplusiii  uu  (iucn.   I'atag..  Urug.,  Argent.,   Chile.— Mt. 

Sv>ii:-yapha  i^-auiDioidts  Hlanch.  Chile. — Mt,  Mf.  A  Palaearctic  genus;  Mex., 
S.   Anicr. 

Rhopalocera. 

Pvrtv/ii/s  cnrvi-  Iluebn.  Widespread  in  S.  America  and  probably  introduced 
in  Juan   bY'rnande/.. — Mt. 

Diptera  [84, 142). 

Acroctyidac  [208^. 

-V  Opcodes    knsclicli   Sabr. — Mt.    A    temperate    genus,  recorded   from  all  con- 
tinents, but  only  2  species  known   from  the  mainland  of  S.  America. 
AiitlioDiyzidae  [12S]. 

Anthony zii  cursor  (Kieffer).   Cosmopol.,  also  S.  Chile. — Mt. 
Calliplioridac  {2-;-;) . 

L  'ally}itropyga  huuteralis  (Walker)  Souza  L.  et  Alb.  (C.  Selkirk!  Enderl),  Chile 
C<)ncej)ci6n). — Mt.   Ml". 

raraludlia  fuh'icrura  (Desvoidy)  Aub.  et  Baxt.— Mt,  SC,  Mf. 

Sarcojicsia  cliloroi^astcr  (Wiedem.)  Arrib.   Chile,  I{^aster  I. — SC,  Mf. 

Sarc())ies}onii)na  hicolor  Souza  L.  et  Alb.  Chile  (Santiago). — Mt,  SC.  A  mono- 
t\-pical   genus. 

Cecidoniyidae  [84,   106). 

+  ^- Tsadaria  pallida  iMulerl. — Mt,  Mf.   Related  to   Canipyloniyza  Meigen. 
(.  hloropuiac  [2()g). 

Ilippilates  ausiralis  Sabrosky  [//.  (Cadrema)  nietallicus  P3nderl.  non  Beck. 
[^  II.  flai'ipcs  (Loew)  Sabr.)].  I^xuad.,  Peru,  Argent.,  Chile. — Mt,  probably  ad- 
ventitious. 

L  uliiidae. 

Ciilcx  intcrfor  Dyar.  — Mt.   Mf. 
J) (die hop odidac  [126). 

^  Ilyilrophorus  hu.ululi  llarmston. — Mt. 

//.  polioi^asUr  (I'hil.)   llarmston.   Chile. — Mt,   SC. 
rSy)npyc?ins  fdiiandtrjrnsis   llarmston.   Near  a   Chilean  sp. — Mt,  Mf. 

liphydridac  [28(/). 

Diniciocnia  cars/a  Iv.d.    Wulp)   Wirth.   Argent.,   Uruguay.— Mt,   SC. 
-V  Discoceriiia  fu))iipc)nus  Wirth.  —  Mt.   Near  a  Chilean  sp. 
Ilyadina  certa  Crcsson.   Chile.  —  Mt,   Mf. 
Ilydrellia  vulgaris  Cresson.   (iuatem.,  l^oliv.,   Chile. — Mt. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  303 

+  Scaiella  a7igustipennis  VVirth. — Mf.    An  almost  Cosmopolitan  genus. 

+  5.  argeniifacies  Wirth. — Mt. 

+  S.  brachyptera  Wirth. — Mt. 

+  5.  decemguitata  Wirth. — Mt,  SC,  Mf. 

^S.  discalis  Wirth. — Mt. 

+  5.  fernandezensis  Wirth. — Mt. 

+  5.  kuscheli  Wirth.— Mt,  Mf. 

+  S.  lutea  Wirth.— Mt. 

■vS.  niarginalis  Wirth. — Mt. 

+  5.  niasatierrensis  Wirth. — Mt. 

■vS.  minima  Wirth. — Mt. 

+  5.  7ia7iopiera  Wirth.— Mt. 

+  5.  pallida  Wirth.— Mt. 

+  S.  pilimana  Wirth.— Mf. 

+  S.  stenoptera  Wirth. — Mt. 

+  S.  vittata  Wirth. — Mt. 

+  Scatophila  fei'naftdesiana  Wirth. — SC. 

vS.  medifemur  Wirth.  Chile  (Coquimbo). — Mt,  SC. 

Heleidae  [288). 

+  Dasyhelea  australis  Wirth. — Mt,  Mf.  Near  a  Chilean  sp.,  genus  cosmop. 

+  Forcipo7Jiyia   te7ttiisqica77iipes    Wirth. — Mt.   A  widespread  genus  (N.  and  S. 
Amer.,  Eur.,  Afr.,  Austral.);   one  species  common  to  Paraguay  and  Australia. 

+  F.  sa7ictaeclarae  Wirth. — Mt,  SC. 

Helo77iyzidae  (128). 

Blaesochaetophora  picticoimis  (Bigot)  Henn.   S.   Chile. — Mt. 

Prosopa7ttru7n    flavifro7is    Tonn.     et    Mall.    (Cnemospathis    Baeckstroemi   et 
Schoenemanni  Enderl.).  Chile,  S.  Africa,  New  Zealand. — Mt,  Mf. 

Lo7ichaeidae  (128). 

Lo7ichaea  patago7iica  Malloch.   Chile. — Mt. 
Mnscidae  (128). 

Austrocoe7tosia  ig7wbilis  (Stein)  Hennig.  Chile. — Mt. 

Ci'aspedockaeta  Ii77ibi7iervis  (Macq.)  Hennig.  S.  Chile. — Mt,  Mf. 

Delia  platiira  v.  sa7icti-jacobi  (Bigot)  Hennig.   Chile. — Mt,  Mf. 

Euryo77i7na  peregri7iU77t  (Meigen)  Hennig.  Peru,  Chile. — Mf. 

Fa7mia  a7tthraci7ia  (Walker)  Hennig.  Chile. — Mt. 

F.  ca7ialicularis  (L.)  Cosmopol.,  also  in  Chile. — Mt,  Mf. 

F.  pu7ictive7ttris  Malloch.  S.  Chile. — Mt. 

Fucellia  i7tte7n7iedia  Lundbeck  (Egeria  masatierrana  et  masafuerana  Enderl.), 
Eur.,  Oceania. — Mt,  Mf. 

Hydrotaea  cya7ieive7itris  Macq.   Chile. — Mt,  Mf. 

Li77i7iophora  patag07tica  Malloch.  S.  Chile,  Patag. — Mt. 

■\- NotoschoeTiomyza  kuscheli  Hennig. — Mt,  Mf. 

Ophyra  caerulea  Macq.   Centr.  Chile  to  Fueg. — Mt. 


304  C.   SKOTTSBERG 

-\-ScliO€)W})iyzJ}ia  eiudejii  liennig. — Mf. 

+  Sylliifniopliora  lispofi'nfia  Hennig. — Mt,  Mf. 

MycetophiUdac  [100]. 

-.  lixicJiia  furcilla  h'reeni. — Mf.   Near  a  Chilean  species;   world-wide  genus. 
-  Leia    tiiallcolns    Freem. — Mt.    Allied    to    a    species    reported    from   Bolivia, 
Peru  and  Brazil;   the  genus  world-wide. 

Macrocera  fujicrca  Freem.   Chile.^ — Mt.  A  world-wide  genus. 

-r  Mvcetophila  a)iij;ustifuyca  P^ndcrl. — Mt,  Mf.  The  genus  world-wide. 

M.  co)iiftra  Freem.   Chile. — Mf. 

J/,  corjiuta  Freem.   Chile. — Mt. 

iM.  fliU'oluuata  hVeem.   Chile.— Mt,   Mf. 

M.  (.')  iusccta  iM-eem.   Chile.— Mt. 

M.  spiiiosa  Freem.   Chile. — Mf. 

+  J/.  subfiiDWsa   Freem. — Mt. 

k- Paraleia  iiephrodops  (r^nderl.  s.n.   Selkirkius)  Freem. — Mt. 

/'.   Jiubilipcuuis  Walker.   Chile. — Mf.  The  genus  neotrop.,  Austral.,  Tasm. 

SciopJiila  ocJircata  Phil.   Chile. — Mt.  A  w^orld-wide  genus. 

Piophilidae  [12S). 

Piophjla  case?  (L.).   Cosmopol.,  domestic. — Mt,   Mf. 
P.  foi'colata  Meigen.   Cosmopol. — Mt. 

Phoridae  [S4). 

-f  Lioyella  juajifcDiajidczica  Enderl. — Mt,  Mf.  A  F^uropean  genus. 
P/iryjuidae  (i2cS'). 

Plirync  fuscipcuuis  Macq.   S.  Chile.— Mt,  Mf. 

l^latypizidac. 

Microsaiiia  pallipes  Meigen.   Cosmopol. — Mf. 
Psycliodidat-  (cS'y,  212). 

PsycJioda    ci}ierea    l^anks.    Cosmopol.,   also  Chile. — Mt.  A   widespread  genus. 
\  l\  t>nuat}eyyc7isis  Satchell. — Mt,  Mf.  Possibly  =  the  following. 
vP.   ))U)nitissima  ICnderl. — Mt. 
/'.  sci>tyi)ii  Tonnoir.   Widespread  in  temperate  regions. — Mt,  Mf. 

Say  cop  hagidac  ( i* -f -f ) . 

Jfypopy^Q^ia  nay'ia  (Walker)  Townsend.   Chile.— Mt,   SC,  Mf. 
Scatopsidac  [S^,  roo). 

■^  ^  Masatieyya  fcyyugbiea    luiderl. — Mt.   Related  to  the  European  Rhaeboza 
linderl.   I^'kkemax  does  not  mention   Masaiicyya. 

Scatopse  fuscipes  Meigen. — Mt,   Mf.   A  world-wide  genus. 

S.   iioiata  (L.).   Cosmopol.,  introduced   with   the  traffic. — Mt. 

Sciayidae  [ioo\ 

+  Brcidysia  fnsca  I^Veem. — Mt,  Mf.   A  world-wide  genus. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  305 

+  B.  media  Freem, — Mt,  Mf. 

+  Merianina  kusckeli  Freem. — Mt,  Mf.  Another  species  in  Brazil. 

+  Psiiosciara  nitens  Freem. — Mt. 

Sunuliidae  (2g4). 

+  Gigantodax  kusckeli  Wygodz.— Mt,  Mf.  An  essentially  Chilean-Patagonian 
genus  with  single  species  as  far  north  as  Mexico. 

Sphaeroceridae  {84,  204). 

Archiborborus  submaculatus  Duda.  S.  Chile,  Patag. — Mt. 

+  +  Gyretria  binodatipes  Enderl. — Mt.  The  genus  is  perhaps  identical  with 
Skottsbergia  Enderl.  and  this  is  merged  into  Leptocera  by  RICHARDS  who, 
however,  does  not  list  the  two  Gyretria  species  described  by  Enderlein,  but 
not  found  in  Dr.  KuscilEL's  material. 

+  G.  crassicosta  Enderl. — Mf. 

Leptocera  brachystoma  (Stenhammar)  Richards.  Cosmopol.,  also  in  Chile. — 
Mt,  Mf.  The  genus  is  widespread  and  well  represented  in  S.  America. 

+  Z.  cultellipennis  (Enderl.  ut  Skottsbergia)  Richards. — Mt. 

L.  darivini  Richards.  Chile,  Argent. — Mt,  Mf. 

L.  divergeiis  Duda.  Peru,  Boliv.,  Chile,  Argent. — Mt,  SC. 

+  Z.  duplicata  Richards. — Mt. 

+  Z.  ellipsipemiis  Richards. — Mt. 

L.  flavipes  (Meigen)  Richards.  Eur.,  N.  Afr.— Mt. 

L.  mediospinosa  Duda.  Cosmopol. — Mt. 

L.  pectinifera  (Villen.)  Richards.  Eur.,  Falkl.  Is.— Mt. 

L.  pulchripes  Duda.  Argent.,  Parag.,  Urug. — Mt. 

+  +Phthitia  alexandri  Richards. — Mt.  The  genus  must  be  very  u^diV  Leptocera. 

+  Pk.   selkirki  (Enderl.  ut  Pterodrepana)  Richards. — Mt. 

+  /%.  venosa  Enderl. — Mt. 

Syrphidae  [96). 

Allograpta  exotica  (Wiedem.)  (A.  Skottsbergi  Enderl.).  Neotropical.  An 
American  genus  of  at  least   16  species,  the  majority  in  S.  America. — Mt,  SC,  Mf. 

^■A.  robinsoniana  Enderl. — Mt,  Mf. 

Melanostoma  fenestratum  (Macq.)  Fluke.    Chile. — Mt.  An  American  genus. 

+  J/.  Lundbladi  (Enderl.  ut  Carposcalis)  Fluke. — Mt. 

Eristalis  tenax  (L.).  Cosmopol,  also  in  Chile. — Mt,  Mf. 

Sterphus  aurifrons  Shannon.   Chile. — Mt. 

Tachinidae  (70). 

Lrtcamyia  chilensis  Aldrich.  Chile. — Mt,  SC,  Mf.  A  S.  American  genus. 
+  Pkanfasiosiphona  kusckeli  Cortes. — Mt.  A  Centr.  American  genus. 

Tendipedidae  (288). 

Anatopynia  vittigera  Edw.  S.  Chile,  Patag. — Mf.  A  world-wide  genus. 

-^Clunio  fuscipenitis  Wirth. — Mf.  A  large,  wide-ranging  genus. 

+  Hydrobaenus  fernandezensis  Wirth. — Mt,  Mf.  The  genus  world-wide. 

20  —  557857   The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.   Vol.  I 


3o6  C.  SKOTTSBKRG 

//.  pnitontni  (Goett^eb.)  Coe.   luigland.   Chile,   Patag. — Mt. 
//.  pusilliis  (luiton)  Coc.   luiglaiul,  Kerguelen. — Mt. 

-^ l\)tio>io)uus  acutus  W'irth.— Mf.  A  genus  of  numerous  species  in  southern 
S.   America,   few  elsewhere  (luir.,   \.   Amer.). 

rP.  t/isiish'/us  W'irth.— Mt.  Mf. 
P.  kirfi-ri  ((iarrett)  lulwards.   l^rit.   Columb.,  Chile,  Eur.— Mf. 

rP.  kuscheli  Wirtii.— Mf. 
/'.  )iii^rinNs  lulwartls. — S.   Chile. — Mf. 

+  /'.  Si'lkirki  W'irth.— Mt.  Mf. 

'Pit?iytars/(S  ffar/pis  (Meigen)  Townes.  \.  Amer.,  S.  Amer.,  also  Chile, 
luir.— Mf. 

'J'i/'iilii/iu-  (7). 

l-lrioptcra  pilipcs  (Fabricius).   Cosmoj)ol.— Mt,   SC,  Mf. 

+  Lii/io?iiij  (I)icranomyia)  affabilis  Alex. — Mt,  Mf.  A  very  large  and  wide- 
ranging  genus. 

■tL.  a))ipliio)iis  Alex. — Mt. 

-f  A.  axierasta  Alex.— Mt. 

+  A.  Jiarpax  Alex. — Mt. 

+  A.   kuscluliaiia  Alex. — Mt. 

-tL.   iiiasafuerae  Alex. — Mf. 

+  A.  pidcsti'is  Alex.— Mf. 

+  A.  selhiiki  Alex.— :\lt.  Mf. 

+  A.  siuardoi  Alex. — Mt. 

L.  tyilubeyculata  Alex.   S.   Chile,   Patag. — Mt,   SC  (an  endemic  variety). 

+  A.   I'oiatrix  Alex. — Mt. 

+  A.   luucris  Alex.— Mf. 

+  A.  yioiqucinia  Alex. — Mt. 

-V  MolophilHs  ainpliacajitlius  Alex. — Mt.  A  world-wide  genus  well  developed 
on   the  mainland. 

+  J/.  ivitiinonis  Alex. — Mt,   Mf. 

+  J/.  apprcssospiiius  Alex. — Mt. 

+  J/.   arcifirus  Alex. — -Mf. 

■V  M .   civiopus  Alex. — Mt. 

+  J/.  iiffocajiiis  /\lcx. — Mf. 

■\M.  (iistifurcus  Alex.- Mt. 

+  J/.  filioliis  Alex.— Mt. 

+  J/.  films  Alex.— Mt. 

+  J/.   uiasafucrac  Alex. — Mf. 

+  J/.  )}U(ltifi(ius  Alex. — Mt. 

+  J/.   fNpfiuii/s  Alex. — Mt. 

+  J/.  pcclijiij'crus  Alex. — Mt. 

+  J/.  rcctispiiius  Alex. — Mt. 

-f  J/,  sclkirkiainis  (Knderl.   ut  Archimolophilus)  Alex.— Mt.     . 

-fJ/.  .sVj'.r  Alex.— Mf. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  ,  307 

+  M.  tridens  Alex.— Mt. 
■vM.  variants  Alex.— Mf. 
-{■M.  yunqiiensis  Alex. — Mt. 

+  Skan?to?toinyia   kuscheli  Alex. — Mf.    An  American  genus,  well  represented 
also  in  Chile  and  extending  north  to  Canada. 
■\-Sh.  masatierrae  Alex. — Mt. 
■\-Sh.  selkirkiana  Alex. — Mt. 
+  Tipula  baecksU'oemi  Alex. — Mt.  The  genus  world-wide. 

Coleoptera. 

As  yet  little  has  been  published  about  the  beetles  collected  by  Dr.  KUSCIIEL; 
to  judge  from  what  is  known  the  number  of  island  species  no  doubt  will  be 
multiplied. 

Anobiidae  [igs\ 

Anobiuni  punctatMin  De  Geer.  Cosmopol,  introduced. — Mt.  Beside  the  typical 
species  an  endemic  ssp.  described  by  PiC  (Mt). 

A.  striatum  01.  Cosmopol.,  introduced. — Mt. 

+  Calymmaderus  atronotatus  Pic. — Mt,  Mf.  Near  a  Chilean  species.  Numerous 
species  in  N.  and  S.  America. 

+  ^ Masatierrum  inipressipeime  Pic. — Mt,  Mf.  A  genus  near  Megorama  Fall., 
a  small  N.  American  genus. 

Stegobium  (Sitodrepa)  paniceum  (L.).  Cosmopol.,  domestic. — Mf. 

Xyletomerus  pubescens  ssp.  kuscheli  Pic. — Mt,  the  ssp.  endemic.  A  north 
American  genus. 

X.  pubescois  Md,x.  piceitarsis  Pic  (fumosus  var.,  Pic). — Mt.  The  variety  endemic. 

A  nth  ribiidae  [133] . 

+  +  Opisolia  lenis  Jordan. — Mt.  Related  to  Eucyclotropis  Jordan  (Centr.  and 
S.  Amer.). 

Bostrychidae  [166). 

Neoterius  pulvinatus  Blanch.  Chile. — Mt.  A  small  genus  reported  from  Peru 
and  Chile. 

Prostepha7ius  sulcicollis  Fairm.  et  Germ.  Chile. — Mt. 

Carabidae  (5,  260). 

Bembidimn  inconstaris  Solier.  Chile. — Mt.  A  world-wide  genus. 

B.  punctigerum  Solier.  Chile. — Mt. 

Laemosthenes  complanatus  Dejean  (Pristonychus,  Gory,  Andrewes).  Cosmopol., 
also  Chile;  introduced.— Mt,  Mf. 

+  Metius  eurypterus  Putzeys. — Mf.  A  S.  American  genus,  mostly  in  the  far 
south. 

M.  flavipes  Dejean.  Chile.— Mt,  SC. 

+  M.  kuscheli  Straneo.— Mt. 

+  J/.  ovalipennis  Straneo. — Mf. 


jo8  C.   SKOTTSBERG 

+  Plerostichus  kusclicli  Straneo. — Mt.   A   world-wide  genus. 

+  /V.  selkirki  Aiulrcwes.— Mt.   SC. 

+  /V.  skottshfyi:;i  Andrewes. — Mt. 

+  /V.   H'alkiri  Andrewes. — Mt. 

+  +  Iraclnsivus  hasalis  Straneo. — Mt.  The  genus  presumably  endemic  [260. 

■3«). 

■T  '/'.  bicolor  Straneo. — Mt. 
+  r.  emdi'iii  Straneo. — Mt. 
-f  /".   kuscluli  Straneo.— Mt. 
+  7".  (>i'ij/ipiN}iis  Straneo.— Mt. 
f  '/'.  pallipcs  (ierm.— Mt,  Mf. 
+  /'.  pH)ntii:;ey  Andrewes. — Mf. 
+  7.  sericeus  Andrewes. — Mt. 

+  Trechisihus  bacckstfoe)}ii  (Andr.)  Straneo. — Mf.   An  American   genus. 
7".  fe»ioralis  Germ.  Chile. — Mt  (end.  ssp.),   SC. 
+  7".   kusclicli  Jean n el. — Mt. 
r  /  'ariopalpus  crusoci  Keed. — Mt,   SC. 

(  Inysoinclidac  [2S2,   iS]. 

^  vMiJiotula  fcffiivnicziiVia  l^echyne. — Mt.  The  genus  is  related  to  Hyp7io- 
pJiila  (W.   ICur. -Japan). 

\-M.  kuschcU  Hechyne.— Mt. 

fj/.  iiitois  W'eise. — Mt. 

Cioidac  [166). 

■  (  is  hi))iaculatiis  Germ.— Mt.   A  world-wide  genus. 
+  C  fcy}ia}idczia)ins  Lesne. — Mt. 
r  C '.   nifus  (jerm. —  Mt. 

i  Icridac  [21^;). 

Xccrohia  rufipcs   De  (ieer.   Cosmopol. — Mt,   introduced. 

C  Occific/Iidnc  [2iS2). 

liriopis  opposita  (nicr.  Chile. — ^It.  The  genus  ranges  from  Vancouver  I.  to 
Patag.   and   I'"uegia. 

C(dydiidac  [2oiA. 

^  /[ycHonicrodcs  uiasd/ucroisis  Poj)e. — Mf.  lieside  the  two  Juan  Fernandez 
species  there  is  a  third   in    New   Zealand. 

'r I\   ni(isa/icyrc>!S/s   I'ope. — Mt. 

V  Pyotoincrus  iiisularis  (ironvelle. — Mt.  The  genus  N.  and  S.  Amer.,  E.  Ind., 
Japan,   .Australia,    \.   Zeal,   (numerous),   .Samoa. 

4  7'.  j^cnudiiii  Pope. — Mf. 

C  yyptophagidac  {j;6]. 

Cyyptopliai::us   atomayioidcs    (iron v.    (Selkirki    l^ruce').    Chile.— Mt.    A    world- 
'   Letter   19.9.  1954. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  309 

wide  genus,  the  subg.  Mjiionomus,  where  the  island  species  belong,  in  Eur.,  N.  and 
Centr.  Afr.,  Macaron.,  Centr.  Asia,  probably  also  N.  Amer. 

+  C.  Skottsbergi  Bruce. — Mt. 

+  C.  splendens  Bruce. — Mt. 

+  +  Cryptosomatula  longicornis  Bruce. — Mf. 

+  Loberosche?na  convexum  Bruce. — Mt.  Beside  the  island  species  3  in  Bo- 
livia and  2  in   Chile. 

+  Z.  discoideum  Bruce. — Mt. 

Curculionidae  [12).  Dr.  Kuschel,  who  specializes  in  this  family,  estimates  the 
number  of  species  collected  by  him  to  exceed  one  hundred. 

+  +  Arwletkrus  gracilis  Auriv. — Mt. 

+  -v Apteronanus  dendroseridis  Auriv. — Mt. 

+  A.  (.^)  gunnerae  Auriv. — Mt. 

Arajnigus  Fulleri  Horn.  A  widespread  noxious  beetle. — Mt. 

+  Caulophilus  (.?)  nigrirostris  Auriv. — Mt.  The  genus  in  southern  U.S.A., 
Centr.  and  S.  America.    It  does  not  exist  on  Juan  Fernandez  (KuscilEL  in  litt.). 

+  Cyphometopus  masafuerae  Auriv. — 'Mf.  3  species  in  Chile,  where  the  island 
species  most  likely  also  occurs  (Kuschel  in  litt.). 

■^  -^Juanobia  ruficeps  Auriv. — Mt. 

+  ^Juanorhinus  Robinsoni  Auriv. — SC. 

Otiorrhynchus  rugosostriatus  Goeze.  W.  and  S.  Europe. — Mt,  accidentally 
introduced  via  Chile. 

+  -v  Pachy stylus  dimidiaius  Wollaston. — Mt. 

+  P.  nitidus  Auriv. — Mt. 

+  Pachy tragus  crassirostris  Wollaston. — Mt. 

+  Pentarthrujn  affijte  Wollaston. — Mt.  A  widespread  genus,  found  on  many 
oceanic  islands. 

+  P.  nigropiceum  (Phil.)  Auriv. — Mt. 

+  P.  nitidum  Wollaston. — Mt. 

+  /^.  rufoclavatum  Auriv. — Mt.   Close  to  P.  ^//V^/^' Broun  from  New  Zealand. 

+  +Platynanus  arenarius  Auriv. — Mt. 

+  P.  Baeckstroemi  Auriv. — Mt. 

+  P.  hirsutissimus  Auriv. — Mt. 

+  P.  quadratifer  Auriv. — Mt. 

+  P.  sericatus  Auriv. — Mt. 

+  /^  Skottsbergi  Auriv.— Mt. 

+  Strongy/optei'us  nitidirostris  Auriv. — Mt.  The  genus  is  also  found  in  Chile 
and  New  Caledonia. 

S.  ovatus  Boh.  Chile.— Mt,  Mf. 

Dermestidae  (104). 

Dermestes  vulpinus  L.   Cosmopol.,  introduced. — Mt. 

Dytiscidae  (2^p,  ii?)- 

Afiisomeria  bisiriata  Brulle.  Chile. — Mt.  The  genus  known  from  Tristan  da  Cunha. 


3IO  C.   SKOTTSBERG 

^Lancctcs  Baeckstrocnii  Zimmerm. — Mf.  Tlie  genus  austral  bicentric. 
RliaJitHs    sigjiatiis   ssp.    knsclieli  Guignot. — Mt,    Mf.    An    endemic    variety  of 
a  Chilean  species. 

Latlnidiidae  [ig6). 

r  Co)ii)ioi)ius    curtipe}uiis    I'ic. — Mt.    Near   C.  dijuidiaius  Belon  (Boliv.,  Chile). 
An   essentially   luiropcan   genus. 

C.  suhfasciatus  Reitt.   Chile.— Mt. 

Milasidae  [joS). 

^ rscudodiiurctiis  Sclk'nki  Flet. — Mt.   An  Argentine  genus. 

Mycetophaoidae  [ic)6). 

Mvit'/op/iajrus  c/iili?isis  Phil.  Chile. — Mt.  The  genus  in  Kur.,  Asia,  Afr.  and  Amer. 

Xitidididae  [nj6,  loj). 

rC)iips  acuta  Ciillogly.— Mt.  A   Chilean  genus. 

4-r.   atrata  (iillogly.— :\Tt. 

t  L '.  dii'vrsa  Pic— Mt. 

+  6".  fi'Diandczia  (jillogly. — ?^It. 

•C.   niucrojiis  (iillogl)-. — Mt. 

Scarahaiidae  (J'//). 

Aphodius  iiranariiis  L.   Cosniopol.,  domestic. — Mt. 

Plfuropliarus   cacsus    Creutz.   Reported    from   X.  Amer.,   Chile,  Eur.,  Orient, 
.Madagascar. — Mt,   introduced. 

Scolytidac  \^^o^)). 

(iiiathotriclius  tortliyloidcs  Schedl.  Chile. — Mt.  The  genus  in  N.  and  S.  America. 

IVilocotribus  ivillei  Schedl.  Peru  and  Chile. — Mt. 
Sti  ip  //]  'li)iida  e  ( J* -r ) . 

■  Atluta    Kob'nisoui   l^ernhauer. — Mt.   Near  a  Brazilian  species.   A  cosmopoli- 
tan  genus  of  about   2000  sj)ecies. 

■  liltusis  sonirufa   h'airni.  et  (ierm. — Mt.  A  genus  of  about   1 50  species,  S. 
Amer.  (also  Chile),   .Austral.,   N.   Zealand. 

\Mid(>)i    irusdi-aiius    Bcrnhauer. — Mt.    A    cosmoj)olitan    genus  of  about  500 
sj)ccics. 

r  Ocyusa  l^acckstrocDu  Bcrnhauer. — ?^It.  About  40,  mainly  Palaearctic. 
IViilo)itlius    nitidiponiis    Solier.    Chile. — Mt.    A  cosmopolitan  genus  of  about 
(Soo  species. 

I   7'rooop/ti(>fns  Sko/Zs/xr^i;-/  Bernhauer. — Mt.    A  cosmopolitan  genus  of  about 
350  -^P- 
I I'DuiocJiilidac  ( / 9"^ ). 

+  r/ianodi-s/a    crihraria    (Blancli.).    Includes    Ph.  auirulata   Reitt.— Mt,   SC.   A 
Chilean   genus. 

+  /'//.   robust  a  Pic. — Mt. 

+  /V/.   rarici^ata  Germ.-Mt. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  3II 

Tenebrionidae  [lO/f.,  ig6). 

Blapstinus  punctulatus  Solier.  S.  Amer.,  also  Chile.— Mt,  SC.  Numerous  spe- 
cies in  N.  and  Centr.  Amer.,  some  in  S.  Amer. 

^Enneboeus  Baeckstroemi  Pic— Mt.  Near  a  species  from  Panama.  The  genus 
Mex.-Colomb.,  Tasmania. 

Nycterinus  gracilipes  Phil.  Chile.— Mt.  Numerous  species  in  Chile. 


Hymenoptera. 

Aphelinidae  (211). 

Aphelinus  jucmidus  Gahan.  N.  America. — Mt,  Mf. 

Bethylidae  [igi). 

+  Cephalonomia  skottsbergi  Brues. — Mt. 

+  +Lepidosternopsis  kuscheliana  Oglobin. — Mt. 

+  Perisieroia  maculicornis  Oglobin. — SC.  A  widely  distributed  genus  of  about 
25  species. 

+  P.  sanctae  clarae  Oglobin. — SC. 

Braconidae  [i8y). 

+  Apa?iteles  evadite  Nixon. — Mt,  Mf.  A  wide-spread  genus. 

^■A.  morroensis  Nixon. — SC. 

Aphaereta  minuta  (Ns.). — Mt. 

+  Opms  kuscheli  Nixon. — Mt,  Mf. 

^0.  scabriventris  Nixon. — Mt. 

Dryijiidae  (igi). 

■^Haplogonatopns  insularis  Oglobin.^ — Mt.  A  genus  of  6  species  (N.  Amer,, 
J.  Fern.,  Australia,  Pacif.  Is.). 

+  +  Idologonatopus  nigrithorax  Oglobin. — Mf.  A  genus  related  to  the  former. 

Elachertidae  [211). 

+  +Kus  chelae  her ius  acrasia  De  Santis. — Mt,  Mf. 

■^ Pseudelachertus   semijiavus    De    Santis. — Mt.     The    genus   otherwise   Aus- 
tralian. 

Encyrtidae  [211). 

+  Hemencyrtus  kuscheli  De  Santis. — Mt,  Mf.  A  neotropical  genus. 

Eniedontidae  (211). 

+  Achrysochris  bicarinata  De  Santis. — Mt,  Mf. 

Eupai^acrias  phytomyzae  (Brethes)  De  Santis.  Chile,  Argentina. — Mt,  introduced. 
+  Omphalomorphella  elachertiformis  De  Santis.    Said  to  come  near  an  Aus- 
tralian sp.— Mt,  Mf. 

Eulophidae  (211). 

■V Diaulomyia  calvaria  De  Santis. — Mt,  Mf.  Allied  sp.  in  Australia. 


312  C.  SKOTTSBERG 

Formicidae  [2SJ). 

Ponera  trigoua  Mayr  var.  opacior  Forel.  \.  Amer.,  W.  Ind.,  Chile,  Argent. 
— Mt.  The  typical  sfjecies  in   Ikazil.  Probably  spread  with  the  traffic. 

Proiolepis  obscura  Mayr  ssp.  raga  Forel.  Melanesia. — Mt.  The  typical  spe- 
cies Java,  Australia,   Hawaii,  another  variety  N.   Guinea  and  Melanesia. 

Tet}a))ionu))i  guiiueiise  (Fabricius).  An  African  ant,  now  widely  spread  with 
the  human  traffic. — Mt. 

IcJniciDnonidae  {2o6\ 

Rnicospilus  purgatus  Say.  Temperate  N.  and  S.  America. — Mt,  Mf.  An  al- 
most world-wide  genus  (Amer.,   luir.,  Afr.,  Austral.,  N.   Zeal.,  Hawaii). 

-\-  llcniiteles  Bacckstroe})ii  Roman. — Mt.  An  almost  world-wide  genus,  less 
rich   in  the  tropics. 

+  //.  luasafuerae  Roman. — Mf. 

+  flo/ocn'UDia    {^)  juaniaua  Roman. — Mt.   The  genus  is  known   from  Europe. 

Mettlia  (Paniscus)  gerliugi  Schrottky.  Chile. — Mt.  Range  of  genus  very 
wide,  including  S.   Amer.,  Falkl.   Is.,  Rodriguez,  Austral.,  N.  Zeal. 

Stilpiius  gagatcs  Grav.  var.  Robi)iso7ii  Roman. — Mf.  The  typical  species  in 
luirope,  the  genus  also  in  N.  America  and  Greenland. 

Tyiptognathus  aequiciiicius  Spin.  Chile. — Mt. 

MyDiaridae  [igo). 

Aiiagnis  incaniatus  Hal.  Palaearctic. — Mf,  undoubtedly  introduced. 

+  +  CroimoDiytiiar  fcrnandczi  Oglobin. — Mt. 

■V  C.  jniperfcctus  Oglobin. — ]\It. 

+  +  Xcsopolynema  caudatuui  Oglobin. — Mt. 

P()ly}u))ia  fiiscipes  Hal.  Palaearctic. — Mf,  supposed  to  have  been  accidentally 
introduced. 

+  V ScolopsoptcroJi  kiiscJieli  Oglobin. — Mt. 

Rhynchota. 

Heteroptera. 

A)itli()coyidac  {2 1\ 

-^ Hucliivianiclla  dcria  Ik-rgroth.— Mt.  Related  to  B.  continua  B.  White  from 
Madeira;  other  species  reported   from  Tasmania  and  Hawaii. 

Lyctocoris  catupcslris   Vi\h\.   Cosmopolitan,   probably  adventitious. — Mt. 

Lygacidac  {2/,   j6i). 

'  V  Micrymcuus  kuschcli  Kormilev.— Mt.  Most  nearly  related  to  Metagerra 
H.  White   from   New   Zealand   (Kokmilfa). 

+  J/.  seclusus  Ik-rgroth. — Mt. 

vXysius  Bacckstrooni  Hergroth.— Mt,  Mf.  An  almost  cosmopolitan  genus 
witii  numerous  species  in  Xew  Zealand  and  Melanesia,  east  to  Samoa;  greatest 
concentration  in  Hawaii.  X.  Pcrrcks/rorifii  is  closer  to  X.  Ilutioni  V>.  White  from 
Xew  Zealand   than   to  any  American   species  (KoKMlLKv). 

+  V  Robnisowclioris  tingitoidcs  Kormilev. — Mt.   P^orms  a  separate  tribe. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  313 

Miridae  (50). 

+  Derophthabna  ferna7ideziana  Carv. — Mt.  A  neotropical  genus  (Braz.,  Argent., 
Urug.). 

+  +Kusche liana  masatierrensis  Carv. — Mt. 

Nabididae  [21). 

Nabis  (Reduviolus)  pujictipennis  Blanch.  Chile. — Mt,  Mf.  A  world-wide  genus. 
Reduviidae  (2g2,  jio). 

Empicoris  (Ploeariodes)  rubrornaculaius  (Blackb.).  Almost  cosmopolitan. — Mt, 
probably  adventitious. 

+  Metapterus  additius  Wygodz. — Mt.  A  wide-ranging  genus  (Amer.,  also 
Chile,  S.  Eur.,  N.  Afr.,  W.  Asia). 

■vM.  kuscheli  Wygodz.— Mt. 

^M.  fnasatierrensis  Wygodz. — Mt. 

Ploiaria  chilensis  (Phil.)  Kuschel. — Mt,  Mf.  Almost  cosmopolitan;  also  in 
Chile  and  probably  adventitious  in  Juan  Fernandez. 

Homoptera. 

Apkididae  (communicated  by  Dr.  Kuschel):  4  introduced  species  on  garden  plants. 
Cicadellidae  (57). 

+  +Evansiella  kuscheli  China. — Mt. 

Delphacidae  (314). 

+  Nesosydne  sappho  Fennah. — Mt.  A  genus  known  before  from  south  and 
central  Pacific  islands  including  Hawaii,  but  never  reported  from  America. 

-f-A^.  minos  Fennah. — Mt,  Mf. 

+  N.  oreas  Fennah. — Mt. 

+  N.  calypso  Fennah. — Mt,  Mf. 

+  N.  philoctetes  Fennah.— Mt,  Mf. 

+  N.  vulca7i  Fennah. — Mt. 

^Delphacodes  kuscheli  Fennah. — Mt.  A  widely  distributed  genus. 

■vD.  (Sogata)  selkirki  (Muir)  Fennah. — Mt. 

Jassidae  [21). 

-f  +  Alloproctus  amandatus  Bergroth. — Mf. 

Some  zoogeographical  statistics. 

Orthoptera. — Of  the  four  species  known  2  are  endemic  but  of  American 
affinity,  one  a  Chilean  species  and  one  reported  from  Galapagos  Is.  and  Easter  I. 

A^^^r^/>/^r^.— Endemism  strong,  4  species  of  5,  2  forming  an  endemic  genus, 
the  fifth  an  American  species.  Relations  presumably  Andean. 

Lepidoptera.—0{  the  26  indigenous  species  18  (69  %)  are  endemic,  and  there 
are  four  endemic  genera.  Of  these,  Apothetoeca  and  FernaTidocrambus  are  related 
to  world-wide  genera  also  represented  in  S.  America,  Hoplotarsia  to  an  American 


314  C.   SKOTTSBERG 

genus;  the  systematic  position  of  JuaJiia  has  not  been  stated.  Together  they 
include  6  species.  Of  tlie  remaining  12,  7  belong  to  genera  with  a  large  to  world- 
wide area  including  at  least  some  part  of  America,  and  one  belongs  to  an  American 
genus.  Kulia  (3  sp.)  and  Lohopliora  are  essentially  boreal.  The  8  non-endemics 
are  found  in   Chile  or  in  some  other  part  of  S.  America. 

The  total  absence  of  all   indigenous  Rhopalocera  is  remarkable. 

Diptera. — At  present  157  named  species  belonging  to  27  families  have  been 
reported.  Nine  or  ten  species  at  least  have  been  introduced  with  the  human  traffic. 
147  are  thought  to  be  indigenous  and  of  these  94,  64%,  are  endemic.  Con- 
sidering our  insufficient  knowledge  of  the  dipterofauna  of  Chile,  too  much  weight 
should  not  be  laid  on  these  figures,  but  even  if  quite  a  few  of  the  insular  en- 
demics will,  in  the  future,  be  discovered  on  the  mainland,  I  trust  that  a  fair 
number  will  remain,  sufficient  to  show  the  peculiar  character  of  the  fauna.  Be- 
sides it  can  be  foreseen  that  Dr.  Kusc;ilEL's  new  material  will  bring  to  light 
some  remarkable  additions. 

The  fauna  is  not  a  haphazard  crowd  of  wind-drifted  flies.  It  gets  its  stamp 
less  from  the  few  endemic  genera — of  10  new  genera  proposed  by  Enderlein 
only  4  remain — than  from  the  presence  of  six  non-endemic,  S.  American  or 
more  wide-ranging  genera  with  six  or  more  species  each,  MolopJiilus  with  19 
(all  end.),  Scatella  with  16  (all  end.),  Lii)wiiia  with  13  (12  end.),  Leptocera 
with  10  (3  end.),  MycctopJiila  with  7  (2  end.)  and  Podoiwmus  with  6  (4  end.), 
together  71   species  of  which   56  (79%)  are  endemic. 

The  Xeotropical-Chilean  character  of  the  fauna  is  obvious.  This  is  what  we 
ex[)ect  (juite  ai)art  from  what  we  may  think  about  the  history  of  the  fauna,  but 
the  almost  total  absence  of  even  a  small  austral-circumpolar  or  Pacific  element 
is  noteworthy;  the  only  examples  w^ould  be  Prosopantruni  flavipes  (austral- 
tricentric]  and  Inicellia  jjitcDJicdia,  said  to  be  distributed  over  "Oceania".  There 
are  some  striking  cases  of  disjunction,  suggesting  bipolarity  {IJoyella,  Hydrobacniis, 
Podouoifius  Kiejferi],  but  the  distances  will  perhaps  be  lessened  when  the  dis- 
tribution  becomes  better  known. 

CoUoptoa. — I  want  to  emphasize  that  of  Dr.  Klschel's  collections  ony  5 
families  have  been  worked  out;  it  is  to  be  regretted  that  no  list  of  the  Cur- 
culionids  is  available.  On  the  other  hand  I  believe  that  the  beetles  inhabiting 
the  Chilean  mainland  are  better  known  than  the  flies  and  some  other  insect 
groups  so  that  the  proportion  between  endemics  and  non-endemics  will  not 
undergo  very  great  change  in   the  future. 

The  number  of  named  s|)ccies  hitherto  reported  from  Juan  Fernandez  is 
103,  belonging  to  19  families,  [)erhaj)s  little  more  than  V'.i  of  the  species  found 
there.  Eleven  species  are  anthropochorous.  Of  the  remaining  92  no  less  than  74  are 
endemic — 80  "o,  only  20%  having  been  found  elsewhere.  Future  research  will 
alter  these  figures,  I  suj)p()se,  a  number  of  island  endemics  will  be  stated  to 
extend  to  Chile  and  vice  versa,  but  on  the  other  hand  we  have  good  reason 
to  expect  that  practically  all  Curculionids  collected  but  not  yet  described  will 
j)rove  to  be  endemic;  of  22  indigenous  species  enumerated  by  AURIVILLIUS  21 
were  described  as  new. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  315 

So  far  49  indigenous  genera  are  cited,  of  which  lO  are  endemic;  five  of 
these  are  CurcuHonids. 

The  non-endemic  species  are,  with  one  exception — Xyletomerus  piibescejis,  a 
N.  American  species  represented  by  2  endemic  varieties — also  found  in  Chile. 
As  yet  no  austral-bicentric  or  -tricentric  species  have  been  found.  However, 
Pycnomerodes  with  2  species  in  Juan  Fernandez,  i  in  New  Zealand  and  none 
elsewhere,  as  well  as  Pyawmerus,  Stroiigylopterus,  Eleusis  and  Enneboeus  suggest 
the  existence  of  a  small  austral,  possibly  Antarctic  element,  even  if  the  area 
in  cases  extends  north  of  the  Equator. 

Hymenoptera. — The  35  registered  species,  5  of  them  adventitious,  cannot 
represent  but  a  minor  portion  of  the  fauna.  Just  as  in  all  other  insect  groups 
endemism  is  strong,  23  species  are  endemic  [']6.6%]  and  of  the  26  genera  6.  The 
affinities  were,  as  a  rule,  not  indicated  by  the  authors;  very  likely  they  are, 
with  some  exceptions,  with  S.  American  forms.  Haplogonatus  is  essentially  south- 
ern, Prenolepis  obscura  is  a  southern,  mainly  Pacific  ant.  ApJielms  jucu7idus  and 
Stilpnus  gagates  are  said  to  be  boreal,  but  in  these  as  in  other  similar  cases 
the  possibility  of  accidental  introduction  must  be  considered. 

Hemiptera. — The  21  indigenous  species — there  are  7  adventitious  ones — are 
by  no  means  a  fair  representation  of  the  Rhynchota  inhabiting  the  islands.  Dr. 
Kuschel's  collection  contains  twice  as  many  species,  more  than  half  of  them 
endemic;  of  the  21  named  species  20  are  restricted  to  Juan  Fernandez.  Where 
3  of  the  4  endemic  genera  have  their  relatives  I  cannot  tell,  but  the  fourth,  Micry- 
menus,  is  most  nearly  allied  to  a  genus  in  New  Zealand.  Buchaiianiella  is  quoted  for 
Madeira,  Tasmania  and  Hawaii,  but  not  from  America,  Nysius,  a  world-wide 
genus,  has  a  stronghold  in  New  Zealand  and  in  the  Pacific,  and  the  single  is- 
land species  stands  nearer  to  a  species  endemic  in  New  Zealand  than  to  its 
American  congeners.  It  seems  likely  that  we  have  to  do  with  an  austral- 
antarctic  element. 

Mollusca. 

The  following  synopsis  is  based  on  information  supplied  by  Professor  NlLS 
Odhner  who  put  his  unique  knowledge  of  this  group  at  my  disposal.  The 
additions  and  changes  in  his  earlier  list  (i8g)  are  entirely  due  to  him,  and  I 
thank  him  for  invaluable  assistance.  Possibly  the  new  material  brought  back  by 
our  1954-55  survey  will  give  additional  taxonomic  results,  but  they  will  not  alter 
the  zoogeographical  position  of  the  fauna. 

E7idodontidae . 

-f  +A?nphidoxa  helicophantoides  Pfeifif. — Mt.  The  genus  (only  2  species  known) 
is  related  to  Stephanoda. 

+  A.  marmorella  Pfeiff. — Mt. 

+  Charopa  (Endodonta,  s.  lat.)  involuta  Odhner.— Mt.  Numerous  species, 
Polyn.,  N.  Guin.,  Austral.,  N.  Zeal. 

-f  C.  occulta  Odhner. — Mt. 

-f  C.  skottsbergi  Odhner. — Mf. 


3l6  C.   SKOTTSBERG 

^Pioictuni  conicion  Odhner. — Mt.  A  genus  of  a  rather  small  number  of  spe- 
cies, in  various  parts  of  the  world. 

^P.  depressu))!   Odhner. — Mt. 

^-  Radiodiscus  inasafucrae  ((Odhner)  Pilsbry. — Mf.  An  American  genus  of 
few  species,  found  in  X.  America  (Arizona)  and  in  S.  America,  south  to  Patagonia. 

■^StepJuDwda  iirctispira  PfeitT. — Mt.  About  30  species  in  western  S.  America 
(Chile,  south  to   Fuegia). 

+  .V.  ceratoides  Pfeiff.— Mt. 

+  .V.  quadrata   b'erussac. — Mt. 

+  5.  sclkirki  \\.  A.  Smith.— Mt. 

-S.  tcssellata  Muehlf.— Mt,   Mf. 

HilicJdae. 

Helix  aspcrsa  Muell.   Cosmopol.,  introduced  to  Chile. — Mt. 
Li))iacida€. 

Agrioliiuax  agrestis  L.  Very  widespread,  introduced  to  Chile. — Mt,  Mf. 

Liuiax  arhoruui  Buch. -Chant.  As  the  former. — Mt,  Mf. 

Mi/ax  gagatcs  Draparn.  As  the  former. — Mt,  Mf. 

Sitcciiicidae  (subfam.   Succineinae). 

+  Succiuea  cuuiingi  Reeve. — Mt.  Succhiea  is  taken  in  its  old  sense;  it  has 
been  split  up,  and  the  Juan  Fernandez  species  belong  to  a  group  which  must 
bear  a  diflerent  name.  It  is  reported  from  N.  America,  Panama,  Galap.  Is.,  St. 
Helena.  S.  Africa,   Hawaii  and  Tahiti. 

+  6.  fcr)ia7idi  Reeve  — Mt. 

.V.  fragilis  King.  (syn.   S.  texta  Odhner).  Hawaii. — Mt. 

+  .V.  gayajia  (D'Orbigny)  (Odhner. — Mt. 

+  .V.  Diasafucrae  Odhner. — Mf. 

+  .S".  pifigujs  (Pfeiff.)   Reeve. — Mf. 

+  .S".  seuiiglohosa  Pfeiff.— Mt,   SC. 

Tor)iatclli)iidae. 

^  +  I''ir?ia?idfzia  buliniojdes  Pfeiff.  (inch  consimilis  Reeve). — Mt. 
+ /'\  couifcra  Reeve. — Mt. 

.'  +  /'".    cyli)idrella    Odhner. — Mt.   Possibly  identical  with  Torvatellina  (F21asma- 
tina)  tunita  Anton,  credited  to  Oi)aru   I.* 
+  /'".  diapluDia  King. — Mt. 
+  /'\   expivisa   Pilsbrw — Mt. 
+ /'\   inorjiata   Pilsbry. — Mt. 
^  F.   lo)iga   Pilsbry. — Mt. 
+  /'".  pliilippiiUia   Pilsbrw — Mt. 
+  /'".  splendid  a  Anton. — .Mt. 
f/'\  tryojii  I'ilsbry. — Mt. 
+  /'\   /('ils()?ii  I'ilsJMy.— Mt. 

'   Rapa  seems  to  be  tlie  name  commonly  used. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  317 

+  Tornatellina  aperta  Odhner. — Mt.  A  Pacific  genus  of  over  50  species,  ranging 
from  E.  Ind.  and  Japan  over  Micron.,  Polyn.  (inch  Hawaii),  Melan.  (Kermadec 
Is.,  N.  Caled.)  to  Austral,  and  N.  Zeal. 

T.  bilamellata  Anton.  Recorded  from  Oparu  I. 

+  T.  callosa  Odhner. — Mt. 

+  T.  conica  Anton. — Mt. 

+  T.  plicosa  Odhner. — Mt. 

T.  reclusiaia  Petit. — Mf.  According  to  Odhner  probably  identical  with  T.  iiir- 
rita  Anton. 

+  T.  trochiformis  (Beck)  Pfeiff. — Mt. 

T.  trochlearis  (Beck)  Pfeiff.  Oparu  I. — Mt. 

+  Tornatellinops  minuta  (Anton)  Pilsbry  et  Cooke. — Mt.  A  Pacific  genus  of 
22  species,  reported  from  Japan,  Philipp.  Is.,  Polyn.  and  N.  Zeal. 

Zonitidae. 

Hyalinia  alliaria  Miller.  A  widely  distributed,  anthropochorous  species. — Mt. 
H.  cellaria  Miller.  As  the  former.— Mt.  Mf. 

Forty-six  species  are  enumerated;  of  these  6  have  been  introduced  through 
the  human  traffic.  Of  the  remaining  40,  35  (87.5%)  are  supposed  to  be  en- 
demic; 13  belong  to  the  two  endemic  genera.  The  occurrence  oi  Succinea  fragilis 
in  Hawaii  and  Juan  Fernandez  and  nowhere  else  in  surprising,  and  4  species 
are  credited  to  Oparu  (Rapa)  Island,  but  the  distribution  is  perhaps  not  too  well 
known.  The  poverty  of  Masafuera,  where  only  5  species  have  been  collected,  3 
of  them  restricted  to  this  island,  is,  I  daresay,  only  apparent.  Additional  forms 
have  been  found  later  and  still  await  study.  On  Santa  Clara  only  empty  shells 
of  a  Masatierran  species  were  found.  This  islet  seems  entirely  unfit  for  land-shells. 

Only  two  well-defined  geographical  groups  are  distinguished,  to  which  a 
third  of  wider  extension  is  added. 

I.  American  element. — 8   sp. 
Amphidoxa  (2),  Radiodiscus  (i),  Stephanoda  (5). 

II.  Pacific  element. — ^23   sp. 

Charopa  (3),  Fernandezia  (11),  TornatelHna  (8),  Tornatellinops  (i). 

III.  Austral  (or  more  widespread)   element. — 9  sp. 
Punctum  (2),  Succinea  (7,  see  above). 


Chapter  IV. 

Continental  and  Oceanic  islands. 

For  a  clear  distinction  between  the  two  main  kinds  of  islands  WALLACE  [2^8) 
is  as  a  rule  referred  to  as  the  leading  authority.  From  a  geographic-geological 
viewpoint  an  island,  usually  neovolcanic  or  coralline,  which  does  not  stand  upon 
a  continental    shelf,    is  called  oceanic.    If  situated  on  the  shelf  there  is  a  strong 


3l8  C.   SKOTTSBERG 

j)ossibility  that,  at  some  period  of  its  existence,  it  has  formed  part  of  the  con- 
tinent. This  is  e.g.  the  case  witli  the  Falkland  Islands.  A  truly  oceanic  island 
lacks  a  continental  basement  of  old,  granitic  or  sedimentary  rocks;  at  least,  their 
presence  has  not  been  demonstrated.  It  is,  on  all  sides,  surrounded  by  deep  water 
and  a  rise  of  perhaj:)s  thousands  of  metres  is  required  to  bring  it  into  contact 
with  a  continent.  Mavr  (/J^),  however,  argues  that,  from  a  biological  viewpoint, 
every  island,  whether  situated  on  a  continental  shelf  or  not,  is  oceanic  which  has 
received  its  entire  living  world  across  the  open  ocean — consequently  it  must  be 
shown  that  every  kind  of  organism  present  on  the  island  has  or  once  had  the 
faculty  of  migrating  across  the  sea  and  establishing  itself,  either  the  species  ac- 
tually found  or  their  ancestors. 

It  goes  without  saying  that  the  answer  to  the  question  "continental  or  oceanic?" 
should  in  the  first  place  be  looked  for  in  the  history  of  the  oceans.  With  regard 
to  the  Pacific  our  knowledge  of  its  origin  and  history  is  incomplete,  and  even 
if  modern  ()ceanograj)hical  research  has  supplied  a  wealth  of  information  on  the 
hydrography,  the  nature  of  the  sediments  and  so  on,  large  parts  of  the  southern 
Pacific  are  little  known  and  soundings  so  few  that  we  cannot  form  but  a  very 
general  idea  of  the  bathymetrical  conditions  and  the  configuration  of  the  bottom. 
The  northern  half  is  of  course  far  better  known.  As  it  is,  we  must  admit  that 
little  or  nothing  has  come  to  light  that  is  opposed  to  the  theory  of  the  perma- 
nence of  the  Pacific  Ocean.  It  is,  with  few  exceptions,  from  the  biologists'  camp 
that  the  theory  has  been  attacked,  particularly  by  phytogeographers;  the  majority 
of  zoologists  seem  to  accept  the  conclusions  arrived  at  by  physiographers  and 
geologists.  It  is  easy  to  understand,  however,  that  many  biogeographers,  struck 
by  the  [)er{)lexing  disjunctions  in  the  distribution  of  plants  and  animals,  started 
to  build  bridges  across  wide  expanses  of  sea,  in  cases  with  a  generosity  that  led 
to  absurdities.  I  have  no  reason  to  enter  into  details,  our  problem  concerns  Ant- 
arctica, southern  South  America  and  Juan  P>rnandez,  but  even  so  it  seems  worth 
while  to  (juote  a  number  of  modern  scientists,  mainly  geographers  and  geologists, 
who  have  expressed  their  opinion  on  the  nature  and  history  of  the  Pacific  Ocean. 

Geotectonics  of  the  Pacific  Basin. 

H.MI.F.V  Willis  (2S4)  thinks  that  a  suboceanic  pressure  works  against  the  con- 
tinents surrounding  the  Pacific,  resulting  in  an  expansion  of  the  suboceanic  mass 
and  a  deepening  of  the  basin  which,  in  its  turn,  has  a  displacing  effect  on  the 
continental  margin.   lie  summarizes  p.   367-368: 

The  consideration  of  the  general  facts  of  the  geotectonics  of  the  Pacific  basin  thus 
leads  us  to  regard  the  great  ocean  as  a  dynamic  realm,  within  which  the  peculiar  char- 
acteristics of  its  rocks  have  facilitated  the  internal  forces  of  the  earth.  The  effects  have 
i)een  as  a  whole  to  deepen  the  basin  in  conse([uence  of  the  expansion  of  the  under- 
lying ro(  ks.  The  expansion  has  in  turn  crushed  the  continental  margins  and  raised  the 
great  cordilleras.  Geologic  studies  of  the  mountain  ranges  have  demonstrated  that  the 
actual  orogenic  j)criod  began  in  the  Jurassic  or  possibly  somewhat  earlier  in  the  Mesozoic. 
Of  the  earlier  periods  we  know  but  little,  but  the  fragmentary  records  indicate  that 
})eriods  of  orogenic  activity  alternated  with  those  of  (piiescence. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  3I9 

Willis  did  not  question  the  permanence  of  the  basin;  what  interests  us  here 
more  particularly  is  his  belief  in  the  instability  of  the  marginal  regions. 

In  H.  E.  Gregory's  view  the  Pacific  basin  inside  the  deep  troughs  is  an  old 
sink  and  it  follows  that  all  the  islands  within  this  sink  are  truly  oceanic.  Con- 
sidering geological  evidence  alone  there  has  been  no  significant  change  in  the 
position  of  Polynesian  land  masses  since  Pleistocene,  most  likely  since  early  Ter- 
tiary time:  "There  is  no  geologic  evidence  that  any  Polynesian  island  stood  in 
Jurassic  or  Cretaceous  seas"  (775.  1673).  Still  he  thought  that  due  regard  should 
be  taken  to  objections  raised  by  other  branches  of  science,  and  he  did  not  extend 
the  unaltered  permanent  basin  outside  the  deep  troughs.  When,  a  little  later, 
another  prominent  geologist,  J.  W.  Gregory  (116),  expressed  a  different  opinion, 
this  attracted  a  good  deal  of  attention.  The  Pacific  had,  he  says,  been  claimed 
to  have  existed  in  its  present  shape  and  size  throughout  geological  time,  a  hypo- 
thesis almost  universally  adopted  by  geophysicists  and  geologists,  but  from 
a  biological  viewpoint  this  theory  did  not  satisfy.  GREGORY  was  no  believer  in 
large-scale  transmarine  migration  of  either  plants  or  animals  and  consequently 
inclined  to  consider  the  arguments  put  forth  by  the  opponents  to  the  permanence 
theory.  He  counted  with  a  number  of  Pacific  seas  separated  by  stretches  of  land, 
and  he  looked  upon  the  region  where  atolls  serve  as  proofs  of  subsidence,  a  sub- 
sidence which  gradually  enlarged  the  basin  until  it  reached  its  present  size,  as 
originally  continental. 

Andrews  [6)  who  was  a  firm  believer  in  successful  transoceanic  migrations 
of  all  kinds  of  organisms  and  knew  more  about  geology  than  most  biogeogra- 
phers,  shared  H.  E.  Gregory's  opinion:  islands  situated  within  the  area  bounded 
by  the  ocean  deeps  such  as  Hawaii,  Marquesas,  Society  and  Juan  Fernandez,  all 
differing  in  their  geological  structure  from  the  continents,  are  oceanic.  Parts 
of  Andrews'  interesting  paper  deserve  to  be  quoted  here. 

An  examination  of  the  continental  blocks  proper  and  the  great  western  island 
arcs  suggests  that  they  have  had  similar  histories,  whatever  great  differences  may  other- 
wise exist  between  them.  Japan,  Eastern  Australia,  New  Zealand,  New  Guinea,  Fiji  .  .  . 
may  be  taken  as  examples.  Each  has  a  foundation  of  ancient  folded  and  metamor- 
phosed sediments,  such  as  conglomerates,  grits,  quartzites,  sandstones,  slates,  shales, 
and  limestones,  and  each  of  these  foundations  has  been  subjected  to  marked  plutonic 
intrusions  of  granitoid  nature.  Upon  this  foundation  have  been  accumulated  sediments 
similar  to  those  mentioned  above,  together  with  lavas  not  only  of  basic  but  also  of 
acid  types.  These,  in  turn,  have  been  folded,  overthrust,  and  invaded  by  plutonic  rocks. 
This  generalization  is  true  even  though  as  yet  no  consensus  exists  concerning  the  age, 
or  ages,  of  the  folded  sediments  and  plutonic  intrusives  of  the  foundadon  rocks.  It 
would  appear,  however,  that  the  foundation  rocks  of  the  island  arcs  which  occur  mar- 
ginally to  the  continents  of  Asia  and  Australia  are  not  as  old  as  the  earliest  members 
of  the  continental  nuclei.  This  leads  to  the  consideration  of  island  arcs  situated  more 
centrally  within  the  Pacific  Ocean.  P'or  this  purpose,  these  may  be  considered  as  in- 
cluding all  the  Pacific  islands  lying  oceanward  of  the  great  island  arcs  mentioned  above. 
The  principal  examples  include  the  Hawaiian  Islands,  the  Marquesas,  Juan  Fernandez, 
Easter  Island,  the  Society  Islands,  the  Cook  Group,  the  Line  Group,  Micronesia,  Samoa, 
and  the  Ladrone,  Caroline,  and  Pellew  groups;  Tonga  and  the  Hebrides  occupy  a 
peculiar  position,  mentioned  below  (p.  202—3). 


320 


SKOTTSBERG 


Field  observations  show  that  those  islands  have  had  histories  which  present  marked 
diftercnces  from  those  of  the  continents  and  their  marginal  island  arcs.  'J'hus,  they 
appear  to  be  comjiosed  almost  entirely  of  volcanic  material,  mainly  basic,  together  with 
"coral  reef"  formations,  whereas  granitoid  intriisives  and  acid  lavas  are  lacking,  to- 
gether with  the  sediments  invarial)ly  associated  with  "continental"  areas.  And  not  only 
is  this  so,  but  the  volcanic  ejectamenta  of  these  inner  groups  do  not  appear  to  con- 
tain fragments  of  granitoids  and  sediments  such  as  might  be  expected  from  volcanos 
discharging  through  a  foundation  of  rock  formations  such  as  compose  the  continents 
(P-  203). 

Another  interesting  feature  is  the  ])eculiar  topograj)hy  of  that  portion  of  the  Pacific 
floor  which  sej)arates  the  great  island  arcs  and  lands  of  "continental''  character  from 
the  more  central  grouj)s.  Thus,  on  the  American  side  of  the  Pacific,  the  "continental" 
lands  are  separated  from  the  groups — such  as  Hawaii,  the  Marcjuesas,  the  Society  Is- 
lands, and  Juan  I'crnandez — by  a  series  of  deep  discontinuous  ocean  trenches,  prac- 
tically collinear  (j).  203). 

The  question  is  whether  these  trenches  are  of  quite  the  same  nature  and 
date  from  the  same  j^eriod  as  the  deeper  trenches  arranged  oceanward  from  the 
great  western  island  arcs.  It  is  unfortunate  for  the  advocates  of  a  "continental" 
origin  of  the  Juan  Fernandez  flora  that  these  islands  are  situated  on  the  wrong 
side  of  the  trench.  However,  the  Galapagos  Islands  occupy  a  similar  position, 
and  still  they  have  been  claimed,  on  good  grounds,  once  to  have  been  united 
with  Central  America.  In  this  connection  another  quotation  from  ANDREWS  with 
regard  to  the  New   Hebrides  and  Tonga  is  of  interest. 

Island  groups  which  are  difficult  to  place  exactly  in  this  scheme  are  the  New 
Hebrides,  Tonga,  and  possibly  the  Pellews  and  the  Ladrones.  A  profound  deep  lies 
between  New  Caledonia  and  the  New  Hebrides,  and  this  is  suggestive  of  a  noncon- 
tinental  origin  of  the  group.  On  the  other  hand,  the  occurrence  of  mineral  deposits 
such  as  copj)er,  iron,  and  nickel,  of  large  kauri,  fig,  myrtaceous,  and  other  trees,  and 
of  animals  such  as  lizards,  turtles,  ducks,  pigeons,  and  parrots,  suggests  that  they  may 
well  have  formed,  at  some  earlier  time,  j)ortions  of  a  continental  margin  which  later 
be<  amc  involved  in  a  j)Owerful  movement  within  the  marginal  Pacific,  resulting  in  the 
gradual  submergence  of  these  outer  ))ortions,  the  present  Hebrides,  Tonga,  and  so  on, 
being  built  upon  su(  h  sinking  area.  This  certainly  is  suggested  for  the  New  Hebrides 
and  for  Tonga,  while  the  Pellew  and  Ladrone  islands  also  have  had  complex  histories, 
which  would  well  repay  close  attention,  in  their  structural,  petrological  and  biological 
aspects   (p.  203). 

An  additional  j)C)int  of  interest  is  the  association  of  great  ocean  deeps  with  youthful 
volcanic  zones,  and  inasnuu  h  as  the  trough  and  crest  of  an  earth  undulation  are  parts 
of  the  same  strut  ture,  it  is  a  legitimate  inference  that  the  great  Pacific  deeps  or  trenches 
are   relatively   youthful   structures  (p.   203). 

Andkkw.s  summarized  his  idea  of  the  Pacific  basin  in  a  number  of  points, 
which,  with  very  slight  verbal  alterations,  form  the  introduction  to  his  paper  on 
the  origin   of  the   Pacific  insular  floras  (7.  613-14): 

1.  The  continents  bordering  the  Pacific  have  been  larger,  at  various  times,  than 
they   are   at   |)resent. 

2.  The  great  bordering  island  arcs  of  the  Pacific — such  as  the  Aleutians,  Japan, 
the  IMiilippines.  the  Netherlands  Fast  Indies,  New  (Guinea,  Fiji,  New  Caledonia,  and 
New  Zealand— have  been  connected  directly  with  the  continental  lands.  Certain  of  these 


DERIVATION  OF  THE  FLORA  AND  FAUNA  321 

island  groups — such  as  Fiji,  New  Caledonia,  and  New  Zealand — appear  to  have  been 
isolated  at  much  earlier  periods  than  others,  such  as  New  Guinea,  the  East  Indies, 
and  Japan. 

3.  The  ancient  borderlands  of  the  continents  have,  in  part  at  least,  suffered  un- 
dulatory  submergence.  Compensatory  forms  are  the  deep  ocean  trenches,  on  the  one 
hand,  and  the  mountain  ranges  of  the  continents,  on  the  other. 

4.  The  Pacific  is  a  relatively  deep  and  unstable  area,  whereas  the  Atlantic — with 
the  exception  of  the  broad  intersected  belt  of  activity  directed  toward  the  equator — 
has,  on  the  whole,  been  relatively  stable  since  the  Palaeozoic. 

5.  The  western  area  of  the  Pacific  appears  to  possess  a  more  complex  structure 
than  the  eastern,  owing  to  the  earth's  rotation,  the  width  and  weakness  of  the  Pacific 
base,  and  the  resistance  opposed  to  this  activity  by  the  stable  continental  masses  of 
Eurasia  and  the  Australian— Sahul  area. 

6.  The  islands  of  the  Pacific  lying  within  the  area  bounded  by  the  ocean  trenches 
have  not  had  continental  histories,  nor  do  they  appear  ever  to  have  had  actual  and 
direct  land  connections  with  the  continents. 

7.  The  New  Hebrides  and  Tongan  Islands,  and  possibly  also  the  Pellew  and  Mari- 
anne groups,  appear  to  partake  in  some  measure  of  the  nature  both  of  "continental" 
and  "oceanic"   islands. 

We  shall  have  occasion  to  return  to  Andrews'  opinion  on  the  origin  and  his- 
tory of  Pacific  floras.  Here  it  seems  convenient  to  draw  attention  to  the  numerous 
submarine  cones  called  "guyots"  recently  discovered  in  the  sea  between  Hawaii 
and  the  Marianas  and  discussed  by  Hess  (1J2).  About  160  flat-topped  peaks, 
presumably  truncated  volcanic  islands,  rise  from  9000  to  150CO  feet  above  the 
deap-sea  bottom.  In  most  cases  their  flattened  summit  was  sounded  in  about  800 
fathoms.  Hess'  working  theory  is  that  they  were  formed  on  land,  sunk  to  their 
present  level  and  levelled  by  sea  action — this  would  mean  that  they  stood  with 
their  summit  at  sea  level  long  enough  to  be  exposed  to  wave  action.  Thus, 
drowned  reefs  could  be  expected,  but  no  such  are  reported:  the  guyots  are, 
Hess  thinks,  very  ancient  structures  dating  from  a  "proterozoic  episode  of  vul- 
canism",  they  are  of  pre-Cambrian  age  and  consequently  do  not  lend  themselves 
to  biological  speculations.  There  may,  however,  have  existed  later  islands,  both 
Paleozoic  and  Mesozoic,  but  all  have  disappeared  beneath  the  surface  of  the  ocean, 
either  built  up  again  by  reef-building  organisms  or  sunk  to  a  depth  where  these 
cannot  live.  The  high  volcanic  islands  are  very  young,  perhaps  not  even  Tertiary 
but  Pleistocene  or  recent,  because  the  rocks  could  rarely  be  proved  to  be  of  Ter- 
tiary age.  We  shall  see  by  and  by  that  the  endemic  insular  floras  and  faunas  cannot 
be  anything  like  recent.  "Oceanic  islands",  Hess  continues,  "are  and  have  always 
been  slowly  sinking  relative  to  sea  level"  as  a  consequence  of  the  accumulation 
of  bottom  sediments  causing  the  water  level  to  rise.  The  red  clay  increases  i  cm 
in  loooo  years,  the  globigerina  ooze  the  same  amount  in  5000  years.  Thus  millions 
of  years  are  needed  to  account  for  even  a  very  moderate  submergence.  Besides, 
raised  shore-lines  are,  in  many  cases,  unmistakable  proofs  of  local  emergence. 
The  melting  of  the  great  inland  ices  ought  to  have  had  a  greater  influence. 

Finally,  let  us  listen  to  one  of  the  foremost  authorities  on  geophysics,  GUTEN- 
BERG (i2j).  He  finds  (p.  7)  that  there  is 

21  -557857    The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.    Vol.  I 


02  2  t'.  SKOTTSBERG 

growing  accumulation  of  evidence  that  the  Pacific  basin  shows  unique  features  which 
are  not  duplicated  in  any  other  oceanic  or  continental  area  of  the  earth.  'J'here  is  no 
feature  on  the  surface  of  the  earth  which  compares  in  dimensions  and  importance  with 
the  Marshall  line,  within  which  the  younger  eruptive  rocks  are  basaltic  rather  than 
andesistic.  This  discontinuity  in  the  material  of  the  crustal  layers  is  called  here  the 
boundarv   of  the    Pacitic    l>asin. 

This  is  in  conforniit}-  with  what  I  have  quoted  above  from  other  sources. 
However, 

certain  areas  of  the  I'acific  Ocean  (near  its  borders,  for  example),  at  least  part  of  the 
region  between  South  America  and  the  Easter  Island  rise,  or  between  the  Marianas 
and  the  Asiatic  continent,  show  indications  of  continental  layers.  For  the  latter,  petro- 
graphical   and   geophysical   evidence  agree. 

As  seen  on  the  niaj),  (il  TKXHKRC]  goes  a  good  way  beyond  the  Juan  Fer- 
nandez-San Anibrosio  rise,  but  the  blaster  Island  shield  which,  excepting  the  vi- 
cinity of  this  island,  is  covered  by  very  deep  water,  belongs  to  the  wide  basaltic 
centre,  where  continental  layers  are  lacking — in  contrast  to  the  Atlantic  where 
"granitic  la\-ers  of  the  continents  continue  far  out  under  the  bottom  .  .  .  probably 
at  least  some  continental  rocks  underlie  its  bottom  throughout  its  area". 

Turning  to  tiie  speculations  of  biologists  I  shall  quote  some  representatives 
from  the  two  op|)osite  camps.  Arldt  [6')  did  not  draw  his  conclusions  merely 
from  facts  of  distribution  but  compiled  a  wealth  of  geological,  palaeontological, 
bathymetrical  dates  and  so  on,  and  constructeci  a  series  of  maps  illustrating  the 
distribution  of  land  and  sea  through  earth's  history.  A  Cretaceous  Oceania  united 
South  America  with  Australia  +  New  Zealand,  it  disappeared  during  F^ogene  and 
left  the  west  coast  of  South  America  in  the  same  position  as  to-day.  F>om  what 
he  sa)-s  about  Juan  Fernandez  it  appears  that  he  regarded  these  islands  as  con- 
tinental 'see  below  p.  376),  while  still  admitting  the  possibility  of  oversea  migra- 
tion from  the  coast.  Camphkll  was  for  a  long  time  a  supporter  of  the  land-bridge 
theorv.  He  regarded  the  Hawaiian  Islands  as  formerly  much  larger  and  more  closely 
connected  with  land  masses  to  the  southwest,  having  become  isolated  during 
early  Tertiary  time  coincident  with  the  ui)lift  of  the  great  Cordilleras  (</./).  Later, 
when  discussing  the  Australasian  element  in  the  Hawaiian  flora  he  expresses  him- 
selt  very  positively:  "We  are  justified  in  assuming  the  former  existence  of  land- 
masses  of  consideral)le  size,  connecting  more  or  less  directly  both  Australia  and 
New  Zealand  with  Hawaii'  (./-r.  22i);  and  when,  for  the  third  time,  he  took  up 
the  history  of  the   Hawaiian    flora,   he  expressed   himself  as  follows  (46.  l8l): 

We  may  assume  that  the  Hawaiian  Arc  hij)elag(),  as  it  now  exists,  is  but  a  remnant  of 
a  much  larger  landmass  which  has  been  in  subsidence  for  a  long  period,  and  that  extensive 
subsidence  has  also  o(  (  urrcd  throughout  Polynesia,  and  to  a  lesser  degree  in  Australasia. 
One  argument  tor  this  assumjition  is  the  great  development  of  coral  reefs  in  the  Pacific, 
espec  iaily  in  Polvuesia  and  northeastern  Australia.  The  existence  of  active  coral  reefs 
involves  continuous  subsidcMKc  and  the  absence  of  large  land-masses  in  mid-Pacific,  with 
the  innumerable  small  coral  islands  and  reefs,  can  be  explained  most  satisfactorily  on  the 
theory  that  the  latter  are  remnant  of  sul)mergcd  land-masses  of  large  size — possible  even 
of  continental  dimensions. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  323 

It  is  surprising  that,  fourteen  years  later,  he  vvas  converted  to  wegenerianism 
in  the  form  modified  by  Du  ToiT,  thinking  that  "this  theory  would  best  explain 
most  of  the  problems  in  the  geographical  distribution  of  the  floras  of  the  Southern 
Hemisphere"  (^7.70) — he  seems  to  have  forgotten  how  badly  Hawaii  fits  into 
the  picture. 

GuiLLAUMiN,  in  his  paper  on  the  floristic  regions  of  the  Pacific  {iiS),  devotes 
a  chapter  to  its  geotectonic  history. 

Un  vaste  continent  parallele  a  I'equateur  a  du  relier  les  regions  australiens,  canaque, 
et  neozelandaise  a  I'Amerique  du  Sud  tandis  que  la  region  malayo-polynesienne  et  le 
domain  hawaiien,   formant  peut-etre  un  continent,  en  etaient  separes. 

The  dissolution  and  disappearance  of  this  hypothetical  continent  took  place 
before  the  end  of  the  Jurassic,  so  that  its  direct  influence  on  the  distribution  of 
angiosperms  must  have  been  slight;  Arldt's  "Ozeanis"  was  more  useful.  The 
Melanesian  extension  of  Australia-New  Zealand,  including  Fiji,  Kermadcc,  Tonga 
etc.,  persisted  longer  and  is  perhaps  better  founded,  but  when  GuiLLAUMiN  thinks 
that  his  hypotheses  "ne  sont  pas  en  contradiction  avec  les  donnees  geologiques 
ni  avec  ce  qu'on  salt  du  relief  sousmarin  du  Pacific",  he  moves  on  unsafe  ground. 
Most  authors  who  deny  any  considerable  reduction  of  the  surface  of  the  Pacific 
admit  that  Melanesia  forms  an  exception.  Thus  Guppy  (121)  who  otherwise  is  a 
firm  believer  in  oversea  migration  thinks  that  "we  should  rope  in  Fiji  with  all 
the  large  islands  westward  and  southward  as  originally  continental"  because  they 
lie  inside  the  gymnosperm  line.  All  non-coniferous  ones  are  excluded.  Germain 
(/05. 973)  goes  much  farther.  Discussing  the  Hawaiian  region  he  thinks  that  it 
may  have  extended  far  eastward.  The  distribution  of  some  animals,  for  instance 
the  eel,  proves  that  Hawaii  vvas  "part  of  an  oriental  Pacific  continent,  perhaps 
also  united  with  the  Juan  Fernandez  and  Easter  islands",  and  he  suggests  that 
many  of  the  Polynesian  elements  in  Hawaii  travelled  by  way  of  South  America 
and  Juan  Fernandez  (p.  1009). 

Among  botanists  Setchell,  pointing  to  the  evidence  furnished  by  geology, 
expresses  his  opinion  on  Pacific  paleogeography  in   the  followings  words. 

I  see  no  necessity  of  postulating  any  fundamental  changes  from  the  point  of  view 
of  the  permanence  of  the  Pacific  Ocean  as  such,  and  the  purely  volcanic  origin,  prob- 
ably in  Tertiary  times,  of  the  islands  existing  in  it,  in  much  the  same  position  as  we 
now  find  them  (279.301). 

Another  prominent  botanist  who  has  strong  claims  to  the  title  "Defender  of 
the  Oceanic  Faith",  is  Fosberg.  He  refuses  to  recognize  any  Pacific  islands  at 
all  as  continental,  even  Fiji  (pc?.  164  etc.);  though  the  flora  is  "plutot  denature 
continentale"  it  is  not  necessary  to  count  with  terrestrial  connections  to  explain 
its  characteristics.  Among  zoologists,  Mayr  (779)  and  Zimmerman  (2^8)  belong  to  the 
same  camp,  and  so  do  most  of  them,  but  it  happens  that  certain  animal  groups^ 
particularly  the  land  molluscs,  offer  serious  difficulties  and  have  led  otherwise 
conservative  zoogeographers  to  take  refuge  in  "a  mid-Pacific  land".  So  for  in- 
stance MUMFORD  (i8j.  247) : 


324  C.  SKOTTSBKRG 

Kvideiue  with  regard  to  the  nature  and  distribution  of  land  snails  is  of  the  utmost 
imi)ortance,  for  it  is  from  the  nature  of  the  land-snail  fauna  in  Polynesia  that  Mr.  Pilshry 
has  rea(  hed  the  conclusion  that  the  whole  of  Polynesia,  with  Hawaii,  was  once  a  great 
continental  land  mass.  l\utida- — an  ancient  generalized  type  of  land  snail — is  wide- 
spread in  Polynesia  and  Melanesia  and  not  elsewhere ;  whereas  many  groups  (e.g.  He- 
li(  idae  and  Arionidae)  found  widely  sj)read  in  the  world  are  absent  in  Polynesia,  etc. 
Professor  Buxton  is  of  the  opinion  that  Pilsbry  has  been  more  successful  than  any  other 
writer  in   establishing  a  case   for  the  early   existence   of  a   mid-Pacific  continent. 

Hut  how  early  or  how  late.^  To  explain  the  absence  of  modern  land  molluscs 
we  may  have  to  j^o  back  to  late  Tertiar)'  times  only,  and  we  should  need  other 
proofs  of  a  distribution  of  land  and  sea  sufficiently  different  from  the  present  one; 
perhaps  the\-  will  be  found.  Ikit  we  cannot  simply  fill  out  the  Pacific  basin  with 
land  and  leave  the  surrounding  continents  unaltered.  Often  enough  due  regard 
was  not  taken  to  such  circumstances.  SwEZKV  (^^2),  who  for  his  own  part  believed 
that  the  entire  insect  fauna  of  Mawaii  owed  its  presence  to  accidental  migration, 
(juotes  two  of  the  authors  of  "P^auna  Mawaiiensis"  who  did,  in  his  opinion  with- 
out an\-  reason,  build  bridges  where  they^  found  that  they  needed  them,  Meyrick 
and  Lord  \\'ALSiN(;iiAM.  Mkvrkk,  an  authority  on  Microlepidoptera,  when  stating 
that  among  the  endemic  Ilaw^aiian  genera  three  were  of  south  Pacific  affinity,  pos- 
tulated "the  former  existence  of  a  considerable  land  area  (now  submerged)  be- 
Iwcen  New  Zealand  and  South  America",  a  land  mass  which  compares  to  Arldt's 
South  Pacific  bridge— such  a  land  still  exists  and  is  not  submerged :  Antarctica! 
—and  he  also  believed  in  a  "Palaeonesia"  extending  from  Rapa  to  Hawaii  and 
from  Pitcairn  Island  to  the  Society  and  Cook  groups.  Little  room  is  left  for  the 
water  of  the  ocean,  but  he  does  not  argue  that  these  land  masses  were  contem- 
j)()rancous.  Lord  \\'.\i,siN(;iiAM  who  based  his  opinion  on  the  distribution  and 
relationships  of  the  Microlepidoptera  regarded  the  Hawaiian  Islands  as  representing 
the  summits  of  mountain  ranges  formerly^  belonging  to  a  continent,  "a  lost  Paci- 
fica";  if  not  accepted,  "some  other  theory  possibly  even  less  acceptable  must  be 
devised   — e(iuall\'   be)'()n(l  the  possibility  of  exact  j)roof 

It  is  cas\-  to  understand  that,  in  all  these  discussions  and  speculations,  Hawaii 
in  its  isolated  j)osition  and  with  its  rich  flora  and  fauna  should  be  the  object  of 
the  main  interest,  and  I  shall  end  this  chapter  with  a  review  of  PL.  H.  l^RVAN's  recent 
contribution,  "'Phe  Hawaiian  Chain"  [40),  ecpially  instructive  as  popularly  written. 
Hkn  AN  believes  in  a  continental  Melanesia  but  regards  Hawaii  as  oceanic,  but 
this  does  not  prevent  him  from  accepting  the  Leeward  islands  as  remnant  of  a 
long,  deeply  submerged  ridge,  nor  from  admitting  that  the  Hawaiian  Islands  proper 
may  have  been  greater  and,  in  part  at  least,  united.  The  chain  is  supposed  to 
have  emerged  first  at  its  extreme  western  end,  where  we  now  only  find  the  rem- 
nants of  once  larger  islands;  it  a|)peared  some  time  during  Tertiary,  the  forma- 
tion proceetling  toward  the  east,  with  the  island  of  Hawaii,  where  ejection  of 
lava  still  occurs,  as  the  youngest  link,  the  chain  having  been  completed  by  the 
end  of  the  Pliocene.  PRVAN,  who  has  a  wide  knowledge  of  the  Hawaiian  fauna,  is 
no  friend  of  land  bridges  and  less  so  of  submerged  continental  masses,  but  un- 
like many  other  zoologists  he  admits  that  it  is  very  difficult  to  explain  the  fauna 


DERIVATION  OF  THE  FLORA  AND  FAUNA  325 

without  the  help  of  some  kind  of  closer  contact  with  other  lands.  He  looks 
around  for  bridges  in  the  shape  of  "stepping  stones",  just  as  Mayr  does,  "tem- 
porarily connected  or  sufficiently  close  together".  "It  is,"  he  continues,  "a  long 
distance  between  Guam  and  Fiji  and  Tahiti  and  Hawaii,  but  if  there  were  numer- 
ous other  islands  spread  conveniently  between  .  .  .";  speaking  of  the  weevils  of 
Necker  Island  and  of  Nihoa,  Wake  and  Laysan  where,  with  the  exception  of  Nihoa 
with  its  endemic  palm  and  Laysan,  once  the  home  of  an  endemic  form  of  San- 
tahun,  no  suitable  host  plants  exist,  he  finds  strong  evidence  for  their  represent- 
ing "the  last  remnants  of  former  forest  insects,  surviving  along  a  route  of  migra- 
tion, a  land  bridge  of  the  past" — this  applies,  I  daresay,  to  an  earlier  connection 
between  the  links  of  the  broken  Hawaiian  chain.  However,  it  seems  to  me  that 
rows  of  "conveniently  spaced  stepping  stones",  sufficient  to  offer  routes  of  migra- 
tion from  several  directions,  involve  tectonic  movements  of  considerable  magnitude. 
Few  conscientious  bridge-builders  would  argue  that,  for  instance,  a  solid  land 
mass  extended  from  Melanesia  and  Indonesia  to  Hawaii  as  a  continuous  open 
road,  it  might  have  risen  gradually  from  west  to  east — when  the  land  upon 
which  the  present  Hawaiian  Islands  were  built,  was  above  the  sea,  the  western 
part  of  the  bridge  had  disappeared;  all  that  was  left  was  a  detached,  "advanced" 
portion  of  a  borderland,  the  home  of  a  facies  of  the  Australian-Malaysian  fauna 
and  flora,  which  gradually  took  possession  of  the  rising  volcanic  soil  of  Hawaii. 


Chapter  V. 

The  Pacific  Ocean  and  Continental  Drift. 

It  serves  no  purpose  to  dwell  here  at  any  length  on  Wegener's  original 
hypothesis,  with  which  every  biogeographer  is  familiar,  but  it  may  be  useful  to 
scrutinize  its  bearing  on  Pacific  problems  in  general  and  Juan  Fernandez  in  par- 
ticular. Before  the  breaking  up  of  Pangaea,  the  Pacific  Ocean  was  twice  as  wide 
as  now,  an  enormous  water  desert  where  no  islands  enlivened  the  seascape.  The 
entire  sial  crust  revolved  west,  where  festoons  were  successively  split  off  from 
the  Asiatic-Australian  land  mass,  got  stuck  in  the  sima  forming  one  island  arc 
after  the  other,  bordered  on  their  outside  by  deep  trenches.  The  Americas  trav- 
elled at  a  greater  speed  away  from  Europe-Africa,  and  the  Pacific  became  less  wide. 
It  is  true  that  most  of  the  island  chains  in  the  Pacific  trend  NW-SE,  but  there 
are  many  islands  that  do  not  follow  this  pattern,  among  them  Juan  Fernandez; 
nevertheless,  even  if  he  did  not  expressly  state  this,  it  seems  clear  that  WEGENER 
regarded  all  of  them  to  be  of  the  same  origin.  In  an  earlier  paper  [2J1)  I  re- 
ferred briefly  to  what  he  said  about  Juan  Fernandez;  here  I  shall  quote  him  in  full 
{280.  116). 

Die  pazifischen  Inseln  (mitsamt  ihrem  submarinen  Unterbau)  warden  in  der  Ver- 
schiebungstheorie  als  von  den  Kontinentalschollen  abgeloste  Randketten  betrachtet,  die 
bei  der  allgemeinen,  vorwiegend  westlich  gerichteten  Bewegung  der  Erdkruste  liber  den 
Kern  allmahlich  nach  Osten  zuriickgeblieben  sind.  Ihre  Heimat  ware  hiernach,  ohne  auf 


326 


C.   SKOTTSBERG 


F^inzelheiten  einzugehen,  aiif  der  asiatischen  Seile  des  Ozeans  zu  siichen,  der  sie  jedenfalls 
in  den  betrachtcten  geologischen  Zeitcn  erheblich  niiher  als  heute  gelegen  hal)en  miis- 
sen.  Die  biologischen  Verhaltnisse  scheinen  dies  zu  l)estiitigen.  So  haben  nach  G  rise- 
bach  iind  I)  rude  die  Hawaiinseln  eine  Flora,  die  am  niichsten  verwandt  nicht  mit 
Nordamerika  ist,  das  ihnen  doch  am  niichsten  liegt,  und  von  dem  heute  Lult-  und 
Meeresstromung  herkommen,  sondern  mit  der  alten  Welt.  Die  Insel  Juan  Fernandez 
zeigt  nach  Skottsl)erg  gar  kcine  X'erwandtschaft  mit  der  doch  so  nahen  Kiiste  von 
('hile,  sondern  mit  Feuerhmd,  Antarktika,  Xeuseehind  und  den  anderen  pazifischen 
Insehi.  Doch  sei  hcrvorgehoben,  dass  die  biologischen  Verhaltnisse  auf  Inseln  allge- 
mein   schwerer  zu   dcuten   sintl  als   diejenigen   auf  grosseren   Landraumen. 

In  the  3r(l  edition  j).  59  we  read  after  "Jnseln":  "Dies  passt  vorziiglich  zu  unserer 
Vorstelluni^,  class  Siidanicrika,  nach  Westen  wandernd,  sich  ihr  erst  in  letzter 
Zcit  so  wcit  i^enahert  bat,  dass  der  Morenunterschied  auffallend  wird."  In  the 
following  editions  this  sentence  was  excluded. 

Certainly  I  never  said  anything  like  that  and  I  fail  to  see  where  Wegp:ner 
got  bis  strange  ideas;  just  as  many  other  writers  I  have  pointed  out  that  the  Andean- 
Chilean  element  is  stronger  than  any  other.  F.vcn  to  a  firm  believer  in  the  festoon 
theory  the  Juan  FYTnandez  and  Desventuradas  Islands  ought  to  ofifer  insuperable 
difficulties.  \\'K(;i:m:r  built  his  theory  on  the  island  arcs  accompanying  the  Asiatic- 
Australian  continental  border;  geologically  these  arcs  are  continental,  but  when 
he  came  to  island  chains  like  Hawaii,  the  Marshall  Islands  and  the  Society  Islands 
— and  we  can  add  Marquesas,  Tuamotu  etc. — all  of  which  are  situated  outside 
the  decj)  trenches,  neovolcanic  and  regarded  as  built  up  from  the  depths  of 
the  ocean  "he  was  driven  to  assume  that  they  have  a  sialic  basement  hidden 
under  tlie  basaltic  layers.  He  thinks  that  this  assumption  is  supported  by  pend- 
ulum observations,  the  force  of  gravitation  being  greater  over  the  islands  than 
over  the  open  ocean  where,  of  course,  a  sial  cover  is  incompatible  with  his  dis- 
j^lacement  theory. 

\\'K(;i:M.k's  theories  were  taken  up  by  Dr  Toil'  and  presented  in  a  modi- 
fied   form   ((S7);    I   shall  (juote  his  attitude  toward   the  festoons. 

As  Wj'.cini.r  has  observed,  they  are  all  comj)arable  in  size,  regular,  linked  to- 
gether en  (,(  helon  and  convex  to  the  Pacific;  each  shuts  off  a  large  ])ortion  of  sea  and 
tronts  an  ex  eani<  deep,  while  the  concave  side  bears  a  row  of  volcanoes.  To  Suess  we 
owe  the  (onception  of  the  deve]oi)ment  of  successive  arcuate  asymmetrical  fold-waves  mi- 
grating outwards  from  the  more  stable  "Amphitheatre  of  Irkutsk",  which  led  to  ])rogressive 
e\i)ansion  of  .Asia  towards  the  Tac  ific.  While  the  hypothesis  has  since  had  to  be  ap- 
jircc  iatcl\  modihef],  its  fundamental  ideas  have  been  brilliantly  confirmed  by  subsequent 
investigations...  Significant  are  the  (x^eanic  fossae  that  immediately  front  the  convex 
sides  of  the  arcs — foredeeps  subsiding  in  nih'ance  of  the  outward-moving  geoantoclines 
and    incidentally    tracts   ot    marked    coastal    instability    (pp.    186-187). 

How  tar  (lid  this  outward  movement,  these  advance-folds  proceed?  Does  Du 
Toil  allow  all  the  Pacific  island  chains  to  be  linked  up  here?  When  the  great 
W'NW  swing  of  Asia  is  rcj)laced  by  an  expansion  toward  the  Pacific,  the  system 
of  rifts  in  the  ocean  floor,  oxer  which  the  island  chains  as  claimed  by  most 
geologists  were  formed,  did  not  exist,  because  there  was  no  tension  to  account 
for  them;   instead,  series  of  ri|>ples  were  crumpled  up  on  the  floor.  The  transfor- 


DERIVATION  OF  THE  FLORA  AND  FAUNA  327 

mation  of  the  geanticline  structures  into  rows  of  islands  is  explained  by  Du  ToiT 
as  follows;  their  crest 

could  become  deepened  by  crustal  tension  and  broken  into  segments  to  form  an  island 
chain  before  vanishing  .  .  .  limbs  were  intermittently  built  up  and  destroyed  during  the 
Cretaceous-Tertiary  through  stretching  in  the  direction  of  their  length  while  they  were 
still  compressed  by   forces  at  right  angles  thereto  (p.   293). 

Trying  to  apply  these  ideas  to  South  America,  which  according  to  Du  ToiT 
as  well  as  WEGENER  was  pressing  into  the  Pacific  basin,  the  Juan  Fernandez- 
Desventuradas-Merriam  ridge  could  be  compared  to  an  advance-fold.  But  it  is  not 
convex  to  the  Pacific,  nor  fronted  by  a  fossa — this  is  situated  on  the  wrong 
side  and  may  well  stand  in  causal  relation  to  the  upheaval  of  the  Cordillera.  To 
think  that  the  submarine  ridge  emerging  in  the  Juan  Fernandez  and  Desventu- 
radas  Islands  is  the  easternmost  advance-wave  from  a  western  borderland  seems 
too  phantastic. 

Du  Toit's  idea  of  the  geological  character  of  the  ocean  floor  differs  from 
Wegener's.  Seismographic  records,  he  says,  scarcely  bear  out  that  the  Pacific 
floor  must  be  composed  of  basalt — the  records  could  readily  accord  with  a 
granitic  layer  up  to  about  10  km  thick  (p.  212).  He  was  no  believer  in  a  more 
or  less  unlimited  oversea  migration  of  plants  and  animals,  nor  in  land-bridges, 
and  he  critizises  J.  W.  GREGORY  and  the  bridge-constructing  biogeographers:  they 
are  wrong,  and  the  displacement  hypothesis  interprets  otherwise.  But  when  he 
speaks  of  the  extensive  "march  into  the  ocean  of  crustal  waves,  thereby  leaving 
their  parent  continents  far  in  the  rear"  and  of  the  "rhythmic  intrusion,  culminat- 
ing in  the  three  migrations  of  the  Cretaceo-Eocene,  mid-Tertiary  and  late  Ter- 
tiary" (p.  214),  these  advance-folds,  when  crumpled  up  from  the  ocean  floor,  were 
absolutely  devoid  of  every  sign  of  terrestrial  life  and  without  a  trace  left  of  the 
sial  cover.  I  fail  to  see  that  they  can  solve  any  biogeographical  problems — we 
have  to  fall  back  on  oversea  dispersal.  Wegener's  festoons  were  at  least  split 
off  from  the  borders  of  a  continent  and  left  behind  with  their  fauna  and  flora.  With 
regard  to  Juan  Fernandez  we  shall  perhaps  be  able  to  find  a  less  adventurous 
explanation  of  its  history. 

Two  years  after  the  appearance  of  his  book,  Du  ToiT  summarized  his  theo- 
ries in  a  paper  which  I  think  it  is  worth  while  to  quote  [82.  75-76).  The  base- 
ment of  the  Melanesian  islands  is,  he  says,  for  the  most  part  continental;  the 
ocean  floor  consists  of  a  relatively  thin  structure  of  sial  underlain  by  sima,  but 
this  does  not  allow  us  to  regard  the  sial  as  continental,  because  it  may  be  a 
product  of  magmatic  differentiation  from  the  sima.  He  points  to  the  parallelism 
between  the  great  Tertiary  folding-zones,  most  evident  along  the  west  coast  of 
the  Americas,  and  the  trend  of  the  coast  line,  and  he  thinks  that  the  "compres- 
sive phases"  were  contemporaneous  all  around  from  New  Zealand  across  Antarc- 
tica to  South  America.  Coming  back  on  the  advance-folds  he  remarks  that  some 
of  Gregory's  hypothetical  bridges  or  land-masses  could  well  have  been  of  this 
nature.  The  procedure  is  illustrated  by  a  map  showing  the  pressure  direction 
and  the  formation  of  island  arcs — except  on  the  American  side,  where  the  sea 
is  a  blank. 


328  C.   SKOTTSBERG 

Xumerous  biologists  have  found  a  solution  of  all  or  most  of  their  difficulties 
to  explain  the  present  distribution  of  the  organic  world  in  the  theory  of  conti- 
nental drift,  combined  with  large-scale  pole-wanderings.  In  view  of  this  the  opi- 
nions expressed  by  modern  geologists  and  physiographers  cannot  be  passed  in 
silence.  A  symi)osium,  arranged  in  1950  (dj)  offers  an  opportunity  to  get  ac- 
quainted with   their  attitude. 

J.   II.  T.   L'MlKiKOXK,    TJic  case  for  the  cvust-substratiiui  theory,  pp.  67-71. 

Si  Kss'  terms  sial  and  sima  were  petrographic.  WEGENER  attributed  different 
physical  properties  to  these  types  of  rock;  sial  should  be  rigid  but  elastic,  sima 
viscous.  These  statements  lack  foundation.  Sima  (basalt)  has  a  higher  melting- 
point,  a})[)roximately  I300°C,  sial  (granites)  approximately  700°C.  The  crystal- 
line crust  of  the  sima  layer  is  at  least  as  strong  as  the  continental  sial.  Still  the 
sial  blocks  were  supposed  to  advance  through  the  sima.  Umbgrove  concludes 
that  continental  drift  is  impossible  at  present.  This  granted,  was  it  perhaps  pos- 
sible in  bygone  times.'  The  answer  is  fetched  from  the  Atlantic  and  Indian  oceans 
with  the  intervening  African  continent;  if  the  floor  of  the  oceans  originated  as 
thought  Wegener,  the  processes  must  have  taken  place  during  early  Precambrian. 
U.Ml'.fiKovE  asks  if  not  the  thick  blanket  of  sediments  would  have  been  squeezed 
and  [)iled  up  in  front  when  America  ploughed  westward.  He  calculates  that, 
considering  the  size  of  the  westward  drift,  a  plateau  200  km  wide  at  sea  level 
would  have  been  formed  in  front;  instead,  "the  continental  slope  is  one  of  the 
steepest  in  the  world  and  is  fronted  by  deep-sea  troughs" — here  we  have,  how- 
ever, to  consider  the  late  upheaval  of  the  Andes.  And  if,  as  Wegener's  theory 
recjuires,  the  Atlantic  originated  in  comparatively  recent  times,  how  are  we  to 
explain  the  enormous  thickness  of  its  bottom  sediments,  according  to  Hans  Pet- 
ri'.KSsoN  a  maximum  of  loooo  feet,  "representing  a  time-span  of  300  to  400 
million  years",  which  would  bring  us  back  to  the  Palaeozoic.  If  continental  drift 
e\er  occurred,  UMliGROVE  asserts,  it  took  place  some  3000  million  years  ago  and 
C()nsc(iuently  loses  every  shade  of  interest  to  the  biogeographer. 

HakoM)  Ji-.iikess,   Mechanical  aspects  of  continental  drift  and  alteriiatii'e  tJieories. 

ji;ill<i:\  s  definitely  rejects  We(;ener's  theory  on  geophysical  grounds; 
W  l.GlNl.K  gave  to  the  sima  properties  which  the  material  has  not;  basic  rocks 
are  stronger  than  acidic,  sima  stronger  than  sial.  The  strength  of  the  ocean  floor 
must  be  overcome,  if  drift  shall  result.  "I  seriously  suggest",  he  concludes,  "that 
no  more  time  be  spent  on  discussion  of  this  theory  until  a  mechanism  for  it  is 
produced;  what  it  has  done,  and  continues  to  do,  is  to  distract  attention  from 
the  serious   problems   of  geophysics"   (p.    80). 

S.    \\  .    W'ooi.iiKiix;!;,     TJu-    hearing   of  Late-tertiary  history  on  vertical  and  Jiori- 
zo}ital  ino/'cinoits  of  the  co)iti?ients. 

The  nature  ot  the  incchanisni  of  the  "uplift"  may  remain  in  doubt;  its  reality  can 
hardly  he  gainsaid  .  .  .  in  so  tar  as  such  comj)lementary  foundering  took  place  during 
Tertiary  times,  it  is  not  directly  relevant  with  the  drift  problem  .  .  .  Vertical  movements 
otter  an   alternative   solution   to   some   of  the  biological   problems.   Biological    evidences 


DERIVATION  OF  THE  FLORA  AND  FAUNA  329 

of  former  facilities  for  exchange  of  biota  are  in  themselves  neutral  in  deciding  which 
alternative  should  be  preferred  without  paying  attention  to  the  ambiguity  of  much  of 
the  geological  evidence  and  the  grave  geophysical  difficulties  (p.   8i). 

The  progress  of  knowledge  of  geophysics  certainly  seems  to  point  to  thinking  in 
terms  of  land  bridges  and  their  coastal  margins. 

The  later  findings  of  seismology,  indicating  three  distinct  rock-shells  each  prob- 
ably capable  of  both  vitreous  and  crystalline  states,  carry  the  possibility,  as  Jeff'reys 
has  clearly  insisted,  of  vertical  movements  on  a  major  scale.  Such  strong  vertical  move- 
ments are  indeed  not  merely  to  be  expected;  they  are  widely  and  unargubly  evidenced 
by    the    facts  of  geomorphology  no  less  than  those  of  stratigraphical  geology  (p.   82). 

R.  F.  Joyce,    The  relation  of  the  Scotia  Arc  to  Pangaea,  pp.   82-88. 

Joyce  made  an  attempt  to  reconstruct  Wegener's  Pangaea  in  Lower  Palaeo- 
zoic time.  To  him  continental  drift  is  a  possibility  only,  he  says  that  if  Pangaea 
did  in  fact  exist,  his  arrangement  of  the  actual  continents  and  islands — as  usual, 
the  Pacific  island  world  is  not  involved — at  the  opening  of  the  Palaeozoic  era  is 
more    in    accord  with  the  known  data  than  in  Wegener's  reconstruction  (p.  87). 

Two  biogeographers  took  part  in  the  symposium. 

H.  E.   HiNTON,    The    Wegener-Du   Toit  theory  of  conti^iental  displacement  and  the 

distribution  of  animals,  pp.  74-79. 

HiNTON  rejects  the  liberal  construction  of  bridges  to  suit  the  demands  of 
specialists.  Beside  the  usual  apprehensions  he  adds  the  following. 

A  further  objection  to  the  past  existence  of  transoceanic  bridges  ...  is  the  nearly 
complete  absence  of  deep-sea  deposits  on  the  existing  continents,  since  we  would  expect 
some  of  the  latter  to  have  been  also  elevated  from  the  sea  floor  (p.  75). 

This  seems  reasonable;  nobody  earnestly  proposes  to  fill  the  surface  of  the 
globe  with  land,  leaving  no  or  little  space  for  the  water.  Bridges  of  any  consider- 
able size  cannot  have  been  contemporaneous,  and  transgression  on  part  of  what 
is  now  land  seems  inevitable,  if  tracts  of  ocean  floor  were  exposed.  We  have 
ample  proofs  that  considerable  transgressions  occurred,  but  these  ancient  seas  may 
have  been  too  shallow  to  be  of  much  importance  as  a  support  of  the  bridge 
hypothesis.  HiNTON  thinks  that  we  require  relatively  few  inter-continental  con- 
nections, the  most  important  being  Brazil-Africa  and  Australia-Antarctica,  but  not 
in  the  form  of  bridges— of  such,  he  admits  isthmian  and  shelf-bridges,  nothing 
more.  Thus,  for  those  who  insist  on  direct  inter-continental  contact,  the  only 
hypothesis  at  their  service  is  the  drift  theory,  and  he  asserts  that  most  of  the 
modern  biogeographers  accept  the  broad  outlines  of  this  theory,  unless  it  is  claimed 
that  sliding  occurred  long  before  the  end  of  the  Mesozoic.  As  we  have  seen, 
however,  this  is  exactly  what  has  been  claimed.  For  his  own  part  he  puts  much 
faith  in  chance  dispersal   across  broad  stretches  of  open  water. 

R.  Good,    The  distribution  of  the  Flowering  Plants  in  relation  to  theories  of  con- 
tinental drift. 
After  some  remarks  on  the  general  distribution  of  plant  families,  grasses  and 

Compositae  taking  the  lead  almost  everywhere — which  seems  quite  natural  in  view 


330 


C.   SKOTTSBERG 


of  the  size  and  enormous  climatic  amplitude  of  these  families — the  preponder- 
ance, in  nearly  all  floras  of  dicotyledons  (2  V2  to  4  times  as  many  as  the  mono- 
cots),  and  the  absence  of  an\'  stron<^ly  marked  South  Pacific  flora  south  of  Hawaii 
and  east  of  Xew  (Guinea,  New  Caledonia  and  New  Zealand  (which  ought  to  show, 
I  suppose,  that  the  island  swarms  are  not  fragments  of  a  larger  land  mass),  GooD 
continues  j).   74: 

A  coiiimon  expression  of  the  theory  of  continental  drift  postulates  that  the  sunder- 
int;  of  various  coiiiinental  masses  began  in  a  relatively  remote  geological  period  and 
that  it  iuis  continued  without  notable  cessation  ever  since.  The  distribution  of  Angio- 
sperms  does  not  seem  to  be  in  special  accordance  with  any  such  particular  course  of 
events.  It  suggests  more  strongly  that  sundering  occurred  after  these  plants  as  a  group 
had  become  well  diffused,  let  us  say  by  the  Eocene.  Similarly  I  know  of  nothing  in 
the  distribution  of  Angiosperms  to  show  that  drift  is  still  going  on.  Can  it  be  that 
continental  drift  has  in  fact  been,  not  a  persistent  process,  but  an  intermittent  feature 
of  geological  time,  as  for  instance  have  the  great  glaciations?  If  so,  and  if  it  can 
further  be  shown  that  the  most  recent  of  these  drift  ages  took  place  not  earlier  than 
the  beginning  of  the  Tertiary,  then  at  last  the  plantgeographers  will  have  to  hand  a 
ma^tcr-kcy  to   most  of  his  perplexities. 

It  will  be  difficult  to  shape  a  key  that  will  fit  a  lock  constructed  in  this  fashion. 
As  the  glacial  periods  were  interrupted  by  interglacial  periods,  so  the  periods 
of  sundering  should  have  been  interrupted  by  periods  not  of  a  standstill  but  of 
the  land-masses  coming  into  contact  again,  last  time  during  the  Jurassic-Creta- 
ceous era,  which  saw  the  origin,  evolution  and  dispersal  over  the  globe  of  the 
Angiosperms,  to  be  followed  by  the  last  drift  age. 

With  regard  to  the  relations  and  disjunctions  between  South  America  (with 
Juan  I'>rnandez,  etc.)  and  Antarctica,  Australia  and  New  Zealand,  vertical  move- 
ments seem   to  otTer  a  less  distant  possibility. 

In  a  most  cleverly  written  chapter  Gooi)  [log.  344-360)  discussed  land- 
bridge  versus  continental  drift.  If  we  cannot,  he  says,  accept  the  former,  nor 
put  our  trust  in  disj^ersal,  a  changing  position  of  the  continents  is  the  only  way 
out  of  the  difficulties.  This  may  be  true,  but  we  have  seen  that  this  theory  does 
not  help  us  to  solve  the  problem  concerning  the  oceanic  islands.  GORDON  (//?) 
|)()ints  out  that  the  occurrence  of  a  small  but  important  subantarctic  element  in 
the  Pacific,  reaching  north  to  the  Hawaiian  Islands  where  it  is  better  displayed 
than  in  anv  of  the  Polynesian  or  Melanesian  groups,  makes  it  impossible  to  deny 
both  land  bridges  and  the  efficiency  of  transoceanic  dispersal  without  providing 
for   nngration    with    the   help   of  shifting   continents,    (iooi) 

has  run  into  an  impissc  f)vcr  the  Pacific  islands  like  Hawaii.  He  has  rejected  the  land- 
bridge  hvpothcsis  in  fivour  of  ( outineutal  drift....  Pnit  he  excludes  continental  drift  so 
tar  as  the  ishmds  arc  (  oik  rrned,  for  he  accepts  them  as  truly  oceanic,  not  continental  frag- 
ments. \'et  he  will  not  ;i!  i  -pt  oversi-as  migration.  Well,  I  can't  see  what  explanation 
remains,  if  all  the-e  tlinr  ;>!.■  e\(  hided,  but  the  plants  are  there  (p.   148). 

W  ri.l  I  {j<j/\  found  that  the  biogeographers  have  good  reason  to  support 
\\  i;(;i:m;Iv  s  tluM)r\-;  much  s|)eaks  against  it,  but  he  trusts  that  the  difficulties 
will  be  overcome  by  and  1)\-.  He  recognized  that,  with  regard  to  the  Pacific, 
they  are  ver>-  considerable  and   call    for  a  modification  of  the  theory. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  33I 

Chapter  VI. 

Transoceanic  migration. 

In  some  of  my  earlier  papers  I  touclied  upon  the  great  problem  of  long- 
distance dispersal  and  the  supposed  efficiency  of  the  transporting  agents;  see  for 
instance  2ji.  20-30,  where,  however,  only  the  flora  was  concerned.  This  time 
also  the  faunas  are,  to  some  extent  at  least,  considered,  and  I  shall  quote  a  num- 
ber of  authors,  old  and  modern,  who  have  expressed  their  opinion  for  or  against 
overseas  transport  as  the  only  possible  means  by  which  the  isolated  islands  of  the 
Pacific  have  received  their  indigenous  flora  and  fauna. 

Advocates  of  large-scale  overseas  migration. 

Among  earlier  authors  Engler  [8^)  and  Grisebacii  exercised  great  influence 
on  their  contemporaries.  They  divided  the  world  into  flora  domains,  regions  and 
districts,  characterized  by  a  combination  of  certain  important  elements  and  by  a 
greater  or  lesser  degree  of  endemism,  but  still  they  never  doubted  the  facility  with 
which  plants  travelled  across  the  oceans;  progressive  endemism  was  the  unavoid- 
able corollary,  but  relict  endemism  was  recognized  as  important.  The  attitude  of 
this  school  is  adequately  expressed  by  GriSEBACH  (j^j.  469): 

So  merkwiirdig  es  auch  sein  mag,  dass  sogar  einzelne  Holzgewachse  sich  hier 
iiber  das  Stille  Meer  verbreitet  haben,  so  ist  ihre  Wanderung  doch  aus  der  antarktischen 
Meeresstromung,  den  herrschenden  Westwinden,  oder  durch  Mithilfe  der  Seevogel, 
vielleicht  auch  durch  alte  Verkehrswege  wohl  hinlanglich  zu  erklaren,  ohne  dass  die 
Annahme  von  Landverbindungen  in  der  Vorwelt  gerechtfertigt  ware,  die  durch  keine 
geologische  Tatsache  gestiitzt  wird. 

By  some  the  case  of  Krakatau  was  quoted  as  a  proof  that  plants  and  animals 
are  able  to  travel  across  water  barriers;  it  is  mentioned  by  Hayek  [304),  but 
with  reservation : 

Wie  die  Besiedeliing  einer  Insel  erfolgt,  haben  die  oben  angefiihrten  Beobachtungen 
bei  der  Wiederbesiedelung  des  Krakatau  gelehrt  .  .  .  Aber  die  Entfernung  des  Krakatau 
von  den  nachst  gelegenen  Inseln  ist  keine  allzugrosse,  sie  betragt  nur  etwa  18—40  Km, 
also  Entfernungen  die  auch  durch  die  Flugtiere  nicht  allzuschwer  iiberbriickt  werden 
konnen  (p.    251). 

When  we  have  to  deal  with  islands,  separated  by  thousands  of  miles  from 
all  continents,  the  difficulties  are  of  much  greater  magnitude,  and  he  continues: 

Und  doch  miissen  wir  annehmen,  dass  auch  die  weit  entfernt  gelegenen  Inseln  ihre 
Pflanzendecke  von  den  zunachst  gelegenen  Festlandern  (und  Inseln)  erhalten  haben, 
wenn  auch  vor  undenklichen  Zeiten  und  ganz  allmahlich.  Dafur  spricht  auch  der  Um- 
stand,  dass  die  Flora  dieser  Inseln  keineswegs  von  der  ubrigen  Flora  der  Erde  grund- 
verschieden  ist,  sondern  denselben  Pflanzenfamilien  angehort  wie  diese,  demnach  von 
derselben   abstammen   muss. 

The    only    exception    known    (at   that  time)  was  Juan  Fernandez  (Lactoridaceae). 
But  he  admits  that  there  are  grave  difficulties: 


332  C.   SKOTTSBERG 

Selbst  bei  Inseln,  die  erst  in  relativ  junger  Zeit  vom  Festlande  abgetrennt  worden 
sind,  ist  ein  weitcrer  Aiistaiisch  der  Florenelemente  zum  mindesten  wesentlich  erschwert 
iind   eine   weitere   /uwandcriing   von  Elementcn  der  Festlandsflora  wenig  wahrscheinlich. 

If  tliis  be  true,   lunv  was  immigration  over  tbousands  of  miles  ever  possible? 

Several  writers  who  have  {)aid  special  attention  to  Pacific  problems  occupy, 
more  or  less  do^^matically,  the  same  standpoint  as  Havek.  Setciiell,  with  whom 
I  had  the  j)ri\ile<;e  to  discuss  this  subject  on  various  occasions,  was  already  quoted 
{).  271  ;  I  shall  add  here  what  he  says,  in  the  same  paper,  about  migration  (2ig.  300). 
He  found  that  1  was  "too  narrow"  in  my  allowances  for  migration  possibilities; 
he  belie\ed  in  "migration  over  very  considerable  breadth  of  barrier,  whether  of 
sea  or  land",  and  absence  was  not  a  result  of  failure  to  migrate  successfully,  but 
could  be  explained  by  obstacles  to  establishment.  He  regarded  the  oceanic  islands 
as  Tertiar)',  but  in  his  summary  pp.  307-309  admitted  the  possibility  of  their 
being  considerabl}'  older,  late  Mesozoic  or  early  to  middle  Tertiary,  which  would 
give  time  for  extensive  progressive  evolution  of  endemic  taxa  and  for  the  dying- 
out  of  their  continental  ancestors;  or  they  had  developed  in  other  directions, 
making  the  relationships  difficult  or  impossible  to  recognize.  Geologists,  however, 
refuse  to  give  even  the  Hawaiian  Islands  a  greater  age  than  late  Tertiary  or  even 
Pleistocene.  It  goes  without  saying  that  travel  facilities  are  difi'erent  in  different 
cases;  spore-plants  are  supposed  to  spread  more  freely  than  seed-plants,  but  even 
these  are  supposed  to  be  quite  capable.  Thus  Stebbins  (ji'p.  537): 

'{"he  seeds  of  j)lants  may  occasionally  be  transported  over  many  hundreds  of  miles 
of  ocean  and  may  establish  themselves  on  Oceanic  islands  like  Hawaii,  Juan  Fernandez, 
St.    Helena  and   the   Canary    Islands. 

r'l.oKiN  has,  he  writes,  shown  that  conifers  of  the  south  hemisphere  have  migrated 
freely  from  Australasia  to  South  Ainerica  and  vice  versa,  w-hereas  mammals  are 
unable  to  pass  and  are  absent  from  oceanic  islands — but  is  it  not  customary  to 
place  them  on  a  j)ar.'  S'lEHl^.lxs'  Antarctic  connection  does  not  include  land-bridges, 
for  "it  existed  for  plants,  but  not  for  vertebrates"  (but  what  about  birds?).  He 
looks  for  assistance  in  lost  islands  between  Antarctica  and  New  Zealand;  on  the 
o{)|)osite  side  the  width  of  oj)en  water  is  not  so  great,  and  seeds  can  still  be  carried 
from   South   America  to  Antarctica  without  much   difficulty. 

As  mentioned  before,  no  botanist  has  greater  confidence  in  long-distance  dis- 
persal  than    h'()Siii;K(; : 

.  .  .  transo(eani<;  migration  across  at  least  2500  miles  without  stej^ping-stones  is  not 
only   a   possibility   hut   a   relatively   common   occurrence   (99.867). 

Im)SI!I:i<(.s  subject  was  the  American  element  in  the  Hawaiian  flora,  but  in  order 
to  exj)lain  the  j)resence  f)f  the  dominant  Australasian  element  we  must  count  with 
still  gieater  distances.  AxKl.RoD  (7^)  quoting  F'o.SBERG  takes  a  modified  position. 
In  case  ot  distances  not  exceeding  some  200  or  300  miles  there  are  no  difficulties, 
"a  coni|)]ete  flora  can  transgress  such  a  barrier  without  the  loss  of  any  significant 
floristic  units  '.  A  greater  distance  results  in  "waif  assemblages",  but  many  will 
find   it  impossible  to  regard   e.g.   the   Hawaiian   flora  as  a  haphazard  accumulation 


DERIVATION  OF  THE  FLORA  AND  FAUNA  333 

of  waifs  and  their  descendants.  What  AxELROD  says  about  migration  probabiHties 
during  different  geological  epochs  is  of  greater  interest. 

Since  plants  are  controlled  largely  by  climate,  and  since  climate  has  been  changing 
during  geologic  time,  it  follows  that  plants  comprising  different  communities  have  had 
different  possibilities  at  different  times  .  .  .  probabilities  for  long-distance  migration  were 
much  higher  for  tropical  plants  in  the  Eocene  than  they  are  to-day.  Temperate  forest 
species  had  a  m.uch  higher  probability  from  late  Cretaceous  to  middle  Tertiary,  it  is 
low  now.  Steppe  plants  had  a  higher  probability  during  Pliocene  than  now.  Desert  spe- 
cies have  a  higher  probability  to-day  than   at  any  time  before. 

His  conclusions  are  drawn  from  the  size  and  area  of  populations  shifting  with  the 
extension  of  climatic  regions. 

For  my  own  part  I  have  expressed  my  opinion  on  overseas  migration  in  the 
Pacific  on  various  occasions  [2JI,  318,  248)  and  I  am  not  going  to  repeat  the  dis- 
cussion here.  My  general  conclusion  was  that  the  effect  of  transoceanic  migration 
has  been  largely  overestimated. 

GUPPY  (121),  who  allowed  birds,  winds  and  currents  to  stock  all  oceanic 
islands  with  plants,  arrived  at  the  conclusion  that  this  traffic  was  a  thing  of  the 
past  and  that  migration  had  practically  ceased  altogether.  I  expressed  my  doubts 
that  it  had  ever  been  effective,  in  any  case  with  regard  to  seed-plants. 

I  have  already  remarked  that  Setctiell  laid  stress  upon  what  he  called  the 
CEB  (climatic-edaphic-biotic)  factor  complex.  The  main  difficulty  for  the  vagabond 
plants  was  not  to  cover  the  distance,  be  it  ever  so  great,  but  to  become  a  successful 
member  of  a  community  already  established  in  the  place  where  it  happened  to 
alight,  and  this  difficulty  increased  as  time  went  by;  most  surfaces  of  the  earth, 
he  says,  are  already  stocked  with  closed  vegetation,  making  it  impossible  for  new 
arrivals  to  gain  a  foothold  {2ig.  300).  His  ideas  are  clearly  expressed  in  218  (\).  874). 

As  the  islands  have  become  more  and  more  completely  stocked  each  with  its  quota 
of  plants  and  animals  and  have  undergone  various  vicissitudes,  particularly  of  elevation, 
erosion,  etc.  its  hospitality  to  migrating  germules  necessarily  has  become  less  and  less, 
the  Biotic  factor  has  become  more  complex  and  the  Edaphic  factor  has  also  suffered  change. 

In  my  view  the  result  could  just  as  well  be  the  opposite,  for  these  "vicis- 
situdes", emergence,  erosion,  volcanic  activity  and  so  forth  create  new  soil,  a  more 
varied  topography,  a  multitude  of  different  habitats,  all  of  which  ought  to  give 
newcomers  increased  opportunities  to  get  established. 

With  Setchell,  Andrews  underlines  the  importance  of  CEB;  genera 
expected  to  occur  in  Hawaii  but  absent  "were  not  amenable  to  germination 
and  survival  after  transport". 

Long  before  Setciiell,  J.  D.  Hooker,  Wallace  and  others  had  paid 
attention  to  the  obstacles  for  the  successful  establishment  of  newcomers,  WALLACE 
believed  that  St.  Helena  had  become  stocked  with  plants  during  early  Tertiary 
time;  later  there  was  no  room  left,  and  the  flora  had  changed  so  completely  that 
no  plant  was  recognized  as  an  insular  form  of  a  continental  species. 

We  meet  with  Setchell's  line  of  thinking  in  a  recent  paper  by  W.  B.  Taylor 
{26J.  572).  In  recent  volcanic  islands  are  many  unstocked  habitats  to  begin  with, 


334  ^-   SKOTTSBERG 

but  each  new  species  would  mean  competition,  and  the  entry  of  an  additional 
species  would  be  very  difhcult  and  consequently  of  rare  occurrence.  This  may 
be  so,  but  tiiere  are  nian\-  communities  of  a  more  open  character  than  the  forest, 
and  even  a  closed  forest  is  not  like  a  tin  packed  with  cigarettes;  young  secondary 
forest  associations,  steppes  and  savannas  ought  to  offer  good  housing  grounds 
to  an  intruder,  suj)posing  that  he  likes  climate  and  soil.  Experience  shows  that 
numerous  aggressive  plants  brought  by  man,  not  only  herbs  but  also  trees  and 
shrubs,  tind  suitable  living  conditions  even  in  undisturbed  natural  communities. 
In  Juan  Fernandez  I  have  had  occasion  to  follow  the  invasion  by  Aristotelia  maqui 
(chilensis)  and  to  witness  the  fabulous  ease  with  which  it  crowds  out  the  native 
vegetation,  and  to  observe  I'i^Jii  Moliiiac  springing  up  on  the  ridges  where  the 
j)lant  cover  was  o|)en  and,  from  there,  to  enter  the  dense  native  brushwood. 
And  they  are  only  two  of  the  many  successful  weeds,  a  third  one,  equally  dan- 
gerous but  of  quite  recent  introduction,  is  Rubus  nluiifolius.  All  three  have  fleshy 
fruits  and  are  eaten  by  man  and  birds  and  propagate  themselves  rapidly.  They 
are  conunon  on  the  opposite  mainland  (where,  of  course,  the  brambleberry  was 
introduced  from  luirope),  but  man,  no  bird,  carried  them  across  to  the  islands. 
Similar  examples  are,  I  presume,  oftered  by  almost  all  oceanic  islands.  I  just  hap- 
j)ene(l  to  read  a  book  on  Cape  Verde  Islands,  where  a  naturalist  tells  us  about 
La)ita)ia  caniara  sj^reading  like  wildfire  and  menacing  the  little  there  is  left  of 
natural  savanna  and  steppe  [326). 

Most  zoologists  favour  the  theory  of  long-distance  dispersal.  Mammals  are, 
as  a  rule  also  bats,  flying  foxes  and  the  like,  excluded,  but  of  birds  some  are  able 
to  cover  very  large  distances,  winged  insects  are  carried  ofl"  to  distant  places 
where  they  never  wanted  to  go,  and  so  forth.  I  shall  quote  some  zoogeographers 
who,  with  reference  to  the  Pacific,  have  expressed  their  opinion  on  overseas  transport. 

Pl-.KKINS,   in   his  introduction  to  Fauna  Haw-aiiensis  (j^j.  XLVl),  wrote: 

All  the  islands  being  volcanic  and  having  been  built  up  from  a  great  depth  of  ocean 
at  various  j)eriods,  their  entire  fauna  naturally  originated  from  immigrants  derived  from 
other  lands.  These  immigrants  must  have  arrived  either  by  flight,  like  the  birds,  or  in 
drift   Hke   the   flightless  insects  and   jirobably  the  land  Mollusca. 

Dritting  logs  were  often  regarded  as  an  important  kind  of  conveyance,  but  they 
come  from  North  America  and  what  they  bring  of  animals,  PERKINS  says,  would 
ser\e  no  |)urpose  because  it  is  unlikely  that  the  passengers  would  become  acclim- 
atized in  ilawaii.  The  fauna  must  have  come  from  the  warmer  parts  of  America, 
from  Australia,  Polynesia  etc.  "at  rare  intervals  from  the  F^ocene  until  now".  If 
we  have  to  belie\e  the  geologists,  no  Ilawaii  existed  in  the  F^ocene — and  how 
did  those,  who  arrive  "now",  manage  to  become  endemic  genera  and  species? — 
non-endemic  (lowering  plants  not  brought  by  man  are  few. 

In  some  instances  (ii  LICK  [Tig]  admits  the  possibility  of  land  connections, 
but  Ilawaii,  Juan  hernandez,  (ialaj)agos,  St.  Helena,  etc.  etc.,  have  always  been 
isolated.  'I  he  (juestion  whether  their  fauna  shows  that  "the  ancestors  possessed 
an  almost  inconceivable  capacity  for  passing  uninjured  over  vast  stretches  of  ocean" 
is    answered    in    the  affirmative.    It   is  significant  that  the  Galapagos  archipelago 


DERIVATION  OF  THE  FLORA  AND  FAUNA  335 

"was  successfully  reached  by  a  giant  tortoise";  already  Wallace  entertained  the 
same  idea — a  strictly  terrestrial  animal  crossing  the  ocean. 

Some  biogeographers  prefer  one  dispersal  agent,  some  another,  most  have 
confidence  in  all,  but  it  goes  without  saying  that  different  types  of  plants  and 
animals  have  availed  themselves  of  different  kinds  of  transport,  I  shall  quote 
GULICK  (iig.414)  first. 

It  is  possible  to  go  far  toward  a  first  diagnosis  of  the  degree  of  a  land's  insularity  by 
noting  how  exclusively  it  is  peopled  by  types  with  a  known  capacity  for  colonizing  across 
vast  expanses  of  ocean.  Our  summary  up  to  this  point  reveals  very  nearly  which  these 
forms  may  be.  Quite  a  majority  of  them,  both  plants  and  animals,  show  characters  that  har- 
monize with  wind-storm  transportation.  A  respectable  majority  of  the  larger-seeded  palms 
and  some  tough-lived  earth-inhabiting  invertebrates,  suggest  transportation  by  water  or  on 
drift-wood.  Such  seeds  and  invertebrate  eggs  as  can  withstand  the  digestive  tracts  of  a  bird, 
have  a  very  substantial  travelling  radius  by  that  means,  easy  500  miles  in  the  routine 
seasonal  migrations,  and  possibly  stretching  in  the  extremest  cases  to  almost  transoceanic 
distances.  .  .  .  Dioecious  plants  and  separate-sexed  animals  are  statistically  at  a  disadvan- 
tage, as  compared  to  the  reversed  condition,  because  of  their  poorer  chance  of  achieving 
fertilization.    The  ability  to  take  a  journey  in  a  gravid  condition  helps  the  chances  greatly. 

"Types  with  a  known  capacity  for  colonizing" — GULlCK  proceeds  from 
what  should  be  proved,  for  their  occurrence  on  isolated  islands  is  in  itself  no 
proof  of  oceanity.  Under  his  angle  the  great  number  of  dioecious  endemic 
phanerogams  in  Hawaii  ought  to  have  surprised  him.  It  almost  seems  as  if  he 
believed  that  entire  specimens  with  roots  and  all  managed  to  reach  a  distant 
island  and  get  established;  surely,  if  only  a  male  or  an  unfertilized  female  arrived) 
all  was  in  vain  until  a  mate  of  the  opposite  sex  turned  up;  a  pregnant  female 
would  of  course  do  better  (bye  the  bye,  WALLACE  tells  a  story  of  a  pregnant 
boa  constrictor  arriving  on  a  West  Indian  island  with  drift-wood  and  in  good 
condition).  I  guess  we  can  leave  these  chances  aside,  for  plants  spread  by 
means  of  seeds,  and  whether  wind-blown,  epizoic  or  endozoic  (provided  they 
do  not,  as  many  assert,  discharge  their  droppings  soon  after  the  departure), 
there  is  every  chance  that  more  than  one  seed  of  the  same  kind  is  brought;  a 
single  many-seeded  berry  is  enough,  and  a  bird  picking  drupes  fills  his  stomach. 
A  seed  portion  of  a  dioecious  species  gives,  under  ordinary  conditions,  50  %  of 
each  sex.  In  the  Hawaiian  flora  we  find,  GULiCK  says  (p.  418),  "a  preponderance 
of  plants  spread  by  wind-carried  spores  and  minute  seeds";  species  with  drupes 
and  berries  are,  however,  numerous.  As  an  example  of  a  presumably  definite 
case  of  bird  rather  than  wind  carriage  he  mentions  the  Hawaiian  species  of 
Vaccinium,  which  he  derives  from  North  America.  Their  presence  is  most  in- 
teresting, "as  the  distances  involved  must  be  very  close  to  the  extreme  physio- 
logical maximum  that  land  birds  can  traverse,  and  still  carry  fruit  seeds  in  their 
droppings".  To  me  it  appears  as  a  bad  case  of  constipation.  Besides,  the  Ha- 
waiian Vaccinia  are  not  related  to  North]  American  groups  but  belong  to  a 
special  section. 

Mayr,  an  extreme  "oceanist"  who  refuses  to  admit  land  connections  for 
either  Fiji  or  New  Caledonia,  in  his  paper  on  the  Pacific  bird  fauna  [lyg]  includes 


336  C.   SKOTTSBERG 

a  general  survey  of  the  dispersal  chances  for  other  animals  and  also  for  plants. 
The  special  instance  cited  (p.    197)  is  not  very  convincing. 

liirds  are  excellent  rivers  and  thus  cai)able  of  ra])id  and  active  spreading  .  .  .  capable 
of  crossing  considerable  stretches  of  open  sea  to  settle  in  new  territories.  There  is  abun- 
dant evidence  of  this,  such  as  the  resettlement  of  Krakatau  Island,  the  recent  arrival  of 
Australian  birds  in  New  Zealand,  and  the  colonization  of  untjuestionably  oceanic  islands. 

Not  even  the  arrival  of  Australian  birds  in  New  Zealand  brings  conviction; 
the  colonization  of  "unquestionably  oceanic  islands"  certainly  does,  if  we  can 
prove  it.    Mavr  continues  p.    198: 

'i'he  possil)ility  of  transport  by  floats  or  in  logs  is  not  to  be  underestimated.  Many 
tro])ical  currents  have  a  sj)ced  of  at  least  2  knots,  that  is,  about  50  miles  a  day,  or  1000 
miles  in  three  weeks.  It  is  probably  not  a  great  task  for  a  wood-boring  insect  to  survive 
3  weeks  in  a  drifting  log.  Air  currents  are,  however,  of  uncomparably  greater  importance 
than  sea  currents.  Even  slight  winds  are  of  great  influence  on  the  distribution  of  floating 
and  flying  animals,  as  recent  investigations  have  shown.  It  is  astonishing  how  rich  the 
"aerial  plankton"  is,  even  up  to  altitudes  of  1000  meters  and  more.  Normal  winds  would, 
of  course,  not  account  for  the  spreading  of  molluscs,  flightless  insects,  and  other  small  in- 
vertebrates. However,  most  of  the  islands,  with  which  we  are  concerned,  are  situated 
within  the  zone  of  tropical  hurricanes,  the  lifting  force  of  which  is  quite  extraordinary.  .  .  . 

The  fact  that  there  are  small  molluscs  and  flightless  insects  on  such  typical  oceanic 
islands  as  Kaster  Island,  juan  Fernandez  and  Saint  Helena  is  almost  unassailable  proof 
that  such  a  method  of  disj:)ersal  is  a  reality.  Tropical  hurricanes  carry  for  hundreds  and 
even  thousands  of  miles.  .  .  .  I'he  result  of  the  recent  surveys  in  the  Hawaiian  Islands, 
the  Mar([uesas,  and  on  Samoa  indicate  that  there  are  indeed  very  few  animals  that 
cannot  be  transported  across  considerable  stretches  of  the  sea  by  winds,  waves,  other 
animals   or   man. 

On  p.   201   Mavr  adds  some  general  remarks: 

The  means  of  dispersal  of  most  plants  and  animals  are  much  more  extensive  than 
was  formerly  realized,  and  even  rather  irregular  distributions  can  be  explained  without 
the  hclj)  of  land  bridges.  Dispersal  across  the  sea  is,  of  course,  most  obvious  for  birds, 
and  ornithologists  were  among  the  first  who  accepted  the  ideas  of  the  permanency  of 
continents  and  oceans.  Most  entomologists  are  also  beginning  to  realize  that  they  can 
solve  most  of  their  distribution  difficulties  without  land  bridges.  The  conchologist, 
hf)wcver,  ])Ostu]ates  even  today  continental  connections  between  all  or  nearly  all  the  is- 
hmds  where  land  shells  exist. 

As  we  shall  see  below  ([).  350)  Mayr  declared  himself  unable  to  explain  how 
land  shells  arc  (lisj)crsecl. 

It  seems  that,  also  with  regard  to  the  birds,  Mavr  contradicts  himself,  for  in 
the  same  paper  (j).  Kj.S)  he  asserts  that  most  birds,  particularly  on  tropical  islands, 
|)recisely  the  islands  we  are  discussing,  are  extraordinarily  sedentary,  and  as  an 
examj)le  he  mentions  that  of  265  species  known  from  that  part  of  New  Guinea 
which  is  ()j)posite  New  Britain,  a  distance  of  45  miles,  only  80  occur  on  New 
Britain,  and  the  situation  in  Western  Papuan  islands  is  even  more  conspicuous; 
he  mentions  two  islands  only  2  miles  apart,  with  rather  different  fauna.  "Literally 
hundreds  of  similar  instances  could  be  listed  ...  all  of  them  indicating  the  sedentary 
habits."  One  is  likely  to  remember  Cirri'V's  fruit-eating  pigeons  which  were 
thought  to  be  res|K)nsible  for  the  dissemination  across  the  Pacific  of  seeds  too 
large  for  other  birds.  The  "pigeons"   are,  now  at  least,  restricted  in  range  and  of 


DERIVATION  OF  THE  FLORA  AND  FAUNA  337 

very  little  use  on  longer  distances.  I  cannot  help  drawing  the  conclusion  from 
this  that  the  sedentary  habit  was  acquired  after  the  great  colonization  had  taken  place. 
Few  phytogeographers  have  had  greater  faith  in  the  capacity  of  wide-ranging 
marine  birds  to  carry  diaspores  than  Grisebach.  In  his  discussion  of  bipolar 
species  found  in  the  far  north  and  the  far  south  but  not  at  all  in  intermediate 
zones  he  selected  Geiitiajia  prosirata  Haenke  as  the  best  example.  Its  distribu- 
tion is  due,  he  says,  to  the  wanderings  of  Diomedea  exulans  which, 

abweichend  von  der  Lebensweise  der  meisten  anderen  Zugvogel,  iiber  beide  Hemispharen, 
von  Kap  Horn  bis  zu  den  Kurilen  und  Kamtschatka,  wandert  und  die  Standorte  jener 
Pflanze  in  der  arktischen  und  antarktischen  Flora  in  Verbindung  setzt.  Mil  der  Beute, 
die  dieser  Vogel  verschlingt,  kann  er  auch  Samen  von  Pflanzen,  welche,  mil  den  Fliissen 
ins  Meer  gespult,  in  den  Magen  der  Fische  ubergehen,  in  einzelnen  Fallen  ausstreuen, 
so  dass  sie  an  fernen  Kiisten  aus  seinem  Dunger  aufkeimen  [32^.  469). 

I  have  not  come  across  any  comments  on  this  bold  theory.  It  is  difficult 
to  take  it  seriously,  but  to  Grisebach  the  only  gap  in  his  argumentation  was 
that  nobody  had  happened  to  witness  such  an  event.  If  he  is  wrong,  he  asks, 
why  is  there  no  trace  of  this  Gentiana  in  the  Andes,  where  it  would  thrive  just 
as  well  as  in  the  Alps  and  in  the  mountains  of  Asia?  To  this  should  be  re- 
marked that  G.  prosirata  is  a  polymorphous  species  of  wide  range  and  that  it 
does  occur  in  the  Andes  from  Colombia  to  Chile,  suggesting  that  it  has  mi- 
grated south  along  the  mountains  without  the  assistance  of  the  albatross. 

Wallace  (^/c^'.  259)  tells  us,  on  the  authority  of  MoSELEY,  naturalist  to  the 
"Challenger"  expedition,  of  the  great  albatross  breeding  on  Marion  Island  in  the 
midst  of  dense,  low  herbage;  I  can  add  that  this  bird  also  breeds  on  South 
Georgia  and  on  some  other  southern  islands,  but  as  far  as  I  know  they  do  not 
shift  breeding  places,  and  even  if  they  did,  they  do  not  go  on  shore  between 
the  breeding  seasons.  TAYLOR,  in  his  important  paper  on  Macquarie  Island  [26 j), 
tells  us  about  a  giant  petrel  which  was  captured,  tagged  and  released  on  this 
island  and  shot  on  South  Georgia,  8000  km  away,  four  months  later,  but  these 
birds  are  often  seen  on  land  where  they  attack  the  penguin  chickens;  this  was 
at  least  the  case  on  Paulet  Island  in  the  Antarctic.  Whether  they  aid  in  the 
dispersal  of  diaspores  is  unknown.^  Taylor  quotes  an  observation,  made  on 
Macquarie,  that  seeds  were  found  adhering  to  the  feet  of  an  albatross.  These 
birds,  when  building  their  nests,  regurgitate  an  oily  fluid  which  makes  seeds  stick 
to  their  feet.  Macquarie  Island  was  ice-covered  during  the  Glacial  epoch  and  the 
plants,  perhaps  with  the  exception  of  some  cryptogams,  must  have  arrived  since 
the  retreat  of  the  ice.  The  vascular  flora  consists  of  35  species,  all  except  4 
occurring  in  the  New  Zealand  subantarctic  area — the  3  species  with  a  claim 
to  be  regarded  as  endemic  should  be  reinvestigated — while  those  4  species  are 
found    in    subantarctic    South    America,  from  where  they  are  derived.     All  Mac- 

^  According  to  Taylor  (p.  570)  the  two  truly  Antarctic  phanerogams,  Deschampsia  ant- 
arctica  and  Colobanthus  crassifolius,  are  very  rare  in  the  Antarctic  and  reproduce  only  vegeta- 
tively.  I  do  not  know  where  he  obtained  this  information.  They  are  scarce  but  have  been 
reported  from  many  localities  along  the  coast  and  adjacent  islands  of  Palmer  Land  between 
lat.  62  and  68  and,  in  favourable  situations  at  least,  both  of  them  flower  and  produce  ripe 
seeds — see  my  paper  in  Botan.  Tidsskrift  vol.  51,   1954- 

22  -  557857    The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.    Vol.  I 


338 


C.   SKOTTSBERG 


quarie  plants  have,  Tavlor  states,  propagules  suited  to  bird  transport.  It  is  sur- 
prising that  Wkrhi  [jiy),  who  made  a  detailed  study  of  the  Kerguelen  flora, 
asserts  that  not  one  of  the  flowering  plants  possesses  any  special  dispersal  mech- 
anism for  either  wind,  water  or  bird  carriage.  Still,  the  two  islands  have  some  9 
species  in  common  ( -,v).  Taylor's  conclusion  that,  "if  long-distance  dispersal 
has  occurred  on  Macquarie  Island,  then  it  could  well  have  occurred  elsewhere" 
is  certainly  correct;  we  know,  for  one  thing,  many  wide-spread  sea-side  plants 
and  a  number  of  widely  dispersed  aquatic  species,  possibly  transported  by 
migratory  birds.  South  Georgia  is  in  much  the  same  situation  as  Macquarie, 
but  still  rather  heavily  glaciated  thanks  to  its  great  altitude,  and  the  possibility 
that  any  higher  plants  survived  the  Glacial  epoch  is  very  small  indeed,  whereas 
intlications  that  many  mosses  and  lichens  date  from  preglacial  times  are  strong. 
The  vascular  flora  is  poorer  than  on  Macquarie,  and  there  are  no  endemic  species. 
When  Ta\i,()R  accuses  me  of  having  argued  against  all  overseas  migration  also 
n  this  case  he  must  have  misunderstood  me.  I  expressly  took  this  possibility 
into  account  in  the  paper  he  quotes  (226). 

W'ali.ack  calls  attention  to  sea  birds  breeding  on  islands  in  the  tropics; 
Phaeioji  makes  its  nests  on  the  Hawaiian  Islands  in  4000  ft.  altitude  and  also  in 
the  highland  of  Tahiti,  and  such  birds  would  account  for  the  similarity  of  the 
mountain  floras.  In  reality  these  floras  have  practically  nothing  in  common. 
Miss  GiUHS  (?.?/),  discussing  the  origin  of  the  montane  flora  of  Fiji,  refused  to 
regard  birds  as  capable  agents;   wind  may  have  been  more  efficient. 

No  modern  zoologist  has  tried  to  defend  the  theory  of  unlimited  overseas 
migration  with  greater  zeal  than  Zimmerman.  In  his  Introduction  to  "Insects  of 
Hawaii"   we  read: 

There  is  no  evidence  whatsoever  to  support  the  contention  that  they  (i.e.  the  Ha- 
waiian Is.)  are  of  continental  origin  or  character,  or  that  they  were  ever  joined  together 
in  an  elongate  subcontinental  land-mass  or  even  in  a  continuous  subaerial  mountain 
range   {2()8.  6). 

And,  in  opposition  to  certain  other  biologists  he  refuses  to  regard  the  islands  as 
old,  they  are  at  most  Pliocene  and  no  part  of  them  older  than  five  million  years; 
most  of  the  lava  is  younger,  the  bulk  of  the  land  Pleistocene.  Pie  is  opposed  to 
my  ideas  but  he  thinks  that  the  explanation  offered  by  him  will,  partially  at  least, 
reconcile  the  differences  between  us.  In  an  earlier  paper  (Amer.  Naturalist  'j^y 
1942),  to  which  he  refers,  he  spoke  of  former  high  islands,  other  than  those  found 
on  maps,  which  once  existed;  once  more  the  "stepping-stones  routes"  are  called 
to  life.  Atolls  are  the  remnant  of  many  of  them,  or  reefs  like  among  the  Leeward 
islands  of  Hawaii,  and  such  preexisting  islands  would  account  for  the  immigration 
from  all  directions.  He  does  not  call  for  jumping  of  thousands  of  miles  of  open 
sea,  but  rather  for  series  of  shorter  over-water  steps.  I  am  afraid  that  we  need 
some  substantial  refurnishing  of  the  Pacific  basin  to  supi)ly  a  sufficient  number 
of  intermediate  stations.  Not  all  of  these  routes  were,  he  says,  available  at  the 
same  time,  and  this  would  explain  the  apparent  difference  in  age  of  various 
sections  of  the  biota  (pp.  51,  52).    Most  of  the  roads  were  cut  off  in  Pliocene  and 


DERIVATION  OF  THE  FLORA  AND  FAUNA  339 

early  Pleistocene,  some  before  Pliocene — when  no  land  existed  where  stands  the 
Hawaiian  chain,  if  geologists  are  right.  UsiNGER  (^/j),  describing  the  distribu- 
tion of  Heteroptera,  also  looks  for  convenient  stepping-stones. 

Divergent  opinio7is  on  the  means  of  transport. 

The  dispersal  agents  universally  recognized  as  important  are  air  currents, 
especially  monsoon  and  trade  winds,  and  cyclonic  storms,  ocean  currents,  birds, 
and  last  not  least  man,  who  with  his  domestic  animals  and  goods  has  become 
more  and  more  important,  whether  he  brings  plants,  seeds  and  animals  to  extend 
their  range — and  many  of  them  become  naturalized — or  carries  diaspores  on  his 
body  and  his  belongings  unaware.  It  is  as  a  rule  not  difficult  to  find  out  where 
we  have  to  do  with  human  action,  but  we  shall  limit  ourselves  here  to  a  discus- 
sion of  natural  factors  of  distribution.  This  subject  has  been  treated  by  innumer- 
able writers  in  biology  and  a  wealth  of  material  was  compiled  by  RiDLEY  (20^), 
a  firm  believer  in  the  great  value  of  all  kinds  of  dispersal  mechanisms,  some  of 
which  are,  of  course,  very,  wonderful.  WULFF  (2^1),  in  his  chapter  "Natural  factors 
for  distribution"  is  more  critical.  Of  animals  only  the  birds  deserve  to  be  men- 
tioned, but  the  plumage  is  no  good  for  carrying  diaspores  any  considerable  dis- 
tance, especially  over  the  sea,  and  the  extreme  marine  birds,  the  strongest  flyers, 
have  no  contact  with  land  outside  their  own  breeding-places.  WULFF  reduces  the 
part  taken  by  birds  to  almost  nothing,  but  I  believe  that  we  have  good  reason 
to  count  with  the  migratory  birds  in  certain  cases.  Water  transport  is  responsible 
for  the  diffusion  of  litoral  halophytes  but  rarely  for  migrations  of  inland  species. 
Wind  is  important  only  on  short  distances,  at  least  for  seed-plants;  special  devices 
do  not  help  very  much.  Even  Ridley  concluded  that  winged  or  plumed  diaspores 
are  not  carried  very  far;  spore-plants  are  more  easily  spread.  Wulff  remarks 
that  according  to  Bentham  Leguminosae  and  Labiatae  hold  their  ground  just 
as  well  if  not  better  than  Compositae.  Altogether,  if  dispersal  by  natural  factors 
had  the  significance  ascribed  to  it,  the  vegetation  of  the  globe,  within  a  certain 
climatic  zone,  would  be  homogeneous  and  the  sporeplants  at  least  ought  to  be 
cosmopolitan,  but  they  are  "localized  in  definite  areas,  their  distribution  paralleling 
that  of  flowering  plants"  (p.  128).  GoOD  observes  [log)  that  we  have  no  proofs 
that  species  equipped  with  special  dispersal  mechanisms  are  more  widely  distri- 
buted than  others.  That  certain  plants  with  such  devices  show  very  wide  areas 
whereas  others  without  them  are  rare  and  local  means  nothing,  because  the  reverse 
is  also  true;    climatic  and  edaphic  factors  should  always  be  taken  into  account. 

The  relative  value  of  the  dispersal  agents  is  put  to  the  test  when  we  deal 
with  oceanic  islands.  Setchell  (21'/)  was  inclined  to  give  considerable  credit  to 
migratory  birds.  The  occurrence  of  identical  species  of  flov.ering  plants  in  Arizona 
and  Argentina  and  in  California  and  Chile  could  be  explained  by  bird  transport, 
and  bipolarity  had  originated  in  the  same  way.  Bird  transport  helped  him  to 
understand  the  remarkable  disjunct  areas  of  arborescent  Compositae  and  Lobe- 
liaceae;  their  birth  place  was  in  the  Antarctic,  and  they  had  been  carried  by 
birds  to  New  Zealand,  Australia,  Malaysia,  Polynesia,  Hawaii,  South  America  and 
the  high  African  mountains — we  can  add  Saint  Helena,  Juan  Fernandez  and  Des- 


340 


C.   SKOTTSBERG 


venturadas.  In  this  case  it  is  not  the  question  of  identical  species  and  rarely  of 
j^enera.  It  is  not  improbable  that  the  secret  of  their  origin  and  early  history  lies 
hidden  in  the  far  soutii,  but  tiiis  is  all  we  can  say.  Hemslev  (727.  66)  when  dis- 
cussing the  endemic  Coni[)ositae  of  Saint  Helena  and  Juan  Fernandez  said  that 
"wind  seems  at  first  to  be  the  most  probable  agent";  still  he  doubted  its  efficiency. 
In  Setciiell's  view  storms  seem  to  ofler  more  than  a  possibility  in  many  in- 
stances, particularly  tropical  cyclones  and  vertical  thrombs  able  to  carry  even 
lieavy  diaspores  to  a  great  altitude.  In  another  paper  (218)  he  points  out  that 
plant  distribution  in  the  Pacific  has  been  from  west  to  east  against  the  prevailing 
winds  and  currents  and,  in  the  case  of  Hawaii,  has  given  much  better  results 
than  the  expected  east-west  route;  the  "frequent  cyclonic  storms"  are  responsible 
for  this  anomal}'  together  with  adverse  biotic  factors,  but  I  fail  to  see  why  they 
shouldn't  ofter  the  same  difficulties  for  diaspores  coming  from  the  west. 

Andrews  (^.615)  paid  s})ecial  attention  to  the  occurrence  of  scattered  "Ant- 
arctic "  genera  and  species  in  and  around  the  Pacific  and  combines  their  distri- 
bution  with  the  direction  of  ocean  currents: 

In  the  South  Pacific  the  westerly  current  sweeps  by  Australia,  New  Zealand,  Tahiti, 
and  the  whole  of  the  west  const  of  South  America,  where  it  is  joined  by  the  cold 
uprising  water  along  the  South  American  coast.  This  gives  rise  to  the  north-moving 
Peruvian  C'urrent  which  sweeps  by  Juan  Fernandez,  Peru,  Central  America,  and  Mexico, 
whence  there  is  a  detlection  westerly  toward  the  Hawaiian  Islands  and  the  tropics. 
This  knowledge  of  the  general  circulation  within  the  Pacific  appears  to  throw  a  flood 
of  light  on  the  occurrence  of  the  Australian,  New  Zealand  and  western  South  American 
elements  in  the  Hawaiian  flora,  such  as...  (2  5  genera  are  enumerated).  The  influence 
of  ocean  currents  is  suggested  i)articularly  in  the  peculiar  distribution  oi  Acaena,  Gunnera, 
Acrfcra,    On-obolus,  Sautalinn,   Sophora,   and   so   on. 

Juan  hY-rnandez  lies  outside  the  Peruvian  current,  but  also  the  outer  island,  Masa- 
fuera,   is  reached  by  drift-wood.  Its  origin  has  not  been  investigated. 

A  look  at  a  current-chart  shows  that  Andrews'  reasoning  has  its  weak 
points;  besides,  I  cannot  see  that  the  genera  he  mentions  are  thalassochorous.  I 
would  recommend  the  reader  to  take  a  look  at  the  many  distribution  maps  pub- 
lished in  a  paper  read  on  the  same  occasion  when  Andrews  presented  his  argu- 
mentation   (  iVeS). 

\\  ith  regard  to  the  cyclonic  storms  several  authors  have,  as  we  have  seen, 
em[)hasized  their  j)r()minent  role  in  the  violent  dispersal  of  both  plants  and  ani- 
mals; they  are,  in  fact,  considered  to  be  the  only  imaginable  force  by  which 
larger  objects  are  trans|)()rte(l,  and  it  is  useless  to  deny  that  such  events  have 
taken  j)lace  and  still  take  })lace,  even  if  it  is  difficult  to  find  definite  proofs  that 
the  transport  did  lead  to  the  establishment  of  an  immigrant  from  afar.  Most 
authors  who  have  taken  refuge  in  c>'clones  have,  however,  expressed  themselves 
in  general  terms  without  a  clear  idea  of  the  extension  of  the  cyclonic  belts  and 
the  trend   of  the  cyclones. 

In  two  j)apers  \'lsiiER  has  summarized  his  studies  on  cyclonic  storms  in  the 
Pacific.  Three  chief  centres  of  origin  are  distinguished  [322):  (i)  Western  N.Pacific, 
originating   some  distance  east  of  the  Philippines  in   lat.   8°   to  25°;    (2)  Western 


DERIVATION  OF  THE  FLORA  AND  FAUNA  34I 

S.  Pacific,  particularly  between  Australia  and  Samoa  in  lat.  io°  to  25°;  (3)  Eastern 
N.  Pacific  off  the  west  coast  of  Mexico  and  Central  America.  Occasionally  tropical 
storms  develop  near  Hawaii  and  over  Australia.  The  normal  course  of  (i)  is  WNW, 
recurring  NE,  and  of  (2)  WSW,  recurring  SE.  To  what  extent  plant  distribution 
runs  parallel  to  cyclone  tracks  has,  as  far  as  I  am  aware,  not  been  investigated. 
As  ViSHER  says,  most  of  the  cyclones  originate  over  the  sea  "well  out  in  the 
ocean"  (p.  87).  They  hit  many  of  the  Polynesian  islands,  and  possibly  collect 
diaspores  on  one  and  deliver  them  on  another,  but  a  frequent  dispersal  of  species 
in  this  way  does  not  appear  very  probable.  I  wonder  whether  there  is  in  Hawaii, 
with  its  90  %  endemics,  a  single  flowering  plant  likely  to  have  been  borne  there 
by  a  cyclonic  storm.  ViSHER  is,  however,  opposed  to  land  connections  in  the 
Pacific,  with  one  exception:  "it  is  known  that  Australia  was  formerly  connected 
with  Asia  by  way  of  the  East  Indies  and  New  Caledonia"  (j^J.  77).  With  regard 
to  Polynesia  he  points  to  the  west-east  hurricanes  and  their  colonizing  power. 
Sea  carriage  also  comes  into  the  picture,  violent  cloudbursts  may  accompany  the 
storm,  brooks  are  transformed  into  swift  rivers  carrying  plant  material,  tree  trunks 
and  soil,  forming  rafts,  which  are  washed  out  into  the  ocean. 

A  recent  paper  by  Bergeron  (j^^)  gives  a  somewhat  different  aspect.  His 
map  shows  the  two  areas  in  the  Pacific  north  and  south  of  the  equator,  where 
hurricanes  arise,  and  how  they  move.  The  Hawaiian  islands  lie,  as  a  rule,  outside 
the  tracks.  The  direction  north  of  the  equator  is  NW  or  WNW  all  through  the 
hurricane  belt,  and  there  is  no  sign  of  an  easterly  direction  enabling  plants  and 
animals  to  be  carried  all  over  Polynesia  as  far  north  and  east  as  Hawaii,  as  ViSllER 
supposed.  South  of  the  equator  the  trend  is  S  and  SE;  the  east  of  the  Pacific 
is  not  reached  and  Juan  Fernandez  lies,  in  longitude  as  well  as  in  latitude,  away 
from  any  cyclonic  belt. 

It  has  often  been  stated  that  spore-plants,  theoretically  at  least,  have  much 
greater  facilities  to  colonize  on  long  distances,  but  it  has  been  shown  that  in 
reality  their  capacity  is  more  limited  than  was  formerly  assumed. 

COPELAND  (68.  165-166)  expressed  his  opinion  on  the  diffusion  of  fern  spores: 

Fern  spores  are  carried  across  water  by  the  wind — ten  miles  of  water  is  no  barrier 
at  all  to  their  spread.  One  hundred  miles  may  be  one  hundred  times  as  great  a  barrier, 
because  the  spores  must  hit  a  target,  a  suitable  place  to  germinate  and  grow.  Still, 
ferns  spread  readily  across  seas  this  wide.  A  thousand  miles  makes  the  obstacle  again 
one  hundred  times  as  great  ...  the  limited  viability  of  the  spores,  the  chance  of  falling 
or  being  washed  out  of  the  air,  and  the  chance  of  very  different  climate  at  such  dis- 
tance, increase  it  materially.  Ferns  rarely  jump  a  thousand  miles  of  ocean.  Still  a  number 
of  species  are  believed  to  have  crossed  the  south  Atlantic,  and  I  believe  that  three 
genera,  Plagiogyria,  Coniogramtne  and  Loxogra?nme,  have  flown  the  north  Pacific  from 
Japan  to  Mexico,  each  in  one  single  instance.  It  is  not  exacdy  impossible  that  direct 
colonization  has  occurred  between  Chile,  New  Zealand,  Tasmania,  the  Cape,  and  Tristan 
da  Cunha. 

It  should  be  mentioned  that  CllRlST  (59),  with  his  unique  knowledge  of  the 
distribution  of  ferns,  pointed  to  the  insignificance  of  spore  dispersal  as  an 
explanation  of  the  origin  of  widely  disjunct  areas. 


342 


C.   SKOTTSBERG 


The  fact  that  Irmsciikr  tried  to  prove  that  the  distribution  of  plants  strongly 
su|:)ports  \\'K(;fnkr's  hypothesis  does  not  lessen  the  value  of  what  he  says  about 
the  limited  capacity  of  plants  to  migrate.  It  is  small  in  the  flowering  plants  [14.3.  291): 

Dass  bei  den  Hliitenptlanzen  die  J^eforderung  der  Samen  und  Friichte  durch  Wind, 
Wasser  und  I'iere  ganz  wescntlich  eingeschrankt  werden  muss  und  fiir  geschlossene 
i-'orniationen  auf  grossen  Kntfernungen  hin  nicht  in  Frage  kommt,  ist  heute  von  den 
rtlan/.engeograi)hcn   allgeniein   anerkannt. 

Irmsciikr  is  too  optimistic,  we  have  seen  that  there  are  phytogeographers 
to  whom  overseas  migration  is  not  only  possible  and  undoubtedly  happens,  on 
rare  occasions  at  least,  but  rather  of  quite  common  occurrence.  And  with  regard 
to  s{)ore-plants  their  distribution  should,  if  this  be  correct,  show  quite  different 
distribution  patterns  than  they  actually  do.  The  bryophytes  are  no  exception  to 
the  rule.  iRMSCiil'.R  remarks  that  already  in  1903  SlEPllANl  denied  that  liverworts 
are  able  to  make  long  and  successful  jumps.  Attention  should  be  paid  to  Domin's 
valuable  j)aper  [yd]  in  which  he  brings  together  numerous  facts  illustrating  the  same 
definite  distribution  patterns  in  this  as  in  other  groups,  and  Miss  FULFORl)  (e.g. 
10  ;]  has  arrived  at  the  same  conclusion.  We  know  that  the  spores,  in  many  cases 
at  least,  are  extremely  sensitive  to  changed  conditions  and  lose  their  viability 
very  ra[)idly  when  exposed  to  the  air — a  promising  field  for  experiments.  Regard- 
ing mosses  I  refer  to  Herzog's  work  [i2g)  where  he  speaks  against  the  belief  in 
the  importance  and  great  range  of  dispersal  through  the  air.  We  find,  IRMSCIIER 
says,  the  same  disjunctions,  the  same  part  areas  (Teilareale)  in  angiosperms, 
gymnosperms,  ferns  and  bryophytes,  and  he  continues  p.  292  : 

Dass  diese  vier  in  ihren  Verbreitungsmitteln  so  verschiedenen  Pflanzengruppen  die- 
seli)cn  \'erhrcitungszuge  ihrer  Disjunktelemente  ergeben,  zeigt  wohl  einwandfrei,  dass 
hierfur  der  "Wind"  ebenso  wie  andere  iiussere  Krafte  als  Ursache  abzulehnen  sind. 
Wiiren  sic  in  aiisschlaggcbender  Weise  an  der  Ausbreitung  beteiligt,  miisste  die  Besie- 
dclung  entsj)rechend  der  Verschiedenheit  der  Friichte,  Samen  und  Sporen  ebenso  ver- 
schiedcnartig  ausgetallen  scin,  d.  h.  in  den  einzelnen  Gruppen  dieser  biologischen  Ver- 
schiedenheit entsprcchcnde  charakteristische  Merkmale  zeigen.  Dies  ist  aber  nicht  der 
i''all.  Der  alien  vicr  druppen  ge  m  e  i  n  sam  en  hochdisjunkten  Ausbildung  so  vieler  Areale 
nuiss   vichnehr  cine  andere    Ursache  zu   (irunde  liegcn  .  .  . 

This  conniion  cause  was,  in  Irmsciier's  opinion,  continental  displacement  in  the 
sense  of  Wi;(;ener.  Those  who  disagree  with  him  will  have  to  look  for  vertical 
movements,   emergence  and   submergence  of  land. 

Lichens,  fungi  etc.  were  not  included  in  Irmsc:her's  discussion.  A  survey  of 
their  distribution  patterns  is  something  to  be  asked  for.  Lichens  are  said  to  de- 
|)en(l  on   their  vegetative  reproduction   bodies  more  than   on  spores. 

It  is  maintained  that,  with  certain  exceptions,  terrestrial  animals  spread 
less  easily  than  plants.  1  have  consulted  a  number  of  zoogeographers  in  order  to 
learn  their  opinion  on  the  mode  of  transport  likely  to  be  used  by  invertebrates 
in  their  sup[K)sed  ocean   voyages.   Birds  etc.   are  left  aside  here. 

Numerous  insects,  butterflies,  moths,  flies,  hymenoptera,  grashoppers,  cock- 
roaches, as  well  as  spiders,  myriapods,  etc.  follow  man  from  land  to  land,  from 
island    to    island.    This  is,  I  daresay,  the  only  safe  way  for  such  animals  to  get 


DERIVATION  OF  THE  FLORA  AND  FAUNA  343 

abroad.  For  those,  and  they  are  in  overwhelming  majority,  which  are  not  an- 
thropochorous,  the  chance  to  cover  large  distances  ought  to  be  very  small.  How- 
ever, Zimmerman,  in  his  admirable  introduction  to  the  Insects  of  Hawaii,  surveys 
one  order  after  the  other  and  finds  nothing  that  speaks  against  his  belief  in  the 
permanent  isolation  of  the  Hawaiian  as  well  as  all  other  oceanic  islands,  and 
consequently  concludes  that,  man-borne  species  excepted,  all  the  ancestors  of  the 
Hawaiian  insects  were  carried  there  by  natural  agents.  Dispersal  with  the  aid  of 
birds  is  of  slight  importance,  but  he  remarks  that  sea  birds  nest  in  forests  on 
the  islands;  I  do  not  think  that  this  means  very  much,  because  they  are  stationary 
(comp.  above,  p.  337).  Marine  drift  is  probably  the  least  successful  of  all  meth- 
ods, he  thinks.  Thus,  the  bulk  of  the  insect  fauna  was  and  is  wind-borne,  a 
traffic  going  on  without  interruption.  Zimmerman  refers  to  experiments  clearly 
showing  that  both  winged  and  unwinged  insects  are  carried  by  air  currents  to 
great  heights,  14,000-15,000  feet  (p.  58).  These  are  largely  abnormal  conditions 
and  due  to  cyclonic  storms,  which  account  for  the  dispersal  all  over  the  mid- 
Pacific.  The  result  is,  as  expected,  a  disharmonious  fauna,  where  large  groups 
common  to  all  continents  are  lacking:  "they  have  been  eliminated  by  the  selective 
agents  of  oversea  dispersal".  It  would  be  interesting  to  know  why  all  represent- 
atives of  large  and  otherwise  widely  distributed  insect  groups  are  excluded  from 
the  passenger  list.  Besides,  would  not  disharmony  result  even  if  land  connections 
once  existed.^ — the  islands  have  remained  isolated  for  a  long  time,  perhaps 
millions  of  years,  while  migration,  favoured  by  climatic  and  edaphic  changes 
affecting  the  general  character  and  composition  of  the  vegetation  has  continued 
over  land  on  the  continents. 

HiNTON  [dj)  who,  as  we  have  seen,  was  opposed  against  both  continental 
drift  and  land  bridges,  believed  that  wind-borne  and  raft-borne  transportation 
across  the  oceans  must  have  been  of  common  occurrence.  If  this  be  so,  why  did 
no  snakes,  frogs  or  gymnosperm  cones  ever  get  aboard  the  rafts  and  arrive  at 
distant  islands.^  Rafts  formed  by  large,  uprooted  trees  are  observed  in  big  rivers 
like  the  Amazon  or  Orinoco,  and  I  guess  that  an  analysis  of  their  composition 
would  reveal  the  presence  of  a  rather  varied  fauna.  WALLACE'S  Boa  constrictor 
was  referred  to  above  (p.  335).  Snakes  and  giant  spiders  are  often  found  hidden 
in  banana  trunks  imported  to  Europe  from  tropical  America.  On  the  other  hand, 
the  chances  for  the  formation  of  substantial  rafts  are  small  within  the  tropical 
Pacific,  where  no  big  rivers  empty. 

W/iat  kind  of  invertebrates  are  likely  to  withstand  transoceariic  migration? 
HiNTON  has  an  answer  ready: 

The  chance  of  accidental  dispersal  varies  according  to  the  nature  of  the  group. 
Colonization  of  the  Hawaiian  and  other  islands,  always  far  removed  from  any  conti- 
nent, provides  us  with  absolute  proof  of  the  kind  of  animals  and  plants  that  can  with- 
stand long  distance  wind  or  raft  transportation  across  the  oceans. 

This  sounds  quite  simple,  but  really  is  a  very  complicated  problem,  to 
which  sufficient  attention  has  not  been  paid.  The  possibilities  vary  according  to 


344  C.  SKOTTSBERG 

size,  flight  capacity,  habits,  mode  of  reproduction,  sensitiveness  to  changes  of 
miUeu,  and  so  forth,  and  I  am  afraid  that  we  have  httle  knowledge,  founded  on 
facts,  in   most  cases. 

liarth-iroDus  have  been  carried  all  over  the  world  with  the  human  traffic. 
It  is  noteworthy  that  as  a  rule  no  truly  indigenous  species  are  found  on  oceanic 
islands,  where,  if  land  connections  had  existed,  they  could  be  expected,  and  even 
if  the  transport  of  eggs  or  living  animals  were  effected  only  by  means  of  rafts 
stocked  with  earth  and  plant  material,  as  some  believe,  they  ought  to  be  present, 
but,  as  far  as  I  am  aware,  nobody  has  witnessed  such  a  transport.  The  presence 
of  endemic  leeches  on  Samoa  and  Juan  Fernandez  (Masafuera)  can  be  understood 
only  if  the  leeches  are  carried  about  on  birds  acting  as  hosts,  otherwise  I  cannot 
see  how  they  would  be  able  to  survive;  they  are  very  sensitive  to  exposure.  To 
these  in  particular  I  should  like  to  apply  what  GULICK,  without  referring  to  any 
special  group  of  animals,  wrote  (z/^.  405): 

How  is  it  possible  at  all  for  creatures  that  would  die  almost  at  touch  of  sea  water  to 
precede  man  by  a  million  years  on  islands  standing  solitary  in  mid-ocean?  Are  their 
remote  homes  really  the  left-over  fragments  of  ancient  inter-continental  land  bridges,  or 
are  these  creatures  prima  facie  evidence  that  their  ancestors  possessed  an  almost  incon- 
ceivable capacity  for  passing  uninjured  over  vast  stretches  of  open  ocean?  The  extremes 
of  hypothesis  that  have  been  proposed  in  response  to  this  dilemma  show  us  how  difficult 
it  has  been  to  find  a  solution. 

Freshwater  criistaceaiis  occur  on  many  islands,  both  traffic-borne  and  indigenous 
species;   only  the  raft  theory  would  account  for  their  spread. 

It  is  supposed  that  cocoons  of  spiders  are  transported  by  wind,  webs  are 
torn  loose  with  cocoons  attached  and  carried  up  into  the  air,  where  a  storm  takes 
care  of  them.  Theoretically  this  is  not  impossible,  but  whether  the  contents  stand 
a  journey  of  thousands  of  miles  is  doubtful.  Adventitious  species  are  found  on 
Pacific  islands,  but  the  bulk  of  the  spider  faunas  is  indigenous  and  endemism  is 
high.  Hkki.anI),  pointing  to  the  general  distribution  of  genera  and  the  high  spe- 
cific endemism,   is  in   favour  of  former  land  connections  [2j.  1052). 

L'isolation  doit  etre  assez  ancien  pour  ce  que  cette  fauna  ait  pu  acquerir  les  carac- 
tcres  d'endemisme  (ju'elle  presente.  Un  botaniste  a  fixe  vers  le  Pliocene  cet  isolement, 
mais  jc  serais  porte  a  croire  (pi'il  est  i)lutc)t  ])lus  ancien,  en  me  basant  sur  la  lenteur 
de   revolution   des   Araignees. 

AI.\\R,   who  rejects  all   land  connections,  remarks  (27^.214): 

Considering  the  haj)ha7,ard  manner  by  which  these  oceanic  islands  receive  their  popu- 
lations, it  is  rather  astonishing  how  similar  the  faunas  of  the  various  islands  are.  Ber- 
i.ANi),  on  the  basis  of  the  distribution  of  spiders,  has  come  to  the  conclusion  that  the 
fauna  of  all  Polynesia  is  so  uniform  as  to  suggest  that  these  islands  are  but  fragments 
of  a  single  land  mass.  This  view  is  similar  to  Pilsbry's,  founded  on  Mollusca.  Actually, 
this  j)aradox  of  tlie  similarity  of  the  fiiunas  of  oceanic  islands  is  solved  in  quite  dif- 
ferent manner.  Of  all  the  j)Ossil)lc  families,  genera  and  species  of  the  Papuan  Region 
that  are  theoretically  in  a  position  to  colonize,  only  a  small  fraction  will  eventually 
avail   themselves  of  the  opportunity. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  345 

This  is  quite  true,  but  will  not  the  result  become  much  the  same  with  land 
connections?  The  main  source  of  the  fauna  is  the  same,  the  similarity  is  a  con- 
sequence, and  local  endemism  is  a  result  of  isolation. 

Acarids  present  the  same  problem.  MUMFORD  [183)  remarks  that  is  is  very 
difficult  to  compile  lists  of  species  for  the  Pacific  islands  and  that,  at  present, 
no  safe  conclusion  can  be  drawn  with  regard  to  the  distribution  of  genera.  This 
is  true,  but  we  know  that  numerous  indigenous  species  occur  and  that  local  en- 
demism seems  to  be  high.  How  these  extremely  delicate  little  animals  would  be 
able  to  stand  long  overseas  voyages  is  difficult  to  imagine.  They  are  plentiful  in 
humid  forest  soils  on  Juan  Fernandez  and  all  the  species  except  two  adventitious 
ones  are  endemic.  Little  is  known  of  their  relationships.  If  washed  down  from 
the  hill-sides  and  carried  out  into  the  sea,  they  will  die — only  wind  transport 
remains.  Pseudoscorpio7is  may  be  more  resistant  to  both  salt  water  and  desicca- 
tion, but  their  pronounced  endemism  bears  witness  of  long  isolation.  All  the  false 
scorpions  recorded  from  Juan  Fernandez  are  endemic,  and  there  is  one  endemic 
genus.    They  were  unknown  when  MuMFORD  (I.e.  246)  wrote: 

As  Chamberlain  points  out,  it  is  doubtful  whether  anything  like  true  insular  endemism 
occurs  in  most  species  of  false  scorpions  because  of  the  ease  with  which  they  are  distrib- 
uted. 

They  are  well  adapted  to  be  carried  about  by  man.  Among  the  Myria- 
poda  are  many  local  endemic  species.  These  creatures  are,  according  to  my  own 
experience,  rather  tough  and  might  be  able  to  spread  by  the  same  methods  as 
earth-worms  and  land  crustaceans.  It  is  not  very  probable  that  they  are  blown 
from  continents  to  distant  islands.  Cosmopolitan  forms  are  probably  adventitious. 
If  this  applies  also  to  Collembola  I  do  not  know;  very  wide-spread  species  may 
be  so  ancient  that  they  have  attained  their  distribution  when  the  map  of  the  globe 
was  quite  different  from  the  present  one.  It  is  unlikely  that  they  are  able  to 
migrate  overseas.  Lindsay  [i6y.  7 1 9)  writes : 

The  primitively  wingless  Collembola  seem  to  constitute  better  material  for  distribution 
studies  than  any  other  insect  order,  because  migration  by  flight  is  imp'ossible  and  the 
delicate  integument  makes  it  very  unlikely  that  the  insect  could  be  carried  any  appre- 
ciable distance  by  the  sea. 

We  can  safely  add  that,  if  blown  out  over  the  ocean,  they  would  soon  perish. 
Zimmerman  suspects  that  none  of  the  32  species  recorded  for  Hawaii  is  indige- 
nous there;  thus,  all  are  supposed  to  have  been  imported  by  the  traffic.  How- 
ever, 3  species  are  supposed  to  be  endemic  in  Juan  Fernandez.  Two  species  of 
the  Hawaiian  Thysanuva  are  "possibly  endemic"  (Zimmerman)  but  perhaps  adven- 
titious. These  creatures  do  not  appear  to  be  fit  for  long-distance  dispersal,  and 
the  two  species  known  from  Juan  Fernandez  are  endemic.  One  of  them  belongs 
to  a  monotypical  genus  with  Australian  affinities,  a  disjunction  not  easily  bridged 
over  without  land  connections. 

Getting  to  the  true  insects,  their  mode  of  dispersal  certainly  varies  a  great 
deal.  Whereas  butterflies,  moths,  flies,  hymenoptera  etc.  are  known  to  be  storm- 
driven  and  eventually  carried   far,  heavy  beetles,  even  if  they  be  properly  winged. 


346  C.   SKOTTSBERG 

are  unfit  for  long  journeys;  whether  they  are  transported  by  waves  and  currents  and 
able  to  stand  immersion  in  salt  water  during  weeks  and  months  I  do  not  know. 
The  endemic  Sightless  insects  have,  just  as  the  flightless  birds,  given  cause  to 
much  s[)eculati()n.  They  are  supposed  to  descend  from  winged  species;  arrived,  it 
is  said,  on  an  oceanic  island,  they  had  the  choice  of  losing  their  wings  or  being 
blown  oft'  the  island  and  lost  altogether.  ZIMMERMAN  does  not  favour  this  view, 
for  they  may  as  well  have  lost  their  power  of  flight  on  the  mother  continent, 
which  did  not  prevent  them  to  be  carried  off  by  a  hurricane.  Mayr  agrees  with 
him,  comp.  quotation  p.  336.  Very  well,  but  is  it  possible  to  imagine  a  flightless 
rail  carried  a  thousand  miles  across  an  ocean.' 

There  are  numerous  Ortlioptcra  on  Hawaii,  most  of  them  endemic  species  or 
even  belonging  to  endemic  genera.  So  far  only  4  species  have  been  reported  from 
Juan  Fernandez,  two  of  them  endemic.  I  presume  that  Zimmerman  regards  these 
insects  as  normally  wind-borne.  The  relations  of  the  Hawaiian  Gryllidae  are  with 
Indo  Pacific  forms,  so  they  have  had  a  long  way  to  go.  The  four  species  of 
tciDiitcs  found  in  Hawaii  are  adventitious,  which  should  indicate  that  they  are 
unable  to  reach  oceanic  islands  without  human  assistance,  but  the  single  species 
discovered  in  Juan  Fernandez  (Masatierra)  is  endemic.  Either  did  it,  or  its  ancestor, 
arrive  over  land,  or  a  colony  was  carried  in  a  floating  log,  which  may  seem  un- 
likely, or  a  storm  brought  a  winged,  swarming  couple  which  founded  a  new  colony 
— even  less  probable.  The  termites  are  a  very  ancient  order  and  date,  it  is  said, 
back  to  the  Mesozoic  at  least. 

Mallophaga  are  spread  with  their  bird  hosts.  All  the  endemic  Hawaiian  species 
live  on  the  Drepanididae,  and  the  marine  birds  here  and  on  Juan  Fernandez  are 
infested  with  widespread  forms.  Tliysanoptera  seem  to  be  easily' spread  with  human 
traffic.  Most  of  the  90  species  found  in  Hawaii  are  adventitious,  few  indigenous. 
Two  of  the  4  species  in  Juan  h'ernandez  are  endemic.  How  these  delicate  insects 
manage  to  get  about  and  to  reach  oceanic  islands  I  cannot  tell. 

Tlie  Hawaiian  islands  have  a  rich  and  peculiar  fauna  of  Xcuroptera,  some 
"among  the  most  aberrant  of  all"  (ZIMMERMAN  p.  "j6).  Of  the  five  species  re- 
corded for  Juan  Fernandez  4  are  endemic,  one  of  them  belonging  to  an  endemic 
genus.  Wind  drift  must  be  taken  into  account,  but  it  cannot  be  very  effective.  The 
situation  remains  tlie  same  when  we  get  to  the  Lepidoptera,  about  lOOO  species 
in  Hawaii,  of  which  85  %  have  not  been  found  elsewhere.  Wind  drift  or  immature 
stages  (eggs,  etc.)  carried  with  plant  material  are  the  only  possibilities,  but  they 
cannot  be  very  great.  vSo  far  only  26  indigenous  species  are  known  from  Juan 
I^'ernandez,  70%  of  them  endemic;  when  the  list  of  Dr.  Kusciiel's  collection,  which 
contains  over  50  species,  has  been  published,  the  figures  will  undergo  alteration. 
In  his  survey  of  the  Pacific  lepidoptera  Swezev  [262)  includes  the  Galapagos 
Islands,  but  does  not  mention  Juan  Fernandez.  The  Pacific  islands  were,  he  says 
p.  319,  j)opulated  from  the  Malayan  and  Oriental  regions,  and  the  fauna  arrived 
in  the  main  by  accident,  winds,  ty[)hoons  etc,  or  with  plant  material  brought  by 
currents.  A  comparable  numerical  development  of  species  per  genus  has  taken 
place  in  no  other  islands  than  the  Hawaiian,  and  the  author  infers  that  they  have 
a  more    ancient  fauna,  descended  from  ancestors  that  arrived  at  a  more  remote 


DERIVATION  OF  THE  FLORA  AND  FAUNA  347 

time.  This  does  not  accord  with  Zimmerman's  statement  that  no  primitive  family 
is  represented  in  Hawaii.  Probably  the  Hawaiian  Diptera  have  been  less  collected; 
Zimmerman,  who  is  responsible  for  the  figures  quoted,  indicates  about  400  species, 
of  which  60  %  are  endemic.  In  relation  to  their  small  size  the  Juan  Fernandez 
Islands  seem  to  be  richer  with  about  i  50  indigenous  species  (64  %  endemic).  Our 
knowledge  of  the  dipterous  fauna  is  in  the  main  due  to  Dr.  KuscilEl/s  collec- 
tions; the  25—30  species  not  yet  reported  on  will  raise  the  total  number. 

Over  1600  species  of  Colcopiera  are  reported  from  Hawaii,  of  which  about 
'j^  %  are  endemic,  and  there  are  numerous  endemic  genera.  Numerous  species 
found  elsewhere  are  adventitious.  In  relation  to  its  size,  Juan  Fernandez  cannot 
be  called  poor  (see  above  p.  307,  etc.),  and  endemism  is  just  as  high  here.  I  presume 
that  Zimmerman  and  others  regard  wind  as  the  principal  dispersal  agent,  though 
not  for  all  kinds  of  beetles,  because  we  have  to  do  with  many  different  types 
of  animals  and  of  habits  and  habitats.  It  is  difficult  to  imagine  how  a  flightless 
beetle  would  be  able  to  keep  afloat  in  the  air  for  thousands  of  miles;  he  must 
have  had  a  great  need  of  the  numerous  "stepping  stones"  postulated  by  ZIMMER- 
MAN. It  should  perhaps  be  mentioned  that  GULICK  (77^.414)  wrote  that  "it  can 
hardly  be  doubted  that  some  carrion-feeding  insects  have  been  distributed  by 
adhering  to  sea  birds".  I  doubt  that  this  ever  happened,  but  as  he  had  just  dis- 
cussed the  transport  of  "invertebrate  eggs"  in  the  digestive  tract  of  birds,  I  sup- 
pose that  he  means  eggs  of  necrophilous  flies  or  beetles  which,  brought  across 
the  sea,  were  deposited  on  another  carrion  and  thus  became  established  on  an 
oceanic  island. 

The  taeevils,  a  most  important  and  interesting  feature  in  isolated  island 
faunas,  are  often  dependent  on  definite  host  plants,  and  are  thought  to  sail  along 
on  logs  as  stowaways.  Evidently  the  Curculionidae  have  become  something  of  a 
stumbling-block.  UsiNGER  (2yj)  came  to  the  conclusion  that  they  must  possess 
some  unknown  special  means  of  dispersal.  Brinck,  who  discussed  the  coleopterous 
fauna  of  Tristan  da  Cunha  (316.  97—104),  another  isolated  volcanic  group  of  islands 
where  geologists  failed  to  discover  any  traces  of  land  connections,  states  that 
the  fauna  contains  endemic  elements  and  ofl"ers  examples  of  remarkable  disjunc- 
tions. It  must  have  originated  from  extinct  faunas  of  neighbouring  continents  if 
not  of  submerged  lands.  The  only  natural  agent  capable  of  transportation  is  the 
wind,  but  Brinck  is  convinced  that  "at  present  no  beetles  are  invading  the  islands 
by  natural  means"  (p.  103),  and  the  reason  is  not  adverse  conditions,  for  several 
species,  introduced  with  the  human  traffic,  have  become  naturalized.  The  unavoid- 
able conclusion  is  that  dispersal  agents,  man  excepted,  are  insufficient — and  they 
were  the  same  in  the  past.  The  original  beetle  fauna  has  survived  from  pregla- 
cial  time,  an  hypothesis  that  nobody  would  feel  inclined  to  reject,  but  it  does 
not  help  us  to  understand  by  what  means  it  was  able  ever  to  arrive.  The  weevils 
of  Hawaii  have  had  a  long  way  to  come;  according  to  ZIMMERMAN  the  ancestors 
as  a  rule  came  from  the  south  Pacific  or  Indo-Pacific  regions.  Generally  he  regards 
also  the  peculiar  genera  to  have  originated  in  Hawaii  or,  eventually,  in  one  of 
the  lost  islands  serving  as  intermediate  stations,  but  among  the  weevils  are  some 
that  defy  all  explanations: 


348  C.   SKOTTSBERG 

Xt'sotflcus  is  evidently  a  relict  endemic  genus  of  four  closely  allied  species,  and 
there  aj)pears  to  be  nothing  like  it  elsewhere  .  .  .  Oodemus  with  its  58  species  ...  is 
the  largest  genus  of  the  Hawaiian  Cossoninae  .  .  .  together  with  its  close  ally  Atiotherus 
(3  species)  endemic,  and  I  know  of  no  genus  or  group  of  genera  from  any  region  from 
which   is   might   have   come.    It   is   an   anomaly. 

Such  cases  brini^  us  back  to  times  long  before  the  formation  of  the  present 
Hawaiian  chain,  and  similar  cases  are  found  also  in  Juan  Fernandez,  e.g.  the 
endemic  tribe  yidviorJuui  of  .Vl  KI\ILL11  s.  Other  examples  are  offered  by  many 
other  insect  groups.  And,  leaving  them  aside  for  a  moment,  is  not  the  endemic 
Hawaiian  bird  family  Drepanididae  another  anomaly.^  Whereas  Brvan  [j^g.  188) 
finds  a   Malayan  origin   most  acceptable,  GULICK  writes  (7/^.420): 

The  history  of  Hawaiian  land  birds  must  have  begun  with  the  arrival  of  some  form 
of  troi)i(al  American  honey  creeper,  which  became  in  due  time  the  progenitor  of  all  18 
genera   and   40   sj)ecies   of  the    Drepanididae. 

Me  seems  to  have  forgotten  that  the  islands  are  claimed  to  have  cHved  out  of 
the  ocean  in  late  Pliocene  and  Pleistocene  times. 

I lyinoioptcra  are  plentiful  in  Hawaii,  about  600  native  species,  among  which 
endemics  are  numerous,  and  the  ancestors  are  supposed  to  have  come  from  the 
south  and  southwest  Pacific,  in  exceptional  cases  from  Asia  and  the  Orient.  This 
order  is  as  yet  little  known  in  Juan  Fernandez,  see  above  p.  315.  Wind  drift  seems 
to  be  the  only  possible  mode  of  transport  unless  infected  larvae  of  butterflies  etc. 
arrived  with  drift-wood,  which  does  not  seem  very  probable.  Ants  are  easily  spread 
with  the  traffic.  Of  the  3  species  known  in  Juan  Fernandez  only  one  appears  to 
have  arrived   without  human  assistance. 

Among  the  Heteroptera  in  Hawaii,  over  200  species  and  80  %  endemic,  ZlM- 
MKRMAN  pays  special  attention  to  the  genus  Xysius,  which  has  its  greatest  known 
diversity  in  those  islands.  All  are  endemic  and  include  the  most  divergent  of  all 
Xysius  species.  The  ancestors  are  supposed  to  have  come  from  the  south  and 
west  Pacific.  The  genus  is,  according  to  UsiNGER  (2'/j)  common  in  the  Australian 
and  Oriental  regions,  extending  through  Melanesia  to  Fiji  anci  Samoa  without  a 
single  re[)resentative  east  of  this  line,  but  important  in  the  Hawaiian  chain;  the 
author  seems  to  have  overlooked  its  occurrence  in  Juan  Fernandez.  Xysius  is 
su()posed  to  have  reached  Hawaii  by  a  circuitous  route  over  open  water  and  the 
Leeward  Hawaiian  islands.  This  route  is  indicated  by  a  submarine  ridge  of  con- 
siderable depth  and  may  once  have  been  interrupted  by  island  peaks  such  as 
Wake  Island;  thence  it  is  followed  to  the  Marianas  and  Caroline  Islands  and  even- 
tually to  the  rich  Papuan  and  Australian  regions,  l^ut  in  other  cases  it  is  less 
eas)-  to  construct  a  suitable  route:  "The  presence  of  twenty  very  unique  genera 
in  Hawaii  and  their  absence  from  old,  high  islands  along  the  very  route  they  are 
said  to  have  traveled  is  inexi)lical)le  by  present  theories"  (I.e.  315).  Why  not 
presume  that  all   related  genera  have  died  out.-,  which  is  the  easiest  explanation. 

Xysius  is  a  widespread  genus,  well  developed  also  in  New  Zealand,  and  the 
single  Juan  I^'ernandez  species  is  claimed  to  be  related  to  another  from  New  Zea- 
land. We  know  numerous  examples  of  the  same  kind  in  other  animal  groups  and 
particularly  among  the  plants  we   have  called  Antarcto-tertiary. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  349 

The  Homoptera  present  the  same  picture  in  Hawaii,  about  350  native  spe- 
cies, most  of  them  endemic  and  suggesting  a  southwest  Pacific  ancestry.  Of  the 
species  reported  from  Juan  Fernandez  only  one-third  have  been  described,  all 
endemic  and  as  far  as  we  know  with  their  relatives  in  the  south  and  central  Pacific. 

No  order  characteristic  of  insular  faunas  has  aroused  greater  interest  among 
biogeographers  than  the  Pacific  larid  molluscs,  nowhere  more  wonderfully  devel- 
oped than  in  the  Hawaiian  chain.  In  the  discussion  of  land  connections  they 
occupy  a  central  position. 

Wallace  found  that  the  wide  distribution  of  the  land  snails  is  "by  no 
means  so  easy  to  explain  as  that  of  the  insects";  the  chances  have  been  "rare 
and  exceptional",  possibly  eggs  stuck  to  the  feet  of  aquatic  birds,  or  the  animals 
themselves  were  storm-carried,  "attached  to  leaves  and  twigs" — this  would  be 
the  only  means  by  which  viviparous  forms  could  be  transported. 

GULICK  [iig]  treats  the  land-shell  problem  at  some  length.  Speaking  of  Easter 
Island  he  asserts  that  "hurricanes  spread  gravid  land-snails  as  dust  over  almost 
as  great  distance  as  plant  seeds  can  be  blown",  but  he  gives  no  facts  to  support 
this  very  positive  statement.  The  land-shells  of  Juan  Fernandez  and  Saint  Helena 
are  then  remarked  upon. 

At  least  three  elements  can  have  derived  their  ancestry  only  from  Polynesia,  fully 
3400  miles  away,   unless  Easter  Island  served  as  a  way  station. 

The  archaic  complexion  of  the  snail  fauna  is  not  necessarily  very  significant,  as 
younger  continental  forms  do  not  for  the  most  part  yield  minute,  easily  wind-blown 
species. 

Archaic — exactly,  malacologists  emphasize  that  more  modern  types  do  not  occur 
on  oceanic  islands  unless  brought  by  man,  and  this  has,  naturally  enough,  been 
used  as  an  argument  in  favour  of  early  land  connections  before  the  modern  types 
existed.  In  Gulick's  view  the  size,  not  the  age,  decides. 

Large,  softskinned  creatures  invariably  make  a  poor  showing  .  .  .  large  helices  ex- 
amplify  this  disability  so  excellently  that  their  failure  to  arrive  is  a  sort  of  negative 
criterium  for  insularity  (p.   414). 

But  neither  are  all  continental  species  large,  nor  all  insular  ones  small,  and  the 
oceanic  snail  fauna  includes  many  forms  that  cannot  spread  like  dust.  GULiCK 
remarks  on  the  genus  Partula  that  "its  120  geographically  restricted  species  mostly 
weigh  too  much  and  are  too  tender  to  fit  easily  into  theories  of  transport  by 
air  or  sea",  and  this  makes  him  take  into  account  the  possibilities  of  "land  ridges" 
to  facilitate  transport. 

"It  is  evident",  he  says  p.  419,  "that  the  vast  diversification  is  a  proof  of 
the  great  local  antiquity  of  these  families,  and  hence  of  a  considerable  antiquity 
of  their  island  habitat."  This  is  not  true  of  the  islands  as  they  appear  now^  and 
as  they  have  stood  for  probably  millions  of  years. 

Land  snails,  just  as  weevils,  are  dispersed  by  some  unknown  method,  UsiN- 
GER  thinks  (I.e.  315),  while  Mayr,  as  already  quoted,  regarded  the  presence  of 
small  molluscs  on  oceanic  islands  as  a  proof  of  the  efficiency  of  hurricanes,  but 
later  on  expressed  himself  as  follows. 


350 


C.   SKOTTSBERG 


It  seems  to  me  that  the  wide  acceptance  of  land  bridges  by  conchologists  is  chiefly 
due  to  three  reasons:  (i)  our  almost  complete  ignorance  of  the  means  of  disi)ersal  of 
snails,  (2)  our  lack  of  knowledge  of  the  speed  of  speciation  in  snails,  and  (3)  faulty  classi- 
fication, i)articularly  generic  classification.  A.  Gulick  has  already  directed  attention  to  the 
])resence  of  snails  on  most  oceanic  islands.  'J'hey  were  unquestionably  carried  there  by 
some  unknown  means  of  transjiortation.  .  .  .  To  me  it  seems  incomparably  simpler  to 
assume  a  still  unknown  method  of  transportation  than  a  land  bridge  that  is  unsupported 
by  any  other  fact. 

Simple,  no  doubt,  but  we  cannot  get  away  from  the  problem  by  an  "ignoramus". 

It  is  eas\'  to  under.stand  that  the  presence  of  fairly  large  forms  of  land-shells 
on  distant  islands  has  caused  a  good  deal  of  trouble.  ZIMMERMAN  tried  to  find 
a  wa\'   out  of  the  difficulties. 

It  has  been  said  that  large  snails  such  at  the  Hawaiian  achatinellids  and  amastrids 
are  particularly  unsuited  for  overseas  distribution.  However,  if  we  approach  the  problem 
differentlv,  different  conclusions  may  be  reached.  If,  as  I  believe,  the  large  Hawaiian 
snails  have  evolved  from  small  or  minute  ancestors,  then  the  argument  based  upon 
their  large  si/e  loses  its  weight.  However,  if  small  snails  can  be  distributed  overseas, 
then  what  is  to  j)revent  eggs  or  tiny,  immature  specimens  of  large  species  from  being 
similarly   transported?   (I.e.  61). 

In  passing,  ZiMMKRMAX  quotes  H.  B.  13.\KKR,  who  thought  that  land-shells 
are  carried  along  by  migratory  birds.  After  these  speculations  it  is  refreshing  to 
read  Hrvax  (^o.  9): 

The  j)resence  of  certain  kinds  of  plants  and  animals  found  in  Hawaii  and  related 
to  species  in  the  southwest  Pacific  is  hard  to  explain  by  any  known  means  of  drift, 
either  over  the  sea  or  through  the  air.  Land  snails  constitute  one  such  group.  These 
mollusks,  whi(  h  breathe  directly  from  the  air,  would  drown  in  water,  particularly  salt 
water;  yet  they  must  have  moisture.  They  cannot  stand  long  exposure  to  the  sun,  but 
live  on  the  leaves  and  trunks  of  forest  ])lants  or  beneath  fallen  leaves  and  trash  on  the 
ground.  How  did  their  ancestors  reach  Hawaii  if  they  could  neither  swim  nor  drift? 

( ^f^poju'iits  ai^^aiiist  the  doi^iua  of  colonization  across  tJic  oceans. 

Many  biogeographcrs  have  arrived  at  the  conclusion  that  the  natural  dispersal 
agents  cannot  be  made  responsible  for  the  distribution  of  all  kinds  of  biota  across 
very  wide  expanses  of  open  water.  Most  of  the  authors  are  botanists,  which  is  sur- 
prising because  the  chances  should  be  greater  for  seeds  and  spores  than  for  eggs 
or  individuals  of  delicate  creatures.  To  ask  for  land  bridges,  or  for  extension  of 
continental  margins  later  submerged  l)ut  leaving  behind  land  fragments,  is  to  refuse 
to  accept  overseas  migration  as  the  only  possible  means  of  colonization.  Conse- 
(piently,  a  number  of  authors  have  already  been  quoted  in  the  chapter  on  the 
history  of  the  Pacific  basin,  (iooD,  Ikmsciikk,  WTij-f,  Cami'BELL,  etc.,  as  well 
as  my  own  contributions  to  the  discussion,  the  latest  in  195 1  [2^8).  I  shall  add 
here  that  I  never  disclaimed  every  possibility  of  migration  over  wide  expanses 
of  water;  see  226  and  2:12.  X'arious  writers  have  compared  isolated  peaks  on 
continents  with  oceanic  islands  and  have  stated  that  under  present  conditions  an 
exchange  of  biota  is  improbable.  \\\x  SlKKMS  expressed  his  opinion  when  dealing 
with    the    Malaysian    mountain    flora,    in    very  plain  terms  [2^8)\   too  many  facts 


DERIVATION  OF  THE  FLORA  AND  FAUNA  35I 

show  the  impossibility  of  attributing  any  importance  worth  mentioning  to  long- 
distance dispersal;  the  theory  is,  when  it  comes  to  migrations  of  floras,  "not 
worth  a  straw",  and  Reiche  (20j),  referring  to  the  history  of  the  Chilean  flora, 
called  it  "eine  Kette  von  Unwahrscheinlichkeiten".  Gordon  (iij)  was  strongly 
inclined  do  deny  ist  value  and  believed  that  plants,  and  more  so  plant  associa- 
tions, advance  slowly  over  land  and  do  not  jump  thousands  of  miles;  he  could 
not,  however,  help  paying  some  attention  to  "Nature's  great  Krakatau  experi- 
ment", of  which  enough  has  been  said. 

Cain  (42)  thinks  that  "migration  is  usually  not  a  random  matter"  (p.  162) 
and  that  long-distance  dispersal  rarely  has  resulted  in  migration  and  establishment, 
nor  does  it  explain  the  discontinuous  areas.  In  reality  he  belongs  to  the  trust- 
worthy opponents.  One  of  the  reasons  advanced  by  him  is,  however,  not  con- 
clusive: 

The  phenomenon  of  local  races  (subspecific  endemics)  is  entirely  opposed  to  the 
idea  of  long-distance  dispersal,  for  such  variation  depends  upon  isolation  which  would 
not  exist  if  long-distance  dispersal  were  generally  effective  (p.  161). 

Those  who  are  in  favour  of  the  theory  emphasize  the  haphazard  character 
of  the  procedure;  success  may  follow  once  and  never  more.  If  the  immigrant 
belongs  to  a  polymorphous  species,  with  intermediate  forms  between  the  sub- 
species or  varieties,  it  may,  isolated  as  it  remains  from  the  rest  of  the  population, 
stand  out  as  a  separate  insular  taxon. 

Among  the  zoologists  few,  mainly  malacologists,  are  in  favour  of  land  con- 
nections, but  some  entomologists  agree  with  them.  Berland  was  already  quoted. 
As  a  rule  a  connection  South  America-Antarctica-Australia  (or  New  Zealand) 
is  asked  for,  but  to  Enderlein  this  bridge  was  not  sufficient: 

Die  zahlreichen  endemischen  Gattungen  (9)  und  Arten  (2)  zeigen,  dass  die  Juan 
Fernandez-Inseln  ein  Refugium  fiir  die  Reste  der  Faunen  unfangreicherer  untergegan- 
gener  Gebiete  darstellen,  die  nicht  mit  dem  neotropischen  Gebiet  in  Verbindung  gestan- 
den  haben  (Enderlein  S4.  643). 

Did  Enderlein  dream  of  a  submerged  Pacific  continent.?  Many  of  the  island 
flies  are  also  native  of  the  mainland  of  Chile,  others  have  their  relations  there, 
and  if  the  islands  were  formerly  connected  with  some  other  land,  it  was  with 
South  America.  Besides,  the  majority  of  Enderlein's  new  genera  have  been 
reduced  to  synonymy  by  later  authors,  his  speculations  have  little  weight,  and 
Wygodzinsky  (2^^.  81),  referring  to  Gigantodax  kiiscJieli,  arrived  at  a  different 
conclusion.    Gigaiitodax  is  an  exclusively  South  American  genus. 

Chapter  VII. 
Biological  characteristics  of  isolated  islands. 

In  his  classical  "Lecture  on  insular  floras"  [138)  J.  D.  Hooker  formulated, 
in  very  lucid  terms,  the  special  features  of  island  floras  and  his  opinion  on  their 
evolution.  As  examples  he  chose  Macaronesia,  St.  Helena,  Ascension  and  Ker- 
guelen  Island.  Their  peculiarities  were  stated  under  five  items. 


352 


C.   SKOTTSBKRG 


1.  /;/  all  casts  ciDisidired  flcrislic  relaliois  exist  beiwcen  the  island  and  one 
niot/ier  coiitinejit. 

2.  rite  floras  of  all  the  islands  in  question  are  more  temperate  in  character 
tha)i  that  of  the  mother  cojitiiioit  on  the  same  latitude. 

This  ina\'  be  true  in  most  cases  thanks  to  the  influence  of  the  surrounding 
ocean;   it  holds  <;o()tl   for  Juan   P'ernandez. 

3.  All  these  islands  show  many  biological  peculiarities  by  which  they  are 
distifii^uished. 

Tlie  chstin<^uishino  characteristics  are  mainly  expressed  in  endemism.  HoOKER 
referred  the  etidemics  to  two  cate<^()ries,  such  as  do  not  show  affinity  to  the  plants 
on  the  mother  continent,  and  such  as,  even  if  belongini^  to  endemic  genera,  are 
related  to  continental  ones. 

If  we  turn  to  the  Pacific  where,  for  obvious  reasons,  only  the  high  volcanic, 
well  watered  islands  are  considered,  we  find  that  all  of  them  are  distinguished  by 
numerous,  in  man\'  cases  also  very  remarkable  endemics.  Hawaii  stand  out  above 
the  others  and  so  do  Juan  Fernandez  and  Desventuradas;  the  floras  of  Tahiti, 
Samoa,  Marcjuesas  etc.,  as  well  as  of  Micronesia,  are  less  independent.  The  degree 
of  s|)atial  isolation  is  not  conclusive;  the  flora  of  Juan  Fernandez  is  more  peculiar 
than  that  of  Marquesas  which  are  situated  much  farther  away  from  any  continent, 
and  this  holds  good  not  only  for  the  angiosperms  but  also  for  the  ferns.  F^n- 
(lemism  in  angi()sj)erms  is  69%  in  the  former  and  50  in  the  latter;  of  the  ferns 
about   30  "o   in   each,   but  only  Juan   P^rnandez  has  an  endemic  genus. 

4.  the  i!;e}ieral  rule  is  that  the  species  also  fotaid  in  the  mother  continent  are 
the  most  abitndant,  the  peculiar  species  are  rarer,  the  peculiar  j^enera  of  contineiital 
affinity  rarer  stilf  but  the  plajits  with  710  affijiity  elseivJiere  are  often  i>ery  commoii. 

This  is,  I  supj)ose,  true  of  the  islands  examined  by  HOOKKR,  with  the 
excej)tion  of  St.  Helena  before  the  arrival  of  man,  but  not  of  Hawaii,  nor  of 
Juan  l''ernandez.  Some  of  the  continental  species — Libertia  and  a  few  grasses — 
are  abundant,  while  others  are  rare,  all  according  to  the  supply  of  suitable 
habitats;  atuong  the  j)cculiar  species  of  continental  affinity  are  many  quite 
common  ones,  e.g.  the  endemic  species  of  Acaena,  Driniys,  Dysopsis,  Es- 
callojiia,  (iuniicra,  Myrceui^enia  (the  leading  forest  tree  on  Masafuera),  Pernettya, 
Rhafhithannius,  (  ]<^n/\  flrijrcroji  fruticosus,  Uncijiia  l)ouglasii\  and  of  the  peculiar 
genera  allied  to  South  yVmerican  ones,  Xot/iomyrcia  is  the  leading  forest  tree 
on  Masatierra,  where  Ochaga/da  is  also  common.  To  these  may  be  added  such 
common  endemics  as  lioehmeria  and  the  species  of /v/^^'-<'7;'rt:  and  Cop rosm a,  unde^v 
the  assumption  that  related  species  of  lloehnieria,  and  re{)resentatives  of  Coprosma 
and  i-agara,  once  belonged  to  the  neotropical  flora.  The  plants  of  no  affinity  in 
the  luother  countr\-,  that  is  the  nearest  continent,  are  as  a  rule  very  local,  few 
are  common   and   many  extremely  rare. 

Fossil)])-  I  have  misunderstood  llooKKK  here.  When  we  say  that  a  species 
is  abundant  in  a  country  we  mean  that  it  is  copious;  if  we  call  it  common,  it 
is  widely  spread;  if  we  call  it  rare,  it  has  been  rejjorted  from  a  small  number 
of  localities  only;  and  if  we  use  "sj)ecies"  in  plural  we  mean  the  same  thing. 
Hut  what  if  HooKHK   with   "abundant"   and   "common"   wanted  to  say  that  these 


DERIVATION  OF  THE  FLORA  AND  FAUNA  353 

Species  were  the  most  numerous  in  the  flora,  and  that  the  "rare"  ones  were  few 
in  number?  If  this  was  what  HoOKER  meant,  the  result  will  be:  Species  also  found 
on  the  continent:  46;  endemic  species  allied  to  South  American  or  other  conti- 
nental species:  71;  endemic  genera,  related  to  continental  genera:  5  (6  species); 
endemic  genera  not  related  to  continental  ones:  12  (24  species).  Still,  this  is  per- 
haps to  give  a  wrong  interpretation  to  Hooker's  words.  There  are  very  few 
native  non-endemics  on  St.  Helena,  and  they  are  of  course  not  in  the  majority 
in  the  Canary  Islands,  nor  are,  in  the  latter  place,  the  isolated  endemic  genera 
and  species  very  many.  In  this  case  HoOKER  says  "plants",  not  "genera"  and 
"species".  If,  to  this  group,  as  represented  in  Juan  Fernandez,  we  add  isolated 
endemic  species,  not  related  to  any  species  in  the  mother  continent,  although  the 
genera  occur  there,  this  group  comprises  54  plants,  or  more  than  ^/g  of  the  angio- 
sperms,  while  group  2  is  reduced  from  71  to  40.  The  figures  would  be:  46,  40, 
6,   54,  thus  conforming  much  better  to  Hooker's  rule. 

5.  Indigenous  annual  plants  are  extremely  rare  or  altogether  abse7it. 

Here  "rare"  must  mean  few  species,  and  Therophytes  are  very  few  in  Juan 
Fernandez  and  some  of  the  registered  species  perhaps  not  originally  native. 

How  were  plants  transported  to  distant  islands.^^  Hooker's  answer  is:  either 
across  the  sea  or  over  submerged  bridges,  and  he  adds:  "the  naturalist,  who  takes 
nothing  for  granted,  finds  insuperable  obstacles  to  the  ready  acceptance  of  either". 
The  situation  is  still  the  same  90  years  after  Hooker. 

Hooker  regarded  the  isolated  island  plants  as  "relics  of  a  far  more  ancient 
vegetation  than  now  prevails  on  the  mother  continent",  but  he  most  certainly 
never  wanted  to  say  that  the  continental  flora  was  altogether  younger  than  the 
insular  but  that  species,  now  restricted  to  the  island,  formerly  occurred  on  the 
mother  continent,  having  become  replaced  there  by  younger  species.  He  based 
his  opinion  on  the  fact  that  Macaronesian  relicts  had  been  found  as  fossils  in 
Tertiary  deposits  on  the  continent.  Time  has  not  permitted  me  to  collect  modern 
data,  and  I  can  only  suppose  that  some  of  the  old  determinations  still  hold  good. 
The  vegetation  of  Europe  has  undergone  great  changes  "within  the  lifetime  of 
these  Atlantic  island  species";  they  once  grew  in  Europe,  but  were  driven  out 
from  there  to  be  preserved  on  the  islands,  which  they  had  reached  "when  condi- 
tions may  have  been  very  different  from  what  they  are  now". 

The  theory  of  a  continental  Macaronesia,  including  Madeira  and  the  Azores, 
goes  back  to  Forbes'  theory  of  the  former  connection  between  the  British  Islands 
and  the  mainland,  definitely  proved  ages  ago.  FORBES  went  further  and  revived 
the  old  idea  of  a  lost  Atlantis,  still  favoured  by  many. 

It  is  interesting  to  follow  Hooker's  discussion  with  his  friend  Darwin  on 
island  problems.  Darwin  believed  in  the  efficiency  of  dispersal  agents  to  carry 
plants  and  animals  across  wide  expanses  of  sea,  and  his  arguments  made  such 
a  deep  impression  on  HoOKER  that  he  became  almost  convinced.  Still,  he  hesi- 
tated, and  certain  serious  difficulties  prevented  him  from  fully  accepting  Darwin's 
ideas.  The  composition  of  the  flora  of  the  Azores  was  not  what  we  had  reason 
to  expect  from  the  direction  of  winds  and  currents.  The  Macaronesian  Ornis  is 
almost  the  same  as  in  Europe  and  undoubtedly  came  from  there,  but  the  flora 

23  -  557857   The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.    Vol.  I 


354 


SKOTTSBERG 


is  considerably  diftcrcnt;  thus  it  may  be  argued  that  "the  birds  and  plants  do 
not  come  under  the  same  category".  Darwin  replied  that 

the  migration  of  birds  is  continuous  and  frecjuent,  and  the  individuals  surviving  and 
breeding,  they  keej)  uj)  the  specific  type,  and  do  not  give  origin  to  local  varieties; 
whilst  the  transport  of  seeds  is  casual  and  rare,  and  very  few  surviving,  these  not  being 
crossed  by  the  original  stock,  in  the  process  of  time  give  rise  to  varieties,  etc.,  and 
do   not   j)crj)etuate  the  continental   races  (p.    lo). 

This  is  the  situation  in  a  nut-shell,  and  Darwin's  arguments  are  repeated  by 
scores  of  biogeographers  to  this  very  day. 

Also  St.   Helena,  Ascension  and  Kerguelen  made  HoOKER  hesitate: 

They  St.  Helena  and  Ascension]  have  no  land  birds,  but  an  African  vegetation; 
and  though  nearly  midway  between  Africa  and  America,  they  have  scarcely  a  single 
American  tvpe  of  flowering  ]:)lants;  and  Kerguelen's  Land  has  a  flora  of  whose  ele- 
ments  most  have  emigrated  not  from  the  nearest  land,   but  from  the  most  distant  (p.  lo). 

11i:msi,i:v  [12'j.  59)  remarks  that  Hooker  seems  to  have  forgotten  the 
Compositae  in  St.  Helena,  most  of  them  showing  American  affinity. 

Kerguelen's  nearest  land  is  Antarctica,  but  not  a  single  flowering  plant  is 
known  from  the  coast  south  of  Kerguelen.  Africa  as  a  mother  country — it  goes 
without  saying  that  subtropical  or  warm  temperate  plants  cannot  endure  a 
subantarctic  climate,  and  only  the  most  distant  lands,  Tierra  del  Fuego  and  the 
Falkland  Islands,  were,  thanks  to  the  strong  and  constant  west  wind  drift,  re- 
garded as  a  mother  country.  Fven  if  South  Georgia  served  as  an  intermediate 
station,  the  distances  are  very  great;  besides,  we  do  not  look  west  for  the  an- 
cestors of  the  peculiar  endemics  in  the  Kerguelen  area.  'V\Tere  the  capacity  of 
the  dis{)ersal  agents  appeared  to  be  inadequate,  HoOKER  was  strongly  inclined 
to  look  for  better  land  connections.  The  existence  of  identical  Macaronesian 
species  on  Madeira  and  the  Canary  Islands  can,  he  says,  hardly  be  explained 
without  the  help  of 

intermediate  masses  of  land,  as  the  Salvages  (supj)osing  them  to  have  been  larger)  .  .  . 
the  only  conceivable  means  of  interisland  transport  .  .  .  and  if  intermediate  islands  are 
granted  (and    Mr.    Darwin   freely  admits  these),   why  not  continents? 

He  must  have  found  that  the  distance  between  Madeira  and  the  Canaries  is  too 
large  to  permit  direct  transport  of  diaspores  under  present  wind  and  current 
conditions;  in  the  Kerguelen  case  they  are,  at  least,  favourable.  Nevertheless, 
later  on  Hooker's  faith  in  transoceanic  migration  was  not  as  steadfast  as  before; 
the  case  of  Kerguelen   troubled   him  (.i""^?^): 

Turning  to  the  natural  agents  of  disi)ersion,  winds  are  no  doubt  the  most  power- 
lul,  and  sufficient  to  account  for  the  transport  of  Cryptogamic  spores;  these,  almost 
throughout  the  year,  blow  from  Fuegia  to  Kerguelen  Island,  and  in  the  opposite 
direction  only  for  very  short  i)eriods,  but  appear  quite  insufficient  to  transport  seeds 
over   4000   miles   (j).    13). 

Various  j)henomenons  .  .  .  common  to  .  .  .  Kerguelen,  the  Crozets  and  Marion, 
favour  the  supposition  of  these  all  having  been  peopled  with  land  plants  from  South 
.\merica    by    intermediate    tracts    of  land  that  have  now  disai)peared;  in  other  words, 


DERIVATION  OF  THE  FLORA  AND  FAUNA  355 

that  those  islands  constitute  the  wrecks  of  either  an  ancient  continent  or  an  archipelago 
which  formerly  extended   further  westwards  ...   (p.  15). 

Gulick's  paper  "Biological  peculiarities  of  oceanic  islands"  does  not,  con- 
trary to  its  title,  contain  a  review  of  the  special  characteristics  of  island  biota; 
his  object  was  to  expound  and  defend  the  theory  of  permanent  isolation  of  is- 
lands like  the  Hawaiian,  Galapagos,  Juan  Fernandez,  St.  Helena,  etc.,  which  are 
said  to  offer  irrefutable  proofs  of  true  oceanity.  Of  this  enough  has  been  said 
already;  I  shall  return  to  Hooker's  five  points,  to  which  others  may  be  added. 

Endemism. — The  occurrence  of  numerous  genera  and  species  restricted  to 
oceanic  islands  has  caused  much  discussion.  "Reichtum  an  Endemismen  ist  iiber- 
haupt  der  hervorragendste  Charakterzug  der  Insel-Floren",  Hayek  wrote  [304).  It 
is,  however,  equally  pronounced  in  continental  districts  like  the  Cape  region,  south- 
western Australia,  western  China,  California  or  Chile,  where  local  concentrations  of 
endemics  are  found. 

If  insular  endemics  show  distant  affinity  only  or,  in  extreme  cases,  no  af- 
finity at  all,  to  continental  taxa,  they  are  looked  upon  as  relicts;  as  the  islands 
are  geologically  young,  the  endemics  have  not  evolved  there  but  must  have  im- 
migrated from  some  mother  country,  where  they  have  become  extinct.  They  may, 
however,  have  undergone  some  change  after  their  arrival  to  the  island.  There 
is  also  a  possibility  that  the  continental  progenitor  has,  in  its  turn,  changed  in 
a  different  direction,  making  its  descendants  so  unlike  that  their  relations  are 
obscured.  Species  only  slightly  different  from  continental  ones  are  much  more 
numerous  than  the  relicts;  they  are  supposed  to  have  originated  in  the  islands 
and  give  examples  of  so-called  progressive  endemism.  As  Ciiristensen  {60.  149) 
pointed  out,  another  alternative  leading  to  the  establishment  of  endemic  species 
should  be  considered.  On  the  continent,  from  where  a  plant  found  its  way  to 
an  isolated  island,  opportunities  for  crossing  with  other  species  often  exist, 
eventually  leading  to  the  disappearance  of  the  original  taxon  with  its  special 
characteristics.  Its  island  offshoot  does  not  share  its  fate  but  remains  true  to  the 
original  type.  The  island  form  did  not  originate  through  a  genetic  change  of 
the  continental  species:  it  represents  the  surviving  species  and  is,  as  it  were,  a  relict. 
This  does  not  apply  to  the  pteridophytes.  Crosses  are  extremely  rare,  the  fern 
species  represent,  in  a  high  degree,  pure  lines,  whence  it  follows  that  insular  en- 
demics are  much  rarer  than  among  the  phanerogams.  This  is  true,  but  it  is 
usually  explained  as  a  result  of  the  enormous  spore  production  and  the  facility 
with  which  they  spread. 

Opinions  about  the  true  nature  of  systematically  isolated  taxa  vary.  GUPPY 
{122)  regarded  them  as  either  highly  specialized  products  of  the  islands,  "the  first 
of  their  race",  or  modified  forms  of  allied  continental  genera,  the  majority  of  which 
had  passed  away,  "the  last  of  their  race"  and  probably  doomed;  to  him  the 
islands  appealed  "more  as  registers  of  past  floral  conditions  in  the  continents 
than  as  representing  their  present  state" — this  in  accordance  with  Hooker's 
views.  The  Age-and-area  theory  of  WiLLlS  [286)  claims  that  wides  are  older 
than  endemics,  a  rule  with  few  exceptions;  in  another  paper  [283)  he  states  that 
"insular  endemic  genera  are  as  a  rule  young  beginners,  not  relics".  I  have  dis- 


350  C.   SKOTTSBERG 

cussed  his  theories  in  an  earher  paper  [jji)  to  which  I  refer.  RlDLEV  {20j)  called 
the  island  endemics  "newborn  s[)ecies  ",  admittinf^  that  all  did  not  fall  within  this 
cate^ors-  but  were  "epibiotics,  relics  at  the  end  of  their  species  life  .  .  .  unable  to 
reach  another  suitable  spot  for  their  |j;rowth".  It  is  noteworthy  that  so  many  of  the 
"e[)ibiotics"  are  Composites,  famous  among  diffusionists  for  their  alleged  effective 
dispersal  mechanisms;  in  RiDi.KV's  eyes  they  are,  perhaps,  pseudo-relicts.  Cain  (^2) 
takes  more  or  less  the  same  position:  "the  relic  nature  of  an  endemic  should 
never  be  accepted  without  some  form  of  positive  evidence"  (p.  227);  proofs  are  hard 
to  tind,  no  island  cases  as  clear  as  Giiikgo  or  Mctascquoia  are  known.  An  endemic 
inhabiting  a  strikingly  limited  area  may  be  a  young  species  that  had  no  time  to  spread, 
or  it  may  be  too  stenotoj)ic,  but  others  are  what  Cain  (p.  230)  calls  "senescent";  such 
species  occup\-  a  small  area,  are  relatively  constant,  ecologically  of  narrow^  amplitude 
and  show  low  competitive  ability.  They  are  unable  to  "penetrate  the  prevailing 
habitats  that  are  dominated  by  the  typical  vegetation  of  the  region"  but  behave 
just  as  stenotoj)ic  young  beginners.  If  indeed  old  and  senescent  they  ought  to 
siiow  some  primitive  characteristics.  According  to  Cain  senescent  species  con- 
stitute "an  anomalous  element  in  the  flora  of  a  given  region";  this  may  be  true, 
for  in  many  instances  they  survive  from  an  earlier  climatic  period  and  are  barely 
able  to  hold  their  ground  under  the  changed  conditions.  Nevertheless  there  are 
cases  when  such  anomalous  species  form  the  typical  vegetation  of  a  certain  habi- 
tat and  where  nobody  would  dream  of  regarding  them  as  young  beginners;  the 
'^ Robi)iso)iia  assemblage"   in   Masatierra  offers  a  good  example  (see  2ji). 

Xuiubcr  of  species  per  genus.— \Nq  know  that  island  floras  contain  a  fair  num- 
ber of  monotypical  genera,  many  genera  that  are  large  elsewhere  but  repre- 
sented on  a  given  island  by  a  single  species,  and  few  with  many  species,  so  that 
tlie  numerical  relation  between  species  and  genus  approaches  i  and  does  not 
exceed  2.  and  this  has  been  regarded  as  a  good  proof  that  the  island  is  truly 
oceanic  and  has  been  peopled  -accidentally  by  waifs  and  strays.  This  rule  is  not 
without  exceptions,  among  which  the  Ilawaiian  flora  is  the  most  striking.  Fos- 
i:Kk(;,  who  contributed  a  cha|)ter  on  the  higher  flora  to  Zimmerman's  book  (2^8), 
indicates  <S3  families,  216  genera  and  1729  species  of  angiosperms.  The  figure 
for  the  genera  may  be  a  little  too  low.  With  regard  to  species  all  depends  on 
the  species  concept.  Hiil.Kr.RAND  (307)  was  conservative;  from  his  Flora  Drude 
^.^^\^-  ^?>^^)  g'^'t  the  ratio  6.2  :  i,  but  scores  of  well-marked  species  have  been  de- 
scribed since  i  8<S,S,  and  in  addition,  particularly  during  the  last  two  or  three  decades, 
a  large  number  of  taxa  that  are  little  more  than  microspecies;  unfortunately 
no  case  of  apomixis  has  been  found  as  yet.^  It  is  difficult  to  know  which  way 
to  take  out  of  this  maze;  neither  the  role  played  by  hybridism  nor  the  existence 
ot  modifications  due  to  environment  has  been  duly  considered.  Based  on  FOS- 
i;i:r(;'s  figures,  the  relation  species  :  genus  is  8:1,  and  this  is  not  at  all  what  one 
expects    to   find   in   an   oceanic   flora.   Nobody  is,  I   think,   likely  to  disclaim  New 

1  .AinDiiK^  taxononiists  particularly  responsible  for  this  alarming^  increase  PL.  P2.  Sheri-F  and 
H.  .St.  joiix  should  be  mentioned  first,  but  some  others  have  also  contributed  and  the  writer 
cannot  plead  innocent.  Shkrik,  in  addition  to  numerous  new  species,  has  described  an  endless 
number  of  varieties. 


PERIVATION  OF  THE  FLORA  AND  FAUNA  357 

Zealand's  continentality.  According  to  CnEESF:MAN  the  number  of  families  is  97, 
of  genera  382  and  of  species  141 5;  the  relation  in  question  is  3.7  :  i.  I  cannot 
attach  much  importance  to  such  figures.  If  an  island  is  a  remnant  of  a  sub- 
merged land-mass  upon  which  during  the  process  of  sinking  lava  was  ejected  until 
the  old  foundation  disappeared,  only  a  small  part  of  the  flora  and  fauna  will  sur- 
vive the  catastrophe,  and  unless  progressive  endemism  comes  to  play  a  role,  the 
living  world  will  present  a  picture  of  disharmony,  with  a  reduced  number  of 
families  and  a  low  species  :  genus  ratio.  Climatic  changes  will  create  a  similar 
situation,  if  a  portion  of  a  continental  flora  has  lost  its  connection  with  the  con- 
tinent. The  Falkland  Islands  rest  on  the  continental  shelf,  they  are  formed  by  old 
sediments  on  a  granitic  foundation  and  are,  universally  I  think,  classified  as  con- 
tinental islands,  but  their  angiospermic  flora  is  disharmonic  and  fragmentary:  38 
families,  97  genera  and  143  species;  the  ratio  is  1.47  :  i.  There  is  a  single  endemic 
genus  and  a  small  number  of  endemic  species.  The  poverty  is  due  to  a  severe 
climate  and  to  losses  suffered  during  the  pleistocene  period  of  solifluction,  con- 
temporaneous with  the  glaciation  on  the  mainland.  The  constitution  of  a  flora,  con- 
tinental or  insular,  depends  on  a  combination  of  many  factors,  geological,  his- 
torical, climatic,  genetic  and  so  forth. 

Absence  of  large,  ividespread  and  '' successfur'  families. — The  lack  of  conifers 
in  islands  demonstrates,  it  is  said,  that  such  islands  are  not  continental,  for  in 
all  continents  gymnosperms  are  plentiful;  "cones  do  not  float",  and  the  seeds, 
winged  or  unwinged,  have  no  chances  to  be  carried  very  far,  but  it  is  surprising 
that  also  Taxads  and  many  Podocarps,  adapted,  as  it  were,  to  endozoochorous 
bird  dispersal,  are  conspicuous  by  their  absence.  Consequently,  islands  where  cone- 
bearing  species  exist  are  regarded  as  continental:  New  Zealand,  Tasmania,  Nor- 
folk Island,  New  Caledonia,   Fiji,  etc. 

Exceptions  occur,  islands  looked  upon  as  permanently  isolated,  Bermuda, 
the  Azores,  Madeira  and  the  Canaries,  have  at  least  some  species  oi  Jimiperus; 
perhaps  junipers  possess  some  kind  of  dispersal  capacity  and  do  not  count,  but 
we  cannot  get  away  from  Finns  cana^^iensis. 

Other  large  and  wide-spread  families  very  poorly  represented  on  isolated  is- 
lands are  Leguminosae,  Araceae  and  Orchidaceae.  Peas  are  a  staple  food  of  many 
birds  but  few  if  any  able  to  pass  their  digestive  tract  unharmed;  they  are  too 
heavy  to  be  carried  any  distance  by  wind  and,  notorious  beach  plants  excepted, 
will  sink  in  water.  The  dry  pods  would  float  but  they  do  not  fall  off  but  open 
on  the  plant  to  discharge  the  seeds.  Nevertheless  SopJiora  sect.  Ediuardsia  has  a 
number  of  closely  related  species  scattered  over  the  south  hemisphere  on  islands 
as  remote  as  Diego  Alvarez,  Rapa,  Easter  and  Marquesas,  with  one  species,  more 
well-marked  than  the  rest,  on  Hawaii.  This  is  a  very  puzzling  case  of  disjunct 
distribution,  but  it  is  not  expedient  to  draw  any  far-reaching  conclusions  from 
this  unique  case.  The  seeds  are  of  the  size  of  a  small  pea  and  not  equipped  with 
a  capacity  to  travel  greater  than  in  hundreds  of  leguminous  plants,  which  are 
within  reach  but  never  crossed  any  water  barriers,  nor  used  any  land  bridges. 
Did  they  not  yet  exist,  were  they  not  within  reach  when  connection  was  estab- 
lished   or  have  they  all  died  out  in  the  islands.^  Each  alternative  seems  equally 


35^ 


SKOTTSBERG 


improbable.  Araceac  are  pleiitihil  in  luiinid  tropical  climates,  and  some  islands, 
the  Hawaiian  for  instance,  ouL^ht  to  offer  a  suitable  environment;  still  there  is  not 
a  sin<^le  native  species.  And  it  is  strange  that  orchids,  with  their  dust-like  seeds, 
should  be,  if  not  altogether  absent,  so  few,  becoming  fewer  and  fewer  as  we  proceed 
east  from  Mala\'sia  and  Melanesia;  cf.  ?/cV,  map  21.  However,  the  seeds,  even 
if  carried  far,  rarely  are  able  to  retain  their  viability  long  enough  to  become 
established,  should  they  haj)pen  to  strike  a  spot  where  they  can  germinate.  I  be- 
lieve that  also  where  we  find  a  very  rich  orchid  flora  most  of  the  species  occupy 
restrictetl  areas;  their  advance  over  land  is  slow.  A  species  introduced  on  pur- 
pose will,  in  rare  instances,  become  naturalized,  but  I  cannot  remember  ever 
luning  heard  of  an  adventitious  orchid.  Taking  all  these  circumstances  into  account, 
the  poverty  of  distant  islands,  even  if  temporarily  connected  with  other  lands, 
is  perhaps  not  altogether  incomj)rehensible.  Besides,  the  submerged  links  may, 
for  all  we  know,  have  been  {)oor  in  legumes,  arum  lilies  and  orchids.  CAMPBELL, 
admitting  that  the  total  absence  oi  Araucana,  Agat/us,  Podocarpus,  Ficus  and  Ara- 
icac  in  oceanic  islands  of  the  Pacific  is  a  valid  objection,  not  easy  to  explain, 
suggests  that  perchance  they  once  did  exist  there  but  were  destroyed  by  volcanic 
eruptions  [^6.  181). 

'J he  prcpcnidcraiicc  of  woody  plants  in  oceanic  islands. — Hemsley  (727.  31) 
em|)hasized  the  prominent  part  taken  by  arboreous  and  shrubby  species  in  many 
islands  and  pointed  out  that  in  some  cases  they  belong  to  otherwise  herbaceous 
families.  Many  peculiar  genera  of  Compositae  are  confined  to  islands  such  as 
Hawaii,  (ialapagos,  and  Juan  Fernandez,  and  are  scattered  over  Polynesia  (see 
,VcS'.  map  20),  and  in  the  Atlantic  (Macaronesia,  St.  Helena)  and  Indian  oceans 
(.Socotra,  the  Seychelles,  Mascarene  Islands);  arboreous  Compositae  are,  as  HemS- 
l.KV  just])'  remarks,  by  no  means  restricted  to  island  habitats  but  numerous  in 
tropi(  al  regions  of  the  continents,  particularly  in  America.  Both  Hawaii  and  Juan 
I'Y-rnandez  offer  good  examples  of  woody  plants  belonging  to  otherwise  herba- 
ceous families  or  genera,  but  Hkmsi.kv's  statement  p.  31  about  Gwinera\\\]\x2S\. 
I'V-rnandez  "caulescent  species  unknown  elsewhere" — is  erroneous,  for  they  are 
just  as  caulescent  in   Hawaii;   besides,  they  are  not  woody,  but  herbaceous. 

Scarcity  of  herbaceous  species,  especially  of  tJierophytes. — Native  annuals  and 
biennials  are  rare  in  islands.  They  cannot  have  been  less  capable  of  migrating 
across  the  sea  than  trees  or  shrubs,  nor  could  soil  or  climate  prevent  them  from 
getting  established;  this  is  at  once  disproved  by  the  countless  herbaceous  weeds 
brought  by  man  which  threaten  to  overrun  so  many  islands.  The  reason  must 
be  historical.  If  it  is  true  that  life  forms  with  a  woody  stem  are  the  most  ancient, 
and  if  tlie  original  stock  making  u|)  island  floras  dates  back  to  before  the  rise  of 
herbs,  a  period  during  which  the  distribution  of  land  and  sea  was  another  than 
now,  we  can  exj)lain  the  high  {)r()p()rtion  of  arborescent  species  in  islands  iso- 
lated since  millions  of  \'ears. 

1-j'ideiice  for  a  greater  antiquity  of  lignifed  angiosper)ns. — The  angiosperms 
originated  in  the  Mesozoic  and  are  traced  back  to  the  Lower  Cretaceous.  The 
tree  form  was  the  res|)onse  to  a  warm  and  humid  climate.  These  problems  were 
subjected   to  a  comprehensive  analysis  by  SiNNOTT  and  I.  W.  Bailey  [22f),  whom 


DERIVATION  OF  THE  FLORA  AND  FAUNA  359 

I  shall  quote  at  some  length.  They  state  that  evidence  for  a  more  recent  origin 
of  herbaceous  plants  is  furnished  by  palaeontology,  anatomy,  phylogeny  and  phyto- 
geography.  Evidence  from  paleobotany  is,  however,  not  conclusive,  because  herbs 
are  much  less  fit  to  be  preserved  as  fossils,  but  it  is  true  that  they  increase  in 
number  in  the  younger  formations.  Evidence  from  anatomy  and  phylogeny  are 
said  to  be  positive;  within  a  group  including  both  ligneous  and  herbaceous  spe- 
cies, the  former  show  other  primitive  characters;  of  the  Leguminosae  the  more 
primitive  members  are  all  woody  (Mimosaceae,  Caesalpiniaceae),  the  proportion 
of  herbs  being  vastly  greater  among  the  Papilionaceae.  The  authors  continue,  p.  572  : 

It  is  generally  admitted  that  endemic  species  of  a  flora  ...  are  for  the  most  part 
more  ancient  than  the  non-endemic  element,  for  they  must  either  have  had  their  origin  in 
the  region  ...  a  process  usually  requiring,  a  long  time — or  else  they  must  be  remnants 
of  an  older  vegetation  which  has  elsewhere  become  extinct.  Endemic  genera  and 
finally  endemic  families  are  in  this  way  regarded  as  progressively  more  ancient  portions 
of  the  flora. 

I  doubt  that  this  statement  is  of  general  application;  we  cannot  argue  that, 
in  a  given  flora,  all  endemics  are  more  ancient  than  the  wides,  because  a  widely 
distributed  species  may  have  remained  unchanged  for  millions  of  years  and  be 
older  than  another  which,  for  various  reasons,  happens  to  have  become  greatly 
restricted  in  range  and  endemic  in  a  small  area.  WiLLlS  and  his  school  are  dia- 
metrically opposed  to  the  opinion  of  SiNNOTT  and  Bailey  and  neither  is  in  pos- 
session of  the  absolute  truth. 

Proceeding  to  an  analysis  of  certain  insular  floras  deserving  special  attention 
the  authors  assume  that,  if  woody  types  are  more  ancient  than  herbaceous,  a 
flora  which  has  been  for  a  long  time  isolated  ought  to  contain  a  large  proportion 
of  woody  endemics,  and  this  is  what  they  do  find.  Their  analysis  of  the  Juan 
Fernandez  flora  was  based  on  JOHOw's  work,  but  this  has  long  been  out  of  date 
and  the  figures  are  incorrect  (new  ones  will  be  found  in  Chapter  IX),  so  I  shall 
leave  this  subject  aside  here.  The  general  conclusions  in  the  chapter  "Discussion 
of  Isolated  Insular  Floras"   deserve  to  be  quoted  in  full  (p.   579). 

It  is  thus  very  clear  that  woody  plants  constitute  a  more  conspicuous  element  in 
the  flora  of  isolated  oceanic  islands  than  in  the  flora  of  adjacent  continental  areas  from 
which  their  vegetation  has  possibly  been  derived,  and  also  that  the  most  ancient  por- 
tion of  the  island  floras,  if  endemism  is  to  be  regarded  as  a  criterion  of  antiquity,  is 
much  more  woody  than  the  recently  acquired  elements.  Annual  herbs,  which  seem  to 
be  the  last  step  in  reduction,  are  almost  entirely  absent  from  insular  floras,  as  has 
been  noted  by  Darwin,  Hooker,  and  others.  Since  the  vegetation  of  these  isolated 
oceanic  islands  is  to  be  regarded  as  more  ancient  in  its  composition  than  that  of  larger 
land  areas,  it  may  be  looked  upon  as  a  vestige  of  an  earlier  and  much  more  uni- 
form flora  which  flourished  over  the  earth  during  the  middle  or  latter  part  of  the  Ter- 
tiary, and  before  the  great  flood  of  herbaceous  vegetation,  developed  chiefly  in  the 
north  temperate  lands,  had  spread  over  the  globe.  This  conclusion  is  strengthened  by 
the  many  similarities  which  these  widely  separated  island  floras  bear  to  one  another. 

Some  of  the  statements  made  by  these  authors  are  questionable,  but  in  the 
main  I  agree;  only  I  prefer  to  say  "early  to  middle  Tertiary".  Darwin,  who  was 


36o 


SKOTTSBERG 


the  first  to  note  the  predominance  of  trees  on  oceanic  islands,  tried  to  explain 
it  as  a  result  of  natural  selection.  SlNNOTT  and  Baii.EV's  criticism  is  absolutely 
C()nvincin<^. 

The  authors  also  paid   attention  to  The  Ancient  Flora  of  Antarctica,  p.  592: 

It  seems  to  be  a  reasonably  safe  conclusion  that  all  genera  commonly  designated 
as  "Antarctic"'  from  their  confinement  to  the  temperate  region  of  the  southern  hemi- 
sphere,  were  iiiiiabitants  of  the   ancient   Antarctic  continent. 

The  authors  made  an  attempt  to  reconstruct  this  flora  and  came  to  the  con- 
clusion that  "two  thirds  of  its  endemic  dicotyledons  were  woody  plants"  (p.  599). 

In  the  authors'  opinion  most  of  the  herbaceous  vegetation  originated  in  the 
north;  this  may  be  so  because  in  the  latter  part  of  the  Tertiary  period,  with  the 
increasing  differentiation  of  climatic  regions,  the  temperate  and  cold-resistant  flora 
is  supposed  to  have  taken  j)ossession  of  large  areas,  but  these  were  concentrated 
in  the  north;  excej)t  the  Antarctic  continent  there  was  not  much  land  in  the 
far  south. 

Several  authors  have  expressed  the  same  opinion  as  SlNNOTT  and  Bailev. 
Irms(IIi:k  (//;)  was  convinced  that  most  of  the  primitive  angiosperms  were  trees 
and  that  the  relation  between  tropical  and  temperate  genera  and  species  point  in 
this  direction;  to  take  one  example:  it  is  generally  acknowledged  that  the  her- 
baceous, mainly  temperate  fanfily  Cruciferae  descends  from  woody  tropical  Cap- 
paridaceae  (IkMSCllKR  might  have  called  attention  to  the  miniature  Lepidiuni  trees 
in   Hawaii): 

Auch  sonst  ist  die  Abstammung  gemiissigter  Sippen  von  tropischen  nachgewiesen 
worden.  .  .  .  Diesc  Gegeniiberstellung  der  gemiissigten  und  tropischen  Sipj)en  lehrt  aber 
auch  ohnc  wciteres,  dass  mit  der  Anpassung  von  Formen  an  die  extratroi)ischen  Zonen 
zuglcicii  (lie  F  m  priigung  des  Ty  ])us  des  H  o  1  z  ge  vviic  h  ses  in  den  des  Krautes 
vor  sich  ging,  somit  let/,  tere  als  die  j  linger  en  Formen  die  Abkomm- 
1  i  n  g  e   \ 0  n    1 1  o  1  /.  g  e  w  ii  c  h  s  e  n   s  i  n  d   (l.  2  oq). 

Dass  (lie  Thcorie  dcr  Abstammung  der  iibrigen  Wuchsformen  von  der  Gestalt  des 
troj)is(  hen  liaunics  immer  mchr  an  l>oden  gewinnt,  geht  zum  Beispiel  aus  dem  Buche 
von  j'.cws  hers  or:  "The  mcgatherm  hygrophilous  forest  of  the  tropics  is  probably  the 
most  an(  ient  t\  pc  of  habitat  .  .  .  and  most  recent  of  all  (life-forms)  is  the  annual  type" 
(II.  321). 

The  investigation  of  Ji:ki"RKV  and  ToKRKV  [144)  led  to  the  conclusion  that 
"the  origin  of  the  herbaceous  type  in  dicotyledons  is  from  woody  or  arboreal 
forms",  and  S 1 1  x  K\\  i:i.L,  in  his  monograph  of  Chacnaciis  [2^g\  Compositae,  33 
sp.),  states  that,  "within  a  family  or  genus,  woody  perennial  species  are  more 
j)rimitive  than  herbaceous  annual  s|)ecies '.  An  evolution  in  the  reversed  direction 
is,  however,  postulated  by  HlKl  iix.sox  [140.  I.  4)  in  the  Polycarpicae.  He  regards 
the  herbaceous  Kanunculaccae  as  the  most  j)rimitive;  a  woody  structure  is,  in  this 
order,  secondarw  The  current  oj)inion  is  that  the  woody  Magnoliaceae  are  among 
the  oldest   living   angiosperms. 

The  rosette  trees.  '\\\(i  common  t\'|)e  of  this  interesting  growth-form  is  char- 
acterized by  a  candelabrum  like  mode  of  branching,  the  two  or  three  (rarely  more) 
innovations    situated    on   practically  the  same  level  at  the  base  of  a  terminal  in- 


DERIVATION  OF  THE  FLORA  AND  FAUNA  361 

florescence;  in  some  cases  cauliflory  is  observed.  A  much  more  uncommon  type 
is  the  unbranched  stem  terminated  by  a  tuft  of  large  leaves  (if  entire  =  Grise- 
BACH's  Clavija  form,  j^j.  I.  ii),  either  pollacanthic  with  lateral  inflorescences 
or  hapaxanthic  when,  after  a  number  of  years,  a  terminal  inflorescence  ends  the 
life  of  the  individual;  this  type  is  the  most  unusual  of  all.  All  kinds  have  in  com- 
mon the  very  short  internodes  and  the  short-lived  leaves,  a  self-evident  condition 
for  the  formation  of  a  compact  tuft.  A  reduction  of  the  rosette  tree  to  herbaceous 
state  would  result  in  a  compact  caudex  multiceps  or  a  single  basal  rosette;  an 
hapaxanthic  tree  would  become  a  therophyte. 

These  growth  forms  are  by  no  means  restricted  to  island  habitats  but  are 
also  found  in  all  continents,  they  belong  to  many  different  genera  and  families 
and  are  characteristic  of  such  dicotylous  families  as  Araliaceae,  Caricaceae,  Theo- 
phrastaceae  etc.;  typical  examples  are  found  in  Compositae  (e.g.  Espeletia,  species 
of  Senecio),  Epacridaceae  [Dracophyllum],  Lecythidaceae  (Grias),  Meliaceae  (Ca- 
rapa),  Rutaceae  [Spatkelia],  Sapindaceae,  and  so  forth.  Among  the  monocotyle- 
dons the  vast  majority  of  palms  belong  here,  the  Pandanaceae  should  of  course 
be  mentioned,  and  well-known  examples  are  scattered  through  the  Liliiflorae  (spe- 
cies of  Aloe,  Cordyline,  species  of  Yucca,  Dracaena,  Fourcroya  etc.),  Ravenala  in 
the  Musaceae,  and  Piiya  Rainiondii  in  Bromeliaceae.  Hapaxanthic  trees  are  few; 
Krause  [162)  called  attention  to  the  interesting  rutaceous  Sohnreya  excelsa  Krause 
from  Amazonas. 

Thus,  even  if  rosette-trees  are  more  or  less  widely  distributed  over  the  globe 
in  warmer  regions,  it  is  a  fact  that  they  are  a  particularly  conspicuous  feature  in 
island  floras.  In  the  Pacific  they  are  very  plentiful  in  the  Hawaiian  Islands,  espe- 
cially among  the  Compositae  and  Lobeliaceae,  also  the  hapaxanthic  type  repre- 
sented; they  occur  in  the  Galapagos  Islands  (Grisebach  II.  512),  we  find  many 
in  New  Caledonia,  they  constitute  a  large  proportion  of  the  poor  Juan  Fernandez 
flora  (see  Chapter  IX),  and  they  are  scattered  over  Oceania.  In  the  Atlantic  they  are 
numerous  in  Macaronesia  (e.g.  Campa7iula  Vidalii  on  the  Azores,  MusscJiia  on  Ma- 
deira, species  of  Aeoniuni,  Dracaena,  EcJiiiwi,  Melanoselinum,  Sonchus,  Sinapide7i- 
dru7n,  etc.).  I  suppose  that  some  are  found  also  on  the  islands  of  the  Indian  Ocean, 
but  I  have  no  reliable  information.  I  may  be  entirely  wrong,  but  I  have  the  idea 
that  this  is  an  old-fashioned  growth  form,  a  relict  element  in  island  floras,  and  that 
the  high  volcanic  islands  of  the  Pacific  and  the  Atlantic,  which  are  so  like  each 
other  in  geology  and  topography,  are,  in  their  present  shape,  of  approximately  the 
same  age  and  date  back  to  a  period  of  land  submergence  and  great  volcanic 
activity.  Perhaps  also  the  "inland  islands",  the  high  volcanic  mountains  of  Africa, 
deserve  to  be  mentioned  in  this  connection,  famous  as  they  are  for  their  magni- 
ficent tree  Lobelias  and  Senecios. 

Rosette  trees  seem  to  favour  open,  sunny  situations;  this  is  certainly  the  case 
in  Juan  Fernandez,  where  they  are  definitely  adapted  to  such  habitats.  Single 
specimens  of  a  few  species  are  sometimes  found  growing  in  the  shade,  with  the 
result  that  the  internodes  become  much  longer,  the  dense  tuft  dissolves  and  the 
formation  of  flowers  is  suppressed.  ScillMPER  believed  that  they  are  adapted  to 
a   very    windy    climate    and  fit  to  withstand  the  pressure  of  strong  winds  better 


362  C.   SKOTTSBERG 

than  trees  of  a  more  ordinary  type,  but  this  theory  has  been  refuted  [22^),  and 
recent  observations  in   the  field  have  not  made  me  alter  my  opinion. 

Ab:se)ice  of  quadrupeds.  The  only  mammals  regarded  as  possibly  native  in 
isolated  islands  are  bats,  but  many  islands  do  not  have  any.  Storm-drift  has  been 
postulated,  but  if  the  distance  be  very  great,  the  animals  might  be  without  food 
too  long  unless  Nature  provides  a  fair  supply  of  air-born  insects.  To  other  land 
mammals  wide  stretches  o{  open  water  are  an  absolute  obstacle  and  their  absence 
is  regardeil  as  one  of  the  safest  proofs  of  the  permanent  isolation  of  oceanic  is- 
lands, just  as  the  presence  of  endemic  foxes  in  the  Falkland  Islands  (now  extinct) 
supports  the  theory  of  an   earlier  connection   with   South   America. 

However,  we  cannot  know  that  indigenous  mammals  never  existed  on  islands, 
graduall}'  having  become  extinct  when  the  submergence  of  land  had  proceeded 
and  grazing  grounds  became  smaller  and  smaller.  It  would  appear  that  this  theory 
is  contradicted  b)'  the  fact  that  herbivorous  mammals  have  been  introduced  by 
man  to  many  islands  and  do  make  a  living  there  also  when  not  tended,  but  if 
allowed  to  naturalize  and  multi{)ly  unrestrainedly  in  virgin  surroundings,  their 
ravages  would  perhaps  prove  catastrophal  to  the  native  vegetation  and,  as  a  con- 
secjuence  thereof,  to  themselves.  They  would  die  out  and  leave  the  flora  to  re- 
cover. We  must  remember  that  the  islands  are  small,  most  of  them  very  small, 
that  herbixorous  animals  need  space  and  that  pasture  lands  as  they  exist  now 
are  a  result  of  cleared  forest  soil  and  introduction  of  innumerable  alien  weeds  or 
cultigens.  Another  explanation  of  the  absence  of  mammals  is  that  the  islands  were 
cut  off  so  early  that  mammals  had  not  yet  taken  possession  of  the  earth  or  were 
not  universally  distributed  and  perhaps  not  within  reach.  But  these  are  mere  wild 
specuhitions.  Xo  fossil  remains  have  been  discovered,  nor  can  they  be  expected 
on  purely  volcanic  islands.  We  cannot  attack  this  problem  with  a  hope  of  success 
as  long  as  we  know  little  or  nothing  of  the  geograj)hical  history  of  islands  and 
archi|)elagc)es.  Hut  wc  have  better  remember  that  continental  islands  such  as  New 
Zealand,  New  Caledonia,  etc.  are  in  the  same  precarious  position  with  regard  to 
mammals  as  the  Hawaiian  Islands. 

-Xative  reptiles  and  amphibians  are  also  absent.  CamI'BKIJ.  (./^)  mentions  in 
j)assing  that  tiicre  arc  half  a  dozen  lizards  in  Hawaii,  but  all  are  species  wide- 
spread in  the  .South  Pacific;  most  likely  they  were  introduced  with  the  early  human 
immigrants  or  j)erha])s  later.  That  there  are  no  frogs  or  toads  would  indicate, 
Cami'I'.kij,  thinks,  that  the  archi])elago  became  isolated  before  the  modern  kinds 
of  these  animals  had  been  developed.  lUit,  as  MlMFORD  points  out,  many  con- 
tinental islands  lack  all  lower  vertebrates  (/cV-,\  24<S). 

1  low  can  we  exj)lain  all  these  i)eculiarities  in  oceanic  islands.^  HooKKR's  answer 
was  this: 

Thus,  according  to  the  hyi)()thc,sis  of  trans-oceanic  nn'gration,  and  the  theory  of  the 
derivative  orij^nn  of  spec  ics,  wc  can  mulcrstand  why  the  ancient  tyj)es  .  .  .  should  have 
survi\c(l  on  tlie  islands  to  wliic  h  hut  few  of  the  superior  race  had  j)enetrated ;• — we  can 
understand  how  it  comes  about,  that  so  many  continental  species  and  genera  are  represented 
on   the   island   by   similar   hut   not   identical   sj)ecies   and   genera,   and  that  there  is  such 


DERIVATION  OF  THE  FLORA  AND  FAUNA  363 

a  representation  of  genera  and  species  in  the  separate  islands  of  a  group; — we  can  under- 
stand why  we  find  in  the  Atlantic  island  Floras  of  such  a  graduated  series  of  forms, 
ascending  from  variety  to  genus,  without  those  sharp  lines  of  specific  distinction  that 
continental  plants  exhibit; — why  whole  tribes  are  absent  in  the  Islands;  why  their  Floras 
are  limited,  and  species  few  in  proportion  to  genera; — why  so  many  peculiar  genera 
tend  to   grotesque  or  picturesque  arborescent  forms  ...  (p.    ii). 

All  this  is,  perhaps,  not  so  easy  to  understand  as  Hooker,  impressed  by 
Darwin's  theory,  thought.  Finally  I  shall  quote  the  summary  of  Wallace's  General 
remark  on  Oceanic  Islands  [2^8.  329-330),  to  bring  this  chapter  to  a  close: 

They  all  agree  in  the  total  absence  of  indigenous  mammalia  and  amphibia,  while 
their  reptiles,  when  they  possess  any,  do  not  exhibit  indications  of  extreme  isolation  and 
antiquity  [for  the  moment  Wallace  seems  to  have  forgotten  the  giant  tortoises  and  the 
peculiar  endemic  lizards  in  the  Galapagos,  typically  oceanic  islands  in  his  opinion]. 
Their  birds  and  insects  present  just  the  amount  of  specialisation  and  diversity  from  con- 
tinental forms  which  may  be  well  explained  by  the  known  means  of  dispersal  acting 
through  long  periods;  their  land  shells  indicate  greater  isolation,  owing  to  their  admittedly 
less  effective  means  of  conveyance  across  the  ocean;  while  their  plants  show  most  clearly 
the  effects  of  those  changes  of  conditions  which  we  have  reason  to  believe  have  occurred 
during  the  Tertiary  epoch,  and  preserve  to  us  in  highly  specialised  and  archaic  forms 
some  record  of  the  primeval  immigration  by  which  islands  were  originally  clothed  with 
vegetation. 

Chapter  VIII. 

Evolution  in  Oceanic  islands. 

Ever  since  the  high  proportion  of  endemic  organisms  in  oceanic  islands  and 
especially  the  occurrence  of  systematically  isolated  genera  were  first  noticed,  an 
explanation  of  this  condition  has  been  sought.  Most  authors  have,  as  we  have  seen, 
assumed  that  the  islands  never  had  been  connected  with  a  continent  and  that, 
consequently,  their  entire  living  world  had  developed  from  a  limited  number  of 
ancestors  carried  across  the  water.  Even  if  the  islands  had  emerged  as  late  as  during 
the  Pliocene,  the  time  was  thought  to  have  been  sufficient  for  immigrants  to  get 
transformed.  It  was  a  peculiar  island  world,  the  Galapagos  Archipelago,  which  gave 
birth  to  Darwin's  theory  of  origin  of  species  through  variation  and  natural  selec- 
tion, and  Hooker,  when  dealing  with  a  number  of  islands,  was  convinced  that 
Darwin  had  given  the  solution  to  their  problems  and  that  their  status  strongly 
supported  his  theory: 

And  if  many  of  the  phenomena  of  oceanic  island  Flora  are  thus  well  explained  by 
aid  of  the  theory  of  the  derivative  origin  of  species,  and  not  at  all  by  any  other  theory, 
it  surely  is  a  strong  corroboration  of  that  theory.  Depend  upon  it,  the  slow  but  steady 
struggle  for  existence  is  taking  advantage  of  every  change  of  form  and  every  change  of 
circumstance  to  which  plants  no  less  than  animals  are  exposed;  and,  variation  and  change 
of  form  are  the  rules  in  organic  life.  ...  By  a  wise  ordinance  it  is  ruled,  that  amongst 
the  living  beings  like  shall  never  produce  its  exact  like;  as  no  two  circumstances  in 
time  or  place  are  absolutely  synchronous,  or  equal,  or  similar,  so  shall  no  two  beings 
be  born  alike;  that  a  variety  in  the  environing  conditions  in  which  the  progeny  of  a  living 
being  may  be  placed,   shall   be  met  by  variety  in  the  progeny  itself.  A  wise  ordinance 


364 


C.   SKOTTSBERG 


it  is,  that  ensures  the  succession  of  beings,  not  by  multiplying  absolutely  identical  forms, 
but  by  varying  these,  so  that  the  right  form  may  fill  its  right  place  in  Nature's  ever 
varying   economy  (/rtV.  11-12). 

As  we  have  seen  already,  Hookkr's  first  point  when  summarizing  the  charac- 
teristics of  island  floras  was  that  also  their  most  peculiar  genera  had  their  origin 
in  a  HKUiier  continent,  that  no  oceanic  island  had  been  a  centre  for  special  creative 
forces.  (iklSKliACll  (II.  494)  may  have  been  the  first  to  express  another  opinion. 
Referring  to  the  systematically  isolated  endemics  in  St.  Melena,  he  argued  that  they 
never  had   iiad  any  allies  elsewhere: 

Na(  h  diesen  'I'hatsachen  kann  von  einem  Stammkontinent  weder  in  dem  Sinne  die 
Kcde  sein,  class  die  Flora  von  daher  durch  natiirliche  Einvvanderungen  bereichert  wurde, 
noch  als  ob  die  endemischen  Arten  aus  Umbildungen  von  Pflanzen  hervorgegangen  wiiren, 
die  in  einer  tViiheren  Periode  von  auswiirts  dahin  gelangten.  Nur  von  gewissen  klima- 
tischen  .\nalogieen  ist  ihrc  Organisation  der  Ausdruck.  S.  Helena  verhalt  sich  demnach 
ganz  verschieden  von  den  Kap-\'erden  und  liefert  den  voUgUltigen  Beweis,  dassdie  Entste- 
hung  der  Pflanzen  auf  Inseln,  cbenso  wohl  wie  auf  Kontinenten,  unabhiingig  von  anderen 
W'getationscentren  mogliih  war.  W'arum  sollte  auch  der  geographische  Umfang  eines 
(".ebiets  auf  die  Kriiftc,  welche  die  Organisationen  erzeugt  haben,  von  Einfluss  sein?  In 
dem  kleinsten  Raunie,  wie  im  grossten,  konnten  sie  in  besonderer  Weise  sich  entfalten: 
nur   werden   sie   im   erstercn    Palle   weniger  zahlreich   sein   miissen. 

In  his  discussion  of  the  Galapagos  flora  he  admits  that  what  he  terms  vicarious 
species,  endemic  in  tiie  islands,  may  have  arisen  through  transformation  of  American 
species,  but  he  doubts  it. 

Diejcnigen,  welche  annehmen,  dass  die  vikariierenden  Arten  aus  Umbildungen  von 
eingcwanderten  hervorgegangen  sind,  konnen  auch  unter  den  endemischen  (iewiichsen 
der  (ialaj)agos  lieispiele  genug  anfiihren,  dass  eine  nahe  Verwandtschaft  dieselben  mit 
Ainerika,  als  ihrem  vorausgesetzten  Stammkontinent,  verbinde.  Aber  allgemein  liisst  sich 
dieser  ( lesichtspunkt  niclit  durchfiihren.  (ierade  unter  den  geselligen  Holzgewiichsen  ,  .  . 
finden  wir  die  eigentiimlichsten  Erzeugnisse,  die  Scalesien  und  andere  Synanthereen,  die 
nach  ihrer  systematischen  Stelhmg  dem  Festlande  ebenso  fremdartig  gegeniiber  stehen 
wie  (he  Lobeliaceen  des  Sandwich-Archipels.  .  .  .  Alle  P>eredsamkeit,  womit  die  Abstam- 
uiung  flcr  Vegetation  ozeanischer  Inseln  von  den  Kontinenten  vertheidigt  zu  werden 
ptlegt,  kann  die  Thatsache  nicht  verdunkeln,  dass  in  solchen  Fallen  [the  genus  Scalesia 
et(  .  arc  mentioned  die  Organisationen  nicht  anzugeben  sind,  aus  deren  Variation  man 
sie  hervorgegangen  \orstellen  mochte.  Die  nahe  Verwandtschaft  hingegen,  welche  zwischen 
vielen  endemischen  Frzeugnissen  des  Archipels  und  denen  der  amerikanischen  Floren 
unleugbar  besteht,  kann  aus  dem  Pildungsgesetz  der  riiumlichen  Analogien  ebenso  wohl, 
als  aus  ei?iem  genctis(  hen  /usanmienhang  abgeleitet  werden.  Und  warum  sollte  iiberhaupt 
das  Festland  vor  den  Inseln  den  \'orzug  selbstiindig  entstandener  Organisationen  gehabt 
haben,  deren  crste  Erzeugung  in  den  friihesten  Perioden  der  Erdgeschichte  jeder  Moglich- 
keit  einer  \'ariation  vorausging?  waruin  sollte  sich  nicht  spiiter  und  an  verschiedenen 
(  )rten  sich  wiederholt  haben,  was  urs])riingli( :h  moglich  war  und  wovon  nur  die  Bedin- 
gungen   ein   noch   ungelostes    Kiithsel    geblieben   sind?  (II.  512—513). 

Such  ideas  have  little  more  than  historical  interest.  Drudk,  another  leading 
authority  on   plant  geograph)-,   was  more   in   accordance  with   modern  thought: 

.  .  .  dass  die  Flora  der  Inseln  nicht  nur  als  Transformationen  der  jetzt  lebenden  Kon- 
tinentaltloren  erfasst  werden  darf,  sondern  dass  auf  vielen  Inseln  unzweifelhaft  eine  Weiter- 


DERIVATION  OF  THE  FLORA  AND  FAUNA  365 

entwickelung  alter,  vielleicht  den  Charakter  einer  alteren  tertiaren  Periode  repriisentieren- 
der  Stammfloren  stattgefunden  hat,  welche  sich  hier  im  Schiitze  der  Abgeschiedenheit 
fern   von    dem    Einfluss  kontinentaler  Umwalzungserscheiniingen  sich  erhalten  konnten 

[jos.  128). 

Drude  agreed  with  Hooker  that  extinct  species,  known  as  fossils  in  Europe, 
were  the  ancestors  of  species  now  endemic  in  Macaronesia. 

With  regard  to  the  Pacific  islands  several  authors  have  discussed  the  question 
to  what  extent  endemism  has  been  of  a  progressive  kind  and  if  not  only  species  but 
also  isolated  genera  have  evolved  on  the  islands  from  a  limited  number  of  un- 
known ancestors.  Most  of  them,  in  spite  of  their  firm  belief  that  the  islands  are,  geologi- 
cally spoken,  very  recent,  regard  the  insular  biota  as  a  local  product;  the  first  arrivals 
had  had  time  to  give  rise  to  new  genera,  those  that  came  a  little  later  became 
new  species,  still  more  recent  ones  varieties  of  a  continental  species,  and  such  as 
arrived  in  our  era  have  not  had  time  to  change  but  are  expected  to  do  so,  because 
isolation  in  a  new  and  strange  environment  makes  them  adapt  themselves  by  changing 
their  genetic  structure.   Hayek  (jo^.  252)  expressed  this  very  clearly: 

Dass  diese  eingewanderten  Elemente  infolge  ihrer  Isolierung  eigene  Entwickelungs- 
richtungen  einschliigen,  die  sich  in  einer  oft  auftallend  grossen  Zahl  von  Endemismen 
aussern,   ist  ja  selbstverstandlich. 

Among  botanists  of  the  latest  decennia  Andrews  (6)  may  be  chosen  as  a 
representative  of  the  school  of  "rapid  adaptive  radiation". 

If  a  newcomer  belongs  to  a  primary  form  of  a  virile  genus  such  as  Acacia,  Coprosma, 
etc.  it  may  be  expected  rapidly  to  become  differentiated  into  varieties  and  species.  As 
Bentham  pointed  out  long  ago,  the  geographic  station  of  a  waif  or  colonist  imposes 
variations  upon  it  almost  from  the  moment  of  its  arrival.  Eucalypts  planted  in  New 
Zealand,  California,  and  other  places  present  marked  differences  from  the  forms  the  same 
species  possess  in  Australia.  In  the  second  ])lace,  if  the  plant  assemblage  into  which 
the  waif  or  colonist  arrives  be  a  result  of  long-continued  struggle  for  existence  such 
as  occurs  commonly  in  Holarctica  and  the  cosmopolitan  tropics,  then  the  opportunity 
for  the  development  of  new  forms  is  remote,  unless  the  new  arrival  itself  is  a  plastic 
form,  and  a  grand  example  of  the  survival  of  the  fittest.  If,  on  the  other  hand,  the 
newcomer  belongs  to  an  agressive  species  in  its  own  continental  setting,  then  it  has, 
all  other  things  being  equal,  an  excellent  chance  of  survival  and  of  differentiating  into 
new  forms  (p.  617). 

Andrews,  himself  an  Australian,  speaking  on  the  Hawaiian  flora,  mentioned 
Acacia,  Coprosma  and  other  genera  of  Australian  or  southwest  Pacific  origin  which 
developed  new,  endemic  forms  in  Hawaii,  where  they  take  a  prominent  part  in  the 
vegetation.  But  when  it  comes  to  Eucalypts  planted  in  California,  nobody  will,  I 
suppose,  consider  this  example  as  a  proof  of  the  origin  of  new  taxa  under  the 
influence  of  new  surroundings. 

There  is,  he  continues,  a  difference  in  the  physical  character  of  islands;  some 
did  not  encourage  the  colonist  to  vary  and  evolve,  others  did. 

If  the  island  ...  be  very  small  and  of  negligible  relief,  it  again  has  but  little  op- 
portunity for  differentiation,  and,  furthermore,  if  the  island  be  even  large  and  high,  but 
the  time  be  short  .  .  .  the  response  will  have  been  but  slight  (i.e.). 


366 


SKOTTSBERG 


If,  on  the  other  hand,  the  island  group  lies  in  the  tropics  and  is  large  in  area,  the 
islands  of  the  group  being  close  together,  if  the  vertical  relief  be  very  great  indeed  .  .  . 
if  the  {)recijMtation  be  very  variable  ...  if  the  soil  be  rich,  but  variable  in  porosity;  if 
the  j)lant  assemblages  into  which  the  waifs  or  colonists  entered  are  not  the  end  result 
of  severe  plant  competition;  then  the  stage  is  set  for  the  rapid  differentiation  of  primary 
types  of  agressive  genera. 

'i'he  evidence  available  suggests  that  these  genera  never  existed  as  such  on  the  land 
from  where  their  immediate  predecessors  were  derived,  but  that  virile  types,  of  the  families 
concerned,  arrived  as  waifs  from  Malaysia,  Australia,  New^  Zealand,  Central  and  North 
America.  Once  they  found  themselves  removed  from  their  former  severe  competition 
with  other  plants,  they  gave  rise  to  the  vigorous,  endemic  Hawaiian  genera  (p.  6i8). 

What  .Vndki-.ws  depicts  is  an  island  which  has  reached  maturity.  It  has  risen 
from  the  bottom  of  the  ocean  to  an  altitude  where  the  moisture  of  the  trade  winds 
is  condensed,  where  there  are  leeward  and  windward  slopes,  different  habitats  and 
a  rich  soil.  This  island  looks  back  on  a  very  long  history,  but  life  begins  to  arrive 
long  before  the  island  has  come  to  rest.  Aerial  plankton  will  bring  microscopic 
green  and  bluegreen  algae,  bacteria,  spores  of  all  kinds  and  also  air-borne  seeds, 
and  some  drift  may  be  washed  up  on  the  shores.  But  to  begin  with  there  is  little 
inimidit)'  and  hardly  anything  we  can  call  soil,  we  need  water  before  even  the 
most  j)rimitive  organisms  can  exist,  so  that  the  island  becomes  fit  to  receive  its 
first  settlers,  microscopic  algae,  then  mosses  and  lichens  and  mycelia  of  fungi,  to 
form  soil  where  the  first  seeds  can  germinate  and  start  to  form  an  incipient  vege- 
tation cover  and  an  abode  for  a  soil  fauna  and  flora.  Lava  cracks  in  cooling,  some 
water  ma)'  stand  in  the  fissures  which  form  the  starting  point  for  further  develop- 
ment, a  spectacle  we  have  before  our  eyes  where  streams  of  lava  are  still  formed. 
Of  higher  plants,  ferns  are  likely  to  be  among  the  first  to  get  established — GUPPY 
even  spoke  of  the  "era  of  ferns";  a  halophytic  Asplcniuui  is  the  only  living  thing 
observed  on  the  far-fiung  reef  Sala  y  Gomez  between  Easter  Island  and  South 
America. 

An  island  in  a  comparatively  recent  stage,  where  there  is  plenty  space  for 
new  settlers,  let  it  be  that  little  comfort  is  as  yet  offered,  would,  we  should  think, 
otter  good  opportunities  for  "virile  and  agressive"  immigrants  to  get  a  foothold 
and  to  become  the  ancestors  of  the  most  ancient  element  in  the  flora,  but  this  is 
not  what  Andki.ws  sa\s.  In  order  to  start  an  evolution  of  new  species  and  genera 
a  ver\'  great  vertical  relief,  a  variable  preci[)itation  and  a  rich  soil  are  the  conditions, 
l3ut  in  oriler  t(;  get  a  rich  soil  cover  we  must  have  a  closed  vegetation  cover,  also 
forest.  Wa I.I.AC!;  [2yS.  295),  sj)eaking  of  St.  Helena,  very  rightly  said  that  "no 
soil  could  be  retained  unless  protected  by  the  vegetation  to  which  it  in  great  part 
owed  its  origin",  and  the  same  is  true  everywhere.  If  the  change  in  environment, 
the  new  living  conditions,  are  the  cause  of  variation,  why  did  they  not  act  until 
the  island  was  already  more  or  less  stocked  with  plants.-  Were  there  no  virile 
species  among  the  earliest  immigrants  which  took  possession  of  the  land  and  formed 
the  oldest  element  of  the  flora.'  Uas  it  lost  among  the  later  arrived  aggressive 
newcomers.-  It  is  calculated  that  about  90%  of  the  Hawaiian  angiosperms  are  en- 
demic, most  of  them  belonging  to  endemic  genera  or  to  species  very  different  from 
their  continental  congeners.   \)o  some  of  the  most  remarkable  monotypical  genera 


DERIVATION  OF  THE  FLORA  AND  FAUNA  367 

represent  true  relics,  descendants  from  the  earliest  settlers,  whereas  the  other  are 
examples  of  progressive  endemism?  This  is  often  said,  but  it  is  not,  as  far  as  I 
can  see,  what  Andrews  means. 

In  the  opinion  of  zoologists  still  infected  with  Lamarckian  ideas  environment 
is  the  direct  cause  of  new  hereditary  characters.  Time  after  time  we  are  told  that 
as  soon  as  a  "germ"  happens  to  land  on  an  island,  it  gives  rise  to  something 
new.  However,  this  does  not,  Mayr  says,  imply  that  every  little  island  is  turned 
into  a  centre  of  evolution  {lyg.  216). 

The  small  and  usually  rather  isolated  islands  of  Polynesia  have  not  only  not  been 
new  centres  of  evolution,  like  the  Galapagos  or  Hawaiian  Islands,  but,  on  the  contrary, 
there  is  good  evidence  that  many  of  them  are  "traps".  Species  that  reach  these  islands 
are  doomed  to  extinction. 

This  is  peculiar.  There  are  many  small  islands  stocked  with  both  non-endemic 
and  endemic  species,  many  of  the  latter  stenotopic,  it  is  true,  but  quite  able  to 
hold  their  own  as  long  the  environment  remains  unchanged.  It  is  a  truism  that, 
if  a  newcomer  lands  on  an  island  where,  for  climatic  or  other  reasons  it  cannot 
live,  it  is  doomed  to  disappear  pretty  quickly,  but  there  is  no  reason  why,  once 
established  and  able  to  reproduce  itself,  it  would  become  extinct  as  long  as  the 
habitat  does  not  undergo  any  change  for  the  worse.  It  is  generally  known  among 
ornithologists,    Mayr  says,  that  island  birds  are  very  vulnerable.   He  continues: 

The  recent  considerations  of  Sewall  Wright  have  given  us  a  possible  key  to  this  curi- 
ous phenomenon.  Apparently  in  these  isolated  populations  there  is  more  gene  loss  than 
gene  mutation.  The  species  are  therefore  adjusted  to  an  exceedingly  narrow  limit  of 
environmental  conditions.  They  are  unable  to  respond  to  any  major  change  of  conditions 
and  must  die  if  such  a  change  occurs,  or  are  crowded  out  if  competitors  arrive. 

We  cannot  be  sure  that  mutations,  should  they  be  induced,  would  do  them 
any  good  and  we  need  not  assume  a  "gene  loss"  to  explain  why  organisms  unable 
to  escape  to  a  more  favourable  habitat  are  bound  to  become  exterminated  as  a 
result  of  "major  changes  of  conditions". 

Zimmerman  {2^8)  is  one  of  the  prominent  defenders  of  the  idea  that,  in  oceanic 
islands,  as  exemplified  by  the  Hawaiian  chain,  a  small  number  of  immigrants  has 
given  rise  to  a  comparatively  rich  fauna  and  flora;  the  proportion  between  genera 
and  species  attains  figures  expected  under  continental  conditions  but  certainly  not 
in  oceanic  islands.  He  admits  that  everything  did  not  necessarily  happen  on  the 
present  islands  as  we  behold  them,  for  differentiation  may  have  begun  on  some 
distant  land  and  proceeded  in  the  course  of  migration,  with  Hawaii  as  the  terminus. 
He  picked  out,  as  an  example,  a  large  curculionid  genus  ranging  from  Australia 
to  Micronesia  and  east  to  Marquesas;  most  islands  or  archipelagoes  have  their  own 
endemic  species,  which  have  developed  on  their  respective  islands.  He  thinks  that, 
if  within  a  varying  population,  one  pregnant  female,  not  carrying  the  gene  consti- 
tution of  the  entire  population,  gets  isolated  on  an  oceanic  island,  she  may  stand 
out  as  distinct  from  the  average — and  if  this  sequence  of  events  "be  accompanied 
by  conditions  conducive  to  isolation  and  survival,  rapid  and  diversified  speciation 
may    follow"    (p.  125).  This  process  is  repeated  on  island  after  island,  and  "the 


368  C.   SKOTTSBERG 

intensity  of  divergence  will  be  increased".  It  is  not  clear  to  me  why  the  intensity 
would  be  increased.  Now,  if  sufficient  time  has  elapsed  and  the  original  sources 
lia\e  been  eroded  down  and  perhaps  become  sterile  atolls,  their  faunas  will  have 
been  exterminated  and  on  the  newer  islands  segregates  without  obvious  ancestral 
relation  will  be  left.  This  development  explains  why  Hawaii  has  so  many  isolated 
endemic  t\'pes.  The  living  world  in  Hawaii  is  older  than  the  rock — in  a  way,  he 
sa)-s.  ( )uite  true,  but  it  is  not  true  that  this  possibility  has,  as  he  says,  been  entirely 
o\erl()oke(l  in  previous  discussions;  I  think  that  1  have,  on  repeated  occasions, 
exj)ressed  myself  ver\'  clearly  on  this  point,  even  if  1  do  not  agree  with  ZIMMERMAN 
when   it  comes  to  exj)lain   why  and  how  it  happened. 

The  biota  as  we  know  it  today  is  in  part  the  ultimate  product  of  a  progressional 
devclopnient  which  has  moved  and  evolved  along  great  insular  archipelagos  over  periods 
of  time  much  longer  than  the  ages  rec^uired  for  the  development  of  the  main  Hawaiian 
Islands  and  their  contemporary  biota.  Various  genera  and  stem  forms  of  groups  of  species 
iiKu  have  evolved  in  islands — now  atolls  such  as  some  of  the  leeward  Hawaiian  chain, 
the  great  Micronesian  archi])elagos,  the  Line  Islands — which  form  the  approach  to  Hawaii. 
However,  some  of  the  genera  and  the  bulk  of  species  known  today  have  originated  on 
our  present  main  islands  (p.  125)  ...  in  contemporary  Hawaii  there  are  preserved  rem- 
nants of  a  biota  which  has  in  ])art  develoj)ed  by  unique  methods  and  in  which  are  preserved 
forms  whicli  are  the  end  ])roducts  of  species  chains  that  carry  back,  through  archi- 
])elagos   now   worn   away,   to  geological  ages  indeterminant  (p.   126). 

In  few  words,  we  have  to  do  with  relict  as  well  as  progressive  endemism — 
nobody  objects  to  that.  A  genus  may  be  an  ancient  relic,  while  the  actual  species 
are  the  result  of  more  recent,  progressive  differentiation.  On  the  other  hand,  there 
is  no  reason  why  not  a  species  could  be  immensely  old  without  having  undergone 
any  j)crceptible  change. 

Zimmi:rm.\x  does  not  hesitate  to  conjure  up  all  the  sunk  archipelagos  he  needs, 
if  only  land  connections  arc  left  out  of  the  discussion.  Once  more  he  describes 
his  vision  on  p.  127  which  I  shall  permit  myself  to  quote,  with  the  obvious  risk 
of  tiring  out  the  reader. 

I  believe  that  the  great  atoll  chains  of  the  Pacific  may  hold  some  of  the  now  hidden 
(lues  to  the  stories  of  the  magnificent  biological  development  of  Polynesia.  Many  of  the 
peculiar  endemic  groups  of  the  Hawaiian  and  southeastern  Polynesian  islands  owe  their 
existence,  if  not  their  very  origin,  to  ancient  high  islands  of  the  one-time  splendid  archi- 
l)clagos  marked  by  (lusters  or  coral  reefs.  Surviving  lines  of  middle  Tertiary  and  of 
|)erhaj)s  even  older  continental  faunas  nuiy  have  had  their  germ  plasm  filtered  down 
through  su(  cessivcly  changing  generations  which  have  passed  successfully  through  island 
maturity  and  degradation  to  atoll  formation  and  have  carried  over  to  new  high  islands 
in  different  archipelagos.  Thus,  some  supposedly  old  types  such  as  certain  land  molluscs 
could  have  maintained  themselves  (but  evolving)  in  insular  isolation  through  long  periods 
ot  time  while  tlieir  continental  j)rogenitors  became  extinct  or  restricted  under  continental 
conditions. 

I  cannot  think  of  what  kind  of  higher  organisms  would  have  passed  success- 
fully through  island  degradation  down  to  atoll  stage;  they  must  have  left  for  new 
high  islands  long  before  their  abode  became  uninhabitable.  V.ven  if,  as  ZIMMERMAN 
thinks,   much   of  the  evolution   took  place  during  migration  from  island  to  island, 


DERIVATION  OF  THE  FLORA  AND  FAUNA  369 

specific  segregation  was  mainly  effected  after  arriving  at  the  final  station,  in  this 
case  Hawaii.  This  happened  yesterday  or  the  day  before,  geologically  spoken: 
"the  rate  of  erosion  is  such  that  these  main  islands  could  not  have  stood  here  as 
they  are  longer  than  from  a  period  late  in  the  Tertiary"  (p.  121).  "Explosive 
speciation"  set  in  during  late  Pliocene  and  must  have  increased  during  "the  great 
Pleistocene  erosion  which  has  left  such  a  spectacular  and  rugged  topography  in 
its  wake.  New  land  open  to  colonization  is  conducive  to  speciation"  (p.  122). 

This  late  and  rapid  differentiation  is  illustrated  by  ZIMMERMAN  for  the  land 
snails  pp.  98-101 : 

Helicidae.  59  species  developed  from  one,  or  possibly  two  original  immigrant 
stocks. 

Pupillidae.  Possibly  4  ancestral  species  gave  rise  to  the  86  Hawaiian  forms. 

Cochliocopidae-Cochliocopinae.  One  immigrant  of  Cochliocopa  stock  could  have 
given  rise  to  the   142  forms. 

C.-Amastrinae.  294  forms  apparently  developed  from  one  basic  stock. 

Tornatellinidae-Tornatellininae.  117  forms  derived  from  4  or  fewer  ancestral 
forms. 

T.-Achaiinellinae.  It  appears  certain  that  this  subfamily  had  its  origin  and 
development  in  the  Hawaiian  area  and  all  of  the  215  forms  may  have  been  derived 
from  a  common  tornatellinid  ancestor. 

FOSBERG,  in  a  chapter  contributed  to  Zimmerman's  book,  tried  to  fix  the 
number  of  ancestors  of  the  Hawaiian  angiosperms.  His  method  is  quite  simple: 
if  the  species,  few  or  many,  of  a  certain  genus  present  the  appearance  of  a  more 
or  less  homogeneous  group,  only  a  single  ancestor  is  made  responsible  for  the 
segregation;  if  subgenera  or  sections  are  distinguished,  we  must  count  with  the 
same  number  of  ancestors  as  of  taxonomic  groups  within  the  genus.  The  method 
seems  a  little  too  easy;  possibly  we  are  confronted  with  a  rather  complicated 
question,  the  solution  of  which  I  am  not  going  to  attempt. 

Setchell    (^218)  uttered  some  sensible  words  on  migration  and  endemism: 

Where  endemism  of  the  degree  of  ordinal  or  family  endemism  occurs  on  oceanic 
islands,  we  may  feel  strongly  inclined  to  believe  that  evolution  of  such  degree  took 
place  on  the  continental  area  which  was  the  source  of  the  original  migration  and  not 
on  the  island  where  now  found,  the  original  becoming  later  extinct,  leaving  the  migrant 
as  an  endemic.  The  same  is  true  of  generic  endemism  or  even  specific  endemism  of  a 
strong  type,  that  is  when  representing  an  isolated  or  aberrant  species  under  the  genus 
(p.  874).  .  .  .  To  assume  that  insular  conditions  originate  new  forms  is  to  overlook  what 
has  taken  place  on  continents  (p.  875). 

This  is,  however,  what  so  many  authors  do.  They  claim  that  oceanic  islands 
follow  their  special  laws,  that  a  plant  or  an  animal  which  happens  to  land  far  away, 
will,  as  it  were,  lose  its  balance;  hidden  factors,  repressed  as  long  as  they  lived 
on  their  fatherland  under  "severe  competition",  are  set  free  and  allow  them  to 
develop  their  inherent  possibilities,  they  are  not  subjected  to  any  struggle  for 
existence  in  their  new  environment.  Says  ZIMMERMAN:  "The  environment,  of  course, 
plays  an  all-important  part  in  the  development  of  species  ...  it  is  generally  agreed 
that  profound  changes  have  been  effected  on  organisms  by  environment"  (p.  187). 

24  -  557857   The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.   Vol.  I 


370 


SKOTTSBERG 


This  is  pure  Lamarckian  langua^^^c.  \Vc  may  say,  of  course,  that  change  of  environ- 
ment may  cause  mutations,  but  our  experience  tends  to  prove  that  only  a  fraction 
of  I  °o  are  \aluable,  the  remainder,  if  not  deleterious,  at  least  indifferent.  Mutations, 
gene  losses.  Inbridization,  polyploidy  and  so  forth,  all  may  have  their  share,  but 
they  are  only  ripples  on  the  surface.  Xo  theory  has  been  able  to  penetrate  to 
the  nucleus  of  the  j)r()blem.  Anyhow  we  have  no  reason  to  think  that,  in  its  pro- 
duction of  species  and  genera,  families  and  orders.  Nature  has  followed  other  lines 
in  islands  than  in  continents.  A  prominent  Swedish  geneticist,  after  a  life-time's 
s|)eculati()n  on  the  causes  generally  accepted  as  responsible  for  the  "origin  of  species", 
rejected  all  of  them;  unfortunately  he  threw  the  egg  away  with  the  shell  and  con- 
vinced himself  that  there  had  been  no  evolution  at  all.  He  did  not,  however, 
revert  to  an  omnipotent  creator  but  invented  a  new  and  entirely  revolutionary  theory; 
if  there  ever  was  a  stillborn   one,   it  was  this  (iS'6). 

I  guess  we  can  take  it  for  granted  that  no  peculiar,  outstanding  types  were 
created  on  young  volcanic  islands,  whatever  their  present  faunistic  and  floristic 
status  may  be  like.  Their  basic  stock  is  much  older  than  the  rocks  and  inseparably 
connected  with  the  great  continental  faunas  and  floras.  Wallace,  however,  made  a 
distinction  between  island  and  continental  history.  He  emphasized  that  evolution 
has  recjuired  an  enormously  long  time  to  produce  the  present  status,  that  from 
the  Cretaceous  until  now  nothing  of  a  revolutionary  character  had  happened,  families, 
genera  antl  in  cases  even  species  still  living  date  from  early  Tertiary  at  least — 
but  on   islands  the  great  period  of  creation  was  repeated  during  the  last  epochs. 

l^'inally  I  shall  quote  some  selected  passages  from  Stebhixs'  book  on  variation 
and  evolution   Uiy). 

The  ditterentiation  of  orders  and  families  of  flowering  plants  through  the  action  of 
natural  selection  under  ])resent  conditions  is  well-nigh  impossible.  .  .  .  All  the  trends 
leading  to  the  difterentiation  of  families  of  flowering  plants  probably  took  place  simul- 
taneously and  at  a  relatively  early  stage  of  angiosperm  evolution.  For  instance,  both 
distributional  and  ])aleontological  evidence  indicates  that  the  Com])Ositae,  the  most  highly 
sj)e(  iali/.ed  family  of  dicotyledons,  already  existed  in  the  latter  part  of  the  Cretaceous 
])erio<l,  and  distributional  evidence  indicates  a  similar  age  for  the  most  advanced  families 
of  niono( otyledons,  the  Orchidaceae  and  Cramineae.  It  is  likely,  therefore,  that  the 
major  part  of  angios])erm  evolution,  involving  the  ])rincipal  trends  in  the  modification 
of  the   flowers,   took    i)lace   during  the   Meso/.oic  era   (pp.  501-502). 

The  g\-nmosj)erms  prove,  he  remarks,  that  neither  great  antiquity  nor  rigid 
stability  necessarily  leads  to  senescence,  but  here,  as  in  other  cases,  a  stenotopic 
character  has  resulted  in   restricted  areas  and  also  in  extinction. 

Isolated  records  excepted — the  latest  discovery  is  a  palm  from  a  Triassic  stratum, 
if  correctly  determined — angios|)ermic  fossils  are  found  in  greater  quantity  from 
younger  Cretaceous  and  then  in  many  surprisingly  modern  types,  l^^vidence  is  strong. 
Si  KP.I'.INS  has  found,  that  evolution  was  rapid  during  one  period  and  slower  during 
another.  At  the  end  f)f  the  Cretaceous  and  the  beginning  of  the  l^Locene  the  number 
of  modern  types  increased  rapidly.  In  eocene  deposits  in  North  America  and  Eurasia 
"the  majority  of  the  s[)ecies  belonged  to  or  closely  approximated  modern  genera" 
(p.  520).  This  conclusion,  mainly  based  on  leaf  impressions,  is  strengthened  by  the 


DERIVATION  OF  THE  FLORA  AND  FAUNA  37I 

famous  London  clay  from  lower  Eocene,  where  a  wealth  of  fruits  and  seeds  were 
found.  The  corresponding  fossil  floras  of  California  include  an  increasing  number 
of  plants  that  cannot  be  distinguished  from  living  genera,  and  abundant  evidence 
shows  that  since  late  Pliocene  "new  species  of  woody  plants  have  been  added  to 
the  Californian  flora  only  in  such  large  and  complex  genera  as  Eriogonum,  Ceanoihus, 
Arctostaphylos,  and  various  Conipositae''  (p.  521).  The  fossil  floras  of  East  Asia  point 
in  the  same  direction,  the  rate  of  evolution  on  woody  plants  has,  since  the  middle 
of  this  period,  been  as  slow  or  slower. 

The  most  likely  inference  on  the  basis  of  all  available  evidence  is  that  most  of  the 
woody  species  of  to-day  have  existed  for  five  million  years  or  more,  and  that  the  evolution 
of  the  genetic  isolation  mechanism  separating  them  took  place  largely  during  the  early 
and  middle  parts  of  the  Tertiary  period  (p.  522). 

With  regard  to  the  herbaceous  floras  there  is  evidence  for  a  rather  rapid  evo- 
lution during  the  later  Tertiary.  Stebbins  quotes  the  results  obtained  by  Elias 
in  fossil  caryopses  dating  from  lower  Miocene  to  middle  Pliocene  and  showing  a 
distinct  progression  until  the  most  recent  ones  cannot,  to  judge  from  the  illustrations 
redrawn  by  Stebbins,  be  distinguished  from  Stipa  or  Piptochaetium,  an  evidence 
from  palaeontology  which  strengthens  the  theory  of  SiNNOT'r  and  Bailey  already 
referred  to. 

Isolation  as  a  cause  of  evolution. — There  is  no  reason  why  a  species,  which 
has  migrated  to  an  island,  should  change  and  give  rise  to  new  forms  only  because 
it  is  spatially  isolated.  A  form  of  a  varying  population  which  gets  isolated  on  an 
island,  may  possess  characters  making  it  stand  out  as  a  more  or  less  well-marked 
form.  As  Jordan  {131, 132)  says,  isolation  is  not  the  direct  cause  of  the  origin. 
Spatial  relation,  Stebbins  remarks, 

may  persist  over  long  periods  of  time  without  causing  the  isolated  populations  to  diverge 
from  each  other  enough  to  become  recognizably  distinct  or  even  different.  .  .  .  the  nature 
of  many  distributions  strongly  suggests  that  some  of  these  disjunct  segments  of  the  same 
species  have  been  isolated  from  each  other  for  millions  of  years.  There  is  some  reason 
for  believing,  therefore,  that  geographic  isolation  alone  does  not  result  even  in  the 
formation  of  subspecies.  ...  In  small  populations,  which  are  particularly  frequent  on 
oceanic  islands,  spatial  isolation  is  the  usual  precursor  to  divergence  in  non-adaptive 
characters  by  means  of  genetic   drift  or  random   fixation   (p.  197). 

Stebbins  also  points  out  that  spatially  isolated  races  or  species  usually  are 
separated  also  by  ecological  barriers,  because  different  areas  also  differ  ecologically; 
distance  may  not  be  the  original  isolating  factor,  separation  and  differentiation  may 
be  due  to  the  selective  effect  of  ecological  and  climatic  factors  before  geographical 
separation  occurred.  The  Hawaiian  Islands  offer  a  wide  field  for  a  study  of  isolation 
and  segregation  resulting  in  local  endemism  (^^^).  The  continental  flora  inherited 
by  a  Great  Hawaii  included  numerous  populations  gradually  taking  possession  of 
Ihe  new  soil,  a  process  of  very  long  duration,  volcanic  activity  progressing  from 
west  to  east,  and  when  separation  took  place,  different  forms  and  species  became 
isolated  on  different  islands.  The  wonderful  systems  of  valleys,  effectively  separated 
by  high  and  very  steep  ridges,  furthermore  promoted  segregation.  Little  has  been 


372  C.   SKOTTSBERG 

done  to  examine  the  <^enoty})ical  constitution  of  this  astonishing  assemblage  of 
\oca\  forms;  some  may,  for  ail  we  know,  be  ecotypes,  but  experimental  studies 
have  not  been  made.  A  limited  number  of  hybrids  have  been  described,  but  we 
know  nothing  of  their  behaviour.  Anyhow,  the  floristic  difference  between  the  islands 
is  very  striking  and  hardly  in  favour  of  the  theory  of  transoceanic  migration,  for 
tlie  majority  t)f  organisms  seem  to  have  been  unable  to  cross  even  the  straits 
separating  the  islands,  and  biotic  factors  preventing  establishment  can  hardly  be 
made  responsible  in  so  many  cases. 


Chapter  IX. 
Juan  Fernandez — oceanic  or  continental  ? 

To  W'ai.i.ack  as  well  as  to  the  majority  of  biogeographers  the  Juan  Fer- 
nandez Islands  were  typically  oceanic  in  spite  of  their  moderate  distance  from 
South  America.  They  had  the  advantage  of  antiquity,  WALLACE  remarks,  for 
the  means  of  transmission  had  formerly  been  greater  than  now,  their  surface  was 
varied,  soil  and  climate  favourable,  "offering  many  chances  for  the  preservation 
and  increase  of  whatever  plants  and  animals  had  chanced  to  reach  them"  (p.  287), 
I  lad  the  character  of  Masafuera  been  known  to  him  he  might  have  been  less 
optimistic.  The  land-shell  fauna,  entirely  endemic,  testified  to  the  great  age  of 
the  islands,  for  none  had  been  introduced  for  so  long  a  period  that  all  which 
did  come  had  given  rise  to  new  forms — or  were  the  last  of  a  fauna  extinct 
on   the  continent. 

Joiiow  (/jo)  based  his  opinion  on  Wallace;  when  discussing  the  dispersal 
agencies  and  the  morphology  of  the  diaspores  his  starting-point  was  /wo  islands, 
Masatierra  })lus  Santa  Clara  and  Masafuera,  which  has  risen  separately  from  the 
deef)  sea.  It  is  strange  that  he  never  thought  of  another  possibility,  because  the 
geologist  who  went  with  him  and  who  wrote  a  chapter  on  the  geology  of  Masa- 
tierra, claimed  to  have  discovered,  in  one  place,  a  fundament  of  rocks  older  than 
the  omnij)rcsent  young  basalt — ^JoilOW  could  not  know  that  the  interpretation 
of  this  stratum  was  false,  as  later  shown  by  OUENSEL  (J02).  After  his  visit  to 
the  ncs\  enturadas  Joiiow  modified  his  opinion;  the  submarine  ridge  uniting  these 
little  islands  with  Juan   T'ernandez  had   then  been   discovered: 

Die  iiiUer  [.'Icicher  P.reite  mit  dem  Hafen  Caldera  und  in  derselben  Entfernung 
vom  Kontinent  wic  Juan  I'crnandc/.  ^elegene  Inselgruppe  ist  vulkanischen  Ursprungs 
und  stcilt,  wie  die  von  dcin  Mitglicd  der  Expedition,  Herrn  Chaigncau,  ausgeliihrten 
Eotungen  crgahcn,  die  iiher  Wasscr  befindiichen  hochsten  (iipfel  einer  im  Ubrigen 
iinterseeisch  verlaufenden  liergkcttc  dar,  welcher  auch  die  Inseln  der  Juan-Fernandez- 
(iru])pe  als  siidli^  hstc  (iiptel  angehoren.  Aus  dem  Vergleiche  der  Eloren  und  Faunen 
heider  .\r(  hipelc,  \vcl(  he  trot/,  der  grossen  klimatischen  Verschiedenheiten  frappante 
X'crwandtschaft  autweisen,  crgicbt  .sich  mit  zwingender  Notwendigkeit  die  Hypothese, 
dass  die  zwei  Inselgriii)i)en  in  der  Vorzeit  mit  einander  in  Landverbindung  gestanden 
hahen  und  dass  ihre  Isolierung  die  Folge  einer  stattgehabten  Senkung  jener  Bergkette 
ist  isso-  259). 


DERIVATION  OF  THE  FLORA  AND  FAUNA 


373 


In  this  assumption  he  may  be  quite  right,  but  it  finds,  as  already  pointed 
out  by  Reiche  (20J.  269),  httle  support  in  the  flora,  for  the  two  groups  have 
only  a  single  species  in  common  (2^1),  and  the  endemic  genera  and  species  are 
very  different.  JOHOW's  belief  in  their  permanent  isolation  from  the  continent 
remained  unchanged. 

In  his  tables,  Continjente  A  and  B,  pp.  218-220,  is  a  column  indicating 
(not  always  with  sufficient  accuracy)  the  nature  of  the  fruits  or  seeds,  and  another 
stating  the  probable  dispersal  agent.  Of  the  143  vascular  plants  known  at  that 
time,  all  the  ferns  and  34  phanerogams  were  supposed  to  have  been  wind-borne, 
61  had  been  transported  by  birds,  either  inside  or  adhering  to  their  feet  or 
plumage;  only  one,  SopJiora  ''tetraptera\  with  winged  pods,  had  drifted  with 
the  current.  Five  [Eryngium  bupleuroides  and  sarcophyllum,  Apiuin  fernandezianum, 
Colletia  spartoides  and  Fagara  (Zanthoxylum)  mayu\  offered  too  serious  difficul- 
ties to  make  their  presence  in  the  islands  explicable,  but  I  cannot  find  that  they 
are  more  "impossible"  than  many  of  the  others.  In  an  earlier  paper  (^j-j)  I  sur- 
veyed the  nature  of  the  diaspores;  I  shall  not  return  to  this  question  here  but 
only  repeat  that  almost  one  third  of  the  flowering  plants  show  no  special  adap- 
tation to  any  particular  mode  of  dispersal  across  the  water,  and  I  shall  add  a 
few  remarks  on  some  knotty  cases.  Among  the  island  Compositae,  a  family  known 
to  be  well  adapted  to  wind  dispersal,  are  several  species  which  lose  their  pappus 
when  the  achenes  are  still  enclosed  in  the  involucre,  or  where  it  is  reduced  to 
uselessness.  Only  one  species  of  Sophora,  called  tetraptera,  was  recognized  by 
JOHOW  and  recorded  from  Chile,  Juan  Fernandez,  Easter  Island  and  New  Zealand. 
Genuine  tetraptera  is  restricted  to  New  Zealand,  but  this  is  of  less  importance, 
the  pods  have  narrow  wings  (poorly  developed  in  some  forms)  which  were  sup- 
posed to  help  the  pods  to  keep  afloat,  but  as  they  open  on  the  tree  and  drop 
their  rather  heavy  seeds  to  the  ground,  the  wings  serve  no  purpose.  Halorrhagis 
is  a  somewhat  similar  case.  JOHOW,  and  others  before  and  after  him,  identified 
the  plant  found  on  Masatierra  with  H.  erecta  (alata),  a  New  Zealand  species  with 
four  narrow  wings  on  the  fruit,  and  he  did  not  hesitate  to  regard  it  as  wind- 
borne.  His  erecta  is,  however,  an  endemic  species;  there  are  two  more  on  Masa- 
fuera  (see  22g,  with  illustrations),  and  their  fruits  are  quite  or  almost  unwinged. 
There  can  be  no  question  of  an  adaptation  to  wind  carriage.  The  same  applies 
to  Selkirkia,  another  of  JOHOw's  anemochorous  plants.  Edible  fruits,  if  not  too 
big,  will  be  swallowed  by  birds,  but  the  indigestable  stones  and  hard-coated 
seeds  will  be  dropped  before  the  islands  are  reached,  and  the  diet  of  the  wide- 
ranging  and  fast-flying  sea  birds  is  another.  Epizoic  transport  is  possible  in  a 
small  number  of  cases,  but  otherwise  the  dispersal  mechanisms,  many  of  them 
quite  wonderful,  serve  to  maintain  the  population  within  its  range,  and  not  to 
stock  distant  islands. 

Other  advocates  of  the  permanent  isolation  of  Juan  Fernandez  are  e.g.  Plate 
{199),  GouRLAY  [301),  Burger  [41),  and  Goetsch  [124).  Plate  drew  his  con- 
clusions from  a  comparison  between  the  littoral  faunas  of  Juan  Fernandez  and 
the  opposite  coast: 


374 


C.   SKOTTSBERG 


Die  Seichtwasser-Faiina  weist  deutlicher  als  die  geologischen  Verhiiltnisse  darauf 
hin,  <lass  Masatierra  in  der  That  als  eine  ozeanische  Insel  anzusehen  ist,  die  nie  in 
Zusamnienhang   niit   deni    I-"estland   gestandcii   haben   kann  .  .  .   (p.    228), 

but  this  did  not  prevent  liiin  from  believing  that  Masatierra  and  Masafuera  once 
formed  a  sini^le  lari^e  island,  in  spite  of  the  distance  (92  miles)  and  the  deep 
water  that  sej)arates  them  (p.  222).  No  more  will  be  said  here  about  a  connection 
with  the  mainland;  I  have  already  discussed  tliis  question  from  an  algologist's 
view|)<)int,  and   with   a  difterent  result  (2jS). 

The  reason  why  (ioLRl.AV  rejected  all  land  connections  was  his  conviction 
of  the  easiness  with  which  plants  and  animals  are  transported;  the  under-title  of 
his  article  reads  "Plants  make  ocean  voyages".  He  had  little  faith  in  the  birds, 
wind  and  sea  were  the  principal  means  of  transportation,  and  as  so  many  others 
he  j)()inted  to  Krakatau  as  the  classical  example.  The  distance  was  very  small, 
it  is  true,  but  winds  and  currents  were  favourable  along  the  coast  region  of 
South  .America,  and  the  flora  was  derived  from  southern  Chile.  Of  the  genera  and 
sj)ecies  cjuite  without  relations  not  only  in  Chile,  but  in  all  America,  he  said 
notliing. 

Hi  K(;i:k,  who  had  visited  the  islands,  followed  JoiJOW;  they  were  typically 
oceanic.  After  telling  us  about  some  of  the  most  remarkable  endemics,  he  exclaims: 

W'er  brachte  die.sen  entlegenen  Stiitten  solch  kostliches  Geschenk?  Die  Stromung, 
die  \'(')gel  iind  vor  allem  die  Winde.  Sie  beluden  sich  niit  Sporen  und  Samen.  .  .  . 
selhst  voiii  I'euerlande  empfangenes  gedieh.  Doch  aiis  viel  weiteren  Fernen  kamen  die 
Kinwandcrer,  auch  vom  tropischen  x^merika,  Polynesien,  ja  sogar  von  Aiistralien  und 
Neuseeland  und   den   Inseln  des  Indischen  Ozeans  (pp.    17-18). 

The  majority  of  the  newcomers  remained  true  to  their  stock  in  spite  of  the 
changed  conditions,  but  others  changed  and  some  took  such  a  fancy  to  the 
climate  that  from  being  herbs  they  became  trees.  1  shall  leave  these  speculations 
without  comment;  we  have  better  reason  to  observe  GOETSCIl's  paper  which, 
in  sj)ite  of  being  fairly  recent,  contains  many  amazing  statements  not  supported 
by  facts.  Ignoring  OLl'.NSlll/s  report  on  the  geology,  he  tells  us  that  the  islands 
rest  "auf  einem  Sockel  von  grimlichem  Andesit,  der  auch  die  Hauptmasse  der 
Anden  bildet".  lie  had  seen  my  writings  on  the  flora;  the  use  he  made  of  them 
mav  l)e  illustrated  by  a  couple  of  examples.  Among  the  plants  introduced  from 
luirope  he  mentions  "Aromo  aus  Castilien" — this  is  a  local  name  for  the  en- 
demic Arjai-a  fcriiaiideziaiuA  Me  rejects  all  land  connections,  the  sandal-wood 
came  from  the  Past  Indies;  he  had  not  observed  that  it  belongs  to  a  quite  dif- 
ferent section  than  SautaliiDi  albiDu.  The  arborescent  Conipositae  and  Plaiitago  had 
originated  in  the  islands,  '' Plaiitago  feriiaiidezia  wx^iX  Skottsbcrgii,  \-2  v[\\\oz\\vc\\\. 
20  cm  langen  und  3  ^2  cm  breiten,  an  der  Spitze  des  Stammes  stehenden  Blat- 
tern'  .  lie  overlooked  that  the  so-called  /'.  Skoitsbergi'i  is  a  modest  annual  and 
a  form  of  the  common  Chilean  /rianata,  a  variable  species.  But  GoETSCII  knows 
what  happened  in  the  islands:  "Dass  der  europiiische  Wegerich  auf  Juan  F'ernandez 
mcterhohe  HUitenahren  und  fusslange  Blatter  tragt,  weist  auf  die  P^ntwickelung 
hin,  die  seine   X'erwandten   einstmal  nahmen"   (p.  29).  This  refers  to  P.  lanceolala., 


DERIVATION  OF  THE  FLORA  AND  FAUNA  375 

which  luxuriates  in  the  islands,  but  cannot  prove  that  the  arborescent  P.  fer7ian- 
dezia  evolved  from  one  of  the  many  Chilean  species  with  which,  from  a  system- 
atic viewpoint,  it  has  nothing  to  do. 

GOETSCH  was,  perhaps,  more  familiar  with  zoology,  but  where  he  got  the 
impression  that  the  lower  fauna  was  extremely  poor,  I  cannot  tell,  unless  Joiiow's 
meagre  list  was  his  only  source.  A  considerable  number  of  land  shells  and  in- 
sects had  been  described,  and  everything  tended  to  show  that  they  represented 
but  fragments  of  the  fauna.  The  most  efficient  dispersal  agent  was,  in  this  case, 
the  current. 

Wir  wissen,  dass  die  warmen  Stromungen  Ozeaniens  bis  an  die  Insel  reichen 
und  australische  Pflanzen  und  Tiere  verfrachten  konnen.  Aus  diesem  Grunde  muss 
zwar  die  Insel  auch  ausserhalb  des  sog.  Humboldtstroms  liegen,  der  von  der  Antarctis 
bis  zum  Aequator  und  der  siidamerikanischen  Westkiiste  lauft.  Wir  wissen  aber  nun 
seit  neueren  Untersuchungen,  dass  dieser  sog.  Humboldtstrom  keineswegs  ein  conti- 
nuierliches  Fliessen  in  nordlicher  Richtung  ist,  sondern  dass  Oberflachen-Wasser  vom 
Lande  wegstromt  und  durch  kaltes  aus  tieferen  Schichten  ersetzt  wird.  Der  chilenische 
und  patagonische  Einfluss  ist  dadurch  gesichert. 

From  what  he  just  said  we  learn  that  the  Humboldt  current,  whatever  it  is, 
and  the  upwelling  cold  water  do  not  reach  as  far  as  Masatierra.  On  the  other 
hand,  it  is  not  true  that,  as  GOOU  says  {log.  220),  Juan  Fernandez  lies  entirely 
outside  the  Pacific  beach  drift.  Drift-wood  is  found  on  the  western  side  of  Masa- 
fuera,  but  it  shows  very  rough  handling  and  must  have  been  months  or  rather 
years  under  way,  and  if  one  has  seen  the  place  one  feels  convinced  that  not  a 
single  plant  owes  its  presence  in  the  island  to  this  mode  of  conveyance,  not  to 
speak  of  the  fauna.  But  GOETSCli  believes  that  also  the  tropical  element  in  the 
flora  has  arrived  by  sea;  he  continues: 

Wir  wissen  endlich,  dass  vom  Norden  her  dann  dem  Humboldtstrom  eine  warme 
Meeresstromung  entgegenwirkt,  der  sogenannte  Nino,  der  sich  periodisch  in  einzelnen 
Jahren  so  geltend  machen  kann,  dass  seine  Wirkung  bis  Valparaiso  gespiirt  wird.  .  .  . 
Damit  findet  auch  der  tropische  Einschlag  in  Fauna  und  Flora  seine  Erklarung  (p.  38). 

This  cannot  refer  to  the  warm  currents  of  Oceania  mentioned  before,  because 
they  came  from  Australia  and  did  not  bring  the  neotropical  element.  Does  he 
think  of  the  equatorial  counter  current  which,  occasionally,  would  extend  its 
influence  to  the  South  American  coast  and  become  deflected  south,  thereby 
fetching  new  passengers.^  This  current  does  not  reach  the  coast. 

Other  authors  have  acted  as  spokesmen  for  the  bridge-builders.  W.  A.  BRYAN — 
not  having  access  to  his  book  I  quote  a  newspaper  article  written  by  him  (joo) — 
believes  that  a  "prehistoric  continent"  once  embraced  the  Pacific  islands,  a  theory 
based  on  the  distribution  of  the  land-snails.  Juan  Fernandez  belonged  to  the 
same  geological  period  as  Hawaii  and  their  living  world  proves  that  at  some 
time  they  were  united.  The  general  biological  character  is,  he  says,  the  same, 
but  there  is  a  slight  difference  in  the  species.  For  details  I  refer  the  reader  to 
261,  where  I  have  reviewed  his  book.  Bryan  was  professor  of  zoology  and  geology 
in  the  University  of  Hawaii  and  had  visited  Juan  F'ernandez,  but  he  does  not 
appear  to  have  profited  very  much  by  his  visit. 


376 


C.   SKOTTSBERG 


Where  HkANClll  [^^oj)  got  his  idea  of  the  island  flora  I  cannot  tell,  in  any 
case  not  from  facts,  in  spite  of  having  visited  Masatierra: 

Coiiio  curiosidad  cientifica  se  piiede  decir  que  Masatierra  no  seria  sino  una  de 
las  tantas  vetas  del  gran  continente  sumergido  en  el  Oceano  Pacifico  en  epocas  en 
(jue  el  continente  sudamericano  era  un  lecho  de  mar,  y  per  consiguiente  la  isla  serfa 
mas  vieja  (pie  el  suelo  de  la  madre  patria.  Como  comprobaci6n  estil  la  ausencia  de 
los  \olcanes,  los  esarpados  farallones  que  muestran  las  capas  geol6gicas,  el  desliza- 
miento  de  las  rocas  de  la  epoca  glacial,  y  sobre  todo  la  flora  unica,  di versa  del  conti- 
nente y  (juc   i)uede  solo  acercarse  a   la   flora   polinesica  y  australiana  (p.    14). 

I  guess  we  can  agree  with  him  that  this  is  all  very  curious,  but  it  is  not 
scientific. 

To  complete  the  picture  we  have  better  consult  Arldt's  great  work  (<?) 
in  which  the  author  draws  his  conclusions  from  the  land  fauna.  Arldt  was  one 
of  the  great  bridge-builders,  but  his  Pacific  bridges  were  of  old  date,  and  he 
thought  that  Juan  I'^ernandez  had  been  isolated  since  late  Cretaceous  times,  so 
that  onl}'  the  most  ancient  element  was  supposed  to  have  arrived  over  land  (p.  322). 

Wie  Tristan  da  Cunha  einen  Rest  des  Siidatlantis  darstellt,  so  die  Juan  Fernandez- 
(iruppe  einen  der  Ozeanis  der  siidpazifischen  Landbriicke.  Das  beweisen  nicht  sicher 
die  \'6gel  .  .  .  auch  nicht  die  Schmetterlinge,  von  denen  die  Nymphaliden  durch  die 
chilenische  J^ranieis  carye  vertreten  sind,  die  Pyraliden  durch  Scoparia  ragonoti, 
ncben  denen  noch  weitere  chilenische  Formen  genannt  werden.  Alle  hiitten  auch  trans- 
marin  die  Inseln  erreichen  konnen.  Wichtiger  sind  die  Landschnecken,  deren  Gattungen 
fast  alle  auch  in  Chile  vorkommen.  Vertreten  sind  z.  B.  die  Succineiden  und  Ferrusa- 
ciden,  sowie  die  Kndodontiden.  Diese  sind  hier  mit  Aniphidoxa  vertreten.  Deren  typische 
Cntergattung  ist  auf  Juan  Fernandez  endemisch.  Stephatwda  besitzt  6  endemische  Arten 
aut  den  Inseln,  andere  auf  Chiloe,  in  Chile,  Patagonien,  Feuerland,  Argentinien,  Para- 
guay, Brasilien  und  eine  auch  auf  Kcrguelen.  Dagegen  gehort  die  auf  Juan  Fernandez 
endemische  J^'ernandezia  zu  den  sonst  hawaiischen  Amastriden,  die  auf  Hawaii  auch 
ihre  ein/igen  niiheren  Verwandten  in  den  Achatinellidcn  besitzen.  Hier  kann  nur  eine 
Au>l)rcitung  iiber  Land  angenommen  werden.  Von  anderen  Landtieren  findet  sich  auf 
Juan  Fernandez  v..  W.  noch  der  'J'auscndfiissler  Gecphilus  latkollis.  Endlich  sind  sie  auch 
von  cinem  ( )ligo(  haeten  erreicht  worden,  der  Ocnerodrilinen  Kerria  saltensis,  ^\^  dMQ\\ 
in  Chile  vorkommt.  Hier  wiire  eine  jiingere,  iiberseeische  Einwanderung  denkbar.  Aber 
Kc) ) id  ist  au<  h  alt  genug,  dass  sie  auf  dem  Landwege  nach  Juan  Fernandez  gekommen 
scin    kann.    Auch    die   Flora  der   Inseln   zeigt  interessante  Beziehungen   (])p.    324-325). 

As  could  be  expected,  Arldt's  south  Pacific  bridge  was  supported  mainly  by 
tile  land  molluscs,  and  they  made  him  extend  his  bridge  across  the  equator  to 
ilawaii.  There  is  in  the  Juan  P'crnandez  flora  some  perplexing  affinities  with 
Hawaii  that  scctn  to  defy  all  attempts  to  an  explanation.  The  occurrence  of  two  (not 
only  one)  endemic  species  of  XcsogeopJiilus  (formerly  a  subgenus  of  6'r^////7//'j)  is  in- 
teresting, Kcrria  is,  j)erhaps,  less  important.  Other  striking  cases  of  disjunction  were 
already  referred  to  in  the  lists  of  evertebrates  above.  It  is  a  pity  that  Arldt  over- 
looked the  leech  Xi'sophilaoiioji,  but  also  from  what  he  says  about  the  butterflies  it  is 
evident  that  the  numerous  zoological  papers  forming  vol.  II  of  "The  Natural  History 
of  Juan  F'ernandez  and  blaster  Island"  had  escaped  his  notice.  The  leech  would, 
I  sup|)ose,  have  furnished  him  with  one  of  the  most  eloquent  evidences  of 
former    land    connections.    Xor    did  he  know  the  botanical  volume  of  this  work, 


DERIVATION  OF  THE  FLORA  AND  FAUNA  377 

for  he  gives  a  summary  of  my  19 14  paper  (22^).  He  agrees  on  the  great  age 
of  the  "Palaeopacilic  element";  second  comes  the  small  neotropical,  third  the 
large  Chilean  group:  "hier  liegt  also  wohl  zum  grossen  Teil  spate,  iiberseeische 
Einwanderung  vor".  This  was,  he  found,  even  more  true  of  the  subantarctic 
element,  "das  iiberhaupt  keine  endemische  Arten  aufzuweisen  hat" — very  few 
were  known  then,  but  later  some  endemic  species  were  discovered. 


The  question  to  which  extent  Juan  Fernandez  presents  the  biological  pecu- 
liarities regarded  as  characteristic  of  oceanic  islands — see  Chapter  VII — will  now 
be  answered.  We  have  seen  that  endemism  is  very  high  among  the  phanerogams 
and  that  the  various  kinds  distinguished  by  HoOKER  are  represented;  we  have 
even  a  primitive  endemic  family,  Lactoridaceae,  we  have  a  proportionately  large 
number  of  peculiar  genera,  some  of  them  quite  isolated,  particularly  among  the 
Compositae  {Centaur odendro7i,  Dendroseris,  Hesperosei'is,  Phoenicoseris,  Rea,  Rhe- 
tinodendron^  Robinsonia,  SympJiyochaeta,  Yunqueci)  but  also  in  other  families, 
Cuniiiiia  (Labiatae),  Juania  (Palmae),  Megalacfme  and  Podophorns  (Gramineae); 
further  there  are  a  few  endemic  genera  closely  related  to  South  American  ones, 
Nothomyrcia  (Myrtaceae),  OcJiagavia  (Bromeliaceae),  and  Selkirkia  (Boraginaceae). 
As  among  the  genera,  so  we  find  species  of  a  strong  character  in  Chenopodium, 
Coprosma,  Eryngium,  EupJirasia,  Fagara,  Peperomia,  Plaiitago,  Sanialum,  Urtka, 
Wahleiibergia,  well-marked  but  not  very  aberrant  species  in  Berberis,  Boehmeria, 
Car  ex,  Cladium,  Colletia,  Erjgeron,  Escallonia,  Gunner  a,  Halorrhagis,  Hespero- 
greigia,  Ranunculus,  R/iaphitkaninus,  Solanum  and  Ugni,  and  many  not  very 
different  from  their  continental  congeners  in  Abrotanella,  Acaena,  Apium,  Azara, 
Cardami7te,  Cliusquea,  Drimys,  Dysopsis,  Galium,  Luzula,  Margyricarpus,  Myrc- 
eugenia,  Perneitya,  Pkrygilanthus,  Sopkora,  Spergularia  and  Uncinia.  Species 
undoubtedly  native  but  also  found  elsewhere  do  not  number  more  than  46, 
and  in  several  cases  their  citizenship  is  open  to  question.  Endemism  among 
the  ferns  is  not  so  high,  but  there  is  one  very  aberrant  genus  [Thyrsopteris]  and 
several  peculiar  species.  Bryophytes  and  lichens  will  not  be  considered;  they  were 
not  included  in  HoOKER's  paper  and  I  have  not  had  occasion  to  compare 
them  with  other  island  floras. 

There  are  no  conifers,  a  single  leguminous  genus  [Sopkora]  with  two  species, 
and  no  orchids.  The  proportion  genus  :  species  is  i  :  1.65.  Mammals,  batrachians, 
reptiles  and  fresh-water  fishes  are  absent.  The  earth-worms  are  supposed  to  be 
adventitious  with  the  possible  exception  of  Kerria  saltensis.  In  all  these  respects 
Juan  Fernandez  agrees  with  the  character  attributed  to  oceanic  islands. 

The  high  percentage  of  woody  plants  was  emphasized  by  SiNNOTT  and 
Bailey,  but  as  the  literature  on  which  they  based  their  figures  is  quite  out  of 
date,  a  new  table  was  prepared. 

The  object  of  Sinnott  and  Bailey  was,  as  we  have  seen  above,  to  show 
that  the  woody  plants  increase  in  number  with  the  rising  degree  of  endemism, 
and  that  the  endemic  genera  were  trees  or  shrubs  and  formed  a  more  ancient 
element    than    the    herbaceous    plants    which    during    former  epochs  were  few  in 


378 


C.   SKOTTSBERG 

Table  VL 
Percentage  of  \v()ocl\-  and  licrbaceous  species  in  Juan  Fernandez. 


Indi.utMious  species     .... 

Not  eiitleniic       

ICndeinie,  ^enus  not  eiuleniic  .     .         71 
Als;)  tlie  <'enus  endemic    .     . 


Total 

\N'()()( 

ly% 

Herbaceous  % 

147 

68 

46.3 

79 

1 
53-7 

46 

5 

10.9 

41* 

89.1 

71 

37 

52.1 

34* 

47-9 

30 

27 

90.0 

3 

10. 0 

Four  species  suffruticose. 


comparison,  and  the  Juan  hVrnandez  Islands  were  regarded  as  supporting  their 
h>-|)othesis;  this,  to  judge  from  tlie  table,  they  certainly  do.  The  five  woody 
non-endemic  species  are  liiiipctnDu  nibnun  and  Saliconiia  fruticosa  (low,  erect 
shrubs),  and  Myrtcola  unnimularia,  Riibiis  gcoidcs  and  Calystegia  tuguriorimi 
(trailing). 

The  rosette  tree  form  is  observed  in  16  genera  with  together  31  species, 
belonging  to  si.x  families,  Boraginaceae,  Bromeliaceae,  Chenopodiaceae,  Compositae, 
Plantaginaceae  and  Umbelliferae;  the  palm  Juania  is  of  course  excluded.  As  I 
have  paid  special  attention  to  them  in  another  paper  (-^J/),  where  they  were 
well   illustrated,   no  more  will  be  said  here. 

Annual  and  biennial  herbs,  the  therophytes  of  Raunklkr,  are  not  completely 
lacking,  but  they  are  ver\'  few:  Cardamiiie  chenopodiifolia,  C/iae/otropis  (2  species), 
Par'utaria,  Plauiaoo  tyuncata,  I'etrao^oma  and  Urtica  Masafuerae.  In  2^1  I  listed 
Cliartntropis  among  the  hemicryj)tophytes;  they  give  the  impression  of  lasting 
more  than  one  year  antl  are  found  green  at  all  seasons,  and  the  same  may  be 
true  of  Parictaria.  I  am  not  at  all  sure  that  PUxutago  iruncala  is  native.  The 
Caydanu}ic  has  been  seen  twice,  last  time  in  1872,  the  Crtica  not  since  1854, 
when  it  was  discoxercd.  It  may  be  that  it  is  an  ephemerous  plant  and  disappears 
in  early  spring,  a  season   when   very   few   botanists  have   visited  the  islands. 

Thus,  the  "oceanic  peculiarities"  are  all  there,  but  possibly  some  of  them 
can  be  exj)lained  otherwise.  iMulemism  of  a  very  high  degree  within  a  small 
area  is  no  monopoly  of  isolated  islands;  it  will  be  sufficient  to  mention  the  Cape 
fiorii  or  southwestern  Australia.  The  large  proj)ortion  of  woody  plants  can  be 
understood  if  the  islands  became  isolated  before  the  myriads  of  herbs,  particularly 
the  annuals  of  Central  Chile,  had  evolved.  (iRlSKr,.\(ll's  "Clavija"  and  related 
life  forms  are  not  confuied  to  oceanic  islands;  if  my  interpretation  of  this  mor- 
phological t\'pe  as  an  evidence  of  anti(juity  is  correct,  we  can  understand  why 
it  takes  such  a  pronuncnt  part  in  old  island  fioras.  The  reason  why  therophytes 
are  almost  wanting  is  not  climatic.  This  is  amply  |)roved  by  the  innumerable 
anniuil  weeds  introduced  with  the  traffic  and  thriving  only  too  well.  The  climate 
is  of  a  modified  Mediterranean  tyj)e,  and  from  a  purely  climatic  viewpoint  we 
should  ex{)ect  a  large  percentage  of  native  annuals  and  biennials;  this  question 
was    discussed    at  some  length   in   ^-r/.  827-830.    Chile  has  hundreds  of  endemic 


DERIVATION  OF  THE  FLORA  AND  FAUNA  379 

therophytes,  but  they  have  not  spread  to  the  islands  although  many  have  special 
dispersal  mechanisms,  and  they  were  not  available  at  the  time  when  the  supposed 
connection  with  the  mainland  existed  before  the  final  uplift  of  the  Andes. 

The  existence  of  many  aberrant  genera  and  species  and,  above  all,  the 
marked  difference  between  the  phanerogams  of  Masatierra  and  Masafuera  shows 
that  there  is  no  exchange  between  the  two  islands  in  spite  of  the  very  moderate 
distance;  even  the  360  miles  separating  Masatierra  from  Chile  should,  in  the 
eyes  of  the  diffusionists,  amount  to  little  if  sufficient  time  be  granted.  The  Mar- 
quesas Islands,  situated  much  farther  away  from  the  continent,  have  a  less  peculiar 
angiospermous  flora  than  Juan  Fernandez;  the  very  opposite  ought  to  be  expected 
if  overseas  migration  had  played  the  dominant  role.  The  absence  of  the  flora 
of  Central  Chile  speaks  against  the  efficiency  of  the  natural  dispersal  agents. 
These  circumstances  are  in  favour  of  the  opinion  that  the  volcanic  islands  arose, 
not  from  the  depths  of  the  ocean,  but  on  a  piece  of  land  formerly  connected 
with  South  America  and  not  sunk  until  the  newborn  islands,  now  reduced  to 
ruins,  had  become  a  refuge  for  the  ancient  continental  fauna  and  flora. 

It  is  also  true  that  several  large  and  widespread  families  are  lacking,  such 
as  leguminous  plants  (with  one  exception)  and  lilies,  well  developed  in  Chile, 
but  they  belong  to  the  modern  Chilean  flora. 

The  plant  world  of  oceanic  islands  is  described  as  a  haphazard  collection 
of  waifs  and  strays,  and  this  is  said  to  explain  why  so  few  genera  contain  more 
than  a  couple  of  species.  But  would  not  the  result  be  the  same  if  the  actual 
islands  originated  through  volcanic  activity  on  a  sinking  land.?  Chance  would 
decide  what  took  possession  of  the  new  soil,  and  different  sets  find  a  refuge 
on  Masatierra  and  Masafuera. 

We  know  that  of  the  mammals  introduced  by  man  the  goat  thrives  and 
multiplies  since  400  years  and  quickly  became  naturalized.  There  were  no  goats 
on  the  mainland  when  the  land-bridge  existed,  but  there  may  of  course  have 
been  some  primitive  mammals;  if  any  of  them  reached  the  islands,  they  have 
disappeared  long  ago.  The  islands,  as  we  see  them,  appear  never  to  have  offered 
great  possibilities  for  the  subsistence  of  a  mammalian  fauna.  They  are  very  small 
and  as  there  was  litde  open  land  there  cannot  have  been  any  grazing  grounds 
worth  mentioning  before  man  altered  the  landscape.  Even  after  the  ground  has 
been  cleared  on  all  the  lower  slopes,  pasture  is  miserable,  and  one  valley  after  the 
other  has  been  turned  into  a  desert  by  the  ravages  of  sheep  and  cattle.  If  left 
to  run  wild  and  multiply,  the  final  result  can  be  foreseen.  Carnivorous  animals 
need  a  prey,  and  there  was  none.  The  same  is  true  of  snakes  which  should 
thrive  well  now  since  the  domestic  rats  and  mice  have  been  introduced. 

Chile's  mammalian  fauna  is  poor  and  nobody  knows  if  the  huemul,  the  pudu, 
the  Chilean  rodents  and  small  marsupials  would  be  able  to  make  a  living  on 
Juan  Fernandez.  Reptiles  are  poorly  represented  in  Chile,  there  is  not  a  single 
tortoise,  very  few  snakes  and  a  dozen  lizards.  Amphibians  are  few,  but  toads 
and  frogs  occur.  Among  the  invertebrates  are  several  orders,  the  chances  of  which 
successfufly  to  get  transported  across  the  sea  are  very  doubtful  or,  as  far  as  we 
can  see,  none  at  all. 


38o 


C.   SKOTTSBERG 


The  insular  peculiarities  as  displayed  in  Juan  Fernandez  (and  in  other  similar 
cases)  do  not,  I  think,  permit  us  to  take  a  definite  position  against  the  hypo- 
thesis of  former  land  connections. 


Chapter  X 
The  Chilean  coast  line  and  the  history  of  the  Andes. 

I^^rom  a  look  at  the  map  we  easily  get  the  impression  that  the  trend  of  the 
South  American  west  coast  is  a  j)roduct  of  the  rise  of  the  Andes,  because  this 
enormous  uplift  must  have  been  compensated  by  the  submergence  of  old  border 
lands  and  by  the  formation  of  a  deep  trench,  and  that  these  movements,  which 
certainly  were  of  very  great  magnitude,  may  have  extended  its  effects  west  as 
far  as  to  the  region  where  we  find  the  Juan  Fernandez  and  Desventuradas  Islands 
forming  the  ex{)osed  summits  of  a  submarine  ridge.  I  shall  call  this  ridge  the 
Chaigneau    Ridge  after  the  Chilean   navy  officer  who  was  the  first  to  survey  it. 

flic  Cliaigticau  Ridge. — The  two  archipelagoes  lie  within  the  2000  m  line, 
San  I'elix  and  San  Ambrosio  on  a  plateau  rising  above  the  400  m  curve  and 
extending  a  long  way  toward  Juan  Fernandez,  as  seen  from  CllAlGNEAU's  table 
( -T^)  which  is  reproduced  here  with  the  soundings  rearranged  from  N.  to  S.  accord- 
ing to  latitude  and  with  the  addition  of  some  figures  from  the  latest  chart. 

The  Meniivn  Ridge. — 160  miles  NW.  of  San  Felix-San  Ambrosio  another 
ridge,  called  the  Mcrriam  Ridge,  was  discovered  during  the  U.S.  "Carnegie" 
campaign  1928-29  (//).  It  extends  between  25°  3'. 2  S.,  82°  20'  W.,  and  24°  54'  S., 
"^i"  13'  \\'.;  the  depths  found  were  1445  and  1260  m,  respectively.  The  bank 
rises  3000  m  above  the  bottom  and  is  only  10  miles  wide.  Along  the  most 
elevated  part  i  1 86,  1188  and  1168  m  were  found.  From  the  latter  spot  a  series 
of  soundings  was  taken  SIv  of  the  ridge,  showing  the  rapid  increase  of  the  depth: 
3  miles  1260  m,  9  miles  2751  m,  20  miles  3620  m,  and  32  miles  4115  m;the 
Merriam  ridge  is  separated  from  the  Chaigneau  ridge  by  deep  water.  Toward 
W'XW.   the  sloj)e  is  more  gradual  until  a  depth  of  3000  m  is  reached. 

North  of  the  Merriam  ridge,  in  21^40'  S.,  8i°4o'  W.,  approximately,  a  sudden 
rise,  bounded  by  the  2000  m  curve  and  surrounded  by  deep  water,  has  been 
discovered  (see  map).  Mere  the  bottom  rises  to  972  m  below  the  surface.  I  do 
not  know  if  this  remarkable  j)lace  has  a  name. 

The  Caniegie  Kidfi^e. — During  the  cruise  of  the  "Carnegie"  two  soundings 
about  100  miles  (jff  the  coast  of  Fcuador,  lat.  l°32'  S.,  long.  82°  16',  gave  I  5 1  5 
and  1454  m,  respective!)-,  indicating  a  rise  of  1800  m  above  the  bottom,  but 
before  the  entire  distance  along  the  coast  has  been  surveyed  we  do  not  know  if 
a  series  of  ridges,  jirobably  much  less  well-marked  than  the  Chaigneau-Merriam 
ridge,  can  be  traced  all   the  way   between   lat.    35°   and   the  equator. 

Between  these  ridges  and  the  coast  is  the  deep  trench  (greatest  depth 
7635  m),  and  south  of  the  latitude  of  Juan  Fernandez  depths  exceeding  5000  m 
are  still  found,  but  farther  south  the  4000  m  curve  is  soon  reached  and  the  trench 
disappears.    Older    maps,    two    of   which    were    reproduced    in    22y.  44,  45,    show 


DERIVATION  OF  THE  FLORA  AND  FAUNA 


381 


Table  VII . 

The  Chaigneau  Ridge. 
r  =  rock,  s  =  sand. 


Depth  in 
m 

Bottom 

S.  lat. 

W.  long. 

Remarks 

38 

. — . 

26°  14' 

79°  56' 

14  km  NE.  San  Felix 

80 

— 

26°  15' 

80°    4'.5 

4.5  km  N.  San  Felix 

69 

— 

26°  i7'.5 

80°    8'.5 

4  km  W.  San  Felix 

179 

— 

26°  i8'.5 

80°    8' 

5  km  W.  San  Felix 

220 

r 

26°  i8'.5 

79°  57' 

Between  San  Felix  and  San  Ambrosio 

400 

s 

26°  i8'.5 

79°  49'.3 

5  km  NE.  San  Ambrosio 

250 

s 

26°  19' 

79°  52' 

3  km  N.  San  Ambrosio 

150 

s 

26°  19' 

79°  51' 

3  km  N.  San  Ambrosio 

215 

s 

26°  19' 

79°  49'.2 

4.5  km  NE.  San  Ambrosio 

182 

— 

26°  25' 

79°  56'.5 

6  km  S.  San  Ambrosio 

148 

— 

26°  26' 

79°  52' 

8  km  S.  San  Ambrosio 

105 

s 

26°  30' 

79°  49' 

15.5  km  S.  San  Ambrosio 

400 

26°  36'.5 

80°    8'.5 

33  km  S.  San  Felix 

550 

27°  49'.5 

80°  14' 

c.  165  km  S.  San  Felix 

675 

28°    3'.5 

80°  16' 

c.  200  km  S.  San  Felix 

660 

28°33'.5 

80°  11' 

c.  260  km  S.  San  Felix 

660 

29°  14' 

80°  15' 

c.  340  km  S.  San  Felix 

1300 

30°  49' 

80°  24'.3 

1430 

31°  26' 

80°  10' 

1800 

— 

32°  45'.5 

80°  18' 

Between  Masatierra  and  Masafuera,  about  i 

°  N. 

1800 

— 

34°  34' 

80°  36' 

About  100  km  SSE.  Masafuera 

Juan  Fernandez  situated  on  a  "lobe"  with  somewhat  shallower  water,  less  than 
3000  m  deep,  extending  NW.  from  the  coast  and  suggesting  a  possible  former 
connection,  but  recent  charts  are  less  unambiguous.  The  2000  m  curve  makes, 
however,  a  bulge  around  lat.  38°,  where  a  depth  of  only  1238  m  is  indicated 
while  deeper  water,  1400-1500  m,  is  met  with  near  the  coast.  Soundings  be- 
tween the  islands  and  the  line  where  the  trough  stops  are  too  few  to  be  of  much 
value,  but  in  all  probability  the  connection,  if  it  did  exist,  should  be  looked 
for  farther  south.  This  is,  as  we  shall  see,  also  the  opinion  of  the  geologists. 
They  do  not  hesitate  to  regard  the  banks  just  described  as  an  extension  from 
the  continent.  Nobody  will,  I  suppose,  argue  that  every  marked  rise  of  the  ocean 
floor  is  a  sign  of  sunken  land;  the  majority  of  oceanographers  prefer  to  call 
them  independent  products  of  volcanic  action.  Some,  perhaps  most  of  them, 
never  reached  the  surface,  some  have  done  so,  but  were  broken  down,  but  many 
are  still  above  the  water  and  form  the  Pacific  islands.  But  even  the  advocates 
of  the  permanence  of  this  largest  of  basins  admit  that  its  margins  are  zones  of 
considerable  disturbance.  If  it  can  be  proved,  or  at  least  be  made  probable,  that 


382  <-"•   SKOTTSBERG 

the  deep  trench  following  the  trend  of  the  Andes,  is  a  consequence  of  the  gigan- 
tic mountain-builchng  processes,  tlie  submarine  ridges  west  of  the  trench  can- 
not hel|>  to  get  iniphcated.  Tiiis  idea  is  by  no  means  new,  it  has  been  expressed 
by  many:  "the  width  of  Soutli  America  may  well  be  a  good  deal  less  now  than 
before  the  Andes  were  uplifted",  as  G()(JI)  says  (lop.  349),  but  some  of  these 
writers  did  not  tr\'  to  penetrate  the  complicated  geographical-geological  history 
of  til  is  region  of  great  tectonic  disturbance.  This  is  true  of  myself,  when  I  tried 
to  describe  what  I  imagined  having  occurred  (22/.  43),  but  it  does  not  apply  to 
Ikms(IIi:k  (//i')  who  took  pains  to  inform  himself  of  the  history  of  the  Andes 
as  told  b\'  geologists.  That  they  arose  in  a  geosyncline  is  proved  by  the  Jurassic 
and  Cretaceous  beds  now  elevated  thousands  of  meters  and  covering  the  older 
eruptives.  To  the  east  of  the  depression  land  had  existed  since  the  Permian,  to 
the  west  was  a  Tacihc  land  mass  of  hypothetical  width;  one  opinion  regarded 
the  Coast  Range  as  belonging  to  this  land.  Irmscher,  who  took  \\T:c;ener's 
sitle.  did  not  ask  for  any  large-scale  subsidence  correlated  with  the  uplift  of  the 
Andes,  because  the  resistance  of  the  sialic  crust  to  the  westward  drift  of  South 
America  was  sufticient  to  account  for  uplift  and  folding.  Consequently,  he  was 
unwilling  to  accept  Pexck's  intrusion  theory;  if,  in  the  future,  it  should  appear 
essential  to  accept  a  land  mass,  it  could  be  nothing  more  than  a  narrow  strip 
which,  perha{)s,  had  been  connected  with   California  (p.  45). 

l'i.N(  K  stellte  (lie  Aiisbildiint(  dor  ozeanischen  Tiefen  am  Rande  des  sudamerika- 
nischen  Kontincntcs,  also  das  \'ersinken  angrenzender  'j'eile  des  Pazifiks,  der  aufwarts 
bewegten  andinen  Scholle  gegeniiber  und  schliesst,  dass  die  niichstliegende  Erklarung 
liir  die  \'olumenanderungen  unter  der  festen  Kruste  in  dort  stattfindenden  Massenver- 
schicbungen  zii  suchcn  ist.  Wenn  Massen  aus  der  pazifischen  Region  in  die  andine 
iihertrcten,   so   nuiss  in   crstcrer  die   Kruste  nachsinken   (p.    51). 

The  cause  of  the  uplift  and  folding  was,  according  to  Penck,  a  result  of 
the  intrusion  of  the  andesitic  magma,  which  lifted  the  mountains  but,  Irmscher 
remarks,  this  could  not  be  the  only  source  of  the  tangential  pressure: 

i)ie  ine(  hanis(  hen  Ursachcn  des  l-altenvorganges  sind  zweifellos  anderer  Natur,  und 
(lie  .Magniaintnision  ist  genau  so  eine  Uirkung  derselben  wie  die  P'allung.  Denn  es  steht 
test,  dass  die  ( ichirgshildung  niit  dem  Kmpordringen  des  Magmas  synchron  ist  und  die 
Intrusion  somit  gleic  hzcitig  niit  der  angcnommenen  annuihlichen  Lostrennung  Siidamerikas 
von  .Atrika. 

Where  does  the  (■iiaigneau-Merriam  bank  come  in.-  \Ve(;ener  left  it  unexplained, 
according  to   1)1    Tori    it  was  an   "advance  foltl". 

l^EKRV  and  SiN(;i:w.\i,i)  (j^),  in  their  review  of  the  tectonic  history  of  South 
America,  describe  the  develoj)ment  in   the   following  terms. 

Some  students  regard  the  Cordillera  de  la  Costa  in  Chile  as  remnants  of  an  an- 
(  ient  massif,  the  bulk  of  \\hi(  h  has  been  downfaultcd  beneath  the  waters  of  the  Pa- 
cific. It  (onsists  of  (rystalline  ro(  ks  both  igneous  and  metamorphic  and  these  have 
(onimonly  been  assumed  to  be  of  great  age — even  Archean.  This  inference  of  antiquity 
rests,  not  upon  their  known  rehitions,  but  upon  the  geosynclinal  nature  of  the  Andean 
seas   which   are  clearly  ej)icontinental   and   not  shelf  seas. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  383 

The  sediments  in  this  geosynchne  are  compressed  between  the  Brazihan 
massif  and  another  massif  in  the  west,  the  vestiges  of  which  should  be  looked 
for  in  the  coast  range.  Anyhow,  "that  there  was  land  to  the  west  of  the  Western 
Andes  cannot  be  doubted".  These  authors  date  the  Concepcion-Arauco  series 
to  the  older  Miocene  and  they  do  not. regard  the  flora  as  a  coast  flora. 

Let  us  now  turn  to  Bruggen,  author  of  a  modern  handbook  on  the  geology 
of  Chile  (jji).  To  begin  with  I  shall  allow  myself  to  quote  Florin's  summary 
(^5.  4-6)  of  Bruggen's  earlier  writings. 

Until  Middle  Tertiary  times  the  great  Andes  and  the  Coastal  Range  were  ...  a 
continuous  upfolded  mountain  chain  subjected  to  powerful  denudation.  .  .  .  The  principal 
uplift  of  the  Andes  in  Chile  occurred  in  the  Middle  Cretaceous,  and  altered  the 
palaeogeographical  features  of  the  Andean  region  considerably.  The  old  geosynclinal  had 
been  turned  into  a  continental  area,  at  the  western  verge  of  which  the  border  of 
the  Pacific  Ocean  at  the  end  of  the  Cretaceous  occupied  approximately  the  same  line 
as  to-day.  Marine  deposits  of  Danian  ...  as  well  as  Palaeocene  age  are  lacking,  and 
at  the  beginning  of  the  Tertiary  period  the  continent  probably  extended  further  to 
the  west. 

In  the  Eocene  and  Oligocene,  respectively,  subsidences  took  place  and  the  ocean 
encroached  more  and  more  on  the  land.  The  Concepci6n-Arauco  coal  measures 
are  coastal  deposits,  which  have  been  called  the  Concepcion  Series  by  Bruggen. 
This  series,  about  400  m.  thick,  rests  unconformably  on  marine  strata  of  Upper 
Cretaceous  (Senonian)  age.  The  shales  containing  fossil  plants  occur  in  conjunction 
with  intercalated  coal  seems.  .  .  .  The  base  of  this  section  is  of  marine  origin,  and 
in  addition  marine  layers  are  intercalated  here  and  there  in  its  middle  part,  which 
is  otherwise  generally  built  up  of  freshwater  deposits.  According  to  Bruggen  the 
Concepci6n  Series  is  overlain  by  the  deposits  of  the  marine  Navidad  Series,  which 
is  upper  Oligocene  or  Lower  Miocene  in  age.  Bruggen  came  to  the  conclusion  that 
the  Concepci(3n  Series  belongs  to  the  Eocene,  basing  this  on  stratigraphical  as  well 
as  on  zoo-palaeontological  evidence.  The  sediments  of  this  series  were  according  to 
him  deposited  on  a  broad,  slowly  sinking  coastal  plain,  and  subsequently  subjected 
to  considerable  tilting  and  faulting,  probably  in  the  Miocene. 

Berry  regarded  both  the  Concepcion  and  the  Navidad  series  as  belonging  to 
the  Lower  Miocene  or  possibly  Upper  Oligocene,  but  according  to  Bruggen 
this  dating  holds  good  for  the  latter  only,  while  the  former  is  much  older,  and 
whereas  BrCggen  thinks  that  the  Eocene  coal  flora  was  deposited  in  extensive 
coastal  swamps,  Berry  regarded  it  as  neither  limnic  nor  littoral,  but  inhabiting 
a  lowland  area  away  from  the  coast.  FLORIN  found  that  the  plant  remains  were 
laid  down  in  the  vicinity  of  the  sea  and  that  they  are  too  well  preserved  to 
have  been  transported  any  great  distance. 

If  it  is  true  that  the  coast-line,  at  the  end  of  the  Cretaceous,  occupied  the 
same  position  as  to-day,  one  is  inclined  to  believe  that  the  palaeogeography 
was  different  before  the  great  uplift  occurred  in  the  Middle  Cretaceous,  when  a 
large  scale  subsidence  ought  to  have  taken  place.  The  oscillations  along  the 
coast  of  central  Chile  during  later  times  could  hardly  have  involved  the  area 
where  the  submarine  ridges  are  found,  so  that  the  possibility  to  link  them  to 
that  part  of  the  continent  as  late  as  that  is  small.  Consequently  attention  has 
been  directed  farther  south,  as  already  suggested.  West  Patagonia  is  a  region 
of  considerable    and    late    subsidence;    the    longitudinal    valley    of  central    Chile 


284  C.   SKOTTSBERG 

disappeared  under  water  to  form  the  long  series  of  the  Patagonian  channels,  the 
Andean  valle\-s  became  fiords  and  the  broad  dissected  fringe  of  islands  and 
skerries  also  give  evidence  to  what  has  happened.  The  weight  of  the  inland  ice 
during  the  perioils  of  glaciation  must,   however,   also  be  taken  into  account. 

The  development  of  the  coastal  region,  as  told  by  BrCggen  1950,  is  ex- 
})lained  by  facts  which,  if  they  have  been  correctly  interpreted,  open  wide  per- 
spectives   to  the  biologist,  even   if  serious  difficulties  still  have  to  be  overcome. 

HkiciCKN  begins  by  stating  that  "el  mar  del  Eoceno"  ended  somewhere 
in  the  latitude  of  Arauco  (38°),  because  a  continental  mass,  "la  Tierra  de  Juan 
Fernandez"  still  existed  (p.  50),  and  pp.  56-59  he  relates  the  history  of  this  land. 
North  of  Rio  Maulh'n  (about  42°)  is  a  zone  of  dislocations  foreign  to  the  struc- 
ture of  the  Andes,  and  this  zone  coincides  with  the  direction  of  a  broad  sub- 
marine ridge  which  branches  off  from  the  continent;  on  this  ridge  are  situated 
the  Juan  PY^nandez  Islands  and,  farther  north,  San  Ambrosio  and  San  Felix. 
Taking  the  2000  m  cur\e  as  a  boundary,  the  ridge  extends  south  to  the  Magellan 
Straits;  we  observe  e.g.  in  the  island  Diego  de  Almagro  the  same  northwest 
direction  that  we  tind  in  the  Tertiary  deposits  of  Parga  and  other  places  in  the 
zone  north  of  Rio  Maulh'n.  To  this  must  be  remarked  that  the  2000  m  line  sur- 
rounds the  Chaigneau  ridge  and  that  in  order  to  unite  it  with  West  Patagonia 
the  3000  m  curve  has  to  be  used.  This  is  also  seen  from  I^rCggen's  map,  prob- 
ably copietl  from  Si  pax.  The  absence  of  marine  sediments  of  Eocene  age 
shows  (p.  59)  that  the  Juan  Fernandez  land  was,  at  that  time,  united  with  the 
continent,  but  that,  during  the  Oligocene,  subsidence  set  in  is  evident  from  the 
extension  of  the  marine  Xavidad  series  south  to  45°,  and  this  was,  as  we  have 
heard,  referred  to  Upper  Oligocene  or  Power  Miocene.  Also  after  the  separation 
the  Juan  Fernandez  land  continued  to  exist  until  finally,  presumably  with  the  late 
Tertiary  uplift  of  the  yVikIcs,  the  last  rest  disappeared,  but  not  before  considerable 
magma  ejections  had  given  birth  to  the  two  archipelagoes.  To  judge  from  the 
degree  of  denudation  and  in  view  of  the  recent  volcanic  activity  close  to  Masa- 
tierra  and  on   San   h'elix^  the  islands  are  young,  probably  Pliocene, 

(  iiando  cxistt'a  todavia  im  resto  de  la  antigua  Tierra  de  Juan  Fernandez,  de  la  cual 
inm/xro  la  flora  del  lioccno?  Ciiando  mas  tarde  se  hundi6  tambien  este  resto,  sobre- 
salian  solanicntc  las  i)artc's  volcanicas,  constituyendo  las  islas  actuales  de  Juan  Fernandez, 
(lue   Servian    j)ara   rcfugif)   de   la   flora. 

This  is  the  process  as  I  have  described  it  (.^27.  43)  and  BrCgcjKN  is  of  the 
same  oj)inion  from  the  geologist's  viewpoint.  And  if  we  go  back  to  Hooker's 
lecture,  we  shall  [wmX  that  the  same  idea,  ai)i)lied  to  a  different  region,  was  famil- 
iar with   him. 

1  This  refers  to  the  siil,>inarinc  eruption  in  1835  "^<ir  die  coast  of  Masatierra;  whether 
any  signs  are  left  I  cannot  tell,  for  tlie  place  has  not  been  sounded.  This  eruption  was  simulta- 
neous with  an  (Mrthi|uake  in  ("oik  epcion.  I'ossilily  there  was  a  connection  with  the  tsunami  of 
\'allenar  in  1922  and  the  eruption  at  .San  Felix  three  months  later,  when  gas  was  ejected  on 
the  island,   killing   a  great  number  of  sea-birds. 

2  My  italics. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  385 

With  regard  to  the  objection  that  oceanic  islands  are  volcanic,  and  hence  prob- 
ably not  the  mountain-tops  of  sunk  continents  and  that  they  contain  no  fossil  mam- 
mals, we  have  in  the  Malay  Archipelago,  vast  areas  of  land  which  if  submerged  (and 
they  are  exposed  to  constant  subsidences  and  risings)  would  leave  only  isolated  vol- 
canic peaks,  such  as  oceanic  islands  present.  Were  such  an  area  to  be  submerged, 
leaving  exposed  the  volcanic  peaks  of  Java  and  the  Moluccas  &c,  &c,  should  we  expect 
to  find  either  recent  or  fossil  terrestrial  mammals  upon  them?  Nor  should  it  be  over- 
looked that,  as  a  general  rule,  islands  diminish  in  size  and  numbers  toward  the 
centres  of  the  great  oceans,  which,  taken  with  the  admission,  that  the  great  islands 
adjacent  to  the  continents  were  previously  united  to  them,  would  favour  the  hypo- 
thesis that  all  may  have  been  so.  And  finally,  we  have  instances  of  continental  distri- 
bution, presenting  facts  so  analogous  to  oceanic,  and  hitherto  so  utterly  unexplicable, 
on  any  hypothesis  of  migration  that  does  not  embrace  immense  geological  changes, 
that  we  can  scarcely  avoid  coupling  the  phenomena  they  present  with  those  of  oceanic 
islands  (p.    1 1). 

However,  HoOKER  did  not  stop  here.  When  he  threw  the  pros  and  cons 
into  the  balance,  the  scale  of  scruples  sank: 

On  the  other  hand,  to  my  mind,  the  great  objection  to  the  continental  extension 
hypothesis  is,  that  it  may  be  said  to  account  for  everything,  but  to  explain  nothing; 
it  proves  too  much,  whilst  the  hypothesis  of  trans-oceanic  migration,  though  it  leaves 
a  multitude  of  facts  unexplained,  offers  a  rational  solution  of  many  of  the  most  puzz- 
ling phenomena  that  oceanic  islands  present:  phenomena  which,  under  the  hypothesis 
of  intermediate  continents,  are  barren  facts,  literally  of  no  scientific  interest — are 
curiosities  of  science,   no  doubt,   but  are  not  scientific  curiosities. 

This  was,  I  am  afraid,  to  say  too  much — but  are  not  Hooker's  words  a 
good  expression  of  the  biogeographer's  dilemma.? 

The  latest  brief  outline  of  the  history  of  the  Andes  is  found  in  Goodspeed's 
monograph  of  Xicotiaiia  [ii2)\  it  was,  partly  at  least,  based  on  WEEKS'  paper 
{2jg).  The  sequence  of  events  especially  refers  to  the  central  Andes,  with  which 
the  history  of  the  southern  Andes  is  said  to  agree.  Four  major  periods  of  uplift 
are  recognized,  the  final  one  during  the  Pliocene-Pleistocene,  with  an  average 
elevation  of  3500  m.  Coincident  with  the  uplift  was  a  compensory  coastal  sink- 
ing. The  troughing  along  the  coast  ceased  with  the  foundering  of  the  Pacific 
fore-lands  at  the  close  of  the  Miocene  and  early  Pliocene.  These  lands  are  sup- 
posed to  have  extended  an  unknown  distance  westward  (p.  31).  It  is  possible 
that  the  border-lands  extended  all  the  way  north  along  Central  America,  Mexico 
and  California,  where  are  found  islands  in  a  position  corresponding  to  that  of 
the  Galapagos  and  Juan  Fernandez  Islands. 

From  what  has  been  said  above  I  can  only  find  that  there  is  good  geo- 
logical evidence  for  a  former  extension  west  of  the  continent,  uniting  southern 
Chile  with  a  "Juan  Fernandez  land".  Before  attempting  to  trace  the  sequence 
of  immigration  of  the  various  floristic  elements  and  of  their  fate  we  ought  to 
know  something  of  the  fossil  floras  of  Chile  and,  further,  to  devote  a  chapter 
to  the  supposed  Antarctic  migration  routes. 


25  ~  557857    The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.    Vol.  I 


^Sb  <-•   SKOTTSBERG 

Chapter  XI. 

The  Tertiary  floras  of  Chile  and  Patagonia. 

'rcrtiar\'  plant  fossils  have  been  discovered  in  the  coast  region  of  south  Chile, 
in  Patagonia  along  the  Andes  south  to  the  Magellan  Straits,  and  in  Tierra  del 
Fuego.  The  richest  localities  are  on  the  coast  of  the  province  of  Arauco,  37°-37°30' 
s.  hit.,  in  combination  with  the  coal  seams  of  Lota,  Lebu  etc.,  and  east  of  Lake 
Xahuclluiapi   on   the  Argentine  side. 

I'lu-  C  'o)iLt'pcio)i-Ayauco  series. — The  fossil  flora  was  examined  by  ILngkliiardt, 
who  described  more  than  one  hundred  species  of  angiosperms,  and  later  by  BERRY 
\2j].  The  determinations  are,  just  as  in  all  the  other  cases,  based  on  leaf  impres- 
sions, but  although  l^l'.KKV  found  that  KxoKLllARDT's  determinations  "are  usually 
to  be  relied  upon"  ({x  75),  very  many  of  them  are,  as  well  as  his  own,  open  to 
doubt.  The  species  were  as  a  rule  referred  to  living  genera,  belonging  to  many 
more  or  less  important  tropical-subtropical  families,  Annonaceae,  Apocynaceae, 
Hignoniaceae,  Bombacaceac,  Caesalpiniaceae,  Combretaceae,  Erythroxylaceae,  Lau- 
raceae,  Lec\'thi(.laceae,  Myrtaceae,  Palmae,  Sapindaceae,  Styracaceae,  Vochysiaceae, 
and  so  forth.  In  general  character  the  flora  approaches  that  of  the  Amazon  basin, 
e.xtended,  the  relief  of  the  Andes  being  low  at  that  time,  to  the  west  coast  of 
the  continent  and  reaching  south  at  least  to  40°  s.  lat.  The  flora  contains  "no 
eleiuents  of  the  flora  of  Central  Chile"  (p.  106) — Cassia  is,  however,  one  of  the 
genera  mentioned,  another  is  Myraugcuia,  represented  by  several  species  in  cen- 
tral and  south  Chile  (to  know  Myrtaceae  without  flower  and  fruit  is  well-nigh 
impossible).  Hl'kKV  referred  the  flora  to  the  Lower  Miocene,  "it  is  surely  not  so 
okl  as  lv)cene",  and  also  younger  than  the  XotJiofagus  flora  in  the  Magellanian 
zone  (j).  115);   this  he  considered  to  be  of  Low^er  Oligocene  age. 

There  are  two  gynuiosperm  genera  in  Berrv's  list  which  seem  to  disturb 
the  imj)ression  of  an  otherwise  homogeneous  neotropical  assemblage,  Araucaria 
and  St<//a'i(f.  The  material  was  reexamined  by  FLORIN  (pj)  who  showed  that 
Araiuana  ara!/i(>r>fsis  Berry  is  a  species  of  Podocarpus,  and  Sequoia  cJiilensis 
Fngclhardt  p.  p.  another.  These,  together  with  a  fern  described  by  HallE  (Lygo- 
({n())i,  ;i2]  are  important  additions  to  the  Arauco  flora.  F^LORIN,  in  accordance 
with   l^KrciCl.N,  ref(,'rs   it  to  the  I'>)cene  and  characterizes  it  as  follows. 

The  ( oinpo.sition  of  the  fossil  c:onifer  vegetation,  and  the  distributional  aspects,  of 
it>  <  onstitucnts,  iii(!i(  ;itc  that  it  derives  from  a  \varni-teiuj)crate  or  sul)troj)ical  rain-forest, 
more  particularly  a  lowland  i)od()c.arj)-evcrgreen  dicotylous  broad-leaved  tree  forest, 
growint,'  on  tin-  ( oastal  plain  or  j)erhaps  partly  on  low  hills  not  far  from  the  coast.  The 
(  limatc  was  ])r()hal)ly  (  harac  tcri/.ed  hy  great  humidity  and  rather  uniform  temperature, 
it    was    trostless,    and    wanner  than    the   i)rescnt   climate   of  the   same   district  (p.  26). 

The  possibility  that  plant  material  from  the  uplands  had  been  carried  down 
and  become  nnxcd  with  material  frf)m  the  coastal  plain  is  contradicted  by  the 
state  ol    preservation   which   is  the  same  in   all  cases  (p.  26). 

'I he  rhiiileufu  flora. — The  fossiliferous  beds  of  Rio  Pichileufu  are  situated 
in  41     s.  lat.  about  30  miles  east  of  Lake  .Xahuelhuapi  in  a  treeless  steppe  country. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  387 

Berry  {2^)  distinguished  more  than  130  species,  most  of  them  referred  to  still 
living  genera  and  belonging  to  48  families  and  21  orders.  The  general  character 
is  subtropical,  and  the  following  families  may  be  mentioned:  Anacardiaceae,  An- 
nonaceae,  Apocynaceae,  Asclepiadaceae,  Bignoniaceae,  Burseraceae,  Caesalpinia- 
ceae,  Celastraceae,  Cochlospermaceae,  Erythroxylaceae,  Euphorbiaceae,  Flacourt- 
iaceae,  Icacinaceae,  Lauraceae,  Loganiaceae,  Meliaceae,  Mimosaceae,  Monimiaceae, 
Moraceae,  Myristicaceae,  Myrtaceae,  Nyctaginiaceae,  Rubiaceae,  Rutaceae,  Sapinda- 
ceae,  Sapotaceae,  Sterculiaeae,  Styracaceae,  Symplocaceae,  Vitaceae.  Of  conifers 
we  find  Araucaria  pichileufuensis,  Fiizroya  tertiaria,  Libocedrus  prechilensis  and  2 
Podocarpus\  further,  there  is  a  species  of  Zaniia  and  Ginkgo  patagonica,  and  of 
ferns  3  species,  one  of  them  a  Dicksonia.  Araucaria  and  Libocedrtis  are,  according  to 
Florin,  correctly  named  (pj),  Ginkgo  should  be  called  Ginkgoites\  Fitzroya  belongs 
to    Podocarpus,    and   one  of  the  Podocarpus  sp.  belongs  to  Acmopyle  of  PiLGER. 

There  is  no  trace  of  Fagaceae  and  Berry  referred  the  flora  to  Low^er  Mio- 
cene and  regarded  it  as  contemporaneous  with  the  Arauco  flora;  they  have  20 
species  in  common,  the  general  character  is  the  same  and  is  said  to  bear  witness 
of  the  same  climate.  The  relief  of  the  Andes  was  low,  no  rain-shadow  existed, 
prevailing  westerly  winds  carried  abundant  moisture  across  the  country,  there 
was  rain  forest  where  now  we  have  dry  grass-land.  Still,  there  is  a  difference 
between  Arauco  and  Pichileufu.  Berry  [zf]  pointed  out  that  the  present  South 
Chilean  rain  forest  flora  is  not  represented  in  the  Arauco  flora  whereas  the 
Pichileufu  beds  contain  such  Chilean  genera  as  Azara,  Berberis,  Maytenus  and 
Myrceugenia  and,  in  addition,  the  following  Antarcto-tertiary  genera:  Drimys, 
Embothrium,  Eucrypliia,  Laurelia,  Libocedrus  and  Lontatia — provided  that  the 
determinations  are  correct.  Nevertheless  the  age  is  supposed  to  be  the  same. 
Lower  Miocene  according  to  Berry,  Eocene  according  to  Florin,  thus  older 
than  the  Araucaria-Nothofagus  beds  of  Magallanes.  It  is  surprising  that,  if  the 
two  floras  are  of  exactly  the  same  age,  the  advancing  Antarctic  flora  had  not 
found  its  way  to  the  coast  of  Chile;  Pichileufu  ought  to  be  younger,  but  perhaps 
still  Eocene,  a  period  of  very  great  length. 

The  Chalia  flora. — Of  considerable  interest  was  the  discovery,  in  Santa  Cruz 
Territory  in  the  valley  of  Rio  Chalia  about  51°  s.  lat.,  of  a  fossil  flora  similar 
to  the  Arauco  and  Pichileufu  floras  and  proving  that  the  subtropical  vegetation 
had  extended  far  south.  Araucaria  and  Nothofagus  are  absent,  the  only  conifer 
found,  Fitzroya  tertiaria,  is,  as  shown  by  FLORIN,  a  Podocarpus.  Of  angiosperm 
families  Anacardiaceae,  Annonaceae,  Bignoniaceae,  Lauraceae,  Monimiaceae,  Myr- 
taceae, Sterculiaceae  etc.  are  represented,  of  Chilean  genera  Laurelia  and  Peumus 
may  be  mentioned.  The  age  is  early  Miocene  according  to  BERRY  (jj^).  Eocene 
according  to  Frenguelli  [337),  the  climate  warm  temperate. 

Berry  {313)  regarded  all  the  fossil  floras  containing  an  abundance  of  Faga- 
ceae [Nothofagus,  according  to  DUSEN  also  Fagus,  which  is  questionable)  as  of 
approximately  the  same  age  and  older  than  the  Concepcion-Arauco  series. 

The  Nirihuao  flora. — Three  localities  close  together  on  Nirihuao  river  near 
Lake  Nahuelhuapi.  Some  ferns,  among  them  Alsophila  australis,  also  known  from 
Seymour  Island,  further  Zamia,  Araucaria  Nathorstii,  Fagus  (?)  and  one  species 


388  C.   SKOTTSBERG 

o{  Xo/Ziofcii^iis.  A^e  proposed  b\'  HKkRV  (jJ4)  Lowest  Miocene  or  Upper  Oligocene, 
then   \-oun<^er  than   the  Ma^allancs  flora. 

r/if  Mdi^iil/cvns  fiord. — DusKX  [yQ)  distin^uislied  two  plant-bearing  horizons, 
an  upper  Ayaucafia  horizon  and  a  lower  Xothofagus  horizon.  Of  the  remaining 
dicots'lons  genera  none  were  identified  with  living  ones;  DUSEX  preferred  to  call 
them  liscalliuiiipJirUiDii.  I lydnvioeiphylliDu  etc.  There  is  no  obviously  tropical 
eletnent,  it  is  a  temperate  flora.  DlSKX  regarded  the  two  horizons  as  distinctly 
different  in  age,  dating  the  upper  to  Lower  ^Miocene,  the  lower  to,  perhaps, 
Oligocene.  Hkrrv.  who  doubted  the  correctness  of  this  distinction,  regarded  them 
as  older  than  the  Concepcion-Arauco  flora,  which  seems  improbable. 

riie  Snuioin-  flora. — To  judge  from  DusKX's  description  (cVo)  the  tropical 
element  is  not  conspicuous,  whereas  the  actual  South  Chilean  forest  flora  is  well 
represented:  Araucaria  (nearly  related  to  A.  araiicaua),  Drimys,  Xotfiofagus., 
L'aldcluria,  lAUirelia,  I.oDiatia,  all  supposed  to  be  of  Antarctic  parentage;  lean 
see  little  reason  for  I^KKR\'s  assertion  that  the  Seymour  flora  contains  "a  large 
element  of  subtropical  or  warm  temperate  types  like  those  found  to-day  in  south- 
ern Hrazil"  together  with  "another  large  element  of  forms  suggestive  of  the 
existing  temperate  flora  of  Southern  Chile  and  Patagonia";  the  former  was  sub- 
tropical and  coastal,  the  latter  temj^erate  and  montane,  washed  down,  DuSEN 
thought,  from  the  mountains  and  embedded  together  with  the  leaves  of  the  low- 
land trees.  The  age  was  estimated  to  be  Upper  Eocene.  Florin's  discovery  of  a 
species  of  Aonopyle  [I^liyllites  sj).,  DlSEx)  is  of  particular  interest  [338]. 

We  have  seen  that  Berry  considered  the  Xotfiofagus  beds  to  be  older  than 
the  Arauco-Pichileufu  deposits.  y\ll  the  local  fossil  floras  of  Patagonia  are,  he 
says  [333],  older  than  the  marine  Patagonian  transgression  and  undoubtedly  pre- 
Miocene.  P'rexcueli,!  distinguished  three  epochs:  (a)  Late  Cretaceous  to  early 
lv)cene,  with  a  tropical  flora,  known  from  the  Chali'a  beds;  (b)  an  intermediate 
period  with  subtropical  and  temperate  types  [Xotfiofagus)  mixed;  to  this  he  would, 
1  suppose,  refer  the  Seymour  flora;  (c)  the  youngest  epoch,  Miocene-Pliocene, 
evident!)'  extending  into  Pleistocene:  a  temperate  flora,  now  ranging  along  both 
sides  of  the  southern  Andes  and  characterized  by  the  dominance  of  A^otfiofagus 
and  ot  a  number  of  conifers.  The  more  or  less  corresponding  development  of 
the  Andes  was  according  to  I5ERRV  (2S):  (i)  I'^ocene-early  Miocene:  low  relief, 
no  high  continuous  mountains,  followed  by  (2)  a  period  of  great  uplift;  (3)  late 
Miocene  to  early  Pliocene:  mature  erosion,  low  relief;  (4)  late  Pliocene  to  Pleis- 
tocene: extensive  uplift,  beginning  of  the  formation  of  the  Chilean  and  Peruvian 
deeps,  where  earlier  there  was  land;  (5)  submergence  of  the  coastal  plain.  Finally, 
but  not  mentioned  by  HlRRV,  the  series  of  glacial  and  interglacial  periods,  a 
most   important   factor   of  disturbance. 

'I  his  is  tlie  background  against  which  we  have  to  discuss  the  history  of  the 
Juan    I'ernandez   flora. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  389 

Chapter  XII. 
Antarctica  as  a  source  of  the  present  circumpolar  floras. 

Much  has  been  written  on  this  subject  and  it  is  not  necessary  to  review 
the  entire  hterature,  which  has  been  done  already  by  several  authors,  but  the 
Antarctic  problems  are  so  important  when  it  comes  to  an  analysis  of  Juan  Fernan- 
dez that  they  cannot  be  passed  in  silence.  J.  D.  Hooker  was  the  first  to  survey 
all  the  lands  scattered  around  the  Antarctic,  Tierra  del  Fuego  and  the  Falkland 
Islands,  Kerguelen  Island,  Tasmania,  New  Zealand  and  its  subantarctic  dependen- 
cies; he  was  struck  by  the  discontinuous  distribution  of  many  genera,  families 
or  even  species— Hemsley,  i2y  (a),  and  the  author  [228)  have  given  lists  of 
such  genera  and  species — he  drew  the  consequences,  although  nothing  was  known 
then  about  the  vanished  flora  of  the  large,  ice-covered  continent,  nor  of  the  palaeo- 
geography  of  the  adjacent  zone,  and  the  idea  that  Antarctica  had  formerly  ex- 
tended farther  north  and  that  the  sporadic  southern  islands  eventually  were 
remnants  of  larger  land  masses  entered  his  mind.  Since  that,  the  various  subant- 
arctic and  austral  floras  have  become  very  well  known,  and  the  cases  of  remark- 
able disjunctions  have  multiplied.  To  those  who  adhere  strictly  to  the  hypothesis 
of  long-distance  dispersal  across  the  oceans  this  means  nothing  more  than  further 
proofs  that  they  are  right,  for  the  west-wind  drift  explains  everything.  Fortunately, 
the  land  bridge  between  South  America  and  Antarctica  rests  on  solid  foundation, 
geographical  as  well  as  geological.  In  his  important  paper  of  1929  [137)  HOLTE- 
DAHL  has  shown  that  the  old  idea  of  land  connection  between  Tierra  del  Fuego 
and  Graham  land  (Palmer  peninsula)  by  way  of  the  Burdwood  Bank,  Shag  Rocks, 
South  Georgia  and  the  South  Sandwich  and  South  Orkney  Islands,  which  had 
been  doubted  by  some,  holds  good:  we  have  to  do  with  a  mountain  range,  a 
continuation  of  the  South  American  Andes,  bordered  by  deep  water  which,  on 
the  Pacific  side,  has  the  character  of  an  abysmal  trench  but  which  exhibits  old 
sediments  to  such  an  extent  that  we  are  forced  to  postulate  land  where  there  is 
now  deep  sea.  The  South  Sandwich  Islands,  being  entirely  neovolcanic,  have  the 
appearance  of  an  "oceanic"  archipelago,  but  they  were  built  up  during  late  Ter- 
tiary times  over  an  older  foundation — a  parallel  to  the  history  of  Juan  Fernandez 
and  of  many  other  islands. 

It  is  .  .  .  quite  evident  that  the  South  Shetland  land  mass  has  once  had  several 
times  the  width  that  it  has  to-day.  .  .  .  With  their  large  amount  of  terrigenous,  clastic 
sediments  etc.,  the  South  Orkneys  and  also  South  Georgia  agree  with  the  folded  ranges  of 
the  continent.  In  fact,  in  order  to  explain  these  masses  of  sediments  we  must  necessarily 
assume  land  to  have  been  present  where  there  is  now  deep  sea. 

We  need  not  assume  that  all  the  links  of  the  Scotia  or,  as  it  is  also  called. 
South  Antillean  Arc  were  united  at  the  same  time,  let  it  be  that  this  is  possible 
or  even  probable;  if  so,  there  was  no  communication  by  water  between  the  At- 
lantic and  the  Pacific,  which  undoutedly  must  have  had  its  consequences  to  the 
water  circulation,  a  question  I  am  quite  unfit  to  discuss.  To  judge  from  the  compo- 


390  C.   SKOTTSBERG 

sition  of  the  local  subantarctic  floras,  rich  in  herbaceous  plants  belonging  to  many 
different  families  and  orders,  land  communication  must  have  persisted  to  middle 
or  even  late  Tertiary  times.  That  the  connection  goes  far  back  is  shown  by  the 
occurrence  of  a  Gondwana  flora  on  both  sides  of  the  Drake  passage  (Graham 
land,  Falkland  Islands),  and  the  Cretaceous  rocks  of  South  Georgia  prove,  as 
HOLTEDAIIL  says,  the  existence  of  a  land  mass  of  considerable  size  where  there 
is  now  sea.  Joyce  (<5-f)  remarks  that  "there  is  good  evidence  that  the  Scotia  Arc 
with  its  extension  into  West  Antarctica  has  persisted  as  a  structural  feature  since 
Lower  Palaeozoic  times". 

The  Scotia  passage  offers  one  of  the  migration  routes  over  land  that  we  are 
in  need  of,  but  another  passage,  the  iMacquarie  route  between  East  Antarctica 
and  Australia-New  Zealand  is  required  to  make  the  trans-antarctic  route  complete. 
There  are  intermediate  islands,  Macquarie,  Auckland  and  Campbell  Islands,  with 
a  subantarctic  flora  suggesting  former  connections,  and  there  are  tracts  with  shal- 
lower water  between  Tasmania  and  the  continent,  but  for  want  of  geological  evi- 
dence this  bridge  is  hypothetical,  and  many  authors  prefer  to  speak  of  submerged 
intermediate  islands  sufficiently  close  to  facilitate  the  spread  of  organisms  able 
to  cross  moderate  water  barriers.  As  AxELROD  says  [14..  183): 

Archijielagoes  of  only  slightly  greater  extent  than  those  now  present  could  account 
for  the  continuity  of  the  Antarcto-Tertiary  Flora  in  all  these  regions  during  the  early 
and   middle   Cenozoic. 

Florin,  referring  to  the  present  and  former  distribution  of  conifers,  expressed 
himself  in  similar  terms. 

Antarctica  has  ])layed  an  important  role  in  the  development  and  distribution  of 
the  southern  group  of  conifers.  The  data  related  to  its  distributions  considered  in  this 
paper  seem  most  readily  explained  by  assuming  land  connections,  or  at  least  much 
closer  proximity  between  Antarctica  and  the  adjacent  southern  ends  of  South  America, 
Australia,   New  Zealand   and   South   Africa  (95.92), 

and  after  the  discovery  of  a  Tertiary  species  q{  Acuwpylc  in  Patagonia  and  a  sec- 
ond fossil  form   in  the  TLocene  of  Seymour  Island,   he  wrote  [33S.  136): 

Fur  cine  ehemalige  Verkniipfung  der  australischen  Region  mit  der  Antarktis  spricht 
auch  die  Verhreitung  der  Gattung  Acmopyle.  Dass  also  die  Antarktis  in  diesem  Falle 
als  cine  alte  Vermittlerin  zwischen  der  australischen  Region  und  Siidamerika  gedient 
hat,   muss   meines   Erachtens   angenommen   werden. 

However,  one  of  the  supposed  links,  Macquarie  Island,  does  not,  it  seems, 
possess  any  plants,  perhaps  not  even  mosses  or  lichens,  dating  back  to  the  height 
of  the  Glacial  lY^riod,  fcjr  this  island  was,  at  least  during  maximum  glaciation, 
entirely  ice-covered  and  must,  Tavlor  says,  have  received  its  present  plant  world, 
very  poor  it  is  true,  in  postglacial  time  from  the  north  and  across  a  considerable 
stretch  of  open  sea  [263).  In  Antarctica  proper  the  situation  is,  with  regard  to 
lichens  and  mosses,  different.  Dahe  [^2.  231),  basing  his  opinion  on  the  discovery 
of  numerous  endemic  lichens  and  of  a  few  mosses  not  very  far  from  the  south 
pole,  concludes  that  part  of  the  flora  survived   the  glaciations.   Here,   where  high 


DERIVATION  OF  THE  FLORA  AND  FAUNA  39I 

mountain  ranges  are  found  right  along  the  coast,  the  inland  ice,  even  during 
maximum  glaciation,  cannot  have  covered  everything.  The  flora  may  not  have 
been  as  rich  in  species  as  it  is  now  but  perhaps  a  little  more  varied  than  one 
has  been  inclined  to  think.  Many  species  are  also  found  in  the  subantarctic 
zone,  and  further  research  work  will,  perhaps,  reduce  the  number  of  endemic 
species. 

Florin  spoke,  as  we  have  seen,  of  a  "proximity  to  South  Africa".  This  is 
where  most  biogeographers  hesitate,  in  spite  of  such  eloquent  facts  as  the  distri- 
bution of  Restionaceae,  Proteaceae  and  other  families,  and  the  occurrence  of  a 
subgenus  of  Gunnera  on  African  soil.  HoOKER,  it  is  true,  had  a  vision  of  a  larger 
Kerguelen  land,  but  this  was  still  a  long  way  off  from  Africa.  Perhaps  GULICK 
{^iig)  ought  to  be  mentioned  here;  he  opposed  the  continental  nature  of  isolated 
islands,  but  he  was  tempted  to  exempt  what  he  called  "continental  outsiders", 
"Kerguelen,  Crozet,  St.  Paul  and  two  or  three  more";  the  sea  lacked  the  deepness 
of  a  typical  ocean,  and  sediments  occurred  on  Kerguelen;  for  these  reasons  he 
admitted  a  possible  former  existence  of  a  "northward  lobe  of  the  Antarctic  con- 
tinent". The  lichenologist  C.  W.  DODGE  has  taken  up  this  question;  the  lichen 
flora  of  "Kerguelia"  presents  features  of  great  antiquity  as  well  as  of  prolonged 
isolation  (5  endemic  genera),  and  the  angiosperms  include  such  aberrant  types  as 
Pringlea  and  Lyallia\  it  should  be  remembered  that  Werth  was  opposed  to  over- 
seas dispersal.  Kerguelen  is  volcanic,  but  old,  the  oldest  lavas  dating  from  late 
Mesozoic  or  early  Tertiary  times,  and  on  them  fluviatile  sediments  and,  on  top 
of  these,  Oligocene  strata  rest.  Erosion  broke  down  the  island  during  Miocene- 
Pliocene,  but  renewed  volcanic  activity  followed  from  the  end  of  Pliocene  into 
Pleistocene.  The  Gaussberg-Kerguelen  ridge  connects  Antarctica  with  Kerguelen + 
Heard  Island;  an  elevation  of  400  fathoms  would  be  sufficient  to  unite  the  two 
islands,  a  rise  of  100  fathoms  would  transform  the  Crozet  group  into  a  single 
island,  and  DODGE  supposes  that  there  is  a  submarine  connection  between  Ker- 
guelen and  the  Crozet  swell.  Kerguelia  in  its  prime  would  include  all  the  islands, 
also  Marion  and  Prince  Edward.  The  great  difficulty,  the  extension  to  South 
Africa,  remains.  So  much  seems  to  be  certain  that,  if  this  bridge  did  exist, 
separation  took  place  early,  long  before  the  other  Antarctic  connections  were 
broken  off. 

I  think  that  the  majority  of  phytogeographers  agree  with  MERRILL  who,  in 
his  last  work  [306),  wrote  that 

there  is  no  reason  whatever  to  doubt  the  validity  of  this  ancient  Antarctic  route  of 
migration  of  various  families  and  genera  of  plants;  certainly,  no  experienced  phytogeo- 
grapher  would  question  the  validity  of  this  route,  for  it  is  as  thoroughly  established 
as  its  more  evident  equivalent  by  what  is  now  the  Arctic  region  (p.  178). 

Most  botanists  have  drawn  their  conclusions  from  the  present  distribution 
of  the  plants  and  this  is,  as  a  rule,  all  they  can  do,  because  few  have  left  any 
traces  of  their  distribution  in  earlier  epochs.  Nevertheless  we  have  no  good  reason 
to  doubt  the  important  part  taken  by  Antarctica  in  the  history  of  the  south 
hemisphere,    but   the  proofs  that  such  was  the  case.  Berry  emphasizes  [28.  34), 


392  C.   SKOTTSBERG 

must  rest  on  palaeobotanical  evidence,  and  it  happens  that  the  fossil  records  are 
at  variance  with  current  ideas.  The  occurrence  of  Ayaiicaria,  Driniys,  Laiwe/ia, 
XoiJiofagus  etc.  in  Tertiary  deposits  on  Seymour  Island  would  lead  us  to  infer 
that  they  are  of  Antarctic  origin  and  have  radiated  from  there,  but  Araucaria 
once  iiad  a  world-wide  distribution,  Drhuys  belongs  to  an  order — Magnoliales — 
of  Holarctic  range,  the  same  is  true  of  Fagaceae,  and  even  Laurclia  is,  Berrv 
points  out,  open  to  doubt,  in  spite  of  the  fact  that  the  Monimiaceae  are  a  south- 
ern family.  Many  other  genera,  called  Antarctic  on  the  strength  of  their  modern 
distribution,  are  known  as  fossils  in  the  north  temperate  and  Arctic  zones.  '' Araji- 
caria  stands  as  a  per[)etual  warning  against  forgetting  that  the  past  is  the  key 
to  the  {:)resent",  l^KRRV  wrote  (I.e.  36).  A  Holarctic  genus  may  have  reached  New 
Zealand  or  Australia  as  well  as  Patagonia  from  the  north,  never  having  used  an 
Antarctic  route,  and  without  leaving  a  trace  of  its  wanderings.  On  the  other  hand, 
the  little  we  know  about  the  preglacial  vegetation  of  Antarctica  is  sufficient  to 
prove  that  this  large  land  mass,  just  as  every  other  part  of  the  globe,  was  inhab- 
ited by  a  rich  and  varied  flora,  that  it  may  have  been  a  primary  centre  of 
evolution,  that,  in  other  instances,  it  served  as  a  secondary  centre  and  that  it  was 
a  much-trodden  road  between   America  and  Australia-New  Zealand. 

Miss  GlHHS  appears  to  have  been  one  of  the  very  few  experienced  phyto- 
gcogra[)hers  who  refused  to  regard  Antarctica  either  as  a  centre  or  as  a  migra- 
tion route  over  land;  it  had  always  been  surrounded  on  all  sides  by  water  and 
no  other  agents  than  "the  wild  west  wind"  [106.  103)  and  a  pole-ward  north-west 
wind,  coming  from  Asia,  were  needed  to  explain  every  distribution  pattern.  The 
southern  focus  of  development  was  not  Antarctica  but  the  mountains  of  New 
(kiinea.  The  highland  of  Tasmania,  the  subject  of  her  survey,  had  received 
nothing  from  the  south,  all  the  so-called  Antarctic  plants,  genera  lika  Ahroiajtclla, 
Astdia,  C'arp/ia,  Colobajiihus,  Coprosnia,  Driinys,  Gaimardia,  Gunner  a,  Lage?io- 
pliora,  Xotliofagus,  Orcobolus  and  so  on,  had  come  from  New  Guinea,  and  from 
there  they  had  radiated  to  Polynesia,  Hawaii,  Juan  P^ernandez,  Tierra  del  PTiego 
and,  I  presume,  Antarctica.  Had  she  lived  to  hear  of  the  discovery  of  A7v//^<7^//i' 
in  New  (luinea,  where  more  species  have  been  found  than  anywhere  else,  and 
in  .\ew  Caledonia,  and  of  the  rich  development  of  the  Winteraceae  in  New  Guinea, 
she  wf)ul(l  have  regarded  such  finds  as  a  forcible  proof  of  the  correctness  of  her 
opinion. 

Many  /ooge()gra{)hers  have  looked  with  much  suspicion  at  the  Antarctic 
continent  as  a  centre  of  radiation.  Sl.Ml'Sox,  in  his  review  of  the  theories  involving 
Antarctica  in  the  distribution  of  vertebrates  (^^,?),  concluded  that  dispersal  had 
been,  in  all  cases,  from  north  to  south;  not  even  the  Scotia  Arc  had  ever  been 
used  as  a  route  of  migration.  His  reasoning  is  logical  and  often  conclusive.  The 
invertebrates  are,  however,  left  aside.   To  quote  part  of  his  summary  (p.  T^y): 

There  is  no  known  bioti(  fact  that  demands  an  Antarctic  land-migration  route  for 
its  explanation  and  there  is  none  that  it  more  simply  explained  by  that  hypothesis 
than  by  any  other.  The  affinities  of  the  southern  faunas  as  a  whole  are  what  would 
he  exj)ected  from  the  present  northern  connection  known,  or  with  considerably  prob- 
ability   inferred,    to    have    existed    at   ai)propriate  times  in   the   past.   'J'here  are  certain 


DERIVATION  OF  THE  FLORA  AND  FAUNA  393 

troublesome  anomalies  and  exceptions  in  the  evidence,  but  none  of  these  can  be  ade- 
quately explained  by  postulating  an  Antarctic  connection.  The  general  weight  of  evi- 
dence is  against  such   a  connection. 

In  scientific  theory  the  best-supported  and  most  nearly  self-sufficient  hypothesis 
should  be  preferred  and  unnecessary  additional  hypotheses  should  be  rejected  or  held 
in  abeyance.  On  this  basis  the  Antarctic  migration  route  hypothesis  remains  simply  a 
hypothesis  with  no  proper  place  in  scientific  thinking. 

To  this  I  shall  make  a  few  remarks.  If  Simpson  had  said  "no  fact  involving 
the  vertebrates"  instead  of  "no  biotic  fact" — very  well,  let  us  assume  that  the 
routes  across  from  and  to  either  America  or  New  Zealand  were  impassable  to 
mammals,  reptiles,  amphibia  and  flightless  land-birds,  birds  with  good  flight  capa- 
city would  have  found  little  difficulty  to  cross,  and  many  biotic  facts  are  known 
that  clearly  speak  in  favour  of  an  Antafctic  migration  route  for  invertebrates  and 
plants.  Simpson  must  have  thought  that  either  did  the  Antarctic  continent  never 
possess  a  fauna  of  land  vertebrates,  or,  if  it  did  have  one,  it  had  evolved  inde- 
pendently of  all  other  faunas  and  disappeared  without  leaving  a  single  trace. 
Only  penguins  are  known  in  a  fossil  state  in  Antarctic  Tertiary  deposits.  Until 
fossil  land  vertebrates  are  discovered,  the  question  of  the  former  existence  of  an 
Antarctic  fauna  of  terrestrial  vertebrates  must  be  left  open. 

Among  the  invertebrates  are  many  examples  of  a  discontinuous  distribution 
most  readily  understood  if  Antarctica  is  taken  into  account.  Several  were  men- 
tioned in  the  chapter  devoted  to  the  composition  of  the  fauna  of  Juan  Fernandez, 
a  few  more  may  be  quoted  here.  Berland,  dealing  with  the  Pacific  spider  fauna  (2j): 

Nous  avons  tire  de  notre  etude  cette  notion  importante  que  la  liaison  entre  I'Aus- 
tralie  et  I'Amerique  a  eu  lieu  par  une  terre  antarctique  dont  les  temoins  restent  ac- 
tuellement,  mais  ni  par  la  Nouvelle  Caledonie,  ni  par  la  Nouvelle  Zelande,  ni  par  le 
groupe  Samoa-Tonga-Fiji,  et,  par  voie  de  consequence,  qu'elle  n'a  pas  eu  lieu  par  le 
centre  du  Pacifique  (p.  1053), 

Berlioz  (2^),  with  examples  off'ered  by  the  distribution  and  affinities  of  beetles, 
states  that  the  group  of  Buprestidae,  forming  "le  noyau  essentiel"  in  the  Bupres- 
tid  fauna  of  Australia,  has  mainly  South  American  affinities  (several  genera), 
and  that 

la  faune  des  Lucanides  d'Australie  et  de  Papouasie  presente  avec  celles  de  I'Amerique 
du  Sud  surtout  de  la  region  andine  et  patagonienne,  des  affinites  aussi  etroites  que 
curieuses. 

Finally  Lindsay  (idy),  calling  attention  to  the  Subantarctic  Collembola, 
extremely  delicate  creatures  "supposed  not  to  be  carried  any  appreciable  distance 
either  by  wind  or  sea,  thus  being  important  proofs  of  former  land  connections", 
mentions  a  genus  of  3  species  of  which  one  is  Fuegian,  one  recorded  from 
the  Scotia  Arc,  and  one  found  in  New  Zealand.  And  other  similar  examples  may 
be  found. 


394 


C.   SKOTTSBERG 


Chapter  XIII. 

The  history  of  Juan  Fernandez — a  tentative  sketch. 

In  my  first  [)aper  on  tlie  Botany  of  Juan  Fernandez  (22/^),  where  only  the 
vascular  plants,  as  known  at  that  time,  were  included,  I  expressed  my  view  on 
the  history  of  the  flora  in   the  following  words: 

Frcilich  haben  wir  keine  Ahnung  davon,  wie  schnell  Arten  oder  (lattiingen  ent" 
stehen,  aber  wir  konnen  uns  kaum  denkcn,  dass  in  der  kurzen  Zeit,  die  seit  der  Ent" 
stehung  der  jetzigen  Inseln  verHossen  ist,  sich  Typen  wie  Lactoris  oder  Rohinsojiia  aiis 
"Keimen"  entwickelten,  die  nach  den  Inseln  gebracht  wiirden,  um  sich  in  ungestorter 
Isolierung  umzutormcn  .  .  .  Ich  bin  der  Meinung,  dass  das  alte  Klement  nicht  auf  Masa- 
ticrra  oder  Masafucra  entstand,  sondern  alter  ist  als  die  jetzigen  Inseln,  und  dass  es 
wenig  wahrschcinlich  ist,  dass  die  alten,  endemischen  (oder  andere,  eng  vervvandte) 
(iattungcn  und  Arten  von  Juan  Fernandez,  noch  nachdem  die  Inseln  gebildet  waren, 
die  vielen  vermeintlichen  Ursprungsorte  bewohnten,  und  dass  Veriinderungen  in  der 
I'flanzenwelt  von  Polynesian,  Neuseeland,  Chile  u.s.w.  in  (puirtiirer  Zeit  die  isolierte  Stel- 
lung  bewirkt  haben.  Ich  glaube  also,  dass  in  vor-  und  friihtertiarer  Zeit  grossere  Ent- 
wickelungszentra  existierten,  und  dass  ihre  Flora  nunmehr  als  ein  altpazifischer  Rest 
fortlebt.  Auch  eine  Resttlora  wird  sich  aus  vielen  Familien  und  (iattungen  aber  ver- 
ba! tnismiissig   wenigen   Arten   zusammensetzen. 

At  that  time  our  knowledge  of  the  flora  was  incomplete;  much  fresh  and  new 
material  was  added  in  19 16-17.  I  had  fixed  my  attention  on  what  I  called  the 
"Old  Pacific  element",  but  did  not  venture  to  look  for  an  exact  site  of  an  evolu- 
tion centre,  though  the  possibility  of  Antarctica  as  an  important  source  of  genera 
and  families  had  been  pointed  out  before;  the  first  sign  of  the  presence  of  there- 
tofore unknown,  subantarctic  flora  in  Masafuera  had  been  observed,  but  I  had  no 
reason  to  link  it  with  the  Old  Pacific  types.  During  the  19 16-17  survey  a  rather 
strong  Antarctic-bicentric  group  took  shape,  and  in  my  1925  sketch  of  the  history 
of  the  flora  (.?.?/)  not  only  was  this,  but  also  the  Old  Pacific  plants  claimed  to 
have  "reached  Juan  F'ernandez  over  South  America,  where  they  have  disappeared" 
(p.  31).  My  object  this  time  is  to  see  if  we  can  approach  these  problems  in  other 
than   general    terms. 

Our  starting  point  is  the  "Tierra  de  Juan  P'ernandez"  of  BrCckien,  forming 
a  westward  extension  or  lobe  of  South  America,  reaching  the  actual  site  of  Juan 
PY'rnandez  and  Desventuradas  Islands,  as  indicated  by  the  bathymetrical  conditions, 
something  in  keeping  with  the  peninsula  of  Lower  California  and  separated  from 
the  coast  by  a  broad,  toward  the  south  gradually  narrowed  bay.  However,  if  we 
remember  that  the  deep  trough  is  supposed  to  have  originated  with  the  final 
uj)lift  of  the  Andes,  the  j)resent  coast  line  ought  to  be  recent  all  along,  but  this 
is  not  in  accordance  with  the  opinion  that  the  P^ocene  Arauco  flora  was  a  coast 
flora.  The  conclusion  would  be  that  the  great  depths  were  initiated  already  during 
Cretaceous  times,  getting  dec|)er  and  deeper  with  the  successive  periods  of  uplift. 
The  Juan  PY'rnandez  land  or  peninsula  formed  part  of  the  neotropical  P2ocene  flora 
region  which  extended  from  Venezuela  and  Brazil  to  south  Chile  and  east  across 
the  mountains,  during  this  era  of  low  relief,  as  shown  by  the  fossiliferous   beds 


DERIVATION  OF  THE  FLORA  AND  FAUNA  395 

of  Pichileufii.  As  the  Concepcion-Arauco  flora  it  was  a  subtropical  rain  forest 
flora  with  podocarps,  tree  ferns,  evergreen  dicotyledonous  trees  and  lianas,  be- 
longing to  some  30  tropical  families;  the  species  were,  with  two  exceptions,  referred 
by  Berry  to  still  existing  genera — if  the  determinations  are  reliable.  This  is 
perhaps  more  than  we  can  expect;  all  we  can  say  is  that  Berry  was  a  man  with 
a  wide  experience  of  both  fossil  and  living  tropical  plants  and  that  undoubtedly 
many  of  the  families  listed  and  perhaps  also  a  fair  number  of  the  genera  are 
correctly  placed.  Two  are  found  in  the  present  flora  of  Juan  Fernandez.  To  Azai-a 
celastriniformis  and  tertiaria — and  the  fossil  does  suggest  Azara — BERRY  remarks 
{28.  107): 

As  a  recent  form  occurs  on  the  island  of  Juan  Fernandez  one  can  predicate  a  con- 
siderable antiquity  'for  the  genus,  which  is  more  than  verified  by  the  present  fossil  forms. 

His  Berberis  corynibosiflora  is  of  still  greater  interest: 

I  have  seen  leaves  of  all  the  South  American  species  and  the  most  similar  is  Berberis 
corymbosa  Hook,  et  Arn.  of  Juan  Fernandez  (i.e.  75). 

I  have  compared  his  illustrations  with  the  island  species  and  I  am  willing  to 
testify  to  the  striking  similarity  between  them.  A  revision  of  these  most  important 
fossil  floras,  with  application  of  modern  technique,  is  eagerly  longed  for. 

Anyhow,  the  neotropical  character  of  the  old  flora  has  been  safely  estab- 
lished, and  Arauco  and  Pichileufu  agree  in  their  general  composition;  20  species 
occur  in  both.  If  this  flora  extended  to  Juan  Fernandez,  this  land  must  have  been 
sufficiently  high  to  force  the  prevailing  westerly  winds  to  unload  part  of  their 
moisture  and  to  give  rise  to  altitudinal  belts.  The  flora  most  likely  had  its  special 
distinctive  marks.  In  view  of  the  very  large  area  it  inhabits,  it  cannot  have  been 
uniform,  and  different  floristic  provinces  showed  special  features  and  had  their 
own  endemics.  If  anything  still  survives  in  identical  or  very  similar  form  can  only 
be  a  subject  of  conjecture  and  is  not  demonstrated  by  leaf  impressions.  Two  of 
the  genera  reported  from  the  mainland,  Azara  and  Myrceugenia,  still  occur  in 
Chile  and  Juan  Fernandez,  and  I  am  inclined  to  believe  that  the  endemic  element 
in  the  insular  forest  flora  dates  back  to  early  Tertiary  times,  genera  like  Podo- 
phorus,  Megalachne,  Jiiaiiia,  Ochagavia,  Nothornyrcia  and  Selkirkia,  and  species 
of  Chusquea,  Hesperogreigia,  Urtica,  Phrygilanthus,  Cheiiopodinin,  Colletia,  Dys- 
vpsis,    Ugni,  Eryngium,  RJiaphithamnus,  Solanum  and  Nicoiiana. 

Centaurodeiidron,  Yunquea  and  the  four  endemic  Cichoriaceous  genera  stand 
apart.  They  are  montane  and  we  have  no  clue  at  all  to  their  history,  but  w^e  can 
take  it  for  granted  that  they  are  not  "new  beginners",  but  old  relicts,  without 
any  near  relatives  anywhere.  Whether  we  assume  that  they  arose  in  the  islands 
and  have  left  no  marks  in  the  continent,  or  derive  the  four  dendroseroid  genera, 
which  form  a  natural  group,  from  Antarctic  ancestors,  a  possibility  certainly  not 
ofl"ered  by   Ceiitaiirodendron  and    Yunquea,  we  are  victims  of  wild  speculation. 

Few  endemic  ferns  belong  to  the  neotropical  element,  Trichomanes  bigae, 
Dryopteris  maequalifolia,  Asple7t2uni  macrosorum  and  stellaium,  Pellaea  chilensis, 


396 


C.   SKOTTSBERG 


Polvpodhini  interniednnji  and  Ophioglossiim  ferjiiutdezianmn,  but  PrZ/^rrt' and  OpJiio- 
glossum  do  not  grow  in  the  forest,  the  former  inhabiting  the  dry  coast  chffs,  the 
hitter  tlie  open  grass  kind,  and  they  may  have  their  own  history. 

How  and  when  the  non-endemic  South  American  species  reached  Juan  Fer- 
nandez is  hard  to  tell.  They  are  temperate  and,  with  the  exception  of  il/j'r/^^f^/rt 
)ni))n)nilaria,  herbaceous.  Myrteola  is  the  only  member  of  this  group  that  extends 
south  to  the  subantarctic  zone.  Of  the  others  Dantlionia,  Koeleria,  Stipa,  Piptochae- 
tiii))i,  lUfocliaris,  JuJiciis  procerus  (also  douibeyanus  and  iiubricatusi),  LUiertia, 
Pipiroj}iia  fcniandcziaua,  Parictaria  and  Miniulus  show,  in  their  mode  of  occur- 
rence, every  sign  of  being  indigenous.  Advocates  of  transoceanic  dispersal  would 
not  hesitate  to  call  them  "late  arrivals  which  have  not  had  time  to  change",  and 
a  direct  transport  is  not  altogether  impossible.  It  would  be  interesting  to  know 
if  a  grass  land  existed  when  Great  Juan  Fernandez  was  connected  with  the  mainland, 
but  unfortunately  we  do  not  even  know  the  extension  of  the  Stipeiuvi  when  the 
early  voyagers  reported  on  the  vegetation  and  already  found  the  introduced  Avena 
barbata  in  dominance  in  the  treeless  western  part  of  Masatierra.  A  remark  made 
by  1^rC(;gex  deserves  to  be  quoted  in  this  connection.  A  current  coming  from 
the  south  swept  past  the  shore  of  Great  Juan  Fernandez.  When,  during  the  Na- 
\idad  transgression,  separation  from  Chile  occurred, 

.  .  .  cl  mar  del  polo  sur  entro  en  comunicacion  con  el  mar  que  banaba  las  costas  de  Chile 
Central  y  cl  primer  antecesor  de  la  corriente  de  Humboldt  llevo  las  aguas  inas  frescas  hacia 
el  norte,  dando  principio  a  la  gran  zona  desertica. 

HrCG(;kn  seems  to  have  forgotten  that  at  that  time  Antarctica  was  covered, 
not  by  an  inland  ice,  but  by  luxuriant  vegetation,  and  that  the  sea  cannot  have 
been  cold;  however,  there  must  have  been  an  uppwelling  of  cold  water,  and  the 
dry  climate  of  the  basal  belt  may  have  been  as  unfavourable  for  tree  growth  as 
it  is  now.  Nevertheless  1  cannot  believe  that  the  present  steppe-like  communities 
date  back  to  early  or  even  middle  Tertiary  time.  On  the  other  hand  it  seems 
(luite  unlikely  that  the  s[)ecies  of  Stipa,  PiptocJiaetiu))i,  DantJionia,  etc.  were  in- 
troduced with  the  traffic  while,  in  this  respect,  Chaetoiropis  (the  endemic  nature 
of  i  h.  inibcrbis  (juestionable),  the  two  Cypcrus,  jf uncus  capillaceus,  Paronychia, 
Ccnlclla.  Ilcdyotis  and  Plantago  iruncata  are  under  suspicion. 

Of  the  ferns  found  elsewhere  Polypodium  lanceolalurn  is  pantropical  and  old 
enough  to  have  belonged  to  the  ancient  flora,  and  this  may  be  true  also  of 
7'richouiaues  exsccfuiu,  Hy))ieiiopJiyllum  spp.,  Adianiuni  chilcnse,  Ptcris  chiknsis  and 
scmiad)iata.  and  lilaplioglossui)i\  1  have  suggested  that  the  latter  was  carried 
directly  to  Masatierra  by  a  northerly  storm,  and  this  could  have  been  the  case 
also  with  Polypodium  Masafucrac,  observed  a  single  time  lOO  years  ago  and  never 
again.  P.  tric/ioiuauoides  remains  doubtful  in  spite  of  the  specimens  still  extant 
and  labelled  Juan    hY'rnandez  {2^(j.  766). 

The  presence  of  a  large,  presumably  boreal  element  is  not  difficult  to  explain, 
for  it  extends  all  along  the  Andes  to  the  far  south.  Many  of  the  species  are 
endemic,  Agrostis  juasafucraua,  2  sj).  of  Spergularia,  2  Berberis,  Cardamine  Krues- 


DERIVATION  OF  THE  FLORA  AND  FAUNA  397 

selii,  Galium  masafuerajtuni  and  6  Erigeron.  The  Boreal  character  of  these  genera 
is  recognized.  This  element  extended  south  along  the  precursors  of  the  late  Tertiary 
mountains;  the  species  of  Agrostisi^^  Spergularia,  Berberis,  Cardamine  and  Ga- 
lium have  their  closest  relatives  either  in  the  tropical  Andes  [Berberis]  or  in  Chile. 
Erigerofi  deserves  special  attention  on  account  of  the  large  number  of  Andean 
species  nearly  related  to  each  other  but  less  so  to  the  insular  group,  which  shows 
a  remarkable  differentiation:  E.fruticosus  and  its  cognate  luteoviridis,  the  three 
herbaceous  rosette  herbs,  and  the  peculiar  E.  rupicola  of  the  coast  rocks;  of  these 
fruticosus  is  found  on  both  islands  and  the  other  species  endemic  on  Masafuera. 

Nine  non-endemic  species,  all  found  in  Chile,  also  belong  here,  Trisctum,  Carex 
Banksii,  Pa^'onychia,  Cardami^ie  flaccid  a,  Callitriche,  Rubus,  Empeirum,  Calysiegia 
and   Gnaphalium,  and,  among  the  ferns,  perhaps   Cysiopteris.  Rubus geoides  forms 
together  with  R.  radicans  an  isolated  section  but  has  a  more  southerly  distribution 
but  all  may  have  reached  Juan  Fernandez  from   South  Chile. 

Finally,  Cuminia  remains  to  be  accounted  for.  Whether  w^e  link  it  with  the 
palaeotropical  Prasioideae,  which  seems  to  be  the  best  way,  or  with  Bystropogon, 
it  stands  out  as  an  isolated  relict  genus. 

I  have  distinguished  a  large  Antarcto-tertiary  element,  over  40  %  of  the 
angiosperms  and  60  %  of  the  ferns.  Among  the  former  there  are  three  or,  if  Lac- 
toris  is  kept  aside,  two  groups;  one  of  them  (i)  is  still  represented  in  South 
America,  the  other  (2)  not.  In  (i)  tw^o  types  can  be  distinguished,  (a)  not  confined 
to  subantarctic  or  alpine  habitats  and  demanding  a  milder  climate;  to  this  lot  I 
refer  Uncinia  Douglasii  and  costata,  Drimys,  Phrygila7itkus,  Escallonia,  Margyri- 
carpus,  Sophora,  Guiuicra,  Apium,  Pernettya,  and  possibly  Planiago  fernandezia. 
All  have  relatives  in  Chile.  To  these  are  added  the  species  also  occurring  on  the 
mainland:  Danthonia,  Koeleria,  Juncus,  Libertia,  Acaena  ovalifolia  (indigenous?)^ 
Centella  and  Nertera\  the  two  grasses  are,  however,  only  tentatively  referred  to 
this  element. 

The  occurrence  of  an  Antarctic  element  in  the  Eocene  flora  of  the  mainland 
has  been  demonstrated.  Berry  lists  Araucaria,  Libocedrus,  Drimys,  Embothrium, 
Laurelia  and  Eucryphia  from  the  Pichileufu  beds;  even  if  the  "magnolia  stock" 
is  of  Boreal  origin  this  does  not  exclude  the  possibility  that  the  Winteraceae 
radiated  from  Antarctica,  a  parallel  case  to  Fagaceae  and  Nothofagus. 

If  we  follow  COPELANI)  many  of  the  ferns  also  belong  to  i  a:  Hymenoglossum, 
five  species  of  Hymeiwphyllum,  Lopkosoria,  Polystichum,  Bleclnium,  Hypolepis, 
Histiopteris,   Gleichenia  pedalis  (indigenous.-)  and  Lycopodium  scariosum. 

Group  1  b  includes  the  so-called  Subantarctic-Magellanian  element  inhabiting 
Fuegia,  the  Falkland  Islands,  etc.  ranging  north  along  the  Andes  and  belonging 
to  a  well-knowm  circumpolar  assemblage  of  genera  and  species.  Here  we  find 
Oreobolus,  Uncijiia  brevicaulis,  phleoides  and  tenuis,  and  Lagenophora  hirsuta, 
further  three  endemic  species,  Acaeiia  masafuerana,  Abrotaiiella  crassipes,  both 
with  near  relatives  in  West  Patagonia-Fuegia,  finally  Agrostis  masafuerana,  if  its 
relation  to  the  bicentric  A.  magellanica  is  confirmed  after  monographic  treat- 
ment. I  brought  it  to  the  boreal  group.  The  following  pteridophytes  are  attached 


398  C.   SKOTTSBERG 

here:   Serpyllopsis,   lIy))uniopliyllui)i  falklandicuni,  Polypodium  (Grammitis)  niagel- 
latiiciiiu,   GlcicJioria  quadripartiia,  and  Lycopodium  inag€lla7naivi. 

Did  this  Antarctic  element,  mostly  not  endemic,  extend  to  Great  Juan  Fer- 
nandez or  did  it  arrive  after  the  separation  from  the  mainland  took  place,  per- 
haps even  after  that  the  volcanic  islands  had  been  formed?  The  same  question 
was  raised  when  we  discussed  the  non-endemic  neotropical-temperate  Stipa,  Pipto- 
cliactiu)n,  Myrtio/a,  Rn/uis,  and  so  forth;  the  Boreal  group  is,  as  we  have  seen, 
also  inxolved.  Is  it  probable  that  also  this  flora  dates  back  to  early  or  middle 
Tertiary  times:  This  seems  unlikely.  The  subantarctic  species  are,  with  the  excep- 
tion of  GrauDJiitis,  restricted  to  the  highland  of  Masafuera,  but  may  have  occurred 
also  on  Masatierra  when  the  islands  stood  higher.  Either  we  must  assume  that 
the  land  connection  with  the  continent  persisted  much  longer  than  is  otherwise 
probable,  or  those  species  have  immigrated  across  the  water  in  late  Pliocene  or 
in  postglacial  time.  I  have  suggested  this  on  repeated  occasions  {2JI,  2^2,  340). 
However,  we  must  not  forget  that  we  have  to  do,  not  with  stray  colonists,  but 
with  plant  communities  composed  by  flowering  plants,  ferns,  bryophytes  and 
lichens.  Unfortunately  nobody  beheld  the  vegetation  before  introduced  species, 
AuthoxiDitJniui  odoratiiDi  and  Rumex  acctosella,  had  invaded  the  highland  and 
changed  the  entire  aspect  beyond  recognition. 

To  distinguish,  among  the  bryophytes,  the  old  element  which  undoubtedly 
must  have  formed  an  important  part  of  the  subtropical  forest  flora  is  more  than 
I  can  undertake;  we  do  not  know  if  the  many  endemic  species  are  relicts  or  not; 
in  fact,  we  do  not  even  know  if  they  are  endemic  until  the  opposite  mainland 
has  been  well  explored.  So  much  can  be  said  that  species  with  a  pronounced 
tropical  distribution  are  few,  but  if  we  add  the  Chilean  species  extending  through 
the  Valdivian  and  Magellanian  forest  zones,  this  South  American  group  makes 
u[)  about  40  "o  of  the  mosses  and  30%  of  the  hepatics,  endemic  species  of  Ameri- 
can affinity  included.  Perhaps  half  a  dozen  mosses,  not  counting  the  few  that 
accompany  man  wherever  he  goes,  and  some  liverworts,  have  a  wide  distribution 
outside  America.  The  dominant  element  is  Antarctic;  this  was  emphasized  by 
such  authorities  as  Cakdoi'  and  Hkrz()(;:  56%  of  the  mosses  and  68%  of  the 
hej)atics  were  referred   to  the  Antarcto-tertiary  element. 

When  it  comes  to  distinguishing  corresponding  groups  among  the  lichens  we 
move  on  very  unsafe  ground,  but  there  are  indications  that,  beside  a  large  South 
.American  element,  we  also  ha\e  an  Antarctic  group  to  which  no  less  than  58 
species  were  referred.  They  are  austral-  or  subantarctic  bicentric  or  tricentric, 
but  many  of  them  range  north  into  lower  latitudes.  A  conspicuous  part  is  formed 
by  Stictaceae.  The  great  bulk  of  the  family  is  by  no  means  southern,  Lobaria, 
Sticla  and  PscudocypIuUaria  are  frequent  in  tropical,  subtropical  and  temperate- 
f)ceanic  climates  throughout,  but  the  austral-circumpolar  species  are  so  many  that 
we  cannot  exclude  Antarctica  as  a  possible  source. 

We  shall  {)roceed  to  group  2.  It  comprises  the  genera  or  species  which  are 
foreign  to  the  South  American  flora  and  have  their  relations  in  Australasia.  JoilOW 
recogni/x'd  very  few;  he  had,  just  as  several  later  authors,  no  other  explanation 
to  offer  than  that  they  had  arrived  across  the  Pacific  from  Australia,  New  Zealand, 


DERIVATION  OF  THE  FLORA  AND  FAUNA  399 

the  East  Indies,  etc.  without  reaching  the  coast  of  Chile.  They  are  much  more 
numerous  than  JoilOW  thought,  28  species:  Cladium,  Carex  bertero7iiana,  Pepero- 
mia  berteroana,  margaritifera  and  Skottsbergii,  Boehmeria,  Santalum,  Ranmuulns, 
Fagara  (2),  Halorrhagis  (3),  Euphrasia,  Coprosma  (2),  Waklenbergia  (5),  Robinso- 
7iia  (5),  Syniphyochaeta  and  Rhctmodendron,  the  three  last  genera  endemic.  Objec- 
tions may  be  raised  against  including  Carex  and  Euphrasia;  the  section  to  which 
Carex  bertero7iiana  was  referred  by  KCkenthal  is  almost  confined  to  New  Zealand 
and  barely  represented  in  Australia,  Tasmania  and  Norfolk  Island,  but  one  little 
known  Chilean  species  is  included,  and  Euphrasia  formosissima  is  distantly  related 
to  E.  perpusilla  of  South  Chile.  The  species  of  Wahlenbergia  are  puzzling,  but 
I  have  given  my  reasons  for  bringing  them  here  as  representing  an  African  sector. 
The  most  eloquent  members  are,  perhaps,  Santalum,  Ranunculus,  Halorrhagis  and 
Coprosma. 

This  element  is  conspicuous  also  among  the  ferns:  the  extremely  old  Thyr- 
sopteris,  ArtJiroptcris,  entirely  unfamiliar  with  the  neotropical  flora,  Dickso7tia, 
Blechnum  ScJiottii  and  Pteris  berteroana. 

I  never  looked  in  earnest  for  a  direct  road  across  the  south  Pacific  from 
Australasia  to  Juan  Fernandez,  a  route  which  ought  to  have  had  South  America 
as  its  terminus.  I  preferred  to  think  that  the  group  in  question  reached  the  islands 
over  the  Scotia  bridge  and  South  America  where,  however,  it  had  become  extinct. 
To  prove  this  we  must  turn  to  palaeontological  evidence.  The  Eocene  beds  on 
the  mainland  contain  leaves  of  many  different  plants,  and  it  is  not  impossible 
that  a  revision  of  the  material  will  contribute  to  a  solution  of  the  problem.  In 
the  lists  published  by  Berry  two  items  call  for  attention,  Cyatheoides  tJiyrsopt- 
eroides  in  the  Arauco  flora,  and  Coprosma  from  Pichileufu,  but  the  material  is 
sterile.  It  is  true  that,  to  judge  from  Berry's  illustrations,  Cyatheoides  suggests 
Thyrsopteris,  but  the  author  later  [28.  57)  compared  it  with  h\s  Dickso7iia  patago- 
nica,  which  was  found  with  sori  and  undoubtedly  belongs  to  the  Cyatheaceae. 
Thyrsopteris-like  fossils  have  been  reported  from  various  places  in  the  north 
hemisphere.  He  described  2  species  of  Coprosma,  based  on  leaf  impressions 
which,  as  far  as  I  can  see,  tell  us  little  about  their  systematic  position.  To 
C.  spathulifolia  he  remarks: 

These  tiny  leaves  have  occasioned  a  good  deal  of  trouble,  as  the  South  American 
representatives  of  the  genus  are  not  similar  to  the  fossil.  .  .  .  The  Chilean  species  are 
not  closely  similar.  .  .  . 

and  to   C.  incerta,  a  most  appropriate  name: 

.  .  .  they  are  so  much  like  the  endemic  species  of  Coprosma  of  the  Juan  Fernandez 
Islands  and  several  forms  from  the  Hawaiian  Islands  that  I  feel  constrained  so  to 
identify  them,   at  least  tentatively.  .  .  . 

I  cannot  find  that  they  agree  better  with  Coprosma  than  with  many  other 
genera.  When  Berry  gives  the  distribution  of  the  genus  as  "from  the  Malayan 
archipelago  through  the  Pacific  islands  to  Chile"  he  includes  Juan  Fernandez  under 
Chile  where,  politically,  the  islands  belong,  for  there  are  no  species  on  the  mainland. 


400 


C.   SKOTTSBERG 


Jt  is  very  easy  to  construct  a  liypothetical  passage  by  which  the  ""  Coprosnia 
group"  readied  Juan  Fernandez  without  crossing  the  Antarctic  or  encroaching  very 
much  upon  the  surface  of  the  Pacific  Ocean.  If  we  have  reason  to  think  that  South 
America  extended  farther  west  I  cannot  see  why  this  wasn't  tiie  case  also  with  West 
Patagonia  and  Tierra  del  h\iego,  a  region  which  undoubtedly  has  undergone  con- 
siderable submergence;  we  have  to  count  with  a  wide  Scotia  bridge  and  an  exten- 
sion of  Palmer  (Graham)  Land,  where  the  geographical-geological  situation  is  the 
same  as  in  South  America  and  where  the  uplift  of  the  mighty  "Antarctandes" 
ought  to  have  been  accompanied  by  submergence  of  the  fore-land.  Plants  and 
animals  could  have  travelled  by  a  circuitous  route  from  the  New  Zealand  region 
over  West  Antarctica  to  Juan  P^rnandez  without  finding  their  way  east  to  what 
is  n(nv  Chile.  This  south  Pacific  path  was  suggested  above  when  I  tried  to  divide 
the  angiosperms  according  to  their  supposed  primary  sources;  the  species  involved 
are  enumerated  under  \:  3,  forming  a  group  "as  far  as  known  without  continental 
American  affinities,  either  suggesting  an  ancient  Antarcto-Pacific  track  east  from 
Australasia  without  reaching  America,  or  having  arrived  along  the  road  over  the 
Scotia    Arc  without    leaving  any  traces   in   the  present  American  flora"'  (p.  269). 

This  idea  of  a  South  Pacific  track  is  not  new.  It  was  postulated  by  Arldt 
as  a  South  Pacific  bridge  and  it  finds  an  expression  in  Croizat's  South  Pacific 
base-line,  which,  however,  if  it  existed,  hardly  permits  us  to  draw  such  far-reaching 
conclusions  as  he  did.  In  the  case  of  Juan  Fernandez  only  13  genera  are  concerned, 
belonging  to  11  families,  the  ferns  not  included,  but  they  make  up  20%  of  the 
angiosperms,  and  others  may  have  existed  that  disappeared  later. 

It  remains  to  see  if,  among  the  cryptogams,  a  '' Coprosnia  group"  can  be 
recognized.  In  IlKRZ()(;'s  paper  on  the  Hepaticae  [130)  a  single  species  is  indicated 
as  restricted  to  Xew  Zealand  and  Juan  PY^nandez,  Palla2>iciuia  xiphoidcs,  and  two 
endemic  s{)ecies  are  said  to  have  their  nearest  allies  not  in  South  America  but 
in  Xew  Zealand.  \o  case  equal  to  Pallainci)iia  is  found  among  the  mosses,  but 
several  endemic  species  are  considered  to  be  related,  not  to  American  ones,  but 
to  s{)ecies  inhabiting  the  south-west  Pacific  region. 

With    regard    to  tlie  faioia  I  shall  confine  myself  to  some  general  remarks. 

The  few  land-birds  are  of  neotropical  origin.  Of  the  seven  species,  three, 
liusttpluvius  i:;aleyilus,  the  owl  and  the  thrush,  occur  in  identical  forms  on  the 
mainland;  the  remainder  are  either  endemic  varieties  or  endemic  species.  The 
most  divergent  is  linstipluvms  fcniaiidoisis.  (ioiri  sen's  idea  that  it  originated  in 
the  islands  as  a  mutation  of  /:".  o^alcritus  is  contradicted  by  the  fact  that  they  are 
not  at  all  closely  related  but  even  brought  to  different  genera  by  some  ornitho- 
logists. The  former  is  a  relict,  the  latter  i)erhaj)s  a  late  immigrant.  Of  the  breeding 
sea  birds  the  genus  l^teroihoiiia  forms  an  austral-circumpolar  element  and  may,  in 
j)reglacial  time,   have  inhabited  the  coasts  of  Antarctica  and  adjacent  islands. 

Little  can  be  said  as  }'et  about  the  invertebrates.  The  endemic  leech,  Ncso- 
philac))i())i,  is  an  important  case  of  non-American  ancestry,  and  the  only  terrestrial 
amphi[)od  is  bicentric.  The  spider  fauna  is  an  appendix  to  the  fauna  of  South 
America,  but  with  special  features;  there  is  no  endemic  genus,  but  specific  endem- 


DERIVATION  OF  THE  FLORA  AND  FAUNA  40I 

ism   is   high.  In  the  fauna  of  the  mainland  Antarctic  affinities  have  been  stated 
to  occur  (Berland  2j.  1044): 

The  New  Caledonia,  New  Zealand  group  has  affinities  with  the  Malaysian  region 
and  still  more  with  Australia.  But  the  small  islands  situated  south  of  New  Zealand, 
namely  the  Campbell,  Auckland  and  Macquarie  Islands,  are  different;  they  present, 
rather  abundantly,  a  group  of  spiders,  Cybaeinae,  relatives  of  which  are  found  in  the 
extreme  south  of  South  America;  these  spiders  are  not  present  in  New  Zealand,  but 
are  allied  to   Australian  and  Tasmanian   species. 

This  is  an  interesting  observation,  for  it  is  known  that  connections  between 
East  Antarctica  and  lands  to  the  north  have  been  looked  for  both  with  New  Zea- 
land and  over  Tasmania  with  East  Australia. 

Among  the  millipedes  Aulacodesimis  and  Nesogeophilus  are  austral  genera 
and  the  species  endemic;  Schizotaenia  alacer  is  known  from  Chile,  but  the  genus 
is  eminently  austral-bicentric.  For  the  same  reason  the  endemic  genus  of  Thysa- 
nura  merits  to  be  noticed. 

A  very  great  number  of  insects  have  been  reported  from  Juan  Fernandez, 
most  of  them  endemic,  also  many  of  the  genera.  The  majority  has  been  described 
only  recently  and  very  often  nothing  was  said  about  their  relations;  where  they  were 
stated  they  are,  as  a  rule,  to  be  found  in  South  America.  Isolated  forms  are 
plentiful  and  bear  witness  of  a  long  history.  For  the  single  termite  an  Antarctic 
ancestry  is  postulated.  Most  of  the  butterflies  collected  have  not  yet  been  de- 
scribed. Diptera  are  numerous  and  largely  allied  to  American  genera  or  species, 
and  the  non-endemic  forms  mostly  Chilean.  I  have  not  been  able  to  get  a  proper 
insight  into  the  distribution  of  the  many  genera  found  elsewhere.  Little  can  be 
said  about  the  beetles  until  Dr.  Kuschel's  material  has  been  described.  Two 
presumably  austral-bicentric  cases  are  noticed,  Pycnoinerodes  and  Eleusis.  As  in 
so  many  oceanic  islands  there  is  in  Juan  Fernandez  a  remarkable  display  of  endemic 
wingless  Curculionids,  living  on  the  endemic  plants  of  South  American  or  Antarctic 
parentage,  and  examples  of  strict  specialization  are  known.  Host  and  lodger  look 
back  upon  a  long  common  history,  but  whether  this  implies  a  common  original 
ancestral  home  or  adaptation  in  the  islands  I  cannot  tell.  Of  Hymenoptera,  Haplo- 
gonatus,  Prenolepis  and,  perhaps,  Metelia  show  Antarctic  connections. 

The  antiquity  of  the  endemic  land  shells  cannot  be  disputed.  The  Tornatellids 
are  an  ancient  Pacific  group  and  their  presence  in  Juan  Fernandez  as  well  as  in 
other  isolated  islands  and  archipelagoes  has  been  considered  a  proof  of  former 
land  connections.  Their  display  in  Hawaii  is  unparalleled,  Germain  summarized 
his  opinion  on  the  evolution  of  the  Hawaiian  fauna  in  the  following  terms  which, 
mutatis  mutandis,  apply  also  to  Juan  P^ernandez  [lOj.  995). 

S'il  est  bien  ainsi,  cette  famille  primordiale  [the  ancestors  of  Achatinellidae,  Amas- 
tridae,  Leptachatinidae  and  Tornatellinidae]  doit  avoir  une  tres  ancienne  origine  et 
remonter  au  Paleozoique.  .  .  .  le  peuplement  malacologique  de  I'archipel  des  Hawaii 
est  fort  ancien  et  doit  remonter  a  des  temps  primaires.  II  n'a  put  se  faire,  comme  le 
preuvent  les  developpements  precedents  sur  la  repartition  des  genres  et  des  especes, 
que  sur  une  aire  contenue,  ce  qui  exclut  la  possibilite  de  considerer  les  iles  Hawaii 
comme  le  resultat  de  I'activite  des  volcans  sous-marins. 

26  —  557857    The  Nat.  Hist,  of  Juan  Fernandez  and  EaJter  Isl.    Vol.  I 


4o: 


C.   SKOTTSBERG 


Finally,  let  us  try  to  reconstruct  the  history  of  our  islands,  beginning  with 
the  time  when  there  existed  a  "Tierra  de  Juan  Fernandez"  in  BrCggen's  sense. 
It  must  have  become  isolated  and  reduced  in  size  rather  early.  We  do  not  know 
if  the  fauna  included  vertebrates  other  than  birds;  if  it  did  they  did  not  survive 
the  long  volcanic  period.  Unfortunately  we  know  too  little  of  their  history  in 
Chile,  when  they  hrst  appeared  in  modern  forms,  and  so  forth.  The  absence  of 
all  gymnosperms  is  difficult  to  explain.  The  Eocene  flora  of  Chile  contained  several, 
Araucaria,  Libocedrus  and  Podocarpus,  all  still  living  there  and  accompanied  by 
I'itzrova,  SaxcgotJiaca,  Pilgerodoidroji  and  Dacrydhim,  and  even  if  no  close  land 
connection  existed,  some  of  them  ought  not  to  have  had  much  greater  difficulties 
to  get  transported  across  the  water  barrier  than  some  of  the  angiosperms  found 
on  the  islands.  If,  on  the  other  hand,  a  land  bridge  existed,  I  can  see  no  obvious 
reason  why  conifers  did  not  use  it  or,  if  they  did,  why  they  didn't  take  possession 
of  the  new  volcanic  soil.  Introduced  araucarias,  pines  and  cypresses  do  well  on 
Masatierra.  The  only  possibility,  remote  perhaps,  could  be  that  they  had  not  been 
able  to  spread  as  far  as  to  Juan  Fernandez  when  the  connection  was  cut.  Their 
absence  gives  us  no  clue  to  the  time  when  this  happened. 

Another  element  in  the  Chilean  flora  would  seem  to  come  to  our  rescue,  the 
XotJiofagiis  flora.  In  my  sketch  of  1925  [2JI.  33)  I  expressed  the  idea  that  "the 
connections  between  the  islands  and  the  main  land  were  severed  before  the  south 
Chilean  flora  assumed  its  present  composition,  and  also  before  the  advancing 
XotJiofiii^us  flora  reached  these  latitudes",  and  BrCcgen  is  of  the  same  opinion. 
He  states  that  tropical  South  America  is  the  ancestral  home  of  much  of  the 
Chilean   forest  flora,  and  adds: 

Pero,  la  actual  flora  de  Chile  central  contiene,  ademas,  una  mezcia  con  una  flora 
dc  (lima  mas  fresco,  (jue  despues  de  la  separaci6n  de  la  Tierra  de  Juan  Fernandez 
inmif^ro   y   (jue  se  caracteriza  por  los  generos  Nothofagus,  Araucaria,   etc. 

Araiicaria  was  found  in  the  Pichileufu  beds  which  are  supposed  to  be  Eocene, 
whereas  the  southern  beeches  appear  in  this  latitude  considerably  later.  In  the 
Magellanian  region  Dl  SKN,  as  we  have  seen,  distinguished  two  horizons,  a  lower, 
Oligocene,  with  Xoiliofagus,  and  an  upper.  Lower  Miocene,  \m\\\\  Araucaria.  Accord- 
ing t(j  Hi<i'(;(;kn  the  separation  of  Juan  Fernandez  from  Chile  took  place  during 
the  Oligocene  Xavidad  transgression.  We  would  think  that,  if  the  bridge  lasted 
longer,  the  XotJiofagus  flora  ought  to  have  invaded  Juan  Fernandez  and  to  have 
found  suitable  stations  in  the  montane  belt,  and  that  the  rising  volcanic  islands 
offered  acceptable  habitats  for  a  genus  of  such  wide  physiological  amplitude. 

Anyhow,  during  the  final  upheaval  of  the  Andes  when,  in  any  case,  most 
of  the  land  disappeared,  leaving  the  submarine  ridge  with  the  rising  volcanic 
masses  standing.  Central  Chile  definitely  ceased  to  be  a  source  of  the  island  flora. 
This  Chilean  flora  is,  as  Reiciie  pointed  out,  a  product  of  the  Andes  with  addition 
of  an  Antarctic  element. 

Submergence  resulted  in  the  displacement  of  the  vegetation  belts  and  in  much 
loss  of  life  when  the  volcanic  eruptions  began  and  lava  flows  and  other  ejected 
material  covered  part  of  the  country.   Long  before  that,   important  changes  had 


DERIVATION  OF  THE  FLORA  AND  FAUNA  403 

taken  place  as  a  consequence  of  the  cooling  of  the  chmate  in  post-Eocene  time, 
and  with  the  widening  space  of  sea  between  the  islands  and  the  mainland  the 
insular  climate  became  more  and  more  oceanic. 

The  rise  of  the  Cordillera  was  an  affair  of  great  magnitude  and  long  duration, 
and  millions  of  years  passed  before  the  last  remnant  of  the  Juan  P^ernandez- 
Desventuradas  ridge  disappeared.  Two  eruption  centres  were  formed,  Masatierra  + 
Santa  Clara,  and  Masafuera.  The  first  two  form  an  arc,  suggesting  a  remnant  of 
a  gigantic  crater,  but  the  place  must  be  properly  sounded  in  order  to  allow  us  to 
prove  or  disprove  this  hypothesis.  I  cannot  even  guess  where  the  eruption  centre 
of  Masafuera  should  be  looked  for.  Abrasion  has,  as  could  be  expected,  forced 
the  coast  escarpment  back  more  on  the  west  than  on  the  east  side,  reducing  the 
area  of  the  island.  A  description  of  the  topography  of  the  islands  is  found  in  this 
volume,  pp.  89-168.  There  is  no  sign  of  recent  activity  anywhere,  no  parasitic 
cones,  no  hot  springs.  When  the  map  in  "Der  neue  Brockhaus"  (1938)  marks 
Juan  Fernandez  as  a  seat  of  active  volcanoes,  this  applies,  I  suppose,  to  the 
submarine  eruption  in  1835  near  Masatierra  and  to  the  phenomena  observed  at 
San  Felix  some  years  ago  in  connection  with  an  earthquake  on  the  mainland, 
but  otherwise  nothing  in  the  way  of  activity  has  been  recorded  after  the  tsunami 
in  I75i>  when  also  an  earthquake  occurred,  and  other  earthquakes  were  registered 
in   1809,    1822   and   1835  (27 j). 

As  I  said,  I  believe  that  Masatierra  and  Masafuera  represent  two  separate 
centres;  the  distance  between  them  is  92  nautical  miles  and  the  sea  is  deep. 
Whether  they  are  of  exactly  the  same  age  and  became  extinct  at  the  same  time 
is  difficult  to  tell;  to  judge  by  the  topography,  Masafuera  makes  an  impression 
of  being  much  less  eroded,  and,  as  a  consequence,  younger,  but  the  difference  is, 
I  think,  mainly  due  to  differences  in  the  basalt;  the  petrographical  structure  is 
not  quite  the  same.  It  is  supposed  that  the  eruptions  began  during  Pliocene  and 
lasted  a  very  long  time  and  that  part  of  the  submarine  ridge  was  still  above  water 
when  the  main  eruption  centres  became  extinct.  Otherwise  the  result  would  have 
been  two  lifeless  islands  without  a  sign  of  the  old  endemic  biota.  The  flora  and 
fauna  were  inherited  from  the  sinking  land.  The  process  is  easily  observed  in 
many  volcanic  islands  and  nowhere  to  greater  advantage  than  in  the  island  of 
Hawaii.  As  soon  as  the  lava  has  cooled  down,  plants  get  established,  microscopic 
algae,  modest  tufts  of  mosses  and  particularly  lichens  of  the  genus  Stereocmilon 
[34-1),  but  a  conditio  sine  qua  7wn  is  that  moisture  is  available,  that  erosion  sets 
in  and  soil  is  formed.  Even  under  very  favourable  conditions  and  with  the 
sources  for  repopulation  next  door,  it  will  take  a  long  time  before  a  closed  vege- 
tation cover  gets  established.  With  regard  to  Juan  Fernandez,  nothing  much 
could  happen  until  the  islands  had  been  built  up  to  a  considerable  altitude, 
undoubtedly  greater  than  now,  when  streams  rushed  down  the  mountain  slopes 
and  started  to  excavate  valleys,  for  no  plant  cover,  not  to  speak  of  forest  growth, 
could  get  established  until  erosion  and  abrasion  had  done  part  of  their  w^ork. 
This  means  that  a  good  deal  of  the  fundament,  now  hundreds  of  metres  below 
the  surface  of  the  ocean,  was  still  exposed  and  retained  a  portion  of  the  original 
flora  and  fauna,  which  became  the  principal  source  of  the  flora  and  fauna  of  the 


404 


C.  SKOTTSBERG 


volcanic  soil.  It  goes  without  saying  that  chance  played  a  dominant  role  and  that 
the  fragmentary  character  of  the  island  world  is  easy  to  understand.  It  is  also 
possible  to  understand  why  different  species  happened  to  become  isolated  on 
Masatierra  and  Masafuera.  According  to  Joiiow  Masafuera  was  populated  through 
overseas  transport  from  Masatierra,  and  this  explained  why  the  former  was  so 
much  poorer,  but  only  50%  of  the  vascular  plants  found  on  Masafuera  were  known 
to  him.  Other  reasons  for  the  dissimilarities  between  the  islands  are  differences 
in  the  topography,  particularly  in  altitude,  Masatierra  is  915,  Masafuera  1570  m 
high.  If  we  could  lower  Masafuera  650  metres,  the  entire  highland  region  wnth 
its  special  flora  and  fauna  would  disappear.  The  question  of  the  origin  of  the 
alpine  flora  is,  as  we  have  seen,  difficult  to  answer.  Did  it  exist  in  the  islands 
before  the  separation  from  Chile  took  place,  and  were  there  any  habitats  where 
it  could  thrive.-  Many  of  the  species  are  of  Antarctic  origin  and  immigrated  to 
the  extreme  south  of  America,  with  or  perhaps  after  the  NoiJiofagus  flora,  where 
they  abound  in  the  bogs  of  the  rainy  zone  and  in  the  mountains  above  the 
timberline.  And  if,  as  was  explained  above,  the  NotJiofagus  flora  never  had  an 
opportunity  to  spread  to  Juan  Fernandez  over  land,  nor  were  those  Magellanian 
plants  able  to  come.  There  are  several  montane  plants  in  Masafuera  which  un- 
doubtedly date  back  to  early  times,  but  they  are  of  different  origin:  the  species 
of  liriireron,  Euphrasia  fonnosissi))ia,  McgalacJme  masafueraiia,  PJwenicoseris  regia^ 
]\anu7iculus  capranuH  and  Robbisoiiia  Masafuerae,  all  of  them  peculiar  endemics, 
and  of  these  Ranuuculus  and  JMegalacJine  are  found  only  along  the  high  ridge 
above  1300  m.  luen  if  some  of  them  are  of  Antarctic  ancestry,  their  history  is 
another.  When  it  comes  to  the  Magellanian  group,  Orcoboliis,  LageiiopJiora^ 
(ileichcnia,  and  so  forth,  it  is  difficult  to  exclude  the  possibility  that  they  be 
glacial  or  postglacial  immigrants.  Carcx  Baiiksii,  limpeirnni  ruhruni — only  a  single 
plant  seen  —  (niliuui  juasafueranimi,  (iiiapJialiujii  spiciforinc,  Myrtcola  yiuDiDiulai'ia 
and  Rubus  gcoides  are,  as  was  already  told,  not  of  Antarctic  origin,  but  may  have 
extended  far  south  after  the  recession  of  the  ice  and  accompanied  Orcobohis,  etc. 
And  we  cannot  refuse  to  admit  that  various  Chilean  ferns,  bryophytes  and  lichens, 
belonging  to  the  forest,  were  transported  across  from  South  Chile,  for  even  if  the 
{:)revailing  winds  are  westerly,  storms  from  other  directions  occur,  and  there  has 
been  j^ilenty  time.  .Still,  the  floristic  difference  between  the  two  islands  is  a  warning, 
not  to  put  too  much  faith  in  the  efficiency  of  the  natural  agents,  among  which, 
in   this  case,   the   wind  stands   foremost. 

The  biological  differences  between  Juan  Fernandez  and  the  mainland  increased 
during  the  Ice  Age.  West  Patagonia  and  l^\iegia  were  covered  by  inland  ice  [idf); 
if  small,  ice-free  refuges  occurred  has  not  been  definitely  stated,  but  is  not  alto- 
getlier  improbable;  their  part  taken  in  the  repopulation  of  the  country  was,  however, 
of  minor  importance.  With  the  north-south  trend  of  the  Cordillera,  the  road  toward 
north  lay  open,  and  the  big  island  of  Chiloe  was  not  ice-covered,  allowing  the 
subantarctic  fiora  and  fauna  to  survive,  perhaps  also  some  of  the  hardy  trees  and 
shrubs.  The  high  crests  of  the  Andes  in  Central  Chile  were  covered  by  glaciers 
descending  into  the  valleys;  scjueezed  between  the  mountains  and  the  coast  a 
migration  back  and   forth   went  on   during  the  successive  interglacial  and  glacial 


DERIVATION  OF  THE  FLORA  AND  FAUNA  405 

periods.  At  times  the  subantarctic  bog  and  heath  must  have  occupied  considerable 
areas  north  of  their  present  range,  but  during  all  this  shifting  to  and  fro  many 
species  may  have  been  lost,  some  of  them  surviving  on  Masafuera.  The  glaciers 
did  not  come  down  to  the  coast  of  Central  Chile,  but  the  climate  was  cold  and 
numerous  ancient  stenotopic  types,  some  of  them  surviving  on  Juan  Fernandez, 
perished. 

The  influence  of  the  glacial  periods  cannot  have  been  very  destructive  on 
the  islands.  Some  plant  species  reduced  their  range  and  became  rare  or  extinct; 
several,  apparently  with  a  very  narrow  physiological  amplitude,  are  on  the  verge 
of  extinction  today.  In  Masafuera,  the  timberline  was  lowered,  I  suppose,  and  the 
forest  patches  in  the  valleys  came  down  toward  the  sea.  Masatierra  has  no  climatic 
upper  timberline.  In  a  depression  on  the  summit  of  the  highest  peak.  El  Yunque, 
Driniys,  JiiaJiia,  Cuminia,  Escallonia,  Rhethiodendron,  Dicksonia,  etc.  etc.,  luxuriate 
more  than  anywhere  else,  thanks  to  the  constant  humidity. 


Part  2. 
EASTER  ISLAND. 

Chapter  XIV. 

Composition,  distribution  and  relationships  of  the  Flora. 

I.  Angiospermae. 
Gramineae. 

PaspaluDi   L.   More  than   200;   trop.   to  temp.,  most  numerous  in  Amer. 

forstcria)un)i  Muegge.   X.   Caled. 

scrobiculatnui   L.   var.  orbicitlare  (Forst.)  Domin.  X.  Guin.,  Austral.,  N.  Caled. ^ 
Polyn. 
Axouopus  l^eauv.  About  75,   the  majority  in   X.   and  S.  Amer. 

paschalis    Pilger.    Related,    according    to    I^ILGKR,    to  A.  scoparius  (Fluegge) 
Pilger,  a  S.   American  species. 
Siipa  L.i 

horridula  Pilger.   Related  species  in   S.   Amer.  and  Austral. 
Sporoboliis  R.   Hr.   Over   lOO,  trop.-subtrop.,   most  numerous  in  Amer. 

elou(^atus    R.    Br.    [as   indicus  (L.)  R.  Br.  in  2jo\.  S.  As.,  Malays.,  Austral., 
X.   Zeal. 
Calauiai:;rostis  Adans.  About   150,   mostly  temp.,   north  and  south. 

rctyofracta  (Willd.)   Link.  (Agrostis  W'illd.,  A.  filitbrmis  (Forst.)  Spreng.)  Aus- 
tral., 'i'asm.,  X.  Zeal.,   Polyn.,   Hawaii. 
DichclacJuic  ICndl.   2,  Austral.,  X.  Zeal. 

sc/Nna  (R.   \^v.)   Hook.   fil.   Austral.,   X.   Zeal. 
nauthojiia   DC. 

paschalis  Pilger.   Perhaps  most  nearly  related  to  /).  cJiilciisis  13esv. 
I'j-(li^ras/is  Host.   ()\er  200,  trop.-subtrop. 

i-loiigata  Jacfj.   \\.   Ind.   to   Malays,   and   Polyn. 

Cyperaceae. 

C)pcri(s  L. 

cragrostis  Lam.   \^ery  likely  introduced,   and   this  may  be  the  case  also  with 
the  other  species. 

^  The  area  of  genera  and  species  also  found  in  Juan  Fernandez  was  indicated  in  Pt.  i  and 
is   not  repeated  liere. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  407 

polystachyus  Rottb.  Trop.-subtrop.,  wide-spread. 

cylindrostachys  Boeck.  As  the  former. 

brevifolhis  (Rottb.)  Hassk.  Pantrop. 
Scirpus  L. 

ripariiis  Presl.  N.  Amer.,  Calif,  to  S.  Amer.,  south  to  Fueg.,  Falkland.  I  can- 
not find  that  var.  paschalis  Kiiekenth.  deserves  to  be  distinguished. 

Juncaceae. 

J  uncus  L. 

plebeius  R.  Br.  Colomb.-Urug.,  Austral,  Tasm.,  N.  Zeal. 

Piperaceae. 

Peperomia  Ruiz  et  Pav. 

rejlexa    Dietr.    Pantrop.,    very    widespread,    north    to    Hawaii    and    south    to 
N.  Zeal.;  also  on  Rapa  and  possibly  Pitcairn. 

Chenopodiaceae. 

Chenopodiiim  ambiguiun  R.  Br.  Austral.,  Tasm.,  N.  Zeal. 

Polygonaceae. 

Polygo7imn  L.  About   150,  world-wide. 

acuminatum    H.  B.  K.    W.    Ind.,    Centr.    and    S.  Amer.,  Galap.  Is.,  trop.  and 
S.  Afr.,  Orient. 

Nyctaginiaceae. 

Boerhaavia  L.  About  20.  Pantrop. 

diffusa    L.    Widespread  in  the  Pacific  and  a  common  weed  in  the  Old  and 
New  World;   probably  of  aboriginal  introduction. 

Aizoaceae. 

Tetragonia  L. 

expansa  Murr. 

Cruciferae. 

Nasturtium  R.  Br.   About  50.  Widespread,  mostly  temp. 
sarmentosum  (Sol.)  O.  E.  Sch,  Austral.-Polyn.,  Hawaii. 

Leguminosae. 

Caesalpiiiia  L.  At  least  90-100;  pantrop. 

bonduc  (L.)  Roxb.  Trop.  As.-Polyn. 
Soph  or  a  L. 

toromiro  (Phil.)  Skottsb.   Nearly  related  to  the  species  from  J.  Fern. 

Euphorbiaceae. 

Euphorbia  L.    1 500-1600;  world-wide. 

hirta  L.  Indomal.-Polyn.,  often  adventitious. 

serpe7is  L.  As  the  former,  also  common  in  trop.  Amer. 


408  C.   SKOTTSBERG 

Umbelliferae. 

ApiuDi  L. 

prostration  Labill.   Austral-circuiii[).,   incl.  A.  atistrale  Thouars. 

Primulaceae. 

Satiiolus  L.  9  (12?),    I    cosmop..  2  (5?)  X.   Amer.,   3  southern  S.  Amer.,  i  S.  Afr., 
I    W.  Austral.,  and  tlie  following. 

repejis  (Forst.)  Pers.   Austral  circump. 

Gentianaceae. 

Erythraea  Neck.  (Centaurium   Hill.)  30-40,  subtrop.-temp. 
australis  R.   Br.   Austral.,   \.   Caled.,  Fiji. 

Convolvulaceae. 

Calystegia  R.  l^r.   7-8,  temp.-subtrop. 

stpiuin   (L.)  R.   Br.  forma.  All  continents,  also  reported  from  Australia,  but 
possibly  introduced   in  the  s.   hemisph. 
Ipoiuaca  L. 

pes  caprae  (L.)  Roth.  Pantrop. 

Solanaceae. 

LyciuDi   L.  About   100,  most  numerous  in   S.  Amer. 

carolhiiamiui  Walt.  var.  scDuiuiccnsc  (Gray)  L.  C.   Hitchc.  Rapa,  Hawaii. 

The  flora  is  extremely  poor,  not  much  richer  than  in  the  low  coral  islands: 
16  families,  26  genera  and  31  species,  and  I  am  not  at  all  sure  that  all  of  them 
are  indigenous  and  did  exist  here  before  man  appeared  on  the  scene;  some  may 
have  been  accidentally  or  purposely  introduced  by  the  aborigines,  by  the  American 
whalers  and  in  modern  times.  With  regard  to  the  "endemic"  Solamini  Insulae 
PascJialis  Bitter,  see  2^c)\  it  was  used  as  medicine.  P'our  species  are  endemic 
(13  %),  but  three  of  them  belong  to  large  grass  genera  needing  monographic  study. 
FoRSiKK  mentions  {347),  beside  some  cultigens,  only  Panictmi  filifonue  Jacq. 
(-- Digitaria  sanguinalis),  a  common  weed,  SJicfficldia  (  =  Samolus)  rcpens,  Aveim 
filiformis  ( -- Calamagrostis  retrofracta)  and  Solanioii  nigruvi;  in  his  journal  (j^<?) 
he  refers  to  Apiuni^  which  he  knew  from  New-  Zealand,  to  ''JMivwsa'  (  =  Sophora), 
and  also  to  the  former  occurrence  of  sandalnwod.  In  regard  to  the  distribution 
of  SiDitalion  in  Southeastern  l\)lynesia,  where  S.  hisulare  Bert.,  including  varieties, 
is  known  from  Tahiti,  Kaiatea,  Marquesas,  and  Austral  Is.,  and  5.  li€ndersone7isc 
F.  B.  R.  Brown  (very  chxse  to  the  former)  is  found  on  Henderson  Island  in  the 
extreme  south-east  of  the  island  swarm,  it  did  not  seem  incredible  that  sandal- 
wood once  occurred  on  l^Laster  Island,  perhaps  introduced  by  the  aborigines;  not 
very  long  ago  S.  yasi  Seem,  was  introduced  to  Tonga  and  is  well  established  there. 
The  matter  is  sufficiently  interesting  to  be  discussed  here. 

Tradition  tells  that  when  Hotu  Matua,  the  legendary  hero  of  the  liaster 
islanders,  took  possession  of  the  island,  he  brought  various  useful  trees  and  other 
plants,    a    story    first    told    by    Fokster    (,?^<^'.  583).    Mhtraix   [180.  i^-xy)    enu- 


DERIVATION  OF  THE  FLORA  AND  FAUNA  409 

merates  mahute  [Brotissonetia  papyrifera),  makoi  [Thcspcsia  populnea),  hauhau 
[Triumfetta  semitfiloba),  toromiro  [Sophora),  naunau  (Saiiialum),  and  marikuru 
[Sapijtdus  saponaria).  None  are  of  American  origin;  all  point  towards  Malaysia- 
Polynesia. 

In  his  description  of  a  beautiful  wooden  hand  presented  to  him  by  one  of  the 
natives  and  now  in  the  British  Museum,  FORSTER  says  that  "the  wood  of  which 
it  was  made  was  the  rare  perfumed  wood  of  Taheitee,  with  the  chips  of  which 
they  communicate  fragrance  to  their  oils",  undoubtedly  sandalwood,  with  which 
FORSTER  was  familiar.  The  natives  were  expert  in  the  art  of  wood-carving,  but 
the  hand  is  very  unlike  all  other  objects,  which  were  made  of  Sophora  wood, 
called  toromiro,  while  the  name  of  the  tree  with  the  fragrant  wood  was  naunau. 
Cooke  {34.3.  722)  translated  this  word  "bastard  sandalwood".  This  is  the  name 
given  to  Myoporum  sandvicense  Gray  in  Hawaii,  where  the  native  name  is  naio 
(naeo,  naieo);  Santalmn  is  called  iliaJii,  in  Marquesas  puahi,  on  Rapa  eahi,  in 
Tahiti  and  Tonga  ahi.  Whether  or  not  the  words  naunau  and  naio  have  the 
same  base  I  cannot  tell;  on  Raivavae,  Rapa  and  New  Zealand  the  name  of 
Myoporum  is  ngaio,  whereas  true  sandalwood,  in  this  case  Mida,  is  called 
maire,  the  Hawaiian  name  for  the  fragrant  Alyxia  oliviformis  Gaud.,  transferred 
to  Mida  in  New  Zealand,  where  no  Alyxia  occurs. 

Our  main  sources  of  Easter  Island  ethnology,  Roggeveen,  La  Perouse, 
Cook,  Mrs.  Routledge,  etc.  do  not  mention  sandalwood,  while  Metraux  pays 
special    attention   to  this  subject,  referring  to  FORSTER  {180.  17-18). 

My  informant  gave  "sandal"  as  the  Spanish  equivalent  of  the  word  naunau — 
the  correct  Spanish  word  is  sdndalo — and  remarked  that  the  tree  had  entirely  disap- 
peared since  the  time  of  Salmon  (between  1880  and  1890).  The  last  one  on  the  island 
grew  near  Vai-mata,  but  died  recently  "because  there  were  no  more  kings".  From  the 
distribution  of  the  sandalwood  it  seems  likely  that  the  naunau  was  the  true  sandal- 
wood and  not  the  bastard.  Sandalwood  "is  found  on  the  atolls  of  Elizabeth  and  Ducie, 
the  nearest  islands  to  Easter  Island". 

Elizabeth  I.  is  another  name  for  Henderson,  a  rocky  coral  island;  Ducie  is 
not  quoted  by  BROWN  [33.  ill).  To  judge  from  the  distribution,  jiaunau  could 
just  as  well  refer  to  Myoporum,  a  genus  represented  by  endemic  species  in  Rapa 
— where,  as  we  shall  learn  presently,  7mu  means  something  quite  different — 
and  Austral  Is.  but  never  reported  from  Easter  Island.  The  wood,  slightly  fra- 
grant, is  said  sometimes  to  have  been  used  as  a  substitute  for  sandalwood  in 
Hawaii.  Metraux's  informant  seems  to  have  been  convinced  of  the  earlier  ex- 
istence of  real  Santalum  on  his  island.  If  so,  and  if  it  had  been  brought  from 
Tahiti,  it  ought  to  have  had  the  same  name. 

The  story  does  not  end  here.  One  of  my  correspondents,  Mr.  Paul  H.  Steele 
of  Sacramento,  Cal.,  kindly  called  my  attention  to  a  book  written  by  Padre 
Sebastl\N  Englert,  who  has  lived  more  than  1 5  years  on  Easter  Island  [86], 
a  place  in  which  Mr.  Steele  is  particularly  interested.  As  I  had  no  opportunity 
to  consult  the  book,  Mr.  STEELE  copied  and  sent  me  the  following  remarkable 
passage. 


^lO  C.   SKOTTSBERG 

yau  o  Naunaii  (Santaliim):  arbiisto  de  la  familia  de  las  santaldceas,  tambien  11a- 
mado  nan  opata,  poniue  creci'a  en  los  barrancos  (opata)  de  la  costa,  entre  rocas  y 
I)iedras.  Ahora  ha  desaparecido.  I.os  ultimos  ejemplares  (jiie  algunos  de  los  natives 
actuales  recuerdan  haber  visto  todavi'a,  se  ban  secado  hace  unos  50  anos.  El  nau  opata 
daba,  como  friitos,  niieces  del  tamano  de  castafias,  los  "mako'i  nau  opata".  Carl 
Friedrich  Behrens  nombra  nueces  entre  los  friitos  que  los  islenos  les  regalaron  en  gran 
numero  a  el  y  a  sus  compaiieros.  Hotii  Matiia  y  su  gente  parecen  haber  trafdo  gran 
cantidad  de  cstas  nueces,  i)orciue  de  ellas  se  alimentaron  en  los  primeros  meses  des- 
pues  de  haber  llegado  a  la  isla.  Al  excavar  la  tierra  en  cuevas  (jue  estaban  antigua- 
niente  habitadas  se  encuentran  cascaras  de  estas  nueces.  Estas  cascaras  generalmente 
no  cstan  ([uebradas  sino  ([ue  han  sido  abiertas  en  forma  de  un  pecjueno  circulo,  para 
ser  usadas  por  los  ninos  en  el  juego  del  trompo.  La  madera  del  arbusto  se  utilizaba, 
j)or  su  cx([uisito  aroma,  para  confeccionar  un  perfume,  como  los  veremos  en  otro  capitulo. 

This  description  does  not  at  all  fit  either  Santaluin  or  Myoportwi.  The  fruit 
of  SaiitaluDi  is  an  ellipsoid  drupe  with  a  thin  fleshy  mesocarp  and  a  very  hard 
endocarp,  and  I  have  never  seen  or  heard  of  a  kind  the  size  of  a  castafia  (chest- 
nut); in  the  largest  I  have  measured  (S.  pyrularhan  Gray)  the  drupe  was  16-18  mm 
long  and  the  stone  12-15  mm.  According  to  HlLLEBRANl)  (joy.  390)  the  drupe 
measures  up  to  24  mm  in  length,  but  I  have  not  seen  any  as  large  as  that.  Nor 
have  I  ever  heard  that  the  kernel  is  used  as  food;  the  idea  that  Hotu  Matua's 
party  could  have  maintained  itself  for  months  on  nothing  else  is  preposterous, 
and  I  fail  to  see  how  the  stone  could  be  used  as  a  whipping-top.  The  Jiatinau  which 
grew  along  the  coast  and  produced  the  "cascaras"  found  in  the  caves  cannot 
have  been  a  species  of  Sajitaluui.  Fortunately  this  could  be  proved.  Mr.  STEELE 
had  told  me  that  Father  FLnglert  had  sent  him  two  shells  for  his  collection 
of  l^aster  Lsland  curios  and  had  promised  him  more,  of  which  he  intended  to 
send  me  samples.  As  time  went  by  and  no  more  came  I  asked  him  the 
favour  of  sending  me  one  of  his  precious  specimens  as  loan,  and  he  willingly 
consented.  It  is  a  hard,  brown  and  smooth,  almost  globular  shell,  2.5  cm  high, 
3  cm  wide,  2  mm  thick,  with  a  large  irregular  hole  in  the  basal  part.  It  has 
nothing  whatever  to  do  with  Saiitaliini.  A  passage  in  Me'I'RAUX's  book,  p.  353, 
j)ut  me  on  the  track.  He  quotes  a  song  which  the  children  used  to  sing  when 
the  tops  were  spinning,  and  it  tells  that  the  spinning-tops  were  made  of  viakoi 
—  'riicspesia  popiibiea  cajxsules!  A  comparison  with  herbarium  material  showed 
that  Mr.  Steele's  specimen  is  a  typical  capsule  of  yy/r.sy^ri-/"^,  one  of  the  "nuts" 
found  in  the  caves.  They  made  \ery  poor  food  but  good  spinning-tops.  We 
did  not  see  Thcspesia  on  the  island,  but  Metrai  X  observed  it  growing  on  the 
cliffs  at  Poike  (the  eastern  headland).  F^vidently  the  word  nau  or  nauuau  has 
been  altogether  misplaced  by  the  blaster  islanders,  although  both  Metraux 
and  bLxcLERi-  were  told  that  it  was  the  name  of  Santaluni.  We  find  the  same 
word  in  Tahiti  and  the  Tuamotu  Islands  for  Lcpidiu))i  bide7itaiuni  Montin,  and 
this  is  called  iiaupata,  strikingly  like  FLxglert's  }iau  opata,  in  Marquesas;  in 
Tahiti  naupata  means  Scaci'ola  frutcscens,  which  is  called  naupaka  in  Hawaii  and 
}ii^ai{}ii!;au  in  Rarotonga,  and  on  Rapa  }iau  is  used  for  So)icJius  oleracens.  All 
these  {plants  were  used  as  medicine. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  4 II 

Geographical  elements. 

In  1934  I  made  an  attempt  to  arrange  the  vascular  plants  according  to  their 
distribution,  with  the  following  result  (2jg.  278-279,  translated  from  French). 

1.  Australian-Polynesian   element  (12). 

a)  Species    found    elsewhere:    Paspalum  (2),  Sporobolus,  Calamagrostis,  Dichelachne, 
Eragrostis,  Juncus,   Peperomia,   Chenopodium,  Nasturtium,  Erythraea. 

b)  Endemic:   Stipa  horridula. 

2.  Palaeantarctic  element  (3). 

a)  Species  found  elsewhere:   Apium,   Samolus. 

b)  Endemic:   Sophora  toromiro. 

3.  American  element  (6). 

a)  Species  found  elsewhere:   Cyperus  vegetus,   Scirpus,   Polygonum,  Lycium. 

b)  Endemic:   Axonopus  paschalis,   Danthonia  paschalis. 

4.  Wide-spread  tropical  element  (10):  Cyperus  (3),  Boerhaavia,  Tetragonia,  Caesalpinia, 

Euphorbia  (2),   Calystegia,  Ipomaea. 

Going  into  more  detail,  I  have  tried  to  rearrange  the  angiosperms  in  the 
following  manner. 

I.  Palaeotropical  element. —  22    sp.   (70%). 

1.  Pantropical  (6):  Cyperus  polystachyus,  cylindrostachys  and  brevifolius, 
Peperomia  reflexa,  Boerhaavia  diffusa,  Ipomaea  pes  caprae. 

2.  Malaysian- Polynesian  (9):  Paspalum  forsterianum  and  scrobiculatum,  Sporo- 
bolus elongatus,  Calamagrostis  retrofracta,  Eragrostis  elongata,  Nasturtium  sarmen- 
tosum,  Caesalpinia  bonduc,  Euphorbia  hirta  and  serpens. 

3.  Australian-Polynesian  (7): 

a.  Endemic:  Stipa  horridula  (.^). 

b.  Not  endemic:  Dichelachne  sciurea,  Chenopodium  ambiguum,  Tetragonia 
expansa,  Erythraea  australis,  Calystegia  sepium  (.'). 

c.  Also  in  tropical  America:  Juncus  plebeius. 

II.  Austral-circumpolar  element. — 4   sp.   (12.9%). 

a.  Endemic:  Danthonia  paschalis,  Sophora  toromiro. 
h.  Not  endemic:   Apium  prostratum,  Samolus  repens. 

III.  Neotropical  element. —  5    sp.   (16.1%). 

a.  Endemic:  Axonopus  paschalis. 

b.  Not  endemic:  Cyperus  eragrostis,  Scirpus  riparius,  Polygonum  acumina- 
tum,  Lycium  carolinianum  var. 

I  am  strongly  inclined  to  regard  Cyperus  eragrostis  as  a  late  immigrant  from 
Chile.  Lycium  carolinianum  var.  sandvicense  is  a  puzzling  case,  a  form  of  a 
North  American  species  but  only  found  in  Hawaii,  Rapa  and  Piaster  Island.  It 
is  a  halophyte  confined  to  the  beach,  but  one  might  expect  it  also  to  occur  on 
the  shores  of  California  etc.,  from  where  it  has  not  been  reported.  Beach  plants 


412  C.   SKOTTSBERG 

are  included  in  all  the  groups,  in  I  Ipouiaca,  Caesalpinia,  CJienopodiuni  and  Tetra- 
gonia,  in  II  Apiuiii  and  Sainolus,  in  III  Lycium\  together  they  are  7,  or  21%  of 
the  angiosperms. 

If  we  want  to  trace  an  ancient  Antarctic  source,  just  as  we  did  in  Juan 
Fernandez,  the  Antarcto-tertiary  group  would  consist  of  Daiitlionia,  Jwicus, 
SopJiora,  Apiuui  and  Sauiolus. 

To  judge  by  the  geographical  position  of  Easter  Island  the  dominance  of  a 
Palaeotropical-Pacific  element  was  expected.  The  island  certainly  is  remote  from 
the  large  cluster  of  islands  of  South-eastern  Polynesia,  but  much  more  distant 
from  South  America.  Consequently,  the  presence  of  a  neotropical  element  is 
surprising.  Future  researches  will  perhaps  show  that  Siipa  Jiorridula  should  be 
added,  which  makes  little  difference  as  long  as  the  endemic  Axonopus  is 
claimed  to  be  of  neotropical  parentage,  and  Scirpiis  riparius  and  Polygo7uan 
acuuiinatuin  remain  American.  Their  mode  of  occurrence  and  ecology  oblige  us  to 
regard  them  as  truly  indigenous,  unless  they  have  been  intentionally  introduced 
in  prehistoric  time  during  one  of  the  mythical  cruises  which,  according  to  Hever- 
DAHL,  put  Piaster  Island  in  contact  with  Peru.  A  direct  transport  of  seeds  across 
the  ocean  without  man's  assistance  is  difficult  to  imagine,  and  it  is  futile  to 
speculate  in  land  connections. 

II.  Pteridophyta  (^?,  2^g). 

Polypodiaceae  (in   the   old   sense). 
Aspleiiiuui  L. 

adiautoidcs  (L.)  C.  Chr.  var.  squamulosum  C.  Chr.  et  Skottsb.  The  species 
trop.  As.-Austral,  X.  Zeal,  and  Polyn.;  also  E.  Afr.  and  adjacent  islands;  the 
variety  endemic. 

obljqiiiDn  P^orst.  Austral-circump.;  the  only  plant  reported  from  Sala  y  Gomez. 

Dai'allia    Sm.    About    40;     wide-spread,    mainly    southern;    As.-Polyn.,    S.  Afr., 
Aladag.,  a  solitary  species  Ibero-Afr.   and  Macaron. 
solida  (Forst.)  Sw.   Indomal.-Polyn.-Austral. 

Doodia  R.  Pr.  12.  "New  Zealand  and  Juan  Fernandez  to  Hawaii  and  Australia" 
(COPELAM)  6g.  158),  but  "Juan   P^ernandez"   is  a  mistake  for  Easter  I. 

paschalis    C.    Chr.   et  Skottsb.   Close  to  /X  blechjwidcs  A.   Cunn.  (Austral.). 

Dryopteris  A  dans. 

Espinosai  Hicken.  l^elongs  to  a  neotropical  group. 
goigylodcs  (Schk.)   ().  K.   Pantrop. 
de)itata  (Forsk.)  C.   Chr.   Pantrop. 

ElapJioglossu))i  Schott. 

Skottsbergii  Krajina.   Related  to  Ji.  tahitensc  Brack. 

Microlcpia  Presl.  45-50;  a  pantrop.  genus  ranging  north  to  Japan  and  south  to 
Madag.  and  X.   Zeal. 

strigosa  (Thunb.)  Presl.  Indomal.-Polyn. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  413 

Polyp  odium  L. 

scolopendria  Burm.  Palaeotrop.,  wide-spread; 
Polystichuin  Roth. 

Fuentesii  Espinosa.  Belongs  to  the  P.  vestitum  assemblage, 
Vittaria  Sm.  About  80,  pantrop.-subtrop. 

elongata  Sw,  Indomal.— Polyn.-Austral. 

Ophioglossaceae. 

Ophioglossu7n  L. 

lusitanicum  L.  subsp.  coriaceum  (Cunn.)  Clausen.  Boliv.-Chile,  Austral.,  Tasm., 
N.  Caled.,  N.  Zeal. 

reticulaiwn  L.  S.E.  As.,  Philipp.,  N.  Guin.,  Melan.-Polyn.,  east  to  Mangareva; 
Mascarene  Is. 

Psilotaceae. 

Psilotum  Sw.   2   wide-spread  sp. 
jiudurn  (L.)  Griseb.  Pantrop. 

Fifteen  species  are  listed,  4  of  them  endemic  (26.6%),  the  endemic  variety 
of  Asplenium  adiantoides  not  counted. 

Geographical  elements. 

In  my  earlier  subdivision  {2jg)  the  following  groups  were  distinguished. 

1.  Australian-Polynesian  element  (6). 

a)  Species    found  elsewhere:  Asplenium  adiantoides,  Elaphoglossum  tahitense,  Micro- 
lepia  strigosa,  Vittaria  elongata,   Ophioglossum  coriaceum. 

b)  Endemic:   Doodia  paschalis. 

2.  Palaeantarctic  element  (i). 

Found  elsewhere:   Asplenium  obtusatum. 

3.  American   element  (i). 

Endemic:   Dryopteris  Espinosai. 

4.  Wide-spread  tropical  element  (4):   Dryopteris  gongylodes  and  parasitica  (=dentata), 

Polypodium  phymatodes,   Ophioglossum  reticulatum. 

The  list  contains  12  species;  two  have  been  reported  later  and  some  nomen- 
clatural  changes  have  been  made.  A  new  geographical  arrangement  follows  here. 

I.  Palaeotfopical  element. —  i  i  sp.    (733%). 

1.  Paiitropical  (3):  Dryopteris  gongylodes  and  dentata,  Psilotum  nudum. 

2.  Palaeotropical  (i):   Asplenium  adiantoides  (endemic  variety). 

3.  Malaysian- Poly 7iesian  (6). 

a.  Endemic:  Elaphoglossum  Skottsbergii. 

b.  Not  endemic:    Davallia   solida,  Microlepia  strigosa,  Polypodium  scolo- 
pendria, Vittaria  elongata,  Ophioglossum  reticulatum  (also  Mascarene  Is.). 

4.  Aiistralian-Polynesia7i  (i). 
Endemic:  Doodia  paschalis. 


414 


C.  SKOTTSBERG 


II.  Austral-circumpolar  element. — 3  sp.  (20%). 

a.  luidemic:  Polysticluim  Fuentesii. 

b.  Not  endemic:  Asj)leniiini  obliquum,  Ophioglossum  lusitanicum  subsp. 
coriaceum. 

III.  Neotropical  element. —  i  sj).  {6.7  %). 
luidemic:   Dryopteris  l^^.spinosai. 

We  find  the  same  dominance  of  a  palaeotropical  element  as  among^  the  an- 
(^iosperms.  OpJiioglossum  Iusita)iicu})i  is  reported  from  Atlantic  Europe,  the  Medi- 
terranean and  Macaronesia,  whereas  the  subspecies  is  austral-bicentric,  an  example 
of  a  remarkable  discontinuous  distribution.  The  American  element  is  represented 
by  a  single  endemic  species  and  there  is  no  fern  corresponding  to  Polygo7ium 
acuDiJiiaiuDi  and  Scirpus  ripariiis.  Polystichinn  Fuentesii  and  Asple7nwn  obliquiwi 
are,  })erhaps,  Antarctic,  and  the  latter  was  classified  as  such  above  (p.  282),  but  it 
is  a  seaside  plant.  If  we  follow  CoPEl.AXi),  Aspleimmi,  Doodia  and  Davallia  are  of 
Antarctic  origin. 

III.  Musci  (jj,  26^). 

I  have  to  thank  Dr.  HERMAN  Persson  for  kind  assistance  in  finding  out 
about  distribution. 

Treuiatodon  Michx  (Dicranaceae).   About  70,   mainly  trop.,  south  to  N.   Zeal. 
pascuanus  Ther. 

Campy lopus  l^rid. 

iutroflexiis  (Hedw.)  Mitt.   See  p.   227. 
turficola  Broth. 
hygrophilus  l^roth. 
dicranodontioides  Broth. 
saxicola  Broth. 

Fissi(/e7is  Med w . 

pascuanus  l^roth. 
PtycJiomiiriuDi  (Bruch)   I^'uernr. 

subcylindricutn  Thcr. 

JTeisia  Hedw.  (Trichostomaceae).  About  30,   widely  distributed  (luir.,  N.  Amer., 
K.  As.,  X.   Afr.). 

flavipes  Hook.   fil.   et  W'ils.  Java,   Ceylon,   E.  Austral.,  Tasm.,  N.   Zeal. 

Bvyu))i  Dill. 

a)\e;e)iteuin  L.  var.  laiiatuui  (Palis.)  Bryol.  eur.  Widely  spread  in  warmer 
countries,  the  species  cosmoj). 

PJiilonoiis  Brid. 

laxissima  (C.   M.)  Ikyol.  jav.   V..  Ind.,  Madag. 
Papillaria  C.   M.  (Meteoriaceae).  About  70;   a  pantrop.  genus. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  415 

pascuana  Ther.  ex  Broth.  Related  to  P.  crocea  (Hpe)  Jaeg.  Ind.,  Ceylon, 
Java,  Philipp.,  E.  Austral.,  N.  Zeal.,  Kermadec  Is.,  Fiji). 

Fabronia  Raddi  (Fabroniaceae).  90-100,  half  of  them  Amer.;  trop.-subtrop. 

macroblepharoides  Broth.  Related  to  F.  macroblepharis  Schwaegr.  (Brazil); 
other  related  sp.   in  Afr.  and  Austral. 

Rhacopilum  Palis. 

cuspidigeruni  Schwaegr.  Norfolk  I.,  N.  Caled.,  Samoa,  Hawaii. 

Besides,  Brotiierus  mentions    Weisia  sp.  and  Macromiirhini  sp. 

Fourteen  species  are  reported,  9  of  them  endemic  (64.3  %),  a  high  figure, 
but  there  can  be  no  doubt  that  only  a  part  of  the  mosses  has  been  collected, 
and  very  likely  the  proportion  of  endemics  will  be  reduced.  Of  greater  interest 
are  the  endemic  aquatic  species  of  Campylopus  of  the  crater  lakes,  possibly  relicts 
from  an  era  when  the  island  was  larger  and  higher  than  now  and  enjoyed  a  more 
humid  climate.  Probably  many  mosses  and  other  cryptogams  disappeared  with 
the  forest  groves.  Of  the  known  species  Campylopus  inlrojiexus,  Weisia  flavipes 
and  Rhacopilum  cuspidigerum  have  a  southern  distribution,  Philonoiis  laxissima 
is  palaeotropical  and  Bryum  argcntejim  cosmopolitan  and  perhaps  anthropochorous; 
the  endemic  species  are  supposed  to  have  tropical  relatives. 

IV.  Hepaticae  (/jo). 

Frulla7iia  Raddi. 

lageiiifera  Schwaegr.  Known  before  from  the  type  locality  only,  said  to  be 
the  Falkland  Is.,  but  this  statement  is  subject  to  doubt;  possibly  Ins.  Marianae 
is  meant  instead  of  Maclovianae. 

Conditions  are  not  favourable  to  hepatics,  but  more  species  will  be  found. 
A  Lejeunea   was    collected,    but   the   material  seems  to  have  been  lost  (I.e.  699). 

V.  Lichenes  (2^7). 

My  thanks  are  due  to  Dr.  A.  H.  Magnusson  and  Dr.  R.  Santesson,  who 
helped  with  information  on  the  distribution. 

Arthonia  Ach.  About  500,  the  majority  trop.-subtrop. 

fuscescens  Fee.  Apparently  only  reported  once  before  (trop.  Amer.). 
Opegrapha  Humb.  About  280,  mostly  warmer  climates. 

paschalis  Zbr. 
Grapliis  (Adans.)  Ach. 

lineola  Ach.  Trop.-subtrop. 
Diploschistes  Norm.  About  30,  cold  to  temp,  regions,  trop.  mountains. 

ariactinus  ILhx.  Described  from  Japan. 

scruposus  (L.)  Norm.  Wide-spread  N.  and  S.  temp. 
Heppia  Naeg.  About  40,  mostly  warm  and  dry  regions. 

Guepini  Nyl.  N.  Amer.,  Eur. 


41  6  C.  SKOTTSBERG 

Lecidea  (Ach.)  Th.   Fr. 
paschalis  Zbr. 

Cladoiiia  (Hill)  \'ainio. 

pityvea  (Flk.)  Fr.   Cosinop. 
Acarospora  ^lass. 

Skottsbergii  Zbr. 
rmDiclia  (Ach.)  De  Not. 

reticulata    Ta\'l.    \.    and    S.    Amer.,   W.   and   S.   luir.,  Afr.,  F>.   As.,   Austral. 

co)ispcrsa  (I^^hrh.)  Ach.   var.  lusitaiia  (Xyl.).   S.   Eur.;  the  species  cosmop. 

Usnea   Wigg. 

subtonilosa  (Zbr.)  Motyka  [344).  Masafuera.  Described  as   U.  Steineri  var.  by 
Zaiilhrl  (KNKR,  who  also  distinguished  var.  tincta  Zbr.   and  quoted  it  for  Easter 
Island;   this  is  called    l'.  inicta  by  MoTYKA,  who  records  it  for  S.  Amer.  only. 
Caloplaca  Th.   Vx. 

rub'Dia  Zbr.  J.   Fern. 

lucois  (Xyl.)  Zbr.  Tata*^.,  Falkl.,   S.  Georgia. 
Biullia  De  Not. 

stcllulata  (Tayl.)  Mudd.   Cosmop. 

fcvjiaiideziana  Zbr.  J.   Fern. 

Iialopliiloides  Zbr.   var.  The  typical  sp.  J.   Fern. 

paschalis  Zbr. 

glazjouajia  M.   Arg.   Brazil. 
R'niod'nia  (S.   Gray)  ^lass.  About  300;   very  widely  distributed. 

Perousii  Zbr. 
/V.ivV/r  (Fr.)  Xyl. 

entcroxantJia  Xyl.   forma.   S.W.   luir.,  Japan. 
Pliyscia  (Schreb.)  Vainio. 

picta  (Sw.)  Xyl.   Wide-spread  trop.-subtrop. 
Anaptycliia  Koerb. 

spcciosa  (W'ulf.)  Mass.   Widely  distributed;    in  Amer.  south   to  Fueg, 

These  23  sj)ecies.  5  regarded  as  endemic,  represent,  I  am  sure,  only  a  minor 
j)art  of  the  lichen  flora  and  do  not  lend  themselves  to  geographical  speculations. 
There  are  several  strange  cases  of  disjunction  serving,  J  daresay,  to  illustrate  our 
insufficient  knowledge  of  the  distribution   of  lichens. 

V.  Fungi. 

Our  collection  contained  a  single  s[)ecics,  lu)i>istella  piisilla  Eloyd,  known 
before   from   Australia  [102). 


DERIVATION  OF  THE  FLORA  AND  FAUNA  417 

Chapter  XV. 
Composition,  distribution  and  relationships  of  the  Fauna. 

Indigenous  vertebrates,  birds  excepted,  lacking.  The  principal  occupation  in 
the  island  is  farming,  cattle  and  sheep  are  plentiful  and  roam  over  the  island 
which,  with  the  exception  of  outlying  rocks,  has  lost  its  primitiveness.  As  a  con- 
sequence of  the  changes  in  the  plant  cover,  particularly  the  extermination  of  the 
indigenous  trees,  also  the  fauna  was  impoverished,  while  through  the  introduction 
of  useful  plants,  numerous  weeds  and  all  kinds  of  goods  many  foreign  insects 
and  other  invertebrates  made  their  appearance,  as  the  lists  below  will  show.  As 
little  research  work  has  been  done  hitherto,  many  more  species  will  probably  be 
found,  indigenous  as  well  as  introduced. 

Aves  (775). 

Sterna  lunata  Peale.  Molucc,  Polyn.,  Fiji,  Hawaii. 

Ajious  stolidus  (L.)  imicoior  Nordmann.  Sala  y  Gomez.  The  typical  species 
trop.-subtrop.,  but  not  observed  on  the  coast  of  America. 

Procelsterna  coerulea  (Benn.)  skottsber^gi  Loennb.  Typical  coerulea  on  Christmas 
I.,  4  other  subspecies  scattered  over  the  Pacific. 

Gygis  alba  (Sparrm.)  j'oyana  Matthews.  With  the  typical  species  wide-spread 
trop. 

Pterodroma  heraldica  Salvin  paschae  Loennb.  The  typical  species  S.W.  Pacific. 

Sula  cyanops  (Sundev.).  Trop.  seas  throughout  the  world. 

According  to  the  natives  some  other  sea  birds  occur,  but  there  are  no  land 
birds. 

Oligochaeta  [i8i). 

Pheretina  califoriiica  (Kbg).  Introduced.  Reported  from  Calif.,  Mex.,  Madeira 
and  Lower  Egypt. 

Araneida  {^22). 

Scytodes  liigtibris  Thorell.  Burma,  N.  Caled.,  perhaps  all  over  Oceania. 

Pholciis phalangioides  Fuessli.  Eur.,  now  spread  over  a  large  part  of  the  globe. 

Theridium  tepidariorum  C.  Koch.   Cosmop. 

+  Tetragnata  Paschae  Berland.  A  large  cosmop.  genus. 

Corinna  cetrata  Simon.  N.   Caled. 

Hasarius  Adansoni  Audouin.   Cosmop. 

Plexippus  Paykulli  Audouin.   Cosmop. 

Possibly  all  the  spiders  are  adventitious  (Berland  I.e.).  Two  species  were 
determined  as  to  genus  only. 

27  ~  557857  The  Nat.  His*,  of  Juan  Fernandez  and  Easter  Isl.  Vol.  I 


41  8  C.  SKOTTSBERG 

Myriapoda  {2'/^. 

PacJivnicriitni  fen-ug'nieuni  Latz.  Wide-spread  in  luir.,  undoubtedly  introduced, 
probably  from  Chile. 

OytJio))i()ypJia  gracilis  Koch,   Latz.  Apparently  wide-spread;   introduced. 
A  third  sj^ecies,   belonging  to  Lajiiyctes,  could  not  be  named. 

Collembola  (216). 

liiitoDiobyya  Diultifaciala  (Tullb.).  N.  and  S.  Amer.,  Eur.,  X.  As.,  N.  Zeal, 
J.   Viixw.    Introduced. 

Embioptera  [222). 

Oligotojfia    \\)sscl€ri  (Krauss).   Ceylon,   Sumatra,  Java. 

Insecta. 
Odonata   (225). 
PciDitala  flai'cscois  Fabr.   Amer.,  Afr.,  As.,  Austral. 

Orthoptefa-Dermaptera  (225,   20 1). 

I  am  indebted  to  Dr.  Princis  for  information  on  the  nomenclature  and  dis- 
tribution. 

Anisolabis  Boniiansi  Scudd.  Galap.  Is.,  Masatierra. 

OiiycJiostylus  uotulaius  (Stal;  Allacta,  22^.  297).  Formosa,  Malays.,  X.  Guin., 
X.   Caled.,   Samoa,  Tahiti,   Marquesas,   Hawaii.  Introduced   from  Tahiti? 

Periplancta  Australasiac  (Fabr.).  Probably  originally  African,  now  cosmop.; 
the  genus  Afr. -Orient. 

Diploptcra  punctata  (I^schtz,  D.  dytiscoides,  22j^.  297).  Ind.  Ceylon,  Burma, 
Malays.,  .Austral.,  Samoa,  .Marcjuesas,  Hawaii;  the  genus  Oriental.  Accidentally 
introduced.' 

McUviozostcria  pliilpotti  {^\\d.\\).  X.  Zeal.  An  Australian  genus,  represented  on 
some   Pacific   islands. 

lUatclla  raga  Hcb.  X.  Amer.;  Asia.'  The  genus  probably  Oriental.  Probably 
adventitious. 

Ofthoptera-Saltatoria  (j'V). 

(iyylliis  occaiiicus  Lc  (iuillou.  Malaysia  and  Japan  to  Polyn.  CllOTARD  regards 
its  presence  on  luister  Island  as  a  j)r()of  of  the  facility  with  which  certain  insects 
are  transpcjrted  large  distances;  nothing  will  prevent  animals,  he  says,  to  be  carried 
across  the  Pacific  fr(jm  yXustralia.  In  this  case,  however,  I  guess  that  man  has 
been   the  agent. 

Thysanoptera   [j). 

I laplotlirips  usitatus  Pagn.   \ar.   incrinis  Ahlb.  The  typical  species  in  Hawaii. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  419 

Neuroptera  (92). 

Chrysopa  laiiata  Banks.  Wide-spread  in  S.  Amer.  and  also  found  on  Hawaii. 
A  wide-ranging  genus  of  several  hundred  sp. 
-t-  Chr.  Skottsbergi  Esben-Peters. 

Lepidoptera  (zj). 

Agrotis  ypsiloii  Rott.  Cosmop. 
Cirphis  Loreyi  Dup.  Widely  distributed. 

Achaea  melicerta  Drury.  On  all  islands  in  the  Pacific  and  Indian  oceans. 
Phytometra  chalcytes  Esp.  Eur.,  As.,  most  islands  of  the  Pacific  and  Indian 
oceans. 

It  is  not  probable  that  any  of  these  Noctuidae  are  indigenous. 

Diptera  [84). 

Sarconesia  chlorogaster  (Wied.).   Chile,  J.   Fern.  Introduced. 

+  + Lip  Sana  insulae-paschalis  Enderl. 

Leptocera  (Coprophila)  ferruginaia  (Stenh.)  var.  insulae-paschalis  Enderl.  The 
typical  species,  known  from  Eur.,  Ind.  and  S.  Amer.,  lives  in  horse-dung  and 
was  spread  with  the  horse.  The  variety  was  picked  from  the  carcass  of  a  sheep 
and  has,  Enderlein  remarks  (I.e.  679),  perhaps  developed  after  the  arrival  in 
Easter  Island  where,  however,  there  are  many  horses. 

Coleoptera. 

Cii  rculionidae  (/.?) . 

Aramigus  Fulleri  Horn.  A  noxious  beetle,  probably  of  N.  Amer.  origin,  in- 
troduced into  many  countries  and  on  some  isolated  islands. 

+  Pentarthruni  paschale  Auriv. 

Areocerus  fasciculates  Deg.   Cosmop.,  introduced. 
Dytiscidae  [2gg). 

■V Bides sus  Skottsbergi  ZxvnmQvni.  The  occurrence  of  an  endemic  aquatic  beetle 
in    the    crater  lake  of  Rano  Kao  among  the  endemic  hygrophilous  mosses  is  of 
interest. 
Elateridae  [308). 

Siinodactylus  Delfijii  Fleut.   Chile.-^  Austral.,  N.  Guin.,  N.  Brit.,  Solomon  Is., 
Hawaii. 
Staphylinidae  [2^). 

Pliilonthus  longicorjiis  Steph.  Cosmop. 

Hymenoptera. 

Formicidae  {283). 

Ponera  trigona  Mayr.  var.  opacior  Forel.  N.  Amer.,  W.  Ind.,  Chile.  The 
typical   species  in  Brazil,  a  subspecies  in  Austral. 

Cardiocondyla  nuda  Mayr.  subsp.  minuta  Forel.   Hawaii.  The  typical  species 
Ind.,  Ceylon,  Austral.,  N.  Guin.  One  subsp.  is  Mediterranean. 
27* -557857 


420  C.  SKOTTSBERG 

Tetramorium  guiuecnse  Fabr.  Probably  of  African  origin,  now  pantrop.;  in 
hothouses  in  the  temp,  region. 

T.  siniil/iuiioH  F.   Smith.  As  the  former. 

Plagiolepis  niactai'ishi  Wheeler.   Formosa,  Hawaii,  Society  Is. 

Prenolepis  hourbojiica  Forel  subsp.  Skottsbergi  Wheeler.  The  typical  species 
known  from  Chagos,  Nicobar  and  Seychelle  Is.,  and  E.  Afr.,  Pemba  I.  Other 
subspecies  in  Ind.,   Comoro  Is.,   E.  As.,  Philipp.  Is.  and  Hawaii. 

Ants  are  easily  carried  about  by  man,  but  it  seems  likely  that  Piaster  Island 
also  has  indigenous  forms. 
Vespidae  (206). 

Polistes  Jiebraeus  F.  \i.  Afr.,  Madag.,  Ind.,  China;  Tahiti? 

Hemiptera  (21). 

Clerada  apicicoiiiis  Sign.   Wide-spread,  introduced. 
Redui'iolus  capsiformis  Germ.  As  the  former. 

Mollusca  {i8g). 

Umax  arboruDi  Bruch-Chant.   Cosmop.,  introduced. 

Milax  gagates  Drap.  As  the  former. 

Melampiis  philippi  Kuester.    Peru. 

+  J/.  pascus  Odhner.  The  genus  distributed  over  the  Pacific  from  Hawaii 
to  N.   Caled.    S.  Amer. 

Tor7iatellbwps  nnprcssa  Mouss.  (Syn.  Pacificella  variabilis  Odhner  I.e.).  Dis- 
tributed from  P^iji  to  Easter  Island.  Perhaps  introduced  with  living  plant  material. 

The  fauna,  as  known  hitherto,  presents  the  same  picture  of  extreme  poverty 
as  the  flora,  and  even  if  future  researches  will  double  the  number  of  species  and 
reveal  the  occurrence  of  groups  not  yet  recorded,  a  considerable  portion  will  con- 
sist of  late  immigrants.  The  known  endemics  are  very  few  and  one  or  two  of 
them  questionable,  and  our  experience  from  the  old  list  of  Juan  P'ernandez  Diptera 
(S4)  bodes  no  good  for  the  single  endemic  genus.  Altogether  half  a  dozen  en- 
demic species  and  some  endemic  forms  of  lower  category  have  been  described, 
and  of  the  species  found  elsewhere  some  are,  perhaps,  indigenous.  An  example 
of  remarkable  discontinuous  distribution  is  offered  by  Melanozosteria  pJiilpotti, 
New  Zealand  and  ICaster  Island ;  possibly  it  will  be  discovered  in  intermediate 
stations,  but  such  stations  are  difficult  to  find  in  other  cases  where  Piaster  Island 
is  the  terminus:  Anisolabus  BovDiausi,  Galapagos  Islands  -f  Juan  P^rnandez,  C/iry- 
sopa  lajiata.  South  America  and  Hawaii,  Poiicra  trigo)ia,  America,  and  llaplotJirips 
7U)tatus,  Hawaii.  A  direct  overseas  transport  is  not  very  probable,  and  I  cannot 
tell  if  these  animals  are  likely  to  have  been   introduced  with  the  traffic. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  421 

Chapter  XVI. 
The  biogeographical  history  of  Easter  Island. 

The  composition  of  the  present  fauna  and  flora  does  not  help  us  to  throw 
any  Hght  on  the  earlier  history  of  Easter  Island,  and  we  do  not  know  what  they 
were  like  before  the  arrival  of  aboriginal  man  many  centuries  ago.  We  know  that 
the  island  became  densely  populated,  that  the  natural  resources,  evidently  poor, 
were  exploited,  the  soil  cultivated  wherever  this  was  possible  and  a  number  of 
useful  plants  introduced  from  other  parts  of  Polynesia;  tradition  tells  that  the  first 
colonists  arrived  from  Rapa,  but  other  opinions  have  also  been  expressed.  ROGGE- 
VEEN,  the  discoverer  of  the  island  in  1722,  did  not  bring  a  naturalist,  but  to 
judge  by  his  narrative  the  island  must  have  looked  much  the  same  as  when 
Sparrman  and  the  FORSTERS,  who  came  with  CoOK  in  1774,  made  the  first  bio- 
logical observations.  Forster  collected  and  cited  a  few  species  (346)  and  men- 
tions, in  his  narrative  (j^/),  "Mimosa"  [SopJwra  toromiro)  and  Apiurn,  which  he 
had  observed  before  in  New  Zealand.  If  there  had  been  other  indigenous  trees, 
they  had  disappeared;  Broussonetia papyrifera,  Thespesia populnea  and  very  likely 
also  Triunifetta  semitriloba  had  been  introduced  but  were  scarce.  For  wood  the 
natives  depended  on  SopJiora,  and  most  of  this  was  gone  already.  FORSTER 
found  the  place  very  barren,  but  on  p.  578  he  speaks  of  a  hillock  covered  with 
toromiro,  and  later  on  another  similar  hill  is  mentioned  (p.  592),  but  all  the  trees 
were  low,  not  over  9  or  10  feet,  the  main  trunk  of  the  biggest  as  thick  as  a 
man's  thigh.  No  wonder  that  the  single  canoe  seen  was  a  patchwork  of  pieces  2 
or  3  feet  long,  and  so  was  the  paddle.  The  population  did  not  exceed  700.  When 
Thompson  and  Cooke  [343)  visited  the  island  in  1886,  groups  of  trees  were  ob- 
served in  some  places: 

In  other  parts  of  the  island  may  be  seen,  in  places  in  considerable  numbers,  a 
hardwood  tree,  more  properly  bush  or  brush,  called  by  the  natives  toromiro.  These 
must  have  flourished  well  at  one  time,  but  are  now  all,  or  nearly  all,  dead  and  decaying 
by  reason  of  being  stripped  of  their  bark  by  the  flocks  of  sheep  which  roam  at  will 
all  over  the  island.  None  of  the  trees  are,  perhaps,  over  10  feet  in  height,  nor  their 
trunks  more  than    2    or   3   inches  in   diameter  (p.  705). 

The  last  specimens  of  toromiro  are  restricted  to  the  inside  of  the  crater  Rano 
Kao.  Easter  Island  was  made  a  national  park  in  order  to  protect  the  unique  stone 
monuments,  and  is  a  bird  sanctuary,  but  otherwise  nature  is  not  preserved  but  the 
land  grazed  over  without  restriction  as  far  as  I  am  aware. 

Among  the  many  isolated  islands  of  the  Pacific,  Easter  occupies  a  rather 
unique  position.  Oceanic  islands  belong  to  two  main  categories,  high  volcanic  and 
low  coralline;  only  the  former  are  of  greater  biological  interest  and  possess  the 
standard  set  of  "peculiarities"  described  by  Hooker,  Wallace  and  others.  Easter 
Island  seems  to  form  a  type  by  itself.  It  is  volcanic  and  cannot  be  called  low,  for 
the    highest  mountain   is   530  m  high  and  some  of  the  craters  reach  an  altitude 


422  C.  SKOTTSBERG 

•of  300-400  m,  sufficient,  one  would  think,  to  create  a  humid  montane  belt  with 
fairly  luxuriant  arboreous  vegetation,  but  of  tliis  there  is  nothing,  in  any  case 
nothing  left.  Rains  are  frecjuent,  and  the  amount  of  precipitation  is  not  small, 
but  evaporation,  favoured  by  high  temperatures  and  the  strong  S.E.  trade  wind, 
is  great  and  most  of  the  water  rapidly  disappears  underground.  The  climax  vege- 
tation is  an  oceanic  steppe-like  meadow  or  grass  heath,  as  some  would  prefer 
to  call  it  (J42].  The  flora  docs  not  present  many  of  the  characteristics  of  oceanic 
islands.  There  are  no  endemic  genera,  no  peculiar  endemic  species,  no  prepon- 
derance of  woody  {)lants;  Sof^Jiora  is  the  only  tree  and  Lychnn  the  only  shrub; 
the  flora  is  herbaceous,  comj^rising  few  therophytes  but  many  annual  weeds.  The 
ratio  species  :  genus  is  1.2  :  i.  With  the  exception  of  Gra7uin€ae,  which  dominate, 
and  Cvpcraccac  (some  of  these  perhaps  not  indigenous)  most  large  and  world-wide 
families  are  absent,  even  Compositae;  there  are  no  conifers,  no  orchids  but  a  fair 
number  of  ferns:   in  these  respects  the  island  conforms  to  typical  oceanic  islands. 

On  the  other  hand,  luister  Island  has  little  in  common  with  the  low  islands, 
atolls  or  other  coralline  structures  with  no  rock  foundation  exposed,  where  en- 
<lemics  are,  as  a  rule,  absent  and  the  fragmentary  flora  consists  of  species  easily 
transported  by  the  natural  agencies  and  by  man. 

The  geographical  position  is  unfavourable  to  immigration,  the  chances  for 
the  arrival  of  seeds  from  America  small,  the  distance  being  3700  km,  and  1850 
km  separate  I^^.aster  from  Pitcairn,  the  nearest  basaltic  islet.  On  the  other  hand, 
the  chances  for  establishment  ought  to  have  been  good:  new  surroundings,  though 
perhaps  not  very  varied,  plenty  of  space,  no  competition,  conditions  furthering 
the  evolution  of  new  s})ecies  and  genera  as  many  biogeographers  believe,  but 
nothing  like  that  seems  to  have  happened  on  Easter  Island.  The  objection  will 
be  raised  that  we  do  not  know,  for  the  bulk  of  the  original  fauna  and  flora  may 
luue  been  destroyed  by  man  and  will  remain  unknown  —  but  is  it  not  surprising, 
if  this  be  true,  that  no  peculiar,  systematically  isolated  form  was  preserved  to 
our  da\-s.-  After  all,  |)erhaps  not.  Truly,  in  many  high  and  well  populated  Poly- 
nesian islands  endemics  are  plentiful,  growing  on  the  elevated  ridges,  on  the 
precij)itous  mountain  sides,  in  the  deep  recesses  of  the  gorges  where  cascades 
tumble  down  or  even  in  less  inaccessible,  but  uninhabited  places,  but  the  topo- 
graphy of  I'.aster  Island  is  different  and  I  can  see  no  reason  why  not  man  and 
his  animals  could  have  succeeded  to  exterminate  practically  everything  of  the 
original  nature  except  the  lichens  and  mosses  covering  the  rocks  and  a  few  herbs 
and   grasses. 

Distance  is  not  the  only  factor,  time  is  another;  the  island  is,  somebody 
will  say,  perha|)s  too  )-oung.  It  has  a  youthful  appearance,  the  craters  are  well 
|)reserve(l,  but  the\-  are  secondary  and  not  responsible  for  the  origin  of  the  island, 
and  their  well-preserved  shape  is  no  proof  of  youth.  There  is  no  sign  of  recent 
activity — I^^)KsrKR's  idea  that  the  decay  during  the  1 8th  century  of  the  old  ab- 
original culture  was  due  to  some  volcanic  catastrophe  lacks  foundation.  As  there 
are  no  permanent  streams,  erosion  must  be  slight.  I'he  various  raiws  and  other 
cones  may  be  old  enough  and  the  foundation,  on  which  they  stand,  very  ancient. 
As  BkifJGKX  says  (.?,v.  290): 


DERIVATION  OF  THE  FLORA  AND  FAUNA  423 

A  pesar  de  que  los  volcanes  .  .  .  tienen  sus  formas  muy  bien  conservadas,  no 
existe  en  la  tradicion  de  los  nativos  ningun  recuerdo  de  una  erupcion.  La  isla  pro- 
duce la  impresi6n  de  tratarse  del  resto  de  un  segmento  de  un  enorme  volcdn  central, 
en  cuya  superficie  inclinada  se  han  formado  los  crateres  actualmente  visibles  como 
conos  adventicios  superiores. 

Easter  Island  is  small,  but  not  very  small,  117  sq.  km,  and  GULICK  (iig) 
called  it  "the  wave-worn  remnant  of  an  island  that  could  once  have  claimed  about 
twice  that  area".  Lack  of  time  cannot  have  prevented  the  island  to  reach  matu- 
rity and  to  acquire  some  internal  harmony  of  the  flora  and  fauna.  Still,  they  are 
very  disharmonious.  The  general  situation  of  isolated  islands  is  clearly  set  forth 
by  GuLiCK  (I.e.  413-414): 

It  is  evident  that  mature  groups  of  islands  will  attain  an  internal  harmony  from 
the  standpoint  of  the  systematist.  But  this  harmony,  instead  of  reflecting  the  pre-existent 
harmony  of  some  continental  source  (as  is  the  case  for  continental  islands  or  land-bridge 
remnants),  will  be  recognizably  derivable  by  descent  from  a  quite  limited  number  of 
original  importations,  at  the  start  distinctly  miscellaneous  and  "disharmonic",  as  was 
observed  to  be  the  condition  in  Bermuda  and  St.  Helena.  Large  series  of  related  or 
previously  associated  forms  will  be  found  from  the  beginning  in  continental  islands, 
but  their  counterpart  must  be  brought  into  existence  de  novo  if  the  group  is  truly 
oceanic.  But  this  distinction,  obvious  in  theory,  is  in  practice  very  difficult  to  recog- 
nize, unless  the  oceanic  condition  is  really  extreme. 

Easter  Island  complies  with  this  condition  and  is  often  described  as  an  ex- 
ample of  a  truly  oceanic  island;  this  is  the  general  opinion,  but  it  was  told  above 
(Chapter  IV)  that  there  is  no  lack  of  theories  according  to  which  Easter  is  a  rem- 
nant of  a  land  mass  of  continental  size,  a  mid-Pacific  continent  or  a  land  bridge 
uniting  the  Australian-New  Zealandic  area  with  South  America  and  Hawaii;  Arldt, 
Germain,  Guillemin,  J.  W.  Gregory,  Meyrick,  Pilsbry  etc.  were  quoted.  It 
is  not  improbable  that  the  island  is  the  rest  of  a  somewhat  larger  piece  of  land, 
but  this  is  all  we  can  say.  The  bathymetrical  conditions — see  map — hardly  tempt 
us  to  construct  bridges,  even  if  not  all  signs  of  submergence  are  lacking.  The 
3000  m  curve  forms  a  large,  almost  closed  crescent,  on  the  north  extremity  of 
which  Easter  Island  rises.  E.N.E.  is  Sala  y  Gomez,  extending  S.W.-N.E.,  1200  m 
long,  150  m  broad  and  30  m  high,  and  this  tiny  islet  is  the  only  visible  part  of  a 
larger  reef  running  in  the  direction  N.E.  ^4  N.  and  called  Scott  Reef,  where  the 
smallest  depth,  1950  m  from  the  islet,  is  35  m  only;  between  this  place  and  the 
islet  a  series  of  soundings  gave  55,  60,  49  and  46  m.  The  bank  extends  at  least 
a  couple  of  km  west  of  the  islet  with  depths  of  42  to  68  m,  and  nowhere  within 
this  range  a  greater  depth  than  95  m  was  found  [172.  24).  The  scale  of  my  map  is 
too  small  to  give  these  details.  Proceeding  east  we  find,  in  about  the  same  dis- 
tance from  Sala  y  Gomez  as  this  is  from  Easter  Island  another  submarine  ridge  run- 
ning N.-S.,  bounded  by  the  1000  m  curve  and  with  depths  as  modest  as  862  and 
308  m.  Further  east  again,  in  97°3o'  w.  1.,  ?ipproximately,  is  another  ridge  trending 
W.-E.,  where  the  smallest  figure  is  497  m.  None  of  these  shallow  areas  were,  I 
believe,  known  to  the  bridge-builders.  They  must  be  welcome  also  to  those  who 
look,  if  not  for  sunken  continents,  at  least  for  submerged  islands  used  as  way 
stations    in    the    migrations.    However,    all   this  means  a  possible  extension  east. 


424 


C.  SKOTTSBERG 


towards  South  America  but  this  is  not  where  we  ought  to  look  for  the  vanished 
world  that  could  have  inhabited  a  greater  and  higher  Easter  Island;  rather,  we 
have  better  look  in  the  opposite  direction. 

It  is  FoRRKsr  Brown's  merit  to  have  pointed  to  the  large  insular  world 
known  as  Southeastern  Polynesia  as  as  important  floristic — and,  I  presume,  faun- 
istic — centre,  and  from  his  Flora  (jj)  and  the  reports  published  by  members  of 
other  recent  survey  parties,  sponsored  by  the  Bishop  Museum  in  Honolulu,  the 
following  data  were  compiled.  The  Society  Islands  are  excluded  and  only  the 
angiosperms  considered.  Included  are  the  Marquesas,  Tuamotu  and  Austral  or 
Tubuai  Islands,  and  the  more  isolated  Rapa  (Oparu),  Mangareva  (Gambler),  Pitcairn 
and   Henderson  (PLlizabeth)  Island. 

The  Marquesas  Islands  are  high,  800-1200  m,  and  belong,  to  judge  from  the 
geology  and  topography,  to  the  same  generation  of  Tertiary  islands  as  Tahiti, 
Juan  Fernandez,  Macaronesia,  etc.  The  Tuamotus  proper  are  atolls.  Of  the  Austral 
Islands  the  high  basaltic  are  considered,  Rimatara,  95  m,  Rurutu,  410  m,  Tubuai, 
400  m,  and  Raivavae  (Vavitao),  440  m;  further,  the  following  outlying  islands  are 
included:  Rapa  (Oparo),  640  m,  Mangareva  (Gambler),  400  m,  Pitcairn,  350  m> 
and  Henderson  (Elizabeth),  an  islet  of  raised  coral  said  to  be  only  25  m  high 
(  lY f )  but  nevertheless  the  home  of  an  endemic  Stvitaliwi  (compare  Laysan  of  the 
Leeward  Hawaiian  Islands  with  S.  cllipiicuui  var.  laysanense).  The  Marquesas  flora 
is  considered  to  be  well  known  and  the  same  may  be  true  of  the  flora  of  the 
otiicr  islands,  even  if  no  complete  lists  have  been  published;  I  suppose  that  all 
the  novelties  have  been  described,  but  my  figures  for  wide-spread  species  are, 
perhaps,  too  low.  There  is  a  difference  between  my  figures  and  those  given  by 
Brown,  because  varieties  are  counted  by  him  as  units  equal  to  species,  which 
e.\[)lains  why  his  figures  for  the  endemics  are  so  high. 

The  largest  families  are  Rubiaceae  (36),  Cyperaceae  (25),  Compositae  (19), 
lui[)horbiaceae  (16),  Gramineae  (13),  Leguminosae  (12),  and  Piperaceae  (10).  Other 
large  and  imj)ortant  families,  such  as  Araliaceae,  Cruciferae,  Ericaceae,  Malvaceae, 
Myrtaceae,  Orchidaceae,  Sapindaceae,  etc.,  are  represented  by  fewer  species.  We 
iiave  every  reason  to  believe  that  the  flora  has  suffered  losses  after  man  had 
taken   })ossessic)n   of  the  soil. 

The  t(jtal  number  of  presumably  indigenous  species — many  of  aboriginal 
introduction  and  not  few  later  arrivals  have  become  naturalized — is  282,  of  which 
156  are  endemic  within  the  area.  The  genera  are  145,  of  which  only  3  are  endemic 
according  to  Brown.  The  ratio  species  :  genus  is  almost  2:1.  No  genus  is  very 
large,  the  Largest  is  Psychotria  with  12  sjjecies,  and  10  have  from  5  to  10  species 
each.  The  distribution  of  the  species  and  the  number  of  local  endemics  are  indi- 
cated  in   Table  VIII.   The  figures  do  not  pretend  to  be  exact. 

A  large  proportif)n  of  endenfics  and  of  woody,  arboreous  or  fruticose  species 
— suffruticose  excluded — are  characteristic  of  oceanic  floras  of  considerable  anti- 
c}uity.  Of  the  156  endemic  species  113  (72.4%)  are  woody,  of  the  126  found 
elsewhere  69  (54.8  %).  The  herbaceous  species  are,  with  very  few  exceptions,  peren- 
nial. Systematically  isolated  tyi)es  are  few,  and  even  the  Marquesas  Islands  can- 
not, in  this  respect,   be  compared  with   either  Hawaii  or  Juan   P^ernandez. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  425 

Table  VIII.      . 
Distribution  of  angiosperms  in  Southeastern  Polynesia. 

Number  of      Number  of  %  Woody  species 

species  endemics  endemics  Number                  % 

Marquesas 151  79  52.3  100  66.2 

Tuamotu 44  3  7.0  23  52.3 

Austral 80  12  15.0  53  66.7 

Rapa 89  44  49.4  62  69.6 

Mangareva 28  3  10.7  17  60.9 

Pitcairn 26  2  7.7  19  73.1 

Henderson 21  3  14.3  i^  71  ^^ 

Total 282  15,6  55.3  182  64.5 

Brown  (jj)  regards  southeastern  Polynesia,  with  the  Tuamotus  in  the  centre, 
as  an  old,  submerged  region: 

Affinities  point  to  the  Tuamotuan  region  as  one  of  the  ancient  mid-oceanic  centers 
of  origin  for  a  large  part  of  the  dicotyledonous  flora  of  southeastern  Polynesia  (HI.  6); 

and,  speaking  of  the  distribution  of  Fitchia,  he  writes  (I.e.  364): 

The  grouping  and  affinity  of  these  allied  species  strongly  suggest  the  Tuamotuan 
region  as  the  center  of  origin  at  a  rather  remote  period,  possibly  at  the  dawn  of  the 
Tertiary  or  somewhat  earlier,  when  it  may  be  assumed  that  high  (pre-Tuamotuan)  islands 
existed  in  place  of  the  low  (Tuamotuan)  atolls  of  the  present. 

Within  the  Marquesas  archipelago  the  sea  is  shallow: 

Apparently,  an  emergence  of  100  meters  would  cause  land  to  appear  in  six  places; 
an  additional  emergence  of  300  meters  would  unite  or  bring  into  close  contact  all 
land  areas  of  the  archipelago.  .  .  .  Botanical  evidences,  outlined  in  an  earlier  paper,^ 
indicate  that  the  islands  were  at  one  time  1000  to  2000  meters  higher  than  at  pre- 
sent .  .  .  (I.e.  I.  17). 

The  floristic  affinities  of  this  region  is  with  Malaysia-Melanesia-Australia; 
there  is  no  neotropical  element  in  spite  of  the  prevailing  direction  of  winds  and 
currents.  Brown,  who  could  be  expected  also  to  look  toward  America,  remarks: 

The  Cichorieae,  to  which  Fitchia  belongs,  are  best  represented  in  Europe  and 
America,  pointing  to  a  more  remote  American  center  of  origin  for  the  pre-Tuamotuan 
ancestral  stock. 

This  brings  up  the  Dendroseris-Thamnosei'is  "^xohX&va.TX^^  iowx  <\^wdiXQ's^xQ\(\ 
genera  form  a  very  natural  group,  Thamnoseris  of  Desventuradas  Islands  stands 
apart,  and  so  does  Fitchia.  If  they  are,  at  least  distantly,  related,  and  isolated 
from  all  other  Cichoriaceous  genera,  then  the  possibility  of  an  Antarcto-tertiary 
ancestry  should  be  considered.  I  have,  however,  referred  the  Dendroseris  assem- 
blage to  an  ancient  neotropical  element,  absent  from  the  present  continental  flora, 


^  Proceed.  2d  Pan-Pacif.  Sci.  Congr.,  Vol.  2,   1923. 


426 


C.  SKOTTSBERG 


and  TJiamuoscris  finds  no  better  place,  but  to  derive  FitcJiia  from  an  American 
source  seems  little  inviting.  Another  solution  is,  perhaps,  in  sight.  Professor  GUN- 
XAR  KRnr.MAX  kindly  told  me  that,  to  judge  by  the  pollen  morphology,  Fitchia 
may  have  to  be  removed  from  the  Cichoriuui  subfamily  where  J.  D.  HoOKER 
placed  it  next  to  Dcudroscyis  and  where  it  has  remained. 

There  is  in  the  Pacific  Ocean  no  island  of  the  size,  geology  and  altitude  of 
l^aster  Island  with  such  an  extremely  poor  flora  and  with  a  subtropical  climate 
favourable  for  plant  growth,  but  nor  is  there  an  island  as  isolated  as  this,  and 
the  conclusion  will  be  that  poverty  is  a  result  of  isolation — even  if  man  is  re- 
sponsible for  the  disappearance  of  part  of  the  flora,  it  cannot  have  been  rich; 
the  Marquesas  Is.,  which  have  been  inhabited  longer,  I  believe,  and  formerly 
had  a  large  native  population,  still  preserve  a  fairly  rich  and  varied  angiospermic 
flora,  half  of  which  is  endemic.  The  distances  are  too  great  to  be  overcome 
except  on  very  rare  occasions.  The  nearest  land  is  to  the  west,  the  small  most 
easterly  islets  of  the  Alangareva  (Gambler)  group,  but  winds  (S.E.  trade-wind)  and 
currents  are  unfavourable  for  transport  from  \V.,  and  Piaster  Island  appears  to  lie 
away  from  the  cyclonic  tracks.  Beach  drift  is  responsible  for  the  arrival  of  several 
s{)ecies,  Ipoinaca,  Caesalpijiia,  Cheuopodium,  Tciragonia,  Eryt/iraea,  Apium,  Samo- 
lus,  LyciuDi  and  perhaps  some  grasses  and  species  of  Cypcrus,  altogether  about 
^'3  of  the  angiosperms.  Storms  bring  light  diaspores,  but  it  is  noteworthy  that 
CoDipositac  are  absent.  I  can  find  no  special  adaptations  for  bird  carriage,  but  the 
j)ossibility  of  rare  cases  of  epizoic  transport  cannot  be  excluded.  However  that 
may    be,  Piaster    is  a  good   example  of  an  island  peopled  by  "waifs  and  strays". 

Affinities  are,  as  we  have  seen,  with  Malaysia-Australia  or  pantropical,  whereas 
the  well-marked  east  Polynesian  flora  has  contributed  nothing,  not  even  its  leading 
famil}'  Rubiaceae,  rich  in  drupe-fruited  forms.  Sophora  torouiiro  is  allied  to  vS".  ''tetra- 
ptcrd^  of  Raivavae  and  Rapa;  I  cannot  tell  if  this  is  the  true /^/r^//rr^,  a  native 
of  New  Zealand,  but  I  do  not  think  it  is,  and  as  Brown's  description  (III.  120) 
shf)ws,  it  differs  much  from  toroniiro,  which  comes  very  close  to  S.  masafuerana. 
Neither  is  of  American  ancestry:  sect.  Edivardsia  is  austral-circumpolar  and  gene- 
rallv  regarded  to  be  of  Antarctic  origin  or,  at  least,   history. 

With  the  exception  of  Lyciian  carolinianiwi  var.  sa7idvicense,  supposed  to 
belong  to  the  beach  drift,  there  is,  if  FitcJiia  is  definitely  excluded,  no  American 
element  in  the  fiora  of  southeastern  Polynesia,  nor  is  it  expected  there.  It  is,  as 
we  have  seen,  found  in  I^aster  Island.  Of  the  3  endemic  grasses,  Stipa  was  ten- 
tatively brought  to  the  palaeotropical  element,  Axonopus  to  the  neotropical.  Dan- 
tlionia  is  an  austral-circum{)olar,  tricentric  genus.  Three  American,  not  endemic 
si)ecies.  Cypcrus  rraj^rostis,  Scirpus  riparius  and  Polygomim  acumiiiatmn,  remain 
to  be   accounted   for. 

\'{  I^^aster  Island  once  had  a  richer  flora  is  an  open  question.  According  to 
newspa[)er  reports  a  palynological  survey  of  the  swamp  in  the  crater  of  Rano 
Kao  was  planned  for  IllA  I.rdaiii/s  recent  survey.  The  thickness  of  the  loose, 
water-soaked  Cauipylopus  peat  was  not  measured  by  me;  it  is  a  somewhat  dan- 
gerous quagmire  which  cannot  be  bored  with  the  usual  methods,  but  samples  may 


DERIVATION  OF  THE  FLORA  AND  FAUNA  427 

be    dug    out  from  different  depths,  and  if  pollen  of  species  not  growing  on  the 
island  are  found,  some  light  will  be  thrown  on  the  history  of  the  flora. 

The  map  accompanying  this  paper  was  prepared  by  the  Oceanographical 
Institute  in  Goteborg.  I  am  greatly  indebted  to  the  Director,  Professor  Hans 
Pettersson,  and  to  Dr.  Borje  Kullenberg  for  valuable  assistance. 

May,   1956. 


Additions. 

P.  251. 

Usnea  Gaudichaudii  Motyka,  known  from  the  "espinal"  of  Central  and  North 
Chile,  is  quoted  for  Juan  Fernandez  (Masatierra)  by  MOTYKA  34^.  600  as  found 
by  Bertero,  1830.  As  its  occurrence  there  seems  little  probable — Bertero 
collected    also    near   Valparaiso,  etc. — I  have  excluded  the  species  from  my  list. 

To  Chapters  IV  and  XVI. 

A  recent  paper  by  R.  FURON,  "Importance  paleogeographique  des  mouve- 
ments  de  subsidence  du  Pacific  Central"  (Rev.  gen.  des  Sciences  62,  1955),  should 
be  noted  here.    His  object  is  expressed  in  the  following  terms: 

Constatant  combien  les  biogeographes  manquent  de  documentation  geologique,  il  nous  a 
paru  utile  de  regrouper  les  notions  acquises  au  cours  de  ces  dernieres  annees,  notions  qui 
eclairent  fort  bien  I'historie  du  Pacifique  depuis  le  Cretace  (p.  307). 

Whereas  the  Galapagos  Islands  show,  he  says,  a  purely  oceanic  type  of 
rocks,  andesitic  basalt  and  tuff  are  found  on  Easter  Island  and  andesite  and 
trachyte  on  Pitcairn.  Contrary  to  what  was  told  above,  corals  of  Cretaceous  age 
were  dredged  in  a  depth  of  2000  m  on  one  of  the  Central  Pacific  guyots.  The 
deep  borings  through  atolls,  the  latest  on  Eniwetok  in  1953,  have  penetrated 
through  coral  formations  dating  from  Pleistocene  to  Eocene  to  the  bedrock  of 
basalt,  indicating  a  subsidence  since  the  end  of  the  Cretaceous  of  2000  m.  Perhaps 
other  parts  of  the  Pacific  would  give  still  greater  figures. 

To  Chapter  V. 

In  the  Proceedings  of  the  Cotterwood  Natur.  Field  Club  31,  1955,  T.  A. 
Sprague  gives  an  account  of  the  Drift  Theory  of  Du  ToiT  and  finds  that,  from 
a  botanist's  viewpoint,  this  theory  "offers  the  best  explanation  hitherto  brought 
forward  of  the  major  problems  of  biogeography".  With  reference  to  the  physical 
side  of  the  drift  process  he  quotes  Holmes'  "Principles  of  Physical  Geography" 
(1944).  In  the  case  of  Juan  Fernandez,  which  certainly  is  one  of  the  minor  pro- 
blems, I  cannot  see  that  the  drift  theory  offers  an  acceptable  solution. 

To  Chapter  VII,   p.   360. 

An  important  paper  by  H.  M0LIIOLM  Hansen,  "Life  forms  as  age  indica- 
tors", Ringkjobing   1956,   confirms  the  opinions  of  SiNNOTT  &  Bailev  and  others. 


Bibliography. 

r.   Ar.r.AVKs,  H.  dfs.   De  speciebiis  generis  lichenum   Cladoniae  ex  Insulis  Tristan  da 

Ciinba.    Results   Norwcg.   Kxj).   to...,   no.    4.    1940. 
Adams,  Lk  Rov.  The  theory  of  isolation  as  apj)lied  to  plants.  Science  22,  1905. 
Ahi.I'.krc,  O.  Thysanoptera   from  Juan   Fernandez  and  Easter  Island.  Nat.  Hist.  Juan 

I'ern.  Ill,  1922. 
Alkxandkk,  C.  r.  Tipulidae  (l)iptera).  (insect.  J.  Fern.  5.)  Rev.  Chil.  de  Entomol.  2, 

1952. 
Andrfavks,   H.  T.   Coleoptera-Carabidae  of  the  Juan  Fernandez  Islands.  Nat.  Hist. 

Juan  Fern.  Ill,   i  93  i . 
Andrkws,  F.  C.  Origin   of  the  Pacific  Insular  Floras.  Proceed.  6th  Pacif.  Sci.  Congr. 

1939,   IV.   Berkeley    1940. 

The   structure  of  the   Pacific  Pasin.   Ibid.   VI. 

Ari.dt,   Th.    Die  Entwickelung  der  Kontinente  und  ihrer  Febewelt.   Leipzig  1907, 

2d  ed.  (vol.    I   only),   Berlin    1938. 
Arnf.ll,  S.    List    of   Hepaticae    collected    in  Marion   Island  by  Mr.   K.   W.   Rand. 

Svensk   bot.   tidskr.    47,    1953. 

Hepaticae  of  Chile  and  Argentina  collected  by  R.  Santesson.  Ibid.  49,  1955. 

An. I',  J.  P.  Preliminary   results   of  Ocean  magnetic  observations  on  the  Carnegie  .  .  . 

October  1928  to  January  1929.  Terrestrial  Magnetism  and  Atmospheric  Electri- 
city, March    1929. 

12.  AuRixir.Lius,  Chr.  Coleoptera-Curculionidae  von  Juan  Fernandez  und  der  OsterTnsel. 

Nat.    Hist.  Juan   Yern.   Ill,    1926. 

13.  AuRiviFijis,  CiiR.,  Prout,  I>.  B.  cv  Mfvrick,  E.  Lepidoptera  von  Juan  Fernandez  und 

der   ( )stcr-Insel.   Ibid.    1922. 

14.  AxKFRoi),  D.J.   Variables   affecting  the  probabilities  of  dispersal.   Proceed,  of  the 

Symposium  on  the  South  Atlantic  Basin,  1949.  Bull.  Amer.  Mus.  Nat.  Hist.  99, 
1952. 
15. A   theory  of  Angiosi)erm   evolution.   Evolution   \T,    1952. 

16.  Bai.F()i;r-1')R()\vnk,  J.  A(|uati(-,   Coleoptera   of  Oceania.   Occ.   Pap.   B.  P.  Bishop  Mus. 

Honolulu   XVIII:  7,    1945. 

17.  Bariram,  E.  V>.  Mosses  of  Chile  and  Argentina  mainly  collected  by  R.  Santesson. 

Svensk   bot.   tidskr.    46,    1952. 

18.  Bfchvnf,  J.  Alticidae   (Coleojitera).   (insect.  J.   I'ern.  i  i .)  Rev.  Chil.  de  Entomol.  2, 

I  ()  5  2 . 

19.  P)FiKR,  M.  I'seudoscorpione   von  Juan   Fernandez.    Rev.   Chil.   de  Entomol.  4,  1955. 

20.  P)FN'iHAM,  (j.  Notes  on   the  classification,  history  and  geographical  distribution  of  the 

Compositae.  Journ.    Linn.   Soc.   P)Ot.    13,  1873. 

21.  Bfrgrdih,  E.  Hemiptera   from  Juan    I'crnandez  and   Easter   Island.  Nat.  Hist.  Juan 

Fern.    Ill,    1924. 

22.  Bfrlaxi),  L.   Araignees  de  I'ile  de  Pa(|ues  ct  dcs  iles  Juan  Fernandez.   Nat.   Hist. 

Juan  Fern.   Ill,    1924. 

23. •  Remar(|ues  sur  la  rej)artition  et  les  affinites  des  Araignees  du  Pacifique.  Proceed. 

3d   Pacif.   Sci.   Congr.    1926.   Tokyo    1928. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  429 

24.  Berlioz,  J.  Quelqiies  considerations  entomologiques  et  conclusions  siir  le  role  bio- 

geographique    de  la    «Ligne  de  Wallace*;   Compte  rendu  Soc.  de  Biogeogr. 
30^  annee,  no.  258,  1953. 

25.  Bernhauer,  M.   Coleoptera-Staphylinidae   von    den  Juan  Fernandez-Inseln  und  der 

Osterinsel.  Nat.   Hist.  Juan  Fern.  Ill,    192 1. 

26.  Berry,  E.  W.  The  origin  and  distribution  of  the  family  Myrtaceae.  Bot.  Gaz.  59,  19  15. 
27. The  Flora  of  the  Concepcion-Arauco  coal  measures  of  Chile.  Johns  Hopkins 

Univ.   Stud.,   Geol.   4.   Baltimore   1922. 

28. Tertiary  Flora  from  the  Rio  Pichileufu,  Argentina.  Geol.  Soc.  of  Amer.  Spec. 

Pap.   No.    12.   Baltimore    1938. 

29.  Berry,  E.  W.  &  Singewald,  J.  T.  jr.  The  tectonic  history  of  Western  South  America. 

Proceed.  3d  Pacif.   Sci.  Congr.    1926.  Tokyo    1928. 

30.  Bews,  J.  W.  The  World's  Grasses.   London    1929. 

31.  Bitter,  G.  Die  Gattung  Acaena.  Bibl.  Botan.  74,    1910-11. 

32.  BouLY  DE  Lesdain,  M.  Lichens  recueillis  en  1930  dans  les  iles  Kerguelen,  Saint-Paul 

et  Amsterdam.   Ann.   Crypt,  exot.   4,    1931. 
;^;^.  Brotherus,  V.  F.  Musci  Insulae-Paschalis.  Nat.   Hist.  Juan  Fern.  II,    1924. 
34. The  Musci  of  the  Juan  Fernandez  Islands.   Ibid. 

35.  Brown,  F.  B.  H.  Flora  of  Southeastern  Polynesia.   I  (1931),   II  (1931),  III  (1935). 

Bernice   P.  Bishop  Mus.  Bull.  84,  89,  130.  Honolulu. 

36.  Bruce,  N.  Coleoptera-Cryptofagidae  von  Juan  Fernandez.  Nat.  Hist.  Juan  Fern.  Ill, 

1940. 

37.  Brues,  C.  T.  a  new  species  of  parasitic  Hymenoptera  from  Juan  Fernandez.  Ibid. 

1924. 

38.  Burtt,  B.  L.  &  Hill,  A.  W.  The  genera  Gaultheria  and  Pernettya  in  New  Zealand, 

Tasmania  and  Australia.  Linn.   Soc.  Journ.  Bot.   49:611,    1935. 

39.  Bryan,  E.  H.  jr.  A  summary  of  the  Hawaiian  birds.  Proceed.  6th  Pacif.  Sci.  Congr. 

1939,   IV.  Berkeley    1940. 
40. The  Hawaiian  Chain.   Honolulu    1954. 

41.  Burger,  O.  Die  Robinson-Insel.   Leipzig   1909. 

42.  Cain,  S.  A.  Foundations  of  Plant  Geography.  New  York  and  London    1944. 

43.  Camp,  W.   Distribution  patterns  of  modern  plants  and  the  problems  of  ancient  dis- 

persals. Ecol.    Monogr.    17,    1947. 

44.  Campbell,  D.  H.   The  Derivation  of  the  Flora  of  Hawaii.  Leland  Stanford  jr.  Univ. 

Publ.    1 919. 

45.  — ■ —  Australian  element  in  the  Hawaiian  Flora.  Amer.  Journ.  of  Bot.    15,    1928. 
46. The   Flora  of  the  Hawaiian  Islands.   Quarterly  Rev.  Biol.   8,    1933. 

47. Continental  drift  and  plant  distribution.  Science  95,    1942. 

48.  Cardot,  J.  La  Flore  bryologique  des  Terres  Magellaniques,  de  la  Georgie  du  Sud  et 

de  i'Antarctide.  Wiss.  Ergebn.  schwed.  Sudpolar-Exp.   1 901-1903.  IV:  8,  1908. 

49.  Cardot,  J.  &  Brothf;rus,  V.  F.  Les  Mousses.  Bot.  Ergebn.  schwed.  Exp.  nach  Pata- 

gonien  und  dem  Feuerlande  1907-1909.  K.  Sv.  Vet.-akad.  Handl.  63:  10,  1923. 

50.  Carvalho,  J.  C.  M.  Miridae  (Hemiptera).  (Insect.  J.  Fern.  3.)  Rev.  Chil.  de  Entomol. 

2,    1952. 

51.  Catalogue  of  Birds  of  the  British  Museum  (Natural  History)  V  (1881),  XVI  (1892). 

52.  Chaigneau,  J.  F.  Cordon  submarino  paralelo  a  las  costas  de  Chile  entre  las  islas  Juan 

Fernandez  y  San   Ambrosio.   Anuario  hidrogr.  Marina  de  Chile  22,  1900. 

53.  Chaney,  R.  W.  Tertiary  centres  and  migration  routes.  Ecol.  Monogr.    17,  1947. 

54.  Cheeseman,  T.  F.  The  vascular  flora  of  Macquarie  Island.  Scient.  Reports  Australas. 

Antarct.  Exp.  1911-14,  VII.  1919. 
55. Manual  of  the  New  Zealand  Flora.    2d  ed.  Wellington    1925. 

56.  Chilton,  Ch.  A    small    collection   of  Amphipoda  from  Juan  Fernandez.  Nat.  Hist. 

Juan  Fern.   Ill,    192 1. 

57.  China,  W.  E.  Homoptera-Cicadellidae.  Rev.  Chil.   de  Entomol.   4,    1955. 

28  -  557857  The  Nat.  Hist,  of  Juan  Fernandez  and  Easter  Isl.  Vol.  I 


43©  C.  SKOTTSBERG 

58.  Chopard,  L.  Gryllides    de  Juan  Fernandez  et  de  Tile  de  Paques.  Nat.   Hist.  Juan 

P>rn.  Ill,    1924. 

59.  Christ,  H.  Geographic  dcr  Fame.  Jena  1910. 

60.  Christenskn,  C.  Demonstration  of  C.  Skottsberg's  fern  collection  from  Juan  Fernan- 

dez.  Botan.   Tidsskr.    37,    1920. 
61. The  Pteridophytes  of  Tristan  da  Cunha.  Res.  Norweg.  Exp.  1 937-1 938,  no.  6. 

Oslo    1940. 
62.   Chrisik.nskn,  C.  &:  SKorisuERG,  C.  The  Pteridophyta  of  the  Juan  P>rnandez  Islands. 

Nat.  Hist.  Juan  Fern.  II,  1920. 
63. The  Ferns  of  Easter  Island.  Ibid. 

64.  CL.AUSEN,  R.  T.  A  monograph  of  the  Ophioglossaceae.  Mem.  Torrey  Bot.  Club,  10, 

1938. 

65.  Continental  Drift.  A  Symposium  on  the  Theory  of — .  The  Advancement  of  Science 

VHI:  29,  195  I. 

66.  CoPELANi),  E.  B.  Genera   Hymenophyllacearum.  Philipp.  Journ.  Sci.  67,  1938. 
67. Fern  evolution   in   Antarctica.   Ibid.  70,  1939. 

68. Antarctica    as  the  source  of  existing  ferns.   Proceed.   6th  Pacif.   Sci.  Congr. 

1939,  IV.  Berkeley    1940. 
69. Genera  Filicum.   Annal.   Crypt,   et  Phytopath.    5.   Waltham,   Mass.    1947. 

70.  CoRrEs,  R.  Tachynidae  (Diptera).  (insect.  J.  Fern.  9.)  Rev.  Chil.  de  Entomol.  2,  1952. 

71.  Croizat,  L.  Manual  of  Phyto-Geography.  The  Hague  1952. 

72.  Dahl,  E.  On  different  types  of  unglaciated  areas  during  the  ice  ages  and  their  sig- 

nificance to  phytogeography.  New  Phytologist  45,  1916. 

73.  Darp.ishire,  O.  V.  The   Lichens.   Wiss.  Ergebn.  schwed.  Siidpolar-Exp.    1901-1903, 

IV:  I.   Stockholm    191 2. 

74.  Dodge,  C.  W.  Lichens  and  Lichen  Parasites.  Reports  Brit.  Austral.  N.  Zeal.  Antarct. 

Research   Exp.  1929-31.  Ser.  B.  VII.  Adelaide  1948. 

75.  DoMiN,  K.  Monograi)hische  Ubersicht  der  Gattung  Centella  L.  Bot.  Jahrb.  fiir  System. 

etc.  ed.  by  A.  Engler,  41,  1908. 

76. Grundzuge  der  })flanzengeographischen  Verbreitung  und  Gliederung  der  Leber- 

moose.  Mem.  Soc.  Roy.  de  Sciences  de  Boheme  1923.  Prague  1924. 

77.  Du  RiETZ,  G.  E.  Two  new  species  of  Euphrasia  from  the  Philippines  and  their  phyto- 
gcographical  significance.   Svensk  bot.  tidskr.  25,  1931. 

78. Problems  of  bipolar  plant  distribution.  Acta  Phytogeogr.  Suec.  XIII,  1940. 

79.  Dlsex,  p.  Uber  die  tertiare  Flora  der  Magellanslander.  Wiss.  Ergebn.  schwed.  Exp. 
1 89  5- 1  897,  I,  1907. 

80. Uber  die  tertiare  Flora  der  ScymourTnsel.  Wiss.  Ergebn.  schwed.  Siidpolar- 
Exp.  1901-1903,  III,  1910. 

81.   Du  ToiT,  L.  Our  Wandering  Continents.  Edinburgh  and  London  1937. 

82. Observations  on  the  evolution  of  the  Pacific  Ocean.  Proceed.  6th  Pacif.  Sci. 

Congr.  1939,  I-  Berkeley  1940. 

83.  Emerson,  E.  A.  A  new  'J'ermite  from  the  Juan  Fernandez  Islands.  Nat.  Hist.  Juan 

Fern.  Ill,  i  924. 

84.  Enderi.ein,  G.  Die   DijUcrenfauna  der   fuan  P'ernandez-Inseln   und  der   Oster-Insel. 

Ibid.  1938. 

85.  Engi.ek,  a.  Versuch  einer  Entwickelungsgeschichte  dcr  Pflanztnwelt  seit  der  Tertiar- 

periode.  II.  1882. 

86.  EN(;i.ER'r.  S.  La  Tierra  de  Hotu  Matua.  Valparaiso  1948. 

87.  Epi.ing,  C.  Synopsis  of  the  South  American  Labiatae.  Fedde,  Repert.  Beih.  85,  1935. 

88.  • The  Labiatae  of  Chile.  Rev.  Universitaria  22,  1937. 

89.  — • — ■  The  distribution  of  the   American   Labiatae.  Proceed.  6th  Pacif.  Sci.  Congr. 

1939,  IV.  Berkeley  1940. 

90. ■  Distribuci6n  geogrdfica  y  parentesco  de  las  Labiadas  de  la  America  del  Sur. 

Rev.  Universitaria  25,  1940. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  43 1 

91.  Ermel,  a.  Eine  Reise  nach  der  Robinson-Insel.  Hamburg  1889. 

92.  Esben-Petersen,  p.  More  Neuroptera  from  Juan  Eernandez  and  Easter  Island.  Nat. 

Hist.  Juan  Fern.  Ill,  1924. 

93.  Evans,  A.  W.  The  Thallose  Hepaticae  of  the  Juan  Fernandez  Islands.  Ibid.  II,  1930. 

94.  Fassett,  N.  C.  Callitriche  in  the  New  World.  Rhodora  53,  1951. 

95.  Florin,  R.  The  Tertiary  fossil  conifers  of  South  Chile  and  their  phytogeographical 

significance.  K.  Sv.  Vet.-akad.  Handl.  ser.  3,  19,  1940. 

96.  Fluke,  C.  L.  Syrphidae.  (insect.  J.  Fern.  18.)  Rev.  Chil.  de  Entomol.  4,  1955. 

97.  FosBERG,  F.  R.  Derivation  of  the  Flora  of  the  Hawaiian  Islands.  See  Zimmerman,  E.  C. 
98. Lignes  biogeographiques  dans  I'ouest  du  Pacifique.  Compte  rendu  Soc.  de  Bio- 

geogr.  29  annee,  no.  256,  1952. 
99. The  American  element  in  the  Hawaiian  Flora.  Proceed.  7th  Intern.  Botan. 

Congr.  1950.  Stockholm  1953. 
100.   Freeman,  P.  Mycetophilidae,  Sciaridae,  Cecidomyidae  and  Scatopsidae.  (insect.  J. 

Fern.  13.)  Rev.  Chil.  de  Entomol.  3,  1953. 
loi.   Fries,  R.  E.  Die  Myxomyceten  der  Juan  Fernandez-lnseln.  Nat.  Hist.  Juan  Fern.  II, 

1920. 

102.  Fries,  Th.  jr.  Die  Gasteromyceten  der  Juan  Fernandez- und  Osterinseln.  Ibid.  1922. 

103.  FuLFORD,  M.  Some  distribution  patterns  of  South  American  Leafy  Hepaticae.  Proceed. 

7th  Intern.  Botan.  Congr.  1950.  Stockholm  1953. 

104.  Gebien,  H.  Coleoptera-Tenebrionidae  von  Juan  Fernandez.  Nat.  Hist.  Juan  Fern.  Ill, 

192  I. 
T05.   Germain,  L.  L'origine  et  revolution  de  la  faune  de  Hawaii.  Proceed.  3d  Pacif.  Sci. 
Congr.   1926.  Tokyo  1928. 

106.  GiBBS,  L.  Notes  on  Phytogeography  and  Flora  of  the  mountain  summit  plateau  of 

Tasmania.  Journ.  of  Ecology  8,    1920. 

107.  GiLLOGLY,  L.  R.  Coleoptera-Nitidulidae.  (insect.  J.  Fern.  24.)  Rev.  Chil.  de  Entomol. 

4,  1955- 

108.  Good,  R.  See  Continental  Drift. 

109. The  Geography  of  the  Flowering  Plants.  2d  ed.  London  1953. 

no.   GooDALL,  J.  D.,  Johnson,  A.  W.,  Philippi,  R.  A.  Las  Aves  de  Chile.  I,  1946.  II, 

195  I.  Buenos  Aires. 
III.   GooDSPEED,  T.  H.  Plant  Hunters  in  the  Andes.  New  York  1941. 
112. The  genus  Nicotiana.  Waltham,  Mass.  1954- 

113.  Gordon,  H.  D.  The  problem  of  sub-antarctic  plant  distribution.  Report  Austral,  and 

New  Zeal.  Ass.  Advancem.  Sci,  27,  1949- 

114.  Grant,  Adele  L.  A  monograph  of  the  genus  Mimulus.  Ann.  Miss.  Botan.  Card.  XI, 

1924. 

115.  Gregory,  H.  E.  Types  of  Pacific  Islands.  Proceed.  3d  Pacif.  Sci.  Congr.  1926,  2. 

Tokyo  1928. 

116.  Gregory,  J.  W.  Theories  of  the  origin  of  the  Pacific.  Proceed.  Geol.  Soc.  London 

86,  1930. 

117.  GuiGNOT,  F.  Dytiscidae  (Coleoptera).  (Insect.  J.  Fern.  10.)  Rev.  Chil.  de  Entomol.  2, 

1952. 

118.  GuiLLAUMiN,  A.   Les  regions  floristiques  du  Pacific  d'apr^s  leur  endemisme  et  la 

repartition  de  quelques  plantes  phanerogames.  Proceed.  3d  Pacif.  Sci.  Congr. 
1926.  Tokyo  1928. 

119.  GuLiCK,  A.  Biological  peculiarities  of  oceanic  islands.  Quart.  Review  Biol.  7,  1932. 
120    GuNDERSEN,  A.  Families  of  Dicotyledons.  Waltham,  Mass.  1950. 

121.  GUPPY,  H.  B.  Observations  of  a  naturalist  in  the  Pacific  between  1896  and  1899.  11. 

London  1906. 
122. The  island  and  the  continent.  Journ.  of  Ecol.  7,  1919. 


432  C.  SKOTTSBERG 

123.  GuTKNBKRG,  B.  Geophvsical   and   geological  observations  in  the  Pacific  area  and 

tectonic  hypotheses.  Proceed.  7th  Pacif.  Sci.  Congr.  1949,  II.  Wellington  1953. 

124.  GoKTSCH,  W.  Die    Robinson-Insel  jiian  Fernandez  iind  ihre  biogeographischen  Pro- 

bleme.  Phoenix  19,  1933. 

125.  Handschin,  C.  Neuroptera.   (Insect.  J.  Fern.    15.)  Rev.  Chil.  de  Fntomol.  4,  1955. 

126.  Harmstox.  F.  C.  r)olichoj)odidae.  (Insect.  J.  Fern.  17.)  Ibid. 

127.  Hkmsi.f.v,  \V.  B.  (a)   Rej)ort  on  the   Present  State  of  Knowledge  of  Various  Insular 

Floras,   (b)   Report  on  the  Botany  of  Juan  Fernandez  and  Masafuera.  Rep, 
Sci.  Res.  H.  M.  S.  Challenger.  Bot.  I,  1885. 

128.  Hf.nnig,  W.   Phryneidae,   Helomyzidae,    Lonchaeidae,  Piophilidae,   Anthomyzidae, 

Muscidae.  (insect.  J.  Fern.  16.)  Rev.  Chil.  de  Entomol.  4,  1955. 

129.  Hf.rzog,  Th.  (ieographie  der  Moose.  Jena  1926. 

130.  — - —  Die  foliosen  Lebermoose  der  Juan  Fernandez-Inseln  und  der  Oster-Insel.  Nat, 

Hist.  Juan  Fern.  II,   1942, 
131. Zur  Bryophytenrtora  Chiles.  Revue  Bryol.  et  Lichenol.  XXIII,  1954, 

132.  Hfss,  H.  H.  Drowned  ancient  islands  of  the  Pacific.  Amer.  Journ.  Sci.  244,  1946. 

133.  Hkkfx,  C.  M.  Polypodiacearum  argentinarum  catalogus.  Rev.  Mus.  de  La  Plata  15, 

I  908. 
I  34,   Hill,  A.  W.  Antarctica  and  j)roblems  of  geographical  distribution.  Proceed,  5th  Intern, 
Bot,  Congr.  1926.  Ithaca  1929. 

135.  HiNTON,  H.  E.  See  Continental  Drift. 

136.  HoDGSOX,  E.  Amy  &  Saixsburv,   G,  O.  K.  Bryophytes  collected  by  G.  E.  Du  Rietz 

on  the  Antipodes  Islands.  Svensk  bot,  tidskr.  42,  1948, 

137.  HoLTFDAHL,  O.  On  the  geology  and  physiography  of  some  Antarctic  and  Sub-Ant- 

arctic islands,   Scient.   Res.  Norweg.  Antarct.  Exp.  No.  3,  Ed.  by  Det  norske 
videnskaj)sakademi.   Oslo  1929. 

138.  Hooker,  J.  I).  Lecture  on   Insular  Floras  before  the  British  x\ssociation  at  Notting- 

ham, 1866.  Also  in  Gard.  Chronicle,  Jan.  1867. 

139.  Hue,  A.-M.  Lichens.  Deuxieme  Exj)ed.  Antarct.  Fran^aise  1908-1910.  Paris  1915. 

140.  HuTCHhxsoN,  J.  The  Families  of  Flowering  Plants.  I.  1926.  II.  1934. 

141.  Huxley,  J.  Species  formation   and  geographical  isolation.  Proceed.   Linnean  Soc, 

Lond.  1938. 

142.  IxsEcros.  Los   Insectos  de  las  islas  Juan  Fernandez.  Ed.  by  Dr.  G,  Kuschel,  1-12. 

Rev.  Chil.  de  Entomol,  2,1951.  —  13,14,  vol.  3,  1953.  —  i  5-27,  vol,  4,  1955. 

143.  Irmschek,  E.  Pflanzenverbreitung  und  Entwickelung  der  Kontinente.  I-II.  Mitteil, 

Hamburg.  Inst,  allgem.  Botan.  5,  1922;  8,  1929. 

144.  Jeffrey,  E.  C.  <S:  Torrey,  P..  S,  Transitional   Herbaceous  Dicotyledons.   Annals  of 

Botany  35,  192  i. 

145.  Jeffreys,  H.  See  Continental  Drift. 

146.  JnERixG,  H.  Das  neotropische  Florengebiet  und  seine  Geschichte.   Bot,  Jahrb,  fiir 

System,  etc.  ed.  by  A.  Exgler,  17,  1893,  Beibl,  42. 

147.  JonAXSSON,  L.  Ein  neuer  Landblutegel  aus  den  Juan  P'ernandez-Inseln,  Nat,  Hist. 

Juan  Fern,  III,  1924. 

148.  JoHXSTOx,  I.  M.  A   revision  of  the  South   American   Boraginoideae.   Contrib.  Gray 

Herb.  78,  1927. 
149. Papers  on  the   Flora  of  Northern  Chile,  i.  The  coastal  Flora  of  the  Depart- 
ments of  Chanaral  and  Taltal.  Ibid.  85,  1929. 

150.  JoHow,  F.  Estudios  sobre  la  Flora  de  las  Islas  de  Juan  Fernandez,  Santiago  1896. 

151.  JoRDAX,  D.  S.  The  origin   of  species  through  isolation.  Science  22,  1905. 

1  52. Isolation  with  segregation  as  a  factor  in  organic  evolution.  Annual  Rep.  Smith- 
son.  Inst.  1925.  Washington  1926. 

153-  JoRDAX,  K.  Coleoptera-Anthribidae  from  Juan  Fernandez.  Nat.  Hist.  Juan  Fern,  III^ 

1931- 

154-  Joyce,  L  R.  T.  See  Continental  Drift, 


DERIVATION  OF  THE  FLORA  AND  FAUNA 


433 


155.  Just,  Th.  Geology  and  Plant  Distribution.  Ecol.  Monogr.  17,  1947. 

156.  Kausel,  E.  Contribuci6n  al  Estudio  de  las  Mirtdceas  Chilenas  I-II.  Rev.  Argent,  de 

Agron.  9,  1942. 

157. Notas  Mirtol6gicas.  Lilloa  13,  1947. 

158. Revisi6n  del  genero  Escallonia.  Darwiniana  10,  1953. 

159.  Kkissler,  K.  Ascomyceten,  Fungi  imperfecti  und  Uredineen  von  Juan  Fernandez. 

Nat.  Hist.  Juan  Fern.  II,  1928. 
160. Nachtrag  zur  Pilzflora  von  Juan  Fernandez.  Ibid. 

161.  KoRMiLEV,  N.  A.  Lygaeidae  (Hemiptera).  (insect.  J.  Fern,  i.)  Rev.  Chil.  de  Entomol. 

2,    1952. 

162.  Krause,  K.  Uber  einen  hapaxanthen  Baum.  Mitteil.  deutsch.  dendrol.  Ges.  1921. 

163.  Krausel,  R.  Pflanzenwanderungen  im  Tertiar.  Proceed.  7th  Intern.  Botan.  Congr. 

1950.  Stockholm  1953. 

164.  KuKENTHAL,  G.  Vorarbeiten  zu  einer  Monographic  der  Rhynchosporoideae.  VIII. 

Fedde,  Repert.  48,  1940.  —  XII.  Ibid.  51,  1942. 

165.  Eawrence,  G.  H.  M.  Taxonomy  of  Vascular  Plants.  New  York  1951. 

166.  Lesne,  p.   Colepptera-Bostrychidae  et  Cioidae  de  Juan  Fernandez.  Nat.  Hist.  Juan 

Fern.   Ill,    1923. 

167.  Lindsay,  A.  Biology  and  biogeography  of  the  Antarctic  and  Subantarctic  Pacific. 

Proceed.  6th  Pacif.  Sci.  Congr.  1939,  II.  Berkeley  1940. 

168.  Looser,  G.  Nueva  localidad  cerca  de  Antofagasta  del  Helecho  Polypodium  Masa- 

fuerae  Philippi.  Rev.  Universit.  28.  Santiago  1943. 

169. Los  Blechnum  (Filices)  de  Chile.  Ibid.  32,  1947. 

170. Comparacion  de  las  Bromeliiiceas  de  Chile  con  las  de  Argentina  y  Peru.  Rev. 

Chil.  de  Hist,  y  Geogr.  n.  iio  (1947),  1948. 
171. El  genero  Polypodium  L.  y  sus  representantes  en  Chile.  Rev.  Universit.  ;^6. 

Santiago  195  i. 

172.  Lopez,  J.  E.  Esploraci6n  de  las  Islas  esporadicas  al  occidente  de  la  costa  de  Chile. 

Anuario  Hidrogrdf.  de  la  Marina  de  Chile.  II.  1876. 

173.  LouRTEiG,  Alicia.  Ranunculaceas  de  Sudamerica  templada.  Darwiniana  9,  1952. 

174.  LoNNBERG,  E.  The  Birds  of  the  Juan  Fernandez  Islands.  Nat.  Hist.  Juan  Fern.  Ill, 

1 920. 
175. Notes  on  Birds  from  Easter  Island.  Ibid.  192  i. 

176.  Mackenzie  Lamb,  I.  A  monograph  of  the  Lichen  genus  Placopsis  Nyl.  Lilloa  13, 

1947. 

177.  Massee,  G.  Lichenes.   The  Subantarctic  Islands  of  New  Zealand.  Wellington  1909. 

178.  Mattfeld,  J.  Compositae.  Nova  Guinea  14:4.  Leiden  1932. 

179.  Mayr,  E.  The  origin  and  history  of  the  Bird  Fauna  of  Polynesia.  Proceed.  6th  Pacif. 

Sci.  Congr.  1929,  IV.  Berkeley  1940. 

180.  Metraux,  a.  Ethnology  of  Easter  Island.  Bull.  Bishop  Mus.  Honolulu  1940. 

181.  Michaelsen,  W.  Oligochaeten  von  Juan  P>rnandez  und  der  Oster-Insel.  Nat.  Hist. 

Juan  Fern.  Ill,  192  i . 

182.  Montagne,  C.  Lfquenes,  in  Gay,  Hist.  fis.  y  polit.  de  Chile.  Botanica  8.  Paris  1852. 

183.  MuMFORD,  E.  P.  The  present  status  of  studies  of  faun  al  distribution  with  reference 

to  Oceanic  Islands.  Proceed.  6th  Pacif.  Sci.  Congr.  1939,  IV.  Berkeley  1940. 

184.  MiJLLER,  J.  Lichens.  Miss.  Scient.  du  Cap  Horn  1882-83.  V.  Paris  1889. 

185.  Navas,  L.  Neuropteres  des  lies  Juan  Fernandez  et  de  I'lle  de  Paques.  Nat.  Hist. 

Juan  Fern.  Ill,  1921. 

186.  NiLSSoN,  H.  Syntetische  Artbildung.  Lund  1953. 

187.  Nixon,  G.  E.  J.  Hymenoptera-Braconidae.  (insect.  J.  Fern.  26.)  Rev.  Chil.  de  Ento- 

mol. 4,  1955. 

188.  Nybelin,  O.   Nesophilaemon  n.  g.   fiir  Philaemon  skottsbergi  L.  Johansson.   Zool. 

Anzeiger  142,  1943. 


434 


C.  SKOTTSBERG 


189.  Odhnkr,  X.  Hj.  Mollusca  from  Juan  Fernandez  and  Easter  Island.  Nat.  Hist.  Juan 

Fern.  Ill,  1922.  Addenda,  ibid.  1926. 

190.  Oglobin,  a.  Mymaridac  (Hymenoptera).  (Insect.  J.  Fern.  1  2.)  Rev.  Chil.  de  Entoniol 

2,  1952. 
191. Bethylidae  y   Dryinidae  (Hymenoptera).  (Insect.  J.  Fern.  14.)  Ibid.  3,  1953 

192.  Oi.ivER,  W.  R.  B.  The  genus  Coprosma.  Bull.  Bishop  Mus.  Honolulu  132,  1935. 

193.  Pkrkins,  R.  C.  L.    Introduction,    being    a    review   of  the   land-fauna    of  Hawaiia 

Fauna  Hawaiiensis  I.  Cambridge  19  13. 

194.  Pfeiffkk,  H.  Oreobolus  R.  Br.,  eine  merkwiirdige  Cyperaceengattung.  Fedde,  Repert 

23,  1927. 

195.  Pic,  M.  Coleoptera-Anobiida  de  Juan   Fernandez.   Nat.  Hist.  Juan  Fern.  HI,  1924 
196. Coleoptera-Clavicornia  et  autres  de  Juan  Fernandez.  Ibid. 

197.   PiLGF.R,  R.   Uber  einige  Gramineae  der  Skottsbergschen  Sammlung  von  Juan  Fer 

nandez.  Fedde,  Repert.  16,  1920. 
198. *;*.    Das    System  der  Gramineae   unter   Ausschluss    der   Bambusoideae.    Bot 

Jahrb.  fur  System,  etc.  ed.  by  A.  Engler,  76,  1954. 

199.  Pi, .ATE,  L.  Zur  Kenntnis  der  Insel  Juan  Fernandez.   Verhandl.   Ges.  fiir  Erdkunde 

Berlin  XXIII,  1896. 

200.  Pope,  R.  D.  Coleoptera-Colydiidae.  (insect.  J.  Fern.  25.)  Rev.  Chil.  de  Entomol.  4, 

1955- 

201.  Princis,  K.  Uber  einige  neue  bzw.  wenig  bekannte  Blattarien.  Arkiv  f.  Zool.  41,  1948. 

202.  Ramiu),  B.  Historia  da  flora  do  planalto  riograndense.  Anais  Bot.  Barbosa  Rodriguez 

V,  1953- 

203.  Reiche,  K.  (irundziige  der  Pflanzenverbreitung  in  Chile.  Engler  &  Drude,  Vege- 

tation der  Erde  7.  Leipzig  1907. 

204.  Richards,  O.  W.  Sphaeroceridae.  (insect.  J.   P>rn.   21.)  Rev.  Chil.  de  Entomol.  4, 

^955- 

205.  Ridley,  H.  N.  The  Dispersal  of  Plants  throughout  the  World.  London  1930. 

206.  Roman,  A.  Ichneumoniden  von  Juan  Fernandez.  Nat,  Hist.  Juan  Fern.  HI,  1924. 

207.  RoMELL,  L.  Bacidiomycetes  from  Juan  Fernandez,  Ibid.  II,  1928. 

208.  Sahroskv,  C.  W.  A  new  species  of  Ogcodes  from  the  Juan  Fernandez  islands.  Rev. 

Chil.  de  ?>ntomol.  1,  1951. 
209. ■  Chloropidae.  (insect.  J.  I^ern.  19.)  Ibid.  4,  1955. 

210.  Sainshurv,  G.  O.  K.  a    Handbook  of  the  New  Zealand  Mosses.   Bull.   Roy.   Soc. 

of  N.  Zeal.  no.  5,  1955. 

211.  Saxtis,  L.  de.  Hymenoptera-Eulophidae,   Entedontidae  (Chalcidoidea  I).  (Insect.  J. 

Fern.  27.)  Rev.  Chil.  de  F'.ntomol.  4,  1955. 

212.  Satchell,  G.  H.  Psychodidae  (Diptera).  (insect.  J.  Fern.  8.)  Rev.  Chil.  de  Entomol. 

2,  1952. 

213.  Schenkling,  S.  Coleoptera-Cleridae  von  Juan  Fernandez.  Nat.  Hist.  Juan  Fern.  IH^ 

1931- 

214.  ScHMii),  F.  Trichoptera.  (insect.  J.  Fern.  4.)  Rev.  Chil.  de  Entomol.  2,  1952, 

215.  Schmidt,  A.  Coleoptera-Scarabaeidae  von  Juan  Fernandez.  Nat.  Hist.  Juan  Fern.  HI, 

1931- 

216.  ScHf)Tr,  H.  Collembola    aus  den  Juan  FernandezTnseln  und  der  Osterinsel.   Nat. 

Hist.  Juan  Fern.  Ill,  1921. 

217.  Setchell,  W.  A.  Les  migrations  des  oiseaux  et  la  dissemination  des  plantes.  Compte- 

rendu  sommaire  Soc.  de  Biogeogr.  22,  1926. 

218.  — — ■  Migration  and  endemism  with  reference  to  Pacific  Insular  floras.  Proceed.  3d 

Pacif.  Sci.  Congr.  1926.  Tokyo  1928. 

219. Pacific  Insular  floras  and  Pacific  paleogeography.  Amer.  Naturalist  69,  1935. 

220.   Seward,  A.  C.  A  study  in  contrasts.  The  present  and  past  distribution  of  certain 

ferns.  Linn.  Soc.  Journ.  of  Bot.  46,  1922. 


DERIVATION  OF  THE  FLORA  AND  FAUNA 


435 


22  1.  Seward,  A.  C.  &  Conway,  Verona.  A  phytogeographical  problem:  fossil  plants  from 
the  Kerguelen  Archipelago.  Ann.  of  Bot.  48,  1934. 

222.  SiLVESTRi,  F.  Thysanura  and  Embioptera.  Nat.  Hist.  Juan  Fern.  Ill,  1924. 

223.  Simpson,  G.  G.  Antarctica    as  a  faimal  migration  route.  Proceed.   6th   Pacif.   Sci. 

Congr.  1939.  Berkeley  1940. 

224.  SiNNOTT,  E.  W.  &  Bailey,  I.  W.  The  origin  and  dispersal  of  Herbaceous  Angio- 

sperms.  Ann.  of  Bot.  28,  1914. 

225.  SjosTEDT,  Y.  Odonata.  Orthoptera.  Nat.  Hist.  Juan  Fern.  Ill,  1924. 

226.  Skottsberg,  C.  Die  Gefasspflanzen  Sudgeorgiens.  Wiss.  Ergebn.  Schwed.  Sudpolar- 

Exp.  IV.  Stockholm  1905. 

227. Studien  uber  die  Vegetation  der  Juan  Fernandez-Inseln.  K.  Sv.  Vet.-akad. 

Handl.  51,  1914. 

228. Notes  on  the  relations  between  the  floras  of  Subantarctic  America  and  New 

Zealand.  Plant  World  18,  19 15. 
229. The  Phanerogams  of  the  Juan  Fernandez  Islands.  Nat.  Hist.  Juan  Fern.  II, 

192  2. 
230.   — —  The  Phanerogams  of  Easter  Island.  Ibid. 
231. Juan  Fernandez  and  Hawaii.  A  phytogeographical  discussion.  Bull.  Bishop 

Mus.  16.  Honolulu  1925. 
232.  — —  Einige  Bemerkungen  uber  die  alpinen  Gefasspflanzen  von  Masafuera.  Veroff". 

Geobot.  Inst.  Riibel  3.  Zurich  1925. 

233- Einige  Pflanzen  von  der  Oster-Insel.  Acta  Horti  Gotob.  3,  1927. 

234. Remarks  on  the  relative  independency  of  Pacific  Floras.  Proceed.  3d  Pacif. 

Sci.  Congr.  1926.  Tokyo  1928. 
235. Pollinationsbiologie   und  Samenverbreitung  auf  den  Juan  Fernandez-Inseln. 

Nat.  Hist.  Juan  Fern.  II,  1928. 
236. The  geographical  distribution  of  the  sandalwoods  and  its  significance.  Proceed. 

4th  Pacif.  Sci.  Congr.    1929.   III.   Batavia  1930. 
237. Notes  on  some  recent  collections  made  in  the  Island  of  Juan  Fernandez.  Acta 

Horti  Gotob.  4,  1929. 
238. Marine  algal  Communities  of  the  Juan  Fernandez  Islands.  Nat.  Hist.  Juan 

Fern.  II,  1941. 

239- I-e  peuplement  des  iles  pacifiques  du  Chili.  Soc.  de  Biogeogr.  IV,  1934. 

240.   • — ■ —  Greigia  Berteroi  Skottsb.   and  its  systematic  position.  Acta  Horti  Gotob.  11, 

1936. 
241. Die  Flora  der  Desventuradas-Inseln.  Goteb.  K.  Vet.  o.  Vitterh.  Samh.  Handl. 

5:6  foljden,  ser.  B,  bd  5:6,   1937. 
242. Geographical  isolation  as  a  factor  in  species  formation.  Proceed.  Linn.  Soc. 

of  Lond.,  Sess.  150,  1938. 
243. On  Mr.  C.  Bock's  collection  of  plants  from  Masatierra  (juan  Fernandez)  with 

remarks  on  the  flower  of  Centaurodendron.  Acta  Horti  Gotob.  12,  1938. 

244. Hawaiian  Vascular  Plants.  Ibid.  15,  1944. 

245. •  Peperomia  berteroana  Miq.  and  P.  tristanensis  Christoph.,  an  interesting  case 

of  disjunction.  Ibid.  16,  1947. 

246. Fine  kleine  Pflanzensammlung  von  San  Ambrosio.  Ibid.  17,  1947- 

247. The  genus  Peperomia  in  Chile.  Ibid. 

248. Biogeografiska  Stillahavsproblem.  Statens  naturv.  forskn.-rad,  arsbok  4  (i949~ 

50).  Stockholm  195  i. 
249. A  supplement  to  the  Pteridophytes  and  Phanerogams  of  Juan  Fernandez  and 

Easter  Island.  Nat.  Hist.  Juan  Fern.  II,  195  i. 
250. On  the  supposed  occurrence  of  Blechnum  longicauda  C.  Chr.  in  Brazil.  Svensk 

bot.  tidskr.  48,  1954. 
251. The  Vegetation  of  the  Juan  Fernandez  Islands.Nat.Hist.JuanFern.il,  1953. 


436  C.  SKOTTSBERG 

252.  Slkumer,  H.  Revision  der  Oattung   Pernettya.  Notizbl.   Botan.  Gartens  Berlin  12, 

1935- 

253.  Smith,  A.  C.  Taxonomic    notes   on  the  old  world  species  of  Winteraceae.   journ 

Arnold  Arbor.  24,  1943. 
254. The  American  species  of  Drimys.  Ibid. 

255.  Sorz.A  LopFs,  H.  iv  Alhuqufrque,  I).  Calliphoridae,  Sarcophagidae.  (Insect.  J.  Fern. 

22.)  Rev.  Chil.  de  Entomol.  4,  1955. 

256.  Si'AKRE,  B.  Review  of  Croizat,  Manual  of  Phytogeography.  Rev.  Universitaria  38, 

1053- 

257.  SrEP.HiNS,  G.  L.  JR.  Additional  evidence  for  a  holarctic  dispersal  of  flowering  plants 

in  the  Mesozoic  era.  Proceed.  6th  Pacif.  Sci.  Congr.  1939,  IV.  Berkeley  1941. 

258.  SiEEMs,  C.  G.  G.  J.  VAN.  On  the  origin  of  the  Malaysian  mountain  flora.  Bull.  Jard. 

bot.  lUiitenzorg,  Ser.  3,  13-14,  1935-36. 

259.  S'locKWELE,  P.  A  revision  of  the  genus  Stenactis.  Contrib.  Dudley  Herbar.  1940. 

260.  Straneo,  S.  L.  c^'  Jeaxnef,,  R.   Goleoptera-Carabidae.    (insect.  J.   Fern.    23.)  Rev. 

Chil.  de  Entomol.  4,  1955. 
261'.   SvENsoN,  H.  Monogra])hic  studies  in  the  genus  Eleocharis  V.  Rhodora  41,  1939. 

262.  SwEZEV,  O.  H.  Distribution  of  Lepidoptera  in   Pacific  Island  Groups.  Proceed.  6th 

Pacif.  Sci.  Congr.  1939,  IV.  Berkeley  1940. 

263.  Taveor,  \V,  B.  An  example  of  long  distance  dispersal.  Ecology  35,  1954. 

264.  Theriot,  J.  Mousses  recoltees  dans  Pile  Mas  a  Tierra  (Juan  Fernandez)  en  1927, 

par  M.  Gualterio  Looser.  Rev.  Chil.  de  Hist.  Nat.  31,  1927. 
265. Mousses  de  Pile  de  Paques.  Revue  bryol.  et  lichenol.  Ser.  II:  10,  1937. 

266.  Thompson,  G.  B.  Anoplura   from  Juan  Fernandez.  Nat.  Hist.  Juan  P'ern.  Ill,  1940. 

267.  Tonnoir,  a.  L.  Australian  Mycetophilidae.   Proceed.   Linn.   Soc.   N.   S.  Wales  54, 

1929. 

268.  'Prac.ardh,  I.  Acarina  from  the  Juan  P^rnandez  Islands.  Nat.  Hist.  Juan  Fern.  Ill, 

1931. 

269.  'PiRMEE,  J.  M.  Repartition  geographique  des  Eryngium  II.  Bull.  Mus.  Nat.  d'Hist. 

Nat.  Paris  1949. 

270.  'PuvAMA,  'P.  On  Santalum  boninense  and  the  distribution  of  the  species  of  Santalum. 

Journ.  Japan.  Bot.  15,  1939. 
271. •  On  genus  Haloragis  and  Micronesian  si)ecies.  Ibid.  16,  1940. 

272.  Ump.grove,  J.  H.  'P.  See  Continental  Drift. 

273.  IsiNCER,  R.  L.  Distribution  of  the   Heteroj)tera  from  Oceania.  Proceed.  6th  Pacif. 

Sci.  C'ongr.  1939,  IV.  Berkeley  1940. 

274.  Vkrhoekf,  K.  L'ber  Myriapoden  von  Juan  P>rnandez  und  der  Osterinsel.  Nat.  Hist. 

Juan  Fern.  Ill,  1924. 

275.  \'n)AL  (ioRMAZ,  F.  Geografia  naiitica  de  la  Republica  de  Chile.  Anuario  Hidrogrdf. 

de  la  Marina  de  Chile  7,  1881. 

276.  \'iERiE\ppER,  F.  Uber  echten  und  falschen  Vikarismus.  Osterr.  botan.  Zeitschr.  68, 

1919. 

277.  Wahri'.erc;,  R.  Einige  terrestre  Isoi)oden  von  den  Juan  Fernandez-Inseln.  Nat.  Hist. 

Juan  I'ern.  Ill,  192 2. 

278.  Waeeace,  a.  R.  Island  Life.  London  1880.  3d  ed.  191 1. 

279.  Weeks,  L.  (i.  Palcogeography  of  South  America.  Bull.  Geol.  Soc.  of  Amer.  59,  1948. 

280.  Wegener,  A.  Die  Entstehung  der  Kontinente  und   Ozeane.  Braunschweig.  3d  ed. 

1922,    5th   ed.    1936. 

281.  Weinl\r(  K,  H.  Studies  in  Juncaceae  with  special  reference  to  the  species  in  P^thiopia 

and  the  Cape.  Svensk  bot.  tidskr.  40,  1946. 

282.  Weise,  J.  Coleoptera-Chrysomelidae  and  Coccinellidae  von  Juan  Fernandez.   Nat. 

Hist.  Juan  Fern.  Ill,  1924. 

283.  Wheeler,  W.  M.  Formicidae  from  Piaster  Island  and  Juan  P'ernandez.  Ibid. 


DERIVATION  OF  THE  FLORA  AND  FAUNA  437 

284.  Willis,  B.  Geotectonics  of  the  Pacific.  Proceed.  3d  Pacif.  Sci.  Congr.  1926.  Tokyo 

1928. 

285.  Willis,  J.  C.  Endemic  genera  of  plants  in  their  relation  to  others.  Annals  of  Bot.  35, 

192  I . 
286. Age  and  Area.  A  Study  in  the  Origin  and  Distribution  of  Plants.  1922. 

287.  Winkler,  H.  Geographic,  in  Manual  of  Pteridology,  ed.  by  F.  Verdoorn.  The  Hague 

1938. 

288.  WiRTH,  W.  W.  Heleidae    and    Tendipedidae.    (insect.  J.  Fern.   7.)   Rev.  Chil.  de 

Entomol.  2,  1952. 
289. Ephydridae.  (insect.  J.  Fern.  20.)  Rev.  Chil.  de  Entomol.  4,  1955. 

290.  WooLDRiDGE.  See  Continental  Drift. 

291.  WuLFF,  E.  V.  An  Introduction  to  Historical  Plant  Geography.  Waltham,  Mass.  1943. 

292.  Wygodzinsky,  P.  Contribuci6n  al  conocimiento  del  genero  Metapterus  Costa  de  las 

Americas  y  de  Juan  Fernandez.  Rev.  Chil.  de  Entomol.  i,  195  i. 

293. Thysanura  from  Juan  Fernandez  Island.  Ibid. 

294. Simuliidae  (Diptera).  (insect.  J.  Fern.  6.)  Rev.  Chil.  de  Entomol.  2,  1952, 

295.   Zahlbruckner,  a.  Die    Flechten.   Bot.  Ergebn.  schwed.  Exped.   nach  Patagonien 

und  dem  Feuerlande.  VI.  K.  Sv.  Vet.-akad.  Handl.  57:6,  1917. 

296. Die  Flechten  der  Juan  Fernandez-Inseln.  Nat.  Hist.  Juan  Fern.  II,  1924. 

297. Die    Flechten    der   Osterinsel,  nebst  einem  Nachtrag  zur  Flechtenflora  von 

Juan  Fernandez.  Ibid.  1926. 

298.  Zimmerman,  E.  C.  Insects  of  Hawaii.  I.  Introduction.  Honolulu  1948. 

299.  ZiMMERMANN,  A.  Coleoptera-Dytiscidae  von  Juan  Fernandez  und  der  Osterinsel.  Nat. 

Hist.  Juan  Fern.  Ill,  1924. 

300.  Bryan,  W.  A.  in  "El  Mercurio",  Valparaiso,  Apr.  23,  1920. 

301.  Gourlay,  W.  B.  in  "The  South  Pacific  Mail",  Valparaiso,  Feb.  2,  1928. 

302.  QuENSEL,  P.  Die  Geologie  der  Juan  Fernandez-Inseln.  Bull.  Geol.  Inst.  Upsala  11, 

1913- 

303.  Branchi,  E.  C.  La  Isla  de  Robinson.  Valparaiso    1922. 

304.  Hayek,  A.  Allgemeine  Pflanzengeographie.  Berlin    1926. 

305.  Drude,  O.  Handbuch  der  Pflanzengeographie.  Stuttgart   1890. 

306.  Merrill,  E.  D.  The  Botany  of  Cook's  Voyages.  Chronica  Botanica  14:  5/6.  Waltham, 

Mass.    1954. 

307.  Hillebrand,  W.  The  Flora  of  the  Hawaiian  Islands.  Heidelberg  1888. 

308.  Fleutiaux,  E.  Coleoptera-Serricornia  de  Juan  Fernandez  et  de  Pile  de  Paques.  Nat. 

Hist.  Juan   Fern.  Ill,  1924. 

309.  ScHEDL,  K.  E.  Chilenische  Borkenkafer.  Rev.  Chil.  de  Entomol.  4,  1955. 

310.  Wygodzinsky,  P.  Reduviidae  y  Cimicidae  (Hemiptera).  (Insect.  J.  Fern.  2.)  Rev. 

Chil.  de  Entomol.  2,  1952. 

311.  Arwidsson,  Th.  Einige  parasitische  Pilze  aus  Juan  P'ernandez  und  der  Osterinsel. 

Svensk  bot.  tidskr.  34,  1940. 

312.  Halle,  T.  G.    A    fossil  fertile  Lygodium  from  the  Tertiary  of  South  Chile.   Ibid. 

313.  Malta,  N.  Die  Gattung  Zygodon  Hook,  et  Tayl.  Latv.  Univ.  Botan.  Darza  Darbi 

n.  I.  Riga  1926. 

314.  Fennah,  R.  G.  Homoptera-Delphacidae.  Proceed.  Entomol.  Soc.  of  London,  Ser.  B 

24,  1955- 

315.  Berry,  E.  W.  Fossil  plants  from  Chubut  Territory  collected  by  the  Scarritt  Patagonian 

Expedition.  Amer.  Mus.  Novit.  536,  1932. 

316.  Brinck,  p.  Coleoptera  of  Tristan  da  Cunha.  Results  Norweg.  Scient.  Exped.  1937-38 

no.  17.  Oslo  1948, 

317.  Werth,  E.  Die  Vegetation  der  subantarktischen  Inseln  Kerguelen,  Possession  und 

Heard-Eiland  II.  Deutsche  Sudpolar-Exped.  1901-02.  Bd.  VIII.  1911. 

318.  Skottsberg,  C.  The  Flora  of  the  Hawaiian  Islands  and  the  History  of  the  Pacific 

Basin.  Proceed.  6th  Pacif.  Sci.  Congr.  1939,  IV.  Berkeley  1940. 


438  C.  SKOTTSBERG 

31Q.   Stebbins,  G.  L.  jr.  Variation  and  Evolution  in  Plants.  New  York  1950. 

320.  LiNDHERG,  H.  De  bortglomda  oarna.  Helsingfors  1955. 

321.  GiBBS,  Lilian.  A  Contribution  to  the  Montane  Flora  of  Fiji.  Journ.  Linn.  Soc.  Bot. 

39,  19C9. 

322.  ViSHKR,  S.  S.  Tropical  cyclones  of  the  Pacific.  Bernice  P.  Bishop  Mus.  Bull.  20,  1925. 
323. Tropical  cyclones  and  the  dispersal  of  life  from  island  to  island  in  the  Pacific. 

Amer.  Naturalist  59,  1925. 

324.  Bergeron,  T.   The  problem  of  tropical  hurricanes.  Journ.  R.  Meteor.  Soc.  80:  344. 

1950. 

325.  Grisebach,  a.  Die  Vegetation  der  Erde.    2d  ed.  Leipzig.   I,  1872,  11,  1884. 

326.  Skottsberg,  C.   On  Scirpus  nodosus  Rottb.   Acta  Soc.  pro  Fauna  et  Flora  Fenn. 

72,  1956. 

327.  Santesson,  R.  The  South  American  Cladinae.  Arkiv  for  Bot.  30,  1942. 


—  The  South  American  Menegazziae.   Ibid. 


329.  • Contributions  to  the  Lichen  Flora  of  South  America.   Ibid.   31,   1944. 

330.  JoHOw,  F.   Ueber  die  Resultate  der  Expedition  nach  den  Islas  Desventuradas  (San 

Ambrosio  und  San  Felix).  Verhandl.   d.  deutsch,  wissensch.  Vereins  zu  San- 
tiago  IIL  Valparaiso  1898. 

331.  Bruggen,  J.  Fundamentos  de  la  Geologia  de  Chile.   Santiago    1950. 

332.  Martin,  C.   Landeskunde  von  Chile.   Hamburg    1909. 

;^^;^.   Stephani,  F.  Bot.  Ergebn.  schwed.  Exped.  nach  Patagonien  und  dem  Feuerlande. 

2.  Die  Lebermoose.  K.  Sv.  Vet.-akad.  Handl.  46,  191 1. 
334.   I^ERRV,  E.  W.  Tertiary    fossil  plants  from  the  Argentine  Republic.   Proceed.   U.S. 

Nation.  Mus.  73,  1928. 
335. A  Miocene  flora  from   Patagonia.  Johns   Hopkins  Univ.   Stud.,  Geol.  no.  6, 

1925.   Not  seen,   only  a  preliminary  notice  in  Proceed.  Nat.   Acad,  of  Sci. 

11:  7,    1925. 
336.   Frenguelli,  J.    Recientes    progresos   en   el  conocimiento  de  la  geologia  y  paleo- 

geograffa  de  Patagonia.   Rev.  Mus.  de  Eva  Peron  (La  Plata),  N.S.  IV,  Geol., 

1953- 

337. l>a  Flora  f6sil   de  la  regi6n  del  alto   Rio  Chalia  en  Santa  Cruz  (Argentina). 

Notas  del  Mus.   de  Eva  Per6n   (La  Plata)  XVL  98,    1953. 

338.  Florin,  R.    Die  heutige   und  friihere  Verbreitung  der  Gattung  Acmopyle  Pilger. 

Svensk   bot.  tidskr.  34,  1940. 

339.  Hooker,  J.  D.  Observations  on  the  Botany  of  Kerguelen  Island.  Philos.  Trans.  Roy. 

Soc.  of  London  168,    1879. 

340.  Skottsberg,  C.  Antarctic  Plants  in  Polynesia.   Essays  in   Geobotany  in  honor  of 

W.  A.   Setchell.   Berkeley    1936. 
341. Plant   succession  on  recent  lava  flows  in  the  island  of  Hawaii.   Goteb.  K. 

Vet.  o.  Vitterh.  Samh.  Handl.,  6:e  foljden,  ser.  B,  bd  1:8,  1941. 
342. The    Vegetation  of  Easter  Island.  Nat.  Hist.  Juan  Fern.  II,  1927. 

343.  Cooke,  (i.  H.   Te  pito  te  henua,  known  as  Rapa  Nui.   Rep.  Smithson.  Inst.  1897. 

Rep.  U.S.  National  Mus.  Part  I.   Washington  1899. 

344.  MoTVKA,  J.  Lichenum  generis  Usnea  studium  monographicum.  Lw6w  1938. 

345.  Brigham,  W.  T.  An  index  to  the  islands  of  the  Pacific  ocean.  Mem.  Bishop  Museum 

Vol.  I:  2.  Honolulu  1900. 

346.  Forster,  (r.  Florulae  insularum  australium  Prodromus.  Goettingen  1776. 

347. A   Voyage   round  the  world  .  .  .  commanded  by  Capt.  James  Cook  1772,  3,  4 

and  5.  Vol.  I.  London  1777. 
348. De  plantis  esculentis  insularum  oceani  australis  Commentatio  botanica.  Berlin 

1786. 


Contents. 

Part  I.  The  Juan  Fernandez  Islands. 

Chapter  I.   Composition,   distribution  and  relationships  of  the  Flora      193 

Chapter  II.   Sources  of  the  island  flora  as  judged  by  the  total  distribution  of  the 
geographical  elements,  with  special  reference  to  the  composition  of  the  Chilean 

flora 256 

Chapter  III.  Composition,   distribution  and  relationships  of  the  Fauna 292 

Chapter  IV,   Continental  and  Oceanic  islands 317 

Chapter  V.  The  Pacific  Ocean  and  Continental  Drift      325 

Chapter  VI.  Transoceanic  migration 331 

Chapter  VII.  Biological  characteristics  of  isolated  islands      35 1 

Chapter  VIII,  Evolution  in  Oceanic  islands      363 

Chapter  IX.  Juan  Fernandez — oceanic  or  continental? 372 

Chapter  X.  The  Chilean  coast  line  and  the  history  of  the  Andes 380 

Chapter  XI.  The  Tertiary  floras  of  Chile  and  Patagonia 386 

Chapter  XII.  Antarctica  as  a  source  of  the  present  circumpolar  floras 389 

Chapter  XIII.  The  history  of  Juan  Fernandez — a  tentative  sketch 394 

Part  2.  Easter  Island. 

Chapter  XIV.   Composition,   distribution  and  relationships  of  the  Flora 406 

Chapter  XV.  Composition,   distribution  and  relationships  of  the  Fauna 417 

Chapter  XVI.  The  biogeographical  history  of  Easter  Island 421 

Bibliography      428 

Printed  October  26th,  1956. 


/ 


P^L 


^»'  \-C 


QH 
198 
J8S$5 
v.l 

BioMed 


Skottsberg,   Carl  Johan  Frederik 

The  natural  history  of  Juan 
Fernandez  and  Easter  Island. 


PLEASE  DO  NOT  REMOVE 
CARDS  OR  SLIPS  FROM  THIS  POCKET 


UNIVERSITY  OF  TORONTO  LIBRARY