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U.  S.  DEPARTMENT   OF   AGRICULTURE 

BUREAU   OF   BIOLOGICAL   SURVEY 

K.    \V.  NELSON,   Chief 


NORTH  AMERICAN  FAUNA 

No.   41 

[Actual  date  of  publication,  February  9,  1918] 


REVIEW  OF  THE  GRIZZLY  AND  BIG  BROWN 
BEARS  OF  NORTH  AMERICA 

(GENUS  URSUS) 

WITH  DESCRIPTION  OF  A  iNEW  GENUS,  VETULARCTOS 


C.  HART    MERRIAM 

CONSULTING   BIOLOGIST,  BIOLOGICAL   SURVEY 
RESEARCH   ASSOCIATE,  SMITHSONIAN   INSTITUTION 


•  / 


WASHINGTON 

GOVERNMENT  URIXTINi;    OFFICE 

1918 


NORTH  AMERICAN  FAUNAS. 

Copies  of  North  American  Faunas  not  out  of  print  are  for  sale,  at  the  prices 
named,  by  the  Superintendent  of  Documents,  Government  Printing  Office,  Wash- 
ington, D.  C. 

No.  1.  Revision  of  the  North  American  Pocket  Mice.  By  C.  Hart  Merriam.  Pp. 
36,  pis.  4.     1S89 Price,  10  cents. 

No.  2.  Descriptions  of  Fourteen  New  Species  and  One  New  Genus  of  North 
American  Mammals.    By  C.  Hart  Merriam.    Pp.  52,  pis.  8,  figs.  7.    1889. 

Price,  10  cents. 

No.  3.  Results  of  a  Biological  Survey  of  the  San  Francisco  Mountain  Region 
and  Desert  of  the  Little  Colorado,  Arizona.  By  C.  Hart  Merriam  and 
Leonhard  Stejneger.    Pp.  136,  pis.  14,  maps  5  (colored),  figs.  2.    1890. 

Price,  25  cents. 

No.  4.  Description  of  Twenty-six  New  Species  of  North  American  Mammals. 
By  C.  Hart  Merriam.    Pp.  60,  pis.  3,  figs.  3.    1890 Price,  10  cents. 

No.  5.  Results  of  a  Biological  Reconnaissance  of  South-Central  Idaho.  By 
C.  Hart  Merriam  and  Leonhard  Stejneger.  Descriptions  of  a  New  Genus  and 
Two  New  Species  of  North  American  Mammals.  By  C.  Hart  Merriam.  Pp. 
132,  pis.  4  (1  colored),  figs.  4.    1891 Price,  15  cents. 

No.  6.  Not  issued. 

No.  7.  The  Death  Valley  Expedition :  A  Biological  Survey  of  Parts  of  Cali- 
fornia, Nevada,  Arizona,  and  Utah.  Part  II. — 1.  Birds,  by  A.  K.  Fisher. 
2.  Reptiles  and  Batrachians,  by  Leonhard  Stejneger.  3.  Fishes,  by  Charles 
H.  Gilbert.  4.  Insects,  by  C.  V.  Riley.  5.  Mollusks,  by  R.  E.  C.  Stearns. 
6.  Desert  Trees  and  Shrubs,  by  C.  Hart  Merriam.  7.  Desert  Cactuses  and 
Yuccas,  by  C.  Hart  Merriam.  8.  List  of  Localities,  by  T.  S.  Palmer.  Pp.  402, 
pis.  15,  maps  5,  figs.  2.    1893 [Out  of  print.] 

No.  8.  Monographic  Revision  of  the  Pocket  Gophers,  Family  Geomyidse  (exclu- 
sive of  the  species  of  Thomomys) .  By  C.  Hart  Merriam.  Pp.  258,  pis.  20, 
figs.  71,  maps  4  (colored).    1895 Price, 35 cents. 

No.  9.  Not  issued.  ) 

No.  10.  Revision  of  the  Shrews  of  the  American  Genera  Biarina  and  Notiosorex. 
By  C.  Hart  Merriam.  The  Long-tailed  Shrews  of  the  Eastern  United  States. 
By  Gerrit  S.  Miller,  jr.  Synopsis  of  the  American  Shrews  of  the  Genus 
Sorex.    By  C.  Hart  Merriam.    Pp.  124,  pis.  12,  figs.  3.    1895 Price,  15  cents. 

No.  11.  Synopsis  of  the  Weasels  of  North  America.  By  C.  Hart  Merriam.  Pp. 
44,  pis.  6,  tigs.  16.    1S96 *_ Price,  10  cents. 

No.  12.  The  Genera  and  Subgenera  of  Voles  and  Lemmings.  By  Gerrit  S. 
Miller,  jr.     Pp.  84,  pis.  3,  figs.  40.     1896 Price,  10  cents. 

No.  13.  Revision  of  the  North  American  Bats  of  the  Family  Vespertilionidae. 
By  Gerrit  S.  Miller,  jr.    Pp.  140,  pis.  3,  figs.  40.    1897 Price,  10  cents. 

No.  14.  Natural  History  of  the  Tres  Marias  Islands,  Mexico.  Prepared  under 
the  direction  of  C.  Hart  Merriam.  General  Account  of  the  Islands,  with  Re- 
ports on  Mammals  and  Birds,  by  E.  W.  Nelson.  Reptiles,  by  Leonhard 
Stejneger.  Notes  on  Crustacea,  by  Mary  J.  Rathbun.  Plants,  by  J.  N.  Rose. 
Bibliography,  by  E.  W.  Nelson.    Pp.  97,  pi.  (map),  figs.  2.    1899. 

Price,  10  cents. 

No.  15.  Revision  of  the  Jumping  Mice  of  the  Genus  Zapus.  By  Edward  A. 
Preble.    Pp.  42,  pi.,  figs.  4.    1899 Price,  5  cents. 

No.  16.  Results  of  a  Biological  Survey  of  Mount  Shasta,  California.  By  C.  Hart 
Merriam.    Pp.  179,  pis.  5,  figs.  46.    1899 [Out  of  print.} 

No.  17.  Revision  of  American  Voles  of  the  Genus  Microtus.  By  Vernon  Bailey. 
Pp.  88,  pis.  5-  figs.  17.     1900 [Out  of  print.  I 

No.  18.  Revision  of  the  Pocket  Mice  of  the  Genus  Perognathus.    By  Wilfred  H. 

Osgood.     Pp.  72.  pis.  4  (inch  2  maps),  figs.  15.     1900 Price,  10  cents. 

(Continued  on  page  3.  of  cover.) 


North  American  Fauna  No.  41.  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  I. 


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U.  S.  DEPARTMENT  OF  AGRICULTURE 

BUREAU  OF  BIOLOGICAL  SURVEY 

E.   W.  NELSON,   Chief 


NORTH  AMERICAN  FAUNA 

No.   41 

[Actual  date  of  publication,  February  9,  1918] 


REVIEW  OF  THE  GRIZZLY  AND  BIG  BROWN 
BEARS  OF  NORTH  AMERICA 

(GENUS  URSUS) 

WITH  DESCRIPTION  OF  A  NEW  GENUS,  VETULARCTOS 


C.  HART    MERRIAM 

CONSULTING  BIOLOGIST,  BIOLOGICAL  SURVEY 
RESEARCH  ASSOCIATE,  SMITHSONIAN  INSTITUTION 


WASHINGTON 

GOVERNMENT  PRINTING  OFFICE 

1918 


/  t/6 


LETTER  OF  TRANSMITTAL. 


United  States  Department  or  Agriculture, 

Bureau  of  Biological  Survey, 
/  Washington,  D.  C,  July  18, 1917. 
Sir  :  I  have  the  honor  to  transmit  for  publication  as  North  Ameri- 
can Fauna  No.  41  a  review  of  the  grizzly  and  big  brown  bears  of 
North  America,  by  Dr.  C.  Hart  Merriam,  consulting  biologist  and 
former  chief  of  the  Biological  Survey  and  research  associate  of  the 
Smithsonian  Institution.  This  review  was  prepared  and  originally 
submitted  in  September,  1916,  but  before  composition  was  begun 
it  was  recalled  from  the  printer  in  order  that  additional  informa- 
tion, developed  by  the  discovery  of  new  material,  might  be  included. 
The  work  is  based  largely  upon  material  in  the  collection  of  the 
Biological  Survey.  Up  to  20  years  ago  only  8  species  of  grizzly 
and  big  brown  bears  were  known,  but  since  then,  largely  through 
the  investigations  of  Dr.  Merriam,  the  number  of  recognizable  forms 
has  increased  to  86.  Additional  study  and  material  may  solve  cer- 
tain points  now  in  doubt,  but  it  is  not  deemed  advisable  to  delay 
further  the  publication  of  our  present  state  of  knowledge  of  this 
group  of  America's  historic  big  game  animals,  now  vanished  from 
great  stretches  of  their  former  domain.  This  review  will  be  of 
material  assistance  to  students  and  others  interested  in  our  native 
wild  life,  past  and  present. 
Respectfully, 

E.  W.  Nelson, 
Chief,  Biological  Survey. 
Hon.  Davto  F.  Hpu?ton^  ,     r  o«    :   .«.  :    . 

Secretary  &f,Agnb&1iWe,:  I  ..; 
2 


CONTENTS. 


Introduction •. 7 

Geographic  distribution 9 

Sexual  differences 10 

Age  differences 10 

Material  examined 10 

Technical  terms 11 

Classification  of  grizzly  and  big  brown  bears 12 

Relative  values  of  cranial  and  dental  characters 13 

List  of  species  and  subspecies,  with  type  localities 14 

Descriptions  of  species  and  subspecies 17 

Horribilis  group 17 

Planiceps  group 34 

Arizonx  group 53 

Hylodromus  group 77 

Horrixus  group 84 

Stikeenensis  group 88 

Alascensis  group 94 

Richardsoni  group 99 

Kidderi  group 106 

Innuitus  group 110 

Townsendi  group 115 

Dalli  group 116 

Gyas  group 124 

Kenaiensis  group 127 

Vetularctos,  a  new  genus  related  to  Ursut 131 

Index 135 

3 


ILLUSTRATIONS. 


PLATES. 

[Plate  I,  frontispiece;  Plates  II-XVI  following  page  134.] 

Plate  I.  Wild  grizzly  in  northwestern  Wyoming.    From  photograph  by  Frederick 
K.  Vreeland. 
II.  Skull  of  Ursus  gyas,  old  S ,  from  Bear  Bay,  Alaska  Peninsula.    No.  91690. 

III.  Skull  of   Ursus  middendorffi,  old   S,  from  Chiniak  Point,  Kodiak  Island, 

Alaska.    No.  96509. 

IV.  Skull  of    Ursus  kenaiensis,  old    d1,  from  Kenai  Peninsula,  Alaska.    No. 

210291. 
V.  Skull  of  Ursus  sheldoni,  adult  c?,  from  Montague  Island,  Prince  William 
Sound,  Alaska.    No.  137318  (type). 
VI.  Skull  of  Ursus  shirasi,  old  <?,  from  Admiralty  Island,  Alaska.     No.  203030 

(type). 
VII.  Skull  of  Ursus  innuitus,  old   <?,  from  Golofnin  Bay,  Alaska.    No.  179780 

(type). 
VIII.  Skull  of  Ursus  kidderi  kidderi,  old   <?,  from  Belkofski,  Alaska  Peninsula. 
No.  91698. 
IX.  Skull  of  Ursus  stikeenensis,  adult  <? ,  from  Tatletuey  Lake,  near  head  of 

Skeena  River,  British  Columbia.    No.  202794  (type). 
X.  Skull  of  Ursus  ophrus,  old  cf ,  from  eastern  British  Columbia.    No.  210252 

(type). 
XI.  Skull  of  Ursus  rungiusi  rungiusi,  old   d\  from  Indian  Point  Creek,  near 

Barkerville,  British  Columbia.    No.  209899. 
XII.  Skull  of  Ursus  alascensi-s,  old  d,  from  Unalaklik  River,  Alaska.    No.  76466 
(type). 

XIII.  Skull  of  Ursus  nortoni,  old  9  ,  from  Yakutat,  Alaska.    No.  178763  (type). 

XIV.  Skull  of   Ursus  horribilis  horribilis,  old  c? ,  from  Missouri  Breaks,  eastern 

Montana.     No.  202739. 
XV.  Skull  of  Ursus  horriseus,  old  <$ ,  from  Coppermines,  New  Mexico    No.  990 

(type). 
XVI.  Skull  of  Ursus  arizonse,  adult  <? ,  from  Escudilla  Mountains,  Apache  County, 
Arizona.     No.  177332  (type). 

[The  plates  of  skulls,  owing  to  restriction  of  space,  are  limited  to  the  side  view,  and 
consequently  in  some  cases  are  misleading.  Skulls  having  similar  profiles  often  differ  sur- 
prisingly when  viewed  from  above  or  below,  as  would  be  seen  at  a  glance  were  it  prac- 
ticable to  give  two  views.  J 

& 


No.  41.  NORTH  AMERICAN  FAUNA.         Feb.,  1918. 

REVIEW  OF  THE  GRIZZLY  AND  BIG  BROWN 
BEARS  OF  NORTH  AMERICA 

(GENUS  URSUS) 
WITH  DESCRIPTION  OF  A  NEW  GENUS,  VETULARCTOS. 


By  C.  Hart  Merbiam. 


INTRODUCTION. 

When  Audubon  and  Bachman  published  their  great  work  on  the 
Mammals  of  North  America  (1846-1854),  and  in  fact  up  to  the  year 
1857,  it  was  commonly  believed  by  naturalists  as  well  as  by  hunters 
and  the  public  generally  that  there  was  only  a  single  species  of 
grizzly  bear — the  one  described  by  Lewis  and  Clark  in  1804-5,  and 
named  Ursus  horribilis  by  Ord  in  1815.  Baird,  in  1857,  described 
another  species,  from  Coppermines,  New  Mexico,  which  he  named 
Ursus  horriceus. 

Nearly  40  years  later,  in  my  "  Preliminary  Synopsis  of  the  Ameri- 
can Bears,"1  eight  grizzlies  and  big  brown  bears  were  recognized, 
of  which  five  were  described  as  new.  It  was  not  then  suspected 
that  the  number  remaining  to  be  discovered  was  anything  like  so 
great  as  has  since  proved  to  be  the  case.  The  steady  influx  of  speci- 
mens resulting  from  the  labors  of  the  Biological  Survey,  supple- 
mented by  the  personal  efforts  of  a  number  of  hunter-naturalists, 
brought  to  light  many  surprises,  most  of  which  have  been  published ; 
and  beginning  in  the  spring  of  1910,  a  fund  placed  at  my  disposal 
made  it  possible  to  offer  hunters  and  trappers  sufficient  inducement 
to  tempt  them  to  exert  themselves  in  securing  needed  specimens.  As 
a  result,  the  national  collection  of  bears  has  steadily  grown  until,  in 
number  of  species  represented,  in  completeness  of  series,  and  in  num- 
ber of  type  specimens,  it  now  far  excels  all  other  collections  in  the 
world  together.     Nevertheless  there  are  many  gaps  in  the  series. 

1  Proc.  Biol.  Soc.  Washington,  X,  pp.  65-S3,  AprU  13,  1896. 


8  NORTH   AMERICAN  FAUNA.  [No.  41. 

Knowledge  of  the  big  bears  is  by  no  means  complete  and  many  years 
must  pass  before  the  last  word  on  the  subject  will  be  written.  Many 
bears  now  roaming  the  wilds  will  have  to  be  killed  and  their  skulls 
and  skins  sent  to  museums  before  their  characters  and  variations  will 
be  fully  understood  and  before  it  will  be  possible  to  construct  accu- 
rate maps  of  their  ranges.  Persons  having  the  means  and  ambition 
to  hunt  big  game  may  be  assured  that  bears  are  still  common  in  many 
parts  of  British  Columbia,  Yukon  Territory,  and  Alaska,  and  that 
much  additional  material  is  absolutely  required  to  settle  questions 
still  in  doubt. 

Among  the  localities  from  which  specimens  are  greatly  needed 
may  be  mentioned  Lynn  Canal  and  Lituya  Bay,  Alaska,  and  in  fact 
the  entire  coast  strip  between  Cross  Sound  and  Yakutat  Bay;  the 
Cook  Inlet  and  Susitna  regions;  the  mountains  between  the  Yukon 
and  Tanana;  the  Endicott  Range  and  other  mountains  between  the 
Yukon  and  the  Arctic  coast  all  the  way  from  Seward  Peninsula 
to  the  Alaska- Yukon  boundary;  the  Rocky  Mountains  of  Canada, 
from  British  Columbia  northward,  including  the  sources  of  the 
Pelly,  Macmillan,  Stewart,  and  Porcupine  Rivers;  the  Mackenzie 
River  and  Great  Bear  Lake  region;  the  southwest  corner  of  Yukon 
Territory;  the  western  part  of  Alberta;  and  the  interior  of  British 
Columbia.  In  the  United  States,  skulls  of  adult  males  are  much 
needed  from  all  localities  inhabited  by  grizzly  bears,  particularly  in 
Colorado,  Idaho,  Montana,  and  Wyoming — including  the  Glacier  and 
other  National  Park  regions. 

What  is  most  needed  to-day  in  this  line  is  a  series  of  adults  col- 
lected by  absolutely  trustworthy  persons  and  labeled  on  the  spot 
for  locality  and  sex.  Many  specimens  in  museums  are  not  labeled 
for  sex ;  others  have  the  sex  wrongly  marked ;  and  many  either  lack 
localities  or  the  localities  given  are  open  to  serious  doubt.  A  speci- 
men is  of  little  value  unless  one  can  pin  his  faith  on  the  label. 

Some  writers  have  advanced  the  view  that  the  various  species  of 
bears  freely  interbreed.  Let  those  so  minded  ask  themselves  the 
question,  If  promiscuous  interbreeding  were  to  take  place,  what 
would  become  of  the  species?  From  the  nature  of  the  case,  the  sta- 
bility of  species  depends  on  the  rarity  of  crossings  with  other  species, 
for  if  interbreeding  were  to  take  place  frequently  the  species  so  inter- 
breeding would  of  course  cease  to  exist,  having  merged  into  a  com- 
mon hybrid.  Hybrids  now  and  then  occur,  particularly  in  zoological 
gardens,  but  among  wild  animals  in  their  native  haunts  they  are  ex- 
ceedingly rare. 

The  number  of  species  here  given  will  appear  to  many  as  preposter- 
ous.   To  all  such  I  extend  a  cordial  invitation  to  visit  the  National 


1918.]  INTKODUCTION.  9 

Museum  and  see  for  themselves  what  the  bear  skulls  show.  Recog- 
nition of  species  is  a  matter  of  interpretation.  If  the  material  is 
adequate  there  can  be  little  room  for  difference  of  opinion;  if  in- 
adequate, many  important  points  must  remain  in  doubt.  It  is  not 
the  business  of  the  naturalist  either  to  create  or  to  suppress  species, 
but  to  endeavor  to  ascertain  how  many  Nature  has  established,  and 
having  discovered  this,  to  point  out  their  characters  and  learn  as 
much  as  possible  about  them. 

One  of  the  unlooked-for  results  of  the  critical  study  of  the  Ameri- 
can bears  is  the  discovery  that  the  big  bears,  like  mice  and  other  small 
mammals,  split  up  into  a  large  number  of  forms  whose  ranges  in  some 
cases  overlap  so  that  three  or  more  species  may  be  found  in  the  same 
region. 

Another  surprising  result  is  the  discovery  that  Admiralty  Island  in 
Southeastern  Alaska  appears  to  be  inhabited  by  no  less  than  five 
distinct  species,  each  of  which  is  obviously  related  to  and  representa- 
tive of  an  adjacent  mainland  species.  The  recognition  of  this  very 
remarkable  state  of  affairs  makes  it  possible  to  understand  what 
before  had  seemed  a  most  anomalous  condition,  namely,  the  extraor- 
dinary diversity  or  variability  of  the  skulls  and  teeth  of  the  island 
bears.  It  was  not  until  material  essential  for  the  determination  of 
the  mainland  forms  had  been  collected  that  it  was  possible  to  recog- 
nize and  define  the  island  forms. 

The  varying  degrees  of  divergence  of  the  island  bears  furnish  an 
interesting  index  to  the  relative  time  when  each  obtained  a  foothold 
on  the  island.  In  this  connection  it  is  well  to  remember  that  the 
breadth  of  the  strait  separating  Admiralty  Island  from  the  main- 
land at  its  narrowest  point  does  not  exceed  5  miles. 

The  mainland  big  bears  with  their  representatives  on  Admiralty 
Island  here  provisionally  recognized  are : 

Mainland  Species.  Admiralty  Island  Species. 

Vrsu8  dalli Ursus  shirasi. 

Ursus  stikeenensis Ursus  mirabilis. 

Vrsus  tahltanicus Ursus  insularis. 

Ursus  ktoakiutl Ursus  neglectus. 

Ursus  caurinus Ursus   eulophus. 

GEOGRAPHIC   DISTRIBUTION. 

In  early  days  grizzly  bears  were  common  in  most  parts  of  west- 
ern North  America,  their  range  being  nearly  continuous  from  north- 
ern Mexico  northward  through  the  Western  States  and  western 
Canada  to  northern  Alaska;  but  now  within  the  United  States  sev- 


10  NORTH  AMERICAN   FAUNA.  [No.  41. 

eral  of  the  species  are  extinct,  and  those  still  living  are  confined  in 
the  main  to  remote  or  inaccessible  mountain  ranges,  where  two  or 
more  species  not  infrequently  occur  together.  In  many  cases  the 
original  distribution  areas  overlapped,  as  they  do  to-day  in  parts  of 
British  Columbia,  Yukon,  and  Alaska;  in  other  cases,  owing  to  the 
settlement  of  the  country,  species  inhabiting  easily  accessible  areas 
were  either  exterminated  or  forced  into  mountains  where  they  now 
occupy  the  same  ground  with  other  species,  so  that  it  is  impossible  to 
ascertain  what  the  original  distribution  was.  As  a  rule,  in  cases  where 
two  or  more  species  inhabit  the  same  area,  the  species  occurring  to- 
gether belong  to  different  superspecific  groups.  Thus  in  the  Yellow- 
stone Park  and  Stikine  River  regions  representatives  of  the  kor- 
ribilis,  absarokus,  tahltanicus,  and  chelan  groups  occur,  so  far  as 
known,  in  the  same  localities. 

SEXUAL  DIFFERENCES. 

In  most  species  of  bears  the  males  are  much  larger  than  the  fe- 
males. In  some  the  disparity  in  size  is  very  remarkable,  as  in  mid- 
dendorffi  of  Kodiak  Island  and  magister  of  southern  California. 
In  a  few  cases  the  difference  is  slight,  as  in  kidderi  of  Alaska 
Peninsula. 

AGE   DIFFERENCES. 

Bear  skulls  undergo  a  series  of  changes  from  early  life  to  old  age, 
and  in  most  species  do  not  attain  their  mature  form  until  seven  or 
more  years  of  age.  In  species  having  the  frontal  shield  highly  ele- 
vated, as  in  middendorfji,  kluane,  stikeenensis,  and  mi?'abHis,  the  f  ron- 
tals  reach  their  maximum  of  arching  or  bulging  in  early  adult  life 
(about  the  sixth  year),  after  which  they  gradually  become  flatter. 

MATERIAL    EXAMINED. 

It  is  a  pleasure  to  express  appreciation  of  the  invaluable  assistance 
rendered  by  the  loan  or  presentation  of  skins  and  skulls  of  grizzly 
and  brown  bears  utilized  in  the  preparation  of  the  accompanying 
descriptions.  To  Charles  Sheldon,  G.  Frederick  Norton,  and  the 
late  Charles  R.  Cross,  jr.,  and  to  J.  H.  Kidder,  Robert  P.  Blake, 
Waldo  Emerson  Forbes,  George  Mixter,  Samuel  Mixter,  Dr.  William 
Jason  Mixter,  Charles  S.  King,  and  Homer  E.  Sargent,  special  thanks 
are  due  for  their  generosity  in  placing  at  my  disposal  the  valuable 
specimens  and  notes  obtained  on  their  private  hunting  trips.  Most 
of  these  specimens  have  been  presented  to  the  national  collections. 
And  to  Miss  Annie  M.  Alexander,  of  Oakland,  California,  special 
acknowledgment  should  be  made  for  the  loan  of  her  collection  of 


1918.]  INTRODUCTION.  11 

Alaska  bears,  now  in  the  Museum  of  Vertebrate  Zoology  of  the  Uni- 
versity of  California,  and  second  only  to  that  of  the  Biological 
Survey  and  National  Museum.1 

In  conclusion  it  is  only  proper  to  state  that  the  material  on  which 
the  present  publication  is  based  could  not  have  been  brought  together 
during  my  lifetime,  nor  the  results  prepared  for  publication,  but  for 
the  generous  assistance  of  Mrs.  E.  H.  Harriman  in  establishing 
under  the  Smithsonian  Institution  a  special  research  fund  for  my 
scientific  work. 

TECHNICAL  TERMS. 

In  describing  the  skulls  of  bears  a  few  terms  are  used  in  a  special 
sense  which  it  is  desirable  to  understand. 

The  frontal  shield  is  that  part  of  the  top  of  the  skull  extending 
from  the  base  of  the  rostrum  backward  to  the  meeting  point  of  the 
temporal  impressions.  It  is  elevated  above  the  surrounding  parts 
and  is  sharply  defined.  Its  posterior  point,  confined  between  the  tem- 
poral impressions,  is  longer  in  female  than  in  male  skulls,  and  up 
to  a  certain  limit  becomes  shorter  with  age. 

The  postorbital  processes  stand  out  from  the  sides  of  the  frontal 
shield,  limiting  the  orbits  posteriorly. 

The  term  sulcate  is  applied  to  skulls  having  a  longitudinal  median 
depression  or  groove  in  the  frontal  shield,  usually  shallow  and  rather 
broad  and  without  definite  lateral  limits. 

1  Others  who  have  helped  hy  the  presentation  or  loan  of  material  are :  C.  E.  Aiken. 
Dr.  J.  A.  Allen,  Dr.  R.  M.  Anderson,  Edward  F.  Ball,  Dr.  Arthur  H.  Bannon,  Br.  William 
Bebb,  H.  C.  Beggs,  Mrs.  C.  C.  Beggs,  John  P.  Bird,  W.  C.  Bradbury,  J.  Stanley-Brown, 
Fred  K.  Burnham,  Mr.  and  Mrs.  E.  S.  Cameron,  Dr.  Frank  M.  Chapman,  R.  H.  Chapman, 
Elton  Clark,  James  L.  Clark,  Charles  B.  Cory,  Prof.  Charles  R.  Cross,  Heyward  Cutting, 
Frank  S.  Daggett,  E.  W.  Deming.  Howard  Eaton,  Charles  Farwell  Edson,  Lincoln  Ells- 
worth, Lieut.  G.  T.  Emmons,  J.  D.  Figgins,  J.  Stanley  Foster,  Charles  A.  Gianini,  Dr. 
J.  B.  Girard,  Dr.  Joseph  Grinnell,  Samuel  Henshaw,  Charles  J.  Hittell,  Dr.  W.  J.  Holland, 
Dr.  R.  Houston,  James  T.  Jardine,  Remington  Kellogg,  Francis  Kermode,  Charles  S.  King. 
Paul  Kleineidam,  Prof.  S.  H.  Knight,  Frederick  Lambart,  Edward  H.  Litchfield,  Col.  J.  A. 
McGuire,  John  Murgatroyd,  Prof.  C.  C.  Nutting,  Wilfred  H.  Osgood,  John  M.  Phillips, 
the  late  W.  Hallett  Phillips,  the  late  Warburton  Pike,  Wilson  Potter,  George  D.  Pratt, 
Dr.  E.  P.  Richardson,  Powhatan  Robinson,  Archibald  Rogers,  Carl  Rungius,  Homer  E. 
Sargent,  Prof.  W.  B.  Scott,  George  Shiras  3d,  George  Shiras  4th,  Dr.  H.  A.  Sifton,  Henry 
A.  Stewart,  Dr.  Walter  T.  Swingle,  P.  A.  Taverner,  Prof.  S.  D.  Thacher,  Dr.  Charles  H. 
Townsend,  Frederick  K.  Vreeland,  E.  R.  Warren,  A.  Bryan  Williams,  and  W.  W.  Wood. 

Fullest  acknowledgments  are  due  the  following  institutions  for  their  courtesy  in  loaning 
specimens :  American  Museum  of  Natural  History,  New  York ;  Carnegie  Museum,  Pitts- 
burgh ;  Colorado  Museum  of  Natural  History,  Denver ;  Field  Museum  of  Natural  History, 
Chicago ;  Museum  of  Comparative  Zoology,  Cambridge ;  Museum  of  History,  Science,  and 
Art,  Los  Angeles ;  Museum  of  Vertebrate  Zoology,  University  of  California ;  Peabody 
Museum  of  Salem ;  Peabody  Museum  of  Yale  University ;  Provincial  Museum,  Victoria, 
British  Columbia ;  U.  S.  National  Museum,  Washington ;  Victoria  Memorial  Museum, 
Ottawa ;  Zoological  Society  of  Philadelphia ;  and  the  Museums  of  the  Universities  of  Iowa, 
Kansas,  Nebraska,  and  Wyoming. 

For  the  frontispiece,  showing  a  wild  grizzly  in  a  pine  forest  near  Yellowstone  Park, 
western  Wyoming,  thanks  are  due  Frederick  K.  Vreeland,  who  was  so  fortunate  as  to 
take  the  photograph. 


12  NOBTH   AMERICAN   FAUNA.  [No.  41. 

The  term  dished  means  that  the  nasal  or  fronto-nasal  region  is 
depressed,  producing  a  change  of  angle  from  the  plane  of  the  anterior 
part  of  the  nasals  to  the  plane  (or  slope)  of  the  frontal  shield. 
Some  skulls  are  strongly  dished,  some  are  flat,  while  some  have  the 
fronto-nasal  region  elevated  and  compressed,  giving  a  very  different 
outline  from  that  of  the  normally  dished  skull. 

The  term  braincase  is  loosely  applied  to  the  whole  upper  rounded 
part  of  the  skull  between  the  occiput  and  postorbital  processes,  but 
not  including  the  wedge-shape  posterior  part  of  the  frontal  shield. 
Used  in  this  way,  it  covers  the  parietal  bones  and  posterior  part  of 
the  frontals  on  each  side  of  the  temporal  ridges,  including  not  only 
the  actual  bony  case  inclosing  the  brain,  but  also  its  anterior  con- 
tinuation (the  sinus  case). 

The  term  sinus  case  is  applied  more  definitely  to  the  smoothly 
rounded  part  of  the  frontals  below  the  shield  and  in  direct  continu- 
ation of  the  braincase — the  outer  shell  covering  the  large  sinuses  or 
air  cells  lying  between  the  nasal  chamber  and  the  brain.  It  is  not 
always  discriminated  from  the  braincase. 

The  term  bellied  is  applied  to  the  posterior  part  of  the  inferior 
border  of  the  ramus  of  the  under  jaw  to  indicate  a  swelling  or  thick- 
ening common  in  many  species. 

The  term  subangvZar  border  is  applied  to  the  posterior  part  of  the 
inferior  border  of  the  under  jaw,  immediately  anterior  to  the  angular 
process,  and  usually  set  off  from  the  rest  of  the  ramus  by  a  step  or 
small  tubercle. 

The  term  keeled  is  applied  to  a  not  uncommon  condition  of  the 
upper  part  of  the  sinus  case,  in  which  the  top  or  arch  is  compressed, 
rising  rather  narrowly  into  the  anterior  part  of  the  sagittal  crest 
and  posterior  part  of  the  frontal  shield.  The  condition  is  marked 
in  Ursus  eulophus  and  occurs  in  several  other  species.  In  most 
species,  however,  this  part  of  the  braincase  is  rather  broadly  rounded, 
the  sagittal  crest  rising  abruptly  from  the  median  line. 

Measurements  are  always  in  millimeters  unless  otherwise  specified. 

The  museum  number  of  the  skull,  unless  otherwise  stated,  is  under- 
stood to  be  that  of  the  United  States  National  Museum. 

CLASSIFICATION   OF  GRIZZLY  AND   BIG   BROWN   BEARS. 

The  differences  formerly  supposed  to  exist  between  the  grizzlies 
and  the  big  brown  bears  appear,  in  the  light  of  the  material  now 
available,  to  distinguish  certain  groups  of  species  from  certain  other 
groups,  rather  than  the  grizzlies  collectively  from  the  big  brown 
bears  collectively.  In  other  words,  the  differences  between  the  griz- 
zlies on  the  one  hand  and  the  big  brown  bears  on  the  other  are 


1018.]  INTRODUCTION.  13 

neither  so  great  nor  so  constant  as  at  one  time  believed.  And  there 
are  species  which  in  the  present  state  of  knowledge  can  not  be  posi- 
tively referred  to  either  group.  In  fact,  it  seems  at  least  possible 
that  certain  species  which  appear  to  belong  with  the  grizzlies  are 
closely  related  to  certain  other  species  which  clearly  belong  with 
the  big  brown  bears.  The  typical  brown  bears  differ  from  the 
typical  grizzlies  in  peculiarities  of  color,  claws,  skull,  and  teeth. 
The  color  of  the  former  is  more  uniform,  with  less  of  the  surface 
grizzling  due  to  admixture  of  pale-tipped  hairs;  the  claws  are 
shorter,  more  curved,  darker,  and  scurfy  instead  of  smooth;  the 
skull  is  more  massive;  the  fourth  lower  premolar  is  conical,  lacking 
the  sulcate  heel  of  the  true  grizzlies.  But  these  are  average  differ- 
ences, not  one  of  which  holds  true  throughout  the  group.  Most  of 
the  specimens  in  museums  consist  of  skulls  only,  unaccompanied  by 
skins  or  claws,  leaving  a  doubt  as  to  the  external  characters;  and 
in  old  bears  the  important  fourth  lower  premolar  is  likely  to  be  so 
worn  that  its  original  form  can  not  be  made  out.  And,  worst  of  all, 
some  of  the  grizzlies  lack  the  distinctive  type  of  premolar,  leaving 
only  the  skull  as  a  guide  to  their  affinities.  The  present  classification, 
therefore,  must  be  regarded  as  tentative  and  subject  to  revision. 

RELATIVE  VALUES  OF  CRANIAL  AND  DENTAL  CHARACTERS. 

In  my  judgment  cranial  characters  among  the  bears  of  the  genus 
Ursus  are  more  permanent  and  of  more  significance  from  the  stand- 
point of  classification  than  minor  tooth  characters.  The  teeth  are 
strongly  modified  by  food  and  consequently  in  some  cases  present 
marked  variations  in  the  same  group.  Thus  the  skull  of  adult  male 
chelidonias  from  the  coast  of  southern  British  Columbia  is  almost  in- 
distinguishable from  that  of  imperator  from  the  Yellowstone  Park,  a 
member  of  the  horribilis  group ;  but  imperator  has  very  large  molars, 
nearly  as  big  as  those  of  horribilis  and  bairdi,  while  chelidonias,  be- 
ing a  fish  eater,  has  such  small  molars  that  were  it  not  for  the  skull 
no  one  would  think  of  placing  it  in  the  horribilis  group. 

Cranial  and  dental  characters  among  the  big  bears  are  very  subtle. 
As  a  rule  comparison  of  any  two  skulls  of  essentially  the  same  size 
brings  to  light  so  many  resemblances  that  one  is  likely  to  infer  a  far 
closer  relationship  than  actually  exists.  This  is  because  the  big  bears 
of  the  genus  Ursus  are  such  a  closely  interrelated  group  that  the  re- 
semblances far  outnumber  the  differences.  Hence  the  greatest  caution 
is  necessary  to  avoid  misleading  conclusions. 

The  present  paper  is  merely  a  review  of  the  existing  state  of  knowl- 
edge of  the  grizzlies  and  big  brown  bears  of  America  and  does  not 
include  either  the  polar  or  the  black  bears.    It  is  not  intended  as  a 


14  NORTH   AMERICAN    FAUNA.  [No.  41. 

monographic  revision,  but  aims  to  supply  a  list  of  the  species,  to- 
gether with  descriptions  and  comparisons  of  adult  skulls,  chiefly 
males.  Little  is  said  of  external  characters,  for  the  reason  that  little 
is  known,  only  a  few  skins  with  claws  being  available  for  study. 

List  of  Species  and  Subspecies  of  Grizzly  and  Big  Brown  Bears,  with  Type 

Localities.1 

(Classification  provisional.) 
Horribilis  group  (pp.  17-34): 

Ursus  horribilis  horribilis  Ord Missouri  River,  northeastern  Montana. 

horribilis  bairdi  Merriam Blue  River,  Summit  County,  Colorado. 

horribilis  impera tor  Merriam Yellowstone  National  Park,  Wyoming. 

chelidonias  nobis Jervis  Inlet,  British  Columbia. 

atnarJco  nobis / .  .Atnarko  River,  British  Columbia. 

kwdkiutl  Merriam Jervis  Inlet,  British  Columbia. 

nortoni  Merriam Southeastern  side  Yakutat  Bay,  Alaska. 

warburtoni  Merriam Atnarko  River,  British  Columbia. 

neglectus  Merriam Near    Hawk     Inlet,     Admiralty     Island, 

Southeastern  Alaska. 

californicus  Merriam Monterey,  California. 

tularensis  Merriam Fort  Tejon,  California. 

colusus  Merriam Sacramento  Valley,  California. 

dusorgus  nobis  2 .Jack  Pine  River,     lberta-British  Columbia 

boundary. 
Planiceps  group  (pp.  34-53): 

Ursus  nelsoni  Merriam Colonia  Garcia,  Chihuahua,  Mexico. 

texensis  texensis  Merriam Davis  Mountains,  Texas. 

texensis  navaho  Merriam Navajo  country  near  Fort  Defiance,   Ari- 
zona.    (Probably  Chuska  Mountains.) 

planiceps  nobis Colorado  (exact  locality  uncertain). 

macrodon  nobis Twin  Lakes,  Colorado. 

mirus  nobis Yellowstone  National  Park ,  Wyoming. 

eltonclarhi  Merriam Near  Freshwater  Bay,   Chichagof  Island, 

Alaska. 

tahltanicus  Merriam Klappan  Creek  (=Third  South  Fork  Stikine 

River),  British  Columbia. 

insularis  Merriam Admiralty  Island,  Alaska. 

orgilos  Merriam Bartlett  Bay,  east  side  Glacier  Bay,  South- 
eastern Alaska. 

orgiloides  nobis Italio  River,  Alaska. 

pallasi  Merriam Donjek  River,  southwestern  Yukon. 

rungiusi  rungiusi  nobis Rocky  Mountains,  headwaters  Athabaska 

River,  Alberta. 

rungiusi  sagittalis  iiobis Champagne  Landing,  southwestern  Yukon. 

macfarlani  nobis Anderson    River,    50    miles    below    Fort 

Anderson,  Mackenzie. 

canadensis  Merriam  2 Moose  Pass,  near  Mount  Robson,  British 

Columbia. 

1  Nearly  130  years  ago  Prof.  Zauschner  proposed  the  name  Ursus  saribur  for  an  animal 
'•  from  the  region  of  Canada  "  (Bestlmmung  der  Hundsart  Krokute,  und  der  Barenart 
Saribur,  p.  8,  1788),  but  the  species  appears  to  be  impossible  of  identification. 

*  Reference  to  group  provisional. 


1918.]  LIST  OF   SPECIES  AND   SUBSPECIES.  15 

Arizonse  group  (pp.  53-76): 

Ursus  arizonse  Merriam Escudilla    Mountains,     Apache     County, 

Arizona. 

idahoensis  nobis North  Fork  Teton  River,  eastern  Idaho. 

pulchellus  pulchellus  nobis Ross  River,  Yukon. 

pulchellus  ereunetts  nobis Beaverfoot     Range,     Kootenay     District, 

British  Columbia. 

oribasus  nobis Upper  Liard  River,  Yukon. 

chelan  Merriam East    slope    Cascade    Mountains,    Chelan 

County,  Washington. 

shoshone  Merriam Estes  Park,  Colorado. 

kennerlyi  Merriam Mountains  of  northeastern  Sonora,  near  Los 

Nogales,  Mexico. 

utahensis  Merriam Salina  Creek,  near  Mayfield,  Utah. 

perturbans  nobis Mount  Taylor,  northern  New  Mexico. 

rogersi  rogersi  nobis Upper  Greybull   River,   Absaroka  Moun- 
tains, Wyoming. 

rogersi  bisonophagus  nobis Black   Hills    (Bear   Lodge),    northeastern 

Wyoming. 

pervagor  Merriam Pemberton  Lake  (now  Lillooet  Lake),  Brit- 
ish Columbia. 

caurmus  Merriam Berners  Bay,  east  side  Lynn  Canal,  South- 
eastern Alaska. 

eulophus  Merriam Admiralty  Island,  Southeastern  Alaska. 

klamathensis  Merriam  * Beswick,  near  mouth  Shovel  Creek,  Kla- 
math River,  northern  California. 

mendocinensis  Merriam  ' Long  Valley,  Mendocino  County,  Califor- 
nia. 

magister  Merriam  l Los  Biacitos,  Santa  Ana  Mountains,  South- 
ern California. 
Hylodromus  group  (pp.  77-84): 

Ursus  hylodromus  Elliot Rocky  Mountains,  western  Alberta. 

kluane  kluane  Merriam McConnell  River,  Yukon. 

kluane  impiger  nobis Columbia  Valley,  British  Columbia. 

pellyensis  nobis Ketza  Divide,  Pelly  Mountains,  Yukon. 

andersoni  nobis ■ Dease  River,  near  Great  Bear  Lake,  Mac- 
kenzie. 
Eorrixus  group  (pp.  84-88): 

Ursus  apache  Merriam Whorton  Creek,  south  slope  White  Moun- 
tains, eastern  Arizona  (a  few  miles  west 
of  Blue). 

horriseus  Baird Coppermines,  southwestern  New  Mexico. 

henshawi  Merriam Southern   Sierra   Nevada,    near    Havilah, 

Kern  County,  California. 
Stikeenensis  group  (pp.  88-94): 

Ursus  stikeenensis  Merriam Tatletuey  Lake,  tributary  to  Finlay  River, 

near  head  Skeena  River,  British  Colum- 
bia. 

crassodon  nobis Klappan  Creek  (=Third  South  Fork  Stilrine 

River),  British  Columbia. 

crassus  nobis  ' Upper  Macmillan  River,  Yukon. 

mirabilis  Merriam  1 Admiralty  Island,  Alaska. 

absarokus  Merriam  * Little   Bighorn   River,   northern   Bighorn 

Mountains,  Montana. 

'Reference  to  group  provisional. 
64854°— 18 2 


16  •  NORTH   AMERICAN   FAUNA.  [No.  41. 

Alascensis  group  (pp.  94-99): 

Ursus  alascensis  Merriam Unalaklik  River,  Alaska. 

t  klat  Merriam Head  of  Toklat  River,  north  base  Alaska 

Range,  near  Mount  McKinley,  Alaska. 

latifrons  Merriam Jasper  House,  Alberta. 

Richardsoni  group  (pp.  99-106): 

Ursus  richardsoni  Swainson Shore  of  Arctic  Ocean,  west  side  Bathurst 

Inlet,  near  mouth  of  Hood  River. 

russelli  Merriam  1 West  side  Mackenzie  River  delta,  Canada. 

phxonyx  Merriam Glacier     Mountain,     Tanana     Mountains, 

Alaska  (about  2  miles  below  source  of 
Comet  Creek,  near  Fortymile  Creek,  be- 
tween Yukon  and  Tanana  Rivers). 

internationalis  Merriam Alaska- Yukon  boundary,  about  50  miles 

south  of  Arctic  coast. 

ophrus  Merriam Eastern  British  Columbia  (exact  locality 

unknown). 

•washake  Merriam North    Fork    Shoshone    River,    Absaroka 

Mountains,  western  Wyoming. 
Kidderi  group  (pp.  106-110): 

Ursus  kidderi  kidderi  Merriam Chinitna  Bay,  Cook  Inlet,  Alaska. 

kidderi  tundrensis  Merriam Shaktolik  River,  Norton  Sound,  Alaska. 

eximius  Merriam Head  of  Knik  Arm,  Cook  Inlet,  Alaska. 

Innuitus  group  (pp.  110-115): 

Ursus  innuitus  Merriam Golofnin  Bay,  south  side  Seward  Penin- 
sula, northwestern  Alaska. 

cressonus  Merriam Lakina  River,  south  slope  Wrangell  Range, 

Alaska. 

alexandrse  Merriam  * Kusilof  Lake,  Kenai  Peninsula,  Alaska. 

Townsendi  group  (pp.  115-116): 

Ursus  townsendi  Merriam Mainland   of   Southeastern  Alaska  (exact 

locality  uncertain). 
Dalli  group  (pp.  116-124): 

Ursus  dalli  Merriam Yakutat  Bay  (northwest  side),  Alaska. 

hoots  Merriam Clearwater  Creek,  a  north  branch  of  Stikine 

River,  British  Columbia. 

sitkensis  Merriam Sitka  Islands,  Alaska. 

shirasi  Merriam Pybus  Bay,  Admiralty  Island,  Alaska. 

nuchek  Merriam  1 Head     of    Nuchek    Bay,     Hinchinbrook 

Island.  Prince  William  Sound,  Alaska. 
Gyas  group  (pp.  124-127): 

Ursus  gyas  Merriam Pavlof  Bay,  Alaska  Peninsula. 

middendorffi  Merriam Kodiak  Island,  Alaska. 

Kenaiensis  group  (pp.  127-131): 

Ursus  kenaiensis  Merriam Cape  Elizabeth,  extreme  west  end  Kenai 

Peninsula,  Alaska. 

sheldoni  Merriam Montague  Island,  Prince  William  Sound, 

Alaska. 


Vetularctos  genus  nobis  fpp.  131-133): 

Vetularctos  inopinatus  nobis Rendezvous  Lake,  northeast  of  Fort  Ander- 
son, Mackenzie. 
1  Reference  to  group  provisional. 


1918.J  HORRIBILIS  GROUP.  17 

DESCRIPTIONS  OF  SPECIES  AND  SUBSPECIES. 
Horribilis  Group. 
URSUS  HORRIBILIS  HORRIBILIS  Obd. 

Big  Plains  Gbizzly. 
(Pi.  XIV.) 

Ursus  horribilis  Ord,  Guthrie's  Geography,  2d  Amer.  Ed.,  pp.  291,  300,  1815 
(Rhoads'  reprint,  1894).  Based  on  the  white  bear  of  Lewis  and  Clark, 
particularly  the  one  from  eastern  Montana  killed  May  5,  1805,  on  the 
Missouri  River,  near  the  mouth  of  Poplar  River  (called  by  them  Porcupine 
River,  but  not  the  same  as  the  Porcupine  of  to-day,  which  is  about  50  miles 
farther  west). 

Ursus  ferox  Rafinesque,  Amer.  Monthly  Mag.,  I,  p.  437,  Oct.,  1817  (nomen 
nudum). 

"  Ursus  ferox  Lewis  &  Clarck,"  Desmarest,  Mammalogie,  p.  164,  Paris,  1820 
(under  U.  cinereus). 

Ursus  cinereus  Desmarest,  Mammalogie,  p.  164,  Paris,  1820. 

Ursus  griseus  Choris,  Voyage  Pittoresque  autour  du  Monde,  folio  colored  plate 
and  text  (unpaged),  Paris,  1822. 

Ursus  candescens  Ham.  Smith,  in  Griffith's  Ouvier,  II,  p.  229  and  facing  plate, 
1827 ;  ibid.  V,  p.  112,  1827. 

Type  locality. — Missouri  Kiver,  a  little  above  mouth  of  Poplar 
River,  northeastern  Montana. 

Characters. — Size  huge;  skull  long  and  massive;  claws  long,  mod- 
erately or  slightly  curved,  and  smoothly  polished;  usually  streaked 
lengthwise  with  whitish  or  yellowish,  which  increases  with  age  until 
in  some  old  individuals  the  claws  are  almost  wholly  white  or  whitish. 
Color  variable,  usually  light. 

Cranial  characters. — Old  male  (topotype,  No.  202739)  from  Breaks 
of  Missouri  River,  about  100  miles  north  of  Fort  Miles,  eastern  Mon- 
tana, April  4,  1890;  killed  and  presented  by  E.  S.  Cameron  and  wife: 
Skull  huge  (total  length,  400  mm.=15|  inches)  ;  vault  of  cranium 
moderately  arched;  zygomata  rather  squarely  spreading  posteriorly, 
moderately  outbowed,  the  squamosal  part  rather  broadly  expanded 
vertically ;  frontal  shield  rather  broad,  flattish,  sloping  gradually  up- 
ward to  apex,  the  posterior  part  broader  than  usual  (not  cut  away  on 
sides  by  incurving  temporal  ridges)  ;  sagittal  crest  long,  straight  on 
top,  high  posteriorly,  reaching  forward  over  posterior  fourth  of  fron- 
tals;  postorbital  processes  large,  flat,  and  horizontally  outstanding; 
rostrum  high,  ascending  posteriorly ;  f ronto-nasal  region  rather  high 
but  distinctly  dished  and  somewhat  sulcate;  palate  long,  concave  in 
cross  section ;  postpalatal  shelf  broad  and  flat,  well  rounded  on  sides ; 
underjaw  large  and  rather  massive;  coronoid  blade  high,  its  apex  not 
strongly  produced  posteriorly;  ramus  rather  flat;   diastema  long. 


18  NORTH   AMERICAN   FAUNA.  [No.  41. 

Teeth  large,  especially  lower  canine  and  last  upper  and  middle  lower 
molar.  Skull  of  female  (No.  13245,  from  head  of  Big  Porcupine 
Creek  [not  Porcupine  River]  between  Musselshell  and  Yellow- 
stone Rivers,  eastern  Montana)  :  Of  generalized  grizzly  type; 
large  and  rather  massive;  vault  of  cranium  rather  flat;  braincase 
rather  broad;  frontal  shield  of  moderate  breadth,  slightly  convex, 
slightly  depressed  or  sulcate  between  orbits,  elongate-lyrate  pos- 
teriorly; postorbital  processes  weak  and  slightly  decurved;  sagittal 
crest  not  yet  reaching  fronto-parietal  suture;  muzzle  moderate  or 
rather  short;  zygomata  moderately  spreading  and  rather  angular; 
palate  rather  narrow.     Teeth  large ;  M-2-  large  and  subrectangular. 

Cranial  comparisons. — Adult  male  (No.  202739,  topotype)  com- 
pared with  two  adult  males  of  bairdi,  one  from  near  Cheyenne,  south- 
eastern Wyoming  (Yountz),  the  other  from  Blue  River,  Summit 
County,  Colorado  (No.  203805):  Size  slightly  larger;  fronto-nasal 
region  (interorbitally)  dished  instead  of  elevated,  not  compressed  in 
front  of  orbits;  frontal  shield  slightly  broader,  rising  from  plane 
of  nasals  (in  bairdi  continuing  plane  of  nasals)  ;  palate  longer  and 
broader;  postpalatal  shelf  much  broader  and  more  rounded  on  sides; 
mastoids  short,  appressed,  vertical  (in  bairdi  longer  and  divergent) ; 
meatus  tube  compressed  between  mastoid  and  glenoid  (in  bairdi  not 
compressed);  underjaw  longer;  ramus  longer  and  flatter  (much 
less  swollen  on  outer  side)  ;  coronoid  blade  less  falcate  (notch  shal- 
lower) ;  lower  canine  and  last  upper  molar  larger. 

Male  adult  compared  with  male  adult  absarokus  (type)  :  Size  de- 
cidedly larger;  vault  of  cranium  less  arched;  frontal  shield  rising 
more  gradually;  rostrum  longer;  palate  broader  and  much  longer; 
ramus  much  longer;  meatus  tube  compressed  between  mastoid  and 
glenoid  (in  absarokus  not  compressed).  Teeth  much  larger,  espe- 
cially lower  canine,  last  upper  molar,  and  middle  lower  molar. 

Remarks. — Until  recently  the  absence  of  authentic  specimens  from 
the  neighborhood  of  the  type  locality,  in  connection  with  the  pres- 
ence of  several  species  of  grizzly  in  Montana,  caused  an  embarrassing 
uncertainty  as  to  which  species  was  entitled  to  the  name  horribilis. 
But  the  slow  accumulation  of  material  during  the  long  period  in 
which  I  have  been  engaged  in  a  study  of  the  group  made  it  possible 
to  map  the  ranges  of  some  of  the  species  with  some  degree  of  con- 
fidence; and  finally,  through  the  generosity  of  Mr.  and  Mrs.  E.  S. 
Cameron,  of  Marsh,  Montana,  I  have  been  presented  with  a  splendid 
skull  of  an  old  male  horribilis  from  the  Breaks  of  the  Missouri, 
about  100  miles  north  of  Fort  Miles,  Montana  (practically  the  type 
locality).  This  skull  proves  that  the  huge  buffalo-killing  grizzly 
of  the  Great  Plains  bordering  the  Missouri  in  eastern  Montana  and 
the  Dakotas — the  "White  Bear"  of  Lewis  and  Clark — is  really  the 
species  to  which  Ord  in  1815  gave  the  name  Ursus  horribilis. 


1918.]  HOREIBILIS  GKOUP.  19 

Skull  measurements. — Old  male  (No.  202739,  from  Missouri  Breaks, 
eastern  Montana):  Basal  length,  351;  occipito-nasal  length,  350; 
palatal  length,  192 ;  zygomatic  breadth,  247 ;  interorbital  breadth,  86. 

URSUS  HORRIBILIS   BAIRDI   Mebriam.1 

Baied  Grizzly. 

Ursus  bairdi  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  192-193,  August 
13,  1914. 

Type  locality. — Blue  River,  Summit  County,  Colorado. 

Type  specimen. — No.  203805,  S  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection  (=3147,  Warren  collection). 

Range. — Southern  Rocky  Mountain  region  from  San  Juan  Moun- 
tains, southwestern  Colorado,  northward  through  Wyoming  to  Mon- 
tana, and  perhaps  to  southeastern  British  Columbia.  Probably  a 
mountain  animal,  while  its  neighbor  horribilis  was  a  plains  species. 

Characters. — Size  large — in  the  Rocky  Mountain  region  exceeded 
only,  if  at  all,  by  horribilis;  skull  long,  with  narrow  elevated  fronto- 
nasal region ;  claws  of  moderate  length,  smooth. 

Cranial  characters. — Old  male  (type)  from  Blue  River,  Colorado, 
and  a  closely  similar  old  male  from  Sabille  Hole,  Laramie  County, 
southeastern  Wyoming,  a  little  north  of  Cheyenne :  Size  large ;  fronto- 
nasal region  high  and  rather  narrow;  rostrum  rather  long,  narrow, 
and  strongly  compressed  in  front  of  orbits;  face  long  sloping;  frontal 
shield  flat,  short  pointed,  faintly  depressed  medially ;  shield  and  nasals 
in  essentially  same  plane  except  that  anterior  third  of  nasals  is 
slightly  upturned;  postorbital  processes  large,  outstanding,  flat,  and 
blunt;  orbital  rims  prominent;  sagittal  crest  moderately  high  pos- 
teriorly; temporal  impressions  short,  incurved,  beaded;  zygomata 
strongly  outbowed,  squarely  spreading  posteriorly;  lachrymal  duct 
notching  orbital  rim ;  squamosal  shelf  short,  arched  over  meatus,  the 
free  edge  thickened ;  palate  and  postpalatal  shelf  of  moderate  breadth ; 
mastoids  rather  long,  divergent.    Molars  large,  especially  M-2-. 

Immature  males,  up  to  at  least  the  fifth  or  sixth  year,  have  rather 
narrow  convex  frontal  shields  with  weak  decurved  postorbital  proc- 
esses, and  may  be  recognized  at  once  by  the  form  of  the  f ronto-nasal 
region,  which  is  high,  narrow,  and  strongly  pinched  in  immediately 
in  front  of  the  orbits. 

Cranial  comparisons. — Old  male  compared  with  old  male  horribilis 
(No.  202739,  from  Missouri  Breaks,  eastern  Montana)  :  Size  essentially 
the  same;  frontal  shield  slightly  lower  posteriorly,  higher  anteriorly, 
the  point  lyrate  and  more  slender ;  f  ronto-nasal  region  elevated  (never 
dished  or  sulcate  between  orbits)  ;  base  of  rostrum  much  more  com- 

1  Named  for  S.  F.  Baird,  former  Secretary  of  the  Smithsonian  Institution  and  founder 
of  the  U.  S.  National  Museum. 


20  NORTH  AMERICAN  FAUNA.  [No.  41. 

pressed  laterally  in  front  of  orbits ;  palate  narrower ;  mastoids  longer 
and  more  divergent;  meatus  tube  broadly  rounded  and  free  (not 
compressed  between  mastoid  and  glenoid) ;  under  jaw  shorter;  ramus 
more  swollen  on  outer  side ;  inferior  border  of  ramus  shorter  and  less 
upturned;  coronoid  blade  more  recurved,  the  apex  narrower  and 
reaching  farther  backward.  Teeth  similar  but  slightly  smaller; 
M1-  and  M>  smaller  and  less  massive. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  348;  occipito- 
nasal  length,  325;  palatal  length,  157;  zygomatic  breadth,  235;  inter- 
orbital  breadth,  81. 

URSUS  HORRIBILIS  IMPERATOR  Mereiam. 

/ 
Yellowstone  Pabk  Big  Grizzly. 

Ursus  imperator  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  180-181, 
August  13,  1914. 

Type  locality. — Lake  Hotel,  Yellowstone  National  Park,  Wyoming. 

Type  specimen. — No.  176297,  $  old,  U.  S.  National  Museum.  Died 
in  National  Zoological  Park. 

Range. — Yellowstone  National  Park,  Wyoming;  limits  unknown. 

Characters. — Size  large;  skull  massive,  with  large  horizontally  out- 
standing postorbitals ;  closely  similar  to  horribilis. 

Cranial  characters. — Old  male  (type)  and  a  much  finer  male  (No. 
216205)  :  Size  large  (only  slightly  smaller  than  horribilis)  ;  skull 
massive;  vault  of  cranium  rather  high;  fronto-nasal  region  moder- 
ately dished;  sagittal  crest  long,  high,  and  straight;  frontal 
shield  exceedingly  short,  flattish,  shallowly  sulcate  medially, 
slightly  swollen  over  orbits,  ending  in  short  obtuse  point  about  30 
millimeters  behind  plane  of  postorbitals;  postorbitals  long,  peglike, 
horizontally  outstanding;  lachrymal  duct  opening  on  orbital  rim; 
zygomata  moderately  outstanding  and  outbowed,  the  squamosal  base 
broadly  expanded;  squamosal  shelves  broad,  arched  over  meatus; 
palate  moderate;  postpalatal  shelf  rather  long;  occipito-sphenoid  91 
mm.;  mastoids  short;  underjaw  large,  long,  and  massive,  with  high 
vertical  coronoid  blade.     Teeth  large :  M.2-  with  long  heel. 

Cranial  comparisons. — Two  old  males  (Nos.  176297  and  216205) 
from  near  northern  boundary  of  Yellowstone  National  Park1  com- 
pared with  an  old  male  topotype  of  horriMUs  (No.  202739)  from 
Missouri  Breaks,  eastern  Montana :  Size  only  slightly  less,  general 
appearance  essentially  same;  frontal  shield  slightly  broader  inter- 
orbitally  and  decidedly  broader  across  postorbital  processes;  postor- 

1  While  both  are  old  males,  the  type  is  the  older  of  the  two  and  the  recessive  changes 
incidental  to  senility  have  already  begun.  It  differs  from  the  other  in  having  the  frontal 
shield  shorter  pointed  posteriorly  (with  consequent  lengthening  of  sagittal  crest),  the 
rostrum  narrower,  the  fronto-nasal  region  more  strongly  dished. 


1918.]  HORRIBILIS  GROUP.  21 

bital  processes  more  widely  outstanding  horizontally;  palate  and 
postpalatal  shelf  narrower;  mastoids  less  appressed  (apex  farther 
from  glenoid  surface,  leaving  broader  space  for  tube  of  auditory 
meatus).  Last  upper  molar  somewhat  smaller,  heel  equally  long  but 
narrower  posteriorly ;  middle  lower  molar  narrower. 

Old  males  (type  and  topotype)  compared  with  old  male  bairdi: 
Size  essentially  same;  frontal  shield  broader  throughout;  distance 
between  tips  of  postorbital  processes  much  greater;  fronto-nasal 
region  more  dished  (in  bairdi  elevated)  and  less  compressed;  inion 
more  strongly  developed. 

Old  male  (type)  compared  with  adult  male  absarokus  (type)  : 
Size  slightly  larger;  frontal  shield  broader  and  flatter;  postorbital 
processes  longer  and  more  widely  outstanding;  rostrum  higher  and 
more  nearly  horizontal  (in  absarokus  lower  anteriorly  and  more 
sloping) ;  sagittal  crest  longer;  squamosal  root  of  zygoma  very  much* 
more  expanded  vertically;  palate  much  longer. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  336 ;  occipito- 
nasal  length,  317;  palatal  length,  191;  zygomatic  breadth,  232;  inter- 
orbital  breadth,  90.  Another  and  more  perfect  old  male  from  Yel- 
lowstone Park  (No.  216205):  Basal  length,  340 j1  occipito-nasal 
length,  332 ;  palatal  length,  186 ;  zygomatic  breadth,  230 ;  interorbital 
breadth,  90. 

URSUS  CHELIDONIAS  sp.  NOV. 
Jeevis  Inlet  Grizzly. 

Type  No.  223133,  $  old,  U.  S.  National  Museum,  Biological  Sur- 
vey collection.  Collected  on  river  at  head  of  Jervis  Inlet,  British 
Columbia,  in  1916,  by  Forrest  and  Fred  Johnstone. 

Characters. — Size  very  large;  external  characters  unknown;  skull 
of  male  strikingly  like  that  of  imperator  but  teeth,  especially  upper 
molars,  very  much  smaller,  resembling  those  of  kwakiutl. 

Cranial  characters. — Adult  male  (type)  :  Skull  very  large,  massive, 
flat  on  top,  with  horizontally  outstanding  or  slightly  uplifted  post- 
orbital processes ;  shield  moderately  broad,  flat,  the  point  long,  reach- 
ing back  to  fronto-parietal  suture;  rostrum  elevated  in  same  plane 
with  shield;  sagittal  crest  short,  confined  to  parietals;  zygomata 
moderately  outstanding  and  outbowed ;  nares  small ;  postpalatal  shelf 
rather  broad;  occipito-sphenoid  unusually  long  (102)  ;  basicranial 
axis  flat ;  mastoids  long  and  spreading ;  meatus  tubes  large  and  free ; 
underjaw  large  and  massive ;  coronoid  blade  high  and  nearly  vertical ; 
ramus  rather  broad  posteriorly,  its  inferior  border  concave  under 
anterior  molar,   convex   behind   plane   of   last   molar;   subangular 

1  In  part  restored. 


22  NORTH  AMERICAN  FAUNA.  [No.  41. 

border  short ;  canines  moderate,  rather  small  for  size  of  skull ;  upper 
and  lower  molars  small;  heel  of  M>  short  and  rather  broadly 
rounded ;  last  lower  premolar  of  grizzly  type. 

Cranial  comparisons. — Ursus  chelidonias  requires  comparison  with 
only  two  species,  imperator  and  warburtoni.  Adult  male  (type) 
compared  with  equally  old  male  imperator  from  Hell  Roaring  Creek 
near  northern  boundary  of  Yellowstone  National  Park:  Size 
slightly  greater;  general  form  and  proportions  almost  identical; 
point  of  shield  somewhat  longer ;  rostrum  more  completely  in  frontal 
plane;  zygomata  more  outbowed  anteriorly;  basicranial  axis  flatter 
(in  imperator  somewhat  arched)  ;  mastoids  longer  and  more  diver- 
gent; meatus  tube  longer;  postpalatal  shelf  broader;  coronoid  blade 
higher;  canines  about  same  size;  molars  (both  upper  and  lower) 
smaller,  the  difference  most  marked  in  M1,  M1,  and  MT. 

Compared  with  two  adult  males  of  warburtoni,  one  (No.  223946) 
from  Iskut  River  near  its  junction  with  the  Stikine;  the  other  (No. 
210142)  from  Chilkat  River  valley,  Southeastern  Alaska:  Skull  as  a 
whole  slightly  larger ;  frontal  shield  more  nearly  horizontal,  somewhat 
broader,  and  completely  flat  (in  warburtoni  slightly  convex)  ;  zygo- 
mata more  outbowed  anteriorly;  postpalatal  shelf  broader;  basi- 
cranial axis  flatter;  occipito-sphenoid  longer  (102  mm.,  contrasted 
with  95);  palate  longer;  postpalatal  shelf  broader;  under  jaw  more 
massive;  coronoid  blade  higher  (posterior  part  of  jaw  more  uplifted, 
raising  condyle  and  coronoid) ;  upper  canines  conspicuously  more 
slender;  lower  canines  much  smaller;  upper  and  lower  molars 
smaller ;  middle  lower  molar  not  only  shorter  but  differing  markedly 
in  proportions,  the  posterior  moiety  small  and  narrow  (14.5)  in  com- 
parison with  the  anterior  (17)  ;  PM^more  distinctly  of  grizzly  type; 
MTwith  saddle  open  (cusplet  on  inner  side  nearly  obsolete). 

Skull  measurements. — Old  male  (type)  :  Basal  length,  346 ;  occipito- 
nasal  length,  337 ;  palatal  length,  191 ;  zygomatic  breadth,  240 ;  inter- 
orbital  breadth,  91. 

URSUS  ATNARKO  sp.  NOV. 

Atnabko  Gbizzly. 

Type  No.  211452,  6*  ad.,  U.  S.  National  Museum,  Biological  Sur- 
vey collection.  From  Lonesome  Lake,1  Atnarko  River,  one  of 
the  upper  forks  of  the  Bella  Coola,  British  Columbia.  Collected  in 
September,  1915,  by  E.  H.  Edwards. 

1  Lonesome  Lake  la  about  30  miles  from  the  Junction  of  the  Whitewater,  or  Talchawko, 
and  the  Atnarko,  which  two  rivers  unite  to  form  the  Bella  Coola.  The  Bella  Coola  Is 
about  45  miles  long.  Lonesome  Lake  is  nearly  on  the  fifty-second  parallel,  and  by  the 
river  about  75  miles  from  the  head  of  Burke  Channel. 


1918.]  HOREIBILIS  GROUP.  23 

Characters. — Size  large;  external  characters  unknown;  skull  of 
male  long  and  narrow,  similar  in  general  to  that  of  kwakiutl  but 
much  narrower;  skull  of  female  surprisingly  different  from  that  of 
kwakiutl,  being  long,  low,  and  very  narrow,  while  kwakiutl  is  ex- 
ceptionally high  and  broad. 

Cranial  characters. — Adult  male  (type)  :  Skull  long,  low,  and 
narrow,  highest  in  posterior  frontal  region;  braincase  anteriorly 
keeling  into  sagittal  crest;  shield  narrow  and  rather  sharp  pointed 
posteriorly ;  postorbitals  moderate,  horizontally  outstanding ;  rostrum 
long  and  slender,  rising  gradually  into  plane  of  shield ;  zygomata  not 
widely  outstanding ;  sagittal  crest  long ;  palate  and  postpalatal  shelf 
long  and  narrow;  lower  jaw  long;  coronoid  blade  moderate;  sub- 
angular  border  short.  Dentition  weak:  canines  small  for  size  of 
skull  (as  in  kwakiutl)  ;  molars  even  smaller  than  in  kwakiutl. 
Adult  female  (based  on  3  adult  and  old  females,  Nos.  223177,  223182, 
and  223183)  from  Atnarko  River:  Skull  long,  low,  slender,  nearly 
flat  on  top;  shield  long  and  narrow,  not  rising  above  plane  of  ros- 
trum; postorbitals  weak,  outstanding;  rostrum  long  and  slender, 
passing  insensibly  into  frontal  plane;  zygomata  appressed;  palate 
long  and  narrow ;  coronoid  blade  moderate ;  subangular  border  short. 
Dentition  weak. 

Cranial  comparisons. — Compared  with  kwakiutl  the  skull  as  a  whole 
is  longer  and  narrower;  frontal  shield  much  narrower  and  more 
evenly  sloping  (rising  less  abruptly  from  rostrum) ;  postorbitals 
much  less  widely  outstanding;  posterior  frontal  region  higher,  keel- 
ing into  sagittal  crest;  palate  and  postpalatal  shelf  notably  longer 
and  narrower. 

Adult  and  old  female  compared  with  old  female  kwakiutl  of 
slightly  greater  basal  length:  Breadth  very  much  less  (across  post- 
orbitals 105  or  less,  contrasted  with  130) ;  shield  low,  narrow,  and 
flat  instead  of  broad,  high,  and  rising  abruptly  from  rostrum ;  post- 
orbitals much  smaller  and  weaker;  zygomata  appressed  instead  of 
outstanding;  palate,  postpalatal  shelf,  and  palatal  notch  very  much 
narrower ;  sagittal  crest  much  weaker. 

Remarks.— The  males  of  kwakiutl  and  atnarko  are  so  much  alike 
that  their  differences  would  naturally  be  regarded  as  of  only  sub- 
specific  weight;  but  the  females  are  so  strikingly  different  that  it 
seems  necessary  to  give  them  independent  specific  rank.  Their  spe- 
cific distinctness  is  the  more  certain  by  reason  of  the  geographic 
contiguity  of  the  localities  where  they  were  killed,  the  type  localities 
being  on  the  same  watercourse  and  not  more  than  75  miles  apart. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  345 ; * 
occipito-nasal  length,  325;  palatal  length,  199;  zygomatic  breadth, 
214;  interorbital  breadth,  75. 

1  Partly  restored. 


24  NORTH  AMERICAN   FAUNA.  [No.  41. 

URSUS  KWAKIUTL  Merriam. 

Kwakitjtl  Grizzly. 

Ursus  kwaJciutl  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  143-144,  Sep- 
tember 6,  1916. 

Type  locality. — Jervis  Inlet,  coast  of  southern  British  Columbia. 

Type  specimen. — No.  211748,  $  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.    Collected  May  17,  1916,  by  Fred  Mansell. 

Range. — Coast  region  of  British  Columbia  from  southwestern  cor- 
ner (Burrard  Inlet,  Howe  Sound,  Jervis  Inlet)  northwesterly  to  or 
beyond  the  lower  Bella  Coola. 

Characters. — Size  large;  color  dark;  ears  densely  furred;  claws 
unknown ;  skull  long,  but  little  arched. 

Color. — Skin  of  head  of  adult  male  (type) :  Nose  brown ;  head 
and  face  from  front  of  eyes  posteriorly  very  dark  brown,  darkest 
on  ears,  slightly  grizzled  on  occiput  by  golden-tipped  hairs. 

Cranial  characters. — Adult  male  (type):  Size  large;  skull  long, 
rather  low  and  narrow,  with  long  high  rostrum,  gradually  ascend- 
ing frontal  shield,  rather  low  fronto-parietal  region,  and  strongly 
outstanding  postorbitals.  Frontal  shield  of  moderate  breadth,  shal- 
lowly  sulcate  medially,  swollen  over  orbits,  short  pointed  posteriorly ; 
rostrum  long,  high,  and  rather  narrow;  nasals  flattened,  nearly 
horizontal;  fronto-nasal  region  sloping  gently  in  plane  of  shield; 
braincase  long,  its  anterior  part  keeling  into  elongate  sagittal  crest; 
zygomata  moderately  spreading  (becoming,  of  course,  more  strongly 
outbowed  in  old  age)  ;  palate  long,  concave,  slightly  arched  antero- 
posteriorly ;  postpalatal  shelf  large  and  broad,  nearly  square ;  denti- 
tion rather  light  for  size  of  skull;  underjaw  long,  its  inferior  border 
slightly  sinuous  (slightly  bellied  under  last  molars) ;  coronoid  blade 
broad  at  base,  not  very  high,  the  apex  slightly  produced  posteriorly 
but  falling  far  short  of  plane  of  condyle. 

Old  female  (No.  215432)  :  The  skull  of  an  old  female  from  Kwatna, 
on  the  lower  Bella  Coola,  is  obviously  of  this  species.  It  is  large 
and  rugged  for  a  female,  with  strongly  dished  fronto-nasal  region 
and  broad  massive  frontal  shield  rising  abruptly  from  a  very  small 
rostrum;  shield  sulcate,  swollen  between  sulcus  and  orbits,  lyrate 
pointed,  ending  at  fronto-parietal  suture ;  postorbitals  large  and  out- 
standing; zygomata  outstanding;  sagittal  crest  strongly  developed 
for  a  female;  palate  and  postpalatal  shelf  very  broad;  teeth  small; 
PMT badly  worn  but  apparently  subcorneal,  as  in  a  male  (No.  215433) 
from  the  same  locality. 

Cranial  comparisons. — The  only  species  requiring  comparison  with 
kwakiutl  are  the  much  larger  nortoni  and  the  very  different  pervagor. 
Adult  male  (type)  compared  with  adult  male  nortoni  (No.  213705) 
from  southeast  side  Yakutat  Bay:  Similar  in  general  but  much 


1918.]  HORRIBILIS  GROUP.  25 

shorter;  rostrum  slightly  longer;  hraincase  materially  shorter; 
shield  much  less  elevated;  fronto-nasal  region  much  less  dished; 
postpalatal  length  much  less.  Compared  with  male  pervagor:  Skull 
longer  and  less  highly  arched;  braincase  and  rostrum  materially 
longer ;  frontal  shield  shorter  pointed  posteriorly ;  rostrum  more  ele- 
vated anteriorly;  nasals  more  nearly  horizontal,  shorter  posteriorly, 
longer  anteriorly;  postpalatal  notch  longer;  under  jaw  decidedly 
longer  and  more  massive,  but  inferior  border  of  ramus  from  symphy- 
sis to  tubercle  of  essentially  same  length;  subangular  border  much 
longer.  Dentition  heavier  (both  upper  and  lower  canines,  incisors, 
and  molars  larger). 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  340; 
occipito-nasal  length,  330;  palatal  length,  184;  zygomatic  breadth, 
212 ;  interorbital  breadth,  85. 

URSUS  NORTONI 1  Mebriam. 

Yakutat  Grizzly. 

(PI.  XIII.) 

Ursus  nortoni  Merrtam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  179-180,  April 
13,  1914. 

Type  locality. — Near  Yakutat  village,  southeastern  side  Yakutat 
Bay,  Alaska. 

Type  specimen. — No.  178763,  $  ad.,  U.  S.  National  Museum, 
Biological  Survey  collection.  (Mother  of  $  cub  of  second  year, 
No.  178764.)    Collected  May  15,  1910,  by  G.  Frederick  Norton. 

Range. — Limited  apparently  to  coastal  plain  on  southeastern  side 
of  Yakutat  Bay. 

Characters. — A  true  grizzly,  apparently  of  the  califomicus- 
kwakiutl  group;  of  large  size,  with  smoothly  polished  horny  claws. 
Fourth  lower  premolar  large  and  of  ultra-grizzly  type  (long  heel 
with  continuous  sulcus  and  complete  lateral  ridges  ending  in  up- 
turned posterior  cusplets)  ;  molars,  especially  M-2-,  small  for  size  of 
skull.  Coloration  normal  for  a  grizzly;  skull  large,  massive,  and 
very  broad. 

Color. — Of  pale  grizzly  type;  head  grizzled  yellowish  or  golden 
brown;  muzzle  pale  brown;  neck  and  shoulders  to  middle  of  back 
pale  buffy  from  the  long  whitish  buff-tipped  hairs,  giving  the  skin 
viewed  from  behind  a  decidedly  whitish  appearance;  hinder  back 
and  rump  dark,  well  washed  with  pale  brown  tips;  lower  part  of 
legs  and  feet  dark  brown;  back  of  fore  feet  browner  and  not  so 
dark ;  underchin  and  throat  pale  soiled  buffy  whitish. 

1  Named  for  G.  Frederick  Norton,  who  collected  and  presented  the  type  specimen. 


26  NOBTH  AMERICAN  PATJNA.  [No.  41. 

Cranial  characters. — Adult  male  (No.  213705,  from  southeast  side 
of  Yakutat  Bay,  northeast  of  Yakutat  village)  :  Skull  large, 
long,  massive,  and  strongly  dished;  braincase  exceptionally  long; 
facial  part  of  skull  relatively  short;  frontal  shield  rather  broad, 
moderately  convex  transversely  and  sulcate  medially  (swollen 
between  sulcus  and  postorbitals),  rising  strongly  and  abruptly  from 
rostrum;  postorbital  processes  well  developed,  slightly  arched;  ros- 
trum rather  short,  horizontal;  zygomata  moderately  spreading, 
outbowed;  sagittal  crest  highly  developed;  squamosal  shelves  broad 
and  long;  palate  and  postpalatal  shelf  rather  long  and  of  moderate 
breadth;  palate  troughed  between  molar  series;  mastoids  moderate; 
underjaw  massive;  coronoid  blade  high,  broad,  and  not  recurved; 
ramus  long  and  straight.  Teeth  small  for  size  of  skull;  M-2-  rather 
short,  the  heel  evenly  emarginate  and  broadly  rounded  posteriorly. 

Adult  female  (type)  :  Rather  massive,  moderately  dished,  broad, 
with  large  broadly  rounded  braincase,  bowed  (but  not  widely  out- 
bowed  and  not  angular),  moderately  spreading  zygomata;  short 
weak  sagittal  crest  confined  to  posterior  half  of  parietals;  rather 
broad  lambdoid  crest  defining  a  rather  broadly  open  groove  or  sulcus 
continuous  with  that  of  the  deeply  sulcate  and  broadly  expanded  hori- 
zontal shelf  of  the  squamosal ;  squamosal  base  of  zygoma  moderately 
expanded;  frontal  shield  broad  and  long,  its  lyrate  point  reaching 
back  to  middle  of  parietals,  broadly  sulcate  between  orbits;  fronto- 
nasal region  moderately  dished;  postorbital  processes  large  and 
broadly  outstanding,  infraorbital  process  of  jugal  and  infra jugal 
process  of  maxillary  well  developed  (probably  not  constant)  ;  ros- 
trum broad,  short,  and  rather  depressed;  lachrymal  opening  within 
orbit;  palate  and  postpalatal  shelf  broad,  the  palate  rather  deeply 
concave  from  incisive  foramina  to  plane  of  hind  molars ;  postpalatal 
notch  broad  and  rather  squarely  truncate;  occipito-sphenoid  length 
90  (about  equal  to  distance  from  posterior  rim  of  alveolus  of  outer 
incisor  to  front  of  last  upper  molar)  ;  basioccipital  considerably 
longer  than  basisphenoid ;  occiput  broad,  rather  low,  truncate ;  coro- 
noid blade  rather  high.  Basicranial  axis  nearly  straight  and  hori- 
zontal, parallel  to  palatal  axis.  Dentition  light  in  both  sexes ;  canines 
and  molars  rather  small  for  size  of  skull ;  PM^  of  ultra  grizzly  type, 
with  slender  cone,  well-developed  completely  sulcate  heel  ending  pos- 
teriorly in  pair  of  upturned  cusplets ;  PM>  large  and  broad ;  heel  of 
M^  rather  short,  not  strongly  narrowed  posteriorly,  third  cusp 
small;  incisors  rather  large. 

Cranial  comparisons. — Male  adult  (type)  compared  with  male 
adult  kwakiutl  (type) :  Similar  in  general  hut  very  much  longer  both 
basally  and  on  top;  rostrum  slightly  shorter;  braincase  materially 
longer;  frontal  shield  rising  higher  and  more  abruptly  from  ros- 


1918.] 


HOBRTBILIS  GROUP.  27 


tram;  fronto-nasal  region  more  conspicuously  dished;  postpalatal 
length  very  much  greater;  under  jaw  longer  and  more  massive; 
ramus  longer;  subangular  border  shorter;  coronoid  blade  much 
larger  and  higher ;  teeth  closely  similar,  but  upper  canines  and  upper 
molars  slightly  larger. 

Specimens  examined. — About  a  dozen  specimens  have  been  ex- 
amined, mainly  females  and  young,  all  from  the  coastal  plain  south- 
east of  Yakutat  Bay.  Three  (including  type)  were  collected  by  G. 
Frederick  Norton,  and  by  him  presented  to  the  Biological  Survey; 
others  were  collected  by  Miss  Annie  M.  Alexander,  and  are  in  the 
Museum  of  Vertebrate  Zoology,  University  of  California ;  still  others 
(including  a  splendid  adult  male)  were  obtained  from  Yakutat  In- 
dians by  E.  M.  Axelson  and  purchased  by  me.  The  localities  are: 
Peninsula  between  Yakutat  and  Disenchantment  Bays  (northeast  of 
Yakutat  village)  ;  near  Yakutat  village;  front  of  Yakutat  Glacier; 
Ankow  River ;  Setuk  River ;  and  Anklin  River. 

Skull  measurements. — Adult  male  (No.  213705)  :  Basal  length, 
353;  occipito-nasal  length,  346;  palatal  length,  185;  zygomatic 
breadth,  241;  interorbital  breadth,  91.  Adult  female  (type)  :  Basal 
length,  306;  occipito-nasal  length,  284;  palatal  length,  165;  zygo- 
matic breadth,  210 ;  interorbital  breadth,  80.5. 

URSUS  WARBURTONI *  Mekriam. 

Warbtjrton  Pike  Geizzly. 

Ursus  Jcwakiutl  toarburtoni  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  p.  145, 
September  6,  1916. 

Type  locality. — Atnarko  River,  British  Columbia. 

Type  specimen. — No.  210576,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  July  15,  1915,  by  E.  H. 
Edwards. 

Range. — Coast  region  (but  perhaps  not  the  immediate  coast  strip) 
of  Southeastern  Alaska  and  adjacent  parts  of  British  Columbia  from 
Chilkat  River  southeasterly  to  Atnarko  River,  one  of  the  upper 
forks  of  the  Bella  Coola  (skulls  of  adult  males  examined  from  At- 
narko River,  Stikine  River,  Iskut  River  near  junction  with  Stikine, 
and  Chilkat  River  valley). 

Cranial  characters. — Adult  males :  Skull  large  and  massive,  rather 
long  and  flattish  on  top,  not  arched.  Similar  to  male  Jcwakiutl  but  with 
much  broader  (less  peglike)  and  flatter  postorbitals,  flatter  frontal 
shield  (not  deeply  concave  in  old  age),  much  shorter  sagittal  crest, 
somewhat  heavier  dentition,  especially  broader  and  more  massive 

1  Named  in  honor  of  the  late  Warburton  Pike,  author  of  The  Barren  Grounds  of  North- 
ern Canada,  and  The  Subarctic  Forest,  who  obtained  a  fine  large  typical  skull  (No. 
223946)  on  the  Iskut  a  few  miles  from  its  junction  with  the  Stikine. 


28  NORTH   AMERICAN   FAUNA.  [No.  41. 

M^.  Easily  distinguished  from  stikeenensis  of  the  same  region  by 
the  much  greater  length  of  skull  and  under  jaw  and  lesser  elevation  of 
frontal  region. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  340;  oc- 
cipito-nasal  length,  324 ;  palatal  length,  185 ;  zygomatic  breadth,  233  ; 
interorbital  breadth,  85.  Old  male  (No.  223946)  from  Iskut  Eiver, 
a  branch  of  the  Stikine:  Basal  length,  326;  occipito-nasal  length, 
340;  palatal  length,  176;  zygomatic  breadth,  230;  interorbital 
breadth,  86.5. 

URSUS  NEGLECTUS  Merriam. 

Admiralty  Island  Grizzly. 

Ursus  kwakiutl  neglecims  Merriam,  Proc.  Biol.   Soc.  Washington,  XXIX,  pp. 
144-145,  September  6,  1916. 

Type  locality. — Near  Hawk  Inlet,  Admiralty  Island,  Southeastern 
Alaska. 

Type  specimen. — No.  209889,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.    Collected  April,  1914,  by  W.  H.  Spaulding. 

Cranial  characters. — Skull  rather  large  and  massive,  flat  topped, 
with  rather  broad  outstanding  postorbitals.  Frontal  shield  moder- 
ate, flattish,  shallowly  sulcate,  rather  short  pointed;  fronto-nasal 
region  elevated  in  plane  of  shield  and  slightly  compressed ;  rostrum 
moderate,  high;  zygomata  moderately  outstanding  and  somewhat 
bowed;  postpalatal  shelf  short  and  broad;  notch  rather  broad. 
Underj  aw  rather  short,  ramus  rather  flat  and  broad  vertically, 
especially  posteriorly,  strongly  bellied  posteriorly;  coronoid  blade 
high  and  narrow,  its  apex  not  reaching  plane  of  condyle.  Teeth 
moderate. 

Cranial  comparisons. — Ursus  neglectus  requires  comparison  with 
both  kwakiutl  and  warburtoni.  Old  male  (type)  compared  with 
adult  male  kwakiutl  (type  from  Jervis  Inlet)  :  Size  similar;  shield 
flatter,  more  nearly  horizontal  and  less  sloping  anteriorly ;  postorbit- 
als broader  and  flatter  on  top ;  fronto-nasal  region  elevated  instead  of 
depressed;  rostrum  larger  and  more  elevated;  zygomata  less  out- 
bowed;  nasals  more  produced  anteriorly,  projecting  broadly  over 
nares  (about  7  mm.  beyond  premaxillse  at  point  of  contact) ;  nares 
more  truncate:  underjaw  and  subangular  border  much  shorter; 
coronoid  blade  narrower.  Upper  canines  and  crown  of  last  upper 
molar  longer. 

Old  male  (type)  compared  with  three  old  males  of  warburtoni 
(from  Atnarko  and  Iskut  Rivers  and  Chilkat  Valley)  :  Size  slightly 
smaller  but  occipito-sphenoid  length  same ;  top  of  skull  more  nearly 
horizontal  (shield  anteriorly  and  rostrum  more  elevated) ;  zygomata 


1918.]  HOERIBILIS  GROUP.  29 

less  outbowed;  nasals  more  projecting  anteriorly;  palate  shorter; 
postpalatal  shelf  broader,  flatter,  and  much  shorter;  mastoids 
shorter;  under  jaw  slightly  smaller;  subangular  border  much  shorter; 
coronoid  blade  narrower  and  more  nearly  vertical.  Canines  (espe- 
cially upper)  more  slender;  M^  narrower  and  less  massive  but 
difference  not  great. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  322 ;  oc- 
cipito-nasal  length,  325 ;  palatal  length,  177 ;  zygomatic  breadth,  229 ; 
interorbital  breadth,  83. 

URSUS  CALIFORNICUS  Merbiam. 
California  Coast  Grizzly. 

[Ursus  horriceus]  subspecies  califonvicus  Merriam,  Proc.  Biol.  Soc.  Washing- 
ton, X,  pp.  76-77,  April  13,  1896. 

Ursus  californicus  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  186,  188, 
August  13,  1914. 

Type  locality. — Monterey,  California. 

Type  specimen. — Skull  No.  3630,  $  old,  U.  S.  National  Museum. 

Range. — Humid  coast  region  of  California  from  San  Francisco 
Bay  south  about  to  San  Luis  Obispo  (apparently  passing  into 
tularensis  in  the  dryer  interior) . 

Characters. — Size  large;  claws  long  and  smooth;  pelage  variable. 
Dentition  heavy ;  last  upper  molar  large,  its  heel  long  and  broad. 

Cranial  characters. — Adult  male:  Skull  long  and  narrow;  vault  of 
cranium  flat — not  arched;  frontal  shield  flat  (or  concave  medially), 
short  pointed  posteriorly,  gradually  sloping  almost  in  same  plane 
with  rostrum  (dishing  slight)  ;  temporal  impressions  beaded;  postor- 
bitals  massive  and  moderately  spreading ;  zygomata  slightly  bowed ; 
rostrum  long,  slightly  compressed  in  front  of  orbits;  nasals  nearly 
horizontal,  slightly  rising  posteriorly  in  plane  of  shield;  frontal 
shield  moderately  broad,  slightly  swollen  over  orbits,  the  point  lost 
in  sagittal  crest  25  to  50  mm.  anterior  to  fronto-parietal  suture; 
postorbital  processes  strongly  developed,  subtriangular,  rather  mas- 
sive, outstanding,  and  slightly  decurved  over  orbits;  sagittal  crest 
strongly  developed  and  nearly  straight;  squamosal  shelf  broad; 
squamosal  base  of  zygoma  vertically  expanded  in  aged  skulls ;  frontal 
part  of  braincase  in  young-adult  skulls  somewhat  elevated  and  tend- 
ing to  "keel"  into  crest;  mastoid  processes  long;  interpterygoid 
fossa  long  and  usually  narrow;  under  jaw  long  and  massive,  ramus 
broad  vertically.  Teeth  large  and  heavy ;  M-3-  subrectangular,  heel 
long,  broad  posteriorly.  Skull  of  female:  Similar  to  that  of  male, 
but  smaller. 

Cranial  comparisons. — Adult  and  old  male  compared  with  adult 
male  klamathensis  (type)  :  Skull  longer ;  zygomatic  breadth  essen- 


30  NORTH  AMERICAN   FAUNA.  [No.  41. 

tially  the  same;  face  and  rostrum  longer;  rostrum  more  compressed 
in  front  of  orbits;  fronto-nasal  region  more  dished;  frontal  shield 
more  concave  medially  and  more  swollen  at  orbital  rims ;  postorbital 
processes  much  more  massive,  slightly  arched,  the  tips  slightly  de- 
curved;  temporal  beads  more  strongly  incurved;  sagittal  crest 
shorter  posteriorly;  lambdoid  crest  more  outstanding  laterally;  oc- 
cipital overhang  much  less;  squamosal  shelf  behind  zygoma  much 
shorter;  occipito-sphenoid  and  palate  longer;  mastoids  much  longer 
and  more  strongly  outstanding.  Under  jaw  much  longer.  Dentition 
heavier,  the  last  upper  and  middle  lower  molar  in  particular  much 
larger ;  heel  of  M1  very  broad  and  broadly  rounded  posteriorly. 

Old  male  from  Monterey,  compared  with  old  male  colusus  (type, 
from  Sacramento  River)  :  Frdnto-nasal  region  slightly  higher  and 
less  depressed;  rostrum  higher,  less  depressed,  and  less  horizontal; 
postorbitals  much  larger  and  more  swollen;  orbital  rims  more 
swollen ;  orbits  more  nearly  vertical,  squamosal  base  of  zygoma  more 
expanded;  palate  narrower  anteriorly;  ramus  much  broader  verti- 
cally, its  inferior  border  less  convex  in  middle  part  and  more  bellied 
posteriorly;  apex  of  coronoid  more  produced  posteriorly;  angular 
process  larger  and  longer;  mastoids  longer  and  directed  less  ante- 
riorly, not  reaching  so  near  glenoid  process.  Dentition  stronger  and 
strikingly  different :  canines  larger ;  upper  incisors  and  both  upper  and 
lower  molars  very  much  larger;  heel  of  M^  large  and  not  normally 
emarginate. 

/Skull  measurements. — Old  male  (type):  Basal  length,  361;  oc- 
cipito-nasal  length,  346;  palatal  length,  196;  zygomatic  breadth, 
224 ;  interorbital  breadth,  82. 

URSUS  TULARENSIS  Mebbiam. 

Tejon  Gbizzlt. 

Ursus  californicus  tularensis  Merriam,  Proc.  Biol.   Soc   Washington,  XXVII, 
p.  188,  August  13,  1914. 

Type  locality. — Fort  Tejon,  Canada  de  las  Uvas,  Tehachapi 
Mountains,  California. 

Type  specimen. — No.  3536,  £  old,  U.  S.  National  Museum.  Col- 
lected by  John  Xantus. 

Range. — Dry  chaparral  hills  of  interior  coast  ranges  between  the 
San  Joaquin  Valley  and  Los  Angeles  plain,  comprising  the  Teha- 
chapi, Tejon,  Sierra  Madre,  and  San  Gabriel  Ranges,  and  probably 
San  Bernardino  Mountains  also,  and  ranging  northward  an  un- 
known distance,  doubtless  covering  the  San  Rafael  and  Gabilan 
Ranges,  and  southern  part  of  the  Diablo  Range ;  limits  unknown. 

Characters. — Size  large,  but  smaller  than  californicus;  claws  of 
grizzly  type,  but  those  of  a  female  dark,  thick,  and  broad  for  a  fe- 


1918.]  HORRIBILIS  GROUP.  31 

male,  rather  straight  (tips  worn  off),  straighter  and  broader  than  in 
female  magister. 

Color. — Nearly  full-grown  male  killed  by  Walter  Richardson,  near 
head  of  Tejunga  Canyon,  San  Gabriel  Mountains,  Southern  Cali- 
fornia, in  1897:  General  color  very  dark  brown,  almost  dusky; 
grizzled  on  upperparts  by  admixture  of  pale-tipped  hairs;  muzzle 
reddish  brown. 

Cranial  characters. — Old  male:  Skull  large,  rather  broad  and  flat 
frontally,  moderately  dished,  moderately  high,  with  large  outstand- 
ing postorbitals  and  beaded  temporal  ridges.  Frontal  shield  rather 
broad,  swollen,  and  somewhat  elevated  on  orbital  rims  and  base  of 
postorbitals,  depressed  interorbitally,  sloping  gradually  into  rostrum ; 
rostrum  large  and  rather  high ;  f ronto-nasal  region  not  depressed ; 
sagittal  crest  rather  short,  elevated  and  produced  posteriorly ;  occipi- 
tal overhang  marked;  zygomata  moderately  spreading,  subtriangu- 
lar;  palate  and  postpalatal  shelf  broad  and  flat,  mastoids  rather  long 
and  inclined  strongly  forward.  Under  jaw  large  and  rather  mas- 
sive; ramus  broad  vertically,  strongly  bellied  posteriorly;  coronoid 
rather  high,  its  apex  not  strongly  recurved.  Teeth  large;  M>  sub- 
rectangular,  the  long  heel  only  slightly  emarginate  on  outer  side. 
Adult  and  old  females :  Skulls  more  easily  distinguished  than  in  males 
from  their  nearest  relative,  calif ornicus.  The  skull  of  the  female  is 
much  more  like  the  normal  female  grizzly  type,  not  resembling  the 
males  as  does  the  female  of  calif  ornicus. 

Cranial  comparisons. — Adult  and  old  females  compared  with  fe- 
male calif  ornicus:  Skull  decidedly  smaller;  frontal  shield  behind 
plane  of  postorbital  processes  more  elevated  and  convex — not  de- 
pressed and  concave  as  in  calif  ornicus;  point  of  shield  longer  and 
broader ;  orbital  rims  more  swollen ;  sagittal  crest  shorter  and  lower ; 
palate  at  least  10  mm.  shorter;  occipito-sphenoid  about  10  mm. 
shorter;  underjaw  smaller  and  lighter;  last  upper  molar  decidedly 
smaller.  Normal  M^  subrectangular  as  in  calif  ornicus  (in  henshawi 
subtriangular  and  small). 

Adult  and  old  males  compared  with  male  calif  ornicus:  Similar  but 
smaller;  base  of  cranium  shorter;  palate  slightly  shorter;  occipito- 
nasal  length  decidedly  less;  braincase  decidedly  shorter;  zygomatic 
breadth  same  or  slightly  greater — the  skull  as  a  whole  relatively 
broader  than  in  calif  ornicus;  postpalatal  shelf  broader;  underjaw 
materially  shorter;  M1  decidedly  smaller  (both  shorter  and  nar- 
rower) ;  lower  series  of  teeth  smaller,  MT  and  M^  particularly 
smaller,  much  narrower  and  less  massive;  heel  of  last  upper  molar 
less  broad  than  in  calif  ornicus. 

Adult  male  compared  with  henshawi  (type)  :  Skull  larger  and 
more  massive;  fronto-nasal  region  much  higher  and  much  less 
dished;  rostrum  larger,  higher,  and  not  depressed;  zygomata  more 
64854°— 18 3 


32  NORTH   AMERICAN   FAUNA.  [No.  41. 

broadly  spreading;  postpalatal  shelf  broader;  coronoid  larger  and 
less  falcate ;  ramus  broader  vertically ;  last  upper  molar  much  larger, 
the  heel  long  and  broad  posteriorly,  contrasted  with  the  short  sub- 
triangular  heel  of  henshawi. 

Skull  measurements. — Average  of  two  old  males  from  Fort  Tejon, 
California:  Basal  length,  329;  occipito-nasal  length,  320.5;  palatal 
length,  179.5 ;  zygomatic  breadth,  228 ;  interorbital  breadth,  78.  Old 
female  from  Fort  Tejon:  Basal  length,  296;  occipito-nasal  length, 
287;  palatal  length,  162;  zygomatic  breadth,  187;  interorbital 
breadth,  75. 

URSUS  COLUSUS  Merriam. 

Sacramento  Valley  Grizzly. 

Ursus  colusu.s  Merriara,    Proc.   Biol.    Soc.   Washington,   XXVII,   pp.   187-188, 
August  13,  1914. 

Type  locality. — Sacramento  River  valley,  California  (probably 
between  Colusa  and  Sacramento). 

Type  specimen. — No.  3837,  $  old,  U.  S.  National  Museum.  Col- 
lected by  the  Wilkes  U.  S.  Exploring  Expedition  and  marked  "  C.  P. 
Ex.  Ex.  6.16  "  (the  numerals  uncertain,  there  being  indication  of  a 
figure  in  front  of  the  first  6).  The  words  "Grizzly  Bear,  Sacra- 
mento "  are  written  on  the  right  parietal  in  pencil. 

Range. — Sacramento  (and  perhaps  also  San  Joaquin)  Valley  and 
adjacent  foothills;  westerly  in  the  hot  inner  coast  mountains  to 
Dobbins  Creek  canyon  on  the  boundary  between  southeastern  Hum- 
boldt and  southwestern  Trinity  Counties. 

Characters. — Size  large,  external  characters  unknown.  Skull 
large  and  long,  resembling  that  of  californicus,  but  teeth  smaller 
and  last  upper  molar  very  different. 

Cranial  characters. — Old  male:  Skull  large,  long,  and  low,  the 
frontal  shield  flat,  postorbital  processes  moderate  or  small,  fronto- 
nasal region  moderately  dished,  palate  long,  sagittal  crest  high, 
temporal  ridges  beaded,  and  occipital  overhang  pronounced.  In  gen- 
eral resembling  largest  skulls  of  calif omicus  but  having  very  much 
smaller  teeth  and  differing  in  numerous  minor  cranial  characters. 
Young-adult  female:  One  from  San  Jose  Mission  (No.  1143,  Yale 
Museum)  appears  to  be  a  not  quite  grown  female  colusus.  The  last 
upper  molar  is  short  and  subtriangular,  the  heel  emarginate  on 
outer  side. 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
calif ornicus  (from  Monterey)  :  Fronto-nasal  region  slightly  lower  and 
more  depressed ;  rostrum  lower,  more  depressed  and  more  nearly  hori- 
zontal ;  postorbitals  much  smaller  and  less  swollen ;  orbital  rims  less 
swollen;  orbits  less  nearly  vertical;  squamosal  base  of  zygoma  less  ex- 


1918.]  HORRIBILIS   GROUP.  33 

panded ;  palate  broader  anteriorly ;  ramus  much  less  broad  vertically, 
its  inferior  border  more  convex  in  middle  part  and  less  bellied  pos- 
teriorly ;  apex  of  coronoid  less  produced  posteriorly ;  angular  process 
smaller  and  shorter;  mastoids  shorter  and  directed  more  anteriorly, 
reaching  nearer  to  glenoid  process.  Dentition  weaker  and  strikingly 
different :  canines  too  badly  broken  to  admit  of  satisfactory  compari- 
son, but  obviously  smaller ;  upper  incisors  and  both  upper  and  lower 
molars  very  much  smaller ;  heel  of  M^  small  and  strongly  emarginate 
on  outer  side,  of  same  size  and  approximately  same  form  as  in  kla- 
mathensis  but  even  more  strongly  constricted  on  outer  side  immedi- 
ately behind  second  cusp ;  lower  molars  of  same  size  as  in  klamathen- 
sis  though  the  jaw  is  much  longer. 

Old  male  (type)  compared  with  klamathensis :  Postorbitals, 
length  of  sagittal  crest,  form  of  zygomata,  and  dentition  essentially 
the  same,  but  skull  longer;  vault  of  cranium  much  lower;  frontal 
shield  narrower  and  much  shorter;  temporal  beads  much  more 
strongly  incurved;  fronto-nasal  region  dished  instead  of  elevated; 
rostrum  much  lower,  more  depressed,  and  more  nearly  horizontal; 
ramus  longer,  less  broad  vertically ;  diastema  much  longer. 

Skull  measurements. — Old  male  (type):  Basal  length,  352;  oc- 
cipito-nasal  length,  337 ;  palatal  length,  185 ;  zygomatic  breadth,  228 ; 
interorbital  breadth,  82.5. 

URSUS  DUSORGUS  sp.  nov.1 
Rindsfoos  Grizzly. 

Type  No.  217426,  $  old,  U.  S.  National  Museum.  From  head  of 
Jack  Pine  River  near  Mount  Bess,  Alberta  (close  to  British  Columbia 
boundary).  Collected  September  4,  1916,  and  presented  to  the  Na- 
tional Museum  by  William  Rindsfoos. 

Cranial  characters. — Skull  large,  broad,  and  massive,  with  flat 
gradually  sloping  frontal  shield  and  high  sagittal  crest;  shield 
broad,  short  pointed,  slightly  thickened  at  orbits,  with  horizontally 
outstanding  slightly  elevated  postorbitals;  fronto-nasal  region 
faintly  dished,  the  broad  slightly  depressed  rostrum  rising  imper- 
ceptibly into  shield;  sinus  case  keeling  into  anterior  part  of  sagittal 
crest,  the  crest  rising  above  plane  of  top  of  skull  and  point  of  f  rontals 
and  reaching  forward  nearly  half  way  from  fronto-parietal  suture 
to  postorbitals;  temporal  impressions  strongly  beaded;  zygomata 
broadly  outstanding;  palate  and  postpalatal  shelf  rather  broad; 
underjaw  massive;  ramus  strongly  bellied  posteriorly;  coronoid 
rather  high.    Teeth  moderate;  last  upper  molar  long. 


1  Tentatively  included  in  horribilis  group.     (See  Introduction,  pp.  12-13.) 


34  NORTH   AMERICAN   FAUNA.  [No.  41. 

Cranial  comparisons. — So  far  as  known  dusorgus  requires  com- 
parison with  only  a  single  species — imperator  of  western  Montana 
and  the  Yellowstone  Park  region.  Old  male  (type)  compared  with 
old  male  imperator  (No.  216205)  :  Similar  in  general  but  size  some- 
what less;  vault  of  cranium,  frontal  region,  and  rostrum  lower; 
shield  flatter,  more  strongly  sloping,  and  shorter  pointed;  rostrum 
more  depressed;  sagittal  crest  much  higher  anteriorly,  rising  well 
above  point  of  shield;  palate  shorter;  meatus  tube  longer;  ramus  of 
jaw  shorter.     Teeth  similar  but  M-1  smaller. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  324;  oc- 
cipito-nasal  length,  322.5 ;  palatal  length,  180 ;  zygomatic  breadth, 
227;  interorbital  breadth,  86.      / 

Planiceps  Group. 

URSUS  NELSONI *  Merriam. 

Nelson  Grizzly. 

Vrsus  nelsoni  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  190-191, 
August  13,  1914. 

Type  locality. — Colonia  Garcia,  Chihuahua,  Mexico. 

Type  specimen. — No.  99657,  9  ad.,  U.  S.  National  Museum 
Biological  Survey  collection.  Collected  November  13,  1899,  by  H.  A. 
Cluff. 

Range. — Sierra  Madre  of  Mexico  from  northwestern  Chihuahua 
and  northeastern  Sonora  south  to  southern  Durango. 

Characters. — Smallest  of  the  grizzly  bears.  General  color,  pale 
buffy  yellowish,  varying  to  grayish  white,  grizzled  from  darker  color 
of  underfur.  Specimens  in  worn  pelage  vary  to  yellowish  brown  and 
even  rusty.  Hairs  of  throat  and  flanks  longer  than  elsewhere ;  belly 
sparsely  haired,  lacking  the  thick  underfur  of  upperparts.  Claws 
long,  smooth,  and  moderately  curved;  brownish  horn  color  streaked 
with  yellowish.  Longest  claw  56  mm.  from  upper  base  to  tip  (tip 
worn  off). 

Color. — Type  specimen  in  fresh  fall  pelage:  Muzzle  pale  brown, 
much  darker  around  eyes;  top  of  head  yellowish  buff;  back  grayish 
brown,  heavily  overlaid  with  pale  buffy  gray  tips  (color  more  buffy 
across  shoulders,  more  whitish  gray  on  back)  ;  hump  dark  brown, 
small;  rump  grizzled  grayish  and  brown,  the  light-tipped  hairs 
failing  posteriorly  and  on  the  sides,  the  dark  brown  ground  color 
passing  into  blackish  brown  on  thighs,  legs,  feet,  and  tail;  forelegs 
also  blackish  brown;  lips  and  point  of  chin  yellowish  buff,  followed 
by  area  of  dark  brown;  underneck  and  underparts  generally  long 
haired  and  grizzled,  the  prevailing  color  yellowish  buff. 

*  Named  for  E.  W.  Nelson,  who  collected  the  original  series  in  the  type  region. 


1918.]  PLANICEPS  GROUP.  36 

Cranial  characters. — Adult  male:  Skull  small  and  wolflike.  Simi- 
lar in  general  to  texensis,  but  smaller  and  more  wolflike;  rostrum 
strikingly  narrow;  frontal  shield  flat,  narrow,  and  only  faintly 
sulcate  medially;  temporal  impressions  not  beaded;  occipito-sphe- 
noid  length  82  mm.  (in  $  texensis,  86) ;  palate  very  short  (149  mm. 
contrasted  with  111  in  texensis)  ;  posterior  root  of  zygoma  slender; 
postorbital  process  slender,  peglike,  and  outstanding  horizontally. 
Teeth  small. 

Skull  measurements.— Old  male  (No.  16025,  Field  Mus.  Nat.  Hist., 
Chicago,  from  Casas  Grandes,  Chihuahua):  Basal  length,  284; 
occipito-nasal  length,  264;  palatal  length,  149;  zygomatic  breadth, 
199;  interorbital  breadth,  65.  Average  of  three  adult  females  from 
Colonia  Garcia,  Chihuahua :  Basal  length,  260 ;  occipito-nasal  length, 
243;  palatal  length,  138;  zygomatic  breadth,  168;  interorbital 
breadth,  61. 

URSUS  TEXENSIS  TEXENSIS  Merbiam. 

Texas  Gbizzly. 

Ursus  horriceus  texensis  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  191, 
August  13,  1914. 

Type  locality. — Davis  Mountains,  Texas. 

Type  specimen. — No.  203198,  $  old,  U.  S.  National  Museum, 
Biological  Survey  collection.  Killed  November  2,  1890,  by  C.  O. 
Finley  and  John  Z.  Means. 

Range. — Restricted,  so  far  as  known,  to  Davis  Mountains,  Texas, 
and  mountains  of  southern  Colorado. 

Characters. — Size  small;  external  characters  unknown.  Affinities 
with  shoshone  rather  than  with  horriceus. 

Cranial  characters. — Old  male  (type)  :  Size  small;  frontal  shield 
low,  narrow,  flat  (slightly  depressed  medially),  short  pointed  pos- 
teriorly, gently  sloping  in  same  plane  with  rostrum;  postorbital 
processes  small,  peglike,  horizontally  outstanding;  fronto-nasal  re- 
gion elevated  in  fronto-facial  plane;  rostrum  narrow;  zygomata 
rather  broadly  outbowed,  moderately  expanded  vertically,  lachrymal 
duct  wholly  anterior  to  orbit ;  sagittal  crest  long  and  nearly  straight ; 
occipital  overhang  marked;  palate  broad,  deeply  excavated  between 
molars  (may  be  abnormal)  ;  postpalatal  shelf  broad  and  flat;  post- 
palatal  notch  broad  and  short;  meatus  tube  slightly  recurved  and 
markedly  upturned.  Under  jaw  light;  inferior  border  of  ramus 
long,  slightly  bellied  posteriorly;  subangular  border  rather  short; 
coronoid  blade  high,  its  recurved  apex  falling  short  of  plane  of 
condyle.  Teeth  rather  small;  M-2-  small,  with  small  heel  obliquely 
narrowed  on  outer  side. 

Old  female  (No.  213002,  from  Navajo  Range,  near  Cromo,  Colo- 
rado) :  Skull  short,  dished,  the  braincase  broad;  frontal  shield  nar- 


36  NORTH   AMERICAN   FAUNA.  [No.  41. 

row ;  zygomata  broadly  spreading,  slightly  bowed ;  occiput  truncate ; 
and  postorbitals  slender,  widely  outstanding.  Rostrum  small,  slen- 
der, tapering,  rising  posteriorly  into  flat  lyrate  shield;  postorbital 
processes  long,  slender,  and  directed  forward  as  well  as  outward; 
braincase  rather  broad  and  depressed;  palate  rather  short;  post- 
palatal  shelf  broad  and  flat.  Teeth  moderate;  canines  small  and 
slender;  molars  nearly  as  large  as  in  male,  and  M>  of  same  form. 

Cranial  comparisons. — Adult  male  (type)  compared  with  adult 
male  planiceps  (type)  :  Size  smaller  throughout;  shield  much  nar- 
rower, less  flat,  and  less  nearly  horizontal ;  f  ronto-nasal  region  dished 
(in  planiceps  not  dished);  rostrum  much  smaller  and  narrower; 
palate  deeply  concave  between  posterior  molars  (in  planiceps  flat) ; 
ramus  of  under  jaw  more  tapering  anteriorly;  angular  processes  not 
bellied  (in  planiceps  bellied) ;  teeth  smaller. 

Old  male  (type)  compared  with  male  shoshone:  Size  smaller; 
vault  of  cranium  lower  and  more  nearly  horizontal;  frontal  shield 
less  sloping;  anterior  part  of  braincase  broadly  depressed  (not 
compressed  or  keeling  into  crest)  ;  palate  broader.    Teeth  smaller. 

Old  male  (type)  compared  with  old  male  horriceus  (type)  :  Skull 
similar  in  basal  length,  but  materially  smaller,  lower,  flatter,  nar- 
rower, more  smoothly  rounded,  broader  across  squamosals,  much 
less  massive,  and  with  wholly  different  postorbitals.  Postorbitals 
small,  slender,  peglike,  and  horizontally  outstanding  instead  of 
large,  broad,  massive,  and  decurved;  frontal  shield  narrower  and 
flatter;  orbital  rims  less  swollen;  rostrum  smaller  (shorter  and  more 
slender)  ;  fronto-nasal  region  even  less  dished ;  palate  somewhat 
broader  between  molars  and  deeply  excavated  between  last  molars 
(slightly  depressed  in  horriceus)  ;  interpterygoid  canal  shorter;  zygo- 
mata more  widely  outstanding  posteriorly  and  bowed  (in  horriceus 
more  angular  and  more  outstanding  anteriorly) ;  inferior  border  of 
jaw  (symphysis  to  subangular  tubercle)  much  longer.  Canines  and 
molars  smaller. 

Old  female  (from  Navajo  Mountains)  compared  with  old  female 
horriceus  (from  mountains  north  of  Silver  City,  N.  Mex.)  :  Skull  as 
a  whole  and  rostrum  shorter;  occiput  doubtless  more  truncate  (that 
of  horriceus  sawed  off)  frontal  shield  shorter  and  more  dished; 
postorbitals  longer,  more  slender,  and  directed  anteriorly  as  well  as 
outward;  zygomata  much  more  broadly  spreading,  more  swollen  at 
anterior  base,  and  distinctly  bowed  instead  of  angular;  palate 
broader,  flat  instead  of  concave;  lower  jaw  thicker  and  heavier. 

Skull  measurements.— Old  male  (type) :  Basal  length.  308 ;  occi- 
pito-nasal  length,  301;  palatal  length,  171;  zygomatic  breadth,  218; 
interorbital  breadth,  71. 


1918.]  PLANICEPS  GROUP.  37 

URSUS  TEXENSIS  NAVAHO  Merriam. 

Navaho  Grizzly. 

Ursus   navaho   Merriam,   Proc.   Biol.    Soc.   Washington,   XXVII,  pp.    191-192, 
August  13,  1914. 

Type  locality. — Navajo  country  near  Fort  Defiance,  Arizona  (Moll- 
hausen) ;  type  probably  killed  in  1856  in  Chuska  Mountains,  on  boun- 
dary between  northeastern  Arizona  and  northwestern  New  Mexico. 

Type  specimen. — No.  3500,  $  old,  U.  S.  National  Museum. 

Range. — Probably  restricted  to  the  isolated  Chuska  Mountains 
(including  the  so-called  Lukachukki  and  Tunitcha  elevations,  and 
perhaps  also  the  neighboring  Carriso  Mountains  on  the  north). 

Characters. — Size  small;  external  characters  unknown;  skull 
short,  broad  and  slightly  dished. 

Cranial  characters. — Old  male  (type;  badly  damaged,  only  the 
front  part  with  zygomata  and  underjaw  remaining)  :  Skull  short; 
zygomata  broadly  spreading  and  outbowed,  the  anterior  root  much 
swollen,  posterior  root  not  expanded;  frontal  shield  flat,  short 
pointed;  postorbital  processes  peglike  and  outstanding;  temporal 
impressions  finely  beaded;  rostrum  short  and  small;  nares  rather 
small  and  nearly  vertical;  lachrymal  duct  opening  on  orbital  rim, 
but  more  in  than  out;  palate  broad  for  so  small  a  skull;  postpalatal 
shelf  broad ;  coronoid  blade  high.  Teeth  small :  M1  very  small ;  M-2- 
short,  with  broadly  rounded  heel  (tooth  nearly  as  broad  posteriorly 
as  in  middle  and  anteriorly). 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
texensis  (type)  :  Size  somewhat  larger;  frontal  shield  and  rostrum 
broader;  jugal  longer  posteriorly.  Palate  and  postpalatal  shelf 
much  shorter  (157  mm.  contrasted  with  173),  and  less  deeply  concave 
between  posterior  molars ;  postorbital  processes  slightly  larger ;  ante- 
rior root  of  zygoma  much  more  swollen  (in  texensis  hardly  swollen)  ; 
nares  much  more  truncate;  underjaw  straighter  (less  upcurved 
posteriorly)  ;  canines  about  same  size;  molar  series  slightly  shorter; 
M.2-  of  same  length  but  heel  broadly  rounded  posteriorly  instead  of 
obliquely  truncate,  the  sides  of  tooth  nearly  parallel;  M1,  MT,  and 
PMA  smaller. 

Skull  measurements. — Old  male  (type)  :  Palatal  length,  157.5 ; 
interorbital  breadth,  81.  Skull  too  badly  broken  to  furnish  other 
measurements. 

URSUS  PLANICEPS  sp.  nov. 
Flat-Headed  Grizzly. 

Type  No.  13289,  $  ad.  (rather  old),  U.  S.  National  Museum,  from 
Colorado  (exact  locality  unknown).    Collected  by  Dr.  F.  V.  Hayden.1 

1  Dr.  Hayden  worked  in  Colorado  in  1869,  mainly  in  the  mountains  and  foothills  of  the 
east-central  part  of  the  State.  Inasmuch  as  two  other  grizzlies,  hairdi  and  macrodon, 
inhabit  the  higher  mountains  of  Colorado  it  seems  highly  probable  that  the  home  of 
planiceps  was  in  the  foothills  or  along  the  western  edge  of  the  plains. 


38  NORTH   AMERICAN   FAUNA.  [No.  41. 

Cranial  characters. — Adult  male  (type)  :  Skull  rather  large,  low, 
and  flat ;  shield  broad,  flat,  and  nearly  horizontal ;  postorbitals  hori- 
zontally outstanding ;  rostrum  broad,  somewhat  depressed ;  zygomata 
broadly  outstanding  and  outbowed;  palate,  postpalatal  shelf,  and 
postpalatal  notch  broad;  occipito-sphenoid  long  (95  mm.)  Under- 
jaw  rather  long;  ramus  flat  and  rather  thin;  condyle  high,  vertical; 
subangular  border  short;  angular  process  bellied  on  underside. 
Canines  moderate;  molars  rather  large  but  much  smaller  than  in 
macrodon. 

Cranial  comparisons. — Ursus  planiceps  requires  comparison  with 
its  two  neighbors,  macrodon  and  texensis,  of  which  its  affinities  are 
closest  with  macrodon.  Old  male  (type)  compared  with  old  male 
macrodon:  Size  and  vault  of  cranium  about  same;  shield  slightly 
broader  and  flatter;  rostrum  broader;  zygomata  very  much  more 
broadly  outstanding  and  outbowed;  palate,  postpalatal  shelf,  and 
palatal  notch  much  broader;  posterior  part  of  underjaw  more  up- 
curved,  lifting  condyle  and  coronoid ;  upper  canines  about  same  size ; 
lower  canines  and  upper  and  lower  molars  decidedly  smaller. 

Old  male  (type)  compared  with  old  male  texensis  (type)  :  Size 
larger  throughout;  shield  much  broader,  flatter,  and  more  nearly 
horizontal ;  f ronto-nasal  region  more  elevated  in  plane  of  shield  (not 
dished  as  in  teaensis) ;  rostrum  much  larger,  broader,  and  more  mas- 
sive; zygomata  about  the  same  but  squamosal  arm  longer;  palate 
flat  (in  texensis  deeply  concave  between  posterior  molars) ;  occipito- 
sphenoid  longer;  underjaw  much  more  massive;  ramus  less  tapering 
anteriorly;  angular  processes  strongly  bellied  (in  texensis  not  bel- 
lied) ;  teeth  larger. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  319.5;  occi- 
pito-nasal  length,  305 ;  palatal  length,  173 ;  zygomatic  breadth,  215 ; 
interorbital  breadth,  75. 

URSUS  MACRODON  sp.  nov. 
Twin  Lakes  Grizzly. 

Type  skull  No.  15707  (skin  No.  12678),  $  old,  U.  S.  National 
Museum.  From  Twin  Lakes,  Colorado,  July  28,  1876.  Collected  by 
C.  W.  Derry. 

Characters. — Old  male  (type)  :  Size  large;  hump  evident;  claws 
(worn  short)  large,  broad,  mainly  yellowish  on  top,  horn  color  on 
sides.    Skull  rather  large  and  flat ;  last  upper  molar  very  large. 

Color. — Type  specimen:  Peculiar  for  a  grizzly,  resembling  some 
of  the  Alaska  brown  bears;  general  body  color  rich  brown,  almost 
reddish  brown,  darker  on  hump  and  lightly  washed  with  pale  tipped 
hairs  on  upper  part  of  back;  head  rather  uniform  brown.  Adult 
female  (No.  203178)  from  South  Branch  Williams  Kiver,  Colorado, 


1918.]  PLANICEPS   GROUP.  39 

killed  September  13,  1895 :  General  color  very  dark,  almost  black. 
Muzzle  reddish  brown  with  a  golden  tinge  on  top,  becoming  very 
dark  brown  on  cheeks,  chin,  and  around  eyes ;  top  of  head  dusky,  over- 
laid by  deep  rich  glossy  hazel  or  between  hazel  and  chestnut;  top  of 
neck  and  upperparts  generally  blackish,  moderately  washed  with 
golden-tipped  hairs;  hump  marked  and  nearly  black;  legs  and  feet 
nearly  black ;  the  long  hairs  below  ears  and  on  sides  of  neck  washed 
with  golden.  Claws  long,  slender,  and  smoothly  polished;  dark 
horn  color,  paler  toward  tips  and  on  sides.  Longest  claw  from  upper 
base  61  mm.  (tip  worn  off;  greatest  breadth  of  claws  8  mm.). 

Cranial  characters. — Old  male  (type)  :  Skull  long,  low,  flat,  and 
rather  narrow;  shield  narrow,  flat,  nearly  horizontal,  sloping  grad- 
ually into  rostrum  without  noticeable  dishing ;  zygomata  not  broadly 
outstanding;  squamosal  root  long  but  not  vertically  expanded;  palate 
long  and  narrow;  postpalatal  notch  narrow;  underjaw  long;  ramus 
straight,  swollen  on  outer  side  below  premolars ;  coronoid  blade  high ; 
subangular  border  rather  short ;  angular  process  bellied ;  canines  and 
molars  large ;  M-2-  40  mm.  in  length ;  heel  long,  slightly  emarginate. 

Female  from  South  Branch  Williams  River,  Colorado  (No.  203178, 
September  13,  1895,  collected  and  presented  by  J.  P.  Bird)  :  Skull 
similar  to  that  of  male  but  much  smaller,  with  correspondingly  nar- 
rower shield  and  rostrum,  and  slightly  more  dished  fronto-nasal  re- 
gion; shield  low,  narrow,  flat,  and  gently  sloping,  the  point  slender 
and  remarkably  short  .for  a  female,  ending  anterior  to  fronto-parietal 
suture;  sagittal  crest  proportionately  long  but  low  anteriorly;  post- 
orbital  processes  small,  slightly  elevated,  and  directed  slightly  back- 
ward; orbital  rims  somewhat  thickened;  rostrum  small;  nasals  an- 
teriorly horizontal,  posteriorly  rising  slightly  and  passing  into  frontal 
shield  in  same  plane ;  zygomata  moderately  spreading,  angular ;  palate 
and  postpalatal  shelf  short.  Canines  very  small;  molars  large;  M> 
38  mm.,  which  is  materially  larger  than  in  males  of  texensis  and 
planiceps. 

Cranial  comparisons. — Skull  of  old  male  (type)  similar  in  general 
to  old  male  texensis  (type)  but  slightly  larger,  with  longer,  flatter,  and 
more  nearly  horizontal  frontal  shield,  and  somewhat  higher  fronto- 
nasal region  and  rostrum;  shield  somewhat  longer  pointed;  fronto- 
nasal region  elevated  in  plane  of  shield  (not  dished  as  in  texensis) ; 
postorbitals  more  broadly  outstanding;  rostrum  broader  and  some- 
what longer ;  opening  of  lachrymal  duct  within  orbital  rim  (in  texen- 
sis anterior  to  rim);  zygomata  less  widely  outstanding;  squamosal 
root  longer;  postpalatal  shelf  narrower;  palate  flatter;  underjaw  more 
massive;  ramus  less  tapering  anteriorly;  inferior  border  of  angular 
process  strongly  bellied  or  keeled  (may  be  individual) ;  canines  and 
molars  very  much  larger  throughout  (especially  M>). 


40  NORTH   AMERICAN   FAUNA.  [No.  41. 

Remarks. — In  size  and  general  appearance  macrodon  resembles 
planiceps,  but  the  frontal  shield  is  more  nearly  horizontal ;  postorbital 
processes  less  widely  outstanding;  zygomata  much  less  broadly  out- 
standing and  not  howed;  squamosal  arm  of  zygoma  longer;  canines 
and  molars,  particularly  M-2- ,  decidedly  larger.  It  may  prove  to  inter- 
grade  with  planiceps. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  312 ;  occipito- 
nasal  length,  302 ;  palatal  length,  171 ;  zygomatic  breadth,  202 ;  inter- 
orbital  breadth,  72. 

URSUS  MIRUS  sp.  nov. 

Yellowstone  Pabk  Grizzly. 

/ 

Type  No.  206595,  $  ad.  (rather  old),  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  From  Slough  Creek,  Yellowstone  Na- 
tional Park,  March  27,  1915.    Collected  by  Henry  Anderson. 

Cranial  characters. — Old  male  (type)  :  Size  medium;  skull  long, 
rather  narrow  and  low  arched ;  shield  rather  narrow  and  flat,  nearly 
horizontal  postorbitally,  anteriorly  sloping  gently  into  rostrum; 
postorbitals  long,  slender,  and  horizontally  outstanding;  rostrum 
high,  subterete  above ;  nares  high ;  zygomata  widely  outbowed ;  squa- 
mosal root  arched  and  broadly  expanded;  palate  and  postpalatal  shelf 
narrow ;  mastoids  vertical ;  underjaw  moderate,  swollen  on  lower  part 
of  outer  side  under  diastema;  subangular  border  rather  long  and 
sloping  upward  posteriorly.    Teeth  of  medium  size. 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
tahltanicus  (type)  :  Basal  length  and  vault  of  cranium  essentially 
same ;  occipito-nasal  length  greater ;  shield  and  rostrum  flatter ;  zygo- 
mata more  outbowed;  squamosal  arm  of  zygoma  longer  and  more 
broadly  expanded;  underjaw  slightly  longer;  coronoid  blade  nar- 
rower in  middle  part ;  inferior  border  of  ramus  of  same  length ;  sub- 
angular  border  longer. 

Old  male  (type)  compared  with  old  male  planiceps  (type)  :  Size 
essentially  same  except  that  the  rostrum  is  decidedly  narrower  and 
more  elevated  anteriorly;  point  of  shield  slightly  more  elevated; 
shield  more  sloping;  postorbitals  longer;  rostrum  narrower,  higher, 
and  subterete  instead  of  flattened  above ;  palate  and  postpalatal  shelf 
narrower ;  zygomata  even  more  strongly  outbowed ;  occipito-sphenoid 
shorter;  subangular  border  of  lower  jaw  longer;  angular  processes  not 
bellied;  teeth  about  same  size. 

Old  male  (type)  compared  with  old  male  texensis  (type)  :  Similar 
in  general  characters  but  somewhat  larger;  vault  of  cranium  slightly 
higher ;  rostrum  much  higher  anteriorly  and  more  nearly  horizontal ; 
nares  much  higher ;  postorbitals  more  slender  and  much  more  widely 
outstanding  (121  contrasted  with  104) ;  posterior  frontal  region  less 


1918.]  PLANICEPS   GEOUP.  41 

elevated ;  zygomata  more  outbowed  (less  triangular) ;  squamosal  root 
of  zygoma  much  more  broadly  expanded ;  postpalatal  shelf  narrower; 
underjaw  larger  and  longer;  ramus  less  tapering  anteriorly;  sub- 
angular  border  somewhat  longer. 

Old  male  (type)  compared  with  old  male  shoshone  (type),  with 
which  it  agrees  in  basal  length:  Vault  of  cranium  lower;  shield 
lower  posteriorly  and  much  less  strongly  sloping;  rostrum  higher 
anteriorly  and  more  nearly  horizontal;  nares  higher;  sinus  case  not 
definitely  keeling  into  sagittal  crest;  zygomata  much  more  broadly 
outstanding  and  outbowed;  squamosal  root  much  more  broadly  ex- 
panded ;  postpalatal  length  less ;  canines *  and  molars  closely  similar. 

Compared  with  old  male  idaJwensis  (type)  :  Skull,  palate,  and 
rostrum  longer ;  basicranium  and  vault  less  arched ;  squamosal  arm  of 
zygomata  longer  and  more  broadly  expanded. 

Remarks. — The  cranial  characters  indicate  that  mints  of  the 
Yellowstone  Park  region  is  rather  closely  related  to  tahltanicus  of 
the  Stikine  region,  and  also,  though  apparently  less  closely,  to 
planiceps  of  Colorado  and  southern  Wyoming.  It  is  so  much  smaller 
and  has  teeth  so  much  smaller  than  horribilis  and  imperator  that 
comparison  in  detail  is  unnecessary;  and  compared  with  absarokus 
the  skull  is  so  much  lower,  flatter,  and  narrower  and  the  teeth  so 
much  smaller  that  the  two  can  not  be  confused. 

Skull  measurements. — Old  male  (type):  Basal  length,  315;  oc- 
cipito-nasal  length,  307 ;  palatal  length,  174 ;  zygomatic  breadth,  220 ; 
interorbital  breadth,  77. 

URSUS  ELTONCLARKI  ■  Merbiam. 

Sitka  Grizzly. 

Ursus  eltonclarJci  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  175-176, 
August  13,  1914. 

Type  locality. — Near  Freshwater  Bay,  Chichagof  Island,  the  more 
northern  of  the  Sitka  Islands,  Alaska. 

Type  specimen. — No.  179066,  $  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  May  19,  1912,  by  Elton  Clark 
and  by  him  presented  to  the  Biological  Survey. 

Range. — The  Sitka  Islands,  Baranof  and  Chichagof. 

Characters. — A  grizzly  of  medium  or  rather  small  size;  skull 
small,  long,  narrow,  and  rather  low,  with  flat  frontal  shield.  Claws 
of  true  grizzly  type,  smoothly  polished,  strongly  curved  and  rather 
short:  longest  claw  (in  type  specimen)  from  upper  base  70  mm.; 
dark  bluish  or  plumbeous  horn  color  streaked  with  whitish  or 
yellowish. 


1  In  the  type  specimen  of  shoshone  the  canines  are  absent,  but  they  are  present  in  a 
young  male,  No.  113410,  from  Marvine,  Colorado,  affording  the  desired  comparisons. 
*  Named  in  honor  of  Elton  Clark,  of  Boston,  who  killed  and  presented  the  type  specimen. 


42  NORTH  AMERICAN   FAUNA.  [No.  41. 

Color. — Color  of  type  very  dark  and  rich.  Nose  pale  brown,  dark- 
ening just  in  front  of  eyes;  face,  head,  and  throat  rich  dark  choco- 
late brown,  with  golden-brown  wash  in  front  of  ears ;  ears  and  patch 
under  each  ear  dusky;  occiput  and  neck  grizzled  golden-brown; 
back  pale,  overlaid  by  buffy  tips ;  legs  and  feet  varying  from  blackish 
brown  to  brownish  black. 

Cranial  characters. — Adidt  male  (type)  :  Skull  elongate,  narrow ; 
zygomata  moderately  spreading,  outbowed,  rounded  posteriorly, 
squamosal  part  not  vertically  expanded ;  frontal  shield  in  same  plane 
with  rostrum,  narrow,  low,  flat,  or  slightly  concave,  acutely  rather 
short  pointed  posteriorly,  the  point  entering  sagittal  crest  about  one- 
third  the  distance  from  fronto-parietal  suture  to  postorbital 
process;  sagittal  crest  moderate,  reaching  more  than  halfway  from 
occiput  to  postorbital  processes;  postorbital  processes  rather  thick, 
outstanding;  fronto-nasal  region  elevated  (not  dished) ;  rostrum 
high  and  sloping  gently  upward  in  plane  of  frontals;  lachrymal 
opening  within  orbit;  palate  long  and  narrow;  postpalatal  shelf 
long;  postpalatal  notch  long  and  narrow;  occipito-sphenoid  short, 
about  80  mm. ;  basisphenoid  deeply  concave,  without  trace  of  median 
ridge.  Under  jaw  long.  Teeth  moderate;  canines  rather  long;  PM¥ 
with  moderately  sloping  heel  slightly  upturned  at  tip,  sulcus  very 
shallow;  MT  rather  short,  with  cusplet  on  inner  side  of  saddle 
posteriorly;  My  with  anterior  moiety  decidedly  longer  than  pos- 
terior and  twin  cusps  of  entoconid  small  and  not  deeply  notched; 
PMA  large  and  rather  broad ;  M1  broad  and  rather  short ;  M-2-  broad 
in  anterior  half,  then  narrowing  strongly,  the  heel  obliquely  truncate 
on  outer  side ;  cusps  rather  weak. 

Adult  female:  Similar,  but,  much  smaller,  distinctly  dished,  point 
of  shield  lyrate,  zygomata  more  angular. 

Cranial  comparisons. — Adult  male  (type)  compared  with  adult 
orgilos  (type)  :  Size  essentially  same;  vault  of  cranium  higher;  ros- 
trum and  fronto-nasal  region  longer  and  more  elevated;  postorbital 
processes  heavier  and  shorter;  occipito-sphenoid  much  shorter  (80 
mm.  contrasted  with  90) ;  mandible  more  massive.  Canines  larger 
and  longer;  My  shorter;  M1  and  MA  shorter  and  broader  (M* 
broader  in  middle.) 

Remarks. — Ursus  eltonclarki  falls  in  the  tahltanicus  group.  Oddly 
enough,  it  resembles  mirus,  a  geographically  remote  member  of  the 
group  from  the  Yellowstone  Park  country,  much  more  closely  than  it 
does  tahltanicus.  It  agrees  with  mirus  essentially  in  narrowness  of 
skull  as  a  whole,  elevation  and  narrowness  of  rostrum  and  narrow- 
ness of  palate,  and  even  exceeds  mirus  in  narrowness  of  shield  and 
postpalatal  notch.  Even  the  under  jaw  agrees  surprisingly  with  that 
of  mirus,  and  the  molar  series  are  of  approximately  the  same  length, 
but  the  lower  canines  are  materially  larger. 


1918.J 


PLANICEPS   GROUP.  43 


Skull  measurements. — Adult  male  (type)  :  Basal  length,  316; 
occipito-nasal  length,  322;  palatal  length,  173;  zygomatic  breadth, 
215;  interorbital  breadth,  69. 

URSUS  TAHLTANICUS  Meeriam. 

Tahltan  Grizzly. 

Ursus  tahltamcus  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  181-182, 
August  13,  1914. 

Type  locality.— Klappan  Creek  (=Third  South  Fork  Stikine 
River),  British  Columbia. 

Type  specimen. — No.  179928,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  in  September,  1906,  and  pre- 
sented by  G.  Frederick  Norton. 

Range. — Middle  and  upper  Stikine-Skeena  region,  limits  uncer- 
tain. 

Characters. — Size  medium;  skull  of  male  slightly  smaller  than 
male  stikeenensis  and  canadensis;  skull  of  female  about  same  size  as 
female  stikeenensis,  decidedly  larger  than  female  canadensis.  Color 
of  type  very  dark. 

Color. — Type  specimen:  Black  (head  absent,  but  entire  body  and 
legs  almost  coal  black,  lightly  grizzled  on  shoulders  and  anterior 
part  of  back  by  tips  of  golden  brownish). 

Cranial  characters. — Adult  and  old  males:  Rostrum  rather  narrow 
(in  old  age  compressed  in  front  of  orbits)  ;  vault  of  cranium  low, 
flattish;  frontal  shield  of  moderate  breadth,  flat  or  depressed  (some- 
times sulcate  medially),  rather  short  pointed,  sloping  gradually  to 
plane  of  muzzle,  and  only  slightly  dished  in  fronto-nasal  region; 
postorbital  processes  peglike,  horizontally  far  outstanding;  sagittal 
crest  low,  reaching  forward  a  little  more  than  halfway  from  inion 
to  postorbitals ;  zygomata  broadly  spreading  and  bowed;  palate  of 
medium  breadth;  postpalatal  shelf  broad  and  flat;  underjaw  short — 
much  shorter  than  in  stikeenensis,  shoshone,  and  canadensis — its 
ramus  bellied  and  upcurved  posteriorly,  not  flattened.  Teeth  rather 
small ;  canines  and  molars  much  smaller  than  in  stikeenensis. 

Female:  Skull  relatively  large  (about  same  length  as  female 
stikeenensis  but  much  narrower) ;  much  larger  than  female  sho- 
shone and  canadensis;  vault  of  cranium  moderately  elevated,  with 
tendency  to  a  fronto- parietal  hump;  facial  part  of  skull  large  and 
long;  frontal  shield  long,  rather  narrow,  lyre  shaped  posteriorly, 
meeting  sagittal  crest  some  distance  posterior  to  fronto-parietal  su- 
ture; fronto-nasal  region  well  dished;  palate  and  postpalatal  shelf 
long;  zygomata  moderately  spreading  and  conspicuously  outbowed; 
underjaw  long.  Young  females  are  best  told  by  the  teeth,  especially 
the  lower  canines.     In  tahltanicus  the  lower  canines  are  shorter  and 


44  NORTH   AMERICAN   FAUNA.  [No.  41. 

more  slender  than  those  of  stikeenensis.  In  the  latter  species  they 
are  longer  and  thicker,  especially  thick  in  middle. 

Cranial  comparisons. — Old  male  (type)  compared  with  adult  male 
orgilos  (type)  :  Size  essentially  same  (basal  length  slightly  less)  ; 
top  of  skull  higher;  frontal  shield  broader  and  less  flat;  postorbital 
processes  less  slender ;  palate  and  postpalatal  shelf  shorter ;  zygomata 
less  bowed  but  more  widely  outstanding;  underjaw  shorter,  much 
more  massive;  inferior  border  of  ramus  shorter  and  more  bellied 
posteriorly;  coronoid  blade  more  recurved. 

Adult  male  compared  with  adult  male  stikeenensis  (both  inhabit- 
ing the  same  region)  :  Basal  length,  zygomatic  breadth,  and  frontal 
breadth  essentially  the  same,  but/frontal  region  much  lower,  frontal 
shield  not  rising  abruptly  at  orbits,  but  sloping  gently  in  plane  of 
rostrum;  rostrum  narrower,  higher,  and  less  nearly  horizontal  (ap- 
pearing longer)  ;  postorbitals  much  smaller  and  more  horizontally 
outstanding;  palate  shorter;  underjaw  shorter.    Dentition  lighter. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  305 ; x  occi- 
pito-nasal  length,  304 ;  palatal  length,  163 ;  zygomatic  breadth,  220 ; 
interorbital  breadth,  84. 

URSUS  INSULARIS  Merriam. 
Island  Gbizzly. 

Ursus  eltonclarhi  insularis  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  p. 
141,  September  6,  1916. 

Type  locality. — Admiralty  Island,  Alaska. 

Type  specimen. — No.  205186,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection,  1914.  Purchased  from  W.  H.  Case,  of 
Juneau. 

Characters. — An  island  grizzly,  apparently  related  to  tahltanicus 
and  orgilos  of  the  mainland,  and  to  eltonclarki  of  Baranof  and  Chich- 
agof  Islands;  in  some  respects  a  miniature  of  hoots.  PM^  indis- 
tinctly of  grizzly  type.    External  characters  unknown. 

Cranial  characters. — Adult  male  (type)  :  Size  medium;  frontal 
shield  broad  and  flattish,  with  exceptionally  large,  broad,  and  flat 
postorbitals;  zygomata  broadly  outstanding;  ramus  of  underjaw 
strongly  upturned  posteriorly,  elevating  coronoid  and  condyle.  Teeth 
of  medium  size. 

Cranial  comparisons. — Adult  male  (type)  compared  with  old  male 
tahltanicus  (type)  :  Basal  length,  zygomatic  breadth,  breadth  and 
form  of  frontal  shield,  breadth  across  postorbitals,  truncation  of 
nasals,  and  length  of  jaw  essentially  same,  but  postorbitals  very 
much  larger,  especially  broader  and  more  decurved;   fronto-nasal 

Restored. 


1918.]  PLANICEPS   GROUP.  45 

region  more  dished;  rostrum  slightly  longer;  palate,  postpalatal 
shelf,  and  postpalatal  notch  narrower;  underjaw  more  massive  and 
more  upturned  posteriorly;  ramus  longer;  coronoid  much  higher; 
subangular  border  shorter.  Molar  series  of  approximately  same 
length  but  more  massive ;  M1  longer,  M2-  shorter. 

Adult  male  (type)  compared  with  adult  male  eltonclarki  (type)  : 
Similar  in  general,  agreeing  essentially  in  basilar  length,  zygomatic 
breadth,  length  and  narrowness  of  palate  and  postpalatal  shelf,  and 
narrowness  of  postpalatal  notch,  but  differing  widely  in  other  charac- 
ters. Vault  of  cranium  less  highly  arched ;  frontal  shield  and  post- 
orbitals  very  much  broader  and  more  massive  (shield  interorbitally 
82  mm.  contrasted  with  69),  less  flat,  faintly  sulcate  medially;  post- 
orbital  processes  very  much  larger,  broader,  more  widely  outstanding 
(from  tip  to  tip  120  mm.  contrasted  with  101)  and  more  decurved; 
rostrum  lower,  broader,  and  shorter;  nasals  shorter  (89  contrasted 
with  105 — probably  not  constant).  Underjaw  more  massive;  inferior 
border  of  ramus  longer,  more  swollen  and  more  upcurved  posteriorly ; 
outer  side  of  ramus  not  depressed  or  excavated  below  anterior  base 
of  coronoid;  coronoid  blade  narrower  and  higher.  Canines  (both 
upper  and  lower)  somewhat  shorter;  molars,  especially  M-1,  My,  and 
Mj  decidedly  larger. 

Compared  with  orgilos  and  tahltanicus  of  the  mainland:  Easily 
distinguished  by  great  size  of  postorbitals  and  upturning  of  posterior 
part  of  ramus. 

Compared  with  kwakiutl:  Quickly  told  by  general  shortness  of 
skull,  including  palate  and  underjaw,  and  by  large  size  and  breadth 
of  postorbitals. 

SkuU  measurements. — Old  male  (type)  :  Basal  length,  311;  occip- 
ito-nasal  length,  310;  palatal  length,  171;  zygomatic  breadth,  216; 
interorbital  breadth,  82. 

URSUS  ORGILOS  Merriam. 

Glacier  Bay  Grizzly. 

Ursus  orgilos  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  176.  August  13, 
1914. 

Type  locality. — Bartlett  Bay,  east  side  of  Glacier  Bay,  Southeast- 
ern Alaska. 

Type  specimen. — No.  180280,  probably  $  ,  rather  old,  U.  S.  Na- 
tional Museum,  Biological  Survey  collection.  Collected  August  22, 
1912,  by  A.  Hasselborg. 

Cranial  characters. — Size  medium ;  skull  long,  rather  narrow,  low, 
flat  on  top,  slightly  dished.  Frontal  shield  rather  narrow,  flat,  concave 
between  orbits,  acute  pointed  posteriorly ;  postorbital  processes  long, 
slender,  outstanding,  slightly  decurved  and  recurved  (posteriorly) ; 


46  NORTH    AMERICAN    FAUNA.  [No.  41. 

rostrum  normal  or  rather  small ;  nasals  nearly  horizontal,  slightly  up- 
lifted and  decurved  anteriorly;  braincase  long  and  low;  squamosal 
shelves  long ;  zygomata  moderately  spreading  and  strongly  outbowed, 
the  broadest  part  more  anterior  than  usual;  sagittal  crest  straight 
and  nearly  horizontal,  reaching  only  to  fronto-parietal  suture;  palate 
long  and  rather  narrow  ;  postpalatal  shelf  long,  flat,  and  rather  broad 
for  size  of  skull ;  jugal  broad  anteriorly,  rising  well  above  lachrymal 
duct;  lachrymal  duct  opening  within  orbit;  occipito-sphenoid  long 
(90  mm.)  ;  underjaw  long;  coronoid  blade  broad  at  base,  moderately 
high,  the  apex  moderately  recurved.  Teeth  of  medium  or  rather 
small  size. 

Cranial  comparisons. — Ursus  orgilos  needs  to  be  distinguished 
from  its  neighbors,  orgiloides,  iahltanicus,  and  pallasi.  Compared 
with  orgiloides,  with  which  it  agrees  essentially  in  size  of  skull  and 
length  of  braincase :  Shield  narrower,  gently  sloping  instead  of  rising 
strongly  from  rostrum;  rostrum  longer  and  narrower;  palate  nar- 
rower. Compared  with  tahltanicus:  Skull  and  shield  narrower; 
postorbitals  more  slender  and  delicate;  zygomata  much  less  out- 
standing; underjaw  more  slender  and  delicate.  Compared  with  pal- 
lasi: Skull  as  a  whole,  braincase,  and  posterior  part  of  frontal  shield 
much  longer;  shield  less  elevated  above  rostrum;  sagittal  crest 
shorter. 

Skidl  measurements — Probably  old  male  (type)  :  Basal  length, 
316;  occipito-nasal  length,  304;  palatal  length,  177;  zygomatic 
breadth,  207 ;  interorbital  breadth,  75. 

URSUS  ORGILOIDES  sp.  nov. 
Alsek  Gbizzly. 

Type  No.  223275,  probably  $  ,  U.  S.  National  Museum,  Biological 
Survey  collection.  From  Italio  River,  Alaska.  Collected  November, 
1916.    Purchased  from  E.  M.  Axelson,  of  Yakutat. 

Range. — Coast  strip  southeast  of  Yakutat  Bay.  Specimens  have 
been  received  from  near  Yakutat  village  and  from  Ankow  and  Ank- 
lin  Rivers  and  mouths  of  Alsek  and  Italio  Rivers. 

Cranial  characters. — Skull  of  medium  size,  long,  low,  and  smoothly 
rounded,  rather  narrow,  with  long  braincase,  long-pointed  shield, 
rather  short  rostrum,  and  large  broadly  rounded  lambdoid  crest; 
shield  medium,  rising  strongly  from  horizontal  rostrum,  sulcate  inter- 
orbitally  and  swollen  between  sulcus  and  orbits;  point  ending  at  or 
near  fronto-parietal  suture;  postorbital  processes  moderate, somewhat 
decurved;  sagittal  crest  rather  short;  occipital  overhang  and  inion 
well  developed;  rostrum  short,  broad  for  size  of  skull;  nares  trun- 
cate; zygomata  not  widely  outstanding  and  not  bowed:  palate  long, 


1918.]  PLANICEPS   GROUP.  47 

of  moderate  breadth;  mastoids  appressed;  meatus  tube  large  and 
free;  underjaw  long;  coronoid  blade  moderate;  apex  only  slightly 
recurved.    Teeth  rather  small. 

Cranial  comparisons. — Adult  male  (type)  compared  with  the  type 
of  orgilos:  Size,  length  of  braincase,  and  length  of  postorbital  part 
of  frontal  shield  essentially  same;  shield  broader,  rising  strongly 
from  rostrum  instead  of  sloping  gradually  into  rostrum,  sulcate 
anteriorly  and  swollen  between  sulcus  and  orbits;  rostrum  shorter 
and  broader;  lachrymal  duct  opening  on  orbital  rim  instead  of 
within  rim ;  zygomatic  arches  more  subtriangular  (less  bowed)  ; 
palate  broader;  meatus  tube  much  larger;  lambdoid  crest  more 
highly  developed  and  more  broadly  rounded.  Teeth  about  same 
size;  PM¥  distinctly  of  grizzly  type  (in  orgilos  subcorneal). 

Compared  with  the  type  of  pallasi,  which  it  strongly  resembles: 
Length  about  an  inch  greater  (mainly  in  posterior  part  of  skull)  ; 
braincase  and  point  of  shield  much  longer;  sagittal  crest  much 
shorter;  occipito-sphenoid  and  palate  longer;  mastoids  appressed 
instead  of  divergent;  underjaw  much  longer,  but  inferior  border 
of  ramus  of  essentialty  same  length;  subangular  border  slightly 
longer. 

Remarks. — Unfortunately  there  is  possible  doubt  as  to  the  sex  of 
the  type  specimens  of  orgilos  and  orgiloides,  though  both  are  believed 
to  be  males.  Both  skulls  have  the  appearance  of  males,  except  that 
in  orgiloides  the  point  of  shield  is  longer  than  usual  in  males  and 
the  canines  are  small  for  males  of  corresponding  size. 

Skull  measurements. — Adult  male  (type):  Basal  length,  316.5; 
occipito-nasal  length,  300;  palatal  length,  169;  zygomatic  breadth, 
218 ;  interorbital  breadth,  81.5. 

URSUS  PALLASI  Mebriam. 

Pallas  Gbizzly. 

Ursus  paZlasi  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  149-150,  Sep- 
tember 6,  1916. 

Type  locality. — Donjek  River,  southwestern  Yukon. 

Type  specimen. — No.  205160,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.    Collected  August,  1913,  by  T.  A.  Dixon. 

Range. — Southwest  corner  of  Yukon  Territory,  east  of  the  St. 
Elias  Range  (Kluane  Lake,  Donjek  River,  St.  Clair  River)  and 
adjacent  eastern  border  of  Alaska;  easterly  to  McConnell  River 
and  Teslin  Lake  and  south  into  northern  British  Columbia. 

Cranial  characters. — Old  male  (type)  :  Size  small,  one  of  the 
smallest  of  the  grizzlies;  skull  moderately  elevated,  flattish  on  top, 
with  relatively  broad  frontal  shield  rising  strongly  at  orbits.  Frontal 
64854°— 18 4 


48  NORTH   AMERICAN   FAUNA.  [No.  41. 

shield  flattish,  exceedingly  short  pointed  posteriorly,  faintly  de- 
pressed medially  between  orbits,  slightly  swollen  on  sides  of  median 
depression,  strongly  sloping  to  rostrum;  postorbital  processes  small, 
peglike,  horizontally  outstanding;  fronto-nasal  region  strongly 
dished;  rostrum  short,  somewhat  depressed  and  pugged;  nasals  ris- 
ing anteriorly;  nares  small  and  subtruncate;  sagittal  crest  long, 
reaching  to  halfway  between  fronto-parietal  suture  and  plane  of  post- 
orbitals; zygomatic  arches  moderately  outstanding,  narrow,  and 
slender,  not  expanded  vertically;  palate  and  postpalatal  shelf  short 
and  broad;  mastoids  long  and  spreading.  Underjaw  long  for  size  of 
skull;  coronoid  narrow  above,  the  apex  not  reaching  plane  of  con- 
dyle ;  teeth  rather  large  for  size  of  skull. 

Old  female  (No.  205162)  from  St.  Clair  River,  Yukon;  collected 
September  6,  1914,  by  A.  Hoyt:  Size  very  small;  frontal  shield 
remarkably  broad  for  so  small  a  skull,  convex  and  medially  sulcate 
interorbitally ;  fronto-nasal  region  strongly  dished;  postorbitals 
small,  outstanding;  rostrum  short  and  depressed;  palate  and  post- 
palatal  shelf  short  and  broad ;  zygomata  rather  strongly  outstanding, 
subtriangular.  Underjaw  very  small  and  light;  coronoid  moderate, 
apex  strongly  recurved;  teeth  very  small,  nearly  as  small  as  in 
nelsoni. 

Cranial  comparisons. — Ursus  palTasi,  owing  to  the  number  of 
species  occurring  in  or  adjacent  to  its  range,  should  be  known  from 
tahltanicus,  orgilos,  orgiloides,  pulchellus,  and  kluane.  It  may  be 
related  to  tahltanicus  but  is  easily  distinguished  by  its  low  broad 
rostrum,  strongly  dished  fronto-nasal  region,  very  short  shield,  less 
elevated  midfrontal  region,  long  and  rather  high  sagittal  crest,  and 
larger  teeth;  while  tahltanicus  has  a  higher  and  narrower  rostrum, 
sloping  fronto-nasal  region,  much  longer  shield,  higher  midfrontal 
region,  shorter  and  lower  sagittal  crest,  and  smaller  teeth.  From 
orgilos  and  orgiloid.es  it  may  be  told  by  the  shortness  of  the  skull  as 
a  whole,  shortness  of  braincase  and  point  of  frontal  shield,  and 
greater  length  of  sagittal  crest.  From  pulchellus  it  differs  in  lower 
vault  of  cranium,  much  broader  shield  and  rostrum,  shorter  rostrum, 
broader  palate  and  postpalatal  shelf,  longer  underjaw,  and  smaller 
molars.  From  kluane  it  differs  strikingly  in  much  smaller  size, 
much  lower  arch  of  cranium,  conspicuously  shorter  braincase  and 
sagittal  crest;  very  much  smaller,  narrower,  and  less  strongly  de- 
curved  postorbitals ;  very  much  less  elevated  and  more  dished  fronto- 
nasal region;  much  lower  rostrum;  much  shorter  palate  and  much 
shorter  underjaw. 

Adult  female  compared  with  adult  female  kluane:  Size  of  skull 
essentially  same  or  slightly  smaller;  canines  smaller;  molars  very 
much  smaller. 


1918.]  PLAN7CEPS   GROUP.  49 

Skull  measurements. — Old  male  (type)  :  Basal  length,  302.5;  occi- 
pito-nasal  length,  279;  palatal  length,  159;  zygomatic  breadth,  209; 
interorbital  breadth,  72.5. 

URSUS  RUNGIUSI  RUNGIUSI  *  sp.  nov. 
Rungius  Geizzlt. 

Type  No.  179893,  $  young-adult,  U.  S.  National  Museum,  Biolog- 
ical Survey  collection.  Collected  September,  1910,  in  Rocky  Moun- 
tains on  headwaters  of  Athabaska  River,  Alberta,  by  Carl  Rungius, 
and  by  him  presented  to  the  Biological  Survey. 

Characters. — Young -adult  male  (type)  :  Size  small;  skull  low  and 
flat,  with  low  depressed  braincase,  very  broad  depressed  sinus  case, 
medium  or  narrow  and  exceptionally  short  frontal  shield,  long  sagittal 
crest,  and  long,  peglike  outstanding'postorbital  processes.  Fully  adult 
males  differ  somewhat.  An  old  male  from  Fortress  Lake,  head  of 
Athabaska  River  (No.  40091,  Amer.  Mus.  Nat.  Hist.),  killed  in  May, 
1916,  by  Malcolm  S.  Mackay;  another  old  male  (No.  1919,  Ottawa 
Museum)  from  Kootenay  Pass,  Alberta;  and  a  fully  adult  male  (No. 
209899)  from  Indian  Point  Creek,  near  Barkerville,  B.  C,  present 
the  following  characters:  Skull  small,  low,  and  nearly  flat;  shield 
low,  of  medium  breadth,  broadly  flat-concave  between  orbits,  sloping 
gradually  into  rostrum,  very  short  pointed  posteriorly;  postorbitals 
peglike,  outstanding,  and  elevated;  orbital  rims  slightly  thickened 
and  everted;  rostrum  small,  narrow,  and  rather  high  for  so  small 
a  skull;  zygomata  widely  outbowed;  palate  and  postpalatal  shelf 
narrow;  base  of  cranium  rather  narrow;  mastoids  appressed  (not 
outstanding);  underjaw  rather  light;  apex  of  coronoid  recurved. 
Dentition  moderate.  PM¥  distinctly  of  grizzly  type.  Canines  and 
upper  molars    rather  large  for  so  small  a  skull. 

Cranial  comparisons. — Adult  male  (No.  209899),  from  Indian 
Point  Creek,  B.  C,  compared  with  adult  male  macfarlani  (type)  : 
Size  slightly  less;  skull  as  a  whole  much  lighter;  rostrum  much 
smaller  and  less  elevated;  fronto-nasal  region  slightly  dished; 
zygomata  much  less  widely  outbowed;  palate  and  postpalatal  shelf 
much  narrower;  sagittal  crest  shorter;  base  of  skull  narrower; 
underjaw  much  less  massive;  apex  of  coronoid  more  recurved. 
Dentition  lighter  (except  M-2-,  which  is  of  about  same  size  in  both) ; 
MT  less  swollen;  PM¥  distinctly  of  grizzly  type  (not  conical  as  in 
macfarlani) . 

Adult  male  (No.  209899,  from  Indian  Point  Creek,  B.  C.)  compared 
with  adult  male  ophrus  (type)  :  Size  smaller;  vault  of  cranium  very 

1  Named  for  the  artist,  Carl  Rungius,  of  New  York,  who  collected  and  presented  the 
type  specimen. 


50  NORTH   AMERICAN   FAUNA.  [No.  41. 

much  lower  and  flatter;  frontal  shield  broadly  concave,  not  deeply 
sulcate  medially ;  postorbital  processes  more  slender  and  horizontal ; 
orbital  rims  slightly  or  not  swollen ;  f ronto-nasal  region  not  notably 
dished ;  rostrum  lower ;  nares  smaller ;  sagittal  crest  less  highly 
developed  and  straighter  (less  convex) ;  zygomata  very  much  less 
outbowed  and  only  slightly  arched  (in  ophrus  very  highly  arched) 
mastoids  much  shorter;  under  jaw  much  shorter;  coronoid  about  the 
same  size ;  upper  canines  and  upper  and  lower  molars  about  the  same 
size;  lower  canines  smaller. 

Adult  and  old  males  compared  with  old  male  hylodrormis  (No. 
205170),  from  Selkirk  Mountains,  B.  C:  Skull  shorter;  shield  flat- 
concave,  rising  at  orbits  and  postorbitals  (in  hylodromus  decurved 
laterally)  ;  posterior  part  of  shield  very  much  shorter;  vault  of 
cranium  much  lower;  braincase  more  depressed;  rostrum  smaller, 
narrower,  more  nearly  horizontal;  zygomata  more  outbowed  (less 
triangular)  ;  sagittal  crest  longer;  postpalatal  shelf  narrower. 

Skull  measurements. — Young-adult  male  (type)  :  Basal  length, 
293;  occipito-nasal  length,  282;  palatal  length,  162;  zygomatic 
breadth,  190;  interorbital  breadth,  73.  Adult  male  (No.  209899) 
from  Indian  Point  Creek,  B.  C. :  Basal  length,  294  ;x  occipito-nasal 
length,  278.5;  palatal  length,  161;  zygomatic  breadth,  198;  inter- 
orbital breadth,  75.  Old  male  (No.  40091  Amer.  Mus.  Nat.  Hist.) 
from  head  Athabaska  River,  B.  C. :  Basal  length,  295 ;  occipito-nasal 
length,  276;  palatal  length,  166;  zygomatic  breadth,  214;  inter- 
orbital breadth,  75. 

URSUS  RUNGIUSI  SAGITTALIS  subsp.  nov. 
Crested  Gbizzly. 

Type  No.  210705,  $  ad.  (rather  old),  U.  S.  National  Museum, 
Biological  Survey  collection.  From  Champagne  Landing,  south- 
western Yukon.  Collected  in  the  fall  of  1915.  Purchased  from 
Mackay  &  Dippie. 

Cranial  characters. — Adult  male  (type)  :  Size  small;  skull  low  and 
narrow;  shield  flat  or  flat-concave,  narrow,  short  pointed,  sloping 
gradually  into  rostrum;  postorbitals  slender,  outstanding;  rostrum 
slender ;  nares  large  and  oblique ;  zygomata  moderately  outstanding, 
subtriangular ;  sagittal  crest  long,  high,  and  arcuate ;  palate  narrow ; 
underjaw  long  and  slender;  coronoid  small  and  falcate;  subangular 
border  short.    Dentition  moderate. 

Cranial  comparisons. — Adult  male  (type)  compared  with  adult 
male  rungiusi  (No.  209899)  :  Skull  similar  in  general  but  shield  nar- 
rower and  flatter;  postorbitals  more  slender;  orbital  rims  not  thick- 
ened or  everted;  nares  larger  and  more  oblique;  sagittal  crest  very 

1  Restored. 


1918.]  PLANICEPS   GROUP.  51 

much  higher,  rising  well  above  point  of  shield;  squamoso-j ugal 
suture  much  shorter;  unclerjaw  longer;  coronoid  smaller  and  more 
falcate;  subangular  border  shorter.  Canines  slightly  larger;  M2 
somewhat  smaller. 

Adult  male  (type)  compared  with  orgilos  (type)  :  Skull  about  an 
inch  shorter;  braincase  very  much  shorter:  point  of  shield  much 
shorter;  sagittal  crest  longer,  much  higher,  and  arcuate  instead  of 
straight;  rostrum  much  smaller;  squamosal  arm  of  zygoma  much 
shorter;  underjaw  and  ramus  much  shorter;  subangular  border  nearly 
the  same ;  coronoid  blade  narrower. 

Skull  measurements. — Adult  male,  old :  Basal  length,  295 ; *  oc- 
cipito-nasal  length,  281 ;  palatal  length,  160 ;  zygomatic  breadth,  200 ; 
interorbital  breadth,  71.5. 

URSUS  MACFARLANI  *  sp.  nov. 
MacFabiane  Bear. 

Type  No.  6551,  $  ad.,  U.  S.  National  Museum.  Collected  on 
Anderson  River,  50  miles  below  Fort  Anderson,  Mackenzie,  May 
8,  1863,  by  R.  MacFarlane.     (Original  No.  551.) 

Characters. — External  characters  unknown.  Relationship  ap- 
parently with  rungiusi. 

Cranial  characters. — Size  medium;  skull  of  adult  male  (type) 
massive,  low,  broad,  flat ;  tip  of  nose  to  point  of  shield  in  same  plane 
without  trace  of  dishing.  Frontal  shield  low,  flat-concave,  rather 
broad,  exceptionally  short  posteriorly,  shallowly  concave  between 
orbits,  not  rising  above  plane  of  rostrum;  postorbitals  large,  hori- 
zontally outstanding  and  slightly  elevated;  rostrum  broad  and  high, 
large  for  size  of  skull;  zygomata  widely  outbowed;  sagittal  crest 
long  and  low;  palate  and  postpalatal  shelf  broad;  underjaw  massive, 
coronoid  blade  high,  narrow,  nearly  vertical,  obtusely  rounded  above ; 
subangular  border  exceptionally  short.  Dentition  heavy:  Canines 
and  molars  large  for  size  of  skull;  PMT  subcorneal;  MT  swollen; 
M-2-  moderate,  the  heel  slightly  emarginate  and  rather  broadly 
rounded  posteriorly.  In  immature  and  young-adult  males  the  frontal 
shield  is  less  flat,  the  sides  (between  sulcus  and  orbits)  strongly 
swollen  (as  shown  by  No.  7146,  from  Franklin  Bay;  and  2773, 
Ottawa  Museum,  from  Stapylton  Bay). 

Cranial  comparisons. — Adult  male  (type)  compared  with  adult 
male  rungiusi  (No.  209899)  from  Indian  Point  Creek  near  Barker- 
ville,  B.  C. :  Skull  slightly  larger  and  much  more  massive,  with  higher 

1  Partly  restored. 

2  Named  in  honor  of  Roderick  MacFarlane,  who  collected  the  specimen  and  presented  it 
to  the  Smithsonian  Institution. 


52  NORTH   AMERICAN   FAUNA.  [No.  41. 

and  much  broader  rostrum,  and  much  more  widely  outbowed  zygo- 
mata; sagittal  crest  longer;  palate,  postpalatal  shelf,  and  base  of 
skull  much  broader;  underjaw  much  more  massive;  coronoid  blade 
more  nearly  vertical  and  less  recurved  at  apex.  Dentition  heavier: 
Canines  and  molars,  upper  and  lower,  larger;  MT  more  swollen;  PM? 
subconical  (not  at  all  of  grizzly  type). 

Skull  measurements. — Adult  male  (type):  Basal  length,  303; 
occipito-nasal  length,  283;  palatal  length,  164;  zygomatic  breadth, 
218 ;  interorbital  breadth,  79. 

URSUS  CANADENSIS  Meebiam.1 
Canada  Gkizzly. 

Vrsus  shoshone  canadensis  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp. 
184-185,  August  13,  1914. 

Type  locality. — Moose  Pass,  near  Mount  Robson,  British  Columbia. 

Type  specimen. — No.  174511,  $  ad.,  U.  S.  National  Museum.  Col- 
lected by  N.  Hollister  and  Charles  D.  Walcott,  jr.,  July  23,  1911. 
(Original  No.  3792,  Hollister  catalogue.) 

Range. — Eastern  British  Columbia;  limits  unknown  (type  from 
near  Mount  Robson ;  and  an  adult  female  from  Kootenay  Lake ) . 

Characters. — Size  medium;  color  brown,  grizzled  with  buff;  claws 
short  for  a  grizzly,  rather  thick,  moderately  curved,  pale  yellowish 
on  upper  surface  and  tips,  brownish  horn  color  on  sides. 

Color. — Muzzle  very  pale  drab  brown,  changing  to  darker  brown 
on  head,  face,  and  chin,  darkest  around  ears;  top  of  head,  cheeks 
posteriorly,2  ears,  back,  and  thighs  washed  with  buffy  whitish  from 
abundance  of  buffy-tipped  hairs;  foreleg  and  lower  part  of  hind  leg 
and  feet  very  dark  (almost  blackish  brown)  ;  long  hairs  of  throat 
and  axillary  region  pale  yellowish,  of  rest  of  underparts  dark 
brown. 

Cranial  characters. — Adult  male  (type)  :  Skull  of  medium  size, 
rather  long,  low,  and  narrow,  flat  in  frontal  region,  with  long,  high 
sagittal  crest  rising  above  general  level  of  top  of  cranium.  Frontal 
shield  small,  flat,  narrow,  faintly  depressed  medially,  short  pointed 
posteriorly,  gently  sloping ;  postorbitals  slender,  peglike,  horizontally 
outstanding;  rostrum  relatively  high,  tapering  anteriorly;  sagittal 
crest  remarkably  long,  arcuate,  and  high,  reaching  nearly  to  midway 
between  fronto-parietal  suture  and  plane  of  postorbitals;  zygomata 
moderate,  slightly  outbowed,  and  only  slightly  expanded  vertically; 
palate  excavated  between  molar  series;  postpalatal  shelf  rather  long 
and  broad;  notch  medium  or  narrow.     Underjaw  rather  massive; 

1  Tentatively  included  in  planiceps  group.     (See  Introduction,  pp.  12—13.) 

2  The  old  whitish-tipped  hairs  of  the  old  coat  have  fallen  out  on  the  cheeks  and  anterior 
part  of  head  nearly  to  ears. 


1918.]  ARIZONA   GROUP.  53 

ramus  moderately  bellied  posteriorly;  coronoid  blade  broad  and 
rather  short,  the  apex  cutting  plane  of  condyle.  Dentition  rather 
heavy;  canines  thick  and  short;  molars  broad. 

Adult  female  (No.  209902  from  Kootenay  Lake,  British  Colum- 
bia) :  Skull  rather  small,  long,  narrow,  low,  and  slightly  dished. 
Frontal  shield  narrow,  flattish,  slightly  depressed  medially  between 
orbits,  long  pointed,  the  point  reaching  to  midparietal  region;  post- 
orbitals  moderate;  rostrum  long  and  slender,  tapering;  zygomata 
moderately  spreading.  Under  jaw  similar  to  that  of  male  but  much 
smaller;  coronoid  blade  relatively  narrower  and  higher.  Dentition 
heavy;  teeth  similar  to  those  of  male  and  only  slightly  smaller; 
canines  large  and  swollen;  molars  broad. 

Cranial  comparisons. — Adult  male  (type)  compared  with  adult 
male  macrodon  (type)  :  Size  essentially  same;  shield  more  strongly 
sloping ;  postorbitals  more  slender ;  rostrum  lower  and  more  tapering ; 
sagittal  crest  higher ;  inion  much  more  strongly  produced ;  subangular 
border  of  ramus  shorter ;  angular  process  not  bellied ;  canines  smaller ; 
M^  and  M-  decidedly  smaller. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  313;  oc- 
cipito-nasal  length,  312 ;  palatal  length,  171 ;  zygomatic  breadth,  208 ; 
interorbital  breadth,  74. 

Arizonae  Group. 

URSUS  ARIZONA  Meebiam. 

Aeizona  Geizzly. 

(PI.  XVI.) 

Vrsus   arizonw  Merriam,   Proc.   Biol.    Soc.   Washington,   XXIX,    pp.    135-136, 
September  6,  1916. 

Type  locality. — Escudilla  Mountains,  Apache  County,  Arizona. 

Type  specimen. — No.  177332,  $  ad.,  U.  S.  National  Museum, 
Biological  Survey  collection.  Collected  September  3,  1911,  by  C.  H. 
Shinn. 

Cranial  characters. — Size  rather  large;  skull  as  a  whole  rather 
long  and  narrow,  with  broad  rostrum;  vault  of  cranium  moderately 
elevated  but  not  arched,  highest  about  two-thirds  distance  from  plane 
of  postorbitals  to  fronto-parietal  suture;  frontal  shield  rather  nar- 
row, nearly  flat,  gently  sloping  in  plane  of  rostrum,  the  posterior 
point  in  type  specimen  reaching  to  about  25  mm.  in  front  of  parietals 
(in  older  specimens  shorter)  ;  postorbitals  broad  and  broadly 
rounded,  nearly  horizontal,  but  not  widely  projecting;  fronto-nasal 
region  and  rostrum  elevated  and  swollen,  continuing  plane  of  frontal 
shield  without  trace  of  dishing,  tapering  anteriorly;  zygomata  not 


54  NORTH   AMERICAN    FAUNA.  [No.  41. 

widely  outstanding,  bowed,  anterior  roots  swollen ;  palate  rather  short 
and  broad;  postpalatal  shelf  broad;  meatus  tube  long;  coronoid 
blade  rather  broad  above,  its  recurved  apex  cutting  plane  of  con- 
dyles. Teeth  rather  small  for  size  of  skull;  canines  of  good  size; 
molars  rather  small  for  size  of  skull,  especially  last  upper  molar. 

Cranial  comparisons. — Adult  male  (type)  compared  with  old  male 
apache  (type)  :  Basilar  length,  occipito-nasal  length,  length  of  palate, 
interorbital  breadth,  and  occipito-sphenoid  length  essentially  the 
same;  zygomata  very  much  less  outstanding  and  subtriangular  in- 
stead of  bowed ;  frontal  shield  flatter,  of  essentially  the  same  breadth 
interorbitally,  but  very  much  narrower  across  postorbital  processes; 
postorbital  processes  much  less  broadly  outstanding;  orbital  rims 
less  swollen;  fronto-nasal  regu^/much  more  elevated  and  swollen; 
rostrum  much  larger,  broader,  more  swollen,  and  tapering  instead 
of  depressed  basally,  narrow,  and  horizontal ;  palate  and  postpalatal 
shelf  much  broader.  Under  jaw  weaker;  ramus  less  broad  vertically; 
coronoid  blade  less  high;  molars  slightly  larger;  heel  of  M-2-  longer, 
more  distinctly  emarginate  on  outer  side  (less  tapering). 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  326;  oc- 
cipito-nasal length,  323;  palatal  length,  175;  zygomatic  breadth,  208; 
interorbital  breadth,  82. 

URSUS  IDAHOENSIS  sp.  nov. 
Idaho  Grizzly. 

Type  No.  93,  $  old,  Merriam  collection  (=187888  U.  S.  National 
Museum),  from  North  Fork  Teton  River,  eastern  Idaho,  September 
23, 1874.  Killed  by  the  late  Richard  Leigh  (better  known  as  "  Beaver 
Dick"). 

Characters. — Size  rather  large,  about  equaling  arizonw  (much 
smaller  than  horribilis  and  subspecies,  and  shorter  than  rogersi, 
bisonophagus,  and  perturbans)  ;  frontal  shield  convex  both  longi- 
tudinally and  transversely;  fronto-nasal  region  elevated  and  some- 
what compressed  (much  as  in  rogersi,  but  rostrum  much  shorter)  ; 
postorbital  processes  rather  weak  and  decurved  as  in  rogersi  and 
arizonce;  sagittal  crest  rather  short  and  not  strongly  developed; 
zygomata  moderately  outbowed ;  coronoid  blade  rather  high,  but  less 
high  than  in  bisonophagus  and  rogersi;  ramus  flattish,  broad  verti- 
cally, but  much  less  broad  and  massive  then  in  rogersi;  dentition 
rather  heavy;  M1  large,  its  heel  elongate,  emarginate,  but  not  nar- 
rowed posteriorly,  slightly  everted ;  canines  rather  small,  about  as  in 
rogersi  and  arizonce. 

An  old  female  (No.  160153)  from  Wallowa  Mountains,  Oregon,  is 
assumed  to  be  typical  of  idahoensis.  It  has  a  long,  low,  slender, 
smoothly  rounded  skull  with  narrow  zygomatic  arches,  narrow 
palate,  rather  broad  postpalatal  shelf,  and  very  small  teeth. 


1918.]  ARIZONA   GROUP.  55 

Skull  measurements. — Old  male  (type)  :  Basal  length,  317;  oc- 
cipito-nasal  length,  318;  palatal  length,  177;  zygomatic  breadth, 
206;  interorbital  breadth,  81. 

URSUS  PULCHELLUS  PULCHELLUS  sp.  NOV. 
Upper  Yukon  Gkizzly. 

Type  No.  221599,  $  ad.,  U.  S.  National  Museum,  Biological  Sur- 
vey collection,  from  Ross  River,  Yukon  Territory,  Canada.  Col- 
lected July  20,  1916,  by  Fred  E.  Enevoldsen. 

Characters. — Size  small;  frontal  shield  and  rostrum  narrow; 
vault  of  cranium  well  arched;  base  of  cranium  moderately  arched; 
shield  rising  rather  strongly  from  plane  of  rostrum,  convex  trans- 
versely but  shallowly  sulcate  medially,  short  pointed  posteriorly 
(point  ending  about  halfway  between  plane  of  postorbitals  and 
fronto-parietal  suture);  rostrum  slender  and  high;  fronto-nasal 
region  dished;  nasals  nearly  horizontal;  sagittal  crest  only  slightly 
developed,  not  high  posteriorly,  somewhat  decurved;  occiput  rather 
low  and  shortly  truncate;  palate  and  postpalatal  shelf  rather  nar- 
row; zygomata  subtriangular ;  squamosal  root  expanded  vertically; 
occipito- sphenoid  short  (80  mm.) ;  under  jaw  short,  its  inferior  bor- 
der straight;  coronoid  rather  low,  broad  basally,  apex  strongly  re- 
curved; subangular  border  short.  Teeth,  particularly  molars,  large 
for  so  small  a  skull;  last  upper  molar  broadly  quadrate  anteriorly, 
the  heel  abruptly  and  strongly  emarginate,  narrowly  rounded  pos- 
teriorly ;  M1  large,  broad,  and  massive. 

Cranial  comparisons. — Ursus  pulchellus  requires  comparison  with 
its  near  relative  ereunetes,  and  also  with  pallasi  and  kluane.  Adult 
male  (type)  compared  with  adult  male  ereunetes  (type)  :  Size 
slightly  smaller;  fronto-nasal  region  more  dished;  shield  less  flat; 
zygomata  broad  (in  ereunetes  slender);  occipito-sphenoid  shorter; 
M1  larger  and  more  massive. 

Compared  with  old  male  pallasi  (type) :  Basal  length  slightly  less; 
occipito-nasal  length  same;  zygomatic  breadth  less;  cranium  higher 
and  more  arched;  frontal  shield  and  rostrum  conspicuously  nar- 
rower; palate  and  postpalatal  shelf  narrower;  rostrum  longer, 
higher,  and  more  slender;  squamosal  root  of  zygoma  more  broadly 
expanded  vertically;  mastoids  less  elongate;  underjaw  shorter  and 
less  massive.    Upper  molars  larger  and  more  massive. 

Compared  with  adult  male  kluane :  Similar  in  general  appearance 
but  skull  as  a  whole,  braincase,  palate,  and  underjaw  very  much 
shorter,'  frontal  shield  very  rmich  narrower;  inferior  border  of  ramus 
more  abruptly  upcurved;  subangular  border  more  nearly  horizontal 
and  much  more  sharply  defined. 


56  NORTH   AMERICAN   FAUNA.  [No.  41. 

Remarks. — The  type  of  pulchellus  came  from  Ross  River,  a  north- 
ern tributary  of  the  Pelly.  Skulls  of  males  from  the  southwestern 
corner  of  Yukon  (Donjek  River  and  Champagne  Landing)  differ  in 
greater  occipito-nasal  length,  more  highly  arched  cranium,  more  ele- 
vated rostrum,  and  less  deeply  emarginate  heel  of  1VP-. 

I  refer  to  pulchellus  an  adult  female,  No.  204187,  from  McConnell 
River,  Yukon,  and  a  still  older  female,  No.  215113,  from  Ross  Moun- 
tains. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  292;  oc- 
cipito-nasal length,  281 ;  palatal  length,  160 ;  zygomatic  breadth,  196 ; 
interorbital  breadth,  66. 

URSUS  PULCHELLUS/EREUNBTES  subsp.  nov. 

KOOTENAY   GRIZZLT. 

Type  No.  222323,  $  ad.,  from  Beaverfoot  Range,  Kootenay  Dis- 
trict, British  Columbia.  Collected  October  1,  1916,  by  George  Hill, 
of  Field,  British  Columbia. 

Characters. — Size  rather  small;  occiput  shortly  truncate;  vault  of 
cranium  and  basicranial  axis  well  arched;  fronto-nasal  region  ele- 
vated, sloping  gradually  upward,  not  dished;  rostrum  small,  rather 
narrow,  rising  gradually  into  shield;  point  of  shield  ending  about 
two-thirds  distance  from  plane  of  postorbitals  to  fronto-parietal 
suture;  zygomata  slender,  rather  broadly  outbowed  for  size  of  skull. 
Last  upper  molar  broad  anteriorly,  the  anterior  part  of  cingulum 
on  inner  side  produced,  the  heel  emarginate,  narrowing  posteriorly, 
subtriangular ;  middle  lower  molar  large  and  massive. 

Skull  in  general  similar  to  that  of  pulchellus  but  slightly  larger, 
with  more  elevated  fronto-nasal  region,  flatter  shield,  much  more 
slender  zygomata,  and  much  longer  occipito-sphenoid ;  teeth  as  in 
pulchellus  except  that  MMs  smaller,  and  M^  is  less  quadrangular 
anteriorly. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  297; 
occipito-nasal  length,  278;  palatal  length,  165;  zygomatic  breadth, 
203 ;  interorbital  breadth,  72. 

URSUS  ORIBASUS  sp.  NOV. 
Liasd  River  Grizzly. 

Type  No.  223991,  $  ad.  (rather  old),  U.  S.  National  Museum, 
Biological  Survey  collection.  From  Upper  Liard  River,  Yukon, 
near  British  Columbia  boundary.  Killed  by  J.  Thompson  in  the 
spring  of  1916.     (Purchased  from  William  Drury,  of  Whitehorse.) 

Characters. — Adult  male  (type) :  Size  large;  hump  absent  or  in- 


1918.]  ARIZONA   GROUP.  57 

conspicuous;  color  dark;  claws  long  (longest  90  mm.1)  and  unusually 
straight;  top  convex  in  section,  dark  horn  color,  paler  at  tips  and 
along  upper  surface;  skull  long,  narrow,  and  arched,  with  elevated 
straight-sloping  fronto-nasal  region  (much  like  that  of  ereunetes 
but  much  longer  posteriorly).  Frontal  shield,  rostrum,  and  molar 
teeth  narrower  than  in  any  other  member  of  the  shoshone  group. 

Color. — General  ground  color  dark  brown  to  dusky ;  muzzle  dull 
golden  brown,  becoming  much  darker  between  eyes;  a  dark  ring 
around  each  eye;  cheeks  chestnut  brown;  top  of  head,  nape,  and 
shoulders  strongly  washed  with  yellowish  buffy;  back  washed  with 
soiled  buffy;  rump  dark  brownish  dusky;  legs  and  feet  dusky 
blackish. 

Cranial  characters. — Adult  male  (type)  :  Skull  rather  large,  long, 
narrow,  rather  strongly  ;:  relied  both  above  and  below,  with  high 
straight-sloping  (not  dished)  fronto-nasal  region.  Frontal  shield 
narrow,  flat,  gently  sloping,  the  point  reaching  two-thirds  distance 
from  postorbitals  to  parietals ;  postorbitals  rather  small,  horizontally 
outstanding;  rostrum  narrow  and  high,  in  same  plane  with  frontal 
shield;  braincase  and  sagittal  crest  long;  inion  and  occipital  over- 
hang marked;  zygomata  well  outstanding,  strongly  subtriangular, 
squamosal  base  broadly  expanded ;  basicranium  and  palate  arched ; 
palate  and  postpalatal  shelf  narrow ;  occipito-sphenoid  92  mm. ;  mas- 
toids large,  divergent.  Under  jaw  long;  ramus  long  and  flat;  sub- 
angular  border  short  and  broad;  coronoid  blade  narrow.  Canines 
rather  long ;  molars  narrow  and  rather  small ;  MA  small,  narrowly 
triangular,  the  heel  small,  thin,  and  pointed;  PM^  imperfectly  of 
grizzly  type. 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
chelan  and  idahoensis  (both  types)  :  Length,  height,  and  arching 
essentially  same;  shield  narrower,  flatter,  and  more  nearly  hori- 
zontal ;  sinus  case  and  rostrum  narrower ;  top  of  rostrum  higher  and 
more  completely  in  fronto-nasal  plane ;  zygomatic  arches  shorter  and 
more  angular  (less  outbowed),  the  inclosed  space  (temporal  fossa) 
much  smaller.  Under  jaw  shorter  and  weaker;  subangular  border 
shorter;  coronoid  blade  narrower. 

Old  male  (type)  compared  with  adult  male  ereunetes  (type) : 
Similar  in  general  appearance  but  length  much  greater;  shield  flat- 
ter; rostrum  more  elevated;  braincase  much  longer;  sagittal  crest 
much  longer  and  higher;  inion  more  strongly  produced;  squamosal 
base  of  zygoma  broadly  expanded  (in  ereunetes  not  expanded).  Ca- 
nines longer;  molars  smaller  and  very  much  narrower. 

1  Claw  of  second  or  index  finger  longest,  but  second,  third,  and  fourth  practically  sub- 
equal  ;  claw  of  thumb  very  long.  Claw  measurements  of  right  hand  (those  of  left  hand 
more  worn  at  tips)  from  upper  exposed  base  to  tip  :  First,  82  mm. ;  second,  90  ;  third,  88  ; 
fourth,  89  ;  fifth,  68. 


58  NORTH   AMERICAN   FAUNA.  [No.  41. 

Remarks. — Ursus  oribasus  appears  to  be  closely  related  to  pulchel- 
lus  and  ereunetes,  both  of  which  have  decidedly  smaller  skulls  with 
larger  teeth.  It  is  related  also  to  idahoensis  and  chelan,  which  are 
about  the  same  size,  but  have  much  broader  skulls  and  differ  other- 
wise as  already  pointed  out. 

Skull  measurements. — Adult  male  (type) :  Basal  length,  310;  oc- 
cipito-nasal  length,  304 ;  palatal  length,  172 ;  zygomatic  breadth,  215 ; 
interorbital  breadth,  75. 

URSUS  CHELAN  Mereiam. 

Chelan  Grizzly. 

Ursus  chelan  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  136-137,  Sep- 
tember 6,  1916.  / 

Type  locality. — East  slope  Cascade  Mountains,  northern  Chelan 
County,  Washington. 

Type  specimen. — No.  205185,  $  old,  U.  S.  National  Museum, 
Biological  Survey  collection.  Killed  in  Township  30  N,  Range  16 
East,  Willamette  Meridian,  Wenatchee  National  Forest.  Collected 
September  1, 1913,  by  D.  S.  Rice. 

Range. — Cascade  and  Cassiar  Mountains  from  northern  Washing- 
ton to  upper  Stikine  River  and  Dease  Lake,  British  Columbia. 

Cranial  characters. —  (External  characters  unknown).  Skull  of 
medium  or  rather  large  size;  facial  axis  strongly  deflected  from 
basicranial  axis;  vault  of  cranium  well  arched,  highest  over  poste- 
rior frontal  region;  sagittal  crest  long,  high,  arcuate,  rising  ante- 
riorly above  general  level  of  top  of  cranium.  Affinities  apparently 
with  hylodromus  on  the  one  hand,  and  with  shoshone  and  pervagor  on 
the  other.  Frontal  shield  narrow,  flattened,  short  pointed  poste- 
riorly, ending  about  midway  between  fronto-parietal  suture  and 
plane  of  postorbitals,  slightly  sulcate  medially ;  postorbital  processes 
rather  broad,  flat,  outstanding  horizontally  (not  depressed  or  de- 
curved)  ;  fronto-nasal  region  including  posterior  two-thirds  of  nasals 
sloping  strongly,  forming  part  of  long  fronto- facial  plane;  rostrum 
small,  short,  somewhat  depressed,  sloping  anteriorly  to  nares,  grad- 
ually rising  posteriorly  into  frontal  plane;  braincase  long,  arched, 
frontal  part  keeled  into  sagittal  crest;  palate  arched  antero-poste- 
riorly,  slightly  concave;  postpalatal  shelf  broad;  zygomata  broadly 
spreading,  rounded  and  outbowed  posteriorly,  vertically  expanded 
and  strongly  arched ;  mastoids  of  medium  length,  spreading ;  under- 
jaw  massive;  ramus  swollen  on  outer  side  over  roots  of  M-j  and 
MT,  bellied  under  last  molars;  coronoid  blade  high,  its  anterior  bor- 
der rather  strongly  recurved,  the  apex  overarching  high  coronoid 
notch  but  barely  reaching  plane  of  front  of  condyle;  teeth  rather 
small  for  size  of  skull  (so  badly  worn  in  type  specimen  that  propor- 
tions of  canines  can  not  be  determined). 


1918.]  ARIZONA   GROUP.  59 

Cranial  comparisons. — Old  male  chelan  differs  from  all  its  rela- 
tives in  the  degree  of  deflection  of  the  facial  part  of  the  skull,  and  ex- 
ceeds all  except  Kylodromus  in  the  arching  of  the  palate. 

Old  male  (type)  compared  with  pervagor  (type)  :  Basal  length 
slightly  less;  zygomatic  breadth  greater;  frontal  shield  shorter 
pointed  and  flatter;  postorbitals  flatter  and  apparently  broader; 
sagittal  crest  longer,  higher  anteriorly  and  more  convex;  rostrum 
shorter  and  more  strongly  sloping  anteriorly;  nares  more  truncate; 
zygomata  more  strongly  outbowed,  more  arched,  more  expanded 
vertically,  squamosal  arm  longer  (squamoso-jugal  suture  much 
longer)  ;  palate  shorter  and  more  strongly  arched ;  under  jaw 
shorter,  its  inferior  ramus  much  shorter;  coronoid  blade  slightly 
higher  and  more  falcate.  Some  of  these  differences  may  be  due  to 
age,  the  skull  of  the  type  of  chelan  being  very  old,  while  the  type  of 
pervagor  is  only  adult.  However,  it  differs  rather  conspicuously 
from  equally  old  male  pervagor  from  Bridge  River  (No.  4,  Provin- 
cial Museum,  Victoria,  B.  C.)  as  will  be  seen  from  the  following: 
Basal  length  20  mm.  less;  occipito-nasal  length  slightly  greater; 
zygomatic  breadth  less;  facial  part  strongly  deflected  (in  pervagor 
not  deflected)  ;  palate  arched  (in  pervagor  not  arched) ;  frontal 
shield  shallowly  sulcate  medially  (in  pervagor  broadly  concave) ; 
postorbitals  broad,  flat,  horizontally  outstanding  (in  pervagor  long, 
peglike,  uplifted,  and  arched)  ;  braincase  and  sagittal  crest  arched 
(in  pervagor  straight  and  nearly  horizontal) ;  occipital  overhang 
much  greater;  zygomata  less  widely  outbowed. 

Old  male  (type)  compared  with  old  male  Kylodromus  (No.  205170) 
from  Selkirk  Mountains,  British  Columbia :  Size  larger  (basal 
length  only  slightly  greater  but  occipito-sphenoid  and  occipito- 
nasal  lengths  much  greater,  and  skull  as  a  whole  distinctly 
larger)  ;  vault  of  cranium  decidedly  more  highly  arched ;  facial 
angle  more  strongly  deflected  from  basicranial  axis;  zygomata 
much  more  widely  spreading  and  outbowed  and  much  more 
arched;  frontal  shield  rising  less  abruptly  from  rostrum,  more 
evenly  sloping,  rising  higher  posteriorly,  and  much  shorter  pointed  ; 
braincase  and  sagittal  crest  much  longer,  the  crest  higher  and  con- 
vex or  arcuate  anteriorly;  occipital  overhang  greater;  palate  more 
strongly  arched  (antero-posteriorly)  ;  mastoids  longer  and  strongly 
spreading.  Underjaw  longer  and  more  massive,  more  swollen  on 
outer  side  below  middle  and  posterior  molars;  its  inferior  border 
more  bellied  posteriorly ;  coronoid  blade  very  much  higher. 

Old  male  (type)  compared  with  old  male  washake  (type)  :  Size 
slightly  larger  (basal  length  essentially  the  same,  but  upper  part 
of  skull  much  longer)  ;  vault  of  cranium  more  highly  arched;  frontal 
shield  continuing  to  rise  posteriorly  (instead  of  flattened)  and  much 
shorter  pointed ;   rostrum  decidedly  broader ;  postorbital  processes 


60  NORTH  AMERICAN  FAUNA.  [No.  41. 

not  elevated;  lachrymal  duct  within  orbit  (not  cutting  rim  as  seen 
from  front);  braincase  compressed  and  keeled  anteriorly  (in 
washake  depressed) ;  sagittal  crest  much  longer  and  convex  instead 
of  straight;  squamosal  arm  of  zygoma  longer  and  more  broadly  ex- 
panded vertically;  palate  concave  and  arched  antero-posteriorly,  in- 
stead of  flat;  postpalatal  shelf  longer  and  less  broadly  flattened ;  oc- 
cipito-sphenoid  longer ;  mastoids  longer  and  more  spreading.  Molars 
smaller. 

Skull  measurements. — Old  male  (type):  Basal  length,  314;  oc- 
cipito-nasal  length,  323 ;  palatal  length,  170 ;  zygomatic  breadth,  225 ; 
interorbital  breadth,  86. 

URSUS  SHOSHONE  Merriam. 

Shoshone  Grizzly. 

Ursus  shoshone  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  184,  August 
13,  1914. 

Type  locality. — Estes  Park,  Rocky  Mountains  of  northern  Colo- 
rado. 

Type  specimen. — No.  203185,  $  old,  U.  S.  National  Museum, 
Biological  Survey  collection. 

Range. — Mountains  of  Colorado  and  Wyoming. 

Characters. — Size  medium  or  rather  large,  but  much  smaller  than 
horribilis  and  bairdi — skull  about  same  size  as  absarokus,  but  nar- 
rower and  widely  different.    External  characters  unknown. 

Cranial  characters. — Adult  nude  (type)  :  Skull  rather  long  and 
high,  with  flattish,  short-pointed,  long-sloping  frontal  shield  con- 
tinuing plane  of  rostrum  to  highest  point,  about  midway  between 
postorbitals  and  fronto-parietal  suture;  zygomata  moderately 
spreading,  outbowed;  anterior  (frontal)  part  of  braincase  keeling 
into  sagittal  crest ;  sagittal  crest  long  and  high ;  lambdoid  crest  high ; 
postorbital  processes  peglike,  outstanding,  rather  slender;  nasal 
region  slightly  dished  and  sulcate  in  middle  third  (nasals  dipping 
toward  one  another — may  be  individual) ;  rostrum  of  moderate 
breadth,  strongly  ascending  in  plane  of  frontal  shield ;  palate  slightly 
dished  between  posterior  molars ;  postpalatal  shelf  broad ;  postpalatal 
notch  long  and  narrow;  lachrymal  duct  cutting  orbital  rim  but 
mainly  on  inner  side.  Under  jaw  rather  long;  ramus  broad  vertically, 
flattish,  highest  posteriorly;  coronoid  blade  high,  rather  strongly 
sloping,  the  apex  cutting  plane  of  condyle.  Canines  (absent  in  type 
but  present  in  other  males)  slender;  molars  medium;  M-3-  large  with 
long  heel. 

Adult  female  (No.  203761,  from  Fort  Fred  Steele,  Wyoming)  : 
Skull  long,  low,  and  rather  slender;  frontal  shield  narrow,  long, 
lyre  pointed  posteriorly,  flat  interorbitally ;  postorbitals  slender  and 


1918.]  ARIZONA    GROUP.  61 

outstanding;  rostrum  slender;  sagittal  crest  short,  reaching  only 
halfway  from  inion  to  fronto-parietal  suture;  lambdoid  crest  mod- 
erate; palate  dished  between  posterior  molars;  postpalatal  shelf 
broad ;  notch  moderate ;  meatus  tube  short ;  ramus  flat  and  light ;  apex 
of  coronoid  produced  posteriorly,  overhanging  deep  coronoid  notch. 
Teeth  rather  small ;  upper  molars  relatively  large,  M>  with  long  heel, 
cut-turned  posteriorly ;  lower  molars  narrow. 

Cranial  comparisons. — Adult  male  compared  with  adult  male  hor- 
riceus:  Rostrum,  nasals,  and  frontal  shield  more  elevated  and  much 
more  strongly  ascending  posteriorly  (less  flattened  and  less  nearly 
horizontal)  ;  lambdoid  much  more  strongly  developed. 

Adult  male  compared  with  adult  male  absarokus:  Frontal  shield 
flat,  rising  gradually  in  long  continuous  slope  to  highest  point,  about 
25  mm.  anterior  to  fronto-parietal  suture  (in  absarokus  arched, 
strongly  convex,  and  rising  suddenly  to  highest  point,  immediately 
behind  orbits)  ;  postorbital  processes  more  slender;  fronto-nasal  re- 
gion only  slightly  dished ;  rostrum  more  slender ;  braincase  narrower ; 
breadth  across  squamosal  shelves  less. 

Remarks. — Skulls  from  the  Wind  River  and  Absaroka  Mountains 
have  the  last  upper  molar  smaller,  the  heel  less  strongly  developed. 

Skull  measurements. — Old  male  (type):  Basal  length,  320;  occi- 
pito-nasal  length,  317;  palatal  length,  166;  zygomatic  breadth,  208; 
interorbital  breadth,  78. 

URSUS  KENNERLYI1  Mebriam. 

Sonoea  Grizzly. 

Ursus  kennerlyi  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  194,  August 
13,  1914. 

Type  locality. — Mountains  of  northeastern  Sonora,  near  Los 
Nogales,  Mexico. 

Type  specimen.— Skull  No.  2086,  $  old ;  skin  No.  1047,  U.  S.  Na- 
tional Museum.    Collected  in  June,  1855,  by  Dr.  C.  B.  Kennerly. 

Range. — Nothing  is  known  of  the  range  of  kennerlyi  except  that 
the  type  specimen  came  from  mountains  near  Nogales,  Sonora.  Its 
affinities  with  utahensis  suggest  that  formerly  it  may  have  had  a  dis- 
connected distribution  northward  in  the  mountains  of  central 
Arizona. 

Characters. — Size  rather  small.  Ursus  kennerlyi  is  a  strongly 
marked  member  of  the  arizonce-utahensis  group,  most  nearly  related 
to  utahensis  but  very  much  smaller,  although  the  teeth  are  about  same 
size.  The  skull,  though  that  of  an  old  male,  agrees  in  size  (length) 
with  that  of  female  utahensis. 

1  Named  In  honor  of  Dr.  C.  B.  Kennerly,  who  collected  the  type  specimen. 


62  NORTH   AMERICAN   FAUNA.  [No.  41. 

Color. T- The  prevailing  color  is  dull  pale  brownish  yellow  with 
amber  tinge.  The  tips  only  are  of  this  color,  the  basal  and  larger 
portion  being  of  a  dark  chestnut-brown,  passing  into  blackish,  which 
extends  nearly  to  yellowish  tips,  the  blackish  predominating  along 
median  line  of  back  and  posteriorly;  legs  blackish  brown  slightly 
tinged  with  chestnut  (Baird). 

Cranial  characters. — Size  rather  small;  skull  long,  narrow,  and 
high,  but  not  much  arched ;  rostrum  narrow  and  high,  in  same  plane 
with  shield;  fronto-nasal  region  strongly  elevated,  making  a  con- 
vexity slightly  above  otherwise  continuous  plane  of  rostrum  and 
frontal  shield ;  rostrum  and  fronto-nasal  region  subterete,  constricted 
(but  not  strongly  pinched  in)  in  front  of  orbits ;  nares  much  higher 
than  broad;  frontal  shield  flaf,  rather  short  pointed  posteriorly, 
passing  into  sagittal  crest  about  one-third  distance  from  fronto- 
parietal suture  to  postorbitals ;  postorbital  processes  long,  rather 
slender,  outstanding,  and  slightly  decurved;  frontal  part  of  brain- 
case  elevated ;  zygomata  moderately  spreading  and  outbowed ;  palate 
long,  somewhat  concave ;  squamosal  shelves  broad ;  coronoid  broadly 
falcate;  ramus  flat  and  broad  vertically.  Teeth  of  medium  size 
(badly  worn). 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
horriceus  (type) :  Size  essentially  the  same;  vault  of  cranium  higher 
over  posterior  frontal  region;  frontal  shield  slightly  convex  (not 
sulcate  or  depressed  between  orbits  or  elsewhere),  longer  pointed 
posteriorly;  fronto-nasal  region  markedly  elevated  instead  of  de- 
pressed; rostrum  much  higher  posteriorly,  rising  in  same  plane  with 
frontal  shield  and  strongly  compressed;  postorbital  processes  more 
slender  and  less  decurved;  sagittal  crest  much  shorter;  angle  of  jaw 
shorter ;  inferior  border  of  ramus  decidedly  longer ;  apex  of  coronoid 
more  slender.  Teeth  so  badly  worn  that  detailed  characters  are  lost ; 
in  size,  however,  they  agree  essentially  with  those  of  horriceus  except 
that  the  large  lower  premolar  is  decidedly  smaller. 

Skull  measurements. — Old  male  (type):  Basal  length,  314;  * 
occipito-nasal  length,  306;  palatal  length,  165;  zygomatic  breadth, 
205 ;  interorbital  breadth,  75. 

URSUS  UTAHENSIS  Merriam. 

Utah  Grizzly. 

Ursus  utahensis  Merriam,   Proc.  Biol.  Soc.  Washington,  XXVIT,  pp.  193-194, 
August  13,  1914. 

Type  locality.— North  Fork  Salina  Creek,  10  or  12  miles  southeast 
of  Mayfield,  Utah. 

1  Restored. 


1918.]  ARIZONA   GROUP.  63 

Type  specimen— No.  180193,  $  old,  U.  S.  National  Museum, 
Biological  Survey  collection.  Collected  May  22,  1911,  by  Mart 
Martenson. 

Range. — Southern  Wasatch  and  Pine  Valley  Mountains ;  limits  un- 
known. 

Characters. — Size  large;  coloration  apparently  normal.  Skull 
long,  narrow,  and  high,  but  not  arched;  fronto-nasal  region  high 
and  very  narrow — strongly  pinched  in. 

Color. — Skin  of  head  of  male  killed  on  Pine  Valley  Mountain, 
southwest  Utah  (obtained  from  forest  ranger,  September  24,  1907,  by 
Clarence  Birdseye;  original  No.  989)  :  Muzzle  pale  brown;  face  and 
throat,  except  pale  lip  edgings  and  long  hairs  of  median  line  of 
throat,  dark  brown,  becoming  grizzled  posteriorly ;  top  of  head  very 
dark;  grizzled  posteriorly  by  brown-tipped  hairs. 

Cranial  characters. — Adult  male  (type,  and  equally  old  male  from 
northeast  corner  Sevier  National  Forest)  :  Size  large;  skull  very  long, 
high,  and  exceedingly  narrow;  zygomata  moderately  spreading  and 
outbowed;  frontal  shield  narrow,  flattened  posteriorly,  falling  away 
laterally  immediately  in  front  of  orbits,  leaving  a  high  fronto-nasal 
ridge ;  short  pointed  posteriorly ;  sagittal  crest  long  and  high,  reach- 
ing anteriorly  nearly  to  midway  between  fronto-parietal  suture  and 
plane  of  postorbital  processes;  postorbital  processes  very  long, 
slender,  peglike,  and  horizontally  extended;  rostrum  long,  high, 
rather  narrow,  and  strongly  compressed  below  nasals;  palate  and 
postpalatal  shelf  exceedingly  long;  postpalatal  shelf  and  notch  nar- 
row; interpterygoid  fossa  exceptionally  deep;  basisphenoid  strongly 
concave.  Under  jaw  very  long,  ramus  flat  and  exceedingly  broad 
vertically;  coronoid  blade  high  and  moderately  recurved.  Dentition 
light  for  so  large  a  skull;  canines  rather  small;  upper  and  lower 
molariform  series  medium  or  rather  small;  middle  lower  molar  de- 
cidedly narrow ;  M-2-  small,  the  heel  narrowed  on  outer  side. 

An  imperfect  skull  of  an  old  male  (No.  167390)  from  Pine  Valley 
Mountain,  southwest  Utah,  differs  from  the  type  in  having  still 
smaller  teeth  both  above  and  below,  the  molars,  fourth  premolar,  and 
canines  being  but  little  larger  than  those  of  the  female  from  the  type 
locality. 

Adult  female  (No.  180207,  from  type  locality)  :  Similar  in  general 
to  male,  but  much  smaller  and  somewhat  less  extreme.  Skull  long 
and  narrow ;  f rontals  and  fronto-nasal  region  essentially  the  same  but 
sagittal  crest  shorter;  zygomata  relatively  as  well  as  actually  much 
narrower  (much  less  spreading)  and  not  outbowed;  rostrum  narrow- 
est anteriorly ;  molars  smaller ;  canines  much  smaller. 

Cranial  comparisons. — Old  male  (type)  compared  with  adult  and  old 
males  of  bairdi  (the  only  neighboring  species  of  approximately  same 
size)  :  Eostrum  longer  and  decidedly  narrower;  base  of  rostrum  in 
64854°— 18 5 


64  NORTH   AMERICAN   FAUNA.  [No.  41. 

front  of  orbits  more  compressed;  postorbital  processes  longer  and 
more  slender;  palate  longer;  postpalatal  shelf  narrower;  interptery- 
goid  canal  much  deeper;  ramus  of  jaw  longer,  decidedly  broader 
(vertically),  flatter,  and  much  thinner  under  M-y  and  M? ;  upper 
molariform  teeth,  middle  lower  molar,  and  lower  canines  much 
smaller. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  348 ;  occipito- 
nasal  length,  337;  palatal  length,  194;  zygomatic  breadth,  226;  inter- 
orbital  breadth,  79. 

URSUS  PERTURBANS  sp.  NOV. 
Mount  Taylob  Gbizzlt. 

Type  No.  222102,  $  old,  U.  S.  National  Museum,  Biological  Sur- 
vey collection.  Collected  near  Mount  Taylor,  northern  New  Mexico, 
July  9,  1916,  by  Ed.  Anderson. 

Characters. — Size  very  large;  skull  long  and  narrow,  with  nar- 
rowly spreading  zygomata  and  exceedingly  high  sagittal  crest ;  affini- 
ties with  utahensis,  idahoensis,  and  more  remotely  with  arizonoe. 
Claws  moderate,  slightly  curved,  mainly  ivory  whitish  on  top, 
darker  on  sides. 

Colon/'. — Adult  male  (type)  :  General  ground  color  dusky;  face  and 
head  dark  brown,  becoming  dusky  around  eyes;  body  dusky,  back 
grizzled  with  dark  golden  tips;  legs  and  feet  black. 

Cranial  characters. — Adult  male  (type)  :  Skull  conspicuously  long 
and  narrow,  the  narrowness  marked  in  braincase,  frontal  shield, 
rostrum,  palate,  postpalatal  notch,  and  basicranial  axis;  postorbital 
processes  moderately  outstanding  horizontally,  frontal  shield  broadly 
and  shallowly  concave,  becoming  flat  in  old  age,  very  short  poste- 
riorly, the  point  entering  sagittal  crest  in  midfrontal  region  (less 
than  halfway  from  postorbital  processes  to  fronto-parietal  suture) ; 
f ronto-nasal  region  very  slightly  dished ;  rostrum  narrow  and  rather 
high  in  type  specimen  (somewhat  broader  in  very  old  skull  from 
Datil  Mountains,  No.  140086) ;  frontal  part  of  braincase  keeling  into 
crest;  underjaw  long;  coronoid  blade  high;  ramus  straight,  its  in- 
ferior border  slightly  concave  under  MT,  only  slightly  upcurved 
posteriorly,  and  not  broadly  expanded  vertically;  diastema  long. 

Teeth  of  medium  size,  about  as  in  idahoensis  (much  smaller  than 
in  horribUis  and  bairdi) ;  heel  of  M-2-  long,  flat,  emarginate,  and 
slightly  everted,  resembling  that  of  idahoensis;  PM¥  strikingly 
small — much  smaller  than  in  any  other  known  grizzly  and  no  larger 
than  in  some  of  the  black  bears  (Euarctos),  its  crown  falling  below 
plane  of  molar  crowns.  Upper  canines  rather  small,  as  in  idahoensis 
and  arizonw;  lower  canines  more  slender  than  in  these  species. 


1918.]  ARIZONJE   GROUP.  65 

The  skull  of  a  still  older  male  (No.  140086)  killed  some  years  ago 
at  Kid  Springs,  Datil  Mountains,  New  Mexico,  10  miles  northeast  of 
Datil,  and  secured  for  the  Biological  Survey  by  N.  Hollister,  in 
October,  1905,  resembles  the  type  in  essential  characters,  but  is  even 
longer  and  owing  to  greater  age  has  the  frontal  shield  flatter,  the 
fronto-nasal  region  less  dished,  the  sagittal  crest  even  more  highly 
developed,  the  interpterygoid  fossa  even  longer  and  narrower. 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
utahensis  (type)  :  Size  and  general  appearance  similar,  but  underjaw 
widely  dissimilar ;  nasals  and  fronto-nasal  region  less  elevated ;  ros- 
trum smaller;  sagittal  crest  more  highly  developed;  zygomata  less 
spreading;  palate  and  postpalatal  shelf  shorter;  postpalatal  shelf 
less  narrowed;  underjaw  very  much  smaller,  shorter,  and  lighter, 
the  ramus  much  less  broadly  expanded  vertically,  its  inferior  border 
shorter  and  less  upcurved  posteriorly;  coronoid  blade  much  smaller 
and  lower.  Canines  much  smaller;  molars  decidedly  larger,  espe- 
cially M^;  heel  of  M^  much  larger  and  broader  posteriorly. 

Remarks. — Ursus  perturbans  appears  to  have  affinities  in  several 
directions.  In  length  and  slenderness  of  skull  it  resembles  utahensis 
more  closely  than  any  other  species,  but  the  underjaw  differs  amaz- 
ingly from  that  of  utahensis,  being  relatively  small  and  light,  while 
that  of  utahensis  is  large  and  remarkably  broad  vertically.  In 
dental  characters,  especially  the  form  of  M-2-,  it  resembles  idahoensis. 
In  the  great  development  of  the  sagittal  crest  it  exceeds  all  known 
grizzlies,  not  excepting  horribilis. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  338;  oc- 
cipito-nasal  length,  339;  palatal  length,  182.5;  zygomatic  breadth, 
210;  interorbital  breadth,  83. 

URSUS  ROGERSI  ROGERSI "  sp.  nov. 
Rogers  Grizzly. 

Type  No.  222983,  $  ad.,  U.  S.  National  Museum,  Biological  Survey 
collection.  Collected  high  up  on  Greybull  River,  Absaroka  Moun- 
tains, Wyoming,  in  the  fall  of  1890,  by  Archibald  Rogers,  and  by  him 
presented  to  the  Biological  Survey. 

Characters. — Skull  very  large  and  long,  the  length  of  the  adult 
male  equalling  or  slightly  exceeding  that  of  horribilis,  bairdi,  utahen- 
sis, and  perturbans ;  fronto-nasal  region  elevated  and  compressed, 
forming  part  of  long  fronto-nasal  plane,  as  in  bairdi  and  utahensis; 
frontal  shield  rather  narrow,  faintly  convex  transversely;  post- 
orbitals  rather  weak  and  somewhat  decurved ;  sinus  case  keeling  into 

1  Named  for  Archibald  Rogers,  of  New  York,  who  collected  and  presented  the  type 
specimen. 


66  NORTH   AMERICAN   FAUNA.  tNo.  41. 

sagittal  crest;  rostrum  long  and  rather  slender,  high  posteriorly, 
strongly  sloping;  palate  narrow  and  very  long;  interpterygoid  fossa 
narrow  and  very  long ;  zygomata  moderate,  rather  low,  as  in  utahensis 
and  idahoensis — not  arched  as  in  horribilis,  bairdi,  and  imperator; 
underjaw  long  and  massive  with  ramus  rather  broadly  expanded 
posteriorly,  but  very  much  less  so  than  in  utahensis.  Canines  small; 
molars  moderate.  Relationships  with  arizonaz,  bisonophagus,  and 
idahoensis,  but  size  much  larger;  and  also  with  utahensis,  with  which 
it  agrees  essentially  in  size. 

Cranial  comparisons. — Adult  male  (type,  perhaps  not  quite  fully 
adult)  compared  with  males  of  the  three  related  forms,  arizonce 
(type,  adult),  bisonophagus  (Jype,  young-adult)  and  idahoensis 
(type,  old)  :  General  appearance  similar,  but  skull  as  a  whole,  palate, 
and  interpterygoid  fossa  much  longer;  underjaw  very  much  longer, 
larger,  and  more  massive ;  coronoid  blade  broader  and  higher ;  canines 
of  approximately  same  length  but  lower  canines  more  massive 
basally ;  molars  in  general  similar,  but  last  upper  molar  longer,  with 
heel  rather  broadly  rounded  posteriorly,  instead  of  emarginate  or 
sub  triangular ;  MT  smaller;  crown  of  M^much  longer.  More  detailed 
comparisons  seem  unnecessary,  though  it  may  be  remarked  that  from 
idahoensis,  its  nearest  neighbor  on  the  west,  it  differs  markedly  not 
only  in  greater  size,  but  also  in  very  much  longer,  larger,  and  more 
massive  underjaw,  and  in  the  following  dental  characters:  M> 
smaller ;  heel  of  M  ^  broader  and  not  emarginate ;  M-g-  much  longer. 

Adult  male  (type)  compared  with  old  male  utahensis  (type) :  Size 
essentially  the  same  though  the  basal  length  is  greater;  frontal 
shield  somewhat  broader;  postorbitals  less  strongly  developed, 
shorter,  depressed  instead  of  horizontally  outstanding ;  rostrum  larger 
and  less  compressed;  interpterygoid  fossa  longer  and  less  deep, 
palate  and  postpalatal  shelf  broader;  underjaw  of  essentially  same 
length,  but  ramus  less  broadly  flattened.  Canines  apparently  less 
elongate  (in  utahensis  broken) ;  last  upper  molar  larger,  the  heel 
broader  and  more  broadly  rounded  posteriorly ;  crowns  of  middle  and 
last  lower  molars  longer. 

Skull  measurements. — Adult  male  (type) :  Basal  length,  353 ;  oc- 
cipito-nasal  length,  345 ;  palatal  length,  193 ;  zygomatic  breadth,  211 ; 
interorbital  breadth,  86. 

URSUS    ROGERSI    BISONOPHAGUS    subsp.    nov. 

Black  Hells  Gbizzly. 

Type  No.  181089,  $  young-adult,  U.  S.  National  Museum,  Biologi- 
cal Survey  collection.  From  Bear  Lodge,  Sundance  National  Forest, 
Black  Hills,  northeastern  Wyoming.  Collected  in  February,  1887, 
by  Paul  Kleineidam. 


1918.]  ABIZONiE   GROUP.  67 

Range. — Black  Hills  of  South  Dakota  and  adjacent  northeast  cor- 
ner of  Wyoming. 

Characters. — Size  large;  skull  long,  slender,  and  rather  low, 
smoothly  rounded  on  sides,  with  weak  decurved  postorbitals,  and 
elevated  fronto-nasal  region.  Affinities  with  arizonm  and  rogersi. 
Claws  of  moderate  length,  strongly  curved,  smoothly  polished,  dark 
horn  color,  marked  toward  tips  with  pale  yellowish,  and  most  of  them 
with  whitish  (superficially)  on  upper  side  of  basal  half. 

Color. — Type:  Muzzle  pale  brown  (apparently  old  pelage) ;  head 
and  face  blackish,  becoming  slightly  grizzled  posteriorly  and  on 
lower  part  of  cheeks  by  wash  of  yellowish-brown-tipped  hairs ;  entire 
body,  legs,  and  feet  very  dark  brown  overlaid  on  back  by  wash  of 
light  tips. 

Cranial  characters. — Young-adult  male  (type)  :  Similar  in  general 
to  rogersi.  Viewed  from  above:  Closely  similar  except  for  smaller 
size  and  differences  in  the  development  of  certain  parts  attributable 
in  the  main  to  lesser  age  (shield  more  convex  transversely;  post- 
orbitals slightly  less  outstanding;  fronto-nasal  region  slightly 
higher,  almost  forming  a  hump).  Viewed  from  below:  Palate  and 
postpalatal  shelf  very  much  shorter/  postpalatal  notch  less  narrow; 
underj aw  smaller  and  lighter;  canines  longer;  M1  slightly  larger; 
M-2-  with  heel  subtriangular,  strongly  narrowed  on  outer  side  as  in 
arizonce  (in  rogersi  not  narrowed  but  rather  broadly  rounded) ;  MT 
slightly  larger;  MT  much  smaller.  Fully  adult  skulls  would  doubt- 
less show  other  differences. 

Cranial  comparisons. — Young-adult  male  (type)  compared  with 
adult  male  arizonw  (type)  :  Skull  and  teeth  similar  but  skull  longer 
and  narrower  anteriorly;  vault  of  cranium  slightly  less  arched; 
rostrum  narrower ;  fronto-nasal  region  slightly  more  compressed  and 
more  elevated,  continuing  frontal  plane;  frontal  shield  slightly  nar- 
rower, somewhat  more  convex  transversely,  its  sides  more  smoothly 
rounded  (doubtless  because  slightly  younger)  ;  postorbital  processes 
less  developed;  lambdoid  crest  more  strongly  developed;  postpalatal 
shelf  smaller.  Underj  aw  slightly  longer;  inferior  border  of  ramus 
longer  and  more  upcurved  posteriorly;  ramus  more  broadly  ex- 
panded vertically;  coronoid  blade  higher  and  flatter  (fossa  less 
deeply  excavated),  its  anterior  border  more  nearly  vertical;  upper 
molars  somewhat  larger ;  middle  lower  molars  smaller ;  upper  canines 
essentially  the  same;  lower  canines  somewhat  longer  and  more 
slender. 

Remarks. — The  range  of  bisonophagus  appears  to  be  completely 
isolated  from  that  of  its  nearest  relative  rogersi  of  the  mountains  be- 
tween Yellowstone  Park  and  Bighorn  Basin.  On  the  other  hand,  its 
range  appears  to  overlap  parts  of  those  of  absarokus,  horribilis,  and 
bairdi.    From  bairdi.  which  it  resembles  in  form  of  skull,  it  is  easily 


68  NORTH   AMERICAN   FAUNA.  [No.  41. 

distinguished   by   decidedly   smaller   size   and   very   much   smaller 
canines  and  molars. 

Skull  measurements. — Young-adult  male  (type)  :  Basal  length, 
331 ; x  occipito-nasal  length,  323 ;  palatal  length,  173 ;  zygomatic 
breadth,  200 ;  interorbital  breadth,  80. 

URSUS  PERVAGOR  Mebeiam. 

Lillooet  Grizzly. 

Ursus  pervagor  Merriam,  Proc.  Biol.   Soc.  Washington,  XXVII,  pp.   186-187, 
August  13,  1914 

Type  locality. — Pemberton  Lake  (now  Lillooet  Lake),  British 
Columbia. 

Type  specimen. — No.  187887,  $  ad.,  U.  S.  National  Museum 
(=No.  6510,  Merriam  collection).  Collected  in  May,  1883,  by  John 
Fannin. 

Range. — Interior  of  southwestern  British  Columbia;  known  only 
from  Lillooet  Lake  and  Bridge  River. 

Characters. — Size  rather  large.     External  characters  unknown. 

Cranial  characters. — Adult  male  (type)  :  Size  large;  skull  long, 
rather  narrow,  high,  moderately  dished;  zygomata  moderately 
spreading  and  outbowed ;  frontal  shield  of  moderate  breadth,  rather 
flat,  strongly  sloping,  shallowly  sulcate  medially,  swollen  on  each 
side  just  behind  plane  of  postorbitals ;  postorbital  processes  outstand- 
ing, thick,  peglike;  postpalatal  shelf  rather  broad  and  flat;  frontal 
part  of  braincase  elevated  and  compressed,  supporting  posterior  part 
of  frontal  shield ;  palate  long ;  squamosal  shelf  long ;  mastoids  long ; 
interpterygoid  fossa  short  and  rather  broad;  under  jaw  long;  coro- 
noid  broad  and  rather  vertical;  ramus  long,  swoll  n  on  outer  side. 
Teeth  small,  particularly  the  canines  and  lower  molars. 

Cranial  comparisons. — Adult  male  (type)  compared  with  adult 
male  canadensis  (type)  :  Considerably  larger ;  vault  of  cranium  much 
higher;  frontal  shield  much  broader,  higher,  more  strongly  sloping, 
and  less  flat;  frontal  part  of  braincase  elevated  and  compressed, 
rising  strongly  to  temporal  impressions  (in  canadensis  not  com- 
pressed except  at  sagittal  crest)  ;  postorbital  processes  much  larger; 
rostrum  more  strongly  ascending;  occipito-sphenoid  longer  (98  mm. 
against  90)  ;  underjaw  much  longer;  coronoid  blade  higher,  its  apex- 
less  recurved.  Canines  about  same  size ;  large  upper  premolar,  upper 
molars,  and  MT  and  MT  decidedly  smaller. 

Adult  male  (type)  compared  with  adult  male  caurinus:  Frontal 
shield  broader,  less  elevated  posteriorly;  fronto-nasal  region  less 
strongly  dished;  coronoid  blade  less  nearly  vertical;  upper  canines 
slightly  shorter;  lower  canines  conspicuously  smaller  and  shorter. 

1  Restored. 


1918.]  ARIZONA   GROUP.  69 

Adult  male  (type)  compared  with  adult  male  eulophus  (type  and 
older  skulls)  :  Similar  in  general,  both  having  the  fronto-facial 
region  long  sloping  and  flattish,  but  differing  in  many  characters. 
The  skull  of  pervagor  differs  from  that  of  eulophus  in  being  slightly 
shorter,  less  highly  arched,  frontal  shield  less  elevated  posteriorly; 
postorbitals  larger,  more  horizontally  outstanding;  sagittal  crest 
shorter  and  straighter,  not  arched;  braincase,  rostrum,  and  palate 
shorter;  squamosal  shelf  shorter;  postpalatal  shelf  broader  and 
shorter;  interpterygoid  notch  broader;  under  jaw  smaller  and  less 
massive ;  inferior  border  of  ramus  shorter  and  less  broadly  expanded ; 
coronoid  lower,  the  apex  less  narrowed  and  less  recurved,  with  less 
development  of  inferior  ridge  of  fossa;  teeth  (canines  and  molars) 
slightly  smaller. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  330; 
occipito-nasal  length,  322;  palatal  length,  178;  zygomatic  breadth, 
224 ;  interorbital  breadth,  81. 

URSUS  CAURINUS  Merbiam. 

Lynn  Canal  Gbizzly. 

Vrsus  caurinus  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  187,  August  18, 
1914. 

Type  locality. — Berners  Bay,  east  side  of  Lynn  Canal,  Southeast- 
ern Alaska. 

Type  specimen. — No.  176591,  2  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.     Collected  June  8,  1911  by  A.  Hasselborg. 

Range. — Coast  of  mainland  of  Southeastern  Alaska  from  Cnilkat 
River  valley  and  Lynn  Canal  south  an  unknown  distance. 

Characters. — Very  closely  related  to  eulophus  of  Admiralty  Island. 
Size  rather  large;  skull  long  and  rather  narrow;  canines  long,  the 
lower  ones  massive;  claws  smoothly  polished. 

Color. — Upperparts  yellowish  buff;  face  and  most  of  head  pale 
brown  or  drab;  ears,  hump,  and  underparts  conspicuously  darker; 
legs  and  feet  dark  brown  or  brownish  black. 

Cranial,  characters. — Male  and  female :  Both  skulls  long  and  nar- 
row, strongly  arched  posteriorly,  moderately  dished;  frontal  shield 
of  medium  breadth,  strongly  ascending;  postorbital  processes  weak 
and  decurved  except  in  old  age,;  frontal  part  of  braincase  elevated, 
forming  an  uplifted  base  for  posterior  part  of  frontal  shield,  behind 
which  it  keels  into  sagittal  crest  as  in  eulophus;  palate  long;  post- 
palatal  shelf  rather  narrow;  squamosal  shelf  long;  lachrymal  duct 
opening  within  orbital  rim;  ramus  of  jaw  broad  and  flattened, 
notably  higher  posteriorly  than  anteriorly.  Teeth  rather  small  for 
size  of  skull;  last  upper  molar  long  and  narrow,  tapering  posteriorly. 


70  NORTH  AMERICAN   FAUNA.  [No.  41. 

Unfortunately,  no  fully  adult  male  caurmus  has  been  obtained, 
but  I  have  secured  a  skull  of  a  young-adult  male  (No.  205169)  from 
Berners  Bay,  Lynn  Canal,  and  another  (No.  210140)  from  Chilkat 
River  valley.  These  present  the  following  characters:  Skull  long, 
high,  and  narrow,  rather  highly  arched  and  dished;  frontals  rising 
rather  strongly  from  rostrum;  frontal  shield  rather  narrow,  long 
sloping,  convex  transversely  in  these  youngish  skulls  (doubtless  flat- 
tish  in  adults),  slightly  sulcate  medially;  long  pointed,  the  point 
nearly  reaching  parietals;  postorbital  processes  rather  broad  and 
slightly  decurved  (doubtless  more  outstanding  with  age) ;  fronto- 
nasal region  dished;  rostrum  moderate;  nasals  nearly  horizontal 
anteriorly,  rising  posteriorly  inj:o  frontal  shield;  frontal  part  of 
braincase  compressed  and  elevated,  keeling  into  temporal  impressions 
and  anterior  part  of  sagittal  crest ;  squamosal  shelves  long.  Under- 
jaw rather  long;  ramus  moderate,  its  inferior  border  bellied  pos- 
teriorly; coronoid  blade  moderate  and  rather  vertical;  apex  not 
strongly  recurved ;  palate,  postpalatal  shelf  and  notch  medium ;  teeth 
rather  small  for  size  of  skull ;  molars  only  slightly  larger  than  those 
of  female ;  M-2-  rather  narrow,  with  narrow  slightly  everted  heel ; 
canines,  especially  lower  canines,  decidedly  thicker  than  those  of 
female;  MT rather  swollen  in  both  male  and  female;  M-j-  small  and 
narrowest  posteriorly;  PMT  with  main  cusp  rather  small,  conical, 
anterior,  with  gradually  sloping  incompletely  sulcate  heel,  rarely 
with  traces  of  posterior  cusplets,  sometimes  with  anterior  cusplet 
on  inner  side  of  cingulum. 

The  Chilkat  Valley  male  is  a  year  older  than  the  Berners  Bay 
skull,  and  broader  across  the  frontals;  postorbital  processes  more 
outstanding,  ramus  of  underjaw  more  bellied  posteriorly;  coronoid 
blade  broader  above;  MT smaller  and  thinner. 

Cranial  comparisons. — Vrsus  caurinus  appears  to  be  rather  closely 
related  to  pervagor  of  the  Lillooet  region  in  the  interior  of  British 
Columbia,  and  to  eulophus  of  Admiralty  Island,  Southeastern  Alaska, 
but  unhappily  no  skulls  of  fully  adult  males  are  available  for  com- 
parison. 

Young-adult  male  (No.  210140,  from  Chilkat  River  valley)  com- 
pared with  adult  male  eulophus  (type)  and  with  a  young  adult 
eulophus  (No.  203284),  both  from  Admiralty  Island:  Frontal  shield 
narrower  anteriorly  and  much  narrower  posteriorly  (narrowed  be- 
hind postorbitals  by  usual  incurving  temporal  ridges,  while  in 
eulophus  the  ridges  are  nearly  straight  and  the  posterior  part  of 
shield  correspondingly  broader)  ;  fronto-nasal  region  more  strongly 
dished;  underjaw  longer;  ramus  thicker  (more  swollen  on  outer 
side)  ;  coronoid  blade  broader  above  and  much  less  recurved.  Canines 
essentially  same;  MT  smaller;  M-2-  broader  in  middle  and  posteriorly. 

Adult  female  (type)  compared  with  young-adult  female  eulophus 
(No.  137470) :  Basal,  occipito-nasal,  and  occipito-sphenoid  lengths 


1918.]  ARIZONA  GROUP.  71 

essentially  same;  vault  of  cranium  much  less  arched;  frontal  shield 
flatter,  narrower  interorbitally  and  postorbitally ;  postorbitals  much 
smaller  and  outstanding  instead  of  strongly  decurved;  fronto-nasal 
region  scarcely  dished ;  rostrum  lower ;  nasals  smaller ;  palate  shorter ; 
under  jaw  more  massive;  coronoid  broader  and  less  recurved;  MT 
thicker;  M^  narrower  throughout,  the  heel  narrowed  on  outer  side. 

Young-adult  male  compared  with  adult  male  pervagor:  Frontal 
shield  narrower,  more  elevated  posteriorly ;  fronto-nasal  region  more 
strongly  dished ;  coronoid  blade  more  nearly  vertical.  M-2-  narrower, 
with  narrower  heel.  Upper  canines  slightly  longer;  lower  canines 
conspicuously  longer  and  larger. 

Skull  measurements. — Adult  female  (type):  Basal  length,  295; 
occipito-nasal  length,  285;  palatal  length,  161;  zygomatic  breadth, 
196 ;  interorbital  breadth,  66. 

URSUS  EULOPHUS  Mebriam. 

Admiralty  Island  Crested  Bear. 

Ursus  eulophus  Merriam,  Proc.  Biol.  Soc.  Washington,  XVII,  p.  153,  October  6, 
1904. 

Type  locality. — Admiralty  Island,  Southeastern  Alaska. 

Type  specimen. — No.  81102,  $  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.    Collected  in  1896  by  Lieut.  G.  T.  Emmons. 

Range. — Admiralty  Island. 

Characters. — Size  large;  color  rich  dark  brown;  claws  blue-black, 
of  moderate  length ;  skull  long,  rather  narrow,  and  high,  with  weak 
decurved  postorbital  processes. 

Color.1 — General  color  of  head  and  body  in  fresh  pelage,  rich  dark 
brown  or  seal  brown;  muzzle  paler;  legs,  feet,  and  belly  dusky  or 
blackish;  neck  and  shoulders  sometimes  grizzled  by  admixture  of 
yellowish-tipped  hairs. 

Cranial  characters. — Adult  males:  Skull  large,  long,  high,  and 
rather  narrow;  frontal  shield  long  and  rather  narrow,  shallowly 
grooved  medially,  gradually  sloping  (not  abruptly  elevated) ;  tem- 
poral impressions  long  and  only  slightly  incurved,  meeting  at  fronto- 
parietal suture;  postorbital  processes  weak  and  decurved;  fronto- 
nasal region  elevated  in  plane  of  shield;  rostrum  long  and  high; 
anterior  third  of  nasals  horizontal,  posterior  two-thirds  rising  in 
fronto-nasal  plane;  zygomata  moderately  spreading,  subangular; 
squamosal  root  only  slightly  expanded;  palate  and  postpalatal  shelf 
long  and  narrow ;  ascending  arms  of  maxillae  long,  passing  far 
beyond  nasals  and  reaching  back  over  anterior  two-thirds  of  orbit; 
nares  rather  small;  braincase  long  anteriorly,  rather  narrow,  com- 

1  Color  from  skins  in  Mus.  Vert.  Zool.,  Univ.  California,  obtained  on  Admiralty  Island 
by  Miss  Annie  M.  Alexander. 


72  NORTH   AMERICAN   FAUNA.  [No.  41. 

pressed,  and  keeling  into  sagittal  crest;  sagittal  crest  high,  curved, 
and  relatively  short;  ramus  of  jaw  high  vertically,  its  inferior  border 
strongly  bellied  posteriorly;  coronoid  blade  large  and  moderately 
high.    Canines  long ;  molars  rather  small  for  so  large  a  skull. 

Cranial  comparisons. — Adult  male  (type)  compared  with  adult 
male  pervagor  (type)  :  Skull  longer;  arch  of  cranium  higher;  frontal 
shield  longer  sloping,  rising  higher  posteriorly;  rostrum  longer; 
braincase  longer;  sagittal  crest  more  convex;  palate  and  postpalatal 
shelf  longer;  shelf  and  notch  narrower;  under  jaw  larger,  longer, 
more  massive ;  inferior  border  of  ramus  longer,  more  strongly  bellied 
posteriorly  and  broader  vertically ;  coronoid  fossa  deeper,  its  inferior 
border  much  more  sharply  defined  by  strongly  developed  ridge  for 
muscular  attachment.    Canines  and  molars  larger. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  346;  oc- 
cipito-nasal  length,  343;  palatal  length,  190;  zygomatic  breadth, 
221 ;  interorbital  breadth,  81. 

URSUS  KLAMATHENSIS  Meeeiam.1 

Klamath  Gbizzly.  • 

Vrsus  klamathensis  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  185- 
186,  August  13,  1914. 

Type  locality. — Beswick,  near  mouth  of  Shovel  Creek,  Klamath 
Eiver,  northern  California. 

Type  specimen. — No.  178735,  $  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  and  presented  by  Charles  Far- 
well  Edson. 

Range. — Siskiyou  Mountains  of  northern  California  and  southern 
Oregon,  ranging  north  in  recent  times  to  Fort  Klamath  region  and 
Rogue  River  valley ;  in  early  days  to  lower  Willamette  Valley  (pre- 
sumably same  species) ;  south  in  Sierra  Nevada  an  unknown  distance. 
(Skull  from  lower  McCloud  River  referred  to  this  species.) 

Characters. — Size  of  male  large ;  skull  in  general  of  the  idahoensis 
type,  but  larger  and  with  heavier  canines.  Claws  moderate,  rather 
strongly  curved,  horn  color,  washed  with  yellowish  basally  and  with 
pale  yellowish  markings  at  tips,  marked  longitudinally  with  fine 
parallel  striae.    Skin  characters  unknown. 

Cranial  characters. — Skull  large  and  high,  highest  about  40 
mm.  in  front  of  fronto-parietal  suture;  rostrum  long,  high,  and 
ascending  in  plane  of  frontal  shield;  fronto-nasal  region  elevated, 
scarcely  if  at  all  dished ;  frontal  shield  broad,  flat,  sloping,  and  rather 
short  pointed;  postorbital  processes  moderate,  peglike,  horizontally 
outstanding ;  sagittal  crest  long  and  well  developed ;  lachrymal  duct 

»  Tentatively  included  in  arizonce  group.     (See  Introduction,  pp.  12-13.) 


1918.]  ARIZONA   GROUP.  73 

opening  on  orbital  rim  (rather  posteriorly  than  anteriorly) ;  zygo- 
mata moderately  spreading,  only  moderately  expanded  vertically; 
anterior  (frontal)  part  of  bra  incase  keeling  into  sagittal  crest; 
occiput  produced  posteriorly  (overhang  much  greater  than  in 
calif  ornicus) ;  squamosal  shelves  long  and  broad;  palate  rather  flat, 
slightly  arched  lengthwise;  postpalatal  shelf  broad  and  flat;  occipito- 
sphenoid  short  for  so  large  a  skull  (length  89  mm.) ;  basioccipital 
very  broad  anteriorly;  mastoids  vertical  and  short.  Under  jaw  long; 
ramus  exceptionally  broad  and  flat  vertically;  coronoid  blade  large 
and  high,  broad  basally.  Canines  very  large ;  molars  moderate ;  last 
upper  molar  relatively  small,  the  heel  emarginate  or  obliquely  trun- 
cate on  outer  side;  middle  lower  molar  with  anterior  moiety  much 
larger  than  posterior.  Large  upper  premolar  absent  and  no  trace 
of  alveolus. 

Cranial  comparisons. — Curiously  enough  klamathensis  does  not  re- 
quire close  comparison  with  any  of  the  other  species  inhabiting  Cali- 
fornia, its  only  near  relatives  being  members  of  the  shoshone- 
idahoensis  group  of  the  Rocky  Mountains,  and  pervagor  of  interior 
British  Columbia. 

Adult  male  (type)  compared  with  old  male  idahoensis  (No.  187888, 
=No.  93,  Merriam  collection,  from  North  Fork  Teton  River, 
eastern  Idaho)  :  Similar  in  general  form  and  proportions  but  larger 
( condylobasilar  length  350  mm.  contrasted  with  335) ;  vault  of  cra- 
nium somewhat  higher;  frontal  shield  broader  and  flatter;  palatal 
length  about  the  same;  postpalatal  length  much  greater  (150  con- 
trasted with  135 )  ;  opening  of  lachrymal  duct  slightly  more  posterior ; 
occipital  overhang  greater;  basioccipital  anteriorly  very  much 
broader;  ramus  of  under  jaw  longer  and  much  broader  vertically; 
coronoid  blade  higher.  Canines  larger  and  longer ;  last  upper  molar 
shorter. 

Adult  male  (type)  compared  with  adult  male  pervagor  (type) : 
Similar  in  size  and  general  characters;  vault  of  cranium  slightly 
higher ;  frontal  shield  flatter  and  somewhat  broader ;  postorbitals  not 
quite  so  large;  fronto-nasal  region  more  elevated  (in  pervagor 
slightly  dished);  rostrum  broader  anteriorly;  occipital  overhang 
greater;  ramus  of  under  jaw  much  more  broadly  expanded  vertically 
and  flatter;  canines  and  molars  very  much  larger. 

Adult  male  (type)  compared  with  old  male  henshawi  (type)  :  Ursus 
klamathensis  and  U.  henshawi  belong  to  widely  different  groups  and 
do  not  require  detailed  comparison.  U.  klamathensis  may  be  distin- 
guished at  a  glance  by  its  much  larger  size,  much  higher  vault  of 
cranium,  highly  elevated  and  continuously  sloping  fronto-nasal 
region  and  rostrum,  and  peglike  postorbitals — in  striking  contrast 
to  the  much  smaller,  lower,  and  strongly  dished  skull  of  henshawi, 


74  NORTH   AMERICAN   FAUNA.  [No.  41. 

with  its  low  depressed  rostrum  and  large  broadly  rounded  post- 
orbitals. 

Adult  male  (type)  compared  with  adult  male  californicus  (from 
coast  region  south  of  San  Francisco  Bay)  :  The  differences  are 
marked  in  the  skull  and  striking  in  the  teeth.  In  klamathensis  the 
vault  of  the  cranium  is  lower  posteriorly  and  higher  anteriorly ;  the 
frontal  shield  flatter  laterally;  the  rostrum  shorter;  the  base  of  the 
cranium  (occipito-sphenoid)  decidedly  shorter.  The  last  upper  and 
middle  lower  molars  are  widely  different,  the  heel  of  M>  in  cali- 
fornicus large,  long,  and  broad  posteriorly,  while  in  klamathensis  it 
is  small  and  emarginate  on  outer  side;  the  anterior  part  of  M7  in 
californicus  is  normal,  while  in  klamathensis  it  is  disproportionately 
large. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  331 ;  oc- 
cipito-nasal  length,  337;  palatal  length,  175;  zygomatic  breadth,  223; 
interorbital  breadth,  85.5. 

URSUS  MENDOCINENSIS  Meekiam.1 

Mendocino  Grizzly. 

Ursus  mendocinensis  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  145-146, 
September  6,  1916. 

Type  locality. — Long  Valley,  Mendocino  County,  California. 

Type  specimen.— No.  206625,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Obtained  through  Charles  J.  and  Frank 
H.  Hittell. 

Characters. — Size  rather  large,  about  equaling  klamathensis  but 
apparently  smaller  than  californicus  and  colusus;  external  characters 
unknown.  Affinities  with  klamathensis,  with  which  it  may  inter- 
grade  at  the  north. 

Cranial  characters. — Skull  short,  broad,  highly  arched,  and 
strongly  dished,  with  widely  outstanding  zygomata  and  truncate  occi- 
put. Frontal  shield  of  moderate  breadth,  short  pointed  posteriorly, 
slightly  convex  between  orbits,  strongly  sloping  to  rostrum ;  rostrum 
short,  broad,  and  strongly  depressed ;  postorbital  processes  moderate, 
sub-peglike,  horizontally  outstanding;  sagittal  crest  high,  thick, 
humped  anteriorly,  short  posteriorly ;  occiput  obliquely  truncate ;  oc- 
cipital overhang  slight  compared  with  that  of  californicus  and  colu- 
sus; palate  short  and  rather  broad;  postpalatal  shelf  of  moderate 
breadth,  flat;  postpalatal  notch  moderate;  mastoids  rather  short, 
directed  anteriorly.  Under  jaw  absent.  Teeth  gone  except  left  hind 
molar,  which  is  short,  with  small  heel,  obliquely  truncate  on  outer 
side  (as  in  klamathensis). 

1  Tentatively  included  in  arizonw  group.      (See  Introduction,  pp.  12-13.) 


1918.]  ARIZONA   GROUP.  75 

Cranial  comparisons. — Similar  in  general  to  klamathensis  but 
frontonasal  region  strongly  dished,  rostrum  shorter,  broader,  flatter 
on  top,  and  depressed  instead  of  elevated;  zygomata  more  widely 
outstanding;  palate  broader;  occipital  overhang  less. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  327;  occi- 
pito-nasal  length,  323;  palatal  length,  183;  interorbital  breadth,  84.5. 

URSUS  MAGISTER  Merriam.1 

Southeen  California  Grizzly. 

Ursus  magister  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  189,  August 
13,  1914. 

Type  locality. — Los  Biacitos,  head  of  San  Onofre  Canyon,  Santa 
Ana  Mountains,  Southern  California. 

Type  specimen. — No.  160155,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Killed  in  August,  1900  or  1901,  by  Henry 
A.  Stewart  and  by  him  presented  to  the  Biological  Survey. 

Range. — Santa  Ana  or  Trabuco  Mountains,  Cuyamaca  and  Santa 
Rosa  Mountains,  and  probably  San  Jacinto  Mountains.  Believed  to 
be  extinct. 

Characters. — Size  of  male  huge  (estimated  weight  over  1,400 
pounds),  largest  of  known  grizzlies,  considerably  larger  than  cali- 
fomicus  of  the  Monterey  region,  and  even  than  horribilis,  the 
great  buffalo-killing  grizzly  of  the  Plains  (only  equaled  by  the 
largest  alexandrce  of  Kenai  Peninsula) ;  sexual  disparity  great;  skull 
of  female  hardly  half  the  bulk  of  male;  skull  of  male  of  a  rather 
generalized  type;  not  dished.  Claws  of  old  female  from  head  of 
Trabuco  Canyon,  Santa  Ana  Mountains,  exceedingly  long,  strongly 
curved,  mainly  yellowish  above. 

Color. —  (Old  female  from  head  of  Trabuco  Canyon) :  General 
color  dusky  or  sooty  all  over  except  head  and  grizzling  of  back. 
Muzzle  gray  or  mouse  brown,  palest  above;  top  of  head  and  neck 
very  dark  brown,  sparsely  grizzled  with  pale-tipped  hairs;  back 
dusky  grizzled  with  grayish;  legs  and  underparts  wholly  blackish. 

Cranial  characters. — Adult  male  (type)  :  Skull  exceedingly  large, 
long;  vault  of  cranium  arched,  but  not  abruptly;  rostrum  long  and 
high ;  f ronto-nasal  region  elevated,  in  same  plane  with  frontal  shield 
and  rostrum;  frontal  shield  flattish-convex,  faintly  sulcate  medially 
and  slightly  swollen  on  each  side  between  postorbital  processes,  the 
point  decurved  and  reaching  fronto-parietal  suture;  zygomata  only 
moderately  spreading,  angular,  the  posterior  root  expanded  and 
rising  abruptly  from  plane  of  squamosal  shelf;  sagittal  crest  rather 
short;  palate  scooped  out  anteriorly,  forming  a  basinlike  depression 
surrounding    the    anterior    palatine    foramina;    occipito-sphenoid 

1  Tentatively  included  in  arizona  group.     (See  Introduction,  pp.  12-13.) 


76  NORTH   AMERICAN   FAUNA.  [No.  41. 

length  103.5  mm.  Underjaw  long;  ramus  broad  and  flat  vertically, 
its  inferior  border  moderately  bellied  and  incurved  posteriorly; 
coronoid  blade  large,  its  apex  strongly  recurved,  cutting  plane  of 
middle  of  condyle.  Teeth  large  and  broad  but  by  no  means  dis- 
proportionate to  large  size  of  skull ;  M-1  broad ;  last  upper  molar 
absent,  but  from  its  alveolus  and  its  form  in  female,  obviously  broad, 
short,  strongly  triangular,  the  heel  small,  narrowed  posteriorly,  ob- 
liquely emarginate  on  outer  side;  PM¥  broad,  with  rather  short 
slightly  sloping  heel,  narrow  imperfect  sulcus  without  posterior 
cusplets;  MT  apparently  normal  (much  worn  in  type  specimen) ; 
middle  lower  molar  absent  in  type  specimen  but  apparently  normal 
(judging  from  the  female,  in  which,  however,  it  is  badly  worn). 

Female  of  extreme  age  (No.  156594,  from  Trabuco  Canyon,  killed 
January  5,  1908,  by  Andrew  Joplin  and  Edward  Adkinson)  :  Size 
small;  rostrum  short  and  depressed;  fronto-nasal  region  strongly 
dished;  frontal  shield  flattish,  slightly  sulcate  interorbitally,  short 
pointed,  beaded  posteriorly  by  elevated  temporal  impressions,  rising 
rather  abruptly  at  orbits;  sagittal  crest  long  and  nearly  horizontal; 
palate  and  postpalatal  shelf  broad,  flat  posteriorly,  concave  ante- 
riorly. 

Cranial  comparisons. — Ursus  magister  does  not  require  close  com- 
parison with  any  other  species.  While  the  largest  skulls  of  old  male 
calif ornicus  equal  it  in  basal  length,  they  are  so  much  lower,  nar- 
rower, and  smaller  in  every  way  that  detailed  comparisons  are  un- 
necessary. The  species  which  it  most  nearly  resembles  is  bairdi  from 
the  mountains  of  Colorado,  but  the  resemblance  is  not  close.  It 
differs  from  bairdi  in  somewhat  larger  size,  much  more  highly 
arched  vault  of  cranium,  much  broader  and  more  strongly  sloping 
frontal  shield,  more  posterior  mastoids,  longer  underjaw  with  much 
more  broadly  flattened  and  less  massive  ramus,  and  in  important 
tooth  characters. 

Between  the  two  geographically  is  utahensis,  which,  like  magister, 
has  the  ramus  of  the  underjaw  very  broadly  flattened  vertically,  but 
in  form  of  cranium  utahensis  goes  to  the  opposite  extreme,  the  frontal 
shield,  rostrum,  and  braincase  being  exceptionally  narrow,  and  the 
fronto-nasal  region  compressed  and  elevated. 

Flesh  measurements. — Old  male  (type)  :  Height  at  shoulder  from 
flat  of  foot  4  ft.  (=1,220  mm.) ;  total  length,  snout  to  tail,  9£  ft.1 
(=2,900  mm.);  sole  of  largest  foot  without  claws:  length  12  in. 
(=305  mm.)  ;  breadth  8  in.  (=204  mm.).  Length  of  old  female  from 
Trabuco  Canyon,  measured  in  the  flesh  by  Andrew  Joplin,  6  ft.  3  in. 

Skull  measurements. — Old  male  (type):  Basal  length,  365;  oc- 
cipito-nasal  length,  366;  palatal  length,  197;  zygomatic  breadth,  236; 
interorbital  breadth,  97. 

*  Apparently  an  error ;  possibly  intended  for  snout  to  claws  of  extended  hind  foot. 


1918.]  HYLODROMUS  GROUP.  77 

Hylodromus  Group. 

URSUS  HYLODROMUS  Elliot. 

Forest  Grizzly. 

(Plate  XI.) 

Ursus  hylodromus  Elliot,  Field  Columb.  Mus.  Pub.  87,  Zool.  Ser.  Ill,  pp  257- 
258,  December,  1913.  (Described  as  a  black  bear!)  Purchased  from 
Mackay  &  Dippie,  taxidermists,  Calgary. 

Ursus  selkirki  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  p.  150,  September 
6,  1916.  (From  Selkirk  Mountains,  Upper  Columbia  River,  British  Co- 
lumbia.) 

Type  locality. — Eocky  Mountains  of  western  Alberta  (precise  lo- 
cality unknown). 

Type  specimen. — No.  19065,  5  young-adult,  Field  Museum  of  Nat- 
ural History. 

Range. — Rocky  Mountain  region  of  western  Alberta  and  eastern 
British  Columbia,  including  Selkirk  Range. 

Characters. — Size  of  male  large,  of  female  small;  external  charac- 
ters unknown. 

Cranial  characters. — Young -adult  female  (type *) :  Size  small,  skull 
short  posteriorly  (occiput  less  extended  than  in  most  species) ;  mod- 
erately arched  and  dished;  zygomata  not  outstanding  and  only 
slightly  bowed;  frontal  shield  of  moderate  breadth,  rising  rather 
strongly  from  rostrum,  its  apex  sublyrate ;  postorbital  processes  large, 
outstanding;  rostrum  rather  high,  strongly  ascending  posteriorly; 
sagittal  crest  short,  reaching  only  two-thirds  distance  from  occiput 
to  fronto-parietal  suture;  squamosal  shelf  narrow  and  nearly  hori- 
zontal; postpalatal  shelf  rather  broad. 

Old  male  (No.  205170)  from  Selkirk  Mountains,  upper  Columbia 
River,  British  Columbia  (assumed  to  be  typical)  :  Size  medium;  skull 
long,  low  arched,  highest  immediately  in  front  of  fronto-parietal 
suture,  and  of  medium  breadth;  braincase  and  palate  arched;  shield 
flattish,  long  pointed,  sloping  gradually  from  point  to  rostrum, 
faintly  sulcate  medially;  postorbitals  broadly  subtriangular,  flat  on 
top  and  slightly  decurved,  convex  posteriorly,  concave  anteriorly; 
rostrum  moderate,  nearly  horizontal ;  f ronto-nasal  region  rising  very 
gradually  into  shield ;  sagittal  crest  short ;  palate  of  medium  breadth, 
slightly  troughed  and  arched ;  postpalatal  shelf  broad  and  flat ;  zygo- 
mata subtriangular,  not  widely  outstanding,  strongly  arched  antero- 
posterior^; squamosal  root  long  but  not  broadly  expanded;  mas- 

1  Through  the  courtesy  of  the  officials  of  the  Field  Museum  I  have  had  the  privilege  of 
examining  the  type  skull  Of  hylodromus  and  comparing  it  with  skulls  in  the  National 
Museum  collection. 


78  NORTH   AMERICAN   FAUNA.  [No.  41. 

toids  short,  not  divergent;  meatus  tube  large  and  free;  ramus  of 
underjaw  straight;  coronoid  blade  low,  broad  in  middle  part.  Teeth 
of  medium  size  (too  badly  worn  to  admit  of  description). 

Cranial  comparisons. — Old  male  (No.  205170,  from  Selkirk  Moun- 
tains) compared  with  old  male  kluane  (type)  :  Size  smaller;  vault  of 
cranium  materially  lower ;  top  of  skull  lower  and  flatter  throughout ; 
shield  narrower,  much  flatter,  and  longer  pointed ;  sagittal  crest  much 
shorter  and  lower ;  occipital  overhang  and  inion  much  less  developed ; 
zygomata  more  triangular  (less  bowed)  ;  braincase,  palate,  and 
underjaw  much  shorter;  coronoid  blade  lower;  subangular  border 
shorter  and  more  strongly  defined.  Teeth  very  badly  worn  but  ca- 
nines and  molars  evidently  mucl>  smaller. 

Compared  with  idahoensis  and  chelan  (both  types)  :  Similar  in 
general  but  smaller;  vault  of  cranium  much  lower;  frontal  shield 
somewhat  narrower,  longer  pointed  posteriorly ;  postorbital  processes 
broader  basally  but  less  widely  outstanding;  sagittal  crest  much 
shorter,  its  anterior  part  less  distinctly  keeled  from  sinus  case; 
zygomata  shorter  and  more  sharply  triangular;  underjaw  shorter; 
coronoid  blade  lower;  subangular  notch  and  border  similar. 

Compared  with  latifrons,  whose  range  it  approaches  on  the  north 
but  with  which  it  does  not  appear  to  be  related,  it  is  easily  dis- 
tinguished by  smaller  size,  flatter  and  very  much  narrower  frontal 
shield,  more  elevated  and  evenly  sloping  fronto-nasal  region,  less 
outbowed  and  more  triangular  zygomata,  much  shorter  underjaw  and 
ramus,  and  much  lower  coronoid  blade. 

Remarks. — The  type  specimen  of  hylodromus  is  a  skull  of  a  young- 
adult  female  from  western  Alberta,  exact  locality  unknown.  Until 
recently  so  few  males  have  been  available  from  this  region  that  I  was 
long  in  doubt  as  to  which  was  its  proper  mate.  In  the  light  of  pres- 
ent material,  however,  it  has  been  possible  to  match  up  males  and 
females  of  most  of  the  species  of  eastern  British  Columbia  and  west- 
ern Alberta  with  some  confidence,  and  I  now  feel  reasonably  certain 
that  the  type  specimen  of  the  species  described  by  me  as  Ursus  sel- 
hirhi  from  the  Selkirk  Mountains  on  the  upper  Columbia  River  is 
in  reality  an  old  male  hylodromus,  the  name  selkirki  thus  falling  as  a 
synonym. 

Skull  measurements. — Old  male  (No.  205170)  from  Selkirk  Moun- 
tains: Basal  length,  305;  occipito-nasal  length,  306;  palatal  length, 
169;  zygomatic  breadth,  206;  interorbital  breadth,  74.  Female 
young-adult  (type):  Basal  length,  275;  occipito-nasal  length,  257; 
palatal  length,  154 ;  zygomatic  breadth,  173 ;  interorbital  breadth,  69. 


1918.]  HYLODROMUS   GROUP.  79 

URSUS  KLUANE  KLUANE  Meebiam. 

Kxuane  Grizzly. 

Ursus  kluane  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  141-143,  Sep- 
tember 6,  1916. 

Type  locality. — McConnell  River,  Yukon. 

Type  specimen. — No.  204188,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  by  Smith  and  Geddis  July  15, 
1914. 

Range. — Southwest  corner  of  Yukon  Territory  east  of  the  St.  Elias 
Range,  extending  northwesterly  in  Alaska  to  Mount  McKinley  region 
(head  of  Toklat),  easterly  in  Yukon  Territory  to  McConnell  River 
(north-northeast  of  Teslin  Lake)  and  probably  south  into  northwest 
corner  of  British  Columbia. 

Cranial  characters. — Adult  male:  Skull  medium,  rather  long,  nar- 
row, somewhat  arched  and  dished,  with  long  braincase,  long  convex 
sagittal  crest,  and  unusually  broad  decurved  postorbitals.  Frontal 
shield  of  medium  width,  strongly  convex  both  transversely  and 
antero-posteriorly,  rising  rather  strongly  from  rostrum,  slightly  sul- 
cate  medially  and  moderately  swollen  over  orbits ;  very  short  pointed, 
the  point  ending  about  midway  between  parietals  and  plane  of  post- 
orbitals; postorbitals  broad,  decurved,  strongly  convex  anteriorly, 
concave  posteriorly ;  f ronto-nasal  region  somewhat  depressed ;  ros- 
trum high  and  narrow,  rounded  above  (subterete)  ;  nares  truncate; 
sagittal  crest  very  long  and  arcuate;  occipital  overhang  and  inion 
well  developed;  zygomata  not  widely  outstanding,  somewhat  bowed, 
rounded  posteriorly;  palate  moderate;  postpalatal  shelf  large  and 
broad;  notch  rather  broad  and  short;  mastoids  long  and  divergent; 
underjaw  rather  long;  coronoid  blade  high  and  narrow,  the  apex 
rather  strongly  recurved;  teeth  too  badly  worn  to  admit  of  descrip- 
tion (apparently  large  for  size  of  skull). 

Adult  female:  Size  small,  nearly  as  small  as  female  pallasi; 
f ronto-nasal  region  moderately  dished  and  usually  sulcate ;  braincase 
moderately  arched,  highest  just  in  front  of  fronto-parietal  suture; 
temporal  impressions  meeting  over  anterior  part  of  parietals  (prob- 
ably somewhat  more  anteriorly  in  old  skulls)  ;  zygomata  moderately 
outbowed,  subtriangular ;  frontal  shield  of  medium  breadth,  lyrate 
pointed  posteriorly;  postorbital  processes  rather  broad  for  so  small 
a  skull,  moderately  decurved;  underjaw  short;  coronoid  blade  broad 
basally  and  rather  short.  Teeth  (canines,  incisors,  and  molars) 
rather  large  for  size  of  skull,  decidedly  larger  than  in  pallasi;  molars, 
both  upper  and  lower,  very  much  larger. 
64854°— 18 6 


80  NORTH   AMERICAN   FAUNA.  [No.  41. 

Cranial  comparisons. — The  species  requiring  comparison  with 
hluane  are  toklat,  pulchellus,  and  pallasi.  Old  male  (type)  com- 
pared with  old  male  toklat,  from  Alaska  Range,  near  north  base  of 
Mount  McKinley:  Size  slightly  larger;  occipito-nasal  length,  length 
of  braincase,  and  length  of  sagittal  crest  very  much  greater ;  frontal 
shield  more  convex  transversely;  postorbital  processes  much  larger 
and  broader;  rostrum  higher,  more  rounded  on  top;  nares  more 
squarely  truncate;  underjaw  longer;  inferior  border  of  ramus  more 
convex  posteriorly ;  coronoid  blade  decidedly  higher,  narrower  above, 
the  apex  more  strongly  recurved;  teeth  badly  worn  in  both,  but 
canines  decidedly  longer  in  kluane;  molars  apparently  somewhat 
larger.  / 

Compared  with  adult  male  pulchellus :  Similar  in  general,  but  skull 
as  a  whole,  braincase,  palate,  and  underjaw  very  much  longer; 
frontal  shield  very  much  broader;  inferior  border  of  ramus  less 
abruptly  upcurved;  subangular  border  less  nearly  horizontal  and 
much  less  sharply  defined. 

Adult  female  compared  with  adult  female  toklat  (comparison 
hardly  necessary  because  of  the  great  difference  in  size) :  Basal  length 
at  least  20  mm.  less ;  vault  of  cranium  and  frontal  shield  lower ;  brain- 
case less  constricted  anteriorly ;  posterior  part  of  shield  much  longer 
and  broader,  reaching  or  passing  the  f ronto-parietal  suture ;  sagittal 
crest  much  shorter;  postpalatal  shelf  less  broad;  underjaw  and 
inferior  border  of  ramus  shorter ;  coronoid  blade  about  same  height ; 
canines  about  same  size;  molariform  series  (upper  and  lower)  about 
same  length,  but  proportions  of  individual  teeth  different:  M^-much 
larger ;  M>  with  shorter  heel ;  MT  larger. 

Old  male  (type)  compared  with  old  male  pallasi  (type)  :  Size  de- 
cidedly greater;  skull  about  an  inch  longer  and  much  more  highly 
arched,  with  conspicuously  longer  braincase  and  longer  sagittal  crest; 
crest  strongly  arched  instead  of  nearly  straight;  postorbitals  very 
much  larger,  broader,  and  more  strongly  decurved;  fronto-nasal 
region  much  more  elevated  and  less  dished;  rostrum  much  higher, 
rounded  above  instead  of  depressed ;  palate  much  longer,  more  arched 
and  more  concave;  underjaw  much  longer;  coronoid  higher. 

Adult  female  compared  with  adult  female  pallasi:  Skulls  very 
much  alike  in  size  and  appearance  (that  of  kluane  slightly  larger), 
but  teeth  strikingly  different.  In  kluane,  canines  larger;  molars  very 
much  larger. 

Skull  measurements. — Old  male  (type):  Basal  length,  317;  oc- 
cipito-nasal length,  324;  palatal  length,  177;  zygomatic  breadth,  210; 
interorbital  breadth,  85. 


1918.]  HYLODROMUS   GROUP.  81 

URSUS  KLUANE  IMPIGER  subsp.  nov. 

Industrious  Geizzlt. 

Type  No.  210708,  $  ,  not  quite  fully  adult.  From  Columbia  Valley, 
British  Columbia.    Collected  in  April,  1914,  by  Mackay  &  Dippie. 

Cranial  characters. — Adult  male:  Similar  in  general  to  kluane  (No. 
221620,  $  ad.,  Kluane  River),  but  shield  narrower  and  less  sulcate, 
less  swollen  over  orbits ;  nasals  less  elevated ;  palate  much  shorter  and 
narrower;  postpalatal  shelf  narrower;  under  jaw  shorter;  subangular 
border  shorter;  subangular  notch  and  space  much  more  strongly  de- 
fined. Teeth  smaller  throughout,  especially  M>  and  My,  but  My 
swollen  as  in  kluane. 

Female  young-adult  from  Brisco,1  Columbia  Valley  (No.  210707)  : 
Vault  well  arched;  shield  posteriorly  broadly  lyrate,  elevated  by 
rising  sinus  case;  rostrum  long  and  slender;  fronto-nasal  region 
dished;  palate  long,  troughed;  underjaw  long  and  straight.  Teeth 
large. 

Cranial  comparisons. — Male  young-adult  (type)  compared  with 
old  male  hylodromus  from  Selkirk  Mountains  (No.  205170) :  Vault 
of  cranium  much  more  highly  arched  and  narrower,  the  narrow 
frontal  shield  rising  higher  and  more  strongly,  but  not  abruptly, 
from  rostrum;  shield  strongly  convex  (in  old  hylodromus  nearly 
flat)  ;  postorbitals  much  more  slender,  outstanding,  depressed,  and 
somewhat  decurved  as  in  kluane';  fronto-nasal  region  more  dished; 
palate  narrower;  inferior  border  of  underjaw  much  longer;  sub- 
angular  border  shorter;  coronoid  blade  higher. 

Female  young-adult  (No.  210707)  compared  with  female  kluane: 
Skull  as  a  whole,  rostrum,  occipito-sphenoid,  palate,  and  underjaw 
much  longer ;  ramus  straighter ;  subangular  border  same ;  teeth  essen- 
tially same  except  that  MT  is  smaller.  Compared  with  female  hylo- 
dromus (type)  :  Skull  as  a  whole,  rostrum,  occipito-sphenoid,  palate, 
and  underjaw  longer  (but  not  so  much  longer  as  in  comparison  with 
female  kluane) ;  ramus  straighter;  subangular  border  shorter;  teeth 
similar. 

Remarks. — Ursus  hylodromus,  U.  impiger,  and  U.  kluane  form  a 
rather  closely  related  group  ranging  from  western  Alberta  and  south- 
eastern British  Columbia  northwesterly  to  southwestern  Yukon. 
From  the  material  now  in  hand  impiger  appears  to  be  most  nearly 

1  Skulls  of  females  identified  as  impiger  have  been  examined  from  Brisco,  Columbia 
Valley,  British  Columbia  (No.  210707)  ;  Morley,  Alberta  (No.  210706)  ;  Jasper,  Alberta 
(No.  222745)  ;  and  headwaters  North  Fork  Blackfoot  River,  western  Montana  (No. 
203188). 


82  NORTH   AMERICAN   FAUNA.  [No.  41. 

related  to  kluane,  but  additional  skulls  of  adults  may  show  that  its 
affinities  with  hylodromus  are  equally  close. 

/Skull  measurements. — Adult  male  (type):  Basal  length,  309;  oc- 
cipito-nasal  length,  293;  palatal  length,  162;  zygomatic  breadth,  197; 
interorbital  breadth,  72. 

URSUS  PELLYENSIS  sp.  nov. 
Pelly  Grizzly. 

Type  No.  215477,  $  young-adult.  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  From  the  Ketza  Divide,  Pelly  Moun- 
tains, Yukon.    Collected  September  30,  1915,  by  Fred  E.  Enevoldsen. 

Characters. — Skull  of  medium  size,  rather  long  and  narrow,  ap- 
parently related  to  hylodromus :  Frontal  shield  narrow,  long  pointed 
(point  reaching  fronto-parietal  suture  but  in  older  specimens  ending 
much  more  anteriorly),  rising  strongly  from  rostrum;  rostrum 
rather  narrow  and  high;  fronto-nasal  region  dished;  postorbitals 
broad,  strongly  decurved  (in  old  skulls  doubtless  more  outstanding)  ; 
sinus  case  keeling  into  posterior  part  of  shield;  palate  moderate 
(rather  broad  for  so  narrow  a  skull),  slightly  arched  and  slightly 
troughed;  under  jaw  long  for  size  of  skull;  inferior  border  of  ramus 
long,  not  upturned;  subangular  border  short;  coronoid  rather  high; 
canines  long,  the  lower  ones  large  for  size  of  skull ;  molars  long  and 
narrow,  especially  MT  and  MT;  anterior  part  of  M¥  exceptionally 
long  and  posterior  part  relatively  narrower  than  in  most  species; 
PM?  subconical. 

Female  skull  (based  on  four  rather  old  specimens  from  Pelly  and 
Ross  Mountains,  Nos.  215710, 215711, 215713,  and 221600)  :  Size  small; 
frontal  shield  of  moderate  breadth,  rather  short  pointed  posteriorly, 
entering  sagittal  crest  anterior  to  fronto-parietal  suture;  postorbital 
processes  well  developed,  outstanding ;  rostrum  rather  small ;  frontal 
shield  rising  moderately  from  rostrum,  shallowly  sulcate  medially; 
zygomata  subtriangular.  Dentition  moderate;  heel  of  last  upper 
molar  slightly  emarginate,  rounded  posteriorly;  canines  small  and 
slender. 

Cranial  comparisons. — Young-adult  male  (type)  compared  with 
young-adult  male  hylodromus  (No.  210708  from  Columbia  Valley, 
British  Columbia)  :  Size  smaller  (basal  length  about  10  mm.  less)  ; 
vault  of  cranium  less  highly  arched;  sinus  case  more  definitely  keeled 
to  support  posterior  part  of  shield;  postorbitals  broader,  more  de- 
curved,  less  outstanding;  palate  broader;  molar  series  about  same; 
upper  canines  essentially  same;  lower  canines  larger;  upper  molar 
series  essentially  same  length;  lower  molar  series  longer  and  nar- 
rower; MT  much  longer  and  more  slender. 


1918.]  HYLODROMUS  GROUP.  83 

Young-adult  male  (type)  compared  with  adult  male  pulchellus 
from  the  neighboring  Eoss  River  (No.  .221599)  :  Size  slightly 
greater;  shield  and  rostrum  broader  (in  pulchellus  exceptionally 
narrow);  postorbitals  broader  and  more  decurved;  palate  longer 
and  broader;  postpalatal  notch  much  shorter;  coronoid  blade  larger; 
molars,  upper  and  lower,  strikingly  narrower  and  less  massive. 

Female  pellyensis  compared  with  female  toklat:  Size  smaller;  vault 
much  lower  and  depressed — sinus  case  not  keeled  into  front  of  sagit- 
tal crest;  fronto-nasal  region  less  strongly  depressed  and  dished; 
rostrum  and  nasals  flatter;  palate  about  same  length;  postpalatal 
length  much  less;  under  jaw  much  shorter.  Canines  smaller;  molar 
series  shorter;  heel  of  M-2-  less  elongate. 

Skull  measurements. — Young-adult  male  (type)  :  Basal  length, 
299;  occipito-nasal  length,  294;  palatal  length,  168;  zygomatic 
breadth,  186 ;  interorbital  breadth,  73. 

URSUS  ANDERSONI1  sp.  Nov.* 
Anderson  Bear. 

Type  No.  34402,  $  ad.  (rather  old),  American  Museum  of  Nat- 
ural History.  Collected  on  east  branch  Dease  River 3  near  Great 
Bear  Lake,  Mackenzie,  May  12,  1911,  by  Dr.  R.  M.  Anderson. 

Characters. — Size  medium  or  rather  large;  frontal  region  includ- 
ing postorbitals  rather  flat  and  narrow  as  in  pellyensis  (in  strong 
contrast  with  the  broader  swollen  frontals  and  elevated  outstanding 
postorbitals  of  richardsoni)  ;  vault  moderately  arched. 

Cranial  characters. — Frontal  shield  rather  narrow,  strongly  slop- 
ing, flattened,  short  pointed  posteriorly  ;  postorbitals  moderate  (short 
in  contrast  with  those  of  richardsoni),  horizontal  (not  decurved); 
sagittal  crest  long  and  strongly  developed,  humped  anteriorly,  pro- 
jecting posteriorly  in  prominent  inion;  rostrum  narrow,  rather 
high,  compressed  below  middle  of  nasals;  orbits  strongly  sloping 
(retreating)  ;  occipital  overhang  marked;  squamosal  shelf  long;  pal- 
ate long  and  narrow;  postpalatal  shelf  moderate;  zygomata  well  out- 
standing and  somewhat  bowed.  Underjaw  moderate ;  coronoid  blade 
strongly  recurved,  apex  sharp  pointed.  Molars  broad  and  rather 
large:  MA  and  M„  together  62  mm.  (in  type  skull)  ;  lower  molars  74 
mm.    Canines  large  (too  much  broken  to  afford  measurements). 

Cranial  comparisons. — The  adult  male  resembles,  though  not  very 
closely,  both  pellyensis  and  tahltanicus.  Compared  with  the  type 
specimen  of  pellyensis:  Size  essentially  same;  shield  shorter  and 
flatter  with  more  horizontally  outstanding  postorbitals   (differences 

1  Named  for  Dr.  R.  M.  Anderson,  who  collected  the  type  specimen. 

2  Tentatively  included  in  hylodromus  group.      (See  Introduction,  pp.  12-13.) 

8  Not  to  be  confused  with  the  better-known  Dease  River  of  northern  British  Columbia. 


84  NORTH   AMERICAN   FAUNA.  [No.  41. 

attributable  in  large  part  at  least  to  difference  in  age) ;  zygomata 
much  less  broadly  outstanding  (also  attributable  to  age)  ;  postpalatal 
notch  decidedly  longer;  coronoid  blade  lower.  Canines  apparently 
larger  (badly  broken) ;  molars  broader  and  more  massive.  Com- 
pared with  an  old  male  (No.  134486)  from  Macmillan  River,  pre- 
sumably old  petty  ensis,  it  is  much  smaller  and  narrower;  shield 
narrower,  flatter,  and  shorter  posteriorly;  postorbitals  flatter  and 
more  nearly  horizontal  on  top;  rostrum  narrower;  sagittal  crest 
longer  and  rising  higher  anteriorly;  palate,  postpalatal  shelf,  di- 
astema, and  occipito-sphenoid  much  shorter;  under  jaw  much  shorter; 
coronoid  lower.    Dentition  heavier ;  canines  and  molars  larger. 

Compared  with  old  male  tahltdnicus  (type),  with  which  it  agrees 
essentially  in  size,  height  of  vault  of  cranium,  slope  and  flatness  of 
shield,  form  of  rostrum,  spread  and  form  of  zygomata,  and  size  and 
form  of  underjaw,  it  differs  as  follows:  Shield  narrower  both  inter- 
orbitally  and  across  postorbitals;  point  of  shield  much  shorter; 
fronto-nasal  region  slightly  dished  (in  tahltanicus  not  dished)  ; 
sagittal  crest  much  longer,  higher  anteriorly,  and  keeled  from  sinus 
case.    Dentition  heavier ;  canines  and  molars  much  larger. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  300;  oc- 
cipito-nasal  length,  285 ;  palatal  length,  165 ;  zygomatic  breadth,  211 ; 
interorbital  breadth,  76. 

Horriaeus  Group. 

URSUS  APACHE  Merriam:. 

Apache  Grizzly. 

Ursus  apache  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  134-135,  Sep- 
tember 6,  1916. 

Type  locality. — Whorton  Creek,  south  slope  of  White  Mountains, 
eastern  Arizona  (a  few  miles  west  of  Blue). 

Type  specimen. — No.  212436,  $  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.    Collected  April  3,  1913,  by  B.  V.  Lilly. 

Cranial  characters. — Skull  short,  broad,  and  low,  rather  massive, 
moderately  dished,  with  broad  frontal  shield  and  exceedingly  broad 
outstanding  postorbitals.  Frontal  shield  broad,  shallowly  sulcate 
medially  between  orbits ;  very  slightly  and  rather  flatly  swollen  over 
orbits;  long  pointed  posteriorly,  meeting  short  sagittal  crest  at 
fronto-parietal  suture;  rostrum  short,  high,  and  rather  narrow; 
zygomata  strongly  outbowed  and  outstanding  anteriorly  as  well  as 
posteriorly ;  ramus  of  jaw  rather  short,  bellied  under  last  molars; 
coronoid  blade  high,  sloping  strongly  outward,  the  apex  overarching 
shallow  coronoid  notch,  but  not  cutting  plane  of  condyle;  dentition 
moderate. 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
horriceus  (type)  :  Size  somewhat  greater ;  vault  of  cranium  materially 


1918.]  HOKBLEUS  GROUP.  85 

higher;  shield  much  broader  both  interorbitally  and  across  postor- 
bital  processes;  the  posterior  part  much  longer  and  horizontal  or 
sloping  posteriorly  (in  horriceus  short  and  sloping  strongly  forward 
from  apex  of  posterior  point);  sagittal  crest  very  much  shorter; 
palate  broader  and  flat  (in  horriceus  concave  and  arched) ;  zygomata 
broadly  outbowed  (in  horriceus  narrowly  triangular)  ;  meatus  tube 
strongly  compressed  between  mastoid  and  glenoid  (in  horriceus  large 
and  free)  ;  underjaw  much  more  massive;  ramus  longer;  broadly  ex- 
panded vertically  and  upcurved  posteriorly;  coronoid  blade  higher. 
Teeth  apparently  similar  (in  horriceus  badly  worn). 

Compared  with  adult  male  absarokus,  to  which  it  appears  to  be  re- 
lated, the  skull  of  apache  differs  as  follows :  Vault  of  cranium  lower, 
less  arched;  frontal  shield  broader  and  flatter;  postorbitals  much 
broader  and  flatter,  standing  out  more  horizontally ;  f ronto-nasal  re- 
gion more  depressed;  rostrum  shorter;  orbits  notably  smaller  (lower 
vertically);  squamosal  trough  shorter  antero-posteriorly ;  zygomata 
very  much  more  strongly  outbowed  and  conspicuously  more  out- 
standing anteriorly;  underjaw  and  inferior  border  of  ramus  shorter; 
coronoid  blade  of  equal  height;  teeth  slightly  smaller  (difference 
slight). 

Old  male  (type)  compared  with  adult  male  arizonce  (type)  :  Basal 
length,  occipito-nasal  length,  length  of  palate,  interorbital  breadth, 
and  occipito-sphenoid  length  essentially  the  same;  zygomata  very 
much  more  outstanding  and  bowed  instead  of  subtriangular ;  frontal 
shield  less  flat,  of  essentially  the  same  breadth  interorbitally  as  in 
arizonce,  but  very  much  wider  across  postorbital  processes,  rising 
strongly  from  plane  of  rostrum;  postorbital  processes  much  more 
broadly  outstanding;  orbital  rims  more  swollen;  fronto-nasal  region 
much  less  elevated;  rostrum  much  smaller,  narrower,  less  swollen, 
depressed  basally,  and  horizontal  instead  of  tapering;  palate  and 
postpalatal  shelf  much  narrower.  Underjaw  stronger;  ramus 
broader  vertically;  coronoid  blade  higher;  molars  slightly  smaller; 
heel  of  M>  shorter,  less  distinctly  emarginate  on  outer  side  (more 
tapering) . 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  325 ; 
occipito-nasal  length,  315;  palatal  length,  171.5;  zygomatic  breadth, 
234 ;  interorbital  breadth,  89. 

URSUS  HORRI^US  Baibd. 

New  Mexico  Grizzly. 

(PI.   XV.) 

Ursus   horribilis   var.    horriceus   Baird,    Mammals   North   Amer.,    Pacific    RR. 

Reports,  VIII,  pp.  224,  225,  1857  (name,  type  locality,  and  measurements). 
Ursus  horriMUs  var.  horriceus  Baird,  Mammals  Mexican  Boundary  Survey,  pp. 

24-29,  1859.     (Full  description,  including  a  specimen  from  Nogales,  since 

made  the  type  of  another  species — kennerlyi.) 


86  NORTH   AMERICAN   FAUNA.  [No.  41. 

Ursus    {Darin)    horriaceus  Gray,   Catalog.   Cam.,   Pach.,   and   Edent,   British 
Museum,  p.  229,  1869. 

Type  locality. — Coppermines,  southwestern  New  Mexico. 

Type  specimen. — No.  990,  $  old,  U.  S.  National  Museum.  Col- 
lected in  1855,  by  J.  H.  Clark. 

Range. — Parts  of  New  Mexico,  south  to  Casas  Grandes,  Chihuahua, 
Mexico;  probably  extending  into  eastern  Arizona. 

Characters. — Size  medium;  external  characters  unknown;  skull 
low  and  flat  with  broad  outstanding  postorbital  processes. 

Cranial  characters. — Old  male  (type)  :  Skull  long,  low,  narrow,  and 
flat;  frontal  shield  short  pointed  posteriorly,  temporal  impressions 
conspicuously  beaded,  curving  strongly  inward,  and  meeting  half- 
way between  plane  of  postorbital  processes  and  fronto-parietal 
suture ;  frontal  shield  flatfish,  shallowly  concave,  swollen  over  orbits ; 
postorbital  processes  large,  broad,  widely  outstanding  and  moderately 
decurved;  fronto-nasal  region  slightly  dished;  rostrum  narrow;  sag- 
ittal crest  long  and  rather  high ;  occipital  overhang  and  inion  marked ; 
zygomatic  arches  angular,  rather  squarely  but  not  widely  spreading 
posteriorly;  squamosal  root  of  zygoma  moderately  expanded  ver- 
tically; lachrymal  duct  cutting  orbital  rim  anterior  to  orbit;  palate 
and  interpterygoid  fossa  rather  narrow.  Teeth  of  medium  size 
(badly  worn  in  type). 

An  old  female  (No.  67405,  from  mountains  north  of  Silver  City, 
New  Mexico,  near  type  locality),  collected  in  1893  by  Dr.  A.  K. 
Fisher,  is  assumed  to  be  horriceus.  Unfortunately  the  occipital  region 
is  absent  so  that  measurements  of  length  can  not  be  taken.  Skull  low, 
with  moderately  spreading  angular  zygomata,  flat  long-pointed 
frontal  shield,  elevated  flat  fronto-nasal  region  (in  plane  of  shield), 
and  narrow  rostrum,  without  trace  of  dishing  except  slight  change  of 
angle  at  middle  of  nasals;  highest  point  of  cranium  about  25  mm. 
anterior  to  fronto-parietal  suture;  postorbital  processes  moderate  or 
weak,  subtriangular,  slightly  decurved;  lachyrmal  duct  cutting 
orbital  rim  anteriorly;  rostrum  somewhat  compressed  a  little  below 
nasals;  palate  short  (145),  concave  between  last  molars;  postpalatal 
shelf  moderately  broad,  short,  and  flat.    Teeth  small. 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
apache  (type)  :  Size  somewhat  smaller ;  vault  of  cranium  lower ;  shield 
much  narrower,  the  point  much  shorter;  palate  concave  and  arched 
instead  of  flat ;  zygomata  narrowly  triangular  instead  of  broadly  out- 
bowed;  ramus  of  jaw  shorter. 

Old  male  (type)  compared  with  adult  male  shoshone  (type)  :  Size 
essentially  the  same  (basal  length  slightly  less  but  occipito-nasal 
length  same);  vault  of  cranium  much  lower  and  flatter;  frontal 
shield  much  lower  and  more  nearly  horizontal,  less  strongly  sloping, 


1918.]  HORRIiEUS  GROUP.  87 

slightly  broader,  more  swollen  over  orbits,  and  broadly  depressed 
interorbitally  instead  of  flat  or  slightly  convex ;  postorbital  processes 
very  much  larger,  broader,  and  more  widely  outstanding;  frontal 
part  of  braincase  broader  and  depressed — not  keeling  into  sagittal 
crest;  zygomata  angular — not  outbowed;  occipito-sphenoid  shorter; 
lachrymal  duct  more  anterior;  inferior  border  of  ramus  shorter; 
coronoid  blade  less  high. 

Old  female  compared  with  old  females  of  nelsoni:  Size  larger 
(basal  length  about  10  mm.  longer)  ;  vault  of  cranium  higher  over 
posterior  frontal  region;  frontal  shield  broader,  flatter  anteriorly; 
zygomatic  arches  slightly  more  spreading.  Teeth,  especially  canines 
and  molars,  materially  larger. 

Skull  measurements. — Old  male  (type):  Basal  length,  312;  oc- 
cipito-nasal  length,  310;  palatal  length,  169;  zygomatic  breadth,  207; 
interorbital  breadth,  79. 

URSUS  HENSHAWI *  Merriam. 

Henshaw  Grizzly. 

Ursus  henshawi  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  190,  August 
13,  1914. 

Type  locality. — Southern  Sierra  Nevada,  near  Havilah,  Kern 
County,  California. 

Type  specimen. — No.  15671,  $  old,  U.  S.  National  Museum.  Col- 
lected in  1875  by  Dr.  J.  T.  Rothrock  and  Henry  W.  Henshaw. 

Range. — Lower  slopes  of  southern  part  of  Sierra  Nevada;  limits 
unknown. 

Characters. — Size  rather  small — by  far  the  smallest  of  the  Cali- 
fornia grizzlies;  size  and  general  cranial  characters  as  in  horriceus,  but 
fronto-nasal  region  strongly  dished  and  rostrum  strongly  depressed. 
Last  upper  molar  short  and  broad,  the  heel  short  and  subtriangular. 
Skin  characters  unknown. 

Cranial  characters. — Old  male  (type)  :  Skull  long,  narrow,  and 
rather  low;  zygomata  subtriangular,  narrowly  spreading;  frontal 
shield  gently  sloping,  flat-concave ;  postorbital  processes  massive,  and 
somewhat  arched ;  orbital  rims  swollen ;  fronto-nasal  region  strongly 
depressed  and  dished ;  rostrum  low,  depressed. 

Cranial  comparisons. — Compared  with  an  equally  old  male  hor- 
riceus  (type,  from  Coppermines,  New  Mexico)  :  Surprisingly  similar 
in  general,  with  similar  broad  outstanding  postorbitals,  but  rostrum 
strongly  depressed ;  nasals  flattened  and  horizontal  anteriorly ;  fronto- 
nasal region  concave  and  strongly  dished  (in  horriceus  rather  high 
and  not  dished)  ;  frontal  shield  strongly  and  broadly  concave  between 
orbits  and  between  postorbital  processes  (only  faintly  depressed  medi- 

*  Named  for  Henry  W.  Henshaw,  formerly  chief  of  the  Biological  Survey. 


88  NORTH  AMEBIOAN   FAUNA.  [No.  41. 

ally  in  horriceus) ;  more  strongly  sloping  anteriorly;  postorbital  pro- 
cesses shorter  and  blunter;  orbital  rims  more  prominent,  relatively 
thin,  somewhat  everted,  continuing  to  lachrymal  notch — their  promi- 
nence anteriorly  due  in  part  to  presence  of  a  broad  sulcus  in  ascending 
arm  of  maxillary  immediately  in  front  of  orbit ;  lachrymal  duct  open- 
ing in  orbit  posterior  to  orbital  rim  (in  horriceus  on  or  anterior  to 
rim) ;  anterior  nares  broader  than  high  (contrary  true  in  horriceus) ; 
lambdoid  crest  higher ;  sagittal  crest  and  inion  much  shorter ;  occipi- 
tal overhang  much  less;  palate  decidedly  broader;  interpterygoid 
canal  shorter;  mastoid  processes  much  longer  and  more  divergent; 
anterior  part  of  pterygoids  more  broadly  expanded  vertically  and 
articulating  with  a  like  expansjxm  of  posterior  arms  of  palatines 
(probably  not  constant).  Upper  molars  decidedly  broader  (canines 
broken  off). 

Compared  with  two  old  males  of  tularensis  (type,  No.  3536,  and 
No.  3537,  from  Fort  Tejon,  California)  :  Size  smaller;  occipito- 
sphenoid  length  and  frontal  breadth  essentially  the  same;  vault  of 
cranium  and  rostrum  very  much  lower;  fronto-nasal  region  more 
deeply  concave,  more  strongly  sloping,  and  strikingly  more  dished; 
rostrum  smaller,  lower,  and  strongly  depressed  instead  of  elevated; 
occipital  overhang  and  development  of  inion  conspicuously  less; 
posterior  part  of  sagittal  crest  not  elevated  or  produced.  Under- 
jaw  decidedly  shorter,  ramus  much  less  broadly  flattened  vertically 
and  more  strongly  bellied  posteriorly.  Last  upper  molar  much 
smaller,  the  heel  conspicuously  shorter  and  narrowed  on  outer  side. 

Skull  measurements. — Old  male  (type):  Basal  length,  318;  oc- 
cipito-nasal  length,  305 ;  palatal  length,  173 ;  zygomatic  breadth,  204 ; 
interorbital  breadth,  76. 

Stikeenensis  Group.  • 

URSUS  STIKEENENSIS  Merbiam. 

Stikine  Gbizzlt. 

(PI.  IX.) 

Ursus  stikeenensis  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  178-179, 
August  13,  1914. 

Type  locality. — Tatletuey  Lake,  tributary  to  Finlay  River,  near 
head  of  Skeena  River,  northern  British  Columbia. 

Type  specimen. — No.  202794,  &  ad.,  U.  S.  National  Museum, 
Biological  Survey  collection.  Collected  September  23,  1913,  by 
Charles  R.  Cross,  jr.,  and  Edward  A.  Preble ;  original  No.  5772. 

Range. — Region  about  head  of  Finlay  River,  and  Dease  Lake 
region,  northern  British  Columbia,  and  northerly  in  Yukon. 


1918.]  STTKEENENSIS  GROUP.  89 

Characters. — Adult  male  (type)  :  Size  medium ;  skull  short,  broad, 
and  highly  arched;  face  strongly  pugged  from  abrupt  rising  of 
frontal  region;  claws  short  and  strongly  curved  for  a  grizzly  (longest 
60  mm. — tips  worn  by  digging),  dark,  marked  with  yellowish  on 
tips  and  sides.  Upper  molars  large.  Total  length  before  skinning 
1,830  mm. ;  hind  foot  267  mm. ;  estimated  height  at  shoulder  990  mm. 
(39  inches). 

Color. — Type  specimen:  General  ground  color  dark  brown,  griz- 
zled and  washed  with  pale-tipped  hairs.  Muzzle  pale  brown,  becom- 
ing darker  between  eyes  and  on  sides  of  face;  top  of  head  in  front 
of  ears  washed  with  yellowish  brown,  almost  forming  a  golden 
brown  band  between  the  darker  ungrizzled  frontal  region  and  the 
more  whitish  color  of  the  neck;  general  ground  color  of  body  very 
dark,  bountifully  overlaid  on  neck  and  back  by  pale  buffy-tipped 
hairs  which  give  a  whitish  cast  to  the  neck ;  ears  dark ;  legs  and  feet 
blackish ;  underparts  dark  brown ;  hairs  on  sides  of  throat  long  and 
grizzly. 

Cranial  characters. — Adult  male  (type):  Size  medium  or  large; 
skull  short  posteriorly,  broad,  highly  arched,  strongly  dished,  with 
abruptly  ascending  frontals  and  large  molar  teeth.  Rostrum  short, 
broad,  broadening  and  strongly  ascending  posteriorly;  nasals 
strongly  upturned  posteriorly;  frontal  shield  rather  broad,  rising 
abruptly  and  swollen  in  front  of  and  above  orbits,  sulcate  medially, 
short  pointed,  the  point  ending  in  sagittal  crest  about  30  mm.  ante- 
rior to  fronto-parietal  suture;  postorbital  processes  large,  broad, 
subtriangular,  and  decurved  (more  broadly  rounded  and  more  out- 
standing in  older  skull)  ;  braincase  long  for  size  of  skull;  zygomata 
rather  broadly  spreading,  subtriangular,  not  much  expanded  verti- 
cally; palate  and  postpalatal  shelf  broad.  Underjaw  and  inferior 
border  of  ramus  very  short.    Molars  large  and  broad. 

Cranial  comparisons. — Adult  male  compared  with  adult  male 
absarokus :  Size  nearly  the  same  though  absarokus  appears  the  larger ; 
basal  length  and  frontal  breadth  approximately  the  same,  but  oc- 
cipito-nasal  length  much  greater  in  absarokus.  In  stikeenensis  ros- 
trum lower,  flatter,  and  more  nearly  horizontal ;  frontal  shield  and  pos- 
terior part  of  nasals  rising  much  more  abruptly ;  frontals  much  more 
swollen  in  front  of  upper  part  of  orbits ;  point  of  shield  much  shorter 
(ending  midway  between  plane  of  postorbitals  and  fronto-parietal 
suture,  while  in  absarokus  it  reaches  posteriorly  to  suture) ;  inion 
short;  palate  broader;  underjaw  shorter;  coronoid  lower. 

Adult  male  compared  with  adult  male  tahltanicus  (both  inhabit- 
ing the  same  region)  :  Basal  length,  zygomatic  breadth,  and  frontal 
breadth  essentially  the  same,  but  frontal  region  much  higher  and 
rising  abruptly  at  orbits  instead  of  sloping  gently  in  plane  of  ros- 
trum; rostrum  broader,  flatter,  and  more  nearly  horizontal  (appear- 


90  NORTH   AMERICAN   FAUNA.  [No.  41. 

ing  shorter) ;  postorbitals  much  larger  and  less  horizontally  outstand- 
ing; palate  longer;  under  jaw  longer.     Dentition  heavier. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  321;  occi- 
pito-nasal  length,  305;  palatal  length,  171;  zygomatic  breadth,  217; 
interorbital  breadth,  84. 

URSUS  CRASSODON  sp.  kov. 
Big-Tooth  Grizzly. 

Type  No.  171049,  $  old,  U.  S.  National  Museum,  Biological  Sur- 
vey collection.  Collected  on  Klappan  Creek  (=Third  South  Fork 
Stikine  Eiver),  September,  1907,  by  Dr.  E.  P.  Richardson,  of  Boston. 

Cranial  characters. — Old  maZe/( type)  :  Frontal  shield  broad,  lyrate 
posteriorly,  the  point  reaching  or  slightly  passing  fronto-parietal 
suture;  shield  rising  abruptly  from  rostrum,  sulcate  anteriorly  but 
nearly  flat  between  postorbitals;  postorbitals  broadly  rounded  and 
horizontally  outstanding;  rostrum  of  medium  breadth,  nearly  hori- 
zontal ;  braincase  short ;  sinus  case  rising  to  support  postorbital  part 
of  shield;  zygomata  moderately  outstanding,  subtriangular,  the  squa- 
mosal root  not  broadly  expanded;  squamosal  shelf  narrow;  palate 
and  postpalatal  shelf  moderate;  ramus  short;  coronoid  blade  rather 
small,  moderately  recurved  at  apex;  canines  rather  large;  molars 
enormous  for  size  of  skull ;  MA  and  M2-  very  broad ;  heel  of  M-2-  sub- 
triangular,  narrowing  posteriorly. 

I  refer  to  this  species  two  young  males  from  Yukon,  one  (No. 
209896)  from  White  River,  the  other  (No.  1839,  Ottawa  Museum) 
from  Wolf  Lake  near  Teslin  Lake,  both  in  the  third  year;  and  an 
old  female  (No.  202792)  from  Tatletuey  Lake  on  the  upper  Finlay. 
Besides  these,  two  very  old  male  skulls  from  southern  Yukon  (No. 
223760  from  Glenlyon  Mountains  and  No.  223767  from  Quiet  Lake 
at  head  of  Big  Salmon  River)  are  provisionally  referred  to  the 
species,  but  not  without  considerable  hesitation.  Both  are  high, 
short,  and  broad  as  in  crassodon,  but  the  teeth  are  too  much  worn  to 
admit  of  reliable  comparison.  One  (No.  223760)  is  so  extremely 
aged  that  the  roots  of  the  canines  have  worked  down  and  become 
absorbed  basally,  leaving  the  middle  part  of  the  rostrum  narrower 
than  normal. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  320 ;  *  oc- 
cipito-nasal  length,  306 ; 1  palatal  length,  175 ;  zygomatic  breadth, 
222 ;  interorbital  breadth,  84. 

URSUS  CRASSUS  sp.  nov.1 
Thickset  Grizzly. 
Type  No.  225473,   $   ad.,  U.  S.  National  Museum,  Biological  Sur- 
vey collection.    From  upper  Macmillan  River,  Yukon.     Collected  in 
September,  1916,  by  William  Drury. 

1  Partly  restored. 

*  Tentatively  included  in  stikeenensis  group.      (See  Introduction,  pp.  12-13.) 


1918.]  STTKEENENSIS  GROUP.  91 

Characters. — Size  large;  hump  apparent  but  not  conspicuous; 
general  color  dark;  claws  of  medium  length,  narrow,  moderately 
curved,  smooth,  whitish  above,  with  dark  sides. 

Color. — Muzzle  pale  brownish;  top  of  head  and  neck  strongly 
washed  with  yellowish  or  golden  buffy;  shoulders  and  back  lightly 
tipped  with  same  on  dark  background ;  legs  and  feet  dusky. 

Cranial  characters. — Adult  male  (type)  :  Skull  rather  large,  short, 
broad,  rather  high,  but  not  highly  arched,  and  unusually  massive. 
Shield  broad,  convex,  faintly  sulcate,  sloping  gradually  into  rostrum, 
the  point  reaching  fronto-parietal  suture;  postorbitals  rather  broad, 
decurved  in  convexity  of  shield;  rostrum  large  and  high,  rising  in 
plane  of  shield;  sagittal  crest  short;  zygomata  moderate;  palate  nar- 
row and  strongly  troughed;  postpalatal  shelf  short;  underjaw  rather 
short,  massive ;  coronoid  blade  high  and  nearly  vertical ;  subangular 
border  short.  Teeth  large;  canines  massive;  Ma  very  long,  its  heel 
exceptionally  long  and  flat,  slightly  emarginate  and  everted. 

Cranial  comparisons. — The  skull  of  the  adult  male  (type)  viewed 
from  above  bears  a  striking  likeness  to  that  of  adult  male  hoots,  but 
when  turned  over  the  likeness  ceases.  It  agrees  with  hoots  essentially 
in  size,  massiveness,  arching  of  vault  of  cranium,  and  frontal  breadth ; 
but  frontal  shield  is  more  convex;  postorbitals  more  decurved;  ros- 
trum somewhat  more  ascending  (higher  posteriorly)  ;  orbital  rims 
less  swollen ;  palate  decidedly  narrower  and  troughed  instead  of  flat ; 
coronoid  more  nearly  vertical;  subangular  border  shorter;  teeth 
strikingly  larger  throughout — incisors,  canines,  and  molars.  The 
teeth  of  hoots  are  small  for  size  of  skull,  those  of  crassus  exception- 
ally large. 

Male  adult  (type)  compared  with  male  adult  crassodon  (type)  : 
Size  slightly  greater;  frontal  shield  much  broader,  more  gently  slop- 
ing, and  more  convex  transversely,  with  decurved  postorbitals  (in 
crassodon  shield  flattish,  nearly  horizontal,  with  horizontally  out- 
standing postorbitals  and  descending  abruptly  to  rostrum)  ;  rostrum 
very  much  larger  and  higher  and  sloping  gradually  into  shield; 
ramus  much  longer ;  subangular  border  shorter ;  coronoid  larger  and 
higher.  Upper  canines  about  same;  lower  canines  much  larger; 
upper  and  lower  molar  series  of  about  same  length  but  upper  molars 
much  narrower  and  less  massive. 

Remarks. — The  skull  of  an  immature  male  (No.  6552)  collected  by 
R.  MacFarlane,  May  1,  1863,  on  Anderson  River,  50  miles  southeast 
of  old  Fort  Anderson,  has  very  large  teeth,  especially  M-2-,  thus  dif- 
fering widely  from  any  adult  Barren  Ground  bear  thus  far  examined. 
Two  still  younger  skulls  collected  on  the  Barren  Grounds  in  1911, 
by  Dr.  R.  M.  Anderson  (No.  34411  Amer.  Mus.  Nat.  Hist,  from  Hor- 
ton  River,  and  34413  from  Langton  Bay),  also  have  the  crown  of  M* 
very  long.    These  three  skulls  I  provisionally  refer  to  crassus. 


92  NORTH  AMERICAN  FAUNA.  [No.  41. 

It  is  a  singular  fact,  in  view  of  the  wide  dissimilarity  of  the  skulls, 
that  the  teeth  of  male  crassus  and  male  kluane  are  very  much  alike. 

Skull  measurements. — Adult  male  (type):  Basal  length,  325; 
occipito-nasal  length,  322;  palatal  length,  171;  zygomatic  breadth, 
224;  interorbital  breadth,  94. 

URSUS  MIRABILIS  Merriam.* 
Strange  Grizzly. 

Ursus  mirabilis  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  p.  146,  September 
6,  1916. 

Type  locality. — Admiralty  Island,  Alaska. 

Type  specimen. — No.  137471,  ^  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.     Collected  June  26,  1905,  by  Cyrus  Catt. 

Characters. — A  true  grizzly,  of  medium  size,  related  to  stikeenensis 
of  the  mainland,  and  having  the  same  high  bulging  forehead;  exter- 
nal characters  unknown. 

Cranial  characters. — Adult  male  (type)  :  Skull  of  medium  size, 
short,  with  rather  broadly  spreading  zygomata  and  highly  arched 
(almost  domed)  frontal  region.  Frontal  shield  of  moderate  breadth, 
short  pointed  posteriorly,  rising  abruptly  from  rostrum,  convex  both 
antero-posteriorly  and  transversely,  slightly  sulcate  medially,  mod- 
erately swollen  on  each  side  of  sulcus ;  postorbital  processes  moderate, 
broader  than  peglike,  slightly  decurved  (continuing  convexity  of 
frontals)  ;  fronto-nasal  region  strongly  dished;  rostrum  high,  nar- 
row, strongly  sloping ;  zygomata  rather  broadly  outstanding,  slightly 
bowed;  palate  and  postpalatal  shelf  rather  broad;  under  jaw  short; 
ramus  bellied  posteriorly;  coronoid  blade  high  and  rather  vertical, 
the  apex  not  reaching  plane  of  condyle.  Upper  canines  rather  long; 
molars  rather  broad  and  short. 

Cranial  comparisons. — Ursus  mirabilis  requires  comparison  with 
only  a  single  species — stikeenensis  of  the  neighboring  mainland :  Size 
smaller ;  frontal  shield  narrower  and  more  bulging  anteriorly,  rising 
more  abruptly  from  rostrum;  rostrum  narrower,  materially  higher, 
and  more  sloping;  zygomata  more  widely  outstanding;  occipital 
overhang  more  pronounced;  palate  and  postpalatal  shelf  similar; 
underjaw  less  massive;  coronoid  blade  narrower  and  higher;  teeth 
similar,  but  heel  of  last  upper  molar  much  shorter. 

Skull  measurements. — Adult  male  (type):  Basal  length,  308; 
occipito-nasal  length,  310;  palatal  length,  168;  zygomatic  breadth, 
230;  interorbital  breadth,  81. 

*  Tentatively  included  in  stikeenensis  group.      (See  Introduction,  pp.  12-13.) 


1918.]  STTKEENENSIS  GROUP.  93 

URSUS  ABSAROKUS  Merbiam.1 

Absaroka   Grizzly. 

Ursus  absarokus  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p  181,  August 
13,  1914. 

Type  locality. — Near  head  of  Little  Bighorn  River,  northern  part 
of  Bighorn  Mountains,  Montana. 

Type  specimen. — No.  67391,  $  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  in  May,  1893;  purchased  for 
Biological  Survey  by  J.  Alden  Loring. 

Range. — Laramie  and  Bighorn  Mountains,  eastern  Wyoming, 
Black  Hills  region,  South  Dakota,  and  northward  along  Little  Mis- 
souri to  Missouri  and  Yellowstone  Rivers. 

Characters. — Size  large,  but  much  less  than  horribilis  and  with 
much  smaller  molars — especially  M^. 

Color. — Head  of  young-adult  (No.  203524)  killed  by  Howard 
Eaton  on  the  Little  Missouri  at  mouth  of  Bear  Creek,  near  Middle  or 
"  Bullion "  Butte,  October  27,  1880 :  Muzzle  pale  brown,  changing 
to  grizzled  dark  brown  on  head  and  face;  a  large  patch  of  dark 
brown  free  from  grizzling  on  side  of  face  extending  from  eye  to 
angle  of  jaw ;  chin  and  gular  region  dark  brown  (except  anterior  part 
of  chin,  which  has  not  yet  molted  the  pale  old  coat) ;  top  and  sides 
of  neck  and  doubtless  body  also,  strongly  grizzled. 

Cranial  characters. — Adult  male  (type  specimen,  and  other  males 
from  Bighorn  and  Laramie  Mountains)  :  Vault  of  cranium  rather 
highly  arched;  zygomata  moderately  outstanding  and  slightly 
bowed;  frontal  shield  rather  broad,  rising  strongly  from  rostrum, 
convex  both  antero-posteriorly  and  transversely ;  slightly  swollen  on 
each  side  of  median  line  just  behind  plane  of  orbits,  faintly  de- 
pressed medially  between  orbits;  postorbital  processes  large,  out- 
standing, and  slightly  decurved,  the  tips  bluntly  rounded;  sagittal 
crest  nearly  straight,  reaching  anteriorly  to  fronto-parietal  suture; 
rostrum  large  and  high,  rising  strongly  into  frontal  shield;  post- 
palatal  shelf  broad  and  flat;  nasals  large  and  long;  top  of  coronoid 
high  and  broadly  rounded,  its  apex  short,  not  reaching  posteriorly  to 
plane  of  condyle.  Teeth  rather  large,  especially  M-2-  which  is  long 
and  broad,  the  heel  emarginate  on  outer  side. 

Cranial  comparisons. — Adult  male  (type)  compared  with  adult 
male  shoshone  (type)  :  Basal  length  essentially  the  same  but  skull 
somewhat  larger  and  more  massive,  broader,  highest  point  more 

1  Tentatively  included  in  stiheenensis  group.     (See  Introduction,  pp.  12-13.) 


94  NORTH   AMERICAN   FAUNA.  [No.  41. 

anterior;  frontal  shield  broader,  domed  (strongly  convex  both  an- 
tero-posteriorly  and  transversely)  and  rising  rather  abruptly  from 
plane  of  rostrum  (in  shoshone  flat)  ;  point  of  shield  much  longer  pos- 
teriorly, reaching  to  or  nearly  to  fronto-parietal  suture.  Molars 
larger — especially  MA 

Skull  measurements. — Adult  male  (type):  Basal  length,  322; 
occipito-nasal  length,  322;  palatal  length,  172;  zygomatic  breadth, 
218;  interorbital  breadth,  88. 

Alascensis  Group. 

URSUS  ALASCENSIS  Merriam. 

/ 
Alaska  Grizzly. 

(PI.  XII.) 

Ursus  horribilis  alascensis  Merriam,  Proc.  Biol.  Soc.  Washington,  X,  pp.  74-75, 
April  13,  1896. 

Type  locality. — Unalaklik  River,  Alaska. 

Type  specimen.1 — No.  76466,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  in  1895,  by  the  late  Rudolf 
Neumann,  of  Iliuliuk,  Alaska. 

Range. — Norton  Sound  region,  Alaska  (Unalaklik  and  Shaktolik 
Hills)  southerly  over  the  Nushagak  and  Kuskokwim  Rivers  to 
Chinitna  on  Cook  Inlet.    Limits  unknown. 

Cranial  characters. — Old  male  (type)  :  Size  small;  braincase  broad 
anteriorly;  frontal  shield  rather  broad,  flattish,  very  short  pointed 
posteriorly  with  correspondingly  elongate  sagittal  crest,  moderately 
sulcate  interorbitally,  otherwise  convex  in  cross  section,  rising  rather 
strongly  from  facial  plane;  postorbitals  rather  small,  moderately 
decurved;  fronto-nasal  region  moderately  dished;  zygomata  sub- 
triangular,  not  broadly  outstanding;  palate  rather  broad  and  con- 
cave; postpalatal  shelf  broad  and  short;  notch  rather  broad;  jaw 
rather  long;  inferior  border  of  ramus  long,  moderately  convex 
posteriorly ;  coronoid  blade  moderate,  rather  high,  the  apex  recurved. 
Teeth  moderate;  last  upper  molar  broad  and  short,  with  short 
obliquely  truncate  heel ;  first  lower  molar  sinuous,  a  strong  concavity 
on  outer  side. 

Adult  female:2  Long  and  high;  vault  of  cranium  well  arched, 
highest  about  midway  of  frontals;  frontal  shield  rather  narrow,  not 
flattened  but  arching  high  above  facial  plane;  swollen  between  me- 

1  In  describing  this  bear  20  years  ago  I  neglected  to  designate  a  type,  and  the  original 
material  included  skulls  of  more  than  one  species.  I  take  this  opportunity  therefore  to 
fix  the  type  of  Ursus  alascensis  on  one  of  the  original  specimens  (No.  76466,  U.  S.  Na- 
tional Museum,  Biological  Survey  collection)  and  to  redefine  the  species. 

2  In  the  absence  of  skulls  of  adult  females  from  the  type  region  it  is  assumed  that 
females  from  the  Nushagak,  Kuskokwim,  and  Chinitna  Rivers  are  fairly  typical  of 
alascensis. 


1918.]  ALASCENSIS  GROUP.  95 

dian  sulcus  and  orbits;  postorbital  processes  decurved;  palate  and 
postpalatal  shelf  long.    Last  upper  molar  with  short  heel. 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
tohlat  (No.  158811,  collected  by  Charles  Sheldon  at  head  of  Toklat 
River  near  northern  base  of  Mount  McKinley)  :  Size  and  general 
appearance  essentially  the  same,  but  vault  of  cranium  less  elevated ; 
frontal  shield  narrower  and  shorter  (the  point  reaching  about  half- 
way from  plane  of  postorbitals  to  parietals) ;  sagittal  crest  longer ; 
zygomata  much  less  broadly  outstanding,  less  arched;  underjaw  de- 
cidedly longer;  inferior  border  of  ramus  longer;  coronoid  decidedly 
higher ;  heel  of  last  upper  molar  very  much  shorter. 

Adult  female  compared  with  adult  female  toklat:  Skull  decidedly 
larger;  vault  of  cranium  much  higher  and  more  arched  (in  tohlat 
low  and  flat),  highest  about  middle  of  frontals  instead  of  at  hinder 
end;  rostrum  larger  (longer,  broader,  and  higher)  ;  face  more  slop- 
ing (nasals  less  nearly  horizontal) ;  frontals  much  more  swollen  be- 
tween sulcus  and  orbits ;  postorbital  processes  more  decurved ;  palate, 
postpalatal  shelf,  and  occipito-sphenoid  notably  longer;  last  upper 
molar  very  much  shorter. 

Adult  male  compared  with  adult  male  tundrensis  of  same  region : 
Size  smaller  (basilar  and  occipito -nasal  lengths  fully  an  inch  less)  ; 
skull  less  massive;  frontals  narrower,  rising  more  abruptly  from 
facial  plane  and  more  swollen  over  orbits;  rostrum  more  slender; 
palate  less  flattened ;  ramus  of  jaw  less  swollen  posteriorly ;  coronoid 
less  falcate;  canines  smaller,  decidedly  shorter,  and  more  curved; 
molars  decidedly  smaller. 

Skull  measurements — Old  male  (type) :  Basal  length,  310;  oc- 
cipito-nasal  length,  298.5;  palatal  length,  166;  zygomatic  breadth, 
206;  interorbital  breadth,  79. 

URSUS  TOKLAT  Meebiam. 

Toklat  Gbizzly. 

Ursus   toklat   Merriam,   Proc   Biol.    Soc.   Washington,   XXVII,   pp.   182-183, 
August  18,  1914. 

Type  locality. — Head  of  Toklat  River,  north  base  of  Alaska  Range, 
near  Mount  McKinley,  Alaska. 

Type  specimen. — No.  158813,  9  ad.,  U.  S.  National  Museum, 
Biological  Survey  collection.  Collected  May  24,  1908,  by  Charles 
Sheldon,  and  by  him  presented  to  the  Biological  Survey;  original 
No.  324.     Mother  of  cub  No.  158814  (original  No.  325). 

Range. — So  far  as  known,  restricted  to  Alaska  Range. 

Characters. — Size  medium;  skulls  of  both  male  and  female  rather 
highly  arched  and  dished,  that  of  female  much  smaller  and  lower 
than  male.  Last  upper  molar  large;  heel  very  long.  Claws  horny 
and  smooth. 

64854°— 18 7 


96  NORTH   AMERICAN   FAUNA.  [No.  41. 

Color. — Color  variable,  upperparts  ranging  from  ordinary 
"  grizzly  bear  color  "  to  creamy  white ;  claws  usually  dark  through- 
out, but  in  many  cases  more  or  less  marked  with  white. 

Cranial  characters. — Adult  males:  Size  medium;  rostrum  high, 
subterete  above,  nearly  horizontal;  frontal  shield  abruptly  elevated, 
convex  or  domed,  swollen  over  orbits,  sulcate  medially,  rather  long 
pointed  (reaching  fronto-parietal  suture)  ;  postorbital  processes 
rather  small  and  strongly  decurved;  squamosal  shelves  short;  palate 
arched  and  excavated  longitudinally;  postpalatal  shelf  rather  broad; 
zygomata  very  broadly  spreading  and  angular  (zygomatic  breadth 
in  proportion  to  basal  length  much  greater  than  in  any  other  true 
grizzly,  about  equaling  that  of  sheldoni)  ;  sagittal  crest  short.  Last 
upper  molar  large,  its  heel  normally  very  long.  Adult  female:  Skull 
rather  long  and  narrow,  with  broadly  spreading  zygomata  and 
strongly  dished  f ronto-nasal  region ;  frontal  shield  rather  flat,  sulcate 
between  orbits,  varying  from  lyre  pointed  to  short  pointed;  post- 
orbital  processes  outstanding  horizontally;  rostrum  rather  slender, 
nearly  horizontal.  In  most  specimens  the  highest  point  of  cranium 
culminates  in  a  rather  abrupt  change  of  angle  at  or  near  the  fronto- 
parietal suture,  forming  a  sort  of  "  hump,"  a  condition  usual  also  in 
skulls  of  female  grizzlies  from  the  upper  Yukon  and  northern 
British  Columbia. 

Cranial  comparisons. — Old  male  compared  with  old  male  alascen- 
sis:  Frontal  region  more  elevated;  sagittal  crest  shorter;  zygomatic 
breadth  much  greater ;  heel  of  last  upper  molar  much  longer. 

Adult  female  compared  with  female  alascensis:  Size  smaller; 
vault  of  cranium  decidedly  lower,  highest  over  posterior  part  of 
frontals  instead  of  over  middle  of  frontals;  frontal  shield  narrow, 
flattened,  sloping  (not  arched  and  not  materially  swollen  over  or- 
bits);  postorbital  processes  outstanding  horizontally;  palate,  post- 
palatal  shelf,  and  occipito-sphenoid  shorter;  M>  with  heel  much 
longer. 

Adult  female  compared  with  adult  female  phmonyx:  Size  de- 
cidedly greater;  teeth  much  larger,  especially  the  canines  and  last 
upper  molars. 

Adult  female  compared  with  adult  female  kluane:  Length  much 
greater;  facial  part  of  skull  about  the  same;  braincase  and  post- 
palatal  parts  much  longer;  vault  of  cranium  higher;  sagittal  crest 
longer  and  higher,  reaching  forward  over  posterior  frontals;  point 
of  shield  much  shorter;  underjaw  and  inferior  border  of  ramus  much 
longer ;  teeth  about  the  same  size  but  M1  usually  smaller ;  heel  of  M-2- 
longer. 

Skull  measurements. — Old  male  (No.  158811,  from  head  Toklat 
River,  Alaska) :  Basal  length,  304;  occipito-nasal  length,  298;  palatal 


1918.]  AXASCENSIS  GBOUP.  97 

length,  169.5 ;  zygomatic  breadth,  222 ;  interorbital  breadth,  80.  Old 
female  (type) :  Basal  length,  283;  occipito-nasal  length,  267;  palatal 
length,  157;  zygomatic  breadth,  187;  interorbital  breadth,  73. 

URSUS  LATIPRONS  Mebeiam. 

Bboad-Fbonted  Gbizzlt. 

Ursus  phcBonyx  latifrons  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp. 
183-184,  August  13,  1914. 

Type  locality. — Jasper  House,  Alberta. 

Type  specimen. — No.  75612,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  September  15,  1895,  by  J.  Alden 
Loring.     Original  No.  3270. 

Range. — Rocky  Mountains  of  western  Alberta  and  eastern  British 
Columbia  from  Jasper  House  northwesterly  to  region  between  head- 
waters of  Parsnip  and  Great  Bend  of  Fraser  River  and  thence  to 
extreme  headwaters  of  Stikine  River;  limits  of  range  unknown. 

Characters. — Size  medium  or  rather  large.  External  characters 
unknown.    Affinities  with  the  geographically  remote  toklat. 

Cranial  characters. — Adult  male  (type)  :  Size  medium;  base  elon- 
gate posteriorly;  vault  of  cranium  rather  low,  only  slightly  arched, 
top  flattish;  frontal  shield  very  broad  (interorbitally  93  mm.,  be- 
tween tips  of  postorbitals  130),  rising  from  rostrum  rather  abruptly, 
broadly  but  shallowly  depressed  medially,  swollen  over  orbits,  the 
point  ending  in  sagittal  crest  some  distance  (apparently  more  than 
an  inch)  anterior  to  fronto-parietal  suture;  postorbital  processes 
broad,  outstanding,  and  only  slightly  depressed ;  f ronto-nasal  region 
slightly  dished;  rostrum  rather  high,  nearly  horizontal,  rounded 
above;  postpalatal  shelf  rather  broad;  notch  rather  narrow;  zygo- 
mata strongly  outbowed  but  not  widely  spreading;  under  jaw  long; 
coronoid  blade  broad  basally,  rather  low,  its  anterior  border  sloping 
strongly  backward.  Teeth  too  badly  worn  to  admit  of  description. 
Adult  female  (No.  209378)  collected  by  F.  K.  Vreeland,  September 
17,  1915,  near  head  of  Big  Salmon  or  North  Fork  Fraser  River  be- 
tween Big  Bend  of  the  Fraser  and  headwaters  of  the  Parsnip :  Simi- 
lar in  general  to  that  of  male  with  the  usual  sexual  difference  in 
frontal  shield,  which  is  lyrate  pointed  posteriorly;  frontal  shield 
broad — broadly  depressed  medially  between  orbits,  swollen  over  or- 
bits; postorbital  processes  strongly  developed,  outstanding,  slightly 
decurved ;  sagittal  crest  short,  high  posteriorly,  covering  about  three- 
quarters  of  suture  between  parietals;  postpalatal  shelf  rather  broad 
and  flat;  mastoids  short  and  appressed;  postpalatal  notch  short,  of 
moderate  breadth;  zygomata  moderately  outstanding,  less  bowed 
than  in  the  male;  coronoid  blade  broad  basally,  rather  low,  recurved 


98  NORTH   AMERICAN   FAUNA.  [No.  41 

at  apex;  canines  of  medium  size;  molars  large,  especially  last  upper 
molar. 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
toklat  (from  type  locality)  :  Frontal  shield  much  broader  and  less 
elevated;  braincase  less  arched;  palate  flatter  (in  toklat  distinctly 
arched  antero-posteriorly) ;  zygomata  more  rounded  basally  and  more 
outbowed  (less  angular) ;  underjaw  slightly  longer;  inferior  border 
of  ramus  much  longer ;  coronoid  blade  broader  and  higher.  The  basal 
part  of  the  skull  is  longer  in  latifrons  than  in  toklat.  This  is  apparent 
whether  the  skull  is  viewed  from  above  or  below.  In  latifrons,  seen 
from  above,  the  distance  from  occipital  crest  to  postorbital  process  is 
materially  greater ;  while  seen  from  bejow,  the  distance  from  occipital 
condyle  to  postpalatal  notch  is  likewise  greater.  Another  difference 
appears  rather  strikingly  when  the  skull  is  viewed  from  below:  in 
latifrons  the  squamosal  base  of  the  zygoma  slopes  gradually  forward, 
while  in  toklat  the  curve  is  much  shorter  and  more  abrupt,  so  that 
the  bases  of  the  zygomata  stand  out  much  more  squarely,  practically 
at  right  angles  to  cranial  axis. 

While  skulls  of  adult  male  latifrons  differ  strikingly  from  those 
of  male  toklat,  skulls  of  the  adult  females  are  surprisingly  alike, 
agreeing  in  general  appearance,  basal  length,  breadth  of  palate,  and 
large  size  of  last  upper  molar,  though  in  latifrons  this  tooth  is  ac- 
tually broader  than  in  most  specimens  of  toklat.  The  two  agree 
essentially  also  in  lower  molars  and  canines.  The  underjaw  in  lati- 
frons, however,  is  shorter,  and  coronoid  blade  lower  than  in  toklat. 

Old  male  compared  with  old  male  kluane:  Frontal  region  much 
broader  and  flatter ;  postorbital  processes  much  less  decurved ;  sagit- 
tal crest  shorter  and  straight  instead  of  arched;  rostrum  less  ele- 
vated and  less  narrowly  rounded  above;  nasals  flatter;  zygomata 
more  widely  spreading  and  more  strongly  outbowed  posteriorly; 
coronoid  blade  broader  at  base. 

Adult  female  compared  with  adult  female  kluane:  Skull  much 
larger,  longer,  and  more  massive;  frontal  shield  irmch  broader; 
vault  of  cranium  flatter  (less  arched  over  posterior  frontals) ;  ros- 
trum higher;  zygomata  more  widely  outstanding;  palate  much 
broader;  last  upper  molar  much  larger:  lower  molars  and  canines 
approximately  same  size.  While  the  skull  of  female  latifrons  is 
much  larger  than  that  of  female  kluane,  the  underjaw  is  only  slightly 
larger. 

Adult  female  compared  with  adult  female  phwonyx:  Size  much 
larger;  frontal  region  much  broader;  rostrum  broader;  nasals  more 
nearly  horizontal;  arch  of  cranium  more  depressed ;  underjaw  longer; 
coronoid  blade  broader;  molars  much  larger.  (Comparison  of  males 
is  unnecessary,  the  male  of  phmonyx  having  a  large,  broad,  massive 
skull  resembling  that  of  daMi.) 


1918.]  RICHARDSONI  GROUP.  99 

Skull  measurements. — Old  male  (type)  :  Basal  length,  324;  oceip- 
ito-nasal  length,  312;  palatal  length,  173;  zygomatic  breadth,  214.5; 
interorbital  breadth,  94. 

Richardsoni  Group. 

URSUS  RICHARDSONI  Swainson. 

Babren  Ground  Beab. 

Vrsus  Richardsoni  Swainson,  Animals  in  Menageries,  pp.  54-56,  1838. 

Type  locality. — Shore  of  Arctic  Ocean,  on  west  side  of  Bathurst 
Inlet  near  mouth  of  Hood  River.1 

Characters. — Size  medium;  color  variable,  from  yellowish  to 
grizzly  brown ;  f oreclaws  of  medium  length,  smooth ;  skull  medium 
or  small,  with  broadly  spreading  zygomata. 

Cranial  characters. — Adult  males:  Two  adult  or  rather  old  male 
skulls  collected  by  Dr.  R.  M.  Anderson  in  1915,  and  loaned  me  by  the 
Museum  of  the  Geological  Survey  of  Canada,  at  Ottawa  (one  No. 
2774  from  Dolphin  and  Union  Straits,  the  other  No.  2776  from 
near  mouth  of  Coppermine  River),  are  assumed  to  be  typical:  Size 
medium ;  cranium  high  in  relation  to  size,  but  not  much  arched ;  basi- 
cranium  flat;  shield  rising  abruptly  from  rostrum,  of  medium 
breadth,  broadly  and  strongly  sulcate  medially,  swollen  over  orbits, 
short  pointed;  postorbital  processes  massive,  outstanding,  arched 
over  orbits  and  slightly  decurved;  orbits  nearly  vertical;  rostrum 
moderate,  rather  high  and  narrow,  sloping  into  sulcus  of  shield; 
orbits  prominent,  rising  well  above  fronto-nasal  plane;  zygomata 
moderate,  somewhat  outbowed;  nares  subrectangular,  broader  than 
high ;  palate  rather  short ;  postpalatal  shelf  moderate  or  rather  broad ; 
squamosal  shelf  weak  posteriorly;  underjaw  rather  short;  infra- 
angular  border  of  ramus  short;  apex  of  coronoid  only  slightly  re- 
curved, falling  short  of  plane  of  condyle;  angular  process  projecting 
beyond  condyle.  Teeth  medium ;  canines  rather  large ;  M-2-  with  mod- 
erate heel,  narrowing  posteriorly ;  cusps  of  posterior  molars  weak, those 
of  inner  side  of  M-2-  nearly  obsolete  (in  striking  contrast  with  the 
highly  developed  cusps  of  both  upper  and  lower  molars  of  pellyensis) . 

An  exceedingly  old  male  in  the  National  Museum  (No.  6255), 
collected  on  Anderson  River  by  R.  MacFarlane,  has  the  shield  less 
swollen-  over  and  in  front  of  orbits,  and  even  shorter  pointed  poste- 

1  Swainson's  description  was  based  on  and  largely  quoted  from  Richardson's  account  of 
"  an  old  and  lean  male,  killed  on  the  shores  of  the  Arctic  Sea  on  the  1st  of  August,  1821.'' 
(Article  on  the  Barren  Ground  Bear,  in  Fauna  Boreali-Americana,  pp.  21-24,  1829.)  And 
in  Franklin's  "  Narrative  of  a  Journey  to  the  Shores  of  the  Polar  Sea,"  p.  373,  1823, 
under  date  of  August  1,  1821,  the  party  being  at  the  mouth  of  Hood  River  on  Bathurst 
Inlet,  the  killing  of  a  lean  male  brown  bear  is  chronicled  in  some  detail.  It  appears 
therefore  that  the  type  locality  of  Vrsus  richardsoni  is  Hood  River,  Bathurst  Inlet. 


100  NORTH   AMERICAN   FAUNA.  [No.  41. 

riorly;  rostrum  more  depressed;  sagittal  crest  longer  and  somewhat 
higher  (but  still  not  high)  and  somewhat  humped  over  posterior 
frontals;  zygomata  more  broadly  outstanding. 

SkiiM  measurements. — Old  male  (No.  6255)  from  Anderson  River: 
Basal  length,  311;  occipito-nasal  length,  286;  palatal  length,  163; 
zygomatic  breadth,  227;  interorbital  breadth,  72.5.  Old  male  (No. 
2774,  Ottawa  Museum)  from  Dolphin  and  Union  Strait:  Basal 
length,  316;  occipito-nasal  length,  290;  palatal  length,  170;  zygo- 
matic breadth,  211;  interorbital  breadth,  79.  Adult  male  (No.  2776 
Ottawa  Museum)  from  near  mouth  Coppermine  River:  Basal  length, 
299;    occipito-nasal   length,    288;    palatal    length,    163;    zygomatic 

breadth,  217 ;  interorbital  breadth,  82.5. 

/ 

URSUS  RUSSELLI  *  Mebbiam.* 
Mackenzie  Delta  Gbizzly. 

Ursus  russelli  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  178,  August  13, 
1914. 

Type  locality. — West  side  Mackenzie  River  delta,  Canada. 

Type  specimen. — No.  21301,  $  old,  University  of  Iowa  Museum. 
Collected  June  28,  1894,  by  Frank  Russell.  Mounted  skin  with 
skull  separate. 

Range. — Lower  Mackenzie  region ;  limits  unknown. 

Characters. — Size  rather  small.  Color  a  curious  pale  drab-brown, 
somewhat  darker  on  legs  and  feet;  ears  conspicuously  hairy.  Claws 
smooth,  moderately  to  strongly  curved;  brownish  horn  color  with 
paler  (almost  amber)  tips.  Teeth  large;  last  upper  molar  of  great 
size  and  peculiar  glassy  texture  in  type  skull.    Affinities  uncertain. 

Cranial  characters. — Adult  male  (type)  :  Skull  of  medium  size, 
about  equaling  old  male  of  alascensis;  rather  short;  fairly  broad 
across  zygomata ;  frontal  shield  sloping  strongly  upward,  moderately 
sulcate,  swollen  over  orbits,  but  orbits  not  everted;  posterior  point 
of  shield  rather  short,  ending  about  one-third  the  distance  from 
fronto-parietal  suture  to  postorbital  processes;  postorbital  processes 
peglike,  standing  out  nearly  horizontally — not  depressed  as  in 
alascensis;  muzzle  rather  narrow  and  high;  zygomata  slender,  the 
posterior  roots  not  expanded  vertically ;  palate  flat,  not  excavated  or 
arched  as  in  several  species;  underjaw  massive,  heavier  under  Mj 
and  My  than  in  alascensis;  coronoid  blade  falcate  but  not  nar- 
rowly so. 

Cranial  comparisons. — From  richardsoni,  its  neighbor  on  the  east, 
with  which  it  agrees  in  size  and  in  certain  dental  characters,  it  differs 
in  much  more  highly  vaulted  cranium;  more  highly  sloping   (less 

1  Named  for  Frank  Russell,  who  collected  the  type  specimen. 

•  Tentatively  included  In  riohardaoni  group.      (See  Introduction,  pp.  12-18.) 


1918.]  EICHAEDSONI  GROUP.  101 

nearly  horizontal)  braincase;  much  more  elevated  frontal  region,  and 
very  much  narrower  rostrum.  The  frontal  shield  is  much  longer  than 
in  richardsoni,  the  temporal  impressions  curving  backward  to  meet 
one  another  about  one-third  the  distance  between  postorbital  processes 
and  f ronto-parietal  suture,  instead  of  turning  abruptly  inward ;  post- 
orbital  processes  more  slender  than  in  richardsoni/  sagittal  crest 
shorter  and  less  nearly  horizontal;  upper  molars  very  much  larger, 
particularly  broader.  Zygomatic  arches  bowed  outward  in  both  spe- 
cies— not  sharply  angular  as  in  many  large  bears. 

Skull  measurements. — Old  male  (type):  Basal  length,  310;  oc- 
cipito-nasal  length,  300 ;  palatal  length,  163 ;  zygomatic  breadth,  220 ; 
interorbital  breadth,  79.5. 

URSUS  PHJEONYX  Mebeiam. 

Tanana  Geizzly. 

Ursus  Jwrribilis  pkceonyx  Merriam,  Proc.  Biol.  Soc.  Washington,  XVII,  p.  154. 
October  6,  1904. 

Type  locality. — Glacier  Mountain,  Tanana  Mountains,  Alaska 
(about  2  miles  below  source  of  Comet  Creek,  near  Fortymile  Creek, 
between  Yukon  and  Tanana  Rivers). 

Type  specimen. — No.  133231,  2  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.    Collected  July  12,  1903,  by  W.  H.  Osgood. 

Range. — Tanana  Mountains  between  Tanana  and  Yukon  Rivers. 

Characters. — Size  of  male  large;  of  female  small  (sexual  dis- 
parity great,  much  greater  than  in  datti). 

Color. — Upperparts  varying  from  creamy  or  buffy  to  dark 
"  grizzly  color  " ;  underparts  and  muzzle  pale  brown ;  legs  very  dark 
brown,  varying  to  blackish  brown;  claws  horny  and  smooth,  usually 
dark  but  sometimes  marked  with  whitish.  Last  upper  molar  of  me- 
dium size  or  rather  small. 

Cranial  characters. — Old  male  (No.  201586,  from  Ketchumstock, 
assumed  to  be  typical  of  phwonyx) :  Size  medium  or  rather  large; 
skull  broad  and  short,  moderately  arched  and  dished,  with  broadly 
outstanding  zygomata  and  rather  deeply  sulcate  strongly  sloping 
frontal  shield.  Frontal  shield  of  moderate  breadth,  strongly  swollen 
over  orbits  and  bases  of  postorbitals,  short  pointed  posteriorly,  the 
point  ending  midway  between  fronto-parietal  suture  and  plane  of 
postorbitals;  postorbitals  triangular,  broad  basally,  convex  poste- 
riorly, straight  or  concave  anteriorly;  rostrum  moderate,  nearly 
horizontal,  sulcate- depressed  on  top;  fronto-nasal  region  distinctly 
dished ;  squamosal  shelves  broad  and  rather  short,  the  outer  margin 
arched  and  upturned;  middle  part  of  zygomata  moderately  ex- 
panded vertically;  palate  broad  and  short;  postpalatal  shelf  broad; 
notch  relatively  narrow.    Under  jaw  long  for  size  of  skull,  massive; 


102  NORTH   AMERICAN   FAUNA.  [No.  41. 

coronoid  blade  high,  rather  narrow,  and  rather  vertical,  the  recurved 
apex  not  reaching  plane  of  condyle.    Teeth  medium. 

Adult  female  (type):  Size  small;  vault  of  cranium  well  arched 
posteriorly,  highest  point  25  to  30  mm.  in  front  of  fronto-parietal 
suture;  braincase  rather  broad,  not  much  constricted  anteriorly, 
not  compressed,  and  with  no  tendency  to  keel  into  anterior  part  of 
sagittal  crest;  frontal  shield  moderately  flattened,  shallowly  sulcate 
medially  and  slightly  swollen  on  each  side  of  median  depression, 
gently  sloping  into  rostrum  posteriorly,  the  lyrate  point  reaching 
nearly  to  parietals ;  rostrum  rather  small ;  zygomata  moderately  out- 
standing; palate  medium;  postpalatal  shelf  rather  broad  and  short. 
Under  jaw  long;  inferior  border  of  ramus  long  and  straight;  coro- 
noids  rather  small,  the  apex  strongly  recurved,  cutting  plane  of 
condyle. 

Cranial  comparisons. — Old  female  (type)  compared  with  adult 
female  kluane:  Size  somewhat  larger;  braincase  broader  anteriorly; 
frontal  shield  broader;  palate  and  postpalatal  shelf  broader;  oc- 
cipito-sphenoid  and  postpalatal  lengths  much  greater.  Under  jaw 
much  longer ;  inferior  border  of  ramus  longer  and  straighter.  Teeth 
too  badly  worn  to  admit  of  description  but  apparently  about  same 
size  as  in  kluane. 

Old  female  (type)  compared  with  adult  female  toklat  from  Alaska 
Range,  near  Mount  McKinley:  Basal  length  somewhat  less;  brain- 
case less  contracted  and  much  broader  anteriorly  (not  compressed 
and  with  no  tendency  to  keel  into  anterior  part  of  sagittal  crest) ; 
frontal  shield  less  deeply  sulcate  anteriorly  and  much  longer  pointed 
posteriorly;  fronto-nasal  region  less  depressed;  under  jaw  shorter; 
ramus  straighter  and  lighter;  coronoid  much  smaller,  narrower,  and 
lower;  upper  canines  apparently  about  the  same;  last  upper  molar 
smaller ;  lower  canines  more  slender ;  lower  molars  apparently  about 
the  same,  but  so  completely  worn  off  in  type  specimen  that  compari- 
son is  impossible. 

Remarks. — The  skull  of  the  adult  male  phmonyx  (if  the  sexes  are 
correctly  mated)  is  large,  broad,  and  rather  massive,  requiring  com- 
parison with  only  a  single  known  species,  Ursus  dalli.  The  female 
on  the  other  hand  is  small  and  resembles  in  a  general  way  the 
females  of  the  still  smaller  kluane,  pulchellus,  and  pallasi. 

Skull  measurements. — Old  male  (No.  205186,  from  Ketchumstock, 
Alaska)  :  Basal  length,  327;  occipito-nasal  length,  309 ;  palatal  length, 
176;  zygomatic  breadth,  242;  interorbital  breadth,  85.  Old  female 
(type) :  Basal  length,  280;  occipito-nasal  length,  267;  palatal  length, 
148;  zygomatic  breadth,  189;  interorbital  breadth,  68. 


1918.]  RICHARDSONI  GROUP.  103 

URSUS  INTERNATIONALIS  Meeeiam. 

AXASKA  BOUNDAEY  GbIZZLT. 

Urstts  internationalis  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  177- 
178,  August  13,  1914. 

Type  locality. — Alaska- Yukon  Boundary,  about  50  miles  south  of 
Arctic  coast  (lat.  69°  00'  30"). 

Type  specimen. — No.  1763,  $  ad.,  Ottawa  Museum.  Killed  July  3, 
1912,  by  Frederick  Lambart,  of  Canadian  Boundary  Survey. 

Range. — Eegion  bordering  Arctic  coast  along  international  bound- 
ary, and  doubtless  adjacent  mountains,  between  the  coast  and  the 
Yukon-Porcupine;  limits  unknown. 

Characters. — Size  medium  or  rather  large;  affinities  doubtful. 
Color  a  peculiar  pale  yellowish  brown.  Head  strongly  arched ;  muz- 
zle and  frontal  region  broad.  Large  lower  premolar  strictly  conical, 
without  heel,  as  in  the  brown  bears. 

Cranial  characters. — Skull  of  medium  size,  massive,  strongly 
arched  and  dished,  highest  over  anterior  part  of  braincase;  frontal 
shield  broad,  very  short  pointed  posteriorly,  sulcate  medially  and 
swollen  over  orbits ;  postorbitals  bluntly  rounded,  strongly  decurved, 
not  widely  projecting;  fronto-nasal  region  strongly  dished;  rostrum 
large  and  broad;  sagittal  crest  long  but  feebly  developed;  zygomata 
subtriangular,  not  widely  outstanding,  and  not  much  expanded  ver- 
tically; palate  and  postpalatal  shelf  rather  broad;  notch  moderate. 
Teeth  rather  small  for  size  of  skull;  heel  of  last  upper  molar  small 
and  obliquely  truncate  on  outer  side;  large  lower  premolar  strictly 
of  brown-bear  type — a  single  cone  without  heel,  sulcus,  or  posterior 
cusplets;  first  lower  molar  broad  and  somewhat  sinuous;  middle 
lower  molar  narrow  and  short  posteriorly. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  309;  oc- 
cipito-nasal  length,  293;  palatal  length,  169;  zygomatic  breadth, 
203.5;  interorbital  breadth,  82. 

URSUS  OPHRUS1  Meebiam. 

High-Bbow  Grizzly. 
(Pl.X.) 

Ur8U8  ophrus  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  148-149,  Sep- 
tember 6,  1916. 

Type  locality. — Eastern  British  Columbia  (exact  locality  un- 
known). 

Type  specimen. — No.  210252,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.    Collected  in  1915,  by  E.  W.  Darbey. 

1  Ophrus,  with  reference  to  the  unusual  brows. 


104  NORTH   AMERICAN   FAUNA.  [Mo.  41. 

Cranial  characters. — Skull  short,  strongly  dished,  remarkably  high, 
the  deeply  sulcate  frontal  shield  rising  abruptly  high  over  orbits, 
with  thickened  brows  and  large  outstanding  arched  postorbital  proc- 
esses. Frontal  shield  of  moderate  breadth ;  deeply  and  broadly  con- 
cave between  orbits,  swollen  over  orbits  and  passing  out  into  strongly 
outstanding  postorbitals,  short  pointed  posteriorly;  fronto-nasal 
region  deeply  sulcate;  middle  part  of  nasals  flat;  sagittal  crest  high 
and  reaching  anteriorly  nearly  midway  from  fronto-parietal  suture 
to  plane  of  postorbitals;  rostrum  rather  small  and  narrow;  palate 
rather  narrow;  postpalatal  shelf  rather  broad;  zygomata  broadly 
and  strongly  outbowed;  mastoids  rather  long.  Under  jaw  long,  its 
inferior  margin  rather  long  and  nearly  straight ;  subangular  tubercle 
considerably  posterior  to  inferior  dental  foramen;  coronoid  blade 
broad,  its  apex  only  moderately  recurved,  ending  anterior  to  plane 
of  condyle;  canines  of  medium  size,  the  lower  ones  rather  massive; 
molars  of  medium  size,  the  upper  rather  small  for  size  of  skull. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  323 ; l  oc- 
cipito-nasal  length,  304;  palatal  length,  175;  zygomatic  breadth,  229; 
interorbital  breadth,  85. 

URSUS  WASHAKE  Merbiam. 

Washakie  Grizzly. 

Ursus   ivashake  Merriam,   Proc.   Biol.   Soc.   Washington,   XXIX,   pp.   152-154, 
September  6,  1916. 

Type  locality. — North  Fork  Shoshone  River,  Absaroka  Mountains, 
western  Wyoming  (between  Bighorn  Basin  and  Yellowstone  National 
Park). 

Type  specimen. — No.  213005,  $  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Killed  September,  1913,  by  Col.  J.  A. 
McGuire. 

Cranial  characters. — Old  male  (type)  :  Size  medium,  about  equal- 
ing male  shoshone  and  male  horriwits;  skull  rather  short  and  high, 
moderately  arched,  with  broad,  elevated  postorbitals  and  rather 
broadly  outbowed  zygomata.  Frontal  shield  rather  narrow,  sloping 
strongly  upward  anteriorly,  highest  at  postorbital  processes,  hori- 
zontal posteriorly,  broadly  concave  between  postorbital  >  processes ; 
postorbital  processes  large,  broad,  subtriangular  as  viewed  from 
above,  outstanding,  elevated  and  slightly  arched,  rising  well  above 
frontal  plane  and  passing  anteriorly  into  thickened  orbital  rims; 
fronto-nasal  region  dished  (change  of  angle  about  middle  of  nasals) ; 
rostrum  rather  small,  strongly  compressed  horizontally  between 
nasals  and  roots  of  canines,  making  nasals  appear  elevated ;  anterior 
nares  small;  zygomata  rather  slender,  broadly  spreading,  rounded 

1  Restored. 


1918.]  RICHARDSONI   GROUP.  105 

and  strongly  outbowed  posteriorly,  only  slightly  expanded  verti- 
cally; sagittal  crest  low;  postpalatal  shelf  broad,  flat,  and  rather 
short;  occipito-sphenoid  87  mm.  (=  distance  from  front  of  canine 
to  or  slightly  beyond  middle  of  M1).  Underjaw  moderate;  ramus 
bellied  posteriorly;  coronoid  blade  high  and  rather  falcate,  the  apex 
cutting  plane  of  condyle  (line  from  apex  to  tip  of  angular  process 
passing  well  behind  condyle).  Teeth  moderate  or  rather  large; 
M^  large. 

Cranial  comparisons. — Old  male  (type)  compared  with  adult  male 
ophrus  (type)  :  Size  about  the  same,  but  appearing  smaller;  frontal 
shield  less  elevated  and  less  deeply  concave;  fronto-nasal  region 
elevated  instead  of  sulcate-dished ;  zygomata  less  widely  outbowed; 
postorbitals  much  broader  and  less  elevated;  orbital  rims  less 
swollen;  postpalatal  shelf  shorter  and  broader;  mastoids  shorter; 
nares  smaller  and  lower.  Underjaw  more  massive;  inferior  border 
of  ramus  more  swollen  and  much  more  bellied  posteriorly ;  coronoid 
blade  higher  and  more  falcate,  the  apex  reaching  much  farther  pos- 
teriorly (cutting  plane  of  hinder  part  of  condyle) . 

Compared  with  adult  male  canadensis  (type),  to  which  it  is  not 
related  but  with  which  it  agrees  essentially  in  basal  and  occipito- 
sphenoid  length :  Frontal  shield  less  flat,  more  elevated  laterally, 
highest  at  postorbitals  instead  of  at  posterior  point;  fronto-nasal 
region  more  dished ;  rostrum  smaller,  narrower  basally,  more  strongly 
compressed  below  nasals;  postorbital  processes  very  much  larger, 
broader,  and  more  massive,  elevated,  arched,  and  subtriangular, 
instead  of  slender  and  narrowly  peglike;  zygomata  more  outbowed 
and  arched;  sagittal  crest  low  and  straight  instead  of  high  and 
arched;  inion  less  developed;  braincase  anteriorly  broader  and  more 
depressed — not  tending  to  "  keel  "  into  sagittal  crest  as  in  canadensis; 
occipito-nasal  length  less,  although  basal  length  of  skull  is  essentially 
the  same  in  both.  Underjaw  longer;  inferior  border  of  ramus 
shorter  and  more  strongly  bellied;  coronoid  blade  higher  and  more 
falcate,  its  apex  reaching  farther  posteriorly;  distance  from  angle 
to  subangular  process  much  greater;  diastema  in  both  jaws  much 
longer.    Last  upper  and  middle  lower  molars  not  quite  so  broad. 

Compared  with  adult  male  absarokus  (type)  :  Size  smaller;  vault 
of  cranium  decidedly  lower ;  braincase  anteriorly  broadly  depressed ; 
frontal  shield  narrower,  lower,  and  flatter,  concave  instead  of  convex 
between  postorbital  processes;  postorbital  processes  (viewed  from 
above)  broadly  triangular,  uplifted,  and  somewhat  arched  instead 
of  pegshape;  orbital  rims  more  thickened  and  elevated;  rostrum 
smaller,  lower,  more  slender,  and  much  more  compressed  horizontally 
between  nasals  and  roots  of  canines,  making  the  nasals  appear 
elevated;  sagittal  crest  lower;  occipito-sphenoid  shorter;  occiput 
lower;  anterior  nares  smaller.    Underjaw  shorter;  inferior  border  of 


106  NORTH   AMERICAN   FAUNA.  [No.  41 . 

ramus  much  shorter;  coronoid  blade  more  falcate,  its  apex  reaching 
farther  posteriorly;  angular  process  more  slender  and  more  pro- 
duced posteriorly  (line  connecting  apex  of  coronoid  with  angle  pass-)! 
ing  well  behind  condyle — in  dbsarohus  cutting  condyle  near  middle).! 
Molars  smaller. 

Skull  measurements. — Adult  male  (type):  Basal  length,  310;  oc- 
cipito-nasal  length,  305;  palatal  length,  170;  zygomatic  breadth,  217; 
interorbital  breadth,  76. 

Xidderi  Group. 
URSUS  KIDDERI   KIDDERI1  Merbiam. 

Kidder   Beab. 

(PI.  VIII.) 

Ursus  kidderi  Merriam,  Proc.  Biol.  Soc.  Washington,  XV,  p.  78,  March  22,  1902. 

Type  locality. — Chinitna  Bay,  Cook  Inlet,  Alaska. 

Type  specimen. — No.  1L6562,  $  young  (not  fully  grown),  U.  S. 
National  Museum,  Biological  Survey  collection.  Collected  June  9, 
1901,  by  James  H.  Kidder. 

Range. — Alaska  Peninsula  for  its  entire  length. 

Characters. — Size  medium — small  compared  with  gyas  of  the 
same ,  region ;  sexual  disparity  small,  female  nearly  as  large  as 
male;  skull  of  adult  male  hardly  half  the  bulk  of  male  gyas; 
skull  of  adult  female  nearly  the  same  size  as  that  of  female  gyas; 
color  yellowish  brown ;  claws  rather  short,  blue-black. 

Color. — June  specimens  (in  left-over  winter  pelage) :  General 
color  yellowish  brown,  darkest  on  belly  and  legs,  legs  much  darker 
than  body.  Most  of  the  Kidder  bears  in  the  National  Zoological 
Park  are  pale  buffy  yellowish,  or  yellowish  cream  color. 

Cranial  characters. — Adult  moJe:  Skull  long,  rather  low,  narrow, 
and  massive;  frontal  shield  narrow,  moderately  sloping;  swollen 
over  orbits  and  rather  deeply  sulcate  or  troughed  medially;  fronto- 
nasal region  slightly  dished;  rostrum  and  nasals  high  and  rather 
long;  postorbital  processes  peglike,  outstanding;  braincase  elon- 
gate; zygomata  angular,  only  moderately  spreading;  sagittal  crest 
long;  palate  long;  mastoids  long;  underjaw  long,  with  long  ramus 
and  moderate  or  low  coronoid  blade.  Adult  female:  Skull  in  general 
like  that  of  male  but  slightly  smaller  and  with  more  slender  rostrum 
and  slightly  smaller  teeth.  The  sagittal  crest  extends  much  farther 
forward  than  in  the  females  of  most  species,  in  this  respect  also 
resembling  the  male.  The  sexual  difference  in  size  of  teeth  appears 
to  be  covered  by  individual  variation. 

1  Named  for  James  H.  Kidder,  who  collected  and  presented  the  type  specimen. 


1918.1  KIDDERI   GEOUP.  107 

Cranial  comparisons. — Adult  male  and  female  compared  with  adult 
female  gyas :  Adult  males  require  no  comparison,  owing  to  the  great  dif- 
ference in  size  of  skull  and  teeth,  but  with  the  females  the  case  is  very 
different,  the  size  being  essentially  the  same.  In  fully  adult  females 
\kidderi  is  easily  distinguished  by  the  lowness  of  the  vault  of  the 
!  cranium  and  greater  length  of  the  sagittal  crest — female  gyas  being 
rather  highly  domed  and  having  the  short  sagittal  crest  of  most 
female  bears. 

Males  of  Mdderi  are  sometimes  hard  to  tell  from  females  of  gyas, 
but  in  the  case  of  fully  adult  skulls  they  may  be  distinguished  as 
follows:  Male  kidderi  averages  longer,  both  in  basal  and  occipito- 
nasal  length,  is  much  less  highly  arched,  and  is  more  obliquely  trun- 
cate posteriorly  so  that  the  occiput  overhangs,  giving  the  effect  of  a 
longer  braincase.  The  rostrum  also  is  somewhat  longer.  The 
frontal  shield  is  quite  different,  being  very  much  shorter  posteriorly, 
ending  about  an  inch  in  front  of  the  parietals,  whereas  in  female 
gyas  the  posterior  point  of  the  shield  extends  posteriorly  to  about  the 
same  distance  behind  the  fronto-parietal  suture.  Thus  in  adult  male 
kidderi  the  frontal  shield  is  horizontal  or  slopes  forward  from  its 
most  posterior  point,  while  in  adult  female  gyas  it  slopes  backward 
for  some  distance — from  a  point  at  least  an  inch  in  front  of  the 
fronto-parietal  suture — the  shield  thus  overreaching  the  highest 
point  of  the  arch  of  the  skull  and  sloping  downward  in  both  direc- 
tions. The  occipito-sphenoid  length  is  slightly  greater  in  female 
gyas  than  in  male  kidderi. 

Comparison  of  cubs  of  the  second,  third,  and  fourth  years:  Cubs  of 
kidderi  after  the  molars  are  in  position  and  the  permanent  canines 
partly  exposed,  are  easily  told  from  those  of  male  gyas  of  corre- 
sponding age  by  the  relatively  small  size  of  the  teeth,  especially  the 
canines,  molars,  and  outer  upper  incisors.  It  is  not  so  easy,  however, 
to  tell  them  from  female  gyas,  and  in  some  cases  it  may  be  impossible. 
The  most  constant  character  appears  to  be  the  length  of  the  canines, 
and  this  can  not  be  determined  in  cubs  less  than  three  or  three  and  a 
half  years  of  age.  The  canines  are  slightly  longer  in  male  kidderi 
than  in  female  gyas,  and  their  diameter  also  is  usually,  but  not 
always,  greater.  As  a  rule,  also,  the  crowns  of  the  molars  are  larger, 
especially  longer,  in  male  kidderi  the  combined  length  of  the  first 
and  second  lower  molars  averaging  about  4  mm.  greater  than  in 
female  gyas.  As  the  teeth  become  worn  with  use  the  differences 
become  less  and  less  obvious,  so  that  after  the  third  year  the  distinc- 
tions are  not  easy  of  recognition. 

Skull  measurements. — Average  of  3  males  from  Belkofski,  Alaska 
Peninsula:  Basal  length,  330;  occipito-nasal  length,  312;  palatal 
length,    177;    zygomatic    breadth,    207;    interorbital    breadth,    74. 


108  NORTH  AMERICAN  FAUNA.  [No.  41. 

Average  of  4  females  from  Pavlof  Bay,  Alaska  Peninsula:  Basal 
length,  330;  occipito-nasal  length,  312;  palatal  length,  181.5;  zygo- 
matic breadth,  218 ;  interorbital  breadth,  80.5. 

URSUS  KIDDERI  TUNDRENSIS  Mebbiam. 

TtJNDBA  BEAB. 

Ursus  kidderi  tundrensis  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  196, 
August  13,  1914. 

Type  locality. — Shaktolik  Eiver,  Norton  Sound,  Alaska. 

Type  specimen. — No.  76470,  $  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  by  natives,  September,  1894, 
and  secured  through  the  late  Rudolf  Neumann,  then  of  Iliuliuk, 
Unalaska. 

Range. — Tundra  region  of  northwestern  Alaska  from  Shaktolik 
River  on  Norton  Sound,  southerly  across  the  lower  Yukon,  Kus- 
kokwim,  and  Nushagak  Rivers  to  Bristol  Bay  and  north  side  of 
base  of  Alaska  Peninsula. 

Characters. — Size  medium  (small  in  contrast  with  gyas),  about 
equaling  kidderi.  External  characters  unknown,  but  doubtless  little 
different  from  kidderi.     Known  to  the  natives  as  "  Red  Bear." 

Cranial  characters. — Size  medium,  about  as  in  kidderi;  skull  rather 
long  and  heavy;  frontals  broad  and  flat,  broadly  but  not  deeply 
sulcate  medially;  postorbital  processes  small;  coronoid  blade  falcate 
and  rather  high. 

Cranial  comparisons. — Similar  to  kidderi  in  essential  cranial  and 
dental  characters,  but  differing  in  having  the  frontal  shield  and  post- 
orbital  processes  very  much  broader  and  flatter,  and  the  frontal  sulcus 
less  marked.  As  in  kidderi  there  is  little  difference  in  the  sexes 
except  that  the  females  have  narrower  muzzles  and  narrower  frontal 
shields.  A  young  male  (No.  16375)  from  Andreafski  on  the  lower 
Yukon,  collected  by  E.  W.  Nelson,  has  somewhat  shorter  canines 
than  the  others. 

From  alascensis,  the  grizzly  of  the  same  general  region,  skulls  of 
tundrensis  (adult  males  in  both  cases)  may  be  distinguished  by  the 
foll®wing  characters:  Size  larger  (basilar  and  occipito-nasal  lengths 
fully  an  inch  greater) ;  skull  as  a  whole  much  more  massive ;  frontals 
broader,  rising  less  abruptly  from  facial  plane,  less  swollen  over 
orbits;  rostrum  less  slender;  palate  more  flat;  ramus  of  jaw  thicker 
under  MT  and  MT;  coronoid  blade  more  falcate;  canines  larger, 
decidedly  longer,  and  somewhat  less  curved ;  molars  decidedly  larger. 

Skull  measurements. — Adult  male  (type) :  Basal  length,  333 ; 
occipito-nasal  length,  317;  palatal  length,  178;  zygomatic  breadth, 
228;  interorbital  breadth,  93. 


1918.]  KIDDERI   GROUP.  109 

URSUS  EXIMIUS  Merkiam. 

Knik  Beab. 

Ursus  eximius  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  139-140,  Sep- 
tember 6,  1916. 

Type  locality. — Head  of  Knik  Arm,  Cook  Inlet,  Alaska. 

Type  specimen. — No.  122495,  $  ad.,  U.  S.  National  Museum.  Col- 
lected by  G.  W.  Palmer. 

Characters. — Size  rather  large ;  color  uniform  rich  dark  brown  sug- 
gesting seal  brown;  muzzle  brown,  paler  than  rest  of  head;  back  of 
head  and  neck  lightly  sprinkled  with  pale-tipped  hairs;  claws  of 
medium  thickness,  only  slightly  curved,  decidedly  short,  probably 
from  wear,  smooth,  very  dark  horn  color,  becoming  paler  on  sides 
toward  tip.  Skull  long  and  narrow,  with  narrow  highly  arched 
frontals.    Related  to  kidderi. 

Cranial  characters. — Adult  male  (type) :  Skull  long,  extremely 
narrow  in  fronto-nasal  region,  rather  highly  arched  and  strongly 
dished.  Frontal  shield  exceedingly  narrow,  convex,  shallowly  sulcate 
medially,  strongly  arched  anteriorly,  horizontal  posteriorly,  long- 
pointed,  the  point  nearly  reaching  f ronto-parietal  suture ;  postorbital 
processes  slender,  peglike,  moderately  outstanding;  fronto-nasal 
region  strongly  dished;  rostrum  long,  narrow,  high,  compressed  be- 
tween nasals  and  canine  roots;  zygomata  moderately  outstanding, 
subtriangular ;  palate  long  and  narrow;  postpalatal  shelf  relatively 
broad ;  notch  rather  broad ;  mastoids  long,  strongly  divergent ;  under- 
jaw  long,  moderately  massive,  the  ramus  broad  vertically;  coronoid 
blade  rather  broad,  the  apex  not  strongly  recurved ;  teeth  of  medium 
size ;  M>  with  rather  long  heel,  not  much  narrowed  posteriorly ;  M> 
relatively  large  and  broad;  PM4  a  single  cone  without  distinct  heel 
but  sulcate  posteriorly. 

Adult  female  (No.  205176,  from  type  locality) :  Skull  long  and 
narrow ;  vault  of  cranium  moderately  arched,  the  highest  part  form- 
ing a  hump  at  f ronto-parietal  suture ;  frontal  shield  narrow,  flattish, 
sulcate  medially,  the  point  reaching  f ronto-parietal  suture;  postor- 
bitals  weak,  subtriangular,  not  decurved ;  fronto-nasal  region  moder- 
ately dished ;  rostrum  narrow,  compressed  between  nasals  and  canine 
roots;  zygomata  moderately  spreading,  subtriangular;  postpalatal 
shelf  relatively  broad;  notch  moderate;  inferior  border  of  ramus 
convex  from  plane  of  front  molar  posteriorly ;  coronoid  blade  broad 
and  low. 

Cranial  comparisons. — Ursus  eximius  appears  to  be  related  to  only 
a  single  species,  kidderi  of  Alaska  Peninsula.  Adult  male  (type) 
compared  with  a  series  of  kidderi  from  various  points  on  Alaska 
Peninsula:  Size  about  the  same;  vault  of  cranium  more  highly 
arched;  frontal  shield  narrower,  more  strongly  convex  in  cross  sec- 


110  NORTH  AMERICAN   FAUNA.  INo.  41. 

tion,  less  deeply  silicate;  postpalatal  processes  more  slender,  peglike, 
and  outstanding;  fronto-nasal  region  more  dished;  rostrum  more 
slender;  nasals  longer  posteriorly,  more  completely  wedge-shaped, 
reaching  posteriorly  to  plane  of  postpalatal  processes;  mastoids 
longer  and  more  divergent. 

Adult  female  (No.  205176,  from  head  of  Knik  Arm)  compared 
with  adult  female  Mdderi:  Size  materially  smaller;  frontal  shield 
and  rostrum  much  narrower;  vault  of  cranium  notably  higher  over 
f ronto-parietal  suture ;  braincase  narrower ;  nasals  longer  posteriorly ; 
under  jaw  smaller  and  lighter. 

Adult  male  (type)  compared  with  adult  male  alascensis  (No. 
76466,  from  Unalaklik  River,  Norton  Sound,  Alaska)  :  Skull  much 
longer,  more  highly  arched,  and  narrower  throughout.  Frontal 
shield  much  more  elevated,  narrower,  and  longer  posteriorly ;  fronto- 
nasal region  more  strongly  dished;  rostrum  narrower  and  higher; 
lambdoid  crest  more  strongly  developed ;  palate  and  postpalatal  shelf 
much  longer;  occipito-sphenoid  much  longer;  mastoids  much  longer 
and  strongly  divergent;  under  jaw  longer;  coronoid  blade  much 
higher;  teeth  larger;  heel  of  M^  much  longer. 

Adult  female  compared  with  female  alascensis:  Length  essentially 
the  same;  skull  narrower  throughout;  frontal  shield  lower,  much 
narrower  and  flatter,  rising  less  abruptly  from  rostrum ;  fronto-nasal 
region  sulcate  but  less  strongly  dished;  rostrum  slightly  more 
slender;  postpalatal  shelf  narrower.  Underjaw  about  same  length; 
inferior  border  of  ramus  more  evenly  convex  (less  abruptly  bellied) ; 
coronoid  blade  broader;  canines  about  same  size;  molars  somewhat 
larger. 

Skull  measurements. — Adult  male  (type):  Basal  length,  331; 
occipito-nasal  length,  319;  palatal  length,  185;  zygomatic  breadth, 
215;  interorbital  breadth,  71. 

Innuitus  Group. 

URSUS  INNUITUS  Mebbiam. 

Innuit  Beab. 

(PI.  VII.) 

Ursus  innuitus  Merriam,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  177,  August  13, 
1914. 

Type  locality. — Golofnin  Bay,  south  side  of  Seward  Peninsula, 
northwestern  Alaska. 

Type  specimen. — No.  179780,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.    Collected  in  1886,  by  Edward  F.  Ball. 

Range. — Coastal  region  of  Norton  Sound,  Alaska,  from  Unalaklik 
northward  and  westward ;  limits  unknown. 


1918.]  INNUITUS   GROUP.  Ill 

Characters. — Size  large;  external  characters  unknown.  Molars 
large  and  massive,  especially  M-2-.  Large  lower  premolar  subcorneal, 
apparently  of  the  brown  bear  type.  But  in  the  north  the  grizzly 
type  of  premolar  often  fails  in  true  grizzlies ;  hence  not  having  seen 
the  claws,  it  is  at  present  impossible  to  say  whether  innuitus  is  a 
brown  bear  or  a  grizzly. 

Cranial  characters. — Old  male  (type):  Skull  large;  basal  length 
essentially  the  same  as  in  horrihilis  and  alexandraz  but  occipito-nasal 
length  much  less,  owing  to  shortness  of  occiput ;  f ronto-nasal  region 
strikingly  dished:  rostrum  short,  exceedingly  broad  (of  same  breadth 
as  in  alexandrce,  very  much  broader  than  in  horrihilis),  strongly  de- 
pressed; frontal  shield  exceedingly  broad  interorbitally,  rising  high 
and  abruptly  from  rostrum,  nearly  horizontal  behind  plane  of  post- 
orbital  processes,  rather  deeply  sulcate  medially  and  strongly 
swollen  over  orbits;  postorbital  processes  large,  subtriangular,  out- 
standing and  decurved;  nasals  nearly  horizontal;  palate  and  post- 
palatal  shelf  broad;  postpalatal  notch  of  medium  width;  zygomata 
broadly  spreading  and  somewhat  outbowed  posteriorly,  acute  ante- 
riorly ;  nares  broader  than  high ;  sagittal  crest  short,  extending  only 
about  25  mm.  beyond  fronto-parietal  suture,  straight  (not  arched), 
high  posteriorly ;  lambdoid  crest  large  and  full ;  coronoid  blade  nar- 
row and  high;  ramus  long  and  flat.  Canines  badly  broken,  appar- 
ently long;  last  lower  premolar  broad,  broader  posteriorly  than  ante- 
riorly, the  cusp  small  and  sloping  posteriorly  without  heel  or  mar- 
ginal cusplets,  but  with  pit  and  indication  of  narrow  sulcus;  molars 
exceptionally  large  and  broad,  the  last  upper  one  with  heel  strikingly 
long  and  broad,  agreeing  almost  exactly  with  that  of  true  horrihilis 
from  eastern  Montana. 

Two  youngish  skulls  from  Unalaklik  (No.  82024,  third  year  and 
No.  210554,  fourth  year)  are  believed  to  be  females  of  this  species. 
They  are  not  old  enough  to  show  adult  cranial  characters  except  that 
the  postpalatal  shelf  is  broad  and  flat  and  the  notch  broad,  but  the 
teeth  are  perfect,  full  grown,  and  unworn.  Canines  of  medium  size 
(in  the  type  badly  broken  and  hence  not  available  for  comparison) ; 
molars  large  but  smaller  and  less  massive  than  those  of  type;  M4 
large,  cusps  on  inner  side  nearly  obsolete  (presenting  little  more 
than  an  undulating  line),  heel  long  and  broad  with  large  flat  gran- 
ular grinding  surface. 

Cranial  comparisons. — The  only  species  needing  comparison  with 
innuitus  are  alexandrw  and  cressonus.  The  old  male  skull  resembles 
certain  old  skulls  of  alexandrai  from  Kenai  Peninsula,  but  differs 
rather  strikingly  in  truncation  of  occiput  (the  occipital  overhang 
being  very  much  less)  and  corresponding  shortening  of  braincase 
and  sagittal  crest.  The  crest  moreover  is  straight  instead  of  arcuate 
64854°— 18 8 


112  NORTH   AMERICAN   FAUNA.  [No.  41. 

or  arched,  and  the  molars,  especially  M1  and  M A  are  very  much 
larger. 

Compared  with  old  male  cressonus  (type) :  Basal  length  and 
zygomatic  breadth  essentially  same ;  occipito-nasal  length  much  less ; 
frontal  shield  broader,  much  less  elevated  over  orbits ;  rostrum  much 
broader  and  lower;  occipital  overhang  much  less;  braincase  and 
sagittal  crest  much  shorter;  postpalatal  shelf  broader  and  flatter. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  353 ;  occipito- 
nasal  length,  331;  palatal  length,  194;  zygomatic  breadth,  251; 
interorbital  breadth,  104. 

URSUS   CRESSONUS   Mebbiam. 

Chitina  Beak. 

Ursus  cressonus  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  137-139, 
September  6,  1916. 

Type  locality. — Lakina  River,  south  slope  of  Wrangell  Range, 
Alaska. 

Type  specimen. — No.  206529,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  by  Captain  J.  P.  Hubrick,  of 
McCarthy,  Alaska,  1914. 

Range. — Chitina  River  valley  and  adjacent  slopes  of  Skolai  and 
Wrangell  Mountains,  westerly  doubtless  through  Chugach  Moun- 
tains to  the  west  side  of  Cook  Inlet;  occurs  as  far  south  as  the 
Iliamna  region. 

Cranial  characters. — Old  male  (type) :  Skull  peculiar  and  distinc- 
tive; size  large  (basal  length  357  mm.);  skull  long,  narrow,  high, 
and  strongly  dished ;  frontal  shield  highly  elevated,  rising  abruptly 
from  rostrum,  rather  broad,  deeply  sulcate  throughout  medially, 
swollen  over  orbits,  short  pointed  posteriorly ;  orbits  nearly  vertical ; 
postorbital  processes  small  and  strongly  decurved;  fronto-nasal  re- 
gion sulcate  and  strongly  dished;  rostrum  rather  short  and  narrow; 
nasals  horizontal  except  posteriorly,  where  they  rise  strongly ;  brain- 
case  exceedingly  long;  sagittal  crest  high  posteriorly,  straight,  and 
long,  reaching  anteriorly  to  halfway  between  parietals  and  plane  of 
postorbitals ;  zygomatic  arches  moderately  spreading,  subtriangular 
(not  outbowed),  expanded  vertically;  palate  and  postpalatal  shelf 
relatively  long  and  narrow  for  so  large  a  skull;  postpalatal  notch 
rather  narrow  and  short;  occipito-sphenoid  long  (about  103  mm.); 
mastoids  outstanding;  anterior  nares  rather  small,  subtruncate,  and 
broader  than  high  in  type  skull,  higher  and  less  truncate  in  the 
Iliamna  skulls.  Under  jaw  absent  in  type  specimen,  but  in  an  old 
male  from  Iliamna  on  north  side  of  Cook  Inlet  (No.  209885)  which 


1918.]  INITUITUS  GROUP.  113 

in  most  respects  closely  matches  the  type,  the  ramus  is  broadly  flat- 
tened vertically,  much  broader  posteriorly  than  anteriorly,  and  the 
coronoid  blade  is  high  and  rather  vertical.  In  younger  skulls  from 
Iliamna  the  coronoid  is  broader  basally  and  less  high.  Canines  large 
and  massive;  molars  moderate.  The  last  upper  molar  is  large  in 
the  type,  smaller  and  more  cut  away  on  outer  side  of  heel  in  the 
Iliamna  specimens.  No.  209885  from  Iliamna  agrees  with  the  type, 
except  that  the  nares  is  higher  and  less  truncate,  and  the  last  upper 
molar  smaller,  with  heel  more  cut  away  on  outer  edge. 

Old  female  (No.  209881,  from  head  of  Chitina  Eiver,  80  miles  from 
McCarthy,  Alaska;  collected  by  Capt.  J.  P.  Hubrick)  :  Size  medium; 
cranium  moderately  arched;  frontal  shield  broad,  deeply  sulcate  an- 
teriorly, strongly  swollen  over  and  posterior  to  orbits,  the  point 
lyrate  and  reaching  parietals;  postorbitals  rather  large,  blunt,  and 
somewhat  decurved;  fronto-nasal  region  strongly  dished  and  de- 
pressed medially ;  rostrum  rather  large  and  high,  nearly  horizontal ; 
palate  and  postpalatal  shelf  broad;  postpalatal  notch  moderate  and 
rather  broad.  Underjaw  long;  coronoid  blade  high  and  rather 
narrow,  its  apex  only  slightly  recurved.  Dentition  light;  canines 
small  and  short;  molars  rather  narrow,  apparently  normal  (too 
badly  worn  to  admit  of  description,  except  that  the  heel  of  M>  is 
moderately  long  and  rather  broadly  rounded  posteriorly). 

Cranial  comparisons. — Old  male  (type)  compared  with  old  male 
dalli:  Size  about  the  same;  vault  of  cranium  and  frontal  shield  much 
more  elevated,  less  flat,  less  nearly  horizontal,  and  much  more  swollen 
over  orbits;  shield  more  deeply  sulcate;  postorbitals  weak  and  de- 
curved  (in  dalli  larger  and  more  horizontally  outstanding) ;  fronto- 
nasal region  more  strongly  dished;  rostrum  narrower  and  longer; 
zygomata  much  less  widely  outstanding  and  much  less  bowed ;  palate 
longer ;  molars  very  much  larger. 

Old  male  (type)  compared  with  adult  male  nuchek  (type)  :  Size, 
elevation  of  vault  of  cranium,  and  zygomatic  breadth  about  the  same ; 
frontal  shield  somewhat  broader,  much  more  highly  arched,  much 
more  swollen  over  orbits,  much  more  deeply  sulcate  medially,  shorter 
and  more  acutely  pointed  posteriorly ;  postorbitals  smaller  and  more 
decurved;  fronto-nasal  region  strongly  dished;  rostrum  more  de- 
pressed; nares  more  truncate;  last  upper  molar  of  normal  form, 
large,  and  with  long  posteriorly  rounded  heel,  differing  widely  from 
the  short,  broad-in-the-middle,  obliquely  truncate  tooth  of  nuchek. 

Skull  measurements. — Old  male  (type):  Basal  length,  357;  oc- 
cipito-nasal  length,  354 ;  palatal  length,  199 ;  zygomatic  breadth,  244 ; 
interorbital  breadth,  97. 


114  NORTH   AMERICAN    FAUNA.  I  No.  41 

URSUS  ALEXANDRA1  Merbiam.* 
Alexander  Grizzly. 

Ursus  alexandrw  Merrlam,  Proc.  Biol.  Soc.  Washington,  XXVII,  pp.  174-175, 
August  13,  1914. 

Type  locality. — Kusilof  Lake,  Kenai  Peninsula,  Alaska. 

Type  specimen. — Skull  No.  4752,  $  old,  Museum  of  Vertebrate 
Zoology,  University  of  California ;  original  No.  218.  Collected  Sep- 
tember, 1906.     (Skull,  skin,  and  skeleton  complete.) 

Range. — Kenai  Peninsula,  Alaska. 

Characters. — Size  very  large ;  skull  long  and  narrow ;  rostrum  ex- 
ceptionally broad  for  a  grizzly;  pelage  very  uniform  in  color, 
scarcely  or  not  grizzled;  clav^s  enormous  (second  foreclaw  of  type 
specimen  measuring:  length  from  upper  base,  91  mm;  height  at 
base,  25;  breadth,  11.5).  The  longest  claw  in  a  specimen  collected 
by  Wilson  Potter  measures  120  mm.;  in  a  male  killed  by  Dall  De- 
Weese,  110  mm. 

Color. — Type,  very  old  male,  in  fresh  short  fall  pelage:  General 
color  pale,  almost  grayish  brown,  becoming  yellowish  brown  between 
ears,  contrasting  with  pale  brown  of  muzzle;  legs  and  feet  only 
slightly  darker  than  back;  entire  animal  remarkably  unicolor;  under- 
fur  plumbeous,  crinkled,  and  wooly.  Another  male,  killed  by  Wilson 
Potter,  of  Philadelphia,  in  May,  1912  (belonging  to  skull  No.  181102, 
presented  by  Wilson  Potter),  is  pale  buffy  inclining  to  light  reddish 
brown  throughout,  without  grizzly  appearance;  legs  only  slightly 
darker.  One  killed  by  Dall  DeWeese,  of  Canyon  City,  Colorado, 
September  7,  1897,  is  described  by  him  as  "grayish-yellow,"  with 
legs  and  sides  chocolate-brown. 

Cranial  characters. — Skull  large,  long,  rather  narrow,  with  mod- 
erately spreading  zygomata,  short  flattish  frontal  shield,  outstanding 
postorbitals  (with  age)  ;  very  broad  rostrum  for  a  grizzly,  and  long 
and  high  sagittal  crest.  Frontal  shield  not  markedly  elevated  above 
plane  of  rostrum;  posterior  root  of  zygoma  not  expanded.  Canines 
large  and  long. 

Cranial  comparisons. — Adult  male  compared  with  adult  male 
kenaicnds:  Basal  length,  palate,  and  occipito-sphenoid  length  essen- 
tially the  same ;  skull  as  a  whole  much  narrower,  frontal  shield  inter- 
orbitally  and  across  postorbital  processes  much  narrower,  flatter,  more 
nearly  horizontal,  not  materially  elevated  above  plane  of  rostrum; 
zygomata  much  less  widely  spreading,  squamosal  part  much  narrower 
(not  expanded)  ;  sagittal  crest  much  longer,  reaching  anteriorly  over 
posterior  third  of  frontals  (in  kenaiensis  ending  on  or  near  fronto- 
parietal suture) ;  posterior  third  of  frontals  compressed,  rising  in 

1  Named  for  Annie  M.  Alexander,  founder  of  the  Museum  of  Vertebrate  Zoology,  Uni- 
versity of  California,  Berkeley. 

*  Tentatively  included  in  innuitus  group.      (See  Introduction,  pp.   12-13.) 


1918.]  TOWNSENDI  GROUP.  115 

a  keel  to  sagittal  crest.  Canine  teeth,  both  upper  and  lower,  but 
especially  the  lower,  much  larger  and  longer. 

Remarks. — The  skull  of  alexandrce  is  of  a  generalized  type,  lacking 
the  special  distinctive  features  that  characterize  several  of  its  neigh- 
bors— as  kenaiensis,  sheld&ni,  and  others — none  of  which  are  true 
grizzlies.  Among  the  grizzlies  it  stands  alone  in  the  great  breadth  of 
the  rostrum,  which  in  bears  of  its  size  is  only  exceeded  by  the  widely 
different  kenaiensis.  Ursus  alexandrce  attains  the  largest  size  known 
among  the  grizzly  bears,  the  biggest  skulls  equaling  those  of  the 
huge  magister  of  Southern  California. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  355 ;  occi- 
pito-nasal  length,  358;  palatal  length,  191;  zygomatic  breadth,  252; 
interorbital  breadth,  87. 

Townsendi  Group. 

URSUS  TOWNSENDI1  Mebbiam. 

Town  send  Beab. 

Ursus  townsendi  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  151-152, 
September  6,  1916. 

Type  locality. — Mainland  of  Southeastern  Alaska  (exact  locality 
uncertain). 

Type  specimen. — No.  216643,  $  old,  U.  S.  National  Museum. 
Purchased  at  Sitka,  in  1889,  by  Dr.  Charles  H.  Townsend. 

Cranial  characters. — Skull  large,  long,  massive,  rather  low,  and 
flat  topped,  dished,  with  extremely  small  teeth.  Shield  broad,  flat, 
slightly  depressed  medially,  the  point  ending  anterior  to  parietals, 
sides  reaching  out  broadly  into  very  broad  postorbitals,  strongly 
sloping  to  rostrum;  rostrum  moderate,  flat  or  depressed  on  top; 
nares  truncate;  zygomata  moderately  outstanding  and  moderately 
bowed;  squamosal  base  broadly  and  abruptly  expanded  vertically; 
palate  and  postpalatal  shelf  moderate;  notch  rather  narrow;  mas- 
toids long;  occipito-sphenoid  95  mm.;  basisphenoid  rather  deeply 
concave.  Underjaw  long;  ramus  broad  and  flat  vertically;  coronoid 
of  moderate  height,  narrowing  above,  sloping  strongly  backward, 
apex  cutting  plane  of  posterior  part  of  condyle ;  upper  two-thirds  of 
anterior  border  strongly  inflected. 

Cranial  comparisons. — Old  male  (type)  compared,  with  male 
caurinus :  Skull  much  larger,  broader,  more  massive,  and  less  arched ; 
teeth  smaller.  Frontal  shield  very  much  broader  interorbitally  and 
postorbitally   (interorbitally  91  mm.  contrasted    with    81    or    less; 

*  Named  for  Dr.  Charles  H.  Townsend,  formerly  naturalist  of  the  Fish  Commission 
steamer  Albatross,  now  director  of  the  New  York  Aquarium,  who  collected  the  specimen 
and  presented  it  to  the  U.  S.  National  Museum. 


116  NORTH   AMERICAN   FAUNA.  |No.41. 

across  postorbitals  130  contrasted  with  116)  ;  postorbitals  very  much 
broader  and  flatter;  rostrum  more  nearly  horizontal;  nares  truncate 
instead  of  sloping;  zygomata  more  widely  outstanding  and  more 
broadly  expanded  vertically. 

Skull  measurements. — Old  male  (type):  Basal  length,  348;  occi- 
pito-nasal  length,  353;  palatal  length,  183;  zygomatic  breadth,  245; 
interorbital  breadth,  91.5. 

Dalli  Group. 

URSUS  DALLI1  Merriam. 

Dall  Brown  Bear. 

Urgus  dalli  Merriam,  Proc.  Biol.  Soc.  Washington,  X,  pp.  71-73,  April  13,  1896. 

Type  locality. — Yakutat  Bay  (northwest  side),  Alaska. 

Type  specimen. — No.  75048,   $  old,  U.  S.  National  Museum. 

Range. — Malaspina  Glacier  and  region  northwest  of  Yakutat  Bay, 
Alaska. 

Characters. — Size  very  large ;  skull  without  very  pronounced  char- 
acters, although  differing  sufficiently  from  its  neighbors.  General 
color  dark  brown,  strongly  grizzled. 

Color. — Adult  male  (from  Malaspina  Glacier,  belonging  to  skull 
No.  210293,  killed  by  G.  Frederick  Norton) :  Muzzle  pale  brown,  be- 
coming much  darker  on  head  and  sides  of  face;  general  body  color 
dark  brown,  moderately  grizzled  with  pale-tipped  hairs,  the  grizzling 
beginning  on  top  of  head  a  little  in  front  of  plane  of  ears  and  ex- 
tending posteriorly  to  middle  part  of  back. 

Cranial  characters. — Adult  male:  Skull  large,  vault  of  cranium 
fairly  high  but  not  arched ;  frontal  shield  broad,  rather  flat,  broadly 
sulcate  or  concave  medially,  rather  short  pointed  posteriorly,  with 
large,  rounded,  broadly  outstanding  postorbital  processes;  fronto- 
nasal region  moderately  dished;  rostrum  broad  and  short,  in  some 
skulls  slightly  depressed;  sagittal  crest  of  medium  length,  rather 
high  and  nearly  straight;  zygomata  rather  broadly  outstanding  and 
strongly  outbowed;  squamosal  root  expanded  in  old  age;  squamosal 
shelf  broad ;  braincase  narrow  anteriorly,  tending  to  keel  into  sagittal 
crest;  nares  small. 

Note. — The  type  skull  of  dalli  (No.  75048,  $  old)  is  abnormal: 
abnormally  large,  abnormally  high  (vault  of  cranium  abnormally 
arched) ;  fronto-nasal  region  abnormally  elevated — not  dished  as 
usual;  and  underjaw  abnormally  long.  No.  75047  (old  $  )  and  No. 
210293  (adult  $  )  are  far  more  typical. 

1  Named  for  William  H.  Dall,  of  the  Smithsonian  Institution. 


1918.]  DALLI  GROUP.  117 

Adult  females  (No.  140085,  from  Copper  River  delta,  July,  1905, 
A.  G.  Maddren;  and  old  female,  No.  210308,  from  Bering  Lake, 
1915,  J.  L.  Hill) :  Size,  medium;  skull  broad  and  short  for  a  female; 
moderately  arched,  and  moderately  dished,  with  rather  broad  frontal 
shield  and  broad  rostrum.  Frontal  shield  rather  broad,  strongly 
sulcate  medially,  swollen  over  orbits;  postorbital  processes  rather 
weak  and  strongly  decurved ;  point  of  shield  ending  at  f  ronto-parietal 
suture;  fronto-nasal  region  strongly  sloping,  rostrum  large,  broad, 
and  rather  short;  sagittal  crest  confined  to  parietals,  high  for  a 
female;  postpalatal  shelf  medium  or  broad,  strongly  rounded  on 
sides;  zygomata  moderately  spreading,  subtriangular,  the  posterior 
base  somewhat  bowed  and  vertically  expanded;  under  jaw  rather 
short ;  coronoid  high,  falcate,  the  apex  cutting  or  overreaching  plane 
of  condyle;  teeth  medium;  heel  of  M>  rather  long,  slightly  emar- 
ginate  on  outer  side,  the  extreme  tip  with  tendency  to  turn  outward. 

Cranial  comparisons. — Adult  and  old  males  (Nos.  75047  and 
210293)  compared  with  adult  male  nuchek  (No.  146459,  type) :  Size 
about  the  same  (basal  length  essentially  same,  but  occipito-nasal 
length  decidedly  less) ;  frontal  shield  broader  and  more  acutely 
pointed ;  vault  of  cranium  less  high  and  more  nearly  horizontal ;  post- 
orbitals  more  outstanding;  rostrum  slightly  shorter  and  more  de- 
pressed (appearing  broader) ;  zygomata  more  outbowed  (less  tri- 
angular) ;  palate  somewhat  shorter;  mastoids  more  appressed,  closer 
to  glenoid  processes,  constricting  meatus  tube  (in  nuchek  more  out- 
standing, leaving  wide  postglenoid  space  with  correspondingly  large 
open  meatus) ;  coronoid  blade  broader  above  (less  falcate).  Canines 
almost  the  same;  molars  decidedly  smaller  and  less  massive  and  in 
details  quite  different  (as  stated  under  nuchek). 

Adult  female  (No.  140085,  from  Copper  River  delta)  compared 
with  female  nuchek  (No.  44049,  from  near  Mount  St.  Elias) :  Skulls 
so  different  as  not  to  require  close  comparison,  that  of  dalli  being 
massive,  broadly  arched  or  domed,  and  with  massive  underjaw,  while 
that  of  nuchek  is  light,  slender,  and  narrow,  with  low  narrow  flat- 
tened frontal  region,  long  slender  rostrum,  and  light  underjaw.  The 
teeth  also  differ  strikingly. 

Adult  and  old  male  compared  with  old  male  cressonus  (type)  : 
Size  about  the  same;  vault  of  cranium  and  frontal  shield  much  less 
elevated,  flatter,  more  nearly  horizontal,  and  much  less  swollen  over 
orbits;  shield  less  deeply  sulcate;  postorbitals  larger  and  more  hori- 
zontally outstanding  (in  cressonus  weak  and  decurved) ;  fronto-nasal 
region  less  strongly  dished;  rostrum  broader  and  shorter;  zygomata 
much  more  widely  outstanding  and  much  more  bowed;  palate 
shorter;  molars  very  much  smaller. 


118  NORTH   AMERICAN   FAUNA.  [No.  41. 

Adult  and  old  female  compared  with  old  female  cressonus:  Skull 
larger,  broader,  and  more  highly  arched ;  frontal  shield  broader,  less 
deeply  sulcate  and  less  swollen  over  orbits ;  f ronto-nasal  region  more 
strongly  dished ;  rostrum  smaller  and  lower ;  zygomata  less  outstand- 
ing, less  arched,  much  less  expanded  vertically  and  more  sharply 
angular;  underjaw  much  shorter  and  lighter;  coronoid  smaller  and 
lower.    Teeth  smaller  throughout. 

Adult  and  old  male  compared  with  adult  and  old  male  kenaiensis: 
Size  the  same  or  somewhat  smaller ;  skull  much  less  massive ;  braincase 
conspicuously  narrower ;  frontal  shield  interorbitally  narrower,  shorter 
pointed  posteriorly;  f ronto-nasal  region  normally  more  strongly 
dished ;  postorbital  processes  less  broadly  rounded ;  rostrum  less  mas- 
sive and  less  elevated ;  sagittal  crest  much  longer ;  occipito-sphenoid 
shorter ;  mastoids  usually  less  outstanding. 

Skull  measurements. — Old  male  (No.  75047,  from  Yakutat  Bay, 
Alaska)  :  Basal  length,  345  ;x  occipito-nasal  length,  342 ;  palatal  length, 
190;  zygomatic  breadth,  263;  interorbital  breadth,  91.5.  Old  male 
(No.  210293,  from  Malaspina  Glacier)  :  Basal  length,  345;  occipito- 
nasal  length.  338;  palatal  length,  188;  z}Tgomatic  breadth,  248;  inter- 
orbital breadth,  96. 

URSUS  HOOTS'  Merbiam. 
Stikine  Bbown  Beab. 

Vrsus  hoots  Merrlam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  140-141,  Sep- 
tember 6,  1916. 

Type  locality. — Clearwater  Creek,  a  north  branch  of  Stikine  River, 
British  Columbia. 

Type  specimen. — No.  206136,  $  ad.,  U.  S.  National  Museum, 
Biological  Survey  collection.  Collected  by  John  Hyland;  presented 
by  Lincoln  Ellsworth. 

Cranial  characters. — Size  medium  or  large;  skull  massive;  slightly 
dished,  rather  short,  flattish  on  top,  very  broad  across  frontals  and 
rostrum.  Frontal  shield  broad,  nearly  flat,  long  pointed;  broadly 
and  shallowly  sulcate  medially  as  far  back  as  posterior  plane  of  post- 
orbitals;  postorbitals  large,  broad,  and  horizontally  outstanding; 
f ronto-nasal  region  sloping ;  rostrum  broad  and  rather  high ;  palate 
and  postpalatal  shelf  broad;  postpalatal  notch  moderate;  sagittal 
crest  short,  ending  at  fronto-parietal  suture;  zygomata  moderately 
outbowed,  not  broadly  spreading;  underjaw  rather  massive;  ramus 
broad  vertically,  its  inferior  border  upcurved  posteriorly;  coronoid 
blade  broad  at  base,  rather  high  and  subfalcate,  the  apex  curving 
strongly  backward,  cutting  plane  of  condyle;  dentition  remarkably 

1  Restored. 

*  Hoots,  the  native  Indian  name  for  the  big  brown  and  grizzly  bears  of  tbe  coast  region. 


1918.]  DALLI  GROUP.  119 

light  for  so  large  a  skull ;  canines  and  molars  (both  upper  and  lower) 
surprisingly  small. 

Remarks. — Ursus  hoots  does  not  appear  to  be  related  to  any  of 
the  other  mainland  species  except  the  newly  discovered  crassus  from 
the  mountains  on  the  upper  Macmillan  River,  Yukon,  from  which 
it  may  be  distinguished  at  a  glance  by  the  small  size  of  its  molar 
teeth.  It  may  be  related  also  to  sitkensis  of  Baranof  and  Chi- 
chagof  Islands,  but  differs  in  somewhat  smaller  size;  less  elevated 
posterior  frontal  region;  broader  postorbital  processes;  less  broadly 
spreading  zygomata ;  shorter  and  less  spreading  mastoids ;  less  nearly 
vertical  and  more  strongly  recurved  coronoid  blade,  the  apex  over- 
arching a  well-defined  coronoid  notch;  smaller  molars  (both  upper 
and  lower) ;  and  smaller  upper  incisors.  The  large  lower  premolar 
has  the  upturned  heel  of  the  Sitka  bear,  but  lacks  the  posterior  sulcus 
and  pair  of  cusplets  of  the  grizzlies.  Two  additional  old  male  skulls 
of  hoots  have  been  recently  received  from  the  Stikine  River  region. 
One  of  these  (No.  224841)  from  Clearwater  branch  of  Stikine — the 
type  locality — agrees  closely  with  the  type  specimen  in  size  and  form, 
but  has  slightly  larger  and  especially  broader  molars  (difference 
most  marked  in  M  ).  The  skull  is  older  than  that  of  the  type  and 
the  mastoids  are  more  strongly  developed  and  divergent.  The  other 
(No.  224839)  is  still  older  and  was  killed  low  down  the  Stikine.  It 
is  of  approximately  the  same  size  as  the  type,  but  considerably  older ; 
the  frontal  shield  is  more  broadly  concave  interorbitally  and  shorter 
pointed  posteriorly;  sagittal  crest  more  strongly  developed;  squa- 
mosal base  of  zygoma  much  more  broadly  expanded;  mastoids  much 
longer  and  strongly  divergent;  postpalatal  shelf  flatter;  coronoid 
blade  broader  at  and  above  middle ;  molars  slightly  larger.  An  adult 
female  from  the  type  locality  (No.  180883,  from  Clearwater  branch 
of  Stikine)  resembles  the  male  type  very  closely  except  for  the 
smaller  size  of  skull  and  teeth,  and  therefore  needs  no  special  descrip- 
tion. A  feature  deserving  mention  is  that  in  both  sexes  the  point 
of  the  frontal  shield  ends  at  the  fronto-parietal  suture. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  333;  oc- 
cipito-nasal  length,  325;  palatal  length,  179;  zygomatic  breadth,  228; 
interorbital  breadth,  96. 

URSUS  SITKENSIS  Merbiam. 

Sitka  Bbown  Bear. 

Ursus  sitkensis  Merriam,  Proc.  Biol.  Soc.  Washington,  X,  p.  73,  April  13,  1896. 

Type  locality. — Sitka  Islands,  Alaska. 

Type  specimen. — No.  187891,  $  ad.,  U.  S.  National  Museum  (No. 
6543,  Merriam  collection).  Collected  by  an  Indian;  purchased  at 
Sitka,  Alaska,  and  presented  to  C.  Hart  Merriam  by  J.  Stanley- 
Brown. 


120  NORTH   AMERICAN   FAUNA.  [No.  41. 

Range. — Sitka  Islands  (Baranof  and  Chichagof),  Alaska. 

Characters. — Size  large;  coloration  very  dark;  claws  of  moderate 
length,  curved,  dark  blue-black,  scurfy ;  skull  broad  and  massive. 

Color. — Dark;  muzzle  dark  brown,  sometimes  chocolate  brown  or 
even  sooty,  paler  in  faded  summer  pelage ;  head  and  body  very  dark 
brown  or  even  dusky,  varying  to  dull  brown  in  summer,  washed  on 
back  of  head,  neck,  and  shoulders  with  yellowish  or  golden. 

Cranial  characters. — Adult  male:  Skull  large,  massive,  dished, 
vault  moderately  elevated,  frontal  shield  (normally)  broad,  strongly 
sloping,  sulcate  medially ;  postorbitals  outstanding,  broadly  rounded ; 
rostrum  normally  rather  short  and  somewhat  depressed;  sagittal 
crest  massive,  straight;  zygomata  broadly  outstanding  and  out- 
bowed  ;  palate  moderate ;  postpalatal  shelf  normally  rather  long  and 
of  medium  breadth ;  notch  moderate  or  rather  narrow ;  basioccipital 
broad ;  mastoids  long  and  spreading. 

Cranial  comparisons. — Adult  and  old  males  compared  with  adult 
and  old  male  dalli:  Frontal  shield  higher  posteriorly  (in  dalli  highest 
at  or  immediately  behind  postorbitals),  flatter  and  more  sloping  at 
base  of  postorbitals,  thus  tilting  postorbital  plane  forward  (in  dalli 
looking  more  directly  upward) ;  a  distinct  thickening  or  hump  pres- 
ent on  each  side  of  median  sulcus  behind  plane  of  postorbitals  (lack- 
ing in  dalli) ;  basioccipital  usually  broader;  inion  less  strongly  de- 
veloped; mastoids  longer,  usually  more  outspreading  and  more  dis- 
tant from  glenoid  processes,  leaving  broader  space  for  audital  canal ; 
auditory  meatus  larger;  rostrum  somewhat  broader  basally;  dis- 
tance from  last  lower  molar  to  middle  of  condyle  usually  less;  last 
upper  molar  somewhat  longer  and  more  nearly  rectangular,  the  outer 
side  of  heel  less  oblique ;  MT  normally  with  open  saddle  between  pos- 
terior and  anterior  parts  (in  dalli  a  cusplet  occupies  the  inner  side  of 
the  saddle) ;  PM¥  in  sitkensis  normally  tricuspidate  as  seen  in  profile, 
there  being  both  anterior  and  posterior  cusplets  on  the  cingulum  (in 
dalli  the  anterior  and  posterior  cusplets  are  absent  and  the  main  cusp 
is  larger  and  higher  and  slopes  posteriorly  without  horizontal  heel). 

Skull  measurements. — Average  of  2  adult  males  from  Chichagof 
Island:  Basal  length,  358.5;  occipito-nasal  length,  354.5;  palatal 
length,  189;  zygomatic  breadth,  260;  interorbital  breadth,  93.5. 

URSUS  SHIRASI1  Merbiam. 

Shieas  Brown  Beab. 

(PL  VI.) 

Uraus  shirati  Merriam,  Proa  BloL  Soc.  Washington,  XXVII,  p.  195,  August 
13,  1914. 

Type  locality. — Pybus  Bay,  Admiralty  Island,  Alaska. 

1  Named  for  George  Shiras,  4th,  who  collected  and  presented  the  specimen. 


1918.]  DALLI   GROUP.  121 

Type  specimen. — No.  203030,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Killed  September  16,  1913,  by  George 
Shiras,  4th,  and  presented  to  the  Biological  Survey. 

Range. — Restricted  to  Admiralty  Island. 

Characters. — A  huge  brown  bear  larger  than  the  largest  sit- 
kensis;  head  highly  arched;  color  black,  except  muzzle,  which  is 
dull  brown ;  claws  dark  blue- black,  dull,  slightly  scurfy  (not  smoothly 
polished  as  in  the  grizzlies),  rather  strongly  curved  and  of  mod- 
erate length  (middle  claw  over  curve,  92  mm.;  from  top  of  base  to 
apex,  75),  fourth  and  fifth  rounded  off  on  outer  side. 

Color. — Entire  animal,  except  muzzle,  coal  black,  showing  when 
examined  closely  a  brownish  wash  along  middle  of  back ;  muzzle  from 
nose  pad  to  between  eyes  dull  brown. 

Cranial  characters. — Old  male  (type)  :  Skull  large,  broad,  massive, 
strongly  dished,  and  highly  arched;  zygomata  large,  broadly  out- 
bowed  and  rounded  anteriorly  as  well  as  posteriorly;  frontal  shield 
remarkably  short  and  broad  (nearly  twice  as  broad  as  long),  deeply 
and  broadly  concave  in  cross  section,  with  huge  uplifted1  broadly 
outstanding  postorbital  processes  which  arch  over  the  orbits  and  are 
strongly  decurved  apically,  completely  roofing  the  orbits;  temporal 
ridges  beaded,  short,  meeting  far  forward  (at  least  25  mm.  in  front 
of  fronto-parietal  suture)  ;  sagittal  crest  long  and  high,  humped 
over  fronto-parietal  suture;  fronto-nasal  region  strongly  concave; 
rostrum  broad  and  short,  rising  strongly  to  meet  frontal  shield; 
palate  broad;  basioccipital  and  basisphenoid  subequal;  ramus 
strongly  bellied  under  posterior  molars;  coronoid  blade  broad  and 
high. 

Dental  characters. — Dentition  heavy;  canines  large,  the  upper  47 
mm.  high  above  enamel  line  of  outer  side;  molars  large  and  rather 
broad;  M1  with  large  and  broad  heel;  MT  with  strongly  developed 
cusplet  on  inner  side  of  saddle. 

Remarks. — Ursus  shirasi  is  a  very  large  member  of  the  brown 
bear  group.  Whether  it  is  always  black,  like  the  type  specimen,  is 
not  known.  But  of  all  the  American  bears  its  skull  is  the  most 
striking  and  distinctive.  The  short  broad  frontal  shield  rising  on 
each  side  into  huge  postorbital  processes,  which  arch  broadly  over 
the  orbits,  serve  to  distinguish  it  at  a  glance  from  all  other  species, 
rendering  close  comparisons  unnecessary.  In  this  connection  it  is 
interesting  to  observe  that  shirasi  and  its  neighbor  eulophus,  an  in- 
habitant of  the  same  island,  present  opposite  extremes  of  departure 
from  the  normal  ursine  type — eulophus  having  a  long  narrow  skull 
with  slender  elongate  rostrum,  long  and  narrow  frontal  shield,  and 

1  Additional  skulls  of  adult  males  recently  received  have  the  postorbitals  equally  large 
but  less  uplifted,  not  rising  above  frontal  plane. 


122  NORTH   AMERICAN   FAUNA.  [No.  41. 

insignificant  postorbital  processes,  while  shirasi  has  an  exceptionally 
broad  skull  with  broad  short  rostrum,  excessively  broad  and  short 
frontal  shield,  and  huge  massive  postorbital  processes. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  355;  oc- 
cipito-nasal  length,  348;  palatal  length,  191;  zygomatic  breadth, 
259;  interorbital  breadth,  104.5. 

URSUS  NUCHEK  Mereiam.1 

Nuchek  Brown  Bear. 

Ursus  nuchek  Merriam,  Proc.  Biol.  Soc.  Washington,  XXIX,  pp.  146-148,    Sep- 
tember 6,  1916. 

Type  locality. — Head  of  Nuchek  Bay,  Hinchinbrook  Island,  Prince 
William  Sound,  Alaska. 

Type  specitnen. — No.  146459,  $  old,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.  Collected  September  15, 1905,  by  C.  Swan- 
son. 

Range. — Prince  William  Sound  easterly  to  Mount  St.  Elias ;  limits 
unknown. 

Characters. — Size  large;  external  characters  unknown;  skull  long, 
narrow,  and  moderately  high ;  molars  peculiar. 

Cranial  characters. — Adult  male  (type)  :  Large,  elongate;  frontal 
shield  relatively  narrow,  flattish,  moderately  depressed  between 
orbits;  orbital  rims  thickened;  postorbital  processes  broad  and  flat- 
tish, moderately  outstanding;  posterior  part  of  shield  broad,  ending 
about  two-thirds  distance  from  plane  of  postorbitals  to  fronto- 
parietal suture ;  sagittal  crest  rather  long,  straight,  high  posteriorly ; 
rostrum  long,  high,  rather  narrow ;  f ronto-nasal  region  sloping  in 
facial  plane;  nasals  slightly  elevated  anteriorly;  zygomata  moder- 
ately spreading,  subtriangular,  not  much  expanded  vertically;  post- 
palatal  shelf  moderate,  its  sides  rounded;  notch  long  and  narrow; 
anterior  nares  small;  meatus  tube  short  and  large.  Underjaw  mas- 
sive; coronoid  blade  narrow  and  falcate.  Teeth  of  medium  size; 
molars  broad  (more  massive  than  in  dalli) ;  last  upper  molar  excep- 
tionally short,  broadest  in  middle,  heel  short  and  obliquely  truncate 
on  outer  side ;  M1  large,  much  broader  posteriorly  than  anteriorly ; 
middle  lower  molar  peculiar:  twin  cusps  of  entoconid  very  small, 
low,  and  close  together;  main  cusp  of  inner  side  large  and  high,  re- 
ducing the  posterior  moiety  of  the  tooth  to  about  a  third  the  length 
of  the  crown  instead  of  about  half  as  usual. 

Young -adult  female  (No.  44049,  from  Chaix  Hills  near  Mount  St. 
Elias,  Alaska;  killed  July  4,  1891,  by  the  late  Prof.  I.  C.  Russell) : 
Skull  long,  narrow,  rather  low,  with  narrow  frontals,  narrow  ros- 

*  Tentatively  included  in  dalli  group.      (See  Introduction,  pp.  12-13.) 


1918.]  DALLI   GROUP.  123 

trum,  and  moderately  outstanding  subtriangular  zygomata.  Frontal 
shield  flattish,  medially  depressed  interorbitally,  sloping  gradually 
into  rostrum,  rather  short  pointed  posteriorly  (ending  about  15  mm. 
in  front  of  parietals;  in  fully  adult  and  old  females  it  would  be 
still  shorter) ;  postorbital  processes  moderate,  horizontally  outstand- 
ing, the  tips  rounded  (not  fully  grown) ;  palate  concave,  postpalatal 
shelf  rather  long  and  broad;  notch  rather  narrow;  basisphenoid 
strongly  concave  antero-posteriorly  and  transversely;  underjaw  long 
and  slender.  Canines  long  and  slender ;  molars  and  large  premolars 
with  rather  high  cusps ;  last  upper  molar  short,  much  broader  in  mid- 
dle than  anteriorly,  heel  short  and  obliquely  truncate  on  outer  side. 

Cranial  comparisons. — Ursus  nuchek  evidently  overlaps  the  range 
of  dalli  and  may  come  in  contact  with  rressonus,  necessitating  com- 
parisons with  both. 

Adult  male  (type)  compared  with  adult  and  old  male  dalli  (Nos. 
75047  and  210293)  :  Size  about  the  same;  basal  length  essentially  the 
same,  but  occipito-nasal  length  decidedly  greater;  skull  appearing 
longer  and  narrower;  more  elevated  behind  orbits  and  much  more 
strongly  sloping  posteriorly;  frontal  shield  narrower,  the  point 
broader  posteriorly;  vault  of  cranium  higher  and  less  nearly  hori- 
zontal ;  postorbitals  less  outstanding ;  f  ronto-nasal  region  less  dished ; 
rostrum  longer  and  not  depressed  (appearing  narrower)  ;  zygomata 
less  outbowed  (more  triangular)  ;  palate  somewhat  longer;  post- 
palatal  notch  longer  and  narrower;  mastoids  less  appressed,  leaving 
wide  postglenoid  space  with  correspondingly  large  open  meatus  (in 
dalli  closer  to  glenoid  process,  pressing  on  and  contracting  meatus 
tubes);  coronoid  blade  narrower  above  (more  falcate).  Canines 
about  the  same ;  molars,  both  upper  and  lower,  decidedly  larger  and 
more  massive  and  in  details  quite  different :  M^  exceptionally  short 
and  much  broader  in  middle  than  elsewhere,  the  heel  short  and 
obliquely  truncate  on  outer  side;  M1  large,  much  broader  posteriorly 
than  anteriorly;  middle  lower  molar  peculiar,  the  twin  cusps  of 
entoconid  very  small,  low,  and  close  together;  metaconid  exception- 
ally large  and  high,  reducing  the  posterior  moiety  of  the  crown  to 
about  one-third  its  length,  instead  of  about  half  as  in  dalli  and  most 
species. 

Young-adult  female  (No.  44049,  from  near  Mount  St.  Elias)  com- 
pared with  adult  female  dalli  (No.  140085,  from  Copper  Eiver 
delta)  :  Skulls  so  strikingly  different  as  not  to  require  close  compari- 
son, that  of  nuchek  being  light,  slender,  narrow,  with  low  narrow 
flattened  frontal  region,  long  slender  rostrum,  and  light  underjaw, 
while  that  of  female  dalli  is  massive,  broadly  arched  or  domed,  and 
with  massive  underjaw.    The  teeth  also  differ  strikingly. 


124  NORTH  AMERICAN   FAUNA.  INo.  41. 

Young-adult  female  (No.  44049)  contrasted  with  old  female  cres- 
sonus  (No.  209881)  :  Size  slightly  smaller  (when  fully  adult  probably 
the  same) ;  frontal  shield  much  narrower  and  flatter,  much  less 
deeply  sulcate,  much  less  swollen  over  orbits,  and  much  shorter  pos- 
teriorly; fronto-nasal  region  in  same  plane  (in  2  cressonus  strongly 
dished);  sagittal  crest  longer;  nares  smaller;  canines  (upper  and 
lower)  much  longer;  molars  more  massive;  M^  extremely  short, 
much  the  broadest  in  middle,  with  short  obliquely  truncate  heel  (in 
cressonus  normal). 

Female  (No.  44049)  compared  with  female  kenaiensis  (No. 
133244)  :  Basal  length  essentially  the  same;  cranium  narrower,  with 
narrower  braincase,  narrower  shield,  and  narrower  rostrum;  zygo- 
mata less  broadly  spreading  (would  be  more  broadly  spreading  with 
age)  ;  occipito-sphenoid  shorter;  palate  essentially  same  length  but 
narrower;  postpalatal  shelf  narrower;  ramus  more  slender  (con- 
spicuously thinner  below  M-y  and  M7) ;  its  inferior  border  straighter, 
less  upcurved  posteriorly;  coronoid  lower  and  less  narrowed  above; 
cusps  of  larger  premolars  above  and  below  much  more  highly  de- 
veloped ;  main  cusp  of  upper  premolars  very  much  higher  relative  to 
posterior  cusp ;  molar  cusps  also  more  strongly  developed ;  last  upper 
molar  shorter  and  of  peculiar  form,  as  in  the  male. 

Skull  measurements. — Old  male  (type)  :  Basal  length,  360  ;*  occi- 
pito-nasal  length,  358;  palatal  length,  191;  zygomatic  breadth,  248; 
interorbital  breadth,  88. 

Gyas  Group. 

URSUS  GYAS  Mebbiam. 

Peninsula  Giant  Beab. 

(Pi.  II.) 

Ursus  dalli  gyas  Merriam,  Proc.  Biol.  Soc.  Washington,  XV,  p.  78,  March  22, 

1902. 
Ursus  merriami  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XVI,  p.  141,  April  12,  1902. 

Type  locality. — Pavlof  Bay,  Alaska  Peninsula. 

Type  specimen. — No.  91669,  $  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection. 

Range. — Entire  length  of  Alaska  Peninsula  from  Cook  Inlet  to 
Isanotski  Strait  and  adjacent  Unimak  Island. 

Characters. — Size  huge,  either  largest  living  bear  or  second  only  to 
the  great  Kadiak  bear  {middendorffi) .  Claws  rather  long  and 
smooth,  dark  when  young,  pale  when  old.  Color  variable,  from 
grizzled  brown  to  pale  yellowish.  Skull  of  male  large,  long,  and 
massive,  but  not  highly  arched.    Sexual  disparity  great. 

1  Restored. 


1918.]  GYAS  GROUP.  125 

Cranial  characters. — Adult  males :  Skull  large,  long,  and  massive ; 
frontal  region  moderately  elevated,  sloping  gradually  into  rostrum, 
strongly  depressed  or  troughed  medially;  slightly  swollen  over  or- 
bits and  bases  of  postorbitals ;  postorbitals  rather  large,  subtriangu- 
lar,  moderately  outstanding;  zygomata  moderately  outstanding  and 
bowed;  palate  long  and  relatively  narrow.  Underjaw  long  and 
massive;  coronoid  blade  very  broad  basally.  Molars  light  for  so 
large  a  skull.  Viewed  from  behind,  the  posterior  frontal  region,  with 
its  depressed  median  trough  and  massive,  outstanding,  and  elevated 
postorbital  processes,  suggests  the  spread  wings  of  a  bat  or  a 
butterfly. 

Skulls  of  adult  males  which  it  seems  necessary  to  call  gyas  present 
a  surprisingly  wide  range  in  size  and  form.  Among  them  are  three 
quite  different  types  which  if  isolated  would  undoubtedly  develop 
into  very  distinct  species. 

1.  Typical  form,  with  large  elongate  skull. — Skull  and  teeth  large  and  mas- 
sive ;  zygomata  widely  spreading ;  sagittal  crest  high ;  f rontals  moderately  ele- 
vated, broadly  sulcate  medially,  swollen  laterally ;  postorbital  processes  thick, 
massive,  subtriangular,  and  decurved ;  nasal  opening  and  rostrum  proportionate 
to  size  of  skull  (but  anterior  nares  very  much  smaller  than  in  middendorffl). 
Teeth  large;  lower  canines  averaging  about  19  mm.  in  diameter  at  base  of 
enamel.  This  large  form  ranges  over  the  entire  length  of  Alaska  Peninsula 
from  Cook  Inlet  (where  it  was  obtained  at  Chinitna  by  Kidder  and  Blake)  to 
Morzhovoi  Bay  and  Unimak  Island.  Departures  from  normal:  The  series  of 
skulls  at  hand  shows  two  prominent  departures  from  typical  gyas — one  larger 
and  more  massive,  the  other  smaller  and  lighter. 

2.  Giant  form,  with  exceptionally  broad  rostrum. — Huge  skulls  with  broad 
massive  rostrum  and  exceedingly  heavy  jaws.  This  type  is  represented  in  the 
Biological  Survey  collection  by  two  specimens,  No.  91694  from  Cold  Bay,  and 
No.  91704  from  Belkofski  Bay.  The  entire  skull  is  larger  and  more  massive,  but 
the  differences  are  most  pronounced  in  the  face  and  jaws.  The  frontal  shield 
and  postorbital  processes  are  not  broader  than  in  some  skulls  of  typical  gyas, 
but  the  postorbital  processes  are  abruptly  de flexed  at  the  tips,  forming  a  thick 
massive  hook  over  the  orbit,  much  as  in  middendorffi.  The  rostrum  is  extraor- 
dinarily broad  and  massive,  giving  the  skull,  viewed  from  the  front,  a  most 
peculiar  aspect.  The  occipital  flange  (lambdoid  crest)  also  is  largely  de- 
veloped. The  peculiarities  of  the  underjaw  are  as  striking  as  those  of  the 
rostrum.  The  inferior  part  of  the  ramus  is  greatly  thickened  anteriorly,  and 
the  posterior  half  is  strongly  everted,  forming  a  broad  lip,  unlike  anything 
seen  in  typical  gyas.  By  reason  of  this  peculiarity  the  jaws  flare  strongly  out- 
ward under  the  last  molars,  and  the  flaring  is  so  great  that  it  is  conspicuous 
even  when  looked  at  from  above. 

3.  Small  form,  with  narrow  rostrum. — Smaller  skulls,  with  narrower  ros- 
trum, more  slender,  horizontally  outstanding  postorbital  processes  and  much 
smaller  canines  (exemplified  by  Nos.  82003  and  82004  from  Pavlof  Bay,  and 
No.  91699  from  Belkofski  Bay).  Compared  with  typical  gyas,  the  skull  as  a 
whole  is  considerably  smaller,  shorter,  and  lighter ;  vault  of  cranium  rather 
more  flattened;  rostrum  decidedly  narrower;  canine  teeth  above  and  below 
decidedly  more  slender ;  last  upper  molar  narrower.  But  the  most  conspicuous 
difference  is  in  the  postorbital  processes,  which  instead  of  being  broadly  tri- 


126  NORTH   AMERICAN   FAUNA.  lNo.41. 

angular,  massive,  and  decurved,  are  elongate,  peglike,  and  stand  out  hori- 
zontally. 

The  above  description  applies  to  fully  adult  males  of  the  form  described  by 
Allen  under  the  name  Ursus  merriami,  the  type  of  which  was  an  immature 
male.  Unfortunately  for  merriami,  the  typical  form  seems  to  be  connected 
with  gyas  by  a  series  of  intergrades.  Thus,  skull  No.  862,  Field  Mus.  Nat. 
Hist.,  has  the  small  teeth  and  narrow  muzzle  of  merriami,  but  the  postorbital 
processes  are  broader  posteriorly  and  slightly  decurved;  and  Nos.  91691, 
147630,  91675,  U.  S.  Nat.  Mus.,  Biological  Survey  collection,  and  No.  4585,  Mus. 
Vert.  Zool.,  Univ.  California,  complete  the  chain  of  intergrades,  so  that  it  is 
difficult  to  tell  just  where  to  draw  a  line  between  them. 

Skulls  of  adult  males  of  the  small  form  have  been  examined  from  various 
localities  from  Cold  Bay  and  Ugashik  Lake  and  River  westerly  to  Pavlof, 
Belkofski,  Bear  Bay,  and  Tonki  Point. 

Adult  females:  Skull  of  moderate  size,  conspicuously  smaller  than 
male;  frontal  region  normally  elevated,  domed,  and  rounded  off,  the 
postorbital  processes  somewhat  decurved,  the  frontal  shield  sulcate 
medially.  Skulls  of  adult  females  differ  among  themselves  in  degree 
of  elevation  and  doming  of  frontal  shield,  depth  of  median  groove, 
relative  massiveness,  and  other  characters,  but  in  the  present  imper- 
fect state  of  knowledge  it  is  impossible  to  assign  positively  any  par- 
ticular female  to  either  of  the  above-described  types  of  males. 

Cranial  comparisons. — Old  male  compared  with  old  male  mid- 
dendorffi  of  essentially  same  size :  Vault  of  cranium  much  less  highly 
arched  and  never  domed;  postorbital  processes  more  strongly  de- 
veloped and  less  decurved ;  zygomata  less  widely  outstanding  and  far 
less  bowed  (ratio  of  zygomatic  breadth  to  basal  length  much  less) ; 
anterior  nares  normal,  not  flaring;  coronoid  blade  less  high.  Last 
upper  molar  shorter  and  of  different  form,  the  heel  obliquely  trun- 
cate on  outer  side,  narrowing  posteriorly ;  lower  molars  broader  and 
heavier. 

Skull  measurements. — Old  male  (type):  Basal  length,  380;  oc- 
cipito-nasal  length,  394 ;  palatal  length,  206 ;  zygomatic  breadth,  286 ; 
interorbital  breadth,  96. 

URSUS  MIDDENDORFFI  Mebbiam. 

Kadiak  Beab. 

(Pi.  III.) 

Ursus  middendorffl  Merrlam,  Proc.  Biol.  Soc.  Washington,  X,  pp.  67-69,  April 

13,  1896. 
Ursus  kadiaki  Kleinschmidt,  Outdoor  Life,  XXVII,  p.  3,  January,  1911. 

Type  locality. — Kodiak  Island,  Alaska. 

Type  specimen. — No.  54793,  $  young-adult,  U.  S.  National  Mu- 
seum, Biological  Survey  collection.  Collected  July  3,  1893,  by  B.  J. 
Bretherton  (original  No.  176). 

Range.— Kodiak  and  adjacent  islands,  Afognak  and  Shuyak;  not 
known  from  mainland. 


1918.]  KENAIENSIS  GROUP.  127 

Cranial  characters. — Size  huge;  skull  of  male  exceedingly  broad, 
high,  and  relatively  short;  frontal  shield  domed,  sulcate  medially 
and  swollen  over  orbits  (obliquely  flattened  in  extreme  age),  rather 
short  pointed  posteriorly,  passing  into  sagittal  crest  in  fully  adult 
skulls  anterior  to  fronto-parietal  suture;  fronto-nasal  region  dished; 
postorbitals  decurved  and  weak,  small  for  size  of  skull;  rostrum 
rather  short  and  of  medium  breadth;  anterior  nares  flaring;  zygo- 
mata extraordinarily  outstanding  and  strongly  outbowed ;  palate 
rather  broad;  postpalatal  shelf  rather  narrow;  mastoids  long  and 
divergent  Underjaw  large,  massive,  and  rather  short;  coronoid 
blade  high  and  moderately  recurved;  molars  small  for  size  of  skull. 

Cranial  comparisons. — Old  male  compared  with  old  male  gyas  of 
essentially  same  size:  Vault  of  cranium  much  more  highly  arched  and 
usually  domed;  postorbital  processes  weak  and  decurved;  zygomatic 
arches  much  more  widely  outstanding  and  far  more  strongly  bowed 
(ratio  of  zygomatic  breadth  to  basal  length  much  greater) :  anterior 
nares  strongly  flaring  (in  gyas  not  flaring) ;  coronoid  blade  higher. 
Last  upper  molar  with  heel  more  broadly  rounded;  lower  molars 
narrower. 

Flesh  measurements. — Young-adult  male  killed  by  J.  H.  Kidder 
on  Shuyak  Island,  off  Afognak,  Alaska,  July,  1901 :  Total  length, 
nose  to  end  of  tail  vertebrae,  8  ft.;  nose  to  base  of  tail.  7  ft.  8 %  in.; 
height  at  shoulders,  4  ft.  34  in.;  length  of  forefoot,  including  claws, 

I  ft.  2£  in.;  hind  foot,  1  ft.  4  in.;  width  of  forefoot,  84,  in.:  width  of 
hind  foot,  7|  in.;  length  of  fore  claws.  44,  in.;  girth  of  body  behind 
shoulders,  5  ft.  If  in.;  girth  of  neck,  3  ft.  2  in.;  girth  of  head  at  ears, 
3  ft.  If  in.;  nose  to  tip  of  hind  foot  (animal  lying  on  one  side),  9  ft. 

II  in.    The  body  after  the  skin  was  removed  was  the  size  of  a  big  ox. 
Skull  measurements. — No.    134407,   largest   of  the   males:   Basal 

length,  392;  occipito-nasal  length,  369;  palatal  length,  211;  zygomatic 
breadth,  306;  interorbital  breadth,  101. 

Kenaiensis  Group. 

URSUS  KENAIENSTS  Mekriam. 

Kenai  Giant  Beau. 

(PI.  IV.) 

Ursus  lenaiensis  Merriam,  Proc.  Biol.  Soc,  Washington,  XVII,  p.  154,  October  6, 
1904. 

Type  locality.— Cape  Elizabeth,  at  extreme  west  end  of  Kenai 
Peninsula,  Alaska. 

Type  specimen— No.  128672,  9  ad.,  U.  S.  National  Museum,  Bio- 
logical Survey  collection.    Collected  in  1903  by  C.  A.  Lambert. 

Range. — Kenai  Peninsula, 
C4854°— 18 9 


128  NORTH  AMERICAN  FAUNA.  [No.  41. 

Characters. — Size  large;  appearance  that  of  a  big  grizzly;  colora- 
tion rather  dark;  claws  moderately  curved,  dark,  usually  marked 
with  whitish  streaks  on  sides  and  near  tip;  longest  claw  in  three 
adults  82-90  mm.  Skull  broad  and  massive,  that  of  male  strikingly 
larger  than  female. 

Color.— Old  male  killed  in  October,  1912,  by  Wilson  Potter,  of 
Philadelphia  (skull  1S1099)  :  Muzzle  pale  fulvous-brown;  cheeks 
and  forehead  similar  but  hairs  longer  and  with  pale  tips;  ground 
color  of  top  of  head,  neck,  and  back  much  darker  but  deeply  washed 
on  tips  with  buffy  or  buffy  whitish,  giving  these  parts  the  look  of  a 
grizzly;  legs  and  feet  (but  not  belly)  much  darker. 

Cranial  characters. — Adult  mate:  Skull  large,  broad,  and  massive, 
with  broad  frontal  shield  and  rostrum,  widely  outbowed  zygomata, 
large  outstanding  bluntly  rounded  postorbitals,  and  small  anterior 
nares.  Frontals  broad  throughout  (interorbitally,  postorbitally,  and 
across  postorbital  processes) ;  frontal  shield  well  elevated  above  plane 
of  rostrum,  shallowly  sulcate  medially  between  orbits,  slightly  swollen 
over  orbits,  long  pointed  posteriorly ;  sagittal  crest  short,  not  reach- 
ing anteriorly  beyond  f ronto-parietal  suture ;  braincase  broad  anteri- 
orly; rostrum  broad  throughout  but  much  broader  posteriorly  than 
anteriorly;  zygomata  abruptly  and  widely  outbowed,  the  squamosal 
root  vertically  expanded;  palate  very  broad;  underjaw  large  and 
massive,  coronoid  blade  large  and  high,  scarcely  recurved.  Canines 
small  and  short.  Adult  female:  Skull  broad,  flat,  and  massive,  with 
exceedingly  broad  rostrum,  broadening  posteriorly;  zygomata 
broadly  spreading;  jugal  broad  anteriorly;  frontals  flattened,  de- 
pressed, low  posteriorly ;  postorbital  processes  large,  blunt,  and  hori- 
zontally outstanding;  palate  exceedingly  broad;  nasals  large  and 
broad;  anterior  nares  small.    Canines  small. 

Cranial  comparisons. — Adult  male  compared  with  adult  male 
alescandrce :  Length  essentially  the  same,  but  henaiensis  much  broader 
and  more  massive  throughout;  frontal  shield  broader  throughout 
(interorbitally,  postorbitally,  and  across  postorbital  processes),  more 
elevated,  convex  (not  flat)  antero-posteriorly,  shallowly  sulcate  inter- 
orbitally, slightly  swollen  over  orbits;  longer  pointed  posteriorly, 
with  correspondingly  shorter  sagittal  crest  ending  anteriorly  at 
f ronto-parietal  suture;  postorbital  processes  more  massive  and  out- 
standing; frontal  part  of  braincase  not  keeled  or  compressed;  rostrum 
much  broader,  especially  posteriorly;  nasals  shorter  anteriorly;  zygo- 
mata much  more  widely  and  abruptly  spreading  posteriorly,  outbowed 
instead  of  angular;  squamosal  part  rising  abruptly  and  broadly  ex- 
panded vertically,  differing  strikingly  from  the  more  slender  and 
gently  curving  form  in  alexondrce;  palate  broader;  mastoids  more 
outstanding;  ramus  longer  and  more  massive;  coronoid  blade  more 


1918.]  KENAIENSIS  GROUP.  129 

nearly  vertical,  higher,  broader  in  upper  third,  less  recurved,  not 
ending  in  posterior  point.  Canines,  both  upper  and  lower,  smaller 
and  shorter. 

Adult  female  compared  with  adult  female  alexandrm:  Skull 
shorter,  both  basall y  and  on  top ;  vault  of  cranium  and  rostrum  lower ; 
fronto-nasal  region  more  dished;  rostrum  very  much  broader  basally, 
its  sides  sloping  anteriorly;  braincase  shorter  (less  occipital  over- 
hang) ;  sagittal  crest  lower;  occipito-sphenoid  and  base  of  skull 
shorter;  palate  broader;  ramus  more  massive.  Canines  smaller  (the 
lower  notably  more  slender). 

Adult  and  old  males  compared  with  adult  and  old  male  dalli: 
Size  the  same  or  somewhat  larger;  skull  much  more  massive;  brain- 
case  conspicuously  broader;  frontal  shield  interorbitally  broader, 
longer  pointed  posteriorly;  fronto-nasal  region  normally  less 
strongly  dished;  postorbital  processes  more  broadly  rounded;  ros- 
trum more  massive  and  more  elevated;  sagittal  crest  much  shorter; 
occipito-sphenoid  longer;  mastoids  usually  more  outstanding. 

Skull  measurements. — Average  of  2  old  males  from  Kenai  Pen- 
insula: Basal  length,  3G7;  occipito-nasal  length,  3G0;  palatal  length, 
197;  zygomatic  breadth,  2G3.5;  interorbital  breadth,  205.  Adult 
female  (type)  :  Basal  length, -288.5 ;  occipito-nasal  length,  285;  palatal 
length,  158;  zygomatic  breadth,  214;  interorbital  breadth,  78. 

UKSUS  SHELDONI  *  Mekkiaji. 

Montague  Island  Bear. 

<ri.v.) 

Ursus  slicldoni  Merriam,  Proc.   Biol.   Soc   Washington,  XXIII,  pp.   127-130, 
September  2,  1910. 

Type  locality. — Montague  Island,  Prince  "William  Sound,  Alaska. 

Type  specimen. — No.  137318,  $  young-adult,  U.  S.  National 
Museum,  Biological  Survey  collection.  Collected  May,  1905,  by 
Charles  Sheldon. 

Characters. — Size  large;  teeth  and  claws  of  the  grizzly  type;  color 
variable,  from  dark  to  light  brown.  Skull  broad  and  massive;  vault 
of  cranium  domed ;  hairs  over  shoulders  elongated  to  form  a  small 
but  distinct  hump. 

Color. — General  color  brownish,  varying  from  pale  to  dark,  the 
hairs  of  the  back  sometimes  yellowish  tipped,  those  of  the  head  griz- 
zled; color  darkest  (almost  blackish)  on  belly,  legs,  and  feet;  ears 
dark  with  whitish  tips.    An  old  she-bear  killed  by  Sheldon,  May  18, 

1  Named  for  Charles  Sheldon,  of  New  York,  who  collected  and  presented  the  type  and 
Other  specimens. 


130  NOBTH   AMERICAN   FAUNA.  I  No.  41. 

1905,  is  very  pale  grizzled  gray  on  the  upperparts,  and  only  moder- 
ately darker  on  the  legs  and  feet.  The  cub  of  this  bear,  killed 
the  same  day,  Avas  in  its  second  year  (about  1G  months  old)  and 
is  very  pale — almost  buffy  gray — with  dark  feet  and  legs,  and  a 
strongly  marked  hump. 

Cranial  characters. — Adult  male  (type)  :  Large,  massive,  excep- 
tionally broad,  Avith  broadly  outboAved  zygomata.  Frontal  shield 
long  and  broad,  nearly  horizontal  from  postorbitals  posteriorly  (act- 
ually sloping  downAvard  posteriorly) ;  broadly  sulcate  medially  be- 
tAveen  postorbital  processes,  moderately  sAvollen  on  each  side,  th?  pos- 
terior part  long  and  broad  (temporal  ridges  in  type  specimen  not 
curving  inward  as  in  most  skulls),  the  point  falling  over  middle  of 
parietals  (in  old  age  doubtless  more  anterior) ;  sagittal  crest  excep- 
tionally short,  confined  in  type  skull  to  posterior  half  of  parietals; 
fronto-nasal  region  elevated,  forming  a  convexity  instead  of  the  usual 
depression  b?tAveen  plane  of  rostrum  and  that  of  frontal  shield; 
rostrum  broad  and  short;  nasals  strongly  sloping,  dished  at  junction 
of  anterior  and  middle  thirds,  the  anterior  third  horizontal  or 
slightly  upturned ;  palate  and  postpalatal  shelf  very  broad :  notch 
rather  broad;  zygomata  broadly  outstanding,  rounded  posteriorly, 
someAvhat  bowed,  not  much  expanded  vertically  (doubtless  more 
broadly  expanded  in  old  age) ;  squamosal  shelves  exceptionally 
broad;  braincase  narroAvest  on  anterior  part  of  parietals  instead  of 
on  frontals.  Und^rjaAv  large  and  massive;  ramus  broad  vertically, 
broadest  under  posterior  molars;  coronoid  blade  elevated,  moderate, 
vertical,  the  apex  only  slightly  recurved.  Dentition  light;  canines 
and  molars  small  for  size  of  skull. 

Old  female  (No.  137316,  mother  of  cub  No.  137315)  from  Mon- 
tagu? Island,  May  18,  1905,  collected  and  presented  by  Charles  Shel- 
don :  Skull  of  medium  size,  broad,  flattish  on  top.  Frontal  shield  flat, 
nearly  horizontal,  the  posterior  part  broad  and  long,  reaching  past 
middle  of  parietals;  postorbital  processes  strongly  developed,  almost 
peglike,  horizontally  outstanding;  fronto-nasal  region  elevated  in 
fronto-facial  plane;  rostrum  short,  rather  broad;  palate  and  post- 
palatal  shelf  broad;  zygomata  broadly  outstanding,  subtriangular, 
rounded  posteriorly.  Underjaw  rather  massive;  coronoid  moderate, 
the  apex  only  slightly  recurved;  teeth  small  for  size  of  skull;  brain- 
case  very  broad. 

Cranial  comparisons. — Skull  in  general  similar  to  that  of  kenaien- 
sis  but  basisphenoid  broader  and  flatter,  its  length  nearly  equal  to 
that  of  basioccipital;  posterior  roots  of  interpterygoid  fossa  more 
Avidely  spreading;  condyle  of  jaw  more  exserted  (in  kenaiensis  ses- 
sile), reaching  so  far  back  that  a  line  dropped  from  peak  of  coronoid 
to  tip  of  angle  touches  or  traverses  it  (in  kenaiensis  this  line  passes 
freely  behind  the  condyle) ;  coronoid,  in  females  of  same  age,  smaller 


1018.]  GENUS  VETULARCTOS.  131 

and  lower,  its  area  for  muscular  attachment  less;  ramus  of  jaw 
strongly  bellied  posteriorly,  its  inferior  border  below  the  coronoid 
strongly  convex  downward  and  curving  evenly,  with  only  a  very 
slight  break,  to  angular  process.  (In  kenaiensis  the  inferior  border 
of  ramus  is  nearly  straight,  not  appreciably  bellied  under  coronoid, 
and  ends  abruptly  in  a  step  or  jog  at  some  distance  behind  the  angle.) 

In  general  form  and  appearance  skulls  of  females  closely  resemble 
those  of  female  kenaiensis,  differing  chiefly  in  the  characters  above 
mentioned  and  in  certain  dental  peculiarities,  notably  the  smaller 
size  and  more  pointed  heel  of  the  last  upper  molar,  and  the  oblique 
truncation  of  M^. 

Dental  characters. — Teeth  in  general  of  the  grizzly  type.  Last 
(fourth)  lower  premolar  normally  with  horizontal  heel,  slightly  up- 
turned at  posterior  end,  the  shallow  median  sulcus  reaching  from 
cusp  to  end  of  heel,  its  defining  ridges  ending  in  slightly  developed 
posterior  cusplets.  (In  kenaiensis  the  last  lower  premolar  is  more 
conical,  the  heel  sloping,  the  sulcus  incomplete,  with  only  a  single 
posterior  cusplet — on  inner  side  of  main  cusp  posteriorly.)  First 
upper  molar  peculiar,  having  both  ends  obliquely  truncate  and  paral- 
lei,  sloping  strongly  from  outer  angles  backward  and  inward;  in- 
ner row  of  cusps  pushed  back  so  that  each  falls  behind  plane  of 
corresponding  cusp  on  outer  side;  tooth  as  a  whole  more  nearly  rec- 
tangular, its  inner  corners  more  nearly  square  (less  rounded),  and 
inner  side  more  flattened  and  much  less  convex  than  in  kenaiensis. 

In  the  females  the  last  lower  molar  is  conspicuously  smaller  than  in 
kenaiensis,  and  the  last  upper  molar  is  smaller,  narrower,  more  wedge- 
shape,  and  more  pointed  posteriorly.  In  one  of  the  males  it  is  simi- 
lar. In  the  other  three  males  the  last  upper  molar  is  larger  and  less 
acute  posteriorly  than  in  the  females,  and  the  third  cusp  on  the  inner 
side  is  better  developed. 

Skull  measurements. — Adult  male  (type)  :  Basal  length,  Sr>9 ;  occip- 
ito-nasal  length,  315;  palatal  length,  198;  zygomatic  breadth,  270; 
intcrorbital  breadth,  102.5. 

VETULARCTOS,  A  NEW  GENUS  RELATED  TO  URSUS. 

Generic  characters. — Skull  like  that  of  Ursus.1  Teeth  in  the  main 
like  those  of  Ursus,  but  Mj  quite  different,  presenting  a  broad  flat 
squarish  grinding  surface  with  suppression  of  the  hypoconid  and 
entoconid,  and  absence  of  the  usual  posterior  cusp  and  notch  on 
outer  side.  M-  with  outer  cusps  (paracone  and  metacone)  normal ; 
inner  cusps  (protocone  and  hypocone)  obsolete.  Mj  with  anterior 
part  (protoconid  and  metaconid)  normal  but  rather  low;  posterior 

1  The  type  is  a  young-adult  female.     Skulls  of  adult  males  may  show  cranial  differences. 


132  NORTH   AMERICAN   FAUNA.  lNo.41. 

part  occupying  more  than  half  of  crown,  a  flat  rectangular  platform, 
truncate  posteriorly,  and  bordered  by  a  low  marginal  rim  bearing  a 
single  small  cusp  on  inner  side  immediately  behind  the  metaconid; 
hypoconid  absent ;  entoconid  obsolete ;  outer  side  of  tooth  nearly  flat, 
without  sulcus  or  reentrant  angle  and  without  notch  between  pro- 
toconid  and  hypoconid. 

In  the  reduction  or  suppression  of  the  inner  cusps  of  M1  and  Ma, 
and  of  both  inner  and  outer  cusps  of  the  large  posterior  moiety  of 
M^,  together  with  the  absence  of  a  reentrant  angle  and  notch  on 
the  outer  side  of  M7,  V etularctos  resembles  Arctotherium  and  Trem- 
arctos.  The  details  of  these  and  other  teeth  in  the  three  genera, 
however,  are  quite  different  and  ,ihe  genera  have  little  in  common. 
Nevertheless  the  resemblances  are  sufficient  to  suggest  that  V etularctos 
may  claim  a  rather  ancient  line  of  descent,  from  which  Arctotherium, 
and  Tremarctos  also  arose — a  line  quite  different  from  the  one 
culminating  in  Ursxis  proper. 

Type  species. — V etularctos  inopinatus. 

VETULARCTOS  INOPINATUS  sp.  NOV. 
Patriakchal  Beak 

Type,  skull  No.  7149  (skin  No.  870G),  9  nearly  adult,  U.  S. 
National  Museum,  from  Rendezvous  Lake,  northeast  of  Fort  Ander- 
son, Mackenzie.  Collected  June  24,  1864,  by  R.  MacFarlane.  Origi- 
nal No.  1979.    Teeth  practically  unworn. 

Color. — General  color  varying  from  whitish  buff  to  pale  yellowish 
buff  (yellowest  on  back  of  head  and  neck),  darkening  to  dull  reddish 
brown  on  ankles,  feet,  and  median  line  of  belly.  The  pale  body  color 
covers  the  entire  body  from  between  eyes  to  base  of  tail  and  reaches 
down  over  thighs  and  upper  part  of  legs.  Muzzle  golden  brown, 
becoming  dull  fulvous-brown  around  eyes;  top  of  head  from  be- 
tween eyes  posteriorly  soiled  buff;  long  hairs  of  cheeks  washed  with 
buffy;  ears  pale  buffy.  Fur  everywhere  full,  soft,  and  woolly; 
basal  fur  of  upperparts  varying  from  grayish  to  grayish  brown,  but 
distal  half  or  more  than  half,  pale  buffy,  so  the  animal  as  a  whole 
appears  to  be  buffy  whitish. 

Cranial  and  dental  characters. — Skull  small,  moderately  arched 
above;  basicranial  axis  arched,  palate  strongly  arched  and  slightly 
concave  longitudinally.  Teeth  peculiar,  presenting  a  combination  of 
long  canines  and  well-developed  cusps  for  seizing  (main  cusp  of 
PM-*  and  PM4,  outer  cusps  of  M-1-  and  M^  and  anterior  cusps  of  MT) 
with  broadly  flattened  surfaces  for  crushing  (in  M-3-,  M7,  and  M7). 
Incisors,  canines,  and  premolars  as  in  JJrsus;  posterior  molars 
peculiar,  showing  a  strong  tendency  toward  the  suppression  of  <*usps, 
particularly  those  of  the  inner  side.     Crown  of  MA  with  outer  cusps 


1918.1  GENUS  VETULAECTOS.  133 

(paracone  and  metacone)  normal;  inner  cusps  (protocone  and  hypo- 
cone — slightly  worn)  low,  scarcely  rising  above  level  of  median 
part  of  tooth ;  no  middle  cusplet.  Crown  of  M-2-  with  outer  cusps 
(paracone  and  metacone)  normal;  inner  cusps  (protocone  and 
hypocone)  obsolete,  represented  by  a  low  ridge  scarcely  projecting 
above  the  granular  surface  of  the  interior  of  the  crown.  Crown  of 
My  normal,  but  anterior  part  higher  than  posterior.  Crown  of 
M^  very  remarkable  (see  description  of  genus).  Crown  of  M,  flat, 
with  only  insignificant  marginal  thickenings  to  represent  protoconid 
and  metaccnid,  the  flat  crown  forming  a  direct  continuation  of  the 
large  crushing  platform  of  the  preceding  tooth. 

/Skull  measurements. — Young-adult  female  (type) :  Basal  length, 
268;  cccipito-nasal  length,  255;  palatal  length,  148;  zygomatic 
breadth,  181;  interorbital  breadth,  70. 

Tooth  measurements.— PU±  17  x  14;  M*,  22.5  x  17;  M^,  35.5  x  19; 
PMT,  14x8;  MT,24.5xl2;M7,  27x17;  M3,  21x16;  upper  molariform 
series,  73;  upper  molars,  59;  lower  molars,  72;  diameter  of  lower 
canine,  13;  upper  incisors  (series),  40. 


North  Arr  <:•  eai  U.  S.  Dept.  Aj 


Plate  II. 


3     fc 


North  American  Fauna  No.  41,  U.  S.  Dept.  Agr.      Biological  Survey. 


Plate  III. 


North  American  Fauna  No.  41.  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  IV. 


North  American  Fauna  No.  41,  U.  S.  Dept.  Agr.      Biological  Survey. 


Plate  V. 


North  American  Fauna  No.  41.  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  VI 


rr     o 
3     7\ 


X         33 

CO     3 


North  American  Fauna  No.  41,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  VII. 


North  American  Fauna  No.  41,  U.  S.  Dept.  Agr.      Biological  Survey. 


Plate  VIII. 


r>    2 


North  American  Fauna  No.  41,  U.  S,  Dept.  Agr.     Biological  Survey. 


Plate  IX. 


uj      g 


co     s3 


=>     25 


North  American  Fauna  No.  41,  U.  S.  Dept  Agr.     Biological  Survey. 


Plate  X. 


O     £. 


u-    S 


North  American  Fauna  No.  41,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  XI. 


o    £ 


North  Amerxan  Fauna  No.  41.  U.  S   Deot   Agr.     Biological  Survey.  PLATE  XII. 


UJ 

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P. 

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North  American  Fauna  No.  41,  U.  S.  Dept.  Agr      Biological  Survey. 


Plate  XIII. 


North  American  Fauna  No.  41,  U.  S    Dept.  Agr.     Biological  Survey 


Plate  XIV. 


North  American  Fauna  No.  41,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  XV. 


North  American  Fauna  No.  41,  U.  S,  Dept.  Agr.     Biological  Survey. 


Plate  XVI, 


2      £ 


INDEX. 


[New  species  in  bold-faced  type;  synonyms  in  italics;  principal  references  In  bold-faced 

figures.) 

A. 

absarokus,   18,   21,  41,  00,  CI,  C7,  85,  80, 

93-94,  103,  100. 
Acknowledgments.   10-11. 
alascensis.  94-05,  0G,  100,  10S.  110. 
alexandra?,  75.  Ill,  114-115,  128,  129. 
andersonl,  83-84. 
apache,  54.  84-85,  80. 
Arctotbei-ium,  genus,   132. 
arizona*.  53-54,  54,  G4,  00,  G7,  85. 
atuarko,  22-23. 

B. 

bairdi.  13.  18,  19-20,  21,  37  (footnote),  G3, 

04,  05,  60,  07,  70. 
blsonophiigus,  54,  60,  G7-CS. 

C. 

callfornicus.  29-30,  30,  31,  32,  73,  74,  75,  70. 
canadensis,  43,  52-53,  08,  103. 
candescent,  17. 
caurlnus,  0,  CS,  69-71. 
cbelan,  57,  58,  58-60,  78. 
chelidonias,  13,  21-22. 
cinereus,  17. 
Classification,  12-13. 
colusus,  30,  32-33,  74. 
Crama!  characters,  13. 
crassodon,  90,  01. 
crassos,  90-92,  119. 

cressonus,  111,  112,  112-113,  117,  118,  123, 
124. 

D. 

dalH,   9,   98,    301,   102,   113,    11G-118,   120, 

122,  123,  129. 
Danis,  86. 

Dental  characters,  13. 
dnsorgus,  33-34. 


eltonclarki,  41-43,  44,  45. 
creosotes,  55,  56,  57,  58. 
Euarctos,  04. 

eulophus.  9,  6!>.  70,  71-72,  121. 
eximius,  109-110. 


F. 


ferox,  17. 


Geographic  distribution,  9-10. 
grisevs,  17. 
Groups,  12-13. 

absarokus.  10. 

alascensis.  94-99. 

arizona?,,  53-76. 

arizona?-utahensis.  Gl. 

californicus-kwakiutl,  25. 

chelan.  10. 

dalli,  116-124. 

g.vas,  124-127. 

horria>us,  84-88. 

horribilis,  10.  13.  17-34. 

bylodromus.  77-84. 

innuitus   110-115. 

kenaiensis.    127-131. 

kidderi.   106-110. 

planiceps.  34-53. 

richardsoni.  99-100. 

shoshone,  57. 

shoshone-idahoensis,  73. 

stikeenensis.  88-94. 

tahltanicus,  10,  42. 

townsendi,    115-116. 
gyas,  106,  107,  108,  124-126,  127. 


64S540— 1S- 


-10 


henshawl.  31,  73,  87-88. 

hoots,  44,  91,  118-119. 

horr,aceu8,  80. 

borriaeus,   7,  35,  36,  61,  62,  84,  85,  85-87, 

87,  104. 
horribllis,  7,  33,  17-19,  19,  20,  41,  54,  64, 

05.  Go.  07.  75,  93,  111. 
hylodrumus,  50,  59,  77-78,  81,  82. 

I. 

Idahoensls,  41,  54-65,  57,  58,  64,  65,  60,  72, 

73,  78. 
imperator,  13,  20-21,  21,  22,  34,  41,  00, 
iraplger,  81-82. 
innuitus.   110-112. 
inoplnatus,  Vehilarrtos,  132-133. 
insularis,  9,  44-45. 
internationaiis,  103. 

K. 

kadiaki,  126. 

kenaiensis,    114,    115,    118,    124,    127-129, 

130,  131. 
kennerlyl.  61-62,  85. 
kidderi,  10,  106-108,  5  08.  109,  110. 
klamathensis,  29,  33,  72-74,  74,  75. 
kluane,  10,  48,  55,  78,  79-80,  81,  82,  92,  96, 

1)8,  102. 
kwakiutl,  9,  21,  23,  24-25,  26,  27,  28,  45. 

U 

latifrons.  78,  97-99. 

List  of  species  and  subspecies,  14-16. 

M. 

macfarlani,  49,  51-52. 

macrodon,  37   (footnote),  38,  38-40,  53. 

maglster,   10,  31,   75-76. 

Material  examined,   10-11. 

mendocinensis,  74-75. 

mcrriami,  124.  126. 

middendorffl.  10,  124,  125,  126,  126-127. 

mirabilis.  !),  10,  92. 

mirus,  40-41,  42. 

N. 

navaho.  37. 
neglectus,  9,  28-29. 
nelsoni,  34-35,  87. 
nortoni,  24,  25-27. 
nuchek,  113,  117,  122-124, 

O. 

ophrns,  49,  50,  103-104,  105. 
ors'.loides,  46.  46-47,  48. 
orgilos,  42,  44,  45,  45-46,  47,  48,  51. 
oribasus,  56-58. 

P. 

pallasi.  4G.  47.  47-49,  55,  79,  80,  102. 

pellyensls,  82-83,  83.  84.  99. 

prrturbans,  54,  64-65,  6~>. 

pervagor,  24,  25,  58.  50.  68-69,  70,  71,  73. 

phjeonvx.  98.  98,  101-102. 

planiceps,  36.  37.  37-38,  39.  40,  41. 

pulchcllus,  48,  55-56,  56,  58,  80,  83,  102. 

R. 

richardsoni.  83.  99-100,  100. 
rogersl,  54.  65-66,  67. 
riinsrlnsl,  49-50,  50,  51. 
russelli,  100-101. 


135 


136 


NOBTH  AMERICAN  FAUNA. 


(No.  41. 


taglttalls,  50-51. 

aaribur,  14  (footnote). 

selkirki,  77,  78. 

Sexual  differences,  10. 

sheldoni.  00.   115,  129-181. 

shirasi,  0,   1 20-1 22. 

Shoshone,  35,  30,  41,  43,  58,  60-61,  86,  93, 

104. 
sitkensis,  119,  119-120. 
stikeenensis,  9,  10,  28,  43,  44,  88-90,  92. 

T. 

tahltanicus,  9,  40,  41,  42,  43-44,  44,  45,  4G, 

48.  83.  84,  89. 
Technical  terms,  11-12. 
texensis.  35,  35-36,  37,  38,  39,  40. 
toklat,  80.  83,  95,  96-97,  97,  98,  102. 
townsendi,  115-116. 

Tremarctos,  genus,   132.  / 

tularensis,  30-32,  88. 
tundrensis,  95,  108. 

V. 

Ursus,  genus,  13,   14-16,  17-131,  181,  132. 
absarokus,    18,  21.   41,   60,  01,  67,  85, 

89,  93-94,  105,  106. 
alascensis,  94-95,  96,  100,  108,  110. 
alexandra;,  75,  111,  114-115,  128,  129. 
andcrsonl,  83-84. 
apache,  54,  84-85,  86. 
arizonae,  53-54,  54,  64,  66,  67,  85. 
atnarko,  22-23. 
bairdi,  13,  IS,  19-20,  21,  37  (footnote). 

63,  64.  G5,  06,  07,  76. 
blsonophagUH,  54,  66,  67-68. 
californicus,  29-30,  30,  31,  32,  73,  74, 

75.  76. 
canadensis,  43,  52-63,  68,  105. 
candescens,  17. 
eaurlnus,  9,  68,  69-71. 
chelan,  57,  58,  68-60,  78. 
chelidonia*.  13,  21-22. 
cinereus,  17. 
colusus,  30,  32-33,  74. 
crassodon,  90,  01. 
crassus,  90-92,  119. 
cressonus,  111,  112,  112-113,  117,  118, 

123,  124. 
dalli,    9,    98,    101.    102,    113,    116-118, 

120,  122    123,  129. 
dnsorsus,  33-34. 
eltonclarkl.  41-43,  44,  45. 
erfunetes,  55,  56,  57,  58. 
eulophus,  9,  69,  70,  71-72,  121, 
eximius.   109-110. 
ferojr,  17. 
griseus,  17. 

gyas.  106,  107,  108,  124-126,  127, 
henshawi.  31,  73,  87-88. 
hoots,  44,  91,  118-119, 
horriaceu8,  86. 
horriaMis,    7.    35.    36,    61,    62,    84,    85, 

85-87,  87,  104. 
horribilis,  7,  13.  17-19,  19,  20,  41,  54, 

64.  65,  66,  67,  75,  93,  111, 
hylodrorous.  50.  59,  77-78,  81,  82. 
Idahoensis,  41.  54-65,  57,  58,  64,  65,  66, 

72,  73,  78. 
imperator,  13,  20-21,  21,  22,  34,  41,  66. 


Ursus  Implgrr,  81-82. 
Innuitus,  110-112. 

insularis  '.),  44-45. 

intemationalis,  103. 

kudiaki,  12U. 

kenaiensis,  114,  115,  118,  124,  127-129, 

130.  131. 
kennerlyi,  61-62,  85. 
kidderi,  10,  100-108,  10S,  109,  110. 
klamatbensis,  20,  33,  72-74,  74,  75. 
khiaae.    Id,   48,   55,   7S,    79-80,  81,  82, 

1)2.  96.  08,  102. 
kwakiutl,  9,  21,  23.  24-25,  20,  27,  28,  45. 
latifrons,  78,  97-99. 
macfarlanl,  40,  51-52. 
macrodou,  37   (footnote),  38,  39-40,  53. 
magister.  10,  31,  75-76. 
mendocinensls,  74-75. 
merriami,  124,   126. 
mlddendorffi,  10,  124,  125,  126,  126-12S. 
mirabilis,  !».  10,  92. 
mirus,  40-41,  42. 
navabo.  37. 
neglectus,  0,  28-29. 
nelsoni,  34-35,  87. 
nortoni,  24,  25-27. 
nuchek,  113,  117,  122-124. 
opnrus,  49,  50.  103-104,  105. 
orgiloldes,  46,  46-47,  4S. 
orgilos,  42    44,  45,  45-46,  47,  48,  51. 
oribaeus,  56-58. 

pallasi,  46,  47,  47-49,  55.  79,  80,  102. 
pellyensls,  82-83,  83,  84,  09. 
pertnrbans,  54,  64-65,  65. 
pervagor,    24,    25,    58,    59,    68-69,    70, 

71,  73. 
pha?onyx,  96,  98,  101-102. 
planleeps,  36,  37.  37-38,  SO,  40,  41. 
pulchellus,  48,  55-56,  56,  58,  80,  83,  102 
ricbardsoni,  83.  99-100,  100. 
rogersl,  54,  65-66,  67. 
rungiusi,  49-50,  50,  51. 
russelli.  100-101. 
•agittalis,  50-51. 
saribur,  14  (footnote). 
telkirki,  77,  78. 
sheldoni.  96.  115.  129-131. 
shirasi.  9,   120-122. 
Shoshone,  35,  36,  41,  43,  58,  60-61,  86, 

03.  104. 
sitkensis,  119.  119-120. 
stikeenensis,  9,  10,  28.  43,  44,  88-90,92. 
tahltanicus,  9,  40.  41,  42,  43-44,  44,  45, 

46.  48.  83,  84,  89. 
texensis.  35,  35-36,  37,  38,  39,  40. 
toklat,  80.  83,  95,  95-97,  97,  98,  102. 
townsendi,  115-116. 
tularensis,  30-82,  88. 
tundrensis,  95,  108. 
utahensis,  61,  02-64,  64,  65,  66,  70. 
warburtoni,  22.  27-28.  28. 
washake,  59,  60.  104-106. 
utahensis,  61,  62-64,  64,  65,  66,  76. 

V. 

Yetulareto*,  genus    16.  131-183. 
iBoplnattiB,  132-133. 

W. 

warburtoni,  i2.  27-28,  28. 
washake,  59,  60,  104-100. 


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4 


LETTER  OF  TRANSMITTAL. 


United  States  Department  or  Agriculture, 

Bureau  of  Biological  Survey, 
Washington,  D.  C,  November  23,  1916. 
Sir:  I  have  the  honor  to  transmit  for  publication  as  North 
American  Fauna  No.  42,  a  report  on  life  zone  investigations  in 
Wyoming,  by  Merritt  Gary,  Assistant  Biologist  of  the  Biological 
Survey.  The  report  is  based  on  the  results  of  natural  history  ex- 
plorations conducted  in  recent  years  by  Survey  field  parties  in  all 
the  important  physiographic  areas  of  the  State.  The  first  section 
characterizes  the  five  transcontinental  life  zones  represented  in 
Wyoming,  defines  their  extent  and  limits,  and  discusses  their  eco- 
nomic possibilities.  The  second  consists  of  notes  on  the  distribution 
and  abundance  of  conspicuous  trees  and  shrubs  observed  during  the 
progress  of  the  survey.  Of  particular  importance  in  connection 
with  this  report,  as  well  as  with  others  yet  to  be  made  on  the  dis- 
tribution of  the  birds  and  mammals  of  Wyoming,  is  the  accompany- 
ing map,  which  shows  in  detail  the  extent  and  boundaries  of  the 
life  zones  which  traverse  the  State. 
Respectfully, 

Henry  W.  Henshaw, 
Chief,  Biological  Survey. 
Hon.  David  F.  Houston, 

Secretary  of  Agriculture. 


CONTENTS. 


Introduction 7 

Physiography  and  climate 9 

Life  zones 12 

Upper  Sonoran  Zone 13 

Great  Plains  Division 14 

Great  Basin  Division 17 

Green  River  Valley 18 

Bed  Desert 19 

Wind  River  Basin 21 

Bighorn  Basin 23 

Characteristic  species 24 

Mammals 25 

Breeding  birds 26 

Reptiles 27 

Amphibians 27 

Plants 28 

Agricultural  utility 30 

Transition  Zone 31 

Characteristic  species 32 

Mammals 34 

Breeding  birds 34 

Plants 36 

Agricultural  utility : 37 

Canadian  Zone 38 

Characteristic  species 42 

Mammals 42 

Breeding  birds 43 

Plants 44 

Hudsonian  Zone 46 

Characteristic  species 47 

Mammals 48 

Breeding  birds 49 

Plants 49 

Arctic- Alpine  Zone 50 

Characteristic  species 51 

Mammals 51 

Breeding  birds 51 

Plants 52 

Importance  of  Boreal  zones  to  Wyoming  and  adjoining  areas 52 

Notes  on  the  distribution  of  conspicuous  trees  and  shrubs 55 

Index 83 

3 


ILLUSTRATIONS. 


PLATES. 

Page. 

Plate  I.  Map  of  Wyoming,  showing  life  zones Frontispiece. 

II.  Fig.  1. — Chugwater  Valley,  showing  growth  of  juniper,  mountain 
mahogany,  and  box  elder.  Fig.  2.— Western  edge  of  Great  Plains 
area,  showing  scattered  yellow  pine  and  juniper 16 

III.  Fig.  1. — Plains  yucca  in  flower,  Chugwater  Valley.     Fig.  2. — Nearer 

view  of  same 16 

IV.  Fig.  1. — Bluffs  along  Green  River,  showing  growth  of  greasewood. 

Fig.  2.— Desert  vegetation,  Red  Desert,  chiefly  saltbush 24 

V.  Fig.  1. — Yellow  pine  forest,  north  base  of  Laramie  Peak.     Fig.  2. — 

Sagebrush  plain,  Wind  River  Basin 24 

VI.  Fig.  1. — Sagebrush  in  Wind  River  Valley,  lower  Transition  Zone. 

Fig.  2. — Garfield  Peak,  Rattlesnake  Mountains,  showing  sagebrush .        32 
VII.  Fig.   1.— Narrow-leaved   cottonwoods  and   yellow  pines,   Rawhide 
Butte,  edge  of    Great  Plains,  lower  border  of  Transition  Zone. 
Fig.  2. — Plain  at  southwestern  base  of  Wind  River  Range,  showing 

rabbit  brush 32 

VIII.  Fig.  1.— Snake  River  Valley,  Jackson  Hole,  showing  mixed  forest  con- 
ditions at  lower  edge  of  Canadian  Zone.     Fig.  2. — Heavy  Engel- 

mann  spruce  forest,  Wyoming  Range 40 

IX.  Fig.  1. — Sylvan  Lake,  Yellowstone  National  Park,  in  Engelmann 
spruce  belt.     Fig.  2. — Heavy  stand  of  lodgepole  pine,  north  slope 

of  Ferris  Mountains 40 

X.  Fig.  1.— Engelmann  spruces, timberline  on  Whirlwind  Peak,  Absaroka 

Range.     Fig.  2. — White-barked  pines,  same  locality 48 

XI.  Fig.  1.— Lower  edge  of  Hudsonian  Zone,  Wind  River  Range  south  of 
Fremont  Peak.  Fig.  2.—  Timberline,  east  slope  of  Needle  Moun- 
tain, showing  growth  of  spruce,  fir,  and  willow 48 

XII.  Fig.  1. — Arctic-Alpine  Zone,  Wind  River  Range  south  of  Fremont 

Peak.     Fig.  2. — Arctic-Alpine  Zone,  Absaroka  Range 48 

XIII.  Fig.  1.-— Teton  Range,  Mount  Moran  south  to  Grand  Teton.     Fig.  2. — 

Snow  in  lower  Hudsonian  Zone,  east  slope  of  Bridger  Peak,  Sierra 
Madre,  July,  1911 52 

XIV.  Fig.  1.— Heavy  forest  of  Engelmann  spruce,  Wyoming  Range.     Fig. 

2. — Rank  vegetation,  Canadian  Zone  forest  floor,  Absaroka  Range. .        52 
XV.  Fig.  1.— Mountain   mahogany,  west  slope  of  Bighorn   Mountains. 

Fig.  2. — Nearer  view  of  same 68 

TEXT  FIGURES. 

Fig.  1 .  Map  of  Wyoming  showing  routes  and  collecting  localities  of  Merritt  Cary 
and  other  members  of  the  Biological  Survey,  mainly  from  1909  to 
1915 9 

2.  Irrigation  canal  traversing  original  sage  plain,  Bighorn  Basin 25 

3.  Forest  of  white-barked  pine  just  below  timberline,  Absaroka  Range. .        55 

5 


6  ILLUSTRATIONS. 

Page. 

Fig.   4.  Forest  of  Engelmann  spruce,  Ferris  Mountains 56 

5.  Fringe  of  blue  spruce,  base  of  Wind  River  Range 57 

6.  Forest  of  Douglas  spruce,  Ferris  Mountains 58 

7.  Grove  of  aspen  poplar,  base  of  Laramie  Peak 60 

8.  Large  narrow-leaved  cottonwood,  upper  Wind  River  Valley 61 

9.  Copses  of  gray-leaved  willow  at  timberline,  Absaroka  Range 62 

10.  Thicket  of  canoe  birch,  Bear  Lodge  Mountains 64 

11.  Clumps  of  mountain  birch,  base  of  Shirley  Mountains 65 

12.  Fringe  of  alder  on  Pacific  Creek 66 

13.  Flowering  raspberry,  base  of  Casper  Mountains 70 

14.  Mountain  balm  in  flower,  Sierra  Madre 74 

15.  Canadian  buffaloberry  in  aspen  woods,  Wyoming  Range 75 

16.  Red  elderberry  in  bloom,  Sierra/Madre 77 

17.  Black  sagebrush,  upper  Wind  River  Valley 80 


No.  42.  NOKTH  AMERICAN  FAUNA.  Oct.,  1917. 

LIFE  ZONE  INVESTIGATIONS  IN 
WYOMING. 


By  Meeritt  Caky. 


INTRODUCTION. 

Wyoming  is  among  the  foremost  of  our  States  in  its  wealth  of 
natural  scenery,  culminating  in  the  grandeur  of  Yellowstone  Na- 
tional Park,  one  of  the  wonders  of  the  world.  In  addition  to  this 
distinction  it  posseses  vast  open  plains  and  lofty  mountains  whence 
flow  the  headwaters  of  mighty  river  systems  emptying  far  away 
to  the  west  into  the  Pacific  Ocean,  to  the  southeast  into  the  Gulf 
of  Mexico,  and  to  the  southwest  into  the  Gulf  of  California.  The 
various  slope  exposures  of  its  mountain  ranges,  the  fertility  of  its 
intervening  valleys  or  basins,  and  the  aridity  of  its  desert  spaces 
present  a  study  of  geographic  and  vertical  distribution  of  wild 
life  that  is  in  many  particulars  unique. 

The  study  of  geographic  and  vertical  distribution  of  life  with 
the  governing  factors  and  attendant  problems  is  valuable  as  a  matter 
of  scientific  research  and  in  the  attainment  of  practical  knowledge. 
The  Biological  Survey  has  been  making  detailed  investigations  of 
the  transcontinental  life  belts,  or  zones,  of  North  America  for  some 
years,  and  this  work  has  been  carried  on  with  special  reference  to 
their  practical  value.  It  has  become  increasingly  evident  that  life 
zones  furnish  a  fairly  accurate  index  to  average  climatic  conditions 
and,  therefore,  are  useful  as  marking  the  limits  of  agricultural 
possibilities,  so  far  as  these  are  dependent  upon  climate.  The 
knowledge  thus  gained  has  been  published  and  made  available  as 
the  investigations  have  progressed  and  the  life  zones  have  been 
mapped.1 

The  opening  up  to  agriculture  of  the  arid  and  semiarid  West 
through  irrigation  and  efficient  methods  of  conserving  the  natural 

*For  detailed  discussion  and  classification  of  the  life  zones  of  North  America  see 
Merriam,  C.  Hart,  Life  Zones  and  Crop  Zones  of  the  United  States  (Bull.  10,  Biological 
Survey,  U.  S.  Dept.  Agr.,  1898)  ;  also  Bailey,  Vernon,  Biological  Survey  of  Texas  (North 
Amer.  Fauna  No.  25,  1905)  ;  Fourth  Provisional  Zone  Map  of  North  America,  prepared 
by  the  Biological  Survey,  1910 ;  Cary,  Merritt,  Biological  Survey  of  Colorado  (North 
Amer.  Fauna  No.  33,  1911)  ;  and  Bailey,  Vernon,  Life  Zones  and  CroD  Zones  of  New 
Mexico  (North  Amer.  Fauna  No.  35,  1913). 

7 


8  NORTH    AMERICAN    FAUNA.  [No.  42. 

rainfall  offers  a  favorable  field  for  the  practical  application  of  this 
knowledge.  A  wide  range  of  altitude  and  a  correspondingly  varied 
climate  and  physiography  include  from  two  to  six  of  the  major  life 
zones  in  each  of  the  several  States,  and  the  zonal  boundaries  are 
on  the  wdiole  well  marked  by  reason  of  the  usually  rapid  or  abrupt 
changes  in  elevation.  New  areas  are  continually  being  reclaimed. 
while  in  practically  all  the  Western  States  large  districts  await 
future  development.  This  is  especially  true  of  the  Rocky  Mountain 
States,  where  general  agriculture  has  been  least  developed. 

In  Wyoming,  agriculture  has  made  rapid  strides  during  the  past 
few  years,  but  it  has  not  yet  advanced  much  beyond  the  experimental 
stage,  and  the  possibilities  are  somewhat  limited  by  a  cool  climate 
due  to  high  average  base  level.  It  appears  unlikely  that  crop  pro- 
duction will  in  future  greatly  exceed  the  local  demand.  There  are, 
however,  certain  restricted  areas  of  low  elevation  and  moderate 
climate  where  a  variety  of  crops  and  some  of  the  hardier  fruits  have 
proved  decidedly  successful.  A  special  value  attaches  to  these  low- 
lying  districts  inasmuch  as  they  are  immediately  surrounded  or 
bordered  by  extensive  nonagricultural  areas  where  mining,  lumber- 
ing, and  stock  raising  are  the  principal  industries.  The  melting 
snows  of  Wyoming  mountains  furnish  an  unfailing  supply  of  water 
for  irrigation  purposes,  and  Federal  and  private  irrigation  projects 
have  already  reclaimed  considerable  sections.  Much  valuable  agri- 
cultural land  in  the  valleys  and  basins  awaits  future  development. 
Useless  experimentation  might  be  avoided  or  a  more  favorable  loca- 
tion secured  if  the  prospective  as  well  as  the  resident  agriculturist, 
and  especially  the  horticulturist,  would  become  familiar  with  the 
groups  of  native  species  of  mammals,  birds,  reptiles,  and  plants 
which  have  proved  to  be  closely  associated  with  the  areas  of  success- 
ful production  of  particular  crops  in  other  parts  of  the  arid  West. 

Natural  history  explorations  carried  on  in  recent  years  by  the 
Biological  Survey  in  all  the  important  physiographic  areas  of  Wyo- 
ming warrant  the  present  report  on  the  life  zones  with  the  accom- 
pan}Ting  map  (PI.  I).  Sufficient  material  has  been  gathered  also 
for  inclusion  of  notes  on  the  distribution  of  conspicuous  trees  and 
shrubs  and  for  later  reports  on  the  mammals  and  birds  of  the  State.1 

*The  present  report  combines  the  results  of  field  investigations  for  the  Biological  Sur- 
vey conducted  at  various  times  by  Dr.  C.  Hart  Mcrriam,  Vernon  Bailey,  B.  II.  Dutcher, 
J.  Alden  Loring,  Edward  A.  Preble,  Alexander  Wetmore,  II.  E.  Anthony,  Stanley  G. 
Jewett,  and  D.  D.  Streeter,  jr.  ;  besides  those  made  by  the  author  since  1909  (see  map 
of  Wyoming,  showing  routes  of  field  parties,  fig.  1).  Lists  and  other  publications  bearing 
cm  the  distribution  of  the  Wyoming  fauna  and  flora,  although  few  in  number,  have  been 
freely  consulted.  For  identification  of  many  of  the  plants  collected  in  the  survey  the 
author  is  indebted  to  Dr.  J.  N.  Rose  and  Paul  C.  Standley,  of  the  F.  S.  National 
Herbarium,  and  to  F.  V.  Coville,  curator  of  the  National  Herbarium,  who  has  named  the 
Ribes.  The  few  reptiles  and  amphibians  collected  have  been  identified  by  Dr  Leonhard 
Stejneger,  of  the  U.  S.  National  Museum. 


1917.] 


PHYSIOGRAPHY   AND   CLIMATE. 


PHYSIOGRAPHY  AND  CLIMATE. 

In  common  with  other  States  of  the  Rocky  Mountain  region, 
Wyoming  has  a  varied  physiography  and  climate  and  great  natural 
resources.  The  surface  features  may  be  classified  broadly  as  moun- 
tains, plains,  and  valleys  or  basins. 

The  continental  watershed  formed  by  the  main  chain  of  the 
Rockies  enters  the  State  through  Yellowstone  Park  near  the  mid- 
western  boundary  of  the  forested  plateau  and  maintains  a  general 
southeasterly  trend  along  the  lofty  crests  of  the  Absaroka  and  Wind 
River  Ranges,  lowering  in  the  Red  Desert  region  to  arid  plains  and 


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Fig.   1. — Map  of  Wyoming  showing  routes  and  collecting  localities  of  Merritt  Cary  and 
other  members  of  the  Biological  Survey,  mainly  from  1909  to  1915. 

alkaline  basins  at  7,000  feet  elevation,  but  again  rising  to  the  sum- 
mits of  the  Sierra  Madre,  where  it  crosses  the  southern  boundary 
of  the  State  near  its  middle  point.  This  watershed  divides  the 
Great  Plains  from  the  Great  Basin,  and  on  its  slopes  in  northwest 
Wyoming  rise  the  headwaters  of  the  Colorado  in  Green  River,  the 
Columbia  in  Snake  River,  and  the  Missouri  in  Yellowstone  and 
Madison  Rivers. 

The  mountains  of  Wyoming,  massed  largely  in  the  northwest, 
occupy  approximately  a  fourth  of  the  total  area.  Chief  among  them 
are  the  Absaroka,  Wind  River,  Gros  Ventre,  and  Teton  Ranges  in 


10  NORTH   AMERICAN   FAUNA.  [No.  42. 

the  northwest;  the  Bighorn  Mountains  in  the  central  northern  por- 
tion; and  the  Sierra  Madre  and  the  Medicine  Bow  Ranges  at  the 
south.  Most  of  these  are  heavily  forested  groups  of  great  elevation, 
whose  summits  and  crests  reach  far  above  timberline  and  are 
usually  snow  capped  even  in  midsummer.  All  belong  to  the  Rocky 
Mountain  system  except  the  Bighorn  Mountains,  which  alone  are 
detached  from  the  main  chain.  Gannett  Peak  in  the  Wind  River 
Mountains,  at  13,785  feet,  is  the  highest  point  in  the  State,  exceeding 
the  height  of  its  close  neighbor,  Fremont  Peak  (13,730  feet)  ;  and  of 
the  Grand  Teton  (13,717  feet)  in  the  Teton  Range. 

The  Ferris,  Green,  Seminole,  Shirley,  and  Rattlesnake  Ranges  are 
small  separated  groups  along  the  upper  Platte  and  Sweetwater 
Valleys  and  in  the  northern  borders  of  the  Red  Desert,  lying  a  lit- 
tle east  of  the  continental  watershed  but  indicating  the  general 
course  of  the  Rockies.  These  differ  greatly  in  configuration,  but 
are  usually  characterized  by  densely  forested  northern  slopes  and 
summits  and  abrupt  barren  southern  exposures  facing  the  desert. 
In  this  region  also  are  the  huge  bare  granite  heaps  and  domes  rising 
from  the  sandy  plain  north  of  the  Sweetwater,  known  collectively 
as  the  Granite  Mountains.  Together,  these  small  ranges  are  a  pleas- 
ant relief  to  a  generally  barren  landscape. 

The  southwest  corner  of  Wyoming  is  much  broken  by  the  northern 
timbered  shoulders  of  the  Uinta  Mountains  and  by  barren  elevated 
ridges  and  mesas  on  either  side  of  Green  River  and  east  of  Bear 
River.  In  the  southeast  the  Laramie  Mountains  are  a  foothill  spur 
of  the  Medicine  Bows  continuous  north  to  the  Laramie  River  Gap 
and  thence  in  more  broken  formation  extending  to  the  Platte  Val- 
ley near  Casper.  Conspicuous  groups  of  outlying  hills  at  the  east 
and  northeast  are  the  pine-clad  Hartville  Mountains  north  of  Guern- 
sey and  the  densely  forested  Black  Hills  and  Bear  Lodge  Mountains. 

Although  well  supplied  with  mountains,  Wyoming  is  perhaps  bet- 
ter known  for  its  vast  open  plains.  These  are  either  level  or  rolling, 
lying  mainly  between  4,500  and  7,000  feet  elevation,  and  are  dis- 
tinguished by  characteristic  types  of  vegetation,  as  the  sage  plains  of 
the  high,  arid,  interior  plateaus;  and  the  grassy  plains  to  the  east 
and  northeast,  which  are  part  of  the  Great  Plains.  These  treeless 
expanses  were  ranged  long  before  historic  times  by  great  bands  of 
buffalo,  and,  succeeding  these,  by  countless  herds  of  cattle  and  sheep, 
and  their  great  grazing  value  is  well  attested  by  the  long  and  bitter 
warfare  for  their  possession  between  cattle  barons  and  flockmasters 
which  marked  the  days  of  the  open  range.  At  present  dry  farming 
is  greatly  restricting  the  cattle  range  on  the  Cheyenne  and  Lusk 
Plains  and  elsewhere  along  the  eastern  edge  of  the  State.  The  ex- 
tensive arid  sage  plains  farther  west,  however,  are  mainly  utilized 
for  sheep  grazing,  to  which  they  are  peculiarly  adapted. 


1917.]  PHYSIOGRAPHY  AND  CLIMATE.  11 

The  numerous  valleys  of  Wyoming  are  "well  watered,  and  with 
their  rich  soils  and  low  elevations  (chiefly  below  5,000  feet)  include 
the  areas  of  greatest  agricultural  importance  and  promise.  Most 
important  are  those  of  the  North  Platte,  Laramie,  Cheyenne,  Belle 
Fourche,  Powder,  Bighorn,  Wind,  Sweetwater,  and  Green  Rivers.1 
The  Bighorn  and  Wind  River  Valleys  are  extensive  basins  of  low 
altitude  and  mild  climate,  well  suited  to  the  production  of  certain 
fruits  and  other  crops.  The  more  elevated  valleys  of  the  Sweetwater 
and  Green  Rivers  are  mainly  devoted  to  stock  raising. 

The  Red  Desert  is  an  extensive  barren  alkaline  plain  or  basin  of 
great  aridity  lying  mainly  west  of  the  continental  watershed  in  the 
southern  part  of  the  State.  Without  perennial  streams  and  with 
soils  strongly  alkaline,  it  would  appear  to  have  no  agricultural 
future.  Alkali-resistant  desert  shrubbery  and  the  moderate  winter 
climate  of  this  region  nevertheless  combine  to  furnish  an  excellent 
winter  range  for  sheep,  and  it  has  long  been  thus  utilized  by  flock- 
masters. 

Wyoming's  lowest  elevation  is  in  the  extreme  northeast,  and  its 
highest  is  in  the  northwest.  Plains  and  plateaus  occupy  much  of  its 
southern  half.  The  altitudinal  extremes  are  3,100  feet  (approxi- 
mate), where  the  Belle  Fourche  River  crosses  the  eastern  boundary; 
and  13,785  feet,  on  the  summit  of  Gannett  Peak  in  the  Wind  River 
Range. 

The  climate  of  the  State  is  mainly  arid,  the  rainfall  ranging  from 
12  to  15  inches  in  the  semiarid  eastern  Great  Plains  area  to  under 
10  inches  in  the  extreme  arid  central  desert  region  (Bighorn  Basin2 
and  Red  Desert).  A  heavier  precipitation  in  the  Bear  Lodge  and 
Black  Hills  districts  at  the  northeast  (15  to  20  inches)  admits  of 
tolerably  successful  agriculture  without  irrigation.  The  high  tim- 
bered mountain  ranges  receive  a  great  deal  of  moisture,  not  only  as 
winter  snows,  but  also  during  summer  as  frequent  heavy,  dashing 
rains.3 

The  elevated  base  level  of  Wyoming  (about  6,000  feet)  insures  a 
generally  cool  climate.  Warm  summers  (mean  summer  temperature 
about  65°  F.)  with  a  long  growing  season  and  moderate  winters 
with  light  snowfall  are  the  rule  only  at  the  lower  levels  in  the  north 
and  east.  The  high  interior  valleys,  plains,  and  plateaus  are  marked 
by  short,  cool  summers  (mean  summer  temperature  about  55°  F.), 
with  prevalent  late  spring  frosts,  and  by  long  winters  with  tolerably 
heavy  snowfall  and  frequent  cold  winds.  The  snowfall  is  excessive 
in  the  mountainous  country  of  the  northwest,  where  occasionally  very 

1  Most  of  the  valleys  are  treated  in  some  detail  under  their  respective  zones. 

2  The  lowest  parts  of  the  Bighorn  Basin  often  receive  less  than  G  inches  of  annual 
rainfall. 

3  The  precipitation  usually  given  for  the  mountains  is  over  18  inches,  but  data  are 
lacking  for  the  higher  altitudes,  where  it  must  be  much  greater. 


12  NORTH   AMERICAN    FAUNA.  [No.  42. 

low  temperatures  are  recorded,1  but  the  winter  season  as  a  whole  is 
perhaps  less  severe  than  on  the  high  wind-swept  plains. 

For  a  State  with  an  arid  climate  Wyoming  is  exceptionally  well 
watered,  and  among  its  natural  resources  none  is  more  essential  to 
its  future  development  than  its  rivers  and  streams.  The  Snake, 
Yellowstone,  Bighorn,  and  Green  Rivers  rise  in  the  mountains  of 
the  northwest;  the  Tongue,  Powder,  Belle  Fourche,  and  Cheyenne 
Rivers,  with  their  numerous  tributaries,  head  in  the  Bighorn  Moun- 
tains and  the  elevations  of  the  northeast ;  while  the  North  Platte  and 
Laramie  Rivers,  which  describe  long,  circuitous  courses  in  the  south- 
eastern part  of  the  State,  have  their  sources  in  the  high  ranges  of 
Colorado. 

LIFE  ZONES  OF  WYOMING. 

Wyoming  has  a  generous  representation  of  animal  and  plant  life. 
This  is  largely  due  to  the  varied  climate  resulting  from  a  difference 
in  altitude  within  its  borders  of  nearly  10,700  feet;  and  in  a  lesser 
degree  to  a  difference  in  latitude  of  4  degrees,  and  a  wide  range  of 
local  physiographic  conditions. 

The  life  zones  range  from  Upper  Sonoran  (the  western  arid 
subdivision  of  the  Upper  Austral  Zone)  at  the  lowest  and  warmest 
elevations,  through  the  Transition,  Canadian,  and  Hudsonian,  to  the 
Arctic-Alpine  Zone  on  the  crests  of  the  highest  mountain  ranges. 
Of  the  seven  North  American  transcontinental  life  zones,  only  the 
Lower  Sonoran  and  the  Tropical  are  unrepresented ;  and  the  Upper 
Sonoran  Zone,  while  covering  large  areas,  is  represented  only  by  its 
upper,  cooler  part. 

The  five  zones  present  in  Wyoming  are  briefly  characterized  as 
follows:  Upper  Sonoran,,  the  zone  of  broad-leaved  cottonwood, 
juniper,  saltbush,  and  yucca,  occupying  most  of  the  valleys  and  low 
plains;  Transition,  the  zone  of  yellow  pine,  narrow-leaved  cotton- 
wood,  and  pure  sagebrush,  embracing  the  high  plains,  the  basal 
slopes  of  the  mountains,  and  all  except  the  highest  foothills;  Cana- 
dian, the  Boreal  forest  belt  of  spruce,  fir,  lodgepole  pine,  and  aspen, 
covering  the  middle  mountain  slopes  and  highest  foothill  ranges; 
Hudsonian,  the  narrow  zone  or  belt  of  white-barked  pine  and 
dwarfed  spruce  and  fir,  in  the  timberline  region;  and  Arctic -Alpine, 
the  treeless  zone,  on  mountain  crests  and  peaks  above  timberline. 

Zonal  boundaries  and  sequence  usually  are  well  marked  on  ranges 
rising  abruptly  from  a  low  base,  as  on  the  western  slope  of  the 
Bighorn  Mountains,  where  a  vertical  interval  of  about  9,500  feet 
may  be  traversed  in  15  or  20  miles;  and  on  the  Wind  River  Range 
southwest  of  Lander.     Under  gradual  change  in  altitude,  however, 

*  — 45°  F.  is  sometimes  recorded  in  Jackson  Hole,  and  though  data  are  lacking,  still 
lower  temperatures  undoubtedly  are  reached  on  the  high  ranges. 


1917.]  UPPEE   SONORAN   ZONE.  13 

as  on  the  open  plains,  the  passage  from  the  Upper  Sonoran  to  the 
Transition  Zone  is  often  scarcely  noticeable,  and  in  mapping  zones 
generally  the  limits  in  many  places  must  be  more  or  less  arbi- 
trarily fixed.  In  a  survey  of  native  mammals,  birds,  reptiles,  and 
plants  made  at  a  given  locality,  a  marked  preponderance  of  charac- 
teristic species  of  a  zone  decides  the  zonal  position ;  while  a  nearly 
equal  representation,  or  a  marked  absence,  of  species  peculiar  to  two 
adjoining  zones  is  indicative  of  an  intermediate  position,  or  the  ap- 
proximate boundary. 

The  several  zones  occupy  areas  of  great  irregularity  in  "Wyoming 
owing  to  the  very  broken  configuration,  and  their  vertical  boundaries 
are  subject  also  to  the  usual  variation  resulting  chiefly  from  differ- 
ences in  latitude,  base  level,  and  slope  exposure.  In  general,  zone 
levels  are  lowest  in  the  north  because  of  lower  base  and  higher 
latitude,  and  highest  in  the  south  where  the  base  level  is  more  ele- 
vated.1 

UPPER   SONORAN   ZONE. 

The  arid  subdivision  of  the  Upper  Austral  Zone,  the  Upper 
Sonoran,  occupies  nearly  a  third  of  Wyoming  (about  30,000  square 
miles),  and  covers  all  the  lower  levels.  Fully  half  this  area  lies 
below  5,000  feet  altitude  in  the  eastern  and  northern  sections,  and 
most  of  it  is  well  characterized.  On  the  high  plains  and  deserts  of 
the  south  the  Sonoran  element  is  present  between  5,500  and  6,500  feet 
elevation,  mainly  in  dilute  form.  Low  altitude,  a  warm  climate  and 
long  growing  season,  and  extensive  open,  level,  or  gently  rolling 
areas  of  rich  soils  combine  to  make  the  Upper  Sonoran  the  chief 
zone  of  crop  production,  dependent  as  in  other  sections  of  the  arid 
West  upon  careful  conservation  and  distribution  of  the  natural  water 
supply.  All  areas  adapted  to  any  extent  to  horticulture  lie  within 
this  zone  and,  because  of  its  agricultural  importance,  the  limits  and 
characterization  are  somewhat  detailed.1 

The  Upper  Sonoran  areas  of  Wyoming  are  mainly  broad  exten- 
sions of  the  zone  from  lower  elevations  on  the  south,  east,  and  north. 
Those  entering  from  the  south  comprise  a  narrow  strip  of  desert  valley 
along  both  sides  of  Green  River  north  nearly  to  Labarge;  and  in 
the  Red  Desert  a  broad  region  of  barren  plains  and  alkaline  de- 
pressions which  reaches  eastward  to  include  a  small  area  in  the 
upper  Platte  Valley  both  north  and  south  of  Fort  Steele.  The  above 
areas  are  narrowly  connected  along  Bitter  Creek,  and  more  broadly 
over  the  Snake  River  Valley  in  northwest  Colorado,  and  together 
they  form  the  northernmost  extensions  of  the  important  Green  River 
Upper  Sonoran  area. 

1  For  detailed  boundaries  of  the  zonal  areas  of  Wyoming  see  the  zone  map  (frontis- 
piece). 


14  NORTH  AMERICAN   FAUNA.  [No.  42. 

Two  large  tracts  of  Upper  Sonoran  country  in  the  east  are  part 
of  the  Great  Plains  area.  The  southernmost  of  these  is  approxi- 
mately bounded  on  the  west  by  the  5,000  to  6,000  foot  basal  plains 
flanking  the  Laramie  and  Hartville  Mountains,  and  extends  north 
to  the  narrow  strip  of  Transition  Zone  along  the  northern  escarp- 
ment of  the  Lusk  Plains.  In  the  North  Platte  Valley  the  zone  con- 
tinues narrowly  through  the  canyons  above  Guernsey,  then  in  greater 
width  to  Casper,  and  in  dilute  character  to  the  Seminole  Mountains 
and  to  Splitroek  in  the  Sweetwater  Valley.  North  of  the  Lusk  Plains 
the  Upper  Sonoran  Zone  includes  most  of  the  Cheyenne  River 
drainage  lying  between  the  pure  sage  plains  of  the  northern  central 
section  and  the  yellow  pine  country  of  the  Black  Hills.  The  Belle 
Fourche  Valley  in  the  northeast  also  carries  a  narrow  Upper  Sonoran 
strip  around  the  northern  and  western  bases  of  the  Bear  Lodge  Moun- 
tains, which  widens  above  Moorcroft  and  extends  nearly  to  the  Pump- 
kin Buttes. 

A  broad  band  of  this  zone  entering  the  State  from  the  north  along 
the  Bighorn  River  and  Clarks  Fork  covers  a  large  extent  of  low- 
altitude  country  in  the  arid  Bighorn  and  Wind  River  Basins  below 
5,500  or  6,000  feet  elevation.  These  two  areas,  of  which  the  Bighorn 
Basin  is  the  larger  and  agriculturally  the  more  important,  are  nar- 
rowly connected  through  the  canyon  south  of  Thermopolis,  but  are 
otherwise  separated  by  the  elevated  Transition  Zone  ridge  of  the  Owl 
Creek  Mountains.  East  of  the  Bighorn  Mountains,  the  valleys  of 
the  Tongue,  Powder,  Little  Powder,  and  Little  Missouri  Rivers  carry 
narrow  tongues  of  Upper  Sonoran  Zone  some  distance  into  the  State, 
separated  more  or  less  widely  by  low,  open  or  pine-clad  Transition 
Zone  watersheds. 

In  Wyoming,  as  in  other  States  traversed  by  the  continental 
watershed,  the  Upper  Sonoran  Zone  is  best  treated  under  its  two 
main  subdivisions,  the  Great  Plains  and  the  Great  Basin. 

Great  Plains  Division — Upper  Sonoran  Zone. 

Great  uniformity  of  surface  features  and  characterization  marks 
the  Great  Plains  area  from  the  Dakotas  to  Texas  and  west  to  the 
eastern  foothills  of  the  Rocky  Mountains.  It  is  a  vast  level  or  undu- 
lating region  of  abundant  grasses  and  moderate  rainfall,  entirely  open 
except  along  streams,  which  are  usually  fringed  with  deciduous  trees 
and  shrubbery,  and  also  in  the  rougher  parts  near  the  foothills  where 

1  The  subdivisions  are  based  upon  differences  in  climate,  configuration,  and  native 
species,  and  are  not  to  be  confused  with  the  physical  Great  Plains  and  Great  Basin 
areas  as  generally  understood  and  with  which  they  are  not  coextensive.  The  imperfect 
characterization  of  the  Great  Basin  division  in  Wyoming,  particularly  as  regards  its 
mammal  and  bird  life,  is  due  to  high  altitude,  and  in  the  Red  Desert  region  in  part  to  the 
iu fusion  of  Great  Plains  species  as  a  result  of  the  continuity  of  the  Sonoran  areas  on 
either  slope  of  the  Continental  Divide. 


1917.]  UPPER  SONORAN   ZONE.  15 

junipers  and  pines  often  occur  in  scattered  growth.  Areas  of  firm 
soils  alternate  with  tracts  of  sandhills  or  rolling  sandy  country,  and 
as  the  foothills  are  approached  there  are  scattered  areas  of  rough 
bad  lands,  the  bluffs  along  streams  become  rocky,  while  numerous 
talus  ridges,  and  clay,  chalk,  or  rock  buttes  of  usually  irregular  but 
in  some  places  of  strikingly  symmetrical  outline,  stand  up  from  the 
plain  (PL  II).  The  important  streams  of  the  Great  Plains  have 
their  sources  in  the  Rocky  Mountains  and  course  through  val- 
leys usually  shallow  and  often  sandy,  which  show  little  erosion.  This 
type  of  country  is  well  marked  along  the  eastern  edge  of  Wyoming 
as  far  north  as  Lusk. 

The  greatest  elevation  of  the  Upper  Sonoran  Zone  is  in  the  south, 
where  on  the  firm-soil  plains  of  the  Cheyenne  region  it  becomes  dilute 
at  5,500  feet  elevation,  and  its  upper  limits  are  indicated  at  approxi- 
mately 6,000  feet,  chiefly  by  absence  of  characteristic  species.  Exact 
delimitation  is  difficult  on  these  open  grassy  plains,  which  extend 
on  a  gradual  incline  to  the  7,000-foot  base  of  the  Laramie  Moun- 
tains. The  Chej^enne  Plains  descend  steadily  northward  to  the 
Platte  Valley  at  a  little  over  4,000  feet,  the  surface  meanwhile  be- 
coming broken  and  the  soil  sandy.  The  Sonoran  element  is  very 
pronounced  in  the  warm  valleys  of  the  Platte  drainage,  as  in  the 
Chugwater  Valley  below  Bordeaux ;  in  the  Laramie  Valley  at  Wheat- 
land, Jetsam,  and  Uva ;  in  Rawhide  Valley  below  Patrick ;  in  Goshen 
Hole;  and  along  the  North  Platte  below  Guernsey;  and  agriculture 
under  irrigation  in  these  districts  is  correspondingly  successful  and 
varied. 

North  of  the  Platte  Valley,  and  jutting  squarely  against  the  eastern 
bases  of  the  Hartville  Mountains  and  the  Rawhide  Butte,  is  an  exten- 
sive grassy  plateau  with  an  elevation  of  from  4,800  to  5,000  feet, 
extending  east  into  Nebraska  and  breaking  sharply  at  the  north 
toward  the  Cheyenne  River.  Upper  Sonoran  species  predominate 
up  to  5,000  feet,  but  the  region  is  near  the  upper  edge  of  the  zone, 
since  rocky  buttes,  ridges,  gulches,  and  cool  northern  declivities 
carry  a  scrubby  growth  of  yellow  pine,  Rocky  Mountain  juniper, 
red  currant,  mountain  mahogany,  and  other  Transition  Zone  vege- 
tation. This  plateau,  often  known  as  the  Lusk  Plains,  is  character- 
ized mainly  by  its  luxuriant  growth  of  nutritious  grasses  a>nd  has 
long  been  noted  as  choice  cattle  range.  Extensive  areas  on  both  the 
Cheyenne  and  Lusk  Plains  are  now  utilized  in  dry  farming,  to  which 
the  soil  and  climate  are  well  adapted. 

Characteristic  associations  of  Upper  Sonoran  species  mark  the 
lower  portions  of  the  Cheyenne  and  Lusk  Plains  and  the  North 
Platte  Valley.  Large  groves  of  broad-leaved  cottonwoods  (chiefly 
Populus  occidentalis)  are  on  the  North  Platte  and  Laramie  Rivers 


16  NORTH   AMERICAN-  FAUNA.  [No.  42. 

and  especially  on  Sibylee  Creek,  while  in  addition  to  cottonwoods 
the  usual  fringe  along  streams  consists  of  willows,  box  elder,  ash,1 
flowering  currant,  and  wolf  berry.  Common  shrubs  or  shrubby  plants 
on  dry  flats,  in  gulches,  and  on  rocky  or  gravelly,  slopes  are  saltbush, 
rabbit  brush,  narrow-leaved  sagebrush,  yucca  (PI.  Ill),  bush  morn- 
ing-glory, sand  cherry,  and  skunk  bush. 

Some  of  the  most  conspicuous  and  characteristic  flowering  plants 
are  Eriogonwm  annuum,  Eumex  venosus,  Abronia  elliptica,  Arge- 
mone  intermedia,  Cleome  serrulata,  Polanisia  trachysperma,  Luplnus 
plattensis,  Astragalus  crassicarpus  and  A.  mollissimus,  Psoralea 
(spp.),  P  etalostemon  (spp.),  Linum  rigidum,  Croton  texensis,  Ment- 
zelia  decapetala,  Opuntia  polyacantha,  Marmllaria  vimpara  and  M. 
missouriensis,  Anogra  albicaulis,  Meriolix  serrulata,  Lithospermum 
gmelini,  Llppla  cuneifolla,  Verbena  hastata  and  V.  bracteosa,  Phy- 
salis  lanceolata,  Solanum  rostratum,  Pentstemon  angustifolius,  Plan- 
tago  purshl,  Liatrls  punctata,  Grindclia  squarrosa,  Ratlbida  colum- 
narts,  Ilelianthus  annuus  and  H.  petiolarls,  Hymenopappus  ftlifolius, 
Carduus  plattensis,  and  Lygodesmia  rostrata. 

Mammals  which  especially  mark  this  region  as  Upper  Sonoran 
are  the  Kennicott  ground  squirrel,  prairie-dog,  Great  Plains  grass- 
hopper mouse,  prairie  harvest  mouse,  Colorado  bushy-tailed  wood 
rat,  Hayden  field  mouse,  yellow  pocket  gopher,  sage  pocket  gopher, 
Wyoming  kangaroo  rat,  Kansas  pocket  mouse,  Bailey  cottontail, 
black-footed  ferret,  northern  plains  mole,  and  California  bat. 

Characteristic  breeding  birds  of  the  plains  are  the  mourning  dove, 
burrowing  owl,  Arkansas  kingbird,  Bullock  oriole,  bronzed  grackle, 
lazuli  bunting,  lark  bunting,  western  grasshopper  sparrow,  western 
lark  sparrow,  white-rumped  shrike,  yellow  warbler,  long-tailed  chat, 
western  mockingbird,  catbird,  and  brown  thrasher. 

Reptiles  are  poorly  represented  on  the  plains  of  eastern  Wyoming. 
The  few  conspicuous  snakes  and  lizards  include  the  plains  rattle- 
snake, prairie  bull  snake,  hog-nosed  snake,  blue  racer,  garter  snakes, 
desert  horned  lizard,  sand  swift,  scaly  lizard,  six-lined  lizard,  and 
many-lined  skink. 

The  Upper  Sonoran  area  north  of  the  Lusk  Plains  is  open,  but 
much  rougher  in  configuration,  and  the  shrubby  type  of  vegetation 
becomes  increasingly  prominent,  especially  in  the  valleys  below 
4,500  feet  elevation,  where  the  Sonoran  element  is  strongest.  Grass 
and  cactus  flats  alternate  with  tracts  of  sagebrush'  rabbit  «brush,  and 
greasewood  over  much  of  the  drainage  basin  of  Cheyenne  River,  and 
in  the  low  Belle  Fourche  and  Little  Missouri  Valleys  in  northeast 
Wyoming.  The  watersheds  between-  the  valleys  are  either  ranges 
of  rolling  grassy  hills  or  abrupt  barren  ridges  of  bad  lands  of  about 

1  Uncommon. 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  II. 


Fig.  1.— Chuqwater  Valley  below  Chugwater. 
Juniper  and  mountain  mahogany  ( Cercocarpus  parvifolius)  on  bluffs,  and  box  elders  in  valley. 


Fig.  2.— Western  Edge  of  Great  Plains  Area. 
Scattered  yellow  pines  and  junipers  in  butte  country  southwest  of  Guernsey  (4,800  feet). 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  III. 


Fig.  1  .—Plains  Yucca  (Yucca  glauca)  in  Flower  in  Chuqwater  Valley  near 
Bordeaux,  July  1,  1909. 


Fig.  2.— Nearer  View  of  Same. 


1917.]  UPPER  SONORAN   ZONE.  17 

5,000  feet  elevation.  Extensive  breaks  with  more  or  less  yellow  pine, 
juniper,  and  Transition  Zone  shrubbery  margin  the  Cheyenne  River 
basin,  especially  on  the  south,  while  the  watercourses  usually  are 
bordered  narrowly  with  gnarled  cottonwoods,  willows,  box  elders, 
and  other  deciduous  trees  and  shrubs,  as  along  plains  streams  gen- 
erally. 

Sagebrush  country  rapidly  takes  the  place  of  grassy  plains  west  of 
Lusk,  varied  in  the  upper  valley  of  the  Platte  with  greasewood, 
rabbit  brush,  saltbush,  and  other  desert  shrubbery,  which  becomes 
increasingly  common  westward.  Shrubby  Sonoran  vegetation  fills 
also  most  of  the  narrow  extensions  of  the  Yellowstone  Valley  Upper 
Sonoran  Zone  in  the  valleys  of  the  Little  Powder,  Powder,  and 
Tongue  Rivers  east  of  the  Bighorn  Mountains. 

Although  the  shrubby  type  of  desert  vegetation,  more  barren 
surface,  and  greater  aridity  of  the  upper  Platte  Valley,  and  the 
Upper  Sonoran  areas  between  the  Black  Hills  and  Bighorn  Moun- 
tains, would  seem  to  place  them  with  the  Great  Basin  division  of 
the  zone,  their  mammals,  birds,  and  herbaceous  plants  are  mainly 
those  of  the  Great  Plains.  Species  which  are  absent  or  rare  on  the 
plains  farther  south  include  among  plants  the  greasewood,  black 
sagebrush,  several  species  of  rabbit  brush,  white  sage,  saltbushes, 
and  stanleya;  and  among  mammals,  the  Maximilian  and  Sweet- 
water pocket  mice.1 

Upper  Sonoran  areas  of  eastern  Wyoming  are  within  the  semi- 
arid  region  and  receive  in  most  sections  a  mean  annual  rainfall  of 
from  12  to  15  inches.  This  is  sufficient  for  a  luxuriant  growth  of 
the  best  varieties  of  range  grasses  in  all  sections  with  suitable  soils, 
but  will  not  admit  of  agriculture  apart  from  irrigation  or  dry- 
farming  methods,  except  possibly  around  the  base  of  the  Black 
Hills,  where  the  precipitation  is  a  little  greater.  Under  irrigation 
the  rich  alluvial  soils  of  the  valleys  yield  abundant  crops,  while  the 
moderate  returns  from  the  soil  secured  most  years  in  the  dry- 
farming  communities  scattered  over  the  plains  are  inducing  a 
steadily  increasing  settlement  of  these  districts. 

Great   Basin   Division — Upper   Sonoran   Zone. 

The  Green  River  Valley,  Red  Desert,  and  the  Bighorn  and  Wind 
River  Basins  have  a  barren  surface  which  shows  everywhere  much 
erosion,  especially  along  the  larger  streams,  which  in  many  places 
flow  through  rugged  canyons.  These  open  interior  areas  of  from 
4;000  to  6,500  feet  elevation  lie  within  the  arid  region  of  slight 
rainfall,  and  the  scanty  vegetation  is  of  the  shrubby,  bunchlike, 

1A  number  of  mammals  and  birds  and  a  few  plants  found  commonly  on  the  Lusk  Plains 
and  southward  do  not  reach  this  region. 

74440°— 17 2 


18  NORTH   AMERICAN   FAUNA.  [No.  42. 

desert  type  so  characteristic  of  the  Great  Basin  region  as  a  whole 
(PI.  IV,  fig.  2),  in  marked  contrast  with  the  grassy  plains  of  eastern 
Wyoming.  A  narrow  fringe  of  scattering  junipers  usually  marks 
the  upper  border  of  the  zone  on  the  rough  lower  margins  of  these 
desert  valleys  and  basins,  while  considerable  areas -of  junipers  with 
scattering  pinyons  partly  fill  the  upper  part  of  the  zone  in  lower 
Green  River  Valley,  and  in  the  extreme  southwestern  borders  of  the 
Red  Desert.1  The  Great  Basin  division  is  better  characterized  in 
Wyoming  by  vegetation  and  climate  than  as  a  faunal  region,2  al- 
though a  few  Great  Basin  species  of  both  mammals  and  breeding 
birds  occur  in  the  Green  River  Valley  and  on  the  Red  Desert.  The 
mammals  and  birds  of  the  Wind  River  and  Bighorn  Basins  are 
mainly  those  common  to  the  Great  Plains. 

The  great  dearth  of  rainfall  in  these  desert  areas  precludes  ordi- 
nary agriculture  and  even  dry  farming.  Effective  agriculture  is 
therefore  possible  through  irrigation  alone. 

Green  River  Valley — Upper  Sonoran  Zone. 

The  Upper  Sonoran  area  extending  north  along  Green  River  nearly 
to  Labarge  is  considered  apart  from  the  Red  Desert,  with  which  it 
is  connected,  on  account  of  different  topographic  features  and  a 
stronger  characterization  in  its  lower  part,  near  the  Utah  boundary, 
where  the  elevation  is  only  5,800  feet  in  the  river  valley.  In  this 
section  it  is  a  rough  incised  region  of  rocky,  juniper-clad  ridges  and 
dry,  open  canyons  or  narrow  valleys  reaching  gradually  down  to 
Green  River  from  high,  bordering  Transition  Zone  hills  and  mesas; 
and  of  precipitous,  brightly  colored  canyons  whose  various  shades 
of  red  and  yellow  are  in  strong  contrast  with  the  intense  black  of  the 
juniper  ridges.  The  steep,  hot,  protected  slopes  of  this  broken  can- 
yon region  carry  the  Upper  Sonoran  Zone  regularly  as  high  as  7,000 
feet. 

Its  species  are  characteristic  of  the  juniper  and  pinyon  belt  of  the 
Great  Basin,  but  are  in  less  variety  than  in  this  part  of  the  zone 
farther  south,  and  some  of  them  reach  but  a  short  distance  into  the 
State.  Junipers,  scattering  pinyons,  mountain  mahogany,  shadscale, 
syringa,  and  cactuses  comprise  the  principal  vegetation  on  ridges  and 
dry  slopes ;  saltbushes,  grayia,  yellow  cleome,  eriogonum,  and  cactuses 
are  common  shrubs  and  plants  on  the  open  sand  or  adobe  flats  in  the 
valleys  between  the  ridges;  the  skunk  bush  and  flowering  currant 
form  characteristic  shrubbery  on  the  bluffs  immediately  along  Green 
River ;  while  extensive  flats  in  the  wider  parts  of  the  river  valley  are 

1  The  belt  of  junipers  and  pinyons,  which  in  regions  farther  south  is  usually  present  in 
good  width  and  characterizes  the  upper  part  of  this  division  of  the  zone,  is  but  poorly 
indicated  in  Wyoming,  where  the  zone  is  more  often  open  throughout. 

2  See  footnote,  p.  14, 


1917.]  UPPER   SONORAN   ZONE.  19 

densely  covered  with  grease  wood  (PL  IV,  fig.  1)  and  glasswort  in 
damp,  alkaline  spots.  ' 

The  gray  titmouse,  Baird  wren,  and  piny  on  jay  are  common  birds 
in  the  juniper  growth,  and  probably  breed,  as  does  also  the  sage  spar- 
row, on  the  greasewood  flats.  Among  the  characteristic  Upper  So- 
noran  mammals  of  this  region  are  the  least  and  rock  chipmunks, 
golden-breasted  canyon  mouse,  True's  cliff  mouse,  Green  River  pocket 
gopher,  and  Great  Basin  spotted  skunk.  The  reptiles  and  amphibians 
include  the  plains  rattlesnake,  scaly  rock  lizard,  Stansbury  sand  liz- 
ard, short-horned  lizard,  and  the  spadefoot  toad. 

North  of  the  canyon  region,  which  extends  to  within  15  or  20 
miles  of  the  town  of  Green  River,  the  Upper  Sonoran  Zone  spreads 
out  in  dilute  form  to  include  the  lower  valleys  of  the  Black  Fork  and 
Big  Sandy,  connecting  eastward  through  the  Bitter  Creek  drainage 
with  the  larger  area  on  the  Red  Desert.1  As  a  narrowing  strip  along 
Green  River  it  continues  to  the  warm  valley  flats  between  Fontenelle 
and  Labarge.  This  is  an  open,  deeply  eroded  region  of  barren  val- 
leys and  bench  lands,  of  bare  mesas  and  variously  colored  bad  lands, 
buttes,  and  bluffs,  whose  zonal  position  is  best  defined  by  its  conspicu- 
ous vegetation,  chiefly  that  common  to  the  Red  Desert.  Upper 
Sonoran  species  of  mammals  and  breeding  birds  are  comparatively 
few  in  number  in  this  high  part  of  the  zone,  where  saltbushes,  grayia, 
small  brown  sagebrush,  rabbit  brush,  and  other  desert  shrubbery, 
and  such  plants  as  yellow  cleome  and  stanleya  give  way  to  the  pure 
sage  and  rabbit  brush  plains  of  the  Transition  Zone  at  about  6,500 
feet  elevation. 

The  Green  River  Valley  is  rather  bleak  and  inhospitable  and  does 
not  seem  to  invite  agricultural  development.  In  character  it  resem- 
bles the  Red  Desert  in  many  ways,  but  is  less  intensified,  particularly 
in  the  aridity  and  the  alkalinity  of  its  soils.  Owing  to  erosion, 
there  is  little  valley  land,  and  this  little  is  partly  used  in  the  cultiva- 
tion of  forage  crops.  At  the  north  a  limited  agriculture  is  possible, 
wherever  lands  can  be  brought  under  irrigation,  on  the  small  areas  of 
bench  lands  which  lie  along  Green  River  and  its  few  permanent  side 
streams.  The  rainfall  is  insufficient  for  a  good  growth  of  forage 
grasses,  yet  large  numbers  of  sheep  subsist  upon  the  desert  shrub- 
bery and  scanty  grass.  Sheep  grazing  appears  to  be  the  most  prac- 
ticable industry,  and  the  rough  character  of  lower  Green  River  Val- 
ley, with  its  many  sheltered  canyons,  warm  protected  slopes,  and 
mild  climate,  peculiarly  fits  it  for  a  winter  range. 

Red  Desert — Upper  Sonoran  Zone. 

Fairly  constant  surface  features  obtain  over  the  open  elevated 
region  of  undulating  plains  and  alkaline  depressions  or  basins,  known 

1  See  frontispiece   (map). 


20  NORTH   AMERICAN    FAUNA.  [No.  42. 

as  Red  Desert,  which  is  particularly  characterized  by  great  aridity, 
saline  or  other  strongly  alkaline  soils,  and  dearth  of  permanent  sur- 
face water.  The  only  relief  to  the  general  monotony  and  barrenness 
is  in  scattering  buttes,  occasional  ranges  of  low  hills,  bluffs  along  dry 
washes,  or  summits  of  distant  bordering  hills  and  mountains.  The 
impression  of  barrenness  is  only  intensified  by  the  prevalent  dull 
greenish  gray  or  occasional  light-gray  hues  of  desert  shrubbery.1 
The  continental  watershed  extends  across  the  north-central  portion 
at  an  elevation  of  less  than  7,000  feet  where  lowest,  being  crossed  by 
the  Union  Pacific  Railroad  at  Creston  at  7,000  feet.  Eastward  the 
altitude  decreases  slightly  towapd  the  North  Platte  Valley  (6,500 
feet),  and  westward  slopes  through  its  only  conspicuous  drainage 
area,  Bitter  Creek  Valley,  to  6,000  feet  on  Green  River.  The  climatic 
features  are  hot  sunny  days,  cool  nights,  very  slight  rainfall  during  a 
short  summer,  and  a  long,  moderately  open  winter.  The  heaviest 
snows  are  often  in  spring  and  are  more  beneficial  to  vegetation  than 
summer  rains,  which  are  either  largely  lost  through  rapid  evapora- 
tion or,  in  the  case  of  the  occasional  heavier  showers,  quickly  run  off 
the  barren  slopes  and  fill  the  dry  gulches  with  muddy  torrents. 
Frosts  are  not  uncommon  during  the  height  of  the  growing  season. 

Dilute  Upper  Sonoran  Zone,  poorly  characterized  apart  from  vege- 
tation, covers  the  lower  portion  of  the  Red  Desert  to  6,500  or  6,800 
feet  altitude,  a  total  area  of  between  5,000  and  6,000  square  miles,  in- 
cluding the  North  Platte  Valley  from  the  Platte  Canyon  to  above 
Fort  Steele.  The  conspicuous  and  dominant  vegetation  is  Upper 
Sonoran  desert  shrubbery — the  various  species  of  saltbush,  the  white 
sage,  greasewood,  grayia,  kochia,  rabbit  brush,  the  black  sagebrush,2 
and  low  desert  sages;  with  scattering  desert  junipers  on  many  of  the 
bluffs.  A  willow  (Salix  fluviatilis)  is  not  uncommon  on  creeks  of  the 
Bitter  Creek  drainage.  Such  plants  as  prickly-pear  cactus,  yellow 
cleome,  stanleya,  many  alkali-resistant  members  of  the  goosefoot 
family,  and  scattering  grasses,  conspicuous  among  which  are  wheat 
grasses  and  giant  rye-grass,  are  abundant  and  characteristic. 

The  small  striped  ground  squirrel,  pocket  mice,  long-eared  bat, 
least  chipmunk,  Green  River  pocket  gopher,  kangaroo  rat,  Bailey 
cottontail,  and  Great  Basin  spotted  skunk  are  Upper  Sonoran  mam- 
mals inhabiting  the  Red  Desert. 

The  variety  of  bird  life  is  very  limited,  and  few  characteristically 
Sonoran  birds  breed  here.  The  sage  sparrow  and  western  lark  spar- 
row are  perhaps  most  nearly  restricted  to  the  zone.  Other  birds 
found  in  abundance  during  the  breeding  season,  the  Brewer  spar- 

1  The  name  "  Red  Desert,"  originally  applied  to  a  restricted  area  of  reddish  soil  along 
the  Union  Pacific  Railroad,  does  not  convey  a  correct  impression  of  this  region  as  a 
whole,  where  desert  vegetation  instead  of  soil  lends  the  characteristic  colors. 

2  Abundant  also   in  Transition   Zone. 


1917.  J  UPPER   SONORAN   ZONE.  21 

row,  thick-billed  redwing,  western  meadowlark,  western  nighthawk, 
sage  thrasher,  and  white-rumped  shrike,  breed  also  on  the  surround- 
ing sage  plains  of  the  Transition  Zone. 

This  arid,  barren  waste  is  naturally  unsuited  to  human  habitation. 
Not  only  do  the  extensive  alkaline  deposits  in  the  desert  basins 
tend  to  make  soil  conditions  over  large  areas  unsuitable  for  crops, 
but  even  the  underground  water  over  practically  the  entire  region 
is  strongly  alkaline  and  unfit  for  use.  These  conditions,  combined 
with  a  deficient  rainfall  and  absence  of  perennial  streams,  give  no 
promise  of  an  agricultural  future.  The  few  ranches  are  situated 
mainly  along  the  skirts  of  the  desert,  where  a  number  of  small  creeks 
coursing  down  from  higher  country  furnish  sufficient  water  for  small 
fields  and  garden  patches  before  being  lost  in  the  sand.  Others  are 
found  at  the  few  widely  separated  springs  which  rise  in  different 
parts  of  the  desert.  A  ready  and  remunerative  market  for  garden 
truck  is  furnished  by  the  large  coal-mining  town  of  Eock  Springs, 
and  trucking  is  carried  on  in  that  district  wherever  there  is  sufficient 
water  for  irrigating  a  small  garden  patch. 

The  chief  value  of  the  Red  Desert,  aside  from  the  extensive  coal 
fields  in  its  western  part,  is  as  a  winter  range  for  the  hundreds  of 
thousands  of  sheep  which  spend  the  summer  on  the  higher  plains 
and  in  the  hills  and  mountains  of  central  and  western  Wyoming. 
The  abundant  sagebrush,  greasewood,  and  saltbushes,  particularly 
the  Nuttall  saltbush  (see  PL  IV,  fig.  2)  and  other  alkali-resistant 
shrubs  and  plants,  afford  an  abundance  of  winter  forage,1  while 
plenty  of  water  for  stock  is  insured  by  the  snows  which  drift  before 
the  frequent  winds  and  permit  browsing  in  the  cleared  spaces.  Its 
peculiar  adaptation  to  the  winter  feeding  of  sheep,  long  appreciated 
by  the  flockmasters,  gives  the  Red  Desert  region  a  special  importance 
as  a  necessary  complement  to  one  of  Wyoming's  great  industries. 

Wind  River  Basin — Upper  Sonoran  Zone. 

The  Upper  Sonoran  area  drained  by  Wind  River  and  its  affluents 
lies  mainly  between  5,000  and  6,000  feet  elevation.  It  extends  from 
the  southern  escarpment  of  the  Owl  Creek  Mountains  to  the  north- 
eastern base  of  the  lofty  Wind  River  Range,  and  at  the  east  and 
southeast  to  the  broad  tract  of  high,  rolling,  sagebrush  plains  which 
separate  it  from  the  Upper  Sonoran  areas  along  the  Platte  and  on 
the  Red  Desert.  In  its  surface  features,  climate,  vegetation,  and 
animal  life  this  region  is  generally  similar  to  the  Bighorn  Basin, 
with  which  it  is  connected  narrowly  through  the  rugged  canyon 
which  cleaves  the  Owl  Creek  Mountains  and  carries  the  waters  of  the 

1  For  a  full  discussion  of  the  pasture  value  of  the  alkaline  desert  basins  see  Nelson, 
The  Red  Desert  of  Wyoming  and  its  Forage  Resources,  Bull.  18,  Div.  of  Agrostology, 
U.  S.  Dept.  Agr.,  1898. 


22  NORTH  AMERICAN   FAUNA.  [No.  42. 

Bighorn.  The  greater  elevation  of  the  Wind  Kiver  Basin  results, 
however,  in  a  weaker  characterization  of  the  zone,  while  extensive 
areas  of  rough  bad  lands,  which  fill  much  of  the  upper  (western) 
part  between  Wind  Eiver  and  the  Owl  Creek  Mountains  and  form 
in  many  places  the  watersheds  between  streams  elsewhere,  greatly  re- 
strict the  agricultural  lands  and  confine  them  largely  to  the  valleys. 
Owing  to  its  proximity  to  the  mountain  mass  of  the  Wind  River 
Range,  the  western  edge  of  this  area  receives  more  moisture  than  the 
central  and  eastern  portions,  where  the  rainfall  seldom  exceeds  10 
inches. 

The  Upper  Sonoran  element  is  most  pronounced  in  the  lower  val- 
leys below  Riverton,  and  on  the  plains  and  bad  lands  which  jut 
against  the  southern  bases  of  the  Owl  Creek  Mountains.  The  vege- 
tation on  the  sandy  plain  north  of  Shoshoni  is  typical  of  the  lower 
elevations  generally,  consisting  of  saltbush,  greasewood,  rabbit  brush, 
small  brown  sagebrush,  spiny  sagebrush,  yucca,  and  prickly-pear 
cactus,  with  skunk  bush  and  juniper  on  bluffs,  and  broad-leaved 
cottonwood,  buffaloberry,  flowering  currant,  and  wolfberry  along 
streams.  Along  the  eastern  base  of  the  Wind  River  Range  and  on 
the  barren  slopes  and  gulches  of  the  higher  bad  lands  dilute  Upper 
Sonoran  Zone  reaches  to  nearly  6,000  feet  altitude,  and  here,  as  well 
as  on  the  southern  side  of  the  Owl  Creek  Mountains,  includes  the 
fringe  of  scattering  junipers  which  is  more  or  less  evident  on  the 
margins  of  the  open  basin.  The  effect  of  slope  exposure  on  zone  level 
is  especially  noticeable  on  the  warm  southern  slopes  of  the  Owl  Creek 
Mountains,  where  the  zone  is  carried  regularly  to  6,500  feet,  at  least 
500  feet  higher  than  on  the  cooler  basal  slopes  of  the  Wind  River 
Mountains,  on  the  opposite  side  of  the  basin. 

The  generally  weak  character  of  the  Upper  Sonoran  Zone  over  the 
Wind  River  Basin  is  evident  from  the  paucity  of  characteristic  zone 
species  of  mammals  and  breeding  birds,  which  include,  among  mam- 
mals, the  pale  chipmunk,  Great  Plains  grasshopper  mouse,  kangaroo 
rat,  Bailey  cottontail,  Great  Basin  spotted  skunk,  and  California 
bat ;  and  among  birds,  the  mourning  dove,  burrowing  owl,  Arkansas 
kingbird,  Bullock  oriole,  western  lark  sparrow,  white-rumped  shrike, 
and  long-tailed  chat.  The  plains  rattlesnake  and  the  desert  short- 
horned  and  scaly  rock  lizards  are  the  most  noticeable  reptiles. 

Extensive  coal  and  oil  fields  are  among  the  natural  resources  of 
the  Wind  River  Basin.  Agriculture  is  largely  supplemental  to  cattle 
and  sheep  raising.  Under  irrigation  the  arable  valley  lands  and  some 
of  the  lower  bench  lands  are  producing  fine  crops  of  alfalfa  and  other 
forage,  and  also  cereals.  The  growing  season  is  short  for  tomatoes 
and  tender  vegetables,  but  most  kinds  do  well  and  are  raised  exten- 
sively near  Lander,  Fort  Washakie,  Riverton,  and  elsewhere.    Apple 


1917.]  UPPER  SONORAN   ZONE.  23 

growing  has  proved  decidedly  successful  in  certain  protected  valleys 
at  the  base  of  the  mountains  southwest  of  Lander,  and  small  fruits 
are  grown  on  a  considerable  scale  at  Lander  and  elsewhere.  With  an 
abundant  supply  of  water  in  the  Wind  River  Range,  the  reclamation 
and  productiveness  of  considerable  areas  of  both  bench  and  valley 
lands  await  the  construction  of  additional  and  especially  higher  irri- 
gation canals. 

Bighorn  Basin — Upper   Sonoran   Zone. 

The  drainage  basin  of  the  Bighorn  River  is  a  large  open  area  of 
3,500  to  5,500  feet  elevation  lying  between  the  high  Bighorn  and 
Absaroka  Ranges  in  northwestern  Wyoming;  extending  from  the 
open  Transition  Zone  ridge  of  the  Owl  Creek  Mountains  northward 
beyond  the  Montana  line.  It  is  a  warm,  protected  section  of  low 
altitude  and  extreme  aridity,  but  with  an  abundance  of  permanent 
streams,  while  the  generally  barren  surface  of  its  valley  and  low 
bench  lands  or  scattered  tracts  of  bad  lands  supports  a  scanty  desert 
vegetation.  The  rainfall  is  often  as  light  as  5  or  6  inches  at  Basin 
and  Lovell,  in  the  low  central  and  northern  sections,  and  less  than 
10  inches  elsewhere.  The  Upper  Sonoran  Zone  in  this  region  covers 
an  area  of  fairly  regular  outline  about  100  miles  in  length  from 
north  to  south  by  60  miles  in  breadth,  approximately  6,000  square 
miles.  The  zone  is  strongly  characterized  in  the  valleys  and  over 
the  lower  portions  generally,  reaching  its  upper  limits  at  a  little 
over  5,000  feet  on  the  bordering  sage  slopes  west  and  south,  and  east- 
ward at  anywhere  from  5,500  to  6,000  feet  on  the  abrupt  hot  slopes 
along  the  western  bases  of  the  Bighorn  Mountains,  where  it  includes 
an  irregular  belt  of  scattered  junipers  varying  from  500  to  1,000  feet 
in  vertical  breadth. 

The  fauna  and  flora  of  the  Bighorn  Basin  are  derived  alike  from 
the  Great  Basin  and  Great  Plains  regions,  and  include  few  if  any 
species  not  common  to  some  of  the  other  Sonoran  areas  of  Wyoming. 
Over  this  region  the  Upper  Sonoran  Zone  is  variously  characterized 
as  to  vegetation  by  a  rank  growth  of  broad-leaved  cottonwood,  wil- 
low, buffaloberry,  skunk  bush,  and  flowering  currant  along  most  of 
the  streams;  greasewood,  rabbit  brush,  and  Suceda  on  adobe  river 
flats;  saltbushes,  rabbit  brush,  spiny  sagebrush,  prickly-pear  cactus, 
and  such  plants  as  Cleome  lutea,  Psoralea  tenuiflora,  and  Plantago 
purshi  on  firm-soil  benches,  with  Grayia  spinosa,  Polarmia  trachys- 
perma,  Lupinus  pusillus,  yucca,  sand  dock,  and  a  small  yellow- 
flowered  Malacothrix  added  in  sandy  areas;  and  by  a  scattering 
growth  of  juniper  and  skunk  bush  on  bad  lands  bluffs  and  on  the 
rough  southern  and  especially  eastern  margins  of  the  basin.  Ex- 
tensive barren  flats  midway  between  Greybull  and  Cody  have  an 


24  NORTH   AMERICAN   FAUNA.  [No.  42. 

almost  pure  growth  of  prickly-pear  cactus,  while  similar  tracts  near 
Frannie  are  likewise  clothed  with  the  Nuttall  saltbush. 

Representative  breeding  birds  include  the  mourning  dove,  bur- 
rowing owl,  Arkansas  kingbird,  Bullock  oriole,  bronzed  grackle, 
western  lark  sparrow,  sage  sparrow,  house  finch,  lazuli  bunting, 
white-rumped  shrike,  yellow  warbler,  western  yellow-throat,  long- 
tailed  chat,  catbird,  brown  thrasher,  and  western  marsh  wren. 

A  few  characteristic  mammals  of  the  Bighorn  Basin  are  the  pale 
chipmunk,  black-tailed  prairie-dog,  Great  Plains  grasshopper  mouse, 
Hayden  field  mouse,  sage  pocket  gopher,  kangaroo  rat,  Maximilian 
pocket  mouse,  Bailey  cottontail,  and  the  brown  and  California  bats. 
Several  reptiles,  the  prairie  bull  snake,  plains  rattlesnake,  horned 
lizard,  and  scaly  rock  lizard,  are  common  over  much  of  the  region. 

This  huge  depression  between  lofty  mountain  ranges  is  a  highly 
favored  region  of  great  promise.  While  physically  and  climatically 
the  best  suited  to  general  agriculture  of  any  of  the  low-altitude  areas 
of  western  Wyoming,  it  is,  perhaps,  best  known  for  its  adaptation 
to  horticulture  and  for  the  rapid  strides  already  made  in  the  success- 
ful production  of  high-grade  apples  and  other  fruits.  The  warm, 
sheltered  valleys  and  hot  Upper  Sonoran  slopes  along  the  western 
bases  of  the  Bighorn  Mountains,  especially  toward  the  northern  end 
of  the  basin,  are  highly  favorable  to  fruit  culture.  It  is  in  these 
situations  that  the  older  bearing  orchards  are  chiefly  located  and  the 
best  results  have  thus  far  been  obtained.  Young  orchards  are  now 
extensively  planted  throughout  the  lower  open  portions  of  the  basin 
and,  although  more  exposed  than  nearer  the  mountains,  nevertheless 
give  promise  of  handsome  returns  under  proper  care  and  due  atten- 
tion to  local  conditions. 

The  Bighorn  River  and  its  principal  tributaries,  the  Shoshone  and 
Greybull  Rivers,  and  Shell,  No  Wood,  and  Owl  Creeks,  fed  by  the 
melting  snows  of  high  mountain  ranges,  carry  an  abundance  of  water, 
amply  sufficient  under  proper  storage  control  for  watering  all  ir- 
rigable lands  in  their  drainages  (fig.  2).  Private  irrigation  projects 
have  already  reclaimed  considerable  portions  of  the  broad  and  fertile 
stream  valleys,  together  with  the  lowest  of  the  adjoining  bench 
lands,  and  the  Federal  Shoshone  project  has  opened  large  tracts  in 
Shoshone  Valley.  The  higher  bench  lands,  at  present  utilized  in 
sheep  grazing,  await  the  construction  of  more  storage  dams  and 
higher  irrigation  canals  before  they  can  be  made  productive. 

Characteristic  Species — Upper  Sonoran  Zone. 

The  delimitation  of  life  zones  is  based  upon  the  combined  ranges  of 
characteristic   species   of   mammals,   breeding  birds,   reptiles,    and 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  IV. 


Fig.  1.— Bluffs  alonq  Green  River  near  Utah  Boundary. 

Dense  growth  of  greasewood  (Sarcobatus  vermiculatus)  at  left. 


12c~ 


■  ^-lOi  '■•'»-"" 


■i 


Fiq.  2— Desert  Vegetation,  Red  Desert,  South  of  the  Ferris  Mountains. 

Chiefly  saltbush  (Atriplex  nuttalli). 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  V. 


Fig.  1.— Yellow  Pine  Forest  near  Springhill,  Northern  Base  of  Laramie  Peak 

(6,500  Feet). 


Fig.  2.— Sagebrush  Plain  near  Fort  Washakie,  Wind  River  Basin. 


1917.] 


UPPER  SONORAN   ZONE. 


25 


plants. x    Species  which  in  Wyoming  best  mark  the  Upper  Sonoran 
Zone  throughout.2  or  in  restricted  areas  of  its  Great  Plains  or  Great 


Fig.  2. — Irrigation  canal  traversing  original  sage  plain  on   bench  north  of   Burlington, 

in  the  Bighorn  Basin. 

Basin  subdivisions,  are  included  in  the  following  lists.  Many  species 
of  wide  zonal  dispersion  occur  commonly  in  the  Upper  Sonoran  Zone 
in  Wyoming,  but  as  these  give  no  definition  to  the  zone  they  are  not 
listed. 

Mammals — Upper  Sonoran  Zone. 

[Species  marked  T.  occur  also  in  the  Transition  Zone.] 


Antilocapra  americana  americanu,  An- 
telope.    T. 

Eutamias  pallidus  pallidas,  Pale  Chip- 
munk.    T. 

Eutamias  minimus  minimus,  Least 
Chipmunk. 

Eutamias  minimus  pictus,  Sagebrush 
Chipmunk. 

Eutamias  dorsalis  utahensis,  Utah 
Rock  Chipmunk. 

Callospermophilus  lateralis  ivortmani, 
Wortman  Mantled  Ground  Squir- 
rel.    T. 

Citellus  obsoletus,  Kennicott  Ground 
Squirrel. 

Citellus  tridecemlineatus  pallidus,  Pale 
Striped  Ground   Squirrel. 


Citellus  tridecemlineatus  parvus,  Small 

Striped   Ground   Squirrel. 
Cynomys     ludovicianus     ludovicianus, 

Black-tailed  Prairie-dog. 
Onychomys  leucogaster  arcticeps,  Great 

Plains  Grasshopper  Mouse. 
Peromyscus    manieulatus    o  s  g  o  o  di  , 

Black-eared  White-footed  Mouse.    T. 
Peromyscus  manieulatus  nebrascensis, 

Yellow  White-footed  Mouse. 
Peromyscus  crinitus  auripectus,Go\i\e\\- 

breasted  Canyon  Mouse. 
Peromyscus    truei    truei,    True's    Cliff 

Mouse. 
Reithrodontomys      megalotis      dyehei, 

Prairie  Harvest  Mouse. 


1  Species  of  insects  and  other  groups  might  be  equally  useful  in  the  determination  of 
zones  if  their  ranges  were  better  known. 

2  See  frontispiece   (map). 


26 


NORTH   AMERICAN   FAUNA. 


so.  41 


Mammals — Upper  Sonoran  Zone — Continued. 


Neotoma  cincrca  orolestes,  Colorado 
Bushy-tailed  Wood  Rat.     T. 

Microtus  ochrogaster  haydeni,  Hayden 
Field  Mouse. 

Fiber  eibethicus  tinnamominus,  Great 
Plains  Muskrat.     T. 

Geomys  lutescens,  Yellow  Pocket 
Gopher. 

Thomomys  otitis,  Green  River  Pocket 
Gopher. 

Thomomys  talpoides  bullatus,  Sage 
Pocket  Gopher. 

Perodipns  ordii  luteolus,  Wyoming 
Kangaroo  Rat. 

Perognathus  Mspidus  paradoxus,  Kan- 
sas Pocket  Mouse. 

Perognathus  fasciatus  fasciatus,  Maxi- 
milian Pocket  Mouse.     T. 

Perognathus  fasciatus  Vitus,  Sweetwa- 
ter Pocket  Mouse. 

Perognathus  flainis  piperi,  Cheyenne 
Pocket  Mouse. 


Perognathus  eallistus,  Red  Desert 
Pocket  Mouse. 

Lepus  californieus  melanotis,  Black- 
tailed  Jack  Rabbit. 

Sylvilagus  auduboni  baileyi,  Bailey 
Cottontail. 

Sylvilagus  floridanus  si  m  His,  Nebraska 
Cottontail. 

Canis  nebracensis,  Plains  Coyote.     T. 

Vulpes  velox  velox,  Kit  Fox,  Swift.    T. 

Procyon  lot  or  lot  or,  Raccoon.    T. 

Taxidea  taxus  taxus,  Badger.     T. 

Spilogale  gracilis  saxatilis.  Great  Ba- 
sin Spotted  Skunk. 

Mustela  nigripes.  Black-footed  Ferret. 

Scalopus  aquaticus  caryi,  Northern 
Plains  Mole. 

Myotis  californieus  caHfornicus,  Cali- 
fornia Bat. 

Myotic  evotis,  Long-eared  Bat. 

Myotis  longicrus  interior,  Long-legged 
Bat.     T. 

Eptesicus  fuscus  fusciis.  Brown  Bat.    T. 


Breeding  Birds — Upper  Sonoran  Zone. 
[Species  marked  T.  breed  also  in  the  Transition  Zone.] 


Querquedula  discors,  Blue-winged  Teal. 
T. 

Querquedula      ci/anoptera,     Cinnamon 
Teal.     T. 

Botaurus  lentiginosus,  Bittern.     T. 

Ardea  herodias.  Great  Blue  Heron.     T. 

Nycticorax   nycticorax  narius,   Black- 
crowned  Night  Heron.     T. 

Rail  us  Virginian  us,  Virginia  Rail.1 

Porzana  Carolina,  Carolina  Rail,  Sora. 
T. 

Fulica  americana,  Coot.    T. 

Bartramia  longicauda,  Upland  Plover. 
T. 

Numenius  americanus,  Long-billed  Cur- 
lew.    T. 

Zenaidura       macroura       carolinensis, 
Mourning  Dove. 

Archibuteo    ferrugi/neus,    Ferruginous 
Rough-legged   Hawk.     T. 

SiJeotiito  cunicularia  hypugcea,  Burrow- 
ing Owl. 

Chordeiles  rirginianus  hcnriji.  Western 
Nighthawk.     T. 

1  Observed  during 


Tyrannus  tiirannus.  Kingbird.     T. 

Tyrannus  rerticalis,  Arkansas  King- 
bird. 

Tyrannus  vociferans,  Cassin  Kingbird. 

Corrus  brachyrhynchos  hesperis,  West- 
ern Crow.     T. 

Cyanocephalus  cyanocephalus,  Pinyon 
Jay. 

Xanthocephalus  xanthoeephalus,  Yel- 
low-headed   Blackbird.      T. 

Agelaius  phoerdceus  fortis,  Thick-billed 
Redwing.     T. 

Sturnclla  neglecta,  Western  Meadow- 
lark.     T. 

Icterus  bullocki,  Bullock  Oriole. 

Quiscalus  quiscula  aneus,  Bronzed 
Grackle. 

Carpodacus  mexieanus  frontalis.  House 
Finch. 

Annnodrainus  sarannaruiih  bimacula- 
tus,  Western  Grasshopper  Sparrow. 

Chondestes  grammacus  strigatus,West- 
ern  Lark  Sparrow. 

Spisella  breweri,  Brewer  Sparrow.     T. 

breeding  season. 


1917.] 


UPPER  SONORAN   ZONE. 


27 


Breeding  Birds — Dppeb   Sonoran  Zone — Continued. 


AmpMspiza  neradensis,  Sage  Sparrow. 

Passerina  amcena,  Lazuli  Bunting. 

Spiza  americana,  Dickcissel. 

Calamospiza  melanocorys,  Lark  Bunt- 
ing.    T. 

Lanius  ludovicianus  excubitorides, 
White-rumped  Shrike.     T. 

Dendroica  (estiva  ce&tiva,  Yellow 
Warbler.     T. 

Oeothlypis  trichas  Occident alis,  West- 
ern Yellow-throat.     T. 

Icteria  rirens  longicauda,  Long-tailed 
Chat. 

Oreoscoptes  montanus.  Sage  Thrasher. 
T. 


Mimus  polyglottos  leucopterus,  West- 
ern Mockingbird. 

Dumetella  carolinensis,  Catbird. 

Toxostoma  rufum,  Brown  Thrasher. 

Thryomanes  betoicM  bairdi,  Baird 
Wren.1 

Telmatodytes  palustris  plexitis.  West- 
ern Marsh  Wren. 

Boeolophus  inomatus  griseus,  Gray 
Titmouse.1 

Psaltriparus  plumbeus,  Lead-colored 
Bush-tit.1 

Polioptila  ccBrulea  obscura,  Western 
Gnatcatcher.1 


Reptiles — Upper  Sonoran  Zone. 

[Species  marked  T.  occur  also  in  the  Transition  Zone.] 

Lizards. 


Holbrookia  maculata.  Sand  Swift. 
Uta  stansburiana,  Stansbury  Lizard. 
Sceloporus  consobrinus,  Scaly  Lizard. 
Sceloporus      graciosus,      Scaly      Rock 

Lizard. 
Phrynosoma      ornatissimum,      Desert 

Short-horned  Lizard.     T. 


Cnemidophorus    sexlineatus,    Six-lined 

Lizard. 
Eumeces     multivirgatus,     Many-lined 

Skink. 


Snakes. 


Thamnophis    sirtalis    parietalis,    Red- 
barred  Garter  Snake.     T. 
Thamnophis  radi.v.  Garter  Snake.     T. 
Bascanion  constrictor.  Blue  Racer. 
Pituophis  sayi,  Prairie  Bull  Snake. 


Liopeltis  vemalis,  Smooth  Green 
Snake.     T. 

Heterodon  nasicus,  Hog-nosed   Snake. 

Crotalits  confluentus,  Plains  Rattle- 
snake. 


Amphibians — Upper  Sonoran  Zone. 

[Species  marked  T.  occur  also  in  the  Transition  Zone.] 

Toads  and  frogs. 


Scaphiopus      hammondi      bombifrons, 

Spadefoot  Toad. 
Bufo    lentiginosus    woodhousei,    Toad. 

T. 


Bufo  cognatus.  Toad. 

Rana   pipiens,   Leopard   Frog.     T. 


Salamanders. 


Ambystoma  tigrinum.  Tiger  Salaman- 
der.    T. 


Probably  breeds. 


28 


NORTH   AMERICAN   FAUNA. 


[No.  42. 


Plants — Upper  Sonoran  Zone. 

[Species  marked  T.  occur  also  in  the  Transition  Zone.] 

Trees  and  shrubs. 


Pinus  edulis,  Pinyon,  Nut  Pine. 

Junipcrus  knighti,  Desert  Juniper. 

Junipcrus  monosperma,  One-seeded 
Juniper. 

Populus  occidentalis,  Broad-leaved 
Cottonwood. 

Populus  acuminata,  Lance-leaved  Cot- 
tonwood. 

Salia  amygdaloides,  Peach-leaved  Wil- 
low. 

Salix  fluviatilis,  Sand-bar  Willow. 

Ulmus  americana?  Elm.     T. 

Acer  negundo,  Box  Elder.     T. 

Fraxinus  lanceoiata,  Ash.     T. 

Cercocarpus  parvifolius,  Mountain  Ma- 
hogany.    T. 

Sarcobatus  vermiculatus,  Greasewood. 

Atriplex  canescens,  Saltbush,  Gray 
Shadscale. 

Atriplex  confertifoUa,  Spiny  Saltbush. 

Atriplex  nuttalli,  Nuttall  Saltbush. 

Atriplex  pabularis,  Nelson  Saltbush. 

Atriplex  argentea,  Silvery  Saltbush. 

Eurotia  lanata,  White  Sage,  Winter 
Fat. 

Grayia  spinosa,  Grayia. 

Kochia  americana,  Kochia,  White 
Sage. 

Philadelphus  occidentalis,  Western 
Syringa. 

Ribes  longiflorum,  Flowering  Currant. 

Prunus  americana,  Wild  Plum.     T. 

Prunus  besseyi,  Sand  Cherry. 


Amorpha     canescens,     False     Indigo, 

Shoestring.     T. 
Amorpha  nana,  False  Indigo. 
Schmaltzia  trilobata,  Skunk  Bush. 
Sell maltzia  glabra,  Smooth  Sumac.     T. 
Rhus  rydbergi,  Western  Poison  Ivy.    T. 
Vitis  vuipina,  Wild  Grape.     T. 
Paj-thenocissus       vltacea,        Virginia 

Creeper.     T. 
Lepargyrea  argentea,  Buffaloberry.     T. 
Symphoricarpos      occidentalis,      Wolf- 
berry.     T. 

Gutierrezia  sarothrw,  Rabbit  Brush. 
Chrysothamnus      graveolens,    ,  Rabbit 

Brush. 
Chrysothamnus      plattensis,       Rabbit 

Brush. 
Chrysothamnus        linifolius,        Rabbit 

Brush. 
Chrysothamnus    stcnophyllus,    Rabbit 

Brush. 
Tctradymia  inermis,  Rabbit  Brush. 
Tctradymia  spinosa,  Rabbit  Brush. 
Tetradymia  nuttalli,  Rabbit  Brush. 
Artemisia      fllifolia,      Narrow  -  leaved 

Sagebrush. 
Artemisia  spinescens,  Spiny  Sagebrush, 

Budbrush. 
Artemisia     pedatiflda,     Small     Brown 

Sagebrush. 
Artemisia,     tridentata,     Black     Sage- 
brush.    T. 


Herbaceous  plants. 


Tradescantia  occidentalis,  Spiderwort. 
Yucca  glauca,  Plains  Yucca. 
Eriogonum  effnsum,  Eriogonum. 
Eriogonum  annuum,  Eriogonum. 
Eriogonum  campanulatum,  Eriogonum. 
Eriogonum  corymbosum,  Eriogonum. 
Eriogonum  multiccps,  Eriogonum. 
Rumex  venosus,  Sand  Dock. 
Suwda  diffusa,  Sea  Blite. 
Suceda  moquini,  Shrubby  Blite. 
Endolepsis  sucTcleyana. 
Salicornia  rubra,  Glasswort.     T. 


Abronia  fragrans. 

Abronia  elliptica. 

Argemonc  intermedia,  Prickly  Poppy. 

Argemone  hispida..  Prickly  Poppy. 

Stanleya  tomentosa,  Stanleya. 

Stanleya  intcgrifolia,  Stanleya. 

Cleome  lutea,  Yellow  Cleome. 

Cleome  serrulata,  Red  Cleome,  Honey 

Plant. 
Polanisia  trachysperma.  Clammy-weed. 
Ijiipinus  plattensis.  Lupine. 
Lupiwus  pusillus,  Small  Lupine. 


>  Northeast  Wyoming  only. 


1917.] 


UPPER   SONORAN    ZONE. 


29 


Plants — Upper  Sonokan  Zone — Continued. 
Herbaceous  plants — Continued. 


Astragalus  crassicarpus,  Buffalo  Bean, 
Ground  Plum. 

Astragalus  mollissimus,  Milk  Vetch. 

Astragalus  missouricnsis,  Milk  Vetch. 

GlycyrrMza  lepidota,  Wild  Licorice.    T. 

Psoralea  tenuiflora,  Psoralea. 

Psoralea  Unearifolia,  Narrow-leaved 
Psoralea. 

Psoralea  lanceolata,  Psoralea. 

Psoralea  hypogca,  Psoralea. 

Psoralea  esculcnta,  Psoralea. 

Parosela  enncandra,  Dalea. 

Pctalostemon  oligophyllus,  White  Prai- 
rie Clover. 

Petalostcmon  eandidus,  Prairie  Clover. 

Petalostcmon  purpureas,  Purple  Prai- 
rie Clover. 

Petalostcmon  villosus,  Silky  Prairie 
Clover. 

Lathyrus  ornatus,  Wild  Pea,  Vetch- 
ling. 

Linum  rigidum,  Wild  Flax. 

Euphorbia  margiiiata,  Snow-on-the- 
inountain. 

C rot  on  texensis,  Croton. 

Malvastrum  coccincum,  False  Mallow. 

Malvastrum  dissection,  False  Mallow. 

Mentzclia  decapetala,  Loasa. 

Mentzclia  nuda,  Loasa. 

Mentzelia  albicanlis,  Loasa. 

Mentzclia  he ricaulis.   Yellow  Loasa.    T. 

Ma  miliaria  missouricnsis,  Ball  Cactus. 
T. 

Mamillaria  vivipara,  Ball  Cactus. 

Eehinocereus  viridiflorus,  Green- 
flowered  Petaya.     T. 

Opuntia  polyacantha.  Prickly  Pear. 

Opuntia  rutila,  Prickly  Pear. 


Anogra  albicaulis,  White  Evening 
Primrose. 

Galpinsia  la  v  and  ulw  folia.. 

Meriolix  serrulata. 

Oaura  coccinca. 

Asclepias  speciosa,  Milkweed. 

Asclepias  pumila,  Milkweed. 

I  pom  oca  leptophylla,  Bush  Morning- 
glory. 

Gilia  polycladon. 

Lithospcrmum  angustifolium,  Grom- 
well.     T. 

Lithospcrmum  gmcliiii,  Gromwell. 

Oreocarya  flava. 

Lippia  cuneifolia. 

Verbena   hastata,   Blue   Vervain. 

Verbena  bractcosa,  Low  Vervain. 

Physalis  lanceolata,  Ground  Cherry. 

Solarium  rostratum,  Buffalo  Bur. 

Pcntstcmon  albidus,  Beard-tongue. 

Pentstemon  angustifolius,  Beard- 
tongue. 

Plant  a  go  purshi,  Plantain. 

Liatris  punctata,  Blazing  Star. 

Grindelia  squarrosa,  Gum  Plant. 

Sideranthns  spinulosus. 

Solid  a  go  mollis,  Goldenrod. 

Solid  a  go  canadensis  gilvocanescens, 
Goldenrod. 

Solidago  rigida,  Goldenrod. 

Ratibida  columnaris,  Cone  Flower. 

Helianthus  annuiis.  Sunflower. 

Helianthus  petiolaris,  Sunflower. 

Hymenopappus  fllifolius. 

Cardans  plattensis,  Thistle. 

Malacothrix  sonehoides. 

Lygodesmia  juncea. 

Lygodesmia  rostrata. 


Grasses. 


Andropogon  scoparius,  Bluestem. 
Andropogon  halli,  Bluestem. 
Panieum  virgatum,  Panic  Grass. 
Aristida  longiscta,  Wire  Grass.     T. 
Stipa  comata.  Feather  Grass. 
Stipa  viridula,  Feather  Grass.     T. 
Oryzopsis  micrantha,  Rice  Grass. 
Eriocoma  cuspidata,  Indian  Millet. 


Muhlenbergia  pungens,  Dropseed  Grass. 
Sporobolus     airoides,     Fine-top     Salt 

Grass. 
Sporobolus  cryptandrus,  Bunch  Grass. 
Calamovilfa    longifolia,    Reed    Grass, 

Sand  Grass. 
Sehedonnardus      paniculatus,      Crab 

Grass. 


30 


NORTH    AMERICAN   FAUNA. 


[No.  42. 


Plants — Upper  Sonoran  Zone — Continued. 
Grasses — Continued. 


Spartina  gracilis.  Marsh  Grass. 
Bouteloua   oligostachya,   Grama 

Grass.     T. 
Bouteloua  hirsuta,  Grama  Grass. 
Atheropogon   curtipendula,  J?all 

Grama.     T. 
Bulbilis  dactyloides,  Buffalo  Grass. 
Munroa      squarrosa,      False      Buffalo 

Grass. 
Eatonia  obtusata. 
Distichlis  spicata,  Salt  Grass. 
Poa  fendlcriana,  Spear  Grass.     T 


/ 


Poa  sheldoni,  Spear  Grass.    T. 
Festuca  octoflora,  Fescue  Grass. 
Agropyron  spicatum,  Wheat  Grass. 
Agropyron  smithi,  Wheat  Grass.     T. 
Sitanion     longifolium,     Long  -  bearded 

Rye  Grass.     T. 
sitanion  Itystri.r.     T. 
Ely  in  us  canadensis,  Wild  Rye.    T. 
Elymus  condcnsatus,  Giant  Rye 

Grass.     T. 
Elymus  salinus,  Desert  Rye  Grass. 


Agricultural  Utility  of  the  Upper  Sonoran  Zone. 

The  growing  of  lm7  and  forage  crops  as  supplemental  to  stock 
raising  and  a  limited  planting  of  cereals  and  vegetables  for  ranch 
use  long  constituted  the  chief  agricultural  endeavors  in  even  the 
most  favored  sections  of  Wyoming.  The  establishment  of  agricul- 
ture as  a  separate  industry  in  this  arid  region  is  comparatively  recent, 
the  result  of  greatly  increased  irrigation  facilities  and  of  more  effi- 
cient methods  of  conserving  the  rainfall  in  the  less  arid  dry-farming 
districts. 

The  Upper  Sonoran  areas  of  Wyoming  lie  mainly  in  the  upper 
and  cooler  parts  of  the  zone  and  are  therefore  climatically  unsuited 
to  a  variety  of  tender  crops,  and  especially  fruits,  which  are  suc- 
cessfully grown  elsewhere  in  its  lower  and  warmer  portions.  They 
are  proving  well  adapted,  howTever,  to  most  of  the  standard  varieties 
of  wheat,  oats,  rye,  emmer,  and  other  cereals,  which  yield  abundantly 
under  irrigation  and  moderately  under  dry  farming.  None  but  the 
early  varieties  of  corn  mature,  and  these  are  raised  to  a  small  extent 
only  in  the  warmest  parts,  as  in  the  lower  Platte  and  Bighorn 
Valleys.  Alfalfa,  the  staple  forage  crop,  produces  regularly  two 
and  at  the  lowest  elevations  often  three  or  even  four  crops  a  year. 
Sugar  beets  are  raised  extensively  in  the  Bighorn  Basin,  the  lower 
Platte  Valley,  and  near  Wheatland.  The  sugar  content  of  Wyo- 
ming beets  is  high,  and  with  the  construction  of  local  sugar  factories 
this  crop  should  become  one  of  the  most  important  and  profitable. 
Potatoes  yield  abundantly  and  are  well  adapted  to  the  dry-farming 
districts  along  the  eastern  edge  of  the  State,  where  they  are  pro- 
duced on  a  large  scale.  Vegetables  of  most  sorts  thrive  under  irri- 
gation, and  trucking  is  profitably  carried  on  in  many  valleys.  The 
tender  sorts  usually  succeed  in  the  warmest  localities,  but  can  not  be 
raised  generally,  because  of  the  short  growing  season. 


7.]  TRANSITION    ZONE.  31 

Fruit  growing  has  been  attempted  on  a  commercial  scale  mainly 
the  Bighorn  Basin,  near  Lander,  and  in  the  Wheatland  district, 
Lere  many  of  the  hardier  varieties  of  apples,  cherries,  and  plums, 
well  as  the  small  fruits,  raspberries,  strawberries,  grapes,  currants, 
oseberries,  and  dewberries,  are  moderately  successful  under  irri- 
tion.  Even  in  these  sheltered  and  favored  districts  the  physical 
d  climatic  conditions  are  so  localized  that  successful  horticulture 
11  perforce  be  limited  to  comparatively  small  areas  where  there  is 
lple  protection  and  the  greatest  freedom  from  the  destructive  late 
osts,  which  kill  so  much  of  the  fruit  blossom  in  this  zone  through- 
t  the  Eocky  Mountain  region.  Late-flowering  and  frost-resistant 
rieties  succeed  best. 

TRANSITION  ZONE. 

Fully  half  of  Wyoming,  or  about  50,000  square  miles,  is  in  the 
■ansition  Zone,  the  zone  intermediate  between  the  Boreal  and  Aus- 
al  regions,  which  is  here  in  its  greater  part  open  and  treeless,  and 
us  less  conspicuously  characterized  than  in  the  southern  Rocky 
Mountain  region,  where  it  is  very  generally  marked  by  extensive  for- 
ests of  yellow  pine.    In  Wyoming  the  zone  comprises,  broadly,  vast 
interior  sagebrush  plains  (see  PL  V,  fig.  2)  and  watersheds,  plateaus, 
and  high-altitude  basins  in  the  central  and  southwest  sections;  and 
elevated  grassy  plains  to  the  east  and  southeast.    It  also  includes  all 
except  the  higher  summits  of  the  pine-clad  foothill  ranges  in  the 
eastern  part  of  the  State,  and  the  open  basal  sagebrush  slopes  of 
the  high   mountain  ranges   farther  west.     In   greater   detail,1   the 
Transition   Zone  in   Wyoming  embraces  the   following   important 
areas : 

The  Cheyenne  Plains  above  5,500  feet  elevation ;  Lusk  Plains  above 
5,000  feet ;  all  of  the  Laramie  Plains  and  Shirley  Basin ;  upper  valley 
of  the  North  Platte  above  6,500  feet;  upper  Green  River  Basin, 
Bear  River  region,  and  extensive  tracts  on  the  borders  of  the  Red 
Desert  above  6,500  feet;  sage  plains  between  Casper  and  Lander 
above  6,000  feet;  Wind  River  Valley  above  6,000  feet;  mainly  open 
borders  of  the  Bighorn  Basin,  5,500  to  7,000  or  7,500  feet;  all  open 
or  partly  timbered  watersheds  between  the  Bighorn  and  Bear  Lodge 
Mountains  above  4,500  feet;  basal  and  middle  slopes  of  Bear 
Lodge  Mountains  and  northern  Black  Hills,  4,000  to  5,500  feet; 
and  southern  Black  Hills  region,  4,500  to  6,000  feet.  Limited  areas 
in  Jackson  Hole  and  Salt  River  Valley  are  eastward  continuations 
of  the  Snake  River  Transition  Zone,  and  narrow  strips  of  the  zone 
enter  Yellowstone  Park  for  a  short  distance  along  the  Yellowstone, 

i  See  zone  map  (frontispiece). 


32  NORTH    AMERICAN    FAUNA.  [N- 

penetrating  the  valleys  of  the  Gardiner,  Lamar,  and  other  tributa 
and  also  along  the  Gallatin,  Madison,  and  Fall  River  valleys. 

The  foregoing  well  illustrates  the  effect  of  both  base  level 
latitude  on  the  vertical  position  of  the  Transition  Zone.     A 
level  of  over  6,000  feet  in  the  southwest  elevates  the  zone  i 
of  the  region  contiguous  to  the  Red  Desert  and  Green  River  ~\ 
to  between  7,000  and  8,500  feet  on  warmest  (southwest)  slope 
0,000  to  7,500  feet  on  coldest  (northeast)  slopes.    In  extreme 
eastern    Wyoming,    adjacent   to    the   low-altitude    northern 
Plains,  the  zone  level  attains  only  4,500  to  6,000  feet  on  sou 
slopes  and  4,000  to  5,500  feet  on  northeast  slopes.1     Much 
2,000-foot  variation  in  zone  level  results  from  difference  i 
elevation,  while  probably  800  feet  is  normal  depression  in  th( 
east  due  to  higher  latitude.     Locally,  mountain  slopes  of  unuo,. 
exposure  and  warmth  carry  zones  abnormally  high,  and  if  very 
steep  and  abrupt  the  horizontal  contraction  is  also  very  marked. 
This  is  well  shown  in  the  Transition  belt  on  the  abrupt,  hot,  south- 
west slopes  of  the  Bighorn  Mountains  east  of  Greybull  and  Ionia. 
Varied  physical  conditions,  and,  to  a  certain  extent,  deforestation, 
affect  both  the  elevation  and  the  horizontal  as  well  as  the  vertical 
width  of  life  zones,  especially  in  mountainous  districts.     In  the 
main,  however,  the  Transition  Zone  is  maintained  in  fairly  uniform 
elevation  and  width  along  the  bases  of  the  Wyoming  ranges. 

Characteristic  Species — Transition  Zone. 

The  Transition  Zone  in  Wyoming  is  conspicuously  marked  only 
along  its  upper  border,  where  mostly  open  sage  slopes  give  way  to 
the  aspen  and  coniferous  forest  belt  of  the  Canadian  Zone.  Incon- 
spicuous vegetation  is  characteristic  of  the  lower  part  on  the  open 
plains,  where  the  zonal  position  is  further  indicated  by  either  the 
absence  or  a  marked  paucity  of  Sonoran  species.  As  elsewhere  in 
the  Rocky  Mountain  region,  its  fauna  and  flora  are  fairly  constant 
throughout,  but  include  many  species,  both  Boreal  and  Austral,  from 
the  adjoining  zones. 

Sagebrush,  yellow  pine,  and  grasses  are  prominent  types  of  vege- 
tation in  the  Wyoming  Transition  area.  The  sagebrush  (PI.  VI), 
the  most  widely  distributed  shrub,  usually  occurs  in  pure  growth, 
while  the  yellow  pines  are  restricted  largely  to  the  lower  mountains, 
foothills,  and  rough  tracts  in  the  eastern  half  of  the  State.  Con- 
siderable Douglas  spruce  and  scattering  Rocky  Mountain  white  pines 

1  On  exposed  slopes  of  mountains  the  difference  in  zone  level  on  the  warm  and  cold 
sides  is  usually  as  much  as  1,000  feet.  So  much  variation  is  not  found  in  the  low  hill 
country  of  northeastern   Wyoming. 


North  American  Fauna  No.  42,  U.  S.  Dept  Agr.     Biological  Survey. 


Plate  VI. 


Fig.  1.— Sagebrush  in  Wind  River  Valley. 
Lower  part  of  Transition  Zone,  near  Circle. 


Fig.  2.— Garfield  Peak,  Rattlesnake  Mountains. 
Sagebrush  covers  much  of  this  barren  8,000  to  9,000  foot  range. 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  VII. 


'■-*ts£SBSi 


~!y*&$& 


Fig.  1.— Rawhide  Butte,  Lower  Border  of  Transition  Zone,  at  Edge  of  Great 

Plains. 

Narrow-leaved  cottonwoods  on  Rawhide  Creek,  southern  base  of  the  butte,  and  yellow  pines  on 

hills  (5,000  feet). 


Fig.  2.— Southwestern  Base  of  Wind  River  Range  near  Big  Sandy. 

Rabbit  brush  ( Chrysotltamnus  and  Tetradymia  inermis)  on  the  plain  (7,500  feet). 


1917.] 


TRANSITION   ZONE.  33 


occur  in  this  zone  in  the  mountains  of  western  Wyoming,  and  the  bur 
( •  ,k  is  a  common  tree  on  the  Bear  Lodge  Mountains  and  elsewhere  in 
northern  Crook  County.  On  streams  along  the  bases  of  the  moun- 
tains generally  the  zone  is  marked  by  narrow-leaved  cotton  wood  (PL 
VII,  fig.  1),  diamond  willow,  and  usually  by  a  dense  shrubbery  of 
Rocky  Mountain  birch,  black  and  red  haws,  cornel,  wild  gooseberry 
and  currant,  serviceberry,  and  silverberry;  on  foothill  and  lower 
mountain  slopes  both  in  the  forest  as  undershrubs  and  in  the  open, 
by  Rocky  Mountain  and  creeping  junipers,  Bebb  willow,  barberry, 
wild  red  currant,  mountain  mahogany,  kunzia,  ninebark,  wild 
cherry,  mountain  and  large-toothed  maples,  mountain  balm,  bear- 
berry,  mountain  snowberry,  and  several  high  plains  species  of  sage- 
brush and  rabbit  brush  (PL  VII,  fig.  2)  ;  and  throughout  by  a  great 
many  herbaceous  plants. 

A  considerable  number  of  birds  of  both  general  and  restricted 
breeding  range  within  the  zone  characterize  this  area  in  Wyoming 
during  the  nesting  season.  Representative  species  are  the  sage  hen, 
sharp-shinned  hawk,  saw-whet  owl,  Lewis  woodpecker,  white- 
throated  swift,  Wright  flycatcher,  magpie,  piny  on  jay,  McCown 
iongspur,  white- winged  junco,  mountain  song  sparrow,  arctic  and 
,  ireen-tailed  towhees,  western  tanager,  plumbeous  vireo,  Macgillivray 
warbler,  Rocky  Mountain  and  pygmy  nuthatches,  and  willow  thrush. 

Mammals  wholly  or  chiefly  restricted  to  the  Transition  Zone  in 
different  parts  of  Wyoming  include  the  plains  white-tailed  deer; 
Black  Hills  red  squirrel;  Wyoming  and  Uinta  ground  squirrels; 
white-tailed  prairie-dog;  bushy-tailed  wood  rat;  pygmy  and  Uinta 
field  mice;  Coues,  Black  Hills,  Fort  Bridger,  and  pygmy  pocket 
gophers;  Uinta  pocket  mouse;  prairie  jumping  mouse;  white-tailed 
lack  rabbit;  Black  Hills  cottontail;  northern  plains  skunk;  and  long- 
legged  bat. 

The  following  reptiles  and  amphibians  are  apparently  more  abun- 
dant in  the  Transition  than  in  the  Upper  Sonoran  Zone  in  Wyoming: 
Western  garter  snake  (Thamnophis  ordinoides  vagrans),  a  toad 
(Bufo  boreas),  and  a  frog  (Rana  pretiosus).  Others  of  regular  oc- 
currence in  at  least  the  lower  part  of  the  zone  include  the  horned 
lizard  (Phrynosoma  ornatissimum) ,  scaly  rock  lizard  (Sceloporus 
grariosus),  garter  snakes  (Thamnophis  sirtalis  parietalis  and  T.  ra- 
dli'),  smooth  green  snake  (Liopeltis  vernalis),  western  toad  {Bufo 
lentiginosus  woodhousei) ,  frogs  (Rana  pipiens  and  Chorophilus  tri- 
seriatus),  and  tiger  salamander  (Ambystoma  tigrinum). 
74440°— 17 3 


34 


NORTH   AMERICAN   FAUNA. 


[No.  42. 


Mammals — Transition  Zone. 

[Species  marked  U.  occur  also  in  the  Upper  Sonoran  Zone;  those  marked  C,  also  in  the 

Canadian.] 


Cervus  canadensis  canadensis,  Elk.     C. 
Odocoilcns      virginiamts      niacrourus. 

Plains  White-tailed  Deer. 
Odocoileus  hemionus   hemionus,   Mule 

Deer.     U.,  C. 
Ovis  canadensis  auduboni,  Bad  Lands 

Sheep.1     U. 
Sciurus  hudsonicus  dakotensis,  Black 

Hills  Red  Squirrel. 
Glaucomys  sabrinus  canescens,  Flying 

Squirrel. 
Callospermophilus    lateralis    lateralis, 

Say  Ground  Squirrel.     C. 
Callospermoph  Hits   lateralis    wort  mani, 

Wortman  Ground  Squirrel.     U. 
Citellus  richardsoni  clcgans,  Wyoming 

Ground  Squirrel.     U. 
Citellus  armatus,  Uinta  Ground  Squir- 
rel.    C. 
Cynomys     leucurus,     White  -tailed 

Prairie-dog.     U. 
Onychom ys     h  ucogaster     brevicaudus, 

Idaho  Grasshopper  Mouse.     U. 
Onychomys   leucogaster   tnissouriensis, 

Northern  Grasshopper  Mouse.     U. 
Peromyseus  leucopus  aridulus,  White- 
footed   Mouse. 
Ncotoma-  cinerea  cinerea,  Bushy-tailed 

Wood  Rat. 
Neotoma    cinerea,    orolestes,    Colorado 

Bushy-tailed  Wood  Rat.     17. 
Microtus     pennsylvanicus     modestus, 

Saguache  Meadow  Mouse.     U. 
Microtus       montanus       caryi,       Uinta 

Meadow  Mouse. 
Microtus    pan  per  rim  us,    Pygmy    Field 

Mouse. 
Fiber    zibethicus    osoyoosensis.    Rocky 

Mountain  Muskrat.     C. 


('((■star  canadensis,  Beaver.     C. 
Thomomys  talpoides  nebulosus,  Blark 

Hills  Pocket  Gopher.     C. 
Thomomys     talpoides     clusius,    Coues 

Pocket  Gopher. 
Thomomys      bridgeri,      Fort     Bridger 

Pocket  Gopher. 
Thomomys    pygmceus,    Pygmy    Pocket 

Gopher. 
Perognathvs     parvus     clarus,     Uinta 

Pocket  Mouse. 
Zapus    hudsonius    campestris,    Prairie 

Jumping  Mouse. 
Erethizon   epixanthum,    Yellow-haired 

Porcupine.     C. 
Lcpus    toiniscndi    eampanius,    White- 
tailed  Jack  Rabbit. 
Lcpus   townsendi    townsendi,   Western 

White-tailed  Jack  Rabbit. 
Sylvilagus     nuttalli     grangcri,     Black 

Hills  Cottontail. 
Felis  hippolestes,  Mountain  Lion.     C. 
Lynx  uinta.  Mountain  Wildcat. 
Cants  nubilus,  Buffalo  Wolf.     U. 
Canis  testes,  Mountain  Coyote. 
Taxidca  taxus  taxus.  Badger.     U. 
Spilogale     tenuis,     Rocky     Mountain 

Spotted  Skunk. 
Mephitis    hudsonica,    Northern    Plains 

Skunk. 
Mustela   arizonensis,  Arizona   Weasel. 

C. 
Mustela  vison  encrgumenos,  Mink.     C. 
Ursus  horrlbilis,  Grizzly  Bear. 
Corynorhinus  macrotis  palleseens.  Big- 
eared  Bat.     U. 
Myotis  longicrus  interior,  Long-legged 

Bat.     U. 


Breeding  Birds — Transition  Zone. 

[Species  marked  U.  breed  also  in  the  Upper  Sonoran  Zone;  those  marked  C,  also  in  the 

Canadian.] 


Colymbus  nigricollis  californicus, 
American  Eared  Grebe.     U. 

Anas  platyrynchos.  Mallard.     U. 

Cliaulelasmus  streperus,  Gadwall.     U. 

Ncttion  carolinense^  Green-winged 
Teal. 


Steganopus  tricolor,  Wilson  Phalarope. 

U. 
Recurvirostra   americana,   Avocet.      U. 
Gallinago  delicata,  Wilson  Snipe.     C. 
Ca  top  troph  or  us   semi  pa  I  ma  tus   inorna- 

tus.  Western  Willet. 
1  Apparently  nearing  extinction. 


1917.] 


TRANSITION  ZONE. 


35 


Breeding  Birds — Transition  Zone — Continued. 


Podasocys  montanus,  Mountain  Plover. 
V. 

Pedioscetes   phasianellus   columbianus, 
Columbian  Sharp-tailed  Grouse. 

Centrocercus  urophasiarms,  Sage  Hen. 

Accipiter  velox,  Sharp-shinned  Hawk. 
C. 

Accipiter  cooperi,  Cooper  Hawk. 

Butco  swainsoni,  Swainson  Hawk. 

Cryptoglaux  acadica,  Saw-whet  Owl. 

Otus  asio  maxwelliw,  Rocky  Mountain 
Screech  Owl.     U. 

Asio  tcilsonianus,  Long-eared  Owl. 

Coccyzus      erythropthalmus,       Black- 
billed  Cuckoo.    U.1 

Dryobates    villosus   monticola,    Rocky 
Mountain  Hairy  Woodpecker.     G. 

Dryobates  pubcscens  homorus,  Batch- 
elder  Woodpecker. 

Melanerpes       erythroccphalus,       Red- 
headed Woodpecker.     U. 

Asyndesmus      lewisi,      Lewis      Wood- 
pecker. 

Phalwnoptilus   nuttalli  nuttalli,   Poor- 
will.     U. 

Aeronautes       melanoleucus,       White- 
throated  Swift.     U. 

Stellula   calliope,   Calliope    Humming- 
bird.2 

Sayornis  sayus,  Say  Phoebe.     U. 

Myiochanes  richardsoni,  Western  Wood 
Pewee. 

Empidonax  minimus,  Least  Flycatcher. 

Empidonax      icriyliti,      Wright      Fly- 
catcher. 

Empidonax  hammondi,  Hammond  Fly- 
catcher.    C. 

Otocoris    alpestris    leucolcema,    Desert 
Horned.  Lark.     U. 

Pica  pica  hudsonia,  Magpie. 

Cyanoceplwlus   eyanocepliaius,   Pinyon 
Jay.     V. 

Dolichonyx  oryzirorus.  Bobolink.     U. 

Euphagus       eyanocephalus,       Brewer 

Blackbird.     U. 
Calcarius    ornatus,    Chestnut-collared 
Longspur.     U. 

Rhynchoplianes      mccowni,      McCown 
Longspur. 


Pooecetes  gramineus  confinis,  Western 
Vesper  Sparrow.     U. 

Passerculus  sandwichensis  alaudimis, 
Western  Savannah  Sparrow.     U. 

Spisella  passerina  arizonw,  Western 
Chipping  Sparrow. 

Spizella  breweri,  Brewer  Sparrow.     U. 

Junco  aikewi,  White-winged  Junco. 

Melospiza  melodia  montana,  Mountain 
Song  Sparrow. 

PassereUa  iliaca  schistacca,  Slate-col- 
ored Fox  Sparrow. 

Pipilo  maculatus  arcticus,  Arctic 
Towhee. 

Oreospiza  chlorura,  Green  -  tailed 
Towhee. 

Zamelodia  melanoeeplxala,  Black- 
headed  Grosbeak.     U. 

Piranga  ludoviciana,  Western  Tana- 
ger. 

Tachycineta  thalassina  lepida.  North- 
ern Violet-green  Swallow.     C. 

Vireosylva  gilva  sioainsoni,  Western 
Warbling  Vireo. 

Lanirireo  solitarius  pi  umbras.  Plum- 
beous Vireo. 

Dendroica  auduboni,  Audubon  War- 
bler.    C. 

Dendroica  nigrescens,  Black-throated 
Gray  Warbler.3 

Seiurus  aurocapillus.  Oven-bird.     C. 

Oporornis  tolmiei,  Macgillivray  War- 
bler.    C. 

Oreoscoptes  montanus,  Sage  Thrasher. 
U. 

Salpinctes  obsoletus,  Rock  Wren.    U. 

Troglodytes  aedon  parkin  a  ni.  Western 
House  Wren.     U. 

Sitta  carolinensis  nelsoni.  Rocky 
Mountain  Nuthatch. 

Sitta  pygmwa,  Pygmy  Nuthatch. 

Penthestes  atricapillus  septentnonalis. 
Long-tailed  Chickadee.     U. 

Hylocichla  fuscescens  salieicola,  Wil- 
low Thrush. 

Planesticus  m i<ira$orius  propinqv »••>•. 
Western  Robin.    C. 

Sialia  currucoides,  Mountain  Blue- 
bird.    C. 


1  Observed  during  breeding  season. 

2  Taken  during  breeding  season. 

3  Probably  breeds. 


36 


NORTH   AMERICAN   FAUNA. 


[No.  42. 


Plants — Transition  Zone. 

[Species  marked  17.  occur  also  in  the  Upper  Sonoran  Zone;  those  marked  C,  also  in  the 

Canadian.] 

Trees  and  shrubs. 


Pinus  scopulorum,  Yellow  Pine,  Rock 
Pine. 

Junipcrus    scopulorum,    Rocky    Moun- 
tain Juniper.     V. 

Junipcrus    sabina,    Creeping    Juniper, 
Trailing  Savin. 

Populus    angustifolia,    Narrow-leaved 
Cottonwood. 

Salix  bebbiana,  Bebb  Willow.    C. 

Salix  cordata  watsoni,  Diamond  Wil- 
low.    U. 

Salix  mackenziana,  Diamond  Willow. 

Salix  scoulcriana,  Willow. 

Betula    fontinalis,     Rocky     Mountain 
Birch. 

Quercus  macrocarpa,  Bur  Oak. 

Herberts    aquifolium,    Barberry,    Ore- 
gon Grape. 

Grossularia  incrmis,  Gooseberry. 

Ribes  inebrious,  Red  Currant. 

Ribes  americanum,  Currant. 

Edwinia  americana. 

Ccrcocarpus  ledifolius,  Mountain  Ma- 
hogany. 

Ccrcocarpus  intricatus,  Mountain  Ma- 
hogany. 

Ccrcocarpus  parvifolius,  Mountain  Ma- 
hogany.    U. 

Kunzia  tri&entaia. 

Holodiscus  dumosus. 

Opulaster  monoyynus,  Ninebark. 

Opulaster  pubescens,  Ninebark. 

Opulaster  malvaceus,  Ninebark. 

Rubus     deliciosus,     Flowering     Rasp- 
berry. 

iclanchier  alni  folia,  Serviceberry. 
iclanchier  elliptica,  Serviceberry. 
nelanchier  oreophila,  Serviceberry. 
•atcegus  rivularis,  Black  Hawthorn. 

i    "atcegus  cerronis,  Hawthorn. 


Crataegus  sheridana,  Red  Hawthorn. 

Prunus  melanocarpa,  Chokecherry. 

Prunus      poinsylvanica,      Wild      Red 
Cherry.     C. 

Acer  glabrum,  Mountain  Maple.     C. 

Acer    grandidentatum,     Barge-toothed 

Maple. 
jDeanothus  velutinus,  Mountain  Balm. 

Ceanothus  fendleri,  Wild  Tea  Bush. 

Ceanothus  mollissimus. 

Elwagnus  argentea,  Silverberry. 

Cornus  stolonifera,  Cornel. 

Cornus  instolonea,   Cornel. 

Arctostaphylos  uva-ursi,  Bearberry. 

Sambucus  canadensis,  Elderberry. 

Sambucus      melanocarpa,      Mountain 
Black  Elderberry.     C. 

Viburnum  lentago,  Sweet  Viburnum. 

Symphoricarpos    rotundifolius,    Snow- 
berry. 

Symphoricarpos   orcophilus,   Mountain 
Snowberry. 

Symphoricarpos      pauciflorus,      Snow- 
berry. 

Lonicera  glaucescens,  Douglas  Honey- 
suckle. 

Chrysothamnus  wyomingensis.  Rabbit 
Brush. 

Chrysothamnus   pulcherrimus,    Rabbit 
Brush. 

Chrysothamnus  parryi,  Rabbit  Brush. 

Chrysothamnus  frigidus,  Rabbit  Brush. ' 

Artemisia     tridentata,     Black     Sage- 
brush.    U. 

Artemisia  cana,  Gray  Sagebrush.     U. 

Artemisia  trifida,  Sagebrush. 

Artemisia     arbuscula,     Brown     Sage- 
brush. 

Artemisia  frigida.  Sagebrush. 

Artemisia  ludoviciana,  Sagebrush.     V. 


Calochortus  gnnnisoni,  Mariposa  Lily. 
Calochortus  nuttalli,  Mariposa  Lily. 
Zygadenus     venenosus,     Poison     Ca- 
mas.    U. 
Iris  missouriensis,  Blue  Flag. 
Corallorhiza  multiflora.  Coral  Root.    C. 
Humulus  lupulus,  Wild  Hop. 


Herbaceous  plants. 

Letcisia  rediriva.  Bitter  Root. 
Arenaria  congesta,  Sandwort.    C. 
Clematis  ligust  id  folia,  White  Virgin's 

Bower. 
Clematis     douylasi.     Purple     Virgin's 

Bower. 
Clematis   occidentalis,   Virgin's  Bower. 


1917.] 


TRANSITION"   ZONE. 


37 


Plants — Transition  Zone — Continued. 
Herbaceous  plants — Continued. 


Anemone  cylindrica,  Anemone.     U. 

Cyrtorhyncha  ranunculina,  Nuttall 
Buttercup. 

Physaria  didymocarpa,  Double  Blad- 
der-pod. 

Drymocallis  glandulosa,  Glandular 
Cinquefoil.     C. 

Potentilla  effusa,  Cinquefoil. 

Thermopsis  rhombifolia,  Yellow  Ther- 
mopsis. 

Lupinus  argenteus,  Silvery  Lupine. 

Astragalus  succulentus,  Ground 
Plum.     U. 

Astragalus  drummondi,  Drummond 
Milk  Vetch. 

Astragalus  hypoglottis,  Milk  Vetch.    U. 

Astragalus  flexuosus,  Milk  Vetch.     U. 

Astragalus  nitidus,  Milk  Vetch. 

Astragalus  calycosus,  Milk  Vetch. 

Aragallus  lamberti.     U. 

Psoralea  urgophylla,  Silvery  Psoralea. 

Hcdysarum  cinerascens,  Hedysaruin. 

Hcdysarum  uintahense,  Uinta  Hedy- 
sarum.    C. 

Ticia  americana,  Vetch.    U. 

Geranium  fremonti,  Geranium.     C. 

Geranium  richardsoni,  Geranium.    C. 

Linum  lernisi,  Wild  Flax. 


Savastana  odorata,  Holy  Grass. 
Stipa  nelsoni,  Feather  Grass. 
Muhlenbergia  comata,  Dropseed  Grass. 
Agrosiis  hiemalis,  Bent  Grass.     C. 
Koeleria  cristata,  June  Grass.     U. 
Poa     longipedunculata,     Long-stalked 

Spear  Grass.     C. 
Poa  lucida,  Spear  Grass. 
Particular ia  nervata,  Manna  Grass. 


Opuntia  fragilis,  Small-jointed  Cactus. 
V. 

Echinocactus  simpsoni,  Simpson  Ball 
Cactus.     C. 

Pachylophus  montanus,  Evening  Prim- 
rose.    U. 

Harbouria  trachy < pleura,  Water  Hem- 
lock.    C. 

Aralia  nudicaulis,  Wild  Sarsaparilla. 

Gentiana  affinis,  Gentian.     C. 

Ap'ocynum  androswm  i  folium,  Indian 
Hemp. 

Phlox  depressa,  Wild  Phlox. 

Phlox  hoodi,  Wild  Phlox. 

Gilki  congesta,  Gilia. 

Phacelia  linearis,  Phacelia. 

Phacelia  heterophylla,  Phacelia. 

Oreocarya  virgata,  Oreocarya. 

Monarda  menthwfolia,  Horse  Mint.     U. 

Pentstemon  laricifolius,  Beard-tongue. 

Adenostcgia  ramosa.     U. 

Scutellaria  brittoni,  Skullcap.     U. 

Mimulus  floribundus,  Monkey  Flower. 

Orthocarpus  luteus,  Yellow  Orthocar- 
pus. 

Campanula  rotundifolia,  Harebell.     C. 

Antennaria  rcflexa,  Everlasting. 

Balsamorrhiza  sagittata,  Balsam  Boot. 

Balsamorrhiza  iticaua,  Balsam  Boot. 

Chwnactis  douglassi. 


Festuca    ovina    duriuscula,    Fescue 
Grass. 

Bromus  marginatus,  Brome  Grass. 
Bromus  porteri,  Brome  Grass. 
Agropyron  pscudorcpcns,  Wheat  Grass. 

C. 
Agropyron  caninum,  Wheat  Grass. 
Agropyron  spicatum,  Wheat  Grass.     U. 


Agricultural  Utility  of  the  Transition  Zone. 

The  elevated  Transition  area  with  its  vast  extent  of  grazing  lands 
is  now,  as  in  the  past,  the  center  of  the  sheep  and  cattle  industries  of 
Wyoming,  and  there  are  also  extensive  coal  and  oil  fields  in  various 
stages  of  development.  The  principal  timberlands  in  the  Transition 
Zone  are  of  small  extent  and  lie  to  the  east  and  northeast.  They 
comprise  a  moderate  growth  of  yellow7  pine  on  the  Laramie   (see 


38  NORTH    AMERICAN    FAUNA.  [No.  42. 

PL  V,  fig.  1),  Casper,  and  Bear  Lodge  Mountains,  and  heavy  forests 
of  this  valuable  timber  on  the  western  slopes  of  the  Black  Hills.1 

The  climate  is  cool  but  dry  and  healthful,  with  a  rainfall  varying 
from  10  to  12  inches  in  the  arid  parts,  12  to  15  inches  in  the  east-cen- 
tral and  southeast  sections,  and  17  to  21  inches  in  the  Black  Hills  and 
Bear  Lodge  districts  at  the  northeast  where  crops  succeed  moderately 
in  all  except  the  sagebrush  lands.  The  snowfall  is  rather  heavy 
throughout.  This  quantity  of  moisture,  though  not  large,  in  a  meas- 
ure counterbalances  the  cool  climate  of  the  Transition  Zone  and,  com- 
bined with  the  much  greater  area,  gives  this  zone  an  agricultural 
value  in  Wyoming  comparable  to  that  of  the  warmer  Upper  Sonoran. 
Large  areas  of  excellent  grass  land  in  the  mountain  valleys  and  on 
the  higher  plateaus  in  the  upper  part  of  the  zone  are  ideal  either  as 
cattle  range  or  for  the  summer  grazing  of  sheep,  while  a  fair  growth 
of  range  grasses  generally  obtains  even  in  the  lower,  more  arid  sec- 
tions. The  rich  soils  yield  abundantly  either  under  dry  farming 
or  irrigation  wherever  there  has  been  a  proper  selection  of  hardy, 
quick-maturing  crops  adapted  to  a  usually  short  growing  season. 

High-altitude  farming  has  been  thoroughly  tested  at  the  Wyo- 
ming Experiment  Station  at  Laramie  for  a  series  of  years.  Very 
favorable  results 2  were  obtained  with  a  great  variety  of  vegetable  and 
cereal  crops  and  even  certain  fruits,  in  experiments  conducted  in  the 
heart  of  the  Transition  Zone  at  over  7,000  feet  elevation,  where  bleak 
winds  are  unusually  prevalent.  Apples  and  small  fruits  which  are 
grown  with  considerable  difficulty  on  the  Laramie  Plains  succeed 
admirably  in  many  localities  with  greater  natural  protection.  Suc- 
cessful farming  districts  in  the  Wyoming  Transition  include  sections 
of  the  Laramie  Plains,  the  Platte  Valley  above  Saratoga,  Bear  Val- 
ley and  Fort  Bridger  region  in  Uinta  County,  Salt  River  Valley  in 
Lincoln  County,  the  eastern  base  of  the  Bighorn  Mountains  in  Sheri- 
dan and  Johnson  Counties,  and  the  Sundance  region  in  Crook  County. 
In  most  of  these,  oats  and  hardy  cereals,  alfalfa  (two  cuttings),  field 
peas,  potatoes,  and  hardy  vegetables  are  raised  to  great  perfection. 
Hay,  forage,  garden  vegetables,  and  a  limited  crop  of  small  grain  are 
very  generally  grown  on  stock  ranches  even  in  the  colder  parts  of 
the  zone. 

CANADIAN  ZONE. 

The  Canadian  Zone,  the  region  of  coniferous  Boreal  forest,  is  the 
most  important  of  the  Boreal  transcontinental  life  areas.  It  extends 
far  southward  in  the  principal  mountain  masses  of  the  Western 

1  Open  sage  country  usually  fills  the  zone  on  the  basal  slopes  of  the  high  ranges  in 
western  Wyoming.  The  growth  of  Douglas  spruce  along  its  upper  border  in  this  region 
is  generally  scattering,  while  the  yellow  pine  is  of  rare  occurrence. 

2  Discussed  in  bulletins  of  the  Wyoming  Experiment  Station,  which  contain  also  valu- 
able cultural  and  other  data  based  on  tests  made  at  the  station  at  Laramie  and  at  the 
experimental  farms  at  Sundance  and  Saratoga. 


1917.]  CANADIAN   ZONE.  39 

States,  and  over  much  of  the  Rocky  Mountain  region  covers  the 
middle  slopes  of  the  high  ranges  and  the  summits  and  upper  slopes 
of  mountains  of  medium  elevation.  It  is  uniformly  and  conspicu- 
ously characterized  from  Montana  to  Colorado  by  forests  of  spruce, 
fir,  lodgepole  pine,  and  aspen,  and  by  a  large  variety  of  Boreal  under- 
shrubs  and  plants. 

In  Wyoming  the  greatest  extent  of  Canadian  Zone  country  is  in 
the  mountainous  northwest.  Here  the  zone  includes  most  of  the 
extensive  undulating  forested  plateau  of  Yellowstone  Park;  large 
rolling  or  hilly  tracts  of  mixed  forest  and  open  country  on  the 
borders  of  Jackson  Hole  (PI.  VIII,  fig.  1),  in  the  basin  of  Hoback 
River,  at  the  head  of  Green  River,  and  on  the  southern  end  of  the 
Wyoming  Range ;  and  the  forested  slopes  of  the  Wind  River,  Absa- 
roka,  Teton,  Gros  Ventre,  Salt  River,  Snake  River,  and  Wyoming 
Ranges  (PL  VIII,  fig.  2)  from  near  their  bases  to  the  upper  limit 
of  large  tree  growth  at  9,000  to  10,000  feet  elevation.  Elsewhere 
the  Bighorn  and  Medicine  Bow  Ranges  and  the  Sierra  Madre  are 
extensively  Canadian,  and  the  Laramie  Mountains  and  Casper  Range 
have  considerable  areas  on  their  summits.  Elevations  of  medium 
altitude  which  are  capped  with  Canadian  Zone  forests  and  also  have 
small  areas  on  their  cool  slopes  are  the  Black  Hills,  the  Bear  Lodge, 
Rattlesnake,  Green,  Ferris,  Seminole,  and  Shirley  Mountains,  the 
northern  shoulders  of  the  Uinta  Mountains,  and  high  plateaus  along 
the  southern  boundary  of  the  State  between  Green  River  and  the 
Red  Desert.  Traces  of  the  zone,  indicated  usually  by  aspens  and 
lodgepole  pines,  or  merely  by  dense,  scrubby  thickets  of  aspens,  are 
on  the  upper,  cool  slopes  of  the  Aspen  Mountains,  the  Bear  River 
Divide,  a  few  desert  peaks  along  the  continental  watershed  between 
South  Pass  and  Steamboat  Mountain,  on  Heart  Mountain  north  of 
Cody,  and  on  Pyramid  and  Heaths  Peaks  along  the  upper  Platte. 

The  lofty  Wind  River  Range  of  comparatively  straight  axis  pre- 
sents a  graphic  view  of  the  Canadian  forest  belt,  which  is  main- 
tained usually  at  uniform  elevation  and  in  full  vertical  width  (about 
2,000  feet)  on  both  slopes,  its  lower  border  sharply  defined  where 
the  forests  meet  the  open  basal  sagebrush  country  of  the  Transition 
Zone.1  The  upper  border  of  the  Canadian  Zone  is  obscure  in  these 
mountains,  as  elsewhere,  the  change  from  heavy  forest  growth  to 
the  narrow  Hudsonian  timberline  belt  of  dwarfed  forest  being 
gradual  and  almost  imperceptible. 

The  main  forest  composition  on  the  Wyoming  mountains  is  very 
uniform,  with  forests  of  lodgepole  pine  and  aspen  in  the  lower  half 
of  the  zone  and  a  heavy  stand  of  Engelmann  spruce,  or  more  often 

1  The  lower  edge  of  the  Canadian  Zone  is  less  clearly  marked  on  the  eastern  slopes  of 
the  Bighorn  Mountains  and  elsewhere  in  eastern  Wyoming  where  forests  of  yellow 
pine  fill  much  of  the  upper  Transition. 


40  NORTH   AMERICAN    FAUNA.  [No.  42. 

a  mixed  forest  of  spruce  and  fir,  higher  up.  Other  trees  of  less 
extensive  growth  and  more  restricted  distribution  are  the  Douglas 
spruce  in  the  lower  part  of  the  zone,  chiefly  in  the  northwestern 
mountains;  the  blue  spruce  and  balsam  poplar  fringing  streams 
along  the  lower  edge  of  the  zone  in  the  south  and  west,  and  in  the 
northwest,  respectively ;  and  the  canoe  birch  in  the  Black  Hills  and 
Bear  Lodge  Mountains.  Fire-swept  tracts  usually  are  first  covered 
with  a  dense  growth  of  young  aspens,  which  are  of  more  rapid 
growth  than  conifers.  In  the  Sierra  Madre  the  fresh  growth  on 
burned-over  areas  is  occasionally  of  fir  over  original  lodgepole  pine 
forest. 

In  the  lower  part  of  the  Canadian  forest  belt  are  considerable 
areas  of  partly  open  mountain  meadows  and  parks,  and  more  rarely, 
of  open  slopes.  Characteristic  tracts  are  the  8,000-foot  watershed 
between  the  Hoback  and  Green  Eivers,  where  groves  of  aspen  inter- 
mixed with  a  little  lodgepole  pine  and  fir  alternate  over  a  gently 
rolling  country  with  open  parks  covered  with  low  matted  sagebrush, 
Frasera,  and  Balsamorrhiza,  or  with  beautiful  grassy  meadows  bril- 
liantly colored  in  summer  with  flowers  of  shrubby  cinquefoil,  lark- 
spur, lupine,  geranium,  iris,  and  painted  cup;  the  grassy  meadows 
and  bordering  sage  benches  of  the  Du  Noir  Valley  at  the  head  of 
Wind  River;  extensive  willow-grown  meadows  and  flats  at  the  north- 
ern end  of  Jackson  Hole  and  in  Yellowstone  Park;  and  open  grass 
or  sage  slopes  on  the  western  side  of  the  Bighorn  Mountains  be- 
tween 8,000  and  9,000  feet  elevation.  On  the  moderately  inclined 
eastern  side  of  the  Wyoming  Range  southwest  of  Bigpiney  are 
unusually  open  mountain  slopes.  Here  grass  and  sage  country  ex- 
tend in  many  places  on  ridges  and  south  slopes  to  the  10,000-foot 
crest  of  the  range,  alternating  regularly  with  dense  tracts  of  Douglas 
spruce,  lodgepole  pine,  and  Engelmann  spruce  forest  on  all  north 
slopes  and  in  gulches. 

The  usual  factors  of  base  level,  latitude,  slope  incline  and  ex- 
posure, and,  to  a  certain  extent,  air  currents,  affect  in  varying  degree 
the  altitude  of  the  Canadian  Zone.  Base  level  appears  to  be  more 
potent  than  latitude  in  Wyoming.  The  variation  from  8,500  to  10,- 
500  feet  on  southwest  slopes  and  7,500  to  9,500  feet  on  northeast 
slopes  in  the  mountains  along  the  southern  boundary  and  bordering 
Green  River  Basin,  to  8,000  to  9,500  feet  on  southwest  slopes  and 
7,000  to  9,000  feet  on  northeast  slopes  in  the  northern  mountains,  is 
largely  latitudinal.  The  depression  to  6,000  feet  on  the  cap  of  the 
Black  Hills  and  Bear  Lodge  Mountains,  however,  is  probably  due 
to  the  low  base  level  of  the  adjacent  plains  on  the  north  and  east. 

The  high  level  of  the  Canadian  Zone  (above  8.000  feet)  on  the 
warm  western  slope  of  the  Bighorn  Mountains  east  of  Ionia  is  due 
to  bold  southwest  exposure,  which  more  than  offsets  the  lowering 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  VIII. 


Fig.  1.— Snake  River  Valley  (6,500  Feet)  near  Moran,  Jackson  Hole. 

Mixed  forest  conditions  at  lower  edge  of  Canadian  Zone  are  shown.    Scattered  groves  of  aspen, 
balsam  poplar,  and  blue  spruce  on  valley  flats;  aspen  and  lodgepole  pine  forests  on  hills. 


Fig.  2.— Heavy  Engelmann  Spruce  Forest  Wyoming  Range  West  of  Merna 

(9,000  Feet). 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  IX. 


Fig.  1.— Sylvan  Lake,  Yellowstone  National  Park,  in  the  Enqelmann  Spruce 

Belt. 


Fiq..2.— Heavy  Stand  of  Lodgepole  Pine,  North  Slope  of  Ferris  Mountains 

(8,500  Feet). 


1917.]  CANADIAN    ZONE.  41 

influence  of  the  low  altitude  of  the  Bighorn  Basin  at  their  western 
bases.  The  Ferris  and  Green  Mountains,  small  isolated  ranges  on 
the  northern  border  of  the  Red  Desert,  show  graphically  the  effect 
of  slope  exposure  on  zone  level.  The  Ferris  Range,  of  9,500  feet 
elevation,  is  narrow  and  sharp-crested,  with  very  abrupt  southwest 
slopes  facing  the  desert.  These  hot,  exposed  slopes  are  open  Transi- 
tion Zone,  and  crowd  the  Canadian  element  above  9,000  feet,  where 
it  is  indicated  just  below  the  summit  by  thickets  of  aspen  and  lodge- 
pole  pine  in  gulches,  and  by  Rocky  Mountain  white  pine  on  exposed 
slopes  and  ridges.  Just  over  the  9,500-foot  crest  on  the  cool  exposure 
are  the  Engelmann  spruces  and  firs  of  the  Canadian  Zone,  somewhat 
dwarfed  at  first  but  soon  encountered  in  large  and  dense  growth. 
The  spruces  and  firs,  with  lodgepole  pines  and  aspens  lower  down, 
form  a  heavy  forest  on  the  northeast  slopes  down  to  7,800  feet,  while 
descending  tongues  in  gulches  along  cold  streams  extend  500  feet 
lower.  The  Canadian  Zone  on  the  Ferris  Range  thus  exhibits  about 
the  maximum  variation  in  zone  level  due  to  slope  exposure — fully 
1,500  feet. 

In  Jackson  Hole,  the  Yellowstone  Valley,  along  both  forks  of  the 
Shoshone  River,  and  at  the  heads  of  many  other  narrow  valleys 
deeply  penetrating  the  mountain  mass  of  northwest  Wyoming  the 
Canadian  element  reaches  a  low  elevation,  and  on  cold  slopes  is 
often  unmixed  with  Transition  species  at  6,000  feet.  Over  this  re- 
gion the  mean  summer  temperature  is  low,  the  cold  air  of  the  sur- 
rounding mass  of  Boreal  country  settling  into  the  valleys  and  can- 
yons at  night,  and  frequent  frosts  occur  during  the  warmest  months. 
Furthermore,  these  mainly  steep-walled  valleys  receive  a  minimum  of 
sunlight,  and  many  slopes  are  shaded  during  the  warmest  part  of  the 
day.  Under  these  conditions  the  zone  occasionally  has  a  vertical 
breadth  of  nearly  3,000  feet,  since  the  factors  which  cause  the  abnor- 
mal depression  of  its  lower  boundary  do  not  appreciably  affect  the 
upper  limit. 

The  cooling  influence  of  cold  streams  and  of  descending  cold  air 
currents  which  flow  down  gulches  and  canyons  regularly  carries  nar- 
row tongues  of  Canadian  Zone  far  below  the  average  level  on  moun- 
tain slopes.  This  is  very  noticeable  on  some  of  the  streams  at  the 
southwestern  base  of  the  Wind  River  Range.  The  clear  icy  waters 
of  Pine  Creek,  the  outlet  of  Fremont  Lake,  carry  a  broad  fringe  of 
lodgepole  pines,  aspens,  and  a  pure  Canadian  undergrowth  almost  to 
its  junction  with  the  New  Fork  at  7,000  feet  elevation,  fully  1,000 
feet  below  the  mean  lower  border  of  the  zone  on  the  southwest  slope 
of  these  mountains.  The  waters  of  the  Big  Sandy,  40  miles  to  the 
southeast,  also  carry  Canadian  species  far  down  into  open  Transition 
sagebrush  country.  Streams  which  break  out  of  the  steep  northern 
escarpment  of  the  Casper  Range  through  deep,  shaded  gulches  are 


42  NORTH  AMERICAN   FAUNA.  [No.  42. 

heavily  fringed  with  aspen  groves  and  Canadian  undershrubs  on  the 
otherwise  open  6,000-foot  basal  slopes,  1,000  feet  below  the  usual 
altitude  of  these  species  on  the  northern  slopes  of  this  range. 

The  Canadian  Zone  is  marked  in  Wyoming  as  a  cool  region  of  con- 
siderable humidity,  but  the  percentage  of  bright  sunny  days  through- 
out the  year  is  high.  Much  of  the  precipitation  is  in  the  form  of 
frequent  sharp  showers  in  summer  and  heavy  snows  in  winter,  but 
there  are  occasional  rains  of  greater  duration  in  spring  and  fall. 
The  chief  value  of  this  zone  is  its  natural  adaptation,  in  abundant 
forest  and  plant  cover  and  cool  summer  temperature,  to  moisture 
retention.1  The  extensive  forests,,  its  greatest  natural  resource,  are 
now  largely  under  Federal  control.  These  are  of  great  value,  espe- 
cially the  forests  of  lodgepole  pine  and  Engelmann  spruce  (PI.  IX), 
but  large  areas  are  as  yet  inaccessible.  Winter  temperatures  are  low, 
occasionally  reaching  — 45°  F.,  and  in  the  small  areas  physically 
adapted  to  agriculture  the  prevalent  summer  frosts  preclude  the 
growing  of  more  than  a  few  of  the  hardiest  crops  and  vegetables  for 
ranch  use. 

Characteristic    Species — Canadian    Zone. 

A  rich  fauna  and  flora  uniformly  characterize  the  Canadian  Zone 
in  Wyoming.  Forest  species  predominate,  many  of  them  identical 
with  or  closely  related  to  Boreal  types  of  transcontinental  range  from 
Labrador  to  Alaska ;  others  are  peculiar  to  the  Rocky  Mountain  re- 
gion ;  and  a  small  number  have  a  restricted  range.  The  canoe  birch, 
beaked  hazelnut^  Douglas  honeysuckle,  northern  chipmunk,  and  a 
few  other  species  of  wide  Boreal  dispersion  occur  only  in  the  out- 
lying Black  Hills  and  Bear  Lodge  Mountains  at  the  northeast,  and 
the  northern  varying  hare  only  in  the  Bighorn  Mountains,  but 
elsewhere  the  characteristic  species  are  very  uniformly  distributed  in 
the  different  mountain  ranges.  With  the  exception  of  many  of  the 
breeding  birds,  which  merely  make  their  summer  home  in  the  Cana- 
dian Zone,  all  are  species  adapted  to  a  region  of  long,  cold  winters 
and  short,  cool  summers. 

Mammals — Canadian  Zone. 

[Species   marked   T.   occur  also  in   the  Transition   Zone;    those   marked   H.,   also   in   the 

Hudsonian.] 


Cervus     canadensis    canadensis,    Elk. 

T,  H. 
Aices      americanus      shirasi,      Shiras 

Moose. 
Schtrits    frcnionti    frcmonti,    Fremont 

Spruce  Squirrel. 
Sciurus    hudsonims    baileyi,    Bighorn 

Red  Squirrel. 


Sciurus  hudsonicus  rentonan,  Rocky 
Mountain  Red  Squirrel. 

Glaucomys  sabrinus  bangsi,  Rocky 
Mountain  Flying  Squirrel. 

Eutamias  borealis,  Northern  Chip- 
munk. 

Eutamias  operariits,  Colorado  Chip- 
munk. 


i  See  pp.  52-53. 


1917.] 


CANADIAN    ZONE. 


43 


Mammals — Canadian  Zone — Continued. 


Eutamias  umbrinus,  Uinta  Chipmunk. 
Eutamias      liiteiventris,      Buff -bellied 

Chipmunk.     T. 
Callospermophilus    lateralis    lateralis, 

Say  Ground  Squirrel.     T. 
Callospermophilus  lateralis  caryi,  Wind 

River  Mantled  Ground  Squirrel. 
Callospermophilus     lateralis     castanu- 

rus,   Chestnut-tailed   Ground    Squir- 
rel. 
Marmata       flaviventris       nosophora, 

Golden-mantled  Marmot.     H. 
Marmot  a    flaviventris     luteola,     Park 

Marmot.     H. 
Marmota    flaviventris    dacota,    Black 

Hills  Marmot.     T. 
Peromyseus     maniculatus     artemisiw, 

Sagebrush  White-footed  Mouse.     '/. 
Peromyseus       maniculatus       ruflnus, 

Tawny  White-footed  Mouse. 
Phenacomys       orophilus,       Mountain 

Phenacomys. 
Evotomys    gapperi    galei,    Gale    Red- 
backed  Mouse. 
Evotomys     brevicaudus,    Black    Hills 

Red-backed  Mouse. 
Mierotus      mordax     mordax,      Rocky 

Mountain  Meadow  Mouse. 
Mierotus       longicaudus,       Long-tailed 

Meadow  Mouse.     T. 
Mierotus    richardsoni    macropus,   Big- 
footed  Meadow  Mouse. 
Mierotus   nanus   nanus,   Dwarf   Field 

Mouse.     T,  H. 
Castor  canadensis,  Beaver.    T. 
Thomomys    fossor,    Colorado    Pocket 

Gopher. 
Thomomys       uinta,       Uinta       Pocket 

Gopher.     H. 
Thomomys      fuscus      fuscus,      Brown 

Pocket  Gopher. 


Thomomys  talpoides  caryi,  Bighorn 
Pocket  Gopher.     II. 

Zapus  princeps  princeps,  Rocky  Moun- 
tain Jumping  Mouse. 

Erethizon  epixanthum,  Yellow-haired 
Porcupine.      T. 

Lepus  americanus  americanus,  North- 
ern Varying  Hare. 

Lepus  bairdii  bairdii,  Snowshoe  Rab- 
bit. 

Fells  hippolestes,  Mountain  Lion.     T. 

Lynx  canadensis  canadensis,  Canada 
Lynx. 

Vulpes  maerourus.  Mountain  Red  Fox. 

Mustela  arizonensis,  Arizona  Wea- 
sel.    T. 

Mustela  cicognanii  leptus,  Dwarf  Wea- 
sel. 

Mattes  caurina  origenes,  Rocky  Moun- 
tain  Marten. 

Lutra  canadensis  canadensis,  Otter.    T. 

Gulo  luscus,  Wolverene. 

Ursus  americanus,  Black  Bear.     T. 

Ursus  imperator,  Yellowstone  Park 
Grizzly   Bear.     T. 

Ursus  washake,  Washakie  Grizzly 
Bear,  Silver-tip.     T. 

Sorex  personatus  personatus,  Masked 
Shrew.     H. 

Sorex  obscurus  obscurus,  Rocky 
Mountain  Shrew.    H. 

Sorex  vagrans  dobsoni,  Dobson 
Shrew. 

Neosorex  palustris  navigator,  White- 
bellied  Water  Shrew. 

Nycteris  cinerea.  Hoary  Bat.     T. 

Lasionycteris  noctivagans,  Silver- 
haired   Bat.      T. 

Myotis  lucifugus  carissima.  Little 
Brown  Bat.     T. 


Breeding  Birds — Canadian  Zone. 

[Species   marked   T.  breed  also  in   the  Transition   Zone;   those   marked   H.,   also   in   the 

Iludsonian.] 


Pelecanus  erythrorhynchos,  White  Pel- 
ican. 

Mergus  americanus,  Merganser.     T. 

Clangula  islandica,  Barrow  Golden- 
eye.     H. 

Charitonetta  albeola,  Buffle-head. 


Rranta  canadensis,  Canada  Goose.  T. 
Olor  buccinator,1  Trumpeter  Swan.  T. 
Dendragapus  ob  s  cur  u  s  richardsoni, 

Richardson  Dusky  Grouse.     T. 
Picoides    arcticus,    Arctic    Three-toed 

Woodpecker. 


1  Probably  no  longer  breeding  in  Wyoming. 


44 


NORTH   AMERICAN   FAUNA. 


[No.  42. 


Breeding  Birds — Canadian  Zone — Continued. 


Picoides  americanus  dorsalis,  Alpine 
Three-toed  Woodpecker.    H. 

Sphyrapicus  varius  nuchalis,  Red- 
naped  Sapsucker.     T. 

Sphyrapicus  thyroideus,  Williamson 
Sapsucker. 

Selasphorus  platycercus,  Broad-tailed 
Hummingbird.     T. 

Kuttallornis  borealls,  Olive-sided  Fly- 
catcher. 

Empidonax  difflcilis,  Western  Fly- 
catcher.    T.  / 

Cyanocitta  stelleri  annectens,  Black- 
headed  Jay. 

Perisorcus  canadensis  capitalis,  Rocky 
Mountain  Jay.     H. 

Carpodacus  cassini,  Cassin  Purple 
Finch. 

Loxia  curvirostra  minor.  Crossbill. 

Spinus  pinus,  Pine  Siskin. 

Zonotrichia  leucophrys,  White-crowned 
Sparrow.     H. 

J  unco  hyemalis  mearnsi,  Pink-sided 
Junco.     H. 


J  unco  phceonotus  caniceps,  Gray-head- 
ed  Junco.     H. 

Mclospiza  lincolni,  Lincoln  Sparrow. 

Wilsonia  pusilla  pileolata,  Pileolated 
Warbler.    H. 

Cinclus  mexicanus  unicolor,  Dipper, 
Water  Ouzel.     T. 

N annus  hiemalis  pacificus,  Western 
Winter  Wren. 

Sitta  canadensis,  Red-breasted  Nut- 
hatch. 

Penthestes  gambeli,  Mountain  Chicka- 
dee.    T. 

Regulus  satrapa,  Golden  -  crowned 
Kinglet.     H. 

Reyulus  calendula,  Ruby -crowned 
Kinglet. 

Myadestes  townsendi,  Towusend  Soli- 
taire.    T. 

Ilylocichla  ustulata  swainsoni,  Olive- 
backed  Thrush.     T. 

HylocicJila  guttata  auduboni,  Audubon 
Hermit  Thrush. 

Sialia  currucoides,  Mountain  Blue- 
bird.    T. 


Plants — Canadian  Zone. 

[Species   marked   T.   ooeur  also   in   the   Transition    Zone;   those  marked   IT.,  also   in   the 

Hudsonian.] 

Trees  and  shrubs. 


Pinus  murrayana,  Lodgepole  Pine. 

Pinus  flexilis.  Rocky  Mountain  White 
Pine.     T,  H. 

Picea  engelmanni,  Engelmann  Spruce. 

Picea  parry  ana,  Blue  Spruce. 

Abies  (concolor  ?),  White  Fir. 

Pseudotsuga  mueronata,  Douglas 
Spruce.     T. 

Juniperus  sibirica,  Low   Juniper.     H. 

Populus  tr<  tiiuloides,  Aspen. 

Populus   balsamifera,  Balsam   Poplar. 

Salix  pyrifolisi  obscura,  Willow. 

Salix  nelsoni,  Nelson  Willow.     H. 

Betula  papyrifera,  Canoe  Birch. 

Betula  glandulosa,  Dwarf  Birch. 

Corylus  rostra ta,  Hazelnut.     T. 

Alnus  tenuifolia,  Alder. 

Ribes  petiolare,  Mountain  Black  Cur- 
rant. 


Ribes  wolfl,  Blue  Currant. 

Ribes  lacustre,  Currant. 

Ribes  viscosissimuin.  Currant. 

Spircea  lucida,  Meadowsweet.     T. 

Rubus  parviflorus,  Thimbleberry. 

Rubus   strigosus.    Red    Raspberry.     T. 

Dasiphora  fruticosa.  Shrubby  Cinque- 
foil.     H. 

Rosa  sayi,  Rose.     T. 

Morbus  scopulina.  Mountain  Ash. 

Pachystima  myrsin ites. 

Rhamnus  alnifolia.  Buckthorn. 

Lepargyrea  canadensis,  Canadian  Buf- 
faloberry. 

Ledum  glandulosum,  Labrador  Tea. 

Mensiesia  ferrugin  ra. 

Vaccinium  oreophilum,  Mountain  Blue- 
berry. 


1917.] 


CANADIAN    ZONE. 


45 


Plants — Canadian  Zone — Continued. 
Trees  and  shrubs — Continued. 


Vaccinium  erythrococcum,  Red  Bil- 
berry.    H. 

Vaccinium   occidentale,   Blueberry. 

Vaccinium  cwspitosum,  Blueberry. 

Sambucus  microbotrys,  Red  Elder- 
berry. 


Lonicera  glaucescens,  Douglas  honey- 
suckle. 

Lonicera  involucrata,  Involucred  Fly- 
honeysuckle. 

Lonicera  utahensis,  Honeysuckle. 


Herbaceous  plants. 


Veratrum  tenuipetalum,  White  Helle- 
bore. 

Zygadenus  elegans,  Beautiful  Camas. 
H. 

Calochortus  pavonaccus,  Yellow  Mari- 
posa. 

Erytlironium  parriflorum,  Dog-tooth 
Violet.     H. 

Streptopus  amplexifolius,  Twisted- 
stalk. 

Disporum  trachyearpum.     T. 

Limnorchis   borealis,  Bog  Orchid. 

Calypso  bulbosa,  Calypso. 

Rum  ex  paucifolius,  Dock.     H. 

Claytonia  rosea,  Spring  Beauty.     H. 

Aetata  arguta,  Baneberry. 

Aetata  rubra,  Baneberry. 

Aquilegia  cwrulea,  Blue  Columbine. 

Aquilegia  flavescens,  Yellow  Colum- 
bine. 

Aquilegia  oreophila,  Columbine.     H. 

Anemone  lithophila,  Anemone. 

Aconitum  columhianum.  Monkshood. 

Clematis  pseudalpina,  Purple  Virgin's 
Bower. 

Ranunculus  inamemus,  Crowfoot. 

Thlaspi  glaucum,  Penny  Grass.     H. 

Parnassia  flmbriata,  Grass-of-Parnas- 
sus. 

lleuehera  parvifolia,  Alum  Root.     II. 

Micranthes  arguta,  Saxifrage. 

Fragaria  paueiflora.  Strawberry.     T. 

Sieversia  cUiata.,  Mountain  Avens. 

Astragalus  alpinus.  Milk  Vetch. 

Trifolium  rydbergi,  Clover.     T. 

Trifolium  ancmophilum.  Clover.     T. 

Geranium  parryi,  Geranium. 


T. 


T. 


Geranium  cwspitosum,  Geranium. 

Viola  canadensis  rydbergi,  Violet. 

Viola,  bellidifolia,  Violet.     H. 

Epilobium  spp. 

Heracleum  lanatum. 

Pyrola  secunda,  Wintergreen. 

Pyrola  uliginosa,  Wintergreen. 

Pyrola.  chlorantha,  Wintergreen. 

Pyrola  elliptica,  Wintergreen. 

Pyrola  pieta,  Painted  Wintergreen. 

Pterospora  andromeda,  Pinedrops. 

Chimaphila  umbellata,  Pipsissewa. 

Moneses  uniflora,  One-flowered  Win- 
tergreen. 

Gentiana  forwoodi,  Closed  Gentian. 

Gentiana  elegans,  Mountain  Fringed 
Gentian. 

Frasera  speeiosa,  Frasera. 

Pentstemon  glaucus.  Beard-tongue. 

Pentstemon  fruticosus,  Beard-tongue. 
T. 

Mimulus  langsdorfii,  Monkey  Flower. 

Mini  til  us  letcisi,  Crimson  Monkey 
Flower. 

Elephant  clla  groenlandica,  Elephant- 
head.     H. 

Pedieularis  raeemosa,  Purple  Louse- 
wort. 

Castilleja  sulphured.  Painted  Cup. 

Linnwa  americana,  Twinflower. 

Valeriana  aeutiloba,  Valerian.     If. 

Anaphalis  snbalpina.  Pearly  Everlast- 
ing. 

Antennaria  puleherrima,  Everlasting. 
T. 

Wyethia  ample.ricaulis. 

Arnica  cordifolia,  Heart-leaved  Arnica. 


46 


NORTH   AMERICAN   FAUNA. 


[No.  42. 


Plants — Canadian  Zone — Continued. 
Grasses. 


Phleum  alpinum,  Alpine  Timothy.     H. 
Xlopecurus      occidental  is,       Mountain 

Foxtail. 
Catamagrostis   canadensis    acuminata, 

Mountain  Reed  Grass. 
Deschampsia   ccespitosa,    Tufted    I  lair 

Grass.     T. 


Danthonia  intermedia,  Mountain   Oat 

Grass. 
Poa  reflecea,  Mountain  Bluegrass.    II. 
Festuca  thurberi,  Thnrber  Fescue. 
Agropyron    violaceum,    Wheat    Grass. 

H. 
Elymus  glaueus,  Rye  Grass.     T. 


HUDSONIAN  ZONE. 

On  the  high  mountain  ranged  of  the  Western  States  the  Canadian 
forest  belt  is  fringed  along  its  upper  edge  by  the  narrow  Hud- 
sonian  strip  of  dwarfed  forest  and  depauperate  vegetation,  which 
gives  way  to  bare  Alpine  slopes  along  a  sharply  defined  climatic 
boundary  known  as  timberline,  corresponding  to  the  transconti- 
nental limit  of  trees  at  the  southern  edge  of  the  Arctic  tundras. 
The  Hudsonian  Zone  occupies  a  relatively  small  area  in  Wyoming. 
It  encircles  the  Alpine  summits  and  crests  of  the  northwestern 
ranges,  including  the  Bighorn  Mountains,  and  small  areas  cap  all 
ranges  or  isolated  peaks  which  approximate  timberline  altitudes.  A 
belt  bounds  the  Alpine  cap  of  the  Snowy  Mountains  at  the  northern 
end  of  the  Medicine  Bow  Range,  and  small,  widely  separated  tracts 
or  islands  are  on  Bridger  Peak  and  neighboring  summits  of  the 
Sierra  Madre,  on  Elk  Mountain,  and  on  Laramie  Peak. 

In  the  southern  mountains  the  forests  commence  to  dwarf  almost 
imperceptibly  at  from  about  10,000  to  10,500  feet  altitude,  accord- 
ing to  slope  exposure,  the  elevation  decreasing  with  higher  latitude 
to  9,000  or  9,500  feet  in  northern  Wyoming.  The  vertical  breadth 
of  the  Hudsonian  belt  varies  with  slope  and  soil  conditions  from  a 
few  hundred  to  1,000  feet.  The  peculiar  ruggedness  and  broken, 
incised  character  of  many  mountains  greatly  obscures  this  belt, 
while  in  places  sheer,  perpendicular  cliffs  and  avalanches  greatly 
contract  it  or  even  narrowly  interrupt  continuity.  Avalanches 
sweep  away  every  vestige  of  tree  growth  and  in  many  places  replace 
the  original  soil  with  extensive  fields  of  slide  rock.  The  result  is 
not  a  climatic  change,  however,  and  can  not  be  considered  in  delimit- 
ing the  zone.  The  Hudsonian  strip  is  usually  widest  in  cold  gulches 
with  abundant  soil,  and  is  narrow  on  exposed,  scantily  soiled  ridges. 
In  fact,  soil  conditions  often  counterbalance  the  elevating  influence 
of  slope  exposure,  and  tree  growth  may  be  found  as  high  on  cold 
well-soiled  slopes  as  on  warm  rocky  inclines. 

An  almost  Arctic  climate  prevails  in  the  Hudsonian  Zone,  which 
in  winter  is  buried  under  deep  snow  and  in  summer  is  flecked  with 
huge  drifts,  many  of  which  never  entirely  leave  protected  gulches. 
The   deep-soiled   slopes   are   thoroughly   saturated    in    summer   by 


1917.]  HUDSONIAN   ZONE.  47 

melting  snow  and  frequent  showers  and  squalls  of  snow  or  sleet,  and 
bogs  and  small  lakes  abound  in  all  level  situations  and  natural 
basins.  Exposed  to  high  winds  throughout  the  year,  the  conifers 
are  in  ragged,  fantastic,  and  usually  one-sided  growth,  and,  along 
with  the  shrubs  and  many  larger  plants,  evidence  an  adverse  cli- 
mate in  stunted  and  otherwise  deficient  development.  (PI.  X.)  The 
middle  of  May  found  the  timberline  region  on  the  Wind  River 
Range  near  Dubois  still  in  the  grasp  of  wintry  weather,  with  few 
plants  in  flower  and  the  low  willows  and  shrubby  cinquefoil  not 
yet  in  leaf;  while  on  the  Bighorn  Mountains  east  of  Hyattville  the 
alpine  willows  had  not  put  forth  leaves  by  June  5.  Many  plants 
were  past  flowering,  and  herbaceous  vegetation  was  partly  dried 
up  on  the  Wyoming  Range  at  10,400  feet  altitude,  August  9;  and 
the  timberline  slopes  on  the  Tetons  were  sere  and  brown  on  August 
30  except  for  the  hardy,  late-flowering  blue  gentians. 

Although  the  climate  is  rigorous  for  eight  months  of  the  year 
and  myriads  of  mosquitoes  greet  one  in  summer,  this  highly  inter- 
esting region  well  repays  the  arduous  climb  entailed  to  reach  its 
confines.  The  gently  rounding  crests  of  the  Wyoming  Range  south 
of  Hoback  Peak  are  peculiarly  attractive.  Grassy  openings  and 
parks  mingle  with  scattered  clumps  and  mats  of  Engelmann  spruce 
and  alpine  fir;  fields  of  scarlet  painted  cup,  blue  larkspur,  white 
columbine,  and  purple  lupine  enliven  a  landscape  flecked  with  white 
banks  of  melting  snow;  and  a  moderate  incline  adds  to  the  beauty 
of  the  region.  The  Hudsonian  area  on  the  southwest  side  of  the 
Wind  River  Range  is  of  very  different  character.  South  of  Fre- 
mont Peak  (PI.  XI,  fig.  1)  it  occupies  a  sloping  granitic  plateau  a 
mile  or  two  wide,  and  between  10.500  and  11.200  feet  altitude,  at  the 
base  of  abrupt  Alpine  peaks  of  nearly  14,000  feet  elevation.  Its  ex- 
ceedingly rough  surface  of  a  seemingly  endless  succession  of  bare 
granite  hummocks,  studded  with  countless  clear,  snow-fed  lakes  in 
rocky  basins,  makes  travel  extremely  arduous.  Vegetation  is  scanty, 
and  coniferous  growth  is  very  scattering  over  this  unusually  rocky 
region.  Hudsonian  vegetation  is  most  abundant  on  deep-soiled 
slopes,  as  on  Whiskey  Mountain  in  the  Wind  River  Range  south  of 
Dubois,  and  along  the  eastern  slope  of  Needle  Mountain  in  the 
Absaroka  Range  (PI.  XI,  fig.  2). 

Characteristic   Species — Hudsonian   Zone. 

Trees  of  the  timberline  belt  in  Wyoming  are  the  alpine  fir,  white- 
barked  pine,  and  Engelmann  spruce.  The  first  two  are  characteristic 
Hudsonian  trees,  but  the  spruce  extends  up  from  the  Canadian  Zone. 
The  spruces  and  firs,  mainly  on  cool  exposures,  exhibit  to  a  marked 
degree  the  depressed  growth  due  to  high  altitude  and  are  usually 
prostrate  mats  at  extreme  timberline.  The  white-barked  pines  of  the 
northwest  ranges  show  less  dwarfing,  but  usually  are  very  ragged 


48  NORTH    AMERICAN    FAUNA.  [No.  42. 

and  one-sided  as  a  result  of  the  prevalent  winds.  They  occur  in 
scattering  growth  or  as  compact  tongues  push  up  warm  exposed 
slopes  and  ridges,  and  even  form  forests  of  considerable  size  a  little 
below  timberline,  as  on  Whirlwind  Peak,  in  the  Absaroka  Range. 
The  Rocky  Mountain  white  pine,  although  a  tree  usually  found  only 
at  a  lower  altitude,  was  found  near  timberline  on  the  Bighorn  Moun- 
tains, and  also  on  the  exposed  10,000-foot  summit  of  Laramie  Peak, 
where  its  characteristic  growth  is  in  depressed  mats  at  the  base  of  a 
low  central  bole  6  feet  or  more  in  height.  Prostrate  clumps  of  low 
juniper  are  common  in  slide  rock  at  timberline  on  the  Sierra  Mad  re, 
on  the  Bighorn  and  Wind  River  Ranges,  and  elsewhere. 

Although  prominently  characterized  by  depauperate  vegetation, 
the  grassy  slopes  between  the  scattered  clumps  of  conifers  are  hand- 
somely carpeted  with  a  wealth  of  small  flowering  plants.  Conspicu- 
ous flowers  of  spring  and  early  summer,  as  observed  on  the  Wind 
River  and  Bighorn  Ranges,  include  globe  flower,  mountain  cowslip, 
shooting  star,  columbine,  spring  beauty,  and  various  mertensias, 
Jacob's  ladder,  forget-me-nots,  buttercups,  saxifrages,  and  drabas. 
In  early  autumn  the  timberline  region  on  the  Wyoming  and  Ab- 
saroka Ranges  was  brilliant  with  flowering  mats  of  lupine,  larkspur, 
painted  cup,  mountain  heath,  and  mountain  laurel,  with  the  more 
scattering  Parry  primroses,  harebells,  gentians,  phloxes,  and  ryd- 
bergias. 

Many  of  the  plants  of  the  Hudsonian  Zone  and  most  of  the  birds 
and  mammals  occur  also  in  the  adjoining  Canadian  or  Arctic- Alpine 
Zones.  Comparatively  few  species  are  closely  restricted  to  this 
narrow  area. 

Mammals — Hudsonian  Zone. 

Mammals  having  their  center  of  abundance  at  or  near  timberline 
are  the  mountain  sheep  (Oris  canadensis  canadensis) ,  the  timberline 
chipmunk  (Eutamias  oreocetes),  marmots  (Marmota  flaviventris 
nosophora  and  M.  f.  luteola),  and  the  pika.  or  coney  (Ochofona 
uinta).  A  number  of  species  range  into  this  region  from  the  Cana- 
dian Zone,  or  make  their  homes  in  both  areas.  Those  occurring  thus 
with  some  regularity  are : 


Sciurus  hudsonievs  ventorum,  Rocky 
Mountain  Red  Squirrel. 

Callospermophilns  lateralis:  eargi.  Wind 
River  Mantled  Ground  Squirrel. 

Evotoings  gapperi  galei,  Gale  Red- 
backed  Mouse. 

Microtus  mordax  mordax,  Rocky  Moun- 
tain Meadow  Mouse. 

Microtus  nanus  nanus.  Dwarf  Field 
Mouse. 

Thomomys  uinta,  Uinta  Pocket  Gopher. 


Thomomys  talpoidcs  caryi.  Bighorn 
Pocket  Gopher. 

Lepus  bairdii  bairdii,  Snowshoe  Rab- 
bit. 

Canis  testes,  Mountain  Coyote. 

Vulpes  macrourus,  Mountain  Red  Fox. 

Mattes  catirina  origenes,  Rocky  Moun- 
tain Marten. 

Boreas  obscurus  obscurus,  Rocky  Moun- 
tain Shrew. 

Sorex  personatus  personatus.  Masked 
Shrew. 


North  American  Fauna  No.  42,  U.  S.  Dept  Agr.     Biological  Suivey, 


Plate  X. 


Fig.  1.— Engelmann  Spruces  at  Timberline,  West  Slope  of 
Whirlwind  Peak,  Absaroka  Range  (10,000  Feet). 


Fig.  2.— White-Barked  Pines,  Same  Locality. 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  XI. 


Fig.  1.— Lower  Edge  of  Hudsonian  Zone,  Wind  River  Range  South  of  Fremont 

Peak  (10,500  Feet). 


Fig.  2.— East  Slope  of  Needle  Mountain,  Absaroka  Range. 

Growth  of  Engelinann  spruce,  alpine  fir,  and  grav-leaved  willow  (Salix  glaucops)  &t  timberline 

(10,000  feet). 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  XII, 


Fig.  1.— Arctic-Alpine  Zone,  Wind  River  Range. 
South  of  Fremont  Peak,  from  11,000  feet  elevation,  July  17, 1911. 


Fig.  2.— Arctic-Alpine  Zone,  Absaroka  Range. 

Between  the  Greybull  and  the  South  Fork  of  Shoshone  River,  from  east  slope  of  Needle  Mountain 

(10,500  feet),  July  11,  1910. 


1917.] 


HUDSON' I  AX    /ONE. 


41) 


BBEEDING    BlKDS — HUDSONIAN    ZONE. 

[Species  marked  C.  breed  also  iu  the  Canadian  Zone.] 


Picoides  americawus  dorsaliSj  Alpine 
Three-toed  AYoodpecker.    C. 

Perisoreus  canadensis  capitalis,  Rocky 
Mountain  Jay.     C. 

Nucifraga  colutnbiana,  Clark  Nut- 
cracker.1 

Pinicola  enucleator  montana,  Rocky 
Mountain  Pine  Grosbeak. 

Zonotrichia  leucophrys,  White-crowned 
Sparrow.     C. 


Junco  hyemalis  meamsi,  Pink-sided 
Junco.     C. 

Junco  phwonotus  caniceps,  Gray-headed 
Junco.     C. 

Wilsonia  pusilla  pileolata,  Pileolated 
Warbler.     C. 

Certhia.  familiaris  molilalia.  Rocky 
Mountain   Creeper.     C. 

Regulus  satrapa,  Golden-crowned  King- 
let.   C. 


Plants — Hudsonian  Zone. 

[Species    marked    ('.    occur   also    in    the    Canadian   Zone;    those    marked    A.,   also    in    the 

Arctic-Alpine.] 


Vuius  albicaulis,  White-barked  Pine. 

ricea  engelmanni,  Engelmann  Spruce 
(dwarf).     C. 

Abies  lasiocarpa,  Alpine  Fir  (dwarf). 
C. 

Juniperus  sibirica,  Low  Juniper.    G. 

Salix  glaueops,  Gray-leaved  Willow. 

Salix  chlorophylla,  Willow.     C. 

Salix  sarimontana,  Willow.     A. 

Ribes  montii/cnum,  Bristly  Red  Cur- 
rant. 

Rubus  strigosus,  Red  Raspberry.     C. 

Polygonum  bistortoides,  Twisted  Po- 
lygonum.    C. 

Claytonia  rosea,  Spring  Beauty. 

Calandrinia  pygmcea,  Alpine  Bitter 
Root. 

Cerastium  beeringianum,  Mouse-ear 
duckweed. 

Caltha  leptoscpala,  Mountain  Cow- 
slip. 

Trollius  albiflorus,  Globe  Flower. 

Aquilegia  saximontana,  Columbine. 

Delphinium  subalpinum,  Larkspur. 

A  lie  hi  one  tetouensis,  Anemone. 

A  n  cm  one  globosa,  Anemone.     G. 

Ranunculus  alpeophilus,  Buttercup. 

Ranunculus  calthcefolius,  Buttercup. 

Thlaspi  glaucum,  Penny  Grass. 

Draba   luteola,   Whitlow  Grass. 

Clementsia  rhodantha,  Red  Orpine.     .4. 

Leptasea  hirculus,  Saxifrage. 

Potentilla  glaucophylla,  Cinquefoil. 
C. 


Drymocallis  pseudorupestris,  Avens. 
Lupinus  ccespitosus,  Lupine.     G. 
Lupinus   laxiflorus,   Lupine.     C. 
Trifolium  dasyphyllum,  Dwarf  Clover. 

A. 
Trifolium  parryi,  Parry  Dwarf  Clover. 

A. 
Hedysa  ru  m  s  ulphun  set  m  s. 
Angelica  roseana. 
Phyllodoce     empetriformis,     Mountain 

Heath. 

Kahnia   polifolia,  Mountain   Laurel. 
Primula  parryi,  Parry  Primrose. 
.  [iid  rosace  su  b  u  m  bell  a  ta. 
Dodecatheon  radicatum,  Shooting  Star. 
C. 

Gcntiana  calycosa,  Gentian. 
fit ut in  mi   st  rift  i  flora.   Gentian. 
Sircrtia  congesta,  A. 
Swertia  palustris,  G. 
Phlox  qwstritosa,  Phlox.     ('. 
Polemonium  viscosum,  Jacob's  Ladder. 
Polemonium  mellitum,  Jacob's  Ladder. 
Phacelia   sericea,    Silky   Phacelia.    A. 
Myosotis  alpestris,  Forget-me-not.    A. 
Uertensia  tweedyi,  Lungwort.     G. 
Pentstemon  alpinus,  Beard-tongue. 
Veronica  toormskjoldi,   Alpine    Speed- 
well. 

Veronica    serpyllifolia,    Speedwell.     C. 
Castilleja  spp.,  Painted  Cup. 
Campanula  parryi,  Parry   Harebell. 
Rydbergia  grandiftora,  Rydbergia.     A. 


Young  noted  in  summer. 


74410° 


50 


NORTH    AMERICAN    FAUNA. 


[  No.  42. 


Plants — Hudson i an  Zonk — Continued. 


Townsendia  parryi,  Parry  Townsendia. 

A. 
Senecio  frembnti,  Paintbrush.    A. 
Senedo  crassulus,  Paintbrush. 
Antennaria  rcfU.ra.  Everlasting. 
Erigeron  compositus,  Fleabane.     C. 
Erigeron  salsuginosus,  Fleabane. 


Chwnactis  alpina,  Alpine  Chsenactis. 

Trisetum  subspicatum,  Oat  Grass. 

Poa  epilis,  Bluegrass.    A. 

Phleum  alpinum,  Alpine  Timothy.     C 

Carex  nigricans,  Sedge. 

Carex  nova,  Sedge. 

Juncus  subtriflorus,  Rush. 


ARCTIC-ALPINE   ZONE. 

The  area  above  timberline  on  the  highest  mountains,  the  Arctic- 
Alpine  Zone,  corresponds  in  climate,  and  in  plant  species  especially, 
to  the  barren  grounds  of  the  Arctic.  It  is  not  continuous  from  one 
range  to  another  in  Wyoming,  for  even  in  the  mountainous  northwest 
the  areas  are  separated,  often  widely,  by  Canadian  Zone  valleys  and 
forested  divides  of  medium  elevation. 

A  wide,  almost  unbroken  stretch  of  this  zone  caps  the  massive 
Wind  River  Range  for  its  entire  length  (PL  XII,  fig.  1),  and  there 
is  a  broken,  irregular  area  of  equal  extent  on  the  main  crest  and 
primary  spurs  of  the  rugged  Absaroka  Range  (PL  XII,  fig.  2). 
The  Gros  Ventre  Range  is  capped  by  a  succession  of  Alpine  plateaus 
sloping  moderately  toward  the  north,  while  the  lofty  peaks  and 
jagged  crests  of  the  Tetons  form  a  narrow  strip  of  Alpine  country. 
Broad  Alpine  areas  on  the  Bighorn  Mountains  lie  south  of  the  gap 
at  the  head  of  Tongue  River,  and  include  all  elevated  summits  of 
the  Cloud  Peak  group.  There  are  traces  of  the  zone,  too  small  for 
plotting  on  the  map  (see  frontispiece),  on  a  few  mountains  in  the 
eastern  and  northwestern  borders  of  Yellowstone  Park.  In  southern 
Wyoming  the  Arctic-Alpine  Zone  is  restricted  to  the  lofty  plateau 
on  the  Snowy  Mountains,  at  the  northern  end  of  the  Medicine  Bow 
Range. 

This  high-altitude  area  is  a  bleak,  wind-swept  region  of  excessive 
snowfall  in  winter  and  frequent  squalls  of  rain,  sleet,  or  snow  in  the 
short  summer,  and  arctic  temperatures  prevail  throughout  the  year. 
On  all  the  ranges  snow  fills  the  gulches  and  partly  covers  cold  slopes 
and  declivities  even  in  the  warmest  months,  while  such  massive, 
elevated  ranges  as  the  Wind  Rivers.  Absarokas,  and  Tetons  carry  ex- 
tensive snow  fields,  and  even  a  few  perennial  ice  fields  or  glaciers  in 
protected  Alpine  valleys. 

The  Arctic- Alpine  Zone  is  conspicuously  marked  by  the  absence  of 
tree  growth,  which  ceases  at  its  lower  border.  The  altitude  of 
timberline  varies  with  latitude  and  slope  exposure  from  10.500  or 
11,000  feet  in  the  Sierra  Mad  re  and  the  Medicine  Bow  Range  at  the 
south,  to  9,500  or  10,000  feet  in  the  Bighorn  Mountains  and  in  the 
Yellowstone  Park  region. 


1917.]  ARCTIC-ALPINE   ZONE.  51 

Characteristic  Species — Arctic-Alpine   Zone. 

The  season  of  plant  growth  is  from  the  middle  of  May  until  August. 
but  during  this  brief  period  the  bleak  slopes  and  even  the  fields 
of  slide  rock  for  1,500  or  2,000  feet  above  timberline  are  bedecked 
with  a  profusion  of  bright-hued  Alpine  flowers.  A  luxuriant  growth 
of  Alpine  grasses  and  sedges  obtains  wherever  there  is  any  depth  of 
soil,  and  furnishes  rich  pasturage  for  mountain  sheep  and  a  few  other 
mammals  which  spend  the  summer  in  this  usually  forbidding  region. 
In  Wyoming  plant  growth  rapidly  decreases  in  size  above  12,000  feet. 

The  low  shrubby  or  matted  growth  usually  extending  from  500  to 
1,000  feet  above  the  limit  of  trees  consists  chiefly  of  dense  thickets  of 
willow  (Salix  glaucops)  and  copses  of  shrubby  cinquefoil  (Pasi- 
phora  frutiaosa)  which  push  up  the  bottoms  and  along  the  margins 
of  wet  gulches  and  basins;  mats  of  mountain  heath  (Phyllodoce  em- 
petriformis)  and  alpine  avens  on  rocky  slopes  and  ridges;  and  spiny 
red  currant  {Ribes  montigenum)  and  dwarfed  raspberry  (Rubus 
strigosns)  in  slide  rock.  Dwarf  alpine  willows  mat  the  ground  in 
places  for  an  indefinite  distance  above  timberline,  and  may  even 
reach  the  highest  summits  with  the  mosses  and  lichens. 

A  large  number  of  characteristic  Arctic- Alpine  herbaceous  plants 
mark  the  zone  in  Wyoming,  but  only  four  species  of  breeding  birds 
are  peculiar  to  it,  and  no  mammals. 

Mammals — Arctic-Alpine  Zone. 

The  few  mammals  found  in  the  Arctic- Alpine  Zone  in  Wyoming 
belong  to  lower  zones.  Among  those  attracted  in  summer  to  its 
grassy  slopes  and  crests  are  the  elk  {Cervu-s  canadensis  canadensis), 
mountain  sheep  (Ovis  canadensis  canadensis),  timberline  chipmunk 
(Eutamias  oreoeete*),  and  Wind  River  mantled  ground  squirrel  (Cal- 
lospermophilus  1.  caryi).  Marmots  (Marmota  f.  noso/>hora),  pocket 
gophers  (Thonwniys  uinta  and  T.  f.  fuscus),  meadow  mice  (Microtus 
m.  mordax),  coneys  {Ochotona  uinta).  and  rarely  the  Uinta  spermo- 
phile  (('/fellas  armatus),  apparently  are  resident  in  the  lower  part  of 
the  zone  in  different  localities.  The  coyote  (Canis  It  stes?)  and  moun- 
tain red  fox  (Vulpes  maerourus)  range  at  various  times  into  Alpine 
country  in  search  of  prey.  On  the  Wyoming  mountains  mammals  are 
rarely  met  with  above  12.000  feet. 

Breeding  Birds— Arctic-Alpine  Zone. 


Lciteosticte  atrata,'  Black  Rosy  Finch. 
Anthus  rubesccns?  Pipit,  Titlark. 
Otocoris   alpestris   leucolwma*   Deserl 
Horned  Lark. 


Lagopus  lencurus  leucurus,1  White- 
tailed  Ptarmigan. 

Leucosticte  australis?  Brown-capped 
Rosy   Finch. 

1  On  the  Medicine  Bow  Range. 

2  On  the  Teton,  Wind  River,  and  Absaroka  Ranges. 

3  Throughout   the  mountains. 

4  One  of  the  few  nesting  birds  of  the  Arctic-Alpine,  but  belonging  chiefly  to  the  Transi 
tion  and  Upper  Sonoran  Zoms. 


52 


NORTE    AMERICAN    FAUNA. 


[No.  12 


Plants — Abctic- Alpine  Zone. 

[Species  marked  //.  occur  also  in  the  Hudsonian  Zone.] 
Aragallus  nanus,  Loco. 


Salix  petrophila,  Rock  Willow. 

Salix  latent.  Rock  Willow. 

Salix  nivalis,  Alpine  Willow. 

Salix  saximontdna,  Creeping  Wil- 
low.   //. 

Oxyria  digyna,  Mountain  Sorrel.    IT. 

Polygonum  viviparum,  Alpine  Knot- 
weed. 

Claytonia  megarrhiza,  Arctic  Spring 
Beauty. 

Spraguea  multiceps.  / 

Silt  lie  acaulis,  Stemless  Catchfly.     II. 

Alsinopsis   obtusiloba,    Sandwort. 

Paronychia  pulvinata,  Whitlowwort. 

Ranunculus  adoneus,  Buttercup.    II. 

Thalictrum  alpinum,  Alpine  Meadow 
Rue. 

\ni his  lyalli. 

Smelowskia  americana. 

Draba  caria,  Whitlow  Cress. 

Draba  crassifolia,  Whitlow  Cress. 

Draba  densifolia;  Whitlow  Cress. 

Parrya  nudicaulis,  Purple  Parrya. 

Rhodiola  integrifolia,  Rosewort. 

Saxifraga  cernua,  Arctic  Saxifrage. 

Leptasea  flagellaris,  Saxifrage. 

Boykinia  heuclieriformis,  Saxifrage. 
Boykinia.     //. 

Dryas  octopetala,  Alpine  Avens. 

Sibbaldia   procumbens,   Sibbaldia.     H. 

Sievcrsia  t  u  r  b  i  n  a  t  a  ,  Mountain 
Avens.     //. 

Lupinus  m  o  n  ticola,  Mountain  Lu- 
pine.    IF. 


Aragallus  lagopus,   Loco. 

Bupleurum  americanum,  Thorough- 
wax. 

(Jreoxis  alpina. 

Androsace  carinata. 

Gentiana  romanzovi,  Dwarf  Closed 
Gentian. 

Polemonium  confertum,  Jacob's  Lad- 
der.    H. 

Eritrichium  argenteum,  Alpine  Forget- 
me-not. 

Mertensia  brevistyla  and  others. 

Pellicula  ris  parryi,  Parry  Louse- 
wort.     H. 

Besseya  alpina. 

Campanula  tttti flora,  Arctic  Harebell. 

Solidago  decumbens,  Goldenrod. 

Erigcron  pinnatisectus,  Fleabane.     //. 

Erit/cron  radicatus,  Fleabane.     11. 

Toncstus  pygmcBUS. 

Achillea  alpicola,  Alpine  Yarrow. 

Artemisia  scopulorum,  Alpine  Sage- 
brush. 

Festuca  brachyphylla,  Alpine  Fescue. 

Poa  arctica,  Arctic  Bluegrass. 

Poa  lettermanni,  Bluegrass. 

Poa  alpina,  Alpine  Bluegrass.     II. 

Carex  engelmanni,  Sedge. 

Carer  albo-nigra,  Sedge. 

Caret-  nubicola,  Sedge. 

Carer  at  rata.  Black  Sedge. 

Carex  ph&ocephala,  Sedge. 

Juncoides  spicatum,  Wood  Rush.    II. 


IMPORTANCE   OF   BOREAL  ZONES   TO   WYOMING   AND   ADJOINING 

AREAS. 

Climatically  and  physically  unsuited  to  agriculture,  the  high  alti- 
tude Canadian,1  Hudsonian,  and  Arctic- Alpine  Zones  nevertheless  are 
not  only  a  valuable  but  an  essential  complement  to  the  lower  agricul- 
tural areas  of  Wyoming  and  most  adjoining  States.  As  the  chief 
sources  of  three  great  river  systems — the  Columbia,  the  Missouri,  and 
the  Colorado — their  importance  is  far  from  local.  The  great  value  of 
the  Boreal  zones  lies  in  their  peculiar  adaptation  to  moisture  con- 
servation. This  is  accomplished  climatically  on  the  bleak  and  bar- 
ren Alpine  slopes  and  summits,  and  by  plant  and  forest  cover  and 
climate  combined  in  the  Hudsonian  and  Canadian  Zones. 


1  See  p.  42. 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  XIII. 


Fig.  1.— Teton  Range,  Mount  Moran  South  to  Grand  Teton. 
Photograph  taken  from  foot  of  Jackson  Lake,  June  5, 1911,  by  Edward  A.  Preble. 


Fig.  2.— Snow  in  Lower  Part  of  Hudsonian  Zone  (10,800  Feet). 
East  slope  of  Bridger  Peak,  Sierra  Madre,July  7, 1911. 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  XIV. 


Fig.  1.— Heavy  Forest  of  Engelmann  Spruce,  Canadian  Zone,  Wyoming  Range 
West  of  Merna  (9,000  Feet). 


Fig.  2.— Rank  Vegetation  on  Floor  of  Canadian  Zone  Forest,  Grinnell  Creek, 
Absaroka  Range  (7,500  Feet). 


1917.]  BOREAL    ZONES.  53 

A  heavy  mantle  of  snow  covers  the-entire  Arctic-Alpine  cap  of  the 
main  ranges  for  seven  or  eight  months  of  the  year  (  PI.  XIII,  fig.  i), 

while  huge  drifts  and  snow  fields  remain  in  protected  spots  through- 
out the  summer.  In  the  timberline  region  shaded  gulches  are  banked 
with  snow  until  August,  and  it  is  not  uncommon  to  find  snowdrifts 
in  the  forest  depths  of  the  Canadian  Zone  until  midsummer  (PI. 
XTII5  fig.  2),  although  most  of  the  snow  on  the  lower  slopes  goes  off 
in  flood  in  May  and  early  June.  The  summer  precipitation  is  heavy, 
consisting  of  rain  in  the  forest  belt,  and  frequent  showers  and  squalls 
of  rain,  sleet,  and  snow  on  the  peaks  and  higher  slopes. 

The  rotting  vegetation  and  mellow  soil  of  the  cool  mountain 
forests  are  specially  adapted  to  the  retention  of  this  moisture,  much 
of  which  works  down  from  the  slowly  melting  snow  banks  in  the 
Alpine  area.  The  shaded,  mossy  forest  floor  soon  becomes  saturated, 
and  the  water,  percolating  through  the  leaves  and  loose  soil,  finds  its 
way  gradually  through  rock  crevices  into  ravines  and  depressions, 
finally  flowing  clear  and  sparkling  into  the  streams  lower  down.  Thus 
is  insured  an  abundance  of  pure  mountain  water  to  the  arid  but  fertile 
valleys  and  plains,  and,  what  is  of  greatest  importance,  a  fairly  uni- 
form volume  in  the  streams  toward  the  end  of  the  growing  season,  the 
period  when  most  required  by  crops.  The  conversion  through  irriga- 
tion of  portions  of  the  valleys  of  the  Snake.  Yellowstone,  Bighorn, 
Green,  and  North  Platte  Rivers  in  Wyoming,  and  especially  in 
neighboring  States,  as  well  as  of  vast  tracts  as  yet  undeveloped,  into 
rich  agricultural  districts  is  made  possible  through  the  combined 
agency  of  the  climate  and  the  forest  and  plant  cover  of  the  Boreal 
zones  of  Wyoming. 

The  Canadian  Zone  has  large  tracts  of  forest,  mainly  of  lodgepole 
pine  and  Engelmann  spruce  (PL  XIV,  fig.  1),  with  considerable 
Douglas  spruce  on  the  lower  slopes.  While  these  are  useful  for  lum- 
ber and  other  utilities,  the  intrinsic  value  is  small  in  comparison  with 
the  permanent  service  they  are  naturally  fitted  to  perform  in  connec- 
tion with  the  agricultural  utilization  of  the  arid  regions.  Fortun- 
ately, most  of  the  timberlands  of  the  Canadian  Zone  in  Wyoming  are 
already  included  in  national  forests.  Forest  control  with  a  view  to 
their  conservation  is  therefore  most  timely. 

While  the  higher  mountain  slopes  of  Wyoming  afford  a  rich  pas- 
turage during  the  summer  months  for  many  hundreds  of  thousands 
of  sheep,  careful  regulation  of  sheep  grazing  is  of  the  utmost  impor- 
tance, as  the  natural  plant  cover  is  a  vital  factor  in  catching  and 
holding  moisture.  (PL  XIV,  fig.  2.)  Once  this  is  badly  broken  up 
or  removed  by  overgrazing  the  rains  go  off  with  a  rush,  carrying 
much  of  the  soil  with  them.  This  results  in  dry,  barren  slopes,  dirty 
streams,  and  a  greatly  diminished  flow  of  water  in  the  lower  country 
during  much  of  the  growing  season,  unless  there  are  adequate  water- 


54  NORTH    AMERICAN    FAUNA.  [No.  42. 

storage  facilities.  Mountains  which  hare  been  extensively  grazed  in- 
clude the  Wyoming  and  Salt  River  Ranges,  the  southern  end  of  the 
Wind  River  Range  below  the  Big  Sandy.  Sierra  Madre,  and  the 
mountains  on  either  side  of  the  Bighorn  Basin  at  its  southern  end. 
The  mountain  meadows  and  parks  with  their  luxuriant  grasses  con- 
stitute an  ideal  summer  range  for  cattle,  and  are  extensively  utilized 
for  this  purpose. 

Wyoming  offers  many  attractive  regions  to  the  tourist,  the  sports- 
man, and  to  those  in  search  of  health  or  recreation.  As  a  permanent 
pleasure  ground  the  mountainous  region  at  the  northwest  is  a  val- 
uable asset,  and  is  perhaps  unsurpassed  in  extent  and  rugged 
grandeur.  Dashing  trout-filled  streams  add  to  the  attractiveness 
of  a  section  full  of  wild  charm  and  beauty,  while  the  dense  forests 
of  Yellowstone  Park  and  the  northern  end  of  Jackson  Hole  afford  a 
safe  retreat  and  breeding  range,  under  Federal  and  State  protection, 
for  many  thousands  of  elk  and  other  large  game  animals,  and  insure 
the  best  of  hunting  in  season  in  districts  adjacent  to  these  protected 
areas. 


NOTES  ON  THE  DISTRIBUTION  OF  CONSPICUOUS  TREES  AND 
SHRUBS  OF  WYOMING. 

The  following  annotated  list  of  Wyoming  trees  and  shrubs,  while 
very  incomplete,  includes  principally  the  more  conspicuous  and  char- 
acteristic zone  species,  and  should  add  to  the  knowledge  of  their  dis- 
tribution within  the  State.  It  is  based  chiefly  on  notes  and  specimens 
collected  by  Biological  Survey  field  parties. 

Pinus  albicaulis  Eugelmann.    White-Barked  Pine.  (Fig.  3.) 

The  small  white-barked  pine  in  Wyoming  is  peculiar  to  the  high  altitudes  of 
the  northwest,  where  it  is  a  characteristic  tree  of  the  Hudsonian  Zone  just 
below  timberline.  It  oc- 
curs on  all  the  lofty 
mountains,  finding  its 
southern  limits  in  the 
main  chain  of  the  Rock- 
ies on  the  Wind  River 
and  Salt  River  Ranges. 
At  timberline  it  is  often 
the  most  abundant  tree, 
especially  on  parts  of  the 
Absaroka  Range,  but  usu- 
ally shares  this  bleak 
region  with  dwarfed  al- 
pine firs  and  Engelmann 
spruces,  pushing  up  dry 
slopes  and  crests  of  ex- 
posed ridges  in  ascend- 
ing tongues,  while  the 
spruces  and  firs  occupy 
wet  gulches  and  the 
deeper-soiled  slopes. 

Pinus  flexilis  James. 

Rocky  Mountain  White 

Pine. 

The  Rocky  Mountain 
white  pine  has  a  general 
dispersion  in  upper 
Transition  and  Canadian 
Zones  in  all  except  the 
northeast  corner  of  the 
State.  It  has  its  center 
of  abundance  with  the  Douglas  spruce  along  the  lower  edge  of  the  Cana- 
dian forest  belt,  occupying  the  ridges  and  dry  slopes,  while  the  spruces  are 
in  gulches  and  on  steep,  cold  exposures.  There  are  few  gravel  or  rocky  ridges 
on  the  high  central  and  western  plains  and  deserts  that  do  not  have  more  or 
less  of  this  pine  in  scattered  and  usually  somewhat  ragged  growth.  In 
Wyoming  it  is  a  small  tree,  rarely  attaining  a  height  of  more  than  30  or  40 
feet  or  a  diameter  above  1*  feet  under  most  favorable  conditions.     Its  usual 

55 


Fig.  3. — Forest  of  white-barked  pine  (Pinus  albicaulis) 
just  below  timberline  on  west  slope  of  Whirlwind  Peak, 
Absaroka  Range  (9,800  feet). 


56 


NORTH    AMERICAN    FAUNA. 


[No.  42. 


habit  of  growth  is  scattering  and  patchy,  lmt  hi  deep-soiled  mountain  districts 
it  often  produces  groves  <>('  considerable  extent. 

Pinus  scopulorum  (Engelmann)   Lemmon.    Rocky  Mountain  Yellow  Pine. 

The  distribution  in  Wyoming  of  the  Rocky  Mountain  yellow  pine,  a  well- 
known  Transition  Zone  tree,  is  mainly  west  to  the  eastern  slopes  of  the 
Bighorn  Mountains  and "  the  region  of  the  upper  Platte,  as  follows :  Black 
Hills,  he-ivy  open  forest  to  6,000  or  6,500  feet  altitude;  Bear  Lodge  Moun- 
tains, moderate  growth  with  oaks,  3,500  to  6.000  feet;  Colony,  low  ridges; 
throughout  the  borders  of  Cheyenne  River  drainage,  in  scattering  pockets  and 
fringe;  watersheds  between  Belle  Fourche  and  Tongue  Rivers,  in  thin  forest 

over  roughest  sections ; 
lower  eastern  slopes  of 
Bighorn  Mountains,  tol- 
erably wide  belt  up  to 
6,000  or  7,000  feet;  foot- 
hill region  of  Casper 
Mountains  south  through- 
out the  length  of  the 
Laramies,  including  the 
Hartville  group  east  of 
the  Platte,  in  usually 
good  growth ;  divide 
southeast  of  Efell ;  Pine 
Mountain  south  of  Na- 
trona ;  borders  of  North 
Platte  Valley,  between 
Alcova  and  Leo.  and 
along  north  base  of  Shir- 
ley Mountains,  scatter- 
ing trees ;  Seminole  Can- 
yon on  the  Platte,  heavier 
growth  ;  Roc  k  Rive  r, 
ridges ;  Woods,  Medicine 
Bow  Range,  some  at 
lower  edge  of  coniferous 
forest ;  basal  slopes  of 
Sierra  Madre  south  of 
Downington ;  and  can- 
yons near  the  mouth 
of  Grand  Encampment 
River.  This  pine  was  not  found  in  western  Wyoming,  although  doubtless  there 
are  scattering  trees  on  the  basal  slopes  of  some  ranges. 

The  yellow  pine  yields  valuable  lumber  mainly   in   the  Black   Hills,   Bear 
Lodge  and  Bighorn  Mountains,  and  on  the  Laramies  in   the  region   north  of 
Laramie  Peak.     Elsewhere  its  growth  is  generally  more  or  less  scrubby. 
Pinus  murrayana  Balfour.     Lodgepole  Pine. 

The  coniferous  element  of  the  Canadian  forest  belt  in  Wyoming  is  chiefly 
of  lodgepole  pine  and  Engelmann  spruce.  The  pines,  although  occurring 
throughout  the  full  width  of  the  Canadian  Zone,  reach  their  greatest  abundance 
and  heaviest  and  purest  stand  in  its  lower  half.  The  best  forests  are  on  the 
main  ranges,  varying  in  elevation  from  8.500  to  10,000  feet  on  the  Sierra 
Madre  at  the  south;  8,000  to  9,500  feet  on  the  Wind  River  Range;  and  7,000 
to  9,000  feet  in  Yellowstone  Park,  the  northern  Absarokas,  and  on  the  eastern 
slopes  of  the  Bighorns. 


BIIG60M 

Fia.  4. —  Forest  of  Engelmann  spruce   (Picea  engelmanni) 

on  north  slope  of  Ferris  Mountains   (9,000  feet). 


1917.] 


TREES   AND   SHRUBS. 


57 


The  lodgepole  pine  forests  of  Wyoming  are  of  gveal  value,  bul  have  been 
extensively  levied  upon  for  ranch  fences,  railroad  ties,  mine  lagging,  and 
lumber,  and  extensive  tracts  have  been  destroyed  by  fire.  The  deforested 
areas  are  now  largely  grown  up  with  second-growth  pine  and  aspen. 

Pinus  edulis  Engelmann.     Pinyon ;  Nut  Pine. 

The  pinyon,  a  small  representative  of  the  Upper  Sonoran  Zone,  barely 
enters  Wyoming  in  the  lower  valley  of  Green  River.  Scattering  trees  were 
found  at  7,000  feet  elevation  on  the  north  face  of  a  juniper  ridge  3  miles 
north  of  the  Utah  boundary  and  the  same  distance  east  of  Green  River.  The 
pinyon   may   possibly   occur   elsewhere   in   this    rough   juniper-clad    ridge   and 


Fig.  5.— Frini 


of  blue  spruce    (Picea  parryana)   on   the  Big  Sandy,   southwest  base  of 
Wind  River  Range    (7,500   feet). 


mesa  country,  but  it  was  not  detected  from  Green  River  east  to  Red  Creek, 
and  thence  north  to  Rock  Springs. 

Picea  engelmanni  (Parry)  Engelmann.     Engelmann  Spruce.  (Fig.  4.) 

The  Engelmann  spruce  is  the  principal  conifer  in  the  upper  part  of  the 
Boreal  forest  belt  on  the  high  ranges  of  northwest  Wyoming,  and  also  on  the 
Bighorn,  Sierra  Madre,  and  Medicine  Bow  Ranges.  Next  to  the  lodgepole 
pine  it  is  the  most  abundant  forest  tree  of  Yellowstone  Park.  Although  of 
regular  occurrence  at  timberline  in  a  dwarfed  state,  this  spruce  belongs  to 
the  Canadian  Zone,  attaining  its  maximum  growth  on  cold,  damp  slopes  and 
in  bogs  between  8,000  and  9,000  feet.  It  does  not  attain  its  best  development 
in  dry  situations  or  on  warm  slopes,  where  it  is  found  at  a  somewhat  higher 
level. 

Picea  parryana   (Andree)    Sargent.     Blue  Spruce.  (Fig.  5.) 

The  blue  spruce  occurs  chiefly  in  western  Wyoming  north  to  Jackson  Lake 

and  the  head  of  Wind  River,  but  was  also  noted  at  Woods  and  in  gulches  south 


58 


NORTH    AMERICAN    FAUNA. 


[No.  42. 


of  Sherman,  and  should  be  present  elsewhere  in  the  southern  mountains.  It 
inhabits  the  margins  of  cold  streams  in  the  lower  border  of  the  Canadian  forest 
belt  between  7.000  and  8.000  feet  elevation  (6.500  to  7.500  feet  in  Jackson  Hole), 
forming  usually  a  most  attractive  fringe  of  scattering  symmetrical  trees. 

Abies  lasiocarpa  (Hooker)  Nuttall.     Alpine  Fir. 

The  alpine  fir  has  much  the  same  distribution  and  vertical  range  as  the  Engel- 
mann  spruce,  although  generally  less  abundant  and  growing  in  scattered  thicket 
formation,  rarely  forming  a  heavy  forest.  With  the  spruce  it  extends  regularly 
to  timberline  on  the  Wyoming  ranges,  where  it  is  the  more  common  of  the  two 
and  forms  dense  prostrate  mats  on  the  bleak,  wind-swept  slopes.     This  fir  is 

partial  to  cold  gulches, 
stream  banks,  and  damp 
spots  generally.  It  may 
be  seen  to  best  advantage 
in  typical  growth  as  dark 
scattered  clumps  in  aspen 
woods.  While  usually  a 
small  tree,  it  attains  large 
size  in  gulches  and  on 
streams  in  the  higher 
portions  of  Yellowstone 
Park,  where  it  forms  for- 
ests of  considerable  ex- 
tent above  8,000  feet 
altitude.  It  was  observed 
on  all  the  mountains  ex- 
cept the  Bear  Lodge  and 
Black  Hills  groups  at  the 
northeast. 

Abies  (concclor?)  Lind- 
ley.  White  Fir. 
The  white  fir  (appar- 
ently Abies  ooncolor) 
forms  small  forests  be- 
low 9,000  feet  elevation 
on  the  northern  shoul- 
ders of  the  Uinta  Moun- 
tains west  of  Lonetree, 
and  is  present  also  in 
fairly  good  stand  with 
aspens  on  the  summit  of  the  plateau  west  of  Maxon.  Firs  which  were  com- 
mon at  8,000  feet  on  the  western  slopes  of  the  Salt  River  Range  near 
Afton  also  had  smooth,  dark-gray  bark  and  were  unquestionably  distinct 
from  the  light-barked  A.  lasiocarpa  on  the  upper  slopes  of  the  same  mountains 
above  9,000  feet.  Unfortunately  no  specimens  were  preserved  from  the  above 
localities. 

Pseudotsuga  mucronata  (Rafinesque)  Sudworth.  Douglas  Spruce.  (Fig.  6.) 
The  Douglas  spruce  has  a  wide  distribution  in  upper  Transition  and  lower 
Canadian  Zones  from  the  Laramie  and  Bighorn  Mountains  westward.  It  was 
not  observed  on  the  Black  Hills  or  Bear  Lodge  Mountains.  This  spruce  grows 
chiefly  at  the  lower  edge  of  the  lodgepole  pine  belt  on  the  main  ranges  of 


Bl  1631 

pIG-   c». — Forest    of   Douglas   spruce    {Pseudotsuga   mucro 
naia),  north  slope  of  Ferris  Mountains  (8,500  feet). 


1017.1  TREKS   AND    SHRUBS.  59 

western  Wyoming,  teaching  down  t<>  open  sagebrush  slopes  on  their  basal 
Hanks,  and  on  (he  Bighorns  and  La  ramies,  extending  down  into  the  upper  pari 
of  the  yellow  pine  belt.  II  varies  locally  as  to  abundance  from  a  thin  fringe 
of  scattering  trees  and  thickets  on  cool  declivities  and  in  gulches  (which  is 
usual),  to  a  wide  belt  of  heavy  pure  forest  of  lumber  size. 

Juniperus  scopulorum  Sargent.     Rocky  Mountain  Juniper. 

The  most  conspicuous  and  widely  distributed  juniper  in  the  State  is  the 
Rocky  Mountain  juniper,  occurring  from  the  barren  rocky  ridges,  canyons, 
gulches,  and  badlands  bluffs  on  the  Sonoran  plains  and  deserts,  up  through 
the  Transition  Zone,  where  it  mingles  with  yellow  pines,  Douglas  spruces,  and 
Rocky  Mountain  white  pines  on  the  lower  mountain  slopes.  This  juniper  is  of 
scattering  growth  along  the  bases  of  mountains  and  on  the  margins  of  the 
desert  basins  and  valleys,  and  does  not  form  a  well-defined  belt  along  the 
upper  edge  of  the  Upper  Sonoran  Zone  as  it  often  does  in  the  southern  Rocky 
Mountains. 

Juniperus  knighti  A.  Nelson.     Desert  Juniper. 

The  distribution  of  the  desert  juniper  is  imperfectly  known,  owing  to  very 
scattering  held  work  in  the  difficult  Red  Desert  region  which  it  inhabits.  The 
few  localities  from  which  there  are  specimens  indicate  a  restricted  range, 
mainly  in  the  Upper  Sonoran  Zone.  S.  G.  Jewett  collected  the  species  at 
Mountainview,  on  May  27,  1913,  and  it  appears  to  be  the  dominant  juniper 
at  Rock  Springs,  near  Carter,  and  in  the  badlands  to  the  south  and  east  of 
Lyman.  <  >n  the  Green  River  bluffs  near  the  Utah  boundary  Juniperus  mono- 
sperma  is  not  uncommon,  and  on  the  higher  borders  of  the  Red  Desert 
J.  scopulorum  is  the  common  species.  The  desert  juniper  is  usually  of  shrubby 
stature,  branching  from  the  base,  and  scarcely  attains  the  dignity  of  a  tree. 
Nelson  found  the  species  on  the  sandstone  bluffs  of  the  Bitter  Creek  drainage, 
and  records  specimens  from  Point  of  Rocks  and  Rock  Springs.1 

Juniperus  monosperma  Engelmann.     One-Seeded  Juniper. 

The  one-seeded  juniper  is  found  at  the  northern  base  of  Owl  Creek  Moun- 
tains west  and  southwest  of  Thermopolis ;  at  Hailey.  southeast  of  Lander: 
along  the  Platte  near  Alcova  in  canyons  and  on  dry  slopes  up  to  0,800  feet 
elevation;  and  on  the  rough  breaks  along  the  lower  Green  River  Valley,  par- 
ticularly on  the  east  side  between  Sage  Creek  and  the  Utah  boundary.  It  is 
tolerably  common  on  the  Snake  River  bluffs  near  Baggs  and  is  probably  the 
species  which  forms  a  considerable  belt  along  the  western  bases  of  the  Sierra 
Madre.  It  was  noted  only  in  the  Upper  Sonoran  Zone,  extending  up  to  7,500 
feet  on  the  hot  slopes  east  of  Green  River  near  the  State  line. 

Juniperus  sibirica  Burgsdorff.     Low  Juniper. 

The  low  juniper,  a  graceful  evergreen,  is  a  characteristic  undershrub  in  the 
Boreal  forest  belt  throughout  the  Wyoming  mountains.  On  most  of  the  loftier 
ranges  it  extends  to  timberline,  where  it  forms  dense  prostrate  mats  among 
lo-ks.  Its  center  of  abundance  is  in  the  Engelmann  spruce  and  lodgepole  pine 
forests  of  the  Canadian  Zone. 

Juniperus  communis  Linnaeus.     Mountain  Juniper. 

Not  infrequent  in  mountain  forests  at  lower  elevations  than  the  low  juniper. 
The  shrubby  mountain  juniper  was  noted  as  follows:  Foothills  west  of  Wheat- 
land, 5,500  feet;  Springhill;  near  Sundance;  Wolf,  north  base  of  Bighorn 
Mountains.  0,700  feet:  head  of  Pat  O'Hara  Creek,  northwest  of  Cody. 

1  Bull.  13,  Div.  of  Agrost.,  T\  S.  Dept.  Agr.,  p.  54,  1898. 


00 


NORTH    AMERICAN    FAUNA. 


[No.  42. 


Juniperus  sabina   Liiuueus.     Creeping  Juniper;  Trailing  S:ivin. 

Mats  of  the  creeping  juniper,  or  trailing  savin,  are  conspicuous  <>n  dry  gravel 
ridges  and  exposed  points  and  summits  of  high  hills  along  the  northern  edge 
of  Wyoming  between  the  Bighorn  and  Bear  Lodge  Mountains.  The  species 
extends  west  at  least  to  the  eastern  base  of  the  Absaroka  Range,  but  is  of  more 
general  distribution  in  the  northeast.  It  was  not  found  in  other  sections  of 
the  State.  The  habitat  of  this  species  is  in  the  Transition  and  lower  Canadian 
Zones. 

Populus  tremuloides  Michaux.     Aspen  Poplar.  (Fig.  7.) 

The  Boreal  aspen  poplar  is  generally  dispersed  at  suitable  elevations,  but 

does  not,  as  a  rule,  reach  large  size  in  the  Wyoming  mountains.     It  commonly 

occurs  in  thickets  10  to 
20  feet  high  on  damp, 
cold  slopes,  as  under- 
growth in  the  coniferous 
forests,  or  as  first  growth 
on  burned-over  tracts. 
Low,  scraggy  thickets  of 
aspen  are  on  most  of  the 
cold  slopes  and  draws 
above  7,500  feet  altitude 
on  barren  elevations  of 
the  southwest,  as  the 
Aspen  Mountains,  Bear 
River  Divide,  and  the 
high  plateaus  east  of 
Green  River  near  the 
southern  boundary  of  the 
State.  Beautiful  aspen 
groves  were  noted,  how- 
ever, between  8,000  and 
9.000  feet  on  the  lower 
southern  slopes  of  the 
Wind  River  Range  east 
of  Leckie,  on  the  divide 
between  Hoback  and 
Green  Rivers,  and  on  the 
northern  shoulders  of  the 
Uinta  Mountains  south- 
east     of     Hilliard.     The 


Bl  1622 

Fig.    7. — Grove    of    aspen    poplar     (Populus    tremuloides) 
near  Springhill,  north  base  of  Laramie  Peak  (0,000  feet) 


aspen  is  perhaps  the  best  characterizing  tree  of  the  Canadian  Zone. 

Populus  balsamifera  Linnaeus.     Balsam  Poplar. 

The  balsam  poplar  inhabits  the  borders  of  many  of  the  larger  streams  in  the 
northwestern  mountains,  being  largely  confined  to  the  Canadian  Zone  between 
6,500  and  7,500  feet  elevation.  It  occurs  south  in  the  Jackson  Hole  country  at 
least  to  the  Gros  Ventre  River.  Scattering  trees  are  found  also  in  wet  gulches 
at  the  southern  base  of  the  Bear  Lodge  Mountains  near  Sundance  at  about 
5,000  feet  elevation. 
Populus  occidentalis    ( Rydberg  >    Britton.     Broad-Leaved  Cottonwood. 

The    broad-leaved    Cottonwood,    characteristic    of    the    Upper    Sonoran    Zone, 
forms  the  principal  fringe  on  the  streams  of  eastern  and  northern  Wyoming, 


1917.] 


TBEES   AND   SHRUBS. 


01 


attaining  perfection  of  growth  in  the  lowest  and  warmest  valleys.  Large  groves 
border  the  Bighorn,  Belle  Fourche,  and  especially  the  streams  of  the  lower 
Platte  drainage"  There  is  a  great  deal  of  cottonwood  growth  on  the  Cheyenne 
River  and  other  streams  southwest  of  the  Black  Hills,  hut  in  this  section  the 
species  grows  in  a  very  stunted  state,  low  and  irregular  and  very  thick  at  the 
the  base.  In  the  Wheatland  district  it  occurs  in  places  in  heavy  growth  with 
Populus  acuminata,  and  along  the  base  of  the  mountains  generally  meets  and 
commingles  with  the  narrow-leaved  Transition  species,  /'.  angustifolia. 

Populus  acuminata  Rydberg.     Lance-Leaved  Cottonwood. 

On  the  Chugwater,  Sibylee,  and  other  tributary  streams  of  the  Laramie  ami 
North  Platte  Rivers  east  of  the  mountains  the  lance-leaved  cottonwood  forms 
in  many  places  a  heavy 
fringe  with  Populus  occi- 
dentalis,  though  by  no 
means  so  generally  dis- 
tributed as  the  latter  spe- 
cies. A  splendid  growth 
of  the  lance-leaved  cot- 
tonwood is  on  Sibylee 
Creek  southwest  of 
Wheatland.  The  species 
was  not  noted  in  north- 
ern Wyoming,  but  at  the 
west  scattering  trees 
are  on  Green  Rive  r, 
just  north  of  the  Utah 
line. 

Populus      angustifolia 

James.   Narrow-Leaved 

Cottonwood.      ( Fig.  8. ) 

The  narrow-leaved  cot- 
tonwood inhabits  the  bor- 
ders of  mountain  streams 
in  the  Transition  Zone, 
mainly  at  elevations  from 
G.(i(Mi  to  7,500  feet  at  the 
west,  extending  down  to 
5,000  feet  in  the  central 
districts,  and  to  4,500 
feet  at  the  eastern  base 
of  the  Bighorn  Moun- 
tains. It  is  apparently  absent  from  the  northeastern  part  of  the  State.  Impor- 
tant streams  bordered  with  good  growth  include  the  Bear,  Green.  Wind,  and 
Greybull  Rivers,  the  north  and  south  branches  of  the  Shoshone  River  above  the 
forks,  Snake  River  below  the  mouth  of  the  Gros  Ventre,  and  the  upper  North 
Platte  down  to  18  miles  above  Casper.  Unusually  tine  groves  are  on  Wolf  and 
Big  Goose  Creeks,  at  the  eastern  base  of  the  Bighorns. 


Fig.  S. — Large  narrow-lea ved  cottonwood   (Puiiiilits  angus- 
tifolia) on  sage  flat  in  upper  Wind  River  Valley. 


Salix  amygdaloides  Anderson.     Peach-Leaved  Willow. 

The  large  peach-leaved  willow  occurs  in  scattered  clumps  along  streams  in 
the  Upper  Sonornn  Zone,  chiefly  at  the  east  and  north,  as  follows:  Chugwater 
Creek  and  affluents;  Little  Bear  Creek  northeast  of  Meadow;  Sibylee  Creek 
west  of  Wheatland  ;  Rawhide  Creek  to  base  of  Rawhide  Butte  ;  near  Lusk  ;  Hay 


62 


NORTH    AMERICAN    FAUNA. 


[No.  l: 


Creek  east  of  Aladdin;  Wind  Creek  northeast,  of  Moorcroft  :  Sheridan :  Arvada  : 
Clear  Creek,  Clearmont  to  Buffalo;  Bighorn  River  and  tributaries  near  Ther- 
mopolis;  and  Green  River  near  Utah  boundary. 

Salix  bebbiana  Sargent.     Bebb  Willow. 

The  Bebb  willow  is  characteristic  on  foothill  and  lower  mountain  slopes  up 
to  about  9,000  feet  altitude,  and  there  are  few  if  any  ranges  in  Wyoming  where 
it  is  not  present.  The  usual  growth  is  in  scattered  clumps  about  springs  and 
bogs,  but  occasionally  there  is  a  heavier  stand  on  the  margins  of  mountain 
streams.  The  height  attained  seldom  exceeds  12  or  15  feet,  and  8  or  10  feet  is 
usual.     Specimens  were  taken  on  the  Laramie.  Bear  Lodge,  Bighorn,  and  Wy- 


FiGi  9. — Terraced  copsca  of  gray-leaved   willow    {Salix  glaucops)    at   tiuibcrline,  Needle 

Mountain,  Abwuroka  Range. 


oming    Ranges,   and  at   Evanston,   while  it  was  observed   at  a   wide  range  of 
localities.     Nelson  records  it  from  Creston,  on  the  open  Red  Desert.1 

Salix  pyrifolia  obscura  Anderson.     Willow. 

Another  willow,  Salix  pyrifolia  obscura,  was  collected  in  the  upper  forests 
on  the  Wyoming  Range  west  of  Alerna,  between  9.000  and  10.000  feet  altitude, 
in  the  Canadian  Zone.     It  is  tolerably  common  at  this  locality. 

Salix  nelscni  Ball.     Nelson  Willow. 

Vernon  Bailey  collected  the  Nelson  willow  along  the  Iludsonian  crest  of  the 
Salt  River  Range  at   10,000  feet  elevation,  August  20.  1911. 

Salix  glaucops  Anderson.     Gray-Leaved  Willow.  (Fig.  9;  Rl.  XI,  fig.  2.  i 

The  gray-leaved   willow    is  a   low  species  of  Alpine  bogs,   growing   in   dense 

copses   about   2   feet   high   near   timberline  on   the   Bighorn.    Wind    River,    and 


1  Bull.   13,  Div.  of  Agrost.,  U.   S.  Dept.  Agr.,  p.  59,   1898. 


1917.]  TREES    AND    SHRUBS.  63 

Absaroka  Ranges,  and  doubtless  elsewhere.  On  Whiskey  Mountain,  south  of 
Dubois,  a  few  blossoms  were  out  on  May  14,  1910,  but  the  leaves  had  not  un- 
folded. The  species  was  not  in  leaf  by  June  5  at  timberline  on  the  Bighorn 
above  Hyattville.  It  was  very  abundant  on  Needle  Mountain,  Absaroka  Range, 
in  beautiful  terraced  copse  formation,  and  likewise  on  the  Wind  River  Moun- 
tains south  of  Fremont  Peak. 

Salix  petrophila  Rydberg.     Rock  Willow. 

The  rock  willow  was  collected  on  the  Alpine  slopes  above  timberline  at  the 
head  of  Bull  Creek,  Wind  River  Range,  in  August,  1893,  by  Vernon  Bailey. 

Salix  tenera  Anderson.    Alpine  Rock  Willow. 

Low  dense  mats  of  the  Alpine  rock  willow,  which  is  very  abundant  on  Whirl- 
wind Peak,  in  the  Absaroka  Range,  cover  portions  of  these  slopes  between  10,000 
and  11,000  feet  elevation. 

Salix  nivalis  Hooker.    Alpine  Willow. 

The  dwarf  alpine  willow  is  tolerably  common  on  the  bleak  slopes  above 
timberline  on  the  Wind  River  Range,  occurring  among  the  rocks  in  dense 
creeping  mats  a  few  inches  high.  It  is  abundant  south  of  Fremont  Peak  from 
11,500  feet  upward. 

Salix  saximontana  Rydberg.    Net-Veined  Willow. 

More  generally  dispersed  on  the  Wyoming  ranges  than  the  other  Alpine  wil- 
lows, but  similar  in  habit  of  growth,  Salix  saximontana  occasionally  extends 
a  little  below  timberline.  It  is  especially  abundant  at  the  northern  end  of 
the  Teton  Range,  where  the  creeping  mats  are  very  extensive,  and  push  down 
the  cool  Hudsonian  slopes  to  9,500  feet  altitude. 

Ostrya  virginiana   (Miller)    Willdenow.     Ironwood. 

Vernon  Bailey  reports  the  ironwood  as  abundant  in  Sand  Creek  canyon  above 
Beulah,  at  the  northern  base  of  the  Black  Hills.  The  species  <-losely  approaches 
the  Wyoming  boundary  in  northwestern  Nebraska,  8  or  10  miles  east  of  Kirt- 
ley,  Wyo.,  where  it  is  not  infrequent  in  wooded  canyons  along  the  northern 
escarpment  of  Pine  Ridge. 

Corylus  rostrata  Alton.    Beaked  Hazelnut. 

The  beaked  hazelnut  is  abundant  on  the  upper  slopes  of  the  Bear  Lodge  Moun- 
tains and  Black  Hills,  forming  dense  undergrowth  in.  aspen  and  birch  thickets 
between  5,500  and  6,000  feet  altitude.  It  appears  to  be  absent  from  the  mairi 
ranges  in  Wyoming. 

Betula  papyrifera  Marshall.    Canoe  Birch.  (Fig.  10.) 

The  canoe  birch  apparently  reaches  its  southern  limits  on  the  Bear  Lodge 
Mountains  and  northern  Black  Hills.  Over  this  region  it  grows  to  medium  size, 
and  with  the  aspen  occurs  in  dense  thickets  on  cool,  shaded  slopes  and  in  damp 
spots  as  low  as  5,000  feet  altitude,  and  on  Sundance  Creek  scattering  trees  of 
good  size  are  found  at  4,700  feet.  On  the  dry  upper  slopes  of  the  Bear  Lodge 
Mountains  this  birch  becomes  very  scrubby.  It  is  apparently  absent  from  the 
Bighorn  Mountains  and  the  ranges  of  western  Wyoming. 

Betula  fontinalis  Sargent.     Rocky  Mountain  Birch.  (Fig.  11.) 

The  Rocky  Mountain,  or  black,  birch  borders  often  in  dense  growth  most  of 
the  streams  on  the  basal  slopes  of  the  mountains,  and  under  the  cooling  influ- 
ence of  the  mountain  water  extends  some  distance  out  onto  the  plains.  At  the 
base  of  the  Bear  Lodge  Mountains  it  is  common  at  4,000  feet  altitude,  but  farther 


64 


NORTH    AMERICAN    FAUNA. 


[No.  42. 


west  it  is  usually  present  between  6,000  and  7,500  feet.     On  warm  exposed  slopes 
of  the  Wind  River  Range  north  of  Big  Sandy  it  extends  to  8,000  feet. 

Betula  glandulosa  Michaux.     Dwarf  Birch. 

The  dwarf  birch  was  noted  only  in  the  mountain  valleys  of  the  northwest, 
from  Yellowstone  Park  (West  Gallatin  and  Lewis  River  meadows)  south  to 
the  Wyoming  Range  (South  Piney  canyon  at  7,500  feet  altitude),  in  the  Canadian 
Zone.  This  little  birch  is  especially  abundant  in  the  extensive  willow  bogs  and 
swamps  bordering  Jackson  Lake,  where  it  occurs  in  dense  thickets  3  or  4  feet 
high.  Its  leaves  had  mostly  turned  to  a  deep  Indian  red  and  a  few  were  falling 
in  Jackson  Hole  by  September  13,  1910.     It  was  noted  also  as  follows:  Head  of 

Pacific  Creek,  near  Two 
Ocean  Pass;  and  Horse 
Creek  meadows,  at 
Merna,  8,000  to  9,000 
feet. 

Alnus  tenuifolia  Nut- 
tall.  Alder.  (Fig.  12.) 
The  alder  is  found  in 
a  growth  of  varying 
density  on  the  upper 
reaches  of  cold  mountain 
streams,  and  with  many 
other  Canadian  Zone 
species  follows  the  cold 
conditions  on  their  mar- 
gins down  for  some  dis- 
tance into  the  Transition 
Zone.  It  is  most  abun- 
dant in  the  mountainous 
northwest,  and  was  not 
noted  in  the  Bear  Lodge 
Mountains  or  on  the 
northern  groups  of  the 
Laramie  Range. 


Ulmus 
nseus. 
The 


Fig. 


10. — Thicket  of  canoe  birch    (Betula  papyrifera)    in 
gulch   near    Sundance,   Bear   Lodge   Mountains. 


americana    Lin- 
Elm, 
elm      penetrates 


Wyoming  for  a  short 
distance  in  the  low  val- 
leys at  the  northeast.  It  was  common  and  growing  to  large  size  on  Sand  Creek 
to  10  miles  above  Beulah  at  the  northern  base  of  the  Black  Hills,  and  on  Hay 
Creek  west  to  Aladdin  and  Eothen,  while  it  is  reported  at  Hulett  in  the  Belle 
Fourche  Valley.     A'ernon  Bailey  found  it  on  Little  Powder  River  near  Morse. 

Quercus  macrocarpa  Michaux.     Bur  Oak. 

The  bur  oak  extends  into  Wyoming  from  the  northeast  and  is  found  in 
abundance  over  a  small  area  in  Crook  County,  principally  east  of  the  Belle 
Fourche  River  and  north  of  Linden  and  Inyankara.  It  occurs  in  scattered 
groves  on  the  partially  open  basal  flanks  of  the  Bear  Lodge  Mountains  and  at 
the  northern  base  of  the  Black  Hills,  extending  to  the  dry  Bear  Lodge  summits 
at  0,000  feet  elevation,  where  Vernon  Bailey  found  scrubby  thickets  4  or  5  feet 


1917.] 


TREES    AND    SHRUBS. 


65 


high,  loaded  with  acorns,  in  August,  1913.  In  some  of  the  stream  valleys  the 
bur  oak  grows  to  large  lumber  size,  notably  on  Sand  Creek  above  Beulah,  and 
many  trees  were  there  noted  with  clean  straight  trunks  of  good  height  from  3 
to  4  feet  in  diameter  at  the  base.  The  vertical  range  of  this  oak  is  about  the 
same  as  that  of  the  yellow  pine,  with  which  it  usually  commingles  in  this 
region.  Vernon  Bailey  noted  a  little  oak  growth  near  the  head  of  the  Little 
Missouri  River,  apparently  its  western  limit  in  the  State. 

Atriplex  canescens  James.     Saltbush ;  Gray  Shadscale. 

The  various  saltbushes  are  characteristic  Sonoran  species  of  the  arid  Great 
Basin  region,  and  barely  enter  the  Great  Plains  area  on  some  of  the  dry  valley 


:M 

n  Jfi  * 

■ 

■jii. 

■>£  ■  I  ^fl^iPS 

Uti 

■ 

' 

:     •    ■ 

-^. 

Fig.   11. 


-Clumps  of  Rocky  Mountain  birch    (Bctula  fontinalis)    15  feet  high,  at  north 
base  of  Shirley   Mountains. 


flats  at  the  eastern  base  of  the  foothills.  They  are  mostly  alkali-resistant  and, 
with  the  possible  exception  of  Atriplex  canescens,  furnish  valuable  winter  forage 
for  sheep  in  the  central  desert  sections.  Of  the  three  principal  shrubby  species 
found  in  Wyoming,  .4.  canescens  is  the  least  abundant.  It  is  apparently  absent 
from  the  Red  Desert  proper  where  A.  confertifoMa  and  .4.  nuttalli  abound,  but 
extends  farther  to  the  east  than  either  of  the  foregoing.  Atriplex  canescens  oc- 
curs in  very  dense  growth  3  feet  or  more  high  on  dry  flats  along  the  Chugwater 
at  Bordeaux,  and  also  in  the  sand  along  the  Laramie  at  Uva,  but  is  in  scatter- 
ing growth  elsewhere. 

Atriplex  confertifolia  S.  Watson.     Round-Leaved  Saltbush. 

The  round-leaved   saltbush   is   a   low   stocky   shrub   1    or   2   feet   high,    very 
abundant  on  sandy  and  alkaline  soils  up  to  7,500  feet  elevation  on  warm  slopes 
east  of  Green  River  near  the  Utah  line  ;  bad  lands  south  of  Lyman,  to  7,000  feet ; 
74440°— 17 5 


66 


NORTH   AMERICAN    FAUNA. 


[No.  42. 


Bear  River  below  Evanston,  to  6.600  feet ;  Fossil ;  Bigpiney ;  Dubois ;  Trout 
Creek,  north  Shoshone  Valley ;  Bighorn  slopes  above  Hyattville,  to  5,500  feet ; 
north  base  of  Rattlesnake  Mountains,  to  6,500  feet ;  Splitrock,  Sweetwater  Val- 
ley ;  east  to  Fort  Steele,  Shirley,  Old  Fort  Fetterman,  and  Arvada. 

Atriplex  nuttalli  S.  Watson.    Nuttall  Salthush.  (PI.  IV,  fig.  2.) 

The  Nuttall  saltbush,  a  low-spreading  species,  commonly  known  as  "salt 
sage  "  to  the  sheepmen,  is  of  great  economic  value,  as  it  affords  the  chief  winter 
food  to  the  flocks  on  the  Red  Desert.  It  has  about  the  same  range  as  Atriplex 
confertifolia,  but  is  seldom  found  in  sand,  being  partial  to  dry  adobe  and 
saline  flats,  where  it  forms  often  the  dominant  plant  growth.     Extensive  flats 

at  Frannie  and  else- 
where in  the  Bighorn 
Basin  are  covered  with 
a  pure  and  uniform 
growth  of  this  saltbush. 
The  observed  eastern 
limits  are  Walcott ;  Lit- 
tle Medicine  Bow  River 
west  of  Marshall ;  Indian 
Creek  at  State  line  north 
of  Kirtley ;  Newcastle ; 
and  Colony. 

Grayia   spinosa   Moquin. 
Grayia. 

A  characteristic  spiny 
shrub  of  the  Sonoran 
desert  tracts  from  the 
Sweetwater  Valley  west- 
ward, especially  abun- 
dant in  sandy  or  adobe 
soils  at  the  lowest  levels. 
On  the  sandy  hummocks 
between  Frannie  and 
Garland  at  the  end  of 
June  dense  clusters  of 
flat,  winged  seeds  were 
borne  in  great  profusion 
on  the  grayia  bushes. 
These  varied  from  green- 
ish to  pink  or  purple- 
brown  and  lent  a  peculiar  hue  to  the  landscape.  This  shrub  has  considerable 
forage  value,  as  its  thick  leaves  and  seeds,  gathered  into  drift  piles  under  the 
bushes,  are  eagerly  eaten  by  sheep  in  winter. 

Grayia  spinosa  was  common  from  Independence  to  Splitrock;  Dry  Lake  to 
Lorey ;  on  first  and  second  benches  above  the  Platte  at  Fort  Steele ;  Worland ; 
Manderson ;  Bonanza ;  Greybull  west  nearly  to  Cody ;  Rock  Springs  region ; 
Green  River  near  Utah  line;  Carter  to  Lyman;  bad  lands  south  of  Lyman; 
Cumberland ;  Fontenelle  to  Opal ;  and  Green  River  flats  north  nearly  to 
Labarge. 

Sarcobatus  vermiculatus  Torrey.    (Jreasewood.  (PI.  IV,  ivj;.  1.) 

This  common  desert  shrub  is  of  wide  dispersion  in  the  more  arid  por- 
tions of  the  Upper  Sonoran  Zone  and  was  noted  at  numerous  localities.     It 


Fig.    12. 


BIISC9 

-Fringe    of    alder    (Alnns    tenuifolia)    on    Pacific 
Creek  below  Two  Ocean  Pass. 


1017.]  TREES   AND    SHRUBS.  07 

forms  a  very  rank  growth  on  the  dry  alkaline  lake  basins  and  flats  of  central 
Wyoming,  especially  in  the  Red  Desert  region,  where  its  dark-green  foliage 
and  large  size  make  it  the  most  conspicuous  of  the  alkali-resistant  shrubs. 
The  greasewood  follows  dry  adobe  valley  flats  to  about  7,000  feet  elevation. 

Berberis  aquifolium  Pursh.     Barberry;  Oregon  Grape. 

The  Oregon  grape,  a  low  undershrub,  is  characteristic  of  dry  forested  or 
partly  wooded  slopes  throughout  the  State,  mainly  in  the  Transition  Zone.  It 
is  commonly  associated  with  the  bearberry,  with  which  it  penetrates  to  the 
lower  edge  of  the  Canadian  forest  belt.  On  the  Ferris  Mountains  it  was  noted  at 
8,700  feet  altitude,  and  on  the  west  side  of  the  Salt  River  Range  was  common 
to  9,000  feet. 

Ribes  longiflorum  Nuttall.     Flowering  Currant. 

In  early  summer  the  brush  fringe  along  streams  at  the  lower  elevations  is 
enlivened  by  the  yellow  bloom  of  the  flowering  currant.  This  shrub  is  a  char- 
acteristic Sonoran  species,  but  is  perhaps  most  conspicuous  at  the  base  of  the 
mountains,  where  it  penetrates  the  foothills  for  some  distance  in  warm  stream 
valleys.  Its  upper  limits  were  observed  as  follows:  Foothills  west  of  Wheat- 
land, to  5,300  feet  altitude;  eastern  base  of  Bighorn  Mountains;  Greybull  River 
at  Meeteetse;  Pat  O'Hara  Creek  to  6,000  feet;  south  slope  of  Owl  Creek  Moun- 
tains to  7,000  feet;  Bull  Creek,  Wind  River  Valley;  streams  of  Salt  River 
Valley;  Evanston  and  west  slope  of  Bear  River  Divide  to  7,500  feet;  Mountain- 
view  ;  and  Henry's  Fork  of  Green  River  to  Burntfork  P.  O. 

Ribes  inebrians  Lindley.     Red  Currant. 

The  red  currant  is  widely  distributed  in  the  Transition  Zone  over  most  of 
Wyoming,  extending  regularly  to  the  lower  edge  of  the  Canadian  Zone.  Flower- 
ing specimens  were  taken  in  the  Laramie  foothills  west  of  Islay  on  June  16, 
1909,  and  near  Merna,  Wyoming  Range,  at  8,000  feet  altitude,  as  late  as  Au- 
gust 10,  1911.     This  species  is  partial  to  rocky  situations. 

Ribes  montigenum  McClatchie.     Bristly  Red  Currant. 

A  high-altitude  species,  the  bristly  red  currant  has  a  general  distribution  on 
the  higher  Wyoming  ranges.  It  is  most  abundant  near  timberline,  where  it 
occurs  either  as  scattering  bushes  in  slide  rock,  or  in  dense  patches  a  foot 
or  two  high  on  deep-soiled  slopes,  as  on  the  Wyoming  Range  west  of  Merna.  It 
was  in  flower  on  the  summit  of  Bridget'  Peak  in  the  Sierra  Madre,  on  July 
7,  1911,  and  still  so  on  the  high  Wind  River  Range  south  of  Fremont  Peak  on 
July  18.  The  red,  edible  fruit  was  abundant  and  fully  ripe  in  the  timberline 
region  on  the  Teton  Range  on  August  30,  1910.  It  was  found  in  abundance  on 
the  high  ridge  extending  north  from  Needle  Mountain,  in  the  Absaroka  Range, 
between  10,000  and  11,000  feet  elevation ;  and  also  on  the  Salt  River  Range, 
above  9,000  feet. 

Ribes  lacustre  (Persoon)  Poiret. 

This  species  occurs  in  Canadian  Zone  forests  in  northwestern  Wyoming.  It 
was  still  in  flower  near  the  upper  end  of  Fremont  Lake  on  July  15,  1911,  but 
was  bearing  fruit  on  Grinned  Creek.  Absaroka  Range  (8,000  feet  altitude), 
July  30,  1910.     Specimens  were  collected  al  both  localities. 

Ribes  petiolare  Douglas.     Mountain  Black  Currant. 

The  mountain  black  currant,  a  wide-ranging  Boreal  species,  was  collected 
on  the  summit  of  the  Bear  Lodge  Mountains  on  June  20,  1912,  where  it  was  not 
uncommon  on  the  dry  crests  at  6,000  feet  elevation.     Flowering  specimens  were 


68  NORTH   AMERICAN    FAUNA.  [No.  42. 

also  taken  in  the  Canadian  Zone  forest  near  the  upper  end  of  Fremont  Lake, 
Wind  River  Range,  July  15,  1911,  and  fruiting  specimens  at  Tower  Falls,  Yel- 
lowstone Park,  August  11,  1910.  It  is  especially  abundant  in  the  Yellowstone 
Park  forests. 

Ribes  viscosissimum  Pursh.     Currant. 

This  currant  is  characteristic  of  the  Canadian  Zone  forest  belt  in  northern 
Wyoming.  Flowering  specimens  were  taken  at  8,500  feet  elevation  on  the 
head  of  Shell  Creek,  Bighorn  Mountains,  June  5,  1910;  near  the  upper  end  of 
Fremont  Lake,  Wind  River  Range,  July  15,  1911 ;  and  at  8,000  feet  on  Grinned 
Creek,  Absaroka  Range,  as  late  as  July  30,  1910.  Vernon  Bailey  collected  the 
species  on  Wolf  Creek,  northern  slope  of  the  Bighorns,  August  10,  1913. 

Edwinia  americana  (Torrey  and  Gray)   Heller.     Edwinia. 

The  low  flowering  edwinia  was  found  only  on  the  crest  of  the  Laramie 
Range  east  of  Laramie,  between  8,500  and  9,000  feet  elevation.  Its  handsome 
white  cyinous  bloom  enlivened  the  rock  ledges  and  cliffs  on  June  18,  1909. 

Cercocarpus  ledifolius  Nuttall.     Mountain  Mahogany. 

The  mountain  mahogany  is  a  small  evergreen  tree  or  stout  shrub  peculiar 
to  rocky  plateaus  and  ridges  and  warm  exposed  basal  slopes  of  mountains. 
In  Wyoming  a  scattering  distribution  is  indicated  from  the  lower  Green  River 
Valley  north  to  Jackson  Hole,  in  the  Transition  Zone.  Most  of  the  rocky 
ridges  adjacent  to  Green  River  near  the  Utah  boundary  are  clothed  up  to 
7,500  feet  elevation  with  dense  scrubby  thickets  from  3  to  6  feet  in  height. 
The  species  was  not  found  farther  north  in  the  Green  River  country,  btit 
enters  the  State  from  the  west  along  Snake  River,  and  covers  in  good  growth 
the  more  exposed  of  the  lower  western  slopes  of  the  Salt  River  Range  east 
of  Afton  and  Smoot  up  to  7,500  feet.  Its  upper  limits  in  the  Snake  River 
drainage  are  reached  apparently  near  Jackson,  where  Edward  A.  Preble  found 
a  considerable  growth  on  a  warm  slope  at  7,000  feet  in  a  tributary  gulch  of 
Cache  Creek. 

Cercocarpus  intricatus  Watson.  Mountain  Mahogany.  (PI.  XV,  figs.  1  and  2.) 
An  abundant  evergreen  shrub  on  warm  open  slopes  of  the  Bighorn  Mountains. 
(in  the  eastern  side  scattering  bushes  dot  t lie  bare  Tongue  River  bluffs  at 
Rancbester.  The  species  forms  dense  thickets  3  feet  high  at  about  5,000  feel 
altitude  near  Eaton's  Ranch,  and  thence  it  ascends  dry,  rocky  ridges  to  6,500 
feet  on  bare  exposed  points  in  the  mountains  south  of  Wolf.  On  the  warm 
western  side  of  the  Bighorns  above  Hyattville  Cercocarpus  intricatus  was  not 
seen  below  5.800  feet,  but  between  6,000  and  6,500  feet  the  dry,  hot,  reddish 
slopes  were  dotted  with  this  intricately  branched,  steely  gray  shrub. 

Cercocarpus  parvifolius  Nuttall.    Mountain  Mahogany.  (PI.  II,  fig.  1.1 

This  mountain  mahogany  occurs  mainly  in  the  Transition  Zone  in  Wyoming, 
although  in  the  southern  Rocky  Mountains  its  center  of  abundance  is  in  the 
juniper  and  pinyon  belt  of  the  Upper  Sonoran  Zone.  It  is  partial  to  warm, 
cocky  situations,  either  partly  open  foothill  slopes  or  outlying  ridges  and 
buttes  on  adjacent  plains  or  deserts.  The  largest  growth  of  mountain  mahogany 
observed  was  fully  9  feet  high,  on  the  pine-clad  foothills  southwest  of  Wheat- 
land, at  5,300  feet  altitude.  The  usual  height  attained  is  from  4  to  0  feet. 
Unusually  extensive  thickets  grow  on  open  ridges  paralleling  I  he  Horse  Creek 
valley  between  l>a\is  Ranch  and  Meadow.  This  shrub  flowers  early  in 
.lime,  but  occasionally  a  little  later.  A  flowering  specimen  from  Steamboat 
Mountain  was  collected  June  26,  1913.     The  distribution  of  Cercocarpus  parvi- 


North  American  Fauna  No.  42,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  XV. 


■*».«!«:-. 


♦9i 


^.*i*:  .vl^.'f  >\^/,  ^i^**^' 


Fig.  1.— Mountain  Mahogany  (Cercocarpus  intricatus). 
•  West  slope  of  Bighorn  Mountains  above  Hyattville  (6,000  feet). 


Fig.  2.— Nearer  View  of  Same. 


1017.]  TREES   AXD   SHRUBS.  69 

f  alius  as  noted  by  Survey  parties  is  entirely  south  of  ;i  line  from  Newcastle 
in  the  northeast  to  Evanston  in  the  southwest. 

Kunzia  tridentata  (Pursh)  Sprengel.    Antelope  Brush. 

The  antelope  brush  is  a  common  shrub  on  dry  open  Transition  slopes  in 
southern  and  western  Wyoming,  being  especially  abundant  on  arid  hills,  pla- 
teaus, and  occasional  sand  ridges  in  the  borders  of  the  Red  Desert  at  eleva- 
tions between  7,000  and  8,500  feet.  It  was  not  noted  north  of  the  Wind  River, 
Rattlesnake,  and  Casper  Ranges,  but  it  is  not  uncommon  on  dry  slopes  and 
sagebrush  flats  south  of  Buffalo  Creek  and  the  Gros  Ventre  River  in  Jackson 
Hole.  With  the  sagebrush  {Artemisia  tridentata)  this  shrub  regularly  ascends 
warm  slopes  and  occurs  in  dry  parks  and  openings  to  the  lower  edge  of  the 
Canadian  Zone.  Its  vertical  range  varies  from  4,500  feet  on  ridges  along 
the  North  Platte  near  Glendo  to  9,000  feet  on  the  summits  of  the  Laramie 
and  Ferris  Ranges,  and  also  on  the  warm  southern  slopes  of  the  Wind  Rivers 
north  of  Big  Sandy. 

Holodiscus  dumosus  (Nuttall)  Heller. 

Common  locally  on  rocky  slopes  in  some  of  the  dry  desert  mountains  and 
in  the  rougher  borders  of  the  Red  Desert,  mainly  in  the  Transition  Zone. 

Opulaster  pubescens  Rydberg.     Ninebark. 

On  dry,  steep,  basal  mountain  slopes  at  the  head  of  Pat  O'Hara  Creek,  north- 
west of  Cody,  the  ninebark  forms  a  low  but  exceedingly  dense  chaparral 
among  partly  dead  Douglas  spruce  and  white  pine  forests  up  to  6,500  feel 
elevation.  Along  the  northern  base  of  the  Bighorn  Mountains  near  Wolf  it 
was  likewise  partial  to  dry  slopes  between  5.000  and  7,000  feet. 

Opulaster  monogynus  (Torrey)  Kuntze.     Western  Ninebark. 

The  western  ninebark,  a  Transition  Zone  shrub,  is  tolerably  common  on  the 
Laramie  Mountains  and  on  outlying  ridges  of  the  plains  region  to  the  eastward, 
at  elevations  between  5,000  and  7,000  feet. 

Opulaster  malvaceus   (Greene)   Kuntze.     Ninebark. 

This  ninebark  is  very  abundant  and  conspicuous  on  the  warm  western 
slopes  of  the  Salt  River  Range  near  Afton,  but  was  not  found  elsewhere.  Its 
vertical  range  in  these  mountains  is  from  their  bases  up  to  7,100  feet  in 
canyons  and  to  7.500  feet  on  warm  slopes,  and  like  the  other  Wyoming  species 
of  the  genus  it  is  mainly  restricted  to  the  Transition  Zone. 

Spiraea  lucida  Douglas.     Meadowsweet. 

The  handsome  flowering  meadowsweet  is  sparingly  present  across  Wyoming 
at  the  north  in  the  lower  part  of  the  Canadian  forest  belt.  Alexander  Wetmore 
collected  it  at  7.000  feet  elevation  on  the  east  side  of  Teton  Pass  on  September 
9.  1910,  and  I  have  observed  it  down  to  5,800  feet  in  the  Bighorn  Mountains 
near  Wolf,  and  in  abundance  on  the  Bear  Lodge  Mountains  above  5,500  feet. 

Vernon  Bailey  reports  a  pink-flowered  species,  probably  Spiraea  densiflora, 
between  7,000  and  8,000  feet  on  the  eastern  slope  of  the  Bighorns,  above  Wolf. 

Dryas  octopetala  Linnaeus.     Alpine  Avens. 

The  low  alpine  avens  is  restricted  to  Arctic  and  Alpine  regions.  In  Wyoming 
I  collected  it  on  the  Bighorn  Mountains,  where  it  was  in  characteristic  pros- 
trate matted  growth  on  the  rocky  slopes  above  timberline  at  the  head  of 
Trapper  Creek,  being  not  yet  in  flower  on  June  10,  1910.  Vernon  Bailey 
collected  it  in  August,  1893,  on  the  high  Wind  River  Range  at  the  head  of 
Bull  Creek. 


70 


NORTH    AMERICAN    FAUNA. 


[No.  42. 


Rubias  deliciosus  James.     False  Raspberry. 

The  handsome  false  raspberry  was  abundant  and  still  flowering  on  the  pine- 
clad  foothill  ridges  southwesl  of  Wheatland  on  July  4.  1909,  at  elevations  be- 
tween 5,000  and  5,500  feet.  This  is  probably  near  its  northern  limit,  as  it  was 
not  noted  in  the  Laramie  Peak  region  nor  on  any  of  the  ranges  farther  west. 

Rubus  parviflorus  Nuttall.     Flowering  Raspberry ;  Thimbleberry.      ( Fig.  13. ) 

The  flowering  raspberry  is  a  tolerably  common  fruiting  species  in  Cana- 
dian Zone  woods  on  most  of  the  Wyoming  ranges,  especially  at  the  north. 
The  large  leaves,  showy  white  flowers  in  June  and  early  July,  and  in  most  years 
an  abundance  of  handsome  red  berries  in  late  August,  make  the  thimbleberry 
very  conspicuous  in  the  woodlands  between  7,000  and  0,000  feet  elevation.     The 


BII64S 

Fn;.  13. — Flowering  raspberry   (Rubua  parviflorus)   In  cool  gulch,  north  base  of  Casper 

Mountains  ((1.500  feet). 

large  berries,  which  are  pleasantly  flavored,  though  somewhat  dry  and  seedy, 
are  eaten  by  many  native  birds  and  mammals. 

Rubus  strigosus  Michaux.     Red  Raspberry. 

The  red  raspberry  is  abundant  at  numerous  localities  in  the  mountain  dis- 
tricts from  the  lower  edge  of  the  forest  belt  to  timberline.  Its  observed  vertical 
range  is  from  4,500  feet  at  Wolf,  at  the  north  base  of  the  Bighorn  Mountains, 
to  11,700  feet,  above  timberline,  south  of  Fremont  Peak,  Wind  River  Range. 
It  reaches  its  best  growth  and  bears  most  abundantly  on  rocky,  partially  for- 
ested slopes  in  upper  Transition  and  lower  Canadian  Zones  on  the  northwestern 
ranges.  In  the  heavy  forests  between  7,000  and  8,000  feet  elevation  on  Grin- 
ned Creek,  in  the  Absaroka  Range,  the  fruit  was  just  ripening  on  August  1, 
1910,  while  on  the  western  slope  of  the  Salt  River  Range  the  bushes  were  full 


1917.]  TREKS   AND   SHRUBS.  71 

of  berries  on  August  20,  1911.  in  the  timberline  region  the  species  occurs  in 
a  dwarfed  state,  usually  uol  over  a  foot  nigh  in  fields  of  slide  rock.  Rock- 
strewn  paths  of  avalanches  throughout  the  mountains  support  more  or  less 
raspberry  growth. 

Dasiphora  fruticosa  (Linnaeus)  Rydberg.    Shrubby  Cinquefoil. 

The  low  shrubby  cinquefoil  inhabits  cold  mountain  bogs  and  meadows  and 
is  more  common  on  the  ranges  aleag  the  western  edge  of  the  State  than  farther 
oast.  It  is  especially  abundant  in  the  cold  stream  meadows  along  the  northern 
base  of  the  Uinta  Mountains,  in  the  upper  Green  River  Basin,  and  at  the  head 
of  Wind  River;  in  the  willow  swamps  of  Jackson  Hole  and  Yellowstone  Park; 
and  in  the  timberline  region  on  the  Wind  River  and  Absaroka  Ranges.  It 
extends  above  timberline  on  deep-soiled  slopes,  but  at  the  higher  altitudes  is 
depauperate,  from  a  few  inches  to  a  foot  in  height. 

Amelanchier   alnifolia   Nuttall.      Serviceberry. 

Several  species  of  serviceberry  are  characteristic  of  the  Transition  Zone  of 
Wyoming,  but  Amelanchier  alnifolia  appears  to  lie  of  widest  range.  On  the  less 
arid  foothills  and  lower  mountain  slopes  at  the  east  and  north,  where  it  is  most 
abundant,  it  is  usually  the  only  species  present,  but  in  the  central  and  southern 
districts  it  occurs  in  places  with  .1.  oreophila,  .1.  elliptica,  and  possibly  others.1 
Typical  specimens  are  from  the  Bear  Lodge  Mountains.  It  was  noted  at  locali- 
ties too  numerous  for  inclusion. 

Amelanchier  elliptica  A.  Nelsou.     Serviceberry. 

Taken  only  in  the  southwest,  where  it  is  the  predominating  species  of  Ame- 
lanehier  in  the  western  Transition  borders  of  the  Red  I  >esert  region  at  elevations 
up  to  8,000  feet.  It  was  observed  at  a  number  of  points  on  the  Bear  River  Divide 
from  Hilliard  north  to  Cokeville.  and  also  in  (he  badlands  south  of  Lyman. 
S.  G.  Jewett  collected  flowering  specimens  at  Spring  Valley  on  June  9,  1912,  and 
also  on  Steamboat  Mountain  on  June  2.1. 

Crataegus  cerronis  A.  Nelson.     Hawthorn. 

Several  species  of  hawthorn  are  represented  in  Wyoming,  where  they  form  a 
characteristic  fringe  on  streams  in  the  Transition  Zone.  Crataegus  cerronis  is 
apparently  the  predominating  form  in  the  Bighorn  Mountains,  and  possibly 
east  to  the  Black  Hills.  It  was  blooming  profusely  on  .lime  0.  1912,  in  the 
heavy  deciduous  fringe  along  Wolf  Creek,  at  the  northern  base  of  the  Bighorns, 
up  to  4,800  feet.2    Flowering  examples  were  collected  at  this  point. 

Crataegus  rivularis  Nuttall.     Black  Hawthorn. 

The  black  hawthorn  is  common  in  southwestern  Wyoming  north  to  the  Salt 
River  Valley  and  Green  River  Basin  and  east  at  least  to  the  upper  Platte,  at 
elevations  from  6.000  to  8,000  feet. 

Sorbus  scopulina  Greene.    Mountain  Ash. 

In  the  Wyoming  ranges  the  mountain  ash  is  usually  ft  low  or  medium-sized 
shrub  3  or  4  feet  high,  growing  in  scattered  clumps  or  occasionally  forming 
small  thickets,  although  in  the  Shirley  Mountains  a  few  clumps  0  feet  or  more 
in  height  were  noted.  It  is  very  generally  distributed  in  the  forests  of  the 
Canadian  Zone  up  to  8.500  or  9,000  feet  altitude. 

1  Data  collated  under  Amelanchier  alnifolia  may  include  some  related  to  other  species, 
being  based  mainly  on  field  identifications.  Unfortunately,  tew  specimens  of  Amelan- 
chier have  been  preserved. 

2  Vernon  Bailey  found  the  hawthorn  up  to  0,000  feet  near  Wolf. 


72  NORTH    AMERICAN    FAUNA. 


12. 


Primus  americana  Marshall.     Wild  Red  Plum. 

Scattered  scrubby  thickets  of  wild  red  plum  are  met  with  in  gulches  and  on 
streams  of  tbe  northeastern  counties  e;ist  of  tbe  Bighorn  Mountains,  the  species 
being  perhaps  most  abundant  on  the  basal  slopes  of  the  Bear  Lodge  Mountains 
and  at  the  northern  base  of  the  Black  Hills. 

Prunus  pennsylvanica  Linmeus  f.    Wild  Red  Cherry. 

The  wild  red  cherry  is  sparingly  present  in  northern  Wyoming  from  the  up- 
per slopes  of  the  Black  Hills  and  Bear  Lodge  Mountains  west  nearly  to  Yellow- 
stone Park.  On  the  dry,  partially  open  summit  of  the  Bear  Lodge  Mountains 
it  grows  in  a  dwarfed  state.  Shrubs  not  over  2  feet  high  were  flowering  pro- 
fusely at  6,000  feet  altitude  on  June  20,  1912,  and  considerable  fruit  was  found 
by  Vernon  Bailey  near  Welcome  late  in  August,  1913.  Fruit  was  reddening  at 
the  end  of  July,  1910,  on  the  North  FofK  of  the  Shoshone  River. 

Prunus  melanocarpa   (A.  Nelson)   Rydberg,     Chokecherry. 

The  chokecherry  is  the  predominating  species  of  Prunus,  being  omnipresent 
in  the  foothills  and  mountains  below  8,000  feet  elevation,  and  on  the  Laramie 
Range  reaching  9.000  feet.  It  is  most  abundant  on  dry,  open,  or  partly  forested 
Transition  Zone  slopes,  where,  however,  it  occurs  usually  in  low  scraggy 
thickets.  It  attains  its  perfection  in  damp  gulches  and  along  streams.  The 
fragrant  white  bloom  is  conspicuous  in  May  and  early  in  June. 

Amorpha  canescens  Pursh.     False  Indigo. 

The  low  silvery  false  indigo,  a  shrub  of  the  open  Great  Plains  area,  barely 
enters  the  State  at  the  northeast  in  the  Belle  Fourche  Valley.  In  June,  191:2. 
it  was  common  near  Aladdin  on  open  grassy  slopes  bordering  Hay  Creek 
valley. 

Rhus  rydbergi  Small.    Western  Poison  Ivy. 

The  western  poison  ivy  was  occasionally  noted  at  the  eastern  base  of  the 
mountains:  base  to  summit  of  Rawhide  Butte,  near  Lusk;  and  Wolf  Creek, 
Bighorn  Mountains,  below  Eaton's  Ranch. 

Schmaltzia  glabra  (Linmeus)   Small.     Smooth  Sumac. 

The  smooth  sumac  was  noted  by  Vernon  Bailey  on  the  lower  eastern  slopes 
of  the  Bighorn  Mountains  near  Wolf.  August  10,  1913.  It  was  sparingly  present 
between  4,000  and  6.000  feet  elevation  in  this  locality,  but  has  not  been  noted 
elsewhere  in  Wyoming  by  Survey  tield  parties. 

Schmaltzia  trilobata  (Nuttall)   Small.     Skunk  Bush. 

The  skunk  bush,  characteristic  of  the  Upper  Sonoran  Zone,  fringes  water- 
courses at  the  lower  elevations  up  to  5,000  or  (i.OOO  feet,  and  on  especially  warm 
slopes  has  been  noted  at  a  little  over  7,000  feet.  The  heaviest  growth  is  along 
streams,  as  at  Greybull.  where  there  are  dense  thickets  S  or  10  feet  high  in 
the  cottonwood  growth  along  the  Bighorn.  A  scrubby  growth  clothes  the  hot 
slopes  in  the  bad  lands  between  Greybull  and  Ionia  to  a  considerable  elevation. 
On  Horse  Creek  north  of  Cheyenne,  and  also  near  Wheatland  and  Cassa,  it 
grows  on  valley  flats  with  sagebrush.  In  the  warm  districts  generally  it  oc- 
curs in  scattering  growth  in  dry  gulches  and  on  exposed  faces  of  bluffs. 

Pachystima  myrsinites   (Pursh)   Rafinesque. 

A  low  evergreen  shrub  of  the  Canadian  forest  belt  in  Wyoming,  chiefly  in  the 
western  and  southwestern  mountains.  Its  vertical  limits  are  approximately 
6,000  and  10,000  feet,  but  it  is  most  abundant  between  7,000  and  9,000  feet. 
« >n    the   Salt    River   Range   near  timberline   it   was   growing   abundantly    in   a 


1917.]  TREES  AND   SITRUBS.  73 

dwarfed  state  a  few  inches  high.  The  small  green  flowers  appear  quite  early, 
the  flower  buds  being  evidenl  on  a  specimen  collected  al  Jackson,  May  4,  L911, 
by  Edward  A.  Preble. 

Acer  negundo  Linnaeus.    Box  Elder. 

The  box  elder  was  noted  chiefly  along  streams  and  in  wet  draws  of  the 
Great  Plains  area,  but  also  at  Bonanza  in  the  Bighorn  Basin,  and  between 
Dixon  and  Baggs  along  the  southern  boundary  of  the  State. 

Acer  glabrum  Torrey.    Mountain  Maple. 

The  handsome  mountain  maple  inhabits  cool  slopes,  gulches,  and  damp  spots 
generally  from  the  base  of  the  mountains  to  8,000  or  9, (too  feet  elevation,  being 
most  abundant  in  southeastern  Wyoming.  It  was  not  noted  in  the  Black  Hills 
region,  although  found  sparingly  in  cool  north  gulches  at  Squaw  Butte,  on  the 
Wyoming-Nebraska  line  east  of  Kirtley.  Its  usual  height  is  5  or  6  feet  in  the 
mountains,  and  it  rarely  exceeds  8  feet  even  along  streams. 

Acer  grandidentatum  Nuttall.     Large-Toothed  Maple. 

Scattered  clumps  of  the  large-toothed  maple,  10  or  12  feet  in  height,  are  on 
the  warm  lower  western  slopes  of  the  Salt  River  Range  near  Afton.  The  verti- 
cal range  in  these  mountains  is  from  their  6,300-foot  bases  up  to  7.300  feet,  in 
the  Transition  Zone,  and  it  does  not  here  attain  its  maximum  growth.  On  the 
Salt  River  Range  it  is  associated  with  the  mountain  maple,  but  does  not  occur  as 
high  as  the  latter. 

Rharnnus  alnifolia  L'Heritier.     Buckthorn. 

The  buckthorn  reaches  a  short  distance  into  Wyoming  at  the  west,  in  the 
region  contiguous  to  Snake  River,  but  was  not  detected  elsewhere.  It  is  a 
conspicuous  shrub  on  stream  margins  and  in  wet  willow  bottoms  in  the  borders 
of  Jackson  Hole,  and  was  abundant  on  Pacific  Creek,  15  miles  northeast  of 
Moran.  It  was  bearing  its  large  ripe  black  berries  in  Webb  Canyon  in  the 
Teton  Mountains,  at  6.700  feet  elevation,  September  1,  1910,  and  was  collected 
by  Alexander  Wetmore  on  Trail  Creek,  near  Teton  Pass,  late  in  September. 

Ceanothus  velutinus  Douglas.    Mountain  Balm.  (Fig.  11.) 

Throughout  the  mountains  one  of  the  most  characteristic  shrubs  is  the  moun- 
tain balm,  in  many  places  forming  a  dense  chaparral  2  or  3  feet  high  on  dry. 
open  or  partly  forested  slopes  mainly  in  the  Transition  Zone.  It  was  found  in 
greatest  abundance  on  dry  summits  of  medium  elevations,  as  the  Bear  Lodge. 
Casper.  Shirley,  Ferris,  and  Rattlesnake  Mountains,  on  the  8,000-foot  divide 
between  the  head  of  Salt  River  and  Smiths  Fork,  and  on  Little  Mountain,  the 
elevated  plateau  between  Maxon  and  the  Green  River  Valley.  The  vertical 
limits  are  from  5,000  feet  near  AVolf,  at  the  eastern  base  of  the  Bighorn  Moun- 
tains, to  9,500  feet  on  Bridger  Peak  in  the  Sierra  Madre.  The  white,  sweet- 
scented  flowers  were  conspicuous  in  Sierra  Madre  forests  west  of  Grand  En- 
campment between  8,500  and  9,500  feet,  July  7.  1911,  and  the  species  was  still  in 
partial  flower  on  the  Wind  River  Range  north  of  Rig  Sandy  July  26,  although 
past  flowering  near  Fremont  Lake  on  the  same  range  by  July  20. 

Ceanothus  fendleri  Gray.     Wild  Tea  Bush. 

The  low,  much-branched  wild  tea  bush  is  common  on  dry,  warm  slopes  near 
Springhill,  north  of  Laramie  Peak.  It  was  found  in  open  yellow-pine  forest  up 
to  7,400  feet  elevation,  but  was  not  noted  elsewhere  within  the  State. 

Ceanothus  mollissimus  Torrey.    Wild  Tea  Bush. 

Vernon  Bailey  found  this  wild  tea  bush  on  Big  Goose  Creek,  at  the  eastern 
base  of  the  Bighorn  Mountains.     It  is  apparently  rare  in  Wyoming. 


74 


NORTH    AMERICAN    FAUNA. 


[No.  42. 


Elseagnus  argentea  Pursh.     Silverberry. 

The  silverberry  occurs  across  Wyoming  ;it  the  west,  where  it  forms  a  scatter- 
ing silvery  fringe  on  stream  banks  mainly  in  the  Transition  Zone.  It  is  espe- 
cially abundant  on  streams  of  the  upper  Green  River  Basin  and  on  tributaries 
of  Wind  and  Snake  Rivers.  Elsewhere  it  was  observed  at  Maxon,  Lone- 
tree,  Cokeville,  Meeteetse,  on  the  South  Fork  of  the  Shoshone  River,  Pat  O'Hara 
Creek,  and  on  Gardiner  River,  near  Mammoth  Hot  Springs. 

Lepargyrea  argentea  (Nuttall)  Greene.    Buffaloberry. 

While  abundant  locally  on  many  streams  at  the  base  of  the  mountains,  espe- 
cially at  the  north,  the  buffaloberry  is  by  no  means  generally  distributed  over 
the  State,  and  was  not  detected  in  the  southwest.     The  localities  where  it  was 


Mountain   balm    iCeanothux   velutinus)    in   flower 
(9,000  feet). 


B I 9636 

Siena   Madre,  July   7,   1911 


found  are  mainly  Upper  Sonoran  :  Sibylee  Creek,  Casper,  Belle  Fourche  River, 
Little  Missouri  River,  Powder  River,  Clear  Creek,  streams  near  Sheridan  and 
Ranchester,  up  Wolf  Creek  to  4,500  feet  elevation,  Alcova,  Casper  Creek,  north 
slope  Rattlesnake  Mountains,  Snake  River  at  Baggs,  Wind  River  Basin  (mainly 
below  6,000  feet),  and  streams  of  Bighorn  Basin. 

Lepargyrea  canadensis  (Linnaeus)  Green.  Canadian  Buffaloberry.  (Fig.  15.) 
The  Canadian  buffaloberry  is  a  characteristic  and  widely  distributed  under- 
shrub  in  the  forests  of  the  Canadian  Zone  between  8,000  and  10,000  feet  eleva- 
tion. Although  usually  found  on  mountain  slopes,  it  is  abundant  in  balsam 
poplar  growth  along  Snake  River  and  other  streams  of  Jackson  Hole. 

Phyllodoce  empetriformis  (Smith)  Don.     Mountain  Heath. 

The  mountain  heath,  a  characteristic  evergreen  of  the  Hudsonian  Zone,  was 
found  locally  abundant  at  widely  separated  localities  in  the  high  ranges  of 


1917.] 


TREES   AXD   SHRUBS. 


75 


northwestern  Wyoming.  Near  Fremont  Peak  it  was  abundant  in  bogs  and 
rocky  situations  from  10,500  feet  altitude  to  a  little  above  timberline,  and 
occasional  at  10,000  feet,  on  July  20  the  dainty  rose-colored  flowers  were 
still  open  on  the  Wind  River  Mountains,  bid  on  August  1  it  was  nearly  through 
flowering  on  Whirlwind  Peak,  in  tbe  Absaroka  Range.  At  timberline  on  the 
Tetons  no  flowers  remained  on  the  branches  by  August  30. 

Menziesia  ferruginea  Smith.     Rustyleaf. 

This    handsome    and    conspicuous    shrub    was    observed    only    in    the    dense 
coniferous  forests  of  Yellowstone  Park,  where  it  is  common  near  Sylvan  Pass, 


1'ic.    15. — Canadian    buffaloberry    {Lepargyrea    canadensis) 
Range  near  Merna   C8.000  feet). 


B 1 3685 

aspen    woods,    Wyoming 


Loin'  Star  Geyser,  below  Norris  Basin,  Thumb,  Lewis  ttiver,  and  Snake  River 
near  Soldier  Station. 

Kalmia  polifolia  Wangenheim.     Laurel. 

The  wet  bogs  and  mossy  margins  of  the  numerous  snow-fed  lakes  on  the 
Wind  River  Range  south  of  Fremont  Peak  were  bright  with  pretty  pink- 
purple  flowering  mats  of  laurel  in  mid-July,  1911.  The  species  was  in  flower 
on  July  18  in  the  Hudsonian  Zone  from  10,000  to  11,000  feet  elevation,  but  was 
not  detected  above  timberline.  With  Phyllodoce  em  pet  ri  for  mis  it  often  formed 
dense  mats,  from  a  few  inches  to  nearly  a  foot  in  height.  Vernon  Bailey  col- 
lected the  laurel  in  the  timberline  region  on  the  Wind  River  Mountains  above 
Bull  Lake  in  1893,  and  at  the  head  of  Raven  Creek,  in  Yellowstone  Park, 
in  1915. 


76  NORTH    AMERICAN    FAUNA.  [No.  42. 

Arctostaphylos  uva-ursi    (Linnaeus)    Sprengel.     Red   Bearberxy. 

The  low  trailing  red  bearberry,  with  its  leathery  evergreen  leaves,  dainty 
rose-hued  blossoms,  and  red  berries,  forms  beautiful  mats  in  the  open  yellow- 
pine  forests  of  the  Black  Hills,  Laramie  Peak  region,  and  on  the  eastern 
slopes  of  the  Bighorn  Mountains.  While  most  abundant  in  the  Transition  Zone. 
it  occurs  occasionally  in  the  coniferous  forests  higher  up,  as  on  the  upper 
slopes  of  Laramie  Peak  and  on  the  Wind  River  Range  south  of  Dubois.  It  is 
very  abundant  in  the  woodlands  of  Jackson  Hole,  and  also  on  the  Bear  Lodge 
Mountains ;  at  from  5,000  to  8,000  feet  elevation  on  the  eastern  slope  of  the 
Bighorn  Mountains  near  Wolf ;  and  from  8,000  to  9,500  feet  on  their  western 
slope  above  Hyattville;  on  the  Rattlesnake,  Shirley,  and  Green  Mountains; 
south  of  Fremont  Peak  and  on  the  head  of  the  Big  Sandy,  Wind  River  Range : 
Needle  Mountain  and  Grinned  Creek,  £bsaroka  Range;  Merna;  Evanston ;  and 
north  slopes  of  Uinta  Mountains  west  of  Lonetree. 

Vaccinium  oreophilum  Rydberg.     Mountain  Blueberry. 

The  mountain  blueberry  reaches  perfection  of  growth  in  the  cool,  damp 
forests  of  the  Canadian  Zone  at  about  8,000  feet  elevation.  In  many  localities 
it  occurs  with  Vaccinium  erythrococcum,  but  it  is  less  abundant  and  usually 
not  found  so  high  up.  It  was  observed  oh  the  Bear  Lodge,  Shirley,  Casper, 
Ferris,  and  Bighorn  Mountains ;  Grinned  Creek,  Absaroka  Range ;  forests  of 
Yellowstone  Park;  Teton  Pass  and  Moose  Creek,  Teton  Range;  wooded  hills 
bordering  Jackson  Lake;  head  of  Pacific  Creek;  Snake  River  west  of  Jackson; 
Salt  River  Range  near  Afton ;  and  on  the  Wyoming  Range  near  Merna. 

Vaccinium  erythrococcum  Rydberg.     Red  Bilberry.  (PI.  IX,  fig.  2.) 

The  low,  small-leaved  red  bilberry  was  noted  on  all  the  higher  elevations 
of  Wyoming.  It  is  most  abundant  in  the  upper  Canadian  Zone,  where  it 
densely  carpets  the  coniferous  forest  floor,  and  it  is  the  only  Vaccinium  present 
in  the  Hudsonian  belt  of  dwarfed  forest  higher  up.  In  early  August  the  small 
juicy  red  berries  are  greatly  relished  by  dusky  grouse  and  other  fovest  birds. 

Vaccinium  occidentale  Gray.     Western  Blueberry. 

The  shrubby  western  blueberry  was  encountered  in  northwestern  Wyoming, 
from  Lewis  Lake,  Yellowstone  Park,  south  to  Jackson  Lake,  and  east  on  the 
Wind  River  Range  to  Fremont  Peak.  It  attains  a  height  of  nearly  2  feet  in 
Yellowstone  Park,  growing  in  dense  clumps  in  lodgepole  pine  forests  between 
7,000  and  8,000  feet  elevation.  Its  small  blue  berries  were  ripe  south  of  Lewis 
Lake  on  August  18,  1910.  South  of  Fremont  Peak  it  was  growing  in  bogs  a 
little  below  timberline,  at  10,500  feet. 

Fraxinus  lanceolata  Borekhausen.     Green  Ash. 

The  green  ash  occurs  sparingly  in  the  low  stream  valleys  of  extreme  eastern 
and  northern  Wyoming  west  to  the  base  of  the  Bighorn  Mountains.  It  was 
found  at  Newcastle;  Sand  Creek  to  10  miles  above  Beulah;  Hay  Creek  to 
Aladdin  and  Eothen;  groves  on  the  Belle  Fourche  flats  at  Devils  Tower;  Little 
Missouri  River;  gulches  near  Morse;  Powder  River  near  Montana  boundary; 
Prairie-dog  Creek;  Sheridan;  Big  Goose  Creek;  Wolf  Creek,  to  4,500  feet  eleva- 
tion, near  Eaton's  Ranch. 

Sambucus  microbotrys  Rydberg.     Red  Elderberry.  (Fig.  16.) 

The  red  elderberry  is  a  characteristic  shrub  of  the  Canadian  Zone  coniferous 

forest  on  the  main  ranges  of  Wyoming.     Usually  it  is  the  only  elderberry  present 

on  the  higher  slopes,  but  lower  down   occurs  with  its  black-fruited  relative, 


l'.KT.I 


TREES    AND    SHRUBS. 


77 


Sambucus  melanocarpa.  The  red  elder  was  not  taken  in  the  foothill  ranges  of 
eastern  Wyoming  nor  in  the  Bighorn  Mountains,  but  it  has  a  general  distribu- 
tion from  7,0(10  to  9,."i00  feet  elevation  in  the  mountains  farther  west. 

Sambucus  melanocarpa  (J ray.     Black  Elderberry. 

The  black  elderberry  is  of  lower  vertical  range  than  the  red  species,  occurring 
mainly  in  the  Transition  Zone,  and  from  our  observations  was  not  so  generally 
distributed.  It  was  noted  on  Laramie  Peak,  north  slope,  rare  at  8,000  feet 
elevation;  Shirley  Mountains,  north  escarpment,  7,600  feet  to  summit;  Ferris 
Mountains,  in  dense  coniferous  forest,  8,000  to  9,000  feet ;  Salt  River  Range, 
7.000  feet.  Vernon  Bailey  found  it  bearing  ripe  berries  on  August  10,  at  7,500 
feet,  on  a  southwest  slope  in  the  Bighorn  Mountains  near  Wolf. 


Pig.  10. — Red  elderberry   (Sambucus  microbotrys)    in  bloom,  Sierra  Madie,  July  7,   1911 

(10,000  feet). 

Sambucus  canadensis  Linnaeus.     Elderberry. 

The  large  elderberry,  Sambucus  canadensis,  was  detected  only  at  the  southern 
base  of  the  Bear  Lodge  Mountains  near  Sundance,  in  the  Transition  Zone.  Its 
handsome  flat-topped  cymes  of  white  flowers  were  very  conspicuous  in  the 
canyon  along  Sundance  Creek,  at  4,700  feet  elevation,  on  June  20,  1912. 

Viburnum  lentago  Linnaeus.     Sweet  Viburnum. 

Vernon  Bailey  found  the  sweet  viburnum  not  uncommon  in  August,  1913.  in 
Hie  shaded  canyons  and  gulches  along  Sand  Creek  above  Beulah,  at  the  northern 
base  of  the  Black  Hills.  He  also  reports  it  from  the  Belle  Fourche  Valley  near 
Devils  Tower,  and  along  Big  Goose  Creek  at  the  cast  base  of  the  Bighorn 
.Mountains, 


78  NORTH   AMERICAN    FAUNA.  [No.  42. 

Symphoricarpos  occidentalis  Hooker.     Wolfberry. 

The  wolfberry  occurs  over  much  of  the  Upper  Sonoran  Great  Plains  area, 
forming  dense  thickets  2  feet  in  height  in  gulches  and  along  si  reams,  and  in 
many  localities  constituting  the  principal  shrubby  growth.  The  species  regu- 
larly extends  west  to  the  bases  of  the  Laramie  and  Bighorn  Mountains,  and 
was  found  also  in  Wind  River  Basin,  and  at  the  eastern  base  of  the  Absaroka 
Range  northwest  of  Cody. 

Symphoricarpos  pauciflorus   (Robbins)   Britton.     Few-Flowered  Snowberry. 

The  few-flowered  snowberry  is  higher  ranging  than  the  wolfberry  (Symphori 
earpos  occidentalis)  and  occurs  mainly  in  the  Transition  Zone  in  northern 
Wyoming.  It  was  recorded  at  various  altitudes  from  4,700  feet  to  8,500  feet 
in  the  Bear  Lodge,  Bighorn,  Absaroka,  and  Wyoming  Ranges. 

Symphoricarpos  oreophilus  Gray.     Mountain  Snowberry. 

The  mountain  snowberry  is  the  predominating  species  of  its  genus  in  Wyoming, 
where  it  is  rarely  absent  from  dry,  rocky,  open,  or  partly  forested  slopes  and 
ridges  in  the  Transition  Zone.  On  some  of  the  southern  and  western  ranges  it 
is  common  on  exposed  points  in  dry  forest  openings  up  to  9,000  feet  elevation. 
This  shrub  enters  the  lower  portions  of  Yellowstone  Park  at  the  north  and 
south,  and  occurs  in  the  open  sections  of  Jackson  Hole. 

Symphoricarpos  rotundifolius  Gray.     Round-Leaved  Snowberry. 

The  round-leaved  snowberry  is  apparently  uncommon.  It  is  represented  by  a 
specimen  from  Rawhide  Butte,  taken  July  23,  1000,  where  scattering  bushes 
were  noted  from  base  to  summit  in  the  Transition  Zone. 

Lonicera  glaucescens  Rydberg.    Douglas  Honeysuckle. 

The  Douglas  honeysuckle  reaches  Wyoming  in  the  extreme  northeast,  where 
it  is  found  sparingly  along  streams  and  on  damp  slopes  above  4,700  feet  eleva- 
tion near  Sundance,  in  the  Bear  Lodge  Mountains. 

Lonicera  utahensis  Watson.     Utah  Honeysuckle. 

The  Utah  honeysuckle  abounds  in  the  mountain  forests  of  extreme  western 
and  northwestern  Wyoming.  From  the  forests  of  Yellowstone  Park,  where  it 
is  especially  abundant,  it  occurs  south  at  least  to  the  Salt  River  Mountains, 
reaching  its  upper  limits  on  the  western  slope  of  this  range  at  9,000  feet  eleva 
tion,  and  on  the  eastern  slope  of  the  Teton  Range  at  about  8,000  feet. 

Lonicera  involucrata  Banks.    Involucred  Fly  Honeysuckle. 

In  Wyoming  the  involucred  fly  honeysuckle  occurs  commonly  in  all  the  prin- 
cipal ranges  with  the  possible  exception  of  the  Bighorn  Mountains.  It  becomes 
increasingly  abundant  westward,  and  is  one  of  the  most  characteristic  shrubs 
on  the  banks  of  cold  streams  up  to  8.000  or  9.000  feet  elevation  in  the  Canadian 
forest  belt,  ripening  its  blackish  berries  about  the  first  of  August. 

Chrysothamnus  graveolens  (Nuttall)  Greene.    Rabbit  Brush. 

The  plains,  deserts,  and  basal  mountain  slopes  of  Wyoming  are  rich  in  species 
of  Chrysothamnus,  some  of  them  among  the  best  characterizing  shrubs  of  the 
Upper  Sonoran  or  Transition  Zones.  The  meager  data  secured  by  Survey  par- 
ties relates  mainly  to  species  which  are  of  importance  in  determining  life  zones. 
€.  graveolens  is  a  conspicuous  shrub,  often  several  feet  high,  growing  in  Upper 
Sonoran  gulches,  desert  arroyos,  and  on  dry  slopes,  principally  in  eastern  and 
northern  Wyoming.  It  is  especially  abundant  on  flats  along  Bighorn  River  iv 
the  lower  portions  of  the  Bighorn  Basin.  The  species  occurs  at  Cassa  ;  Fori 
Fetterman;  Casper;  along  the  North  Platte  near  Alcova ;  Hay  Creek  east  of 


1917.]  TEEES   AND   SHRUBS.  79 

Aladdin;  Wind  < 'reek  northeast  of  Moorcroft;  Buffalo;  gulches  ;it  south  base  <>f 
Owl  Creek  Mountains;  Badwater  River  north  of  Shoshoni;  Worland;  Mander- 
son ;  Greybull ;  Frannie  to  Garland ;  and  east  of  Cody. 

Chrysothamnus  linifolius  Greene.     Rabbit  Brush. 

Like  Chrysothamnus  graveolcns,  this  species  is  a  fair-sized  shrub,  inhabiting 
principally  the  cut  banks  of  adobe  along  desert  washes  and  streams  in  the 
Upper  Sonoran  Zone,  as  along  Bear  Creek,  south  of  Cassa;  Rock  Springs;  and 
warm  pockets  along  Green  River  between  Fontenelle  and  Labarge.  Vernon 
Bailey  collected  a  specimen  at  Arvada,  on  Powder  River,  August  13,  1913. 
Nelson  found  the  species  at  Point  of  Rocks  and  Granger.1 

Chrysothamnus  plattensis  Greene.     Rabbit  Brush. 

Chrysothamnus  plattensis  is  a  low,  spreading  Upper  Sonoran  species  of  rabbit 
brush  of  the  North  Platte  and  tributary  valleys  at  the  southeast.  It  was  noted 
up  to  6,200  feet  elevation  on  Horse  Creek,  near  Meadow,  and  also  at  Snow's 
Ranch,  on  Rawhide  Creek  below  Patrick. 

Chrysothamnus  stenophyllus  Greene.     Rabbit  Brush. 

This  is  a  common  species  of  rabbit  brush  over  most  of  the  sandy  plains  and 
ridges  of  central  Wyoming,  in  the  Upper  Sonoran  Zone,  as  al  Casper,  Bfell,  and 
in  Sweetwater  Valley  from  Independence  to  Splitrock.  Nelson  records  it  from 
Point  of  Rocks,  on  the  Red  Desert.1 

Chrysothamnus  stenophyllus  Greene.     Rabbit  Brush. 

This  species  was  taken  in  the  Upper  Sonoran  Zone  in  the  Belle  Fourche 
Valley  at  Moorcroft,  August  15,  1913,  by  Vernon  Bailey. 

Chrysothamnus  howardi   (Parry)    Greene.     Rabbit  Brush. 

A  specimen  of  Chrysothamnus  howardi  was  collected  by  Vernon  Bailey  at 
Arvada,  on  Powder  River,  August  13,  1913.     It  is  usually  of  the  higher  plains. 

Chrysothamnus  wyomingensis  A.  Nelson.     Rabbit  Brush. 

Mainly  of  the  high  Transition  plains  and  basal  mountain  slopes  at  the  north- 
It  is  very  abundant  at  the  northern  base  of  the  Bighorn  Mountains  at  Wolf, 
where  a  specimen  was  collected  on  June  6,  1912,  and  is  apparently  the  species 
so  abundant  on  the  basal  slopes  of  the  Bear  Lodge  Mountains.  Vernon  Bailey 
collected  it  at  Arvada,  on  Powder  River. 

Chrysothamnus  pulcherrimus  A.  Nelson.     Rabbit  Brush. 

This  rabbit  brush  is  common  in  the  Transition  Zone  on  the  high  central 
plains  and  in  the  upper  Green  River  Basin,  growing  on  open  slopes  with  sage- 
brush regularly  to  8.000  feet  altitude.  It  is  especially  abundant  on  both  slopes 
of  the  mountains  near  Laramie  Peak,  on  the  northern  Laramie  Plains,  and 
in  Shirley  Basin.  It  was  noted  along  Little  Medicine  Bow  River  west  of  Mar- 
shall at  7,000  feet. 

Artemisia  tridentata  Nuttall.     Black  Sagebrush.  (Fig.  17.) 

The  most  widely  distributed  shrub  in  Wyoming  is  the  black  sagebrush.  It 
is  omnipresent  in  open  country  east  to  the  edge  of  the  Great  Plains,  where  its 
eastern  limits  are  marked  by  Grin  Junction  (North  Platte  Valley),  Lost  Spring. 
Indian  Creek  north  of  Kirtley  at  State  line,  Clifton,  Newcastle,  Wind  Creek 
northeast  of  Moorcroft,  and  Colony.  It  was  not  found  east  of  the  Laramie 
Mountains  in  southeast  Wyoming.     From  4.000  feet   elevation  at  the  east  this 

1  Bull.  13,  Div.  <>f  Agrost.,  U.  S.  Dopt.  Agr.,  p.  66,  1898. 


80 


NORTH   AMERICAN    FAUNA. 


[No.  42. 


sagebrush  extends  to  timberline  on  some  of  the  ranges,  attaining  9.800  feet 
on  the  Bighorns  above  Hyattville,  and  10,000  feet  on  the  Wind  Rivers  south 
of  Dubois  and  on  the  Wyoming  Range  west  of  Bigpiney  and  Merna. 

Artemisia  cana  Pursh.     Gray  Sagebrush. 

The  gray  sagebrush  is  generally  dispersed  and  especially  common  in  the 
mountain  valleys  and  parks,  but  less  abundant  than  the  black  species.  It  ex- 
tends into  the  edge  of  the  Great  Plains  area.  In  the  valleys  east  of  the  Laramie 
Mountains  this  is  usually  the  only  shrubby  sage  present,  and  it  forms  much 
of  the  scattering  growth  on  the  open  stretches  between  the  Black  Hills  and 
Bighorn  Mountains.     Westward  it  rapidly  gives  way  to  the  black  sagebrush. 


Fig.  17. — Black  sagebrush  (Artemisia  tridentata)   10  feet  high,  upper  Wind  River  Valley. 


Artemisia  trifida  Nuttall.     Sagebrush. 

A  small  shrubby  sage,  sparingly  present  in  open  Transition  country  chiefly 
at  the  west,  as  on  the  summit  of  the  Owl  Creek  Mountains;  Bear  River  Divide 
north  of  Evanston,  7,500  feet  elevation;  benches  above  Cokeville;  and  Salt 
River  Valley.  Vernon  Bailey  found  the  species  in  open  parks  in  the  mountains 
south  of  Douglas,  near  Valley,  and  in  Jackson  Hole. 

Artemisia  fllifolia  Torrey.     Narrow-Leaved  Sagebrush. 

The  narrow-leaved  sagebrush  is  a  characteristic  shrub  of  the  Great  Plains 
Sonoran  area,  occurring  in  the  North  Platte  drainage  chiefly  in  sandy  tracts  as 
far  west  as  Guernsey.  A  very  dense  growth  2  or  3  feet  high  covers  the  Rawhide 
flats  at  Snow's  Ranch  south  of  Patrick,  and  it  is  abundant  in  the  sandy  valley 
of  the  Chugwater  east  of  Wheatland, 


1917.]  TREES    AND    SHRUBS.  81 

Artemisia  arbuscula  Nuttall.     Brown  Sagebrush. 

The  brown  sagebrush  is  uncommon,  according  to  the  author's  observations. 
Vernon  Bailey  reports  it  at  Hams  Fork  Station  (Moyer  Junction)  and  Valley, 
and  collected  specimens  on  the  Owl  Creek  Mountains  in  1S93  and  in  Salt  River 
Valley  on  August  15,  1911,  all  Transition  Zone  localities  in  western  Wyoming. 

Artemisia  spinescens  Eaton.     Spiny  Sage ;  Bud  Brush. 

A  low  spinescent  shrubby  sage,  abundant  at  many  localities  in  the  Upper 
Sonoran  desert  region  at  the  southwest.  It  is  common,  also,  on  sandy  benches 
near  Shoshoni  and  in  the  lower  parts  of  the  Bighorn  Valley  near  Greybull  and 
Manderson.  It  occurs  at  many  stations  with  the  less  shrubby  Artemisia 
ijedati/ida,  with  which  it  is  sometimes  confused. 

Artemisia  frigida  Willdenow.    Pasture  Sage. 

The  pasture  sage,  a  low  silvery  sage,  is  scarcely  a  shrub,  but  is  included  as 
a  characteristic  Transition  species  of  the  genus.  It  has  a  very  wide  range 
over  Wyoming  on  high  plains,  bare  ridges  and  plateaus,  and  open  mountain 
slopes  generally. 

Tetradymia  spinosa  Hooker  and  Arnott.    Spiny  Rabbit  Brush. 

The  members  of  the  genus  Tetradymia  are  characteristic  shrubs  of  the  Great 
Basin  Division  of  the  Upper  Sonoran  Zone,  and  do  not  enter  the  Great  Plains 
area.  They  occur  in  varying  abundance  and  scattered  bunchlike  growths  over 
the  dry  hills  and  plains  of  the  arid  central  desert  section,  from  the  Green  River 
Valley  and  Red  Desert  north  to  the  Bighorn  Basin. 

Tetradymia  spinosa  is  the  more  widely  distributed  species  below  6,500  feet 
altitude.  It  is  common  at  Green  River,  Superior,  Rock  Springs  region,  warm 
river  flats  (Fontenelle  to  Labarge),  sand  flats  north  of  Opal,  Shoshoni,  Wor- 
land,  Bonanza,  Manderson,  Greybull  to  Cody,  and  Frannie  to  Garland. 

Tetradymia  nuttalli  Torrey  and  Gray.    Nuttall  Rabbit  Brush. 

The  Nuttall  rabbit  brush  is  more  spiny  than  the  preceding  species.  It  was 
collected  only  at  Fort  Steele,  where  it  was  abundant  in  the  warm,  lower  parts 
of  the  North  Platte  Valley.  Nelson  mentions  specimens  from  Bitter  Creek 
and  Green  River.1 

Tetradymia  inermis  Nuttall.     Rabbit  Brush.  (PI.  VII,  fig.  2.) 

This  high-ranging  rabbit  brush  grows  abundantly  on  the  sandy  plains  of 
the  upper  Green  River  Basin  to  an  elevation  of  7,500  feet.  It  was  not  found 
at  the  lower  levels,  but  was  common  on  sand  flats  at  Eden  and  Big  Sandy  and 
also  on  Little  Piney  Creek.  Nelson  collected  specimens  at  Bitter  Creek  on  the 
Red  Desert.' 


1  Bull.  13,  Div.  of  Agrost.,  U.  S.  Dept.  Agr.,  p.  67,  1898. 
74440°— 17 Q 


INDEX 


[Pages  containing  the  principal  reference  to  a  species  in  bold-faced  figures.] 


A. 


Abies  (concolor?),  44,  58. 

lasiocarpa,  49,  58. 
Abronia  elliptica,  16,  28. 

fragrans,  28. 
Accipiter  cooperi,  35. 

velox,  35. 
Acer  glabrum,  36,  T3. 

grandidentatuni,  36,  73. 

negundo,  28,  73. 
Achillea  alpicola,  52. 
Acknowledgments,  8    (footnote). 
Aconitum  columbianum,  45. 
Actsea  arguta,  45. 

rubra,  45. 
Adenostegia  ramosa,  37. 
Aeronautes  melanoleucus,  35. 
Agelaius  phceniceus  fortis,  26. 
Agriculture,  Arctic-Alpine  Zone.     See  Boreal 
Zones. 

Boreal  Zones,  52-54. 

Canadian    Zone,    42.      See   also   Boreal 
Zones. 

Hudsonian  Zone.     See  Boreal  Zones. 

Transition  Zone,  37-38. 

Upper  Sonoran  Zone,  30-31. 
Great  Basin  Division,  18. 
Bighorn  Basin,  24. 
Green  River  Valley,  19. 
Red  Desert,  21. 
Wind  River  Basin,  22-23. 
Great  Plains  Division,  17. 
Agropyron  caninum,  37. 

pseudorepens,  37. 

smithi,  30. 

spicatum,  30,  37. 

violaceum,  46. 
Agrostis  hiemalis,  37. 
Alces  americanus  shirasi,  42. 
Alder,  44,  64. 
Alnus  tenuifolia,  44,  64. 
Alopecurus  occidentalis,  46. 
Alsinopsis  ohtusiloba,  52. 
Alum   root,   45. 
Ambystoma  tigrinum,  27,  33. 
Amelanchier  alnifolia,  36,  71. 

elliptica,    36, 

oreophila,   36. 
Ammodramus  savannarum  bimaculatus,  26. 
Amorpha  canescens,  28,  72. 

nana,    28. 
Amphibian  lists,  19,  27,  33. 
Amphispiza  nevadensis,  27. 


Anaphalis  suhalpina,  4."i. 
Anas  platyrhynchos,  34. 
Andropogon  halli,  29. 

scoparius,  29. 
Androsace  carinata,  52. 

subumbellata,  49. 
Anemone,  37,  45,  49. 
Anemone  cylindrica,  37. 

globosa,  49. 

lithophila,   45. 

tetonensis,  49, 
Angelica   roseana,  49. 
Anogra  albicaulis,  16,  29. 
Antelope,    25. 
Antennaria  pulcherrima,  45. 

reflexa,  37,  50. 
Anthus  rubescens,   51. 
Antilocapra  americana  americana,  25. 
Apocynum  adrosa^mifolium,  37. 
Aquilegia  cserulea,  45. 

flavescens,  45. 

oreophila,  45. 

saximontana,  49. 
Arabia  lyalli,  52. 
Aragallus  lagopus,  52. 

lamberti,  37. 

nanus.  52. 
Aralia  nudicaulis,   37. 
Archibuteo  ferrugineus,  26. 
Arctostaphylos  uva-ursi,  36,  76. 
Ardea  herodias,  26. 
Arenaria   congesta,   36. 
Argemone  hispida,  28. 

intermedia,    16,    28. 
Aristida    longiseta,    29. 
Arnica,  heart-leaved.  45. 
Arnica  cordifolia,  45. 
Artemisia  arbuscula,  36,  81. 

cana,  36,  80. 

filifolia,  28,  80. 

frigida,  36,  81. 

ludoviciana,  36. 

pedatifida,  28,  81. 

scopulorum,  52. 

spinescens,  28,  81. 

tridentata,  28,  36,  79-SO 

trifida,  36.  80. 
Asclepias  pumila,  29. 

speciosa,  29. 
Ash,  16,  28. 

green,  76. 

mountain,  44,  71. 
Asio  wilsonianus,  35. 
Aspen,  39,  40,  41,  42,  44,  60. 

83 


84 


NORTH    AMERICAN    FAUNA. 


[No.  4: 


Astragalus  alpinus,  45. 

calycosus,  37. 

crassicarpus,   16,  29. 

drummondi,  37. 

flexuosus,  37. 

hypoglottis,  37. 

missouriensis,  29. 

mollissimus,  16,  29. 

nitidus.  37. 

succnlentus,  37. 
Asyndesmus  lewisi,  35. 
Atheropogon  curtipendula,  30. 
Atriplex  argentea,  28. 

canescens,  28,  65. 

confertifolia,  28,  65-66. 

nuttalli,  28,  66. 

pabularis,  28. 
Avens,  49. 

alpine,  51,  52,  69. 

mountain,  45,  52. 
Avocet,  34. 

B. 

Badger,  26,  34. 

Baeolophus  inornatus  griseus,  27. 
Balm,  mountain,  33,  36,  73. 
Balsamorrhiza,  40. 
incana,  37. 
sagittata,  37. 
Balsam  root,  37. 
Baneberry,  45. 
Barberry.  33,  36,  67. 
Bartramia   longicauda,  26. 
Bascanion  constrictor,  27. 
Bat,  big-eared.   :'.4. 
brown,  24,  26. 
little,  43. 
California,  16,  22,  24,  26. 
boary,  43. 
long-eared,  20,  26. 
long-legged,  26,  33,  34. 
silver-haired,  43. 
Bean,  buffalo,  29. 
Bear,  black,  43. 
grizzly,  34. 

Washakie,  43. 
Yellowstone  Park,  43. 
Bearberry.  33,  36. 

red,  76. 
Beard-tongue,  29,  37,  45,  49. 
Beaver,  34,  43. 
Berberis  aquifolium,  36,  67. 
Besseya  alpina,  52. 
Betula  fontinalis,  36,  63-64. 
glandulosa,  44,  64. 
papyrifera,  44,  63. 
Bilberry,  red.  45.  76. 
Birch,  canoe,  40,  42,  44,  63. 
dwarf,  44,  64. 

Rocky  Mountain,  33,  36,  03-64. 
Bird  lists,  16,  19,  20-21,  22,  24,  26-27, 

34-35,  43-44,  49,  51. 
Bitter  root,  36. 
alpine,  49. 
Bittern,  26. 
Blackbird,  Brewer,  35. 
yellow-headed,  26. 


33, 


Bladder-pod,  double,  37. 
Blazing  star,  29. 
Blite,  sea,  28. 

shrubby,  28. 
Blueberry,  45. 

mountain,  44,  76. 
western,  76. 
Bluebird,  mountain,  35,  44. 
Bluegrass,  50,  52. 
alpine,  52. 
arctic,  52. 
mountain,  46. 
Bluestem,  29. 
Bobolink,  35. 

Botaurus  lentiginosis,  26. 
Bouteloua  hirsuta,  30. 

oligostachya,  30. 
Boykinia  heucheriformis,  52. 
Branta  canadensis,  43. 
Bromus  marginatus,  37. 

porteri,  37. 
Brush,  antelope,  69. 
bud,  81. 

rabbit,    16,    17,    19,   20,   28,   36,   78-79, 
79,  81. 

Nuttall,  81. 
spiny,  81. 
Buckthorn,  44,  73. 
Budbrush,  28. 
Buffaloberry,  22,  28,  74. 

Canadian,  44,  74. 
Buffle-head,   43. 
Bufo  boreas,  33. 
cognatus,  27. 

lentiginosis  woodhousei,  27,  33. 
Bulbilis  dactyloides,  30. 
Bunting,   lark,   16,   27. 
lazuli,   16,   24,   27. 
Bnpleurum  americanum,  52. 
Bur,  buffalo,  29. 
Bush,  skunk,  28. 
Bush-tit,   lead-colored,   27. 
Buteo  swainsoni,  35. 
Buttercup,  48,  49,  52. 
Nuttall,  37. 


C. 


Cactus,  18. 
ball,  29. 

Simpson,  37. 
prickly-pear,  20,  22,  23. 
small-jointed,  37. 
Calamagrostis  canadensis  acuminata,  46. 
Calamospiza    melanocorys,    27. 
Calamovilfa  longifolia,  29. 
Calandrinia  pygmsea.  49. 
Calcarius  ornatus,   35. 
Calochortus  gunnisoni,  36. 
nuttalli,   36. 
pavonaceus  45. 
Callospermophilus  lateralis  caryi,  43,  48,  51. 
castanurus,  43. 
lateralis,  34,  43. 
wortmani,  25,  34. 
Caltha  leptosepala,  49. 
Calypso  bulbosa,  45. 


1917.] 


INDEX. 


85 


Camas,  beautiful,  45. 

poison,  36. 
Campanula  parryi,  49. 

rotundifolia,  37. 

uniflora,   52. 
Canis  lestes,  34,  48,  51. 

nebracensis,   26. 

nubilus,  34. 
Canyon  mouse.     See  Mouse. 
Carcluus  plattensis,  16,  29. 
Carex  albo-nigra,  52. 

atrata,  52. 

engelmanni,  52. 

nigricans,  50. 

nova,   50. 

nubicola,  52. 

phseocephala,  52. 
Carpodacus   cassini,   44. 

mcxicanus  frontalis,  26. 
Castilleja,  49. 

sulphurea,  45. 
Castor  canadensis,  34,  43. 
Catbird,  16,  24,  27. 
Catchfly,   stemless,  52. 

Catoptrophorus  semipalmatus  inornatus,  34. 
Ceanotbus  fendleri,  36,  73. 

mollissimus,  36,  73. 

velutinus,  36,  73. 
Centrocercus  urophasianus.  35. 
Cerastium  beeringianum,  49. 
Cercocarpus  intricatus,  36,  68. 

ledifolius,  36,  68. 

parvifolius,  28,  36,  68-69. 
Certbia  familiaris  montana.  49. 
Cervus  canadensis  canadensis,  34,  42,  51. 
Chaenactis   alpina,   50. 

douglassi,  37. 
Charitonetta  albeola,  43. 
Cbat,  long-tailed,   16,  22,  24,  27. 
Chaulelasmus  streperus,   34. 
Cberry,  ground,  29. 

red,  wild,  36,  72. 

sand,  16,  28. 

wild,   33. 
Chickadee,  long-tailed,  35. 

mountain,  44. 
Chickweed,  mouse-ear,  49. 
Chimaphila  umbellata,  45. 
Chipmunk,  buff-bellied,  43. 

•Colorado,   42. 

least,  19,  20,  25. 

northern,   42. 

pale,  22,  24,  25. 

rock,   Utah,   19,   25. 

sagebrush,  25. 

timberline,  48,  51. 

Uinta,  43. 
Chokecherry,  36,  72. 
Chondestes  grammacus  strigatus,  26. 
Chordeiles  virginianus  henryi,  26. 
Chorophilus  triseriatus,  33. 
Chrysothamnus  frigidus,  36. 

graveolens,  28,  78-79. 

howardi,  79. 

lanceolatus,  79. 

linifolius,  28,  79. 

parryi,  36. 


Chrysothamnus  plattensis,  28,  79. 

nulcherrimus,  36,  79. 

stenophyllus,  28,  79. 

wyomingensis,  36,  79. 
Cinclus  mexicanus  unicolor,  44. 
Cinquefoil,  37,  49. 

glandular,  37. 

shrubby,  40,  44,  47,  51,  71. 
Citellus  armatus,  34,  51. 

obsoletus,  25. 

ricbardsoni  elegans,  34. 

tridecemlineatus  pallidus,  2.~>. 
parvus,  25. 
Clammy-weed,  28. 
Clangula  islandica,  43. 
Claytonia  megarrhiza,  52. 

rosea,  45,  49. 
Clematis  douglasi,  36. 

ligusticifolia,  36. 

occidentalis,  36. 

pseudalpina,  45. 
Clementsia  rhodantba,  49. 
Cleome.  red,  28. 

yellow,  18,  19,  20,  28. 
Cleomelutea,  23.  28. 

serrulata,  16.  28. 
Cliff  mouse.     See  Mouse. 
Climate,  11-12. 
Clover,  45. 

dwarf,  49. 

Parry,  49. 

prairie,  29. 

purple,  29. 
silky,  29. 
white,  29. 
Cnemidophorus  sexlineatus,  27. 
Coccyzus  erythropthalmus,  35. 
Columbine,  45.  48,  49. 

blue,  45. 

white,  47. 

yellow,  45. 
Colymbus  nigricollis  californicus,  34. 
Cone  flower,  29. 
Coney,  48,  51. 
Coot,  26. 

Corallorhiza  multiflora,  36. 
Coral  root,  36. 
Cornel.  33,  36. 
Cornus  instolonea,  36. 

stolonifera,  36. 
Corvus  brachyrhynchos  hesperis,  26. 
Corylus  rostrata,  44,  63. 
Corynorhinus  macrotis  pallescens,  34. 
Cottontail,  Bailey,  16,  20,  22,  24,  26. 

Black  Hills.   33,   34. 

Nebraska,   26. 
Cottonwood,      broad-leaved,      15,      22,      28, 
60-61. 

lance-leaved,  28,  61. 

narrow-leaved,  33,  36,  61. 
Coyote,  51. 

mountain,  34,  48. 

plains,    26. 
Cowslip,  mountain,  48,  49. 
Crataegus  cerronis,  36,  71. 

rivularis,  36,  71. 

sheridana,  36. 


86 


NORTH   AMERICAN   FAUNA. 


[No.  42. 


Creeper,  Rocky  Mountain,  49. 

Virginia,  28. 
Cross,  Whitlow,  52. 
Crossbill,  44. 
Crotalus  confluentus,  27. 
Croton,  29. 

texensis,  16,  29. 
Crow,  western,  26. 
Crowfoot,  45. 
Cryptoglaux  acadica,  35. 
Cuckoo,  black-billed,  35. 
Curlew,  long-billed,  26. 
Currant,  36,  44,  68. 

black,  mountain,  44,  G7-G8. 

blue,   44. 

flowering,  16,  18,  22,  28,  G7. 

red,  15,  36,  67. 

bristly,  49,  G7. 
spiny,  51. 
wild,  33. 
Cyanocephalus  cyanocephalus,  26,  35. 
Cyanocitta  stelleri  annectens,  44. 
Cynomys  leucurus,  34. 

ludovicianus  ludovicianus,  25. 
Cyrtorbyncha  ranunculina,  37. 

D. 

Dalea,  29. 

Danthonia  intermedia,  46. 
Dasiphora  fruticosa,  44,  51,  71. 
Deer,  mule,  34. 

white-tailed,  plains,  33,  34. 
Delphinium  subalpinum,  49. 
Dendragapus  obscurus  riehardsoni,  43. 
Dendroica   sestiva   sestiva,   27. 

auduboni,  35. 

nigrescens,  35. 
Deschampsia  csespitosa,   46. 
Dickcissel,  27. 
Dipper.  44. 

Disporum  tracbycarpum,  45. 
Distichlis  spicata,  30. 
Dock,  45. 

sand,  23,  28. 
Dodecatheon  radicatum,  49. 
Dolichonyx  oryzivorus,  35. 
Dove,  mourning,  16,  22,  24,  26. 
Draba,  48. 

cana,  52. 

crassifolia,  52. 

densifolia,   52. 

luteola,  49. 
Dryas  octopetala,  52,  69. 
Drymocallis  glandulosa,  37. 

pseudorupestris,    49. 
Dryobates  pubescens  bomorus,  35. 

villosus  monticola.  35. 
Dumotella  carolinensis.  27. 

E. 

Eatonia  obtusata,  30. 
Echinocactus  simpsoni,  37. 
Echinocereus  viridiflorus,  29. 
Edwinia,  68. 

americana,  36.  68. 
Elseagnus  argentea,  36,  74. 
Elder,  box.  16,  28,  73. 


Elderberry,  36,  77. 
black,  77. 

mountain,  36. 
red,  45,  76-77. 
Elephantella  groenlandica,  45. 
Elephant-head,  45. 
Elk,  34,  42,  51,  54. 
Elm,  28,  64. 
Elymus   canadensis,   30. 
condensatus,  30. 
glaucus,   46. 

salinus,  30. 
Empidonax  difficilis,  44. 

hammondi,  35. 

minimus,  35. 

wiighti,  35. 
Endolepsis  suckleyana,  28. 
Epilobium,  45. 
Eptesicus  fuscus  fuscus,  26. 
Erethizon  epixanthum,  34,  43. 
Erigeron  compositus,  50. 

pinnatisectus,  52. 

radicatus,   52. 

salsuginosus,  50. 
Eriocoma   cuspidata,  29. 
Eriogonum,  18,  28. 

annuum,  16,  28. 

campanulatum,   28. 

corymbosum,   28. 

effusum,  28. 

multiceps,  28. 
Eritricbium  argenteum,  52. 
Erythronium   parviflorum,  45. 
Eumeces  multivirgatus,  27. 
Euphagus  cyanocephalus,  35. 
Euphorbia   marginata,  29. 
Eurotia  lanata,  28. 
Eutamias  borealis,  42. 

dorsalis  utahensis,  25. 

luteiventris,  43. 

minimus   minimus,  25. 
pictus,  25. 

operarius,   42. 

oreocetes,  48,  51. 

pallidus  pallidus,  25. 

umbrinus,   43. 
Everlasting,  37,  45,  50. 

pearly,  45. 
Evotomys  brevicaudus,  43. 

gapperi  galei,  43,  48. 


Felis  hippolestes,  34,  43. 
Ferret,  black-footed,  16,  26. 
Fescue,  alpine,  52. 

Thurber,   46. 
Festuca  brachyphylla,  52. 

octoflora.  30. 

ovina  duriuscula,  37. 

tburberi.  46. 
Fiber  zibethicus  cinnamominus,  26. 

osoyoosensis.  34. 
Field  mouse.  See  Mouse. 
Finch,  house,  24.  26. 

purple,  Cassin.  44. 

rosy,  black,  51. 

brown-capped,  51. 


1917.] 


INDEX. 


87 


Fir,  39,  40,  41. 

alpine,  47,  58. 
(dwarf),  49. 

white,  44,  5S. 
Flag,  blue.  36. 
Flax,  wild,  29,  37. 
Fleabane,  50,  52. 
Flycatcher,  Hammond,  35. 

least,  35. 

olive-sided,  44. 

western,  44. 

Wright,  33,  35. 
Forget-me-not,  48,  49,  52. 
Fox,  kit,  26. 

red,  mountain,  43,  48,  51. 
Foxtail,  mountain,  46. 
Fragaria  pauciflora,  45. 
Frasera,  40,  45. 

speciosa,  45. 
Fraxinus  lanceolata,  28,  70. 
Frog,  33. 

leopard,  27. 
Fulica  amevicana,  26. 

G. 

Gadwall,  34. 
Gallinago  delicata,  34. 
Galpinsia  lavaudulcefolia,  29. 
Gaura  coccinea,  29. 
Gentian,  37,  48,  49. 

blue,  47. 

closed,  45. 

dwarf,  52. 

fringed,  mountain,  45. 
Gentiana  afflnis,  37. 

calycosa,  49. 

elegans.  45. 

forwoodi,  45. 

romanzovi,  52. 

strictiflora,  49. 
Geomys  lutescens,  26. 
Geothlypis  trichas  occidentalis,  27. 
Geranium,  37,  40,  45. 

coespitosum,  45. 

fremonti,  37. 

parryi,  45. 

richardsoni,  37. 
Gilia,  37. 

congesta,  37. 

polycladoh,  29. 
Glasswort,  19,  28. 
Glaucomys  sabrimis  bangsi,  42. 

canescens,  34. 
Globe  flower,  48,  49. 
Glycyrrhiza  lepidota,  29. 
Gnatcatcher,  western,  27. 
Golden-eye,  Barrow.  43. 
Goldenrod,  29,  52. 
Goose,  Canada,  43. 
Gooseberry,  36. 

wild,  33. 
Gopher.     See  Pocket  gopher. 
Grackle,   bronzed,  16,  24,  26, 
Grama,  tall,  30. 
Grape.  Oregon,  36,  67, 

wild,  28. 


Grass,  bent,  37. 

brome,  37. 

buffalo,  30. 
false,  30. 

bunch,  29. 

crab,  29. 

dropseed,  29,  37. 

feather,  29,  37. 

fescue,  30,  37. 

grama,  30. 

hair,  tufted,  46. 

holy,  37. 

June,  37. 

manna,  37. 

marsh,  30. 

oat,  50. 

mountain,  46. 

panic,  29. 

penny,  45,  49. 

reed,  29. 

mountain,  46. 

rice,  29. 

rye,  46. 

desert,  30. 
giant,  20,  30. 
long-bearded,  30. 

sand,  29. 

salt,  30. 

fine-top,  29. 

spear,  30,  37. 

long-stalked,  37. 

wheat,  20,  30,  37,  46. 

whitlow,  49. 

wire,  29. 
Grasshopper  mouse.     See  Mouse. 
Grass-of-Parnassus,  45. 
Grayia,  18,  19,  20,  28,  66. 

spinosa,  23,  28,  66. 
Greasewood,  17,  19,  20,  21,  28,  66-67. 
Grebe,  eared,  American,  34. 
Grindelia  squarrosa,  16,  29. 
Gromwell,  29. 
Grosbeak,   black-headed,  35. 

pine,  Rocky  Mountain,  48. 
Grossularia  inermis,  26. 
Ground  squirrel.     See  Squirrel. 
Grouse,  dusky,  76. 

Richardson,  43. 

sharp-tailed,  Columbian,  35. 
Gulo  luscus,  43. 
Gum  plant,  29. 
Gutierrezia   sarothrae,  28. 


Harbouria  trachypleura,  37. 
Hare,   varying,   northern,  42,  43, 
Harebell,  37,  48. 

arctic,  52. 

Parry,  49. 
Harvest  mouse.     See  Mouse. 
Haw,  black,  33. 

red,   33. 
Hawk,  Cooper,  35. 

rough-legged,  ferruginous,  26. 

sharp-shinned,  33,  35. 

Swainson,   35. 


88 


NORTH   AMERICAN   FAUNA. 


[No.  42. 


Hawthorn,  36,  71. 

black,  36,  71. 

red,  36. 
Hazelnut,  44. 

beaked,  42,  63. 
Heath,  mountain,  48,  49,  51,  74-78. 
Hedysarum,  37. 

Uinta,  37. 
Hedysarum  cinerascens,  37. 

sulphurescens,  49. 

uintahense,  37. 
Helianthus  annuus,  16,  29. 

petiolaris,  16,  29. 
Hellebore,  white,  45. 
Hemlock,  water,  37. 
Hemp,  Indian,  37. 
Hen,  sage,  33,  35. 
Heracleum  lanatum,  45. 
Heron,  blue,  great,  26. 

night,  black-crowned,  26. 
Heterodon  nasicus,  27. 
Heuchera  parvifolia,  45. 
Ilolbrookia  maculata,  27. 
Holodiscus  dumosus,  36,  69. 
Honey  plant,  28. 
Honeysuckle,  45. 

Douglas,  36,  42,  45,  78. 

involucred  fly,  45,  78. 

Utah,  78. 
Hop,  wild,  36. 
Hummingbird,  broad-tailed,  44. 

calliope,  35. 
Humulus  lupulus,  36. 
Hylocichla  fuscescens  salicicola,  35. 

guttata  auduboni,  44. 

ustulata  swainsoni,  44. 
Hymenopappus  filifolius,  16,  29. 

I. 

Icteria  virens  longicauda,  27. 
Icterus  bullocki,  26. 
Indigo,  false,  28,  72. 
Ipomsea  leptophylla,  29. 
Iris,  40. 

missouriensis,  36. 
Ironwood,  63. 
Ivy,  poison,  western,  28,  72. 

J. 

Jacob's  ladder,  48,  49,  52. 
Jay,  black-headed,  44. 

pinyon,  19,  26,  33,  35. 

Rocky  Mountain,   44.  49. 
Juiico,   gray-headed,  44,  49. 

pink-sided,  44,  49. 

white-winged,  33,  35. 
Junco  aikeni,  35. 

hyemalis  mearnsi,  44,  49. 

phaeonotus  caniceps,  44,  49. 
Juncoides  spicatum,  52. 
Juneus  snbtriflorus,  50. 
Juniper,  17,  18,  22,  23. 

creeping,  33,  36.  60. 

desert,  20,  28,  5J>. 

low,  44,  48,  49,  59. 

mountain,  59. 


Juniper,  one-seeded,  28,  59. 

Rocky  Mountain,  15,  33,  36,  59. 
Juniperus  communis,  59. 

knighti,  28,  59. 

monosperma,  28,  59. 

sabina,  36,  60. 

scopulorum,  36,  59. 

sibirica,  44,  49,  59. 

K. 

Kalmia  polifolia,  49,  75. 
Kangaroo  rat.  See  Rat. 
Kingbird,   26. 

Arkansas,  16,  22,  24,  26. 

Cassin,  26. 
Kinglet,   golden-crowned,   44,   49. 

ruby-crowned,  44. 
Knotweed,  alpine,  52. 
Kochia,  20,  28. 

americana,  28. 
Kceleria  cristata,  37. 
Kunzia,    33. 

tridentata,  36,  69. 


Lagopus  leucurus  leucurus,  51. 
Lanius  ludovicianus  excubitorides,  27. 
Lanivireo  solitarius  plumbeus,  35. 
Lark,  horned,  desert,  35,  51. 
Larkspur,  40,  48,  49. 

blue,  47. 
Lasionycteris  noctivagans,  43. 
Lathyrus  ornatus,  29. 
Laurel,  75. 

mountain,  48,  49. 
Ledum  glandulosum,  44. 
Lepargyrea  argentea,  28,  74. 

canadensis,  44,  74. 
Leptasea  flagellaris,  52. 

hirculus,  49. 
Lepus  americanus  americanus,  43. 

bairdii  bairdil,  43,  48. 

californicus  melanotis,  26. 

townsendi   campanius,  34. 
townsendi,  34. 
Leucosticte  atrata,  51. 

australis,  51. 
Lewisia  rediviva,  36. 
Liatris  punctata,  16,  29. 
Lichen,  51. 
Licorice,  wild,  29. 
Life  zones.     See  Zones. 
Lily,  mariposa,  36. 
Limnorchis  borealis,  45. 
Linna?a  americana,  45. 
Linum  lewisi,  37. 

rigidum,  16,  29. 
Lion,  mountain,  34,  43. 
Liopeltis  vernalis,  27,  33. 
Lippia  cuneifolia,  16,  29. 
Lithospermum  angustifolium,  29. 

gmelini,  16,  29. 
Lizard,  horned,  24,  33. 
desert,  16. 

rock,  scaly,  19,  22,  24,  27,  33. 

scaly,  16,  27. 


1917.] 


INDEX. 


89 


Lizard,  short-horned,  19. 
desert,  22,  27. 

six-lined,   16,  27. 

Stansbury,  19,  27. 
Loasa,  29. 

yellow,  29. 
Loco,  52. 
Longspur,   chestnut-collared,   35. 

McCown,  33,  35. 
Lonicera  glaucescens,  30,  45,  7S. 

involucrata,  45,  78. 

utahensis,  45,  7S. 
Lousewort,  Parry,  52. 

purple,  45. 
Loxia  curvirostra  minor,  44. 
Lungwort,  49. 
Lupine,  28,  40,  48,  49. 

mountain,  52. 

purple,  47. 

silvery,  37. 

small,  28. 
Lupinus,  16. 

argenteus,  37. 

esespitosus,  49. 

laxiflorus,  49. 

montieola,  52. 

plattensis,  28. 

pusillus,  23,  28. 
Lutra  canadensis  canadensis,  43. 
Lygodesmia  juncea,  29. 

rostra ta,  10,  29. 
Lynx,  Canada,  43. 
Lynx  canadensis  canadensis,  43. 

uinta,  34. 

M. 

Magpie,  33,  35. 

Mahogany,    mountain.    15,    18,    28,    33,    36, 

68,  68-69. 
Malacothrix,  23. 

sonchoides,  29. 
Mallard,  34. 
Mallow,  false,  29. 
Malvastrum  coccineum,  29. 

dissectum,  29. 
Mamillaria  missouriensis,  10,  29. 

vivipara,  16,  29. 
Mammal  lists,  16,  19,  20,  22,  24,  25-26,  33. 

34,  42-43,  48,  51. 
Maple,  large-toothed,  33,  36,  73. 

mountain,  33,  36,  73. 
Mariposa,  yellow,  45. 
Mariposa  lily,  36. 
Marmot,  48,  51. 

Black  Hills,  43. 

golden-mantled,  43. 

park.  43. 
Mar m ota  flaviventris  dacota,  43. 
luteola,  43,  4S. 
nosophora,  43,  48,  51. 
Marten,  Rocky  Mountain,  43,  48. 
Martes  caurina  origenes,  43,  48. 
Meadowlark,  western,   21,   26. 
Meadow  mouse.    See  Mouse. 

rue,  alpine,   52. 
Meadowsweet,  44,  69. 
Mi  lanerpes  erythrocephalus,  35. 


Melospiza  lincolni,  44. 

melodia  montana,  35. 
Mentzelia  albicaulis,  29. 
decapetala,  16,  29. 
lsevieaulis,  29. 
nuda,  29. 
Menziesia  ferruginea,  44,  75. 
Mephitis  hudsonica,  34. 
Merganser,  43. 
Mergus  americanus,  43. 
Meriolix  serrulata,  10,  29. 
Mertensia,  48. 

brevistyla,  52. 
tweedyi,    49. 
Micranthes  arguta,  45. 
Microtus  longicaudus,  43. 
montanus  caryi,  34. 
mordax  mordax,  43,  4S,  51. 
nanus  nanus,  43,  48. 
ochrogaster  haydeni,  20. 
pauperrimus,  34. 
pennsylvanicus  modestus,  34. 
richardsoni   macropus,  43. 
Milkweed,  29. 
Millet,    Indian,   29. 
Mimulus  floribundus,  37. 
langsdorfii,  45. 
lewisi,   45. 
Mimus  polyglottos  leucopterus,  27. 
Mink,    34. 
Mint,  horse,  37. 
Mockingbird,  western,  16,  27. 
Mole,  northern  plains,  16,  20. 
Monarda  rnenthsefolia,  37. 
Moneses  uniflora,  45. 
Monkey   flower,  37,   45. 

crimson,  45. 
Monkshood,    45, 
Moose,   Shiras,  42. 
Morning-glory,  bush,  16,  29. 
Moss,  51. 

Mouse,   canyon,  golden-breasted,   19,   25. 
cliff,  True's,  19,  25. 
field,  dwarf,  43,  48. 

Hayden,  16,  24,  26. 
pygmy,   33,  34. 
Uinta,  33. 
grasshopper,  Great  Plains,  16,  22,  '..'4, 
25. 

Idaho,  34. 
northern,  34. 
harvest,  prairie,  10,  25. 
«  jumping,  prairie.  33,  34. 
Rocky    Mountain,   43. 
meadow,  51. 

big-footed,  43. 
long-tailed,  43. 
Rocky  Mountain,  43,  4S. 
Saguache,    34. 
Uinta,  34. 
pocket,  20. 

Cheyenne,   20. 
Kansas,  16,  26. 
Maximilian,  17,  24,  26. 
Red  Desert.  26, 
Sweetwater,   17,  26. 
Uinta,    33,    .14. 


90 


NORTH   AMERICAN    FAUNA. 


[No.  42. 


Mouse,  red-backed,  Black  Hills,  43. 
Gale,  43,  48. 
white-footed,  34. 

black-oared,   25. 
sagebrush,  43. 
tawny,  43. 
yellow,  25. 
Muhlenbergia   comata,  37. 

pungens,  29. 
Munroa  squarrosa,  30. 
Muskrat,  Great  Plains,  26. 

Rocky  Mountain,  34. 
Mustela  arizonensis,  34,  43. 
cicognanii  leptus,  43. 
nigripes,  26. 
vison  energumenos,  34. 
Myadestes  townsendi,  44. 
Myiochanes   richardsoni,  35. 
Myosotis  alpestris,  49. 
Myotis  californicus   californicus,   26. 
evotis,   26. 

longicrus  interior,  26,  34. 
lucifugus  carissima,  43. 


X. 


Nannus  hiemalis  pacificus,  44. 
Neosorex  palustris  navigator,  43. 
Neotoma  cinerea  cinerea,  34. 

orolestes,  26,  34. 
Nettion  carolinense,  34. 
Nighthawk,  western,  21,  26. 
Ninebark,  33,  36,  69. 

western,  69. 
Nucifraga  columbiana,  49. 
Numenius  americanus,  26. 
Nutcracker,   Clark,   49. 
Nuthatch,  pygmy,  35. 

red-breasted,  44. 

Rocky  Mountain,  35. 
Nuttallornis  borealis,  44. 
Nycteris  cinerea,  43. 
Nycticorax  nycticorax  nsevius,  26. 


Oak,  bur,  33,  36,  64-65. 

Ochotona   uinta,  48,  51. 
Odocoileus  hemionus  hemionus,  34. 

virginianus  macrourus,  34. 
Olor  buccinator,  43. 
Onychomys  leucogaster  arcticeps,  25. 
brevicaudus,  34. 
missouriensis,  34. 
Oporornis  tolmiei,  35. 
Opulaster  malvaceus,  36,  69. 

monogynus,  36,  69. 

pubescens,  36,  69. 
Opuntia  fragilis,  37. 

polyacantha,  16,  29. 

rutila,   29. 
Orchid,  bog,  45. 
Oreocarya  flava,  29. 

virgata,    37. 
Oreoscoptes  montanus,  27,  35. 
Oreospiza  chlorura,  35. 
Oreoxis  alpina.  52. 


Oriole,  Bullock,  16,  22,  24,  26. 

Orpine,    red,   49. 

Ortbocarpus,  yellow,  37. 

Orthocarpus  luteus,  37. 

Oryzopsis  micrantha,  29. 

Ostrya  virginiana,  G3. 

Otocoris  alpestris  leucolsema,  35,  51. 

Otter,  43. 

Otus  asio  maxwellise,  35. 

Ouzel,  water,  44. 

Oven-bird,  35. 

Ovis  canadensis  auduboni,  34. 

canadensis,  48,  51. 
Owl,  burrowing,  16,  22,  24,  26. 

long-eared,  35. 

saw-whet,  33,  35. 

screech,  Rocky  Mountain,  35. 
Oxyria  digyna,  52. 

P. 

Pachylophus  montanus,  37. 
Pachystinia  myrsinites,  44,  72-73. 
Paintbrush,  50. 
Painted  cup,  40,  45,  48,  49. 

scarlet,  47. 
Panicularia  nervata,  37. 
Panicum  virgatum,  29. 
Parnassia  flmbriata,  45. 
Paronychia  pulvinata,  52. 
Parosela  enneandra,  29. 
Parrya,  purple,  52. 
Parrya  nudicaulis,  52. 
Parthenocissus  vitacea,  28. 
I'asserculus  sandwichensis  alaudinus,  35. 
Passerella  iliaca  schistacea,  35. 
Passerina  amoena,  27. 
Pea,  wild,  29. 

Pear,  prickly,  29.     (See  also  Cactus.) 
Pedicularis  parryi,  52. 

racemosa,  45. 
Pedicecetes  phasianellus  columbianus,  35. 
Pelecanus  erythrorhynchos,  43. 
Pelican,  white,  43. 
Pentbestes  atricapillus  septentrionalis,  35. 

gambeli,  44. 
Pentstemon  albidus,  29. 

alpinus,  49. 

angustifolius,  16,  29. 

fruticosus,  45. 

glaucus,  45. 

laricifolius,  37. 
Perisoreus   canadensis   capitalis,  44,  49, 
Perodipus  ordii  luteolus,  26. 
Perognathus  callistus,  26. 

fasciatus  fasciatus,  20. 
litus.  26. 

fiavus  piperi,  26. 

hispidus  paradoxus,  26. 

parvus  clarus,  34. 
Peromyscus  crinitus   auripectus,  25. 

leucopus  aridulus,  34. 

maniculatus  artemisi»,  43. 
nebrascensis,  25. 
osgoodi,  25. 
rufinus,  43. 

truei  truei,  25. 


1017.] 


INDEX. 


91 


Petalostemon,  16. 

candidus,  29. 

oligopbyllus,  29. 

purpureus,  29. 

villosus,  29. 
rptaya,   green-flowered,  29. 
Pewee,  wood,  western,  35. 
Phacelia,  37. 

silky,  49. 
Phacelia  heterophylla,  37. 

linearis,  37. 

sericea,  49. 
Phalaenoptilus  nuttalli  nuttalli,  35. 
Phalarope,  Wilson,  34. 
Phenacomys,  mountain,  43. 
Phenacoruys  orophilus,  43. 
Pbiladelphus  occidentalis,  28. 
Phleum  alpinum,  46,  50. 
Phlox,  48,  49. 

wild,  37. 
rhlox  caespitosa,  49. 

dcpressa,  37. 

hoodi,  37. 
Phoebe,  Say,  35. 

Phrynosoma  ornatissimum,  27,  33. 
Phyllodoce  ernpetriformis,  49,  51,  74-75. 
Pbysalis  lanceolata,  16,  29. 
Pbysaria  didymocarpa,  37. 
Physiography,  9-12. 
Pica  pica  hudsonia,  35. 
Picea  engelmanni,  44,  49,  57. 

parryana,  44,  57-58. 
Picoides  americanus  dorsalis,  44,  49. 

arcticus,  43. 
Pika,  48. 

Pine,    lodgepole,    39,    40,    41,    42,    44,    53, 
56-57. 

nut,  28,  57. 

rock,  36. 

white.  Rocky  Mountain,  32,  41,  44,  48, 
55-56. 

white-barked,  47,  49,  55. 

yellow,    15,   17,   36,  38,   39    (footnote). 
Rocky  Mountain,  56. 
Pinedrops,  45. 

Pinicola  enucleator  montana,  49. 
Pinus  albicaulis,  49,  55. 

edulis,  28,  57. 

flexilis,  44,  55-56. 

murrayana,  44,  56-57. 

scopulorum,  36,  56. 
Pinyon,  18,  28,  57. 
I'ipilo  maculatus  arcticus,  35. 
Pipit,  51. 
Pipsissewa,  45. 
Piranga  ludoviciana,  35. 
1'ituophis  sayi,  27. 

Planesticus  migratorius  propinquiiS4-35. 
Plant   lists,   15-16,   17,   18,   19,  20,"  22,   23, 
28-30,   32-33,   36-37,    39-42,    44-46,   47- 
48,  49-50,  51,  52,  55-81. 
Plantago  purshi,  16,  23;  -0. 
Plantain,  29. 
Plover,  mountain,  35. 

upland,  26. 


Plum,  ground,  29,  37. 

red,  wild,  72. 

wild,  28. 
Poa  alpina,  52. 

arctica,  52. 

epilis,  50. 

fcndleriana,  30. 

lettermanni,   52. 

longipedunculata,  37. 

lucida,  37. 

reflexa,  '46. 

sheldoni,  30. 
Pocket  gopher,  51. 

Bighorn,  43,  48. 

Black  Hills,  33,  34. 

brown,  43. 

Colorado,  43. 

Coues,  33,  34. 

Fort  Bridger,  33,  34. 

Green  River,  19,  20,  26. 

pygmy,  33,  34. 

sage,  16,  24,  26.  " 

Uinta,  16,  43,  48. 

yellow,    26. 
Pocket   mouse.     See  Mouse. 
Podasocys   montanus,  35. 
Polanisia   trachyspeima,    10,   23,   28. 
Polemonium  confertum,  52. 

mellitum,  49. 

viscosum,   49. 
Polioptila  ca?rulea  obscura,  27. 
Polygonum,  twisted,  49. 
Polygonum  bistortoides,  49. 

viviparum,  52. 
Pocecetes  gramineus  conflnis,  35. 
Poor-will,  35. 
Poplar,  aspen.  60. 

balsam,  40,  44,  60. 
Poppy,  prickly,  28. 
Populus  acuminata,  28.  61. 

augustifolia,  36,  61. 

balsamifera,  44.  60. 

occidentalis,  15,  28,'  60-61. 

tremuloides,  44,  60. 
Porcupine,   yellow-haired,  34,  43. 
Porzana    Carolina,   26. 
Potentilla   effusa,   37. 

glaucophylla,  49. 
Prairie-dog,    16. 

black-tailed,  24,  25. 

white-tailed,  33,  34. 
Primrose,  evening,  37. 
white,  .28. 

Parry,   48,   49. 
Primula  parryi,  49. 
Procyon  lotor  lotor,  26. 
Primus  americana,  28,  72. 

besseyi,  28. 

melanocarpa,  36,  72. 

pennsylvanica,  36,'  -72. 
Psaltriparus  plumbeus,  27. 
Pseudotsuga  mucronata,  44,  58—59. 
Psoralea,  16,  29. 

narrow-leaved,  29. 

silvery,  37. 


92 


NORTH   AMERICAN    FAUNA. 


[No.  42. 


Psoralea  argophylla,  37. 

esculenta,   29. 

hypogea,  29. 

laneeolata,  29. 

linearifolia,  29. 

tenuiflora,  23,  29. 
Ptarmigan,  white-tailed,  51. 
Pterospora  andromeda,  45. 
Pyrola  chlorantha,  45. 

elliptica,    45. 

picta,    45. 

secunda,   45. 

uliginosa,   45. 

Q. 

Quercus  macrocarpa,  36,  64—65. 
Querquedula  cyanoptera,  26. 

discors,    26. 
Quiscalus  quiscula  seneus,  26. 

R. 

Rabbit,  jack,  black-tailed,  26. 
white-tailed,  33,  34. 
western,  34 

snowshoe,  43,  48. 
Raccoon,    26. 
Racer,   blue,   16,   27. 
Rail,  Carolina,  26. 

Virginia,    26. 
Rallus  virginianus,  26. 
Rana  pipiens,  27.  33. 

pretiosus,  33. 
Ranunculus  adoneus,  52. 

alpeophilus,  49. 

calthaefolius,  49. 

inamoenus,  45. 
Raspberry,  dwarfed,  51. 

false.  70. 

flowering,  36,  70. 

red,  44,  49,  70-71. 
Ratibida  columnaris,  16,  29. 
Rat,  kangaroo,  20,  22,  24. 
Wyoming,  16,  26. 

wood,  bushy-tailed,  33,  34. 

Colorado,  16,  26,  34. 
Rattlesnake,  plains,  16,  19,  22,  24,  27. 
Recurvirostra  americana,  34. 
Red-backed  mouse.     See  Mouse. 
Redwing,  thick-billed,   21,  26. 
Regulus  calendula,  44. 

satrapa,  44,  49. 
Reithrodontomys   megalotis  dychel,  25. 
Reptile  lists,  16,  19,  24.  27,  33. 
Rhamnus  alnifolia,  44,  73. 
Rhodiola  integrifolia,  52. 
Rhus  rydbergi,  28.  72. 
Rhynchophanes  mccowni,  35. 
Ribes  americanum.  36. 

inebrians,  36,  67. 

lacustre.  44,  67. 

longiflorum,  28,  67. 

montigenum,  49,  51,  67. 

petiolare,  44,  67-68. 

viscosissimum,  44,  68. 

wolfi,   44. 
Robin,  western,  35. 


Rosa  sayi,  44. 

Rose,   44. 

Rosewort,  52. 

Rubus  deliciosus,  36,  70. 

parviflorus,  44.  70. 

strigosus,  44,  49,  51,  70-71. 
Rumex  paucifolius,  45. 

venosus,   16,  28. 
Rush,   50. 

wood,  52. 
Rustyleaf,  75. 
Rydbergia,   48,   49. 

grandiflora,  49. 
Rye,  wild,  30. 

S. 

Sage,  desert,  low,  20. 

pasture,  81. 

spring,  81. 

white,    17,    20,    28. 
Sagebrush,  21,  36,  80. 

alpine,   52. 

black,  17,  20,  28,  36,  79-SO. 

brown,  36,  81. 

small,  19,  22,  28. 

gray,  36,  80. 

narrow-leaved,  16,  28,  80. 

spiny,  22,  28. 
Salamander,    tiger,   27,   33. 
Salicornia    rubra,    28. 
Salix  amygdaloides,  28,  61-62. 

bebbiana,  36,  62. 

chlorophylla,  49. 

cordata  watsoni,  36. 

fluviatilis,  20,  28. 

glaucops,  49,  51,  62-63. 

mackenziana,  36. 

nelsoni,  44,  62. 

nivalis,  52,  63. 

petrophila,  52,  63. 

pyrifolia  obscura,  44,  62. 

saximontana,  49,  52,  63. 

scouleriana,  36. 

tenera,  52,  63. 
Salpinctes  obsoletus,  35. 
Saltbush,  16,  17,  18,  19,  20,  21,  28,  65. 

Nelson,  28. 

Nuttall,  21,  24,  28,  66. 

round-leaved,  65—66. 

silvery,  28. 

spiny,  28. 
Sambucus  canadensis,  36,  77. 

melanocarpa,  36,   77. 

microbotrys,  45,  76-77. 
Sandwort,    36,  52. 
Sapsucker,   red-naped,  44. 

Williamson,  44. 
Sarcobatus  vermiculatus,  28,  66-67. 
Sarsaparilla,  wild,  37. 
Savastana  odorata,  37. 
Savin,  trailing,  36.  60. 
Saxifraga  cernua,  52. 
Saxifrage,  45,  48,  49,  52. 

arctic,  52. 
Sayornis  sayus,  35. 
Scalopus   aquaticus   caryi,  26. 
Scaphiopus  hammondi  bombifrons,  27. 


1917.] 


INDEX. 


93 


Sceloporus  consobrinus,  27. 

graciosus,  27,  33. 
Schedonnardus  paniculatus,  29. 
Schmaltzia  glabra,  28,  72. 

trilobata,  28,  72. 
Sciurus  fremonti  fremonti,  42. 

hudsonicus  baileyi,  42. 
dakotensis,  34. 
ventorum,  42,  48. 
Scutellaria  brittoni,  37. 
Sedge,  50,  52. 

black,  52. 
Seiurus  aurocapillus,  35. 
Selasphorus  platycercus,  44. 
Senecio  crassulus,  50. 

fremonti,  50. 
Serviceberry,  33,  36,  71. 
Shadscale,  18. 

gray,  28,  65. 
Sheep,  bad  lands,  34. 

mountain,  48,  51. 
Shoestring,  28. 
Shooting  star,  48,  49. 
Shrew,  Dobson,  43. 

masked,  43,  48. 

Rocky  Mountain,  43,  48. 

water,  white-bellied,  43. 
Shrike,  white-rumped,  16,  21,  22,  24,  27. 
Shrubs,  distribution,  55-81. 
Sialia  currucoides,  35,  44. 
Sibbaldia  procumbens,  52. 
Sideranthus  spinulosus,  29. 
Sieversia  ciliata,  45. 

turbinata,  52. 
Silene  acaulis,  52. 
Silverberry,  33,  36,  74. 
Silver-tip,  43. 
Siskin,  pine,  44. 
Sitanion  hystrix,  30. 

longifolium,  30. 
Sitta  canadensis,  44. 

carolinensis  nelsoni,  35. 

pygmsea,  35. 
Skink,  many-lined,   16,  27. 
Skullcap,  37. 
Skunk,  northern  plains,  33,  34. 

spotted,  Great  Basin,  19,  20,  22,  26. 
Rocky  Mountain,  34. 
Skunk  bush,  16,  18,  72. 
Smelowskia  americana,  52. 
Snake,  bull,  prairie,  16,  24,  27. 

garter,  16,  27,  33. 
red-barred,  27. 
western  33. 

green,  smooth,  27,  33. 

hog-nosed,  16,  27. 
Snipe,  Wilson.  34. 
Snowberry,  36. 

few-flowered,  78. 

mountain,  33,  36,  78. 

round-leaved,  78. 
Snow-on-the-mountain,  29. 
Solanum  rostratum,  16,  29. 
Solidago  canadensis  gilvocanescens,   29. 

decumbens,  52. 

mollis,  29. 

rigida,  29. 


Solitaire,  Townsend,  44. 
Sora,  26. 

Sorbus  scopulina,  44. 
Sorex  obscurus  obscurus,  43,  48. 
personatus   personatus,  43,  48. 
vagrans  dobsoni,  43. 
Sorrel,  mountain,  52. 
Sparrow,  Brewer,  20,  26,  35. 
chipping,  western,  35. 
fox,  slate-colored,  35. 
grasshopper,  western,   16,   26. 
lark,   western,   16,    20,    22,    24,    26. 
Lincoln,  44. 
sage,  19,  20,  24,  27. 
Savannah,   western,   35. 
song,  mountain,  33,  35. 
vesper,  western,   35. 
white-crowned,  44,  49. 
Spartina  gracilis,  30. 
Speedwell,  49. 

alpine,  49. 
Speotyto  cunicularia  hypugsa,  26. 
Spermophile,  Uinta,  51. 

See  also   Squirrel. 
Sphyrapicus  thyroideus,   44. 

varius  nuchalis,  44. 
Spiderwort,  28. 
Spilogale  gracilis  saxatilis,  26. 

tenuis,  34. 
Spinus  pinus,  44. 
Spiraea  lucida,  44,  6J>. 
Spiza  americana,  27. 
Spizella  breweri,  26,  35. 

passerina  arizona?,  35. 
Sporobolus  airoides,  29. 

cryptandrus,  29. 
Spraguea   multiceps,  52. 
Spring  beauty,  45,  48,  49. 

arctic,  52. 
Spruce,  blue,  40,  44,  57-58. 

Douglas,  32,  38   (footnote),  40,  44,  53, 

58-59. 
Engelmann,    39,    40,    41,    42,    44,    47, 
53,  57. 

(dwarf),  49. 
Squirrel,  flying,  34. 

Rocky  Mountain,  42. 
ground,  chestnut-tailed,  43. 
Kennicott,  16,  25. 
mantled,    Wind    River,    4.'!,   4S,   51. 

Wortman,  25,   34 
Say,  34,  43. 
striped,  20. 
pale,  25. 
small,  25. 
Uinta,  33,  34. 
Wyoming,  33,  34. 
red,  Black  Hills,  33,  34. 
Bighorn,  42. 
Rocky  Mountain,  42,  48. 
Spruce,  FremoDt,  42. 
Stanleya,  17,  19,  20,  28. 
integrifolia,  28. 
tomentosa,  28. 
Steganopus  tricolor,  34. 
Stellula  calliope,  35. 


94 


NORTH    AMERICAN    FAUNA. 


[No.  42. 


Stipa  comata,  29. 

nelsoni,  37. 

viridula.  29. 
Strawberry,  4."). 
Streptopus  amplexifolius,  45. 
Sturnella  neglecta,  26. 
Suseda,  23. 

diffusa,  28. 

moquini,  28. 
Sumac,  smooth,  28,  72. 
Sunflower,   29. 

Swallow,  violet-green,  northern,  35. 
Swan,  trumpeter,  43. 
Swertia  congesta,  49. 

palustris.  49. 
Swift,  26. 

sand.   16,  27. 

white  throated,  33,  35. 
Sylvilagus  auduhoni  baileyi,  26. 

floridanus  similis,  26. 

nuttalli  grangeri,  34. 
Symphoricarpos  occidentalis,  28,  7S. 

oreophilus,  36,  78. 

pauciflorus,  36,  7S. 

rotundifolius,  36,  78. 
Syringa,  18. 

western,  28. 


Taehycineta  thalassina  lepida,  35. 
Tanager,  western,  33,  35. 
Taxidea  taxus  taxus,  26,  34. 
Tea,   Labrador,   44. 
Tea  bush,  wild,  36,  73. 
Teal,  blue-winged,  26. 

cinnamon,   26. 

green-winged,  34. 
Telmatodytes   palustris  plesius,   27. 
Tetradymia   inermis,  28.   81. 

nuttalli,  28,  81. 

spinosa,  28,  81. 
Thalictrum  alpinum,   52. 
Thamnophis  ordinoides  vagrans,  33. 

radix,  27,  33. 

sirtalis  parietalis,  27,  33. 
Thermopsis,  yellow,  37. 
Thermopsis  rhombifolia,  37. 
Thimbleberry,  44,  70. 
Thistle,    29. 

Thlaspi  glaucum,  45,  49. 
Thomomys  bridgeri,  34. 

fossor,    43. 

fuscus   fuscus,   43,   51. 

ocius,   26. 

pygmaeus,  34. 

talpoides  bullatus,  26. 
caryi,  43,  48. 
clusius,  34. 
nebulosus,  34. 

uinta,  43,  48,  51. 
Thoroughwax,  52. 
Thrasher,  brown,  16,  24,  27. 

sage,  21.  27,  35. 
Thrush,  hermit,  Audubon,  44. 

olive-backed,  44. 

willow,    35. 
Thryomanes  bewicki  bairdi,  27. 


Timothy,  alpine,  46,  50. 
Titlark,    51. 
Titmouse,  gray,  10,  27. 
Toad,   27,   33. 

spadefoot,   19,  27. 

western,  33. 
Tonestus  pygmanis,  52. 
Towhee,  arctic.  33,  35. 

green-tailed,  33,  35. 
Townsendia.  Tarry,  50. 
Townsendia  parryi,  50. 
Toxostoma   rufum,  27. 
Tradescantia  occidentalis,  28. 
Trees  and  shrubs,  distribution,  55-81. 
Trifolium  anemophilum,  45. 

dasyphyllum,  49. 

parryi,    49. 

rydbergi,   45. 
Trisetum  subspicatum,  50. 
Troglodytes  aedon  parkmani,  35. 
Trollius  albiflorus,  40. 
Twinflower,  45. 
Twistedstalk,   45. 
Tyrannus  tyrannus,  26. 

verticalis,    20. 

vociferans,  26. 

U. 

Ulmus  americana,  28,  64. 
Ursus  americanus,  43. 

horribilis,  34. 

imperator,   43. 

washake,    43. 
Uta   stansburiana,    27. 


Vaccinium  cn?spitosum,  45. 

erythrococcum,  45,  76. 

occidentale,  45,  76. 

oreophilum,  44,  76 
Valerian,   45. 
Valeriana  acutiloba.  45. 
Veratrum  tenuipetalum,  45. 
Verbena  bracteosa,  16,  29. 

hastata,   16,   29. 
Veronica  serpyllifolia,  49. 

wormskjoldi,  40. 
Vervain,  blue,  29. 

low,   29. 
Vetch,   37. 

milk,    29,    37,   45. 
Drummond,  37. 
Vetchling,    20. 
Viburnum,  sweet.  36,  77. 
Viburnum  lentago,  36.  77. 
Vicia    americana,    37. 
Viola  bellidifolia.  4.",. 

canadensis  rydbergi,  45. 
Violet,  45. 

dog-tooth,   45. 
Vireo,  plumbeous,  33,  35. 

warbling,  western,  35. 
Vireosylva  gilva  swainsoni,  35. 
Virgin's  bower,  36. 

purple,  36,  45. 

white,  36. 


1917.] 


INDEX. 


95 


Vitis  vulpina,  28. 

Vulpes  macrourus,  43,  4S,  51. 

velox  velox,  26. 

W. 

Warbler,  Audubon,  "5. 

gray,  black-throated,  35. 
Macgillivray,  33,  35. 
pileolated,  44,  49. 

yellow,  16,  24.  27. 
Weasel,  Arizona,  34,  43. 

dwarf,  43. 
White-footed  mouse.     See  Mouse. 
Whitlowwort,  52. 
Wildcat,  mountain,  34. 
Willet,  western,  34. 
Willow,   16,  20,  36,  44,  40,  51,  62. 

alpine,  47,  51,  52,  63. 

Bebb,  33,  36,  62. 

creeping,  52. 

diamond,  33,  36. 

gray-leaved,  49,  62-63. 

low,  47. 

Nelson,  44,  62. 

net-veined,  63. 

peach-leaved,  28,  61-62. 

rock,  52,  63. 
alpine,  63. 

sand-bar,  28. 
Wilsonia  pusilla  pileolata,  44,  49. 
Winter  fat,  28. 
Wintergreen,  45. 

one-flowered,  45. 

painted,  45. 
Wolf,  buffalo,  34. 
Wolfberry,  16,  22,  28,  78. 
Wolverene,  43. 
Woodpecker,  Batchelder,  35. 

hairy,  Rocky  Mountain,  35. 


Woodpecker,  Lewis,  33,  35. 

red-headed,  35. 

three-toed,    alpine,  44,  49. 
arctic.  4:'.. 
Wood  rat.     See  Rat. 
Wren,  Baird,  19,  27. 

house,  western,  35. 

marsh,  western,  24,  27. 

rock,  35. 

winter,  western,  44. 
Wyethia  amplexicaulis,  45. 


Xanthocephalus  xanthoccphalus,  26. 


Yarrow,  alpine,  52. 
Yellow-throat,  western,  114,  117. 
Yucca,   16,  22.   2::. 

plains,   28. 
Yucca  glauca,  28. 


Z. 


Zamelodia  melanocephala,  35. 
Zapus  hudsonius  campestris,  34. 

princeps  princeps,  43. 
Zenaidura  macroura  carolinensis,  26. 
Zone,  Arctic-Alpine,  12,  50-52. 

Canadian,   12,   38-46. 

Hudsonian,   12,  46-50. 

Transition,  12,  31-38. 

Upper  Sonoran,  12,  13-31. 
Zones,  Boreal,  52-54. 

life,   12-13. 
Zonotrichia  leucophrys,  44,  49. 
Zygadenus  elegans,  45. 

venenosus,  36. 


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V 


U.  S.  DEPARTMENT  OF  AGRICULTURE 

BUREAU  OF  BIOLOGICAL  SURVEY 

E.  W.  NELSON,   Chief 


NORTH  AMERICAN  FAUNA 

ISTo.  43 

[Actual  date  of  publication,  September  23,  1918] 


THE  RICE  RATS  OF  NORTH  AMERICA 


(Genus  ORYZOMYS ) 


BY 

EDWARD  A.  GOLDMAN 

ASSISTANT   BIOLOGIST,    BIOLOGICAL   SURVEY 


WASHINGTON 

GOVERNMENT  PRINTING   OFFICE 

1918 


t^  3  .Ulf 

LETTER  OF  TRANSMITTAL. 


United  States  Department  of  Agriculture, 

Bureau  of  Biological  Survey, 

Washington,  D.  C,  April  26, 1917. 
Sir:  I  have  the  honor  to  transmit  for  publication  as  North  American  Fauna  No.  43  the  results  of  a 
study  of  North  American  forms  of  the  rice  rats  (genus  Oryzomys),  by  Edward  A.  Goldman,  assistant 
biologist  of  this  bureau.  This  constitutes  a  revision  of  this  group,  based  chiefly  upon  material  in  the 
collection  of  the  Biological  Survey.  Rice  rats  are  distributed  from  the  latitude  of  Maryland  and  Delaware 
south  through  parts  of  the  Southeastern  States,  Mexico,  and  Central  America  to  South  America,  where 
they  reach  their  highest  development.  While  not  so  injurious  to  agriculture  as  some  other  rodents,  they 
consume  in  the  aggregate  large  quantities  of  forage  when,  like  cottonrats  and  meadow  mice,  they  increase 
locally  to  excessive  numbers.  A  knowledge  of  their  distribution,  as  presented  in  thisreport  and  its  accom 
panying  maps,  will  aid  in  studies  made  to  control  the  depredations  of  rice  rats.  Owing  to  their  nocturnal 
habits  the  animals  are  little  known,  and  their  economic  relations  should  be  better  understood. 
Respectfully,  . 

E.  W.  Nelson, 
Chief,  Biological  Survey. 
Hon.  D.  F.  Houston, 

Secretary  of  Agriculture. 


CONTENTS. 


Page. 

Introduction 3 

Genus  Oryzomys 11 

Subgenus  Oryzomys 17 

Subgenus  Oligoryzomys 87 

Subgenus  Melanomys 94 

Index 99 


ILLUSTRATIONS. 


[Following  page  98. 

Plate  I.  Skulls  (dorsal  and  central  views)  of  Oryzomys. 
II.  Skulls  (dorsal  and  ventral  views)  of  Oryzomys. 

III.  Skulls  (dorsal  and  ventral  views)  of  Oryzomys. 

IV.  Skulls  (dorsal  and  ventral  views)  of  Oryzomys. 

V.  Skulls  and  mandibles  (lateral  views)  of  Oryzomys. 
VI.  Upper  and  lower  molars  (partially  worn  crowns)  of  Oryzomys. 

TEXT  FIGURES. 

Page. 

Fig.   1.  Upper  and  lower  molar  crowns  of  Oryzomys,  showing  nomenclature  of  cusps 11 

2.  Distribution  of  subspecies  of  Oryzomys  palustris 22 

3.  Distribution  of  Oryzomys  couesi  and  closely  related  species 28 

4.  Distribution  of  the  Oryzomys  meianotis  group ." 47 

5.  Distribution  of  the  Oryzomys  alfaroi  group 56 

6.  Distribution  of  Oryzomys  talamancx 72 

7.  Distribution  of  subspecies  of  Oryzomys  bombycinus 75 

8.  Distribution  of  the  Oryzomys  devius  group 79 

9.  Distribution  of  subspecies  of  Oryzomys  tcctus 83 

10.  Distribution  of  subspecies  of  Oryzomys  fulvescens 89 

11.  Distribution  of  subspecies  of  Oryzomys  caliginosus  (exclusive  of  South  America) 96 

2 


No.  43.  NORTH  AMERICAN  FAUNA.     Sept.  23,  1918. 

THE  RICE  RATS  OF  NORTH  AMERICA. 

(Genus  Oryzomys.) 


By  Edward  A.  Goldman. 


INTRODUCTION. 

The  rice  rats  of  the  genus  Oryzomys  belong  to  the  murine  sub- 
family Cricetina?,  which  includes  also  a  number  of  closely  allied 
generic  groups  of  American  rodents,  as  the  vesper  rats  (Nyctomys) , 
deer  mice  (Peromyscus) ,  grasshopper  mice  (Onychomys) ,  and  others. 
By  reason  of  their  small  size  and  nocturnal  habits  all  these  rodents 
largely  escape  observation,  and  the  economic  importance  resulting 
from  their  excessive  numbers  and  wide  distribution  is  not  generally 
realized.  The  vernacular  name  "rice  meadow-mouse,"  bestowed 
on  Oryzomys  palustris  by  Bachman,  because  originally  it  was  found 
in  the  rice  fields  of  South  Carolina  and  Georgia,  has  led  to  the 
appellation  "rice  rat"  commonly  applied  to  all  species  of  the  genus, 
although  many  inhabit  regions  where  rice  is  not  cultivated. 

Owing  to  similarity  in  ratlike  form  and  general  appearance,  most 
of  the  genera  with  which  Oryzomys  is  allied,  including  species  with 
widely  differing  habits,  are  scarcely  recognizable  by  external  char- 
acters alone,  and  reliance  must  be  placed  on  distinguishing  features 
exhibited  by  skulls  and  teeth.  The  genus  Oryzomys  comprises  a 
somewhat  composite  assemblage  of  species  presenting  considerable 
diversity  in  general  characters,  but  having  the  essential  dental 
arrangement  repeated  with  remarkable  fidelity  throughout  the 
series. 

The  general  range  of  the  genus  is  from  New  Jersey  and  the  central 
part  of  the  Mississippi  Valley  southward,  the  group  being  represented 
in  nearly  every  part  of  South  America  and  apparently  reaching  its 
greatest  development  there.  More  than  150  species  and  subspecies 
have  been  described,  of  which  number,  however,  some  have  been  seg- 
regated in  the  erection  of  closely  allied  genera,  and  others  are  doubt- 
fully allocated.  Many  regions  remain  unexplored  and  the  number 
of  forms  assignable  to  Oryzomys  as  now  restricted  will  probably  far 
outnumber  those  of  any  other  genus  of  American  rodents.  In  the 
present  revision  are  treated  the  North  American  continental  species 

3 


4  NORTH    AMERICAN    FAUNA.  [No.  43. 

as  far  as  the  eastern  border  of  Panama,  and  those  of  outlying  islands. 
Fifty-one  forms  of  21  species  are  recognized,  two  of  which  are  char- 
acterized for  the  first  time.  These  are  comprised  in  the  three  sub- 
genera Oryzomys,  Oligoryzomys,  and  Melanomys. 

HABITS    AND    ECONOMIC    RELATIONS. 

The  habits  of  all  the  rice  rats  are  somewhat  similar,  but  differ  in 
details  in  conformity  with  varying  environmental  conditions.  In 
general,  a  preference  is  shown  for  meadows  or  marshy  areas,  com- 
monly in  the  vicinity  of  water  at  rather  low  elevations;  but  some 
species  have  ascended,  especially  along  the  courses  of  streams,  to 
high  altitudes;  others,  departing  farther  from  the  semiaquatic 
environment,  have  entered  the  forest  and  become  partially  scan- 
sorial.  They  are  nocturnal  and,  like  many  other  small  rodents,  thus 
escape  ordinary  observation,  most  species  coming  out  and  wander- 
ing here  and  there  through  marsh,  meadow,  and  herbaceous  or 
even  shrubby  vegetation.  Fairly  well-worn  runways  are  sometimes 
made  along  the  edges  of  water  or  form  general  routes  through  dense 
vegetation,  but  these  are  not  so  well  defined  as  those  of  the  cotton 
rats  (Sigmodon),  which  often  share  the  same  local  habitat.  The 
more  aquatic  species  readily  enter  the  water  and  swim  and  dive 
freely. 

The  nests,  made  of  plant  fibers  of  many  kinds,  are  placed  in  shallow 
burrows  or  sheltered  places,  commonly  under  massed  vegetation  on 
or  near  the  ground,  but  the  site  chosen  may  be  in  a  tangled  clump  of 
flags  or  marsh  grasses  standing  in  the  water.  From  3  to  7  young 
are  produced  at  a  birth,  4  or  5  being  the  usual  number.  In  Ory- 
zomys palustris  many  young  are  brought  forth  in  the  months  of 
April  and  May,  but  in  this  and  other  species  their  appearance  at  all 
seasons  is  ample  evidence  that  there  is  no  definite  breeding  season, 
except  possibly  near  the  extreme  northern  limit  of  the  group. 

The  rice  rats  feed  extensively  on  green  or  succulent  plants,  food 
habits  shared  with  the  cottonrats  (Sigmodon)  and  meadow  mice  ( Mi~ 
crotus).  In  early  morning  many  freshly  cut  grass  stems  in  the 
meadows  evidence  their  nocturnal  activity  in  favored  places.  While 
green  food  forms  the  normal  ration  of  most  species,  drier  foods,  as 
seeds,  also  are  eaten  to  some  extent,  and  as  with  some  other  small 
rodents  meat  is  relished  at  times.  Specimens  may  be  taken  in  meat- 
baited  traps,  and  individuals  thus  caught  are  sometimes  devoured 
by  their  own  kind.  Bachman,1  who  studied  the  habits  of  Oryzomys 
palustris,  records  that  those  kept  in  captivity  "  fed  on  grains  of  various 
kinds,  but  always  gave  the  preference  to  small  pieces  of  meat."  In 
their  natural  habitat  he  observed  them  scratching  up  the  recently 

1  Audubon,  J.  J.,  and  Bachman,  John,  Quadr.  North  Amer.,  Ill,  p.  215,  1854. 


1918.]  INTRODUCTION.  5 

planted  rice.  He  found  that  they  begin  feeding  on  rice  when  it  is 
in  the  milky  state  and  continue  gathering  the  scattered  grains  in 
the  fields  during  autumn  and  winter.  He  also  observed  them  feed- 
ing on  the  large  seeds  of  the  gama  grass  ( Tripsacum  dactyloides) ,  on 
those  of  the  wild  rye  (Elymus  virginicus) ,  and  at  certain  seasons  on 
those  of  the  marsh  grass  (Spartina  glabra).  Bachman  further  states 
that  the  rice  rat  "  sometimes  retires  to  the  shore  for  food,  but  has  no 
disrelish  to  the  small  Crustacea  and  mollusks  that  remain  on  the 
mud  at  the  subsiding  of  the  tide." 

Species  inhabiting  forested  areas  usually  become  very  abundant  in 
clearings  where  a  ground  cover  is  allowed  to  grow.  Rice  rats  appar- 
ently are  not  so  injurious  to  crops  as  some  other  rodents,  but  like 
the  cotton  rats  and  meadow  mice  increase  locally  to  excessive  num- 
bers and  then  consume,  in  the  aggregate,  very  large  quantities  of 
forage.  Methods  of  poisoning  that  have  proved  effective  in  checking 
the  ravages  of  meadow  mice  could  probably  be  utilized  with  similar 
success  in  the  control  of  rice  rats.  Owing  to  their  nocturnal  habits 
these  mice  are  preyed  upon  by  owls  and  doubtless  by  many  carnivor- 
ous mammals.  The  rice  rats  and  allied  members  of  the  great  murine 
family  to  which  they  belong  are  the  most  numerous  of  American 
mammals,  and  their  economic  relations  should  be  better  known. 

GENERAL    CHARACTERS. 

The  rice  rats  as  a  whole  present  a  wide  range  of  variation  in  external 
appearance.  Some  of  the  larger,  more  robust  species,  including 
Oryzomys  palustris,  have  not  infrequently  been  mistaken  for  Old 
World  rats  of  the  genus  Rattus,  which  have  followed  civilized  man 
in  his  migrations  and  now  not  only  universally  infest  his  structures, 
but  in  favorable  districts  invade  the  fields  in  close  competition  with 
native  rodents.  Superficial  resemblances  are  often  striking,  the 
bodily  proportions  and  color  and  texture  of  pelage  being  very  similar. 
On  the  other  hand,  many  species  are  widely  different  from  the  Old 
World  rats,  in  rich  tawny  coloration  or  character  of  pelage,  and  are 
more  likely  to  be  confused  with  allied  American  murine  genera. 
The  smallest  North  American  species  (subgenus  Oligoryzomys)  simu- 
late in  color,  delicate  structure,  and  great  length  of  tail  some  of  the 
harvest  mice  (Beithrodontomys) ;  another  group  (subgenus  Melanomys) 
is  more  robust,  the  tail  very  short,  and  general  proportions  Akodon- 
like. 

In  Oryzomys  and  allied  genera  superficial  resemblances  frequently 
mask  the  more  essential  features,  and  external  characters,  especially 
size  and  color,  are  less  dependable  than  cranial  modifications  in  tracing 
relationships  of  species.  The  group  alignment,  however,  is  often 
indicated  by  proportions  of  body  and  limbs;  color  and  length  of 
general  pelage;  length  of  vibrissa?;  hairiness  of  ears,  feet,  and  tail; 


6  NORTH    AMERICAN    FAUNA.  [No.  43. 

presence  or  absence  of  prominent  tufts  of  digital  bristles;  and  form 
of  claws.  Aside  from  the  general  form  and  angularity  of  the  skull, 
the  principal  characters  of  taxonomic  value  are  the  following:  Size 
and  form  of  incisors  and  molars;  depth  and  arrangement  of  reentrant 
angles,  form  and  position  of  enamel  islands,  and  development  of 
cusps  in  molar  crowns ;  length  and  form  of  anterior  palatine  foramina ; 
length  of  palatal  bridge;  position  of  lachrymals;  form  of  maxillae, 
premaxillae,  frontals,  parietals,  and  interparietal;  and  size  and  form 
of  audital  bullae. 

PELAGE. 

The  pelage  is  rather  harsh  in"  texture  throughout  the  genus,  but 
varies  greatly  in  length  in  the  component  groups.  The  overfur  is 
longer,  the  vibrissas  shorter,  and  the  underfur  denser  and  more  woolly 
in  semiaquatic  species.  Groups  in  which  the  pelage  is  normally  short 
tend  to  develop  longer  fur  in  the  forms  which  range  at  high  altitudes : 
but  several  species,  some  with  pelage  much  longer  than  others,  may 
occur  together  at  the  same  elevations.  The  ears  are  rather  small  and 
in  typical  Oryzomys  and  various  groups,  including  the  subgenus  Oligo- 
ryzomys,  the  hairy  eovering  is  moderately  long,  coarse,  and  not 
sharply  differentiated  from  the  body  fur;  in  the  0.  talamancse  and 
other  groups  the  ears  are  minutely  pilose,  in  marked  contrast  with  the 
general  pelage.  The  claws  on  the  toes  of  the  hind  feet  in  the  semi- 
aquatic  or  more  strictly  terrestrial  species  are  relatively  long  and 
straight,  broad,  and  obtuse,  and  only  partially  overlapped  by  the 
digital  bristles,  while  in  the  less  aquatic  or  scansorial  species  they  are 
short  and  recurved,  compressed  and  sharp  pointed,  and  prominent 
digital  tufts  project  beyond  the  ends.  In  0.  palustris  the  vibrissas 
scarcely  reach  from  the  muzzle  to  the  ears;  in  0.  pirrensis  and  0. 
hombycinus  they  extend  over  the  shoulders,  while  in  various  species 
they  are  intermediate  in  length. 


In  coloration  of  upperparts  the  range  of  variation  in  the  rice  rats  is 
from  pale  shades  of  buff  or  gray  to  rich  tawny  or  russet,  more  or  less 
mixed  with  black,  especially  over  the  median  part  of  the  dorsum. 
The  underparts  usually  are  dull  white  or  buff,  without  a  sharp  line 
of  demarcation  along  the  sides,  the  plumbeous  basal  color  showing 
through  (a  few  species  exhibiting  basally  white  areas).  The  tail  is  not 
very  sharply  bicolor,  but  usually  is  brown  above  and  lighter  below, 
at  least  basally,  the  epidermis  and  scanty  investing  hairs  of  about  the 
same  tone. 

MOLT. 

,As  breeding  begins  during  adolescence  and  is  continuous  through- 
out the  year,  and  as  individuals  arrive  at  maturity  at  all  seasons, 
there  is  no  very  definite  period  for  molting,   although  the  more 


1918.]  INTRODUCTION.  7 

northerly  forms  tend  in  winter  to  acquire  a  longer  pelage,  which  be- 
comes abraded  in  summer.  As  a  rule  the  new  coat  seems  to  replace 
the  old  almost  imperceptibly,  but  adults  in  apparently  fresh  and 
in  obviously  worn  pelages  may  often  be  seen  together. 


The  age  of  individuals  is  indicated  approximately  by  the  degree 
of  wear  on  the  molar  crowns.  The  shearing  of  the  slopes,  beginning 
early  in  life,  becomes  distinctly  noticeable  as  full  growth  is  attained, 
and,  progressing  rapidly  across  the  summits  of  the  tubercles,  in 
advanced  age  results  in  the  obliteration  of  all  trace  of  enamel 
arrangement. 

VARIATION. 

Variation  in  the  rice  rats  is  assignable  to  several  categories,  of 
which  perhaps  the  most  obvious  are  individual  and  geographic. 

INDIVIDUAL   VARIATION. 

By  individual  variation  is  meant  all  the  degrees  of  divergence 
from  a  typical  mean  exhibited  by  large  series  of  conspecific  skins 
and  skulls  from  any  given  locality.  The  range  of  this  variation, 
especially  in  general  size,  is  extraordinary;  in  many  species  of  corre- 
sponding age  and  sex  it  exceeds  10  per  cent  both  in  external  and 
cranial  dimensions.  The  typical  mean,  therefore,  may  be  difficult 
to  determine  when  a  small  series  of  examples  exhibits  preponder- 
ance toward  either  of  the  extremes;  and  conclusions  based  on  the 
dimensions  of  a  small  number  of  individuals  are  likely  to  be  mis- 
leading. While  males  average  slightly  larger  than  females,  sexual 
differences  in  size  appear  to  be  negligible.  Cranial  variations  in 
proportions  and  in  the  form  of  individual  bones  are  noticeable,  but 
usually  within  rather  circumscribed  limits.  Some  skulls  are  decid- 
edly broader  and  more  massive  than  others  of  the  same  age  and  sex. 
Thickness  of  rostrum  is  usually,  but  not  always,  associated  with 
breadth  of  frontal  region  and  braincase.  General  expansion  of  the 
braincase  commonly  results  in  increased  breadth  across  the  posterior 
part  of  the  frontals.  The  interparietal  is  variable  in  form,  as  are  the 
parietals  in  the  extent  of  encroachment  of  the  lateral  wings  on  the 
squamosals.  The  size  of  the  molar  teeth  and  of  the  audital  bullae 
is  fairly  constant. 

Individual  variation  in  color  is  much  less  than  in  size.  Much  of 
the  variation  in  color  observable  is  due  to  age  or  condition  of  pelage. 
The  older  adults  tend  at  all  seasons  to  exhibit  more  rufescent  tones 
than  the  younger.  A  rusty  reddish  appearance  is  often  due  to  much- 
worn  pelage.     There  are  no  distinct  color  phases. 


NORTH   AMERICAN    FAUNA.  [No.  43. 


GEOGRAPHIC   VARIATION. 


Geographic  variation,  or  the  tendency  of  species  to  subdivide  into 
regional  or  more  or  less  localized  forms,  is  very  great  in  the  genus 
Oryzomys.  Of  the  numerous  forms  first  described  as  distinct  species 
a  considerable  number  prove  when  better  known  to  be  geographic 
representatives  of  widely  ranging  specific  types,  presenting  differ- 
ential characters  associated  in  part  with  environmental  conditions. 
The  intergradation  and  subspecific  position  of  many  such  forms  can 
be  demonstrated  beyond  any  reasonable  doubt  in  some  instances, 
and  in  others  may  be  safely  assumed,  in  the  light  of  knowledge  of 
the  essential  characters  of  the  /particular  group.  Since  the  distri- 
bution of  the  rice  rats  is  mainly  at  low  elevations,  it  is  not  surprising 
to  find  that  species  maintaining  the  same  characters  over  extensive 
areas  near  sea  level,  where  nearly  uniform  topographic  and  climatic 
conditions  prevail,  become  locally  modified  on  ascending  to  high 
altitudes.  Thus,  0.  couesi  and  0.  alfaroi  are  represented  by  widely 
ranging  coastal  forms  and  by  more  localized  high-mountain  races. 
That  forms  inhabiting  mainly  open,  arid  regions  are  paler  than  those 
inhabiting  humid  or  heavily  forested  areas  is  well  illustrated  by  the 
distribution  of  the  races  of  0.  couesi  in  Mexico.  The  dark  typical 
form  occupies  the  relatively  humid  area  near  the  Gulf  of  Mexico  and 
the  Caribbean  Sea,  while  the  pale  subspecies,  0.  c.  mexicanus, 
pushes  far  northward  along  the  arid  Pacific  coast,  the  point  of  diver- 
gence being  near  the  Isthmus  of  Tehuantepec.  Species  reaching 
high  altitudes  tend  to  develop  structural  as  well  as  color  differences, 
while  varying  climatic  conditions  at  low  elevations  are  apt  to  result 
mainly  in  color  modifications.  Insular  species  are  usually  related  to 
those  inhabiting  the  adjacent  mainland,  but  may  exhibit  very  dis- 
tinctive characters,  unless  the  islands  are  very  near  the  coast. 

HISTORY   AND   MATERIAL. 

A  species  of  Oryzomys  was  first  described  under  the  name  Mus  palus- 
tris  from  New  Jersey,  by  Richard  Harlan,1  in  1S37.  He  compared  the 
animal  with  the  Norway  rat,  and  owing  to  superficial  resemblances 
regarded  it  as  congeneric  with  the  rats  of  the  Old  World.  It  was 
erroneously  referred  to  the  genus  Arvicola  in  1854  by  Bachman,2  who 
mentions  having  obtained  specimens  as  early  as  the  winter  of  1816. 
According  to  Bachman,  these  specimens  were  described  by  him  (but 
the  description  not  published)  in  May,  1836.  One  was  sent  to  the 
Academy  of  Natural  Sciences  of ,  Philadelphia  for  comparison  with 
material  there,  and  on  the  basis  of  this  and  an  example  in  the  Academy 
collection  Dr.  Harlan  felt  authorized  to  publish  his  Mus  palustris. 

i  Silliman's  Amer.  Journ.  Sci.,  XXXI,  p.  385,  1837. 

*  Audubon,  J.  J.,  and  Bachman,  John,  Quadr.  North  Amer.,  Ill,  p.  214,  1854. 


1918.]  INTRODUCTION".  9 

In  renaming  the  species  Arvieola  oryzivora,  Bachman  assumed  that 
Mus  palustris  Harlan  was  preoccupied  by  Arvieola  palustris  Harlan,1 
an  obvious  error,  as  the  two  were  not  congeneric. 

In  1857  Baird,2  recognizing  distinctive  characters,  used  Oryzomys 
as  a  full  generic  name  for  the  group,  with  Mus  palustris  Harlan  as 
type,  apparently  inadvertently,  however,  as  on  a  later  page  of  his 
publication 3  he  accorded  it  only  subgeneric  value  under  Hesperomys 
of  Waterhouse.  In  this  course  he  was  followed  by  Coues  in  1877.4 
Thirteen  years  later  the  same  author5  raised  the  name  to  generic 
rank.  Meanwhile  Hesperomys  fulvescens  Saussure  6  and  Hesperomys 
couesi  Alston  7  had  been  described,  but  their  real  generic  position 
was  not  determined  until  later.  New  species  were  added  at  inter- 
vals, but  of  the  relationships  of  the  North  American  members  of  the 
genus  little  was  known  until  1901,  when,  as  a  result  of  study 
mainly  of  material  which  had  accumulated  in  the  collection  of  the 
Biological  Survey,  Merriam8  published  a  synopsis  of  the  forms 
inhabiting  the  United  States  and  Mexico.  Thirty-five  species  and 
subspecies  were  recognized  by  him,  of  which  20  were  new.  The 
species  were  divided  into  natural  groups  for  the  first  time,  and  their 
salient  characters  pointed  out.  Short  papers,  largely  descriptive  of 
new  species,  by  Thomas,9  Allen,10  Bangs,11  Elliott,12  and  Goldman,13 
have  since  appeared.  The  larger  collections  now  available  render  it 
possible  to  determine  the  status  of  nearly  all  names,  and  the  relation- 
ship, especially  of  the  more  austral  species,  to  South  American  forms. 

The  present  revision  is  the  result  of  a  study  of  the  rice  rats  in  the 
Biological  Survey,  the  Merriam,  and  other  collections  in  the  United 
States  National  Museum,  now  numbering  1,050  specimens,  aug- 
mented by  563  from  other  American  museums,14  the  assemblage 
including  the  types  or  topotypes  of  most  of  the  species.     The  location 

1  Harlan,  Richard,  Fauna  Americana,  p.  13G,  1825. 

2  Baird,  S.  F.,  Mamm.  North  Amer.,  p.  459,  1857. 

*  Op.  cit.,  p.  482. 

*  Coues,  Elliott,  Monogr.  North  Amer.  Rodentia,  p.  113,  1877. 
» Coues,  Elliott,  Century  Diet.,  IV,  p.  4165,  1890. 

«  Saussure,  H.de,  Rev.  et  Mag.  de  Zool.,  ser.  2,  XII,  p.  102,  March,  1860. 

i  Alston,  E.  R.,  Proe.  Zool.  Soc,  London,  p.  756,  187G. 

8  Merriam,  C.  Hart.,  Proc.  Washington  Acad.  ScL,  III,  pp.  273-295,  July  26,  1901. 

»  Thomas,  O.,  Ann.  Mag.  Nat.  Hist.,  ser.  7,  VIII,  pp.  251-253,  Sept.,  1901. 

io  Allen,  J.  A.,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXIV,  pp.  654-657,  Oct.  13,  1908  (including  Oryzomys  ochra- 
ceus  [=Nectomys  alfari],see  Goldman,  Proc.  Biol.  Soc,  Washington,  XXIX,  p.  127,  June  6,  1916);  ibid., 
XXXIII,  pp.  99-100,  Apr.  30,  1910;  ibid.,  XXXII,  pp.  533-554,  Nov.  17,  1913. 

U  Bangs,  O.,  Bull.  Mus.  Comp.  Zool.,  XXXIX,  pp.  33-36,  Apr.  1902. 

12  Elliott,  D.  G.,  Field  Columb.  Mus.  publ.  71,  Zool.  ser.  Ill,  p.  145,  Feb.,  1903;  ibid.,  Ill,  pp.  266-267, 
Mar.  1904. 

is  Goldman,  E.  A.,  Smiths.  Misc.  Coll.,  LVI,  no.  36,  pp.  5-8,  Feb.  19,  1912;  ibid.,  LX.,  no.  22,  pp.  5-6, 
Feb.  28,  1913;  Proc.  Biol.  Soc.  Washington,  XXVIII,  pp.  127-130,  June  29,  1915. 

n  For  the  use  of  material  generously  loaned  and  for  other  courtesies  the  author's  thanks  are  due  to  Dr.  J.  A. 
Allen,  American  Museum  of  Natural  History;  to  Mr.  Samuel  Henshaw  and  Mr.  Outraru  Bangs,  Museum 
of  Comparative  Zoology;  to  Mr.  W.  H.  Osgood,  Field  Museum  of  Natural  History;  to  Mr.  Witmer  Stone, 
Academy  of  Natural  Sciences  of  Philadelphia;  to  Mr.  W.  E.  Clyde  Todd  and  Mr.  O.  P.  Murie,  Carnegie 
Museum;  and  to  Mr.  CD.  Bunker,  Kansas  University  Museum.  In  addition  he  is  indebted  to  Mr.  Oldfield 
Thomas  for  critical  notes  and  comparisons  of  specimens  with  types  in  the  British  Museum. 


10  NORTH   AMERICAN    FAUNA.  [No.  43. 

of  specimens  examined  in  collections  other  than  those  in  the  United 
States  National  Museum  is  indicated  by  footnotes. 

EXPLANATIONS. 

MEASUREMENTS . 

All  measurements  of  specimens  are  in  millimeters.  With  a  few 
exceptions,  usually  stated,  the  external  measurements  were  taken  in 
the  flesh  by  the  collector,  as  follows :  Total  length,  nose  to  end  of  ter- 
minal tail  vertebra;  tail  vertebrse,  upper  base  of  tail  to  end  of  terminal 
tail  vertebra;  hind  foot,  heel  to  end  of  longest  claw.  While  adult 
males  average  slightly  larger  tfyan  females,  the  difference  is  scarcely 
appreciable  and  in  the  small  series  usually  available  may  be  ignored. 
The  external  and  cranial  measurements  given,  therefore,  are  of  series 
which  may  include  specimens  of  both  sexes.  Of  many  species  and 
subspecies  so  few  nearly  typical  examples  are  available  that  the  meas- 
urements given  may  not  represent  the  normal  range  of  individual  va- 
riation, and  too  broad  generalizations,  therefore,  should  not  be  based 
upon  them.  The  following  cranial  measurements  were  taken  with  a 
vernier  caliper  by  the  author : 

Greatest  length. — Length  from  tip  of  nasals  to  supraoccipital  in 
median  line  over  foramen  magnum. 

Zygomatic  breadth. — Greatest  distance  across  zygomata. 

Interorbital  breadth. — Least  distance  between  orbits. 

Width  of  braincase. — Distance  between  outer  sides  of  squamosals 
at  the  slight  constriction  over  auditor}^  meatus  and  immediately  in 
front  of  lateral  occipital  crests. 

Nasals. — Greatest  length  of  nasals. 

Anterior  palatine  foramina. — Greatest  length  of  large  palatal 
foramina. 

Palatal  bridge. — Distance  from  excavated  posterior  border  of  palate 
to  posterior  end  of  either  large  palatal  foramen. 

Upper  molar  series. — Greatest  length  of  maxillary  toothrow  at 
alveolar  border. 

COLORS. 

The  names  of  colors  used  in  descriptions  are  mainly  those  of 
Ridgway.1  A  few  other  modifying  or  comparative  terms,  however, 
have  been  employed,  usually  when  some  special  difficulty  was 
encountered  in  naming  an  indefinite  hue  or  tone. 

TEETH. 

The  nomenclature  of  the  principal  tooth  elements  used  in  the  text 
is  given  in  figure  1 .2 

1  Ridgway,  Robert,  Color  Standards  and  Color  Nomenclature,  1912. 

2  For  the  homology  and  nomenclature  of  the  molar  crown  divisions  the  writer  is  indebted  to  Messrs. 
Gerrit  S.  Miller,  jr.,  and  James  W.  Gidley,  whose  extensive  researches,  still  in  progress,  in  the  phylogeny 
of  the  rodents,  enable  them  to  render  an  authoritative  opinion. 


1918.] 


GENUS    ORYZOMYS. 


11 


ZONAL   DIVISIONS. 


Aside  from  the  well-known  extratropical  North  American  zonal 
divisions,  all  references  in  the  text  under  "Geographic  distribution" 
are  to  less  well-known  tropical  divisions,  the  Lower  and  Upper 
Tropical  Zones,  which  may  be  roughly  denned  as  follows:  The  Lower 
Tropical  Zone,  extending  in  tropical  Middle  America  from  sea  level  to 
elevations  varying  mainly  in  accordance  with  latitude  and  local 
topographic  conditions.  South  of  the  twentieth  parallel  this  zone 
reaches  to  about  3,000  or  3,500  feet  altitude.  Above  these  limits  it 
is  replaced  by  the  Upper  Tropical  Zone,  which  extends  to  about  7,000 


pre Protoconule . 

pr Protocone. 

hy Hypocone. 


prcd Protoconulid. 

prd Protoconid. 

hyd Hypoconid. 


pad Paraconid. 

med Metaconid. 

mesd Mesostylid. 

end Entoconid. 

ensd Entostylid. 


pas Parastyle. 

sec.  pas Secondary 

parastyle. 

pa Paracone. 

ms Mesostyle. 

me Metacone. 

mts Metast  yle . 

Fig.  1.— Molar  teeth  of  typical  Oryzomys  with  outlines  accentuated  to  show  principal  crown  elements. 
A,  Right  upper  molars;  B,  left  lower  molars. 

or  8,000  feet.  Each  of  these  zones  is  readily  separable  into  sub- 
divisions on  the  basis  of  moisture,  and  are  denominated,  respectively, 
the  Humid  and  Arid  Lower  Tropical  and  the  Humid  and  Arid  Upper 
Tropical  Zones. 

GENUS  ORYZOMYS  Baird. 

Oryzomys  Baird,  Mamm.  North  Amer.,  pp.  xlii,  458,  482,  1857  (subgenus  of  Hes- 
peromys  Waterhouse).     Type  Mus  palustris  Harlan. 

Oryzomys  Coues,  Century  Diet.,  IV,  p.  4164,  1890. 

Oligoryzomys  Bangs,  Proc.  New  England  Zool.  Club,  I,  p.  94,  Feb.  23,  1900  (sub- 
genus).   Type  Oryzomys  navus  Bangs. 

Melanomys  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  7,  X,  p.  248,  Sept.  1902  (subgenus). 
Type  Oryzomys  phseopus  Thomas. 

Melanomys  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXXII,  p.  533,  Nov.  17,  1913  (genus). 

Geographic  distribution. — Eastern  and  central  United  States  from 
New  Jersey,  Kentucky,  Illinois,  Kansas,  and  Texas  across  Mexico 
to  Lower  California  and  south  through  South  America  to  Cape  Horn. 


12  NORTH  AMERICAN   FAUNA.  [No.  43. 

Generic  characters. — Form  murine;  pelage  slightly  hispid,  but  not 
bristly  or  spiny;  tail  usually  long,  but  varying  from  about  three- 
fourths  length  of  head  and  body  to  one-fourth  longer,  the  annulated 
scales  distinctly  visible  through  short,  sparse  investing  hairs;  ears 
varying  from  small  to  rather  large,  and  finely  to  coarsely  haired; 
soles  of  hind  feet  naked  to  heels,  normally  6-tuberculate ;  mamma?  8: 
pectoral,  2-2;  inguinal,  2-2. 

Skull  relatively  thin  and  smoothly  rounded;  supraorbital  and 
temporal  ridges  prominent  or  obsolescent;  interparietal  variable  in 
size  and  contour ;  zygomata  slender,  depressed  to  near  level  of  molars ; 
antorbital  foramen  subcylindrical  above,  more  or  less  abruptly  con- 
stricted to  a  narrow  slitlike  opening  below;  outer  wall  of  antorbital 
foramen  rather  broad,  but  varying  in  extent  of  anterior  projection, 
the  upper  border  rounded  or  slightly  angular;  palate  reaching 
posteriorly  beyond  plane  of  last  molars,  the  interpterygoid  border 
concave  or  presenting  a  slight  median  projection;  palatal  pits 
present;  audital  bullas  rather  slightly  inflated,  tapering  anteriorly 
and  oblique  in  position. 

Molar  crowns  low  (slightly  higher  in  Melanomys  than  in  the 
other  subgenera)  with  prominent  cusps,  cones,  or  tubercles,  and 
well-developed  styles,  the  principal  cusps  arranged  nearly  opposite 
in  two  longitudinal  series;  upper  molars  3-rooted;  lower  molars 
2-rooted.  First  upper  molar  with  parastyle  and  paraconule  strongly 
developed,  partially  divided  near  middle,  and  extended  across 
internally  for  a  distance  equal  to  three-fourths  or  more  of  the  greatest 
transverse  diameter  of  tooth;  secondary  parastyle  usually  prominent; 
mesostyles  and  metastyles  in  all  the  upper  molars  present  as  small 
cusps  or  represented  by  enamel  ridges  connecting  with  principal 
inner  cusps;  inner  cusps  of  first  and  second  upper  molars  with  oblique 
commissures ;  third  upper  molar  with  posterior  portions  more  or  less 
obsolete,  the  hypocone  varying  in  development  and  the  metacone 
usually  indicated  only  by  a  low  abbreviated  enamel  fold.  Lower 
molar  cusp-arrangement  similar  to  upper,  but  paraconid  and  para- 
conulid  in  anterior  tooth  faintly  or  not  at  all  notched  longitudinally; 
protoconids  and  hypoconids  in  first  and  second  molars  with  oblique 
commissures  as  in  the  homologous  elements  in  upper  teeth;  meso- 
stylids  and  entostylids  joined  by  transverse  enamel  ridges  with  outer 
principal  cusps;  second  lower  molar  with  a  moderately  developed 
protoconulid;  third  lower  molar  with  entoconid  and  entostylid  com- 
pletely fused  or  obsolescent.  Front  of  upper  incisors  without 
grooves. 

Remarks. — When  the  molars  of  Oryzomys  are  opposed  the  cusps 
in  the  upper  jaw  enter  behind  the  corresponding  cusps  in  the  lower 
jaw.     The  parastyle  is  overlapped  by  the  paraconid,  the  trenchant 


1918.]  GENUS   OKYZOMYS.  13 

anterior  border  of  which  shears  on  its  anterior  surface.  The  para- 
conule  of  the  anterior  molar  and  the  protocones  and  hypocones  of 
the  other  molars  are  broadly  beveled  or  sheared  internally,  as  are 
the  protoconids  and  hypoconids.  The  paracones  and  metacones,  on 
the  other  hand,  are  at  first  sheared  posteriorly,  while  the  metaconids 
and  entoconids  are  sheared  anteriorly,  but  progressive  wear  extend- 
ing across  the  summits  tends  to  level  the  entire  series  uniformly. 

The  genus  presents  complex  relationships  to  various  murine  genera. 
Among  its  nearer  relatives  is  Nectomys,  which  the  more  typical 
forms  much  resemble  in  external  appearance  as  well  as  in  the  general 
form  of  the  cranium.  In  dentition  the  two  genera  are  also  similar, 
but  in  typical  Oryzomys,  while  the  molar  crowns  as  a  whole  are 
decidedly  lower,  the  cusps  or  cones  are  higher,  more  conical,  with 
summits  more  fully  covered  with  enamel.  In  Nectomys,  on  the 
other  hand,  the  low  cusps  tend  to  fuse  with  the  stjdes  and  dentine  is 
exposed  at  an  earlier  age.  Oryzomys  differs  from  Nectomys  in  other 
dental  details,  especially  the  development  of  the  paraconulid  of  the 
second  lower  molar.  The  subgenus  Melanomys,  however,  exhibits  a 
somewhat  intermediate  condition;  the  molar  crowns  as  a  whole  are 
somewhat  higher  than  in  typical  Oryzomys,  and  lower  than  in  Nec- 
tomys; the  cones  are  high  as  in  Oryzomys,  but  in  sculpture,  especially 
the  early  exposure  of  dentine,  approach  those  of  Nectomys.  Another 
character  suggesting  gradation  of  Melanomys  toward  Nectomys  is 
the  position  of  the  lachrymal,  its  attachment  being  mainly  with  the 
maxilla  as  in  that  genus,  instead  of  about  equally  with  maxilla  and 
frontal  as  in  typical  Oryzomys.  Agreement  with  Oryzomys  is  shown 
in  the  more  essential  dental  details,  including  the  marked  develop- 
ment of  the  paraconulid  in  the  second  lower  molar,  an  element 
absent  in  Nectomys. 

Comparison  with  various  other  genera  reveals  obvious  resem- 
blances and  points  of  difference  in  varying  combination.  Oryzomys 
agrees  closely  with  Neacomys  in  cranial  and  dental  characters,  but 
the  latter  genus  exhibits  a  departure  in  its  grooved  and  bristly  or 
spiny  pelage.  The  generic  name  Nesoryzomys  based  on  rice  rats 
inhabiting  the  Galapagos  Islands  does  not  seem  well  founded,  as  the 
palate  and  dentition  are  Oryzomys-]ik.Q  and  in  the  short  tail  and 
reduced  interorbital  space,  alleged  generic  characters,  it  is  not  widely 
different  from  some  of  the  continental  species  of  Oryzomys. 

In  dentition  Oryzomys  is  similar  to  RMpidomys,  but  the  molar 
cusps  are  more  crowded  and  the  parastyle  and  protoconule  are  less 
distinctly  separated  by  a  median  notch.  The  posterior  part  of  the 
palate,  however,  presents  features  at  variance  in  the  two  genera. 
Unlike  that  of  Oryzomys  the  palate  in  Rhipidomys  is  excavated  be- 
tween the  posterior  molars,  and  the  palatal  pits  are  separated  by  the 


14  NORTH  AMERICAN   FAUNA.  [No.  43. 

interpterygoid  fossa.  The  genus  Nyctomys  is  allied  to  Rhipidomys, 
but  exhibits  a  wider  departure  from  Oryzomys.  The  molar  crown 
arrangement  is  similar  in  all  three  genera,  but  in  Nyctomys  the  cusps 
are  more  angular,  the  styles  and  commissures  less  developed,  and  the 
enamel  covering  maintained  until  later  in  life  than  in  the  others. 
In  the  posterior  excavation  of  the  palate  Nyctomys  more  nearly 
resembles  Rhipidomys;  but,  in  the  remarkable  lateral  development  of 
the  interparietal  to  a  broad  line  of  contact  with  the  squamosals  and 
consequent  isolation  of  supraoccipital  and  parietals,  Nyctomys  differs 
notably  from  both  Rhipidomys  and  Oryzomys. 

Some  sections  of  the  large  genus  Peromyscus  are  not  very  widely 
different  from  Oryzomys  in  deritition,  but  the  principal  cusps  are 
obliquely  placed,  the  protoconule  much  more  developed,  and  the 
paracone  and  metacone  shear  mainly  internally  instead  of  poste- 
riorly as  in  Oryzomys.  The  general  form  of  the  skull  in  Peromyscus 
is  distinctive;  the  palate  ends  near  the  posterior  plane  of  the  molars 
and  the  palatal  pits  present  in  Oryzomys  are  absent. 

In  external  appearance  the  genus  Reithrodontomys  bears  a  remark- 
able resemblance  to  the  oryzomine  subgenus  Oligoryzomys;  and  in 
the  Aporodon  section  of  Reithrodontomys,  in  which  the  styles  are 
present,  an  approach  in  molar  pattern  is  manifested;  but  in  Reith- 
rodontomys generic  distinction  is  lodged  in  the  grooved  upper  in- 
cisors. The  genus  Zygodontomys,  until  recently  associated  with 
Oryzomys,  is  similar  in  outward  appearance  and  in  general  form  of 
skull,  but  in  height  of  molar  crowns,  absence  of  style  ridges,  and 
longitudinal  instead  of  oblique  commissures  of  cusps  it  departs 
widely  from  Oryzomys  and  more  nearly  approaches  ATcodon. 

SUBGENERA    AND    MINOR    GROUPS. 

Of  the  three  subgenera  into  which  the  North  American  species  of 
the  genus  Oryzomys  are  here  divided,  Oligoryzomys  and  Melanomys  each 
contain  a  single  group  of  closely  related  forms,  while  the  subgenus 
Oryzomys  is  divisible  into  seven  rather  well-defined  minor  groups,  or 
assemblages,  of  species  or  subspecies  (excluding  0.  victus,  whose 
exact  relationships  are  unknown).  These  assemblages  are  usually 
distinguishable  by  external  characters,  but  recourse  to  cranial  struc- 
ture is  sometimes  necessary  in  order  to  make  accurate  determina- 
tions. 

(1)  The  0.  palustris  group  is  characterized  by  large,  robust  form, 
small,  coarsely  haired  ears,  short  vibrissas,  and  the  absence  of  con- 
spicuous tufts  of  silvery  bristles,  which  in  all  the  other  groups  pro- 
ject beyond  the  ends  of  the  longer  claws  on  the  hind  feet.  The  skull 
is  broad  with  short  rostrum,  very  long  anterior  palatine  foramina 
(normally  reaching  anterior  plane  of  first  molars),  and  large  audital 


1918.]  GENUS   ORYZOMYS.  15 

bullae.  The  principal  reentrant  angles  normally  reach  less  than  half- 
way across  the  molar  crowns.  In  the  moderately  worn  crown  of  the 
second  upper  molar  a  crescentic  central  enamel  island  extends  along 
the  postero-internal  base  of  the  paracone. 

(2)  The  0.  melanotis  group  comprises  slender,  medium-sized,  rich 
ochraceous-buffy  or  ochraceous-tawny  species  with  large  ears  clothed 
externally  with  short,  fine  dusky  hairs  and  internally  with  similarly 
short,  fine  rufescent  hairs.  The  skull  is  narrow  with  elongated  ros- 
trum, short  anterior  palatine  foramina,  and  small  audital  bullae. 
The  dentition  departs  slightly  from  the  0.  palustris  type ;  the  enamel 
arrangement  is  very  similar  but  the  reentrant  angles  extend  farther 
across  the  molar  crowns. 

(3)  The  0.  dlfaroi  group  includes  small,  dark-colored  forms  with 
short  pelage,  comparatively  large,  conspicuous  ears  clothed  exter- 
nally and  internally  with  short,  fine  blackish  hairs.  The  skull  is 
small  and  delicate  in  structure.  The  dentition  is  similar  to  that  of 
the  0.  melanotis  group. 

(4)  The  0.  talamancse  group  bears  much  superficial  resemblance 
to  the  0.  alfaroi  group,  but  the  members  are  usually  brighter,  more 
tawny  in  color.  More  distinctive  characters  are  exhibited  by  the 
skull  and  teeth,  especially  the  molar  crown  arrangement.  In  the 
grinding  surface  of  the  second  upper  molar  the  dentine  ridge  con- 
necting paracone  and  protocone,  owing  to  more  posterior  position, 
eliminates  the  large  central  enamel  island  present  in  the  0.  alfaroi 
group,  and  the  crown,  of  the  third  lower  molar  is  much  more  than 
half  cleft  by  the  outer  reentrant  angle  (about  half  clef t  in  the  0.  alfaroi 
group). 

(5)  The  0.  bombycinus  group  is  easily  recognized  by  very  long 
pelage,  that  of  the  back  measuring  about  12  millimeters.  The  supra- 
orbital vibrissa?  reach  the  remarkable  length  of  50  to  70  millimeters. 
The  dentition  is  about  as  in  the  0.  talamancse,  group. 

(6)  The  0.  devius  group  is  distinguished  by  very  large  but  rather 
slender  form,  relatively  long  tail  (much  longer  than  head  and  body), 
and  dark  general  coloration.  The  dentition  is  similar  to  that  of  the 
0.  talamanCse  and  0.  bombycinus  groups. 

(7)  The  0.  tectus  group  may  be  known  by  large  size,  rich  tawny 
coloration,  small  ears  clothed  with  rather  coarse  hairs  of  general  body 
color,  and  short,  stout  hind  feet.  The  skull  is  broad,  with  short  ros- 
trum and  prominently  projecting  supraorbital  ridges.  The  dentition 
is  much  as  in  the  0.  talamancse  group. 

In  the  present  revision  44  species  and  subspecies  are  assigned  to 
the  typical  subgenus  Oryzomys,  5  forms  are  placed  in  the  subgenus 
Oligoryzomys,  and  2  in  the  subgenus  Melanomys. 


16  NORTH  AMERICAN   FAUNA.  [No.  43. 

List  of  North  American  Species  and  Subspecies,  with  Type  Localities. 

Subgenus  ORYZOMYS. 

Oryzomys  palustris  group: 

Oryzomys  paluslris  palustris  (Harlan) . .  "  Fastland, ' :  near  Salem,  New  Jersey. 

palustris  natator  Chapman Gainesville,  Florida. 

palustris  coloratus  Bangs Cape  Sable,  Florida. 

palustris  texensis  Allen Rockport,  Texas. 

couesi  couesi  (Alston) Coban ,  Guatemala. 

couesi  richmondi  Merriam Escondido  River,  Nicaragua. 

couesi  zygomaticus  Merriam Nenton,  Guatemala. 

couesi  mexicanus  Allen Hacienda  San  Marcos,  Jalisco,  Mexico. 

couesi  aztecus  Merriam Yautepec,  Morelos,  Mexico. 

couesi  crinilus  Merriam Tlalpam,  Federal  District,  Mexico. 

couesi  regillus  Goldman /. . .  Los  Reyes,  Michoacan,  Mexico. 

couesi  albiventer  Merriam Ameca,  Jalisco,  Mexico. 

couesi  peragrus  Merriam Rio  Verde,  San  Luis  Potosi,  Mexico. 

couesi  aquaticus  Allen Brownsville,  Texas. 

fulgens  Thomas "  Mexico. ' ' 

gatuncnsis  Goldman Gatun,  Canal  Zone,  Panama. 

cozumelx  Merriam Cozumel  Island,  Mexico. 

antillarum  Thomas Jamaica. 

peninsulas  Thomas Santa  Anita,  Lower  California,  Mexico. 

nelsoni  Merriam Maria  Madre  Island,  Mexico. 

Oryzomys  melanotis  group: 

Oryzomys  melanotis  melanotis  Thomas.  Mineral  San  Sebastian,  Jalisco,  Mexico. 

melanotis  colimcnsis,  nobis Armeria,  Colima,  Mexico. 

rostratus  rostratus  Merriam Metlaltoyuca,  Puebla,  Mexico. 

rostratus  megadon  Merriam Teapa,  Tabasco,  Mexico. 

rostratus  yucatanensis  Merriam Chichen  Itza,  Yucatan,  Mexico. 

Oryzomys  alfaroi  group : 

Oryzomys  alfaroi  alfaroi  (Allen) San  Carlos,  Costa  Rica. 

alfaroi  dariensis  Goldman Cana,  Panama. 

alfaroi  angusticeps  Merriam Volcan  Santa  Maria,  Guatemala. 

alfaroi  rhabdops  Merriam Calel,  Guatemala. 

alfaroi  caudatus  Merriam Comaltepec,  Oaxaca,  Mexico. 

alfaroi  palatinus  Merriam Teapa,  Tabasco,  Mexico. 

alfaroi  saturatior  Merriam Tumbala,  Chiapas,  Mexico. 

alfaroi  cliapmani  Thomas Jalapa,  Vera  Cruz,  Mexico. 

alfaroi  dilutior  Merriam Huauchinango,  Puebla,  Mexico. 

guerrerensis  Goldman Omilteme,  Guerrero,  Mexico. 

hylocetes  Merriam Chicharras,  Chiapas,  Mexico. 

Oryzomys  talamancss  group: 

Oryzomys  talamancee  Allen Talamanca,  Costa  Rica. 

Oryzomys  bombycinus  group: 

Oryzomys       bombycinus       bombycinus 
Goldman Cerro  Azul,  Panama. 

bombycinus  alleni  Goldman Tuis,  Costa  Rica. 

Oryzomys  devius  group: 

Oryzomys  devius  Bangs Boquete,  Panama. 

pirrensis  Goldman Mount  Pirre,  Panama. 

Oryzomys  tectus  group : 

Oryzomys  tectus  tectus  Thomas Bugaba,  Panama. 

tectus  frontalis  Goldman Corozal,  Canal  Zone,  Panama. 

Oryzomys  victus  1  Thomas St.  Vincent,  Lesser  Antilles. 

iNot  examined  and  group  association  not  determined. 


1918.]  SUBGENUS   ORYZOMYS.  17 

Subgenus  OLIGORYZOMYS. 

Oryzomys  fulvescens  fulvescens  (Saussure)..  Orizaba,  Vera  Cruz.  Mexico. 

fulvescens  lenis  Goldman Loa  Reyes,  Michoacan,  Mexico. 

fulvescens  mayensis,  nobia Apazote,  Campeche,  Mexico. 

fulvescens  costaricensis  Allen El  General,  Coata  Rica. 

fulvescens  vegetus  Bangs Boquete,  Panama. 

Subgenus  MELANOMYS. 

Oryzomys  caliginosus  idoneus  Goldman Cerro  Azul.  Panama. 

caliginosus  chrysomelas  Allen Suerre,  Costa  Rica. 

Key  to  Subgenera. 

a1.  Lachrymal  articulating  about  equally  with  frontal  and  maxilla  anteriorly;  tail 
about  equal  to  or  longerthan  head  and  body. 
b1.  Second  upper  molar  with  central  enamel  island  normally  elongated  or  absent; 
supraorbital  and  temporal  ridgea  present;  hind  foot  usually  more  than 

25 Oryzomys  (p.  17). 

b2.  Second  upper  molar  with  central  enamel  island  normally  circular;  supraorbital 
and  temporal  ridgea  absent;  hind  foot  usually  less  than  25. 

Oligoryzomys  (p.  87). 
a?.  Lachrymal  articulating  almost  entirely  with  maxilla  anteriorly;  tail  about  three- 
fourths  length  of  head  and  body Melanomys  (p.  94). 

Subgenus  ORYZOMYS  Baird. 

Subgeneric  characters. — Color  of  upperparts  usually  contrasting 
strongly  with  that  of  underparts;1  feet  (epidermis  and  hairs),  includ- 
ing toes,  whitish,  yellowish,  or  brownish;1  tail  about  equal  to  or 
longer  than  head  and  body;1  anterior  border  of  lachrymal  articulating 
about  equally  with  maxilla  and  frontal;1  supraorbital  and  temporal 
ridges  usually  prominent;2  secondary  parastyle  well  developed;1 
slightly  worn  crown  of  second  upper  molar  with  central  enamel  island 
elongated  or  absent;2  upper  incisors  decidedly  curved  backward  near 
points.1 

Key  to  Species  of  the  Subgenus  Oryzomys. 
[Typical  adults.] 

a1.  Habitat  North  America  and  Greater  Antilles. 
b1.  Hind  foot  without  prominent  tufts  of  digital  bristles  projecting  beyond  ends  of 
three  median  claws. 
c1.  Habitat  Rio  Grande  Valley  south  to  Panama,  and  including  islands  near 
coasts  of  Mexico  and  West  Indies. 
d1.  Habitat  continental. 

el.  Supraorbital  ridges  not  projecting  prominently  over  orbita. 
f1.  Head  and  shoulders  not  distinctly  grayiah. 

gl.  Upperparts  richer  ochraceous-tawny.  ("Mexico.").  0.fulgens(p.41). 
g2.  Upperparts  duller  ochraceous-tawny  or  ochraceous-buffy.     (Southern 

Texas  to  Costa  Rica. ) O.  couesi  (p.  28) . 

f2.  Head  and  shoulders  distinctly  grayish.      (Lower  California.) 

O.  peninsulse  (p.  45). 

i  Contrasting  with  Melanomys,  » Contrasting  with  Oligoryzomys* 

14521°— 18— 2 


18  NORTH   AMERICAN    FAUNA.  [No.  43. 

e2.  Supraorbital  ridges  projecting  prominently  over  orbits.    (Panama.) 

O.  gatunensis  (p.  42). 
d2.  Habitat  insular. 
e1.  Habitat  islands  off  coasts  of  Mexico. 
P.  TJpperparts ochraceous-buffy .  (Maria Madre Island.).  0.nelsoni(p.46). 
P.  Upperparts  ochraceous-tawny.  (Cozumel Island.).  O. cozumelae (p. 43). 

e2.  Habitat  Greater  Antilles.     (Jamaica.)...^ O.  antillarum  (p.  44). 

c2.  Habitat  United  States  north  of  Rio  Grande  Valley O.  palustris  (p.  21). 

b2.  Hind  foot  with  prominent  tufts  of  digital  bristles  projecting  beyond  ends  of  three 
median  claws. 
c1.  Ears  clothed  externally  with  fine  blackish  hairs  contrasting  with  color  of  head; 
hind  foot  long  and  narrow. 
d1.  Hind  foot  less  than  33. 
e1.  Supraorbital  vibrissse  less  than  40. 

*.  Ears  clothed  internally  with  fine  blackish  hairs. 

gl.  Second  upper  molar  with  central  enamel  island  present ;  third  lower 
molar  with  outer  reentrant  angle  extending  about  half  way  across 
crown. 
ft1.  Upperparts  darker  ochraceous-buff  or  ochraceous-tawny. 
i1.  Zygomata   not  wider   anteriorly  than   posteriorly,  or   if  wider 
zygomatic  breadth  more  than  14.     (Atlantic  slope  in  eastern 
and  southern  Mexico  and  south  to  Panama.) 

O.   alfaroi  (p.  58). 
i2.  Zygomata  wider  anteriorly  than  posteriorly;  zygomatic  breadth 
14  or  less.     (Pacific  slope  of  mountains  of  southern  Chiapas.) 

O.  hylocetes  (p.  70). 
ft2.  Upperparts  paler  ochraceous-buff  or  ochraceous-tawny.     (Pacific 
slope  of  Sierra  Madre  in  Guerrero  and  Oaxaca.) 

O.  guerrerensis  (p.  69). 

g2.  Second  upper  molar  with  central  enamel  island  absent;  third  lower 

molar  with  outer  reentrant  angle  extending  more  than  hah  way 

across  crown.     (Costa  Rica  and  Panama. ).0  .  talamancse  (p.  73). 

P.  Ears  clothed  internally  with  buffy  or  rusty  reddish  hairs. 

g1.  Size  larger;  hind  foot  30  or  more.     (Atlantic  coast  of  Mexico.) 

O.  rostratus  (p.  52). 
g2.  Size  smaller;  hind  foot  less  than  30.     (Pacific  coast  of  Mexico.) 

O.  melanotis  (p.  49). 

e2.  Supraorbital  vibrissse  more  than  50 O.  bombycinus  (p.  76). 

d2.  Hind  foot  33  or  more. 
e1.  Color  paler;  supraorbital  ridges  absent.     (Costa  Rica  and  western  Panama.) 

O.  devius  (p.  80). 
e2.  Color  darker;  supraorbital  ridges  present.     (Eastern  Panama.) 

O.  pirrensis  (p.  81). 
c2.  Ears  clothed  externally  with  coarse  tawny  hairs  not  contrasting  with  color  of 
head;  hind  foot  short  and  broad.     (Costa  Rica  and  Panama.) 

O.  tectus  (p.  84). 
a2.  Habitat  Lesser  Antilles.     (St.  Vincent.) O.  victus  (p.  86). 

Oryzomys  palustris  Group. 

Geographic  distribution. — Coastal  areas  from  southern  New  Jersey 
to  southern  Texas ;  north  in  the  Mississippi  Valley  to  southern  Illinois, 
and  southward  from  the  Rio  Grande  Valley  on  the  east,  and  southern 


1918.]  ORYZOMYS  PALUSTRIS   GROUP.  19 

Sinaloa  on  the  west,  through  Middle  America  to  Panama,  with  out- 
lying forms  inhabiting  southern  Lower  California,  the  Tres  Marias 
Islands,  and  Cozumel  Island.  Confined  mainly  to  the  vicinity  of 
water  at  low  elevations,  but  ranging  up  to  over  7,000  feet  altitude  in 
the  marshy  bottom  of  the  Valley  of  Mexico  (see  maps,  figs.  2  and  3). 

General  characters. — Size  large;  form  robust;  tail  usually  equal  to 
or  longer  than  head  and  body  (shorter  in  some  examples),  thinly  but 
rather  distinctly  haired;  ears  small  and  inconspicuous,  well  haired 
internally  as  well  as  externally;  general  pelage  long,  rather  coarse  and 
rigid;  the  underfur  somewhat  woolly;  vibrissas  short,  the  longest 
arising  from  muzzle  scarcely  as  long  as  head;  hind  feet  broad,  the 
upper  surface  rather  well  haired  and  under  surface  naked  and  coarsely 
granular  anteriorly,  becoming  smooth  along  outer  side  of  large  pos- 
terior tubercle ;  inner  edge  of  plantar  surface  overlapped  by  fringing 
bristles ;  toes  of  hind  feet  webbed  near  base ;  the  claws  long,  relatively 
straight,  and  projecting  well  beyond  overlapping  bristles.  Color  of 
upperparts  presenting  a  wide  range  of  variation  from  grizzled  grayish 
brown,  or  pale  buff,  to  rich  ochraceous-buff  or  ochraceous-tawny,  more 
or  less  heavily  overlaid  with  black;  underparts  ranging  from  white  to 
light  ochraceous-buff. 

Slcull. — Size  large,  with  rostrum  short  and  braincase  high  and  well 
arched ;  outer  wall  of  antorbital  foramen  with  superior  border  extend- 
ing well  forward,  the  foramen  appearing  as  a  deep  circular  notch  as 
viewed  from  above;  frontal  region  rather  broad,  the  lateral  margins 
trenchant,  somewhat  upturned,  and  projecting  as  supraorbital  ridges, 
frontals  usually  encroaching  in  a  narrow  point  posteriorly  along  the 
median  line  between  the  parietals;  temporal  ridges  well  developed 
anteriorly  along  parieto-squamosal  borders,  usually  becoming  indis- 
tinct posteriorly  in  crossing  lateral  wings  of  parietals  to  low  lambdoid 
crest;  interparietal  small,  sub  triangular,  the  anterior  border  a  nearly 
straight  fine  and  the  posterior  with  an  ill-defined  median  angle;  ante- 
rior palatine  foramina  narrow  and  much  elongated,  about  equal  in 
length  to  palatal  bridge,  normally  reaching  posteriorly  to  anterior 
plane  of  first  molars,  the  median  septum  with  posterior  or  maxillary 
portion  contracted  and  anterior  or  premaxillary  section  expanded 
above;  palatal  pits  large  and  normally  oval  in  outline;  interpterygoid 
fossa  moderately  broad;  sphenopalatine  vacuities  large  in  0.  palus- 
tris,  absent,  or  present  as  very  narrow  slits,  in  0.  couesi  and  related 
forms;  audital  bullae  large,  the  swollen  portion  projecting  anteriorly 
beyond  anterior  plane  of  basioccipital ;  basioccipital  narrow;  angle  of 
mandible  rather  broad  and  projecting  posteriorly;  coronoid  process 
large,  rising  high  above  condyle;  dentition  moderately  heavy;  third 
lower  molar  rather  short  and  broad;  mandibular  toothrow  only 
slightly  narrower  posteriorly  than  anteriorly;  inner  reentrant  angles 
in  upper  molars  and  outer  reentrant  angles  in  lower  molars  reaching 


20  NORTH  AMERICAN    FAUNA.  [No.  43. 

less  than  half  way  across  moderately  worn  crowns;  second  upper 
molar  with  a  somewhat  crescentic  enamel  island,  or  furrow,  along 
postero-internal  base  of  paracone,  becoming  restricted  and  finally 
obliterated  through  extended  wear;  tubercle  over  root  of  lower  incisor 
prominent. 

Remarks. — The  0.  palustris  group  includes  0.  palustris  of  the  United 
States,  and  0.  couesi  and  nearly  related  Middle  American  forms. 
Aside  from  the  darker,  more  brownish  colors  which  usually  charac- 
terize 0.  palustris,  in  contrast  with  the  brighter,  more  ochraceous 
buffy  or  rufescent  tones  of  0.  couesi  and  its  relatives,  these  sections  of 
the  group  also  differ  notably  in  cpality  of  pelage,  the  overfur  being 
longer  and  projecting  farther  beyond  the  underfur  in  the  former  than 
in  the  latter.  The  forms  of  the  two  sections  of  the  group  agree  closely 
in  essential  cranial  details,  but  skulls  of  subspecies  of  0.  palustris  are 
usually  recognizable  by  the  large  size  of  the  sphenopalatine  vacuities. 
Members  of  the  group  as  a  whole  are  distinguished  externally  from 
those  of  other  Middle  American  groups  of  the  same  subgenus  Ly  the 
small  size  and  internal  as  well  as  external  hairiness  of  the  ears. 

Key  to  Species  and  Subspecies  of  the  O.  palustris  Group. 

[Typical  adults.] 

a1.  Upperparts   mainly  grayish    or  brownish,  or  if  ochraceous-buffy  or  ochraceous- 
tawny  underparts  not  distinctly  buffy.     (United  States  north  of  Rio  Grande 
Valley  [0.  palustris].) 
b1.  Upperparts  mainly  grayish  or  brownish. 
c1.  Color  darker.     (Atlantic  coast  region  from  southern  New  Jersey  to  northern 
Florida;   Alabama;  southeastern  Mississippi;   central  Tennessee;   southern 
Kentucky;    southern  Illinois,  and  parts  of  southeastern  Missouri.) 

O.  p.  palustris  (p.  22). 

c2.  Color  paler.     (Coast  region  of  Texas  from  Nueces  Bay  northward;  Louisiana; 

western  Mississippi;  southern  and  eastern  Arkansas;  extreme  southeastern 

Missouri;  southeastern  Kansas.) O.  p.  texensis  (p.  27). 

b2.  Upperparts  mainly  clay  color  or  ochraceous-tawny. 

c1.  Color  duller,  less  distinctly  ochraceous-tawny.     (North-central  Florida.) 

O.  p.  natator  (p.  24). 
c2.  Color  brighter,  more  distinctly  ochraceous-tawny.     (Southern  Florida.) 

O.  p.  coloratus  (p.  26). 
a2.  Upperparts  mainly  ochraceous-buffy  or  ochraceous-tawny.     (Rio  Grande  Valley 
to  Panama  and  islands  near  coasts  of  Mexico  and  West  Indies  [0.  couesi  and 
related  forms].) 
b1.  Habitat  continental. 
c1.  Supraorbital  ridges  not  projecting  prominently  over  orbits. 
dl.  Head  and  shoulders  not  distinctly  grayish. 

e1.  Upperparts    duller  ochraceous-tawny    or    ochraceous-buffy.     [0.   couesi 
and  subspecies.] 
f1.  Underparts  normally  white. 

g1.  Size  larger;  hind  foot  usually  35  or  more;  upper  molar  series  usually 
more  than  5. 


1918.] 


OKYZOMYS   PALUSTBIS   GROUP.  21 


h1.  Upperparts  more  intense  ochraceous-buff.     (Northwestern  Michoa- 

can) O.  c.  regillus(p.  37). 

h2.  Upperparts  less  intense  ochraceous-buff.     (Central  Jalisco.) 

O.  c.  albiventer  (p.  38). 
g2.  Size  smaller;    hind  foot  usually  less  than  35;    upper  molar  series 
usually  less  than  5. 
hl.  Upperparts  darker  ochraceous-buff.     (Pacific  coastal  region  from 
southern  Sinaloa  to  southeastern  Oaxaca.) 

O.  c.  mexicanus  (p.  33). 
h2.  Upperparts    paler    ochraceous-buff.     (Morelos;    southern  Puebla; 
northern  Oaxaca;  northeastern  Guerrero.).  O.  c.  aztecus  (p.  35). 
f2.  Underparts  normally  buffy. 
g1.  Size  larger;    hind  foot  usually  more  than  33;    upper  molar  series 
usually  more  than  4.8. 
h1.  Size  smaller;  hind  foot  averaging  less  than  35. 

i  K  Color  darker.     (San  Luis  Potosi.) O.  c.  peragrus  (p.  39). 

i2.  Color  paler.     (Rio  Grande  Valley.). .  O.  c.  aquaticus  (p.  39). 
h2.  Size  larger;   hind  foot  averaging  about  36.     (Valley  of  Mexico.) 

O.  c.  crinitus  (p.  36). 
g2.  Size  smaller;    hind  foot  usually  less  than  33;    upper  molar  series 
usually  more  than  4.8. 
h1.  Upperparts  normally  ochraceous-buffy. 

i1.  Color  darker.     (Northern  Vera  Cruz   to  northwestern  Costa 

Rica.) O.  c.  couesi  (p.  29). 

i2.  Color   paler.     (Southwestern    Gautemala  and    south-centra] 

Chiapas.) O.  c.  zygomaticus(p.  32). 

h2.  Upperparts  normally   ochraceous-tawny.     (Lowlands  of   eastern 

Nicaragua.) O.  c.  richmondi  (p.  32). 

e2.  Upperparts  richer  ochraceous-tawny.    (Mexico.) O.  fulgens(p.  41). 

cP.  Head  and  shoulders  distinctly  grayish.     (Lower  California.) 

O.  peninsulee  (p.  45). 
c2.  Supraorbital  ridges  projecting  prominently  over  orbits.     (Panama.) 

O.  gatunensis  (p.  42). 

b2.  Habitat  insular. 

c1.  Habitat  off  east  coast  of  Mexico.     (Cozumel  Island.). .  O.  cozumelse  (p.  43). 

c2.  Habitat  off  west  coast  of  Mexico.     (Maria  Madre  Island.).  O.  nelsoni  (p.  46). 

c3.  Habitat  West  Indies.     (Jamaica.) O.  antillarum  (p.  44). 

ORYZOMYS  PALUSTRIS  (Harlan). 
[Synonymy  under  subspecies.] 

Geographic  distribution. — Atlantic  and  Gulf  coastal  areas  from 
southern  New  Jersey  (not  yet  recorded  from  Delaware  and  Maryland, 
but  doubtless  occurs  there),  to  southern  Texas,  and  north  through  the 
Mississippi  Valley  to  southern  Kentucky,  southern  Illinois,  and 
eastern  Kansas  (fig.  2).  Altitudinal  range  from  sea  level  up  along 
streams  to  about  500  feet  altitude  (rarely  to  1,000  feet),  mainly  in  the 
Lower  Austral  Zone,  but  reaching  into  the  Upper  Austral  Zone  in  the 
more  northerly  localities,  and  into  the  Tropical  Zone  in  southern 
Florida. 

General  characters. — Similar  in  general  to  O.  couesi,  but  pelage 
longer;  colors  usually  darker  and  duller  grayish  brown  instead  of 


22 


NORTH   AMERICAN    FAUNA. 


[No.  43. 


ochraceous-buffy    or   ochraceous-tawny;   skull    differing   in    various 
details,  but  dentition  about  the  same. 

Color. — Upperparts  in  general  varying  from  grizzled  grayish  brown 
or  pale  buff,  to  tawny-olive,  clay  color,  and  ochraceous-tawny;  the 
face,  top  of  head,  and  back  heavily  lined  or  overlaid  with  black; 
sides  paler,  owing  to  a  thinner  admixture  of  blackish  hairs;  under- 
pays white,  varying  to  buffy  white  and  rarely  to  pale  buff;  outer 
sides  of  ears  dusky,  the  inner  sides  clothed  with  grayish  or  rusty  hairs; 
feet  whitish ;  tail  varying  from  brownish  above  and  whitish  below  to 
dusky  all  round.      Young  (in  first  pelage):  Varying  from  grayish 

brown  to  dull  tawny- 
olive  above,  whitish 
below. 

STcull. — (For  gen- 
eral outlines  see  under 
0.  palustris  group.) 
In  general  form  closely 
resembling  that  of  0. 
couesi,  but  sphenopa- 
latine vacuities,  ab- 
sent or  much  reduced 
in  size  in  that  species, 
large  and  widely  open; 
antorbital  foramen 
with  anterior  border 
less  rounded  above, 
less  inclined  or  pro- 
duced forward  at  the 
base,  in  many  exam- 
ples somewhat  excised 
or  tending  to  develop 
a  point  as  in  Sigmodon. 
Remarks. — 0.  palustris  is  divisible  into  four  geographic  races  which 
form  a  closely  intergrading  series.  The  species  apparently  attains 
its  largest  size  in  Florida  and  along  the  coast  of  the  Gulf  of  Mexico  east 
of  the  Mississippi  delta. 

ORYZOMYS  PALUSTRIS  PALUSTRIS  (Harlan). 

Swamp  Rice  Rat. 

[PI.  I,  figs.  1,  la;  PI.  V,  figs,  1,  4;  PI.  VI,  figs.  1,  la.] 

Mus  palustris  Harlan,  Silliman's  Amer.  Journ.  Sci.,  XXXI,  p.  38!'),  1837. 

Arvicola  oryzivora,  Bachman,  in  Audubon  and  Bachman,  Quadr.  North  Amer.,  Ill, 

p.  214,  1854.     Type  from  St.  Johns  Parish,  South  Carolina. 
IJ[esperomys]  palustris  Wagner,  in  Suppl.  Schreber's  Saugthiere,   III,  p.  543,  1843. 
Oryzomys  palustris  Baird,  Mamm.  North  Amer.,  p.  459,  1857. 


1.  O. p. palustris.    2.  O.p.natator.    3.  O.p.coloratus.    4.  O.p.texensis- 
Fig.  2.— Geographic  distribution  of  subspecies  of  Oryzomys  palustris. 


1918.]  ORYZOMYS  PALUSTEIS  GROUP.  23 

Type  locality. — "Fastland/'  near  Salem,  Salem  County,  New  Jersey. 

Type. — Not  known  to  exist. 

Geographic  distribution. — Atlantic  coastal  areas  from  southern  New 
Jersey  (not  yet  known  from  Delaware  or  Maryland,  but  doubtless 
occurs  there)  south  to  northeastern  Florida,  thence  westward  through 
southern  Georgia  to  the  Gulf  coast  of  Alabama  and  Mississippi,  and 
north  through  Alabama  and  western  Tennessee  to  southwestern  Ken- 
tucky, southern  Illinois,  and  parts  of  southeastern  Missouri.  Alti- 
tudinal  range  from  sea  level  up  along  streams  to  about  500  feet  (rarely 
to  1,000  feet),  mainly  in  Lower  Austral  Zone,  but  reaching  into  Upper 
Austral  Zone  in  southern  New  Jersey,  southeastern  Kentucky,  and 
southeastern  Missouri  (Marble  Hill). 

General  characters. — Size  usually  smaller  and  color  more  brownish, 
less  tawny,  than  in  0.  p.  natator  and  0.  p.  coloratus;  skull  less  massive. 
Closely  resembling  0.  p.  texensis,  but  darker,  more  brownish  than 
topotypes  of  the  latter  form;  skull  broader. 

Color. — Fresh  pelage  (December):  Upperparts  grizzled  grayish 
brown  or  pale  buff,  the  brownish  or  buffy  tone  most  intense  on  rump, 
darkened  on  face,  top  of  head,  and  back  by  overlying  blackish  hairs; 
feet  whitish;  tail  brownish  above,  whitish  below,  becoming  in  some 
specimens  dark  all  round  near  tip.  Young  (in  first  pelage) :  Grayish 
brown  above,  dull  whitish  below. 

Skull. — (For  general  outlines  see  under  O.  palustris  group.)  Very 
similar  to  those  of  0.  p.  natator  and  O.  p.  coloratus,  but  narrower; 
braincase  decidedly  narrower;  zygomata  less  widely  spreading; 
frontal  region  narrower,  the  supraorbital  borders  less  projecting. 
Compared  with  that  of  0.  p.  texensis  the  skull  differs  mainly  in  some- 
what larger  average  size;  zygomata  usually  more  widely  spreading. 

Measurements. — Average  of  four  adults  from  Greenwich,  N.  J.  (near 
type  locality):  Total  length,  242  (237-245);  tail  vertebrae,  112  (109- 
116);  hind  foot,  31  (30-31.5).  An  adult  from  Pope  Creek,  Va.:  260; 
130;  33.  Average  of  four  adults  from  Georgetown,  S.  C:  257  (233- 
273);  125  (113-132);  32.5  (31-33).  Average  of  three  adults  from 
Bon  Secour,  Ala.:  265  (250-280);  131  (125-138);  30.5  (30-31.5). 
Adult  from  Bayou  La  Batre,  La.:  233;  116;  30.  Adult  from  Marble 
Hill,  Mo.:  252;  115;  29.  Adult  from  Olive  Branch,  111.:  255;  113 
29.  Adult  from  Barbourville,  Ky.:  270;  133;  30.5.  Skull  (two 
adults  from  Greenwich,  N.  J.:  Greatest  length,  32.4,  31;  zygomatic 
breadth,  17.2,  16.6;  interorbital  breadth,  5.2,  5.1;  width  of  braincase, 
11.9,  12;  nasals,  12.7,  12;  anterior  palatine  foramina,  6.7,  6.7; 
palatal  bridge,  6.2,  6.2;  upper  molar  series,  4.5,  4.6. 

Remarks. — 0.  p.  palustris  passes  into  0.  p.  texensis  in  the  Mississippi 
Valley.  Specimens  from  Marble  Hill,  Mo.,  and  Olive  Branch,  111., 
however,  seem  referable  to  the  typical  form.     A  tendency  to  develop 


24  NORTH   AMERICAN   FAUNA.  [No.  43. 

the  ruddy  color  and  more  massive  skull  of  0.  p.  natator  is  exhibited  by 
specimens  from  New  Berlin  and  Burnside  Beach,  northern  Florida, 
and  intergradation  of  the  two  forms  seems  evident  in  that  part  of  the 
State.  No  rice  rats  have  been  recorded  from  Delaware  or  Maryland, 
but  the  favorable  character  of  the  country  and  the  narrowness  of  the 
gap  between  collecting  stations  to  the  north  and  south  point  to 
probable  continuity  of  range.  Specimens  from  South  Carolina 
apparently  representing  Arvicola  oryzivora  of  Audubon  and  Bachman 
are  inseparable  from  typical  palustris. 

Specimens  examined. — Total  number,  233,  as  follows: 

Alabama:  Autaugaville,  1;  Bayoi/La  Batre,  2;  Bon  Secour,  3;  Elmore,  1;  Flor- 
ence, 1;  Gallion,  1;  Hayneville,  2;  Huntsville,  4;  Jackson,  2;  Mobile,  1;  Mont- 
gomery, 1;  Mount  Weogufka,  1;  Reform,  1;  Sand  Mountain  (near  Carpenter), 
1;  Seale,  1. 

Florida:  Burnside  Beach,  9;1  New  Berlin,  8.2 

Georgia:  Cumberland  Island,  14;3>  i  Hursman  Lake,  l;1  Mcintosh  County,  1; 
Okefinokee  Swamp,  l;2  Ossabaw  Island,  12;1  Riceboro,  7;5  Saint  Marys,  20;3>  6 
Savannah,  10;  Toccoa,  2. 

Illinois:  Olive  Branch,  3. 

Kentucky:  Barbourville,  3. 

Missouri:  Marble  Hill,  1. 

Mississippi:  Biloxi,  1. 

New  Jersey:  Cedar  Creek,  l;7  Greenwich,  13. 8 

North  Carolina:  Coinjock,  1;  Pea  Island,  2;1  Raleigh,  31.9» 10 

South  Carolina:  Beaufort  County,  2;  Calhoun  Falls,  2;2  Easley,  1;  Frogmore, 
l;3  Georgetown,  11;  Plantersville,  7;  Saint  Helena  Island,  1;  Society  Hill,  3. 

Tennessee:  High  Cliff,  1;  Lawrenceburg,  2. 

Virginia:  Dismal  Swamp,  20;  Pope  Creek  (5  miles  southeast  of  Colonial 
Beach),  2;  Smith  Island,  6;  Suffolk,  1;  Wallops  Island,  3;  Warsaw  (4  miles 
southwest),  5;  Wreck  Island,  1. 

ORYZOMYS  PALUSTRIS  NATATOR  Chapman. 

Central  Florida  Rice  Rat. 

Oryzomys  palustris  natator  Chapman,  Bull.  Amer.  Mus.  Nat.  Hist.,  V.,  p.  44,  March 
17,' 1893. 

Type  locality. — Gainesville,  Alachua  County,  Florida. 

Type. — No.  }%%%,  <$  adult,  American  Museum  of  Natural  History; 
collected  by  F.  M.  Chapman,  January  31,  1889. 

Geographic  distribution. — Central  Florida,  north  of  Everglades; 
Austro riparian  division  of  Lower  Austral  Zone. 

1  Collection  Mus.  Comp.  Zool.  7  Collection  Acad.  Nat.  Sci.  Philadelphia. 

2  Collection  Field  Mus.  Nat.  Hist.  8  Five  in  collection  Acad.  Nat.  Sci.  Philadelphia. 
»  Collection  Amer.  Mus.  Nat.  Hist.  9  Five  in  collection  Amer.  Mus.  Nat.  Hist. 

<  Eleven  in  collection  Mus.  Comp.  Zool.  10  Two  in  collection  Field  Mus.  Nat.  Hist.;  6  in 

6  Five  in  collection  Field  Mus.  Nat.  Hist.  Mus.  Comp.  Zool. 

6  Three  in  collection  Field  Mus.  Nat.  Hist.;  15  in 
Mus.  Comp.  Zool. 


1918.]  ORYZOMYS   PALUSTEIS   GROUP.  25 

General  characters. — Most  like  0.  p.  coloratus;  differing  usually  in 
less  intense  tawny  suffusion  of  upperparts,  especially  cheeks  and  sides 
of  body.  Size  larger,  color  more  tawny,  and  skull  more  massive  than 
usual  in  0.  p.  palustris  or  0.  p.  texensis. 

Color. — Fresh  pelage  (December) :  Upperparts  varying  from  grizzled 
grayish  brown  or  pale  buff  to  tawny-olive,  clay  color,  or  ochraceous- 
tawny,  deepest  and  richest  on  lower  part  of  back  and  rump,  becoming 
paler  and  more  buffy  on  sides,  and  darkened  dorsally  by  admixture 
of  blackish  hairs ;  face  grayish  or  pale  buffy  beneath  overlying  dusky 
hairs;  underparts  usually  white,  but  in  some  specimens  more  or  less 
suffused  with  pale  buff;  feet  white;  tail  brownish  above,  whitish  be- 
low, becoming  in  some  specimens  dark  all  round  toward  tip.  Young 
(in  first  pelage) :  Brownish  or  dull  tawny-olive  mixed  with  black  above, 
whitish  below. 

STcull. — Similar  to  those  of  0.  p.  palustris  and  0.  p.  texensis,  but 
broader;  braincase  decidedly  broader;  zygomata  more  widely  spread- 
ing; frontal  region  broader,  the  supraorbital  borders  more  projecting. 
Comparison  with  the  skull  of  0.  p.  coloratus  reveals  no  appreciable 
difference. 

Measurements. — Type:  Total  length,  295;  tail  vertebrae,  143;  hind 
foot  (dry  skin),  33.  Average  of  10  adults  (type  and  9  topotypes): 
276.7  (271-300);  142.9  (132-156);  34.3  (32.5-37).  SJcull  (average  of 
6  adults,  type  and  5  topotypes):  Greatest  length,  32.4  (31.4-33.7); 
zygomatic  breadth,  17.2  (16.4-17.7);  interorbital  breadth,  5.4  (5.1- 
6.1);  width  of  braincase,  12.4  (12-12.8);  nasals,-  12.7  (12-13.5);  an- 
terior palatine  foramina,  6.9  (6.4-7.2);  palatal  bridge,  6.1  (5.7-6.5); 
upper  molar  series,  4.7  (4.5-4.9). 

Remarks. — In  northern  Florida,  not  far  to  the  northward  of  the 
type  locality,  0.  p.  natator  passes  into  0.  p.  palustris,  as  shown  by 
specimens  from  New  Berlin  and  Burnside,  which,  however,  seem  more 
properly  placed  with  the  latter  form.  In  the  vicinity  of  Lake  Oke- 
chobee  natator  merges  with  0.  p.  coloratus,  a  richer  colored  form 
inhabiting  the  southern  part  of  the  State. 

Specimens  examined. — Total  number,  121,  as  follows: 

Florida:  Anastasia  Island,  2;1  Canaveral,  4;  Cape  Canaveral,  3;  Cartersville,  1; 
Crystal  River,  l;1  Enterprise,  26 ;2  Espanita,  3;3  Fort  Kissimmee,  1;  Gaines- 
ville, 19  (type  and  topotypes);4  Geneva,  1;  Kissimmee,  2;  Kissimmee  River, 
2;  Lake  Harney,  11;  Lake  Kissimmee,  19;  Micco,  9;5  Mullet  Lake,  1;  Oak 
Lodge,  9;1  Ocala,  2;  Tarpon  Springs,  l;6  Titusville,  4. 

i  Collection  Mus.  Comp.  Zool. 

2  Fourteen  in  collection  Amer.  Mus.  Nat.  Hist.;  10  in  Field  Mus.  Nat.  Hist.;  2  in  Mus.  Comp.  Zool. 

s  Collection  Field  Mus.  Nat.  Hist. 

*  Nine  in  collection  Amer.  Mus.  Nat.  Hist.;  5  in  Field  Mus.  Nat.  Hist.;  5  in  Mus.  Comp.  Zool. 

6  Three  in  collection  Amer.  Mus.  Nat.  Hist.;  3  in  Field  Mus.  Nat.  Hist. 

«  Collection  Acad.  Nat.  Sci.  Philadelphia. 


26  NORTH   AMERICAN   FAUNA.  [No.  43. 

ORYZOMYS  PALUSTRIS  COLORATUS  Bangs. 

Everglades  Rice  Rat. 

(PI.  I,  figs.  2,  2a.) 

Oryzomys  palustris  coloratus  Bangs,  Proc.  Boston  Soe.  Nat.  Hist.,  XXVIII,  p.  189, 
March,  1898. 

Oryzomys  natator  floridanus  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  277,  July 
26,  1901.  Type  from  Everglade,  Florida,  No.  71349,  $  ad.,  U.  S.  Nat.  Mus.  (Bio- 
logical Survey  collection);  collected  by  J.  Alden  Loring,  March  29,  1895. 

Type  locality. — Cape  Sable,  Monroe  County,  Florida. 

Type. — No.  4470,  <?  adult,  Museum  of  Comparative  Zoology  (col- 
lection of  E.  A.  and  O.  Bangs) ;  collected  by  C.  L.  Brownell,  April  17, 
1895. 

Geographic  distribution. — Tropical  southern  Florida,  north  to  Lake 
Okechobee. 

General  characters. — Closely  resembling  0.  p.  natator;  differing  in 
more  intense  tawny  suffusion  of  upperparts;  size  about  the  same. 
Size  larger  than  usual  in  0.  p.  palustris  and  0.  p.  texensis,  and  color 
much  more  tawny  than  either. 

Color. — About  as  in  0.  p.  natator,  but  general  tone  of  upperparts 
slightly  richer,  more  tawny  or  rufescent. 

STcull. — Like  that  of  0.  p.  natator. 

Measurements. — Type:  Total  length,  301;  tail  vertebrae,  150;  hind 
foot,  35.  Average  of  three  adult  topotypes:  296  (278-305);  144 
(133-152);  33.4  (33.4-33.4).  STcull  (average  of  four  adults,  type 
and  three  topotypes):  Greatest  length,  32.2  (31.8-32.7);  zygomatic 
breadth,  17.1  (16.8-17.5);  interorbital  breadth,  5.8  (5.8-5.9);  width 
of  braincase,  12.4  (12.2-12.5);  nasals,  12.4  (12.1-12.6);  anterior 
palatine  foramina,  6.6  (6.3-7);  palatal  bridge,  6.1  (5.7-6.6);  upper 
molar  series,  4.8  (4.7-4.9). 

Remarks. — O.  p.  coloratus  requires  close  comparison  with  0.  p. 
natator,  from  which  it  apparently  differs  only  in  color.  The  richer 
tone  in  coloratus  is  most  noticeable  when  specimens  are  turned  on 
their  sides,  and  the  cheeks  and  flanks  contrasted  with  those  of  exam- 
ples of  natator.  As  Bangs  rightly  states,  coloratus  "occupies  only 
the  southern,  tropical  part  of  the  Florida  peninsula." 

In  describing  "Oryzomys  natator  fioridanus,^  Merriam  overlooked 
the  name  coloratus,  which  had  already  been  applied  to  the  animal 
of  the  region;  the  two  are  clearly  synonymous. 

Specimens  examined. — Total  number,  50,  as  follows: 

Florida:  Cape  Sable,  11  (type  and  topotypes);1  Eden,  1;  Everglade,  16  (includ- 
ing type  of  "floridanus");  Flamingo,  13 ;2  Juno  (Lake  Worth),  5;  Jupiter^;1 
Miami,  1;  Miami  River,  1. 

i  Collection  Mus.  Comp.  Zool.       2  Twelve  in  collection  Mus.  Comp.  Zool.;  1  in  Amer.  Mus.  Nat.  Hist. 


1918.]  ORYZOMYS   PALUSTRIS   GROUP.  27 

ORYZOMYS  PALUSTRIS  TEXENSIS  Allen. 

Texas  Rice  Rat. 

Oryzomys  palustris  texensis  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  VI,  p.  177,  May  31, 
1894. 

Type  locality. — Rockport,  Aransas  County,  Texas. 

Type. — No.  ^-ftf,  c?  ad.,  American  Museum  of  Natural  History; 
collected  by  H.  P.  Attwater,  November  15,  1893. 

Geographic  distribution. — From  Corpus  Christi  Bay  north  and  east 
along  the  Gulf  coast  of  Texas  and  Louisiana  to  the  delta  of  the  Mis- 
sissippi, thence  north  in  the  Mississippi  Valley  to  southeastern  Mis- 
souri; general  range  reaching  southeastern  Kansas,  probably  by  way 
of  the  Arkansas  River  valley  through  Oklahoma  (not  yet  known 
from  Oklahoma) ;  altitudinal  range  in  Austroriparian  Zone,  mainly 
below  500  feet,  but  extending  up  to  about  1,000  feet  in  Kansas. 

General  diameters. — Closely  resembling  0.  p.  palustris,  but  typical 
examples  paler;  skull  usually  narrower.  Color  paler,  less  rufescent, 
and  skull  decidedly  narrower,  less  massive,  than  in  O.  p.  natator  and 
O.  p.  color atus. 

Color. — About  as  in  0.  p.  palustris,  but  averaging  slightly  paler. 
An  ochraceous-tawny  suffusion  of  upperparts  and  underparts  is 
shown  in  rare  examples. 

Skull. — About  like  that  of  O.  p.  palustris,  but  averaging  somewhat 
smaller,  with  less  widely  spreading  zygomata.  Similar  to  that  of 
0.  p.  natator,  but  narrower;  braincase  decidedly  narrower;  zygomata 
less  widely  spreading;  frontal  region  narrower,  the  supraorbital 
borders  less  projecting. 

Measurements. — Type:  Total  length,  256 ;  tail  vertebrae,  139;  hind 
foot,  30.5.  Average  of  eight  adult  topotypes:  242  (226-279);  120 
(108-133);  29  (28.5-30.5).  Skull  (average  of  5  adults— type  and  4 
topotypes):  Greatest  length,  31  (30-32.1);  zygomatic  breadth,  16 
(15.2-16.8);  interorbital  breadth,  5.3  (5.2-5.4);  width  of  braincase, 
11.8  (11.1-12.3);  nasals,  12.5  (12-12.9);  anterior  palatine  foramina, 
6.1  (5.7-6.5);  palatal  bridge,  5.8  (5.5-6.1);  upper  molar  series,  4.4 
(4.3-4.7). 

Remarks. — Specimens  from  the  type  locality  are  paler  than  those 
from  other  localities  in  the  immediate  vicinity,  some  of  which  are 
practically  indistinguishable  from  many  typical  examples  of  0.  p. 
palustris.  Moreover,  in  cranial  characters,  especially  in  width  of 
braincase  and  outward  spread  of  zygomata  (characters  winch  dis- 
tinguish 0.  p.  texensis  from  0.  p.  natator  and  0.  p.  coloratus),  texensis 
very  closely  approaches  palustris.  The  skull  of  the  type  and  larger 
topotypes  of  texensis  seem  inseparable  from  some  of  the  smaller  skulls 
of  comparable  age  from  the  region  of  the  type  locality  of  palustris. 
The  cranial  difference  noted,  however,  affects  the  majority  of  indi- 


28 


NORTH   AMERICAN    FAUNA. 


[No.  43. 


viduals,  and  together  with  a  tendency  toward  pallid  coloration 
exhibited  by  animals  inhabiting  a  wide  area,  seems  to  entitle  texensis 
to  recognition  as  a  separate  form.  The  few  specimens  available  from 
Arkansas,  western  Tennessee,  and  extreme  southeastern  Missouri 
seem  referable  to  texensis,  but  approach  palustris  so  closely  that  they 
might  with  nearly  equal  propriety  be  assigned  to  that  subspecies. 
Specimens  examined. — Total  number,  110,  as  follows: 

Lake  City,  1;  Wilmot,  1. 


3;  Lake  Catherine, 


Arkansas:  Camden,  2;  Delight,  1; 

Kansas:  Neosho  Falls,  2.1 

Louisiana:  Burbridge,    IS;2  Gibson,   4;2  Houma,    1;  Iowa, 
6;3  Main  Pass,  2;  Mermenton,  4;4  New  Orleans,  2. 

Mississippi:  Fayette,  1.  / 

Missouri:  Kennett,  2;  Portageville,  1. 

Tennessee:  Arlington,  1. 

Texas:  Corpus  Christi,  7;5  Matagorda,  7;  Matagorda  Island,  2;  Matagorda  Penin- 
sula, 1;  Nueces  Bay,  5;  Padre  Island,  4;  Port  Lavaca,  4;  Rockport,  24  (type 
and  topotypes);6  Sabine,  1;  Victoria,  1;  Virginia  Point,  1 ;  Wharton  County ,  l.3 


Fig.  3. — Geographic  distribution  of  Oryzomys  couesi  and  related  species. 

ORYZOMYS  COUESI  (Alston). 

[Synonymy  under  subspecies.] 

Geographic  distribution. — River  valleys  and  marshy  areas  from 
southern  Texas  on  the  east  and  southern  Sinaloa  on  the  west,  south 
through  central  and  southern  Mexico,  Guatemala,  Honduras,  and 
Nicaragua  to  northern  Costa  Rica.  Altitudinal  range  from  sea  level, 
regardless  of  latitude,  to  over  7,000  feet  in  the  Valley  of  Mexico; 
zonal  range  mainly  Tropical  but  reaching  into  Sonoran  Zone  on  the 
tableland  of  Mexico  (fig.  3). 


1  One  in  Mus.  Comp.  Zool. 
*  Collection  Mus.  Comp.  Zool. 
3  Collection  Amer.  Mus.  Nat.  Hist. 
«  Collection  Field  Mus.  Nat.  Hist. 


'  One  in  collection  Amer.  Mus.  Nat.  Hist. 
6  Nineteen  in  collection  Amer.  Mus.  Nat.  Hist.;  2 
in  Field  Mus.  Nat.  Hist.;  1  in  Mus.  Comp.  Zool. 


1918.]  ORYZOMYS  PALUSTRIS  GROUP.  29 

General  characters. — Similar  in  general  to  0.  palustris,  but  pelage 
shorter;  colors  usually  brighter  and  richer,  ochraceous-buffy  or 
ochraceous-tawny  instead  of  grayish  brown ;  skull  differing  in  various 
details,  but  dentition  about  the  same. 

Color. — Upperparts  varying  from  light  buff  or  pinkish  buff  through 
ochraceous-buff  to  ochraceous-tawny,  more  or  less  suffused  with 
tawny  or  russet,  the  general  shade  paler  on  cheeks,  shoulders,  and 
along  sides,  becoming  deeper,  more  intense,  and  darkened  on  face, 
top  of  head,  and  back  by  admixture  of  black  hairs;  underparts 
varying  from  nearly  pure  white  through  light  buff  to  light  ochraceous- 
buff,  more  or  less  suffused  with  pale  salmon  color;  outer  sides  of  ears 
blackish,  inner  sides  moderately  clothed  with  short  hairs  varying 
from  grayish  to  ochraceous-buff  or  rusty  reddish;  feet  wnite;  tail 
dark  brownish  above,  whitish  or  light  brownish  below. 

Shutt. — (For  general  outlines  see  under  0.  palustris  group.)  In 
general  form  closely  resembting  that  of  0.  palustris,  but  sphenopala- 
tine vacuities,  large  in  that  species,  absent  or  much  reduced  in  size; 
antorbital  foramen  with  anterior  border  more  rounded  above,  more 
inclined  or  produced  forward  at  the  base,  not  excised  or  tending  to 
develop  the  somewhat  Sigmodon-Yike  point  often  present  in  palustris; 
interparietal  with  a  more  evident  posterior  angle. 

Remarks. — Ten  geographic  races  of  0.  couesi  appear  to  be  recog- 
nizable in  the  area  between  the  valley  of  the  Bio  Grande,  Texas,  and 
northern  Costa  Rica.  While  all  the  more  minute  steps  of  inter- 
gradation  are  not  always  shown  by  the  material  now  available, 
mainland  forms  throughout  this  wide  interval  agree  so  closely  in  all 
essential  details  that  they  seem  safely  assignable  to  a  single  species- 
Several  outlying  insular  forms  (and  0.  peninsulse  of  Lower  California) 
exhibit  more  distinctive  characters  and  are  accorded  specific  rank, 
but  they  are  clearly  related  to  the  widely  ranging  0.  couesi  section  of 
the  0.  palustris  group. 

ORYZOMYS  COUESI  COUESI   (Alston). 
Coues  Rice  Rat. 

(PI.  I,  figs.  3,  3a.) 

Hesperomys  couesi  Alston,  Proc.  Zool.  Soc.  London,  1876,  p.  756. 

Oryzomys  couesi  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  6,  XI,  p.  403,  May  1893  (type 

and  locality  fixed). 
Oryzomys  jalaps  Allen  and  Chapman,  Bull.  Amer.  Mus.  Nat.  Hist.,  IX,  p.  206,  June 

16, 1897.    Type  from  Jalapa,  Vera  Cruz,  Mexico  (altitude 4,400  ft.).  No.  fHiHr,  S 

ad.,  Amer.  Mus.  Nat.  Hist.;  collected  by  F.  M.  Chapman,  April  16,  1897. 
Oryzomys  jalapx  rufinus  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  285,  July  26, 

1901.     Type  from  Catemaco,  Vera  Cruz,  Mexico  (altitude  1,000  feet).     No.  65499, 

$  ad.,  U.  S.  Nat.  Mus.  (Biological  Survey  collection);   collected  by  Nelson  and 

Goldman,  April  27,  1894. 
Oryzomys  teapensis  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  286,  July  26,  1901. 

Type  from  Teapa,  Tabasco,  Mexico.     No.  99973,    <?   subad.,  U.  S.  Nat.  Mus. 

(Biological  Survey  collection);  collected  by  Nelson  and  Goldman,  April  4.  1900, 


30  NORTH   AMERICAN    FAUNA.  [No.  43. 

Oryzomys  goldmani  Merriam.  Proc.  Washington  Acad.  Sci.,  Ill,  p.  288,  July  26,  1901. 

Type  from  Coatzacoalcos,  Vera  Cruz,  Mexico  (near  sea  level).    No.  78110,  £  ad., 

U.  S.  Xat.  Mus.  (^Biological  Survey  collection^;  collected  by  Nelson  and  Goldman, 

April  11,  1S96. 
Oryzomys  jalap-  apatdius  Elliot.  Field  Columb.  Mus..  publ.  90.  zool.  ser.,  Ill,  p.  266, 

March  8.  1904.     Type  from  San  Carlos.  Vera  Cruz,  Mexico.     No.  13107,   $  ad., 

Field  Mus.  Nat.  Hist.;   collected  by  X.  G.  Buxton,  March  1,  1903. 
Oryzomys  richarc 's>" i  Mien.  Bull.  Arner.  Mus.  Nat.  Hist.,  XXVIII,  p.  99,  April  30, 

1910.     Type  from  Pena  Blanca.  Xicaragua.     No.  29800,  <?  ad.,  Amcr.  Mus.  Xat. 

Hist.;   collected  by  Wm.  B.  Richardson,  May  25,  1909. 

Type  locality. — Coban.  Guatemala. 

Type. — In  British  Museum:  collected  by  Osbert  Salvin. 

Geographic  distribution. — From  northern  Vera  Cruz  southeastward 
through  eastern  Puebla,  eastern  Oaxaca,  northern  and  extreme 
southern  Chiapas,  Tabasco,  Campeche,  Yucatan,  Quintana  Roo, 
Guatemala,  Honduras,  and  Nicaragua,  to  northwestern  Costa  Rica; 
altitudinal  range  from  sea  level  to  about  5,000  feet  mainly  in  Humid 
Lower  Tropical  Zone. 

General  characters. — Size  about  as  in  0.  c.  zygomaticus,  O.  c.  mexi- 
canus  and  0.  c.  richmondi;  color  slightly  darker  than  in  zygomuticus, 
decidedly  darker  than  in  mexicanus,  and  decidedly  paler  than  in 
ri-chmondi;  skull  about  like  those  of  mexicanus  and  richmondi; 
sphenopalatine  vacuities  absent  or  represented  by  very  narrow  slits 
as  usual  in  the  0.  couesi  section  of  the  O.  palustris  group.  Similar  hi 
general  to  O.  c.  peragrus,  but  somewhat  smaller,  with  upperparts, 
especially  cheeks,  shoulders,  and  sides,  more  ochraceous-buffy:  skull 
less  massive. 

Color. — Fresh  pelage:  Upperparts  varying  from  ochraceous-buffy 
to  ochraceous-tawny,  deepened  in  rare  examples  to  light  cinnamon- 
brown,  lightest  on  cheeks,  shoulders,  and  along  sides,  the  face,  top 
of  head,  and  back  much  darkened  by  black  hairs;  underparts  varying 
from  light  buff  to  light  ochraceous  buff  (rarely  dull  white);  outer 
sides  of  ears  blackish,  the  inner  sides  clothed  with  short  ochraceous- 
buffy  hairs;  feet  white;  tail  brownish  above,  dull  yellowish  below 
proximally,  becoming  light  brownish  toward  tip. 

STcuU. — About  as  in  0.  c.  zygomaticus,  0.  c.  mexicanus,  and  O.  c. 
richmondi;  differing  mainly  in  smaller  general  size,  decidedly  nar- 
rower braincase,  and  smaller  molars  than  those  of  0.  c.  crinitus,  O.  e. 
albivtnter,  and  other  Mexican  tableland  forms.  Similar  to  that  of 
0.  c.  peragrus,  but  braincase  narrower. 

Measurements. — Average  of  4  adults  from  Tumbala,  Chiapas: 
Total  length,  252  (242-265) ;  tail  vertebra?,  130  (127-135) ;  hind  foot, 
30.7  (30-31).  Average  of  10  adults  from  Yaruca,  Honduras:  267.5 
(255-2S0);  138(130-145);  29.1  (2S-32).  Average  of  seven  adults 
from  Orizaba,   Vera   Cruz:     263    (248-294);     14S    (139-174);     33.1 


1918.]  OEYZOMYS  PALUSTRIS  GROUP.  31 

(32-34.5).  Skull  (average  of  5  adults  from  Yaruca,  Honduras): 
Greatest  length,  30.5  (29.9-31.3);  zygomatic  breadth,  16  (15.5-17.2); 
interorbital  breadth,  4.8  (4.5-5.1);  width  of  braincase,  11.4  (11.2- 
11.6);  nasals,  11.9  (11.4-12.4);  anterior  palatine  foramina,  6 
(5.5-6.2);  palatal  bridge,  5.5  (5.3-6.1);  upper  molar  series,  4.7 
(4.5-4.8). 

Remarks. — In  the  absence  of  material  from  the  type  locality  aa 
fixed  by  Thomas,  specimens  from  Tumbala,  Chiapas;  and  Yaruca, 
Honduras,  which  agree  closely  with  his  description,  are  assumed  to 
represent  typical  couesi  and  have  been  used  as  a  basis  for  comparison. 
Individual  variation  in  size,  color,  and  cranial  details  exhibited  by 
every  large  series  of  0.  c.  couesi  is  very  striking,  but  the  form  main- 
tains with  remarkable  constancy  its  essential  characters  throughout 
its  wide  range.  Examples  from  northern  Vera  Cruz  and  eastern 
Puebla  present  the  same  general  variations  and  are  not  satisfactorily 
separable  from  those  from  Honduras  and  Nicaragua.  This  variation 
has  resulted  in  the  publication  of  several  names  based  on  characters 
which  prove  to  be  inconstant  in  the  large  number  of  specimens 
passed  in  review.  Specimens  from  various  localities  indicate  direct 
intergradation  with  0.  c.  zygomaticus,  0.  c.  mexicanus,  and  0.  c. 
peragrus. 

Specimens  examined. — Total  number,  199,  as  follows: 

Campeche:  La  Tuxpeiia,  1. 

Chiapas:  Chicharras,  4;  Tumbala,  6. 

Costa  Rica:  Bahia  de  Salinas,  l.1 

Guatemala:  Jacaltenango,  3. 

Honduras:  Yaruca,  35. 2 

Nicaragua:  Chontales,  22;1  Matagalpa,  4;1  Ocotal,  2;1  PenaBlanca,  8  (including 
type  of  "richardsoni") ;*  Quilali,  l;1  Rio  Coco,  14;1  Rio  Grande,  l;1  Rio  San 
JuandelNorte,  l;1  RioTuma,  2;1  San  Juan,  l^Tuma,  S^Uluce,  l;1  Vijagua^.1 

Oaxaca:  Comaltepec,  1;  Guichicovi,  3;  Reforma,  l3;  Santo  Domingo  (mountain 
near),  8;  Tuxtepec,  2. 

Puebla:  Huauchinango,  1;  Metlaltoyuca,  11. 

Quintana  Roo:  Santa  Lucia,  3.4 

Tabasco:  Teapa,  3  (including  type  of  "teapensis"). 

Vera  Cruz:  Achotal,  4;3  Buena  Vista,  2;  Catemaco,  2  (including  type  of 
"rufinus");  Jalapa,  4  (including  type  of  "jalapx");1  Pasa  Nueva,  l;1  Coat- 
zacoalcos,  3  (including  type  of  '  'goldmani' ' ) ;  Jico,  2  ;3  Mjrador,  1 ;  Motzorongo, 
1;  San  Carlos,  3  (including  type  of  "ajmtclius")-?  Orizaba,  16;5  Rivera  (75 
miles  south),  1;  Papantla,  1-  Teocelo,  If  Tlacotalpam,  7;  Ubero,  1. 

Yucatan:  Rio  Lagartos,  2.3 


i  Collection  Amer.  Mus.  Nat.  Hist. 

5  Twenty-six  specimens  in  Mus.  Comp.  Zool.;  6  in  Field  Mus.  Nat.  Hist. 

s  Collection  Field  Mus.  Nat.  Hist. 

*  Collection  Mus.  Comp.  Zool. 

5  Two  in  Field  Mus.  Nat.  Hist. 


32  NORTH   AMERICAN    FAUNA.  [No.  43. 

ORYZOMYS  COUESI  RICHMONDI  Merriam. 

Richmond  Rice  Rat. 

Oryzomys  richmondi  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  284,  July  26, 1901. 

Type  locality. — Escondido  River  (50  miles  above  Bluefields),  Nica- 
ragua. 

Type. — No.  |f f^g,  c?  adult,  United  States  National  Museum  (Bio- 
logical Survey  collection) ;  collected  by  Charles  W.  Richmond,  June 
21,  1892. 

Geographic  distribution. — Low  river  valleys  of  eastern  Nicaragua; 
Humid  Lower  Tropical  Zone. 

General  characters. — Size  and  proportions  about  as  in  0.  c.  couesi; 
color  decidedly  darker. 

Color. — Similar  to  that  of  0.  c.  couesi,  but  decidedly  darker,  more 
regularly  ochraceous-tawny,  the  back  and  upper  part  of  sides  more 
heavily  darkened  by  admixture  of  black  hairs;  underparts  light 
ochraceous-buff. 

Skull. — As  in  0.  c.  couesi. 

Measurements. — Type:  Total  length,  295;  tail  vertebrae,  150;  hind 
foot,  33.5.  Average  of  10  adults  (type  and  nine  topotypes):  275.8 
(255-295);  137  (124-151);  30.9  (29-33.5).  SJcull  (average  of  5 
adults — type  and  4  topotypes):  Greatest  length,  31.8  (31.2-33.3); 
zygomatic  breadth,  16.9  (16.4-17.9);  interorbital  breadth,  5.1  (5-5.3); 
width  of  braincase,  11.7  (11-11.9);  nasals,  12.3  (11.6-13);  anterior 
palatine  foramina,  6(5.8-6.2);  palatal  bridge,  5.8  (5.7-5.9);  upper 
molar  series,  4.5  (4.3-4.6). 

Remarks. — This  form,  the  darkest  of  the  0.  couesi  series,  is  known 
only  from  low  elevations  in  eastern  Nicaragua,  where  Richmond 
found  it  inhabiting  banana  plantations.  Although  much  darker  in 
general  tone  of  upperparts  than  most  examples  of  0.  c.  couesi  from 
adjacent  territory,  close  agreement  in  all  other  important  respects 
points  to  complete  intergradation  with  the  latter. 

Specimens  examined. — Total  number,  35,  as  follows: 

Nicaragua:  Escondido  River  (50  miles  above  Bluefields,  12 2  [type  and  topotypes] ; 
45  miles  above  Bluefields,  18;  40  miles  above  Bluefields,  3;  25  miles  above 
Bluefields,  1;  16  miles  above  Bluefields,  1). 

ORYZOMYS  COUESI  ZYGOMATICUS  Merriam. 

Guatemalan  Rice  Rat. 

Oryzomys  zygomaticus  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  285,  July  26, 
1901. 

Type  locality. — Nenton,  Guatemala  (altitude  3,000  feet). 

Type. — No.  76794,  s  adult  United  States  National  Museum  (Bio- 
logical Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A.  Gold- 
man, December  15,  1895. 


1918.]  OEYZOMYS  PALUSTEIS  GROUP.  33 

Geographic  distribution. — Known  only  from  the  Chiapas  River 
valley  in  southwestern  Guatemala  and  south-central  Chiapas;  Arid 
Lower  Tropical  Zone. 

General  characters. — Closely  resembling  0.  c.  couesi  and  0.  c.  mexi- 
canus,  but  upperparts  slightly  paler  than  in  the  former  and  slightly 
darker  than  in  the  latter. 

Color. — About  as  in  0.  c.  couesi  but  averaging  slightly  paler,  the 
general  tone  ochraceous-buff  as  in  the  palest  examples  of  0.  c.  couesi; 
underparts  nearly  pure  white  in  the  type,  varying  to  light  buff  in 
other  examples. 

SJcull. — As  in  0.  c.  couesi. 

Measurements. — Type:  Total  length,  290;  tail  vertebrae,  152;  hind 
foot,  33.  Skull  (type):  Greatest  length,  30.9;  zygomatic  breadth, 
26.9;  interorbital  breadth,  4.8;  width  of  braincase,  11.6;  nasals,  11.6; 
anterior  palatine  foramina,  6;  palatal  bridge,  5.9;  upper  molar  series, 
4.6. 

Remarks. — 0.  c.  zygomaticus  seems  to  be  a  slightly  differentiated  and 
rather  localized  form  intermediate  in  color  and  geographic  position 
between  0.  c.  couesi  and  0.  c.  mexicanus.  It  doubtless  intergrades 
directly  with  couesi,  the  type  of  which  came  from  a  higher  elevation 
about  100  miles  to  the  eastward  in  central  Guatemala.  Three  speci- 
mens from  Jacaltenango,  at  about  5,500  feet  altitude,  only  a  few 
miles  to  the  southeast,  are  appreciably  darker  in  color  and  seem  to 
represent  typical  couesi.  Near  the  Pacific  Coast  in  southwestern 
Chiapas,  zygomaticus  doubtless  passes  into  mexicanus,  which  differs 
mainly  in  having  slightly  paler  color. 

Specimens  examined. — Total  number,  5,  as  follows: 

Chiapas:  Ocuilapa,  1. 

Guatemala:  N  en  ton,  4  (type  and  topotypes). 

ORYZOMYS  COUESI  MEXICANUS  Allen. 

Mexican  Rice  Rat. 

Oryzomys  mexicanus  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  IX,  p.  52,  March  15,  1897. 
Oryzomys  bulleri  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  IX,  p.  53,  March  15,  1897.     Type 

from  Valle  de  Banderas,  Tepic,  Mexico,  No.  ffff,  <?  Bubad.,  Amer.  Mus.  Nat. 

Hist.;  collected  by  Audley  C.  Buller,  February  2,  1893. 
Oryzomys  rufus  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  287,  July  26,  1901. 

Type  from  Santiago,  Tepic,  Mexico  (altitude  200  feet).     No.  91404,  9  old,  U.  S. 

Nat.  Mus.   (Biological    Survey  collection);    collected   by    E.  W.  Nelson   and 

E.  A.  Goldman,  June  20,  1897. 

Type  locality. — Hacienda  San  Marcos,  Tonila,  Jalisco,  Mexico  (alti- 
tude 3,500  feet). 

Type.— No.  -ff§-£,  c?  adult,  American  Museum  of  Natural  History; 
collected  by  Audley  C.  Buller,  December  30,  1889. 
14521°— 18 3 


34  NORTH   AMERICAN    FAUNA.  [No.  43. 

Geographic  distribution. — Pacific  coastal  plains  and  basal  mountain 
slopes  from  southern  Sinaloa  to  southeastern  Oaxaca,  Mexico;  alti- 
tudinal  range  from  sea  level  to  about  1,000  feet  (rarely  to  3,500  feet), 
mainly  in  Arid  Lower  Tropical  Zone. 

General  characters. — Size  and  proportions  about  as  in  0.  c.  couesi; 
color  of  upperparts  rather  decidedly  paler,  more  ochraceous-buffy; 
underparts  usually  white,  but  varying  to  light  buff,  or  light  ochra- 
ceous-buff,  the  normal  shades  in  couesi.  Slightly  paler  than  0.  c. 
zygomaticus;  slightly  darker  than  0.  c.  aztecus,  with  smaller  molar 
teeth.  Similar  in  color  to  0.  c.  alhiv enter,  but  size  smaller,,  and  skull 
differing  in  detail. 

Color. — Fresh  pelage:  Upperparts  in  general  varying  shades  of 
ochraceous-buff,  more  or  less  suffused  with  tawny  in  old  adults,  be- 
coming warm  buff  on  cheeks,  shoulders,  and  along  lower  parts  of 
sides;  the  face,  top  of  head,  and  back  moderate^  darkened  by 
blackish  hairs;  underparts  usually  nearly  pure  white,  but  varying  to 
light  buff;  outer  sides  of  ears  dusky,  the  inner  sides  clothed  with 
grayish  or  rusty  reddish  hairs;  feet  white;  tail  brownish  above,  dull 
yellowish  below  on  proximal  portion,  becoming  light  brownish  toward 
tip.  Young  (in  first  pelage) :  Upperparts  near  tawny-olive,  the  gen- 
eral tone  darker  and  duller  than  in  adults. 

Skull. — About  as  in  O.  c.  couesi.  Differing  from  those  of  O.  c. 
aztecus,  O.  c.  regillus,  and  O.  c.  albiventer  most  noticeably  in  smaller 
size  of  molar  teeth,  the  toothrows  being  correspondingly  shortened. 

Measurements. — Average  of  10  adults  from  Escuinapa,  Sinaloa: 
Total  length,  251.4' (239-273);  tail  vertebrae,  137.4  (127-165);  hind 
foot,  28.9  (27-35).  SkuU  (average  of  same):  Greatest  length,  31.2 
(29.8-33);  zygomatic  breadth,  17  (16.1-17.8);  interorbital  breadth, 
4.8  (4.5-5.3) ;  width  of  braincase,  11.6  (11.3-12.2);  nasals,  12.2  (11.1- 
13.4);  anterior  palatine  foramina,  6.1  (5.8-6.7);  palatal  bridge,  5.5 
(4.8-6);  upper  molar  series,  4.4  (4.2-4.8). 

Remarks. — The  narrow  distribution  area  of  0.  c.  mexicanus  along 
the  west  coast  of  Mexico  somewhat  parallels  that  of  0.  c.  couesi  along 
the  east  coast;  and  as  in  that  form,  wide  range  of  individual  varia- 
tion in  size  and  color  has  resulted  in  the  publication  of  names  which 
appear  to  be  based  on  unstable  characters.  While  individuals  vary 
notably  in  size  and  contour,  as  shown  by  every  large  series,  skulls  of 
mexicanus  and  couesi  seem  indistinguishable,  their  general  characters 
being  maintained  with  remarkable  uniformity  throughout  the  com- 
bined ranges  of  the  two  forms.  Intergradation  of  mexicanus  with 
couesi  seems  to  be  indicated  by  specimens  from  the  Isthmus  of 
Tehuantepec,  and  with  0.  c.  aztecus  by  examples  from  the  valley  of 
the  Balsas  River  in  Guerrero. 


1918.]  OEYZOMYS   PALUSTKIS   GROUP.  35 

Specimens  examined. — Total  number,  106,  as  follows: 

Colima:  Armeria,  11;  Hacienda  Magdalena,  2. 

Guerrero:  Ometepec,  7. 

Jalisco:  Hacienda  San  Marcos,  1  (type).1 

Michoacan:  La  Huacana,  5. 

Oaxaca:  Huilotepec,  2;  Juchitan,  1;  Llano  Grande,  5;  Pluma,  3;  Puerto  Angel,  8; 

Reforma,  l;2  Santa  Efigenia,  1;  Tehuantepec,  1. 
Sinaloa:  Escuinapa,  47;1  Mazatlan,  l;3  Rosario,  2.4 
Tepic:  San  Bias,  4;  Santiago,  2  (including  type  of  "rufus");  Valle  de  Banderas, 

2  (including  type  of  "bulleri"). 

ORYZOMYS  COITESI  AZTECUS  Merriam. 

Aztec  Rice  Rat. 

Oryzomys  crinitus  aztecus  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  282,  July  26, 
1901. 

Type  locality. — Yautepec,  Morelos,  Mexico  (altitude  4,000  feet). 

Type. — No.  51173,  6"  adult,  United  States  National  Museum  (Bio- 
logical Survey  collection);  collected  by  E.  W.  Nelson  and  E.  A.  Gold- 
man, January  16,  1893. 

Geographic  distribution. — Interior  river  valleys  of  Morelos,  southern 
Puebla,  northern  Oaxaca,  and  northeastern  Guerrero,  Mexico;  alti- 
tudinal  range  from  about  3,000  to  at  least  4,000  feet  in  Arid  Lower 
Tropical  Zone. 

General  characters. — A  pale  form  with  white  underparts  and  rather 
heavy  dentition.  Closely  resembling  O.  c.  mexicanus,  but  paler  in 
color  and  with  heavier  dentition  than  usual  in  that  subspecies.  Dif- 
fering from  O.  c.  crinitus  in  paler  upperparts,  white  instead  of  buffy 
underparts,  and  in  cranial  details. 

Color. — As  in  the  paler  examples  of  O.  c.  mexicanus,  the  prevailing 
tone  beneath  the  dark  hairs  on  the  back  pale  ochraceous-buff ;  under- 
parts nearly  pure  white. 

Skull. — About  like  that  of  O.  c.  mexicanus,  but  molar  teeth  larger. 
Similar  to  that  of  0.  c.  crinitus,  but  braincase  narrower  and  molars 
slightly  smaller. 

Measurements. — Type:  Total  length,  290;  tail  vertebrae,  154;  hind 
foot,  35.  Two  adults  from  Puente  de  Ixtle,  Morelos:  318,  313;  160, 
170;  34,  33.  Skull  (type) :  Greatest  length,  32.4;  zygomatic  breadth, 
17;  interorbital  breadth,  5.2;  width  of  braincase,  12.2;  nasals,  12.9; 
anterior  palatine  foramina,  6.4;  palatal  bridge,  6.2;  upper  molar 
series,  5. 

Remarks. — O.  c.  aztecus  is  the  palest  form  of  the  0.  couesi  series 
but  the  light  ochraceous  tone  of  the  upperparts  is  very  closely  ap- 
proached by  the  paler  examples  of  O.  c.  mexicanus,  O.  c.  albiventer, 

1  Collection  Amer.  Mus.  Nat.  Hist.  » Collection  Mus.  Comp.  Zool. 

»  Collection  Field  Mus.  Nat.  Hist.  <  One  in  collection  Mus.  Comp.  Zool. 


36  NORTH   AMERICAN   FAUNA.  [No.  43. 

and  0.  c.  aquaticus.  Specimens  from  the  valley  of  the  Balsas  River 
in  Guerrero  and  from  northern  Puebla  approach  mexicanus  in  denti- 
tion, as  well  as  color,  and  might  with  nearly  equal  propriety  be  re- 
ferred to  that  subspecies.  The  characters  separating  aztecus  and 
0.  c.  crinitus  are  rather  slight,  as  indicated  in  the  original  descrip- 
tions, and  it  seems  best  to  regard  both  as  forms  of  0.  couesi. 
Specimens  examined. — Total  number,  20,  as  follows: 

Guerrero:  Balsas,  l;1  Tlalixtaquilla,  1. 

Morelos:  Cuernavaca,  1;  Puente  de  Ixtle,  6;2  Yautepec,  5  (type  and  topotypes). 

Oaxaca:  Cuicatlan,  2. 

Puebla:  Piaxtla,  4. 

/ 
ORYZOMYS   COUESI   CRINITUS  Merriam. 

Valley  of  Mexico  Rice  Rat. 

Oryzomys  crinitus  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  281,  July  26,  1901. 

Type  locality. — Tlalpam,  Federal  District,  Mexico  (altitude  7,500 
feet). 

Type. — No.  50182,  J1  subadult  (molars  unworn),  United  States 
National  Museum  (Biological  Survey  collection) ;  collected  by  E.  W. 
Nelson  and  E.  A.  Goldman,  November  30,  1892. 

Geographic  distribution. — Known  only  from  type  locality,  at  about 
7,500  feet  altitude  in  the  Valley  of  Mexico;  Upper  Sonoran  Zone. 

General  characters. — A  large,  moderately  dark  form  with  light 
buffy  underparts  and  heavy  dentition.  Similar  to  0.  c.  aztecus  and 
O.  c.  albiventer,  but  upperparts  darker  than  either  and  underparts 
buffy  instead  of  white ;   differing  also  in  cranial  details. 

Color. — Very  similar  to  that  of  0.  c.  mexicanus,  but  general  tone  of 
upperparts  slightly  darker;  underparts  light  buffy  as  in  some  exam- 
ples of  mexicanus. 

Skull. — Rather  large  with  broad  braincase  and  heavy  dentition. 
Very  similar  to  those  of  0.  c.  aztecus,  0.  c.  albiventer,  and  O.  c.  regillus, 
but  frontal  region  usually  broader  posteriorly;  dentition  about  the 
same;    interparietal  rather  large. 

Measurements. — Type:  Total  length,  307;  tail  vertebra?,  161 
hind  foot,  37.  Adult  topotype:  280;  148;  35.  Shull  (type) 
Greatest  length,  32.4;  zygomatic  breadth,  17;  interorbital  breadth,  5 
width  of  braincase,  12.3;  nasals,  12.4;  anterior  palatine  foramina 
6.7;  palatal  bridge,  5.7;   upper  molar  series,  4.9. 

Remarks. — In  the  vicinity  of  marshes  along  the  southern  border  of 
the  Valley  of  Mexico,  O.  c.  crinitus  occurs  at  about  7,500  feet,  the 
highest  altitude  attained  by  any  known  member  of  the  0.  couesi 
section  of  the  genus.  The  exact  relationship  of  this  form  to  O.fulgens 
remains  to  be  determined,  since  it  possesses  some  of  the  characters 

i  Collection  Field  Mus.  Nat.  Hist.  » Three  in  collection  Field  Mus.  Nat.  Hist. 


1918.]  0RYZ0MYS  PALUSTRIS   GROUP.  37 

ascribed  to  the  latter  species,  the  exact  habitat  of  which  is  unknown. 
It  is  closely  allied  to  the  other  forms  inhabiting  river  valleys  of  the 
plateau  region  of  Mexico — 0.  c.  aztecus,  0.  c.  albiventer,  and  0.  c. 
regillus. 

Specimens  examined. — Three,  from  type  locality. 

ORYZOMYS  COUESI   REGILLUS  Goldman. 

Michoacan  Rice  Rat. 

(PI.  I,  figs.  4, 4a.) 

Oryzomys  couesi  regillus  Goldman,  Proc.  Biol.  Soc.  Washington,  XXVIII,  p.  129, 
June  29,  1915. 

Type  locality. — Los  Reyes,  Michoacan,  Mexico. 

Type. — No.  125945,  d  adult,  United  States  National  Museum 
(Biological  Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A. 
Goldman,  February  17,  1903. 

Geographic  distribution. — Plateau  region  of  northwestern  Michoacan, 
Mexico;  altitudinal  range  from  about  3,000  to  4,000  feet,  mainly  in 
Arid  Lower  Tropical  Zone. 

General  characters. — A  large  form  closely  allied  to  O.  c.  albiventer; 
upperparts  darker,  more  rufescent  in  color.  Similar  in  general  to 
O.  c.  mexicanus,  but  larger  and  richer  colored;  cranial  details  also 
distinctive. 

Color. — Much  as  in  O.  c.  mexicanus  and  O.  c.  albiventer  but  upper- 
parts  darker  and  more  rufescent,  the  general  tone  rich  ochraceous- 
buff,  the  back  and  rump  strongly  suffused  with  tawny  and  lined  with 
black  hairs  as  usual  in  the  group;  underparts  varying  from  nearly 
pure  white  to  light  buff. 

STcull. — Like  that  of  O.  c.  albiventer;  similar  to  those  of  O.  c. 
mexicanus  and  0.  c.  couesi,  but  larger,  with  broader  braincase  and 
heavier  dentition. 

Measurements. — Type:  Total  length,  305;  tail  vertebrae,  169; 
hind  foot,  36.  Average  of  three  adult  topotypes:  308  (285-320); 
168  (155-180);  35  (34-36).  Skull  (average  of  4  adults— type  and  3 
topotypes);  Greatest  length,  33.1  (32.5-33.5);  zygomatic  breadth, 
18.1  (17.5-18.5);  interorbital  breadth,  4.9  (4.6-5.3);  width  of  brain- 
case,  12.5  (12.4-12.5);  nasals,  12.9  (12.5-13.3);  anterior  palatine 
foramina,  6.2  (6-6.5);  palatal  bridge,  6.5  (6.4-6.6);  upper  molar 
series,  5.1  (4.9-5.3). 

Remarks. — This  handsome  rice  rat  differs  only  in  color  from  its 
near  geographic  neighbor,  O.  c.  albiventer,  and  the  two  doubtless 
intergrade  in  northern  Michoacan. 

Specimens  examined. — Total  number,  13,  as  follows: 
Michoacan:  Los  Reyes,  11  (type  and  topotypes);  Querendaro,  1;  Zamora,  1. 


38  NORTH    AMERICAN    FAUNA.  [No.  48. 

ORYZOMYS   COUESI  ALBIVENTER  Merriam. 

White-Bellied  Rice  Rat. 

Oryzomys  albiventer  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  279,  July  26,  1901- 
Oryzomys  molestus  Elliot,  Field  Columb.  Mus.,  publ.  71,  zool.  ser.,  Ill,  p.  145,  Feb- 
ruary, 1903.    Type  from  Ocotlan,  Jalisco,  Mexico  (altitude  5,000  feet).    No. 
8667,  c?  old,  Field  Mus.  Nat.  Hist.;   collected  by  F.  E.  Lutz,  June,  1901. 

Type  locality. — Ameca,  Jalisco,  Mexico  (altitude  4,000  feet). 

Type. — No.  82236,  <?  adult,  United  States  National  Museum 
(Biological  Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A. 
Goldman,  February  6,  1897. 

Geographic  distribution. — River  valleys  of  the  plateau  region  in 
central  Jalisco,  Mexico;  altitudinal  range  from  about  4,000  to  5,000 
feet  mainly  in  Lower  Sonoran  Zone. 

General  characters. — Color  about  as  in  O.  c.  mexicanus,  but  size 
usually  larger  and  cranial  details  distinctive.  Similar  in  size  and 
proportions  to  O.  c.  regillus  and  O.  c.  crinitus,  but  paler  than  either, 
the  upperparts  lacking  the  rich  rufescent  tone  of  the  former,  and  the 
underparts  normally  white  instead  of  buffy  as  in  the  latter. 

Color. — About  as  in  O.  c.  mexicanus,  underparts  normally  white,  but 
varying  to  pale,  creamy  buff. 

SJcuU. — About  like  that  of  O.  c.  regillus;  similar  to  that  of  O.  c. 
crinitus,  but  frontal  region  usually  narrower  posteriorly.  Compared 
with  those  of  0.  c.  couesi  and  0.  c.  mexicanus  the  skull  is  decidedly 
broader,  with  heavier  dentition. 

Measurements. — Type:  Total  length,  288;  tail  vertebrae,  153; 
hind  foot,  37.5.  Average  of  10  adults  (type  and  9  topotypes):  294 
(276-314);  161  (151-173);  36.6  (34-40).  Skull  (average  of  10 
adults — type  and  9  topotypes):  Greatest  length,  33.1  (31.5-34.4); 
zygomatic  breadth,  17.9  (17.3-18);  interorbital  breadth,  4.8  (4.1-5.2); 
width  of  braincase,  12.3  (11.5-12.8);  nasals,  13  (12.3-14.2);  anterior 
palatine  foramina,  6.3  (5.9-6.6);  palatal  bridge,  6.3  (5.8-6.7); 
upper  molar  series,  5.3  (5-5.5). 

Remarks. — While  O.  c.  albiventer  is  usually  larger  in  general  size, 
with  broader  skull  and  decidedly  larger  molar  teeth  than  O.  c.  mexi- 
canus, occasional  examples  of  the  two  forms  are  difficult  to  distin- 
guish and  point  to  probable  intergradation  in  western  Jalisco. 
" Oryzomys  molestus"  of  Elliot  is  based  on  an  unusually  large  old 
adult  which  is  clearly  referable  to  albiventer,  as  are  four  examples 
in  the  Biological  Survey  collection  from  the  same  locality. 

Specimens  examined. — Total  number,  18,  as  follows: 
Jalisco:  Ameca,  12  (type and  topotypes);   La  Barca,  1;   Ocotlan,  5  (including 
type  of  ''molestus.") 1 

i  Collection  Field  Mus.  Nat.  Hist. 


1918.1  ORYZOMYS  PALUSTRIS   GROUP.  39 

ORYZOMYS  COUESI  PERAGRUS  Merriam. 

Rio  Verde  Rice  Rat. 

Oryzomys  mexicanus  peragrus  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  283, 
July  26,  1901. 

Type  locality. — Rio  Verde,  San  Luis  Potosi,  Mexico. 

Type. — No.  82119,  6*  subadult,  United  States  National  Museum 
(Biological  Survey  collection);  collected  by  E.  W.  Nelson  and  E.  A. 
Goldman,  January  8,  1897. 

Geographic  distribution. — Known  only  from  type  locality,  at  about 
3,000  feet  altitude  on  the  Rio  Verde  River,  in  southern  San  Luis 
Potosi;  Lower  Sonoran  Zone. 

General  characters. — Closely  allied  to  0.  c.  couesi  but  slightly  larger, 
with  upperparts  more  pinkish  instead  of  ochraceous-buffy  in  general 
tone;  skull  more  massive.  Similar  to  0.  c.  aquaticus  but  upperparts 
usually  darker,  the  ground  color  less  ochraceous-buffy  and  the  dorsal 
area  more  heavily  lined  with  black. 

Color. — Similar  to  that  of  0.  c.  couesi  but  upperparts  paler,  less 
ochraceous-buffy,  the  cheeks,  shoulders,  and  sides  near  pinkish  buff; 
lips  and  chin  whitish. 

SJcull. — About  like  that  of  0.  c.  aquaticus;  larger  and  more  massive 
than  that  of  0.  c.  couesi,  with  broader  braincase  and  broader  inter- 
orbital  space. 

Measurements. — Type:  Total  length,  294;  tail  vertebra?,  167;  hind 
foot,  35.  Average  of  2  adult  topotypes:  274  (265-283);  151  (143- 
160);  33.5  (33-34).  STcull  (average  of  3  adults— type  and  2  topo- 
types): Greatest  length,  31.5  (31-32.2);  zygomatic  breadth,  16.6 
(16.1-17.1);  interorbital  breadth,  5.5  (5.4-6.2);  width  of  braincase, 
12.1  (12-12.3);  nasals,  12.1  (11.8-12.5);  anterior  palatine  foramina, 
5.8  (5.7-5.8);  palatal  bridge,  5.9  (5.7-6.2);  upper  molar  series,  4.9 
(4.7-5). 

Remarks. — As  nearly  as  can  be  determined  by  the  three  known 
specimens,  all  young  adults,  0.  c.  peragrus  is  somewhat  intermediate 
in  characters,  as  well  as  geographic  position,  between  O.  c.  couesi  and 
0.  c.  aquaticus.  It  differs  in  color  from  both,  but  in  cranial  details 
approaches  the  latter  form. 

Specimens  examined.— Three,  from  type  locality. 

ORYZOMYS  COUESI  AQUATICUS  Allen. 
Rio  Grande  Rice  Rat. 

(PI.  I,  figs.  5,  5a.) 

Oryzomys  aquaticus  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  Ill,  p.  289,  June  30,  1891. 

Type  locality. — Brownsville,  Cameron  County,  Texas. 
Type. — No.  %m,  e  adult,  American  Museum  of  Natural  History; 
collected  by  F.  B.  Armstrong,  March  6,  1891. 


40  NORTH   AMERICAN    FAUNA.  [No.  43. 

Geographic  distribution. — Rio  Grande  Valley,  from  Camargo, 
Tamaulipas,  to  Gulf  coast  near  Brownsville,  Texas;  altitudinal  range 
from  sea  level  to  about  300  feet  in  lower  Sonoran  Zone. 

General  characters. — Size  large;  closely  resembling  0.  c.  peragrus 
but  upperparts  usually  'paler,  less  pinkish,  more  ochraceous-buffy; 
underparts  buffy,  as  in  peragrus.  Similar  in  general  to  0.  c.  couesi 
but  larger;  general  color  above  paler  ochraceous-buff;  skull  relatively 
broader  and  more  massive. 

Color. — Fresh  pelage:  Upperparts  near  ochraceous-buff,  palest  on 
cheeks,  shoulders,  and  along  sides;  the  face,  top  of  head,  and  back 
obscured  by  dusky  hairs;  underparts  varying  from  light  buff  to 
warm  buff,  becoming  in  some'  specimens  more  or  less  distinctly 
whitish  on  lips,  chin,  and  throat;  feet  whitish;  tail  brownish  above, 
pale  yellowish  below,  becoming  pale  brownish  toward  tip.  Young 
(in  first  pelage):  Upperparts  near  tawny-olive;  underparts  with  a 
pale  buffy  wash,  the  plumbeous  basal  color  of  the  fur  showing 
through. 

Skull. — About  like  that  of  0.  c.  peragrus;  decidedly  larger  and 
heavier  throughout  than  that  of  0.  c.  couesi,  with  broader  braincase 
and  more  widely  spreading  zygomata;  ascending  branches  of  p  re- 
maxillae  exhibiting  a  tendency  to  exceed  nasals  in  posterior  extent 
(nasals  usually  exceeding  premaxillae  in  forms  of  0.  couesi). 

Measurements. — Type:  Total  length,  286;  tail  vertebrae,  140;  hind 
foot,  31.2  (dry  skin).  Average  of  5  adult  topotypes:  297  (283-310); 
161  (138-180);  34.5  (32-38).  Shall  (average  of  same):  Greatest 
length,  33  (32-35.4);  zygomatic  breadth,  18  (16.7-18.8);  interorbital 
breadth,  5  (4.7-5.2);  width  of  braincase,  12  (11.6-12.7);  nasals,  12.7 
(11.6-13.8);  anterior  palatine  foramina,  6.3  (6.1-6.6);  palatal  bridge, 
6  (5.4-6.6);  upper  molar  series,  5.1  (4.8-5.3). 

Remarks. — The  Rio  Grande  Valley,  inhabited  by  O.  c.  aquaticus, 
marks  the  extreme  northern  limit  of  the  general  range  of  the  0.  couesi 
series.  No  rice  rats  are  yet  known  from  the  interval  of  coastal  plain 
in  Tamaulipas,  but  close  resemblance  in  all  essential  respects  points 
to  intergradation  through  O.  c.  peragrus  with  typical  couesi.  In 
general  size  and  contour  the  skull  of  aquaticus  is  not  widely  different 
from  those  of  0.  c.  albiventer,  0.  c.  regillus,  and  O.  c.  crinitus  of  the 
Mexican  plateau  region,  but  it  maintains  the  lighter  dentition  of 
typical  couesi. 

Specimens  examined. — Total  number,  41,  as  follows: 

Texas:  Brownsville,    37    (type    and    topotypes);1    Lomita  Ranch   (Hidalgo 

County),  1. 
Tamaulipas:  Camargo,  1;  Matamoros,  2. 

i  Fourteen  in  Amer.  Mus.  Nat.  Hist.;  3  in  Kansas  Univ.  Mus.;  2  in  Field  Mus.  Nat.  Hist.;  2  in  Mus. 
Comp.  ZooL;  2  in  Acad.  Nat.  Sci.  Philadelphia. 


1918.]  ORYZOMYS  PALUSTRIS  GROUP.  41 

ORYZOMYS  FULGENS  Thomas. 
Thomas  Rice  Rat. 
Oryzomys  fulgens  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  6,  XI,  p.  403,  May,  1893. 

Type  locality. — "Mexico."  Southern  Mexico,  exact  locality  un- 
known (probably  in  or  near  Valley  of  Mexico) . 

Type. — 70.6.20.3,  <?  adult,  British  Museum;  purchased  of  Geale, 
collected  by  A.  Boucard. 

Geographic  distribution. — Range  unknown. 

General  characters. — From  original  description:  "Size  large.  Fur 
very  thick,  coarse  and  woolly.  General  colour  above  bright  fulvous, 
brighter  than  in  any  other  Central- American  species;  anterior  half  of 
the  body,  including  the  head,  rather  paler  and  duller  than  the  pos- 
terior half.  Ears  decidedly  small,  broadly  rounded,  thinly  haired, 
their  hairs  practically  the  same  colour  as  those  of  the  head  in  general, 
so  that  they  are  not  distinguishable  by  colour  at  a  distance.  Lips, 
chin,  throat,  and  inguinal  region  whitish,  belly  with  a  strong  suffusion 
of  fawn,  which  reaches  a  maximum  on  the  breast  between  the  fore 
legs;  passage  of  upper  colour  into  lower  quite  gradual.  Outer  sides 
of  limbs  like  back,  inner  sides  whitish;  upper  surfaces  of  hands  and 
feet  thinly  clothed  with  pale  silvery-fawn  hairs.  Tail  long,  thinly 
haired,  the  scales  not  hidden  by  the  hairs;  above  blackish,  below  yel- 
lowish, darkening  towards  the  tip." 

Skull. — From  original  description:  "Skull  readily  distinguishable 
from  all  allied  species  by  its  great  breadth,  the  bold  expansion  of  the 
zygomata,  and  especially  by  the  evenly  incurved  outline  of  the 
supraorbital  edges;  in  all  other  species  these  edges  form  two  approxi- 
mately straight  lines  diverging  from  the  narrowest  interorbital 
point,  but  in  0.  fulgens  the  whole  inner  wall  of  the  orbit  forms  one 
even  curve,  the  breadth  at  the  posterior  end  of  the  olfactory  cham- 
ber being  scarcely  greater  than  at  the  anterior  end.  Nasals  broad 
and  flattened.  Frontal  premaxillary  processes  very  narrow  and 
barely  attaining  to  the  same  level  as  the  back  of  the  nasals.  Anterior 
palatine  foramina  large,  widely  open,  their  posterior  margin  just 
level  with  the  front  of  ml ." 

Measurements. — Dry  skin  of  type  (from  original  description): 
Head  and  body,  160;  tail,  151;  hind  foot,  37.5  (c.  u.).  Skull  (type): 
Upper  length/  31.8;  zygomatic  breadth,  17.8  (c);  nasals,  13.2; 
interorbital  breadth,  4.8;  diastema,  9.1;  anterior  palatine  foramina, 
7.2;  upper  molar  series,  5.2. 

Remarks. — The  type  of  0.  fulgens  I  have  been  unable  to  examine, 
and  none  of  the  more  recently  accumulated  material  from  Mexico 
can  at  present  be  assigned  with  certainty  to  that  species.  Thomas's 
full  description,  above  quoted,  and  comparisons  kindly  made  for  me 

1  To  back  of  parietal  suture  only. 


42  NORTH   AMERICAN    FAUNA.  [No.  43. 

by  W.  H.  Osgood,  however,  seem  to  indicate  that  this  is  a  member  of 
the  widely  dispersed  0.  couesi  section  of  the  genus.  Some  of  the 
characters  given  are  shared  in  common  by  various  forms  now  recog- 
nized. In  size  and  color  arrangement  it  closely  approaches  0.  c. 
crinitus,  of  the  Valley  of  Mexico,  and  may  be  identical  with  that 
form,  but  the  intensity  of  color  and  details  presented  by  the  incom- 
plete skull,  especially  the  form  of  the  interorbital  region,  appear  to 
be  distinctive  and  the  exact  position  of  fitlgens  remains  to  be 
determined. 

ORYZOMYS  GATUNENSIS  Goldman. 

Gatun  Rice  Rat. 
/ 
(PI.  I,  figs.  6, 6a.) 

Oryzomys  gatunensis  Goldman,  Smiths.  Misc.  Coll.,  LVI,  no.  36,  p.  7,  February  19, 
1912. 

Type  locality. — Gatun,  Canal  Zone,  Panama. 

Type. — No.  171034,  d  young  (about  two-thirds  grown),  United 
States  National  Museum  (Biological  Survey  collection) ;  collected  by 
E.  A.  Goldman  March  7,  1911. 

Geographic  distribution. — Known  only  from  type  locality,  near  sea 
level;  Humid  Lower  Tropical  Zone. 

General  characters. — A  dark-colored  form  externally  similar  to  0.  c. 
richmondi,  but  with  distinctive  cranial  characters. 

Color. — Fresh  pelage:  Upperparts  near  ochraceous-tawny,  palest 
on  cheeks,  shoulders,  and  along  sides;  the  face,  top  of  head,  and 
back  much  darker  by  admixture  of  black  hairs;  underparts  light 
ochraceous-buffy;  outer  sides  of  ears  blackish,  the  inner  sides  clothed 
with  ochraceous-buffy  hairs;  feet  thinly  covered  with  very  short 
whitish  or  grayish  hairs;  tail  light  brownish  above,  somewhat  paler 
below. 

Skull. — Similar  in  general  to  that  of  0.  c.  richmondi,  but  frontal 
region  broader,  the  lateral  margins  more  developed  as  supraorbital 
shelves;  interparietal  much  less  extended  antero-posteriorly ;  nasals 
more  prolonged  posteriorly  be3rond  premaxillse;  dentition  about  as 
in  richmondi. 

Measurements. — Type:  Total  length,  224;  tail  vertebrae,  115;  hind 
foot,  31.5.  Skull  (type):  Greatest  length,  27.7;  zygomatic  breadth, 
14.5;  interorbital  breadth,  5.3;  width  of  braincase,  1 1 .4 ;  nasals,  10.2; 
anterior  palatine  foramina,  5.6;  palatal  bridge,  5.2;  upper  molar 
series,  5. 

Remarks. — This  species  requires  comparison  with  only  0.  c.  rich- 
mondi, with  which  it  is  nearly  identical  in  color  and  general  external 
appearance.  The  skull,  however,  differs  in  apparently  important 
respects  from  those  of  all  the  forms  of  0.  couesi.  Especially  notice- 
able is  the  lateral  development  of  the  supraorbital  ridges,  a  character 


1918.]  ORYZOMYS   PALUSTRIS   GROUP.  43 

which  in  fully  adult  examples  would  doubtless  be  more  pronounced ; 
the  reduced  antero-posterior  extent  of  the  interparietal  seems  to  be 
another  distinguishing  feature.  On  the  other  hand  the  material 
representing  0.  gatunensis  is  scanty  and  the  range  of  individual  varia- 
tion being  undetermined,  intergradation  with  couesi  and  richmondi 
in  Costa  Rica  or  western  Panama  seems  not  improbable. 
Specimens  examined. — Two,  from  type  locality. 

ORYZOMYS  COZUMEL^]  Merriam. 
Cozumel  Rice  Rat. 

(PI.  I,  figs.  7,  7a.) 
Oryzomys  cozumelx  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  280,  July  26,  1901. 

Type  locality. — Cozumel  Island,  off  east  coast  of  Quintana  Roo, 
Mexico. 

Type. — No.  108462,  <?  adult,  United  States  National  Museum 
(Biological  Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A. 
Goldman,  April  8,  1901. 

Geographic  distribution.— Known  only  from  type  locality. 

General  characters. — Size  large;  tail  much  longer  than  head  and 
body;  color  dark.  A  large  species  allied  to  0.  c.  couesi  of  the  neighbor- 
ing mainland,  differing  in  larger  size,  darker  color,  and  proportionately 
longer  tail. 

Color.— Worn  pelage:  Upperparts  between  ochraceous-tawny  and 
cinnamon-brownish,  palest  on  cheeks,  shoulders,  and  sides,  darkened 
on  face,  top  of  head,  and  back  by  a  brownish  admixture;  underparts 
between  light  buff  and  light  ochraceous-buff ;  feet  whitish;  tail  dark 
brownish  above,  dull  yellowish  below  basally,  becoming  dusky  all 
around  toward  tip. 

Skull. — In  general  outline  closely  resembling  that  of  0.  c.  couesi 
but  larger  and  more  massive;  rostrum  and  anterior  roots  of  zygomata 
relatively  slightly  heavier;  dentition  about  as  in  couesi. 

Measurements. — Type:  Total  length,  332;  tail  vertebrae,  182;  hind 
foot,  35.  Average  of  6  adult  topotypes:  306  (285-327);  172  (163- 
177);  34.3  (33-35.5).  STcidl  (average  of  6  adults — type  and  5  topo- 
types): Greatest  length,  32  (30.5-34.4);  zygomatic  breadth,  16.9 
(15.9-17.9);  interorbital  breadth,  5  (4.7-5.6);  width  of  braincase, 
12.1  (11.5-12.6);  nasals,  12.5  (11.7-13.6);  anterior  palatine  foramina, 
6.3  (6.2-6.7);  palatal  bridge,  5.8  (5.7-6.1);  upper  molar  series,  5 
(4.8-5.3). 

Remarks. — This  insular  species  is  clearly  an  offshoot  of  O.  couesi, 
the  widely  ranging  mainland  form.  Its  divergence  from  typical 
couesi  is  mainly  in  the  direction  of  larger  general  size  and  the  develop- 
ment of  a  relatively  longer  tail. 

Specimens  examined. — Nine,  from  type  locality. 


44  NORTH   AMERICAN    FAUNA.  [No.  43. 

ORYZOMYS  ANTILLARUM  Thomas. 
Jamaican  Rice  Rat. 

Oryzomys  ant'dlarum  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  7,  I,  p.  177,  February,  1898. 

Type  locality. — Jamaica. 

Type— No.  45.10.25.48,  British  Museum;  collected  by  P.  H. 
Gosse. 

Geographic  distribution. — Known  only  from  Jamaica. 

General  characters. — Allied  to  0.  couesi;  size  and  proportions  about 
as  in  the  typical  subspecies;  color  apparently  similar;  skull  differing 
in  rather  slight  details. 

Color. — From  original  description  of  type:  "General  colour  dull 
rufous,  rather  (though  not  prominently)  richer  on  the  rump  and 
greyer  on  the  head;  black  lining  of  back  not  prominently  marked. 
Belly  dull  yellowish,  not  sharply  denned,  the  hairs  slaty  grey  basally. 
No  blackish  ring  round  eyes.  Ears  small,  their  visible  external  sur- 
face blackish  and  internal  yellowish,  but  in  neither  case  very  strongly 
contrasting  with  the  general  colour.  Hands  and  feet  dull  whitish 
above.  Tail  apparently  about  as  long  as  head  and  body,  very  thinly 
haired,  almost  naked,  pale  brownish  above,  rather  lighter  below." 
Two  specimens  in  the  United  States  National  Museum,  collected 
many  years  ago,  are  very  tawny  above,  but  appear  to  have  been  im- 
mersed in  alcohol,  and  the  naturally  ruddy  tone  thereby  intensified. 

Slcull. — Closely  resembling  that  of  0.  c.  couesi,  but  nasals  reaching 
farther  posteriorly  beyond  premaxillse  (nasals  and  premaxillse  more 
nearly  conterminous  in  couesi);  maxillary  arm  of  zygoma  heavier; 
anterior  palatine  foramina  shorter  than  usual  in  couesi;  dentition 
about  the  same. 

Measurements. — From  original  description  of  type  (measured  in 
skin):  "Head  and  body  (apparently  stretched)  130  millim. ;  tail  (im- 
perfect at  tip)  130;  hind  foot  without  claws  (moistened),  28."  *  *  * 
Skull  (type):  "Basal  length  (c.)  26,  basilar  length  (c.)  24;  greatest 
breadth  17;  nasals  12.6x4.1;  interorbital  breadth  5.2;  breadth  of 
braincase  on  squamosals  12.9;  interparietal  2.8  x  8.5;  palate  length 
from  henselion  14 ;  diastema  8.3 ;  palatal  foramina  5.7  x  2.1 ;  length  of 
upper  molar  series  4.6".  An  adult  from  Metcalfe  Parish  (dry  skin): 
Total  length,  252;  tail  vertebras,  122;  hind  foot  (c.  u.),  29.2.  STcuU 
(of  same):  Greatest  length,  30.5;  zygomatic  breadth,  16.6;  interor- 
bital breadth,  5.1 ;  width  of  braincase,  12.2 ;  nasals,  12.6 ;  anterior  pala- 
tine foramina,  5.7;  upper  molar  series,  4.5. 

Remarks. — The  relationship  of  the  Jamaican  rice  rat  to  0.  couesi 
was  pointed  out  by  Thomas  in  Ms  original  account  of  the  species.  In 
view  of  its  isolation,  the  general  agreement  in  all  the  more  important 
respects,  even  to  size,  a  character  usually  subject  to  modification  in 
insular  forms,  is  remarkable,  and  suggests  the  possibility  that  0.  aniil- 


1918.]  ORYZOMYS   PALUSTRIS   GROUP.  45 

larum,  may  have  been  transported  from  some  point  on  the  coast  of  the 
North  American  mainland  within  a  comparatively  recent  period. 
As  suggested  by  Thomas,  however,  the  fact  that  no  specimens  appear 
to  have  been  taken  since  1877,  while  rats  of  the  Old  World  have 
devastated  the  island,  to  be  persecuted  in  their  turn  by  the  mon- 
goose, introduced  in  1872,  renders  it  probable  that  the  rice  rat  has 
been  exterminated  there. 

Specimens  examined. — Total  number,  2,  as  follows: 
Jamaica:  Metcalfe  Parish,  1;  Spanishtown,  1. 

ORYZOMYS  PENINSULA  Thomas. 

Lower  California  Rice  Rat. 

(PI.  I,  figs.  8,  8a.) 

Oryzomys  peninsulse  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  6,  XX,  p.  548,  December, 
1897. 

Type  locality. — Santa  Anita,  Lower  California,  Mexico. 

Type. — Male  adult,  British  Museum. 

Geographic  distribution. — Known  only  from  very  limited  marshy 
areas  near  sea  level  in  extreme  southern  Lower  California;  Arid  Lower 
Tropical  Zone. 

General  characters. — General  size  and  proportions  about  as  in 
0.  c.  mexicanus;  color  similar,  but  anterior  part  of  body,  especially 
head  and  shoulders,  strongly  suffused  with  gray;  skull  rather  broad, 
with  squarely  spreading  zygomata  and  large  interparietal. 

Color. — Fresh  pelage:  Upperparts  between  warm  buff  and  pale 
ochraceous-buff  (more  or  less  distinctly  tawny  in  worn  pelage  of  old 
adults)  most  intense  on  rump,  becoming  light  buffy  grayish  on  head, 
fore  limbs,  shoulders,  and  sides;  underparts  overlaid  with  white, 
the  basal  color  of  the  fur  usually  plumbeous,  except  on  chin  and 
throat,  where  it  is  pure  white  to  roots;  outer  sides  of  ears  brownish, 
inner  sides  clothed  with  grayish  or  ochraceous-buff y  hairs;  feet  white; 
tail  light  brownish  above,  yellowish  below,  becoming  more  or  less 
distinctly  brownish  toward  tip.  Young  (in  first  pelage) :  Upper- 
parts  more  fuscous,  the  head  and  shoulders  lacking  the  grayish  suffu- 
sion so  noticeable  in  adults;  underparts  dull  buffy  or  soiled  white, 
becoming  pure  white  on  chin  and  throat. 

STcull. — Similar  to  that  of  0.  c.  mexicanus,  but  broader;  zygomata 
more  widely  and  squarely  spreading;  braincase  less  flattened,  higher, 
more  rounded  or  inflated;  interparietal  larger;  anterior  palatine 
foramina  much  broader,  more  widely  open;  lateral  wings  of  parietals 
extending  farther  below  temporal  ridges  at  expense  of  squamosals; 
ascending  branches  of  premaxillse  usually  reaching  posteriorly  beyond 
nasals  (nasals  usually  exceeding  premaxillse  in  posterior  extent  in 
mexicanus) . 


46  NORTH  AMERICAN    FAUNA.  [No.  43. 

Measurements. — Average  of  nine  adults  from  San  Jose  del  Cabo, 
Lower  California  (near  type  locality) :  Total  length,  283  (270-305) ; 
tail    vertebra?,    143    (138-156);   hind   foot,    32.2    (31.5-34).     STcull 
(average  of  eight  adults  from  San  Jose  del  Cabo,  Lower  California) 
Greatest  length,  32.5  (31.5-33.9);  zygomatic  breadth,  17.9  (17.1-19) 
interorbital  breadth,  5.1  (4.9-5.3) ;  width  of  braincase,  12  (11.4-12.2) 
nasals,    13    (12.6-13.4);   anterior  palatine  foramina,   6.6    (6.2-7.4) 
palatal  bridge,  6.2  (6-6.4);  upper  molar  series,  4.8  (4.7-5.1). 

Remarks. — 0.  peninsulas,  is  allied  to  0.  c.  mexicanus  of  the  adjacent 
mainland  coast  of  Mexico,  but  important  differential  cranial  charac- 
ters are  numerous.  The  range  of  the  species  seems  to  be  centered  in 
the  marshes  near  the  mouth  of  the  San  Jose  River  at  San  Jose  del 
Cabo,  extending  a  few  miles  inland  along  the  course  of  the  stream. 
The  neighboring  coast  is  extremely  arid  and  unsuitable  for  habitation 
by  an  Oryzomys. 

Specimens  examined. — Total  number,  16,  as  follows: 

Lower  California:  San  Jose  del  Cabo,  15;  Santa  Anita,  1  (topotype). 

ORYZOMYS  NELSONI  Merriam. 

Nelson  Rice  Rat. 

(PI.  II,  figs,  l,  la.) 

Oryzomys  nelsoni  Merriam,  Proc.  Biol.  Soc.  Washington,  XII,  p.  15,  January  27,  1898. 

Type  locality. — Maria  Madre  Island,  Tres  Marias  Islands,  off  coast 
of  Tepic,  western  Mexico  (altitude  800  feet). 

Type. — No.  89200,  o*  adult,  United  States  National  Museum  (Bio- 
logical Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A.  Gold- 
man, May  13,  1897. 

Geographic  distribution. — Known  only  from  type  locality,  Maria 
Madre  Island,  where  it  inhabits  moist  places  on  the  upper  slopes  at 
about  800  feet  altitude;  Arid  Lower  Tropical  Zone. 

General  characters. — Size  very  large;  tail  much  longer  than  head  and 
body;  color  of  upper  parts  rich  ochraceous-buff ;  skull  rather  long  and 
narrow,  but  massive.  Allied  to  0.  c.  mexicanus,  but  differing  widely 
in  details  of  structure. 

Color. — Fresh  pelage:  Upperparts  rich  ochraceous-buff,  most  intense 
on  rump,  paling  to  warm  buff  on  head,  shoulders,  and  lower  parts  of 
sides ;  somewhat  darkened  on  face,  top  of  head,  and  back  by  dusky 
hairs;  underparts  white;  outer  and  inner  sides  of  ears  thinly  clothed 
with  grayish  hairs ;  tail  light  brownish  above  and  all  round  near  tip, 
becoming  yellowish  below  on  basal  portion. 

STcull. — Massive,  upper  outline  rising  high  over  anterior  roots  of 
zygomata,  the  rostrum  very  heavy  and  strongly  decurved.  Some- 
what similar  in  general  to  that  of  0.  c.  mexicanus,  but  much  larger 
and  heavier;  rostrum  much  more  swollen  and  decurved;  zygomata 


1918.] 


OKYZOMYS   MELANOTIS  GROUP. 


47 


heavier,  but  relatively  less  widely  spreading,  the  sides  more  nearly 
parallel;  supraorbital  ridges  less  divergent  posteriorly;  interparietal 
larger;  anterior  palatine  foramina  relatively  shorter;  dentition  about 
as  in  mexicanus. 

Measurements. — Type:  Total  length,  324;  tail  vertebras,  190;  hind 
foot,  38.  Two  adult  topotypes:  344,  320;  191,  185;  39,  37.  Skull 
(average  of  3  adults — t}^pe  and  2  topotypes) :  Greatest  length,  35.8 
(34.5-37.8);  zygomatic  breadth,  18.7  (18.2-19);  interorbital  breadth, 
5.4  (5.2-5.6);  width  of  braincase,  12.9  (12.8-13.1);  nasals,  14.5 
(14.2-14.8);  anterior  palatine  foramina,  6.3  (6.1-6.7);  palatal  bridge, 
6.9  (6.6-7.3);  upper  molar  series,  5.3  (5.1-5.4). 

Remarks. — The  isolation  of  0.  nelsoni  on  an  island  70  miles  off  the 
mainland  coast  has  resulted  in  the  evolution  of  a  well-marked  species, 


i*i/ 


■*"/* 


^T: 


HI  DA*-6?. 


C?    /    HON  D. 


1.  0.  TO.  melanotis.  4.  0.  r.  megadon. 

2.  0.  m.  colimensis.  5.  O.  r.  yucatanensis. 

3.  O.  r.  rostratus. 

Fig.  4. — Geographic  distribution  of  the  Oryzomys  melanotis  group. 

but  agreement  in  essential  characters  with  forms  of  0.  couesi  places 
it  in  that  widely  distributed  section  of  the  genus.  It  differs  strikingly 
from  all  the  mainland  forms  in  the  remarkable  development  of  the 
rostrum. 

Specimens  examined. — Four,  from  type  locality. 

Oryzomys  melanotis  Group. 

Geographic  distributio7i — Mainly  lower  elevations  near  the  Pacific 
coast  of  Mexico  from  southern  Sinaloa  to  Colima,  and  along  the  gulf 
coast  from  southern  Tamaulipas  to  the  Yucatan  peninsula;  altitudinal 
range  from  sea  level  to  about  3,000  feet;  Arid  and  Humid  Lower 
Tropical  Zones  (fig.  4). 

General  characters. — Size  small,  medium,  or  large;  form  rather 
slender;  tail  about  same  length  as  head  and  body  or  somewhat  longer, 


48  NORTH   AMERICAN    FAUNA.  [No.  43. 

scantily  and  indistinctly  haired;  ears  large  and  conspicuous,  thinly 
clothed  externally  with  short,  fine  dusky  hairs  and  internally  with 
similarly  short,  ruf escent  hairs ;  general  pelage  short,  rather  harsh, 
and  lacking  the  woolly  quality  of  0.  palustris;  vibrissas  about  as  long 
as  head;  toes  of  hind  feet  more  or  less  distinctly  webbed  at  base,  the 
three  longest  bearing  tufts  of  silvery  bristles  which  project  beyond 
ends  of  claws;  claws  short,  recurved,  compressed,  and  sharp  pointed. 
Color  of  upperparts  varying  from  ochraceous-buff  to  rich  ochraceous- 
tawny,  lined  with  black;  underparts  white  or  buffy  whitish. 

Skull. — Size  small,  medium,  or  large,  with  rostrum  long  and  brain- 
case  rather  narrow ;  maxillary  /arm  of  zygoma  heavy ;  outer  wall  of 
antorbital  foramen  with  projecting  border  rounded  or  sloping  for- 
ward, the  antorbital  fossa  deep  and  conspicuous,  but  less  evenly 
circular  than  in  0.  palustris  as  viewed  from  above;  nasals  and  pre- 
maxillse  about  conterminous  posteriorly;  frontal  region  wide  or 
narrow, constricted  near  middle,  the  lateral  margins  ridged  or  smooth; 
temporal  ridges  moderately  developed  anteriorly  along  parieto- 
squamosal  sutures,  becoming  indistinct  posteriorly  in  crossing  lateral 
wings  of  parietals;  interparietal  large;  anterior  palatine  foramina 
short  and  broad,  much  shorter  than  palatal  bridge,  not  normally 
reaching  anterior  plane  of  first  molars;  palatal  pits  and  sphenopala- 
tine vacuities  large;  interpterygoid  fossa  broad;  audital  bullas  small, 
the  inner  sides  extensively  overlapped  by  mastoids;  basi-occipital 
broad;  angle  of  mandible  broad,  the  inferior  border  strongly  turned 
inward;  coronoid  process  short  and  low,  owing  largely  to  high,  thin 
commissural  border  extending  to  condyle ;  molars  small,  approaching 
the  0.  palustris  type  but  smaller,  second  upper  molar  more  evenly 
cleft  by  inner  reentrant  angle,  and  third  lower  molar  more  deeply 
incised  by  outer  reentrant  angle.  The  skulls  of  0.  melanotis  and  its 
allies  differ  from  those  of  the  0.  palustris  group  most  noticeably  in 
the  elongation  of  the  rostrum,  shortness  of  anterior  palatine  foramina 
in  relation  to  palatal  bridge,  small  size  of  audital  bulla?,  and  the  dental 
details  noted. 

Remarks. — Unlike  0.  palustris  and  its  allies,  which  live  on  the 
ground  mainly  in  open  marshy  places  or  resort  to  the  vicinity  of 
water,  members  of  this  group  favor  more  forested  areas,  where  they 
are  partially  scansorial  in  habits,  and  occur  on  well-drained  but  moist 
mountain  slopes  sometimes  at  points  distant  from  water. 

In  external  appearance,  forms  of  the  0.  melanotis  group  superficially 
resemble  those  of  0.  couesi,  and  the  two  often  occur  together  at  the 
same  localities.  In  contrast  with  0.  couesi,  however,  the  upperparts 
are  usually  more  rufescent,  the  ears  are  much  larger,  less  conspicu- 
ously hairy,  and  tufts  of  silvery  bristles  project  beyond  the  points 
of  the  three  longer  claws  on  the  hind  foot  (claws  reaching  beyond 
bristles  in  couesi). 


1918.]  ORYZOMYS  MELANOTTS  GROUP.  49 

Key  to  Species  and  Subspecies  of  the  O.  melanotis  Group. 

a1.  Size  larger;   hind  foot  30  or  more.     (Atlantic  coast  of  Mexico  [0.  rostratus].) 
b1.  Upper  molar  series  4  or  more. 

c1.  Color  darker.     (Tabasco;  Campeche.) O.  r.  megadon  (p.  54). 

c2.  Color  paler.     (Southern  Tamaulipas;    Vera  Cruz;    northern  Puebla;  north- 
eastern Oaxaca.) O.  r.  rostratus  (p.  53). 

b2.  Upper  molar  series  less  than  4.    (Yucatan.) O.  r.  yucatanensis  (p.  55). 

a2.  Size  smaller;   hind  foot  less  than  30.     (Pacific  coast  of  Mexico  [0.  melanotis}.) 

b1.  Color  darker.     (Sinaloa;  Tepic;  Jalisco.) O.  m.  melanotis  (p.  50). 

b2.  Color  paler.    (Colima.) O.  m.  colimensis  (p  51). 

ORYZOMYS  MELANOTIS  Thomas. 

[Synonymy  under  subspecies.] 

Geographic  distribution. — Pacific  coastal  plains  and  basal  mountain 
slopes  from  southern  Sinaloa  south  through  western  Tepic  and  Jalisco 
to  Colima;  altitudinal  range  from  sea  level  to  3,000  feet;  Arid  Lower 
Tropical  Zone. 

General  characters. — Size  small  or  medium;  general  color  above 
varying  from  rich  ochraceous-buffy  to  pale  ochraceous-tawny ;  simi- 
lar to  small  forms  of  0.  rostratus  in  general  external  appearance  but 
skull  differing  in  important  details. 

Color. — Fresh  pelage:  Upperparts  varying  from  ochraceous  -  buff 
to  pale  ochraceous-tawny  (sometimes  becoming  intense  tawny  or 
rusty  reddish  in  the  worn  pelage  of  old  adults),  rather  sparingly 
lined  with  black  hairs,  which  becoming  less  numerous  along  cheeks 
and  sides,  leave  them  a  purer,  brighter  color  than  the  back;  under- 
pays pale  buff y  white,  the  dark  basal  color  of  the  fur  showing  through ; 
outer  sides  of  ears  blackish,  the  inner  sides  thinly  clothed  with  rusty 
reddish  hairs;  a  more  or  less  conspicuous  patch  of  light-colored  fur 
under  base  of  ear;  feet  (epidermis)  dull  yellowish,  thinly  covered 
above  with  short  white  hairs;  tail  (epidermis)  brownish  above,  be- 
coming light  brownish  or  dull  yellowish  below  except  near  lip,  which 
is  usually  dark  all  around.  Young  (in  first  pelage):  Upperparts 
duller  and  darker,  the  general  tone  browner  than  in  adults. 

SlcuU. — Similar  in  general  to  that  of  O.  rostratus,  but  upper  outline 
less  elevated  over  anterior  part  of  frontals;  braincase  relatively 
higher,  the  parietal  region  more  expanded;  rostrum  more  slender, 
less  decurved;  frontals  broader,  with  narrow  and  delicate  but  rather 
well-developed  supraorbital  borders;  parietals  more  squarely  truncate 
anteriorly,  the  fronto-parietal  sutures  more  widely  divergent;  tem- 
poral ridges  tending  to  spread  more  widely  posteriorly,  rejoining 
squamosals  after  crossing  slightly  developed  lateral  wings  of  parietals; 
interparietal  larger,  with  a  less  evident  posterior  angle;  outer  wall 
of  antorbital  foramen  narrower,  less  extended  anteriorly;  anterior 
palatine  foramina  short  and  wide  as  in  rostratus;  dentition  about  the 
same. 

14521°— X8 4 


50  NORTH   AMERICAN    FAUNA.  [No.  43. 

Remarks. — Two  closely  allied  geographic  races  of  0.  melanotis 
occupy  a  part  of  the  arid  coast  region  of  western  Mexico,  where  they 
appear  to  be  completely  isolated  from  their  relatives  (subspecies  of 
0.  rostratus)  along  the  Gulf  coast  of  eastern  Mexico.  The  not  very 
distant  relationship  of  0.  melanotis  to  0.  rostratus  is  evidenced  in 
numerous  characters,  but  the  two  species  appear  to  be  distinct.  In 
general  external  appearance,  including  color,  they  are  much  alike; 
in  fresh  pelage  0.  melanotis  may  usually  be  distinguished  by  the 
whitish  subauricular  spots;  the  skulls  are  easily  separable  by  the 
characters  pointed  out. 

ORYZOMYS  MELAN0TIS  MELANOTIS  Thomas. 
Jalisco  Rice  Rat. 

(PI.  II,  figs.  2,  2a;  PI.  V,  fig.  5;  PI.  VI,  figs.  2,  2a.) 
Oryzomys  melanotis  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  6,  XI,  p.  404,  May,  1893. 

Type  locality. — Mineral  San  Sebastian,  Jalisco,  Mexico. 

Type. — 93.3.6.25,  d  old,  British  Museum;  collected  by  Dr.  Audley  C. 
Buller,  January  25,  1893. 

Geographic  distribution. — Coastal  plains  and  basal  mountain  slopes 
in  southern  Sinaloa,  Tepic,  and  Jalisco;  altitudinal  range  from  sea 
level  to  about  3,000  feet;  Arid  Lower  Tropical  Zone. 

General  characters. — Size  medium;  color  of  upperparts  near  pale 
ochraceous-taAvny;  closely  allied  to  0.  m.  colimensis  but  larger  and 
darker  colored;  externally  similar  to  pale  examples  of  0.  rostratus; 
skull  with  long,  slender  rostrum  and  high-arched  braincase. 

Color. — Fresh  pelage:  Upperparts  pale  ochraceous-tawny  (becoming 
intense  tawny  or  rusty  reddish  in  the  worn  pelage  of  old  adults), 
rather  sparingly  lined  with  black  hairs,  which  becoming  less  numerous 
along  cheeks  and  sides  leave  them  a  purer,  brighter  color  than  on 
the  back;  underparts  white  or  pale  buffy  white,  the  dark  basal  color 
of  the  fur  showing  through;  outer  sides  of  ears  blackish,  inner  sides 
thinly  clothed  with  rusty  reddish  hairs;  a  more  or  less  conspicuous 
patch  of  whitish  fur  under  base  of  ear;  feet  whitish;  tail  brownish 
above,  becoming  light  brownish  or  yellowish  below  except  near  tip, 
which  is  usually  dark  all  around.  Young  (in  first  pelage) :  Upper- 
parts  duller  and  darker,  the  general  tone  browner  than  in  adults. 

STcull. — Similar  to  that  of  O.  m.  colimensis,  but  larger,  with  com- 
paratively smaller  molar  teeth. 

Measurements. — Average  of  five  adult  topotypes:  Total  length, 
235  (228-244);  tail  vertebrae,  128  (124-134);  hind  foot,  28.2  (27.5-29). 
STcuU  (two  adult  topotypes):  Greatest  length,  28.5,  27.9;  zygomatic 
breadth,  14.4,  14.5;  interorbital  breadth,  4.8,  5.2;  width  of  brain- 
case,  10.8,  10.7;  nasals,  11.5,  10.3;  anterior  palatine  foramina,  4.3, 
5.2;  palatal  bridge,  5.4,  5.4;  upper  molar  series,  4,  4.3. 


1918.]  ORYZOMYS   MELANOTIC  GROUP.  51 

Remarks. — Specimens  from  localities  near  sea  level  in  Jalisco  and 
Tepic  are  larger  than  those  from  the  type  locality  at  3,000  feet  alti- 
tude on  the  slope  of  the  mountains,  and  may  represent  a  slightly 
different  form.  In  size  they  contrast  strongly  with  the  small  form 
0.  m.  colimensis  inhabiting  the  coast  of  Colima. 

Specimens  examined. — Total  number,  13,  as  follows: 
Jalisco:  Ixtapa,  2;  San  Sebastian,  6  (type  and  topotypes). 
Sinaloa:  Los  Limones,  l.1 
Tepic:  San  Bias,  2;  Santiago,  2. 

ORYZOMYS  MELANOTIS  COLIMENSIS,  subsp.  nov. 

Colima  Rice  Rat. 

(PI.  II,  figs.  3, 3a.) 

Type  locality. — Armeria,  Colima,  Mexico  (altitude  about  100  feet). 

Type.— No.  !ftff>  9  adult>  United  States  National  Museum  (Bio- 
logical Survey  collection);  collected  by  E.  W.  Nelson,  March  2,  1892. 
Original  number  1987. 

Geographic  distribution. — Forested  coastal  plains  and  basal  moun- 
tain slopes  in  the  State  of  Colima,  Mexico  ;  altitudinal  range  from 
sea  level  to  1,500  feet;  Arid  Lower  Tropical  Zone. 

General  characters. — A  small  form  closely  allied  to  O.  m.  melanotis; 
differing  mainly  in  smaller  size  and  paler  color;  general  tone  of  upper- 
parts  ochraceous-buff  instead  of  ochraceous-tawny  as  in  melanotis. 

Color. — Upperparts  in  general  near  ochraceous-buff,  the  top  of 
head  and  back  sparingly  mixed  with  black,  giving  a  lined  effect; 
cheeks,  shoulders,  and  sides  paler  than  back,  the  general  tone  light 
ochraceous-buff;  underparts  whitish  or  pale  buffy  whitish;  ears 
blackish  or  brownish,  thinly  clothed  on  outer  sides  with  short  dusky 
hairs,  and  on  inner  sides  with  buffy  or  rusty  reddish  hairs ;  a  patch  of 
whitish  fur  under  base  of  ear  as  in  0.  m.  melanotis;  feet  yellowish, 
thinly  covered  above  with  short  white  hairs;  tail  brownish  above, 
yellowish  below  to  near  tip,  which  is  dusky  all  around. 

Skull. — About  like  that  of  O.  m.  melanotis,  but  smaller;  molar  teeth 
actually  about  the  same  size  as  those  of  melanotis,  and  therefore  rela- 
tively larger. 

Measurements. — Type:  Total  length,  216;  tail  vertebrae,  116; 
hind  foot,  26.  Adult  topotype:  220;  118;  28.  Skull  (average  of 
two  adults,  type  and  topotype):  Greatest  length,  27  (26.3-27.7); 
zygomatic  breadth,  14  (13.9-14.2);  interorbital  breadth,  4.7  (4.5- 
4.9);  width  of  braincase,  10.2  (9.9-10.6);  nasals,  10.7  (10.3-11.1); 
anterior  palatine  foramina,  4.4  (4.3-4.5);  palatal  bridge,  5.4  (5.4- 
5.5);  upper  molar  series,  4  (3.9-4.1). 

i  Collection  Amor,  Mus.  Nat.  Hist. 


52  NORTH   AMERICAN    FAUNA.  [No.  43. 

Remarks. — This  small  form  is  known  only  from  the  State  of  Colima, 
but  probably  ranges  to  the  southward  along  the  coast  of  Michoacan. 
An  adult  example  from  Hacienda  Magdalena  is  dark  in  color  and  in 
this  respect  approaches  0.  m.  melanotis,  but  agrees  otherwise  with 
the  small  series  of  topotypes  from  near  the  Pacific  coast. 

Specimens  examined. — Total  number,  4,  as  follows: 
Colima:  Armeria,  3  (type  and  topotypes);  Hacienda  Magdalena,  1. 

ORYZOMYS  ROSTRATUS  Merriam. 

[Synonymy  under  subspecies.] 

Geographic  distribution. — Coastal  plains  and  basal  mountain  slopes 
from  extreme  southeastern  Tamaulipas  through  northern  Puebla, 
Vera  Cruz,  northeastern  Oaxaca,  Tabasco,  Campeche,  and  Yucatan 
to  northern  Quintana  Roo;  altitudinal  range  from  sea  level  to  about 
1,500  feet;  Arid  and  Humid  Lower  Tropical  Zones. 

General  characters. — A  rather  large,  rufescent  species,  not  very 
unlike  0.  melanotis  in  external  appearance,  but  cranial  characters 
distinctive.  (For  additional  characters,  excepting  specific  color,  see 
under  0.  melanotis  group.) 

Color. — Upperparts  varying  from  ochraceous-buff  to  rich  intense 
ochraceous-tawny,  purest  and  brightest  along  cheeks  and  sides;  the 
face,  top  of  head,  and  back  moderately  lined  with  black  hairs,  which 
alter  the  general  tone;  underparts  white  or  pale  buffy  white,  the 
plumbeous  basal  color  usually  showing  through;  outer  sides  of  ears 
blackish,  the  inner  sides  thinly  and  inconspicuously  clothed  with  pale 
buffy  or  rusty  reddish  hairs;  feet  (epidermis)  dull  yellowish,  thinly 
covered  with  short  white  hairs;  tail  (epidermis)  varying  from  nearly 
uniform  brownish  throughout  to  irregularly  yellowish  on  under  side. 
Young  (in  first  pelage) :  Upperparts  darker  and  less  rufescent  than  in 
adults. 

STcull. — Similar  in  general  to  that  of  0.  melanotis,  but  upper  outline 
more  elevated  over  anterior  part  of  frontals;  braincase  relatively 
lower,  the  parietal  region  less  expanded;  rostrum  heavier,  more 
decurved;  frontals  narrower  posteriorly;  parietals  less  squarely 
truncate  anteriorly,  the  fronto-parietal  sutures  less  widely  divergent; 
temporal  ridges  tending  to  spread  less  widely  posteriorly;  interpa- 
rietal smaller,  with  a  more  evident  posterior  angle;  outer  wall  of 
antorbital  foramen  broader,  more  extended  anteriorly;  anterior  pala- 
tine foramina  short  and  wide  as  in  melanotis;  dentition  about  the 
same. 

Remarks. — Three  geographic  races  of  0.  rostratus  are  recognizable, 
all  of  which  closely  intergrade  and  differ  rather  slightly  in  average 
size,  color,  or  cranial  details.  Viewed  as  a  whole,  the  forms  exhibit  a 
progressive  decrease  in  size  from  west  to  east,  typical  rostratus  being 
the  largest  and  0.  r.  yucatanensis  the  smallest  of  the  series. 


1918.]  ORYZOMYS   MELANOTIS  GROUP.  53 

ORYZOMYS  ROSTRATUS  ROSTRATUS  Merriam. 
Metlaltoyuca  Rice  Rat. 

(PI.  II,  figs.  4,  4a.) 
Oryzomys  rostratus  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  293,  July  26,  1901. 

Type  locality.— Metlaltoyuca,  Puebla,  Mexico  (altitude  800  feet). 

Type. — No.  93112,  <?  old,  United  States  National  Museum  (Bio- 
logical Survey  collection);  collected  by  E.  W.  Nelson  and  E.  A. 
Goldman,  February  5,  1898. 

Geographic  distribution. — Forested  coastal  plains  and  basal  moun- 
tain slopes  in  southeastern  Tamaulipas,  northern  Puebla,  Vera 
Cruz,  and  northeastern  Oaxaca;  altitudinal  range  from  sea  level  to 
about  1,500  feet;  Arid  and  Humid  Lower  Tropical  Zones. 

General  characters. — Similar  to  O.  r.  megadon  and  0.  r.  yucatanensis, 
but  averaging  larger  than  either;  color  rather  pale,  much  as  in  yuca- 
tanensis, slightly  paler,  less  rufescent  than  usual  in  megadon;  skull 
broad. 

Color. — Fresh  pelage:  Upperparts  rich  ochraceous-tawny,  purest 
and  brightest  along  cheeks  and  sides ;  the  face,  top  of  head,  and  back 
moderately  lined  with  black  hairs  which  darken  the  general  tone; 
underparts  nearly  pure  white  in  some  specimens,  pale  buffy  white 
in  others,  the  dark  basal  color,  however,  usually  showing  through; 
outer  sides  of  ears  blackish,  the  inner  sides  thinly  clothed  with  pale 
buffy  or  rusty  reddish  hairs;  feet  (epidermis)  dull  yellowish,  thinly 
covered  with  short  white  hairs;  tail  varying  from  nearly  uniform 
brownish  throughout  to  irregularly  yellowish  on  under  side. 

Skull. — Closely  resembling  those  of  O.  r.  megadon  and  O.  r.  yuca- 
tanensis in  general  form,  but  larger  and  relatively  broader  than 
either,  the  greater  breadth  most  conspicuous  in  the  braincase; 
dentition  about  as  in  megadon,  decidedly  heavier  than  in  yucatanensis. 
Similar  to  that  of  O.  talamancx,  but  narrower;  zygomata  less  squarely 
spreading,  the  sides  more  divergent  anteriorly;  frontal  region  nar- 
rower, the  supraorbital  ridges  weakly  developed;  parietals  with 
lateral  wings  less  developed  below  temporal  ridges;  interparietal 
smaller.  Contrasted  with  that  of  0.  melanotis,  the  skull  is  larger, 
the  upper  outline  more  arched  over  anterior  part  of  frontals;  brain- 
case  relatively  lower  and  natter;  frontals  relatively  narrower  pos- 
teriorly; interparietal  smaller. 

Measurements. — Type:  Total  length,  277;  tail  vertebrae,  141;  hind 
foot,  32.5.  Average  of  seven  adult  topo types:  255  (240-270);  136 
(125-145);  31.9  (30-33).  Slull  (average  of  7  adults— type  and  6 
topotypes):  Greatest  length,  31.4  (30.8-33.3);  zygomatic  breadth, 
16.1  (15.4-17.4);  interorbital  breadth,  5  (4.6-5.2);  width  of  braincase, 
11.5  (11.4-11.7);  nasals,  12.8  (11.5-13.8);  anterior  palatine  foramina, 
5.2  (4.4-6);  palatal  bridge,  6.6  (6.3-7.3);  upper  molar  series,  4.3 
(4.1-4.4). 


54  NORTH   AMERICAN    FAUNA.  [No.  43. 

Remarks. — While  0.  r.  rostratus  is  distinguished  by  somewhat 
larger  general  size,  and  the  greater  breadth  of  the  braincase  is  an 
especially  noticeable  cranial  feature,  the  wide  range  of  individual 
variation  shown  in  large  series  of  specimens  renders  the  smaller 
examples  difficult  to  separate  from  some  of  those  of  0.  r.  megadon. 
Specimens  from  Pasa  Nueva  and  Achotal,  Vera  Cruz,  are  rather 
small  and  grade  toward  megadon.  Those  from  Alta  Mira,  Tamaulipas, 
marking  the  extreme  northern  limit  of  the  known  range  of  rostratus, 
average  slightly  paler  than  typical  examples. 

Specimens  examined. — Total  number,  48,  as  follows: 

Oaxaca:  Santo  Domingo  (mountains  near),  5. 

Puebla:  Metlaltoyuca  14  (type  and  topotypes). 

Tamaulipas:  Alta  Mira,  5. 

Vera  Cruz:  Achotal,  8;1  Motzorongo,  1;  Pasa  Nueva,  6;2  San  Carlos,  9.1 

ORYZOMYS  ROSTRATUS  MEGADON  Merriam. 
Tabasco  Rice  Rat. 

(PI.  II,  figs.  5,  5a.) 

Oryzomys  rostratus  megadon  Merriam,  Proc  Washington  Acad.  Sci.,  III,  p.  294,  July 
26,  1901. 

Type  locality. — Teapa,  Tabasco,  Mexico. 

Type. — No.  99978,  <?  old,  United  States  National  Museum  (Bio- 
logical Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A. 
Goldman,  March  24,  1900. 

Geographic  distribution. — Heavily  forested  coastal  plains  and  lower 
mountain  slopes  in  Tabasco  and  Campeche;  altitudinal  range  from 
sea  level  to  at  least  500  feet;  Humid  Lower  Tropical  Zone. 

General  characters.- — Closely  resembling  O.  r.  rostratus  and  0.  r. 
yucatanensis,  but  color  slightly  richer,  more  tawny  than  either ;  skull 
smaller  than  that  of  rostratus  and  larger  than  that  of  yucatanensis. 

Color. — About  like  that  of  0.  r.  rostratus,  but  slightly  darker  and 
richer,  more  intense  ochraceous-tawny. 

Skull. — Similar  to  that  of  0.  r.  rostratus,  but  smaller  and  narrower; 
differing  from  that  of  0.  r.  yucatanensis  mainly  in  larger  size  and 
heavier  dentition. 

Measurements. — Type:  Total  length,  272;  tail  vertebrae,  140; 
hind  foot  32.5.  Average  of  2  adult  topotypes:  236.5  (236-237) ;  121 
(120-122);  30  (29-31).  Skull  (average  of  3  adults— type  and  2 
topotypes):  Greatest  length,  30.5  (29.1-32.3);  zygomatic  breadth, 
15.6  (15-16.5);  interorbital  breadth,  4.6  (4.4-4.9);  -  width  of  brain- 
case,  10.6  (10.5-10.7);  nasals,  11.5  (11.3-12);  anterior  palatine 
foramina,  4.9  (4.6-5.1);  palatal  bridge,  6.3  (6-6.8);  upper  molar 
series,  4.3  (4.1-4.6). 

1  Collection  Field  Mus.  Nat.  Hist.  s  Collection  Amer.  Mus.  Nat.  Hist. 


1918.]  ORYZOMYS   MELANOTTS   GROUP.  55 

Remarks. — 0.  r.  megadon  appears  to  be  a  rather  localized  form.    A 
series  of  specimens  from  Apazote,  Campeche,  agree  most  closely,  as  a 
whole,  with  typical  megadon,  but  some  are  rather  pale  in  color,  and  in 
the  reduced  size  of  molar  teeth  also  approach  0.  r.  yucatanensis. 
Specimens  examined. — Total  number,  18,  as  follows: 
Campeche:  Apazote,  11;  Champoton,  1. 
Tabasco:  Teapa  6  (type  and  topotypes). 

ORYZOMYS  ROSTRATUS  YUCATANENSIS  Merriam. 
Yucatan  Rice  Rat. 

(PI.  II,  figs.  6, 6a.) 

Oryzomys  yucatanensis  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  294,  July  26, 
1901. 

Type  locality. — Chichen  Itza,  Yucatan,  Mexico. 

Type. — No.  108139,  S  adult,  United  States  National  Museum 
(Biological  Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A. 
Goldman,  February  9,  1901. 

Geographic  distribution. — Forested  lowlands  of  Yucatan  and  north- 
ern Quintana  Roo ;  mainly  Arid  Lower  Tropical  Zone. 

General  characters. — Similar  to  0.  r.  rostratus  and  0.  r.  megadon  but 
somewhat  smaller  than  either;  skull  small,  with  very  small  molar 
teeth. 

Color. — Much  as  in  0.  r.  rostratus  but  decidedly  paler,  the  general 
tone  ochraceous-buff  instead  of  rich  ochraceous-tawny. 

Skull. — Most  closely  resembling  that  of  0.  r.  megadon,  but  smaller, 
with  short  anterior  palatine  foramina  and  decidedly  smaller  molar 
teeth. 

Measurements. — Type:  Total  length,  235;  tail  vertebrae,  119; 
hind  foot,  32.  Adult  from  Puerto  Morelos,  Quintana  Roo,  255;  136; 
31.  Skull  (type):  Greatest  length,  28.7;  zygomatic  breadth,  15; 
interorbital  breadth,  4.6;  width  of  braincase,  10.8;  nasals,  11.8;  ante- 
rior palatine  foramina,  4.2 ;  palatal  bridge,  5.9 ;  upper  molar  series,  3.7. 

Remarks. — This  rather  small  pale  form  of  0.  rostratus  apparently 
passes  into  0.  r.  megadon  in  southern  Campeche.  The  specimens 
from  Apazote  are  referred  to  megadon,  with  which  the  majority  agree 
most  closely,  but  several  are  indistinguishable  in  color  from  0.  f. 
yucatanensis,  and,  as  they  present  no  wide  departure  in  cranial 
details,  might  be  assigned  to  that  form  but  for  the  presence  of  the 
larger  or  darker  examples  in  the  same  series. 

Specimens  examined. — Total  number,  5,  as  follows: 

Quintana  Roo:  Puerto  Morelos,  1. 

Yucatan:  Chichen  Itza,  4  (type  and  topotypes). 


56 


NORTH   AMERICAN    FAUNA. 


[No.  43. 


Oryzomys  alfaroi  Group. 

Geographic  distribution. — Mountainous  districts  from  northern 
Puebla,  Mexico,  south  through  Central  America  and  northwestern 
South  America  to  Ecuador;  altitudinal  range,  from  about  1,000  to 
10,000  feet,  mainly  in  Humid  Upper  and  Lower  Tropical  Zones 
(fig.  5). 

General  characters. — Size  small;  form  slender;  tail  usually  longer 
than  head  and  body,  thinly  and  rather  indistinctly  haired;  ears 
large  and  conspicuous,  thinly  clothed  externally  and  internally  with 
short,  fine,  blackish  hairs;  general  pelage  usually  rather  short  and 
harsh,  becoming  longest  in  f  orms'ranging  at  high  elevations ;  vibrissa? 


1.  O.  a.  alfaroi.  7.  O.a.  saturatior. 

2.  O.a.dariensis.  8.  O.a.chapmani. 

3.  O.  a.  angusticeps.  9.  O.  a.  dilutior. 

4.  O.a.rhabdops.  10.  O.guerrerensis. 

5.  O.  a.  caudatus.  11.  O.  hylocetes. 

6.  O.  a.  palatinus. 
Fig.  5. — Geographic  distribution  of  the  Oryzomys 

alfaroi  group. 

about  as  long  as  head;  toes  of  hind  feet  inconspicuously  webbed  at 
base,  the  longest  three  bearing  tufts  of  silvery  whitish  bristles  which 
project  beyond  the  ends  of  the  claws  as  in  the  0.  talamancse  group; 
claws  short  and  recurved,  compressed  and  sharp  pointed.  General 
color  dark,  the  upperparts  usually  near  ochraceous-tawny,  heavily 
mixed  with  black;  underparts  dull  whitish  or  pale  buffy. 

Skull. — Small  and  delicate  in  structure,  with  rostrum  elongated, 
the  braincase  moderately  broad  and  rather  low;  maxillary  arm  of 
zygoma  weakly  developed;  outer  wall  of  antorbital  foramen  with 
anterior  border  rounded,  the  antorbital  notch  only  moderately  deep 
as  viewed  from  above;  nasals  and  premaxillaB  ending  posteriorly  in 
about  the  same  plane ;  frontal  region  broad,  the  lateral  margins  very 
slightly  elevated  to  form  delicate  supraorbital  ridges;  temporal 
ridges  slightly  developed  anteriorly  along  parieto-squamosal  sutures, 


1918.]  ORYZOMYS   ALFAKOI   GROUP.  57 

usually  becoming  obsolescent  posteriorly  in  crossing  lateral  wings  of 
parietals;  interparietal  large,  somewhat  irregularly  pointed-elliptical 
in  form;  anterior  palatine  foramina  short  and  broad,  shorter  than 
palatal  bridge  and  not  usually  reaching  anterior  plane  of  first  molars; 
palatal  pits  rather  large  and  rounded;  sphenopalatine  vacuities 
present  but  small;  interpterygoid  fossa  broad;  audita!  bullae  small ; 
angle  of  mandible  short  and  broad,  the  inferior  border  even  more 
strongly  turned  inward  than  in  the  0.  talamancse  group;  coronoid 
process  with  free  portion  short  owing  to  high  connecting  ridge  reach- 
ing nearly  to  summit  of  condyle ;  molars  small,  the  reentrant  angles 
shallow  and  the  crown  arrangement,  especially  the  enamel  islands  in 
the  second  upper  molars,  much  as  in  the  0.  palustris  group,  but  inner 
reentrant  angles  in  first  upper  molars  broader.  More  reliable  dis- 
tinguishing characters  are  presented  by  the  skull  and  teeth;  the 
molars  are  less  deeply  cleft  by  reentrant  angles  in  the  0.  alfaroi 
group,  and  the  enamel  island  present  at  the  postero-internal  base  of 
the  paracone  of  the  second  upper  molar  (moderately  worn)  is  absent 
in  the  talamancse  group. 

Remarks. — This  group,  comprised  of  a  series  of  small,  dark-colored, 
closely  allied,  and  somewhat  localized  forms  typified  by  0.  alfaroi, 
extends  in  an  irregular  chain  along  the  backbone  of  the  continent 
from  southern  Mexico  to  northwestern  South  America,  one  form  at 
least  ranging  as  far  south  as  Ecuador.  While  the  0.  palustris  and 
0.  talamancse  groups  range  mainly  at  low  elevations,  or  are  restricted 
to  the  vicinity  of  water  at  the  higher  levels,  alfaroi  and  its  allies  often 
inhabit  well-drained  but  moist  mountain  slopes,  where  they  com- 
monly ascend  to  high  elevations. 

0.  alfaroi  and  its  relatives  approach  the  members  of  the  0.  talamancse 
group  in  general  external  characters,  and  the  two  partially  overlap  in 
geographic  range.  In  appearance  individuals  may  be  much  alike, 
both  groups  having  large  ears,  appearing  nearly  naked,  and  slender 
limbs;  forms  of  the  alfaroi  group  are,  however,  usually  smaller  and 
decidedly  darker  in  general  color;  the  ears  are  smaller,  but  not  mark- 
edly different  in  color. 

Key  to  Species  and  Subspecies  of  the  O.  alfaroi  Group. 

a1.  Zygomata  broader  posteriorly  than  anteriorly. 

ft1.  Upperparts  darker  ochraceous-buff  or  ochraceous-tawny. 
c1.  Upperparts  less  intense  ochraceous-buff  or  ochraceous-tawny. 
d1.  Median  dorsal  area  not  distinctly  blackish. 
el.  Size  smaller;  total  length  less  than  240. 
f1.  Skull  broader;  zygomatic  breadth  13  or  more. 
g1.  Anterior  palatine  foramina  usually  4  or  more. 
ft1.  Rostrum  more  massive.     (Northern  Puebla.)  O.  a.  dilutior  (p.  68). 
ft2.  Rostrum  less  massive.     (Central  Vera  Cruz.) 

O.  a.  chapmani  (p.  67). 


58  NORTH   AMERICAN    FAUNA.  [No.  43. 

g2.  Anterior  palatine  foramina  usually  less  than  4.     (Mountains  of  Hon- 
duras; Nicaragua;  Costa  Rica;  western  Panama.) 

O.  a.  alfaroi  (p.  59). 
f2.  Skull  narrower;  zygomatic  breadth  less  than  13.     (Mountains  of  south- 
ern Tabasco  and  northwestern  Chiapas.). . .  O.  a.  palatinus  (p.  65). 
e2.  Size  larger;  total  length  240  or  more. 
f1.  Pelage  short;  tail  about  140.     (Mountains  of  northeastern  Oaxaca.) 

O.  a.  caudatus  (p.  64). 
f2.  Pelage  long;  tail  less  than  135.     (High  mountains  of  southwestern  Gua- 
temala and  of  central  and  southern  Chiapas.) 

O.  a.  angusticeps  (p.  62). 
d?.  Median  dorsal  area  distinctly  blackish.     (Mountains  of  northern  Chiapas.) 

/  O.  a.  saturatior  (p.  66). 

c2.  Upperparts  more  intense  ochraceous-tawny.     (Eastern  Panama.) 

O.  a.  dariensis  (p.  61). 
b2.  Upperparts  paler  ochraceous-buff  or  ochraceous-tawny.     (Pacific  slope  of  Sierra 

Madre  in  Guerrero  and  Oaxaca.) O.  guerrerensis  (p.  69). 

a2.  Zygomata  broader  anteriorly  than  posteriorly. 

bl.  Size  larger;  total  length  225  or  more.     (Southeastern  Guatemala.) 

O.  a.  rhabdops  (p.  63). 
b2.  Size  smaller;  total  length  less  than  225.     (Southern  Chiapas.) 

O.  hylocetes  (p.  70). 

ORYZOMYS  ALFAROI  Allen. 

[Synonymy  under  subspecies.] 

Geographic  distribution. — Heavily  forested  mountain  slopes  from 
northern  Puebla  south  through  southern  Mexico  and  Central  America, 
at  least  to  Colombia  and  Ecuador;  altitudinal  range  from  about  1,000 
to  10,000  feet,  mainly  in  Humid  Upper  and  Lower  Tropical  Zones. 

General  characters. — Size  small;  color  dark;  skull  light  and  rather 
delicate  in  structure.  Similar  in  general  to  0.  guerrerensis,  but  color 
much  darker,  and  skull  differing  in  detail.  (For  additional  general 
characters  see  under  O.  alfaroi  group.) 

Color. — Fresh  pelage:  Upperparts  varying  from  dark  ochraceous- 
buff  to  ochraceous-tawny,  tawny,  russet,  or  cinnamon-brown,  usually 
heavily  mixed  with  black,  this  color  often  predominant  over  dorsum, 
the  lighter  element  purest  and  becoming  more  or  less  ochraceous- 
buffy  on  cheeks,  shoulders,  and  sides;  underparts  dull  white  or  buffy, 
thinly  overlying  the  dark  plumbeous  basal  color;  nose  blackish;  inner 
and  outer  sides  of  ears  thinly  clothed  with  very  short  black  hairs, 
light  ochraceous-buffy  subauricular  spots  present  in  some  forms, 
absent  in  others;  feet  dull  whitish,  the  elongated  silvery  tufts  on  toes 
of  hind  feet  projecting  beyond  points  of  longer  claws;  feet  (epidermis) 
yellowish,  thinly  clothed  above  with  very  short  glossy  white  hairs;  tail 
nearly  naked,  brownish  or  blackish  above,  yellowish  below  basally, 
becoming  dark  all  around  toward  tip.  Young  (in  first  pelage) :  Upper- 
parts  blackish,  finely  and  inconspicuously  mixed  with  ochraceous- 
tawny;  underparts  darker  than  in  adults,  the  dark  basal  color  thinly 


1918.]  OKYZOMYS  ALFAROI   GROUP.  59 

overlaid  with  white;  feet  varying  from  dull  whitish  to  brownish;  tail 
blackish. 

Skull. — Similar  in  general  to  that  of  0.  guerrerensis,  but  more 
elongated;  braincase  higher,  less  flattened.  In  the  anteriorly  spread- 
ing zygomata,  one  form  of  0.  alfaroi  (0.  a.  rhabdops)  approaches 
0.  hylocetes,  but  the  latter  is  much  smaller,  with  very  small  teeth,  and 
present  material  seems  to  indicate  specific  distinctness.  (For  addi- 
tional characters  see  under  0.  alfaroi  group.) 

Remarks. — All  the  North  American  members  of  the  0.  alfaroi  group, 
excepting  0.  hylocetes  and  0.  guerrerensis,  and  the  South  American 
forms  O.palmirse  and  0.  gracilis,  appear  to  be  assignable  subspecifically 
to  0.  alfaroi.1  While  complete  intergradation  may  not  be  shown  by 
the  material  examined,  the  more  essential  characters  prevail  with 
such  uniformity  throughout  the  series  as  to  leave  little  room  for  doubt 
of  its  existence.  The  accession  of  new  material  may  not  improbably 
show  that  liylocetes  and  guerrerensis  are  also  geographic  races  of 
alfaroi.  Typical  alfaroi  presents '  closer  resemblance  to  the  geo- 
graphically distant  race  0.  a.  chapmani,  of  Vera  Cruz,  than  to  the 
annectent  forms  inhabiting  the  high  mountains  of  Chiapas  and 
Guatemala.  This  resemblance  between  the  more  widely  removed 
subspecies  may  be  due  to  the  fact  that  the  intermediate  races  occupy 
more  diversified  areas,  most  of  them  having  ascended  to  high  eleva- 
tions where  peculiar  environmental  conditions  would  conduce  to 
differentiation. 

OKYZOMYS  ALFAROI  ALFAROI  (Allen). 

Alfaro  Rice  Rat. 

(PI.  ill,  figs.  1,  la.) 

Hesperomys  (Oryzomys)  alfaroi  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  Ill,  p.  214,  April 

17,  1891. 
Oryzomys  alfaroi  Allen,  Abstr.  Proc.  Linn.  Soc.   New  York,  1893-94,  p.  36,  July.  20, 

1894. 
Oryzomys  alfaroi  incertus  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXIV,  p.  655,  October 

13,  1908.     Type  from  Rio  Grande,  Nicaragua,  No.  28584,  S  ad.,  Amer.  Mus.  Nat. 

Hist.;  collected  by  W.  B.  Richardson,  March  28,  1908. 

Type  locality. — San  Carlos,  Costa  Rica. 

Type. — No.  ff-jHr,  9  subadult,  American  Museum  of  Natural  His- 
tory; collected  by  Anastasio  Alfaro,  December,  1888. 

Geographic  distribution. — Heavily  forested  mountainous  portions 
of  Honduras,  Nicaragua,  Costa  Rica,  and  western  Panama;  alti- 
tudinal  range  from  about  1,000  to  4,000  feet;  mainly  in  Humid 
Lower  Tropical  Zone. 

1  The  South  American  forms  will  therefore  stand  as  follows: 

Oryzomys  alfaroi  palmirse  Allen Miraflores,  Colombia. 

Oryzomys  alfaroi  gracilis  Thomas Concordia,  Medellin,  Colombia. 


60  NORTH   AMERICAN    FAUNA.  [No.  43. 

General  characters. — A  rather  small  form  with  short  pelage,  resem- 
bling 0.  a.  dariensis,  but  coloration  duller,  lessrufescent;  skull  differ- 
ing in  slight  details.  Very  similar  in  general  to  0.  c.  chapmani,  but 
color  usually  duller;  skull  with  shorter  anterior  palatine  foramina. 
Smaller  than  0.  a.  angusticeps  and  0.  a.  rhabdops,  with  shorter  pelage 
than  either. 

Color. — Upperparts  varying  from  ochraceous-buff  to  dull  ochra- 
ceous-tawny,  heavily  mixed  with  black.  (Other  colors  as  given 
under  0.  alfaroi.) 

STcull. — Size  medium  for  the  group,  rather  narrow  and  elongated 
with  narrowly  spreading  zygomata  and  short,  wide  anterior  palatine 
foramina.  In  general  form  very  similar  to  that  of  0.  a.  dariensis, 
but  braincase  and  frontal  region  usually  broader;  apparently  differing 
from  those  of  0.  a.  angusticeps  and  0.  a.  chapmani  most  noticeably 
in  shorter  anterior  palatine  foramina.  Compared  with  that  of  0.  a. 
rhabdops  the  skull  is  decidedly  narrower,  with  smaller,  less  inflated 
braincase,  and  zygomata  much  less  divergent  anteriorly. 

Measurements. — Average  of  6  adults  from  Tuis,  Costa  Kica:  Total 
length,  210  (188-221);  tail  vertebras,  109  (102-114);  hind  foot,  26.3 
(26-27).  STcull  (average  of  same) :  Greatest  length,  26.7  (25.6-28.3) ; 
zygomatic  breadth,  13.5  (13.2-14.1);  interorbital  breadth,  5.2  (5.1- 
5.5);  width  of  braincase,  10.5  (10.1-10.8);  nasals,  10.9  (10.3-11.5); 
anterior  palatine  foramina,  3.9  (3.5-4.9) ;  palatal  bridge,  5.4  (5.3-5.7) ; 
upper  molar  series,  3.7  (3.6-3.8). 

Remarks. — While  a  considerable  gap  separates  the  known  geo- 
graphic ranges  of  0.  a.  alfaroi  and  0.  a.  dariensis,  these  forms  exhibit 
such  close  approach  in  size,  color,  and  cranial  details  that  intergrada- 
tion  may  be  safely  assumed.  Specimens  from  Yaruca,  Honduras,  are 
referable  to  alfaroi,  but  in  the  larger  size  shown  by  some  examples  are 
not  very  unlike  O.  a.  angusticeps  and  0.  a.  rhabdops.  Three  speci- 
mens from  Managua,  Nicaragua,  recorded  by  Thomas  *  as  0.  gracilis, 
were  probably  assignable  to  alfaroi. 

Allen's  "0.  a.  incertus"  was  based  on  rather  richly  colored  speci- 
mens which  are  well  within  the  range  of  individual  variation  exhibited 
by  typical  alfaroi,  as  the  accession  of  additional  material  indicated 
to  him.2 

The  general  range  of  alfaroi  overlaps  that  of  0.  talamancse,  and  as  it 
may  closely  resemble  that  species  in  general  appearance  some  con- 
fusion of  the  two  has  resulted.  While  many  examples  may  be 
inseparable  in  color,  alfaroi  is  a  smaller  animal  than  talamancse.  The 
skull  of  alfaroi  is  distinguished  by  its  smaller  size  and  more  delicate 
structure,  the  maxillary  arm  of  the  zygoma  is  more  slender  and  the 
teeth  are  decidedly  smaller;  the  second  upper  molar  has  a  large, 

i  Thomas,  Oldfield,  Ann.  Mag.  Nat.  Hist.,  ser.  6,  XVI,  p.  57,  July,  1895. 
»  Allen,  J.  A.,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXVIII,  p.  99,  Apr.  30,  1910. 


1918.] 


ORYZOMYS  ALFAROI   GROUP.  61 


elongated  enamel  island  at  the  posterointernal  base  of  the  paracone, 
and  the  internal  reentrant  angle  extends  less  than  halfway  across 
the  crown  of  the  moderately  worn  tooth,  while  in  talamancx  the 
enamel  island  mentioned  is  absent  or  represented  only  by  a  small 
island  near  the  apex  of  the  reentrant  angle,  which  in  this  species 
reaches  halfway  across  the  molar  crown. 

Specimens  examined. — Total  number,  86,  as  follows : 

Costa   Rica:   San  Carlos,  3  (type  and  topotypes);  l  Tuis,  15;  1   exact   locality 

unknown,  7.1 
Honduras:  Yaruca,  19. 2 
Nicaragua:  Chontales,  6;1  Jalapa,  2;1  Jicaro,  l;1  Jinotega,   l;3  Rio  Coco,  5;1 

Rio  Grande,  3  (including  type  of  "incertus");1  San  Juan,  5;1  Tuma,  2;1 

Uluce,  2;1  exact  locality  unknown,  l.1 
Panama:  Boquete,  14.* 

ORYZOMYS  ALFAROI  DARIENSIS  Goldman. 
Darien  Rice  Rat. 

(PI.  Ill,  figs.  2,  2a;  PI.  V,  figs.  G;  PI.  VI,  figs.  3,  3a.) 

Oryzomys  alfaroi  dariensis  Goldman,  Proc.  Biol.  Soc.  Washington,  XXVIII,  p.  128, 
June  29,  1915. 

Type  locality. — Cana,  eastern  Panama  (altitude  2,000  feet). 

Type. — No.  178660,  $  adult,  United  States  National  Museum 
(Biological  Survey  collection);  collected  by  E.  A.  Goldman,  March  4, 
1912. 

Geographic  distribution. — Heavily  forested  mountain  slopes  in 
eastern  Panama  at  2,000  feet  altitude,  and  probably  adjacent  por- 
tions of  Colombia;  Humid  Lower  Tropical  Zone. 

General  characters. — A  small  form  closely  allied  to  0.  a.  alfaroi; 
color  of  upperparts  richer,  more  rufescent;  skull  usually  narrower. 
Similar  to  the  South  American  forms  O.  a.  gracilis  and  to  0.  a.  pal- 
mirse,  but  color  more  rufescent  and  skull  differing  in  detail. 

Color. — Fresh  pelage:  Upperparts  between  ochraceous-tawny  and 
tawny,  finely  mixed  with  black;  becoming  paler  and  ochraceous- 
buffy  on  cheeks,  shoulders,  and  lower  part  of  sides;  underparts,  feet, 
and  tail  as  given  under  O.  alfaroi. 

STcull. — About  like  that  of  O.  a.  alfaroi,  but  braincase  and  frontal 
region  usually  narrower.  Closely  resembling  that  of  O.  a.  palmirx, 
of  South  America,  but  shorter,  with  more  widely  spreading  zygomata 
and  smaller  teeth.  Compared  with  Ecuadorean  specimens  assumed 
to  represent  O.  a.  gracilis,  the  skull  is  more  massive,  with  more 
widely  spreading  zygomata. 

1  Collection  Amer.  Mus.  Nat.  Hist. 

!  Fourteen  specimens  in  Mus.  Comp.  Zool.;  2  in  Field  Mus.  Nat.  Hist. 

»  Collection  Field  Mus.  Nat.  Hist. 

*  Eleven  specimens  in  Mus.  Comp.  Zool.;  2  in  Field  Mus.  Nat.  Hist.;  1  in  Amer.  Mus.  Nat.  Hist. 


62  NORTH  AMERICAN   FAUNA.  [No.  43. 

Measurements. — Type:  Total  length,  203;  tail  vertebrae,  107;  hind 
foot,  25.5.  Average  of  5  adult  topotypes:  220  (212-226);  113  (107- 
117) ;  24.6  (23-26).  SJcull  (average  of  6  adults,  type  and  5  topotypes) 
Greatest  length,  27.3  (26.5-27.8);  zygomatic  breadth,  14.3(14-14.7) 
interorbital  breadth,  4.8  (4.6-5);  width  of  braincase,  10.3  (10-10.5) 
nasals,  11.1  (10.6-11.4);  anterior  palatine  foramina,  4.3  (4-4.8) 
palatal  bridge,  5.6  (5-5.9);  upper  molar  series,  3.7  (3.6-3.9). 

Remarks. — This  small,  slender  rice  rat  differs  from  typical  0.  a. 
alfaroi,  of  Costa  Rica,  mainly  in  richer,  more  tawny  coloration.  It  is 
closely  allied  to  the  Colombian  form  described  as  0.  palmirse  and  the 
latter  is  clearly  assignable  to  subspecific  rank,  if  it  does  not  prove 
to  be  identical  with  0.  a.  gracilis,  the  type  of  which  came  from 
farther  north  in  the  Cauca  Valley.  Comparison  with  specimens  from 
northern  Ecuador,  assigned  to  gracilis  by  Mr.  Oldfield  Thomas,  and 
reference  to  the  original  description  of  that  species  indicate  that  the 
two  are  very  nearly  related.  The  description  of  the  color  of  gracilis, 
however,  seems  to  apply  to  the  Ecuadorean  specimens,  or  to  palmirse, 
rather  than  to  the  Darien  animal.  Moreover,  the  skull  0.  a.  dariensis 
is  distinguished  from  that  of  gracilis,  as  here  understood,  by  the 
greater  lateral  expansion  of  the  zygomata. 

0.  talamancse  also  occurs  at  the  type  locality  of  dariensis  and  the 
two  are  superficially  much  alike.  The  smaller  size,  especially  the 
smaller  hind  foot,  usually  distinguishes  dariensis  externally,  while  the 
skull  is  smaller,  more  delicate  in  structure,  the  maxillary  arm  of  the 
zygoma  more  slender  and  the  molar  teeth  much  smaller;  the  second 
upper  molar  has  a  large  elongated  enamel  island  at  the  postero-inter- 
nal  base  of  the  paracone,  and  the  internal  reentrant  angle  extends  less 
than  half  way  across  the  crown  of  the  moderately  worn  tooth,  much 
as  in  0.  palustris.  In  talamancse  the  enamel  island  is  absent  and  the 
reentrant  angle  reaches  halfway  across  the  molar  crown. 

Specimens  examined. — Eleven,  from  type  locality. 

ORYZOMYS  ALFAROI  ANGUSTICEPS  Merriam. 

Volcan  Santa  Maria  Rice  Rat. 

Oryzomys  angusticeps  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  292,  July  26, 
1901. 

Type  locality. — Volcan  Santa  Maria,  Guatemala  (altitude,  9,000  feet). 

Type. — No.  76816,  c?  adult,  United  States  National  Museum  (Bio- 
logical Survey  collection);  collected  by  E.  W.  Nelson  and  E.  A.  Gold- 
man, January  22,  1896. 

Geographic  distribution. — Heavily  forested  slopes  of  high  moun- 
tains in  southwestern  Guatemala  and  central  and  southern  Chiapas; 
altitudinal  range  from  about  8,000  to  9,500  feet. 

General  characters. — Similar  to  0.  a.  alfaroi  but  larger,  with  longer 
pelage;  anterior  palatine  foramina  longer.  Closely  approaching 
0.  a.  rhabdops  in  size,  but  color  darker  and  skull  differing  in  detail. 


1918.]  ORYZOMYS  ALFAROI   GROUP.  63 

Color. — Lighter  element  in  upperparts  pale  cinnamon  brownish, 
heavily  mixed  with  black,  the  general  tone  appreciably  darkened  by 
the  dark  plumbeous  basal  color  of  the  long  pelage;  underparts  and 
sides  of  muzzle  light  ochraceous-buffy;  ears,  feet,  and  tail  as  given 
under  0.  alfaroi. 

Skull. — Similar  to  that  of  0.  a.  alfaroi,  but  larger,  zygomata  more 
squarely  spreading  anteriorly,  the  sides  more  nearly  parallel;  frontal 
region  usually  narrower;  anterior  palatine  foramina  longer.  Ap- 
proaching that  of  0.  a.  rhabdops  in  size,  but  zygomata  less  divergent 
anteriorly. 

Measurements. — Type:  Total  length,  245;  tail  vertebrse,  134;  hind 
foot,  29.  STcull  (type):  Greatest  length,  28.6;  zygomatic  breadth, 
14;  interorbital  breadth,  4.6;  width  of  braincase,  10.3;  nasals,  10.6; 
anterior  palatine  foramina,  5;  palatal  bridge,  5.4;  upper  molar 
series,  3.6. 

Remarks. — Aside  from  its  geographic  neighbor,  0.  a.  rJiabdops, 
no  other  North  American  form  of  the  0.  alfaroi  group  attains  so  high 
an  altitude;  like  rhabdops  it  has  developed  a  longer  pelage  than 
forms  ranging  at  lower  elevations.  The  skull  of  the  type  of  0.  a. 
angusticeps  seems  to  be  abnormally  narrow;  the  topotypes  are  all 
young  and  their  skulls  of  little  value  for  comparative  purposes.  An 
apparent  tendency  toward  greater  breadth  shown  in  the  skulls  of 
specimens  from  San  Cristobal  and  Pinabete,  Chiapas,  is  believed  to 
be  within  the  probable  range  of  individual  variation;  on  the  other 
hand,  they  do  not  differ  appreciably  from  some  skulls  of  rliabdops,  and 
suggest  intergradation.  In  size  and  color  the  Chiapas  examples  are 
very  similar  to  the  topotypes. 

Specimens  examined. — Total  number,  16,  as  follows: 

Chiapas:  Pinabete,  2;  San  Cristobal,  10. 

Guatemala:  Volcan  Santa  Maria  4  (type  and  topotypes). 

ORYZOMYS  ALFAROI  RHABDOPS  Merriam. 

Calel  Rice  Rat. 
(PI.  Ill,  figs.  3,  3a.) 
Oryzomys  rhabdops  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  291,  July  26,  1901. 

Type  locality. — Calel,  Guatemala  (altitude,  10,000  feet). 

Type. — No.  76813,  S  adult,  United  States  National  Museum 
(Biological  Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A. 
Goldman,  January  15,  1896. 

Geographic  distribution. — -Known  only  from  type  locality,  at  about 
10,000  feet  altitude  on  the  upper  slope  of  the  high  mountains  in 
southwestern  Guatemala. 

General  characters. — Size  large;  pelage  very  long  for  a  member  of  the 
O.  alfaroi  group,  the  longer  hairs  on  dorsum  reaching  about  11  mm.; 


64  NORTH   AMERICAN    FAUNA.  [No.  43. 

color  rather  pale,  upperparts  near  ochraceous-tawny;  skull  broad 
with  anteriorly  divergent  zygomata.  Similar  in  general  to  0.  a. 
angusticeps,  but  color  paler,  and  skull  differing  in  detail. 

Color. — Fresh  pelage:  Upperparts  near  ochraceous-tawny,  richest 
and  purest  on  cheeks,  shoulders,  and  sides,  the  back  darkened  by 
black  hairs,  but  less  copiously  than  in  most  forms  of  0.  alfaroi;  sides 
of  muzzle  and  underparts  in  general  light  ochraceous-buffy;  nose 
deep  black,  except  extreme  tip  which,  like  lips,  is  whitish;  ears  black, 
with  more  or  less  distinct  light  ochraceous-buffy  subauricular  spots; 
feet  and  tail  as  given  under  0.  alfaroi.  Young  (in  first  pelage) : 
Upperparts  blackish,  inconspicuously  tinged  with  ochraceous-tawny; 
underparts  plumbeous,  lightly  overlaid  with  light  ochraceous-buff. 

Skull. — Skull  broad,  with  large  inflated  braincase  and  anteriorly 
diverging  zygomata.  Similar  to  that  of  0.  a.  angusticeps,  but 
zygomata  wider  anteriorly  than  posteriorly  (sides  more  nearly  parallel 
in  angusticeps).  In  general  form,  especially  the  anteriorly  spreading 
zygomata,  similar  to  that  of  0.  hylocetes,  but  much  larger,  with  shorter 
anterior  palatine  foramina. 

Measurements. — Type:  Total  length,  255;  tail  vertebrae,  141;  hind 
foot,  29.5.  Adult  topotype:  255;  138;  28.5.  Skull  (average  of 
two  adults — type  and  topotype) :  Greatest  length,  28.6  (28.4-28.8) ; 
zygomatic  breadth,  14.8  (14.6-15.1);  interorbital  breadth,  4.8 
(4.S-4.9);  width  of  braincase,  10.9(10.8-11.1);  nasals,  11  (10.5-11.5); 
anterior  palatine  foramina,  4.3  (4.2-4.5) ;  palatal  bridge,  5.7  (5.5-5.9) ; 
upper  molar  series,  3.8  (3.7-3.9). 

Remarks. — No  other  North  American  form  of  the  genus  is  known  to 
range  10,000  feet  above  sea  level,  but  a  near  geographic  neighbor, 
0.  a.  angusticeps,  which  has  also  pushed  up  above  the  Tropical  Zones 
on  mountains  of  similar  elevation,  approaches  and  may  equal  this 
altitude.  The  two  forms  are  evidently  closely  allied.  Distinguishing 
cranial  characters  are  developed  mainly  in  adults,  the  skulls  of  most 
of  the  younger  examples  being  apparently  inseparable.  In  the 
remarkable  anterior  expansion  of  the  zygomata  the  skull  of  O.  a. 
rhabdops  resembles  that  of  0.  hylocetes,  but  the  latter  appears  to  be  a 
very  distinct  form. 

Specimens  examined. — Fourteen,  from  type  locality. 

ORYZOMYS  ALFAROI  CAUDATUS  Merriam. 
Comaltepec  Rice  Rat. 

(PI.  Ill,  figs.  4,  4a.) 
Oryzomys  chapmani  caudatus  Merriam,  Proc.  Washington  Acad.  Sci  ,  III,  p.  289,  July 
26,  1901. 

Type  locality. — Comaltepec,  Oaxaca,  Mexico  (altitude  3,500  feet). 

Type. — No.  68641,  <?  adult,  United  States  National  Museum  (Bio- 
logical Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A.  Gold- 
man, July  31,  1894. 


1918.]  OEYZOMYS  ALFAROI   GROUP.  65 

Geographic  distribution. — Mountains  of  northeastern  Oaxaca;  alti- 
tudinal  range  from  3,500  to  6,500  feet;   Humid  Upper  Tropical  Zone. 

General  characters. — Size  largest  of  the  0.  alfaroi  group;  color 
dark;  pelage  short.  Similar  in  general  to  0.  a.  chapmani,  but 
decidedly  larger,  with  the  lighter  colored  admixture  in  upperparts  of 
ochraceous-buffy  instead  of  ochraceous-tawny;  skull  larger  and  more 
massive  than  usual  in  the  alfaroi  group. 

Color. — Fresh  pelage:  Upperparts  ochraceous-buffy,  heavily  mixed 
with  black,  the  latter  color  predominating,  especially  along  lower  part 
of  back;  underparts  and  sides  of  muzzle  light  ochraceous-buffy; 
lips  and  chin  whitish;  nose  black;  ears,  feet,  and  tail  as  given  under 
0.  alfaroi. 

Skull. — Size  very  large  and  structure  heavy  for  0.  alfaroi.  Very 
similar  in  general  form  to  that  of  0.  a.  chapmani,  but  much  larger 
with  more  swollen  rostrum. 

Measurements. — Type:  Total  length,  257;  tail  vertebrae,  141;  hind 
foot,  30.  Skull  (type) :  Greatest  length,  29;  zygomatic  breadth,  15; 
interorbital  breadth,  5;  width  of  braincase,  11.3;  nasals,  11.4; 
anterior  palatine  foramina,  4.8;  palatal  bridge,  5.2;  upper  molar 
series,  3.9. 

Remarks. — 0.  a.  caudatus  was  based  on  a  single  specimen  which 
seems  to  indicate  a  subspecies  still  larger  than  0.  a.  rhabdops  and 
0.  a.  angusticeps,  but  the  extent  of  individual  variation  remains  to  be 
determined.  Although  not  very  widely  differing  from  angusticeps  it 
seems  to  be  most  closely  allied  to  0.  a.  chapmani,  the  greater  size 
being  the  chief  distinguishing  character.  An  immature  example 
from  6,500  feet  altitude  at  Totontepec,  Oaxaca,  is  referable  to  the 
same  form. 

Specimens  examined. — Total  number,  2,  as  follows: 
Oaxaca:  Comaltepec,  1  (type);  Totontepec,  1. 

ORYZOMYS  ALFAROI  PALATINUS  Merriam. 
Teapa  Rice  Rat. 
(PI.  Ill,  figs.  5,  5a.) 
Oryzomys  palatinus  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  290,  July  26,  1901. 

Type  locality. — Teapa,  Tabasco,  Mexico  (altitude,  3,000  feet). 
Type.—  No.  99977,   9  adult,  United  States  National  Museum  (Bio- 
logical Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A.  Gold- 
man, April  1,  1900. 

Geographic  distribution. — Forested  mountain  slopes  in  southern 
Tabasco  and  northwestern  Chiapas;  known  altitudinal  range  from 
about  3,000  to  3,500  feet;  Humid  Upper  Tropical  Zone. 

General  characters. — A  small  form,  similar  to  O.  a.  chapmani  in 
color,  but  skull  narrower,  lighter,  and  more  like  that  of  the  darker 
subspecies  O.  a.  saturatior. 

14521°— 18 5 


66  NORTH   AMERICAN   FAUNA.  [No.  43. 

Color.— Type  in  fresh  pelage:  Upperparts  rich  ochraceous-tawny, 
finely  and  uniformly  lined  with  black  from  nose  over  top  of  head 
and  back  to  near  base  of  tail,  the  tawny  element  becoming  purer  and 
paler  on  cheeks  and  along  lower  part  of  sides;  underparts  and  feet 
dull  whitish;  tail  dark  brownish,  except  basal  half  of  under  side, 
which  is  dull  yellowish.  Specimens  from  Ocuilapa,  Chiapas,  appar- 
ently referable  to  this  form,  are  indistinguishable  in  color  from  0.  a. 
chapmani. 

SJcull. — Size  small,  general  form  long  and  narrow,  structure  light. 
Scarcely  distinguishable  from  some  of  the  narrower  skulls  of  0.  a. 
saturatior;  zygomata  very  slender  as  in  that  form.  Similar  in 
general  to  that  of  0.  a.  chapmani,  but  narrower,  maxillary  arm  of 
zygoma  more  slender. 

Measurements. — Type:  Total  length,  209;  tail  vertebrae,  106; 
hind  foot,  25.  Adult  from  Tumbala,  Chiapas:  222;  116;  27.  STcull 
(type):  Greatest  length,  26.7;  zygomatic  breadth,  12.6;  interorbital 
breadth,  4.8;  width  of  braincase,  10.2;  nasals,  10.2;  anterior  palatine 
foramina,  3.8;  palatal  bridge,  5.5;  upper  molar  series,  3.8. 

Remarks. — In  0.  a.  palatinus  the  general  color  of  0.  a.  chapmani 
seems  to  be  combined  with  the  narrower,  lighter  skull  of  the  darker- 
colored  form  0.  a.  saturatior.  The  scanty  material  available  indicates 
that  intergradation  of  palatinus  with  saturatior  is  probable,  since  the 
differential  characters  are  slight  and  both  forms  inhabit  the  northern 
slope  of  the  same  mountain  range. 

Specimens  examined. — Total  number,  5,  as  follows: 

Chiapas:  Ocuilapa,  3;  Tumbala,  1. 
Tabasco:  Teapa,  1  (type). 

ORYZOMYS  ALFAROI  SATURATIOR  Merriam. 

Dusky  Rice  Rat. 

Oryzomys  chapmani  saturatior  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  290, 
July  26,  1901. 

Type  locality. — Tumbala,  Chiapas,  Mexico  (altitude,  5,000  feet). 

Type. — No.  76183,  ?  adult,  United  States  National  Museum  (Bio- 
logical Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A.  Gold- 
man, October  23,  1895. 

Geographic  distribution. — Forested  northern  slope  of  mountains  of 
northern  Chiapas,  at  5,000  feet  altitude,  limits  of  range  unknown; 
Humid  Upper  Tropical  Zone. 

General  characters. — Size  rather  small — about  as  in  O.  a.  alfaroi  and 
0.  a.  chapmani;  color  very  dark.  Closely  allied  to  O.  a.  palatinus 
and  chapmani,  but  darker  than  either,  the  back  blackish  and  under- 
parts usually  distinctly  buffy. 

Color. — Upperparts  cinnamon  brownish,  very  heavily  mixed  with 
black,  the  top  of  head  and  median  dorsal  area  blackish ;  underparts, 


1918.]  ORYZOMYS  ALFAROI   GROUP.  67 

except  chin  and  throat,  varying  from  light  to  dark  ochraceous-buff ; 
chin  and  throat  more  or  less  distinctly  whitish,  the  fur  in  some 
examples  pure  white  to  roots  along  a  narrow  median  line;  ears  black; 
feet  brownish  to  toes,  the  toes  dull  yellowish,  those  of  hind  feet  bear- 
ing the  silvery  terminal  bristles  present  throughout  the  group;  tail 
usually  dark  all  around,  but  in  some  examples  yellowish  on  under 
side  at  base.  Young  (in  first  pelage) :  Upperparts  nearly  black,  the 
cinnamon  brownish  hairs  inconspicuous  and  restricted  mainly  to 
the  sides;  underparts  dark  plumbeous,  very  thinly  overlaid  with 
ochraceous-buff;  feet  and  tail  blackish. 

Skull. — Size  small  and  structure  light.  About  like  that  of  0.  a. 
palatinus,  but  usually  broader.  Similar  to  that  of  0.  a.  chapmani, 
but  averaging  slightly  smaller,  with  less  swollen  rostrum  and  more 
slender  zygomata. 

Measurements. — Type:  Total  length,  218;  tail  vertebras,  120;  hind 
foot,  25.5.  Average  of  10  adults  (type  and  9  topotypes) :  208.4  (195- 
225);  111.1  (105-122);  26.1  (25-28).  Skull  (average  of  3  adults- 
type  and  2  topotypes):  Greatest  length,  25.3  (25.2-25.4);  zygomatic 
breadth,  12.8  (12.4-13.5);  interorbital  breadth,  4.7  (4.5-4.8);  width 
of  braincase,  10.1  (10-10.3);  nasals,  9.8  (9.5-10.2);  anterior  palatine 
foramina,  4.2  (3.7-4.5);  palatal  bridge,  5.1  (4.9-5.3);  upper  molar 
series,  3.6  (3.5-3.6). 

Remarks. — The  very  dark  colors  of  O.  a.  saturatior  appear  to  be 
directly  due  to  environmental  conditions.  The  area  inhabited  by 
this  form  is  heavily  forested  and  excessively  humid ;  torrential  rains 
occur  during  the  wet  season  and  fog  enshrouds  the  mountains  nearly 
throughout  the  year.  Two  specimens  from  Tumbala,  with  narrow 
skulls  and  slender  rostra,  approach  O.  a.  palatinus,  whose  range  is 
doubtless  contiguous,  and  point  to  intergradation  with  that  form. 

Specimens  examined. — Seventeen,  from  type  locality. 

ORYZOMYS  ALFAROI  CHAPMANI  Thomas. 
Chapman  Rice  Rat. 

Oryzomys  chapmani  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  7,  I,  p.  179,  February,  1898. 

Type  locality. — Jalapa,  Vera  Cruz,  Mexico  (altitude,  4,400  feet). 

Type.— 97.9.9.30,  British  Museum;  collected  by  Frank  M.  Chap- 
man, March  31,  1897. 

Geographic  distribution. — Forested  eastern  slopes  of  the  Mexico 
plateau  region  in  central  Vera  Cruz;  known  altitudinal  range  from 
4,400  to  6,000  feet;  Humid  Upper  Tropical  Zone. 

General  characters. — Closely  resembling  O.  a.  alfaroi;  size  and 
color  very  similar,  but  general  tone  of  upperparts  averaging  slightly 
richer,  more  tawny;  cranial  characters  distinctive.  Size  about  as  in 
O.  a.  dilutior,  but  color  slightly  more  tawny;  skull  much  less  massive; 


68  NORTH    AMERICAN    FAUNA.  [No.  43. 

differing  from  0.  a.  caudatus  in  much  smaller  size  and  more  tawny 
color. 

Color. — Fresh  pelage:  Upperparts  near  ochraceous-tawny  (becom- 
ing russet  in  some  old  adults),  finely  and  abundantly  mixed  with 
black,  the  tawny  element  purer  but  somewhat  paler  and  grading 
toward  dark  ochraceous-buff  along  cheeks  and  lower  part  of  sides; 
underparts  whitish  or  pale  buffy,  the  plumbeous  basal  color  showing 
through  except  over  a  narrow  median  area  on  throat  and  chest, 
where  the  fur  in  some  specimens  is  pure  white  to  roots;  ears  black, 
light  subauricular  spots  usually  present;  feet  and  tail  as  given  under 
0.  alfaroi.  Young  (in  first  pelage) :  Upperparts  blackish,  inconspicu- 
ously mixed  with  ochraceous-tawny;  underparts  darker  than  in 
adults,  the  dark  basal  color  less  heavily  overlaid  with  white;  feet 
and  tail  brownish  (becoming  paler  in  adults). 

Skull. — Size  and  general  form  much  as  in  0.  a.  alfaroi,  but  tending 
to  be  shorter  and  relatively  broader,  the  zygomata  more  widely  or 
squarely  spreading  anteriorly;  frontal  region  usually  narrower;  ante- 
rior palatine  foramina  longer,  commonly  reaching  anterior  plane  of 
first  molars;  dentition  about  the  same.  Similar  to  that  of  0.  a. 
dilutior,  but  less  massive;  rostrum  less  swollen  and  decurved;  maxil- 
lary arm  of  zygoma  more  slender.  Much  like  that  of  0.  a.  caudatus, 
but  much  smaller. 

Measurements. — Average  of  10  adult  topotypes:  Total  length,  224 
(202-265);  tail  vertebras,  117.9  (108-132);  hind  foot,  25.1  (24-27). 
Skull  (average  of  7  adult  topotypes) :  Greatest  length,  26.4  (25.4- 
27.3);  zygomatic  breadth,  13.7  (13.1-14.4);  interorbital  breadth,  4.7 
(4.4-4.8) ;  width  of  braincase,  10.7  (10.2-11);  nasals,  10.3  (9.6-10.9); 
anterior  palatine  foramina,  4.4  (4-4.7);  palatal  bridge,  5.3  (5-5.6); 
upper  molar  series,  3.8  (3.7-3.9). 

Remarks. — The  general  resemblance  of  O.  a.  chapmani  to  O.  a. 
alfaroi  is  noteworthy,  in  view  of  their  geographic  separation  and  the 
occurrence  of  apparently  annectent  forms  that  differ  considerably 
from  both.  The  annectent  forms,  however,  inhabit  diversified  areas, 
some  having  ascended  to  high  elevations,  where  their  differentiation 
has  probably  been  the  result  of  rather  local  environmental  conditions. 

Specimens  examined. — Total  number,  25,  as  follows: 
Vera  Cruz:  Jalapa,  17  (type  and  topotypes);1  Jico,  6;  Mirador,  1;  Teocelo,  l.2 

ORYZOMYS  ALFAROI  DILUTIOR  Mereiam. 
Puebla  Rice  Rat. 

(PI.  Ill,  figs.  6,  6a.) 
Oryzomys  chapmani  dilutior  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  290,  July 
26,  1901. 

Type  locality. — Huauchinango,  Puebla,  Mexico  (altitude  5,000 
feet). 

1  Thirteen  in  collection  Amer.  Mus.  Nat.  Hist.  '  Collection  Mus.  Comp.  Zool. 


1918.]  ORYZOMYS  ALFAROI   GROUP.  69 

Type. — No.  93124,  <?  adult,  United  States  National  Museum  (Bio- 
logical Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A.  Gold- 
man, January  10,  1898. 

Geographic  distribution. — Eastern  slope  of  Mexican  plateau  region 
at  5,000  feet  altitude  in  northern  Puebla;  limits  of  range  unknown; 
Humid  Upper  Tropical  Zone. 

General  characters. — Similar  in  size  to  0.  a.  chapmani,  but  color 
slightly  paler,  less  tawny;  skull  more  massive. 

Color. — About  as  in  0.  a.  chapmani,  but  slightly  paler,  the  general 
tone  less  tawny. 

SJcull. — Most  like  that  of  0.  a.  chapmani,  but  more  massive;  ros- 
trum more  swollen,  the  upper  outline  more  strongly  decurved;  maxil- 
lary arm  of  zygoma  heavier. 

Measurements. — Type:  Total  length,  223;  tail  vertebrae,  117;  hind 
foot,  28.  An  adult  topotype:  225;  125;  27.5.  SJcull  (type  and  adult 
topotype):  Greatest  length,  27.6,  27.4;  zygomatic  breadth,  15,  14.2; 
interorbital  breadth,  4.9,  4.6;  width  of  braincase,  10.8,  10.4;  nasals, 
10.3,  10.5.;  anterior  palatine  foramina,  4.6,  4.4;  palatal  bridge,  5.2,  5; 
upper  molar  series,  4,  4. 

Remarks. — The  range  of  O.  a.  diluiior  marks  the  northern  limit  of 
the  O.  alfaroi  group.  The  subspecies  appears  to  be  a  well-marked 
form,  requiring  close  comparison  only  with  0.  a.  chapmani. 

Specimens  examined. — Three,  from  type  locality. 

ORYZOMYS  GUERRERENSIS  Goldman. 

Guerrero  Rice  Rat. 

(PI.  Ill,  figs.  7,  7a.) 

Oryzomys  guerrerensis  Goldman,  Proc.   Biol.  Soc.  Washington,  XXXVIII,  p.   127, 
June  29,  1915. 

Type  locality. — Omilteme,  Guerrero,  Mexico  (altitude  8,000  feet). 

Type. — No.  127517,  <$  adult  (molars  moderately  worn),  United 
States  National  Museum  (Biological  Survey  collection);  collected 
by  E.  W.  Nelson  and  E.  A.  Goldman,  May  20,  1903. 

Geographic  distribution. — Forested  Pacific  slope  of  Sierra  Madre  in 
Guerrero  and  Oaxaca,  Mexico;  altitudinal  range  from  3,000  to  about 
8,000  feet;  Humid  Upper  Tropical  Zone. 

General  characters. — Size  small;  color  averaging  palest  of  the  North 
American  forms  of  the  O.  alfaroi  group.  Similar  in  general  to  O.  a. 
chapmani,  but  color  paler  and  skull  smaller  and  natter. 

Color. — Upperparts  varying  from  dark  ochraceous-buff  to  dark 
ochraceous-tawny,  purest  on  cheeks,  shoulders,  and  sides;  the  face, 
top  of  head,  and  back  darkened  by  a  moderate  admixture  of  black 
hairs;  underparts  dull  grayish  white,  the  dark  basal  color  of  the  fur 
everywhere  showing  through,  except  in  2  examples  out  of  8,  in  which 


70  NORTH   AMERICAN    FAUNA.  [No.  43. 

the  fur  is  pure  white  to  roots  along  a  narrow  median  line  on  chin  and 
throat;  outer  and  inner  sides  of  ears  well  clothed  with  deep  glossy- 
black  hairs;  feet  whitish;  tail  brownish  above,  irregularly  yellowish 
below  to  near  tip,  which  is  dusky  all  around.  Young  (in  first  pelage) : 
As  in  0.  a.  chapmani. 

Skull. — Small  and  short,  the  braincase  broad  and  somewhat  flat- 
tened. Similar  to  that  of  0.  a.  chapmani,  but  usually  smaller  and 
flatter;  zygomata  tending  to  curve  evenly  outward,  the  sides  less 
nearly  parallel;  sides  of  rostrum  more  tapering  anteriorly;  ascending 
branches  of  premaxillse  usually  broader  posteriorly;  maxillary  arm  of 
zygoma  more  slender;  incisors  smaller. 

Measurements. — Type:  Total  length,  220;  tail  vertebrae,  118;  hind 
foot,  27.  Adult  from  Pluma,  Oaxaca:  221 ;  116;  26.5.  Skull  (type) : 
Greatest  length,  26.3;  zygomatic  breadth,  14;  interorbital  breadth, 
4.9;  width  of  braincase,  10.6;  nasals,  10.1;  anterior  palatine  foramina, 
4;  palatal  bridge,  5.5;  upper  molar  series,  3.8. 

Remarks. — The  range  of  0.  guerrerensis  marks  the  northern  limit  of 
the  distribution  of  the  0.  alfaroi  group  along  the  western  slope  of  the 
mountains  bordering  the  Pacific  coast  of  Mexico.  As  in  other  south- 
ern groups,  the  general  range  of  0.  alfaroi  and  its  allies  seems  to  bifur- 
cate north  of  the  Isthmus  of  Tehuantepec,  the  forms  which  pass  farther 
to  the  northward  along  the  eastern  and  western  slopes  of  the  interior 
plateau  region  being  isolated  and  developing  differential  characters. 
The  ranges  of  0.  guerrerensis  and  0.  a.  chapmani  appear  to  be  com- 
pletely separated,  and  while  these  forms  differ  appreciably  in  numer- 
ous details,  they  agree  rather  closely  in  the  more  essential  features, 
and  may  prove  to  intergrade  through  the  much  larger  form  0.  a. 
caudatus.  The  latter,  however,  as  at  present  understood,  exhibits  so 
wide  a  departure  from  guerrerensis  that  close  comparison  seems  un- 
necessary. 

Specimens  examined. — Total  number,  8,  as  follows: 

Guerrero:  Omilteme,  6  (type  and  topotypes). 
Oaxaca:  Pluma,  2. 

ORYZOMYS  HYLOCETES  Merriam. 

Chiapas  Rice  Rat. 

(PI.  Ill,  figs.  8,  8a.) 

Oryzomys  hylocetes  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  291,  July  26,  1901. 

Type  locality. — Chicharras,  Chiapas,  Mexico  (altitude  3,500  feet). 

Type. — No.  77605,  d1  old,  United  States  National  Museum  (Bio- 
logical Survey  collection);  collected  by  E.  W.  Nelson  and  E.  A. 
Goldman,  February  14,  1896. 

Geographic  distribution. — Heavily  forested  Pacific  slope,  at  3,500 
feet,   of  mountains  along  continental  divide  in  extreme   southern 


1918.]  ORYZOMYS   TALAMANC^E   GEOUP.  71 

Chiapas,  and  doubtless,  adjacent  portions  of  Guatemala;  Humid 
Upper  Tropical  Zone. 

General  characters. — Size  small;  color  dark;  pelage  short;  skull  short 
and  relatively  broad  across  anterior  roots  of  zygomata.  Somewhat 
similar  to  0.  a.  angusticeps  and  0.  a.  rhabdops,  but  much  smaller 
than  either,  and  cranial  characters  distinctive. 

Color. — About  as  in  0.  a.  saturatior. 

STcull.—  General  form  short,  with  posteriorly  narrow  braincase  and 
anteriorly  divergent  zygomata;  anterior  palatine  foramina  rather 
long,  but  not  reaching  anterior  plane  of  first  molars;  molars  small. 
In  size  the  skull  is  perhaps  nearest  to  that  of  0.  a.  saturatior,  but  it 
differs  in  the  anterior  expansion  of  the  zygomata  and  departs  from 
that  of  its  larger  and  nearer  geographic  neighbor  0.  a.  angusticeps  in 
the  same  respect.  The  skull  is  similar  in  general  outline,  especially 
the  form  of  zygomata,  to  that  of  0.  a.  rhabdops,  but  very  much 
smaller. 

Measurements. — Type:  Total  length,  217;  tail  vertebrae,  118;  hind 
foot,  27.  SJcull  (type):  Greatest  length,  26.2;  zygomatic  breadth, 
13.6;  interorbital  breadth,  4.6;  width  of  braincase,  9.9;  nasals,  9.7; 
anterior  palatine  foramina,  4.5;  palatal  bridge,  5.3;  upper  molar 
series,  3.5. 

Remarks. — Two  examples  only  of  0.  liylocetes  are  known,  one  old 
and  in  worn  pelage,  the  other  very  young,  and  the  range  of  individual 
variation  the  form  may  present  can  not  now  be  determined.  The 
species  inhabits  the  lower  slope  of  the  mountains,  while  the  upper 
levels  of  the  same  range  are  occupied  by  0.  a.  angusticeps,  a  much 
larger  form,  with  longer  pelage  and  apparently  representing  a  distinct 
specific  type;  but  only  through  the  accession  of  new  material  can 
the  exact  relationship  of  the  two  be  made  clear.  The  skull  in  general 
contour  bears  a  striking  resemblance  to  that  of  the  otherwise  different 
Guatemalan  form,  0.  a.  rhabdops. 

Specimens  examined. — Two,  from  type  locality. 

Oryzomys  talamancse  Group. 

Geographic  distribution. — Forested  districts  in  eastern  Costa  Rica 
and  Panama,  and  south  in  South  America  at  least  to  southern  Brazil; 
altitudinal  range  from  sea  level  to  3,000  feet  in  Panama;  Arid  and 
Humid  Lower  Tropical  Zones  (fig.  6). 

General  characters. — Size  large;  form  rather  slender;  tail  about  equal 
to  or  somewhat  exceeding  head  and  body,  scantily  and  indistinctly 
haired;  ears  large  and  conspicuous,  thinly  clothed  externally  and 
internally,  with  very  short,  fine,  almost  microscopic  hairs,  general 
pelage  short,  rather  harsh,  and  lacking  the  woolly  quality  of  0. 
palustris;  vibrissas  about  as  long  as  head;  toes  of  hind  feet  more  or 
less  distinctly  webbed  at  base,  the  longest  three  bearing  conspicuous 


72  NORTH   AMERICAN    FAUNA.  [No.  43. 

tufts  of  silvery  bristles  which  extend  beyond  the  points  of  the  claws; 
claws  short,  recurved,  compressed,  and  sharp  pointed.  Color  of  up- 
perparts  varying  from  rich  ochraceous-tawny  to  cinnamon  brown  or 
russet,  lined  with  black;  underparts  white  or  buffy  whitish. 

SJcull. — Size  large,  with  rostrum  long  and  braincase  low  and  some- 
what flattened;  maxillary  arm  of  zygoma  heavy;  outer  wall  of  ant- 
orbital  foramen  with  projecting  border  rounded  or  sloping  forward, 
the  antorbital  notch  rather  shallow  as  viewed  from  above;  nasals  and 
premaxillse  about  conterminous  posteriorly;  frontal  region  wide,  the 
lateral  margins  trenchant,  a  more  or  less  conspicuous  depression  on 
median  line  close  to  posterior  enpls  of  nasals;  temporal  ridges  moder- 
ately developed  anteriorly  along  parieto-squamosal  sutures,  becoming 
indistinct  posteriorly  in  crossing  lateral  wings  of  parietals;  interpa- 
rietal large;  anterior  palatine  foramina  short  and  broad,  much  shorter 
than  palatal  bridge,  not  reaching  anterior  plane  of  first  molars;  palatal 
pits   small;    spheno-palatine   vacuities    small;    interpterygoid    fossa 

broad;  audital  bullae  small,  the 
inner  sides  largely  overlapped 
by  mastoids;  basi-occipital 
broad;  angle  of  mandible  mod- 
erately broad,  the  inferior  bor- 
der strongly  turned  inward; 
coronoid  process  rather  short 
and  low;  molars  moderately 
heavy;  the  inner  reentrant  an- 
gles in  upper  molars  and  outer 
reentrant  angles  in  lower  mo- 
lars, deeply  cutting  the  crowns, 

Fig.  6. — Geographic  distribution  of  Oryzomys  talamancse.  , .,      -,  .        .,  i      •  i 

mandibular  tooth  row  decid- 
edly narrower  posteriorly  than  anteriorly;  tubercle  over  root  of  lower 
incisor  weakly  developed.  Contrasted  with  those  of  the  0.  palustris 
group,  skulls  of  0.  talamancse  and  its  allies  differ  especially  in  greater 
length  of  rostrum,  shorter  anterior  palatine  foramina  in  relation  to 
length  of  palatal  bridge,  smaller  size  of  audital  bullae,  and  in  dental 
detail;  the  upper  molars  are  more  deeply  cleft  by  inner  reentrant 
angles  and  the  lower  molars  by  outer  reentrant  angles;  the  second 
upper  molar  differs  especially  in  the  more  nearly  equal  size  of  the 
protocone  and  hypocone  (protocone  larger  than  hypocone  in  palus- 
tris),  owing  to  more  central  position  of  inner  reentrant  angle,  and  in 
the  absence  of  the  large,  elongated  enamel  island  extending,  in  the 
moderately  worn  tooth  of  palusiris,  along  the  postero-internal  base 
of  the  paracone,  or  the  reduction  of  this  island  to  a  smaller  one,  ap- 
pearing in  talamancse  near  the  mesostyle;  the  third  lower  molar  is 
more  triangular  in  outline  than  in  palustris,  the  posterior  division 
being  decidedlv  narrower  than  the  anterior. 


1918.]  ORYZOMYS  TALAMANCJE   GROUP.  73 

Remarks. — The  group  name  used  is  that  of  the  single  North  Ameri- 
can species,  0.  talamancse  of  Costa  Rica.  This  species  typifies  a  series 
of  wide  distribution  in  South  America;  0.  mollipilosus  and  0.  medius 
are  closely  allied  Colombian  and  Venezuelan  forms,  and  others  of 
this  unrevised  group  range  far  south  in  Brazil. 

The  0.  talamancse  group  is  more  nearly  related  to  the  0.  bombycinus 
group  than  to  any  other  North  American  section  of  the  genus,  and 
representatives  of  the  two  sometimes  share  the  same  local  habitat. 
Members  of  the  talamancss  group  are  externally  separable  by  the  short- 
ness of  their  pelage,  while  the  heavier  rostrum  and  the  extension 
of  the  lateral  wings  of  the  parietals  below  the  temporal  ridges  are 
distinguishing  cranial  characters.  In  dentition  the  two  groups  agree 
very  closely. 

ORYZOMYS  TALAMANCA  Allen. 
Talamanca  Rice  Rat. 

(PI.  IV,  figs.  3,  3a;  PI.  V,  fig.  7;  PI.  VI,  figs.  4,  4a.) 

Oryzomys  talamancse  Allen,  Proc.  U.  S.  Nat.  Mus.,  XIV,  p.  193,  July  24,  1891. 

Oryzomys  panamensis  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  7,  VIII,  p.  252,  September, 
1901.  Type  from  City  of  Panama,  Panama.  No.  0.5.1.67,  $,  British  Mus.;  col- 
lected by  E.  Andre\  February  25,  1899. 

Oryzomys  carrikeri  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXIV,  p.  656,  October  13, 
1908.  Type  from  Rio  Sicsola,  Talamanca,  Costa  Rica.  No.  25976,  9  ad.,  Amer. 
Mus.  Nat.  Hist. ;  collected  by  M.  A.  Carriker,  jr.,  August  18,  1904. 

Type  locality. — Talamanca,  Costa  Rica  (probably  near  Sipurio,  in 
the  valley  of  the  Rio  Sicsola). 

Type. — No.  ||f||,  c?  adult  (molars  much  worn),  United  States 
National  Museum;  collected  by  W.  M.  Gabb. 

Geographic  distribution. — Heavily  forested  regions  from  eastern 
Costa  Rica  eastward  through  Panama  to  near  Colombian  frontier; 
altitudinal  range  from  sea  level  to  3,000  feet;  Arid  and  Humid  Lower 
Tropical  Zones. 

General  characters. — Externally  similar  to  O.  mollipilosus  of  South 
America;  skull  narrower.  (For  additional  general  characters  see 
under  0.  talamancx  group.) 

Color. — General  color  of  upperparts  varying  from  pale  ochraceous- 
tawny  to  cinnamon  brown  or  russet,  becoming  lighter  and  in  some 
specimens  ochraceous-buffy  on  cheeks,  shoulders,  and  sides;  under- 
pays dull  white  or  buffy  whitish,  the  plumbeous  basal  color  showing 
through;  ears  brownish,  indistinctly  clothed  externally  with  very 
short  dusky  hairs  and  internally  with  almost  microscopic  buffy  or 
grayish  hairs;  feet  (epidermis)  dull  yellowish,  thinly  covered  above 
with  short  glossy  white  hairs,  the  hind  feet  with  tufts  of  silvery 
bristles  projecting  beyond  claws  of  longer  digits;  tail  (epidermis) 
dark  brownish  above,  varying  from  light  brownish  to  dull  yellowish. 


74  NORTH   AMERICAN   FAUNA.  TNo.  43. 

Skull. — About  like  that  of  0.  mollipilosus,  but  braincase  and  frontal 
region  decidedly  broader. 

Measurements. — Type  (dry  skin) :  Total  length,  233 ;  tail  vertebrae, 
115;  hind  foot,  30.  Average  of  three  adults  from  Rio  Sicsola,  Costa 
Rica:  251  (240-265);  124  (118-133);  29.8  (29-30.5).  Skull  (type): 
Greatest  length,  31.5;  zygomatic  breadth,  15.5;  interorbital  breadth, 
5.3;  width  of  braincase,  11.2;  nasals,  12.8;  anterior  palatine  foramina, 
4.3 ;  palatal  bridge,  7.1 ;  upper  molar  series,  4.6. 

Remarks. — Specimens  from  the  Canal  Zone  and  as  far  east  as 
extreme  eastern  Panama  seem  referable  to  typical  0.  talamancx  of 
Costa  Rica.  Two  examples  from  Gatun,  Canal  Zone-,  have  been  sub- 
mitted to  Mr.  Oldfield  Thomas  for  comparison  with  the  type  of  "  0. 
jianamensis"  in  the  British  Museum.  The  result  of  his  examination 
he  has  kindly  written  as  follows:  "We  have  only  one  specimen  of 
0.  panamensis  [the  type]  and  it  is  both  larger  and  more  rufous  than 
your  specimens.  But  it  is  older;  the  skull  agrees  in  general  characters 
and  the  toothrow  is  of  exactly  the  same  length.  As  to  the  colour  I 
think  the  difference  is  only  due  to  the  coming  on  of  the  faded  fulvous 
stage  found  in  the  old  specimens  of  most  species  of  Oryzomys.  Per- 
sonally I  should  certainly  refer  your  specimens  to  panamensis." 
Since  the  examples  used  for  comparison  are  regarded  as  fairly  typical 
of  talamancx  I  conclude  that  panamensis  must  be  placed  in  the  syn- 
onymy of  that  species. 

The  type  and  two  topotypes  of  "0. carrikeri"  Allen  agree  essentially 
with  the  type  of  talamancx.  The  exact  locality  of  the  latter  is  not 
definitely  known,  but  Gabb,  the  collector,  worked  mainly  near  Sipurio 
and  probably  secured  the  specimen  there.  The  Carriker  collection 
came  from  about  halfway  between  Cuabre  and  the  mouth  of  the  Rio 
Sicsola,  the  two  localities  being,  on  this  assumption,  not  far  apart  in 
the  same  river  valley. 

The  ranges  of  talamancse  and  0.  alfaroi  overlap,  and  owing  to  their 
superficial  resemblance  the  two  have  sometimes  been  confused; 
talamancx  is  a  larger  animal  than  alfaroi,  with  a  longer  hind  foot.  The 
skulls,  however,  present  the  safest  distinguishing  characters,  that  ot 
talamancx  being  more  massive,  with  heavier  maxillary  arms  of  zygo- 
mata, and  heavier  dentition;  the  first  and  second  upper  molars, 
besides  differing  in  details  of  crown  arrangement,  are  more  deeply  cleft 
by  inner  reentrant  angles  and  all  the  lower  molars  by  outer  reentrant 
angles,  while  in  alfaroi  they  more  nearly  approach  the  condition 
shown  in  0.  palustris. 

Specimens  examined. — Total  number,  20,  as  follows: 

Costa  Fvica:Boruca,  l;1  Rio  Sicsola,  3  (including  type  of ' '  carrikeri ") ;2  Talamanca, 

1  (type). 
Panama:  Cana,  7;  Cerro  Brujo,  1;  Divala,  1; J  Gatun,  6. 

1  Collection  Mus.  Comp.  Zool.  2  Collection  Amer.  Mus.  Nat.  Hist. 


1918.] 


ORYZOMYS   BOMBYCINUS   GROUP. 


75 


Oryzomys  bombycinus  Group . 

Geographic  distribution. — Forested  regions  from  southern  Costa 
Rica  south  through  Panama  at  least  to  northern  Ecuador ;  altitudinal 
range  from  near  sea  level  to  at  least  3,000  feet;  Humid  Lower  Tropical 
Zone  (fig.  7). 

General  characters. — Size  large;  form  rather  slender;  tail  about 
equal  to  or  slightly  longer  than  head  and  body,  scantily  and  very 
indistinctly  haired;  pelage  of  upperparts  very  long  (that  of  back 
measuring  about  12  mm.),  underfur  not  woolly  as  in  0.  palustris; 
supraorbital  vibrissa?  50  to  70 
mm.  in  length  (exceeding  those 
arising  from  the  muzzle) ,  reach- 
ing posteriorly  to  sides  of  body ; 
ears  rather  large  and  naked 
appearing,  the  short,  fine  hairs 
scattered  over  inner  and  outer 
surfaces  almost  microscopic ; 
hind  feet  slender,  very  scantily 
haired  above,  the  plantar  sur- 
face generally  smooth,  but  in 
some  examples  indistinctly 
granular  between  interdigital 
tubercles ;  toes  of  hind  feet  more 
or  less  distinctly  webbed  at 
base,  the  longest  three  bearing  conspicuous  tufts  of  silvery  bristles  pro- 
jecting beyond  ends  of  claws  as  in  the  0.  talamancse  group ;  claws  short, 
recurved,  compressed,  and  sharp  pointed.  Colors  dark;  upperparts 
near  cinnamon  brown  or  russet;  underparts  overlaid  with  dull  white. 

STcuM. — Size  rather  large,  with  rostrum  long  and  narrow,  the  nasals 
attenuate  and  slightly  exceeding  premaxillaB  in  posterior  extent; 
braincase  broad  and  moderately  expanded;  zygomata  slender;  outer 
wall  of  antorbital  foramen  with  projecting  upper  anterior  border 
well  rounded,  the  antorbital  notch  broad  but  not  deeply  cutting 
zygoma  as  viewed  from  above;  frontal  region  moderately  broad  and 
somewhat  flattened,  the  small  depression  on  median  line  near  ante- 
rior border  deep  and  conspicuous,  and  more  or  less  involving  ends 
of  nasals ;  lateral  margins  of  f rontals  rising  in  distinct,  but  narrow, 
compressed  ridges;  temporal  ridges  well  developed,  bounding  parie- 
tals  laterally  as  far  as  lateral  extension  of  supraoccipital,  where 
they  turn  abruptly  and  are  continued  downward  along  the  anterior 
border  of  that  segment;  parietals  without  lateral  wings  extending 
across  temporal  ridges;  interparietal  large,  reaching  transversely 
nearly  across  posterior  parietal  border;  anterior  palatine  foramina 
short,  not  reaching  anterior  plane  of  first  molars;  palatal  pits 
small;  spheno-palatine  vacuities  very  small  or  absent;  interpterygoid 


1.  O.  6.  bombycinus.         [        2.  O.  b.  alleni. 

Fig.  7. — Geographic  distribution    of    subspecies   of 

Oryzomys  bombycinus. 


76  NORTH    AMERICAN    FAUNA.  [No.  43. 

fossa  broad ;  audital  bulla?  small ;  basi-occipital  broad ;  angle  of  mandi- 
ble broad,  the  inferior  border  strongly  turned  inward;  coronoid  process 
rather  large  and  broad  across  base;  molars  moderately  heavy,  the 
crown  arrangement  of  the  0.  talamancse  type,  the  crowns  deeply  incised 
by  reentrant  angles  and  the  mandibular  toothrowmuch  narrower  pos- 
teriorly; second  upper  molar  about  as  in  talamancse,  the  elongated 
enamel  island  present  in  0.  palustris,  0.  melanotis  and  0.  alfaroi  groups, 
being  absent  or  displaced  by  a  bending  backward  of  the  commissure  of 
the  paracone  and  protocone ;  third  lower  molar  long  and  narrow,  the 
large  outer  reentrant  angle  extending  more  than  halfway  across  the 
crown. 

Remarks. — Forms  of  this  group  differing  only  subspecifically  among 
themselves  evidently  range  from  Costa  Rica  to  Ecuador  and  may 
extend  much  farther;  the  group,  therefore,  seems  to  be  mainly  South 
American  in  distribution.  Owing  to  lack  of  definite  knowledge  of 
the  relationships  of  the  South  American  forms  it  seems  best  to  treat 
the  North  American  representatives  as  specifically  distinct  until  a 
revision  of  the  South  American  species  can  be  undertaken. 

In  many  important  characters,  especially  of  the  skull  and  dentition, 
the  0.  hombycinus  group  approaches  the  0.  talamancse,  group  and  the 
two  sometimes  occur  together  in  the  same  localities.  The  hombycinus 
group  is  easily  recognizable  externally  by  the  much  greater  length 
of  the  pelage,  while  the  skull  is  sufficiently  distinguished  by  the  slender 
rostrum  and  the  absence  of  the  lateral  wings  of  the  parietals,  which  in 
the  talamancse  group  extend  across  the  temporal  ridges  at  the  expense 
of  the  squamosals.  Unlike  many  groups  of  the  genus  which  show  a 
preference  for  open  or  partially  open  situations  0.  hombycinus  seems 
to  be  at  home  in  the  depth  of  the  forest. 

ORYZOMYS  BOMBYCINUS  Goldman. 

[Synonymy  under  subspecies.) 

Geographic  distribution. — Heavily  forested  areas  from  southern 
Costa  Rica  to  eastern  Panama,  and  probably  western  Colombia; 
altitudinal  range  from  sea  level  to  at  least  3,000  feet;  Humid  Lower 
Tropical  Zone. 

General  characters. — (See  under  0.  hombycinus  group). 

Color. — General  color  of  upperparts  varying  from  ochraceous- 
tawny  to  cinnamon  brown  or  russet  finely  mixed  with  black;  darkest 
on  face,  top  of  head,  and  back,  the  lighter  element  restricted  to  tips 
of  hairs  and  the  dark  basal  color  showing  through,  becoming  fighter 
and  approaching  ochraceous-buff  or  pale  ochraceous- tawny  on  cheeks, 
shoulders,  and  sides;  underparts  dull  whitish;  ears  black,  the  outer 
and  inner  sides  thinly  clothed  with  very  short  dusky  hairs;  feet 
(epidermis)  dull  yellowish,  the  hairs  whitish;  tail  dark  brownish  above, 
light  brownish  below.      Young   (about  two-thirds  grown) :  Upper- 


1918.]  ORYZOMYS  BOMBYCINUS   GROUP.  77 

parts  blackish,  nearly  pure  black  on  face,  top  of  head,  and  back, 
becoming  lighter  and  more  distinctly  lined  with  ochraceous-buffy 
hairs  on  cheeks,  shoulders,  and  sides. 

Skull. — (See  under  0.  hombycinus  group.) 

Remarks. — Two  closely  allied  forms  inhabit  Panama  and  Costa 
Rica,  and  may  not  improbably  prove  to  be  geographic  races  of 
0.  nitidus  of  Peru,  since  Ecuadorean  specimens  believed  to  be  near 
nitidus  show  close  alliance  to  the  North  American  forms. 

Key  to  Subspecies  of  O.  bombycinus. 

a1.  Braincase  less  inflated.     (Panama.) O.  b.  bombycinus  (p.  77). 

a2.  Braincase  more  inflated.     (Costa  Rica.) O.  b.  alleni  (p.  78). 

ORYZOMYS  BOMBYCINUS  BOMBYCINUS  Goldman. 

Long-Haired  Rice  Rat. 

(PI.  II,  figs.  7,  7a;  PI.  V,  fig.  8;  PI.  VI,  figs.  5,  5a.) 

Oryzomys  bombycinus  Goldman,  Smiths.  Miec.  Coll.,  LVI,  No.  36,  p.  6,  February  19, 
1912. 

Type  locality. — Cerro  Azul,  near  headwaters  of  Chagres  River, 
Panama  (altitude,  2,500  feet). 

Type.— No.  171105,  6*  adult,  United  States  National  Museum 
(Biological  Survey  collection);  collected  by  E.  A.  Goldman,  March 
26,  1911. 

Geographic  distribution. — Mountains  of  east-central  Panama;  alti- 
tudinal  range  from  1,000  to  3,000  feet;   Humid  Lower  Tropical  Zone. 

General  characters. — Size  about  as  in  0.  b.  alleni,  color  identical; 
skull  with  much  natter,  less  expanded  braincase. 

Color. — (See  under  0.  bombijcinus.) 

Skull. — Similar  to  that  of  0.  b.  alleni,  but  braincase  lower,  flatter, 
much  less  distended,  especially  anteriorly;  frontal  region  narrower 
posteriorly. 

Measurements. — Type:  Total  length,  252 ;  tail  vertebra?,  127;  hind 
foot,.  32.  Skull  (type  and  adult  topotype):  Greatest  length,  31.3, 
31.7;  zygomatic  breadth,  15.4,  16.1;  interorbital  breadth,  5.5,  5.4; 
width  of  braincase,  11.2,  11.6;  nasals,  12.2,  12.2;  anterior  palatine 
foramina,  5,  4.5;  palatal  bridge,  6.2,  6.1 ;  upper  molar  series,  4.7,  4.4. 

Remarks. — The  Panama  form  apparently  differs  from  the  Costa 
Rican  animal  only  in  cranial  characters,  and  the  specimens  on  which 
it  is  based  closely  resemble  Ecuadorean  examples  believed  to  be 
near  0.  nitidus.  The  four  Panama  specimens  are  very  slightly  paler 
in  color  than  five  from  Ecuador,  with  skulls  also  similar,  but  the 
zygomata  more  strongly  bowed  outward,  the  sides  less  nearly  parallel. 

Specimens  examined. — Total  number,  4,  as  follows: 
Panama:  Cerro  Azul,  3  (type  and  topotypes);  Cerro  Brujo,  1. 


78  NORTH  AMERICAN   FAUNA.  [No.  43. 

ORYZOMYS  BOMBYCINUS  ALLENI  Goldman. 
Allen  Rice  Rat. 

(PI.  II,  figs.  8,  8a.) 

Oryzomys  nitidus  alleni  Goldman,  Proc.  Biol.  Soc.  Washington,  XXVIII,  p.  128, 
June  29,  1915. 

Type  locality. — Tuis  (about  35  miles  east  of  Cartago),  Costa  Rica. 

Type. — No.  -f-ffl,  S  sub  adult  (teeth  slightly  worn),  American 
Museum  of  Natural  History;  collected  by  George  K.  Cherrie,  July 
15,  1894. 

Geographic  distribution. — Mountainous  portions  of  southern  Costa 
Rica;  altitudinal  range  from  8<3o  to  about  2,000  feet;  Humid  Lower 
Tropical  Zone. 

General  characters. — Size  about  as  in  0.  b.  bombycinus;  color  iden- 
tical;  skull  with  braincase  higher,  much  more  fully  expanded. 

Color. — As  in  0.  b.  bombycinus.     (See  under  0.  bombycinus  group.) 

Skull. — Similar  to  that  of  0.  b.  bombycinus,  but  braincase  higher, 
more  arched,  much  more  distended,  especially  anteriorly;  frontal 
region  broader  posteriorly. 

Measurements. — Type  tyoung):  Total  length,  218;  tail  vertebrae, 
111,  hind  foot,  29.  Skull  (type):  Greatest  length,  28.2;  zygomatic 
breadth,  14.3;  interorbital  breadth,  5.5;  width  of  braincase,  11.6; 
nasals,  10.9;  anterior  palatine  foramina,  3.5;  palatal  bridge  5.5; 
upper  molar  series,  4.3. 

Remarks. — Cranial  comparisons  are  necessary  in  order  to  distin- 
guish this  Costa  Rican  form  from  0.  b.  bombycinus  of  Panama. 
While  the  interorbital  breadth  is  about  the  same,  the  greater  anterior 
expansion  of  the  braincase  has  resulted  in  broadening  the  posterior 
part  of  the  frontal  region. 

Specimens  examined. — -Total  number,  4,  as  follows 
Costa  Rica:  Tuis,  3  (type  and  topotypes);  Guapiles,  l.1 

Oryzomys  devius  Group. 

Geographic  distribution. — High  mountains  of  central  Costa  Rica  to 
Colombia  and  Venezuela;  altitudinal  range  from  4,000  to  at  least 
5,000  feet;  Humid  Upper  Tropical  Zone  (fig.  8). 

General  characters. — Size  very  large,  but  form  rather  slender;  tail 
much  longer  than  head  and  body,  indistinctly  haired;  ears  rather 
large,  blackish,  the  short  hairs,  thinly  distributed  over  inner  and  outer 
sides,  of  about  the  same  color  as  the  epidermis;  pelage  long,  but 
somewhat  rigid;  vibrissa  long  (about  reaching  shoulders);  hind  feet 
long  and  narrow,  thinly  haired  above,  the  plantar  surface  between 
the  large  conspicuous  tubercles  smooth;  toes  of  hind  feet  with  the 
slight  basal  webbing  usual  in  the  genus,  the  longest  three  tufted  with 

If  i  Collection  Carnegie  Mus. 


1918.] 


ORYZOMYS  DEVIUS   GROUP. 


79 


silvery  bristles  as  in  0.  talamancse  and  other  groups;  claws  short, 
recurved,  compressed  and  sharp  pointed  as  in  other  partially  scan- 
sorial  sections  of  the  genus.  Colors  dark;  upperparts  between  tawny 
and  russet ;  underparts  varying  from  ochraceous-tawny  to  dull  white, 
the  basal  color  plumbeous  except  in  some  examples,  which  tend  to 
exhibit  irregular  areas  on  throat  where  the  fur  may  be  pure  white 
to  roots. 

STcull. — Size  very  large,  with  rostrum  long  and  heavy;  nasals 
broad  and  reaching  posteriorly  slightly  beyond  premaxillfe;  braincase 
moderately  broad  and  inflated;  zygomata  heavy;  the  maxillary  arm 
extensively  overlapping  outer  side  of  jugal,  in  some  examples  in  con- 
tact with  squamosal,  the  gap  usual  in  the  genus  being  completely 
bridged;  outer  wall  of  antorbital  foramen  with  projecting  upper 
anterior  border  rounded,  the  antorbital  notch  broad  as  viewed  from 
above,  but  less  deeply  cutting 
zygoma,  the  aperture  appear- 
ing less  evenly  circular  than  in 
the  0 .  palustris  group;  frontal 
region  narrow,  especially  pos- 
teriorly, the  anterior  median 
depression  rather  inconspicu- 
ous ;  lateral  margins  of  f  rontals 
smoothly  rounded,  or  rising  in 
slightly  upturned  ridges;  tem- 
poral ridges  moderately  devel- 
oped, crossing  lateral  wings  of 
parietals  to  supraoccipital  and 
continuing  thence  downward  in 
prominent  crests  along  occipito- 
squamosal  border;  interparietal  very  large,  extending  transversely 
nearly  across  posterior  parietal  border,  pointed-elliptical  in  outline 
owing  to  convexity  of  anterior  margin ;  anterior  palatine  foramina  very 
short,  not  reaching  anterior  plane  of  first  molars,  narrow  anteriorly, 
very  broad  and  gaping  widely  open  posteriorly,  palatal  pits  large  and 
irregular  in  form;  spheno-palatine  vacuities  absent;  interpterygoid 
fossa  moderately  broad  and  extending  well  forward,  the  anterior  bor- 
der closely  approaching  posterior  plane  of  last  molars;  audital  bulla? 
variable  but  rather  small;  angle  of  mandible  broad,  the  inferior  border 
strongly  incurved ;  coronoid  process  large  and  strongly  upturned,  the 
high  thin  connecting  ridge  extending  to  near  summit  of  condyle ; 
molars  heavy  and  approaching  the  0.  talamancse,  type,  the  crowns  of 
the  upper  series  deeply  incised  by  inner  reentrant  angles,  and  of  lower 
series  by  outer  reentrant  angles;  as  in  the  talamancse  group  the 
second  upper  molar  is  evenly  cleft  by  inner  reentrant  angle  and  lacks 


1.  O.  devius.  2.  O.  pirrensis. 

Fig.  8. — Geographic  distribution  of  the  Oryzomys  devius 

group. 


80  NORTH   AMERICAN   FAUNA.  [No.  43. 

the  elongated  enamel  island  extending  in  the  moderately  worn  crown 
of  0.  palustris  and  other  groups  along  the  postero-internal  base  of  the 
paracone;  mandibular  toothrow  very  narrow  posteriorly,  the  third 
molar  elongated  and  deeply  cleft  by  outer  reentrant  angle. 

Remarks. — The  0.  devius  group,  as  it  may  be  called,  includes  two 
northern  representatives  of  a  widely  distributed  South  American 
section  of  the  genus  which  in  that  country  has  sometimes  been 
denominated  the  0.  meridensis  group,  from  the  name  of  a  Vene- 
zuelan species. 

These  northern  forms  generally  lack  the  irregular  but  extensive  pure 
white  pectoral  and  inguinal  areas  exhibited  by  the  allied  Venezuelan 
and  Colombian  forms  0.  meridensis  and  0.  maculiventer,  although  a 
tendency  to  develop  them  is  shown  in  0.  devius. 

The  North  American  species  are  externally  easily  recognizable 
among  their  congeners  of  the  general  region  by  the  combination  of 
large  size  with  dark  color  and  very  long  tails.  Except  for  the  more 
hispid  pelage  they  are  superficially  much  like  the  species  of  Pero- 
myscus  (subgenus  Megadontomys)  occurring  at  the  same  localities 
with  them. 

Key  to  Species  of  the  O.  devius  Group. 

a1.  Supraorbital  ridges  prominent.     (Eastern  Panama.) O.  pirrensis  (p.  81) . 

a2.  Supraorbital  ridges  not  prominent.    (Western  Panama.) O.  devius  (p.  80). 

ORYZOMYS  DEVIUS  Bangs. 

Chiriqui  Rice  Rat. 
(PL  IV,  figs.  1,  la.) 

Oryzomys  devius  Bangs,  Bull.  Mus.  Comp.  Zool.,  XXXIX,  p.  34,  figs,  13, 14,  April, 
1902. 

Type  locality. — Boquete,  Volcan  do  Chiriqui,  Panama  (altitude, 
5,000  feet). 

Type.— No.  10324,  ?  adult,  Museum  of  Comparative  Zoology 
(Bangs  collection) ;  collected  by  W.  W.  Brown,  jr.,  January  29,  1901. 

Geographic  distribution. — Forested  slopes  of  high  mountains  in 
central  Costa  Rica  and  western  Panama;  altitudinal  range  from  4,000 
to  at  least  5,000  feet;  Humid  Upper  Tropical  Zone. 

General  characters. — Size  large,  about  as  in  0.  pirrensis  of  eastern 
Panama,  but  color  slightly  paler,  more  tawny;  skull  more  smoothly 
rounded;  audital  bullae  much  larger. 

Color. — Upperparts  dark  tawny,  inclining  toward  russet  and  rather 
heavily  mixed  with  black  along  median  line  of  dorsum,  becoming  light 
tawny,  or  in  the  paler  examples  dark  ochraceous-buffy  along  lower 
part  of  sides;  throat  whitish  (the  fur  pure  white  to  roots  over  a 
^mall  area  in  one  example),  rest  of  underparts  overlaid  with  ochra- 
ceous-buffy in  three  examples  and  with  dull  white  in  the  other  three 
examined;  nose  and  ears  blackish;  feet  dull  yellowish  or  light  brown- 


1918.]  ORYZOMYS  DEVIUS   GROUP.  81 

ish;  tail  (epidermis)  dark  brownish  above,  paler  below.  Young  (in 
first  pelage) :  Upperparts  blackish,  the  tawny  element  appearing 
rather  inconspicuously  along  sides;  underparts  (in  single  specimen 
examined)  thinly  overlaid  with  dull  white. 

Slcull. — Similar  to  that  of  0.  pirrensis,  but  more  smoothly  rounded, 
the  supraorbital  and  temporal  ridges  weakly  developed  or  absent; 
nasals  slightly  longer,  ending  posteriorly  in  plane  of  lachrymals;  zygo- 
mata less  widely  spreading;'  audital  bullae  decidedly  larger.  Compared 
with  that  of  0.  meridensis,  the  skull  is  larger,  with  longer  nasals  and 
larger  audital  bulla?. 

Measurements. — Type:  Total  length,  335;  tail  vertebra?,  ISO;  hind 
foot,  33.  Two  adult  topotypes:  345,  360;  185,  195;  36,  35.  Slcull 
(average  of  three  adults,  type,  and  two  topotypes) :  Greatest  length, 
36.6  (35.8-37.5);  zygomatic  breadth,  18.4  (18-19);  interorbital 
breadth,  5.7  (5.5-6);  width  of  braincase,  12.6  (12.2-12.9);  nasals, 
14.3  (13.8-15);  anterior  palatine  foramina,  5.4  (5.2-5.6);  palatal 
bridge,  7.8  (7.8-7.9);  upper  molar  series,  5.7  (5.6-5.8). 

Remarks. — 0.  devius  is  clearly  allied  to  0.  pirrensis  of  eastern 
Panama,  but  possesses  very  distinctive  cranial  characters,  and  there 
is  no  hint  of  intergradation.  Both  are  inhabitants  of  high  moun- 
tains, and  their  ranges  are  apparently  separated  by  the  intervening 
lowlands  in  the  vicinity  of  the  Canal  Zone. 

Specimens  examined. — Total  number,  8,  as  follows: 

Costa  Rica:  Volcan  Irazu,  2.1 

Panama:  Boquete,  6  (type  and  topotypes).2 

ORYZOMYS  PIRRENSIS  Goldman. 

Mount  Pirre  Rice  Rat. 

(PI.  IV,  figs.  2,  2a;  PI.  V,  fig.  9;  PI.  VI,  figs.  6,  6a.) 

Oryzomys  pirrensis  Goldman,  Smiths.  Misc.  Coll.,  LX,  No.  22,  p.  5,  February  28,  1913. 

Type  locality. — Head  of  Rio  Limon,  Mount  Pirre,  Panama  (altitude 
4,500  feet). 

Type.— No.  178993,  <?  adult,  United  States  National  Museum  (Bio- 
logical Survey  collection) ;  collected  by  E.  A.  Goldman,  April  29,  1912. 

Geographic  distribution. — Steep,  heavily  forested  slopes  of  high 
mountains  at  4,500  feet  altitude  in  eastern  Panama,  and  probably 
adjacent  portions  of  Colombia;  Humid  Upper  Tropical  Zone. 

General  characters. — Size  large,  about  as  in  0.  devius  of  western 
Panama,  but  color  slightly  darker,  more  russet;  skull  more  angular; 
audital  bulla?  decidedly  smaller. 

Color. — Fresh  pelage:  Upperparts  between  tawny  and  russet,  heav- 
ily mixed  with  black  along  the  broad  median  line  from  top  of  head  to 

1  One  specimen  in  Mus.  Comp.  Zool. 

2  Four  specimens  in  Mus.  Comp.  Zool.;  2  in  Field  Mus.  Nat.  Hist. 

14521°— 18 6 


82  NORTH    AMERICAN    FAUNA.  [No.  43. 

base  of  tail,  becoming  lighter,  purer  tawi^  on  cheeks,  shoulders,  and 
sides;  throat  whitish  or  grayish,  rest  of  underparts  usually  overlaid 
with  ochraceous-tawny,  but  varying  to  dull  white,  the  basal  color  of 
the  fur  everywhere  deep  plumbeous;  nose  and  ears  blackish;  fore  feet 
blackish,  becoming  lighter  on  toes;  hind  feet  dark  brown,  thinly 
clothed  with  short  hairs  to  toes,  the  toes  dull  yellowish;  epidermis  of 
tail  dark  brown  above,  usually  somewhat  paler  below.  Young  (in 
first  pelage) :  Upperparts  darker  than  in  a*dults,  the  blackish  element 
in  the  pelage  predominant;  underparts  with  a  thinner  ochraceous- 
tawny  wash. 

Skull. — Similar  to  that  of  0.  devius,  but  more  angular,  the  supra- 
orbital and  temporal  ridges  well  developed;  nasals  slightly  shorter, 
not  reaching  posteriorly  to  plane  of  lachrymals;  zygomata  more 
widely  spreading;  audital  bulla?  decidedly  smaller.  In  small  size  of 
audital  bullas  the  skull  agrees  with  those  of  0.  meridensis  and  0.  macu- 
liventer,  but  contrasts  with  both  in  larger  general  size  and  angularity. 

Measurements. — Type:  Total  length,  340;  tail  vertebrae,  185;  hind 
foot,  38.  Average  of  five  adult  topotypes:  314  (309-322);  164  (159- 
170) ;  35.7  (34-37).  Skull  (average  of  six  adults,  type,  and  five  topo- 
types):  Greatest  length,  36.8  (34.9-38.5);  zygomatic  breadth,  19.2 
(17.8-20);  interorbital  breadth,  5.8  (5.5-6);  width  of  braincase,  12.4 
(11.8-12.9);  nasals,  13.7  (13-14);  anterior  palatine  foramina,  5.8  (5.4- 
6);  palatal  bridge,  7.5  (7.3-7.8);  upper  molar  series,  5.7  (5.5-5.9). 

Remarks. — In  external  appearance  0.  pirrensis  differs  only  slightly 
from  0.  devius  of  western  Panama.  The  skull,  however,  combines 
the  large  general  size  of  that  species  with  the  smaller  general  dimen- 
sions and  small  audital  bullse  of  0.  meridensis  and  0.  maculiventer;  it 
differs  from  both  in  the  development  of  the  supraorbital  and  temporal 
ridges.  In  color  of  upperparts  0.  pirrensis  is  similar  to  the  South 
American  species,  but  the  underparts  show  no  trace  of  the  pure  white 
pectoral  and  inguinal  areas  which  are  so  conspicuous  in  the  latter 
forms. 

Specimens  examined. — Eight,  from  type  locality. 

Oryzomys  tectus  Group. 

Geographic  distribution. — Southern  Costa  Rica  and  southeastward 
through  Panama  at  least  to  Colombia  and  Venezuela;  altitudinal 
range  from  near  sea  level  to  about  5,000  feet  (fig.  9). 

General  characters. — Size  large,  form  rather  robust;  tail  about  equal 
to  or  somewhat  exceeding  head  and  body,  scantily  haired,  the  epi- 
dermal scales  small;  ears  small,  moderately  clothed  externally  and 
internally  with  comparatively  coarse  hairs  of  general  body  color; 
general  pelage  long,  coarse,  and  rigid;  vibrissas  somewhat  longer  than 
head;  hind  feet  short  and  relatively  broad,  well  haired  above,  the 
longest  three  toes  bearing  conspicuous  tufts  of  silvery  bristles,  which 


1918.] 


ORYZOMYS   TECTUS   GROUP. 


83 


project  beyond  ends  of  claws;  claws  short,  strongly  curved,  com- 
pressed, and  sharp-pointed.  Color  of  upperparts  between  rich  tawny 
and  ochraceous-tawny,  mixed  with  black,  the  tawny  element  pre- 
dominating ;  underparts  varying  from  nearly  pure  white  to  warm  buff. 

Skull. — Size  large  and  angular,  with  rostrum  short  and  braincase 
low  and  flattened;  outer  wall  of  antorbital  foramen  rounded  above, 
the  forward  projection  moderate;  nasals  short,  ending  posteriorly  in 
the  anterior  plane  of  orbits;  premaxillse  about  conterminous  with 
nasals  posteriorly,  the  ends  slightly  expanded,  not  beveled  externally ; 
frontals  very  broad,  the  lateral  margins  overhanging  as  supraorbital 
shelves;  temporal  ridges  prominent,  extending  posteriorly  to  supra- 
orbital border;  mastoid  process  of  squamosal  short  and  stout,  owing 
to  slight  excision  of  squamosal  margin;  interparietal  large,  pointed- 
elliptical,  the  anterior  angle  well  developed;  lachrymal  very  small; 
anterior  palatine  foramina 
short,  moderately  broad  anteri- 
orly and  posteriorly,  reaching 
or  nearly  reaching  anterior 
plane  of  first  molars;  palatal 
pits  normally  small;  spheno- 
palatine vacuities  absent  or 
very  small ;  audital  bullae  small ; 
ramus  of  mandible  short,  the 
angle  broad;  coronoid  process 
broad  and  strongly  hooked,  a 
high  trenchant  ridge  connecting 
with  condyle;  tubercle  over 
root  of  lower  incisor  large. 
Molars  similar  to  those  of  the 
0.  talamancse  group;  approaching  the  0.  palustris  type  in  general  char- 
acters, but  second  upper  molar  with  inner  lobes  more  nearly  equal 
in  extent  (the  anterior  slightly  the  larger  in  palustris),  and  central 
enamel  island  present  in  palustris  usually  fused  with  enamel  fold 
separating  paracone  and  parastyle;  third  lower  molar  more  triangular 
in  outline,  the  posterior  lobe  narrower  and  the  outer  reentrant  angle 
more  nearly  dividing  crown. 

Remarks.— -O.  tectus  is  typical  of  a  group  including  also  0.  flavicans, 
0.  palmarius,  and  other  extralimital  forms  distinguished  by  rich 
coloration,  coarsely  haired  ears,  and  short  stout  hind  feet.  In  external 
appearance  they  are  not  very  unlike  some  species  of  Rhipidomys, 
but  have  shorter,  less  hairy  tails  and  lack  the  dark  metapodial  mark- 
ings usually  present  in  that  genus;  in  general  characters  they  appar- 
ently approach  the  section  assigned  to  generic  rank  by  Thomas  under 
the  name  (E corny s,1  but  in  the  more  essential  respects  scarcely  exhibit 


1.  O.  t.  tectus.  |  2.  0.  t.  frontalis. 

Fig.  9. — Geographic  distribution  of  subspecies  of  Oryzo- 

mys  tectus. 


I  Thomas,  Oldfield,  Ann.  Mag.  Nat.  Hist.,  ser.  7,  XVIII,  p.  444,  Dec.  1906. 


84  NORTH   AMERICAN    FAUNA.  [No.  43. 

a  wider  departure  from  typical  Oryzomys  than  several  groups  usually- 
assigned  to  the  genus. 

ORYZOMYS  TECTUS  Thomas. 

[Synonymy  under  subspecies.] 

Geographic  distribution. — Southern  Costa  Rica,  Panama,  and  prob- 
ably adjacent  portions  of  Colombia;  vertical  range  from  near  sea 
level  to   about  2,000  feet,   mainly  in  Arid  Lower  Tropical  Zone. 

General  characters. — Similar  to  0.  jlavicans  of  Venezuela,  but 
larger;  color  darker;  skull  larger  with  laterally  expanded  frontal 
region.     (For  additional  generaKcharacters  see  under  0.  tectus  group.) 

Color. — Fresh  pelage:  General  color  of  upperparts  varying  shades 
of  tawny  and  ochraceous-tawny,  rather  inconspicuously  darkened  on 
face,  top  of  head,  and  back  by  an  admixture  of  dusky  hairs,  becom- 
ing paler,  more  ochraceous-buffy  on  cheeks,  shoulders,  and  sides; 
underparts  varying  from  near  pure  white  to  warm  buff;  ears  clothed 
internally  and  externally  with  short  tawny  hairs;  feet  whitish;  tail 
varying  from  uniform  dark  brownish  all  around  to  dark  brownish 
above,  and  whitish  or  yellowish  below. 

SJcull. — Size  large,  with  remarkably  broad  frontal  region.  Similar 
in  general  to  0.  Jlavicans,  but  decidedly  larger;  frontal  region  broader, 
the  lateral  margins  shelving  farther  over  orbits,  not  strongly  upturned 
as  in  Jlavicans.     (For  additional  characters  see  under  0.  tectus  group.) 

Remarks. — 0.  tectus  is  clearly  allied  to  0.  Jlavicans  Jlavicans,  O.f. 
illectus,  and  0.  palmarius,  all  South  American  forms  in  which  the 
supraorbital  ridges  are  well  developed,  but  are  compressed  and  not 
widely  expanded  and  Tylomys-like  as  in  the  forms  of  tectus.  This 
difference,  although  rather  striking,  is  a  relative  instead  of  absolute 
character,  and  close  agreement  in  other  essential  respects  even  sug- 
gests probable  intergradation. 

Key  to  Subspecies  of  O.  tectus. 

a1.  Upperparta   more   distinctly    tawny.      (Western    Panama   and   southern    Costa 

Rica.) O.  t.  tectus  (p.  84). 

a2.  Upperparts  less  distinctly  tawny.     (Eastern  Panama.) O.  t.  frontalis  (p.  85). 

ORYZOMYS  TECTUS  TECTUS  Thomas. 
Bugaba  Rice  Rat. 

Oryzomys  tectus  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  7,  VIII,  p.  251,  September,  1901. 

Type  locality. — Bugaba,  Chiriqui,  Panama  (altitude  800  feet). 

Type. — No.  0.7.11.43,  $  ,  British  Museum;  collected  by  H.  J. 
Watson,  September  15,  1898. 

Geographic  distribution. — Pacific  slope  of  western  Panama  and 
southern  Costa  Rica  at  about  800  feet  altitude,  limits  of  altitudinal 
range  unknown ;  Arid  Lower  Tropical  Zone. 


1918.]  OEYZOMYS   TECTUS   GROUP.  85 

General  characters. — Closely  resembling  0.  t.  frontalis,  but  upper- 
parts  richer  tawny;  underparts  more  extensively  buffy;  lips,  chin, 
and  throat  buffy  instead  of  white  as  in  frontalis;  skull  differing  in 
rather  slight  details. 

Color. — Upperparts  near  tawny  rather  thinly  mixed  with  black, 
general  color  darkest  over  dorsum,  becoming  paler  and  rich  ochra- 
ceous-buffy  on  cheeks  and  sides ;  underparts,  including  lips,  chin,  and 
throat  overlaid  with  warm  buff;  ears  clothed  with  tawny  hairs;  feet 
whitish;  tail  (epidermis  and  hairs)  brownish  above,  whitish  or  dull 
yellowish  below  to  near  tip,  which  is  dark  all  around. 

Skull. — Very  similar  to  that  of  0.  t.  frontalis,  but  frontals  more 
extended  posteriorly  on  median  line  between  parietals;  interparietal 
smaller. 

Measurements. — From  original  description  of  type:  "Head  and 
body,  140  millim. ;  tail,  142;  hind  foot,  s.  u.  27,  c.  u.  29.5;  ear,  18. 
Skull:  Tip  of  nasals  to  back  of  interparietal,  33;  greatest  breadth,  17; 
nasals,  11.6  X  4 ;  interorbital  breadth,  6.5 ;  palate  length,  13.8;  diastema 
8.1;  palatal  foramina,  5X2.2;  length  of  upper  molar  series,  4.9." 
A  rather  young  example  from  Boruca,  Costa  Rica:  Total  length,  258; 
tail  vertebrae,  140;  hind  foot,  27. 

Remarks. — The  type  of  0.  t.  tectus  has  not  been  examined  by  me, 
but  specimens  from  Boruca,  Costa  Rica,  are  believed  to  be  typical. 
They  are  distinguished  from  0.  t.  frontalis  of  eastern  Panama  mainly 
by  richer  general  coloration.  No  other  member  of  the  0.  tectus  group 
ranges  so  far  into  the  Central  American  Subregion. 

Specimens  examined. — Two,  as  follows: 

Costa  Rica:  Boruca,  2. 

ORYZOMYS  TECTUS  FRONTALIS  Goldman. 

Corozal  Rice  Rat. 

(PI.  IV,  figs.  4,  4a;  PI.  V,  fig.  10;  PI.  VI,  figs.  7,  7a.) 

Oryzomys frontalis  Goldman,  Smiths.  Misc.  Coll.,  LV1,  No.  36,  p.  6,  February  19,  1912. 

Type  locality. — Corozal,  Canal  Zone,  Panama  (altitude  100  feet). 

Type. — No.  171531,  ?  adult,  United  States  National  Museum 
(Biological  Survey  collection);  collected  by  E.  A.  Goldman,  June 
20,  1911. 

Geographic  distribution. — Forested  Pacific  slope  of  Panama  from 
the  Canal  Zone  to  near  Colombian  frontier,  and  probably  adjacent 
Colombian  territory;  altitudinal  range  from  sea  level  to  about  2,000 
feet;   Arid  and  Humid  Lower  Tropical  Zones. 

General  characters. — Similar  in  general  to  0.  t.  tectus,  but  upperparts 
duller,  less  distinctly  tawny;  underparts  less  extensively  buffy;  lips, 
chin,  and  throat  white  instead  of  buffy  as  in  tectus;  skull  differing  in 
rather  slight  details. 


86  NORTH   AMERICAN    FAUNA.  [No.  43. 

Color. — Upperparts  between  tawny  and  ochraceous-tawny,  rather 
sparingly  mixed  dorsaUy  with  black,  the  general  color  becoming  paler, 
purer,  and  ochraceous-buffy  on  cheeks  and  sides;  underparts  in 
general  white  in  some  examples,  thinly  overlaid  with  warm  buff 
across  the  abdomen  in  others,  the  lips,  chin,  and  throat  in  all  speci- 
mens examined  nearly  pure  white  to  roots  of  hairs;  ears  and  feet  as 
in  0.  t.  tectus;  tail  usually  unicolor,  dark  brownish,  but  in  some 
examples  becoming  lighter  on  under  side  near  base. 

STcull. — Closely  resembling  that  of  0.  t.  tectus,  but  frontals  less 
extended  posteriorly  on  median  line  between  parietals ;  interparietal 
larger. 

Measurements. — Type:  Total  length,  309;  tail  vertebrae,  161 ;  hind 
foot,  30.     Average  of  five  adults  from  Cana,  Panama:  288  (281-293) ; 
156  (152-164);    27.8  (26.5-29).     Skull  (average  of  same) :    Greatest 
length,    32.8    (31.1-34.1);     zygomatic    breadth,    17.1     (16.6-17.5) 
interorbital  breadth,  6.1  (5.3-6.8);  width  of  braincase,  12.2  (12-12.6) 
nasals,  11.4  (10.5-12.4);    anterior  palatine  foramina,  5.3  (4.9-5.6) 
palatal  bridge,  6.4  (5.6-6.9);    upper  molar  series,  5.2  (4.8-5.3). 

Remarks. — Two  specimens  of  0.  t.  frontalis  from  eastern  Panama 
have  been  submitted  to  Mr.  Oldfield  Thomas,  who  has  kindly  com- 
pared them  with  the  type  and  topotypes  of  0.  t.  tectus  in  the  British 
Museum.  While  inclined  to  regard  them  as  referable  to  the  same 
form  he  writes  that  "they  are  not  quite  so  rich  in  colour  as  our 
specimens."  Two  examples  from  Boruca,  Costa  Rica,  not  far  from 
the  type  locality  and  in  the  same  general  f  aunal  area,  and  believed  to 
be  fairly  typical  of  tectus,  exhibit  a  type  of  coloration  and  slight 
cranial  details  indicating  that  eastern  and  western  Panama  are 
inhabited  by  closely  allied  but  easily  recognizable  forms. 

Specimens  examined. — Total  number,  12,  as  follows: 

Canal  Zone:  Corozal,  1  (type). 
Panama:  Cana,  11. 

ORYZOMYS  VICTUS »  Thomas. 

St.  Vincent  Rice  Rat. 

Oryzomys  victus  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  7,  I,  p.  178,  February,  1898. 

Type  locality. — St.  Vincent,  Lesser  Antilles. 

Type. — No.  97.12.26.1,  9  adult,  British  Museum;  collected  by 
H.  H.  Smith,  presented  by  F.  DuCane  Godman. 

Geographic  distribution. — Known  only  from  St.  Vincent. 

General  characters. — From  original  description  of  type:  "Size  and 
proportions  about  as  in  the  larger  members  of  the  0.  longicaudatus 
group.  General  colour  dark  rufous,  but  evidently  affected  by  the 
spirit  in  which  the  specimen  has  been  preserved.     Under  surface 

1  Group  association  undetermined. 


1918.]  SUBGENUS  OLIGOEYZOMYS.  87 

buffy  white,  the  bases  of  the  hairs  slate-colour.  Eyes  without  darker 
rims.  Ears  short,  the  anterior  part  of  their  backs  brown,  not  strik- 
ingly contrasting  with  the  general  colour  of  the  head.  Hands  and 
feet  thinly  clothed  with  fine  silvery  hairs.  Tail  almost  naked,  brown 
above,  slightly  paler  below.     Mammae  2-2=8." 

SlcuU. — From  original  description  of  type:  "Skull  with  the  general 
shape  of  the  South-American  0.  longicaudatus,  the  braincase  being 
similarly  lengthened  as  compared  with  the  broadened  braincase  of 
the  Central-American  0.  melanotis  and  its  allies.  Compared  with  a 
Rio  Janeiro  example  it  is  larger,  more  rounded,  the  supraorbital 
edges  less  sharply  square,  but  the  parietal  ridges  thicker  and  better 
developed.  Molars  larger  and  stouter,  palate  ending  only  just 
behind  the  back  of  m3." 

Measurements. — From  original  description  of  type  (measured  in 
spirit):  "Head  and  body,  96  millim.;  tail,  121;  hind  foot  without 
claws,  25;  with  claws,  26.7;  ear,  14.  Skull:  Basilar  length,  21.4; 
basal  length,  23.8;  greatest  breadth,  15.1;  nasals,  11.2x3.4;  inter- 
orbital  breadth,  4.5;  interparietal,  3.2x10;  palate  length  from 
henselion,  12.3;  diastema,  7.8;  palatal  foramina,  5.4x1.8;  length 
of  upper  molar  series,  4.1." 

Remarks. — -The  type  of  0.  victus  has  not  been  examined  by  me.  It 
was  originally  compared  mainly  with  a  South  American  species,  but 
the  true  affinities  remain  to  be  determined.  As  in  the  case  of  0. 
antillarum,  of  Jamaica,  this  rice  rat  seems  likely  to  be  endangered  by 
the  presence  of  the  mongoose,  if  it  has  not  already  been  exterminated 
since  the  introduction  of  that  indiscriminately  destructive  animal. 

Subgenus  OLIGORYZOMYS  Bangs. 

Oligoryzomys  Bangs,  Proc.  New  England  Zool.  Club.,  I,  p.  94,  February  23,  1900  (sub- 
genus).    Type  Oryzomys  navus  Bangs. 

GeograpJiic  distribution. — Southern  Mexico  south  through  Central 
America  to  undetermined  limits  in  South  America. 

Subgeneric  characters. — Size  very  small;  hind  foot  usually  less  than 
25;  form  slender  and  Reithrodontomys-lake;  ears  rather  large  and 
coarsely  haired;  tail  much  longer  than  head  and  body;  four  longer 
toes  of  hind  feet  bearing  tufts  of  silvery  bristles  projecting  beyond 
ends  of  claws. 

Skull  delicate  in  structure,  smoothly  rounded;  supraorbital  and 
temporal  ridges  absent;  interorbital  region  narrow,  the  constriction 
about  equal  to  width  of  rostrum  between  antorbital  foramina ;  outer 
wall  of  antorbital  foramen  projecting  slightly  forward,  as  viewed  from 
above ;  angle  of  mandible  placed  well  within  vertical  plane  of  condyle ; 
molars  with  small  accessory  cusps  present  as  in  subgenus  Oryzomys, 
but  reentrant  angles  usually  broader,  the  salient  angles  formed  by 
worn  crowns  of  tubercles  less  evenly  rounded;    upper  molars  early 


88  NORTH   AMERICAN    FAUNA.  [No.  43. 

exhibiting  small,  circular  central  enamel  islands  which  persist  until 
obliterated  by  wear  in  extreme  old  age;  second  upper  molar  with 
protocone  and  hypocone  of  about  equal  size,  the  inner  reentrant 
angle  central  in  position;  third  lower  molar  with  outer  reentrant 
angle  extending  less  than  halfway  across  crown;  tubercle  over  root 
of  lower  incisor  large. 

Remarks. — The  subgenus  Oligoryzomys  includes  a  series  of  forms 
easily  distinguishable  among  their  North  American  congeners  by 
diminutive  size  and  external  resemblance  to  species  of  the  genus 
Reiihrodontomys .  Oligoryzomys,  however,  departs  from  the  subgenus 
Oryzomys,  as  currently  restricted,  mainly  in  a  combination  of  relative 
rather  than  absolute  characters.  The  molar  crowns  differ  in  details 
of  enamel  arrangement,  the  second  upper  especially,  in  the  early 
appearance  of  a  single,  persistent,  normally  circular  enamel  island  in 
the  broad  central  space  between  the  apex  of  the  inner  reentrant  angle 
and  the  base  of  the  paracone.  In  the  subgenus  Oryzomys  this  molar 
crown  varies  in  pattern;  the  more  typical  forms  normally  present,  in 
early  stages  of  wear,  an  elongated,  crescentic  enamel  island  in  the 
central  space,  but  in  more  divergent  forms  the  enamel  island  may  be 
absent  or  tend  to  unite  with  the  long,  deep  enamel  fold  between  the 
paracone  and  parastyle.  Several  groups  seem  at  least  as  fully 
entitled  to  subgeneric  recognition  as  Oligoryzomys,  but  the  problem  of 
further  subgeneric  divisions  can  best  be  solved  when  more  compre- 
hensive study  of  the  genus  is  undertaken. 

ORYZOMYS  FULVESCENS  (Saussure). 
[Synonomy  under  subspecies.) 

Geographic  distribution. — From  southern  Mexico  south  through 
Guatemala,  Honduras,  Nicaragua,  and  Costa  Kica  to  Panama,  and 
probably  portions  of  Colombia ;  altitudinal  range  from  near  sea  level 
to  about  5,500  feet;  mainly  Arid  and  Humid  Lower  Tropical  Zones 
(fig.  10). 

General  characters. — {See  subgeneric  characters  under  subgenus 
Oligoryzomys.) 

Color. — Upperparts  varying  from  light  ochraceous-buff  to  ochra- 
ceous-buff  or  tawny,  most  intense  on  rump,  palest  and  purest  on 
cheeks,  shoulders,  and  sides;  the  face,  top  of  head,  and  back  mod- 
erately darkened  by  blackish  hairs;  underparts  varying  from  nearly 
pure  white  in  general  to  light  ochraceous-buff  on  abdomen,  inguinal 
region,  and  inner  sides  of  hind  limbs ;  the  lips,  throat,  and  fore  limbs 
white ;  outer  sides  of  ears  blackish,  inner  sides  clothed  with  ochra- 
ceous-buffy  hairs;  vibrissas  black  (about  as  long  as  head);  feet  white, 
the  hind  feet  with  tufts  of  silvery  hairs  projecting  beyond  claws  of 
longest  four  digits;  tail  dark  brownish  above,  lighter  brown  or  yel- 
lowish below,  except  toward  tip,  which  is  dusky  all  around.     Young 


1918.] 


OEYZOMYS  FULVESCENS. 


89 


(in  first  pelage) :  Usually  darker  than  adults,  the  ochraceous-buff  or 
tawny  element  less  developed. 

Skull. — (See  subgeneric  characters  under  subgenus  Oligoryzomys.) 
Remarks. — The  North  American  forms  of  the  subgenus  are  all 
referable  to  a  single  species  whose  range  probably  extends  well  into 
South  America.  They  agree  closely  among  themselves  in  essential 
characters,  and  by  small  size  and  slender  form  are  sufficiently  distin- 
guished from  other  sections  of  the  genus  inhabiting  the  same  region. 


1.  O././ulvcscens.  4.  O.f.  costaricensis. 

2.  O.f.lenis.  5.  O./.vegetus. 

3.  O.f.  mayensis. 

Fig.  10. — Geographic  distribution  of  subspecies  of 
Oryzomy  s  fulvescens. 


Key  to  Subspecies  of  O.  fulvescens. 
a1.  Size  smaller;  hind  foot  less  than  24. 
b1.  Upperparts  darker  ochraceous-buff. 
c1.  Upper  molar  series  shorter.     (Southern  Tamaulipas;  eastern  Oaxac a;  Chiapas; 

northern  Honduras.) O.  f.  fulvescens  (p.  89). 

c2.  Upper  molar  series  longer.     (Southwestern  Panama;  Costa  Rica;  Nicaragua.) 

O.  f.  costaricensis  (p.  92). 
b2.  Upperparts  paler  ochraceous-buff. 
cl.  Skull    broader;  zygomatic    breadth    11.5    or    more.     (Michoacan;  Guerrero; 

southern  Oaxaca.) O.  f.  lenis  (p.  91). 

c2.  Skull  narrower;  zygomatic  breadth  less  than  11.5.     (Yucatan;  Campeche.) 

O.  f.  mayensis  (p.  92). 
a2.  Size  larger;  hind  foot  24  or  more.     (Mountains  of  western  Panama.) 

O.  f.  vegetus  (p.  93). 

ORYZOMYS  FULVESCENS  FULVESCENS  (Saussure). 

Vera  Cruz  Pygmy  Rice  Rat. 

[PI.  IV,  figs.  5,  5a;  PI.  V,  figs.  2, 11;  PI.  VI,  figs.  8,  8a.] 

Hesperomys fulvescens  Saussure,  Rev.  et  Mag.  Zool.,  ser.  2,  XII,  p.  102,  March,  1860. 
Oryzomys  fulvescens  Allen  and  Chapman,  Bull.  Amer.  Mus.  Nat.  Hist.,  IX,  p.  204, 
June  16,  1897. 

Type  locality. — Orizaba,1  Vera  Cruz,  Mexico. 


1  Type  locality  fixed  by  Merriam,  Proc.  Washington  Acad.  Sci.,  Ill,  p.  295,  July  26.  1901. 


90  NORTH   AMERICAN    FAUNA.  [No.  43. 

Type. — In  Geneva  Museum  of  Natural  History. 

Geographic  distribution. — Southern  Tamaulipas,  Vera  Cruz,  eastern 
Oaxaca,  Chiapas,  and  east  through  central  Guatemala  to  eastern 
Honduras;  altitudinal  range  from  near  sea  level  to  about  5,500  feet; 
mainly  Arid  and  Humid  Lower  Tropical  Zones. 

General  characters. — Size  small;  molar  tooth  series  short.  Similar 
to  O.f.  lenis  but  upperparts  darker  ochraceous-buff,  the  general  tone 
less  yellowish;  skull  narrower  and  less  massive.  Size  about  as  in 
O.f.  costaricensis ,  but  upperparts  usually  less  tawny;  molar  series 
shorter. 

Color. — Fresh  pelage:  Upperparts  near  ochraceous-buff,  most  in- 
tense and  in  some  examples  suffused  with  tawny  on  rump,  palest  and 
purest  on  cheeks,  shoulders,  and  sides;  the  face,  top  of  head,  and  back 
moderately  darkened  by  blackish  hairs;  underparts  varying  from 
nearly  pure  white  throughout  in  rare  examples  to  warm  buff  on 
abdomen,  inguinal  area,  and  inner  sides  of  hind  limbs,  the  white  ap- 
pearing only  on  lips,  chin,  and  inner  sides  of  hind  limbs;  outer  sides  of 
ears  blackish,  inner  sides  clothed  with  ochraceous-buffy  hairs;  feet 
whitish;  tail  brownish  above,  yellowish  below,  except  toward  tip, 
which  is  dusky  all  around.  Young  (in  first  pelage) :  Darker  than 
adults,  the  ochraceous-buff  of  upperparts  restricted  to  narrow  tips  of 
hairs,  thus  permitting  plumbeous  basal  color  to  show  through  and 
alter  general  tone. 

Skull. — About  like  that  of  O.f.  costaricensis,  but  molar  tooth  series 
shorter.  Contrasted  with  0.  f.  lenis,  the  skull  is  narrower,  with 
less  widely  spreading  zygomata;  maxillary  arms  of  zygomata  and 
ascending  branches  of  premaxillae  less  broad  and  heavy. 

Measurements. — Average  of  10  adults  from  Orizaba,  Vera  Cruz: 
Total  length,  174.7  (168-205);  tail  vertebras,  107.1  (96-118);  hind 
foot,  22  (21-23).  Skull  (average  of  same):  Greatest  length,  21.9 
(21.1-22.5);  zygomatic  breadth,  11.5  (11-11.7);  interorbital  breadth, 
3.5  (3.4-3.9);  width  of  braincase,  9.5  (9.3-9.8);  nasals,  7.9  (7.6-8.3); 
anterior  palatine  foramina,  3.5  (3.3-3.6);  palatal  bridge,  3.7  (3.5-4); 
upper  molar  series,  2.9  (2.9-3). 

Remarks. — The  differential  characters  are  rather  slight,  and  0.  f. 
fulvescens  may  safely  be  assumed  to  intergrade  with  O.f.  lenis  near 
the  Pacific  coast  in  the  vicinity  of  the  Isthmus  of  Tehuantepec,  with 
0.  f.  costaricensis  in  eastern  Honduras  or  western  Nicaragua,  and 
with  the  paler  form  inhabiting  the  Yucatan  Peninsula,  in  southern 
Campeche.  A  single  specimen  from  Patuca,  Honduras,  the  only  one 
available  from  that  country,  has  a  short  molar  series  and  seems  refer- 
able to  the  subspecies  fulvescens,  the  known  range  of  which  is  thus 
materially  extended  eastward  from  western  Guatemala. 

Specimens  examined. — Total  number,  73,  as  follows: 
Chiapas:  Huehuetan,  4;  Ocuilapa,  1;  Tumbala,  11;  Yajalon,  1. 
Guatemala:  Jacaltenango,  3;  Nenton,  1. 


1918.]  OEYZOMYS  FULVESCENS.  91 

Honduras:  Patuca,  2. 

Oaxaca:  Santo  Domingo  (mountains  near),  2;  Tuxtepec,  1. 
Tamaulipas:  Alta  Mira,  1. 

Vera  Cruz:  Jalapa,  24;1  Jico,  2;  Mirador,  1;  Orizaba  (type  locality  by  fixation), 
17;  Pasa  Nueva,  l;2  Santiago  Tuxtla,  1. 

ORYZOMYS  FULVESCENS  LENIS  Goldman. 

Los  Reyes  Pygmy  Rice  Rat. 

(PI.  IV,  figs.  6,  6a.) 

Oryzomys  fulvescens  lenis  Goldman,  Proc.  Biol.  Soc.  Washington,  XXVIII,  p.  130, 
June  29,  1915. 

Type  locality. — Los  Reyes,  Michoacan,  Mexico. 

Type. — No.  125941,  <$  adult,  United  States  National  Museum  (Bio- 
logical Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A.  Gold- 
man, February  14,  1903. 

Geographic  distribution. — Coastal  plains  and  basal  mountain  slopes 
in  Michoacan,  Guerrero,  and  Oaxaca;  altitudinal  range  from  near  sea 
level  to  about  3,000  feet,  mainly  in  Arid  Lower  Tropical  Zone. 

General  characters. — Similar  to  O.  f.  fulvescens,  but  upperparts 
paler  ochraceous-buff ,  the  general  tone  more  yellowish ;  skull  broader 
and  more  massive. 

Color. — Fresh  pelage:  Upperparts  between  ochraceous-buff  and 
light  ochraceous-buff  becoming  warm  buff  on  cheeks,  shoulders,  and 
lower  part  of  sides;  the  face,  top  of  head,  and  back  sparingly  lined 
with  dark  hairs;  underparts,  ears,  feet,  and  tail  as  in  O.f.  fulvescens. 

SJcull. — Broader  than  that  of  O.  f.  fulvescens;  zygomata  more 
widely  spreading;  maxillary  arms  of  zygomata  and  ascending  branches 
of  premaxillse  broader  and  heavier;  dentition  rather  heavy,  but 
equaled  in  some  examples  of  O.  f.  fulvescens. 

Measurements. — Type:  Total  length,  198;  tail  vertebra?,  115;  hind 
foot,  23.  Adult  topotype:  188;  115;  23.  Skull  (type  and  adult 
topotype):  Greatest  length,  22.6,  21.8;  zygomatic  breadth,  12.3, 
11.9;  interorbital  breadth,  3.4,  3.5;  width  of  braincase,  10,  10.3; 
nasals,  8.6,  7.8;  anterior  palatine  foramina,  3.9,  3.5;  palatal  bridge, 
4.1,  3.7;  upper  molar  series,  3,  3. 

Remarks. — The  general  distribution  area  of  0.  fulvescens  is  divided 
north  of  the  Isthmus  of  Tehuantepec  by  the  high,  interior  plateau 
region,  the  western  arm  representing  the  range  of  O.f.  lenis.  Speci- 
mens from  Guerrero  and  Oaxaca  approach  subspecies  fulvescens  in 
characters. 

Specimens  examined. — Total  number,  4,  as  follows: 
Guerrero:  Ometepec  (near),  1. 
Michoacan:  Los  Reyes,  2  (type  and  topotypes). 
Oaxaca:  Pluma,  1. 

i  Sixteen  in  Amer.  Mus.  Nat.  Hist.;  2  in  Mus.  Comp.  Zool.         2  Collection  Amer.  Mus.  Nat.  Hist. 


92  NORTH   AMERICAN    FAUNA.  [No.  43. 

ORYZOMYS   FULVESCENS  MAYENSIS,  subsp.  nov. 
Maya  Pygmy   Rice   Rat. 

Type  locality. — Apazote  (near  Yohaltum),  Campeche,  Mexico 
(altitude  200  feet) . 

Type. — No.  107979,  <?  adult,  United  States  National  Museum 
(Biological  Survey  collection) ;  collected  by  E.  W.  Nelson  and  E.  A. 
Goldman,  January  5,  1901.     Original  number  14405. 

Geographic  distribution. — Peninsula  of  Yucatan  and  Campeche; 
altitudinal  range  from  near  sea  level  to  about  300  feet;  Arid  Lower 
Tropical  Zone. 

General  characters. — Similar  to  O.  f.  fulvescens,  but  paler  colored; 
skull  slightly  narrower. 

Color. — Fresh  pelage:  Upperparts  near  ochraceous-buff  shading 
toward  light  ochraceous-buff,  darkest  and  rather  sparingly  lined 
with  black  over  dorsum,  becoming  clearer  and  paler  on  cheeks, 
shoulders,  and  sides;  underparts,  ears,  feet,  and  tail  as  in  O.f.  ful- 
vescens. 

STcull. — About  like  that  of  O.  f.  fulvescens,  but  zygomata  and 
braincase  slightly  narrower. 

Measurements. — Type:  Total  length,  177;  tail  vertebrae,  101,  hind 
foot,  22.  Average  of  three  adult  topotypes:  189  (184-196);  113 
(108-119) ;  23  (22-24).  Skull  (average  of  three  adults,  type  and  two 
topotypes):  Greatest  length,  21.1  (20.4-21.8);  zygomatic  breadth, 
11  (10.8-11.2) ;  interorbital  breadth,  3.6  (3.5-3.8) ;  width  of  braincase, 
9  (8.9-9.1);  nasals,  8.1  (7.4-8.4);  anterior  palatine  foramina,  3.3 
(3.1-3.5);  palatal  bridge,  4  (3.6-4.4);  upper  molar  series,  2.9  (2.9-3). 

Remarks. — The  pale  color  of  this  form  seems  to  be  due  to  the  same 
environmental  conditions  that  have  been  operative  in  the  evolution 
of  pallid  races  in  other  mammalian  groups  inhabiting  Yucatan.  In 
this  region  pale  coloration  is  apparently  associated  with  the  arid 
climate  and  white  limestone  formation,  outcropping  or  thinly  over- 
laid with  soil  over  large  areas  which  have  risen  less  than  300  feet  above 
the  level  of  the  sea.  At  Chichen  Itza  the  pygmy  rice  rat  was  taken  in 
the  vicinity  of  imposing  Maya  Indian  ruins,  a  circumstance  that  sug- 
gested the  subspecific  name. 

Specimens  examined.— Total  number,  7,  as  follows: 

Campeche:  Apazote,  4  (types  and  topotypes). 
Yucatan:  Chichen  Itza,  1;  Tunkas,  2. 

ORYZOMYS  FULVESCENS  COSTARICENSIS  Allen 
Costa  Rican  Pygmy  Rice  Rat. 

Oryzomys  costaricensis  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  V,  p.  239,  September  22, 

1893. 
Oryzomys  (Oligoryzomys)  nicaragux  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXVIII,  p. 

100,  April  30,  1910.     Type  from  Vijagua,  Nicaragua,  No.  29543,   c?   ad.,  Amer. 

Mus.  Nat.  Hist. ;  collected  by  W.  B.  Richardson,  March  24, 1909. 


1918.]  ORYZOMYS   FULVESCENS.  93 

Type  locality. — El  General,  Costa  Rica  (altitude,  2,150  feet). 

Type. — No.  f§-fl>  ?  subadult,  American  Museum  of  Natural  His- 
tory, collected  by  H.  Pittier,  February,  1891. 

Geographic  distribution. — Southwestern  Panama  east  to  Canal  Zone, 
Costa  Rica,  and  north  to  northern  Nicaragua;  altitudinal  range  from 
near  sea  level  to  at  least  2,000  feet. 

General  characters. — Similar  to  0.  f  fulvescens,  but  upperparts 
usually  more  tawny;  molars  larger.  Also  similar  to  O.f.  vegetus,  but 
smaller  and  averaging  paler. 

Color. — About  as  in  0.  f.  fulvescens,  but  upperparts  usually  more 
distinctly  tawny. 

Skull. — Closely  resembling  that  of  O.f.  fulvescens,  but  molars 
larger,  the  toothrows  decidedly  longer.  General  size  smaller  than 
that  of  O.f.  vegetus;  molars  about  the  same. 

Measurements. — Two  adults  from  Buenos  Aires,  Costa  Rica: 
Total  length,  183,  190;  tail  vertebrae,  100,  108;  hind  foot,  20,  23. 
Skull  (same):  Greatest  length,  21.1,  21.5;  zygomatic  breadth,  11.4, 
11.4;  interorbital  breadth,  3.7,  3.7;  width  of  braincase,  9.9,  9.6; 
nasals,  8.2,  8;  anterior  palatine  foramina,  3.6,  3.7;  palatal  bridge, 
4,  4.2;  upper  molar  series,  3.2,  3.2. 

Remarks. — The  general  dimensions  of  O.f.  costaricensis  are  about 
the  same  as  those  of  O.f.  fulvescens;  the  color  difference  is  slight  but 
the  upperparts  are  usually  more  distinctly  tawny.  The  most  dis- 
tinctive feature  seems  to  be  the  larger  molars  and  resulting  elongation 
of  the  toothrows,  a  character  shared  with  the  larger  and  darker  form 
O.f  vegetus.  uO.  nicaraguae"  was  based  on  scanty  material  from 
Nicaragua  which  is  not  satisfactorily  separable  from  costaricensis. 
The  type,  apparently  full  grown  but  with  molars  rather  slightly  worn, 
is  not  so  tawny  as  most  examples  of  costaricensis,  but  this  color  ele- 
ment is  usually  less  distinct  in  younger  individuals.  The  toothrows 
are  long,  as  in  costaricensis. 

Specimens  examined. — Total  number,  19,  as  follows: 

Costa  Rica:  Boruca,  7;1  Buenos  Aires,  2;2  Cerro  de  Buena  Vista,  l;2  El  General, 

3  (type  and  topotypes);2  La  Carpintera,  l.2 
Nicaragua:  San  Rafael  del  Norte,  l;2  Vijagua,  1  (type  of  "nicaraguse  ").2 
Panama:  La  Chorrera,  l;2  Old  Panama,  2.2 

ORYZOMYS  FULVESCENS  VEGETUS  Bangs. 

Volcan  Chiriqui  Pygmy  Rice  Rat. 

Oryzomys  (Oligoryzomys)  vegetus  Bangs,  Bull.  Mus.  Comp.  Zool.,  XXXIX,  text  fig.  15, 
p.  35,  April,  1902. 

Type  locality. — Boquete,  Volcan  de  Chiriqui,  Panama  (altitude, 
4,000  feet). 

i  Six  in  Araer.  Mus.  Nat.  Hist.;  1  in  Field  Mus.  Nat.  Hist.         2  Collection  Amer.  Mus.  Nat.  Hist. 


94  NORTH    AMERICAN    FAUNA.  [No.  43. 

Type. — No.  10298,  9  old  adult,  Museum  of  Comparative  Zoology 
(Bangs  collection) ;  collected  by  W.  W.  Brown,  jr.,  April  16,  1901. 

Geographic  distribution. — Known  only  from  the  vicinity  of  type 
locality;  Upper  Tropical  Zone. 

General  characters. — Size  larger  and  color  usually  darker  than  in 
0.  f  costaricensis;  skull  larger,  but  molar  teeth  relatively  small. 

Color. — About  as  in  0 .  f.  fulvescens  and  O.f.  costaricensis,  but  gen- 
eral tone  averaging  darker  than  either,  the  tawny  element  in  upper- 
parts  distinct  as  in  costaricensis;  abdomen  varying  in  the  darkest 
examples  to  a  shade  darker  than  light  ochraceous-buff  of  Ridgway. 

Slcull. — Decidedly  larger  than  that  of  O.f.  costaricensis,  with  molar 
teeth  actually  about  the  same  size,  therefore  relatively  smaller. 

Measurements. — Type:  Total  length,  210;  tail  vertebrae,  120;  hind 
foot,  25.  Average  of  six  adult  topotypes:  216  (205-235);  122  (115- 
130) ;  24.2  (24-25) .  SJcull  (average  of  5  adults — type  and  4  topotypes) : 
Greatest  length,  23.6  (22.7-24.3);  zygomatic  breadth,  12.3  (11.9- 
12.8);  interorbital  breadth,  3.8  (3.7-4.2);  width  of  braincase,  10.3 
(10-10.5) ;  nasals,  8.7  (8.3-9.3) ;  anterior  palatine  foramina,  3.6  (3.4- 
3.9) ;  palatal  bridge,  3.9  (3.8-4.3) ;  upper  molar  series,  3.1  (3.1-3.1). 

Remarks. — Larger  average  size  and  tendency  toward  darker  colora- 
tion usually  distinguish  O.f.  vegetus  from  the  more  northern  forms  of 
the  group,  but  there  seems  to  be  no  sharp  line  of  demarcation  between 
it  and  O.f.  costaricensis.  Both  share  the  heavier  dentition  as  com- 
pared with  0.  f.  fulvescens.  Four  specimens  in  the  topotype  series 
were  referred  by  Bangs  to  costaricensis.  They  are  the  paler  examples 
and  in  color  apparently  do  not  differ  from  costaricensis,  but  the  larger 
size  of  the  skulls  indicates  that  they  belong  with  the  remainder  of  the 
series  of  vegetus. 

Specimens  examined. — Twenty-two,1  from  type  locality. 

Subgenus  MELANOMYS  Thomas. 

Melanomys  Thomas,  Ann.  Mag.  Nat.  Hist.,  ser.  7,  X,  p.  248,  Sept.,  1902;  Novitatea 
Zool.,  X,  no.  1,  p.  41,  Apr.  20,  1903  (subgenus).  Type  Oryzomys  phxopus 
Thomas. 

Melanomys  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXXII,  p.  533,  Nov.  17, 1913  (genus). 

Geographic  distribution. — Andean  region  of  northwestern  South 
America  from  southern  Ecuador  to  northern  Colombia,  and  north- 
westward through  Panama  and  Costa  Rica  to  northern  Nicaragua; 
altitudinal  range  from  near  sea  level  to  about  8,000  feet;  mainly  in 
Upper  and  Lower  Tropical  Zones. 

Subgeneric  characters. — Color  very  dark,  upperparts  and  under- 
parts  not  strongly  contrasted;  form  robust;  tail  about  three-fourths 
length  of  head  and  body,  black  all  around;  feet  (epidermis  and  hair) 
blackish  to  base  of  claws,  which  are  light  horn  color;  hind  feet 
broad,  stout,  the  digital  bristles  not  projecting  be}Tond  ends  of  claws. 

i  Fifteen  in  Mus.  Comp.  Zool.;  7  in  Field  Mus.  Nat.  Hist. 


1918.]  SUBGENUS   MELANOMYS.  95 

Skull  rotund;  rostrum  short,  nearly  straight;  braincase  large  and 
inflated;  frontals  broad,  the  lateral  margins  projecting  as  supraor- 
bital shelves;  zygomata  slender,  but  maxillary  root  decidedly 
expanded  above  along  frontal  and  premaxillary  sutures;  antorbital 
foramen  rather  narrow  above,  but  little  compressed  and  opening 
widely  below,  the  outer  wall  projecting  slightly  forward  as  viewed 
from  above;  lachrymal  articulating  mainly  with  maxilla;  mastoid 
process  of  squamosal  short  and  broad  owing  to  slight  excision  of 
squamosal  border.  Molars  slightly  hypsodont;  parastyle  and  proto- 
cone  of  first  upper  molar  with  oblique  commissures;  secondary 
parastyle  rudimentary  or  fused  with  main  element;  incisors  thin 
and  weak,  the  upper  ones  descending  about  perpendicularly  from 
premaxillae,  the  points  not  decidedly  curved  backward. 

Remarks. — This  group  of  dark-colored  forms,  segregated  by  Thomas 
under  the  name  Melanomys  as  a  subgenus  of  Oryzomys,  was  elevated 
to  generic  rank  by  Allen  hi  1913.  In  general  characters,  and  espe- 
cially dentition,  however,  it  approaches  typical  Oryzomys  so  closely 
that  subgeneric  recognition  seems  better  to  express  the  relationship. 
The  most  notable  dental  differences  appear  to  be  shown  in  the  rather 
high  molar  crowns,  comparatively  straight,  thin  upper  incisors,  and 
the  tendency  toward  suppression  of  the  secondary  parastyle  normally 
present  hi  typical  Oryzomys.  The  secondary  parastyle  is  normally 
fused  with  the  parastyle,  which  is  correspondingly  enlarged.  Among 
the  more  important  cranial  details  are  the  expansion  of  the  maxillary 
root  of  the  zygoma  over  the  antorbital  foramen  and  the  resulting 
alteration  in  the  position  of  the  lachrymal  as  compared  with  typical 
Oryzomys.  The  lachrymal  hi  Melanomys  articulates  almost  entirely 
with  the  maxilla,  its  position  being  anterior  to  the  fronto-m axillary 
suture,  while  in  typical  Oryzomys  it  bridges  this  suture  and  about 
evenly  overlaps  the  maxilla  and  frontal. 

The  subgenus  Melanomys  differs  so  strikingly  from  the  subgenus 
Oligoryzomys  in  appearance  that  no  close  comparison  is  necessary, 
although  most  of  the  characters  of  the  latter  are  found  hi  varying 
combinations  in  the  subgenus  Oryzomys. 

ORYZOMYS  CALIGINOSUS  (Tomes). 

Hesperomys  caliginosus  Tomes,  Proc.  Zool.  Soc.  London,  1860,  p.  263.    Type  from 

coast  of  Ecuador,  probably  Esmeraldas.1    Type  No.  7.1.1.128,  British  Museum; 

collected  by  Louis  Fraser. 
Akodon  caliginosus  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XIV,  p.  45,  Jan.  31,  1901. 
Hesperomys  (Melanomys)  caliginosus  Thomas,  Ann.  Mag.  Nat.  Hist.  ser.  8,  XT,  p.  406, 

Apr.  1913. 
Melanomys  caliginosus  caliginosus  [et  al.  subsp.]  Allen,  Bull.  Amer.  Mus.  Nat.  Hist., 

XXXII,  p.  537,  Nov.  17,  1913. 

»  See  Allen  (op.  cit.,  pp.  533-554,  1913)  for  locality  and  revision  of  group  regarded  as  generically  distinct. 


96 


NORTH   AMERICAN    FAUNA. 


[No.  43. 


Geographic  distribution. — (See  under  subgenus  Melanomys)  (fig.  11). 
General  characters. — (See   subgeneric   characters  under  subgenus 
Melanomys.) 

Color. — Fresh  pelage:  Upperparts  varying  from  tawny  to  russet, 
abundantly  but  finely  mixed  with  black,  the  black  tending  to  become 
predominant  especially  on  rump;  underparts  overlaid  with  varying 
shades  from  dark  ochraceous-tawny  to  cinnamon  brown,  the  dark 
plumbeous  basal  color  showing  through;  vibrissse  black,  scarcely 
reaching  posteriorly  to  ears;  ears,  feet,  and  tail  thinly  clothed  with 
short  hairs,  the  hairs  and  epidermis  blackish.  Young  (in  first  pelage) : 
Similar  to  adults,  but  upperparts  usually  more  distinctly  blackish, 

and  underparts  less  heavily 
overlaid  with  ochraceous- 
tawny  or  cinnamon  brown. 

Skull. — (See  subgeneric 
characters  under  subgenus 
Melanomys.) 

Remarks. — In  a  revision  of 
the"  Melanomys"  group,  Allen  * 
assigns  the  component  parts  to 
several  specific  types  on  the 
basis  of  characters  which  seem 
comparatively  unimportant; 
the  slight  departures  exhibited 
either  in  color  or  cranial  de- 
tails, in  examples  from  widely 
separated  regions,  point  to  the 
probable  inclusion  of  most,  if  not  all,  of  the  forms  in  the  species 
0.  caliginosus.  Two  subspecies  range  within  the  limits  of  the  region 
under  review. 

Key  to  Subspecies  of  O.  caliginosus  (Exclusive  of  South  America). 
a1.  Upperparts  tending  toward  tawny;   frontal  region  narrower.     (Eastern  Panama.) 

O.  c.  idoneus  (p.  96). 

a2.  Upperparts  tending  toward  russet;    frontal  region  broader.     (Western  Panama; 

Costa  Rica;  Nicaragua.) O.  c.  chrysomelas  (p.  97). 

ORYZOMYS  CALIGINOSUS  IDONEUS  Goldman 

Cerro  Azul  Dusky  Rice  Rat. 

(  PI.  IV,  figs.  7,  7a;  PI.  V,  figs.  3,  12;  PI.  VI,  figs.  9,  9a.) 

Oryzomys  idoneus  Goldman,  Smiths.  Misc.  Coll.,  LVI,  No.  36,  p.  5,  January  19,  1912. 

Melanomys  idoneus  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXXII,  p.  548,  November 

17,  1913. 

Type  locality. — Cerro  Azul,  near  headwaters  of  Chagres  River, 
Panama  (altitude,  2,500  feet). 

Type. — No.  171106,  9  adult,  United  States  National  Museum 
(Biological  Survey  collection);  collected  by  E.  A.  Goldman,  March 
26,  1911. 


1.  O.  c.  idoneus.  \  2.  0.  c.  chrysomelas. 

Fig.  11. — Geographic  distribution  of  subspecies  of 
Oryzomys  caliginosus  (exclusive  of  South  American 
forms). 


'See  Allen  (op.cit.,  pp.  533-554,  1913)  for  locality  and  revision  of  group  regarded  as  generically  distinct. 


1918.]  ORYZOMYS   CALIGINOSUS.  97 

Geographic  distribution. — Heavily  forested  mountain  slopes  in 
eastern  Panama;  known  altitudinal  range  from  1,800  to  2,800  feet; 
Humid  Lower  Tropical  Zone. 

General  characters. — Similar  to  0.  c.  caliginosus,  of  Ecuador,  but 
upperparts  paler,  more  tawny,  instead  of  russet  in  color.  Closely 
resembling  0.  c.  chrysomelas,  but  color  paler;  skull  narrower  between 
orbits. 

Color. — Fresh  pelage:  Upperparts  near  tawny  (becoming  most 
intense  in  worn  pelage),  heavily  and  finely  mixed  with  black,  the 
mixture  usually  uniform,  but  black  predominating  on  rump  in  some 
examples;  underparts,  including  inner  sides  of  limbs,  dark  ochra- 
ceous-tawny;  inner  sides  of  ankles  black;  ears,  feet,  and  tail  thinly 
clothed  with  short  hairs,  the  hairs  and  epidermis  blackish.  Young 
(in  first  pelage) :  Upperparts  usually  more  distinctly  blackish  than 
adults,  the  tawny  element  restricted  to  narrower  tips  of  hairs. 

Skull. — Similar  to  that  of  0.  c.  caliginosus,  but  frontal  region 
narrower;  molars  slightly  larger,  the  toothrows  longer;  anterior 
palatine  foramina  usually  shorter.  Closely  resembling  that  of  0.  c. 
chrysomelas,  but  frontal  region  more  constricted,  the  supraorbital 
borders  less  projecting  laterally. 

Measurements. — Type:  Total  length,  218;  tail  vertebrae,  88; 
hind  foot,  30.  Average  of  10  adults  from  Cana,  Panama:  217.9 
(196-240);  93.3  (85-105);  26.7  (26-27.5).  STcuU  (average  of  same): 
Greatest  length,  30.1  (28.6-30.3);  zygomatic  breadth,  16.2  (15.1- 
16.7);  interorbital  breadth,  6.3  (5.9-6.5);  width  of  braincase,  11.8 
(11.2-12.3);  nasals,  11.4  (10.7-12.6);  anterior  palatine  foramina, 
4.6  (4.4-5);  palatal  bridge,  6.1  (5.4-6.8);  upper  molar  series,  4.7 
(4.4-5). 

Remarks. — While  0.  c.  idoneus  differs  from  typical  0.  c.  caliginosus 
very  appreciably  in  color,  close  relationship  is  evidenced  in  cranial 
characters,  some  skulls  being  scarcely  distinguishable.  In  color  it 
closely  approaches  0.  c.  columbianus,  of  the  Santa  Marta  region  of 
Colombia,  but  is  slightly  darker;  the  tail  is  usually  shorter.  In 
cranial  characters  similarity  to  columbianus  is  also  shown,  but  the 
ascending  branches  of  the  premaxillse  are  narrower  posteriorly,  the 
ends  externally  beveled  in  outline  along  the  interdigitating  fronto- 
premaxillary  fine  of  contact. 

Specimens  examined. — Total  number,  47,  as  follows: 

Panama:  Cana,  46;  Cerro  Azul,  1  (type). 

ORYZOMYS  CALIGINOSUS  CHRYSOMELAS  (Allen). 
Costa  Rican  Dusky  Rice  Rat. 

(PI.  IV,  figs.  8,  8a.) 

Hesperomys  (Habrothrix)  caliginosus  Allen,  Bull.  Amer.  Mue.  Nat.  Hist  ,  III   p   210 

Apr.  17,  1891  (Costa  Rica).  '  *'        ' 

Oryzomys  chrysomelas  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  IX,  p.  37,  Mar.  11,  1897. 
Oryzomys  (Melanomys)  chrysomelas  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXIV.  p  654 

Oct.  13,  1908  (Nicaragua).  e 

Zygodontomys  chrysomelas  Bangs,  Bull.  Mus.  Comp.  Zool.,  XXXIX,  p.  37,  April,  1902 

(Panama). 
Melanomys  chrysomelas  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXXII,  p.  547.  Nov  17 

1913.  "      ' 

Zogodontomys  (sic)  chrysomelas  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  XXXII,  p.  547. 

Nov.  17,  1913.  *         ' 

14521°— 18 7 


98  NORTH   AMERICAN    FAUNA.  [No.  43. 

Type  locality. — Suerre,  Costa  Rica  (altitude  probably  between  3,000 
and  4,000  feet). 

Type. — No.  VkW>  $  adult,  American  Museum  of  Natural  History ; 
collected  by  Anastasio  Alfaro,  July  16,  1895. 

Geographic  distribution. — Western  Panama,  Costa  Rica,  and  north 
to  northern  Nicaragua ;  altitudinal  range  from  near  sea  level  to  about 
3,000  feet,  mainly  in  Humid  Lower  Tropical  Zone. 

General  characters. — Most  closely  allied  to  0.  c.  idoneus,  but  slightly 
darker  in  color;  skull  broader  between  orbits. 

Color. — About  as  in  0.  c.  idoneus,  but  rufescent  element  in  upper- 
parts  darker  and  inclining  toward  the  russet  tone  of  the  paler  species 
of  0.  c.  caliginosus. 

Skull. — Similar  to  that  of  0.  c.  idoneus,  but  frontal  region  less  con- 
stricted, the  supraorbital  borders  more  projecting  laterally. 

Measurements. — Average  of  five  adults  from  Escondido  River  (45 
miles  from  Bluefields),  Nicaragua:  Total  length,  210  (202-217);  tail 
vertebrae,  89  (87-90);  hind  foot,  25.9  (25-27).  SJcull  (average  of 
four  adults  from  same  locality):  Greatest  length,  29.1  (28.2-29.5); 
zygomatic  breadth,  15.9  (15.8-16.1);  interorbital  breadth.  6.6  (6.3- 
6.8);  width  of  braincase,  11.8  (11.4-12.2);  nasals,  10.9  (10.1-11.5); 
anterior  palatine  foramina,  4.9  (4.8-5.1);  palatal  bridge,  5.6  (5.3- 
5.8);  upper  molar  series,  4.5  (4.3-4.7). 

Remarks. — The  range  of  0.  c.  chrysomelas  marks  the  northern  limit 
of  the  species  in  Central  America.  The  Costa  Rican  subspecies  is 
most  closely  allied  to  its  near  geographic  neighbor  in  eastern  Panama, 
although  in  color  the  darker  specimens  approach  some  of  the  paler 
examples  of  0.  c.  caliginosus  and  other  South  American  forms. 

Specimens  examined. — Total  number,  61,  as  follows: 

Costa  Rica:  Pacuare,  5;  Port  Limon,  1;  San  Carlos,  l;1  Suerre,  5  (type  and  topo- 

types);2  Talamanca  (probably  near  Sipurio),  4. 
Nicaragua:  Chontales,  3;1  Escondido  River  (45-50  miles  from  Bluefields),  14; 

Greytown,  1;  Savala,  2;1  Pena  Blanca,  3;1  Rio  Coco,  3;1  Rio  Grande,  l;1 

RioTumaS;1  Tuma,  6:1  Vijagua,  6.1 
Panama:  Mosca,  1. 

Plate  I. 

[All  subgenus  Oryzomys.    Natural  size;  all  in  U.  S.  Nat.  Mus.,  Biological  Survey  collection,  except 

figs.  2,  2a.] 

Figs    1   la.  Oryzomys  palustris  palustris  (Harlan).     Georgetown,  South  Carolina,      c? 
adult.     (No.  71367.) 

2,  2a.  Oryzomys  palustris  coloralus  Bangs.     Type.     Cape  Sable,  Florida.     <? 

adult.     (No.  4470,  Mus.  Comp.  Zool.) 

3,  3a.  Oryzomys  couesi  couesi   (Alston).    Yaruca,  Honduras,     c?  adult.     (No. 

131814.) 

4  4a.  Oryzomys  couesi  regillus  Goldman.     Type.     Los  Reyes,  Michoacan.     <? 

adult.     (No.  125945.) 

5  5a.  Oryzomys  couesi  aquaticus  Allen.     Topotype.     Brownsville,  Texas.     <? 

adult.     (No.  65066.) 

6  6a.  Oryzomys  gatunensis  Goldman.     Type.     Gatun,  Canal  Zone,     c?  young. 

(No.  17i034.)  . 

7  7a.  Oryzomys    cozumelse    Merriam.     Type.     Cozumel    Island,    Mexico,     c? 

adult.     (No.  108462.)  .        . 

8  8a    Oryzomys  peninsulse  Thomas.     San  Jose  del  Cabo,  Lower  California.     ? 

adult.     (No.  146618.)  ^^ 

i  Collection  Amer.  Mus.  Nat.  Hist.  „.,,,,       „T  *  ,-,.  t 

»  Four  (including  type)  in  Amer.  Mus.  Nat.  Hist.;  1  in  Field  Mus.  Nat.  Hist. 


North  American  Fauna  No.  43,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  I. 


Skulls  of  Oryzomys. 


1,  la.  O.  p.  palustris. 

2,  2a.  O.  p.  coloratus. 

3,  3a.  O.  c.  couesi. 
4,4a.  O.c.  regillus. 


5,  5a.  O.  c.  aquatieus. 

6,  6a.  O.  gatunensis. 

7,  7a.  O.  eozumelse. 

8,  8a.  O.  peninsulse. 


North  America 

n  Fauna  No.  43,  U.  S.  Dept.  Agr.     1 

iiological  Survey. 

Plate  II. 

Ir- 

II 

»] 

il 

il 

■  <*o 

g    B  2  -*>  : 

\  B  -c»gr 

;     & 

'                                      -~             £Z 

-iBiMfiHBlii  rllHirl 

^          4? 

Ik; 

■  1 '.'  ^  -  '^''^**<Jy?S?*8iw 

*** 

wJ 
1^ 

HI 

lr,il 

■a    nwl 

/a               ^TT^a 

^  ^ 

<4a 

f  1 

^w       r? 

c?                           o 

W } 

r^ 

i> 

.-~. 

Br  & 

Hifl 

TB     Br        ^B 
Jfl      B>       -j  sB 

B   y 

■Fr  s*v-  s    jraH 

~M 

ra1 

^CJi 

W^    7a 

W*« 

Skulls  of  Oryzomys. 


1,  la.  O.  nelsoni. 

2,  2a.  O.  m.  melanotis. 

3,  3a.  O.  m.  colimensis. 

4,  -ia.  O.  r.  rostratus. 


5,  5a.  O.  r.  megadon. 

6,  6a.  O.  r.  yucatanensis. 

7,  7a.  O.  b.  bombycinus. 
8,8a.  O.  b.  alleni. 


Plate  II. 

[All  subgenus  Oryzomys     Natural  size;  all  in  U.  S.  Nat.  Mus.,  Biological  Survey  collection,  except 

figs.  8,  8a.] 

Figs.  1,  la.  Oryzomys  nelsoni  Merriam.    Type.     Maria  Madre  Island,  Mexico,     cf 
adult.     (No.  89200.) 

2,  2a.  Oryzomys  melanotis  melanotis  Thomas.    Topotype.     San  Sebastian,  Ja- 

lisco,    c?  adult.     (No.  88062.) 

3,  3a.  Oryzomys  melanotis  colimensis,   subsp.  nov.     Type.     Armeria,  Colima. 

$  adult.     (No.  mn~) 

4,  4a.  Oryzomys  rostratus  rostratus  Merriam.    Type.    Metlatloyuca,   Puebla. 

$  adult.     (No.  93112.) 
5, 5a.  Oryzomys    rostratus    megadon    Merriam.    Type.    Teapa,    Tabasco.     <? 
adult.     (No.  99978.) 

6,  6a.  Oryzomys  rostratiis  yucatanensis  Merriam.    Type.     Chichen  Itza,  Yuca- 

tan.    <?  adult.     (No.  108139.) 

7,  la.  Oryzomys   bombycinus   bombycinus   Goldman.    Topotype.    Cerro  Azul, 

Panama.     $   adult.     (No.  171103.) 

8,  8a.  Oryzomys  bombycinus  alleni  Goldman.     Type.     Tuis,  Costa  Rica.     <? 

subadult.     (No.  ffft,  Amer.  Mus.  Nat.  Hist.) 


Plate  III. 
[All  subgenus  Oryzornys.    Natural  size;  all  in  U.  S.  Nat.  Mus.,  Biological  Survey  collection. 

Figs.  1,  la.  Oryzornys  alfaroi  alfaroi  (Allen).    Yaruca,  Honduras.     $   adult.     (No. 
131817.) 

2,  2a.  Oryzornys  alfaroi  dariensis  Goldman.    Type.     Cana,  Panama.     $  adult. 

(No.  178660.) 

3,  3a.  Oryzornys  alfaroi  rhabdops  Merriam.        Type.     Calel,   Guatemala,     <? 

adult.     (No.  76813.) 
4, 4a.  Oryzornys  alfaroi  caudatus  Merriam.    Type.     Comaltepec,  Oaxaca.     d 
adult.     (No.  68641.) 

5,  5a.  Oryzornys  alfaroi  palatinus  Merriam.    Type.    Teapa,  Tabasco.     ?  adult. 

(No.  99977.) 

6,  6a.  Oryzornys  alfaroi  dilutior  Merriam.    Type.    Huauchinango,  Puebla.     S 

adult.     (No.  93124.) 
7, 7a.    Oryzornys   guerrerensis    Goldman.     Type.     Omilteme,    Guerrero,      c? 

adult.     (No.  127517.) 
8,  8a.  Oryzornys  hylocetes  Merriam.     Type.     Chicharras,    Chiapas.     S    adult. 

(No.  77605.) 


North  American  Fauna  No.  43,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  III. 


Skulls  of  Oryzomys. 


1,  la.  O.  a.  alfaroi. 

2,  2a.  O.  a.  dariensis. 

3,  3a.  O.  a.  rhabdops. 
4,4a.  O.  a.  caudalus. 


5,  5a.  O.  a.  palatums. 
6, 6a.  O.  a.  dilution 

7,  7a.  O.  guerrerensis. 

8,  8a.  O.  hylocetes. 


North  American  Fauna  No.  43,  U.  S.  Dept.  Agr.      Biological  Survey. 


Plate  IV. 


Skulls  of  Oryzomys. 


1,  la.  O.  devius. 

2,  2a.  O.  pirrensis. 

3,  3a.  O.  talamancae. 

4,  4a.  O.  t.  frontalis. 


5,  5a.  O.  f.  fulvescens. 

6,6a.  O.f.  lenis. 

7,  7a.  O.  c.  idoneus. 

8,  8a.  O.  c.  chrysomelas. 


Plate  IV. 

[Natural  size;  all  in  U.  S.  Nat  Mus.,  Biological  Survey  collection,  except  figs.  1,  la.J 

Figs.  1,  la.  Oryzomys  (Oryzomys)  devius  Bangs.     Type.     Boquete,  Panama.     9  adult. 
(No.  10324,  Mus.  Comp.  Zool.) 

2,  2a.  Oryzomys  (Oryzomys)  pirrensis  Goldman.     Type.     Mount  Pirre,  Panama. 

c?  adult.     (No.  178993.) 

3,  3a.  Oryzomys  (Oryzomys)  talamancse,  Allen.     Gatun,  Canal  Zone,     c?  adult. 

(No.  170981.) 

4,  4a.  Oryzomys  (Oryzomys)  tectus  frontalis  Goldman.     Type.     Corozal,  Canal 

Zone.     ?  adult.     (No.  171531.) 

5,  5a.  Oryzomys  (Oligoryzomys)  fulvescens  fulvescens  (Saussure).     Orizaba,  Vera 

Cruz,     d1  adult.     (No.  63688.) 

6,  6a.  Oryzomys  (Oligoryzomys)  fulvescens  lenis  Goldman.     Type.     Los  Reyes, 

Michoacan.     (No.  125941.) 

7,  7a.  Oryzomys  (Oligoryzomys)  caliginosus  idoneus  Goldman.     Cana,  Panama. 

c?  adult.     (No.  178608.) 

8,  8a.  Oryzomys  (Oligoryzomys)  caliginosus   chrysomelas  (Allen).     Port   Limon, 

Costa  Rica.     <?  adult.     (No.  76265.) 


PLATfe   V. 

[Natural  size;  all  in  U.  S.  Nat.  Mus.,  Biological  Survey  collection.] 

Fig.  1.  Oryzomys  (Oryzomys)  palustris  palustris  (Harlan).     Georgetown,  South  Caro- 
lina.    <?  adult.     (No.  71367.) 

2.  Oryzomys    (Oligoryzomys)  fulvescens  fulvescens    (Saussure).     Orizaba,    Vera 

Cruz.     <?  adult.     (No.  58248.) 

3.  Oryzomys  (Melanomys)  caliginosus  idoneus  Goldman.     Cana,   Panama.     <? 

adult.     (No.  178663.) 

4.  Oryzomys  (Oryzomys)  palustris  palustris  (Harlan).     Greenwich,  New  Jersey. 

<?  adult.     (No.  117384.) 

5.  Oryzomys  (Oryzomys)  melanotis  melanotis  Thomas.     San  Sebastian,  Jalisco. 

<?  adult.  (No.  88061.) 

6.  Oryzomys  (Oryzomys)  alfaroi  dariensis  Goldman.     Cana,  Panama,     d  adult. 

(No.  178657.) 

7.  Oryzomys  (Oryzomys)  talamancx  Allen.     Gatun,  Canal  Zone.      9  adult.     (No. 

170979.) 

8.  Oryzomys  (Oryzomys)  bombycinus  bombycinus  Goldman.     Cerro  Azul,  Panama 

9  adult.     (No.  171103.) 

9.  Oryzomys  (Oryzomys)  pirrensis  Goldman.     Mount  Pirre,  Panama.     <$  adult. 

(No.  178996.) 

10.  Oryzomys  (Oryzomys)  tectus  frontalis  Goldman.     Cana,   Panama.     <$   adult. 

(No.  178649.) 

11.  Oryzomys    (Oligoryzomys)  fulvescens  fulvescens    (Saussure).     Orizaba,    Vera 

Cruz.     <?  adult.     (No.  58248.) 

12.  Oryzomys   (Melanomys)   caliginosus  idoneus  Goldman.     Cana,    Panama.     <? 

adult.     (No.  178684.) 


North  American  Fauna  No.  43,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  V. 


VgBjJJ 


s^M 


Skulls  and  Mandibles  of  Oryzomys. 


1,4.  O.  p.  palustris. 

2. 11.  O.  f.  fulvescens. 

3. 12.  O.  c.  idoneus. 

5.  O.  m.  melanotis. 

6.  O.  a.  dariensis. 


7.  O.  talamancse. 

8.  O.  b.  bombycinus. 

9.  O.  pirrensis. 
10.  O.  t.  frontalis. 


North  American  Fauna  No.  43,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  VI. 


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1,  la.  O.  p.  palustris. 

2,  2a.  O.  m.  melanotis. 

3,  3a.  O.  a.  dariensis. 


Molars  of  Oryzomys. 

4,  4a.  O.  talamancse. 

5,  5a.  O.  b.  bombycinus. 

6,  6a.  O.  pirrensis. 


7,  7a.  O.  t.  frontalis. 

8,  Sa.  O.  f.  fulvescens. 

9,  9a.  O.  c.  idoneus. 


Plate  VI. 

Slightly  retouched  and  enlarged  about   eight  diameters;  all  in  U.  S.  Nat.  Mus.,  Biological  Survey 

collection.] 

Figs.  1,  la.  Oryzomys  (Oryzomys)  palustris  palustris  (Harlan).     Dismal  Swamp,  Vir- 
ginia.    <?  subadnlt.     (No.  75203.) 

2,  2a.  Oryzomys  (Oryzomys)  melanotis  melanotis  Thomas.     Ixtapa,  Jalisco,     cf 

subadult.     (No.  88068.) 

3,  3a.  Oryzomys  (Oryzomys)  alfaroi  dariensis  Goldman.     Cana,  Panama.     <?  sub- 

adult.     (No.  178653.) 

4,  4a.  Oryzomys  (Oryzomys)  talamancn    Allen.     Cana,   Panama.      9    subadult. 

(No.  179601.) 

5,  5a.  Oryzomys   (Oryzomys)    bombycinus   bovibyeinus   Goldman.     Cerro   Azul, 

Panama.      9  adult.     (No.  171103.) 
6,6a.  Oryzomys  (Oryzomys)  pirrensis  Goldman.     Mount  Pine,  Panama.      9  sub- 
adult.    (No.  179030.) 

7,  7a.  Oryzomys  (Oryzomys)  tectus  frontalis  Goldman.     Cana,  Panama.      9  sub- 

adult,    (No.  178646.) 

8,  8a.  Oryzomys  (Oligoryzomys)  fulvescens  fuhescens  (Saussure).     Orizaba,  Vera 

Cruz,     c?  subadult.     (No.  58244.) 

9,  9a.  Oryzomys   (Melanomys)   caliginosus  idoneus   Goldman.     Cana,    Panama. 

9  subadult.     (No.  178668.) 


INDEX. 


[New  names  in  bold-faced  type;  synonyms  in  italics;  principal  references  in  bold-faced  figures.] 


Age,  7. 

Akodon  caliginosus,  95. 

albi venter,  Oryzomys  couesi,30,34,35,36,37,38,40. 

alfaroi,  Hesperomys  (Oryzomys),  59. 

Oryzomys  (species),  58-59,  74. 

Oryzomys  alfaroi,  8, 59-61,  61, 62, 63, 66, 67, 68. 
alleni,  Oryzomys  bombycinus,  77,  78. 
angusticeps,  Oryzomys  alfaroi,  60, 63-63, 64 ,  65, 71 . 
antillarum,  Oryzomys,  44-45, 87. 
apatelius,  Oryzomys  jalapx,  30. 
aquaticus,  Oryzomys  couesi,  36,  39,  39-40. 
Arvicola,  8. 

oryzivora,  9,  22. 

palustris,  9. 
aztecus,  Oryzomys  couesi,  34,  35-36,  36,  37. 


bombycinus,  Oryzomys  (species),  76-77. 

Oryzomys  bombycinus,  77,  78. 
bulleri,  Oryzomys,  33. 


caliginosus,  Akodon,  95. 

Hesperomys,  95;  97. 

Melanomys  caliginosus,  95. 

Oryzomys  (species),  95-96. 

Oryzomys  caliginosus,  97,  98. 
carrikeri,  Oryzomys,  73,  74. 
caudatus,  Oryzomys  alfaroi,  64-65,  68,  70. 
chapmani,  Oryzomys  alfaroi,  59, 60, 65, 66, 67, 67-68, 

69,  70. 
Characters,  general,  5-7. 
chrysomelas,  Melanomys,  97. 

Oryzomys  caliginosus,  97,  97-98. 

Zogodontomys,  97. 

Zygodontomys,  97. 
colimensis,  Oryzomys  melanotis,  50,  51,  51-52. 
Color,  6,  10. 

coloratus,  Oryzomys  palustris,  23,  25,  26,  27. 
columbianus,  Oryzomys  caliginosus,  97. 
costaricensis,  Oryzomys  fulvescens,  90,  92-93,  94. 
couesi,  Hesperomys,  9,  29. 

Oryzomys  (species),  28-29, 30, 36,  42, 44, 47, 48. 

Oryzomys  couesi,  8, 19,  20,  29-31,  32,33, 34,  35, 
37,  38,  39,  40,  43,  44. 
cozumelse,  Oryzomys,  43. 
crinitus,  Oryzomys  couesi,  30, 35, 36-37, 38, 40, 42. 

D. 

dariensis,  Oryzomys  alfaroi,  60,  61-62. 
devius,  Oryzomvs,  80,  80-81,  81,  82. 
dilutior,  Oryzomys  alfaroi,  67,  68,  68-69. 


E. 


Economic  relations,  4-5. 
Explanations,  10-11. 

F. 

flavicans,  Oryzomys  flavicans,  83,  84. 
floridanus,  Oryzomys  natator,  26. 
frontalis,  Oryzomys  tectus,  85,  85-86. 
fulgens,  Oryzomys,  36,  41-42. 
fulvescens,  Hesperomys,  9,  89. 

Oryzomys  (species),  88-89. 

Oryzomys  fulvescens,  89-91,  91,  92,  93,  94. 


G. 

gatunensis,  Oryzomys,  42-43. 
Geographic  variation,  8. 
goldmani,  Oryzomys,  30. 
gracilis,  Oryzomys,  59,  60,  61,  62. 
Groups,  14-15. 

Oryzomys  alfaroi,  15,  56-71,  76. 

bombycinus,  6,  15,  73,  75-78. 

caliginosus,  95-98. 

devius,  15,  78-82. 

fulvescens,  88-94. 

longicaudatus,  86. 

melanotis,  15,  47-55,  76. 

meridensis,  80. 

palustris,  6,14-15,  18-46,  57,  71,  72,  75,  76, 
79v80. 

pirrensis,  6. 

talamancaj,  6,  15,  56,  57,  71-74,  75,  76,  79, 
83 

tectus,  15,  82-86. 
guerrerensis,  Oryzomys,  58,  59,  69-70. 


Habits.  4-5. 

Habrothrix  caliginosus,  97. 

Hesperomys,  9, 11. 

alfaroi,  59. 

caliginosus,  95,  97. 

couesi,  9,  29. 

fulvescens,  9,  89. 

palustris,  22. 
History,  8-10. 
hylocetes,  Oryzomys,  59,  64,  70-71. 

I. 

idoneus,  Melanomys,  96. 

Oryzomys  caliginosus,  96-97,  98. 
illectus,  Oryzomys  flavicans,  84. 
incertus,  Oryzomys  alfaroi,  59,  60. 
Individual  variation,  7. 
Introduction,  3-11. 


jalapx,  Oryzomys,  29. 


lenis,  Oryzomys  fulvescens,  90,  91. 
Life  zones,  11. 

List  of  species  and  subspecies,  16-17. 
longicaudatus,  Oryzomys,  87. 

M. 

maculiventer,  Oryzomys,  80,  82. 

Material,  8-10. 

mayensis,  Oryzomys  fulvescens,  92. 

Measurements,  10. 

medius,  Oryzomys,  73. 

megadon,  Oryzomys  rostratus,  53,  54,  54-55,  55. 

Megadontomys,  80. 

Melanomys,  genus,  11. 

subgenus,  4,  5,  11,  12,  13, 17,  94-98. 
Melanomys  caliginosus,  95. 

chrysomelas,  97. 

idoneus,  96. 
melanotis,  Oryzomys  (species),  49-50,  52,  53, 87. 

Oryzomys  melanotis,  50-51,  51,  52. 
meridensis,  Oryzomys,  80,  82. 

99 


100 


NORTH   AMERICAN    FAUNA. 


[No.  43,  1918.] 


mexicanus,  Oryzomys  couesi,  8,  30,  31,  33,  33-35, 

35,36,37,38,45,46. 
molestus,  Oryzomys,  38. 
mollipilosus,  Oryzomys,  73,  74. 
Molt,  6-7. 
Mus  palustris,  8,  9,  11,  22. 

N. 

natator,  Oryzomys  palustris,  23,  24,  24-25,  26,  27. 

navus,  Oryzomys,  11. 

Neacomys,  13. 

Nectomys,  13. 

nelsoni,  Oryzomys,  46-47. 

Nesoryzomys,  13. 

nicaragux,  Oryzomys  (Oligoryzomys),  92,  93. 

Nyctomys,  14. 

O. 
03eomys,  83. 

Oligoryzomys,  subgenus,  4,  5,  11,  14,  17,  87-94,  95. 
oryzivora,  Arvtcola,  9,  22. 
Oryzomys,  genus,  3,  11-17.  / 

subgenus,  4,  11,  16,  17-87,  87,  88,  95. 
Oryzomys  albiventer,  30, 34, 35, 36, 37,  38, 40. 

alfaroi,  8,  59-61,  61,  62,  63,  66,  67,  68. 

alleni,  77,  78. 

angusticeps,  60,  62-63,  64,  65,  71. 

antillaruni,  44-45,  87. 

apatelius,  30. 

aquaticus,  36,  39,  39-40. 

aztecus,  34,  35-36,  36,  37. 

bombycinus,  76-77,  77,  78. 

bulleri,  33. 

caliginosus,  96,  97,  98. 

carrikeri,  73,  74. 

caudatus,  64-65,  68,  70. 

chapmani,  59,  60,  65,  66,  67,  67-68,  69,  70. 

chrysomelas,  97,  97-98. 

colimensis,  50,  51,  51-52. 

coloratus,  23,  25,  26,  27. 

columbianus,  97. 

costaricensis,  90,  92-93,  94. 

couesi,  8,  19,  20,  28-29,  29-31,  32,  33,  34,  35,  36, 
37,  38,  39,  40,  42,  43,  44,  47,  48. 

cozumela;,  43. 

crinitus,  30,  35,  36-37,  38,  40,  42. 

dariensis,  60,  61-62. 

devius,  80.  80-81,  81,  82. 

dilutior,  67,  68,  68-69.      • 

flavicans,  83,  84. 

floridanus,  26. 

frontalis,  85,  85-86. 

fulgens,  36,  41-42. 

fulvescens,  88-89,  89-91,  91,  92,  93,  94. 

gatunensis,  42-43. 

goldmani,  30. 

gracilis,  59,  60,  61,  62. 

guerrerensis,  58,  59,  69-70. 

hylocetes,  59,  64,  70-71. 

idoneus,  96-97,  98. 

illectus,  84. 

incertus,  59,  60. 

jalapx,  29. 

lenis,  90,  91. 

longicaudatus,  87. 

maculiventer,  80,  82. 

mayensls,  92. 

medius,  73. 

megadon,  53,  54,  54-55,  55. 

melanotis,  49-50,  50-51,  51,  52,  53,  87. 

meridensis,  80,  82. 

mexicanus,  8,  30,  31,  33,  33-35,  35,  36,  37,  38,  45, 
46. 

molestus,  38. 

mollipilosus,  73,  74. 

natator,  23,  24,  24-25,  26,  27. 

navus,  11. 

nelsoni,  46-47. 

nicaragux,  92,  93. 

palatums,  65-66,  66,  67. 

palmarius,  83,  84. 

palmin?,  59,  61,  62. 

palustris,  3,  4,  5,  21-22,  22-24,  25,  26,  27,  29,  48, 
62,  74. 


Oryzomys  panamensis,  73,  74. 
penmsulae,  29,  45-46. 
peragrus,  30,  31,  39,  40. 
pirrensis,  80,  81,  81-82. 
regillus,  34,  36,  37,  37,  38,  40. 
rhabdops,  59,  60,  62,  63,  63-64,  65,  71. 
richardsoni,  30. 
richmondi,  30,  32,  42. 
rostratus,  49,  50,  52,  53-54,  54,  55. 
rufinus,  29. 
rufus,  33. 

saturation  65,  66,  66-67,  71. 
talamancae,  53,  60,  61,  62,  73-74. 
teapensis,  29. 
tectus,  84,  84-85,  85,  86. 
texensis,  23,  25,  26,  27-28. 
vegetus,  93,  93-94. 
victus,  14,  86-87. 
yucatanensis,  52,  53,  54,  55,  55. 
zygomaticus,  30,  31,  32-33,  34. 

P. 

palatums,  Oryzomys  alfaroi,  65-66,  66,  67. 
palmirag,  Oryzomys,  59,  61,  62. 
palmarius,  Oryzomys,  83,  84. 
palustris,  Arvicola,  9. 

Hespcromys,  22. 

Mus,  8,  9,  11,  22. 

Oryzomys  (species),  21-22,  29,  48,  62,  74. 

Oryzomys  palustris,  3,  4,  5,  22-24,  25,  26,  27. 
panamensis,  Oryzomys,  73,  74. 
Pelage,  6. 

peninsulae,  Oryzomys,  29,  45-46. 
peragrus,  Oryzomys  couesi,  30,  31,  39,  40. 
geromyscus,  14,  80. 
phseopus,  Oryzomys,  11,  94. 
pirrensis,  Oryzomys,  SO,  81,  81-82 

R. 

regillus,  Oryzomys  couesi,  34,  36,  37,  37,  38,  40. 

Reithrodontomys,  14,  87,  88. 

rhabdops,  Oryzomys  alfaroi,  59,  60,  62,  63,  63-64, 

65,  71. 
Rhipidomys,  13,  83. 
richardsoni,  Oryzomys,  30. 
richmondi,  Oryzomys  couesi,  30,  32,  42. 
rostratus,  Oryzomys  (species),  49,  50,  52. 

Oryzomys  rostratus,  53-54,  54,  55. 
rufinus,  Oryzomys  jalapx,  29. 
rufus,  Oryzomys,  33. 

S. 

saturatior,  Oryzomys  alfaroi,  65,  66,  66-67,  71. 
Specimens  examined,  9-10. 
Subgenera,  14-15,  17. 

T. 

talamanca?,  Oryzomys,  53,  60,  61,  62,  73-74. 
teapensis,  Oryzomys,  29. 
tectus,  Oryzomys  (species),  84. 

Oryzomys  tectus,  84-85,  85,  86. 
Teeth,  10,  11  (fig.). 

texensis,  Oryzomys  palustris,  23,  25,  26,  27-28. 
Tylomys,  84. 

V. 
Variation,  7-8. 

vegetus,  Oryzomys  fulvescens,  93,  93-94. 
victus,  Oryzomys,  14,  86-87. 


yucatanensis,  Oryzomys  rostratus,  52,  53,  54,  55,  55. 
Z. 

Zogodontomys  chrysomelas,  97. 
Zonal  divisions,  11. 
Zygodontomys,  14. 
chrysomelas,  97. 
zygomaticus,  Oryzomys  couesi,  30,  31,  32-33,  34. 


ADDITIONAL  COPIES 

OF  THIS  PUBLICATION  MAY  BE  PROCURED  FROM 

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V 


North  American  Fauna  No.  44,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate 


Small  Eastern  Flying  Squirrel  (Glaucomys  volans  volans). 
From  life;  Coram,  N.  Y.,  August,  1915;  photographed  by  Francis  Harper. 


U.  S.  DEPARTMENT  OF  AGRICULTURE 

BUREAU  OF  BIOLOGICAL  SURVEY 

E.  W.  NELSON,  Chief 


NORTH  AMERICAN  FAUNA 

No.   44- 

[Actual  date  of  publication,  June  13,  1918] 


REVISION  OF  THE  AMERICAN  FLYING  SQUIRRELS 


ARTHUR    H.    HOWELL 

ASSISTANT   BIOLOGIST,    BIOLOGICAL    SURVEY 


WASHINGTON 

GOVERNMENT  PRINTING  OFFICE 

1918 


J,  S.  SUPT.  OF  DOCUMENT^ 


LETTER  OF  TRANSMITTAL. 


United  States  Department  or  Agriculture, 

Bureau  of  Biological  Survey, 
Washington,  D.  C,  March  9, 1917. 
Sir  :  I  have  the  honor  to  transmit  for  publication  as  North  Ameri- 
can Fauna  No.  44  a  revision  of  the  American  flying  squirrels  (genus 
Glaucomys) ,  by  Arthur  H.  Howell,  assistant  biologist  of  this  bureau. 
The  revision  is  based  largely  upon  material  in  the  collection  of  the 
Biological  Survey.  Flying  squirrels  are  found  over  much  of  North 
America,  and  this  report  on  the  group  gives  needed  information  con- 
cerning the  number  of  forms  and  their  distribution. 

Respectfully, 

E.  W.  Nelson, 
Chief  Biological  Survey. 
Hon.  David  F.  Houston, 

Secretary  of  Agriculture. 


CONTENTS. 


Page. 

Introduction 5 

Habits 5 

Voice 6 

Nests 7 

Breeding 8 

Food 9 

Economic  status 10 

Pelage  and  molt 10 

Explanation  of  cranial  measurements 11 

Material  examined 11 

Genus  Glaucomys 11 

History  and  nomenclature 12 

Generic  names 12 

Specific  names 13 

Generic  characters 14 

List  of  species  and  subspecies,  with  type  localities 16 

Key  to  species  and  subspecies 16 

Descriptions  of  species  and  subspecies 18 

Glaucomys  volans  group 18 

Glaucomys  sabrinus  group  j 29 

Tables  of  cranial  measurements 59 

Explanation  of  plates 62 

Index 63 

3 


ILLUSTRATIONS. 


[Plate  I,  frontispiece ;  Plates  II-VII,  following  page  62.] 
Plate  I.  Small  eastern  flying  squirrel  (Glaucomys  volans  volans). 

II.  Skulls  (dorsal  view)  of  Glaucomys  volans  saturatus,  G.  v.  volans,  G.  v.  texensis, 
G.  v.  goldmani,  G.  v.  querceti,  G.  sabrinus  bangsi,  G.  s.  alpinus,  G.  s. 
canescens,  G.  s.  yukonensis,  G.  s.  macrotis,  G.  s.  sabrinus,  and  G.  s.  mak- 
kovikensis. 

III.  Skulls  (dorsal  view)  of  Glaucomys  sabrinus  stephensi,  G.  s.  flaviventris,  G.  s. 

klamatliensis,  G.  s.  fuliginosus,  G.  s.  lascivus,  G.  s.  zaphseus,  G.  s.  colum- 
biensis,  G.  s.  latipes,  G.  s.  californicus,  G.  s.  olympicus,  G.  s.  oregonensis, 
and  G.  s.  bullatus. 

IV.  Skulls  (ventral  view)  of  Glaucomys  volans  saturatus,  G.  v.  volans,  G.  v. 

texensis,  G.  v.  goldmani,  G.  v.  querceti,  G.  sabrinus  bangsi,  G.  s.  alpinus, 
G.  s.  canescens,  G.  s.  yukonensis,  G.  s.  macrotis,  G.  s.  sabrinus,  and  G.  s. 
makkovikensis . 
V.  Skulls  (ventral  view)  of  Glaucomys  sabrinus  stephensi,  G.  s.  flaviventris,  G.  s. 
klamathensis,  G.  s.  fuliginosus,  G.  s.  lascivus,  G.  s.  zaphseus,  G.  s.  colum- 
biensis,  G.  s.  latipes,  G.  s.  californicus,  G.  s.  olympicus,  G.  s.  oregonensis, 
and  G.  s.  bullatus. 
VI .  Skulls  (lateral  view)  of  Glaucomys  volans  volans,  G.  v.  querceti,  G.  v.  saturatus, 
G.  v.  texensis,  G.  sabrinus  vmcrotis,  G.  s.  sabrinus,  G.  s.  columbiensis,  G.  s. 
oregonensis,  G.  s.  zaphteus,  G.  s.  bangsi,  G.  s.  canescens,  and  G.  s.  alpinus. 
VII.  Skulls  (lateral  view)  of  Glaucomys  sabrinus  lascivus,  G.  s.  flaviventris,  G.  s. 
klamathensis,  G.   s.   bullatus,   G.  s.  californicus,   G.  s.   olympicus,   G.  «. 
fuliginosus,  G.  s.  latipes,  and  G.  s.  stephensi;  mandible  of  G.  s.  sabrinus; 
and  skull  (mandible  and  dorsal,  ventral,  and  lateral  views)  of  Pteromys 
biichneri. 

TEXT    FIGURES. 

Page. 
Fig.  1.  Map  showing  distribution  of  Glaucomys  sabrinus  and  G.  volans 6 

2.  Map  showing  distribution  of  the  subspecies  of  Glaucomys  volans 19 

3.  Map  showing  distribution  of  the  subspecies  of  Glaucomys  sabrinus  except 

G.  s.  bullatus 30 

4.  Map  showing  distribution  of  Glaucomys  sabrinus  bullatus 51 

4 


No.  44.  NORTH  AMERICAN  FAUNA.        June  13, 1918. 

REVISION  OF  THE  AMERICAN  FLYING  SQUIRRELS. 


By  Arthur  H.  Howell. 


INTRODUCTION. 

The  American  flying  squirrels  comprise  two  distinct  groups — the 
small  species  (Glaucomys  volcms)  of  eastern  United  States  and  Mex- 
ico; and  the  larger  species  {G.  sdbrinus)  occupying  western  United 
States  and  Canada,  with  one  form  ranging  into  New  England  and 
the  border  States  along  the  Great  Lakes  (fig.  1). 

HABITS. 

Flying  squirrels  are  almost  exclusively  arboreal,  seldom  descending 
to  the  ground  and  apparently  never  running  for  any  distance  on  its 
surface.  They  are  unsuspicious  and,  being  easily  tamed,  make  very 
attractive  pets.1  Possessed  of  intense  activity,  their  movements  are 
the  acme  of  grace  and  agility.  They  do  not  fly,  in  the  usual  sense 
of  the  term,  but  progress  from  tree  to  tree  by  gliding  with  out- 
stretched membranes  from  an  elevated  position  to  a  point  lower 
down,  usually  near  the  ground.  Just  before  alighting,  the  animal 
checks  its  momentum  by  sweeping  upward  in  a  gentle  curve  and 
alights  on  the  tree  trunk  with  its  head  up.  Ascending  the  tree  by 
climbing,  it  is  ready  for  another  flight.  In  these  gliding  leaps, 
which  may  extend  for  a  distance  of  50  yards  or  more,  the  squirrel  is 
able  to  change  its  course  to  one  side  or  the  other  with  perfect  ease. 

Unlike  all  other  American  squirrels,  the  flying  squirrels  are  strictly 
nocturnal.  During  the  daytime  they  remain  concealed  in  their 
nests  and  are  never  seen  abroad  unless  frightened  from  their  re- 
treats. Usually  they  may  be  driven  readily  from  their  holes  by 
pounding  with  an  ax  or  club  on  the  base  of  the  tree  in  which  they  are 

1  A  pair  of  flying  squirrels  regularly  visit  the  sixth  floor  of  an  apartment  house  front- 
ing the  National  Zoological  Park,  Washington,  D.  C,  and  although  usually  averse  to  the 
glare  of  an  electric  light,  they  will  when  hungry  eat  nuts  in  apparent  unconcern  within 
5  or  6  feet  of  an  observer.  They  run  up  and  down  the  stuccoed  sides  of  the  building 
with  perfect  ease,  doubtless  reaching  the  middle  stories  by  jumping  from  a  near-by  tree. 
Their  visits  usually  are  between  dark  and  midnight. 

5 


6 


NORTH   AMERICAN   FAUNA. 


[No.   44. 


sleeping.  Woodchoppers  frequently  scare  them  out  when  felling 
timber,  and  anyone  who  camps  frequently  in  the  woods  is  likely  to 
hear  them  at  night  running  or  jumping  about  in  the  trees  or  drop- 
ping nuts  to  the  ground. 


IGlAUCOMYS     SABRINUS 

IGlaucomys   VOLANS 


Fio.  1. — Map  showing  the  distribution  of  Glaucomys  saorinus  and  O.  volana,  based  on 
specimens  examined  and  reports  of  occurrence. 

VOICE. 

The  notes  of  Glaucomys  volans  are  described  by  Vernon  Bailey  as 
a  "  fine,  whistling  squeak,"  and  Wood  says  of  the  same  species :  "  The 


1918.]  INTRODUCTION.  7 

flying  squirrel  has  three  quite  diverse  calls  or  cries.  It  utters  the 
usual '  chuck-chuck '  of  squirrels,  the  usual  quick,  sharp  squeak  when 
scolding,  and  also,  more  rarely,  a  clear  musical  note,  commonly  me- 
lodious and  pleasant,  but  occasionally  shrill.  This  resembles  the 
chirp  of  a  bird  and  may  be  kept  up  for  ten  minutes  at  a  time."1 
Seton,  writing  of  the  northern  flying  squirrel,  G.  sabrinus,  in  Mani- 
toba, says :  "  The  cry  of  this  species  is  said  to  be  like  that  of  volans, 
which  is  a  prolonged  squeak  not  unlike  the  complaint  of  a  red-eyed 
vireo  whose  nest  is  threatened." 2 

NESTS. 

These  squirrels  are  found  only  in  wooded  regions,  and  for  homes 
they  depend  chiefly  on  hollow  trees  or  stubs  and  deserted  woodpecker 
nests.  They  often  take  up  their  abode,  also,  in  attics,  in  outbuild- 
ings, or  in  boxes  constructed  for  martins  or  other  birds.  Audubon 
and  Bachman  mention  finding  a  number  of  flying  squirrels  in  crevices 
of  rock  at  Red  Sulphur  Springs,  Va.  In  the  cavity  chosen  for  a 
home  a  nest  is  constructed  of  shreds  of  bark,  dry  leaves,  moss,  feath- 
ers, fur,  or  other  soft  material.  Outside  nests  are  often  built,  or  the 
deserted  nests  of  birds  or  of  other  squirrels  utilized.  Concerning 
this  habit  (in  Glaucomys  volans)  Mearns  says: 

Not  infrequently  it  builds  outside  nests,  and  even  lives  in  them  during  the 
winter.  Some  resemble  the  leaf  nests  of  the  gray  squirrel  externally,  though 
there  is  always  a  warm  lining  within;  other  nests  are  indistinguishable  from 
those  of  red  squirrels;  and  others  still  are  deftly  woven  of  the  softest  possible 
materials.3 

Helme,4  writing  from  Long  Island,  N.  Y.,  and  Rhoads,5  writing 
from  Pennsylvania,  both  mention  this  habit  in  volans,  and  both  state 
that  an  evergreen  tree  usually  is  selected  as  a  site  for  the  nest.  Ever- 
mann  and  Clark G  speak  of  finding  in  Indiana  a  large  globular  nest 
of  fibrous  material  situated  in  the  crotch  of  a  small  oak ;  and  King 7 
describes  a  summer  nest  built  of  small  twigs  and  oak  leaves,  and 
lined  with  grass,  situated  about  10  feet  from  the  ground  in  a  small 
oak. 

The  larger  squirrels  of  the  sabrlnus  group  also  occasionally  con- 
struct outside  nests,  although,  as  is  the  case  with  volans,  hollow  trees 
usually  are  preferred.  J.  Ellis  McLellan,  at  Gold  Beach,  Oreg.,  once 
found  a  flying  squirrel  occupying  a  small  spherical  nest  made  of 
sticks  and  moss  placed  in  a  small  fir  tree.  Vernon  Bailey,  at  Mc- 
Kenzie  Bridge,  Oreg.,  found  soft  nests  of  the  flying  squirrel  in  the 

iWood,  F.  E.,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  VIII,  pp.  535-536,  1910. 

2  Seton,  E.  T.,  Life  Hist.  Northern  Anim.,  I,  p.  441,  1909. 

3  Mearns,  E.  A.,  Bull.  Amer.  Mus.  Nat.  Hist.,  X,  p.  342,  1898. 

*  Helme,  A.  H.,  Abstr.  Proc.  Linn.  Soc.  New  York,  Nos.  13-14,  p.  23,  1902. 
5  Rhoads,  S.  N.,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  p.  389,  1894. 
8  Proc.  Wasbington  Acad.  Sci.,  XIII,  p.  15,  1911. 
7  King,  F.  H.,  Amer.  Nat.,  XVII,  p.  36,  1883. 


8  NORTH   AMERICAN   FAUNA.  [No.  44. 

masses  of  tree  moss  (Ilypnum)  which  cover  the  trunks  and  branches 
of  maple  and  ash  trees  in  the  deepest  forest.  There  were  runways  up 
the  trunks  under  the  mossy  fleece  and  others  where  the  moss  had  been 
worn  away  by  constant  use.  At  Elk  River,  Minn.,  in  November, 
1904,  Charles  Bailey  found  two  of  the  large  flying  squirrels  (G. 
sabrlnus  rnaerotis)  occupying  a  nest  of  sticks  and  moss  in  a  small 
tamarack.  At  Florence,  Mont.,  on  June  11,  1910,  Clarence  Birdseye 
found  in  a  pile  of  cordwood  a  flying  squirrel's  nest  containing  4 
young  with  eyes  not  yet  open. 

Flying  squirrels  apparently  do  not  hibernate,  but  are  known  to  lay 
up  stores  of  food  and  to  remain  for  a  large  part  of  the  winter  within 
their  nests.  During  the  wTinter'and  to  some  extent,  also,  at  other  sea- 
sons they  are  somewhat  gregarious  in  habit.  At  Mullan,  Idaho,  in 
June,  Clark  P.  Streator  started  8  adults  (G.  s.  latipes)  from  a  hollow 
tree  and  secured  6  of  them,  all  of  which  proved  to  be  males.  Seton 
mentions  finding  9  adults  (G.  s.  canescens)  in  a  hollow  stub  at  Car- 
berry,  Manitoba.1  The  same  habit  is  possessed  by  G.  volans;  Prof. 
U.  O.  Cox,  of  Farmland,  Ind.,  found,  in  November,  15  of  these  squir- 
rels in  a  small  rotten  stub  a  little  higher  than  a  man's  head,2  and 
at  Dothan,  Ala.,  in  March,  the  writer  started  6  from  a  similar  stub 
in  a  wooded  swamp.  Audubon  and  Bachman  record  finding  20  in  a 
martin  box  having  8  or  10  apartments,  placed  on  top  of  a  large  locust 
tree,  the  box  being  occupied,  also,  by  bats  and  screech  owls.3  Dr. 
J.  Schneck  states  that  he  once  ran  as  many  as  50  out  of  one  den.4 
Audubon  and  Bachman  narrate  an  interesting  experience  with  flying- 
squirrels  near  Philadelphia,  Pa.,  where  not  less  than  200  were  ob- 
served at  once  about  sunset  engaged  in  sportive  gambols  in  a  grove 
of  trees.5 

BREEDING. 

The  number  of  young  produced  at  a  birth  varies  from  3  to  6 — 
usually  4  or  5.  The  period  of  gestation  is  said  to  be  one  month,6 
but  no  definite  evidence  on  this  point  is  available.  In  the  Northern 
States  the  young  of  GTaueomys  volans  are  brought  forth  in  April, 
and  a  second  brood  may  be  produced  in  August  or  September.  Ever- 
mann  and  Clark 7  describe  finding  small  naked  young  ones  in  Indiana 
on  August  19,  and  Langdon 8  records  young  with  eyes  not  yet  open 
at  Madisonville,  Ohio,  September  7. 

1  Seton,  E.  T.,  Life  Hist.  Northern  Anim.,  I,  p.  441,  1909. 

2  Evermann  and  Clark,  Proc.  Washington  Acad.  Sci.,  XIII,  p.  16,  1911. 
a  Audubon  and  Bachman,  Quadr.  North  Auier.,  I,  p.  220,  1840. 

4  Wood,  F.  E.,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  VIII,  p.  534,  1910. 

5  Audubon  and  Bachman,  Quadr.  North  Amer.,  I,  p.  218,  1840. 

6  D.  W.  C,  On  the  Habits  of  the  American  Flying  Squirrels  (Pteromys  rolucella  Cuv.)  . 
Mag.  Nat.   Hist.,  IX,  pp.   569-572,   1836. 

7  Evermann  and  Clark,  Proc.  Washington  Acad.  Sci.,  XIII,  p.  15,  1911. 

8  Langdon,  Frank  W.,  Journ.  Cincinnati  Soc.  Nat.  Hist.,  Ill,  p.  303,  1881. 


191S.]  INTRODUCTION.  9 

The  large  flying  squirrels  of  the  sabrinus  group  apparently  breed 
somewhat  later  than  voluns.  Seton  records  a  specimen  taken  at 
Winnipeg,  Manitoba,  on  April  21,  which  contained  6  young  ready 
for  birth ;  and  a  nest  of  1  young,  with  eyes  not  opened,  found  May  15  \ 
Merriam  found  nursing  young  of  G.  s.  macrotis  about  a  month  old 
in  northern  New  York  on  June  18.  In  the  Bitterroot  Mountains, 
Mont.,  at  4,500  feet  altitude,  Clarence  Birdseye  took  a  female  of 
G.  s.  barigs?iL,  May  17,  1910,  containing  two  large  embryos;  and  in 
the  Bitterroot  Valley,  near  Florence,  Mont.,  found  a  nest  of  small 
nursing  young  on  June  11. 

FOOD. 

The  food  of  flying  squirrels  consists  in  large  part  of  nuts,  including, 
probably,  most  of  the  native  species — chestnuts,  acorns,  beechnuts,  hick- 
ory nuts,  pecans,  and  others.  Catesby  includes  pine  seeds  and 
persimmon  berries  in  their  diet,  and  Prof.  D.  E.  Lantz  states  that 
they  frequently  gather  and  store  seeds  of  the  cultivated  cherry. 
Stomachs  of  three  specimens  of  volans  taken  in  Virginia  in  January 
contained  finely  chewed  chestnut  meats,  a  few  pieces  of  acorn,  and 
several  beetle  larvae — probably  secured  with  the  nuts.  Several  stom- 
achs taken  at  Red  Fork,  Okla.,  in  June  contained  only  remains  of 
insect  larva?.  Buds  of  trees  are  said  to  be  eaten  in  winter,  and  corn 
or  other  grain  sometimes  is  taken.  Beetles  and  perhaps  other  in- 
sects constitute  a  part  of  the  animals'  fare.  They  have  a  decided  taste 
for  meat,  and  are  so  frequently  caught  by  fur  trappers  in  meat- 
baited  traps  set  for  larger  game  as  to  constitute  a  nuisance. 
Rowley  states  that  he  has  known  the  small  eastern  species  {volans) 
to  gnaw  the  edges  of  meat  hung  from  the  rafters  of  an  outhouse.2 
Flying  squirrels  are  suspected  by  some  observers  of  destroying  the 
eggs  of  wild  birds,  and  Merriam  remarks  that  the  eagerness  with 
which  these  animals  seize  and  feast  upon  a  dead  bird  placed  within 
their  reach  would  indicate  that  they  are  not  strangers  to  such  a 
repast.3  King  states  that  in  confinement  they  eat  birds'  eggs  with 
great  satisfaction,  and  relates  an  instance  of  the  instant  killing  by 
them  of  a  nestling  chipping  sparrow  placed  in  their  cage.  A  large 
moth  introduced  alive  was  also  quickly  captured  and  eaten.4 

A  writer  in  "  Forest  and  Stream,"  describing  the  habits  of  flying 
squirrels  in  captivity,  states  that  they  were  fond  of  parsley,  pansy 
flowers,  and  apple-tree  twigs,  the  last  being'  devoured  with  great 
avidity,  the  leaves  and  bark  stripped  clean  from  the  wood.  This 
writer  states  also  that  the  squirrels  were  fond  of  moths  and  grass- 

1  Seton,  E.  T.,  Life  Hist.  Northern  Anim.,  p.  442,  1909. 

2  Rowley,  J.,  Abstr.  Proc.  Linn.  Soc.  New  York,  Nos.  13-14,  p.  40,  1902. 
s  Merriam,  C.  Hart.,  Trans.  Linn.  Soc.  New  York,  II,  p.  100,  1884. 
'King,  F.  H.,  Anier.  Nat.,  XVII,  pp.  40-41,  1883. 


10  NORTH   AMERICAN   FAUNA.  [No.  44. 

hoppers,  but  refused  crickets,  and  their  preference  in  nuts  was  for 
acorns,  filberts,  and  pecans.1 

Stores  of  food  laid  up  by  flying  squirrels  for  winter  use  have  occa- 
sionally been  found,  indicating  that  the  hoarding  habit  is  probably 
general.  Audubon  and  Bachman  record  an  instance  of  a  martin 
box  occupied  by  a  considerable  number  of  flying  squirrels,  in  which 
was  found  a  quantity  of  hickory  nuts,  chestnuts,  acorns,  and  corn.2 

ECONOMIC    STATUS. 

The  food  habits  of  the  flying  squirrels  are  such  that  they  are  almost 
entirely  harmless.  So  far  as  known  they  do  not  damage  farm  crops, 
and  the  relatively  small  number  of  nuts  which  they  consume  does  not 
seriously  affect  the  total  supply.  They  have  been  suspected  of  de- 
stroying the  eggs  and  young  of  wild  birds,  but  no  definite  evidence 
on  this  point  is  forthcoming.  Occasionally  they  may  take  posses- 
sion of  bird  boxes  or  enter  the  lofts  of  dwelling  houses  and  thus  be- 
come objectionable,  but  ordinarily  their  gentle  and  confiding  ways 
and  their  interesting  habits  make  them  desirable  neighbors. 

PELAGE  AND  MOLT. 

The  pelage  in  members  of  this  genus  is  of  moderate  length,  very 
dense,  fine,  and  silky  in  texture.  Apparently  only  one  complete  molt 
occurs  in  a  year,  in  the  autumn,  from  September  to  November.  The 
new  pelage  usually  appears  first  on  the  sides  and  spreads  thence  over 
the  back,  the  head  and  shoulders  being  the  last  parts  to  be  renewed. 
The  effects  of  wear  usually  are  not  noticeable  until  March,  but  from 
then  till  November  many  worn  specimens  are  to  be  found,  though 
apparently  a  considerable  proportion  retain  the  winter  pelage 
throughout  the  summer  with  little  indication  of  wear.  The  head, 
nape,  and  rump  are  first  subject  to  wear,  though  frequently  the  entire 
upperparts  and  tail  become  much  worn.  In  summer  there  is  a  gen- 
eral tendency  to  a  reddening  of  the  hairs  on  the  upperparts. 

In  Glaucomys  volans  and  in  most,  if  not  all,  of  the  races  of  G. 
sabrinus  the  middle  portion  of  the  soles  in  summer  is  naked,  the  same 
portions  being  furred  in  winter  pelage  in  all  forms  except  the  south- 
ern races  of  volans.  In  one  race  (G.  volans  volans)  the  hind  toes 
are  conspicuously  marked  with  white  in  winter,  but  are  brown  in 
summer.  The  exact  manner  of  this  change  is  not  apparent,  but  it 
probably  is  accomplished  by  a  partial  spring  molt.3 

1 "  C.  C.  H.,"  Forest  and  Stream,  LII,  pp.  125-126,  1899. 

a  Audubon  and  Bachman,  Quad.  North  Amer.,  I,  p.  220,  1846. 

3  For  a  detailed  account  of  the  molt  in  O.  volans  see,  under  that  species,  pp.  20-22. 


1918.]  GENUS  GLAUCOMYS.  11 

EXPLANATION  OF  CRANIAL  MEASUREMENTS. 

The  following  measurements  of  skulls  of  this  genus  have  been  taken 
in  millimeters : 

Greatest  length. — From  anterior  border  of  nasals  to  posterior 
border  of  supraoccipital  in  median  line. 

Zygomatic  breadth. — Greatest  breadth  across  zygomata. 

Mastoidal  breadth. — Greatest  breadth  across  mastoids. 

Inter  orbital  breadth. — Shortest  distance  across  frontals  in  front  of 
postorbital  processes. 

Postorbital  breadth. — Shortest  distance  across  frontals  behind 
postorbital  processes. 

Length  of  nasals. — Greatest  length  of  nasals,  measured  along 
median  line. 

Maxillary  toothrow. — Alveolar  length  of  maxillary  molar-premolar 
toothrow. 

MATERIAL  EXAMINED. 

For  the  present  revision  the  writer  lias  assembled  for  study 
1,052  specimens.  About  half  of  these  are  contained  in  the  United 
States  National  Museum,  including  the  Biological  Survey  and  Mer- 
riam  collections.  The  remainder  have  been  borrowed  from  other 
American  museums  and  from  private  collections,  so  that  practically 
all  the  available  material  in  this  country  has  been  at  the  disposal  of 
the  reviser.1 

Genus  GLAUCOMYS  Thomas. 

Mus  Linnaeus,  Syst.  Nat,  Ed.  10,  p.  63  (part),  1758. 

Sciurus  Pallas,  Nov.  Spec.  Glires,  p.  350  (part),  1778.  Not  Sciurus  Linnaeus,  1758. 
Pteromys  Tiedemann,  Zool.,  I,  p.  451  (part),  1808.  Not  Pteromys  G.  Cuvier,  1800. 
Sciuropterus  Lesson,  Manuel  de  Mamm.,  p.  242,  1827  (part)  ;  Desmarest,  Diet. 

Sci.  Nat.,  p.  140,  1827  (part).     Not  Sciuropterus,  F.  Cuvier,  1825. 
Olaucomys  Thomas,  Ann.  and  Mag.  Nat.  Hist.    (ser.  8),  I,  p.  5,  1908.     Type, 

Mus  volans  Linnaeus. 

1  For  the  loan  of  this  material  the  writer  desires  to  extend  thanks  to  the  owners  and 
custodians  as  follows  :  Dr.  J.  A.  Allen,  of  the  American  Museum  of  Natural  History ; 
Messrs.  Samuel  Henshaw  and  Outram  Bangs,  of  the  Museum  of  Comparative  Zoology ; 
Dr.  Witmer  Stone,  of  the  Academy  of  Natural  Sciences  of  Philadelphia  ;  Messrs.  Charles 
B.  Cory  and  W.  II.  Osgood,  of  the  Field  Museum  of  Natural  History  ;  Dr.  Joseph  Grinnell, 
of  the  Museum  of  Vertebrate  Zoology,  University  of  California;  Mr.  P.  A.  Taverner,  of 
the  Victoria  Memorial  Museum ;  Mr.  F.  Kermode,  of  the  Provincial  Museum,  Victoria, 
British  Columbia  ;  Dr.  W.  J.  Holland  and  Mr.  W.  E.  Clyde  Todd,  of  the  Carnegie  Museum  ; 
Dr.  A.  G.  Ruthven,  of  the  University  of  Michigan  ;  Mr.  C.  D.  Bunker,  of  the  Kansas 
University  Museum  ;  Mr.  Myron  H.  Swenk,  of  the  University  of  Nebraska  ;  Mr.  J.  D. 
Figgins,  of  the  Colorado  Museum  of  Natural  History ;  Mr.  J.  O.  Snyder,  of  Leland 
Stanford,  Jr.,  University ;  Dr.  Barton  W.  Evermann,  of  the  California  Academy  of 
Sciences ;  Mr.  Henry  L.  Ward,  of  the  Milwaukee  Public  Museum ;  Mr.  W.  E.  Saun- 
ders, of  London,  Ontario ;  Mr.  Edward  R.  Warren,  of  Colorado  Springs,  Colo. ; 
Mr.  G.  L.  Kirk,  of  Rutland,  Vt,  and  Dr.  H.  V.  Ogden,  of  Milwaukee,  Wis. ;  and  to 
Mr.  Francis  Harper  for  the  photograph  from  which  the  frontispiece  was  made. 


12  NORTH   AMERICAN   FAUNA.  [No.  44. 

HISTORY  AND  NOMENCLATURE. 

The  small  eastern  flying  squirrel  {Glaucomys  volans),  which  is 
very  common  in  Virginia  and  the  other  South  Atlantic  States,  at- 
tracted the  notice  of  the  earliest  colonists  and  through  their  descrip- 
tions early  became  known  to  naturalists.  Captain  John  Smith,  in 
his  History  of  Virginia,  published  originally  in  1624,  refers  to  the 
animal  in  the  following  words : 

A  Small  beast  they  have  they  call  Assapanick,  but  we  call  them  flying  Squir- 
rels, because  spreading  their  legs,  and  so  stretching  the  largenesse  of  their  skins, 
that  they  have  beene  seene  to  fly  30  or  40  yards.1 

This  species  was  described  more  or  less  fully  also  in  the  seventeenth 
and  eighteenth  centuries  by  numerous  other  writers,  including  Ray,2 
Catesby,3  and  Edwards.4  Both  Catesby  and  Edwards  figured  the 
animal,  and  Catesby  described  its  habits  in  the  following  words : 

These  Squirrels  are  gregarious,  travelling  from  one  Tree  to  another  in  com- 
panies of  ten,  or  twelve  together.  When  I  first  saw  them,  I  took  them  for  dead 
Leaves,  blown  one  Way  by  the  Wind,  but  was  not  long  so  deceived,  when  I 
perceived  many  of  them  to  follow  one  another  in  one  Direction:  They  will  fly 
fourscore  Yards  from  one  Tree  to  another.  They  can  not  rise  in  their  Flight, 
nor  keep  in  a  horizontal  Line,  but  descend  gradually,  so  that  in  Proportion  to 
the  Distance  the  Tree,  they  design  to  fly  to,  is  from  them,  so  much  the  higher 
they  mount  on  the  Tree  they  fly  from      *     *     *. 

The  species  of  eastern  Canada  was  briefly  described  by  Sagard- 
Theodat  in  163G,5  and  that  of  southern  Mexico  by  Hernandez  in  1651.6 

GENERIC    NAMES. 

Placed  first  by  Linnaeus  in  the  genus  Mus,  the  American  flying 
squirrels  were  later  associated  with  Sciurus  by  Gmelin,  Pallas,  and 
Shaw.  In  1800  G.  Cuvier  proposed  a  new  genus,  Pteromys,  to  include 
the  European  flying  squirrel  (Sciurus  volans  Linnaeus)  and  the  large 
East  Indian  species  (Sciurus  yetaurista  Gmelin).  The  American 
forms  were  referred  by  later  writers  to  this  genus.  F.  Cuvier,  in 
1823,7  pointed  out  the  characters  distinguishing  the  American  and 
European  flying  squirrels  from  the  large  Asiatic  species,  and. in  1825 
proposed  the  name  Sciuropterus  for  the  European  animal,8  but  this 
name  was  not  generally  accepted  until  Allen  adopted  it  for  the 
American  species  in  his  monograph  of  the  Sciurido?  (1877). 9     This 

1  Smith,  John,  The  Generall  Historie  of  Virginia,  &c,  p.  27,  1624. 

2  Ray,  John,  Synop.  Quad.,  p.  215,  1693. 

3  Catesby,  Mark,  Nat.  Hist.  Carolina,  II,  rp.  76-77,  1743. 

4  Edwards,  George,  Nat.  Hist.  Birds,  IV,  p.  191,  1751. 

5  Sagard-Theodat,  Gabriel,  Hist.  Canada,  III,  p.  745,  1636. 

6  Hernandez,  F.,  Hist.  Anim.  et  Min.  Novre.,  Hisp.,  p.  9,  1651  :  In  Rerum  Med.  Novte  Hisp. 
Thesaurus. 

7  Cuvier,  P.,  Mem.  Mus.  Hist.  Nat,  Paris,  pp.  125-127,  1823. 

8  Cuvier,  F.,  Dents  des  Mamm.,  p.  255,  1825. 

9  Allen,  J.  A.,  Monogr.  North  Amer.  Rodentia  :  Rept.  U.  S.  Geol.  Surv.  Terr.,  XI,  p.  653, 
1877. 


1918.]  GENUS   GLAUCOMYS.  13 

name  continued  in  current  use  until  1914,  when  Miller1  showed  that 
according  to  modern  rules  for  fixing  types  it  must  give  place  to 
Pteromys  of  earlier  date,  the  type  of  the  latter  having  been  fixed  by 
Fleming,2  in  1822,  on  Sciurus  volans — the  European  frying  squirrel. 
Thomas,3  in  1908,  in  a  revision  of  the  genera  of  flying  squirrels  sepa- 
rated the  xVmerican  forms  subgenerically  from  the  European  members 
of  the  genus  under  the  name  Glaucomys,  and  in  1915  the  present 
writer  raised  the  group  to  generic  rank.4 

SPECIFIC   NAMES. 

Linnaeus,  in  the  10th  edition  of  his  Sy sterna  Naturae,5  described  the 
smaller  American  species  under  the  name  Mus  volans,  basing  his 
description  mainly  on  the  account  of  the  animal  as  given  by  Ray. 
Curiously  enough,  on  a  later  page  of  the  same  work  he  described  the 
European  flying  squirrel  as  Sciurus  volans.  The  two  species  nat- 
urally were  united  in  one  genus  by  later  authors,  and  the  duplica- 
tion of  the  name  volans  necessitated  the  renaming  of  one  of  them. 
Pallas,  therefore,  in  1778,6  proposed  the  name  volucclla  for  the  Amer- 
ican animal,  which  designation  remained  in  general  use  until  1890, 
when  Jordan  revived  the  original  name  volans.7  The  larger  Amer- 
ican species  was  named  Sciurus  hudsonius,  in  1788,  by  Gmelin,8  but 
this  name  proved  to  be  preoccupied  by  Erxleben's  Sciurus  vulgaris 
hudsonicus  applied  to  the  American  chickaree,  so,  in  1801,  Shaw  pro- 
posed in  its  place  the  name  sabrinus.0  During  the  nineteenth  century 
both  these  names  were  used  for  the  northern  flying  squirrel — kud- 
sonius  by  Baird,  Allen,  and  others  and  sahsrinus  b}T  Richardson,  Au- 
dubon and  Bachman,  and  others.  Finally,  in  1891,  the  nomenclatural 
question  involved  having  been  settled,  Rhoads10  revived  the  name 
sabrinus,  which  has  remained  current  ever  since.  In  addition  to 
Pallas's  name  volucella,  the  small  species  Mus  volans  Linnaeus  was 
several  times  renamed  during  the  early  years  of  the  nineteenth 
centur}'.  Tiedemann,  in  1808,  proposed  for  it  the  name  Pteromys 
virginianus,11  and  Oken,  in  1816,  called  the  species  Pteromys  ameri- 
cana.12 

Richardson,  in  1828,13  described  a  new  species,  alpinus,  from  mate- 
rial collected  by  Thomas  Drummond  in  western  Canada,  but  the  fol- 

1  Miller,  G.  S.,  Proc.  Biol.  Soc.  Washington,  XXVII,  p.  216,  1914. 

2  Fleming,  J.,  Philos.  Zool.,  II,  p.  190,  1822. 

•Thomas,  O.,  Ann.  and  Mag.  Nat.  Hist.  (ser.  8),  I,  p.  5,  1908. 
«  Howell,  A.  II.,  Proc.  Biol.  Soe.  Washington,  XXVIII,  p.  109,  1915. 
ELinna>us,  Syst.  Nat,  Ed.  10,  I,  p.  G3,  1758. 
•Pallas,  Nov.  Spec.  Glircs,  p.  351,  1778. 

7  Jordan,  D.  S.,  Man.  Vert.  U.  S.,  Ed.  5,  321,  1890. 

8  Gmelin,  J.  F.,  Syst.  Nat.,  I,  p.  153,  1788. 
0  Shaw,  Gen.  Zool.,  II,  p.   157,  1801. 

10  Rhoads,  S.  N.,  Amer.  Nat.,  XXVIII,  p.  525,  1894. 
u  Tiedemann,  F.,  Zool.,  I,  p.  451,  1808. 

12  Oken,  Lorenz,  Lehrb.  der  Naturg.,  II,  p.  865,  1816. 

13  Richardson,  J.,  Zool.  Journ.,  Ill,  p.  519,  1828. 


14  NORTH   AMERICAN   FAUNA.  [No.  44. 

lowing  year  he  reduced  the  form  to  a  subspecies  of  sabrinus.1  Bach- 
man,  in  1839,  named  oregonensis,  a  new  species  from  the  lower  Colum- 
bia River,  and  after  this  date  no  new  names  were  proposed  for  the 
group  for  a  period  of  more  than  50  years. 

Audubon  and  Bachman  in  their  great  work,  the  Quadrupeds  of 
North  America,2  treated  the  four  species  already  described — volucella, 
sabrinus,  alpinus,  and  oregonensis.  Baird,  in  his  Mammals  of  North 
America,3  recognized  the  same  four  species  (using  the  name  hud- 
sonius,  however,  in  place  of  sabrinus),  but  was  able  to  add  very  little 
to  the  history  of  the  group.  Allen,  in  his  Monograph  of  the 
Sciuridse,4  united  the  four  forms  into  one  species,  volucella,  with 
hudsonius  as  a  variety,  placing  alpinus  and  oregonensis  in  synonymy. 
Bangs,  in  1896,  revised  the  eastern  members  of  the  genus,  raising 
sabrinus  again  to  the  rank  of  a  species  and  describing  as  new,  querceti, 
a  southern  race  of  volans,  and  silus,  a  supposed  new  species  from 
West  Virginia.5  Rhoads,  in  1897,  revised  the  western  forms  of  the 
genus,  reviving  alpinus  as  a  full  species  with  oregonensis  as  a  sub- 
species and  describing  three  additional  races — fuliginosus  from  the 
Cascade  Mountains,  Washington;  bangsi  from  central  Idaho;  and 
calif ornicus  from  the  San  Bernardino  Mountains,  California.6 

During  the  next  eight  years  9  new  forms  were  added  by  various 
authors  to  the  9  already  recognized — one  (goldmwii)  as  a  race  of 
volans  from  southern  Mexico,  two  (macrotis  and  makkovikensis)  as 
races  of  sabrinus,  five  {klamathensis,  olympicus,  lascivus,  Stephens i, 
and  zaphceus)  as  races  of  alpinus,  and  one  (yukonensis)  as  a  full 
species. 

In  1915  Swenk  proposed  as  new  Pteromys  volans  nebrascensis, 
and  the  present  writer  described  two  races  of  volans,  four  of  sabrinus, 
and  one  full  species,  bullatus,7  thus  raising  the  total  number  of  recog- 
nized forms  to  26.  In  the  present  revision  all  of  these  are  recognized 
except  silus  of  Bangs  and  nebrascensis  of  Swenk,  which  are  referred 
to  G.  volans  volans. 

GENERIC  CHARACTERS. 

Form  sciurine;  fore  and  hind  limbs  connected  by  a  broad  fold  of 
skin  extending  from  wrists  to  ankles,  supported  anteriorly  by  a 

1  Fauna  Boreali-Amer.,  I,  p.  195,  1829. 

2  Audubon  and  Bachman,  Quad.  North  Amer.,  I,  pp.  132,  216,  1846 ;  III,  pp.  202,  206, 
1854. 

3  Baird,  S.  F.,  Mamm.  North  Amer. :  Rept.  Expl.  and  Surv.  R.  R.  Pac,  VIII,  pp.  286- 
290,  1857. 

*  Allen,  J.  A.,  Monogr.  North  Amer.  Rodentia  :  Rept.  U.  S.  Geol.  Surv.  Terr.,  XI, 
pp.  653-666,  1877. 

6  Bangs,  O.,  Proc.  Biol.  Soc.  Washington,  X  pp.  162-166,  1896. 

0  Rhoads,  S.  N.f  Proc.  Acad.  Nat.  Sci.  Philadelphia,  pp.  314-327,  1897. 

7  Howell,  A.  II.,  Proc.  Biol.  Soc.  Washington,  XXVIII,  pp.  109-114,  1915  ;  in  the  present 
paper  bullatus  is  treated  as  a  subspecies  of  sabrinus. 


1918.]  GENUS  GLAUCOMYS.  15 

slender  cartilaginous  process  growing  from  the  wrist.  Tail  broad, 
much  flattened,  densely  haired,  its  sides  usually  nearly  parallel  or 
slightly  narrowed  at  each  end,  the  tip  evenly  rounded.  Soles  with 
4  tubercles  at  base  of  toes ;  no  metatarsal  pad ;  palms  with  5  tuber- 
cles— 3  at  base  of  fingers,  one  at  base  of  the  rudimentary  thumb,  and 
one  opposite  the  latter  on  outside  of  wrist.  Mammae,  8,  as  follows: 
Pectoral,  | ;  abdominal,  § ;  inguinal,  ^.  Skull  with  smoothly  rounded 
braincase,  flattened  in  some  species,  subglobular  in  others;  nasals 
abruptly  depressed  at  tip;  dorsal  outline  of  skull  from  nasals  to 
postfrontal  region  nearly  straight,  then  abruptly  depressed  to 
occiput;  frontals  long  and  narrow,  the  interorbital  region  decid- 
edly longer  than  broad;  interorbital  constriction  pronounced, 
usually  with  a  distinct  notch  at  anterior  base  of  postorbital 
processes;  postorbital  processes  broad  at  base,  tapering  abruptly  to 
a  point,  slightly  depressed  at  tip;  zygomata  moderately  expanded 
posteriorly,  less  widely  anteriorly;  audital  bulla)  large,  subcircular 
in  outline,  smoothly  rounded ;  pterygoids  slender  and  rather  low,  the 
hamular  processes  usually  touching  audital  bullae;  dentition,  I,f; 
PM,  -f;  M,  f=22;  maxillary  toothrows  approximately  parallel; 
anterior  upper  premolar  very  small,  terete,  its  crown  simple;  trans- 
verse ridges  of  upper  molariform  teeth  continuous. 

The  skull  of  Glaucomys  differs  from  that  of  Pteromys1  in  the 
following  particulars  (see  PI.  VII,  figs.  11-14)  :  Rostrum  with  sides 
not  parallel,  but  tapering  gradually  from  tip  to  point  of  union  with 
the  zygomata;  nasal  branches  of  premaxillae  relatively  much  wider; 
zygomata  narrower  anteriorly,  not  standing  out  squarely  from  the 
rostrum ;  maxillary  arm  of  zygoma  much  weaker,  the  median  portion 
of  the  arch  nearly  vertical  (nearly  horizontal  in  Pteromys) ;  frontals 
relatively  longer  and  narrower,  the  surface  nearly  flat  or  with 
a  very  slight  depression;  postorbital  processes  broader  at  the  base 
and  less  attenuate;  braincase  more  swollen,  its  outlines  smooth  and 
rounded;  occiput  not  noticeably  depressed;  anterior  palatine  fora- 
mina relatively  smaller;  mandible  relatively  longer  and  more  slen- 
derly built,  the  masseteric  ridge  less  strongly  developed,  terminating 
at  a  point  slightly  posterior  to  the  premolar ;  angular  process  weaker, 
its  border  forming  a  regular  curve  (not  abruptly  bent  outwards,  as 
in  Pteromys)-,  coronoid  process  larger  and  longer;  enamel  pattern 
of  molariform  teeth  simple,  the  transverse  ridges  of  the  upper  molars 
complete  (in  Pteromys  PM>,  MA,  and  M^  having  the  posterior  ridge 
cut  by  a  deep  reentrant  fold). 

1  As  restricted  to  the  small  palaearctic  species  P.  volans,  P.  buchneri,  and  related  forms. 


16  NORTH   AMERICAN    FAUNA.  INo.  44. 

List  of  Species  and  Subspecies,  with  Type  Localities. 

Glaucomys  volans  group: 

G  lav  corny  s  volans  volans  (Linnaeus) Virginia. 

volans  saturatus  Howell Dothan,  Alabama. 

volans  qucrceti  (Bangs) Citronelle,  Florida. 

volans  texensisHoweW Sour  Lake,  Texas  (7  miles  north- 
east ) . 

volans  gold mani  (Nelson) Teopisca,     Chiapas      (20     miles 

southeast). 
Glaucomys  sabrinus  group: 

Glaucomys  sabrinus  sabrinus  (Shaw) Mouth  of  Severn  River,  Ontario. 

sabrinus  makkovikensis  (S^rnborger)  -Makkovik,  Labrador. 

sabrinus  macrotis  (Mearns) Hunter     Mountain      (Catskills), 

Greene  County,  New  York. 

sabrinus  cancscens  Howell Portage  la  Prairie,  Manitoba. 

sabrinus  bangsi  (Rhoads) Idaho  County,  Idaho. 

sabrinus  alpinus  (Richardson) Jasper  House,  Alberta. 

sabrinus  yukonensis  (Osgood) Camp   Davidson,    Yukon   River, 

Yukon. 

sabrinus  zaphams  (Osgood) Helm  Bay,  Cleveland  Peninsula, 

Alaska. 

sabrinus  oregonensis  (Bachman) "Pine   woods   of  the   Columbia, 

near  the  sea  " — probably  near 
the  present  site  of  St.  Helens, 
Oregon. 

sabrinus  columbiensis  Howell Okanogan,  British  Columbia. 

sabrinus  fid iginosus  (Rhoads) Cascade  Mountains,  near  Martin 

Station,  Kittitas  County, 
Washington. 

sabrinus  latipes  Howell Glacier,  British  Columbia. 

sabrinus  olympicus  (Elliot) Happy  Lake,  Olympic  Mountains, 

Washington. 

sabrinus  bullatus Howell Sawtooth  [Alturas]  Lake,  Idaho. 

sabrinus  klamathensis  (Merriam) Fort  Klamath,  Oregon. 

sabrinus  flavivcntris  Howell Head    of    Bear    Creek,    Trinity 

County,  California. 

sabrinus  lascivus  (Bangs) Tallac,    Eldorado    County,    Cali- 
fornia. 

sabrinus  calif  or  nicus  (Rhoads) Squirrel    Inn,     San    Bernardino 

Mountains,  California. 

sabrinus  stephensi  (Merriam) Sherwood,     Mendocino     County, 

California. 

Key  to  Species  and  Subspecies. 

a1.  Hairs  on  helly  white  at  roots  (volans  group). 

b\  Colors  paler  (drab,  pinkish  cinnamon,  or  yellowish  wood  brown). 

&.  Size  larger;  hind  toes  white  in  winter  pelage volans  (p.  19). 

c2.  Size  smaller ;  hind  toes  not  white  in  winter. 
d\  Skull  relatively  short  and  broad  (greatest  length,  32.7-34  mm.)  ;  bulla? 

smaller texensis  (p.  27). 

d*.  Skull  relatively  long  and  narrow   (greatest  length,  33.2-35.3)  ;  bull* 
larger . querceti  (p.  26). 


1918.]  KEY   TO    SPECIES  AND   SUBSPECIES.  17 

6a.  Colors  darker  (dark  sayal  brown;  hair  brown;  pale  snuff  brown). 

c1.  Sides  of  face  marked  with  fuscous goldmani  (p.  28). 

c3.  Sides  of  face  not  marked  with  fuscous saturatus  (p.  24). 

a2.  Hairs  on  belly  gray  at  roots  (sabrinus  group). 

&\  Bulla?  very  large bullatus  (p.  51). 

b2.  Bullae  medium  or  small. 
&.  Soles  yellowish. 

d\  Underparts  strongly  yellowish flaviventris  (p.  54). 

d2.  Underparts  not  strongly  yellowish. 
e\  Underparts  tinged  with  yellow  or  cream-buff. 

f.  Under  side  of  tail  darker  (deep  pinkish  cinnamon), 

columbiensis  (p.  45). 
f.  Under   side   of   tail   paler    (light   pinkish   cinnamon   or   colonial 

buff) klamathensis  (p.  52). 

e*.  Underparts  tinged  with  avellaneous lascivus  (p.  55). 

c*.  Soles  not  yellowish. 

d1.  Underparts   heavily    washed   with    wood    brown   or   some   shade   of 
cinnamon  or  buff. 
e1.  Upperparts  strongly  rufescent  (mikado  brown  or  pecan  brown), 

oregonensis  (p.  44). 
e2.  Upperparts  not  strongly  rufescent. 

f.  Tail  strongly  clouded  with  fuscous  beneath zaphaeus  (p.  43). 

f.  Tail  not  strongly  clouded  with  fuscous  beneath. 
g\  Size  larger  (greatest  length  of  skull  usually  more  than  42  mm.), 

latipes  (p. 48). 
g*.  Size  smaller  (greatest  length  of  skull  usually  less  than  42  mm.). 
h\  Colors  darker  (wood  brown  to  snuff  brown), 

olympicus  (p.  49). 

h2.  Colors  paler  (pale  sayal  brown) fuliginosus  (p.  47). 

d3.  Underparts  whitish,  or  moderately  washed  with  wood  brown  or  some 
shade  of  cinnamon  or  buff. 
e\  Upperparts  vinaceous  or  brownish    (pinkish  cinnamon,   vinaceous- 
cinnamon,  or  sayal  brown). 

f.  Upperparts  very  pale  (light  pinkish  cinnamon) canescens  (p.  37). 

f.  Upperparts  darker  (vinaceous-cinnamon  to  sayal  brown). 
g1.  Underparts  white  or  faintly  washed  with  pinkish  cinnamon  or 
wood  brown. 
h\  Size  larger    (greatest  length   of  skull   in   adults  more  than 
39  mm.). 
i\  Under  side  of  tail  strongly  washed  with  fuscous  or  dark 
brown. 
j1.  Size  larger   (greatest  length  of  skull  usually  more  than 

40.4  mm.) makkovikensis  (p.  34). 

f.  Size  smaller  (greatest  length  of  skull  usually  less  than 

40.4  mm.) sabrinus  (p.  31). 

i*.  Under  side  of  tail  not  strongly  washed  with  fuscous  (pale 
or  deep  pinkish  cinnamon). 

f.  Upperparts  vinaceous-cinnamon columbiensis  (p.  45). 

j'.  Upperparts  sayal  brown stephensi  (p.  57). 

h*.  Size  smaller  (greatest  length  of  skull  less  than  39  mm.), 

macrotis  (p.  35), 

14520°— 18— No.  44 2 


18  NORTH   AMERICAN   FAUNA.  [No.  44. 

g9.  Underparts    moderately    washed    with    pinkish    cinnamon    or 
avellaneous. 
h\  Size  larger  (average  length  of  tail  about  165  mm.), 

yukonensis  (p.  41). 
h'.  Size  smaller  (average  length  of  tail  about  142  mm.), 

bangsi  ( p.  38 ) . 
er.  Upperparts  drab,  gray,  or  wood  brown. 

f.  Size  larger;  tail  heavily  marked  with  fuscous alpinus  (p. 40). 

f.  Size  smaller;  tail  not  heavily  marked  with  fuscous. 

g\  Under  side  of  tail  deep  pinkish  cinnamon bangsi  (p.  38). 

g2.  Under  side  of  tail  pinkish  buff  or  pale  olive-buff. 
h1.  Skull  with  deeper  braincase;  soles  never  washed  with  yellow 

or  buff . stephensi  (p.  57). 

h2.  Skull   with   shallower   braincase ;    soles  often   washed   with 
yellow  or  buff. 

i\  Larger  and  darker lascivus  (p.  55). 

i2.  Smaller  and  paler californicus  (p.  56). 

DESCRIPTIONS   OF   SPECIES  AND  SUBSPECIES. 

Glaucomys  volans  Group. 

GLAUCOMYS  VOLANS   (Linnaeus). 
[Synonymy  under  subspecies.] 

General  characters. — Size  small  (hind  foot  32  mm.  or  less) ;  hairs 
on  underparts  white  to  the  roots,  except  on  hind  legs  and  flying  mem- 
branes, where  they  are  plumbeous  at  base. 

Cranial  characters. — Skull  small  (greatest  length  35.7  mm.  or  less) ; 
rostrum  short ;  audital  bullae  moderately  inflated. 

Color. — Upperparts  drab,  pinkish  cinnamon,  sayal  brown,  pale 
snuff  brown,  hair-brown,  or  yellowish  wood  brown,  the  bases  of  the 
hairs  deep  neutral  gray ;  sides  of  face  smoke  gray,  often  shaded  with 
fuscous  or  buff;  borders  of  flying  membrane  above,  fuscous,  clove 
brown,  or  blackish  brown ;  fore  feet  buffy  white,  hair-brown,  or  gray- 
ish buff ;  hind  feet  hair-brown,  fuscous,  or  mouse  gray,  the  toes  in  one 
race  white  in  winter;  tail  above,  hair-brown,  snuff  brown,  verona 
brown,  fuscous,  or  drab;  beneath,  pinkish  cinnamon,  vinaceous- 
cinnamon,  or  pinkish  buff;  underparts  creamy  white,  the  sides  often 
edged  with  pinkish  cinnamon,  vinaceous-cinnamon,  or  pinkish  buff. 

Geographic  distribution. — Eastern  United  States  and  extreme 
southern  Canada,  from  southern  New  Hampshire,  northern  New 
York  (Lewis  County),  southern  Ontario  (London), central  Michigan, 
northern  Wisconsin  (Burnett  County),  and  central  Minnesota  (Ait- 
kin County),  south  to  the  Gulf  coast  and  southern  Florida  (Fort 
Myers);  west  to  eastern  Nebraska  (Otoe  and  Nemaha  Counties), 
eastern  Kansas  (Douglas  and  Woodson  Counties),  southwestern 
Oklahoma  (Wichita  Mountains),  and  eastern  Texas  (Parker,  Bas- 
trop, and  DeWitt  Counties) ;  also  the  highlands  of  Chiapas  and 


1918.] 


GLAUCOMYS  VOLANS  GROUP. 


19 


Guatemala  and  probably  locally  throughout  eastern  Mexico.  Zonal 
range  from  the  Transition  through  Upper  and  Lower  Austral  Zones 
(%  2). 

GLAUeOMYS  VOLANS  VOLANS  (Linnaeus). 

Small  Eastern  Flying  Squirrel. 

[PI.  I ;  PI.  II,  fig.  2  ;  PI.  IV,  fig.  2  ;  PI.  VI,  fig.  1.] 

[Mus]  volans  Linnaeus,  Syst.  Nat.,  ed.  10,  p.  63,  1758  (based  chiefly  on  Ray  and 

Edwards). 
Sciurus  volucella  Pallas,  Nov.  Spec.  Glh-es,  p.  351,  177S  (apparently  a  renaming 

of  Mus  volans  Linnaeus,  but  based  primarily  on  Edwards  and  Catesby). 
Ptcromys       virginianus 

Tiedemann,      Zool., 

I,  p.  451,  1808   (re- 
naming of  Mus  vo- 
lans Linnaeus). 
Pteromys         americana 

Oken,     Lehrb.     der 

Naturg.,  II,  p.  865, 

1816    (renaming  of 

Sciurus       volucella 

Pallas). 
Pteromys  volucella  Des- 

marest,  Nouv.  Diet. 

d  '  H  i  s  t .       Nat., 

XXVII,       p.       406, 

1818;  Audubon  and 

Bachman,        Quad. 

North  Amer.,   I.   p. 

216,      1846,      plate 

XXVIII ;         Baird, 

M  a  m  m  .     North 

Amer. :  Kept.  Expl. 

and    Surv.      It.    R. 

Pac,   VIII,   p.   286, 

1857. 
Sciuropterus      vol  ucella 

Lesson,   Manual   de 

Mamm.,  p.  242, 1827. 
Sciuropterus  americanus 

Desmarest,  Diet.  Sci. 

Nat.,  p.  140,  1827. 

?  Pteromys  cucullatus  Fischer,  Synop.  Mamm.,  p.  365,  1829  (type  locality,  Vir- 
ginia (?)  ;  based  on  the  "Sciurus,  Virginianus,  volans"  of  Seba).1 
Sciuropterus  volucella,  var.  volucella  Allen,  Proc.  Boston  Soc.  Nat.  Hist.,  XVI, 

p.  189,  1874 ;  Monogr.  North  Amer.  Rodentia :  Rept.  U.  S.  Geol.  Surv.  Terr., 

XI,  p.  655,  1S77. 


Fig.  2. — Map  showing  the  distribution  of  the  subspecies  of 
Glaucomys   volans,  based  on  specimens  examined. 


1  Seba's  figure  does  not  agree  with  any  known  species  of  flying  squirrel ;  Thomas  (Proc. 
Zool.  Soc.  London,  p.  148,  1911)  has  identified  it  as  a  young  Petaurista,  but  the  flattened 
tail  strongly  suggests  Glaucomys ;  Linnaeus  cites  Seba's  description  in  the  synonymy  of 
Mus  volans. 


20  NORTH   AMERICAN   FAUNA.  I  No.   44. 

8[ciuropterus]  volans  Jordan,  Man.  Vert.  U.  S.,  ed.  5,  p.  321,  1890. 

Sciuropterus  situs  Bangs,  Proc.  Biol.  Soc.  Washington,  X,  p.  163,  1896.  Type, 
No.  4931,  Mus.  Comp.  Zool.  (formerly  in  collection  E.  A.  &  O.  Bangs)  ; 
from  top  of  Katis  Mountain,  near  White  Sulphur  Springs,  West  Virginia; 
September  2,  1895. 

Pteromys  volans  nebrascensis  Swenk,  University  Studies  (Lincoln,  Nebraska), 
XV,  p.  151,  "April  "  [=  Sept.  25],  1915.  Type,  No.  286,  Coll.  State  Entomolo- 
gist, Univ.  Nebraska ;  from  Nebraska  City,  Nebr.,  Nov.  26,  1914. 

Type  locality. — Virginia.1 

Distribution. — Northeastern  United  States  and  extreme  southern 
Canada,  from  central  Minnesota,  Wisconsin,  and  Michigan,  southern 
Ontario,  northern  New  York7 (Lewis  County),  and  southern  New 
Hampshire  south  to  North  Carolina  (Raleigh),  Tennessee  (Nash- 
ville), and  northern  Arkansas  and  Oklahoma  (Boston  Mountains) ; 
west  to  eastern  Nebraska  (Otoe  and  Nemaha  Counties)  and  eastern 
Kansas  (Douglas  and  Woodson  Counties). 

Characters. — Size  large  (for  the  group);  colors  pale;  upperparts 
varying  from  drab  to  pinkish  cinnamon;  toes,  in  winter,  usually 
strongly  marked  with  white  and  soles  haired  to  the  bases  of  the  toes. 

Color. — Winter  pelage:  Upperparts  varying  from  drab  through 
numerous  intermediate  shades  to  pinkish  cinnamon,  shaded  along 
sides  with  pinkish  buff ;  hairs  on  back  with  an  indistinct,  subterminal 
band  of  brown  (not  visible  on  the  surface) ,  their  bases  deep  neutral 
gray;  sides  of  face  smoke  gray,  often  washed  with  pinkish  buff; 
eye-ring  fuscous  or  clove  brown;  ears  hair-brown  or  drab,  scantily 
haired ;  upper  surface  of  flying  membrane  fuscous  to  blackish  brown ; 
under  surface  edged  with  light  pinkish  cinnamon ;  underparts  white, 
with  a  creamy  tinge,  the  hairs  white  to  the  roots  (except  on  hind 
legs,  where  their  bases  are  neutral  gray) ;  tail,  above,  snuff  brown  to 
drab,  shading  on  sides  to  hair-brown;  beneath,  light  pinkish  cinna- 
mon to  pinkish  buff,  shaded  in  some  specimens  with  drab-gray ;  front 
feet  buffy  white  or  hair-brown,  sometimes  shaded  with  gray;  hind 
feet  hair-brown  (rarely  mouse  gray),  the  toes  and  inner  borders  of 
feet  white.  Summer  pelage:  Similar  to  the  winter  pelage,  but  gen- 
eral tone  of  upperparts  usually  darker,  caused  in  part  by  reddening 
of  the  hairs  and  in  part  by  wearing  away  of  the  tips,  thus  exposing 
portions  of  the  underfur;  toes  without  white  markings;  middle  por- 
tion of  soles  naked. 

Molt. — From  the  large  series  of  skins  of  this  race  which  have  been 
examined  it  has  been  possible  to  assemble  a  series  of  97  specimens 
representing  every  month  in  the  year  and  showing  fully  all  the  pelage 
gradations  from  one  season  to  another.     The  following  notes  on  the 

1  Fixed  by  Elliot,  Synop.  Mamm.  North  Amer.  :  Field  Columb.  Mus.,  Zool.  Ser.,  II,  p.  109, 
1901  (see  also  Thomas,  Proc.  Zool.  Soe.  London,  p.  148,  1911). 


1618.]  GLAUCOMYS  VOLANS  GROUP.  21 

pelage  of  these  specimens,  arranged  chronologically,  will  serve  to 
make  clear  the  time  and  manner  of  molting : 

December. — Five  specimens,  all  in  full  unworn  pelage ;  hind  feet  white,  the 
soles  fully  haired. 

January. — Seven  specimens,  all  in  full  unworn  pelage  excepting  one  (Virginia, 
January  18),  which  is  slightly  worn  on  rump;  hind  feet  of  all,  white;  soles 
haired. 

February. — Eight  specimens,  all  in  full  unworn  pelage ;  hind  feet  white  and 
soles  haired. 

March. — Fourteen  specimens;  ten  in  full  unworn  pelage;  four  adults  (Wash- 
ington, D.  C,  March  1)  show  considerable  wear  on  head,  nape,  and  rump;  one 
immature  specimen  (White  Sulphur,  West  Virginia,  March  1)  is  slightly  worn 
on  head ;  all  have  white  feet  and  hairy  soles. 

April. — Thirteen  specimens;  four  in  full  pelage  showing  little  wear;  six 
much  worn  on  head  and  nape ;  two  considerably  worn  on  entire  upperparts  and 
one  on  flanks;  all  but  one  have  white  feet  and  all  but  one  (Long  Island,  New 
York,  April  15),  hairy  soles. 

May. — Seven  specimens;  two  young  individuals  (Maryland,  May  13)  in  much- 
worn  pelage  have  brown  feet,  the  soles  hairy ;  one  young  from  Wisconsin  (June 
10)  is  similar,  but  shows  little  wear;  one  adult  (West  Virginia,  May  17)  is  in 
full  pelage,  but  has  brown  feet  and  naked  soles ;  two  adults  in  full  unworn 
pelage,  except  on  head,  have  white  feet  but  naked  soles;  one  adult  (Mount 
Graylock,  Mass.,  May  6)  is  slightly  worn  on  head,  having  white  feet  and 
hairy  soles. 

June. — Eight  specimens  ;  four  moderately  worn ;  three  in  full  unworn  pelage ; 
one  worn  on  head  only;  one  (Washington,  D.  C,  June  20)  practically  same  as 
winter  specimens  (except  feet)  ;  feet  dusky,  without  white  markings  (except 
in  one  from  Massachusetts,  June  27,  in  which  white  on  toes  is  faintly  indicated.) 

July. — Eleven  specimens;  three  young  nearly  full  grown  (July  23,  27); 
two  from  Massachusetts  (July  3,  26),  practically  full-grown  young,  show  con- 
siderable wear ;  adults  show  only  moderate  wear  on  body,  considerable  on 
tail;  one  (July  28,  Washington,  D.  C.)  has  pelage  as  full  as  in  winter,  but 
redder ;  feet  dusky ;  soles  naked. 

August. — Two  specimens;  one  (Lake  George,  New  York,  August  30)  is  some- 
what worn  on  head,  very  little  on  body;  another  (Massachusetts,  August  30) 
shows  considerable  wear  on  upperparts  and  tail ;  feet  dusky. 

September. — Eight  specimens ;  four  show  little  wear ;  two  young  are  consid- 
erably worn;  one  adult  (September  6,  Maryland)  is  much  worn,  especially 
on  nape  and  tail ;  one  (Lake  George,  New  York,  September  15)  has  practically 
completed  the  body  molt,  except  on  head  and  a  small  area  in  middle  of  hinder 
back ;  feet  dusky  and  soles  naked. 

October. — Five  specimens;  one  young  (October  21)  shows  little  wear;  one 
adult  (Sing  Sing,  N.  Y.,  October  17)  apparently  still  retains  the  old  pelage, 
which  is  little  worn  except  on  head  and  tail ;  feet  noticeably  whitened, 
but  soles  naked;  three  adults  (Massachusetts,  October  21;  Virginia,  October 
19)  are  molting  all  over  the  upperparts,  the  feet  still  dusky  and  soles  naked. 

November. — Nine  specimens ;  two  immnture  individuals  from  Maryland  are 
much  worn  and  in  process  of  molting ;  one  adult  from  Maryland  (  $ ,  November 
9)  still  retains  old  pelage  on  back,  being  much  worn  on  head  and  nape,  but 
fresh  pelage  seems  to  be  coming  in  on  sides  of  body ;  feet  drab  without  white 
markings  and  soles  naked;  one  adult  (Washington,  D.  C,  November  1)  has 
completed  the  body  molt  except  on  head  and  a  patch  on  nape;  the  toes  on  hind 


22  NORTH   AMERICAN   FAUNA.  [No.  44. 

feet  are  slightly  whitened,  and  the  hair  is  encroaching  on  the  soles ;  five  adults 
from  Connecticut  (November  26)  are  in  full  winter  pelage,  with  white  feet 
and  hairy  soles. 

Skull. — Largest  of  the  group;  superior  outline  straight  from 
nasals  to  postf rontal  region ;  braincase  moderately  depressed,  squar- 
ish in  general  outline. 

Measurements. — Average  of  1G  adults  from  District  of  Columbia 
and  adjacent  parts  of  Virginia  and  Maryland:  Total  length,  232 
(220-240);  tail  vertebra,  101  (90-110);  hind  foot,  30.4  (29-32). 
Skull:  Average  of  12  adults  from  District  of  Columbia:  Greatest 
length,  34.9  (34.1-35.6)  ;  zygomatic  breadth,  21  (20.4-22.2)  ;  mas- 
toidal breadth,  17.1  (16.5-17.7)  ;  least  interorbital  breadth,  6.9  (6.2- 
7.6)  ;  least  postorbital  breadth,  9.1  (8.6-9.6)  ;  length  of  nasals,  9.6 
(9.2-10.1)  ;  maxillary  toothrow,  6.4  (6.2-6.7). 

Type  specimen. — None  known  to  exist. 

Remarks. — This  flying  squirrel  has  an  extensive  range,  and 
although  individual  variation  within  the  subspecies  is  considerable, 
there  is  practically  no  variation  which  can  be  correlated  with  faunal 
areas.  Large  series  of  specimens  are  available  from  nearly  all  parts 
of  its  range,  and  careful  comparison  fails  to  reveal  any  appreciable 
differences  between  those  of  the  New  England  States  on  the  one 
hand  and  those  of  Minnesota,  Iowa,  Kansas,  and  Nebraska  on  the 
other.  On  the  southern  border  of  the  range,  however,  a  gradual 
darkening  of  the  color  of  the  upperparts  is  noticeable,  and  also  a 
tendency  toward  elimination  of  white  on  the  toes  in  winter  pelage. 
Specimens  from  Virginia  (the  type  region),  however,  do  not  differ 
appreciably  from  New  England  and  New  York  individuals.  Speci- 
mens from  Ealeigh,  N.  C,  and  from  Stillwell,  Okla.,  show  decided 
approach  in  characters  to  saturatus;  a  single  winter  specimen  from 
Gainesville,  Tex.,  although  approaching  texensis  in  skull  characters, 
agrees  almost  exactly  with  typical  volans  in  color  and  in  having 
white  toes.  Since  this  specimen  shows  no  approach  to  saturatus 
(which  occurs  at  Eed  Fork,  in  the  Arkansas  Valley),  it  seems  prob- 
able that  typical  volans  ranges  from  Kansas  into  western  Oklahoma, 
perhaps  as  far  as  the  Wichita  Mountains,  and  thence  into  northern 
Texas.  A  single  specimen  in  summer  pelage  from  Oklahoma  City, 
although  somewhat  indeterminate  in  characters,  seems  to  bear  out 
this  theory  of  distribution.  Additional  material  from  that  part  of 
Oklahoma  is  needed  to  settle  the  question. 

The  type  of  Sciuropterus  silus  Bangs  has  been  examined  and 
proves  to  be  an  immature  individual  of  volans,  evidentty  a  runt.  A 
considerable  series  is  now  available  from  the  type  locality  of  silus, 
and  all  are  perfectly  typical  of  volans. 

The  type  of  Pteromys  volans  nebraseensis  Swenk  has  been  exam- 
ined and  found  to  agree  essentially  with  t}7pical  specimens  of  volans 


1918.]  GLAUCOMYS  VOLANS  GROUP.  23 

from  the  vicinity  of  Washington,  D.  C.  It  is  a  rather  pronounced 
example  of  the  gray  phase,  and,  although  taken  on  November  26, 
the  hind  toes  have  not  yet  acquired  the  clear  white  which  charac- 
terizes the  winter  pelage.  The  hind  feet  are  somewhat  darker  than 
the  average  of  volans — in  this  respect  agreeing  with  saturatus.  The 
tail  measurement  of  the  type  (115  mm.)  is  about  14  mm.  more  than 
the  average,  but  only  5  mm.  more  than  the  extremes  of  volans.  In 
the  dried  skin,  however,  the  tail  does  not  measure  any  longer  than 
in  dozens  of  typical  specimens.  The  skull  and  hind  feet  measure 
exactly  as  in  typical  examples. 
Specimens  examined. — Total  number,  356,  as  follows: 

Connecticut:   East  Hartford,  5;1  Liberty  Hill,  8.2 

District  of  Columbia:  Cleveland  Park,  13;  Washington,  23. 

Illinois:   Belleville,  1;  Chicago,  2;  Golconda,  1  ;3  Olive  Branch,  l;s  Olney, 

2 ;  Parkersburg,  1 ;  Warsaw,  8 ;  Willow  Springs,  1.* 
Indiana :  Denver,  2  ;2' 4  Kankakee,  1  ;3  La  Porte,  1  ;3  Wheatland,  1. 
Iowa :   Hillsboro,  2 ;  Iowa  City,  1  ;5  Knoxville,  3 ;  Thayer,  2.6 
Kansas:   Douglas    County,    5;6    Lawrence,    ll;5    Topeka,    4;8    Woodson 

County,  1.' 
Kentucky:   Eubanks,  3. 
Maryland:  Bacon  Hill,  3;8  Branchville,  1;  Capitol  View,  1;  Garrett  Park, 

1 ;   Kensington,  2 ;   Laurel,  3 ;   Marlboro,  1 ;   Marshall   Hall,  2 ;   Oxon 

Hill,  1 ;  Plummer  Island,  2 ;  Prince  Georges  County,  3 ;  Silver  Spring,  2. 
Massachusetts:   Ipswich,  1;  Mount  Graylock,  1;  Wareham,  4;2  Waverly, 

2  ;2  Wilmington,  2 ;  Woburn,  1. 
Michigan:  Ann  Arbor,  4;4  Brooklyn,  1  ;4  Dexter,  1;*  Iosca,  l;4  Ithaca,  2;4 

Kavanagh  Lake,  3;4  Lodi  Township,  Washtenaw  County,  l;4  Napoleon, 

l;4  Portage  Lake,  2;4  Sand  Point,  10  ;4  Ypsilanti,  1* 
Minnesota:   Aitkin,  1  ;3  Bridgman,   2;    Elk  River,   19;    Fort  Snelling,  3; 

Steele  County,  2.8 
Missouri :   Independence,  1 ;  Stotesbury,  I.2 
New  Hampshire:   Hancock,  2.2 
New    Jersey:   Cape   May   County,   1 ;°   Culvers   Gap,    Sussex   County,   l;8 

Ellenville,  1 ;  Haddonfield,  5  ;8  Lake  Hopatcong,  4  ;8  Seaville,  2.8 
New  York:    Ardsley,  3;3,8  Geneva,  1;   Hastings,   If  Jamaica,   1;    Lake 

George,  5;   Lake  Grove,  2;   Locust  Grove,  5;   Miller  Place,  5;*  Sing 

Sing,  7 ;  Suffolk  County,  l.« 
North  Carolina :   Apex,  1  ;3  Raleigh,  20.10 
Ohio:  Fort  Ancient,  l;9  Madisonville,  3;  Ravenna,  1." 
Oklahoma :  Oklahoma  City,  1 ;  Stilwell,  7. 
Ontario :   Kingsville,  1  ;u  London,  7.11 

1  Three  in  collection  Acad.  Nat.  Scl.  Philadelphia. 

2  Collection  Mus.  Comp.  Zool. 
Collection  Field  Mus.  Nat.  Hist. 
*  Collection  Univ.  Michigan. 

6  Collection  D.  Stoner,  Iowa  City,  Iowa. 

a  Collsction  Kansas  Univ.  Mus. 

T  Collection  Colorado  Mus.  Nat.  Hist. 

8  Collection  Acad.  Nat.  Sci.  Philadelphia. 

8  Collection  Amer.  Mus.  Nat.  Hist. 

10  Nine  in  collection  Amer.  Mus.  Nat.  Hist. 

11  Collection  W.  E.  Saunders,  London,  Out. 


24  NORTH   AMERICAN   FAUNA.  [No.  44. 

Pennsylvania:  Carlisle,  1;  Chester  County,  2;  Drury  Run,  Clinton 
County,  1 ;  Huntingdon  Furnace,  1  ;*  Leasuresville,  3  ;l  Moon  Township, 
Allegheny  County,  1  ;2  Pine  Grove  Furnace,  1." 

Tennessee:   Nashville,  l.3 

Texas:   Gainesville,  2. 

Vermont:   Castleton,  l;4  Rutland,  2.4 

Virginia:  Cherrydale,  1;  Dranesville,  1;  Dunn  Loring,  4;  Falls  Church,  8; 
Fort  Myer,  1 ;  Henrico  County,  1 ;  Maywood,  1 ;  Mount  Rodgers,  Gray- 
son County,  2 ;  Mount  Vernon,  7 ;  Nelson  County.  2  ;3  Suffolk,  1 ;  War- 
wick County,  2. 

West  Virginia:  Travellers  Repose,  2;  White  Sulphur  Springs,  13." 

Wisconsin :  Beaver  Dam,  2  ;"■ '  Cassville,  1  ;3  Delavan,  1 ;  Lake  Koshkonong, 
l;7  Racine,  1.  / 

GLAUCOMYS  VOLANS  SATURATUS  Howell. 

Southeastern  Flying  Squirrel. 

[PI.  II,  fig.  1 ;  PI.  IV,  fig.  1 ;  PI.  VI,  fig.  3.] 

Olaucomys  volans  saturatus  Howell,  Proc.  Biol.  Soc.  Washington,  XXVIII,  p. 
110,  May  27,  1915. 

Type  locality. — Dothan,  Alabama. 

Distribution. — Southeastern  United  States  (excepting  peninsular 
Florida  and  the  coast  region  of  Georgia)  from  South  Carolina  and 
western  North  Carolina  west  to  central  Oklahoma  and  north  in  the 
Mississippi  Valley  to  southwestern  Kentucky. 

Characters. — Similar  in  size  and  skull  characters  to  vola?is,  but 
upperparts  darker  at  all  seasons;  toes  not  conspicuously  whitened  in 
winter.  Compared  with  querceti:  Colors  averaging  darker,  face 
grayer  (less  buffy),  and  hind  feet  grayer  (less  brownish);  soles 
haired  in  winter. 

Color. — Winter  pelage:  Upperparts  pale  snuff  brown  or  dark  sayal 
brown,  varying  to  hair-brown ;  sides  of  face  smoke  gray,  rarely  with 
a  tinge  of  Duff;  upper  surface  of  flying  membrane  dark  clove  brown 
or  blackish  brown ;  hind  feet  hair-brown,  the  toes  washed  with  buff ; 
fore  feet  grayish  buff ;  tail  above,  snuff  brown,  shaded  with  bister  or 
dark  hair-brown  (rarely  solid  hair-brown)  ;  beneath,  pinkish  cinna- 
mon or  vinaceous-cinnamon  (rarely  light  hair-brown,  washed  with 
pinkish  cinnamon) ;  underparts  creamy  white,  the  under  surface  of 
the  membranes  edged  with  light  pinkish  cinnamon  or  light  vinaceous 
cinnamon.     Summer  pelage:  Essentially  as  in  winter,  but  hind  feet 


1  Collection  Acad.  Nat.  Sci.  Philadelphia. 

2  Collection  Carnegie  Mus. 

3  Collection  Mus.  Comp.  Zool. 

*  Collection  G.  L.  Kirk,  Rutland,  Vt. 

8  Ten  in  collection  Mus.  Comp.  Zool. ;  two  in  Field  Mus.  Nat.  Hist. 

6  Collection  Mus.  Vert.  Zool.,  Univ.  California. 

'Collection  E.  R.  Warren,  Colorado  Springs,  Colo. 


1918.]  GLAUCOMYS  VOLANS   GROUP.  25 

fuscous.  Variation  (March  specimens  from  Milton,  Florida)  : 
Upperparts  rich  pecan  brown. 

Skull. — Closely  similar  to  that  of  volans,  but  averaging  smaller; 
practically  same  size  as  that  of  querceti,  but  bullae  averaging  smaller. 

Measurements. — Average  of  12  adults  from  southern  Alabama:1 
Total  length,  226  (211-244)  ;  tail  vertebra?,  100  (84-109) ;  hind  foot, 
30  (28-33).  Skull:  Average  of  10  adults  from  southern  Alabama:2 
Greatest  length,  34  (32.8-34.9)  ;  zygomatic  breadth,  20.3  (19.9-21.1)  ; 
mastoidal  breadth,  17  (16.6-17.5)  ;  least  interorbital  breadth,  7  (6.7- 
7.8) ;  least  postorbital  breadth,  8.9  (8.3-10)  ;  length  of  nasals,  9.4 
(8.8-10)  ;  maxillary  tooth  row,  6.4  (6.2-6.8). 

Type  specimen. — No.  178366,  U.  S.  Nat.  Mus.  (Biological  Survey 
collection) ;  $  ad.,  skin  and  skull;  collected  March  13,  1912,  by  A.  H. 
Howell. 

Remarks. — This  race  is  the  darkest  form  in  the  volans  group,  differ- 
ing in  this  respect  both  from  the  Florida  form  (querceti)  and  from 
the  Texas  form  (texensis).  It  is  much  darker  than  volans,  both  in 
winter  and  in  summer.  Its  range  is  chiefly  in  the  Lower  Austral 
Zone,  but  extends  into  the  Upper  Austral  in  the  mountains  of  eastern 
Tennessee  and  western  North  Carolina.  Intergradation  with  volans 
is  shown  by  a  series  from  Magnetic  City,  foot  of  Roan  Mountain, 
North  Carolina,  and  with  querceti  by  a  series  from  Pinetucky,  Ga. 
The  Pinetucky  specimens  are  typical  of  saturatus  in  skull  characters, 
but  average  paler  in  color.  By  reason  of  lack  of  material  from 
Louisiana,  the  relationship  of  this  race  to  texensis  and  its  western 
limits  in  that  State  can  not  be  defined.  Specimens  from  Eed  Fork, 
Okla.,  are  typical  but  in  the  Boston  Mountains  intergradation  with 
volans  occurs. 

Two  specimens  in  full  winter  pelage  from  Milton,  Fla.,  differ  from 
typical  saturatus  in  having  the  upperparts  pecan  brown  instead  of 
snuff  brown.  These  at  first  were  thought  to  represent  an  undescribed 
race,  but  the  recent  acquisition  of  3  adult  specimens  from  Muscogee, 
Fla.,  only  a  few  miles  from  Milton,  which  do  not  differ  in  any  way 
from  typical  saturatus,  leads  to  the  conclusion  that  the  Milton  speci- 
mens represent  a  color  phase  of  this  race. 

Specimens  examined. — Total  number,  89,  as  follows : 

Alabama:  Ardell,  4;  Autaugaville,  10;  Carlton,  2;  Dothan,  6;  Greensboro, 
2;  Mobile,  4;  Perdido  River,  1;  Sand  Mountain,  near  Carpenter,  2; 
York,  1. 

Arkansas:   Delight,  2. 

Florida:   Milton,  2;  Muscogee,  2. 

Georgia :  Columbus,  1 ;  Okef enokee  Swamp,  2 ;  Pinetucky,  13  ;  Reidsville,  2 ; 
Young  Harris,  1. 

Kentucky:   Hickman,  1. 

1  Dothan,  Autaugaville,  Carlton,  and  York. 

2  Dothan,  Autaugaville,  Carlton,  and  Greensboro. 


26  NORTH  AMERICAN   FAUNA.  I  No.  44. 

Louisiana:   Mer  Rouge,  2;  Powhatan  Plantation,  near  Gibson,  l.1 

Mississippi:   Columbus,  1;  Washington,  1. 

North  Carolina:   Cranberry,  1;  Magnetic  City,  foot  of  Roan  Mountain,  9. 

Oklahoma:   Red  Fork,  4. 

South  Carolina :   Calhoun  Falls,  4  ;2  Cleora,  1 ;  Greenville,  1 ;  Plantersville, 

3 ;  Santee,  1. 
Tennessee:   Watauga  Valley,  2. 

GLAUCOMYS  VOLANS  QUERCETI  (Bangs). 

Florida  Flying  Squirrel. 

[PI.  II,  fig.  9  ;  PI.  JV,  fig.  9 ;  PI.  VI,  fig.  2.] 

Sciuropterus  volans  querceti  Bangs,  Proc.  Biol.  Soc,  Washington,  X,  p.  166, 

Dec.  28,  1896. 
[Sciuropterus  rolucella]  querceti  Trouessart,  Cat.  Mamm.,  I,  p.  402,  1897. 

Type  locality. — Citronelle,  Florida. 

Distribution. — Peninsular  Florida  (south  at  least  to  Fort  Myers) 
and  the  coast  region  of  Georgia. 

Characters. — Very  similar  in  size  and  color  to  volans,  but  upper- 
parts  averaging  slightly  darker;  toes  without  conspicuous  white 
markings  in  winter;  soles  partly  naked  at  all  seasons;  skull  with 
larger  audital  bullse. 

Color. — Winter  pelage:  Upperparts  sayal  brown  to  yellowish 
wood  brown,  shaded  on  sides  and  fore  legs  with  pinkish  buff  or  light 
pinkish  cinnamon;  upper  surface  of  flying  membrane  blackish 
brown;  ears  fuscous;  sides  of  face  smoke  gray,  often  clouded  with 
fuscous  or  buff;  hind  feet  pale  fuscous,  the  toes  grayish  buff;  front 
feet  whitish  buff;  tail  above,  snuff  brown  or  verona  brown,  shading 
(when  worn)  to  dark  hair-brown ;  beneath,  pinkish  buff  or  pinkish 
cinnamon;  underparts  creamy  white,  often  irregularly  shaded  with 
pinkish  buff  or  pinkish  cinnamon,  particularly  along  edges  of  flying 
membranes.  Summer  pelage:  Essentially  the  same  as  in  winter,  but 
averaging  slightly  browner  (less  drab)  above. 

Skull. — Closely  similar  to  that  of  volans,  but  averaging  slightly 
smaller,  with  larger  audital  bullae. 

Measurements. — Average  of  8  adults  from  Florida : 3  Total  length, 
229  (221-253);  tail  vertebras,  94  (81-115);  hind  foot,  31  (30-32). 
Skull:  Average  of  13  adults  from  Florida:4  Greatest  length,  34.3 
(33.2-35.3)  ;  zygomatic  breadth,  20.5  (19.1-21.3) ;  mastoidal  breadth, 
17.4  (16.8-18.4);  least  interorbital  breadth,  7  (6.5-7.3);  least  post- 
orbital  breadth,  8.5  (8.2-8.9)  ;  length  of  nasals,  9.4  (8.7-10) ;  maxil- 
lary toothrow,  6.4  (5.9-7.1). 

1  Collection  Mus.  Comp.  Zool. 

2  Collection  Field  Mus.  Nat.  Hist. 

3  Citronelle  and  Enterprise. 

4  Citronelle,  Enterprise,  Lake  Harney,  and  New  Berlin. 


1918.]  GLAUCOMYS  VOLANS  GROUP.  27 

Type  specimen. — No.  2451,  Mus.  Comp.  Zool.  (formerly  same  num- 
ber, collection  E.  A.  and  O.  Bangs)  ;  2  ad.,  skin  and  skull ;  collected 
September  17,  1894,  by  F.  L.  Small. 

Remarks. — The  Florida  flying  squirrel  resembles  volans  very 
closely  in  color,  but  is  easily  distinguished  in  winter  pelage  b}T  the 
absence  of  white  markings  on  the  toes  and  by  naked  soles.  Compared 
with  saturatus  it  averages  considerabty  paler.  The  large  size  of  the 
audital  bullae  seems  to  be  only  an  average  character,  for  while  pro- 
nounced in  some  specimens,  others  do  not  differ  in  this  respect  from 
typical  volans.  The  type  specimen  has  larger  bulla?  than  have  two 
topotypes  and,  indeed,  can  scarcely  be  matched  in  this  character  by 
any  of  the  Florida  specimens  examined.  Intergradation  with  satu- 
ratus takes  place  in  southeastern  Georgia.  Specimens  from  St. 
Marys,  Ga.,  agree  in  color  with  querceti,  but  have  smaller  bullrc; 
some  individuals  in  the  series  have  the  hind  toes  marked  with  buffy 
white,  but  less  extensively  than  in  volans.  Considerably  more  mate- 
rial from  Florida  and  Georgia  is  needed  to  show  the  average  char- 
acters and  exact  distribution  of  this  race. 

Specimens  examined. — Total  number,  67,  as  follows : 

Florida :  Citronelle,  3  ;*  Enterprise,  10  ;2  Fort  Myers,  1  ;*  Gainesville,  2  ;s 
Hernando  County,  2;  Lake  Harney,  2;  Nassau  County,  2  ;3  New  Berlin, 
13  ;*  Ocala,  1 ;  Shell  Hammock,  2 ;  Tarpon  Springs,  9." 

Georgia :   Mcintosh  County,  1  ;x  Montgomery,  4  j1  St.  Marys,  15.1- 3- B> ' 

GLAUCOMYS  VOLANS  TEXENSIS  Howell. 

Texas  Flying  Squirrel. 

[PI.  II,  fig.  5  ;  PI.  IV,  fig.  5 ;  PI.  VI,  fig.  4.] 

Olaucomys  volans  texensis  Howell,  Proc.  Biol.  Soc.  Washington,  XXVIII,  p.  110, 
May  27,  1915. 

Type  locality. — Sour  Lake,  Texas  (7  miles  northeast). 

Distribution. — Eastern  Texas,  west  to  Aledo,  Gurley,  Elgin,  and 
Cuero ;  eastern  limits  of  range  unknown. 

Characters. — Similar  in  color  and  size  to  volans;  upperparts 
slightly  more  ochraceous;  toes  without  conspicuous  white  markings; 
skull  decidedly  shorter  and  broader.  Compared  with  querceti:  Sim- 
ilar in  color,  but  skull  shorter,  with  smaller  bullae.  Compared  with 
saturatus :  Colors  much  paler ;  skull  shorter  and  broader. 

Color. — Winter  pelage  (March)  :  Upperparts  yellowish  wood 
brown  to  drab,  shaded  on  sides  and  fore  legs  with  pinkish  buff;  upper 
surface  of  flying  membrane  fuscous-black;  ears  fuscous;  front  feet 

1  Collection  Mus.  Comp.   Zool. 

2  Three  in  collection  Anier.  Mus.  Nat.  nis.  ;  two  in  Field  Mus.  Nat.  Hist. 
8  Collection  Amer.  Mus.  Nat.  Hist. 

4  Four  in  collection  Amer.  Mus.  Nat.  Hist. ;  three  in  Acad.  Nat.  Scl.  Philadelphia. 

6  Collection  Field  Mus.  Nat.  Hist. 

6  Collection  Mus.  Vert.  Zool.,  Univ.  California. 


28  X0K1H  AMIKkWX  FAUNA.  tNo.  44. 

buffy  white:  hind  foot  hair-brown  or  drab,  the  toes  washed  with 

bully  white;  tail  above,  fuscous,  shaded  with  snuff  brown:  beneath, 
light  pinkish  cinnamon;  underparts  creamy  white.  Summer  pelage: 
A  specimen  from  Colmesniel,  Tex.  (July  BO),  agrees  essentially  with 

the  grayest  April  specimen  from  Sour  Lake:  a  much  worn  specimen 
from  Tarkington  (November  23)  is  slightly  more  ochraceous  than 

spring  specimens. 

Skull. — Similar  to  that  of  safuratus.  but  shorter  and  relatively 
broader:  bra  incase  more  rounded,  deeper,  and  more  abruptly  de- 
pressed posteriorly.  Differs  from  querceti  in  the  same  characters 
and  also  in  having  smaller  audita]  bullae. 

M  ensure  ?n<nte. — Average  of  0  topotypes  (adult)  :  Total  length. 
229  (219-236) :  tail  vertebrae,  104  (97-111) :  hind  foot,  30.8  (30-31). 
Skull:  Average  of  5  topotypes  (adult)  :  Greatest  length.  33.5  (32.7- 
S4)  :  zygomatic  breadth.  -JO. 9  (20.6-21.1)  :  mastoidal  breadth,  17.2 
(17.1-17.4)  :  least  interorbital  breadth.  7.2  (6.9-7.3)  :  least  postorbi- 
tal  breadth.  8.6  (8.2-S.9)  :  length  of  nasals.  9.4  (9.2-9.8)  :  maxillary 
toothrow,  0.3  (0.1-6.4). 

Type  specimen. — Xo.  136400,  U.  S.  Xat.  Mus.  (Biological  Survey 
collection)  :  3  ad.,  skin  and  skull:  collected  March  15.  1905.  by  J.  H. 
Gaut. 

Remarks. — The  Texas  flying  squirrel  closely  resembles  both  volant 
and  quereeti  in  color,  but  differs  from  them  in  skull  characters.  It  is 
much  paler  than  ioturatus,  which  occupies  the  eastern  Gulf  States 
from  Louisiana  to  Georgia.  The  material  at  hand  is  too  scanty  to 
show  clearly  the  relationships  and  exact  range  of  this  form.  Inter- 
gradation  with  oolans  is  indicated  by  an  intermediate  specimen  from 
Gainesville.  Tex.  (referred  to  colons),  and  doubtless  intergradation 
takes  place  also  with  satiwatus  where  their  ranges  meet. 

>    eclmens  txam'neJ. — Total  number.  10.  as  follows: 

Texas:  Colmesniel.  1 :  Sour  Lake.  6  :  Tarkington.  1 :  Texarkana.  1 :  Troup,  1. 

GLAUCOMYS  VOLANS  GOLDMANI  (Nelson). 

Mexican  Flyixg  Squirrel, 

[PI.  II,  fig.  6;  PI.  IV,  fig.  6.] 

Sciuroptcrus  vohms  goidmani  Nelson,  Proo.  Biol.   Soc.  Washington,  XVII,  p. 
14S,  Oct.  6,  1901 

Type  locality. — Teopisca.  Chiapas.  Mexico   (20  miles  southeast). 

Distribution. — Known  only  from  type  locality — probably  the  high- 
lands of  Chiapas  and  Guatemala. 

Characters* — Closely  similar  in  color  to  saturatus.  but  top  and 
sides  of  nose  whitish,  sides  of  face  marked  with  fuscous,  and  upper 
surface  of  flying  membrane  more  brownish:  soles  naked:  skull  essen- 
tiallv  like  that  of  solans. 


391%.]  GLAUCOMVS    SABKINUS    GROUP.  29 

Color. — Spring  pelage  (April  specimen,  worn  on  nape  and  tail 
only):  Upperparte   dark  sayal  brown:   sides   of   face   smoke   gray. 

strongly   varied   with   fuscous;   eye-ring  fnscoa  flesh    color, 

shaded  with  fuscous:  top  and  sides  of  nose  creamy  white;  upper 
surface  of  flying  membrane  clove  brown:  hind  feet  fuscous,  the  toes 
faintly  shaded  with  buff;  front  feet  buffy  white;  tail,  above,  snuff 
brown;  beneath,  pinkish  cinnamon  or  pale  cinnamon  buff;  under- 
parts  creamy  white,  irregularly  washed  with  pinkish  cinnamon,  most 
strongly  along  edges  of  membranes. 

Skull. — Essentially  the  same  as  that  of  uolans — slightly  larger  than 
that  of  saturatne. 

Measurement*. — Type  and  topotype :  Total  length.  237,  238 ;  tail 
vertebrae.  112. 107:  hind  foot.  30.  31.5.  SkuU:  Greatest  length.  35.1, 
34.9:  zygomatic  breadth.  21.3.  21/.*:  mastoidal  breadth.  17.8.  17.5: 
least  interorbital  breadth,  8.7,  9.2;  least  postorbital  breadth,  8.7,  9.2; 
length  of  nasals,  10.3.  0.0:  maxillary  toothrow,  6.4,  0.3. 

Type  specimen. — No.  132833.  U.  S.  Nat.  Bins.  I  Biological  Survey 
collection)  :  $  ad.,  skin  and  skull:  collected  April  8.  1004,  by  E.  A. 
Goldman. 

Remarks. — This  race,  although  widely  separated  geographically 
from  the  United  States  forms  of  the  group,  bears  so  close  a  resem- 
blance to  saiuratus  that  a  subspecific  designation  seems  best  to  ex- 
press its  relationship.  From  no  point  between  southern  Texas  and 
Chiapas  are  specimens  of  the  genus  known,  and  Nelson  and  Goldman 
in  their  travels  in  Mexico  heard  of  flying  squirrels  in  only  a  few  local- 
ities. The  animals  were  reported  to  occur  in  the  Pinal  de  Amoles 
range  of  mountains  in  eastern  Queretaro,  but  whether  these  are  refer- 
able to  this  race  is  unknown.  Tomes,  under  the  name  Pteromya 
volucella.  records  a  specimen  from  Duenas.  Guatemala,  which  un- 
doubtedly is  referable  to  the  present  form.1 

Specimen*  examined. — Two,  from  type  locality. 

Glaucomys  sabrinus  Group. 

GLAUCOMYS  SABRINUS    (Shaw). 

[Synonymy  under  subspecie.-.] 

General  characters . — Size  medium  to  large  (hind  foot  34  mm.  or 
more)  ;  hairs  on  underparts  plumbeous  at  roots. 

Cranixd  characters. — Skull  medium  to  large  (greatest  length  36 
mm.  or  more)  ;  rostrum  usually  rather  long  (except  in  macrotis)  ; 
audital  bullae  moderately  or  in  some  races  considerably  inflated. 

Color. — TTpperparts  cinnamon,  pinkish  cinnamon,  vinaceous-cinna- 
mon,  vinaceous-fawn,  cinnamon-buff,  avellaneous,  drab,  cinnamon- 
drab,  wood  brown,  sayal  brown,  snuff  brown,  mikado  brown,  or  pecan 

1  Tomes,  Robert  F.,  Proc.  Zool.  Soc.  London,  1861,  p.  281. 


30 


NORTH   AMERICAN    FAUNA. 


[No.  44. 


brown;  sides  (and  in  some  races  front)  of  face  smoke  gray  or  pale 
gray,  often  washed  with  buff,  cinnamon,  or  fuscous;  eye-ring  fuscous; 
borders  of  flying  membrane  above,  clove  brown,  fuscous,  or  blackish 


Fig. 


3. — Map  showing  the  distribution  of  the  subspecies  of  Glaucomys  sabrinus   (except 
G.  s.  bull  at  us,  see  fig.  4,  p.  51).  based  on  specimens  examined. 


brown ;  forefeet  hair-brown,  wood  brown,  drab,  or  mouse  gray ;  hind 
feet  hair-brown,  fuscous,  clove  brown,  wood  brown,  snuff  brown,  or 
mouse  gray,  the  soles  buffy  white,  yellowish  white,  creamy  white, 


1918.]  GLAUCOMYS  SABRINUS  GROUP.  31 

drab,  mouse  gray,  cinnamon-buff,  colonial  buff,  olive-ochre,  olive- 
buff,  or  oil  yellow;  tail  above,  cinnamon,  hair-brown,  wood  brown, 
sayal  brown,  snuff  brown,  clove  brown,  fuscous,  or  fuscous-black, 
usually  darkest  at  the  tip ;  tail  beneath,  pinkish  cinnamon,  vinaceous- 
cinnamon,  pinkish-,  ochraceous-,  cinnamon-,  colonial-,  cream-,  or 
olive-buff,  drab,  drab-gray,  wood  brown,  avellaneous,  clove  brown,  or 
fuscous-black,  frequently  edged  with  hair-brown  or  fuscous;  under- 
pays white,  creamy  white,  or  buffy  white,  more  or  less  washed  or 
clouded  with  pinkish  cinnamon,  pinkish-,  ochraceous-,  cream-,  cinna- 
mon-, colonial-,  or  cartridge-buff,  straw  yellow,  greenish  yellow, 
Naples  yellow,  wood  brown,  or  avellaneous. 

Geographic  distribution. — Wooded  parts  of  Alaska,  Canada, 
northeastern  United  States,  and  mountains  of  western  United  States, 
from  the  Yukon  Valley,  Cook  Inlet,  Upper  Mackenzie  River,  Fort 
Anderson,  Great  Slave  Lake,  northern  Manitoba,  northern  Quebec 
(Hamilton  River),  and  Labrador  (Makkovik),  south  to  northern 
Massachusetts,  southern  New  York  (Catskill  Mountains),  Pennsyl- 
vania (Erie  ?),  central  Michigan  (Montcalm  County),  northern  Wis- 
consin, central  Minnesota  (Elk  River),  South  Dakota  (Black  Hills), 
Utah  (Uinta1  and  Wasatch  Mountains2),  Idaho  (Sawtooth  Moun- 
tains), and  southern  California  (Sierra  Nevada,  San  Bernardino, 
and  San  Jacinto  Mountains)   (figs.  3  and  4). 

GLAUCOMYS  SABRINUS  SABRINUS  (Shaw). 

Hudson  Bay  Flying  Squirrel. 

[PI.  II,  fig.  11 ;  PI.  IV,  fig.  11 ;  PI.  VI,  fig.  6 ;  PI.  VII,  fig.  10.] 

Sciurus  hudsonius  Gmelin,  Syst.  Nat.,  I,  p.  153,  1788  (not  Sciurus  vulgaris  hud- 

sonicus  Erxleben,  1777). 
Sciurus  sabrinus  Shaw,  Gen.  Zool.,  II,  p.  157,  1801. 
?  Pteromys  canadensis  Geoffroy,  Cat.  Mamm.  Mus.  Hist.  Nat,  Paris,  p.  170, 

1803.    Type  locality,  North  America  [probably  Quebec,  Canada.3] 

1  Allen  (Monogr.  North  Amer.  Rodentia,  p.  664,  1877)  records  a  specimen  from  Uinta 
Mountains  which  has  since  disappeared  from  the  collection  ;  this  specimen  was  taken 
September  20,  1870,  by  H.  D.  Schmidt,  near  the  head  of  Smiths  Fork,  Utah  (see  Prelim. 
Kept.  U.  S.  Geol.  Surv.  Wyoming,  etc.,  pp.  41-44,  461,  1871). 

2  F.  E.  Crandall,  of  Victor,  Idaho,  states  that  while  living  in  Emery  County,  Utah,  he 
often  saw  flying  squirrels  and  knew  of  their  being  killed  by  loggers  when  felling  trees. 
His  report  indicates  that  some  form  of  the  group  ranges  much  farther  south  than  has 
been  supposed. 

3  The  description  under  this  name  was  based  on  a  specimen  (No.  CCCLI)  in  the  Paris 
Museum,  but  as  I  am  informed  by  Dr.  A.  Menegaux  in  a  recent  letter,  this  specimen  is 
not  now  in  the  museum.  The  description  is  inadequate,  but  the  size  of  the  animal  is 
given  as  150  mm.  This  measurement  probably  refers  to  the  length  of  head  and  body, 
and  if  so  agrees  well  with  that  of  certain  specimens  of  sabrinus  from  southern  Ontario. 
It  is  too  large  for  any  member  of  the  volans  group.  The  specimen  doubtless  was  sent 
from  Canada  (probably  from  the  vicinity  of  Quebec),  as  suggested  by  the  common  name, 
"  Le  Polatouche  du  Canada,"  and  the  name  canadensis  may  thus  be  considered  a  syno- 
nym of  sabrinus  of  two  years  earlier  date. 


32  NOETH  AMERICAN   FAUNA.  {No.  44. 

Ptcromys  sabrinus  Richardson,  Zool.  Journal,  III,  p.  519,  1828 ;  Fauna  Boreali- 

Amer.,  I,  p.  193,  1829;  Audubon  and  Bachman,  Quad.' North  Amer.,  Ill,  p. 

202,  1853,  plate  CXLIII. 
Fteromys  hudsonius  Fischer,  Synop.  Mannn.,  p.  365,  1829 ;  Baird,  Mamm.  North 

Amer. :  Rept.  Expl.  and  Surv.  R.  R.  Pac,  VIII,  p.  288,  1857. 
Sciuropterus   volucella  var.    hudsonius   Allen,    Proc.   Boston    Soc.   Nat.    Hisc., 

XVI,  p.  289,  1874 ;  Allen,  Monogr.  North  Amer.  Rodentia :  Rept.  U.  S.  Geol. 

Surv.  Terr.,  XI,  p.  655,  1S77. 
Sciuropterus  volucella  sabrinus  Rhoads,  Amer.  Nat.,  XXVIII,  p.  525,  1894. 
Sciuropterus  sabrinus  Bangs,  Proc.  Biol.  Soc.  Washington,  X,  p.  162,  1896. 

Type  locality. — Mouth  of  Severn  River,  Ontario,  Canada. 

Distribution. — Interior  of  Canada,  from  Fort  Simpson  (possibly 
Fort  Anderson),  Mackenzie,  and  lower  Churchill  River,  west  side  of 
Hudson  Bay,  south  to  northern  Minnesota,  extreme  northwestern 
Wisconsin,  southern  Ontario  (vicinity  of  Lake  Nipissing) ,  and  south- 
ern Quebec  (Lake  Edward). 

Characters. — Size  medium  (for  the  group)  ;  general  tone  of  upper- 
parts  vinaceous-cinnamon ;  sides  of  face  smoke  gray;  tail  strongly 
shaded  with  fuscous;  underparts  whitish,  shaded  with  drab;  skull 
of  medium  size,  rather  long  and  narrow,  with  narrow  braincase. 

Color. — Winter  pelage:  Upperparts  deep  vinaceous-cinnamon,  the 
sides  faintly  washed  with  light  pinkish  cinnamon,  the  underfur  dark 
mouse  gray;  upper  surface  of  flying  membrane  fuscous,  the  color 
extending  over  legs  and  becoming  paler  on  feet;  toes  shaded  with 
grayish  white;  front  and  sides  of  face  smoke  gray  or  pale  smoke 
gray;  eye-ring  fuscous;  underparts  soiled  whitish,  faintly  washed 
with  pale  yellowish  and  more  or  less  strongly  shaded  with  drab  or 
hair-brown,  especially  along  sides,  the  underfur  deep  neutral  gray; 
tail,  above,  hair-brown  or  fuscous,  often  shaded  near  the  base  with 
vinaceous-cinnamon,  the  tip  usually  clear  fuscous;  beneath,  drab, 
shaded  with  pale  wood  brown  and  bordered  on  sides  and  tip  with 
hair-brown  or  fuscous ;  soles  buffy  white  on  inner  side,  drab  on  outer 
side.    /Slimmer  pelage :  Similar  to  the  winter  pelage. 

Skull. — Size  medium  (larger  than  macrotis,  smaller  than  alpinus) ; 
braincase  rather  narrow  and  elongate;  zygomata  not  widely  ex- 
panded, contracted  anteriorly;  dorsal  outline  nearly  straight  from 
nasals  to  postf rontal  region ;  braincase  moderately  depressed ;  audital 
bullae  of  medium  size. 

Measurements. — Adult  male  from  Cochrane,  Ontario:  Total 
length,  315;  tail  vertebrae,  140;  hind  foot  (dry),  40;  ear  from  notch 
(dry),  18.  Adult  female  from  Lake  Edward,  Quebec:  315;  148;  43; 
20.5.  Average  of  4  adults  from  Norway  House,  Manitoba:  Hind 
foot  (dry),  39.5  (38.5-41).  Skull:  Subadult  male  from  Cochrane, 
Ontario:  Greatest  length,  40;  zygomatic  breadth,  24;  mastoidal 
breadth,  18.1 ;  least  interorbital  breadth,  7.3 ;  least  postorbital  breadth, 
8.9;  length  of  nasals,  11.5;  maxillary  toothrow,  7.8.    Two  adults 


1918.]  GLAUCOMYS  SABRINUS  GROUP.  33 

from  vicinity  of  Great  Slave  Lake,  Mackenzie :  Greatest  length,  39.9, 
40.2;  zygomatic  breadth,  24.5,  24.4;  mastoidal  breadth,  18.8,  19.2; 
least  interorbital  breadth,  8.3,  8.2 ;  least  postorbital  breadth,  9.4,  9.2 ; 
length  of  nasals,  12,  12.8;  maxillary  toothrow,  7.8,  8.1. 

Type  specimen. — Not  noAv  known  to  exist.1 

Remarks. — The  Hudson  Bay  flying  squirrel — the  first  form  of  the 
group  to  receive  a  name — is  still  imperfectly  known,  no  specimens 
from  the  type  locality  being  as  yet  available.  A  good  series  of  skins 
from  Norway  House,  Manitoba,  doubtless  represent  the  typical  form, 
and  these  have  been  used  in  making  comparisons,  but  there  are  no 
skulls  in  any  collection  from  nearer  the  type  locality  than  Great 
Slave  Lake  on  one  side  and  Cochrane,  Ontario,  on  the  other.  Since 
these  skulls,  however,  agree  essentially  in  characters,  they  are  assumed 
to  be  typical.  Two  winter  specimens  from  Cochrane,  Ontario,  differ 
from  Norway  House  specimens  in  being  slightly  paler  and  less  vina- 
ceous  above,  having  the  hind  feet  mouse  gray  and  the  tail  dark  hair- 
brown,  with  scarcely  a  trace  di  vinaceous ;  whether  these  specimens 
represent  the  typical  race  or  an  unrecognized  form  can  not  now  be 
decided. 

The  present  race  has  a  very  wide  range  and  apparently  inter- 
grades  with  all  the  surrounding  forms — with  makhovilcensis  in  east- 
ern Quebec,  with  macrotis  in  northern  Minnesota,  northern  Wisconsin, 
northern  Michigan,  and  southern  Ontario ;  with  canescens  in  southern 
Manitoba,  with  alpinus  in  southern  Mackenzie  and  Alberta,  and  prob- 
ably with  bangsi  in  southwestern  Alberta  and  northwestern  Montana. 
Specimens  snowing  intergradation  with  alpinus  have  been  examined 
from  Fort  Providence  and  Fort  Simpson,  Mackenzie ;  and  from  Cas- 
cade Rapid,  Grand  Rapids,  Edmonton,  and  Red  Deer,  Alberta.  The 
northern  specimens  differ  from  typical  sabrinus  in  being  paler  and 
grayer ; 2  those  from  Red  Deer  are  nearly  typical  in  color,  but  have 
skulls  closely  resembling  those  of  alpinus.  Specimens  from  Trout 
Creek,  Ontario;  Hinckley  and  Itasca  County,  Minn.;  and  Gordon, 
Wis.,  are  intermediate  between  sabrinus  and  macrotis.  A  specimen 
from  Godbput,  Quebec,  agrees  with  sabrinus  in  color,  but  has  a  skull 
equaling  that  of  makkovikensis  in  size.  Specimens  from  Rat  Portage, 
Ontario,  and  Winnipeg,  Manitoba,  are  paler  than  the  typical  form, 
thus  showing  approach  to  canescens. 

A  specimen  in  worn  and  faded  pelage,  without  tail  or  skull,  from 
Fort  Liard,  Mackenzie,  is  provisionally  referred  to  this  race.  Better 
material  from  that  region  may  show  the  form  occurring  there  to  be 
alpinus. 


1The  names  hudsonius  and  sabrinus  were  based  on  Forster's  account  (Philos.  Trans., 
LXII,  p.  379,  1772)  of  a  specimen  sent  to  the  Royal  Society  from  the  mouth  of  Severn 
River. 

2  Some  of  these  intermediates  bear  a  surprisingly  close  resemblance  in  color  to  batigsi. 

14520°— 18— No.  44 3 


34  NORTH   AMERICAN   FAUNA.  [No.  44. 

Specimens  examined. — Total  number,  72,  as  follows: 
Alberta:   Calgary,  1;  Cascade  Rapid,  Athabaska  River,  1;  Didsbury,  1; 

Edmonton,  4  ;*  Grand  Rapids,  Athabaska  River,  1 ;  Gull  Lake,  1  ;a  Red 

Deer,  2.2 
Mackenzie:  Big  Island,   Great  Slave  Lake,  3;   Fort  Anderson,  1  ;8  Fort 

Liard,  l;4  Fort  Providence,  5;B  Fort  Resolution,  1  ;e  Fort  Simpson,  2; 

Hay  River,  Great  Slave  Lake,  2. 
Manitoba :   "  Hudson  Bay,"  1 ;  Norway  House,  9 ;  Oxford  House,  1 ;  Red 

River  Settlement,  3;  Selkirk  Settlement,  1;  Winnipeg,  2. 
Minnesota:   Hinckley,  2;  Itasca  County,  8.7 
Ontario:   Cochrane,  2;8- 9  Hannah  Bay  (near  Moose  Factory),  1  ;2  Matawag- 

amingue   [=Ft.  Mattagami],  5;  Moose  Factory,  l;2  Rat  Portage,  1; 

Trout  Creek,  2.8 
Quebec :  Godbout,  2 ;  Lake  Edward,  1  ;10  Tadousac,  l.6 
Saskatchewan:   Cumberland  House,  1. 
Wisconsin :   Gordon,  1  ;7  Nemakagan  River,  Burnett  County,  l.11 

GLAUCOMYS  SABRINUS  MAKKOVIKENSIS   (Sornbokgek). 

Labrador  Flying  •Squirrel. 

TPl.  II,  fig.  12 ;  PI.  IV,  fig.  12.] 

Sciuropterus  sabrinus  makkovikensis  Sornborger,  Ottawa  Nat.,  XIV,  p.  48,  June 
6,  1900. 

Type  locality. — Makkovik,  Labrador. 

Distribution. — Coast  region  of  Labrador  and  eastern  Quebec;  ex- 
act limits  unknown. 

Characters. — Similar  to  sabrinus,  but  larger  and  darker,  with 
darker  face,  tail,  and  feet. 

Color. — Adults:  Upperparts  dark  vinaceous-cinnamon  (averaging 
a  shade  darker  than  in  sabrinus)  ;  sides  of  face  smoke  gray;  upper 
surface  of  flying  membrane  clove  brown ;  feet  clove  brown  to  fuscous, 
the  soles  buffy  white  and  toes  washed  with  the  same  color;  tail 
ftiscous-black  or  clove  brown,  both  above  and  below,  moderately 
mixed  (chiefly  near  the  base)  with  pinkish  cinnamon;  underparts 
soiled  whitish,  faintly  washed  with  light  pinkish  cinnamon  or 
pinkish  buff.  Young :  Upperparts  wood  brown,  shaded  with  vinace- 
ous-cinnamon ;  tail  nearly  uniform  fuscous-black. 

Skull. — Similar  to  that  of  sabrinus,  but  slightly  larger. 

Measurements. — Average  of  5  specimens  (adult  and  subadult) 
from  Labrador:  Total   length,   309    (293-330);   tail   vertebrae,   138 

I  Collection  Victoria  Mem.  Mus. 
-  Collection  Mus.  Comp.  Zool. 

3  Fragments  of  skeleton  ;  provisionally  referred. 

4  Provisionally  referred. 

6  Three  in  collection  Victoria  Mem.  Mus. 

6  Collection  Amer.  Mus.  Nat.  Hist. 

7  Collection  Dr.  II.  V.  Ogden,  Milwaukee,  Wis. 
*  Collection  Carnegie  Museum. 

8  Collection  W.  E.  Saunders,  London,  Ont. 
10  Collection  Field.  Mus.  Nat.  Hist. 

II  Collection  Milwaukee  Pub.  Mus. 


1918.]  GLAUCOMYS  SABRINUS  GROUP.  35 

(128-146)  ;  hind  foot, 41.8  (40-45).  Skull:  Average  of  6  adults  from 
Labrador:  Greatest  length,  40.7  (40.4-41.3)  ;  zygomatic  breadth,  24.5 
(23.5-25.2)  ;  mastoidal  breadth,  18.9  (18.6-19.1)  ;  least  interorbital 
breadth,  7.8  (7.3-8.2)  ;  least  postorbital  breadth,  10  (9.7-10.3)  ;  length 
of  nasals,  11.4  (10.7-12.2) ;  maxillary  toothrow,  7.8  (7.3-8.1). 

Type  specimens. — Cotypes,  Nos.  10476,  10477,  Mus.  Comp.  Zool. 
(formerly  Nos.  1540,  1541,  collection  J.  D.  Sornborger)  ;  skins  and 
skulls,  the  skulls  imperfect ;  collected  in  1899  by  Bev.  W.  W.  Perritt. 

Remarks. — The  Labrador  flying  squirrel,  like  certain  other  species 
from  this  region,  is  larger  and  darker  than  the  form  from  the  in- 
terior of  Canada.  It  is  very  much  larger  than  macrotis,  of  New 
England.  Intergradation  with  sab?inus  apparently  takes  place  in 
the  vicinity  of  Godbout,  Quebec.  There  is  no  material  to  show  what 
form  occupies  the  interior  of  eastern  Quebec. 

Specimens  examined. — Total  number,  15,  as  follows : 

Labrador:   Cartwriglit,  1;  L'Anse  au  Loup,  l;1  Makkovik,  9;1  Paradise,  3. 
Quebec :   Northwest  River,  1. 

GLAUCOMYS  SABRINUS  MACROTIS  (Mearns). 

Mearns  Flying  Squirrel. 

[PI.  II,  fig.  10;  PI.  IV,  fig.  10  ;  PI.  VI,  fig.  5.] 

Sciiiropterus  volucella  liudsonius  Merriam,  Trans.  Linn.  Soc.  New  York,  II,  p. 

108,  1S84  (not  Sciurus  hudsonius  Gmelin). 
Sciuropterus  sabrinus  macrotis  Mearns,  Proc.  U.  S.  Nat.  Mus.,  XXI,  p.  353,  Nov. 

4,  1898. 

Type  locality. — Hunter  Mountain  (Catskills),  Greene  County,  New 
York;  altitude  3,300  feet. 

Distribution. — Nova  Scotia,  New  Brunswick,  Maine,  New  Hamp- 
shire, Vermont,  northern  Massachusetts,  Boreal  portions  of  New 
York,  northern  Pennsylvania  ( ? ) ,  southern  Ontario,  northern  part 
of  Michigan,  and  northeastern  Wisconsin ;  west  to  Elk  Kiver,  Minne- 
sota. 

Characters. — Similar  to  sabrinus,  but  smaller;  ears  averaging 
slightly  longer;  upperparts  and  hind  feet  paler  and  underparts 
whiter. 

Color. — Winter  pelage  (adult)  :  Upperparts  cinnamon  or  pinkish 
cinnamon  (rarely  light  pinkish  cinnamon),  shading  on  sides  to 
pinkish  buff ;  sides  of  face  pale  smoke  gray,  this  color  extending  back 
on  sides  of  neck  beyond  the  ears ;  top  of  nose  frequently  tinged  with 
gray;  upper  surface  of  membranes  clove  brown;  fore  feet  drab, 
clouded  with  dull  white ;  hind  feet,  above,  light  hair-brown  or  mouse 
gray ;  beneath,  soiled  whitish,  shaded  with  drab  or  buff ;  tail,  above, 
dull  cinnamon,  more  or  less  mixed  with  hair-brown  or  fuscous,  the 
general  tone  varying  from  hair-brown  to  pale  snuff  brown ;  beneath, 

1  Collection  Mus.  Comp.  Zool. 


36  NORTH   AMERICAN   FAUNA.  [No.  44. 

pinkish  cinnamon  or  light  pinkish  cinnamon;  underparts  white, 
irregularly  shaded  with  light  pinkish  cinnamon.  Summer  pelage: 
Similar  to  the  winter  pelage,  but  upperparts  usually  slightly 
darker — pale  orange-cinnamon.  Young  (October)  :  Upperparts  be- 
tween wood  brown  and  fawn  color,  otherwise  as  in  adult. 

Skull. — Similar  to  that  of  sabrinus,  but  decidedly  smaller;  brain- 
case  less  elongate,  more  nearly  spherical.  Compared  with  G.  volans 
volans:  Slightly  larger  (greatest  length  36  mm.  or  more  in  macrotis; 
35.6  mm.  or  less  in  volans) ;  fronto-parietal  region  more  elevated; 
braincase  deeper  and  less  flattened. 

Measurements. — Average  of  16  adults  from  New  York  and  New 
England:1  Total  length,  275.6  (263-290)  ;  tail  vertebrse,  126.4  (115- 
135);  hind  foot,  36  (34-38);  ear  from  notch  (dry),2  18.8  (16-20). 
Skull:  Average  of  11  adults  from  same  region:  Greatest  length, 
37.3  (36-38.7)  ;  zygomatic  breadth,  22.6  (21.9-23.5) ;  mastoidal 
breadth,  17.6  (17.2-18)  ;  least  interorbital  breadth,  6.9  (6.2-7.3)  ; 
least  postorbital  breadth,  9.2  (8.6-9.9)  ;  length  of  nasals,  10.7 
(10-11.5)  ;  maxillary  toothrow,  7  (6.4-7.7). 

Type  specimen. — No.  83152,  U.  S.  Nat.  Mus. ;  9  ad.,  skin  and  skull ; 
collected  August  31,  1896,  by  Edgar  A.  Mearns. 

Remarks. — The  Mearns  flying  squirrel  is  a  strongly  marked  form 
having  a  rather  extensive  range  in  the  Northeastern  States  and 
southern  Canada,  from  Minnesota  to  Nova  Scotia.  It  is  not  known 
from  any  point  south  of  the  Catskills  in  New  York  and  the  vicinity 
of  Erie,  Pa.,3  but  may  be  expected  to  occur  in  the  mountainous  parts 
of  Pennsylvania  and  possibly  farther  south  in  the  Alleghenies. 

Intergradation  with  sabrinus  is  shown  by  specimens  from  southern 
Ontario  (Gooderham)  and  northern  Michigan  (Porcupine  Mountains 
and  Vermilion).  Two  specimens  from  Elk  River,  Minn.,  show  ap- 
proach to  canescens,  one  of  them  being  almost  as  pale  as  that  form. 
The  skulls  are  nearly  t}7pical  of  macrotis.  Specimens  from  Maine 
have  somewhat  shorter  ears  than  those  from  New  York,  Massachu- 
setts, and  New  Hampshire. 

The  range  of  this  form  overlaps  that  of  G.  volans  volans  for  a  con- 
siderable distance  in  southern  New  England,  New  York,  Michigan, 
Wisconsin,  and  Minnesota.  Although  some  specimens  approach 
volans  rather  closely  in  size  and  resemble  it  in  the  shape  of  skull  and 
shortness  of  rostrum,  they  are  always  readily  separable  (except,  per- 
haps, in  the  young)  from  the  latter  species  by  the  plumbeous  bases 
of  the  hairs  on  the  belly.  The  upperparts,  also,  are  decidedly  more 
vinaceous  and  less  drab  than  in  volans. 

1  Hunter  Mountain  and  Peterboro,  N.  Y. ;  Wilmington,  Mass. ;  Ossipee,  N.  H.  ;  Green- 
ville, Bucksport,  and  Moosehead  Lake,  Maine. 

2  Seven  specimens  from  New  York,  Massachusetts,  and  New  Hampshire. 

3  The  specimens  labeled  as  from  Erie  are  without  further  data,  but  are  said  to  have 
been  in  the  collection  of  Geo.  B.  Sennett ;  they  may  not  have  been  collected  in  the  immedi- 
ate vicinity  of  Erie.     There  are  no  other  Pennsylvania  records. 


19187]  GLAUCOMYS  SABRINUS  GROUP.  37 

Specimens  examined. — Total  number,  78,  as  follows: 

Maine:  Bucksport,  3;1  Greenville,  8;1  Lincoln,  6;2  Moosehead  Lake,  If 
Steuben,  1 ;  Third  Mopang  Lake,  2.2 

Massachusetts:   Lunenburg,  1;  Wilmington,  3;  Winchendon,  1. 

Michigan :  Le  Roy,  1  ;3  Montcalm  County,  1  ;3  Palmer,  1 ;  Porcupine  Moun- 
tains, 3;3  Vermilion,  3.3 

Minnesota:   Elk  River,  2. 

New  Brunswick:   No  specific  locality,  1. 

New  Hampshire:   Ossipee,  4. 

New  York :  Adirondack  Mountains,  1 ;  Big  Moose  Lake,  3 ;  Hunter  Moun- 
tain, Greene  County,  1 ;  Locust  Grove,  1 ;  Peterboro,  1. 

Nova  Scotia:   Annapolis,  3  ;3,  *  Halifax,  3;  Kings  County,  5.5 

Ontario :  Gooderham,  3  ;6  Maganetewan,  1  ;2  New  Edinburgh.l  ;5  Woodstock, l.T 

Pennsylvania:   Erie  [=mountains  near?],  2.8 

Vermont:   Rutland,  2;9  Sherburne,  1.* 

Wisconsin :  Clarks  Lake,  Door  County,  1 ;  Kelley  Brook,  1  ;10  Lakewood,  2  ; 
Langlade  County,  1  ;"■  Mamie  Lake,  Vilas  County,  2. 

GLAUCOMYS  SABRINUS  CANESCENS  Howell. 

Pallid  Flying  Squikkel. 

[PI.  II,  fig.  7 ;  PI.  IV,  fig.  7  ;  PI.  VI,  fig.  11.] 

Olaucomys  sabrinus  canescens  Howell,  Proc.  Biol.  Soc.  Washington,  XXVIII, 
p.  Ill,  May  27,  1915. 

Type  locality. — Portage  la  Prairie,  Manitoba. 

Distribution. — Southern  Manitoba;  eastern  North  Dakota;  Black 
Hills,  S.  Dak.;  and  Bear  Lodge  Mountains,  Wyo. ;  exact  limits 
unknown. 

Characters. — Similar  to  microtis,  but  much  paler,  with  grayer 
head  and  larger  skull.  Compared  with  sabrinus:  Size  smaller; 
upperparts  and  feet  paler;  underparts  whiter. 

Color. — Winter  pelage:  Upperparts  light  pinkish  cinnamon,  shad- 
ing to  pale  pinkish  cinnamon  on  sides;  front  and  sides  of  face  (some- 
times nearly  the  whole  head)  pale  smoke  gray;  ears  pale  fuscous; 
eye-ring  and  upper  side  of  flying  membrane  fuscous;  feet  hair- 
brown,  the  toes  marked  with  grayish  white;  tail,  above,  dark  cin- 
namon, shaded  with  hair-brown;  beneath,  light  pinkish  cinnamon, 
edged  with  hair-brown ;  underparts  and  soles  creamy  white. 

1  Collection  Mus.   Coinp.   Zool- 

2  Collection  Acad.  Nat.  Sci.  Philadelphia. 
s  Collection  Univ.  Michigan. 

4  Collection  E.  R.  Warren,  Colorado  Springs,  Colo. 

6  Collection  Victoria  Mem.  Mus. 

8  Collection  Field  Mus.  Nat.  Hist. 

7  Collection  W.  E.  Saunders,  London,  Ont. 

8  Collection  Amer.   Mus.   Nat.   Hist. 
•Collection  G.  L.  Kirk,  Rutland,  Vt. 

10  Collection  Milwaukee  Pub.  Mus. 

u  Collection  Dr.  H.  V.  Ogden,  Milwaukee,  Wis. 


38  NORTH   AMERICAN   FAUNA.  [No.  44. 

Skull. — Similar  to  that  of  macrotis,  but  slightly  longer  and  rela- 
tively narrower,  with  longer  nasals;  decidedly  smaller  than  that  of 
sdbrinus. 

Measurements. — Average  of  3  adults:1  Total  length,  297  (290- 
306) ;  tail  vertebrae,  138  (133-146) ;  hind  foot,  37.7  (37-38).  Skull: 
Two  topotypes  (adult):  Greatest  length,  38.5,  38.8;  zygomatic 
breadth,  22.8,  22.9;  mastoidal  breadth,  17.5,  17.6;  least  interorbital 
breadth,  9.2,  9.5 ;  least  postorbital  breadth,  9.4,  9.2 ;  length  of  nasals, 
11.5,  11.2 ;  maxillary  toothrow,  7.7,  7.6. 

Type  specimen. — No.  7663,  Field  Mus.  Nat.  Hist. ;  5  subad.,  skin 
and  skull ;  collected  February  2f,  1900,  by  G.  F.  Dippie. 

Remarks. — This  is  the  palest  of  the  races  of  sdbrinus,  occupying 
chiefly  the  thinly  timbered  parts  of  southern  Manitoba  and 
eastern  North  Dakota.  The  limits  of  its  range  are  not  known,  and 
may  extend  westward  to  Saskatchewan.  Two  specimens  from  the 
Bear  Lodge  Mountains,  Wyo.,  are  referred  to  this  race,  but  a  larger 
series  may  show  characters  to  separate  the  form  in  that  region. 
These  two  specimens  show  some  approach  to  bangsi  in  skull  char- 
acters and  in  a  slightly  more  vinaceous  coloring  on  the  upperparts. 
One  of  them  has  a  wash  of  pinkish  cinnamon  on  the  underparts. 
In  a  specimen  from  Portland,  N.  Dak.,  the  upperparts  are  a  deeper 
shade  of  pinkish  cinnamon  and  the  skull  is  larger,  showing,  appar- 
ently, intergradation  with  sdbrinus.  Two  specimens  in  worn  pelage 
from  Pembina,  N.  Dak.,  are  also  considered  intermediate  between 
canescens  and  sabrinus. 

Specimens  examined. — Total  number,  10,  as  follows : 

Manitoba :  Carberry,  1 ;  Portage  la  Prairie,  2.2 

Minnesota:  Breckenridge,  1. 

North  Dakota :   Grafton,  1 ;  Pembina,  2 ;  Portland,  1. 

Wyoming:  Bear  Lodge  Mountains  (Middle  Fork  Hay  Creek),  2. 

GLAUCOMYS  SABRINUS  BANGSI   (Rhoads). 

Bangs  Flying  Squirrel. 

[PI.  II,  fig.  3;  PI.  IV,  fig.  3;  PL  VI,  fig.  10.] 

^Sciuropterus  alpinus  bangsi  Rhoads,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  June, 
1897,  p.  321. 

Type  locality. — Idaho  County,  Idaho. 

Distribution. — Mountains  of  central  Idaho,  eastern  Oregon,  south- 
western Montana,  and  western  Wyoming,  north  to  vicinity  of  Flat- 
head Lake,  Montana ;  southern  limits  unknown. 

Characters. — Similar  in  size  and  color  to  sdbrinus,  but  upperparts 
averaging  more  drab  (less  vinaceous  or  ochraceous)  and  underparts 
more  clouded  with  pinkish  cinnamon  (never  yellowish  white,  as  in 

1  Two  from  type  locality  ;  one  from  Portland,  N.  Dak. 

2  Collection  Field  Mus.  Nat.  Hist. 


1918.]  GLAUCOMYS  SABRINUS  GROUP.  39 

sabrinus) ;  feet  grayer  (less  brownish).  Compared  with  alpinus: 
Upperparts  decidedly  more  vinaceous,  tail  paler  and  much  less 
clouded  with  fuscous. 

Color. — Winter  pelage:  Upperparts  pale  wood  brown  or  avella- 
neous,  shading  in  some  specimens  to  vinaceous-cinnamon;  feet  pale 
hair-brown,  shading  to  drab-gray,  the  toes  often  grayish  white ;  tail 
above,  wood  brown,  tinged  with  cinnamon  and  more  or  less  shaded 
with  hair-brown  or  fuscous;  beneath,  light  pinkish  cinnamon, 
shaded  with  dark  hair-brown;  underparts  whitish,  strongly  washed 
with  pinkish  cinnamon  or  avellaneous.  Variation  (May  specimen 
from  Lake  Como,  Mont.)  :  Upperparts  pale  orange-cinnamon;  feet 
pale  fuscous. 

Skull. — Very  similar  to  that  of  sabrinus;  slightly  smaller  than  that 
of  alpinus,  with  smaller  bulla?. 

Measurements. — Average  of  6  (adult  and  subadult)  from  Idaho 
and  Montana : x  Total  length,  315  (304-327) ;  tail  vertebrae,  142  (137- 
148)  ;  hind  foot,  39.5  (37-41).  Skull:  Average  of  7  (adult  and  sub- 
adult)  from  same  localities:  Greatest  length,  39.9  (38.2-41.2); 
zygomatic  breadth,  24.3  (23.3-25)  ;  mastoidal  breadth,  18.7  (18.3- 
19.1)  ;  least  interorbital  breadth,  7.5  (6.9-8)  ;  least  postorbital 
breadth,  9  (8.4-9.7)  ;  length  of  nasals,  12.2  (11.1-12.5)  ;  maxillary 
toothrow,  8.1  (7.6-8.7). 

Type  specimen. — No.  6959,  Mus.  Comp.  Zool.  (formerly  same  num- 
ber, collection  E.  A.  and  O.  Bangs) ;  $  subad.,  skin  and  skull ; 
collected  March  8,  1897,  by  Harbison  and  Bargamin,  Raymond, 
Idaho. 

Remarks. — This  subspecies  is  the  smallest  of  the  Rocky  Mountain 
forms  of  sabrinus.  It  agrees  with  the  typical  race  in  skull  characters, 
but  shows  some  approach  in  color  to  alpinus,  differing  from  the  latter, 
however,  in  color  of  tail  and  skull  characters.  Intergradation  with 
sabrinus  is  indicated  by  a  specimen  from  Paola,  Mont.  With  lalipes, 
whose  range  meets  (and  possibly  overlaps)  that  of  bangsi,  there 
seems  to  be  no  intergradation.  The  specimen  from  Paola  shows  no 
approach  to  latipes,  while  one  from  Nyack,  a  few  miles  farther 
north,  is  clearly  referable  to  the  latter  form.  Specimens  from  west- 
ern Wyoming  (Pahaska,  Kendall,  Pacific  Creek)  are  slightly  larger 
than  typical  specimens,  but  do  not  differ  appreciably  in  color.  An 
August  specimen  from  Anthony,  Oreg.,  provisionally  referred  to  this 
race,  has  the  upperparts  rich  orange-cinnamon,  much  darker  than  any 
of  the  Idaho  series,  and  quite  different  from  another  August  specimen 
from  Bourne,  Oreg.,  which  is  in  the  normal  wood-brown  phase. 

Specimens  examined. — Total  number,  31,  as  follows: 
Idaho:   Idaho  County,  2;2  Ketchum,  2;2  Sawtooth  Lake,  2. 

1  Idaho  County  (type  and  topotype),  Ketchum,  and  Sawtooth  Lake,  Idaho;  Florence, 
Mont. 

s  Collection  Mus.  Comp.  Zool. 


40  NORTH   AMERICAN   FAUNA.  [No.  44. 

Montana :   Florence,  6 ;  Paola,  1 ;  Rock  Creek,  near  Lake  Como,  1. 
Oregon:   Anthony,  9;1  Bourne,  1. 

Wyoming:   Kendall    (12  miles  north,  at  7,700  feet  altitude),  2;   Pacific 
Creek,  1 ;  Pahaska,  2 ;  Wind  River  Mountains,  near  Dubois,  2. 

GLAUCOMYS  SABRINUS  ALPINUS    (Richardson). 

Richardson  Flying  Squirrel. 

[PI.  II,  fig.  4  ;  PI.  IV,  fig.  4  ;  PI.  VI,  fig.  12.] 

Pteromys  alpinus  Richardson,  Zool.  Journal,  III,  p.  519,  1828;  Audubon  and 
Bachman,  Quad.  North  Amer.,  Ill,  p.  206,  1854  ;  Baird,  Mamm.  North  Amer. : 
Rept.  Expl.  and  Surv.,  R.  R.  Pac,,  VIII,  p.  289,  1S57. 

Pteromys  sabriiius  var.  /3  alpinus  Richardson,  Fauna  Boreali-Amer.,  I,  p.  195, 
1829. 

[Sciuropterus  volucella]  alpinus  Trouessart,  Bull.  Soc.  Angers,  X,  p.  67,  1880. 

Sciuropterus  alpinus  Rhoads,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  p.  319,  1897. 

Type  locality. — Jasper  House,  Alberta.2 

Distribution. — Rocky  Mountain  region  of  Alberta  and  British 
Columbia,  from  vicinity  of  Henry  House  north  at  least  to  Peace 
River  and  Babine  Lake,  British  Columbia  (limits  of  range  unknown) . 

Characters. — Similar  to  sabrinus,  but  upperparts  much  grayer  (less 
vinaceous),  tail  darker,  and  skull  larger,  with  broader  braincase. 

Colo/r. — Winter  pelage :  Upperparts  light  drab ;  sides  of  face  pale 
smoke  gray ;  eye-ring  fuscous ;  ears  edged  with  blackish  brown ;  upper 
surface  of  flying  membrane  blackish  brown;  hind  feet  hair-brown; 
fore  feet  similar  but  slightly  paler;  soles  and  palms  soiled  whitish; 
tail  wood  brown,  much  mixed,  both  above  and  below,  with  fuscous 
or  clove  brown,  strongest  on  sides  and  tip ;  underparts  soiled  whitish, 
sometimes  irregularly  marked  (chiefly  along  median  line)  with  light 
pinkish  cinnamon. 

Skull. — Similar  to  that  of  sabrinus,  but  larger,  with  broader  brain- 
case  ;  postorbital  constriction  relatively  narrow. 

Measurements. — Two  adults  from  Henry  House,  Alberta:  Total 
length,  292,  343;  tail  vertebra?,  123,  155;  hind  foot,  41.5,  13;  average 
of  3  adults  from  Stuart  Lake,  B.  C. :  322 ;  149 ;  41.7.  Skull:  Average 
of  3  adults  from  Henry  House  and  Jasper  House,  Alberta :  Greatest 
length,  41.5  (41.2^2) ;  zygomatic  breadth,  25.4  (25.1-25.6) ;  mas- 
toidal breadth,  19.4;  least  interorbital  breadth,  8.3  (8.2-8.8);  least 
postorbital  breadth,  9.2  (9.1-9.6)  ;  length  of  nasals,  12.5  (12.4-12.6) ; 
maxillary  toothrow,  7.8  (7.5-8.1). 

1  Seven  in  collection  Amer.  Mus.  Nat.  Hist.  ;  two  in  Mus.  Vert.  Zool.,  Univ.  California. 

2  The  habitat  is  given  by  Richardson  in  the  original  description  as  "  Vallies  in  the 
Rocky  Mountains."  Later  (Fauna  Boreali-Amer.,  I,  p.  195,  1829)  he  speaks  of  specimens 
from  the  head  of  the  Elk  [=Athabaska]  River,  and  from  the  souih  branch  of  the  Mac- 
kenzie [=Liard,  specimens  probably  from  Fort  Nelson].  As  Richardson  speaks  of  Drum- 
mond  as  the  discoverer  of  the  species,  the  vicinity  of  Jasper  House,  on  the  headwaters  of 
the  Athabaska,  near  which  place  Drummond  made  extensive  collections,  may  be  considered 
the  type  locality. 


1918.]  GLAUCOMYS   SABRINUS  GROUP.  41 

Type  specimen. — None  known  to  exist. 

Remarks. — This  handsome  race,  recognized  as  distinct  by  Richard- 
son nearly  a  century  ago,  has  until  recently  been  imperfectly  known. 
Baird,  in  1857,  admitted  the  species  but  was  in  doubt  as  to  its  real 
characters.  Allen,  in  his  Monograph  of  the  Sciuridss  (1877)  placed 
it  in  synonymy.  Rhoads,  in  1897,1  restored  it  to  specific  rank,  evi- 
dently using  New  England  specimens  of  macrotis  in  comparison  and 
considering  them  typical  of  sabrinus.  Additional  material  now 
available,  both  of  typical  alpinus  and  of  sabrinus  from  various  parts 
of  its  range,  shows  conclusively  that  the  two  forms  are  subspecincally 
related.  As  already  shown  under  sabrinus  (p.  33),  specimens  from 
western  Mackenzie  and  northern  Alberta  are  intermediate  in  color 
between  alpinus  and  sabrinus  and  those  from  southern  Alberta  have 
skulls  nearly  typical  of  alpinus  but  agree  with  sabrinus  in  color.  Of 
two  specimens  from  Peace  River  Canyon,  B.  C,  one  is  typical  of 
alpinus,  while  the  other  closely  matches  sabrinm  in  color.  The  pres- 
ent form  intergrades,  also,  with  zaphceus  and  columbiensis  in  British 
Columbia,  and  possibly  with  bangsi  in  southern  Alberta.  Several 
specimens  from  Stuart  Lake,  B.  C,  are  intermediate  in  color  between 
alpinus  and  columbiensis,  while  one  specimen  is  so  dark  as  to  suggest 
intergradation  with  zaphams. 

Specimens  examined. — Total  number,  23,  as  follows: 

Alberta:  Henry  House,  3;  Jasper  House,  2;  no  specific  locality,  I.2 
British  Columbia:  Babine  Lake,  8:  Cariboo,  1; 2  Peace  River  Canyon  (near 
Hudsons  Hope),  2;  Stuart  Lake,  6.s 

GLAUCOMYS  SABRINUS  YUKONENSIS   (Osgood). 
Yukon  Flying  Squirrel. 
[PI.  II,  fig.  8;  Pi.  IV,  fig.  8.] 

Sciuropterus  yukonensis  Osgood,  North  Amer.  Fauna  No.  19,  p.  25,  Oct.  6,  1900. 

Type  locality. — Camp  Davidson,  Yukon  River,  Yukon  (near 
Alaska-Canada  boundary). 

Distribution. — Yukon  River  region,  from  vicinity  of  Mayo  Lake, 
Yukon  (head  of  Stewart  River),  to  Tanana,  Alaska;  exact  limits 
unknown. 

Characters. — Similar  to  sabrinus  in  color,  but  larger,  with  longer 
tail,  broader  hind  foot,  and  larger  skull.  Compared  with  alpinus: 
Upperparts  more  ochraceous  (less  drab) ;  tail  longer  and  more 
vinaceous. 

Color. — Upperparts  pinkish  cinnamon  to  vinaceous-cinnamon  ; 
sides  of  face  pale  smoke  gray,  sometimes  clouded  with  light  pinkish 

1  Proc.  Acad.  Nat.  Sci.  Philadelphia..  1897,  p.  319. 

2  Collection  Field  Mus.  Nat.  Hist. 

•Two  in  collection  Provincial  Mus.,  Victoria,  B.  C. ;  one  in  Acad.  Nat.  Sci.  Philadelphia. 


42  NORTH  AMERICAN   FAUNA.  [No.  44. 

cinnamon;  eye-ring  clove  brown  or  fuscous-black;  upper  surface  of 
flying  membrane  clove  brown  or  fuscous ;  legs  similar,  partially  over- 
laid with  the  color  of  the  back;  feet  wood  brown,  hair-brown,  or 
mouse  gray ;  soles  and  palms  cinnamon-buff  or  whitish,  clouded  with 
mouse  gray ;  tail  above,  sayal  brown,  shaded  with  fuscous,  becoming 
pure  fuscous  at  the  tip ;  beneath,  vinaceous-cinnamon  or  light  pinkish 
cinnamon,  edged  with  fuscous  or  hair-brown;  underparts  soiled 
whitish,  moderately  washed  with  light  pinkish  cinnamon  or  avella- 
neous. 

Skull.— Closely  similar  to  /that  of  alpinus;  braincase  slightly 
larger;  nasals  slightly  shorter. 

Measurements. — Type :  Total  length,  365 ;  tail  vertebrae,  180 ; *  hind 
foot  (dry),  41;  ?  adult  from  Tanana,  Alaska:  324;  150;  42.  Skull: 
Average  of  3  adults  from  Alaska:2  Greatest  length,  41.3  (41-41.5) ; 
zygomatic  breadth,  25.2  (25.1-25.3) ;  mastoidal  breadth,  19.6  (19.5- 
19.7)  ;  least  interorbital  breadth,  8.1  (7.8-8.3) ;  least  postorbital 
breadth,  9.5  (9.3-10.3) ;  length  of  nasals,  11.9  (11.6-12.1) ;  maxillary 
toothrow,  8.3  (7.9-8.7). 

Type  specimen. — No.$£$$# ,  U.  S.  Nat.  Mus. ;  9  ad.,  skin  and  skull ; 
collected  December  8,  1890,  by  R.  E.  Carson. 

Remarks. — This  is  one  of  the  larger  members  of  the  group,  about 
equaling  latipes  and  bullatus  in  external  measurements.  The 
hind  feet,  although  about  the  same  length  as  those  of  alpinus, 
are  considerably  broader  and  heavier,  and  the  tail  averages  longer. 
Although  the  present  form  resembles  sdbrinus  in  color,  its  skull 
is  practically  identical  with  that  of  alpinus.  No  specimens  are 
available  from  the  large  area  between  the  known  ranges  of  alpinus 
and  yukonensis,  but  in  view  of  the  close  relationship  of  these  two 
forms,  as  shown  by  the  skull  characters,  it  seems  probable  that  they 
will  be  found  to  intergrade,  and  for  that  reason  yukonensis  is  here 
treated  as  a  subspecies  of  sdbrinus.  Osgood  states 3  that  flying  squir- 
rels have  been  taken  in  the  Knik  district,  Cook  Inlet,  but  until  speci- 
mens can  be  secured  from  that  region  their  subspecific  identity  must 
remain  in  doubt. 

Specimens  examined. — Total  number,  10,  as  follows: 

Alaska:  Big  Chena  River  (65  miles  from  mouth),  1;*  Chicken,  1;  Tanana, 

3;  Toklat  River  (head),  1. 
Yukon:   Camp  Davidson   (near  Alaska-Canada  boundary),  2;   Fortymile, 

1  ;5  Mayo  Lake,  1.° 

1  In  a  topotype,  the  tail   (measured  from  dry  skin)  is  approximately  160. 

2  Camp  Davidson,  Tanana,  and  head  of  Toklat  River. 

3  Osgood,  W.  H.,  North  Amer.  Fauna  No.  21,  p.  63,  1901. 
*  Skull  only. 

8  Collection  Mus.  Vert.  Zool.,  Univ.  California. 
0  Collection  Victoria  Mem.  Mus. 


1918.]  GLAUCOMYS  SABRINUS  GROUP.  43 

GLAUCOMYS  SABRINUS  ZAPH^EUS    (Osgood). 

Axaska  Coast  Flying  Squirrel. 

[PI.  Ill,  fig.  6 ;  PI.  V,  fig.  6 ;  PI.  VI,  fig.  9.] 

Sciuropterus   alpinus  zaphcetis   Osgood,   Proc.   Biol.   Soc.   Washington,   XVIII, 
p.  133,  Apr.  18,  1905. 

Type  locality. — Helm  Bay,  Cleveland  Peninsula,  Alaska. 

Distribution. — Coast  region  of  southeastern  Alaska  and  northern 
British  Columbia ;  limits  of  range  unknown. 

Characters. — Similar  to  alpinus,  but  upperparts  browner  (less 
grayish)  and  underparts  darker.  Compared  with  oregonensis: 
Upperparts  paler;  underparts  grayer  (without  cinnamon  or  buff). 

Color. — Winter  pelage:  Upperparts  sayal  brown;  sides  of  face 
pale  smoke  gray,  often  clouded  with  fuscous-black,  the  blackish  eye- 
ring  often  pronounced;  upper  surface  of  flying  membrane  fuscous; 
feet  dark  hair-brown  or  fuscous,  the  soles  whitish  tinged  with  hair- 
brown  on  outer  margin ;  tail  above,  fuscous,  tinged  with  sayal  brown ; 
beneath,  dull  vinaceous-cinnamon,  much  clouded  with  fuscous  on  sides 
and  tip ;  underparts  soiled  whitish,  strongly  washed  with  avellaneous 
or  wood  brown,  strongest  along  sides.  Summer  pelage:  Upperparts 
yellowish  wood  brown ;  underparts  washed  with  pinkish  cinnamon. 

Skull. — Similar  to  that  of  alpinus,  but  braincase  narrower,  bullae 
slightly  smaller,  and  upper  toothrow  slightly  longer. 

Measurements. — Average  of  6  topotypes  (adult)  :  Total  length, 
30T  (292-311)  ;  tail  vertebras,  144  (133-152) ;  hind  foot  (dry  skin), 
40.7  (39-42)  ;  average  of  3  topotypes  (adult)  i1  306;  145;  40.  Skull: 
Average  of  6  topotypes  (adult  and  subadult)  :  Greatest  length,  40.1 
(39.3-11.5)  ;  zygomatic  breadth,  24.7  (23.7-26.2) ;  mastoidal  breadth, 
18.7  (18.2-19.1) ;  least  interorbital  breadth,  7.8  (7.4-8.3) ;  least  post- 
orbital  breadth,  9  (8.7-9.8)  ;  length  of  nasals,  12.5  (12.2-12.9) ;  maxil- 
lary toothrow,  7.8  (8-8.6). 

Type  specimen. — No.  136137,  U.  S.  Nat.  Mus.  (Biological  Survey 
collection);  5  ad.,  skin  and  skull;  collected  January  21,  1905,  by 
Cyrus  Catt. 

Remarks. — This  subspecies  is  intermediate  in  color  between  alpinus 
and  oregonensis,  being  nearer  the  latter  in  the  color  of  the  upper- 
parts,  but  lacking  the  deep  cinnamon-buff  on  the  underparts.  In 
skull  characters  it  is  close  to  alpinus  and  intergrades  with  it  in  the 
interior  of  British  Columbia.  The  northern  and  southern  limits  of 
its  range  are  unknown,  but  very  probably  it  intergrades  with  ore- 
gonensis in  southwestern  British  Columbia  and  possibly  with 
yukonensis  in  Yukon. 

1  Measured  in  flesh  by  Frank  Stephens. 


44  NORTH   AMERICAN    FAUNA.  [No.  44. 

Specimens  examined.— Total  number,  14,  as  follows: 

Alaska:  Bradflelcl   Canal,    l;1    Etolin    Island,    l;1    Helm    Bay,    Cleveland 

Peninsula,  10 ; 2  Tongass,  1. 
British  Columbia:   Nass  River,  1. 

GLAUCOMYS  SABRINUS  GREGONENSIS  (Bachman). 

Bachman  Flying  Squirrel. 

[PI.  Ill,  fig.  11 ;  PI.  V,  fig.  11  ;  PI.  VI,  fig.  8.] 

Pteromys  oregonensis  Bachman,  Journ.  Acad.  Nat.  Sci.  Philadelphia,  VIII, 
p.  101,  1839 ;  Audubon  and  Bachman,  Quad.  North  Araer.,  I,  p.  132,  1846. 

[Sciuropterus  volucella]  oregonentis  [sic]  Trouessart,  Bull.  Soc.  Angers,  X, 
p.  G7,  1S80. 

Sciuropterus  alpinus  oregonensis  Rhoads,  Proc.  Acad.  Nat.  Sci.  Philadelphia, 
1S97,  p.  324. 

Type  locality. — "  Pine  woods  of  the  Columbia  near  the  sea " — 
probably  near  the  present  site  of  St.  Helens,  Oregon. 

Distribution. — Coast  region  of  Oregon,  Washington,  and  southern 
British  Columbia ;  northern  and  southern  limits  unknown. 

Characters. — Similar  to  zapluvus,  but  colors  more  rufescent,  both 
above  and  below  (upperparts  rich  reddish  brown,  underparts  cinna- 
mon or  buff)  ;  sides  of  face  more  buffy  (less  grayish)  ;  skull  slightly 
smaller. 

Color. — Winter  pelage:  Upperparts  mikado  brown  or  pecan  brown ; 
sides  of  face  smoke  gray  or  grayish  white,  often  much  mixed  with 
buff  or  cinnamon ;  upper  surface  of  flying  membrane  clove  brown  or 
fuscous;  hind  feet  snuff  brown  or  pale  fuscous,  the  soles  pinkish 
buff  shaded  with  drab;  fore  feet  hair-brown  or  drab;  tail  above, 
snuff  brown,  shaded  on  sides  and  tip  with  fuscous  or  clove  brown; 
beneath,  cinnamon-buff;  underparts  heavity  washed  with  pinkish 
cinnamon  or  cinnamon-buff. 

Skull. — Similar  to  that  of  zapka:us,  but  averaging  smaller,  with  the 
zygomata  less  widely  expanded.  Closely  similar  to  that  of  fuligi- 
nosus,  but  averaging  slightly  smaller. 

Measurements. — Average  of  4  adults  from  Oregon : 3  Total  length, 
300  (294-310)  ;  tail  vertebrae,  127  (122-138)  ;  hind  foot,  38.6  (38-40). 
Skull:  Average  of  8  adults  from  Oregon : 4  Greatest  length,  39.9  (38.5- 
40.7)  ;  zygomatic  breadth,  23.7  (23-24.9)  ;  mastoidal  breadth,  18.3 
(17.6-19) ;  least  interorbital  breadth,  7.9  (7.3-8.8)  ;  least  postorbital 
breadth,  8.6  (8.1-9)  ;  length  of  nasals,  12.2  (10.9-12.9)  ;  maxillary 
toothrow,  8.2  (7.9-8.7). 

1  Collection  Mus.  Vert.  Zool.,  Univ.  California. 

2  Four  in  collection  Mus.  Vert.  Zool.,  Univ.  California. 

3  Portland,  Yaquina  Bay,  and  Gold  Beach. 

4  Portland  (3),  Latourelle  Falls,  Yaquina  Bay,  Gold-Beach,  Salem,  and  lower  Columbia 
River   (type). 


1918.]  GLAUCOMYS   SABRINUS   GROUP.  45 

Type  specimen. — No.  235,  Acad.  Nat.  Sci.  Philadelphia;  adult, 
skin  and  skull;  skin  (dismounted  and  skull  removed)  in  fair  con- 
dition, possibly  somewhat  faded ;  skull  nearly  perfect,  but  portion  of 
occiput  cut  away ;  collected  in  1839  by  J.  K.  Townsend. 

Remarks. — The  Bachman  flying  squirrel  is  easily  recognized  by 
its  rich  coloration,  rufescent  above  and  buffy  below.  It  occupies  the 
humid  northwest  coast  belt,  but  does  not  range  high  in  the  moun- 
tains. Intergradation  with  fuliginosus  is  shown  by  a  specimen  from 
Marmot,  Oreg.  (west  slope  of  Mount  Hood),  and  with  columbiensis 
by  specimen's  from  Sumas,  Chilliwack,  and  Stave  Lake,  B.  C.  These 
latter  specimens  agree  with  oregonensis  in  the  color  of  the  upper- 
parts,  but  are  much  paler  beneath.  A  specimen  from  Sulphur 
Springs,  Oreg.,  is  considerably  paler  above  than  typical  specimens. 
Specimens  from  Yaquina  Bay  and  Gold  Beach,  Oreg.,  have  smaller 
skulls  than  the  type  and  other  specimens  from  the  Columbia  River. 

Specimens  examined. — Total  number,  41,  as  follows: 

British  Columbia:  Agassiz,  l;1  Chilliwack,  1;  Harrison,  l;2  Mission,  1  ;3 
Stave  Lake,  1  ;4  Sumas,  3.4 

Oregon:  Brownsville,  1  ;B  Clackamas  River,  1;  "Columbia  River,"  3;8  Elk 
Head,  1;  Gold  Beach,  1;  Latourelle  Falls  (18  miles  east  of  Portland), 
1 ;'  Marmot,  1 ;  Portland,  4  ;  Port  Orford,  4 ;  Salem,  1  ;T  Sulphur  Springs, 
Benton  County  (near  Albany),  1;  Yaquina  Bay,  1;  no  specific  local- 
ity, 2. 

Washington :  Camp  Skagit,  1 ;  Fort  Bellingham,  1 ;  "  Lewis  River,"  1  ;4 
Olympic  Mountains  (north  Fork  Skokomish  River),  1;  Puget  Sound,  6; 
no  specific  locality,  1. 

GLAUCOMYS  SABRINUS  COLUMBIENSIS  Howell. 

Okanagan  Flying  Squikeel. 

[PI.  Ill,  fig.  7  ;  PI.  V,  fig.  7  ;  PI.  VI,  fig.  7.] 

Glaucomys  sabrinus  columbiensis  Howell,  Proc.  Biol.  Soc.  Washington,  XXVIII, 
p.  Ill,  May  27,  1915. 

Type  locality. — Okanogan,  British  Columbia. 

Distribution. — Interior  valleys  and  foothills  of  southern  British 
Columbia  and  northern  Washington,  from  Shuswap  Lake  and  Cran- 
brook,  British  Columbia,  south  to  Lake  Chelan,  Washington. 

Characters. — Decidedly  paler  than  oregonensis,  both  above  and 
below;  similar  to  Mamathensis,  but  upperparts  more  vinaceous  and 
tail  much  darker;  very  similar  to  sabrlnus,  but  soles  of  hind  feet 
often  yellow  (as  in  klamathensis) ,  and  skull  larger.     Compared  with 

1  Collection  W.  E.  Saunders,  London,  Ont. 

2  Collection  Provincial  Mus.,  Victoria,  B.  C. 

3  Collection  Victoria  Mem.  Mus. 

4  Collection  Mus.  Comp.  Zool. 

6  Collection  Leland  Stanford,  Jr.,  Univ. 
6  Collection  Acad.  Nat.  Sci.  Philadelphia. 
1  Collection  Game  Dept.  Oregon. 


46  NORTH   AMERICAN   FAUNA.  [No.  44. 

fuliginosus :  Upperparts  more  vinaceous  (less  brownish) ;  underparts 
paler;  tail  more  brownish. 

Color. — Winter  pelage:  Upperparts  between  vinaceous-cinnamon 
and  vinaceous-f awn ;  sides  of  face  smoke  gray,  shaded  with  fuscous; 
front  of  face  sometimes  marked  with  grayish  white;  upper  surface 
of  flying  membrane  clove  brown ;  feet  hair-brown  or  pale  fuscous,  the 
soles  deep  colonial  buff,  or  sometimes  grayish  white;  tail  above, 
snuff  brown,  shading  to  fuscous  on  sides  and  tip;  beneath,  pinkish 
cinnamon,  more  or  less  clouded  (often  strongly)  with  fuscous;  under- 
parts white,  tinged  with  straw  yellow  or  light  pinkish  cinnamon. 

Skull. — Closely  similar  to  that  of  alpinus,  but  averaging  slightly 
narrower,  with  flatter,  shallower  braincase;  similar  to  that  of  ore- 
gonensis,  but  slightly  larger,  with  shorter  toothrow ;  very  similar  to 
that  of  fuliginosus,  but  averaging  relatively  broader. 

Measurements. — Two  topotypes  (subadult)  :  Total  length,  313;  tail 
vertebrae,  143;  hind  foot,  42.  Skull:  Average  of  7  (adult  and  sub- 
adult)  from  southern  British  Columbia.1  Greatest  length,  41.1 
(40.3-41.6)  ;  zygomatic  breadth,  24.5  (23.6-25.2)  ;  mastoidal  breadth, 
19.1  (18.8-19.5) ;  least  interorbital  breadth,  7.8  (7.3-8.6) ;  least  post- 
orbital  breadth,  9.1  (8.4-9.7) ;  length  of  nasals,  12.6  (12-13.3)  ;  maxil- 
lary toothrow,  7.8  (7.6-8.2). 

Type  specimen. — No.  94310,  U.  S.  Nat.  Mus.  (Biological  Survey 
collection)  ;  $  subad.,  skin  and  skull ;  collected  May  9,  1898,  by  Allan 
Brooks. 

Remarks. — This  subspecies,  like  most  of  the  races  inhabiting  the 
dry  interior  valleys  of  southern  British  Columbia,  is  considerably 
paler  than  the  coast  form  (oregonensis) .  Intergradation  with  the 
latter  is  shown  by  a  series  of  specimens  from  Sumas  and  Chilliwack, 
B.  C.j  and  with  fuliginosus  by  a  series  from  mouth  of  Salmon  River, 
B.  C.  Two  specimens  in  the  latter  series  are  referable  to  the  pres- 
ent form,  although  having  somewhat  darker  underparts.  Other 
specimens,  labeled  as  from  the  same  locality  but  possibly  taken  at  a 
higher  altitude,  agree  essentially  with  fuliginosus,  to  which  race  they 
are  referred.  Intergradation  with  aljnnus  is  shown  by  specimens 
from  Stuart  Lake,  B.  C.  (referred  to  alpinus).  One  specimen  from 
Cranbrook,  B.  C,  is  considerably  grayer  above  than  the  typical  form, 
thus  suggesting  intergradation  with  latipes,  with  which  it  agrees  in 
all  other  respects.     Another  from  the  same  place  is  typical. 

Specimens  examined. — Total  number,  11,  as  follows: 

British  Columbia:  Cranbrook,  2;  Grand  Prairie  (22  miles  northwest  of 
Okanogan),  1  ;2  Okanogan,  2;  (Okanogan  Falls  (Shuttleworth  Creek), 
l;2  Penticton,  l;3  Salmon  River  (mouth),  2;3  Vernon,  1. 

Washington:   Chelan  Mountains,  1.* 

1  Okanogan,  Vernon,  Penticton,  Cranbrook,  and  mouth  of  Salmon  River. 
1  Collection  Provincial  Mus.,  Victoria,  B.  C. 
s  Collection  Victoria  Mem.  Mus. 
♦Collection  Amer.  Mus.  Nat.  Hist. 


1918.]  GLAUCOMYS  SABEINUS  GROUP.  47 

GLAUCOMYS  SABRINUS  FULIGINOSUS  (Rhoads). 

Cascade  Flying  Squibrel. 

[PL  III,  fig.  4  ;  PI.  V,  fig.  4  ;  PL  VII,  fig.  7.] 

Sciuropterus  alpinus  fuliginosus  Rhoads,  Proc.  Acad.  Nat.   Sci.  Philadelphia, 
June,  1897,  p.  321. 

Type  locality. — Cascade  Mountains,  near  Martin  Station,  Kittitas 
County,  Washington  (altitude  about  8,000  feet). 

Distribution. — Cascade  Range,  from  southern  British  Columbia 
south  through  Washington  and  Oregon  to  the  Siskiyou  Mountains, 
California. 

Characters. — Similar  to  columbiensis,  but  upperparts  more  brown- 
ish (less  vinaceous)  ;  underparts  darker,  and  tail  paler  (less  brown- 
ish). Compared  with  oregonensis:  Colors  much  less  rufescent,  espe- 
cially on  upperparts  and  underside  of  tail. 

Color. — Winter  pelage:  Upperparts  pale  sayal  brown  (between 
sayal  brown  and  wood  brown),  the  sides  sometimes  faintly  washed 
with  pinkish  cinnamon;  upper  surface  of  flying  membrane  clove 
brown;  sides  of  face  smoke  gray  or  pale  smoke  gray,  sometimes 
washed  with  light  buff  or  pale  fuscous ;  eye-ring  fuscous ;  ears  hair- 
brown;  hind  feet  dark  hair-brown  (approaching  fuscous),  the  toes 
marked  with  buffy  white,  the  soles  buffy  white  or  yellowish  white, 
edged  with  hair-brown ;  tail  above,  dark  hair-brown,  shading  to  fus- 
cous, sometimes  sparingly  mixed  with  dull  cinnamon  or  pinkish  cin- 
namon ;  beneath,  pale  ochraceous-buff  or  light  ochraceous-buff ,  edged 
on  sides  with  fuscous,  the  bases  of  hairs  pale  smoke  gray ;  underparts 
pinkish  cinnamon  or  light  pinkish  cinnamon,  the  throat  often  creamy 
white. 

Skull. — Closely  similar  to  that  of  columbiensis,  but  averaging  rela- 
tively narrower ;  similar  to  that  of  oregonensis,  but  slightly  larger. 

Measurements. — Average  of  3  topotypes  (adult)  :  Total  length, 
308  (304-317)  ;  tail  vertebrae,  144  (140-153)  ;  hind  foot,  40.7  (40-42). 
Skull:  Average  of  10  (adult  and  subadult)  from  Easton  and  Keeche- 
lus  Lake,  Wash.:  Greatest  length,  41.4  (40.2-42.5);  zygomatic 
breadth,  24.1  (23.5-24.9)  ;  mastoidal  breadth,  18.7  (17.9-19.3) ;  least 
interorbital  breadth,  8.3  (7.7-9)  ;  least  postorbital  breadth,  9.1  (8.4- 
9.6)  ;  length  of  nasals,  12.9  (12.3-13.3)  ;  maxillary  toothrow,  8.4 
(8.2-8.4) . 

Type  specimen. — No.  8058,  Acad.  Nat.  Sci.  Philadelphia  (formerly 
No.  1058,  collection  S.  N.  Rhoads)  ;  $  subad.,  skin  and  skull ;  col- 
lected in  March,  1893,  by  Allan  Rupert. 

Remarks. — This  subspecies,  occupying  the  Cascade  Range  in  Wash- 
ington and  Oregon,  is  markedly  paler  than  the  coast  form  (oregon- 
ensis).  It  intergrades  with  the  latter  on  the  western  slopes  of  the 
range,  with  klamathensis  on  the  eastern  slopes  in  Oregon,  with  colum- 


48  NORTH   AMERICAN   FAUNA.  [No.  44. 

biensis  in  southern  British  Columbia,  and  with  flaviventris  in  north- 
western California. 

Specimens  from  Vida  and  Belknap  Springs,  Oreg.,  have  browner 
backs  and  paler  bellies  than  the  typical  series,  thus  showing  approach 
to  oregonensis.     A  specimen  from  Crater  Lake,  Oreg.,  approaches 
klamathensis  in  having  paler  and  more  yellowish  underparts.    Speci- 
mens from  the  Siskiyou  Mountains,  Cal.,  are  intermediate  between 
fuliginosus  and  flaviventris,  agreeing  with  the  former  in  the  color 
of  the  upperparts  and  feet  and  in  skull  characters,  but  having  the 
underparts  paler  and  more  yellowish  than  in  typical  specimens. 
Specimens  examined. — TotaFnumber,  68,  as  follows : 
British  Columbia:   Hope,  7  ;*■ 2  Salmon  River  (mouth),  6.3 
California:   Preston  Peak,  1;  Siskiyou  Mountains,  5. 

Oregon:   Belknap  Springs,  2  ;  Crater  Lake,  1 ;  McKenzie  Bridge,  2 ;  Vida,  3.4 
Washington:  Bumping  Lake,  Yakima  County,  1;  Cowlitz  River,  1;  Easton, 
18 ;  Keechelus  Lake,  14 ;  Martin  Station,  3 ;  Potato  Hill,  15  miles  north 
of  Goldendale,  1 ;  Signal  Peak,  Yakima  Indian  Reservation,  3. 

GLAUCOMYS  SABRINUS  LATIPES  Howell. 

Broad-Footed  Flying  Squirrel. 

[PL  III,  fig.  8 ;  PL  V,  fig.  8 ;  PL  VII,  fig.  8.] 

Olaucomys  sabrinus  latipes  Howell,  Proc.  Biol.  Soc.  Washington,  XXVIII,  p.112, 
May  27,  1915. 

Type  locality. — Glacier,  British  Columbia. 

Distribution. — Selkirk  Range,  and  other  ranges  in  southeastern 
British  Columbia,  higher  mountains  of  northern  Idaho  and  north- 
western Montana ;  south  to  Mullan  and  Orofino,  Idaho. 

Characters. — Similar  to  fuliginosus  but  larger;  upperparts  aver- 
aging darker  and  grayer;  feet  larger  and  darker.  Compared  with 
alpinus  and  bang  si:  Size  larger;  upperparts  more  brownish  (less 
drab)  ;  underparts  darker. 

Color. — Winter  pelage:  Upperparts  drab,  more  or  less  mixed  with 
sayal  brown ;  sides  of  face  smoke  gray ;  ears  fuscous ;  upper  surface  of 
flying  membrane  dark  clove  brown ;  hind  feet  fuscous ;  fore  feet  dark 
hair-brown,  the  toes  shaded  with  buffy  white;  tail  above,  fuscous 
shaded  with  sayal  brown;  beneath,  pinkish  buff  or  light  ochraceous- 
buff,  more  or  less  mixed  with  fuscous;  underparts  light  ochraceous- 
buff  or  cinnamon-buff.  Summer  pelage:  Upperparts  chiefly  sayal 
brown  with  a  drab  tinge,  otherwise  as  in  winter. 

Skull. — Similar  in  general  to  that  of  fuliginosus,  but  larger  (about 
equaling  that  of  bullatus) ;  longer  and  relatively  narrower  than  that 
of  alpinus. 

1  Collection  Mus.  Coinp.  Zool. 

2  Collection  E.  R.  Warren,  Colorado  Springs,  Colo. 

3  Collection  Victoria  Mem.  Mus. 

1  Two  in  collection  Game  Dept.  Oregon. 


1918.]  GLAUCOMYS  SABKINUS   GROUP.  49 

Measurements. — Average  of  10  adults:1  Total  length,  342  (315- 
368)  ;  tail  vertebra?,  153  (145-161) ;  hind  foot,  41.4  (38-44).  Skull: 
Average  of  7  adults:1  Greatest  length,  42.9  (41.6-44.2) ;  zygomatic 
breadth,  24.5  (24-25.4)  ;  mastoidal  breadth,  19.5  (19-20)  ;  least  in- 
terorbital  breadth,  8.3  (8-8.8)  ;  least  postorbital  breadth,  8.8  (8-9.1)  ; 
length  of  nasals,  13.3  (12.5-14) ;  maxillary  toothrow,  8.3  (7.9-9). 

Type  specimen. — No.  68753,  U.  S.  Nat.  Mus.  (Biological  Survey 
collection),  5  ad.,  skin  and  skull;  collected  August  13,  1894,  by  J. 
Alden  Loring. 

Remarks. — This  subspecies  is  one  of  the  largest  of  the  American 
flying  squirrels,  nearly  equaling  bullatus  in  external  measurements 
and  size  of  skull.  Although  evidently  closely  related  to  fuliginosus 
of  the  Cascades,  intermediate  specimens  are  unknown,  but  are  likely 
to  be  found  when  further  collecting  is  done  in  British  Columbia. 
The  present  form  apparently  does  not  intergrade  either  with  alpinus 
of  the  northern  Eockies  or  with  bangsi  of  the  Bitterroot  and  Saw- 
tooth Ranges  in  Idaho  and  Montana.  From  the  latter  race, 
the  range  of  which  meets  that  of  latipes  in  northern  Montana, 
this  form  differs  in  much  larger  size  and  darker  colors.  The 
range  of  latipes  is  apparently  confined  to  the  mountains,  since  an- 
other form  {columbiensis)  occurs  in  the  Okanogan  Valley  and  at 
Cranbrook,  B.  C.  One  specimen  from  the  latter  locality  nearly 
matches  specimens  of  latipes  in  color  of  the  upperparts,  but  in  other 
characters  agrees  with  columbiensis. 

Specimens  examined. — Total  number,  23,  as  follows : 

British  Columbia:  Glacier,  1;  Sehoonover  Mountain,  near  Okanogan  (4,000 

feet  altitude),  2;2  Shuswap,  1. 
Idaho :   Coolin,  4 ;  Mullan,  7 ;  Oroflno,  1 ;  Priest  Lake,  1. 
Montana :   Nyack  1 ;  Stanton  Lake,  5. 

GLAUCOMYS  SABRINUS  OLYMPICUS  (Elliot). 

Olympic  Flying  Squirrel. 

[PI.  Ill,  flg.  10  ;  PI.  V,  fig.  10 ;  PI.  VII,  fig.  6.] 

Sciuropterus  alpinus  olympicus  Elliot,  Field  Columb.  Mus.,  Publ.  30,  Zool.  Ser., 
I,  p.  225,  Feb.  1,  1899. 

Type  locality. — Happy  Lake,  Olympic  Mountains,  Washington. 

Distribution. — Olympic  Peninsula,  Washington,  and  south  along 
the  coast  to  southern  Oregon;  occurring  in  some  localities  with 
oregonensis. 

1  From  Glacier,  B.  C. ;  Coolin,  Idaho  ;  and  Stanton  Lake  and  Nyack,  Mont. 

2  Collection  Provincial  Mus.,  Victoria,  B.  C. 

14520°— 18— No.  44 4 


50  NORTH   AMERICAN   FAUNA.  [No.  44. 

Characters. — Similar  to  fuliginosus,  but  upperparts,  face,  and  feet 
much  darker;  colors  much  less  rufescent  than  in  oregonensis,  both 
above  and  below ;  skull  similar  to  that  of  fuliginosus,  slightly  larger 
than  that  of  oregonensis. 

Color. — Worn  summer  pelage:  Upperparts  wood  brown;  sides  of 
face  smoke  gray  shaded  with  cinnamon ;  eye-ring  and  upper  surface 
of  flying  membrane  fuscous-black;  ears  pale  fuscous;  feet  fuscous, 
the  soles  shaded  with  buffy  white;  tail  above,  fuscous;  beneath,  dull 
light  pinkish  cinnamon,  edged  with  fuscous;  underparts  whitish, 
heavily  washed  with  pinkish  cinnamon.  Winter  pelage  (specimens 
from  Ocean  View,  Oreg.):1  Upperparts  snuff  brown;  tail  above, 
snuff  brown  heavily  shaded  with  fuscous  or  fuscous-black;  feet  deep 
mouse  gray ;  otherwise  as  in  summer. 

Skull. — Essentially  the  same  as  in  fuliginosus;  very  similar  to 
that  of  oregonensis,  but  averaging  slightly  larger. 

Measurements. — Average  of  4  topotypes  (subadult)  :  Total  length, 
335  (328-346) ;  tail  vertebra?,  162  (159-166) ;  hind  foot,  37.3  (35-39). 
Skull:  Greatest  length,  41  (40.3-41.6);  zygomatic  breadth,  23.9 
(23.4-24.7)  ;  mastoidal  breadth,  18.6  (18.4-19) ;  least  interorbital 
breadth,  8.2  (7.6-8.5)  ;  least  postorbital  breadth,  9.1  (8.8-9.4) ;  length 
of  nasals,  12.4  (11.2-13.2)  ;  maxillary  toothrow,  8.4  (8.3-8.5). 

Type  specimen. — No.  5902,  Field  Mus.  Nat.  Hist. ;  $  subad.,  skin 
and  skull ;  collected  September  10,  1898,  by  D.  G.  Elliot. 

Remarks. — The  Olympic  flying  squirrel  is  apparently  the  darkest 
of  the  American  forms.  It  is  most  nearly  related  to  oregonensis, 
from  which  it  differs  in  much  less  rufescent  coloration.  The  material 
at  hand  is  insufficient  to  show  clearly  whether  the  two  races  inter- 
grade.  The  occurrence  of  typical  specimens  of  both  forms  in  the 
Olympic  Mountains  and  at  Gold  Beach,  Oreg.,  argues  for  their  spe- 
cific distinctness,  but,  on  the  other  hand,  a  series  from  Ocean  View, 
Oreg.,  strongly  suggests  intergradation,  some  of  the  specimens  being 
clearly  intermediate  in  color  between  olympicus  and  oregonensis. 
In  view  of  the  close  cranial  relationship  which  olympicus  bears  to 
both  oregonensis  and  fuliginosus  it  seems  best  to  unite  it  with  them 
as  a  subspecies  of  sahrinus.  To  clear  up  the  problem  much  additional 
material  is  needed — particularly  a  good  series  of  winter  specimens 
from  the  type  region.  At  present  the  only  available  specimens  in 
winter  pelage  are  those  from  Ocean  View,  Oreg.,  which,  as  already 
explained,  may  be  shading  toward  oregonensis. 

Specimens  examined. — Total  number,  15,  as  follows: 
Oregon :   Gold  Beach,  1  ;2  Ocean  View,  9.s 
Washington :   Olympic  Mountains,  4  ;2  Seattle,  1." 

1  Possibly  not  typical — see  Remarks. 

2  Collection  Field  Mus.  Nat.  Hist. 

3  Collection  Game  Dept.  Oregon. 


1918.] 


GLAUCOMYS   SABEINUS  GROUP. 


51 


GLAUCOMYS  SABRINUS  BULLATUS  Howell. 

Sawtooth  Mountains  Flying  Sqtjierel. 

[PL  III,  fig.  12  ;  PL  V,  fig.  12  ;  PL  VII,  fig.  4.] 

Glaucomys    bicllatus   Howell,   Proc.   Biol.    Soc.   Washington,    XXVIII,   p.    113, 
May  27,  1915. 

Type  locality. — Sawtooth  [Alturas]  Lake,  Idaho. 

Distribution. — Sawtooth  Mountains,  Idaho,  north  to  Cranbrook, 
British  Columbia,  and  west  to  the  Blue  Mountains,  Oregon  (fig.  4). 

Characters. — Similar  to  klamathensis,  but  larger,  with  much  larger 
bullae;  upperparts  more  pinkish,  underparts  tinged  with  avel- 
laneous  instead  of  yellow,  tail  darker  above  and  more  vinaceous  be- 
low, and  soles  of  hind  feet  smoke  gray  instead  of  olive-ochre.  Similar 
in  color  to  bangsi,  but  upper- 
parts  decidedly  more  pinkish 
(less  vinaceous  or  drab)  ;  gray  on 
face  purer  and  more  extensive; 
size  much  larger. 

Color. — Winter  pelage:  Upper- 
parts  between  pinkish  cinnamon 
and  cinnamon-buff,  shaded  in 
some  specimens  with  wood  brown, 
becoming  paler  on  the  face ;  sides 
of  face  and  neck  pale  smoke  gray, 
this  color  extending  back  to  a 
point  slightly  behind  the  ears; 
ears  dark  hair-brown  edged  with 
fuscous  and  often  partly  clothed 
with  smoke  gray  hairs ;  upper  sur- 
face of  flying  membrane  between 
hair-brown  and  fuscous;  feet  be- 
tween mouse  gray  and  hair-brown, 

the  fore  feet  and  toes  of  hind  feet  often  marked  with  grayish  white, 
the  soles  pale  smoke  gray ;  tail  above,  pinkish  cinnamon  much  mixed 
with  fuscous,  becoming  dark  mouse  gray  at  tip ;  beneath,  dull  light 
pinkish  cinnamon;  underparts  creamy  white,  strongly  shaded  with 
avellaneous  or  light  pinkish  cinnamon,  becoming  pure  avellaneous 
on  sides.  Summer  pelage:  Hind  feet  somewhat  paler  (about  mouse 
gray)  but  otherwise  as  in  winter. 

Skull. — 'Size  large  (averaging  largest  of  the  races  of  sab  Anus) ; 
braincase  relatively  narrow  and  very  deep,  abruptly  depressed  pos- 
teriorly; interorbital  region  depressed  and  fronto-parietal  region 
markedly  elevated;  interorbital  region  broad,  the  interorbital  notch 
obsolete  or  much  reduced;  audital  bullae  very  large;  basioccipital 
relatively  narrow ;  molar  teeth  massive. 


Fig.  4. — Map  showing  the  distribution 
of  Glaucomys  sabrinus  bullatus,  based 
on  specimens  examined.  This  form 
occurs  in  some  localities  with  others 
of  the  same  species  (see  fig.  3, 
P.  30). 


52  NORTH   AMERICAN   FAUNA.  TNo.  44. 

Measurements. — Type:  Total  length,  340;  tail  vertebrae,  150;  hind 
foot,  46.  Average  of  6  adults  from  Ketchum,  Idaho:  Total  length, 
336  (327-354) ;  tail  vertebra?,  142  (135-150) ;  hind  foot,  42.5  (39-45). 
Skull:  Average  of  type  and  of  5  adults  from  Ketchum,  Idaho :  Great- 
est length,  43.3  (42.4-44.2)  ;  zygomatic  breadth,  25  (24.4-25.5) ;  mas- 
toidal breadth,  19.3  (19-20) ;  least  interorbital  breadth,  8.9  (8.1-9.6) ; 
least  postorbital  breadth,  9.4  (9-9.7)  ;  length  of  nasals,  13.6  (13.5- 
13.9) ;  maxillary  toothrow,  9.1  (8.8-9.3). 

Type  specimen. — No.  fffrii  U.  S.  Nat.  Mus.  (Biological  Survey 
collection)  :  9  ad.,  skin  and  skull;  collected  September  28,  1890,  by 
Vernon  Bailey  and  B.  H.  Dutcner. 

Remarks. — This  race  is  the  largest  and  one  of  the  handsomest  of 
the  American  flying  squirrels.  It  was  originally  believed  to  be  a  dis- 
tinct species,  since  it  occurs  at  the  same  localities  with  bangsi  in  Idaho 
and  with  latipes  in  British  Columbia,  and  does  not  intergrade  with 
them,  but  recently  acquired  material  from  eastern  Oregon  shows  in- 
tergradation  with  klamathensis.  An  adult  from  Beech  Creek,  Oreg., 
agrees  well  with  typical  bullatus  in  color,  and  its  skull  is  slightly 
larger  than  the  largest  of  the  Idaho  series;  the  braincase,  however., 
is  relatively  broader  and  flatter.  Another  adult  from  Cornucopia, 
Oreg.,  approaches  klamathensis  in  having  the  belly  and  the  soles 
tinged  with  colonial  buff.  A  specimen  from  Cranbrook,  British 
Columbia,  differs  from  the  Idaho  series  in  having  the  underparts  a 
slightly  darker  shade  of  cinnamon  and  the  tail  more  extensively 
shaded  with  fuscous. 

Specimens  examined. — Total  number,  14,  as  follows : 

British.  Columbia:   Cranbrook,  1. 

Idaho :   Ketchum,  6  ;*  Sawtooth  Lake,  2. 

Oregon :   Anthony,  1  ;2  Beech  Creek,  1 ;  Cornucopia,  3. 

GLAUCOMYS  SABRINUS  KLAMATHENSIS   (Meebiam). 

Klamath  Flying  Squirrel. 

[PI.  Ill,  fig.  3  ;  PI.  V,  fig.  3  ;  PI.  VII,  fig.  3.] 

Sciuropterus  alpinus  klamathensis  Merriam,  Proc.  Biol.  Soc.  Washington,  XI. 
p.  225,  July  15, 1897. 

Type  locality. — Fort  Klamath,  Oregon. 

Distribution. — Central  Oregon,  chiefly  east  of  the  Cascades;  north- 
ern and  eastern  limits  unknown. 

Characters. — Similar  to  fuliginosus  but  upperparts  averaging 
slightly  paler ;  gray  on  face  purer  and  more  extensive  (less  mixed  with 
buff);  underparts  paler,  usually  washed  with  pale  j'ellow;  soles 
strongly  tinted  with  yellow;  audital  bulla?  larger.    Similar  to  bullatus 

1  Collection  Mus.  Comp.  Zool.  2  Collection  Amer.  Mus.  Nat.  Hist 


1918.]  GLAUCOMYS  SABRINUS  GROUP.  53 

but  bullae  smaller,  colors  paler,  and  underparts  and  soles  shaded 
with  yellowish. 

Color. — Winter  pelage:  Upperparts  cinnamon-drab  or  pale  sayal 
brown  shading  in  some  specimens  to  light  yellowish  drab;  sides  of 
face  smoke  gray,  often  considerably  washed  with  pinkish  cinnamon ; 
upper  surface  of  flying  membrane  fuscous;  ears  drab  or  pale  hair- 
brown;  hind  feet  hair-brown,  the  soles  olive-ochre  or  deep  colonial 
buff ;  fore  feet  light  brown  mixed  with  buffy  white ;  tail  above,  sayal 
brown  or  dull  cinnamon,  much  mixed  with  hair-brown,  the  tip  nearly 
pure  hair-brown;  beneath,  pinkish  buff,  light  pinkish  cinnamon,  or 
colonial  buff,  moderately  shaded  with  hair-brown,  chiefly  on  sides; 
underparts  whitish,  rather  heavily  washed  with  colonial  buff  or 
cream-buff,  shading  to  light  pinkish  cinnamon  along  sides. 

Skull. — Similar  to  that  of  fuliginosus  but  with  larger  bullae ;  simi- 
lar to  that  of  hullatus  but  smaller,  with  smaller  bullae  and  flatter 
braincase. 

Measurements. — Average  of  9  topotypes  (adult  and  subadult)  : 
Total  length,  319  (300-336) ;  tail  vertebrae,  144  (135-154)  ;  hind  foot, 
40.4  (39.5-42).  Skull:  Average  of  6  topotypes  (adult  and  subadult)  : 
Greatest  length,  40.9  (40.2-42.3)  ;  zygomatic  breadth,  23.6  (23.2-25) ; 
mastoidal  breadth,  18.9  (18.6-19.6)  ;  least  interorbital  breadth,  7.8 
(7-9)  ;  least  postorbital  breadth,  9.2  (8.6-10)  ;  length  of  nasals,  12.7 
(11.8-13.9)  ;  maxillary  toothrow,  8.6  (8.1-9). 

Type  specimen. — No.  87310,  U.  S.  Nat.  Mus.  (Biological  Survey  col- 
lection) ;  9  ad.,  skin  and  skull;  collected  January  11,  1897,  by  B.  L. 
Cunningham. 

Remarks. — This  race  occupies  the  interior  of  Oregon,  chiefly  east 
of  the  Cascades,  but  extending  into  the  eastern  foothills  of  that  range 
and  occasionally  as  high  as  Crater  Mountain  (4  miles  south  of  Crater 
Lake) .  It  intergrades  with  fuliginosus  in  the  Cascades  and  with  flavi- 
ventris  in  northeastern  California.  A  series  of  specimens  from  Davis 
Mountain,  Crook  County,  Oreg.,  exhibit  the  characters  of  the  sub- 
species in  a  more  pronounced  way  than  the  series  from  the  type 
locality.  In  color  they  are  more  drab  above  and  have  the  underparts 
and  feet  more  intensely  yellowish;  the  skulls  average  somewhat 
larger.  A  specimen  from  Paulina  Lake,  Oreg.,  differs  from  typical 
specimens  in  having  gray  instead  of  yellow  feet,  and  approaches 
fuliginosus  also  in  the  color  of  the  back  and  underparts. 

Specimens  examined. — Total  number,  27,  as  follows : 

Oregon:  Crater  Mountain  (4  miles  south  of  Crater  Lake),  1;  Davis  Moun- 
tain, Crook  County,  7  ;*  Foi*t  Klamath,  17 ;  Paulina  Lake,  1 ;  "  Upper 
Klamath"  [Lake],  1." 

'Collection  Game  Dept.  Oregon.  2 Collection  Field  Mus.  Nat.   Hist. 


54  NORTH  AMERICAN   FAUNA.  [No.  44. 

GLAUCOMYS  SABRINUS  FLAVIVENTRIS  Howell. 

Yellow-Bellied  Flying  Squirrel. 

[PI.  Ill,  fig.  2  ;  PI.  V,  fig.  2  ;  PL  VII,  fig.  2.] 

Glaucomys  sabrinus  flaviventris  Howell,  Proc.  Biol.  Soc.  Wash.,  XXVIII,  p.  112, 
May  27,  1915. 

Type  locality. — Head  of  Bear  Creek,  Trinity  County,  California 
(altitude,  6,400  feet). 

Distribution. — Northern  California,  from  the  Trinity  Mountains  in 
Siskiyou  and  Trinity  Counties  .east  to  the  Warner  Mountains,  Modoc 
County. 

Characters. — Similar  to  klamathensis  but  smaller,  with  much 
smaller  audital  bullae;  underparts  and  feet  more  strongly  suffused 
with  yellow  or  buff ;  tail  darker  beneath. 

Color. — Summer  pelage :  Upperparts  pale  sayal  brown  varying  in 
some  specimens  to  drab ;  sides  of  head  smoke  gray,  often  tinged  with 
buff;  ears  hair-brown;  upper  surface  of  ftying  membrane  fuscous; 
feet  hair-brown,  often  tinged  with  pale  greenish  yellow,  the  soles  oil 
yellow ;  tail  above,  dark  buffy  brown,  shading  to  pale  fuscous  at  tip ; 
beneath,  brownish  cream-buff,  bordered  on  sides  with  pale  fuscous  or 
hair-brown;  underparts  whitish,  usually  heavily  washed  with  pale 
greenish  yellow  or  Naples  yellow,  shading  to  wood  brown  along  sides ; 
a  small  white  patch  on  throat,  the  hairs  white  to  the  roots. 

Skull. — Similar  to  that  of  klamathensis,  but  decidely  smaller,  with 
much  smaller  bulla?;  much  smaller  than  that  of  fuliginosus;  closely 
similar  to  that  of  lascivus. 

Measurements. — Average  of  5  topotypes  (adult)  :  Total  length,  306 
(286-322);  tail  vertebra?,  133  (122-140);  hind  foot,  40.4  (40-42). 
Skull:  Greatest  length,  39.5  (38.7-40.4)  ;  zygomatic  breadth,  23.5 
(23.2-24.2) ;  mastoidal  breadth,  17.9  (17.6-18.7) ;  least  interorbital 
breadth,  7.2  (6.3-8.3) ;  least  postorbital  breadth,  8.9  (8.2-9.8)  ;  length 
of  nasals,  12.3  (11.8-12.6) ;  maxillary  toothrow,  8.4  (7.7-8.8). 

Type  specimen. — No.  13319,  Mus.  Vert.  Zool.,  Univ.  California; 
$  ad.,  skin  and  skull;  collected  August  13,  1911,  by  Annie  M. 
Alexander. 

Remarks. — This  race  is  apparently  most  nearly  related  to  lascivus 
of  the  Sierra  Nevada,  from  which  it  differs  widely  in  color  of  under- 
parts. It  differs  so  much  from  oregonensis,  both  in  color  and  cranial 
characters,  that  intergradation  between  them  seems  very  improbable. 
Specimens  from  the  Warner  Mountains,  Cal.,  are  intermediate  be- 
tween this  form  and  klamathensis ;  in  color  they  resemble  flaviventris, 
but  one  of  the  two  individuals  has  a  skull  equaling  that  of  klamath- 
ensis, though  with  smaller  bulla?.  A  large  series  from  Rush  Creek, 
Siskiyou  County,  average  slightly  less  yellowish  beneath  and  have 
somewhat  larger  skulls  than  tj'pical  specimens,  some  of  them  equaling 


1918.]  GLAUCOMYS   SABRINUS  GROUP.  55 

skulls  of  fuliginosus  in  size.     Intergradation  with  lascivus  occurs  in 
the  region  around  Mount  Lassen. 

Specimens  examined. — Total  number,  27,  as  follows : 

California:1  Bear  Creek,  Trinity  County  (head,  altitude,  6,400  feet),  5; 
Castle  Lake,  Siskiyou  County,  2;  Grizzly  Creek,  Trinity  County  (alti- 
tude, 6,000  feet),  1;  Jackson  Lake,  Siskiyou  County,  3;  Rush  Creek, 
Siskiyou  County,  14 ;  Warner  Mountains,  2. 

GLAUCOMYS  SABRINUS  LASCIVUS  (Bangs). 

Sierra  Flying  Squirrel. 

[PI.  Ill,  fig.  5  ;  PI.  V,  fig.  5  ;  PI.  VII,  fig.  1.] 

Sciuropterus  alpinus  lascivus  Bangs,  Proc.  New  England  Zool.  Club,  I,  p.  69, 
July  31,  1899. 

Type  locality. — Tallac,  Eldorado  County,  California. 

Distribution. — Sierra  Nevada  Range  and  northward  to  eastern 
Shasta  County,  California. 

Characters. — Similar  to  flaviventris,  but  slightly  smaller,  with 
smaller  hind  foot;  underparts  without  yellow  suffusion.  Compared 
with  klamathensis:  Size  smaller;  color  above  more  drab  (less 
brownish)  ;  underparts  and  feet  less  yellowish.  Similar  in  color  to 
stephensi,  but  paler. 

Color. — Winter  pelage :  Upperparts  wood  brown  shading  to  drab ; 
sides  of  face  smoke  gray,  often  shaded  with  pale  fuscous;  upper 
surface  of  flying  membrane  blackish  brown ;  ears  fuscous-black ;  hind 
feet  hair-brown,  the  toes  shaded  with  whitish  or  buff;  fore  feet  mouse 
gray;  tail  above,  fuscous-black  mixed  with  wood  brown  or  pale 
snuff  brown;  beneath,  drab-gray  shading  to  avellaneous;  underparts 
grayish  white  faintly  washed  with  avellaneous.  Summer  pelage: 
Upperparts  more  brownish,  usually  rich  wood  brown;  underparts 
faintly  washed  with  cartridge  buff  or  light  pinkish  cinnamon ;  upper 
surface  of  membranes  clove  brown;  hind  feet  pale  hair-brown,  the 
soles  whitish  drab  or  faintly  tinged  with  olive-buff;  under  side 
of  tail  pinkish  buff. 

Skull. — Essentially  like  that  of  flaviventris;  much  smaller  than 
that  of  klamathensis  and  of  fuliginosus. 

Measurements. — Average  of  5  topotypes  (adult  and  subadult)  :  Total 
length,  303  (295-320) ;  tail  vertebras,  133  (125-150)  ;  hind  foot,  40.2 
(39-43).  Average  of  11  from  Cisco,  Cal. :  292;  122;  37.5.  Skull: 
Average  of  4  topotypes  (adult  and  subadult)  :  Greatest  length,  40.4 
(39.5^tl.5);  zygomatic  breadth,  23.5  (23-24.1);  mastoidal  breadth, 
18.2  (17.7-18.8)  ;  least  interorbital  breadth,  7.4  (7.1-7.8)  ;  least  postor- 
bital  breadth,  9.2  (9-9.5)  ;  length  of  nasals,  13  (12.7-13.2)  ;  maxil- 
lary toothrow,  8.1. 

1  All  in  collection  Mus.  Vert.  Zool.,  Univ.  California. 


56  NORTH   AMERICAN   FAUNA.  I  No.  44. 

Type  specimen. — No.  9186,  Mus.  Comp.  Zool.  (formerly  in  collec- 
tion of  E.  A.  and  O.  Bangs)  ;  5  subad.,  skin  and  skull;  collected 
August  28,  1898,  by  W.  W.  Price  and  P.  O.  Simons. 

Remarks. — The  Sierra  flying  squirrel  is  closely  related  to  flavi- 
ventris  of  northern  California,  but  differs  markedly  from  it  in  the 
color  of  the  underparts,  tail,  and  feet.  Intergradation  between  the 
two  forms  is  shown  by  specimens  from  Mount  Lassen,  Dana,  Pratt- 
yille,  and  Castle  Lake,  Siskiyou  County,  which  approach  ftaviventris 
in  having  the  soles  yellowish  and  the  underparts  tinged  with  the 
same.  One  specimen  from  Echo,  close  to  the  type  locality,  also  has 
yellow  soles  and  a  buffy  tail,  but  no  yellow  on  the  belly.  In  a  large 
series  from  Cisco,  Cal.,  most  of  the  specimens  show  indications  of 
being  stained  on  the  underparts  with  soot.  Intergradation  with 
calif omicus  is  probable,  but  there  is  no  material  available  from  the 
southern  Sierra  to  show  it.  This  race  is  very  similar  in  color  to 
alpinws  of  the  northern  Rocky  Mountains,  but  is  of  much  smaller  size 
and  has  a  paler  tail. 

Specimens  examined. — Total  number,  34,  as  follows: 

California:  Blue  Canyon,  3;1  Cisco,  IS;1  Dana,  1;  Echo,  l;2  Fort  Crook, 
1;  Fresno  [=inoiantains  near  ?],  1  (skeleton)  ;  Kings  River  Canyon,  1 ; 
Mill  Creek,  south  base  Mount  Lassen,  1 ;  Mount  Tallac,  4 ; 2' 3  Prattville 
(12  miles  northeast),  1;  Quincy,  1;  Red  Point,  Placer  County,  1.* 

GLAUCOMYS  SABRINUS  CALIFORNICUS   (Rhoads). 

San  Bernardino  Flying  Squirrel. 

[PI.  Ill,  fig.  9  ;  PI.  V,  fig.  9  ;  PI.  VII,  fig.  5.] 

Sciuropterus  alpinus  californicus  Rhoads,  Proc.  Acad.  Nat.  Sci.,  Philadelphia, 

p.  323,  June,  1S97. 
Sciuropterus  californicus  Grinnell,  Univ.  California.  Publ.  Zool.,  V,  pp.  138-139, 

1908. 

Type  locality. — Squirrel  Inn,  San  Bernardino  Mountains,  Cali- 
fornia (altitude,  5,200  feet). 

Distribution. — San  Bernardino  and  San  Jacinto  Mountains,  Cali- 
fornia. 

Characters. — Similar  to  lascivus,  but  upperparts  paler  and  more 
grayish ;  face  between  eyes  usually  washed  with  gray ;  skull  smaller. 

Color. — Unworn  pelage  (April  to  August)  :  Upperparts  light 
drab,  shaded  in  some  specimens  with  yellowish  wood  brown;  sides 
of  face  pale  smoke  gray ;  front  of  face  often  washed  with  gray ;  ears 
hair-brown;  upper  surface  of  flying  membrane  fuscous;  feet  hair- 
brown  or  mouse  gray,  the  toes  whitish  or  buffy  white,  the  soles  buffy 
white  or  olive-buff;  tail  above,  hair-brown,  sparingly  mixed  with 

1  Collection  Mus.   Vert.   Zool.,  Univ.   California. 

2  Collection  Mus.  Comp.  Zool. 

s  Collection  Acad.  Nat.  Sci.  Philadelphia. 
4  Collection  Leland  Stanford,  Jr.,  Univ. 


1918.]  GLAUCOMYS   SABRINUS  GROUP.  57 

sayal  brown ;  beneath,  pinkish  buff  or  pale  olive-buff,  edged  on  sides 
with  hair-brown;  underparts  soiled  whitish,  faintly  shaded  with 
cartridge  buff. 

Skull. — Similar  to  that  of  lascivus,  but  averaging  smaller,  with 
shorter  nasals  and  smaller  bulla?. 

Measurements. — Average  of  8  adults  from  San  Bernardino  Moun- 
tains, Cal. :  Total  length,  297  (280-312)  ;  tail  vertebrae,  137  (127-149)  ; 
hind  foot,  37.6  (36-39).  Skull:  Average  of  5  adults  from  same 
locality:  Greatest  length,  39  (37.3-40);  zygomatic  breadth,  22.8 
(22.3-23.5);  mastoidal  breadth,  17.7  (17.2-18.1);  least  interorbital 
breadth,  7.8  (7.4-8.2)  ;  least  postorbital  breadth,  8.7  (7.9-9.6) ;  length 
of  nasals,  11.9  (11.3-12.4)  ;  maxillarv  toothrow,  8.1  (7.5-8.6). 

Type  specimen. — No.  10487,  Acad.  Nat.  Sci.  Philadelphia  (for- 
merly No.  3487,  collection  S.  N.  Rhoads) ;  $  ad.,  skin  and  skull ; 
collected  June  5,  1896,  by  R.  B.  Herron. 

Remarks. — This  subspecies  is  the  grayest  of  the  races  of  sabrinus, 
being  considerably  paler  than  aZpinus.  It  is  closely  related  to 
laschnts  of  the  Sierra,  and  probably  intergrades  with  it,  though  no 
intermediate  specimens  have  thus  far  been  collected. 

Specimens  examined. — Total  number,  9,  as  follows : 
California :  San  Bernardino  Mountains,  8 ; "  San  Jacinto  Mountains,  l.2 

GLAUCOMYS  SABRINUS  STEPHENS!  (Mekriam). 

California  Coast  Flying  Squirrel. 

[PI.  Ill,  fig.  1  ;  PI.  V,  fig.  1  ;  PI.  VII,  fig.  9.] 

Sciuropterus  oregonensis  stephensi  Merriam,  Proc.  Biol.  Soc.  Washington,  XIII, 
p.  151,  June  13,  1900. 

Type  locality. — Sherwood,  Mendocino  County,  California. 

Distribution. — Coast  region  of  northern  California;  limits  of  range 
unknown. 

Characters. — Similar  to  lascivus,  but  upperparts  more  reddish 
(less  drab)  ;  soles  whiter  (never  tinged  with  yellowish)  and  toes  less 
distinctly  marked  with  buffy  or  white;  skull  with  very  deep  brain- 
case.  Compared  with  oregonensis :  "Upperparts  and  tail  less  reddish; 
underparts  whiter.  Compared  with  fuliginosus :  Size  much  smaller ; 
color  above  in  fresh  pelage  somewhat  grayer  (less  ochraceous)  and 
underparts  paler. 

Color. — Fresh  winter  pelage:  Upperparts  wood  brown;  sides  of 
face  pale  smoke  gray ;  upper  surface  of  flying  membrane  fuscous  or 

1  Three  in  collection  Acad.  Nat.  Sci.  Philadelphia ;  three  in  Mus.  Vert.  Zool.,  Dniv. 
California  ;  one  in  Mus.  Comp.  Zool. 

a  Collection  Mus.  Vert.  Zool.,  Univ.  California. 


58  NORTH  AMERICAN   FAUNA.  [No.  44. 

fuscous-black ;  feet  hair-brown,  the  soles  buffy  white ;  tail  above,  hair- 
brown,  shaded  with  fuscous-black,  darkest  at  tip ;  beneath,  pale  pink- 
ish buff  edged  with  hair-brown;  underparts  whitish,  irregularly 
washed  with  light  pinkish  cinnamon.  Worn  winter  pelage  (May)  : 
Upperparts  decidedly  more  brownish,  about  sayal  brown;  tail  above, 
somewhat  browner;  beneath,  light  pinkish  buff,  edged  with. fuscous. 
Summer  pelage  (July)  :  Similar  to  spring  pelage  but  underside  of 
tail  darker  (pinkish  buff). 

Skull. — Similar  to  that  of  oregonensis  and  of  lascivus,  but  averag- 
ing slightly  smaller,  with  relatively  shorter  nasals  and  slightly 
smaller  bulla? ;  braincase  deeper  and  more  abruptly  depressed. 

Measurements. — Average  of  3  adults  from  Laj^tonville,  Cal. : 
Total  length,  307  (300-310)  ;  tail  vertebra?,  143  (133-149)  ;  hind  foot, 
39.3  (38-40)  Skull:  Average  of  3  adults  from  Laytonville,  Cal.: 
Greatest  length,  40  (39.7-40.5)  ;  zygomatic  breadth,  23.5  (23-2*4) ; 
mastodial  breadth,  18.3  (18.3-18.4)  ;  least  interorbital  breadth,  7.9 
(7.6-8.4)  ;  least  postorbital  breadth,  9.2  (8.6-9.9) ;  length  of  nasals, 
11.7  (11.1-12.6) ;  maxillary  toothrow,  7.9  (7.3-8). 

Type  specimen.— -No.  99830,  IT.  S.  Nat.  Mus.  (Biological  Survey 
collection)  ;  $  subad.,  skin  and  skull;  collected  May  10,  1894,  by  F. 
Stephens. 

Remarks. — This  race,  as  indicated  by  its  skull,  is  closely  related  to 
oregonensis;  in  color  it  resembles  lascivus  rather  closely,  and  doubt- 
less intergrades  with  both  it  and  aregonensis.  The  type  (from  Sher- 
wood) and  one  nearly  adult  specimen  from  La3^tonville  have  skulls 
in  which  the  occipital  region  is  very  abruptly  decurved,  but  in  two 
other  adults  from  the  same  locality  this  character  is  less  pronounced. 
A  specimen,  without  skull,  from  8  miles  southeast  of  Cecilville — 
within  the  known  range  of  flaviventris — agrees  essentially  with  the 
type  of  stephensi,  and  thus  suggests  that  intergradation  does  not 
occur  between  these  two  races. 

Specimens  examined. — Total  number,  6,  as  follows: 
California:  Cecilville  (8  miles  southeast),  1  ;*  Eureka,  l;a  Laytonville,  3;3 
Sherwood,  1. 

1  Collection  Mus.  Vert.  Zool.,  Univ.  California. 

2  Collection  Leland  Stanford,  Jr.,  Univ. 

3  Collection  California  Acad.  Sci. 


TABLES   OF   MEASUBEMENTS. 


59 


TABLES   OF   CRANIAL  MEASUREMENTS. 
Table  I. — Glaucomys  volans. 


Species  and  locality. 


•S 

■3 

03 
CD 

X5 

03 

03 

.a 
?, 

3 
S 

eg 

o 

Ph 

Mm. 

Mm. 

7 

9.1 

7.3 

9.3 

7 

9.2 

6.2 

8.6 

6.5 

8.4 

7.5 

8.9 

6.9 

9.1 

7 

9.4 

6.0 

8.2 

6.9 

8.6 

6.8 

8.3 

7.2 

8.7 

6.8 

9.1 

7.2 

8.3 

7.3 

9 

7.3 

8.4 

7 

8.2 

7.3 

8.9 

7.3 

8.9 

6.9 

8.5 

7.2 

8.7 

7.2 

9.2 

Remarks. 


Glaucomys  volans  volans: 
Washington,  D.  C 

Do 

Do 

Do 

Lake  George,  N.  Y... 
Wilmington,  Mass. . . 
Elk  River,  Minn 

Do 

Glaucomys  v.  querceti: 

Citronelle,  Fla 

Do 

Lake  Harney,  Fla 

Enterprise,  Fla 

Glaucomys  v.  saturatus: 
Dothan,  Ala 

Do 

Do 

Greensboro,  Ala 

Glaucomys  v.  texensis: 

Sour  Lake,  Tex 

Do 

Do 

Do 

Glaucomys  v.  goldmani: 
Teopisca,  Chiapas 


Mm. 
9.6 
9.8 

10.1 
9.2 
9.6 
9.9 
9.1 
9.9 


9.6 
9.6 


9.7 
9.5 
10 
9.7 


9.5 
9.2 


9.2 
9.2 


10.3 
9.9 


Mm. 
6.2 
6.3 
6.5 
6 

5.6 
6.6 
6.5 
6.4 


6.3 
6.4 
6.5 
6.4 


6.4 
6.5 
6.7 
6.2 


6.4 
6.3 


6.2 
6.1 


6.4 
6.3 


Adult. 

Subadult. 

Adult. 

Do. 

Do. 

Do. 

Do. 

Do. 


Adult;  type. 
Adult. 
Subadult. 
Do. 


Adult. 
Adult;  type. 
Adult. 
Do. 


Subadult. 
Old    adult; 

type. 

Adult. 

Do. 


Adult;  type. 
Adult. 


1  Collection  Mus.  Comp.  Zool. 
Table  II. — Glaucomys  sabrinus. 


Species  and  locality. 


Remarks. 


Glaucomys  sabrinus  sabrinus: 

Cochrane,  Ontario 

Lake  Edward,  Quebec 

Big  Island,  Mackenzie 

Winnipeg,  Manitoba 

Glaucomys  s.  makkovikensis 

Makkovik,  Labrador 

Do 

Glaucomys  s.  macrotis: 

Hunter  Mountain,  N.  Y 

Peterboro,  N.  Y 

Ossipee;  N.  H 

Greenville,  Me 

Glaucomys  s.  canescens: 

Portage  la  Prairie,  Manitoba. 
Do 


Mm. 
40 
39.4 
40.2 
39.4 

40.8 
40.4 

37.5 

36.7 

36 

38.7 


3    38.5 
9    38.8 


Mm. 

24 

24.2 

24.2 

24.3 

24.7 
24.4 

22.4 
22.4 
22.3 
23.5 

22.8 
22.9 


Mm. 
18.1 
18.5 
19.2 
18 


18.6 

17.5 
17.7 
17.2 
17.9 

17.5 
17.6 


Mm. 
7.3 
8.1 

8.2 

7.8 

7.3 
7.6 

7 

7.1 
6.9 
6.8 

7.7 
7.6 


Mm. 
8.9 
9.7 
9.2 
9.3 

10 

9.7 

9.3 
9.2 
8.6 


9.2 
9.5 


Mm. 
11.5 
11.4 
12.8 
12 

12.2 
11.1 

10.6 
10.5 
10.5 
10.1 

11.5 
11.2 


Mm. 
7.8 
7.3 

8.1 

7.7 

7.4 
7.9 

7.1 
6.4 
7.2 

6.7 

7.7 
7.6 


Subadult. 
Adult. 
Subadult. 
Do. 

Adult. 
Do. 

Adult;  type. 
Adult. 

Do. 

Do. 

Do. 
Adult;  type. 


1  Collection  Carnegie  Mus.;  not  numbered. 
»  Collection  Field  Mus.  Nat.  Hist. 


1  Collection  Mus.  Comp.  Zool. 


60 


NORTH   AMERICAN   FAUNA. 
Table  II. — Glaucomys  sabrinvs — Continued. 


[No.  44. 


No. 


Species  and  locality. 


16959 


31674 
169169 


174398 
174397 

3.5320 
180338 

136137 
136138 
2S790 

3  253 

«  146 
141952 

69416 

94310 

s  1026 
20G543 


93282 
93144 

68573 
159393 
159394 


6  5905 
6  5904 

31675 
18499 
18495 


193991 
87305 

213319 

2  13321 
213318 

19186 

19187 

3  11458 

» 10487 

310486 
18646 


Glaucomys  s.  bangsi: 
Idaho  Co.,  Idaho. 


Sawtooth  Lake,  Idaho. . . 

Florence,  Mcnt 

Glaucomys  s.  alpinus: 

Jasper  House,  Alberta . . . 

Henry  House,  Alberta. . . 

Do 

Glaucomys  s.  yukonensis: 

Camp  Davidson,  Alaska. 

Tanana,  Alaska 

Glaucomys  s.  zaphseus: 

Helm  Bay,  Alaska 

Do..: 

Bradfield  Canal,  Alaska.. 
Glaucomys  s.  oregonensis: 

Columbia  River,  Oreg 

Salem,  Oreg 

Portland,  Oreg 

Gold  Beach,  Oreg 

Glaucomys  s.  columbiensis: 

Okanogan,  B .  C 


Penticton,  B.  C 

Cranbrook,  B.  C 

Glaucomys  s.  fuliginosus: 
Martin  Station,  Wash. 


Keechelus  Lake,  Wash 

Do 

Glaucomys  s.  latipes: 

Glacier,B.C 

Coolin,  Idaho 

Do 

Glaucomys  s.  olympicus: 

Olympic  Mountains,  Wash. 


Do 

Do 

Glaucomys  s.  bullatus: 

Sawtooth  Lake,  Idaho. 

Ketchum,  Idaho 

Do 

Glaucomys  s.  ~klamathensis: 
Fort  'Klamath,  Oreg. . . 


Do. 
Do. 


Glaucomys  s.  flavivcntris: 

Bear  Creek,  Trinity  Co.,Cal. 

Do 

Do 

Glaucomys  s.  lascivus: 

Mount  Tallac,  Cal 


Do. 
Do. 


Glaucomys  s.  californicus: 

San  Bernardino  Mountains. 
Cal. 

Do 

Do 

Glaucomys  s.  stephensi: 

Sherwood,  Cal 


Mm. 

38.8 

40.9 
41.2 

41.1 
41.2 

42 

41.5 
41 

41.5 
40.1 
41.2 

10. 5+ 
40.7 
40.5 
38.5 

40.5 

41.2 
41.6 

39.5 

42.5 
41.9 

44.2 

43.7 
41.6 


41.6 
40.5 

44 

44.2 

42.6 


40.2 
42.3 

40 
40.4 

38.7 

39.5 


9  1.5 

9  j40.4 

?  39.3 

9  37.3 

9  40 

9  38.4 


Mm. 

23.7 

24.8 
24.7 

25.4 

25.1 
25.0 

25.3 
25.1 

26.2 
24.2 
24.7 

24.8 
24.9 
23.7 
23 

24.4 

25.2 
24.6 

23.7 

24.3 
24.4 

25.1 

24.2 

24 

23.7 

24.7 
23.7 


Mm. 

18.6 

18.7 
19.1 

19.4 
19.4 
19.4 

19.5 

19.7 

19.1 
18.2 
18.9 

19 

18.6 
19 
18.2 

18.9 

19.4 
19.5 


19.1 
18.8 

20 

19.5 

19 


19 
IS.  5 


25      "  19 
25.  5     19.  4 
24.4     19 


23.4     19 
25         19.6 


23.4 
24.2 
23.3 

23 

23.3 
24.1 


17.9 
18.7 
17.6 

17.7 

18.8 
18.4 


23.5     17.6 


22.3     IS.  1 
22.9  '  18 


22.5     17.4 


Mm. 

7.4 


7.8 
7.8 


8 
8.2 


7.8 
8.3 

8.3 
7.4 
8.4 

S.5 
8.1 
8.8 
7.6 

7.7 

7.7 
8.6 


8.6 


8.3 

8 


8.4 
7.6 


9.6 
8.1 


7.8 
8.3 

7.9 
8.3 
7.2 

7.8 

7.2 
7.6 

8.2 

7.4 
7.4 

7.4 


Mm 
8.4 
9 


9 
9.1 

9.6 

10.3 


9 

8.8 

9.2 

8.9 

8.7 

9 

8.1 

9.2 

8.4 
9 

9.3 

9.6 


9.6 
9.2 


8.7 

10 
9.4 

9.3 

9.S 
8.7 

9.5 

9.2 
9 

9.2 

7.9 
8.4 


Mm 

lL.fi 

11.8 
12.4 

12.4 
12.6 
12.4 

12 
11.6 

12.9 
12.3 
12.6 

12.4 
12.9 
12.5 
10.9 

12 

12.9 
12.6 

12.6 

13 
13 

14 

13.6 

12.5 


12.5 
12.5 

13.9 
13.5 
13.9 

12.6 

12.7 
13.9 

12.3 
11.8 
12.6 

12.8 

13.2 
12.7 

11.7 

11.3 
12.1 

11.7 


Mm 

8.7 


8.1 
7.5 
7.9 

7.9 

8.  4 


8.3 

7.8 

8.1 

8.7 

8 

7.9 

7.7 

7.7 
8 

8.6 

8.8 
8.2 

8.8 
9 


i  Collection  Mus.  Comp.  Zool. 

2  Collection  Mus.  Vert.  Zool.,  Univ.  California. 

8  Collection  Acad.  Nat.  Sci.,  Philadelphia. 


«  Collection  Game  Dept.  Oregon. 
6  Collection  Victoria  Mem.  Mus. 
6  Collection  Field  Mus.  Nat.  Hist. 


EXPLANATION  OF  PLATES. 

[All  natural  size.] 
/ 
Plate  II. 

Fig.    1.  Glaucomys  volans  saturatus,   $   ad.,  Dothan,  Ala.   (No.  178363,  U.  S. 

Nat.  Mus.,  Biological  Survey  collection). 
Fig.    2.  Glaucomys  volans  volans,   2    ad.,  Cleveland  Park,  D.  C.    (No.  155517, 

U.  S.  Nat.  Mus.). 
Fig.    3.  Glaucomys  sabrinus   bangsi,  ad.,   Florence,   Mont.    (No.   168243,  U.   S. 

Nat.  Mus.,  Biological  Survey  collection). 
Fig.    4.  Glaucomys    sabrinus    alpinus,     2     ad.,    Henry    House,    Alberta     (No. 

174398,  U.   S.  Nat.  Mus.). 
Fig.    5.  Glaucomys  volans  texensis,   2   ad.,  Sour  Lake,  Tex.  (No.  136402,  U.  S. 

Nat.  Mus.,  Biological  Survey  collection). 
Fig.    6.  Glaucomys  volans   goldmani,    2    ad.,  Teopisca,   Chiapas    (No.   132834, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.    7.  Glaucomys    sabrinus    cancsccm,    2    ad.    (type),    Portage    la    Prairie, 

Manitoba  (No.  7663,  Field  Mus.  Nat.  Hist.). 
Fig.   8.  Glaucomys  sabrinus  yukonensis,  2   ad.  (type),  Camp  Davidson,  Alaska 

(No.  35320,  U.   S.  Nat.  Mus.). 
Fig.    9.  Glaucomys  volans  querceti,    2    ad.    (type),  Citronelle,  Fla.    (No.  2451, 

Mus.   Comp.   Zool.). 
Fig.  10.  Glaucomys  sabrinus  macrotis,   2  ad.   (type),  Catskill  Mountains,  N.  Y. 

(No.  83152,  U.  S.  Nat.  Mus.). 
Fig.  11.  Glaucomys  sabrinus  sabrinus,    $   subad.,  Cochrane,  Ontario   (Carnegie 

Mus.,  not  numbered). 
Fig.  12.  Glaucomys    sabrinus    makkovikensis,    ad.,    Makkovik,    Labrador    (No. 

13449,  Mus.  Comp.  Zool.). 
62 


North  American  Fauna  No.  44,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  II. 


Skulls  of  Glaucomys  volans  and  G.  sabrinus. 


1.  G.  v.  saturatus. 

2.  G.  v.  volans. 

3.  G.  s.  bangsi. 

4.  G.  s.  alpinus. 


5.  G.  v.  texensis. 

6.  G.  v.  goldmani. 

7.  G.  s.canescens. 

8.  G.  s.  yukonensis. 


9.  G.  v.  querceti. 

10.  G.  s.  macrotis. 

11.  G.  s.  sabrinus. 

12.  G.  s.  makkovikensis. 


North  American  Fauna  No.  44,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  III. 


1.  G.  s.  stephensi. 

2.  G.  s.  flaviventris. 

3.  G.  s.  klamathensis. 

4.  G.  s.  fuliginosus. 


Skulls  of  Glaucomys  sabrinus. 

5.  G.  s.  lascivus. 

6.  G.  s.  zaphanis. 

7.  G.  s.  columbiensis. 

8.  G.  s.  latipes. 


9.  G.  s.  californicus. 

10.  G.  s.  olympicus. 

11.  G.  s.  oregonensis. 

12.  G.  s.  buUatus. 


Plate  III. 

Fig.    1.  Glaucomys  sabrinus  stephensi,   $   ad.,  Laytonville,  Cal.   (No.  331,  Cal. 

Acad.  Sci.) 
Fig.    2.  Glaucomys  sabrimis  flaviventris,    $    ad.    (type),  Bear  Creek,  Trinity 

County,  Cal.  (No.  13319,  Mus.  Vert.  Zool.). 
Fig.    3.  Glaucomys  sabrinus  klamathcnsis,   2    ad.   (type),  Fort  Klamath,  Oreg. 

(No.  87310,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.   4.  Glaucomys  sabrinus  fuliginosus,    2    ad.,  Easton,   Wash.    (No.   116864, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.    5.  Glaucomys  sabrinus  lascivus,  2  ad.,  Cisco,  Cal.  (No.  19495,  Mus.  Vert. 

Zool.). 
Fig.    6.  Glaucomys  sabrinus  zaphosus,    2    ad.   (type),  Helm  Bay,  Alaska   (No. 

136137,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.    7.  Glaucomys  sabrinus  columbiensis,    2    ad.,  Penticton,  B.  C.    (No.  1026, 

Victoria  Mem.  Mus.). 
Fig.    8.  Glaucomys  sabrinus  latipes,  2    ad.   (type),  Glacier,  B.  C.   (No.  68573, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.    9.  Glaucomys  sabrinus  calif  ornicus,  2  ad.  (type),  San  Bernardino  Moun- 
tains, Cal.  (No.  10487,  Acad.  Nat.  Sci.  Philadelphia). 
Fig.  10.  Glaucomys  sabrinus  olympicus,    $    subad.,  Olympic  Mountains,  Wash. 

(No.  5905,  Field  Mus.  Nat.  Hist.). 
Fig.  11.  Glaucomys   sabrinus   oregonensis,   ad.    (type),   Columbia   River,   Oreg. 

(No.  253,  Acad.  Nat.  Sci.  Philadelphia). 
Fig.  12.  Glaucomys  sabrinus  bullatus,  $   ad.,  Ketchum,  Idaho   (No.  8499,  Mus. 

Comp.   Zool.). 


/ 

Plate  IV. 

Fig.    1.  Glaucomys  volans  saturatus,  $  ad.,  Dothan,  Ala.  (No.  178363,  U.  S.  Nat. 

Mus.,  Biological  Survey  collection). 
Fig.    2.  Glaucomys  volans  volans,    2    ad.,  Cleveland  Park,  D.  C.   (No.  155517, 

U.  S.  Nat.  Mus.). 
Fig.    3.  Glaucomys  sabrinus  bangsi,  ad.,  Florence,  Mont.  (No.  168243,  U.  S.  Nat. 

Mus.,  Biological  Survey  collection). 
Fig.    4.  Glaucomys  sabrinus  alpinus,  2  ad.,  Henry  House,  Alberta  (No.  174398, 

U.  S.  Nat.  Mus.). 
Fig.   5.  Glauccmiys  volans  texensis,   2  ad.,  Sour  Lake,  Tex.  (No.  136402,  U.  S. 

Nat.  Mus.,  Biological  Survey  collection). 
Fig.    6.  Glaucomys  volans  goldmani,    2    ad.,   Teopisca,   Chiapas    (No.   132834, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.    7.  Glaucomys  sabrinus  canescens,  2  ad.  (type),  Portage  la  Prairie,  Mani- 
toba (No.  7663,  Field  Mus.  Nat.  Hist.). 
Fig.    8.  Glaucomys  sabrinus  yukonensis,  2   ad.  (type),  Camp  Davidson,  Alaska 

(No.  35320,  U.  S.  Nat.  Mus.). 
Fig.    9.  Glaucomys  volans  querceti,   2   ad.  (type),  Citronelle,  Fla.     (No.  2451, 

Mus.  Coinp.  Zool.). 
Fig.  10.  Glaucomys  sabrinus  macrotis,  2   ad.  (type),  Catskill  Mountains,  N.  T. 

(No.  83152,  U.  S.  Nat.  Mus.). 
Fig.  11.  Glaucomys  sabrinus  sabrinus,   $   subad.,  Cochrane,  Ontario   (Carnegie 

Mus.,  not  numbered). 
Fig.  12.  Glaucomys    sabrinus    makkovikemis,    ad.,    Makkovik,    Labrador    (No. 

13449,  Mus.  Comp.  Zool.). 


North  American  Fauna  No.  44,  U.  S.  Dept  Agr.     Biological  Survey. 


Plate  IV. 


Skulls  of  Glaucomys  volans  and  G.  sabrinus. 


1.  G.  v.  saturatus. 

2.  G.  v.  volans. 

3.  G.  s.  bangsi. 

4.  G.  s.  alpinus. 


5.  G.  v.  texensis. 

6.  G.  v.  goldmani. 

7.  G.  s.  canescens. 

8.  G.  s.  yukonensis. 


9.  G.  v.  querceti. 

10.  G.  s.  macrotis. 

11.  G.  s.  sabrinus. 

12.  G.  s.  makkovikensL 


North  American  Fauna  No.  44,  U.  S 

.  Dept.  Agr.      Biological  Survey. 

?^v2w  its'  -;;r£ 

...<  ^^^        \^B    *  ^ 

Plate  V. 

B    *1 

BB  *  V  v  '  mfl  '-£§ 

>:.Ms    Jr            USB 

H^^F   %J|E^|  HI 

'ill 

K  ii 

Mm 

I    IB 

|&gSa8B&£  .   ■  ,  -     -  " 

Bll 

IT '  ill 

■  B 

"-B  ^  B "' 
Vj/^^B  fj  B 

n  « 

i^r;  tI''.' 

Iy          .*     BAB !  •  ■ 

f    II     1       TC 

B1  ii 

■  ^^  si 

it  *r?j|ji 

B*  O  *  kfl 

B    j  ;    IB 

BB      J  ■>      B     B 

1.  G.  s.  stephensi. 

2.  G.  s.  flaviventris. 

3.  G.  s.  klamathensis. 

4.  G.  s.  fuliginosus. 


Skulls  of  Glaucomys  sabrinus. 

5.  G.  s.  lascivus. 

6.  G.  s.  zaphseus. 

7.  G.  s.  columbiensis. 

8.  G.  s.  latipes. 


9.  G.  s.  californicus. 

10.  G.  s.  olympicus. 

11.  G.  s.  oregonensis. 

12.  G.  s.  bullatus. 


Plate  V. 

Fig.   1.  Glaucomys  sabrinus  stephensi,   $   ad.,  Laytonville,  Cal.,   (No.  331,  Cal. 
Acad.  Sci.). 

Fig.   2.  Glaucomys  sabrinus  flaviventris,    $    ad.    (type),  Bear  Creek,  Trinity 
County,  Cal.  (No.  13319,  Mus.  Vert.  Zool.). 

Fig.   3.  Glaucomys  sabrinus  klamathensis,   9   ad.  (type),  Fort  Klamath,  Oreg. 
(No.  87310,  TJ.  S.  Nat.  Mus.,  Biological  Survey  collection). 

Fig.    4.  Glaucomys  sabrinus  fuliginosus,    9    ad.,   Easton,   Wash.    (No.  116864, 
U.  S.  Nat.  Mus.,  Biological  Survey  collection). 

Fig.   5.  Glaucomys  sabrinus  lascivus,  9  ad.,  Cisco,  Cal.  (No.  19495,  Mus.  Vert. 
Zool.). 

Fig.   6.  Glaucomys  sabrinus  zaphceus,    9   ad.   (type),  Helm  Bay,  Alaska   (No. 
136137,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 

Fig.    7.  Glaucomys  sabrinus  columbiensis,  9  ad.,  Penticton,  B.  C.  (No.  1026,  Vic- 
toria Mem.  Mus.). 

Fig.   8.  Glaucomys  sabrinus  latipes,   9   ad.   (type),  Glacier,  B.  C.   (No.  68573, 
U.  S.  Nat.  Mus.,  Biological  Survey  collection). 

Fig.   9.  Glaucomys  sabrinus  calif ornicus,  9  ad.  (type),  San  Bernardino  Moun- 
tains, Cal.  (No.  10487,  Acad.  Nat.  Sci.  Philadelphia). 

Fig.  10.  Glaucomys  sabrinus  olympicus,   $   subad.,  Olympic  Mountains,  Wash. 
(No.  5905,  Field  Mus.  Nat.  Hist.). 

Fig.  11.  Glaucomys  sabrinus  oregoncnsis,   ad.    (type),   Columbia  River,   Oreg. 
(No.  253,  Acad.  Nat.  Sci.  Philadelphia). 

Fig.  12.  Glaucomys  sabrinus  bullatus,   $   ad.,  Ketchum,  Idaho  (No.  8499,  Mus. 
Comp.  Zool.). 
14520°— 18— No.  44 5 


Plate  VI. 

Fig.    1.  Glaucomys  volans  volans,    2    ad.,  Cleveland  Park,  D.  C.   (No.  155517, 

U.  S.  Nat.  Mus.). 
Fig.    2.  Glaucomys  volans  querceti,    2    ad.    (type),  Citronelle,  Fla.   (No.  2451, 

Mus.  Comp.  Zool.). 
Fig.    3.  Glaucomys  volans  saturatus,   S    ad.,  Dothan,  Ala.    (No.  178363,  U.  S. 

Nat.  Mus.,  Biological  Survey  collection). 
Fig.    4.  Glaucomys  volans  texcnsis,   2  ad.,  Sour  Lake,  Tex.   (No.  136402,  U.  S. 

Nat.  Mus.,  Biological  Survey  collection). 
Fig.    5.  Glaucomys  sabrinus  macrotis,   2  ad.   (type),  Catskill  Mountains,  N.  Y. 

(No.  83152,  U.  S.  Nat.  Mus.). 
Fig.    6.  Glaucomys  sabrinus  sabrinus,   S   subad.,  Cochrane,  Ontario   (Carnegie 

Museum,  not  numbered). 
Fig.    7.  Glaucomys  sabrinus  columbicnsis,    2    ad.,  Penticton,  B.  C.    (No.  1026, 

Victoria  Mem.  Mus.). 
Fig.    S.  Glaucomys  sabrinus  oregonensis,  adult   (type),  Columbia  River,  Oreg. 

(No.  253.  Acad.  Nat.  Sci.  Philadelphia). 
Fig.    9.  Glaucomys  sabrinus  zaphwus,    2    ad.    (type),  Helm  Bay,  Alaska   (No. 

136137,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  10.  Glaucomys   sabrinus   bangsi,   ad.,   Florence,   Mont.    (No   168243,   U.   S. 

Nat.  Mus.,  Biological  Survey  collection). 
Fig.  11.  Glaucomys  sabrinus  cancsccns,  2  ad.  (type),  Portage  la  Prairie,  Mani- 
toba (No.  7663,  Field  Mus.  Nat.  Hist). 
Fig.  12.  Glaucomys  sabrinus  alpiiws,  2  ad.,  Henry  House,  Alberta  (No.  174398, 

U.  S.  Nat.  Mus.). 


North  American  Fauna  No.  44,  U.  S.  Dept  Agr. 

Biological  Survey.                                                  PLATE  Vl„ 

■    ^5 

•  W             '.  ^g  1     §*  ■""    'V   ■■■'    "'•  '/'■■-; 

ill  l^jfc^w'---: f 

'  Hub!                Ib^_ 

'•v;'.:',^:^."  '  .'"   ;  :' --';' '"-  -  tM  -•' 

Skulls  of  Glaucomys  volans  and  G.  sabrinus. 


1.  G.  v.  volans. 

2.  G.  v.  querceti. 

3.  G.  v.  saturatus. 

4.  G.  v.  texensis. 

5.  G.  s.  macrotis. 

6.  G.  s.  sabrinus. 


7.  G.  s.  columbiensis. 

8.  G.  s.  oregonensis. 

9.  G.  s.  zaphaeus. 

10.  G.  s.  bangsi. 

11.  G.  s.  canescens. 

12.  G.  s.  alpinus. 


North  American  Fauna  No.  44,  U.  S.  Dept.  Agr.     Biological  Survey. 


Plate  VII. 


Skulls  of  Glaucomys  sabrinus  and  Pteromys  buchneri. 


1.  G.  s.  lascivus. 

2.  G.  s.  flaviventris. 

3.  G.  s.  klamathensis. 

4.  G.  s.  bullatus. 

5.  G.  s.  californicus. 

6.  G.  s.  olyrnpicus. 


7.  G.  s.  fuliginosus. 

8.  G.  s.  latipes. 

9.  G.  s.  stephensi. 

10.  G.  s.  sabrinus. 

11,  12,  13,  14.  P.  biickneri. 


Plate  VII. 

Fig.    1.  Glaucomys  sabrinus  lascivus,   2  ad.,  Cisco,  Cal.  (No.  19495,  Mus.  Vert. 

Zool.). 
Fig.    2.  Glaucomys  sabrinus  fiaviventris,    $    ad.    (type),  Bear  Creek,  Trinity 

County,  Cal.  (No.  13319,  Mus.  Vert.  Zool.). 
Fig.    3.  Glaucomys  sabrinus  klamathcnsis,   $   ad.   (type),  Fort  Klamath,  Oreg. 

(No.  87310,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.    4.  Glaucomys  sabrinus  bullatus,   $  ad.,  Ketchum,  Idaho  (No.  8499,  Mus. 

Comp.  Zool.). 
Fig.    5.  Glaucomys  sabrinus  calif  ornicus,   9  ad.  (type),  San  Bernardino  Moun- 
tains, Cal.  (No.  104S7,  Acad.  Nat.  Sci.  Philadelphia). 
Fig.    6.  Glaucomys  sabrinus  olympicus,   $   subad.,  Olympic  Mountains,  Wash. 

(No.  5905,  Field  Mus.  Nat.  Hist.). 
Fig.    7.  Glaucomys   sabrinus  fuliginosus,    $    ad.,  Easton,   Wash.    (No.   116864, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.    8.  Glaucomys  sabrinus  latipes,    $   ad.   (type),  Glacier,  B.  C.    (No.  68573, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.    9.  Glaucomys  sabrinus  stephensi,   $   ad.,  Laytonville,  Oul.   (No.  331,  Cal. 

Acad.  Sci.). 
Fig.  10.  Glaucomys  sabrinus  sabrinus,   $   subad.,  Cochrane,  Ontario   (Carnegie 

Mus.,  not  numbered). 
Figs.  11,  12,  13,  14.  Pteromys  buchneri,   S   ad.,  Shansi  Mountains,  China   (No. 

172618,  U.  S.  Nat.  Mus.). 


IND 

[Synonyms 


alpinus,  Glaucomys,  40. 

Pteromys,  40. 

Sciuropterus,  40. 
atnericana,  Pteromys,   19. 
americanus,  Sciuropterus,  19. 

B. 

bangsi,  Glaucomys,  38. 

Sciuropterus,  38. 
Breeding,   8. 

biichneri,  Pteromys,  15   (footnote), 
bullatus,  Glaucomys,  51. 


californicus,  Glaucomys,   56. 

Sciuropterus,  56. 
canadensis,  Pteromys,  31. 
canescens,  Glaucomys,  37. 
Characters,   generic,    14. 
columbiensis,   Glaucomys,   45. 
cucullatus,  Pteromys,  19. 


E. 


Economic  status,  10. 

F. 

flaviventris,   Glaucomys,   54. 
Food,  9. 

fuliginosus,   Glaucomys,   47. 
Sciuropterus,  47. 

G. 


Generic  names,   12. 
Glaucomys,  genus,  11. 

cranial  measurements,  tables,  59. 
generic   characters,   14. 
history,   12. 
key  to  species,   16. 

list  of  species,  16. 

nomenclature,   12. 

specific  names,  13. 
Glaucomys  alpinus,  40. 

bangsi,   38. 

bullatus,   51. 

californicus,  56. 

canescens,  37. 

columbiensis,   45. 

flaviventris,    54. 

fuliginosus,  47. 

goldmani,  28. 

klamathensis,  52. 

lascivus,  55. 

latipes,  48. 


EX. 

in   italics.] 

Glaucomys  macrotis,  35. 

makkovikensis,    34. 

olympicus,  49. 

oregonensis,  44. 

querceti,  26. 

sabrinus,  31. 

saturatus,  24. 

stephensi,  57. 

texensis,  27. 

volans,  19. 

yukonensis,  41. 

zaphseus,  43. 
goldmani,  Glaucomys,  28. 

Sciuropterus,  28. 


Habits,  5. 

History,  12. 

hudsonius,  Pteromys,  32. 

Sciuropterus,  32,  35. 

Sciurus,  31. 

K. 

Key  to  species,  16. 
klamathensis,  Glaucomys,  52. 
Sciuropterus,  52. 


lascivus,  Glaucomys,  55. 

Sciuropterus,  55. 
latipes,  Glaucomys,  48. 
List  of   species,   16. 

M. 

macrotis,  Glaucomys,  35. 

Sciuropterus,  35. 
makkovikensis,  Glaucomys,  34. 

Sciuropterus,  34. 
Material  examined,  11. 
Measurements,  cranial,  explanation  of,  11. 

tables  of,  59. 
Molt,  10,  21. 
Mus,  genus,  11. 

volans,  19. 

N. 


nebrascensis,  Pteromys,  20. 
Nests,  7. 
Nomenclature,  12. 


olympicus,  Glaucomys,  49. 
Sciuropterus,  49. 


63 


64 


NORTH    AMERICAN    FAUNA. 


[No.  44,  3918] 


oregonensis,  Glaucomys,  44. 
Pteromys,  44. 
Sciuropterus,  44. 

P. 

Pelage,  10. 

petaurista,  Sciurus,  12. 
Pteromys,  generic  characters,  15 

biichneri,   15    (footnote). 

volans,  15   (footnote). 
Pteromys,   genus,   11. 
Pteromys  alpinus,  40. 

amcricana,    19. 

canadensis,  31. 

cucullatits,  19. 

hudsonius,   32. 

nebrascensis,   20. 

oregonensis,  44. 

sabrinus,   32. 

virginianus,  19. 

volucella,  19. 

Q. 

querceti,    Glaucomys,    26. 
Sciuropterus,  26. 

S. 

sabrinus,    Glaucomys,    31. 

Pteromys,    32. 

Sciuropterus,    32. 

Sciurus,    31. 
saturatus,   Glaucomys,   24. 
Sciuropterus,    genus,    11. 
Sciuropterus  alpinus,  40. 

americanus,    19. 

bangsi,    38. 

calif ornicus,    56. 

fuliginosus,    47. 

goldmani,    28. 

hudsonius,    32,    35. 

klamathensis,    52. 

lascivus,  55. 

macrotis,   35. 

makkovikensis,  34. 

olympicus,  49. 


Sciuropterus  oregonensis,  44. 

querceti,  26. 

sabrinus,  32. 

sjfes,   20. 

stephensi,  57. 

volans,  20. 

volucella,  19. 

yukonensis,  41. 

zaphmus,   43. 
Sciurus,   genus,    11. 
Sciurus  hudsonius,  31. 

sabrinus,   31. 

petaurista,   12. 

volans,  12,   13. 

voZitceHa,    19. 
silus,   Sciuropterus,  20. 
Specific   names,    13. 
stephensi,   Glaucomys,  57. 

Sciuropterus,    57. 


texensis,    Glaucomys,    27. 

V. 

virginianus,  Pteromys,   19. 

Voice,   6. 

volans,   Glaucomys,    19. 

JfMS,    19. 

Pteromys,  15   (footnote). 

Sciuropterus,   20. 

Sciurus,  12,    13. 
volucella,   Pteromys,   19. 

Sciuropterus,    19. 

Sciurus,  19. 

Y. 

yukonensis,  Glaucomys,  41. 
Sciuropterus,    41. 


zaphrpus.    Glaucomys,   43. 
Sciuropterus,    43. 


ADDITIONAL  COPIES 

OF  TUTS  PUBLICATION  MAY  BE  PROCURED  FROM 

THE  SUPERINTENDENT  OF  DOCUMENTS 

GOVERNMENT  PRINTING  OFFICE 

"WASHINGTON,  D.  C. 

AT 

15  CENTS  PER  COPY 


V 


URE 


LETTER  OF  TRANSMITTAL. 


United  States  Department  of  Agriculture, 

Bureau  or  Biological  Survey, 
Washington,  D.  C,  January  18,  1921. 
Sir:  I  have  the  honor  to  transmit  herewith,  recommending  that 
it  be  published  as  No.  45  in  the  series  of  the  North  American  Fauna, 
a  report  on  the  biological  survey  of  Alabama,  by  Arthur  H.  Howell' 
assistant  biologist  of  this  bureau.  This  treats  of  the  life  zones  and  the 
mammals  of  the  State.  A  further  report  on  the  birds  has  been  pre- 
pared for  separate  publication.  Both  are  based  on  natural  history 
explorations  conducted  during  recent  years  by  the  bureau.  The  first 
part  of  the  present  report  defines  the  physiographic  features  of  the 
State  and  characterizes  the  two  transcontinental  life  zones  repre- 
sented; it  is  accompanied  by  a  map  showing  the  boundaries  of  each 
zone.  The  second  part  consists  of  notes  on  the  distribution,  abun- 
dance, and  habits  of  the  native  mammals  of  the  State,  of  which  there 
are  65  forms  now  known,  10  having  been  added  as  a  result  of  the 
Survey  investigations,  3  of  them  new  to  science.  This  report  will  be 
of  material  assistance  to  farmers,  students,  and  others  interested  in  the 
habits  and  economic  relations  of  our  native  wild  animals. 
Respectfully, 

E.  W.  Nelson, 

tt       t^  m  n  r  Chief  of  Bureau. 

Hon.  E.  T.  Meredith, 

Secretary  of  Agriculture. 


CONTENTS. 

Page. 

introduction 5 

'art  I. — Physiography  and  life  zones  of  Alabama 7 

Physiographic  features  of  the  State 7 

Life  zones 10 

Upper  Austral  Zone 10 

Lower  Austral  Zone 11 

'art  II.- — Report  on  the  mammals  of  Alabama 17 

History  of  Alabama  mammalogy 17 

Annotated  list  of  mammals 18 

Didelphiidae:  Opossums IS 

Talpidae:  Moles 20 

Soricidae  :  Shrews 21 

Vespertilionidae  :  Typical  bats 23 

Molossidae  :  Free-tailed  bats 28 

Ursidae  :  Bears , 29 

Canidae:  Wolves  and  foxes 30 

Procyonidae  :  Raccoons 34 

Mustelidae  :  Weasels,  minks,  skunks,  otters,  etc 35 

Felidae  :  Cougars,  wildcats,  etc 41 

Muridae:  Mice,  rats,  etc 43 

Geomyidae:   Pocket  gophers 59 

Sciuridae:  Marmots,  squirrels,  etc 60 

Petauristidae :  Flying  squirrels 67 

Castoridae  :  Beavers 67 

Leporidae:  Hares  and  rabbits 70 

Cervidae :  Deer 75 

Bibliography 77 

Index 81 

3 


ILLUSTRATIONS. 


PLATES. 


Page. 

Plate  I.  Map  of  Alabama,  showing  life  zones Frontispiece. 

II.  Fig.  1. — Eastern  slopes  of  Sand  Mountain,  from  Trenton  Val- 
ley. Fig.  2,-^-Gulch  of  Long  Island  Creek,  Sand 
Mountain 8 

III.  Fig.  1. — Rock    ridge    near   summit    of   Choccolocco    Mountain. 

Fig.  2. — Choccolocco  Valley,  near  Piedmont 

IV.  Fig.  1. — Rocky  slopes  near  base  of  Talladega  Mountains.     Fig. 

2. — Upper  slopes  of  Choccolocco  Mountain 8 

V.  Fig.  1. — Tennessee  River  near  Bloomfield.    Fig.  2. — Sipsey  Fork 

at  mouth. of  Bear  Branch S 

VI.  Fig.  1. — Pine    forest   near   Jasper.     Fig.    2. — Turpentine    "or- 

chard," near  Mobile 8 

VII.  Fig.  1. — Catoma    Swamp,   near   Teasley   Mill.     Fig.   2. — River- 
bottom  swamp,  near  Seale 8 

VIII.  Fig.  1.— Sand  dunes,  Orange  Beach.     Fig.  2.— Swamp  on  Dau- 
phin Island,  overflowed  by  sand 12 

IX.  Fig.  1.— Stiggins  Lake,  Tensaw  River  delta.     Fig.  2.— Marsh  on 

bay  side  of  Dauphin  Island 12 

X.  Fig.  1. — Mounds  of  pocket  gopher  in   open  pine  forest.     Fig. 

2. — Mounds  of  pocket  gopher  in  sandy  field 60 

XL  Fig.  1. — Dense  canebrake  bordering  Catoma  Creek.     Fig.  2. — 

Beaver  dam  in  Catoma  Creek 68 

TEXT  FIGURES. 

Fig.  1.  Map  of  provisional  distribution  of  races  of  the  old-field  mouse  in 

Alabama 45 

2.  Map  of  distribution  of  the  white-footed  mouse  in  Alabama 47 

3.  Map  of  distribution  of  races  of  the  cotton  mouse  in  Alabama 49 

4.  Map  of  distribution  of  wood  rats  in  Alabama 53 

5.  Map  of  distribution  of  muskrats  in  Alabama 57 

6.  Map  of  distribution  of  the  Alabama  pocket  gopher  in  Alabama 60 

7.  Map  of  distribution  of  the  southern  woodchuck  in  Alabama 61 

8.  Map  of  distribution  of  fox  squirrels  in  Alabama 66 

9.  Map  of  distribution  of  the  New  England  cottontail  rabbit  in  Ala- 

bama          72 

10.  Map  of  distribution  of  swamp  and  marsh  rabbits  in  Alabama 74 

4 


No.  45.  NORTH  AMERICAN  FAUNA.  October  28, 1921. 


A  BIOLOGICAL  SURVEY  OF  ALABAMA. 


By  Arthur  H.  Howell. 


INTRODUCTION. 

In  the  work  of  mapping  the  life  zones  of  North  America  and 
the  distribution  of  mammals  and  birds — a  project  upon  which  the 
Biological  Survey  has  been  engaged  for  many  years — the  South- 
eastern States  have  until  very  recent  years  received  little  attention. 
The  more  conspicuous  birds  and  mammals  found  in  that  region  were 
known  in  a  general  way,  but  exact  knowledge  of  their  distribution 
was  lacking  and,  as  a  result,  the  boundaries  of  the  life  zones  could  be 
drawn  only  tentatively. 

To  secure  definite  information  regarding  the  distribution,  abun- 
dance, and  economic  relations  of  the  birds  and  mammals  and  at  the 
same  time  to  study  the  distribution  of  the  plant  life  of  this  region, 
a  biological  survey  of  Alabama  was  carried  on  during  a  period 
of  several  years.  From  these  investigations,  and  with  the  aid  of 
Mohr's  detailed  report  on  the  plant  life,1  it  has  been  possible  to  define 
the  ranges  of  the  mammals  and  birds  and  the  boundaries  of  the 
natural  life  zones  within  the  State.  The  report  on  the  bird  life  has 
been  prepared  and  tentative  arrangements  made  for  its  publica- 
tion; the  reports  on  life  zones  and  on  the  mammals  are  presented 
herewith. 

The  field  collecting  on  which  the  present  report  is  chiefly  based 
was  carried  on  by  the  writer,  with  the  assistance  of  James  S.  Gut- 
sell,  Ernest  G.  Holt,  James  L.  Peters,  and  Luther  J.  Goldman  during 
parts  of  the  years  1908  and  1911-1916.  Collections  had  previously 
been  made  by  C.  S.  Brimley  in  1890,  and  by  Russell  J.  Thompson 
in  1892.  The  writer's  work  covered  every  month  in  the  year  and 
practically  all  parts  of  the  State.  Valuable  assistance  was  rendered 
also  by  numerous  residents  of  the  State,  who  were  instrumental  in 

1Mohr,  Charles,  Plant  life  of  Alabama:  Contr.  U.  S.  Nat.  Herb.,  vol.  6,  1901. 

5 


6  NORTH  AMERICAN   FUNA.  [No.  45 

securing  needed  specimens  and  information.  Among  those  who 
rendered  especially  valuable  aid  were  Lewis  S.  Golsan,  of  Pratt- 
ville;  Edward  S.  Graves,  of  Bentonsport,  Iowa  (formerly  residing 
on  Sand  Mountain,  near  Carpenter,  Ala.)  ;  Peter  A.  Brannon,  of 
Montgomery;  and  H.  P.  Loding,  of  Mobile. 


PART  I.  PHYSIOGRAPHY  AND  LIFE  ZONES  OF  ALABAMA. 

PHYSIOGRAPHIC  FEATURES  OF  THE  STATE. 

Alabama  presents  a  wide  diversity  of  physiographic  conditions, 
ranging  from  the  low  sandy  islands  and  pine-covered  flats  of  the 
coast  region  through  the  fertile  valleys  and  rolling  hills  of  the 
central  parts,  to  the  rugged  and  rock-crowned  ranges  of  the  north- 
east, reaching  in  Clay  County  an  altitude  of  2,400  feet  above  sea 
level.  The  principal  natural  divisions  of  the  State  are  the  fol- 
lowing:2 Mountain  Region,  Tennessee  Valley,  Warrior  Basin  and 
Tableland,  Central  Prairie  Belt,  Coast  Pine  Belt,  and  Coast  Plain  or 
Gulf  Strip. 

Mountain  Region. 

The  Mountain  Region  comprises  the  southernmost  spurs  of  the 
Appalachian  system  and  covers  approximately  the  northeastern 
portion  of  the  State  from  northern  Coosa  County  northward.  The 
principal  ranges  are  the  Talladega  Mountains,  in  Talladega  and 
Clay  Counties  (Cheaha  Peak,  altitude  2,407  feet)  ;  Choccolocco 
Mountain,  in  Calhoun  County  (2,074  feet)  ;  Oakey  Mountain,  in 
Cleburne  County  (1,945  feet)  ;  Mount  Weisner,  in  Cherokee  County 
(1,900  feet)  ;  Lookout  Mountain  (1,847  feet,  near  Mentone) ;  Sand 
Mountain  (1,785  feet,  near  Carpenter)  (the  last  two  being  parallel 
ridges  extending  from  the  vicinity  of  Gadsden  and  Attalla  north- 
eastward through  Dekalb  County),  and  the  Cumberland  Plateau, 
in  Jackson  and  Marshall  Counties. 

On  most  of  the  mountains  the  soil  is  thin  and  rocky,  supporting  a 
rather  open,  stunted  growth  of  pine,  red  cedar,  oak,  chestnut,  hickory, 
and  other  trees,  and  it  is  on  the  upper  slopes  of  these  ridges  that  the 
Upper  Austral  Zone  reaches  its  southern  limit. 

Tennessee  Valley. 

The  Tennessee  River  flows  through  a  broad,  rolling  valley,  from 
12  to  15  miles  wide,  crossing  the  State  from  the  northeastern  to  the 
northwestern  corner.  Now  largely  cleared  of  timber  and  under  culti- 
vation, this  valley  is  one  of  the  most  important  agricultural  sections 
of  the  State. 

-  Adapted  in  part  from  Mohr's  map  of  the  floral  areas  of  Alabama,  in  "  Plant  Life  of 
Alabama"  (1901),  where  will  be  found  a  more  detailed  discussion  of  the  physiographic 
conditions. 

7 


8  NORTH   AMERICAN   FAUNA.  [No.  45 

Warrior  Basin  and  Tableland. 

The  Warrior  Tableland  comprises  the  elevated  plateau  in  the  north 
central  part  of  the  State,  covering  part  or  all  of  Winston,  Cullman, 
Marshall,  Blount,  Etowah,  Walker,  Jefferson,  Tuscaloosa,  Shelby, 
and  Bibb  Counties.  It  is  a  region  of  low  hills,  interspersed  with 
narrow  valleys,  many  of  .the  streams  flowing  through  deep  gorges 
bordered  with  high  cliffs.  A  large  proportion  of  this  area  is  heavily 
forested,  the  prevailing  trees  being  chestnut  oak  (Quercus  prinus), 
Spanish  oak  {Quercus  digitata),  pines  (Pinus  echinata,  Pinus  taeda, 
Pinus  virginio/nu,  and  Pinus  palustris),  hickories,  chestnut,  and  sour 
gum  {Nyssa  sylvatica) .  On  the  moist  slopes  of  the  ravines  a  dense 
growth  of  shrubs  is  usually  found,  including  laurel  (Kalmia),  aza- 
leas, huckleberries,  and  the  handsome  oak-leaved  hydrangea  (Hy- 
drangea quercifolia). 

Central  Prairie  Belt. 

The  Prairie  Region  of  Alabama  comprises  a  belt  from  35  to  45 
miles  wide  crossing  the  State  from  Sumter  County  to  Eussell  County, 
and  including  also  part  or  all  of  Pickens,  Greene,  Hale,  Perry,  Dallas, 
Autauga,  Lowndes,  Montgomery,  Macon,  and  Bullock  Counties.  It 
consists  of  a  rolling  plain,  bordered  on  the  north  and  south  by  pine- 
covered  hills.  Although  it  contains  extensive  tracts  of  prairie,  it  is 
by  no  means  exclusively  of  this  character,  but  includes  many  small 
areas  of  forested  land,  extensive  wooded  swamps,  and  numerous 
canebrakes. 

Many  varieties  of  trees  are  found  in  this  region,  among  the  more 
prominent  of  which  are  the  post  oak  (Quercus  minor),  Texas  white 
oak  (Quercus  durandi),  water  oak  (Quercus  nigra),  overcup  oak 
(Quercus  lyrata),  pecan,  hickories,  elms,  hawthorns,  red  cedar,  and 
several  species  of  pine  (Pinus  taeda,  Pinus  echinata,  Pinus  palustris, 
and  Pinus  glabra). 

Coast  Pine  Belt. 

The  Coast  Pine  Belt  extends  across  the  State  from  Choctaw  to 
southern  Barbour  and  Henry  Counties,  merging  on  the  north  into 
the  Prairie  Belt  and  on  the  south  into  the  Coast  Plain.  Of  its  floral 
characters,  Mohr  says :  "  Broadly  stated,  it  consists  in  the  increased 
frequency  of  types  which  are  at  home  in  the  Louisianian  Area,  and  in 
distinction  from  regions  farther  north,  of  a  tree  growth  in  which, 
though  otherwise  similar,  the  long-leaf  pine  in  its  highest  development 
predominates.  *  *  *  Fully  one-half  of  the  area  of  this  region  is 
under  cover  of  the  long-leaf  pine  *  *  *."3  Other  characteristic 
trees  and  shrubs  are  the  shortleaf  pine  (Pinus  echinata),  southern 
spruce  pine  (Pinus  glabra),  basswood  (Tilia  heteroplxylla) ,  magnolia 

3  Mohr,  Charles,  Plant  life  of  Alabama,  pp.  106,  110,  1901. 


North  American  Fauna  No.  45,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  II. 


B  I  5 1  03 

Fig.  I. — Eastern  Slopes  of  Sand  Mountain,  from  Trenton  Valley. 

Rocky  escarpment,  heavily  timbered,  except  on  steepest  slopes. 


SI  7307 

Fig.  2. — Gulch  of  Long   Island  Creek,  Sand  Mountain. 

Hardwood  timber  in  the  bottom  of  the  gulch;  shortleaf  pines  (Pinus  virginiana)  on  top  of  plateau. 


North  American  Fauna  No.  45,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  III. 


Fig.  i.— Rock  Ridge  near  Summit  of  Choccolocco  Mountain. 

Longleaf  pine  (Pinus  palustris)  growing  on  summit. 


Fig.  2.-Choccolocco  Valley,  near  Piedmont.    Dugger  Mountain  in 

Distance. 


North  American  Fauna  No.  45,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  IV. 


Fig.  I.— Rocky  Slopes  near  Base  of  Talladega  Mountains. 

Young  growth  of  longleaf  pine  (Pinus  pahistris). 


Fig.  2.— Upper  Slopes  of  Choccolocco  Mountain. 

Timber  chiefly  oaks  and  longleaf  pine. 


North  American  Fauna  No.  45,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  V. 


Fig.    I.— Tennessee    River    near    Bloom  field. 

Rock  bluff  covered  with  dense  growth  of  mixed  timber. 


Fig.  2.-Sipsey    Fork   at    Mouth   of   Bear   Branch. 

Wooded  hills,  with  narrow  valley  bottoms  under  cultivation. 


North  Ame 


rican  Fauna  No.  45,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  V I . 


F|G    |  -pine  Forest  near  Jasper. 


FIG.  2.-TURPENTINE  "ORCHARD"   NEAR    MOBILE. 

Pure  stand  of  longleaf  pine. 


North  American  Fauna  No.  45,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  VII. 


Fig.  I.— Catoma  Swamp,  near  Teasley  Mill. 

Spruce  pine,  beech,  oaks,  hickories,  etc. 


Fig.  2.— River-bottom  Swamp  near  Seale. 

Undergrowth  of  palmetto  (Sabal  minus)  in  hardwood  forest. 


1921.]  PHYSIOGKAPHY  AND   LIFE   ZONES   OF   ALABAMA.  9 

{Magnolia  foetida),  red  bay  (Persea  borbonia),  beech,  water  oak 
{Quercus  nigra),  laurel  oak  {Quercus  laurifolia) ,  storax  bush  {Sty- 
rax  grandiflora) ,  sweet  illicium  {Illicium  floridanum) ,  white  and  red 
flowered  buckeyes,  and  the  Carolina  allspice  {Calycanthus  ftoridus). 

Coast  Plain  or  Gulf  Strip. 

The  Coast  Plain,  as  used  here,  includes  the  Lower  Coast  Pine 
Belt,  the  Coast  Plain,  and  the  Littoral  Belt  of  Mohr's  "Map 
of  the  Floral  Areas  of  Alabama."  It  comprises  the  sand-dune  and 
salt-marsh  areas  of  the  coast  and  outer  islands,  the  coastal  pine  flats, 
the  extensive  river  swamps  around  the  head  of  Mobile  Bay,  and  the 
rolling  sandy  uplands  of  the  southern  tier  of  counties. 

In  the  hill  country,  which  attains  an  altitude  of  250  or  300  feet 
above  sea  level,  the  longleaf  pine  {Pinus  palustris)  is  the  prevailing 
tree,  with  some  loblolly  pine  {Pinus  taeda) ,  interspersed  with  a  rather 
sparse  and  stunted  growth  of  turkey  oak  {Quercus  catesbaei),  blue 
jack  {Quercus  brevifolia),  huckleberry  bushes,  and  the  heatherlike 
shrub  Ceratiola  ericoides.  In  the  depressions  between  the  hills  and 
about  the  heads  of  the  numerous  streams  rising  in  this  area  are  found 
many  small  swamps,  in  which  is  a  dense  growth  of  white  bay  {Mag- 
nolia virginiana) ,  water  oak  (  Quercus  nigra),  laurel  oak  {Quercus 
laurifolia),  southern  black  gum  {Nyssa  biflora),  swamp  maple, 
white  cedar  {Chamaecyparis  thyoides),  pond  cypress  {Taxodium 
ascendens),  and  scattering  pines. 

On  the  flats  along  the  coast  is  found  an  open  forest  of  pines, 
chiefly  the  swamp  pine  {Pinus  elliottii).  In  the  hammocks — slightly 
elevated  tracts  rising  from  the  swampy  flats — occurs  a  profusion  of 
large  timber  trees,  including  the  magnolia  {Magnolia  foetida) ,  beech, 
holly  {Ilex  opaca),  water  oak  {Quercus  nigra),  laurel  oak  {Quercus 
laurifolia),  and  the  magnificent  live  oak  {Quercus  virginiana),  a 
tree  characteristic  of  the  coastal  region.  About  the  borders  of  the 
hammocks  and  in  the  small  swamps  is  found  a  dense  growth  of  shrubs, 
including  the  titi  {Gliftonia  nionophylla)  and  the  leatherwood 
{Cyrilla  racemiflora) . 

Extensive  river  swamps  cover  the  delta  formed  by  the  Mobile  and 
Tensaw  Rivers,  much  of  the  land  being  subject  to  long-continued 
overflows.  Here  grow  magnificent  forests  of  cypress  {Taxodium  dis- 
tichum),  tupelo  gum  {Nyssa  aquatica),  black  gum  {Nyssa  biflora), 
red  maple  {Acer  drummondii) ,  water  hickory  {Hicoria  aquatica), 
water  oak  {Quercus  nigra),  overcup  oak  {Quercus  lyrata),  water  ash 
{Fraxinus  caroliniana) ,  green  ash  {Fraxinus  lanceolata) ,  swamp 
Cottonwood  {Populus  heterophylla) ,  and  other  trees,  all  heavily 
festooned  with  the  long  Spanish  "  moss  "  and  hung  with  tangles  of 
grape  vine  and  other  climbing  plants. 


10  NORTH   AMERICAN   FAUNA.  [No.  45 

On  the  shores  of  the  coastal  bays  and  on  the  outer  islands  are  ex- 
tensive salt  marshes,  largely  occupied  by  the  black  rush  {J uncus 
roemerianus)  and  inhabited  by  great  numbers  of  the  Louisiana  clap- 
per rail  {R alius  crepitans  saturatus) . 

On  Dauphin  Island  is  a  considerable  forest  of  pines  and  hard- 
woods. Petit  Bois  Island  supports  only  a  small  tract  of  pines,  but 
parts  of  the  island  are  covered  with  dense  thickets  of  yaupon  (Ilex 
vomitorla).  The  lower  parts  of  both  Dauphin  and  Petit  Bois 
Islands  are  given  over  to  rolling  sand  dunes,  and  the  shores  are 
bordered  with  extensive  mud  flats. 

LIFE  ZONHS  OF  ALABAMA. 

Only  two  of  the  seven  transcontinental  life  zones  of  North  America 
are  represented  in  Alabama — the  humid  divisions  of  the  Upper  Aus- 
tral and  Lower  Austral  Zones,  known,  respectively,  as  the  Carolinian 
and  Austroriparian  (or  Louisianian),  and  corresponding  in  the  West 
to  the  arid  divisions,  the  Upper  Sonoran  and  the  Lower  Sonoran,  re- 
spectively. The  Lower  Austral  Zone  covers  the  greater  part  of  the 
State  and  is  subdivided  into  two  fairly  well-marked  regions. 

Upper  Austral  Zone. 

The  humid,  or  Carolinian,  division  of  the  Upper  Austral  Zone 
occurs  in  rather  dilute  form  on  the  higher  mountains  in  the  north- 
eastern part  of  Alabama,  reaching  its  southern  limit  in  the  Talladega 
Mountains  of  Clay  County.  This  zone  is  chiefly  confined  to  the 
upper  slopes,  above  1,500  feet  altitude,  and  is  therefore  of  little  agri- 
cultural importance  in  the  State.  Its  most  important  crops  are  hay, 
corn,  oats,  wheat,  tobacco,  cowpeas,  white  potatoes,  sweet  potatoes, 
apples,  pears,  peaches,  and  plums.  A  large  number  of  plants  and  a 
smaller  number  of  birds  and  mammals  occurring  in  this  zone  serve 
to  mark  approximately  its  southern  boundary.  The  most  important 
species  in  Alabama  are  the  following : 

Characteristic  Breeding  Birds — Upper  Austral  Zone. 


Ruffed  grouse   (Bonasa  umbellus  um- 

bellus). 
Sparrow    hawk     (Cerchneis    sparveria 

sparveria )  .* 
Whip-poor-will    (SetocJialchis  vocifera 

vooifera ) . 
Phoebe  (Sayomig  phoebe)* 


Scarlet  tanager  {Piranga  olivacea).* 
Worm-eating     warbler     (Helmitheros 

vermivorus) . 
Ovenbird    (Seiurus  aurocapilliis  auro? 

capilhis). 
Southern  robin    (Planesticus  migrato- 

riits  achrusterus). 


Characteristic  Mammals — Upper  Austral  Zone. 


New  York  weasel  (Mustcla  novebora- 

censis ) . 
Allegheny  cliff  rat  (Neotoma  pcnnsyl- 

■vanica). 


Southern    woodehuck,    or    ground-hoj 
(Marmota  monax  monax).6 


4  Breeds  rarely  in  the  Lower  Austral  Zone. 

5  Occurs  also  in  mountainous  parts  of  the  Lower  Austral  Zone. 


1921.] 


PHYSIOGRAPHY  AND  LIFE   ZONES   OF   ALABAMA. 


11 


Characteristic  Plants — Upper  Austral  Zone.' 


Trees  and  shrubs. 


Hemlock  (Tsuga  canadensis) . 
Butternut  (Juglans  cinerea). 
Cherry  birch  (Betula  tenia). 
Scarlet  oak  (Quercus  cocomea). 
Chestnut  oak  (Quercus  prinus). 
Hairy    syringa     (Philadelphus    hirsu- 

tus). 
Mountain  black  cherry   (Prunus  sero- 

tina  neomontana). 
Clammy  locust  (Robinia  visoosa). 
Rose  acacia  (Robinia  hispida). 
Mountain  privet  (Ilex  monticola). 


Bitter-sweet  (Celastrus  scandens). 

Fetid  buckeye  (Acsculus  glabra). 

Blue  grape  (Vitis  bicolor). 

Sweet-scented  azalea  (Azalea  arbores- 
cens ) . 

Mountain  blueberry  (Vaccinium  palli- 
dum ) . 

Blue  ash  (Eraxinus  quadrangulata) . 

Withe-rod   ( Viburnum  cassinoides ) . 

Bush  honeysuckle  (Diervilla  rivu- 
laris). 


Herbaceous  plants. 


Moccasin  flower  (Cypripedium  acaule). 
Showy  orchid   (Orchis  spectabilis). 
Virginian  spring  beauty  (Claytonia  vir- 

ginica). 
Round-leaved  campion  (Silene  rotundi- 

folia). 
Smooth  meadow-rue  (Thalictrum  dioi- 

cum). 
Sweet  wood-violet  (Viola  leconteana) . 
Pale  violet  (Viola  striata). 
Long-spurred  violet    (Viola  rostrata). 
Colored  willow-herb    (Epilobium  colo* 

ratum). 
Spikenard  (Aralia  racemosa). 
Canada     lovage      (Ligusticum     cana- 

dense ) . 
Sweet  cicely  (Osmorrhiza  claytoni). 
Spotted  wintergreen  (Chimaphila  mac- 

ulata ) . 
Four-leaf       loosestrife       (Lysimachia 

quadrifolia) . 
Mountain  steironema  (Steironema  ton- 
sum  ) . 


Four-leaved  milkweed  (Asclepias  quad- 
rifolia). 

Creeping  purple  phlox  (Phlox  rep- 
tans). 

Virginia  lungwort  (Meriensia  virgin- 
ica). 

Bradbury  horsemint  (Monarda  brad- 
buriana). 

Gerardia  (Agalinis  asperula). 

Narrow-leaved  houstonia  (Eoustonia 
tenui  folia) . 

Upland  boneset  (Eupatorium  sessili- 
folium). 

White  goldenrod  (Solidago  bicolor). 

Broad-leaved  goldenrod  (Solidago  flexi- 
c«  al is). 

Hispid  goldenrod  (Solidago  hispida). 

Slender  aster  (Aster  gracilis). 

Sky-blue  aster  (Aster  azureus). 

Canada  leaf-cup  (Polymnia  canaden- 
sis). 

Mountain  cone-flower  (Rudbeckia  mon- 
ticola). 


Lower  Austral  Zone. 

The  Austroriparian  (Louisianian)  Division  of  the  Lower  Austral 
Zone  occupies  all  the  lowlands  of  Alabama  and  the  lower  slopes  of 
the  mountains  below  1,500  feet  altitude,  thus  including  practically- 
all  the  agricultural  lands  in  the  State.  The  principal  crops  in  this 
zone  are  cotton,  corn,  oats,  sorghum,  peanuts,  sweet  potatoes,  water- 


6  Partial  list. 


12 


NORTH   AMERICAN   FAUNA. 


[No.  45 


melons,  cantaloupes,  strawberries,  peaches,  figs,  and  pecans.  Lumber 
and  turpentine  also  are  important  products. 

A  subdivision  of  the  Lower  Austral  Zone,  known  as  the  Semi- 
tropical  or  Gulf  Strip,  is  recognized  as  occupying  the  southern  tier 
of  counties  from  about  the  latitude  of  Castleberry  southward,  its 
northern  boundary  practically  coinciding  with  that  of  Mohr's 
"  Lower  Coast  Pine  Belt."  In  this  belt  many  varieties  of  subtropi- 
cal fruits  flourish  which  will  not  live  farther  north. 

The  Lower  Austral  Zone  in  Alabama  is  characterized  by  the 
presence  of  a  large  number  of  southern  animals  and  plants,  among 
which  the  following  are  the  mcjst  important : 

Characteristic  Breeding  Birds — Lower  Austral  Zone. 

(a)   Species  Ranging  Nearly  Throughout  the  Lower  Austral  Zone  and  Limited 

Northward  by  It. 


Black  vulture  (Coragyps  urubu  urubu). 
Florida    red-shouldered   hawk    (Buteo 

Uneatus  alleni). 
Florida     barred     owl      (Striw     varia 

alleni). 
Florida  screech  owl   (Otus  asio  asio). 
Southern    hairy     woodpecker     {Dryo- 

bates  villosus  auduboni). 
Downy  woodpecker   {Dryobates  pubes- 

cens  pubescens) . 
Red-cockaded    woodpecker    (Phrenopi- 

cus  borealis). 
Flicker   (Colaptes  auratus  auratus). 
Chuck-will's-widow  (Antrostomus  car- 

olinensis) . 
Florida  blue  jay   (Cyanocitta  cristata 

cristata)? 
Southern        meadowlark        (Sturnella 

magna  argutula). 

(6)  Species  Ranging  North  to  About  the  Middle  of  the  State  (Autauga  and  Hale 

Counties). 


Bachman  sparrow   {Peueaea  aestivalis 

bachmani)  .* 
Alabama    towhee     (Pipilo    erythroph- 

thalmus  canaster) . 
Blue  grosbeak   (Ouiraca  caerulea  cae- 

rulea )  .8 
Swainson       warbler       (Limnothlypis 

swainsonii) . 
Bachman    warbler    {Yermivora    bach- 

manii). 
Yellow-throated     warbler     (Dendroica 

dominiea  dominica).* 
Florida  white-breasted  nuthatch  (Sitta 

cai'olincnsis  carolinensis ) ." 
Brown-headed     nuthatch     (Sitta     pu- 

silla). 


Ground  dove    (Chanuepelia   passerina 

passerina). 
Little  sparrow  hawk   {Cerchneis  spar- 

veria  paula). 
Florida  nighthawk   (Chordeiles  minor 

ehapmani).10 


Florida  red-wing   (Agelaius  phocniecus 

phoeniceus) . 
Loggerhead    shrike    (Lanius    ludovici- 

an us  ludo  vieia nus ) . 
Florida  yellow-throat    (Geothlypis   tri- 

chas  ignota). 


7  Formerly  known  as  Cyanocitta  cristata  flurincola. 

8  Occurs  also  in  the  Upper  Austral  Zone. 

9  Formerly  known  as  Sitta  carolinensis  atkinsi. 

10  Possibly  throughout  the  zone. 


North  American  Fauna  No.  45,  U.  S.  Dept.  Agr.,  Biological  Survey.  PLATE  VI  I  I 


Fig.  I— Sand  Dunes,  Orange  Beach. 

Thickets  of  yaupon  (Ilex  vomitoria)  and  dwarf  live-oak  (Quercus  virginiana  maritima);  home  of 
the  white-fronted  beach  mouse  (Peromyscus  polionotus  albifrons). 


Fig.  2.— Swamp  on  Dauphin   Island,  Overflowed  by  Sand. 

Black  rush  (Juncus  roemerianus)  in  foreground;  swamp  pine  (Pinus  cllinttii)  in  background. 


North  American  Fauna  No.  45,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  IX. 


Fig.  I— Stiggins  Lake,  Tensaw  River  Delta. 

Shores  bordered  with  cypress  (  Taxodium  distichum)  and  various  hardwood  trees. 


Fig.  2.— Marsh  on  Bay  Side  of  Dauphin   Island. 

Dense  growth  of  black  rush  (Juncus  roemerianus). 


1921.] 


PHYSIOGRAPHY   AND  LIFE   ZONES   OF   ALABAMA. 


13 


(c)  Species  Confined  to  the  Semi-tropical,,  or  Gulf,  Strip. 


Louisiana  clapper  rail  (Rallus  crepi- 
tans saturatus). 

Purple  gallinule  (lonornis  martini- 
cus). 

Florida  grackle  (Quiscalus  quiscula 
quiscula). 


Boat-tailed    grackle     (Mcgaquiscalus 

major  major). 
Howell    seaside    sparrow    (Passerher- 

bulus  maritimus  hoirelli).11 
Marian    marsh    wren     (Telmatodytes 

palustris  marianae). 


Characteristic  Mammals — Lower  Austral  Zone. 

(a)   Species  Ranging  Nearly  Throughout  the  Lower  Austral  Zone  and  Limited 

Northward  by  It. 


Carolina  short-tailed  shrew  (Blarina 
brevicauda  carolinensis) . 

Evening  bat   (Nycticcius  humeraUs)  .a 

Alabama  weasel  (Mustela  peninsulae 
olivacea). 

Old-field  mouse  (Peromyscus  poliono- 
tus polionotus). 

Cotton  mouse  (Peromyscus  gossy- 
pinus  subspp.). 

Southern  golden  mouse  {Peromyscus 
nuttalli  aureolus). 


Swamp  rice   rat    (Oryzomys  palustris 

palustris)?2 
Cotton    rat    (Sigmodon    hispidus    his- 

pidus ) . 
Florida  wood  rat   (Neotoma  floridana 

florid  ana ) . 
Alabama  pocket  gopher  (Geomys  tuza 

mobilensis)  .1S 
Swamp    rabbit    (Sylvilagus    aquaticus 

aquations). 


(b)   Species  Confined  to  the  Semitropical,  or  Gulf,  Strip. 


Mahogany  bat   (Nycteris  seminola). 

LeConte  free-tailed  bat  (Nyctinomus 
cynocephalus) ." 

White-fronted  beach  mouse  (Peromys- 
cus polionotus  albifrons). 

Louisiana  muskrat  (Fiber  rivalicius). 


Bayou  gray  squirrel  (Sciurus  caroli- 
n  en  sis  fuliginosus ) . 

Coast  swamp  rabbit  (Sylvilagus  aqua- 
ticus littoralis). 

Marsh  rabbit  (Sylvilagus  palustris 
palustris ) . 


Characteristic  Plants — Lower  Austral  Zone.15 

(o)   Species  Ranging  Nearly  Throughout  the  Lower  Austral  Zone  and  Limited 

Northward  by  It. 


Trees  and  shrubs. 


Loblolly  pine  (Pinus  taeda). 
Longleaf  pine   (Pinus  palustris). 
Spruce  pine  (Pinus  glabra). 
Swamp  cypress  (Taxodium  distichum). 
Saw  palmetto  (Serenoa  scrrulata). 
Pecan  (Hicoria  pecan). 
Nutmeg  hickory    (Hicoria  myristicae- 
formis). 


Water  hickory  (Hicoria  aquatica). 
Southern    wax    myrtle    (Myrica    ceri- 

fcra). 
Overcup  oak  (Q  it  ere  us  lyrata). 
Texas  white  oak  (Quercus  durandi). 
Turkey  oak  (Quercus  catesbaci). 
Swamp   Spanish  oak    (Quercus   pago- 

dae  folia). 


11  Cf.  Griscom,  Ludlow,  and  J.  T.  Nichols,  Abstr.  Proc.  Linnaean  Soc.  New  York,  no.  32, 
p.  22,  Nov.  S,  1920. 

12  Occurs  locally  in  the  Upper  Austral  Zone. 

13  Ranges  north  to  Tuscaloosa  County  only. 

14  Ranges  north  to  Greensboro. 

15  Partial  list. 


14 


NORTH   AMERICAN   FAUNA. 


[No.  45 


Water  oak  (Quercus  nigra). 
Laurel  oak  (Quercus  laurifoHa). 
Upland    willow    oak     (Quercus    brevir 

folia). 
Winged  elm  (Ulmus  alata). 
Planer  tree  (Planera  aquatica). 
Southern  hackberry   (Ccltis  mississip- 

piensix). 
Magnolia   (Magnolia  foetida). 
Sweet-shrub ;     calycanthus     ( Calycan- 

thus  floridus). 
Red  bay  (Per sea  borbonia). 
Swamp  red  bay  (Persea  pubescetis) . 
Prairie  plum  (Primus  umbellata) .    / 
Southern     prickly-ash     (Zantho.ryhun 

clava-herculis ) . 
Sebastiana   (Sebastiana  ligustrina). 
Leatherwood   (Cyrilla  racemiflora) . 
Tall  inkberry  (Ilex  lucida). 
Pale-bark  maple  (Acer  Icucoderme) . 
Florida  maple  (Acer  floridanum). 
Drummond    red    maple    (Acer    drum- 

mondU). 
Southern  basswood  (Tilia  australis). 
Virginia    stewartia    (Stetoartia   mala- 

codendron) . 


Southern  St.  Peterswort  (Ascyrum  hy- 

perieoides) . 
Southern  black  gum  (Nyssa  biflora). 
Tupelo  gum  (Nyssa  aquatica). 
Fetterbush  (Pieris  nitida). 
Evergreen  blueberry  (Vaccinium  myr- 

sinites). 
Elliott  blueberry  (Vaccinium'  clliottii). 
Fuscous  blueberry    (Vaccinium  fusca- 

tum). 
Southern      buckthorn      (Bumelia      ly- 

cioides ) . 
Southern  silver-bell  tree  (Halesia  dip- 

tera). 
Powdery     storax      (Styrax     pulveru- 

lenta). 
Large-flowered  storax  (Styrax  grandi- 

folia). 
Water  ash  (Fraxinus  caroliniana) . 
Devil- wood   (Osmanthus  americana). 
Yellow  jessamine  (Gelsemium  semper- 

virens). 
Catalpa   (Catalpa  catalpa). 
Opossum  haw   (Viburnum  nitidum). 


Herbaceous  plants. 


Cane  (Arundinaria  macrospcrma). 
Downy  eriogonum   (Eriogonum  tomen- 

tosum). 
Red-flowered      pitcher-plant       (Sarra- 

cenia  rubra). 
Short-leaved    sundew    (Drosera   brevi- 

folia). 
Sensitive  pea   (ChamaecHsta  nniltiphi- 

nata). 
Lanceolate  false-indigo   (Baptisia  lan- 

ceolata). 
Round-leaved     rattlebox      (Crotalaria 

rolundifolia). 
Spreading  lupine    (Lupinus  diffusus). 
Hairy  lupine  (Lupinus  vVlosus). 
Pine -barren    prairie  -  clover    (Kuhni- 

stcra  pinnata) . 
Wild  indigo   (Indigofera  caroliniana). 
Hispid  goat's-rue  (Cracca  hispidula). 
Narrow-leaved   tick-trefoil    (Meibomia 

tcnuifolia). 
La-rge  -  flowered     milkwort     (Poly gala 

grandiflora) . 
Queen's-delight    (StiUingia  sylvatica). 
Rough    rose-mallow    (Hibiscus   acule- 

aHis). 


Carolina  violet  (Viola  Carolina). 

Lance-lea ved  meadow-beauty  (Rhexia 
lanccolata). 

Deer  grass  (Rhexia  glabella). 

Hoary  ludwigia   (Ludwigia  pilosa). 

Miterwort   ( GynOdton  inn  mitreola ) . 

Rabbit's-milkweed  (Asclepias  humi- 
strata ) . 

Carolina  morning-glory  (Ipomaea  caro- 
liniana ) . 

Low  breweria  (Breiceria  huinistrata). 

Florida  phlox   (Phlox  floridana). 

Spanish   "cypress"    (Gilia  rubra). 

Carolina  vervain  (Verbena  caro- 
Unensis). 

Swamp  basil  (Hyptis  rugosa) . 

Carolina  thyme  (Clinipodmm  Caro- 
linian um). 

Azure  salvia  (Salvia  azurea). 

Fascicled  gerardia  (Agalinis  fasei- 
culata). 

Southern  bluehearts  (Buchnera  elon- 
gata). 

Fimbriated  ruellia   (Ruellia  humilis). 

Southern  bluets  (Houstonia  pusilla). 


1921.] 


PHYSIOGRAPHY  AND  LIFE   ZONES   OF   ALABAMA. 


15 


Narrow  wild  lettuce    (Lactuca  gram- 

inifolia) 
Narrow-leaved      vernonia      ( Yernonia 

angustifolia). 
Blazing-star  ( Lacinaria  elegans). 
Vanilla  plant    (Trilisa  odoratissima) . 
Hoary-leaved  golden  aster  (Chrysopsis 

trichophylla) . 
Hoary      golden      aster       (Chrysopsis 

pilosa). 
Narrow-leaved     goldenrod      {SoUdago 

angustifolia). 
Short-leaved       goldenrod       ( SoUdago 

brachyphylla) . 

(b)  Species  of  the  Subtropical  Division,  or  Gulk  Strip,  of  the  Lower  Austral,  Zone. 

Trees  and  shrubs. 


Mouse-ears    (Sericocarpus   bifoliatus). 
Wandlike  aster  (Aster  purpuratus). 
Seaside  aster  (Aster  exilis). 
Southern     daisy     fleabane     (Erigeron 

quercifolius) . 
Rosinweed    (Silphium  asteriscus). 
Oxeye   (Heliopsis  minor). 
False       sunflower        (Tetragonotlucn. 

Jtelianthoides). 
Short-leaved       heleniuin       (Helenium 

brevifolium). 


Swamp  pine  (Pivus  elUottii). 

Sand  pine  (Pinus  clausa). 

Pond  cypress   (Taxodium  asecndens). 

Coast  red  cedar  (Juniperus  barbaden- 

sis). 
Bartrani      wax-myrtle      (Myrica      in- 

odora). 
Dwarf    live-oak    (Quercus    Virginian  a 

maritima). 
Twin  live-oak  (Quercus  gcminata) . 
Seaside     scrub-oak     (Quercus    niyrti- 

folia). 
Narrow-leaved        papaw         (Asimina 

angustifolia) . 

Hcrbaccou 

Elliott  yellow-eyed  grass  (Xyris  el- 
Uottii). 

Creeping  orchis  (Habenaria  repens). 

Small  yellow  pondlily  (Nymphaea 
chartacea ) . 

Sea  rocket  (Cakile  chapmani) . 

Parrot-beak  (Sarracenia  psittacina) . 

Drummond  trumpet-leaf  (Sarracenia 
drummondi). 

Capillary  sundew  (Drosera  capillar  is). 

Pursh  rattlebox   (Crotalaria  purshii). 

Slender  prairie-clover  (Kuhnistera 
gracilis). 

Goat's-rue  (Cracca  smallii). 

Prostrate  goat's-rue  (Cracca  chryso- 
phylla ) . 

Acute-leaved  vetch   (Vicia  acutifoUa). 

Florida  milk-pea  (Galactia  floridana). 

Erect  milk-pea  (Galactia  erecta). 


Deer    plum    (Chrysobalanus    oblongh- 

folius). 
Ceratiola   (Ceratiola  ericoides). 
Titi   (Cliftonia  monophylla) . 
Low     andromeda     (Pieris    phillyrcae- 

folia ) . 
Hoary  huckleberry  (Gaglussacia  fron- 

dosa  tomentosa). 
Scarlet     balm      (Clinopodium      cocci- 

■ncum). 
Seaside  balm    (Conradina  canescens). 
Carolina     boxthorn      (Lycium     caro- 

linianum). 

s  plants. 

Pine-barren  pitcheria  (Pitcheria  galac- 
tioides). 

Chapman  milkwort  (Polygala  chap- 
mani). 

Dwarf  St.  Peter's- wort  (Ascyrum  pu- 
ns Hum). 

Opaque  St.  John's-wort  (Hypericum 
opacum ). 

Seaside  rock-rose  (Helianthcmum  are- 
nicola). 

Divaricate  pinweed  (Lechea  divari- 
cata). 

Crowfoot  cactus  (Opuntia  pes-corvi). 

Swamp  meadow  -  beauty  (Rhexia 
stricta). 

Narrow-leaved  gaura  (Gaura  angusti- 
folia). 

Large-leaved  sabbatia  (Sabbatia  ma- 
crophylla). 


16 


NORTH   AMERICAN   FAUNA. 


[No.  45 


Gentian    sabbatia    (Lapithea   gentian- 

oides). 
Michaux     milkweed      (Aselepias     mi- 

cha/uxn). 
Goat's-foot     morning-glory     (Ipomoea 

pcs-caprae) . 
Seaside    morning-glory     (Ipomoea    lit- 

toralis). 
Fog-fruit  (Lippia  nodi  flora). 
Large-flowered    pennyroyal    (Diceran- 

dra  lincari folia). 
Narrow-leaved  ground-cherry  (Physalis 

angustifolia) . 
Florida  toad-flax   (Linaria  floridaha). 
Rough     hedge-hyssop     (Soplironanthe 

hispida). 
Leafless  gerardia  (Agalinis  aphylla). 
Night-blooming  ruellia   (Ruellia  nocti- 

flora). 


Round-leaved  houstonia  (Houstonia 
procumbens). 

Short-leaved  lobelia  {Lobelia  brevi- 
folia ) . 

Azure-flowered  stokesia  (Stokesia 
laevis). 

Slender  blazing-star  (Lacinaria  gra- 
cilis). 

Large-flowered  golden  aster  (Chrysop- 
sis  oligantha). 

Few-flowered  goldenrod  ( Chrysoma 
pauciflosculosa) . 

Small-leaved  aster  (Aster  adnatus). 

Rayless sunflower  (Hclianthus  radula). 

Lance-leaved  Indian  plantain  (Mesa- 
denia  lanceolata ) . 

Smooth  thistle  (Cirsium  nuttalli). 

LeConte  thistle  (Cirsium  lecontei). 


PART  II.  REPORT  ON  THE  MAMMALS  OF  ALABAMA. 

HISTORY  OF  ALABAMA  MAMMALOGY. 

Alabama  was  visited  in  the  early  years  of  its  settlement  by  num- 
bers of  travelers  and  a  few  naturalists,  some  of  whom  have  published 
accounts  of  their  journeys,  but  the  references  in  their  works  to  mam- 
mals are  for  the  most  part  brief  and  incidental. 

Apparently  the  first  naturalist  to  visit  the  State  was  William 
Bartram,  who,  in  the  summer  and  fall  of  1776  (or  possibly  1777,  the 
date  not  being  clear  from  his  narrative),  in  the  course  of  extended 
travels  in  the  Southern  States  passed  through  Alabama  from  the  old 
Muscogee  town  of  Uche,  en  the  "  Chata  Uche  "  River,  to  Tallassee, 
on  the  Tallapoosa  River,  thence  southward  along  the  general  course 
of  the  Alabama  River  to  "  Taensa  "  and  Mobile.  His  narrative,1 
although  replete  with  interesting  descriptions  of  the  flora,  contains 
only  a  few  brief  references  to  the  larger  mammals,  such  as  wolves, 
bears,  "tygers"  (cougars),  and  deer.  In  1820  Adam  Hodgson  made 
an  extended  journey  through  the  Southern  States,  crossing  Alabama 
twice — first  from  Ouchee  Bridge,  in  Russell  County,  to  Blakely  and 
Mobile,  and  later  from  Franklin  County  eastward  to  Madison  County, 
via  Tuscumbia,  Muscle  Shoals,  xYxhens,  and  Huntsville.  In  his  nar- 
rative he  refers  casually  to  "  panthers  "  (cougars) ,  gray  foxes,  and 
bears.2  In  1830  James  Stuart  journeyed  across  Alabama  from  Fort 
Mitchell  to  Montgomery,  thence  to  Mobile.  Apparently,  the  only 
mammals  which  attracted  his  attention  were  deer,  which  he  mentions 
incidentally.3  Two  years  later,  in  1832,  C.  D.  Arfwedson  covered 
practically  the  same  route  and  likewise  in  his  narrative  mentions 
only  deer.4  In  1856  Charles  Lanman  published  an  account  of  his 
"  Adventures,"  in  which  are  included  four  chapters  on  Alabama,  with 
a  few  casual  references  to  mammals.5 

Audubon  and  Bachman,  in  their  monumental  work  on  the  North 
American  quadrupeds6 — the  first  and  in  some  respects  the  best  ever 
published   on   the  subject— have   given   us   many   excellent  biogra- 

1  Bartram,  William,  Travels,  pp.  520,  London,  1792. 

-  Hodgson,  Adam,  Letters  from  North  America,  vol.  1,  pp.  117-154,  262-27.%  London, 
1824. 

3  Stuart,  James,  Three  years  in  North  America,  vol.  2,  pp.  164-226,  Edinburgh,  1888. 

4  Arfwedson,  C.  D.,  The  United  States  and  Canada,  vol.  2,  pp.  1-47,  London,  1834. 

6  Lanman,  Charles,  Adventures  in  the  wilds  of  the  United  States  and  British  American 
Provinces,  vol.  2,  pp.  146-1S8,  1856. 

6  Audubon,  J.  J.,  and  John  Bachman,  Quad.  North  ximer.,  3  vols.,  New  York,  1846-1854. 

49663°— 21 2  17 


18  NORTH   AMERICAN   FAUNA.  [No.  45 

phies  of  southern  mammals,  largely  based  on  Bachman's  careful 
observations  in  South  Carolina,  but  containing  numerous  incidental 
references  to  Alabama  mammals.  In  Baird's  great  work  on  North 
American  mammals7  only  a  few  Alabama  specimens  are  listed. 
Gosse,  in  his  "  Letters," s  referred  briefly  to  a  number  of  mammals 
and  gave  interesting  accounts  of  deer  hunting  and  of  a  nocturnal 
"  possum  "  hunt. 

Later  references  to  Alabama  mammals  are  found  chiefly  in  mono- 
graphs and  technical  revisions,  and  as  a  rule  do  not  treat  at  all  of 
habits  or  distribution.  In  1909,  however,  the  present  writer  pub- 
lished a  paper  containing  brie^f  notes  on  the  distribution  of  southern 
mammals  based  on  a  field  trip  made  during  the  summer  and  fall  of 
1908.  This  paper — apparently  the  first  local  list  from  the  State — 
contains  records  of  20  species  occurring  in  Alabama.9  In  1916  John 
H.  Wallace,  jr.,  published  in  his  Fifth  Annual  Eeport  of  the  Depart- 
ment of  Game  and  Fish  of  the  State  of  Alabama  a  list  of  the  mam- 
mals of  Alabama,  comprising  50  species.  The  paper  is  evidently 
based  on  a  nominal  list  furnished  by  the  Biological  Survey;  the 
annotations  are  descriptive  and  biographical  and  contain  no  infor- 
mation as  to  distribution. 

The  present  report,  resulting  from  the  field  investigations  carried 
on  by  the  Biological  Survey  from  1908  to  1916,  comprises  an  an- 
notated list  of  65  forms,  10  not  heretofore  recorded  from  the  State 
and  3  new  to  science.10 

ANNOTATED  LIST  OF  MAMMALS. 

Order  MAESUPIALIA. 

Family  DIDELPHIIDAE:  Opossums. 

Didelphis  virginiana  virginiana  Kerr. 

Virginia  Opossum. 

Didelphis  virginiana  Kerr,  Animal  Kingdom,  p.  193,  1792. 

Opossums  are  common  and  generally  distributed  throughout  Ala- 
bama, dwelling  chiefly  in  timbered  regions,  either  in  swampy  bottom 
lands  or  the  drier  upland  woods  and  in  ravines  among  the  hills. 
Two  specimens  from  Ardell  and  one  from  Leighton  are  typical 
virginiana,  as  was  also  a  captive  specimen  examined  at  Piedmont, 
but  there  are  no  specimens  available  to  show  where  this  race  merges 
into  pigra  of  the  coast  region.     The  species  is  reported  from  Sand 

7Baird,  S.  F.,  Mamm.  North  Amer.  :  Rept.  Expl.  Surv.  R.  R.  Tac,  vol.  8,  1857. 

8  Gosse,  P.  EL,  Letters  from  Alabama,  pp.  306,  London,  1859. 

B  Howell,  A.  H.,  Notes  on  the  distribution  of  certain  mammals  in  the  southeastern  United 
States:  Proc.  Biol.  Soc.  Washington,  vol.  22,  pp.  55-68,  1909. 

10  The  new  forms  are  Fcalopus  aquaticus  hoioelH  Jackson,  Mustela  peninsulae  olivacea 
Howell,  and  Glaucomys  volans  saturatus  Howell. 


1921.]  MAMMALS   OF   ALABAMA OPOSSUMS.  19 

Mountain,  Woodville,  Elkmont,  Fort  Payne,  Mount  Weogufka,  Nat- 
ural Bridge,  Greensboro,  Wilsonville,  Talladega  Mountains,  Au- 
burn, Tidewater  (Tuscaloosa  County),  and  other  places. 

The  'possum  is  a  stupid,  clumsy  creature,  with  scarcely  enough 
sense  to  keep  out  of  reach  of  a  cur  dog;  it  often  blunders  into  any 
sort  of  a  trap.  Young  individuals  are  frequently  taken  in  spring- 
wire  rat  traps  set  for  small  mammals,  and  the  adults  in  small  steel 
traps.  Opossum  hunting  is  a  favorite  sport  in  the  South,  and  thou- 
sands of  the  animals  are  captured  every  season  by  local  hunters 
without  seriously  reducing  their  numbers.  This  is  due,  doubtless, 
to  the  great  fecundity  of  the  species.  It  is  reported  to  breed  three 
or  more  times  in  a  season,  usually  (according  to  Bachman)  in  March, 
May,  and  July,  and  to  produce  from  6  to  13  young  in  a  litter. 

The  animals  are  chiefly  nocturnal,  remaining  during  the  daytime 
in  their  retreats  in  hollow  logs  or  in  old  nests  of  the  gray  squirrel  or 
the  Florida  wood  rat.  At  Leighton,  in  July,  1911,  while  hunting 
for  wood  rats  along  the  osage-orange  hedges,  the  writer  poked  a  full- 
grown  opossum  out  of  a  large  rat  nest  10  feet  up  among  the  thorny 
branches  of  the  hedge  trees. 

Audubon  and  Bachman  describe  the  feeding  habits  of  the  opossum 
as  follows: 

It  enters  the  cornfields  (maize),  crawls  up  the  stalks,  and  sometimes  breaks- 
them  down  in  the  manner  of  the  raccoon,  to  feed  on  the  young  and  tender  grains ; 
it  picks  up  chestnuts,  acorns,  chinquapins  and  beechnuts,  and  munches  them 
in  the  manner  of  the  bear.  "We  have,  on  dissection,  ascertained  that  it  had  de- 
voured blackberries,  whortleberries,  and  wild  cherries,  and  its  resort  to  the 
persimmon  tree  is  proverbial.  It  is  also  insectivorous,  and  is  seen  scratching  up 
the  leaves  in  search  of  worms  and  the  larvae  of  insects,  of  which  it  is  very  fond. 
In  early  spring  it  lays  the  vegetable  kingdom  under  contribution  for  its  support, 
and  we  have  observed  it  digging  up  the  roots  of  the  small  atamasco  lily 
(Zepherina  atamasco),  and  the  young  and  tender  shoots  of  the  China  brier 
(Smilax  rotund i folia) ,  as  they  shoot  out  of  the  ground  like  asparagus.  It  is 
moreover  decidedly  carnivorous,  eating  young  birds  that  it  may  detect  on  the 
ground,  sucking  the  eggs  in  all  the  partridge,  towhee-bunting  and  other  nests 
it  can  find  in  its  persevering  search.  It  destroys  mice  and  other  rodentia, 
and  devours  whole  broods  of  young  rabbits,  scratching  about  the  nest  and 
scattering  the  hair  and  other  materials  of  which  it  was  composed.  "We  have 
observed  it  squatting  in  the  grass  and  brier  thickets  in  Carolina,  which  are  the 
common  resort  of  the  very  abundant  cotton  rat  (Siamodon  hispidum),  and 
from  patches  of  skin  and  other  mutilated  remains  we  satisfied  ourselves  that 
the  opossum  was  one  among  many  other  species  designed  by  Providence  to  keep 
in  check  the  too  rapid  increase  of  these  troublesome  rats.  We  must  admit  that  it 
sometimes  makes  a  sly  visit  to  the  poultry  house,  killing  a  few  of  the  hens  and 
playing  havoc  among  the  eggs.  The  annoyances  of  the  farmer  however  from 
this  mischievous  propensity,  are  not  as  great  as  those  sustained  from  some  of 
the  other  species,  and  can  not  for  a  moment  be  compared  with  the  destruction 
caused  by  the  weasel,  the  mink,  or  the  skunk." 

11  Audubon  and  Bachman,  Quad.  North  Amer.,  vol.  2,  pp.  112-113,  1851. 


20  NORTH   AMERICAN   FAUNA.  [No.  45 

Didelphis  virginiana  pigra  Bangs. 

Florida  Opossum. 

Didelphis  virginiana  pigra  Bangs,  Proc.  Boston  Soc.  Nat.  Hist.,  vol.  28,  p.  172, 
1S98. 
The  Florida  opossum,  characterized  by  darker  colors,  with  less 
white  on  the  toes,  occupies  the  southern  part  of  the  State.  Two  speci- 
mens from  Castleberry  and  one  from  Bon  Secour  are  referable  to  this 
subspecies,  but  the  northern  limits  of  its  range  are  at  present  uncer- 
tain. Allen  refers  four  skulls  from  Sylacauga  to  this  form.  Re- 
ports of  the  occurrence  of  opossums  at  Seale,  Hayneville,  Catoma 
Creek  (Montgomery  County),  Myrtlewood,  Thomasville,  Carlton, 
Mobile,  and  Alabama  Port  should  probably  be  assigned  to  the  present 
form. 

Order  INSECTIVORA. 

Family  TALPIDAE :  Moles. 

Scalopus  aquaticus  howelli  Jackson. 

Howell  Mole. 

Scalopus  aquaticus  hoirclU  Jackson,  Proc.  Biol.  Soc.  Washington,  vol.  27,  p.  19, 
1914. 

The  Howell  mole,  which  is  a  form  of  the  common  mole,  ranges  over 
practically  the  whole  State  and  inhabits  a  great  variety  of  situations. 
Sandy  or  loamy  soil  is  preferred  by  the  mole,  and  its  wanderings  take 
it  into  meadows,  gardens,  cultivated  fields,  river  bottoms,  mountain 
slopes,  and  both  coniferous  and  hardwood  forests.  The  tunnels  are 
usually  near  the  surface  and  appear  as  a  network  of  tortuous  ridges 
or  sometimes  as  a  single  ridge  running  approximately  straight  for 
some  distance.  During  periods  of  drought  moles  burrow  deeper  and 
at  times  almost  desert  the  surface  ridges.  They  practically  never 
leave  their  burrows  and  when  forcibly  brought  to  the  surface  at 
once  seek  safety  by  burrowing  again  into  the  soil.  Although  pos- 
sessing rudimentary  eyes,  moles  probably  are  unable  to  distinguish 
objects  by  sight,  but  they  may  be  able  to  perceive  light  from  darkness. 

The  mole  is  considered  a  pest  by  most  farmers  and  is  accused  of 
destroying  newly  planted  corn  and  injuring  potatoes  and  other  field 
crops,  as  well  as  garden  bulbs.  Its  diet,  however,  is  composed  largely 
of  insects  and  earthworms,  and  much  of  the  damage  attributed  to  it 
is  really  done  by  pine  mice,  which  follow  its  tunnels  and  attack  the 
plants  or  seeds  near  which  they  lead.  It  is  true,  nevertheless,  that 
moles  do  cause  both  annoyance  and  damage  by  uprooting  lawns  and 
by  tunneling  among  the  roots  of  plants,  thereby  aiding  the  incursions 
of  injurious  rodents. 


1921.]  MAMMALS   OF   ALABAMA SHREWS.  21 

The  form  of  the  common  mole  inhabiting  Alabama  is  intermediate 
in  size  between  the  eastern  mole  (Scalopus  aquaticus  aquations)  and 
the  Florida  mole  (Scalopus  a.  australis),  but  it  is  usually  paler  than 
either,  with  a  flat  skull  and  a  long,  narrow  rostrum.  Specimens  have 
been  taken  at  Huntsville,  Sand  Mountain  (near  Carpenter),  Auburn, 
Ardell,  Autaugaville,  Greensboro,  Cottondale,  Eutaw,  and  Castle- 
berry.  Evidence  of  the  presence  of  moles  has  been  noted  at  Elkmont, 
Natural  Bridge,  Talladega  Mountains,  Seale,  Dothan,  Carlton,  Mo- 
bile, Bayou  Labatre,  Orange  Beach,  and  many  other  places. 

Family  SORICIDAE:  Shrews. 

Blarina  brevicauda  carolinensis  (Bacliman). 

Carolina  Short-tailed  Shbew. 

Sorex  carolinensis  Bacliman,  Journ.  Acad.  Nat.  Sci.  Philadelphia,  vol.  7,  pt.  2, 
p.  366,  1837. 

The  Carolina  short-tailed  shrew  is  the  largest  of  the  three  species 
of  shrews  found  in  the  State ;  it  occurs  in  all  sections,  but  less  abun- 
dantly in  the  south.  In  Bear  Swamp,  near  Autaugaville,  a  number 
of  specimens  were  trapped  by  the  writer  around  rotten  logs  in  the 
swamp,  and  at  York  a  number  were  secured  at  rotten  logs  in  weedy 
fields.  In  Bucks  Pocket,  on  Sand  Mountain,  two  were  caught  in  the 
same  spot  beside  a  log  in  a  wooded  ravine.  L.  J.  Goldman  found  the 
animals  plentiful  in  pine  woods  at  Ardell,  and  at  Greensboro,  Jack- 
son, and  Seale  individuals  have  been  taken  in  fields  of  broom  sedge. 
Single  specimens  have  been  taken  also  by  J.  L.  Peters,  at  Alabama 
Port,  and  by  H.  P.  Loding,  at  Spring  Hill. 

In  the  shrews  the  senses  of  hearing,  smell,  and  touch  are  acute, 
while  that  of  sight  is  practically  useless,  serving  apparently  only  to 
distinguish  light  from  darkness.  The  little  creatures  are  exceedingly 
fierce  and  voracious,  easily  capturing  and  killing  animals  larger  than 
themselves. 

The  present  species,  like  most  of  its  tribe,  is  almost  wholly  noc- 
turnal. It  lives  chiefly  in  hollow  logs  and  stumps  and  in  underground 
burrows,  usually  in  moist  or  peaty  soil.  It  also  makes  shallow  run- 
ways under  the  surface  vegetation  and  uses  the  burrows  and  runways 
of  other  animals. 

Studies  of  the  northern  race  of  this  species  (Blarina  brevicauda 
brevicauda),  by  A.  Franklin  Shull,  showed  that  its  food  consists 
mainly  of  meadow  voles  (Microtus),  mice,  insects,  earthworms,  and 
land  snails.  The  snails  were  hoarded  in  little  piles  near  the  entrances 
to  the  burrows,  and  also  moved  back  and  forth  into  the  galleries. 
The  burrows  were  found  to  be  25  to  30  millimeters  in  diameter,  enter- 
ing the  ground  at  a  steep  angle,  and  extending  from  15  to  40  centime- 


22  NORTH   AMERICAN   FAUNA.  [No.  45 

ters  (6  to  15f  inches).  The  nests  were  "  usually  made  of  grass,  sedge, 
and  leaves  of  nettle,  goldenrod,  or  ash,  arranged  in  the  form  of  a 
hollow  ball,  the  shell  of  which  was  1  to  3  cm.  thick." 12 

Bachman  mentions  having  found  two  nests  of  this  species  composed 
of  root  fibers  and  withered  blades  of  grasses  about  a  foot  beneath  the 
surface.  One  nest  contained  five,  the  other  six  young.  On  another 
occasion  in  one  of  the  tunnels  of  the  mole  he  found  "  a  small  cavity 
containing  a  hoard  of  coleopterous  insects,  principally  composed  of  a 
rare  species  (Scarabaeus  tityus),  fully  the  size  of  the  animal  itself; 
some  of  them  were  nearly  consumed,  and  the  rest  mutilated,  although 
still  living."  13  , 

Cryptotis  parva   (Say). 

Least  Shrew. 
Sorex  parvus  Say,  Long's  Exped.  Rocky  Mountains,  vol.  1,  p.  163,  1823. 

The  least  shrew  is  the  smallest  of  the  short-tailed  shrews,  and  but 
little  larger  than  the  Bachman  shrew  (Sorex  longirostris) .  Although 
few  specimens  have  been  taken,  it  is  apparently  generally  distributed 
over  most  of  the  State  and  in  some  localities  is  not  uncommon.  In 
the  Merriam  collection  is  one  taken  at  Mobile  in  1890  by  the  late 
Dr.  Charles  Mohr.  E.  Gr.  Holt  secured  seven  specimens,  December 
14,  1911,  from  beneath  an  overturned  stack  of  old  alfalfa  hay  in  an 
open  field  at  Barachias.  H.  P.  Loding  captured  two  around  manure 
piles  in  his  garden  in  the  suburbs  of  Mobile.  Three  skulls  of  this 
species  were  found  in  the  stomachs  of  two  barred  owls  (Strix  varia 
alleni)  killed  at  Autaugaville  in  December,  1911.  Specimens  have 
been  trapped  on  Sand  Mountain  (near  Carpenter),  on  Cane  Creek, 
Marshall  County  (near  Oleander),  and  at  Woodville  and  Alabama 
Port. 

Kelatively  little  is  known  about  the  habits  of  this  little  shrew.  It 
is  believed  to  be  partly  diurnal,  and  is  an  inhabitant  of  dry,  grassy 
fields,  rather  than  the  woods  or  marshes  preferred  by  most  other 
shrews.  It  is  frequently  taken  in  runways  of  cotton  rats  (Sigmodon) 
and  may  also  make  small  runways  of  its  own.  A  nest  found  under 
a  log  at  Victoria,  Texas,  by  J.  D.  Mitchell  was  built  of  coarse,  broad- 
leaved  grass  and  lined  inside  with  fine  grass,  the  outside  neatly  and 
strongly  woven. 

Sorex  longirostris  Bachman. 

Bachman  Shrew. 

Sorex  longirostris  Bachman,  Journ.  Acad.  Nat.  Sci.  Philadelphia,  vol.  7,  pt.  2, 
p.  370,  1837. 

The  Bachman  shrew,  a  tiny  long-tailed  species,  even  smaller  in 
bulk  than  the  least  shrew  (Cryptotis  parva),  is  the  smallest  mammal 

12  Shull,  A.  Franklin,  Amer.  Nat.,  vol.  41,  pp.  495-522,  1907. 

13  Audubon  and  Bachman,  Op.  cit.,  vol.  2,  p.  177. 


1921.]  MAMMALS   OF   ALABAMA BATS.  23 

found  in  Alabama,  and  likewise  the  rarest.  It  is  known  only  from 
a  single  specimen  taken  from  the  stomach  of  a  barred  owl  killed  by 
Lewis  S.  Golsan  on  the  borders  of  Bear  Swamp,  Autauga  County, 
December  1,  1911.  The  writer,  in  company  with  Mr.  Golsan,  trapped 
carefully  for  several  days  in  and  around  Bear  Swamp,  where  the 
owl  responsible  for  the  single  Alabama  specimen  was  killed,  but  no 
shrews  of  this  genus  were  secured. 

Originally  described  from  the  lower  Santee  River,  South  Carolina, 
the  species  has  since  been  taken  in  North  Carolina,  Virginia,  Mary- 
land, Indiana,  Illinois,  and  Georgia.  Although  ranging  over  a  wide 
area,  it  is  apparently  uncommon  or  else  difficult  to  trap,  since  com- 
paratively few  specimens  have  been  taken. 

Little  is  known  of  the  habits  of  this  shrew,  but  apparently  they 
are  similar  to  those  of  the  other  long-tailed  shrews,  which  are  found 
chiefly  in  moist  woodland,  living  in  or  beneath  rotten  logs  or  stumps 
or  in  crevices  of  rocks.  One  of  the  original  specimens,  described  by 
Bachman,  was  found  by  laborers  while  digging  a  ditch  through 
grounds  nearly  overflowed  with  water.  Another  was  taken  from 
the  throat  of  a  hooded  merganser,  presumably  killed  in  a  swamp  or 
rice  marsh.14  In  southern  Illinois,  Edmund  Heller  secured  a  number 
of  specimens  in  low,  swampy  woods ;  but  Brimley,  at  Raleigh,  North 
Carolina,  states  that  the  species  is  "  found  on  comparatively  high 
ground,  not  in  swamps  nor  on  the  edges  of  them     *     *     *."  15 

Order  CHIROPTERA. 

Family  VESPERTILIONIDAE:  Typical  Bats. 

Myotis  grisescens  Howell. 

Gkay  Bat. 

Myotis  grisescens  Howell,  Proc.  Biol.  Soc.  Washington,  vol.  22,  p.  46,  1909. 

The  gra}^  bat,  a  medium-sized  species,  is  abundant  at  several  locali- 
ties along  the  Tennessee  River,  but  has  not  been  taken  elsewhere 
in  Alabama.  It  was  first  discovered  in  Nickajack  Cave,  Tennessee, 
which  is  in  the  north  face  of  Sand  Mountain,  about  half  a  mile  north 
of  the  Alabama-Georgia-Tennessee  boundary,  where  in  August,  1908. 
the  writer  collected  the  series  from  which  the  species  was  described.10 
Colonies  have  since  been  found  near  Fort  Deposit,  on  the  Tennessee 
River,  and  at  Rogersville. 

The  gray  bat  lives  in  caves,  both  winter  and  summer,  usually  in 
large  colonies.     In  Nickajack  Cave  large  clusters  hung  from  the 

14  Audubon  and  Bachman,  Op.  cit.,  vol.  3,  p.  250. 
^Brimley,  C.  S.,  Amer.  Nat,  vol.  31,  p.  448,  1897. 
18  Howell,  A.  H.,  Op.  cit.,  pp.  45-47. 


24  NORTH   AMERICAN    FAUNA.  [No.  45 

high  ceiling  of  a  large  chamber  about  300  yards  from  the  entrance, 
and  smaller  numbers  were  found  in  crevices  in  the  ceiling  close  to  the 
entrance.  In  the  cave  near  Fort  Deposit  the  bats  occupied  a  small, 
low  chamber,  reached  by  following  a  narrow,  winding  passage 
for  about  half  a  mile.  They  hung  from  the  ceiling  in  one  compact 
mass,  3  or  4  feet  square  and  several  bats  deep.  On  the  evening  of 
June  18  the  bats  were  observed  coming  out  of  this  cave  about  7 
o'clock;  they  swarmed  out  in  large  numbers,  feeding  in  the  mouth 
of  the  cave  and  among  the  trees  on  the  river  bank.  Of  the  18  speci- 
ments  collected  in  this  cave,  all  were  males;  while  in  Saltpetre  Cave, 
near  Rogersville,  of  62  specimens  collected  by  Holt,  46  were  females. 
Holt,  writing  of  his  visit  to  the  Rogersville  Cave,  says : 

The  bats  were  not  hanging  in  clusters,  but  thousands  of  them  lined  the  ceiling 
in  a  solid  sheet,  hanging  separately  head  downward.  A  couple  of  shots  were 
sufficient  to  cause  pandemonium,  and  immediately  I  found  myself  in  almost 
total  darkness  (my  assistant  having  retreated  around  a  corner  with  the  lan- 
tern), surrounded  by  a  swirling  mass  of  squeaking  bats.  They  were  every- 
where, the  flying  thousands  filling  the  air,  and  in  their  panic  rushing  against  me 
and  sticking  all  over  my  head  and  body ;  I  had  to  keep  kicking  to  prevent  them 
crawling  up  my  breeches  legs.  On  my  way  out  the  scattered  bats  seemed  to 
fill  the  whole  cave.  Their  droppings  covered  the  floor  in  places  to  the  depth  of 
several  feet. 

At  Limekiln,  on  the  Tennessee  River  north  of  Leighton,  we  ob- 
served a  number  of  bats,  believed  to  be  of  this  species,  coming  out  of 
a  small  cave  on  the  river  bluff,  from  which  issues  also  a  cold  spring. 

[Myotis  lucifugus  lucifugus  (LeConte). 
Little  Brown  Bat. 

V[espertilio]  lucifugus  LeConte,  McMurtrie's  Cuvier,  Animal  Kingdom,  vol.  1,  p. 
431,  1831. 
The  little  brown  bat  has  an  extensive  range  from  northern  Canada  south  to 
Florida,  but  apparently  is  absent  or  very  scarce  in  Alabama.  Miller  records 
a  skin  from  Greensboro  in  the  Merriam  collection,"  but  the  specimen  can  not 
be  located,  nor  is  there  any  record  of  it  in  Dr.  Merriam's  catalogue.  There  is 
considerable  doubt,  therefore,  as  to  the  occurrence  of  this  species  in  the  State.] 

Lasionycteris  noctivagans  (LeConte). 

Silver-haired  Bat. 

Y[est)ertilio']  noctinuians  LeConte,  McMurtrie's  Cuvier.  Animal  Kingdom,  vol.  1, 
p.  431,  1831. 

The  silver-haired  bat  is  a  northern-breeding  species,  and  occurs  in 
Alabama  only  as  a  migrant  or  possibly  a  winter  resident.  It  is  known 
at  present  only  from  five  specimens  taken  by  the  writer,  as  follows : 
Squaw  Shoals,  April  11,  1912  (2)  ;  Autaugaville,  April  17,  1912  (2)  ; 

17  Miller,  G.  S..  North  Amer.  Fauna  No.  13,  p.  62,  1897. 


1921.]  MAMMALS    OF    ALABAMA BATS.  25 

and  Sand  Mountain,  near  Carpenter,  October  29,  1916  (1).  At 
Squaw  Shoals  a  specimen  was  seen  which  had  been  taken  from  a  hol- 
low tree  by  J.  T.  Winchester,  in  the  fall  of  1911. 

These  are  the  most  southerly  records  for  the  species.  It  breeds  in 
the  northern  United  States  and  south  to  North  Carolina  and  possibly 
northern  Georgia 1S  and  migrates  in  autumn  to  the  Southern  States 
and  even  to  Bermuda.  The  silver-haired  bat  is  a  tree-dwelling 
species.  Vernon  Bailey  states  that  in  Minnesota  it  roosts  under  bark 
on  old  trees,  and  C.  S.  Brimley,  in  North  Carolina,  has  on  several 
occasions  taken  specimens  in  winter  in  hollow  trees.  Its  flight  is 
rather  slower  and  less  erratic  than  that  of  the  red  bat. 

Pipistrellus  subflavus  subflavus  (F.  Cuvier). 

Southeastern  Pipistreixe. 

Y[espertilio~\  subflavus  F.  Cuvier,  Nouv.  Ann.  Mus.  Hist.  Nat.,  Paris,  vol.  1,  p. 
17,  1832. 

The  southern  pipistrelle  is  the  smallest  and  one  of  the  commonest 
of  the  bats  in  the  South.  It  ranges  to  Florida  and  southern  Loui- 
siana, but  apparently  is  scarce  or  absent  in  southern  Alabama.  Speci- 
mens have  been  collected  at  Leighton,  Elkmont,  Stevenson,  Hunts- 
ville,  Fort  Payne,  Bucks  Pocket  (near  Grove  Oak),  and  Greensboro. 
A  number  of  small  bats  seen  at  Abbeville  were  thought  to  be  of  this 
species. 

This  little  bat  may  usually  be  distinguished  by  its  small  size  and  its 
erratic,  butterfly-like  flight.  It  frequents  caves  in  winter  and  to  a 
less  extent  in  summer.    Hahn  says  of  it,  as  observed  in  Indiana : 

Pipistrellus  subflavus  is  solitary  in  habit.  Occasionally  two  are  found  side  by 
side,  though  I  have  never  seen  them  clinging  to  each  other  except  in  mating. 
However  they  do  not  avoid  the  vicinity  of  others  of  their  own  kind  or  other 
species.  This  species  seems  to  prefer  the  side  walls  of  the  higher  passages.  I 
have  never  seen  it  suspended  from  the  roof  except  where  there  was  a  crevice  or 
prominent  ledge.10 

In  Bucks  Pocket,  on  Sand  Mountain,  a  semitorpid  individual  was 
found  November  5,  1916,  clinging  to  the  side  of  a  large  rock  in  a  cool 
ravine.  Hahn  mentions  seeing  one  under  a  ledge  of  rock  near 
Mitchell,  Indiana,  in  February.20 

Eptesicus  fuscus  fuscus  (Beauvois). 
Large  Brown  Bat. 
Vespertila  [sic]  fuscus  Beauvois,  Cat.  Peale's  Mus.,  Philadelphia,  p.  14,  1796. 

The  large  brown  bat  is  moderately  common  in  the  northern  part  of 
the  State,  but  has  thus  far  never  been  taken  in  the  southern  part.    It 

18  Specimens  are  known  from  Bertie  County,  North  Carolina,  July  1,  1891,  and  December 
26,  1S92  (Brimley,  Amer.  Nat.,  vol.  31,  p.  239,  1897)  ;  Highland,  North  Carolina,  April  18, 
18S6  (Merriam  collection)  ;  Toccoa,  Georgia,  May  26,  1916  (Biological  Survey  collection). 

19  Hahn,  W.  L.,  Biological  Bulletin   (Indiana  Univ.),  vol.  15,  No.  3,  p.  145,  1908. 

20  Op.  cit,  p.  144. 


26  NORTH   AMERICAN   FAUNA.  [No.  45 

has  been  taken  during  the  breeding  season  at  Leighton,  Sand  Moun- 
tain (near  Carpenter),  Erin,  and  Greensboro;  two  specimens  taken 
at  Autaugaville,  January  17  and  February  26,  1912,  may  have  been 
migrants. 

This  species  frequents  caves  to  some  extent  in  winter,  but  in  sum- 
mer is  most  often  found  concealed  about  buildings,  either  in  the 
attic,  in  some  dark  corner  under  the  eaves,  or  behind  a  shutter.  In  a 
schoolhouse  at  Leighton  on  July  3,  1913,  Holt  found  eight  individuals 
hanging  to  the  wall  of  the  open  vestibule,  just  beneath  the  ceiling. 
They  were  seen  there  about  9  p.  m.,  but  were  not  to  be  found  during 
the  day.  Dr.  Fisher  states  that  in  southern  New  York  this  species  is 
the  last  to  make  its  appearance  in  the  evening  and  that  its  favorite 
hunting  grounds  are  fields  well  surrounded  by  trees.21 

Nycteris  borealis  borealis  (Miiller). 
Red  Bat. 
Vespertilio  borealis  Miiller,  Natursyst,  Suppl.,  p.  21,  1776. 

The  red  bat  is  one  of  the  commonest  bats,  ranging  practically  all 
over  the  State,  except  in  the  coast  region,  where  its  place  is  largely 
taken  by  the  mahogany  bat  {Nycteris  seminola).  Specimens  have 
been  collected  at  Leighton,  Sand  Mountain  (near  Carpenter),  Gun- 
tersville,  Attalla,  Logan,  Mount  Weogufka,  Squaw  Shoals,  Greens- 
boro, Hayneville,  Autaugaville,  Barachias,  Point  Clear,  Ashford, 
Castleberry,  and  Abbeville.  In  the  coast  region  it  is  of  rare  occur- 
rence and  the  few  specimens  taken  there  (Point  Clear,  Apr.  21,  29, 
1892;  Ashford,  Nov.  27,  1916)  may  have  been  migrants. 

The  red  bat,  though  sometimes  found  in  caves  in  winter,  is  in  sum- 
mer a  dweller  in  trees  or  garrets.  On  one  occasion  at  Lake  Grove, 
New  York,  in  summer,  the  writer  came  upon  a  number  of  these  bats 
closely  bunched  together  hanging  to  a  low  branch  of  a  small  hickory 
tree.  This  is  one  of  the  first  bats  to  appear  in  the  evening,  being  seen 
usually  half  an  hour  or  more  before  dark  and  sometimes  also  during 
the  middle  of  the  day.  On  Sand  Mountain,  April  16,  1914,  one  was 
seen  about  noon  feeding  in  bright  sunlight ;  it  flew  leisurely  up  and 
down  over  a  creek  for  a  period  of  20  minutes  or  more.  Holt  reports 
seeing  one  drinking,  swallow-fashion,  over  the  Tennessee  River. 

This  bat  is  known  to  perform  extended  migrations  and  has  even 
been  observed  migrating  during  the  daytime.22 

Nycteris  seminola  (Rhoads). 

Mahogany  Bat. 

AtalapJia  borealis  seminola  Rhoads,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  p.  32, 
1895. 

The  mahogany  bat  is  a  dark-colored  species  occurring  commonly  in 
the  coast  region,  where  it  takes  the  place  of  the  red  bat  of  the  more 

"-1  Fisher,  A.  K.,  Forest  and  Stream,  vol.  16,  p.  490,  1881. 

-Miller,  G.  S.,  Science  (N.  S.),  vol.  5,  pp.  541-543,  April  2,  1897;  Mearns,  E.  A., 
Bull.  Amer.  Mus.  Nat.  Hist.,  vol.  10,  p.  345,  1898. 


1921.]  MAMMALS   OF   ALABAMA BATS.  27 

northern  counties.  Specimens  have  been  taken  at  Point  Clear,  April 
25,  1892 ;  Orange  Beach,  January  24,  28,  1912 ;  and  Mobile,  May  10, 
1911,  August  1,  1913,  and  February  27,  1916.  It  ranges  sporadically 
also  as  far  north  as  Autaugaville  and  even  Fort  Payne,  where  speci- 
mens were  taken  April  17,  1912,  and  June  30,  1911,  respectively. 
These  show  no  evidence  of  intergradation  with  borealis,  and  the  ma- 
terial at  present  available  indicates  that  this  form  is  a  distinct 
species.  Apparently  it  does  not  differ  materially  in  habits  from  its 
near  relative,  the  red  bat.  Like  the  latter  species,  it  is  an  early  flier, 
often  seen  abroad  shortly  after  sundown.  At  Orange  Beach,  the 
last  week  in  January,  one  was  seen  flying  a  few  minutes  after  5  p.  m. 
During  the  winter  season  it  apparently  feeds  only  on  mild  days.  In 
South  Carolina  the  writer  found  this  bat  resting  in  long  bunches  of 
Spanish  moss. 

Nycteris  cinerea   (Beauvois). 

Hoaky  Bat. 

Vespertilio  cinereus  Beauvois,  Cat.  Peale's  Mus.,  Philadelphia,  p.  15,  1796. 

The  large  and  handsome  hoary  bat  occurs  in  the  State  only  as  a 
rare  migrant ;  only  two  specimens  have  thus  far  been  taken — one  by 
Russell  J.  Thompson,  at  Point  Clear,  Mobile  Bay,  April  19,  1892, 
the  other  by  Lewis  S.  Golsan,  at  Autaugaville,  January  15,  1912.  Its 
summer  home  is  chiefly  within  the  Canadian  Zone,  from  northern 
New  York  and  Wisconsin  northward,  but  during  the  winter  season 
it  migrates  southward  for  long  distances,  having  been  taken  in 
central  Mexico,  southern  Lower  California,  and  the  Bermuda 
Islands.  Little  is  known  of  its  habits,  but,  like  its  smaller  relative, 
the  red  bat,  it  is  undoubtedly  a  tree-dweller.  Merriam  writes  of  its 
habits  as  observed  in  the  Adirondack  region  of  New  York : 

The  Hoary  Bat  can  be  recognized,  even  in  the  dusk  of  evening,  by  its  great 
size,  its  long  and  pointed  wings,  and  the  swiftness  and  irregularity  of  its 
flight.  It  does  not  start,  out  so  early  as  our  other  bats,  and  is  consequently 
much  more  difficult  to  shoot.  The  borders  of  woods,  water  courses,  and  road- 
ways through  the  forest  are  among  its  favorite  resorts,  and  its  nightly  range  is 
vastly  greater  than  that  of  any  of  its  associates.  While  the  other  species 
are  extremely  local,  moving  to  and  fro  over  a  very  restricted  area,  this  traverses 
a  comparatively  large  extent  of  territory  in  its  evening  excursions,  which  fact 
is  probably  attributable  to  its  superior  power  of  flight.22 

Nycticeius  hunieralis  (Rafinesque). 

Evening  Bat. 

Vespertilio  humeralis  Rafinesque,  Anier.  Monthly  Mag.,  vol.  3,  p.  445,  1818. 

The  evening  bat  is  abundant  and  generally  distributed  throughout 
the  State.     Specimens  have  been  taken  at  Leighton,  Guntersville, 

22  Merriam,  C.  nart,  Mamm,  Adirondack  region  :  Trans.  Linnaean  Soc.  New  York,  vol.  2, 
p.  78,  1884. 


28  NORTH   AMERICAN   FAUNA.  [No.  45 

Ardell,  Squaw  Shoals,  Erin,  Mount  Weogufka,  Auburn,  Greensboro, 
Hayneville,  Seale,  Abbeville,  Bon  Secour,  Point  Clear,  and  Castle- 
berry.  In  flight  it  appears  of  small  or  medium  size  and  of  a  very 
dark  color.  Like  the  red  bat,  it  begins  to  hunt  considerably  before 
dark ;  at  Guntersviile,  in  the  middle  of  June,  the  first  ones  were  seen 
flying  about  7  o'clock  in  the  evening.  Compared  with  that  of  other 
species,  its  flight  is  rather  slow  and  steady,  often  close  to  the  earth, 
though  sometimes  at  a  moderate  height.  In  winter  this  bat  probably 
hibernates  in  the  same  manner  as  other  species.  A  specimen  was 
taken  at  Bon  Secour  on  October  19,  1908;  and  at  Autaugaville  the 
writer  saw  several  flying  on  November  20  and  21,  1916. 

Corynorhinus  macrotis    (LeConte). 

LeConte  Big-eared  Bat. 

Plec[otus]   macrotis  LeConte,  McMurtrie's  Cuvier,  Animal  Kingdom,  vol.  1,  p. 
431,  1831. 

The  LeConte  big-eared  bat  is  apparently  of  rather  local  distribu- 
tion and  nowhere  very  common  in  the  State.  Only  a  few  specimens 
have  thus  far  been  captured — one  each  from  Huntsville,  Town  Creek, 
and  Greensboro,  and  two  from  Autaugaville.  The  Huntsville  speci- 
men was  obtained  in  a  small  outhouse  on  Monte  Sano  (altitude  1,600 
feet),  having  been  stupefied  by  the  smoke  of  a  fire  built  in  the  house, 
so  that  it  dropped  from  the  ceiling,  to  which  it  was  clinging.  F.  W. 
McCormack  described  some  very  long-eared  bats  which  he  caught 
some  years  ago  in  the  garret  of  an  old  mill  on  Town  Creek;  later 
a  specimen  was  obtained  (July  4,  1913)  from  this  mill  by  E.  G.  Holt. 
Miller  lists  a  skin  in  the  Merriam  collection  from  Greensboro,  but 
the  specimen  can  not  now  be  located,  nor  is  there  any  other  record 
of  it.23  L.  S.  Golsan  obtained  two  specimens  at  Autaugaville,  Au- 
gust 22  and  September  3,  1912.  Harm  states  that  in  Indiana  this 
species  has  been  found  only  in  dimly  lighted  parts  of  caves  near  the 
entrances,  where  it  was  clinging  to  the  side  walls,  its  long  ears 
folded  down  along  the  neck.24 

Family  MOLOSSIDAE  :  Free-tailed  Bats. 

Nyctinomus  cynocephalus   (LeConte). 

LeConte  Fkee-tailed  Bat. 
Nyct[icea]  cynocephala  LeConte,  McMurtrie's  Cuvier,  Animal  Kingdom,  vol.  1, 
p.  432,  1831. 

The  LeConte  free-tailed  bat  probably  occurs  more  or  less  com- 
monly over  at  least  the  southern  part  of  Alabama,  but  thus  far  it  has 
been  taken  only  at  Greensboro  and  Orange  Beach.     It  is  primarily 

23  Miller,  G.  S.,  North  Amer.  Fauna  No.  13,  p.  52,  1897. 

M  Hahn,  W.  L.,  Biological  Bull.   (Indiana  Univ.),  vol.  15,  no.  3,  p.  145,  1908. 


1921.]  MAMMALS    OF  ALABAMA BEARS.  29 

a  house  bat,  often  infesting  the  attics  of  dwellings  in  such  numbers 
as  to  become  a  decided  nuisance.  It  hides  also  in  the  crevices  of 
wooden  bridges.  Vernon  Bailey  states  that  the  species  possesses  a 
rank,  musky  odor,  so  strong  as  to  be  noticeable  from  the  outside  of 
an  infested  house.  The  late  Dr.  W.  C.  Avery  caught  about  400 
specimens  by  setting  nets  on  the  outside  of  an  old  building  at  Greens- 
boro; 30  specimens,  secured  July  31  and  August  1, 1890,  are  preserved 
in  the  Biological  Survey  collection.  At  Orange  Beach,  January  24 
and  28,  1912,  two  specimens  shot  by  local  hunters  were  secured.  The 
writer  has  shot  many  bats  in  various  parts  of  the  South,  but  as  yet 
has  never  taken  this  species. 

Order  CARNIVORA. 

Family  URSIDAE :  Bears. 

TJrsus  fioridanus  Merriam. 

Florida  Black  Bear. 

Vrsus  fioridanus  Merriam,  Proc.  Biol.  Soc.  Washington,  vol.  10,  p.  81,  1896. 

Bears  doubtless  ranged  over  all  of  Alabama  in  early  times,  but 
at  present  are  exterminated  everywhere  except  in  the  swamps  of 
the  southern  counties.  In  the  big  swamps  bordering  the  Tensaw  and 
Mobile  Rivers  they  are  still  common  and  a  number  are  killed  there 
ever}-  fall.  A.  J.  Mclntyre,  of  Carlton,  is  reported  to  have  killed 
in  recent  years  over  100  bears  and  to  have  caught  10  cubs.  In  April, 
1911,  he  killed  an  old  male,  estimated  to  weigh  500  pounds,  which 
for  25  years  had  been  known  to  settlers  in  that  region  as  "  Old  Nub- 
foot,"  being  recognized  by  his  peculiar  track.  This  bear  had  be- 
come very  bold  in  raiding  hog  pens  and  was  charged  with  the  de- 
struction of  a  considerable  number  of  calves  and  hogs.  Bears  are 
occasionally  found  in  the  swamps  of  southern  Mobile  County,  and 
are  reported  fairly  numerous  near  Bayou  Labatre,  where  they  do 
considerable  damage  to  stock.  In  November,  1915,  T.  J.  Pendarvis 
killed  a  large  male,  near  Irvington,  estimated  to  weigh  400  pounds. 
The  animal  was  baited  in  the  swamp  through  which  he  had  been 
passing  and  was  shot  by  moonlight  from  a  place  of  concealment  near 
the  bait.  This  bear  was  said  to  have  a  regular  trail  through  the 
swamps  and  to  cross  a  public  road  not  over  3  miles  from  Bayou 
Labatre  by  climbing  a  wire  fence. 

At  Bon  Secour  a  single  bear  was  reported  about  1905 — the  only 
one  for  many  years.  At  Ashford,  Houston  CoumVy,  bears  are  rarely 
found,  but  during  a  big  freshet  in  July,  1916,  the  tracks  of  one 
were  seen  in  a  field  near  a  swamp. 

Four  skulls  of  the  southern  Alabama  bear  have  been  examined 
and  they  indicate  that  the  form  found  there  is  intermediate  between 


30  NORTH   AMERICAN   FAUNA.  [No.  45 

floridanus  and  luteolus.  The  skull  of  an  old  adult  male  from  Carl- 
ton agrees  in  general  with  luteolus  of  comparable  age,  from  Louisi- 
ana, but  differs  in  having  smaller  teeth,  a  somewhat  shorter  muzzle, 
and  a  shorter  palate.  Compared  with  the  type  of  floHdanus  (from 
Dade  County,  Florida)  and  with  an  adult  from  New  Smyrna, 
Florida,  this  skull  differs  in  having  the  palatal  floor  more  level,  the 
molars  somewhat  larger,  the  muzzle  shorter  and  relatively  broader, 
and  the  frontal  region  slightly  less  elevated.  An  old  adult  (prob- 
ably female)  from  the  same  locality  and  a  young  adult  male  from 
Bayou  Labatre  agree  with  floridanus  in  size  of  teeth,  depression  of 
palatal  floor,  shape  of  muzzle,  and  elevation  of  frontal  region.  The 
single  skin  from  Alabama  (BayW  Labatre)  is  glossy  black  all  over, 
except  the  muzzle,  which  is  brownish  black  above,  the  sides  of  the 
face  ochraceous-tawny. 

The  extermination  of  the  bears  of  northern  Alabama  makes  it 
impossible  to  say  what  race  was  found  there;  quite  likely  typical 
amerlcanus  ranged  at  least  in  the  mountainous  parts. 

Family  CAXIDAE :  Wolves  and  Foxes. 
Canis  floridanus  Miller. 

SOUTHEASTKKX    WOLF. 

Canis  floridanus  Miller,  Proc.  Biol.  Soc.  Washington,  vol.  25,  p.  95,  1912. 

Wolves  in  former  times  doubtless  ranged  over  the  greater  part 
of  Alabama,  but  are  now  on  the  verge  of  extinction.  Their  last 
stronghold  appears  to  be  the  rough,  hilly  country  stretching  from 
Walker  County  northwestward  to  Colbert  County. 

It  was  reported  that  three  or  four  wolves  were  killed  in  September, 
1912,  on  the  Blackwater  near  South  Lowell  by  the  O'Rear  brothers, 
and  during  the  spring  of  1914  the  animals  were  heard  howling 
(according  to  reports)  at  points  in  Walker  County  within  12  miles 
of  Jasper.  C.  H.  Harbison,  living  on  Ryan  Creek,  in  western  Cull- 
man County,  states  that  wolves  were  destructive  to  stock  in  that 
region  in  1915,  and  that,  after  eating  up  most  of  the  sheep  and  lambs, 
they  left  the  country  and  moved  westward  into  Winston  and  Marion 
Counties.  This  migration  is  apparently  confirmed  by  the  appearance 
of  the  animals  about  that  time  in  Franklin  and  Colbert  Counties. 
D.  L.  Paden,  whose  home  is  in  the  hill  country,  12  miles  south  of 
Cherokee,  writes  (in  1917)  that  the  wolves  came  into  that  country 
"  several  years  ago " ;  that  they  killed  thousands  of  dollars'  worth 
of  sheep  and  goats,  and  practically  stopped  the  raising  of  these 
animals;  and  that  some  calves  also  were  killed  by  them.  In  April, 
1917,  in  the  same  locality,  G.  G.  Paden  killed  an  adult  male  wolf, 
the  skin  and  skull  of  which  were  secured  by  the  Biological  Survey. 


1921.]  MAMMALS   OF   ALABAMA WOLVES.  31 

This  is  the  only  specimen  from  the  Southeastern  States,  excepting  a 
single  one  from  Florida,  that  has  come  into  the  possession  of  any 
museum,  and  is  therefore  of  extreme  interest.  It  may  be  described 
as  follows* 

Upperparts  mixed  blackish  and  cinnamon-buff;  shoulders,  sides  of  neck 
and  body,  and  fore  back  washed  with  light  buff ;  top  and  sides  of  head  grizzled 
iron  gray  (produced  by  a  mixture  of  black  and  light  buff),  tinged  medially 
with  cinnamon-buff;  muzzle  light  pinkish  cinnamon,  shading  to  light  buff 
around  edges  of  lips  and  on  sides  of  face  beneath  the  eyes ;  ears  cinnamon-buff, 
shaded  with  blackish ;  front  legs  and  feet  light  buff,  washed  with  cinnamon-baff 
and  with  a  faint  median  stripe  of  blackish;  hind  legs  and  feet  light  buff, 
strongly  washed  with  cinnamon ;  tail  mixed  cinnamon  and  light  buff,  shaded 
above  and  on  tip  with  blackish ;  underparts  mixed  light  buff  and  cinnamon- 
buff,  washed  with  whitish.  Compared  with  the  type  of  Cants  floridanus  (from 
Putnam  County,  Florida)  the  Alabama  specimen  differs  in  the  following  partic- 
ulars ;  Top  and  sides  of  head  more  grayish ;  muzzle  slightly  paler ;  ears  less 
tawny ;  fore  legs  paler  and  with  less  extensive  median  black  line ;  hind  legs 
paler  (less  strongly  ochraceous).  It  differs  from  Michigan  specimens  (lycaon?) 
in  being  slightly  paler,  especially  on  the  fore  legs  and  muzzle,  with  the  ears  less 
intensely  tawny. 

The  skull  (adult  $  )  agrees  essentially  with  that  of  floridanus  (type,  adult  5  ), 
but  is  slightly  larger  and  relatively  narrower  across  postorbital  processes,  with 
a  slightly  longer  rostrum.  It  agrees  in  dental  characters  with  the  Louisiana 
wolf  (specimens  from  Tallulah,  Louisiana),  but  differs  in  being  relatively 
broader  with  a  shorter,  broader  rostrum.  Compared  with  lycaon  of  eastern 
Canada  (specimens  from  40  miles  northeast  of  Mattawa,  Quebec)  it  differs  as 
follows :  Skull  of  nearly  the  same  size,  but  with  braincase  and  rostrum  rela- 
tively narrower;  upper  premolars  and  carnassial  longer  and  relatively  narrower, 
the  latter  with  antero-internal  heel  more  strongly  developed ;  first  upper  molar 
broader,  with  posterior  middle  cusp  more  strongly  developed. 

Wolves  ranged  in  some  numbers  in  the  Talladega  and  Choccolocco 
Mountains  until  about  1896,  in  which  year,  in  northern  Clay  County, 
17  were  said  to  have  been  killed  in  two  days.  The  last  one  killed  in 
that  region  was  about  1905  and  it  was  mounted  and  exhibited  in 
Talladega  for  a  number  of  years.  Wolves  were  heard  howling  on 
Duggar  Mountain  near  Piedmont  as  late  as  1914,  but  at  the  present 
time  they  are  almost,  if  not  quite,  exterminated  in  that  region.  A 
black  wolf  Was  reported  to  have  been  killed  on  La  Grange  Mountain, 
near  Leighton,  about  1906  and  more  recently  another  was  seen  there. 

Strangely  enough,  in  the  big  swamp  country  in  Baldwin  and  Mobile 
Counties,  where  deer  are  still  numerous,  wolves  apparently  were  ex- 
terminated many  years  ago ;  the  last  one  of  which  there  is  record  was 
reported  killed  near  Carlton  about  1894. 

Little  is  known  of  the  habits  of  the  Alabama  wolves  except  that 
they  roamed  the  mountains  in  small  droves  and  fed  considerably  on 
the  smaller  domestic  animals — sheep,  goats,  pigs,  and  sometimes 
calves.  Bartram,  writing  in  1791,  mentions  the  occurrence  of  wolves 
on  the  lower  Tombigbee  Kiver,  and  describes  pups  of  a  litter  found 


32  NORTH   AMERICAN   FAUNA.  [No.  45 

in  July,  1776,  on  the  Tallapoosa  River,  near  Coolome,  as  "  half  the 
size  of  a  small  cur  dog  and  quite  black." 25 

Vulpes  fulva   (Desmarest). 
Red  Fox. 

Canis  fuhnis  Desmarest.  Mammalogie,  vol.  1,  p.  203,  1820. 

The  red  fox  apparently  is  not  native  in  the  Southern  States,  but 
has  been  introduced  or  has  extended  its  range  into  that  region  within 
historic  times.  Eegarding  the  early  history  of  the  animal  in  America, 
Audubon  and  Bachman  make  the  following  statements : 

Red  Poxes  have  gradually  migrated  from  the  Northern  to  the  Southern  States. 
This  change  of  habitation  may  possibly  be  owing  to  the  more  extensive  cultiva- 
tion to  which  we  have  alluded  *  *  *  as  a  reason  for  this  species  having 
become  more  numerous  than  it  was  before  the  Revolution.  This  idea,  however, 
would  seem  to  be  overthrown  by  the  continued  abundance  of  Gray  Foxes  in  the 
Eastern  States.  In  the  early  history  of  our  country  the  Red  Fox  was  unknown 
south  of  Pennsylvania,  that  State  being  its  Southern  limit.  In  process  of  time 
it  was  found  in  the  mountains  of  Virginia,  where  it  has  now  become  more 
abundant  than  the  Gray  Fox.  A  few  years  afterwards  it  appeared  in  the  more 
elevated  portions  of  North  Carolina,  then  in  the  mountains  of  South  Carolina, 
and  finally  in  Georgia,  where  we  have  recently  observed  it. 

This  species  was  first  seen  in  Lincoln  County,  Georgia,  in  the  year  1840, 
since  then  it  has  spread  over  the  less  elevated  parts  of  the  country,  and  is  not 
rare  in  the  neighborhood  of  Augusta.  We  are  informed  by  Mr.  Beile,  an  intelli- 
gent observer  of  the  habits  of  animals,  that  on  one  occasion  near  Augusta,  as 
he  was  using  a  call  for  wild  turkeys,  a  little  before  sunrise,  in  the  vicinity  of 
Augusta,  two  Red  Foxes  came  to  the  call,  supposing  it  to  be  that  of  a  wild 
turkey,  and  were  both  killed  by  one  discharge  of  his  gun.26 

Whether  the  species  extended  its  range  naturally  into  Alabama  is 
not  known.  Numerous  reports  from  residents,  however,  indicate 
that  foxes  from  other  States  have  been  imported  and  liberated  in 
many  widely  separated  localities.  Statements  to  this  effect  have 
been  made  by  people  at  Dean,  Wilsonville,  Castleberry,  Abbeville, 
and  Orange  Beach.  Red  foxes  are  reported  to  occur  in  small  num- 
bers also  at  Elkmont,  Leighton,  Sand  Mountain,  Woodville,  Hunts- 
ville,  Fort  Payne,  Mount  Weogufka,  Jackson,  Dothan,  and  Bon 
Secour,  and  in  Elmore,  Wilcox,  and  Montgomery  Counties.  Two 
skulls  from  Alabama  (Tennessee  River,  north  of  Leighton;  Catoma 
Creek,  Montgomery  County)  agree  essentially  with  a  specimen  from 
Virginia  assumed  to  be  typical  fulva. 

The  den  of  the  red  fox  is  said  to  be  in  caves  or  clefts  of  rocky 
bluffs,  sometimes  in  old  stumps  or  hollow  logs,  or  in  burrows  in  the 
ground.     Merriam  describes  the  animal's  feeding  habits  as  follows : 

25  Bartram.  William,  Travels,  pp.  396;  410,  1702. 

26  Audubon  and  Bachman,  Op.  cit.,  vol.  2,  pp.  269-270. 


1921.]  MAMMALS   OF   ALABAMA FOXES.  33 

He  is  both  nocturnal  and  diurnal  in  habits,  and  preys  upon  skunks,  wood- 
chucks,  muskrats,  hares,  rabbits,  squirrels,  mice,  and  small  birds  and  eggs. 
He  is  a  well-known  and  much-dreaded  depredator  of  the  poultry  yard,  destroy- 
ing, with  equal  alacrity,  turkeys,  ducks,  geese,  hens,  chickens,  and  doves;  and 
has  been  known  to  make  off  with  young  lambs.  He  will  also  eat  carrion,  and 
even  fish,  and  is  said  to  be  fond  of  ripe  grapes  and  strawberries.87 

TTrocyon  cinereoargenteus  cinereoargenteus  (Schreber). 
Gray  Fox. 
Cams  cinereo-argenteus  Schreber,  Saugthiere,  vol.  3,  pi.  92,  1775. 

The  gray  fox  is  reported  from  all  sections  of  the  State  and  is  still 
common  in  many  localities,  though  much  reduced  in  numbers  in  the 
more  thickly  settled  regions.  Fox  hunting  with  hounds  is  a  favorite 
sport  and  has  resulted  in  extermination  of  the  species  in  some  locali- 
ties. The  gray  fox  is  a  sly  and  cunning  creature,  but  he  lacks  the 
wonderful  shrewdness  and  sagacity  of  the  red  fox,  which  has  made 
the  latter  so  popular  as  an  object  of  the  chase.  Audubon  and  Bach- 
man  give  a  good  account  of  the  animal's  habits,  from  which  the  fol- 
lowing is  quoted : 

The  Gray  Fox  is  shy  and  cowardly,  and  the  snap  of  a  stick  or  the  barking  of  a 
dog  will  set  him  off  on  a  full  run.  Although  timid  and  suspicious  to  this  degree, 
his  cunning  and  voracity  place  him  in  a  conspicuous  rank  among  the  animals 
that  prey  upon  other  species  weaker  than  themselves.     *     *     * 

In  the  Southern  States  this  species  is  able  to  supply  itself  with  a  great  variety 
and  abundance  of  food,  and  is  consequently  generally  in  good  condition  and 
often  quite  fat.  We  have  followed  the  track  of  the  Gray  Fox  in  moist  ground 
until  it  led  us  to  the  scattered  remains  of  a  marsh  hare,  which  no  doubt  the  fox 
had  killed.  Many  nests  of  the  fresh  water  marsh  hen  (Rallus  elegans)  are 
torn  to  pieces  and  the  eggs  devoured  by  this  prowler.  In  Pennsylvania  and 
New  Jersey,  the  meadow  mouse  (Arvicola  Pennsylvanica)  is  often  eaten  by  this 
species ;  and  in  the  Southern  States,  the  cotton-rat  and  Florida  rat,  constitute 
no  inconsiderable  portion  of  its  food.  We  have  seen  places  where  the  Gray  Fox 
had  been  scratching  the  decayed  logs  and  the  bark  of  trees  in  order  to  obtain 
insects. 

Although  this  Fox  is  nocturnal  in  his  habits  we  have  frequently  observed  him 
in  search  of  food  at  all  hours  of  the  day ;  in  general,  however,  he  lies  concealed 
in  some  thicket,  or  in  a  large  tuft  of  tall  broom-grass,  till  twilight  invites  him 
to  renew  his  travels  and  adventures. 

On  a  cold  starlight  night  in  winter,  we  have  frequently  heard  the  hoarse 
querulous  bark  of  this  species ;  sometimes  two  of  them,  some  distance  apart, 
were  answering  each  other  in  the  manner  of  the  dog. 

Although  we  have  often  seen  this  Fox  fairly  run  down  and  killed  by  hounds, 
without  his  having  attempted  to  climb  a  tree,  yet  it  not  infrequently  occurs 
that  when  his  strength  begins  to  fail  he  ascends  one  that  is  small  or  sloping, 
and  standing  on  some  horizontal  branch  20  or  30  feet  from  the  ground,  looks 
down  on  the  fierce  and  clamorous  pack  which  soon  comes  up  and  surrounds 
the  foot  of  the  tree.  *  *  *  We  were  unable  to  obtain  any  information  in 
regard  to  the  manner  in  which  the  Fox  climbs  trees,  as  he  does  not  possess  the 


27  Merriam,  C.  Hart,  Trans.  Linnaean  Soc.  New  York,  vol.  1,  p.  45,  1882, 
49663°— 21 3 


34  NORTH   AMERICAN    FAUNA.  [No.  45 

retractile  nails  of  the  cat  or  the  sharp  claws  of  the  squirrel,  until  we  saw  the 
animal  in  the  act.  At  one  time  when  we  thus  observed  the  Fox,  he  first  leaped 
onto  a  low  branch  4  or  5  feet  from  the  ground,  from  whence  he  made  his 
way  upward  by  leaping  cautiously  and  rather  awkwardly  from  branch  to 
branch,  till  he  attained  a  secure  position  in  the  largest  fork  of  the  tree,  where 
he  stopped.  On  another  occasion,  he  ascended  in  the  manner  of  a  bear,  but  with 
far  greater  celerity,  by  clasping  the  stem  of  a  small  pine.  We  have  since  been 
informed  that  the  Fox  also  climbs  trees  occasionally  by  the  aid  of  his  claws,  in 
the  manner  of  a  raccoon  or  a  cat.28 

This  genus  has  not  been  revised,  and  material  is  lacking  from  east- 
ern United  States  to  determine  the  ranges  of  the  various  forms.  Two 
skulls  (adult  $  )  from  Cullman  County  agree  essentially  with  skulls 
from  Maryland,  assumed  to  be  typical  elnereoargenteus ;  an  adult  (un- 
sexed)  from  Orange  Beach  has  somewhat  larger  molar  teeth,  in  this 
respect  apparently  approaching  subspecies  foridaaius.  The  skin  of 
the  latter  does  not  differ  appreciably  from  those  of  typical  individ- 
uals. Immature  specimens  have  been  examined  from  Autaugaville, 
and  the  animal  is  reported  by  residents  to  occur  at  Leighton,  Wood- 
ville,  Huntsville,  Elkmont,  Sand  Mountain,  Fort  Payne,  Piedmont, 
Talladega  Mountains,  Natural  Bridge,  Ardell,  Wilsonville,  York, 
Reform,  Hayneville,  Jackson,  Seale,  Dothan,  Castleberry,  Carlton, 
Mobile,  and  Bayou  Labatre.  At  Abbeville  the  native  fox  is  said  to 
have  about  disappeared,  and  fox  hunters  have  imported  and  turned 
loose  a  number  from  Kentucky  and  Tennessee. 

Family  PROCYONIDAE:  Raccoons. 

Procyon  lotor  lotor   (Linnaeus). 

Raccoon. 

[Ursus]  lotor  Linnaeus,  Syst.  Nat.,  ed.  10,  vol.  1,  p.  48,  1758. 

The  "  coon  "  is  one  of  the  commonest  and  best  known  of  the  south- 
ern mammals.  'Coon  hunting  is  one  of  the  most  popular  sports  and 
many  animals  are  captured  in  nocturnal  hunts;  thousands  are  trapped 
also  every  season  by  boys  and  by  professional  trappers,  so  that  in 
many  localities  the  animal  is  now  becoming  scarce. 

Raccoons  dwell  chiefly  in  timbered  swamps  and  river  bottoms, 
where  they  forage  extensively  along  the  banks  of  the  streams. 
Although  good  swimmers  they  do  not  take  to  the  water  from  pref- 
erence, but  usually  cross  a  stream  on  a  fallen  tree.  On  the  coast  the 
salt  marshes  prove  attractive,  for  in  them  'coons  find  an  abundance  of 
shellfish  and  crustaceans,  which  are  favorite  articles  of  diet.  In 
January,  1912,  while  hunting  with  dogs  on  a  marsh  island  near  the 
mouth  of  Perdido  Bay,  the  dogs  ran  down  and  captured  a  'coon  that 
was  hiding  in  the  marsh  grass.     'Coon  tracks  are  numerous  through- 

*  Audubon  and  Bachman,  Op.  cit.,  vol.  1,  pp.  164,  186-167, 


1921.]  MAMMALS   OF   ALABAMA WEASELS.  35 

out  these  wet  marshes  and  along  the  shores  of  the  bays.  In  the  big 
swamps  of  the  Tensaw  and  Mobile  Rivers,  'coons  are,  or  have  been, 
very  numerous.  In  March,  1912,  going  up  the  Tensaw  in  a  launch,  the 
writer  shot  two  out  of  the  tops  of  tall  trees  in  the  swamp,  where  they 
were  curled  up  asleep.  The  raccoon  is  by  no  means  confined  to  the 
swamps,  however,  but  makes  frequent  visits  to  the  adjacent  corn- 
fields and  feasts  upon  the  ripening  ears  in  the  milk  stage,  often  doing 
considerable  damage. 

The  diet  of  the  raccoon  includes  a  great  variety  of  food ;  wild  fruits 
and  berries  are  eagerly  sought  when  available ;  shellfish,  crustaceans, 
fish,  small  mammals,  reptiles,  and  birds  are  secured  as  opportunity 
offers;  and  occasional  raids  are  made  on  the  farmers'  hen  roosts. 
Audubon  and  Bachman  speak  of  the  animal's  fondness  for  birds'  eggs 
and  for  those  of  the  soft-shelled  turtle,  and  of  its  habit  of  feeding  on 
fresh-water  mussels  and  the  small  oysters  growing  in  shallow  waters 
at  the  mouths  of  the  rivers,  known  locally  as  "  raccoon  oysters."  Stone 
and  Cram  speak  of  its  feeding  habits  as  follows: 

From  the  accounts  of  numerous  eye-witnesses  it  would  appear  to  be  a  pretty 
regular  practice  with  them  to  lie  in  wait  at  the  edge  of  the  water  and  hook  out 
any  fish  that  comes  within  reach  by  a  smart  stroke  of  the  fore  paw  with  claws 
extended. 

Being  night  wanderers,  they  undoubtedly  often  manage  to  surprise  sleeping 
birds,  both  on  the  ground  and  among  the  branches,  as  it  is  a  common  custom 
with  them  in  thick  woods  to  travel  for  long  distances  among  the  tree-tops  with- 
out once  descending  to  earth,  robbing  the  nests  of  birds  and  squirrels  on  the 
way.30 

Specimens  examined  from  Huntsville,  Sylacauga,  Barachias, 
Castleberry,  Hurricane,  and  Orange  Beach  are  all  referable  to 
t37pical  lotor;  the  skulls  show  no  approach  to  the  characters  of  the 
Florida  form  (elucus),  although  the  skins  are  somewhat  more 
ochraceous  above  than  Maryland  and  Virginia  specimens. 

Family  MUSTELIDAE:  Weasels,  Minks,  Skunks,  Otters,  etc. 

Mustela  noveboracensis   (Emmons). 

New  York  Weasel. 

Putorius  noveboracensis  Emmons,  Kept.  Quadr.  Massachusetts,  p.  45,  1840. 

The  New  York  weasel  occupies  the  northeastern  United  States, 
extending  south  into  Alabama  in  the  mountains  of  the  northern 
counties.  It  is  definitely  known  only  from  a  single  specimen 
trapped  by  the  writer  at  the  base  of  a  cliff  on  Lookout  Mountain, 
near  Fort  Payne,  at  an  altitude  of  1,500  feet.  Records  of  weasels 
from  Sand  Mountain,  Gunters  Mountain,  Talladega  Mountains,  Elk- 
mont,  and  Woodville  are  provisionally  referred  to  this  species.     A 

M  Stone,  Witmer,  and  William  Everitt  Cram,  American  animals,  p.  250,  1902. 


36  NOETH   AMERICAN   FAUNA.  [No.  45 

resident  of  Carpenter  (Jackson  County)  related  that  he  had  seen  a 
den  of  weasels  in  a  rock  pile  at  that  place ;  two  weasels  were  reported 
to  have  been  killed  in  April,  1910,  in  a  sawmill  slab  pile  at  Dean. 
Stone  and  Cram  describe  the  feeding  habits  of  this  weasel  as  follows : 

In  winter  the  larger  weasels  kill  large  numbers  of  gray  rabbits.  *  *  *  In 
summer  they  catch  grasshoppers,  crickets,  and  beetles  of  various  sorts,  and  rob 
every  bird's  nest  they  find.  Ground-feeding  birds  are  especially  liable  to  be 
caught  by  them,  and  they  have  even  been  seen  to  spring  into  the  air  and  catch 
birds  on  the  wing.31 

Mustela  peninsulae  olivacea  Howell. 

Alabama  Weasel. 

Mustela  peninsulae  olivacea  Howell,   Proc.  Biol.   Soe.   Washington,  vol.  26,  p. 
139, 1913. 

Weasels  are  apparently  scarce  everywhere  in  the  Southern  States 
and  specimens  are  difficult  to  obtain.  The  Alabama  weasel  is  re- 
lated to  the  Florida  weasel  and  not  to  the  Alleghenian  species.  It 
inhabits  practically  the  whole  of  the  State  except  the  mountainous 
regions  of  the  northeastern  part,  but  the  limits  of  its  range  are  at 
present  unknown.  Specimens  have  been  examined  from  Autauga- 
ville,  Ardell,  Leighton,  and  Muscle  Shoals ;  the  animals  are  reported 
from  Greensboro,  Myrtlewood,  Oakchia,  Reform,  Teaslej^  Mill  (Mont- 
gomery County),  Seale,  Jackson,  Castleberry,  Newville,  Dothan, 
Abbeville,  and  Point  Clear. 

This  is  a  rather  large  weasel.  The  upperparts  in  winter  are  buffy 
brown,  with  a  tinge  of  olivaceous;  in  summer  mummy  brown,  de- 
cidedly darker  than  the  winter  pelage;  the  underparts  are  washed 
with  straw  color  or  buff. 

Little  is  known  of  the  habits  of  this  species.  It  lives  in  the  drier 
parts  of  the  timbered  swamps,  making  its  den  usually  under  the 
roots  of  a  tree  or  in  a  hollow  stump,  and  ranges  also  into  the  rocky 
hill  country,  where  it  inhabits  crevices  in  cliffs  or  rock  piles.  J.  S. 
Tharp,  of  Ashford,  states  that  he  once  saw  two  of  these  weasels 
run  under  the  roots  of  a  large  tree  in  a  swamp  near  Dothan.  The 
late  Dr.  W.  C.  Avery,  of  Greensboro,  writes  that  the  weasel, 
although  not  common,  at  times  proved  very  destructive  to  chickens 
and  young  pigeons,  cutting  the  throats  and  sucking  the  blood  of  a 
dozen  or  more  fowls,  the  bodies  of  which  it  never  touched  except  to 
drink  the  blood.  L.  S.  Golsan,  who  secured  the  type  specimen  of  this 
race  near  Autaugaville,  states  that  these  weasels  have  been  found  in 
nests  of  the  swamp  wood  rat  (Neotoma  foridana).  He  says  they 
take  to  trees  almost  as  readily  as  squirrels. 

31  Stone  and  Cram,  Op.  cit.,  pp.  236,  237, 


1921.]  MAMMALS   OF  ALABAMA — SKUNKS.  37 

Mustela  vison  mink  Peale  and  Beauvois.82 

Mink. 

Mustela  mink  Peale  and  Beauvois,  Catalog  Peale's  Mus.,  p.  39,  1796. 

Minks  are  generally  distributed  over  most  of  Alabama,  except  the 
most  southern  counties,  where  they  are  very  scarce.  In  many  locali- 
ties they  have  been  almost  exterminated,  and  the  recent  high  prices 
of  fur  have  led  to  a  great  decrease  in  their  numbers  everywhere. 

A  series  of  12  skulls  from  Barachias  and  Autaugaville  and  4  from 
Winston  and  Cullman  Counties  are  practically  typical  of  this  sub- 
species. A  series  of  5  skulls  from  the  Mobile  River  swamps  is 
also  referable  here,  though  showing  approach  to  subspecies  vulgivaga, 
of  Louisiana,  in  the  small  size  of  the  females;  the  single  male  skull 
is  typical  mink. 

The  species  is  reported  also  from  Leighton,  Elkmont,  Woodville, 
Piedmont,  Natural  Bridge,  Greensboro,  Reform,  Wilsonville,  Tal- 
ladega Mountains,  Mount  Weogufka,  Hayneville,  Thomasville,  Myr- 
tlewood,  Carlton,  Jackson,  and  Abbeville.  At  Dothan,  Orange 
Beach,  and  Bayou  Labatre  it  occurs  rarely. 

Minks  are  semiaquatic  in  habit,  always  found  along  streams  or  in 
swamps  or  marshes.  They  are  rapid  swimmers,  perfectly  at  home 
in  the  water,  and  able  to  capture  with  comparative  ease  good-sized 
fish.  They  feed  also  on  frogs,  lizards,  crawfish,  rats,  mice,  and  rab- 
bits, and  are  destructive  of  ducks  or  other  marsh-dwelling  birds, 
and,  to  a  less  extent,  of  upland  birds.  Audubon  and  Bachman  state 
that  in  the  South  the  cotton  rat  furnishes  a  considerable  part  of 
their  food,  and  mention  on  one  occasion  seeing  a  mink  issuing  from 
a  hole  in  the  earth  dragging  by  the  neck  a  large  Florida  wood  rat. 
Minks  are  often  very  destructive  of  poultry,  especially  ducks.  Dr. 
Avery  mentions  a  case  where  a  mink  visited  a  poultry  house  at 
Greensboro  several  times  in  a  single  week  and  dragged  off  at  least  a 
dozen  small  hens,  which  it  devoured  beneath  an  outhouse. 

Spilogale  putorius  (Linnaeus). 

Alleghenian  Spotted  Skunk. 

Viverra  putorins  Linnaeus,  Syst.  Nat.,  ed.  10.  vol.  1,  p.  44,  1758. 
Spilogale  ringens  Merriam,  North  Ainer.  Fauna  No.  4,  p.  9,  1890  (Greensboro, 
Alabama). 

The  little  spotted  skunk  occurs  commonly  over  the  greater  part  of 
the  State,  where,  in  common  with  the  large  skunks,  it  is  known  as 
"polecat."  It  lives  chiefly  about  cultivated  lands,  the  borders  of 
brushy  swamps,  and  in  waste  lands  generally,  though  not  in  wet 
swamps  or  in  heavy  timber.     Its  burrows  are  dug  beneath  the  roots 

32  For  use  of  this  name,  cf.  Hollister,  Proc.  Biol.  Soc.  Washington,  vol.  27,  p.  215,  1914. 


38  NORTH   AMERICAN   FAUNA.  (No.  45 

of  a  tree  or  stump,  a  rock  or  slab  pile,  or  in  a  cut  bank.  These 
skunks  are  much  more  agile  than  their  larger  relatives  (Mephitis), 
and  readily  climb  small  trees  or  bushes  when  pursued  by  dogs. 

They  subsist  largely  on  insects  (particularly  beetles  and  grass- 
hoppers), mice  and  other  small  mammals,  lizards,  salamanders, 
crawfish,  and  occasionally  small  birds,  and  are  sometimes  destructive 
to  poultry  and  occasionally  to  peanuts.  Henry  Golson,  of  Autauga- 
ville,  informed  the  writer  in  November,  1916,  that  these  little  skunks 
had  gained  an  entrance  to  a  storehouse  in  a  field  where  he  keeps 
peanuts,  peas,  etc.,  and  had  eaten  a  considerable  quantity  of  the 
peanuts;  he  caught  several  of  the  animals  on  the  pile  of  peanuts; 
and  quantities  of  chewed  shells  which  they  had  left  were  in  evidence. 

This  observation  is  confirmed  by  one  related  by  the  late  Dr.  Avery, 
of  Greensboro,  in  a  letter  dated  November  7,  1893,  stating — 

Last  summer  I  was  told  that  something  was  destroying  the  ground  peas  of  a 
neighbor.  Some  steel  traps  were  set  and  a  little  striped  skunk  was  caught. 
*  *  *  The  skunks  were  eating  the  ground  peas  and  eight  were  caught  in  my 
traps  in  less  than  two  months.  The  stomachs  of  these  skunks  contained  ground 
peas  mixed  with  the  debris  of  insects.  I  found  a  persimmon  seed  in  the  rec- 
tum of  a  skunk  of  which  I  made  a  skin. 

The  late  Dr.  Charles  Mohr,  in  a  letter  dated  June  1,  1890,  gives  in- 
teresting original  notes  on  the  habits  of  this  little  skunk  near  Mobile ; 
he  says: 

I  will  give  some  notes  on  the  habits  of  the  small  striped  skunk,  and  in  par- 
ticular describe  the  manner  in  which  it  disposes  of  the  eggs  which  it  is  very- 
fond  of  stealing  at  its  nightly  visits  to  the  chicken  house.  Its  proceedings,  in 
reaching  its  aim,  seem  to  me  strange,  in  fact  almost  incredible.  I  have,  however, 
the  facts  from  an  intelligent  and  close  observer,  a  personal  friend  of  mine,  one 
whose  word  I  can  not  doubt.  He  says  the  skunk  is  unable  to  open  the  egg  by 
the  aid  of  its  teeth  or  to  take  hold  of  it  with  its  mouth  in  order  to  carry  it 
away.  It  removes  the  egg  from  the  nest,  rolls  it  with  the  front  paws  to  a  place 
presenting  a  solid,  hard  surface,  then  the  egg  is  taken  in  its  paws,  the  animal 
assumes  an  erect  posture,  lifts  it  from  the  ground,  then  lets  it  drop  from  the 
height  of  its  body  to  insure  its  breakage  in  striking  the  hard  ground. 

This  animal  is  at  nearly  all  seasons  a  frequent  visitor  to  the  basements  of 
the  premises  not  only  in  the  country  but  in  the  suburbs  of  the  city.  If  not 
disturbed  it  becomes  quite  fearless.  My  friend  tells  me  it  hunts  mice,  young 
rats,  roaches,  etc.,  and  in  that  way  is  of  benefit  when  there  are  no  hen  nests 
to  be  depredated  upon. 

Specimens  of  the  little  spotted  skunk  have  been  examined  from 
Leighton,  Greensboro,  Autaugaville,  Prattville,  Mobile,  and  Ashford. 
It  is  reported  to  occur  also  at  Fort  Payne,  Piedmont,  Ardell,  Squaw 
Shoals,  Talladega  Mountains,  Reform,  Jackson,  Castleberry,  and  Bon 
Secour.    It  seems  to  be  unknown  at  Orange  Beach. 


1921.]  MAMMALS   OF   ALABAMA SKUNKS.  39 

Mephitis  mephitis  nigra  (Peale  and  Beauvois).83 
Eastern  Skunk. 

Viverra  nigra  Peale  and  Beauvois,  Cat.  Peale's  Mus.,  p.  37,  1796. 

The  common  skunk,  or  "  polecat,"  is  found  in  moderate  numbers 
throughout  the  State,  but  the  recent  high  prices  of  fur  have  led  to  a 
considerable  reduction  in  its  numbers.  Skunks  den  in  burrows  of  their 
own  construction  or  in  deserted  burrows  of  other  animals,  choosing 
for  their  headquarters  a  thicket  or  brushy  fence  corner  in  a  field,  a 
rock  pile  or  slab  pile,  a  gully  or  wash,  or  some  similar  protected  loca- 
tion. They  frequently  burrow  beneath  a  house,  barn,  or  shed,  and  in 
such  situations  are  liable  to  become  very  objectionable  to  human  resi- 
dents. 

The  food  of  skunks  consists  largely  of  insects,  small  mammals, 
reptiles,  and  amphibians.  Almost  any  animal  food  is  relished  by 
them  and  even  carrion  is  frequently  eaten.  While  occasional  raids 
are  made  on  the  poultry  yard  or  on  the  nests  of  wild  birds,  their  de- 
struction of  insects  and  noxious  mammals  doubtless  more  than  offsets 
any  damage  they  may  do  to  poultry  or  game. 

Specimens  examined  from  Ardell,  Reform,  Squaw  Shoals,  and 
Jackson  indicate  that  the  skunk  of  northern  and  central  Alabama  is 
fairly  typical  of  nigra.  As  in  other  regions,  there  is  great  variation 
in  the  coloration,  ranging  from  an  animal  with  broad  white  stripes 
the  entire  length  of  the  body  to  one  nearly  black  except  for  a  patch 
of  white  on  the  nape  and  shoulders. 

Skunks  are  reported  as  occurring  in  moderate  numbers  at  Leighton, 
Elkmont,  Woodville,  Sand  Mountain,  Fort  Payne,  Talladega  Moun- 
tains, Anniston,  Jasper,  Auburn,  York,  Thomasville,  and  Teasley 
Mill  (Montgomery  County). 

Mephitis  mephitis  elongata  Bangs. 

Florida  Skunk. 

Mephitis  mephitica  elongata  Bangs,  Proc.  Boston  Soc.  Nat.  Hist.,  vol.  26,  p.  531, 
1895. 

The  Florida  skunk,  characterized  by  long  tail  and  rather  heavy 
skull,  occurs  in  the  coast  region  of  Alabama.  Three  specimens  from 
Perdido  Bay  are  all  that  are  available,  so  that  the  northern  limits  of 
the  range  of  the  subspecies  are  unknown.  Skunks  reported  from 
Ashford,  Bon  Secour,  Bayou  Labatre,  Mobile,  and  Castleberry  are 
provisionally  referred  to  this  race.  On  Dauphin  Island,  in  February, 
1912,  numerous  tracks  of  this  animal  were  seen  about  logs  and  drift- 
wood along  the  shores  of  the  Sound. 

88  For  use  of  this  name,  cf.  Hollister,  Proc.  Biol.  Soc.  Washington,  vol.  27,  p.  215,  1914. 


40  NORTH   AMERICAN   FAUNA.  [No.  45 

Lutra  canadensis  canadensis   (Schreber). 
Otter. 

Mustcla  lutra  canadensis  Schreber,  Siiugthiere,  pi.  126  B,  1776. 

The  otter  is  generally  distributed  in  the  State,  but  in  most  places 
occurs  rather  sparingly.  It  is  perhaps  most  numerous  in  the  middle 
and  southern  counties,  but  is  everywhere  decreasing  rapidly  through 
persistent  trapping.  It  is  an  exceedingly  shy  and  retiring  creature, 
choosing  for  its  home  the  most  remote  and  unfrequented  swamps, 
streams,  or  ponds. 

The  species  is  reported  to  occur  at  Muscle  Shoals,  Ardell,  Myrtle- 
wood,  Oakchia,  Wilsonville,  Mount  Weogufka,  Hajmeville,  Teasley 
Mill  (Montgomery  County),  Jackson,  Carlton,  Mobile,  Bayou 
Labatre,  Bon  Secour,  Point  Clear,  Orange  Beach,  Ashford,  and  Abbe- 
ville. A  trapper  at  Whistler  caught  four  otters  in  the  creek  near  his 
home  during  the  winter  of  1911-12;  Will  Matthews  shot  four  at  one 
time  near  Castleberry,  in  1911,  as  they  were  playing  around  a  log 
in  the  creek. 

The  only  specimen  at  hand  is  one  caught  by  a  trapper  in  Mobile 
River,  near  the  Louisville  &  Nashville  Railroad  bridge,  in  January, 
1917,  and  with  so  little  material  it  is  impossible  satisfactorily  to  define 
the  status  of  the  Alabama  otters.  The  single  specimen  is  very  dark 
blackish  brown,  shaded  about  the  lips  and  throat  with  grayish,  thus 
agreeing  in  color  with  the  otters  of  the  Northern  States,  and  differing 
from  the  Florida  form  (vaga),  which  is  decidedly  redder;  the  skull 
of  the  Alabama  specimen,  however,  approaches  that  of  vaga  in  being 
relatively  long  and  narrow,  with  the  postorbital  region  constricted. 
This  specimen  weighed  IT  pounds  in  the  flesh.  A  skin,  taken  near 
Jackson,  which  I  examined  in  a  dealer's  store,  was  also  of  the  same 
dark  color,  and  the  animals  referred  to  above,  killed  by  Mr.  Mat- 
thews, were  described  as  being  black. 

Otters  are  almost  as  aquatic  as  seals,  which,  indeed,  they  somewhat 
resemble  in  appearance  and  actions  in  the  water.  In  the  Southern 
States,  according  to  Audubon  and  Bachman,  the  young  are  brought 
forth  about  the  middle  of  March.34  The  habits  of  the  species  are  well 
described  by  Stone  and  Cram,  as  follows : 

When  traveling  overland  otters  follow  the  smoothest  course  they  can  find, 
going  round  stumps  and  hummocks  and  beneath  logs  in  preference  to  climbing 
over  them.  Following  the  same  course  week  after  week,  often  in  families  of 
four  or  five  together,  they  soon  establish  a  distinct  path  clear  of  obstacles; 
crooked  and  tortuous  yet  keeping  to  the  same  general  direction,  and  in  most 
cases  leading  to  some  rapid  or  springhole  beneath  the  bank  where  the  water 
seldom  freezes. 

Otters  are  beautiful  swimmers ;  they  glide  and  shoot  along  through  the  water, 
twisting  and  turning  like  the  fish  they  so  delight  in  chasing.     I  have  seen  one 

31  Audubon  and  Bachman,  Op.  cit.,  vol.  2,  p.  10. 


1921.]  MAMMALS  OF  ALABAMA — COUGARS.  41 

pursuing  a  muskrat,  as  a  pickerel  pursues  a  shiner,  splashing  through  the  shal- 
low water  where  the  stream  had  overflowed  its  banks.  At  times  both  would  be 
invisible  beneath  the  surface  for  several  minutes,  to  appear  again  perhaps  out 
in  the  current  at  a  distance,  the  muskrat  always  diving  and  dodging  for  its  life. 

Otters  will  also  catch  wild  ducks  on  the  water,  raising  and  seizing  them  from 
beneath.  They  catch  their  fish  by  fairly  swimming  them  down  in  spite  of  all 
their  twisting  and  darting.     *     *     * 

The  otter's  home  is  a  den  beneath  the  bank,  usually  with  the  entrance  under 
water  for  safety.  This  is  evidently  not  regarded  as  absolutely  essential,  how- 
ever, for  otters  have  been  known  to  have  their  nests  in  caves,  high  up  in  the 
banks  and  at  the  bottom  of  hollow  trees.     *     *     * 

They  get  the  greatest  fun  from  sliding;  where  the  bank  is  sufficiently  steep 
and  slanting  they  make  a  roundabout  path  leading  up  to  the  top  of  the  bank 
and  from  there  they  slide  down  the  slippery  surface  into  the  water  one  after  an- 
other like  boys  sliding  down  hill  on  the  snow.34 

Family  FELIDAE :  Cougars,  Wildcats,  etc. 

Felis  couguar  Kerr. 

Cougar  ;  Panther  ;  Puma. 

Felis  couguar  Kerr,  Animal  Kingdom,  p.  151,  1792. 

The  cougar,  or  "  panther,"  as  this  animal  is  usually  called,  doubtless 
in  early  times  occupied  the  greater  part  of  the  State ;  it  is  now  nearly, 
if  not  quite,  exterminated.  Bartram,  writing  in  1791,  speaks  of 
"  tygers  "  as  occurring  on  the  lower  Tombigbee  River,  this  statement 
probably  referring  to  the  cougar.35  Hodgson  records  one  killed  in 
March,  1820,  on  "  Ouchee  Creek,"  and  mentions  seeing  a  mounted 
specimen  near  Blakeley,  Baldwin  County,  the  same  year.36  Hallock 
in  1877  reported  the  "  panther  "  occasional  in  Dekalb  County.37  An 
old  resident  of  Sand  Mountain,  near  Carpenter,  reported  seeing 
one  there  some  20  years  ago  [1896].  Recent  reports,  although  rather 
indefinite,  indicate  that  a  very  few  may  still  remain  in  the  big  swamps 
of  the  southern  counties.  Tracks  of  two  of  these  animals  were  seen 
by  an  experienced  trapper  about  1912  in  Big  lichee  Creek  Swamp, 
near  Seale,  and  one  is  reported  to  have  been  seen  about  1905  in  the 
region  of  Nigger  Lake,  Baldwin  County.  Lack  of  any  specimens 
from  the  State  makes  it  impossible  to  say  which  form  of  the  species 
occurs  here. 

The  cougar  is  a  shy,  retiring  beast,  of  cowardly  disposition,  inhabit- 
ing the  wildest  parts  of  the  forest,  the  cliffs  among  the  mountains, 
and  the  deep  canebrakes  of  the  river-bottom  swamps.  With  the 
advent  of  settlers  it  quickly  disappeared  from  the  vicinity  of  civiliza- 
tion and  retired  to  more  secluded  regions. 


34  Stone  and  Cram,  American  animals,  pp.  220-223,  1902. 

85  Bartram,  William,  Travels,  p.  410,  1792. 

36  Hodgson,  Letters  from  North  America,  vol.  1,  pp.  123,  149,  1S34. 

87  Hallock,  Charles,  Sportsman's  Gazetteer,  p.  3  [of  Sportsman's  Directory],  1877. 


42  NORTH   AMERICAN   FAUNA.  [No.  45 

Roosevelt  describes  the  habits  of  the  cougar  as  follows : 

In  its  essential  habits  and  traits,  the  big,  slinking,  nearly  uni-colored  cat 
seems  to  be  much  the  same  everywhere,  whether  living  in  mountain,  open  plain, 
or  forest,  under  arctic  cold  or  tropic  heat.  When  the  settlements  become  thick, 
it  retires  to  dense  forest,  dark  swamp  or  inaccessible  mountain  gorge,  and  moves 
about  only  at  night.  In  wilder  regions  it  not  infrequently  roams  during  the  day 
and  ventures  freely  into  the  open.  Deer  are  its  customary  prey  where  they  are 
plentiful,  bucks,  does,  and  fawns  being  killed  indifferently.  Usually  the  deer 
is  killed  almost  instantaneously,  but  occasionally  there  is  quite  a  scuffle,  in 
which  the  cougar  may  get  bruised,  though,  as  far  as  I  know,  never  seriously. 
It  is  also  a  dreaded  enemy  of  sheep,  pigs,  calves,  and  especially  colts,  and 
when  pressed  by  hunger  a  big  male  cougar  will  kill  a  full-grown  horse  or  cow, 
moose  or  wapiti.  It  is  the  special  enemy  of  mountain  sheep.  In  1886,  while 
hunting  white  goats  north  of  Clarke's  fork  of  the  Columbia,  in  a  region  where 
cougar  were  common,  I  found  them  preying  as  freely  on  the  goats  as  on  the 
deer.  It  rarely  catches  antelope,  but  is  quick  to  seize  rabbits,  other  small 
beasts,  and  even  porcupines. 

No  animal,  not  even  the  wolf,  is  so  rarely  seen  or  so  difficult  to  get  without 
dogs.  On  the  other  hand,  no  other  wild  beast  of  its  size  and  power  is  so 
easy  to  kill  by  the  aid  of  dogs.  There  are  many  contradictions  in  its  character. 
Like  the  American  wolf,  it  is  certainly  very  much  afraid  of  man ;  yet  it  habitu- 
ally follows  the  trail  of  the  hunter  or  solitary  traveller,  dogging  his  footsteps, 
itself  always  unseen.  I  have  had  this  happen  to  me  personally.  When  hungry 
it  will  seize  and  carry  off  any  dog;  yet  it  will  sometimes  go  up  a  tree  when 
pursued  even  by  a  single  small  dog  wholly  unable  to  do  it  the  least  harm.  It 
is  small  wonder  that  the  average  frontier  settler  should  grow  to  regard  almost 
with  superstition  the  great  furtive  cat  which  he  never  sees,  but  of  whose  pres- 
ence he  is  ever  aware,  and  of  whose  prowess  sinister  proof  is  sometimes  afforded 
by  the  deaths  not  alone  of  his  lesser  stock,  but  even  of  his  milch  cow  or  saddle 
horse.38 

Mr.  Roosevelt  states  also  that  the  cougar  has  been  known  to  attack 
human  beings,  but  authentic  instances  of  such  attacks  are  exceedingly 
rare. 

Lynx  ruffus  floridanus  Rafinesque. 

Florida  Wildcat  ;  Bobcat  ;  Catamount. 

Lynx  floridanus  Rafinesque,  Amer.  Monthly  Mag.,  vol.  2,  p.  46,  1817. 

Wildcats  occur  in  moderate  numbers  throughout  the  wilder  parts 
of  the  State.  In  the  mountains  they  live  about  the  gulches  and  rocky 
bluffs,  while  in  the  lowlands  they  seek  the  shelter  of  the  swamps  and 
of  brushy  thickets.  On  the  outer  beach  bordering  Perdido  Bay  nu- 
merous tracks  were  seen  in  the  palmetto  scrub,  and  the  animals  are 
trapped  every  winter  in  this  vicinity.  Specimens  from  Orange  Beach 
and  Castleberry  agree  closely  with  specimens  of  floridcmus  from 
Florida ;  the  race  is  characterized  by  dark  colors,  with  abundant,  dis- 
tinct blackish  spots  on  the  body  and  legs  and  narrow  blackish  streaks 
along  the  median  line  of  the  back. 

38  Roosevelt,  Theodore,  With  the  cougar  hounds :  Scribner's  Mag.,  vol.  30,  no.  4,  pp. 
431-432,  October,  1901. 


1921.1  MAMMALS   OF   ALABAMA MICE.  43 

Whether  this  race  ranges  throughout  the  State  or  whether  typical 
rujfus  occurs  in  the  northern  part  can  not,  in  the  absence  of  speci- 
mens,' be  determined.  Wildcats  are  reported  from  many  localities, 
including  Sand  Mountain,  Talladega  Mountains,  Mount  Weogufka, 
Piedmont,  Ardell,  Natural  Bridge,  Autaugaville,  Hayneville,  Seale, 
Oakchia.  Myrtlewood,  Jackson,  Carlton,  Abbeville,  Dothan,  and 
Mobile. 

These  cats  destroy  many  young  pigs  and  some  lambs,  kids,  and 
poultry.  Hunters  secure  the  animals  both  by  chasing  with  hounds 
and  by  trapping.  Audubon  and  Bachman  describe  the  habits  of  the 
wild  cat  in  the  Southern  States  as  follows : 

It  is  abundant  in  the  Caneb rakes  (patches  or  thickets  of  the  Miegia  macro- 
sperma,  of  Michaux,  which  often  extend  for  miles,  and  are  almost  impassable) 
bordering  the  lakes,  rivers,  and  lagoons  of  Carolina,  Louisiana,  and  other  South- 
ern and  South  Western  States.  This  species  also  inhabits  the  mountains  and 
the  undulating  or  rolling  country  of  the  Southern  States,  and  frequents  the 
thickets  that  generally  spring  up  on  deserted  cotton  plantations,  some  of  which 
are  two  or  three  miles  long,  and  perhaps  a  mile  wide,  and  afford,  from  the  quan- 
tity of  briers,  shrubs,  and  young  trees  of  various  kinds  which  have  overgrown 
them,  excellent  cover  for  many  quadrupeds  and  birds.  In  these  bramble-covered 
old  fields,  the  "  Cats  "  feed  chiefly  on  the  rabbits  and  rats  that  make  their  homes 
in  their  almost  impenetrable  and  tangled  recesses ;  and  seldom  does  the  cautious 
Wild  Cat  voluntarily  leave  so  comfortable  and  secure  a  lurking  place,  except  in 
the  breeding  season,  or  to  follow  in  very  sultry  weather,  the  dry  beds  of  streams 
or  brooks,  to  pick  up  the  cat-fish,  etc.,  or  cray-fish  and  frogs  that  remain  in  the 
deep  holes  of  the  creeks,  during  the  drought  of  summer. 

The  Wild  Cat  not  only  makes  great  havoc  among  the  chickens,  turkeys,  and 
ducks  of  the  planter,  but  destroys  many  of  the  smaller  quadrupeds,  as  well  as 
partridges,  and  such  other  birds  as  he  can  surprise  roosting  on  the  ground. 
*     *     * 

The  domicile  of  the  Wild-Cat  is  sometimes  under  an  old  log,  covered  with 
vines  such  as  the  Smilax,  Ziziplwis  volubilus,  Rubus,  etc.,  but  more  commonly  in 
a  hollow  tree.  Sometimes  it  is  found  in  an  opening  twenty  or  thirty  feet  high, 
but  generally  much  nearer  the  ground,  frequently  in  a  cavity  at  the  root,  and 
sometimes  in  the  hollow  trunk  of  a  fallen  tree,  where,  after  collecting  a  con- 
siderable quantity  of  long  moss  and  dried  leaves  to  make  a  comfortable  lair,  it 
produces  from  two  to  four  young.  These  are  brought  forth  in  the  latter  end  of 
March  in  Carolina ;  in  the  Northern  States,  however,  the  kittens  appear  later, 
as  we  have  heard  of  an  instance  in  Pennsylvania  where  two  young  were  found 
on  the  15th  day  of  May,  apparently  not  a  week  old.38 

Order  RODENTIA. 

Family  MURIDAE:  Mice,  Rats,  etc. 

Reithrodontomys  humulis  merriami  Allen. 

Meekiam  Harvest  Mouse. 

Reithrodontomys  merriami  J.  A.  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  vol.  7,  p. 
119,  1895. 

The  Merriam  harvest  mouse  is  known  from  only  a  few  localities, 
but  probably  ranges  nearly  throughout  the  State.     It  inhabits  old 

39  Audubon  and  Bachman,  Op.  cit.,  vol.  1,  pp.  6,  13. 


44  NORTH  AMERICAN   FAUNA.  [No.  45 

fields  and  the  brushy  borders  of  cultivated  land,  selecting  usually 
for  its  home  a  thick  tangle  of  matted  grass,  weeds,  or  briers,  often 
in  a  wet  bottom  or  the  edge  of  a  marsh. 

At  York,  in  February,  1912,  these  mice  were  numerous  in  wet 
patches  of  broom  sedge  and  about  the  brushy  borders  of  fields.  Holt 
has  taken  a  number  of  specimens  on  the  prairie  at  Barachias  in  dry 
broomsedge  fields,  where  they  travel  in  runways  made  by  cotton 
rats.  A  few  were  caught  by  him  also  at  Carlton,  Jackson,  and  Dean 
(Clay  County) ;  and  the  writer  took  one  near  Autaugaville. 

Heithrodontomys  humulis  liumulis  (Bachman). 

Eastebn  Harvest  Mouse. 

Mus  humulis  Bachman,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  p.  97,  1841. 

Two  specimens  of  the  eastern  harvest  mouse,  trapped  in  weedy 
fields  on  Sand  Mountain,  in  the  extreme  northeastern  corner  of 
Jackson  County,  at  an  elevation  of  1,500  feet,  are  decidedly  paler 
than  specimens  from  the  central  part  of  the  State,  and  although 
not  typical  of  humulis,  they  are  best  referred  to  that  form.  The 
older  one  of  the  two  (an  adult  female)  is  less  intensely  brownish 
than  typical  examples  of  humulis,  being  strongly  shaded  with  fus- 
cous; the  younger  is  uniform  fuscous  above. 

In  habits  this  subspecies  does  not  differ  from  merriami,  which  occu- 
pies most  of  Alabama.  Audubon  and  Bachman  state  that  its  food 
consists  largely  of  the  seeds  of  wild  grasses,  they  having  found  in 
its  nest  small  stores  of  seeds  of  broom  grass,  crab  grass,  and  other 
meadow  grasses.40 

Peromyscus  polionotus  polionotus  (Wagner). 

Old-field  Mouse. 

Mus  polionotus  Wagner,  Wiegmann's  Arch.  f.  Naturg.,  IX,  vol.  2,  p.  52,  1843. 

The  little  old-field  mouse  occurs  rather  commonly  in  suitable  situa- 
tions throughout  the  eastern,  central,  and  northeastern  parts  of  the 
State.  It  ranges  northward  to  the  extreme  northeastern  corner  of 
Jackson  County,  but  apparently  is  not  found  north  of  the  Tennessee 
River.  Its  western  limit  is  not  known  with  certainty;  it  has  been 
taken  at  Centerville,  Bibb  County,  but  rather  careful  trapping  at 
Reform,  Pickens  County,  and  Jackson,  Clarke  County,  failed  to 
secure  it  (fig  1). 

At  Abbeville  it  was  found  abundant  in  a  neglected  sandy  field  on 
a  hillside,  where  cactus  grows  in  abundance  but  grasses  only  sparsely. 
Where  cactus  was  most  abundant  the  mice  were  most  numerous. 
Their  burrows  were  found  at  frequent  intervals,  each  one  marked 

40  Audubon  and  Bachman,  Op.  cit.,  vol.  2,  p.  105. 


1921.] 


MAMMALS   OF   ALABAMA MICE. 


45 


by  a  heap  of  soil,  usually  about  6  or  8  inches  across  and  2  inches  high. 
The  entrance  holes  were  of  irregular  shape,  about  1^  inches  in 
diameter;  a  few  were  found  plugged  with  sand,  but  the  majority 
were  open.  No  definite  trails  were  observed  in  this  colony  and  the 
soil  was  too  hard  to  show  tracks  well. 

In  the  sandy  fields 
of  Autauga  County 
this  species  is  appar- 
ently more  abundant 
than  in  other  sec- 
tions visited.  Here 
they  live  chiefly  in 
cultivated  land,  par- 
ticularly in  corn- 
fields, but  range  also 
into  open  timbered 
tracts.  Numerous 
burrows,  scratched 
out  by  the  mice,  are 
seen  in  both  these 
situations,  and  fresh 
scratchings  observed 
in  the  cornfields  seem 
to  indicate  that  the 
mice  dig  up  some 
freshly  planted  corn. 
The  holes  are  in  some 
cases  considerably 
larger  than  seems 
necessary  for  the 
mouse  to  enter  and 
the  piles  of  dirt  in- 
dicate burrows  of 
considerable  depth. 
L.  S.  Golsan,  how- 
ever, frequently  has 
dug  out  the  mice 
with  his  hands. 

On  Sand  Mountain  these  mice  were  found  in  fair  numbers  in  two 
localities — Bucks  Pocket,  near  Grove  Oak,  and  at  a  point  about  9 
miles  southeast  of  Carpenter  Station.  Here  they  live  in  cultivated 
fields  of  cotton  and  corn,  preferring  sandy  soils,  but  ranging  into 
some  that  are  rather  clayey  and  also  to  some  extent  into  weedy  hedge- 
rows and  the  edges  of  open  timber.  In  Greenbrier  Cove,  on  Cane 
Creek,  Marshall  County,  they  were  common  in  similar  situations. 


nfa  Beach 


Fig.    1. — Provisional   distribution    of    races   of    the   old-field 
mouse    in    Alabama. 

1.  Peromyscus  polionotus  polionotus. 

2.  Peromyscus  polionotus  albifrons. 
(Limits  of  range  imperfectly  known.) 


46  NOETH   AMERICAN   FAUNA.  [No.  45 

Little  is  known  of  the  food  habits  of  this  species,  but  it  is  believed 
to  feed  largely  on  the  seeds  of  grasses,  weeds,  and  grain;  several 
stomachs  from  Abbeville  contained  remains  of  blackberries  with 
other  finely  chewed  food. 

The  Alabama  examples  of  this  race  agree  well  with  specimens  from 
Georgia — the  type  region. 

Peromyscus  polionotus  albifrons  Osgood. 

White-feonted  Beach  Mouse. 

Peromyscus  polionotus  albifrons  Osgood,  North  Ainer.  Fauna  No.  28,  p.  108, 1909. 

The  white-fronted  beach  mouse,  a  pale  race  of  the  old-field  mouse, 
is  confined  in  Alabama,  so  far  as  is  known,  to  the  drifting  sand  dunes 
along  the  coast  of  Baldwin  County ;  it  may  occur,  however,  in  sandy 
tracts  at  some  distance  from  the  coast  and  doubtless  intergrades  with 
Peromyscus  p.  polionotus  in  some  of  the  southern  counties.  Mobile 
Bay  apparently  forms  a  barrier  to  its  westward  distribution,  as  no 
signs  of  its  presence  have  been  found  on  the  beaches  of  Mobile  County 
or  on  the  Gulf  coast  of  Mississippi  (fig.  1). 

In  the  dunes  which  extend  along  the  Gulf  coast  from  the  Little 
Lagoon  eastward  to  the  mouth  of  Perdido  Bay  these  mice  occur  in 
abundance  in  situations  where  their  pale  colors  harmonize  with  the 
very  light-colored  sand.  They  seem  to  be  most  numerous  in  the 
line  of  dunes  nearest  the  surf,  where  the  cover  is  very  sparse,  con- 
sisting of  stunted  live-oak  bushes,  yaupon,  pokeberry,  patches  of 
"  sea  oats  "  ( Uniola  paniculata) ,  and  a  few  low  herbaceous  plants. 
Here  their  tracks  and  trails  are  seen  everywhere  in  the  sand,  leading 
in  and  out  among  the  clumps  of  sea-oats  or  from  one  clump  of  bushes 
to  another. 

The  mice  dig  little  burrows  in  the  sand,  usually  beneath  a  bush, 
the  entrance  holes  being  usually  small  and  round,  though  sometimes 
larger  than  the  size  of  the  animal.  Most  of  these  entrances  are  left 
open,  but  a  few  were  found  closed  with  sand.  On  the  rolling  sand 
flats  nearer  the  bay,  where  the  growth  of  bushes  and  palmetto  scrub 
is  more  dense,  the  mice  were  found  in  smaller  numbers.  The  stom- 
ach of  one  contained  remains  of  red  berries,  and  of  another  finely 
chewed  vegetable  matter,  probably  seeds. 

Peromyscus  leucopus  leucopus  (Rafinesque). 

White-footed  Mouse. 

Musculus  leucopus  Rafinesque,  Aruer.  Monthly  Mag.,  vol.  3,  p.  446,  1818. 

The  white- footed  mouse  is  one  of  the  commonest  and  most  widely 
distributed  species  of  eastern  United  States,  the  typical  race  ranging 


1921.] 


MAMMALS   OF   ALABAMA MICE. 


47 


from  Virginia  to  Arkansas  and  Louisiana ;  in  Alabama,  however,  it  is 
restricted,  so  far  as  known,  to  the  northern  half  of  the  State,  from 
Montgomery  County  northward,  and  is  in  general  less  numerous  than 
in  the  more  northern  parts  of  its  range.  Specimens  have  been  taken 
at  Muscle  Shoals,  Leighton,  Woodville,  Stevenson,  Sand  Mountain 
(near  Carpenter),  Erin  (Clay  County),  Choccolocco  Mountain, 
Greensboro,  and 
Barachias  (fig.  2). 

This  species  lives 
in  a  great  variety  of 
situations,  but  usu- 
ally in  or  near  tim- 
ber tracts.  It  is  not 
so  fond  of  the 
swampy  bottom- 
lands as  its  relative, 
the  cotton  mouse,  but 
is  parial  to  upland 
woods,  the  borders 
of  cultivated  fields 
and  brushy  hedge 
rows;  it  ranges  also 
up  to  the  summits  of 
the  mountains,  where 
it  lives  in  rock  piles 
and  crevices  in  the 
cliffs.  At  Leighton, 
while  hunting  for 
wood  rats  (Neotoma) 
along  the  osage- 
orange  hedges,  the 
writer  punched  two 
of  these  mice  out 
of  a  large  rat  nest 
about  10  or  12  feet 
high,    among    the 

branches  of  one  of  the  trees.  This  species,  like  the  cotton  mouse, 
often  dwells  in  hollow  logs  or  stumps,  or  sometimes  in  hollow  trees 
at  no  great  distance  from  the  ground.  At  times  it  appropriates  the 
deserted  nest  of  a  bird  in  a  bush  or  low  tree,  adding  to  it  sufficient 
material  to  make  a  warm,  covered  nest;  it  is  said,  also,  occasionally 
to  construct  a  complete  nest  of  its  own  in  the  branches  of  a  bush,  5 
to  15  feet  from  the  ground.  Audubon  and  Bachman  describe  such 


n$e  Be»ch 


Fig.  2. — Distribution  of  the  white-footed  mouse  (Pcromyscus 
leucopus  leucopus)   in  Alabama. 


48  NORTH   AMERICAN   FAUNA.  [No.  45 

nests  as  being  of  oval  shape,  7  inches  long  and  4  inches  broad,  com- 
posed of  dried  moss,  shreds  of  grape  vine  bark,  and  dry  leaves.41 
The  food  of  this  mouse  is  described  by  the  same  authors  as  follows : 

In  its  wild  state  it  is  continually  laying  up  little  stores  of  grain  and  grass 
seeds.  We  have  seen  it  carrying  in  its  mouth  acorns  and  chinquapins.  In  the 
Northern  States  these  little  hoards  are  often  composed  wholly  of  wheat;  in 
the  South,  of  rice.  This  species,  like  all  rats  and  mice,  is  fond  of  Indian-corn, 
from  which  it  only  extracts  the  choicest,  sweetest  portions,  eating  the  heart 
and  leaving  the  rest,  untouched.42 

Stone  and  Cram  state  that  the  white- footed  mouse  climbs  for  rose 
hips  and  red  alder  berries  and  also  gathers  and  stores  the  seeds  of 
the  garget.43 

Peromyscus  gossypinus  gossypinus  (LeConte). 
Cotton  Mouse. 

Hespcromys  gossypinus,  LeConte,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  vol.  6, 
p.  411,  1853. 

The  cotton  mouse  is  probably  the  most  abundant  and  widely  dis- 
tributed mammal  in  the  State,  where  it  is  often  called  "  wood  rat." 

The  name  "  cotton  mouse  "  suggests  a  habitat  in  cotton  fields,  but 
this  species  is  scarcely  ever  found  in  such  places,  unless  the  fields  are 
on  the  borders  of  a  timbered  swamp.  It  is  a  typical  timber  mouse 
and  most  abundant  in  the  heavy  swamps  of  the  river  bottoms;  it  is 
found  also  in  upland  timber,  and  wherever  rocky  ledges  or  bluffs 
occur  it  utilizes  the  shelter  afforded  by  the  crevices  or  caves  in  these 
formations.  In  the  swamps,  hollow  trees  or  logs,  or  rotten  stumps 
and  woodpiles  furnish  convenient  retreats.  LeConte,  who  discovered 
and  named  this  species,  says  of  it : 

It  forms  its  nest  under  logs  and  under  the  bark  of  decaying  trees,  generally 
of  cotton,  frequently  using  more  than  a  pound  of  this  material  for  its  purpose.44 

Little  is  known  of  the  food  habits  of  this  mouse,  but  in  view  of  its 
close  relationship  to  Peromyscus  leucopus  its  food  is  probably  similar, 
consisting  largely  of  wild  berries,  nuts,  seeds,  and  such  insects  as  can 
be  obtained.  The  mice  are  frequently  captured  in  traps  baited  with 
either  salt  or  fresh  meat. 

The  typical  race  (gossypinus)  has  been  taken  at  Orange  Beach, 
Point  Clear,  Ashford,  Castleberry,  Carlton,  and  Jackson.  It  may 
range  somewhat  farther  north  than  these  records  indicate,  but 
material  is  lacking  to  show  its  exact  northern  limit  (fig.  3).  Inter- 
gradation  with  the  northern  race  (megacephalus)  takes  place  in  the 
region  between  Castleberry  and  Montgomery. 

41  Audubon  and  Bachruan,  Op.  cit.,  vol.  1,  p.  302. 

42  Op.  cit.,  p.  303. 

43  Stone  and  Cram,  American  animals,  pp.   132-133,  1902. 

«*  LeConte,  John,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  vol.  6,  p.  411,  1853. 


1921.] 


MAMMALS   OF   ALABAMA — MICE. 


49 


Peromyscus  gossypinus  megacephalus  (Rhoads). 

Rhoads  Cotton  Mouse. 

Sitomys  megacephalus  Rhoads,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  p.  254,  1894. 

The  Rhoads  cotton  mouse,  distinguished  chiefly  by  larger  size,  occu- 
pies the  northern  half  of  the  State,  south  at  least  to  Autauga  County. 
The  type  specimen 
came  from  Wood- 
ville,  and  a  fine 
series  of  32  topo- 
types  in  winter  pel- 
age (March)  secured 
by  the  Biological 
Survey  includes  a 
number  of  old  adults 
as  well  as  younger 
individuals.  Speci- 
mens have  been 
taken,  also,  at  Muscle 
Shoals,  Leighton, 
Scottsboro,  Sand 
Mountain  (near  Car- 
penter Station), 
Cane  Creek  (Mar- 
shall County),  Gun- 
tersville,  Bucks 
Pocket,  Fort  Payne, 
Attalla,  Ardell,  Mel- 
ville, Squaw  Shoals, 
Erin  (Clay  County) , 
Autauga  ville,  E 1  - 
more,  and  Mont- 
gomery (fig.  3). 

The  series  of  12 
from  Autaugaville 
(taken  chiefly  i  n 
and  around  Bear 
Swamp)  is  clearly 
referable  to  megace- 


-Distribution    of    races    of    the    cotton    mouse    in 
Alabama. 

1.  Peromyscus  gossypinus  megacephalus. 

2.  Peromyscus  gossypinus  gossypinus. 
(Unshaded  area  represents  region  of  intergradation  from 

which  no  specimens  have  been  examined.) 


phalus,  but  two  specimens  from  Montgomery  indicate  that  in  this 
region  intergradation  with  typical  gossypinus  takes  place.  A  series 
of  5  specimens  from  Guntersville  also  are  decidedly  smaller  than 
typical  megacephalus  (the  hind  foot  averaging  about  21.5  mm.),  and 
thus  strongly  approach  gossypinus;  their  assignment  to  that  race, 
however,  is  inconsistent  with  our  knowledge  of  the  ranges  of  the  two 
forms  in  the  State,  the  latter  being  found  chiefly  in  the  coast  region. 

49663°— 21 4 


50  NORTH   AMERICAN   FAUNA.  [No.  45 

Occasional  specimens  taken  in  the  range  of  this  race  seem  to  be 
intermediate  in  characters  between  it  and  leucopus,  which  also  occu- 
pies in  part  the  same  territory.  These  specimens — two  from  Ardell 
and  one  each  from  Dean  and  Tidewater — resemble  leucopus  rather 
closely  in  external  appearance  and  measurements,  but  have  decidedly 
larger  skulls;  the  writer  regards  these  individuals  as  hybrids  be- 
tween the  two  species. 

In  habits  this  race  is,  of  course,  quite  similar  to  the  typical  form 
of  the  species  (gossypinus) .  Since  its  habitat  includes  much  hilly 
and  mountainous  country,  where  cliffs  and  rocky  bluffs  abound,  it  is 
very  frequently  found  about  caves,  crevices,  and  "  rock  houses  "  in 
company  with  the  Florida  wood  rat  or  the  Allegheny  cliff  rat.  In 
the  Big  Bear  Swamp  it  was  common  in  brushy  thickets,  living  in 
rotten  logs  and  stumps  not  only  around  the  borders  but  in  the  wet 
interior  of  the  swamp.  On  one  occasion  (Oct.  15,  1911)  L.  S.  Golsan 
captured  a  female  of  this  species  which  was  carrying  a  young  one  in 
her  mouth.  In  the  stomach  of  a  rattlesnake  killed  on  the  rocky, 
pine-covered  slope  of  Choccolocco  Mountain,  near  Piedmont,  Octo- 
ber 20,  1916,  the  writer  found  the  remains  of  a  mouse  of  this  species. 

Peromyscus  nuttalli  aureolus  (Audubon  and  Bachman). 

Southebn  Golden  Mouse. 

Mus  (Calomys)  aureolus  Audubon  and  Bachman,  Proc.  Acad.  Nat.  Sci.  Phila- 
delphia, vol.  1,  p.  98,  1841. 

The  golden  mouse  occurs  nearly  throughout  the  lowlands  of  the 
State,  being  confined  here,  as  elsewhere,  almost  exclusively  to  the 
Lower  Austral  Zone.  Specimens  examined  from  Huntsville,  Ardell, 
Seale,  Dothan,  Brewton,  Mobile,  and  Point  Clear  agree  well  with 
specimens  of  this  race  from  Florida. 

This  mouse  lives  chiefly  in  canebrakes  and  swampy  woodland, 
more  rarely  in  dry,  thickety  flatwoods  or  among  the  hills.  It  is  some- 
what arboreal  in  habits,  and  builds  for  itself  a  globular  or  oval- 
shaped  nest  in  a  bunch  of  cane,  a  bush,  or  a  tree.  The  nest  is  beau- 
tifully woven  of  grass  or  shreds  of  bark  and  lined  with  fur  or  fine 
grass  stalks.  Audubon  and  Bachman  state  that  they  have  observed 
this  mouse  run  up  tall  trees  with  great  agility  and  conceal  itself  in 
a  hole  at  least  30  feet  from  the  ground.  In  a  swamp  at  Dothan, 
Alabama,  the  writer  caught  one  as  it  was  running  up  the  trunk  of 
a  tree.  Dr.  A.  K.  Fisher  found  a  nest  in  the  Dismal  Swamp,  Vir- 
ginia, in  dead  brush,  about  2  feet  from  the  ground;  when  the  nest 
was  touched  a  female  golden  mouse  with  two  young  clinging  to  her 
teats  dropped  to  the  ground,  and  after  a  few  moments  ran  up  the 
stem  of  a  bush  and  reentered  the  nest. 


1921.]  MAMMALS  OF  ALABAMA — MICE.  51 

Oryzomys  palustris  palustris  (Harlan). 

Swamp  Rice  Rat. 

Mus  palustris  Harlan,  Silliman's  Amer.  Journ.  Sci.,  vol.  31,  p.  386,  1837. 

The  rice  rat,  or  marsh  rat,  apparently  is  generally  distributed  in 
the  State,  living  in  wet,  marshy  areas  in  fields,  as  well  as  in  wooded 
swamps  and  in  the  coastal  salt  marshes.  It  is  particularly  numerous 
in  the  latter  situations,  as  on  Blakely  Island,  in  Mobile  River,  and 
about  Little  Lagoon,  on  the  coast  of  Baldwin  County.  Fishermen 
who  ply  their  trade  at  night  on  the  Lagoon  state  that  these  rats  are 
common  there  and  that  at  night  they  often  come  around  the  tents 
on  the  shore  of  the  lagoon  to  feed  on  scraps  of  fish  and  other  food 
thrown  out  by  the  men. 

Although  living  to  some  extent  in  cultivated  lands,  this  rat  prob- 
ably is  not  sufficiently  numerous  to  do  any  appreciable  damage  to 
crops.  It  is  similar  in  appearance  to  the  common  barn  rat,  but 
slightly  smaller,  with  a  slenderer  tail  and  whiter  belly.  Specimens 
have  been  taken  at  Huntsville,  Sand  Mountain  (near  Carpenter), 
Reform.  V/eogufka,  Elmore,  Gallion,  Autaugaville,  Seale,  Mobile, 
Baj'ou  Labatre,  and  Little  Lagoon,  Baldwin  County.  Audubon  and 
Bachman  describe  the  habits  of  this  species  in  South  Carolina  as 
follows : 

The  Rice  Meadow-Mouse,  as  its  name  implies,  is  found  in  particular  locali- 
ties in  the  banks  of  the  rice-fields  of  Carolina  and  Georgia.  It  burrows  in 
the  dykes  or  dams  a  few  inches  above  the  line  of  the  usual  rise  of  the  water. 
Its  burrow  is  seldom  much  beyond  a  foot  in  depth.  It  has  a  compact  nest  at 
the  extremity,  where  it  produces  its  young  in  April.  They  are  usually  four 
or  five.  *  *  *  We  have  observed  it  scratching  up  the  rice  when  newly 
planted  and  before  it  had  been  overflowed  by  the  water.  When  the  rice  is  in 
its  milky  state  this  animal  commences  feeding  on  it,  and  continues  during 
the  autumn  and  winter,  gleaning  the  fields  of  the  scattered  grains.  We  have 
also  seen  its  burrows  in  old  banks  on  deserted  rice-fields,  and  observed  that 
it  had  been  feeding  on  the  large  seeds  of  the  Gama  grass  {Tripsicum  dacty- 
loides),  and  on  those  of  the  wild  rye  (Elymus  Viryinicus) .  *  *  *  Its  nest 
is  suspended  on  a  bunch  of  interlaced  marsh  grass.  In  this  situation  we 
observed  one  with  five  young.  At  certain  seasons  this  little  animal  feeds  on 
the  seeds  of  the  marsh  grass  (Spartina  glabra).  When  these  fall  it  sometimes 
retires  to  the  shore  for  food,  but  has  no  disrelish  to  the  small  Crustacea  and 
mollusks  that  remain  on  the  mud  at  the  subsiding  of  the  tide. 

This  species  swims  rapidly,  and  dives  in  the  manner  of  the  European  water 
rat  (Arvicola  amphibia),  or  of  our  Arvicola  Pennsylvanica.  In  an  attempt 
at  capturing  some  alive,  they  swam  so  actively,  and  dived  so  far  from  us,  that 
the  majority  escaped.  Those  we  kept  in  captivity  produced  young  in  May  and 
September;  they  were  fed  on  grains  of  various  kinds,  but  always  gave  the 
preference  to  small  pieces  of  meat.48 


<6  Audubon  and  Bachman,  Op.  cit.,  vol.  3,  p.  215. 


52  NORTH  AMERICAN   FAUNA.  [No.  45 

Sigmodon  hispidus  hispidus  Say  and  Ord. 

Cotton  Rat. 

Sigmodon  hispidus  Say  and  Ord,  Journ.  Acad.  Nat.  Sci.  Philadelphia,  vol.  4, 
pt.  2,  p.  354,  1825. 

The  cotton  rat  is  perhaps  the  most  abundant  rodent  on  the 
farm  lands  of  Alabama.  It  lives  in  grassy  fields,  brushy  pastures, 
marshes,  and  along  the  brushy  or  weed-grown  borders  of  cultivated 
fields,  practically  in  all  sections  except  on  the  mountains.  It  makes 
well-defined  beaten  trails  through  the  grass  and  under  fallen  vegeta- 
tion and  brush  piles,  and  apparently  lives  both  in  shallow  burrows 
and  in  grass  nests  constructed  above  ground.  Its  food  consists 
mainly  of  the  stems  and  seeds  of  various  wild  grasses  or  other  plants, 
and  it  doubtless  consumes  some  grain  also,  but  in  this  section  of 
the  country  is  not  accused  of  damaging  crops  to  any  extent.  It  is 
partly  diurnal  and  may  often  be  seen  in  broad  daylight  scurrying 
across  a  road  or  other  open  space.  The  only  vernacular  name  I  have 
heard  applied  to  it  is  "  gopher  rat."  The  species  has  been  found  in 
abundance  at  Leighton,  Woodville,  Arclell,  Dean,  Barachias,  Jack- 
son, Carlton,  Bon  Secour,  Ashford,  and  Bayou  Labatre;  specimens 
have  been  taken  also  on  Sand  Mountain  (near  Carpenter),  at  Pied- 
mont, Cane  Creek  (Marshall  County),  Greensboro,  Gallion,  Cath- 
erine, Auburn,  Seale,  Castleberiw,  and  Abbeville. 

Neotoma  floridana  floridana  (Ord). 
Florida  Wood  Rat. 
Mus  floridana  Ord,  Bull.  Soc.  Philom.  Paris,  p.  181,  1818. 

The  Florida  wood  rat  occurs  abundantly  over  the  greater  part  of 
Alabama  from  the  Tennessee  Valley  southward.  It  dwells  in  a 
variety  of  situations,  but  is  most  frequently  found  in  wooded  bot- 
toms or  swamps,  where  it  constructs  medium-sized  nests  of  sticks, 
leaves,  and  rubbish,  placed  on  or  near  the  ground  in  hollow  logs  or 
trees.  In  the  cultivated  lands  of  the  Prairie  Belt  and  the  Tennessee 
Valley  the  rats  live  chiefly  in  the  osage  orange  hedges,  where  they 
build  very  large  nests  in  the  branches  of  the  trees,  often  10  or  15 
feet  above  the  ground.  When  driven  from  their  nests  they  run 
swiftly  and  deftly  about  among  the  branches.  In  many  places  it  is 
a  common  pastime  for  boys  to  poke  the  rats  out  of  their  nests  and 
hunt  them  with  dogs.  Wherever  rocky  bluffs  or  cliffs  occur  within 
the  range  of  this  species,  the  rats  may  be  found  dwelling  in  them, 
seeming  to  prefer  (as  do  most  members  of  the  genus)  rocks  to  trees 
or  logs.  When  occupying  such  situations  they  do  not  build  large 
nests,  but  are  content  to  carry  a  small  quantity  of  sticks  and  rubbish 
into  crevices  in  the  rocks  or  into  caves.  On  Sand  Mountain,  in 
niches  in  the  cliffs  in  Long  Island  Creek  Gulch,  I  found  small  nests  or 
"  forms  "  made  of  grass  and  small  sticks,  similar  to  those  made  by 


1921.] 


MAMMALS  OF  ALABAMA — BATS. 


53 


Neotoma  pennsylvanica.  These,  I  believe,  were  constructed  by  the 
rats  and  probably  are  used  at  night  as  resting  places,  as  were  those  of 
pennsylvanica  at  Mammoth  Cave,  Kentucky,  in  the  daytime. 

Judging  from  the  accumulation  of  hickor3T-nut  shells  found  about 
the  cliffs  occupied  by  these  wood  rats,  nuts  must  form  an  important 
item  of  their  food  in  winter.  Papaw  seeds  have  also  been  found 
near  their  dens.  In 
summer  the  rats 
doubtless  feed 
largely  on  succulent 
herbs  and  berries. 

The  range  of  this 
species  in  Alabama, 
so  far  as  worked  out, 
seems  to  be  strictly 
complementary  t  o 
that  of  pennsyl- 
vanica, the  latter 
being  found  only 
north  of  the  Ten- 
nessee River,  and 
floridana  only  south 
of  it.  At  Muscle 
Shoals  pennsyl- 
vanica lives  in  the 
cliffs  on  the  north 
shore  and  floridana 
in  the  cliffs  directly 
opposite.  The  pres- 
ent species  has  been 
taken  also  at  Leigh- 
ton,  Sand  Mountain 
(Long  Island  Creek, 
near  Carpenter), 
Choccolocco  Moun- 
tain (2,000  feet  alti- 
tude), Dean   (Cedar 

Mountain,  1,500-2,000  feet),  Lock  14  on  Warrior  River,  Autaugaville, 
Barachias,  Castleberry,  Mobile,  Point  Clear,  and  Orange  Beach 
(%  4). 

Neotoma  pennsylvanica  Stone. 

Allegheny  Cliff  Rat. 
Neotoma  pennsylvanica  Stone,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  p.  16,  1893. 

The  Allegheny  cliff  rat,  a  large  gray  species,  ranges  from  southern 
New  York  and  Pennsylvania  southward  to  northern  Alabama,  where 


Fig.  4. — Distribution  of  wood   rats   in  Alabama. 

1.  Neotoma  pennsylvanica. 

2.  Neotoma  floridana  floridana. 


54  NORTH  AMERICAN  FAUNA.  [No.  45 

apparently  it  is  limited  by  the  Tennessee  River.  It  has  been  taken 
at  Woodville,  Monte  Sano  (near  Huntsville),  Gunters  Mountain 
(near  Fort  Deposit),  and  at  Muscle  Shoals  (north  shore)   (fig.  4). 

It  occurs  in  scattering  colonies,  chiefly  about  cliffs  on  the  mountain 
sides,  but  sometimes  descends  to  low  altitudes  along  streams,  where 
it  dwells  in  rock  bluffs  or  in  caves.  It  is  never  found,  however,  in 
lowland  swamps,  as  is  its  relative,  the  Florida  wood  rat.  This 
species  carries  into  its  dens  in  the  crevices  of  the  cliffs  considerable 
quantities  of  sticks,  leaves,  nut  shells,  and  other  rubbish,  but  does  not 
construct  large  stick  nests  such  ^as  the  rats  of  the  floridanus  group  use. 
At  Mammoth  Cave,  Kentucky,  the  writer  found  these  wood  rats 
living  in  small  nests  or  "  forms  "  made  of  finely  shredded  cedar  bark, 
placed  on  ledges  or  on  the  floor  of  a  small  cave.40 

The  food  of  this  species  is  mainly  of  vegetable  origin  and  consists 
in  part  of  hickory  nuts,  acorns,  chestnuts,  and  the  tender  leaves  and 
stems  of  herbaceous  plants.  The  cliff  rats,  dwelling  as  they  do  in 
the  remoter  parts  of  the  mountains,  rarely  cause  any  damage  to  man's 
industries.  Sometimes,  however,  where  the  cliffs  in  which  they  live 
are  close  to  farm  buildings,  the  rats  may  enter  corn  cribs  and  destroy 
some  of  the  corn. 

Pitymys  pinetorum  pinetorum  (LeConte). 

Pine  Mouse. 

Psammomys  pinetorum  LeConte,  Ann.  Lye.  Nat.  Hist.  New  York,  vol.  3,  p.  133, 
1830. 

Pine  mice  occur  in  moderate  numbers  over  the  greater  part  of  the 
State,  at  least  as  far  south  as  Houston  County.  Specimens  from 
Autaugaville,  Prattville,  Greensboro,  and  Ashford  are  provisionally 
referred  to  the  typical  race,  which  occupies  the  South  Atlantic 
coastal  plain  of  the  Carolinas  and  Georgia,  but  additional  material 
from  southern  Alabama  may  render  necessary  a  different  decision  as 
to  the  status  of  this  form.  In  color,  and  size  of  ears,  the  specimens 
agree  well  with  pinetorum,  and  the  skulls,  although  decidedly  smaller, 
have  the  V-shaped  interpterygoid  notch  characteristic  of  that  race. 

These  mice  are  moderately  common  in  sandy  fields  near  Autauga- 
ville, and  one  was  taken  also  in  a  grassy  bog  in  the  heavy  timber 
on  the  borders  of  Bear  Swamp.  The  name  "  pine  mouse  "  is  some- 
what a  misnomer,  for,  although  the  animal  occasionally  lives  in  open 
pine  lands,  it  is  more  often  found  in  cultivated  fields,  meadows,  or  even 
in  wet  bottom-land  timber.  It  is  largely  subterranean  in  habit, 
and  for  that  reason  is  not  very  well  known  to  most  people.  It  is  a 
sleek,  silky  little  mouse  of  a  rich  tawny-brown  color,  with  a  blunt 
nose,  small,  beady  eyes,  and  a  very  short  tail.    It  makes  little  bur- 

«  See  Howell,  Proc.  Biol.  Soc.  Washington,  vol.  23,  pp.  27-28,  1910. 


1921.]  MAMMALS   OF   ALABAMA MUSKRATS.  55 

rows  and  runways  in  the  grass  of  meadows  or  under  the  dead  leaves 
and  rubbish  in  the  woods,  and  also  makes  extensive  use  of  the  run- 
ways of  the  common  mole.  This  habit  has  resulted  in  fastening  on 
the  innocent  mole  the  responsibility  for  the  damage  done  by  the 
pine  mouse  to  vegetables,  bulbs,  etc. 

This  species  is  probably  the  most  destructive  of  any  of  the  native 
field  rodents  of  the  Eastern  States.  It  attacks  white  potatoes,  sweet 
potatoes,  bulbs,  strawberry,  blackberry,  and  other  plants,  nursery 
stock,  orchard  trees,  and  stores  of  vegetables  of  all  kinds.  Freshly 
planted  seeds  of  melons  or  cantaloupes  are  extensively  eaten,  as 
well  as  the  roots  of  the  growing  plants.  The  damage  to  orchards, 
which  consists  in  gnawing  off  the  bark  near  the  surface  of  the  ground, 
is  often  extensive,  and  in  the  apple-growing  regions  of  Virginia  and 
West  Virginia,  where  the  mice  are  abundant,  presents  a  serious 
problem. 

Pitymys  pinetorum  auricularis  (Bailey). 

Bluegrass  Vole  ;  Pine  Mouse. 

Microtus  pinetorum  auricularis  Bailey,  Proc.  Biol.   Soc.  Washington,  vol.  12, 
p.  90,  1898. 

The  bluegrass  vole,  an  interior  race  of  the  pine  mouse,  occupies 
the  lower  Mississippi  Valley,  ranging  northward  through  northern 
Alabama,  Tennessee,  and  Kentucky.  Typical  specimens  have  rather 
larger  ears  than  the  other  subspecies.  Specimens  examined  from 
Ardell,  Woodville,  and  Sand  Mountain  (near  Carpenter)  are  re- 
ferred to  this  raCe  because  of  the  close  resemblance  in  skull  char- 
acters, the  interpterygoid  fossa  being  U-shaped  rather  than  V-shaped. 
In  habits  this  race  probably  does  not  differ  much  from  the  other  sub- 
species. On  Sand  Mountain  their  signs  were  rather  numerous  in 
cultivated  land,  where  as  usual  these  rodents  travel  mainly  in  mole 
runways.  At  Ardell  and  Woodville,  L.  J.  Goldman  found  them  in 
heavy  timber  and  trapped  specimens  under  rotten  logs. 

Fiber  zibethicus  zibethicus  (Linnaeus). 

Muskeat. 

[Castor]  zibethicus  Linnaeus,  Syst.  Nat.,  ed.  12,  vol.  1,  p.  79,  1766. 

The  common  muskrat  of  the  Northern  States  occurs  in  moderate 
numbers  in  most  of  the  streams  and  ponds  in  the  northern  part  of 
Alabama,  south  as  far  at  least  as  Clarke,  Lowndes,  and  Pike  Coun- 
ties. In  the  southern  part  of  the  State  no  muskrats  are  found,  ex- 
cept in  southern  Mobile  County,  where  Fiber  rivalicius  occurs  (fig. 
5).  It  is  difficult  to  understand  why  the  present  species  does  not 
range  all  the  way  to  the  coast,  as  conditions  in  the  southern  counties 
seem  as  well  suited  to  its  needs  as  in  the  more  northern  parts. 


56  NORTH   AMERICAN    FAUNA.  [No.  45 

Specimens  have  been  examined  from  Reform  and  Autaugaville, 
and  the  animal  is  reported  from  Leighton,  Elkmont,  Scottsboro, 
Guntersville,  Erin  (Clay  County),  Ardell,  Squaw  Shoals  (Warrior 
River),  Greensboro,  Mount  Weogufka,  Barachias,  Seale,  Hayne- 
ville,  Teasley  Mill  (Montgomery  County),  Myrtlewood,  and  Coffee- 
ville.  Inquiries  made  of  many  residents  and  trappers  indicate  that 
the  species  occurs  at  Jackson  only  sporadically.  Most  of  the  people 
questioned  had  never  seen  a  muskrat,  but  two  persons  had  seen  a 
few  taken  there. 

C.  W.  Howe,  a  trapper  of  long  experience,  states  that  muskrats 
first  appeared  near  Linwood,  ^ike  County,  in  1906,  and  are  now 
quite  plentiful  on  the  upper  Conecuh  River  between  Troy  and  Union 
Springs,  where  in  1898  none  were  to  be  found.  He  states  also  that 
in  1912  he  found  unmistakable  signs  of  muskrats  in  Little  River,  on 
the  northern  line  of  Baldwin  County.  The  animals  are  very  scarce 
so  far  south,  and  Mr.  Howe  thinks  this  colony  probably  drifted 
down  from  a  point  higher  up  the  river.  In  the  spring  of  1916  he 
saw  a  single  individual  still  farther  south,  in  Mobile  River  at 
Twelve  Mile  Island,  where  by  the  light  of  a  jack  lantern  it  was 
plainly  seen  swimming  in  the  river  at  night.  As  no  other  signs  of 
muskrats  have  ever  been  found  in  that  vicinity,  this  individual  doubt- 
less was  a  stray.  Muskrats  are  chiefly  of  nocturnal  habit  and  are 
rarely  seen  abroad  in  daylight  except  in  the  dusk  of  evening,  when 
they  emerge  from  their  dens  and  begin  to  feed. 

The  muskrat's  method  of  constructing  its  house  varies  in  differ- 
ent parts  of  its  range.  In  a  region  of  ponds  and  marshes  it  con- 
structs large  domed  houses  of  rushes,  sticks,  leaves,  mud,  and  other 
rubbish.  In  creeks  and  rivers  where  the  banks  are  steep,  however, 
it  tunnels  into  the  banks  from  beneath  the  surface  of  the  water  and 
constructs  its  nest  in  a  cavity  excavated  at  the  upper  end  of  the  tun- 
nel, above  high-water  mark.  This  appears  to  be  the  usual  habit  of 
the  species  in  Alabama. 

The  food  of  the  muskrat  consists  largely  of  the  stems,  leaves,  and 
roots  of  various  aquatic  plants,  including  rushes,  sedges,  arrowhead 
(Sagittaria) ,  pondlilies,  golden  club  {Orontium) ,  and  various  marsh 
grasses.  It  is  said  to  feed  on  timothy,  redtop  (Agrostis),  and 
clover,  as  well  as  on  Indian  corn  and  various  garden  vegetables. 
Joel  Burgess,  of  Dean,  Clay  County,  states  that  he  has  known  the 
muskrats  to  cut  100  or  more  stalks  of  corn  in  three  nights  and  drag 
them  into  the  creek.  The  animal  varies  its  fare  in  some  localities 
by  feeding  on  mussels;  and  piles  of  shells  are  often  found  about 
shallow  places  in  the  rivers,  where  they  have  been  left  by  muskrats. 


1921.] 


MAMMALS   OF   ALABAMA MUSKRATS. 


57 


Fiber  rivalicius  Bangs. 
Louisiana  Mitskrat. 

;s,  Proc.  Boston  Soc   Nat.  Hist.,  vol.  26,  p.  541, 


Fiber  zibethicus  rivalicius  Bam 
1895. 

The  small,  dark-colored  Louisiana  muskrat  is  found  at  present  in 
Alabama  only  in  the  coast  region  west  of  Mobile  Bay  (fig.  5).  The 
animal  has  apparently  extended  its  range  into  the  State  within  very 
recent  times,  since 
it  is  unknown  to 
most  hunters  and 
trappers  in  the  re- 
gion where  it  is  now 
found,  and  many  of 
those  who  are  ac- 
quainted with  it 
state  that  it  first  ap- 
peared there  shortly 
after  a  big  storm  in 
1906. 

This  species  is  not 
as  yet  widely  dis- 
persed, as  it  is  known 
at  present  only  from 
the  coast  marshes 
about  Bayou  La- 
batre.  Signs  of  its 
presence  were  seen 
also  in  Three  Mile 
Creek,  near  Mobile, 
in  1913,  by  the  late 
Kenneth  Beale,  but 
no  specimens  have 
been  taken  there. 
It  will  undoubtedly 
spread  out  and  be- 
come more  abundant 
along  the  coast  of 
Mississippi  Sound  and  the  western  shore  of  Mobile  Bay,  and  there 
seems  to  be  no  reason  why  it  should  not  extend  its  range  into  the 
big  marshes  of  the  Delta  region  and  along  the  eastern  shore  of  the  Bay. 

At  Bayou  Labatre,  residents  reported  a  number  of  muskrat  houses 
on  the  marshes  between  the  mouth  of  the  bayou  and  Little  River, 
but  a  storm  and  high  tide  in  the  fall  of  1915  apparently  broke  up  this 
colony,  and  in  December  of  that  year  search  for  them  in  that  locality 
was  unsuccessful.    A  considerable  colony,  however,  was  located  close 


Fig. 


— Distribution  of  muskrats  in  Alabama. 

1.  Fiber  zibethicus  zibethicus. 

2.  Fiber  rivalicius. 


58  NORTH   AMERICAN   FAUNA.  [No.  45 

to  the  town  in  a  little  marsh  between  the  public  road  and  the  pine 
woods.  The  animals  had  constructed  four  large  houses  of  rushes, 
the  largest  fully  6  feet  in  diameter  and  3  or  4  feet  high.  For  quite 
a  space  around  each  house  all  the  growing  rushes  had  been  cut  close 
to  the  ground,  and  the  marsh  was  honeycombed  with  underground 
trails  leading  in  various  directions  from  the  houses  to  the  banks  of 
the  bayou,  most  of  the  exits  being  under  water  except  at  very  low  tide. 
This  species  is  a  valuable  fur  bearer,  and  as  it  increases  in  numbers 
doubtless  will  provide  a  profitable  industry  for  the  residents  of  the 
coast  counties.  In  the  rice-growing  districts  of  Louisiana  it  is  a 
serious  pest  by  reason  of  its  habit  of  burrowing  into  dikes  and  levees. 

Rattus  norvegicus  (Erxleben). 
Norway  Rat  ;  Barn  Rat  ;  "  Gopher  Rat." 

[Hits]  norvegicus  Erxleben,  Syst.  Itegni  Anim.,  vol.  1,  p.  381,  1777. 

The  common  Norway  rat — an  exotic  species,  imported  from  the 
Old  World — is  widely  distributed  in  the  State,  and  is  everywhere  a 
serious  pest.  It  lives  chiefly  in  towns  and  around  farm  buildings, 
but  often  ranges  out  into  the  fields  for  a  considerable  distance,  par- 
ticularly along  ditch  banks  and  about  the  borders  of  marshy  bottoms. 
In  the  open  it  digs  burrows  in  the  banks  of  ditches  or  streams,  and 
becomes  practically  a  "  wild  "  animal.  It  does  an  immense  amount 
of  damage  to  stores  of  feed  or  grain,  to  corn  in  cribs  or  in  the  field, 
to  rice  and  sugar  cane,  and  to  young  poultry.  The  losses  resulting 
from  its  depredations  are  much  greater  than  those  occasioned  by  all 
the  native  species  of  rodents  combined. 

Rats  are  responsible  also  for  the  dissemination  of  bubonic  plague, 
trichinosis,  and  other  serious  diseases.  Every  effort,  therefore, 
should  be  made  to  reduce  their  numbers  and  to  exclude  them  from 
storehouses  and  farm  buildings.47 

Rattus  rattus  rattus  (Linnaeus). 

Black  Rat. 

[Mus]  rattus  Linnaeus,  Syst.  Nat.,  ed.  10,  vol.  1,  p.  61,  1758. 

The  black  rat  was  introduced  into  North  America  soon  after  the 
settlement  of  the  early  colonies,  and  before  long  was  established 
in  the  towns  and  villages.  After  the  arrival  of  the  brown  rat,  how- 
ever, it  began  to  decrease  in  numbers  and  at  present  is  rare  and 
local  in  the  eastern  United  States.  Specimens  were  taken  near 
Greensboro  in  1891  and  on  Lookout  Mountain,  near  Fort  Payne,  in 

47  Suggestions  for  destroying  rats  are  contained  in  Farmers'  Bulletin  896,  TJ.  S.  Dept. 
Agr.,  "  House  Rats  and  Mice,"  by  David  E.  Lantz ;  pp.  24,  1917.  Cf.  also  Separate  725, 
Yearbook  1917,  U.  S.  Dept.  Agr.,  pp.  235-251,  1918,  "  The  House  Rat :  the  Most  Destruc- 
tive Animal  in  the  World,"  by  the  same  author. 


1921.]  MAMMALS  OF   ALABAMA — POCKET   GOPHERS.  59 

1911.     It  is  slenderer  and  longer  tailed  than  the  brown  rat,  and  is 
said  to  live  almost  exclusively  in  the  walls  of  houses. 

Rattus  rattus  alexandrinus  ( Geoff roy ) . 

Roof  Rat. 

Mus  alexandrinus  Geoffroy,  Descr.  de  l'Egypte,  Mamm.,  p.  733,  1818. 

The  roof  rat  is  believed  to  be  a  native  of  Egypt.  It  resembles  the 
black  rat  closely  in  size  and  proportions,  but  differs  in  color,  being 
of  about  the  shade  of  the  brown  rat,  but  with  whiter  or  more  yellow- 
ish underparts.  It  is  well  established  in  the  Southern  States,  and 
has  been  taken  in  Alabama  at  Greensboro,  Castleberry,  Bayou  La- 
batre,  and  Abbeville.  Roof  rats  live  chiefly  about  the  roofs  of 
dwellings  or  outhouses,  but  occasionally  wander  out  into  the  fields. 
At  Abbeville  the  writer  observed  them  on  a  moonlight  evening  in 
June  climbing  silently  and  dexterously  up  and  down  the  branches  of 
an  oleander  bush  growing  close  to  the  hotel  porch. 

Mus  musculus  musculus  Linnaeus. 

House  Mouse. 

[Mus]  musculus  Linnaeus,  Syst.  Nat.,  ed.  10,  vol.  1,  p.  62,  1758. 

The  house  mouse,  a  species  introduced  from  Europe,  is  abundant 
and  generally  distributed  throughout  the  State.  It  is  by  no  means 
confined  to  the  vicinity  of  buildings  but  ranges  widely  in  waste  lands 
and  cultivated  fields.  So  far  as  known,  however,  it  does  not  seri- 
ously damage  farm  crops  except  as  they  are  stored  in  granaries  or 
shocks.  In  houses  and  farm  buildings  it  is  a  serious  pest,  though 
to  a  less  extent  than  its  larger  relative,  the  brown  rat. 

Family  GEOMYIDAE:  Pocket  Gophers. 

Geomys  tuza  mobilensis  Merriam. 

Alabama  Pocket  Gopher  ;  "  Salamander." 

Geomys  tuza  mobilensis  Merriam,  North  Amer.  Fauna  No.  8,  p.  119,  1895. 

The  pocket  gopher  (usually,  though  incorrectly,  known  in  the 
South  as  "salamander  ")  is  found  rather  locally  in  southern  Alabama, 
east  of  the  Tombigbee  River,  and  ranges  north  to  Tuscaloosa  County 
(fig.  6).  The  Tombigbee- Warrior  Basin  marks  the  western  limit 
of  its  range,  and  no  member  of  the  genus  is  again  encountered  until 
after  crossing  the  Mississippi  River. 

The  usual  habitat  of  the  pocket  gopher  is  in  sandy  pine  flats,  but 
in  Tuscaloosa  County,  near  Lock  14,  Warrior  River  (the  most  north- 
ern point  from  which  the  species  is  known),  it  is  found  on  gravelly 
ridges  in  mixed  timber — longleaf  pine,  oaks,  etc.    Its  presence  is 


60 


NORTH   AMERICAN   FAUNA. 


[No.  45 


easily  detected  by  the  numerous  mounds  of  earth  which  it  throws  up 
at  frequent  intervals,  usually  in  a  more  or  less  continuous  line.  Only 
rarely  ma}'  the  pocket  gopher  be  seen  above  ground,  and  then  only  for 
a  few  seconds  as  it  pushes  the  earth  from  its  burrow.     By  digging 

open  the  mound, 
however,  until  a 
clear  tunnel  is  ex- 
posed, and  setting  a 
steel  trap  or  pocket 
gopher  trap  in  the 
runway,  the  animal 
may  easily  be  cap- 
tured. 

Unlike  its  western 
relative,  the  Ala- 
bama pocket  gopher 
is  not  injurious  to 
cultivated  crops.  It 
lives  almost  entirely 
in  waste  lands,  in 
poor,  sandy,  or  grav- 
elly soil,  and  is  said 
to  desert  a  locality 
as  soon  as  cultivation 
is  begun. 

Specimens  have 
been  examined  from 
Orange  Beach,  Point 
Clear,  B  r  e  w  t  o  n, 
Steadham,  Seale, 
and  Lock  14,  War- 
rior River.  Evi- 
dences of  the  ani- 
mal's presence  have 
been  seen  at  Castleberry,  Evergreen,  Andalusia,  Ewell,  Thomasville, 
Magnolia,  Selma,  Kingston,  and  Chehaw. 

Family  SCIURIDAE :  Marmots,  Squirrels,  etc. 

Marmota  monax  monax  (Linnaeus). 

Southern  Woodchtjck  ;   Ground-hog. 

[Mus]  monax  Linnaeus,  Syst.  Nat.,  ed.  10,  vol.  1,  p.  60,  1758. 

The  woodchuck,  or  ground-hog,  as  it  is  usually  called  in  the  South, 
occurs  plentifully  in  the  rough,  hilly  country  of  northern  Alabama 
as  far  south  as  Tuscaloosa  and  Coosa  Counties  and  the  Talladega 


Fig.  6. 


-Distribution  of  the  Alabama  pocket  gopher  {Geafmya 
tuza  mobilensis)  in  Alabama. 


North  American  Fauna  No.  45,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  X. 


Fig.  I.— Mounds  of  Pocket  Gopher  (Geomys  tuza  mobilensis)  in  Open 

Pine  Forest. 

Gravelly  hills  near  Lock  14,  Warrior  River. 


<s>.       ^k 

s^tiM 

;. 

■'A 

| 

*■■  "'""   tlliiiE*.' 

Fig.  2.— Mounds  of  Pocket  Gopher  in  Sandy  Field. 

Waste  land  near  Point  Clear. 


1921.] 


MAMMALS   OF  ALABAMA — MARMOTS. 


61 


Mountains  of  Talladega  and  Clay  Counties  (fig.  7).  Stragglers  are 
sometimes  found  somewhat  farther  south.  L.  S.  Golsan  states  that  a 
single  individual — the  only  one  ever  known  in  the  vicinity — was  killed 
near  Autaugaville  a  number  of  years  ago  (about  1890) ;  and  Peter 
Brannon  states  that  he  has  seen  the  woodchuck  a  few  times  at  Seale. 
In  the  South  woodchucks  live  chiefly  in  rocky  bluffs  along  streams 
or  on  mountain  sides,  their  burrows  usually  being  constructed  under 
bowlders    or    lederes 


of  rock  or  beneath 
roots  of  trees — al- 
most always  on  a 
steep,  wooded  slope. 
From  these  retreats 
they  pay  visits  to 
near-by  fields  and  do 
considerable  damage 
to  such  farm  crops 
as  oats,  hay,  corn, 
and  pumpkins,  and 
to  beans  and  other 
garden  vegetables. 

The  woodchuck  is 
a  famous  sleeper, 
spending  usually 
from  four  to  six 
months  during  the 
winter  season  in  its 
burrow  in  a  state  of 
torpor.  In  the 
Northern  States  it 
retires  in  fall  often 
as  early  as  the  last  of 
September,  remain- 
ing until  the  middle 
or  last  of  the  fol- 
lowing March ;  in 
the  South  it  remains 
out  later,  perhaps  till  November 
ruary. 

Specimens  of  the  woodchuck  have  been  examined  from  Ardell, 
Cullman  County,  and  the  species  is  reported  also  from  Leighton, 
Monte  Sano,  Sand  Mountain,  Woodville,  Guntersville,  Natural 
Bridge,  Piedmont,  Attalla,  Jasper,  Squaw  Shoals,  Mount  Weogufka, 
and  the  Talladega  Mountains,  near  Erin. 


nje  Beach 


Fig.  7.- 


Distribution  of  the  southern  woodchuck  (Alarmota 
monax  monax)  in  Alabama. 


and   reappears   early    in   Feb- 


62  NORTH   AMERICAN   FAUNA.  [No.  45 

Tamias  striatus  venustus  Bangs. 

.  South westebn  Chipmunk. 

Tamias  striatus  venustus  Bangs,  Proc.  Biol.  Soc.  Washington,  vol.  10,  p.  137, 
1896. 

The  chipmunk,  or  ground  squirrel,  occurs  abundantly  in  certain  of 
the  northern  counties  of  the  State  and  sparingly  in  the  central  part 
as  far  south  as  Carlton,  Castleberry,  and  Dothan.  It  is  apparently 
shyer  and  less  noisy  than  the  chipmunk  of  New  England,  but  this 
may  be  due  in  part  to  its  relative  scarcity. 

In  most  localities  in  the  State  where  the  species  is  reported  to 
occur,  it  has  been  possible  usually  to  see  only  an  occasional  individual 
and  that  for  only  a  brief  moment.  The  usual  haunts  of  the  chip- 
munk are  the  heavily  forested  hillsides  and  rocky  mountain  slopes 
where  there  is  an  abundance  of  brush  and  fallen  timber,  over  and 
beneath  which  it  loves  to  romp  and  in  the  shelter  of  which  it  finds 
safe  retreats.  In  the  low  country  chipmunks  are  found  in  the  moist 
bottomland  woods  and  even  in  timbered  swamps.  They  live  chiefly 
on  or  near  the  ground  and  make  burrows  in  the  side  of  a  hill  or 
bank,  beneath  a  rock  or  the  roots  of  a  tree,  or  in  a  decaying  stump. 
Audubon  and  Bacliman  describe  a  burrow  which  was  excavated  in 
January  (locality  not  stated,  but  probably  New  York)  when  the 
snow  lay  about  5  inches  deep  on  a  piece  of  ground  covered  with 
leaves  to  a  depth  of  8  inches : 

The  hole  descended  at  first  almost  perpendicularly  for  about  three  feet.  It 
then  continued  with  one  or  two  windings,  rising  a  little  nearer  the  surface 
until  it  had  advanced  about  8  feet,  when  we  came  to  a  large  nest  made  of  oak 
leaves  and  dried  grasses.  Here  lay,  snugly  covered,  three  Chipping  Squirrels. 
Another  was  subsequently  dug  from  one  of  the  small  lateral  galleries,  to  which 
it  had  evidently  retreated  to  avoid  us.  They  were  not  dormant,  and  seemed 
ready  to  bite  when  taken  in  the  hand ;  but  they  were  not  very  active,  and  ap- 
peared somewhat  sluggish  and  benumbed,  which  we  conjectured  was  owing  to 
their  being  exposed  to  sudden  cold  from  our  having  opened  their  burrow. 

There  was  about  a  gill  of  wheat  and  buckwheat  in  the  nest;  but  in  the  gal- 
leries we  afterwards  dug  out,  we  obtained  about  a  quart  of  the  beaked  hazel 
nuts  (Corylus  rostrata),  nearly  a  peck  of  acorns,  some  grains  of  Indian  corn, 
about  two  quarts  of  buckwheat,  and  a  very  small  quantity  of  grass  seeds.48 

From  this  account  it  is  evident  that  the  chipmunk  does  not  hiber- 
nate in  the  true  sense,  but  stores  up  food  for  winter  use.  The  same 
authors  state  that  in  Louisiana  they  caught  a  chipmunk  which  had 
in  its  cheek  pouches  no  less  than  16  chinquapin  nuts  and  that  another 
received  from  Pennsylvania  contained  "  at  least  one  and  a  half  table- 
spoonfuls  of  bush  trefoil  {Hedysamm  cannabinum)  in  its  widely  dis- 
tended sacks."49  Merriam  states  that  the  chipmunk  feeds  upon  a 
variety  of  nuts  and  roots,  corn  and  other  grain,  and  the  larvae  of  cer- 

43  Audubon  and  Bacbman,  Op.  cit.,  vol.  1,  pp.  69-70. 
»  Op.  cit,  p.  69. 


1921.]  MAMMALS    OF   ALABAMA SQUIRRELS.  63 

tain  insects ;  and  that  in  the  Adirondack  region  of  New  York  beech- 
nuts form  its  principal  food.50 

Chipmunks  with  varying  quantities  of  nuts  and  grain  in  their 
pouches  have  been  observed  by  Mr.  Sewell,  of  Dean;  one  had  90 
grains  of  small  corn ;  and  others,  2  chestnuts  and  1  or  2  chinquapins ; 
5  chinquapins ;  2  hickory  nuts ;  and  5  peanuts  in  the  shell.  In  Bucks 
Pocket,  on  Sand  Mountain,  chipmunks  were  feeding  on  acorns,  one 
individual  taken  having  five  large  pieces  of  acorn  in  its  pouches. 
At  this  locality  these  animals  were  more  numerous  than  the  writer 
ever  saw  them  elsewhere  in  the  South.  In  the  timber  at  the  foot  of 
the  steep  slopes  of  the  canyon  they  were  constantly  seen  and  heard. 
Their  call  notes,  which  were  heard  throughout  the  day,  were  of  two 
forms — one  a  low  chuck  or  cluck,  the  other  a  high-pitched  tchip, 
much  like  the  alarm  note  of  the  hooded  warbler.  When  badly  fright- 
ened the  chipmunks  uttered  a  hurried  chippering  note,  twitched  their 
tails  nervously,  and  scurried  into  their  retreats. 

The  species  is  reported  to  occur  commonly  in  the  Talladega  Moun- 
tains, at  Guntersville,  and  at  Elkmont ;  in  smaller  numbers  at  Leigh- 
ton,  Huntsville,  Squaw  Shoals,  Mount  Weogufka,  and  Greensboro; 
and  sparingly  at  Jackson,  Carlton,  Castleberry,  Booth,  Wilsonville, 
Seale,  Teasley  Mill,  and  Dothan. 

The  Alabama  series  as  a  whole  is  intermediate  between  Tamias 
striatus  striatus  and  T.  s.  venustus,  but  rather  nearer  the  latter.  The 
skulls  are  decidedly  larger  than  typical  striatus  from  the  Carolinas 
(Roan  Mountain  specimens)  and  agree  with  those  of  venustus  from 
Arkansas.  In  color,  some  individuals  agree  with  venustus  in  having 
lighter  gray  backs  and  brighter  rumps  than  striatus,  while  others 
are  almost  as  dark  as  typical  striatus.  The  grayest  specimens  are 
from  Ardell,  Guntersville,  Bucks  Pocket,  and  Talladega  Mountains. 
Those  from  Woodville  and  Greensboro  approach  striatus  in  color. 
The  series  differs  also  from  striatus  in  having  a  larger  hind  foot 
(average  36.6  mm.  instead  of  33.6),  this  being  even  larger  than 
typical  venustus  (which  measures  35.5). 

Sciurus  carolinensis  carolinensis  Gmelin. 
Giiay  Squirrel;  "Cat  Squirrel." 
[Sciurus]  carolinensis  Gmelin,  Syst.  Nat.,  vol.  1,  p.  148,  178S. 

The  gray  squirrel  is  one  of  the  most  abundant  and  generally  dis- 
tributed mammals  in  the  State.  It  inhabits  for  the  most  part  moist 
bottom  lands  and  swamps,  where  there  is  an  abundance  of  oak,  hick- 
ory, and  other  nut-bearing  trees.  It  is  found  also  in  lesser  numbers 
on  the  wooded  hills  and  the  lower  slopes  of  the  mountains,  but  does 
not  range  at  all  into  pine  timber. 

50  Merriam,  C.  Hart,  Mamm.  Adirondack  region :  Trans.  Linnaean  Soc.  New  York,  vol. 
2,  pp.  135-136,  1884. 


64  NOETH   AMERICAN   FAUNA.  [No.  45 

Gray  squirrels  live  both  in  hollow  trees  and  in  nests  among  the 
branches,  constructed  of  leaves,  Spanish  moss,  etc.  The  young,  usu- 
ally five  or  six  in  number,  are  brought  forth  early  in  spring,  and 
sometimes  a  second  litter  is  born  in  the  summer.  This  species  is  said 
to  be  irregularly  migratory,  its  wanderings  probably  induced  by  a 
search  for  desirable  feeding  places.  Acorns  and  hickory  nuts  form 
its  principal  food,  but  it  is  fond  also  of  the  fruit  of  the  great  mag- 
nolia {Magnolia  foetida).  It  is  most  active  during  the  early  morn- 
ing hours  and  about  sundown ;  and  Audubon  states  that  it  is  to  some 
extent  nocturnal,  having  frequently  been  observed  by  moonlight. 

The  flesh  of  the  gray  squirrel  is  tender  and  juicy  and  is  highly 
esteemed  for  food.  Thousands  are  shot  by  local  hunters,  and  in  many 
localities  the  species  has  been  greatly  reduced  in  numbers.  As  its 
tameness  renders  it  usually  an  easy  mark  to  one  who  understands  its 
habits,  if  it  is  not  afforded  better  legal  protection  it  will  before  many 
years  become  very  scare. 

Typical  specimens  have  been  examined  from  Sand  Mountain  (near 
Carpenter),  Lookout  Mountain  (near  Fort  Payne),  Arclell,  Greens- 
boro, Barachias,  Castleberry,  Point  Clear,  Bon  Secour,  and  Orange 
Beach.  Four  specimens  from  the  Tensaw  River  swamps  above  Hur- 
ricane and  two  from  Stiggins  Lake  are  intermediate  between  caro- 
linensis and  fuliginosus.  The  species  is  recorded  also  from  Cherokee, 
Muscle  Shoals,  Elkmont,  Guntersville,  Talladega  Mountains,  Pied- 
mont, Mount  Weogufka,  Squaw  Shoals,  Natural  Bridge,  Auburn, 
Autaugaville,  Reform,  Oakchia,  Seale,  Teasley  Mill,  Jackson,  Carl- 
ton, Abbeville,  and  Ashford. 

Sciurus  carolinensis  fuliginosus  Bachman. 

Bayou  Gray  Squirrel. 

Sciurus  fuliginosus  Bachman,  Proc.  Zool.  Soc.  London,  p.  97,  1838. 

The  bayou  gray  squirrel  ranges  from  the  coast  of  Louisiana  to 
the  head  of  Mobile  Bay,  being  confined  to  the  shores  of  the  bayous 
and  the  deep  cypress  swamps.  At  Stiggins  Lake  (3  miles  east  of 
Mount  Vernon) ,  these  squirrels  were  abundant  on  May  26,  1914,  and 
of  three  specimens  obtained,  one  is  typical  of  fuliginosus,  the  others 
intermediate  between  it  and  carolinensis.  A  typical  specimen  was 
taken  also  at  Chuckvee  Bay,  May  12,  1911,  and  one  at  Bayou  Labatre, 
January  6,  1917. 

Sciurus  nig-er  niger  Linnaeus. 

Carolina  Fox  Squirrel. 

[Sciurus]  niger  Linnaeus,  Syst.  Nat.,  ed.  10,  vol.  1,  p.  64,  1758. 

The  Carolina  fox  squirrel  is  the  form  ranging  through  South 
Carolina,  southern  Georgia,  and  northern  Florida ;  it  enters  Ala- 
bama only  in   the   southeastern   counties,   ranging  west   to   Castle- 


1921.]  MAMMALS   OF   ALABAMA SQUIRRELS.  65 

berry,  where  it  intergrades  with  the  interior  form,  texianus  (fig.  8). 
Six  specimens  from  Abbeville  are  typical  niger,  with  grayish  backs 
and  whitish  underparts ;  one  is  solid  light  gray  above  except  the  head, 
which  is  black,  this  one  agreeing  perfectly  with  specimens  from  South 
Carolina,  the  type  region.  Of  11  specimens  from  Castleberry,  5  are 
typical  ?iiger,  2  typical  texianus,  and  4  intermediate,  perhaps  nearer 
texianus.  This  squirrel  is  reported  to  occur  in  some  numbers  at 
Headland,  and  rarely  at  Seale. 

The  fox  squirrel  is  a  lover  of  the  dry  pine  forests  and  is  never 
found  in  the  low  bottoms  inhabited  by  the  gray  squirrel.  It  is  a  very 
shy  species  and  is  seldom  seen  moving  about  except  by  hunters  who 
remain  concealed  near  its  haunts.  It  is  easily  "treed,"  however,  by 
the  aid  of  a  good  squirrel  dog,  and  in  most  places,  through  persistent 
hunting,  has  become  very  scarce.  When  surprised  on  the  ground 
(so  Bachman  tells  us),  these  squirrels  will  usually  run  for  a  consider- 
able distance  to  reach  a  tree  containing  a  hollow,  in  which  they  take 
refuge.  Their  usual  nesting  place  is  in  a  hollow  tree,  but  outside 
nests,  composed  of  sticks,  leaves,  and  moss  placed  in  the  forks  of  a 
tree,  are  used  for  temporary  homes  during  a  part  of  the  year. 

This  squirrel  is  wholly  diurnal  and  is  said  to  remain  in  its  retreats 
until  the  sun  is  well  up.  Its  alarm  note  is  rather  harsh  barking,  not 
unlike  that  of  the  gray  squirrel,  but  usually  recognizable.  Audubon 
and  Bachman  have  given  a  good  account  of  its  feeding  habits,  as 
follows : 

The  food  of  the  Fox  Squirrel  is  various ;  besides  acorns,  and  different  kinds  of 
nuts,  its  principal  subsistence  for  many  weeks  in  autumn  is  the  fruit  extracted 
from  the  cones  of  the  pine,  especially  the  long-leaved  pitch  pine  (Pinus 
palustris).  Whilst  the  green  corn  is  yet  in  its  milky  state,  this  Squirrel  makes 
long  journeys  to  visit  the  fields,  and  for  the  sake  of  convenience  frequently 
builds  a  temporary  summer  house  in  the  vicinity,  in  order  to  share  with  the  little 
Carolina  squirrel  and  the  crow  a  portion  of  the  delicacies  and  treasures  of  the 
husbandman ;  *  *  *  The  Fox  Squirrel  does  not  appear  to  lay  up  any  winter 
stores — there  appears  to  be  no  food  in  any  of  his  nests,  nor  does  he,  like  the 
red  squirrel  (Sciurus  hudsonius),  resort  to  any  hoards  which  in  the  season  of 
abundance  were  buried  in  the  earth,  or  concealed  under  logs  and  leaves.  During 
the  winter  season  he  leaves  his  retreat  but  seldom,  and  then  only  for  a  little 
while  and  in  fine  weather  in  the  middle  of  the  day.  He  has  evidently  the 
power,  like  the  marmot  and  raccoon,  of  being  sustained  for  a  considerable  length 
of  time  without  much  suffering  in  the  absence  of  food.  When  this  animal  makes 
his  appearance  in  winter,  he  is  seen  searching  among  the  leaves  where  the  wild 
turkey  has  been  busy  at  work,  and  gleaning  the  refuse  acorns  which  have 
escaped  its  search ;  at  such  times,  also,  this  squirrel  does  not  reject  worms  and 
insects  which  he  may  detect  beneath  the  bark  of  fallen  or  decayed  trees. 
Toward  spring  he  feeds  on  the  buds  of  hickory,  oak,  and  various  other  trees,  as 
well  as  on  several  kinds  of  roots,  especially  the  wild  potato  (Apios  tuberosa). 
As  the  spring  advances  farther  he  is  a  constant  visitor  to  the  black  mulberry 
tree  {Morus  rubra),  where  he  finds  a  supply  for  several  weeks.  From  this  time 
till  winter,  the  fruits  of  the  field  and  forest  enable  him  to  revel  in  abundance.61 


51  Audubon  and  Bachman,  Op.  clt.,  vol.  2,  p.   136. 
49663°— 21 5 


66 


NORTH   AMERICAN   FAUNA. 


[No.  45 


Sciurus  niger  texianus  Bachman. 

Bachman  Fox  Squirrel. 

Sciurus  texianus  Bachman,  Proc.  Zool.  Soc.  London,  p,  86,  1838. 

The  Bachman  fox  squirrel  ranges  from  southern  Louisiana  through 
Mississippi  and  the  greater  part  of  Alabama  (except  the  south- 
eastern counties)  to  eastern  Tennessee  (fig.  8).    It  is  a  richly  colored 

form,  somewhat  re- 
sembling the  com- 
mon fox  squirrel  of 
the  northern  Missis- 
sippi Valley  (ruft- 
venter) ,  but  differ- 
ing in  having  the 
head  black  and  the 
nose  and  ears  white. 
Six  specimens 
from  Autaugaville 
and  two  from  Sand 
Mountain  (near  Car- 
penter) are  typical 
texianus,  except  that 
one  from  the  latter 
place  has  buffy  ears ; 
the  underparts  vary 
from  light  ochrace- 
ous-buff  to  deep  och- 
raceous-orange ;  one 
from  Autaugaville  is 
nearly  black.  One 
specimen  from  Jack- 
son is  very  dark,  the 
underparts,  legs,  tail, 
and  ears  being  rich 
tawny,  and  the  back 
overlaid  with  the 
same  color;  another 
from  the  same  place, 
however,  is  of  the 
normal  buff  color,  with  buff-tipped  ears.  Two  specimens  from  Orange 
Beach  and  two  from  Castleberry  also  have  the  upperparts  rather 
heavily  tinged  with  tawny.  Four  specimens  from  Castleberry  are 
clearly  intermediate  between  texianus  and  niger,  showing  more  or 
less  gray  on  the  sides.  Others  from  this  locality,  as  already  stated, 
are  typical  niger. 


nje  Besch 


Fig.  8. 


-Distribution  of  fox  squirrels  in  Alabama. 

1.  Sciurus  niger  niger, 

2.  Sciurus  niger  texianus. 

(Unshaded  area  indicates  region  from  which  no  specimens 
have  been  examined.) 


1921.]  MAMMALS   OF   ALABAMA BEAVERS.  67 

This  race  has  practically  the  same  habits  as  the  Carolina  fox 
squirrel ;  in  the  mountainous  regions  of  the  State,  however,  it  is  not 
confined  to  pine  timber,  but  ranges  as  well  into  the  mixed  woods  of 
pine,  hickory,  and  oak.  Like  the  other  forms  of  the  species,  it  is 
never  found  in  wet  bottoms,  but  always  on  dry  uplands  or  mountain 
slopes.  It  is  reported  to  occur  plentifully  at  or  near  Cherokee,  Scotts- 
boro,  Piedmont,  Talladega  Mountains,  and  Mount  Weogufka,  and 
rarely  at  Huntsville,  Woodville,  Fort  Payne,  Ardell,  Reform,  Myrtle- 
wood,  Carlton,  and  Bayou  Labatre. 

Family  PETAURISTIDAE :  Flying  Squirrels. 

Glaucomys  volans  saturatus  Howell. 

SOUTHEASTEKN   FLYING   SQUIRREL. 

Glaucomys  volans  saturatus  Howell,  Proc.  Biol.  Soc.  Washington,  vol.  28,  p.  110, 
1915. 

Flying  squirrels  occur  commonly  in  all  parts  of  the  State,  but  on 
account  of  their  nocturnal  habits  are  seldom  seen  unless  driven  from 
their  retreats.  They  inhabit  chiefly  hollow  trees  or  stumps,  but  fre- 
quently take  up  their  abode  in  an  attic  or  outbuilding.  If  one  wishes 
to  observe  their  interesting  and  graceful  movements,  he  may  easily 
dislodge  them  from  their  retreats  in  hollow  trees  by  rapping  sharply 
on  the  trunk  with  an  axe  or  club.  Launching  from  a  point  well  up 
in  a  tree  the  squirrels  glide  swiftly  to  a  lower  point  on  a  near-by  tree, 
then  scramble  up  among  the  branches  and  are  ready  for  another 
flight. 

Flying  squirrels  feed  chiefly  upon  various  nuts,  berries,  and  seeds ; 
they  are  fond  of  meat  and  occasionally  destroy  insects.  They  are 
looked  upon  with  disfavor  by  many  people  by  reason  of  their  depre- 
dations on  pecan  groves  or  on  stores  of  nuts  in  the  lofts  of  houses; 
but  the  damage  done  by  them  is  usually  inconsiderable. 

Specimens  of  this  dark  southern  race  have  been  examined  from 
Sand  Mountain  (near  Carpenter),  Ardell,  Greensboro,  York,  Au- 
taugaville,  Carlton,  Dothan,  Mobile,  and  Perdido  River,  and  there 
are  records  of  its  occurrence  at  many  other  localities. 

Family  CASTORIDAE:  Beavers. 

Castor  canadensis  carolinensis  Rhoads. 

Cabolina  Beaver. 

Castor  canadensis  carolinensis  Rhoads,  Trans.  Amer.  Philos.  Soc,  N.  S.,  vol.  19, 
p.  420,  1894. 

The  beaver  in  early  times  doubtless  occupied  practically  the  entire 
State ;  with  the  coming  of  the  settlers  the  animals  decreased  greatly 
in  numbers,  but  in  some  localities  have  held  their  own  remarkably 


05  NORTH   AMERICAN   FAUNA.  [No.  45 

well,  even  in  well-settled  farming  districts.  Apparently  they  are 
more  numerous  at  present  in  the  central  part  of  the  State,  in  Mont- 
gomery and  Lowndes  Counties,  than  in  either  the  wild  hill  country 
of  the  northern  part  or  the  big  swamps  of  the  south. 

Old  residents  in  Montgomery  County,  near  Teasley  Mill,  stated 
that  in  the  early  days  of  the  settlement  of  the  country  beavers  were 
found  in  abundance,  often  building  extensive  dams  and  forming  large 
ponds  in  the  swampy  bottoms.  Although  comparatively  few  were 
trapped,  they  are  said  to  have  disappeared  entirely  from  this  region 
for  a  number  of  years.  About  1908  they  appeared  again,  working  up 
from  the  lower  stretches  of  Catoma  Creek,  and  at  present  they  seem 
to  be  confined  to  this  creek  where  it  runs  through  Catoma  Swamp. 

In  December,  1913,  E.  G.  Holt  visited  this  locality  and  secured  one 
young  beaver,  and  in  April  and  May,  1914,  the  writer  spent  several 
days  there  and  trapped  an  adult  female.  At  the  point  where  the 
adult  specimen  was  trapped  were  several  "  slides  "  on  the  steep  bank 
of  the  creek  and  two  well-worn  trails  leading  through  a  dense 
canebrake  to  a  shallow  slough.  The  animals  were  living  in  the 
creek  where  it  runs  through  the  timber  and  beside  cultivated 
fields.  Here  the  stream  is  not  over  10  to  20  feet  wide  and  for  the 
most  part  shallow,  though  in  places  perhaps  8  or  10  feet  deep. 
Beavers  had  constructed  small  dams  across  the  creek,  generally  utiliz- 
ing the  trunks  of  fallen  trees  (not  of  their  own  cutting)  for  a  founda- 
tion, filling  in  the  space  beneath  the  logs  with  sticks,  brush,  cane 
stalks,  leaves,  and  mud.  They  had  constructed  no  houses,  but  were 
living  in  burrows  in  the  banks  of  the  creek.  Occasional  small  trees 
had  been  felled  by  them,  the  largest,  a  cottonwood  27  inches  in  cir- 
cumference ;  the  end  of  this  log,  which  rested  in  the  middle  of  the 
stream,  showed  marks  of  the  beavers'  teeth  where  they  had  gnawed 
the  bark ;  a  few  other  trees  growing  along  the  creek  showed  marks  of 
their  gnawings — a  pine  {Pinus  glabra),  a  water  beech  (Carpfoms 
caroliniana) ,  and  a  sweet  gum  {Liquidamber  styracifiua) . 

In  April,  1915,  L.  J.  Goldman  visited  the  region  about  Hayneville, 
Lowndes  County,  and  noted  considerable  evidence  of  the  occurrence 
of  beavers  in  numbers.  Several  dams  were  found  at  intervals  along 
Big  Swamp  Creek  and  in  Jones  Lake.  At  the  latter  place  three  houses 
had  been  constructed  of  sticks,  cane  stalks,  and  mud,  but  they  were 
not  in  use  at  the  time,  although  they  had  been  occupied  the  previous 
summer.  Wherever  cornfields  bordered  the  lake  Mr.  Goldman  found 
deeply  worn  trails  leading  into  the  fields,  and  he  was  informed  that 
in  summer  beavers  resort  regularly  to  the  corn  patches  and  consume 
a  considerable  quantity  of  corn. 

Beavers  are  occasionally  surprised  in  the  fields  even  in  day- 
light, and  a  large  one  was  reported  to  have  been  taken  in  that  manner 
on  Pintlala  Creek  in  the  spring  of  1915.    At  Teasley  Mill  a  planter 


North  American  Fauna  No.  45,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  XI. 


Fig.  I.— Dense  Cane  Brake  Bordering  Catoma  Creek. 


Fig.  2.— Beaver  Dam  in  Catoma  Creek. 

Views  in  Catoma  Swamp  near  Teasley  Mill,  Montgomery  County. 


1921.]  MAMMALS  OF  ALABAMA — BEAVERS.  69 

told  of  finding  a  beaA^er  in  a  cornfield,  some  100  yards  from  the 
creek,  standing  up  reaching  for  an  ear  of  corn.  On  moonlight  nights 
it  is  often  possible  by  remaining  concealed  near  the  creek  to  observe 
beavers  as  they  swim  by  in  the  stream.  During  periods  of  low  water, 
by  cutting  the  dams  and  lowering  the  stream  so  that  the  entrances  to 
the  burrows  are  exposed,  they  can  be  driven  from  their  dens  with  the 
aid  of  dogs  and  easily  secured.  At  present  there  seems  to  be  very  little 
trapping  done  for  their  fur,  but  the  flesh  is  eagerly  sought  for  food, 
especially  by  negroes  on  the  plantations.  Beavers  have  been  protected 
by  many  of  the  planters,  which  doubtless  is  the  reason  they  have  not 
been  exterminated.  They  are  now  protected,  by  law  between  March  1 
and  November  1  each  year. 

B.  I.  Garner,  a  trapper  working  in  Autauga  County,  caught  two 
beavers  on  the  Alabama  River,  12  miles  below  Prattville,  in  Novem- 
ber, 1916.  A  colony  was  reported  to  be  living  on  Beaver  Creek, 
Conecuh  County,  a  few  miles  from  Evergreen,  in  1919,  where  they  are 
said  to  occupy  a  body  of  water  covering  many  acres  of  ground.52 
Beavers  were  common  until  a  few  years  ago  in  Big  Uchee  Creek, 
Russell  County.  Hundreds  are  said  to  have  been  taken  in  a  pond  in  a 
dense  canebrake  about  12  miles  northwest  of  Seale,  the  last  having 
been  trapped  about  1912.  A  small  colony  was  still  located  on  the 
Moreland  plantation,  about  8  miles  northeast  of  Seale,  in  1915.  In 
1912  a  few  were  reported  living  on  Horse  Creek,  near  Thomasville, 
and  on  Beaver  Creek,  near  Myrtlewood.  In  1911,  a  small  colony  was 
reported  near  Mount  Weogufka,  and  in  1915  L.  J.  Goldman  found 
signs  of  a  small  colony  on  the  lower  part  of  Rock  Creek,  Winston 
County,  where  during  the  year  previous  3  or  4  had  been  caught. 

In  the  northern  part  of  the  State  beavers  seem  almost  to  have 
been  exterminated;  they  formerly  occurred  in  small  numbers  in  the 
Tennessee  River  at  Muscle  Shoals,  but  disappeared  about  1895 ;  they 
lived  about  the  "towheads"  (small  islands)  in  the  river  and  bur- 
rowed into  the  banks,  but  did  not  build  dams.  A  few  were  reported 
in  1916  in  Big  Wills  Creek,  near  Collinsville.  They  disappeared 
from  Talladega  Creek,  near  Dean,  about  1896. 

In  the  southern  counties  rather  vague  reports  of  the  beaver  have 
been  gathered,  indicating  that  a  few  may  still  remain  near  Jackson, 
Abbeville,  and  Bayou  Labatre.  A  professional  trapper  reports  tak- 
ing three  in  Bassett  Creek,  near  Jackson,  in  February,  1913,53  and 
C.  W.  Howe,  also  a  trapper  of  long  experience,  relates  that  in  1915 
he  noticed  a  few  signs  of  beavers  in  Little  River,  on  the  northern  line 
of  Baldwin  County. 

Besides  the  adult  female  and  young  male  from  Montgomery 
County,  there  has  been  obtained  through  C.  H.  Harbison  an  adult 

62  Montgomery    (Ala.)    Advertiser,   October  24,   1919. 

53Kuhn,  A.  L..  Hunter-Trader-Trapper,  vol.  28,  pp.  41-42,  June,  1914. 


70  NORTH   AMERICAN    FAUNA.  [No.  45 

male  skull  taken  at  Ardell,  Cullman  County.  Comparison  with 
specimens  of  canadensis  from  interior  Canada  (Oxford  House) 
shows  that  the  present  form  has  a  shorter  and  relatively  broader 
skull ;  heavier,  broader,  and  more  widely  expanded  zygomata ;  wider 
braincase  and  interorbital  region;  and  larger  audital  bullae.  The 
lower  jaw  is  relatively  heavier,  the  inferior  border  of  the  angle  flat- 
tened into  a  broad  ledge,  and  the  coronoid  process  heavy. 

The  adult  female  specimen  taken  in  Catoma  Creek  weighed  38^ 
pounds ;  one  taken  at  Jackson,  by  A.  L.  Kuhn,  weighed  54  pounds ; 5* 
and  one  killed  on  Pintlala  Creek  was  reported  to  weigh  65  pounds, 
which  probably  represents  aborit  the  maximum  size  attained  by  this 
species.  External  measurements  of  the  adult  from  Catoma  Creek  are 
as  follows :  Total  length,  1,035  mm. ;  length  of  tail,  290  mm. ;  greatest 
breadth  of  tail,  163  mm. ;  hind  foot,  170  mm. 

Order  LAGOMORPHA. 

Family  LEPORIDAE:  Hares  and  Rabbits. 

Sylvilagus  floridanus  mallurus  (Thomas). 

Eastern  Cottontail. 

L[epus]  n[uttalli]  mallurus  Thomas,  Ann.  and  Mag.  Nat.  Hist.,  ser.  7,  vol.  2, 
p.  320,  1898. 

The  cottontail  rabbit  occurs  abundantly  in  all  parts  of  Alabama ;  it 
lives  chiefly  in  old  fields,  brier  patches,  and  the  brushy  borders  of 
woodland,  and  to  some  extent  in  swampy  bottoms.  Cottontails  are 
chiefly  nocturnal  in  habit,  and  usually  spend  most  of  the  daylight 
hours  concealed  in  a  "  form  "  in  a  tussock  of  grass  or  a  bunch  of  weeds 
or  briers,  or  beneath  a  small  bush.  When  startled  from  their  retreats, 
they  run  with  great  swiftness,  twisting  and  doubling  through  the 
brush,  and  seek  shelter  in  a  near-by  thicket  or  a  hollow  log  or  stump. 
They  make  occasional  inroads  upon  the  farmer's  kitchen  garden 
and  often  seriously  damage  orchards  and  nursery  stock  by  gnawing 
the  bark,  coming  out  to  feed  in  the  early  morning  and  again  about 
sundown.  Nelson,  in  his  revision  of  the  rabbits  of  North  America,55 
referred  most  of  the  Alabama  cottontails  to  the  subspecies  alacer,  the 
type  locality  of  which  is  Stillwell,  Oklahoma;  more  abundant  mate- 
rial, now  available,  however,  indicates  that  they  should  be  placed  with 
mallurus,  the  form  ranging  throughout  the  South  Atlantic  States. 
They  agree  very  closely  with  this  race  in  color  and  differ  only  in 
having  slightly  smaller  audital  bullae. 

Specimens  have  been  examined  from  Sand  Mountain  (near  Car- 
penter), Huntsville,  Talladega  Mountains  (Erin),  Ardell,  Auburn, 

54  Kuhn,  Arthur  L.,  Op.  cit. 

«  Nelson,  E.  W„  North  Amer.  Fauna  No.  29,  pp.  174-176,  1909. 


1921.]  MAMMALS   OF  ALABAMA RABBITS.  71 

Carlton,  Castleberry,  Mobile,  Alabama  Port,  Bon  Secour,  Point 
Clear,  Orange  Beach,  and  Dolhan.  The  species  is  reported,  also, 
from  Leighton,  Elkmont,  Natural  Bridge,  Attalla,  Greensboro,  An- 
niston,  Barachias,  Seale,  Abbeville,  Bayou  Labatre,  and  many  other 
places. 

Sylvilagus  transitionalis  (Bangs). 

New  England  Cottontail. 

Lepus  sylvaticus  transitionalis  Bangs,  Proc.  Boston  Soc.  Nat.  Hist.,  vol.  26,  p. 
405,  1895. 

The  New  England  cottontail  ranges  from  New  York  and  New 
Hampshire  southward  to  northern  Georgia  and  Alabama,  inhabiting 
chiefly  mountain  slopes  and  the  rough  foothill  country.  By  reason  of 
its  close  external  resemblance  to  the  common  eastern  cottontail  {Syl- 
vilagus  ftoridanus  mallurus)  it  is  difficult  to  distinguish  between  the 
two  except  by  critical  examination  of  specimens,  and  as  a  result  little 
is  known  of  the  peculiar  habits  or  exact  distribution  of  the  New  Eng- 
land cottontail.  It  resembles  the  common  rabbit  rather  closely,  but 
differs  in  smaller  size,  shorter  ears,  a  greater  amount  of  black  on  the 
back,  and  in  peculiar  skull  characters.  Its  range  overlaps  that  of 
mallurus  with  no  indications  of  intergradation.  It  is  said  to  be  more 
of  a  forest-inhabiting  species  than  is  the  common  rabbit. 

At  present,  transitionalis  is  known  from  only  one  locality  in  Geor- 
gia (Brasstown  Bald  Mountain,  where  it  was  taken  by  the  writer  in 
1908)  and  three  localities  in  Alabama — Erin  and  Dean,  at  the  foot 
of  the  Talladega  Mountains,  in  Clay  County,  and  Ardell,  in  the 
rough  hill  country  along  Sipsey  Fork,  Cullman  County — from  each 
of  which  a  single  specimen  has  been  obtained  (fig.  9).  More  exten- 
sive collecting  in  the  northern  part  of  the  State  will  doubtless  result 
in  a  considerable  extension  of  its  known  range. 

Sylvilagus  aquaticus  aquaticus  (Bachman). 

Swamp  Rabbit  ;  "  Cane-cutter." 

Lepus  aquaticus  Bachman,  Journ.  Acad.  Nat.  Sci.  Philadelphia,  vol.  7,  p.  319, 
1837. 

The  large  swamp  rabbit  is  abundant  and  generally  distributed  over 
the  lowlands  of  the  State  except  in  the  extreme  southern  counties  east 
of  Mobile  Bay,  which  region  is  occupied  by  the  eastern  marsh  rabbit 
(Sylvilagus  palustris)  (fig.  10).  It  is  found  in  all  the  river  swamps 
and  ranges  up  along  the  small  streams  to  the  very  foot  of  the  moun- 
tains, where  it  reaches  an  altitude  of  1,200  feet  in  Clay  County.  In 
the  Tennessee  Valley  it  is  reported  as  far  north  as  Crowtown  Island 
and  Big  Crow  Creek,  near  Stevenson. 

Although  living  by  choice  in  the  wet  bottoms,  these  rabbits  when 
pursued  will  sometimes  leave  the  swamp  and  cross  the  dry,  wooded 


72 


NORTH  AMERICAN   FAUNA. 


[No.  45 


hillsides.  They  usually  remain  hidden  most  of  the  day  in  thick 
brush,  but  occasionally  one  may  be  seen  sitting  in  an  opening  in  the 
swamp,  and  if  the  observer  remains  quiet  it  will  often  wait  a  few 
seconds  before  running  off,  or  rarely  it  will  stop  after  running  a  few 
steps  and  look  around.  When  pursued  by  dogs  these  rabbits  make 
very  fast  time  and  usually  have  no  difficulty  in  distancing  the  pack. 
When  hard  pressed,  however,  they  are  quite  likely  to  take  refuge  in 

a  hollow  tree,  up 
which  they  scramble 
for  a  distance  of  sev- 
eral feet.  In  such 
situations  hunters 
often  secure  them  by 
inserting  in  the  hol- 
low a  slender  switch, 
which  being  twisted 
into  the  fur  and 
gently  pulled,  usu- 
ally induces  the  rab- 
bit to  come  down  far 
enough  to  be  seized 
by  hand  or  by  a  dog. 
When  running  from 
hounds  they  very 
frequently  jump  into 
a  creek  and  swim 
across  or  some  dis- 
tance up  or  down  the 
stream.  During  pe- 
riods of  high  water 
these  rabbits  are 
often  compelled  to 
seek  safety  on  the 
high  ridges  or  small 
islands  in  the  bot- 
toms, and  at  such 
times  large  numbers 
are  sometimes  killed  by  hunters  with  guns  or  clubs. 

Bachman  states  that  this  rabbit  feeds  on  the  roots  of  various 
aquatic  plants,  especially  on  a  species  of  iris  growing  in  water.  The 
common  name  in  use  for  this  animal  in  many  parts  of  the  South — 
"  cane-cutter " — indicates  its  fondness  for  the  stems  of  the  cane 
{Arundinaria) ,  which  grows  in  abundance  in  its  favorite  haunts. 
Where  cultivated  fields  adjoin  the  swamps,  swamp  rabbits  often  for- 


ran $e  B«ach 


Fig.  9.- 


-Distribution  of  the  New  England  cottontail  rabbit 
(Sylvilagus  transitionalis)    in   Alabama. 


1921.]  MAMMALS   OF   ALABAMA RABBITS.  73 

age  in  corn  or  other  crops  and  at  times  causes  considerable  damage. 
Bachman  states  that  the  young,  which  are  from  four  to  six  in  num- 
ber, "  are  frequently  found  in  nests  formed  of  leaves  and  grasses, 
placed  on  hillocks  in  the  swamps,  or  in  the  hollow  of  some  fallen 
tree." 56 

Bachman  described  this  species  in  1837  from  specimens  sent  to  him 
by  Dr.  J.  M.  Lee  and  Capt.  Benjamin  Logan  from  western  Alabama. 
Typical  specimens  have  been  examined  from  Ardell,  Dean,  Hunts- 
ville,  Reform,  Greensboro,  Auburn,  Teasley  Mill,  Castleberry,  Carl- 
ton, and  Abbeville.  The  animal  is  recorded  also  from  Elkmont, 
Leighton,  Fort  Payne,  Piedmont  (Nance  Creek),  Natural  Bridge, 
Squaw  Shoals,  Weogufka,  York,  Autaugaville,  Jackson,  and  Seale. 
Specimens  from  Stockton  and  Whistler  are  approaching  the  sub- 
species littoralis.  Its  southern  limit  is  apparently  between  Castle- 
berry and  Flomaton  and  between  Abbeville  and  Dothan.  As  yet,  it 
never  has  been  taken  at  the  same  locality  with  palustris. 

Sylvilagus  aquaticus  littoralis  Nelson. 
Coast  Swamp  Rabbit. 
Syfttilagus  aquaticus  littoralis  Nelson,  North  Amer.  Fauna  No.  29,  p.  273,  1909. 
The  coast  race  of  the  swamp  rabbit  is  confined  to  a  narrow  belt 
along  the  coast  from  Matagorda  Bay,  Texas,  eastward  to  the  west 
side  of  Mobile  Bay.  It  ranges  but  a  short  distance  from  the  coast 
marshes  (specimens  from  so  near  as  Whistler  being  referable  to  the 
interior  form,  aquaticus)  and  is  practically  limited  to  the  tidewater 
region.  In  the  marshes  of  Blakely  Island,  opposite  Mobile,  it  is 
very  numerous  and  three  specimens  taken  there  are  typical  of  the 
race.  Specimens  from  the  Tensaw  River,  four  miles  north  of  Hurri- 
cane, however,  are  intermediate  between  littoralis  and  aquaticus,  but 
are  best  referred  to  the  former.  The  species  occurs  only  sparingly 
about  Bayou  Labatre  and  the  coast  region  westward;  it  is  occasion- 
ally taken  on  Grand  Batture  Island.  In  habits  this  rabbit  differs 
very  little  from  the  typical  race,  but  those  observed  in  the  Blakely 
Island  marshes  seemed  less  fleet  of  foot  and  very  averse  to  leaving 
the  cover  of  the  marsh.  In  fact,  about  the  only  way  to  get  them 
out  of  the  rushes  is  to  set  fire  to  the  vegetation ;  when  driven  out  by 
the  flames,  they  skulk  along  with  ears  laid  back  and  dart  into  the 
nearest  patch  of  cover. 

Sylvilagus  palustris  palustris  (Bachman). 

Marsh  Rabbit. 

Lepus  palustris  Bachman,  Journ.  Acad.  Nat.  Sci.  Philadelphia,  vol.  7,  p.  194, 
1837. 

The  marsh  rabbit  occurs  in  Alabama  only  in  the  southern  counties 
east  of  Mobile  Bay ;  its  northern  limit  has  not  been  definitely  ascer- 

56  Audubon  and  Bachman,  Op.  cit.,  vol.  1,  p.  289. 


74 


NORTH   AMERICAN   FAUNA. 


[No.  45 


tained,  but  is  roughly  indicated  by  records  of  its  occurrence  at  Gor- 
don, Ashford,  Dothan,  Orange  Beach,  and  Flomaton  (fig.  10). 

In  the  salt  marshes  of  Perdido  Bay  and  Bon  Secour  River  the 
rabbits  are  abundant,  and  there  their  droppings  and  well-beaten  trails 
may  be  seen  everywhere.  In  localities  back  from  the  coast  they  live 
in  the  small,  wet,  timbered  swamps  which  are  found  at  frequent 

intervals  along  the 
streams  and  around 
ponds  in  the  timber. 
They  never  leave  the 
cover  of  the  swamps 
unless  driven  out  by 
dogs  or  fire,  and  on 
such  occasions  they 
quickly  seek  the 
nearest  thicket  or 
take  refuge  in  a 
hollow  log  or  tree. 
Their  short  legs  pre- 
vent them  from  at- 
taining much  speed ; 
when  pursued  they 
depend  for  escape  on 
their  facility  in  dou- 
bling and  twisting. 
When  driven  from  a 
marsh  by  the  firing 
of  the  dry  grass, 
often  many  are 
killed  by  men  wait- 
ing for  them  with 
clubs.  In  the  marsh 
they   are  frequently 


Fig.    10. — Distribution 


o£    swamp 
Alabama. 


and    marsh    rabbits    in 


1.  Sylvilagus   aquaticus  aquatic-us. 

2.  Sylvilagus  aquaticus   littoralis. 

3.  Sylvilagus   palustris   palustris. 


caught  by  dogs. 
They  take  to  the 
water  readily  and 
swim  easily  and 
swiftly. 

Bachman  states  that  the  marsh  rabbit  feeds  on  various  grasses  and 
gnaws  the  twigs  of  young  sassafras  and  of  the  pond  spice  {Benzoin) . 
He  has  also  found  it  digging  for  the  bulbs  of  the  wild  potato  {Apios 
apios)  and  the  atamasco  lily.57  Nelson  states  that  the  marsh  rabbits 
examined  from  Bon  Secour  are  intermediate  between  palustrus  and 
paludicola,  but  nearer  the  former.58 

CT  Audubon  and  Bachman,  Op.  cit,  vol.  1,  p.  155. 

68  Nelson,  E.  W.,  North  Amer.  Fauna  No.  29,  p.  268,  1909. 


1921.]  MAMMALS   OF   ALABAMA DEER.  75 

Order  ARTIODACTYLA. 

Family  CERVIDAE :  Deer. 

Odocoileus  virginianus  virginianus  (Boddaert). 

Virginia  Deek. 

Cervus  virginianus  Boddaert,  Elenchus  Animalium,  vol.  1,  p.  136,  1785. 

Deer  once  ranged  in  large  numbers  over  all  of  Alabama,  but  they 
are  now  exterminated  in  all  but  the  wilder  and  more  inaccessible 
parts.  Their  former  abundance  is  attested  by  the  records  of  early 
travelers.  Bartram,  who  visited  the  State  in  1776,  notes  that  in  the 
region  of  the  lower  Tombigbee  River,  about  300  deer  were  killed 
annually  by  one  planter;59  and  Gosse  records  them  as  being  very 
abundant  in  Dallas  County  in  1858,  when  70  were  killed  in  a  single 
hunt.60 

The  only  part  of  the  State  where  deer  are  still  abundant  is  in  the 
big  wooded  swamps  of  the  lower  Tensaw  and  Mobile  Rivers.  In 
that  region  a  number  are  killed  during  the  open  season  every  fall. 
They  are  hunted  with  dogs,  each  hunter  selecting  a  "  stand,"  where  he 
remains  in  expectation  that  the  deer  will  pass  within  range  of  his 
gun — usually  a  shotgun  loaded  with  buckshot.  The  deer  take  readily 
to  the  water  and  swim  easily  from  one  island  to  another  in  this  great 
swamp ;  in  this  way  they  are  able  to  keep  ahead  of  the  dogs,  but  are 
often  shot  while  swimming  a  creek  or  river  or  when  crossing  an 
opening  in  the  timber.  Deer  are  still  found  in  moderate  numbers 
in  the  sandhills  and  swamps  of  southern  Baldwin  County.  In  1915 
they  were  reported  to  be  numerous  between  Foley  and  the  shores  of 
Perdido  Bay  and  were  occasionally  seen  near  Orange  Beach,  on  the 
south  side  of  the  bay.  'Twenty  years  ago  or  more  they  were  common 
in  the  sandhills  and  small  swamps  of  Mobile  County,  but  now  ap- 
parently all  have  been  exterminated  from  that  region. 

Small  numbers  of  deer  still  range  over  the  hill  country  from  Win- 
ston to  Colbert  Counties;  three  were  killed  in  November,  1916,  on 
Buzzard  Roost  Creek,  south  of  Barton.  Reports  during  the  same 
year  indicated  that  a  few  deer  still  remained  in  northern  Cleburne 
County  near  Borden  Springs.  In  Russell  County  they  are  said  to 
have  been  exterminated  within  the  last  decade.  A  specimen  in  the 
National  Museum  collection  was  taken  at  Selma  in  1891. 

Specimens  have  been  examined  from  Orange  Beach,  Gravine  Island 
(Tensaw  River),  Mount  Vernon  Barracks,  Selma,  and  Barton. 
These  all  agree  rather  closely  with  a  skin  of  virginianus  from  Hali- 

59  Bartram,  W.,  Travels,  p.  410,   1792. 

«°  Gosse,  P.,  Letters  from  Alabama,  p.  266,  1859. 


76  NORTH  AMERICAN   FAUNA.  [No.  45 

fax,  North  Carolina,  and  differ  from  skins  of  osceola  in  darker  color. 
The  skulls  also  of  both  the  Barton  and  the  Tensaw  River  specimens 
agree  essentially  with  the  two  skulls  of  virgvnicmus  examined  (Hali- 
fax, North  Carolina,  and  Meadow  Creek  Mountain,  Greenbrier 
County,  West  Virginia)  ;  compared  with  skulls  of  osceola  from 
Florida,  these  Alabama  specimens  (as  well  as  those  from  North 
Carolina  and  West  Virginia)  differ  but  little  in  size  or  other  charac- 
ters, except  that  in  osceola  (as  stated  by  Bangs  in  the  original  de- 
scription) the  nasal  and  premaxillary  bones  meet,  while  in  vir- 
gvniomus  they  are  separated  by  an  arm  of  the  maxillary.  In  describ- 
ing osceola,  Bangs  made  comparison  with  the  large  northeastern 
deer,  since  separated  by  Miller  under  the  name  borealis,  so  that  the 
differences  between  the  Florida  deer  and  typical  virgmianus  are  much 
less  than  would  be  inferred  from  reading  Bangs's  description. 

The  horns  of  the  Alabama  deer  are  of  medium  size,  decidedly 
smaller  than  those  of  the  Louisiana  deer  (Odocoileus  louisianae)  ; 
the  breadth  of  beam  in  a  fully  adult  buck  from  Tensaw  River  is  14f 
inches.  The  hoofs  on  the  hind  feet  in  the  specimen  from  the  Colbert 
County  hills  are  decidedly  shorter  and  broader  than  those  from  the 
swamps  and  sandhills  of  the  coast  or  than  those  of  the  Florida  deer. 

The  general  color  tone  of  the  upperparts  and  legs  in  winter  pelage 
is  cinnamon-drab,  much  mixed  with  blackish,  the  muzzle  and  ears 
light  drab ;  in  the  Barton  specimen  there  is  a  band  of  blackish  along 
the  middle  line  of  the  back  and  the  upper  surface  of  the  tail  is  cin- 
namon, becoming  fuscous  toward  the  tip;  in  the  Baldwin  County 
specimens  the  surface  hairs  on  the  tail  are  likewise  cinnamon,  but  by 
reason  of  wearing  away  of  the  tips  the  basal  color  (blackish  brown 
or  clove-brown)  appears  as  a  solid  band.  The  summer  coat  is  a 
bright  tan  color,  very  different  from  the  drab  winter  coat. 


BIBLIOGRAPHY. 


1792.  Bartram,  William.  Travels  through  North  and  South  Carolina,  Georgia, 
East  and  West  Florida,  the  Cherokee  Country,  the  extensive  terri- 
tories of  the  Muscogulges,  or  Creek  Confederacy,  and  the  country  of 
the  Chactaws.  8vo.,  London,  pp.  520.  Reprinted  from  original  edition, 
pp.  522,  Philadelphia,  1791. 

Account  of  the  author's  travels  in  Alabama  from  Uche  town  to 
Mobile  and  of  an  excursion  from  that  place  up  the  Tensaw  River. 
Mention  of  a  litter  of  young  wolves  seen  near  the  Tallapoosa  River. 
Casual  reference  to  deer,  bears,  "  tygers  "  (cougars),  and  wolves. 

1824.  Hodgson,  Adam.  Letters  from  North  America  written  during  a  tour  of 
the  United  States  and  Canada.    2  vols.,  8vo.,  London. 

Account  (in  vol.  1,  pp.  117-154,  262-273)  of  a  journey  across  Ala- 
bama; casual  references  to  "panthers"  (cougars),  gray  foxes,  and 
bears. 

1833.  Stuart,  James.    Three  years  in  North  America.    2  vols.,  8vo.    Edinburgh. 

Account  (in  vol.  2,  pp.  164-226)  of  a  journey  through  Alabama  in 
1830 ;  casual  mention  of  deer. 

1834.  Arfwedson,  C.  D.     The  United  States  and  Canada  in  1832,  1833,  and 

1834.    2  vols.,  8vo.     London. 

Account  (in  vol.  2,  pp.  1-^7)  of  a  journey  in  1832  across  Alabama; 
casual  mention  of  deer  seen  in  the  Alabama  River. 
1846-54.     Audubon,  John  James,  and  John  Bachman.     The  viviparous  quad- 
rupeds of  North  America.    3  vols.,  royal  8vo.,  New  York. 

Numerous  incidental  references  to  mammals  occurring  in  Alabama. 

1856.  Lanman,  Charles.     Adventures  in  the  wilds  of  the  United  States  and 

British  American  Provinces,  2  vols.,  8vo.,  Philadelphia. 

Four  chapters  (in  vol.  2,  pp.  146-188)  on  Alabama,  with  a  few 
casual  references  to  mammals. 

1857.  Baird,  S.  F.     Mammals  of  North  America :  Report  of  explorations  and 

surveys  to  ascertain  the  most  practicable  route  for  a  railroad  from 
the  Mississippi  River  to  the  Pacific  Ocean.    Vol.  VIII,  4to. 
A  few  specimens  listed  from  Alabama. 
1859.     Gosse,  Philip  Henry.    Letters  from  Alabama  (U.  S.)  chiefly  relating  to 
natural  history.    16mo.,  London,  pp.  xii,  306. 

Brief  references  to  a  number  of  mammals  and  extended  accounts 
of  deer  hunting  and  of  a  nocturnal  'possum  hunt. 
1861.     Gesner,  William.     On  the  habits  of  the  pouched  rat,  or  salamander 
(Geomys  pineti) ,  of  Georgia.     Ann.  Rept.  Smiths.  Inst,  for  1860,  pp. 
431^33. 

Description  of  pocket  gopher  mounds  in  Russell  County. 
1864.     Allen,  Harrison.     Monograph  of  the  bats  of  North  America.     Smiths. 
Misc.  Coll.,  vol.  7,  art.  I,  pp.  i-xxiii,  1-85. 

Nycteris  borealis  and  Corynorhinus  macrotis  recorded  from  Eutaw. 

77 


78  NORTH   AMERICAN   FAUNA.  [No.  45 

1877.     Coues,  Elliott.    Fur-bearing  animals;  a  monograph  of  North  American 
Mustelidae,  etc.     Misc.  Pub.  No.  8,  U.  S.  Geol.  Surv.  Terr.,  pp.  348, 
pis.  20. 
Record  of  Putorius  vison  from  Tuscaloosa. 
1877.     Coues,   Elliott,    and   J.   A.    Allen.     Monographs    of   North   American 
Rodentia.     Rept.  U.  S.  Geol.  Surv.  Terr.,  vol.  11,  pp.  1091,  pis.  7. 
Contains  a  very  few  records  of  Alabama  mammals. 

1889.  Merriam,  C.  Hart.     Report  of  the  ornithologist  and  mammalogist.    Ann. 

Rept.  Dept.  Agr.  for  1888,  pp.  477-536. 

Mention  of  a  mink  killing  poultry  at  Greensboro. 

1890.  Merriam,  C.  Hart.     Contribution  toward  a  revision  of  the  little  striped 

skunks  of  the  genus  Spilogale,  with  descriptions  of  seven  new  species. 
North  Amer.  Fauna  No.  4,  pp.  1-15. 

Original  description  of  Spilogale  ringens  [=putorius]  from  Greens- 
boro. 

1891.  True,   Frederick  W.     The  puma  or  American  lion    (Felts  concolor  of 

Linnaeus).    Rept.  U.  S.  Nat.  Mus.  for  1888-89,  pp.  591-608,  pi.  XCIV. 
Record  of  the  puma  from  Dekalb  County. 

1893.  Allen,  Harrison.     A  monograph  of  the  bats  of  North  America.     Bull. 

No.  43,  U.  S.  Nat.  Mus.,  pp.  198,  pis.  38. 

Corynorhinus   macrotis    and    Atalapha    noveboracensis    [=Nycteri$ 
borealis]  recorded  from  Eutaw. 

1894.  Chapman,  Frank  M.     Remarks  on  certain  land  mammals  from  Florida, 

with  a  list  of  the  species  known  to  occur  in  the  State.     Bull.  Amer. 
Mus.  Nat.  Hist.,  vol.  6,  pp.  333-346. 

Records  of  beavers  and  muskrats  from  Alabama ;  remarks  on  the 
type  specimen  of  Peromyscus  megacephalus  from  Woodville. 

1894.  Rhoads,  Samuel  N.    Descriptions  of  four  new  species  and  two  new  sub- 

species of  white-footed  mice  from  the  United  States  and  British  Co- 
lumbia.    Proc.   Acad.  Nat.   Sci.   Philadelphia,   for  1894,   pp.  253-261. 
Original  description  of  Sitomys   [Peromyscus]    megacephalus  from 
Woodville. 

1895.  Merraim,  C.  Hart.    Monographic  revision  of  the  pocket  gophers,  family 

Geomyidae    (exclusive   of  the   species   of   Thomomys).     North  Amer. 
Fauna  No.  8,  pp.  258.  pis.  19,  4  maps. 

Original  description  of  Geomys  tusa  mobilensis  from  Point  Clear, 
Mobile  Bay. 

1895.  Merriam,   C.   Hart.     Revision  of  the  shrews  of  the  American  genera 

Blarina  and  Notiosorex.     North  Amer.  Fauna  No.  10,  pp.  5-34. 

Records   of  Blarina   brevicauda   carolinensis   and   Cryptotis   parva 
from  Alabama. 

1896.  True,  Frederick  W.     A  revision  of  the  American  moles.     Proc.  U.  S. 

Nat.  Mus.,  vol.  19,  pp.  1-111 ;  pis.  4. 

Scalopus  aquaticus  recorded  from  Woodville. 

1897.  Miller,   Gerrit  S.,  Jr.     Revision  of  the  North  American  bats  of  the 

family  Vespertilionidae.    North  Amer.  Fauna  No.  13,  pp.  135,  pis.  3. 

Five  species  of  bats  listed  from  Alabama. 
1900.     Bailey,  Vernon.     Revision  of  American  voles  of  the  genus  Mierotus. 
North  Amer.  Fauna  No.  17,  pp.  88,  pis.  5. 

Microtus  pinetorum  auricularis  recorded  from  Greensboro. 


1921.]  BIBLIOGRAPHY.  79 

1901.     Allen,  J.  A.     A  preliminary  study  of  the  North  American  opossums  of 
the  genus  Didelphis.    Bull.  Amer.  Mus.  Nat.  Hist.,  vol.  14,  pp.  149-188 ; 
pis.  XXII-XXV. 
Didelphis  virginiana  pigra  recorded  from  Sylacauga. 

1901.  Howell,   Arthur   H.     Revision   of  the   skunks   of  the   genus    Chincha 

[=  Mephitis].    North  Amer.  Fauna  No.  20,  pp.  47,  pis.  8. 
Mephitis  elongata  recorded  from  Baldwin  County. 

1902.  Bailey,  Vernon.     Synopsis  of  the  North  American  species  of  Sigmodon. 

Proc.  Biol.  Soc.  Washington,  vol.  15,  pp.  101-116. 
Records  of  Sigmodon  hispidus  from  Alabama. 
1906.     Howell,  Arthur  H.     Revision  of  the  skunks  of  the  genus  Spilogale. 
North  Amer.  Fauna  No.  26 ;  pp.  55,  pis.  10. 

Spilogale  putorkis  recorded  from  Greensboro  and  Mobile. 
1909.     Howell,  Arthur  H.     Notes  on  the  distribution  of  certain  mammals  in 
the  southeastern  United  States.    Proc.  Biol.  Soc.  Washington,  vol.  22, 
pp.  55-68. 
Records  of  twenty  species  from  Alabama. 
1909.     Nelson,  E.  W.     The  rabbits  of  North  America.     North  Amer.   Fauna 
No.  29 ;  pp.  314,  pis.  13,  fig.  19. 

Records  of  four  forms  of  Sylvilagus  from  Alabama. 

1909.  Osgood,  Wilfred  H.    Revision  of  the  mice  of  the  American  genus  Pero- 

myscu*.    North  Amer.  Fauna  No.  28 ;  pp.  285 ;  pis.  8. 
Records  of  five  forms  of  Peromyscus  from  Alabama. 

1910.  Goldman,  Edward  A.     Revision  of  the  wood  rats  of  the  genus  Neotoma. 

North  Amer.  Fauna  No.  31,  pp.  124,  pis.  8. 

Records  of  Neotoma  floridana  rubida  and  Neotoma  Pennsylvania 
from  Alabama. 

1911.  Hollister,  N.    A  systematic  synopsis  of  the  muskrats.    North  Amer. 

Fauna  No.  32,  pp.  47,  pis.  6. 

Fiber  zibethicus  zibethicus  recorded  from  Reform. 
1913.     Hollister,  N.     A  synopsis  of  the  American  minks.     Proc.   U.   S.  Nat. 
Mus.,  vol.  44,  pp.  471-480. 

Mustela  vison  lutreocephala  recorded  from  Alabama. 

1913.  Howell,    Arthur    H.     Description    of   a    new   weasel    from    Alabama. 

Proc.  Biol.  Soc.  Washington,  vol.  26,  pp.  139-140. 

Original  description  of  Mustela  peninsulae  olivacea  from  Autauga- 
ville., 

1914.  Howell,  Arthur  H.     Revision  of  the  American  harvest  mice   (genus 

Reithrodontomys).    North  Amer.  Fauna  No.  36,  pp.  97,  pis.  7. 
Records  of  Reithrodontomys  humulis  merriami  from  Alabama. 

1914.  Jackson,  Hartley  H.  T.     New  moles  of  the  genus  Sealopus.    Proc.  Biol. 

Soc.  Washington,  vol.  27,  pp.  19-22. 

Original   description  of  Sealopus  aquaticus   howelli  from  Autauga- 
ville. 

1915.  Howell,  Arthur  H.     Descriptions  of  a  new  genus  and  seven  new  races 

of  flying  squirrels.     Proc.  Biol.  Soc.  Washington,  vol.  28,  pp.  109-113. 

Original  description  of  Glaucomys  volans  saturatus  from  Dothan. 
1915.     Howell,  Arthur  H.     Revision  of  the  American  marmots.     North  Amer. 
Fauna  No.  37,  pp.  80,  pis.  15. 

Marmota  monax  monax  recorded  from  Ardell. 


80  NORTH   AMERICAN   FAUNA.  [No.  45 

1915.  Jackson,   Hartley   H.   T.    A   review    of   the   American   moles.    North 

Amer.  Fauna  No.  38,  pp.  100,  pis.  6. 

Records  of  Scalopus  aquations  howelli  from  Alabama. 

1916.  Wallace,  John  H.,  Jr.    The  mammals  of  Alabama.    Fifth  Bien.  Kept, 

Dept.  Game  and  Fish,  State  of  Alabama,  pp.  8-21. 

Fifty  species  listed ;  the  annotations  are  of  a  general  nature  and 
contain  no  definite  information  as  to  occurrence  in  the  State. 
1918.     Goldman,  Edward  A.    The  rice  rats  of  North  America  (genus  Oryzomys). 
North  Amer.  Fauna  No.  43,  pp.  100,  pis.  6. 
Records  of  Oryzomys  palustris  palustris  from  Alabama. 
1918.     Howell,  Arthur  H.     Revision  of  the  American  flying  squirrels.     North 
Amer.  Fauna  No.  44,  pp.  64,  pis.  7. 

Records  of  Glaucomys  voluns  saturatus  from  Alabama. 


INDEX. 


A. 
Acacia,  rose,  11. 
Acer  drummondii,  9,  14. 

floridanum,  14. 

leucoderme,  14. 
Aesculus  glabra,  11. 
Agalinis  aphylla,  10. 

fasciculata,  14. 
Allspice,  Carolina,  9. 
Andromeda,  low,  15. 
Antrostomus  Caroline n six,  12. 
A  plos  apios,  74. 

tuberoxa,  65. 
Arabia  racemoxa,  11 . 
Arundinaria  macroxperma,  14. 
Axclepiax  humixtrata,  14. 

michauxii,  16. 

quadrifolia,  11. 
Ascyrumhypericoidex,  14. 

pumilum,  15. 
Ash,  blue,  11. 

green,  9. 

water,  9,  14. 
./isimma  anguxtifolia,  Id. 
ulster  adnatux,  10. 

azureux,  11. 

exilix,  15. 

graeilix,  11. 

purpuratux,  15. 
Aster,  hoary  golden,  15. 

hoary-leaved  golden,  15. 

large-flowered  golden,  16. 

seaside,  15. 

sky-blue,  11. 

slender,  11. 

small-leaved,  16. 

wandlike,  15. 
Atalapha  borealix  seminola,  26. 
yljaZea  arborexcenx,  11. 
Azalea,  sweet-scented,  11. 

B. 

Balm,  scarlet,  15. 

seaside.  15. 
Baptixia  lanceolata,  14. 
Basil,  swamp,  14. 

49663°— 21 6 


Basswood,  8. 

southern,  14. 
Bat,  evening,  13,  27-28. 

gray,  23-24. 

hoary,  27. 

large  brown,  25-20. 

LeConte  big-eared,  28. 

LeOonte  free-tailed,  13,  28-29. 

little  brown,  24. 

mahogany,  13,  20-27. 

red,  26. 

silver-haired,  24-25. 
Bay,  red,  9,  14. 

swamp  red,  14. 

white,  9. 
Bear,  Florida  black,  29-30. 
Beaver,  Carolina,  07-70. 
Beech,  9. 

water,  68. 
Benzoin,  74. 
Betula  lenta,  11. 
Birch,  cherry,  11. 
Bitter-sweet,  11. 

Blarina  brevicauda  carolinenxix,    13, 21-22. 
Blazing-star,  15. 

slender,  16. 
Blueberry,  Elliott,  14. 

evergreen,  14. 

fuscous,  14. 

mountain,  11. 
Bluehearts,  southern,  14. 
Blue  jack,  9. 
Bluets,  southern,  14. 
Bobcat,  42-43. 

Bonaxa  umbellus  umbellux,  10. 
Boneset,  upland,  11. 
Boxthorn,  Carolina,  15. 
Brevjerla  humistrata,  14. 
Breweria,  low,  14. 
Buchnera  elongata,  14. 
Buckeye,  9. 

fetid,  11. 
Buckthorn,  southern,  14. 
Bumelia  lycioides,  14. 
Buteo  lineatux  alleni,  12. 
Butternut,  11. 

81 


82 


NORTH   AMERICAN   FAUNA. 


[No.  45 


C. 

Cactus,  crowfoot,  15. 
Cahile  chapmani,  15. 
Calycanthvs,  14. 

floridm,  9,  14. 
Campion,  round-leaved,  11. 
Cane,  14. 
Cane-cutter,  71. 
Canis  cinereo-argenteus,  33. 
fioridanus,  30-32. 
fulvus,  32. 
Carpinus  caroliniana,  68. 
Castor  canadensis  carolinensis,  67-70.     / 

zibcthicus,  55. 
Catalpa  catalpa,  14. 
Catamount,  42-43. 
Cedar,  coast  red,  15. 
red,  7,  8. 
white,  9. 
Celastrus  scandens,  11. 
Celtis  mississippiensis,  14. 
Ceratiola  ericoides,  9,  15. 
Cerchneis  sparveria  sparveria,  10. 
Cervus  virginianus,  75. 
Chamaecrista  multipinnata,  14. 
Chamaecy parts  thyoides,  9. 
Cherry,  mountain  black,  11. 
Chestnut,  7,  8. 
Chimaphila  maculata,  11. 
Chipmunk,  southwestern,  62-63. 
Chrysobalanus  oblongifolius,  15. 
Chrysoma  pauciflosculosa,  16. 
Chrysopsis  oligantha,  16. 
pilosa,  15. 
trichophylla,  15. 
Chuck-will's-widow,  12. 
Cirsium  leconlei,  16. 

nuttalli,  16. 
Claytonia  virginica,  11. 
Cliftonia  monophylla,  9,  15. 
Clinipodium  carolinianum ,  14. 

coccineum,  15. 
Colaptes  auratus  auratus,  12. 
Cone-flower,  mountain,  11. 
Conradina  canescens,  15. 
Coragyps  urubu  urubu,  12. 
Corylus  rostrata,  62. 
Corynorhinus  macrotis,  28. 
Cottontail,  eastern,  70-71. 

New  England,  71. 
Cottonwood,  swamp,  9. 
Cougar,  41-42. 
Cracca  chrysophylla,  15. 
hispidula,  14. 
smallii,  15. 


Crotalaria  purshii,  15. 

rotundifolia,  14. 
Cryptotis  parva,  22. 
Cyanocitta  cristata  cristata,  12. 
Cynoctonum  mitreola,  14. 
Cypress,  9. 

pond,  9,  15. 

swamp,  13. 
Cypripedium  acaule,  11. 
Cyrilla  racemiflora,  9,  14. 

D. 

Deer  grass,  14. 

Deer,  Virginia,  75-76. 

Dendroica  dominica  dominica,  12. 

Devil-wood,  14. 

Dicerandra  linear  if olia,  16. 

Didelphis  virginiana  pigra,  20. 

virginiana  virginiana,  18-19. 
Diervilla  rivularis,  11. 
Drosera  brevifolia,  14. 

capillaris,  15. 
Dryobates  pubescens  pubescens,  12. 

villosus  auduboni,  12. 

E. 

Elm,  winged,  14. 
Epilobium  coloratum,  11. 
Eptesicus fuscus fuscus,  25-26. 
Erigeron  quercifolius,  15. 
Eriogonum,  downy,  14. 
Eriogonum  tomentosum,  14. 
Eupatorium  sessilifolium,  11. 

F. 

False-indigo,  lanceolate,  14. 
.FVfrs  couguar,  41-42. 
Fetterbush,  14. 
Fiber  rivalicius,  13,  57-58. 

zibethicus  zibethicus,  55-57. 
Fleabane,  southern  daisy,  15. 
Flicker,  12. 
Fog-fruit,  16. 
Fox,  gray,  33-34. 

red,  32-33. 
Fraxinus  caroliniana,  9,  14. 

lanceolata,  9. 

quadrangulata,  11. 

G. 

Galactia  erecta,  15. 
Jloridana,  15. 
Gallinule,  purple,  13. 
Gaura  angustifolia ,  15. 


1921.] 


INDEX. 


83 


Gaura,  narrow-leaved.  15. 
Gaylussacia  frondosa  tomentosa,  15. 
Gelsemium  sempervirens,  14. 
Geomys  tuza  mobilensis,  13,  59-60. 
Geothlypis  trichas  ignola,  12. 
Gerardia,  fascicled,  14. 

leafless,  16. 
Gilia  rubra,  14. 

Glaucomys  volans  saturatus,  67. 
Goat's-rue,  15. 

hispid,  14. 

prostrate,  15. 
Goldenrod,  broad-leaved,  11. 

few-flowered,  16. 

hispid,  11. 

narrow-leaved,  15. 

short-leaved,  15. 

white,  11. 
Gopher,  Alabama  pocket,  13,  59-60. 
Grackle,  boat-tailed,  13. 

Florida,  13. 
Grape,  blue,  11. 
Grosbeak,  blue,  12. 
Ground-cherry,  narrow- leaved,  16. 
Ground  hog,  10,  60-62. 
Grouse,  ruffed,  10. 
Guiraca  caerulea  caerulea,  12. 
Gum,  sour,  8. 

southern  black,  9,  14. 

sweet,  68. 

tupelo,  9,  14. 


Habenaria  repens,  15. 

Hackberry,  southern,  14. 

Halesia  diptera,  14. 

Haw,  opossum,  14. 

Hawk,  Florida  red-shouldered,  12. 

sparrow,  10. 
Hazelnut,  62. 
Hedge-hyssop,  rough,  16. 
Hcdysarum  cannabinum,  62. 
Helenium  brevifolium,  15. 
Helenium,  short-leaved,  15. 
Helianihcmum  arenicola,  15. 
Helianthus  radula,  16. 
Heliopsis  minor,  15. 
Helmitheros  vermivorus,  10. 
Hemlock,  11. 

Hespcromys  gossypinus,  48. 
Hibiscus  aculealus,  14. 
Hickory,  nutmeg,  13. 

water,  9, 13. 


Hicoria  aquatica,  9, 13. 

myristicaeformis,  13. 

pecan,  13. 
Holly,  9. 

Honeysuckle,  bush,  11. 
Horsemint,  Bradbury,  11. 
Houstonia,  narrow-leaved,  11. 

round-leaved,  16. 
Houstonia  procumbens,  16. 

pusilla,  14. 

tenuifolia,  11. 
Huckleberry,  hoary,  15. 
Hydrangea,  oak -leaved,  8. 
Hydrangea  quercifolia,  8. 
Hypericum  opacum,  15. 
Hyptis  rugosa,  14. 


Ilex  lucida,  14. 

monticola,  11. 

opaca,  9. 

vomitoria,  10. 
Illicium  fioridanum,  9. 
Illicium,  sweet,  9. 
Indian-plantain,  lance-leaved,  16. 
Indigo,  wild,  14. 
Indigofera  caroliniana,  14. 
Inkberry,  tall,  14. 
Ionornis  martinicus,  13. 
Ipomaea  caroliniana,  14. 

littoralis,  16. 

pes-caprae,  16. 


Jay,  Florida  blue,  12. 
Jessamine,  yellow,  14. 
Juglans  cinerea,  11. 
Juncus  roemerianus,  10. 
Juniperus  barbadensis,  15. 

K. 

Kahnia,  8. 

Kuhnistera  gracilis,  15. 
pinnata,  14. 


Lacinaria  elegans,  15. 

gracilis,  16. 
Lactuca  graminifolia,  15. 
Lanius  ludovicianus  ludovicianus,  12. 
Lapithea  gentianoides,  16. 
Lasionycteris  noctivagans,  24-25. 


84 


NORTH   AMERICAN    FAUNA. 


[No.  45 


Laurel,  8. 

Leaf-cup,  Canada,  11. 
Leatkerwood,  9,  14. 
Lcchea  divaricala,  15. 
Lepus  aquaticus,  71. 

nuttalli  mallurus,  70. 

palustris,  73. 

sylvaticus  transitionalis,  71. 
Lettuce,  narrow  wild,  15. 
Ligusticum  canadense,  11. 
Lily,  atamasco,  74. 
Limnothlypis  swainsonii,  12. 
Linaria  floridana,  16. 
Lippia  nodiflora,  1G. 
Liquidamber  styraciflua,  68. 
Live-oak,  dwarf,  15. 

twin,  15. 
Lobelia  brevifolia,  16. 
Lobelia,  short-leaved,  16. 
Locust,  clammy,  11. 
Loosestrife,  four-leaved,  11. 
Lovage,  Canada,  11. 
Ludwigia,  hoary,  14. 
Ludwigia  pilosa,  14. 
Lungwort,  Virginia,  11. 
Lupine,  hairy,  14. 

spreading,  14. 
Lupinus  diffusus,  14. 

villosus,  14. 
Lutra  canadensis  canadensis,  40^11. 
Lycium  carolinianum,  15. 
Lynx  ruffus  floridanus,  42-43. 
Lysimachia  quadrifolia,  11. 

M. 

Magnolia,  8,  14,  '64. 
Magnolia  foetida,  9,  14,  64. 

virginiana,  9. 
Maple,  Drummond  red,  14. 

Florida,  14. 

pale-bark,  14. 

red,  9. 

swamp,  9. 
Marmota  monax  monax,  10,  60-62. 
Meadow-beauty,  lance-leaved,  14. 

swamp,  15. 
Meadowlark,  southern,  12. 
Meadow-rue,  smooth,  11. 
Megaquiscalus  major  major,  13. 
Meibomia  tenuifolia,  14. 
Mephitis  mephitis  elongata,  39. 

mephitis  nigra,  39. 
Mertensiavirginica,  11. 
Mesadenia  lanceolata,  16. 


Microtus  pinetorum  auricularis,  55. 
Milk-pea,  erect,  15. 

Florida,  15. 
Milkweed,  four-leaved,  11. 

Michaux,  16. 
Milkwort,  Chapman,  15. 

large-flowered,  14. 
Mink,  37. 
Miterwort,  14. 
Moccasin  flower,  11. 
Mole,  Howell,  20-21. 
Monarda  bradburiana,  11. 
Morning-glory,  Carolina,  14. 

goat's-foot,  16. 

seaside,  16. 
Morus  rubra,  65. 
Mouse,  cotton,  13,  48. 

eastern  harvest,  44. 

house,  59. 

Merriam  harvest,  43-44. 

old-field,  13, 44-46. 

pine,  54-55. 

Rhoads  cotton,  49-50. 

southern  golden,  13,  50. 

white-footed,  46-48. 

white-fronted  beach,  13,  46. 
Mouse-ears,  15. 
Mulberry,  black,  65. 
Mus  alexandrinvA,  59. 

(Calomys)  aurcolus,  50. 

floridana,  52. 

humulis,  44. 

monax,  60. 

musculus  musculus,  59. 

norvegicus,  58. 

palustris,  51. 

polionotus,  44. 

rattus,  58. 
Musculus  leucopus,  46. 
Muskrat,  55-57. 

Louisiana,  13,  57-58. 
Mustela  Intra  canad,ensis,  40. 

noveboracrnsis,  10,  35-36. 

peninsulac  olivacea,  13,  36. 

vison  mink,  37. 
My otis  grisescens,  23-24. 

lucifugus  lucifugus,  24. 
Myrica  cerifera,  13. 

inodora,  15. 

N. 

Neotoma,  47. 

floridana  floridana,  13, 36, 52-53. 
pennsylvanica,  53-54. 


1921.] 


INDEX. 


85 


Nuthatch,  brown-headed,  12. 

Florida  white-breasted,  12. 
Nycteris  borealis  bvrealut,  26. 

dnerea,  27. 

seminola,  13,  26-27. 
Nycticea  cynocephala,  28. 
Nycticeius  humeralis,  13,  27-28. 
Nyctinomus  cynocephalus,  13, 28-29. 
Nymphaea  chartacea,  15. 
Nyssa  aquatica,  9,  14. 

biflora,  9, 14. 

sylvatica,  8. 

Oak,  chestnut,  8, 11. 

laurel,  9, 14. 

live,  9. 

overcup,  8,  9,  13. 

post,  8. 

scarlet,  11. 

Spanish,  8. 

swamp  Spanish,  13. 

Texas  white,  8,  13. 

turkey,  9, 13. 

Upland  willow,  14. 

water,  8,  9,  14. 
Odoccrileus  virginianus  virgvnianus,  75-76. 
Opossum,  Florida,  20. 

Virginia,  18-19. 
Opuntia  pes-corvi,  15. 
Orchid,  showy,  11. 
Orchis,  creeping,  15. 
Orchis  spectabilis,  11. 
Oryzomys  palustris  palustris,  13, 51. 
Osmanthus  amcricana,  14. 
Osmorrhiza  claytoni,  11. 
Otter,  40-41. 
Otus  asio  asio,  12. 
Ovenbird,  10. 
Owl,  barred,  22. 

Florida  barred,  12. 

Florida  screech,  12. 

Oxeye,  15. 

P. 

Palmetto,  saw,  13. 

Panther,  41-42. 

Papaw,  narrow-leaved,  15. 

Parrot-beak,  15. 

Passerherbulus  maritimus  howelli,  13. 

Pea,  sensitive,  14. 

Pecan,  13. 

Pennyroyal,  large-flowered,  16. 

Peromyscus  gossypinus,  13. 

gossypinus  gossypinus,  48. 

gossypinus  megacephalus,  49-50. 

leucopus  leucopxis,  46-48. 


Peromyscus  nuttalli  aureolus,  13,  50. 
polionotus  albifrons,  13,  46. 
polionotus  polionotus,  13,  44-46. 
Persea  borbonia,  9,  14. 

pubescens,  14. 
Peucaea  aestivalis  bachmani,  12. 
Philadelphits  hirsutus,  11. 
Phlox,  creeping  purple,  11. 

Florida,  14. 
Phlox  Jloridana,  14. 

reptans,  11. 
Phoebe,  10. 

Phrenopicus  borealis,  12. 
Physalis  angustifolia,  16. 
Pieris  nitida,  14. 

phillyreaefolia,  15. 
Pine,  loblolly,  9,  13. 
longleaf,  9,  13. 
sand,  15. 
shortleaf,  8. 
southern  spruce,  8. 
spruce,  13. 
swamp,  9,  15. 
Pinus  clausa,  15. 
echinata,  8. 
elliottii,  9,  15. 
glabra,  8,  13,  68. 
palustris,  8,  9,  13,  65. 
taeda,  8,  9,  13. 
virginiana,  8. 
Pinweed.  divaricate,  15. 
Pipilo  erythrophthalmus  canaster,  12. 
Pipistrelle,  southeastern,  25. 
Pipistrellus  subflavus  subflavus,  25. 
Piranga  olivacea,  10. 
Pitcheria  galactioides,  15. 
Pitcheria,  pine-barren,  15. 
Pitcher-plant,  red-flowered,  14. 
Pitymys  pinetorum  auricularis,  55. 

pinetorum  pinetorum,  54-55. 
Planera  aquatica,  14. 
Planer-tree,  14. 

Planesticus  migratorius  achrusterus,  10. 
Plecotus  macrotis,  28. 
Plum,  deer,  15. 
prairie,  14. 
Pocket  gopher,  Alabama,  13,  59-60. 
Poly  gala  chapmani,  15. 

grandiflora,  14. 
Polymnia  canadensis,  11. 
Pondlily,  small  yellow,  15. 
Potato,  wild,  65,  74. 
Prairie-clover,  pine-barren,  14. 

slender,  15. 


86 


NORTH   AMERICAN    FAUNA. 


[No.  45 


Prickly-ash,  southern,  14. 
Privet,  mountain,  11. 
Proeyon  lotor  lotor,  34-35. 
Prunus  serotina  neomontana,  11. 

umbellata,  14. 
Psammomys  pinetorum,  54. 
Puma,  41-42. 
Putorius  noveboracensis ,  35. 

Q. 

Queen 's-delight,  14. 
Quercus  brevifolia,  9,  14. 

catesbaei,  9,  13. 

coccinea,  11. 

durandi,  8,  13. 

geminata,  15. 

laurifolia,  9,  14. 

fr/rata,  8,  9,  13. 

minor,  8. 

myrtifolia,  15. 

m'^ra,  8,  9,  14. 

pagodaefolia,  13. 

prinus,  8,  11. 

virginiana,  9. 

virginiana  maritima,  15. 
Quiscalus  quiscula  quiscula,  13. 

R. 

Rabbit,  coast  swamp,  13,  73. 

marsh,  13,  73-75. 

swamp,  13,  71-73. 
Rabbit's-milkweed,  14. 
Raccoon,  34-35. 
Rail,  Louisiana  clapper,  10,  13. 
Rallus  crepitans  saturatus,  10,  13. 
Rat,  Allegheny  cliff,  10,  53-54. 

barn,  58. 

black,  58-59. 

cotton,  13,  22,  52. 

Florida  wood,  13,  52-53. 

gopher,  52,  58. 

Norway,  58. 

roof,  59. 

swamp  rice,  13, 51. 

wood,  36,  37. 
Rattlebox,  Pursh,  15. 

round-leaved,  14. 
Rattlesnake,  50. 
Rattus  norvegicus,  58. 

rattus  alexandrinus,  59. 

rattus  rattus,  58-59. 
Reithrodontomys  humulis  humulis,  44. 

humulis  merriami,  43-44. 


Rhexia  glabella,  14. 

lanceolata,  14. 

stricta,  15. 
Robin,  southern,  10. 
Robinia  hispida,  11. 

viscosa,  11. 
Rock-rose,  seaside,  15. 
Rose-mallow,  rough,  14. 
Rosinweed,  15. 
Rudbeckia  monticola,  11. 
Ruellia,  fimbriated,  14. 

night-blooming,  16. 
Ruellia  humilis,  14. 

noctiflora,  16. 
Rush,  black,  10. 

S. 

Sabbatia,  gentian,  16. 

large-leaved,  15. 
Sabbatia  macro phylla,  15. 
St.  John's- wort,  opaque,  15. 
St.  Peter's-wort,  dwarf,  15. 

southern,  14. 
Salamander,  59-60. 
Salvia,  azure,  14. 
Salvia  azurea,  14. 
Sarracenia  drummondi,  15. 

psittacina,  15. 

rubra,  14. 
Sayornis  phoebe,  10. 
Scalopus  aquaticus  howelli,  20-21. 
Scarabaeus  tityus,  22. 
Sciurus  carolinensis  carolinensis,  63-64. 

carolinensis  fuliginosus,  13,  64. 

niger  niger,  64-66. 

niger  texianus,  66-67. 
Scrub-oak,  seaside,  15. 
Sea  rocket,  15. 
Sebastiana,  14. 
Sebastiana  ligustrina,  14. 
Seiurus  aurocapillus  aurocapillus,  10. 
Serenoa  serrulata,  13. 
Sericocarpus  bifoliatus,  15. 
Setochalcis  vocifera  voci/era,  10. 
Shrew,  Bachman,  22-23. 

Carolina  short-tailed,  13,  21-22. 

least,  22. 
Shrike,  loggerhead,  12. 
Sigmodon,  22. 

hispidum,  19. 

hispidus  hispidus,  13,  52. 
Silene  rotundifolia,  11. 
Silphium  aster  iscus,  15. 


1921.] 


INDEX. 


87 


Silver-bell  tree,  southern,  14. 

Sitomys  megacephalus,  49. 

Sitta  carolinensis  carolinensis,  12. 

pusilla,  12. 
Skunk,  Alleghenian  spotted,  37-38. 

eastern,  39. 

Florida,  39. 
Solidago  angustifolia,  15. 

bicolor,  11. 

brachyphylla,  15. 

jlexicaulis,  11. 

hispida,  11. 
Sophronanthe  hispida,  16. 
Sorex  carolinensis,  21. 

longirostris,  22-23. 

parvus,  22. 
Spanish-cypress,  14. 
Sparrow,  Bachman,  12. 

Howell  seaside,  13. 
Spikenard,  11. 
Spilogale  putorius,  37-38. 

ringens,  37. 
Spring  beauty,  Virginian,  11. 
Squirrel,  Bachman  fox,  66-67. 

bayou  gray,  13,  64. 

Carolina  fox,  64-66. 

cat,  63-64. 

gray,  63-64. 

southeastern  flying,  67. 
Steironema,  mountain,  11. 
Steironema  tonsum,  11. 
Stewartia  malacodendron,  14. 
Stewartia,  Virginia,  14. 
Stillingia  sylvatica,  14. 
Stokesia,  azure-flowered,  16. 
Stokesia  laevis,  16. 
Storax,  bush,  9. 

large-flowered,  14. 

powdery,  14. 
Strix  varia  alleni,  12,  22. 
Sturnellu  magna  argutula,  12. 
Styrax  grandiflora,  9,  14. 

pulverulenta,  14. 
Sundew,  capillary,  15. 

short-leaved,  14. 
Sunflower,  false,  15. 

rayless,  16. 
Sweet  cicely,  11. 
Sweet-shrub,  14. 
Sylvilagus  aquaticus  aquaticus,  13,  71-73. 

aquaticus  littoralis,  13,  73. 

floridanus  mallurus,  70-71. 

palustris  palustris,  13,  73-75. 

transitionalis ,  71. 
Syringa,  hairy,  11. 


Tamias  strlatus  venustus,  62-63. 
Tanager,  scarlet,  10. 
Taxodium  ascendens,  9,  15. 

distichum,  9,  13. 
Telmatodytes  palustris  marianae,  13. 
Tetragonoiheca  helianihoides,  15. 
Thalictrum  dioicum,  11. 
Thistle,  LeConte,  16. 

smooth,  16. 
Thyme,  Carolina,  14. 
Tick-trefoil,  narrow-leaved,  14. 
Tilia  australis,  14. 

heterophylla,  8. 
Titi,  9,  15. 

Toad-flax,  Florida,  16. 
Towhee,  Alabama,  12. 
Trilisa  odoratissima ,  15. 
Trumpet-leaf,  Drummond,  15. 
Tsuga  canadensis,  11. 

IT. 

Ulmus  alata,  14. 

Uniola  paniculata,  46. 

Urocyon  cinereoargenteus  cinereoargenteus, 

33-34. 
Ursus  floridanus ,  29-30. 

lotor,  34. 

V. 
Vaccinium  elliottii,  14. 

fuscatum,  14. 

myrsinites,  14. 

pallidum,  11. 
Vanilla  plant,  15. 
Verbena  carolinensis,  14. 
Vermivora  bachmanii,  12. 
Vernonia  angustifolia,  15. 
Vernonia,  narrow-leaved,  15. 
Vervain,  Carolina,  14. 
Vespertilafuscus,  25. 
Vespertilio  borealis,  26. 

cinereus,  27. 

humeralis,  27. 

lucifugus,  24. 

noctivagans,  24-25. 

subflavus,  25. 
Vetch,  acute-leaved,  15. 
Viburnum  cassirwides,  11. 

nitidum,  14. 
Vicia  acutifolia,  15. 
Viola  Carolina,  14. 

leconteana,  11. 

rostrata,  11. 

striata,  11. 


88 


NORTH   AMERICAN    FAUNA. 


[No.  45, 1921. 


Violet,  Carolina,  14. 

long-spurred,  11. 

pale,  11. 
Vitis  bicolor,  11. 
Viverra  nigra,  39. 

putorius,  37. 
Vole,  bluegrass,  55. 
Vulpesfulva,  32-33. 
Vulture,  black,  12. 

W. 

Warbler,  Bachman,  12. 

Swainson,  12. 

worm-eating,  10. 

yellow-throated,  12. 
Wax-myrtle,  Bartram,  15. 

southern,  13. 
Weasel,  Alabama,  13,  36. 

New  York,  10,  35-36. 
Whip-poor-will,  10. 


Wildcat,  42^3. 

Willow-herb,  colored,  11. 

Wintergreen,  spotted,  11. 

Withe-rod,  11. 

Wolf,  southeastern,  30-32. 

Woodchuck,  southern,  10,  60-62. 

Woodpecker,  downy,  12. 

red-cockaded,  12. 

southern  hairy,  12. 
Wood- violet,  sweet,  11. 
Wren,  Marian  marsh,  13. 


X. 


Xyris  elliottii,  15. 


Yaupon,  10. 

Yellow-eyed  grass,  Elliott,  15. 

Yellow-throat,  Florida,  12. 

Z. 

Zantho.iylum  clava-herculis,  14. 


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North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  I. 


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U.  S.  DEPARTMENT  OF  AGRICULTURE 

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NORTH  AMERICAN  FAUNA 

IsTo.   46 

[Actual  date  of  publication,  June  20,  1923] 


A  BIOLOGICAL  SURVEY  OF  THE  PRIBILOF 
ISLANDS,  ALASKA 

I.  BIRDS  AND  MAMMALS 

By  Edward  A.  Preble,  Assistant  Biologist,  Division  of  Biological 

Investigations,  and  W.  L.  McAtee,  In  Charge  Food  Habits 

Research;  Bureau  of  Biological  Survey 

II.  INSECTS,  ARACHNIDS,  AND  CHIL0P0DS 

By  Various  Entomologists,  With  an  Introduction 
by  W.  L.  McAtee,  Biological  Survey 


WASHINGTON 

GOVERNMENT  PRINTING  OFFICE 

1923 


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V*H\f   \uy 

LETTER  OF  TRANSMITTAL. 


United  States  Department  of  Agriculture, 

Bureau  of  Biological  Survey, 
Washington,  D.  C,  May  8, 1922. 

Sir  :  I  have  the  honor  to  transmit  herewith,  recommending  that  it 
be  published  as  No.  46  in  the  series  of  the  North  American  Fauna, 
a  report  on  a  biological  survey  of  the  Pribilof  Islands,  Bering  Sea, 
Alaska,  by  Edward  A.  Preble,  assistant  biologist,  and  W.  L.  McAtee, 
in  charge  of  food  habits  research,  of  this  bureau,  assisted  by  a 
number  of  specialists  in  entomology.  This  report  treats  of  the  birds 
and  mammals  of  the  group,  together  with  the  insects  and  related 
invertebrates,  and  the  life-zone  relationships  of  the  region.  The 
Pribilof  Islands  constitute  a  national  reservation  and  are  especially 
noteworthy  as  being  the  sole  breeding  place  of  the  largest  herds  of 
fur  seals  in  the  world,  which  are  the  property  of  the  United  States 
Government  and  are  managed  by  the  Bureau  of  Fisheries  of  the 
Department  of  Commerce.  Numbers  of  valuable  blue  foxes,  which 
by  selective  breeding  under  normal  conditions  have  been  developed 
to  a  point  of  excellence  nowhere  else  attained,  are  also  found  on  the 
islands,  as  well  as  two  herds  of  reindeer.  The  present  report  will  be 
of  great  assistance  to  students  and  others  interested  in  the  natural 
history  of  Alaska. 

Respectfully,  E.  W.  Nelson, 

Chief  of  Bureau. 

Hon.  Henry  C.  Wallace, 

Secretary  of  Agriculture. 
n 


CONTENTS. 

Page. 

Part    I.  Birds  and  Mammals  of  the  Pribilof  Islands 1 

Introduction.    By  Edward  A.  Preble 1 

Life  zone  relationships.    By  Edward  A.  Preble 5 

Birds.    By  Edward  A.  Preble  and  W.  L.  McAtee 10 

Mammals.     By  Edward  A.  Preble 102 

Bibliography.    By  Edward  A.  Preble 121 

Part  II.  Insects,  Arachnids,  and  Chilopods  of  the  Pribilof  Islands 129 

Introduction.    By  W.  L.  McAtee 129 

Apterygota.    By  W.  L.  McAtee 139 

Orthoptera.    By  Morgan  Hebard 140 

Mallophaga.    By  G.  F.  Ferris 141 

Anoplura.    By  W.  L.  McAtee 142 

Homoptera.     By  Edith  M.  Patch 143 

Heteroptera.    By  W.  L.  McAtee 145 

Trichoptera.     By  Nathan  Banks 146 

Lepidoptera.    By  Wm.  T.  M.  Forbes 147 

Coleoptera.     By  H.  F.  Wickham 150 

Mecoptera.     By  Nathan  Banks 15S 

Diptera   (Tipulidae  and  Rhyphidae).     By  Charles  P.  Alex- 
ander   „ 159 

Diptera   (except  Tipulidae,  Rhyphidae,  and  Calliphoridae). 

By  J.  R.  Malloch 170 

Diptera  (Calliphoridae).    By  W.  R.  Walton 228 

Hymenoptera.     By  Henry  L.  Viereek 229 

Arachnida.     By  Nathan  Banks 237 

Chilopoda.    By  Ralph  V.  Chamberlin 240 

Index  to  Part  I 245 

m 


ILLUSTRATIONS. 


Page. 

Plate    I.  Harem  of  fur  seal  (Callorhinus  alascanus) Frontispiece 

II.  Map  of  Bering  Sea 2 

III.  Map  of  St.  Paul  Island 2 

IV.  Map  of  St.  George  Island 3 

V.  Topography  of  St.  Paul  Island 3 

VI.  Some  breeding  Pribilof  birds 20 

VII.  Pribilof  fur  seals  and  blue  fox 108 

VIII.  Details  of  plant  louse  (Macrosiphum  constrictum) 144 

IX.  Details  of  scorpion  fly  (Mecoptera)  and  mites  (Arachnida) 158 

X.  Details  of  crane  flies  (Tipulidae) 168 

XL  Details  of  crane  flies   (Tipulidae) 169 

XII.  Two  flies,  Coclopa  frigida  and  Pogonota  kincaidi  (Cyclorrhapha)  _  226 

XIII.  Details  of  flies  and  midges  (Nematocera  and  Brachycera) 226 

XIV.  Details  of  flies  (Cyclorrhapha) 227 

XV.  Details  of  flies  (Cyclorrhapha) 227 

IV 


No.  46.  NORTH  AMERICAN  FAUNA.  June  20,  1923 


A  BIOLOGICAL  SURVEY  OF  THE  PRIBILOF 
ISLANDS,  ALASKA. 


Part  I.  BIRDS  AND  MAMMALS. 

By  Edward  A.  Preble,  Assistant  Biologist,  Division  of  Biological  Investigations, 
and  W.  L.  McAtee,  In  Charge  Food  Habits  Research;  Bureau  of  Biological 
Survey. 


INTRODUCTION. 

By  Edward  A.  Preble. 

THE   ISLANDS. 

The  Pribilof  Islands  consist  of  five  small  islands  situated  in  Bering 
Sea,  in  latitude  57°  north  and  longitude  170°  west,  200  miles  from 
the  nearest  lands,  being  this  distance  north  of  the  Aleutian  Chain 
and  approximately  the  same  distance  south  of  St.  Matthew  Islands. 
The  largest  and  best  known  of  the  group  are  St.  Paul  and  St. 
George,  the  former,  about  13  miles  long,  with  a  coast  line  of  45 
miles;  and  the  latter,  12  miles  long,  with  a  coast  line  of  30  miles 
(Pis.  II,  III,  and  IV).  The  smaller  islands  are  all  close  to  St.  Paul — 
Walrus,  about  6  miles  to  the  east ;  Otter,  about  4  miles  to  the  south ; 
and  Sealion  Rock,  a  mere  rock  close  to  the  south  shore   (PI.  III). 

The  group  is  of  volcanic  origin  and  the  shore  lines  are  mainly 
rocky,  in  many  places  precipitous  (PI.  V).  On  St.  Paul  these  rough 
stretches  are  interrupted  by  sandy  beaches,  which  often  extend  for 
miles;  on  St.  George,  the  beaches  are  few  in  number,  and  not  ex- 
tensive. Back  from  the  shore  the  land  usually  rises,  gradually  slop- 
ing plateaus  being  surmounted  by  rocky  or  sandy  eminences,  in  some 
cases  the  craters  of  extinct  volcanoes.  These  hills  are  most  numer- 
ous on  St.  Paul,  where  two  attain  elevations  of  590  and  665  feet, 
respectively.  The  highest  hill  on  St.  George,  Ulekiah,  has  an  eleva- 
tion of  945  feet. 

The  Pribilofs  were  discovered  by  Russian  fur  traders  in  1786, 
while  searching  for  the  summer  home  of  the  hordes  of  fur  seals 
which  had  been  noted  pushing  northward  each  spring  past  the 
Aleutians.  The  islands  were  then  uninhabited,  but  were  at  once 
colonized  by  Aleutian  natives,  planted  there  by  the  Russians  to  col- 

1 


2  NORTH   AMERICAN   FAUNA.  [No.  46 

lect  the  skins  of  the  myriads  of  seals.  In  1799  the  islands  passed  into 
the  hands  of  the  Russian  American  Co.,  and  there  remained  until 
1867,  when  the  United  States  purchased  Alaska  from  Eussia. 

The  Pribilof  Islands  now  constitute  a  national  reservation.  In 
1870  St.  Paul  and  St.  George  Islands  were  made  in  effect  a  fur-seal 
reservation,  under  the  administration  of  the  Secretary  of  the 
Treasury.  The  remaining  islands,  Walrus  and  Otter,  were  set  aside 
by  Executive  order  as  the  Pribilof  Bird  Reservation  in  1909,  and  in 
1910  jurisdiction  over  the  entire  group  was  transferred  to  the  Secre- 
tary of  Commerce. 

The  three  larger  islands — St/Paul,  St.  George,  and  Otter — though 
supporting  no  tree  growth,  and  only  a  few  low  shrubs,  are  thickly 
clothed  with  herbaceous  vegetation,  the  moist  summers  inducing  a 
luxuriance  almost  tropical.  The  list  of  plants  includes  many  notable 
species,  which,  because  of  the  peculiar  summer  climate,  have  a  very 
long  season  of  flowering.  In  consequence,  large  fields  of  arctic  pop- 
pies, lupines,  saxifrages,  and  other  flowers  of  great  beauty  are  found 
throughout  the  summer.  Certain  annuals  attain  so  rank  a  growth 
as  to  add  seriously  to  the  labor  of  foot  travel  over  the  rougher  areas. 

SCIENTIFIC   OBSERVATIONS   AND    COLLECTIONS. 

The  teeming  hordes  of  fur  seals  and  the  swarms  of  breeding  sea 
birds  seem  to  have  interested  the  early  inhabitants  merely  in  a 
utilitarian  way.  It  was  not  until  1840  that  any  work  enumerating 
even  the  more  conspicuous  species  of  birds  and  mammals  appeared. 
(See  Veniaminof,  1840,  in  Bibliography.)  William  Palmer  (1899, 
p.  360)  summed  up  the  history  of  the  ornithological  collections  and 
observations  made  on  the  Pribilofs  up  to  1899,  as  follows : 

Coinde  in  1860  made  a  list  of  but  nine  species  that  were  collected  by  an 
officer  of  the  Russian  Navy,  Mr.  Warneck,  in  1852.  The  jealous  care  exercised 
by  the  Russians  for  the  preservation  of  the  seals  prevented  any  outsiders  from 
studying  the  avifauna,  so  that  it  was  not  until  the  islands  passed  under  the 
control  of  this  Government,  and  Mr.  Elliott  made  his  investigations,  that  any 
great  effort  was  made  to  study  the  bird  life.  Dr.  W.  H.  Dall  had  indeed  in 
1S68  spent  a  short  time  at  the  islands  and,  together  with  the  officers  of  his 
surveying  vessel,  had  collected  some  specimens,  one  of  which  was  described  as 
new  by  Prof.  S.  F.  Baird,  but  the  first  systematic  investigation  was  made  by 
Mr.  Henry  W.  Elliott  [in  1872  and  1873].  The  result  of  this  gentleman's  work 
was  a  list  of  40  species,  based  on  copious  notes  and  numerous  specimens  which 
were  named  and  elaborated  by  Dr.  Elliott  Coues  in  Mr.  Elliott's  report  for 
1873  [=1874],  and  which  was  reprinted  in  1875.  Mr.  Elliott  made  another 
more  extensive  elaboration  in  his  monograph  of  the  seal  islands  in  1882.  Be- 
sides the  above,  various  other  Government  expeditions  which  have  visited  the 
waters  of  Bering  Sea  for  different  reasons  during  the  past  16  years  have 
generally  touched  at  the  seal  islands  and  given  several  naturalists  opportu- 
nities for  collecting  and  noting  the  bird  life.  Thus,  Mr.  L.  M.  Turner  in  1878, 
Dr.  T.  H.  Bean  in  1S80,  Mr.  E.  W.  Nelson  in  1881,  Lieut.  J.  E.  Lutz  in  18S4, 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey.  PLATE   I  I. 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey.  PLATE    III. 


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North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  V. 


Fig.   I. — Winter  on  St.  Paul  Island. 

View  looking  toward  head  of  lagoon  from  beach  near  Village  Landing.    Photograph  by  G 

Hanna. 


Pallas 


Fig.  2. — Black  Bluffs,  Near  Village,  St.  Paul  Island. 

This  is  the  most  notable  of  the  exposures  of  fossiliferous  rock  on  the  Pribilofs.    Photograph  by 

A.  ChristofFersen. 


Fig.  3. — Surf-Lashed  Shore,  St.  Paul  Island. 

In  these  seething  waters  the  young  seals  play  for  hours.    Photograph  by  A.  Christoffersen. 


1923.]  PRIBILOF   ISLANDS   SURVEY,   INTRODUCTION.  3 

Mr.  C.  H.  Townsend  in  1885  to  1896,  myself  in  1890,  Messrs.  F.  W.  True  and 
D.  W.  Prentiss,  jr.,  in  1895,  Mr.  F.  A.  Lucas  in  1896-97  have  spent  from  a  day 
or  two  to  several  months  on  the  islands. 

Palmer  spent  the  period  from  May  27  to  August  11,  1890,  on  the 
Pribilofs,  mainly  on  St.  Paul.  His  time  was  spent  largely  in  the 
study  of  the  birds,  but  owing  to  regulations  then  in  vogue  regarding 
the  use  of  firearms  his  opportunities  for  collecting  specimens  were 
somewhat  restricted.  His  report,  which  is  a  painstaking  and 
scholarly  work,  has  remained  the  only  completely  annotated  list  of 
the  birds  of  the  group  up  to  the  present  time. 

In  1910,  when  the  Pribilof  Islands  were  placed  in  charge  of  the 
Bureau  of  Fisheries  of  the  Department  of  Commerce,  in  furtherance 
of  the  policy  to  make  a  thorough  study  of  the  life  of  the  islands,  the 
breeding  place  of  the  fur  seal,  Dr.  Walter  L.  Hahn  was  appointed 
naturalist.  Evermann  (1913,  p.  15)  thus  refers  to  his  work,  and 
that  of  his  successor : 

Dr.  Walter  L  Hahn,  at  that  time  head  of  the  department  of  biology  in  the 
State  normal  school  at  Springfield,  S.  Dak.,  was  appointed  naturalist  in  the  sum- 
mer of  1910.  He  arrived  at  St.  Paul  Island  August  24  and  immediately  entered 
upon  his  duties  with  an  energy  and  intelligence  which  could  scarcely  be  excelled. 
His  untimely  death  on  May  31,  1911,  from  exposure  in  the  ice-cold  water  of 
the  village  lagoon,  resulting  from  the  capsizing  of  a  boat,  was  a  severe  loss 
to  the  fur-seal  service  and  to  biological  science.  During  his  few  months  on 
St  Paul  Island,  Doctor  Hahn,  from  the  voluminous  notes  and  records  which 
he  left,  appears  to  have  been  indefatigable  in  his  field  work  and  marvelously 
painstaking  in  recording  his  observations.  His  notes,  typewritten  up  to  noon 
of  the  very  day  of  his  death,  record  a  vast  number  of  new  and  important 
observations  on  the  fur  seals  and  blue  foxes.  They  also  teem  with  records  of 
interesting  observations  on  the  birds  and  other  animals  of  the  islands. 

Mr.  Millard  C.  Marsh,  pathologist  of  the  Bureau  of  Fisheries,  was  appointed 
to  the  vacancy  caused  by  the  death  of  Doctor  Hahn.  He  reached  St.  Paul 
Island  August  23,  1911,  and  took  up  and  continued  with  commendable  industry 
and  intelligence  the  work  so  ably  begun  by  his  predecessor. 

As  a  result  of  this  work  Evermann  (1.  c.)  was  able  to  add  18  spe- 
cies of  birds  to  the  Pribilof  list ;  four  of  these  were  for  the  first  time 
added  to  the  North  American  list.  In  addition,  the  present  writer 
has  been  able,  through  the  courtesy  of  the  Bureau  of  Fisheries,  to  use 
the  notes  on  occurrence  of  many  species  recorded  by  Hahn  and 
Marsh.  Copies  of  the  notes  in  the  St.  Paul  Island  log  relating  to 
many  species  of  birds  and  mammals,  carefully  transcribed  by  Hahn, 
and  arranged  under  specific  headings,  have  also  been  available. 

Alvin  G.  Whitney,  school-teacher  on  St.  Paul  Island  from  the  sum- 
mer of  1912  to  the  summer  of  1914,  made  considerable  collections  of 
insects  and  shells  and  some  other  invertebrates,  but  his  contributions 
to  ornithology  and  mammalogy  were  small. 

The  next  naturalist  to  work  on  the  Pribilofs  was  G.  Dallas  Hanna. 
He  arrived  on  St.  Paul  in  July,  1913,  and  was  transferred  during  the 
next  month  to  St  George.     Although  his  regular  duties  as  school- 


4  NORTH  AMERICAN   FAUNA.  [No.  46 

teacher  and  volunteer  radio  operator  involved  long  hours,  he  found 
time  to  make  notes  and  collections.  In  early  August  of  1914  he 
was  transferred  to  St.  Paul,  where  he  remained  (with  the  exception 
of  two  trips  to  St.  George  in  the  summer  of  1915)  until  September, 

1915,  when  he  returned  to  the  United  States.  During  all  this  period 
he  devoted  most  of  his  spare  time  to  ornithology.  In  1916,  mainly 
as  a  result  of  his  own  work,  he  added  13  species  to  the  Pribilof  list 
in  a  paper  published  in  The  Auk. 

In  the  meantime  the  visit  of  the  writer  and  his  colleagues,  Wilfred 
H.  Osgood,  of  the  Field  Museum,  Chicago,  111.,  and  Prof.  George  H. 
Parker,  of  Harvard  Universit/,  Cambridge,  Mass.,  on  a  special  in- 
vestigation of  the  fur-seal  herd  in  the  summer  of  1914,  afforded  an 
opportunity  for  further  study  of  the  fauna.  It  is  needless  to  state 
that  the  extremely  interesting  wealth  of  bird  life  on  these  islands, 
then  for  the  first  time  observed  by  the  writer,  determined  him  to 
prepare  a  fully  annotated  list  of  the  birds,  and  this  forms  a  part  of 
the  present  report. 

Mr.  Hanna  returned  to  St.  Paul  in  June,  1916,  and  remained  there 
(excepting  short  visits  to  St.  George  each  summer  and  a  residence 
there  from  October,  1916,  to  May,  1917)  until  September,  1918.  As 
a  result  of  the  work  done  there,  he  was  able  to  publish  a  paper 
adding  21  species  of  birds  to  the  Pribilof  list,  4  of  which  were  new 
to  North  America.  He  later  spent  two  summer  seasons  on  the 
islands  (June  to  September,  1919  and  1920)  on  seal  work,  and  each 
season  made  additions  to  the  list.  In  1919  he  published  a  list  of  all 
the  birds  known  at  that  time  to  have  occurred  on  the  Pribilofs,  129 
in  number,  in  which  the  various  species  are  classed  as  breeders,  regu- 
lar migrants,  or  transients,  and  the  author  first  reporting  the  species 
from  the  group  is  indicated.  In  addition,  he  has  published  other 
papers  of  a  general  nature  or  relating  to  certain  species,  and  he  has 
very  generously  furnished  the  writer  with  numberless  unpublished 
notes  on  the  occurrence  of  birds  and  mammals,  extending  practically 
throughout  his  period  of  service  on  the  islands.  Without  his 
records  and  those  of  Hahn,  above  referred  to,  the  preparation  of 
the  present  lists  would  scarcely  be  justified.  Notes  on  a  number  of 
species  of  birds  made  by  H.  P.  Adams  on  St.  George  Island  in  1915, 

1916,  and  1917,  have  also  been  available.  All  such  notes,  when  lack- 
ing citations,  are  understood  to  be  hitherto  unpublished. 

The  foregoing  account  has  been  written  mainly  from  the  stand- 
point of  the  ornithologist;  since  the  birds  are  so  abundant  and  so 
far  outnumber  the  other  vertebrates  in  species  and  individuals,  it  is 
natural  that  they  should  have  received  the  greater  share  of  atten- 
tion. Practically  all  workers,  however,  have  done  some  work  on  the 
mammals.  In  the  bibliography  (pp.  121-128),  therefore,  the  titles 
are  merged  in  one  list;  indeed,  many  articles  refer  to  both  groups. 


LIFE  ZONE  RELATIONSHIPS. 

By  Edwaed  A.  Pbeble. 
While  the  Pribilof  group  as  regards  its  zoogeography  is  most 
closely  related  to  the  Arctic,  there  are  many  elements  that  forbid 
its  unqualified  reference  to  that  zone.  The  small  size  of  the  islands, 
their  distance  from  other  lands,  and  probably  to  some  extent  the 
ocean  currents  have  all  been  factors  which  helped  to  determine  its 
present  faunal  and  floral  characteristics.  The  severe  winds  which 
sweep  the  islands  have  also  had  their  influence.  Other  important 
contributing  causes  have  been  the  original  freedom  of  the  islands 
from  human  habitation,  and  their  rugged  topography,  combined  with 
an  abundant  food  supply,  which  have  favored  their  selection  as 
breeding  stations  for  myriads  of  aquatic  birds  and  a  number  of 
species  of  mammals.  All  these  factors  have  combined  to  produce  a 
fauna  and  flora  peculiar  in  many  respects. 

MAMMALS. 

Among  land  mammals  we  find  only  three  species,  all  more  or  less 
peculiar  to  the  group.  Of  these  the  foxes  (Alopex  pribilofenxis) 
were  probably  originally  brought  to  the  islands  by  floating  ice  and 
have  become  slightly  differentiated  from  the  original  mainland 
stock.  The  shrew  (Sorex  pribilof ensis)  and  the  lemming  (Lemmas 
nig?%ipes):  peculiar  to  St.  Paul  and  St.  George  Islands,  respectively, 
have  been  so  modified  that  their  origin  can  not  be  determined.  The 
fox  and  lemming,  however,  and  perhaps  the  shrew,  belong  to  groups 
which  are  essentially  Arctic  in  distribution.  All  the  other  mammals, 
excepting  those  introduced  by  man,  are  aquatic  and  can  not  be  dis- 
cussed in  terms  applicable  to  the  distribution  of  the  others.  The 
boundaries  of  their  habitats  are  probably  determined  by  the  tem- 
perature of  the  water,  acting  to  some  extent  directly,  but  more 
especially,  through  their  food  supply.  Of  the  strictly  aquatic 
mammals,  the  bowhead  whale  (Balaena  mysticetus) ,  if  correctly 
identified,  is  near  the  southern  limit  of  its  range ;  all  the  others,  with 
the  exception  of  the  killer  whale  (Orcinus  rectipinna),  which  visits 
the  Arctic  Ocean  in  summer,  are  near  their  northern  limit. 

In  regard  to  the  pinnipeds,  less  strictly  aquatic,  the  Pribilofs  are 
the  meeting  point  of  both  southern  and  northern  types.  The  Steller 
sea  lion  (Kumetopias  jubata),  Pribilof  fur  seal  (Callorhinus  alasca- 
nus),  and  Pribilof  harbor  seal  (Phoca  richardii  pribilof  ensis)  are 
at  or  near  their  northern  limit;  while  the  walrus  (Odobenus 
divergens),  ribbon  seal  (Phoca  fasciata),  and  bearded  seal  (Erigna- 
thus  barbatus  nauticus)    there  approach  their  southern  boundary. 

5 


NORTH   AMERICAN"   FAUNA. 


[No.  46 


The  sea  otter  (Latax  lutris  lutris),  now  exterminated  there,  found 
about  the  Pribilofs  one  of  its  northernmost  stations. 


BIRDS. 


In  discussing  the  zonal  relation  of  the  group  as  reflected  in  its  bird 
life  it  is  advisable  to  deal  only  with  species  which  regularly  breed. 
These  are  as  follows: 


Fulmarus  glacialis  rodgersi. 
Phalacrocorax  pelagians  robustus. 
Phalacrocorax  urile. 
Clangula  hyemalis. 
Lobipes  lobatus. 

Arquatella  maritime/,  ptilocnemis. 
Leucosticte  griseonucha. 
Plectrophenax  nivalis  townsendi. 
Calcarius  lapponicus  alascensis. 
Nannus  troglodytes  alascensis. 


Lunda  cin'hata. 

Fratercula  corniculata, 

Phaleris  psittacula. 

Aethia  cristateUa. 

Aethia  pusilla. 

Vria  troille  calif ornica. 

Uria  lomvia  arra. 

Rissa  tridactyla  pollicaris. 

Rissa  brevirostris. 

Larus  hyperboreus  hyperboreus. 

Larus  glaucescens. 

Sixteen  of  these  (excepting  Rissa  brevirostris,  Larus  glaucescens, 
Phalacrocorax  urile,  and  Nannus  t.  alascensis)  breed  commonly  on 
the  St.  Matthew  group  to  the  northward  (a  number  of  them  still 
farther  north,  to  and  beyond  Bering  Strait),  and  one  other,  Plectro- 
phenax nivalis  townsendi,  is  represented  there  by  a  closely  related 
species,  P.  hyperboreus.  Rissa  brevirostris  reaches  its  northern  limit 
on  the  Pribilofs,  and  Larus  glaucescens  in  the  northern  part  of 
Bering  Sea.  Phalacrocorax  urile  is  a  summer  visitor  from  the  Siber- 
ian coast,  and  Nannus  t.  alascensis  is  peculiar,  but  has  close  relatives 
on  the  Aleutian  Islands.  It  will  be  noted  that  these  are  mainly  sea 
birds,  some  of  which  nest  as  far  south  as  California,  but  which,  find- 
ing abundant  food  combined  with  convenient  nesting  sites  on  these 
northern  islands,  breed  successfully  there  also  regardless  of  the  lower 
temperature.  These,  like  the  aquatic  mammals,  can  not  be  regarded 
as  coordinate  in  value  with  land  vertebrates  as  regards  zonal  signifi- 
cance. Clangula  hyemalis  and  Lobipes  lobatus,  which  nest  about 
fresh-water  ponds,  and  Arquatella  m.  ptilocnemis,  Leucosticte  griseo- 
nucha, Plectrophenax  n.  townsendi,  and  Calcarius  I.  alascensis,  are 
distinctly  Arctic  types. 

It  will  thus  be  seen  that  both  the  mammalian  and  avian  inhabitants 
of  the  Pribilofs,  excluding  those  which  can  not  legitimately  be  con- 
sidered as  furnishing  criteria  for  the  definition  of  life  zones  as  cur- 
rently considered,  favor  the  reference  of  this  group  to  some  division 
of  the  Arctic  Zone. 

But  while  the  aquatic  species  of  mammals  and  birds  must  be  con- 
sidered from  a  standpoint  different  from  that  accorded  those  in- 
habiting the  land,  they  can  hardly  be  ignored  in  the  consideration 
of  a  fauna  of  which  they  constitute  an  overwhelming  majority. 
Palmer,  indeed,  our  principal  authority  on  Pribilof  birds,  considered 


1923.]  LIFE   ZONES   OF   THE   PRIBILOF   ISLANDS.  7 

the  breeding  water-birds  as  mainly  characterizing  a  zoogeographic 
area  to  which  he  accorded  the  rank  of  a  subregion  of  the  Holarctic — 
the  Aleutican.  This  is  practically  the  same  as  the  Aleutian  Faunal 
District  of  Nelson,  extended  to  include  the  Pribilofs  and  other  islands 
and  the  immediate  coasts  of  Alaska  and  Siberia  north  to  Bering 
Strait,  and  includes  the  insular  and  coastal  portions  of  the  Alaskan 
Province  of  the  Holarctic  region  of  Newton.  (Dictionary  of  Birds, 
Part  II,  p.  331,  1893.) 

PLANTS. 

The  botany  of  the  Pribilofs  has  been  carefully  studied  by  James 
M.  Macoun,  who  has  published  an  annotated  list  of  184  species,  in- 
cluding the  vascular  cryptogams.  His  paper  includes  a  table,  pre- 
pared in  collaboration  with  Theodor  Holm,  one  of  our  most  eminent 
students  of  Arctic  plants,  which  shows  the  distribution  of  the  va- 
rious species  in  other  parts  of  the  Arctic  region.  Regarding  the 
phaenogams  only,  172  species,  we  find  that  141  species  are  found 
on  the  east  coast  of  Bering  Sea,  and  126  on  the  west  coast,  while  101 
grow  on  both  coasts ;  74  are  found  in  Arctic  America.  Twenty-three 
are  not  known  from  either  the  east  or  west  coasts  of  Bering  Sea,  and 
of  these  23  only  6  are  found  in  Arctic  America ;  of  the  remaining  17 
the  majority  are  species  which  have  been  described  from  the  Pribi- 
lofs, while  the  remainder  are  presumably  found  also  on  the  Aleu- 
tians.1 It  will  thus  be  seen,  as  might  be  expected,  that  the  great 
majority  of  the  Pribilof  plants  are  arctic  or  subarctic  species. 

The  phaenogams  classed  as  Arctic  by  Macoun  and  Holm  are  the 
following : 


Ranunculus  tricophyllus  Chaix. 
Ranunculus  hyperboreus  Rottb. 
Ranunculus  pygmacus  Wahl. 
Ranunculus  reptans  L. 
Ranunculus  eschscholtsii  SchL 
Coptis  trifolia  Salisb. 
Papaver  radicatum  Rottb. 
Draba  hirta  L. 
Draba  wahlenbergii  Hartm. 
Eutrema  edwardsii  R.  Br. 
Cochlearia  officinalis  L. 
Cardamine  bellidifolia  L. 
Cardamine  pratensis  L. 
Silene  acaulis  L. 
Lychnis  apetala  L. 


Arenaria  peploides  L. 

Stellaria  liumifusa  Rottb. 

Stellaria  longipes   Goldie   var.    laeta 

Was. 
Cerastium  alpinum  L. 
Sagina  linnaci  Presl. 
Sagina  nivalis  Fr. 
Rubus  chamaemorus  L. 
Rubus  arcticus  L. 
Potentilla  anserina  L. 
Comarum  palustre  L. 
Potentilla  emarginata  Pursh. 
Saxifraga  hieracifolia  Waldst.  and  Kit. 
Saxifraga  stellaris  L.  var.  comosa  Poir. 
Saxifraga  hirculus  L. 


1  The  late  James  M.  Macoun,  In  1914,  informed  me  that  a  few  species  of  Pribilof  plants, 
evidently  of  Aleutian  origin,  were  known  by  him  to  occur  only  near  Southwest  Point,  St. 
Paul  Island.  Since  Mr.  Macoun  on  that  occasion  added  several  species  to  the  list,  which  I 
do  not  now  remember,  and  which  are  probably  yet  unpublished,  the  particular  ones  re- 
ferred to  by  him  can  not  be  determined.  A  direct  comparison  of  lists  of  the  Pribilof  and 
Aleutian  species  would  be  desirable  in  the  present  connection,  but  I  have  found  no  work 
giving  a  complete  list  of  the  plants  of  the  latter  region. 


8 


NORTH   AMERICAN   FAUNA. 


[No.  46 


Chrysosptenium  alternifolium  L. 
Parnassia  kotzebuei  Cham,  and  Schl. 
Hippuris  vulgaris  L. 
Epilobium  spicatum  Lain. 
Selinum  benthami  Wat. 
Valeriana  capitata  Pall. 
Achillea  millefolium  L. 
Chrysanthemum  arcticum  L. 
Petasites  frigida  Fries. 
Campanula  uniflora  L. 
Pyrola  minor  L. 
Armeria  vulgaris  Willd. 
Veronica  serpyllifoUa  L. 
Pedicularis  sudetica  Willd. 
Ped-lcularis  lanata  Willd. 
Euphrasia  officinalis  L. 
Gynandra  stelleri  Cham,  and  Schl. 
Koenigia  islandica  L. 
Polygonum  viviparum  L. 
Polygonum  bistorta  L. 
Oxyria  reniformis  Hook. 
Rumex  acetosella  L. 
Salix  arctica  Pall. 


Salix  reticulata  L. 
Empetrum  nigrum  L. 
Lloydia  serotina  Reich. 
Streptopus  amplexifolius  DC. 
J  uncus  biglumis  L. 
Eriophorum  polystachyon  L. 
Eriophorum  vaginatum  L. 
Carex  vulgaris  Fries,  and  vara. 
Carex  membranopacta  Bailey. 
Carex  rariflora  Smith. 
Carex  saxatilis  L. 
Hierochloa  pauciflora  R.  Br. 
Alopecurus  alpinus  Smith. 
Arclogrostis  latifolia  Grisb. 
Calamagrostis  purpurascens  V. 
Trisetum  subspicatum  Beauv. 
Phippsia  algida  R.  Br. 
Arctophila  effusa  Lange. 
Dupontia  psilosantha  Rupr. 
Glyceria  angustata  Fries. 
Festuca  ovina  L. 
Elymus  mollis  Trin. 


The  following  species  of  plants  have  been  described  as  new  from 
Pribilof  specimens  (some  of  these  are  not  currently  considered 
valid) : 


Papaver  macounii  Greene. 
Nesodraba  grandis  Greene. 
Cardamine  umbellata  Greene. 
Crysosplenium  beringianum  Rose. 
Primula  macounii  Greene. 
Primula  eximia  Greene. 
Polygonum  macounii  Small. 


Salix  rotundata  Rydberg. 
Salix  cyclophylla  Rydberg. 
Carex  pribylovensis  Macoun. 
Alopecurus   howellii   Vasey   var. 

riami  Beal. 
Elymus  villosissimus  Scribner. 
Calamogrostis  arctica  Vasey. 


mer- 


ZOOGEOGRAPHT  OF  NEIGHBORING  AREAS. 

It  is  scarcely  possible  to  discuss  the  zoogeographic  aspects  of  the 
Pribilofs  without  considering  also  the  neighboring  Aleutian  Peninsula 
and  Islands.  The  latest  treatment  seems  to  be  that  of  Osgood,2  who 
has  discussed  the  zoogeography  of  the  Alaska  Peninsula  at  some 
length,  and  while  assigning  the  treeless  part  of  that  region  to  the 
unqualified  Arctic  Zone  admits  the  possibility  that  a  more  extended 
study  may  show  the  terminal  part  of  the  peninsula  and  the  adjoin- 
ing islands  to  belong  to  a  subdivision  of  the  Arctic.  The  writer  be- 
lieves that  recognition  of  such  a  subdivision  is  desirable,  and  would 
retain  the  name  first  applied  to  it  by  Nelson,  the  Aleutian  Faunal 
District,3  considered  as  a  subdivision  of  the  Arctic  Zone,  and  includ- 

3  A  Biological  Reconnaissance  of  the  Base  of  the  Alaska  Peninsula :  North  American 
Fauna  No.  24,  pp.  21-25,  1904.  I  have  also  had  the  benefit  of  suggestions  from  Dr. 
Alexander  Wetroore,  who  spent  the  summer  of  1911  on  the  Aleutian  Islands  and  the 
Peninsula. 

s  Report  on  Nat  Hist.  Collections  made  in  Alaska,  p.  26,  1887. 


1923.] 


LIFE   ZONES   OF   THE   PRIBILOF   ISLANDS. 


ing  the  western  end  of  the  Alaska  Peninsula,  the  Aleutian  Islands, 
and  the  Pribilofs.  The  St.  Matthew  group  and  perhaps  other 
islands  in  Bering  Sea  might  also  best  be  included  here. 

It  is  evident,  however,  that  this  subdivision,  and  especially  the 
Alaska  Peninsula,  by  no  means  shares  its  characteristic  fauna  and 
flora  exclusively  or  even  principally  with  the  Arctic  proper.  In  the 
first  place  it  should  be  borne  in  mind  that  although  the  summer  tem- 
peratures alone  on  the  Pribilofs  and  at  Unalaska  are  sufficiently  low 
to  limit  the  growth  of  trees,  absence  of  trees  over  large  areas  on  the 
peninsula  is  evidently  due  in  part  to  some  other  factor.  There  is 
little  doubt  that  this  contributing  cause,  as  suggested  by  Osgood,  is 
the  severe  wind  which  sweeps  the  region  at  all  seasons.  This,  how- 
ever, has  not  sufficed  to  prevent  a  luxuriant  growth  of  alders  and 
willows  in  the  more  sheltered  parts  of  the  peninsula  beyond  the  line 
of  tree  growth,  and  these  afford  breeding  sites  for  the  following 
birds,  which  over  most  of  their  ranges  are  not  considered  to  nest 
above  the  Hudsonian  Zone: 


Dendroica  aestiva  rubiginosa. 
Wilsonia  pusilla  pileolata. 
Hylocichla  ustulata  swainsoni. 
Hylocichla  guttata  guttata. 


Pica  pica  hudsonia. 
Zonotrichia  coronata. 
Passerella  iliaca  unalaschensis. 
Melospiza  melodia  insignis. 
Riparia  riparia. 

The  dipper,  or  water  ousel  (Cinclus  mexicanus  miticolor),  a  bird 
nesting  in  the  Canadian  and  Hudsonian  zones,  also  breeds  in  this 
area. 

Opposed  to  this  list  is  the  following,  including  birds  which  may 
be  considered  truly  Arctic  in  their  affinities,  and  most  of  which  nest 
largely  on  the  higher  grounds : 


Plectrophenax  nivalis  toionsendu 
AcantMs  homemanni  exilipes. 
Anthus  spinoletta  rubescens. 


Lagopus  lagopus  albus. 
Lagopus  rupestris  nelsoni. 
Calcarius  lapponicus  alascensis. 
Leucosticte  grisconucha. 

The  remainder  of  the  breeding  land  birds  of  the  Alaska  Peninsula, 
all  wide-ranging  forms,  may  be  classed  as  follows : 

Archibuteo  lagopus     sancti-johannis. 

Breeding  in  Hudsonian  and  Arctic 

Zones. 

Haliaeetus  leuoocephalus     alascanus. 

Breeding  from   Canadian  to  Arctic 

Zones. 


Falco  peregrinus  pealei.  Breeding 
from  Transition  to  Arctic  Zones. 

Corvus  corax  principalis.  Breeding 
from  Canadian  to  Arctic  Zones. 

Passerculus  sandwichensis  sandwich- 
ensis.  Breeding  regularly  to  Hud- 
sonian Zone. 


It  seems  evident,  therefore,  that  while  it  may  be  desirable  to  in- 
clude the  terminal  part  of  the  Alaska  Peninsula  in  the  Aleutian 
Faunal  District,  a  majority  of  its  breeding  land  birds  belong  at  least 
in  part  to  zones  below  the  Arctic. 


BIRDS  OF  THE  PRIBELOF  ISLANDS,  ALASKA. 
By  Edwabd  A.  Pbeble  and  W.  L.  McAtee. 

In  the  annotated  list  of  birds  of  the  Pribilofs,  the  portions  of  the 
accounts  relating  to  distribution,  migration,  nesting,  and  general  habits 
are  by  Edward  A.  Preble.  The  attempt  has  been  made  to  collect  all 
available  data  referring  to  the  various  species,  with  special  reference 
to  dates  of  arrival,  nesting,  departure,  and  occurrence  in  winter,  and 
in  most  cases  the  notes  are  presented  in  the  order  here  indicated.  The 
circumstances  under  which  the  observations  were  made  have  been  de- 
scribed in  detail  in  the  general  introduction,  and  need  not  be  here  re- 
peated. 

BREEDING  BIRDS. 

Of  the  137  species  now  known  to  have  occurred  on  the  Pribilofs, 
only  23  are  known  to  breed ;  of  these  Anas  platyrhyncha  platyrhyn- 
cha and  Nettion  carolinense  are  rare.    The  list  follows : 


Lunda   cirrhata. 

Fratercula  comiculata. 

Phaleris  psittacula. 

Aethia  cristatella. 

Aethia  pusilla. 

JJria  troille  californica. 

Uria   lomvia   arra. 

Rissa  tridactyla  pollicaris. 

Rissa  brevvrostris. 

Larus  hyperboreus  hyperboreus. 

Larus  glaucescens. 

Fulmarus  glacialis  rodgersi. 


Phalacrocorax  pelugicus  robustus. 

Phalaerocorax  urile. 

Anas  platyrhyncha  platyrhyncha. 

Nettion  carolinense. 

Clangula   hyemalis. 

Lobipes   lobatus. 

Arquatella  maritima  ptilocnemis. 

Leucosticte   griseonucha. 

Plectrophenax  nivalis  totnnsendi. 

Calcarius  lapponicus  alascensis. 

Nannus  troglodytes  aluscensis. 


In  addition,  HistHonicus  Mstrionicus  pacificus  is  present  in  abun- 
dance all  summer,  but  has  not  been  found  actually  nesting.  Cepphus 
columba  and  Larus  schistisagus  also  have  been  stated  to  breed,  but  on 
insufficient  evidence. 


REGULAR  MIGRANTS. 


Several  of  these,  even  some  which  breed  abundantly,  occur  in  great- 
er numbers  during  migration.  Examples  are  Larus  hyperboreus  hy- 
perboreus, Anas  platyrhyncha  platyrhyncha,  Histrionicus  Mstrioni- 
cus padficus  and  Arquatella  maritima  ptilocnemis.  Sixteen  others 
are  known  to  be  more  or  less  regular  migrants;  these  are  the  fol- 
lowing : 
10 


1923.] 


BIRDS  OF   THE  PRIBILOF  ISLANDS. 


11 


Xema  sabinii.    Spring  and  fall. 
Sterna  paradisaea.    Spring  and  fall. 
Daflla  acuta  tzitzihoa.     Spring  and  fall. 
Somateria  v-nigra.    Mainly  in  winter. 
Somateria     spectabilis.       Mainly     in 

winter. 
Oidemia  deglandi  dixoni.     Mainly   in 

winter. 
Branta    canadensis    minima.      Spring 

and  fall. 
Philacte  canagica.     Spring  and  fall. 
Phalaropus    fulicarius.      Spring    and 

fall. 


Pisobia  acuminata.     Mainly  in  fall. 
Pisobia  maculata.    Mainly  in  fall. 
Limosa  lapponica  baueri.    Spring  and 

fall. 
Heteroscehis  incanus  incanus.     Spring 

and  fall. 
Pluvialis  dominica  fulva.     Spring  and 

fall. 
Arenaria  interpres  interpres.     Mainly 

in  fall. 
Anthus  spinoletta  rubescens.     Mainly 

in   fall. 


OCCASIONAL  VISITORS. 

All  the  remaining  species  of  the  list  are  transients  which  have  been 
detected  once  or  in  some  cases  a  number  of  times.  A  considerable 
number  of  these,  as  more  observations  are  afforded,  will  probably 
be  transferred  to  the  list  of  regular  migrants.  In  addition  to  those 
migrants  which  occur  mainly  as  winter  visitors  from  farther  north, 
the  following,  of  less  regular  occurrence,  have  also  been  noted  chiefly 
in  winter. 


Colymbus  auritus. 
Cepphus  columba. 
Polysticta  stelleri. 


Arquatella  maritima  couesi. 
Uierofalco  rusticolus  sacer. 
Nyctea  nyctea. 


VISITORS   FROM   EURASIA. 

Those  members  of  the  avifauna  which  may  fairly  be  classed  as 
transients  or  accidental  visitors  number  over  80  species.  Of  these 
the  following  are  Eurasian,  with  general  ranges  as  given;  unless 
otherwise  indicated  specimens  taken  on  the  Pribilofs  have  proved 
to  be  the  first  records  for  North  America : 

Mareca  penelope.  Breeds  in  northern  Eurasia  from  Scandinavia  to  Kamchatka, 
wintering  to  the  southward. 

Eunetta  falcata.  Occurs  regularly  in  eastern  Asia  to  Kamchatka,  Korea,  and 
Japan. 

Nettion  crecca.  Of  general  distribution  in  northern  Eurasia,  breeding  easterly 
to  the  eastern  Aleutian  Islands,  where  it  was  recorded  previous  to  its  capture 
on  the  Pribilofs. 

Fuligula  fuligula.  Breeding  in  Eurasia  eastward  to  Kamchatka,  migrating 
in  winter  to  northern  Africa,  India,  China,  and  Japan. 

Aithyia  ferina.  Breeding  in  Eurasia  east  to  Lake  Baikal,  wandering  in  migra- 
tion and  winter  eastward  to  China  and  Japan. 

Glaucionetta  clangula  clangula.  Breeding  generally  throughout  northern  Eu- 
rasia. 

Limnocryptes  gallinula.  Breeds  in  Eurasia  east  to  the  Kolyma  River.  In  mi- 
gration and  winter  occurs  south  to  India  and  Burma,  and  occasionally  to 
Japan  and  Formosa. 


12  NORTH   AMERICAN   FAUNA.  [No.  46 

Pisobia  subminuta.  Breeds  in  eastern  Siberia,  Kamchatka,  and  the  Commander 
and  Kurile  Islands.  Migrates  in  winter  as  far  as  Australia  and  the  Philip- 
pines. 

Rhyacophilus  glareola.  Breeds  in  northern  Eurasia  eastward  to  Kamchatka 
and  Bering  Island,  wintering  south  to  Africa,  southern  Asia,  Japan,  and  the 
Philippine  Islands.  Recorded  previously  from  Alaska  (Littlejohn,  The  Con- 
dor, vol.  6,  p.  138,  1904,  Sanak  Island). 

Heteroscelus  incanus  brevipes.  Breeds  in  eastern  Siberia  and  Kamchatka. 
Migrates  southwardly  to  China,  Japan,  the  Malay  Peninsula,  and  Australia. 
Taken  at  least  once  on  Bering  Island  in  May. 

Philomuchus  pugnax.  Occurs  in  central  Europe  and  Asia,  eastward  rarely  to 
Japan  and  China,  wintering  in  Africa  and  India.  First  record  for  western 
coast  of  North  America,  but  has  been  taken  many  times  in  the  eastern  United 
States. 

Numenius  tahitiensis.  Occurs  on  many  of  the  Pacific  islands,  mainly  in  winter ; 
has  been  taken  several  times  in  summer  on  the  mainland  of  Alaska,  once  at 
least  previous  to  its  capture  on  the  Pribilofs. 

Thallasoaetus  pelagicus.  Breeds  in  Kamchatka,  and  has  been  taken  on  Bering 
Island. 

Cryptoglaux  funerea  magna.  Described  from  the  Kolyma  River;  probably  of 
general  range  in  northeastern  Siberia. 

Cuculus  canorus  telephonus.  Breeds  in  Kamchatka,  Manchuria,  Japan,  and 
China. 

Micropus  padficus.  Breeds  in  eastern  Asia,  including  Kamchatka,  Japan,  and 
northern  China.     Has  also  been  reported  from  the  Commander  Islands. 

Coccothraustes  coccothraustes  japonicus.  Occurs  in  Japan,  Korea,  northern 
China,  and  eastern  Siberia. 

Fringilla  montif  ring  ilia.  Breeds  in  northern  Europe  and  Asia  east  to  Kam- 
chatka ;  in  migration  to  southern  Asia. 

Pinicola  leucura  kamtschathensis.     Breeds  in  Kamchatka. 

Anthus  spinoletta  japonicus.  Breeds  in  eastern  Siberia  and  Kamchatka  and 
the  Kurile  Islands ;  winters  in  Japan,  China,  and  India. 

In  addition,  Pluvialis  dominion  fulva,  added  to  the  list  of  North  American 
species  from  the  Pribilofs,  has  since  been  found  to  be  the  breeding  bird  of 
northern  and  western  Alaska. 

BIRDS  DESCRIBED  FROM  THE  PRIBILOFS. 

Five  valid  forms  have  been  described  from  Pribilof  specimer 
as  follows : 
Rissa  tridactyla  pollicaris.     Ridgway,  in  Baird,  Brewer,  and  Ridgway,  Watt  t 

Birds  of  North  America,  vol.  2,  p.  202,  1884.     Type  from  St  George  Islam* 

Ranges  generally  over  the  North  Pacific. 
Arquatella  maritime  ptilocnemds.    Coues,   in  Elliott's  Rept.     Seal   Islands  ol 

Alaska,  1874  (unpaged),  p.  182,  1875.    Type  from  St.  George  Island.     Breed 

ing  on  the  Pribilof  and  St  Matthew  Groups,  and  on  St.  Lawrence  Island. 
Plectrophenax  nivalis   toicnsendi.    Ridgway,   Manual  North  American   Birds, 

p.  403,  1887.     Type  from  Otter  Island.     The  breeding  form  on  the  Pribilof 

Aleutian,  and  Commander  Islands. 
Calcarius  lapponicus  alascensis.    Ridgway,  The  Auk,  vol.  15,  p.  320,  October, 

1S98.    Type  from  St.  Paul  Island.     The  breeding  form  on  the  islands  and 

mainland  of  northern  and  western  Alaska. 
Nannus  troglodytes  alascensis.    Baird,  Trans.  Chicago  Acad.  Sci.,  vol.  1,  p.  315, 

1869.    Type  from  St.  George  Island.     Confined  to  the  Pribilof  Group. 


1923.]  BIRDS  OF    THE   PRIBILOF   ISLANDS.  13 

GENERAL  OBSERVATIONS  ON   MIGRATION. 

Although  it  seems  very  remarkable  that  so  many  species  of  birds 
otherwise  known  as  regular  breeders  or  migrants  only  on  the  neigh- 
boring coasts  of  Alaska  or  Asia  have  been  taken  on  the  compara- 
tively small  and  distant  Pribilofs,  the  fact  that  they  have  been 
detected  is  perhaps  more  to  be  wondered  at  than  that  they  have 
reached  there.  During  migration  birds  are  often  driven  out  to  sea 
by  winds  of  unusual  velocity  or  become  bewildered  by  fogs,  while 
some  habitually  cross  wide  expanses  of  ocean.  Such  travelers,  when 
weary  and  hungry,  eagerly  grasp  the  opportunity  to  seek  the  solid 
land.  During  the  rather  brief  periods  when  the  fogs  lift,  both  the 
main  islands  are  visible  for  a  considerable  distance.  But  the  most 
potent  factor  in  aiding  the  birds  to  find  a  landing  place  is  probably 
the  noisy  herds  of  fur  seals  and  sea  lions. 

During  the  seasons  when  most  of  the  smaller  birds  are  migrating 
the  beaches  are  occupied  by  the  thronging  thousands,  whose  com- 
bined voices  produce  a  clamor  which  can  be  heard  above  all  other 
sounds  of  nature  at  a  distance  of  several  miles.  So  far  reaching  is 
this  sound  that  it  is  frequently  useful  to  navigators,  helping  them  to 
locate  the  islands  when  driven  from  their  course  by  winds  and  cur- 
rents. There  is  no  doubt  that  many  storm-driven  birds  are  thus 
guided  to  this  haven,  either  to  be  put  on  record  as  rare  visitors,  or 
after  resting  again  to  try  to  traverse  the  storm-lashed  pathway. 

The  fact  that  the  Pribilofs  have  furnished  so  many  additions  to 
the  list  of  North  American  birds,  and  that  species  from  both  North 
America  and  Asia  continue  to  be  taken  there,  has  suggested  com- 
parison with  Helgoland,  the  islet  in  the  North  Sea  which  has 
proved  such  a  rich  ornithological  field.  However,  when  we  come  to 
consider  the  characteristics  of  these  two  stations,  and  their  data  on 
migration,  it  is  apparent  that  they  have  little  common  ground  for 
comparison. 

ac  Although  the  Pribilofs  and  Helgoland  are  in  nearly  the  same 
latitude,  the  climatic  conditions  and  other  phenomena  affecting  the 
'avifauna  are  very  different.     Helgoland  is  so  situated  relative  to 
•^Scandinavia,   Germany,  Russia,   and  the   other  parts  of  northern 
„.  Europe  as  to  lie  directly  in  the  migration  pathway  of  vast  numbers 
_jpf  birds  which  breed  in  those  comparatively  temperate  lands,  and 
which   winter   in   southern   Europe   and   in   Africa.      Furthermore, 
'these  hordes  of  regular  migrants  carry  along  with  them  many  species, 
'both  northern  and  southern,  which  otherwise  would  not  wander  from 
their  native  haunts,  and  some  of  which  have  been  detected  while 
pausing  on  Helgoland.     Other   causes,   apparently,  have  operated 
in  the  case  of  many  species  which  are  breeders  in  southern  and  east- 
ern Asia  and  in  North  America. 
108731°— 23 2 


14  NORTH   AMERICAN   FAUNA.  [No.  46 

The  Pribilofs,  on  the  other  hand,  lie  in  the  midst  of  Bering  Sea, 
in  an  environment  essentially  Arctic,  and  not  on  a  migration  route 
which  is  followed  by  any  considerable  number  of  birds,  either  those 
passing  between  their  summer  and  winter  homes  in  western  America 
or  those  which  habitually  cross  between  America  and  Asia.  Those 
which  do  use  the  islands  as  a  stepping  stone  are  certain  waders  and 
swimmers  which  nest  in  northwestern  Alaska,  and  winter  in  the 
Aleutians,  the  Pacific  Islands,  or  southeastern  Asia.  The  more 
notable  of  this  class  are  the  following:  Philacte  canagica,  Limosa 
lapponica  baueri,  Heteroscelus  incanus  incaniLS,  Pluvialis  dominica 
fulva,  and  Arenaria  interpres  interpres. 

As  regards  land  birds,  however,  it  is  a  notable  fact  that  none  of 
the  small  species  of  birds  which  are  essentially  Asiatic,  but  which 
cross  each  summer  to  nest  in  Alaska  (including  Budytes  flavus 
alascensis  and  Acanthopneuste  borealis),  have  been  detected  on  the 
Pribilofs. 

A  canvass  of  the  list  of  birds  which  have  been  observed  on  Helgo- 
land, leaving  out  those  crag-nesting  species  which  are  influenced  in 
their  choice  of  homes  by  the  presence  of  suitable  cliffs  and  an  abun- 
dant marine  food  supply,  shows  that  about  283  species  nest  south 
of  the  limit  of  trees,  while  only  83,  or  less  than  23  per  cent  of  the  total 
thus  considered,  are  Arctic,  nesting  north  of  the  tree  limit.  Apply- 
ing the  same  test  to  Pribilof  birds  the  proportion  is  exactly  50  per 
cent  in  each  class.  Helgoland  has  had  the  advantage  of  more  inten- 
sive observations,  continued  over  a  much  longer  period,  but  since 
collections  and  observations  in  both  places  have  been  made  indis- 
criminately it  is  unlikely  that  the  relative  numbers  of  species  nest- 
ing above  and  below  the  limit  of  trees  would  be  found  to  be  ma- 
terially different  were  the  data  more  comparable  in  this  regard. 

It  is  evident,  therefore,  that  the  avifauna  of  the  two  places  is 
radically  different  in  character,  since  we  have  on  Helgoland  an 
overwhelming  majority  of  non- Arctic  species,  many  of  which  are 
wanderers  from  their  regular  homes  in  Africa,  Asia,  or  America, 
while  those  found  on  the  Pribilofs  which  may  be  classed  as  strays 
are  in  all  cases  natives  of  the  neighboring  coasts  of  Asia  and  America. 
Furthermore,  those  transient  land  birds  which  have  been  detected  on 
the  Pribilofs  seem  to  have  reached  there  largely  by  chance  rather 
than  by  following  any  regular  migration  routes. 

FOOD  HABITS  OF  BIRDS. 

The  notes  on  food  habits  of  each  species  and  also  the  following 
general  account  of  the  food  resources  of  Pribilof  birds  and  of  the 
circumstances  attending  the  elaboration  of  the  available  data  on  the 
subject  were  prepared  by  W.  L.  McAtee: 


1923.]  BIRDS  OF    THE   PRIBILOF   ISLANDS.  15 

All  pertinent  previous  reports  on  Pribilof  birds  have  been  ex- 
amined and  remarks  on  bird  food  quoted  from  them  when  desirable. 
Field  notes  of  various  naturalists  who  have  visited  the  islands  also 
have  been  drawn  upon  and  when  used  the  source  is  acknowledged. 
The  contents  of  32  bird  stomachs  from  the  Pribilofs  were  reported 
on  by  Dr.  S.  D.  Judcl  in  William  Palmer's  "Avifauna  of  the  Pribilof 
Islands"  (1899,  pp.  380-427).  In  some  cases  these  stomachs  have 
been  reexamined;  in  others  the  original  report  has  been  referred  to. 
For  the  present  account  a  considerable  collection  of  stomachs  has 
been  available  for  examination,  largely  due  to  the  efforts  of  G. 
Dallas  Hanna.0 

Features  of  the  food  habits  of  birds  of  the  Pribilofs  that  seem  of 
sufficient  interest  to  merit  special  mention  are  the  great  diversity 
of  organisms  fed  upon  by  some  of  the  birds  resorting  to  the  kelp 
beds,  and  the  unusual  preponderance  of  two-winged  flies  in  the  diet 
of  birds  feeding  on  land.  In  both  cases,  the  phenomenon  mentioned 
undoubtedly  is  merely  a  reflection  of  relative  availability  of  food 
items.  It  is  well  known  that  Diptera  are  the  dominant  insects  of 
high  latitudes,  and  collections  made  in  connection  with  the  present 
investigation  emphasize  this  fact.  They  are  the  most  numerous  in- 
sects both  in  species  and  individuals  and  consequently  contribute 
most  voluminously  to  the  subsistence  of  the  chief  predatory  crea- 
tures of  the  region,  namely,  birds. 

The  kelp  pastures  are  the  richest  sort  of  feeding  places  for  omnivo- 
rous birds,  and  such  species  as  the  eider  duck  seem  to  draw  on  all  the 
inhabitants  of  these  beds  for  sustenance.  Representatives  of  most 
of  the  main  branches  of  the  animal  kingdom  in  addition  to  various 
kinds  of  algae  are  frequently  found  in  single  stomachs.  The  fol- 
lowing brief  list  of  the  mixed  contents  of  four  Pacific  eiders  will 
illustrate  how  the  whole  range  of  food  items  is  drawn  upon  by  these 
birds:  Plume-algae  (2  species),  kelp,  hydroids  (2  kinds),  mussels 
(3  kinds),  sea  snails  (7  species),  marine  worms  (2  kinds),  star- 
fishes, sea  urchins,  amphipods  (2  sorts),  hermit  crabs  (2  species), 
barnacles,  sea-squirt,  and  sculpin.  To  some  it  may  be  surprising 
that  so  varied  an  assemblage  of  marine  organisms  is  available  in 

a  Most  of  the  stomach  contents  were  analyzed  by  W.  L.  McAtee,  but  numerous  stomachs 
of  certain  groups  of  birds  were  examined  by  the  following:  Ira  N.  Gabrielson,  finches; 
Charles  C.  Sperry,  shorebirds  ;  and  Alexander  Wetmore,  grebes  and  shoreblrds.  Identifica- 
tions of  many  items  found  in  the  stomachs  were  based  on  the  collections  of  invertebrates 
reported  upon  in  the  appendix  to  the  present  paper,  and  to  the  determiners  thereof  is 
due  much  of  the  credit  for  details  in  the  statements  relating  to  bird  food.  In  addition 
many  other  specimens  were  directly  referred  to  specialists  for  identification.  Those  who 
cooperated  in  this  way  were :  W.  A.  Setchell,  algae ;  Agnes  Chase  and  F.  P.  Metcalf, 
grass  seeds  ;  C.  C.  Nutting,  hydroids  ;  W.  H.  Dall  and  Paul  Bartsch,  mollusks  ;  R.  C.  Osburn, 
bryozoans  ;  A.  H.  Clark,  starfishes,  sea  urchins,  and  ascidians ;  Mary  J.  Rathbun,  crabs ; 
W.  L.  Schmitt,  other  decapods ;  Clarence  L.  Shoemaker,  amphipods ;  H.  A.  Pilsbry, 
barnacles ;  S.  A.  Rohwer,  sawflies  ;  Leon  J.  Cole,  pycnogonids  ;  and  Alexander  Wetmore, 
birds.     To  all  of  these  grateful  acknowledgment  is  made. 


16  NORTH   AMERICAN   FAUNA.  [No.  46 

this  far-northern  region,  and  this,  too,  during  the  winter  months  as 
well  as  the  summer.  It  would  seem  that  these  Boreal  kelp  beds  rival 
in  abundance  and  diversity  of  life  the  famed  tide-pools  of  the 
Tropics.  While  the  birds  frequenting  them  must  necessarily  be  of 
hardy  races  that  can  endure  the  cold  and  buffeting  of  the  wintry 
gales  of  the  Bering  Sea,  it  is  apparent  that  insufficient  and  monoto- 
nous diet  decidedly  is  not  a  characteristic  of  this  environment,  one 
usually  thought  of  as  extremely  rigorous. 

Family  COLYMBIDAE. 

/ 
Colymbus  holboellii  (Reinbardt).     Holboell  Grebe. 

The  Holboell  grebe  was  first  recorded  from  the  islands  by  Coues 
(in  Elliott,  1874,  unpaged,  and  1875,  p.  201),  who  recorded  the 
specimen  as  Podiceps  griseigena.  This  specimen,  still  in  the  U.  S. 
National  Museum  collection  (No.  64302  im.),  has  been  correctly  iden- 
tified as  Colymbus  holboettii  by  Nelson  (1887,  p.  35)  and  Palmer 
(1899,  p.  383).  It  was  taken  by  Elliott  at  St.  George  Island,  June 
22,  1873. 

A  second  specimen  (No.  237485.  male,  U.  S.  Nat.  Mus.)  was  taken 
by  G.  Dallas  Hanna  at  St.  Paul  Island,  December  6,  1914.  It  was 
feeding  in  the  surf  off  Kitovi  Rookery  when  shot. 

Food. — The  stomach  contents  of  the  December  specimen  was  com- 
posed of  a  few  feathers,  and  food  items  as  follows:  Otoliths  of  40 
or  more  small  fishes,  further  unidentified,  99  per  cent;  and  numerous 
amphipods  of  the  family  Pontogeneidae,  1  per  cent. 

Colymbus  auritus  Linnaeus.     Horned  Grebe. 

The  horned  grebe  was  first  recorded  from  the  Pribilofs  by  G.  Dal- 
las Hanna,  who  obtained  specimens  from  both  St.  Paul  and  St. 
George  Islands  in  1913  and  1914,  as  recorded  by  him  (1916,  p.  401). 
Following  are  the  records  of  the  several  specimens:  Female,  shot  in 
Cemetery  Lake,  St.  George,  October  13,  1913  (stomach  contained  a 
quantity  of  decayed  vegetation  from  bottom  of  lake)  ;  female,  in 
surf,  St.  George,  December  19,  1913  (feet  and  bill  slate  color;  eye, 
orange);  male,  November  30,  1914,  Southwest  Bay,  St.  Paul;  one 
(sex  not  recorded),  January  6,  1915,  St.  Paul.  Mr.  Hanna  states 
that  other  single  birds  were  seen  in  the  surf  about  St.  Paul  on  De- 
cember 4  and  13,  1914,  and  in  the  Salt  Lagoon  on  at  least  two  occa- 
sions, and  that  the  natives  consider  it  quite  common  in  winter.  All 
the  specimens  above  listed  are  in  the  collection  of  the  U.  S.  National 
Museum. 

Food. — The  contents  of  the  stomachs  of  the  three  last  mentioned 
have  been  analyzed.  As  in  the  case  of  grebes  collected  elsewhere,  the 
bird's  own  feathers  are  a  prominent  part  of  the  stomach  contents, 


1923.]  BIRDS   OF   THE   PRIBILOF   ISLANDS.  17 

varying  in  the  samples  at  hand  from  5  to  98  per  cent  of  the  entire 
bulk.  Ignoring  these,  the  food  in  these  three  stomachs  was  com- 
posed of  Crustacea,  75  per  cent;  marine  worms  {Nereis),  23.3  per 
cent;  and  fish,  1.7  per  cent.  The  crustaceans  were  exclusively  am- 
phipods  identified  as  Chironcsimus  multiarticulatus  and  Pontoge- 
neia  sp.,  of  the  family  Pontogeneidae,  and  others  of  the  families 
Calliopiidae,  Metopidae,  and  Gammaridae. 

Family  GAVIIDAE. 

Gavia  adamsii  (Gray).     Yellow-billed  Loon. 

The  yellow-billed  loon  is  a  transient  visitor,  first  recorded  from 
the  islands  by  C.  H.  Townsend,  who  says  (1887,  p.  98)  :  "I  saw  a 
skin  of  the  white-billed  loon  at  the  house  of  the  United  States  Treas- 
ury agent  on  St.  Pauls  Island.  It  was  killed  there  by  a  native  in 
August,  1885."  A  second  specimen  was  taken  by  M.  C.  Marsh  on 
St.  Paul,  May  18,  1912,  and  is  now  in  the  Biological  Survey  collec- 
tion. It  is  a  male,  and  measured  in  the  flesh:  Length,  34  inches; 
extent,  61.  The  only  other  record  refers  to  10  individuals  observed 
by  G.  Dallas  Hanna  off  St.  George  Island,  August  18,  1913. 

Gavia  viridigularis  Dwight.     Green-throated  Loon. 

The  green-throated  loon  was  first  recorded  by  Coues  in  Elliott's 
Report  (1874,  unpaged,  and  1875,  p.  201)  under  the  name  Colymbus 
arcticus,  from  a  bird  found  cast  up  nearly  dead  on  the  beach  at 
Zapadni,  St.  George  Island,  June  22,  1873.  This  specimen,  which  is 
still  in  the  collection  of  the  U.  S.  National  Museum  (No.  64303,  male), 
has  been  several  times  recorded  under  the  name  arcticus,  and  has 
played  no  small  part  in  the  claim  of  that  species  to  be  considered  a 
North  American  bird.  This  specimen  has  been  recently  examined 
and  found  to  be  referable  to  the  species  lately  described  by  Dr. 
Jonathan  Dwight,  Gavia  viridigularis  (Auk,  1918,  p.  198),  as,  indeed, 
was  intimated  by  Dwight  in  the  introduction  to  his  article  (p.  196). 
Another  of  the  two  remaining  specimens  which  have  been  referred 
to  G.  arcticus  (No.  76004,  U.  S.  Nat.  Mus.,  St.  Michael,  Alaska,  Aug- 
ust 24,  1877,  E.  W.  Nelson),  is  also  plainly  referable  to  G.  viridi- 
gularis. Both  these  specimens  have  a  decided  green  reflection  on 
the  dark  areas  of  the  throat,  when  viewed  toward  the  light,  while 
in  G.  arcticus  and  all  its  races  these  areas  have  a  purplish  reflection. 
As  far  as  known  Gavia  viridigularis  (type  locality,  Gichega,  north- 
eastern Siberia)  is  confined  to  northeastern  Siberia  and  extreme 
western  Alaska. 

Family  ALCID^E. 

Lunda  cirrhata  (Pallas).     Tufted  Puffin. 

The  tufted  puffin  was  first  recorded  specifically  from  the  islands 
by  Coinde  (1860,  p.  403),  although  Veniaminof  in  1840  (quoted  by 


18  NORTH   AMERICAN   FAUNA.  [No.  46 

Elliott,  1874,  unpaged,  and  1875,  p.  242)  mentions  puffins  as  occur- 
ring and  probably  refers  in  part  to  the  present  species. 

An  abundant  summer  resident  on  the  Pribilofs,  breeding  on  all 
the  five  islands  of  the  group.  It  arrives  about  the  middle  of  May 
and  the  majority  leave  the  islands  by  the  last  of  August. 

Elliott  gives  May  10  as  about  the  time  of  its  arrival ;  the  earliest 
dates  I  have  for  recent  years  are  as  follows:  St.  Paul,  March  5,  1911, 
specimen,  perhaps  wintered  (Hahn)  ;  St.  Paul,  May  20,  1911,  several 
seen  (Hahn)  ;  St.  Paul,  May  20,  1912,  specimen  (Hahn)  ;  St.  Paul, 
May  19,  1914,  eight  observed /(Hanna)  ;  St.  Paul,  May  15,  1915, 
three  seen,  arrivals  (Hanna);  St.  George,  May  21,  1917  (Hanna). 

The  eggs  are  usually  laid  in  burrows  dug  by  the  birds  in  turfy 
soil  at  the  edge  of  cliffs  or  on  the  summits  of  the  smaller  islands. 
Numbers  nest  in  easily  accessible  locations  on  Walrus  Island,  Otter 
Island,  and  Sealion  Rock,  where  they  are  safe  from  the  foxes.  On 
the  other  islands  the  birds  nest  among  the  rocks  for  protection. 
Dates  for  eggs  are  as  follows :  Otter  Island,  June  24,  27,  1884  (Lutz, 
1889,  p.  31)  ;  Sealion  Rock,  June  29,  1914,  some  fresh,  but  mostly 
heavily  incubated  (Preble)  ;  St.  George,  July  1,  1914  (Hanna)  ; 
Walrus  Island,  July  7,  1911,  eggs  being  incubated  (Bent,  1919,  p. 
83);  Otter  Island,  July  4,  1890;  Walrus  Island,  August  7,  1890, 
slightly  incubated  (Palmer,  1899,  p.  384).  That  many  birds  nest 
late  in  the  summer  is  shown  by  the  fact  that,  as  elsewhere  mentioned, 
Hahn  observed  many  young  birds  taken  from  nests  on  AValrus  Island 
as  late  as  October  2. 

The  bulk  of  the  individuals  have  left  the  islands  before  September. 
Later  notable  dates  are  as  follows:  St.  Paul,  early  September,  1910, 
still  common  (Hahn)  ;  Walrus  Island,  October  2,  1910,  about  40 
young  birds  taken  from  nests  by  natives,  adult  and  young  specimens 
taken,  numerous  at  sea  (Hahn)  ;  St.  Paul,  December  8,  1910,  one 
young  bird  found  dead  (Hahn)  ;  St.  George,  September  4,  1913,  150 
observed  (Hanna)  ;  St.  George,  September  15, 1913,25  seen  (Hanna)  ; 
St.  Paul,  September  22,  1914,  75  seen  (Hanna) ;  St.  Paul,  December 
6,  1914,  4  seen  (Hanna)  ;  St.  Paul,  December  13,  1914,  10  seen 
(Hanna). 

The  following  notes  made  by  the  collectors,  and  relating  to  fresh 
specimens,  are  of  interest:  No.  63,  Hahn,  female  adult,  St.  Paul 
Island,  March  5,  1911:  Length,  15i  inches;  extent,  27f ;  bill,  deep 
orange  on  distal  half;  culmen  and  cutting  edges  washed  with  plum- 
beous; iris,  pinkish;  feet,  pale  flesh-color.  No.  187,  Marsh,  female 
adult,  St.  Paul,  May  20,  1912,  length,  15*  inches;  extent,  27ji  No. 
35,  Hahn,  Walrus  Island,  October  2,  1910  (young  bird  with  much 
down  remaining  on  neck  and  rump)  :  Upper  mandible,  plumbeous; 
lower  mandible,  paler  toward  tip;  eyelids,  dusky;  feet,  dusky  gray 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  19 

above,  with  web  sooty  below.  No.  36,  Hahn,  female  adult,  October 
2,  1910,  Walrus  Island:  Length,  15  inches;  extent,  29;  base  of  bill 
horn-color,  tip  deep  orange ;  iris,  yellow ;  feet,  pale  orange. 

Fratercula  corniculata   (Naumann).     Horned  Puffin.  PI.  VI,  fig.  2. 

The  horned  puffin  is  an  abundant  summer  resident  on  the  Pribilof  s, 
arriving  early  in  May  and  usually  departing  in  early  September.  It 
seems  to  be  more  abundant  than  the  tufted  puffin.  Puffins  were  listed 
by  Veniaminof  in  1840 ;  among  the  species  occurring  on  the  islands 
he  could  hardly  have  failed  to  notice  this  conspicuous  one  (Trans,  by 
Elliott,  1874,  unpaged,  and  1875,  p.  242).  First  recorded  specifi- 
cally by  Coues,  in  Elliott  (1.  c,  p.  202). 

Arrival  notes  in  recent  years  are  as  follows :  St.  Paul,  May  9,  1911, 
one  shot  (Hahn)  ;  St.  George,  May  19,  1914,  May  9,  1915,  5  ob- 
served (Hanna)  ;  St.  Paul,  May  10,  1915,  10  observed  (Hanna)  ;  St. 
George,  May  21,  1917,  observed  (Hanna). 

The  bird  nests  on  all  of  the  five  islands  of  the  group,  laying  its 
single  egg  in  a  cavity  among  the  rocks.  The  egg  is  white  with  ob- 
scure spotting  about  the  larger  end.  Dates  for  eggs  are  as  follows : 
St.  George,  July  4, 1873,  Elliott  (Palmer,  1899,  p.  385) ;  Otter  Island, 
June  19,  20,  24,  26,  30,  1884  (Lutz,  1889,  p.  31)  ;  St.  George,  July  1, 
1914  (Hanna).  The  majority  leave  the  islands  by  early  September. 
Some  definite  later  dates  follow :  St.  Paul  Island,  September  1,  1910, 
25  observed;  September  7,  10;  September  8,  6;  September  9,  10; 
September  11,  3;  October  2,  many  observed  on  Walrus  Island  but 
less  numerous  at  sea  than  the  tufted  puffin;  October  18,  1  shot  on 
North  Shore ;  October  19,  observed ;  November  29,  about  20  seen  off 
Kitovi  Rookery,  St.  Paul;  December  6,  specimen  (all  by  Hahn). 
September  28,  1913,  about  500  birds  seen  about  Garden  Cove,  St. 
George  Island,  although  most  of  the  birds  had  then  left  the  cliffs 
on  the  north  side  of  the  island  (Hanna).  St.  Paul  Island,  September 
22,  1914,  100  birds  observed;  December  6,  5  observed;  December  13, 
75  observed,  and  many  found  dead  on  the  beach  (Hanna). 

The  following  notes  were  made  from  fresh  specimens ;  Adult  male, 
St.  Paul  Island,  September  18,  1910,  Hahn;  length  13|  inches:  ex- 
tent, 25;  eyelids  deep  orange;  eye  tuft  plumbeous;  legs  deep  orange; 
bill  lemon  at  base  shading  to  deep  orange  at  tip.  Adult  female,  St. 
Paul  Island,  December  6,  1910,  Hahn;  length,  13  inches;  extent,  24^: 
bill  dusky,  feet  pearl  gray  above,  dusky  below.  Male,  St.  Paul 
Island,  September  23,  1911,  length,  13  inches;  extent,  25^;  bill 
uniform  light  yellowish  on  all  basal  portions  including  collar  of  upper 
mandible,  the  rest  or  terminal  third  dark  orange-red;  the  grooves 
darkened.  No  vermilion  red  on  the  bill ;  edge  of  eyelids  bright  red ; 
angle  of  mouth  lemon-yellow ;  legs  and  feet  orange,  deepest  on  webs  ; 
yellow  on  outer  surface  of  tarsus  and  outer  toes,  and  more  or  less  on 


20  NORTH   AMERICAN  FAUNA.  [No.  4G 

all  toes.  Male,  St.  George  Island,  August  23, 1913,  Hanna;  bill  waxy 
olive-green ;  light  orange  in  corner  of  mouth ;  point  of  upper  mandi- 
ble dark  orange ;  of  lower,  light  orange ;  a  dark  patch  at  edge  of  each 
mandible  near  tip,  darkest  on  posterior  edge. 

Food. — The  stomach  of  a  specimen  collected  December  6,  1910,  was 
reported  in  field  notes  of  W.  L.  Hahn  to  contain  a  few  isopods  and 
fragments  apparently  of  seaweed. 

Phaleris  psittacula  (Pallas).     Paroquet  Auklet  PI.  VI,  fig.  1. 

The  paroquet  auklet,  a  beautiful  and  interesting  species,  the  "  Bail- 
lie  Brushkie  "  of  the  Aleuts,  is  ,&  rather  common  summer  resident  on 
the  Pribilofs.  It  appears  to  have  been  first  recorded  from  the  islands 
by  Veniaminof  in  1840  under  the  name  "  white-breasted  auk  "  (Trans, 
by  Elliott,  1874,  unpaged,  and  1875,  p.  242). 

Spring  records  follow :  St.  Paul  Island,  May  1  and  2,  1900  (com- 
mon, notes  from  island  log  by  Hahn) ;  St.  Paul  Island,  May  1,  1911, 
some  taken  by  natives  (Hahn)  ;  St.  Paul  Island,  April  30,  1912 
(specimen  by  Marsh)  ;  St.  George  Island,  March  28,  1914,  8  observed; 
April  3,  observed ;  April  8,  20  observed ;  April  19,  begun  lighting  on 
cliffs;  April  22,  20  observed  (Hanna)  ;  April  24,  1915,  300  on  Tolstoi 
Cliffs,  St.  Paul  Island;  April  27,  500  on  Tolstoi  Cliffs;  May  4  and  9, 
abundant  (Hanna). 

It  breeds  on  all  the  five  islands  of  the  group,  most  abundantly  on 
St.  George  and  St.  Paul  Islands,  in  large  numbers  on  Otter  Island, 
rather  commonly  at  times  on  Walrus  Island,  and  in  small  numbers 
even  on  Sea  Lion  Bock.  Its  single  white  egg,  seemingly  large  for 
the  bird,  is  usually  laid  beneath  large  bowlders  or  in  practically  in- 
accessible cavities  in  the  cliffs;  in  some  cases  the  egg  is  deposited 
in  a  burrow  apparently  excavated  in  the  loose  soil  topping  the  vol- 
canic cliffs.  Dates  for  eggs  follow :  St.  George  Island,  July  1,  1873 ; 
Walrus  Island,  June  13,  1890  (egg  collection,  U.  S.  Nat.  Mus.) ; 
Otter  Island,  June  20,  23,  1884  (Lutz,  1889,  p.  31)  ;  St.  George 
Island,  July  1,  1914  (Hanna) ;  Pribilof  Islands,  June  8,  July  7,  July 
16  (Bent,  1919,  p.  120). 

Autumn  and  winter  dates  follow:  Elliott  (1875,  p.  205)  states 
that  they  depart  from  the  islands  between  August  20  and  Septem- 
ber 1,  and  this  statement  seems  to  be  generally  true.  Later  detailed 
observations  are:  St.  Paul,  1910:  August  30,  specimen;  August  31; 
September  7,  one  observed ;  October  2,  several  seen  on  way  to  Walrus 
Island;  November  7,  specimen  (Hahn).  St.  George  August  20,  1913, 
not  seen  later  (Hanna).  St.  Paul,  1914:  October  4,  4  observed; 
December  4,  20  seen;  December  13,  200  seen,  flocks  abundant  off- 
shore, many  dead  on  beach  (Hanna).  St.  Paul,  February  18,  1915, 
large  flocks  just  off-shore,  Northeast  Point  (Hanna).  St.  George, 
February  6,  1917,  2  killed  (Hanna). 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey. 


PLATE   VI. 


Fig.  I. — Paroquet  Auklet  (Phaleris  psittacula)  and  Group  of  Least  Auklets 

(Aethia  pusilla). 

Many  of  the  breeding  birds  associate  closely.    Photograph,  by  G.  Dallas  Hanna,  summer,  1920,  on 

St."  George  Island. 


Fig.  2. — Horned  Puffin   (Fratercula  corniculata). 

These  interesting  birds  nest  chiefly  in  clefts  among  the  rocks.    Photograph  by  G.  Dallas  Hanna, 

on  St.  George  Island. 


Fig.  3. — Pacific   Kittiwake   (Rissa  tridactyla  pollicaris)  on   Nesting   Cliff. 

This  is  the  most  abundant  of  the  two  species  nf  kittiwakes.    Photograph  by  G.  Dallas  Hanna,  on 

St.  George  Island. 


1923.]  BIRDS   OF   THE   PRIBILOF   ISLANDS.  21 

The  species  thus  winters  at  sea  in  the  vicinity  of  the  islands,  and 
thence  southward  to  the  latitude  of  Monterey  Bay,  Calif.,  and  the 
Kurile  Islands,  Japan. 

A  female  taken  on  St.  Paul  Island,  May  20,  1911  (Hanna),  meas- 
ured in  the  flesh :  Length,  10  inches,  extent  21 ;  the  bill  was  blood 
orange  in  color;  feet  and  tarsus,  bluish  in  front,  dusky  behind.  A 
male,  taken  April  30,  1912,  measured:  Length,  10§  inches,  extent, 
21 ;  bill,  coral  red,  rostral  shield  darker.  A  female,  same  place  and 
date,  length,  9  J  inches,  extent  20^ ;  bill  coral  red. 

Food.— H.  W.  Elliott  stated  (1874,  unpaged,  and  1875,  p.  205)  that 
the  food  of  the  paroquet  auklet  consists  of  amphipods  and  fish  fry. 
He  says  further  that  "  I  have  never  seen  one  among  the  thousands 
that  were  around  me  when  on  the  islands  '  opening '  the  bivalve 
shells,  such  as  mussels,  etc.,  as  stated  by  Professor  Brandt."  The 
suggestion  of  the  latter  author  no  doubt  was  a  deduction  from  the 
shape  of  the  bill  of  the  paroquet  auklet,  but  with  little  doubt  is 
erroneous.  The  only  stomach  of  this  species  available  was  collected 
at  St-  Matthews  Island,  July  12,  1916  (G.  D.  Hanna),  and  the  content 
consisted  entirely  of  remains  of  small  Crustacea. 

Aethia  cristatella  (Pallas).     Crested  Auklet. 

The  crested  auklet  is  a  fairly  common  summer  resident  on  the 
Pribilof  Group,  nesting  on  all  the  five  islands  excepting  Sealion 
Rock.  It  usually  comes  on  the  cliffs  in  April  or  earlier,  but  this 
date  is  uncertain  and  difficult  to  determine,  since  large  numbers  pass 
the  winter  on  the  ocean  in  the  vicinity  and  are  likely  to  be  seen  in 
the  surf  at  any  time.  It  is  called  by  the  natives  "  Canooskie  "  (Little 
Captain).  The  species  was  first  recorded  from  the  islands  by 
Veniaminof  in  1840,  under  the  name  crested  auk.  (Trans,  by  Elliott, 
1874,  unpaged,  and  1875,  p.  242.) 

Available  spring  records  of  occurrence  later  than  February  are 
as  follows:  St.  Paul  Island,  April  5,  1901,  a  few  brought  in;  St.  Paul, 
April  10,  1906,  some  shot  (notes  taken  from  island  log  by  W.  L. 
Hahn)  ;  April  30,  1911,  5  or  6  shot  by  natives  at  Tolstoi  Point,  St. 
Paul  (Hahn)  ;  St.  Paul,  March  12,  16,  18,  1912,  specimens  (Marsh)  ; 
St.  George,  April  8,  1914,  thousands  observed  (Hanna)  ;  St.  Paul, 
1915 — April  15,  hundreds;  April  18,  many  near  shore;  April  24,  40 
seen;  April  27,  30  seen;  May  6,  abundant  (Hanna). 

This  species  usually  nests  about  the  high  cliffs,  the  single  white 
egg  being  deposited  in  a  deep  and  usually  inaccessible  recess;  oc- 
casionally the  birds  lay  in  the  depths  of  the  bowlder  beaches.  Eggs 
have  been  taken  on  the  following  dates :  St.  George  Island,  June  19, 
20,  and  July  4,  1873 ;  St.  Paul  Island,  July  10,  1895  (collection  U.  S. 
Nat.  Mus.)  ;  Otter  Island,  June  20,  22,  1884  (Lutz,  1889,  p.  31)  ; 
Walrus  Island,  June  16, 1910  (James  Judge). 


22  NORTH  AMERICAN  FAUNA.  [No.  46 

Notes  on  fall  and  winter  occurrence  follow:  St.  Paul  Island, 
January  31,  1904,  and  December  6,  1909,  numbers  seen  (notes  from 
island  log).  St.  George,  vast  numbers  seen  about  end  of  November, 
1905  (Chichester,  1908,  p.  49).  St.  Paul,  1910— probably  5,000  seen 
at  sea  November  8,  usually  flying  at  great  speed  close  to  the  water; 
many  seen  at  sea  November  9,  22,  and  29,  off  St.  Paul  (Hahn).  A 
great  many  flocks  of  hundreds  each  seen  about  1  mile  off  St.  George 
January  4,  1914  (Hanna).  Hundreds  of  thousands  seen  off  St.  Paul 
December  13,  1914,  and  February  18  and  March  5,  1915  (Hanna). 
Taken  on  St.  George,  October/18  and  29,  1915  (H.  P.  Adams). 
Abundant  at  edge  of  ice  pack  off  St.  George,  January  25,  1917 
(Hanna).  Some  killed  at  sea  near  St.  Paul,  February  8,  1918 
(Hanna). 

These  flocks  wintering  at  sea  are  observed  well  into  the  spring. 
Thus  Hanna  noted  them  off  St.  George  Island  March  21  and  April 
8,  1914. 

On  several  occasions  numbers  of  the  birds  have  been  found  dead 
on  the  beaches,  usually  in  the  early  summer.  These  may  represent 
merely  the  normal  mortality  from  the  large  flocks  which  winter  in 
the  vicinity. 

G.  Dallas  Hanna,  who  has  had  exceptional  opportunities  for  ob- 
serving this  species,  thinks  that  great  numbers  of  crested  auklets 
move  northward  in  winter  from  their  principal  breeding  grounds 
south  of  Bering  Sea,  and  that  this  accounts  for  the  immense  num- 
bers sometimes  seen  at  this  season,  when  they  are  much  more 
abundant  than  in  summer.    He  states : 

This  species  evidently  winters  not  far  to  the  south  of  the  Pribilof  Islands, 
since  after  a  severe  storm  from  this  quarter  in  winter  the  birds  are  almost 
always  seen  in  small  numbers  and  occasionally  in  enormous  flocks,  as  in  Feb- 
ruary, 1915,  off  St.  Paul,  and  January,  1917,  off  St.  George.  At  these  times 
the  birds  gather  on  the  water  as  closely  as  they  can  sit,  and  on  both  of  the 
above  occasions  some  flocks  of  the  birds  covered  as  much  as  5  acres.  This  is  a 
conservative  statement,  as  applied  to  the  larger  flocks,  as  the  areas  were  care- 
fully estimated.  At  the  same  time  there  were  many  smaller  flocks  in  the 
vicinity.  The  presence  of  drift  ice  does  not  prevent  them  from  appearing  after 
a  blow;  they  then  gather  in  the  open  spaces  in  their  customary  manner.  (MS. 
notes,    1920.) 

The  flesh  measurements  of  a  number  of  specimens,  recorded  by 
M.  C.  Marsh,  range  as  follows:  Length,  9  to  10£  inches;  extent, 
18f  to  19{J.  No  appreciable  sexual  difference  in  size  is  apparent. 
A  male  (No.  104,  M.  C.  Marsh)  taken  on  St.  Paul  Island  December 
5,  1911,  measured:  Length,  10  inches;  extent,  19.  Notes  on  the  colors 
of  the  soft  parts  follow :  Iris  with  a  ring  of  pale  green  within  yel- 
low, finally  bordered  by  a  black  circle  on  the  edge  of  sclerotic;  legs 
and  feet  pale  bluish,  blackish  behind,  the  web  dusky  in  front;  bill 
plain  brown,  lighter  underneath. 


1923.]  BIRDS   OF   THE  PMBILOF  ISLANDS.  23 

A  summer  female  (No.  349,  Hanna)  taken  on  St.  George  August 
22,  1913,  had  the  bill  lead  color  at  the  tip,  brown  at  base;  iris  white; 
feet  and  toes  slate ;  webs,  black. 

Food. — Seven  well-filled  stomachs  of  crested  auklets  from  the 
Pribilofs  contained  only  amphipods,  identified  as  P arathemisto 
oblivia  in  five  cases  and  as  Socarnea  vahli  in  one.  Another  nearly 
empty  stomach  contained  only  two  bits  of  kelp.  Six  of  the  stomachs 
holding  amphipods  were  collected  in  midwinter,  and  the  following 
note  by  G.  Dallas  Hanna  is  of  interest  as  indicating  the  depth  to 
which  the  birds  must  descend  at  that  season  to  secure  their  staple 
food: 

On  several  occasions  I  have  been  on  fishing  parties  out  among  these  birds 
and  twice  we  pulled  up  codfish  which  had  in  their  stomachs  in  one  case  one, 
and  in  the  other  case  two,  birds  of  this  species.  As  the  cod  is  a  bottom  feeder 
it  seems  evident  that  the  birds  descend  to  30  fathoms,  the  depth  of  the  water 
at  the  localities  concerned.  I  believe  it  is  customary  for  most  of  the  diving 
birds  to  feed  on  the  bottom,  because  I  have  been  unable  to  secure  amphipods 
at  intermediate  depths  in  the  winter  time.  In  summer,  however,  the  crustaceans 
come  nearer  the  surface  and  are  more  easily  secured. 

Aethia  pusilla  (Pallas).     Least  Auklet.  PI.  VI,  fig.  1. 

The  least  auklet,  the  "  Choochkie "  of  the  natives,  is  excessively 
abundant  on  the  Pribilofs  during  the  breeding  season,  being  present 
in  countless  thousands,  probably  millions.  Palmer  estimated  their 
numbers  as  equaling  the  sum  of  all  the  other  species.  They  breed 
on  all  the  islands,  but  most  abundantly  on  St.  George,  where  an  im- 
mense colony  occupies  the  base  and  slopes  of  a  high  hill,  Ulekiah, 
which  rises  near  the  center  of  the  island.  Elsewhere  they  nest  in 
the  cliffs  and  bowlder  beaches  throughout  practically  the  entire  coast 
line  of  the  different  islands.  The  species  was  first  recorded  from  the 
islands  by  Coinde  (1860,  p.  403). 

Apparently  the  birds  resort  to  land  only  to  breed.  A  few  winter 
at  sea  about  the  Pribilofs,  and  these  flocks  are  augmented  in  spring 
by  the  arrival  of  the  myriads  which  have  wintered  to  the  southward. 
A  few  days  after  their  appearance  offshore  they  begin  to  alight  on 
the  cliffs  and  bowlder  beaches,  and  soon  become  abundant  on  their 
breeding  grounds. 

Since  these  birds  form  an  important  and  welcome  addition  to  the 
spring  food  supply  of  the  natives,  their  arrival  is  eagerly  awaited, 
and  many  notes  on  spring  arrival  have  been  recorded.  The  follow- 
ing, referring  to  St.  Paul  Island,  were  taken  from  the  island  log 
by  W.  L.  Hahn:  April  30,  1886,  arrival;  April  25,  1889,  arrival; 
April  18,  1899,  arrival;  April  28,  1900,  small  numbers;  May  1  and  2, 
common;  April  24,  1901,  arrival ;  April  25,  numerous;  April  18,  1902, 
arrival;  April  23,  1903,  arrival;  April  13,  1904,  arrival;  April  19, 
1906,  seen  at  sea ;  May  2,  1908,  arrival ;  April  26,  1909,  arrival ;  May 
10,  1910,  first  seen  flying  over  the  land.     Notes  during  recent  years 


24  NORTH  AMERICAN   FAUNA.  [No.  46 

are:  St.  Paul  Island,  1911— April  18,  arrival;  April  27,  about  100 
seen;  April  28,  first  seen  flying  over  land;  May  9,  many  thousands 
(Hahn).  St.  George  Island,  1914 — April  8,  about  250  seen,  first  date 
when  observed  in  numbers,  though  a  few  had  been  about  all  winter; 
April  14,  upwards  of  10,000  seen,  small  numbers  flying  over  land 
and  large  flocks  along  beach  (Hanna).    St.  Paul  Island,  1915 — March 

30,  flocks  flew  along  nesting  ground;  April  15,  flocks  near  shore; 
April  24,  27,  thousands;  May  6,  abundant  (Hanna).  St.  George, 
1916 — April  16,  seen  at  sea;  April  25,  seen  on  land;  1917 — April  17, 
offshore;  April  23,  on  rocks  (|J.  P.  Adams).  St.  Paul,  1918— April 
10,  near  shore;  April  14,  large  flocks  over  bowlder  beaches  (Hanna). 

Eggs  are  usually  laid  in  June;  dates  of  collection  follow:  Otter 
Island,  June  14,  17,  1884  (Lutz,  1889,  p.  31).  St.  George,  May  28, 
1890;  St.  Paul,  June  5,  1890  (Palmer,  1899,  p.  388).  St.  George, 
May  24,  June  8,  1913,  June  16,  1914  (eggs  well  incubated)  (Hanna). 
St.  Paul,  July  2,  1914  (Preble).  St.  George,  June  5,  1916,  eggs  se- 
cured (H.  P.  Adams). 

After  the  close  of  the  nesting  season  in  July  the  species  rapidly 
becomes  scarce  about  the  shores  of  the  islands.  Hahn's  observations 
in  1910  show  records  as  follows :  August  30,  about  300  seen ;  August 

31,  50;  September  1,  8;  September  10,  1;  September  11,  1;  October 
2,  none  seen  on  St.  Paul  for  some  time,  but  still  numerous  at  sea.  In 
1913,  on  St.  George  Island,  Hanna  saw  none  about  the  land  later 
than  August  20.  In  1914,  on  St.  Paul,  I  noted  them  only  in  small 
numbers  during  August,  observing  a  few,  however,  almost  daily  up 
to  the  time  of  my  departure  on  August  30,  these  late  lingerers  being 
probably  the  late  hatched  birds.  After  my  departure  Hanna  re- 
corded 6  observed  on  October  4,  1,000  on  December  4,  and  500  on  De- 
cember 13.  H.  P.  Adams,  in  1916,  saw  the  last  about  St.  George  on 
August  28,  and  observed  it  at  sea  March  26,  1917.  Hanna  saw  one 
off  St.  George  January  25,  1917. 

Numerous  specimens  taken  by  Hahn  and  Marsh  in  1911  and  1912 
measured  in  the  flesh :  Length,  6f  to  7^  inches;  extent,  12|  to  13TV 

Food. — The  only  information  we  have  on  the  food  of  the  least  auk- 
let  is  H.  W.  Elliott's  credible  statement  (1875,  p.  209)  that  "they 
feed  on  water  shrimps  and  sea  fleas." 

Synthliboramphus  antiquus  (Gmelin).     Ancient  Murrelet. 

The  ancient  murrelet  was  first  recorded  from  the  Pribilofs  by  W. 
H.  Dall  (Dall  and  Bannister,  1869,  p.  310),  who  recorded  it  as  "  com- 
mon at  St.  George."  No  specimens  were  taken  at  that  time,  and  no 
further  particulars  as  to  the  basis  of  the  record  are  available.  The 
first  specimen  was  taken  by  W.  L.  Hahn  on  St.  Paul  Island,  Novem- 
ber 7,  1910;  this  is  now  in  the  collection  of  the  Biological  Survey. 
It  measured  in  the  flesh :  Length,  10  inches ;  extent,  16f . 


1923.]  BIRDS  OF    THE   PRIBILOF   ISLANDS.  25 

Hanna  observed  three  at  St.  Paul  on  December  4,  1914,  and  took 
a  specimen  there  on  April  18,  1918 ;  another  specimen,  a  female, 
taken  on  St.  George  Island,  May  12,  1916,  was  sent  in  by  Dr.  H.  P. 
Adams.  Both  of  these  are  now  in  the  collection  of  the  U.  S.  Na- 
tional Museum. 

Food. — The  stomach  of  the  April  specimen  above  mentioned  con- 
tained remains  of  more  than  800  amphipods  (Pontogeneia  inermis). 

Brachyramphus  marmoratus  (Gmelin).     Marbled  Murrelet. 

The  marbled  murrelet  was  first  recorded  from  the  Pribilofs  by 
Hanna  (1919a,  p.  176,  and  1920a,  p.  251),  on  the  basis  of  a  speci- 
men taken  by  him  on  St.  Paul  Island  January  13,  1918.  It  is  now 
No.  255110  of  the  collection  of  the  U.  S.  National  Museum. 

Food. — The  stomach  of  this  specimen  was  about  half  filled  with 
ground-up  amphipods.  These  constituted  100  per  cent  of  the  food, 
a  bit  of  hydroid  present  being  much  less  than  1  per  cent. 

Cepphus  columba  Pallas.     Pigeon  Guillemot. 

The  pigeon  guillemot  was  first  recorded  from  the  Pribilofs  by 
Nelson,  who  states  "they  nest  on  the  fur  seal  islands"  (1887,  p.  45). 
This  statement  was  based  on  conjecture,  from  his  having  observed 
the  birds  under  the  cliffs  of  St.  George  Island.  Palmer  found  the 
birds  quite  common,  but  very  shy,  on  the  ocean  to  the  southward  of 
Walrus  Island,  on  June  13,  1890  (1899,  p.  389). 

Hanna  has  published  the  following  note,  based  on  his  observation 
of  the  species  (1916,  p.  402)  : 

This  is  a  common  winter  resident,  and  by  April  18,  1915,  some  had  the 
black  summer  plumage  almost  complete  though  still  slightly  speckled  with 
white.  It  is  strange  that  this  bird  should  not  breed  on  the  Pribilofs,  since  it  is 
so  abundant  on  the  neighboring  Aleutians.  The  last  one  seen  in  1915  was 
on  May  26.  In  winter  it  feeds  along  the  surf  line,  is  very  tame,  and  utters  a 
low,  shrill  whistle  when  observed.  A  male  and  a  female  were  taken  on  each 
island.  The  species  has  been  previously  recorded  from  the  islands,  but 
apparently  no  specimens  were  secured  prior  to  my  work  there. 

His  detailed  notes  taken  on  St.  Paul  Island,  follow :  1914 — Decem- 
ber 2,  8  seen ;  December  9,  1 ;  December  4,  6,  and  13,  about  200  seen 
each  day.  1915 — -February  18,  200  seen  at  Northeast  Point;  April  6, 
1;  April  18,  12;  May  10,  <2;  May  18,  2;  May  26,  1. 

According  to  his  field  catalogue,  M.  C.  Marsh  collected  a  male  of 
this  species  on  St.  Paul  Island,  January  6,  1912,  and  made  the  fol- 
lowing notes:  Length,  13  inches;  extent,  23.  Legs  and  feet  orange 
red  in  front,  somewhat  darker  behind;  tarsus  black  behind;  claws 
black ;  mouth  vermilion ;  iris  brown. 

Hanna,  during  1913  to  1915,  observed  the  species  on  a  number  of 
occasions,  as  follows:  St.  George  Island,  1913 — December  19  and  30. 
St.  Paul  Island,  1914,  December  2,  8  seen ;  December  9,  1 ;  December 


26  NORTH   AMERICAN   FAUNA.  [No.  46 

4,  6,  and  13,  about  200  seen  each  day ;  1915 — February  18,  about  200 
seen  at  Northeast  Point.  During  the  same  spring  the  species  was 
noted  as  follows :  April  6, 1 ;  April  18,  12 ;  May  10,  2 ;  May  18,  2 ;  May 
26,  1.  Single  specimens,  now  in  the  U.  S.  National  Museum,  were 
collected  by  Hanna  on  the  following  dates;  St.  George  Island,  De- 
cember 19  and  30,  1913;  St.  Paul  Island,  December  6,  1914;  April  18, 
1915.  The  specimen  last  mentioned  has  nearly  finished  acquiring  the 
summer  plumage,  though  the  underparts  are  still  speckled  with  white. 
H.  P.  Adams  noted  the  species  on  St.  George,  February  4,  1917,  and 
Hanna  on  January  24,  1917.      / 

Food. — Of  the  seven  stomachs  of  the  pigeon  guillemot  from  the 
Pribilof  Islands  only  five  were  well-filled,  and  these  contained  the 
following  food  items:  Amphipods,  56.8  per  cent;  hermit  and  spider 
crabs,  20.8  per  cent ;  isopods,  17.2  per  cent ;  fish,  4  per  cent ;  kelp,  1  per 
cent ;  and  univalves,  0.2  per  cent.  The  amphipods  were  identified  as 
Allorchestes  malleolus  and  the  isopods  as  Idothea  ochotensis.  The 
spider  crab  was  Lithodes  brevipes,  and  the  hermit  crabs  include  two 
species — Hapalogaster  grebnitzkii  and  Dermaturus  mandtii.  No 
fewer  than  13  of  the  last-named  species  had  been  eaten  by  one  bird. 
The  fact  that  almost  no  univalve  fragments  were  present  in  the 
stomachs  of  birds  eating  so  many  hermit  crabs  shows  that  the  crabs 
are  either  pulled  from  the  empty  shells  they  use  for  homes  or  are 
caught  while  transferring  from  one  shell  to  another,  a  thing  they 
frequently  do.  The  fish  remains  in  the  present  series  of  stomachs 
were  unidentified,  and  the  miscellaneous  items  of  small  importance, 
mostly  contained  in  a  single  nearly  empty  stomach,  were  remains  of 
marine  worms,  a  pycnogonid  or  sea  spider,  and  bits  of  kelp. 

Uria  troille  calif ornica  (H.  Bryant).     California  Murre. 

The  California  murre  is  the  less  abundant  of  the  two  species  breed- 
ing on  the  Pribilofs,  occurring  in  some  numbers  among  the  nesting- 
colonies  of  the  Pallas  murre  in  practically  all  the  sites  on  the  islands, 
excepting  Sealion  Rock,  and  in  some  cases  occupying  extensive  areas 
to  the  exclusion  of  the  larger  species.  It  was  first  recorded  from  the 
Pribilofs  by  Elliott  (1874,  unpaged,  and  1875,  p.  210). 

It  is  difficult  to  fix  even  the  approximate  date  of  spring  arrival  on 
the  Pribilofs,  since  most  observers  have  failed  to  distinguish  the  two 
species,  and  the  case  is  further  complicated  by  the  fact  that  both 
species  occur  in  some  numbers  during  the  winter.  The  birds  usually 
begin  perching  on  the  cliffs,  probably  indicating  an  awakening  breed- 
ing interest,  in  late  April  or  early  May. 

The  bird  breeds  on  all  the  islands  of  the  group  excepting  Sealion 
Rock  (Hanna,  1920b,  p.  175).  Mr.  Hanna  states  that  on  St.  Paul 
this  species  nests  principally  at  Southwest  Point,  and  on  the  high 


1923.]  BIRDS   OF   THE   PRIBILOF   ISLANDS.  27 

cliffs  on  the  western  side  of  the  island,  among  the  colonies  of  the 
more  numerous  Pallas  murre,  but  that  it  is  not  found  among  the  ex- 
tensive murre  rookeries  on  Eeef  or  Kitovi ;  this  is  in  agreement  with 
my  own  observations  as  far  as  they  extend.  It  nests  in  great  num- 
bers on  Walrus  Island,  usually  apart  from  the  other  species.  This 
circumstance  has  caused  different  visitors,  judging  from  hurried  ob- 
servation covering  only  parts  of  the  rock,  to  believe  that  the  murre 
rookeries  there  consisted  mainly  of  one  or  the  other  species.  Palmer, 
however,  in  1890,  made  two  trips  to  the  rock,  and  noted  the  Pallas 
murre  exclusively  on  the  western  and  northern  parts  of  the  island, 
and  the  present  species  on  its  southeastern  part.  During  my  own 
visit  there  on  July  16,  1914,  I  noted  that  the  breeding  colonies  on  the 
higher  central  part  of  the  island  were  principally  of  the  present  spe- 
cies. Further  detailed  observations  are  needed  to  show  the  propor- 
tionate numbers  and  the  breeding  areas  of  the  two  species  on  Walrus 
Island,  and  these  may  vary  in  different  years. 

The  first  eggs  are  laid  in  early  June,  and  owing  to  various  causes 
eggs  still  being  incubated  may  be  found  until  near  the  end  of  July ; 
at  the  time  of  our  visit  there  on  July  16,  1914,  young  birds  and  eggs 
seemed  about  equally  numerous.  Eggs  were  taken  on  Otter  Island, 
June  13,  1884  (Lutz~  1889,  p.  31). 

The  birds  leave  the  breeding  rookeries  in  late  August  but  continue 
about  the  islands  during  the  autumn  and  winter,  being  by  far  the 
more  common  of  the  two  species  at  that  season,  and  being  called  the 
winter  arrie  by  the  natives,  many  of  whom  think  it  is  found  only 
at  that  season.  Winter  specimens  in  the  collections  of  the  Biological 
Survey  and  U.  S.  National  Museum  include  the  following :  St.  George 
Island,  male,  December  21,  1913 ;  male,  December  30,  1913 ;  St.  Paul 
Island,  December  6,  1914.  The  species  seems  usually  to  be  seen  only 
in  small  companies  during  the  winter.  However,  on  February  18, 
1915,  Hanna  observed  upwards  of  1,000  about  St.  Paul.  He  also 
found  the  birds  abundant  at  the  edge  of  the  ice-pack  off  St.  George, 
January  25,  1917. 

Food. — Eighteen  stomachs  of  the  California  murre  from  the  Prib- 
ilofs,  mostly  taken  in  winter,  have  been  examined.  Of  these  only 
12  were  well  filled  and  the  contents  of  these  were  almost  exclusively 
amphipods.  The  kind  most  frequently  identified  was  Pontogeneia 
sp. ;  Anonyx  nugax,  Allorchestes  ochotensis,  Hyale  sp.,  J  asset,  sp., 
and  Metopa  sp.  occurring  in  smaller  numbers.  One  of  the  well- 
filled  murre's  stomachs  contained  remains  of  20  or  more  marine 
worms  of  the  family  Nereidae,  and  two  of  them  held  isopods  {Ido- 
Ihea  ochotensis) .  The  nearly  empty  stomachs,  which  experience  has 
shown  do  not  as  a  rule  fairly  represent  normal  food  habits,  yielded 
remains  of  small  sculpins  (Cottidae),  bits  of  barnacles,  mollusks  (in- 


28  NORTH   AMERICAN   FAUNA.  [No.  46 

eluding  Aegobuccinum  oregonense) ,  and  hydroids;  and  the  following 
vegetable  matter:  Algae,  and  glumes  of  grasses   (Phleum  alpinum 
and  Festuca  ovina  var.  violacea). 
Uria  lomvia  arra  (Pallas).    Pallas  Murre. 

The  Pallas  murre  is  a  very  abundant  species,  the  more  numerous 
of  the  two  murres,  and  nests  on  all  the  islands  of  the  Pribilof  group. 
It  was  apparently  first  definitely  recorded  by  Dall  (Dall  and  Ban- 
nister, 1869,  p.  309),  though  Veniaminof  (Trans,  by  Elliott,  1874, 
unpaged,  and  1875,  p.  242),  in  referring  to  "arries"  as  occurring, 
referred,  of  course,  to  this  species  as  well  as  to  the  California  murre. 

Since  the  species  winters  in  small  numbers  about  the  Pribilofs  it 
is  impossible  to  give  definitely  its  usual  date  of  arrival  or  departure, 
especially  since  the  two  species  are  difficult  to  distinguish  at  a  little 
distance.  The  following  notes  taken  by  Hanna  in  1915  on  St.  Paul 
Island  evidently  refer  to  spring  arrival  of  this  species  on  the  nest- 
ing cliffs:  April  15,  150  killed  by  natives,  hundreds  reported  on  the 
cliffs  at  Southwest  Point ;  April  26,  100  killed  in  same  place ;  May  6, 
fairly  common.  On  St.  George,  in  1917,  Hanna  noted  the  first  of 
the  season  on  April  20. 

The  Pallas  murre  is  the  more  numerous  of  the  two  species  on 
all  the  nesting  rookeries.  Egg  laying  begins  in  early  June.  James 
Judge  (MS.  report)  states  that  two  arrie  eggs  were  found  on  Walrus 
Island  on  June  1,  1889,  and  that  the  birds  had  not  begun  to  lay  at 
the  same  place  on  June  8,  1904 ;  on  June  16,  1910,  932  fresh  eggs  were 
found  at  the  same  place;  on  June  21,  1909  (St.  Paul  Island  log) 
about  6,000  eggs  were  obtained  by  the  natives  at  the  same  place. 
(These  notes  may  refer  in  part  to  the  California  murre.)  Definitely 
identified  eggs  of  the  present  species  were  taken  as  follows:  St. 
George,  June  27,  July  1,  1873;  St.  Paul,  July  31,  1890  (Catalogue 
eggs,  U.  S.  Nat.  Mus.)  ;  Otter  Island,  June  13,  30,  1884  (Lutz,  1889, 
p.  31)  ;  Walrus  Island,  June  13,  1890  (Palmer,  1889,  p.  391) ;  St. 
George,  July  1,  1914  (18  eggs  in  collection  U.  S.  Nat.  Mus.  selected 
from  100,  to  show  variation  in  color,  by  G.  Dallas  Hanna) . 

By  the  end  of  August  most  of  the  birds  have  left  the  breeding 
rookeries;  at  this  time  many  late-hatched  young  are  deserted  and 
soon  perish,  the  desire  of  the  mother  to  accompany  the  departing 
flocks  evidently  being  stronger  than  the  parental  instinct.  Hanna 
states  that  on  August  31.  1913,  most  of  the  murres  had  gone,  and 
that  many  young  ones  were  falling  from  the  cliffs. 

A  few  are  observed  about  the  island  during  the  autumn  and 
winter ;  Hanna's  notes  for  1914  record  2  observed  on  October  4,  3  on 
December  4,  and  3  dead  ones  seen  December  13.  As  is  the  case  with 
several  others  of  the  more  abundant  species,  dead  ones  are  frequently 
observed  in  large  numbers.    Notes  taken  from  the  St.  Paul  Island 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  29 

log  by  W.  L.  Hahn  include  the  following  entries  in  the  autumn  of 
1900:  October  25,  hundreds  of  sick  young  ones;  November  2,  hun- 
dreds of  dead  birds;  November  10,  at  least  5,000  dead  on  the  shores 
of  the  lagoon ;  November  24.  both  sides  of  Northeast  Point  covered 
with  dead  ones. 

A  female  specimen  taken  by  M.  C.  Marsh,  February  5.  1912,  meas- 
ured in  the  flesh:  Length,  15^  inches;  extent,  28-|;  bill  jet-black,  ex- 
cepting the  border  of  light  yellow  at  base  of  the  upper  mandible  and 
the  pale  tip ;  legs  and  feet  black  posteriorly,  dusky  and  dark  yellow 
anteriorly. 

Food. — If  the  relatively  small  number  of  stomachs  available  can 
be  taken  as  a  criterion,  the  food  of  the  Pallas  murre  in  the  Pribilof 
region  differs  markedly  from  that  of  the  California  murre.  Of  six 
stomachs  examined,  four  were  well-filled,  and  the  food  in  them  con- 
sisted of  fish  remains,  48.75  per  cent;  flesh  and  mandibles  of  squids, 
26.25  per  cent ;  and  crustaceans,  25  per  cent.  The  nearly  empty  stom- 
achs also  contained  fish  and  squid  remains.  The  crustaceans  which 
completely  filled  one  stomach  were  shrimps  identified  as  Spironto- 
caris  polaris. 

Family  STERCORARIIDAE. 

Coprotheres  pomarinus  (Temminek).*    Pomabine  Jaeger. 

The  Pomarine  jaeger  is  a  somewhat  rare  visitor.  It  was  first 
obtained  by  Elliott,  who  took  a  specimen  on  the  uplands  between 
Kaminista  Lake  and  Polovina,  on  St.  Paul  (Elliott,  1874,  unpaged, 
and  1875,  p.  196).  This  is  No.  62522,  female  adult,  U.  S.  National 
Museum,  June  23,  1872.  Another  specimen  (No.  106857,  male,  June 
16,  1885)  was  obtained  and  reported  by  C.  H.  Townsend  (1887,  p  98). 

Palmer  (1899.  p.  393)  states:  "During  the  summer  of  1890  two 
were  seen  on  St.  George  eating  the  carcass  of  a  fur  seal.  One  of 
these  was  killed  and  preserved  by  Mr.  Ed.  Lavender."  The  final 
disposition  of  this  specimen  is  not  known.  Two  specimens,  a  male  and 
a  female,  were  taken  by  H.  J.  Christolfers  on  St.  Paul  Island,  June 
10,  1916.  These  are  now  in  the  collection  of  the  U.  S.  National  Mu- 
seum. 

Stercorarius  parasiticus  (Linnaeus).     Parasitic  Jaeger. 

The  parasitic  jaeger  was  first  added  to  the  fauna  of  the  islands 
by  Elliott,  who  mentions  seeing  four  or  five  individuals  (1874,  un- 
paged, and  1875,  p.  197).  A  specimen  taken  by  him,  a  female  in  the 
dark  phase  of  coloration,  collected  June  15,  1872,  is  No.  62524,  of 
the  bird  collection  of  the  U.  S.  National  Museum.    This  constituted 

*  Stercorarius  pomarinus  of  the  A.  0.  U.  check  list,  1910. 
108731°— 23 3 


30  NORTH   AMERICAN  FAUNA.  [No.  46 

the  only  record  until  1914,  when  G.  D.  Hanna  on  September  24  took 
a  dark  male  bird  on  St.  Paul  Island,  now  No.  237495,  U.  S.  National 
Museum.  Hanna  observed  another  individual,  also  a  dark-colored 
one,  on  June  14,  1915,  on  the  same  island. 

Food. — The  stomach  of  the  September  specimen  taken  by  Hanna 
was  filled  with  finely  ground  remains  of  small  fishes.  The  grinding 
material  it  contained  was  three  lumps  of  an  old  bryozoan  colony. 

Stercorarius  longicaudus  Vieillot.     Long-tailed  Jaeger. 

Judging  by  the  records  the  long-tailed  jaeger  is  the  most  fre- 
quently observed  of  the  three  jaegers,  and  we  may  perhaps  credit 
Veniaminof  (Trans,  by  Elliott,  1874  unpaged,  and  1875,  p.  242) 
with  first  having  observed  it,  since  he  records  "jaegers"  as  one  of 
the  species  occurring  on  the  Pribilof  Islands.  Elliott  took  one  June 
13,  1872,  which  has  been  several  times  recorded,  and  is  still  in  the 
collection  of  the  U.  S.  National  Museum.  Though  thus  early  noted, 
the  species  appears  not  to  have  been  afterwards  observed  until  recent 
years.  Hanna  shot  one  on  St.  George,  May  23, 1914.  This  was  a  male 
which  had  a  string  about  the  right  foot,  showing  previous  capture. 
Later  in  the  same  summer  Mr.  Hanna  and  the  writer  observed  a 
number  on  both  islands.  We  saw  it  daily  on  St.  George,  July  12, 
13,  and  14,  noting  as  many  as  7  on  the  two  latter  dates,  and  taking 
a  specimen  on  the  14th ;  I  noted  one  near  Kaminista  Lake,  St.  Paul, 
on  July  29.  The  next  year,  on  May  27,  Hanna  took  a  specimen  on 
St.  Paul. 

None  of  the  jaegers  are  known  to  have  bred  on  the  Pribilof s  nor 
to  have  remained  through  the  breeding  season,  although  the  condi- 
tions there  would  seem  to  be  favorable  for  nesting. 

Food. — Five  stomachs  of  the  long-tailed  jaeger  collected  on  the 
Pribilof  Islands  are  at  hand,  of  which  four  are  well  filled,  three 
of  them  with  insects,  and  one  with  the  remains  of  eggshell  and  an 
unhatched  young  of  a  deep-water  duck  (Fuligulinae).  One  of  the 
stomachs  containing  insects  held  more  than  90  individuals  of  4 
species.  The  most  numerous  species  was  the  large  gold  and  green 
ground  beetle  Caratus  truncaticollis,  of  which  one  stomach  contained 
58,  including  one  larva.  Other  ground  beetles  eaten  were  Pterosti- 
chus  pingruidineus,  Pterostichus  sp.,  and  Aviara  brunnipennis. 
One  stomach  contained  3  of  the  rather  large  yellow  and  brown 
moths  (Byphoraia  subnebulosa) ,  one  a  crane-fly  (Tipula  sp.)  with 
eggs,  one  two  flesh-flies  {Cynomyia  hirta)  and  16  larvae  of  this  or 
an  allied  species,  and  another  a  small  parasitic  wasp  (Amblyteles 
sp.).  A  bit  of  fish  bone  was  in  one  of  the  well-filled  stomachs  and 
a  nearly  empty  stomach  contained  a  single  sawfly  {Amauronematus, 
probably  of  the  species  wkitneyi,  described  from  material  in  the 
present  collections). 


1923.]  BIRDS   OF    THE   PBIBILOF   ISLANDS.  31 

Elliott  (1874,  unpaged,  and  1875,  p.  198)  states  that  two  indi- 
viduals seen  by  him  on  July  29,  1872,  [on  St.  Paul]  were  apparently 
feeding  upon  insects,  and  upon  a  small  black  berry  (Empetrum 
nigrum). 

Family  LARIDAE. 

Pagophila  alba  (Gunnerus).     Ivory  Gull. 

An  ivory  gull  obtained  by  True  and  Prentiss  in  1895  was  first 
recorded  by  Palmer  (1899,  p.  395).  Concerning  it,  Prentiss  (1902,  p. 
99)  says:  "One  was  killed  by  the  natives  on  St.  Paul  in  February 
or  March,  1895,  and  prepared  by  Mrs.  Judge,  from  whom  I  obtained 
it.  It  was  not  previously  known  from  the  Islands."  Other  specimens 
were  taken  on  St.  George  Island  by  A.  H.  Proctor  on  February  18, 
1916,  and  by  G.  Dallas  Hanna  on  March  27.  1917.  All  these  speci- 
mens are  in  the  collection  of  the  U.  S.  National  Museum. 

Rissa  tridactyla  pollicaris  Ridgway.     Pacific  Kittiwake.  PI.  VI.  fig.  3. 

Rissa  tridactylus  pollicaris  Ridgway,  Water  Birds  North  Amer.,  vol.  2,  p. 
202,  1884.     (Type  from  St.  George  Island.) 

The  Pacific,  or  western  black-legged,  kittiwake  is  very  abundant 
on  the  Pribilofs,  nesting  on  the  cliffs  of  all  five  islands  of  the  group, 
and  remaining  in  small  numbers  during  the  autumn  and  early 
winter.  The  species  was  first  recorded  from  the  islands  by  Coinde 
(1860,  p.  401),  and  has  been  observed  by  all  later  students  of  the 
avifauna.    It  is  eaten  in  large  numbers  by  the  natives. 

The  following  notes  on  spring  arrival  are  the  only  definite  ones 
at  hand:  St.  Paul  Island,  1909,  first  reported  April  20;  hundreds 
May  1  (St.  Paul  Island  log,  fide  Hahn).  St.  Paul  Island,  1911,  2 
seen,  April  24;  many  shot,  April  30  (Hahn).  St.  George  Island,  1914, 
2  seen  April  8;  10  seen  April  22  (Hanna).  St.  Paul  Island,  1915, 
May  18,  40  killed  at  Northeast  Point;  May  28,  abundant  (Hanna). 
St.  George,  April  16,  1917  (Hanna). 

The  nests  are  built  of  grass  and  moss,  with  some  mud,  and  are 
placed  on  small  platforms  on  the  faces  of  cliffs.  The  following  dates 
will  show  the  period  of  nesting :  St.  Paul,  July  3,  1879 ;  St.  George, 
June  24,  1873  (catalogue  eggs,  U.  S.  Nat.  Mus.).  Otter  Island,  June 
21,  1884,  eggs  taken  (Lutz,  1889,  p.  31).  Palmer  (1899,  p.  396)  gives 
the  following  pertinent  data :  St.  George,  May  28,  1890,  a  few  nests, 
but  no  eggs;  Walrus  Island,  June  8,  1889,  two  eggs  collected  by  C.  H. 
Townsend;  St.  Paul  Island,  August  2,  1890,  most  nests  with  young, 
but  a  few  had  a  young  one  and  an  egg,  or  sometimes  two  or  three 
eggs.  In  1914,  on  St.  George  Island,  Hanna  noted  that  on  July  1 
the  nests  were  being  completed,  and  a  few  had  eggs.  During  the 
same  season,  the  writer  found  a  few  nests  with  eggs  on  Sealion 
Hock,  on  June  29,  and  saw  many  nests  on  Walrus  Island  on  July  16, 


32  FORTH  AMERICAN  FAUNA.  [No.  46 

and  on  Otter  Island,  July  27.  Bent  (1921,  p.  49)  gives  13  records  for 
eggs  on  the  Pribilof s  from  June  10  to  July  7 ;  of  these,  7  records  fell 
between  June  25  and  July  3. 

Hahn  recorded  this  species  as  common  on  St.  Paul  Island  during 
September  and  October,  1910,  and  saw  quite  a  number  in  November. 
He  saw  12  on  December  8,  and  noted  it  as  numerous  December  21  of 
the  same  year,  and  saw  6  individuals  on  January  4,  1911.  Hanna,  on 
St.  Paul  Island,  during  the  autumn  of  1914,  observed  the  species  as 
follows:  September  14,  40;  September  16,  10;  September  20,  2;  Sep- 
tember 22,  100 ;  October  4,  25 ;  October  12,  3. 

A  female  specimen  taken  by  W.  L.  Hahn,  August  31, 1910,  measured 
in  the  flesh :  Length,  15|  inches ;  extent,  39. 

Food. — Three  stomachs  of  St.  Paul  specimens  of  the  Pacific  kitti- 
wake  (taken  in  July  and  August)  have  been  examined  but  yield 
meager  data  on  the  food  habits  of  the  species.  One,  very  well  filled, 
contained  fish  flesh,  with  no  bones,  and  therefore  not  identifiable, 
another  held  fragments  of  Crustacea,  also  unidentifiable,  and  the 
third  only  vegetable  debris,  evidently  algae.  A  more  valuable  stom- 
ach content  collected  at  St.  Matthews  Island,  July  12,  1916,  by  G. 
Dallas  Hanna,  contained  Crustacea  exclusively:  A  spider  crab,  Li- 
thodidae,  50  per  cent;  and  isopods,  including  Idothea  ochotensis,  50 
per  cent. 

Regarding  the  kittiwakes  Mr.  Hanna  says : 

They  both  occasionally  eat  small  fishes  which  come  near  the  surface  of  the 
sea.  At  such  times  they  dive  down  like  a  tern  for  their  prey.  They  also  eat  the 
refuse  from  freshly  cleaned  fish  which  may  be  thrown  into  the  sea,  but  I  have 
never  seen  one  eat  carrion. 

However,  the  record  above  of  fish  flesh  without  bones  is  very  sug- 
gestive that  carrion  sometimes  is  eaten. 

Kissa  brevirostris  (Bruch).     Red-legged  Kittiwake. 

The  red-legged  kittiwake,  an  exceedingly  beautiful  species,  the 
"Goverooskie"  of  the  natives,  is  much  less  abundant  than  its  congener, 
but  is  nevertheless  found  in  large  numbers.  It  breeds  on  all  the 
islands  excepting  Sealion  Rock,  and  appears  to  leave  the  neighbor- 
hood of  the  Pribilofs  in  September,  to  remain  until  April.  It  is  a 
favorite  article  of  food  of  the  natives,  being  more  esteemed  than  the 
black-legged  species.  It  was  first  credited  to  the  islands  by  Coinde, 
who  supposed  it  to  be  an  undescribed  species,  and  named  it  Laru* 
warnecki  (1860,  p.  401). 

Hahn  noted  that  many  were  shot  on  St.  Paul  Island  on  April  30, 
1911.  All  the  rest  of  the  available  notes  on  spring  arrival  are  by 
Hanna.  On  St.  George  Island,  in  1914,  he  observed  4  on  April  8, 50  on 
April  22,  and  upwards  of  1,000  on  May  6.  His  notes  for  1915,  on  St. 
Paul  Island,  are  as  follows :  April  18,  20  individuals  observed ;  April 


1923.]  BIRDS  OF   THE  PBIBILOF  ISLANDS.  33 

24,  6 ;  April  26,  81  killed  by  natives  at  Southwest  Point ;  May  4,  30  ob- 
served, abundant  at  Southwest  Point ;  May  18,  100  observed  at  North- 
east Point ;  May  28,  abundant.  In  1917,  on  St.  George,  he  noted  the 
first  on  April  22. 

The  red-legged  kittiwake  nests  among  the  colonies  of  its  more  nu- 
merous relative.  Prentiss  (1902,  p.  99)  makes  the  following  state- 
ment regarding  its  detailed  distribution  on  St.  Paul  Island,  from  ob- 
servations made  in  the  summer  of  1895 : 

The  distribution  of  the  red-leg  on  the  island  of  St.  Paul  differs  from  that  of 
the  black-leg.  Around  the  village  and  the  southeast  portion  of  the  island  the 
red-leg  is  comparatively  rare,  while  at  Half-way  Point  on  the  east  side  of  the 
island,  there  is  a  marked  increase  in  numbers.  At  the  southwestern  portion 
they  form  nearly  one-half  of  the  kittiwakes.  *  *  *  On  the  north  side  of 
St.  Paul  they  were  numerous,  and  also  on  the  beach  at  Zapadnie. 

Palmer  (1899,  p.  398)  gives  the  measurements  of  eggs  taken  on 
St.  George  Island,  by  Elliott,  June  25,  1872.  Lutz  (1889,  p.  31) 
records  an  egg  taken  on  Otter  Island,  June  29,  1884.  The  catalog 
of  eggs  in  the  U.  S.  National  Museum  notes  eggs  taken  on  St.  Paul, 
July  15,  1895.  Hanna  took  eggs  on  St.  George  Island,  July  3,  1914. 
On  August  31,  he  observed  young  still  in  the  nest;  on  September  11, 
the  young  were  all  on  the  wing.  This  apparently  was  the  signal 
for  the  departure  of  the  species.  On  September  14,  he  records  seeing 
12  individuals;  on  the  16th,  4,  and  on  the  22d,  10.  These  were  the 
last  seen  that  season.    I  find  no  winter  records. 

Food. — Fifteen  stomachs  of  the  red-legged  kittiwake  from  St. 
George  Island  have  been  examined,  eight  of  them  being  half  or 
more  full  and  seven  nearly  empty.  Of  the  food  in  the  former,  25 
per  cent  consisted  of  squid  remains,  37.5  per  cent  of  fish,  and  37.5 
per  cent  of  Crustacea.  The  Crustacea  were  small  shrimplike  forms, 
chiefly  of  the  genus  Tkysanoessa,  and  were  present  to  the  number 
of  several  hundred  in  each  of  two  stomachs.  All  of  the  nearly  empty 
stomachs  contained  squid  mandibles. 

Larus  hyperboreus  hyperboreus  Gunnerus.     Glaucous  Gull. 

The  glaucous  gull  breeds  in  small  numbers  on  the  grassy  summit 
of  Walrus  Island,  among  a  large  colony  of  glaucous-winged  gulls, 
and  during  the  summer  season  may  occasionally  be  observed  on  the 
other  larger  islands.  It  is  found  in  larger  numbers  during  the  fall, 
and  Hanna  (1917,  p.  407)  states  that  in  winter  it  is  the  only  large 
gull  found  on  the  Pribilofs,  having  replaced  entirely  the  glaucous- 
winged  gull  with  which  it  has  been  associated. 

The  species  was  first  recorded  by  Palmer  (1899,  p.  395)  under  the 
name  Larus  oarrovianus.  A  specimen  taken  by  him  there  on  June 
13,  1890,  has  recently  been  referred  by  Oberholser,  as  a  result  of  a 
critical  study  of  the  group,  to  the  typical  form  (1918,  p.  471). 


34  NORTH   AMERICAN  FAUNA.  [No.  46 

On  the  occasion  of  my  visit  to  Walrus  Island  on  July  16,  1914,  a 
few  glaucous  gulls,  about  15  pairs,  were  found  to  be  nesting  on  the 
extreme  grassy  summit  of  the  island.  At  this  time  most  of  the 
nests  contained  well-grown  young.  No  specimens  were  collected, 
and  owing  to  the  brief  time  we  could  stay  on  the  island,  no  detailed 
study  could  be  made  of  this  or  any  particular  species. 

Hanna  observed  the  glaucous  gull  on  St.  Paul  Island,  as  follows: 
1914,  September  14,  1 ;  September  16,  5 ;  September  20,  1 ;  October 
12,  1 ;  October  25,  20 ;  November  27,  10 ;  December  13,  4.  1915  :  Feb- 
ruary 18,  25;  March  5,  2;  March  18,  300;  April  2,  1;  April  24,  10; 
May  4,  30 ;  May  28,  15. 

The  species  evidently  was  common  about  St.  George  Island  in  the 
spring  of  1917,  since  Hanna  sent  in  the  stomachs  of  nine  birds  taken 
on  April  12. 

Food. — Nine  stomachs  of  glaucous  gulls  from  St.  George  Island 
and  one  from  St.  Matthew  Island,  all  well  filled,  were  available  for 
the  present  study.  They  reveal  that  this  species  shares  the  well- 
known  propensity  of  gulls  to  feed  largely  on, carrion.  Fifty-eight 
per  cent  of  the  total  food  of  these  birds  was  fishes,  the  indications 
being  that  most  if  not  all  of  them  were  found  dead.  Ten  per  cent 
of  the  food  consisted  of  a  large  egg,  perhaps  of  a  duck,  found  in  one 
stomach,  another  10  per  cent,  approximately,  of  squid  remains,  a 
slightly  larger  amount  of  invertebrates  including  hydroids,  mollusks, 
marine  worms,  and  a  crab,  and  a  similar  proportion  of  algae.  The 
mollusks  identified  were  the  common  blue  mussel  {Mytilus  edulis), 
a  sea  slug  (Chiton  (?)  submarmorea) ,  and  the  crab  was  a  hermit 
(Dermaturus  mandtii). 

Larus  hyperboreus  barrovianus  Ridgway.     Point  Barrow  Gull. 

A  gull  taken  on  St.  Paul  Island  on  June  21,  1890,  has  been  referred 
by  H.  C.  Oberholser  (1918,  p.  473)  to  this  form,  which  is  character- 
ized as  smaller,  and  having  a  darker  mantle,  than  typical  L.  hyper- 
boreans. This  bird  must  have  been  a  nonbreeding  bird  or  a  wan- 
derer from  some  breeding  colony.  It  was  taken  by  William  Palmer, 
but  for  some  reason  was  not  mentioned  in  his  account  of  the  birds 
of  the  Pribilofs.  The  specimen  is  still  in  the  collection  of  the  U.  S. 
National  Museum. 

Larus  glaucescens  Naumann.     Glaucous-winged  Gull. 

The  glaucous-winged  gull  is  an  abundant  summer  resident,  breed- 
ing on  Walrus  Island,  St.  George  Island,  and  Sealion  Rock,  and  is 
also  observed  in  winter,  sometimes  in  considerable  numbers.  First 
recorded  from  the  Pribilofs  by  Veniaminof  in  1840  (Trans,  by  El- 
liott, 1874,  unpaged,  and  1875,  p.  242),  under  the  name  of  "chickee" 
(the  native  name),  and  noted  by  all  subsequent  observers. 


1923.]  "BIRDS   OF   THE   PRIBILOF   ISLANDS.  35 

Because  of  its  wintering  to  some  extent,  the  date  of  spring  arrival 
is  somewhat  uncertain.  The  following  dates  of  observation  later 
than  late  March  are  available:  St.  George  Island,  1914:  March  28, 
25  seen;  May  6,  30  seen;  April  14,  40  about  stranded  whale;  April 
22,  20.  St.  Paul  Island,  1915 :  April  24,  about  3,000  seen,  appeared 
in  numbers  during  the  last  few  days;  May  4,  20  seen;  May  28, 
abundant. 

The  birds  breed  principally  on  Walrus  Island,  the  foxes  appar- 
ently preventing  them  from  nesting  on  St.  Paul  Island  and  Otter 
Island,  and  on  St.  George  except  in  a  few  places,  as  the  birds  prefer 
flat  turfy  situations  for  nesting.  A  few  pairs  breed  on  the  summit 
of  Sealion  Rock,  a  small  islet  close  to  St.  Paul.  All  through  the 
breeding  season  the  birds  are  common  on  St.  Paul  Island,  feeding 
about  the  shores  and  numerous  ponds,  and  apparently  securing  most 
of  their  nesting  material  there.  On  Walrus  Island,  in  1872,  Elliott 
considered  that  500  or  600  were  nesting;  in  1914  I  estimated  about 
100  pairs.  The  eggs,  usually  2  or  3  in  number,  are  laid  mainly  in 
June.  James  Judge  reported  finding  359  eggs,  all  fresh,  on  Walrus 
Island,  June  16,  1910.  Palmer  (1899,  p.  394)  on  June  13,  1890, 
found  some  fresh  eggs  and  many  well  incubated,  while  a  few  were 
already  hatched.  At  the  same  place  on  July  16,  1914,  I  found  many 
young,  but  only  a  few  eggs,  and  those  usually  about  to  hatch. 

A  few  fall  and  winter  dates  of  observation  follow :  St.  Paul 
Island,  1910,  abundant  during  September  and  October;  less  so  in 
November.  December  8,  12  seen;  December  21  numerous  at  sea; 
1911,  January  4,  3  seen;  January  30,  6  (Hahn).  St.  George:  1913, 
September  4,  8  seen;  September  15,  about  20;  October  9,  4;  October 
18,  50;  November  11,  a  few  along  beach  all  the  time;  1917,  January 
25,  abundant  (Hanna).  M.  C.  March  made  the  following  notes  on 
fresh  specimens  collected  by  him :  St.  Paul  Island,  October  16,  1911, 
female,  length,  23|  inches ;  extent,  53 ;  basal  half  of  bill  whitish, 
scarcely  tinged  with  flesh  color;  terminal  part,  black,  excepting  pale 
tip ;  legs  and  feet  outwardly  dirty  brownish,  paler  on  inner  surface. 
Another  female  specimen,  same  place  and  date,  measured :  Length, 
24  inches;  extent,  52;  iris  purplish-silvery  mottled;  edges  of  eyelids 
pinkish;  bill  whitish  and  yellow,  with  a  blotch  of  orange  at  angle; 
lower  mandible  yellow,  deepest  on  curves;  upper  mandible  lighter 
along  straight  edges  of  commissure  and  above  nostrils ;  legs  and  feet 
faded  pinkish. 

Food. — William  Palmer  in  his  report  on  Pribilof  birds  (1899,  p. 
395)  noted  that  the  glaucous-winged  gull  feeds  "on  pretty  much 
everything  in  the  way  of  offal.  Even  the  dead  seals  are  devoured 
by  them,  and  they  vie  with  the  foxes  in  their  ability  to  search  out 
and  dispose  of  all  animal  matter."     Results  obtained  in  the  present 


36  NORTH   AMERICAN   FAUNA.  [No.  46 

investigation  bear  out  this  bird's  reputation  as  a  scavenger.  Three 
of  the  four  well-filled  stomachs  contained  fish  bones  and  flesh  and 
eggs,  undoubtedly  carrion,  which  amounted  to  41.25  per  cent  of  the 
total  food.  A  rib  of  a  fur  seal  was  the  principal  food  taken  by  one 
bird;  the  bone  was  6  inches  long  and  extended  from  the  throat  to 
the  stomach,  where  the  lower  end  was  undergoing  digestion.  One 
full  and  one  nearly  empty  stomach  contained  exclusively  remains  of 
eggs  and  young  of  a  bird  of  the  auklet  family.  Miscellaneous  con- 
tents were  remains  of  univalve,  sea  urchin,  crab,  and  algae. 

Larus  schistisagus  Stejneger.     Sla't/st-backed  Gull. 

The  slaty-backed  gull  was  included  among  the  birds  of  the 
Pribilofs  by  Palmer,  whose  account  (1899,  p.  394)  may  be  quoted 
in  full: 

Several  of  the  natives  informed  me  that  a  large  black-backed  gull,  different 
from  the  following,  bred  sparingly  on  the  cliffs  of  Otter  Island,  and  that  they 
rarely  visited  St.  Paul.  Daniel  Webster,  an  old  experienced  sealer  who  had 
then  spent  22  years  on  the  islands,  also  told  me  that  a  large  dark-backed  gull  was 
to  be  found  in  small  numbers  on  the  cliffs  of  St.  George.  On  June  11,  on  St. 
Paul  I  saw  three,  and  on  June  12,  another,  which  came  in  over  the  Reef  from 
the  direction  of  Otter  Island.  Several  evenings  after  August  1  I  noticed  some 
large  gulls  flying  slowly  in  from  the  Reef  (on  one  evening  I  counted  seven). 
The  mantle  was  much  darker  than  in  glaucescens.  They  were  probably  this 
species,  the  status  of  which  is  very  uncertain  on  the  American  side  of  Bering 
Sea. 

Gulls  observed  by  Hanna  on  two  occasions  were  apparently  of 
this  species.  On  December  21,  1913,  he  saw  about  25  large  white- 
tailed  gulls  with  slate-colored  backs,  off  St.  George  Island;  and  on 
October  15,  1919,  saw  two  dark-backed  gulls,  with  black-tipped 
wings,  and  slightly  smaller  than  the  glaucous-winged  gull,  about  the 
ship's  anchorage  off  St.  Paul  Island.  He  has  no  evidence  of  the 
breeding  of  the  species  on  Otter  Island. 

Rh.odostetb.ia  rosea  (Macgillivray).     Rosy  Gull. 

A  specimen  of  the  beautiful  rosy  gull  was  shot  by  a  native  on  a 
fresh-water  lake  on  St.  George  Island  May  25,  1911,  and  was  pre- 
served by  A.  H.  Proctor.  The  specimen,  which  is  now  in  the  col- 
lection of  the  Biological  Survey,  was  recorded  by  Evermann  (1913. 
p.  17).  A  second  specimen,  a  female,  taken  on  St.  George  March 
24,  1917,  is  now  in  the  U.  S.  National  Museum. 

Xema  sabinii  (J.  Sabine).     Sabine  Gull. 

The  Sabine  gull  appears  to  be  of  regular  occurrence  in  early  sum- 
mer and  in  the  autumn.  It  was  first  credited  to  the  islands  by 
Palmer  (1899,  p.  398),  who  mentions  one  seen  on  St.  Paul  Island 
by  H.  W.  Elliott  June  4, 1890,  and  one  (perhaps  the  same  individual) 
seen  later  about  the  lagoon  by  himself.     He  also  lists  a  specimen 


1923.]  BIKDS  OF   THE   PRIBIL.OF   ISLANDS.  37 

taken  June  26,  1890,  and  refers  to  others  taken  during  the  summer  of 
1896,  and  one  on  St.  George  Island  by  D.  W.  Prentiss.  Prentiss,  re- 
ferring to  observations  made  in  1895,  says  (1902,  p.  100)  :  "I  saw 
them  occasionally  on  both  St.  Paul  and  St.  George,  and  secured 
several." 

W.  L.  Hahn,  during  the  autumn  of  1910,  observed  the  species  as 
follows  on  St.  Paul  Island:  August  31,  1;  September  1,  7;  Sep- 
tember 7,  2;  September  8,  20;  September  11,  20;  October  2,  1.  A 
specimen  taken  by  him  on  September  8  measured  in  the  flesh: 
Length,  13|  inches ;  extent,  30f . 

G.  Dallas  Hanna,  on  St.  George  Island,  observed  two  individuals, 
collecting  one,  May  19,  1914,  and  on  June  16  saw  three.  In  the  year 
following,  on  St.  Paul  Island,  he  saw  one  on  May  26.  Two  speci- 
mens, an  adult  and  an  immature  female,  were  collected  by  him  on 
St.  Paul  Island,  August  30,  1916.  They  are  now  in  the  collection 
of  the  U.  S.  National  Museum. 

Food. — One  well-filled  stomach  and  crop  of  the  Sabine  gull  taken 
May  19,  1914,  on  St.  George,  gives  evidence  that  this  bird  feeds  upon 
insects  on  the  wing  and  is  skillful  in  the  pursuit.  The  food  con- 
tained in  this  stomach  included  the  remains  of  6  gold  and  green 
ground  beetles  (Carabus  truncaticollis) ,  2  other  ground  beetles 
{Pterostichus  sp.),  and  1  water  beetle,  together  4  per  cent;  a  great 
many  caddisflies,  16  per  cent;  and  numerous  two-winged  flies  (37 
Borborws  annidatus,  5  Leria  fruterna,  3  Scatophaga  dasythrix,  1 
Hydrophorus  innotatus,  and  27  or  more  others),  80  per  cent.  A 
nearly  empty  stomach  taken  at  the  same  time  contained  remains  of 
amphipods,  and  marine  worms  (Nereidae). 

Sterna  paradisaea  Brunnich.     Arctic  Teen. 

The  arctic  tern  is  apparently  of  regular  occurrence  about  the 
Pribilofs  during  migration.  It  was  first  recorded  from  the  group 
by  Palmer  (1899,  p.  398)  on  the  authority  of  F.  A.  Lucas,  who  saw 
two  on  St.  Paul  Island  July  26,  1890,  and  of  Daniel  Webster,  an  old 
sealer,  who  described  a  tern  claimed  to  be  found  every  fall  about 
St.  George.  Later  observations  have  added  to  the  records.  Hahn 
observed  eight  on  St.  Paul  Island  August  30,  1910,  and  one  on 
September  24.  Hanna  saw  four  feeding  in  the  surf  off  St.  George 
Island  May  19,  1914,  and  collected  one.  During  the  following- 
spring,  on  St.  Paul,  he  observed  the  species  as  follows :  May  9,  two : 
May  11,  four;  May  15,  one;  May  26,  one;  June  30,  one.  Two  female 
specimens,  an  adult  and  an  immature,  now  in  the  collection  of  the 
U.  S.  National  Museum,  were  taken  by  Hanna  on  St.  Paul  Island  on 
August  28  and  30.  1916.  He  also  took  a  specimen  on  St.  George, 
August  5,  1920. 


38  NORTH   AMERICAN   FAUNA.  [No.  46 

Food. — Only  two  stomachs  of  the  arctic  tern  from  the  Pribilofs 
(May  19,  1914,  and  August  5,  1920)  have  been  examined;  one  of  them 
was  practically  filled  with  remains  of  small  sculpins,  the  other  with 
amphipods  (46  or  more  Pontogeneia  sp.).  Both  stomachs  contained 
traces  of  univalves,  in  each  case  less  than  1  per  cent. 

Family  DIOMEDEIDAE. 

Diomedea  albatrus  Pallas.     Short-tailed  Albatross. 

The  short-tailed  albatross  used  to  be  of  common  occurrence  about 
the  Pribilof  Islands  according  to  the  testimony  of  the  natives. 
Veniaminof,  writing  in  1840  (Trans,  by  Elliott,  1874  unpaged,  and 
1875,  p.  242)  stated  that  the  albatross  "  is  frequently  to  be  seen  about 
the  beaches. "  Elliott  took  a  specimen  August  12,  1873,  and  was 
informed  by  the  natives  that  20  years  earlier  the  birds  had  been  very 
abundant  but  had  disappeared  with  the  whalers.  Palmer  (1899,  p. 
381)  obtained  an  immature  bird  on  St.  Paul  on  August  4,  1890,  and 
observed  five  others  on  August  10,  off  the  reefs  of  the  same  island. 

I  have  no  recent  records,  excepting  that  of  a  sternum  picked  up  on 
the  beach  at  St.  Paul  by  Hanna. 

Family  PROCELLARIIDAE. 

Fulmarus  glacialis  rodgersi  Cassin.     Rodgers  Fulmar. 

The  Rodgers  fulmar  is  a  rather  common  summer  resident,  but  the 
greater  number  apparently  leave  the  islands  as  soon  as  breeding  is 
over.  A  few  are  seen  during  the  autumn,  but  they  are  very  rare 
later.  The  species  lays  its  single  egg  on  narrow  shelves  on  steep 
cliffs  on  St.  Paul,  Otter,  and  St.  George,  by  far  the  greater  number 
on  the  last.  It  was  first  recorded  from  the  Pribilofs  by  Dall,  who 
procured  a  specimen  on  St.  George  (1869,  p.  323). 

The  birds  usually  arrive  in  March  or  April;  I  have  few  definite 
dates.  On  St.  George  Island  in  1914  Hanna,  on  March  21,  recorded 
having  seen  large  numbers  during  the  two  preceding  days,  "  the 
first  since  January,  when  one  was  seen  " ;  April  3,  numbers  reported 
at  Zapadni;  April  8',  30  observed;  April  22,  20.  In  1917  it  was  first 
seen  at  Zapadni  on  March  8.  Hanna's  notes  for  St.  Paul  are  as  fol- 
lows: 1915 — May  2,  200  reported  at  Southwest  Point;  May  4,  hun- 
dreds; May  10,  abundant.  1918 — February  8,  killed  at  sea  near 
island;  large  numbers  on  high  cliffs. 

The  following  dates  refer  to  nesting:  Otter  Island,  June  10,  1872, 
St.  George,  June  10,  1873,  eggs  in  U.  S.  National  Museum,  collected 
by  Elliott.  Otter  Island,  June  14,  1885,  eggs  taken  by  C.  H.  Town- 
send  (Palmer,  1899,  p.  382).  St.  George  Island,  July  1,  1914,  eggs 
taken  by  Hanna.    On  August  4, 1914,  we  found  nests  with  well  grown 


1923.]  BIRDS   OF   THE   PMBILOF   ISLANDS.  39 

young  on  the  cliffs  near  Zapadni,  St.  George.  H.  P.  Adams  secured 
eggs  on  St.  George,  June  4,  1916. 

Hahn  found  fulmars  common  at  sea  between  Walrus  Island  and 
St.  Paul,  October  2,  1910.  Hanna  saw  a  few  on  St.  George  Island, 
September  11,  1913,  and  two  on  September  28.  On  St.  Paul,  in  1915, 
he  noted  two  on  October  4. 

A  considerable  number  of  birds  in  the  sooty  phase  of  plumage 
occur  in  the  breeding  colonies.  Hanna  thinks  that  on  St.  George 
Island  about  one-fifth  are  dark  birds.  He  has  preserved  two  speci- 
mens of  the  dark  phase  of  this  species,  and  states  (1920a,  p.  249)  : 
"  These  birds  are  often  seen  in  the  large  colonies  which  breed  on  the 
Pribilofs.  They  seem  to  mate  indiscriminately  with  light  colored 
birds  and  in  one  case  a  slate-colored  downy  young  was  seen  which 
had  light-colored  parents."  On  July  12,  1914,  in  a  breeding  colony 
near  the  village  of  St.  George,  I  saw  a  light  and  a  dark  bird,  appar- 
ently mates,  presiding  over  a  nest  containing  a  young  one. 

Food. — Of  19  stomachs  of  the  Rodgers  fulmar  collected  on  St. 
George  Island,  18  were  nearly  empty,  but  whether  empty  or  well- 
filled  the  entire  food  remains  present  were  those  of  squids.  In  one 
case  mandibles  representing  at  least  13  squids  were  present.  G. 
Dallas  Hanna  also  recognized  the  importance  of  squids  in  the  dietary 
of  this  bird  and  gives  the  following  interesting  account  of  another 
food  habit  of  the  species : 

Some  species  of  squid  is  eaten  extensively  and  the  eyes,  which  resemble 
pearls,  are  found  in  the  stomachs  and  the  beaks  have  also  been  secured.  But 
the  huge  brown-rayed  jellyfish  found  in  Bering  Sea  also  is  eaten.  In  the  tide 
rips  and  eddies  about  the  Pribilofs  this  jellyfish  is  often  brought  to  the  surface 
of  the  water.  Fulmars  attack  it  voraciously  when  an  opportunity  is  presented 
and  if  possible  tear  it  to  pieces.  The  birds  then  gather  around  in  large  num- 
bers to  enjoy  the  feast.  It  seems  that  the  attacks  of  the  birds  finding  the 
jellyfish  are  chiefly  an  endeavor  to  so  injure  the  animal  that  it  will  be  unable 
to  dive  to  safety.  I  have  seen  as  many  as  50  birds  in  one  small  circle  eating 
these  jellyfish,  and  I  have  never  seen  anything  else  being  captured  or  eaten 
although  in,  the  course  of  a  day's  fishing  many  thousands  of  the  birds  are  seen 
cruising  over  the  ocean. 

Puffinus  tenuirostris  (Temminck).    Slender-billed  Sheabwater. 

Hanna  (1919a,  p.  176,  and  1920a,  p.  251)  added  the  slender-billed 
shearwater  to  the  Pribilof  fauna,  and  writes  as  follows : 

A  female  slender-billed  shearwater  was  picked  up  on  the  beach  of  St.  Paul 
Island  on  June  4,  1918.  Another  bird  was  seen  shortly  after,  flying  about  2 
miles  out  at  sea.  Mr.  C.  E.  Crompton  told  me  that  numerous  individuals  were 
in  the  vicinity  of  St.  George  Island  at  about  the  same  time.     (1.  c,  p.  251.) 

This  specimen  is  now  No.  255111  of  the  bird  collection  of  the  U.  S. 
National  Museum. 


40  NORTH   AMERICAN   FAUNA.  [No.  46 

Cceanodroma  furcata  (Gmelin).     Fork-tailed  Petrel. 

The  fork-tailed  petrel  has  been  observed  on  a  number  of  occa- 
sions about  the  Pribilofs  during  the  summer,  but  is  not  known  to 
breed.  It  was  first  recorded  from  the  islands  by  Palmer  (1899,  p. 
383)  on  the  strength  of  an  individual  picked  up  on  the  beach  of  St. 
-Paul  by  True  and  Prentiss  in  1895,  and  another  similarly  taken  by 
D'Arcy  Thompson  in  August,   1897. 

M.  C.  Marsh  next  observed  the  species,  collecting  specimens  on 
St.  Paul  Island  on  March  6  and  April  28,  1912.  Hanna  saw  a  gray 
petrel,  which  was  probably  this  species,  on  St.  George  Island  June 
16,  1914,  and  on  July  15  of  the  same  year,  while  crossing  from  St. 
George  to  St.  Paul,  the  writer  observed  a  few  individuals  at  sea. 
Hanna  took  a  specimen,  now  in  the  U.  S.  National  Museum,  at 
Northeast  Point,  St.  Paul  Island,  on  April  24,  1915 ;  it  was  the  only 
one  seen  at  the  time.  He  observed  another  off  St.  George,  May  4, 
1917. 

Marsh  made  the  following  notes  from  the  fresh  specimens:  Male, 
St.  Paul  Island,  March  6,  1912,  length,  9£  inches;  extent,  18fV;  bill, 
legs,  and  feet,  black.  St.  Paul  (sex  not  determined),  April  28,  1912, 
length,  8^f  inches;  extent,  18-J. 

Food. — A  single  stomach  of  the  forked-tailed  petrel  from  the 
Pribilofs  (St.  Paul,  April  24,  1915)  has  been  examined  and  it  was 
practically  empty,  containing  only  traces  of  fish  bones. 

Family  PHALACROCORACIDAE. 

Phalacrocorax  pelagicus  robustus  Ridgway.     Violet-green  Cormorant. 

The  violet-green  cormorant  seems  to  have  been  first  authoritatively 
credited  to  the  Pribilofs  by  Baird,  Brewer,  and  Ridgway  (1884,  vol. 
2,  p.  161),  who  say:  "It  is  also  said  to  be  abundant  at  St.  George's 
Island,  in  Behring's  Sea,  where  Captain  Smith  obtained  several 
examples."  Although  the  bird  was  thus  early  recorded,  its  regular 
presence  has  been  generally  ignored  by  observers,  and  it  remained 
for  G.  Dallas  Hanna  to  discover  that  the  bird  is  rather  common  and 
breeds  in  a  number  of  places.  He  states  (1916,  p.  402)  that  it  is 
common  about  the  Pribilof  Islands  in  winter,  and  that  it  is  recog- 
nized as  differing  from  the  red-faced  cormorant  by  the  natives,  who 
call  it  "  sea  shag." 

We  found  it  nesting  on  Sealion  Rock  in  1914  and  at  a  number  of 
places  on  both  St.  Paul  and  St.  George  Islands,  and  it  is  believed 
also  to  breed  on  Otter  Island. 

Hanna  has  collected  a  number  of  specimens  which  are  in  the  U.  S. 
National  Museum  (one  is  No.  255123,  St.  George,  May  2,  1917)  and 
has  furnished  the  writer  with  the  following  dates  of  occurrence  on 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  41 

St.  Paul  Island:  1914 — September  22,  2  observed;  December  6,  3; 
December  8,  18  killed  at  Northeast  Point  by  natives;  December  13, 
10.  1915— February  18,  10;  March  5,  2;  March  15,  25;  March  18, 
30;  March  24,  3;  May  6,  4  seen  on  Reef  Cliffs;  May  9,  4  seen  on 
Southwest  Point  Cliffs;  May  18,  2  seen  at  Northeast  Point;  these 
May  records  in  all  probability  indicate  birds  settled  on  their  nesting 
cliffs.  Hanna  also  reports  the  species  killed  at  sea  near  the  island 
February  8,  1918. 

Food. — This  cormorant,  like  all  its  race,  is  a  great  lover  of  fish  food, 
but  investigation  discloses  it  is  by  no  means  exclusively  piscivo- 
rous. The  contents  of  21  well-filled  stomachs  from  the  Pribilof s  were : 
Fishes,  nearly  74  per  cent,  and  Crustacea,  about  26  per  cent.  The 
fishes  were  chiefly  sculpins  taken  by  17  birds;  the  species  identified 
were  Myoxocephalus  sp.,  M.  polyacanthocephalus,  and  Megalocottus 
laticeps.  Three  of  these  violet-green  cormorants  had  captured  rock 
eels  (Pholis  fasciatus) ,  in  one  case  to  the  number  of  20,  which  in 
bulk  were  equal  to  about  three-quarters  of  a  pint.  All  but  three  of 
the  stomachs  examined  contained  remains  of  shrimps  of  the  genus 
Spirontocaris,  and  no  fewer  than  30  individual  shrimps  were  taken 
from  a  single  stomach.  Both  Spirontocaris  polaris  and  IS.  groen- 
landica  were  identified,  and  specimens  of  the  latter  species  3£  inches 
in  length  had  been  eaten.  Four  of  the  birds  also  had  devoured 
hermit  crabs,  which  were  identified  as  Dermaturus  mandtii,  Pagumis 
undosus,  and  Hapalogaster  grebnitzkii.  Ten  of  the  latter  were 
present  in  a  single  stomach.  The  nearly  empty  stomachs  also  con- 
tained crustaceans  and  fish  remains,  and  one  well-filled  stomach 
yielded  a  bit  of  sea  lettuce,  no  doubt  accidentally  taken. 

Phalacrocorax  urile   (Gmelin).     Red-faced  Cormorant. 

The  red-faced  cormorant  is  a  rather  common  breeder,  nesting  on 
all  the  islands  of  the  group  excepting  Sealion  Rock.  It  was  prob- 
ably first  noted  from  the  Pribilof s  b}^  Coinde  (1860,  p.  401).  Owing 
to  this  bird  having  been  confused  with  P.  p.  robustus  in  many  cases 
there  are  numerous  notes  of  occurrence  at  various  seasons  which  can 
not  with  safety  be  used  for  either.  In  the  present  account,  therefore, 
all  doubtful  notes  have  been  ignored. 

Apparently  the  bird  does  not  winter  unless  in  very  small  num- 
bers. It  usually  arrives  in  late  April  or  early  May.  Hanna's  spring 
notes  for  1915  on  St.  Paul  Island  are  as  follows :  April  15,  only  one 
P.  urile  among  25  cormorants  shot  by  the  natives  for  food;  April 
24,  30  observed;  May  9,  8  observed  at  Southwest  Point;  May  18,  6 
seen ;  May  28,  8  seen  at  Northeast  Point. 

The  bird  is  an  early  breeder.  Eggs  were  found  on  Otter  Island 
May  22,  and  on  Walrus  Island  on  May  23,  1875  (notes  from  island 
log).     Other  dates  for  eggs  include:  Walrus  Island,  June  1,  1872, 


42  NORTH   AMERICAN   FAUNA.  [No.  46 

eggs  well  incubated  taken  by  Elliott ;  Walrus  Island,  June  13,  1890, 
eggs  more  or  less  advanced  in  incubation  taken  by  Palmer  (1899, 
p.  377).  On  the  occasion  of  our  own  visit  to  Walrus  Island  on  July 
16,  1914,  young  about  one-fourth  grown  were  found. 

The  birds  remain  about  the  islands  in  small  numbers  throughout 
the  autumn.  Hanna's  notes  for  1914  on  St.  Paul  Island  are  as  fol- 
lows: September  20,  2  observed;  September  22,  1;  October  4,  4; 
October  25,  2 ;  November  16,  5 ;  November  27,  6. 

Palmer  (1899,  p.  373)  has  given  a  very  full  account  of  the  species, 
especially  of  the  nesting  and  the/feather  structure. 

Food. — Five  well-filled  and  one  nearly  empty  stomach  of  the  red- 
faced  cormorant  are  at  hand;  the  latter  may  be  dismissed  at  once 
with  the  statement  that  it  contained  one  mandible  of  a  squid,  the 
only  trace  of  this  type  of  animal  in  the  series.  The  food  in  the  other 
5  stomachs  consisted  of  fishes,  57.8  per  cent;  crustaceans,  41.4  per 
cent;  and  algae  and  hydroids,  0.8  per  cent.  The  fishes  included 
sculpins  (Gymnocanthus  pistilliger) ,  rock  eels  (Pholis  sp.),  and  an 
unidentified  kind.  The  crustaceans  were  chiefly  shrimps  of  the 
genus  Spirontocaris  ;  both  S.  groenlandicus  and  S.  polaris  were  iden- 
tified. No  fewer  than  66  of  the  latter  species  were  present  in  one 
stomach.  Hermit  crabs  {Hapalogaster  grebnitzkii)  were  eaten  also 
by  one  bird,  and  a  spider  crab  (Oregonia  gracilis)  by  another.  Hy- 
droids (Abietinaria  sp.),  sea  lettuce,  and  plume  algae  (Ptilota 
asplenoides,  and  Ptilota  sp.)  were  each  found  in  a  single  stomach. 

Family  ANATIDAE. 
Mergus  merganser  americanus  Cassin.     Amebican  Mebganseb. 

The  earliest  record  of  the  American  merganser  is  that  of  Dall 
(1873,  p.  31),  who  stated  that  the  bird  winters  on  the  Pribilofs. 
Palmer  gives  it  as  undoubtedly  occurring  during  migrations  and  in 
winter,  but  gives  no  definite  notes  beyond  referring  to  Dall's  account. 
The  first  known  specimen  from  the  islands  seems  to  have  been  taken 
on  St.  George  Island,  October  13,  1916,  by  Dr.  H.  P.  Adams,  and  is 
now  No.  253784,  female,  of  the  bird  collection  of  the  U.  S.  National 
Museum. 

Mergus  serrator  Linnaeus.     Red-breasted  Mebganseb. 

The  red-breasted  merganser  has  been  detected  only  a  few  times  on 
the  Pribilofs.  It  was  first  recorded  from  the  group  by  Evermann 
(1913,  p.  17),  who  listed  a  female  specimen  taken  on  St.  Paul  Island, 
December  16,  1910.  Hahn,  who  prepared  and  sent  in  the  specimen, 
states  that  it  was  shot  on  the  lagoon  by  Doctor  Morgan.  It  measured 
in  the  flesh :  Length,  19|  inches ;  extent.  29. 

Hanna  states  that  he  examined  a  specimen  of  this  species  which 
was  shot  by  a  native  on  St.  George  Island  in  the  autumn  of  1913. 


1923.]  BIRDS   OF   THE   PRIBILOF   ISLANDS.  43 

Anas  platyrhyncha  platyrhyncha  Linnaeus.     Mallaed. 

The  mallard  is  a  regular  spring  and  autumn  migrant  on  the  Pribi- 
lofs.  A  few  remain  during  the  summer,  and  it  has  occasionally 
been  knoAvn  to  breed.  It  was  first  added  to  the  avifauna  of  the 
islands  by  Elliott,  who  recorded  it  from  both  islands  (1874,  unpaged, 
and  1875,  p.  190). 

Palmer  observed  a  few  during  the  summer  of  1890  (1899,  p.  380). 
The  following  notes  on  occurrence  in  spring,  taken  from  the  St.  Paul 
Island  log  by  W.  L.  Hahn,  are  of  interest:  May  9,  1888,  1  shot; 
about  April  10,  1903,  2  shot;  May  15,  1905,  observed;  May  7,  1908, 
2  killed  (first  for  season)  ;  May  1,  1909,  several  seen.  Hahn  himself 
reported  it  in  1911  as  follows:  April  22,  3  seen;  April  24,  2  seen; 
April  29,  4  seen ;  April  30,  many  reported. 

On  St.  George  Island,  in  1914,  Hanna  noted  the  mallard  as  follows : 
April  22,  9  seen ;  May  19,  6  seen.  On  St.  Paul  Island,  in  1915,  he  re- 
corded the  following  observations:  April  30,  5  seen;  May  2,  8; 
May  4,  15;  May  6,  4;  May  9,  20;  May  17,  4  (paired)  ;  May  27,  2. 
During  more  recent  years  he  has  a  record  of  4  seen  on  St.  George 
April  14,  1917,  and  7  on  St.  Paul  May  16,  1919.  H.  P.  Adams  noted 
the  bird  on  St.  George  September  26  and  27,  1915 ;  April  16  and  19, 
1916;  and  April  26,  1917.  The  following  are  all  the  data  regarding 
breeding  that  I  have  secured.  Elliott  (1875,  p.  190)  states  that  a 
pair  bred  at  Polovina,  St.  Paul  Island,  in  1872.  Hanna  furnishes 
the  following  notes :  In  1917  a  pair  nested  in  a  pond  at  Little  Polo- 
vina, St.  Paul.  In  1918  a  brood  of  8  young  was  hatched  in  the  pool 
above  Ice-house  Lake,  St.  Paul.  These  were  killed,  with  the  parents, 
before  they  were  half  grown.  Hanna  was  told  by  W.  C.  Allis,  who 
has  wide  experience  on  the  islands,  that  it  was  formerly  not  a  rare 
occurrence  to  see  flocks  of  young  mallards  in  the  various  ponds  on 
the  road  to  Northeast  Point,  and  that  they  were  eagerly  hunted  by 
the  natives.  While  on  the  Pribilofs  in  the  summer  of  1914,  I  ob- 
served the  mallard  only  once,  recording  a  male  seen  at  Kaminista 
Lake,  St.  Paul  Island,  on  August  17. 

The  following  dates  of  occurrence  in  autumn  are  available:  St. 
Paul  Island,  several  shot  October  9,  11,  and  15,  1902  (island  log 
notes  transcribed  by  Hahn).  Hahn  himself  thus  recorded  the  bird 
on  St.  Paul  in  the  autumn  of  1910:  October  9,  several  shot  at 
Northeast  Point;  October  11,  1  seen;  October  14,  5;  October  15, 
20  seen  in  two  flocks  of  6  and  14,  flying  in  direction  of  St.  George 
Island,  in  evening;  October  16,  3;  October  19,  5;  October  20,  4;  Oc- 
tober 23.  1 ;  November  9,  3.  On  St.  George  Island,  in  1913,  Hanna 
saw  1  on  September  28,  and  10  on  October  18 ;  on  St.  Paul,  in  1914, 
he  saw  4  on  November  20,  and  2  on  November  21.  In  1916,  on  St. 
George,  Hanna  noted  it  as  follows :  October  26,  still  abundant ;  No- 


44  NORTH   AMERICAN   FAUNA.  [No.  46 

vember  5,  5  seen ;  December  24,  1  killed.  He  also  found  it  abundant 
there  on  April  28,  1917. 

The  only  winter  record  is  of  a  female  taken  by  M.  C.  Marsh  on 
St.  Paul  Island,  January  1,  1912.  The  following  notes  are  from 
the  fresh  specimen:  Length,  21  inches;  extent,  35^;  legs  and  feet, 
orange-red;  bill,  yellow,  blotched  with  black;  nail,  black.  A  male 
taken  by  Hahn  on  St.  Paul,  November  9,  1910,  measured:  Length, 
22|  inches ;  extent,  35£.  A  female  taken  at  the  same  time  measured : 
Length,  20£  inches ;  extent,  33f . 

Food. — Univalves  (Littorinct  atkana)  constituted  three-fourths  of 
the  food  of  two  mallards  collected  in  a  salt  lagoon  on  St.  Paul  Island, 
November  20, 1914,  and  unidentified  vegetable  matter,  the  other  fourth. 
G.  Dallas  Hanna  notes  that  they  usually  feed  in  fresh-water  ponds 
and  that  they  prey  extensively  on  the  larvae  of  blowflies.  W.  L. 
Hahn  examined  a  gullet  which  was  filled  with  larvae  of  Diptera  and 
of  caddisflies. 

Chaulelasmus  streperus  (Linnaeus).     Gad  wall. 

The  gadwall  has  been  taken  on  the  Pribilofs  on  only  two  occa- 
sions. It  was  first  added  to  the  list  by  Evermann  (1913,  p.  17),  who 
recorded  a  female  specimen  taken  by  M.  C.  Marsh  on  St.  Paul  Island, 
November  13,  1911.  The  following  notes,  taken  by  Marsh  from  the 
bird  in  the  flesh,  are  from  his  field  catalogue:  Length,  18f  inches; 
extent,  33f ;  iris,  brown;  lower  mandible,  light  yellow,  tip,  dark; 
upper  mandible,  darker  yellow  along  sides,  black  at  tip,  and  along 
culmen,  shading  into  yellow  on  the  sides ;  legs  and  toes,  light  yellow, 
dark  at  the  joints;  web,  and  toes  below,  black. 

A  second  specimen,  also  a  female,  taken  from  a  flock  of  three  at 
Halfway  Point,  St.  Paul  Island,  May  20,  1919,  has  been  recorded  by 
Hanna  (1920b,  p.  174). 

Mareca  penelope  (Linnaeus).     European  Widgeon. 

The  European  widgeon  was  first  ascribed  to  the  Pribilofs  by  Coues 
(in  Elliott  1874,  unpaged,  and  1875,  p.  190)  on  the  strength  of  an 
adult  male  specimen  taken  on  St.  Paul  Island,  May  27,  1872,  by  El- 
liott, who  also  reported  seeing  a  few  others.  A  second  specimen, 
also  a  male,  taken  by  W.  L.  Hahn  on  St.  Paul,  April  30,  1911,  was 
recorded  by  Evermann  (1913,  p.  17).  Hahn's  catalogue  gives  the 
flesh  measurements  of  this  bird  as  follows:  Length,  19^  inches;  ex- 
tent, 32.  A  third  specimen,  male,  No.  242691,  preserved  by  A.  G. 
Whitney,  is  in  the  collection  of  the  Biological  Survey.  It  was  taken 
by  a  native  on  Ice-house  Lake,  St.  Paul,  May  9,  1913.  The  follow- 
ing notes  were  made  from  the  fresh  specimen:  Length,  17.7  inches; 
tip  of  upper  mandible  for  one-half  inch  black,  this  color  extending 
farther  back  on  the  edges;  remainder  of  upper  mandible  slaty-blue, 


1923.]  BIRDS   OF   THE    PRIBILOF   ISLANDS.  45 

lighter  between  nostrils;  lower  mandible  black;  feet  and  legs  gray- 
ish, webs  blackish.  Parasites  taken  from  this  specimen  proved  to 
be  Trinoton  luridum,  previously  recorded  from  the  same  host. 

Another,  represented  only  by  the  head  and  feet,  was  taken  by  G. 
Dallas  Hanna  on  St.  Paul,  May  1,  1918,  and  is  now  No.  255133  of  the 
bird  collection  of  the  U.  S.  National  Museum. 

By  the  capture  of  this  bird  on  the  Pribilofs,  the  species  was  added 
to  the  North  American  fauna.     It  breeds  in  Eurasia  from  Scandi- 
navia to  Kamchatka,  migrating  southward  in  winter. 
Eunetta  falcata  (Georgi).     Falcated  Teal. 

The  falcated  teal,  a  beautiful  crested  species,  was  added  to  the  avi- 
fauna of  the  Pribilofs  by  Hanna,  who  has  recorded  (1919a,  p.  176, 
and  1920a,  p.  250)  a  specimen  secured  by  him  on  St.  George  Island, 
April  18,  1917.  This  specimen,  which  is  a  male  in  high  plumage, 
is  now  No.  255131  of  the  bird  collection  of  the  U.  S.  National  Mu- 
seum. The  species  ranges  normally  in  Asia  east  to  Kamchatka, 
Korea,  and  Japan.  The  specimen  mentioned  is  the  first  secured  in 
North  America. 

Food. — The  stomach  of  this  rare  straggler  was  about  half  filled 
with  sea  lettuce. 

Nettion  crecca  (Linnaeus).     European  Teal. 

A  male  and  female  of  the  European  teal,  taken  on  St.  Paul  Island 
May  4,  1918,  and  recorded  by  G.  Dallas  Hanna  (1919a,  p.  176,  and 
1920a,  p.  252),  formed  the  first  record  for  the  Pribilofs.  These  are 
now  Nos.  255127  and  255128  of  the  bird  collection  of  the  U.  S.  Na- 
tional Museum.  Another  specimen,  a  male,  was  taken  on  the  same 
island  September  20,  1919,  and  has  also  been  put  on  record  by  Mr. 
Hanna  (1920b,  p.  174).  It  is  in  the  collection  of  the  California 
Academy  of  Sciences. 

The  species  is  of  general  distribution  in  northern  Eurasia,  and 
breeds  regularly  eastward  to  the  eastern  Aleutians,  where  it  had  been 
detected  many  years  before  its  capture  on  the  Pribilofs. 
Nettion  carolinense   (Gmelin).     Green-winged  Teal. 

The  exact  status  of  the  green- winged  teal  on  the  Pribilofs  is  un- 
certain owing  to  the  great  difficulty  of  distinguishing  the  female 
and  young  of  this  species  from  those  of  the  European  teal  (Nettion 
crecca),  even  with  the  birds  in  hand.  Teals  were  definitely  ascer- 
tained to  breed  at  Antone  Lake,  St.  Paul  Island,  by  Mr.  Hanna  and 
myself  in  the  summer  of  1914,  and  an  adult  female  and  her  unfledged 
young  one  were  taken.  We  failed  to  take  or  even  see  a  male  on  this 
occasion,  however,  and  the  specimens  we  secured  can  not  be  cer- 
tainly identified.  Teals  of  one  or  both  species  also  occur  quite 
frequently  in  migration. 
108731°— 23 i 


46  FORTH   AMERICAN  FAUNA.  [No.  46 

The  present  species  was  recorded  from  the  Pribilofs  by  Palmer 
(1909,  p.  380),  who  saw  one  on  a  pond  on  St.  George  Island,  May  28, 
1890.  It  was  later  killed  by  some  one  but  was  not  secured  for  a 
specimen.  The  following  notes  on  the  occurrence  of  teal  are  given 
with  the  understanding  that  they  may  not  be  referable  in  all  cases 
to  the  American  species:  St.  Paul  Island:  May  15,  1905,  seen  on 
Polovina  Lake  (island  log,  note  transcribed  by  Hahn)  ;  February  27, 
1911,  1  killed  on  Lagoon  Reef  by  a  native,  preserved  by  Hahn  (now 
No.  270533,  male,  collection  Biological  Survey)  ;  May  21,  1911,  1 
killed  (reported  to  Hahn)  ;  July  29,  1914,  1  seen  on  Polovina  Lake 
(Preble)  ;  August  22  (4),  and  25  (1),  1914,  (Preble  and  Hanna) ; 
October  17,  1914,  3  seen  (Hanna)  ;  May  18,  1915,  1,  Halfway  Point 
(Hanna). 

On  St.  George  Island  Hanna  observed  3  teals  on  September  4,  1913, 
and  1  on  May  19,  1914,  and  collected  a  fine  male  of  the  American 
form  there  on  May  10,  1917.  Another  male  was  taken  by  him  on  St. 
Paul,  May  26,  1919  (1920b,  p.  174).  These  two  last-mentioned 
specimens  are  in  the  collections  of  the  U.  S.  National  Museum,  and 
of  the  California  Academy  of  Sciences,  respectively. 

Food. — Two  teals  from  Antone  Lake,  St.  Paul  (August  31,  1914, 
G.  D.  Hanna),  identified  as  the  green-winged,  had  been  feeding  on 
seeds  and  foliage  of  white  water  crowfoot  {Batrachium  trichophyl- 
luni)  and  a  pondweed  {Potamogeton  filiformis),  together  with  a  few 
small  Diptera. 

Two  other  teals,  not  certainly  identified  as  to  species,  collected  at 
the  same  lake  in  the  same  month  (August  22,  25,  1914)  had  taken  the 
same  things  in  somewhat  different  proportions,  together  with  cer- 
tain additional  items.  Water  crowfoot  made  the  bulk  of  the  food 
of  one  of  these  birds,  but  a  few  seeds  of  pondweed  and  a  sedge 
(Carex  sp.)  were  present.  The  bird  had  eaten  also  beetles,  flies, 
bivalved  crustaceans  (Ostracoda),  and  mollusks.  The  preponder- 
ant article  of  food  of  the  second  teal  was  rat-tailed  larvae  of  a 
flower  fly  (Eristalinae),  of  which  at  least  16  had  been  taken.  This 
bird  had  eaten  also  lesser  quantities  of  beetles  and  Hymenoptera  and 
of  the  seeds  of  sedge  and  water  crowfoot. 

Spatula  clypeata  (Linnaeus).     Shoveleb. 

The  shoveler  was  added  to  the  Pribilof  list  by  Evermann  (1913, 
p.  17),  who  recorded  a  male  specimen  taken  by  W.  L.  Hahn  on  St. 
Paul  Island,  May  24,  1911.  The  following  notes  are  from  Hahn's 
catalogue:  Length,  19^  inches;  extent,  30;  legs  and  feet,  pinkish- 
orange;  bill,  glossy  black,  except  for  slight  yellow  mottling  below. 
This  specimen  is  now  in  the  Biological  Survey  collection. 

The  only  other  record  seems  to  be  that  of  a  male  bird  observed  by 
the  writer  near  Northeast  Point,  St.  Paul  Island,  June  27,  1914. 


1923.]  BIRDS  OF   THE  PBIBILOF  ISLANDS.  47 

Dafila  acuta  tzitzihoa  (Vieillot).     Pintail. 

Two  male  specimens  of  the  pintail  taken  by  M.  C.  Marsh  on  St. 
Paul  Island,  May  24, 1912,  were  recorded  by  Evermann  (1913,  p.  17). 
Marsh  in  his  field  catalogue  gives  the  following  notes  from  the  fresh 
specimens :  No.  178,  male,  length,  25£  inches,  extent,  35| ;  iris,  brown ; 
bill,  blue,  with  a  broad  black  stripe  down  the  middle  and  a  narrow 
black  stripe  at  base ;  upper  mandible,  black,  edged  forward ;  feet,  gray 
above,  webs  darker.  No.  179,  male,  length,  24^f  inches ;  extent,  34f ; 
colors  of  soft  parts  same  as  No.  178. 

Hanna  observed  9  individuals  at  Webster  Lake,  August  17,  1914, 
and  obtained  1  (No.  237499,  U.  S.  Nat.  Mus.).  He  also  noted  2  on 
St.  Paul  Island,  May  27,  and  2  on  May  31,  1915.  On  St.  George,  in 
1916,  he  noted  it  as  still  common  on  October  26.  He  considers  that 
the  bird  is  a  common  migrant. 

Food. — The  Pribilof  Islands  evidently  do  not  afford  the  vegetable 
food  which  is  most  relished  by  the  pintail  in  its  continental  range, 
for  the  six  stomachs  of  this  species  collected  there  were  filled  entirely 
with  animal  food.  Five  of  them  contained  only  midge  (Chirono- 
midae)  larvae  and  the  other  one  caddis  (Trichoptera)  larvae  and 
their  cases.    Mr.  Hanna  says  that  this  species — 

has  never  been  seen  feeding  anywhere  except  in  fresh-water  ponds.  They  eat 
the  lavae  of  blowflies  from  the  seal  carcasses  extensively.  These  larvae  wander 
from  the  decaying  meat  when  the  time  comes  for  them  to  pupate  and  many  of 
them  are  drowned  in  the  fresh-water  ponds. 

Aristonetta  valisineria  (Wilson).8     Canvasback. 

The  famous  canvasback  was  added  to  the  Pribilof  list  by  G.  Dallas 
Hanna  (1919a,  p.  176,  and  1920a,  p.  252)  on  the  strength  of  a  fine 
adult  male  taken  on  St.  George  Island,  May  18,  1917.  It  is  now  in 
the  collection  of  the  U.  S.  National  Museum,  No.  255149. 

Food. — The  stomach  of  this  specimen  contained  only  caddis  larvae 
and  cases. 

Fulix  marila  (Linnaeus).*     Scaup  Duck. 

A  male  scaup  duck  taken  by  M.  C.  Marsh  on  St.  Paul  Island, 
April  27,  1912,  was  recorded  by  Evermann  (1913,  p.  17).  Marsh 
in  his  field  catalogue  gives  the  following  notes:  Length,  18  inches; 
extent,  31f ;  bill,  pale  blue;  nail,  black;  iris,  yellow.  The  species 
seems  not  to  have  been  otherwise  observed. 

Fuligula  fuligula  (Linnaeus).     Tufted  Duck. 

A  female  tufted  duck  collected  on  St.  Paul  Island,  May  9,  1911, 
by  W.  L.  Hahn,  proved  to  be  the  first  record  for  the  Pribilofs,  and 

1  Marila  valisineria  of  the  A.  O.  D.  check  list,  1910. 
•  Marila  marila  of  the  A.  O.  U.  Check  List,  1910. 


48  NORTH  AMERICAN   FAUNA.  [No.  46 

also  for  North  America.  It  was  formally  recorded  by  Evermann 
(1913,  p.  17).  The  bird  was  accompanied  by  a  male,  which  escaped. 
The  species  breeds  regularly  in  northern  Eurasia  east  to  Kamchatka, 
and  migrates  to  northern  Africa,  India,  China,  and  Japan. 

Food. — Evermann  (I.e.)  quotes  from  Hahn's  field  notes  as  fol- 
lows: "the  stomach  was  full  of  grass,  cress  (?),  small  seeds,  and  a 
few  larvae. " 

Aithyia  ferina  (Linnaeus).    European  Pochard. 

The  European  pochard  was  added  to  the  avifauna  of  the  Pribilofs 
and  North  America  by  Evermann  (1913,  p.  17),  on  the  basis  of  a 
male  specimen  taken  by  M.  C.  Marsh  on  St.  Paul  Island,  May  4, 
1912.  His  catalogue  records  the  following  notes  made  from  the 
fresh  specimen:  Length,  18  inches;  extent,  30-|;  iris,  yellowish-red; 
base  of  bill,  brownish-black,  middle  pale  bluish,  terminal  band  of 
jet  black  sharply  demarcated  from  the  bluish;  legs  and  toes,  pale, 
with  much  grayish,  webs  black.  The  specimen  is  now  No.  239123 
of  the  collection  of  the  Biological  Survey.  The  pochard  breeds  in 
northern  Eurasia  east  to  Lake  Baikal,  and  in  migration  and  in 
winter  wanders  eastward  to  China  and  Japan. 

Glaucionetta  clangula  clangula   (Linnaeus).     European  Goldeneye. 

A  female  goldeneye  (No.  237500,  U.  S.  Nat.  Mus.),  taken  by  G. 
Dallas  Hanna  on  St.  Paul  Island  November  27,  1914,  from  a  flock  of 
four,  proved  to  be  the  typical  form,  and  hence  constitutes  a  new 
record  for  North  America.  It  was  so  reported  by  Mr.  Hanna  (1916, 
p.  400).  The  specimen  was  determined  by  Dr.  H.  C.  Oberholser. 
This    form  breeds  generally  throughout  northern  Eurasia. 

Other  goldeneyes,  assumed  to  belong  to  the  present  form,  were 
observed  by  Mr.  Hanna  on  St.  Paul  Island,  some  in  the  possession 
of  native  hunters,  as  follows:  October  12,  1914,  4;  October  25,  4; 
April  12,  1915,  5. 

Food. — Three  stomachs  from  St.  Paul  (October  12,  1914)  have 
been  examined  and  their  contents  were  amphipods,  55  per  cent; 
fish  remains,  10  per  cent;  marine  worms  (Nereidae),  3.3  per  cent; 
and  seeds  of  pondweed  {Potamogeton  sp.),  31.7  per  cent. 

Glaucionetta  clangula  americana  (Bonaparte).    American  Goldeneye. 

During  recent  years  this  goldeneye  has  been  taken  on  both  St. 
Paul  and  St.  George  Islands  by  G.  Dallas  Hanna,  who  has  put  the 
captures  on  record.  He  took  a  male  specimen  on  St.  George  Island, 
May  6,  1917,  and  a  female  on  St.  Paul,  January  31,  1918  (1919a,  p. 
176,  and  1910a,  p.  252). 

Food. — The  stomach  of  the  specimen  last  mentioned  was  practi- 
cally empty;  however,  the  slight  remains  of  food  represented  three 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  49 

distinct  phyla  of  animals,  namely,  Mollusca  (Modiolaria  verrucosa) , 
barnacles,  and  sea  urchins. 

Charitonetta  albeola  (Linnaeus).     Bufflehead. 

The  beautiful  little  bufflehead  was  first  taken  on  the  Pribilofs  by 
G.  Dallas  Hanna,  and  was  recorded  by  him  (1916,  p.  401).  He 
observed  it  first  on  St.  George  Island,  taking  male  specimens  Novem- 
ber 19,  1913,  and  May  19,  1914.  In  the  spring  of  1915  he  observed 
others  on  St.  Paul  Island  as  follows:  May  19,  2  seen  at  Halfway 
Point;  May  26,  1  Northeast  Point;  May  28,  8  (paired)  on  Webster 
Lake. 

Food. — Only  animal  food  was  present  in  the  two  bufflehead 
gizzards  collected  by  Mr.  Hanna  on  St.  Paul,  May  19,  1914,  and 
January  29,  1918.  One  of  the  stomachs  held  only  caddis  larvae  and 
cases,  while  the  other  contained  ground-up  amphipods,  80  per  cent; 
7  or  more  small  bivalves  (Modiolaria  verrucosa),  18  per  cent;  and 
a  bit  of  barnacle,  2  per  cent. 

Clangula  hyemalis  (Linnaeus).     Old-squaw. 

The  old-squaw  is  resident  about  the  Pribilofs,  being,  however, 
much  more  abundant  in  winter  than  in  summer.  It  breeds  only 
on  St.  Paul,  nesting  about  most  of  the  shallow  fresh-water  ponds 
which  are  scattered  over  the  island.  It  was  first  recorded  from 
the  group  by  Elliott  (1874,  unpaged,  and  1875,  p.  191). 

Since  the  species  is  abundant  during  the  winter  and  spring,  no 
dates  of  arrival  can  be  cited;  Hanna  records  the  birds  as  pairing 
April  18,  1915,  about  which  date  they  begin  acquiring  the  summer 
plumage,  and  as  being  observed  in  pairs  at  various  dates  in  May. 
Eggs  were  found  June  12  and  17,  1890,  by  Palmer  (1899,  p.  378). 
The  egg-laying  period  is  long,  since  downy  young  not  over  two 
days  old  were  taken  August  17,  1897,  by  Snodgrass  and  Greely 
(Grinnell,  1901,  p.  19),  and  the  writer  saw  downy  young  on  Antone 
Lake,  August  25,  1914. 

Many  fall  and  winter  dates  are  at  hand,  but  in  the  case  of  so 
common  a  bird  it  seems  unnecessary  to  give  them  in  detail.  They 
seem  to  indicate  a  gradual  gain  in  numbers  until  the  species  is  seen 
by  hundreds  or  thousands  during  the  winter  and  early  spring. 

A  male,  St.  Paul  Island,  January  7,  1912.  M.  C.  Marsh,  measured : 
Length,  16|  inches;  extent,  28f.  Another  male,  St.  Paul  Island, 
February  5,  1912,  M.  C.  Marsh :  Length,  16  inches ;  extent,  28 J ;  iris 
brown ;  bill  black  with  band  of  rose.  Another  male,  St.  Paul,  March 
27,  1912,  M.  C.  Marsh:  Length,  19  inches;  extent,  28^;  iris  brown, 
basal  half  of  bill  jet-black,  rest  rose-pink  except  black  nail;  lower 
mandible  same  relative  colors;  legs  and  feet  pale  bluish,  webs  black. 
A  female  specimen,  St.  Paul  Island,  February  5,  1912,  M.  C.  Marsh, 


50  NORTH   AMERICAN   FAUNA.  [No.  46 

measured:  Length,  15^  inches;  extent,  27;  iris  brown;  feet  pale 
bluish,  darkly  reticulated.  Another  female,  same  place  and  collector, 
February  23,  1912,  measured:  Length,  15^  inches;  extent,  27|;  bill 
dusky  bluish  except  black  nail,  and  black  bordering  the  feathers  at 
base  of  upper  mandible  and  the  nostrils;  iris  light  brown;  legs  and 
toes  in  front  very  pale-bluish,  black  behind ;  webs  black. 

Food. — Fifteen  well-filled  and  fourteen  nearly  empty  stomachs 
of  the  old-squaw  are  available  to  illustrate  its  food  habits  at  the 
Pribilof  Islands.  From  the  nature  of  the  food  it  is  evident  that 
most  of  it  was  obtained  in  salt /Water,  and  in  this  connection  a  note 
on  the  habits  of  the  species  by  Mr.  Hanna  is  of  interest.  He  says 
the  old-squaw  is — 

much  more  abundant  in  winter  than  in  summer.  It  breeds  in  the  lowland 
fresh-water  lakes  where  several  sets  of  eggs  have  been  collected.  Parent  birds 
have  been  seen  flying  fromr  the  sea  to  the  nesting  site  which  might  indicate 
that  food  is  secured  in  salt  water  at  all  seasons.  In  winter  they  feed  just  out- 
side the  surf  lines,  in  the  open  spaces  between  the  ice  cakes,  and  in  the  Salt 
Lagoon  when  this  is  open. 

The  food  composing  the  15  normal  stomach  contents  studied  con- 
sisted of  amphipods,  74.1  per  cent;  hermit  crabs,  11.2  per  cent;  caddis 
larvae  (the  only  fresh-water  animals  in  the  lot),  6.4  per  cent;  mol- 
lusks,  3  per  cent ;  vegetable  matter,  1.6  per  cent ;  and  numerous  items 
of  lesser  importance.  In  12  cases  amphipods  constituted  the  bulk 
of  the  food  and  were  represented  by  very  numerous  individuals. 
The  species  identified  were  Anonyx  nugax,  Bathymedon  (?)  obtusi- 
frons,  Ca/prella  kincaidi,  Gammarus  pribilofensis,  Hippomedon  sp., 
Orchomenella  minuta,  Pontoporeia  affinis,  and  Tmetonyx  sp.  Other 
crustaceans  eaten  were  the  hermit  crabs  Dermaturus  mandtii  and 
Hapalogaster  grebnitzkii,  remains  of  8  or  more  of  the  former  and  6 
or  more  of  the  latter  were  found  (together)  in  one  stomach.  Barna- 
cles had  been  eaten  by  9  of  the  well-fed  birds,  and  compose  1.2  per 
cent  of  the  total  food ;  only  one  species,  Balanus  balanus,  was  identi- 
fied. Mollusks  composing  3  per  cent  of  the  diet  included  both 
bivalves  and  univalves.  Of  the  former  the  common  blue  mussel 
{Mytilus  edulis)  was  found  in  6  stomachs,  Saxicava  arctica  in  3, 
Modiolaria  verrucosa  in  2,  and  Musculus  sp.  in  1.  The  univalves 
identified  were  Margarites  helicina,  Littorina  atk/ma,  and  Borcotro- 
phon  sp.  Remains  of  a  sea  slug  (Chiton)  occurred  in -one  stomach. 
Among  the  other  items  of  animal  food  were  Foraminifera,  hydroids 
(Abietinaria  sp.),  B^ozoa,  nereid  and  tubicolous  worms,  sand  dol- 
lars (E chinaraehnius  parma),  and  other  sea  urchins. 

The  vegetable  food  taken  by  these  15  old  squaws  comprised  root- 
stocks  and  foliage  of  grass  and  sedge,  seeds  of  sedge  (Carex  sp.)  and 
lupine  (Lupinus  nootkatensis) ,  and  algae.  The  latter  included  kelp, 
plume  algae  (Ceramiaceae),  and  a  filamentous  alga  (Chaetomorpha 
melagonium) . 


1923.]  BIRDS   OF    THE   PBIBILOF  ISLANDS.  51 

The  14  nearly  empt}r  stomachs  contained  chiefly  the  same  items  as 
the  better-filled  ones,  those  of  most  frequent  occurrence  being  amphi- 
pods,  barnacles,  and  mollusks.  One  additional  item,  namely  star- 
fish remains,  was  obtained  from  this  series. 

Histrionicus  histrionicus  pacificus  Brooks.     Harlequin  Duck. 

The  harlequin  is  resident  the  entire  year  on  the  Pribilofs,  being 
most  common  in  spring  and  fall,  fairly  common  in  winter  except 
when  the  pack  ice  is  present,  and  found  in  small  flocks  throughout 
the  summer.  No  absolute  evidence  of  its  breeding  on  the  islands  is 
at  hand.  It  was  first  definitely  recorded  from  the  Pribilofs  by 
Elliott  (1874,  unpaged,  and  1875,  p.  199). 

Being  abundant  during  all  or  most  of  the  winter,  it  can  not  be 
said  to  arrive  in  spring.  Hanna  gives  many  instances  of  occurrence 
in  spring  and  summer,  from  which  the  following  may  be  given : 
St.  Paul  Island,  1915— March  5,  25;  March  18,  6;  April  24,  200; 
April  27,  50;  May  4,  40;  May  28,  10;  July  19,  several  individuals 
seen  which  were  molting  and  could  not  fly;  have  been  abundant  all 
summer.  While  on  the  islands  in  1914  the  writer  saw  flocks  on  the 
surf-washed  rocks  almost  daily,  both  on  St.  Paul  and  St.  George. 
The  notes  most  nearly  giving  evidence  of  breeding  that  I  have  been 
able  to  find  are  these  of  Palmer,  who  says  (1899,  p.  379)  :  "I  was 
told  that  some  young  were  killed  on  St.  George  during  the  summer 
[of  1890]."  Hahn  observed  two  pairs  mating  on  St.  Paul  Island, 
April  24,  1911 ;  these,  however,  may  have  bred  elsewhere. 

Numbers  of  specimens  of  both  sexes  are  in  the  collection  of  the 
U.  S.  National  Museum.  The  following  notes  from  selected  speci- 
mens from  St.  Paul  Island  are  from  the  collectors'  field  catalogues; 
Male,  November  18,  1910,  W.  L.  Hahn;  length,  18  inches;  extent, 
25|;  bill  plumbeous  at  base,  tipped  with  pale  horn  color;  feet  dusky. 
Male,  December  7,  1911,  M.  C.  Marsh;  length,  lfr|  inches;  extent  24|; 
bill  dark  olive,  lighter  underneath;  tip  of  lower  mandible  rosy; 
tarsus  and  toes  pale  greenish,  joints  dark,  webs  black,  obscure  streaks 
of  rose  along  inner  surface  of  tarsus  and  middle  and  inner  toes; 
hallux  lobe  black  on  outer  surface.  Male,  February  23,  1911,  M.  C. 
Marsh;  length,  17  inches;  extent,  27|;  iris  dark  brown;  bill  pale 
bluish,  darkened  at  base;  nail  lighter,  legs  and  feet  dark  brown, 
webs  black.  Male,  March  8,  1912,  Marsh:  Length,  16£  inches;  ex- 
tent, 26|;  iris  brown,  bill  dark  leaden,  rami  below  pale  blue,  mem- 
brane between  black;  legs  and  feet  with  slight  pale  bluish  cast, 
darker  at  joints,  webs  black.  Female,  October  20,  1911,  Marsh: 
Length,  14T9F  inches;  extent,  23;  iris  brown;  bill  leaden,  lighter  un- 
derneath; tarsus  very  pale  yellowish  in  front,  dark  behind;  toes  pale 
yellow,  the  joints  dark,  web  brownish  black. 


52  NORTH   AMERICAN  FAUNA.  [No.  46 

Food. — Mr.  Hanna  notes  that  harlequin  ducks  feed  in  the  kelp  beds 
and  rest  on  the  surf-washed  rocks  the  year  round,  and  it  is  evident 
from  the  contents  of  the  11  well-filled  and  8  nearly  empty  stomachs 
of  the  species  collected  about  the  Pribilofs  that  the  birds  take  full 
advantage  of  the  great  variety  of  invertebrate  life  to  be  found  in 
these  rich  marine  pastures.  The  bulk  of  their  food  is  amphipods 
(51.4  per  cent  of  the  total  contents  of  11  stomachs),  hermit  crabs  25.1 
per  cent,  and  mollusks  19  per  cent.  Other  items,  though  numerous,  in 
every  case  fall  short  of  1  per  cent  of  the  total  food.  Amphipods,  the 
greatest  staple,  are  taken  in  large  numbers,  and  a  variety  of  species 
are  captured.  Identifications  are :  Anonyx  nugax,  Gammarus  pribi- 
lofensis,  Hyale  sp.,  Jassa  marmorata,  Metopa  sp.,  and  Orchomenella 
rninuta.  The  hermit  crabs  in  all  cases  were  Dermaturus  mandtii,  and 
16  of  these  were  represented  in  the  stomach  of  one  duck.  Isopod 
crustaceans  were  found  in  one  gizzard  and  barnacles  in  five.  The 
mollusks  included  widely  diverse  members  of  the  group,  as  sea  slugs 
(Chiton  sp.,  Schizoplax  sp.),  limpets,  sea  snails,  and  mussels.  The 
univalves  identified  were  Littorina  atkama,  Margarites  helicina,  and 
Trichotropis  insignis,  and  the  bivalves  Modiolaria  verrucosa,  Mytilus 
edulis,  and  Saxicava  arctica. 

Among  the  animal  food  items  of  lesser  importance  are  Foramini- 
fera,  hydroids  (Abietinaria  sp.),  tubicolous  and  other  (nereid) 
marine  worms,  sea  spiders  or  pycnogonids  (Ammothea  pribilofensis) , 
sea  urchins,  and  sand  dollars  (Echinarachnius  parma) .  Vegetable 
matter  composed  less  than  1  per  cent  of  the  food  of  the  harlequins 
here  reported  upon  and  all  of  it  may  have  been  taken  accidentally.  It 
included  kelp,  plume  algae  (Ptilota  pectinata,  and  P.  asplenoides) , 
and  other  algae  (Pterosiph&nia  bipinnata). 

Scraps  of  food  in  the  8  nearly  empty  stomachs  of  harlequins  were 
chiefly  of  the  same  items  mentioned  above,  amphipods  and  barnacles 
(in  part  B alarms  balanus)  being  of  most  frequent  occurrence.  Ad- 
ditional foods  from  these  stomachs  were  bryozoans,  the  amphipod 
Pontoporeia  afjinis,  and  fish.  W.  L.  Hahn  examined  a  bird  that  had 
fed  more  extensively  on  isopods  than  any  of  those  here  reported  upon, 
there  being  45  of  these  animals  entire  in  the  gullet  and  stomach  be- 
sides many  fragments. 

Polysticta  stelleri  (Pallas).     Stellee  Eidee. 

The  beautiful  Steller  eider  is  apparently  a  regular  migrant,  and 
early  summer  visitor,  though  the  records  are  somewhat  scattered  and 
meager.  Elliott  shot  2  in  June,  1872,  from  the  Village  Hill  on  St. 
Paul,  and  speaks  of  others  shot  off  East  Point,  St.  George,  in  the 
same  year  (1874,  unpaged,  and  1875,  p.  192).  Palmer  saw  several 
in  company  with  harlequin  ducks,  from  the  Village  Hill,  St.  Paul, 
and  on  June  14  shot  3  males  from  a  flock  of  4  in  St.  Paul  Harbor. 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  53 

Later  he  saw  many  others.  The  species  was  next  detected  by  Marsh, 
who  prepared  2  males  which  were  shot  by  P.  R.  E.  Hatton  on  the 
Lagoon,  St.  Paul  Island,  on  April  23,  1912,  and  took  another  on 
May  9.  These  are  in  the  collection  of  the  Biological  Survey.  Hanna 
was  the  next  observer  to  record  it,  taking  a  female  specimen  on  St. 
Paul,  October  25,  1914,  and  a  male  on  March  21,  1915.  He  also  ob- 
served 4  on  St.  Paul,  April  18,  1915. 

Individuals  of  this  species,  represented  by  stomachs  sent  in  for 
examination,  were  later  taken  as  follows:  St.  George,  February  4 
and  April  12,  1917;  St.  Paul,  December  6,  1918.  The  specimen  taken 
February  4  is  now  No.  255125  of  the  U.  S.  National  Museum  col- 
lection. 

Notes  from  Marsh's  field  catalogue,  taken  from  the  fresh  specimens, 
are  as  follows:  Male,  St.  Paul  Island,  April  23,  1912,  length,  18£ 
inches;  extent,  29T9e;  iris  dark  brown;  bill  pale  blue;  legs  and 
feet  grayish  brown.  Another  male,  same  place  and  date,  measured : 
Length,  17|  inches;  extent,  28.  Notes  on  soft  parts  same  as  pre- 
ceding. The  specimen  taken  May  9,  1912,  measured:  Length,  17 J 
inches;  extent,  28|. 

Food. — The  food  of  the  Steller  duck  in  the  Pribilof  region  is 
known  from  the  examination  of  two  well-filled  stomachs  and  one 
other.  The  content  of  the  well-filled  stomachs  was  some  60  per 
cent  amphipods  and  the  remainder  chiefly  mollusks.  Amphipods 
identified  were  Allorchestes  sp.,  and  Pontoporeia  afflnis.  The  mol- 
lusks were  chiefly  univalves,  Neverita  recluzmna  (25  or  more  in  one 
stomach),  Meltmella  sp.,  and  Cardium  ciliatum  beirg  identified. 
Minor  items  of  food  were  a  bit  of  a  bivalved  molluslc,  and  remains 
of  marine  worms  (Nereidae),  barnacle,  sea  urchin,  ana  algae. 

Arctonetta  fischeri  (Brandt).     Spectacled  Eideb. 

The  spectacled  eider  was  added  to  the  Pribilof  list  by  the  capture  of 
three  female  birds  on  January  13,  1918.  These  were  taken  by  native 
hunters  and  secured  by  G.  Dallas  Hanna,  who  has  put  their  capture 
on  record  (1919a,  p.  176,  and  1920a,  p.  252).  These  specimens,  two  of 
which  were  preserved  as  skins  and  the  other  in  formalin  for  anatomi- 
cal study,  are  now  in  the  collection  of  the  U.  S.  National  Museum. 

Food. — Two  stomach  contents  of  the  spectacled  eider  have  been  ex- 
amined and  were  found  to  be  composed  of  amphipocfo,  about  90  per 
cent;  mollusks,  about  5  per  cent;  and  various  minor  items.  One 
species  of  amphipod  was  determined,  namely,  Batlymedon  obtusi- 
frons,  and  4  of  mollusks.  The  latter  included  the  common  blue  mus- 
sel (Mytilus  edulis)  ;  Modiolaria  verrucosa  and  Rochefortia  sp.,  bi- 
valves; and  Lora  sp.,  univalve.  Other  articles  of  food  in  these  two 
stomachs    were:    Foraminifera,    Bryozoa,    sea    urchin,    sand-dollar 


54  NORTH   AMERICAN"  FAUNA.  [No.  46 

(E chinarachnius  parma) ,  starfish,  crab  {Erimacrus  iseribeckii) ,  bones 
of  a  sculpin,  and  bits  of  kelp. 

Somateria  v-nigra  Gray.     Pacific  Eideb. 

The  Pacific  eider  was  first  credited  to  the  Pribilofs  by  Palmer 
(1899,  p.  380),  who  gives  it  as  a  winter  visitant,  usually  seen  offshore. 
The  first  actual  specimen  seems  to  have  been  taken  by  Hahn  (male, 
St.  Paul  Island,  May  17,  1911).  Marsh  later  collected  female  speci- 
mens at  the  same  place  on  December  23,  1911,  and  January  8,  1912. 

Hanna  states  that  many  are  shot  by  the  natives  when  the  ice  is 
around  the  islands.  He  notes  eiders,  probably  the  present  species,  on 
St.  George  Island,  November  11,  1913,  when  20  were  seen  about  100 
feet  offshore.  Six  others  were  seen  at  the  same  place  March  28,  1914. 
Later  notes,  referring  to  birds  certainly  identified,  are  as  follows :  St. 
Paul  Island,  December  4,  1914,  10  observed ;  December  13,  10 ;  Jan- 
uary 10,  about  100;  February  18,  about  5,000;  March  5,  about  300; 
April  15,  60 ;  April  18,  20 ;  April  24,  200 ;  April  27,  50.  H.  P.  Adams 
noted  the  species  about  St.  George  on  February  10,  1916 ;  and  Janu- 
ary 13,  February  4,  and  March  25,  1917. 

Notes  from  field  catalogues  follow :  St.  Paul  Island,  male,  May  17, 
1911,  W.  L.  Hahn:  Bill,  deep  orange;  feet,  orange.  St.  Paul  Island, 
male,  January  1,  1912,  Marsh:  Length,  24f  inches;  extent,  42$; 
bill,  orange,  paler  terminally;  legs  and  feet,  yellow.  St.  Paul  Island, 
female,  January  8,  1912,  Marsh  :  Length,  23f ;  extent,  39 ;  iris,  brown; 
basal  portion  of  bill,  gray ;  terminal,  pale  yellow ;  legs  and  feet,  dusky 
yellowish;  webs,  darker. 

Food. — Information  obtained  by  examination  of  a  collection  of  44 
well-filled  and  3  other  stomachs  of  the  Pacific  eider  from  the  Pribi- 
lofs is  sufficient  to  give  a  very  good  idea  of  the  food  of  the  species 
in  that  region  during  the  months  in  which  they  were  collected; 
namely,  January,  February,  and  March.  A  wide  variety  of  organ- 
isms inhabiting  the  sea  is  represented  in  the  food,  ranging  from 
algae  and  Foraminifera  through  the  principal  group  of  marine  in- 
vertebrates to  the  Chordata,  or  backboned  animals,  as  ascidians 
and  fishes.  Many  of  the  articles  of  diet  are  not  fed  upon  to  any 
great  extent,  and  the  favorites  seem  to  be  mollusks  and  crustaceans, 
according  to  the  following  schedule  of  the  principal  items :  Mollusks, 
41.2  per  cent;  crustaceans,  28.2  per  cent;  sea  urchins,  9.45  per  cent; 
starfishes,  7.34  per  cent;  and  algae,  7.47  per  cent.  The  mollusks 
included  limpets,  sea  snails,  mussels,  and  sea  slugs.  The  kinds  most 
frequently  taken  were  common  blue  mussels  (Mytilus  edidis),  found 
in  15  stomachs,  and  another  species  of  bivalve  (Modiolaria  verru- 
cosa), found  in  11;  in  two  cases  hundreds  of  the  latter  shellfish  had 
been  eaten.     Besides  these,  identifications  made  of  mollusks  of  the 


1923.]  BIRDS  OF   THE   PRIBILOF  ISLANDS.  55 

various  groups  were:  Aemaea  sp. ;  Amphissa  reticulata,  Borcotro- 
phon  (?)  multicostatus,  Buccinum  plectrum,  Haloconcha  (?) 
minor,  H.  reflexa,  Littorina  atkana,  L.  grandis,  Margarites 
helicina,  Natica  clausa,  Neverita  recluziana,  Nucella  lima,  Plicifiisus 
spitzbergensis,  Trichotropis  insignis,  Volutharpa  ampullacea,  Car- 
dium  ciliatum,  Pecten  islandicus,  Saxicava  arctica,  Chiton  mar- 
moreus,  and  Chiton  sp. 

The  crustaceans  eaten  by  Pacific  eiders  consisted  of  crabs,  13.0G  per 
cent;  amphipods,  12.4  per  cent;  shrimps,  1.5  per  cent;  and  isopods, 
1.3  per  cent.  The  crabs  captured  were  principally  hermit  crabs,  and 
one  species,  Dermaturus  mandtii,  was  found  in  17  stomachs  and  to 
the  number  of  11  specimens  in  one  of  them.  The  other  hermit  crabs 
eaten  were  Hapalogaster  grebnitzkii,  Pagurus  dalli,  and  P.  trigono- 
cheirus.  Spider  crabs  (Oregonia  gracilis)  were  found  in  two  stom- 
achs, and  remains  of  crabs  of  the  family  Xanthidae,  a  group  not 
hitherto  reported  from  the  region,  in  one.  The  amphipods  identified 
belonged  to  a  number  of  species,  among  which  Gammarus  prihilo- 
fensis  occurred  most  frequently,  namely  5  times.  The  other  forms 
determined  were:  Allorchestes  malleolus,  Allorchestes  sp.,  Amphi- 
thoe  rubricata,  Anonyx  nugax,  Atylus  sp.,  Calliopiidae,  Caprella  kin- 
caidi.  Ischyrocerus  sp.,  Metopa  sp.,  Nebalia  sp.,  N eopleustes  assimilis, 
Neopleustes  sp.,  and  Pleustes  panoplus.  The  isopods  eaten  by  Pacific 
eiders  belong  to  the  family  Idotheidae,  of  which  Idothea  ochotensis 
was  specifically  identified.  Thirty  shrimps  (Spirontocaris  polaris) 
were  found  in  one  stomach  and  a  few  minute  bivalved  crustaceans 
(Ostracoda)  in  another. 

Among  birds  thus  far  studied  as  to  food  habits,  preying  upon  sea 
urchins  and  starfishes  is  unusual,  but  Pacific  eiders  in  the  Pribilof 
region  form  an  exception  to  the  rule  in  that  more  than  a  sixth  of 
their  food  is  made  up  of  these  animals.  The  sea  urchins  most  fre- 
quently taken  were  the  sand-dollars  (E chniarachnius  parma),  but 
spheroidal  urchins  (Strongylocentrus  drobachiensis)  of  the  more 
ordinary  type  were  not  neglected.  Sea  urchins  (9.45  per  cent)  were 
found  in  19  stomachs,  and  starfishes  (7.34  per  cent)  in  14 ;  the  latter 
in  5  cases  were  identified  as  Leptasterias  sp. 

Other  articles  of  animal  food  forming  more  than  1  per  cent  of 
the  diet  were  barnacles  and  worms.  The  former  had  been  eaten  by 
14  birds,  and  only  one  species  (Balanus  balanus)  was  identified; 
the  worms  were  of  the  family  Nereidae  in  8  cases,  and  tubicolous 
forms  in  two.  While  amounting  to  less  than  1  per  cent,  other  animals 
eaten  by  the  Pacific  eiders  are  of  considerable  interest  because  they 
are  preyed  upon  by  so  few  birds.  Such  are  the  hydroids,  bryozoans, 
sea  spiders,  and  ascidians.  Hydroids  not  further  identified  were 
found  in  9  stomachs,  Abietinaria  sp.  in  8,  SertulareUa  sp.  in  4,  and 


56  NORTH   AMERICAN   FAUNA.  [No.  46 

Thuiaria  sp.  in  1.  Bryozoans  not  further  determined  were  observed 
in  10  gizzards,  Crista  sp.  in  3,  Gellepora  surcularis  in  2,  and  Menipea 
pribilofi  in  1.  All  of  the  sea  spiders,  found  in  3  stomachs,  belonged 
to  a  single  species,  Ammothea  pribilofensis,  and  the  ascidians  or 
sea  squirts,  found  in  two  gizzards,  were  identical;  they  were  found 
to  be  Boltenia  ovifera.  Foraminifera,  the  only  representatives  of 
the  Protozoa,  or  simplest  animals  identified,  were  obtained  from  5 
gizzards. 

The  percentage  (7.47)  of  marine  algae  in  the  stomach  contents 
of  Pacific  eiders  is  so  high  as  to7  suggest  that  these  plants  are  in  part 
sought  for,  not  all  incidentally  taken,  as  some  of  them,  growing  on 
hermit  crabs  and  their  shelly  homes,  undoubtedly  are.  Algae  chiefly 
with  this  latter  habit  were  plume  algae  (Ceramiaceae),  identified 
as  Ptilota  pectinata  in  6  cases,  P.  asplenoides  in  4,  and  Ptilota  sp.  in 
12.  Coralline  algae  were  found  in  one  stomach,  and  a  single  seed  of 
the  rye  grass  (Ely mm  villosissimus)  in  another. 

Somateria  spectabilis  (Linnaeus).     King  Eider. 

The  beautiful  king  eider,  although  it  appears  to  be  of  regular  oc- 
currence nearly  throughout  the  year,  and  is  usually  more  or  less 
abundant  in  winter,  has  only  recently  been  included  in  the  list,  hav- 
ing been  recorded  by  Evermann  (1913,  p.  17)  from  specimens  taken 
by  M.  C.  Marsh  on  St.  Paul  Island  on  February  4,  and  March  9,  1912. 
The  species  is  present  in  small  numbers  throughout  the  summer,  but 
has  not  been  known  to  breed.  In  1914  it  was  observed  by  the  writer 
on  several  occasions  as  follows:  St.  Paul  Island,  July  19,  10  seen  at 
Northeast  Point ;  August  17,  1  at  Northeast  Point ;  August  24,  1.  St. 
George  Island,  August  4,  1  seen  at  Zapadni.  Hanna  sent  in  for 
examination  the  stomachs  of  three  shot  on  St.  George  Island,  May 
3,  1917. 

I  have  no  dates  of  occurrence  in  early  fall;  Hanna's  notes  for  St. 
Paul  Island,  fall  of  1914,  follow:  November  27,  2  seen;  December  4, 
75;  December  13,  50. 

The  species  is  sometimes,  perhaps  usually,  common  about  the 
islands  in  winter,  especially  when  pack  ice  is  present.  On  March 
29,  1911,  Hahn  noted  that  between  150  and  200  eiders,  nearly  all  of 
which  were  male  king  eiders,  were  shot  on  St.  Paul.  In  1915,  on 
St.  Paul,  Hanna  recorded  the  bird  as  follows :  January  10,  15  seen ; 
February  18,  abundant;  March  5,  about  100  in  the  ice  pack;  April 
6,  1 ;  April  15,  40 ;  April  18,  6  (near  shore)  ;  April  24,  15 ;  May  10, 
14.  H.  P.  Adams,  on  St.  George,  noted  the  bird  on  January  13,  Feb- 
ruary 4,  and  March  23,  1917. 

There  are  numerous  specimens  from  St.  Paul  Island  in  the  collec- 
tion of  the  Biological  Survey,  mainly  taken  by  W.  L.  Hahn  and  M. 
C.  Marsh.     From  their  catalogues  the  following  data  have  been  cop- 


1923.1  BIRDS   OF    THE   PRIBILOF   ISLANDS.  57 

ied:  Male,  March  30,  1911,  Hahn:  length,  22  inches;  extent,  34;  bill 
reddish  at  base,  shading  to  pale  horn  color  at  tip;  caruncle  deep 
lemon;  toes  pale  lemon,  webs  blackish.  Male,  January  7,  1912, 
Marsh :  Length,  22^  inches ;  extent,  38| ;  iris  dark  brown,  processes 
of  bill  yellow,  shading  into  yellow-red  of  bill,  which  is  pale  at  tip; 
legs  and  feet  yellow,  dark  at  joints,  webs  blackish.  Immature  male, 
February  4,  1912,  Marsh :  Length,  21f  inches,  extent,  36f ;  iris  dark 
brown,  processes  of  bill  light  yellow ;  bill  mostly  drab,  nail  darker ; 
feet  dingy  yellowish,  webs  dark.  Female,  February  1,  1912,  Marsh : 
Length,  19|  inches,  extent,  36;  iris  dark  brown;  bill  dusky,  pale  be- 
low and  about  nail;  feet  dusky  reticulated,  on  yellowish  ground. 
Female,  February  4,  1912,  Marsh :  Iris  dark  brown ;  processes  of  bill 
black,  pale  band  back  of  nail,  above  and  below ;  feet  yellowish,  dusky 
reticulated. 

Food. — To  illustrate  the  food  habits  of  the  king  eider  in  the  Prib- 
ilof  area  we  have  had  for  examination  about  the  same  number  of 
stomachs  as  of  the  Pacific  eider,  namely,  44  well-filled  ones  and  9 
others.  The  nature  of  the  food  also  proves  to  be  very  similar,  the 
principal  items  and  their  proportions  being:  Mollusks,  34.36  per 
cent;  crustaceans,  28.2  per  cent;  sea  urchins,  9.54  per  cent;  starfishes, 
2.59  per  cent;  and  algae,  9.88  per  cent.  The  king  eider  takes  fewer 
mollusks  and  a  much  lower  percentage  of  starfishes  than  the  Pacific 
eider,  but  makes  up  for  these  in  part  by  greater  consumption  of  bar- 
nacles and  fishes.  Mr.  Hanna  notes  that  this  species  can  descend  to 
the  bottom  in  30  fathoms  of  water,  as  evidenced  by  the  fact  that  a 
bird  killed  just  after  coming  to  the  surface  of  water  of  this  depth 
had  freshly  swallowed  mollusks  in  its  gullet. 

Like  its  congener,  this  eider  draws  its  shellfish  fare  from  diverse 
branches  of  the  molluscan  kingdom.  Of  the  bivalves  eaten  the 
favorite  was  the  common  blue  mussel  (Mytilus  edulis),  which 
occurred  in  21  stomachs,  and  next  in  preference  was  Modiolaria 
verrucosa,  taken  by  12  birds.  Other  bivalves  eaten  included  the 
scallop  {Pecten  islandicus)  and  Cardium  ciliatum,  Musculus  sp.,  My  a 
intermedia,  Rochefortia  sp.,  and  Saxicava  arctica.  A  single  limpet 
(Acmaea  mitra)  was  identified  in  the  food,  as  also  were  numerous 
sea  snails,  of  which  Plicifusus  spitzbergensis,  found  in  7  stomachs, 
was  of  most  frequent  occurrence.  The  others  identified  are:  Astyris 
sp.,  Borcotrophon  (?)  multico status,  Bucoinum  fischerianum,  Bucci- 
num  sp.,  Cerithiopsis  sp.,  Littorina  atkana,  L.  grandis,  L.  gron- 
landica,  Littorina  sp.,  Machaeroplax  cinereus,  Margarites  helicina, 
Natica  clausa,  N.  operculina,  Neverita  recluziana,  Nucella  lima,  Tri- 
chotropis  insignis,  and  T .  nucellatus.  Sea  slugs,  as  the  creeping 
mollusks  with  segmented  dorsal  mail  may  be  called,  also  were  taken 
by  the  king  eiders,  and  the  following  forms  were  identified :  Chiton 
marmoreus,  Chiton  sp.,  and  Schizoplax  brandtii. 


58  NORTH   AMERICAN  FAUNA.  [No.  46 

The  crustaceans  eaten  by  the  king  eiders  studied  consisted  rather 
more  of  crabs  and  less  of  amphipods  than  in  the  case  of  the  Pacific 
eider.  Crabs  were  slightly  more  than  a  fifth  of  the  total  food,  most 
of  them  being  hermits.  One  species  of  this  group,  Dermaturus 
mandtii,  was  found  in  no  fewer  than  33  of  the  stomachs,  7  individuals 
being  counted  in  one  of  them.  Other  hermit  crabs  identified  were 
Hapalogaster  grebnitzkii,  Pagurus  trigonocheirus,  and  Pagurus  sp. 
Other  crabs  eaten  were  the  spider  crab  (Hyas  coarctatus) ,  the  mud 
crab  (Telmessus  cheiragonus),  and  crabs  of  the  family  Xanthidae. 
Amphipods  were  much  less  prominent  in  number  and  variety  in 
stomachs  of  the  king  eider  than  in  those  of  its  relative,  and  only 
3  forms  were  identified,  namely,  Gammarus  pribUofensis,  Hyale  sp., 
and  Orchomenella  minuta.  A  few  isopods  were  eaten,  among  which 
one  species,  Synidotea  nebulosa,  was  determined;  and  also  a  few 
shrimps  {Spirontocaris  polaris). 

Sea  urchins,  in  this  species,  as  well  as  in  the  Pacific  eider, 
are  a  prominent  element  of  the  food.  The  kinds  eaten  are  the 
same  and  they  occurred  in  the  following  numbers  of  stomachs :  Sea 
urchins  not  further  identified,  19;  E chinardchnius  parma,  9;  and 
tStrongylocentrus  drobachiensis,  2.  Starfishes,  for  some  reason 
comparatively  neglected  by  this  eider,  were  found  in  6  stomachs  and 
only  in  one  case  were  identified  as  to  genus  {Leptasterias) . 

Of  the  lesser  items  of  animal  food,  fishes  and  barnacles  are  the 
most  important.  The  only  fish  remains  at  all  classifiable  were  some 
bculpin  bones.  Three  batches  of  eggs,  not  absolutely  known  to  be 
those  of  fishes,  were  added  to  the  undoubted  fish  remains,  and  make 
a  percentage  of  the  total  food  of  3.84  per  cent.  Barnacles  (2.34  per 
cent)  are  next  in  importance,  being  found  in  18  stomachs;  a  single 
species,  Balaams  rostratus,  was  identified. 

Both  tubicolous  and  nereid  marine  worms  were  eaten,  the  latter 
occurring  in  20  stomachs;  the  percentage  of  worms  in  the  food  was 
1.7  per  cent.  While  some  of  them  occurred  in  numerous  gizzards,  the 
remaining  articles  of  animal  food  in  no  case  amounted  to  as  much  as 
1  per  cent  of  the  total  subsistence.  Hydroids  not  further  identified 
were  found  in  13  stomachs,  and  also  the  following  genera  of  this 
group,  in  the  number  of  gizzards  indicated:  Abietinaria  8,  Sertu- 
larella  4,  and  Thuiaria  3.  Bryozoa  similarly  not  further  named, 
occurred  in  16  gizzards;  those  identified  were  Cellepora  surcularis 
in  2,  and  Menipea  pribilofi  and  Myriozoum  subgracile  in  1  each, 
Sea  spiders  (Pycnogonida) ,  found  in  2  stomachs,  a  sea  squirt 
(Boltenia  ovifera)  in  1,  and  Foraminifera  in  5,  complete  the  list  of 
animal  foods. 

The  vegetable  food  (9.88  per  cent)  consisted  exclusively  of  algae. 
That  not  further  identified,  chiefly  kelp  or  laminar  algae,  was  found 


1923.]  BIRDS   OF   THE   PRIBILOF   ISLANDS.  59 

in  19  stomachs.  Plume  algae  {Ptilota  pectinata  in  17  stomachs,  P. 
asplenoides  in  15,  and  Ptilota  sp.  in  8)  were  a  prominent  article  of 
diet,  and  two  kinds  of  coralline  algae  (Corallina  probably  aovleatu 
and  Amphiroa  cretacea)were  each  found  in  a  single  stomach. 

Oidemia  deglandi  dixoni  Brooks.     Pacific  White-winged  Scoter. 

During  the  past  few  years  the  Pacific  white-winged  scoter  has 
come  to  be  recognized  as  a  more  or  less  regular  migrant  and  winter 
visitor.  G.  Dallas  Hanna  has  put  the  species  on  record  (1919a,  p. 
176),  and  later  (1920a,  p.  252)  writes  as  follows: 

Four  specimens  of  the  western  white-winged  scoter  have  been  taken  on  the 
Pribilofs  during  the  last  three  years.  All  were  females  but  this  is  merely  an 
accidental  circumstance  because  males  have  been  seen.  The  first  specimen  was 
secured  on  October  30,  1916,  at  St.  George  Island  by  the  writer.  The  next 
was  taken  on  November  15,  1916,  at  the  same  island  by  Dr.  H.  P.  Adams, 
formerly  physician  of  the  U.  S.  Bureau  of  Fisheries.  Another  was  secured  at 
St.  George  Island  on  February  4,  1917,  and  one  at  St.  Paul  Island,  February  8, 
1918,  both  by  the  writer. 

Hanna  also  noted  the  bird  as  abundant  at  the  margin  of  the  ice 
pack  off  St.  George  on  January  25,  1917. 

The  species  is  of  regular  occurrence  about  the  islands  in  winter. 
It  has  been  seen  on  several  occasions  before  any  specimens  were 
secured  but  was  not  recorded  because  positive  specific  identification 
could  not  be  made. 

The  specimens  above  mentioned  are  all  in  the  collection  of  the 
U.  S.  National  Museum. 

Food. — The  stomachs  of  2  white-winged  scoters,  1  collected  on 
St.  George  Island,  February  4,  1917,  and  the  other  on  St.  Paul, 
February  8,  1918,  have  been  examined  and  the  food  proved  to  be 
about  equally  divided  between  amphipods  and  bivalves.  None  of 
the  former  were  identified,  and  the  mussels  were  Saxicava  arctica 
principally,  with  a  trace  of  Mytilus  edulis.  Other  items  of  food 
sparingly  represented  were  sea  urchin,  bryozoan,  and  hydroid. 

Chen  hyperborea  hyperborea  Pallas.     Snow  Goose. 

The  snow  goose  was  added  to  the  Pribilof  list  by  G.  Dallas  Hanna 
(1919a,  p.  176,  and  1920a,  p.  251),  who  obtained  an  individual  shot 
by  a  native  at  Northeast  Point,  St.  Paul  Island,  September  16, 
1916.  "Another  bird  was  killed  the  following  year  on  the  same 
island  but  since  it  belonged  clearly  to  the  same  species  and  time  did 
not  permit  of  its  preparation  it  was  not  preserved"  (1.  c,  p.  251). 

The  specimen  from  Northeast  Point  is  now  in  the  collection  of 
the  U.  S.  National  Museum,  and  the  head  of  one  shot  on  St.  George 
by  Mr.  Hanna  is  in  the  Biological  Survey  collection. 


60  NORTH   AMERICAN   FAUNA.  [No.  46 

Anser  albifrons  albifrons  (Scopoli).     White-fbonted  Goose. 

The  white- fronted  goose  was  first  noted  from  the  islands  by 
Palmer,  who  saw  2  on  a  small  pond  near  the  village  on  St.  Paul 
Island,  June  11,  1890.  Two  others  were  seen  by  him  later  (1899,  p. 
380).  The  only  other  records  seem  to  be  those  of  G.  Dallas  Hanna, 
who  saw  1  individual  at  Garden  Cove,  St.  George  Island,  September 
28,  1913,  and  2  individuals  at  Halfway  Point,  St.  Paul  Island,  on 
May  18,  1915. 

Food. — The  contents  of  two  stomachs  of  the  white-fronted  goose 
collected  on  St.  George  Island /in  May  have  been  studied  and  found 
to  consist  of  leaves  of  grass,  99  per  cent ;  and  those  of  saxifrage  and 
chickweed,  1  per  cent. 

Branta  canadensis  hutchinsi  (Richardson).     Hutchins  Goose. 

So  far  as  known  to  the  writer  the  Hutchins  goose  has  been  detected 
only  once  on  the  Pribilofs,  by  G.  Dallas  Hanna.  He  has  reported  the 
occurrence  (1919a,  p.  176,  and  1920a,  p.  251)  : 

A  female  of  this  subspecies  was  shot  and  preserved  on  St.  Paul  Island,  May 
12,  1918.  It  is  considerably  larger  than  the  cackling  goose  which  ordinarily 
comes  to  the  islands  each  spring  and  fall,  and  unlike  the  latter  form  there  is  no 
sharp  demarcation  in  the  coloration  of  the  under  parts  (p.  251). 

The  specimen  above  mentioned  is  now  No.  255152  of  the  bird  collec- 
tion of  the  U.  S.  National  Museum. 

Branta  canadensis  minima  Ridgway.     Cackling  Goose. 

Elliott  secured  a  specimen  of  the  cackling  goose  (62526,  male  adult, 
U.  S.  Nat.  Mus.)  on  St.  Paul  Island,  May  14,  1872.  This  specimen 
was  listed  by  Palmer,  who  also  (1899,  p.  381)  records  individuals 
observed  on  St.  Paul  on  June  25  and  June  28,  1890.  Palmer  (1.  c.) 
and  Hanna  (1919b,  p.  106)  consider  the  bird  a  regular  migrant. 

Hahn  reported  geese  of  this  type,  assumed  to  be  referable  to  the 
present  race,  on  St.  Paul  Island,  as  follows:  September  15,  1910, 
5;  September  24,  1;  October  2,  8;  May  30,  1911,  1.  Hanna  reported 
this  race  as  follows :  St.  Paul,  1915 :  May  14,  8  flew  over  village ; 
May  16,  4  observed;  May  18,  6  in  pairs,  2  killed,  Halfway  Point. 

Branta  nigricans  (Lawrence).     Black  Brant. 

The  black  brant  was  first  recorded  from  the  Pribilofs  by  Dall 
(1874,  p.  275),  who  says,  in  connection  with  the  occurrence  of  the 
bird  on  the  western  Aleutians :  "  It  has  also  been  reported  as  a 
casual  visitor  at  the  Pribiloff  Islands." 

Specimens  were  first  collected  by  Hanna,  and  recorded  by  him 
(1916,  p.  401).  His  first  one,  a  male,  was  taken  at  Bear  Lake,  St. 
George  Island,  September  28,  1913,  and  is  now  No.  242535  of  the 
U.  S.  National  Museum  collection.     Another  individual  was  seen 


1923.]  BIRDS   OF   THE   PRIBILOF   ISLANDS.  61 

at  this  time.  He  states  (1.  c.)  that  small  flocks  and  single  birds 
seem  to  occur  each  spring  and  fall,  and  he  considers  that  this  species, 
is  more  common  than  any  other  goose  excepting  the  emperor. 
During  the  following  year,  on  St.  Paul  Island,  Hanna  observed  the 
species,  as  follows:  September  17,  5  seen,  1  secured;  September  20, 
2  seen;  October  8,  2;  October  25,  4  at  Northeast  Point. 

Notes  taken  from  the  St.  Paul  Island  log  by  W.  L.  Hahn,  refer- 
ring to  black  geese,  undoubtedly  relate  to  this  species.  These  notes 
record  5  individuals  shot  from  a  flock  on  October  7,  1902;  and  10 
shot  April  18,  1905. 

Food. — The  contents  of  a  single  stomach  of  the  black  brant  col- 
lected at  St.  Paul  Island  on  September  17,  1914,  consisted  exclusively 
of  a  filamentous  alga  (C haetomorpha  cannaban). 

Philacte  canagica  (Sevastianoff).     Emperob  Goose. 

The  beautiful  emperor  goose  seems  to  be  of  regular  occurrence  in 
spring  and  fall  on  both  the  larger  islands.  It  was  first  recorded  by 
Elliott,  who  witnessed  the  capture  of  an  entire  flock,  which  landed 
so  exhausted  that  the  natives  were  able  to  take  them  by  hand  (1874, 
unpaged,  and  1875,  p.  189). 

Townsend  (1887,  p.  99)  speaks  of  3  or  4  seen  on  St.  Paul  Island 
in  September  [1885] ;  Palmer  records  1  killed  on  St.  George  Island, 
July  16,  1890.  Hahn  found  notes  in  the  St.  Paul  log  recording  the 
shooting  of  1  on  November  29,  1904,  and  2  on  September  24,  1908; 
and  Hahn  himself  reports  2  seen  on  St.  Paul  Island  May  10,  1911. 

Hanna  has  made  the  most  extensive  observations,  as  follows:  St. 
George  Island,  1913,  December  21,  4  seen ;  1914,  April  8,  3  seen ;  April 
22,  8 ;  May  19,  8  seen,  2  shot,  not  skinned  owing  to  scarcity  of  food 
among  the  natives,  head  and  feet  saved;  1917,  May  10,  2  shot.  St. 
Paul  Island,  1914,  September  14,  6 ;  September  16,  20 ;  September  20, 
2;  September  22,  2;  October  31,  15;  November  4,  1  (specimen)  ;  No- 
vember 26,  3;  1915,  March  20,  5;  May  18  (2,  in  village  cove)  ;  June 
14,  7  pairs;  June  16,  4  pairs.  H.  P.  Adams,  on  St.  George,  noted 
the  bird  on  September  22  and  26,  1915,  and  April  26  and  28,  1917. 

Food. — H.  W.  Elliott  has  reported  the  food  of  the  emperor  goose 
to  consist  of  Mytilus  edulis  and  other  shellfish  (1874,  unpaged,  and 
1875,  p.  190).  Such  food  habits  would  be  a  radical  departure  from 
the  standard  vegetarianism  of  geese  in  general,  and  there  is  no  direct 
evidence  that  they  exist.  Reliable  evidence  points  to  the  emperor 
goose  sharing  the  plant-feeding  proclivities  of  its  allies  as  the  entire 
food  of  5  birds  of  this  species  collected  on  the  Pribilofs  was  marine 
algae  of  the  type  known  as  sea  lettuce,  identified  in  one  case  as  Dilsea 
integra. 

10S7310—  23 5 


62  NORTH   AMERICAN   FAUNA.  [No.  46 

Olor  columbianus  (Ord).     Whistling  Swan. 

The  whistling  swan  was  first  recorded  by  Palmer,  who  says  (1899, 
p.  381)  : 

Several  were  seen  on  St.  George  in  the  fall  of  1889  and  three  rested  on  the 
shore  at  Halfway  Point,  on  St.  Paul,  during  the  same  autumn. 

Hahn  took  the  following  from  the  St.  Paul  Island  log:  October 
9,  1888,  1  killed;  May  15,  1903,  1  shot;  October,  1909,  2  shot;  weight, 
8-|  and  9  pounds,  respectively.  Hahn  himself  recorded  1  seen  by  a 
native,  October  2,  1910;  and  2  reported  May  7,  1911. 

Hanna  (1916,  p.  402)  has  recorded  1  shot  by  a  native  on  a  high 
hill  on  St.  George  Island,  October  10,  1913.  Its  measurements,  re- 
duced to  millimeters,  were  as  follows:  Length,  1,151  mm.;  extent, 
1,780  mm.  The  head  and  foot  of  this  one  were  saved  as  a  specimen. 
H.  P.  Adams  records  the  bird  on  St.  George  as  follows :  October  16, 
1915,  noted;  October  17,  1915,  1  killed. 

Family   GRUIDAE. 

Megalornis  canadensis  (Linnaeus.)8    Little  Brown  Crane. 

The  little  brown  crane,  which  has  several  times  been  observed  in 
early  summer,  was  added  to  the  Pribilof  list  by  Townsend  (1887, 
p.  99).    He  says: 

While  at  St.  Paul  Island  [summer,  1S85]  some  natives  came  to  me  and  re- 
ported having  seen  a  large  long-legged  bird  wading  in  a  shallow  pond  near  the 
village,  and  gave  me  a  large  feather  which  the  bird  dropped  in  its  flight. 

Palmer's  original  notes  follow : 

Jake,  a  well-known  Aleut  sealer,  told  me  on  June  28  [1890],  that  he  had 
seen  3  cranes  that  morning  north  of  Kaminista  Lake.  Going  there,  I  found  5, 
but  they  were  very  wild.  They  remained  in  the  neighborhood  until  after 
August  1,  as  I  saw  them  frequently  picking  up  insects  on  the  tundra.  One  was 
caught  in  a  trap  near  the  village  in  June,  18SS  (1899,  p.  398). 

According  to  the  St.  Paul  Island  log  (notes  transcribed  by  Hahn), 
two  "  blue  cranes "  were  seen  on  the  tundra  September  13  and  14, 
1908.    These  birds  were  almost  certainly  the  present  species. 

Hanna  observed  3  cranes  at  Big  Lake,  St.  Paul  Island,  on  May 
12,  1915,  and  1  on  the  tundra  near  Rocky  Point  Lake  on  June  26.  In 
May,  1919,  he  took  the  first  specimens,  which  he  recorded  as  follows 
(1920b,  p.  174) : 

Two  little  brown  cranes  (Grus  canadensis)  were  collected  on  St.  Paul  Island 
in  May,  1919.  This  species  occurs  in  small  numbers  regularly  during  migrations 
but  the  birds  are  rarely  shot.  The  previous  record  was  based  on  a  single 
feather  picked  up  on  the  tundra. 

Food.— -William  Palmer  stated  (1899,  p.  398)  that  he  saw  little 
brown  cranes  picking  up  insects  on  the  tundra  of  the  Pribilofs,  and 

_  • 

8  Grus  canadensis  of  the  A.  O.  U.  Check  List,  1910. 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  63 

G.  Dallas  Hanna  notes  that  the  two  specimens  collected  by  him  on 
St.  Paul  Island  had  their  crops  filled  almost  to  the  bursting  point 
with  blowfly  larvae  from  the  seal-killing  fields. 

Family  PHALAROPODIDAE. 
Phalaropus  fulicarius    (Linnaeus).     Red   Phalarope. 

The  red  phalarope  is  an  abundant  northward  migrant  in  early 
summer  and  again  appears  on  its  return  late  in  summer.  It  is  occa- 
sionally seen  in  midsummer,  but  has  not  been  found  breeding  on  the 
islands. 

The  earliest  spring  dates  are  from  specimens  taken  by  M.  C.  Marsh 
on  St.  Paul  Island,  May  16  and  31,  1912,  and  by  Hanna  on  St. 
George  May  19  and  23,  1914,  and  the  latter's  note  of  May  26.  1915, 
when  he  saw  large  flocks  crossing  between  St.  Paul  and  Otter  Islands. 
Palmer  collected  several  about  Polovina,  St.  Paul  Island,  between 
June  10  and  July  17,  1890,  but  thought  that  they  did  not  breed. 
Hanna  observed  2  at  Halfway  Point,  St.  Paul,  June  20,  1915,  and  a 
pair  each  on  St.  Paul  and  St.  George  throughout  the  summer  of 
1919,  but  found  no  evidence  of  nesting. 

Fall  migrants  begin  to  appear  in  mid-July.  Palmer  noted  the  first 
fall  arrivals  on  July  17,  1890,  I  saw  the  first  one  on  St.  George 
July  14,  1914,  and  on  July  16  found  hundreds  on  the  water  about 
Walrus  Island.  These  were  largely  still  in  breeding  plumage.  From 
that  date  until  we  left  the  Pribilofs.  the  last  of  August,  I  frequently 
observed  the  birds,  usually  in  small  numbers  but  sometimes  in  large 
flocks. 

During  the  autumn  the  birds  continue  to  be  observed  until  quite 
late.  Hahn's  notes  for  St.  Paul  for  1910  follow:  August  30,  about 
100  seen ;  a  few  to  September  7 ;  September  8,  about  1,000 ;  September 
9,  about  200;  September  11,  about  200;  September  12,  about  50;  Sep- 
tember 15  to  19,  a  few;  October  2,  many  seen  on  way  to  Walrus 
Island ;  October  15,  1 ;  October  19,  3 ;  October  20,  2.  On  January  18, 
1911,  Hahn  saw  one  on  the  water  off  Tolstoi  Point,  St.  Paul.  Hanna 
on  St.  George  records  the  following  observations  in  1913 :  September 
5,  4;  September  15,  25 ;  October  9,  about  100,  some  in  surf  but  mostly 
in  fresh-water  ponds;  October  18,  12.  His  notes  for  St.  Paul,  1914, 
follow :  September  20,  1 ;  October  25,  1  taken  at  Webster  Lake ;  No- 
vember 4,  1  in  surf  at  village  landing.  Specimens  taken  on  St.  Paul 
by  Hahn  and  Marsh,  in  1910,  1911,  and  1912,  measured  in  the  flesh 
as  follows :  Males,  length  7|  to  8|  inches ;  extent,  15^  to  15|. 
Females,  length,  9  to  9| ;  extent,  16£  to  17. 

Food. — The  red  phalarope,  one  of  the  swimming  snipe,  is  repre- 
sented in  the  present  collection  by  53  well-filled  and  2  nearly  empty 
stomachs.  While  the  phalaropes  habitually  alight  and  feed  in  water, 
it  is  evident  from  analyses  of  the  contents  of  these  stomachs  that  they 


64  NORTH   AMERICAN   FAUNA.  [No.  46 

feed  also  on  land,  where  they  must  obtain  most  of  the  flies  and  beetles 
they  eat.  The  principal  food  items  found  in  the  53  stomachs  were : 
Crustaceans,  38.29  per  cent;  flies,  21.69  per  cent;  fishes,  18.18  per 
cent;  caddisflies,  10.15  per  cent;  and  beetles,  9.86  per  cent. 

Of  the  crustaceans  eaten,  30.45  per  cent  of  the  total  of  38.29  per 
cent  were  identified  as  amphipods  and  probably  part  of  the  uniden- 
tified belonged  to  the  same  group.  Amphipods  were  found  in  22 
stomachs,  but  could  be  determined  to  the  species  in  only  2  cases,  these 
being  Gam/tnarus  pribilofensis  and  Pontogeneia  inermis.  Another 
group  of  crustaceans,  while  contributing  only  a  trace  to  the  food  of 
the  red  phalarope,  is  of  considerable  interest  as  an  item  of  avian  diet, 
because  its  members  are  so  small,  bordering  on  microscopic  size. 
These  are  the  water  fleas  (Daphniidae),  which,  as  ordinarily  seen 
by  human  observers,  appear  like  dancing  motes  in  waters  pierced  by 
the  sunbeams  in  which  they  love  to  gather.  The  contribution  of  these 
little  animals  to  the  food  of  the  phalarope  is  their  egg  cases  (ephip- 
pia),  each  a  twin  sac  inclosing  the  two  relatively  large  black  eggs. 
These  were  found  in  6  stomachs,  no  fewer  than  50  of  them  being 
taken  from  one  stomach,  and  in  this  single  instance  they  amounted 
to  5  per  cent  of  the  contents. 

Two-winged  flies  (Diptera)  and  their  larvae  and  pupae  were 
found  in  more  than  half  of  the  gizzards  examined  and  constituted 
21.69  per  cent  of  the  total  food.  The  kinds  most  frequently  taken 
were  members  of  the  dung-fly  family  (Scatophagidae)  and  the  adults 
and  larvae  of  midges  (Chironomidae).  Seventy  of  these  larvae  were 
found  in  a  single  stomach.  The  determinations  of  flies,  other  than 
as  just  mentioned,  were:  Anthomyiidae,  blowflies  (Muscidae),  dung- 
flies  {Scatophaga  <"rinita,  S.  dasythrix,  and  Scatophaga  sp.),  soldier- 
flies  (Stratiomyidae),  and  crane-flies  (Tipulidae). 

Fishes  were  the  next  most  important  element  of  the  food  of  the 
red  phalarope,  being  found  in  15  stomachs  and  composing  18.18  per 
cent  of  the  total  subsistence.  All  taken  were  small  sculpins  (Cotti- 
dae).  Caddisflies  and  their  larvae  were  taken  by  14  birds  and 
formed  10.15  per  cent  of  the  food.  These  live  in  fresh  water  and 
the  larvae  construct  cases  of  a  variety  of  materials  in  which  they 
pass  the  immature  stages. 

The  only  remaining  item  of  any  importance  in  the  diet  of  this 
species  is  beetles  (9.86  per  cent).  Ground  beetles  of  the  genus 
Pterostichus  were  taken  more  frequently  than  any  other  kind,  oc- 
curring in  11  stomachs.  Another  beetle  (Amara  sp.)  of  the  same 
family  was  identified,  and  in  addition  the  following:  Rove  beetles, 
including  Atheta  sp.,  Hadrotes  sp.,  and  Olophrum  fuscum;  diving 
water  beetles,  including  Ilybius  angiustior;  and  Aegialites  calif orni- 


1923.]  BIRDS   OF   THE   PRIBILOF   ISLANDS.  65 

cus,  a  beetle  of  a  peculiar  family  having  a  few  species  along  our 
western  coast. 

Hymenoptera,  mostly  small  parasitic  wasps  (Ples-lgnathus  sp.), 
the  only  other  insects  eaten,  were  found  in  5  stomachs  and  spiders 
in  1.  Mollusks  were  taken  by  two  birds  and  marine  worms  (Nerei- 
dae)  by  one. 

Vegetable  matter  constituted  less  than  half  of  1  per  cent  of  the 
contents  of  the  53  stomachs  of  red  phalaropes  examined,  and  is  to 
be  considered  as  an  accidental  rather  than  genuine  article  of  food. 
It  consisted  entirely  of  seeds,  of  which  those  of  violet  (Viola  sp.) 
were  found  in  5  stomachs,  sedge  (Carex  sp.)  in  2.  and  the  following 
in  1  each:  Crowberry  (Empetrum  nigriim),  lousewort  (Pedlcu- 
laris  sp.),  and  bog  bean  (Menyantlies  trifoliata). 

Lobipes  lobatus  (Linnaeus).     Northern  Phalarope. 

The  northern  phalarope  is  apparently  a  fairly  abundant  migrant 
on  the  Pribilofs,  although  I  have  no  definite  information  on  this 
point.    It  breeds  in  small  numbers  on  both  St.  George  and  St.  Paul. 

Elliott  first  recorded  the  bird  (1874,  unpaged,  and  1875,  p.  181) 
and  took  downy  young  on  St.  George  in  1873.  Palmer  found  the 
birds  nesting  near  Rocky  Point,  St.  Paul,  July  2,  1890,  also  collect- 
ing newly  hatched  young.  I  observed  several,  apparently  nesting, 
near  the  latter  locality  on  June  27,  1914.  Hanna  saw  four  pairs 
near  the  same  place  on  June  14,  1915.  The  earliest  spring  date  is  of 
a  specimen  taken  by  Hanna  on  St.  George,  May  23,  1914.  The  latest 
dates  are  also  represented  by  specimens:  August  15,  1914,  St.  Paul, 
Hanna;  and  August  17,  1897,  St.  Paul,  Greely  and  Snodgrass. 

A  female  specimen  collected  by  Marsh  on  St.  Paul  Island,  June  1, 
1912,  measured  in  the  flesh :  Length,  7f  inches ;  extent,  14^. 

Family   SCOLOPACIDAE. 

Limnocryptes  gallinula  ( Linnaeus L     European  Jacksnipe. 

The  only  record  of  the  European  jacksnipe  as  an  inhabitant  of  the 
Pribilofs  is  based  on  a  specimen  taken  by  G.  Dallas  Hanna  and  re- 
corded by  him  (1920b,  p.  173)  as  follows: 

Some  time  during  the  spring  of  1919,  probably  in  April,  a  native  on  St.  Paul 
Island  secured  one  of  these  birds  and  very  kindly  kept  the  skin  for  me  until 
my  arrival.  The  identification  was  made  by  Mr.  Joseph  Mailliard,  of  the  Cali- 
fornia Academy  of  Sciences.  This  is  a  new  record  for  the  Pribilof  Islands  and 
for  North  America. 

The  specimen  is  deposited  in  the  collection  of  the  California  Acad- 
emy of  Sciences.  The  species  breeds  generally  in  northern  Eurasia 
east  to  the  Kolyma  River,  and  occurs  in  migration  and  in  winter 
south  to  India  and  Burma,  and  occasionally  to  Japan  and  Formosa. 


66  NORTH   AMERICAN   FAUNA.  [No.  46 

Limuodromus   griseus   scolopaceus    (Say).*     Long-billed   Dowitcher. 

A  female  specimen  of  the  long-billed  dowitcher  taken  on  St.  Paul 
Island,  September  18,  1919,  by  G.  Dallas  Hanna,  and  recorded  by  him 
(1920b,  p.  173)  constitutes  the  first  record  for  the  Pribilofs. 

Later  Mr.  Hanna  took  2  additional  specimens,  a  male  and  a  female, 
at  Northeast  Point,  St.  Paul  Island,  August  29,  1920.  These  are  re- 
corded by  Mailliard  and  Hanna  (1921,  p.  95).  All  these  specimens 
are  in  the  collection  of  the  California  Academy  of  Sciences. 

Food. — Two  stomachs,  of  the  two  specimens  last  mentioned  from 
St.  Paul  Island,  have  been  examined  and  their  contents  were  almost 
exclusively  the  larvae  of  midges  (Chironomidae),  of  which  there  were 
more  than  75  in  one  gizzard  and  more  than  100  in  the  other.  Vege- 
table debris,  amounting  to  3  per  cent  by  bulk  of  the  stomach  contents, 
also  was  present,  and  it  probably  was  picked  up  incidentally  with 
the  midge  larvae.  Included  in  the  vegetable  matter  were  seeds  of 
bottle  brush  (Hippuris  vulgaris),  sedge  (Carex  sp.),  and  water 
chickweed  (Montia  fontana). 

Arquatella  maritima  couesi  Ridgway.     Aleutian  Sandpiper. 

The  Aleutian  sandpiper,  which  breeds  mainly  on  the  Alaska 
Peninsula,  visits  the  Pribilofs  in  the  autumn  and  lingers  in  small 
parties  about  the  icy  shores  until  winter.  It  has  also  been  observed 
early  in  spring,  sometimes  about  the  pack  ice.  It  was  first  ascribed  to 
the  islands  by  Seale  (1898,  p.  139),  from  specimens  taken  on  St.  Paul 
Island  in  the  late  summer  of  1897  by  R.  E.  Snodgrass  and  A.  W. 
Greely.  There  is  reason  to  believe,  however,  that  this  record  is  based 
on  mis-identified  specimens.  These  are  now  supposed  to  be  in  the  col- 
lection of  Leland  Stanford  University.  At  the  request  of  the  writer, 
Dr.  Walter  K.  Fisher  kindly  attempted  to  look  up  the  pertinent  speci- 
mens but  was  able  to  find  only  one  which  could  reasonably  be  sup- 
posed to  be  in  part  the  basis  of  the  record.  This  specimen  (No.  3498, 
an  immature  female,  labeled  T.  couesi,  taken  July  25,  1897)  he  kindly 
forwarded  for  examination.  This  proved  to  be  an  undoubted  exam- 
ple of  the  breeding  species  A.  m.  ptilocnemis,  and  is  in  the  usual 
plumage  of  the  immature  bird  at  this  season.  It  is  possible  that  later 
research  will  bring  to  light  other  specimens  of  the  Snodgrass-Greely 
collection  which  are  actually  referable  to  -4.  m.  couesi,  but  the  writer 
is  rather  inclined  to  the  belief  that  they  will  all  prove  to  be  the  breed- 
ing form,  which  gathers  abundantly  on  the  beaches  in  late  summer, 
and  includes  many  small  specimens  which,  without  close  examina- 
tion, might  be  considered  to  be  the  Aleutian  form. 

However  this  may  be,  the  species  has  been  definitely  added  to  the 
list  by  Hanna,  who  took  specimens  on  St.  Paul  Island  in  the  autumn 


'Macrorhamphus  y.  scolopaceus  of  the  A.  O.  U.  check  list,  1910. 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  67 

of  1914  and  in  the  spring  following.  His  notes  of  occurrence  are  as 
follows:  1914,  September  16,  4  observed  (1  taken  now  in  the  collec- 
tion of  the  Biological  Survey)  ;  October  IT,  1  taken  (female  No. 
237527,  U.  S.  Nat.  Mus.)  ;  October  25,  4  seen  at  Northeast  Point;  No- 
vember 16,  8  seen;  November  27,  4  seen  at  Lukanin  beach,  2  taken 
(males,  Nos.  237525  and  237526,  U.  S.  Nat.  Mus.).  1915,  March  5,  1 
seen  in  pack  ice;  April  24,  1  taken  at  Northeast  Point  (No.  237524,  U. 

5.  Nat.  Mus.).  This  last  specimen  has  begun  to  acquire  the  reddish 
summer  feathering  on  the  back  and  the  sides  of  the  breast. 

Mailliard  and  Hanna  (1921,  p.  94)  have  recently  recorded  a  speci- 
men, not  sexed,  taken  on  St.  George  Island,  February  12,  1917.  This 
is  in  the  collection  of  the  California  Academy  of  Sciences. 

Arquatella  maritima  ptilocnemis  (Coues).     Pribilof  Sandpiper. 

Tringa  ptilocnemis  Coues,  in  Elliott.  Report  Seal  Islands,  Alaska,  1874,  un- 
paged;  1875,  reprint,  p.  182  (type  from  St.  George  Island). 

The  Pribilof  sandpiper  breeds  in  some  numbers  on  both  St.  Paul 
and  St.  George.  It  is  absent  from  the  Pribilofs  during  the  colder 
months,  leaving  in  October,  and  returning  in  middle  or  late  April. 

Veniaminof  in  1840  (Trans,  by  Elliott,  1874  unpaged,  and  1875, 
p.  242)  listed  "  a  few  kinds  of  Tringa  "  as  occurring  on  the  Pribilofs, 
and  may  reasonably  be  supposed  to  have  observed  the  only  species  of 
the  group  which  breeds;  the  species,  however,  was  first  definitely 
recorded  by  Dall,  who  obtained  a  specimen  on  St.  George  and  listed 
it  under  the  name  Tringa  maritima  (1869,  p.  291).  Later  both  Dall 
and  Coues  listed  the  species  as  Tringa  crassirostris,  but  Coues,  as- 
sured of  its  novelty,  in  a  footnote  (in  Elliott,  1874,  not  paged,  p.  182 
of  1875  reprint)  gave  it  the  specific  name  it  now  bears.  About  the 
same  time,  Harting  (1874,  p.  243)  bestowed  upon  it  another  name, 
Tringa  gracilis. 

Hanna  in  1914  saw  seven  individuals  at  Zapadni,  St.  George 
Island,  on  April  14.  His  records  of  observations  on  St.  Paul  in  the 
spring  of  1915  are  as  follows:  April  24,  about  50  seen  in  flocks;  a 
male  taken  on  this  date  had  nearly  finished  acquiring  the  breeding 
plumage,  but  still  retained  a  few  traces  of  the  winter  feathering; 
April  26,  about  50  (in  flocks)  ;  May  2,  about  30  (pairing)  ;  May  4,  18 
(about  all  paired)  ;  May  9,  20  (about  all  paired)  ;  May  18,  12  (in 
pairs)  ;  May  28,  8  (in  pairs). 

This  sandpiper  nests  on  the  higher  lichen-covered  parts  of  both 
St.  George  and  St.  Paul,  and  as  far  as  known,  on  only  one  other 
group — St.  Matthew.  The  first  eggs  known,  a  set  of  4,  were  taken 
by  H.  W.  Elliott  and  G.  R.  Adams  on  St.  George,  June  19,  1873. 
Another  set  of  4  was  taken  by  True  and  Prentiss  on  St.  Paul,  July 

6,  1895.     More  recently  a  number  of  sets  have  been  taken  on  St. 


68  FORTH   AMERICAN   FAUNA.  [No.  46 

George  Island  by  G.  Dallas  Hanna  and  others,  and  Mr.  Hanna 
(1921a,  pp.  50-57)  has  published  a  very  full  and  interesting  account 
of  the  nesting  of  the  species,  with  description  and  measurements  of 
the  eggs,  and  the  present  place  of  depository  of  the  23  sets  of  which 
he  has  knowledge. 

The  experience  of  the  writer  is  limited  to  a  few  observations  made 
on  the  two  main  islands  of  the  group  in  the  summer  of  1914.  A 
few  pairs  attending  young  were  found  on  the  breeding  grounds 
near  the  Polovina  Lakes,  St.  Paul,  on  June  28,  and  adults  and  downy 
young  were  collected.  Both  sexes  had  been  brooding  as  shown  by 
the  bare  spots  on  the  sides  of  the  abdomen.  They  were  still  linger- 
ing about  their  breeding  grounds  here  on  July  20,  and  perhaps  later. 
About  August  9  the  birds  began  to  be  common  about  the  beaches, 
the  flocks  there  apparently  being  in  excess  of  the  number  breeding 
on  the  islands,  and  in  all  probability,  therefore,  comprised  in  part 
of  migrants  from  other  breeding  stations.  They  continued  to  be 
abundant  until  my  departure  the  last  of  August.  A  young  female 
bird  taken  August  28  still  retained  some  of  the  down  of  the  juvenal 
plumage  on  the  postauricular  region.  During  our  brief  visits  in 
mid-July  and  early  August  to  St.  George  Island,  where  the  bird 
breeds  more  abundantly,  many  were  observed,  on  the  former  occa- 
sion near  their  breeding  grounds,  and  in  early  August  mainly  about 
the  beaches. 

About  the  middle  of  July,  when  the  earlier  nesting  birds  are  freed 
from  family  cares,  they  begin  to  resort  to  the  beaches  to  feed,  and 
at  night  gather  in  flocks  to  roost  on  some  favorite  rocky  point. 
Later  the  young  join  the  adults  and  the  flocks  increase  in  size  through 
August. 

The  majority  of  the  birds  leave  by  mid-September.  Hahn  in  1910 
observed  a  few  on  St.  Paul  during  September  and  early  October; 
his  latest  date  is  October  20.  Hanna  in  1913  noted  the  species  on 
St.  George  as  follows :  August  30,  20  seen ;  September  4,  20 ;  Sep- 
tember 11,  500;  September  15,  about  75;  September  28,  4.  Hanna's 
notes  for  St.  Paul,  autumn  of  1914,  are  as  follows:  September  14, 
large  flocks  going  south;  September  16,  about  500  individuals  seen, 
the  majority  have  gone;  September  20,  about  200;  September  22, 
about  30 ;  October  4,  20 ;  October  17,  10. 

The  species  winters  as  far  as  known  on  the  mainland  of  south- 
eastern Alaska  and  has  not  been  observed  on  the  Pribilofs  later  than 
the  dates  above  noted,  being  replaced  at  that  season,  in  a  measure, 
by  the  Aleutian  sandpiper,  as  detailed  in  the  account  of  that  species. 

Males  taken  by  Hahn  and  Marsh  measured  in  the  flesh :  Length,  8 
to  9|  inches;  extent,  16  to  16T3e;  females,  length,  9f  to  10£;  extent, 
16|  to  17£.    A  female  taken  on  St.  Paul,  October  19,  1910,  by  Hahn, 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  69 

had  the  bill  horn  color  at  base,  dusky  at  tip;  feet  dusky  greenish, 
tarsus  horn  color. 

Food. — More  stomachs  of  the  Pribilof  sandpiper  have  been  avail- 
able for  examination  than  of  any  other  species,  the  total  being  198, 
of  which  6  were  nearly  empty  (and  consequently  excluded  from  com- 
putations, the  results  of  which  are  here  cited).  The  articles  of  food 
composing  more  than  1  per  cent  of  the  total  were:  Mollusks,  32.63 
per  cent;  crustaceans,  29.15  per  cent;  flies  (Diptera),  23.49  per  cent; 
beetles,  10.29  per  cent;  marine  worms,  1.27  per  cent;  and  vegetable 
matter,  chiefly  algae.  1.21  per  cent. 

The  molluscan  food,  although  the  largest  single  item,  was  little 
varied,  consisting  chiefly  of  univalves,  and  these  mostly  of  the  genus 
Littorina.  Unidentified  species  of  this  genus  were  found  in  33 
stomachs,  and  L.  sitchana  in  53.  Several  stomachs  held  large  num- 
bers of  this  species,  the  maximum  count  being  205.  One  other 
univalve  (Neritina  sitchana)  was  determined  among  the  food,  and  3 
kinds  of  bivalves,  namely,  Pisidium  hannai  in  1  stomach,  blue  mussels 
(Mytilus  edulis)  in  14,  and  Modiolus  modiolus  in  7.  One  of  these 
stomachs  yielded  40  of  the  last-named  species. 

Like  the  molluscan  food  of  the  Pribilof  sandpiper,  the  crustacean 
also  was  very  uniform,  being  almost  entirely  amphipods.  Unidenti- 
fied amphipods  were  found  in  43  stomachs,  Gammarus  pribilof ensis 
in  20,  Gammarus  sp.  in  9,  and  Orchestia  sp.  in  1.  Isopods,  named  as 
Idothea  (?)  ochotensis,  occurred  in  2  gizzards,  and  eggs  of  water 
fleas  (Daphniidae)  in  1. 

The  third  item  in  rank  among  the  food  staples  of  this  sandpiper, 
two-winged  flies  (Diptera,  23.49  per  cent),  includes  larvae,  pupae, 
and  adults  of  a  variety  of  forms.  Crane-flies  (Tipulidae,  Tipula 
sp.),  either  the  larvae  or  adults,  the  latter  often  with  eggs,  were 
most  frequently  identified,  and  various  members  of  the  dung-fly 
family  (Scatophagidae)  next.  Flies  of  the  latter  group  determined 
were :  Scatophaga  crinita,  S.  furcata,  Leria  fratema,  and  L.  leucos- 
toma.  Blowflies,  so  numerous  on  the  Pribilofs  when  large  numbers 
of  rotting  seal  carcasses  are  present,  were  eaten  by  12  of  the  birds. 
As  many  as  23  of  one  of  these  flies  (Calliphora  vomitoria)  were 
found  in  a  single  stomach,  and  no  fewer  than  112  larvae  of  blowflies 
in  another.  Kelp-flies  {Coelopa  eximia,  Coelopa  sp.,  and  Fucellia 
sp.)  occurred  in  10  stomachs. 

Beetles,  last  of  the  major  elements  of  the  Pribilof  sandpipers' 
food,  10.29  per  cent  in  all,  were  a  varied  lot.  Ground  beetles  were 
distinctly  the  favorites,  those  of  the  genus  Pterostichus  being  found 
in  33  stomachs,  and  in  numbers  as  great  as  21,  25,  and  31  in  three 
instances.  Larvae  as  well  as  adult  beetles  of  this  family  were  eaten, 
and  the  following  forms  in  addition  to  the  genus  already  mentioned 


70  NORTH   AMERICAN   FAUNA.  [No.  46 

were  determined:  Amara  brunnip  emits,  Atnara  sp.,  and  Patrobus  sep- 
tentrionis.  A  small  species  of  the  burying  beetle  family  (Silphidae), 
by  name  Lyrosoma  opaca,  was  eaten  by  3  birds,  unidentified  rove 
beetles  (Staphylinidae)  by  7,  and  others  of  the  latter  family  as  fol- 
lows :  Atheta  sp.,  1 ;  Tachinus  apterus,  1 ;  Tachinus  sp.,  8 ;  and  Oloph- 
rum  fuscum,  2.  One  of  the  moss  beetles  {Byrrhus  fasciatus)  was 
found  in  3  stomachs,  a  click  beetle  {Hypnoidus  musculus)  in  2,  a  leaf 
beetle  {Chrysomela  subsulcata)  in  2,  the  beach  beetle  {Aegialites 
calif  ornicus)  in  3,  and  a  weevil  {Lophalophus  inquinatus)   in  3. 

Of  the  remaining  insects  in  the  dietary  of  this  bird,  caddis  larvae 
were  found  in  4  stomachs,  a  plant  bug  {Irbisia  sericans)  in  1,  a 
moth  in  1,  caterpillars  in  2,  and  Hymenoptera  in  several.  Identified 
forms  of  the  last  group  were  all  parasitic  wasps,  representing  the 
following  genera :  Gelis,  Seleucus,  Bathymetis,  Stiboscopus,  Phyga- 
deuon,  and  Ambly  teles. 

Mites  were  found  in  1  stomach  examined  and  spiders  in  5.  Marine 
worms,  chiefly  of  the  family  Nereidae,  composed  1.27  per  cent  of  the 
food,  and  as  many  as  38  individuals  were  represented  in  a  single 
stomach.  A  sculpin  {My otocephalus  sp.)  was  eaten  by  1  of  the 
birds  studied.  It  is  of  interest  to  note  that  material,  no  doubt  taken 
for  grinding  purposes,  included  in  a  few  cases  bits  of  bone,  and  in 
one  instance  a  molar  tooth  of  the  lemming  {Lemmus  nigripes). 

Vegetable  matter,  1.21  per  cent  of  the  total  food  of  the  species, 
consisted  chiefly  of  algae,  including  kelp  and  plume  algae  (Ptilota 
sp.).  Bits  of  moss  also  were  eaten  and  a  few  seeds  of  the  following 
plants:  Grass,  lupine  {Lupinus  nootkatensis) ,  violet  {Viola  langs- 
dorfti,  crowberry  {Empetrum  nigrum),  and  bottle  brush  {Hippuris 
vulgaris). 

Pisobia   acuminata    (Hoi*sfield).     Sharp-tailed    Sandpiper. 

The  sharp-tailed  sandpiper  was  first  recorded  from  the  Pribilofs 
by  Seale  (1898,  p.  139)  on  the  basis  of  a  specimen  taken  by  R.  E. 
Snodgrass  and  A.  W.  Greely  on  St.  Paul  Island  August  19,  1897. 
(Grinnell  (1901,  p.  20)  gives  the  date  of  what  is  apparently  the 
same  specimen  as  August  17.)  Bishop,  touching  at  St.  George  on 
October  3,  1899.  saw  about  a  dozen  and  took  3  (1900,  p.  66).  The 
bird  has  since  been  found  to  be  a  regular  fall  migrant  and  is  some- 
times abundant.  Hahn  recorded  it  in  small  numbers  on  September  7 
and  24,  October  11  and  20,  and  November  9,  1910.  A  male  taken 
by  Hahn  on  St.  Paul  on  October  8  is  in  the  collection  of  the  Bio- 
logical Survey. 

Hanna,  in  1913,  took  specimens  on  St.  George  on  August  30, 
September  5  and  6,  and  October  18.  In  1914,  on  St.  Paul,  he 
observed  a  few  on  August  29,  September  14  and  20,  and  1  on  Octo- 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  71 

ber  12.  The  same  observer  noted  the  species  in  large  flocks,  together 
with  the  pectoral  sandpiper,  on  the  seal-killing  fields  of  St.  Paul 
during  the  early  autumn  of  1919. 

Two  males  taken  by  Marsh  October  11,  1911,  measured,  respec- 
tively :  Length,  9£  and  8f  inches ;  extent,  17|  and  17.  Females  taken 
by  Hahn  measured  as  follows :  Length,  8  inches ;  extent,  14f  and  15£. 

Food. — Eight  well-filled  and  1  nearly  empty  stomach  of  the  sharp  - 
tailed  sandpiper  are  available  to  illustrate  the  food  habits.  This 
number  is  too  small  to  furnish  reliable  results,  and  too  great  depend- 
ence must  not  be  placed  in  data  as  to  the  relative  ranks  of  food  items 
as  here  stated.  The  percentages  found  for  the  limited  material, 
then,  are  flies  (Diptera),  39.1  per  cent;  crustaceans,  18.1  per  cent; 
mollusks,  14.2  per  cent;  caddisflies,  11.8  per  cent;  beetles,  8.8  per 
cent;  Hymenoptera,  1.8  per  cent;  and  vegetable  matter,  3.9  per  cent. 

Mr.  Hanna  notes  that  flocks  of  this  species  frequent  the  seal- 
killing  fields  feeding  on  fly  maggots,  a  statement  receiving  confirma- 
tion from  stomach  analysis.  Blowflies  {Calliphora  vomitoria)  were 
found  in  two  stomachs  and  larvae  of  this  or  an  allied  species  in 
another.  Thirty-three  rat-tailed  larvae  of  flower  flies  (Syrphidae; 
Eristalinae),  and  those  of  crane  flies  (Tipulidae),  and  midges  (Chi- 
ronomidae)  in  1  each. 

Both  caddis  larvae  and  adults  were  eaten,  and  of  the  latter  Chi- 
lostigma  praeteritum  was  identified.  Among  the  beetles,  ground 
beetles  are  well  represented  by  Pterostichus  in  2  stomachs  and  the 
large  gold  and  green  beetle  (Carabus  truncaticollis)  in  1.  Other 
species  of  beetles  eaten,  each  by  1  bird,  except  a  weevil  (Lophalo- 
phus  inquinatus)  bj'  2,  were:  A  predacious  diving  beetle  (Hydro- 
porus  sp.),  rove  beetle  (Tachinus  sp.),  moss  beetle  (Byrrhus 
fasciatus),  click  beetle  (Hypnoidus  musculus),  stores  beetle  (Ptinus 
fur),  and  a  leaf  beetle  (Chrysomela  subsulcata). 

The  few  remaining  insects  found  in  stomachs  of  the  sharp-tailed 
sandpiper  were  parasitic  wasps  {Gelis  sp.,  and  Polyrhembia  sp.) 
and  a  plant  bug  {Irbi&ia  sericans).  Spiders  were  found  in  1  stomach 
and  amphipocls  in  2,  these  comprising  the  bulk  of  the  crustaceans 
eaten.  The  vegetable  matter  in  the  stomachs  consisted  of  plant 
fibers,  further  unidentified,  which  were  probably  taken  accidentally. 

Pisobia  maculata  (Vieillot).     Pectoral  Sandpiper. 

The  pectoral  sandpiper  is  a  migrant  on  the  Pribilofs,  evidently 
occurring  mainly  in  early  autumn,  though  the  first  record  relates  to 
June  specimens.  Palmer  (1899,  p.  404)  obtained  2  specimens  on  St. 
Paul,  June  12,  1890.  The  next  instance  of  its  capture  seems  to  have 
been  recorded  by  Bishop  (1900,  p.  66),  who  mentions  a  specimen  shot 
on  St.  George  Island  by  W.  H.  Osgood,  October  3,  1899.    Hahn  took 


72  NORTH   AMERICAN   FAUNA.  [No.  46 

specimens  on  St.  Paul  on  September  11  and  October  8,  1910;  and 
Marsh  collected  the  species  on  the  same  island  on  October  11,  1911, 
and  May  30,  1912. 

While  on  St.  Paul  Island  in  the  summer  of  1914  the  writer  first 
noted  this  species  on  August  17,  when  four  were  observed  at  Northeast 
Point ;  subsequent  records,  made  near  the  village,  follow :  August  23, 
7  observed ;  August  24,  30 ;  August  25,  6 ;  August  28,  common ;  August 
29,  abundant.  Following  my  departure  on  August  30,  G.  Dallas 
Hanna  noted  it:  September  14,  18;  September  16,  12;  September  20, 
about  50.  / 

Notes  taken  from  the  field  catalogues  follow :  Male,  St.  Paul  Island, 
October  8,  1910,  W.  L.  Hahn:  length,  8f  inches;  extent,  16i ;  bill, 
horn  color  at  base,  dusky  on  terminal  half;  feet,  greenish-yellow. 
Female,  St.  Paul  Island,  September  11, 1910, Hahn:  length, 8|  inches; 
extent,  17.  Male,  St.  Paul,  May  30,  1912,  M.  C.  Marsh :  length,  9f 
inches;  extent,  17f ;  legs  and  feet,  yellowish-red;  bill,  black. 

Food. — Included  in  the  present  study  are  23  stomachs  of  this 
species,  of  which  2  collected  long  ago  were  imperfectly  examined  and 
are  not  available  for  reexamination.  The  food  proportions  here 
given,  therefore,  are  based  on  the  contents  of  21  well-filled  gizzards. 
The  principal  elements  of  the  food  are :  Flies  (Diptera),  54.5  per  cent; 
amphipods,  22.3  per  cent;  vegetable  matter,  chiefly  algae,  10.5  per 
cent;  beetles,  8  per  cent;  Hymenoptera,  2.1  per  cent;  and  bugs 
(Hemiptera),  1.3  per  cent. 

The  Diptera  eaten  were  almost  exclusively  larvae  of  crane-flies 
(Tipulidae;  Tipula  sp.),  of  which  no  fewer  than  123  were  in  a  single 
stomach  among  the  16  in  which  such  larvae  occurred.  Kelp-flies 
{Coelopa  sp.)  were  found  in  2  stomachs.  The  beetles  eaten  were 
mostly  ground  beetles  and  their  larvae,  of  which  the  following  kinds 
were  identified:  Amara  brunnipennis,  A.  glacialis,  Amara  sp.,  and 
Pterosticlius  sp.  Other  beetles  eaten  included  a  predacious  diving 
beetle  (Hydroporus  sp.),  rove  beetles  (Quedius  hyperboreus,  and 
Hadrotes  sp.),  and  weevils  (Lophalophus  inqulnatus).  The  Hy- 
menoptera consumed  were  small  parasitic  wasps  {Amblyteles  sp.  and 
Campoplex  sp.),  and  the  Hemiptera,  the  plant  bug  (Irbis'ia  sericans). 
The  only  other  insects  eaten  were  caddis  larvae.  Mites  and  spiders 
were  found  in  3  stomachs  each,  and  amphipods,  the  only  crustaceans 
eaten,  in  5. 

The  vegetable  food,  while  largely  algae,  included  also  a  few  seeds 
of  grass,  lupine  (Lupinus  nootkatensis) ,  and  violet  {Viola  langs- 
dorfii) . 

Pisobia  bairdi  (Coues).    Baird  Sandpiper. 

The  Baird  sandpiper  was  added  to  the  Pribilof  list  by  Hanna 
(1916,  p.  401),  who  took  3  specimens  at  Kaminista  Lake,  St.  Paul 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  73 

Island,  August  31,  1914.  Mailliard  and  Hanna  (1921,  p.  95)  have 
recently  put  on  record  2  specimens  taken  at  Northeast  Point,  St. 
Paul  Island,  August  12,  1920. 

Food.— The  food  secured  by  Baird  sandpipers  on  the  Pribilofs  is 
known  only  from  the  contents  of  three  stomachs  taken  August  31, 
1914.  They  contained  amphipods  (partly  Gamvmrus  sp.),  73.6  per 
cent;  algae,  11.6  per  cent;  beetles,  10.3  per  cent;  and  a  fly  larva,  1 
per  'cent.  The  beetles  were  ground  beetles  (Pterostiohus  sp.  and 
Patrobus  septentrionis) ,  and  a  weevil  (Lophalophus  inquinatus). 

Pisobia  rainutilla   (Vieillot).    Least  Sandpiper. 

The  first  specimen  of  the  least  sandpiper  known  to  have  been  taken 
on  the  Pribilofs  was  a  female  shot  by  Wilfred  H.  Osgood,  at  Kamin- 
ista  Lake,  St.  Paul  Island,  August  29,  1914.  It  is  now  in  the  col- 
lection of  the  Biological  Survey,  and  has  been  recorded  by  G.  Dallas 
Hanna  (1916,  p.  401).  Hanna  also  observed  2  on  September  14  of 
the  same  year,  and  1  on  July  13,  1915,  also  on  St.  Paul.  A  second 
specimen,  taken  by  Hanna  at  Northeast  Point  on  St.  Paul  Island, 
August  27,  1920,  has  been  recorded  by  Mailliard  and  Hanna  (1921, 

p.  95). 

Food.— The  stomachs  of  the  2  specimens  above  mentioned  were 
examined.  One  of  them  contained  amphipods  exclusively,  and  the 
other  the  following  items:  23  seeds  of  bottle  brush  (Hippuris  vul- 
garis), 50  per  cent;  bits  of  hydroid  stems,  40  per  cent;  and  chitin 
from  the  blue  mussel  {Mytilus  edulis),  10  per  cent. 
Pisobia  subminuta  ( Middendorff  1 .10     Long-toed  Stint. 

The  specimen  of  the  long-toed  stint  taken  by  Townsend  in  1885, 
which  added  a  new  species  "to  the  North  American  list,  still  remains 
the  only  record  for  the  Pribilofs.  The  occurrence  was  first  put  on 
record  by  Ridgway  (1886,  p.  275).  Concerning  the  taking  of  the 
bird,  Townsend  says  (1887,  p.  100)  : 

By  the  capture  of  this  Asiatic  bird  on  Otter  Island,  Alaska,  where  I  shot  an 
adult  female,  in  breeding  plumage,  on  June  8,  1885,  a  species  is  added  to  the 
fauna  of  North  America.  It  was  feeding  in  a  shallow  salt-water  pond,  with 
other  Tringae,  which  I  supposed  to  be  Actodromas. 

The  long-toed  stint  breeds  in  Kamchatka  and  other  parts  of  eastern 
Siberia,  including  the  Commander  and  Kurile  Islands.     In  winter 
it  occurs  as  far  south  as  Australia  and  the  Philippines. 
Pelidna    alpina    sakhalina    (Vieillot).     Red-backed    Sandpiper. 

The  American  dunlin  was  first  observed  on  the  Pribilofs  by  L.  B. 
Bishop  October  3,  1899,  when  during  a  short  visit  to  St.  George 
Island  be  observed  a  few  (1900,  p.  67).    The  first  specimen,  a  male, 

10  Pisobia  damacensis  of  the  A.  O.  U.  Check  List,  1910. 


74  NORTH   AMEPJCAN   FAUNA.  [No.  46 

was  taken  by  Hahn  October  30,  1910,  and  recorded  by  Evermann 
(1913,  p.  17).  It  measured:  Length,  7-J  inches;  extent,  14.  It  is  now 
in  the  collection  of  the  Biological  Survey. 

A  second  specimen,  shot  by  Doctor  Hunter  on  St.  Paul  Island, 
September  3,  1914,  was  preserved  by  Hanna,  and  is  now  in  the  U.  S. 
National  Museum  collection.  The  reddish  brown  of  the  scapulars 
and  tertials  is  nearly  replaced  by  the  gray  of  the  winter  plumage; 
sides  of  breast  with  small  rounded  black  blotches.  Another  indi- 
vidual was  observed  at  the  same  time. 

Food. — The  single  available  stomach  0f  the  red-backed  sandpiper 
(St.  Paul,  September  3,  1914)  was  entirely  filled  with  amphipods. 

Ereunetes  pusillus  (Linnaeus).     Semipalmated  Sandpiper. 

A  female  semipalmated  sandpiper  taken  by  Palmer  on  St.  Paul 
Island,  June  12,  1890,  remains  the  only  record  for  the  Pribilofs.  This 
specimen,  which  was  recorded  by  Palmer  (1899,  p.  405),  was  in  com- 
pany with  two  pectoral  sandpipers,  evidently  just  arrived  from  the 
south. 

Food. — The  stomach  of  this  specimen  was  found  to  contain  remains 
of  the  beach  beetle  (AegiaZites  calif 'ornicus) ,  10 per  cent;  fragments  of 
small  flies  (Diptera),  85  per  cent;  and  two  seeds  (not  identified),  5 
per  cent. 
Limosa  lapponica  baueri  (Naumann).     Pacific  Godwit. 

The  Pacific  godwit  seems  to  be  of  regular  occurrence  in  spring 
and  fall.  It  was  first  recorded  by  Elliott  who  observed  it  in  flocks 
of  a  dozen  to  50  (1874  unpaged,  and  1875,  p.  187).  Subsequent 
records  of  occurrence  are  few.  Palmer  took  specimens  on  Walrus 
Island,  June  13,  and  on  St-  Paul,  July  7,  1890.  Marsh  collected  a 
male  on  St.  Paul,  June  12,  1912;  it  measured:  Length,  15^  inches; 
extent,  27|.  Hanna  collected  an  adult  female  on  St.  George,  Sep- 
tember 7,  1913.  In  1915,  on  St.  Paul,  he  observed  the  bird  as  follows: 
May  20,  4  seen,  1  taken ;  May  26,  14 ;  May  27,  30 ;  May  28,  50  seen 
at  Northeast  Point. 

Food. — Six  stomachs  of  Pacific  godwits  collected  on  St.  Paul  and 
Walrus  Islands  by  William  Palmer  in  1890  have  been  reexamined. 
Dr.  S.  D.  Judd's  analysis  of  their  contents  was  published  by  Palmer 
(1899,  pp.  405,  406),  but  the  present  report  is  an  improvement  in 
some  respects  over  the  original.  The  items  of  the  food  in  rank  by 
bulk,  are  flies  (Diptera),  76.6  per  cent;  beetles,  17  per  cent;  mollusks, 
3.6  per  cent;  marine  worms  (Nereidae),  1.3  per  cent;  and  vegetable 
matter,  1  per  cent.  The  fly  remains  were  very  largely  midge 
(chironomid)  larvae,  but  included  also  a  few  dung-flies  (Scatophaga 
sp.).  The  beetles  included  as  the  principal  item  450  or  more  beach 
beetles  (Aegialites  calif ornicus) ,  which  made  up  85  per  cent  of  the 
contents    of    one    stomach ;    and    as    lesser    items,    ground    beetles 


1923.]  BIRDS  OF   THE   PRIBILOF   ISLANDS.  75 

{Pterostichus  sp.  and  Amara  bru/nnipennis) ,  rove  beetle  (Tachivus 
sp.),  and  a  weevil  (Lophalophus  inguinatios) .  A  caddisfly  was  the 
only  other  insect  eaten.  The  mention  of  tiger  beetles  by  Judd  must 
be  set  down  as  a  misidentification,  based  no  doubt  on  the  jaws  of 
marine  worms  (Nereidae)  which  one  of  the  stomachs  contained.  All 
of  the  mollusks  eaten  by  the  godwits  were  of  a  single  species  of 
univalve,  Littorina  sitchana. 

The  vegetable  matter  consisted  of  unidentified  fibers  and  a  few 
leguminous  seeds,  probably  accidentally  taken. 

Totanus  melanoleucus  (Gmelin).     Greater  Yellow-legs. 

A  greater  yellow-legs  taken  at  Kaminista  Lake,  St.  Paul  Island,  in 
1897,  by  R.  E.  Snodgrass  and  A.  W.  Greely,  was  at  last  accounts  in 
the  collection  of  Leland  Stanford  University.  Alvin  Seale,  who 
first  recorded  it  (1898,  p.  139).  gives  the  date  as  August  23.  Grin- 
nell,  reporting  on  the  same  collection  (1901,  p.  20),  gives  the  date  as 
August  17.  However,  Walter  K.  Fisher,  in  response  to  my  request, 
kindly  looked  up  the  specimen  and  tells  me  that  the  label  bears  the 
date  August  29,  1897.  These  details  are  given  lest  it  be  supposed 
that  there  are  several  specimens;  there  is  apparently  only  one  in- 
volved. 

Hanna  reports  seeing  one  at  close  range  on  St.  George  on  May  28, 
1917,  and  another  on  St.  Paul,  June  10,  1919. 

Totanus  flavipes  (Gmelin).     Lesser  Yellow-legs. 

Palmer  saw  a  lesser  yellow-legs  among  a  flock  of  godwits,  June 
11,  1890,  but  failed  to  secure  it  (1899,  p.  407).  Hanna  took  a  female 
specimen,  now  in  the  collection  of  the  U.  S.  National  Museum,  on  St. 
George  Island,  October  18,  1913,  and  has  recorded  it  (1916,  p.  402). 
These  seem  to  be  the  only  records. 

Rhyacophilus  glareola  (Linnaeus).     Wood  Sandpiper. 

A  specimen  of  the  wood  sandpiper  was  taken  on  St.  George  Island 
May  19,  1914,  by  G.  Dallas  Hanna.  This  specimen,  which  was  par- 
tially demolished  by  a  blue  fox  and  now  consists  only  of  the  wings 
and  one  foot,  is  now  in  the  collection  of  the  Biological  Survey,  and 
has  been  recorded  by  Mr.  Hanna  (1916,  p.  401). 

The  wood  sandpiper  breeds  in  northern  Eurasia  east  to  Kam- 
chatka and  Bering  Island,  and  winters  in  northern  Africa,  south- 
ern Asia,  Japan,  and  the  Philippines.  It  had  been  taken  once  in 
Alaska  previous  to  its  capture  on  the  Pribilofs. 

Heteroscelus  incanus  incanus   (Gmelin)."     Wandering  Tattler. 

The  wandering  tattler  is  a  rather  common  migrant  in  early  and 
late  summer,  the  outward  spring  and  return  fall  movements  being 

11  Heteractitis  incanus  of  the  A.  O.  U.  Check  List,  1910. 


76  NORTH   AMERICAN   FAUNA.  [No.  46 

separated  by  only  a  short  interval.  The  species  was  first  recorded 
from  the  islands  .by  Elliott  (1874,  unpaged,  and  1875,  p.  187).  The 
available  dates  of  occurrence,  divided  as  nearly  as  may  be  into  spring 
and  fall  records,  are  as  follows:  St.  Paul  Island,  May  27,  1872, 
specimen  (Elliott).  Otter  Island,  June  8,  1885,  specimen  (Town- 
send).  St.  Paul  Island,  May  22,  1911,  1  seen;  May  29,  1  seen;  May 
30,  2  seen  (Hahn).  St.  Paul  Island,  May  31,  1915,  1  seen  (Hanna). 
The  earliest  record  marking  the  return  movement  is  that  of  Hanna, 
who  saw  one  on  St.  Paul  Island,  July  7,  1915.  Palmer  observed 
it  on  St.  Paul  Jul}7  10,  1890,  and  took  specimens  there  on  July  29 
and  31.  Hahn  did  not  begin  his  observations  on  St.  Paul  in  1910 
until  about  the  1st  of  September ;  his  notes  on  this  species  are  as  fol- 
lows :  September  2,  2 ;  September  4,  2 ;  September  7,  3 ;  September  8, 
6;  September  9,  3;  September  11,  10;  September  18,  3;  October  4,  1. 
Hanna,  on  St.  George,  observed  4  singly  on  August  24,  1913,  and  col- 
lected 1. 

During  the  late  summer  of  1914  the  writer  did  not  observe  this 
species  until  July  27,  when  one  was  seen  on  Otter  Island.  On  St. 
Paul  a  few  were  seen  between  Northeast  Point  and  the  village  on 
July  29,  and  subsequently  as  follows :  August  15,  1 ;  August  16,  1 ; 
August  23,  2;  August  25,  1;  August  28,  4;  August  29,  3.  One  was 
also  observed  on  St.  George  on  August  5.  Following  my  departure 
from  the  Pribilofs  G.  Dallas  Hanna  recorded  the  species  on  St. 
Paul  on  the  following  dates:  September  14,  8;  September  16,  10; 
September  20,  3 ;  September  22,  2.  Hanna  informs  me  he  took 
specimens  on  St.  Paul  Island  on  August  18  and  27,  1920. 

Food. — The  six  stomachs  of  wandering  tattlers  examined  yielded 
quite  a  variety  of  food  items  of  which  those  in  excess  of  1  per  cent 
of  the  total,  are:  Flies  (Diptera),  46.1  per  cent;  caddisflies,  30.6 
per  cent ;  amphipods,  16  per  cent ;  mollusks,  3.6  per  cent ;  and  beetles, 
1.1  per  cent.  One  stomach  was  filled  with  blowflies  (Callipfiora 
vomitoria),  another  nearly  so  with  dung-flies  (Scatophaga  crinita), 
and  a  third  in  like  measure  with  kelp-flies  (Coelopa  frigida),  and 
crane-fly  larvae.  Two  of  the  stomachs  were  nearly  full  of  caddisfly 
larvae.  Amphipods  (including  Gammawis  sp.)  occurred  in  three  of 
the  gizzards  but  in  quantit}'  only  in  one.  The  mollusks  eaten  by  two 
birds  were  univalves  of  the  genus  Littorina,  23  in  one  of  the  stom- 
achs being  L.  sitchana.  The  only  beetle  was  the  large  gold  and 
green  ground  beetle  (Carabvs  truncaticollis)  and  the  only  other  in- 
sects were  small  parasitic  wasps  {P olyrhembia  sp.).  Concluding  the 
enumeration  of  food  items,  one  stomach  contained  a  few  bones  of  a 
small  fish  and  another  a  little  vegetable  debris. 
Heteroscelus  incanus  brevipes  (Vieillot).     Polynesian  Tattler. 

The  Polynesian  tattler  has  been  taken  on  three  occasions  on  the 
Pribilofs,   the   only    North   American   locality.     G.    Dallas   Hanna 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  77 

first  recorded  it  (1919a,  p.  176,  and  1920a,  p.  250)  and  (1.  c,  p.  250) 
elaborates  the  circumstances  as  follows : 

The  history  of  the  Polynesian  tattler  in  North  America  dates  back  to 
October  4,  1911,  when  a  female  was  secured  on  St.  Paul  Island  by  Mr.  M.  C. 
Marsh,  then  the  naturalist  of  the  fur-seal  service.  The  specimen  was  placed 
in  the  National  Museum  collection  without  being  detected  as  differing  from  the 
wandering  tattler.  It  was  discovered  by  Dr.  H.  C.  Oberholser  while  he  was 
verifying  the  identification  of  a  second  specimen  of  the  same  species,  a  female 
collected  on  St.  Paul  Island,  September  2,  1917,  by  the  writer.  Owing  to  the 
difficulty  of  distinguishing  the  tattlers  it  may  be  that  the  Asiatic  form  comes 
across  Bering  Sea  more  frequently  than  the  records  would  indicate. 

A  third  specimen  was  taken  later  by  Mr.  Hanna,  and  is  referred 
to  as  follows  (1920b,  p.  174)  : 

A  female  Polynesian  tattler  (Heteractitis  brevipes)  was  secured  on  St. 
Paul  Island  on  September  17,  1919,  near  Kitovi  Rookery.  It  makes  the  thii'd 
specimen  taken  in  North  America.  I  had  an  opportunity  to  observe  this  bird 
for  a  while  with  two  wandering  tattlers  in  view  at  the  time.  The  actions  of 
the  two  species  were  practically  the  same.  They  feed  very  close  to  the  sea 
on  rocky  shores  and  when  disturbed  fly  lazily  rarely  more  than  100  yards. 
The  wandering  tattlers  on  this  occasion  appeared  perceptibly  larger  than  the 
Polynesian,  and  the  notes  of  the  two  were  different.  The  latter  uttered  an 
irregular  screech  not  of  the  same  intensity  or  pitch,  whereas  the  former  gave 
its  usual  call,  a  series  of  6  to  10  individual  notes  uttered  in  the  same  pitch  and 
rapid  succession  but  each  of  shortening  duration. 

The  Polynesian  tattler  breeds  in  eastern  Siberia,  including  Kam- 
chatka, and  moves  southward  in  winter  to  China,  Japan,  the  Malay 
Peninsula,  and  Australia.  It  has  also  been  taken  on  Bering  Island 
in  the  spring  migration. 

Philomachus  pugnax  (Linnaeus).12    Ruff. 

An  immature  female  ruff  taken  by  W.  L.  Hahn  on  St.  Paul  Island, 
September  7,  1910,  formed  the  basis  for  Evermann's  account,  credit- 
ing the  species  to  the  Pribilofs  (1913,  p.  18).  This  specimen,  which 
is  now  No.  239169  of  the  collection  of  the  Biological  Survey,  is  the 
first  one  taken  on  the  western  coast  of  North  America.  Hahn's 
field  catalogue  shows  that  it  measured  in  the  flesh :  Length,  llf  inches ; 
extent,  23J. 

The  ruff  is  of  wide  distribution  in  central  Eurasia,  occurring 
eastward  rarely  to  Japan  and  China,  and  in  winter  moving  south- 
ward to  Africa  and  India. 

Numenius  hudsonicus  Latham.     Hudsonian  Curlew. 

The  Hudsonian  curlew  has  been  recorded  a  few  times  from  the 
islands  on  what  appears  to  be  excellent  authority,  but  no  specimens 
have  been  taken.  It  was  first  added  to  the  list  by  Palmer,  who  re- 
corded one  seen  at  close  range  on  St.  Paul  Island  by  F.  A.  Lucas, 

12  Machetes  pugnax  of  the  A.  O.  U.  Check  List,  1910. 
108731  °— 23 6 


78  NORTH   AMERICAN   FAUNA.  [No.  46 

July  16,  1897.  The  St.  Paul  Island  log,  under  date  of  May  27,  1903, 
refers  to  "  curlew  "  as  having  been  "  here  last  few  clays  "  (notes  taken 
by  Hahn).  Hahn  himself  noted  the  bird  as  follows :  August  29,  1910, 
1  seen;  September  2,  1  seen  flying  over  pond;  September  25,  1  seen 
flying  over  lagoon. 

While  on  St.  Paul  Island  in  1914  the  writer  saw  2  curlews,  which 
seemed  to  be  of  this  species,  flying  over  the  shallow  ponds  near  the 
village  of  St.  Paul.  On  May  18,  1915,  G.  Dallas  Hanna  observed 
6  individuals  on  St.  Paul. 

Numenius  borealis  (J.  R.  Forster).     Eskimo  Curlew. 

The  Eskimo  curlew  was  first  recorded  by  Coues  (in  Elliott,  1874, 
unpaged;  1875,  p.  189).    He  says: 

A  single  specimen  only  of  the  Esquimaux  curlew  was  taken  by  Mr.  Elliott  on 
St.  Paul's  Island,  June,  1872.     None  other  than  this  one  was  seen  by  him. 

Palmer  lists  the  specimen  as  No.  62448,  adult  male,  May  26,  1872. 
This  specimen  is  still,  and  will  probably  remain,  the  only  record  for 
the  Pribilofs. 

Numenius  tahitiensis  (Gmelin).     Bristle-thighed  Curlew. 

The  bristle-thighed  curlew,  which  had  previously  been  taken  on 
the  mainland  of  northern  Alaska,  was  added  to  the  Pribilof  list  by 
G.  Dallas  Hanna,  a  specimen  having  been  taken  by  him  on  St.  George 
Island,  May  26, 1917,  and  recorded  by  the  collector  (1919a,  p.  176,  and 
1920a,  p.  252).  This  specimen  is  now  No.  255154  of  the  bird  col- 
lection of  the  U.  S.  National  Museum.  The  species  winters  on 
Hawaii  and  other  islands  of  the  south  Pacific. 

Family  CHARADRIIDAE. 
Pluvialis  dominica  fulva  (Gmelin).1'    Pacific  Golden  Plover. 

The  western  race  of  the  golden  plover  visits  the  Pribilofs  regu- 
larly in  spring  and  fall,  still  in  fair  numbers,  although  it  is  prob- 
ably much  less  common  than  formerly.  It  was  first  recorded  from 
the  group  by  Coinde  (1860,  p.  400),  who  considered  it  identical  with 
European  specimens.  Elliott  (1874,  unpaged,  and  1875,  p.  180) 
observed  it  as  a  spring  and  fall  migrant,  and  the  single  specimen 
preserved  by  him  was  identified  by  Coues  as  Charadrius  fulvus. 

Coinde's  specimens  taken  by  Warneck  seem  to  have  been  unusually 
early  visitors;  they  were  taken  on  St.  Paul  April  18  and  20,  1852. 
Elliott's  specimen  was  collected  on  St.  Paul  May  1,  1873.  Palmer 
(1899,  p.  408)  took  a  single  specimen,  the  only  one  he  saw,  on  St. 
Paul  June  12,  1890.  Hahn  took  1  on  May  7,  1911.  Hanna's  earliest 
date  for  1915  is  May  9,  when  the  natives  reported  seeing  12  at  Half- 
way Point,  St.  Paul ;  he  noted  8  on  May  17,  and  1  on  May  28.  These 
are  all  the  spring  dates  available  to  the  writer. 

11  Charadrtus  dominions  fulvus  of  the  A.  O.  U.  Check  List,  1910. 


1923-1  BIRDS   OF    THE   PRIBILOF   ISLANDS.  79 

Dates  of  arrival  in  the  autumn  are  more  common.  From  the  St. 
Paul  Island  log  Hahn  extracted  the  following  notes:  October  10, 
1897,  several  killed;  October  19,  1901,  many  came.  Hahn  himself, 
in  1910,  observed  the  species  on  St.  Paul  as  follows:  September  3, 
6;  September  4,  1;  September  7,  3;  September  11,  5;  October  3,  1; 
October  11,  4;  October  18,  2;  October  19,  6;  October  20,  6;  October 
23,  2;  October  24,  2.  Hanna,  on  St.  George,  noted  the  species  in  the 
autumn  of  1913  as  follows:  September  4,  2  seen;  September  7,  15 
seen  on  high  tundra,  1  taken ;  September  28,  4 ;  October  9,  4 ;  October 
18,  8  on  uplands.  Pie  also  noted  4  on  St.  George  on  November  5, 
1916. 

My  own  dates  for  St.  Paul  Island  in  1914  seem  to  be  the  earliest 
fall  records  available :  August  7,  1  observed ;  August  10,  small  flock ; 
August  15,  3;  August  17,  4  at  Northeast  Point;  August  24,  2.  Con- 
tinuing observations  after  my  departure,  Hanna  noted  the  bird  as 
follows:  September  3,  4  seen,  1  with  black  belly;  September  14,  6; 
September  20,  30;  October  4,  3;  October  25,  2  at  Northeast  Point. 

Food. — Elliott  and  Hanna  both  note  that  Pacific  golden  plovers 
frequent  the  seal-killing  grounds  on  the  Pribilofs  and  feed  ex- 
tensively upon  the  blowfly  maggots  there.  However,  in  the  two 
stomachs  available  for  examination  (St.  Paul,  June  12,  1890,  Palmer; 
and  September  3,  1914,  Hanna)  none  of  these  larvae  were  found, 
their  food  contents  consisting  of  beetles,  72.5  per  cent ;  flies,  22.5  per 
cent;  Hymenoptera,  4  per  cent;  and  seeds  of  crowberry  {Empetrum 
nigrum),  1  per  cent.  The  beetles  were  chiefly  ground  beetles  (Ptero- 
stichus  sp.,  Amara  sp.,  and  Carabus  truncaticollis) ,  but  rove  beetles 
(Staphylinidae)  and  leaf  beetles  (Chr-ysomela  sub  sulcata)  also  were 
eaten.  The  flies  devoured  were  chiefly  crane-fly  larvae  (Tipulidae), 
and  the  Hymenoptera  were  partly  a  new  species  of  parasitic  wasp 
(Ambly teles  alpestriformis)  and  partly  unidentified.  Plahn  noted 
several  chrysomelid  beetles  and  some  dry  grass  blades  in  a  stomach 
examined  by  him. 

Charadrius  semipalmatus  Bonaparte.     Semipalmated  Plover. 

The  semipalmated  plover  is  known  only  from  a  specimen  taken 
from  a  small  flock  at  Northeast  Point,  St.  Paul  Island,  July  6,  1895, 
by  D.  W.  Prentiss,  jr.,  and  first  published  by  Palmer  (1889,  p.  408). 
Prentiss  later  (1902,  p.  102)  gives  the  following  details  regarding 
its  capture : 

Saw  a  number  on  the  sands  at  Northeast  Point,  on  July  6,  and  as  I  only  had 
my  beansbooter,  I  failed  to  get  more  than  one,  which  was  a  poor  specimen. 
This  is  an  addition  to  the  avifauna  of  the  Islands. 

The  specimen  is  now  No.  153543,  adult  female,  of  the  bird  collec- 
tion of  the  U.  S.  National  Museum. 


80  NORTH   AMERICAN   FAUNA.  [No.  46 

Arenaria  interpres  interpres  (Linnaeus).     Pacific  Turnstone. 

The  Pacific  race  of  the  turnstone,  first  recorded  from  the  Pribilofs 
by  Coinde  (1860,  p.  400),  occurs  only  as  a  migrant,  mainly  in  fall, 
when  it  stops  for  rest  and  food  before  starting  on  its  long  journey 
to  its  wintering  grounds. 

I  have  only  two  dates  of  occurrence  which  may  be  classed  as  spring 
migration.  Hahn  took  a  specimen  May  24,  1911.  It  measured  in 
the  flesh:  Length,  9£  inches;  extent,  19£.  The  other,  taken  by  M.  C. 
Marsh  May  30,  1912,  measured :  Length,  9^  inches ;  extent,  18-^ ;  the 
legs  and  feet  were  orange  red.  These  specimens  are  in  the  collection 
of  the  Biological  Survey. 

The  first  fall  arrivals  come  some  time  in  July ;  the  earliest  date  I 
find  is  July  2,  1895  (Prentiss,  1902,  p.  102).  Hahn,  reaching  St. 
Paul  in  late  August,  1910,  found  the  bird  abundant.  His  notes  for 
the  remainder  of  the  autumn  follow :  September  1  to  15,  still  com- 
mon, flock  seen  leaving  September  10 ;  September  15  to  30,  less  com- 
mon, flock  seen  on  September  24 ;  October  2,  some  on  Walrus  Island ; 
October  3,  7 ;  October  4,  9 ;  October  5,  3 ;  October  6,  2 ;  October  18,  5 ; 
October  19,  1;  October  20,  2  (single  ones).  Hanna,  making  observa- 
tions on  St.  George  in  1913,  noted  the  species  as  follows :  August  30, 
4  seen;  September  4,  2;  September  5,  4;  September  15,  15. 

My  notes  for  St.  Paul  in  1914,  are  the  most  complete  available; 
they  follow:  July  11,  first  observed,  abundant,  and  remained  so  until 
the  17th ;  common  July  18,  19,  20,  21,  24,  25 ;  abundant  July  28  to  30 ; 
common  August  3,  4,  7 ;  abundant  August  9,  and  from  August  21  to  30. 
Continuing  observations  after  my  departure,  Hanna  noted  the  bird 
as  follows:  September  14,  large  flocks;  September  16,  about  500  ob- 
served; September  20,  about  50;  September  22,  5;  October  12,  2; 
October  4,  6 ;  October  17,  8 ;  October  25,  3  at  Northeast  Point ;  October 
24,  6  (all  with  sore  feet). 

Food. — As  in  the  case  of  the  Pacific  golden  plover,  Elliott  and 
Hanna  both  assert  that  turnstones  feed  almost  exclusively  on  mag- 
gots in  the  seal-killing  fields  during  their  stay  on  the  Pribilof 
Islands.14  The  12  well-filled  stomachs  of  the  bird  from  the  islands 
do  not  bear  out  such  far-reaching  statements,  but  these  larvae  did 
compose  most  of  the  contents  of  one  stomach.  The  food  items  of  the 
12  birds  collected  were  found  to  be  two- winged  flies  (Diptera), 
50.83  per  cent;  caddis  larvae,  16.25  per  cent;  beetles,  12.5  per  cent; 
amphipods,  11.66  per  cent:  mollusks,  8.33  per  cent;  and  seeds  of 
crowberry  (Empetmm  nigrum),  a  few  in  one  stomach,  amounting  to 
less  than  1  per  cent  of  the  total  food.     Besides  the  blowfly  larvae 


14  Mr.  Hanna  attributes   (1921,  p.  57)   the  same  habit  to  phalaropes,  and  pectoral  and 
sharp-tailed  sandpipers  also. 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  81 

previously  mentioned,  the  dipterous  food  included  larvae  of  crane 
flies  (numbering  82  and  110  in  two  instances),  remains  of  dung 
flies  (Scatophagidae),  and  other  fly  larvae  and  pupae.  The  beetles 
consumed  were  chiefly  ground  beetles,  those  identified  being  Pteros- 
tichus  sp.,  Amara  brunnipennis,  Amara  sp.,  and  Carabus  truncati- 
collis.  A  rove  beetle  (Tachinus  sp.)  and  a  click  beetle  also  were 
found  in  the  stomachs.  The  remains  of  mollusks  in  the  diet  were 
especially  interesting,  consisting  of  more  than  400  operculi  of  uni- 
valves, apparently  of  a  single  species,  which  completely  filled  one 
gizzard.  These  objects,  each  of  which  forms  a  door  closely  fitting 
the  aperture  of  the  shell  when  the  snail  withdraws  into  its  retreat, 
were  chitinous  in  the  present  instance,  and  evidently  had  strongly 
resisted  digestion;  it  is  probable  they  would  have  been  ejected  in 
mass  had  the  bird  lived.  Alvin  G.  Whitney  in  a  field  note  states 
that  turnstones  searched  over  the  whole  mud  bottom  of  a  large  pond 
for  the  ruby-red  midge  (chironomid)  larvae,  which  were  abundant 
there. 

Family  HAEMATOPODIDAE. 

Haematopus  bachmani  Audubon.     Black  Oyster-catcher. 

A  specimen  of  the  black  oyster-catcher  was  taken  on  St.  George 
Island  January  12,  1917,  and  was  preserved  and  recorded  by  G. 
Dallas  Hanna  (1919a,  p.  176,  and  1920a,  p.  253).  It  constitutes  the 
first  record  for  the  Pribilofs  and  is  now  No.  255153  of  the  bird  col- 
lection  of   the   Biological    Survey.     Hanna   comments    as   follows: 

There  is  good  reason  to  suspect  that  this  species  has  been  shot  on  St. 
George  Island  before.  One  native  told  me  he  had  given  a  bird  like  it  to  a 
Doctor  Mills  several  years  earlier  but  that  it  had  spoiled  before  being  pre- 
pared as  a  specimen  (1.  c,  p.  253). 

However,  a  specimen  (No.  241645)  taken  by  H.  C.  Mills  on  St. 
George  Island,  in  the  spring  of  1913,  is  in  the  Biological  Survey 
collection. 

Family  ACCIPITRIDAE. 

Archibuteo    lagopus    sancti-johannis     (Gmelin).      American     Rough-legged 
Hawk. 

The  American  race  of  the  rough-legged  hawk  was  added  to  the 
Pribilof  list  by  G.  Dallas  Hanna  (1919a,  p.  176,  and  1920a,  p.  253). 
His  account  (1.  c,  p.  253),  in  part,  follows: 

One  specimen,  unsexed,  was  secured  on  St.  George  Island  in  the  fall  of  1917 
by  Mr.  C.  E.  Crompton,  of  the  U.  S.  Bureau  of  Fisheries.  He  has  kindly  con- 
sented to  the  record  being  included  in  this  list. 

This  specimen  is  now  No.  255159  of  the  bird  collection  of  the  U.  S. 
National  Museum. 


82  NORTH   AMERICAN   FAUNA.  [No.  46 

Thallasoaetus  pelagicus  (Pallas).     Kamchatkan  Sea  Eagle. 

G.  Dallas  Hanna  has  put  on  record  the  capture  of  a  Kamchatkan 
sea  eagle  on  St.  Paul,  thereby  adding  it  to  the  Pribilof  and  to  the 
North  American  list  (1919a,  p.  176,  and  1920a,  p.  250).  His  account 
(1.  c,  p.  250),  in  part,  follows: 

A  bird  of  this  species  was  shot  and  wounded  on  St.  Paul  Island,  December 
15,.  1917,  but  fell  into  the  sea.  Five  days  later  it  was  picked  up  on  the  beach 
in  badly  decomposed  condition.  Enough  of  the  specimen  could  be  saved,  how- 
ever, to  enable  the  identification  to  be  made  in  the  National  Museum. 

The  incomplete  skeleton  is  in /the  collection  of  the  above  institu- 
tion. The  species  breeds  in  Kamchatka,  and  has  been  taken  on 
Bering  Island. 

Haliaeetus  leucocephalus  alascanus  C.  H.  Townsend.     Northern  Bald  Eagle. 

The  status  of  the  bald  eagle  in  the  Pribilof  avifauna  is  somewhat 
doubtful,  since  no  specimens  have  been  taken.  Palmer  first  recorded 
it,  having  exhibited  an  Unalaska  specimen,  which  was  recognized 
by  the  St.  Paul  natives  as  being  occasionally  seen  about  the  islands. 

According  to  the  St.  Paul  Island  log  (notes  extracted  by  Hahn) 
eagles  were  seen  on  April  29,  1907,  and  October  16,  1908.  A  large 
bird,  evidently  an  eagle,  was  reported  to  Hanna  as  having  been  seen 
on  St.  George,  December  21,  1913.  These  notes  probably,  though  of 
course  not  certainly,  refer  to  the  common  Alaska  species. 

Family  FALCONIDAE. 
Hierofalco  rusticolus  sacer  (J.  R.  Forster).     American  Gyrfalcon. 

The  gyrfalcon  is  a  more  or  less  regular  visitor  to  the  Pribilofs.  It 
was  first  credited  to  the  group  by  Coues  (in  Elliott,  1874,  unpaged, 
and  1875,  p.  179)  from  a  specimen  in  the  collection  from  St.  Paul 
Island,  taken  in  March,  1873.  The  same  specimen  was  referred  to  by 
Elliott  in  his  subsequent  accounts  of  the  ornithology  of  the  group. 
On  September  24,  1880,  a  female  specimen  was  taken  while  trying 
to  alight  on  a  Coast  Survey  vessel  in  Bering  Sea,  60  miles  east  south- 
east of  St.  George  Island.  This  was  recorded  by  Bean  (1882,  p. 
161).  The  two  specimens  above  noted  formed  the  sole  basis  for 
Palmers  account  of  the  bird  on  the  Pribilofs  (1899,  p.  418). 

No  other  gyrfalcons  seem  to  have  been  taken  on  the  Pribilofs  until 
the  autumn  of  1913,  when  G.  Dallas  Hanna  took  a  female  specimen 
on  St.  George  Island,  November  29.  It  had  been  noted  on  the  cliffs 
along  the  shore  for  about  a  month.  During  the  autumn  of  1914,  on 
St.  Paul,  he  observed  a  few  others,  as  follows:  September  14,  1 ;  Sep- 
tember 20,  1;  October  14,  1  (taken)  ;  October  25,  3;  November  20,  1. 
Another  individual  was  shot  on  St.  George  Island  during  the  winter 
of    1915-16.     During    the    following    winter,    1916-17,    St.    George 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  83 

Island  was  visited  by  so  many  gyrfalcons  that  the  local  wren  (N an- 
nus alascensis)  and  the  rosy  finch  (Leucosticte  griseoucha)  were 
nearly  extirpated.  On  November  30,  6  were  seen,  3  of  which  were 
killed. 

By  offering  a  reward  Hanna  was  able  to  secure  a  series  of  13 
specimens  altogether.  Six  of  these,  taken  during  December,  1916, 
and  January  and  February,  1917,  are  in  the  U.  S.  National  Museum. 
These  are  mainly  immature  birds,  but  at  least  one  in  adult  plumage 
is  included  in  the  series.  Mr.  Hanna  was  told  that  the  birds  were 
also  unusually  common  on  St.  Paul  Island  during  the  same  winter, 
but  their  destructive  habits  seem  not  to  have  resulted  so  disastrously 
to  the  smaller  resident  birds  of  St.  Paul  as  was  the  case  on  St. 
George.  Although  all  the  available  Pribilof  specimens  have  been 
examined  by  the  writer,  his  studies  have  not  progressed  far  enough 
to  permit  a  decision  as  to  the  form  represented  there.  These  birds 
are  all,  of  course,  migrants  from  some  other  region,  but  whether  from 
North  America  or  from  Asia  can  not  be  determined  without  a  more 
exhaustive  study  than  is  justified  in  the  present  connection.  The 
subspecific  designation  above  used  is  therefore  employed  only  tenta- 
tively. It  is  believed,  however,  that  only  one  form  inhabits  north- 
western North  America,  to  which  the  name  Falco  sacer  (Forster, 
Phil.  Trans.,  vol.  62,  pp.  383,  423,  1772)  appears  to  be  applicable. 

Food. — G.  Dallas  Hanna  states  in  field  notes  that  the  stomach  of 
the  bird  killed  on  St.  George  Island,  November  29,  1913,  contained 
feathers  of  a  rosy  finch  (Leucosticte  griseonucha) ,  and  that  three 
birds  shot  there  on  November  30,  1916,  had  eaten  snow  buntings  and 
rosy  finches. 

Falco  peregrinus  anatum  Bonaparte.     Duck  Hawk. 

Palmer  added  the  duck  hawk  to  the  Pribilof  list  (1899,  p.  419). 
In  the  summer  of  1890,  on  St.  Paul,  he  "  dug  the  remains  of  one  from 
a  sand  dune ;  it  had  been  killed  the  previous  December."  Hahn 
found  a  note  on  this  species  in  the  St.  Paul  Island  log  under  date  of 
December  14,  1904 :  "  One  shot  but  lost."  The  writer  saw  a  duck 
hawk  about  the  cliffs  on  Tolstoi,  St.  Paul,  June  26,  1914;  it  was  not 
identifiable  subspecifically. 

Falco  peregrinus  pealei  Ridgway.     Peale  Falcon. 

This  form  of  the  duck  hawk  was  added  to  the  Pribilof  list  by 
Hanna,  who  took  a  female  specimen  on  St.  George  Island,  March  12, 
1914,  and  has  put  the  capture  on  record  (1916,  p.  401) .  The  specimen 
is  now  No.  242571  of  the  Biological  Survey  collection.  According 
to  a  manuscript  report  in  the  Biological  Survey,  H.  P.  Adams  noted 
one  on  St.  George  Island,  January  13,  1917. 


84  NORTH   AMERICAN   FAUNA.  [No.  46 

Family  STRIGIDAE. 

Asio  flammeus  Pontoppidan.     Short-eared  Owl. 

The  short-eared  owl  was  added  to  the  Pribilof  list  by  Palmer 
(1899,  p.  419)  on  the  basis  of  one  seen  by  him  near  the  village  of 
St.  Paul,  June  17,  1890.  The  natives  reported  seeing  the  species 
every  winter. 

W.  L.  Hahn  took  a  specimen  on  St.  Paul  Island,  November  8, 1910, 
which  is  now  in  the  collection  of  the  Biological  Survey.  It  measured 
in  the  flesh:  Length,  14  inches;  extent,  39f.  A  second  specimen, 
also  in  the  Survey  collection,  was  taken  by  Hanna  on  St.  George, 
February  28,  1914. 

Food. — William  Palmer  noted  that  the  bird  under  observation 
by  him  "  often  visited  the  same  place  during  the  night  or  early  in 
the  morning  to  feed  upon  the  least  auklets  which  were  breeding  in 
the  vicinity  "  and,  he  adds : 

I  often  found  on  the  moss  a  round  patch  of  feathers  which  showed  the  fate 
of  a  Choochkie.  A  few  are  seen  every  winter  by  the  natives,  but  they  all 
agree  that  hawks  and  owls  are  more  abundant  on  St.  George,  owing  to  the 
presence  of  numerous  lemmings,  which  are  entirely  absent  from  St.  Paul. 

W.  L.  Hahn  reports  that  the  stomach  of  the  specimen  collected  by 
him  contained  three  shrews  (/Sorex  pribilof 'ensis). 

Cryptoglaux  funerea  magna  Buturlin.     Buturlin  Hawk  Owl. 

A  female  hawk  owl  taken  by  W.  L.  Hahn,  on  St.  Paul  Island, 
January  26,  1911,  was  recorded  by  Evermann  (1913,  p.  18)  under  the 
name  Cryptoglaux  funerea  funerea  and  was  considered  a  new  record 
for  the  Pribilofs  and  for  North  America.  The  specimen,  which  is 
in  the  collection  of  the  Biological  Survey,  has  recently  been  examin- 
ed by  Dr.  H.  C.  Oberholser,  in  the  light  of  additional  material,  and 
found  to  be  referable  to  C.  f.  magna,  described  by  Buturlin  from  the 
Kolyma  River,  Siberia,  and  has  been  recorded  by  him  (1922,  p.  73). 
G .  f.  magna  thus  replaces  C.  f.  funerea  as  a  Pribilof  and  a  North 
American  bird. 

According  to  Hahn's  field  catalogue  this  specimen  measured  in 
the  flesh:  Length,  10\  inches;  extent,  23^.  Its  stomach  was  empty 
save  for  a  few  hairs. 

Cryptoglaux  funerea  richardsoni  Bonaparte.     Richardson  Owl. 

Evermann  added  the  Richardson  owl  to  the  Pribilof  list  on  the 
basis  of  a  male  specimen  taken  by  M.  C.  Marsh  on  St.  Paul  Island, 
February  2,  1912  (1913,  p.  18).  The  field  catalogues  give  the  fol- 
lowing notes  taken  from  the  fresh  specimen:  Length,  9f  inches; 
extent,  22;  iris,  yellow;  cere,  blackish  behind,  pale  forwards;  bill, 
pale  along  culmen  and  gonys,  the  rest  black;  soles,  light  yellowish; 
claws,  black. 


1923.]  BIRDS   OF   THE   PRTBTLOF   ISLANDS.  85 

Notes  taken  from  the  St.  Paul  Island  log  by  Hahn  record  the 
capture  of  an  owl,  which  from  the  detailed  description  could  be  no 
other  than  this  species,  on  March  20,  1905 ;  another  was  shot  on  May 
22,  and  one  was  seen  on  May  23,  of  the  same  year. 

Food. — The  stomach  of  the  specimen  collected  by  Marsh,  Febru- 
ary 2,  1912,  contained  bones  and  fur  of  the  house  mouse. 

Nyctea  nyctea  (Linnaeus).     Snowy  Owl. 

The  boreal  snowy  owl  is  rather  frequently  observed  on  both  St. 
Paul  and  St.  George  during  the  fall  and  winter  months.  On  a  few 
occasions  it  has  been  observed  in  summer,  but  it  has  never  been 
found  to  breed.  It  was  first  recorded  from  the  Pribilofs  by  Elliott 
(1881,  p.  136),  who  states  that  it  was  occasionally  taken  on  either 
island,  but  that  he  saw  none. 

The  following  are  the  definite  records  of  occurrence  that  have 
come  to  the  attention  of  the  writer:  Palmer  took  1  on  St.  Paul  on 
June  13,  1890,  and  states  that  1  was  shot  on  St.  George  on  June  10 
of  the  same  year.  He  is  also  authority  for  the  statement  that  7  in- 
dividuals were  shot  by  one  of  the  Treasury  agents  during  the  win- 
ter of  1884-85  (island  not  stated),  and  that  F.  A.  Lucas  saw  3  dur- 
ing the  summer  of  1896.  W.  L.  Hahn  found  records  in  the  St.  Paul 
Island  log  to  the  effect  that  2  were  seen  December  4,  1904,  and  that 
1  was  shot  March  5,  1905.  Hahn  himself  noted  the  species  on  St. 
Paul  as  follows:  November  5,  1910,  1  taken  near  Cove  salthouse: 
November  8,  1  seen  near  Polovina;  November  9,  2  seen,  1  shot;  Jan- 
uary 18,  1911,  1  seen  at  Black  Bluffs.  M.  C.  Marsh  took  specimens 
on  the  same  island  on  November  20  and  December  10,  1911.  Hanna, 
collecting  on  St.  George  in  the  fall  of  1913,  noted  the  birds  as  fol- 
lows: September  4,  1  seen;  September  28,  1  seen  at  Garden  Cove; 
October  9,  1  seen;  December  21,  1  seen;  he  saw  another  on  St. 
George,  July  20,  1914.  On  St.  Paul,  in  1915,  Hanna  took  a  speci- 
men, now  in  the  U.  S.  National  Museum,  on  February  7,  and  later 
observed  the  species  as  follows:  April  4,  1;  April  16,  1  (pursuing 
crested  auklets)  ;  May  18,  2  seen  on  Hutchinson  Hill.  On  St. 
George,  he  saw  1  on  November  5,  1916. 

The  field  catalogues  show  the  following  measurements  of  speci- 
mens taken  on  St.  Paul:  Female,  November  5,  1910,  Hahn,  length, 
25  inches,  extent,  63-]-;  male,  November  20,  1911,  Marsh,  length,  23, 
extent,  57;  male,  December  10,  1911,  length,  21| ;  extent,  55£. 

Food. — G.  Dallas  Hanna  states  in  a  field  note  that  an  owl  of  this 
species  has  been  known  to  snatch  from  the  water  an  old-squaw  duck 
which  had  been  shot  by  a  hunter.  Usually,  however,  these  owls  re- 
main on  the  highlands  and  apparently  feed  on  the  birds  found  there. 
No  evidence  has  been  seen  that  they  feed  on  the  St.  George  lem- 


86  NORTH   AMERICAN   FAUNA.  [No.  46 

mings.  Two  stomachs  of  the  snowy  owl,  both  from  St.  Paul  (June 
13.  1890,  Palmer;  and  February  7,  1915,  Hanna)  have  been  ex- 
amined, one  of  which  contained  remains  of  the  least  auklet  {Aethia 
pusilta)   and  the  other  a  Pribilof  shrew   (Sorex  pribilofensis). 

W.  L.  Hahn  examined  one  stomach  which  contained  feathers  of 
the  burgomaster  gull;  and  Marsh,  two,  each  of  which  held  three 
house  mice  (Mus  muscvlus). 

Family  CUCULIDAE. 

Cuculus  canorus  telephonus  Heine.,    Kamchatkan  Cuckoo. 

A  Kamchatkan  cuckoo  taken  by  William  Palmer  at  Northeast 
Point,  St.  Paul  Island,  July  4,  1890,  the  capture  of  which  is  detailed 
by  him  (1899,  p.  419),  was  the  first  taken  in  North  America  and 
remains  the  only  record  for  the  Pribilofs.  It  breeds  in  Kamchatka, 
Manchuria,  Japan,  and  China. 

Food. — Palmer  states  (1.  c.  p.  420)  that  the  stomach  of  this  bird, 
which  was  examined  by  Dr.  S.  D.  Judd,  contained  more  than  100 
crane  flies,  many  of  them  females  full  of  eggs. 

Family  PICIDAE. 

Colaptes  auratus  luteus  Bangs.     Northern  Flicker. 

A  specimen  of  the  northern  flicker,  obtained  by  James  Judge  on 
St.  George  Island  in  the  autumn  of  1904  (a  short  time  previous  to 
October  5),  was  sent  to  the  United  States  National  Museum.  It  was 
recorded  by  A.  H.  Clark  (1910,  p.  60)  and  is  catalogued  as  No. 
191769  of  the  bird  collection. 

Family  MICROPODIDAE. 

Micropus  pacificus  (Latham).     Japanese  Swift. 

A  female  specimen  of  the  Japanese  swift  was  taken  by  G.  Dallas 
Hanna  on  St.  George  Island  August  1,  1920.  It  was  flying  about 
over  the  tundra  and  along  the  cliffs.  The  occurrence  has  been 
recorded  by  Mailliard  and  Hanna  (1921,  p.  93),  and  the  specimen  is 
deposited  in  the  Museum  of  the  California  Acadenry  of  Sciences. 

The  Japanese  swift  breeds  over  a  considerable  part  of  eastern 
Asia,  including  Kamchatka,  Japan,  and  northern  China  and  has 
been  reported  from  the  Commander  Islands. 

Food. — The  stomach  of  this  bird  contained  remains  of  numerous 
dung  flies  (Scatophaga  crinita)  and  of  a  few  other  Diptera. 

Family  CORVIDAE. 

Corvus  corax  principalis  Ridgway.     Northern  Raven. 

The  raven  has  only  recently  been  observed  in  its  natural  state  on 
the  Pribilofs.    Elliott  (1874,  unpaged,  and  1875,  p.  178)  is  authority 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  87 

I  for  the  statement  that  the  species  was  several  times  introduced  by 
i  the  Russians,  but  that  none  of  the  birds  remained. 

Dall,  in  an  account  of  the  birds  of  the  western  Aleutians  (1874,  p. 
274),  states  that  a  few  young  ones  were  taken  to  St.  Paul  in  the 
spring  [of  1873]  in  the  hope  that  they  might  be  of  use  in  devouring 
the  remains  of  seals  killed  there,  "  which  cause  a  very  offensive  odor 
during  the  hunting  season." 

G.  Dallas  Hanna  has  recently  recorded  (1920b,  p.  174)  the  occur- 
rence of  the  species  on  St.  Paul  as  follows : 

During  January,  1919,  heavy  ice  surrounded  St.  Paul  Island,  but  the  natives 
were  able  to  get  out  from  shore  in  their  boats  in  the  open  leads  for  the  excellent 
duck  shooting.  On  one  of  these  occasions  three  ravens  were  seen  near  Sea 
Lion  Rock.  One  was  shot  and  wounded  but  could  not  be  secured.  *  *  * 
identification  could  hardly  be  mistaken.  The  occurrence  was  confirmed  by 
several  reliable  men  to  whom  the  species  was  well  known  when  they  were  in 
Unalaska.  So  in  this  case  it  seems  worth  while  to  make  the  record  without 
the  specimen. 

Family  ICTERIDAE. 

Euphagus  carolinus  (Miiller).     Rusty  Blackbird. 

A  specimen  of  the  rusty  blackbird  taken  near  the  lake  at  Northeast 
Point,  St.  Paul  Island,  October  20,  1911,  by  M.  C.  Marsh,  proved  to 
be  the  first  noted  from  the  Pribilofs,  and  was  recorded  by  Evermann 
(1913,  p.  18.)  According  to  field  catalogue  this  bird  measured  in 
the  flesh :  Length,  9T%  inches ;  extent,  14^. 

A  second  specimen,  taken  on  St.  Paul  in  the  autumn  of  1915  and 
presented  by  George  Haley  to  Hanna,  was  sent  to  the  Biological 
Survey.    It  is  now  catalogued  as  No.  242573. 

Family  FRINGILLIDAE. 

Coccothraustes   coccothraustes  japonicus    (Temminck   and   Schlegel).     Japa- 
nese Hawfinch. 

Evermann's  account  (1913,  p.  18)  of  a  male  specimen  of  the 
Japanese  hawfinch  taken  by  M.  C.  Marsh  near  the  village,  St.  Paul 
Island,  November  1,  1911,  added  this  species  to  the  Pribilof  and  to 
the  North  American  lists.  The  following  notes,  taken  from  Marsh's 
field  catalogue,  were  made  from  the  fresh  specimen :  Iris  pale ;  lower 
mandible  pale,  the  upper  rather  bronzed,  its  base  paler;  culmen  black 
near  tip ;  tarsus  and  feet  uniform  pale ;  claws  darker. 

The  species  has  a  range  in  eastern  Asia  including  Japan,  Korea, 
Northern  China,  and  eastern  Siberia. 

Fringilla  montifringilla  Linnaeus.     Brambling. 

A  male  brambling,  the  first  known  to  have  been  taken  in  North 
America,  was  captured  in  the  watch  house  at  Northeast  Point,  St. 
Paul  Island,  on  October  25,  1914.    It  is  now  in  the  collection  of  the 


88  NORTH   AMERICAN   FAUNA.  [No.  46 

U.  S.  National  Museum  and  has  been  recorded  by  G.  Dallas  Hanna, 
who  preserved  the  specimen  (191G,  p.  400). 

The  brambling  has  a  very  extensive  range  in  northern  Eurasia, 
breeding  regularly  east  to  Kamchatka  and  wintering  mainly  in  south- 
ern Asia. 

Food. — The  stomach  of  the  bird  collected  contained  bits  of  seeds  of 
bunchberry  (C'ornus  stiecica),  90  per  cent;  and  fragments  of  bugs 
(Hemiptera)  and  flies  (Diptera),  10  per  cent. 

Pinicola  leucura  kamtschathensis  Dybowski.  Kamchatkan  Pine  Grosbeak. 
A  fine  adult  male  specimen  of  the  Kamchatkan  pine  grosbeak, 
taken  on  the  tundra  on  St.  George  Island  in  October,  1915,  and  pre- 
served by  A.  H.  Proctor,  has  been  recorded  by  J.  H.  Riley  (1917,  p. 
210).  It  forms  the  first  record  for  the  Pribilofs  and  for  North 
America.  It  was  sent  to  the  U.  S.  National  Museum,  where  it  is  now 
catalogued  as  No.  253121  of  the  bird  collection.  As  indicated  by  its 
name,  it  is  the  breeding  form  of  Kamchatka. 

Loxia  leucoptera  leucoptera  Gmelin.     White-winged  Ceossbill. 

A  female  white-winged  crossbill  was  taken  by  G.  Dallas  Hanna 
near  Halfway  Point,  St.  Paul  Island,  August  9,  1920,  and  has  been 
recorded  by  Mailliard  and  Hanna  (1921,  p.  93).  When  shot  the  bird 
was  apparently  feeding  on  the  unripe  seeds  of  wild  parsnip.  It  is 
now  in  the  collection  of  the  California  Academy  of  Sciences. 

Food. — The  stomach  of  this  bird  was  entirely  filled  with  remains 
of  blowflies  (Oalliphora  vomitoria) .  This  is  a  remarkable  meal  for  a 
crossbill  and  no  doubt  reflects  the  predominance  of  blowflies  among 
food  items  available  to  the  bird  at  the  time. 

Leucosticte  griseonucha  (Brandt).     Aleutian  Rosy  Finch. 

The  beautiful  Aleutian  rosy  finch  is  a  rather  common  breeder  on 
the  Pribilofs,  nesting  on  the  three  larger  islands,  St.  George,  St. 
Paul,  and  Otter,  and  remaining  in  small  numbers  throughout  the 
winter.  It  is  the  most  familiar  species  on  the  islands,  especially  on 
St.  George,  where  it  is  most  common.  It  was  first  added  to  the 
Pribilof  fauna  by  Dall,  who  took  specimens  on  St.  George  in  1868 
(Dall  and  Bannister,  1869,  p.  282). 

Although  a  few  may  be  present  in  winter  the  bulk  of  the  summer 
residents  arrive  in  early  spring.  Hahn  recorded  them  as  numerous 
on  St.  Paul  April  4,  1911,  when  they  were  heard  singing  for  the 
first  time,  and  as  evidently  pairing  on  April  5.  Hanna,  making  ob- 
servations on  St.  George  in  1914,  noted  the  birds  as  very  common, 
singing  and  apparently  mating,  on  March  28  and  April  8,  and  esti- 
mated the  number  seen  on  the  latter  date  as  500.  On  April  22  he 
considered  them  much  more  abundant  than  in  winter,  and  on  May 
6  estimated  a  total  of  2,000  birds  seen. 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  89 

On  St.  Paul,  where  the  bird  is  less  abundant,  Hanna's  notes  for 
the  spring  of  1915  follow:  March  31,  30  seen,  first  heard  singing; 
April  6,  30  seen,  singing  general ;  April  18,  60  seen,  paired ;  April  24, 
about  300  seen. 

A  nest  and  a  set  of  5  eggs  were  taken  by  Lutz  on  Otter  Island, 
June  16,  1884  (1889,  p.  31).  Several  sets  of  fresh  eggs  were  col- 
lected by  Hanna  on  St.  George,  June  15,  1914;  on  June  16  a  set  of 
6  about  one-half  incubated  was  found.  A  female  bird  taken  July 
15,  1914,  was  nearly  ready  to  lay  what  was  presumably  her  second 
set  of  eggs.  The  previous  year,  also  on  St.  George,  Hanna  saw  one 
young  bird  not  able  to  fly  on  September  15,  and  a  brood  just  flying 
on  September  18.  H.  P.  Adams  took  eggs  on  St.  George  on  June  4, 
1916. 

During  the  summer  of  1914  the  writer  found  the  bird  common  on 
St.  Paul  Island.  On  June  22  a  nearly  completed  nest  was  found  on  a 
narrow  shelf  beneath  an  arched  rock  about  15  feet  from  the  ground. 
On  July  4  this  nest  contained  its  complement  of  5  eggs.  Another  nest 
found  the  same  day  in  a  small  cavity  on  the  face  of  a  cliff  contained  5 
eggs  which  were  obviously  on  the  point  of  hatching.  The  first  young 
out  of  the  nest  were  seen  on  July  2.  The  bird  continued  to  be  abun- 
dant up  to  the  time  of  my  departure  the  last  of  August.  The  nests 
are  quite  bulky  and  are  built  of  grasses  and  the  dry  stalks  of  various 
herbaceous  plants,  with  a  lining  of  fine  grass  and  feathers.  Hanna 
found  a  nest  on  St.  George  in  1914  which  had  a  lining  of  reindeer 
hair.    The  eggs,  usually  5  or  6,  are  white  with  a  faint  gloss. 

Hahn,  on  St.  Paul,  noted  the  birds  as  common  during  September, 
October,  and  November,  1910 ;  December  3,  20  seen ;  December  8,  a 
flock;  December  16,  numerous;  December  21,  a  few;  January  2,  1911, 
2  seen;  February  1,  1;  February  2,  1;  February  27,  3;  March  2,  30 
or  40. 

Hanna,  making  observations  on  St.  George  in  1913,  saw  upwards 
of  1,000  at  Garden  Cove,  on  September  28 ;  many  were  seen  at  various 
times  during  the  fall  and  winter.  In  1914,  on  St.  Paul,  he  observed 
approximately  the  following  numbers:  September  14,  16,  20,  and 
October  12,  about  150  birds  each  day;  October  17,  20;  October  25, 
150 ;  November  27,  200 ;  December  4,  200.  1915,  January  10,  25  ;  Feb- 
ruary 18,  50;  March  5,  30. 

Ten  male  specimens  collected  and  measured  in  the  flesh  by  M.  C. 
Marsh  on  St.  Paul  in  the  fall  and  winter  of  1911-12,  ranged  in  length 
from  7f5¥  inches  to  8f,  and  in  extent  of  wing  from  12^  inches  to  14|. 
A  male  bird  taken  by  Hanna  on  St.  George,  December  19,  1913,  is  re- 
corded as  having  eyes,  brown;  feet,  black;  and  bill,  yellow,  black 
at  tip. 

In  the  winter  of  1916-17,  owing  to  an  unusual  visitation  of  gyrfal- 
cons,  the  rosy  finches  living  on  the  Pribilofs  were  nearly  all  destroyed. 


90  NORTH    AMERICAN   FAUNA.  [No.  46 

During  a  trip  made  around  St.  George  in  May,  1917,  Hanna  saw  not 
over  4  pairs.  Since  that  time,  however,  probably  through  migra- 
tion, the  birds  have  nearly  or  quite  reached  their  former  numbers. 
Food. — William  Palmer  says  of  the  Aleutian  rosy  finch  on  the 
Pribilofs  (1899,  p.  426)  : 

I  have  picked  from  the  mouth  of  a  freshly  killed  bird  the  most  minute  insects, 
and  have  watched  them  feeding  on  the  drying  carcass  of  a  seal  hanging  outside 
of  the  house  of  an  Aleut,  and  they  do  not  scorn  the  possibilities  afforded  by 
the  decaying  seal  carcasses  on  the  killing  ground. 

The  birds  visited  the  latter  situations,  no  doubt,  for  the  insects  at- 
tracted there,  not  with  any  intent  of  eating  flesh  or  offal.  Indeed, 
they  take  a  surprisingly  low  proportion  of  animal  food,  if  we  may 
rely  on  the  results  from  the  collection  of  22  stomachs  examined  for 
the  present  report. 

The  food  in  these  stomachs  was  found  to  be  vegetable,  75.5  per  cent; 
and  animal,  24.5  per  cent.  The  plant  diet  was  chiefly  seeds,  but  in  a 
few  cases  bits  of  leaves  and  fruiting  capsules  were  eaten.  Seeds  of 
crowberry  (Empetrum  nigrum)  were  found  more  frequently  than 
any  other  (i.  e.,  in  6  gizzards)  and  from  20  to  40  seeds  were  present 
in  certain  of  these  stomachs.  The  largest  numbers  of  seeds  eaten  by 
any  of  these  rosy  finches  were  250  and  450,  in  two  instances,  of  those 
of  brook  saxifrage  {Chrysosplenium  beringianum) .  In  one  case  also 
160  seeds  of  sea  parsley  (Ligusticum  scoticum)  were  contained  in  a 
single  stomach.  Other  seeds  eaten  included  those  of  grass,  rush 
(Juncus  sp.),  sedge  (Carex  sp.),  chickweed  (Alsine  borealis),  butter- 
cup (Ranunculus  sp.),  water  chickweed  (Montia  fontana),  cinque- 
foil  (Potentilla  sp.),  and  bluebell  (Campanula  sp.). 

Of  the  animal  food,  approximately  21  per  cent  of  a  total  of  24.5 
per  cent  consisted  of  two-winged  flies,  2  per  cent  of  beetles,  and  1 
per  cent  of  springtails.  The  flies  consumed  were  chiefly  crane  flies 
(Tipulidae),  and  the  beetles  included  ground  beetles  (Pterostichus 
sp.  and  others),  leaf  beetles  (Chrysomela  sub  sulcata) ,  beach  beetles 
(Aegialites  calif ornicu-s),  and  weevils.  Caterpillars  occurred  in  2 
stomachs  and  springtails  (Aptera:  Collembola)  in  1.  The  latter  in- 
sects were  identified  as  Isotoma  violacca  var.  mucronata,  and  the 
record  is  the  first  of  the  occurrence  of  this  species  on  American  ter- 
ritory. 

Mr.  Hahn  noted  the  rosy  finch  feeding  on  seeds  of  poochka,  or 
wild  parsnip  (C oelopleurum  gmelini),  and  of  rye  grass,  and  Mr. 
Hanna  observed  that  in  winter  they  appeared  to  feed  almost  ex- 
clusively on  the  seeds  of  poochka. 

Acanthis  bornemannii  exilipes   (Coues).     Hoary  Redpoix. 

G.  Dallas  Hanna  has  added  the  hoary  redpoll  to  the  Pribilof  list 
by  recording  a  female  specimen  taken  at  the  village,  St.  Paul  Island, 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  91 

April  21,  1915  (1916,  p.  401).  Later  notes  taken  on  St.  Paul  by  the 
same  observer  include  the  following:  April  20,  1915,  1  seen;  August 
22,  6  seen  in  company  with  a  small  flock  of  A.  linaria. 

Food. — The  single  stomach  of  this  bird  available  for  examination 
(St.  Paul,  April  20,  1915)  contained  seeds  of  a  plant  of  the  pink 
family  {Lychnis  apetala) ,  of  a  chickweed  {Alsine  sp.),  of  a  grass, 
and  of  an  unidentified  kind. 

Acanthis  linaria  linaria  (Linnaeus).     Redpoll. 

The  common  redpoll  was  added  to  the  Pribilof  fauna  by  H.  W. 
Elliott,  who  took  specimens  on  St.  Paul,  June  21,  1872.  These  he 
preserved  in  alcohol,  but  they  were  in  some  way  lost  (1881,  p.  136)  ; 
he  had  previously  mentioned  (1874,  unpaged,  and  1875,  p.  172)  hav- 
ing seen  a  small  flock  of  redpolls  on  St.  Paul  in  late  October,  1872. 
No  others  appear  to  have  been  seen  until  1910,  when  Hahn,  on  St. 
Paul,  observed  redpolls,  probably  of  this  species,  as  follows:  October 
5,  about  50  seen ;  October  19,  about  15 ;  October  20,  3  at  Northeast 
Point.  No  specimens  were  taken  on  these  occasions.  In  1913, 
Hanna,  collecting  on  St.  George,  observed  5  birds  on  September  7, 
and  collected  3.  On  October  18,  9  others,  1  of  which  was  taken, 
were  seen.  These  specimens  are  in  the  collection  of  the  Biological 
Survey.  On  St.  Paul,  in  1914,  Hanna  observed  the  species  as  fol- 
lows: September  14,  2;  October  14,  1;  October  25,  3;  1915:  February 
18,  1 ;  April  18,  1 ;  August  22,  8  seen  and  1  collected  at  Northeast 
Point.  The  bird  is  thus,  in  all  probability,  a  more  or  less  regular 
migrant. 

Food. — G.  D.  Hanna  reports  a  small  flock  observed  feeding  on 
wild  parsnip  on  St.  George  Island,  September  7,  1913.  This  plant 
is  Coelopleurum  gmelini. 

Spinus  pinus  pinus  (Wilson).     Pine  Siskin. 

The  pine  siskin  was  added  to  the  list  of  Pribilof  visitors  by  G.  Dal- 
las Hanna,  who  recorded  the  capture  (1919a,  p.  177,  and  1920a,  p. 
254).    He  comments  (1.  c,  p.  254)  as  follows: 

A  male  pine  siskin  was  secured  from  a  flock  of  12  found  among  the  Northeast 
Point  sand  dunes,  St.  Paul  Island,  on  September  24,  1917.  The  birds  seemed 
to  be  perfectly  satisfied  to  feed  on  the  seeds  of  the  few  ground  plants  which 
grow  there. 

This  specimen  was  deposited  in  the  bird  collection  of  the  U.  S. 
National  Museum,  and  catalogued  as  No.  255168. 

Plectrophenax  nivalis  nivalis  (Linnaeus).    Snow  Bunting. 

A  male  specimen  of  typical  nivalis  taken  by  M.  C.  Marsh  on  St. 
Paul  Island,  March  31,  1912,  was  recorded  by  Evermann  (1913,  p. 
18),  being  thus  added  to  the  Pribilof  list.     A  snowflake  taken  on 


92  NORTH   AMERICAN   FAUNA.  [No.  46 

St.  George  Island,  October  3,  1899,  by  Wilfred  H.  Osgood  also  proves 
to  be  of  the  mainland  race.  It  is  now  No.  165738  of  the  U.  S. 
National  Museum,  Biological  Survey  collection. 

Plectrophenax  nivalis  townsendi  Ridgway.     Pribilof  Snow  Bunting. 

Plectrophcnax  nivalis  townsendi  Ridgway,  Manual  North  Araer.  Birds,  p. 
403,  1887   (type  from  Otter  Island). 

The  breeding  snow  bunting  was  first  definitely  recorded  from  the 
Pribilofs  by  Dall  (in  Dall  and  Bannister,  1869,  p.  283),  under  the 
name  Plectrophanes  nivalis;  it  is  altogether  probable,  however,  that 
Veniaminof  in  1840  (Translation  by  Elliott,  1874,  unpaged,  and  1875, 
p.  242)  referred  to  the  present  species  under  the  name  "  snowfinch." 
The  Pribilof  bird  was  described  as  a  race  of  the  circumpolar  species 
by  Ridgway  from  a  series  of  specimens  which  included  No.  106695 
(IT.  S.  Nat.  Mus.,  Otter  Island,  June  8,  1885,  C.  H.  Townsend),  in- 
dicated as  the  type  specimen.     It  is  a  year-long  resident  of  the  islands. 

The  species  nests  on  St.  George,  St.  Paul,  and  Otter  Islands. 
Palmer  (1899,  p.  425)  has  described  the  nest  and  eggs.  He  took 
sets  on  June  4  and  24,  1890;  these  eggs  and  a  set  taken  on  Otter 
Island,  June  15,  1884  (Lutz,  1889,  p.  31),  furnish  the  only  definite 
nesting  dates  available. 

During  the  breeding  season,  the  snow  buntings  keep  rather  closely 
to  the  more  elevated  parts  of  the  islands  and  are  seldom  seen  about 
the  village  or  the  beaches.  Hahn  noted  the  bird  on  St.  Paul  as  fol- 
lows: Numerous  during  fall  of  1910;  occasional  in  December;  not 
found  in  village  during  mild  weather;  numerous  in  village  Decem- 
ber 21  and  27;  also  observed  there  on  January  2,  4,  23,  and  28,  1911; 
February  4  to  10,  appear  in  the  village  daily;  first  heard  singing 
March  18,  singing  full  song  March  31.  By  April  16  the  birds  had 
lost  the  buffy  tinge  and  were  pure  black  and  white. 

Hanna,  making  observations  on  St.  George  in  1913,  noted  the 
species  frequently  during  the  autumn  and  winter,  stating  that  the 
birds  left  the  higher  parts  of  the  island  about  the  end  of  September. 
They  were  singing  and  mating  on  May  6,  1914.  H.  P.  Adams  noted 
the  species  on  St.  George,  January  13,  1917. 

On  St.  Paul  in  the  autumn  of  1914  Hanna  noted  the  bird  as  fol- 
lows :  October  4,  common  in  village ;  October  17,  2 ;  October  25,  200 ; 
November  27,  20;  December  13,  50.  1915:  January  10,  50;  February 
18,  50;  March  20,  10;  April  6,  4  (singing)  ;  April  18,  30;  April  24, 
30 ;  May  6,  4. 

My  own  experience  with  the  species  is  limited,  being  confined 
to  the  observation  and  collection  of  a  few  specimens  on  both  St. 
Paul  and  St.  George  in  1914.  when  the  birds,  sometimes  accompanied 
by  their  young,  were  found  to  be  fairly  common  on  their  breeding 
grounds. 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  93 

There  are  numerous  specimens  in  the  collection  of  the  Biological 
Survey  and  the  U.  S.  National  Museum.  Specimens  from  St.  Paul 
measured  in  the  flesh  by  Hahn  and  Marsh  varied  as  follows :  Length, 
7  to  7f  inches;  extent,  12i  to  13|. 

Food. — The  stomachs  of  Pribilof  snow  buntings  collected  by  Wil- 
liam Palmer  have  been  reexamined,  but  the  writer  is  unable  to  recon- 
cile great  discrepancies  between  the  findings  of  Dr.  Judd  and  those 
of  the  present  investigation.  He  contents  himself,  therefore,  with  a 
simple  statement  as  to  what  was  identified  in  the  stomachs  available, 
a  total  of  21,  including  two  for  which  percentages  were  not  estimated 
and  which,  therefore,  are  not  included  in  the  calculations  of  food 
proportions.  The  articles  of  food  composing  more  than  1  per  cent 
of  the  total  contents  of  the  19  well-filled  stomachs  were :  Vegetable 
matter,  chiefly  seeds,  50.5  per  cent;  two-winged  flies  (Diptera),  32.52 
per  cent;  beetles,  5.3  per  cent,  and  caterpillars  and  moths,  1.5  per 
cent.  The  seeds  eaten  included  those  of  sedge  {Carex  sp.),  rush 
(Juncus  sp.),  water  chickweed  (Montia  fontana),  buttercup  (Ranun- 
culus sp.),  cinquefoil  (Potentilla  sp.),  crowberry  (Empetrum 
nigrum),  and  lousewort  (Pedicularis  sp.).  From  47  to  81  seeds  of 
the  latter  were  found  in  the  stomachs  of  the  four  birds  eating  them. 

The  Diptera  eaten  by  the  snowflakes  were  largely  crane  flies 
(Tipulidae),  but  dung  flies  (Scatophagidae)  were  eaten  as  well  as 
larvae  of  a  snipe  fly  (Spania  edeta).  Among  the  beetles,  the  leaf 
beetle  (Chrysomela  subsulcata)  was  identified  more  frequently  (in 
5  stomachs)  than  any  other  species,  among  which  were  ground  beetles 
(Amara  sp.  and  Pterostichus  sp.),  rove  beetles  (T echinus  sp.),  and 
beach  beetles  (Aegialites  calif 'ornicus) .  Hymenoptera  (Amblyteles 
sp.  and  a  sawfly)  were  found  in  two  stomachs,  and  a  spider  in  one. 

Plectrophenax  hyperboreus  (Ridgway).     McKay  Snow  Bunting. 

(t.  Dallas  Hanna  has  added  the  McKay  snow  bunting  to  the 
Pribilof  list  (1919a,  p.  176),  and  later  commented  as  follows  (1920a, 
p.   254)  : 

Since  this  species  is  known  to  wander  from  its  only  breeding  place,  St. 
Matthew  Island  group,  to  the  mainland  of  Alaska,  it  has  been  expected  and 
searched  for  on  the  Pribilofs  for  several  years.  But  it  was  not  definitely  known 
to  come  until  March  30,  1918,  when  a  male  in  full  winter  plumage  was  secured 
on  St.  Paul  Island.  It  and  a  female,  which  escaped,  were  found  on  the  top  of 
Rush  Hill,  the  highest  point  of  the  island. 

This  specimen  is  now  catalogued  as  No.  255167  of  the  collection 
of  the  U.  S.  National  Museum. 

Food. — The  stomach  of  the  specimen  above  mentioned  contained 

the  following  food:  Remains  of  more  than   160  seeds,  apparently 

of  a  sandwort  (Arenaria  sp.),  57  per  cent;  14  or  more  rove  beetles 

(Staphylinidae),  8  per  cent;  1  larva  of  a  leaf  beetle  (Chrysomela 

108731°— 23 7 


94  NORTH    AMERICAN    FAUNA.  [No,  4« 

subsulcata),  4  per  cent;  3  sawfly  cocoons,  15  per  cent;  1  moth 
cocoon,  15  per  cent;  4  fly  pupae,  and  1  adult  long- footed  fly  (Hydro- 
phones innotatus),  1  per  cent. 

Calcarius  lapponicus  alascensis  Ridgway.    Alaska  Longspur, 

Calcarius  lapponicus  alascensis  Ridgway,  The  Auk,  vol.  15,  p.  320,  Oct.  1898 
(type  from  St.  Paul  Island). 

The  longspur,  one  of  the  few  small  land  birds  breeding  on  the 
Pribilofs,  is  an  abundant  summer  resident.  It  is  in  some  respects  the 
most  pleasing  of  the  bird  population,  owing  to  its  habit  of  delivering 
its  beautiful  song  on  the  wing,  while  it  slowly  flutters  earthward.  It 
usually  arrives  in  early  May,  and  departs  by  the  end  of  October.  It 
was  first  recorded  by  Elliott  (1874,  unpaged,  and  1875,  p.  178). 

Hahn  noted  the  bird  as  common  and  singing  on  St.  Paul,  May  13, 
1911,  and  as  singing  on  the  wing  on  May  29.  Hanna  observed  it  on 
St.  Paul  in  1915  as  follows:  May  6,  6  seen;  May  9,  15;  May  16,  40; 
May  28,  abundant.  On  St.  George,  in  1917,  he  notes  that  the  birds 
arrived  in  a  body,  already  paired,  on  May  8. 

It  nests  in  late  June  and  July,  and  fresh  eggs  may  be  found  even 
as  late  as  early  August.  Elliott  took  fresh  eggs  on  St.  George  on 
July  7  [1873]  (1874,  unpaged,  and  1875,  p.  178).  Palmer,  in  1890, 
found  nests  on  St.  Paul  as  follows :  June  20,  5  eggs ;  June  21,  5  eggs ; 
June  21,  3  eggs,  2  young;  July  2,  6  young,  well  feathered;  July  5, 
5  young  just  from  the  nest,  Prentiss,  in  1895,  found  the  species  nest- 
ing as  follows :  St.  Paul,  July  1,  5  young  partly  fledged ;  July  2,  6 
eggs,  well  incubated ;  July  3,  6  eggs,  fresh ;  July  6,  3  eggs,  fresh ;  St. 
George,  July  21,  3  eggs,  fresh;  July  22,  1  young,  very  small  (1902, 
p.  102).  The  nest  is  usually  placed  on  a  sloping  place,  sometimes  in 
the  shelter  of  a  tuft  of  grass  or  a  tall  plant. 

During  my  own  visit  to  the  Pribilofs  in  1914  I  found  the  bird 
rather  common  and  quite  generally  distributed  on  St.  Paul,  St. 
George,  and  Otter  Islands,  throughout  my  stay  from  June  21  to  the 
end  of  August.  A  nest  containing  4  fresh  eggs  was  found  on  St. 
George  on  August  4,  at  which  time  the  birds  were  still  giving  their 
pleasant  aerial  song.  During  the  same  autumn,  after  my  departure, 
G.  Dallas  Hanna  observed  large  numbers  of  the  birds  during  Sep- 
tember. On  October  4  he  saw  10,  and  on  October  12  noted  that  all 
had  departed.  In  the  autumn  of  1913,  on  St.  George,  Hanna  had 
observed  the  bird  last  on  September  28,  when  he  saw  about  20.  On 
St.  George,  in  1916,  he  noted  that  it  was  still  abundant  on  October  26. 

Hahn  in  1910  saw  the  species  frequently  during  September  on  St. 
Paul  Island,  and  during  early  October  noted  it  as  follows:  October 
3, 10 ;  October  4,  5,  and  6,  about  50  each  day ;  October  8,  less  common; 
October  10,  observed ;  October  11  (the  last  observation  recorded)  10. 


1923.]  BIRDS   OF    THE   PRIBIL/OF   ISLANDS.  95 

Specimens  taken  on  St.  Paul  by  Hahn  and  Marsh  in  1910  and  1912 
measured  in  the  flesh  as  follows :  Length,  5y\  to  6f  inches ;  extent, 
H  to  lift. 

Food. — The  collection  of  stomachs  of  Alaska  longspurs  from  the 
Pribilof  Islands  consists  of  the  six  collected  by  Palmer  and  ex- 
amined by  Judd,  which  are  said  (1899,  p.  423)  to  have  con- 
tained only  insect  cuticle  and  grinding  material ;  and  18  others,  of 
which  one  contained  food  so  finely  ground  that  percentages  for  its 
constituents  could  not  be  estimated.  The  principal  items  of  diet  of 
the  remaining  17  were:  Seeds  of  various  plants,  26.5  per  cent;  two- 
winged  flies,  38.3  per  cent;  beetles,  15.8  per  cent;  caterpillars  and 
moths,  10  per  cent;  bugs,  6  per  cent;  spiders,  1.4  per  cent;  and 
parasitic  wasps,  1.2  per  cent. 

The  seeds  eaten  by  these  longspurs  were  from  a  variety  of  plants, 
of  which  sedge  (Car  ex  sp.),  cinquefoil  (Potentilla  sp.),  and  violet 
(Viola  langsdorfii)  were  most  frequently  taken.  Others  identified 
were  grass,  water  chickweed  (Montia  fontana),  poppy  (Papaver 
macounii),  brook  saxifrage  (Chrysosplenium  beringianum) ,  crow- 
berry  (Empetrum  nigrum),  and  gentian  (Gentiana  sp.). 

Flies  contributing  to  the  diet  of  the  Alaska  longspur  were  chiefly 
crane  flies  (Tipula  sp.),  but  blowflies  (Cynomyia  hirta  and  Calli- 
phora  vomitoria),  dung  flies  (Scatophaga  crinita),  and  kelp  flies 
(FucelUa  sp.)  also  were  consumed.  The  beetles  were  about  evenly 
divided  between  ground  beetles  (Carabidae),  weevils  (Lophalophus 
inquinatus) ,  and  leaf  beetles  (Chrysomela  sub  sulcata) .  None  of  the 
moths  and  caterpillars  eaten  were  more  closely  identified,  but  among 
the  Hymenoptera,  small  parasitic  wasps  were  named  as  follows : 
Amblyteles,  Campoplex  (?),  and  Tryphon  (?);  and  among  the 
spiders,  the  single  species  Tegenaria  derhami. 

Passerculus    sandwichensis    sandwichensis     Gmelin.     Aleutian     Savannah 
Sparrow. 

Palmer  added  the  Savannah  sparrow  to  the  Pribilof  list  on  the 
basis  of  one  seen  June  3,  1890,  near  Lukanin  Beach,  St.  Paul  Island 
(1899,  p.  422). 

Since  it  seems  more  likely  that  Savannah  sparrows  occurring  on 
the  Pribilof s  should  be  referable  to  the  race  breeding  in  that  region, 
I  include  under  this  heading  those  seen  by  Hanna  on  St.  George 
Island,  August  18, 1913,  when  he  noted  10  individuals.  He  mentions, 
also,  without  giving  details  (1920a,  p.  249),  having  collected  a 
specimen  on  St.  Paul  Island,  probably  referring  to  a  specimen,  now 
No.  255169,  female,  of  the  collection  of  the  U.  S.  National  Museum, 
taken  by  him  on  St.  Paul,  September  24,  1917. 


96  NORTH   AMERICAN   FAUNA.  [No.  46 

Passerculus  sandwichensis  alaudinus  Bonaparte.     Western  Savannah  Spar- 
row. 

A  female  specimen  of  the  western  Savannah  sparrow  taken  on 
St.  George  Island,  September  1,  1913,  by  G.  Dallas  Hanna  has  been 
identified  as  this  form  and  has  been  formally  recorded  by  the  col- 
lector (1916,  p.  402).    It  was  the  only  one  seen  at  the  time. 
Zonotrichia  gambeli  Nuttall.     Gambel  Sparrow. 

Evermann  (1913,  p.  18)   recorded  a  male  Gambel  sparrow  taken 
by  M.  C.  Marsh  on  St.  Paul  Island,  May  24,  1912,  the  only  record 
for  the  Pribilofs.     It  measured  :  Length,  7  inches ;  extent,  9{|. 
Junco  hyemalis  hyemalis  (Linnaeus).     Slate-colored  Junco. 

On  the  same  day  that  he  added  the  pine  siskin  to  the  Pribilof  list, 
G.  Dallas  Hanna  took  specimens  of  the  slate-colored  junco,  before 
unknown  on  the  group.  He  has  recorded  the  occurrence  (1919a,  p. 
177)  and  later  (1920a,  p.  254)  published  particulars,  as  follows: 

The  female  juncos  were  secured  from  a  flock  of  6  at  Northeast  Point,  St. 
Paul  Island,  on  September  24,  1917.  They  were  feeding  about  the  buildings 
there,  apparently  as  contented  as  if  they  were  in  the  midst  of  civilization. 

These  skins  are  now  in  the  collection  of  the  U.  S.  National  Museum, 
being  catalogued  as  Nos.  255170  and  255171. 
Melospiza  melodia  sanaka  McGregor.     Aleutian  Song  Sparrow. 

Among  the  numerous  species  which  have  been  added  to  the  Pribi- 
lof Island  bird  list  by  G.  Dallas  Hanna  is  the  Aleutian  song  sparrow. 
He  has  recorded  its  occurrence  as  follows  (1916,  p.  401)  : 

Two  specimens  were  collected  on  St.  George  Island,  a  male,  October  28,  1913, 
at  the  village;  and  a  female,  December  21,  1913,  at  Garden  Cove.     Both  birds 
were  feeding  on  the  beaches. 
Passerella  iliaca  sinuosa  Grinnell.     Valdez  Fox  Sparrow. 

This  race  of  fox  sparrow  was  added  to  the  Pribilof  list  by  Ever- 
mann, under  the  name  P.  i.  insularis,  on  the  basis  of  a  male  specimen 
collected  on  St.  Paul  Island,  near  the  village,  September  7,  1910,  by 
W.  L.  Hahn.  It  measured  in  the  flesh :  Length,  7f  inches ;  extent,  lOf . 
Dr.  H.  C.  Oberholser.  from  a  recent  critical  comparison  of  this  speci- 
men, identifies  it  as  P.  i.  sinuosa. 

Hahn's  notes  regarding  this  and  other  individuals  observed  follow : 

September  7.  1910,  1  shot  among  the  rocks  near  Gorbatch  Rookery.  High 
northeasterly  winds  had  prevailed  for  several  days.  October  4,  1  seen  near 
where  the  first  was  killed.  Another  seen  among  the  rocks  back  of  Reef  Rookery 
later  the  same  afternoon.     October  15,  1  seen  among  the  rocks  at  Kaminista. 

Passerella  iliaca  unalaschensis  (Gmelin).     Shumagin  Fox  Sparrow. 

This  race,  the  Shumagin  fox  sparrow,  was  added  to  the  Pribilof  list 
by  G.  Dallas  Hanna,  who  thus  records  the  specimen  (1920b,  p.  173)  : 
"  On  May  20,  1919,  a  female  fox  sparrow  was  found  at  Northeast 
Point,  St.  Paul  Island,  feeding  among  the  piles  of  drift  wood.  Mr. 
Mailliard  is  responsible  for  the  identification."  This  specimen  is 
deposited  in  the  collection  of  the  California  Academy  of  Sciences. 


1923.]  BIRDS   OF   THE   PRIBILOF   ISLANDS.  97 

Family  HIRUNDINIDAE. 

Petrochelidon  lunifrons  lunifrons  (Say).     Cliff  Swallow. 

The  cliff  swallow  was  added  to  the  list  of  Pribilof  birds  by  G. 
Dallas  Hanna,  who  thus  records  the  specimen  taken  (1919a,  p.  177, 
and  1920a,  p.  254)  :  "A  cliff  swallow  was  shot  and  skinned  on  St. 
Paul  Island  about  June  10,  1918,  by  a  native  from  whom  the  speci- 
men was  secured.  This  makes  the  second  species  of  swallow  to  be 
collected  on  the  island."  (1.  c,  p.  254.)  The  specimen  is  now  No. 
255172  of  the  bird  collection  of  the  U.  S.  National  Museum. 

Hirundo  rustica  erythrogastra  Boddaert.     Barn  Swallow. 

The  barn  swallow  was  given  a  place  in  the  Pribilof  list  by  Palmer 
on  the  basis  of  a  bird  observed  by  him  near  the  village  of  St.  George, 
May  28,  1890.  The  same  bird  remained  about  the  village  for  nearly 
two  weeks.  Another  was  seen  near  Black  Bluffs,  St.  Paul  Island, 
June  4,  of  the  same  year.  The  species  has  not  been  since  observed. 
Iridoprocne  bicolor  (Vieillot).     Tree  Swallow. 

A  male  specimen  taken  by  John  Hanson,  a  native,  at  Northeast 
Point,  St.  Paul  Island,  May  25,  1922,  and  recorded  by  Mailliard 
(1923,  p.  31),  adds  this  species  to  the  Pribilof  list.  The  specimen  is 
deposited  in  the  California  Academy  of  Sciences. 

Tachycineta  thalassina  lepida  Mearns.     Violet-green  Swallow. 

On  August  22,  1914,  while  G.  Dallas  Hanna  and  myself  were 
traversing  St.  Paul  Island  toward  Zapadni  Rookery,  we  saw  three 
swallows  flying  about  the  cliffs  near  the  head  of  the  lagoon,  and  Mr. 
Hanna  secured  two  of  them.  They  were  at  once  identified  as  this 
species,  thus  first  detected  on  the  Pribilofs.  Mr.  Hanna  has  since 
(1916,  p.  401)  recorded  the  occurrence.  He  also  observed  a  single 
bird  about  some  high  bluffs  on  St.  Paul,  June  22,  1919. 

Food. — The  two  stomachs  of  violet-green  swallows  available  taken 
on  St.  Paul,  August  22,  1914,  were  examined  by  the  late  Prof.  F.  E.  L. 
Beal,  who  recorded  that  they  were  filled  with  finely  ground  Diptera. 

Family  COMPSOTHLYPIDAE. 

Wilsonia  pusilla  pileolata  Pallas.     Pileolated  Warbler. 

Two  mummified  specimens  of  the  pileolated  warbler,  a  male  and 
a  female,  preserved  by  A.  G.  Whitney  on  St.  Paul  Island,  August 
25,  1913,  are  the  first  known  from  the  Pribilofs.  These  specimens, 
as  well  as  an  individual  observed  on  St.  George  Island,  August  20, 
1913,  by  G.  Dallas  Hanna,  were  recorded  by  Hanna  (1916,  p.  402). 

Family  MOTACILLIDAE. 
Anthus  spinoletta  rubescens  (Tunstall).     Pipit. 

The  pipit  seems  to  have  been  first  recorded  from  the  Pribilofs  by 
Nelson  (1887,  p.  208),  who  states  that  the  species  was  taken  on  St. 


98  NORTH   AMERICAN   FAUNA.  [No.  46 

George  Island  on  August  15.  He  evidently  refers  to  a  specimen 
taken  by  W.  H.  Dall  on  St.  George,  August  15,  1868.  This  was 
recorded  by  Palmer  (1899,  p.  421)  as  being  then  in  the  collection  of 
the  U.  S.  National  Museum,  and  formed  the  sole  basis  for  the  inclu- 
sion of  the  bird  in  his  list. 

In  the  course  of  Mr.  Hanna's  work  on  the  Pribilofs  he  has  found 
this  bird  to  be  a  rather  regular  migrant  in  fall,  and  has  taken  it  once 
in  spring. 

In  the  fall  of  1913,  on  St.  George  Island,  Hanna  observed  about 
20  pipits  on  September  1,  takiifg  2  specimens,  and  saw  10  on  Septem- 
ber 4.  In  1914,  on  St.  Paul,  he  noted  the  birds  as  follows:  August 
31,  2  seen,  1  collected;  September  20,  1  taken;  October  12,  1.  The 
specimens  taken  by  him  are  in  the  collection  of  the  Biological 
Survey. 

The  only  spring  record  is  of  a  specimen  taken  by  Hanna  on  St. 
Paul  May  25,  1919,  and  recorded  by  him. 

Food. — This  species  is  reported  by  Hanna  to  feed  during  its 
stay  on  the  islands  in  fall  migration  almost  exclusively  on  maggots 
on  the  killing  fields.  However,  the  contents  of  two  stomachs,  col- 
lected August  31,  1914,  and  September  20,  1916,  contained  no  trace 
of  such  maggots.  The  food  in  these  gizzards  consisted  of  10  per 
cent  vegetable  matter  (seeds  of  a  violet,  Viola  langsdorfii)  and  90 
per  cent  animal  matter.  The  components  of  the  animal  food  were 
beetles  (ground  beetles,  PterosticJius  sp. ;  and  weevils  Lophalophus 
inquinatus),  37  per  cent;  caterpillars,  33.5  per  cent;  plant  bugs 
(Irbisia  sericans),  8  per  cent;  spiders,  7.5  per  cent;  flies,  2.5  per 
cent;  and  Hymenoptera,  1.5  per  cent. 

Anthus  spinoletta  japonicus  (Temminck  and  Schlegel).     Japanese  Pipit. 

G.  Dallas  Hanna  collected  a  Japanese  pipit  on  St.  Paul  Island 
on  August  29,  1916,  thereby  adding  the  species  to  the  list  of  Pribilof 
visitors.  The  specimen,  which  is  now  No.  255173  of  the  bird  collec- 
tion of  the  U.  S.  National  Museum,  was  recorded  by  him  (1919a,  p. 
177,  and  1920a,  p.  251). 

Family  TROGLODYTIDAE. 

Nannus  troglodytes  alascensis  (Baird).     Alaska  Wren. 

Troglodytes  alascensis  Baird,  Trans.  Chicago  Acad.  Sci.,  vol.  1,  315,  186(J 
(type  from  St.  George  Island). 

This  interesting  resident  wren,  the  "  Limmershin  "  (i.  e.,  chew  of 
tobacco)  of  the  natives,  was  first  recorded  from  the  group  by  W.  H. 
Dall  (in  Dall  and  Bannister,  1869,  p.  280)  and  was  formally  charac- 
terized as  a  new  species  by  Baird  in  a  later  article  (1869,  p.  315)  of 
the  same  volume.  The  basis  of  these  accounts,  and  the  type  specimen, 
was  a  bird  taken  by  Dall  on  St.  George  Island  on  August  17, 1868. 


1923.]  BIRDS   OF    THE   PEIBELOF   ISLANDS.  99 

The  species  is  a  year-long  resident  and  varies  greatly  in  abundance 
in  different  years,  becoming  scarce  because  of  especially  severe  winters 
or  an  unusual  visitation  of  predatory  birds,  but  soon  recovering  its 
numbers  evidently  owing  entirely  to  its  rapid  rate  of  increase.  Thus 
Elliott  (1875,  p.  173)  found  it  rather  rare  on  St.  George  in  1873,  but 
quite  numerous  in  1874.  Palmer  observed  a  few  on  St.  George  in 
1890,  and  secured  specimens.  Hanna  took  a  series  of  specimens  on 
St.  George  in  the  autumn  of  1913  and  the  following  winter,  and  his 
notes  show  that  the  birds  were  common,  since  on  one  occasion,  near 
Garden  Cove  on  September  28,  he  estimated  that  he  saw  200  indi- 
viduals. I  saw  many  and  took  a  few  on  St.  George  in  mid-July  and 
early  August,  1914. 

Up  to  1914  no  naturalist  had  ever  seen  one  of  the  birds  on  any 
island  of  the  group  excepting  St.  George,  and  the  natives  declared 
that  it  was  absent  elsewhere.  It  was,  therefore,  a  great  surprise  when 
Mr.  Hanna,  on  October  29,  1914,  took  2  specimens  along  the  shore 
between  East  Landing  ai  d  Reef  Rockery,  St.  Paul  Island.  On  May 
16,  1915,  Hanna  collected  another  specimen  near  Lukanin  Rookery, 
and  the  natives  reported  seeing  2  near  Rock  Point.  During  the  same 
summer,  according  to  notes  kindly  furnished  me  by  Mr.  Hanna,  Mr. 
George  Haley  saw  11  individuals  on  Otter  Island,  this  being  the  first 
record  for  that  place.  The  birds  since  seem  to  have  become  well 
established  there,  as  Hanna  (1920b,  p.  175)  states  that  it  bred  there 
in  1916,  1917,  and  1918.  He  states  also  (1920a,  p.  250)  that  a  wren, 
probably  from  Otter  Island,  was  seen  on  St.  Paul  in  the  summer  of 
1918.  It  seems  likely,  therefore,  unless  the  species  meets  with  a 
reverse  on  Otter  Island  from  some  cause,  that  it  will  in  time  become 
regularly  established  as  a  breeder  on  St.  Paul,  and  that,  therefore, 
the  likelihood  of  the  species  surviving  will  be  strengthened. 

During  the  winter  of  1916-17  St.  George  was  visited  by  an  unusual 
number  of  gyrfalcons,  which  preyed  on  the  wrens  and  rosy  finches  to 
such  an  extent  that  they  were  almost  extirpated.  G.  Dallas  Hanna 
states  that  in  May,  1917,  he  found  not  over  six  pairs  of  wrens  during 
a  trip  made  entirely  around  the  island.  Since  then,  however,  as  else- 
where detailed,  the  species  has  become  at  least  fairly  common  again 
and  has  even  spread  to  the  other  main  islands,  previously  unoccupied. 
For  many  years  after  the  discovery  of  this  species  its  nest  and  breed- 
ing habits  were  almost  unknown.  A  nest  and  incomplete  set  of  eggs 
taken  by  a  native  in  June,  1876,  were  forwarded  with  other  specimens 
to  the  Museum  of  Comparative  Zoology  and  were  described  by  Dr. 
J.  A.  Allen  (1877,  p.  82).  The  attempts  of  various  naturalists,  who 
visited  the  islands  subsequently,  to  find  the  nest  of  this  abundant  but 
elusive  species  failed  entirely  until  1918,  when  Dr.  Harold  Heath,  in 
May  and  June,  by  studying  the  habits  of  the  birds  was  able  to  dis- 
cover upward  of  16  nests.     The  number  of  eggs  was  generally  7,  and 


100  NORTH   AMERICAN   FAUNA.  [No.  46 

this  seemed  to  be  the  maximum  number.  In  the  case  of  a  nest  found 
by  Mr.  C.  E.  Crompton  on  St.  George  later  the  same  summer,  a  sec- 
ond set  of  eggs  was  deposited  within  a  week  after  the  first  brood  of 
young  left  the  nest.  The  habit  of  raising  two  broods  is  almost  unique 
among  the  birds  of  this  far  northern  station. 

The  nests  found  by  Heath  were  all  situated  in  the  faces  of  cliffs 
at  elevations  ranging  from  8  to  100  feet.  They  usually  occupied  a 
crevice  between  shattered  rocks,  or  blowholes  in  the  lava,  or  cavities 
beneath  mossy  banks.  In  several  cases  the  sites  had  been  occu- 
pied in  previous  years.  The  riests  were  usually  globular  with  the 
entrance  at  the  side  and  were  composed  of  a  meshwork  of  grass 
and  roots,  often  roofed  with  moss.  The  lining  was  usually  com- 
posed of  fine  rootlets  and  lichen,  mixed  with  feathers  and  fox  hair, 
and  in  some  cases  reindeer  hair.  A  very  full  and  extremely  inter- 
esting account  of  the  nesting  habits  and  homes  of  this  species,  from 
which  the  above  notes  are  gleaned,  was  published  by  Doctor  Heath 
(1920,  p.  49-55). 

Food. — Of  the  11  stomachs  of  Alaska  wrens  available  9  were 
examined  some  time  ago  by  less  discriminating  methods  than  those 
at  present  in  use,  and  it  is  only  possible,  therefore,  to  indicate  the 
nature  of  the  food  in  very  general  terms.  The  sustenance  was  en- 
tirely animal  and  included  the  following  groups :  Amphipods,  24.1 
per  cent;  two-winged  flies  (partly  Borboridae),  24.1  per  cent; 
beetles  (including  ground  and  rove  beetles),  14.3  per  cent;  bugs 
(Hemiptera),  13.2  per  cent;  caterpillars,  12.9  per  cent;  and  Hy- 
menoptera,  11.4  per  cent. 

A  recently  examined  stomach  contained  the  following  items:  Six 
beetles  of  the  sexton-beetle  family  (Lyrosoma  opaca),  12  per  cent; 
rove  beetles  (Olophrum  fuscum  and  2  Liparocephalus  brevipennis) , 
3  per  cent;  3  small  parasitic  wasps  (including  Phygadeuon  sp.  and 
Plesignatlais  sp.),  1  per  cent;  remains  of  dung  flies  (Scatophaga 
sp.)  and  perhaps  other  flies,  74  per  cent;  one  mite  of  an  undescribed 
genus  of  the  family  Gamasidae,  trace;  and  amphipod  remains,  30 
per  cent. 

Another  stomach,  lately  examined,  taken  October  29,  1914,  con- 
tained remains  of  24  or  more  rove  beetles  (Staphylinidae),  70  per 
cent;  4  beach  beetles  {Aegialites  debilis),  19  per  cent;  1  other  beetle, 
1  per  cent ;  and  a  few  flies,  10  per  cent. 

Family  TURDIDAE. 

Hylocichla  minima  aliciae  (Baird).     Gray-cheeked  Thrush. 

A  chance  specimen  of  the  gray-cheeked  thrush  has  been  taken  by 
G.  Dallas  Hanna,  who  recorded  the  incident  as  follows  (1920a.  p. 
254)  :  "A   female  gray-cheeked  thrush   was  collected  on   St.   Paul 


1923.]  BIRDS   OF    THE   PRIBILOF   ISLANDS.  101 

Island  September  9,  1917.  It  was  found  feeding  on  spaded-up 
ground  about  the  Naval  Radio  Station."  The  specimen  is  deposited 
in  the  collection  of  the  U.  S.  National  Museum  and  is  catalogued  as 
No.  255175. 

Food. — The  contents  of  the  stomach  of  this  bird  were  3  blowflies 
(Calliphora  vomitoria)  and  2  dung  flies  (Scatophagidae),  60  per 
cent;  4  ground  beetles  {Pterostichus  sp.),  1  rove  beetle  (Staphylini- 
dae),  and  1  weevil  (Lophalophus  inquinatus) ,  40  per  cent. 

Planesticus  migratorius  propinquus  Ridgway.    Western  Robin. 

Elliott  observed  a  robin  on  the  hill  back  of  the  village  of  St.  Paul 
on  October  15,  1872.  The  same  bird  was  seen  for  several  days,  but 
the  statement  is  made  that  he  did  not  shoot  the  bird  (1874,  unpaged, 
and  1875,  pp.  170,  172).  In  his  later  account  of  the  occurrence  Elli- 
ott states  (perhaps  through  an  error)  that  a  specimen  was  secured 
in  October,  1872.  The  bird  seen  by  Elliott  remained  the  only  rec- 
ord until  1910,  when  Hahn  recorded  1  found  at  the  watch  house  at 
Northeast  Point  on  October  19.  The  bird  had  been  shot  by  a  native 
two  weeks  previously.  Among  the  specimens  which  had  accumu- 
lated at  St.  Paul  Island  and  which  were  brought  in  to  Washington 
in  1914  were  the  head  and  feet  of  a  robin,  unlabeled,  but  probably 
the  remains  of  this  individual. 

The  above  notes  referring  to  birds  not  identified  subspecifically  are 
placed  under  the  present  heading,  since  a  specimen  taken  by  G.  Dallas 
Hanna  on  St.  Paul  Island  September  15,  1919,  and  recorded  by  him 
(1920b,  p.  175)  proves  to  belong  to  this  race.  This  specimen  was  de- 
posited in  the  collection  of  the  California  Academy  of  Sciences. 

Oenantlie  oenanthe  oenanthe   (Linnaeus).15     Wheateak. 

A  specimen  of  the  wheatear,  taken  by  R.  E.  Snodgrass  and  A.  W, 
Greely  on  St.  Paul  Island,  August  29,  1897,  was  first  recorded  by 
Seale  (1898,  p.  139).  and  thus  added  to  the  list  of  Pribilof  birds. 
Grinnell,  reporting  on  the  collection  of  which  this  specimen  formed 
a  part,  comments  on  this  bird  as  follows  (1901,  p.  20)  : 

An  immature  bird  (No.  3486,  Coll.  L.  S.  Jr.  U.).  taken  on  St.  Paul  Island 
August  29,  forms  the  first  record  for  the  Pribilofs  and  fills  in  another  gap  in 
the  known  range  of  this  remarkably  widespread  species. 

A  second  specimen  was  taken  by  G.  Dallas  Hanna  on  St.  Paul 
Island,  September  1,  1917,  and  has  been  recorded  by  Mailliard  and 
Hanna  (1921,  p.  94).  This  is  now  No.  255174,  male,  of  the  collec- 
tion of  the  U.  S.  National  Museum. 

16  Saxicola  oenanthe  oenanthe  of  the  A.  O.  U.  Check  list,  1910. 


MAMMALS  OF  THE  PRIBILOF  ISLANDS. 
By  Edward  A.  Preble. 

Wliile  the  birds  of  the  Pribilofs  are  of  great  popular  interest, 
the  mammals,  though  fewer  in  number  of  species,  are  of  even  greater 
importance.  The  herd  of  fur  seals  is  one  of  the  most  interesting 
and  valuable  aggregations  of  mammals  in  the  world,  and  their  pres- 
ence is,  of  course,  responsible  for  the  settlement  of  the  islands  and 
the  consequent  comparatively  intensive  study  of  its  fauna  and  flora. 
The  foxes  afford  one  of  the  best  examples  known  of  the  selective 
breeding  of  a  wild  animal  carried  on  under  practically  natural  con- 
ditions. In  the  present  treatment  the  accounts  of  these  species  are 
by  no  means  commensurate  in  volume  with  their  importance.  Since, 
however,  they  have  been  the  subject  of  many  special  reports,  it  has 
seemed  unnecessary  to  treat  them  more  than  briefly  here. 

For  convenience  in  citing  references,  the  titles  referring  to  mam- 
mals are  given  together  with  those  relating  to  birds  in  the  Bibliog- 
raphy (pp.  121-128). 

Family  SORICIDAE. 

Sorex  pribilofensis  Merriam.     Pribilof  Shrew. 

The  Pribilof  shrew,  which  is  confined  to  St.  Paul  Island,  was  first 
discovered  by  William  Palmer,  in  1890,  and  a  series  of  16  specimens 
was  collected.  Other  specimens  were  taken  by  Dr.  C.  Hart  Merriam 
in  the  summer  of  1891.  The  species  was  not  named,  however,  until 
Doctor  Merriam  wrote  his  Synopsis  of  American  Shrews  in  1895, 
when  he  formally  characterized  it  (1895b,  p.  87).  A.  G.  Whitney 
collected  one  in  full  winter  pelage,  and  a  small  series  in  the  summer 
of  1914.  The  writer  also  took  a  few  in  the  summer  of  1914,  and 
during  the  next  year  or  two  G.  Dallas  Hanna  collected  a  large  series. 
Altogether,  about  135  specimens  have  been  collected,  nearly  all  of 
which  are  in  the  U.  S.  National  Museum.  Most  of  these  specimens 
were  taken  in  a  partially  marshy  tract,  grown  up  to  rani?:  grasses, 
bordering  a  shallow  pond  between  the  village  and  East  Landing. 
A  few  have  been  collected  at  Northeast  Point. 

Ten  specimens,  measured  in  the  flesh  by  the  writer,  average  in 
measurements :  Total  length,  103.1  mm. ;  tail  vertebrae,  35.8 ;  hind 
foot,  13.2.  This  series  showed  remarkable  uniformity  in  size,  the 
total  length  ranging  only  from  100  to  106  mm.,  and  the  tail  vertebrae 
only  from  34  to  37. 
102 


1923.]  MAMMALS   OF   THE   PRIBILOF   ISLANDS.  103 

Family  URS'IDAE. 

Thalarctos  maritimus  (Pliipps).     Polae  Bear. 

Polar  bears  have  been  transported  to  the  Pribilofs  on  drift  ice  and 
have  landed  on  a  number  of  occasions.  True  (1899,  p.  354)  merely 
mentions  the  fact  but  gives  no  details. 

W.  L.  Halm  found  in  the  St.  Paul  Island  log,  under  date  of  Sep- 
tember 20,  1874,  an  entry  stating  that  a  party  visited  the  cave  on 
Bogoslof  and  brought  back  a  bear  skull  known  to  have  been  there 
since  the  time  of  the  first  occupation  of  the  island.  Other  notes 
from  the  same  source  record  a  white  bear  seen  at  Halfway  Point 
March  28  and  30,  1880,  and  at  Northeast  Point,  March  13,  1884. 
Frederic  A.  Lucas  (1898,  p.  718)  has  recorded  the  skull  of  a  polar 
bear  from  the  Pribilofs,  but  whether  this  is  the  Bogoslof  specimen 
above  noted  is  not  known.  I  have  been  unable  to  find  any  Pribilof 
specimen  in  the  U.  S.  National  Museum  collection. 

The  following  account  of  a  bear  observed  on  St.  George  Island, 
by  G.  Dallas  Hanna  (1914,  p.  218)  summarizes  the  evidence  relating 
to  that  place: 

Ou  February  14,  1914.  a  polar  bear  was  seen  and  shot  at  by  an  Aleut  at 
Zapadni  Rookery,  St.  George  Island.  The  tracks  were  seen  on  shore  in  the 
snow  by  the  writer  and  others.  This  animal  was  formerly  an  abundant  inhab- 
itant of  the  northern  part  of  Bering  Sea,  but  rarely  came  south  of  St.  Matthew 
Island.  If  native  reports  are  to  be  believed,  it  has  been  seen  on  St.  Paul  Island 
on  at  least  three  different  occasions,  but  this  was  many  years  ago.  From  the 
same  source  of  information  comes  the  only  previous  record  of  the  animal  on  St. 
George.  This  was  about  1820,  when  a  bear  came  ashore  from  the  ice  pack  at 
the  village  and  went  westward  to  a  small  pond  where  it  spent  some  time  on  the 
ice.  Thereafter  this  has  been  known  as  Bear  Lake.  The  numerous  tracks 
along  the  beach  in  the  snow  show  that  the  bear  seen  at  Zapadni  had  been 
ashore  also.  During  the  winter  no  drift  ice  had  been  sighted  from  the  island. 
But  this  could  not  have  been  seen  unless  it  had  come  within  10  miles. 

Family  CANIDAE. 

Alopex  pribilof ensis  (Merriam).     Pribilof  Arctic  Fox;  Blue  Fox. 

PI.  VII,  fig.  3. 

The  foxes  of  the  Pribilof  Islands  have  been  described  as  a  separate 
race,  the  long  period  of  isolation  evidently  having  resulted  in  their 
differentiation  from  the  mainland  stock.  On  the  Pribilofs  the  blue 
phase  of  coloration,  which  is  really  an  abnormal  type  and  which 
occurs  more  or  less  rarely  throughout  the  range  of  the  Arctic  foxes, 
at  least  in  America,  has  become  predominant.  The  white  fox  is 
merely  the  winter  condition  of  the  normal  animal,  which  in  summer 
has  tawny  sides  and  brown  back  and  shoulders.  The  so-called  blue 
fox  is  brownish  or  sooty  in  its  summer  coat  and  lighter  in  winter. 

The  animals  usually  mate  in  March  or  early  April,  and  new-born 
young  have  been  found  from  May  17  to  June  6.    Among  22  litters 


104  NORTH   AMERICAN   FAUNA.  [No.  46 

examined,  the  smallest  contained  5  and  the  largest  11  pups.  A  newly 
born  pup  weighed  2|  ounces.  The  adults  vary  greatly  in  weight, 
according  to  sex  and  condition.  Males  taken  in  midwinter  may 
range  from  5£  to  171  pounds,  and  have  reached  as  high  as  20£ 
pounds.  Females  are  recorded  weighing  from  4  to  15  pounds,  and 
in  one  case  21f  pounds.     (Chichester,  1908,  p.  51.) 

Foxes  have  been  found  on  both  St.  Paul  and  St.  George  Islands 
ever  since  they  were  first  discovered,  and  it  seems  that  the  blue 
phase  of  coloration  has  always  been  predominant  there.  Indeed,  the 
early  accounts  of  the  islands  aver  that  at  the  time  of  their  discovery 
only  the  blue  foxes  were  found,  but  that  white  ones  came  (presum- 
ably on  the  ice)  a  few  years  later.  But  since  white  foxes  are  still 
known  to  come  from  time  to  time  in  the  same  way,  it  is  likely  that 
some  of  this  color  were  always  found  there.  From  the  first  efforts 
have  been  made  to  keep  down  the  number  of  the  less  valuable  white 
foxes.  On  St.  Paul,  where  only  steel  traps  are  used,  these  efforts 
have  been  only  partially  successful,  and,  judging  from  the  catches 
of  the  past  few  years,  about  17  per  cent  of  the  foxes  there  are  white. 
On  St.  George,  however,  where  food  is  much  less  abundant,  the 
foxes  can  be  attracted  to  certain  places  and  then  taken  in  large  cage 
traps.  By  this  means,  a  good  proportion  of  all  the  foxes  of  the  island 
being  annually  handled,  no  white  ones  are  allowed  to  live,  and  a 
supply  of  the  best  blue  ones  are  annually  liberated  as  breeders.  The 
natural  result  of  this  selective  method  has  been  to  improve  the  blue 
ones  and  to  restrict  the  normal  phase  to  a  minimum.  Thus  the 
catch  of  white  foxes  on  St  George,  in  spite  of  the  fact  that  all  white 
ones  taken  are  killed,  has  averaged  during  the  past  few  years  con- 
siderably less  than  1  per  cent  of  the  total. 

We  know  little  about  the  numbers  of  foxes  found  on  the  islands  in 
earliest  times.  During  the  period  from  1842  to  1860,  inclusive,  the 
liussi  an- American  Co.  made  on  the  Pribilofs  an  average  annual  catch 
of  1,829  foxes,  more  that  two-thirds  of  which  came  from  St.  George. 
Figures  from  1861  to  1870  are  not  available.  Between  1871  and  1890, 
24,792  skins  were  taken;  20,412  came  from  St.  George.  From  1890, 
the  foxes  diminished  rapidly,  owing,  no  doubt,  to  the  lessened  num- 
ber of  seals  killed,  the  bodies  of  which  had  formed  their  chief  food. 
Special  feeding  was  then  resorted  to.  but  in  spite  of  this,  and  of  some 
close  seasons,  only  11,250  were  taken  from  the  seasons  1890--91,  to 
1913-14,  inclusive,  a  yearly  average  of  only  469.  During  recent  years 
the  numbers  taken  are  again  increasing  as  shown  by  the  table  given 
below. 

Most  of  the  improvements  in  methods  of  dealing  with  the  foxes  on 
the  Pribilofs  are  the  results  of  the  painstaking  work  of  James  Judge, 
who  studied  the  problem  carefully  on  St.  George  for  many  years  and 
published  two  papers  on  the  subject.     (See  Bibliography.)     From 


1923.] 


MAMMALS   OF   THE   PRIBILOF   ISLANDS. 


105 


these  papers  and  from  his  manuscript  notes  are  taken  much  of  the 
data  on  which  the  present  account,  as  well  as  the  one  referred  to 
later,  are  based. 

The  history  of  the  fox  industry  on  the  Pribilofs  was  discussed  in  a 
fairly  comprehensive  manner  by  the  writer  (in  Osgood  et  al.,  1915,  pp. 
105-116).  In  a  table  there  given  the  numbers  taken  between  1890 
and  1914  are  set  forth  in  detail.  In  order  to  bring  the  subject  up  to 
date  the  following  table,  compiled  from  figures  published  by  the  Bu- 
reau of  Fisheries,  and  so  arranged  as  to  be  readily  comparable  with 
the  table  referred  to,  has  been  prepared. 

Number  of  foxes  taken  on  Pribilof  Islands,  1914-1921. 


Year. 

St.  Paul. 

St.  George. 

Total,  Pribilof  Islands. 

Blue. 

173 
211 
150 
90 
119 
156 
123 
138 

White. 

39 
18 
37 
14 
25 
32 
13 
21 

Total. 

Blue. 

White. 

Total. 

Blue. 

236 
420 
567 
692 
667 
902 
1,125 
712 

White. 

40 
20 
39 
19 
30 
36 
14 
21 

Total. 

1914-15 

212 
229 
187 
104 
144 
188 
136 
159 

63 
209 
417 
602 
548 
746 
1,002 
574 

1 

2 
2 
5 
5 

4 
1 

64 
211 
419 
607 
553 
750 
1,003 
574 

276 

1915-16 

440 

1916-17 

606 

1917-1S 

711 

1918-19 

697 

1919-20 

938 

1920-21 

1  139 

1921-22 

733 

Formerly  even  the  best  fox  skins  from  the  Pribilofs  were  worth 
only  a  few  dollars,  but  during  the  period  of  inflated  prices  following 
the  World  War,  due  partly  to  the  growing  scarcity  of  furs,  large 
sums  have  been  realized.  Thus  665  blue  foxes  sold  at  St.  Louis  Sep- 
tember 10,  1919,  brought  an  average  price  of  $195.90.  On  February 
21,  1921,  901  blue  foxes  brought  an  average  of  $88.12,  and  37  white 
foxes  brought  $35  per  skin.  On  September  28,  1921,  1,125  blue  foxes 
were  sold  at  an  average  price  of  $96.83. 

Vulpes  alascensis  Merriam.     Alaska  Red  Fox. 

Red  foxes  are  said  to  be  sometimes  brought  to  the  Pribilofs  on 
the  pack  ice,  but  I  have  no  definite  data.  True  (1899,  p.  354)  states 
that  Mr.  Palmer  reported  in  1890  that  not  more  than  six  or  seven 
had  been  taken  in  twenty  years. 

Family  MUSTELIDAE. 
Latax  lutris  Linnaeus.     Sea  Otteb. 

At  the  time  of  the  discovery  of  the  Pribilof  Islands  sea  otters  were 
found  in  great  numbers,  but  the  cupidity  of  the  Russian  fur  hunters 
soon  caused  their  extirpation.  According  to  Elliott  (1874,  un- 
paged, and  1875,  p.  54)  as  many  as  5,000  were  killed  on  St.  Paul 
Island  during  the  first  year  of  its  occupation  (1787)  ;  after  this  the 
decline  was  rapid.     Veniaminof,  quoted  by  Elliott   (1.  c,  p.  242), 


106  NORTH   AMERICAN   FAUNA.  [No.  46 

states  that  the  animals  became  scarce  there  in  1811,  and  extinct  within 
the  next  30  years.  Elliott  himself,  in  1872  and  1873,  seems  not  to 
have  observed  any,  nor  to  have  added  anything  to  the  history  of 
the  species  on  the  islands. 

W.  L.  Hahn  found  in  the  St.  Paul  Island  log  entries  to  the  fol- 
lowing effect:     A  sea  otter  was  seen  by  fishermen  September  23, 

1889,  the  first  for  several  years.  On  June  1,  1896,  one  which  had 
been  crushed  by  the  ice  was  found  at  Rocky  Point.  On  December  1 
(apparently  of  the  same  year)  one  was  reported  close  inshore  at 
Southwest  Bay.  True  (1899,  p.  353)  mentions  a  skull  found  on  St. 
Paul  by  C.  H.  Townsend  in  1892,  and  the  Rocky  Point  find,  above 
noted.  There  are  two  skulls  in  the  collection  of  the  U.  S.  National 
Museum,  received  years  ago  from  the  Bureau  of  Fisheries,  which 
may  be  these  specimens.  The  Biological  Survey  collection  contains 
two  large  skulls  collected  by  A.  G.  Whitney  on  St.  Paul  Island,  a 
very  much  weathered  one  picked  up  on  lagoon  beach  on  October  9, 
1913,  soon  after  a  violent  storm,  and  one  which  seems  much  more 
recent  found  at  Rocky  Point  on  June  21,  1914. 

Hanna  notes  that  the  St.  George  Island  log  for  July  24,  1892, 
records  the  finding  of  a  dead  sea  otter  by  a  native  at  Zapadni 
Rookery.     The  skin  of  this  animal  was  sold  for  about  $100. 

Family  ODOBENIDAE. 

Odobenus  divergens  (Illiger).     Pacific  Walrus. 

The  walrus  formerly  resorted  to  the  two  main  islands  of  the 
Pribilofs  in  some  numbers  judging  from  the  reports  of  early  his- 
torians and  by  the  remains  which  are  still  found  there.  There  is 
no  indication,  however,  that  the  animal  ever  bred  in  the  vicinity, 
and  most  of  the  visitors  apparently  were  males.  They  must  have 
occurred  most  commonly  in  the  vicinity  of  Cross  Hill,  near  North- 
east Point,  at  that  somewhat  remote  time  when  the  encroaching 
sand  was  closing  up  the  passage  between  the  main  island  and  that 
portion,  topped  by  Hutchinson  Hill,  which  now  constitutes  North- 
east Point,  and  concerning  which  both  tradition  and  topographic 
features  indicate  former  separation.  Considerable  quantities  of 
bones  and  teeth  have  been  dug  out  of  the  sand  dunes  there,  and  re- 
mains are  still  sometimes  found.  The  last  stand  as  a  regular  haul- 
ing place,  however,  was  Walrus  Island,  where  upwards  of  150  in- 
dividuals were  observed  by  Elliott  in  1872.  These  were  all  males, 
but  he  was  afterwards  informed  that  on  one  occasion  a  single  female 
was  seen  there.     The  animals  continued  to  resort  there  as  late  as 

1890,  when,  according  to  True  (1899,  p.  354),  William  Palmer  ob- 
served 8,  "but  killed  none,  and  they  were  reported  all  killed  the 
following  season. " 


1923.]  MAMMALS   OF   THE   PRIBILOF   ISLANDS.  107 

The  following  notes,  taken  from  the  St.  Paul  Island  log  by  W.  L. 
Hahn,  afford  some  indication  of  the  number  killed,  and  the  diminish- 
ing frequency  of  their  appearance  in  recent  years : 

Walrus  Island :  June  10,  1872,  many  seen ;  June  29,  1873,  a  few ;  August  21, 
1873,  3  killed ;  May  22,  1874,  very  few ;  May  23,  1877,  1  taken ;  June  2,  1876,  3 
shot ;  February  26,  1881,  1  seen ;  May  21,  1881,  60  seen  near  the  island ;  June 
8,  1881,  1  killed;  June  14,  1881,  3  seen;  June  22,  1890,  3  killed;  August  24, 
1890,  1  shot ;  June  10,  1893,  1  seen.  St.  Paul  Island :  February  26,  1875,  3  found 
dead  at  Northeast  Point;  May  26,  1875,  1  found  dead  at  Gorbatch  Rookery; 
December  25,  1877,  1  killed  on  Reef  Rookery;  March  23,  1899,  1  found  dead 
near  Rocky  Point ;  April  22,  1899,  1  found  dead  at  Northeast  Point ;  March  17, 
1900,  1  found  dead  at  Northeast  Point ;  August  4,  1904,  2  found  dead  at  Rocky 
Point ;  November  4,  1904,  1  seen  in  water  near  East  Landing ;  June  23,  1907,  1 
found  dead  at  Northeast  Point. 

During  my  visit  to  the  Pribilofs,  in  1914,  I  saw  a  few  skulls  near 
Northeast  Point,  one  on  Otter  Island,  and  a  number  among  the 
rocks  on  Walrus  Island.  Later  in  the  same  year  Hanna  (1914,  p. 
218)  published  the  following  note : 

During  the  month  of  March,  1914,  walruses  were  sighted  three  different  times 
swimming  along  just  offshore.  They  were  at  no  very  remote  time  abundant 
on  St.  George,  as  the  skulls  may  yet  be  seen  about  the  beaches.  Five  dead  ones 
floated  in  at  Garden  Cove  about  1907. 

In  July,  1918,  Hanna  observed  a  young  male  walrus  asleep  on  a 
surf-washed  rock  at  North  Rookery,  St.  George.  The  animal  was 
not  at  all  wary,  and  Hanna  was  able  to  photograph  it  at  a  distance 
of  a  few  feet,  and  to  retire  without  disturbing  it. 

Family  OTARIIDAE. 

Eumetopias  jubata  (Schreber).     Steller  Sea  Lion. 

Only  a  few  hundred  sea  lions,  found  in  two  small  breeding  colonies, 
are  all  that  now  remain  of  the  many  thousands  which  until  com- 
paratively recent  years  inhabited  the  islands.  Elliott  (1875,  p.  153) 
estimated  that  in  1872  there  were  from  20,000  to  25,000  about  St. 
Paul  Island,  and  7,000  or  8,000  on  St.  George.  They  then  occupied  a 
considerable  area  at  Northeast  Point,  St.  Paul,  and  several  loca- 
tions on  St.  George. 

Elliott  (1.  c,  p.  71)  states  that  a  few  formerly  bred  on  Walrus 
Island,  but  apparently  this  colony  no  longer  exists,  although  a  few 
of  the  animals  sometimes  haul  out  on  rocks  there,  as  well  as  at  many 
points  on  all  the  islands.  According  to  True  (1899,  p.  351)  at  least 
one  harem  persisted  on  Walrus  Island  as  late  as  1890.  He  states  that 
on  St.  George  the  principal  stations  were  near  Tolstoi  Point  and 
East  Rookery.  Sea  lions  are  believed  to  breed  still  at  these  places, 
and  also  at  Garden  Cove. 

Their  breeding  habits  are  similar,  in  general,  to  those  of  the  fur 
seal.    The  harems  are  smaller,  usually  numbering  only  10  or  12  cows 


108  NORTH   AMERICAN   FAUNA.  [No.  46 

to  a  bull.  The  young  are  born  about  the  middle  of  June.  The 
animals  are  much  more  timid  than  the  seals,  and  therefore  their 
intimate  habits  are  more  difficult  of  observation.  The  bulls  fight 
among  themselves  with  great  ferocity.  With  the  exception  of  a  few 
old  bulls  all  the  sea  lions  are  said  to  leave  the  islands  in  winter. 
They  return  in  March,  April,  and  May,  the  old  bulls  first. 

Like  the  fur  seal,  the  difference  in  size  of  the  male  and  female  sea 
lion  is  very  great.  The  breeding  males  have  been  estimated  to  weigh 
from  1,500  to  2,000  pounds,  while  a  cow  may  weigh  up  to  500  or  600. 
I  have  no  data  on  the  weight  of  the  new-born  young.  Young  ap- 
proximately 3  weeks  old,  observed  by  the  writer  at  the  breeding 
rookery  at  Northeast  Point,  on  June  27,  1914,  were  estimated  to  weigh 
about  35  pounds. 

The  animals  were  formerly  of  great  economic  importance  to  the 
natives,  being  used  as  food,  clothing,  and  in  the  construction  of 
their  skin  boats.  They  were  formerly  killed  in  large  numbers,  not 
only  for  use  on  the  Pribilofs  but  for  exportation  to  Unalaska, 
Kodiak,  and  other  points.  Between  1870  and  1890  from  30  to  500 
were  killed  annually.  The  animals  were  formerly  gathered  in  small 
herds,  called  pods,  by  a  party  of  natives  stealing  at  night  between 
a  sleeping  herd  and  the  water,  and  by  suddenly  rising  with  a  clamor 
of  shouting  and  firing  of  pistols,  getting  as  many  as  possible  of  the 
startled  animals  to  start  inland  instead  of  toward  the  water.  When 
once  headed  away  from  the  sea  they  were  easy  of  control,  and  when, 
by  a  number  of  such  sorties,  a  sufficient  number  had  been  collected, 
they  were  started  to  the  village.  This  was  to  save  the  labor  of  trans- 
porting the  skins  and  meat  by  teams.  The  journey  of  11  or  12  miles 
took  from  5  days  to  3  weeks  according  to  the  weather,  for  the  huge 
beasts  could  travel  but  slowly,  and  could  not  be  forced  in  warm 
weather  without  great  mortality.  On  arrival  at  the  village  the 
entire  herd  were  killed  with  guns  and  lances.  Since  about  1882  the 
lessened  number  of  the  animals  has  made  driving  impracticable, 
and  the  killing  has  been  done  at  Northeast  Point.  The  topographic 
conditions  on  St.  George  are  such  that  driving  has  never  been  prac- 
ticable there. 

During  recent  years  they  are  seldom  killed  except  for  their  skins, 
which  are  still  used  to  cover  the  framework  of  the  large  bidarras, 
used  principally  to  land  the  cargoes  from  the  supply  steamers, 
which  must  anchor  some  distance  from  the  wharves.  For  this  pur- 
pose the  young  bulls  only  are  killed,  as  the  hides  of  the  old  bulls 
are  unsuitable.  The  skins  of  12  animals  are  needed  for  covering  one 
boat  and  this  cover  lasts  four  or  five  years. 
Callorhinus  alascanus  Jordan  and  Clark.     Pribilof  Fur  Seal. 

The  fur  seal,  which  seems  to  have  been  directly  responsible  for  the 
discovery  of  the  Pribilof  Islands,  and  which  has  been  the  dominant 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey. 


PLATE    VII. 


Fig.   I. — Bull  Seals  in  Spring,  on  Their  Rookery  Stations. 

The  males  arrive  early  in  spring  and  select  a  home  site,  where  they  are  joined  later  by  the  females. 
Photograph  by  A.  ChristoSersen,  on  Reef  Rookery, 'St.  Paul  Island. 


Fig.  2. — Bull  Seals  in  August,  on  the  Grassy  Levels. 

The  old  males,  which  arrive  in  spring  fat  and  full  of  energy,  are  much  reduced  in  flesh  and  spirits  after 
three  months  of  fasting.    Photograph  by  G.  Dallas  Hanna,  St.  Paul  Island. 


Fig.  3. — Blue  Fox  (Alopex  pribilofensis). 

The  blue  phase  of  coloration  practically  attains  perfection  as  regards  quality 
of  fur  and  freedom  from  infe-ior  strains  on  St.  George  Island.  Photo- 
graph by  G.  Dallas  Hanna,  November,  1916. 


1923.]  MAMMALS   OF   THE   PRIBILOF   ISLANDS.  109 

factor  in  its  subsequent  history,  is  one  of  the  most  interesting  animals 
known.  It  resorts  to  these  islands  only  during  the  breeding  season, 
and  spends  the  remainder  of  the  year  at  sea,  not  being  known  to 
land  at  any  other  place.  Its  migration  carries  it  as  far  south  as 
the  latitude  of  southern  California,  the  females  going  farthest  and 
the  old  bulls  wintering  mainly  south  of  the  Aleutians  or  in  the  Gulf 
of  Alaska. 

On  the  approach  of  the  breeding  season  the  old  bulls  return  to 
the  islands,  the  earliest  arriving  about  the  first  of  May,  and  each 
one  takes  a  position  on  the  rookery  ground,  often  in  the  spot  occupied 
the  previous  year.  (PI.  VII,  fig.  1.)  From  the  time  he  lands  until 
the  end  of  the  breeding  season,  late  in  July  or  early  in  August,  the 
bull  never  leaves  his  station  and  consequently  takes  no  food.  Shortly 
after  the  first  of  June  the  females  begin  to  arrive,  each  one  impelled 
by  her  pregnant  condition  to  seek  a  place  to  bring  forth  her  pup. 
On  arrival  the  females  land  and  join  some  male  which  is  gathering 
a  harem.  (See  frontispiece.)  Before  many  days,  sometimes  almost 
at  once,  the  female  is  delivered  of  her  pup,  and  within  a  few  days  is 
again  impregnated.  She  is  then  allowed  to  depart  in  her  quest  for 
food,  which  she  must  have  in  abundance  in  order  to  nourish  her  pup. 
She  feeds  at  sea,  going  from  50  to  100  miles,  and  after  gorging  on 
fish  remains  in  the  water  until  digestion  has  taken  place.  She  then 
returns  to  the  land,  picks  out  her  own  pup  from  among  the  thousands 
that  swarm  over  the  rookeries,  and  nurses  it  until  she  is  impelled  to 
repeat  her  trip  for  food. 

The  harems,  comprising  from  a  few  to  as  many  as  100  cows  and 
each  attended  by  a  single  bull,  are  crowded  together  until  practically 
all  the  rookery  ground  is  occupied.  As  the  young  grow  larger  they 
wander  farther  afield  during  the  absence  of  their  mothers,  but  the 
female  always  finds  her  own  pup,  and  will  nurse  no  other. 

By  early  August  practically  all  the  cows  have  given  birth,  and 
the  old  bulls,  exhausted  by  their  long  fast  and  their  protracted 
harem  service,  after  a  few  days'  rest,  go  to  sea  to  feed  and  recuper- 
ate (PI.  VII,  Fig.  2).  The  females,  however,  continue  to  nurse 
their  young,  which  learn  to  swim  during  the  late  summer,  until  No- 
vember, when  they,  too,  leaA^e  the  islands. 

The  sexes  differ  greatly  in  size,  the  breeding  bulls  weighing  from 
400  to  500  pounds,  while  the  breeding  cows  vary  from  50  to  60  to 
about  100  pounds,  according  to  age  and  condition.  The  pups  when 
born  weigh  from  10  to  12  pounds.  Just  before  leaving  the  islands 
in  November  the  pups  have  been  found  to  weigh  from  about  25  to  50 
pounds;  when  they  return,  in  the  following  August,  or  when  a  little 
over  a  year  old,  they  weigh  from  33  to  40  pounds,  having  become 
108731°— 23 8 


110  NORTH   AMERICAN   FAUNA.  [No.  46 

longer  but  much  less  fat  since  being  weaned  and  living  the  life  of 
the  adults. 

During  the  early  years  of  the  occupancy  of  the  group,  the  slaugh- 
ter of  the  animals  for  their  valuable  skins  was  pursued  in  a  very 
wasteful  manner,  both  males  and  females  being  killed  in  such  num- 
bers as  to  exceed  the  hunters'  resources  for  the  proper  preservation 
of  the  catch,  and  to  glut  the  market.  The  imperfect  records  of  these 
early  days  indicate  that  more  than  1,800,000  seals  were  killed  be- 
tween 1786  and  1834,  and  the  /herd  was  seriously  depleted.  From 
1835  to  1867  the  females  were  spared,  and  the  killing  was  otherwise 
restricted  so  that  the  herd  gradually  increased.  At  the  time  of  the 
purchase  of  Alaska  by  the  United  States  in  1867  various  estimates 
placed  the  number  of  fur  seals  between  2,000,000  and  5,000,000 
animals. 

In  1870  the  privilege  of  sealing  on  the  islands  was  leased  to  the 
Alaska  Commercial  Co.  for  a  term  of  20  years.  During  this  period 
1,977,377  skins  were  taken  on  the  islands.  At  the  expiration  of  this 
lease  the  North  American  Commercial  Co.  was  awarded  the  seal- 
ing privilege  for  a  further  period  of  20  years,  and  342,651  skins  were 
taken. 

About  1890  the  effect  of  killing  seals  at  sea,  a  large  percentage 
being  females,  the  death  of  which  caused  the  loss  of  unborn  or 
young  pups,  began  to  be  recognized  as  an  intolerable  waste  of 
valuable  life.  As  a  result,  by  agreement  between  the  countries  in- 
terested, pelagic  sealing  was  curtailed,  and  finally,  in  1911,  by  a 
treaty  between  the  United  States,  Great  Britain,  Russia,  and  Japan, 
it  was  abolished  for  a  period  of  15  years.  Under  its  provisions  the 
United  States  and  Russia,  as  guardians  of  the  seal  herds,  agreed  to 
pay  Great  Britain  and  Japan  a  percentage  of  the  value  of  the  seals 
taken  on  land. 

In  1912,  owing  to  the  fear  that  the  seal  herd  had  become  depleted 
to  the  danger  point,  all  killing  was  forbidden  by  law  for  a  period 
of  five  years,  excepting  such  numbers  as  were  needed  by  the  natives 
for  food  purposes.  At  this  time,  more  intensive  studies  of  the  herd 
were  inaugurated,  and  as  a  result  it  became  possible  to  estimate  the 
numbers  of  the  animals  with  more  accuracy  than  had  before  been 
possible.  One  of  the  important  features  was  the  counting  each  year 
of  the  newly  born  pups,  which  afforded  an  index  to  the  increase  of 
the  herd.  This,  and  related  matters,  was  rather  fully  discussed  by 
Osgood,  Parker,  and  Preble  (1915,  pp.  39,  44,  etc.),  the  data  being 
brought  up  to  1914. 

Since  1914  a  summer  survey  of  the  breeding  herd  has  been  made 
annually.  The  following  table,  giving  the  number  of  pups  born 
since  1912,  and  the  estimated  size  of  the  entire  herd,  will  serve  to 


1923.] 


MAMMALS   OF   THE   PRIBILOF   ISLANDS. 


Ill 


show  the  steady  increase  in  numbers.  The  fibres  have  been  taken 
from  publications  issued  by  the  Bureau  of  Fisheries.  Those  for 
the  pups  up  to  1916,  inclusive,  are  actual  counts;  while  from  1917  on 
they  are  based  on  counts  for  some  of  the  rookeries,  the  others  being 
estimated  by  counting  the  harems  and  applying  the  figures  repre- 
senting the  average  harem. 

Count  of  seal  pups,  and  estimated  numbers  of  entire  herd. 


Year. 

Pups. 

Seals  of 
all  ages. 

Year. 

Pups. 

Seals  of 
all  ages. 

1912 

81,984 
92, 268 

93. 250 
103, 527 
116,977 

215, 73S 
268, 305 
294,687 
363,872 
417, 281 

1917 

128, 024 
143, 005 
157, 572 
167, 527 
176, 665 

468, 692 
496  600 

1913 

1918 

1911 

1919 

524  235 

1915 

1920 

552, 718 
581  457 

1916 

1921 

Family  PHOCIDAE. 

Phoca  richardii  pribilofensis  Allen.     Pribilof  Harbor  Seal. 

The  hair  seal  of  the  Pribilofs,  which  has  been  separated  as  a  race 
of  Phoca  richai'dii,  the  harbor  seal  of  the  northwest  coast,  is  found 
in  comparatively  small  numbers  mainly  on  the  larger  islands.  The 
animals  have  their  young  on  the  bare  sea-washed  rocks,  breeding  on 
St.  Paul  mainly  or  entirely  on  the  unfrequented  north  shore,  though 
they  frequently  haul  out  on  the  rocks  at  various  other  places.  T 
have  no  definite  data  on  the  breeding  of  the  species  about  St.  George, 
but  the  animal  is  frequently  seen  there,  and  probably  breeds  about  the 
less  accessible  parts  of  the  shore.  The  young,  which  are  said  by  El- 
liott (1884,  p.  28)  to  weigh  from  3  to  7  pounds  when  born,  are  white 
in  color,  but  change  in  a  few  months  to  the  color  of  the  old  ones, 
Avhich  is  usually  steel  gray,  blotched  and  mottled  with  dark  brown. 

Hahn  collected  notes  from  the  St.  Paul  Island  log,  showing  that 
hair  seals  were  taken  nearly  every  year  by  the  natives.  Under  date 
of  May  30, 1910,  about  50  are  stated  to  have  been  seen  on  Otter  Island. 
On  the  only  occasion  when  we  visited  that  place  in  1914,  on  June  27, 
we  saw  about  a  dozen  at  the  edge  of  the  surf  near  the  landing  place. 

The  flesh  of  this  seal  is  very  much  relished  by  the  natives,  and  the 
few  that  are  taken  are  shot  mainly  for  this  reason. 

True  (1899,  p.  351)  noted  a  few  solitary  examples  at  various  points 
about  St.  Paul  in  1895  and  states  that  three  young  ones  were  brought 
to  the  village. 

Phoca  fasciata  Zimmermann.     Ribbon  Seal. 

This  species  was  recorded  by  True  (1899,  p.  351)  as  follows: 

A  young  female  of  this  species  was  taken  by  one  of  the  sealers  84  miles  west 
of  St.  Paul  in  August,  1896,  so  that  it  may  be  regarded  as  a  probable  occasional, 
visitant  to  the  Pribilofs, 


112  NORTH   AMERICAN    FAUNA.  [No.  46 

Erig-nathus  barbatus  nauticus  (Pallas).     Pacific  Bearded  Seal. 

The  bearded  seal  has  only  recently  been  added  to  the  list  of  Pribi- 
lof  mammals.  G.  Dallas  Hanna  (1921b,  p.  126)  records  a  specimen 
taken  at  St.  George  Island  in  the  winter  of  1917-18  by  C.  E. 
Crompton. 

In  a  recent  note  Hanna  gives  more  data  regarding  this  species. 
The  specimen  referred  to  was  shot  at  East  Rookery  by  a  native,  on 
January  29,  1918.  The  animal  was  in  advanced  age,  judging  by  the 
fact  that  it  had  lost  all  its  teeth7;  it  was  fat,  however,  indicating  that 
it  had  been  able  to  capture  sufficient  food.  Its  sex  was  not  recorded. 
Among  the  detailed  measurements  made  from  the  fresh  specimen 
the  following  may  be  noted:  Total  length  (nose  to  tip  of  tail),  93 
inches;  tail,  7  inches;  hind  foot,  174;  girth  behind  fore  flippers,  60. 

The  seals  referred  to  by  a  reliable  St.  George  native,  who  reported 
to  Hanna  that  several  very  large  hair  seals,  8  to  10  feet  long,  were 
found  on  the  beaches  in  1900,  a  record  year  for  pack  ice,  were  most 
probably  of  this  species. 

Family   MURIDAE. 

Lemmus  nigripes  True.     Pribilof  Lemming. 

The  Pribilof  lemming,  which  is  found  only  on  St.  George  Island, 
was  first  reported  by  Elliott  (1874,  unpaged;  1875,  p.  72),  under  the 
name  Myodes  ohensis,  but  Merriam  in  1891  seems  to  have  been  the 
first  naturalist  to  collect  specimens.  In  1892  or  1893  two  specimens 
were  taken  by  C.  H.  Townsend.  One  of  these,  No.  59152,  U.  S. 
National  Museum,  male,  was  made  the  type  of  the  species,  which 
was  formally  characterized  by  Frederick  W.  True  (1894,  p.  2). 

In  recent  years  Hanna  has  taken  a  series,  and  in  the  summer  of 
1914  the  writer  collected  a  few.  Two  adult  males  then  taken  meas- 
ured, respectively :  Total  length,  152  mm. ;  tail  vertebrae,  21 ;  hind 
foot,  20;  and  154,  19,  23.  Two  adult,  though  not  fully  grown,  fe- 
males measured  140,  18,  21,  and  136,  18,  21.  A  female  measuring 
128,  17,  21,  collected  on  July  23,  contained  three  small  embryos. 

As  in  other  regions,  the  lemmings  of  St.  George  suffer  consider- 
able fluctuations  in  numbers.  Elliott  reported  them  very  abundant 
in  1873;  Palmer  in  1890,  and  True  in  1895,  1896,  and  1897,  reported 
them  scarce.  They  were  rather  common  in  1913  and  1914.  Their 
periodical  scarcity  has  been  attributed  to  the  foxes,  but  G.  Dallas 
Hanna  states  that  he  has  found  no  evidence  that  these  animals  prey 
on  the  lemmings.  It  would  be  very  remarkable,  however,  if  a  fox 
did  not  occasionally  snap  up  such  a  tender  morsel. 

True  (1899,  p.  346),  on  information  furnished  by  Palmer,  stated 
that  a  few  had  been  introduced  on  St.  Paul  from  St.  George,  and  in 
1911  Hahn  was  assured  by  a  native  chief  that  this  had  taken  place 
on  two  occasions,  the  last  time  about  15  years  before,  but  that  he  did 
not  know  of  any  having  been  seen  afterwards. 


1923.]  MAMMALS   OF   THE   PPJBILOF   ISLANDS.  113 

Fiber  zibethicus  zalophus  Hollister.     Alaska  Peninsula  Muskrat. 

The  introduction  of  muskrats  on  the  Pribilofs  was  advocated  some 
years  ago,  and  has  once  been  attempted,  but  was  not  successful. 
Seven  individuals  were  captured  near  Nushagak  in  the  summer  of 
1913  by  G.  Dallas  Hanna  for  planting  on  St.  George  Island.  During 
transit  to  the  islands  the  animals  preyed  on  each  other  until  only  the 
strongest  was  left.  This  was  liberated  in  a  pond  near  the  village, 
but  is  not  known  to  have  survived  the  winter. 

With  a  view  to  introducing  these  animals  most  of  the  ponds  on 
St.  Paul  Island  were  surveyed  by  W.  L.  Halm,  but  were  found  to  be 
too  shallow.  Among  the  ponds  not  critically  examined,  one  in  par- 
ticular, Antone  Lake,  seems  to  the  writer  to  be  suitable,  at  least  as 
far  as  its  physical  characteristics  are  concerned.  The  introduction 
of  the  animals,  however,  is  not  advocated  by  the  writer,  since  it  is 
not  believed  that  they  would  be  of  any  importance  as  food  for 
the  foxes,  and  it  is  unlikely  that  the  comparatively  small  number 
that  could  live  in  the  one  or  two  ponds  of  possible  suitability  would 
prove  economically  profitable. 

Mus  musculus  Linnaeus.    House  Mouse. 

Elliott,  from  observations  made  in  1872-73,  say|,  that  mice  had 
been  brought  to  the  islands  in  ships'  cargoes  long  before  and  were 
a  great  pest  (1875,  p.  73).  True,  from  observations  made  in  1895, 
states  that  the  animal  was  extremely  abundant  about  the  village  of 
St.  Paul,  and  had  been  noted  by  various  observers  on  St.  George 
(1899,  p.  348).  G.  Dallas  Hanna  thinks  that  there  are  now  none 
on  St.  George,  but  there  seems  to  be  no  good  reason  for  this  condi- 
tion, unless  the  pests  are  kept  from  increasing  there  by  the  greater 
number  of  foxes.  On  St-  Paul,  however,  they  were  abundant  in  1914, 
not  only  about  the  village,  but  were  taken  in  my  traps  set  for  shrews 
at  some  distance  from  the  buildings,  and  were  found  even  about  the 
salt  house  at  Northeast  Point. 
Rattus  norvegicus  (Erxleben).    Norway  Rat. 

Elliott  (1875,  p.  73)  states  that  at  the  time  of  his  observations  the 
islands  were  free  from  rats.    True  (1899,  p.  348)  reports  as  follows: 

I  saw  nothing  of  rats  while  on  the  islands  in  1S95,  but  observe  that  Mr. 
Palmer  has  noted  [1890]  that  they  arrive  occasionally  in  ships,  but  do  not  breed. 

While  on  the  islands  in  1914  I  saw  no  evidence  of  rats. 

Family  SCIURIDAB. 

Citellus  plesius  ablusus  Osgood.    Nushagak  Ground  Squirrel. 

Ground  squirrels  have  been  liberated  on  the  Pribilofs  with  a  view 
to  furnishing  an  additional  source  of  food  for  the  foxes.  Preble 
(in  Osgood  et  al.,  1915,  p.  129)  has  summarized  the  matter  as  follows: 

The  introduction  of  ground  squirrels  has  been  attempted  on  two  occasions, 
but  neither  has  proven  a  success.     In  1899  some  were  brought  from  Unalaska 


114  NORTH   AMERICAN   FAUNA.  [No.  46 

and  liberated  on  St.  Paul,  near  the  village.     Tbeir  disappearance  has  been  at- 
tributed to  cats,  but  whether  they  were  eaten  by  cats  or  foxes  is  immaterial. 

In  the  summer  of  1913  the  assignment  of  G.  Dallas  Hanna  for  work  on  St. 
George  Island  afforded  an  opportunity  to  make  another  attempt,  and  22  ground 
squirrels,  including  both  sexes  and  different  ages,  were  captured  at  Nushagak. 
Of  these,  four  died  from  natural  causes  before  their  journey  was  begun.  Various 
circumstances  made  it  impracticable  to  provide  small  cages  for  the  animals,  and 
they  were  shipped  in  a  single  large  crate.  Although  plentifully  supplied  with 
green  food,  they  preyed  on  each  other,  and  while  this  tendency  was  overcome 
to  some  extent  by  supplying  them  with  meat,  the  stock  of  18  had  been  reduced 
to  5  before  they  reached  their  destination.  These  5,  an  adult  female  and  4 
young,  including  both  sexes,  were  liberated  near  the  village  on  St.  George  Island 
in  August.  At  least  two  survived  the  winter,  and  were  seen  on  several  occa- 
sions in  early  May,  1914.  They  were  not  known  to  have  been  observed  later ; 
during  our  visit  in  early  August  none  were  seen,  and  a  careful  search  disclosed 
no  positive  evidence  of  their  presence.  It  is  doubtful  if  any  survived  the  sum- 
mer, and  in  view  of  the  fact  that  numbers  of  foxes  continually  ranged  in  the 
vicinity  of  the  spot  where  they  had  been  observed,  the  destruction  of  the  squir- 
rels would  seem  to  be  inevitable. 

Family  CERVIDAB. 

Eangifer  tarandus  (Linnaeus).    Reindeer. 

Beindeer  wete  introduced  on  the  Pribilofs  in  the  summer  of  1911, 
when  a  herd  of  40  animals  was  brought  from  Unalakleet.  Twenty- 
one  cows  and  four  bulls  were  landed  on  St.  Paul;  one  of  the  bulls 
was  injured  in  landing,  and  died  soon  after.  Of  the  15  landed  on  St. 
George,  3  were  males,  1  adult  and  2  yearlings;  the  adult  bull  met 
with  some  accident  a  few  days  later  and  disappeared.  During  the 
following  summer  18  fawns  were  born  on  St.  Paul,  and  11  on  St. 
George.  The  herds  have  since  continued  to  increase,  and  now  num- 
ber over  400  animals.  Annually  since  1917  a  number  of  the  surplus 
bulls  have  been  killed  for  food,  and  have  furnished  a  welcome  and 
substantial  addition  to  the  tables  of  officials  and  natives. 

Upon  their  introduction,  the  animals  sought  out  the  least  fre- 
quented and  most  favorable  parts  of  the  islands,  over  which  they 
roam  at  will.  The  lichens  on  which  they  mainly  subsist  in  winter 
grow  over  considerable  tracts,  and  a  recent  survey  of  the  situation 
shows  that  large  areas  still  remain  practically  untouched,  so  that 
even  considering  the  comparatively  slow  growth  of  these  plants,  no 
apprehension  need  be  felt  at  present  that  the  herd  is  becoming  too 
large  for  its  natural  food  supply. 

On  each  island  a  corral  has  been  built  into  which  the  respective 
herds  have  been  driven  from  time  to  time  for  examination.  As  the 
herds  have  increased  and  the  commercial  taking  of  seal  skins  has 
been  resumed,  less  attention  could  be  devoted  to  the  reindeer,  be- 
cause of  the  increased  demand  on  the  time  of  the  able-bodied  natives, 
and  in  consequence  the  animals  have  become  somewhat  wild  and 
unmanageable.     Since,  however,  they  are  steadily  increasing,  cost 


1923.] 


MAMMALS   OF   THE   PRIBILOF   ISLANDS. 


115 


nothing  for  maintenance,  and  are  more  and  more  valuable  as  a  source 
of  food,  the  experiment  may  be  considered  highly  successful. 

The  rate  of  growth  of  the  two  herds  since  their  introduction,  under 
practically  natural  conditions  and  without  special  care,  is  of  in- 
terest. 

Grouth  of  reindeer  herd  on  Pribilof  Islands. 


Year. 

St.  Paul. 

St.  George. 

Total. 

1911 

25 

40 

'55 

75 

92 

111 

144 

»158 

6  164 

6  192 

8  250 

15 

25 

40 

58 

62 

85 

2  93 

U14 

6  123 

U25 

»160 

40 

1912 

65 

1913 

95 

1914 

133 

1915 

154 

1916 

196 

1917 

237 

1918 

272 

1919 

287 

1920 .' 

317 

1921 

1410 

1  Approximate. 

2  Excluding  3  killed. 
>  Excluding  2  killed. 


<  Excluding  18  killed. 
6  Excluding  14  killed. 
6  Excluding  22  killed. 

Family  BALAENIDAE. 


i  Excluding  31  killed, 
s  Excluding  32  killed. 
'■  Excluding  19  killed 


Balaena  mysticetus  Linnaeus.     Bowhead  Whale. 

In  a  number  of  instances  whales  have  drifted  ashore  on  the  Prib- 
ilofs,  and  in  some  cases  they  have  been  identified.  St.  Paul  has 
apparently  received  the  larger  share,  a  natural  result  of  its  longer 
and  more  easily  observed  coast  line.  Hahn  collected  notes  from  the 
St.  Paul  Island  log  detailing  some  of  these  occurrences.  On  Jan- 
uary 1,  1890,  a  large  right  whale  was  discovered  at  East  Landing, 
St.  Paul.  The  head  was  gone ;  the  body  was  more  than  60  feet  long. 
Another  carcass  came  ashore  at  Zoltoi,  near  the  village,  on  August 
31  of  the  same  year.  From  the  same  source  Hahn  learned  that  other 
whales  (species  not  stated)  were  stranded  as  follows:  October  18, 
1886,  a  whale  56  feet  long,  bearing  a  harpoon,  drifted  into  English 
Bay.  Others  were  recorded  as  coming  ashore  on  September  13, 
1892;  March  27,  1903;  and  August  9,  1906. 

Dr.  Frederick  W.  True,  himself  a  student  of  this  group,  examined 
the  bones  of  a  number  of  individuals  of  this  species,  and  made  meas- 
urements of  the  right  mandible  of  one  found  on  the  beach  north  of 
Lukanin  (1899,  p.  352).  He  mentions  also  a  bowhead  whale  which 
came  ashore  on  St.  George  in  1889,  from  which  the  natives  secured 
about  1,500  pounds  of  whalebone.  The  remains  of  others  found  by 
True  near  East  Landing  and  at  Zoltoi  were  probably  those  of  indi- 
viduals above  mentioned,  records  of  which  were  found  by  Hahn. 

Scammon  says  (1874,  p.  68)  :  "The  last  seen  of  them  in  high  lati- 
tudes, by  whalemen,  is  on  their  return  from  the  Arctic  Ocean, 
when  they  are  found  in  the  vicinity  of  St.  Paul's  Island,  Behring 
Sea,  in  the  month  of  October,  and  these  are  usually  very  large." 


116  NOKTH   AMERICAN   FAUNA.  [No.  46 

Family  BALAENOPTERIDAE. 

Balaenoptera  velifera  (Cope).     Large  Finback  Whale. 

True  summarized  the  result  of  his  observations  on  the  large  fin- 
back whale  as  follows  (1899,  p.  352)  : 

The  hind  part  of  a  skull  of  a  large  finback,  which  may  be  this  species,  was 
found  by  Mr.  Prentiss  and  myself  on  the  shore  of  the  lagoon  in  1S95.  The 
greatest  breadth  across  the  temporals  was  6.35  feet.  The  height  of  the  occip- 
ital from  the  upper  margin  of  the  foramen  magnum  to  the  nasals  was  3.15 
feet. 

I  saw  nothing  of  these  whales  about  the  Pribilof  Islands  in  the  summer  of 
1S95,  but  when  returning  homeward  observed  them  in  large  numbers  on 
September  3  along  the  south  coast  of  Kadiak.  Mr.  Palmer  remarked  in  1890 : 
"Not  common  about  the  islands  in  summer,  but  a  number  were  seen  after  leav- 
ing St.  George  for  the  south  on  August  11.  A  dead  one  was  stripped  of  its 
'  bone '  on  St.  Paul  last  winter." 

Hahn  found  in  the  St.  Paul  Island  log  a  record  of  a  large  finback 
which  drifted  ashore  at  the  north  end  of  St.  Paul  Island  on  Feb- 
ruary 13,  1884. 

Balaenoptera  davidsoni  Scammon.     Davidson  Lesser  Rorqual. 

True  (1899,  p.  352)  gives  the  following  brief  account  of  the  ror- 
qual, which  comprises  all  the  data  available : 

The  bones  of  a  small  finback,  belonging  without  doubt  to  the  species,  were 
found  at  Rocky  Point,  St.  Paul.  There  were  27  in  all — the  seventh  cervical, 
11  dorsals,  and  15  lumbars  and  caudals. 

Megaptera  versabilis  Cope.     Pacific  Humpbacked  Whale. 

According  to  a  note  taken  by  Hahn  from  the  St.  Paul  Island  log, 
a  female  humpback,  about  two-thirds  grown,  came  ashore  at  Zoltoi 
beach  on  September  27,  1876. 

Family  PHYSETERIDAE. 

Physeter  macrocephalus  Linnaeus.    Sperm  Whale. 

The  only  record  of  the  sperm  whale  on  the  Pribilofs  seems  to  be 
that  of  G.  Dallas  Hanna  (1914,  p.  218),  who  thus  details  the 
occurrence : 

A  sperm  whale  or  cachelot  came  ashore  at  Zapadni  Rookery  April  14,  1914. 
It  was  a  male  47  feet  long  and  had  probably  been  dead  a  week.  About  5  tons 
of  blubber  were  saved  for  fox  food  when  the  head  and  carcass  floated  away. 
Although  other  species  of  whales  are  abundant  about  the  island,  the  cachelot 
had  never  been  seen  before  by  any  of  the  Aleuts. 

In  a  recent  note  Hanna  mentions  a  female  sperm  whale  which 
came  ashore  in  an  advanced  stage  of  decomposition  near  Kitovi 
Rookery,  St.  Paul  Island,  in  the  summer  of  1919. 


1923.]  MAMMALS    OF   THE   PEIBILOF   ISLANDS.  117 

Family   ZIPHIIDAE. 
Berardius  bairdii  Stejneger.     Pacific  Beaked  "Whale. 

The  first  notice  of  the  occurrence  of  this  species  is  apparently  that 

of  True,  who  recorded  it  as  follows : 

Two  large  beaked  whales  were  found  on  the  coast  of  St.  George  Island, 
Pribilof  Group,  Alaska,  in  June,  1903,  by  Mr.  James  Judge,  the  resident 
treasury  agent.  One  of  these,  a  female,  was  reported  by  Mr.  Judge  as  being 
40  feet  2  inches  long.  *  *  *  The  other  specimen,  a  male,  was  25  feet  5 
inches  long. 

These  were  skeletonized  and  sent  to  the  U.  S.  National  Museum, 
and  were  later  exhaustively  discussed  and  figured  by  True,  together 
with  a  third  individual  reported  by  Ezra  W.  Clark  as  occurring  at 
the  same  place  on  August  21,  1909,  but  which,  apparently,  was  not 
preserved  (1910,  pp.  2,  60  et.  seq.). 

Family  DELPHINIDAE. 
Orcinus  rectipinna  Cope.     Killer  Whale. 

Killer  whales  are  often  seen  about  the  Pribilofs  and  are  known  to 
prey  on  seals  and  sea  lions,  both  young  and  adults.  True  (1899,  p. 
353)  reported  as  follows: 

The  skull  of  a  killer  was  brought  from  St.  Paul  Island  in  1895  by  Mr. 
Charles  H.  Townsend,  obtained  from  a  specimen  which  came  ashore  to  the 
south  of  Hutchinsons  Hill.  Many  nominal  species  of  killers  have  been  es- 
tablished, but  it  has  not  yet  been  demonstrated  whether  there  are  really 
several  or  only  one. 

I  saw  two  killers  on  one  occasion  quite  close  inshore  at  St.  Paul  in  1895. 
In  1890  Mr.  Palmer  wrote :  "  A  few  seen  about  the  islands  in  May  and  early  in 
summer.  They  return  in  August.  We  saw  quite  a  number  on  August  12  be- 
tween St.  George  and  Unalaska." 

Most  of  the  natives  have  seen  killers  chasing  sea  lions,  and  have  seen  both 
sea  lions  and  killers  strand  on  the  rocky  shore.  The  majority  of  the  killers 
seen  had  a  large  whitish  blotch  on  each  side  of  the  back,  immediately  behind 
the  dorsal.  In  no  case  was  this  blotch  pure  white,  though  the  center  and  upper 
part  of  it  was  always  lighter  than  the  sides.  The  tip  of  the  dorsal  in  no 
case  turned  over. 

Preble  (in  Osgood,  et  al.,  1915,  p.  72)  has  summarized  the  rela- 
tions of  this  species  to  the  fur  seal,  as  follows : 

WThile  the  young  pups  are  still  about  the  islands  in  autumn  many  are  de- 
stroyed by  killer  whales  (Orca  gladiator),  which  are  frequently  observed 
singly  or  in  small  schools  cruising  about  in  front  of  the  rookeries  and  are 
known  to  prey  especially  on  the  pups.  The  following  actual  records  of  killer 
whales  observed  about  St.  Paul  Island  in  autumn,  selected  from  a  large  num- 
ber of  observations  taken  from  the  island  log  by  the  late  Doctor  Hahn,  indi- 
cate to  some  degree  the  part  played  by  them  in  the  destruction  of  young  seals. 
A  large  school  of  killers  was  seen  near  East  Landing  on  October  21,  1875, 
and  5  near  the  same  place  on  September  21,  1891 ;  1  seen  off  Reef  Rookery  on 
December  2,  1902,  was  playing  havoc  with  a  band  of  seals ;  fragments  of  both 


118  NORTH   AMERICAN   FAUNA.  tNo.  46 

cows  and  pups,  the  work  of  killer  whales,  were  found  strewn  along  the  beach 
at  Northeast  Point  on  November  6,  1904.  In  the  autumn  of  1907  killers  were 
reported  on  numerous  occasions,  and  native  watchmen  at  Northeast  Point  and 
Polovina  reported  considerable  destruction.  A  killer  24  feet  long  was  stranded 
at  Northeast  Point  on  December  16,  190S.  On  November  1,  1913,  G.  Dallas 
Hanna  observed  three  killers  close  to  the  reef  near  the  village  of  St.  George 
preying  on  the  seal  pups.  Two  of  these  came  so  close  to  the  bluffs  that  he 
was  able  to  hit  them  with  a  rifle  and  killed  at  least  one. 

These  records  indicate  that  killer  whales  are  by  no  means  uncommon  about 
the  Pribilofs.  The  stomachs  of  two  killers  examined  by  Captain  Bryant  con- 
tained, respectively,  18  and  24  seal  pups  [Rept.  Fur-Seal  Investigations,  189G-97, 
pt.  3,  p.  93,  1899],  and  it  is  certain  that  the  total  number  of  young  seals  killed 
by  them  must  be  very  great. 

As  possibly  having  some  bearing  on  the  seasonal  movements  of 
killer  whales,  the  dates  on  which  they  have  been  noted  in  the  St. 
Paul  Island  log,  as  extracted  by  Hahn,  may  be  given:  October  21, 
1875;  June  6,  1877;  May  21,  1881;  June  8,  1881  (Walrus  Island); 
June,  1882;  May  15,  1884;  May  21,  1884;  May  19,  1886;  May  28, 
1888 ;  May  31,  1889 ;  September  21,  1889 ;  June  1, 1894;  May  26,  1895 ; 
May  22,  23,  24,  and  26,  1900;  July  18,  1902  (school);  December 
2,  1902 ;  May  5,  1903 ;  November  8,  1904 ;  October  12,  November  16, 
1907 ;  December  16,  1908 ;  December  6,  1909 ;  June  6,  1910 ;  June  21, 
1910;  May  31,  1911  (Hahn). 

G.  Dallas  Hanna,  in  notes  kindly  forwarded  recently,  gives  some 
data  regarding  the  occurrence  of  killer  whales  about  the  islands 
within  the  past  few  years.    A  part  of  his  account  may  be  quoted : 

In  the  fall  schools  of  them  cruise  round  and  round  the  beaches  in  close  forma- 
tion and  actually  devour  seals  by  the  hundreds.  The  official  journals  which 
are  kept  on  the  islands  contain  many  accounts  by  eyewitnesses  of  their  depre- 
dations.   I  once  saw  two  killers  take  three  seals  in  five  minutes. 

The  fall  visit  of  the  animals  coincides  with  the  period  when  the 
young  seal  pups  are  learning  to  swim,  and  it  is  upon  them  that  the 
greatest  damage  is  wrought.  At  times,  however,  they  have  been 
seen  to  capture  older  animals.  In  the  spring  of  1917  a  school  of 
them  maintained  a  stand  between  St.  Paul  Island  and  Sealion 
Rock,  and  for  more  than  a  week  succeeded  in  capturing  or  driving 
away  practically  every  seal  which  approached  the  great  breeding 
grounds  in  the  vicinity. 

Phocaena  phocaena  (Linnaeus).     Harbor  Porpoise. 

True  (1899,  p.  353)  sums  up  the  evidence  regarding  the  occurrence 
of  this  species  in  Bering  Sea  as  follows: 

A  few  bones  of  a  small  porpoise,  apparently  of  this  species,  were  picked  up 
at  St.  Paul  June  3,  1890,  and  two  small  schools  were  seen  in  the  harbor  at 
Unalaska  May  20  and  21,  the  same  year.  A  specimen  of  this  species  was  ob- 
tained by  Mr.  Charles  H.  Townsend  at  Captains  Harbor,  Unalaska,  August 
17,  1895. 


1923.]  MAMMALS   OF   THE   PRIBILOF   ISLANDS.  119 

I  find  in  the  collection  of  the  U.  S.  National  Museum  the  skull  and 
skeleton,  No.  49428,  taken  by  True  and  Prentiss  at  St.  Paul,  in  the 
summer  of  1895.  Another  skull  and  skeleton  No.  217912,  male,  was 
taken  at  Northeast  Point,  St.  Paul,  by  G.  D.  Hanna,  on  July  19, 
1916. 

On  February  12,  1917,  a  school  of  porpoises  was  forced  ashore 
by  the  heavy  ice  in  the  Village  Cove  at  St.  George  Island.  Thirteen 
were  thus  trapped,  12  of  which  were  carefully  measured  and  skele- 
tonized. Eleven  of  these,  10  complete  skeletons  and  1  skull,  are  now 
in  the  collection  of  the  U.  S.  National  Museum. 

Family  ELEPHANTIDAE. 

Elephas  primigenius  Blumenbach.     Mammoth. 

Remains  of  the  mammoth  have  been  found  on  several  occasions. 
The  earliest  specimen  seems  to  have  been  a  tooth  found  on  St.  George 
in  1836.  Dall  (in  Dall  and  Harris,  1892,  p.  266)  refers  to  this  in 
part  as  follows :  "  [The  discovery  of]  a  mammoth  tooth  on  the  island 
of  St.  George  of  the  Pribilof  group,  in  1836,  vouched  for  by  Veniam- 
inoff  (Unal.,  I,  p.  106)."  A  native  chief,  Gromoff,  informed  Dr. 
W.  L.  Hahn  that  he  had  found  two  mammoth  tusks  on  St.  Paul,  one 
on  the  north  shore  and  the  other  at  Northeast  Point.  The  latter  is 
probably  the  one  referred  to  by  Stanley-Brown  (1892,  p.  499),  who 
says : 

There  are  two  fragments  of  paleontologic  evidence  connected  with  the  islands 
which,  as  they  have  been  used  by  writers,  demand  a  cautionary  word.  The 
tusk  of  a  mammoth  was  found  in  the  sands  of  Northeast  Point  on  St.  Paul 
Island,  and  the  tooth  of  one  is  reported  as  coming  from  the  shores  of  St. 
George.  As  there  is  not  a  foot  of  earth  on  either  island,  save  that  which  has 
resulted  from  the  decomposition  of  the  native  rock  and  the  decay  of  vegeta- 
tion, the  value  of  such  testimony  is  questionable. 

Whether  Stanley-Brown  doubts  that  the  specimens  were  actually 
taken  on  the  islands,  or  questions  the  deductions  which  had  been 
drawn  from  their  presence  by  previous  writers  on  the  geology  of  the 
North,  is  not  clear. 

George  M.  Dawson  (1894,  p.  4),  in  the  course  of  a  somewhat  ex- 
tended discussion  of  the  subject  of  mammoth  remains  in  north- 
western America,  refers  to  these  finds  and  to  Stanley-Brown's  com- 
ment, and  professes  not  to  understand  "  the  precise  intention  of  the 
cautionary  remark  just  quoted."  His  conclusion  is  that  the  presence 
of  these  remains  on  the  Pribilofs  indicates  a  former  connection  with 
the  mainland. 

Frederic  A.  Lucas  (1898,  p.  718)  has  put  on  record  two  teeth  of 
the  mammoth  obtained  by  R.  E.  Snodgrass  from  the  cave  on  Bogos- 
lof  Hill,  St.  Paul  Island,  in  1897.    Lucas  (1.  c,  p.  718)  is  also  of  the 


120  NORTH   AMERICAN   FAUNA.  [No.  46 

opinion  that  the  presence  of  the  remains  of  the  mammoth  on  the 
group  argues  for  former  land  connection.  Maddren  (1905,  p.  21), 
in  discussing  this  and  other  finds,  has  described  at  some  length  the 
circumstances  attending  the  collection  of  these  teeth  from  the  testi- 
mony of  Bristow  Adams,  who  was  a  member  of  the -party  which 
made  the  collection. 

G.  Dallas  Hanna,  in  a  recent  paper  on  the  geology  of  the  Pribilofs 
(1919c,  p.  222),  makes  the  following  statement: 

It  should  be  stated  here  that  the  reports  of  the  finding  of  bones  of  fossil 
elephants  on  the  Pribilof  Islands  are  probably  attributable  to  practical  jokes 
which  have  been  played  on  credulous  naturalists  in  the  past.  No  such  bones 
have  thus  far  been  found  that  were  not  planted  by  man,  according  to  reports 
of  eyewitnesses  to  some  of  the  pranks. 

This  statement  can  hardly  be  considered  as  applicable  to  the  find- 
ing of  the  teeth  and  tusks  above  mentioned,  since  their  authenticity 
seems  to  be  as  well  established  as  is  usual  in  such  cases.  It  would 
seem  to  the  writer,  however,  that  these  remains  were  most  probably 
accidentally  transported  to  the  islands,  most  likely  on  floating  ice, 
and  that,  therefore,  their  occurrence  there  has  no  special  geological 
significance. 


BIBLIOGRAPHY. 
By  Edward  A.  Preble. 

The  following  list  of  titles  relate  mainly  to  the  birds  and  mam- 
mals of  the  Pribilof  Islands,  and,  while  known  to  be  incomplete,  is 
believed  to  include  all  the  more  important  works.  It  is  designed 
mainly  to  furnish  complete  citations  of  works  referred  to  in  the 
preparation  of  the  present  lists  of  birds  and  mammals,  and  es- 
pecially the  articles  drawn  upon  for  information.  Most  of  the 
voluminous  literature  relating  to  fur  seals  is  listed  in  Fur  Seals 
and  Other  Life  of  the  Pribilof  Islands,  by  Osgood,  Preble,  and  Parker, 
published  in  1915,  and  no  attempt  has  been  made  to  include  such 
titles  in  the  present  list. 

Allen,  Joel  Asaph. 

1877.  Nest  and  eggs  of  the  Alaskan  wren :  Bull.  Nuttall  Ornith.  Club,  vol. 
2,  no.  3,  p.  82,  July. 

Descriptions  of  first  nest  and  eggs  known,  taken  on  St.  George 
Island,  in  June,  1876. 
1902.  The  hair  seals    (family   Phocidae)    of  the  North  Pacific  Ocean  and 
Bering  Sea :  Bull.  Arner.  Mus.  Nat.  Hist.,  vol.  16,  art.  34,  pp.  459- 
499,  December  12. 

Original  description  of  Phoca  richardii  pribilofensis    (p.  495). 
Mention  of  ribbon  seal   {Histriophoca  fasciata)   from  vicinity  of 
Pribilofs. 
Baibd,  Spencer  Fullerton. 

1869.  On  additions  to  the  bird-fauna  of  North  America,  made  by  the  scien- 
tific corps  of  the  Russo-American  Telegraph  Company :  Trans. 
Chicago  Acad.  Sci.,  vol.  1,  pt.  2,  art.  10,  pp.  311-325. 

Contains  discussion  of  several  species  first  noted  from  the  islands 
by  Dall  and  Bannister,  including  original  description  of  Troglodytes 
aluscensis  from  St.  George  Island. 
Baird,  S.  F.,  T.  M.  Brewer,  and  Robert  Ridgwat. 
1884.  The  water  birds  of  North  America,  vol.  2. 

Original  description  of  Rissa  tridactyla  pollicaris   (p.  202). 
Bean,  Tarleton  Hoffman. 

1882.  Notes  on  birds  collected  during  the  summer  of  1880  in  Alaska  and 
Siberia :    Proc.  U.  S.  Nat.  Mus.,  vol.  5,  pp.  144-173,  July  8,  1882. 
Notes  on  a  few  species  from  the  Pribilofs  or  vicinity. 
Bent,  Arthur  Cleveland. 

1919.  Life  histories  of  North  American  diving  birds,  order  Pygopodes : 
Bull.  107,  U.  S.  Nat.  Mus.     245  p. 

Contains   many    references   to   habits  of  birds   on   the    Pribilof 
Islands. 

121     • 


122  NORTH   AMERICAN   FAUNA.  [No.  46 

Bent,  Arthur  Cleveland — Continued. 

1921.  Life  histories  of  North  American  gulls  and  terns,  order  Longipennes: 

Bull.  113,  U.  S.  Nat.  Mus.    345  p. 

Contains   many    references   to    habits   of  birds   on    the    Pribilof 
Islands. 

1922.  Life   histories   of  North   American   petrels   and   pelicans   and   their 

allies :  Bull.  121,  U.  S.  Nat.  Mus.    343  p. 

Contains  references  to  habits  of  a  few  species  on  the  Pribilofs. 
Bishop,  Louis  Bennett. 

1900.  Birds  of  the  Yukon  Region,  with  notes  on  other  species :  North 
Amer.  Fauna  No.  19,  pp>  47-96. 

Notes  on  a  few  species  taken  on  St.  George  Island  in  October, 
1899. 
Chichester,  Harry  D. 

1908.  [Report  on  condition  of  affairs  on  St.  George  Island,  August  14,  1905, 
to  June  1,  1906].  In  Reports  Relating  to  Alaska  Seal  Fisheries,  by 
W.  I.  Lembkey  and  others;  Senate  Doc.  376,  60th  Cong.,  1st  sess., 
pp.  47-58. 

Records,  in  discussion  of  food  of  foxes,  the  occurrence  of  vast 
numbers  of  "sea  quail"   (Simorhynchus  cristatellus) . 
Clark,  Austin  Hobart. 

1910.  The  birds  collected  and  observed  during  the  cruise  of  the  United 
States  Fisheries  steamer  Albatross  in  the  North  Pacific  Ocean, 
and  in  the  Bering.  Okhotsk,  Japan,  and  Eastern  Seas  from  April 
to  December,  1906:  Proc.  U.  S.  Nat.  Mus.,  vol.  38,  pp.  25-74, 
April  30. 

Records  specimen  of   Colaptes  auratus   luteus  from   St.   George 
Island,  the  first  record  for  the  Pribilofs. 
Coinde,  J.  P. 

1860.  Notice  sur  la  faune  ornithologique  de  l'ile  de  Saint-Paul,  suivie  de 
l'enumeration  de  quelques  especes  d'insectes  (Coleopteres)  des 
Aleoutiennes  et  du  Kamtschatka :  Rev.  et  Mag.  de  Zool.,  vol.  12, 
2e  ser.,  pp.  396-405. 

Earliest  account  of  a  collection  of  birds  from  the  Pribilofs,  nine 
species  taken  by  Doctor  Waraeck  in  1842. 
Dall.  William  Healet. 

1873.  Notes  on  the  avifauna  of  the  Aleutian  Islands  from  Unalaska  east- 

ward:  Proc.  Calif.  Acad.   Sci.,   vol.   5,  1873-74,  pp.  25-35,   Febru- 
ary 8. 

Includes    notes    on    the    occurrence    of   a    few    species    on    the 
Pribilofs. 

1874.  Notes  on  the  avifauna  of  the  Aleutian  Islands,  especially  those  west 

of  Unalaska:  Proc.  Calif.  Acad.  Sci.,  vol.  5,  1873-74,  pp.  270-281, 
Mar.  14. 

Includes    notes    on    the    occurrence    of    a    few    species    on    the 
Pribilofs. 
Dall,  W.  H.,  and  H.  M.  Bannister. 

1869.  List  of  the  birds  of  Alaska,  with  biographical  notes:  Trans.  Chicago 
Acad,  of  Sci.,  vol.  1,  pp.  267-360. 

Contains  first  recorded  instance  of  occurrence  of  several  species 
on  the  Pribilofs. 


1923.]  BIBLIOGRAPHY,   BIRDS   AND   MAMMALS.  123 

Dall,  W.  H.,  and  Gilbert  D.  Harris. 

1892.  Correlation  papers:  Neocene:  Bull.  84,  U.  S.  Geol.  Surv.  349  p.,  3 
pis.,  43  figs. 

Mention  (p.  266),  on  the  authority  of  Veniaminof,  of  a  mammoth 
tooth  found  on  St.  George  Island  in  1836. 
Dawson,  George  Mercer. 

1894.  Notes  on  the  occurrence  of  mammoth  remains  in  the  Yukon  District 
of  Canada  and  Alaska.  Quart.  Journ.  Geol.  Soc.  London,  vol.  50, 
pp.  1-9. 

Includes    discussion   of   probable   origin   of    mammoth    remains 
found  on  the  Pribilofs. 
Dwight,  Jonathan,  Jr. 

1918.  A  new  species  of  loon  (Gavia  viridigularis)  from  northeastern  Si- 
beria :  The  Auk,  vol.  35,  no.  2,  pp.  196-199,  April. 

The  only   loon  of  this   type  from  the  Pribilofs  appears   to  be 
referable  to  this  species. 
Elliott,  Henry  Wood,  and  Elliott  Coues. 

1874.  Report  on  the  Pribilov  Group  or  Seal  Islands  of  Alaska,  by  Henry 

W.  Elliott,  assistant  agent,  Treasury  Department.  Pub.  by  Treas. 
Dept. ;  129  p.  (not  numbered),  2  maps,  45  pis.;  in  appendix  is  an 
article  entitled  "  Ornithology  of  the  Pribilov  Islands,  by  Dr.  Elliott 
Coues,  U.  S.  A.  (based  on  Mr.  H.  W.  Elliott's  manuscripts  and 
collections)." 

Forty  species  are  treated  under  specific  headings  and  two  or 
three  others  are  referred  to  by  Elliott  in  the  introduction  to  the 
formal  list.  Original  description  of  Tringa  ptilocnemis  given  in  a 
footnote.  Bears  date  of  1873  on  title  page,  and  1875  on  cover, 
but  was  issued  early  in  1874. 

1875.  A  report  upon  the  condition  of  affairs  in  the  Territory  of  Alaska, 

by  Henry  W.  Elliott,  special  agent,  Treasury  Department.  Pub. 
by  Treas.  Dept. ;  277  p.,  many  illus. ;  chap.  IX,  "  Ornithology  of 
the  Pribilof  Islands,"  by  Dr.  Elliott  Coues,  pp.  168-212. 

A  reprint,  without  the  illustrations,  of  the  report  (unpaged)  by 
the  same  authors  published  in  1874. 
Elliott,  Henry  W. 

1881.  The  seal-islands  of  Alaska  ;  176  p.,  with  maps  and  text  illus.  Pub- 
lished (together  with  "The  history  and  present  condition  of  the 
fishery  industries,"  by  G.  Brown  Goode)  as  Bull.  Tenth  Census  of 
United  States,  by  Int.  Dept. 

Reprinted,  with  practically  no  changes  and  the  same  pagination, 
as  Spec.  Bull.  176,  U.  S.  Comm.  Fish  and  Fisheries,  1882.  Same 
report  again  issued  in  1884  as  Misc.  Doc.  42,  Pt.  8,  47th  Cong.,  2d 
sess.  (with  three  other  reports)  with  addition  of  an  index,  increas- 
ing the  size  to  188  pages.  This  latter  report  also  issued  as  a 
separate. 

Contains  lists  of  the  mammals  and  birds  of  the  group,  pp.  124- 
136,  in  substance  much  like  the  lists  in  his  earlier  work,  but  re- 
written, and  entirely  by  Elliott. 
Evermann,  Barton  Warren. 

1913.  Eighteen  species  of  birds  new  to  the  Pribilof  Islands,  including  four 
new  to  North  America :  The  Auk,  vol.  30,  no.  1,  pp.  15-18, 
January. 

Based  on  collections  made  by  W.  L.  Hahn  and  M.  C.  Marsh  on 
St.  Paul  Island  from  1910  to  1912. 


124 


NORTH   AMERICAN   FAUNA. 


[No.  46 


1914. 


1916. 


Grin nell,  Joseph. 

1901.  Record  of  Alaskan  birds :     The  Condor,  vol.  3,  pp.  19-23,  January. 

Briefly  annotated  list  of  21  species  in  the  collection  of  Leland 

Stanford  University,  taken  by  R.  E.  Snodgrass  and  A.  W.  Greely 

on  the  Pribilofs  in   the  summer  of  1897.     Tetanus  melanoleucus 

and  Saxicola  ocnanthe  added  to  the  Pribilof  list. 

Hanna,  G.  Dallas.     (See  also  Mailliard  and  Hanna.) 

Interesting    mammals    on    the    Pribilof    Islands:      Proc.    Biol.    Sot, 
Washington,  vol.  27,  p.  218. 

Notes   occurrence   of   polar   bear,   walrus,   and   cachelot   on   St. 
George  in  1914. 
Records  of  birds  new  to  the  Pribilof  Islands,  including  two  new  to 
North  America :     The  Auk,  vol.  33,  no.  4,  pp.  400-403,  October. 

Thirteen  species  added  to  the  Pribilof  list ;  notes  on  eight  other 
species. 
1917.  The  summer  birds  of  the  St.  Matthew  Island  Bird  Reservation :    The 
Auk,  vol.  34,  no.  4,  pp.  403-410. 

Contains  a  few  notes  on  Pribilof  birds. 
1919a.  Additions  to  the  avifauna  of  the  Pribilof  Islands,  Alaska,  including 
species  new  to  North   America :     Journ.   Washington  Acad.   Sci., 
vol.  9,  no.  6,  pp.  176-177,  March  19. 

Announcement  of  21  species  taken  on  the  Pribilofs  for  the  first 
time ;  four  of  these  new  to  North  America.  These  were  later 
formally  recorded,  with  dates,  in  The  Auk,  vol.  37,  1920. 
1919b.  Check  list  of  birds  of  the  Pribilof  Islands,  Alaska,  with  the  names  of 
persons  first  recording  the  species  from  the  Islands :  In  Alaska 
Fisheries  and  Fur  Industries  in  1918.  Bur.  Fisheries,  Doc.  No. 
872,  by  Ward  T.  Bower;  App.  VII,  Rept.  U.  S.  Coram.  Fisheries 
for  1918,  pp.  105-107. 
1919c.  Geological  notes  on  the  Pribilof  Islands,  Alaska,  with  an  account  of 
the  fossil  diatoms:  Amer.  Jour.  Sci.,  vol.  48  (4th  ser.),  pp.  216- 
224,  September. 

Notes   the   occurrence   of   a   few   remains   of   mammals   in   the 
Pliocene  deposits. 
1920a.  Additions  to  the  avifauna  of  the  Pribilof  Islands,  Alaska,  including 
four    species    new    to    North    America :      The    Auk,    vol.    37,    pp. 
248-254,  April. 

Twenty-one  species  added  to  the  Pribilof  list ;  four  of  these  new 
to  North  America.  (These  were  listed  in  Jour.  Washington  Acad. 
Sci.,  vol.  9,  no.  6,  pp.  176,  177,  publ.  March  19,  1919,  in  a  brief  report 
of  the  meeting  of  the  Biological  Society  of  Washington  for  Jan- 
uary 25,  when  the  discoveries  were  announced.)  Notes  on  several 
other  species. 
1920b.  New  and  interesting  records  of  Pribilof  Island  birds:  The  Condor, 
vol.  22,  no.  5,  pp.  173-175,  September  24. 

Three  species  added  to  the  Pribilof  list ;  notes  on  various  other 
species  of  particular  interest ;  table  showing  islands  on  which  the 
various  breeders  nest. 
1921a.  The    Pribilof    sandpiper:  The    Condor,    vol.    23,    no.    2,    pp.    50-57, 
March  31. 

Account  of  the  breeding  distribution,  nesting  habits,  and  eggs  of 
Arquatclla  ptilnocnemis. 


1923.]  BIBLIOGRAPHY,   BIRDS  AND  MAMMALS.  125 

Hanna,  G.  Dallas — Continued. 

1921b.  Natural  history  records  of  Pribilof  Islands :  in  Alaska  Fisheries 
and  Fur  Industries  in  1920,  by  Ward  T.  Bower :  Bur.  Fisheries, 
Doc.  No.  909,  App.  VI,  Kept.  U.  S.  Coram.  Fisheries  for  1921,  pp. 
122-127. 

List  of   more   important   papers   on   natural   history   published 
during  recent  years. 
Harttng,  James  Edmund. 

1874.  On  a  new  species  of  Tringa  from  Alaska :  Proc.  Zool.  Soc.  London, 
for  year  1874,  pp.  242-244. 

Description  of  Tringa  gracilis   (=Arquatella  ptilocnemis)    from 
St.  Paul  Island.     In  a  postscript,  the  author  admits  that  his  sup- 
posed new  species  is  identical  with  T.  ptilocnemis  described  a  few 
months  earlier  by  Coues. 
Heath,  Harold. 

1920.  The  nesting  habits  of  the  Alaska  wren:  The  Condor,  vol.  22,  no.  2, 
March-April,  pp.  49-55. 

Based  mainly  on  observations  made  on   St.  George  Island,  in 
1918. 
Judge,  James. 

1909.  The  blue  foxes  of  the  Pribilof  Islands :  Rept.  Amer.  Breeders'  Asso., 
vol.  5,  pp.  325-340,  1909. 

Historical  account  of  fox  industry,  including  life  history, 
methods  of  trapping,  and  yield.  Relates  mainly  to  St.  George 
Island,  where  the  methods  now  practiced  were  developed,  mainly 
by  the  author. 
1912.  The  blue  foxes  of  St.  Paul  and  Otter  Islands,  Alaska :  Ann.  Rept. 
Amer.  Breeders'  Asso.,  vol.  7,  pp.  275-279,  1912. 

Account  of  feeding,  methods  of  capture,  and  numbers  on  these 
islands. 
Lucas,  Frederic  A. 

1898.  The  occurrence  of  mammoth  remains  on  the  Pribilof  Islands : 
Science  (N.  S.),  vol.  8,  no.  203,  p.  718,  November  18. 

Records   finding   of   remains    of   mammoth   and   polar   bear    in 
Bogoslof  Cave,  St.  Paul  Island,  in  1897. 
Lutz,  John  E. 

1889.  Report  of  Lieut.  J.  E.  Lutz :  Rept.  of  Cruise  of  revenue  steamer 
Corivin  in  Arctic  Ocean,  1884,  pp.  28-35,  1889. 

Records  eggs  or  young  of  fourteen  species  of  birds  taken  on 
Otter  Island  in  1884. 
Maddeen,  Alfred  Geddes. 

1905.  Smithsonian  Exploration  in  Alaska  in  1904,  in  search  of  mammoth 
and  other  fossil  remains.  Pub.  no.  1584,  vol.  49,  Smiths.  Misc. 
Coll. 

Several  finds  of  remains  of  the  mammoth  on  the  Pribilofs  dis- 
cussed (pp.  20-22). 
Mailliard,  Joseph. 

1923.  The  tree  swallow  added  to  the  Pribilof  list:  The  Condor,  vol.  25, 
no.  1,  p.  31,  January. 

Records  a  specimen  of  Iridoprocne  bicolor  taken  on  St.  Paul  Island. 
1087318— 23 9 


126  NORTH  AMERICAN  FAUNA.  [No.  46 

Mailliard,  Joseph,  and  G.  Dallas  Hanna. 

1921.  New  bird  records  for  North  America,   with  notes  on  the  Pribilof 

Island  list :    The  Condor,  vol.  23,  no.  3,  pp.  93-95,  May. 

Adds  two  species,  Micropus  pacificus  and  Loxia  leucoptera,  to 
the  Pribilof  list. 
Meeriam,  Clinton  Hart. 

1895a.  Mammals  of  the  Pribilof  Islands:  Science,  N.  S.,  vol.  1,  no.  25, 
p.  698,  June  21. 

Eight  species  enumerated  in  notice  of  paper  read  by  Doctor 
Merriam  at  meeting  of  the  Biological  Society  of  Washington,  May 
18,  1895.  / 

1895b.  Synopsis  of  American  shrews  of  the  genus  Sorex:  North  Amer. 
Fauna  No.  10,  pp.  57-124,  December  31. 

Contains  original  description  of  Sorex  pribilof  ensis  from   St. 
Paul  Island. 
Nelson,  Edwaed  William. 

1883.  Birds  of  Behring  Sea  and  the  Arctic  Ocean :  Cruise  of  Revenue- 
Steamer  Corurin  in  Alaska  and  the  N.  W.  Arctic  Ocean  in  1881 : 
House  Exec.  Doc.  no.  105,  47th  Cong.,  2d  Sess.,  pp.  55-118. 

Contains  many  references  to  Pribilof  birds,  mainly  from  Dall 
and  Elliott. 
1887.    Report  on  natural  history  collections  made  in  Alaska  between  the 
years  1877  and  1881 :  Arctic  ser.  publ.  issued  in  connection  with 
Signal  Service,  U.  S.  Army,  no.  3 ;  Birds,  pp.  21-226. 
Mention  of  a  few  species  from  the  Pribilofs. 
Oberholsee,  Habby  Chuech. 

1918.  The  subspecies  of  Larus  hyperboreus  Gunnerus:  The  Auk,  vol.  35, 
no.  4,  pp.  467^174,  October. 

Specimens  of  L.  h.  hyperboreus  and  L.  h.  barrovianus  recorded 
from  the  Pribilofs. 

1922.  Notes  on  North  American  birds,  XI:     The  Auk,  vol.  39,  no.  1,  pp. 

72-78,  January. 

The  owl  (no.  239184,  U.  S.  Nat.  Mus.),  taken  on  St.  Paul 
January  26,  1911,  and  recorded  by  Evermann  (The  Auk,  vol.  30, 
p.  18,  January,  1913)  as  Cryptoglaux  funerea  funerea,  proves  to 
be  referable  to  C.  f.  magna  (Buturlin). 
Osgood,  Wilfeed  Hudson,  Edwaed  Alexandeb  Pbeble,  and  Geobge  Howard 
Paekeb. 
1915.  Fur  seals  and  other  life  of  the  Pribilof  Islands,  Alaska,  in  1914: 
Senate  Doc.  980,  63d  Cong.,  3d  Sess.;  also  Doc.  820,  U.  S.  Bur. 
Fisheries,  172  p.,  24  maps,  June  19.  (Document  820  also  reprinted 
with  addition  of  index,  pp.  I-V,  1916.) 

Mainly    on    fur   seals;  brief   treatment  of  other  mammals    and 
birds. 
Palmer,  William. 

1894a.  Do  wading  birds  swim?     The  Nidiologist,  vol.  1,  no.  12,  p.  175. 

Instance  of  swimming  of  Pribilof  sandpiper ;  mention  of  other 
species. 
1894b.  An  Asiatic  cuckoo  on  the  Pribilof  Islands,  Alaska :  The  Auk,  vol.  11, 
no.  3,  p.  325,  October. 

Record  of  a  specimen  taken  by  him  on  St.  Paul  Island,  July  4, 
1S90,  first  record  for  North  America. 


1923.]  BIBLIOGRAPHY,   BIRDS  AND   MAMMALS.  127 

Palmer,  William — Continued. 

1899.  The  avifauna  of  the  Pribilof  Islands:  In  the  Ftir  Seals  and  Fur- 
Seal  Islands  of  the  North  Pacific  Ocean,  pt.  3,  pp.  355-431,  with 
4  pis. 

History  and  detailed  discussion  of  ornithological  work  on  the 
islands,  with  comparative  lists  of  species,  discussion  of  distribu- 
tion and  migration,  a  bibliography,  and  an  annotated  list  of  69 
species. 
Prentiss,  Daniel  Webster,  Jr. 

1902.  Notes  on  birds  of  the  Pribilof  Islands:  The  Osprey,  vol.  1  (new 
series),  no.  7,  pp.  97-103,  July. 

Notes  on  25  species  observed  on  St.  George  and  St.  Paul  during 
the   summer   of   1895.     Specimens   of   Pagophila   alba,   Aegialitis 
semipalmata,  and  Oceanodroma  furcata  recorded  from  the  islands 
for   the  first   time. 
Ridgway,    Robert. 

1SS6.  Tringa  damacensis  (Horsf.)  in  Alaska;  a  sandpiper  new  to  the  North 
American  fauna :  The  Auk,  vol.  3,  no.  2,  p.  275,  April. 

Records  specimen  taken  by  C.  H.  Townsend  on   Otter   Island, 
June  8,  1885. 
1887.  A  manual  of  North  American  birds ;  631  p.,  124  pis. 

Original   description  of  Plectrophenax  nivalis  townsendi,  from 
Otter  Island   (p.  403). 
1898.  New  species,  etc.,   of  American  birds,  II :  The  Auk,  vol.  15,  no.  4, 
pp.  319-324,  October. 

Description   of   Calcarius  lapponicus  alascensis,  from  St.   Paul 
Island  (p.  320). 
Riley,  Joseph  Harvey. 

1917.  A  bird  new  to  the  North  American  fauna :  The  Auk,  vol.  34,  no.  2, 
p.  210,  April. 

Records  specimen  of  Pinicola  c.  kamtschathensis  taken  on   St. 
George  Island,  October,  1915. 
Scammon,  Charles  Melville,  and  W.  H.  Dall. 

1874.  The  marine  mammals  of  the  northwestern  coast  of  North  America, 
described  and  illustrated :  together  with  an  account  of  the  Ameri- 
can whale-fishery.    4to.,  p.  319  4-V,  San  Francisco  and  New  York. 
An  appendix,  by  W.  H.  Dall,  pp.  281-319,  comprises  mainly  a 
catalogue  of  the  Cetacea  of  the  North  Pacific  Ocean. 
Seale,  Alvin. 

1898.  Notes  on  Alaskan  water  birds :  Proc.  Acad.  Nat.  Sci.  Philadelphia, 
1898,  pp.  126-140. 

Includes  brief  annotations  on  eleven  species  taken  on  the  Pribi- 
lofs  by  A.  W.  Greely  and  R.  E.  Snodgrass.  Totanus  melanoleucus 
and  Saxicola  oenanthe  added  to  the  Pribilof  list ;  Tringa  couesi, 
also  added,  evidently  based  on  nri'sidentitication ;  dates  of  speci- 
mens unreliable. 
Stanley-Brown,  Joseph. 

1892.  Geology  of  the  Pribilof  Islands:  Bull.  Geol.  Soc.  Amer.,  vol.  3,  pp. 
496-500,  1892. 

Reference  made  to  discovery  of  remains  of  mammoth  on  the 
islands. 


128  NORTH   AMERICAN  FAUNA.  [No.  46 

Stejneger,  Leonhard. 

1889.  Notes  on  the  downy  young  of  the  parrot  auk  and  of  the  crested  auk : 
Rept.  of  Cruise  of  Revenue-Steamer  Corwin  in  Arctic  Ocean, 
1884,  pp.  125,  126. 

Describes   and   figures  the  young  of  these  species,   not  before 
known,  from  specimens  taken  on  Otter  Island  by  J.  E.  Lutz  in 
1884. 
1885.  Results    of   ornithological   explorations   in   the   Commander    Islands 
and  in  Kamtschatka :  Bull.  29,  U.  S.  Nat.  Mus.,  382  p.,  8  pis. 
A  few  notes  on  Pribilof  birds. 
Townsend,  Charles  Haskins.  / 

1887.  Notes   on   the  natural   history   and   ethnology   of  northern   Alaska  : 

Rept.  of  Revenue  Marine  Steamer  Corwin  in  Arctic  Ocean  in  1885. 

Includes   "  Notes  on  Mammals,   Birds,  and  Fishes  obtained  at 

various  places  between  the  Aleutian  Islands  and  Kotzebuc  Sound  " 

(pp.  96-102).     A  number  of  species  recorded  from  the  Pribilof s. 

True,  Frederick  William. 

1899.  Mammals  of  the  Pribilof  Islands :  In  Fur  Seals  and  Fur-Seal  Islands 
of  North  Pacific  Ocean,  by  David  Starr  Jordan  and  others;  pt.  3, 
pp.  345-354. 

Notes  on   16  species,   native  and   introduced,   found   about   the 
group,  and  two  extirpated  species. 
1904.  Note  on   three   very   large  beaked  whales  from   the  North  Pacific : 
Science  (N.  S.),  vol.  20,  no.  521,  pp.  8S8-889,  December  23. 

Records  two  large  specimens,  the  skeletons  of  which  were  later 
received  by  the  U.  S.  National  Museum,  found  by  James  Judge, 
on  the  shore  of  St.  George  Island  in  June,  1903. 
1910.  An  account  of  the  beaked  whales  of  the  Family  Ziphiidae  in  the 
collection  of  the  United  States  National  Museum,  with  remarks 
on  some  specimens  in  other  American  Museums :  Bull.  73,  U.  S. 
Natl.  Mus.,  89  p.,  42  pis. 

Includes  account  of  two  skeletons  of  Berardius  bairdii  from  St. 
George  Island. 
Turner,  Ltjcien  M. 

1W0.  Contributions    to    the    natural    history    of    Alaska,    extending    from 
May,  1874,  to  August,  1881 :  Arctic  ser.  publ.,  issued  in  connec- 
tion with  Signal  Service,  U.  S.  A. ;  no.  2,  pt.  5 ;  Birds,  pp.  115-196. 
A  few  species  mentioned  as  found  on  the  Pribilofs. 
Veniaminof,  Bishop  Innocent. 

1840.  [Extract  from]  Zabieska  ob  Octrovah  Oonahlalshkenskaho  Otdayla. 
St.  Petersburg,  1840.  Translation  of  "  selections  most  pertinent  to 
subject"  (Pribilof  Islands)  published  by  Elliott  in  his  report 
on  the  condition  of  affairs  in  the  Territory  of  Alaska,  1875,  pp. 
241-244. 

A  paragraph  mentioning  somewhat  over  a  dozen  species  of  birds 
which  visit  the  Pribilof  Islands  seems  to  be  the  earliest  note  on 
the  ornithology  of  the  group. 


A  BIOLOGICAL  SURVEY  OF  THE   PRIBILOF 
ISLANDS,  ALASKA. 


Part  II.  INSECTS,  ARACHNIDS,  AND  CHILOPODS  OF  THE 
PRIBILOF  ISLANDS,  ALASKA. 


•  INTRODUCTION. 

By  W.  L.  McAtee,  In  Charge  Food  HaMts  Research, 
Bureau  of  Biological  Survey. 

The  bulk  of  the  material  upon  which  the  present  report  is  based 
was  collected  by  Alvin  G.  and  Elsie  G.  Whitney  from  October,  1912, 
to  July,  1914,  and  by  G.  Dallas  Hanna  in  1914,  1915,  1916,  and  1917. 
The  collectors  were  employed  on  the  Pribilof  Islands  during  these 
3'ears  by  the  U.  S.  Bureau  of  Fisheries  and  thanks  are  due  the  Chief 
of  that  Bureau  for  turning  over  their  collections  of  invertebrates  to 
the  Biological  Survey.  It  has  been  of  great  assistance  to  have  this 
material  for  working  up  simultaneously  with  the  examination  of  the 
bird  stomachs,  reported  on  in  earlier  pages,  which  also  for  the  most 
part  were  donated  by  the  Bureau  of  Fisheries.  Material  of  both 
classes  from  this  source  was  supplemented  by  collections  made  by 
Edward  A.  Preble,  of  the  Biological  Survey,  in  1914.  The  Whitneys 
and  Hanna  used  a  system  of  lot  numbers  for  their  collections  of  in- 
vertebrates, and  these  numbers  have  in  all  cases  been  placed  on  the 
specimen  labels.  Data  for  lot  numbers  applying  to  more  than  single 
specimens  are  reproduced  on  pages  132  to  138,  as  they  may  be  useful 
in  future  when  these  collections  are  distributed.  All  type  specimens 
mentioned  in  the  following  reports  as  well  as  the  bulk  of  the  remain- 
ing material  will  be  deposited  in  the  U.  S.  National  Museum. 

Previous  general  treatises  upon  the  insects  and  arachnids  of  the 
Pribilof  Islands  are  three  in  number.  The  first  of  these  is  the 
List  of  Insects  Hitherto  Known  from  the  Pribilof  Islands,  which 
appeared  in  the  report  on  The  Fur  Seals  and  Fur-seal  Islands  of 
the  North  Pacific  Ocean,  Part  III,  pages  547-554,  1899;  prepared 
by  E.  A.  Schwarz,  with  the  assistance  in  a  few  groups  of  M.  L. 
Linell,  W.  H.  Ashmead,  D.  W.  Coquillett,  and  Herbert  Osborn. 
The  second  report  was  contained  in  several  of  the  Papers  from  the 
Harriman  Alaska  Expedition,  mostly  published  in  the  Proceedings 

129 


130 


NORTH   AMERICAN   FAUNA. 


[No.  46 


of  the  Washington  Academy  of  Sciences  from  1900  to  1902,  and 
reprinted  with  some  additions  as  volumes  8  and  9,  Reports  of  the 
Harriman  Alaska  Expedition,  1904.  These  articles  appeared  under 
the  authorship  of  the  following  12  entomologists:  W.  H.  Ashmead, 
Nathan  Banks,  A.  N.  Caudell,  O.  F.  Cook,  D.  W.  Coquillett,  R.  P. 
Currie,  H.  G.  Dyar,  J.  W.  Folsom,  Otto  Heidemann,  Trevor  Kin- 
caid,  Theo.  Pergande,  and  E.  A.  Schwarz.  More  recently  a  report 
on  specimens  of  insects  collected  in  the  summer  of  1920  has  appeared 
in  Proceedings  of  the  California  Academy  of  Sciences,  Fourth  Series, 
volume  11,  No.  14,  pages  153-&5,  November  2,  1921.  The  collabo- 
rators in  this  case  were  E.  C.  Van  Dyke,  M.  C.  Van  Duzee,  F.  R. 
Cole,  J.  R.  Malloch,  C.  P.  Alexander,  H.  Frison,  A.  D.  MacGilli- 
vray,  and  E.  P.  Van  Duzee.  • 

Like  its  predecessors,  the  present  report  was  brought  to  comple- 
tion only  through  the  efforts  of  a  number  of  specialists,  and  the 
Biological  Survey  desires  to  put  on  record  its  hearty  appreciation 
of  their  valued  cooperation.  The  authors  of  parts  of  the  following 
report  are  C.  P.  Alexander,  Nathan  Banks,  R.  V.  Chamberlin, 
G.  F.  Ferris,  W.  T.  M.  Forbes,  Morgan  Hebard,  W.  L.  McAtee, 
J.  R.  Malloch,  Edith  M.  Patch,  H.  L.  Viereck,  W.  R.  Walton,  and 
H.  F.  Wickham.  The  bureau  is  also  indebted  to  Dr.  J.  W.  Folsom 
for  assistance  in  identifying  specimens  of  Collembola. 

The  progress  of  knowledge  of  the  insect,  arachnid,  and  myriapod 
fauna  of  the  Pribilof  Group  is  shown  in  the  subjoined  table.  Only 
fully  identified  forms  have  been  included  in  the  tabulation,  generic 
determinations  being  ignored. 

Species  of  insects,  arachnids,  and  cMlopods  reported  from  the  Pribilof  Islands. 


Fur-seal  Islands 
Report,  1899. 

Harriman  Reports, 

1900-1904. 

CaliforniaAcademy 
Report,  1921. 

Present  report. 

Group. 

Number 
of  species. 

Described 
as  new. 

Number 
of  species. 

Described 
as  new. 

Number 
of  species. 

Described 
as  new. 

Number 
of  species. 

Described 
as  new. 

3 

1 

4 

2 

5 

3 

3 

3 

5 

8 

49 

1 

75 

81 

33 

4 

1 

1 

1 

2 
1 
2 
2 
4 

1 

1 

1 

1 

3 

6 

26 

2 

20 

1 

1 

7 
1 
1 

33 

7 

8 
6 

21 

Hymenoptera 

66 

53 

2 

1 

2 

3 

1 

Total 

31 

2 

83 

58 

77 

15 

276 

27 

While  a  comparatively  large  number  of  species  of  insects  are 
known  from  the  Pribilof  Group,  discussion  of  the  relations  of  this 
fauna  to  that  of  other  areas  can  be  only  tentative  for  the  following 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  131 

reasons:  (1)  More  than  a  third  of  the  species  recorded  in  the  fore- 
going table  have  been  described  as  new  from  the  islands;  (2)  very 
few  of  these  have  been  collected  subsequently  elsewhere;  and  (3) 
the  distribution  of  most  of  the  remaining  species  is  very  imperfectly 
known. 

These  limitations  understood,  the  following  may  be  said  of  species 
known  both  from  the  Pribilof  Islands  and  other  areas:  All  of  the 
previously  known  spring-tails  (3  species  of  Collembola)  seem  to  be 
Holarctic  in  distribution.  The  two  species  of  roaches  (Orthoptera) 
are  only  waifs  on  the  Pribilof s  brought  there  on  ships  from  the 
south.  Of  the  5  kinds  of  bird  lice  (Mallophaga),  3  have  been  re- 
corded from  Europe  and  probably  are  Holarctic,  while  2  are  known 
from  other  localities  in  the  Pacific  Ocean.  The  distribution  of  these 
ectoparasites  as  well  as  of  the  sucking  lice  (one  species  on  man  and 
two  on  the  fur  seal)  depends  entirely  on  that  of  their  hosts ;  by  living 
next  the  skin  of  warm-blooded  animals  they  enjoy  a  habitat  almost 
uniform  as  to  temperature  and  other  essentials  and  are  thus  prac- 
tically exempt,  so  far  as.  direct  influence  is  concerned,  from  factors 
bearing  on  the  distribution  of  animals  in  general. 

One  of  the  Homoptera  recorded  from  the  Pribilofs  was  described 
from  Bering  Island,  while  of  the  Heteroptera,  one  plant  bug  is  Hol- 
arctic, another  is  known  from  northwestern  North  America  and  the 
northern  Pacific  Islands,  and  the  bed  bug  exists  practically  every- 
where that  there  are  permanent  habitations  of  man.  Of  the  caddis 
flies  (Trichoptera),  one  is  known  from  the  mainland  of  Alaska  and 
another  is  Holarctic.  Of  the  several  species  of  moths  (Lepidoptera), 
2  occur  in  northwestern  North  America,  2  in  Alaska,  4  in  Arctic 
America,  and  2  are  Holarctic. 

The  distribution  of  the  beetles  is  better  known  than  that  of  most 
of  the  orders;  9  species  recorded  from  the  Pribilofs  occur  also  in 
northwestern  North  America,  14  in  Alaska,  7  in  northern  North 
America,  and  6  in  Alaska  and  Siberia;  while  11  are  Holarctic  and 
2  are  cosmopolitan,  being  carried  about  in  commerce.  Of  the  flies 
(Diptera),  3  species  are  known  from  northwestern  North  America, 
13  from  Alaska,  12  from  northern  North  America,  and  9  are  Hol- 
arctic. 

Hymenoptera  exceed  in  number  of  species  any  other  order  of  in- 
sect on  the  Pribilofs,  but  their  distribution  is  very  imperfectly  known. 
Five  of  those  collected  elsewhere  have  been  taken  in  northern  North 
America,  5  in  Alaska,  and  9  on  other  islands  of  the  northern  Pacific, 
while  1  is  known  to  be  Holarctic. 

Summarizing  the  distributional  data  for  all  of  the  orders,  it  ap- 
pears that  the  largest  number  of  species  the  Pribilof  Islands  are 
known  to  have  in  common  with  another  region  is  35,  with  Alaska; 
30  are  Holarctic;  28  occur  more  or  less  throughout  northern  North 


132  NORTH   AMERICAN   FAUNA.  [No.  40 

America;  15  in  northwestern  North  America  (usually  including 
Alaska)  ;  14  on  other  islands  of  the  northern  Pacific;  6  in  Alaska 
and  Siberia ;  and  6  are  "  tramp  "  species,  parasitic  on  man,  or  dis- 
tributed in  commerce.  As  information  on  the  distribution  of  in- 
sects increases  it  is  probable  that  a  large  proportion  of  the  Pribilof 
fauna  will  prove  to  be  Holarctic. 

Lot  numbers  and  field  notes  of  G.  Dallas  Hanna,  from  St.  George  Island,  191  Jf. 

1.  Diptera,  90  specimens  collected  about  fox  houses  and  on  beach  among  bowl- 

ders. The  most  abundant  species,  the  very  woolly  one,  has  become 
abundant  the  past  few  days  in  the  outhouses,  and  the  fox  and  meat 
houses,  and  among  the  bowlders  on  the  beach.  The  smooth-bodied  spe- 
cies so  common  in  the  lot  sent  to  Washington  in  May  is  becoming 
scarce.     June  4. 

2.  Diptera.  30  specimens,  4  species.    In  the  grass  and  on  very  wet  soggy  ground 

near  village  landing.     June  4. 

3.  Coleoptera,  20  specimens.     Found  living  in  grass,  under  stones,  and  in  sun 

near  village  landing.     June  4. 

4.  Coleoptera,  11  specimens,  1  species.     Found  in  copulation  on  sunny  side  of 

bowlders  about  surf  line,  village  landing.  -  June  4. 

5.  Coleoptera,  17  specimens,  4  species.     In  grass  and  about  stones,  village  land- 

ing.    June  4. 
0.  Spiders.  4  specimens.     Under  stones,  village  landing.     June  4. 

7.  Diptera,  14  specimens.     Found  on  several  species  of  flowers  and  in  grass. 
When  approached  it  darts  into  the  grass  but  never  tries  to  fly  away.     Up- 
lands.    June  8. 

8.  Diptera,  17  specimens.     The  two  largest  from  flowers.    Near  village.    June  8. 

9.  Diptera,  04  specimens.     The  most  abundant  species  is  one  which  is  very 

common  along  the  beaches,  living  upon  the  decaying  marine  algae. 
Near  North  Rookery.     June  10. 

10.  Coleoptera,  5  specimens.     Crawling  over  grass  and  rocks  of  highlands.     2 

copulating,  1  eating  piece  of  dead  bird.     Near  North  Rookery.    June  10. 

11.  Diptera  (minute),  12  specimens  from  near  beach  at  East  Landing.  June  10. 

12.  Coleoptera,  23  specimens.     2  water  beetles  seen  eating  a  dead  earthworm. 

Small  very  black  beetles  found  only  on  bowlders  of  beach  near  East 
Landing.     June  10. 

14.  Coleoptera,  5  specimens.     Found  crawling  over  grass  of  bench  lands.     Not 

yet  seen  on  top  of  high  hills.     June  12. 

15.  Coleoptera,    6    specimens.     Crawling    over    grass    of    bench    lands.     Young 

found  on  wild  parsnip.     June  12. 

16.  Diptera,   4   specimens.     Found    crawling   over   grass   of   dry   bench    lands. 

Not  seen  near  bogs  or  on  top  of  high  hills.     June  12. 

17.  Diptera,  12  specimens,  long  legged.     In  wet  places,  mostly,  but  some  seen 

crawling  over  grass  far  from  water.  None  seen  flying.  Those  with 
wings  best  developed  from  Spring  Creek,  Garden  Cove.  June  14. 
One  seen  with  the  very  fuzzy  fly  in  No.  18  beneath  it;  apparently  they 
were  fighting. 

18.  Diptera,  9  specimens,  5  species.     Vicinity  of  Garden  Cove.     June  14. 

19.  Diptera,  10  specimens.     Vicinity  of  Garden  Cove.     June  14. 
21.  Neuroptera,  3  specimens  from  Garden  Cove.     June  14. 

23.  Diptera,  5  species,  15  specimens  from  Garden  Cove.     June  14. 


1923.]  INSECTS   OF    THE   PRIBILOF   ISLANDS.  133 

24.  Coleoptera,  3  species,  8  specimens  from  Garden  "Cove.     June  14. 

25.  Coleoptera,  5  specimens  from  Garden  Cove.     June  16.     Mrs.  E.  G.  Whitney, 

collector. 

26.  Diptera,  9  specimens  from  Garden  Cove.     June  16.     Mrs.  E.  G.  Whitney, 

collector. 

27.  Diptera,  5  specimens,  from   Garden  Cove,  June  16.     Mrs.  E.  G.  Whitney, 

collector. 

28.  Coleoptera,  10  specimens,  toward  East  Rookery  from  Village ;  2  females  seen 

depositing  eggs  in  wet  packed  earth  which  was  clear  of  vegetation. 
Each  had  made  a  trench  about  2  inches  long,  very  shallow,  and  was 
depositing  the  eggs  in  a  hole  in  the  trench.  The  holes  were  $  to  \ 
inch  deep.     June  16. 

29.  Coleoptera,  9  specimens.     Toward  East  Rookery  from  Village.     June  16. 

30.  Diptera,   7   specimens,    long-legged.     Toward   East    Rookery   from   Village. 

None  seen  with  wings  developed.     June  16. 

32.  Diptera,  several  species,   78   specimens    (small).     Sweepings  toward   East 

Rookery.     June  16. 

33.  Aphididae,  9  specimens.     About  East  Landing.    June  16. 

34.  Diptera,  50  specimens.     Toward  East  Rookery  from  Village.     June  16. 

35.  Diptera,  43  specimens.     Toward  East  Rookery  from  Village.     June  16. 

36.  Diptera,  24  specimens.     Several  species.     Toward  East  Rookery  from  Vil- 

lage.    June  16. 

37.  Diptera,  201  specimens   (small),  1  Hymenoptera.     Staraya  Artel  Rookery, 

June  17. 

38.  Diptera,  7  specimens.     Xeuroptera,  1.     Uplands  toward  Staraya  Artel  Rook- 

ery.    June  17. 

39.  Lepidoptera,  4  specimens    (only  5  seen).     Uplands  toward   Staraya  Artel 

Rookery.     June  17. 

40.  Diptera,  4  specimens    (all   seen)    in   bog  plants,   border  Gull  Lake  beside 

Staraya  Artel  Rookery ;  caught  on  wing.     June  17. 

41.  Diptera,  3  specimens.     Uplands  toward  Staraya  Artel  Rookery.     June  17. 

42.  Diptera,  36  specimens,  spiders  1.     Uplands  toward  Staraya  Artel.     June  17. 

43.  Aphididae,  4  specimens.     Uplands  toward  Staraya  Artel.    June  17. 

44.  Diptera,   6  specimens.     Around  a  sphagnum   bog  \   mile   west  of  Village. 

June  17. 

45.  Caterpillars,    3    specimens   with    eggs    and    cocoons    made.      Preserved    in 

alcohol.     June  18. 

46.  Insects,  30  specimens,  from  toward  East  Rookery.     June  24. 

47.  Insects  and  spiders,  30  specimens  from  toward  North  Rookery.     June  25. 

48.  Lepidoptera,  17  specimens  from  top  of  ridge,  Zapadni  Trail.     June  27. 

49.  Insects,  75  specimens  approximately.     Toward  Zapadni.     June  27. 

50.  Diptera,  13  specimens;  Coleoptera,  1.     Both  common.     Flies  feeding  about 

meat  and  fox  dung.     Toward  Zapadni.     July  4. 

51.  Lepidoptera,  5   specimens.     Cream-colored   species   is  very   rare.     Toward 

Zapadni.     July  4. 

52.  Diptera,  29  specimens.     From  toward  Zapadni  Rookery.    July  4. 

53.  10  Coleoptera  from  trail  toward  Zapadni.     July  4. 

54.  4  wood  ticks,  2  species.     From  Tolstoi  Point.     Found  under  bowlders  just 

above  surf  line  on  beach.    July  9. 

55.  50  insects,  many  species.    Small  black  beetle  is  very  common  in  some  places, 

always  found  at  surf  line  among  bowlders. 


134  NORTH   AMERICAN   FAUNA.  [No.  46 

IjOt  numbers  and  field  notes  of  Alvin  G.  Whitney  and  Elsie  G.  Whitney,  from 
St.  Paul  Island,  unless  otherunse  stated,  1912  to  191^. 

I.  3  caterpillars  in  grass.    Telegraph  Hill.    October  11,  1912. 

4.  Galls  from   Salix.     Adult  insects  had  emerged  during   September.      (Slide 

No.  5  may  be  larvae  from  willow  galls,  No.  14  of  collection).  Flat 
north   of   Telegraph   Hill.      Autumn,    1912. 

5.  9  larvae,  1  spring-tail,  found  in  bottom  of  bag  in  which  willows  and  mosses 

had  been  collected,  March  23.  St.  Paul  Island.  Southwest  Bay  to 
Southwest  Point.     March  23,  1913. 

II.  Mites.     April,  1913.  / 

14.  4  larvae.     Probably  from  galls  on  Salix.     May,  1913. 

21.  2  coleopterous  larvae.     At  roots  of  native  plant.     May  14,  1913. 

22.  5  chrysalids  of  black  and  red  caterpillars.     Caterpillars  were  collected  on 

Lukanin  Hill  about  May  1.  After  two  days  in  the  laboratory,  they 
began  to  spin  cocoons  in  this  bottle,  and  finished  within  a  day,  when 
they  were  placed  in  alcohol.     May  14,  1913. 

23.  3  gnats.     Company  house.     May  16,   1913. 

24.  2  parasitized  dipterous  flies.    1  parasitic  mite  from  one  of  the  flies.    Com- 

pany House  bathroom.     May  16,  1913. 

26.  13  dipterous  flies.     Company  House.  May  16,  1913. 

27.  3  dipterous  flies.     Company  House  bathroom.     May  16,  1913. 

28.  2  dipterous  flies.     Laboratory.     May   16,   1913. 

33.  Pupae  in  grass  stems  growing  in  shelter  of  rock  crevices.  1  hymenopterous 
fly,  which  emerged  from  one  of  these  pupae  in  warmth  of  laboratory. 
Tolstoi  Hill.     Spring    (probably  about  May  20),   1913. 

36.  10  bumblebees.     June  5,  1913. 

37.  2    (parasitized?)    gnats;   nearly  dead  when  found.     Laboratory.     June  5, 

1913. 

39.  6  gnats.     From  a  great  many  on  window,  probably  hatched  from   native 

Sagina  transplanted  to  Laboratory  a  few  days  before.  Laboratory. 
June  5,  1913. 

40.  2  crane  flies.     In  grass,  1  at  Kitovi  and  the  other  on  Reef  Peninsula.     June 

9  and  10,  1913. 

41.  1  fly  and  pupal  skin.     Pupa  collected  June  10,  hatched  June  20.     June  20, 

1913. 

42.  6   flies.     The   first   of   this    species   seen    this    season.     Sand   dunes,    Reef 

Peninsula.     June  11,  1913. 

43.  8  centipedes.     From  crevices  and  holes- in  vesicular  lava,  under  a  thin  layer 

of  soil.  Coll.  by  E.  G.  W.  Centipedes  were  found  also  near  bowlder 
beach  just  northeast  of  Little  Polovina  Rookery,  July,  1914.  (Prof. 
G.  H.  Parker  collected  specimens  on  Akutan  Island,  in  the  Aleutians, 
June,  1914.)     Gorbatch  Cliffs.    June  11,  1913. 

44.  3   fliea     Abdomens  covered   with  white  woolly   hairs.     Sand   dunes,   Reef 

Peninsula.     June  11,  1913. 

45.  2  beetles.     Sand  dunes,  Reef  Peninsula.     June  15,  1913. 

47.  4  Carabidae.     In  grass  and  moss.     Southwest  Bay.    June  17,  1913. 

49.  2  ground  beetles.     A  very  common  form.     Tolstoi  Hill.     June  18,  1913. 

50.  About  20  hymenopterous  flies  and  their  pupa-cases.     Egg  cluster  collected 

by  E.  G.  W.,  May  29,  flies  hatched  out  some  time  before  June  20,  in 
the  pill  box  in  which  collected.     May  29,  1913. 


1923.]  INSECTS   OF    THE   PRIBILOF   ISLANDS.  135 

51.  12   larvae.     Orange-colored    in   life.     Abundant  on   willow  catkins   in    the 

"pussy"  stage,  slightly  before  blossoming.  The  larvae  were  secreted 
among  the  buds  in  the  compact  heads.  These  willows  covered  with 
galls  just  beginning  to  develop.  Flat  north  of  Telegraph  Hill.  June 
22,  1913. 

52.  4  mites.     From  plants.     June,  1913. 

56.  2  weevils.     June  20,  1913. 

57.  2  flies.     Male  and  female  mating.     From  house.     June  24,  1913. 

60.  2  flies ;  1  crane  fly  ;  1  beetle ;  1  spider.  Otter  Island  (6  miles  from  St.  Paul). 
July  3,  1913. 

63.  2  flies.     Company  House.     July  6,  1913. 

64.  2  flies.     Same  2  species  as  No.  63.     Company  House.     July  6,  1913. 

66.  2  flies.  Duplicate  of  species  Nos.  64  and  65.  Several  eggs.  The  eggs  were 
ejected  by  larger  fly  in  its  struggles  to  escape  from  vial.  Company 
House.     July  6,  1913. 

68.  4  aphids.     On  Pedicularis.     July  9,  1913. 

69.  8  aphids.     On  lettuce.     Originally  on  Pedicularis  blooms  brought  to  labora- 

tory June  22,  and  escaped  to  lettuce  bed  where  they  multiplied  rapidly. 

Laboratory  hotbed.     July  10,  1913. 
74.  4  blue  flies.     Collected  by  E.  G.  W.     Zoltoi  Beach.     July  17,  1913. 
?5.  2  mites.     On  Montia  fontana.     July  17,  1913. 

77.  12  larvae.    Abundant  everywhere  around  the  roots  of  grasses  and  of  herbs, 

and  especially  under  beds  of  moss  on  the  roots  of  which  it  feeds,  killing 
the  moss  over  considerable  areas.  Under  such  a  moss  bed  I  found  as 
many  as  20  to  the  square  foot.  This  larva  is  found  over  all  the  island 
in  grassy  or  mossy  places  and  all  through  the  summer  season.  It  must 
also  be  of  considerable  ecological  importance,  because  of  its  food  value 
to  the  birds  and  foxes.  The  foxes  will  dig  over  large  areas  of  moss 
beds  to  feed  on  these  larvae.  Was  unable  to  find  the  species  in  adult 
form.  Could  not  seem  to  raise  adults  in  laboratory  by  keeping  larvae 
with  one  of  the  food  plants.  It  may  possibly  be  the  larval  form  of  the 
crane  fly,  which  is  very  abundant.  Color  not  altered  by  pickling  in 
alcohol.    St.  Paul  Island.    Reef  "  Parade  Ground."    July  18,  1913. 

78.  16  larvae.     Same  as  No.  77.     July  20,  1913. 

SO.  2  flies.    Male  and  female  mating.    Reef  Peninsula.    July  18,  1913. 
87.  7  mites.    This  form  very  abundant,  found  everywhere  in  moss  and  on  flow- 
ering plants.     July,  1913. 

90.  3  cast  larval  skins  of  crane  flies.    Hutchinson  Hill.    July  22,  1913. 

91.  20  gnats.     Big  Lake.     Swarms  of  these  insects  were  being  driven  by  the 

breeze  southeastward  off  of  the  lake.  These  clouds  of  gnats  noticeable 
throughout  July  and  August.    July  22,  1913. 

92.  10  gnats.     Same  as  No.  91.     Big  Lake.     Collected  from  our  coats  as  they 

swarmed  past  us  while  driving  along  the  lake.    July  22,  1913. 

93.  3  small  gnats.     Big  Lake.     Collected  at  same  time  with  larger  ones  from 

lake  shore.    July  22,  1913. 

94.  3  Diptera,  1  bug,  2  Neuroptera,  1  moth,  1  beetle.    Northeast  Point  Peninsula. 

July  22,  1913. 

95.  11  Hymenoptera.     Collected  by  a  native  boy  who  said  he  found  a  nest  of 

them.     July  23,  1913. 

96.  4  caterpillars.     Collected  by  a  native  September  8;  then  left  in  a  dry  box 

until  October  6,  in  the  hope  that  they  would  pupate.  None  did  so ;  sev- 
eral dried  up.  The  specimens  preserved  were  the  ones  left  living  Oato- 
ber  6.     October  6,  1913. 


136  NORTH   AMERICAN   FAUNA.  [No.  46 

98.  About  10  larvae  from  mud  of  dried-up  pond,  where  Leucosticte  had  been 
scratching  for  them.  This  pond,  about  one-fourth  mile  long  and 
eighteen  inches  deep,  was  dry  from  August  5  to  mid- September.  Dur- 
ing this  time  its  whole  mud  bottom  was  scratched  entirely  over  by 
turnstones  to  get  at  these  larvae,  which  were  abundant.  Color  of 
larvae  ruby-red.     Mud  bottom  of  Village  Pond.    August  10,  1913. 

102.  4  water  beetles.    Creek  outlet  of  Antone  Lake.     Fall,  1913. 

104.  33  mites  from  Stellaria.     September  13,  1913. 

105.  6  Mallophaga.     On  choochkie  (Aethia  pusillus).     November  4,  1913. 

106.  11  Mallophaga.     On  Aethia  pusillus.     Same  as  No.  105.    November  4,  1913. 

107.  14   Mallophaga.     On   Aethia  pusillus.     Same  as  Nos.   105   and   106.     No- 

vember   4,    1913. 

108.  Lice  on  Mus  musculus.     November  16,  1913. 

109.  Lice  on  Mus  musculus.    Duplicate  of  No.  108.    November  16,  1913. 

110.  Lice  on  Mus  musculus.    Duplicate  of  No.  108.    November  16,  1913. 

111.  6  caterpillars.    On  a  sandy  road.    Halfway  Point.    August  1,  1913. 

112.  5  caterpillars.    Collected  September  8  by  a  native,  same  as  No.  96,  left  in 

box  to  pupate,  but  dried  up  without  transforming.    October  6,  1913. 

113.  8  bumblebees.     Collected  by  a  native  who  killed  them  in  grease.     North- 

east Point.     May  31,  1913. 

116.  3  caterpillars.     About  300  feet  up  on  bare  south  cinder  slope,  where  the 

snow  was  melted  off  and  the  surface  of  the  cinders  warmed  by  the  sun. 
All  insects  in  the  wet,  half-frozen  tundra  below  were  still  dormant. 
Polovina  Hill.    March  29,  1914. 

117.  2  larvae  and  3  pupal  shells.     Found  close  together  in  moss  on  the  flat 

near  Polovina  Lake.  A  fox  was  digging  up  insects  near  by.  March 
29,  1914. 

118.  3  larvae;  3  rove  beetles.     From  moss  on  Hat  near  Polovina  Lake.    March 

29,  1914. 

119.  3  small  beetles  and  several  mites.    April  5,  1914. 

120.  2  aphids  and  one  empty  aphid  skin,  and  1  hymenopterous  fly.     The  live 

aphids  were  bluish  with  whitish  "  bloom  " ;  collected  in  moss  on  scoria 
bank  back  of  "  Company  House."  April  7,  1914.  The  fly  emerged  ap- 
parently from  one  of  these  aphids  about  April  18,  1914. 

121.  1  beetle.     Village  Hill.     April  15,  1914. 

126.  2  lice  from  Aleut  child's  hair.     April  22,  1914. 

127.  4  Mallophaga,   from   European   widgeon    shot   May   9   at   Icehouse   Lake. 

May  11,  1913. 

128.  2  beetles.     Male  and  female  mating.     From  surface   of  tide  pool  under 

Gorbatch  Cliffs.     May  13,  1914. 

129.  2  insects.     Found  jumping  like  springtails  on  the  bare  sand  dunes,  Dia- 

mond Hill  dunes.     May  16,  1914. 

131.  4  beetles.     May  22,  1914.     5  beetles  from  Zoltoi  Beach  sand  dunes.     May 

19,  1914. 

132.  6  flies ;   8  mites.     From  privy.     The  mites  were  parasitic  on   the  flies, 

sometimes  two  or  three  mites  on  a  single  host.    May  19,  1914. 

133.  3  coleopterous  larvae.    Lukanin  Hill.    May  19,  1914. 

3  coleopterous  larvae.  Zoltoi  Beach  sand  dunes.  May  22,  1914.  These 
beetle  larvae,  resembling  those  of  "  potato  beetles,"  are  exceedingly 
common  all  summer,  and  feed  on  a  large  variety  of  plants,  including 
Pedicularis.  Probably  are  the  larval  stage  of  the  species  in  No.  145 
and  the  largest  species  in  No.  189. 
135.  9  rove  beetles.    Zoltoi  Beach  sand  dunes.    May  22,  1914. 


1923.]  INSECTS   OF    THE   PRIBILOF   ISLANDS.  137 

138.  10  weevils.    May  19,  1914. 

139.  6  mites.    Parasitic  on  flies.    Same  as  in  No.  132.    May  19,  1914. 

141.  7  mites.     Parasitic  on  flies.     Same  as  Nos.  132  and  139.     May  23,  1914. 
143.  6  Diptera.    May  23,  1914. 

146.  6  beetles,  May  19,  1914.    7  beetles  from  Zoltoi  sand  dunes  May  22,  1914. 

7  beetles  from  meadow  by  villages.     July  24,  1914. 

147.  2  beetles,  May  21,  1914.    1  beetle,  Zoltoi  sand  dunes,  May  22,  1914. 

148.  22  rove  beetles.     Zoltoi  sand  dunes.     May  22,  1914. 

149.  8  click  beetles.     This  species  is  found  abundantly  under  spreading  Mer- 

tensia  maritima  plants.    Zoltoi  sand  dunes.    May  22,  1914. 

150.  39  beetles,  Lukanin  Hill,  May  19,  1914.    3  beetles,  Zoltoi  sand  dunes,  May 

22,  1914,  1  beetle,  July  24,  1914. 

152.  13  Diptera.     From  outside  of  laboratory  window.     May  23,  1914. 

153.  4  flies.     From  privy.     May  23,  1914. 

154.  4  Diptera.     May  23,  1914. 

155.  2  Diptera.     Gray  in  color,  with  2  diagonal  bands  on  under  side  of  abdomen. 

Laboratory.     May  23,  1914. 

156.  10  Diptera  and  their  parasitic  mites.     From  outhouse.     May  23,  1914. 

157.  2  ,  male  and  female    (?).     Perhaps  same  species  as  No.  129. 

Bronze-lustered,  wingless  insects,  found  among  the  moss.     Grassy  bank 
beyond  village  wells.     May  23,  1914. 

159.  6  beetles.     These  beetles  were  taken  when  a  very  heavy  wind  had  been 

blowing  all  day,  and  they  had  been  swept  into  the  hollows  among  the 
dunes.  Their  wings  protruded  from  under  the  half-expanded  elytra, 
indicating  that  they  had  been  in  flight.  I  have  never  seen  any  other 
beetles  on  the  Pribilofs  that  could  fly.  This  was  the  only  time  I  saw 
this  species,  which  was  fairly  abundant  in  the  one  locality  this  one 
day.  I  saw  perhaps  twenty  or  more  in  the  sand  hollows.  They  were 
velvety  brown  in  color,  with  blotches  of  darker  brown,  and  when 
touched  drew  their  legs  into  grooves  on  the  body,  so  they  were  then 
as  smooth  as  a  bean.     (Same  as  No.  166.)     Zoltoi  sand  dunes.     May 

23,  1914. 

160.  2   carabid    (?)    larvae.     In    moss.     Grassy    bankside   near   village   wells. 

May  23,  1914. 

167.  5  beetles.    May,  1914. 

168.  12  gnats,  flying  in  a  swarm  by  laboratory.     Spring,  1914. 

169.  1  bed  bug,  from  native  boy's  clothes.     2  (?).     Dark  bluish  in 

life,  except  the  young  ones,  which  are  white.  Probably  same  species 
as  Nos.  13  and  200.  These  insects  are  common  in  damp  herbage,  espe- 
cially in  moss  beds,  and  are  sometimes  seen  in  myriads  in  crevices  of 
low  ledges  along  the  shores  and  crowded  together  on  the  surface  of 
tide  pools.     April,  1914. 

170.  About  25  crane  flies.     1  harvestman.     Crane  flies  were  crawling  every- 

where at  this  time,  and  many  were  mating.  Tolstoi  sand  dunes. 
About  June  1,  1914. 

173.  2  beetles.     Male  and  female,  mating.     (Probably  same  species  as  Nos.  180 

and  181.)     Ledges,  Kitovi  Rookery.     June  16,  1914. 

174.  3  mites,  1  moth,  1  hymenopteron,  1  crane  fly.     Spring,  1914. 

176.  1  fly  and  several  pupa  cases.  The  pupa  cases  were  found  June  14  in  the 
interstices  of  the  nasal  bones  of  a  fur  seal  skull  on  the  killing  field. 
From  these  one  fly  hatched  out  in  the  vial  June  20.  Near  village. 
June  20,  1914. 


138  NORTH   AMERICAN   FAUNA.  [No.  46 

180.  12  beetles.     From  crevices  and  face  of  cliffs  on  southwest  side  of  Sealion 

Rock,  an  islet  less  than  %  mile  long  and  about  i  mile  from  St.  Paul 
Island.  The  center  of  the  island  is  about  60  ft.  high  and  supports  a 
little  grass  (Glyceric/,)  and  a  few  herbs.  The  island  is  similar  in 
character  to  Walrus  Island  (12  miles  distant),  where  a  rare  species 
of  Coleoptera  exists.     Possibly  this  is  the  same  species.     June  29,  1914. 

181.  44  beetles ;  2  beetles ;  larvae.     From  face  of  cliffs  and  crevices  of  rocks 

on  southwest  side  of  island.  Same  as  No.  180,  and  probably  same  spe- 
cies as  No.  173.    Sealion  Rock.    June  29.  1914. 

182.  10  Diptera.     From  grassy  summit,  60  ft.  high.     Sealion  Rock.     June  29, 

1914.  / 

183.  5  moths.  June,  1914. 

184.  3  moths.  June,  1914. 

185.  2  mites.  From  Lunda  cirrhata.    Color,  pale  blue.    June  30,  1914. 

187.  56  beetles   (several  species).     Collected  from  mud  shore  of  village  pond. 

Many  were  mating.    June  30,  1914. 

188.  60  carabid  beetles.     Grass-covered  upland.  East  Landing  to  village  wells. 

These  carabids  are  abundant  and  especially  active  and  noticeable 
during  June  and  July.    June,  1914. 

189.  181  beetles  of  several  species.    Near  village.    June,  1914. 

190.  2  coleopterous  larvae.     June,  1914. 

191.  5  mites.     On  Rissa  t.  pollicaris.    July  4,  1914. 

192.  4  Mallophaga.    On  Rissa  t.  pollicaris.    These  were  from  the  same  bird  as 

the  mites  in  No.  191,  namely,  E.  A.  Preble's  catalog  No.  2239.  July  4, 
1914. 

193.  Lice  on  Sorex  pribilofensis.    July  5,  1914. 

194.  Lice  on  Sorex  pribilofensis.     (Same  as  193.)     July  5,  1914. 

195.  55  Diptera.    This  species  was  very  abundant  and  active  on  Chrysanthemum 

arcticum  flowers  (in  full  bloom  on  this  date)  in  salt  marsh  on  north 
side  of  Salt  Lagoon.  Do  not  think  I've  seen  it  elsewhere  on  the 
Pribilofs.     "  Salt  Lagoon  "  marsh.     July  24,  1914. 

196.  4  rove  beetles.     July  24,  1914. 

197.  4  white  larvae.    1  immature  click  beetle.     (From  underside  of  moss  bed.) 

Reef  "  Parade  Ground."    July  28,  1914. 

200.  2  .     (Probably  same  species  as  Nos.  IS  and  169.)     Found  in 

moss.     Spring,  1914. 

201.  2  click  beetles,  1  small  beetle.     Summer,  1914. 
204.  3  larvae,  1  beetle.     Summer,  1914. 

210.  7  Diptera.     Summer,  1914. 
212.  3  beetles.     Summer,  1914. 


APTERYGOTA.1 

By  W.  L.  McAtee,  Bureau  of  Biological  Survey. 

Family  PODURIDAE. 

Neanura  gigantea  Tullberg. 

Anura  gigantea,  Tullberg,  Tycho.  Collembola  borealia-Nordiska  Collembola. 
Ofversigt  af.  Kongl.  Vetenskaps  Akaderuiens  Forhandlingar,  No.  5,  p.  41, 
PI.  XI,  fig.  59.     1876.     [Siberia.] 

Two  lots  collected  on  St.  Paul  Island  in  spring  of  1914  by  A.  G. 
Whitney,  who  says :  "Dark  bluish  in  life,  except  the  young  which  are 
white.  These  insects  are  common  in  damp  herbage,  especially  in  moss 
beds,  and  are  sometimes  seen  in  myriads  in  crevices  of  low  ledges 
along  the  shore  and  crowded  together  on  the  surface  of  the  tide  pools." 

Aphorura  dentata  Folsom. 

Aphorura  dentata  Folsom,  J.  W.,  Proc.  Washington  Acad.  Sci.,  vol.  4,  pp. 
91-92,  PI.  VII,  figs.  29-36,  1902. 

Originally  described  from  material  including  one  specimen  collected 
on  St.  Paul  Island,  August  1,  1897. 

Isotoma  viridis  Bourlet. 

Isotoma  viridis  Bourlet.  Memoire  sur  les  Podures.  Memoires  de  la  Societe 
Royale  des  Sciences,  de  1' Agriculture  et  des  Arts,  p.  401,  Lille,  1839. 
[France.] 

Two  lots  collected  on  St.  George  Island,  June  8,  10,  1914,  by  G.  D. 
Hanna,  and  two  from  St.  Paul  Island,  by  A.  G.  Whitney,  one  collected 
May  19,  1914.     The  remark,  "  common  in  tundra  moss  "  accompanies 
last  lot. 
Isotoma  violacea  Tullberg  var.  mucronata  Axelson. 

Isotoma  violacea  Tullberg,  Tycho.  op.  cit,  p.  36.     [Siberia.] 
Isotoma  violacea  Tullberg  var.  mucronata  Axelson,  W.  M.  Vorlaufige  Mit- 
theilung   fiber  einige  neue  Collembolen-Formen  aus   Finnland.      Medde- 
landen  af  Societas  pro  Fauna  et  Flora   Fennica,  36,  p.   118,  1899-1900 
(1900).     [Finland.] 

Several  specimens  of  this  variety  representing  a  species  previously 
known,  according  to  Dr.  J.  W.  Folsom,  only  from  Norway,  Sweden, 
Finland,  and  Greenland,  were  found  in  the  stomach  of  a  Leucosticte 
griseonud'ha,  collected  on  St.  George  Island,  August  2,  1920. 

1  The  insects  of  this  order  were  identified  by  Dr.  J.  W.  Folsom.  He  has  recorded  from 
the  Pribilofs  three  of  the  species  here  mentioned,  in  his  admirable  report  on  the  Apterygota 
in  Papers  from  the  Harriman  Alaska  Expedition,  XXVII,  Proc.  Washington  Acad.  Sci., 
vol.  4,  pp.  87-116,  pis.  4-7,  March  27,  1902. 

139 


ORTHOPTERA. 

By  Morgan  Hebard,  Academy  of  Natural  Sciences  of  Philadelphia. 

/ 
Family  BLATTIDAE. 

Subfamily  Panchlorinae. 

Panchlora  cubensis  Saussure. 

P[anchlora]  cubensis  Saussure,  Rev.  et  Mag.  de  Zool.,  (2),  vol.  14,  p.  230, 
1862  [2,  Cuba]. 

A  single  green  cockroach  (  $  )  collected  on  St.  Paul  Island,  in 
1916,  by  G.  Dallas  Hanna  apparently  represents  this  common  and 
widely  distributed  tropical  American  species,  which  is  frequently 
introduced  into  the  temperate  regions  of  North  America  in  bananas. 
As  the  northernmost  point  at  which  a  species  of  the  genus  is  known 
to  be  native  is  Brownsville,  Tex.,  there  is  no  doubt  that  the  present 
is  an  adventive  specimen,  having  been  transported  to  the  Pribilofs 
on  board  ship. 

It  agrees  fully  with  West  Indian  material  before  me,  except  in 
having  the  interval  between  the  eyes  exceptionally  wide  (.5  milli- 
meter) for  females  of  the  species,  nearly  three-quarters  the  occipital 
ocular  width.  The  interocular  width  is  subject  to  variation  in  the 
species,  but  in  the  great  majority  of  females  approximates  one-third 
the  occipital  ocular  width.  For  a  detailed  discussion  of  cubensis 
see  Mem.  Amer.  Ent.  Soc,  No.  2,  p.  198  (1917). 


Blatella  germanica  Linnaeus  is  recorded  as  having  been  Introduced  into  the  Pribilof 
Islands  at  least  twice  and  a  preserved  specimen  has  been  reported  from  St.  George.  ( Van 
Duzee,  E.  P.,  Proc.  Cattf.  Acad.  Sci.,  Fourth  Ser.,  11,  p.  193,  Nov.  1921.)— W.  L.  M. 

140 


MALLOPHAGA. 

By  G.  F.  Ferris,  Assistant  Entomologist,  Stanford  University. 

All  of  the  material  here  reported  on  was  collected  by  A.  G.  Whitney 
on  St.  Paul  Island.  While  the  collection  is  small  it  contains  two 
very  interesting  records,  two  of  the  species  not  having  been  recorded 
previously  from  North  America,  their  host  records  also  being  new. 

Docophorus  lari  Denny. 

Four  specimens  of  this  widely  spread  gull-infesting  species  from 
the  Pacific  kittiwake,  Rissa  tridactyla  poUicaris.  It  has  previously 
been  recorded  from  the  same  host. 

Docophorus  merguli  Denny. 

Several  specimens  from  the  least  auklet,  Aethia  pusilla.  This 
species  has  previously  been  recorded  but  twice,  both  times  from  the 
little  auk  or  rotchie,  Alle  (Mergulus)  alle,  of  Europe.  The  specimens 
at  hand  agree  very  well  with  specimens  from  the  latter,  sent  us  by 
Mr.  Waterston. 

Nirmus  maritimus  Kellogg  and  Chapman. 

Several  specimens  from  Aethia  pusilla.  This  record  also  is  new. 
Another  species  of  Nhvnus  (N.  citrinus  Nitzsch)  has  been  recorded 
from  the  same  host  and  this  record  may,  perhaps,  refer  to  the  same 
species  as  the  two  are  not  very  different. 

Menopon  lutescens  Nitzsch. 

Several  specimens  from  Aethia  pusilla.  This  is  also  a  new  record, 
both  as  to  host  and  locality,  the  species  previously  having  been 
recorded  from  Alca  tor  da  and  Alle  (Mergulus)  alle  of  Europe. 
The  determination  of  any  species  of  Menopon  is  always  attended  with 
uncertainty,  but  this  species  has  been  figured  by  Waterston  (Proc. 
Eoyal  Phys.  Soc.  Edinburgh,  vol.  18,  No.  4,  pp.  266-267,  f .  3,  1912) , 
and  as  it  is  apparently  rather  characteristic  of  the  auklet  group  the 
determination  is  reasonably  safe. 

Trinoton  luridum  Nitzsch. 

Four  specimens  of  this  common  duck-infesting  species  from  the 
European  widgeon,  Mareca  penelope.  It  has  previously  been  re- 
corded from  the  same  host. 

108731°— 23 10  141 


ANOPLTJRA. 

By  W.  L.  McAtee,  Bureau  of  Biological  Survey. 

Family  PEDICULIDAE. 
/ 
Pediculus  capitis  De  Geer. 

Pediculus   (humanus  capitis),  De  Geer,  Charles,  Memoires  pour  servir  a 
L'Histoire  des  Insectes,  Tome  Septieme,  p.  67,  1778. 

Five  specimens  are  in  the  collection,  all  collected  from  the  heads 
of  Aleuts  on  St.  Paul  Island  in  April  and  in  "  summer." 

Family  ECHINOPHTHIRIIDAE. 

Antarctophthirus  callorhini  Osborn. 

Haematopinus  callorhini,  Osborn,  Herbert,  The  Fur  Seals  and  Fur  Seal 
Islands  of  the  North  Pacific  Ocean.     Part  3,  p.  553,  1899. 

This  species  was  originally  described  from  specimens  collected  on 
fur  seals  from  the  Pribilof  Islands. 

Echinophthirius  fluctus  Ferris. 

Echinophthirius  fluctus,  Ferris,  G.  F.,  Anoplura  from  Sea-Lions  of  the  Pa- 
cific Ocean.     Ent.  News,  vol.  27,  no.  8,  pp.  366-370,  Oct.  1916. 

Mr.  Ferris  kindly  gives  me  permission  to  publish  the  fact  that  this 
species,  originally  described  from  the  Steller  sea  lion,  has  also  been 
taken  on  the  fur  seal,  and  must  therefore  occur  on  the  Pribilofs. 

142 


HOMOPTERA. 

By  Edith  M.  Patch,  State  Entomologist,  Maine  Agricultural  Experiment  Station. 

Family  APHIDIDAE. 
(Plate  VIII.) 

Macrosiphum  constrictum,  new  species. 

Alate  viviparous  female. — Beak  short,  reaching  second  coxa.  An- 
tenna 3.09  mm.  in  total  length  with  joints  measuring:  I,  .13  mm. ;  II, 
.07  mm. ;  III,  .78  mm. ;  IV,  .49  mm. ;  V,  .53  mm. ;  VI,  base  including 
sensoria  .22  m.,  spur  .87  mm.  Ill  with  nine  sensoria  in  a  row.  Cor- 
nicle .65  mm.  long,  slightly  but  distinctly  swollen  at  middle  of  distal 
half  and  constricted  a  little  near  the  tip,  where  it  is  marked  by  faint 
reticulations  for  about  .04  mm.  There  is  nothing  particularly  dis- 
tinctive about  the  venation  of  the  wing,  which  is  3.55  mm.  in  length. 

Apterous  viviparous  female. — Beak  short,  reaching  second  coxa. 
Antennae  from  two  individuals  were  measured,  one  of  which  was 
2.72  mm.  long  with  joints  as  follows:  I,  .14  mm.;  II,  .08  mm.;  Ill, 
.7  mm. ;  IV,  .51  mm. ;  V,  .5  mm. ;  VI,  base  including  sensoria  .22  mm., 
spur  .57  mm.  This  antenna  had  nine  sensoria  on  III.  The  other 
antenna  measured  2.93  mm.  with  the  joints  as  follows :  I,  .14  mm. ; 
II,  .10  mm. ;  III,  .72  mm. ;  IV,  .42  mm. ;  V,  .49  mm.  ;VI,  base  includ- 
ing sensoria  .25  mm.,  spur  .81  mm.  Ill  in  this  case  with  six  sensoria. 
Cornicle  .63  mm.  long,  with  shape  and  reticulations  like  that  of  the 
alate  female  except  that  the  bulge  of  the  distal  half  is  slightly  more 
pronounced. 

Apterous  oviparous  female. — Beak  short,  reaching  second  coxa. 
Antenna  2.28  mm.  in  total  length,  with  the  joints  measuring:  I,  .14 
mm. ;  II,  .09  mm. ;  III,  .6  mm. ;  IV,  .38  mm. ;  V.  -39  mm. ;  VI,  base 
including  sensoria  .2  mm.,  spur  .48  mm.  Sensoria  of  III  variable  in 
number.  One  individual  had  two  on  one  side  and  three  on  the  other. 
Cornicle  .57  mm.  long  with  shape  and  reticulation  similar  to  those  of 
the  viviparous  females.  The  tarsus  to  base  of  claw  is  .15  mm.  long. 
The  hind  tibia  is  not  perceptibly  swollen. 

As  aphids  vary  somewhat  in  size  in  different  collections  and  as 
the  number  of  antennal  sensoria  is  subject  to  fluctuation,  the  fore- 
going items  should  be  taken  as  approximate  rather  than  absolute. 

Described  from  one  alate  viviparous,  two  apterous  viviparous,  and 
two  apterous  oviparous  females. 

143 


144  NORTH   AMERICAN   FAUNA.  [No.  46 

Cotypes. — Locality  St.  Paul  Island.  Collection  (Lot  No.  69)  taken 
by  A.  G.  Whitney,  June  22,  1913,  from  Pedicularis,  and  escaped  from 
the  laboratory  to  lettuce  bed,  where  they  multiplied  rapidly.  Speci- 
mens removed  from  lettuce  July  10  comprised  one  apterous  and  one 
alate  female  and  some  nymphs.  Collection  (Lot  No.  68)  taken  by 
A.  G.  Whitney  from  Pedicularis  comprised  two  apterous  oviparous 
females  and  one  apterous  viviparous. 

Metatype,  collected  by  A.  G.  Whitney,  St.  Paul  Island,  on  Saxi- 
fraga,  spring  1914,  comprised  one  nymph  (Lot  No.  164). 

Ideotypes  collected  by  G.  D.  Hanna,  St.  George  Island,  June  16 
(Lot  No.  33)  and  June  17  (Lot  No.  43),  1914.  Nymphs  only.  Food 
plant  not  recorded. 

This  species  shows  certain  resemblances  to  Macrosiphum  antir- 
rhinum (Macchiati)  as  described  and  figured  by  Theobald,1  but  the 
short  beak  of  M ,  constrictum  and  the  antennal  sensoria  of  the  ap- 
terous viviparous  female  and  several  minor  differences  serve  to  dis- 
tinguish it.  The  most  striking  characters  of  M.  constrictum  are  the 
slight  dilation  of  the  cornicle,  with  its  very  restricted  area  of  faint 
distal  reticulation ;  the  short  beak ;  the  long  slender  basal  portion  of 
antennal  joint  VI  and  the  short-pointed  antennal  setae. 


Another  species  of  plant  louse,  Nectarophora  insularis,  was  described  from  St.  Paul 
Island  by  Theo.  Pergande  (Proc.  Washington  Acad.  Sci.,  vol.  2,  p.  515,  Dec.  20,  1900)  ; 
and  a  fulgoroid  leaf-hopper,  Delphax  stejnegeri  Ashmead,  originally  described  from  Bering 
Island,  has  been  recorded  as  occurring  on  the  Pribilofs  (Ashmead.  W.  H.,  Harriman  Alaska 
Expedition,  vol.  8,  p.  130,  1904). — W.  L.  M. 


Explanation  of  Plate  VIII. 

Details  of  Plant  Louse  (Macrosiphum  constrictum). 

Fig.  1.  Antenna  of  alate  female. 

Fig.  2.  Antenna  of  apterous  viviparous  female. 

Fig.  3.  Antenna  and  tarsus  of  apterous  oviparous  female. 

Fig.  4.  Cornicle  of  apterous  viviparous  female. 

Fig.  5.  Cornicle  of  apterous  oviparous  female. 

Fig.  6.  Cornicle  of  alate  female. 

Fig.  7.  Wing  of  female. 

1  Theobald,  Fred  V.    The  British  Species  of  the  Genus  Macrosiphum.     PL  II,  Jour.  Econ. 
Biol.,  vol.  8,  p.  151,  1913. 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey.  PLATE   VI  I  I. 


Details  of  Plant  Louse  (Macrosiphum  constrictum 

(Explanation  on  page  144.) 


HETEROPTERA. 

By  W.  L.  McAtee,  Bureau  of  Biological  Survey. 

Family    MIRIDAE. 
Irbisia  sericans  Stal. 

Leptomerocoris  sericans  Stal.  C.  Beitrag  zur  Hemipteren-Fauna  Siberiens 
und   des  Russischen  Nord-Amerika.     Entomologische  Zeitung    (Stettin) 

19,  p.  188,  1858.     [Sitka.] 

Irbisia  (Leptomerocoris)  sericans  Stal.  Heidemann,  O.,  Papers  from  the 
Harriman  Alaska  Expedition,  vol.  13,  Entomological  Results  (7)  ; 
Heteroptera.     Proc.    Washington  Acad.   Sci.,   vol.   2,   p.   504,   December 

20,  1900. 

This,  the  only  species  of  Heteroptera  in  the  collection,  has  a  wide 
distribution  in  Alaska  and  on  the  neighboring  islands,  and  is  known 
to  occur  as  far  south  as  Oregon.  The  specimens  at  hand  were  ob- 
tained on  St.  Paul  Island  in  June,  July,  and  September,  and  on  St. 
George  Island  in  August. 


Orthoceplialus  saltator  Hahn.  also  of  the  family  Miridae,  was  collected  on  St.  Paul 
Island  by  Barrett-Hamilton  (Schwarz,  E.  A.,  the  Fur  Seals  and  Fur-Seal  Islands  of  the 
North  F-aciflc  Ocean,  Part  3,  p.  552,  1899).  Cimew  lectularius  Linnaeus  also  is  known  to 
be  present  on  the  islands. 

145 


TBICHOPTERA. 

By  Nathan  Banks,  Museum  of  Comparative  Zoology, 
Harvard' 'University. 

Family    LIMNEPHILIDAE. 

Limnephilus  kincaidi  Banks. 

Two  from  St.  George  Island,  August  4  and  September,  and  one 
apparently  this  species  from  St.  Paul,  August  17.  Originally  de- 
scribed from  St.  George  Island. 

Arctoecia  consocia  Walker. 

One  from  St.  Paul  Island,  August  16. 

Asynarchus  simplex  Banks. 

Three  from  St.  Paul  Island,  July  14  and  22,  and  one  apparently 
this  species,  from  St.  George,  August  4.  Originally  described  from 
St.  Paul  Island. 

A  description  of  the  larva  of  this  species  from  the  Pribilofs,  has  been  published  by 
R.  A.  Muttkowski,  Bui.  Wisconsin  Nat.  Hist.  Soc,  vol.  13,  N.  S.,  No.  1,  pp.  42-45, 
March.  1915. — W.  L.  M. 

Chilostigma  praeterita  Walker. 

Three  from  St.  George  Island,  June  17,  and  St.  Paul  Island,  May 
1,  and  "  Summer."  In  these  specimens  the  bristle-bearing  areas  or 
the  posterior  warts,  prothoracic  lobes,  mesothoracic  stripes,  and 
tegulae  are  yellowish  or  reddish.  The  typical  form  was  described 
as  having  these  black,  but  they  are  pale  in  some  of  my  European 
specimens,  and  I  believe  are  normally  pale;  sometimes  drying  out 
dark. 


In  addition  to  the  above-mentioned  species,  Anabolia  simplex  Banks  has  been  recorded 
from  St.  Paul  Island.      (Van  Duzee,  E.  P.,  Proc.  Calif.  Acad.  Sci.  Fourth  Ser.,  11,  p.   193, 
Nov.,   1921.) — W.  L.  M. 
146 


LEPIDOPTERA. 

By  Wm.  T.  M.  Fokbes,  Department  of  Entomology, 
Cornell  University. 

There  are  nine  species  represented  in  this  little  collection,  besides 
some  larvae  which  can  not  be  surely  identified.  The  complete 
absence  of  butterflies  is  surprising,  but  may  perhaps  be  explained 
by  some  peculiarity  of  the  climate.  The  same  thing  occurs  in  Ice- 
land, although  as  a  rule  butterflies  are  common  in  the  Arctic  regions. 

There  seems  to  be  a  slight  tendency  for  the  two  islands,  St.  George 
and  St.  Paul,  to  have  different  local  forms,  but  it  is  not  marked 
enough  to  be  certain.  One  specimen  of  Hyphoraia  subnebulosa  from 
St.  George  is  exceptionally  pale  and  shows  some  differences  of  mark- 
ing but  the  other  is  like  those  from  St.  Paul.  Psychophora  sabinii 
is  represented  by  a  suffused  specimen  from  St.  Paul,  while  all  those 
in  good  condition  from  St.  George  show  crisp  clean-cut  markings, 
but  such  suffusion  occurs  commonly  in  the  Arctic  and  is  likely  to  be 
an  individual  rather  than  a  racial  character. 

Family  ARCTIIDAE. 
Hyphoraia  subnebulosa  Dyar. 

Three  from  St.  Paul,  June;  2,  St.  George,  July.  The  type,  in  the 
U.  S.  National  Museum,  comes  well  within  the  range  of  this  series, 
but  is  in  poor  condition.  The  thorax  of  the  type  in  particular  is 
beyond  description.  As  the  specimens  in  this  series  show  it,  the  col- 
lar is  yellow,  with  some  black  hairs  along  the  posterior  edge,  the  disk 
of  the  thorax  is  red-brown,  concolorous  with  the  wings,  edged  on  each 
side  by  a  broad  yellow  stripe,  each  side  of  which  the  black  under- 
scaling  shows  more  or  less  distinctly  as  a  black  line.  The  upper 
two-thirds  of  the  tegulae  is  chocolate  brown,  while  the  part  just  over 
the  base  of  the  wing  is  black  in  front  and  yellow  behind.  The  hair 
is  loose  and  erect  in  the  male,  but  in  the  female  the  vestiture  of  both 
body  and  legs  is  smooth  and  close,  like  that  of  Apantesis.  The 
series  in  the  National  Museum  (from  a  variety  of  places)  indicates 
that  there  may  be  a  large  number  of  minor  local  forms. 

Family  NOCTUIDAE. 

Agrotiphila  alaskae  Grote. 

Three  males  and  one  female,  from  St.  George,  appear  to  be  of 
this  species.    The  males  are  normal  enough.    The  female  has  reduced, 

147 


148  NORTH  AMERICAN  FAUNA.  [No.  46 

lanceolate  wings  of  half  the  area  of  the  male's,  with  no  black  mark- 
ings at  all.  The  specimen  from  St-  George  is  marked  in  two  shades 
of  pale  olive,  but  one  from  Popof  Island  in  the  National  Museum  is 
purple-brown.  The  antennae  are  serrate  and  fasciculate,  not  simple 
as  Hampson  describes  them.  Superficially  the  moth  could  be  easily 
mistaken  for  Pachnohia  wockei. 

Anarta  richardsoni  Curtis. 

One  from  St.  Paul.  The  hair  on  the  deeply  sunken  eyes  is  sparse 
and  easily  overlooked.  / 

Family   GEOMETRIDAE. 

Psychophora  sabinii  Curtis.      {ScinneHa  Dyar). 

There  can  be  no  doubt  that  this  is  P.  sabinii  of  Curtis,  as  has  been 
most  generally  believed  and  as  has  been  specifically  noted  by  Barnes 
and  McDunnough  since  the  preparation  of  this  manuscript.  They 
have  proposed  for  it  the  genus  Barrovia.  "Psychophora  "  fasciata  has 
nothing  to  do  with  this  and  no  special  resemblance  to  Curtis's  figure. 
In  it  the  pectinations  of  the  antennae  are  as  broad  at  the  base  as 
farther  out,  while  in  our  form  as  well  as  in  Curtis's  figure  they  are 
shown  as  narrowing  at  the  point  of  attachment  and  apparently 
articulated  to  the  shaft.  In  fasciata  the  fringe  is  dark  at  the  base 
with  a  white  tip,  the  hind  wing  is  dark-veined  without  transverse 
lines,  the  fore  wing  has  no  dark  shade  at  the  base;  the  t.  a.  line  is 
angled  on  the  cell  only,  while  the  t.  p.  line  is  not  at  all  toothed.  The 
antennae  are  serrate  at  the  tip.  In  all  these  points  fasciata  differs 
from  Curtis's  figures  and  from  the  specimens  in  this  lot.  Fasciata 
is  a  noctuid,  as  shown  by  the  venation  and  the  large  ocelli. 

The  long  series  from  St.  George  (23  specimens,  part  of  which  look 
as  if  they  were  collected  in  alcohol)  are  very  crisply  marked  and 
look  like  Trichochlamys  polata.  Some  of  them  tend  to  show  the 
four  pale  ovals  in  the  median  area  which  Curtis  figures,  and  one  is 
chocolate  brown  with  contrasting  white-filled  lines.  There  is  some 
tendency  to  suffusion,  especially  in  the  St.  Paul  specimen,  but  noth- 
ing like  the  forms  frigidaria  and  polaris. 

Family  PYRALIDAE. 
Titanio  sp. 
St.  Paul. 

Phlyctaenia  sp. 

Too  poor  to  identify  with  certainty.  It  may  possibly  be  P.  wash- 
ingtonialls  Grote.    St.  George. 


1923.]  INSECTS   OF   THE   PRIBILOF  ISLANDS.  149 

Family  PTEROPHORIDAE. 
Platyptilia  sp. 

It  seems  too  far  north  for  the  Californian  P.  modesta,  but  there 
seems  to  be  nothing  else  as  close  to  it  among  the  species  of  Platyptilia 
with  obsolescent  scale  tuft.  P.  pterodactyla  as  described  by  Walker 
and  as  figured  by  Walsingham  is  a  paler  species  with  a  pure  white 
area  and  a  couple  of  black  dashes  on  the  first  feather,  while  this  spe- 
cies is  almost  immaculate  mouse-gray.  The  usual  black  scale-tuft  in 
the  third  feather  of  the  hind  wing  is  represented  by  four  or  five  scat- 
tered scales.    Three  from  St.  George.    August. 

Family  TORTRICIDAE. 

Sparganothis  moeschleriana  Wocke. 

All  the  specimens  I  have  seen  of  this  species  show  the  venation 
and  other  structures  of  Sparganotkis,  group  C'enopis,  and  the  mark- 
ings would  indicate  the  same  reference;  1,  St.  George,  August. 

TINEID. 
Family  Undetermined. 
Too  poor  to  name,  2  from  St.  George,  June. 


E.  P.  Van  Duzee  lists  from  the  Pribilofs  the  pyralid,  Phlyctaenia  washingtonalis  Grote, 
the  tortricid,  Argyroploce  schulziana  Fabricius  (with  a  query),  and  the  oecophorid,  Bork- 
hausenia  pseudospretella  Stainton.  (Proc.  Calif.  Acad.  Sci.,  Fourth  Ser.,  11,  pp.  194-5, 
Nov.  1921.)  H.  W.  Elliott  stated  (Report  on  the  Seal  Islands  of  Alaska,  Kept.  10th  Cen- 
sus, Vol.  8,  1884,  p.  12)  that  "  a  very  few  species  of  butterflies,  principally  the  yellow 
Nymphalidae,  are  represented  by  numerous  individuals."  However,  no  butterflies  have 
thus  far  been  collected  on  the  Pribilofs,  and  it  is  likely  that  Elliott's  note  was  merely 
from  recollection,  and  perhaps  bad  reference  to  the  rather  profusely  yellow-marked 
Hyphoraia. — W.  L.  M. 


COLEOPTERA. 

By  Prof.  H.  F.  Wickham, department  of  Applied  Zoology, 
University  of  Iowa. 

The  present  collection  of  Coleoptera  is  probably  by  far  the  most 
complete  of  any  ever  brought  out  from  the  Pribilof  Islands.  Many 
of  the  species  were  obtained  in  large  series,  exhibiting  wide  varia- 
tion and  giving  valuable  information  as  to  time  of  occurrence.  The 
fact  that  both  St.  George  and  St.  Paul  were  consistently  worked 
for  material  enables  us  to  form  an  opinion  as  to  the  faunal  relations 
of  these  islands  with  each  other;  and  finally  it  should  be  noted  that 
a  few  species  were  obtained  that  had  not  previously  been  reported 
from  Alaska. 

Two  features  stand  out  very  clearly  from  examination  of  the  mate- 
rial and  data — first  the  long  season  over  which  the  adults  of  many 
species  may  be  found,  and  second,  the  great  individual  variation. 
Color,  sculpture,  size,  and  even  outline  appear  to  have  become  in- 
constant as  if  the  restraints  which  ordinarily  hold  the  species  within 
narrow  limits  had  been  relaxed.  An  explanation  of  this  variability 
which  seems  to  me  probably  fundamental  was  suggested  by  Alexan- 
der Wetmore,  of  the  Biological  Survey — namely,  close  interbreed- 
ing brought  about  by  the  narrow  quarters  to  which  these  insects 
are  confined.  A  contributing  cause  may  perhaps  be  found  in  the 
rigorous  climatic  conditions  which  must  often  subject  the  forming 
chitinous  exoskeleton  of  the  newly  emerged  adult  to  severe  physio- 
logical stress,  resulting  in  modifications  of  the  surface  sculpture  and 
possibly  even  of  its  texture.  Eetardation  or  hastening  of  evapora- 
tion is  known  to  affect  the  intimate  sculpture  of  the  tegument  in 
semiarid  districts  and  may  well  have  an  influence  here. 

Several  species  of  beetles  have  been  found  upon  one  of  these  islands 
which  are  not  yet  known  to  occur  upon  the  other,  but  I  do  not  see 
any  evidence  of  the  development  of  different  races  where  a  given 
species  inhabits  both  St.  Paul  and  St.  George,  even  though  a  large 
percentage  of  the  Coleopterous  fauna  is  incapable  of  flight  and 
probably  rarely  crosses  the  distance  of  forty  miles  or  so  between 
the  land  areas.  A  few  cosmopolitan  beetles  are  becoming  intro- 
duced by  commerce. 
150 


1923.]  INSECTS  OF  THE  PRIBIDOF  ISLANDS.  151 

Family  CARABIDAE. 

Genus  Carabus  Linnaeus. 

Carabus  truncaticollis  Eschseholtz. 

Numerous  specimens  are  contained  in  the  collections  from  St. 
Paul  and  St.  George.  Those  from  the  former  island  were  found  in 
every  month  from  May  to  October,  and  those  from  the  latter  island 
from  April  14  to  September  10.  Taken  as  a  whole,  the  series  ranges 
from  a  vivid  green  through  duller  and  bronze  greens,  red  or  copper 
bronzes  to  a  brown  bronze.  In  general  the  greens  predominate,  but 
the  dates  do  not  indicate  any  relation  between  season  and  color. 
Neither  is  there  any  correlation  between  color  and  locality,  as  speci- 
mens from  either  island  show  practically  all  of  the  intergrades. 
The  legs  vary  from  reddish  yellow,  with  dark  tarsi  and  tibial  apices, 
to  piceous.  The  size  runs  from  15.5  mm.  to  20  mm.  Several  larvae, 
apparently  about  fully  grown,  are  dated  May  23  and  June. 

Genus  Nebria  Latreille. 

Nebria  bifaria  Mannerheim. 

Three  specimens,  St.  Paul,  without  date  other  than  "  Summer, 
1914."  Already  known  from  this  island  as  well  as  from  Alaska, 
Siberia,  and  Kamchatka. 

Genus  Pelophila  Dejean. 

Pelophila  eschscholtzii  Mannerheim. 

Three  specimens,  St.  Paul,  one  without  date,  the  other  two  taken 
August  16;  numerous  specimens  June  30.  The  first-mentioned  ex- 
ample is  distinctly  metallic  in  color  with  greenish  elytral  border; 
the  others  are  nearly  black  above,  the  elytra  castaneous.  Great 
variation  is  shown  in  number  and  position  of  the  elytral  foveae, 
even  on  the  two  sides  of  the  same  individual. 

Genus  Patrobus  Dejean. 

Patrobus  septentrionis  Dejean. 

St.  George  and  St.  Paul.  The  dates  run  through  every  month 
from  June  25  to  September  3.     A  common  and  widespread  species. 

Genus  Pterostichus  Bonelli. 

A  very  fine  series  of  small  pterostichi,  belonging  to  the  subgenus 
Pseudoeryobius,  was  brought  out  by  the  collectors.  It  contains  rep- 
resentatives of  all  the  species  hitherto  known  from  the  Pribilof 
group  and  also  a  few  others  apparently  described  from  different 
parts  of  Alaska.  The  separation  and  identification  of  these  forms 
has  been  a  difficult  task  and  the  results  of  my  study  are  by  no  means 


152  NORTH   AMERICAN   FAUNA.  [No.  46 

satisfactory.  In  common  with  better  known  and  more  readily  rec- 
ognized insects  like  Cardbus  truncaticoUis  and  CKrysomela  subsvl- 
cata,  these  pterostichi  appear  to  vary  enormously  in  size,  color,  and 
depth  of  sculpture.  There  seems,  also,  to  be  some  variation  in 
thoracic  outline;  the  basal  foveae  are  affected  by  the  sculptural 
inconstancy  and  the  number  of  elytral  striae  of  one  individual.  In 
consequence  the  characters  ordinarily  used  for  specific  distinction  be- 
come much  lessened  in  value  and  one  has  to  depend,  in  great  part, 
upon  facies  for  separation.  Nevertheless  the  only  basis  for  assign- 
ing most  of  the  species  to  their  names,  most  of  which  were  given  by 
Dejean  and  the  early  Russian  writers,  is  found  in  these  same  char- 
acters and  the  attempt  has  been  made  to  use  them  with  discrimina- 
tion. A  study  of  the  species  of  Pseudocn/obius  of  both  hemispheres 
is  absolutely  needed  for  the  proper  delimitation  of  our  native  forms. 

Pterostichus  vindicatus  Mannerheim. 

St.  George,  June,  July  16;  St.  Paul,  June.  While  placed  in  this 
species  on  account  of  the  much  finer  elytral  striae,  the  identification 
must  be  considered  provisional.  This  is  the  first  record  for  the 
Pribilofs. 

Pterostichus  ventricosus  Eschscholtz. 

St.  George,  April  17,  May  6,  June  4  and  25,  July  16,  August  4, 
September.     St.  Paul,  June. 

Pterostichus  subexaratus  Mannerheim. 

St.  George,  April  1  and  17,  May  6  and  17,  August  4,  September  2 ; 
St.  Paul,  June.  Some  doubt  attaches  to  this  identification.  The 
species  has  not  hitherto  been  recorded  from  these  islands. 

Pterostichus  pinguedineus  Eschscholtz. 

St.  Paul  and  St.  George,  taken  every  month  from  April  17  to 
September  3.     Evidently  a  common  species. 

Pterostichus  hyperboreus  Mannerheim. 
St.  George,  May  6  and  June  10. 

Pterostichus  similis  Menetries. 

Common  on  both  islands.  Records  from  St.  George  are  June  4, 
10,  14,  19.  25,  July  4  and  16,  August  4 ;  from  St.  Paul,  April  15  and 
"May  to  July  ". 

Pterostichus  quadricollis  Menetries. 

St.  George  and  St.  Paul,  June.     Less  abundant  than  P.  similis. 

Pterostichus  subcaudatus  Mannerheim. 

St.  Paul  and  St.  George,  June.  This  is  the  first  record  for  the 
Pribilof  Islands. 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  153 

Pterostichus  empetricola  Dejean. 

St.  George,  June  4,  10,  25,  July  4  and  16,  August  4;  St.  Paul, 
May  17,  19,  22,  31,  June,  July  13,  September  1  and  3 ;  also  Sea  Lion 
Rock,  June  29.     Fairly  abundant  and  seems  variable  in  size. 

Genus  Amara  Bonelli. 
Amara  brunnipennis  Dejean. 

St.  Paul  and  St.  George.  Dates  are  shown  for  each  month  be- 
tween April  17  and  September  3.  Specimens  run  from  shining 
black  to  some  with  distinct  metallic  tinge  and  several  are  of  the 
form  with  brown  elytra.  There  is  also  a  good  deal  of  variation  in 
the  extent  of  punctuation  of  the  prothoracic  basal  impressions. 
This  insect  belongs  to  the  subgenus  Curtonotus  Stephens. 

Amara  glacialis  Mannerheim. 

Several  specimens  from  St.  Paul,  without  definite  date.  Belongs 
to  the  subgenus  Brady tus  Zimmermann. 

Amara  brunnea  Gyllenhal. 

Many  examples  from  St.  Paul,  May,  June,  and  July.  Belongs  in 
the  subgenus  Acrodon  Zimmermann. 

Family  DYTISCIDAE. 

Genus  Hydroporus  Clairville. 
Hydroporus  sp.  indet. 

A  few  specimens  are  at  hand  from  St.  George  (April  4  and  Sep- 
tember 3)  and  St.  Paul  (June  11).  At  present  it  is  not  feasible  to 
attempt  specific  determination  of  this  insect,  which  is  a  small  black 
species  3.5  mm.  long;  it  is  evidently  close  to  or  identical  with  H. 
nigellus  Mannerheim,  which  figures  in  our  lists  as  a  synonyn  of  H. 
tartaricus  Leconte. 

Hydroporus  sp.  indet. 

Three  examples  of  a  bicolored  species,  5  mm.  in  length,  come 
from  St.  George,  dated  June  10.  They  appear  to  be  related  to  H. 
truncatus  Mannerheim  in  size,  color,  and  oblique  truncation  of  the 
elytral  apices,  but  differ  from  the  description  in  some  details. 

Genus  Agabus  Leach. 
Agabus  sp.  indet. 

Three  females  from  St.  George,  June,  August,  and  September. 
The  male  is  necessary  for  identification. 

Family  SILPHIDAB. 
Genus  Lyrosoma  Mannerheim. 
Lyrosoma  opaca  Mannerheim. 

St.  George,  abundant ;  St.  Paul,  two  specimens ;  taken  every  month 
from  May  17  to  September  6 ;  Sealion  Rock,  June  29. 


154  NORTH   AMERICAN   FAUNA.  [No.  46 

Family    STAPHYLINIDAE. 

Genus  Atheta  Thomson. 

Atheta  (Megista)  nomadica  Casey. 

St.  Paul,  May  22  and  July  4,  many  specimens.  The  determination 
is  due  to  Dr.  A.  Fenyes  who  writes  that  it  is  probably  correct.  He  has 
also  given  us  the  generic  references  for  the  two  species  following. 

Atheta  sp.  indet. 

St.  Paul,  May  22.  / 

Genus  Ocyusa  Kraatz. 
Ocyusa  sp.  indet. 
St.  George,  May  4,  one  specimen  in  bad  condition. 

Genus  Liparocephalus  Maklin. 

Liparocephalus  brevipennis  Maklin. 

St.  Paul,  a  pair,  May  13 :  St.  George,  one  specimen,  September  2. 

Genus  Quedius  Stephens. 

Quedius  hyperboreus  Erichson. 

St.  Paul,  May  22,  one  specimen. 
Quedius  molochinus  Gravenhorst. 

One  specimen,  St.  George,  August  4. 

Genus  Tachinus  Gravenhorst. 

Tachinus  apterus  Maklin. 

About  sixty  specimens  of  Tachinus  belong  to  a  species  which  runs 
close  to  instabUis  Maklin  by  Dr.  Horn's  table.  Some  of  the  females, 
however,  have  the  median  dorsal  lobe  of  the  abdominal  apex  short, 
this  being  the  character  upon  which  T.  apterus  Maklin  is  based. 
Likely  enough,  the  whole  series  might  properly  be  referred  to  T. 
apterus,  which  was  not  known  to  Dr.  Horn.  Specimens  occur  on  both 
Islands,  and  the  dates  cover  April,  May,  June,  July,  and  September. 

Genus  Bryoporus  Kraatz. 

Bryoporus  near  insignis  Maklin. 

A  specimen  of  Bryoporus  collected  on  St.  Paul,  May  22,  may  be 
compared  with  Mycetoporus  insignia  Maklin  from  the  island  of 
Afognak.  The  description  seems  to  agree  in  most  of  the  principal 
points  but  differs  in  regard  to  coloration.  The  St.  Paul  specimen  is 
5.5  mm.  in  length,  very  shining,  piceous  black,  the  elytra  tending  to- 
ward castaneous.  The  antennae  are  blackish,  strongly  thickened  ex- 
ternally and  as  long  as  the  head  and  prothorax  together,  the  two  basal 
joints  yellowish,  third  a  little  darker.  Legs  pale  testaceous,  coxae 
darker,  hind  femora  and  trochanters  infuscate.  Hind  margins  of  ab- 
dominal segments  scarcely  perceptibly  lighter  above,  more  distinctly 


1923.]  INSECTS   OF   THE   PRIBILOF  ISLANDS.  155 

so  beneath.  The  four  punctures  described  by  Maklin  as  being  placed 
slightly  in  front  of  the  hind  margin  of  the  pronotum  are  very  dis- 
tinct, the  lateral  and  apical  series  are  small.  Elytra  sparsely  and 
finely  punctate  on  the  disk,  a  little  more  closely  toward  the  scutellum, 
in  addition  to  showing  the  usual  sutural,  discal,  and  lateral  series  of 
larger  punctures.  I  do  not  make  out  any  sexual  characters.  Myce- 
toporus  and  Bryoporus  are  separated  by  slender  characters  and  have 
been  united  by  Fauvel.  It  is  entirely  probable  that  the  species  in 
hand  is  congeneric  with  that  of  Maklin  though  perhaps  not  con- 
specific. 

Genus  Deliphrum  Erichson. 
Deliphrum  sp.  indet. 

About  a  dozen  specimens,  St.  Paul  and  St.  George,  covering  the 
months  of  April,  May,  June,  August,  and  September.  These  belong 
to  Deliphrum  or  some  closely  allied  genus  but  the  species  can  not  be 
identified  with  any  hitherto  recorded  from  North  America.  Quite 
possibly  it  may  be  described  from  Siberia. 

Genus  Olophrum  Erichson. 

Olophrum  marginatum  Kirby. 
St.  George,  May  6  and  September  2. 

Olophrum  fuscum  Gravenhorst. 
St.  George,  June  and  September ;  St.  Paul,  May. 

Genus  Micralymma  Westwood. 

Micralymma  dicksoni  Maklin. 

St.  George,  April  12,  June  4,  25,  August  16,  September  2;  St. 
Paul,  April  5.  The  identification  of  this  very  interesting  beach- 
inhabiting  insect  is  based  upon  comparison  with  specimens  in  the 
United  States  National  Museum,  having  the  above  specific  label  in 
the  handwriting  of  Mr.  Schwarz. 

Family  LATHRIDIIDAE 
Genus  Enicmus  Thomson. 
Enicmus  protensicollis  Mannerheim. 
St.  Paul,  one  specimen.      May  24. 

Family  BYRRHIDAE 
Genus  Byrrhus  Linnaeus. 
Byrrhus  fasciatus  Fabricius. 

St.  Paul,  several  specimens.  The  only  dates  given  are  May  23 
and  July  13.  These  specimens  agree  very  closely  with  examples  in 
the  United  States  National  Museum  from  Copper  Island,  carrying 
the  above  specific  label. 


156  NORTH   AMERICAN   FAUNA.  [No.  46 

Family  ELATERIDAE. 

Genus  Cryptohypnus  Eschscholtz. 

Cryptohypnus  littoralis  Eschscholtz. 

Several,  from  St.  Paul.  The  dates  given  are  April  30  to  May  22, 
but  most  of  the  specimens  are  simply  marked  "  Summer." 

Genus  Hypnvidus   Stephens. 

Hypnoidus  musculus  Eschscholtz/ 

St.  Paul,  four  specimens.  The  only  definite  date  is  June  20. 
Common  at  various  points  on  the  Alaskan  seacoast,  occurring  under 
shingle  along  the  beaches. 

Family  PTINIDAE. 

Genus  Trigonogenius  Solier. 

Trigonogenius  globulum  Solier. 

St.  Paul,  April;  St.  George,  April  17.  Widely  distributed  by 
commerce. 

Genus  Ptvnus  Linnaeus. 

Ptinus  fur  Linnaeus. 

St.  Paul,  January  6,  May  16 ;  St.  George,  May  17  and  September  3. 
Common  in  houses  over  most  of  the  civilized  world. 

Family  CHRYSOMELIDAE. 

Genus  Chrysomela  Linnaeus. 

Chrysomela  subsulcata  Mannerheim. 

The  collection  contains  a  beautiful  and  extensive  series,  varying 
in  size,  color,  and  depth  of  sculpture.  Some  specimens  are  green, 
others  blue,  while  several  are  decidedly  coppery.  A  few  are  black- 
ish with  very  little  luster.  They  come  from  both  St.  George  and 
St.  Paul  and  were  mostly  collected  in  May,  June,  and  July,  though 
the  dates  run  as  early  as  April  30  and  as  late  as  September  3.  Sev- 
eral larvae  of  different  sizes  bear  the  date  of  May  14-22. 

Family   AEGIALITIDAE. 

Genus  Aegialites  Mannerheim. 
Aegialites  calif ornicus  Motschulsky. 

Numerous  specimens  from  St.  Paul,  St.  George,  and  Sea  Lion 
Hock.  The  dates  run  between  June  4  and  July  8.  Larvae  were  taken 
June  4  and  June  29. 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  157 

Family  CURCTJLIONIDAB. 

Genus  Lophalophus  Leconte. 

Lophalophus  inquinatus  Mannerheim. 

St.  Paul,  fairly  common,  May  19  to  August  16. 

Genus  Trachodes  Germar. 

Trachodes  ptinoides  Germar. 
St.  George,  May  6  to  September  10. 

Genus  Orchestes  Illiger. 

Orchestes  parvicollis  Leconte. 
Three  specimens,  St.  Paul,  spring  of  1914. 


A  specimen  of  Ilybius  angustior  Gyllenhal  was  present  in  the  stomach  of  a  red  phala- 
rope,  collected  on  St.  George  Island  August  2,  1920,  and  one  of  Hadrotes,  not  specifically 
identified,  in  the  stomach  of  a  pectoral  sandpiper  collected  on  St.  Paul  Island  August  22, 
1914. 

In  addition  to  the  beetles  mentioned  in  the  preceding  list,  E.  A.  Schwarz  recorded  (Re- 
port on  Par  Seals  and  Fur  Seal  Islands,  Pt.  3,  pp.  548-549,  1899),  partly  on  the  basis  of 
Wosnesenski's  specimens  as  reported  by  Menetries,  Motschulsky,  and  Mannerheim,  the 
following  species  as  inhabitants  of  the  Pribilofs  : 

Laccophilus  decipiens  Le  Conte.      St.  George.   I  Cercyon  lateralis  Marsham.     St.  Paul. 
Berosus  maculosus  Mannerheim.     St.  George.  |    Hadrotes  crassus  Mannerheim.     St.   George. 

In  his  paper  on  the  Metamorphoses  of  some  Alaska  Coleoptera  (Proc.  Washington  Acad. 
Sci.,  vol.  2,  p.  201,  Nov.  24,  1900)  Trevor  Kincaid  records  also  the  weevil  Lepidophorus 
lineaticollis  Kirby  from  St.  Paul  and  describes  its  larva  and  pupa. 

E.  C.  Van  Dyke  in  the  1921  report  (Proc.  Calif.  Acad.  Sci.,  Fourth  Ser.,  11,  pp.  156- 
166,  Nov.,  1921)  adds  the  following  7  species:  Pterostichus  hudsonicios  Le  Conte,  Amara 
remotestriata  Dejean,  Agabus  hypomelas  Mannerheim,  Atheta  graminicola  Gravenhorst, 
Quedius  fulvicollis  Stephens,  Arpedium  beringenus  Van  Dyke,  and  Lepyrus  palustris  Sco- 
poli. 

An  easily  overlooked  paper  recording  26  species  of  Coleoptera  from  the  Aleutian  Islands, 
but  none  specifically  from  the  Pribilofs,  is  the  following :  Coinde,,  J.  P.,  Notice  sur  le  faune 
ornithologique  de  l'lle  de  Saint-Paul,  suivie  de  l'enumeration  de  quelques  especes  d'insectes 
(Coleopteres)  des  Aleoutiennes  et  du  Kamtschatka,  Rev.  et  Mag.  Zool.,  2'e  Sen,  T.  XII, 
pp.  396-405,  I860.— W.  L.  M. 

108731°— 23 11 


MECOPTEBA. 

By  Nathan  Banks,  Museum  of  Comparative  Zoology, 
Harvard  University. 

PI.  IX,  fig.  8. 

Family  PANORPIDAE. 
Boreus  borealis,  n.  sp. 

Brassy  black;  the  legs,  including  coxae  and  the  pleura,  the  lower 
half  of  the  beak,  the  wings  (except  tip  in  the  male),  and  the  ovi- 
positor (except  tip),  are  yellowish;  the  extreme  tips  of  tibia  and 
tarsal  joints  black;  male  genitalia  pale;  face  hairy,  vertex  shining; 
wings  in  male  very  long  and  slender,  fully  one-half  as  long  as  abdo- 
men, tip  curved  downward  and  ending  in  two  points,  one  more 
slender  than  the  other;  ventral  plate  of  male  truncate,  not  notched 
at  tip;  in  the  female  wings  reach  to  abdomen,  about  twice  as  long 
as  broad,  larger  than  in  allied  species ;  ovipositor  one-half  the  length 
of  the  abdomen,  basal  part  concave  above,  beyond  straight,  below 
hairy. 

Length :  Male,  3.8  mm. ;  female,  with  ovipositor,  5  mm. 

From  St.  Paul  Island,  Bering  Sea,  Alaska,  May  16-23, 1914  (Whit- 
ney coll.).  Differs  from  other  American  species  in  pale  coxae  and 
pleura,  longer  wings,  and  larger  size. 

See  Plate  IX,  Figure  8,  male  gentalia  from  side  and  behind,  and 
side  view  of  wings  and  ovipositor. 


Explanation  of  Plate  IX. 
Details  of  Scorpion  Fly  (Mecoptera)  and  Mites  (Arachnida,  pp.  237-239). 

Fig.  1.  Dermacarus  sp.,  venter  and  tarsus   I — hypopus. 
Fig.  2.  Notaspis  serrifrons,  edge  of  head  and  top  of  cephalothorax. 
Fig.  3.  Hilaira  glacialis,  palpus  of  male. 
Fig.  4.  Lohmannia  scabra,  dorsum. 

Fig.  5.  Parasitus   borealis,  epitomae  showing  variation. 
Fig.  6.  Tyroglyphus  whitneyi,  venter  and  tarsus  I — hypopus. 
Fig.  7.  Parasitus  borealis,  leg  II  of  male. 

Fig.  8.  Boreus  borealis  (Mecoptera)  ;  above,  male  genitalia  from  side  and  be- 
hind ;  below,  side  view  of  wings  and  ovipositor. 
158 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  IX. 


Details  of  Scorpion  Fly  (Mecoptera,  Fig.  8)  and  Mites  (Arachnida,  Figs.  1-7). 

(Explanation  on  page  158.) 


DIPTERA. 

Suborder  Orthorrhapha. 

Division  NEMATOCERA. 

Families  TIPULIDAE  and  RHTPHIDAE. 

By  Charles  P.  Alexander,  Department  of  Entomology, 
Massachusetts  Agricultural  College. 

(Plates  X  and  XI.) 

Our  knowledge  of  the  crane  flies  of  the  Pribilof  Islands  is  due 
largely  to  the  collections  made  by  G.  Dallas  Hanna  on  the  island  of 
St.  George  and  by  Alvin  G.  and  Elsie  G.  Whitney  on  the  island  of 
St.  Paul,  previous  collections  yielding  very  fragmentary  data.  In 
E.  A.  Schwarz's  list1  the  following  records  for  members  of  this 
family  of  flies  occur: 

Trichoeera  sp.  A  single  specimen  collected  by  Mr.  Barrett-Hamilton.  To  this 
or  an  allied  species  I  am  inclined  to  refer  the  "  gnat "  mentioned  by  Mr.  Elliott 
which  "  flits  about  in  large  swarms,  but  it  is  inoffensive  and  seeks  shelter  in 
the  grass." 

Tipulid.  A  single  larva  from  Mr.  Barrett-Hamilton's  collection  from  St.  Paul 
indicates  a  larger  species  than  the  Trichoeera  just  mentioned. 

The  above  material  was  determined  by  the  late  D.  W.  Coquillett ; 
additional  specimens  that  were  collected  by  Mr.  Elliott  in  1895  were 
found  among  the  undetermined  material  in  the  United  States  Na- 
tional Museum  and  will  be  found  recorded  under  Tipula  whitneyi 
on  a  later  page. 

The  crane  fly  fauna  of  these  islands  is  similar  to  that  of  many  other 
wind-swept  islands,  in  the  large  number  of  species  with  the  wings  so 
atrophied  that  the  flies  are  incapable  of  flight.  The  relatively  large 
number  of  species  that  seem  to  be  confined  to  the  Pribilofs  is  partly 
accounted  for  in  this  manner.  The  Alaskan  Tipulidae  named  by  Mr. 
Coquillett  and  now  contained  in  the  National  Museum  have  been 
compared  with  the  Pribilof  material  and  were  found  to  represent 
quite  a  different  fauna,  some  comparisons  to  which  are  noted  in  later 
pages. 

The  notes  of  the  collectors  of  the  material  are  quoted  under  their 
original  numbers. 

1  Sehwarz,  B.  A.,  List  of  insects  hitherto  known  from  the  Pribilof  Islands :  Report  on 
Fur  Seals  and  Fur-seal  Islands,  Part  3,  pp.  550-552,  1899. 

159 


160  NORTH   AMERICAN   FAUNA.  [No.  46 

Family  RHYPHIDAE. 

Genus  Trichocera  Meigen  (1803). 
Trichocera  sp. 

Numerous  specimens  of  these  crane-flies  were  included  in  the  col- 
lection, but  the  systematic  condition  of  the  species  of  the  genus  is 
such  that  it  is  impossible  to  identify  the  insects  at  this  time.  St. 
George  Island,  June  14,  1914;  St.  Paul  Island,  Sept.  30,  1911.  Mr. 
Whitney's  note  on  his  number  1£8  follows : 

"  Spring  1914.  12  gnats  ( ?)  flying  in  a  swarm  by  the  laboratory, 
St.  Paul  Island." 

Family  TIPULIDAB. 

Subfamily  Limnobiinae. 

Tribe  Pediciini. 

Genus  Tricyphona  Zetterstedt   (1840). 

Tricyphona  hannai,  n.  sp. 

Male,  length  about  7.7  mm. ;  wing,  5.8  mm. 

Rostrum  and  palpi  very  short,  dark  brown.  Antennae  short,  dark 
brown,  first  segment  about  half  again  as  long  as  the  second ;  flagellum 
short,  of  an  indeterminable  number  of  segments,  the  basal  segments 
greatly  enlarged,  thence  tapering  to  the  apex ;  the  terminal  segments 
very  small,  more  or  less  fused,  and  provided  with  long  verticils ;  eyes 
small,  hairy;  head  with  the  front  and  vertex  broad,  dark  brown, 
with  a  grayish  yellow  pollen. 

Pronotum  prominent;  thorax  dark  brown  with  a  grayish-yellow 
pollen;  halteres  long,  more  or  less  flattened  and  twisted,  the  knob 
not  prominent;  wings  considerably  atrophied  both  in  length  and 
width,  the  venation  considerably  degenerated  (see  pi.  10,  fig.  1)  ; 
the  color  is  light  brown,  the  disk  darker  anteriorly;  the  costa  is 
incrassated  and  provided  with  several  ranks  of  stout  hairlike  bristles ; 
i?'s  elongated,  straight,  in  a  line  with  i?4+5,  which  is  forked  at  the 
apex,  R4  and  R5  being  separate  at  the  wing-margin;  R2+3  indistinct 
on  its  terminal  portion;  cross- vein  r-m  elongate,  prominent;  media 
with  only  the  upper  branch  clearly  defined,  this  branch  apparently 
unforked;  cubitus  well  developed,  dark  brown,  well  defined;  two 
anal  veins,  the  second  very  long  and  straight;  some  of  the  veins 
with  strong  hairs  or  hairlike  bristles  on  them,  a  group  of  about  five 
in  the  stigmal  region,  about  six  along  R2+3;  a  considerable  series 
on  the  apical  portion  of  R4+5  and  on  Rt  and  R5;  about  twelve  on 
the  upper  branch  of  M,  others  on  the  forks  of  Cu,  and  about  nine, 
evenly  spaced,  on  the  second  anal  vein. 

Abdomen  dark  brown,  with  sparse  long,  yellow  appressed  hairs; 
caudal  and  lateral  margins  of  the  segments  paler;  hypopygium  (see 
PL  X,  fig.  6)  with  the  ninth  tergite  rather  broad,  the  caudal  margin 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  161 

gently  concave;  pleurites  very  short  and  stout,  the  outer  face  with 
numerous  pale  hairs,  the  inner  face  with  numerous  black  spicules; 
appendages  two,  the  dorsal  appendage  a  capitate  lobe  on  a  very 
short  pedicel,  the  head  with  numerous  black  spicules  and  a  few  long 
yellow  hairs;  ventral  appendage  a  flattened  blade-like  subchitinized 
arm ;  ninth  sternite  narrow,  the  caudal  margin  with  the  median  por- 
tion straight  but  slightly  denticulate  at  the  ends. 

Holotype,  $ ,  St.  George  Island,  Bering  Sea ;  June  10,  1914 
(Hanna).  "Lot-number  13.  Found  near  a  pool  in  Sphagnum  bog, 
west  of  village." 

There  can  be  little  doubt  but  that  this  insect  is  a  degenerate  species 
of  Tricyphona  with  the  fused  portion  of  veins  R4+5  of  the  wings 
very  extensive,  and  many  details  of  the  venation  considerably  atro- 
phied or  hypertrophied.  The  insect  is  named  in  honor  of  the  col- 
lector, G.  Dallas  Hanna. 

Subfamily  Tipulinae. 
Tribe  Tipulini. 

Genus  Tipula  Linnaeus   (1758). 
Tipula  whitneyi,  n.  sp. 

Male,  length  13-14  mm. ;  wing,  2.8-5.5  mm. 

Female,  length  19-22  mm. ;  wing,  2.5-3  mm. 

Frontal  prolongation  of  the  head  moderate  in  length,  gray,  nasus 
indistinct;  palpi  dark  brown.  Antennae  dark  brownish-black,  the 
segments  of  the  flagellum  very  slightly  constricted  beyond  the  en- 
larged base ;  the  segments  covered  with  a  short,  dense,  gray  pubescence. 
Head  clear  whitish  gray  with  a  distinct  impressed  median  line. 

Pronotal  scutum  light  gray  with  a  narrow  median  brown  vitta. 
Mesonotal  praescutum  and  scutum  clear  light  gray  without  apparent 
darker  markings  of  any  kind;  scutellum  and  postnotum  brownish 
gray  with  a  narrow  brown  median  vitta ;  pleura  brownish  gray ; 
halteres  short,  yellowish  brown,  the  knob  darker  brown;  legs  with 
the  coxae  prominent,  light  gray,  with  numerous  long  pale  hairs ;  tro- 
chanters reddish  brown;  femora  and  tibiae  reddish  brown,  tipped 
with  dark  brown ;  tarsi  dark  brown  to  black ;  wings  extremely  reduced 
in  both  sexes,  in  some  specimens  a  little  longer  than  in  others,  one 
male  having  the  right  wing  twice  as  long  as  the  left  wing;  in  most 
specimens  the  wings  extend  about  to  the  tip  of  the  first  abdominal 
segment;  wings  light  brown,  the  costal  margin  very  greatly  in- 
crassated,  the  region  immediately  behind  the  costa  with  an  abundance 
of  short  bristles;  venation  (see  PI.  X,  fig.  2)  indistinct,  distorted,  but 
traceable. 

Abdomen  varying  from  brown  to  reddish  brown,  with  a  broad, 
dark  brown  dorso-median  stripe;  first  tergite  largely  dark  brown; 


162  NORTH  AMERICAN   FAUNA.  [No.  46 

lateral  margins  of  the  sclerites  broadly,  caudal  margins  very  nar- 
rowly, pale;  sternites  grayish  brown;  hypopygium  (see  PI.  X,fig.  7) 
with  the  ninth  tergite  (see  PI.  XI,  fig.  13)  prominent,  the  caudal 
margin  with  a  broad  U-shaped  median  notch  which  is  notched  again 
by  a  smaller  W-shaped  incision;  the  lateral  lobes  are  broadly  trun- 
cated, with  the  caudal  margin  shiny,  tumid;  ninth  pleurite  large, 
complete,  situated  on  the  dorso-caudal  face  of  the  ninth  sternite,  the 
ventral  inner  angle  clothed  with  numerous  long  pale  yellow  hairs; 
pleural  appendages  two,  the  outer  appendage  a  slender,  cylindrical 
fleshy  lobe  that  is  clothed  with  comparatively  short  hairs;  inner 
pleural  appendage  large,  prominent,  compressed,  projecting  cephalad 
as  a  narrowed  lobe  which  occupies  the  notch  of  the  tergite ;  near  the 
apex  it  is  split  into  a  smaller  lobe  which  is  deflected  laterad ;  ninth 
sternite  with  a  broad  U-shaped  notch  on  the  caudal  half;  on  the 
cephalic  half  the  margins  of  each  side  are  approximated  but  not 
contiguous,  the  median  area  membranaceous ;  eighth  sternite  with  the 
caudal  margin  simple,  unarmed. 

The  female  is  similar  to  the  male  but  the  dorso-median  abdominal 
vitta  is  often  interrupted  on  the  basal  third  of  each  segment;  the 
wings  are  still  shorter,  extending  to  just  beyond  the  base  of  the  first 
abdominal  segment;  valves  of  the  ovipositor  (see  pi.  11,  fig.  21) 
very  long  and  slender,  the  tergal  valves  slightly  divergent,  enlarged 
basally,  thence  gradually  narrowed  to  the  tip ;  sternal  valves  shorter, 
compressed,  the  apices  rather  blunt. 

Holotype,  S,  St.  George  Island,  Bering  Sea;  June  12,  1914 
(Hanna)  ;  lot  16.  Allotype,  9,  topotypic;  lot  27,  June  16,  1914. 
Paratypes,  35  $  's,  9  %  as  follows :  18  $  's,  8  9  's,  topotypic,  June  12 
to  July  8,  1914  (Hanna) ;  lots  16,  17,  27,  30,  41,  46,  49,  52,  and  55. 
1  $  ,  Otter  Island,  July  3,  1913  (Whitney)  ;  lot  60.  1  <J ,  1  9  ,  St. 
Paul  Island,  June  10,  1913  (Whitney) ;  lot  40.  2  9  's  with  the  last, 
June  1,  1914;  lot  170.  1  $  ,  bred  from  pupa,  with  the  last,  June, 
1914;  lot  186.  1  $,  1  9  (gravid),  St.  Paul  Island,  July  10,  1895 
(H.  W.  Elliott).  1  $  ,  with  the  last,  July  12,  1895;  U.  S.  Nat.  Mus. 
Ace.  No.  30147. 

The  accompanying  collectors'  notes  with  the  above  lot  numbers 
are  as  follows: 

Hanna:  Lot  16.  Found  crawling  over  grass  of  high  beach  lands,  not  seen 
near  bogs  or  on  top  of  high  hills;  lot  30,  toward  East  Rookery  from  village — 
none  seen  with  wings  developed ;  lot  41,  uplands  toward  Staraya  Artel  Rookery ; 
lot  46,  from  toward  East  Rookery ;  lot  49,  toward  Zapadni — damaged  by 
cyanide;  lot  52,  from  toward  Zapadni  Rookery. 

Whitney :  Lot  40.  In  grass,  one  at  Kitovi  and  the  other  on  Reef  Peninsula ; 
lot  60,  Otter  Island  (6  miles  from  St.  Paul). 

This  fly  is  named  in  honor  of  the  collector  of  certain  of  the  para- 
types, Mr.  Alvin  G.  Whitney. 


1923.]  INSECTS   OF   THE  PRIBILOF  ISLANDS.  163 

The  pupal  skin  from  which  one  of  the  paratypes  was  bred  was 
collected  about  June  1  and  the  adult  fly  emerged  early  in  June.  The 
following  notes  on  the  exuvium  are  included : 

Length  about  21.5  mm ;  diameter  about  5  mm. ;  prothoracic  breath- 
ing horns  very  short,  finely  crenulated ;  abdominal  tergites  with  the 
caudal  half  of  each  segment  bearing  four  blunt  tubercles  in  align- 
ment ;  the  eighth  segment  with  a  fleshy  tubercle  on  each  side ;  ninth 
tergite  (see  PI.  XI,  fig.  23)  with  the  caudal  margin  deeply  concave; 
the  lateral  angles  wrinkled;  tergal  valves  very  elongated,  blunt  at 
their  apices;  sternal  valves  shorter;  caudal  half  of  sternite  five  (see 
PI.  XI,  fig.  24)  with  four  subacute  fleshy  tubercles  on  each  side  of 
the  median  line;  sixth  sternite  with  three  similar  tubercles;  seventh 
sternite  with  two  similar  tubercles;  eighth  sternite  with  six  large 
tubercles;  leg  pads  ending  about  at  the  base  of  abdominal  segment 
four;  wing  pads  ending  just  beyond  the  base  of  segment  three. 

Tipula  pribilofensis,  n.  sp.  Pis.  X  and  XI. 

Male. — Length  12.5-13.5  mm.;  wing,  10.5-11.5  mm.;  antennae 
about  5.5  mm. 

Female. — Length,  15.5-19  mm.;  wing,  10-11  mm. 

Frontal  prolongation  of  the  head  rather  short,  dark  brown,  with 
a  dark  gray  bloom  ;  nasus  distinct ;  palpi  short,  dark  brown ;  antennae 
rather  elongated,  black,  the  flagellar  segments  beyond  the  first  deeply 
constricted  at  their  middle ;  head  dark  with  a  dense,  dark  gray  bloom. 

Pronotal  scutum  gray,  the  scutellum  yellowish  on  the  lateral  mar- 
gins, this  color  becoming  confluent  with  the  same  color  of  the  dorso- 
pleural  membranes;  mesonotum  gray,  stripes  not  indicated;  sides 
of  the  scutellum  and  postnotum  more  yellowish ;  pleura  brownish 
gray;  halteres  rather  short,  dull  yellow,  the  knobs  more  brownish; 
legs  with  the  coxae  gray,  trochanters,  femora  and  tibiae  brown,  the 
two  latter  a  little  darkened  at  their  apices;  tarsi  black;  wings  semi- 
atrophied,  the  length  little  reduced  but  the  width  considerably  re- 
stricted so  that  the  venation  is  much  distorted;  color  of  the  wings 
pale  brownish,  the  stigma  distinct,  pale  brown,  not  encroaching  into 
the  base  of  cell  R2;  veins  brown;  venation  as  in  Plate  10,  Figure  3. 

Abdominal  tergites  reddish  yellow  with  three  indistinct  inter- 
rupted brown  lines,  the  lateral  stripes  becoming  distinct  only  on 
the  apical  segments  where  they  suffuse  the  entire  bases  of  the 
sclerites;  ninth  tergite  black;  tergites  with  conspicuous  transverse 
punctured  areas  on  the  basal  half  of  each  segment,  these  areas  in- 
terrupted on  the  mid-dorsal  line;  hypopygium  (see  PI.  X,  fig.  8) 
with  the  ninth  tergite  (see  PI.  XI,  fig.  14)  extensive,  the  caudal  mar- 
gin with  a  very  broad  V-shaped  notch,  the  lateral  angles  prolonged 
caudad  as  shiny  impunctate  horns;  pleural  appendages  two;  the 
outer  appendage  a  conspicuous  elongated  fleshy  lobe,  narrowed  at 


164  NORTH   AMERICAN   FAUNA.  [No.  46 

the  base,  thence  very  slightly  expanded  and  tapering  gradually  to 
the  blunt  apex ;  it  is  clothed  with  abundant  hairs,  on  the  caudal  face 
very  long,  divergent,  on  the  cephalic  and  lateral  faces  short,  more 
appressed;  inner  pleural  appendage  a  complex,  flattened,  chitinized 
lobe  divided  into  two  lobules,  the  ventral  or  caudal  lobule  project- 
ing caudad  as  a  compressed  blade  that  is  blunt  at  the  apex,  the  outer 
face  with  about  eight  short  bristles,  the  inner  face  with  several  long 
pale  hairs;  the  inner  or  dorsal  lobule  jutting  into  the  notch  of  the 
ninth  tergite,  flattened,  compressed,  with  indistinct  parallel  grooves ; 
the  sterno-pleural  suture  is  indistinct ;  at  the  point  where  it  is  usually 
located  a  short,  slender,  fleshy  setigerous  lobe;  eighth  sternite  (see 
pi.  11,  fig.  19)  produced  caudad  as  a  very  flattened,  depressed,  median 
arm  that  is  shaped  like  a  spade ;  the  apex  is  gently  notched  medially 
by  a  broad  U-shaped  incision;  the  caudal  margin  of  this  tongue  is 
fringed  with  delicate  pale  hairs. 

The  female  is  similar  to  the  male;  the  antennal  segments  simple 
throughout;  abdominal  tergites  dark  gray,  the  caudal  margins  of 
the  segments  brighter,  more  yellowish;  OAdpositor  (see  PL  XI,  fig. 
20)  with  the  last  tergite  extremely  elongated,  smooth,  shiny  black, 
chitinized;  tergal  valves  of  the  ovipositor  triangular,  lying  both 
transversely  and  vertically,  short,  acutely  pointed  from  very  broad 
bases,  the  apices  divergent;  the  dorsal  face  smooth,  light  chestnut 
brown ;  the  outer  face  with  a  prominent  median  carina  running  from 
the  base  to  the  apex,  the  remaining  surface  of  this  face  with  a 
roughened  irregular  meshwork  of  raised  lines;  the  ridges  between 
the  three  faces  of  the  valves  with  numerous  fimbriate  hairs;  sternal 
valves  reduced  to  tiny  lobes. 

Holotype,  $  ,  St.  Paul  Island,  June  1,  1914  (Whitney) ;  lot  170. 
"No  170.  About  June  1,  1914.  Tolstoi  sand  dunes.  Crane  flies 
were  crawling  everywhere  at  this  time  and  many  were  mating. 
Allotype,   $  ,  topotypic.    Paratypes,  20   $  's,  3   ?  's,  topotypic. 

Tipula  aleutica,  n.  sp.  Pis.  X  and  XI. 

Male. — Length  about  13-14  mm;  wing,  13.5  mm.  Discolored  by 
cyanide. 

Frontal  prolongation  of  the  head  dark  brown,  short  and  stout: 
nasus  indistinct;  antennae  dark  brown,  rather  short,  the  segments 
not  constricted;  head  dark  grayish  brown  with  abundant  long  pale 
hairs. 

Pronotal  scutum  grayish  brown  with  abundant  long  pale  hairs; 
mesonotal  praescutum  gray  with  blue-gray  stripes,  these  latter  in- 
distinctly margined  with  darker;  the  median  stripe  broadest  at  the 
cephalic  end,  narrowed  at  the  suture,  these  stripes  appearing  to  be 
discolored,  probably  by  the  action  of  cyanide;  scutum  gray,  the 
lobes  blue-gray;    pleura  dull  gray;    halteres  short,    pale   yellowish 


1923.]  INSECTS   OF    THE   PRIBILOF   ISLANDS.  165 

throughout;  legs  with  the  coxae  dull  gray  densely  covered  with  long 
pale  hairs;  trochanters  brown;  femora  and  tibia  light  brownish 
yellow,  the  apices  slightly  darkened;  tarsi  dark  brown;  wings  with 
a  very  faint  brownish  tinge,  the  stigma  brown ;  veins  dark  brown ; 
venation  as  in  Plate  10,  Figure  4. 

Abdomen  brownish  gray,  the  caudal  margins  of  the  segments 
ringed  with  paler;  hypopygium  (see  PL  X,  fig.  9)  with  the  ninth 
tergite  (see  PI.  XI,  fig.  15)  moderately  prominent,  the  caudal  margin 
straight  across,  with  two  lobes,  one  on  either  side  of  the  median 
line;  these  lobes  pale  yellow,  conical,  their  apices  rather  acute,  the 
notch  between  them  narrowly  V-shaped;  ninth  sterno-pleurite 
prominent,  the  pleural  region  partially  separated  from  the  sternite 
by  a  conspicuous  arcuated  suture  beneath;  pleural  appendages  two, 
situated  far  out  near  the  apex  of  the  sterno-pleurite,  the  outer  ap 
pendage  pale,  prominent,  flattened,  a  little  narrowed  toward  the 
blunt  apex;  inner  appendage  of  a  very  simple  structure,  a  pale 
slightly  chitinized  lobe  whose  anterior  angle  is  produced  cephalad 
as  a  long  subacute  lobule,  on  the  outer  face  near  the  caudal  margin, 
a  slender,  acutely  pointed  horn  directed  cephalad;  ninth  sternite 
profoundly  incised  by  a  very  narrow  V-shaped  notch,  the  adjacent 
margins  pale-pubescent,  not  approximated;  eighth  sternite  narrow, 
the  caudal  margin  straight  across,  unarmed. 

Holotype,  8,  St.  George  Island,  June  27,  1914  (Hanna).  "Lot 
49.     Toward  Zapadni." 

This  crane  fly  belongs  to  the  group  of  perlongipes  Johnson,  sul- 
phured Doane,  tenebrosa  Coquillett,  and  kennicotti  Alexander.  The 
only  species  with  which  it  requires  comparison  are  cimmeria  Speiser, 
and  tenebrosa  Coquillett,  and  this  comparison  is  given  herewith,  the 
notes  and  figures  being  based  upon  the  types  in  the  United  States 
National  Museum. 

Tipula  cimmeria  Speiser  (Dem  Kilimandjaro,  dem  Meru  Ex- 
pedition, 10,  Diptera.  4,  Orthorhapha.  Nematocera,  p.  57,  1909)  is 
the  correct  name  for  Tipula  strigata  Coquillett.  Type  number  5205, 
U.  S.  National  Museum,  from  Yakutat,  Alaska,  June  21,  1899,  col- 
lected by  Kincaid. 

The  type  of  strigata  is  a  male;  antennae  rather  short,  scape  dull 
yellow,  flagellum,  dark  brown,  the  segments  a  little  constricted 
beyond  the  base :  Frontal  prolongation  of  the  head  short,  nasus  very 
prominent;  Wing- venation  with  the  basal  deflection  of  R4.,5, 
r-m  and  the  basal  deflection  of  M1+2  almost  in  a  line.  Hypopygium 
with  the  tergite,  pleurite  and  sternite  fused  in  an  almost  continuous 
ring,  the  pleural  suture  well-indicated  beneath;  the  tergo-pleural 
notch  small,  on  the  caudal  margin  only;  ninth  tergite  (see  PI.  XI, 
fig.  16)  subquadrate,  dark  brown,  with  the  caudal  margin  trans- 
versely truncated  and  bearing  a  pair  of  median  lobes   (as  in  the 


166  NORTH   AMERICAN   FAUNA.  [No.  46 

tephrocephala  group)  ;  these  lobes  pale,  darkened  at  their  apices, 
very  closely  approximated  on  the  basal  three-quarters,  the  tips  more 
separated,  the  apices  of  the  lobes  minutely  spiculose;  the  length  of 
these  lobes  is  about  the  same  as  the  length  of  the  tergite;  they  are 
fringed  on  their  outer  lateral  margin  with  long  hairs;  ninth  pleur- 
ite  with  the  suture  conspicuous  beneath,  broadly  U-shaped;  the  ven- 
trocaudal  angle  with  a  tuft  of  long  hairs  which  are  decussate  on  the 
median  line  beneath;  outer  pleural  appendage  (see  PL  X,  fig.  12) 
large,  prominent,  pale,  fleshy,  very  flattened,  elongate,  slightly  con- 
stricted beyond  the  base,  then  expanded,  the  apex  a  little  pointed; 
the  outer  face  with  scanty  strigose  yellow  hairs;  inner  pleural  ap- 
pendage very  large,  powerful,  bilobed,  the  outer  or  caudal  lobe  short, 
subrotund,  the  apex  a  little  truncated,  densely  and  finely  pale  stri- 
gose on  the  inner  face;  inner  lobe  flattened,  compressed;  ninth  ster- 
nite  deeply  divided,  at  the  caudal  angle  just  behind  the  suture  with 
a  sparse  tuft  of  long  pale  hairs,  decussate  on  the  median  line  beneath ; 
near  the  base  of  the  split  a  dense  tuft  of  golden  yellow  hair;  eighth 
sternite  prominent,  straight  across  the  caudal  margin,  unarmed 
with  any  brush  or  tuft. 

Tipula  tenebrosa  Coquillett  was  described  from  Berg  Bay,  Alaska; 
collected  June  10,  1899,  by  Kincaid;  type  number  5206,  U.  S.  Na- 
tional Museum.  The  type  is  a  male ;  the  hypopygium  has  the  ninth 
tergite  (see  PI.  XI,  fig.  17)  large,  convex,  the  caudal  margin  with  a 
prominent  stout  lobe  on  either  side  of  the  median  line,  these  sepa- 
rated by  a  space  equal  to  about  one-half  the  diameter  of  the  lobe; 
the  apices  of  these  lobes  blackened,  minutely  spiculose;  caudal  mar- 
gin of  the  tergite  sloping  obliquely  backward  from  these  lobes ;  notch 
between  the  ninth  tergite  and  the  ninth  pleurite  quite  deep,  but  not 
running  back  to  the  eighth  segment;  ninth  pleurite  incomplete,  the 
pleural  suture  well  indicated  beneath;  the  pleural  region  produced 
caudad  as  a  blunt  triangular  arm  bearing  the  appendages  out  near 
its  apex;  outer  pleural  appendage  (see  PI.  X,  fig.  11)  flattened,  sub- 
quadrate  or  slightly  elongated,  bearing  at  the  base  on  the  inside  the 
inner  pleural  appendage  which  is  flattened,  bilobed,  the  caudal  lobe 
a  short,  blackened,  chitinized  point;  the  caudal  face  of  the  lobe  on 
the  basal  half  is  downy  pubescent;  ninth  sternite  deeply  cleft  on  the 
median  line  beneath  but  the  adjoining  sides  contiguous;  eighth  ster- 
nite prominent,  the  caudal  margin  unarmed.  Coquillett's  descrip- 
tion of  the  hypopygium  does  not  agree  at  all  with  the  type;  the 
outer  pleural  appendages  are  described  as  being  nearly  twice  as  long 
as  wide,  the  lower  outer  angle  considerably  prolonged  beyond  the 
upper  one ;  this  agrees  much  better  with  the  somewhat  similar  Tipula 
ci?)imeria,  discussed  above. 


1923.]  INSECTS   OF   THE   PRIBILOF  ISLANDS.  167 

Tipula  alascaensis,  n.  sp.  Pis.  X  and  XI. 

Male. — Length,  11-13.5  mm. ;  wing,  14.5-15  mm. 

Female. — Length,  15-18  mm. ;  wing,  17.5  mm. 

Frontal  prolongation  of  the  head  bluish  gray,  very  short,  nasus 
indistinct;  palpi  gray,  short;  antennae  very  short,  black,  with  a 
sparse  grayish  bloom;  first  segment  elongated,  longer  than  the  sec- 
and  and  third  together;  the  flagellar  segments  very  short,  slightly 
constricted  beyond  the  basal  swelling ;  head  blue-gray  with  abundant 
long  hairs,  especially  a  tuft  on  the  genae. 

Mesonotum  dark  gray  with  rather  indistinct  stripes,  the  median 
vitta  very  broad,  rapidly  narrowed  behind;  lateral  stripes  narrow, 
beginning  behind  the  conspicuous  pseudosutural  foveae;  thoracic 
interspaces  with  short  pale,  erect  hair;  scutum  and  lateral  por- 
tions of  the  postnotum  with  abundant  erect  black  hairs;  pleura 
dark  gray,  smooth,  a  large  setigerous  area  on  the  mesepisternum 
behind  the  fore  coxae;  halteres  short,  brown,  the  knobs  a  little 
brighter;  legs  with  the  coxae  gray,  clothed  with  abundant  long 
yellow  hairs;  femora  yellowish  brown  tipped  with  dark  brown; 
tibiae  brown  tipped  with  darker  brown;  tarsi  dark  brown;  wings 
fully  developed  in  both  sexes,  strongly  tinged  with  brownish  yellow, 
the  costal  cell  not  different  in  color  from  the  other  cells  of  the  wing ; 
stigma  conspicuous,  oval,  dark  brown ;  small  areas  before  the  stigma 
in  cell  1st  B1  and  beyond  the  stigma  in  cell  2d  B^  and  the  base  of 
R2  slightly  paler;  veins  dark  brown;  venation  as  in  Plate  X, 
Figure  5. 

Abdomen  dark  gray,  the  segments  narrowly  ringed  with  pale 
yellowish  around  the  caudal  margin;  hypopyginm  (see  PI.  X,  fig. 
10)  very  inconspicuous  and  somewhat  concealed;  ninth  tergite  (see 
PI.  XI,  fig.  18)  rather  prominent,  the  caudal  margin  rounded,  with 
a  deep,  narrow  median  notch;  the  lateral  lobes  are  thus  very  broad 
and  somewhat  obliquely  truncated;  dorsal  surface  of  the  sclerite 
densely  hairy;  ninth  pleurite  small,  complete,  situated  on  the  dorso- 
caudal  face  of  the  ninth  sternite;  outer  pleural  appendage  short, 
clavate,  slightly  enlarged  at  the  base,  the  head  rounded,  clothed 
with  abundant  golden  hairs;  inner  pleural  appendage  compressed, 
flattened,  on  the  outer  face  clothed  with  short,  appressed  golden 
hairs;  ninth  sternite  prominent,  with  a  very  deep  median  notch 
whose  margins  are  widely  separated. 

The  female  is  generally  similar  to  the  male;  the  ovipositor  has 
the  last  two  segments  exceedingly  narrowed  as  in  besselsi  Osten 
Sacken  and  pilicops  Alexander;  the  tergal  valves  (see  PI.  XI,  fig.  22) 
acute  but  small,  tapering  gradually  from  the  broad  base,  the  apices 
divergent. 


168  NORTH  AMERICAN   FAUNA.  [No.  46 

Holotype,  male,  St.  George  Island,  June  14,  1914  (Hanna),  Lot 
number  17.  Allotype,  female,  topotypic.  Paratypes,  2  males,  3 
females,  topotypic;  2  females,  topotypic  on  June  16,  1914  (Lot  27). 
"Lot  number  17.  In  wet  places,  mostly,  but  some  seen  crawling 
over  grass  far  from  water.  None  seen  flying.  Those  with  wings  best 
developed  (the  present  species)  from  Spring  Creek,  Garden  Cove. 
One  seen  with  the  very  fuzzy  fly  in  No.  18  (Scatophaga)  beneath  it; 
apparently  both  were  fighting."  "Lot  number  27.  Garden  Cove. 
Mrs.  E.  G.  Whitney."  / 

Tipula,  sp. 

Abundant  larvae  of  an  unknown  species  of  TipvZa  taken  on  St. 
Paul  island  July  18-20,  1913.  Mr.  Whitney's  notes  on  the  species 
are  very  interesting: 

No.  77.  July  18,  1913.  Reef  Parade  Ground.  12  larvae.  Abundant  every- 
where there  around  the  roots  of  grasses,  herbs,  and  especially  under  beds  of 
moss  on  the  roots  of  which  it  feeds,  killing  the  moss  over  considerable  areas. 
Under  such  a  moss  bed  I  found  as  many  as  20  to  the  square  foot.  This  larva 
is  found  all  over  the  island  in  grassy  or  mossy  places  and  all  through  the 
summer  season.  It  must  be  of  considerable  ecological  importance  because  of 
its  food  value  to  the  birds  and  foxes.  The  foxes  will  dig  over  large  areas  of 
moss  beds  to  feed  on  these  larvae.  Was  unable  to  find  the  species  in  adult 
form.  Could  not  seem  to  raise  adults  in  laboratory  by  keeping  larvae  with 
one  of  the  food  plants.  It  may  possibly  be  the  larval  form  of  the  crane  fly, 
which  is  very  abundant.     Color  not  altered  by  pickling  in  alcohol. 

The  identity  of  the  form  with  any  of  the  adult  flies  known  from 
the  Pribilofs  is  very  doubtful.  The  large  size  of  the  larvae  in  mid- 
July  would  imply  a  species  that  emerges  at  or  near  the  very  end  of 
the  growing  season,  and  it  seems  possible  that  they  belong  to  such 
a  species,  as  yet  unknown. 

The  fully  grown  larva  measures  29-30  mm.  in  length  and  about  5 
mm.  in  diameter;  the  form  is  plump,  color  light  brownish  yellow 
without  conspicuous  darker  markings;  hairs  and  setae  sparse;  the 
dorsa  of  the  thoracic  segments  with  hairs  as  in  Plate  11,  Figure  25 ; 
the  abdominal  segments  have  six  bristles  in  alignment,  the  inter- 
mediate four  being  almost  evenly  spaced,  the  outer  one  being  much 
the  strongest;  the  fifth  and  sixth  again  are  weak  and  situated  close 
to  the  strong  bristles  (see  PI.  XI,  fig.  26).  The  stigmal  field  is  sur- 
rounded by  six  weak  teeth,  the  dorsal  pair  closely  approximated,  the 
lateral  pair  being  latero-dorsal  in  position;  the  ventral  pair  very 
broad,  the  inner  face  with  a  broad-triangular  black  chitinized  area. 
Stigmata  large,  separated  by  a  distance  about  equal  to  the  diameter 
of  one  stigma,  located  ventrad  of  the  four  dorsal-lying  teeth  that 
surround  the  stigmal  field ;  gills  fleshy,  not  prominent.  (See  PI.  XI, 
figs.  27,  28.) 

Trioyphona  glacialis  Alexander,  and  Tipula  pribiloiHa  Alexander  have  previously  been 
recorded  by  this  author.  (Proc.  Calif.  Acad.  ScL,  Fourth  Ser.  11,  pp.  183-184,  Nov., 
1921.  )-*-W.  L.  M. 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey. 


PLATE    X. 


Details  of  Crane  Flies  (Tipulidae 

(Explanation  on  page  169.) 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey. 


PLATE    XI. 


Details  of  Crane  Flies  (Tipulidae). 

(Explanation  on  page  169.) 


[No.  40  INSECTS   OF   THE   PRIBILOF   ISLANDS.  169 

Explanation  of  Plates  X  and  XI. 

Plate  X. — Details  of  Crane  Flies  (Tipulidae). 

Fig.    1.  Wing  of  Tricyphona  hannai,  s*p.  n. 

Fig.   2.  Wing  of  Tipula  whitneyi,  sp.  n. 

Fig.    3.  Wing  of  Tipula  pribilofensis,  sp.  n. 

Fig.    4.  Wing  of  Tipula  aleutica,  sp.  n. ;  Ri,  2,  s,  t,  »=Radial  veins;  Mi,  M»,  M«= 

Medial  veins;  Ciii,  Cu2=Cubital  veins;  2d  A=second  anal  vein. 
Fig.    5.  Wing  of  Tipula  alascaensis,  sp.  n. 
Fig.    6.  Hypopygium  of  Tricyphona  hannai;  dorsal  aspect;  pZ=ninth  pleurite; 

<Z=dorsal  appendage;  v=ventral  appendage. 
Fig.    7.  Hypopygium   of   Tipula  whitneyi;  lateral  aspect;   8t,   9t=eighth  and 

ninth  tergites ;  8s=eighth  sternite. 
Fig.    8.  Hypopygium  of  Tipula  pribilofensis ;  lateral  aspect;  lettering  as  in  fig. 

7;  9  st-pl=ninth  sterno-pleurite. 
Fig.    9.  Hypopygium  of  Tipula  aleutica;  lateral  aspect;  lettering  as  in  figs.  7 

and  8. 
Fig.  10.  Hypopygium  of  Tipula  alascaensis;  lateral  aspect;  lettering  as  in  fig. 

7;  9s,  9pl=ninth  sternite  and  pleurite. 
Fig.  11.  Outer  pleural  appendage  of  Tipula,  tenebrosa  Coquillett,  lateral  aspect. 
Fig.  12.  Outer  pleural  appendage  of  Tipula  cimmeria  Speiser ;  lateral  aspect. 

Plate  XI. — Details  of  Crane  Flies  (Tipulidae). 

Fig.  13.  Ninth  tergite  of  Tipula  whitneyi;  dorsal  aspect. 

Fig.  14.  Ninth  tergite  of  Tipula  pribilofensis ;  dorsal  aspect. 

Fig.  15.  Ninth  tergite  of  Tipula  aleutica;  dorsal  aspect. 

Fig.  16.  Ninth  tergite  of  Tipula  cimmeria;  dorsal  aspect. 

Fig.  17.  Ninth  tergite  of  Tipula  tenebrosa;  dorsal  aspect. 

Fig.  18.  Ninth  tergite  of  Tipula  alascaensis ;  dorsal  aspect. 

Fig.  19.  Eighth  sternite  of  Tipula  pribilofensis ;  ventral  aspect  of  the  median  lobe. 

Fig.  20.  Ovipositor  of  Tipula  pribilofensis;  dorsal  aspect. 

Fig.  21.  Ovipositor  of  Tipula  whitneyi;  dorsal  aspect. 

Fig.  22.  Ovipositor  of  Tipula  alascaensis ;  dorsal  aspect. 

Fig.  23.  Pupa  of  Tipula  whitneyi;  end  of  the  abdomen,  dorsal  aspect;  8,  9= 

eighth  and  ninth  segments. 
Fig.  24.  Pupa  of  Tipula  whitneyi;  end  of  the  abdomen,  ventral  aspect;  5,  6,  7= 

fifth,  sixth,  and  seventh  segments. 
Fig.  25.  Larva  of  Tipula  sp. ;  second  thoracic  segment,  dorsal  aspect,  showing 

the  distribution  of  the  setae. 
Fig.  26.  Larva  of  Tipula  sp. ;  fifth  abdominal  segment,  dorsal  aspect,  showing 

the  distribution  of  the  setae. 
Fig.  27.  Larva  of  Tipula  sp. ;  stigmal  field,  caudal  aspect. 
Fig.  28.  Larva  of  Tipula  sp. ;  stigmal  field,  dorso-caudal  aspect 


DIPTERA. 

(Except  TIPULIDAE,  RHYPHIDAE,  and  CALLIPHORIDAE.) 

By  J.  R.  Malloch,  Assistant  Biologist,  Bureau  of  Biological  Survey. 

(Plates  XII-XV.) 

The  present  collection  contains  a  very  large  number  of  specimens 
but  is  not  particularly  rich  in  species.  Moreover,  there  is  nothing  very 
remarkable  in  the  material,  the  only  genus  that  has  not  previously 
been  recorded  from  Alaska  being  Smittia,  a  genus  of  Chironomidae 
described  from  the  Arctic  regions  of  Europe  (Spitzbergen). 

I  have  taken  the  opportunity  of  indicating  in  the  introductory 
notes  to  each  family  what  the  known  larval  habits  of  the  species  are, 
considering  that  this  information  may  have  a  certain  value  in  a  list 
of  this  nature  even  though  it  does  not  refer  directly  to  the  species  in 
the  list. 

The  arrangement  is  that  of  the  Aldrich  Catalogue,  but  there  are 
several  changes  in  generic  names. 

Suborder  Orthorrhapha, 

Division  NEMATOCERA. 

Family  CHIRONOMIDAE. 

The  larvae  of  most  of  the  members  of  this  family  are  aquatic  in 
habit  and  those  in  the  present  oollection  are  representatives  of  that 
section,  belonging  entirely  to  the  subfamily  Chironominae,  though  it 
is  not  improbable  that  some  of  them  may  be  terrestrial  as  is  the  case 
with  C amptocladius  byssinus  and  some  others.  All  of  them  except 
Smittia  may  be  located  generically  by  the  use  of  the  keys  in  my  paper 
on  Illinois  Chironomidae  subsequently  cited.  I  have  made  figures  of 
the  male  genitalia  of  the  species  described  herein  to  prevent  any  mis- 
conceptions by  future  workers  on  the  group  as  to  their  identity. 

Genus  Chironomus  Meigen. 

There  is  a  striking  similarity  in  the  species  of  Chironomus  in  the 
collection.  All  are  black,  have  the  basal  joint  of  the  fore  tarsi  very 
little  longer  than  the  fore  tibiae,  and  the  males  of  all  have  the  fore 
tarsi  with  long,  soft  hairs.  Fortunately,  the  hypopygia  of  the  males 
present  in  their  structure  good  characters  for  differentiation  of  the 
species. 
170 


1923.]  INSECTS   OF   THE  PRIBILOF   ISLANDS.  171 

Synopsis  of  species. 

1.  Males 2. 

Females 4. 

2.  Hypopygium    with    the    superior    processes    indistinguishable,    the    inferior 

pair    very    small,    apex    of    lateral    arm    obtusely    truncate     (PI.    XIII, 

fig.   6) obtusilobus. 

Hypopygium  with  superior  and  inferior  processes  present,  the  latter  well 
developed,  apex  of  lateral  arm  pointed  or  rounded 3. 

3.  Superior  process  of  hypopygium  small,  obtuse,  a  distinct  fasciculus  of  hairs 

above  it  under  the  dorsal  plate,  the  apical  process  of  the  latter  stout,  apex 

of  lateral  arm  rounded  (PI.  XIII,  fig.  16) deviatus. 

Superior  process  of  hypopygium  large,  sickle-shaped,  no  fasciculus  above  it, 
apical  process  of  dorsal  plate  slender,  apex  of  lateral  arm  pointed  (PI. 
XIII,  fig.  13) conformis. 

4.  Small  species  3-4  mm.  in  length ;  mesonotum  with  a  conspicuous  pale  spot  on 

anterior  lateral  angles deviatus. 

Larger  species  5-7  mm.  in  length ;  thorax  black 5. 

5.  Wings  brownish,  veins  very  distinct ;  posterior  branch  of  radius  arcuate  at 

tip conformis. 

Wings  whitish,  veins  indistinct ;  posterior  branch  of  radius  straight  or  very 
slightly  arcuate obtusilobus. 

The  males  of  all  three  species  run  down  to  caption  11  in  my  key  to 
Group  B,  Subsection  I  of  Chironomus.2  The  structure  of  the  hypo- 
pygia  readily  separates  them  from  the  species  therein  included  ex- 
cept in  the  case  of  conformis  which  has  the  hypopygium  similar  to 
that  of  quadripunctatus ;  the  latter,  however,  has  yellow  legs. 

Chironomus  obtusilobus,  n.  sp. 

Male. — Black,  opaque.  Antennal  plumes  fuscous.  Legs  fuscous, 
tibiae  and  tarsi  yellowish  or  pale  brown.  Wings  whitish,  veins 
rather  indistinct,  cross  vein  faintly  darkened.  Halteres  testaceous. 
Hairs  on  abdomen  and  legs  pale  brown. 

Hypopj^gium  as  in  Plate  XIII,  Figure  6,  the  lateral  arms  rather 
stouter  than  normal.  Fore  tarsi  with  long  hairs,  basal  joint  slightly 
longer  than  tibiae  (120:113),  second  joint  not  greatly  longer  than 
third  (35:  30).  Radius  almost  entirely  straight,  ending  as  far  before 
apex  of  wing  as  media  does  behind  it. 

Female. — Agrees  with  the  male  in  color.  Fore  tarsi  without  long 
hairs;  legs  stouter  than  in  male. 

Length,  7 — 8.5  mm. 

Type  locality. — St.  George  Island,  June  17,  1914  (G.  D.  Hanna, 
lot  40),  4  males  caught  on  the  wing  among  bog  plants,  border  of 
Gull  Lake,  beside  Staraya  Artel  Rookery.  Paratypes  from  St.  Paul 
Island,  9  specimens,  July  22,  1913,  Big  Lake  (A.  G.  Whitney,  lot 
92) ,  "  collected  from  our  coats  as  they  swarmed  past  us  while  driving 
along  the  lake";  18  specimens,  same  date  and  place  (A.  G.  Whitney, 

*  Bull.  Illinois  State  Lab.  Nat  Hist.,  vol.  10,  art.  6,  1915,  p.  417. 


172  NORTH   AMERICAN   FAUNA.  [No.  46 

lot  91),  "swarms  of  these  insects  were  driven  by  the  breeze  south- 
eastward off  of  the  lake.  These  clouds  of  gnats  noticeable  through- 
out July  and  August. " 

Chironomus  deviatus,  n.  sp. 

Male. — Black,  shining,  without  pruinescence  on  thorax.  Meso- 
notum  slightly  yellowish  on  anterior  lateral  angles.  Legs  fuscous, 
tibiae  and  tarsi  paler.  Wings  whitish,  veins  pale,  cross  vein  not 
infuscated,  a  black  longitudinal  streak  on  base  of  wing  as  in  Cam- 
optodadius.  Halteres  testaceous.  Antennal  plumes  and  hairs  on 
body  and  legs  fuscous. 

Hypopygium  as  in  Plate  XIII,  Figure  16 ;  the  fasciculus  of  hairs 
below  dorsal  plate  is  more  characteristic  of  the  hypopygia  of  Tany- 
tarsus  than  of  Chironomus  but  the  wings  are  not  hairy.  Basal  joint 
of  fore  tarsi  slightly  longer  than  tibia  (50:42),  second  joint  dis- 
tinctly longer  than  third  (29:21) ;  fore  tarsi  and  mid  and  hind  legs 
with  long  hairs.  Wings  narrow ;  radius  ending  appreciably  farther 
in  front  of  apex  of  wing  than  does  media  behind  it. 

Female. — Differs  from  the  male  in  having  the  anterior  lateral 
margins  of  the  thorax  with  a  conspicuous  yellow  or  greenish  spot,  the 
legs  much  paler,  and  the  base  of  the  wings  noticeably  yellowish. 
Fore  tarsi  without  long  hairs. 

Length,  3.25-4.5  mm. 

Type  locality. — St.  Paul  Island,  Laboratory  (A.  G.  Whitney,  lot 
39),  3  males  and  1  female.  "From  a  great  many  on  window;  prob- 
ably hatched  from  native  Sagina  sod  transplanted  to  laboratory  a 
few  days  before." 

The  specimens  taken  on  this  occasion  did  not  represent  a  single 
species,  as  a  male  of  an  Orthocladius  bears  the  same  lot  number.  I 
regret  that  more  specimens  were  not  taken,  as  it  is  not  improbable 
that  there  were  more  than  2  species  present. 

Chironomus  conforcnis,  n.  sp. 

Male. — Deep  black,  opaque.  Antennal  plumes  fuscous;  thorax 
with  slight  indications  of  3  longitudinal  grayish  pruinescent  lines; 
abdomen  with  faint  brownish  posterior  margins  to  segments;  legs 
fuscous,  tibiae  and  tarsi  yellowish;  wings  slightly  brownish,  veins 
distinct,  cross  vein  darkened;  halteres  yellowish  brown;  hairs  on 
body  black,  on  legs  brownish. 

Hypopygium  similar  to  that  of  decorus  Johannsen,  the  superior 
process  and  apical  portion  of  lateral  arm  as  in  Plate  XIII,  Figure  13. 
Basal  joint  of  fore  tarsi  very  little  longer  than  fore  tibia  (82  :  78), 
second  joint  very  much  longer  than  third  (52  :  30) ;  fore  tarsi  and  mid 
and  hind  legs  with  very  long  and  rather  dense  hairs.  Radius 
slightly  arcuate  apically,  ending  as  far  before  apex  of  wing  as  does 
media  behind  it. 


1923.]  INSECTS  OF   THE  FRIBILOF  ISLANDS.  173 

Female- — Agrees  in  color  with  male.  Fore  tarsi  without  long 
hairs ;  basal  joint  about  as  long  as  fore  tibia. 

Length,  8-9  mm. 

Type  locality.— St.  Paul  Island,  August  16,  1914  (E.  A.  Preble), 
2  males,  1  female ;  paratypes,  1  male,  1  female,  same  island,  June  5, 
1913  (A.  G.  Whitney,  lot  37),  laboratory.  I  have  before  me  a  para- 
type  from  Admiralty  Bay,  Alaska,  June  27,  belonging  to  the  Phila- 
delphia Academy  of  Natural  Sciences.  I  had  purposed  describing 
the  species  from  this  specimen  but  take  the  present  opportunity  of 
doing  so  in  company  with  this  additional  material  and  make  the 
Admiralty  Bay  specimen  a  paratype. 

Genus  Tanytarsus  Van  der  Wtilp. 

There  is  one  species  of  this  genus  amongst  the  present  material,  sim- 
ilatus, which  was  originally  described  by  the  present  writer,  in  the 
paper  cited  under  the  previous  genus,  from  material  obtained  at  Madi- 
son, Wisconsin.  The  species  described  on  a  previous  page  as  Chiron- 
omus  deviatus  has  the  hypopygial  characters  of  Tanytarsus,  but  I 
can  not  detect  any  surface  hairs  on  the  wings  and  pending  the  receipt 
of  better  specimens  I  leave  it  in  Chironomus,  although  inclined  to 
consider  it  as  possibly  belonging  to  the  present  genus. 

Tanytarsus  similatus  Malloch. 

Tanytarsus  similatus,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist,  vol.  10, 
art.  6,  p.  494,  1915. 

In  my  original  description  of  this  species  I  stated  that  the  hypopy- 
gium  was  similar  to  that  of  viridiventris,  differing  in  the  structure 
of  the  superior  and  inferior  processes.  The  dorsal  plate  of  my 
specimen  appeared  to  have  the  apical  extension  broken  off  and  I  did 
not  figure  it.  In  the  example  before  me  I  find  that  the  dorsal  plate 
is  the  same  as  in  the  type  and  presents  an  added  character  for  dis- 
tinguishing the  species.     (PL  XIII,  fig.  11.) 

Locality,  St.  Paul  Island,  1  male  and  2  females,  July  22,  1913 
(A.  G.  Whitney,  lot  93)  ;  Big  Lake,  "  Collected  at  same  time  as  larger 
ones  [Chironomus  conformis~\  from  lake  shore." 

Genus  Orthocladius  Van  der  Wulp. 

Most  of  the  specimens  of  this  family  in  the  present  collection  be- 
long to  the  genus  Orthocladius.  A  few  of  the  examples  are  in  fairly 
good  condition  and  are  identifiable,  but  the  greater  portion  are 
poorly  preserved  and  in  that  condition  are  impossible  of  identifica- 
tion, or  at  least  are  not  in  condition  that  warrants  my  giving  a  spe- 
cific name  to  them,  because  of  the  very  large  number  of  extremely 
closely  allied  forms  occurring  in  the  genus  and  the  liability  to  error. 
It  is  possible  for  me  to  identify  the  following  species. 
108731°— 23 12 


174  NORTH   AMERICAN   FAUNA.  [No.  46 

Orthocladius  obumbratus  Johannsen. 

Orthocladius  obumbratus  Johannsen,  Bull.  86,  New  York  State  Museum, 
p.  281,  1905. 

This  species  is  represented  by  a  large  number  of  specimens  in  the 
collection.  Johannsen  described  the  species  from  examples  obtained 
at  Ithaca,  N.  Y.,  and  Douglas,  Alaska.  A  comparison  of  the  hypo- 
pygia  of  an  Ithaca  specimen  sent  me  by  Prof.  Johannsen  and  one 
of  those  from  St.  George  Island  shows  that  they  are  the  same  species. 
I  give  herewith  (PL  XIII,  fig.  10)  a  figure  of  the  apical  prolonga- 
tion of  the  dorsal  plate  of  the  hypopygium;  in  nivoriundus  and 
most  allied  species  this  plate  is  without  conspicuous  hairs. 

The  data  on  specimens  in  collection  is  as  follows:  St.  George 
Island,  35  specimens  June  17, 1914  (lot  37,  G.  D.  Hanna). 

Orthocladius  nivoriundus  Fitch. 

Chironomus  nivoriundus  Fitch,  Winter  Insects  of  Eastern  New  York,  p. 
274,  1846. 

Not  so  numerously  represented  as  the  foregoing. 
St.  George  Island: 

1  specimen,  June  5,  1913  (lot  39,  A.  G.  Whitney). 
1  specimen,  June  10,  1914  (lot  11,  G.  D.  Hanna). 
8  specimens,  June  17,  1914  (lot  37,  G.  D.  Hanna). 

Orthocladius  sp.  I. 

A  male  specimen  in  rather  poor  condition  has  the  hypopygium 
differing  from  that  of  nivoriundus  in  having  no  distinct  extension 
of  the  dorsal  plate.  This  may  be  the  result  of  an  accident  as  the 
normal  extension  is  easily  broken. 

St.  George  Island:  1  specimen  June  17,  1914  (lot  37,  G.  D.  Hanna). 

Orthocladius  sp.  II. 

A  male  specimen  lacking  front  tarsi  and  otherwise  in  poor  condi- 
tion has  the  hypopygium  very  different  from  that  of  nivoriundus. 
Undoubtedly  the  example  belongs  to  a  distinct  species  that  is  proba- 
bly undescribed. 

The  extension  of  the  dorsal  plate  is  very  long  and  slender,  and 
the  apical  shoe-shaped  portion  of  the  lateral  arm  is  much  stouter 
than  in  any  species  known  to  me. 

St.  George  Island:  1  specimen,  June  14, 1914  (lot  23,  G.  D.  Hanna). 

Orthocladius  sp.  III. 

A  male  specimen  lacking  the  apical  portion  of  the  abdomen  repre- 
sents a  species  unknown  to  me.  Differs  from  the  4  species  already 
mentioned  in  having  the  scutellum  yellow  apically.  The  fore  tarsi 
are  bare,  and  very  long,  the  basal  joint  being  four-fifths  as  long  as 
the  tibia. 

St.  George  Island:  June  5,  1913  (lot  39,  A.  G.  Whitney). 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  175 

Orthocladius  sp.  IV. 

Two  males  of  a  small  species  in  poor  condition.  The  species  re- 
sembles Trichocladius  infuscatus  Malloch,  but  the  legs  are  slightly 
yellowish. 

St.  George  Island:    June  17,  1914  (lot  37,  G.  D.  Hanna). 

Genus  Smittia  Holmgren. 

The  genus  Smittia  has  not  previously  been  recorded  from  America, 
the  only  described  species  having  been  taken  on  the  islands  of  Nova 
Zembla  (longipennis)  and  Spitzbergen  (brevipennis)  in  the  Arctic 
regions  of  the  Old  World.  Although  the  original  description  of  the 
genus  is  not  very  full  I  have  no  hesitation  in  placing  the  present 
species  in  Smittia.  It  is  possible,  though  not  at  all  probable,  that  the 
Alaskan  species  is  the  same  as  that  described  from  Spitzbergen,  but 
I  am  not  able  to  find  in  the  description  of  the  latter  confirmation 
that  is  necessary  to  permit  me  arriving  at  a  decision  that  they  are 
identical.  I  am  therefore  describing  the  present  species  as  new.  I 
have  also  taken  this  opportunity  of  redescribing  the  genus  and  indi- 
cating its  true  position  in  the  Chironominae. 

Description  of  Genus. 

Male. — Antenna  apparently  10-jointed  (2-J-8),  basal  and  apical 
flagellar  joints  elongated,  joints  5,  6,  and  7  not  separated  on  their 
entire  circumference  (PI.  XIII,  fig.  7),  flagellar  hair  very  short,  the 
longest  not  exceeding  the  apical  joint  in  length;  palpi  4-jointed,  base 
slightly  tuberculate,  basal  joint  about  as  broad  as  long  and  half  as 
long  as  second,  joints  2,  3,  and  4  subequal;  fore  tarsi  with  the  basal 
joint  shorter  than  fore  tibia;  hypopygium  as  in  Orthocladius  (sens, 
lat.)  and  Camptocladius,  the  apical  portions  of  lateral  arm  re- 
curved (fig.  5),  venation  as  in  Orthocladius. 

Female. — Differs  from  the  male  in  having  the  antenna  (PI.  XIII, 
fig-  8)  7-jointed  (2+5). 

Keiffer  has  indicated  that  Orthocladiariae  may  be  separated  from 
the  group  which  contains  Chirononnus  by  the  nature  of  the  arma- 
ture of  the  apex  of  the  hind  tibia.  In  Orthocladius  there  is  a  dis- 
tinct spur,  while  in  Chironomus  and  allied  genera  there  is  a  comblike 
series  of  setulae.  In  the  present  species  the  hind  tibial  characters 
and  also  those  of  the  hypopygium  are  those  of  the  Orthocladius  group. 
The  genus  as  a  more  specialized  form  should  be  placed  after  that 
genus  in  our  lists. 

Smittia  arctica,  n.  sp. 

Male  and  female. — Black,  opaque.  Halteres  obscurely  yellowish. 
Legs  piceous.     Wings  whitish,  veins  yellow. 

Male. — Antennal  sensory  organs  pale,  hairlike  (PL  XIII,  fig.  7)  ; 
eyes  bare.    Pronotum  linear,  very  slightly  notched  centrally;  meso- 


176  NOBTH   AMERICAN   FAUNA.  [No.  46 

notum  with  3  longitudinal  series  of  weak  hairs  on  disc,  the  central  se- 
ries weakest.  Abdomen  1.5  times  as  long  as  head  and  thorax  combined ; 
hypopygium  as  Plate  XIII.  Figure  5.  Basal  joint  of  fore  tarsi 
slightly  over  half  as  long  as  tibia  (26:47);  legs  with  very  short 
hairs ;  claws  flattened  at  apices  but  without  distinct  apical  incisions ; 
empodium  slender,  as  long  as  claw,  distinctly  fringed.  Wings  not 
extending  to  apex  of  fifth  abdominal  segment;  costa  extending  to 
apex  (PI.  XIII,  fig.  12).  / 

Female. — Antenna  as  in  Plate  XIH,  Figure  8.  Abdomen  stout. 
Basal  joint  of  fore  tarsi  distinctly  over  half  as  long  as  fore  tibiae 
(15  :25)  ;  claws  acute  at  apices.     Wing  veins  not  so  thick  as  in  male. 

Length,  1.75-2.25  mm. 

Type  specimen. — St.  Paul  Island,  May  23,  1914  (A.  G.  Whitney, 
lot  143). 

Paratopes. — St.  Paul  Island,  6  males  and  7  females,  May  23,  1914, 
from  outside  of  laboratory  window  (A.  G.  Whitney,  lot  152)  ;  1 
female,  May  17,  1913  (A.  G.  Whitney,  lot  29)  ;  1  female,  July  9,  1913 
(A.  G.  Whitney,  lot  67)  ;  St.  George  Island,  2  females,  June  16, 
1914  (G.  D.  Hanna,  lot  32),  taken  by  sweeping  toward  East  Rookery; 
and  1  female,  June  17,  1914  (G.  D.  Hanna,  lot  37),  Staraya  Artel 
Rookery. 

The  type  and  lot  67  are  on  slides,  the  others  are  mounted  on  card 
points. 

In  my  paper  on  the  Chironomidae  of  Illinois3  the  genus  Smittia 
runs  down  to  Orthocladius  (sens,  lat.)  and  is  readily  separated  from 
any  of  the  subgenera  therein  contained  by  the  very  short  wings  and 
the  number  of  antenna!  joints  in  both  sexes. 

Family  MYCETOPHILIDAE. 

The  larvae  of  most  of  the  species  of  this  family  feed  upon  fungi 
and  decaying  vegetable  matter,  some  of  them  occurring  in  colonies 
under  bark  of  dead  trees  or  fence  posts.  The  flies  are  usually  dim- 
cult  to  collect  except  by  sweeping  amongst  overhanging  bushes  or 
grasses  or  at  lights,  though  at  times  they  may  be  found  in  numbers 
on  fungi  or  on  the  inner  sides  of  windows  of  out-houses  or  buildings. 
There  are  only  seven  specimens  of  the  family  in  the  present  collec- 
tion, representing  five  genera.  Unfortunately,  a  specific  identifica- 
tion is  not  possible  in  two  cases  owing  to  the  poor  condition  of  the 
specimens. 

Genus  Macrocera  Meigen. 

There  are  9  species  of  the  genus  Macrocera  described  from  North 
America,  none  of  which  have  been  recorded  from  Alaska.     The  only 

» Bull.  Illinois  State  Lab.  Nat.  Hist,  vol.  10,  art.  6,  1915. 


1923.]  INSECTS  OF   THE  PRIBILOF  ISLANDS.  177 

example  of  the  genus  in  the  present  collection  apparently  belongs  to 
an  undescribed  species. 

Macrocera  beringensis,  n.  sp. 

Male. — Glossy  black-brown.  Mouth  parts  and  basal  2  antennal 
joints  yellowish ;  antennal  flagellum  shining,  black.  Prothorax, 
pleural  sutures,  and  scutellum  yellowish,  remainder  of  thorax  glossy 
black-brown.  Abdomen  unicolorous  black-brown.  Legs  testaceous, 
apices  of  mid  and  hind  coxae,  and  the  tarsi  infuscated.  Wings 
slightly  grayish,  a  large  fuscous  spot  over  petiole  of  media ;  another 
between  the  branches  of  cubitus  touching  the  posterior  branch  along 
its  apical  half  and  not  extending  to  anterior  branch;  a  similarly 
colored,  slightly  curved,  fasciform  spot  between  middle  of  wing  and 
apex,  the  posterior  extremity  of  which  covers  the  apical  half  of  an- 
terior branch  of  cubitus  and  the  anterior  one  extending  to  fork  of 
radius;  apical  spot  rather  faint.  Halteres  testaceous,  more  or  less 
tinged  with  brown. 

Antenna  not  over  1|  times  as  long  as  entire  body,  flagellum  rather 
thick  and  from  fourth  joint  to  apex  distinctly  hairy,  basal  flagellar 
joint  about  one-fourth  longer  than  second  and  slightly  longer  than 
third ;  median  ocellus  as  large  as»lateral ;  f  rons  with  a  distinct  median 
furrow.  Hypopygium  stout,  apex  of  lateral  arms  each  with  a  stout 
black  thorn  on  inner  angle.  Legs  long  but  not  particularly  slender; 
basal  joint  of  fore  tarsus  two-thirds  as  long  as  fore  tibia  and  slightly 
longer  than  the  remaining  tarsal  joints  combined.  Petiole  of  media 
very  short,  about  3  times  as  long  as  its  own  diameter;  costa  extend- 
ing almost  to  apex  of  wing ;  its  last  section  about  4  times  as  long  as 
its  penultimate  one — the  one  preceding  fork  of  radius ;  disk  of  wings 
without  distinct  hairs,  veins  except  the  anal  one  with  setulose  hairs. 

Length,  6  mm. 

Type—  St.  Paul  Island,  Summer,  1914   (E.  A.  Preble). 

Genus  Boletma  Staeger. 

A  single  female  specimen  of  a  species  of  this  genus  is  contained  in 
the  collection  but  is  in  such  poor  condition  that  its  identity  is  un- 
certain. In  most  particulars  it  agrees  with  beringensis  Coquillett. 
It  is  an  abnormal  specimen  in  so  far  as  its  wing  venation  is  con- 
cerned, one  wing  having  the  base  of  the  anterior  branch  of  media 
absent  while  the  other  has  it  present.  As  this  character,  the  absence 
or  presence  of  this  portion  of  this  vein,  is  used  as  a  generic  one, 
this  departure  in  the  present  case  is  worth  recording. 

Locality.— St.  George  Island,  June  27, 1914  (G.  D.  Hanna,  lot  49) ; 
toward  Zapadni. 


178  NORTH   AMERICAN   FAUNA.  [No.  46 

Boletina  obesula  Johannsen. 

Boletina  obesula  Johannsen,  The  Fungus  Gnats  of  North  America,  Bull.  196, 
Maine  Agr.  Exp.  Sta.,  Dec,  1911,  p.  276. 

Recorded  by  Cole  (Proc.  Calif.  Acad.  Sciences,  4th  ser.,  vol.  11, 
p.  1G9,  Nov.,  1921)  from  St.  Paul  Island,  the  record  being  based 
upon  one  female  taken  by  G.  D.  Hanna  June  21,  1920. 

Boletina  sp. 

One  male  of  an  "  undescribe^  species  "  recorded  by  Cole  as  "  too 
poorly  preserved  for  description  "  in  the  same  paper  as  above,  from 
St.  Paul  Island,  June  21,  1920,  G.  D.  Hanna  (p.  170). 

Genus  Rhymosia  Winnertz. 
Rhymosia  sp. 
A  female  specimen  in  poor  condition,  minus  legs  and  antennae. 
St.  Paul  Island,  August  19,  1914  (E.  A.  Preble). 

Genus  Allodia  Winnertz. 
Allodia  subelata,  n.  sp.  PI.  XIII,  fig.  9. 

Male. — Fuscous,  opaque;  flagellum  and  legs  brown;  wings  gray- 
ish, veins  brown;  halteres  testaceous  yellow;  hairs  yellow,  bristles 
blackish. 

Antenna  about  one  and  one-third"  times  as  long  as  head  and  thorax 
together;  frons  with  the  normal  soft  decumbent  hairs,  scutellar 
bristles  4  in  number;  3  bristles  above  base  of  fore  coxa  and  about 
6  at  apex  of  each.  Comparative  lengths  of  fore  tibia  and  fore 
metatarsus  28 :  23 ;  hind  tibial  setulae  very  weak ;  spurs  on  mid  and 
hind  tibiae  long  and  stout.  Hypopygium  as  in  plate  13,  figure  9. 
Furcation  of  media  distinctly  beyond  apex  of  cross  vein,  the  latter 
barely  more  than  half  as  long  as  petiole ;  furcation  of  cubitus  directly 
below  proximal  end  of  cross  vein,  the  angle  very  acute;  anal  vein 
indistinct,  subcostal  vein  very  short,  ending  in  radius. 

Length,  3  mm. 

Type.— St.  George  Island,  June  14,  1914  (G.  D.  Hanna,  Lot.  23) ; 
Garden  Cove. 

This  species  is  closely  allied  to  elata  Johannsen,  differing  in  vena- 
tion and  hypopygial  characters  chiefly. 

Genus  Exechia  Winnertz. 
Exechia  casta  Johannsen. 

Exechia  casta  Johannsen,  The  Fungus  Gnats  of  North  America,  part  4, 
Bull.  200,  Maine  Agr.  Exp.  Sta.,  p.  74,  June,  1912. 

One  male  and  2  females  in  collection  evidently  belong  to  this 
species.  The  data  on  specimens  is  as  follows :  St.  George  Island. 
June  16,  1914  (G.  D.  Hanna,  Lot  32).  Taken  by  sweeping,  toward 
East  Rookery.     I  have  made  a  drawing  of  the  male  hypopygium 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  179 

(PI.  XIII,  fig.  14)  which  shows  some  slight  differences  from  that 
given  by  Johannsen. 

Johannsen's  specimens  were  obtained  from  the  following  localities : 
Black  Kock  Creek,  Dubois  (type),  Dinwiddie  Creek,  Hunters  Creek, 
Wyoming,  in  September. 

Family  SCIARIDAE. 

The  species  of  this  family  represented  in  the  collection  belong  to 
the  genus  Sciara.  The  members  of  this  genus  are  remarkably  similar 
in  general  appearance  and  it  is  only  by  the  use  of  characters  of  wing 
venation  and  of  genitalia  that  they  can  be  differentiated  with  any- 
thing like  certainty  in  the  adult  stage.  The  larvae  which  I  have 
examined  likewise  very  closely  resemble  each  other,  and  those  I  have 
found  were  invariably  feeding  upon  decaying  vegetable  matter, 
manure,  or  in  fungi.  One  species  I  have  reared  from  fallen  plums. 
I  have  recently  described  the  larva  of  a  species  that  is  often  found 
crawling  on  the  surface  of  the  ground  in  immense  numbers  in  the 
form  of  a  rope-like  mass.4  A  number  of  other  species  in  the  family 
are  known  to  have  the  same  habit.  The  imagines,  exclusive  of  the 
Arctic  forms,  have  been  dealt  with  in  an  extensive  paper  by  Prof. 
O.  A.  Johannsen.6 

Genus  Sciara  Meigen. 

There  are  representatives  of  3  species  of  this  genus  in  the  collection. 
One  of  these  is,  I  am  confident,  identical  with  one  described  by 
Rubsaamen;  one  is  evidently  undescribed,  while  the  third  is  in  too 
poor  condition  to  permit  of  its  exact  identity  being  ascertained. 

Sciara  glacialis  FiUbsaamen.  PL  XIII,  fig.  3. 

Sciara  glacialis  Rubsaamen,  Bibl.  Zool.,  Orig. — Abh.  aus  sem  Sessannu.  d. 

Zool.  Suft.  20,  1898,  p.  109. 
Sciara  humicola  Lundbeck,  Vidensk.  Middel.  f.  d.  Naturhist.  Foren.  Kjoben- 
haven,  1898,  p.  252. 

Hypopygium  of  male  as  in  PL  13,  fig.  3.  This  species  is  evidently 
common  on  St.  George  Island,  as  it  is  represented  by  64  specimens  in 
the  collection,  with  data  as  follows : 

3  specimens,  June  10,  1914  (lot  11,  G.  D.  Hanna). 

3  specimens,  June  14,  1914  (lot  23,  G.  D.  Hanna). 
32  specimens,  June  16,  1914  (lot  32,  G.  D.  Hanna). 

1  specimen,  June  16,  1914  (lot  36,  G.  D.  Hanna). 
22  specimens,  June  17,  1914  (lot  37,  G.  D.  Hanna). 
3  specimens,  June  24,  1914  (lot  46,  G.  D.  Hanna). 

Lots  32,  36,  and  46  were  obtained  "toward  East  Rookery;"  lot  11 
from  near  beach  at  East  Landing;  lot  23  from  Garden  Cove;  and 
lot  37  Staraya  Artel  Rookery. 

*  Bull.  Illinois  State  Lab.  Nat  Hist.,  vol.  11,  art  4,  1915. 

s  The  Fungus  Gnats  of  North  America,  Part  IV,  Bull.  No.  200,  Maine  Agr.  Exp.  Sta., 
1912. 


180  NORTH   AMERICAN   FAUNA.  [No.  46 

Sciara  unguicauda,  n.  sp. 

Male. — Brownish  black,  thorax  shining,  abdomen  opaque.  Legs, 
especially  the  fore  pair,  yellowish  brown.  Wings  clear,  veins  pale 
brown.     Halteres  rufotestaceous.     Arms  of  hypopygium  reddish. 

Face  slightly  buccate;  eyes  hairy,  disc  of  mesonotum  with  short 
and  very  sparse  hairs.  Hypopygium  similar  to  that  of  glacialis,  the 
apical  portion  of  lateral  arm  stouter,  and  the  terminal  thorn  very 
strong.  First  branch  of  radius  ends  distinctly  short  of  furcation  of 
media;  costa  extends  half  way  from  apex  of  radius  to  apex  of  an- 
terior branch  of  media;  petiole  of  media  subequal  in  length  to  an- 
terior branch  of  that  vein,  the  branches  not  appreciably  divergent 
apically ;  cross  vein  over  midway  from  base  of  first  branch  of  radius ; 
media  leaves  radius  distinctly  proximad  of  midway  from  base  to 
cross  vein ;  furcation  of  cubitus  slightly  proximad  of  base  of  media. 

Female. — Slightly  paler  in  color  than  the  male.  Apical  plate  of 
genitalia  nearly  twice  as  long  as  wide,  subequal  in  length  to  pre- 
apical,  and  distinctly  longer  than  basal  one. 

Length,  3-4  mm. 

Type.— St.  George  Island,  July  8,  1914  (lot  55,  G.  D.  Hanna). 
Allotype  and  paratypes,  same  data.  Paratypes,  July  4,  1914  (lot  52, 
G.  D.  Hanna)  ;  toward  Zapadni  Rookery.    Seven  specimens. 

Sciara  sp. 

A  single  male  specimen  in  rather  poor  condition  differs  from  the 
other  two  in  structure  of  the  hypopygium  (PI.  XIII,  fig.  4).  In  the 
form  of  the  apical  portion  of  the  hypopygium  it  approaches  closely 
that  of  varians  Johannsen,  a  species  described  from  Lawrence,  Kans. ; 
Ithaca,  N.  Y. ;  and  Moscow,  Idaho.  The  wings  are  in  very  poor  con- 
dition and  the  specimen  is  otherwise  in  such  a  state  that  I  can  not 
give  a  definite  identification.  The  data  connected  with  it  is  as  fol- 
lows: 

St.  George  Island,  July  8,  1914  (lot  55,  G.  D.  Hanna). 

Genus  Neosoiara  Petty. 
Neosciara  sp. 

Cole  (Proc.  Calif.  Acad.  Sciences,  4th  ser.,  vol.  11,  p.  170,  Nov. 
1921)  records  a  female  specimen  "  in  rather  poor  condition,  so  that  its 
identity  is  not  certain  "  but  "  near  tridentata  Rubs.",  St.  Paul  Island, 
June  21,  1920  (G.  D.  Hanna). 

Family  BIBIONIDAE. 

Genus  Dilophus  Meigen. 
Dilophus  tibialis  Loew. 

Diolphus   tibialis   Loew,   H.   Diptera   Amerieae   septentrionalis   indigenae. 
Centuria,  IX,  61,  1869.    Complete  Work,  vol.  2,  p.  200.     [Sitka.] 


1923.]  INSECTS   OF   THE   PRIBILOF  ISLANDS.  181 

A  single  female,  collected  on  St.  Paul  Island,  June  24,  1916,  by  G. 
Dallas  Hanna  is  referred  to  this  species  by  W.  L.  McAtee,  who  notes 
that  it  differs  from  the  typical  form  by  absence  of  yellow  on  body; 
when  males  are  available  the  species  may  prove  to  be  new. 

Division  BRACHYCERA. 

Family  LEPTIDAE. 

This  family  is  represented  in  the  present  collection  by  a  single 
species  of  the  genus  Ptiolina. 

The  larvae  of  the  known  species  of  the  family  are  largely  terres- 
trial, living  in  the  soil  in  woods  or  in  decaying  tree  stumps  and 
feeding  upon  earthworms  and  larvae  of  insects.  One  genus,  Atherix, 
is  aquatic  in  the  larval  and  pupal  stages,  living  in  flowing  water. 
The  genus  Ptiolina  occurs  in  the  larval  stage  in  Europe  in  woods, 
under  moss  or  in  the  earth.  I  have  taken  the  imagines  only  in  very 
marshy  spots  at  rather  high  altitudes. 

Genus  Ptiolina  Zetterstedt. 

This  genus  is  separable  from  Spania  by  the  structure  of  the  third 
antennal  joint  as  pointed  out  by  Verrall.6  It  is  highly  probable  that 
all  of  our  three  North  American  species  previously  described  belong 
to  this  genus  instead  of  Spania.  The  latter  occurs  in  the  same  situ- 
ations as  Ptiolina  in  Europe,  but  is  much  commoner. 

Ptiolina  arctica,  n.  sp. 

Male  and  female. — Brownish  black,  slightly  shining.  Immature 
specimens  yellowish  brown.  Wings  clear  or  slightly  grayish. 
Halteres  brown  or  yellowish. 

Male. — Eyes  large,  closely  contiguous  for  a  short  distance;  ocelli 
situated  upon  a  distinct  elevation ;  space  above  antennae  subtriangu- 
lar;  eyes  widely  divergent  posteriorly  on  sides  of  face;  basal  2  an- 
tennal joints  short,  subequal,  short-haired  above,  third  joint  missing; 
palpi  broad,  rather  hairy.  Thorax  with  rather  short  and  sparse 
hairs,  most  noticeable  on  the  anterior  lateral  and  posterior  portions ; 
scutellum  convex,  rounded  in  outline,  surface  hairs  of  moderate 
length,  not  dense.  Abdomen  more  conspicuously  hairy  than  thorax ; 
hypopygium  chitinised,  lateral  arms  stout,  symmetrical,  rounded 
apically,  not  hairy.  Legs  moderately  stout,  their  surfaces  with  short 
hairs;  mid  and  hind  tibae  with  apical  spurs.  Venation  normal; 
fork  of  third  vein  frequently  evanescent  at  base ;  branches  of  media 
not  fused  at  apex  of  discal  cell;  anal  cell  closed. 

Female. — Agrees  in  color  with  the  male. 

'British  Flies,  vol.  5,  p.  316,  1909. 


182  NORTH   AMERICAN   FAUNA.  [No.  46 

Eyes  very  widely  separated;  third  antennal  joint  much  longer 
than  broad,  its  apex  tapering;  style  terminal,  as  long  as  third  ioint. 
Apex  of  abdomen  slightly  tapering.     Otherwise  as  male. 

Length  7-8  mm. 

Type  locality. — St.  George  Island,  August  4,  1914  (G.  D.  Hanna). 

This  species  is  larger  than  any  of  those  previously  described  from 
North  America.  Coquillett  has  recorded  Spania  edeta  Walker,  from 
Alaska.  This  species  is  velvety  black  and  has  only  a  length  of  4 
mm.  Neither  of  Loew's  species  can  be  identical  with  the  one  here 
described,  differing  both  in  color  and  venation. 

The  specimens  before  me  are  in  poor  condition  either  having  been 
wet  or  at  one  time  in  alcohol,  and  are  much  discolored.  Some  of 
them  were  evidently  immature  when  captured.  The  thorax  may  be 
in  well  preserved  examples  more  or  less  distinctly  vittate  as  traces 
of  vittae  are  visible  in  one  or  two  of  those  in  this  collection.  The 
venation  is  like  that  of  most  Leptidae,  rather  unstable. 

The  larva  of  this  species  has  not  been  described.  A  vial  num- 
bered 14116,  containing  2  larvae  from  the  stomach  of  Plectrophenax 
nivalis  townsendi,  St.  Paul  Island,  June  19,  1890,  is  before  me  and 
the  specimens  are,  I  believe,  referable  to  this  species.  The  descrip- 
tion is  as  follows: 

Length,  8-9  mm.  White,  cephalic  parts  dark  castaneous.  Body 
cylindrical,  tapered  on  prothorax  and  mesothorax,  and  flattened  on 
dorsal  surface  of  apical  segment.  Head  of  the  same  general  structure 
as  that  of  Chrysopila,  differing  in  having  the  labrum  much  broader, 
blunt  at  apex,  *he  sides  slightly  tapered  anteriorly,  and  the  dorsum 
slightly  ridged  transversely  on  anterior  half;  antennae  short  and 
stout,  not  twice  as  long  as  thick,  terminal  joint  very  minute;  between 
the  antennae  and  the  labrum  there  is  on  each  side  a  large  pale  mem- 
branous area,  the  surface  of  which  is  granular;  maxillae  large, 
almost  entirely  pale  and  membranous,  palpi  much  smaller  than  an- 
tennae; mandibles  stout,  slightly  hooked  at  apex;  posterior  dorsal 
arcuate  shield  about  1£  times  as  long  as  broad,  rounded  posteriorly; 
internal  cephalic  rods  extending  to  posterior  margin  of  head. 
Thoracic  and  abdominal  locomotor  organs  not  easily  distinguishable 
in  specimens  owing  to  condition,  but  evidently  consisting  of  slightly 
raised  transverse  areas  on  venter,  similar  to  those  on  Chrysopila, 
which  are  armed  with  very  small,  sharp  toothlike  elevations;  apical 
segment  slightly  longer  than  its  basal  breadth,  rounded  from  near 
middle,  its  apex  with  2  small  upwardly  directed,  slightly  chitinized 
teeth,  which  are  separated  by  a  distance  greater  than  the  height  of 
either  tooth,  dorsum  with  6  longitudinal  grooves  or  furrows  which 
do  not  extend  to  apex,  and  give  a  ridged  appearance  to  the  segment; 
venter  of  apical  segment  with  flat  elevation  extending  from  base  to 


1923.]  INSECTS   OF   THE  PRIBILOF  ISLANDS.  183 

near  apex,  its  apical  outline  in  the  form  of  2  rounded  lobes,  slightly 
cephalad  of  middle  of  this  flat  area  is  the  anal  opening  which  is  very 
distinct,  and  oval  in  shape ;  between  the  apex  of  the  above  elevation 
and  extreme  apex  of  segment  there  is  a  slight  but  distinct  incision 
parallel  to  apical  margin  of  segment. 

Family  EMPIDIDAE. 

The  larvae  of  the  great  majority  of  the  species  in  this  family 
are  terrestrial  and  feed  upon  vegetable  matter  in  the  soil  or  in 
rotten  wood  or  upon  larvae  or  other  small  animals.  One  species 
has  been  found  in  the  larval  and  pupal  stages  in  running  water  in 
New  York  State.  The  adult  females  are  predaceous  but  many 
species  are  found  in  large  numbers  upon  flowers  of  various  plants. 
The  two  genera  represented  in  the  present  collection  are  the  largest 
in  point  of  numbers  in  the  family,  and  are  the  most  widely  dis- 
tributed and  common.  Coquillett  has  described  or  recorded  about 
20  species  of  the  genus  Rhamphomyia  and  seven  species  of  Empis 
from  Alaska. 

Genus  Empis  Linnaeus. 

There  are  four  species  of  this  genus  in  the  collection,  all  in  rather 
poor  condition  and  all  represented  by  females  only.  With  one 
exception  the  species  in  this  collection  are  referable  to  the  group 
that  contains  virgata  Coquillett. 

Coquillett7  recorded  Empis  virgata  Coquillett  from  Alaska  and 
pellucida,  fumida,  and  infumata  were  described  by  him  at  the  same 
time,  all  being  referred  to  as  resembling  virgata  and  only  the  first 
being  described  fully,  the  others  being  briefly  compared  with  it.  I 
had  4  specimens  before  me  that  belong  to  the  same  group  as  pellu- 
cida;  because  of  the  rather  meagre  descriptions  I  was  unable  to 
satisfactorily  identify  the  species.  I  was  obliged,  therefore,  to  borrow 
paratypes  of  Coquillett's  species  from  the  United  States  National 
Museum  for  comparison.  I  have  drawn  up  a  key  to  the  species 
(females),  which  is  presented  herewith,  using  characters  not  men- 
tioned in  the  original  descriptions. 

All  of  the  species  are  black  in  color,  the  legs  sometimes  brownish 
in  peUucida,  the  mesonotum  either  trivittate  or  quadrivittate,  the 
vittse  shining  and  the  intervening  spaces  grayish  pruinose;  legs 
with  very  few  weak  spines,  not  feathered;  halteres  yellow.  There 
is  a  very  great  similarity  between  the  5  species,  and  the  synopsis 
now  given  embodies  practically  all  the  essential  characters  useful 
for  their  separation. 

7  Proc.  Washington  Acad.  Sci.,  vol.  2,  p.  408,   1900. 


184  NORTH  AMERICAN-  FAUNA.  [No.  46 

Key  to  species  of  virgata  group. 

1.  Hind    femora    with    several    distinct   bristles    on    antero-dorsal    surface   of 

apical  third ;  tibial  bristles  strong,  those  on  hind  tibiae  longer  than  the 
diameter  of  the  tibia ;  notopleural  bristles  5  in  number,  the  anterior  one 
weak,  the  posterior  4  strong,  subequal  in  length  and  rather  closely 
placed,  the  series  in  a  straight  line ;  third  antennal  joint  tapering  except 
on  a  short  space  at  base;  fork  of  third  vein  about  twice  as  long  as  the 

section  of  costa  preceding  it fumida  Coquillett. 

Hind  femora  without  distinct  bristles  on  apical  third  of  antero-dorsal  sur- 
face, only  the  normal  short  hairs  present;  hind  tibial  bristles  not  longer 
than  the  tibial  diameter ;  notopleural  bristles  rather  weak,  widely  placed 
and  usually  3  in  number,  fork  of  third  vein  not  twice  as  long  as  section 
of  costa  preceding  it 2. 

2.  Fore  coxa   with   very  few   widely   placed   hairs  on   anterior  surface,   seen 

from  the  side  only  about  a  dozen  visible ;  hind  femora  and  tibiae  slender, 
the  former  almost  nude  on  anterior  surface ;  distance  between  apices 
of  third  vein  and  first  branch  of  media  distinctly  greater  than  that  be- 
tween branches  of  media,  measured  along  margin  of  wing,  because  of 
a   distinct   deflection   of  the   first   branch   of   media   just  below   fork   of 

third pellucida  Coquillett. 

Fore  coxae  with  numerous  rather  long,  soft  hairs  anteriorly 3. 

3.  Thorax  trivittate,   the  vittae  shining,   the  spaces   between   gray   pruinose ; 

wings  slightly  infuscated  at  apices virgata  Coquillett. 

Thorax  quadrivittate,  occasionally  the  vittae  are  not  very  distinct ;  wings 
evenly  lutescent  or  pale  brownish,  not  noticeably  darker  at  apices  than 
elsewhere 4. 

4.  Proboscis  distinctly  more  than  twice  as  long  as  height  of  head ;  fork  of  third 

vein  slightly  bent  in  middle,  base  of  cell  enclosed  by  it  acute  posteriorly. 

infumata  Coquillett. 

Proboscis  less  than  twice  as  long  as  height  of  head ;  fork  of  third  vein  rather 
abruptly  bent  at  middle,  the  base  of  cell  enclosed  by  it  obtuse  posteriorly. 
subinfumata,  n.  sp. 

Empis  infumata  Coquillett. 

Empis  infumata  Coquillett,  Proc.  Washington  Acad.  Sci.,  vol.  2,  p.  409,  1900. 

Two  specimens  in  rather  poor  condition,  having  been  wet,  obtained 
June  27,  1914  (lot  49,  G.  D.  Hanna),  on  St.  George  Island,  are  evi- 
dently referable  to  this  species.  Originally  described  from  Popof 
Island,  Alaska. 

Empis  subinfumata,  n.  sp. 

Female. — A  more  robust  species  than  infumata  with  a  more  intense 
black  coloring,  the  legs  and  palpi  being  entirely  black.  The  thoracic 
hairs  are  more  conspicuous  and  the  legs  are  rather  stouter.  The  hind 
femora  and  tibia  each  possess  a  sulcus  on  the  anterior  surface 
apically,  but  this  may  not  be  evident  in  fresh,  well-matured  examples. 
Other  characters  are  mentioned  in  key. 

Length,  6  mm. 

Type  locality.— St.  George  Island,  June  16,  1914  (lot  35,  G.  D. 
Hanna).     One  specimen. 


1923.]  INSECTS   OF    THE   PRIBILOF   ISLANDS.  185 

Empis  sp. 

A  female  taken  May  1,  1913  (lot  18,  A.  G.  Whitney)  on  St.  Paul 
Island,  very  probably  represents  a  distinct  species.  It  differs  from 
subinfumata  in  having  the  mesonotum  much  more  distinctly  pruin- 
escent  and  in  having  4  notopleural  bristles,  but  is  in  such  poor  condi- 
tion that  I  do  not  deem  it  advisable  to  describe  it  as  new. 

Empis  sp. 

Cole  (Proc.  Calif.  Acad.  Sciences,  4th  ser.,  vol.  11,  Nov.,  1921,  p. 
170)  records  a  male  of  a  species  "  probably  undescribed  "  in  "  poor 
condition"  from  St.  George  Island,  June  30,  1920  (G.  D.  Hanna). 

Possibly  one  of  the  species  listed  above. 

Empis  frontalis  Coquillett. 

Empis  frontalis  Coquillett,  Proc.  Ent.  Soc.  Washington,  vol.  5,  p.  271,  1903. 

This  species  was  originally  described  from  specimens  obtained  on 
St.  George  Island  by  Professor  Kincaid  and  is  represented  in  the  col- 
lection by  2  females  also  from  that  island.  The  data  connected  with 
the  specimens  are  as  follows : 

Jnne  17,  1914  (lot  49,  G.  D.  Hanna). 
August  4,  1914  (G.  D.  Hanna). 

Genus  Rhamphomyia  Meigen. 

This  genus  is  represented  by  one  species  of  which  there  are  56  speci- 
mens in  the  collection.  I  can  not  definitely  associate  it  with  any  de- 
scribed North  American  species  and  describe  it  herewith. 

I  have  reared  one  species  of  this  genus  from  larvae  found  in  rotten 
tree-stumps. 

Rhamphomyia  opacithorax,  n.  sp. 

Male. — Black,  slightly  shining,  the  thorax  almost  entirely  opaque, 
because  of  the  dense  brownish  surface  pruinescence.  Head  entirely 
black.  Mesonotum  not  vittate,  hairs  and  bristles  black.  Abdomen 
not  so  distinctly  pruinescent  as  thorax,  the  hairs  yellowish  or  whitish. 
Legs  entirely  black,  the  femora  glossy ;  surface  hairs  fuscous.  Wings 
whitish  hyaline,  veins  brown,  stigma  brown.  Halteres  brown,  knobs 
fuscous. 

Eyes  contiguous  for  one-half  the  length  of  frons;  third  antennal 
joint  long,  tapering  from  base  to  apex;  style  nearly  one-third  as 
long  as  third  joint;  proboscis  not  over  li  times  as  long  as  height 
of  head.  Mesonotum  with  very  long  upright  hairs  laterally;  acro- 
stichals  2-rowed,  very  weak;  scutellum  with  4  bristles.  Hairs  on 
abdomen  rather  long ;  hypopygium  as  in  Plate  XIII,  Figure  15.  Legs 
moderately  stout;  basal  joint  of  fore  tarsi  slender,  not  as  thick  as 
tibia  at  apex;  hind  femora  and  tibiae  slightly  thickened,  the  lat- 


186  NORTH   AMERICAN   FAUNA.  [No.  46 

ter  appreciably  so  at  apices;  dorsal  surfaces  of  hind  tibiae  clothed 
with  rather  dense  soft  hairs  which  become  longer  from  base  to  apex ; 
basal  joint  of  hind  tarsi  thicker  than  basal  joints  of  other  tarsi 
but  slightly  thinner  than  hind  tibiae  apically,  the  dorsal  surface 
with  long  hairs.  Greatest  length  of  discal  cell  equal  to  length  of 
last  section  of  fifth  vein;  sixth  vein  incomplete;  veins  1  to  3  more 
distinct  than  other  veins. 

Female. — Differs  from  the  male  in  having  the  wings  rather  uni- 
formly pale  brownish,  all  the  veins  equally  distinct,  the  hind  legs 
less  stout  and  without  long  hairs. 

Length,  4.25-5.25  mm. 

Type  locality.— St  Paul  Island,  July  24,  1914  (lot  195,  A.  G. 
Whitney),  56  specimens. 

Rhamphomyia  sp. 

Cole  (Proc.  Calif.  Acad.  Sciences,  4th  ser.,  Vol.  II,  p.  170,  Nov., 
1921)  records  the  occurrence  of  the  female  of  a  species  "very  close 
to  conservativa  Malloch"  from  St.  Paul  Island,  August  10,  1920 
(G.  D.  Hanna). 

Family  DOLICHOPODIDAE. 

A  surprising  feature  of  the  present  collection  is  the  fact  that  there 
is  but  one  species  of  Dolichopodidae  in  it.  The  majority  of  the 
members  of  this  family  are  aquatic,  many  of  the  species  being  con- 
fined to  the  seashore.  The  single  species  represented  in  the  material 
before  me  belongs  to  a  group  that  is  invariably  aquatic  in  habit, 
the  adults  running  with  facility  on  the  surface  of  pools  of  water 
on  or  near  the  seashore.  Several  species  belonging  to  genera  closely 
allied  to  Hydrophones  are  predaceous  and  in  some  manner  the  whole 
family  has  been  credited  by  various  authors  as  being  predaceous, 
which  is  erroneous,  the  great  majority  of  species  in  the  family  feed- 
ing upon  nectar  or  other  liquids. 

Genus  Hydrophorus  Fallen. 

Hydrophorus  fumipennis  Van  Duzee. 

Hydrophorus  fumipennis  Van  Duzee,  Proc.  Cal.  Acad.  Sciences,  vol.  11,  p. 
167,  1921. 
This  species  is  represented  by  60  specimens  taken  on  St.  George 
Island,  by  G.  D.  Hanna,  and  1  specimen  by  the  same  collector  on  St. 
Paul  Island,  August  16,  1015.  The  St.  George  specimens  bear  dates 
and  lot  numbers  as  follows :  2  specimens,  June  4,  Lot  No.  2 ;  4  speci- 
mens, June  10,  Lot  No.  11 ;  9  specimens,  June  16,  Lot  No.  36 ;  4  speci- 
mens, June  17,  Lot  No.  44.  Lot  No.  2  was  taken  in  grass  and  on 
very  wet,  soggy  ground  near  Village  Landing;  Lot  No.  11  near  beach 
at  East  Landing;  Lot  No.  36  toward  East  Rookery  from  Village; 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  187 

and  Lot  44  around  a  sphagnum  bog  one-fourth  mile  west  of  Village. 
Specimens  from  St.  George  without  lot  numbers  are:  13,  September 
3,  1913;  9,  May  4,  1914;  and  19,  April  1,  1914,  G.  D.  Hanna. 
The  species  was  originally  described  from  St.  Paul  Island. 

Suborder  Cyclorrhapha. 
Division  ASCHIZA. 

Family  PHORIDAE. 

The  larvae  of  this  family  have  in  the  different  genera  very  diverse 
habits.  Some  are  internal  parasites  of  living  insects,  larvae,  pupae, 
and  imagines,  but  the  great  majority  are  scavengers,  feeding  upon 
decaying  animal  and  vegetable  matter.  A  summary  of  the  larval 
habits  appears  in  my  paper  on  the  family  printed  in  1912.8 

The  only  species  in  the  present  collection  belongs  to  the  genus 
Aphiochaeta  and  to  the  group  of  that  genus  that  feed  upon  fungi  in 
their  larval  stages. 

Genus  Aphiochaeta  Brues. 

Aphiochaeta  dubitata  Malloch. 

Aphiochaeta  dubitata  Malloch,  Proc.  U.  S.  Nat.  Mus.,  vol.  43,  p.  480,  1912. 

Six  specimens  with  data  as  follows: 
St.  George  Island. 

1  specimen,  June  16  (lot  32,  G.  D.  Hanna). 
3  specimens,  June  17  (lot  37,  G.  D.  Hanna). 

St.  Paul  Island. 

2  specimens,  August  16,  1915  (G.  D.  Hanna). 

Family  SYRPHIDAB. 

The  larvae  of  the  different  genera  of  Syrphidae  have  very  diverse 
habits;  some  are  scavengers,  others  are  aphidophagous,  while  some 
live  in  nests  of  Hymenoptera,  assumably  feeding  upon  the  detritis  of 
the  nests.  The  two  species  in  the  present  collection  are  probably 
scavengers  in  an  aquatic  or  semiaquatic  habitat  in  their  larval  states. 

Genus  Helophilus  Meigen. 

Helophilus  borealis  Staeger. 

Helophilus  borealis  Staeger,  Kroy.  Natur.  Hist.  Tidskrift,  n.  ser.  1,  p.  359, 

1845. 

This  species  which  was  originally  described  from  Greenland  is  rep- 
resented in  the  collection  by  one  female  the  data  for  which  is  August 
1,  1914  (E.  A.  Preble). 

8  Proc.  U.  S.  Nat.  Mus.,  vol.  43,  pp.  411-529,  1912. 


188  NORTH   AMERICAN  FAUNA.  [No.  46 

Helophilus  dychei  Williston. 

Helophilus  dychei  Williston  (in  Hunter),  Can.  Ent.,  vol.  29,  p.  136,  1897. 
This  species,  which  was  originally  described  from  specimens  ob- 
tained at  Sitka,  Alaska,  is  represented  by  2  males  and  1  female  in 
this  collection.    The  data  are  as  follows: 

St.  George,  August  22,  1913  (G.  D.  Hanna). 

St.  George,  June  14,  1914  (Lot  18,  G.  D.  Hanna). 

St.  Paul,  June  5,  1913  (Lot  38,/ A.  G.  Whitney). 

The  thorax  in  both  specimens  is  very  densely  long  pilose  and  the 
2  abbreviated  discal  stripes  are  very  indistinct,  almost  invisible. 

Genus  Pterallastes  Loew. 

Pterallastes  borealis  Cole. 

Pterallastes  borealis  Cole,  Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11,  p.  170, 
Nov.,  1921. 

Described  from  a  male  and  female  collected  on  St.  Paul  Island, 
August  20,  1920  (G.  D.  Hanna).  Closely  related  to  perfidiosus 
Hunter. 

Genus  Syrphus  Fabricius. 

Syrphus  contumax  Osten  Sacken. 

Syrphus  contumax  Osten  Sacken,  Proc.  Boston  Soc.  Nat.  Hist.,  vol.  18,  p. 
148,  Oct.,  1875. 

Recorded  by  Cole  (Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11,  p.  171, 
Nov.,  1921),  from  St.  George  Island,  June  30,  1920  (G.  D.  Hanna). 

Division  SCHIZOPHORA. 

Family  ANTHOMTIIDAE. 

The  larvae  of  the  species  comprising  this  family  are  mostly  phyto- 
phagous or  scavengers,  though  certain  species  are  recorded  as  para- 
sitic upon  nestlings  of  some  birds,  living  attached  to  various  parts 
of  their  bodies.  The  few  species  in  the  collection  before  me  belong 
apparently  to  the  group  that  feeds  upon  decaying  vegetable  matter; 
a  closely  allied  species  was  reared  by  me  from  rotten  wood.  The 
family  is  well  represented  in  North  America,  but  very  few  collectors 
know  more  than  a  small  percentage  of  the  species. 

Subfamily  Phaoniinae. 

Genus  Phaonia  Robineau-Desvoidy. 

This  genus  is  distinguished  from  others  of  the  subfamily  in  this 
paper  by  having  in  both  sexes  a  strong  bristle  beyond  middle 
on  the  postero-dorsal  surface  of  hind  tibia.  A  comprehensive  re- 
vision of  the  genus  by  the  writer  is  now  ready  for  the  press.  There 
is  but  one  species  known  to  me  from  the  Pribilof  Islands. 


J 923.]  INSECTS   OF   THE  PBIBILOF   ISLANDS.  189 

Phaonia  minima  Malloch. 

Phaonia  minima  Malloch,  J.  R.,  Rep.  Can.  Arctic  Exped.,  3,  Part  C,  p.  61, 
1919. 

St.  Paul  Island,  July  19,  1916  (G.  D.  Harma). 

Genus  Helina  Robineau-Desvoidy. 

The  name  Helina  has  been  used  by  me  in  all  of  my  recent  papers 
on  Anthomyiidae  for  a  segregate  of  that  group  listed  under  the 
generic  name  Mydaea  K-D.  by  Stein  and  other  authors.  The  true 
species  of  Mydaea  all  have  the  third  wing-vein  setulose  at  base  above 
and  below,  and  the  females  have  the  penultimate  abdominal  sternite 
with  short  stout  bristles.  Helina  as  at  present  limited  has  no  species 
that  possesses  the  above  combination  of  characters. 

The  species  hannai,  of  which  a  full  descripition  is  presented  herein, 
was  included  in  a  recently  published  synopsis  of  the  genus  by  the 
writer  and  a  description  also  included." 

Helina  borealis  Malloch. 

Aricia  borealis  Malloch,  J.  R.,  Rep.  Can.  Arctic  Expedition,  3,  Part  C,  pp. 
64-5.     1919. 

St.  Paul  Island,  Alaska,  July  19,  1916  (G.  D.  Hanna). 

Helina  hannai  Malloch. 

Helina  hannai  Malloch,  J.  R.,  Can.  Ent.,  vol.  53,  no.  5,  p.  109,  May,  1921. 

Puparium. — Length,  8  mm. ;  diameter  at  middle,  2.5  mm.  Color, 
reddish  testaceous,  distinctly  shining.  Cephalic  extremity  rather 
slender,  glossy ;  anterior  margin  of  first  dorsal  thoracic  segment  sub- 
carinate ;  integument  throughout  with  very  minute  longitudinal  striae, 
which  are  not  continuous  but  in  the  form  of  short,  slightly  irregular 
lines;  transverse  rugae  indistinct  anteriorly,  except  between  the  last 
thoracic  and  first  abdominal  segments,  becoming  noticeable  between 
the  third  and  fourth  abdominal  segments  and  from  that  point  be- 
coming stronger  to  apex  where  they  are  present  in  the  form  of  con- 
spicuous raised  ridges ;  lateral  f asciform  areas  distinct,  margined 
throughout  their  length  by  a  series  of  microscopic  rounded  swellings, 
which  series  is  continued  over  the  venter  in  the  form  of  a  single 
line  along  the  margin  of  each  segment;  each  ventral  segment  has  2 
short  transverse  series  of  similar  raised  areas  on  the  disc  (dorsal 
segments  glued  to  card  and  thus  invisible)  ;  anal  opening  rather 
large,  surrounded  by  a  heart-shaped  slightly  elevated  ridge  (PI.  XV, 
fig.  29) ;  anal  spiracles  consisting  of  3  slit-like  openings  on  a  slightly 
raised  base  (fig.  30). 

Imago. — Male.  Black,  distinctly  shining.  Head  black;  orbits 
with  silvery  pile ;  frontal  stripe  opaque  black.     Thorax  without  dis- 

•Can.  Ent.,  vol.  53,  p.  109,  1921. 
108731°—  23 13 


190  NORTH   AMERICAN   FAUNA.  [No.  46 

tinct  pruinescence,  with  a  faint  bronzy  tinge,  and  indistinctly  trivit- 
tate  anteriorly.  Abdomen,  when  viewed  from  behind,  with  distinct 
brownish  pruinescence  and  continuous  dorso-central  longitudinal 
black  stripe;  in  some  lights  with  a  slight  bronzy  reflection.  Legs 
black.  Wings  slightly  fuscous,  base  yellow;  cross  veins  not  infus- 
cated.    Calyptrae  bright  yellow.    Halteres  brown,  knobs  black. 

Eyes  distinctly  separated,  narrowest  part  of  stripe  about  one-tenth 
the  width  of  head  at  that  point ;  width  of  frons  at  base  of  antennae 
about  one-fourth  that  of  head  at  that  point;  orbital  bristles  strong 
and  of  moderate  length;  third  antennal  joint  over  twice  the  length  of 
second;  arista  very  short  haired;  eyes  with  very  few  microscopic 
hairs;  face  slightly  receding  toward  lower  margin;  cheeks  with 
numerous  bristly  hairs;  proboscis  rather  thick,  of  moderate  length; 
palpi  slightly  dilated  apically.  Acrostichals  irregularly  4-rowed; 
dorso-centrals  2+4.  Abdomen  subcylindrical,  elongate,  slightly  nar- 
rowed apically;  hypopygium  small,  almost  entirely  retracted;  sur- 
face hairs  on  dorsal  segments  strong,  especially  on  apical  2  segments. 
Fore  tibia  with  2-3  weak  bristles  on  the  postero-ventral  surface ;  mid 
tibia  with  2  posterior  bristles ;  hind  tibia  with  2  antero-dorsal  and  2 
antero-ventral  bristles.  Costal  spine  weak;  third  and  fourth  veins 
slightly  divergent  apically,  the  former  with  2-3  bristles  at  base ;  outer 
cross  vein  slightly  undulated;  last  section  of  fourth  vein  less  than 
1.5  as  long  as  preceding  section. 

Female. — Agrees  in  color  with  the  male  except  that  the  abdomen 
is  less  distinctly  pruinescent.  Frons  over  one-third  the  width  of 
head;  ocellar  triangle  produced  in  the  form  of  a  long  slender  point 
nearly  to  anterior  margin ;  upper  orbital  directed  outward,  the  others 
inward ;  cruciate  central  bristles  absent ;  cheeks  higher  than  in  male, 
each  about  one-third  as  high  as  eye.  Abdomen  rather  broad.  In 
other  respects  as  male.    Length,  6.5-7.5  mm. 

Four  specimens,  St.  George  Island,  June  24,  1914  (lot  46,  G.  D. 
Hanna),  1  male  and  puparium,  the  male  in  poor  condition,  June  20, 
1913  (lot  41,  A.  G.  Whitney)  ;  1  male,  abnormal  in  having  4  bristles 
on  antero-dorsal  surface  of  hind  tibia,  June  10,  1914  (lot  9,  G.  D. 
Hanna)  ;  1  male  and  2  females,  June  17,  1914  (lot  42,  G.  D.  Hanna). 

The  example  mounted  along  with  the  empty  puparium  bears  no 
data  that  throws  any  light  upon  the  habits  of  the  larva,  neither  is 
there  any  data  which  indicates  the  habits  of  the  adults  of  the  other 
lots. 

Helina  sp. 

A  female  in  very  poor  condition  differs  distinctly  from  the  fore- 
going; I  am  unable  to  identify  it  with  any  described  species.  Be- 
cause of  its  sex  and  condition,  I  refrain  from  attempting  a  de- 
scription of  the  species. 

Locality,  St.  George  Island,  July  8,  1914  (lot  55,  G.  D.  Hanna). 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  191 

Genus  Mydaea  Robineau-Desvoidy. 

Mydaea  rugia  (Walker). 

Anthomyia  rugia  Walker,  List  of  Insects  in  the  British  Museum,  Diptera, 
Part  4,  p.  923,  1S59. 

I  have  recorded  this  species  from  St.  George  Island,  July  1,  1920 
(Proc.  Calif.  Acad.  Sciences,  4th  ser.,  vol.  11,  Nov.,  1921,  p.  178). 

Genus  Melanochelia  Rondani. 

The  species  of  this  genus  resemble  those  of  Hellna  R-D.  that  have 
the  abdomen  with  paired  dorsal  abdominal  spots,  but  they  differ 
in  having  the  arista  bare  or  pubescent.  The  species  form  a  very 
homogeneous  group  and  although  rather  closely  resembling  the 
above  mentioned  group  in  Helin-a,  they  seem  also  to  differ  largely 
in  habits  of  the  adults.  The  species  of  M elanochelia  are  most  gen- 
erally found  near  streams  or  on  the  shores  of  lakes  or  of  the  sea, 
and  many  species  may  be  taken  upon  the  exposed  surfaces  of  rocks 
in  stream  beds,  especially  in  sunshine,  though  they  are  very  quick 
of  flight  and  difficult  to  detect  because  their  gray  and  black  colors 
blend  very  well  into  the  color  of  the  rocks  upon  which  they  settle. 
One  species  that  I  have  found  commonly  in  Scotland  is  predaceous 
upon  insects,  but  the  majority  of  the  adults  are  flower  frequenters. 
The  larval  habits  are  not  well  known  but  some  are  feeders  upon  de- 
caying animal  and  vegetable  matter. 

Melanochelia  nobilis  Stein. 

Limnophora  nobilis  Stein,  Berl.  Ent.  Zeitschr.,  p.  207,  1898. 

There  are  two  males  and  one  female  of  this  species  in  the  material 
before  me.  The  males,  although  in  rather  poor  condition,  agree  in 
almost  every  respect  with  Stein's  description.  The  specimen  in  the 
best  condition  shows  indications  of  thoracic  vittae  but  in  all  other 
respects  agrees  with  the  original  description.  The  female  I  am  not 
so  certain  of  but  consider  it  very  probably  as  belonging  to  nobilis. 

Localities. — Males,  St.  George  Island,  June  17,  1914  (lot  42,  G.  D. 
Hanna) ;  female,  St.  Paul  Island,  July  21,  1913  (lot  85,  A.  G. 
Whitney).  Lot  42  was  taken  on  uplands  toward  Staraya  Artel;  lot 
85  is  given  as  Webster  House,  Northeast  Point. 

The  original  description  of  this  species  was  made  from  a  male 
obtained  in  Alaska.  Coquillett  subsequently  recorded  it  from  Sitka, 
Seldovia,  and  Popof  Island,  Alaska.10 

Melanochelia  sanctipauli  Malloch. 

Melanochelia   sanctipauli   Malloch,   Proc.   Calif.   Acad.   Sciences,   4th   ser., 
vol.  11,  Nov.,  1921,  p.  180. 

Described  from  St.  Paul  Island,  July  12,  1920  (G.  D.  Hanna). 

10  Proc.  Washington  Acad.  Sci„  vol.  2,  p.  445,   1900. 


192  NORTH   AMERICAN   FAUNA.  [No.  46 

Melanochelia  spinicosta  Malloch. 

Melanochelia  spinicosta  Malloch,  Proc.  Calif.  Acad.  Sciences,  4th  ser.,  vol. 
11,  p.  181,  1921. 
Described  from  St.  Paul  Island,  and  recorded  from  St.  George 
Island,  June  30,  July  12,  and  August  10, 1920  (G.  D.  Hanna). 

Genus  Eriphia  Meigen. 
Eriphia  cinerea  Meigen. 

Eriphia  cinerea  Meigen,  Syst.  Beschr.,  5,  p.  206,  1826. 
Collected  in  1920,  by  G.  D.  Hanna  and  recently  recorded  for  the 
first  time  from  this  hemisphere  (Proc.  Calif.  Acad.  Sciences,  4th  ser., 
vol.  11,  1921,  p.  178). 

Genus  Eupogonomyia  Malloch. 

Eupogonomyia  pribilofensis  Malloch. 

Eupogonomyia  pribilofensis  Malloch,  Proc.  Calif.  Acad.  Sciences,  4th  ser., 
vol.  11,  p.  179,  1921. 

Described  from  St.  Paul  Island,  June  21,  1920  (G.  D.  Hanna). 

Genus  Hydrophoria  Robineau-Desvoidy. 

This  genus  is  represented  by  a  large  number  of  specimens  belong- 
ing to  a  single  species. 

Hydrophoria  alaskensis  Malloch.  (PI.  XV,  fig.  34.) 

Hydrophoria  alaskensis  Malloch,  J.  R.,  Can.  Ent,  vol.  52,  no.  11,  p.  257, 
Nov.,  1920. 

St.  George  Island — 

14  specimens,  June  8,  1914  (lot  7,  G.  D.  Hanna). 
9  specimens,  June  10,  1914  (lot  9,  G.  D.  Hanna). 

1  specimen,  June  16,  1914  (lot  34,  G.  D.  Hanna). 
26  specimens,  June  16,  1914  (lot  35,  G.  D.  Hanna). 

2  specimens,  May  6,  1914  (G.  D.  Hanna). 

1  specimen,  June  16,  1914  (lot  36,  G.  D.  Hanna). 
19  specimens,  June  17,  1914   (lot  42,  G.  D.  Hanna). 

2  specimens,  June  24,  1914   (lot  46,  G.  D.  Hanna). 
2  specimens,  June  25,  1914  (lot  47,  G.  D.  Hanna). 

Lot  7  has  the  following  data :  "  Found  on  several  species  of  flowers 
and  on  grass.  When  approached  it  darts  into  the  grass  but  never 
tries  to  fly  away.  Uplands."  I  am  unaware  of  this  characteristic  in 
other  anthomyids;  it  may  be  due  to  prevailing  strong  winds.  The 
remaining  lots  were  evidently  collected  by  sweeping  and  bear  no 
data  of  interest.  The  places  of  collection  were  mostly  "  towards  East 
Eookery "  and  "  towards  North  Eookery "  with  a  reference  to 
"  Uplands  "  towards  Staraya  Artel  in  the  case  of  Lot  42.  All  the 
specimens  were  taken  on  St.  George  Island.  I  have  recorded  the  oc- 
currence of  this  species  from  St.  Paul  Island,  June  21,  1920,  G.  D. 
Hanna  (Proc.  Calif.  Acad.  Sciences,  4th  ser.,  vol.  11,  1921,  p.  182). 


1923.]  INSECTS   OF   THE   PRIBILOF  ISLANDS.  193 

Genus  Hylemyia  Robineau-Desvoidy. 

This  genus  contains  a  large  number  of  very  closely  allied  species, 
although  the  number  recorded  from  North  America  is  comparatively 
small.  I  have  reared  some  of  the  European  species  from  decaying 
vegetation  and  from  manure,  but  several  occur  in  the  larval  stage 
upon  the  roots  of  various  plants,  wild  and  cultivated.  There  is  a 
single  species  in  the  present  collection  which  appears  to  be  new  to 
science. 

Hylemyia  flavisquama,  n.  sp. 

Male. — Black,  very  slightly  shiny.  Head  black;  frontal  stripe 
orange  red,  orbits  blackish  brown,  distinctly  silky ;  facial  orbits  and 
cheeks  brown,  with  silky  surface ;  face  blackish  brown,  surface  with 
brownish  pile.  Thorax  with  faint  pruinescence,  most  distinct  on  2 
narrow  lines  between  the  acrostichals  and  the  dorso-centrals,  and 
on  the  lateral  anterior  angles.  Abdomen  with  grayish  pruinescence ; 
when  viewed  from  behind  there  is  a  distinct  dorso-central  stripe 
visible  which  is  not  disconnected  at  the  abdominal  sutures  and  is 
laterally  extended  anteriorly  in  the  form  of  a  narrow  stripe  along 
the  fore  margin  of  each  segment.  Legs  black.  Wings  slightly 
grayish  or  fuscous.  Squamae  bright  yellow,  fringes  concolorous. 
Halteres  brown  at  base,  knobs  yellow. 

Eyes  distinctly  separated,  the  narrowest  part  of  frons  about  one- 
fifth  the  width  of  either  eye;  above  bases  of  antennae  the  frons  is 
over  one-fourth  the  width  of  head  at  that  point;  orbital  bristles  very 
long  though  fine,  about  6  pairs  present  on  lower  orbits  in  addition 
to  a  number  of  weaker  hairs ;  face  in  profile  protruding  beyond  eyes 
about  as  far  as  the  width  of  third  antennal  joint,  mouth  margin  pro- 
tuberant ;  antenna  of  moderate  length ;  arista  pubescent ;  cheeks  with 
numerous  long  hairs,  the  upper  ones  upcurved,  those  on  vibrissal 
angle  strong,  the  vibrissae  noticeably  strong;  cheeks  broader  than 
third  antennal  joint;  proboscis  rather  long,  stout;  palpi  slender, 
slightly  dilated  apically,  with  a  number  of  weak  surface  hairs. 
Thoracic  hairs  strong;  2  pairs  of  very  strong  presutural  dorso-cen- 
trals; acrostichals  4-rowed  anterior  to  suture.  Abdomen  equal  in 
length  to  thorax,  rather  broad,  the  surface  with  conspicuous  hairs; 
hypopygium  rather  small,  without  conspicuous  lamellae.  Fore  tibia 
with  a  weak  bristle  at  middle  on  postero-ventral  surface  (usually 
with  a  still  weaker  one  below  it),  and  a  short  one  below  middle  on 
antero-dorsal  surface,  preapical  bristle  usually  duplicated;  antero- 
ventral  surface  of  mid-femur  with  a  graduated  series  of  bristles,  the 
longer  ones  at  the  base;  mid-tibia  usually  with  the  following  bris- 
tles: 1  ante ro- ventral,  2  antero-dorsal,  3^i  postero-dorsal,  and  2-3 
postero-ventral ;  hind  femora  with  a  rather  irregular  series  of  strong 
bristles  on  the  antero-dorsal  and  antero-ventral  surfaces;  hind  tibia 


194  NORTH   AMERICAN   FAUNA.  [No.  46 

with  antero-  and  postero-dorsal  surfaces  armed  with  series  of  rather 
strong  bristles;  postero-ventral  surface  with  usually  2  weak  bristles 
near  base,  the  uppermost  just  above  middle.  Costa  with  black, 
setulose  hairs;  costal  spine  small;  inner  cross  vein  just  beyond  apex 
of  first  vein;  outer  cross  vein  almost  straight,  its  upper  extremity 
much  nearer  apex  of  wing  than  its  lower ;  veins  3  and  4  slightly  con- 
vergent apically ;  last  section  of  4  about  one  and  two-thirds  times  the 
length  of  penultimate  section. 

Female. — Differs  from  the  itfale  in  being  less  intensely  black  and 
in  having  the  surface  of  the  body  parts  more  distinctly  pruinose ;  the 
upper  half  of  the  central  stripe  of  frons  is  velvety  black,  merging 
into  the  bright  orange  of  the  anterior  portion.  The  base  of  the  wing 
is  noticeably  yellowish. 

Frons  over  one-third  the  head  width ;  orbits  each  over  half  as  wide 
as  center  stripe;  upper  2  (or  3)  orbitals  directed  slightly  outward, 
lower  4  slightly  inward ;  cruciate  frontal  bristles  strong ;  head  much 
as  in  the  male  in  other  respects  except  that  the  hairs  on  the  cheeks 
are  less  numerous  and  much  stronger,  2  or  3  slightly  upwardly  di- 
rected bristles  being  noticeably  so.  Thorax  with  less  hair  than  in 
male;  acrostichals  irregularly  4-rowed.  Legs  with  a  similar  arma- 
ture to  those  of  male,  the  pair  of  weak  bristles  on  postero-ventral 
surface  of  fore  tibiae  either  absent  or  represented  by  very  weak  hairs. 

Length,  3.75-4.25  mm. 

Type.— St.  George  Island,  June  14,  1914  (lot  19,  G.  D.  Hanna), 
vicinity  of  Garden  Cove. 

Paratypes : 

5  specimens,  June  16,  1914  (lot  35,  G.  D.  Hanna). 
2  specimens,  June  17,  1914  (lot  42,  G.  D.  Hanna). 

2  specimens,  June  25,  1914  (lot  47,  G.  D.  Hanna). 

3  specimens  (including  allotype),  July  4,  1914  (lot  52,  G.  D.  Hanna). 
13  specimens,  July  8,  1914  (lot  55,  G.  D.  Hanna). 

The  data  contains  no  information  as  to  habits,  the  specimens  being 
recorded  as  from  "  toward  "  the  various  rookeries,  and  all  are  from 
St.  George  Island. 

This  species  has  much  the  appearance  of  Anthomyia  radicum 
Linnaeus,  but  differs  in  having  the  upper  scale  of  squamae  larger  than 
the  under  one.  From  P.  badia  Walker  it  differs  in  having  the  acros- 
tichals 4-rowed  instead  of  2-rowed  and  in  having  the  squamae  yellow 
instead  of  whitish. 

Hylemyia  sp. 

A  male  which  is  rather  smaller  than  the  smallest  specimen  of  the 
preceding  species  and  differs  in  having  the  acrostichals  2-rowed, 
represents  a  distinct  species,  but  the  condition  of  the  specimen  pre- 
cludes my  arriving  at  a  definite  identification. 


1923.]  INSECTS   OF   THE  PRIBILOF  ISLANDS.  195 

Locality. — St.  George  Island,  June  17,  1914  (lot  44,  G.  D.  Hanna). 
The  specimen  was  taken  with  other  species  "  around  a  sphagnum 
bog  i  mile  west  of  Village." 

Genus  Fueellia  Robineau-Desvoidy. 

This  genus  has  been  considered  as  belonging  to  the  Anthomyiidae 
by  several  authorities  while  others  have  placed  it  in  the  Cordyluridae. 
However,  it  is  most  properly  placed  in  the  Anthomyiidae  and  is  dis- 
tinguished from  the  genera  in  that  family  which  have  the  eyes  sepa- 
rated in  both  sexes  by  the  presence  of  a  pair  of  cruciate  bristles  on 
the  center  of  the  frontal  stripe. 

The  genus  is  separable  from  Scatophagidae,  as  are  all  Antho- 
myiidae known  to  me  except  4  species,  by  the  area  below  the  protho- 
racic  spiracles,  above  and  in  front  of  the  prothoracic  and  stigmatal 
bristles,  being  bare  instead  of  covered,  at  least  in  part,  with  long  soft 
hairs. 

Stein11  in  1910  revised  the  genus  and  fully  described  the  known 
species ;  Aldrich  in  1918  12  again  revised  the  genus  for  North  Amer- 
ica and  recorded  13  species  as  occurring  in  the  Western  Hemisphere. 
Of  the  described  species  3  occur  in  the  material  before  me.  Two  of 
these  species,  fucorum  and  a?itennata,  are  mentioned  by  Stein  as 
occurring  on  St.  Paul  Island;  the  third  was  originally  described 
from  Greenland.  A  fourth  species  occurring  in  northern  latitudes 
is  pictipennis  Becker,  recorded  from  Hecla  Haven,  East  Greenland. 

The  species  are  invariably  found  on  or  near  the  shore  either  of 
the  sea  or  rivers,  and  generally  are  common.  Their  habits  are  very 
similar  to  those  of  the  Cordyluridae,  the  larvae  being  recorded  as 
feeding  upon  decaying  drift,  though  there  is  no  record  so  far  as  I 
know  of  the  imagines  being  predaceous.  I  have  taken  specimens  of 
the  genus  at  considerable  distances  from  the  sea  on  the  banks  of  vari- 
ous rivers  and  as  most  entomologists  have  the  impression  that  the 
genus  is  exclusively  maritime  in  habit  it  seems  pertinent  to  put  upon 
record  here  the  capture  by  myself  of  a  male  of  maritima  at  Carmi, 
Illinois,  a  town  on  the  Little  Wabash  River,  several  hundreds  of 
miles  from  the  sea  and  nearly  300  miles  from  Lake  Michigan,  the 
nearest  large  area  of  fresh  water  where  it  also  occurs.  I  have  also 
seen  a  male  of  this  species  taken  by  R.  P.  Dow  at  Claremont, 
N.  H.,  which  is  about  90  miles  from  the  sea  in  a  straight  line. 

Fueellia  fucorum  Fallen. 

Scatomyza  fucorum  Fallen,  Scatomyz.,  5,  1819. 

This  species  is  widely  distributed  in  the  Arctic  regions  and  ex- 
tends as  far  south  as  Friday  Harbor.  Stein  records  it  from  St. 
Paul  Island,  and  Meidnaja,  Bering  Straits,  in  addition  to  Friday 

"Wiener  Ent.  Zeit.,  vol.  29,  p.  11,  1910. 

"Proc.  CaJif.  Acad.  Sci.,  4th  ser.,  vol.  S,  p.  157,  1918. 


196  NORTH   AMERICAN   FAUNA.  [No.  46 

Harbor.    There  are  records  of  the  species  from  as  far  south  as  Porto 
Rico  but  it  is  probable  that  most  of  these  refer  to  maritima.    In 
the  present  collection  the  species  is  represented  by  127  specimens  with 
data  as  follows: 
St.  George  Island : 

16  specimens,  May  6,  1914  (G.  D.  Hanna). 

2  specimens,  June  16,  1914  (lot  34,  G.  D.  Hanna). 

5  specimens,  June  4,  1914  (lot  1,  G.  D.  Hanna). 
50  specimens,  June  10,  1914  (lot  9(/G.  D.  Hanna). 

1  specimen,  June  14,  1914  (lot  19,  G.  D.  Hanna). 

1  specimen,  July  4,  1914  (G.  D.  Hanna). 

St.  Paul  Island : 
14  specimens,  August  1,  1914  (E.  A.  Preble). 
44  specimens,  August  19,  1914  (E.  A.  Preble). 

1  specimen,  Summer,  1914  (lot  210,  A.  G.  Whitney). 
10  specimens,  August  16,  1915  (G.  D.  Hanna). 

2  specimens,  August  26,  1916  (G.  D.  Hanna). 

Lot  1  contains  specimens  collected  about  fox  houses  and  on  beach 
among  bowlders ;  lot  9  contains  specimens  that  are  recorded  as  "  very 
common  along  the  beaches,  living  upon  the  decaying  marine  algae  " ; 
lot  19  is  from  the  vicinity  of  Garden  Cove ;  lot  34,  from  "  toward 
East  Rookery";  the  others  have  no  data  other  than  that  already 
given. 

One  specimen  in  the  last  lot  in  the  series  listed  has  2  large  mites 
attached  to  the  posterior  portion  of  thorax  close  to  base  of  posterior 
coxae. 

Fucellia  ariciiformis  Holmgren. 

Scatophaga  ariciiformis  Holmgren,  Kongl.,  Vetenskap.     Akad.       Forhand- 

lingar,  No.  6,  p.  103,  1872. 
Fucellia  ariciiformis  (Holmgren)  Lundbeck,  Vidensk.    Meddel.    Naturhist. 

Foren.  Kjobenhaven,  p.  292,  1900. 

This  species  was  originally  described  from  Greenland  by  Holmgren 
and  afterwards  recorded  from  there  by  Lundbeck.  Stein  indicated 
in  his  revision  of  the  genus  the  characters  that  are  available  for  dis- 
tinguishing the  sexes  from  those  of  fucorum  to  which  it  is  most 
closely  related.  The  male  possesses  the  tuft  of  short  spines  at  base 
of  hind  femora  but  the  tubercle  at  base  of  hind  femora  in  fucorum 
is  absent  in  ariciiformis.  The  female  differs  from  that  of  fucorum 
in  having  only  one  antero-ventral  midtibial  bristle,  and  the  antero- 
ventral  hind  tibial  bristles  very  much  stronger,  more  numerous,  and 
carried  nearly  to  base. 

There  are  4  specimens  of  the  species  in  the  present  collection,  with 
data  as  follows : 

St.  Paul  Island : 
1  female,  May  23,  1914  (lot  154,  A.  G.  Whitney). 
1  female.  2  males,  August  16,  1915  (G.  D.  Hanna). 


1923.]  INSECTS  OF   THE  PRIBILOF  ISLANDS.  197 

A  series  taken  on  St.  Paul  Island,  August  10,  1920  (G.  D.  Hanna), 
has  been  recorded  by  Cole  (Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11,  p. 
172, 1921). 

Fucellia  antennata  Stein. 

Fucellia  antennata  Stein,  Wien.  Ent.  Zeit.,  vol.  29,  p.  23,  1910. 

This  species  was  originally  described  from  Alaska,  the  localities 
given  being  Sitka,  St.  Paul  Island,  and  Karluk.  It  differs  from  all 
other  species  in  the  genus  in  having  the  antennae  elongated,  the  apex 
of  the  third  joint  being  almost  on  a  level  with  the  upper  mouth 
margin.  The  male  is  further  distinguished  by  having  the  antero- 
ventral  surface  of  the  posterior  femora  with  a  series  of  closely  placed 
bristles  extending  from  slightly  before  the  middle  to  their  apices. 
The  species  has  much  the  same  habitus  as  Amaurosoma,  a  genus  of 
predaceous  cordylurids,  but  the  fore  femora  in  that  genus  are  usually 
armed  on  their  antero-ventral  surfaces  with  a  group  of  setulae  and 
the  cruciate  frontal  bristles  are  absent. 

Amongst  the  material  before  me  there  are  29  specimens  of  anten- 
nata as  follows : 

St.  Paul  Island : 

2  specimens,  May  23,  1914  (lot  155,  A.  G.  Whitney). 

9  specimens,  August  1,  1914  (E.  A.  Preble). 

14  specimens,  August  19,  1914  (E.  A.  Preble). 

2  specimens,  Summer,  1914  (E.  A.  Preble). 

1  specimen,  May  16,  1913  (lot  28,  A.  G.  Whitney). 

1  specimen,  August  16,  1915  (G.  D.  Hanna). 

Only  the  first  and  next  to  the  last  lots  have  any  information  re- 
garding the  exact  place  of  capture  attached  to  them.  The  three  speci- 
mens in  Lot  155  were  taken  in  the  Laboratory,  as  was  also  that  in  Lot 
28;  the  latter  is  pinned  with  a  specimen  of  Scatophaga  dasythrix. 
but  the  data  indicates  nothing  other  than  that  they  were  taken  at  the 
same  time. 

Recorded  by  Cole  (Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11,  p.  172, 
November,  1921),  as  a  "  common  species  on  St.  Paul  Island,  August  10 
to  25,  1920"  (G.  D.  Hanna). 

Fucellia  pictipennis  Becker. 

Recorded  by  Cole  (Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11,  p.  172, 
November,  1921)  from  three  specimens  taken  on  St.  Paul  Island, 
June  21  and  August  25, 1920  (G.  D.  Hanna). 

Family  SCATOPHAGIDAE. 

Some  species  of  the  subfamily  Scatophaginae  are  remarkably 
common,  both  on  the  seashore  and  throughout  areas  remote  from  the 
sea,  while  others  are  invariably  confined  to  the  seashore  or  its  imme- 
diate vicinity.     All,  however,  are  very  similar  in  larval  habits,  feed- 


198  NORTH   AMERICAN   FAUNA.  [No.  46 

ing  upon  manure  or  decaying  vegetable  matter.  The  adults  are,  so 
far  as  is  known,  predaceous,  though  they  are  commonly  found  feed- 
ing both  on  flowers  and  fresh  manure.  The  larval  habits  of  the 
species  of  the  subfamily  Cordylurinae  are  not  very  generally  known. 
Some  of  them  feed  upon  decaying  vegetable  matter;  I  have  reared 
one  species  from  river  drift;  and  one  species,  Hydromyza  confluens 
Loew,  feeds  in  the  stems  of  Nymphaea  advena,  forming  gall-like 
swellings  thereon.  So  far  as  I  know,  the  adults  are  predaceous,  some 
of  the  genera  being  particularly  adapted  for  this  mode  of  life,  the 
fore  legs  being  armed  with  strong  spines  to  aid  in  catching  and  re- 
taining prey,  but  even  where  these  spines  are  present  the  insects 
may  also  be  found  feeding  upon  nectar  of  flowers. 

The  material  belonging  to  this  family  contained  in  the  present 
collection  consists  principally  of  Scatophaginae  belonging  to  the 
genus  Scatophaga.  I  have  taken  pains  to  dissect  the  males  of  the 
species  of  Scatophaga  and  figure  certain  parts  with  a  view  to  facili- 
tating their  identification  for  future  students ;  this  has  resulted  also 
in  confirming  their  identity  in  some  cases  with  species  from  more 
temperate  latitudes. 

Subfamily  Cobdylubinae. 

Cordylura  beringensis,  n.  sp. 

Male. — Black,  slightly  shining.  Anterior  portion  of  frons  red- 
dish, merging  into  whitish  on  sides  at  anterior  margin;  face  and 
cheeks  yellowish  white;  frons,  face,  and  cheeks  with  pale  gray  or 
whitish  pruinescence ;  antennae  black,  apex  of  second  joint  brown; 
proboscis  and  palpi  black.  Thorax  and  abdomen  with  distinct, 
brownish  gray  pruinescence.  Legs  black,  tarsi  reddish  on  the  ven- 
tral surfaces.  Wings  clear,  veins  black;  calyptrae  yellowish  white, 
their  fringes  white.     Halteres  pale  brown,  knobs  reddish. 

Frons  slightly  broader  than  eye,  narrowed  anteriorly ;  orbits  usu- 
ally with  6  pairs  of  bristles,  the  upper  2  pairs  directed  outward,  the 
next  pair  directed  forward,  and  the  lower  pairs  incurved;  third 
antennal  joint  of  moderate  length,  rounded  apically;  arista  plumose; 
vibrissae  very  long;  cheeks  very  narrow;  proboscis  stout;  palpi 
slender,  armed  with  a  pair  of  long  hairlike  bristles.  Two  pairs 
of  dorso-central  bristles  on  thorax  anterior  to  suture;  acrostichals 
2-rowed;  mesopleura  with  4-5  moderately  strong  bristles;  sterno- 
pleurum  with  1  very  strong  bristle ;  scutellum  with  4  marginal  bristles. 
Abdomen  short  and  stout;  hypopygium  very  large,  hairy;  fifth 
ventral  segment  ending  in  2  large  rounded  lateral  lobes.  Legs  stout, 
the  fore  and  hind  femora  especially  so;  all  legs  with  long  and 
strong  bristles ;  fore  and  mid  femora  with  long  pale  hairs  on  ventral 
surfaces,  the  latter  with  2-3  strong  antero-ventral  bristles  near 
apex;  hind  femora  with  a  very  long  downward  directed  hair  near 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  199 

base  on  ventral  surface;  hind  tibia  with  2  antero-ventral,  3  antero- 
dorsal  and  3  postero-dorsal  bristles.  Apical  portion  of  first  wing- 
vein  bristly. 

Length,  5  mm. 

Type.— St.  George  Island,  June  16,  1&14  (lot  35,  G.  D.  Hanna). 
The  type  and  2  paratypes  were  taken  "  toward  East  Rookery  from 
Village."     Other  paratypes  as  follows: 

3  specimens,  June  17,  1914,  uplands  near   Staraya  Artel    (lot  42,   G.   D. 

Hanna). 
1  specimen,  June  17,  1914,  near  a  sphagnum  bog,  i  mile  west  of  Village 

(lot  44,   G.  D.  Hanna). 
1  specimen,   June  25,   1914,   from   toward  North   Rookery    (lot  47,    G.   D. 

Hanna). 

There  is  a  Colorado  species  which  very  closely  resembles  the  above. 
It  differs  in  having  the  legs  with  weaker  bristles,  the  wings  clearer, 
the  inner  cross  vein  more  distinctly  beyond  middle  of  discal  cell,  and 
the  first  vein  almost  bare. 

This  species  has  a  decided  affinity  to  proboscidea  Zetterstedt.  a 
species  that  has  a  northern  range  in  Europe.  It  differs,  however,  in 
being  smaller  and  in  having  the  abdomen  much  less  conspicuously 
hairy,  as  well  as  in  several  other  minor  characters. 

Genus  Allomyella  n.  n.u 

Generic  characters. — Head  about  as  high  as  long,  face  slightly 
retreating  below,  cheek  of  moderate  width,  eye  higher  than  long, 
vibrissa  weak,  below  it  one  bristle;  antennae  of  moderate  length, 
third  joint  subangulate  at  apex  on  upper  side,  arista  bare,  occiput 
with  bristles  along  upper  half  on  eye-margin,  and  below  these 
another  series  or  irregular  group  on  back  of  head.  Dorso-centrals 
5,  the  other  bristles  as  in  Cordylura;  1  sterno-pleural ;  pteropleura 
with  a  few  hairs.  Abdomen  at  least  as  long  as  wings,  broad,  apical 
segment  short,  compressed,  but  little  protruded.  Fore  tibia  with 
short  black  setulae  on  ventral  surface.     First  wing  vein  bare. 

Allomyella  brevipennis,  n.  sp. 

Female. — Black,  abdomen  distinctly  shiny.  Head  black;  frons 
opaque ;  central  stripe  brown,  paler  anteriorly ;  lower  part  of  face 
and  anterior  angles  of  cheeks  yellow ;  antennae  black,  second  joint 
slightly  reddish  apically ;  arista  black ;  proboscis  glossy  black ;  palpi 
brown  at  base,  yellow  apically.  Legs  black,  tibiae  and  tarsi  testa- 
ceous yellow,  the  apices  of  the  latter  slightly  darkened.  Wings 
slightly  brownish,  the  veins  thick  and  slightly  darkened,  faintly 
infuscated    along   their   margins.     Calyptrae    brown,    fringes    yel- 

13  The  generic  name  Allomyia  given  by  me  to  tbis  genus  in  my  paper  on  the  Canadian 
Arctic  Diptera  is  preoccupied  by  Allomyia  Felt  and  the  name  of  my  genus  accordingly  is 
changed  herein. 


200  NOETH  AMERICAN  FAUNA.  [No.  46 

lowish.  Halteres  reddish  yellow.  Hairs  and  bristles  yellow,  the 
stronger  bristles  on  head,  legs,  and  abdomen  blackish,  but  paler  in 
transmitted  light. 

Frons  distinctly  broader  than  eye,  very  slightly  narrowed  ante- 
riorly ;  orbits  each  with  5  bristles,  the  2  upper  ones  directed  outward ; 
arista  bare ;  profile  as  in  Plate  XV,  Figure  28.  Thorax  and  abdomen 
with  rather  numerous  surface  hairs;  pleural  bristles  not  well  distin- 
guished from  the  other  hairs,  the  prothoracic  and  stigmatal  bristles 
weak.  Last  abdominal  segment  distinctly  elongated,  its  lateral  mar- 
gins with  a  number  of  long  bristles.  Legs  rather  stout ;  fore  femora 
distinctly  incrassated ;  all  femora  with  rather  widely  separated  hair- 
like central  bristles ;  fore  tibia  with  4  bristles,  2  on  the  dorsal  surface 
(one  at  middle  and  one  near  tip),  and  2  slightly  below  them  trans- 
versely on  posterior  surface ;  hind  tibia  with  the  following  bristles :  2 
on  postero-dorsal  surface,  one  at  middle  and  the  other  at  one-fourth 
from  base;  2  on  antero-dorsal  surface,  the  upper  in  transverse  line 
with  the  upper  on  postero-dorsal  surface,  the  lower  slightly  below 
middle;  2 — 3  on  antero-ventral  surface  near  apex.  Wings  not  ex- 
tending to  apex  of  abdomen ;  third  and  fourth  veins  divergent,  the 
former  ending  in  apex  of  wing. 

Length,  5.25  mm. 

Type. — St.  George  Island,  June  8,  1914  (lot  8,  G.  D.  Hanna). 

Subfamily  Hydromyzinae. 
Genus  Pogotwta  Becker. 

Pogonota  kincaidi  Coquillett.  (PI.  XII,  fig.2.) 

Pogonota  kincaidi  Coquillett,  Proc.  Washington  Acad.  Sci.,  vol.  2,  p.  455, 
1900. 
This  species  is  represented  by  seven  specimens,  with  data  as  fol- 
lows: 2  males  and  1  female,  St.  George  Island,  June  16,  1914  (G.  D. 
Hanna,  Lot  36)  ;  1  male  and  3  females,  August  16,  1915  (G.  D. 
Hanna).  The  original  description  was  made  from  a  male  and  female 
obtained  by  Prof.  T.  Kincaid  on  Popof  Island,  Alaska,  and  there  are 
no  subsequent  records  of  the  species. 

Subfamily  Scatophaginae. 

Genus  Scatophaga  Meigen. 

There  is  not  a  published  synopsis  of  the  North  American  species 
of  this  genus,  and  in  presenting  one  covering  the  species  contained 
in  the  present  collection  I  hope  to  accomplish  two  objects — to  make 
it  possible  for  future  students  to  recognize  the  forms  recorded,  and 
to  indicate  that,  similar  though  the  species  appear,  there  are  just 
as  good  characters  available  for  their  separation  as  there  are  in 
most  of  the  dipterous  families  and  even  better  than  there  are  in  some. 


1923.]  INSECTS   OF   THE   PRIBILOF  ISLAISTDS.  201 

Becker  in  1894 14  published  the  most  complete  study  of  the  family 
that  has  been  undertaken  up  to  the  present.  In  the  genus  Scatophaga 
the  synoptic  key  is  very  full  and  quite  satisfactory;  but  few  of  the 
species  are  described  fully  in  the  text,  the  author  confining  himself 
to  notes  upon  the  species  and  to  indicating  their  synonymy  except  in 
the  case  of  new  species.  So  far  as  I  know,  the  present  paper  is 
the  first  in  which  use  has  been  made  of  the  characters  of  the  fifth 
ventral  segment  of  the  abdomen  of  the  males  except  that  by  the 
writer  on  the  Diptera  of  the  Canadian  Arctic  Expedition.  The  dif- 
ferences in  structure  in  this  segment  are  very  marked  in  some  of 
the  species,  as  can  be  seen  from  an  examination  of  the  figures. 
There  are  four  distinct  types  of  structure  of  this  segment  in  the 
species  I  have  examined.  The  simplest  form  is  represented  by  1 
species,  crinita,  and  has  on  each  lateral  angle  of  the  fifth  sternite  a 
small  rounded  prolongation ;  in  3  species  there  is  a  very  long  prolonga- 
tion of  this  segment  on  each  side  of  the  median  line;  furcata 
has  a  small,  rather  knoblike  process  on  each  side  of  the  median  line 
on  posterior  margin  of  the  segment;  and  stercoraria  has  a  similar 
pair  to  those  of  furcata  in  addition  to  the  lateral  elongations,  pre- 
senting the  most  elaborate  structure  of  any  species  known  to  me. 
Stercoraria  (PI.  XIV,  fig.  IT)  has  been  recorded  from  Alaska,  but  is 
unrepresented  in  the  present  collection,  the  figure  being  introduced 
here  to  show  the  difference  in  form  of  the  segment.  It  will  be  ob- 
served that  there  is  a  distinct  similarity  in  the  male  hypopygia  of 
Scatophaga  and  Coelopa  (PL  XIV,  figs.  23,  24,  and  25).  I  published 
figures  of  the  fifth  sternites  of  the  males  of  S.  suilla  and  S.  lutaria 
in  my  report  on  the  Diptera  collected  by  the  Canadian  Arctic  Ex- 
pedition, 1919. 

Key  to  species  of  Scatophaga. 

1.  Males    2. 

Females  r 6. 

2.  Mid  and  hind  tibiae  with  a  number  of  strong,  outstanding  bristles  in  addi- 

tion to  the  soft  hairs 3. 

Mid  and  hind  tibiae  with  remarkably  long  soft  hairs,  and  without  strong 
bristles  except  at  apex  and  in  crinita  one  bristle  near  apex  on  posterior 
surface 5. 

3.  Cross  veins  of  the  wings  not  infuscated ;  legs  black,  very  densely  brownish 

pruinose,  bases  of  tibiae   indistinctly   reddish ;   fifth   ventral   segment   of 

abdomen  as  in  Plate  XIV,  Figure  21 islandica. 

Cross  veins  of  the  wings  conspicuously  infuscated ;  legs  either  almost  en- 
tirely yellow,  or  reddish  with  black  femora 4. 

4.  Legs  reddish  or  vinous  colored,  shining,  femora  black;  hairs  on  hind  tibiae 

woolly,  nearly  as-  long  as  the  bristles,  the  latter  rather  slender  ;  fifth  ventral 
segment  of  abdomen  with  a  long,  thornlike  projection  on  each  side  of  the 
median  line  (PI.  XIV,  fig.  20) rubicunda. 

"Berl.  Ent.  Zeitschr.,  vol.  39,  pp.  77-196. 


202  NORTH   AMERICAN   FAUNA.  [No.  46 

Legs  pale  testaceous  or  yellowish,  only  the  fore  femora  in  part  black ;  hairs 
on  hind  tibiae  setulose,  very  much  shorter  than  the  bristles,  the  latter 
very  stout,  fifth  ventral  segment  of  abdomen  with  short  knoblike  process 
on  each  side  of  the  median  line  (PI,  XIV,  fig.  22) furcata. 

5.  Legs  sinning  black ;  abdominal  hairs  yellow,  varying  from  whitish  to  orange ; 

mid  tibiae  with  a  bristle  near  apex  on  the  posterior  surface;  fifth  ventral 

abdominal  segment  as  in  Plate  XIV,  Figure  19 crinita. 

Legs  dull  black ;  abdominal  hairs  black,  occasionally  a  few  of  those  on  the 
ventral  surface  are  yellowish;  mid  tibiae  without  bristle  near  apex;  fifth 
ventral  abdominal  segment  as  in/Plate  XIV,  Figure  18 dasythrix. 

6.  Pteropleura  bare 7. 

Pteropleura  hairy 9. 

7.  Cross  veins  of  the  wings  not  infuscated ;  hairs  on  dorsum  of  abdomen  black, 

contrasting  sharply  with  those  of  venter  and  lower  part  of  pleura,  which 

are  pale  yellow  and  silky ;  legs  black crinita. 

Cross  veins  of  wings  conspicuously  infuscated ;  legs  in  great  part  yellow  or 
reddish - 8. 

8.  Large  species,  averaging  10  mm.  in  length ;  legs  vinous  in  color,  femora  in 

large  part  black ;  all  hairs  on  abdomen  fuscous ;  apical  spur  on  hind  tibia 

very  strong  and  much  curved rubicunda. 

Smaller  species,  averaging  7-8  mm.  in  length ;  legs  yellow,  fore  femora  more 
or  less  blackened  above ;  hairs  on  dorsum  of  abdomen  blackish  and  setulose, 
those  on  venter  yellow  and  soft ;  apical  spur  on  hind  tibia  of  moderate 
strength  and  almost  straight furcata. 

9.  Frons  conspicuously  reddish  in  front ;  tibiae  reddish  or  brownish ;  hairs  on 

mesonotum  not  numerous,  setulose,  those  on  hind  tibia  very  much  shorter 
than  the  intermixed  bristles,  and  setulose;  apex  of  abdomen  with  normal 

hairing islandica. 

Frons  not  noticeably  reddish  in  front ;  tibiae  black ;  hairs  on  mesonotum 
numerous,  long  and  rather  soft,  those  on  hind  tibiae  soft  and  slightly  curled, 
some  of  them  as  long  as  the  intermixed  bristles ;  apex  of  abdomen  with  a 
noticeable  tuft  of  soft  curled  black  hairs dasythrix. 

Scatophaga  islandica  Becker.  PI.  XIV,  fig.  21. 

Scatophaga  islandica  Becker,  Berl.  Ent.  Zeitschr.,  vol.  39,  p.  175,  1894. 

Apical  ventral  segment  of  male  as  Plate  14,  Figure  21. 

This  species  was  originally  described  from  specimens  in  the  Loew 
collection  from  Iceland  and  Labrador.  Coquillett  subsequently  re- 
corded it  from  Commander  Islands  and  Alaska. 

There  are  9  specimens  in  the  present  collection  from  St.  Paul 
Island  with  data  as  follows: 

4  specimens,  August  19,  1914   (E.  A.  Preble). 
1  specimen,  August  1,  1914  (E.  A.  Preble). 
4  specimens,  August  16,  1915  (G.  D.  Hanna). 

Recorded  by  Cole  (Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11,  p.  172, 
Nov.,  1921)  from  St.  Paul  Island,  June  21  and  August  10, 1920  (G.  D. 
Hanna). 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  203 

Scatophaga  rubicunda  Malloch.  PI.  XIV,  fig.  20. 

Scatophaga  rubicunda  Malloch,  J.  R.,  Can.  Arctic  Exp.,  1913-1918,  vol.  3, 
p.  81c  (1919). 

Male. — Opaque  black-brown.  Head  concolorous  with  thorax; 
frontal  stripe  orange  red,  orbits  on  lower  portions  grayish ;  face  and 
cheeks  reddish  orange,  their  surfaces  covered  with  grayish  yellow 
pruinescence,  antennae  brownish  black,  apex  of  second  joint  on  inner 
side  reddish ;  proboscis  glossy  black ;  palpi  reddish  or  yellowish.  Disc 
of  mesonotum  with  indications  of  6  vittae,  a  median  confluent  pair 
on  acrostichal  area,  a  narrow  line  along  the  region  of  the  dorso-cen- 
tral  bristles  and  a  broader  irregular  streak  laterad  of  the  latter. 
Hypopygium  and  apex  of  last  segment  reddish.  Legs  vinous  colored, 
the  femora  except  their  apices  darkened.  Wings  slightly  grayish, 
with  a  yellow  tinge  along  the  costal  region,  the  cross  veins  distinctly 
infuscated,  veins  reddish  brown.  Halteres  reddish.  Bristles  black, 
the  soft  hairs  fuscous. 

Arista  bare,  swollen  at  base,  apical  part  very  slender;  cheek  half 
as  high  as  eye,  anterior  angle  with  a  group  of  8  to  10  long,  black 
hairs;  palpi  elongate,  slightly  leaflike.  Thorax  with  long  soft 
hairs,  acrostichal  bristles  not  distinguishable  from  the  discal  hairs, 
the  dorso-central  bristles  barely  so ;  pteropleura  bare ;  scutellum  with 
4  strong  marginal  bristles  and  numerous  long  hairs.  Abdominal 
hairs  not  so  long  as  in  crinita,  more  dense  and  not  "  crinkly  " ;  fifth 
ventral  plate  as  Plate  XIV,  Figure  20.  Legs  with  long  hairs;  mid- 
tibiae  with  7  long  bristles  distinguishable  from  the  hairs,  in  addi- 
tion to  the  apical  spines,  situated  as  follows:  2  on  the  antero- 
dorsal  surface,  one  above  the  other,  the  upper  one  just  below  mid- 
dle; 3  on  the  postero-dorsal  surface,  the  upper  near  base;  and  2  on 
posterior  surface,  in  transverse  line  with  those  on  the  antero-dorsal 
surface;  hind  tibiae  each  with  6  bristles  in  addition  to  the  hairs 
and  apical  spurs  as  follows :  3  strong  ones  on  antero-dorsal  sur- 
face, the  upper  one  just  above  middle  and  the  lower  one  close  to  tip; 
and  3  long  hairlike  ones  on  postero-dorsal  surface,  the  upper  one 
being  about  one-fourth  from  base  of  tibiae,  the  next  close  to  middle, 
and  the  lowest  one  very  near  to  apex ;  apical  spurs  bent. 

Female. — Similar  to  the  male  in  color.  Differs  from  the  male  in 
having  the  hairs  throughout  shorter,  stronger,  and  less  numerous 
and  the  bristles  stronger.    Length,  9-10  mm. 

Type.— St.  George  Island,  June  16,  1914  (lot  34,  G.  D.  Hanna). 
Paratype,  St.  George  Island,  1  specimen,  same  data  as  type.  Allo- 
type, St.  George  Island,  one  of  2  specimens,  June  4,  1914  (lot  1, 
G.  D.  Hanna)  ;  and  2  specimens  June  8,  1914  (lot  8,  G.  D.  Hanna). 
Lot  34  was  collected  toward  East  Rookery  from  village;  lot  1  about 
fox  houses  and  on  beach  among  boulders  along  with  S.  dasythrix; 


204  NOBTH   AMERICAN   FAUNA.  [No.  46 

and  lot  8,  near  village,  from  flowers.  Recorded  by  Cole  (Proc. 
Calif.  Acad.  Sci.,  4th  Ser.,  vol.  11,  Nov.  1921,  p.  173)  from  St.  George 
Island,  July  1  (G.  D.  Hanna). 

This  species  differs  from  nubifera  Coquillett  in  being  larger  and 
in  the  chaetotaxy  of  the  mid  and  hind  tibiae.  Sometimes  the  femora 
are  entirely  reddish,  and  very  rarely  there  are  two  or  three  long 
hairs  present  on  center  of  pteropleura. 

Scatophagy  furcata  Say.  /  PI.  XIV,  figs.  22  and  23. 

Pyropa  furcata,  Say,  Jour.  Acad.  Sci.  Phila.,  vol.  3,  p.  98,  1823. 
Sc'atophaga  squalida  Meigen,  Syst.  Beschr.  Zwiefl.  Ins.,  vol.  5,  p.  252,  1826. 
Scatophaga  nigricans  (Macquart)  Meigen.  1.  c.  vol.  7,  p.  342,  1838. 
Cordylura  fvscipennis  Zetterstedt,  Ins.  Lapp.,  p.  733,  1840. 
Cordylura  fuscinervis  Zetterstedt.  Dipt.  Scand.,  vol.  5,  pp.  1973,  1975 ;  1845. 
Scatophaga  apicalis  Curtis,  Appendix  to  Narrative  of  2d  Voyage  in  Search 

of  Northwest  Passage,  p.  76,  1835. 
Cleigastra  suisterci  Townsend,  Can.  Ent.,  vol.  23,  p.  153,  1891. 

The  fifth  ventral  segment  of  the  male  and  the  hypopygium  are 
shown  in  Plate  14,  figures  22  and  23. 

This  species  was  first  described  by  Say  from  specimens  obtained 
in  Missouri.  It  occurs  throughout  North  America,  extending  its 
range  well  into  the  Arctic  portions,  and  is  quite  as  common  in  Eu- 
rope as  in  America.  There  is  very  considerable  variation  in  size 
and  color  in  this  species,  which  has  probably  caused  some  confusion 
on  the  part  of  different  authors.  I  have  dissected  many  examples 
of  different  sizes  and  of  varied  shades  of  color  in  an  attempt  to  find 
structural  differences  but  have  failed  to  find  any  that  would  jus- 
tify me  in  separating  even  the  most  extreme  forms.  I  know  that 
the  nature  of  the  pabulum  of  the  larvae  affects  the  appearance  of 
the  resultant  imagines,  those  that  have  had  an  abundance  of  nutri- 
tious food  being  large,  brightly  colored,  and  very  hairy,  while  those 
that  have  had  a  supply  of  rather  dry  and  poor  food  are  smaller, 
darker,  and  less  hairy. 

The  normal  food  of  the  larvae  of  this  species  is  manure. 

In  the  present  collection  there  are  83  specimens  with  data  as  fol- 
lows: 

St.  George  Island. 

1  specimen,  April  24,  1914  (G.  D.  Hanna). 
1  specimen,  May  6,  1914  (G.  D.  Hanna). 
1  specimen,  June  16,  1914  (lot  35,  G.  D.  Hanna). 
1  specimen,  June  17,  1914  (lot  42,  G.  D.  Hanna). 
4  specimens,  June  24,  1914  (lot  46,  G.  D.  Hanna). 
4  specimens,  June  25,  1914  (lot  47,  G.  D.  Hanna). 
14  specimens,  June  27,  1914  (lot  49,  G.  D.  Hanna). 
1  specimen,  July  4.  1914  (lot  52,  G.  D.  Hanna). 
1  specimen,  July  8,  1914  (lot  55,  G.  D.  Hanna). 


1923.]  INSECTS   OF   THE   PRIBIL-OF   ISLANDS.  205 

St.  Paul  Island. 

11  specimens,  August  16.  1914  (G.  D.  Hanna). 
15  specimens,  August  1,  1914   (E.  A.  Preble). 
35  specimens,  August  19,  1914  (E.  A.  Preble). 

3  specimens,  Summer,  1914  (E.  A.  Preble). 
14  specimens,  1916  (G.  D.  Hanna). 

Only  lot  55  is  mentioned  definitely  as  having  been  taken  on  the 
shore;  the  others  are  recorded  as  from  "toward  East  Rookery"; 
"uplands  near  Staraya  Artel";  "toward  North  Rookery,"  and  "to- 
ward Zapadni." 

Recorded  by  Cole  (Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11,  Nov. 
1921,  p.  173)  from  St.  Paul  Island,  June  21  to  August  10,  1920,  and 
St.  George  Island,  June  30  (G.  D.  Hanna). 

Scatophagy  crinita  Coquillett.  PI.  XIV,  fig.  19. 

Scatophaga  crinita,  Coquillett,  Proc.  U.  S.  Nat.  Mus.,  vol.  23,  p.  612,  1901. 

The  fifth  ventral  abdominal  segment  of  the  male  is  as  in  Plate  14, 
Figure  19. 

This  species  was  originally  described  from  specimens  obtained  on 
Bering  Island.  The  series  of  specimens  in  the  present  collection 
shows  a  very  considerable  variation  both  in  the  size  of  the  different 
specimens  and  in  the  color  of  the  hairs  on  the  body,  the  latter  vary- 
ing from  yellowish  white  to  deep  orange.  As  indicated  under  the 
previous  species  there  very  probably  is  some  connection  between  these 
conditions  and  the  nature  of  the  larval  pabulum. 

There  are  89  specimens  in  the  collection  with  data  as  follows: 

St.  Paul  Island. 

1  specimen,  May  23,  1914  (lot  154,  A.  G.  Whitney). 
3  specimens,  June  11,  1913  (lot  44,  A.  G.  Whitney). 
1  specimen,  July  3,  1913  (lot  60,  A.  G.  Whitney). 
1  specimen,  July  22,  1913  (lot  94,  A.  G.  Whitney). 
1  specimen,  Summer,  1914  (E.  A.  Preble). 

1  specimen,  August  16,  1915  (G.  D.  Hanna). 

5  specimens,  June- Aug.  1916  (G.  D.  Hanna). 

St.  George  Island. 

2  specimens,  Sept.  3,  1913  (G.  D.  Hanna). 
2  specimens,  Sept.  6,  1913  (G.  D.  Hanna). 

2  specimens,  May  6,  1914  (G.  D.  Hanna). 

3  specimens,  June  4,  1914  (lot  1,  G.  D.  Hanna). 
1  specimen,    June  8,  1914  (lot  8,  G.  D.  Hanna). 
1  specimen,    June  10,  1914  (lot  9,  G.  D.  Hanna). 

6  specimens,  June  14,  1914  (lot  18,  G.  D.  Hanna). 
1  specimen,    June  14,  1914  (lot  19,  G.  D.  Hanna). 

16  specimens,  June  16,  1914  (lot  34,  G.  D.  Hanna). 
9  specimens,  June  16,  1914  (lot  26,  G.  D.  Hanna). 
1  specimen,    June  16,  1914  (no  number,  G.  D.  Hanna). 
10S7310— 23 14 


206  NORTH   AMERICAN   FAUNA.    .  [No.  46 

7  specimens,  June  17,  1914  (lot  38,  G.  D.  Hanna). 

4  specimens,  June  24,  1914  (lot  46,  G.  D.  Hanna). 
10  specimens,  June  27,  1914  (lot  49,  G.  D.  Hanna). 
15  specimens,  July  4,  1914  (lot  52,  G.  D.  Hanna). 

1  specimen,    July  8,  1914  (lot  55,  G.  D.  Hanna). 

The  specimen  of  lot  60  was  taken  on  Otter  Island  at  the  same 
time  as  the  specimen  of  S.  dasythrix  bearing  the  same  number.  Lots 
44  and  94  were  taken  on  peninsulas  on  St.  Paul  Island.  The  speci- 
men bearing  the  Hanna  lot  number  8  appears  to  have  been  taken 
on  flowers  (see  also  under  S.  Aibicunda).  Judging  from  the  data 
pertaining  to  the  collection  this  species  is  found  both  along  the 
shore  and  on  the  uplands,  many  examples  being  obtained  towards 
East  Rookery,  at  Garden  Cove,  and  towards  Zapadni. 

Scatophaga  dasythrix  Becker.  PI.  XIV,  tig.  18. 

Scatophaga  dasythrix  Becker,  Berl.  Ent.  Zeitschr.,  vol.  39,  p.  173,  1894. 

The  fifth  ventral  abominal  segment  of  the  male  is  shown  in  Plate 
14,  Figure  18. 

This  species  was  originally  described  from  specimens  in  the  col- 
lections of  Loew  and  Schnabl,  obtained  from  Bering  Straits.  Sub- 
sequently Coquillett  recorded  its  occurrence  on  Bering  Island. 

In  the  present  collection  there  are  108  specimens  with  data  as 
follows : 

St.  George  Island. 

1  specimen,  August  16,  1913  (G.  D.  Hanna). 

2  specimens,  May  17,  1914  (G.  D.  Hanna). 

62  specimens,  June  4,  1914  (lot  1,  G.  D.  Hanna). 
1  specimen,  June  10, 1914  (lot  9,  G.  D.  Hanna). 

10  specimens,  June  16,  1914  (lot  34,  G.  D.  Hanna). 
1  specimen,  July  4,  1914  (lot  52,  G.  D.  Hanna). 

St.  Paul  Island. 

1  specimen,  April  13,  1913  (lot  7,  A.  G.  Whitney). 

1  specimen,  May  16,  1913  (lot  28,  A.  G.  Whitney). 

2  specimens,  June  24,  1913  (lot  57,  A.  G.  Whitney). 
1  specimen,  July  6,  1913  (lot  63,  A.  G.  Whitney). 

1  specimen,  July  6,  1913  (lot  64,  A.  G.  Whitney). 

1  specimen,  July  6,  1913  (lot  66,  A.  G.  Whitney). 

2  specimens,  July  18,  1913  (lot  80,  A.  G.  Whitney). 

2  specimens,  July  22,  1913  (lot  94,  A.  G.  Whitney). 

3  specimens,  summer,  1914  (E.  A.  Preble). 
2  specimens,  August  19, 1914  (E.  A.  Preble). 

1  specimen,  August  1,  1914  (E.  A.  Preble). 

2  specimens,  August  16,  1915  (G.  D.  Hanna). 

11  specimens,  July- August,  1916  (G.  D.  Hanna). 

Otter  Island  (6  miles  from  St.  Paul  Island). 

1  specimen,  July  3,  1914  (lot  60,  A.  G.  Whitney). 
Sealion  Rock,  or  "  Sivutch  "  Island. 

10  specimens,  June  29,  1914  (lot  182,  A.  G.  Whitney). 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  207 

This  species  appears  from  the  number  of  examples  and  the  recorded 
data  to  be  the  most  common  of  those  in  the  collection.  Several  exam- 
ples were  taken  in  copula  and  three  with  prey.  The  prey  represents 
3  species :  Fucellia  antennata,  Leria  fraterna,  and  Leria  leucostoma. 
The  specimens  in  lot  182  were  taken  on  the  grassy  summit  of  Sea- 
lion  Rock  at  a  height  of  60  feet. 


D.  W.  Coquillett  records  from  the  Pribilofs  (in  Schwarz,  E.  A.,  In  The  Fur  Seals 
and  Fur-Seal  Islands  of  the  North  Pacific  Ocean,  Part  3,  pp.  550-552,  1899)  one 
species  in  addition  to  those  listed  in  the  present  paper,  which,  however,  may  be  only 
nominally  distinct :  Scatophaga  diadema  Wiedemann. — W.  L.  M. 

Family  HELOMYZIDAE. 

The  species  of  Helomyzidae  are  usually  found  upon  carrion  or  de- 
caying vegetable  matter,  though  certain  species  of  the  genus  Leria 
have  been  recorded  as  frequenting  caves  where  they  feed  upon  fungi 
and  droppings  of  bats.  The  species  recorded  as  frequenting  caves 
are  not  specialized  in  any  manner  that  adapts  them  to  this  habitat 
and  are  found  in  other  situations  also.  A  species  of  the  genus 
Heteromyza  that  has  been  found  in  caves,  I  have  found  on  carrion 
in  an  open  field  in  Scotland.  The  members  of  the  genus  Eccoptomera 
are  found  in  the  larval  and  pupal  stages  in  underground  nests  of 
moles,  and  I  have  found  the  imagines  there  also,  but  I  have  sometimes 
taken  the  latter  by  sweeping  the  undergrowth  in  Scotch  woods. 

The  family  is  not  well  represented  by  species  in  the  material  be- 
fore me  but  there  are  a  large  number  of  specimens. 

A  single  male  of  each  species  in  the  collection  was  taken  as  the 
prey  of  Scatophaga  dasythrix.  These  are  recorded  under  that 
species. 

Aldrich  and  Darlington  have  published  an  extensive  paper  on  the 
family. 15 

Genus  Leria  Robineau-Desvoidy. 

Leria  fraterna  Loew. 

Scoliocentra  fraterna  Loew,  Bert.  Ent.  Zeitschr.,  p.  27,  1863. 

Leria  fraterna   (Loew),  Coquillet,  Proc.  Wash.  Acad.  Sci.,  vol.  2,  p.  457, 

1900. 
Leria  fraterna  Aldrich  and  Darlington,  Trans.  Amer.   Ent.  Soc,  vol.  34, 

p.   79,   1908. 

This  species,  which  was  originally  described  from  Sitka,  Alaska, 
by  Loew,  is  represented  by  147  specimens  in  the  present  collection 
from  the  following  localities: 

St.  George  Island. 

31  specimens,  April-May,  1914  (G.  D.  Hanna). 
23  specimens,  June  4,  1914  (lot  1,  G.  D.  Hanna). 
6  specimens,  June  8,  1914  (lot  8,  G.  D.  Hanna). 

»B  Trans.  Amer.  Ent.  Soc,  vol.  34,  1908. 


208  NORTH  AMERICAN   FAUNA.  [No.  46 

18  specimens,  June  16,  1914  (lot  34,  G.  D.  Hanna). 
1  specimen,  June  16,  1914  (lot  35,  G.  D.  Hanna). 
3  specimens,  June  14,  1914  (lot  19,  G.  D.  Hanna). 

1  specimen,  June  27,  1914  (lot  49,  G.  D.  Hanna). 

2  specimens,  July  4,  1914  (lot  52,  G.  D.  Hanna). 
6  specimens,  June  24  ,1914  (lot  46,  G.  D.  Hanna). 
1  specimen,  June  14,  1914  (lot  23,  G.  D.  Hanna). 

3  specimens,  June  17,  1914  (lot  42,  G.  D.  Hanna). 

1  specimen,  June  10,  1914  (lot  9,  G.  D.  Hanna). 

St.  Paul  Island.  / 

5  specimens,  May  17,  1912  (M.  C.  Marsh). 

13  specimens,  May  16,  1913  (lot  26,  A.  G.  Whitney). 

2  specimens,  May  16,  1913  (lot  24,  A.  G.  Whitney). 

6  specimens,  May  19,  1913  (lot  132,  A.  G.  Whitney). 
8  specimens,  May  23,  1913  (lot  156,  A.  G.  Whitney). 
1  specimen,  May  23,  1913  (lot  154,  A.  G.  Whitney). 

1  specimen,  July  11,  1913  (lot  70,  A.  G.  Whitney). 

2  specimens,  Aug.  1,  1914  (E.  A.  Preble). 

6  specimens,  summer,  1914  (E.  A.  Preble). 

1  specimen  (see  under  Scatophaga  dasythrix,  lot  64,  A.  G.  W.). 

The  collections  were  made  in  various  portions  of  the  islands,  some 
of  them  about  fox  houses  and  on  beaches,  while  others  were  made 
amongst  grass  and  herbage.  No  indication  is  given  as  to  more  exact 
habitats. 

Aldrich  and  Darlington  give  the  following  localities  for  this 
species : 

Moscow,  Idaho. 
St.  Anthony  Park,  Minn. 
Montreal,  Canada. 
Ungava  Bay,  Labrador. 
Hudson  Bay  Territory. 

It  has  also  been  recorded  by  Coquillett  as  occurring  in  Alaska, 
British  Columbia,  and  on  Mount  Washington,  N.  H.  A  female  speci- 
men in  the  collection  of  the  Illinois  State  Laboratory  of  Natural 
History  does  not  differ  materially  from  the  specimens  before  me ;  the 
data  on  this  specimen  is  Algonquin,  111.,  March  21,  1894. 

Leria  leucostoma  Loew. 

Ble,pha?-optera  leucostoma  Loew,  Berl.  Ent.  Zeitschr.,  p.  28,  1863. 

This  species,  which  also  was  originally  described  from  Alaska  by 
Loew,  is  represented  by  18  specimens,  as  follows : 
St.  George  Island. 

2  specimens,   September  2,  1913    (G.  D.  Hanna). 

2  specimens,  April  12,  1914    (G.  D.  Hanna). 

1  specimen,  June  16,  1914  (lot  35,  G.  D.  Hanna). 

2  specimens,  June  8,  1914  (lot  8,  G.  D.  Hanna). 
5  specimens,  July  4,  1914  (lot  52,  G.  D.  Hanna). 
1  specimen,  June  27,  1914  (lot  49,  G.  D.  Hanna). 


1923.]  INSECTS    OF    THE   PRIBILOF   ISLANDS.  209 

St.   Paul  Island. 

August   19,   1915    (G.    D.   Hanna). 
June  29,  1914   (lot  182,  A.  G.  Whitney). 
July  6,  1914    (lot  66,  A.  G.  Whitney). 
July  27,   1914    (E.   A.    Preble). 
August  1  and  19,  1914    (E.  A.  Preble). 
(See  under  Scatophaga  dasythrix,  lot  63,  A.  G.  W.) 

Aldrich  and  Darlington  record  this  species  from  Hampton,  N.  H. ; 
White  Mountains,  N.  H. ;  and  Mount  Constitution,  Wash. ;  Coquillett 
has  recorded  it  from  Alaska  and  White  Mountains,  N.  H. 

Leria  pectlnata  Loew  has  been  recorded  by  Coquillett  from  the 
Pribilofs  (in  Schwarz,  E.  A.,  in  The  Fur  Seals  and  Fur  Seal  Islands 
of  the  North  Pacific  Ocean,  pt.  3,  1899,  pp.  550-552). 

Leria  crassipes  Loew  has  been  recorded  by  Cole  (Proc.  Calif. 
Acad.  Sci.,  4th  ser.,  vol.  11,  p.  173,  Nov.,  1921),  from  St.  Paul  Island, 
July  4  and  August  10  (G.  D.  Hanna).  However,  it  is  doubtful  if 
this  species  occurs  in  North  America;  the  specimens  recorded  are 
almost  without  doubt  leucostoma  Loew. 

Leria  sp. 

A  female  recorded  by  Cole  (Proc.  Calif.  Acad.  Sci.,  4th  Ser.,  vol. 
11,  p.  173,  Nov.,  1921),  as  "  near  iners  Meigen  "  from  St  Paul  Island, 
June  10    (G.  D.  Hanna). 

Family   BORBORIDAE. 

The  species  comprising  this  family  live  in  the  larval  state  in 
manure,  fungi,  carrion,  and  decaying  vegetable  matter.  One  species 
has  been  recorded  as  living  in  ants'  nests  and  another  in  water  col- 
lected in  epiphytic  bromeliads.  From  the  data  accompanying  the 
specimens  in  the  present  collection  I  assume  that  the  species  conform 
to  the  most  general  mode  of  life,  i.  e.,  pass  the  larval  stage  in  decay- 
ing vegetable  matter  or  in  manure. 

Genus  Borborus  Meigen. 

This  genus  is  represented  by  two  species:  annuhis  Walker  and 
subapterus  n.  sp.  The  former  has  the  normal  borborid  habitus  but 
the  latter  has  the  wings  much  abbreviated  and  is  thus  readily 
separated  from  any  described  species  occurring  in  North  America. 
To  facilitate  the  identification  of  annulus,  which  is  rather  poorly 
described  by  Walker,  it  is  redescribed  herewith. 

Cole  records  (Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11,  p.  173,  Nov., 
1921),  two  species  of  this  family  from  the  Pribilof  Islands,  "one  a 
Copromyza  {Borborus)  and  the  other  a  Leptocera,  but  the  specimens 
are  in  poor  condition  for  identification." 


210  NORTH   AMERICAN   FAUNA.  [No.  46 

Borborus  annulus  Walker. 

Borborus  annulus  Walker,  List  Ins.  Brit.  Mus.,  Diptera,  pt.  4,  p.  1129,  1849. 

Male  and  female. — Black,  shining,  with  a  slight  olivaceous  tinge. 
Head  black,  frontal  triangle  and  orbits  slightly  shining,  brownish 
pruinescent,  center  stripe  opaque  black,  anteriorly  reddish,  face 
brown,  reddish  or  yellowish  at  base  of  vibrissae,  distinctly  brownish 
pruinescent;  antennae  black  or  slightly  brownish.  Mesonotum  with 
yellowish  pruinescence  which  does  not  obscure  the  shining  black 
ground  color.  Abdomen  glossy/black.  Legs  black,  yellow  on  apices 
of  coxae,  trochanters,  extreme  bases  of  femora,  bases  of  tibiae 
(broadly),  apices  of  tibiae  (narrowly),  and  entire  tarsi.  In  addition 
to  these  yellow  markings  there  is  generally  a  similarly  colored  narrow 
band  near  the  apices  of  the  middle  and  hind  femora  which  is  not 
mentioned  in  the  original  description.  Wings  slightly  yellowish, 
veins  brown,  crossveins  broadly  infuscated.  Halteres  brownish 
yellow. 

Frons  as  broad  as  its  length  at  center;  orbits  each  with  two  slender 
bristles  directed  outward  over  eyes;  center  stripe  with  numerous 
setulose  hairs,  especially  on  anterior  portions  of  opaque  areas;  an- 
tennae of  moderate  size,  third  joint  rounded  anteriorly;  arista  with 
sparse  but  distinct  pubescence,  entire  length  of  arista  about  twice 
that  of  anterior  width  of  frons;  face  concave  in  profile,  distinctly 
produced  between  antennae  and  with  a  rounded  central  keel ;  labrum 
distinctly  protruded;  proboscis  large  and  fleshy;  vibrissa  long,  buccal 
bristle  shorter  than  vibrissa,  upwardly  directed.  Mesonotum  with 
3  pairs  of  dorso-centrals  and  between  these  4  longitudinal  rows  of 
short  setulose  hairs;  scutellum  with  4  marginal  bristles.  Abdomen 
broad  and  short,  first  visible  segment  elongated;  male  hypopygium 
of  moderate  size,  protuberant,  its  surface  with  numerous  short  hairs. 
Legs  rather  long,  fore  and  mid  femora  slightly  thickened  and  per- 
ceptibly bent;  mid  femora  with  3  bristles  near  apex  on  the  anterior 
surface;  mid  tibiae  with  a  series  of  6-7  short  bristles  from  base  to 
apex  on  antero-dorsal  surface;  hind  tibiae  with  an  outstanding 
setulose  hair  on  the  antero-ventral  surface  beyond  middle,  a  distinct 
but  slender  preapical  dorsal  bristle  and  a  rather  weak  apical  thorn- 
like spur ;  basal  joint  of  hind  tarsi  thickened  and  about  two-thirds  as 
long  as  second.  Distance  from  humeral  cross-vein  to  end  of  first 
vein  about  one-third  as  long  as  next  costal  division  and  slightly 
longer  than  third;  inner  cross- vein  slightly  before  middle  of  discal 
cell;  last  section  of  fourth  vein  slightly  longer  than  preceding  sec- 
tion; outer  cross- vein  upright;  fifth  vein  not  extending  to  margin 
of  wing.     Length  3.5-4  mm. 

Originally  described  from  "  York  Factory  and  St.  Martin  Falls," 
Canada.     Aldrich  states  that  the  last  named  locality  is  now  known 


19^.1  INSECTS   OF   THE   PRIBLLOF   ISLANDS.  211 

as  Martin's  Falls  and  is  located  "  in  longitude  86.30,  latitude  51.30, 
in  other  words,  about  200  miles  north  of  the  northern  arch  of  Lake 
Superior"  (Cat.  Dipt.  N.  Amer.,  p.  66).  Coquillett  has  since  re- 
corded the  species  from  Popof  Island,  Alaska,  and  Schwarz  lists  it 
from  Pribilof  Islands.  The  species  is  represented  in  the  material 
before  me  by  32  specimens  with  data  as  follows : 

St.  George  Island. 

2  specimens,  April  17,  1914  (G.  D.  Hanna). 

13  specimens,  June  4,  1914  (lot  2,  G.  D.  Hanna),  Taken  on  very  wet  soggy 
ground  near  Village  landing. 

3  specimens,  June  14,  1914  (lot  23,  G.  D.  Hanna),  Garden  Cove. 

4  specimens,  June  16,  1914  (lot  36,  G.  D.  Hanna),  Toward  East  Rookery 

from  village. 

2  specimens,  same  date  as  last  (lot  35,  G.  D.  Hanna). 

St.  Paul  Island. 

3  specimens,  May  16,  1913   (lot  27,  A.  G.  Whitney),  taken  in  Company 
House  bathroom. 

4  female  specimens,  May  23, 1914  (lot  153,  G.  D.  Hanna),  from  privy. 

2  specimens  taken  in  the  summer  of  1914,  one  marked  lot  210,  A.  G.  Whit- 
ney, and  other  collected  by  E.  A.  Preble. 

Eorborus  subapterus,  n.  sp.  PI.  XV,  fig.  27. 

Female. — Black,  shining ;  venter  of  abdomen  and  stems  of  halteres 
brown.  Wings  brown.  Frons  distinctly  longer  at  center  than  its 
greatest  width;  frontal  triangle  and  orbits  shining,  center  stripe 
opaque ;  each  orbit  with  2  long  slender  bristles  which  are  very  slightly 
outwardly  directed ;  center  stripe  with  hairs  much  as  in  annulus;  an- 
tennae rather  above  the  average  size,  third  joint  disclike ;  arista  very 
slender,  distinctly  but  sparsely  pubescent,  entire  length  of  arista  about 
\\  times  that  of  anterior  width  of  frons;  hairs  on  basal  joint  of  an- 
tenna long  and  fine ;  face  concave  in  profile,  distinctly  keeled ;  labrum 
much  protruded,  vibrissa  very  long  and  slender;  buccal  bristle  short 
and  hairlike,  upwardly  directed.  Mesonotum  with  numerous  rather 
long  discal  hairs  which  obscure  the  3  pairs  of  dorso-centrals;  scutel- 
lum  with  4  weak  marginal  bristles.  Basal  abdominal  segment  not 
noticeably  elongated;  all  segments  with  very  few  short  hairs.  Legs 
slightly  elongated;  fore  and  hind  femora  noticeably  stronger  than 
mid  pair;  mid  tibiae  with  3  bristles,  a  pair  about  one-fourth  from 
apex,  one  of  which  is  on  the  antero-ventral  and  the  other  on  the 
postero-ventral  surface,  and  one  on  the  dorsal  surface  near  apex ;  hind 
tibiae  with  weak,  hairlike  preapical  bristle,  and  weak,  almost  straight, 
apical  spur;  basal  joint  of  hind  tarsi  much  dilated  and  over  two- 
thirds  as  long  as  second ;  surfaces  of  all  legs  with  numerous  soft  hairs. 
Wings  abbreviated,  extending  to  middle  of  fourth  abdominal  seg- 
ment ;  venation  as  in  Plate  XV,  Figure  27.    Length  3.5  mm. 


212  NOETH   AMERICAN   FAUNA.  [No.  46 

Male. — Agrees  with  the  female  in  color  and  general  structure. 
Differs  in  having  the  abdomen  robust,  and  obtusely  rounded  at  apex, 
with  the  hypopygium  of  moderate  size. 

Type  locality. — St.  George  Island,  June  16,  1914,  "toward  East 
Kookery  from  village."  (Lot  No.  36,  G.  D.  Hanna.)  Two  para- 
types,  St.  George  Island,  along  Garden  Cove  Creek  (G.  D.  Hanna). 

Borborus  pedestris  Meigen,  a  European  species,  has  the  wings 
much  shorter  than  the  present  species,  scarcely  longer  than  the 
scutellum,  and  differs  also  in  color.  Leptocera  nivalis  Haliday,  a 
species  belonging  to  an  allied  genus  and  also  European,  occurs  in 
winter  generally  and  has  the  faculty  of  leaping  exceptionally  de- 
veloped. 

Genus  Leptocera  Olivier. 

This  genus  is  listed  as  Limosina  Macquart  by  Aldrich.  Leptocera 
is,  however,  an  older  name  for  the  same  genus  and  must  replace  it 
in  our  lists  as  indicated  by  Coquillett  in  his  paper  on  "  The  Type- 
species  of  North  American  Diptera." 16  Leptocera  was  erected  in 
1813  and  Limosina  in  1835.  There  is  a  single  species  represented  in 
the  present  material. 

Leptocera  limosa  Fallen. 

Copromyza  limosa  Fallen,  Dipt.  Suec,  Heteroniy,  8,  6   (1820). 

Three  specimens  that  evidently  belong  to  this  species  were  taken 
by  G.  D.  Hanna  on  St.  George  Island ;  two  bear  the  label  Lot  No.  2, 
and  one  Lot  No.  11,  the  former  being  taken  along  with  Borborus 
annulus,  June  4,  1914,  "  on  wet  soggy  ground  near  the  Village  land- 
ing," and  the  latter.  June  10,  1914,  "  near  beach  at  East  landing." 

13 remi  found  the  larvae  in  confervae  in  Europe  (Schiner). 

Aldrich  records  the  species  in  his  Catalogue  from  New  Jersey, 
White  Mountains,  N.  H.,  and  Montreal,  Canada.  I  have  taken  the 
species  in  Illinois  and  it  is  probably  of  general  occurrence  in  the 
United  States. 

Family  PHYCODROMIDAE. 

The  members  of  this  family  resemble  very  closely  the  genus  Bor- 
borus in  general  habits  and  have,  by  some  of  the  older  authors,  been 
treated  as  belonging  to  the  same  family  under  the  name  Copromzi- 
dae.  In  addition  to  resembling  them  in  appearance  they  also  have 
very  similar  habits,  feeding  mainly  upon  decaying  vegetable  matter, 
but  I  have  never  found  species  of  Coelopa  away  from  the  seashore 
or  the  shores  of  a  tidal  river  while  Borborus  and  other  genera  of 
that  family  may  be  met  with  in  almost  any  locality  where  a  suitable 
pabulum  is  obtainable.     Coelopa  is  the  only  genus  so  far  recorded 

18  Proc.  U.  S.  Nat.  Mus.,  vol.  37,  p.  559,  1910. 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  213 

from  North  America.  Of  the  three  species  recorded  two  are  con- 
sidered as  identical  with  two  of  those  occurring  in  Europe  and  those 
are  the  species  represented  in  the  present  collection. 

Genus  Coelopa  Meigen. 

Coelopa  frig-ida  Fallen.  PI.  XII,  fig.  1. 

Copromysa  frigida  Fallen,  Dipt.  Suee.,  Hydrom.,  p.  6,  1S20. 
Coelopa  frigida  Zetterstedt,  Dipt.,  Scand.,  vol.  6.  p.  2472,  1847. 
Coelopa  simplex  Haliday,  Ent.  Mag.,  vol.  1,  p.  1G7,  1833. 

This  species  is  represented  by  28  specimens  in  the  collection.    The 
data  upon  the  labels  are  as  follows: 
St.  Paul  Island. 

7  specimens,  August  19,  1914   (B.  A.  Preble). 

1  specimen,  August  1,  1914  (E.  A.  Preble). 
5  specimens,  summer,  1914    (E.  A.  Preble). 

2  specimens,  summer,  1914  (lot  210,  A.  G.  Wbitney). 

1  specimen,  May  23,  1914  (lot  156,  A.  G.  Wbitney). 

3  specimens,  August  16,  1914   (G.  D.  Hanna). 

St.  George  Island. 

2  specimens,  June  16,  1914   (lot  35,  G.  D.  Hanna). 
1  specimen,  June  17,  1914  (lot  42,  G.  D.  Hanna). 

3  specimens,  June  14,  1914  (lot  19,  G.  D.  Hanna). 

3  specimens,  May  6,  1914  (G.  D.  Hanna). 

Coelopa  eximia  Stenhammer.  PI.  XIV,  fig.  25. 

Copromysa  eximia  Stenhammer,  K.  vetensk,  Akad.  Handl.,  p.  318,  1854. 
Coelopa  frigida  Haliday,  Ent.  Mag,  vol.  1,  p.  167,  1S33. 

This  species  is  represented  by  13  specimens  in  the  collection.  All 
were  taken  on  St.  Paul  Island  and  bear  the  following  data: 

1  specimen,  May  23   (lot  154,  A.  G.  Wbitney). 

2  specimens,  August  1  (E.  A.  Preble). 

2  specimens,  August  19   (E.  A.  Preble). 

1  specimen,  summer,  1914   (lot  210,  A.  G.  Wbitney). 

4  specimens,  August  16,  1915  (G.  D.  Hanna). 

2  specimens,  August  16,  1915   (G.  D.  Hanna). 
1  specimen,  July  7,  1917   (G.  D.  Hanna). 

There  has  been  considerable  confusion  in  this  genus  because  of 
misidentification  by  various  authors  in  the  early  part  of  the  nine- 
teenth century.  Schiner  probably  had  his  synonymy  more  exact  than 
any  previous  author  and  his  names  have  been  accepted  by  the  present 
writer. 

Coquillett  recorded  frigida  and  nitidula  Zetterstedt  from  Alaska.17 
This  nitidula  is  the  species  I  record  as  eximia.  Hagen  has  recorded 
frigida  from  Massachusetts,18  and  Coquillett  has  recorded  it  from 

17  Proc.  Washington  Acad.  Sci.,  vol.  2,  p.  460,  1900. 

18  Can.  Ent.,  vol.  17,  p.  140,  1885. 


214  NORTH   AMERICAN   FAUNA.  [No.  46 

the  Commander  Islands.19  Cole  records  (Proc.  Calif.  Acad.  Sci., 
4th  ser.,  vol.  11,  p.  174,  Nov.  1921),  C.  parvula  Haliday  from  St.  Paul 
Island,  June  8  and  21,  1920  (G.  D.  Hanna).  This  also  is  the  species 
listed  here  as  eximia. 

A  key  for  the  separation  of  the  three  North  American  species  is 
given  herewith  as  there  is  no  previous  synopsis  available  to  students 
which  includes  all  three. 

Key  to  North  American  species  of  Coelopa. 

1.  Males 2. 

Females 4. 

2.  Legs  without  strong  bristles,  covered  with  long  woolly  hairs,  those  on  hind 

tibiae  very  conspicuous ;  cheeks  rather  densely  covered  with  soft  hair,  not 

bristly eximia. 

Femora   and    tibia   with  numerous   strong  bristles;    cheeks   with   numerous 
bristles  which  are  shorter  posteriorly 3. 

3.  Mid  tibiae  with  numerous  strong  bristles  on  entire  surface  except  ventrally, 

the  ventral   surface  with   moderately  long  soft  hairs ;    hind   tibiae  with 
slightly  stronger  bristles  than  mid  pair ;  bristles  on  hind  femora  extending 

to  base frigida. 

Mid  tibiae  with  very  long  woolly  hairs  especially  on  ventral  surface,  only 
2-3  bristles  present  on  apical  half  of  anterior  surface ;  hind  tibiae  with 
very  conspicuously  stronger  armature  than  mid  pair,  the  bases  of 
bristles  slightly  tuberculate ;  bristles  on  hind  femora  confined  to  apical 
half   ( Cresson,  Calif. ) vanduzeei. 

4.  Fore  and  hind  femora  and  tibiae  with  conspicuous  bristles vanduzeei. 

Fore  and  hind  femora  and  tibiae  without  conspicuous  bristles 5. 

5.  Cheeks  and  fore  femora  with  uniform  soft  hairs eximia. 

Cheeks  bristly ;  fore  femora  with  soft  hairs,  conspicuous  among  which  are 

several   longer,    moderately   strong    bristles    on    the    antero-dorsal     sur- 
face  frigida. 

Family  SCIOMYZIDAE. 

There  is  one  species  in  the  collection  that  in  certain  respects  re- 
sembles some  of  the  species  in  the  family  Helomyzidae,  but  which 
because  of  the  absence  of  the  vibrissae  and  of  the  costal  spines  must 
be  placed  in  the  Sciomyzidae.  It  agrees  with  no  described  genus  in 
the  latter  family,  so  that  I  am  forced  to  erect  a  new  genus  for  its 
reception. 

The  larvae  of  the  Sciomyzidae,  so  far  as  I  know  them,  are  aquatic 
or  live  in  damp  earth,  usually  on  the  margins  of  streams  or  ponds, 
and  feed  upon  decaying  vegetable  matter.  There  is  nothing  in  the 
records  pertaining  to  the  present  material  that  indicates  the  larval 
habits  of  the  species,  as  they  bear  only  the  date  of  collection. 

Genus  Pseudosciomyza,  n.  gen. 

Generic  characters. — Frons  broad;  orbits  with  3-4  bristles;  post 
vertical  bristles  divergent;  second  antennal  joint  much  shorter  than 

19  Rept.  on  Fur  Seals  and  Fur  Seal  Islands,  pt.  4,  p.  345,  1899. 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  215 

third;   third  joint  distinctly  longer  than  broad    (arista  missing) 
face  receding  toward  mouth  margin;  eye  small,  round,  barely  highe, 
than  cheek.     Thorax  with  the  following  bristles :  1  humeral,  2  noto 
pleural,  1  presutural,  2  pairs  dorso-central,  1  prealar,  and  2  postalar 
scutellum  with  4  bristles;  propleurum  with  a  bristle;  stigmatal  bris- 
tle absent;  mesopleura  unarmed;  sternopleura  with  numerous  long 
hairs  none  of  which  can  be  considered  as  bristles.     Legs  hairy,  with- 
out distinct  bristles  except  those  at  apices  of  mid  and  hind  tibiae,  the 
former  stronger  than  the  latter  and  noticeably  curved.     Costa  un- 
spined;  inner  cross  vein  before  apex  of  first  vein,  sixth  vein  reach- 
ing margin  of  wing.     Genotype,  the  following  species. 

Pseudosciomyza  hannai  Cole. 

Dryomyza  hannai  Cole,  Proc.  Calif.  Acad.  Sei.,  vol.  11,  p.  174,  1921. 

Male. — Brownish  black,  subopaque,  head  yellowish  brown;  vertex 
and  occiput  with  dense  grayish  pruinescence ;  f rons  orange  brown, 
darker  posteriorly;  orbits  pruinescent;  face  and  cheeks  testaceous 
yellow;  third  antennal  joint  slightly  brownish.  Thorax  with  rather 
dense  yellowish  gray  pruinescence.  Abdomen  more  brownish  than 
thorax,  the  surface  slightly  shining  and  with  but  little  pruinescence ; 
apices  of  segments  narrowly  yellowish.  Legs  reddish  brown; 
femora  slightly  darkened.  Wings  faintly  yellowish,  noticeably  so  on 
each  side  of  humeral  vein  and  between  apices  of  auxiliary  vein  and 
first;  cross  veins  with  very  indistinct  yellowish  marginal  suffusion. 
Halteres  pale  yellow. 

Frons  slightly  over  half  the  width  of  head;  orbits  slightly  differen- 
tiated, each  with  3-4  bristles  which  are  directed  slightly  outward; 
center  stripe  with  numerous  black,  setulose  hairs;  antennae  rather 
short,  not  descending  much  below  level  of  eyes;  face  slightly  carinate. 
upper  mouth  margin  arched;  labrum  small,  protruded  tongue-like; 
proboscis  fleshy;  palpi  rather  broad,  hairy;  disc  of  thorax  with 
numerous  long  setulose  hairs  among  which  it  is  difficult  to  distinguish 
the  bristles;  sternopleurum  with  long  hairs  on  its  entire  surface. 
Abdomen  Avith  long  and  rather  strong  hairs  on  all  segments;  hypo- 
pygium  rather  large,  knob-like.  Legs  stout,  the  femora  noticeably 
so,  hairy;  tarsal  claws  long.  Wings  elongate,  their  length  exceeding 
that  of  insect  from  head  to  tip  of  abdomen,  and  equal  to  3  times 
their  greatest  width;  first  costal  division  about  1.5  that  of  second, 
the  two  combined  exceeding  in  length  that  of  third;  second  vein 
ending  well  in  front  of  wing  tip;  third  vein  ending  very  close  to 
apex  of  wing;  inner  cross  vein  about  three  sevenths  from  apex  of 
discal  cell ;  last  section  of  fourth  vein  about  1.5  that  of  penultimate 
section. 

Female. — See  Cole's  description  (op.  cit.). 


216  NORTH   AMERICAN   FAUNA.  [No.  46 

Length,  5  mm.  St.  Paul  Island,  4  specimens,  August  19,  1914 
(E.  A.  Preble)  ;  1  specimen,  August  1,  1914  (E.  A.  Preble). 

This  genus  has  the  clypeus  well  developed  and  protuberant,  and 
because  of  this  character,  runs  to  the  subfamily  Dryomyzinae  in 
Melander's  recent  key  to  the  genera  of  Tetanoceridae 20.  It  is 
separable  from  Helcomyza  Curtis  by  the  absence  of  costal  spines 
and  the  mid  tibial  bristling  and  from  the  other  two  genera  by  the 
lack  of  posterior  bristles  on  mid  tibiae,  and  bristling  of  scutellum 
and  dorsum  of  thorax.  / 

This  genus  and  species  have  been  in  manuscript  since  1915,  but  the 
species  was  described  by  Cole  from  material  collected  on  St.  Paul 
Island  August  20,  1920.  The  author  of  the  species  indicated  that  its 
position  in  the  genus  Dryomyza,  in  which  he  placed  it,  was  doubtful, 
which  is  correct. 

Family  TRYPETIDAE. 

The  species  comprising  this  family  are  phytophagous  in  the  larval 
stage;  feeding  in  stems,  roots,  leaves,  flowers,  or  fruits,  and  giving 
little  or  no  indications  of  their  presence,  while  others  make  their 
presence  evident  by  the  formation  of  galls  upon  the  roots  or  stems, 
or  by  causing  large  blotches  upon  the  leaves  within  which  they  feed. 
There  is  in  the  collection  before  me  one  species  which  appears  to  be 
undescribed.  Loew  has  recorded  Spilographa  flavonotata  Macquart, 
and  Tephritis  angustipennis  Loew  from  the  Yukon  River,  Alaska; 
and  Coquillett  in  addition  to  describing  Trypeta  flaveola  from  Com- 
mander Islands  has  recorded  species  from  Canada  and  northern  and 
western  States  of  the  Union,  some  of  which  may  occur  in  Alaska,  but 
it  is  very  improbable  that  any  of  them  will  be  found  on  the  islands 
covered  by  the  present  investigation  because  of  their  connection  with 
certain  food  plants  which  do  not  occur  on  these  islands. 

Genus  Acidia  Kobineau-Desvoidy. 

The  genera  Acidia  and  Spilographa  are  in  my  opinion  not  validly 
separable.  The  only  difference  between  the  genotypes,  which  I  have 
examined,  lies  in  the  arrangement  of  the  thoracic  dorsocentral 
bristles.  In  Acidia  the  four  bristles  in  front  of  the  scutellum  are 
more  nearly  in  a  transverse  line  than  they  are  in  Spilographa.  The 
very  faint  distinction  between  the  wTing  markings  of  the  two  so-called 
genera  is  too  trivial  to  warrant  their  generic  separation. 

I  retain  the  generic  name  Spilographa  in  the  references  to  species 
in  the  following  paragraph  as  it  is  under  that  name  that  the  species 
have  been  recorded  in  the  literature. 

The  European  Spilographa  alternata  Fallen  lives  in  rose  hips, 
some  of  the  genus  live  in  fruits  of  Beiberis,  while S.  zoe  Meigen  and  S. 

30  Ann.  Ent.  Soc.  Amer.,  vol.  13,  p.  307,  1921. 


1923.]  INSECTS   OF   THE   PRIBILOF   ISLANDS.  217 

artemisae  Fabricius  in  the  larval  stages  mine  in  leaves  of  Artemisia 
and  allied  plants.  The  North  American  species  electa  Say,  is  found 
in  the  larval  stage  in  berries  of  Solarium  carolinense.  The  larval 
habit  of  the  species  before  me  is  not  known  but  it  is  undoubtedly  a 
phytophagous  species. 

Acidia  uncinata  Coquillett.  PI.  XV. 

Puparium  (PI.  XV,  fig.  37). — Length  4.5  mm.,  diameter  at  middle 
2.25  mm.  Color,  pale  yellowish  white,  slightly  shining.  Surface 
of  segments  very  minutely  transversely  rugulose,  appearing  except 
under  a  high  magnification  as  entirely  smooth.  Dorsal  thoracic 
segments  as  in  Plate  XV,  Figure  33,  the  second  and  third  with  small 
scalelike  setulae  on  their  anterior  margins  arranged  in  short,  slightly 
curved,  transverse  series.  All  segments  each  with  a  slightly  irregu- 
lar transverse  series  of  very  small,  rounded,  raised  areas  which 
are  rather  widely  separated  and  each  of  which  is  armed  at  apex  with 
a  weak  hair.  Apex  of  abdomen  as  in  figure  35 ;  spiracles  slightly 
elevated,  each  with  3  rather  conspicuous,  black,  slits  (PL  XV, 
fig.  35). 

Imago. — Male:  Brownish  testaceous,  shining.  Head  including  an- 
tennae and  palpi  pale  yellowish  testaceous,  center  stripe  of  frons 
opaque,  darker  than  face,  lower  orbits  subopaque,  upper  orbits  and 
triangle  shining.  Thorax  and  abdomen  distinctly  shining,  the  for- 
mer with  slight  yellowish  pruinescence ;  humeri  paler  than  disc  of 
thorax;  postnotum  with  a  large  blackish  spot  on  each  side.  Legs 
yellowish  testaceous.  Wings  with  blackish  or  brownish  markings 
as  follows:  A  brown  spot  beyond  humeral  vein,  a  similarly  col- 
ored spot  filling  the  space  between  apex  of  auxiliary  vein  and  apex 
of  first  vein  and  extending  posteriorly  as  far  as  second  vein  but  not 
connecting  on  the  disc  with  the  fusiform  spot  covering  the  inner 
cross  vein,  the  latter  extending  in  an  almost  straight  line  to  costa, 
filling  the  entire  cell  to  apex  of  second  vein  and  distinctly  indicated 
along  the  anterior  margin  of  the  cell  between  second  and  third  veins, 
apex  of  wing  infuscated,  outer  cross  vein  enclosed  in  a  brown  suf- 
fusion.    Hairs  and  bristles  black. 

Upper  frontal  orbits  elongated,  their  lower  extremities  extending 
beyond  apex  of  ocellar  triangle;  lower  orbits  each  with  3  bristles; 
eye  about  1.5  times  as  high  as  long;  cheek  about  one-sixth  the  height 
of  eye.  Thoracic  chaetotaxy  normal.  Fore  femora  with  ventral 
bristles,  the  other  pairs  unarmed ;  hind  tibiae  without  dorsal  setulae, 
only  a  few  weak  hairs  present.  First  and  third  wing-veins  setulose, 
the  latter  with  setulae  extending  well  beyond  inner  cross  vein;  apex 
of  third  vein  very  noticeably  curved  backward;  inner  cross  vein  at 
less  than  one-third  from  apex  of  discal  cell. 

Length,  .4.75  mm. 


218  NORTH   AMERICAN   FAUNA.  [No.  46 

Locality —St.  Paul  Island,  1  male,  spring  1913  (A.  G.  Whitney, 
Lot  35) ;  1  male  and  empty  puparium,  May  23,  1914  (A.  G.  Whitney, 
Lot  163)  ;  and  1  puparium,  fall  1913  (A.  G.  Whitney,  Lot  103).  Re- 
corded by  Cole  (Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11,  Nov.  1921, 
p.  175)  from  St.  Paul  Island,  August  10  and  12, 1920  (G.  D.  Hanna). 
Originally  described  from  Fort  Wrangel,  Alaska. 

Lot  35  in  list  is  given  as  "  Found  dead  on  moss  and  liverwort 
specimens  from  Tolstoi  Hill."  Lot  163,  "  Grassy  bankside  near  Vil- 
lage wells.  One  cocoon  and  fly.  /These  yellow,  ribbed  cocoons  found 
everywhere  in  lower  part  of  moss  beds  and  among  their  roots;  a 
moist  location."  Lot  103  "  On  C oelopleurum.  These  pupae  common 
everywhere  under  and  amid  thick  damp  moss  on  tundra  all  through 
the  summer  season." 

The  cocoons  referred  to  are  the  puparia  of  this  trypetid.  It  seems 
remarkable  that  the  puparia  should  be  so  very  common  as  stated 
above  and  that  only  2  flies  are  in  the  collection. 

The  puparium  of  this  species  has  a  very  well  defined  lateral  fusi- 
form area,  a  character  that  one  might  in  a  measure  associate  with 
the  family  Ortalidae  if  Bank's  paper  on  the  larvae  of  Diptera  were 
used  as  a  guide  to  the  identit}'  of  this  stage. 21  It  is  therefore  per- 
tinent to  point  out  that  the  paper  in  question  provides  only  charac- 
ters for  the  identification  of  such  species  as  might  reasonably  be 
expected  to  occur  in  the  stomach  of  man  as  accidental  introductions 
with  food  and  is  not  intended  to  cover  the  entire  Muscoidea.  A  num- 
ber of  species  in  Ortalidae  have  no  clearly  defined  lateral  fusiform 
areas  and  the  anal  stigmatical  areas  are  not  noticeably  elevated, 
while  some  Trypeticlae  have  both  distinct  lateral  fusiform  areas 
and  more  or  less  elevated  anal  stigmatical  areas. 

Family  SEPSIDAE. 

The  members  of  this  family  so  far  as  they  are  known  live  in  de- 
caying animal  or  vegetable  matter  or  in  preserved  foods  or  meats. 
The  only  genus  represented  in   the   present   collection  is  PiophUa. 

Genus  PiophUa  Fallen. 

Of  the  species  in  this  genus  one  at  least  is  of  economic  importance 
because  of  its  common  occurrence  in  cheese  and  preserved  meats. 
The  larvae  are  able  to  leap  short  distances  and  the  species  is  popu- 
larly known  as  the  Cheese  Skipper  (PiophUa  casei).  This  species, 
which  probably  originated  in  Europe,  has  been  found  in  human 
graves  and  is  distributed  throughout  the  whole  of  Europe,  and  North 
America  from  Alaska  to  Mexico.  Several  other  European  species 
occur  in  North  America.     One  of  the  species  in  the  present  collec- 

21  U.  S.  Dept.  Agric,  Bur,  Ent.,  Tech.  Ser.  22,  1912. 


1923.]  INSECTS   OF    THE   PRIBILOF   ISLANDS.  219 

tion  was  found  in  the  skull  of  a  dead  seal,  and  in  this  connection 
it  may  be  of  interest  to  record  that  several  species  are  found  in  car- 
casses of  dogs  and  other  animals  that  are  not  uncommon  on  the 
shores  of  rivers,  especially  near  the  sea,  in  Britain.  One  species  I 
haATe  taken  in  such  situations  I  have  met  with  nowhere  else. 

Piophila  anomala,  n.  sp.  PI.  XV. 

Larva. — Not  preserved.  Cephalopharyngeal  skeleton  as  in  Plate 
15,  Figure  36;  dissected  from  puparium. 

Puparium  (PI.  XV,  fig.  31). — Length,  3.5-4  mm.  Reddish  brown, 
slightly  shining.  Surface  with  fine  transverse  rather  irregular 
rugae  (fig.  31).  Anterior  respiratory  organs  very  small.  Segmen- 
tation rather  indistinct.  Posterior  spiracles  with  3  rather  indistinct 
slitlike  openings;  apex  as  in  Plate  XV,  Figure  32.  The  entire  body 
without  distinguishable  hairs  or  setulae. 

Imago. — Male  and  female. — Glossy  black.  Head,  with  the  excep- 
tion of  the  upper  portion  of  frons,  the  occiput  and  posterior  por- 
tion of  cheeks,  reddish  yellow;  palpi  yellow;  third  antennal  joint 
brownish.  Legs  black,  yellow  on  extreme  bases  and  apices  of  all 
femora,  the  bases  of  all  tibiae  and  basal  3  joints  of  mid  and  hind 
tarsi.  Wings  clear,  veins  yellowish.  Calyptrae  whitish.  Halteres 
yellow. 

Male. — Frons  distinctly  narrowed  anteriorly;  orbits  each  with  2 
bristles,  the  anterior  one  weaker  than  the  posterior;  disc  of  frons 
with  a  few  weak  hairs;  second  antennal  joint  with  a  rather  long 
apical  dorsal  hair,  third  joint  rounded;  arista  indistinctly  pubescent; 
vibrissa  as  long  as  arista;  cheek  nearly  half  as  high  as  eye.  Me- 
sonotum  with  2  dorso-centrals ;  scutellum  with  4  bristles.  Hypopy- 
gium  small.  Legs  rather  stout,  fore  tarsi  not  appreciably  thickened. 
Venation  normal. 

Female. — Agrees  in  color  with  the  male.  Ovipositor  long  and 
slender. 

Length,  3.75  mm. 

Type.— St.  George  Island,  July  4,  1914  (Lot  52,  G.  D.  Hanna)  ; 
toward  Zapadni  Rookery.  Allotj^pe  and  puparium,  St.  Paul  Island 
(Lot  176,  A.  G.  Whitney).  The  data  attached  to  this  lot  is  as  fol- 
lows :  "  Near  Village.  One  fly  and  several  pupa  cases.  The  pupa 
cases  were  found  June  14  in  the  interstices  of  the  nasal  bones  of  a 
fur  seal  skull  on  the  '  killing  field.'  From  these  one  fly  hatched  out 
June  20."  Paratypes,  St.  George  Island,  2  specimens,  August  4, 
1914  (G.  D.  Hanna) ;  St.  Paul  Island,  1  specimen,  August  19,  1914 
(E.  A.  Preble). 

This  species  differs  essentially  from  others  in  the  genus  in  having 
the  frontal  orbits  each  with  2  distinct  bristles.  P.  casei  Linnaeus 
usually  has  the  frons  much  darker  and  only  exceptionally  have  the 


220  NORTH   AMERICAN   FAUNA.  [No.  46 

orbits  any  hairlike  setulae.  The  disc  of  thorax  in  anomala  is  glossy 
black,  with  a  slight  bluish  tinge,  and  the  surface  has  rather  uniform 
short  hairs.  In  this  respect  the  species  agrees  closely  with  nigriceps 
Meigen,  but  the  latter  has  the  face  blackened  and  the  scutellum 
flattened  and  transversely  rugulose.  The  scutellum  in  anomala  is 
convex  and  smooth.  P.  casei  differs  from  both  anomala  and  nigri- 
ceps in  having  the  thorax  subopaque,  with  3  slight  longitudinal 
grooves  in  which  there  are  a  series  of  short  hairs,  the  remainder  of 
disc  being  bare  and  with  a  sligjat  olivaceous  tinge. 

The  foregoing  notes  are  drawn  from  a  comparison  with  North 
American  specimens  of  casei  and  nigriceps  named  by  Coquillett.  I 
have  no  European  examples  of  the  species. 

Since  the  completion  of  the  manuscript  of  this  paper  a  revision  of 
the  family  has  appeared  by  A.  L.  Melander  and  A.  Spuler.22  The 
species  described  herein  will  run  down  to  oriens  Mel.  and  Spul.  in 
their  key  to  species  of  Piophila,  but  the  legs  are  differently  colored 
in  my  species,  the  fore  coxae  in  anomala  being  mostly  black  and  the 
mid  and  hind  tibiae  largely  blackened,  whereas  in  oriens  the  fore 
coxae  and  mid  tibiae  are  entirely  yellow  and  the  hind  tibiae  and  the 
tarsi  less  broadly  blackened,  all  of  the  mid  tarsi  and  the  basal  4 
joints  of  the  hind  pair  being  yellow. 

There  is  a  narrow  dorso-central  stripe  on  abdomen  of  anomala 
which  is  transversely  rugose ;  no  mention  is  made  of  this  in  descrip- 
tion of  oriens. 

Cole  records  Piophila  oriens  (Proc.  Calif  Acad.  Sci.,  4th  ser.,  vol. 
11,  Nov.  1921,  p.  176)  from  St.  Paul  Island,  August  10,  1920  (G.  D. 
Hanna) .     Oriens  was  described  from  New  York  and  Massachusetts. 

Piophila  sp. 

A  female  taken  by  G.  D.  Hanna,  June  16,  1914  (lot  36),  on  St. 
George  Island,  differs  from  the  foregoing  in  having  the  antennae 
black;  the  cheeks  higher,  rugose  posteriorly;  the  humeri  and  center 
of  scutellum  slightly  reddish;  and  the  legs  darker.  Unfortunately 
there  is  but  one  poorly  preserved  example,  so  I  refrain  from  giving  it 
a  name. 

In  several  respects  the  specimens  agree  fairly  well  with  the  descrip- 
tion of  pilosa  Staeger,  a  species  recorded  from  Greenland.  The 
male  of  pilosa  is  distinguished  from  allied  species  by  the  rather  con- 
spicuous short  pilosity,  which  is  especially  noticeable  on  the  abdomen, 
and  by  the  black  antennae  and  very  dark  legs.  The  female  is  less 
noticeably  pilose.  Nigeirima  Lundbeck,  a  species  described  from 
Greenland,  differs  from  all  others  so  far  described  in  being  entirely 
black. 

"  Bull.  143,  Washington  Agri.  Exp.  Sta.,  1017. 


1923.]  INSECTS   OF    THE   PRIBILOF   ISLANDS.  221 

Family    EPHYDRIDAE. 

The  species  of  this  family  are  aquatic  in  habit,  the  larvae  being 
found  in  liquids  or  in  mud.  Some  few  species  are  met  with  in  the 
larval  stage  as  miners  in  stems  or  leaves  of  aquatic  or  marsh  plants. 
The  species  in  the  present  collection  are  similar  to  those  that  frequent 
moist  ground,  and  in  all  probability  the  larvae  will  be  found  in  the 
wet  mud  or  water  about  which  the  adults  occur.  Many  of  the 
species  are  flower  frequenters  in  the  adult  stage. 

Genus  Scatella  Robiueau-Desvoidy. 

.This  genus  contains  a  large  number  of  species  which  are  met  with 
even  more  commonly  in  the  Old  World  than  in  the  New.  The  great 
majority  of  the  species  so  far  described  have  the  wings  either  with 
dark  spots  on  a  clear  ground  or  clear  spots  on  a  dark  ground.  The 
species  in  the  present  collection  differs  from  these  groups  in  having 
the  wings  unspotted.  /Scatella  setosa  Coquillett  and  8.  stagnalis 
Fallen,  the  two  species  recorded  from  Alaska,  belong  to  the  group 
with  clear  spots  on  the  wings. 

Scatella  brunnipennis,  n.  sp. 

Male  and  female.  Subopaque  brown.  Face,  yellowish  brown, 
much  paler  than  frons,  the  latter  greenish  anteriorly  in  well  pre- 
served specimens,  cheeks  and  lower  part  of  back  of  head  slightly 
gray  dusted.  Thorax  slightly  shining  on  disc  anteriorly  in  well 
preserved  specimens  with  bluish  or  greenish  luster,  entirely  opaque  on 
pleura;  mesonotum  without  distinct  vittae,  abdomen  brown,  slightly 
shining  at  base,  becoming  glossy  on  second  segment  and  noticeably 
polished  towards  apex,  the  whole  with  a  distinct  bronzy  reflection. 
Legs,  brownish  black,  femora  with  slight  grayish  pruinescence. 
Wings  subfuscous,  unspotted;  veins,  dark  brown.  Halteres  brown 
or  yellow. , 

Frons  seen  from  above  over  3  times  as  wide  as  either  eye ;  2  strong 
orbital  bristles  on  each  side;  center  stripe  above  and  orbits  with  a 
number  of  short  setulose  hairs;  third  antennal  joint  barely  longer 
than  broad;  arista  short,  scarcely  exceeding  length  of  antenna,  its 
pubescence  very  short;  face  very  decidedly  convex,  with  numerous 
short  bristles,  those  on  mouth  margin  and  on  a  line  with  eye  margin 
but  some  distance  from  it,  most  distinct;  no  string  bristle  on  cheek. 
Humeral  area  with  a  few  setae;  notopleural  bristles  2  in  number; 
dorso-centrals  3,  the  anterior  and  middle  pairs  less  widely  separated 
than  posterior  pair,  acrostichals  in  2  regular,  complete  rows;  dorso- 
central  line  filled  in  between  bristles  with  short  setulae;  mesopleura 
108731°— 23 15 


222  NORTH   AMERICAN   FAUNA.  [No.  46 

with  1  strong  bristle  and  a  number  of  hairs,  those  on  posterior  mar- 
gin directed  backward  and  those  on  upper  margin  directed  upward ; 
sternopleurum  with  1  strong  bristle;  scutellum  subtriangular,  flat- 
tened on  disc,  anterior  pair  of  bristles  not  very  much  shorter  than 
posterior  pair.  Abdomen  with  sparse,  short,  surface  hairs.  Legs 
normal.  Wings  slightly  longer  than  entire  insect;  distance  from 
humeral  vein  to  end  of  first  vein  barely  more  than  one-fourth  as 
great  as  next  costal  division;  both  costal  breaks  distinct;  venation 
similar  to  that  of  stagnalis.      / 

Length,  2-3  mm. 

Type  locality. — St.  Paul  Island,  August  16,  1915,  60  specimens 
(G.  D.  Hanna).    Other  paratypes  as  follows: 

St.  Paul  Island. 

11  specimens,  August  1,  1914  (E.  A.  Preble). 
15  specimens,  August  19,  1914  (B.  A.  Preble). 

St.  George  Island. 

8  specimens,  June  4,  1914  (lot  2,  G.  D.  Hanna). 
1  specimen,  June  16,  1914  (lot  32,  G.  D.  Hanna). 

This  species  strongly  resembles  quadrisetosa  Becker,  differing, 
however,  in  the  yellow  instead  of  gray  face  and  the  absence  of  the 
strong  metallic  color  of  the  lower  part  of  the  frons.  Quadrisetosa 
is  a  Norwegian  species  that  has  not  been  recorded  from  this  side  of 
the  Atlantic,  except  by  Cole  (Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11, 
Nov.  1921,  p.  176)  and  of  his  specimens,  which  came  from  St.  Paul 
Island  June  21  and  August  10,  1920  (G.  D.  Hanna)  he  says  they 
"  seem  to  answer  the  description  of  this  form." 

Genus  Parydra  Stenhammer. 

A  genus  which  is  well  represented  in  Europe  and  North  America ; 
the  larvae  live  in  stagnant  water,  and  the  adult  flies  are  found  in 
marshy  situations. 

Parydra  metallica  Cole.  PI.  XV,  fig.  26. 

Parydra  metallica  Cole,  Proc.  Calif.  Acad.  Sci.,  4th  ser.,  vol.  11,  p.  176, 
Nov.  1921. 

This  species  was  in  manuscript  for  several  years  in  this  paper,  but 
has  been  described  by  Cole  as  above. 

It  is  an  aberrant  species  and  may  reasonably  be  removed  from 
Parydra,  but  I  do  not  consider  it  pertinent  to  do  so  in  this  paper. 

Originally  described  from  St.  George  Island,  June  28,  1920 
(G.  D.  Hanna).  In  the  present  collection  it  is  represented  by  three 
specimens  from  the  same  island,  June  4  and  16  (lots  2  and  36,  G.  D. 
Hanna). 


1923.1  INSECTS   OF   THE   PRIBILOF  ISLANDS,  223 

Family  DROSOPHILIDAE. 

The  known  larvae  of  the  species  of  this  family  feed  upon  decaying 
vegetable  matter,  exuding  sap  of  trees,  in  fermenting  liquids,  and 
rarely  in  leaves  of  living  plants. 

There  is  a  single  species  in  the  present  collection. 

Genus  Drosophila  Fallen. 

Drosophila  graminum  Fallen. 

Drosophila  graminum  Fallen,  Geomyzides,  p.  8,  1823. 

A  female  of  this  species  taken  on  St.  Paul  Island,  August  16, 
1915  (G.  D.  Hanna),  has  the  thoracic  stripes  well  defined  and  in 
every  respect  agrees  with  the  dark  forms  occurring  in  the  United 
States. 

Family  AGROMYZIDAE. 

There  is  but  a  single  species  of  this  family  in  the  collection.  It 
belongs  to  the  genus  Phytomyza,  the  species  of  which  are,  so  far 
as  known,  phytophagous  in  the  larval  stage,  usually  mining  in  the 
leaves  of  various  plants,  or  living  in  the  froth  of  Cercopidae. 

Phytomyza  obscurella  Fallen. 

Phytomyza  obscurella  Fallen,  Phytomyzides,  4,  1823. 

I  have  considerable  doubt  about  the  identity  of  this  species. 
Melander  has  had  an  opportunity  of  comparing  Alaskan  and  Euro- 
pean examples  of  this  species  and  considers  the  forms  ilicicola  Loew 
and  nigra  Meigen  as  varieties  of  obscurella,  listing  both  as  oc- 
curring in  Alaska.  The  recorded  food  plants  of  the  varieties  sug- 
gest confusion  of  species — ilicicola  on  holly,  obscurella  on  honey- 
suckle and  elder,  and  nigra  on  Primula  veris  and  Heracleum  sphon- 
dylium.  In  view  of  the  facts  that  I  have  no  European  examples 
of  obscurella  for  comparison  and  that  I  have  no  record  of  the  food 
plant  of  the  Alaskan  species  and  have  not  had  opportunity  to  com- 
pare the  larvae  and  pupae  of  the  different  forms,  I  leave  the  matter 
as  it  is,  merely  calling  attention  to  the  element  of  doubt  in  the 
matter  of  the  recorded  occurrence  of  obscurella  in  Alaska  and  the 
sinking  of  ilicicola  and  nigra,  as  varieties  of  obscurella. 

There  are  7  examples  that  I  place  under  this  species  name  pro- 
visionally.   The  data  are  as  follows: 

St.  George  Island. 

3  specimens,  June  16,  1914  (lot  32,  G.  D.  Hanna). 

4  specimens,  August  16,  1915  (G.  D.  Hanna). 


224  NORTH   AMERICAN   FAUNA.  (No.  46 

Agromyza  parvicella  Coquillett. 

This  species  was  originally  described  from  St.  Paul  Island,23  but 
no  specimens  were  found  in  the  present  collection. 

Immature  Stages  of  Diptera. 

Suborder  Orthorrhapha. 

Division  NEMATOCERA. 

Family   CHIRONOMIDAE. 

There  are  a  few  specimens  of  larvae  of  Chironomidae  in  alcohol, 
brief  descriptions  of  which  are  appended. 

Genus  Chlronomus  Meigen. 
G'hironomus  sp.  I. 

-Length,  10-12  mm.  Color  in  life  red.  Head  about  1^  times  as 
long  as  broad,  tapered  anteriorly,  eye  spot  duplicated;  antennae 
5-jointed,  basal  joint  about  4  times  as  long  as  its  diameter,  second 
joint  as  long  as  diameter  of  basal,  third  joint  about  as  broad  as  long, 
much  shorter  than  fourth  and  subequal  in  length  to  apical  joint; 
J  a  brum  with  4  long  hairs  on  each  side  of  center  anteriorly,  the  down- 
ward projecting  margin  with  fine  teeth;  transverse  comb  consisting 
of  9  rather  large  rounded  teeth;  mandibles  with  3  large  dark  teeth 
and  a  subapical  dorsal  and  median  paler  pair;  labrum  similar  to  that 
of  decorus  Johannsen.  Anal  ventral  blood  gills  absent;  anterior  and 
posterior  pseudopods  well  developed;  dorsal  anal  papillae  large; 
each  armed  with  about  6  long  hairs. 

St.  Paul  Island,  11  specimens,  August  10,  1913  (lot  98,  A.  G. 
Whitney),. 

The  data  for  this  lot  are  as  follows : 

About  10  larvae  from  mud  of  dried-up  pond  where  Leucasticte  had  beeu 
scratching  for  them.  This  pond  about  one-quarter  mile  long  and  18  inches 
deep  was  dry  from  August  5  to  mid-September.  During  this  time  its  whole 
mud  bottom  was  scratched  over  by  turnstones  to  get  at  these  larvae  which 
were  abundant.    Color  of  the  larvae  ruby-red. 

Genus  incertus. 

There  are  several  larvae  in  the  collection  that  I  do  not  know  the 
genus  of.  They  differ  from  any  larva  known  to  me  in  having  the 
apical  abdominal  segment  armed  with  several  concentric  series  of 
stout  hooks,  the  area  so  armed  being  but  slightly  elevated  and  re- 
sembling that  present  on  larvae  of  Simuliidae.  The  head,  however, 
is  of  the  normal  Chironomid  type  and  there  is  no  possible  doubt  as 
to  its  relationship  with  that  family.  The  head  is  short  and  broad, 
slightly  tapering  anteriorly;  the  antennae  are  of  moderate  length, 
the  basal  joint  short  and  stout,  not  twice  as  long  as  its  diameter,  the 

»  Coquillett,  D.  W.,  Joarn.  New  York  Ent.  Soc,  vol.  10,  p.  189,  1902. 


1923.]  INSECTS   OF    THE   PRIBILOF   ISLANDS.  225 

second  very  slender,  about  one-third  as  thick  as  basal  and  about  ono- 
third  longer  than  it,  apical  portion  consisting  either  of  one  joint  or 
2  very  closely  fused,  the  length  of  which  is  about  equal  to  the  di- 
ameter; apex  of  basal  joint  with  a  stout  process  which  tapers  appre- 
ciably apically  and  is  as  long  as  second  joint;  mandibles  with  5  teeth; 
labium  with  a  large  rather  irregularly  rounded  central  tooth  and  a 
much  smaller  rounded  one  on  each  side ;  labrum  with  2  pairs  of  stout 
protruded  ventral  processes. 

Length,  4-6  mm. 

St.  Paul  Island. 

7  specimens,  March  23,  1913  (lot  5,  A.  G.  Whitney). 
1  specimen,  April,  1913  (lot  10,  A.  G.  Whitney). 

The  specimens  in  lot  5  were  found  in  the  bottom  of  a  bag  in 
which  willows  and  mosses  had  been  collected.  It  is  possible  that  this 
is  the  larva  of  Smittia  but  more  data  are  necessary  before  a  reliable 
opinion  can  be  expressed. 

Family  LIMNOBIIDAE. 

Among  the  alcoholic  material  in  the  collection  there  is  a  larva 
that  has  puzzled  me  considerably.  The  head  is  complete  and  in  most 
respects  resembles  that  of  members  of  the  Mycetophilidae,  having  the 
typical  very  short  antennae,  apically  subtruncate.  toothed  mandibles, 
and  tapering  dorsal  cephalic  plate.  Were  I  judging  from  the  head 
alone  I  should  undoubtedly  place  the  species  in  the  Mycetophilidae 
but  the  respiratory  system  to  all  appearance  is  confined  to  prothoracic 
and  anal  spiracles,  the  apical  abdominal  segment  has  4  distinct  finger- 
like protuberances,  and  the  whole  of  the  body  is  covered  with  soft 
decumbent  hairs,  characters  that  associate  it  in  my  mind  with 
Trichocera  of  the  Limnobiidae,  usually  considered  a  subfamily  of  the 
Tipulidae. 

In  the  absence  of  pupae  and  imagines  of  this  species  It  is  not  pos- 
sible for  one  to  place  it  definitely,  as  our  knowledge  of  the  early 
stages  of  the  order,  although  increasing  slowly,  is  not  such  that  we 
can  identify  more  than  a  mere  fraction  of  the  species  in  the  larval 
stage. 

St.  Paul  Island :  1  specimen,  March  23, 1913  (lot  5,  A.  G.  Whitney). 

Found  along  with  some  chironomid  larvae  in  the  bottom  of  a  bag 
in  which  willows  and  mosses  had  been  collected. 

Suborder  Cyclorrhapha. 

Division   SCHIZOPHORA. 

Family  CALLIPHORIDAE. 

A  single  example  of  a  large  calliphorid  puparium  is  in  the  collec- 
tion.    As  no  examples  of  the  adults  of  this  family  are  before  me  I 


226  NORTH   AMERICAN   FAUNA.  [No.  46 

can  not  attempt  to  associate  it  with  any  species.  Muscids  are  treated 
in  the  paper  following. 

Length,  11.5  mm.  Reddish  testaceous.  Cephalic  and  caudal  ex- 
tremities slightly  tapering;  segments  well  differentiated;  anterior 
margins  of  segments  with  very  short  spines;  lateral  fusiform  area 
narrow;  spiracles  with  straight  slits  which  open  almost  directly 
laterad,  disclike  basal  elevation  not  distinguishable;  margin  of 
spiracular  area  elevated  so  that  the  spiracles  are  in  a  cavity;  spi- 
racular  field  with  12  distinct  tubercles  on  margin,  6  above,  the 
largest  being  the  inner  and  outer  pair  in  top  row  and  the  2  outer  on 
each  side  of  bottom  row;  anal  opening  with  a  large  conical  pro- 
tuberance on  each  side. 

St.  Paul  Island:  May  1,  1913  (lot  17,  A.  G.  Whitney). 


Explanation  of  Plates  XII-XV. 
Plate  XII.— Flies   (Cyclorrhapha). 
Fig.  1.  Coelopa  frigida  (male).  Fig.  2.  Poponota  kincaidi  (male). 

Plate  XIII. — Details  of  Flies  and  Midges  (Nematocera  and  Brachycera). 

Fig.     3.  Sciara  glacialis,  hypopygium  of  male,  apical  portion  of  one  side. 

Fig.     4.  Sciara  sp.     Same  as  Figure  3. 

Fig.     5.  Smittia  arctica,  hypopygium  of  male,  one  side. 

Fig.     6.  Chironomus  obtusilobus.     Same  as  Figure  5. 

Fig.     7.  Smittia  arctica,  antennae  of  male. 

Fig.     8.  Smittia  arctica,  antenna  of  female. 

Fig.    9.  Allodia  subelata,  hypopygium  of  male,  one  side. 

Fig.  10.  Orthocladius  obumbratus,  hypopygium  of  male,  apex  of  dorsal  plate. 
Fig.  11.  Tanytarsus  similatus.     Same  as  Figure  10. 

Fig.  12.  Smittia  arctica,  wing  of  male,  with  more  enlarged  section  of  costa. 

Fig.  13.  Chironomus    conformis,   hypopygium    of    male,    superior    process   and 
apex  of  lateral  arm. 

Fig.  14.  Exechia  casta,  hypopygium  of  male,  one  side. 

Fig.  15.  Rhanrphomyia  opacithorax,  hypopygium  of  male,  lateral  view. 

Fig.  16.  Chironomus  deviatus,  hypopygium  of  male,  one  side ;  a,  apex  of  supe- 
rior process. 

Plate  XIV. — Details  of  Flies    (Cyclorrhapha). — Figures  17-21,  apical  ventral 
abdominal  plate  of  males  ;  Figures  23-25,  hypopygia. 

Fig.  17.  Scatophaga  stercoraria.  Fig.  22.  Scatophaga  furcata. 

Fig.  18.  Scatophaga  dasythrix.  Fig.  23.  Scatophaga  furcata. 

Fig.  19.  Scatophaga  crinita.  Fig.  24.  Scatophaga  stercoraria. 

Fig.  20.  Scatophaga  rubicunda.  Fig.  25.  Coelopa  eximia. 

Fig.  21.  Scatophaga  islandica. 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey.  PLATE   XI  I. 


Flies  (Cyclorrhapha). 

Fig.  1.  Coelopa  frigida,  male.    Fig.  2.  Fogonota  kincaidi,  male. 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey.  PLATE   XIII. 


10 


Details  of  Flies  and  Midges  (Nematocera  and  Brachycera). 

(Explanation  on  page  226.) 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey.  PLATE   XIV. 


Details  of  Flies  'Cyclorrhapha  i 

(Explanation  on  page  226.) 


North  American  Fauna  No.  46,  U.  S.  Dept.  Agr.,  Biological  Survey.  PLATE   XV. 


Details  of  Flies  (Cyclorrhapha). 

(Explanation  on  page  227.) 


1923.]  INSECTS   OF   THE   PRIBTLOF   ISLANDS.  227 

Plate  XV.— Details  of  Flies  (Cyclorrhapha). 

Fig.  26.  Parydra  metallica,  head. 

Fig.  27.  Borborus  sabapterus,  wing. 

Fig.  28.  Allomyella  brevipennis,  head. 

Fig.  29.  Helina  liannai,  anal  opening  of  puparium. 

Fig.  30.  Helina  hannai,  apex  of  puparium,  end  view. 

Fig.  31.  Piophila  anomala,  puparium,  dorsal  view. 

Fig.  32.  Piophila  anomala,  end  view. 

Fig.  33.  Acidia  uncinata,  puparium,  dorsal  view  of  thoracic  segments  and  res- 
piratory organs ;  a,  opening  of  latter  enlarged ;  b,  spines  of  segments, 
enlarged. 

Fig.  34.  Hydrophoria  alaskensis,  abdomen  of  male,  lateral  view. 

Fig.  35.  Acidia  uncinata,  apex  of  puparium,  end  view,  and  spiracular  slits  en- 
larged. 

Fig.  36.  Piophila  anomala,  larval  head  parts. 

Fig.  37.  Acidia  uncinata,  puparium,  lateral  view. 


DIPTERA. 

Suborder  Cyclorrhapha. 

Division  SCHIZOPHORA. 

Family  CALLIPHORIDAE. 

By  W.  R.  Walton,  In  Charge  Cereal  and  Forage  Insect  Investigations, 
Bureau  of  Entomology. 

Cynomyia  hirta  Hough. 

This  species  was  originally  described  from  St.  Paul  Island  by 
Hough.  It  has  also  been  recorded  from  Popof  Island  and  Kodiak, 
Alaska,  by  Coquillett.  It  closely  resembles  G.  mortuorum  Linn,  of 
Europe,  but  is  evidently  distinct. 

The  specimens  before  me  exhibit  a  considerable  variation  in  size, 
i.  e.,  10-16  mm.  They  were  collected  on  both  St.  George  and  St. 
Paul  Islands  from  June  21  to  August  26. 

Calliphora  vomitoria  Liunaeus. 

This  species  has  been  recorded  by  Coquillett  as  occurring  in  Alaska 
as  follows:  Sitka,  Kukuk  Bay,  Popof  Island,  and  Seldovia.  This 
ib  the  species  mentioned  in  the  list  of  Diptera  as  Calliphora  obscoena 
Esch.  in  the  report  on  the  Fur  Seal  Islands. 

It  inhabits  also  northern  Europe,  Canada,  and  the  northeastern 
portion  of  the  United  States.  Specimens  were  collected  on  both 
St.  George  and  St.  Paul  Islands,  from  July  3  to  September. 


The  tachinid  Muscopteryx  obsctira  was  described  by  D.  W.  Coquillett  (New  Diptera 
from  North  America,  Proc.  U.  S.  Nat.  Mus.,  vol.  25,  p.  116,  Sept.  12,  1902)  from  St.  Paul 
Island,  and  afterwards  made  the  type  of  a  new  genus,  Alaskophyto,  by  C.  H.  T.  Townsend 
(Can.  Ent.,  vol.  47,  no.  9,  p.  285,  Sept.  1915).— W.  L.  M. 

Cole  records  (Proc.  Calif.  Acad.  Sci.,  4th  Ser.,  vol.  11,  p.  171,  1921)  Didyma  pullata  van 
der  Wulp  from  St.  Paul  Island.  There  is  no  such  species,  but  evidently  pullula  van  der 
Wulp  is  intended.  Van  der  Wulp's  species,  which  is  not  a  Didyma,  is  recorded  from 
Mexico  ;  and  Coquillett's  record  of  it  from  Alaska  is  erroneous.  In  all  probability,  there- 
fore, the  species  does  not  occur  in  the  Pribilofs. — J.  R.  M. 

228 


HYMENOPTERA. 

By  Henry  L.  Viereck,  Assistant  Biologist,  Bureau  of  Biological  Survey. 

In  the  following  list,  records  of  the  species  with  a  reference  let- 
ter24 are  taken  from  published  reports  and  those  without  a  refer- 
ence letter  are  here  published  for  the  first  time.  Where  the  type 
locality  of  a  species  is  in  these  islands  the  species  is  preceded  by  an 

asterisk  (*). 

Suborder  Chalastogastra. 

Superfamily  TENTHREDINOIDEA. 
Family  XYELIDAE. 
MegaxyelaP 

Determined  by  S.  A.  Rohwer.  One  head  from  St.  Paul  Island, 
from  the  stomach  of  Arguatella  ptilocnemis,  No.  14119. 

Family  TENTHREDINIDAE. 

*  Amauronematus  isolatus  Kincaid.  (6) 

Determined  by  S.  A.  Rohwer.  9  ,  St.  Paul  Island,  June,  1913. 
A.  G.  Whitney,  Lot  55. 

♦Amauronematus  wliitneyi  Rohwer. 

Type.— -St.  Paul  Island,  Bering  Sea,  April  15,  1914.  1  9  ,  A.  G. 
Whitney,  Lot  122. 

Amauronematus  sp. 

Determined  by  S.  A.  Rohwer.  Prooably  the  same  as  the  preced- 
ing. St.  Paul  Island,  May  27,  1915,  from  the  stomach  of  Stercora- 
rius  longicaudus  (No.  135031,  Biological  Survey  stomach  collec- 
tion). 

Pachynematus  gotarus  Kincaid. 

Determined  by  S.  A.  Rohwer.  3  9  4  $  ,  St.  George  Island,  June 
27,  1914  (G.  Dailas  Hanna),  2  S  ,  both  Lot  49;  and  2  9  ,  one,  Lot  49; 
one,  Lot  21. 

Pachynematus  sp. 

One  male  "  does  not  seem  to  be  described. "  Determined  by  S.  A. 
Rohwer.     St.  George  Island,  June  14,  1914  (G.  Dallas  Hanna). 

24  The  reference  letters  have  relation  respectively  to  (a)  Ashmead,  W.  H.,  Hymenoptera 
of  Alaska,  Proc.  Washington  Acad.  Sci.,  vol.  4,  pp.  117-274,  May  29,  1902;  and  (6)  Kin- 
caid, T.,  Tenthredinoidea  of  the  Expedition,  same  journal,  vol.  2,  pp.  341-346,  Nov.  24, 
1900.  The  sawflies  here  listed  with  Rohwer  as  author  were  described  in  Proc.  U.  S.  Nat. 
Mus.,  vol.  57,  pp.  212,  216,  1920. 

229 


230  NORTH   AMERICAN   FAUNA.  [No.  46 

Pteronidea  melanostoma  Rohwer. 

Type. — St.    George    Island,    Bering    Sea,    June    14,    1914.     1 9 
G.  D.  Hanna  (Lot  21;  Lot  12  according  to  original  description). 

Dolerus  konowi  MacGillivray. 

Determined  by  S.  A.  Rohwer.  St.  Paul  Island,  summer,  1914. 
A.  G.  Whitney,  Lot  202,  1  9  . 

Dolerus  sp. 

Determined  by  S.  A.  Eohwer.  /  1  9  ,  St.  Paul  Island,  summer,  1914. 
(E.  A.  Preble.) 

Suborder  Clistogastra, 
Superfamily  ICHNEUMONOIDEA. 
Family  VIPIONIDAE. 
*Apanteles  (Protapanteles)  alticola  Ashmead.    (a) 

*Apanteles  (Protapanteles)  congestiformis,  n.  sp. 

Type  locality.— St.  Paul  Island,  May  29,  June  20,  1913  (A.  G.  & 
E.  G.  Whitney,  lot  No.  50),  cocoons  collected  May  29,  1913. 

Female. — Length  2.5  mm.  Related  to  A.  (P.)  carduicola  (Pack- 
ard) and  may  prove  to  be  at  most  a  race  of  A.  (P.)  congestus  (Nees.), 
from  which  it  differs  in  its  mostly  black  fore  femora.  Shiny,  partly 
polished,  partly  pale  sericeous;  head  above  mostly  polished,  indis- 
tinctly sculptured,  face  almost  polished,  finely  indistinctly  punctured, 
without  a  median  welt,  labrum  blackish,  basal  half  of  mandibles 
black,  the  apical  half  mostly  reddish,  antennae  black  throughout 
except  for  the  pale  joint  between  the  pedicel  and  flagel,  palpi  dark 
stramineous ;  thorax  mostly  closely  punctured,  the  punctures  shallow 
and  adjoining  or  nearly  adjoining,  scutel  almost  impunctate,  polished, 
sparsely  punctured,  posterior  half  of  mesopleura  mostly  polished, 
impunctate,  wings  with  a  faint  brownish  tinge,  almost  colorless, 
stigma  brownish  stramineous,  veins  dull  stramineous  with  a  smoky 
tinge,  transverse  cubitus  a  little  longer  than  the  first  abscissa  of  the 
radius,  tegulae  black,  legs  black  or  blackish  except  for  the  distal 
trochanter  of  the  fore  legs,  the  apical  fourth  of  fore  tibiae  and  all 
of  the  remaining  tibiae  and  most  of  the  tarsi  which  are  rather  pale 
brownish  stramineous,  end  joints  of  the  tarsi  blackish  as  are  the 
penultimate  and  antepenultimate  tarsal  joints  of  the  fore  and  mid 
legs,  metapleura  with  the  anterior  half  mostly  polished,  the  pos- 
terior half  not  nearly  so  coarsely  reticulated  as  propodeum;  pro- 
podeum  shiny,  rather  coarsely  reticulated  and  with  a  distinct  median 
longitudinal  carina;  abdomen  with  its  first  and  second  plates  more 
or  less  sculptured,  the  first  plate  finely  wrinkled,  partly  indistinctly 
punctured,  partly  longitudinally  striate,  second  plate  not  so  definitely 


1923.]  INSECTS   OF    THE   PRIBILOF   ISLANDS.  231 

sculptured,  the  succeeding  tergites  highly  polished,  abdomen  black 
throughout,  ovipositor  hardly  exserted. 

Allotopotype. — Essentially  as  in  the  type,  except  that  the  legs  are 
black  or  blackish  throughout  except  for  the  yellowish  annulus  near 
the  base  of  the  tibiae.  Cocoons  imbedded  in  a  mass  of  pale  lemon- 
tinted  floss. 

Family  ALYSIIDAE. 

*Gyrocampa  alaskensis  Ashmead.  (a) 

Family    BANCHIDAB. 

Enizemum  tibiale  (Cresson).(°) 

Family  BRACONIDAE. 

Ichneutes  reunitor  Nees. 

Two  specimens  from  St.  George  Island,  June  14,  16,  1914, 
(G.  Dallas  Hanna,  lots  22  and  36.) 

Family    ICHNEUMONIDAE. 

♦Monoctonus  paulensis  (Ashmead)    (°)    (=ApMdius). 

Many  specimens  from  St.  George  Island.  One,  April  7,  1914 
(emerged  apparently  from  an  empty  aphid  skin  preserved  with 
this  specimen)  (A.  G.  Whitney,  lot  120)  ;  the  remaining  specimens 
were  collected  June  4,  10,  16,  1914,  and  are  labeled,  respectively,  G. 
Dallas  Hanna,  lots  2,  11,  28,  and  32. 

♦Aphidius  propinquus  Ashmead (a)  =  (A.  frigidus  Ashmead). 

Five  specimens  from  St.  George  Island,  June  10,  16,  1914,  August 
16,  1915  (G.  Dallas  Hanna,  lots  11,  12,  and  32). 

*Praon  alaskensis  Ashmead. (a) 
*Catastenus  alaskensis  Ashmead. («) 
Catastenus  trifasciatus  Ashmead.  «*> 
Mesochorus  frontalis  Ashmead.  («■) 
Campoplegidea  laticinctus  Cresson.W 
*Hypocryptus  variegatipes  Ashmead. 
Cteniscus  clypeatus  Cressou.w 
♦Polyblastus  glacialis  Ashmead.W 
Mesoleius  stejnegeri  Ashmead. W 
♦Calliphrurus  minor  Ashmead.  «*> 
*Calliphrurus  affinis  Ashmead.  (°> 
♦Calliphrurus  clypeatus  Ashmead. <«) 
♦Tryphon  alaskensis  Ashmead.  (Q) 
Stenomacrus  borealis  Ashmead.  (°) 
Stenomacrus  sp. 


232  NORTH   AMERICAN   FAUNA.  [No.  46 

Two  specimens  from  St.  George  Island,  June  16,  17,  1914  (G. 
Dallas  Hanna,  lots   32,  37). 

Stenomacrus  sp. 

One  specimen  from  St.  George  Island,  July  8,  1914  (G.  Dallas 
Hanna,  lot  55). 

*Ortbocentrus  nigritus  Aslimead.(°) 
Atmetus  insularis  Ashmead.(a) 
Deleter  flavifrons  Ashrnead.(a)  / 

*Neuroteles  dubiosus  Asbniead.(a) 
*Hypoleptus  alaskensis  Ashmead.<«) 
*Synoplus  pleuralis  Ashmead.(°) 
*Synoplus  brevipennis  Asbmead.<°) 
*Lissonota  alaskensis  Asbmead.W 
*Gelis  obesus  Asbmead=  (Pezomachus) . 
Gelis  sp. 

One   female   from   St.   Paul   Island,  August  31,  1914,   from  the 
stomach  of  Pisobia  aurita  (No.  134907,  Biological  Survey  stomach 
collection). 
*Gelis  nigrellus  Ashmead«J)  =  {Pezomachus). 

Atypical  females  representing  at  most  perhaps  only  a  dark  variety 
of  this  species  were  collected  as  follows :  St.  George  Island,  June  8, 
10,  1914  (G.  Dallas  Hanna,  Lot  Nos.  8  and  11,  respectively) ;  St. 
Paul  Island,  August  16, 1915  (G.  Dallas  Hanna). 

Gelis  sp. 

One  female  from  St.  Paul  Island,  August  31,  1914,  from  the 
stomach  of  Pisobia  aurita  (No.  134907)  represents  a  species  pre- 
sumably related  to  G.  posthumus  Foerster. 

*Mesoleptus  kincaidi  Ashmead(a)  =  (Exolytus). 
Mesoleptus  niger  Ashmead(a)  =  (Exolytus). 
*Mesoleptus  perplexus  Asbmead(a)  =  (Exolytus). 
*Xestopbyes  nigripes  Ashuiead(°)=  (Xestophya) . 

St.  Paul  Island,  Summer,  1914  (A.  G.  Whitney,  Lot  205). 
*Xestopbyes  polita  Asbinead(o)  =  (Xestophya). 
*Polyrbembia  sanctipauli  Asbmead<a>=  (Exolytus). 

Two  specimens  from  St.  Paul  Island,  August  31,  1914,  from  the 
stomach  of  Heteroscelus  incanus  (No.  134917,  Biological  Survey 
stomach  collection). 

Polyrbembia  sp. 

Three  specimens  from  St.  Paul  Island,  August  31,  1914,  from  the 
stomach  Pisobia  aunta  (No.  134907,  stomach  collection). 


1923.]  INSECTS   OF    THE   PRIBILOF   ISLANDS.  233 

Seleucus  sp. 

One  female  from  St.  Paul  Island,  August. 27,  1914,  from  the  stom- 
ach of  Arquatella  philocnemis  (No.  126723,  stomach  collection). 
Apparently  not  represented  in  any  published  key  to  species  of  this 
genus. 

Bachia  nigra  Ashrnead.<°) 
Bachia  sp. 

One  female  from  St.  Paul  Island,  summer  1914.  (E.  A.  Preble.) 
Presumably  related  to  Phygadeuon  longigena  Thomson. 

Scinacopus  sp. 

One  male  and  its  cocoon  from  St.  Paul  Island,  near  Polovina 
Lake,  March  29,  1914  (A.  G.  Whitney,  Lot  115),  "Pupa  in  brown 
papery  cocoon  brought  home  in  bunch  of  lichens  and  hatched  out  in 
vial."    Presumably  related  to  Phygadeuon  perfusor  (Gravenhorst). 

Zaphleges  sp. 

One  male  from  St.  Paul  Island,  August  16,  1915  (G.  Dallas 
Hanna).  Also  presumably  related  to  Phygadeuon  perfusor  (Grav- 
enhorst). 

♦Plesignathus  rubrocinctus  Ashmead<°)  =  (Plesiognathus). 
Plesignathus  sp. 

One  female  from  St.  George  Island,  September  6,  1913  (G.  Dallas 
Hanna).    Presumably  related  to  Phygadeuon  vagans  Gravenhorst. 

Plesignathus  sp. 

One  male  from  St.  George  Island,  September  6,  1913  (G.  Dallas 
Hanna).    Presumably  related  to  Phygadeuon  brachyurus  Thomson. 

*Bathymetis  simulator  Ashmead.W 
Bathymetis  quadriceps  Ashmead.(°) 
^Bathymetis  confusa  Ashrnead.<°) 
*Bathymetis  simillima  Ashmead.(o) 

St.  Paul  Island,  August  16,  1915  (G.  Dallas  Hanna) ;  St.  George 
Island,  August  4,  1914  (G.  Dallas  Hanna.) 

Bathymetis  rubrocincta  Ashmead.(°> 
*Bathymetis  simulans  Ashmead.<°> 
*Bathymetis  imitator  Ashmead.(a) 
♦Bathymetis  nigricornis  Ashmead.(°) 
*Stiboscopus  mandibularis  Ashmead.d) 
*Stiboscopus  alaskensis  Asbmead.d) 
*Stiboscopus  sanctipauli  Ashmead.W 
Stiboscopus  sp. 


234  NORTH   AMERICAN   FAUNA.  [No.  46 

One  male  from  St.  Paul  Island,  August  16,  1915  (G.  Dallas 
Hanna).     Presumably  related  to  Phygadeucm  liostemus  Thomson. 

*Pezoporus  trifasciatus  Ashmea&w *=  (Microcryptus) . 

Stibeutes  nigrita  Ashmead.e*) 

*Isochresta  unicincta  Ashmead.<°> 

♦Theroscopus  rufipes  Ashmead.c*) 

*Habromma  nigrum  Ashmead.c*) 

*Aclastus  rufipes  Ashmead.w 

Acrolyta  aciculata  Ashrnead.O*) 

Caenomeris?  sp. 

One  female,  St.  Paul  Island,  August  16,  1915  (G.  Dallas  Hanna). 
Presumably  related  to  Aclastus  minutus  (Bridgman.) 

♦Spinolia  minuta  Ashmead.W 

*Centeterus  dorsator  Ashmead.(°) 

Amblyteles  (Pterocormus)  alpestriformis,  n.  sp. 

Type  locality.— -St.  Paul  Island,  Telegraph  Hill,  June  22,  1913; 
spring  and  summer  of  1915  (A.  G.  Whitney,  Lot  Nos.  54,  174,  208). 
Presumably  related  to  P.  alpestris  (Holmgren). 

Female. — Length  9  mm. ;  colored  somewhat  like  Pterocormus  ?  dis- 
parilis  (Cresson),  from  the  original  description  of  which  it  differs 
as  follows:  Head  reddish,  except  for  the  cheeks,  malar  space,  and 
antennal  basin,  all  of  which  are  mostly  black;  mandibles  reddish, 
blackish  at  base  and  apex;  palpi  fuscous;  antennae  blackish,  except 
for  the  basal  fourth,  which  is  mostly  reddish,  without  an  annulus; 
apical  third  of  antennae  with  the  joints  slightly  faceted  above; 
thorax  with  its  tegulae  stramineous,  without  a  yellow  line  before  and 
beneath;  scutel  and  postscutel  reddish;  wings  subhyaline,  brownish; 
veins  and  stigma  pale  brownish  stramineous;  legs  mostly  reddish; 
coxae  and  trochanters  mostly  black;  hind  coxae  reddish  above;  end 
joint  of  tarsi  brownish ;  propodeum  slightly  concave  posteriorly,  with- 
out lateral  angles ;  propodeal  carinae  well  defined ;  areola  wider  in 
front  than  behind,  nearly  quadrate,  slightly  rounded,  emarginate  be- 
hind ;  areola  rather  indefinitely,  coarsely  sculptured ;  abdomen  finely 
reticulated,  its  punctures  mostly  from  adjoining  to  two  puncture 
widths  apart;  second  tergite  apically,  fifth  basally,  and  third  and 
fourth  tergites  broadly  down  the  middle,  black ;  second  and  third  ter- 
gites  without  yellow  spots ;  sixth  and  seventh  tergites  with  a  median 
yellow  spot ;  post-petiole  dullish,  finely  sculptured,  almost  impunctate. 

Other  locality. — St.  George  Island,  August  4,  1914  (G.  Dallas 
Hanna). 


1923.]  INSECTS   OF   THE  PRIBILOF   ISLANDS.  235 

Amblyteles  (Pterocormus?)  cervnlus  Provancher.(°) 
Superfamily  CYNIPOIDEA. 
Family  FIGITIDAB. 
Alloxysta  alaskensis  Ashrnead.«») 
Alloxysta  sp. 

From  St.   Paul  Island,  August   15,  1914    (No.    126733). 

♦Tetrarhapta  alaskensis  Ashmead.  (») 
St.  Paul  Island,  August  16,  1915  (G.  Dallas  Hanna). 

Superfamily  CHALCIDOIDEA. 

Family  MYMARIDAE. 
Anaphes  sp. 

Two  specimens  of  a  species  924  \k  long  and  apparently  related  to 
A.  hercules  Girault,  the  largest  North  American  species.  St.  George 
Island,  Staraya  Artel  Rookery,  June  17,  1914  (G.  Dallas  Hanna, 
Lot  37). 

Family  PTEROMALIDAE. 

♦Eutelus  confusus  Ashmead.(°) 
"  Habrocytus  capreae  Swederus." 

This  is  pt-obably  what  is  intended  by  Trvdymns  cap?rae  L.  in 
Schwarz's  report  (Fur  Seals  and  Fur  Seal  Islands,  Part  3,  p. 
550,  1899). 

Asapb.es  sp. 
St.  Paul  Island,  August  16,  1915  (G.  Dallas  Hanna). 

Pterosema  sp. 

One  male  from  St.  Paul  Island,  Tolstoi  Hill,  spring  1913,  "  prob- 
ably about  May  20,"  from  pupae  in  grass  stems  growing  in  shelter 
of  rock  crevices,  emerged  in  warmth  of  laboratory  (A.  G.  Whitney, 
Lot  33). 

Family  MISCOGASTERIDAE. 
Stictomiscbus  sp. 

One  female  from  St.  Paul  Island,  August  16,  1915  £G.  Dallas 
Hanna). 

♦Terobia  vulgaris  Asbmead.w 

Superfamily  SERPHOIDEA. 
Family  CERAPHRONIDAE. 
♦Lygocerus  alaskensis  Asbmead.(»> 


236  NORTH   AMERICAN   FAUNA.  [No.  46 

Family  BELYTIDAE. 

♦Zelotypa  scutellata  Ashrnead.w 
Zelotypa  sp. 

From  St.  Paul  Island,  August  15,  1914  (No.  126733). 

Family  SERPHIDAE. 

♦Serphus  nigripes  Ashmead.W 
St.  Paul  Island,  August  16,  1915  (G.  Dallas  Hanna). 

Serphus  sp. 
St.  George  Island,  August  4,  1914  (G.  Dallas  Hanna). 

SUPERFAMILY   APOIDEA. 

Family  APIDAE. 

Bremus  (Bremus)  kincaidi  Cockerell. 

Many  females  and  workers  from  St.  Paul  Island,  as  follows:  Fe- 
males, October  11,  1912  (A.  G.  Whitney,  Lot  2)  ;  May  31,  June  5, 
1913  (A.  G.  Whitney,  Lots  113  and  36,  respectively)  ;  workers,  July 
23,  1913  (A.  G.  Whitney,  Lot  95). 


In  the  report  on  G.  D.  Hanna's  1920  Pribllof  collection,  A.  D.  MacGillivray  describes 
(Proc.  Calif.  Acad.  Sci.,  Fourth  Ser.,  11,  pp.  188-192,  Nov.,  1921)  6  new  species  of 
sawflies  under  the  following  names :  Pontania  sueta,  P.  stipata,  P.  subatrata,  P.  sub- 
lorata,  Pachynematus  venustus,  and  P.  vermis. — W.  L.  M. 


ARACHNIDA 

By  Nathan  Banks,  Museum  of  Comparative  Zoology. 
Harvard  University. 

(Plate  IX,  figs.  1-7— see  p.  158.) 

Suborder  Acarina. 

Family  BDELLIDAE. 

Bdella  frigida  Banks. 
St.  George  Island,  July  9. 

Cyta  brevirostris  Koch. 
St.  Paul  Island,  April. 

Family  ERYTHRAEIDAE. 

Erythraeus  tonsus  Koch. 
St.  Paul  Island,  July  19,  on  Lathyrus  maritima. 

Family  PARASITIDAE. 

Parasitus  borealis  Koch.  (PI.  IX,  figs.  5  and  7.) 

St.  Paul  Island,  May  16  and  19;  May  23,  on  flies;  June  26. 

Macrocheles  arcticus  Kramer  and  Neuman. 
St.  Paul  Island,  May  23,  in  moss;  September  13,  on  chickweed. 

Family  IXODIDAE. 

Ceratixodes  putus  Cambridge. 

St.  Paul  Island,  July  4,  on  Rissa  t.  pollicaiis,  also  May  17  and  June 
30;  and  St.  George  Island,  Aug.  4. 

Family  ORIBATIDAE. 

Galumna  lucens  Kocb. 
St.  Paul  Island,  July  17,  on  chickweed ;  Sept.  13,  on  chickweed. 

Oribatella  borealis  Banks. 

St.  Paul  Island,  July. 
Notaspis  serrifrons,  n.  sp.  (PI.  ix,  fig.  2.) 

Yellowish,   cephalothorax    subtriangular,   with   large   submedian 
lamellae,  each  ending  in  a  long  bristle,  a  pair  of  short  apical  bris- 
tles, superior  bristles  long  and  fine;   pseudostigmatic  organ  short 
and  capitate ;  the  front  margin  of  cephalothorax  has  a  row  of  about 
108721°— 23 16  237 


238  NORTH   AMERICAN  FAUNA.  [No.  46 

15  sub-equal  teeth.  Abdomen  nearly  globose,  without  hairs;  ventral 
apertures  nearly  circular,  the  genital  fully  twice  its  length  in  front 
of  the  twice  larger  anal  aperture;  coxal  plate  with  two  lines  each 
side;  legs  short,  with  simple  hairs. 

Length  .5  mm. 

From  St.  Paul  Island,  July  (Whitney). 

Scutovertex  nigrof emoratus  Koch. 

Numerous  specimens  from  St^Paul  Island,  Aug.  16. 
Hermannia  reticulata  Thor. 

St.  Paul  Island,  July. 

Nothrus  sp. 

Young  from  St.  Paul  Island,  June. 

Lohmannia  scabra  Koch.  (PI-  IX,  fig.  4.) 

St.  Paul  Island,  April  5 ;  October  on  Erytrichium  chamissonis. 

Family  TYROGLYPHIDAE. 

Dermacarus  sp.  (PI-  EEf  nS- 1.) 

Hypopi  from  St.  Paul  Island,  July  5,  on  Sorex  pribilofensis. 

Tyrog-lyphus  whitneyi,  n.  sp.      ,  (PI.  IX,  fig.  6.) 

Hypopus. — Very  similar  to  T.  lundbecki  Trag.  from  Greenland. 
The  body,  however,  is  not,  or  is  only  a  trifle  broader  at  humeri  than  at 
the  hind  coxae.  The  hind  legs  are  situated  only  a  little  in  front  of 
the  sucker-plate,  and  when  extended  behind  reach  fully  to  tip  of  the 
body.  The  plate  is  similar  to  that  of  T.  lundbechi  except  that  the 
median  pair  of  suckers  are  very  much  larger  than  the  others,  in 
fact  more  than  twice  as  large  as  any  others.  (PI.  IX,  fig.  6.) 
Length,  .2  mm. 
From  St.  Paul  Island,  on  Bombus,  May  31  (Whitney  coll.). 

Family  LISTROPHOKIDAE. 

Myocoptes  musculinus  Koch. 
St.  Paul  Island,  November  16,  on  house  mouse. 

Family    ANALGESIDAE. 
Analges  sp. 
Females  from  St.  Paul  Island,  April,  without  host. 

Suborder  Phalangida. 
Family  PHALANGIIDAE. 

Leptobunus  borealis  Banks. 

Common  on  both  St.  Paul  and  St.  George  Islands  from  the  first  of 
June  till  fall. 


1023.]  ARACHNIDS    OF   THE   PRIBILOF   ISLANDS.  239 

Suborder  Araneida. 

Family  AGELENIDAE. 

Tegenaria  derbami  Scopoli. 

St.  Paul  Island,  May  11,  and  "  Summer " ;  St.   George   Island, 
June  27. 

Family  THERIDIIDAE. 

Bathypbantes  pogonias  Kulczynski. 
From  St.  Paul  Island,  July  3. 

Microneta  ululabilis  Keyserling. 

St.  George  Island,  June  17  and  September. 
Erigone  arctica  White. 

Several  from  St.  Paul  Island,  summer. 

Erigone  psycb.ropb.ila  Tborell. 

Several  from  St.  George  Island,  July  4. 

Erigone  sp. 

Two  females  from  St.  George  Island,  July  4,  and  August  4,  another 
from  St.  Paul,  August  17. 

Hilaira  glacialis  Thorell.  (PI.  IX,  fig.  3.) 

From  St.  Paul  Island,  May  19 ;  I  give  a  figure  of  the  male  palpus. 

Lepthyphantes  sp. 

One  female  St.  George  Island,  August  4. 


Ixodes  arcticus,  a  tick  parasitic  upon  the  fur  seal,  was  described  from  the  Pribilof 
Islands  by  Herbert  Osborn  (The  Fur  Seals  and  Pur-Seal  Islands  of  the  North  Pacific 
Ocean,  Part  3,  pp.  553-554,  1899). 

Opinions  differ  as  to  the  limits  of  the  spider  family  Linyphiidae,  and  Dr.  R.  V.  Cham- 
berlin  has  published  a  paper  on  the  "  Linyphiidae  of  St.  Paul  Island,  Alaska  "  ( Journ. 
New  York  Ent.  Soc,  vol.  29,  no.  1,  pp.  35-43,  pis.  3—4,  March,  1921),  in  which  he  records 
11  species.     Five  of  these  are  described  as  new,  in  addition  to  3  new  genera. 

These  spiders  are  herewith  listed : 


Cornicularia  clavicornis  Emerton. 
Erigone  sibirica  Kulczynski. 
Oedothorax  septentrionalis  Kulczynski. 
Oedothorax  nesides  Chamberlin. 
Aigola  Chamberlin. 
Aigola  pauliana  Chamberlin. 
Aigola  tuoerella  Chamberlin. 


Anitsia  Cliamberlin. 
Anitsia  abjecta  Chamberlin. 
Arctilaira  Chamberlin. 
Arctilaira  bellans  Chamberlin. 
Tmeticus  armatus  Banks. 
Microneta  heathi  Chamberlin. 
Bathyphantes  brevipes  Emerton. 


— W.  L.  M. 


[For  Plate  IX  (Arachnida)  and  explanation  of  plate,  see  page  158.] 


CHILOPODA. 

By  Ralph  V.  Chamberlin,  Museum  of  Comparative  Zoology, 
Harvard  University. 

No  diplopod  has  been  found  dn  the  Pribilof  Islands  and  it  is  not 
likely  that  any  occurs  there  naturally.  Four  species  of  chilopods, 
however,  are  natives  of  the  islands,  two  of  these  being  apparently 
abundant.  One  of  these,  the  geophiloid  Linotaenia,  is  widespread  in 
North  America  from  middle  latitudes  in  the  United  States  north- 
ward through  Canada  to  Alaska  and  the  adjacent  islands,  and,  if  not 
identical  with,  is  certainly  extremely  close  to,  the  widespread  Eura- 
sian species  Linotaenia  attenuatus  (Leach).  The  northern  specimens 
present  certain  differences  from  southern  forms  in  both  hemispheres 
as  pointed  out  below.  The  lithobiid  is  apparently  Asiatic  in  origin 
and  does  not  conform  generically  to  any  truly  North  American  genus 
as  conceived  by  the  writer.  The  other  two  species,  both  geophilomor- 
phous  forms,  appear  to  be  much  less  common.  Pachymerium  ferru- 
gineum  is  a  form  widespread  throughout  the  Eurasian  and  North 
American  regions,  having  a  range  very  similar  to  that  of  the  Lino- 
taenia ;  but  the  fourth  species,  the  Escaryus,  is  thus  far  known  from 
only  a  single  specimen  taken  on  St.  Paul  Island. 

Lithobiomorpha. 

Family  LITHOBIIDAE. 
Ezembius,  gen.  nov. 

Body  conspicuously  narrowed  cephalad ;  the  first  dorsal  plate  nar- 
rower than  the  head  but  wider  than  the  second  plate. 

Head  with  marginal  interruptions.  Antennae  short;  articles  nor- 
mally twenty. 

Ocelli  in  several  series;  single  ocellus  little  if  any  larger. 

Prosternal  teeth  2-f-2  or  3+3;  sinus  V-shaped;  ectal  spines  bristle- 
like. 

Coxal  pores  circular;  3,  3,  3,  3,  to  6,  7,  7,  6  in  number. 

None  of  the  posterior  coxae  laterally  armed;  but  two  or  more 
pairs  commonly  dorsally  armed.  Anal  legs  each  with  two  tarsal 
claws.  Ventral  spines  of  anal  legs  normally  0,  1,  3,  2,  0;  dorsal  1,  0, 
3,  1,  0.  Dorsal  spines  of  penult  legs  1,  0,  3,  1,  1.  Tibiae  of  all  legs 
excepting  the  anal  dorsally  armed,  commonly  the  tibae  of  the  first 
two  pairs  bearing  a  single  spine,  the  others  caudad  to  the  antepenult 
pair  bearing  two  spines. 

Tarsi  in  fully  grown  specimen  more  or  less  clearly  divided. 
240 


1923.]  CHILOPODS  OF  THE  PEIBTLOF  ISLANDS.  241 

Neither  anal  nor  penult  legs  of  male  with  a  definitely  developed 
process  or  lobe. 

Claw  of  female  gonopods  short,  trilobed,  the  lobes  normally  short 
and  inconspicuous  or  the  lobes  obsolete  and  the  claw  entire  or  sub- 
entire;  basal  spines  2-{-2,  slender. 

Small  and  medium  sized  species. 

Genotype. — Lithobius  stejnegeri  Bollman. 

Among  other  species  apparently  congeneric  with  this  type  form 
is  a  group  of  Siberian  species  embracing,  e.  g.,  the  following  de- 
scribed by  Stuxberg:  ostiacorum,  princeps,  sulcipes  (see  below),  and 
scrobiculatus. 

Ezembius  stejnegeri  (Bollman). 

Lithobius  stejnegeri  Bollman,  Bull.  U.  S.  Nat.  Mus.,  46,  p.  199,  1893. 
Lithobius  sulcipes  Bollman,  loc.  cit,  p.  199,  1893. 
Monotarsobius  arcticus  Attems,  Arkiv.  for  Zool.,  V.  no.  3,  p.  19,  1909. 
Lithobius  (Archililhobius)  haasei  Attems,  1909,  op.  cit.,  p.  22,  1909. 
Lithobius  sulcipes  Chamberlin,  Canad.  Ent.,  p.  260,  1911. 

Of  this  species  there  are  in  the  collection  sent  me  for  study  by  the 
Bureau  of  Biological  Survey  eight  specimens  from  St.  Paul  Island, 
four  of  these  being  adult  males,  two  adult  females,  and  two  young 
females  in  the  immaturus  stage,  apparently  differing  by  one  moult. 
There  are  also  five  adults  from  St.  George  Island  (collected  Aug. 
4,  1914,  by  E.  A.  Preble).  I  have  also  studied  several  specimens  of 
this  form  secured  on  St.  Paul  Island  in  1910  by  Prof.  Harold  Heath. 
In  addition  to  the  specimens  from  the  Pribilof  Islands,  I  have  ex- 
amined and  compared  material  from  Bering  Island,  the  type  locality, 
Copper  Island,  Popof  Island,  and  other  islands  of  the  region.  More 
recently  I  have  received  from  the  Bureau  several  additional  speci- 
mens collected  in  the  group  by  G.  Dallas  Hanna. 

After  a  careful  study  I  am  unable  to  detect  more  than  one  species 
in  the  material  from  these  localities,  and  am  convinced  that  the 
sulcipes  of  Bollman,  certainly  the  M  onotarsobius  arcticus  of  Attems, 
and  the  Lithobius  (Archilithobius)  haasei  of  Attems,  all  described 
from  Bering  Island,  are  one  and  the  same  as  E.  stejnegeri,  which  in 
turn  may  prove  to  be  identical  with  L,  sulcipes  Stuxberg  (1875),  and 
likely  with  the  much  earlier  L.  sibiricus  of  Gerstfeldt  (1858),  later 
fixed  by  the  description  of  Haase  (1880).  In  the  present  note,  how- 
ever, I  have  given  above  only  the  synonymy  of  forms  described  or 
recorded  from  the  islands  in  or  adjacent  to  Bering  Sea,  and  reserve 
the  question  of  the  Siberian  species  until  more  abundant  material 
from  that  region  can  be  studied.  The  M.  arcticus  of  Attems  is  sep- 
arated from  stejnegeri  for  the  single  stated  reason  that  the  ventral 
spines  of  the  anal  legs  of  the  latter  species  are  given  as  1,  3,  2,  6  in- 
stead of  1,  3,  2,  0,  as  in  articus.     The  6  in  the  first  formula,  as  might 


242  NOBTH   AMERICAN   FAUNA.  1923.] 

have  been  surmised,  is  clearly  a  misprint  for  0,  the  types  of  stejnegeri 
all  having  the  formula  for  anal  legs  1,  3,  2,  0,  so  that  there  remains 
no  evident  ground  whatsoever  for  maintaining  arcticus.  Attems  re- 
fers his  specimens  to  Monotarsobius ;  but  the  tarsi  in  the  larger 
specimens  of  the  species  are  quite  clearly  biarticulate,  though  in 
smaller  individuals  the  division  in  the  anterior  tarsi  may  not  be  dis- 
tinct or  may  be  but  partial.  Lithobius  (Archilithobius)  haasei 
Attems  was  apparently  based  upon  larger  specimens  of  the  same 
species.  For  this  form  the  prosternal  teeth  are  given  *as  3-J-3,  a 
number  occurring  quite  commonly  in  larger  individuals  from  all  the 
localities  above  mentioned,  while  the  number  may  be  3  on  one  side 
and  2  on  the  other.  The  extra  teeth  appear  on  the  border  of  the 
median  sinus  and  are  at  first  smaller  than  the  others.  The  claw  of 
the  female  gonopods  is  typically  tripartite;  but  the  lobes  are  short 
and  often  are  nearly  or  quite  obliterated  as  such,  leaving  the  claw 
subentire  or  entire. 

Geophilomorpha. 

Family  LINOTAENIIDAB. 

Linotaenia  chionophila  (Wood). 

?  Geophilus  acuminatus  Leach,  Trans.  Linn.  Soc.  Lond.,  XI,  p.  386,  1814. 
Strigamia  chionophila  Wood,  Journ.  Phil.  Acad.,  V,  p.  50,  1862. 
Scolioplanes  acuminatus  Attems,  Archiv.  for  Zool.,  V,  no.  3,  p.  25,  1909. 
Linotaenia  chionophila  Chamberlin,  Canad.  Ent.,  p.  260,  1911. 

Of  this  species  I  have  examined  22  females  and  29  males  collected 
on  St.  Paul  Island  in  1910  by  Prof.  Harold  Heath.  Specimens  from 
Bering  Island  studied  by  Attems  are  by  him  recorded  as  Scolioplanes 
acuminatus  (Leach),  a  species  well  known  in  Europe.  A  specimen 
from  the  same  island  is  listed  by  Bollman  as  L.  chionophila  (Wood). 
Specimens  from  Popof  and  Kadiak  Islands  and  from  Sitka  and 
Lower  Inlet  are  likewise  referable  to  Wood's  species.  There  is  no 
room  for  doubt  that  this  northern  form  is  the  typical  chionophila, 
of  which  the  type  specimen,  a  female,  was  taken  at  Fort  Simpson  on 
the  Red  River  of  the  North  [Fort  Simpson  is  on  Mackenzie  River, 
near  Lat.  62°  E.  A.  P.].  If  Graf  Attems  is  right  in  his  identification 
of  the  specimens  from  Bering  Island,  as  there  is  not  much  room  to 
doubt,  then  the  European  form  is  one  and  the  same  as  the  North 
American,  and  attenuatus  of  Leach  must  replace  chionophila  of 
Wood  as  the  specific  name.  In  view  of  the  different  mode  in  number 
of  pairs  of  legs  and  a  few  other  minor  points,  however,  I  believe  it 
as  well  to  keep  Wood's  name  for  the  present  and  until  the  forms  have 
been  more  intensively  studied  as  to  variation  and  distribution,  espe- 
cially since  such  difficulty  is  often  met  in  separating  closely  allied 
species  of  Linotaenia. 


1923.]  CHILOPODS   OF   THE   PRIBILOF   ISLANDS.  243 

Of  the  22  females  from  St.  Paul  Island  16  have  45  pairs  of  legs 
and  6  have  43.  Wood's  type  has  43  pairs.  Of  the  29  males  from  St. 
Paul  Island  27  have  43  pairs  of  legs,  one  has  45  pairs,  and  one  has 
but  41  pairs.  Attems  states  that  among  his  specimens  from  Bering 
Island  one  male  had  41  pairs  and  one  45  pairs,  the  others  having  43. 
Thus  it  would  seem  that  the  numbers  of  pairs  of  legs  in  the  male  is. 
almost  constant  at  43,  individuals  with  41  or  45  being  occasional; 
while  in  the  female  the  modal  number  is  45,  variation  to  43  being 
frequent. 

In  the  case  of  the  European  specimens  of  L.  attenuatus,  the  num- 
ber of  pairs  of  legs  is  nearly  always  smaller.  In  Austria-Hungary 
Latzel  found  among  60  specimens  studied  that  all  the  males  had  39 
pairs  while  in  the  females  the  number  was  either  41  or  43.  Meinert 
gives  the  number  of  pairs  of  legs  as  constantly  41,  but  gives  the 
number  in  the  female  as  41  or  47  pairs,  one  specimen  having  the  latter 
number.  In  "Die  Myriopoden  Stiermarks"  Attems  states  that  all 
the  males  studied  by  him  from  that  country  had  39  pairs  of  legs,  ex- 
cepting one  which  had  41,  while  all  the  females  had  41.  The  same 
author,  however,  found  among  specimens  from  Transylvania  four 
males  with  37,  two  males  with  35,  and  five  with  but  33  pairs  of  legs ; 
and  of  females  nine  with  39,  one  with  37,  and  seven  with  33  pairs  of 
legs.  It  will  be  noted  then  that  in  European  specimens  of  L.  attenu- 
atus the  most  usual  number  of  pairs  of  legs  in  the  male  is  39  and  that 
in  some  regions  this  number  seems  to  be  nearly  fixed;  but  that  in 
other  places  variation  below  this  number  may  be  frequent  or  the  rule. 
Similarly  the  modal  number  for  the  female  is  41,  but  variation  is 
more  frequent  than  in  the  male,  the  number  sometimes  being  43,  or, 
in  sections  where  the  variation  in  the  number  in  the  male  in  the  minus 
direction  is  frequent,  falling  to  37  and  even  to  33. 

In  the  United  States  specimens  of  L.  chionophila  differ  from  the 
northern  specimens  and  agree  with  the  European  L.  attenuatus  in 
having  the  number  of  pairs  of  legs  in  the  male  most  frequently  39.  In 
the  female  the  number  varies  from  41  to  37.  Of  22  females  from 
Ithaca,  N.  Y.,  I  find  six  to  have  41  pairs,  eight  to  have  39,  and  eight 
to  have  37  pairs. 

It  may  be  noted  that  the  number  of  coxal  pores  in  specimens  of 
attenuatus  from  Europe  and  specimens  of  chionophila  from  the 
United  States  averages  considerably  higher  than  in  the  specimens 
from  the  Pribilof  and  other  islands  of  the  region.  Of  the  22  females 
from  St.  Paul  Island,  eight  have  on  each  side  six  pores,  five  have  five, 
five  have  seven,  three  have  eight,  and  one  has  nine;  of  the  males, 
sixteen  have  six  pores,  ten  have  five,  two  have  seven,  and  one  has 
eight.     The  mode  is  thus  six  pores  on  each  side. 


244  NORTH  AMERICAN  FAUNA.  [No.  46 

Family  GEOPHILIDAE. 
Pachymerium  ferrugineum  (C.  L.  Koch). 

Geophilus  ferrugineus  C.  L.  Koch,  Deutschl.  Crust.,  Myr.,  u.  Arachn.,  Hft. 

3,  Tab.  2,  1835. 
Pachymerium  ferrugineum  C.  L.  Koch,  System  der  Myr.,  p.  187,  1847. 
Mecistocephalus  ferrugineus  Newport,  Cat.  Myr.  Brit.  Mus.,  p.  81,  1856. 
Mecistocephalus  foveatus  McNeill.  Proc.  U.  S.  Nat.  Mus.,  vol.  10,  p.  333,  1887. 
Geophilus   atterwatus  Cook    (nee  Say),  Proc.   U.   S.  Nat.  Mus.,  vol.  18, 

p.  59,  1895. 

One  specimen  of  this  form  was  secured  on  St.  Paul  Island  by  the 
Harriman  Expedition.  It  is  probably  not  uncommon  on  the  Pribilof 
Islands,  since  it  is  common  both  on  the  Asiatic  and  North  American 
mainlands.  It  is  a  widespread  and  abundant  species  both  in  Europe 
and  Asia  and  in  North  America. 

Family  SCHENDYLIDAE. 
Escaryus  albus  Cook. 

Esaryus  albus  Cook,  Harriman  Alaska  Expedition,  vol.  8,  p.  77,  1904. 

The  type  of  this  species,  taken  on  St.  Paul  Island,  is  the  only  speci- 
men thus  far  known. 


INDEX. 


Abietinaria,  42,  50,   52,  55, 

58. 
ablusus,  Citellus,  113. 
Acanthis    hornemanni    exili- 
pes,  9,  90. 

linaria  .(inaria,  91. 
Acanthopneusie  Dorealis,  14. 
acaulis,  Silene,  7. 
Accipitridae,  81. 
acetosella,  Rumex,  8. 
Achillea  millefolium,  8. 
Acmaea,  55. 
Actodromas,  73. 
aculeata,  Corallina,  59. 
acuminata,  Pisobia,   11,  70. 
Adams,  H.  P.,  work  by,  4. 
adamsii,  Gavia,  17. 
Aegialites    californicus,    64, 
65,  70,  74,  90,  93. 

debilis,  100. 
Aegialitis  semipalmata,  126. 
Aegobuccinum       oregonense, 

28. 
Aethia  cristatella,  6,  10,  21. 

pusilla,  6,  10,  23,  86. 
affinis,   Pontoporeia,   50,  52, 

53. 
Aithyia  ferina,  11,  48. 
alascanus,     Callorhinus,     5, 
108. 

Haliaeetus,  9,  82. 
alascensis,  Budytes,  14. 

Calcarius,  6,  9,  10,  12, 
94,  126. 

Nannus,  6,  10,  12,  98. 

Troglodytes,  98,  121. 

Vulpes,  105. 
Alaska  longspur,  94. 

Peninsula'',  8,  9. 

Peninsula  muskrat,  113. 

red  fox,  105. 

wren,  98. 
Alaskan  Province,  7. 
alaudinus,  Passerculus,  96. 
alba,  Pagophila,  31,  126. 
albatross,  Short-tailed,  38. 
albatrus,  Diomedea,  38. 
albeola,  Charitonetta,  49. 
albifrons,  Anser,  60. 
albus,  Lagopus,  9. 
Alcidae,  17. 


[To  Part  I  only,  pages  1-128.] 

Aleutian  Faunal  District,  7, 
9. 

Islands,  8,  9. 

Peninsula,  8. 

rosy  finch,  88. 

sandpiper,  66. 

Savannah  sparrow,  95. 

song  sparrow,  96. 
Aleutican  subregion,  7. 
Algae,  28,  36,  42,  50,  52,  53, 

54,  57,    61,    69,    73. 
Coralline,     59. 
Filamentous,  50. 
Marine,  56. 

Plume,   42,    50,    52,   56, 
59. 
algida,  Phippsia,  8. 
aliciae,  Hylocichla,  100. 
Allorchestes,  53,  55. 
malleolus,  26,  55. 
ochotensis,  27. 
Alopecurus  alpinus,  8. 

howellii  var.    merriami, 
8. 
Alopex  pribilofensis,  5,  103. 
alpestriformis,      Amblyteles, 

79. 
alpinum,  Cerastium,  7. 

Phleum,   28. 
alpinus,  Alopecurus,  8. 
Alsine,   91. 

borealis,    90. 
alternifolium,     Chrysosplen- 

ium,   8. 
Amara,    64,   70,   72,    79,    81, 
93. 

brunnipennis,     30,     70, 

72,  75,  81. 
glacialis,    72. 
Amauronematus       whitneyi, 

30. 
Amblyteles,    30,   70,    72,   93. 

alpestriformis,  79. 
American  goldeneye,  48. 
gyrfalcon,  82. 
merganser,  42. 
rough-legged    hawk,    81. 
americana,  Glaucionetta,  48. 
americanus,  Mergus,  42. 
Ammothea  pribilofensis,   52, 

56. 
Amphipods,   15,   23,   25,   26, 
37,  38,  48,  50,  51,  52,  53, 

55,  58,  59,  69,  71,  72,  73, 
74,  76,  80,  100. 


Amphiroa   cretacea,   59. 
Amphissa  reticulata,   55. 
Amphithoe  rubicata,  55. 
amplexifolius,  Streptopus,  8. 
ampullacea,  Volutharpa,  55. 
Anas     platyrhyncha     platy- 

rhyncha,  10,  43. 
Anatidae,  42. 
anatum,  Falco,  83. 
Ancient  murrelet,  24. 
angustata,  Glyceria,  8. 
angustior,   Ilybius,   64. 
annulatus,  Borborus,  37. 
Anonyx  nugax,  27,  50,  52,  55. 
Anser     albifrons     albifrons, 

60. 
anserina,  Potentilla,  8. 
Anthomyiidae,  64. 
Anthus  spinoletta  japonicus, 
12,  98. 

spinoletta  rubescens,  9, 
11,  97. 
antiquus,    Synthlibo  ram- 

phus,  24. 
apetala,  Lychnis,  7,  91. 
Aptera,  90. 

apterus,  Tachinus,  70. 
Archibuteo    lagopus    sanctl- 

johannis,  9,  81. 
Arctic  tern,  37. 
zone,  6,  8. 
arctica,   Calamogrostis,  8. 
Salix,  8. 

Saxicava,     50,     52,    55, 
57,  59. 
arcticum,      Chrysanthemum, 

8. 
arcticus,  Rubus,  8. 
Arctogrostis  latifolia,   8. 
Arctonetta  fischeri,  53. 
Arctophila  effusa,  8. 
Arenaria,  93. 

interpres   interpres,   11, 

14,  80. 
peploides,  7. 
Aristonetta  valisineria,  47. 
Armeria  vulgaris,  8. 
Arquatella  maritima   couesi, 
11,  66. 
maritima  ptilocnemis,  6, 

10,  12,  67. 
ptilocnemis,  124. 
arra,  TTria,  6,  10,  28. 
Arries,  28. 

245 


246 

Ascidians,  54,  56. 
Asio  flammeus,  84. 
asplenoldes,  Ptilota,   42,  52, 

56,  59. 
assimilis,  Neopleustes,  55. 
Astyris,  57. 
Atheta,  64,  70. 
atkana,    Littorina,    44,    50, 

52,  55,  57. 
Atylus,  55. 

Auk,  White-breasted,  20. 
Auklet,  Crested,  21. 

Least,  23,  86. 

Paroquet,  20. 
auritus,  Colymbus,  11,  16. 


bachmani,    Haematopus,   81. 
Baillie  brushkie,  20. 
Baird  sandpiper,  72. 
bairdi,  Pisobia,  72. 
bairdii,  Berardius,  117,  127. 
Balaeaa  mysticetus,  5,  115. 
Balaenidae,  115. 
Balaenoptera  davidsoni,  116. 

velifera,  116. 
Balaenopteridae,  116. 
Balanus  balanus,  50,  52,  55. 

rosteratus,  58. 
barbatus,   Erignathus,    112. 
Barnacles,    15,    27,    49,    50, 

51,  52,  53,  55,  58. 
Barn   swallow,   97. 
barrovianus,  Larus,  33,  34. 
Bartsch,  Paul,  work  by,  15. 
Bathymedon  obtusifrons,  50, 

53. 
Bathymetis,  70. 
Batracblum      tricbophyllum, 

46. 
baueri,  Limosa,  11,  14,  74. 
Beach  beetles,  70,  74,  90,  93, 

100. 
Bean,  T.   H.,   work  by,   2. 
Bear,  Polar,  103. 
Bearded  seal,  6,  112. 
Beetles,  74. 

Beach,  70,  74,  90,  93, 
100. 

Chrysomelid,   79. 

Click,  71,  81. 

Diving,  64,  71,  72. 

Ground,  30,  37,  72,  73, 
74,  76,  79,  81,  90,  93, 
95,  98,  100,  101. 

Leaf,  70,  71,  79,  90,  93, 
95. 

Moss,  70,  71. 

Rove,  64,  70,  71,  72,  75, 
79,  81,  100,  101. 

Sexton,  100. 

Water,  37. 
bellidifolia,  Cardamine,  7. 
benthami,    Selinum,   8. 


NORTH  AMERICAN  FAUNA. 


[No.  46 


Berardius  bairdii,  117,  127. 
beringianum,    Chrypsospleni- 

um,  8,  90,  95. 
bicolor,  Iridoprocne,  97,  125. 
biglumis,  Juncus,  8. 
bipinnata,  Pterosiphonia,  52. 
Birds,  breeding  distribution, 
6. 

food  habits,  14. 

list,  10. 
bistorta,  Polygonum,   8. 
Bivalves,   49,  53. 
Blackbird,  Rusty,  87. 
Black  brant,  60. 

oyster-catcher,  81. 
Blowflies,  64,  95. 
Bluebell,  90. 
Blue  fox,  103. 

mussels,  34,  50,  53,  54, 
57,   69,  73. 
Bog  bean,  65. 
Boltenia  ovifera,  56,  58. 
Borboridae,  100. 
Borborous  annulatus,  37. 
Borcotrophon,   50. 

multicostatus,  55,  57. 
borealis,  Acantbopneuste,  14. 

Alsine,  90. 

Numenius,  78. 
Bowhead  whale,  5,  115. 
Brachyramphus  marmoratus, 

25. 
Brambling,  87. 
brandtii,  Schizoplax,  57. 
Brant,  Black,   60. 
Branta    canadensis    hutchin- 
si,  60. 

canadensis   minima,   11, 
60. 

nigricans,  60. 
brevipennis,    Liparocephalus, 

100. 
brevipes,  Heteractitis,  77. 

Heteroscelus,  12,  76. 

Lithodes,   26. 
brevirostris,  Rissa,  6,  10,  32. 
Bristle- thighed  curlew,  78. 
Brook   saxifrage,  90,  95. 
brunnipennis,  Amara,  30,  70, 

72,  75,  81. 
Bryozoa,  30,  50,  52,  53,  56, 

58,  59. 
Buccinum  fischerianum,  57. 

plectrum,  55. 
Budytes     flavus     alascensis, 

14. 
Bufflehead,  49. 
Bunchberry,  88. 
Bunting,   Snow,  83,  91. 
Buttercup,  90,   93. 
Buturlin  hawk  owl,  84. 
Byrrhus  fasciatus,  70,  71. 


Cackling  goose,  60. 
Caddis  flies,  37,  44,  47,  50, 
64,  75,  76. 

larvae,  80. 
Calamogrostis  arctica,  8. 

purpurascens,  8. 
Calcarius     lapponicus     alas- 
censis,   6,    9,    10,    12,    94, 
126. 
California  murre,  26. 
californica,  Una,  6,   10,   26. 
californicus,    Aegialites,    64, 

65,  70,  74,  90,  93. 
Calliopiidae,  17,  55. 
Calliphora  vomitoria,  69,  71, 

76,  88,  95,  101. 
Callorhinus      alascanus,     5, 

108. 
Campanula,  90. 

uniflora,  8. 
Campoplex,  72. 
canadensis,  Megalornis,  62. 
canagica,    Philacte,    11,    14, 

61. 
Canldae,  103. 

cannaban,  Chaetomorpha,  61. 
Canooskie,   21. 
Canvasback,  47. 
capitata,  Valeriana,  8. 
Caprella  kincaidi,  50,  55. 
Carabidae,  95. 
Carabus     truncaticollis,     30, 

37,  71,  76,  79,  81. 
Cardamine  bellidifolia,  7. 

pratensis,  7. 

umbellata,  8. 
Cardium    ciliatum,    53,    55, 

57. 
Carex,  65,  66,  90,  93,  95. 

membranopacta,  8. 

pribylovensis,  8. 

rariflora,   8. 

saxatilis,  8. 

vulgaris,  8. 
carolinense,  Nettion,  10,  45. 
carolinus,  Euphagus,  87. 
Caterpillars,  100. 
Cellepora  surcularis,  56,  58. 
Cepphus    columba,    10,    11, 

25. 
Caramiaceae,  50,  56. 
Cerastium  alpinum,  7. 
Cerithiopsis,  57. 
Cervidae,  114. 
Chaetomorpha  cannaban,  61. 

melagonium,  50. 
chamaeinorus,  Rubus,  8. 
Charadriidae,  78. 
Charadrius    dominicus     ful- 
vus,  78. 

fulvus,  78. 

semipalmatus,  79. 
Charitonetta    albeola,    49. 
Chase,   Agnes,   work  by,   15. 


1923.] 


INDEX. 


247 


Chaulelasmus  streperus,  44. 
cheiragonus,  Telmessus,  58. 
Chen  hyperborea  hyperborea, 

59. 
Chickee,  34. 
Chickweed,  60,  90,  91. 

Water,  65,  90,  93,  95. 
Chironesimus       multiarticu- 

latus,  17. 
Chironomidae,    47,    64,    71, 

74,   SI. 
Chitin,  73. 

Chiton,  50,  52,  55,  57. 
marinoreus,  55,  57. 
submarmorea,  34. 
Choochkie,  23,  84. 
Chrysanthemum  arcticum,  8. 
Chrysomela    subsulcata,    70, 

71,   79,   90,   93,   94,   95. 
Chrysomelid  beetles,  79. 
Chrysosplenium     alternifoli- 
um,    8. 

beringlanum,  8,  90,  95. 
ciliatum,    Cardium,    53,    55, 

57. 
Cinclus  mexicanus  unicolor, 

9. 
cinereus,    Machaeroplax,    57. 
Cinquefoil,  90,  93,  95. 
cirrhata,   Lunda,   6,    10,    17. 
Citellus      plesius       ablusus, 

113. 
elangula,    Glaucionetta,    11, 

48.  , 

elangula  hyemalis,  6,  10,  49. 
Clark,   A.   H.,   work   by,   15. 
clausa,  Natica,  55,  57. 
Click  beetle,  71,  81. 
Cliff    swallow,    97. 
clypeata,  Spatula,  46. 
coarctatus,  Hyas,  58. 
Coccothraustes     coccothrau- 

stes  japonicus,  12,  87. 
Cochlearia   officinalis,   7. 
Codfish,    23. 
Coelopa,  eximia,  69. 

frigida,    76. 
Coelopleurum     gmelini,     90, 

91. 
Colaptes  auratus  luteus,  86, 

121. 
Cole,  Leon  J.,  work  by,   15. 
Collembola,    90. 
columba,    Cepphus,    10,    11, 

25. 
columbianus,  Olor,  62. 
Colymbidae,    16. 
Colymbus    auritus,     11,    16. 

holboellii,    16. 
Comarum  palustre,  8. 
comosa,    Saxifraga,    8. 
Compsothlypidae,  97. 
Coprotheres    pomarinus,    29. 
Coptis    trifolia,    7. 
Corallina   aculeata,   59. 
Coralline  algae,   59. 


Cormorant,  Red-faced,  41. 

Violet-green,  40,  41. 
corniculata,    Fratercula,    6, 

10,    19. 
Cornus   suecica,   88. 
coronata,   Zonotrichia,,   9. 
Corvidae,    SO. 
Corvus  corax  principalis,  9, 

86. 
Cottidae,    27. 

Coues,   Elliott,   work  by,  2. 
couesi,  Arquatella,  11,  66. 
Crab,   34,   36,   54. 

Hermit,  15,  26,  34,  42, 

50,  52,  55,  58. 
Mud,    58. 

Spider,   26,    32,   42,   55, 
58. 
Crane,  Little  brown,  62. 
Crane-flies,    30,    64,    69,    71, 
72,    76,    79,    81,    86,    90, 
93,  95. 
crassirostris,  Tringa,  67. 
crecca,   Nettion,   11. 
Crested  auklet,  21. 
cretacea,  Amphiroa,  59. 
crinita,  Scatophaga,  64,   69, 

76,    86,    95. 
Crisia,    56. 

cristatella,  Aethia,  6,  10,  21. 
cristatellus,      Simorhynchus, 

121. 
Crossbill,  White-winged,   88. 
Crowberry,    65,    70,    79,    80, 

90,  93,  95. 
Crowfoot,  Water,  46. 
Crustacea,  17,  21,  32,  33,  41. 
Crustaceans,  41,  42,  54,  57, 

64,  69. 
Cryptoglaux    funerea    fune- 
rea,  84,  125. 

funerea  magna,   12,  84, 

125. 
funerea  richardsoni,  84. 
Cuckoo,  Kamchatkan,  86. 
Cuculidae,  86. 
Cuculus  canorus  telephonus, 

12,  86. 
Curlew,  Bristle-thighed,  78. 
Eskimo,  78. 
Hudsonian,  77. 
cyclopylla,  Salix,  8. 
Cynomyia  hirta,  30,  95. 

D. 

Dafila    acuta    tzitzihoa,    11, 

47. 
Dall,  W.  H.,  work  by,  15. 
dalli,  Pagurus,  55. 
damacensis,  Pisobia,  73. 

Tringa,  126. 
Daphniidae,  64,  69. 
dasythrix,     Scatophaga,    37, 

64. 


Davidson      lesser      rorqual, 

116. 
davidsoni,  B  a  1  a  e  n  o  ptera, 

116. 
debilis,  Aegialites,  100. 
Delphinidae,  117. 
Dendroica    aestiva    rubigin- 

osa,  9. 
Dermaturus   mantii,   26,   34, 

41,  50,  52,  55,  58. 
Dilsea  integra,  61. 
Diomedea  albatrus,  38. 
Diomedeidae,  38. 
Dipper,  9. 
Diptera,  44,  64,   69,  72,   74, 

76,  80,  86,  88,  93. 
divergens,    Odobenus,    5,    6, 

106. 
Diving  beetles,  64,  71,  72. 
dixoni,  Oidemia,  11,  59. 
Dowitcher,  Long-billed,  66. 
Draba  hirta,  7. 

wahlenbergii,  7. 
drobachiensis,    Strongylocen- 

trus,  55,  58. 
Duck,    American    goldeneye, 
48. 

American        merganser, 

42. 
Bufllehead,  49. 
Canvasback,  47. 
European  goldeneye,  48. 
European  pochard,  48. 
European  teal,  45. 
European  widgeon,  44. 
Falcated  teal,  45. 
Gadwall,   44. 
Green-winged  teal,  45. 
Harlequin,   51. 
King  eider,  56. 
Mallard,  43. 
Oldsquaw,   49. 
Pacific  eider,  54. 
Pacific       white  -  winged 

scoter,  59. 
Pintail,  47. 
Red-breasted  merganser, 

42. 
Scaup,  47. 
Shoveler,  46. 
Spectacled  eider,  53. 
Steller  eider,  52. 
Tufted,  47. 
Duck  hawk,  83. 
Dungflies,  64,  69,  74,  76,  81, 

86,  93,  95,  100,  101. 
Dupontia  psilosantha,  8. 

EL 

Eagle,  Northern  bald,  82. 
Echinarachnius    parma,    50, 

52,  54,  55,  58. 
edeta,   Spania,  93. 

edulis,   Mytilus,   34,   50,   52, 

53,  54,  57,  59,  61,  69,  73. 


248 


NORTH  AMERICAN   FAUNA. 


[No.  46 


edwardsii,  Eutrema,  7. 
Eels,  Rock,  41,  42. 
effusa,  Arctophila,  8. 
Eider,   King,  56. 

Pacific,  54. 

Spectacled,  53. 

Steller,  52. 
Elephantidae,   119. 
Elephas  primigenius,   119. 
Elliott,  Henry  W.,  work  by, 

2. 
Elymus  mollis,  8. 

villosissimus,  8,   56. 
emarginata,  Potentilla,  8. 
Emperor  goose,  61. 
Empetrum  nigrum,  8,  31,  65, 

70,   79,   80,  90,   93,   95. 
Epilobium  spicatum,  8. 
Ereunetes  pusillus,  74. 
Erignathus  barbatus  nauti- 

cus,  6,  112. 
Erimacrus  isenbeckii,  54. 
Eriophorum  polystachyon,  8. 

vaginatum,  8. 
Eristalinae,  46. 
erytbrogastra,    Hirundo,   97. 
escbscboltzii,  Ranunculus,  7. 
Eskimo  curlew,  78. 
Eumetopias  jubata,  5,  107. 
Eunetta  falcata,  11,  45. 
Eupbagus  carolinus,  87. 
Euphrasia  officinalis,  8. 
European  goldeneye,   48. 

jacksnipe,  65. 

pochard,  48. 

widgeon,  44. 
Eutrema  edwardsii,  7. 
Evermann,  B.  W.,  work  by, 

3. 
exilipes,  Acanthis,  9,  90. 
eximia,  Coelopa,  69. 

Primula,    8. 

F. 

falcata,  Eunetta,  11,  45. 

Falcated   teal,   45. 

Falco     peregrinus     anatum, 

83. 

peregrinus  pealei,  9,  S3, 
sacer,  83. 
Falcon,  Peale,  83. 
Falconidae,    82. 
fasciata,  Phoca,  6,  111. 
fasciatus,    Byrrhus,    70,    71. 

Pholis,    41. 
Faunal  District,  Aleutian,  7, 

9. 
ferina,  Aithyia,  11,  48. 
Festuca  ovina,  8. 

ovina  violacea,  28. 
Fiber     zibethicus     zalophus, 

113. 
Filamentous  alga,  50. 
filiformis,  Potamogeton,   46. 


Finch,  Aleutian  rosy,  88. 

Rosy,  83. 
fischeri,  Arctonetta,  53. 
fischerianum,  Buccinum,  57. 
flammeus,  Asio,  84. 
flavipes,  Totanus,  75. 
Fleas,    Sea,    24. 

Water,    64,    69. 
Flesh    flies,    30. 
Flicker,   Northern,  86. 
Flower   flies,   71. 
fontana,  Montia,  66,  90,  93, 

95.  , 

Food  habits,  birds,  14. 
Foraminifera,     52,    53,     54, 

5S. 
Forked-tailed  petrel,   40. 
Fox,    5. 

Alaska  red,   105. 
Blue,    103. 
Pribilaf  arctic,  103. 
Fratercula     corniculata,     6, 

10,    19. 
fraterna,  Leria,  37,  69. 
frigida,  Coelopa,  76. 

Petasites,  8. 
Fringilla  montifringilla,   12, 

87. 
Fringillidae,   87. 
Fucellia,   69,  95. 
fulicarius,    Phalaropus,    11, 

63. 
Fuligula  fullgula,   11,   47. 
Pulix    marila,    47. 
Fulmar,  Rodgers,  38. 
Fulmarus      glacialis      rodg- 

ersi,   6,   10,   38. 
fulva,  Pluvialis,  11,   12,  14, 

78. 
fulvus,  Charadrius,  78. 
funerea,      Cryptoglaux,      84, 

125. 
fur,  Ptinus,  71. 
Fur  seal,  Pribilof,  5. 
furcata,    Oceanodroina,    40, 
126. 
Scatophaga,  69. 
fuscum,    Olophrum,    64,    70, 

100. 

G. 
Gabrielson,  Ira  N.,  work  by, 

15. 
Gadwall,  44. 
,gallinula,   Limnocryptes,  11, 

65. 
Gamasidae,    100. 
Gambel  sparrow,   96. 
gambeli,  Zonotrichia,  96. 
Gammaridae,  17. 
Gammarus,   69,   73,    76. 

pribilofensis,  50,  52,  55, 
58,   64,   69. 
Gavia  adamsii,  17. 

vlridigularis,   17,   122. 
Gaviidae,  17. 
Gelis,  70,  71. 


Gentian,  95. 
glacialis,  Amara,  72. 
glareola,    Rhyacophilus,    12, 

75. 
glaucescens,    Larus,    6,    10, 

34. 
Glaucionetta  clangula  ainer- 
icana,    48. 

clangula    clangula,    11, 
48. 
Glaucous  gull,  33. 
Glaucous-winged  gull,  34. 
Glyceria  angustata,   8. 
gmelini,     Coelopleurum,    90, 

91. 
godwit,  Pacific,  74. 
Goldeneye,  American,  48. 

European,   48. 
Goose,  Cackling,  60. 

Emperor,  61. 

Hutchins,  60. 

Snow,  59. 

White-fronted,  60. 
Goverooskie,  32. 
gracilis,  Oregonia,  42,  55. 

Tringa,  67,  124. 
grandis,  Llttorina,  55,  57. 

Nesodraba,  8. 
Grass,  Rye,  56. 
Gray-cheeked  thrush,  100. 
Greater  yellow-legs,  75. 
Grebe,  Holboell,  16. 

Horned,  16. 
grebnitzkii,       Hapalogaster, 

26,  41,  42,  50,  55,  58. 
Green-throated  loon,  17. 
Green-winged  teal,  45. 
griseigena,  Podiceps,  16. 
griseonucha,   Leucosticte,   6, 

9,  10,  83,  88. 
groenlandica,    Spirontocaris, 

41,  42. 
gronlandica,  Littorina,  57. 
grosbeak,  Kamchatkan  pine, 

88. 
Ground  beetles,   30,   72,   73, 
74,  76,  79,  81,  90,  93,  95, 
98,  100,  101. 
Gruidae,  62. 
Guillemot,  Pigeon,  25. 
Gull,  Glaucous,  33. 

Glaucous-winged,  34. 

Ivory,  31. 

Pacific  kittiwake,  31. 

Point  Barrow,  34. 

Red  -  legged  kittiwake, 
32. 

Rosy,  36. 

Sabine,  36. 

Slaty-backed,  36. 
guttata,  Hylocichla,  9. 
Gymnocanthus        pistilliger, 

42. 
Gynandra  stelleri,  8. 
Gyrfalcons,  89. 

American,  82. 


1923.] 


I2STDEX. 


249 


H. 

Hadrotes,  64,  72. 

Haematopodidae,  81. 

Haematopus  bachmani,  81. 

Hahn,   Walter   L.,   work  by, 
3. 

Haliaeetus  leucoc  ephalus 
alascanus,  9,  82. 

Haloconcha  minor,  55. 
reflexa,  55. 

Hanna,  G.  Dallas,  work  by, 
3,  4,  15. 

hannai,  Pisidium,  69. 

Hapalogaster  grebnitzkii,  26, 
41,  42,  50,  55,  58. 

Harbor  porpoise,  118. 
seal,  Pribilof,  5. 

Harlequin  duck,  51. 

Hawfinch,  Japanese,  87. 

Hawk,    American    rough-leg- 
ged, 81. 
Duck,  83. 

Helgoland,  birds  of,  13,  14. 

helicina,  Margarites,  50,  52, 
55,  57. 

Hemiptera,  72,  88,  100. 

Hermit  crab,  15,  26,  34,  42, 
50,  52,  55,  58. 

Heteractitis  brevipes,  77. 
incanus,  75. 

Heteroscelus   incanus   brevi- 
pes, 12,  76. 

incanus,  11,  14,  75. 

hieracifolia,  Saxifraga,  8. 

Hierochloa,  pauciflora,  8. 

Hierofalco   rusticolus   sacer, 
11,  82. 

Hippomedon,  50. 

Hippuris  vulgaris,  8,  6G,  70, 
73. 

hirculus,  Saxifraga,  8. 

hirta,  Cynomyia,  30,  95. 
Draba,   7. 

Hirundinidae,  97. 

Hirundo  rustica  erythrogas- 
tra,  97. 

Histrionicus  histrionicus  pa- 
cificus,  10,  51. 

Hoary  redpoll,  90. 

Holarctic  region,  7. 

Holboell  grebe,  16. 

holboellii,   Colymbus,   16. 

Holm,  Theodor,  work  by,  7. 

Horned  grebe,  16. 
puffin,  19. 

House  mouse,  85,  86,  113. 

hudsonia,  Pica,  9. 

Hudsonian  curlew,  77. 
Zone,  birds,  9. 

hudsonicus,   Numenius,   77. 

humifusa,  Stellaria,  7. 

Hutchins  goose,  60. 

hutchinsi,   Branta,  60. 


Hyale,  27,  52,  58. 
Hyas  coarctatus,  58. 
Hydroids,  15,  25,  28,  34,  42, 

52,  58,  59,  73. 
Hydrophorus   innotatus,    37, 

94. 
Hydroporus,  71,  72. 
hyemalis,    Clangula,    6,    10, 
49. 

Junco,  96. 
Hylocichla  guttata   guttata, 
9. 

minima  aliciae,  100. 
ustulata  swainsoni,   9. 
Hymenoptera,  46,  70,  72,  79, 

93,  98,  100. 
hyperborea,  Chen,  59. 
hyperboreus,    Larus,    6,    10, 
33,  125. 

Plectrophenax,  93. 
Quedius,  72. 
Ranunculus,  7. 
Hyphoraia   subnebulosa,    30. 
Hypnoidus  musculus,  70,  71. 

I. 

Icteridae,    87. 

Idothea   ochotensis,   26,    27, 

32,  55,  69. 
Idotheidae,  55. 
Ilybius  angustior,  64. 
incanus,   Heteractitis,   75. 

Heteroscelus,  11,  14,  75. 
inermis,  Pontogeneia,  25,  64. 
innotatus,  Hydrophorus,  37, 

94. 
inquinatus,  Lophalophus,  70, 

71,  72,  73,  75,  95,  98,  101. 
insignis,   Melospiza,  9. 

Trichotropis,  52,  55,  57. 
integra,  Dilsea,  61. 
intermedia,  Mya,  57. 
interpres,  Arenaria,   11,   14, 

80. 
Irbisia  sericans,  70,  71,  72, 

98. 
Iridoprocne  bicolor,  97,  125. 
Ischyrocerus,    55. 
isenbeckii,  Erimacrus,  54. 
islandica,    Koenigia,    S. 
islandicus,   Pecten,    55,   57. 
Isopods,  26,  27,  32,  52,  55, 

58,  69. 
Ivory    gull,    31. 


Jacksnipe,    European,    65. 
Jaeger,    Parasitic,    29. 

Pomarine,   29. 
Japanese  hawfinch,  87. 

pipit,   98. 

swift,   86. 


japonicus,  Anthus,  12,  98. 

Coccothraustes,   12,   87. 

Micropus,    125. 
Jassa,    27. 

marmorata,    52. 
Jellyfish,   Brown-eyed,   39. 
jubata,  Eunietopias,   5,   107. 
Judd,  S.  D.,  work  by,  15. 
Junco     hyemalis     hyemalis, 

96. 
Junco,   Slate-colored,  96. 
Juncus,    90,    93. 

biglumis,   8. 

K. 

Kamchatkan  cuckoo,   86. 

pine  grossbeak,  S8. 

sea   eagle,    82. 
kamtschathensis,      Pinicola, 

12,    88,    126. 
Kelp,  15,  23,  26,  50,  52,  54, 
58. 

flies,  69,   76,   95. 
Killer  whale,  5,  117. 
kincaidi,  Caprella,  50,  55. 
King    eider,    56. 
Kittiwake,    Pacific,    31. 

Red-legged,    32. 

Western,  31. 
Koenigia   islandica,    8. 
kotzebuei,  Parnassia,  8. 


laeta,   Stellaria.   7. 
Lagopus    lagopus    albus,    9. 

rupestris  nelsoni,  9. 
lanata,  Pedicularis,  S. 
langsdorfii,     Viola,    70,    72, 

95,  98. 
Large  finback  whale,  116. 
Laridae,   31. 
Larus  barrovianus,  33. 

glaucescens,    6,    10,    34. 

hyperboreus,   125. 

hyperboreus    barro- 
vianus, 34. 

hyperboreus     hyperbor- 
eus, 6,  10,  33. 

schistisagus,  10,  36. 

warnecki,    32. 
Latax  lutris,  105. 

lutris  lutris,   6. 
laticeps,  Megalocottus,  41. 
latifolia,  Arctogrostis,  8. 
Leaf  beetle,   70,  71,   79,  90, 

93,  95. 
Least  auklet,  23,  86. 

sandpiper,  73. 
Lemming,   Pribilof,   5,   112. 
Lemmus  nigripes,  5,  70,  112. 
lepida,  Tachycineta,  97. 
Leptasterias,  55,  58. 
Leria  fraterna,  37,  69. 

leucostoma,   69. 


250 


NORTH  AMERICAN   FAUNA. 


[No.  46 


Lesser  yellow-legs,  75. 
Lettuce,  Sea,  41,  42,  45. 
leucoptera,  Loxia,  88,  125. 
Leucosticte    griseonucha,    6, 

9,  10,  83,  88. 
leucostoma,  Leria,  69. 
Life  zones,  5. 
Ligusticum  scoticum,  90. 
lima,  Nucella,  55,  57. 
Limmershin,  98. 
Limnocryptes    gallinula,    11, 

65. 
Limnodromus   griseus   scolo- 

paceus,  66. 
Limosa  lapponica  baueri,  11, 

14,  74. 
Limpets,  52,  54. 
linaria,  Acanthis,  91. 
linnaei,   Sagina,   7. 
Liparocephalus    brevipennis, 

100. 
Lithodes  brevipes,  26. 
Lithodidae,  32. 
Little  brown  crane,  62. 
Littorina,  57,  69,  76. 

atkana,  44,  50,  52,  55, 
57. 

grandis,  55,  57. 

grolandica,  57. 

sitchana,  69,  75,  76. 
Lloydia  serotina,   8. 
lobatus,  Lobipes,  6,  10,  65. 
Long-billed  dowitcher,  66. 
Long-footed   fly,   94. 
longspur,  Alaska,  94. 
Long-toed  stint,  73. 
Loon,  Green-throated,   17. 

Yellow-billed,    17. 
Lophalophus  inquinatus,  70, 
71,    72,    73,    75,    95,    98, 
101. 
Lora,  53. 

Lousewort,   65,  93. 
Loxia  leucoptera,  125. 

leucoptera      leucoptera, 
88. 
Lucas,  F.  A.,  work  by,  3. 
Lunda  cirrhata,  6,  10,  17. 
lunifrons,  Petrochelidon,  97. 
Lupine,  50,  70,  72. 
Lupinus     nootkatensis,     50, 

70,   72. 
luridum,  Trinoton,  45. 
luteus,   Colaptes,  86,   121. 
lutris,  Latax,  6,  105. 
Lutz,  J.   E.,  work  by,   2. 
Lychnis  apetala,  7,  91. 
Lyrosoma  opaca,  70,   100. 

M. 

Maehaeroplax  cinereus,  57. 
Machetes  pugnax,  77. 


Macoun,  James  M.,  work  by, 

7. 
macoun  ii,  Papaver,  8,  95. 
Polygonum,  8. 
Primula,  8. 
macrocephalus,        Physeter, 

116. 
Macrorhamphus   g.    scolopa- 

ceus,  66. 
maculata,  Pisobia,  11,  71. 
magna,  Cryptoglaux,  12,  84, 

125.  / 
Mallard,  43. 
malleolus,    Allorchestes,    26, 

55. 
Mammals,  distribution,  5. 

list,  102. 
Mammoth,  119. 
mandtii,  Dermaturus,  26,  34, 

41,  50,  52,  55,  58. 
Marbled  murrelet,  25. 
Mareca  penelope,  11,  44. 
Margarites  helicina,  50,   52, 

55,  57. 
Marila  marila,  47. 

valisineria,  47. 
marila,  Fulix,  47. 
Marine  algae,   56. 

worms,   15,   17,   26,   34, 

37,  48,  53,  65,  69,  70, 

74,  75. 

maritimus,   Thalarctos,   103. 

marmorata,  Jassa,  52. 

marmoratus,    Brachyramph- 

us,   25. 
marmoreus,  Chiton,  55,  57. 
Marsh,  Millard  C,  work  by, 

3. 
McAfee,  W.  L.,  work  by,  14, 

15. 
McKay  snow  bunting,  93. 
Mcgalocottus  laticeps,  41. 
Megalornis  canadensis,  62. 
Megaptera  versabilis,  116. 
melagonium,   Chaetomorpha, 

50. 
Melanella,  53. 
melanoleucus,    Totanus,    75, 

123,  127. 
Melospiza    melodia    insignis, 

9. 

melodia  sanaka,  96. 
membranopacta,    Carex,   8. 
Menipea  pribilofi,  56,  58. 
Menyanthes  trifoliata,  65. 
Merganser,  American,  42. 

Red-breasted,  42. 
Mergus     merganser     ameri- 

canus,  42. 
serrator,  42. 
merriami,  Alopecurus,  8. 
Metcalf,  F.  P.,  work  by,  15. 
Metopa,  27,  52,  55. 
Metopidae,   17. 


Micropodidae,  86. 
Micropus    pacificus,    12,    86, 

125. 
Midges,  64,  71,  74. 

Ruby-red,   81. 
millefolium,  Achillea,  8. 
minima,  Branta,  11,  60. 
minor,  Haloconcha,  55. 

Pyrola,  8. 
minuta,     Orchomenella,    50, 

52,  58. 
minutilla,  Pisobia,  73. 
Mites,  72. 
Modiolaria  verrucosa,  49,  50, 

52,  53,  54. 
Modiolus  modiolus,  69. 
mollis,  Elymus,  8. 
Mollusca,  49. 

Mollusks,  27,  34,  46,  51,  52, 

54,  57,  69. 

Montia  fontana,  66,  90,  93, 

95. 
montifringilla,  Fringilla,  12, 

87. 
Moss  beetles,  70,  71. 
Motacillidae,  97. 
Moth,  Brown,  30. 

Yellow,  30. 
Mouse,  House,  85,  86,  113. 
Mud  crab,  58. 
multiarticulatus,     Chironesi- 

mus,  17. 
multieostatus,  Borcotrophon, 

55,  57. 
Muridae,  112. 
murre,  California,  26. 

Pallas,  28.   . 
Murrelet,  Ancient,  24. 

Marbled,  25. 
Mus  musculus,  86,  113. 
Muscidae,  64. 
Musculus,  50,  57. 
musculus,  Hypnoidus,  70,  71. 
Muskrat,   Alaska   Peninsula, 

113. 
Mussels,  15,  52,  54. 

Blue,  34,  50,  53,  54,  57, 
69,  73. 
Mustelidae,  105. 
Mya  intermedia,  57. 
Myodes  obensis,  112. 
Myoxocephalus,  41,  70. 

polyacanthoceph- 
alus,  41. 
Myriozoum  subgracile,  58. 
mysticetus,  Balaena,  5,  115. 
Mytilus    edulis,    34,    50,    52, 

53,  54,  57,  59,  61,  69,  73. 

N. 

Nannus  troglodytes  alascen- 

sis,  6,  10,  12,  98. 
Natica  clausa,  55.  57. 
operculina,  57. 


1923.] 


ENDEX. 


251 


nauticus,  Erignathus,  6,  112 
Nebalia,    55. 

nebulosa,   Synidotea,  58. 
Nelson,  E.   W.,  work  by,  2 
nelsoni,    Lagopus,    9. 
Neopleustee,    55. 
assimilis,   55. 
Nereid    worms,    50,    52,    58. 
Nereidae,  37,  48,  53,  65,  70, 

74,    75. 
Nereis,    17. 

Neritina    sitchana,    69. 
Nesodraba  grandis,  8. 
Nettion  carolinense,   10,  45. 

crecca,    11. 
Neverita  recluziana,  53,  55, 

57. 
nigricans,  Branta,  60. 
nigripes,     Lemmus,     5,     70, 

112. 
nigrum,    Empetrum,    8,    31, 
65,  70,  79,  80,  90,  93,  95. 
nivalis,    Plectrophenax,    91. 

Sagina,    7. 
nootkatensis,    Lupinus,     50, 

70,    72. 
North   Sea,   13. 
Northern  bald  eagle,  82. 
flicker,    86. 
phalarope,    65. 
raven,    86. 
norvegicus,  Rattus,  113. 
Norway   rat,    113. 
Nucella  lima,   55,   57. 
nucellatus,  Trichotropis,  5T. 
nugax,   Anonyx,   27,   50,   52, 

55. 
Numenius  borealis,   78. 
hudsonicus,    77. 
tahitiensis,   12,  78. 
Nushagak    ground    squirrel, 

113. 
Nutting,  C.  C,  work  by,  15. 
Nyctea  nyctea,   11,   85. 

O. 

obensis,  Myodes,   112. 
oblivia,    Parathemisto,    23. 
obtusifrons,        Bathymedon, 

50,     53. 
Oceanodroma     furcata,     40, 

126. 
ochotensis,  Allorchestes,  27. 
Idothea,  26,  27,  32,  55, 

69. 
Odobenidae,    106. 
Odobenus    divergens,    5,    6, 

106. 
Oenanthe    oenanthe    oenan- 

the,    101. 
oenanthe,  Saxicola,  101,  123, 

127. 
officinalis,   Cochlearia,  7. 
Euphrasia,    8. 


Oidemia  deglandi  dixoni,  11, 

59. 
Old-squaw    duck,    49,    85. 
Olophrum    fuscum,    64,    70, 

100. 
Olor  columbianus,  62. 
opaca,  Lyrosoma,   70,   100. 
operculina,  Natica,  57. 
Orchestia,   69. 
Orchomenella     minuta,     50, 

52,  58. 
Orcinus  rectipinna,  5,  117. 
oregonense,      Aegobuccinum, 

28. 
Oregonia  gracilis,  42,  55. 
Osburn,  R.  C,  work  by,  15. 
Osgood,    Wilfred    H.,    work 

by,  4. 
Ostracoda,  46,  55. 
Otariidae,  107. 
Otoliths,  16. 
Otter  Island,  desc,  1. 
Otter,  Sea,  6,  105. 
Ousel,  Water,  9. 
ovifera,  Boltenia,  56,  58. 
ovina,    Festuca,    8. 
Owl,  Buturlin  hawk,  84. 
Richardson,  84. 
Short-eared,  84. 
Snowy,  85. 
Oxyria  reniformis,  8. 
Oyster-catcher,  Black,  81. 


Pacific  beaked  whale,  117. 

eider,  54. 

godwit,  74. 

golden  plover,  78. 

humpbacked  whale,  116. 

kittiwake,    31. 

turnstone,  80. 

walrus,  106. 

white-winged  scoter,  59. 
pacificus,    Histrionicus,     10, 
51. 

Micropus,  12,  86. 
Tagophila  alba,  31,  126. 
Pagurus,  58. 

dalli,  55. 

trigonocheirus,  55,  58. 

undosus,  41. 
Pallas  murre,  28. 
Palmer,    William,    work   by, 

2,  3,  15. 
palustre,  Comarum,  8. 
panoplus,  Pleustes,  55. 
Papaver  macounii,  8,  95. 

radicatum,  7. 
paradisaea,   Sterna,  11,  37. 
Parasitic  jaeger,  29. 

wasps,    65,    71,    76,    79, 
100. 


parasiticus,  Stercorarius,  29. 
Parathemisto  oblivia,  23. 
Parker,     George     H.,     work 

by,   4. 
parma,    Echinarachnius,    50, 

52,  54,  55,  58. 
Parnassia  kotzebuei,  8. 
Paroquet  auklet,  20. 
Parsnip,  Wild,  90. 
Passerculus      sandwichensis 
alaudinus,  96. 
sandwichensis  sandwich- 
ensis, 9,  95. 
Passerella  iliaca  sinuosa,  96. 
iliaca  unalaschensis,  9, 
96. 
Patrobus    septentrionis,    70, 

73. 
pauciflora,   Hierochloa,   8. 
Peale  falcon,  83. 
pealei,  Falco,  9,  83. 
Pecten  islandicus,  55,  57. 
pectinata,    Ptilota,    52,    56, 

59. 
Pectorial  sandpiper,  71. 
Pedicularis,   65,   93. 
lanata,  8. 
sudetica,  8. 
pelagicus,  Thallasoaetus,  12, 

82. 
Felidna  alpina  sakhalina,  72. 
penelope,  Mareca,   11,  44. 
peploides,  Arenaria,  7. 
Peta sites  frigida,  8. 
Petrel,  Fork-tailed,  40. 
Petrochelidon  lunifrons  lunl- 

frons,  97. 
Phalacrocoracidae,  40. 
Phalacrocorax   pelagicus   ro- 
bustus,  6,  10,  40. 
urile,  6,  10,  41. 
Phalarope,  Northern,  65. 

Red,  63. 
Phalaropodidae,  63. 
Phalaropus     fulicarius,     11, 

63. 
Phaleris    psittacula,    6,    10, 

20. 
Philacte    canagica,    11,    14, 

61. 
Philomachus  pugnax,  12,  77. 
Phippsia  algida,  8. 
Phleum  alpinum,  28. 
Phoca  fasciata,  6,  111. 

richardii     pribilofensis, 
5,  111. 
Phocaena  phocaena,  118. 
Phocidae,  111. 
Pholis,  42. 

fasciatus,  41. 
Phygadeuon,  70,   100. 
Physeter  macrocephalus,  116. 
Physeteridae,    116. 
Pica  pica  hudsonia,  9. 


252 


NORTH  AMERICAN  FAUNA. 


[No.  46 


Picidae,  86. 
Pigeon  guillemot,  25. 
pileolata,  Wilsonia,  9,  97. 
Pileolated  warbler,  97. 
Pilsbry,  H.  A.,  work  by,  15. 
Pine  siskin,  91. 
pinguidineus,     Pterostichus, 

30. 
Pinicola    enucleator    kamt- 
schathensis,  126. 

leucura     kamtschathen- 
sis,  12,  88. 
Pink    family,    91. 
Pintail,    47. 
pinus,    Spinus,   91. 
Pipit,    97. 

Japanese,   98. 
Pisidium  hannai,  69. 
Pisobia    acuminata,    11,    70. 
bairdi,    72. 
damacensis,    73. 
maculata,    11,    71. 
minutilla,  73. 
subminuta,  12,   73. 
pistilliger,      Gymnocanthus, 

42. 
Planesticus  migratorius  pro- 

pinquus,  101. 
Plant   bug,    71. 
Plants,    distribution,    7. 
platyrhyncha,  Anas,  10,  43. 
Plectrophenax    hyperboreus, 
93. 

nivalis  nivalis,  91. 
nivalis  townsendi,  6,  9, 
10,  12,  92,  126. 
plectrum,   Buccinum,   55. 
Plesignathus,  65,   100. 
Pleustes  panoplus,  55. 
Plicifusus        spitzbergensis, 

55,  57. 

Plover,  Pacific  golden,  78. 

Semipalmated,  79. 
Plume-algae,  15,  42,  50,  52, 

56,  59. 

Pluvialis  dominica  fulva,  11, 

12,   14,   78. 
Pochard,  European,  48. 
Podiceps  griseigena,  16. 
Point  Barrow  gull,   34. 
Polar  bear,  103. 
polaris,     Spirontocaris,     41, 

42,  55,  58. 
pollicaris,   Rissa,    6,   10,   12, 

31,  121. 
polyacanthocephalus,    Myox- 

ocephalus,  41. 
Polygonum  bistorta,  8. 
macounii,    8. 
viviparum,    8. 
Polynesian    tattler,    76. 
Polyrhembia,    71,    76. 
polystachyon,      Eriophorum, 


Polysticta    stelleri,    11,    52. 
Pomarine  jaeger,  29. 
pomarinus,  Coprotheres,  29. 

Stercorarius,  29. 
Pondweed,  48. 
Pontogeneia,  17,  38. 
inermis,   25,    64. 
Pontogeneidae,  16,  17. 
Pontoporeia   afiinis,    50,    52, 

53. 
Poochka,  90. 
Poppy,  95.   / 
Porpoise,  Harbor,  118. 
Potamogeton,  48. 

filiformis,  46. 
Potentilla,  90,  93,  95. 
anserina,  8. 
emarginata,  8. 
pratensis,   Cardamine,  7. 
Preble,  E.  A.,  work  by,  4. 
Prentiss,  D.  W.,  work  by,  3. 
Pribilof  arctic  fox,  103. 
Bird  Reservation,  2. 
fur  seal,  5,  108. 
harbor  seal,  5,  111. 
Islands,  desc,  1. 
lemming,  112. 
sandpiper,   67. 
shrew,  86,  102. 
snow  bunting,  92. 
pribilofensis,  Alopex,  5,  103. 
Ammothea,  52,  56. 
Gammarus,    50,    52,   55, 

58,  64,  69. 
Phoca,   5,  111. 
Sorex,    5,    84,    86,    102, 
125. 
pribilofi,  Menipea,   56,   58. 
pribylovensis,  Carex,  8. 
primigenius,  Elephas,  119. 
Primula  eximia,  8. 

macounii,  8. 
principalis,  Corvus,  9,  86. 
Procellariidae,  38. 
propinquus,  Planesticus,  101. 
psilosantha,    Dupontia,    8. 
psittacula,    Phaleris,    6,    10, 

20. 
Pterosiphonia  bipinnata,  52. 
Pterostichus,  30,  37,  64,  69, 
71,  72,  73,  75,  79,  81, 
90,  93,  98,  101. 
pinguidineus,  30. 
ptilocnemis,     Arquatella,     6, 
10,  12,  67,  124. 
Tringa,   67,    122. 
Ptilota,  56,  59. 

asplenoldes,  42,  52,  56, 

59. 
pectinata,  52,  56,  59. 
Ptinus  fur,  71. 
Puffin,  Horned,  19. 

Tufted,  17. 
Puffinus  tenuirostris,  39. 


pugnax,  Machetes,  77. 

Philomachus,   12.   77. 
purpurascens,       Calamagros- 

tis,  8. 
pusilla,    Aethla,    6,    10,    23, 

86. 
pusillus,  Ereunetes,  74. 
Pycnogonid,  26,  52,  58. 
pygmaeus,  Ranunculus,  7. 
Pyrola  minor,  8. 

Q. 

Quedius  hyperboreus,  72. 


Radicatum,  Papaver,  7. 
Rangifer  tarandus,  114. 
Ranunculus,  90,  93. 
eschscholtzii,  7. 
hyperboreus,  7. 
pygmaeus,  7. 
reptans,  7. 
tricophyllus,  7.  , 
rariflora,  Carex,  8. 
Rat,  Norway,  113. 
Rathbun,  Mary  J.,  work  by, 

15. 
Rattus  norvegicus,  113. 
Raven,  Northern,  86. 
recluziana,  Neverita,  53,  55, 

57. 
rectipinna,  Orcinus,  5,  117. 
Red  phalarope,  63. 
Red-backed  sandpiper,  73. 
Red-breasted  merganser,  42. 
Red-faced  cormorant,  41. 
Red-legged  kittiwake,  32. 
Redpoll,  91. 

Hoary,   90. 
reflexa,  Haloconcha,  55. 
Reindeer,  114. 
reniformis,  Oxyria,  8. 
reptans,  Ranunculus,  7. 
reticulata,  Amphissa,  55. 

Salix,  8. 
Rhodostethia  rosea,  36. 
Rhyacophilus    glareola,    12, 

75. 
Ribbon  seal,  6,  111. 
Richardson  owl,  84. 
•ichardsoni,  Cryptoglaux,  84. 
Riparia  riparia,  9. 
Rissa  brevlrostris,  6,  10,  32. 
tridactyla   pollicaris,   6, 
10,  12,  31,  121. 
Robin,  Western,  101. 
robustus,    Phalacrocorax,    6, 

10,  40. 
Rochefortia,  53,  57. 
Rock  eels,  41,  42. 
Rodgers  fulmar,  38. 
rodgersi,  Fulmarus,  6,  10,  38. 


1923.] 


INDEX. 


253 


Rohwer,  S.  A.,  work  by,  15. 
Rorqual,      Davidson      lesser, 

116. 
rosea,  Rhodostethia,  36. 
rostratus,  Balanus,  58. 
Rosy  finch,  83. 

gull,  36. 
rotundata,  Salix,  8. 
Rove  beetles,  64,  70,  71,  72, 

75,  79,  81,  93,  100,  101. 
rubescens,  Antbus,  9,  11,  97. 
rubiginosa,  Dcndroica.  9. 
rubricata,  Amphithoe,  55. 
Rubus  arcticus,  8. 

cliamaemorus,  8. 
Ruby-red  midge,  81. 
Ruff,  77. 

Rumex  acetosella,  8. 
Rusty  blackbird,  87. 
Rye  grass,  56. 


S. 


Sabine  gull.  36. 
sabinii,  Xema,   11,   36. 
sacer,   Falco,  83, 

Hierofalco,   11,  82. 
Sagina  linnaei,  7. 

nivalis,  7. 
Saint  George  Island,  descr., 

1. 
Saint  Matthew  Islands,  9. 
Saint  Paul   Island,  desc,   1. 
sakhalina,    Pelidna,   73. 
Salix  arctica,  8. 

cyclopylla,  8. 

reticulata,  8. 

rotundata.  8. 
sanaka,  Melospiza,  96. 
sancti-johannis,    Archibuteo, 

9,  81. 
Sand  dollars,  50.  52,  53,  55. 
Sandpiper,   Aleutian,   66. 

Baird,  72. 

Least,  73. 

Pectoral.    71. 

Pribilof.  67. 

Red-backed,  73. 

Semipalmated,  74. 

Sharp-tailed,  70. 

Wood,  75. 
sandwichensis,    Fasserculus, 

91,   95. 
Sandwort,   93. 
Sawfly,  30. 
saxatilis,  Carex,  8. 
Saxicava  arctica,  50,  52,  55, 

57,  59. 
Saxicola  oenanthe  oenanthe, 
101. 

oenanthe,    123,   127. 
Saxifraga  hieracifolia,  8. 

hirculus,  8. 

Btellaris  comosa,  8. 


Saxifrage,  60. 

Brook,  90,  95. 
Scatophaga,  64,  74,  100. 

crinita,   64,   69,   76,  86, 
95. 

dasythrix,  37,  64. 

furcata,   69. 
Scatophagidae,    64,    69,    81, 

93,    101. 
Scaup   duck,  47. 
schistisagus,   Larus,    10,   36. 
Schizoplax,  52. 

brandtii,  57. 
Schmitt,  W.  L.,  work  by,  15. 
Sciuridae,  113. 
scolopaceus,      Limnodromus, 
66. 

Macrorhamphus,  66. 
Scolopacidae,   65. 
scoter,  Pacific  white-winged, 

59. 
scotieum,   Ligusticum,   90. 
Sculpin,    15,  27,   38,  41,   42, 

54,  70. 
Sea  eagle,  Kamchatkan,   82. 

fleas,    24. 

lettuce,   41,  42,   45,   61. 

otter,    6,    1Q5. 

parsley,  90. 

quail,    121. 

shag,  40. 

slug,   34,  50,   54. 

snail,   15,  52,   54,  57. 

spider,  26,  52,  58. 

squirt,   15,  58. 

urchin,    15,    36,   49,    50, 
52,  53,  54,  55,  57,  59. 
Seal,    Bearded,    6,    112. 

Pribilof  harbor,  111. 

Ribbon,    6,    111. 
Sealion,    Steller,    5,    107. 
Sealion    Rock,   desc,    1. 
Sedge,  66,  90,  93,  95. 
Seleucus,   70. 
Selinum    benthami,    8. 
semipalmata,  Aegialitis,  126. 
Semipalmated   plover,    79. 

sandpiper,    74. 
semipalmatus,       Charadrius, 

79. 
septentrionis,   Patrobus,    70, 

73. 
sericans,  Irbisia,  70,  71,  72, 

98. 
serotina,    Lloydia,    8. 
serpyllifolia,   Veronica,   8. 
serrator,    Mergus,    42. 
Sertularella,    55,    58. 
Setchell,  W.  A.,  work  by,  15. 
Sexton-beetle,     100. 
Sharp-tailed    sandpiper,    70. 
Shearwater,     Slender-billed, 
39. 


Shoemaker,       Clarence       L., 

work    by,    15. 
Short-eared    owl,    84. 
Short-tailed  albatross,  38. 
Shoveler,    46. 
Shrew,    5,    84. 

Pribilof,    86,    102. 
Shrimps,    24,    41,   42. 
Shumagin    fox    sparrow,    96. 
Silene    acaulis,    7. 
Silphidae,    70. 
Simorhynchus      cristatellus, 

121. 
sinuosa,   Passerella,   96. 
Siskin,  Pine,  91. 
sitchana,    Littorina,    69,    75, 
76. 

Neritina,    69. 
Slate-eolored  juneo,  96. 
Slaty-backed    gull,    36. 
Slender-billed       shearwater, 

39. 
Slug,   Sea,  34,  50,  54. 
Snails,   Sea,   15,   52,   54,   57. 
Sr.Jpe    fly,    93. 
Snow  buntings,  83,  91. 
McKay,  93. 
Pribilof,   92. 
goose,  59. 
Snowy  owl,  85. 
Socarnea  vahli,  23. 
Soldier-flies,  64. 
Somateria      spectabilis,      11, 
56. 
v-nigra,   11,   54. 
Sorex    pribilofensis,     5,     84, 

86,  102,  125. 
Soricidae,  102. 
Spania  edeta,  93. 
Sparrow,     Aleutian     Savan- 
nah, 95. 
Aleutian  song,  96. 
Gambel,  96. 
Shumagin  fox,  96. 
Valdez   fox,   96. 
Western    Savannah,    96. 
Spatula  clypoata,  46. 
spectabilis,     Somateria,     11,. 

56. 
Spectacled  eider,  53. 
Sperm   whale,   116. 
Sperry,     Charles     C,     work 

by,  15. 
spicatum,   Epilobium,  8. 
Spider,   Sea,  26,   52. 
Spider,  crab,  26,  32,  42,  55,. 

58. 
Spiders,  71,  72,  $3. 
Spinus  pinus  pinus,  91. 
Spirontocaris,  41. 

groenlandica,   41,   42. 
polaris,  41,  42,  55,  58. 
spitzbergensis,  Plicifusus,  55, 
57. 


108731°— 23- 


-17 


254 


NORTH   AMERICAN   FAUNA. 


[No.  46 


Squid,  29,  33,  30,  42. 
Squirrel,   Nushagak    ground, 

113. 
Staphylinidae,     70,    79,    93, 

100,  101. 
Star  fishes,  15,  54,  57. 
Stellaria  humifusa,  7. 
longipes  laeta,  7. 
Steller    eider,    52. 

sea  lion,  5,  107. 
stelleri,   Gynandra,  8. 

Folysticta,  11,  52. 
Stercorariidae,  29. 
Stercorarius  parasiticus,  29. 

pomarinus,  29. 
Sterna  paradisaea,  11,  37. 
Stiboscopus,  70. 
Stint,  Long-toed,  73. 
Stores  beetle,  71. 
Stratiomyidae,  64. 
streprus,    Chaulelasinus,    44. 
Streptopus  amplexifolius,  8. 
Strigidae,  84. 
Strongylocentrus      drobachl- 

ensis,   55,  58. 
subgracile,  Myriozouin,  58. 
submarmorea,  Chiton,  34. 
subminuta,  Pisobia,  12,  73. 
subnebulosa,  Hyphoraia,  30. 
subspicatum,  Trisetum,  8. 
subsulcata,    Chrysomela,   70, 

71,  79,  90,  93,  94,  95. 
sudetica,  Pedicularis,  8. 
suecica,  Cornus,  88. 
surcularis,  Cellepora,  56,  58. 
swainsoni,  Hylocichla,  9. 
Swallow,  Barn,  97. 
Cliff,  97. 
Tree,  97,  125. 
Violet-green,  97. 
Swan,  Whistling,  62. 
Swift,  Japanese,  86. 
Synidotea  nebulosa,  58. 
Synthliboramphus    antiquus, 

24. 
Syrphidae,  71. 

T. 

Tachinus,  70.  71,  75,  81,  93. 

apterus,  70. 
Tachycineta     thalassina     le- 

pida,  97. 
tahitiensis,     Numenius,     12, 

78. 
tarandus,   Rangifer,    114. 
Tattler,  Polynesian,  76. 

Wandering,  75. 
Teal,   Falcated,  45. 

Green-winged,  45. 
telephonus,  Cuculus,  12,  86. 
Telmessus  cheiragonus,  58. 
tenuirostris,  Puffinus,  39. 
Tern,  Arctic,  37. 
Thalarctos  maritimus,  103. 


Thallasoaetus  pelagicus,   12, 

82. 
Thrush,   Gray-cheeked,    100. 
Thuiaria,  56,  58. 
Thysanoessa,  33. 
Tipula,  30,  69,  72,  95. 
Tipulidae,    64t    69,    71,    72, 

79,  90,  93. 
Tmetonyx,  50. 
Totanus  fiavipes,  75. 

melanoleucus,    75,    123, 
127./ 
Townsend,   C.    H.,   work    by, 

3. 
townsendi,   Plectrophenax,  6, 

9,  10,  12,  92,  126. 
Tree  swallow,  97,  125. 
trichophyllum,     Batrachium, 

46. 
Trichoptera,  47. 
Trichotropis  insignia,  52,  55, 

57. 

nucellatus,  57. 
tricophyllus.  Ranunculus,  7. 
trifolia,  Coptis,  7. 
trifoliata,  Menyanthes,  65. 
trigonoeheirus,  Pagurus,  55, 

58. 
Tringa  crassirostris,  67. 
damacensis,   126. 
gracilis,  67,  124. 
ptilocnemis,  67,  122. 
Trinoton  luridum,  45. 
Trisetum   subspicatum,   8. 
Troglodytes    alascensis,    98, 

121. 
Troglodytidae,  98. 
True,   F.   W.,  work  by,  3. 
truncaticollis,    Carabus,    30, 

37,  71,  76,  79,  81. 
Tubicolous    worms,    50,    52, 

58. 
Tufted  duck,  47. 

puffin,  17. 
Turdidae,   100. 
Turner,  L.  M.,  work  by,  2. 
Turnstone,  Pacific,  80. 
Two-winged  flies,  37,  64,  69, 

80. 
tzitzihoa,    Dafila,    11,    47. 

U. 

Clekiah,  5. 

umbellata,    Cardamine,    8. 

unalaschensis,  Passerella,  9, 

9,  96. 
I'nalaska,  9. 
undosus,   Pagurus,   41. 
unicolor,  Cinclus,  9. 
uniflora,  Campanula,  8. 
Fnivalves,  26,  36,  38,  52,  53. 
Uria  lomvia  arra,  6,  10,  28. 
troille  californica,  6,  10, 
26. 


urile,   Phalacrocorax,  6,   10, 

41. 
Ursidae,  103. 


vaginatum,  Eriophorum,  8. 
vahli,   Socarnea,   23. 
Valdez  fox  sparrow,  96. 
Valeriana  capitata,  8. 
valisineria,   Aristonetta,    47. 

Marila,  47. 
velifera,    Balaenoptera,    116. 
Veniaminof,  work  by,  2. 
Veronica  serpyllifolia,  8. 
verrucosa,     Modiolaria,     49, 

50,  52,  53,  54. 
versabilis,  Megaptera,  116. 
villosissimus,  Elymus,  8,  56. 
Viola,   65. 

langsdorfii,   70,    72,    95, 
98. 
Tiolacea,  Festuca,  28. 
Violet,  70,  72,  95,  98. 
Violet-green    cormorant,    40, 
41. 

swallow,  97. 
viridigularis,  Gavia,  17,  122. 
viviparum,  Polygonum,  8. 
v-nigra,  Somateria,   11,  54. 
Volutharpa  ampullacea,  55. 
vomitoria,     Calliphora,     69, 

71,   76,  88,  95,   101. 
vulgaris,   Armeria,  8. 

Carex,  8. 

Hippuris,  8,  66,  70,  73. 
Vulpes  alascensis,  105. 

W. 

wahlenbergii,     Draba,     7. 
Walrus,   5. 

Pacific,  106. 
Walrus    Island,    desc,   1. 
Wandering  tattler,  75. 
Warbler,  Pileolated,  97. 
Warneck  (of  Russian  Navy), 

work    by,    2. 
warnecki,  Larus,  32. 
Wasp,  parasitic,  30,   65,  71, 

76,   79,   100. 
Water    chickweed,     66,     90, 
93,    95. 

crowfoot,    46. 
fleas,   64,   69. 
ousel,    9. 
shrimps,    24. 
Weevil,   70,    71,   72,   73,    75, 

90,    95,    98,    101. 
Western    black-legged    kitti- 
wake,    31. 
robin,     101. 

Savannah    sparrow,   96. 
Wetmore,    Alexander,    work 
by,   15. 


1923.] 


INDEX. 


255 


Whale,  Bowhead,  5,  115. 

Killer,   5,   117. 

Large  finback,  116. 

Pacific   beaked,   117. 

Pacifichumpbacked,  116. 

Sperm,  116. 
Wheatear,    101. 
Whistling  swan,  62. 
White-breasted  auk,  20. 
White-fronted   goose,   60. 
White-winged    crossbill,    88. 
Whitney,  Alvin  G.,  work  by, 

3. 
whitneyi,      Amauronematus, 
30. 


Widgeon,    European,    44. 
Wild   parsnip,   90. 
Wilsonia     pusilla     pileolata, 

9,   97. 
Wood    sandpiper,    75. 
Worms,  Marine,   15,   17,  26, 

34,  37,  48,  53,  65,  69,  70, 

74,    75. 

Nereid,  58. 

Tubicolous,    50,    52,   58. 
Wren,    Alaska,    9S. 

X. 

Xanthidae,    55,    58. 
Xema   sabinii,   11,   36. 


Yellow-billed  loon,   17. 
Yellow-legs,   Greater,  75. 
Lesser,    75. 

Z. 

zalophus,   Fiber,    113. 
Ziphiidae,  117. 
Zone,  Arctic,  6,  8. 

Hudsonian,  9. 
Zonotrichia    coronata,    9. 

gambeli,   96. 
Zoogeography,  8. 


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North  American  Fauna  No.  47,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  I. 


9-.   o 


±  ■* 


U.  S.  DEPARTMENT  OF  AGRICULTURE 

BUREAU  OF  BIOLOGICAL  SURVEY 


NORTH  AMERICAN  FAUNA 

ISTo.  47 

[Actual  date  of  publication,  August  21,  1924  ] 


REVISION  OF  THE  AMERICAN  PIKAS 


(Genus  OCHOTONA) 


By  Arthur  H.  Howell,  Assistant  Biologist 

Division  of  Biological  Investigations 

Bureau  of  Biological  Survev 


WASHINGTON 

GOVERNMENT    PRINTING    OFFICE 

1924 


&*€    ■  ,      /fiZ// 


LETTER  OF  TRANSMITTAL. 

United  States  Department  of  Agriculture, 

Bureau  of  Biological  Survey, 
Washington,  D.  C,  October  25, 1923. 

Sir:  I  have  the  honor  to  transmit  herewith,  with  the  recommendation  that  it  be 
published  as  No.  47  in  the  series  of  the  North  American  Fauna,  a  report  of  a  study, 
which  constitutes  a  revision,  of  the  American  forms  of  the  pikas,  or  "rock  conies" 
(genus  Ochotona),  by  Arthur  H.  Howell,  assistant  biologist  of  this  bureau.  The  report 
is  based  chiefly  on  material  in  the  collection  of  the  Biological  Survey,  gathered  dur- 
ing many  years  of  investigations  of  the  native  mammals  of  North  America.  Pikas 
inhabit  rocky  country,  mainly  in  mountainous  regions,  and  in  America  are  found  in 
such  situations  in  the  western  part,  ranging  from  Alaska  to  California  and  New  Mex- 
ico. The  publication  of  the  report  will  furnish  needful  information  to  mammalogists 
and  to  biological  students  generally  concerning  the  number  of  forms  and  the  habits, 
geographic  distribution,  and  relationships  of  these  interesting  mammals. 
Respectfully, 

W.  C.  Henderson, 

Acting  Chief  of  Burtau. 
Hon.  Henry  C.  Wallace, 

Secretary  of  Agriculture. 
n 


CONTENTS. 

Pag*. 

Introduction 1 

Distribution  and  habitat  of  pikas  in  America 1 

Habits 3 

Pelage  and  molt 5 

Material  examined 6 

Genus  Ochotona  Link 7 

History  and  nomenclature 8 

Subgenus  Pika  Lacepede 9 

List  of  American  species  and  subspecies,  with  type  localities 9 

Key  to  the  American  species 10 

Descriptions  of  American  species  and  subspecies 10 

Ochotona  princeps  (Richardson) 10 

Ochotona  collaris  (Nelson) 34 

Ochotona  schisticeps  (Merriam) 36 

Table  of  cranial  measurements 40 

Explanation  of  plates 52 

Index 57 

in 


ILLUSTRATIONS. 

PLATES. 

Pace. 
Plate     I.  Figgins  pika  (Ochotona  princeps  ftgginsi),  from  Irwin,  Colorado, 

Frontispiece. 
II.  Fig.  1. — Pika  in  rockslide,  Gunnison  county,  Colorado;  Fig.  2. — 

Haystack  of  pika,  Garfield  County,  Colorado 4 

III.  Skulls  (dorsal  views)  of  Ochotona  princeps  and  0.  collaris 52 

IV.  Skulls  (dorsal  views)  of  Ochotona  princeps  and  0.  schisticeps 53 

V.  Skulls  (ventral  views)  of  Ochotona  princeps  and  0.  collaris 54 

VI.  Skulls  (ventral  views)  of  Ochotona  princeps  and  0.  schisticeps 55 

TEXT  FIGURES. 

Fig.  1.  Map  of  the  distribution  of  the  American  species  of  Ochotona  (subgenus 

Pika) 2 

2.  Map  of  the  distribution  of  the  subspecies  of  Ochotona  princeps 11 

3.  Map  of  the  distribution  of  Ochotona  collaris 34 

4.  Map  of  the  distribution  of  the  subspecies  of  Ochotona  schisticeps 37 

IV 


No.  47.  NORTH  AMERICAN  FAUNA.  August  21,  1924 


REVISION  OF  THE  AMERICAN  PIKAS. 

(Genus  Ochotona). 


By  Arthur  H.  Howell. 


INTRODUCTION. 

Pikas,  or  "  conies,"  *  as  they  are  more  frequently  called  in  America, 
are  small,  tailless,  rabbitlike  mammals,  which  on  account  of  their 
shyness  and  the  inaccessible  nature  of  their  haunts  are  comparatively 
little  known.  (See  frontispiece.)  They  are  widely  distributed  in  Asia 
and  extreme  eastern  Europe,  where  the  genus,  Ochotona,  is  represented 
by  three,  or  possibly  four,  subgenera  and  a  large  number  of  species;2 
in  the  Western  Hemisphere  they  are  found  only  in  western  North 
America,  from  Alaska  to  southern  California  and  northern  New  Mex- 
ico. In  this  region  are  found  three  species — Ochotona  princeps,  0. 
collaris,  and  0.  schisticeps , with  numerous  subspecies. 

DISTRIBUTION    AND    HABITAT    OF    PIKAS    IN    AMERICA. 

Pikas  occur  in  western  North  America,  chiefly  in  the  mountains, 
from  the  Mount  McKinley  Range  of  central  Alaska  and  the  Ogilvie 
Mountains  of  central  Yukon  south  interruptedly  to  the  southern  Sierra 
Nevada  in  California  (headwaters  of  Tule  lliver) ,  the  Toyabe  Moun- 
tains in  central  Nevada,  the  Parowan  Mountains  in  southern  Utah, 
and  the  Jemez  and  Pecos  River  Mountains  in  northern  New  Mexico: 
east  to  the  Macmillan  Mountains,  Yukon,  Mount  Inglesmaldie  (near 
Banff),  Alberta,  Belt  Mountains,  Montana,  Big  Horn  Mountains, 
Wyoming,  and  Pikes  Peak,  Colorado ;  and  west  to  the  Chigmit  Moun- 
tains, Alaska,  and  the  Olympic  Mountains,  Washington  (reported,  but 
no  specimens  known). 

As  shown  by  the  accompanying  map  (fig.  1),  pikas  are  absent  from 
wide  areas  within  the  boundaries  of  their  general  range.  In  the  United 
States  this  absence  is  explained  by  the  presence  of  extensive  plains 
and  deserts  unsuited  to  then-  habitat,  but  in  central  and  northern  Brit- 
ish Columbia  a  broad  gap  is  found  between  the  ranges  of  Ochotona 
collaris  and  0.  priuceps  in  a  region  seemingly  well  adapted  to  their 
occupation. 

'  The  name  "cony"  is  applied  both  to  the  English  rabbit  {Oryciolagus  cuniculus)  and  to  the  daman,  or 
Syrian  hyrax  ( Procavia  syriaca) ;  the  name  "  pika,"  which  has  been  in  common  use  in  Europe  for  the  mem- 
bers of  the  genus  Ochoioni  for  many  years,  is  much  to  be  preferred;  the  latter  name  is  fro:n  the  vernacu- 
lar of  the  Tunguses — a  tribe  living  in  northeastern  Siberia— who,  according  to  Pallas  (Reise,  vol.  2,  p.  701 , 
1773j,  call  this  anin.al  "  peeka." 

lFor  the  subgeneric  groups,  see  Lyon,  M.  W.,  Smiths.  Misc.  Coll.,  vol.  45,  p.  438,  1904;   Thomas,  Old 
field,  Proc.  Zool.  Soc.  London,  1908,  p.  982. 


NORTH  AMERICAN  FAUNA. 


[No.  47 


FlG'  VwS^qS?^  °f  ihe  Ain?','ic^n  species  of  Ochotona  (subgenus  Pika)-in  the  western 
.  .n.ted  States,  Canada,  and  Alaska.    1.  o.  collaris.     2.  O.  prtnceps.    3.  0.  scnsticeps 

^ST?f  g  a  Jou™ey  in  1910  through  Canada,  from  Telegraph  Creek,  on 
the  btikine  kiver,  southeastward  to  Peace  Hirer,  by  war  of  the  head- 
waters of  the  Finlay  River  and  Fort  Grahame,  Edward  A.  Preble 
of  the  Biological  Surrey,  failed  to  find  any  eridence  of  the  presence 
of  pikas  in  those  parts.  He  was  informed,  also,  by  Dan  Brown  who 
accompanied  him  on  the  trip,  and  who  has  lired  and  traveled 'for  a 
number  of  years  as  a  hunter  and  prospector  in  this  region  and  fur- 
thermore is  well  acquainted  with  the  pikas  in  Alaska,  that  these  ani- 


1924.]  INTRODUCTION.  8 

mals  are  apparently  absent  from  the  Dease  River  country  of  northern 
British  Columbia,  the  area  drained  by  the  Muddy  (or  Kachika)  and 
the  upper  tributaries  of  the  Stikine.  Interrogation  of  a  number  of 
intelligent  Indians  in  this  region  plainly  showed  that  they  did  not 
know  the  animal.  It  is  not  impossible  that  some  form  of  the  genus 
may  occur  locally  in  the  coast  mountains  of  British  Columbia,  thus  in 
part  bridging  the  gap  between  the  ranges  of  0.  collaris  and  0.  princeps. 

The  American  pikas  are  practically  confined  to  a  single  type  of 
habitat — piles  of  broken  rock.  (PI.  II,  fig.  1.)  Quite  rarely  they  have 
been  found  living  in  piles  of  lumber  or  slabs  (Silverton,  Colorado)  or 
in  log  jams  in  streams  (Goose  Prairie  and  Easton,  Washington). 
Their  selection  of  rock  slides  for  homes  results  in  their  restriction 
chiefly  to  mountain  sides,  but  at  least  two  races  (Ochotona  schisti- 
cevs  taylori  and  0.  s.  goldmani)  are  found  on  the  plains,  living  in  piles 
of  lava  rock  (Madeline  Plains,  California;  Jacob  Ranch,  20  miles 
northeast  of  Adel,  Oregon;  and  the  Snake  River  Desert,  Idaho). 
Usually  they  occur  only  at  rather  high  altitudes,  chiefly  at  or  above 
timberline,  but,  given  proper  conditions,  altitude  apparently  has 
little  to  do  with  their  distribution,  since  they  are  known  to  range 
from  a  few  hundred  feet  above  sea  level  (North  Bend,  on  Fraser 
River,  British  Columbia)  up  to  13,600  feet  (Wheeler  Peak,  New 
Mexico). 

In  rock  slides  the  pikas  live  in  families  or  loose  colonies,  finding  an 
abundance  of  food  in  the  herbage  and  shrubbery  growing  near  at 
hand,  and  enjoying  protection  from  their  enemies  in  the  interstices 
of  the  slides. 

HABITS. 

Pikas  are  chiefly  diurnal,  but  are  frequently  heard  calling  during 
the  night  in  their  retreats  among  the  rocks.  They  are  very  active  and 
agile,  running  with  great  facility  over  the  rough  rocks  among  which 
they  live,  often  making  short  leaps  from  rock  to  rock,  their  furry 
soles  doubtless  making  them  very  sure  footed.  They  do  not  ordi- 
narily climb  except  into  low  shrubbery;  but  they  have  been  observed 
to  walk  out  on  low  horizontal  limbs  of  aspens  to  cut  the  leaves.  The 
gait  of  the  pika  has  been  described  as  "a  slightly  hopping  run" 
(Osgood) ;  Grinnell  and  Storer  state  that  the  hobbling  gait  reminds 
one  somewhat  of  the  hopping  of  a  brush-rabbit  [Sylvilagus  bacJimani], 

During  sunny  days,  pikas  often  sit  hunched  up  and  motionless  for 
considerable  periods  in  some  protected  place  where  they  can  command 
a  wide  view,  and  if  danger  threatens  be  ready  instantly  to  seek  the 
shelter  of  their  rock  slide.  These  observation  posts  are  usuall}* 
marked  by  accumulations  of  excrement  "of  an  o  Watery  spherical 
shape,  like  those  of  a  rabbit  but  much  smaller,  and  by  whitish  stains 
due  to  the  action  of  the  liquid  excrement  on  the  granite."3  The 
animals  are  untidy  in  their  habits,  often  dropping  their  excrement 
promiscuously  on  their  "haystacks." 

Pikas  are  usually  shy  and  difficult  to  approach  except  by  stealth, 
but  at  certain  times  and  places  they  are  quite  unsuspicious.  In  the 
Pecos  River   Mountains,  New  Mexico,  Vernon   Bailey  was    able   to 

'Grirmell  and  Storer,  Sierra  Club  Bull.,  vol.  10,  no.  2,  p.  163,  1917. 


4  NORTH   AMERICAN    FAUNA.  [No.  47 

observe  and  photograph  them  at  a  distance  of  a  few  feet ;  one  indi- 
vidual in  its  haste  to  move  a  bunch  of  "hay"  even  collided  with 
his  boot. 

Little  is  known  of  the  pikas'  habits  in  winter,  but  the  fact  that 
they  store  food  indicates  that  probably  they  do  not  hibernate  but 
remain  more  or  less  active  in  their  retreats  among  the  rocks  while 
the  mountains  are  covered  with  snow.  E.  R.  Warren  mentions  hav- 
ing seen  one  abroad  in  Colorado  in  January;4  and  J.  Aldcn  Loring 
reports  having  heard  several  near  Chama,  New  Mexico,  on  December 
23,  1893.  / 

Ordinarily  the  American  pikas  do  not  dig  burrows,  but  on  Mount 
Rainier,  Washington,  Merriam  found  a  colony  living  in  burrows  under 
large  rocks,  and  in  the  same  region  observed  a  few  of  their  burrows 
among  dwarf  fir  trees  at  some  distance  from  a  mass  of  slide  rock. 
Apparently  nothing  is  known  as  to  the  kind  of  nests  that  pikas  con- 
struct, but  doubtless  in  the  shelter  of  the  rock  slides  some  form  of  nest 
is  used  for  breeding  purposes. 

The  young,  usually  three  or  four,  or  rarely  five,  in  number,  are 
brought  forth  from  late  in  May  or  early  in  June  to  early  in  September. 
Two  young  specimens  from  Mullan,  Idaho,  collected  June  9  and  10, 
appeared  to  be  several  weeks  old  and  must  have  been  born  in  May. 

VOICE. 

The  notes  of  the  pika  are  variously  described  as  a  bleat,' or  bark, 
somewhat  ventriloquial  in  character*.  Grinnell  and  Storer  describe 
the  ordinary  call  as  a  moderately  loud  two-  or  three-syllabled  utter- 
ance with  a  nasal  intonation.  Osgood  characterizes  the  note  of  the 
Alaska  species  as  a  "sharp  metallic  bark";  Vernon  Bailey  describes 
that  of  Ochotona  p.  uinta  and  of  0.  p.  lerfiJii  as  a  nasal  squeak,  some- 
what resembling  the  bleat  of  a  very  young  lamb;  and  Thomas  Drum- 
mond  compares  the  cry  of  0.  princeps  to  that  of  a  rabbit  when  hurt. 
Several  observers  have  testified  to  marked  variation  in  the  notes  of  the 
different  species  of  pikas.  Merriam,  describing  the  notes  of  one  seen 
in  Yosemite  National  Park,  California,  says:  5  "  He  uttered  a  rather 
sharp,  harsh  cry,  something  like  Mh-Jcah,  usually  repeated  twice 
but  sometimes  three  times,  very  different  from  the  high-pitched  eet-eet 
of  the  Idaho  species  [lemhi]  and  less  shrill  than  the  whistling  cry  of  some 
of  the  others."  Writing  of  pikas  as  observed  on  Mount  Rainier, 
Washington,  Merriam  also  says:6  "In  addition  to  their  usual 
'  bleat '  one  of  them  in  a  den  of  rocks  very  near  me  made  a  low,  chat- 
tering cry  suggestive  of  a  wren  and  seemingly  made  at  or  for  the 
benefit  of  another."  Merritt  Gary,  writing  of  a  colony  observed  on  a  cold 
and  windy  day  in  the  Big  Horn  Mountains,  Wyoming,  says: 7  "  Their 
alarm  notes  were  seldom  heard,  but  a  low-pitched,  semirasping,  mew- 
ing note  was  occasionally  uttered  by  one  of  the  little  fellows  as  it  sat  on 
a  sharp-pointed  crag  in  the  midst  of  the  slide  rock,  each  note  being 
accompanied  by  a  forward  movement  of  the  head." 

*  Warren,  E.  R.,  Mammals  of  Colorado,  p.  55,  1910. 
6  Merriam,  C.  Hart,  MS.  notes,  July,  1903. 
•Merriam,  C.  Hart,  MS.  notes,  August.  1897. 
TCary,  Merritt,  MS.  notes,  May,  1910. 


North  American  Fauna  No.  47,  U.  S.  Dept.  Age, 'Biological  Survey.        PLATE  1 1. 


Fig.  I.— Pika  in  Rockslide,  Rednell  Basin,  Gunnison  County, 
Colorado. 


Fig.  2.— Haystack  of  Pika,  Trappers  Lake,  Garfield  County. 
Colorado. 

From  photographs  by  Edward  R.  Warren. 


1924.]  INTRODUCTION. 


FOOD. 


The  most  characteristic  habit  of  the  pikas  is  that  of  storing  food 
for  winter  use.  In  late  summer  and  earty  fall  the  animals  indus- 
triously gather  the  stems,  leaves,  and  flowers  of  a  variety  of  plants 
growing  in  or  near  their  home  rock  slides  and  pile  these  cuttings 
in  miniature  "hay  stacks"  on  shelving  rocks,  in  spaces  between 
rocks,  or  more  rarely  under  logs  or  stumps  in  the  slide,  where  they 
are  exposed  to  the  air  but  well  protected  from  rain  by  overhang- 
ing rocks.  These  stacks  average  about  the  size  of  a  bushel  meas- 
ure and  sometimes  attain  a  height  of  2\  feet.  (See  PI.  II,  fig.  2.) 
The  "hay"  is  gathered  by  the  pikas  often  at  a  distance  of  several 
hundred  feet  from  their  den  and  carried  in  their  mouths  in  small 
bundles,  which  are  deposited  on  the  stack.  Beaten  paths  often  lead 
from  one  stack  to  another  or  from  the  feeding  grounds  to  the 
den.  The  animals  evidently  exercise  some  discrimination  in  selecting 
the  species  of  plants  desired,  and  different  colonies  vary  in  their  choice, 
some  selecting  chiefly  one  or  two  varieties,  others  a  considerable  num- 
ber. In  the  Lemhi  Mountains,  Idaho,  Merriam  found  their  chief  food 
plant  to  be  a  species  of  avens  (Geum  rossii).6 

A  stack  examined  by  Bailey  in  New  Mexico  contained  34  recog- 
nizable species  of  plants,  the  most  common  being  nettles,  chokecherry, 
gooseberry,  currant,  raspberry,  rose,  snowberry,  svringa,  lupine,  aspen, 
and  phacelia.  Other  plants  commonly  utilized  by  pikas  are  golden- 
rod,  fireweed  (Epilobium) ,  lupine,  aster,  avens  (Geum),  gentian,  Indian 
tobacco  (Antennaria),  thistles,  beardtongue  (Pentstemon),  bearberry 
(Arctostaphylos) ,  heather  (Bryanthus),  knotweed  (Polygonum),  saxi- 
frage, huckleberry,  cinquefoil  (Potentilla),  yarrow  (Achillea),  phlox, 
ferns,  and  a  variety  of  grasses  and  sedges.  In  Yellowstone  Park, 
Wyoming,  Bailey  examined  a  pika's  haystack  in  which  grass  formed 
about  half  the  total  material  and  blueberry  bushes  about  a  fourth. 

In  some  localities  pikas  seem  to  have  a  preference  for  the  leaves 
of  trees  and  shrubs.  In  the  Natchez  River  Valley,  Washington,  Mer- 
riam found  their  "hay"  composed  almost  entirely  of  the  branches  of 
chokecherry  (Prunus  demissa)  about  a  foot  and  a  half  long,  with  some 
branches  of  the  service  berry  (Amelanchier) ,  a  little  aspen  (Populus), 
much  wild  rose,  Spiraea  of  two  species,  raspberry,  currant,  and  a  few 
herbaceous  plants. 

In  the  Gros  Ventre  Mountains  near  Jackson  Hole,  Wyoming,  Wet- 
more  found  the  pikas'  "hay"  made  up  largely  of  leaves  of  elders, 
which  in  some  cases  had  been  cut  off  as  high  as  2\  feet  above  the 
ground,  some  of  the  shrubs  being  nearly  stripped  of  their  leaves. 
On  the  lava  beds  of  Oregon,  sagebrush  is  sometimes  used  for  the 
hay.  In  Alaska  Osgood  found  most  of  the  common  plants  in  the 
pika  haystacks,  Dry  as  being  most  frequently  utilized. 

PELAGE    AND   MOLT. 

The  pelage  of  the  pika  is  dense,  soft,  and  silky,  and  in  winter 
moderately  long;  the  feet  and  toes  are  well  haired,  the  soles  and 
palms  with  dense  furcy  pads.  The  basal  portion  of  the  body  hairs 
is  a  uniform  dark  neutral  gray  or  a  deep  neutral  gray  (Ridgway, 

•  North  Amer.  Fauna  No.  5,  p.  73, 1891. 


6  NORTH   AMERICAN   FAUNA.  [No.  47 

"Color  Standards,"  PI.  LIII) ;  the  terminal  or  subterminal  portion 
varies  from  smoke-gray  through  many  different  shades  of  buff  and 
cinnamon  to  tawny;  the  tips  of  many  of  the  hairs  on  the  dorsal  area 
are  fuscous  or  blackish,  differing  quantities  of  this  color,  with  varia- 
tions in  the  shade  of  the  subterminal  portion,  producing  many 
color  tones. 

All  the  American  species  except  Ochotona  collaris  have  two  annual 
molts,  and  in  most  of  the  races  the  winter  and  summer  pelages  differ 
decidedly  in  color.  In  some  species  the  midsummer  molt  begins  early 
in  July  (rarely  early  in  June)  with/certain  individuals,  and  is  frequently 
completed  by  the  20th  of  that  month;  in  others  this  molt  frequently 
does  not  begin  until  the  middle  or  last  of  August  and  is  not  complete 
until  well  into  September.  In  the  Sierra  Nevada  races  of  0.  schisticevs 
this  molt  apparently  occurs  considerably  later  than  in  most  of  the 
other  forms. 

The  new  pelage  usually  appears  first  on  the  head,  the  molt  pro- 
gressing evenly  Dackward  over  the  shoulders  and  back  and  on  the 
sides  to  the  line  of  the  belly;  occasionally  the  molt  is  irregular,  the 
new  hair  appearing  in  isolated  patches  on  the  head,  shoulders,  and 
back;  the  underparts  are  renewed  last. 

The  midsummer  pelage  is  worn  a  comparatively  short  time — 
probably  not  over  two  months  and  apparently  often  much  less  than 
that — for  in  some  species  the  winter  pelage  begins  to  appear  early  in 
September,  and  by  the  last  of  that  month  the  fall  molt  is  practically 
complete.  This  molt  apparently  progresses  in  practically  the  same 
way  as  the  midsummer  molt.  In  the  fresh  winter  pelage  the  fuscous 
or  blackish  tips  of  the  hairs  on  the  back  are  usually  prominent; 
later  in  the  season  these  tips  wear  away  to  some  extent,  leaving  the 
animal  with  a  more  grayish  tone  in  midwinter  than  in  early  fall. 

The  winter  pelage  wears  away  slowly  and  evenly;  specimens  taken 
in  May  or  June  show  only  moderate  wear,  but  when  the  midsummer 
molt  is  delayed  until  late  in  July  or  into  August  the  animals  often 
present  a  greatly  worn,  faded,  and  patchy  appearance. 

The  most  northerly  ranging  American  species — 0.  collaris — ap- 
parently has  but  a  single  annual  molt,  which  extends  from  late  in 
July  to  early  in  September.  No  specimens  of  collaris  taken  later 
than  September  1  and  2  are  available,  but  specimens  taken  on  those 
dates  had  nearly  acquired  the  fresh  pelage,  which  has  every  appear- 
ance of  being  the  same  as  that  of  specimens  taken  late  in  April. 

The  progress  of  the  molts  is  described  in  greater  detail  under  the 
different  species. 

MATERIAL    EXAMINED. 

The  present  revision  is  based  on  a  study  of  1,315  specimens;  of 
these  approximately  two-thirds  are  contained  in  the  United  States 
National  Museum,  chiefly  in  the  Biological  Survey  collection;  the 
remainder  have  been  borrowed  from  other  American  museums  and 
from  private  collections.9 

•  For  the  loan  of  this  material  the  author  desires  to  extend  thanks  to  the  owners  and  custodians,  as 
follows:  H.  E.  Anthony,  of  the  American  Museum  of  Natural  History;  Dr.  Glover  M.  Allen  and 
Samuel  Uenshaw,  of  the  Museum  of  Comparative  Zoology,  Harvard  University:  Dr.  Wilfred  H.  Osgood, 
of  the  Field  Museum  of  Natural  History:  Dr.  Witmer  Stone,  of  the  Academy  of  Natural  Sciences  of 
Philadelphia;  Dr.  Joseph  Orinnell,  of  the  Museum  of  Vertebrate  Zoology,  University  of  California; 
J.  D.  Figgins,  of  the  Colorado  Museum  of  Natural  History;  F>r.  R.  M.  Anderson  and  P.  A.  Taverner,  of 
the  Victoria  Memorial  Museum;  F.  Kermode,  of  the  Provincial  Museum,  Victoria,  British  Columbia; 
Stanley  G.  Jewett,  of  Portland,  Oregon,  and  E.  R.  Warren,  of  Colorado  Springs,  Colorado. 


1924.J  GENUS   OCHOTONA.  7 

EXPLANATION    OP   CRANIAL   MEASUREMENTS. 

Ill  the  study  of  the  skulls  of  this  genus,  the  following  measurements 
were  taken  in  millimeters: 

Occipito-nasal  length. — Greatest  length  from  posterior  border  of 
occiput  to  anterior  tip  of  nasals. 

Zygomatic  breadth. — Breadth  across  zygomata  at  posterior  border 
of  orbits. 

Breadth  of  braincase. — Breadth  across  parietals  at  posterior  border 
of  auditory  meatus. 

Interorbital  breadth. — Least  breadth  of  frontals  between  the  orbits. 
Width  of  'palatal  bridge. — -Antero-posterior  length  of  bony  palate, 
from  posterior  border  of  palatal  foramen  to  anterior  border  01  inter- 
pterygoid  fossa. 

Length  of  nasals. — Greatest  length  (measured  with  vernier  calipers) 
from  plane  of  posterior  border  to  anterior  tip. 

Genus  OCHOTONA  Link. 

Lepus  Pallas,  Reise,  vol.  2,  p.  701,  1773  (part).     Not  Lepus  Linnaeus,  1758. 

Ochotona  Link,  Beytrage  Naturg.,  vol.  1,  pt.  2,  p.  74,  1795.  Type,  Ochotona  minor 
Link=Lepus  ogotona  Pallas=Lepws  dauuricus  Pallas.10 

Pika  Lacepede,  Tabl.  Mamm.,  1799,  p.  9.     Type,  Pika  alpinus=Lepus  alpinus  Pallas. 

Lagomys  G.  Cuvier,  Lemons  Anat.  Comp.,  vol.  1,  tabl.  I,  1800  (not  Lagomys  Storr, 
1780). 

Ogotoma  Gray,  Ann.  Mag.  Nat.  Hist.,  3d  ser.,  vol.  20,  p.  220,  1867.  Type,  Ogotoma 
pallasii=Lagomys  ogotona  Waterhouse  (not  Lepus  ogotona  Pallas).10 

Conothoa  Lyon,  Smiths.  Misc.  Coll.,  vol.  45,  p.  438,  1904  (subgenus).  Type,  Ocho- 
tona roylii  Ogilby. 

Generic  characters. — Form  compact;  tail  very  short — practically 
obsolete:  limbs  short,  the  hind  legs  bub  little  longer  than  fore  legs; 
soles  and  palms  densely  furred;  toes  5  in  front,  4  behind,  armed  with 
curved,  slender  claws;  a  naked  pad  at  base  of  each  toe.  Ears  moder- 
ately large,  suborbicular,  haired  on  both  surfaces;  eyes  rather  small; 
fur  soft  and  moderately  dense.  Mammae  4  or  6:  Pectoral,  $  or  •§; 
inguinal,  \. 

Braincase  rather  flat — not  inflated;  rostrum  relatively  narrow,  the 
nasals  broader  anteriorly  than  posteriorly  or  of  equal  width;  post- 
orbital  processes  lacking;  interorbital  constriction  pronounced;  bony 
palate  relatively  short;  anteroinferior  angle  of  zygoma  with  pro- 
nounced spinous  process;  malar  bone  greatly  prolonged  posteriorly, 
reaching  nearly  to  the  auditory  opening;  maxillary  bones  usually 
without  fenestration;  ascending  ramus  of  mandible  relatively  wide, 
with  the  condyle  well  developed;  anterior  face  of  ascending  ramus 
without  the  groove  found  in  the  Leporidae ;  the  coronoid  process  con- 
sisting of  a  small  tubercle,  often  nearly  obsolete;  a  smaller  tubercle 
often  appearing  lower  down  on  the  ascending  ramus,  near  the  last 
molar;  angular  process  of  mandible  relatively  long,  slender,  and 
recurved,  the  angle  of  the  jaw  pronounced;  mental  foramen  situated 
on  the  side  of  the  horizontal  ramus,  beneath  the  last  lower  molar. 

Dental  formula :  I,  f ;  C,%;  PM,f;  M, -f;  third  upper  molar  lacking; 
molars  with  simple  enamel  pattern,  the  reentrant  angles  without  crena- 
tion;  second  upper  premolar  decidedly  smaller  than  thud,  with  a  reen- 

to Cf.  Thomas,  Proc.  Zool.  Soc.  London,  1908,  p.  109. 


8  NORTH   AMERICAN   FAUNA.  [So.  47 

trant  angle  on  anterior  face;  second  (last)  upper  molar  with  a  project- 
ing enamel  loop  on  intero-posterior  border;  last  lower  molar  simple 
(not  double) ;  cutting  edge  of  first  upper  incisor  V-shaped.11 

HISTORY    AND    NOMENCLATURE. 

The  pikas  were  first  made  known  to  science  in  1769,  when  Pallas  pub- 
lished the  description  of  a  species  from  eastern  Russia  and  southern 
Siberia  under  the  name  Leyus  pusillus.12  During  the  next  few  years 
the  same  author  described  several/  additional  species  from  Asia  and 
gave  full  accounts  of  their  habits  and  anatomy.13 

In  1795,  Link  separated  the  pikas  from  the  hares  under  the  generic 
name  Ochotona*4 — evidently  derived  from  the  Mongolian  vernacular, 
which  Pallas  gives  as  "ochodona"  01  "  ogotona."  15  Link's  name,  how- 
ever, was  overlooked  or  disregarded  by  his  contemporaries,  and  Cuvier's 
name  Lagomys,  proposed  for  the  genus  in  1800, 16  was  currently  used 
for  almost  a  century,  until  Thomas,  in  1896, 17  showed  that  it  was 

Ereoccupied  by  Lagomy?  Storr,  1780,  and  furthermore  was  antedated 
y  Ochotona  Link. 

In  1799,  Lacepede  introduced  the  name  PiJca  in  conjunction  with  one 
of  the  Siberian  species,  alpinus:18  the  name  is  antedated,  of  course,  by 
Ochotona,  but,  as  shown  by  Lyon,19  it  is  available  as  the  name  for  the 
subgenus  containing  all  the  American  species. 

In  1867,  in  a  paper  on  the  skulls  of  hares  and  pikas,  Gray  proposed 
a  new  generic  name,  Ogotoma,  for  the  species  described  by  Waterhouse 
under  the  name  Lagomys  ogotona,  naming  it  Ogotoma  pallasii.20 
Gray's  name  Ogotoma'  has  recently  been  revived  by  Thomas  in  a 
subgeneric  sense.21 

In  1904  Lyon  published  an  important  paper  on  the  "Classification 
of  the  Hares  and  Their  Allies,"  in  which  he  described  in  detail  the 
osteology  of  the  pikas  and  proposed  a  new  subgeneric  name,  Conoihoa, 
based  on  Ochotona  roylii  Ogilby,  from  central  Asia.22 

The  first  American  species  to  be  described  was  Richardson's  Lepus 
{Lagomys)  princeps,  1828,  based  on  specimens  collected  by  Thomas 
Drummond,  in  the  Rocky  Mountains  of  Canada,  near  the  sources  of 
the  Athabaska  River.  In  I860  Lord  proposed  a  second  species, 
Lagomys  minimus,  based  on  specimens  taken  by  himself  on  the  inter- 
national boundary  in  southern  British  Columbia.  This  name,  how- 
ever, was  placed  in  synonymy  by  later  authors  until  revived  by  Bangs 
in  1899.  Thus  princeps  remained  for  over  60  years  the  only  recog- 
nized American  species. 

In  1889  Merriam  named  a  new  species,  schistic.eps,  from  the  Sierra 
Nevada,  and  in  1893  Nelson  described  the  Alaskan  species  under  the 
name  collaris.     In  1899  Bangs  described   two  new  forms — saxatilis, 

11  For  a  fuller  discussion  of  the  osteological  differences  between  Ochr.tona  and  the  I.eporidae,  cf.  Lvon, 
Smiths.  Misc.  Coll.,  vol.  45,  pi).  384-385,  1904. 
'2  Pallas,  P.  S.,  Nov.  Comm.  Acad.  Sci.  Petropolit.,  vol.  13,  p.  531,  1769. 
;3 Pallas,  P.  S.,  Reise,  vol.  2,  p.  701,1773,  and  vol.  3,  p.  692,  1776;  Nov.  Spec.  Quad.,  Glires,p.59,  1778. 

14  Link,  D.  H.  P.,  Bevtrage,  vol.  1.  pt.  2.  p.  74,  1795. 

15  Pallas,  P.  S.,  Reise,  vol.  2.  p.  701:  vol.  3.  p.  692. 

16  Cuvier,  F.,  Lecons  d'  Anat.  Cornp.,  vol.  1  (Tab.),  1800. 
"  Thomas,  O.,  Proc.  Zool.  Soc.  London,  1896,  p.  1026. 

>»  Lacepede,  B.  G.  E.,  Tabl.  ilamm.,  p.  9,  1799. 
;9  Lvon,  M.  \Y.,  Smiths.  Misc.  Coll.,  vol.  45,  p.  332,  1904. 

2»Grav.  J.  E.,  Ann.  and  Mag.  Nat.  Hist.,  3d  ser..  vol.  20,  p.  220,  1867;  cf.  Thomas,  Proc.  Zool.  Soc. 
London',  1908,  pp.  109-110. 
"  Thomas,  Oldfield,  Proc.  Zool.  Soc.  London,  1908,  p.  982. 
«  Lvon.  M.  W.,  Smiths.  Misc.  Coll.,  vol.  45,  p.  438,  1904. 


1924.]  SUBGENUS  PIKA.  9 

from  Colorado,  and  cuppes,  from  southeastern  British  Columbia — and 
in  1905  Allen  described  cinnamomea,  from  southern  Utah.  In  1912 
five  new  species  were  proposed — Jigginsi  by  Allen,  from  Colorado; 
uinta  and  Levis  by  Hollister,  from  Utah  and  Alberta,  respectively; 
and  taylori  and  albatus  by  Grinnell,  both  from  California.  A  species 
from  New  Mexico,  nigrpscens,  was  described  by  Bailey  in  1913,  and 
two  new  races  from  California,  muiri  and  sJipltoni,  by  Grinnell  in  1916 
and  1918,  respectively.  In  1919  the  present  author  described  nine 
new  forms  from  various  parts  of  Canada  and  the  United  States — 
brunnescens,  from  Washington;  fumosa  and  jewetti,  from  Oregon; 
fuscipes,  from  Utah;  incana,  from  New  Mexico;  lemhi,  from  Idaho; 
lutescens,  from  Alberta;  nevadensis,  from  Nevada;  and  ventorum,  from 
Wyoming. 

Two  new  races  are  described  in  this  paper — broolcsi,  from  British 
Columbia,  and  goldmani,  from  Idaho — bringing  the  total  number  of 
recognized  American  forms  to  26.  Further  collecting  in  previously 
unworked  regions  will  doubtless  reveal  a  number  of  additional 
species  or  subspecies. 

Subgenus  PIKA  Lacepede. 

Subgeneric  characters.23 — Skull  very  flat;  interorbital  region  rather 
broad  (not  elevated  and  not  much  constricted,  as  in  Ogotoma)  ;  maxil- 
lary orifice  roundly  triangular  (much  as  in  Ogotoma,  less  elongated 
than  in  Ochotona) ;  incisive  and  palatal  foramina  separate  (as  in 
Ogotoma) . 

Remarks. — The  subgenus  Pilca,  includes  all  the  American  species 
and  is  found  also  practically  throughout  the  range  of  the  genus  in 
Asia  and  eastern  Europe. 

List  of  American  Species  and  Subspecies,  -with  Type  Localities. 

Ochotona  princeps  princeps  (Richardson) . .  Headwaters  of  Athabaska  River,  near  Ath- 

abaska  Pass,  Alberta. 

princeps  lutescens  Howell Mount  Inglestnaldie,  near  Banff,  Alberta. 

princeps  levis  Hollister Chief  Mountain  [Waterton]  Lake,  Alberta. 

princeps  lemhi  Howell Lemhi  Mountains,  west  of  Junction,  Idaho. 

princeps  ventorum  Howell Fremont  Peak,   Wind   River   Mountains, 

Wyoming. 

princeps  uinta  Hollister Uinta  Mountains,  near  head  of  east  fork 

Bear  River,  Utah. 

princeps  nevadcnsis  Howell Ruby  Mountains,  southwest  of  Ruby  Val- 
ley Post  Office,  Nevada. 

princeps  Jigginsi  J.  A.  Allen Pagoda  Peak,  Rio  Blanco  County,  Color- 
ado. 

princeps  saxatilis  Bangs Montgomery,  near  Mount   Lincoln,   Park 

County,  Colorado. 

princeps  incaiva  Howell Pecos  Baldy,  New  Mexico. 

princeps  nigrescens  Bailey Jemez  Mountains,  New  Mexico. 

princeps  cuppes,  Bangs Monashee    Divide,    Gold    Range,    British 

Columbia. 

princeps  fenisex  Osgood "  Ptarmigan  Hill,"  near  head  of  Ashnola 

River,  Cascade  Range,  British  Columbia. 

princeps  brooksi,  nobis Sicarnous,  British  Columbia. 

princeps  brunnescens  Howell Keechelus,  Kittitas  County,  Washington. 

princeps  fumosa  Howell Permilia  Lake,  west  base  of  Mount  Jeffer- 
son, Oregon. 

"The  characters  of  the  subgenus  Qshotom,  as  given  by  Lyon  ( Smiths.  Misc.  Coll.,  vol.  45,  p.  438, 1904) 
are  in  reality  those  of  the  subgenus  Oqofnma  Gray,  as  has  been  pointed  out  by  Thomas  (Proc.  Zool.  Soc. 
London,  1908,  p.  982). 


10  NORTH   AMERICAN    FAUNA.  FNo.  47 

List  of  Species,  with  Type  Localities — Continued. 

Ochotona  collaris  (Nelson) Near  head    of  Tanana  River  (about  200* 

miles  south  of  Fort  Yukon),  Alaska. 

schisticeps  schisticeps  (Merriam) Donner,  California. 

schisticeps  taylori  Grinnell Warren  Peak,  Warner  Mountains,  Modoe 

County,  California. 
schisticeps  goldmani,  nobis Echo  Crater,  Snake  River  Desert  (20  miles 

southwest  of  Arco),  Idaho. 
schisticeps  jewetti  Howell Headof  Pine  Creek,  near  Cornucopia,  Baker 

County,  Oregon. 
schisticeps  muiri  Grinnell  and  Storer.  .Ten  Lakes,  Yosemite  National  Park,  Cal- 

/    ifornia. 
schisticeps  albata  Grinnell Cottonwood  Lakes,  near  Mount  Whitney r 

Inyo  County,  California. 
schisticeps  sheltoni  Grinnell White    Mountains,   near    Big   Prospector 

Meadow,  Mono  County,  California. 

schisticeps  cinnamomea  J.  A.  Allen Beaver  Mountains,  Utah. 

schisticeps  fuscipes  Howell Parowan  Mountains,  Iron  County,  Utah. 

Key  to  the  American  Species. 

a1  Underparts   creamy  white,   without  buffy   wash;    an  indistinct    grayish 

"  collar"  on  shoulders collaris. 

a2  Underparts  washed  with  buff;  no  grayish  "  collar"  on  shoulders. 

b l  Palatal  bridge  narrow  (average  about  1.6  mm.) schisticeps. 

b2  Palatal  bridge  broader  (average  about  2.3  mm.) princeps. 

Descriptions  of  American  Species  and  Subspecies. 

OCHOTONA  PRINCEPS  (Richardson). 
[Synonymy  under  subspecies.] 

General  characters. — Coloration  grayish  or  cinnamon-buff,  more  or 
less  mixed  with  fuscous,  the  general  tone  varying  from  pale  smoke- 
gray,  mouse-gray,  or  drab  to  wood  brown,  cinnamon-buff,  or  cinnamon- 
arab;  in  summer  more  buffy,  ochraceous,  or  tawny  and  more  varied 
with  blackish  or  fuscous.  Skull  relatively  long  and  narrow,  with  nar- 
row interorbital  constriction ;  nasals  usually  rather  long;  palatal  bridge 
wide;  interpterygoid  fossa  U-shaped,  with  the  sides  usually  parallel 
(occasionally  somewhat  spatulate  in  brooksi,  uinta,  and  lemhi). 

Color.—  Upperparts  pinkish  buff,  pinkish  cinnamon,  vinaceous- 
cinnamon,  cinnamon-buff,  ochraceous-tawny,  or  clay  color,  more  or 
less  mixed  with  fuscous  or  blackish;  hinder  back  sometimes  washed 
with  smoke-gray;  sides  pinkish  buff,  pinkish  cinnamon,  vinaceous- 
cinnamon,  cinnamon-bun,  or  ochraceous-tawny;  ears  hair-brown,. 
deep  mouse-gray,  chaetura  drab,  chaetura  black,  or  fuscous  black, 
margined  with  buff  or  buffy  white:  feet  creamy  white,  pinkish  buff, 
light  pinkish  cinnamon,  or  cinnamon-buff;  soles  buffy  white,  hair- 
brown,  drab,  chaetura  drab,  or  fuscous;  palms  buffy  white,  light  drab, 
hair-brown,  or  pale  olive-buff;  underparts  white,  pinkish  buff,  pinkish 
cinnamon,  vinaceous-cinnamon,  or  cinnamon-buff. 

Geographic  distribution. — Rocky  Mountain  region,  from  east-central 
British  Columbia  (headwaters  of  South  Pine  River)  south  to  the 
Pecos  River  Mountains  and  Jemez  Mountains,  New  Mexico,  La  Sal 
Mountains,  Utah,  and  Ruby  Mountains,  Nevada;  east,  to  the  Belt 
Mountains,  Montana,  Big  Horn  Mountains,  Wyoming,  and  Pikes  Peak, 
Colorado;  also  the  Cascade  Mountain  system,  from  southern  British 
Columbia  south  to  the  vicinity  of  Crater  Lake,  Oregon.     (Fig.  2.) 

Remarks. — The  first  described  American  species  of  pika  has  an  ex- 
tensive range  in  the  Rocky  Mountain  region  and  is  separable  into  at 
least  16  races.     Typical  princeps  apparently  ranges  continuously  along 


1924.] 


OCHOTONA   PRINCEPS. 


11 


Fig.  2.— Distribution  of  the  subspecies  of  Ochotona  princeps. 


1.  O.  p.  bronksi. 

2.  O.  p.fenisex. 

3.  0.  p.  brunnescens. 

4.  O.  p.fumosa. 


5.  0.  p.  cuppes. 

6.  O.  p.  princeps. 

7.  O.  p.  lu'escens. 

8.  0.  p.  Zewis. 


9.  0.  p.  lemhi. 

10.  O.  p.  ventorum. 

11.  O.  p.  uinta. 

12.  0.  p.  nevadensis. 


13.  O.  p.  figginsi. 

14.  O.  p.  saxatilis. 

15.  O.  p.  nigrescent. 

16.  O.  p.  mcana. 


the  higher  ridges  of  the  main  divide  from  northern  Alberta  to  southern 
Montana.  A  dark  race  (cuppes)  occupies  most  of  southeastern  British 
Columbia,  on  the  western  slopes  of  the  main  range,  and  a  very  pale 
form  (lutescens)  is  found  on  the  outlying  ranges  of  Alberta,  east  of  the 
main  divide.  In  central  and  western  Montana,  levis,  paler  than 
typical  princeps  but  not  so  pale  as  lutescens,  is  found  in  the  Belt 
Mountains  and  at  Waterton  Lake.  In  central  Idaho,  another  small, 
pale  race  occurs.  In  western  Wyoming  and  northeastern  Utah  two 
rather  large,  richly  colored  forms  (ventorum  and  uinta)  connect  typical 


12  NORTH   AMERICAN   FAUNA.  [No.  47 

princeps  with  the  Colorado  and  New  Mexico  races ;  in  this  region  the 
easterly  races  (saxatilis  and  incana)  are  paler  than  the  more  westerly 
races  (jigginsi  and  nigrescens).  The  Ruby  Mountains  form,  although 
widely  isolated,  is  apparently  closely  related  to  uinta  and  has  no  close 
relationship  with  the  form  in  the  Toyabe  Mountains — 0.  schisticeps 
cinnamomea.  In  southern  British  Columbia  cuppes  grades  into fenisex, 
a  paler  form  occupying  the  mountains  lying  to  the  west  of  the  Okanagan 
Valley,  and  the  latter  passes  into  the  darker  race,  brunnescens,  of  the 
main  Cascade  Range.  In  central  British  Columbia  occurs  a  form 
(brooksi)  darker  than  fenisex  and  somewhat  resembling  brunnescens. 
The  darkest  form  of  all,  fumoffa,  is  found  on  the  west  slopes  of  the 
Cascade  in  Oregon  and  in  the  mountains  around  Paulina  Lake. 

OCHOTONA  PRINCEPS  PRINCEPS  (Richardson). 

Rocky  Mountains  Pika. 

(PI.  Ill,  fig.  1;  PI.  V,  fig.  1.) 

Lepus  (Lagomys)  princeps  Richardaon,  Zool.  Journ.,  vol.  3,  p.  520,  1828;  Fauna  Bor- 

eali-Americana,  p.  227, 1829. 
[Ochotona]  princeps  Trouessart,  Cat.  Mamm.  Viv.  Foss.,  p.  648,  1897. 

Type  locality. — Headwaters  of  Athabaska  River,  near  Athabaska 
Pass,  Alberta.24 

Geographic  distribution. — Rocky  Mountains,  from  eastern  British 
Columbia  (headwaters  South  Pine  River)  and  western  Alberta  (Mus- 
keg Creek)  south  along  the  main  divide  to  eastern  Idaho  and  western 
Montana  (Bitterroot  Mountains).      (Fig.  2,  p.  11.) 

Characters. — Size  medium;  skull  relatively  narrow  with  rather  long 
nasals,  narrow  interpterygoid  fossa,  and  moderately  wide  palatal 
bridge;  winter  pelage  uniform  bully  drab  or  mouse-gray;  summer 
pelage  clay  color  or  pinkish  cinnamon,  heavily  shaded  with  blackish. 

Color.—  Winter  pelage  (October):  Cinnamon  phase:  Upperparts 
mixed  light  pinkish  cinnamon  and  fuscous,  sometimes  washed  on 
hinder  back  with  blackish,  shading  on  lower  sides  to  light  pinkish 
cinnamon;  ears  chsetura  black  or  chaetura  drab,  margined  with  light 
buff;  feet  pinkish  buff;  soles  hair-brown;  palms  light  drab;  under- 
pays whitish,  moderately  washed  with  pinkish  buff  or  cinnamon-buff. 
Drab  phase:  Similar  to  the  cinnamon  phase  but  dorsal  surface  strongly 
washed  with  light  smoke-gray,  especially  on  hinder  back.  Summer 
pelage :  Upperparts  between  pinkish  cinnamon  and  clay  color,  richest 
on  head  and  shoulders,  heavily  shaded,  particularly  on  middle  and 
hinder  back,  with  black  or  fuscous-black:  sides  of  head  and  neck 
clay  color  or  pinkish  buff,  shading  on  sides  of  body  to  light  pinkish 
cinnamon;  underparts  dull  whitish,  moderately  washed  with  light 
pinkish  cinnamon  or  pale  cinnamon-buff;  ears  and  feet  as  in  winter 
pelage.  Young  (specimens  from  Upper  Stillwater  Lake,  Montana, 
July  2  and  3,  apparently  less  than  a  month  old) :  Upperparts  fuscous- 
black,  shaded  with  light  buff;  sides  washed  with  light  pinkish  cinna- 
mon ;  ears  blackish  mouse-gray,  margined  with  buffy  white ;  feet 
soiled  whitish,  washed  with  pale  pinkish  buff;  soles  mouse  gray: 
palms  soiled  whitish;  underparts  wnitish,  washed  with  pinkish  buff. 

**See  remarks  on  pp.  13-14. 


1&24.]  OCHOTONA  PRINCEPS.  13 

Molt. — The  progress  of  the  molts  and  the  pelage  changes  are  well 
shown  by  a  series  of  specimens  from  the  vicinity  of  Henry  House, 
Alberta.  A  specimen  taken  July  3  shows  the  midsummer  molt  begin- 
ning on  the  face,  head,  and  middle  of  fore  back  behind  the  ears,  the 
rest  of  the  upperparts  being  in  moderately  worn,  grayish  winter  pel- 
age; the  underparts  are  worn  and  the  pelage  thin,  with  only  a  slight 
wash  of  light  burl  in  the  median  line;  in  another  taken  July  17  the 
new  pelage  covers  the  entire  head  and  shoulders  and  reaches  nearly  to 
the  middle  of  the  back:  the  entire  underparts  are  densely  clothed 
with  what  appears  to  be  a  new  pelage  heavily  washed  with  buff:  one 
taken  July  13  has  the  summer  pelage  nearly  complete,  covering  the 
whole  dorsal  surface  except  the  rump  and  a  patch  on  one  side  of  the 
hinder  back;  the  ventral  surface  is  also  nearly  completely  renewed: 
a  specimen  taken  September  10  has  the  summer  pelage  greatly  worn 
on  the  back  and  sides,  though  but  little  on  the  belly:  in  one  taken 
the  same  date,  the  winter  pelage  has  covered  the  head,  shoulders,  and 
back  for  two-thirds  of  the  distance  from  nose  to  tail,  but  has  not 
reached  the  lower  sides:  this  pelage  does  not  differ  greatly  from  the 
fresh  summer  pelage  but  is  somewhat  more  grayish  or  vinaceous  and 
less  buffy:  in  a  specimen  taken  September  29  the  winter  pelage  covers 
the  whole  dorsal  surface  and  the  sides,  though  the  pelage  on  the  under- 
parts is  not  completely  renewed;  this  specimen  is  decidedly  more 
grayish  than  that  taken  on  September  10,  and  the  fuscous  or  blackish 
hairs  are  much  less  prominent;  specimens  taken  October  17  and  18 
are  in  full  winter  pelage,  with  drab  or  cinnamon  tones  prevailing. 

Farther  south  the  midsummer  molt  may  in  individual  cases  begin 
early  in  June,  though  as  a  rule  it  occurs  in  Montana  from  late  in  June 
to  early  in  August.  Two  specimens  from  the  Bitterroot  Mountains, 
near  Florence,  Montana,  June  7,  show  the  summer  pelage  just  appear- 
ing in  patches  on  the  shoulders  and  around  the  eyes. 

Skull. — Relatively  long  and  narrow  with  narrow  interorbital  region ; 
nasals  of  moderate  length:  palatal  bridge  rather  wide;  interpterygoid 
fossa  rather  narrow,  the  sides  nearly  parallel. 

Measurements. — Average  of  20  adults  (10  males,  10  females)  from 
vicinity  of  Henry  House,  Alberta:  Total  length,  191.6  (181-204); 
hind  foot,  30.5  (29-33) .  Shull:  Average  of  10  adult  males  from  vicin- 
ity of  type  locality  (Henry  House,  Grand  Cache  River,  and  Smoky 
River,  Alberta) :  Occipito-nasal  length,  43.5  (41.9-45.2);  zygomatic 
breadth,  21.4  (21-22);  breadth  of  braincase,  18  (17.2-19.1);  interor- 
bital breadth,  5.4  (4.5-5.7) :  width  of  palatal  bridge,  2  (1.6-2.3) ;  length 
of  nasals,  14.4  (13.4-15.4).  Average  of  8  adult  females  from  same 
localities:  Occipito-nasal  length,  42.5  (41.1-45);  zygomatic  breadth, 
21.3  (20.8-21.9);  breadth  of  braincase,  17.8  (17.1-18.6):  interorbital 
breadth,  5.5  (5-5.8);  width  of  palatal  bridge,  2.1  (1.8-2.7) ;  length  of 
nasals,  14  (13.1-14.7). 

Remarks.-  -The  Rocky  Mountain  pika  was  the  first  American  species 
to  be  discovered  and  for  many  years  (from  the  time  of  its  description 
in  1828  until  1889)  was  the  only  currently  recognized  species. 

Richardson,  who  described  the  form,  stated  that  "Drummond 
killed  several  [pikas]  near  the  sources  of  the  Elk  [  =  Athabaska] 
73502—24 2 


14  NORTH   AMERICAN   FAUNA.  [No.  47 

River,"25  and  Preble  considers  that  these  specimens  were  undoubt- 
edly the  basis  of  Richardson's  description. 26  The  type  locality,  there- 
fore, may  be  definitely  fixed  as  the  source  of  the  Athabaska  River, 
near  Athabaska  Pass.  The  splendid  series  collected  by  Loring  in 
1895  and  1896  in  the  region  just  north  of  the  type  locality  (from 
Henry  House  to  Muskeg  Creek  and  Smoky  River)  has  been  of  the 
greatest  value  in  studying  the  group  and  working  out  the  pelage 
changes  and  range  of  variation  in  the  species. 

Intergradation  with  lutescens  is  indicated  'by  several  specimens 
from  Rocky  Pass,  Alberta,  which  are  somewhat  paler  than  typical 
princeps. 

Although  no  specimens  are  available  from  the  region  between 
Jasper  Park,  Alberta,  and  northern  Montana,  doubtless  the  species 
ranges  continuously  along  the  higher  ridges.  Specimens  from  Glac- 
ier Park  and  the  Bitterroot  Mountains,  Montana,  are  typical.  A 
series  from  Mullan,  Idaho,  and  Silver  [  =  Saltese],  Montana,  agrees 
with  princeps  in  external  characters,  but  several  of  the  skulls  show 
approach  to  lemhi  in  the  narrowing  of  the  nasals  posteriorly.  A  series 
from  Thompson  Pass,  Montana,  approaches  cuppes  in  having  the 
underparts  somewhat  darker  (more  ochraceous). 

The  northern  limits  of  the  range  of  princeps  are  not  definitely 
known.  S.  Prescott  Fay  and  C.  R.  Cross,  jr.,  in  August,  1914,  noted 
the  species  near  the  head  of  Wapiti  River,  and  again  near  the  head 
of  the  east  branch  of  South  Pine  River,  northern  British  Columbia, 
but  reported  the  animals  very  scarce  in  this  region,  and  did  not  secure 
specimens. 

Richardson  mentions27  having  received  specimens  obtained  by  Mr. 
MacPherson  from  the  " River  of  the  Mountains"  [  =  Liard] — prob- 
ably from  the  vicinity  of  Fort  Nelson — and  later  B.  R.  Ross  reported 
the  species  common  among  the  mountain  ranges  of  the  Liard  River.28 
No  specimens  have  been  examined,  however,  from  this  region  and 
whether  the  animal  found  there  is  princeps  or  not  is  quite  uncertain. 

Specimens  examined. — Total  number,  108,  as  follows: 

Alberta:  Grand  Cache  River  (about  60  miles  north  of  Jasper  House),  1;  Henry 
House  (vicinity),  40;  Jasper  Park.  6;29  Muskeg  Creek  (about  60  miles  north 
of  Jasper  House),  6;  Rocky  Pass,  3; 30  Rocky  River  (near  head),  1;  30 Smoky  River 
(near  head),  4;  Smoky  Valley  (about  50  miles  north  of  Jasper  House),  1. 

British  Columbia:  Mount  Evans  (near  Cranbrook),  l;31  Spillamacheen  River,  2. 30 

Idaho:  Bitterroot  Mountains  (west  of  Hamilton,  Montana \  2;  Mullan,  8;  Thompson 
Pass,  12. 

Montana:  Bass  Creek,  Bitterroot  Mountains  (northwest  of  Stevensville),  7;  Corvallis 
(mountains  15  miles  east),  ] ;  Lake  Como,  Bitterroot  Mountains,  1 ;  Mount  Lo  Lo, 
1;  Piegan  Pass,  Glacier  National  Park,  1;  Silver  [=Saltese],  4;  St.  Marys  Lake 
[mountains  near],  1;  Sweeny  Creek  (7  miles  west  of  Florence),  5. 

*  Richardson,  John,  Fauna  Boreali-Amer.,  vol.  1,  p.  227, 1829. 

"Preble,  E.  A.,  North  Amer.  Fauna  No.  27,  p.  198, 1908. 

"Richardson,  John,  Loc.  cit. 

2«Canad.  Nat.  and  Oeol.,  vol.  7,  p.  141, 1862. 

"Five  in  collection  Victoria  Mem.  Has.;  one  in  collection  Amer.  Mus.  Nat.  Hist. 

'"Collection  Amer.  Mus.  Nat.  Hist. 

"Collection  Provincial  Mus.,  Victoria,  British  Columbia. 


1924.]  OCHOTONA   PRINCEPS.  15 

OCHOTONA  PRINCEPS  LUTESCENS  Howell. 

Alberta   Pika. 

(PL  Ill,  fig.  7;  PI.  V,  fig.  7.) 

Ochotona  princeps  lutescens  Howell,  Proc.  Biol.  Soc  Washington,  vol.  32,  p.  105,  May 
20,  1919. 

Type. — Collected  on  Mount  Inglesmaldie,  near  Banff,  Alberta,  Can- 
ada (at  about  8,000  feet  altitude) ,  July  26,1899,  by  G.  F.  Dippie;  s 
adult,  skin  and  skull;  No.  108650,  U.  S.  National  Museum,  Biological 
Survey  collection;  original  number,  193  (2973,  X  Catalogue). 

Geographic  distribution. — Mount  Inglesmaldie  (near  Banff)  and 
other  mountains  in  Canadian  National  Park,  Alberta,  south  to  Mount 
Forget-me-not  (50  to  75  miles  southwest  of  Calgary).      (Fig.  2,  p.  11.) 

Characters. — Similar  to  princeps,  but  smaller;  coloration  decidedly 
paler,  both  above  and  below. 

Color. — Summer  pelage:  Upperparts  mixed  pinkish  buff  and  fuscous,, 
darkest  in  the  middle  of  the  back,  shading  to  clear  pinkish  buff  along 
sides;  ears  hair-brown,  broadly  margined  with  light  buff  and  clothed 
inside  with  buffy  white  hairs;  feet  creamy  white,  faintly  washed  with 
pinkish  buff;  soles  drab;  palms  pale  olive-buff;  underparts  creamy 
white,  faintly  washed  with  pale  pinkish  buff,  chiefly  along  median  line. 
Worn  winter  pelage  (Mount  Forget-me-not,  Alberta,July  10) :  General 
tone  of  upperparts  smoke-gray,  faintly  washed  with  fuscous;  sides 
washed  with  pinkish  buff;  ears  and  feet  as  in  summer  pelage.  Fresh 
winter  pelage:  Not  represented. 

Skull. — Similar  to  princeps,  but  smaller,  with  shorter  nasals. 

Measurements. — Average  of  8  adults  (4  males,  4  females)  from  Can- 
more,  Alberta:  Total  length,  177  (170-188);  hind  foot,  29.4  (26-31). 
Skull:  Average  of  5  adult  males  from  vicinity  of  type  locality:  Occi- 
pi to-nasal  length,  41.6  (39.3-43.4);  zygomatic  breadth,  20.9  (19.7- 
21.7);  breadth  of  braincase,  17.3  (16.5-18);  interorbital  breadth,  5.4 
(5.1-6);  width  of  palatal  bridge,  1.9  (1.7-2.2):  length  of  nasals,  13.5 
(12—14.1) :  Average  of  3  females  from  same  region:  Occipito-nasal 
length,  41.9  (41.1-43.3) ;  zygomatic  breadth,  20.3  (20-20.5);  breadth 
of  braincase,  17.3  (16.3-18.8);  interorbital  breadth,  5.6  (5.5-5.8); 
width  of  palatal  bridge,  1.8  (1.6-1.9);  length  of  nasals,  12.9  (12.3- 
13.4). 

Remarks. — The  Alberta  pika  is  a  well-marked  race,  characterized 
by  small  size  and  pale  coloration;  it  apparently  has  a  rather  restricted 
range  on  the  easterly*  side  of  the  main  chain  of  the  Rocky  Mountains, 
but  more  extensive  collections  are  needed  to  show  its  northern  and 
southern  limits.  Three  specimens  in  worn  pelage  from  Ptarmigan 
Lake  are  apparently  intermediate  between  lutescens  and  princeps .  A 
series  from  Canmore,  Alberta,  is  intermediate  between  tutesrs,:  s  and 
levis;  in  coloration  above  the  specimens  agree  closely  with  lews,  but 
the  underparts  are  less  buffy  and  the  soles  and  palms  paler,  as  in 
lutescens,  to  which  form  they  are  referred. 

Specimens  examined. — Total  number,  50,  as  follows: 

Alberta:  Banff,  2;32  Canmore,  17;  Mount  Forget-me-not  [50  to  75  mileB  southwest  of 
Calgary],  3;32  Mount  Inglesmaldie  (near  Banff),  25; 33  Ptarmigan  Lake,3.34 

^Collection  Victoria  Mem.  Mus. 

»Ten  in  collection  Field  Mus.  Nat.  Hist.;  6 in  collection  Mus.  Comp.  Zool.;  5  in  collection  Amer.  Mus, 
Nat.  Hist. 

"Collection  Amer.  Mus.  Nat.  Hist. 


10  NORTH   AMERICAN   FAUNA.  [No.  47 

OCHOTONA  PRINCEPS  LEVIS  Hollister. 

HOLLISTER  PlKA. 

Ochotona  levis  Hollister,  Proc.  Biol.  Soc.  Washington,  vol.  25,  p.  57,  April  13,  1912. 

Type. — Collected  at  Chief  Mountain  [Waterton]  Lake,  Alberta, 
August  24,  1874,  by  Dr.  Elliott  Coues;  9  adult,  skin  and  skull; 
No.  ||g%i|U.  S.  National  Museum ;  original  number,  4593. 

Geographic  distribution. — Mountains  of  southern  Alberta  and  Mon- 
tana (east  of  the  main  divide)  from  Waterton  Lake  to  the  Belt 
Mountains.     (Fig.  2,  p.  11)       . 

Characters. — Smaller  and  paler  than  princeps,  similar  to  lutescens, 
but  darker  (more  buffy)  both  above  and  below;  soles  and  palms 
darker. 

Color. — Summer  pelage:  Upperparts  light  pinkish  cinnamon,  mixed 
with  fuscous,  shading  on  lower  sides  to  clear  pinkish  cinnamon  or 
light  pinkish  cinnamon;  ears  dark  hair-brown  or  chaetura  drab, 
margined  with  light  buff;  feet  pinkish  buff  or  cinnamon-buff;  soles 
hair-brown  or  chaetura  drab;  palms  light  drab:  underparts  whitish, 
partially  or  wholly  suffused  with  pinkish  buff,  or  light  pinkish  cinna- 
mon.     Winter  pelage:  Not  represented. 

Skull. —  Closely  similar  to  lutescens;  smaller  than  princeps. 

MeasureTnents. —  Average  of  3  adult  males  from  type  locality:  Total 
length,  183  (180-190);  hind  foot,  30.5.  Adult  $  (type),  168;  28. 
Average  of  7  adult  males  from  Belt  Mountains,  Montana:  173 
(168-180),  30  (29-31).  Skull:  Adult  male  from  Little  Belt  Moun- 
tains, Montana:  Occipito-nasal  length,  41.9;  zygomatic  breadth,  21.9; 
breadth  of  braincase,  17.4;  interorbital  breadth,  6.1;  width  of  palatal 
bridge,  1.9;  length  of  nasals,  14.2.  Adult  female  (type):  Occipito- 
nasal  length,  41.8;  zygomatic  breadth,  20.5;  breadth  of  braincase, 
17.2;  interorbital  breadth,  5.2;  width  of  palatal  bridge,  1.5;  length 
of  nasals,  14.2. 

Remarks. — Intermediate  in  color  between  princeps  and  lutescens, 
lebis  geographically  occupies  a  region  segregated  from  the  range  of 
lutescens  but  adjoining  the  range  of  princeps.  The  specimens  from 
the  type  locality  are  mostly  in  worn  summer  pelage  and  the  series  is 
insufficient  to  snow  the  characters  clearly.  A  series  from  the  Belt 
Mountains,  Montana,  in  fresh  summer  pelage  agrees  closely  with  the 
type  series  and  differs  markedly  from  specimens  of  princeps  from  the 
main  divide  in  Montana  (Piegan  Pass,  Glacier  National  Park,  and 
Bitterroot  Mountains). 

Specimens  examined. — Total  number,  15,  as  follows: 

Alberta:  Chief  Mountain  [Waterton]  Lake,  5. 
Montana:  Belt  Mountains,  1;  Little  Belt  Mountains,  9. 

OCHOTONA  PRINCEPS  LEMHI  Howell. 

Lemhi  Pika. 

(PL  Ill,  %•  9;  PI.  V,  fig.  9.) 

Ochotona  uinta  lemhi  Howell,  Proc.  Biol.  Soc.  Washington,  vol.  32,  p.  106,  Mav  20, 
L919. 

Typ\ — Collected  on  the  Lemhi  Mountains  (10  miles  west  of  Junc- 
tion), Idaho,  August  19,  1890,  by  Clark  P.  Streator;  9  adult,  skin  and 
skull;  No.  flHHrn  U.  S.  National  Museum,  Biological  Survey  collec- 
tion; original  number,  174. 


1924.]  OCHOTONA   PRINCEPS.  17 

Geographic  distribution. — Mountains  of  south-central  Idaho — the 
Lemhi,  Lost  River,  Salmon  River,  and  Sawtooth  Ranges;  eastward 
to  the  Beaverhead  Mountains,  east  of  Leadore;  northern  limits  of 
range  imperfectly  known.     (Fig.  2,    p.  11.) 

Characters. — Size  small;  colors  very  pale;  similar  in  winter  pelage 
to  princeps,  but  upperparts  paler  and  underparts  more  whitish;  in 
summer  decidedly  paler  (less  blackish)  above  and  less  buffy  beneath ; 
similar  in  summer  pelage  to  levis  but  general  tone  above  more  buffy 
and  underparts  more  whitish  or  yellowish  (less  pinkish) ;  skull  small 
and  relatively  narrow,  with  narrow  nasals. 

Color. —  Winter  pelage:  Upperparts  mixed  pinkish  buff  or  cinnamon  - 
buff  and  fuscous  or  fuscous-black,  rather  heavily  washed  in  some 
specimens,  chiefly  on  hinder  back,  with  pale  smoke-gray;  sides  pinkish 
buff  or  cinnamon-buff;  ears  dark  mouse-gray,  margined  with  buffy 
white;  hairs  on  inside  of  ears  buffy  (or  grayish  white,  tipped  with 
fuscous);  feet  grayish  white,  washed  with  pinkish  buff;  soles  hair- 
brown;  palms  grayish  white,  tinged  with  drab;  underparts  grayish 
white,  washed  with  pinkish  buff,  or  in  some  specimens  nearly  solid 
pinkish  buff.  Summer  pelage:  Upperparts  cinnamon-buff  or  pinkish 
buff,  moderately  mixed  with  fuscous;  hinder  back  and  rump  often 
washed  with  smoke-gray;  sides  cinnamon-buff  or  pinkish  buff;  ear's 
dark  mouse-gray  or  chaetura  black;  feet  whitish,  pinkish  buff,  or  deep 
cream  buff;  soles  and  palms  as  in  winter  pelage;  underparts  whitish, 
faintly  or  sometimes  heavily  washed  with  pinkish  buff  or  cream  buff. 

Molt. — The  beginning  of  the  midsummer  molt  is  shown  by  a  speci- 
men from  Elk  Summit,  Idaho,  August  4,  in  which  the  new  summer 
pelage  is  coming  in  on  the  head  and  in  a  small  spot  on  the  middle  of 
the  back.  Specimens  taken  in  the  Lemhi  Mountains  in  August  are 
in  full  summer  pelage,  some  of  them  showing  considerable  wear.  One 
taken  there  on  August  27  shows  the  fresh  winter  pelage  coming  in 
over  the  whole  of  the  fore  back;  another  from  head  of  Wood  River, 
Idaho,  September  25,  is  in  about  the  same  condition  of  molt,  while 
in  two  others  from  the  same  place,  October  5,  the  entire  dorsal  surface 
is  clothed  with  new  winter  pelage. 

Skull. — Averaging  smaller  than  in  princeps,  with  narrower  brain- 
case  and  interorbital  constriction;  nasals  narrowed  posteriorly;  inter- 
pterygoid  fossa  averaging  broader,  with  a  tendency  to  be  more 
spatulate. 

Measurements. — Average  of  10  adults  (8  males,  2  females)  from 
type  locality:  Total  length,  178.7;  hind  foot,  29.5.  Skull:  Average  of 
5  males  and  5  females35  from  Lemhi  Mountains  and  mountains  near 
Birch  Creek:  Occipito-nasal  length,  42.9;  41.4  (40.1-44.1);  zygomatic 
breadth,  21.2;  20.5  (20-22);  breadth  of  braincase,  17.9;  16.8 
(16.1-18.5) ;  interorbital  breadth,  4.9  (4.6-5.3) ;  width  of  palatal  bridge, 
2  (1.7-2.5);  length  of  nasals,  14.2;  14.1  (13.2-14.9). 

Remarks. — The  Lemhi  pika  is  a  strongly  marked  form  whose  rela- 
tionships are  not  as  yet  clearly  understood.  It  undoubtedly  iiUer- 
grades  with  princeps  somewhere  in  central  Idaho  and  probably  also 
with  ventorum  in  southwestern  Montana.  Specimens  of  princeps  from 
Mullan,  Idaho,  and  Silver  [Saltese],  Montana,  show  a  tendency  to  vary 
in  skull  characters  in  the  direction  of  lemhi.™     Two  specimens  from 

35  Averaged  separately,  the  first  measurement  in  each  case  being  that  of  the  males. 

36  As  shown  under  princeps,  p.  14. 


18  NORTH   AMERICAN   FAUNA.  [No.  47 

Seven  Devils  Mountains,  in  western  Idaho,  are  provisionally  referred 
to  lemhi;  their  skulls  are  typical  of  this  form,  but  the  hind  feet  are 
longer  and  the  coloration  of  one  of  them  is  decidedly  darker. 
Specimens  examined.— -Total  number,  40,  as  follows: 

Idaho:  Birch  Creek  [=mountains  east],  4;  Elk  Summit  (about  15  miles  southeast 
of  Warren),  2;  Ketchum,  3;37  Leadore  [=mountains  eastj,  3;  Lemhi  Moun- 
tains38 (10  miles  west  of  Junction),  18;  Little  Lost  River  mountains  (near  Birch 
Creek),  1;  Seven  Devils  Mountains,  2;  Stanley  Lake,  3;39  Wood  River  (near 
head;,  4. 

OCHOTONA  PRINCEPS  VENTORUM  Howell. 

Wyoming  Pika. 

Ochotona  uinta  ventorum  Howell,  Proc.  Biol.  Soc  Washington,  vol.  32,  p.  106,  May  20, 
1919. 

Type. — Collected  on  Fremont  Peak,  Wind  River  Mountains,  Wyo- 
ming, July  19,  1911,  by  H.  E.  Anthony;  c?  adult,  skin  and  skull;  No. 
176778,  U.  S.  National  Museum,  Biological  Survey  collection;  original 
number,  395. 

Geographic  distribution.— Wind  River,  Gros  Ventre,  Absaroka,  and 
Teton  Ranges,  Wyoming,  north  to  the  Beartooth  and  Absaroka 
Ranges  in  southern  Montana.     (Fig.  2,  p.  11.) 

CTioracters. — Similar  to  princeps,  but  coloration  in  summer  pelage 
more  buffy  and  less  varied  with  blackish;  skull  averaging  larger,  with 
longer  nasals  and  broader  interpterygoid  fossa.  Compared  with 
levis,  the  coloration  is  darker  and  more  buffy  and  the  skull  much 
larger. 

Color. —  Winter  pelage  (Gros  Ventre  Range,  near  Jackson,  Wyoming, 
September  19) :  General  tone  of  upperparts  between  drab  and  wood- 
brown,  shading  on  sides  to  light  pinkish  cinnamon;  head  and  back 
washed  with  fuscous,  front  of  face  with  clay  color,  and  rump  with 
blackish;  ears  deep  mouse-gray,  margined  with  buffy  white;  feet 
pinkish  buff;  soles  hair-brown;  palms  buffy  white,  tinged  with  light 
drab;  underparts  soiled  whitish,  faintly  washed  with  pinkish  buff. 
Summer  pelage:  Upperparts  mixed  cinnamon-buff  or  pinkish  cinna- 
mon and  fuscous-black,  the  general  tone  yellowish  wood-brown ;  sides 
pinkish  cinnamon  or  pinkish  buff;  ears  chaetura  drab,  margined  with 
buffy  white;  feet  pinkish  buff  or  pale  pinkish  buff;  soles  hair-brown; 
palms  buffy  white,  tinged  with  light  drab:  underparts  grayish  white, 
moderately  or  sometimes  heavily  washed  with  pinkish  buff  or  light 
pinkish  cinnamon. 

STcull. — Similar  to  princeps,  but  averaging  larger;  nasals  longer  and 
narrower  posteriorly;  interpterygoid  fossa  decidedly  broader  and 
usually  more  spatulate. 

Measurements. — Average  of  12  adults  (7  males,  5  females)  from 
Wind  River  Mountains,  Wyoming:  Total  length,  197  (185-216) ;  hind 
foot,  31.1  (29-33).  Slcull:  Average  of  6  adult  males  from  Wind 
River  Mountains:  Occipito-nasal  length,  45.2  (44-46.1);  zygomatic 
breadth,  22  (21.5-22.6);  breadth  of  braincase,  18  (17.3-18.5);  inter- 
orbital  breadth,  5  (4.6-5.6) ;  width  of  palatal  bridge,  2.2  (2-2.7) ;  length 
of  nasals,   15   (14.4-15.6).     Average  of  4  adult  females  from  same 

"Collection  Mus.  Comp.  Zool. 

33  Recorded  as  "  Salmon  River  Mts.  "  in  North  Amer.  Fauna  No.  5,  p.  74,  1891. 

89  Collection  Amer.  Mus.  Nat.  Hist. 


1924.]  OCHOTONA   PRINCEPS.  19 

localities:  Occipito-nasal  length,  44.5  (43.4-46.7);  zygomatic  breadth, 
21.4  (21.2-21.6);  breadth  of  braincase,  17.9  (17.3-18.5);  interorbital 
breadth,  4.8  (4.6-5.2) ;  width  of  palatal  bridge,  1.8  (1.5-2) ;  length  of 
nasals,  14.5  (13.7-15.1). 

Remarks. — This  race  resembles  princeps  rather  closely  in  coloration, 
but  is  distinguished  by  pronounced  skull  characters.  Intergradation 
between  them  is  not  definitely  shown  by  the  series  in  hand,  but  more 
material  from  extreme  southwestern  Montana  doubtless  will  establish 
beyond  question  their  subspecific  relationship. 

A  small  series  from  Emigrant  Peak,  Montana,  agree  in  color  and 
external  measurements  with  ventorum;  their  skulls,  however,  are 
somewhat  smaller  and  the  nasals  are  shorter  and  broader — characters 
indicating  an  approach  to  princeps  or  possibly  also  to  lemhi.  Speci- 
mens from  the  Absaroka  and  Beartooth  Mountains  have  somewhat 
smaller  skulls  than  the  typical  series  but  agree  with  them  in  having 
long  nasals.  A  consideraole  series  from  the  Teton  Range  is  referable 
to  this  race,  though  perhaps  shading  toward  uinta :  specimens  in 
summer  pelage  agree  well  with  the  Wind  River  series  of  ventorum  but 
have  more  buffy  hind  feet,  a  specimen  from  Teton  Pass  acquiring 
winter  pelage  (September  15)  is  light  drab  above,  more  grayish  than 
specimens  of  ventorum  from  the  Gros  Ventre  Range,  near  Jackson, 
and  not  at  all  like  specimens  of  typical  uinta  in  corresponding  pelage. 

The  form  occurring  in  the  Big  Horn  Mountains  is  provisionally 
referred  to  this  race,  but  material  in  fresh  pelage  may  show  it  to  be 
different.  Specimens  in  worn  winter  pelage  (June)  agree  closely  with 
July  specimens  in  corresponding  pelage  from  the  Wind  River  Moun- 
tains, but  are  slightly  more  buffy  on  the  sides. 

Specimens  examined. — Total  number,  77,  as  follows: 

Montana:  Beartooth  Mountains,  8;  Emigrant  Peak,  7.40 

Wyoming:  Big  Horn  Mountains  (head  of  Trappers  Creek),  7;  Fremont  Peak  (at 
11,500  feet  altitude),  9;  Gros  Ventre  Mountains,  near  Jackson,  3;  Lake  Fork, 
Wind  River  Mountains,  13;  Middle  Piney  Lake,  near  Stanley,  1;  Needle  Moun- 
tain, 4;  Pahaska  (Whirlwind  Peak),  5;  Teton  Mountains  (South  Moose  Creek, 
at  10,000  feet  altitude),  12;  Teton  Pass,  2;  Wind  River  Mountains  (12  miles 
northeast  of  Kendall,  at  11,000  feet  altitude),  3;  Yellowstone  Park,  3  (Mammoth 
Hot  Springs,  1 ;  Upper  Yellowstone,  at  Park  boundary,  2). 

OCHOTONA  PRINCEPS  UINTA  Hollister. 

Uinta  Pika. 

(PI.  Ill,  fig-  3;  PI.  V,  fig.  3.) 

Ochotona  uinta  Hollister,  Proc.  Biol.  Soc.  Washington,  vol.  25,  p.  58.  April  13,  1912- 

Type. — Collected  in  the  Uinta  Mountains,  near  head  of  east  fork 
of  Bear  River,  Utah,41  September  28,  1870,  by  the  Hayden  Expedi- 
tion; adult,  skin  only;  No.  9750,  U.  S.  National  Museum;  original 
number,  567. 

"Two  in  collection  Mus.  Comp.  Zool. 

41  The  itinerary  of  the  Hayden  Expedition  shows  that  the  party  was  camped  on  the  night  of  Septem- 
ber 26-27  on  the  divide  between  the  west  fork  of  Blacks  Fork  and' the  east  fork  of  Bear  River  at  an  alti- 
tude of  11,066  feet,  and  in  all  probability  the  type  of  O.  uinta  was  taken  near  this  camp.  The  original  label 
on  the  type  specimen  records  the  date  of  capture  rs  September  28,  but  a  study  of  the  records  of  the 
expedition  (pp.  52,  495)  shows  that  on  that  date  they  were  at  the  Sulphur  Creek  camp,  at  7,376  feet 
altitude,  probably  much  below  the  range  of  the  pikas.  On  the  label  of  another  specimen  taken  on  the 
same  trip  the  date  has  been  altered  from  September  28  to  27,  which  strengthens  the  view  that  both  speci- 
mens were  taken  at  the  higher  camp  near  timber  line  and  possibly  not  labeled  till  the  next  day  (see 
Prelim.  Kept.  U.  S.  Geol.  Surv.,  Wyoming  [=4th  Ann.  Rept.  U.  S.  Geol.  Surv.],  1871,  pp.  44-54,  495) 


20  NORTH   AMERICAN   FAUNA.  [No.  47 

Geographic  distrihution. — The  Uinta  and  Wasatch  Ranges,  in  Utah, 
and  the  Salt  River  and  Wyoming  Ranges,  in  western  Wyoming. 
(Fig.  2,  p.  11.) 

Characters. — Similar  to  ventorum,  but  coloration  in  winter  pelage 
decidedly  more  buffy  (less  grayish) ;  in  summer  pelage  more  intensely 
buffy,  both  dorsally  and  ventrally. 

Color.—  Winter  pelage:  General  tone  of  upperparts  buffy  wood- 
brown,  shading  on  sides  to  light  vinaceous -cinnamon  or  light  pinkish 
cinnamon;  dorsal  area  moderately  shaded  with  fuscous,  darkest  in 
the  median  line  and  on  top  of  head;  feet  pinkish  buff;  soles  drab  or 
hair-brown ;  palms  soiled  whitish^  ears  fuscous  or  fuscous-black,  mar- 
gined with  light  buff ;  underparts  soiled  whitish,  washed  with  pinkish 
buff  or  light  pinkish  cinnamon.  Summer  pelage  (specimen  from  head 
of  Smith  Fork,  Salt  River  Range,  Wyoming) :  General  tone  of 
upperparts  between  cinnamon-buff  and  pinkish  cinnamon,  moderately 
shaded  (chiefly  on  hinder  back)  with  fuscous-black ;  sides  pinkish  cinna- 
mon :  ears  deep  mouse-gray  or  chaetura  black  edged  with  light  buff ;  feet 
light  ochraceous-buff :  soles  hair-brown;  palms  pinkish  buff;  under- 
parts buffy  white,  washed  with  pinkish  buff  or  light  pinkish  cinna- 
mon, darkest  along  the  median  line. 

Skull. — Closely  similar  to  ventorum;  possibly  averaging  slightly 
broader,  with  broader  nasals. 

Measurements. — Adult  male  and  adult  female  from  Uinta  Moun- 
tains, Utah:  Total  length,  210;  205;  hind  foot,  29.  Average  of  3 
adult  males  from  Salt  River  Range,  Wyoming;  192;  30.3.  Skull: 
Average  of  3  adult  males  from  Uinta  Mountains,  Utah,  and  Salt 
River  Mountains,  Wyoming:42  Occipito-nasal  length,  45.3  (45.3) ;  zygo- 
matic breadth,  21.9  (22.1) ;~  breadth  of  braincase,  18.2  (17.5);  interor- 
bital  breadth,  5.1  (5.1);  width  of  palatal  bridge,  2.2  (2.4);  length  of 
nasals,  15.5  (15.7).  Average  of  5  adult  females  from  same  localities:42 
Occipito-nasal  length,  43.7  (44.2);  zygomatic  breadth,  21.3  (22); 
breadth  of  braincase,  18  (18.4);  interorbital  breadth,  5  (5.1);  width 
of  palatal  bridge,  2  (2.8) :  length  of  nasals,  14.7  (15). 

Remarks. — The  distribution  and  relationships  of  this  race,  which 
is  very  poorly  represented  in  the  material  at  hand,  are  not  accurately 
known.  The  only  specimens  available  from  the  type  region  are  3 
skins  in  fresh  winter  pelage,  without  skulls,  taken  by  the  Hayden 
Expedition,  in  1870,  and  2  skins  in  worn  winter  pelage,  with  skulls, 
taken  by  Vernon  Bailey,  in  1890.  It  is  the  most  uniformly  buffy- 
colored  race  of  all  the  American  forms.  In  the  absence  of  any  speci- 
mens in  summer  pelage  from  the  type  locality,  specimens  from  the 
Salt  River  Range,  Wyoming,  are  used  for  comparison  in  that  pelage. 
These,  however,  may  prove  to  be  intermediate  between  uinta  and 
ventorum,  as  specimens  from  farther  north,  in  the  Teton  Range,  cer- 
tainly are. 

Specimens  examined. — Total  number,  22,  as  follows: 

Utah:   Uinta  Mountains,  5  (near  head  Bear  River,  3;  near  head  Black  Fork,  2); 

Wasatch  Mountains.  I.43 
Wyoming:  La  Barge  Creek,  Salt  River  Mountains  (9,000  feet),  6;  Smith  Fork  (at 

head).  Salt  River  Mountains,  3;  Salt  River  Mountains,  7. 

"Measurements  of  one  topotype  in  parentheses. 
«No  definite  locality. 


1924.]  OCHOTONA  PEINCEPS.  21 

OCHOTONA  PRINCEPS  NEVADENSIS  Howell. 

Ruby  Mountains  Pika. 

(PI.  Ill,  fig.  6;  PI.  V,  fig.  6.) 

Ochotona  uinta  nevadensis  Howell,  Proc.  Biol.  Soc.  Washington,  vol.  32,  p.    107, 
May  20,  1919. 

Type. — Collected  in  the  Ruby  Mountains,  southwest  of  Ruby  Val- 
ley Post  Office,  Nevada  (at  10,500  feet  altitude),  June  21,  1898,  by 
Vernon  Bailey;  $  adult,  skin  and  skull;  No.  94213,  U.  S.  National 
Museum,  Biological  Survey  collection;  original  number,  6580. 

Geoqraphic  distribution. — Known  onlv  from  Ruby  Mountains, 
Nevada.     (Fig.  2,  p.  11.) 

Characters. — Similar  (in  worn  winter  pelage)  to  uinta,  but  dis- 
tinctly paler;  sides  of  nose  and  face  more  extensively  grayish. 

Color. —  Worn  winter  pelage:  Upperparts  mixed  pinkish  buff  and 
fuscous,  darkest  in  the  median  line  and  on  front  of  face;  sides  of  nose 
and  face  smoke-gray;  ears  fuscous,  margined  with  light  buff;  sides 
pinkish  buff;  feet  pale  pinkish  buff;  soles  drab;  palms  soiled  buffy 
white;  underparts  gravish  white,  moderatelv  washed  with  pinkish 
buff. 

Skull.—  Closely  similar  to  uinta. 

Measurements. — One  adult  female  (type)  and  one  adult  male 
(topotype):  Total  length,  200,  190;  hind  foot,  32,  30.  Skull  (same 
specimens):  Occipito-nasal  length,  44.1,  42;  zygomatic  breadth,  21.2, 
21.1;  breadth  of  braincase,  18,  17.9;  interorbital  breadth,  5.2,  5.4; 
width  of  palatal  bridge,  2.8,  2.5;  length  of  nasals,  14.5,  14.3. 

Remarks. — The  Ruby  Mountains  pika  is  closely  related  to  uinta  and 
is  specifically  distinct  from  cinnamomea  of  the  Beaver  Mountains,  Utah, 
and  the  Toyabe  Mountains,  Nevada.  In  the  absence  of  specimens  in 
fresh  pelage,  its  characters  can  not  be  fully  described,  and  additional 
material  may  show  greater  differences  than  are  indicated  by  the  small 
series  examined.  The  skull  of  the  type  (an  adult  female)  resembles 
the  skull  of  uinta  very  closely;  a  single  adult  male  topotype,  however, 
is  considerably  shorter  and  has  the  cranium  noticeably  elevated  in 
the  fronto-parietal  region. 

Specimens  examined. — Total  number,  3,  from  type  locality. 

,      OCHOTONA  PRINCEPS  FIGGINSI  J.  A.  Allen. 
Figgins  Pika. 
(PI.  I;  PI.  IV.,  fig.  4;  PI.  VI,  fig.  4.) 

Ochotona  figginsi  J.  A.  Allen,  Bull.  Amer.  Mue.  Nat.  Hist.,  vol.  31,  p.  103,  May  28. 

1912. 
Ochotona  saxatilis  figginsi  Henderson  and  Cockerell,  Proc.  Biol.  Soc.  Washington, 

vol.  26,  p.  127,  1913. 

Type. — Collected  on  Pagoda  Peak,  Rio  Blanco  County,  Colorado, 
October  30,  1910,  by  J.  D.  Figgins;  9  adult,  skin  and  skull;  No. 
32721,  American  Museum  of  Natural  History;  original  number,  6. 

Geographic  distribution. — Western  Colorado,  from  Gunnison  County 
north  to  eastern  Routt  County  (Hahn  Peak)  and  to  Bridger  Peak, 
Wyoming.     (Fig.  2,  p.  11.) 

Characters. — Similar  to  uinta,  but  winter  pelage  more  grayish  (less 
buffy)  and  more  varied  with  blackish;  summer  pelage  darker  (more 
blackish).  Compared  with  saxatilis,  the  size  is  smaller  and  the 
coloration  darker  and  more  vinaceous. 


22  NORTH  AMERICAN  FAUNA.  [No.  47 

Color. —  Winter  pelage  (November) :  Upperparts  mixed  vinaceous- 
cinnamon  or  light  vinaceous-cinnamon  and  fuscous,  more  or  less 
shaded  with  grayish  white,  the  general  tone  varying  from  drab  to 
cinnamon-drab;  sides  vinaceous-cinnamon  or  light  vinaceous-cinna- 
mon; ears  chaetura  drab,  margined  with  buffy  white;  feet  light  pink- 
ish cinnamon;  soles  hair-brown;  palms  soiled  whitish,  washed  with 
light  drab;  underparts  grayish  white,  faintly  or  sometimes  moder- 
ately washed  with  pinkish  cinnamon.  Specimens  in  fresh  winter 
pelage  (September  21)  are  much  more  blackish  than  November  speci- 
mens, being  mixed  vinaceous-cinnamon  and  fuscous-black  above. 
Summer  pelage  (September) :  Cldsely  similar  above  to  corresponding 
pelage  of  saxatilis;  sides  a  deeper  shade  of  vinaceous-cinnamon ;  under- 
parts moderately,  sometimes  heavily,  washed  with  light  vinaceous- 
cinnamon  or  light  pinkish  cinnamon ;  feet  light  pinkish  cinnamon  or 
cinnamon-buff. 

Skull. — Practically  identical  with  uinta;  similar  to  saxatilis  but 
averaging  smaller,  with  shorter  nasals. 

Measurements. — Average  of  9  adults  (6  males,  3  females)  from  type 
locality:  Total  length,  190.4  (184-197);  hind  foot,  30.1  (28.4-31.8). 
Skull:  Average  of  4  adult  males  from  vicinity  of  type  locality: 
Occipito-nasal  length,  43.9  (42.8-44.7);  zygomatic  breadth,  21.6 
(21-22.3) ;  breadth  of  braincase,  18.3  (17.9-19.2) ;  interorbital  breadth, 
5.1  (4.7-5.5) ;  width  of  palatal  bridge,  2.5  (2-2.8) ;  length  of  nasals, 
14.7  (14.2-15.3).  Average  of  4  adult  females  from  same  region: 
Occipito-nasal  length,  42.9  (42.1-43.7);  zygomatic  breadth,  21.1 
(20.7-21.4);  breadth  of  braincase,  17.8  (17-19);  interorbital  breadth, 
5.3  (5-5.7);  width  of  palatal  bridge,  1.9  (1.7-2);  length  of  nasals,  14.8 
(13.9-15.3). 

Remarks. — Although  separated  from  the  range  of  uinta  by  an  Upper 
Sonoran  and  Transition  area  of  considerable  width,  the  present  form 
nevertheless  is  very  closely  related  to  that  race,  and  the  specific  dif- 
ferences are  overlapped  by  individual  variation.  In  other  words, 
while  the  two  forms  are  readily  distinguishable  when  a  good  series  of 
specimens  is  available,  individual  specimens  occur  in  both  races  which 
are  difficult  to  tell  apart.  The  skulls  of  the  two  forms,  judging  by 
a  rather  meager  series,  are  practically  identical.  For  these  reasons 
it  has  seemed  best  to  unite  saxatilis  and  its  races  with  uinta  and  its 
races  as  subspecies  of  princeps. 

Specimens  from  Mount  Zirkel  are  clearly  intermediate  between 
Jigginsi  and  saxatilis;  those  in  fresh  winter  pelage  are  nearest  to  jig  ginsi 
in  coloration  but  the  skulls  agree  closely  with  those  of  saxatilis;  one 
skin  in  worn  summer  pelage  is  very  similar  to  saxatilis.  Specimens 
from  Gunnison  County  (Irwin  and  Crested  Butte)  are  likewise  inter- 
mediate, some  individuals  agreeing  with  jigginsi,  others  with  saxa- 
tilis; the  skulls  being  all  smaller  than  those  of  saxatilis. 

Specimens  examined. — Total  number,  41,  as  follows: 

Colorado:  Baldy  Mountain  (near  Mount  Zirkel),  3; 44  Crested  Butte,  2;44  Irwin, 
Gunnison  County,  2;45  Mount  Zirkel,  Larimer  County,  6;46  Pagoda  Peak,  16; 47 
Sand  Mountain  (9  miles  southwest  of  Halms  Peak  Post  Office),  1;  Trappers  Lake, 
Garfield  County,  7;44  White  House  Mountain  (near  Marble),  2.44 

Wyoming:  Bridger  Peak,  Sierra  Madre  Mountains,  2. 

44 Collection  of  E.  R.  Warren. 

«  One  in  collection  E.  R.  Warren;  one  in  collection  Mus.  Comp.  Zool. 

46  Collection  Colorado  Mus.  Nat.  Hist. 

47 Twelve  in  collection  Colorado  Mus.  Nat.  Hist.:  3  in  collection  Amer.  Mus.  Nat.  Hist. 


1924.]  OCHOTONA  PEINCEPS.  23 

OCHOTONA  PRINCEPS  SAXATILIS  Bangs. 

Colorado  Pika. 

(PI.  IV,  fig.  1;  PI.  VI,  fig.  1.) 

Ochotona  saxatilis  Bangs,  Proc.  New  England  Zool.  Club,  vol.  1,  p.  41,  June  5,  1899 . 

Type. — Collected  at  Montgomery,  near  Mount  Lincoln,  Park  County, 
Colorado,  July  27,  1871,  by  J.  A.  Allen;48  <?  adult  (skin  only);  No. 
2703,  Museum  of  Comparative  Zoology;  original  number,  945. 

Geographic  distribution. — -Easterly  mountain  ranges  of  middle 
Colorado  from  the  Medicine  Bow  Range  south  to  the  Sangre  de 
Cristo  Range;  northward  to  Medicine  Peak,  Wyoming;  westward  in 
southern  Colorado  to  the  San  Juan  Range  and  to  the  La  Sal  Moun- 
tains in  eastern  Utah;  altitudinal  range  approximately  from  9,000  to 
13,500  feet.49     (Fig.  2,  p.  11.) 

Characters. — Size  large;  similar  in  coloration  to  jigginsi,  but  colora- 
tion in  winter  pelage  paler  and  more  grayish  (less  vinaceous) ;  in  sum- 
mer pelage  slightly  paler  above  and  underparts  less  washed  with 
vinaceous.  Compared  with  ventorvm:  Coloration  in  winter  pelage 
more  grayish  in  general  tone,  the  sides  light  ochraceous-buff  instead 
of  pinkish  cinnamon,  and  the  underparts  more  heavily  washed  with 
buff;  in  summer  pelage  the  upperparts  are  usually  a  deeper  shade  of 
buff  or  cinnamon.  Compared  with  uinta,  the  winter  pelage  is  decid- 
edly more  grayish  (less  buffy). 

Color. —  Winter  pelage:  General  tone  of  upperparts  avellaneous, 
with  a  buffy  ochraceous  cast,  moderately  washed  with  fuscous-black; 
face,  and  head  between  eyes,  dull  cinnamon  buff;  sides  light  pinkish 
cinnamon,  shading  to  pinkish  buff;  ears  chaetura  black,  margined 
with  light  buff  and  clothed  inside  with  buffy- white  hairs;  feet  pink- 
ish buff;  soles  hair-brown;  palms  soiled  whitish,  tinged  with  drab; 
underparts  soiled  whitish,  moderately  washed  with  pinkish  buff. 
Summer  pelage:  Upperparts  pinkish  cinnamon  or  light  pinkish  cinna- 
mon mixed  with  fuscous,  the  cinnamon  shades  usually  prevailing; 
sides  paler,  shading  along  line  of  belly  to  light  pinkish  cinnamon  or 
pinkish  buff;  ears  chaetura  drab,  margined  with  light  buff  and  clothed 
on  inside  with  buffy-white  hairs;  front  feet  pinkish  cinnamon  or 
light  pinkish  cinnamon,  the  palms  soiled  whitish;  hind  feet  light 
pinkish  cinnamon  varying  to  buffy  white,  the  soles  hair-brown; 
underparts  whitish,  washed  with  pinkish  buff  or  pale  cinnamon-buff. 

Molt. — In  a  series  of  24  adults  from  Mount  Lincoln,  Colorado, 
taken  July  20,  1871,  the  midsummer  molt  is  well  advanced,  and  in 
about  half  the  specimens  it  is  nearly  or  quite  complete;  in  several  it 
is  just  beginning  on  the  head,  shoulders,  and  fore  back;  2  adult 
females  in  the  series  show  no  signs  of  molting,  apparently  having 
carried  the  winter  pelage  to  this  date  with  relatively  little  wear;  2 
young  individuals  about  two-thirds  grown,  taken  the  same  da}-, 
also  show  no  sign  of  molting  but  have  a  moderately  full,  uniform 
pelage,  similar  to  that  of  the  adults,  but  duller  in  color. 

In  a  specimen  ( <$  adult)  from  Geneva  Park,  Colorado,  August  31, 
1910,  the  midsummer  molt  has  progressed  on  the  back  about  three- 
fourths  of  the  distance  from  nose  to  tail,  but  on  the  sides  less  than 
half  that  distance,  the  old,  gray  winter  fur  showing  on  rump  in 
strong  contrast  to  the  fresh  ochraceous  summer  coat. 

*Cf.  Allen,  Bull.  Amer.  Mus.  Nat.  Hist.,  vol.  31,  p.  103  (footnote),  1912. 
»  Ci.  Cary,  North  Amer   Fauna  No.  33,  p.  151,  1911. 


24  NORTH   AMERICAN   FAUNA.  [No.  47 

A  specimen  from  Colorado  without  date  shows  the  midsummer 
molt  beginning  on  the  face  and  also  in  isolated  patches  on  the  nape, 
shoulders,  and  back. 

The  beginnings  of  the  fall  molt  are  not  well  shown  by  the  mate- 
rial in  hand;  3  specimens  from  Mount  Lincoln,  taken  September  26 
and  28,  1914,  have  practically  completed  the  molt  on  the  upperparts, 
though  not  on  the  belly;  the  molting  line  may  be  plainly  seen  on  the 
lower  sides. 

Specimens  from  Geneva  Park,  taken  in  November,  have  a  longer, 
fuller  pelage  than  September  specimens  and  the  coloration  is  more 
grayish — the  fuscous  or  blackish  tints  on  the  back  and  the  ochraceous 
on  the  head  and  face  decidedly  less  prominent. 

Skull. — Similar  to  jigginsi,  uinta,  and  vrntorum,  but  averaging 
larger;  nasals  long  and  rather  narrow  posteriorly. 

Measurements. — Average  of  10  adults  (5  males,  5  females)  from 
Mount  Lincoln  region  and  the  Front  Range  (Corona  and  Mount 
Kelso):  Total  length,  195  (177-202);  hind  foot,  32.2  (32-33).  Skull: 
Average  of  6  adult  males  from  the  Front  Range  and  Mount  Lincoln : 
Occipito-nasal  length,  45.5  (44.6-48.1);  zygomatic  breadth,  22  (21.3- 
23.2);  breadth  of  braincase.  18.8  (17.9-19.6);  interorbital  breadth,  5.5 
(5-5.9);  width  of  palatal  bridge,  2.4  (2.1-2.8) ;  length  of  nasals,  15.3 
,'  14.9-16.3) .  Average  of  8  adult  females  from  same  localities:  Occipito- 
nasal  length,  45.3  (44.2-47.3);  zygomatic  breadth,  21.7  (21.1-22.4); 
breadth  of  braincase,  18.5  (17.8-19.3) ;  interorbital  breadth,  5.5  (5.2- 
5.9);  width  of  palatal  bridge,  2.2  (1.8-2.8);  length  of  nasals,  15.7 
(14.9-17). 

Remarks. — The  Colorado  pika  is  the  largest  member  of  the  genus, 
and  is  easily  recognized  either  by  the  large  skull  with  long  nasals  or 
by  the  general  buffy-grayish  coloration.  It  lias  an  extensive  range 
on  the  high  mountains  of  Colorado,  intergrading  -with  Jigginsi  on  the 
westward  and  with  incana  and  nigrescens  on  the  southward. 

A  large  series  from  Silverton,  Colorado,  in  winter  pelage,  although 
practically  t}''pical  in  external  characters,  shows  intergradation  with 
nigrescens  in  cranial  characters,  some  skulls  being  decidedly  smaller 
than  others.  The  average  of  the  total  length  is  about  5  per  cent 
shorter,  but  the  nasals  are  practically  the  same  length  as  those  of 
saxatilis. 

Specimens  examined. — Total  number,  160,  as  follows: 

Colorado:  Alma,  Park  County,  13; 50  Bald  Mountain  [near  Ward,  Boulder  County],  1; 
Berthoud  Pass,  l;51  Boreas  Pass,  Summit  County,  5;52  Boulder  County,  ll;83 
Corona  [near  Arrow,  Grand  County],  11; 54  Crystal  Lake  (5  miles  west  of  Lake 
City), 2;  Dixie  Lake  [near  Arrow,  Grand  County],  4; M  Estes  Park,  5;51  Geneva 
Park  (about  70  miles  west  of  Denver),  6;51  Irwin  Lakes,  1  (skull);  Longs  Peak,  5; 
Middle  Brush  Creek,  Gunnison  County,  2;55  Montgomery  (near  Mount  Lincoln), 
Park  County,  29; 55  Mount  Kelso,  1;  Mount  Lincoln,  8;57  Mount  Meeker,  4;S1 
Mount  Shavano,  3; 55  Pikes  Peak,  5; 55  Silverton,  17;  Ten  Mile  Creek,  Summit 
County,  l;55  Twin  Lakes,  3;  Ward,  3;58  no  specific  locality,  16. 

Utah:  La  Sal  Mountains,  1. 

Wyoming:  Medicine  Bow  Mountains,  2. 

4°  Seven  in  collection  of  E.  R.  Warren,  Colorado  Springs,  Colorado;  3  in  collection  Colorado  Mus.  Nat. 
Hist. 

81  Collection  Amer.  Mus.  Nat.  Hist. 

"Three  in  collection  of  E.  R.  Warren;  1  in  collection  Amer.  Mus.  Nat.  Hist.;  1  in  collection  Mus. 
Vert.  Zool. 

63  Seven  in  collection  Field  Mus.  Nat.  Hist. 

"Collection  Colorado  Mus.  Nat.  Hist. 

65  Collection  of  E.  R.  Warren. 

"Collection  Mus.  Comp.  Zool. 

67  Five  in  collection  of  E.  R.  Warren. 

58  Two  in  collection  Arner.  Mus.  Nat.  Hist. 


1924.]  OCHOTONA  PKINCEPS.  25 

OCHOTONA  PRINCEPS  INCANA  Howell. 
New  Mexico  Pika. 

Ochotona  saxatilis  incana  Howell,  Proc.  Biol.  Soc.  Washington,  vol.  32,  p.  107,  May  20, 
1919. 

Type. — Collected  on  Pecos  Baldy,  New  Mexico  (at  12,000  feet  alti- 
tude) ,  August  10,  1903,  by  Vernon  Bailey;  9  adult,  skin  and  skull; 
No.  128944,  U.  S.  National  Museum,  Biological  Survey  collection; 
original  number,  8079. 

Geographic  distribution. — High  mountains  of  northern  New  Mexico, 
from  recos  Baldy  north  to  the  Culebra  Range  and  probably  to  Sierra 
Blanca,  Colorado  (Fig.  2,  p.  11.) 

Characters. — Similar  in  size  and  coloration  to  saxatilis,  but  general 
tone  of  upperparts  in  summer  pelage  paler  and  less  tawny;  in  winter 
pelage  more  grayish  and  less  ochraceous.  Compared  with  nigrescens, 
the  upperparts  are  paler  and  more  buffy. 

Color. — -Summer  pelage:  General  tone  of  upperparts  between  cin- 
namon-buff and  pinkish  buff,  moderately  washed  with  fuscous  or 
fuscous-black;  sides  cinnamon-buff,  shading  posteriorly  to  pinkish  buff; 
ears  chaetura  black,  margined  with  buffy  white  and  blotched  on  inner 
surface  with  the  same  color;  feet  pinkish  buff,  washed  with  cinna- 
mon-buff; soles  chaetura  drab;  palms  soiled  whitish,  tinged  with  drab; 
underparts  soiled  whitish,  washed  with  pinkish  buff.  Winter  pelage 
(specimen  from  near  Twining,  October  7) :  Upperparts  nearly  uniform 
pale  smoke-gray,  faintly  tinged  on  fore  back  with  buffy  brown;  sides 
of  face  and  shoulders  washed  with  pinkish  buff  and  light  pinkish  cin- 
namon and  faintly  washed  with  fuscous;  ears  deep  mouse-gray,  mar- 
fined  with  light  buff;  hind  feet  soiled  whitish,  tinged  with  pinkish 
uff,  the  soles  hair-brown;  front  feet  pinkish  buff,  the  palms  buffy 
white;  underparts  soiled  whitish,  very  faintly  tinged  with  light  buff. 

SJcull. — Closely  similar  to  saxatilis;  nasals  averaging  slightly  shorter. 

Measurements. — Average  of  7  adults  (4  males,  3  females)  from  Pecos 
River  Mountains  and  Taos  Mountains,  New  Mex. :  Total  length,  190.5 ; 
hind  foot,  30.5.  SJcull:  Average  of  4  adult  males  from  Pecos  Baldy 
and  TruchasPeak:  Occipito-nasal  length,  45  (44.3-45.8);  zygomatic 
breadth,  22  (21.8-22.3);  breadth  of  braincase,  19  (18.8-19.5);  interor- 
bital  breadth,  5.4  (5-5.8) ;  width  of  palatal  bridge,  2.5  (2.2-2.8) ;  length 
of  nasals,  15  (14.3-15.6).  Average  of  3  adult  females  from  Pecos 
Baldy  and  Wheeler  Peak,  New  Mex. :  Occipito-nasal  length,  44.6  (43- 
46.8);  zygomatic  breadth,  21.7  (21.3-22.5);  breadth  of  braincase, 
18.8  (18.3-19.1);  interorbital  breadth,  5.4  (5.2-5.6) ;  width  of  palatal 
bridge,  2.5  (2.2-2.8);  length  of  nasals,  14.5  (13.8-15.8). 

Remarks. — Intergradation  doubtless  takes  place  in  southern  Colo- 
rado between  this  subspecies  and  its  nearest  relative,  saxatilis.  The 
characters  on  which  the  form  is  based  are  to  some  extent  overlap- 
ping, but  the  differences  are  apparent  when  series  in  summer  pelage 
are  compared. 

Specimens  examined. — -Total  number,  13,  as  follows: 

Colorado:  Medano  Creek  (head),  Saguache  County,  l.59 

New  Mexico:    Culebra  Mountain,  1;   Pecos  Baldy,  5;   Truchas  Peak,  4;    Twining 
[=  Wheeler  Peak],  2. 

"Collection  of  E.  R.  Warren. 


26  NORTH   AMERICAN   FAUNA.  [No.  47 

OCHOTONA  PRINCEPS  NIGRESCENS  Bailey. 

Jemez  Mountains  Pika. 

(PI.  IV,  fig.  7;  PI.  VI,  fig.  7.) 

Ochotona  nigrescens,  Bailey,  Proc.  Biol.  Soc.  Washington,  vol.  26,  p.  133,  1913. 

Type. — Collected  in  the  Jemez  Mountains,  New  Mexico  (at  10,000 
feet  altitude),  August  28,  1906,  byVernon  Bailey;  s  adult,  skin  and 
skull;  No.  147976,  U.  S.  National  Museum  (Biological  Survey  collec- 
tion) ;  original  number,  8447. 

Geographic  distribution. — Jemez  Mountains,  New  Mexico,  north  to 
the  southern  end  of  the  San  Juan  Mountains,  Colorado.  (Fig.  2, 
p.  11.) 

Characters. — Similar  in  summer  pelage  to  saxatilis,  but  upperparts 
darker  and  more  grayish  (less  buffy).  Compared  with  jigginsi:  Color 
in  summer  pelage  decidedly  more  grayish  (less  brownish) ;  skull  about 
the  size  of  that  of  Jigginsi,  smaller  than  in  saxatilis  or  incana. 

Color. — Summer  pelage  (August  28) :  Head,  shoulders,  and  fore 
back  mixed  fuscous  and  pinkish  cinnamon,  shaded  with  light  buff; 
hinder  back  smoke-gray,  sparingly  mixed  with  fuscous;  sides  of  face 
and  neck  washed  with  pinkish  cinnamon  and  sides  of  body  with  light 
pinkish  cinnamon;  ears  blackish  on  both  surfaces,  narrowly  margined 
with  light  buff  and  with  a  patch  of  grayish  white  on  inner  surface; 
feet  light  pinkish  buff;  soles  fuscous;  palms  soiled  whitish,  washed 
with  light  drab ;  underparts  grayish  white,  washed  with  light  pinkish 
cinnamon  (not  cinnamon-brown,  as  in  original  description). 

Shall. — Very  similar  to  that  of  jigginsi,  but  slightly  smaller ;  decid- 
edly smaller  than  that  of  saxatilis  or  of  incana. 

Measurements. — Three  adults  (2  males,  1  female)  from  type  locality 
each  measured:  Total  length,  200;  hind  foot,  30.  STcull.  Type: 
Occipito-nasal  length,  43.3;  zygomatic  breadth,  20.9;  breadth  of  brain- 
case,  18;  interorbital  breadth,  5.3;  width  of  palatal  bridge,  2.1 ;  length 
of  nasals,  14.3. 

Remarks. — In  its  typical  form  this  race  occupies  the  isolated  moun- 
tain area  of  the  Jemez  Mountains  in  northern  New  Mexico.  A  series 
from  Navajo  Peaks — a  spur  of  the  San  Juan  Range  in  extreme  south- 
ern Colorado — is  provisionally  referred  to  nigrescens,  though  appar- 
ently it  is  about  as  close  in  characters  to  saxatilis.  In  color  these 
specimens  average  darker  above  than  saxatilis  but  are  not  so  gray  as 
nigrescens;  some  of  the  skulls  are  nearly  typical  of  nigrescens,  while 
others  agree  well  with  saxatilis.  The  typical  winter  pelage  is  unrepre- 
sented; intermediate  specimens  from  Osier,  Colorado,  taken  late  in 
July,  still  retain  some  of  the  worn  winter  pelage,  which  averages  some- 
what darker  than  that  of  typical  saxatilis. 

Specimens  examined. — Total  number,  20,  as  follows: 

Colorado:    Cumbres,  3;    Navajo    Peaks,  Archuleta  County,  4; 60  Osier,  4;60  Upper 

Navajo  River,  6. 6" 
New  Mexico:  Jemez  Mountains,  3. 

M  Collection  Colorado  Mus.  Nat.  Hist. 


1924.]  OCHOTONA   PRINCEPS.  27 

OCHOTONA  PRINCEPS  CUPPES  Bangs. 

Bangs  Pika. 

(PL  III,  fig.  4;  PI.  V,  fig.  4.) 

Ochotona  cuppes  Bangs,  Proc.  New  England  Zool.  Club,  vol.  1,  p.  40,  June  5,  1899. 

Type. — Collected  on  the  Monashee  Divide,  Gold  Range,  British 
Columbia  (at  4,000  feet  altitude),  August  2,  1897,  by  Allan  Brooks; 
S  subadult,  skin  and  skull;  No.  7389,  Museum  of  Comparative 
Zoology;  original  number,  1030. 

Geographic  distribution. — Southeastern  British  Columbia,  northeast- 
ern Washington,  and  northern  Idaho,  from  Glacier  south  to  Cabinet 
Mountains;  west  to  the  Gold  Range,  British  Columbia;  exact  limits 
unknown.     (Fig.  2.  p.  11.) 

Characters. — Similar  to  princeps  but  coloration  darker,  the  upper- 
parts  and  sides  (in  summer  pelage)  suffused  with  ochraceous-tawny 
instead  of  cinnamon-buff,  the  underparts  pinkish  cinnamon  instead  of 
pinkish  buff. 

Color. —  Type  (acquiring summer  pelage):  Head,  shoulders,  and  fore 
back  mixed  ochraceous-tawny  and  fuscous;  hinder  back  and  sides 
(still  retaining  old  worn  pelage)  mixed  pinkish  cinnamon  and  light 
buff,  with  scattering  grayish  hairs  on  dorsal  surface;  sides  of  neck 
ochraceous-tawny;  ears  chaetura  drab,  broadly  edged  with  buffy  white; 
hind  feet  pinkish  buff,  the  soles  chaetura  drab;  front  feet  cinnamon- 
buff,  the  palms  buffy  white,  tinged  with  drab;  underparts  pinkish 
cinnamon,  becoming  ochraceous-tawny  on  throat. 

STcull. — Not  appreciably  different  from  princeps. 

Measurements. — (Subadult  male,  type) :  Total  length,  190,  hind  foot, 
30.  STcull:  (Type  and  an  old  male  from  Glacier,  British  Columbia) : 
Occipito-nasal  length,  43.4,  46;  zygomatic  breadth,  21.9,  22.5;  breadth 
of  braincase,  19, 18.3;  interorbital  breadth,  5;  width  of  palatal  bridge, 
2.3,  3.5;  length  of  nasals,  13.5,  15.4. 

Remarks. — Bangs  pika  is  a  dark  form  of  princeps  occupying,  prob- 
ably, most  of  southeastern  British  Columbia.  It  is  poorly  represented 
in  collections,  and  therefore  its  characters  are  imperfectly  known. 
Five  specimens  from  Glacier,  British  Columbia,  agree  in  coloration 
with  the  type  of  cuppes;  only  one  adult  skull  is  available  from  that 
locality,  however,  and  it  is  considerably  larger  than  the  type — about 
equaling  the  largest  skulls  of  princeps.  Three  specimens  from  Ross- 
land,  British  Columbia,  have  somewhat  lighter  underparts  than  typical 
cuppes. 

A  series  of  5  adults  in  fresh  winter  pelage  from  Cabinet  Mountains, 
Idaho,  is  provisionally  referred  to  this  race,  but  additional  material 
in  winter  pelage  from  the  type  region  of  cuppes  may  show  the  Idaho 
specimens  to  oe  a  recognizable  race.  In  color  these  specimens  are 
decidedly  darker  than  specimens  of  princeps  in  the  same  pelage;81 
the  skulls,  compared  with  those  of  cuppes,  are  relatively  shorter 
and  broader. 

61  Upperparts  mixed  pinkish  cinnamon  and  fuscous  or  fuscous-black,  sometimes  washed  with  smoke-gray ; 
underparts  pate  pinkish  buff;  more  or  less  washed  with  cinnamon-buff;  soles  dark  hair-brown  or  fuscous; 
ears  and  feet  as  in  princeps. 


28  NOETH   AMERICAN"   FAUNA.  [No.  47 

A  series  from  Nelson,  British  Columbia,  agrees  well  in  cranial  char- 
acters with  typical  cuppes,  though  one  specimen  in  the  series  has  a 
skull  like  the  Cabinet  Mountains  series;  the  only  one  in  summer  pelage 
is  somewhat  paler  than  cuppes  and  not  appreciably  different  from 
princeps,  but  others  in  winter  pelage  are  darker  than  princeps  in  the 
same  pelage  and  almost  as  dark  as  the  Cabinet  Mountains  specimens. 

Specimens  examined. — Total  number,  30,  as  follows: 

British  Columbia:  Glacier,  6;62  Monashee  Divide,  Gold  Range,  4;63  Nelson  (Toad 

Mountain,  6  miles  south),  10,  Rossland,  3;M. 
Idaho:  Cabinet  Mountains,  5.  / 

Washington:  Sullivan  Lake,  Pend  Oreille  County,  2. 

OCHOTONA  PRINCEPS  FENISEX  Osgood. 

ASHNOLA   PlKA. 

(PI.  Ill,  fig.  5;  PI.  V,  fig.  5.) 

Lagomys  minimus  Lord,  Proc.  Zool.  Soc.  London,  1863,  p.  98  (not  Lagomys  minimus 

Schinz). 
Ochotona  minimus  Elliot,  Field  Columbian  Mus.,  Zool.  Ser.,  vol.  2,  p.  269,  1901  (not 

Lagomys  minimus  Schinz). 
Ochotona  fenisex  Osgood,  Proc.  Biol.  Soc.  Washington,  vol.  26,  p.  80,  March  22,  1913 

(new  name  for  Lagomys  minimus  Lord). 

Type. — Collected  on  "Ptarmigan  Hill,"  near  head  of  Ashnola  River, 
Cascade  Range,  British  Columbia,65  in  early  fall,  1860  (?),  by  J.  K. 
Lord;  sex  unknown;  skin  No.  62.12.30.11;  skull,  No.  1443a  (osteo- 
logical  collection) ,  British  Museum.6" 

Geographic  distribution. — Interior  mountain  ranges  of  northern  Wash- 
ington and  southern  British  Columbia  from  vicinity  of  Wenatchee, 
Washington,  north  to  the  upper  end  of  Okanagan  Lake.  (Fig.  2, 
p.  11.) 

Characters. — Similar  to  cuppes,  but  coloration  in  summer  pelage  paler 
and  less  ochraceous,  especially  on  head,  shoulders,  and  underparts; 
hind  feet  more  whitish  (less  buffy);  sides  of  nose  washed  with  gray; 
very  similar  to  cuppes  in  winter  pelage,  but  hind  feet  and  underparts 
paler  (less  buffy).  Skull  similar  to  that  of  cuppes,  but  averaging 
slightly  smaller,  with  shorter  nasals. 

Color. —  Winter  pelage  (Okanagan,  British  Columbia,  November) : 
Upperparts  mixea  fuscous  and  light  pinkish  cinnamon,  the  general 
tone  varying  between  hair-brown  and  wood-brown;  sides  pinkish 
cinnamon  or  light  pinkish  cinnamon,  more  or  less  clouded  with  fus- 
cous; sides  of  nose  smoke-gray  or  pale  smoke-gray;  sides  of  face  some- 
times tinged  with  grayish;  ears  chaetura  black,  margined  with  buffy 
white;  feet  white,  the  fore  feet  washed  with  pinkish  buff;  soles  hair- 
brown  or  chaetura  drab;  palms  soiled  whitish,  washed  with  drab; 
underparts  grayish  white,  washed  with  pinkish  buff  or  light  pinkish 
cinnamon.  Specimens  in  fresh  fall  pelage  (Okanagan,  British  Col- 
umbia, September  30)  differ  from  November  specimens  in  having  the 
upperparts  a  paler  shade  of  pinkish  cinnamon  or  light  buff,  rather 
heavily  washed  on  hinder  back  with  blackish,  and  the  underparts 

*>  Three  in  collection  Acad.  Nat.  Sci.  Philadelphia. 

m  Collection  Mus.  Comp.  Zool. 

"Collection  Victoria  Mem.  Mus. 

S5Cf.  Lord,  J.  K.,  Naturalist  in  British  Columbia,  vol.  1,  p.  321,  1866. 

"Selected  by  Oldfleld  Thomas  as  lectotype  from  3  specimens  collected  by  Lord. 


1924.]  OCHOTONA   PRINCEPS.  29 

more  heavily  washed  with  light  pinkish  cinnamon,  becoming  almost 
pinkish  cinnamon  on  chest.  Summer  pelage  (Bald  Mountain,  Wash- 
ington, September  7) :  Upperparts  mixed  pinkish  cinnamon  and  fus- 
cous, the  general  tone  more  tawny  and  less  drabby  than  in  winter 
pelage;  sides  pinkish  cinnamon  or  light  pinkish  cinnamon. 

Molt. — A  specimen  (  <$  subadult)  fromTulameen,  British  Columbia, 
June  11,  shows  the  midsummer  molt  just  beginning  in  a  small  patch 
of  new  hair  on  top  of  the  head;  another  (  9  adult)  from  Okanagan, 
June  14,  has  a  small  patch  of  the  new  pelage  in  the  middle  of  the  nape. 
An  adult  male  from  Horseshoe  Basin,  Washington,  July  22,  shows  the 
fresh  summer  pelage  covering  the  head,  shoulders,  and  fore  back. 

An  adult  male  from  Wenatchee,  Washington,  September  19,  appar- 
ently is  acquiring  winter  pelage,  the  new  hair  covering  most  of  the 
dorsal  area  excepting  the  rump.  In  a  specimen  from  Okanagan, 
British  Columbia,  September  30,  new  winter  pelage  covers  the  entire 
dorsal  surface  and  the  sides,  but  not  the  belly.  Specimens  from  the 
same  locality  in  May  and  June  show  the  same  pelage  as  fall  specimens, 
with  indications  of  moderate  wear. 

Skull. — Similar  to  cuppes,  but  averaging  slightly  smaller,  with 
shorter  nasals. 

Measurements. — Average  of  10  adults  (5  males,  5  females)  from 
Bald  Mountain,  Washington:  Total  length,  183  (165-190) ;  hind  foot, 
31.5  (31-32).  Skull:  Type:  Occipito-nasal  length,  42;  zygomatic 
breadth,  22;  breadth  of  braincase,  18.7;  interorbital  breadth,  6;  width 
of  palatal  bridge,  3;  length  of  nasals,  13.4.  Average  of  6  adults 
(including  the  type)  from  type  region  (Bald  Mountain  and  Horseshoe 
Basin,  Washington) :  Occipito-nasal  length,  42.9  (42-43.8);  zygomatic 
breadth,  21.6  (21-22) ;  breadth  of  braincase,  18.6  (18.2-19.2) ;  interor- 
bital breadth,  5.5  (5.1-6);  width  of  palatal  bridge,  2.6  (2.3-3) ;  length 
of  nasals,  13.8(13-14.6). 

Remarks. — This  species,  the  second  American  pika  to  be  named,  was 
originally  described  by  J.  K.  Lord  in  1863  under  the  name  Lagomys 
minimus.  The  name  was  placed  in  synonymy,  however,  by  Allen  in 
his  monograph  of  the  group  in  1877 67,  and  was  not  again  recognized 
until  revived  by  Bangs  in  1899. 68  Some  years  later,  Osgood  pointed 
out  that  the  name  Lagomys  minimus  was  preoccupied,  and  renamed 
the  species  OcJiotona  fenisex.69 

Bangs  applied  the  name  minimus  to  the  dark,  brownish-colored 
form  inhabiting  the  higher  parts  of  the  Cascade  Range,  of  which  form 
he  had  specimens  from  Hope,  Sumas,  and  the  Mount  Baker  Range, 
British  Columbia.  The  original  description,  however,  applies  rather 
to  the  paler  form  ranging  from  the  north  end  of  Okanagan  Lake  south- 
ward along  the  interior  ranges,  east  of  the  main  Cascade  Range,  to 
Wenatchee,  Washington.  In  order  definitely  to  fix  the  name,  speci- 
mens of  both  the  Cascade  form  and  the  interior  form  were  sent  to 
Oldfleld  Thomas,  who  has  compared  them  with  the  type  of  minimus 
in  the  British  Museum.  He  states  that  the  type  agrees  closely  with 
a  November  specimen  (No.  203214,  U.  S.  Nat.  Mus.)  from  Okanagan, 
British  Columbia,  and  differs  from  specimens  of  the  dark  Cascade  form. 
There  need  be  no  further  doubt,  therefore,  as  to  the  applicability  of 

"Allen,  J.  A.,  Monog.  North  Amer.  Rodentia,  Rept.  U.  S.  Geol.   Surv.  Terr.,  vol.  11,  p.  407,  1877. 
"s Bangs,  O.,  Proc.  New  England  Zool.  Club,  vol.  1,  p.  39,  1899. 
■•  Osgood,  W.  H.,  Proc.  Biol.  Soc.  Washington,  vol.  26,  p.  88,  1913. 

73502—24 3 


30  NORTH   AMERICAN    FAUNA.  [No.  47 

Lord's  name  to  the  pale  interior  form.  This  conclusion  is  furthermore 
confirmed  by  a  series  of  specimens  recently  collected  by  Taylor  and 
Cantwell  on  Bald  Mountain,  Washington — practically  at  the  type 
locality  of  fenis ex. 

A  large  series  in  worn  pelage  (September),  from  Wenatchee,  Wash- 
ington, is  apparently  intermediate  between  fenisex  and  brunnescens; 
in  coloration  the  specimens  are  nearest  to  fenisex:  the  skulls,  however, 
average  decidedly  larger  than  those  of  fenisex,  most  of  them,  indeed, 
about  equaling  those  of  brunnescens. 

Although  no  actual  intergrades  between  this  race  and  cuppes  have 
been  examined,  the  resemblance  between  the  two  forms  is  so  close 
that  it  is  considered  desirable  to  unite  them  into  one  species. 

Specimens  examined. — Total  number,  60,  as  follows: 

British.  Columbia:  Hope  (Roab  Ranch),  l;70  Okanagan,  16;  71  Okanagan  Lake 
(mountains  west  of  Lake,  2,500  feet  altitude),  5;  Tulameen,  3. 72 

Washington:  Bald  Mountain,  Okanogan  County  (head  of  Ashnola  River),  12; 
Barron,  1;  Hidden  Lakes,  1;  Horseshoe  Basin  (near  Mount  Chopaka),  l;73Pasay- 
ten  River  (west  fork,  at  mouth  of  Holman  Creek),  3;  Stehekin,  2;  Wenatchee 
(mountains  near),  15. 

OCHOTONA  PRINCEPS  BROOKSI,  stjbsp.  nov.  74 

Shuswap  Pika. 

Type. — Collected  at  Sicamous,  British  Columbia,  September  24, 
1894,  by  Clark  P.  Streator;  <?  adult,  skin  and  skull;  No.  69275,  U.  S. 
National  Museum,  Biological  Survey  collection ;  original  number,  4319. 

Geographic  distribution. — Interior  of  southern  British  Columbia, 
from  Shuswap  Lake  west  to  Mount  McLean  and  McGillivary  Creek, 
Lillooet  District;  limits  of  range  unknown.     (Fig.  2,  p.  11.) 

Characters. — Similar  in  winter  pelage  to  brunnescejis,  but  general 
tone  more  vinaceous  and  head  more  strongly  cinnamon;  in  summer 
pelage  very  much  paler.  Compared  with  fenisex:  Winter  pelage  dis- 
tinctly more  brownish  or  cinnamon  (less  grayish) ;  summer  pelage 
paler. 

Color. —  Type  (acquiring  winter  pelage):  Head  cinnamon  shaded 
with  fuscous;  rest  of  upperparts  vinaceous-cinnamon  mixed  with 
fuscous-black,  darkest  in  middle  of  back,  where  the  blackish  predomi- 
nates; sides  of  nose  faintly  tinged  with  smoke-gray;  ears  chaetura 
drab,  margined  with  buffy  white;  feet  soiled  whitish,  washed  with 
pinkish  buff;  soles  hair-brown;  palms  soiled  whitish,  washed  with 
drab;  underparts  soiled  whitish,  washed  with  pinkish  cinnamon. 
Summer  pelage  (specimens  from  McGillivary  Creek,  Lillooet  District, 
British  Columbia,  August  17):  Head  pinkish  cinnamon,  shaded  with 
fuscous ;  rest  of  upperparts  light  pinkish  cinnamon  mixed  with  fuscous, 
palest  on  shoulders  and  fore  back;  ears,  feet,  and  underparts  as  in 
winter  pelage. 

Molt. — A  specimen  from  Sicamous,  British  Columbia,  July  19,  is 
acquiring  summer  pelage,  the  new  hair  covering  the  head,  shoulders, 
fore  back,  and  patches  on  the  hinder  back  and  sides;  an  adult  female 
from  Shuswap.  British  Columbia,  September  5,  is  in  complete  summer 
pelage,  moderately  worn  on  the  hinder  back;  an  adult  female  from  the 

70  Collection  Mus.  Conip.  Zool. 

T1  Nine  in  collection  Provincial  Mus., Victoria,  British  Columbia. 

72  Collection  Provincial  Mus. 

"Collection  of  P.  G.  .Tewett. 

'4  Named  ior  Maj.  Allan  Brooks,  of  Okanagan,  British  Columbia. 


1924.]  OCHOTONA   PRINCEPS.  31 

same  locality,  same  date,  is  acquiring  winter  pelage,  the  new  hair 
covering  the  head,  shoulders,  and  more  than  half  of  the  back;  the 
hinder  back  and  sides  are  in  greatly  worn  summer  pelage. 

In  a  series  of  8  adults  from  Sicamous,  September  24-29,  the  new 
winter  pelage  covers  the  greater  part  of  the  dorsal  surface  excepting 
the  rump;  in  one  of  this  series,  however  (male,  September  24),  the  molt 
is  no  further  advanced  than  in  the  September  5  specimen  referred  to 
above. 

Skull. — Closely  similar  to  fenisex  but  nasals  averaging  slightly 
shorter;  slightly  smaller  than  in  cuppes,  with  the  interpterygoid  fossa 
more  spatulate  (the  sides  less  nearly  parallel). 

Measurements. —  Type:  Total  length,  195 ;  hind  foot,  31 .  Average  of 
9  adults  (8  males.  1  female)  from  type  locality:  Total  length,  192.4 
(180-206);  hind  foot,  31.3  (30-33).  SJcull:  Average  of  6  adult  males 
from  type  localitv:  Occipito-nasal  length,  43  (42.3-43.9);  zygomatic 
breadth,  21.5  (21-22);  breadth  of  braincase,  18.1  (17. 7-18.6);  inter- 
orbital  breadth,  5.6  (5.4-6);  width  of  palatal  bridge,  2.2  (2-2.6); 
length  of  nasals,  13.6  (12.8-14.2).  Average  of  3  adult  females  from 
Sicamous  and  Shuswap,  British  Columbia:  Occipito-nasal  length,  42.4 
(41.1-43.5) ;  zygomatic  breadth, 20.4  (19.9-20.8)  ;breadth  of  braincase, 
17.4  (17.2-17!6);  interorbital  breadth,  5.2  (5.1-5.3);  width  of  palatal 
bridge,  2.2  (2-2.6);  length  of  nasals,  13.3  (12.8-13.7). 

Remarks. — Although  occupying  an  area  contiguous  to  that  of  fenisex 
and  apparently  quite  similar  in  topographic  conditions,  this  race  is 
distinguished  by  pronounced  color  differences.  In  winter  pelage  it 
most  nearly  resembles  brunnescens  and  doubtless  intergrades  with 
that  race  where  their  ranges  meet.  From  cuppes  it  differs  not  only 
in  the  more  spatulate  interpterygoid  fossa  but  also  in  being  much  paler 
in  summer  pelage. 

Specimens  examined. — Total  number,  36,  as  follows: 

British  Columbia:  Blackwater  Lake,  Lillooet  District,  4: 75  Griffin  Lake,  2  (skulls) ; 7S 
McGillivary  Creek,  Lillooet  District,  6;  76  Mount  McLean,  Lillooet  District,  1; 76 
Shuswap,  9;  Sicamous,  14.77 

OCHOTONA  PRINCEPS  BRUNNESCENS  Howell. 

Cascade  Pika. 

(PI.  Ill,  fig.  8;  PI.  V,  fig.  8.) 

Ochotona  minimus  Bangs,  Proc.  New  England  Zool.  Club,  vol.  1,  p.  39,  1899  (not 

Lagomys  minimus  Lord). 
Ochotona  fenisex  brunnescens  Howell,  Proc.  Biol.  Soc.  Washington,  vol.  32,  p.  108, 

May  20,  1919. 

Type. — Collected  at  Keechelus,  Kittitas  County,  Washington, 
August  23,  1917,  by  George  G.  Cantwell;  male  subadult,  skin  and  skull; 
No.  227259,  U.  S.  National  Museum,  Biological  Survey  collection; 
original  number,  38  (19568,  X  catalogue). 

Geographic  distribution. — Cascade  Range,  from  southwestern  British 
Columbia  south  to  vicinity  of  Crater  Lake,  Oregon;  interrupted  appar- 
ently in  central  Oregon  (vicinity  of  Mount  Jefferson  and  Three  Sis- 
ters) by  the  range  offumosa.     (Fig.  2,  p.  11.) 

Characters. — Similar  to  fenisex,  but  upperparts  decidedly  more 
brownish  (less  grayish)  and  underparts  more  buffy. 

7i  Collection  Provincial  Mus.,  Victoria,  British  Columbia. 

"Collection  Victoria  Mem.  Mus. 

;?  Four  in  collection  Victoria  Mem.  Mus. 


32  NORTH    AMERICAN    FAUNA.  [No.  47 

Color. —  Winter  pelage  (November  specimens  from  Hope,  British 
Columbia) :  Upperparts  mixed  pinkish  cinnamon  and  fuscous,  the 
latter  color  predominating;  sides  pinkish  cinnamon  or  light  pinkish 
cinnamon;  sides  of  nose  smoke-gray,  washed  with  pinkish  cinnamon; 
ears  fuscous-black,  margined  with  light  buff;  feet  light  pinkish 
cinnamon;  soles  hair-brown;  palms  buffy  white,  tinged  with  light 
drab;  underparts  grayish  white,  moderately  or  sometimes  heavily 
washed  with  pinkish  cinnamon.  Summer  pelage  (July  and  August): 
Very  similar  to  the  winter  pelage  (often  identical)  but  general  tone 
of  upperparts  averaging  deepei^cmnamon  (less  pinkish),  the  sides 
of  the  head  and  body  frequently  rich  cinnamon,  shaded  Avith  fuscous, 
and  the  underparts  sometimes  heavily  washed  with  cinnamon;  soles 
darker — fuscous  or  chaetura  black. 

Molt. — In  an  adult  male  from  Easton,  Washington,  July  10,  fresh 
summer  pelage  covers  the  greater  part  of  the  dorsal  surface  except 
the  hinder  back,  the  molting  line  being  irregular  and  poorly  defined; 
another  from  the  same  locality,  July  19,  is  in  practically  complete 
summer  pelage.  An  adult  male  from  Stevens  Pass,  Washington,  Au- 
gust 23,  is  still  in  badly  worn  winter  pelage,  the  new  summer  coat  hav- 
ing covered  the  head,  shoulders,  and  fore  back.  An  adult  male  from 
Mount  Rainier,  August  31,  shows  new  summer  pelage  covering  the 
head,  shoulders,  and  middle  of  the  back. 

No  specimens  of  this  race  are  at  hand  showing  the  beginnings  of 
winter  pelage,  but  the  fall  molt  probably  occurs,  as  in  the  other  races 
of  the  species,  during  September.  Specimens  taken  in  June  resemble 
those  in  fresh  winter  pelage  but  show  a  moderate  amount  of  wear. 

Skull. — Similar  to  that  of  fenisex,  but  averaging  larger,  with  rela- 
tively shorter  nasals. 

Measurements. — Average  of  13  adults  (8  males,  5  females) :  Total 
length,  203  (185-215);  hind  foot,  33.4  (31-35).  Skull:  Average  of  11 
adult  males  from  Cascade  Range,  Washington:  Occipito-nasal  length, 
44  (42.3-46.2) ;  zygomatic  breadth,  21.9  (21.5-22.3) ;  breadth  of  brain- 
case,  18.7  (18-19.2):  interorbital  breadth,  5.4  (4.9-5.7);  width  of 
palatal  bridge,  2.4  (2.1-3.4) ;  length  of  nasals,  14  (13.1-14.9).  Average 
of  5  adult  females  from  same  region:  Occipito-nasal  length,  43.6  (41.8- 
44.6);  zygomatic  breadth,  21.8(21.7-21.9);  breadth  of  braincase,  18.4 
(17.7-19);  interorbital  breadth,  5.4  (5-5.5);  width  of  palatal  bridge, 
2.4  (2.2-2.7);  length  of  nasals,  14.1  (13.2-14.9). 

Remarks. — The  Cascade  pika  is  a  wide  ranging  form  which  has 
been  insufficiently  characterized  and  erroneously  identified  with 
"minimus"  of  Lord  [=  fenisex).  The  uncertainty  concerning  the 
correct  name  for  this  race  is  in  large  part  due  to  the  lack  of  material 
from  the  type  locality  of  "minimus".  Bangs  assumed  that  his  speci- 
mens from  the  Mount  Baker  Range  represented  "minimus,"  and  no 
one  since  has  questioned  this  decision.  Material  now  in  hand,  how- 
ever, from  the  immediate  vicinity  of  the  type  locality  indicates  that 
the  form  described  by  Lord  is  not  the  same  as  the  form  occupying 
the  higher  parts  of  the  Cascades.  The  latter,  therefore,  has  recently 
been  characterized  by  the  writer  under  the  name  brunnescens. 

This  race  is  subject  to  relatively  little  variation  in  color  over  its 
entire  range,  and  likewise  is  but  little  different  in  summer  and  winter 
pelage.  A  series  from  the  vicinity  of  Crater  Lake,  at  the  southern 
end  of  the  range,  averages  slightly  paler  and  grayer  than  specimens 
from  more  northern  localities,  but  the  differences  are  too  slight  to 


1924.]  OCHOTOXA    PEINCEPS.  33 

warrant  recognition  by  name.  This  series  shows  no  approach  to 
fumosa,  the  range  of  which  apparently  cuts  off  the  southern  Cascade 
colonies  of  brunnescens  from  those  of  northern  Oregon  and  Washington. 
Variation  in  size  of  the  skull  is  considerable  but  apparently  is  individ- 
ual and  not  regional. 

Specimens  examined. — Total  number,  118,  as  follows: 

British  Columbia:  Chilliwack,  6; 7S  Chilliwack  Lake,  1; 79  Hope  (Lake  HouBe),  6; M 
Mount  Baker  Range.  9;S1  Sumas,  I;60  Summit  Lake,  Cascade  Mountains,  l;80 
Tami  Hi  Creek,  3;79  Vancouver,  l.79 

Oregon:  Anna  Creek,  Mount  Mazama,  2;  Crater  Lake,  13;  Diamond  Lake,  1;  Mount 
Hood,  3. 

Washington:  Easton,  12;  Hart  Lake,  3;  Keechelus,  3;  Keechelus  Lake,  l;82  Lyman 
Lake,  Chelan  County,  2;  Mount  Adams.  3;  Mount  Aix,  2;  Mount  Index,  1;  Mount 
Rainier,  24;  Mount  St.  Helens,  8;  Natches  River  (40  miles  above  North  Yakima, 
at  2,800  feet  altitude),  1;  Snoqualmie  Pass,  9;83  Stevens  Pass,  King  County,  1; 
Twin  Sisters  Lakes  (near  Cowlitz  Pass),  1. 

OCHOTONA  PRINCEPS  FUMOSA  Howell. 
Dusky  Pika. 

Ochotona  fenisex  fumosa  Howell,  Proc.  Biol.  Soc.  Washington,  vol.  32,  p.  109, 
May  20,  1919. 

Type. — Collected  at  Permilia  Lake,  west  base  of  Mount  Jefferson, 
Oregon,  October  4,  1897,  by  J.  Alden  Loring;  male  adult,  skin  and 
skull;  No.  91144,  U.  S.  National  Museum,  Biological  Survey  col- 
lection; original  number,  4799. 

Geographic  distribution. — Western  slopes  of  the  Cascade  Mountains 
in  Oregon,  from  the  upper  Clackamas  River  south  at  least  to  Three 
Sisters;  also  mountains  near  Paulina  Lake,  Crook  County.  (Fig.  2, 
p.  11.) 

Characters. — About  the  size  of  brunnescens  with  slightly  narrower 
skull;  coloration  above  more  blackish  and  less  brownish. 

Color. — ■  Winter  pelage:  Upperparts  a  mixture  of  vinaceous-cinna- 
mon  and  black,  the  median  dorsal  area  nearly  solid  black,  shading 
on  sides  to  nearly  clear  vinaceous-cinnamon ;  sides  of  nose  dark 
smoke-gray  and  top  of  head  shaded  with  the  same;  sides  of  neck, 
beneath  ears, washed  with  cinnamon-buff;  ears  chaetura  black,  edged 
with  buffy  white;  feet  cinnamon-buff;  soles  chaetura  drab ;  palms  hair- 
brown;  underparts  soiled  whitish,  strongly  washed  along  middle  of 
belly  with  vinaceous-cinnamon,  shading  on  throat  to  pinkish  cinnamon. 
Summer  pelage  (specimens  from  Three  Sisters,  Oregon,  July  15) : 
Similar  to  the  winter  pelage,  but  upperparts  less  blackish,  less  buffy, 
and  more  strongly  shaded  with  grayish. 

Skull. — Similar  to  brunnescens  but  averaging  slightly  narrower, 
with  slightly  longer  nasals  (at  least  in  males) ;  palate  with  a  very 
small  spine  in  center  of  interpterygoid  fossa. 

Measurements. — Average  of  7  adults  (5  males,  2  females)  from  type 
locality:  Total  length,  196  (184-202) ;  hind  foot,  32.3  (31-34):  Slcull: 
Average  of  5  adult  males  from  type  locality:  Occipito-nasal  length, 
44.3  (43-45.6);  zygomatic  breadth.  21.3  (20.9-22);  breadth  of  brain- 

78 One  in  collection  r"ieid  Mus.  Nat.  Hist.;  one  in  collection  Mus.  Comp.  Zool. 

"Collection  Victoria  Mem.  Mus. 

80Collection  Mus.  Comp.  Zool. 

81  Six  in  collection  Mus.  Comp.  Zoo!.;  3  in  collection  Acad.  Nat.  Sci.  Philadelphia. 

"Collection  S.  <;.  Jewett. 

"Collection  Acad.  Nat.  Sci.  Philadelphia. 


34 


NORTH    AMERICAN    FAUNA. 


[No.  47 


case,  18.3  (18-18.7);  interorbital  breadth,  5.1  (4.7-5.3);  width  of 
palatal  bridge,  2.8  (2.4-3.2);  length  of  nasals,  14.8  (13.6-15.7). 
Average  of  2  adult  females  from  type  locality:  Occipito-nasal  length, 
43.6  (43.3-43.9) ;  zygomatic  breadth,  21.2  (21-21.4) ;  breadth  of  brain- 
case,  18.4  (17.8-18.9);  interorbital  breadth,  5.5  (5.5-5.6);  width  of 
palatal  bridge,  3.4  (2.9-4) ;  length  of  nasals,  13.5  (13.3-13.8). 

Remarks. — Closely  related  to  brunvescens,  which  occupies  the 
summits  of  the  Cascade  Range  in  Oregon,  this  dark  race  apparently 
occupies  the  western  slopes  of  the  range,  descending  in  some  localities 
to  as  low  as  900  feet  altitude.  Apparently  it  crosses  the  range  to  the 
eastward  in  the  vicinity  of  Three  Sisters  and  occupies  also  the 
mountains  around  Paulina  Lake. 

Specimens  examined. — Total  number,  16,  as  follows: 

Oregon:  Clackamas  River,  15  miles  above  Estacada  (at  about  900  feet  altitude),  l;84 
McKenzie  Bridge  (10  miles  southeast,  at  Lost  Creek  Ranger  Station),  1;  Paulina 
Lake,  1;  Permilia  Lake  (west  base  Mount  Jefferson),  9;  Three  Sisters,  4. 

OCHOTONA  COLLARIS  (Nelson). 

Collared  Pika. 

(PI.  Ill,  fig.  2;  PI.  V,  fig.  2.) 

Lagomys  collaris  Nelson,  Proc.  Biol.  Soc  Washington,  vol.  8,  p.  117,  December  21, 
1893. 

Type. — Collected  near  head  of  Tanana  River,  about  200  miles  south 
of  Fort  Yukon,  Alaska,  by  Indians,  for  E.  W.  Nelson  in  the  summer 
of  1880;  adult,  skin  and  skull;  No.  ||||fi  U.  S.  National  Museum; 
original  number,  164. 


Fig.  3. — Distribution  of  Ochotdwa  collaris. 


Geographic  distribution. — Central  and  southern  Alaska  and  Yukon, 
from  Mount  McKinley  and  the  Ogilvie  Range  south  to  Teslin  Lake, 
British  Columbia,  and  White  Pass,  Alaska.      (Fig.  3.) 


»♦  Collection  S.  G.  Jewett. 


1924.]  OCHOTONA   COLLARIS.  35 

Characters. — Similar  to  0.  princeps  princeps  in  winter  pelage  but 
coloration  decidedly  more  grayish  and  less  buffy,  both  above  and 
below;  a  more  or  less  distinct  grayish  patch  on  nape  and  shoulders; 
skull  relatively  broad,  with  broad  interpterygoid  fossa  and  large 
audital  bullae. 

Color. — Fresh  winter  pelage  (specimens  from  mountains  near  Eagle, 
July  31 -September  2) :  Upperparts  mixed  pinkish  buff  and  fuscous 
or  fuscous-black,  the  general  tone  drab  or  light  drab,  shading  to 
smoke-gray;  lower  sides  (next  to  belly)  rather  faintly  washed  with 
pinkish  buff  or  light  pinkish  buff;  an  irregular  patch  or  collar  (indis- 
tinct or  nearly  obsolete  in  some  specimens)  of  pale  smoke-gray  on 
shoulders  just  behind  ears;  ears  chaetura  black,  edged  with  light  buff 
or  buffy  white,  usually  with  a  large  blotch  of  buffy  white  or  pinkish 
cinnamon  on  inner  surface;  feet  whitish,  washed  with  pale  pinkish 
buff;  soles  hair-brown;  palms  buffy  white  or  light  drab;  underparts 
whitish,  faintly  washed  with  pale  pinkish  buff.  Worn  winter  pelage 
(specimens  from  head  of  Toklat  River,  May  3) :  Upperparts  decidedly 
more  grayish  than  in  autumn,  the  general  tone  nearly  smoke-gray, 
washed  with  buff,  and  the  head  and  shoulders  clear  smoke-gray. 

Molt. — The  available  material  strongly  indicates  that  this  species 
has  but  a  single  annual  molt,  which  extends  from  late  July  to  early 
September. 

Specimens  from  Mount  McKinley,  Alaska,  July  28,  show  the  molt 
in  progress,  the  new  pelage  appearing  in  an  irregular  patch  on  the 
middle  dorsal  area  and  a  small  patch  on  the  face  between  the  eyes, 
the  unrenewed  portions  of  the  pelage  being  greatly  worn.  The  molt 
must  in  some  cases  begin  earlier  than  this  for  in  a  specimen  from 
near  Eagle,  Alaska,  July  31,  the  pelage  is  almost  entirely  renewed, 
except  for  two  small  patches  on  each  side  of  the  rump;  in  other  cases 
it  does  not  begin  until  much  later,  as  shown  by  an  adult  female  from 
the  same  locality,  taken  September  1,  in  which  the  new  pelage 
appears  only  in  an  irregular  area  on  the  middle  of  the  back;  another 
specimen  taken  August  30  has  the  entire  back  covered  with  the  fresh 
pelage,  but  the  top  of  the  head  and  the  nape  are  still  in  a  much-worn, 
old  pelage. 

Specimens  from  the  head  of  Toklat  River  in  early  May  are  moder- 
ately worn  and  are  more  uniform  in  coloration,  the  fuscous  hairs  of 
the  fresh  pelage  having  largely  disappeared  and  the  general  tone 
become  more  grayish,  often  with  a  pronounced  buffy  tinge. 

Shull. — Similar  to  princeps  but  broader,  with  shorter  nasals  and 
larger  audital  bullae;  interpterygoid  fossa  broader  and  more  spatu- 
late,  its  sides  not  parallel,  but  expanding  slightly  near  anterior  end 
and  contracting  posteriorly. 

Measurements. — Average  of  15  adults  (10  males,  5  females)  from 
mountains  near  Eagle,  Alaska:  Total  length,  189  (178-198);  hind 
foot,  30  (29-31).  Shull:  Average  of  10  adult  males  from  same 
locality:  Occipi to-nasal  length,  43.6  (42.7-44.2);  zygomatic  breadth, 
21.8  (21.4-22.2);  breadth  of  braincase,  18.6  (18-19.3);  interorbital 
breadth.  5.6  (5.1-6.2);  width  of  palatal  bridge,  2.5  (2.3-2.7);  length 
of  nasals,  13.3  (13-13.6).  Average  of  5  adult  females  from  same 
locality:  Occipito-nasal  length,  42.7  (42-44.1);  zygomatic  breadth, 
21.4  (20.7-22);  breadth  of  braincase,  18.4  (18.4-18.5);  interorbital 
breadth,  5.6  (5.2-5.8);  width  of  palatal  bridge,  2.8  (2.5-3.2);  length 
of  nasals,  13.2  (12.6-13.83 


36  NORTH   AMERICAN   FAUNA.  [No.  47 

Remarks. — The  collared  pika  is  a  well-marked  species  having  an 
extensive  range  in  Alaska  and  Yukon.  Widely  separated  geographi- 
cally from  the  other  American  species  (see  remarks  on  pp.  1-3)  it  is 
apparently  more  closely  related  to  0.  princeps  fenisex  of  southern 
British  Columbia  than  to  typical  princeps  of  the  Rocky  Mountain 
region.  Both  0.  eollaris  and  0.  p.  fenisex  are  characterized  by  a 
relatively  broad  skull,  with  short  nasals  and  broad  interpterygoid 
fossa.  In  external  appearance,  eollaris  is  much  more  grayish  and 
less  buffy  than  either  princeps  or  fenisex. 

Specimens  examined. — Total  number,  118,  as  follows: 

Alaska:  Knik  [=mountains  near],  1;  Mount  McKinley,  5;  Seward  Creek  (moun- 
tains at  head),  89;  Tanana  River  (near  head),  3;  Toklat  River  (head),  6;  White 
Pass,  1. 

British  Columbia:  Bennett,  5;  Tagish  Lake,  2.  85 

Yukon:  Conrod,  l;86  Ogilvie  Mountains  (head  of  Coal  Creek),  4;  Teslin  Lake 
(vicinity),  l.86 

OCHOTONA  SCHISTICEPS  (Merriam) 

[Synonymy  under  subspecies.] 

General  characters. — Head  and  nape  frequently  grayish,  in  marked 
contrast  to  the  rest  of  the  upperparts;  general  tone  of  upperparts 
varying  from  cinnamon-buff  or  avellaneous  to  wood  brown  or  sayal 
6rown.  Skull  averaging  somewhat  smaller,  but  relatively  broader 
than  that  of  0.  princeps  fenisex;  interpterygoid  fossa  about  as  in 
fenisex;  palatal  bridge  averaging  narrower  than  in  any  other  Ameri- 
can species. 

Color. — Head  and  nape  varying  from  pallid  neutral  gray  or  gull 
gray  to  smoke-gray,  always  more  or  less  washed  with  chaetura  drab, 
fuscous,  or  fuscous  black;  head  sometimes  pinkish  cinnamon  or  pink- 
ish buff,  mixed  with  fuscous  or  blackish,  the  nape  only  being  washed 
with  grayish;  rest  of  upperparts  pinkish  buff,  light  pinkish  cinnamon, 
vinaceous-cinnamon,  or  cinnamon,  more  or  less  mixed  with  chae- 
tura drab,  fuscous,  or  fuscous-black;  sides  of  nose  washed  with 
smoke-gray,  pale  smoke-gray,  or  light  buff;  sides  pale  pinkish  buff, 
pinkish  cinnamon,  light  pinkish  cinnamon,  vinaceous-cinnamon, 
or  light  ochraceous-salmon ;  ears  chaetura  drab,  chaetura  black, 
fuscous-black,  or  deep  mouse-gray,  margined  with  grayish  white  or 
light  buff;  feet  grayish  white,  pinkish  buff,  light  pinkish  cinnamon, 
or  cinnamon  buff;  soles  chaetura  drab,  chaetura  black,  hair-brown, 
fuscous,  or  fuscous-black;  palms  grayish  white,  light  drab,  tilleul 
buff,  or  hair-biown;  underparts  pinkish  buff,  pinkish  cinnamon,  vina- 
ceous-buff,  light  ochraceous-salmon,  or  soiled  whitish  with  a  faint 
wash  of  buff  or  cinnamon. 

Geographic  distribution. — The  Sierra  Nevada  of  California  (chiefly 
above  7,000  feet  altitude)  from  the  headwaters  of  Tule  River  north 
to  Mount  Shasta  and  Goose  Nest  Mountain;  the  lava-bed  region  of 
northeastern  California,  southern  Oregon,  and  southern  Idaho,  from 
Madeline  Plains  to  the  Snake  River  Desert,  Idaho ;  the  Blue  Moun- 
tains of  northeastern  Oregon  (except  northern  portion) ;  the  Toyabe 
Mountains  of  central  Nevada,  and  the  Beaver  and  Parowan  Ranges 
of  central  Utah.      (Fig.  4.) 

"Collection  Provincial  Mus.,  Victoria,  British  Columbia. 
•*  Collection  Victoria  Mem.  Mus. 


1924.] 


OCHOTOXA   SCHISTICEPS. 


37 


Fig.  4. — Distribution  of  the  subspecies  of  Ochotona  schisticcps. 


1.  O.  s.jewetti. 

2.  O.  s.  taylori. 


3.  O.  *.  schisticcps. 

4.  O.  s.muiri. 


5.  O.  s.  shettoni. 

6.  O.  s.  albata. 


9.  0.  s.  goldmani. 


7.  O.  s.  cinnamomea. 

8.  O.  s.fuscipes. 


Remarks. — In  the  Sierra  Nevada  system  and  the  mountains  of  the 
Great  Basin  the  species  scliisticeps  is  widely  distributed.  It  is  sep- 
arable into  at  least  9  races,  many  of  which  occupy  isolated  mountain 
areas  or  lava  outcrops  on  the  desert. 

Living  under  such  diverse  conditions,  the  various  forms  have  devel- 
oped striking  differences  in  coloration;  the  darkest  forms  (goldmani 
and  taylori)  are  found  in  the  lava-bed  formations  of  northeastern 
California,  eastern  Oregon,  and  southern  Idaho;  passing  southward 
in  the  Sierra  Nevada  the  pikas  gradually  become  paler,  the  palest 
form  of  all  being  found  in  the  Mount  Whitney  region,  at  the  extreme 
southern  end  of  the  range.  The  form  (cinnamomea)  found  in  the 
Toyabe  Mountains  (and  probably  other  ranges)  of  central  Nevada 
and  in  the  Beaver  Mountains,  Utah,  is  not  unlike  scMsticeps  in  col- 
oration, though  smaller,  but  strangely  enough,  the  form  occurring  in 
the  Parowan  Range — practically  adjacent  to  the  Beaver  Range — is 
larger  and  darker.  The  Blue  Mountains  form  (jewetti)  again  is  paler 
than  typical  schisticcps  in  summer,  but  darker  in  winter. 


38  NORTH   AMERICAN    FAUNA.  [No.  47 

OCHOTONA  SCHISTICEPS  SCHISTICEPS  (Merriam). 

Gray-headed  Pika. 

(PI.  IV,  fig.  2;  PI.  VI,  fig.  2.) 

Lagomys  schisticeps  Merriam,  North  Amer.  Fauna  No.  2.  p.  11,  October  30,  1889. 
Ochotona  schisticeps  Merriam,  North  Amer.  Fauna  No.  16,  p.  99,  1899. 

Type. — Collected  at  Dormer,  California,  June  9,  1888,  by  Charles 
A.  Allen;  j  adult,  skin  and  skull;  No.  186518,  U.  S.  National  Museum 
(formerly  No.  -fl-ft*  Merriam  collection). 

Geographic  distribution. — Boreal  zones  of  the  northern  Sierra 
Nevada,  from  Mount  Shasta  south  to  Donner  Pass;  altitudinal  range 
from  about  5,000  feet  (on  Mill  Creek,  above  Morgan,  Tehama  County) 
to  at  least  9,000  feet.    >  (Fig.  4,  p.  37.) 

Characters. — Coloration  paler  than  in  O.  princeps  brunnescens,  with 
head  and  nuchal  region  more  or  less  strongly  washed  with  grayish; 
skull  smaller,  with  much  narrower  palatal  bridge. 

Color. —  Winter  pelage:  Head  and  shoulders  smoke-gray,  mixed 
with  fuscous;  rest  of  upperparts  vinaceous-cinnamon,  mixed  with 
fuscous,  shading  on  sides  to  clear  vinaceous-cinnamon;  ears  chaetura 
black,  margined  with  light  buff;  feet  soiled  whitish,  washed  with 
pinkish  buff;  underparts  soiled  whitish,  washed  with  pinkish  buff  or 
light  pinkish  cinnamon.  Summer  pelage,:  Very  similar  to  the  winter 
pelage,  but  upperparts  less  uniformly  darkened  with  fuscous  and 
general  tone  nearer  cinnamon  than  vinaceous-cinnamon. 

Molt. — An  adult  female  from  Donner,  California,  August  18,  shows 
fresh  summer  pelage  covering  the  head,  shoulders,  and  middle  of  the 
back  nearly  to  the  rump,  the  sides  and  back  of  the  shoulders  retain- 
ing the  old  pelage;  a  male  from  Mount  Shasta,  September  25,  is  acquir- 
ing winter  pelage,  the  new  hair  covering  the  dorsil  area  to  within  about 
an  inch  of  the  rump,  the  lower  sides  and  underparts  still  retaining  the 
worn  summer  pelage.  An  adult  male  from  Donner,  October  9,  has 
completed  the  fall  molt  over  the  entire  dorsal  area,  but  the  belly  is  still 
in  old  pelage ;  another  from  the  same  locality,  October  8,  is  in  complete 
winter  pelage. 

Skull. — -Similar  to  that  of  O.  princeps  brunnescens  but  averaging 
slightly  smaller;  palatal  bridge  averaging  decidedly  narrower. 

Measurements. — -Average  of  15  adults  (10  males,  5  females)  from 
type  locality;  Total  length,  186  (178-200);  hind  foot,  29.5  (27-31). 
Skull:  Average  of  8  adult  males  from  tvpe  locality:92  occipito-nasal 
length,  42.5  (41.5-45);  zygomatic  breadth,  21.1  (20.4-21.7);  breadth 
of  braincase,  18.8  (18.1-19.4) ;  interorbital breadth,  5.1  (4.8-5.3) ;  width 
of  palatal  bridge,  1.6  (1.2-2.6);  length  of  nasals,  13.7  (13.2-14.5). 
Average  of  2  adult  females  from  tvpe  locality:  Occipito-nasal 
length,  41.8  (41.4-42.2);  zygomatic  breadth,  21  (20.8-21.1);  breadth 
of  braincase,  18.5  (18.1-19);  interorbital  breadth,  5.5  (5.3-5.7); 
width  of  palatal  bridge,  1.35  (1.3-1.4);  length  of  nasals,  13.55 
(13.5-13.6). 

Remarks.— The,  gray-headeJ  pika — the  first  of  the  California  forms 
to  be  characterized — is  the  darkest  of  the  Sierra  races,  though  not 
quite  so  blackish  as  taylori  from  the  lava-bed  region  of  Oregon  and 
northern  California.  It  intergrades  with  taylori  in  the  Mount  Shasta 
region,  as  shown  by  specimens  from  Mount  Shasta  in  fresh  fall  pel- 

"  It  seams  remarkable  that  in  this  series  of  topotrpe  skulls  all  are  considerably  smaller  than  the 
type  and  in  all  the  palatal  bridge  is  much  nirrower;  the  measurements  of  the  type  are  the  maxima 
given  above. 


1924.]  OCHOTOXA   SCHISTICEPS.  39 

age  which  are  slightly  darker  than  typical  schisticeps.  Southward, 
schisticeps  passes  into  nmiri  in  the  region  between  Donner  Pass  and 
Mount  Tallac. 

Specimens  examined. — Total  number,  54,  as  follows: 

California:  Donner,  25; 93  Donner  Pass,  2;94  Lassen  County,  2;  Lassen  Peak,  4; 
Mount  Shasta,  IS;95  Prattville  (12  miles  northeast),  1;  Summit  [=Donner],  4; 
Tahoe,  Placer  County,  1. 

OCHOTONA  SCHISTICEPS  TAYLORI  Grinnell. 

Taylor  Pika. 

(PI.  IV,  fig.  5;  PL  VI,  fig.  5.) 

Ochotona  taylori  Grinnell,  Proc  Biol.  Soc  Washington,  vol.  25,  p.  129,  July  31,  1912. 

Type. — Collected  on  Warren  Peak  (at  9,000  feet  altitude) ,  Warner 
Mountains,  Modoc  County,  California,  July  18,  1910,  by  W.  P.  Taylor 
and  H.  C.  Bryant:  <?  adult,  skin  and  skull;  No.  11292,  Museum  of 
Vertebrate  Zoology,  University  of  California;  original  number,  3885. 

Geographic  distribution. — Northeastern  California  and  southern 
Oregon,  from  Madeline  Plains  north  to  Steens  Mountains,  Oregon, 
and  west  to  Goose  Nest  Mountain,  Siskiyou  County,  California;  alti- 
tudinal  range  from  about  5,000  to  8,000  feet.     (Fig.  4,  p.  37.) 

Characters. — Similar  to  schisticeps  but  smaller;  coloration  in  summer 
pelage  slightly  more  blackish,  especially  on  head;  in  winter  pelage, 
decidedly  darker  above,  the  fuscous  shades  of  schisticeps  largely 
replaced  by  black  in  taylori  and  the  gray  on  head  also  darker. 

Color. — Summer  pelage  (July) :  Upperparts  light  pinkish  cinnamon, 
strongly  shaded  with  fuscous-black;  head  usually  ( ?)  like  back  without 
trace  of  gray  (all  of  the  small  series  from  the  type  locality  are  practically 
without  gray  on  head)  but  in  some  specimens  extensively  sprinkled 
with  whitish  hairs,  producing  a  general  tone  near  neutral  gray;  ears 
chaetura  black,  margined  with  buffy  white ;  soles  fuscous-black ;  palms 
tilleul  buff,  washed  with  drab ;  lower  sides  and  underparts  light  ochrace- 
ous-salmon,  becoming  more  cinnamon  on  the  throat.  Winter  pelage 
(October) :  Face  and  head  pale  gull  gray,  mixed  with  blackish,  produc- 
ing an  "iron-gray"  tone;  nose  sometimes  washed  with  smoke-gray 
and  fuscous;  dorsal  area  pale  pinkish  cinnamon,  heavily  washed  with 
dark  fuscous-black;  sides  between  light  vinaceous-cinnamon  and  light 
pinkish  cinnamon,  with  only  a  slight  mixture  of  fuscous;  ears  blackish, 
margined  with  buffy  white ;  feet  soiled  whitish,  faintly  washed  with 
pinkish  buff;  soles  fuscous-black;  palms  light  drab ;  underparts  soiled 
whitish,  moderately  washed  with  vinaceous-buff. 

Molt. — The  date  of  the  midsummer  molt  in  this  race  is  not  shown 
by  the  series  at  hand;  specimens  from  Goose  Nest  Mountain,  Cali- 
fornia, July  15  and  16,  are  apparently  in  worn  winter  pelage,  while 
one  from  Steens  Mountains,  Oregon,  July  26,  is  in  full  summer  pelage. 
The  fall  molt  apparently  begins  early,  as  shown  by  a  series  from  Steens 
Mountains,  August  26-30,  in  which  the  fresh  winter  pelage  covers 
nearly  all  the  dorsal  surface  of  more  than  half  the  specimens,  the  rest 
being  in  worn  summer  coat. 

"Two  in  collection  Amer.  Mus.  Nat.  Hist. 
"Collection  Mus.  Vert.  Zool. 
"One  in  collection  Mus.  Vert.  Zool. 


40  NORTH   AMERICAN    FAUNA.  [No.  47 

Skull. — Similar  to  scMsticeps  but  averaging  smaller. 

Measurements. — Average  of  9  adults  (6  males,  3  females)  from  vicin- 
ity of  type  localitv  and  southern  Oregon  (Adel  and  Mount  Warner) : 
Total  length,  175  (162-190);  hind  foot,  26.7  (25T28).  Skull:  Two 
adult  males  (type  and  one  from  Madeline  Plains,  California) :  Occipito- 
nasal  length,  41.7,  41.8;  zygomatic  breadth,  20.8,  20.5;  breadth  of 
braincase,  18.7,  18.1;  interorbital  breadth,  5.8,  4.7;  width  of  palatal 
bridge,  1.2,  1.4;  length  of  nasals,  13.6.  Average  of  5  adult  females 
(2  from  type  region  and  3  from  Steens  Mountains,  Oregon.) :  Occipito- 
nasal  length,  41.5  (39.7-42.7);  zygomatic  breadth,  20.3  (19.6-20.8); 
breadth  of  braincase,  17.9  (17.4-18.6);  interorbital  breadth,  4.6  (4.4- 
4.7):  width  of  palatal  bridge,  1.4  (1.2-1.6);  length  of  nasals,  13.5 
(12.5-14.4). 

Remarks. — This  race  is  closely  related  to  scMsticeps,  with  which  it 
intergrades  in  northeastern  California.  Its  range  is  chiefly  in  the  lava- 
bed  region  of  southern  Oregon  and  northeastern  California,  where  it 
descends  to  quite  low  altitudes.  In  summer  pelage  it  differs  but 
little  from  scMsticeps  but  in  winter  pelage  it  is  noticeably  darker. 
Specimens  available  from  the  type  locality  are  all  in  summer  pelage, 
and  judging  from  these  alone,  one  would  hardly  be  justified  in  separa- 
ting the  form  from  scMsticeps;  apparently,  however,  the  series  from 
southern  Oregon  shows  more  distinctive  characters,  especially  in  winter 
pelage. 

Specimens  examined. — Total  number,  35,  as  follows: 

California:  Goose  Nest  Mountain,  3;  Little  Shasta  River  (near  head),  Siskiyou 
County,  l;96  Madeline  Plains  (near  Termo,  5,400  feet  altitude"),  1;  Sugar  Hill, 
Modoc  County,  2; 96  Sugarloaf  Mountain,  Modoc  County,  2;  Warren  Peak,  Warner 
Mountains,  7.96 

Oregon:  Adel,  1;  Jack  Lake  (20  miles  northeast  of  Adel),  2;97  Mount  Warner,  2; 
Steens  Mountains,  14. 

OCHOTONA  SCHISTICEPS  GOLDMANI,  subsp.  nov.  98 

Lava-bed  Pika. 

(PL  IV,  fig.  8;  PI.  VI,  fig.  8.) 

Type. — Collected  at  Echo  Crater,  Snake  River  Desert,  Idaho  (20 
miles  southwest  of  Arco),  June  18,  1921,  by  Luther  J.  Goldman:  o* 
adult,  skin  and  skull;  No.  236408,  TT.  S.  National  Museum  (Biologi- 
cal Survey  collection) :  original  number,  2754. 

Geographic  distribution. — Known  only  from  Echo  Crater  and  Fis- 
sure Crater,  Snake  River  Desert,  Idaho.   (Fig.  4,  p.  37.) 

Characters. — Similar  to  taylori,  but  upperparts  much  darker  (more 
blackish) :  sides  and  underparts  paler  and  more  pinkish  (less  cinna- 
mon) ;  sides  of  nose  with  patches  of  gray ;  skull  broader  interorbitally 
with  shorter  jiasals. 

Color. —  Type  (fresh  summer  pelage):  Upperparts  (including  head) 
mixed  blackish  and  light  ochraceous-buff,  the  black  prevailing;  nape 
with  a  rather  indistinct  patch  of  pale  smoke-gray  (apparently  a 
remnant  of  winter  pelage) ;  sides  of  nose  with  large  patches  of  neutral 

86  Collection  Mus.  Vert.  Zool. 

•'  Collection  S.  G.  Jewett. 

•8  Named  for  Luther  Jacob  Goldman,  who  collected  all  the  specimens  at  present  known. 


1924.]  OCHOTONA   SCHISTIGEPS.  41 

gray;  ears  chaetura  black,  margined  with  creamy  white;  lower  sides 
pinkish  buff,  becoming  light  pinkish  cinnamon  on  sides  of  neck;  feet 
and  underparts  whitish,  moderately  washed  with  pinkish  buff;  soles 
deep  mouse-gray;  palms  soiled  whitish,  washed  with  mouse-gray. 
Worn  winter  pelage  (June  13):  Upperparts  much  less  blackish  and 
sides  paler  than  in  summer  pelage;  general  tone  of  upperparts  near 
hair-brown  (produced  by  a  mixture  of  light  ochraceous-buff  and 
pale  smoke-gray,  moderately  shaded  with  fuscous) ;  sides  pale  pink- 
ish buff,  shading  to  light  pinkish  cinnamon  on  sides  of  neck. 

Molt. — The  midsummer  molt  begins  in  June;  one  specimen  (  9  ), 
taken  June  13,  is  still  in  winter  pelage;  the  other  5  are  acquiring  sum- 
mer pelage,  which  first  covers  the  head  and  progresses  backward  over 
the  body;  in  one  specimen,  taken  June  13,  the  molt  is  practically  com- 
plete; in  others,  taken  June  13  and  June  18,  the  old  pelage  still  shows 
on  the  rump  and  in  an  irregular  patch  of  smoke-gray  on  the  nape. 

Skull. — Similar  to  taylori  but  averaging  longer,  and  broader  inter- 
orbitally;  palatal  bridge  slightly  wider;  nasals  shorter  and  broader, 
expanding  anteriorly. 

Measurements. —  Type:  Total  length,  188;  hind  foot,  31.  Average 
of  6  adults  from  type  locality h  183;  30.2.  SJcull:  Average  of  4  adults : 
Oceipito-nasal  length,  42.6  (41.5-43.9) ;  zygomatic  breadth,  20.6  (20.1- 
21.6):  breadth  of  braincase  18.5(17.5-19);  interorbital  breadth,  5.8 
(5.7-6.1) ;  width  of  palatal  bridge,  1.6  (1.3-1.9) :  length  of  nasals,  13.1 
(12.8-13.4). 

Remarks. — The  lava-bed  pika  is  the  darkest-known  form  in  the 
genus.  In  general  coloration  it  most  resembles  0.  princeps  princeps, 
but  it  is  darker  and  has  prominent  gray  patches  on  the  sides  of  the 
nose,  and  differs  widely  from  that  form  in  cranial  characters. 

It  is  strikingly  different  from  its  nearest  geographical  neighbor — 
0.  princeps  lemhi — which  lives  in  the  Lemhi  Mountains,  about  60  miles 
northeast  of  the  type  locality  of  the  present  form.  Compared  with 
lemhi,  goldmani  is  very  much  darker  in  coloration  and  has  a  broader 
skull  with  shorter,  broader  nasals,  larger  audital  bullae,  and  narrower 
palatal  bridge. 

Its  close  relationship  to  the  schisticeps  group  makes  advisable  its 
designation  as  a  race  of  that  species;  although  possessing  the  diag- 
nostic skull  characters  of  the  group  this  race  apparently  never  has  a 
gray  head  (at  least  in  summer  pelage) . 

The  occurrence  of  a  species  of  Ocnotona  in  the  Snake  River  Desert 
of  Idaho  was  unsuspected;  its  discovery  by  Luther  Goldman  extends 
the  range  of  the  schisticeps  group  eastward  from  the  Steens  Mountains, 
Oregon,  into  central  Idaho.  This  race  doubtless  is  completely  isolated 
from  its  relatives,  occurring  only  in  scattered  lava  piles  on  the  desert. 
Mr.  Goldman  states  that  at  Echo  Crater  the  animals  were  found  in 
small  numbers,  and  that  at  Fissure  Crater,  about  15  miles  south  in 
the  interior  of  the  desert,  abundant  old  sign  was  found  but  at  the  time 
of  his  visit  the  pikas  had  mostly  disappeared,  and  he  secured  only  one 
specimen  as  a  result  of  considerable  effort. 

Specimens  examined. — Total  number,  6,  as  follows: 

Idaho:  Echo  Grater  (20  miles  southwest  of  Arco),  5;  Fissure  Crater  (26  miles  south- 
west of  Arco),  1. 


42  NORTH    AMERICAN    FAUNA.  [No.  47 

OCHOTONA  SCHISTICEPS  JEWETTI  Howell. 
Blue  Mountains  Pika. 

Ocholona  schisticeps  jeuetti  Howell,  Proe.  Biol.  Soc.  Washington,  vol.  32,  p.  109,  May 
20,  1919. 

Type. — Collected  at  head  of  Pine  Creek,  near  Cornucopia,  south 
slope  Wallowa  Mountains,  Baker  County,  Oregon,  September  3,  1915, 
by  Stanley  G.  Jewett;«?  adult,  skin  and  skull;  No.  208352,  U.  S. 
National  Museum,  Biological  Survey  collection :  original  number,  2362. 

Geographic  distribution. — Wallowa  Mountains,  Strawberry  Moun- 
tains, and  southern  portion  of  Blue  Mountains,  northeastern  Oregon; 
altitudinal  range  approximately  from  5,000  to  10,000  feet.     (Fig.  4, 

P-37-) 

Characters.- — Similar  to  schisticeps   but  paler  in   summer   pelage 

slightly  darker  in  winter  pelage.  Compared  with  taylori:  Size  larger; 
coloration  in  winter  pelage  deeper  cinnamon  but  less  blackish;  in  sum- 
mer pelage  decidedly  paler  both  above  and  below. 

Color. — Summer  pelage:  Upperpart slight  pinkish  cinnamon,  mixed 
with  fuscous ;  sides  of  nose  pale  smoke-gray :  sides  of  face,  occiput,  and 
nape  washed  with  the  same;  ears  chaetura  drab,  margined  with  gray- 
ish white  and  faintly  washed  inside  with  the  same;  sides  light  pinkish 
cinnamon:  feet  grayish  white,  washed  with  light  pinkish  cinnamon; 
soles  chaetura  drab;  palms  grayish  white,  washed  with  drab;  under- 
pays grayish  white,  washed  with  light  pinkish  cinnamon.  Winter 
pelage:  Closely  similar  to  schisticeps  in  the  corresponding  pelage,  but 
slightly  darker. 

Molt. — The  midsummer  molt  apparently  occurs  during  July ;  in  a 
specimen  from  Bourne,  Oregon.  August  4,  the  summer  pelage  is  prac- 
tically complete,  except  for  a  patch  on  the  rump.  A  specimen  from 
Anthony,  Oregon,  September  2,  shows  the  beginnings  of  the  fall  molt, 
the  new  winter  pelage  covering  the  head,  shoulders,  and  fore  back,  the 
rest  of  the  upperparts  being  in  badly  worn  summer  pelage. 

Skull. — Closely  similar  to  schisticeps;  larger  than  that  of  taylori. 

Measurements. — Average  of  12  adults  (6  males,  6  females)  from 
Cornucopia,  Bourne,  and  Anthony,  Oregon:  Total  length,  182  (165- 
195);  hind  foot,  31  (30-32).  Skull;  Average  of  16  adults  (7  males, 
9  females)  from  the  type  region  and  Strawberry  Mountains:  Occipito- 
nasal  length,  43.3  (41.8-44.8);  zj'gomatic  breadth,  21.2  (20.6-22.1); 
breadth  of  braincase,  18.1  (16.9-19);  interorbital  breadth,  5.3  (4.9- 
5.8);  width  of  palatal  bridge,  1.8  (1.3-2.4);  length  of  nasals,  14.1  (13.1- 
15.3). 

Remarks. — This  race  most  nearly  resembles  schisticeps  of  the  high 
Sierra;  the  ranges  of  the  two  are  separated,  however,  by  that  of  tay- 
lori, which  occupies  the  intervening  lava-bed  region. . 

The  Wallowa  Mountains  mark  the  eastern  limit  of  O.  schisticeps  in 
this  region;  in  the  Seven  Devils  Mountains — directly  across  the  can- 
yon of  Snake  River  in  Idaho — occurs  a  race  of  0.  princeps,  a  quite 
distinct  species. 

Specimens  examined. — Total  number,  49.  as  follows: 

Oregon:  Anthony.  Baker  County,  15;"  Austin,  1;  Bourne,  9;  Cornucopia  (near  head 
East  Pine  Creek),  6;  Strawberry  Butte,  2;  Strawberry  Mountains,  8;  Wallowa 
Lake,  2;  Wallowa  Mountains,  6. 

"Twelve  in  collection  Amer.  Mus.  Nat.  Hist.;  2  in  collection  Mus.  Vert.  Zool.;  1  in  collection  E.  R. 
Warren. 


1924.]  OCHOTONA    SCHISTICEPS.  43 

OCHOTONA  SCHISTICEPS  MUIRI  Grinnell  and  Stoker. 

YOSEMITE    PlKA. 

(PI.  IV,  fig.  3;  PI.  VI,  fig.  3.) 

Ochotona  schisticeps  muiri  Grinnell  and  Storer,  Univ.  California  Publ.  Zool.,  vol.  17, 
p.  6,  August  23,  1916. 

Type. — Collected  at  Ten  Lakes,  Yosemite  National  Park,  Tuolumne 
County,  California  (at  9,300  feet  altitude),  October  11,  1915,  by  W.P. 
Taylor;  <?  adult,  skin  and  skull;  No.  23480,  Museum  of  Vertebrate 
Zoology,  University  of  California;  original  number,  7720. 

Geographic  distribution. — Central  portion  of  Sierra  Nevada,  from 
Mount  Tallac  south  to  Mammoth  Pass;  altitudinal  range,  7,800  to 
12,800  feet.  (Fig.  4,  p.  37.) 

Characters. — Similar  to  schisticeps  but  paler,  with  smaller  admixture 
of  fuscous  on  upperparts  and  sides;  gray  of  head  and  nape  paler; 
decidedly  paler  than  jewetti  both  in  winter  and  summer. 

Color. —  Winter  pelage:  Vinaceous  phase:  Upperparts  vinaceous- 
cinnamon,  moderately  mixed  on  dorsal  area  with  fuscous;  sides  of 
nose  washed  with  pale  smoke-gray  and  light  buff;  occiput  and  nape 
with  a  rather  indistinct  and  irregular  (often  nearly  obsolete)  wash  of 

f>ale  smoke-gray;  ears  chaetura  black,  margined  with  buffy  white; 
eet  soiled  whitish,  washed  with  pinkish  buff;  soles  hair-brown;  palms 
grayish  white,  washed  with  mouse-gray ;  underparts  heavily  washed 
with  pinkish  cinnamon.  Drab  phase  (type) :  Upperparts  mixed  light 
pinkish  cinnamon  and  fuscous,  producing  a  yellowish  drab  tone;  head, 
face,  and  nape  mixed  pale  smoke-gray  and  fuscous,  washed  with  pink- 
ish buff;  sides  light  pinkish  cinnamon;  underparts  grayish  white, 
moderately  washed  with  pale  pinkish  buff;  ears  and  feet  as  in  vina- 
ceous phase.  Summer  pelage:  Upperparts  cinnamon-buff  or  light 
pinkish  cinnamon,  moderately  mixed  with  fuscous;  sides  and  under- 
parts between  pinkish  buff  and  cinnamon-buff,  or  sometimes  pinkish 
cinnamon;  head,  ears,  and  feet  as  in  winter  pelage. 

Molt. — An  adult  female  from  Mount  Dana,  California,  August  20,  is 
in  badly  worn  winter  pelage,  the  fresh  midsummer  pelage  showing  on 
the  nose  and  face  and  in  small  patches  on  the  middle  of  the  back;  in 
an  adult  male  from  Mount  Lyell,  August  30,  the  new  pelage  covers  the 
head,  shoulders,  and  most  of  the  back  to  the  rump  excepting  a  small 
area  on  the  hinder  back  and  the  greater  part  of  the  sides ;  the  under- 
parts apparently  are  in  fresh  pelage;  an  adult  male  from  Mammoth 
Pass,  July  25,  is  in  nearly  complete  summer  pelage  except  on  the 
rump  and  buttocks. 

This  pelage  must  be  retained  but  a  short  time,  for  specimens  taken 
at  Ten  Lakes,  Yosemite  Park,  October  10  and  11,  have  acquired  a 
fresh  winter  pelage  (differing  in  color  from  the  summer  pelage)  over 
the  entire  dorsal  area,  the  molting  line  showing  clearly  on  the  lower 
sides. 

Slcull. — Similar  to  schisticeps,  but  averaging  slightly  smaller  and 
relatively  narrower. 

Measurements. — Average  of  22  adults  (11  males,  11  females)  from 
Yosemite  National  Park:  Total  length,  188  (180-205);  hind  foot, 
29.7  (27-32).  Skull.  Average  of  20  adults  (10  males,  10  females) 
from  Yosemite  National  Park:  Occipito-nasal  length,  42  (40.2-42.7); 


44  NORTH    AMERICAN    FAUNA.  [No.  47 

zygomatic  breadth,  20.7  (20.1-22.1);  breadth  of  braincase,  17.9 
(17.2-19.1);  interorbital  breadth,  4.9  (4.4-5.7);  width  of  palatal 
bridge,  1.6  (1.2-2);  length  of  nasals,  13.9  (13.1-14.6). 

Remark. — The  Yosemite  pika  is  a  well-marked  form,  distinctly 
paler  than  schisticeps,  but  not  nearly  so  pale  as  albata  of  the  Mount 
Whitney  region.  It  has  a  more  extensive  range  than  any  of  the 
other  Sierra  forms  and  is  apparently  abundant  throughout  its  range. 

Specimens  examined. — Total  number,  100,  as  follows: 

California:  Bishop  Creek  [=  mountains  west],  3;  Bloody  Canyon,  Mono  Lake,  1; ' 
Donohoe  Pass,  Mono  County,  lj'^JEcho,  El  Dorado  County,  4;2  Ellery  Lake, 
Mono  County,  1; '  Heather  Lake,  10;  l> 2  Lake  Tenaya,  6; 3  Lucile  Lake,  El  Dorado 
County,  l;1  Lyell  Canyon,  Yosemite  National  Park.  13; '  Markleeville,  3;  Mono 
Pass,  1;  Mount  Dana,  3;4  Mount  Gibbs,  1;  Mount  Hoffman,  2; l  Mount  Lyell.  10; 
Mount  Parsons,  l;1  Mount  Tallac,  9; 5  Mount  Unicorn,  2;  Owens  River  (head),  1; 
Pine  City,  Mono  County  [near  Mammoth  Pass],  2;  Soda  Springs,  Yosemite  Na- 
tional Park,  3; '  Summit  of  Sierra  (Latitude  39°),  2;1  Ten  Lakes,  Yosemite 
National  Park,  6;1  Tioga  Lake,  Mono  County,  l;1  Vogelsang  Lake,  Yosemite 
National  Park,  12;  *  Washburn  Lake,  Yosemite  National  Park,  l.1 

OCHOTONA  SCHISTICEPS  ALBATA  Grinnell. 

Mount  Whitney  Pika. 

Ochotona  albatus  Grinnell,  Univ.  California  Publ.  Zool.,  vol.  10,  p.  125,  Januarv  31, 

1912. 
Ochotona  schisticeps  albatus,  Grinnell,  op.  cit.,  vol.  17,  p.  6,  1916. 

Type. — Collected  at  Cottonwood  Lakes,  near  Mount  Whitney  (at 
11,000  feet  altitude),  Inyo  County,  California,  September  3,  1911,  by 
Joseph  Grinnell;  9  adult,  skin  and  skull;  No.  16223,  Museum  of 
Vertebrate  Zoology,  University  of  California;  original  number,  1741. 

Geographic  distribution. — Southern  part  of  the  high  Sierra  Nevada, 
from  Kearsarge  Pass  south  to  the  headwaters  of  the  Tule  and  Kern 
Rivers:  altitudinal  range  from  about  8,500  feet  to  13,000  feet.     (Fig. 

4>  P-  37-) 

Characters. — Similar  to  O.  schisticeps  muiri,  but  coloration  decidedly 

paler. 

Color. — Summer  pelage:  Head  and  face  pale  smoke-gray,  moder- 
ately washed  with  chaetura  drab;  dorsal  area  pinkish  buff,  sparingly 
darkened  with  chaetura  drab,  shading  to  pale  pinkish  buff  on  lower 
sides;  ears  between  chaetura  drab  and  deep  mouse-gray ;  feet  grayish 
white,  faintly  washed  with  pinkish  buff;  soles  fuscous  or  dark  hair- 
brown  ;  palms  soiled  whitish,  washed  with  light  drab ;  underparts  gray- 
ish white,  faintly,  or  sometimes  strongly  washed  with  pinkish  buff. 
Worn  winter  pelage  (June) :  Very  similar  to  the  summer  pelage  but 
upperparts,  especially  the  head,  more  grayish  (less  buffy).  Fresh 
winter  pelage:  Not  represented. 

Molt. — Two  adult  males  from  Mount  Whitney,  California,  August 
9,  are  apparently  acquiring  summer  pelage,  the  new  hairs  coming  in 
irregularly  over  the  greater  part  of  the  dorsal  area;  three  adult  females 
from  Mineral  King,  California,  September  9,  likewise  show  a  new  pel- 
age (presumably  the  summer  pelage)  investing  the  head,  shoulders, 
and  middle  of  the  back.  The  beginnings  of  winter  pelage  are  not 
Bhown  by  the  specimens  at  hand. 

1  Collection  Mus.  Vert.  Zool. 

'Collection  Mus.  Comp.  Zool. 

5  Five  in  collection  Mus.  Vert.  Zool. 

*  Two  in  collection  Mus.  Vert.  Zool. 

5  Five  in  collection  Mus.  Comp.  Zool.;  3  in  collection  Acad.  Nat.  Scl.  Philadelphia. 


1924.]  OCHOTONA   SCHISTICEPS.  45 

Skull. — Practically  identical  with  that  of  muiri. 

Measurements. — Average  of  18  adults  (10  males,  8  females)  from 
type  region  (Mount  Whitney  and  Kearsarge  Pass) :  Total  length,  182 
(176-190);  hind  foot,  29.6  (28-31).  Skull:  Average  of  17  adults  (9 
males,  8  females)  from  type  region  (Kearsarge  Pass  to  Mineral  King) : 
Occipito-nasal  length,  42.4  (40.8-43.4) ;  zygomatic  breadth,  20.7  (20- 
21.2);  breadth  of  braincase,  17.9  (17-18.9);  interorbital  breadth,  4.8 
(4.2-5.5);  width  of  palatal  bridge,  1.6  (1.3-2.4);  length  of  nasals, 
13.  9  (13.3-14.4). 

Remarks. — Palest  of  the  American  pikas,  the  nearest  relative  of 
this  race  is  muiri,  of  the  Yosemite  region,  intergradation  with  which 
is  apparently  shown  by  a  series  from  Mineral  King,  some  of  which 
are  typical  of  albata,  while  several  are  scarcely  distinguishable  from 
muiri.  Its  range  is  not  extensive,  being  restricted  to  the  region 
around  Mount  Whitney,  northward  to  Kearsarge  Pass,  between  alti- 
tudes of  about  8,500  feet  (Onion  Valley)  to  13,000  feet  (on  west  slopes 
of  Mount  Whitney).  Grinnell  refers  to  the  aridity  of  the  climate  of 
this  region,  as  compared  with  that  of  the  northern  Sierra  Nevada, 
and  to  its  long  existence  as  a  land  mass  of  high  elevation  as  possible 
explanations  of  its  faunal  distinctness.6 

Specimens  examined. — Total  number,  68,  as  follows: 

California:  Bullfrog  Lake,  Fresno  County,  8;7  Cottonwood  Lakes,  4;7  Cottonwood 
Meadows,  2;  Independence  Creek  (10,000  feet  altitude),  1;  Kaweah  River  (East 
Fork,  near  Mineral  King),  2;  Kearsarge  Pass,  17  ;7  Kern  River  (near  head  of  North 
Fork),  2;  Mineral  King  [East  Fork  Kaweah  River],  20;  Mount  Whitney,  10;8 
Whitney  Creek,  Tulare  County  (altitude  10,850  feet),  2.7 

OCHOTONA  SCHISTICEPS  SHELTONI  Grinnell. 

White  Mountains  Pika. 

Ochotona  schisticeps  sheltoni  Grinnell,  Univ.  California  Publ.  Zool.,  vol.  17,  no.  14, 
p.  429,  April  25,  1918. 

Type. — Collected  in  the  White  Mountains  (at  11,000  feet  altitude), 
near  Big  Prospector  Meadow,  Mono  County,  California,  July  29,  1917, 
by  A.  C.  Shelton;  <?  adult,  skin  and  skull;  No.  27560,  Museum  of 
Vertebrate  Zoology,  University  of  California;  original  number,  3414. 

Geographic  distribution. — The  White  Mountains  of  California  and 
Nevada  (8,000  to  13,000  feet  altitude).     (Fig.  4,  p.37.) 

Characters . — Size  of  taylori:  somewhat  intermediate  in  coloration 
between  schisticeps  and  muiri.  Compared  with  schisticeps:  Dorsal  area 
averaging  slightly  less  cinnamon  in  general  tone  and  in  some  speci- 
mens more  heavily  washed  with  blackish;  nape  a  slightly  paler  shade 
of  gray.  Compared  with  muiri:  Dorsal  area  averaging  slightly  darker, 
some  specimens  decidedly  more  blackish;  sides  deeper  cinnamon. 
Compared  with  taylori:  Coloration  in  summer  pelage  similar,  but 
averaging  decidedly  paler,  especially  on  the  head. 

Color. — Summer  pelage:  Upperparts  (including  head)  light  pinkish 
cinnamon,  more  or  less  washed  with  fuscous-black,  the  general  tone 
varying  from  avellaneous  to  wood  brown ;  occiput  and  nape  washed 
to  a  varying  extent  by  grayish  white  (in  some  specimens  scarcely 
evident);  ears  dark  fuscous-black  (almost  black),  margined  with 
grayish  white:  sides  pinkish  cinnamon  or  light  pinkish  cinnamon ;  feet 

6  Grinnell,  Joseph,  T'niv.  California.  Publ.  Zool.,  vol.  10,  p.  127,  1912. 

'Collection  Mus.  Vert.  Zool. 

•Seven  in  collection  Field  Mus.  Nat.  Hist.;  2  in  collection  Mus.  Comp.  Zool. 

73502—24 4 


46  NORTH   AMERICAN   FAUNA.  [No.  47 

whitish,  more  or  less  washed  with  light  pinkish  cinnamon;  soles 
chaetura  black;  palms  soiled  whitish,  shaded  with  hair-brown;  under- 
parts  whitish,  moderately  washed  with  pinkish  cinnamon  or  light 
pinkish  cinnamon.      Winter  pelage:  Not  represented. 

Skull. — Closely  similar  to  taylori  but  palatal  bridge  broader;  simi- 
lar to  schisticeps  but  averaging  slightly  smaller;  nasals  narrower 
posteriorly;  closely  similar  to  muiri  but  averaging  slightly  shorter 
and  relatively  broader. 

Measurements. — Average  of  11  adults  (7  males,  4  females)  from 
White  Mountains,  California :  Total  length,  175  (167-188) ;  hind  foot, 
27.6  (26-30).  SJcull:  Average  qf  11  adult  males  from  type  region: 
Occipito-nasal  length,  41.8  (40.9-42.8) ;  zygomatic  breadth,  21  (20.3- 
21.8):  breadth  of  braincase,  18.3  (17.3-18.7);  interorbital  breadth,  4.8 
(4.5-5.2);  width  of  palatal  bridge,  1.7  (1.4-2.2);  length  of  nasals, 
13.9  (13.2-14.4);  average  of  10  adult  females  from  tvpe  region: 
Occipito-nasal  length,  40.9  (39.5-42.5) ;  zygomatic  breadth,  20.5  (20.2- 
21.2) ;  breadth  of  braincase,  17.9  (17.3-18.4);  interorbital  breadth,  4.7 
(4.4-5.2) ;  width  of  palatal  bridge,  1.8  (1.4-2.2) ;  length  of  nasals,  13.5 
(13-14.6.) 

Remarks. — The  pika  of  the  White  Mountains  is  somewhat  inter- 
mediate in  color  between  schisticeps  and  muiri,  to  both  of  which  it  is 
very  closely  related;  on  geographic  grounds  it  might  be  expected  to 
be  nearer  albata  of  the  Mount  Whitney  region. 

Specimens  examined. — Total  number,  45,  from  the  type  region.9 

OCHOTONA  SCHISTICEPS  CINNAMOMEA  J.  A.  Allen. 

Beaver  Mountains  Pika. 

(PI.  IV,  fig.  9;  PI.  VI,  fig.  9.) 

Ochotona  cinnamomea  J.  A.  Allen,  Mus.  Brooklyn  Inet.  Arts  and  Sci.,  Science  Bull. 
vol.  1,  no.  5,  p.  121,  March  31,  1905. 

Type. — Collected  at  "Briggs"  [  =  Britts]  Meadows,  Beaver  Moun- 
tains, Utah  (at  11,000  feet  altitude) ,  August  20,  1904,  by  George 
P.  Engelhardt;  adult  (unsexed),  skin  and  skull;  No.  28733,  American 
Museum  of  Natural  History  (formerly  No.  434,  Museum  of  the 
Brooklyn  Institute  of  Arts  and  Sciences). 

Geographic  distribution. — Beaver  Range,  Utah,  and  Toyabe  Moun- 
tains, Nevada;  altitudinal  range  from  8,000  to  11,000  feet.  (Fig.  4, 
p.  37.) 

Characters. — Very  similar  to  schisticeps  but  smaller;  face,  head, 
and  nape  (in  summer  pelage)  averaging  paler;  sides  slightly  richer 
cinnamon. 

Color. — Summer  pelage:  Upperparts  cinnamon  or  vinaceous-cinna- 
mon,  mixed  with  fuscous-black,  shading  on  sides  to  deep  vinaceous- 
cinnamon;  nose  and  face  washed  with  smoke-gray,  pale  smoke- 
gray,  or  light  buff;  top  of  head  fuscous  or  fuscous-black,  washed 
with  light  buff;  nape  with  an  irregular  and  often  indistinct  patch 
of  pallid  neutral  gray;  ears  chaetura  black  or  chaetura  drab,  margined 
with  buffy  white,  the  long  hairs  at  anterior  base  grayish  white;  feet 
grayish,  washed  with  pinkish  buff  or  cinnamon-bun;  soles  hair-brown 
or  fuscous;  palms  buffy  white,  washed  with  drab;  underparts  light 
pinkish  cinnamon,  the  bases  of  the  hairs  deep  neutral  gray.  Winter 
pelage:  Not  represented. 

'Thirty-eight  in  collection  Mus.  Vert.  Zool. 


1924.]  OCHOTONA   SCHISTICEPS.  47 

Molt. — The  date  of  the  midsummer  molt  in  this  race  is  not  indicated 
by  the  material  at  hand;  the  series  from  the  type  locality,  taken  be- 
tween August  14  and  21,  are  all  in  complete  summer  pelage,  more  or  less 
worn.  The  fall  molt  begins  early  in  September,  as  shown  by  a  speci- 
men from  Toyabe  Mountains,  Nevada,  September  5,  in  which  the 
fresh  winter  pelage  covers  the  fore  back  and  part  of  the  head,  the 
occiput  and  nape  being  in  old  worn  pelage.  Another  specimen  from 
the  same  locality,  September  15,  has  practically  the  whole  dorsal 
area  renewed,  but  likewise  shows  a  worn  area  on  the  occiput. 

Skull. — Similar  to  that  of  taylori  but  braincase  averaging  narrower; 
smaller  than  that  of  schisticeps. 

Measurements. — Average  of  11  adults  (5  males,  6  females)  from 
type  locality:  Total  length,  188  (180-200);  hind  foot,  28.4  (27-29). 
Skull:  Average  of  5  adult  males  from  type  locality:  Occipito-nasal 
length,  41.6  (40.7-42.2):  zygomatic  breadth,  20.7  (20.3-21);  breadth 
of  braincase,  17.8  (17.4-18.5);  interorbital  breadth,  5.2  (4.8-5.9); 
width  of  palatal  bridge,  1.6  (1.4-1.7);  length  of  nasals,  13.5  (13-14.1). 
Average  of  two  adult  females  from  type  locality:  Occipito-nasal  length, 
39.9  (39.5-40.3);  zygomatic  breadth,  20.1;  breadth  of  braincase,  17 
(16.5-17.5);  interorbital  breadth,  5;  width  of  palatal  bridge,  1.5 
(1.4-1.7);  length  of  nasals,  12.6  (12.2-13). 

Remarks. — Considering  the  wide  area  of  desert  separating  the  range 
of  cinnamomea  from  that  of  schisticeps  the  differences  between  the  two 
forms  are  remarkably  slight.  A  series  from  the  Toyabe  Mountains, 
central  Nevada,  agrees  closely  with  typical  cinnamomea.  In  this 
series  are  two  specimens  acquiring  winter  pelage  (September  5  and 
15);  this  pelage  differs  from  the  summer  pelage  of  typical  specimens 
in  being  nearer  vinaceous-cinnamon  above ;  others  in  the  series  taken 
in  May  are  decidedly  paler  than  August  specimens  from  the  Beaver 
Mountains,  the  sides  and  upperparts  being  light  pinkish  cinnamon  in 
general  tone,  moderately  washed  on  the  dorsal  area  with  fuscous. 

Specimens  examined. — Total  number,  25,  as  follows: 

Nevada:  Arc  Dome  [Toyabe  Mountains]  5;  Toyabe  Mountains  (South  Twin  River), 4. 
Utah:  Beaver  Mountains  (Britt's  Meadows,  15;  Puffer  Lake,  1),  16. 10 

OCHOTONA  SCHISTICEPS  FUSCIPES  Howell. 
Parowan  Mountains  Pika. 
(PI.  IV,  fig.  6;  PI.  VI,  fig.  6.) 

Ochotona  schisticrps  fuscipes  Howell,  Proc.  Biol.  Soc.  Washington,  vol.  32,  p.  110, 
May  20,  1919. 

Type. — Collected  at  Brian  Head.  Parowan  Mountains,  Iron  County, 
Utah,  September  7,  1908,  by  W.  H.  Osgood;  $  adult,  skin  and  skull; 
No.  158094,  U.  S.  National  Museum.  Biological  Survey  collection; 
original  number,  3475. 

Geographic  distribution. — Known  onlv  from  tvpe  locality.  (Fig.  4, 
p.  37.) 

Characters. — Closely  similar  (in  summer  pelage)  to  schisticeps ,  but 
soles  and  palms  darker;  similar  to  taylori,  but  head  paler  and  more 
grayish,  general  tone  of  upperparts  less  vinaceous  (more  pinkish)  and 

10 One  in  collection  Amer.  Mus.  Nat.  Hist. 


48  NORTH    AMERICAN    FAUNA.  [No.  47 

underparts  paler;  similar  to  cinnamomea  but  larger,  and  coloration 
above  more  blackish  and  less  intensely  cinnamon. 

Color. — Summer  pelage:  Upperparts  mixed  light  pinkish  cinnamon 
and  fuscous-black  (the  blackish  prevailing) ;  tip  of  nose  fuscous-black; 
sides  of  nose  and  face  smoke-gray,  tinged  with  buff  and  washed  with 
fuscous;  back  of  head  and  nape  washed  with  pale  neutral  gray;  ears 
fuscous-black,  margined  with  buffy  white;  sides  pinkish  cinnamon 
mixed  with  fuscous,  becoming  clear  pinkish  cinnamon  along  line  of 
belly;  hind  feet  soiled  whitish,  washed  with  pale  cinnamon-buff,  the 
soles  fuscous ;  front  feet  cinnamon-buff,  the  palms  hair-brown ;  under- 
parts grayish  white,  heavily  washed  with  pinkish  cinnamon.  Winter 
pelage:  Not  represented. 

Skull. — Similar  to  that  of  cinnamomea,  but  larger  and  relatively 
broader,  with  larger  audital  bullae:  very  similar  to  that  of  schisticeps, 
but  braincase  averaging  slightly  narrower. 

Measurements. — Average  of  5  adults  (1  male,  4  females)  from  type 
locality:  Total  length,  200  (195-207);  hind  foot,  30.3  (29-32). 
Skull:  Average  of  5  adults  (1  male,  4  females) :  Occipito-nasal  length, 
42.4  (41.7-43.4);  zygomatic  breadth.  21.3  (20.4-22.2);  breadth  of 
braincase,  17.8  (17.2-18.5);  interorbital  breadth,  5.1  (4.9-5.3) ;  width 
of  palatal  bridge,  1.4  (1.2-1.5);  length  of  nasals,  13.1  (12.3-13.9). 

Remarks. — On  account  of  the  close  physical  connection  existing 
between  the  Parowan  Range  and  the  Beaver  Range,  the  pikas  on  the 
two  ranges  would  reasonably  be  expected  to  be  the  same.  The 
Parowan  Mountain  form,  however,  is  apparently  more  closely  related 
to  scJiisiiceps  of  the  Sierra  Nevada  and  to  taylori  of  the  lava-bed 
region  of  southern  Oregon  than  to  its  near  neighbor,  cinnamomea,  of 
the  Beaver  Mountains.  Its  range  is  separated,  however,  from  the 
ranges  of  the  California  and  Oregon  races  by  a  broad  expanse  of 
desert;  furthermore,  the  Toyabe  Mountains  in  central  Nevada  are 
occupied  by  cinnamomea. 

Specimens  examined. — Total  number,  6,  from  type  locality. 


1924.] 


CRANIAL    MEASUREMENTS. 


49 


Cranial  measurements  of  Ochotona. 


?pecies  and  locality. 


Ochotona  princeps  princeps: 
Henrv  House,  Alberta. 

Do 

Do 

Do 


Ochotona  princeps  lutescens: 

Mount  Inglesmaldie,  Alberta. 


Do. 
Do. 
Do. 


Ochotona  princeps  levin: 

Little  15elt  Mountains,  Montana. 
Chiel  Mountain  Lake,  Montana. 


Ochotona  princeps  lemhi: 

Lemhi  Mountains,  Idaho. 

Do 

Do 

Do 


Ochotona  princeps  ventorum: 

Wind    River   Mountains,   Wyo- 
ming. 

Do 

Do 

Do 


Ochotona  princeps  uinta: 

Uinta  Mountains,  ntah 

Salt  River  Mountains,  Wyoming. 

Uinta  Mountains,  Utah 

Salt  River  Mountains,  Wyoming. 


Ochotona  princeps  nevadtnsis: 
Rubv  Mountains,  Nevada . 
Do 


Ochotona  princeps  figginsi: 
Pagoda  Peak.  Colorado. . . 
Trapper's  Lake,  Colorado. 
Pagoda  Peak,  Colorado . . . 


Do. 


Ochotona  princeps  saxatilis: 

Alma,  Colorado 

Corona,  Colorado 

Long's  Peak,  Colorado 

Mount  Lincoln,  Colorado. 


Ochotona  princeps  incana: 

Pecos  Baldy,  New  Mexico. . . 
Do 

Wheeler  Peak,  New  Mexico. 
Pecos  Baldy,  New  Mexico... 


Ochotona  princeps  nigrescens: 

Jemez  Mountains,  New  Mexico . . 

Ochotona  princeps  cuppes: 

Gold  Range,  British  Columbia... 


Glacier,  British  Columbia <? 


45.2 
43.2 
45 
42 


43.3 

41.9 
43.3 
41.1 


cf  ,  41.6 

<?  i  44.1 

9  41.3 

9  i  42 


46.1 
43.8 

46.7 


45.3 

44 

-!i.2 

43.4 


42 
44.1 


<?  42.8 
3  44.7 
9 


43.7 


46 
48.1 
44.5 
47.2 


44.4 
45.8 
44.1 
46.8 


■<f, 


21.1 
21 

21.9 
21 


21.6 
21 
20.' 5 


21.9 
20.5 


21 
21.3 

20.4 
20.5 


21.5 


22.6 
21.4 


21.1 
21.2 


21.4 
22.3 
20.7 


21.8 
23.2 
21.3 

22.1 


21.8 
22.3 
21.3 
22.5 


21.9 
22.5 


18.3 
17.6 
18.6 
17.4 


17.3 
18.8 
16.9 


17.4 
17.2 


17.6 
18.5 
17 
16.7 


17.8 

IS.  5 
17.3 
18.5 


17.5 
18.:' 
18.4 
18.2 


17.9 
18 


18 

19.2 
17 


18.9 
19.6 
17.9 
18.5 


19.5 
19 
18.3 
19 


19 
18.3 


4.5 
5.6 
5.6 
5.8 


5.3 


5.8 
5.5 


6.1 
5.2 


4.7 
4.9 
4.8 
4.7 


5.1 
4.6 
5 


5.1 
5.6 
5.1 
5.6 


5.4 
5.2 


5.2 
5.5 
5.1 


5.6 
5.9 
5.3 
5.7 


5 

5.8 
5.2 
5.3 


5.3 

5 
5 


2.3 
2 

2.1 
1.8 


1.8 
L6 


1.9 
1.5 


1.7 
2.1 
2.1 
1.8 


2.1 
1.9 
1.9 


2.4 
2.3 
2.8 
1.8 


2.5 

2.8 


1.8 
2.7 
2 

1.7 


2.4 
2.8 
1.8 
2 


2.8 
2.2 
2.2 
2.5 


2.3 
3.5 


Remarks. 


14.5 
13.9 
14.7 
14.2 


13.9 

14.1 

12.9 
12.3 


14.2 
14.2 


13.8 
14.9 
14.3 
14 


14.4 
13.7 
14.9 


15.7 
15 
15 
14.7 


14.3 
14.5 


14.5 
15 

14.8 


15.1 
16.3 
15.3 
16.8 


15 
15 

14 
15.8 


14.3 

13.5 
15.4 


Old  adult. 
Adult. 
Old  adult. 
&S    Do- 


Old   adult; 

tvpe. 
Old  adult. 
Adult. 
Do. 


Do. 

Adult;  type. 


Adult. 
Do. 
Adult;  type. 
Adult. 


Do. 

Do. 

Do. 
Do. 


Do. 
Do. 
Do. 
Do. 


Do. 
Adult;  type. 


Adult. 
Do. 
Subadult; 

type. 
Adult. 


Subadult. 
Old  adult. 
Adult. 
Do. 


Do. 

Do. 

Do. 

Adult;  typ«. 


Do. 


Subadult; 

type. 
Old  adult. 


1  Collection  Amer.  Mus.  Nat.  Hist. 

2  Collection  E.  R.  Warren,  Colorado  Springs,  Colorado. 
*  Collection  Colorado  Mus.  Nat.  Hist. 

'Collection  Mus.  Comp.  Zool. 


50 


NORTH    AMERICAN    FAUNA. 
Cranial  measurements  of  Ochotona — Continued. 


[No.  47 


Species  and  locality. 


O  03 


Remarks. 


0 1443a 
6  848 


69274 
69275 
67130 


227258 
89840 
41538 


91144 
91140 
91143 


131266 
131329 
131298 


186517 


100466 
100250 


811292 
89120 
811294 


236408 
236406 


208352 

207676 
•040548 


8  23480 
109407 
109 10S 


827560 
41420 


158089 
158093 


158094 
158096 
158095 


Ochotona  princcps  fenisex: 

Head  of  Ashnola  River.  British 

Columbia. 
Horseshoe  Basin,  Washington. ... 


Ochotona  princcps  brooksi: 

Sicamous,  British  Columbia. 

Do 

Do 


Ochotona  princcps  brunncscens: 

Keechelus,  Washington 

Mount  Rainier,  Washington. 
Easton,  Washington 


Ochotona  princcps  fvmosa: 
Mount  Jefferson.  Oregon. 

Do 

Do 


Ochotona  collaris: 

Seward  Creek,  near  Eagle.  Alaska 

Do 

Do 


Ochotona  schisliceps  schisticeps 
Donner,  California 


Do 
Do. 


Ochotona  sell  isticeps  tnylori: 

Warner  Mountains,  California. 

Madeline  Plains.  California 

Warner  Mountains,  California. 


Ochotona  schisliceps  goldmani: 
Snake  River  Desert,  Idaho. 
Do 


(Jch  otona  schisliceps  jewetti: 
Cornucopia,  Oregon... 

Bourne,  Oregon 

Anthony,  Oregon 


Ochotona  schisliceps  muiri: 
Ten  Lakes,  California... 
Mount  Lyell,  California. 
Do 


Ochotona  schisliceps  albata: 
•12584  Mount  Whitnev,  California. 

42689  Mineral  King,  California 

•  12590  Mount  Whitney,  California. 


Ochotona  schisliceps  shclloni: 

White  Mountains.  California. 
Do 


Ochotona  schisliceps  cinnamomea: 

Beaver  Mountains,  Utah 

Do 


Ochotona  schisticeps  fuscipes: 
Parowan  Mountains,  Utah. 

Do 

Do 


<?/ 


42 
43.1 


42.3 
43.9 
43.5 


45.9 
42.3 
44.6 


44.5 
45.6 
43.9 


43.1 
44.2 
44.1 


41.5 
42.2 


41.7 
41.8 

42.7 


43.5 
43.9 


42.6 

44 

43.8 


41.6 
42.4 
42.5 


43.3 
41.7 
42.6 


42.8 
40.8 


41.7 

40.3 


21.5 

22 

20.6 


22.3 
21.5 
21.8 


21.3 

22 

21.4 


21.9 

22 

22 


20.9 
20.8 


20.8 
20.5 


20.4 
21.6 


21.6 
21.3 
21.8 


20.4 
21 

20.5 


20 

20.8 

21.2 


21.5 
20.3 


20.9 
20.1 


41.7  21.4 
43.4  !  21.4 
41.7  I  20.4 

I 


18.7 
19 


17.9 
18.4 
17.5 


19.2 
18.7 
18.3 


18.2 
18.1 
IS.  9 


19.1 
18.6 
18.4 


19.4 


18.2 
18.1 


18.7 
18.1 
17.6 


18.8 
18.7 


19 

18.5 

18.2 


17.3 
17.8 
18 


18.2 
17 
IS.  2 


18.7 
18 


17.8 
17.5 


17.9 
18.3 
17.3 


6 

5.1 

5.6 
*5.'i 


5.9 
5.2 


5.3 
5.3 
5.6 


5.1 
6.2 
5.6 


5.3 


4.8 
5.3 


5.8 
4.7 
4.6 


5.8 
6.1 


5.2 

5 

5.8 


4.8 
5.2 
4.6 


5 

4.9 

5.2 


5 

4.7 


5 

5.1 

4.9 


3 
2.8 


2 
2.6 


2.6 

2 

2.4 


2.4 
2.8 
2.9 


2.5 
2.5 
3.2 


1.8 
1.3 


1.2 
1.4 

1.2 


1.3 

l.S 


1.5 
1.4 
1.6 


1.9 
1.2 
1.6 


1.5 
1.6 
2 


1.7 
1.7 


1.6 
1.4 


1.4 
1.2 
1.5 


13.4 
14 


12.8 
14.2 
13.7 


14.4 
13.1 
14.2 


15.4 
15.7 
13.8 


13 

13.4 
12.6 


14.5 


13.7 
13.5 


13.6 
14.4 


13.4 
13 


13.7 
13.8 
15.3 


14.1 
13.6 
14.1 


13.7 
14.1 
13.6 


14 

13.2 


13 

12.2 


12.3 
13.3 
13 


Type. 
Adult. 


Do. 
Adult;  type. 
Adult. 


Old  adult. 

Adult. 
Do. 


Adult;  type. 
Adult. 
Do. 


Do. 
Do. 
Do. 


Old  adult; 

type. 
Adult. 
Do. 


Adult;  type. 
Adult. 
Do. 


Adult;  type. 
Adult. 


Adult;  type. 
Adult. 
Do. 


Adult;  type. 
Adult. 
Do. 


Do. 
Do. 
Do. 


Adult;  type. 
Adult. 


Do. 
Do. 


Adult:  type. 
Adult. 
Do. 


6  Collection  British  Mus.:  measured  bv  Oldfield  Thomas. 

•  Collection  of  S.  G.  Jewett. 

7  Abnormally  broad  for  the  species. 

•  Collection  Mus.  Vert.  Zool..  Univ.  California. 
•Collection  Field  Mus.  Nat.  Hist. 

10  Collection  Amer.  Mus.  Nat.  Hist. 


Plate  III. 

[All  natural  size.] 

Fig.  1. — Ochotona   princeps  princeps,  S   adult,  Henry  House,  Alberta  (No.  81537, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  2. — Ochotona  collaris,  d  adult,  Seward  Creek,  near  Eagle,  Alaska  (No.   131266, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  3. — Ochotona  princeps  uinta,    tf   adult,  Uinta   Mountains,  Utah    (No.    25608, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  4 — Ochotona  princeps  cuppes,  <?  subadult  (type),  Monashee  Divide,  Gold  Range, 

British  Columbia  (No.  7389,  Mus.  Comp.  Zool.). 
Fig.  5. — Ochotona  princeps  fenisex,  <S  adult,  Horseshoe  Basin  [near  Mount  Chopaka], 

Washington  (No.  848,  collection  S.  G.  JewettV 
Fig.  6. — Ochotona  princeps  nevadensis,  9    adult  (type),  Ruby  Mountains,   Nevada 

(No.  94213,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  7. — Ochotona  princeps    lutescens,    <$   adult,  Mount    Inglesmaldie,    near  Banff, 

Alberta  (No.  108652,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  8. — Ochotona  princeps  brunnescens,  c?  adult,  Keechelus,  Washington  (No.  227258, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  9. — Ochotona  princeps  lemhi,    tf   adult,  Lemhi  Mountains,  Idaho  (No.  31204, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 

52 


North  American  Fauna  No.  47,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  III. 


Skulls  of  Ochotona  princeps  and  O.  collaris,  Dorsal  Views. 


1.  O.  p.  princeps. 

2.  O.  collaris. 

3.  O.  p.  uinta. 


4.  O.  p.  cuppes. 

5.  O.  p.  fenisex. 

6.  O.  p.  nevadensis. 


7.  O.  p.  lutescens. 

8.  O.  p.  brunnescens. 

9.  O.  p.  lemhi. 


North  American  Fauna  No.  47,  U.  S.  Dept.  Agr.,  Biological  Survey.  PLATE  IV. 


Skulls  of  Ochotona  princeps  and  O.  schisticeps,  Dorsal  Views. 

1.  O.  p.  saxatilis.  4.  O.  p.  flgginsi.  7.  O.  p.  nigrescens. 

2.  O.s.  schisticeps.  5.  O.  s.  taylori.  8.  O.  s.  goldmani. 

3.  O.s.  muiri.  6.  O.  s.  fuscipes.  9.  O.  s.  cinnamomea 


Plate  IV. 
[All  natural  size.] 

Fig.  1. — Ochotona  princeps  saxatiiis,  9  adult,  Mount  Lincoln,  Colorado  (No.  205793 

U.  S.  Nat.  Mua.,  Biological  Survey  collection). 
Fig.  2. — Ochotona  schisticeps  schisticeps,  adult,  Donner  Pass,  California  (No.  25561, 

Mus.  Vert.  Zool.,  Univ.  of  California;. 
Fig.  3. — Ochotona  schisticeps  muiri,  <?   adult,  Vogelsang  Lake,  Yosemite  National 

Park,  California  (No.  23473,  Mus.  Vert.  Zool.). 
Fig.  4. — Ochotona  princeps  figginsi,  J1  adult,  Trappers  Lake,  Garfield  County,  Colo- 
rado (No.  4031,  collection  of  E.  R.  Warren). 
Fig.  5. — Ochotona  schisticeps  taylori,    $    adult,  Warner   Mountains,  California  (No. 

11294,  Mus.  Vert.  Zool.). 
Fig.  6. — Ochotona  schisticeps  fuscipes,    <?   adult  (type),  Parowan    Mountains,  Utah 

(No.  158094,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  7. — Ochotona   princeps   nigrescens,    $   adult    (type),   Jemez    Mountains,    New 

Mexico  (No.  147976,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  8. — Ochotona  schisticeps  goldmani,  J  adult  (type),  Snake  River  Desert,  Idaho 

(No.  236408,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  9. — Ochotona  schisticeps  cinnamomea,    d   adult,  Beaver  Mountains,  Utah  (No. 

158086,  U.  S.  Nat.  Mus..  Biological  Survey  collection). 

53 


Plate   V. 

[All  natural  size.] 

Fig.  1. — Ochotona  princeps  princeps,    c?   adult,  Henry  House,  Alberta  (No.  81537, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  2. — Ochotona  collaris,  c?  adult,  Seward  Creek,  near  Eagle,  Alaska  (No.  131266, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  3. — Ochotona  princeps  uinta,  d*  adult,  Uinta  Mountains,  Utah  (No.  25608,  U.  S. 

Nat.  Mus.,  Biological  Survey  collection). 
Fig.  4. — Ochotona  princeps  cuppes,  cf  subadult  (type),  Monashee  Divide,  Gold  Range, 

British  Columbia  (No.  7389,  Mus.  Comp.  Zool.). 
Fig.  5. — Ochotona  princeps  fenisex,  <?  adult,  Horseshoe  Basin  [near  Mount  Chopaka], 

Washington  (No.  848,  collection  S.  G.  Jewett). 
Fig.  6. — Ochotona  princeps  nevadensis,  9   adult  (type),  Ruby  Mountains,  Nevada 

(No.  94213,  U.  S.  Nat.  Mus.,  Biological  Survey  collection ). 
Fig.  7. — Ochotona  princeps   lutescens,    &    adult,  Mount   Inglesmaldie,  near  Banff, 

Alberta  (No.  108652,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  8. — Ochotona  princeps  brunnescens,  <?  adult,  Keechelus,  Washington  (No.  227258, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  9 — Ochotona  princeps  lemhi,  cf  adult,  Lemhi  Mountains,  Idaho  (No.  31204,  U.  S. 

Nat.  Mus.,  Biological  Survey  collection). 

54 


North  American  Fauna  No.  47,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  V. 


Skulls  of  Ochotona  princeps  and  O.  collaris,  Ventral  Views. 


1.  O.  p.  princeps. 

2.  O.  collaris. 

3.  O.  p.  uinta. 


4.  O.  p.  cuppes. 

5.  O.  p.  fenisex. 

6.  O.  p.  nevadensis 


7.  O.  p.  lutescens. 

8.  O.  p.  brunnescens. 

9.  O.  p.  lemhi. 


North  American  Fauna  No.  47,  U.  S.  Dept.  Agr.,  Biological  Survey.  PLATE  VI. 


mbbw  h  ,• 

■W  ^Fm 

1  im 

:^g?l. 

bt!  ^r      r   ^B 

Wwt  *1H 

bV1^ 

Bv  w    7              *H 

K/->  11 

pJrl 

Ii  •  iM 

HbIbh 

B^?W 

Hv^H 

blA&T  ixmbi 

^F  .  j»  J<J 

v  ll  ^B 

l^,Jr  -^ 

y  jjP  ■  :?!■ 

Hf       IV  ~^^        "    •  >^HJ 

Br    J?  ii     •      ^Bl 

■  k^T J '  "^  ■ 

Bf  V-^  i      'Ht^^B^ 

BfAl  * 

■#      TfW    «1 

H  Kiy  #cbU 

lf;»p£B| 

BiB'      ^v^jMfct  ^|BJ 

IV  '  *  IWSl  JH 

wMs*^~-  ■  ^«B 

BK^J 

BBW^C      ?^^H 

B^ "^  ^Bk      ■    ^W 

I*fl 

dBT^HJ 

BY     T     ^f 

bP^^ 

Br     -,,*v      ».  ^B 

BT                    *  1H 

Hgi  ^|  ill 

■      ^B^          Bf        IIbVibV 

Iff    *  *  *    *Tb1 

Br    4k '^ 
Wjkjgjwk    tiB>H 

If  *ir    ll 

BiflT   '^jk'^v^iBI 

Vll&H 

IT  -Jl     ■ 

B«^     *'  'JB           HBl 
Bk          £B\           B 

s^SggBBl 

B^     JF' ' '^L,     ^B 

Skulls  of  Ochotona  princeps  and  0.  schisticeps,  Ventral  Views. 

1.  O.  p.  saxatilis.  4.  0.  p.  figginsi.  7.  O.  p.  nigrescens. 

2.  O.  s.  schisticeps.  5.  O.  s.  taylori.  8.  O.  s.  goldmani. 

3.  O.s.  muiri.  6.  O.  s.  fuscipes.  9.  O.  s.  cinnamomea. 


Plate  VI. 

[All  natural  size.] 

Fig.  1. — Ochotona  princeps  saxatilis,  9  adult,  Mount  Lincoln,  Colorado  (No.  205793, 

U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  2. — Ochotona  schisticeps  schisticeps,  adult,  Donner  Pass,  California  (No.  25561, 

Mus.  Vert.     Zool.,  Univ.  of  California). 
Fig.  3. — Ochotona  schisticeps  muiri,    d   adult,  Vogelsang  Lake,  Yosemite  National 

Park,  California  (No.  23473,  Mus.  Vert.  Zool.). 
Fig.  4. — Ochotona  princeps  fig ginsi,  <?  adult,  Trappers  Lake,  Garfield  County,  Colo- 
rado (No.  4031,  collection  of  E.  R.  Warren). 
Fig.  5. — Ochotona  schisticeps  taylori,    9   adult,  Warner  Mountains,  California   (No. 

11294,  Mus.  Vert.  Zool.). 
Fig.  6. — Ochotona  schisticeps  fuscipes,  <$  adult  (type),  Parowan  Mountains,  Utah  (No. 

158094,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  7. — Ochotona  princeps  nigrescens,  <$  adult  (type),  Jemez  Mountains,  New  Mexico 

(No.  147976,  U.  S.  Nat.  Mus.,  Biological  Survey  collection^. 
Fig.  8. — Ochotona  schisticeps  goldmani,  $  adult  (type),  Snake  River  Desert,  Idaho 

(No.  236408,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  9. — Ochotona  schisticeps  cinnamomea,  <?   adult,  Beaver  Mountains,  Utah  (No. 

158086,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 

55 


INDEX 


[New  species  in  bold-face  type.] 


albata,  Ochotona,  44. 
alpinus,  Lepus,  7. 

Pika,  7. 
brooksi,  Ochotona,  30. 
brunnescens,  Ochotona,  31. 
cinnamomea,  Ochotona,  46. 
collaris,  Lagomys,  34. 

Ochotona,  34. 
Conothoa,  7,  8. 
cuppes,  Ochotona,  27. 
dauuricus,  Lepus,  7. 
fenisex,  Ochotona,  28. 
figginsi,  Ochotona,  21. 
fumosa,  Ochotona,  33. 
fuscipes,  Ochotona,  47. 
goldinani,  Ochotona,  40. 
incana,  Ochotona,  25. 
jewetti,  Ochotona,  42. 
Lagomys,  7,  8. 
collaris,  34. 
minimus,  28. 
ogotona,  7,  8. 
schisticeps,  38. 
lemhi.  Ochotona,  16. 
Lepus  alpinus,  7. 
t        dauuricus,  7. 

ogotona,  7. 
:         princeps,  8,  12. 

pusillus,  8. 
levis,  Ochotona,  16. 
lutescens,  Ochotona,  15. 
minimus,  Lagomvs,  28. 

Ochotona,  28,  31. 
minor,  Ochotona,  7. 
muiri,  Ochotona, 43. 
nevadensis,  Ochotona,  21. 
nigrescens,  Ochotona,  26. 
Ochotona  collaris,  34. 
minimus,  28,  31. 
minor,  7. 

princeps  brooksi,  30. 
brunnescens,  31. 
cuppes,  27. 
fenisex,  28. 
figginsi,  21. 
fumosa,  33. 
incana,  25. 
lemhi,  16, 
levis,  16. 
lutescens,  15. 
nevadensis,  21. 
nigrescens,  26. 
princeps,  12. 
saxatilis,  23. 
uinta,  19. 
ventorum,  18. 


Ochotona  roylii,  7,  8. 

schisticeps  albata,  44 
cinnamomea,  46 
fuscipes,  47. 
goldmani,  40. 
jewetti,  42. 
muiri,  43. 
schisticeps,  38. 
sheltoni,  45. 
taylori,  39. 
Ogotoma,  7,  8. 

pallasii,  7,  8. 
ogotona,  Lagomys,  7,  8. 

Lepus,  7. 
pallasii,  Ogotoma,  7,  8. 
Pika,  7,  8,  9. 

alpinus,  7. 
Pika,  Alberta,  15. 

Ashnola,  28. 

Bangs,  27. 

Beaver  Mountains,  4fi. 

Blue  Mountains,  42. 

Cascade,  31. 

Collared,  34. 

Colorado,  23. 

Dusky,  33. 

Figgins,  21. 

Gray-headed,  38. 

Hollister,  16. 

Jemez  Mountains,  26. 

Lava-bed,  40. 

Lemhi,  16. 

Mount  Whitney,  44. 

New  Mexico,  25. 

Parowan  Mountains,  47. 

Rocky  Mountains,  12. 

Ruby  Mountains,  21. 

Shuswap,  30. 

Taylor,  39. 

Uinta,  19. 

White  Mountains,  45. 

Wyoming,  18. 

Yosemite,  43. 
princeps,  Lepus  (Lagomys),  8,  12. 

Ochotona,  12. 
pusillus,  Lepus,  8. 
roylii,  Ochotona,  7,  8. 
saxatilis,  Ochotona,  23. 
schisticeps,  Lagomys,  38. 

Ochotona,  38. 
sheltoni,  Ochotona,  45. 
taylori,  Ochotona,  39. 
uinta,  Ochotona,  19. 
ventorum,  Ochotona,  18. 


ADDITIONAL  COPIES 

OF  THIS  PUBLICATION  MAY  BE  PROCURED  FROM 

THE  SUPERINTENDENT  OF  DOCUMENTS 

GOVERNMENT  PRINTING  OFFICE 

WASHINGTON,  D.  C. 

AT 

15  CENTS  PER   COPY 

V 


North  American  Fauna  No.  48,  U.  S.  Dept.  Agr.,  Biological  Survey 


Plate 


Red  Tree  Mouse  (Phenacomys  longicaudus) 

Photographed  near  Carlotta,  Calif. 


U.  S.  DEPARTMENT  OF  AGRICULTURE 

BUREAU  OF  BIOLOGICAL  SURVEY 


NORTH  AMERICAN  FAUNA 


!N"o.  48 


[Actual  date  of  publication,  October  12,  1926] 


VOLES  OF  THE  GENUS  PHENACOMYS 

I.  REVISION  OF  THE  GENUS  PHENACOMYS 

II.  LIFE  HISTORY  OF  THE  RED  TREE  MOUSE 

(PHENACOMYS  LONGICAUDUS) 


BY 

A.  BRAZIER  HOWELL 

SCIENTIFIC  ASSISTANT,  DIVISION  OF  BIOLOGICAL  INVESTIGATIONS 
BUREAU  OF  BIOLOGICAL  SUBVEY 


WASHINGTON 

GOVERNMENT  PRINTING  OFFICE 

1926 


Is 


SUPERINTENDENT  OF  DOCUMENT* 

/ 

OCT.  26  1926 


LETTER  OF  TRANSMITTAL 


United  States  Department  of  Agriculture, 

Bureau  of  Biological  Survey, 
Washington,  D.  C,  February  10,  1926. 
Sir  :  I  have  the  honor  to  transmit  herewith,  with  the  recommendation  that 
it  be  published  as  No.  48  in  the  series  of  the  North  American  Fauna,  a  report 
on  the  voles  of  the  American  genus  Phenacomys,  by  Alfred  Brazier  Howell,  an 
assistant  of  this  bureau.  The  report  consists  of  two  parts,  (1)  a  technical  revi- 
sion of  the  genus,  and  (2)  the  results  of  a  study  of  one  of  the  more  interesting 
and  scientifically  important  species,  concerning  which  the  author  has  had 
exceptional  opportunities  for  observation. 

These  investigations  constitute  an  intensive  study  of  the  subject  and  are 
based  on  an  examination  of  practically  all  the  specimens  extant  in  public  and 
private  collections  in  this  country  and  Canada. 
Respectfully, 

E.  W.  Nelson,  Chief  of  Bureau. 
Hon.  W.  M.  Jardine, 

Secretary  of  Agriculture. 
ii 


CONTENTS 


Revision  of  the  genus  Phenacomys- 

Introduction 

Geographic  distribution 

Habits 

Material    examined    and    de- 
sired   

Explanations       of      measure- 
ments   and   color   terms 

Genus  Phenacomys  Merriani 

History 

Key  to  the  genus 

Characters  

Variation 

Probable  affinities 

Groups 

List  of  species  and  subspecies, 
with  their  type  localities. 
Key  to  species  and  subspecies — 
Descriptions  of  species  and  sub- 
species  

Phenacomys    intermedins 
group 


Page 

I.  Revision  of  the  genus — Cont'd. 

Descriptions  of  species — Cont'd. 

Phenacomys  ungava  group 25 

Phenacomys  albipes  group 30 

Phenacomys     longicaudus 

group 32 

II.  Life  history  of  the  red  tree  mouse 

(Phenacomys    longicaudus)-  39 

Introduction 39 

Distributional  factors 40 

Investigational  methods 41 

Area  investigated 41 

Nests 43 

Construction  and  site 45 

Young 48 

Pood 50 

Habits 55 

Other  behavior 57 

Enemies 59 

Bibliography  of  Phenacomys 60 

Index 65 


in 


ILLUSTRATIONS 


PLATES 

Page 
Plate  1.  Red  tree  mouse   (Phenacomys  longicaudus) Frontispiece. 

2.  Skulls  of  Phenacomys 36 

3.  Skulls  of  Phenacomys 36 

4.  Typical  habitat  of  Phenacomys  longicaudus 42 

5.  Typical  nesting  tree  of  Phenacomys  lonyieaudus 43 

6.  Fig.  1 — Female  red  tree  mouse  at  nest.     Fig.  2. — Female  red  tree  mouse 

in  fir  tree 40 

7.  Fig.  1. — Young  of  red  tree  mouse.     Fig.  2. — Day's  food  of  one  red  tree 

mouse.     Fig.  3. — Cross  section  of  fir  needle,  a  staple  food  of  Phenaco- 
mys   longicaudus 47 

TEXT    FIGURES 

Fig.  1.  Map  of  approximate  distribution  of  the  genus  Phenacomys 2 

2.  Molars  of  Phenacomys  intermedins  intermedins 8 

3.  Extreme  change  in  enamel  pattern  of  molars  of  Phenacomys  mackenzii,  due 

to  age  and  wear 8 

4.  Extremes  of  variation  of  palates  of  Phenacomys 1<> 

5.  Map  of  distribution  of  the  Phenacomys  intermedins  group 14 

6.  Map  of  distribution  of  the  Phenacomys  ungava  group 24 

7.  Extremes  of  variation  in  anterior  trefoil  and  inclosure  of  triangles  in  first 

lower  molars  of  Phenacomys  mackenzii 29 

8.  Map  of  distribution  of  Phenacomys  albipes 30 

9.  Map  of  distribution  of  the  Phenacomys  longicaudus  group 32 

10.  Extremes  of  variation  in  enamel  pattern  of  Phenacomys  longicaudus 33 

11.  Diagram  of  area  harboring  colony  of  Phenacomys  longicau<kis 42 

IV 


No.  48  NORTH  AMERICAN  FAUNA  October  12,  1926 

VOLES  OF  THE  GENUS  PHENACOMYS 

I.  REVISION  OF  THE  GENUS  PHENACOMYS 

By  A.  Brazier  Howell 


INTRODUCTION 

The  rodent  genus  Phenacomys  belongs  to  the  section  Microti,  sub- 
family Microtinae,  family  Cricetidae,  and  so  far  as  known  it  is 
confined  to  the  New  World.  As  its  main  range  lies  in  regions  com- 
paratively unfrequented  by  man,  and  as  the  animals  are  usually  de- 
cidedly rare,  study  material  has  been  slow  to  accumulate. 

There  has  been  little  need  of  a  colloquial  name  for  this  genus,  but 
when  one  has  been  desired,  the  animals  have  usually  been  called 
"  lemming  mice."  They  are  not  lemmings,  however,  nor  more  closely 
related  to  the  latter  than  are  many  other  genera  of  voles,  and  the 
name  is  misleading.  Because  of  a  diversity  of  habits  and  habitat,  no 
one  name,  strictly  English,  can  well  be  applied,  except  to  two 
arboreal  long-tailed  species,  which  are  termed  "  tree  mice,"  and  it 
is  thought  better  for  the  short-tailed  terrestrial  species  to  retain  the 
Latin  generic  name  for  its  colloquial  equivalent. 

Although  it  is  hoped  that  everything  possible  that  might  prove 
of  real  value  to  the  investigator  has  been  included  in  the  following 
revision,  an  effort  has  been  made  to  exclude  all  minute  descriptions 
of  an  attempted  exactitude  impossible  of  attainment  and  often 
highly  confusing  to  the  student. 

GEOGRAPHIC  DISTRIBUTION 

In  general,  PJienacomys  is  found  in  scattered  localities  through- 
out practically  the  whole  of  the  upper  Canadian  and  the  Hudson- 
ian  Zones  of  Canada;  apparently  it  is  absent  from  Alaska;  and  in 
the  United  States  it  is  found  in  the  Rocky  Mountain  system  as  far 
south  as  northern  New  Mexico,  on  the  boreal  summits  of  isolated 
mountain  ranges  of  the  Great  Basin,  and  south  in  the  Sierra  Nevada 
at  least  to  the  Yosemite  Park,  Calif. ;  and  also  in  the  coastal 
belt  of  Oregon  and  northern  California.     (See  map,  fig.  1.) 

HABITS 

The  habitats  of  members  of  the  genus  are  extremely  diverse  in  char- 
acter and  vary  from  sandy  blueberry  barrens,  boreal  heather  mead- 
ows, dry  woods  of  the  far  north,  and  sphagnum  bogs,  to  a  riparian 
association  on  the  Pacific  coast  and  an  arboreal  habitat  among  ever- 
green forests.  Habits  are  equally  diverse,  but  very  little  is  known 
of  them.  Well-defined  runways  seldom  occur  but  are  occasionally 
made  when  the  animals  are  living  among  beds  of  heather  or  other 
very  dense  growth. 

1 


NORTH   AMERICAN   FAUNA 


[No.  48 


The  short-tailed  species,  usually  living  in  dry  situations  where 
cover  is  ample,  probably  feed  on  a  variety  of  seeds,  tender  growth, 
and  other  relatively  soft  or  brittle  foods  for  which  their  teeth  are 
best  suited.  The  majority  of  existing  specimens  of  Phenacomys 
alhipes  were  obtained  on  the  immediate  banks  of  little  streams  in  the 
forest,  so  that  its  habitat  seems  to  be  chiefly  riparian  in  character, 
but  its  food  is  unknown.  Two  long-tailed  species,  P.  longicaudus  and 
P.  silvicola,  are  definitely,  although  not  exclusively,  arboreal.  Their 
food,  so  far  as  known,  consists  chiefly  of  the-  fleshy  parts  of  the 
needles  of  the  conifers  in  which  they  live,  and  the  medullary  portions 
constitute  the  chief  material  from  which  their  nests  are  constructed. 
Some  tender  bark  also  is  consumed. 


Fig.  1. — Approximate  distribution  of  the  genus  Phenacomys 

In  all  probability,  more  than  one  litter  of  young  is  raised  during 
the  warmer  months  each  year,  but  exact  data  upon  the  subject  are 
lacking.  There  are  records  among  Biological  Survey  field  notes 
of  females  containing  from  4  to  9  embryos ;  the  usual  number  seems 
to  be  5  or  6,  but  is  fewer  in  the  case  of  P.  longicaudus,  which  the 
writer  has  found  to  have  1  to  3  young,  and  which  must  also  have 
fewer  predatory  enemies  to  contend  with. 

It  is  believed  that  no  member  of  the  genus  hibernates,  and  that 
there  is  no  diminution  of  activity  because  of  cold  weather.  With 
rare  exceptions  the  ranges  of  the  short-tailed  species  are  situated  in 
regions  unsuited  to  agriculture,  and  they  could  be  of  little  or  no 
economic  importance  even  were  they  far  more  abundant  than  is 
now  the  case.     One  long-tailed  species,  P.  longicaudus,  is  said  by 


1926]  REVISION    OF   GENUS   PHENACOMYS  6 

Taylor  (1915b,  p.  142)  1  occasionally  to  girdle  and  kill  the  top  of 
its  nest  tree,  but  it  is  not  certain  that  the  mice  are  directly  responsible 
for  this  damage. 

MATERIAL    EXAMINED   AND    DESIRED 

This  revision  is  based  upon  the  study  of  317  specimens  of  the 
genus  Phenacomys,  practically  all  now  available,  except  some  juven- 
ile material  of  the  better-known  forms,  which  it  was  thought  un- 
necessary to  examine.2 

For  the  generic  revision  made  by  G.  S.  Miller,  jr.,  in  1897  (1897b, 
pp.  77-87)  the  material  available  was  so  very  scanty  that  a  proper 
understanding  of  relationships  and  the  degree  of  individual  varia- 
tion was  at  that  time  hardly  possible,  and  in  consequence,  further 
revision  has  been  needed  for  some  years.  Although  material  is  now 
more«abundant,  it  is  far  from  all  that  could  be  desired  in  this  regard 
and  will  probably  remain  so  for  many  years.  Upon  the  study  of 
hundreds  of  additional  specimens,  still  another  revision  will  un- 
doubtedly prove  desirable. 

Examples  of  the  genus  Phenacomys  will  always  be  eagerly  sought 
by  collectors  because  of  their  rarity.  Specimens  are  especially 
needed  from  a  number  of  critical  localities,  for  it  is  folly  to  claim 
that  a  perfect  understanding  of  a  species  or  subspecies  can  be  gained 
from  one  or  two  specimens.  More  material  is  urgently  needed  from 
the  whole  of  the  eastern  half  of  Canada,  especially  Ungava,  Quebec, 
and  the  area  to  the  south  of  Hudson  Bay,  from  the  Hudsonian  Zone 
of  New  Brunswick,  and  from  the  New  England  States. 

In  the  western  portion  of  North  America  a  special  effort  should 
be  made  to  obtain  a  small  series  of  adults  from  the  basaltic  plateau 
to  the  northward  of  Kamloops,  British  Columbia — the  type  locality 
of  intermedins.  Material  from  several  places  in  the  Bitterroot 
Mountains  between  Montana  and  Idaho,  as  well  as  series  from  the 
Lemhi  Mountains  (the  type  locality  of  "  orophilus  ")  of  the  latter 
State,  and  from  some  one  spot  in  Colorado,  are  also  greatly  to  be 
desired.  A  small  series  from  the  Blue  Mountains  of  Oregon  would 
be  worth  an  effort,  as  well  as  from  various  localities  in  the  Cascade 
Mountains  of  that  State. 

Additional  skins  of  albipes,  although  desirable  because  of  their 
rarity,  would  not  add  materially  to  our  S37stematic  knowledge  of  the 
species,  but  specimens  of  silvicola  would  be  of  the  greatest  impor- 
tance. The  recent  discovery  of  that  species  is  an  indication  of  the 
possibilities  of  the  region,  and  all  collectors  in  the  forests  of  the 
humid  Northwest  should  investigate  everything  encountered  that 
might  prove  to  be  the  nest  of  an  arboreal  phenacomys. 

1  Literature  citations  in  parentheses  refer  to  the  bibliography  beginning  on  page  58. 

2  The  writer's  thanks  are  hereby  tendered  G.  S.  Miller,  jr.,  curator  of  mammals,  of  the 
United  States  National  Museum,  for  the  unrestricted  use  of  the  material  in  his  care.  For 
the  loan  of  pertinent  material,  he  is  also  greatly  indebted  to  the  Museum  of  Comparative 
Zoology  (29  specimens),  through  G.  M.  Allen;  the  Academy  of  Natural  Sciences  of  Phila- 
delphia (5  specimens),  through  Witmer  Stone;  the  Field  Museum  of  Natural  History  (10 
specimens),  through  W.  H.  Osgood;  the  Museum  of  Vertebrate  Zoology  (9  specimens), 
through  Joseph  Grinnell ;  the  Victoria  Memorial  Museum  (11  specimens),  through  R.  M. 
Anderson;  the  Provincial  Museum  (8  specimens),  through  Francis  Kermode  ;  the  Ameri- 
can Museum  of  Natural  History  (4  specimens),  through  H.  E.  Anthony:  the  Oregon  Agri- 
cultural College  (1  specimen),  through  H.  M.  Wight;  Washington  State  College  (10 
specimens),  through  W.  T.  Shaw;  and  to  D.  R.  Dickey  (9  specimens),  S.  G.  Jewett  (10 
specimens),  E.  R.  Warren  (2  specimens),  W.  S.  Saunders  (1  specimen),  and  A.  Walker 
(3  specimens).  Grateful  acknowledgment  is  made  also  to  various  members  of  the  staff 
of  the  Biological  Survey  and  to  G.  S.  Miller,  jr.,  for  advice. 


4  NORTH   AMERICAN   FAUNA  [No.  48 

EXPLANATIONS  OF  MEASUREMENTS  AND  COLOR  TERMS 

All  measurements  given  are  in  millimeters.  With  the  exception 
of  a  few  alcoholics  as  noted,  the  external  measurements  were  taken 
by  the  collector  in  the  field  from  the  specimen  in  the  flesh.  Since 
collectors  frequently  measure  the  same  specimen  differently,  too  much 
reliance  should  not  be  placed  on  such  figures,  nor  on  differences  indi- 
cated in  the  size  of  the  sexes  when  very  few  individuals  are  con- 
cerned. For  this  reason  slight  differences  in  body  size  between 
closely  related  forms  are  not  emphasized  unless  supported  by  ample 
evidence. 

Only  those  skulls  (except  types)  were  measured  which  were  suffi- 
ciently adult  to  show  an  indication  beyond  the  mandibular  alveoli 
of  the  rooted  nature  of  the  molars,  the  measurements  being  taken  by 
the  writer  with  dial  calipers  on  which  divisions  were  in  tenths  of  a 
millimeter. 

Condylobasilar  length, — Distance  from  the  posterior  border  of  the 
condyle  to  the  posterior  edge  of  the  alveolus  of  the  incisor  on  the 
corresponding  side  of  the  skull. 

Nasals. — Greatest  length  of  nasals. 

Interorbital  breadth. — Shortest  distance  between  the  orbits. 

Zygomatic  breadth, — Distance  between  the  zygomatic  arches  at 
their  widest  point. 

Lambdoidal  width. — Greatest  width  of  posterior  portion  of  brain 
case  measured  across  the  lambdoidal  ridges. 

Incisive  foramvina. — Greatest  length  of  incisive  foramina. 

Maxillary  tooth  row. — Aggregate  length  of  right-hand  upper 
molar  series. 

Height. — Height  of  skull,  measured  from  a  plane  passing  from  the 
inferior  surface  of  the  bullae  to  the  inferiormost  molar  projection. 

Listed  specimens  are  in  the  United  States  National  Museum  unless 
otherwise  stated. 

For  the  reason  that  it  is  often  impossible  to  apply  a  single  precise 
color  term  to  an  animal  of  the  agouti  pattern,  the  color  names  given 
herein,  including  those  of  Ridgway,3  are  meant  to  be  only  approxi- 
mate. Although  such  terms  as  "  warm  brown  "  and  "  grayish  brown  " 
are  admittedly  unsatisfactory,  it  is  at  least  felt  that  they  do  not 
indicate  a  misleading  exactness  impossible  of  attainment  in  the  case 
of  most  small  mammals. 

Genus  PHENACOMYS  Merriam 
Phenacomys  Merriam,  North  Ainer.  Fauna  No.  2,  p.  28,  1889. 

HISTORY 

The  characters  Avhereby  the  present  genus  is  distinguished  from 
other  microtines  were  first  appreciated  by  C.  Hart  Merriam,  who 
named  Phenacomys  as  a  new  genus  in  1889,  designating  intermedins 
as  the  type  species,  and  at  the  same  time  describing  as  additional 
new  species  ungava,  "  celatus"  and  "  latimanus"  each  from  a  single 
spirit-specimen  of  poor  quality.  In  the  following  year  F.  W.  True 
described  longicaudus  from  an  Oregon  example  with  badly  damaged 

3  Ridgway,  Robert.     Color  standards  and  color  nomenclature.     43  pp.,  illus.     1912. 


1926]  REVISION    OF   GENUS  PHENACOMYS  O 

skull ;  and  in  1891  Merriam  described  "  orophilus  "  from  Idaho,  based 
on  an  immature  individual  of  which  no  adult  topotypes  have  as  yet 
been  collected.  In  1894  J.  A.  Allen  named  "  truei  "  from  a  skin  with 
fragmentary  skull  then  supposed  to  have  come  from  the  Black  Hills 
of  South  Dakota,  but  since  ascertained  to  have  been  taken  in  Wyo- 
ming. In  1895,  S.  N.  Rhoads  described  still  another  species,  from 
southern  British  Columbia,  under  the  name  "  oramontis'''>\  and  in 
1897  Merriam  (1897b)  characterized  " prebleV  from  Colorado.  In 
1899  J.  A.  Allen  named  "  constablei "  from  British  Columbia,  and 
D.  G.  Elliott  (1899  b),  olympicus,  from  Washington,  as  well  as 
"  Microtus  (Lagurus)  pumilus"  which  proved  to  be  a  juvenile 
Phenacomys.  The  following  year  Outram  Bangs  named  crussus  as  a 
subspecies  of  "  celatus"  from  Labrador.  Then,  in  1901,  came  albipes 
by  Merriam,  and  a  year  later,  machenzii  by  E.  A.  Preble  (1902  a). 
In  1921  silvicola  was  described,  and  in  1923,  levis  and  celsus,  these 
three  by  the  writer. 

Thus,  18  forms  of  Phenacomys  have  been  described  as  new,  all 
but  3  of  them  as  full  species.  Of  these,  10  are  herein  recognized, 
the  remainder  being  placed  in  synonymy. 

In  1894,  Nehring  (1894,  p.  346)  referred  to  the  genus  Phenacomys 
some  fossil  material  from  the  late  Pliocene  forest  beds  of  England, 
previously  described  by  Newton  as  Arvicola  intermedins.  This  is 
of  interest  in  the  present  connection,  because  the  teeth  of  this 
animal  have  roots  and  because  it  is  the  only  case  in  which  Old 
World  material  has  been  ascribed  to  this  genus.  The  type  of 
enamel  pattern,  however,  definitely  precludes  the  possibility  of  this 
material  being  referable  to  Phenacomys  as  at  present  defined,  and 
it  has  since  been  included  with  several  other  Pliocene  species  from 
various  parts  of  Europe  in  the  extinct  genus  Mimomys  Major  (1902, 
p.  102).  None  of  these  animals,  however,  is  believed  to  be  directly 
ancestral  to  either  Microtus  or  Phenacomys.  (See  Hinton,  in 
Barrett-Hamilton,  1913,  p.  385.) 

In  Miller's  revision  of  the  genus  Phenacomys  (1897b,  p.  77),  six 
of  the  forms  that  had  then  been  described  were  recognized.  The 
author  of  this  revision  was  seriously  handicapped  in  having  so 
few  specimens  available  (64  in  all),  especially  from  any  single 
locality,  and  a  proper  understanding  of  variations  and  relation- 
ships could  not  be  obtained. 

In  1915,  W.  P.  Taylor  (1915b,  p.  Ill)  described  a  subgenus 
Arborimus  to  include  longicaudus  (the  type  species)  and  albipes. 
In  addition,  several  other  races  were  discussed  and  an  effort  made 
to  define  their  relationships.  Unquestionably,  longicaudus  and 
silvicola,  which  latter  was  unknown  in  1915,  constitute  a  readily 
recognizable  group,  but  the  characters  on  which  the  subgenus  were 
based  are  all  weak,  ill-defined,  and  not,  in  the  opinion  of  the  writer, 
of  sufficient  importance  to  merit  subgeneric  position,  especially 
when  one  considers  what  a  compact  unit-division  is  included  within 
the  genus  as  it  now  stands.  Furthermore,  albipes  is  intermediate 
in  many  respects  between  the  short-tailed  members  of  the  genus  and 
the  arboreal  species,  and  this  fact  would  still  further  complicate  the 
establishment  of  a  subgenus.  In  short,  it  is  considered  that  the 
recognition  of  a  subgeneric  division  within  the  genus  could  be  based 
only  upon  weakly  defined  characters,  and  that  such  a  step  would 
complicate,  rather  than  clarify,  our  understanding  of  the  subject. 
93S900— 26 .2 


b  NORTH   AMERICAN   FAUNA  [No.  48 

KEY  TO  THE   GENUS 

Molars  of  adults  rooted Genera  Ondatra,  Evotomys,  Phenacomys. 

Skull  in  adults  less  than  40  mm.  long Evotomys,  Phenacomys. 

Posterior  border  of  palate  not  shelf-like :  inner  reentrant  angles  of 

lower  molars  much  deeper  than  outer Phenacomys. 

CHARACTERS 

Generic  diagnosis. — The  most  important  characters  by  which  the 
genus  may  be  distinguished  are  the  presence  of  rooted  molars  when 
adult  (versus  all  other  recent  genera  but  Evotomys  and  Ondatra — or 
Fiber  of  authors)  and  by  not  having  a  palate  that  is  transversely 
continuous  and  shelf-like  (versi\^Evoto??iys) .  The  enamel  pattern  of 
the  molars  is  unique,  chiefly  in  the  great  depth  of  the  reentrant  angles, 
as  compared  with  the  external,  of  the  mandibular  teeth,  a  char- 
acter readily  distinguishing  animals  of  this  genus  at  any  age  from 
all  other  Microti.  The  upper  incisors  are  without  grooves,  and  the 
proximal  terminations  of  the  roots  of  the  lower  ones  are  situated  on  • 
the  buccal  side  of  the  molars  and  terminate  posterior  to  the  last  one. 
The  tail  is  longer  than  the  hind  foot,  and  in  the  short-tailed  groups 
does  not  taper  to  an  appreciable  extent.  The  palate  is  normal,  with 
or  without  a  median  projection. 

External  characters. — Externally  there  is  little  to  distinguish 
Phenacomys  from  many  other  microtines.  There  is  variation  in 
the  length  of  the  tail,  from  short  to  long,  and  also  in  the  degree  of 
hairiness  of  this  member.  The  ears  are  rather  small  in  most  cases 
and  hidden  to  a  large  extent  by  the  body  hairs.  The  coloration  varies 
through  a  number  of  shades  of  agouti  gray  and  brown,  and  in  one 
case  it  is  strongly  reddish.  In  some  of  the  races  there  is  a  tendency 
toward  increased  grayness  about  the  head;  one  group  has  the  face 
yellow,  and  two  species,  dusky.  The  feet  of  the  terrestrial  species 
are  whiter  than  is  usually  the  case  with  Microtus,  and  the  tail  more 
sharply  bicolor  in  the  shorter-tailed  ones.  Juveniles  are  usually  con- 
siderably darker  and  more  plumbeous  than  adults,  as  is  normally 
the  case  in  this  subfamily.  In  longicamdus  the  coat  is  inclined  to  be 
short  and  coarse,  but  all  other  known  species  of  the  genus  are  covered 
with  hair  that  is  longer  and  more  silky  than  in  the  majority  of 
meadow  mice. 

On  account  of  the  dearth  of  winter  and  earty-spring  material,  it  is 
impossible  to  be  sure  whether  there  are  one  or  two  annual  molts,  but  as 
the  winter  pelage  is  not  only  longer  but  slightly  paler  than  that  of 
early  summer,  it  may  be  assumed  that  there  are  tvro.  At  any  rate,  the 
autumnal  molt  takes  place  at  almost  any  period  late  in  summer  or 
early  in  fall,  the  exact  time  probably  depending  upon  the  age  of  the 
animal  and  local  conditions  of  temperature.  Usually  the  old  pelage 
becomes  somewhat  worn  and  ragged  and  is  gradually  replaced  by 
the  new  coat,  but  variations  are  noted  in  two  instances.  In  the  case 
of  an  old  male  from  the  Bitterroot  Mountains  near  Mullan,  Idaho, 
July  3,  1920,  most  of  the  pelage  is  decidedly  worn  and  dull,  but 
the  new  coat  has  been  assumed  upon  the  underparts  and  has  begun 
to  appear  in  a  clearly  defined  area  upon  the  top  of  the  head  and  the 
nape.  In  an  immature  male  taken  at  Sawtooth  City,  Idaho,  October 
4,  1895,  this  change  is  reversed,  for  the  top  of  the  head,  from  the 


1926]  REVISION    OF   GENUS   PHENACOMYS  7 

nose  to  the  nape,  is  covered  with  short  hairs  of  the  immature,  sum- 
mer coat,  while  the  remainder  of  the  body  has  assumed  the  full, 
grayer,  and  paler  winter  pelage  of  the  adult.  Winter  pelage  is 
naturally  more  luxuriant  than  summer,  and  it  seems  to  be  paler  as 
well,  but  there  is  so  little  late-fall  material  available  that  any  such 
statement  must  be  offered  with  hesitation. 

Where  collectors'  measurements  indicate  any  difference  at  all  in 
size  between  the  sexes,  it  is  invariably  in  favor  of  the  female.  This 
difference  is  considerable  in  some  cases,  but  varies,  and  the  figures 
here  are  usually  offered  without  comment,  so  that  the  reader  may 
form  his  own  conclusions.  Such  difference  seldom  extends  to  the 
crania. 

It  is  sometimes  difficult  to  distinguish  hip  glands  in  the  case  of 
dried  specimens,  or  even  alcoholics,  and  as  no  observations  in  this 
regard  seem  to  have  been  made  in  the  field  upon  fresh  material, 
there  is  little  on  the  subject  that  can  be  stated  with  certainty. 
Phenacomys  longicaudus  apparently  does  not  have  well-developed  hip 
glands  at  any  age,  but  the  males  of  most,  if  not  all,  of  the  short- 
tailed  species  develop  large  glands  when  fully  adult. 

The  full  number  of  mammae,  as  determined  by  various  members 
of  field  parties  of  the  Biological  Survey,  is  evidently  8,  distributed 
2 :  2  inguinal  and  2  :  2  pectoral  in  the  case  of  the  short-tailed  species ; 
but  this  formula  is  often  found  to  vary,  because  of  the  fact  that  the 
full  complement  of  mammae  does  not  always  develop  in  young  fe- 
males with  small  litters.  In  all  specimens  of  P.  longicaudus  examined 
there  are  but  2  :  2  inguinal  mammae  functional.  The  only  female  of 
albipes  that  has  so  far  been  taken  has  2 : 2  inguinal  and  1 : 1  pectoral 
mammae,  but  the  front  legs  are  dried  in  such  position  that  these  may 
possibly  hide  an  additional  pectoral  pair. 

There  are  six  well-developed  tubercles  on  the  soles  of  the  hind 
feet  of  five  spirit  specimens  of  longicaudus,  three  of  ungava,  and 
two  of  olympicus,  but  there  may  be  some  variation  in  this  respect, 
as  in  many  other  microtines. 

JSkull. — The  skull  of  Phenacomys  is  very  typically  Microtus-like  in 
general  form,  and  some  species  of  Microtus  can  be  found  to  match 
almost  every  one  of  the  characters  of  outline.  On  the  whole,  how- 
ever, the  postorbital  processes  of  the  squamosals  are  more  prominent 
than  in  all  but  a  few  species  of  Microtus.  The  palate  is  essentially 
Microtus-like,  and  in  some  forms  (as  celsus)  it  is  very  simple  and 
without  detail;  in  others  (as  the  ungava  group)  it  is  more  complex, 
with  a  median  process  or  spine  projecting  posteriorly. 

Teeth. — It  has  been  asserted  a  number  of  times  (Major,  Hinton, 
AVinge,  et  al.)  that  the  cheek  teeth  of  the  Microtinae  consist  of  the 
fourth  premolar  and  the  first  and  second  molars.  Although  not  yet 
prepared  to  discuss  this  matter  in  full,  the  writer  is  not  at  all  antag- 
onistic to  this  interpretation  of  the  facts,  but  preliminary  investiga- 
tion has  caused  him  to  doubt  the  conclusiveness  of  the  evidence  on 
which  this  assertion  has  been  based.  Therefore,  the  more  conserva- 
tive terminology  (first,  second,  and  third  molars)  will  be  retained  for 
the  present  (fig.  2). 

In  young  individuals  of  Phenacomys  the  molars  are  rootless,  as 
in  Microtus,  and  at  first  seem  to  grow  from  their  matrices  at  a  rate 
slightly  more  rapid  than  that  at  which  they  are  worn  away.    Shortly 


8 


NORTH   AMERICAN   FAUNA 


[No.  48 


before  full  size  has  been  attained,  however,  it  will  be  found  that  the 
dental  pulp  has  begun  to  fail,  and  a  basal  zone,  devoid  of  crenula- 
tions,  is  then  formed  on  each  molar.  As  this  progresses,  the  proximal 
opening  becomes  slightly  constricted,  especially  at  a  point  near  the 
middle,  at  which  spot  a  bridge  of  dentine  is  speedily  formed.  The 
edges  of  the  two  resulting  openings  then  become  elongated  until 
two  fangs  result,  the  basal  openings  of  which,  with  increasing  age, 
become  entirely  closed.  Adults  in  their  early  prime  often  have  an 
internal,  longitudinal  thickening  upon  the  edge  of  the  anterior  root 
of  the  larger  molars,  most  pronounced  in  the  first  upper  one.  This 
condition — not  confined  to  the  genus,  or  even  the  order  Rodentia — 
is  of  great  interest  as  probably  either  indicating  a  tendency  toward 
the  formation  of  a  third  root  by  a  process  of  division,  or,  what  is 
more  probable,  illustrating  the  manner  in 
which  the  third  root  so  often  found  in  other 
families  and  subfamilies  may  have  been  lost 
through  mergence  with  a  neighbor.  An  exami- 
nation of  considerable  material  does  not  indi- 
cate that  this  thickening  of  the  root  increases 
with  advancing  senescence  of  the  individual; 
on  the  contrary,  the  roots  of  very  old  animals 
are  more  attenuate,  and  it  is  believed  that  this 
may  be  caused  by  a  process  of  absorption  of 
the  thickened  root,  thus,  perhaps,  favoring  the 
theory  that  this  thickening  is  a  relic  of  a 
primitive  character. 

The  point  where  the  longitudinal  striations 
or  foldings  of  the  enamel  upon  the  sides  of  the 
teeth  cease  does  not  appear  beyond  the  alveoli 
until  the  animal  is  fully  adult,  and  this  point 
can  usually  be  seen  first  in  the  case  of  the  man- 
dibular teeth.     The  pattern  of  enamel  folding 


Fig.  2. — Left  upper  and 
right  lower  molar  se- 
ries of  Phenacomysi  far 
termetUus  ■mteitnedms 
(No.  67325),  from 
Beartooth  Mountains, 
Mont.     Enlarged 


Fig.  3. — Extreme  change 
in  enamel  pattern  due 
to  age  and  wear. 
Second  upper  molar, 
right-hand  series,  of 
Plwnacom.i/s  maokenz>U 
(No.  110435).  En- 
larged 


is  constant  and  highlv  characteristic  in  a  ge- 


neric sense  and  little  or  no  variation  that  can 
be  correlated  with  specific  differences  occurs; 
but  there  is  usually  much  individual  variation 
within  certain  limits,  some  of  which  is  due  to  advancing  age,  causing 
a  decrease  in  the  acuteness  of  the  angles.  Finally  the  teeth  may 
become  so  worn  that  the  divisions  of  the  roots  are  pushed  well  beyond 
the  alveoli,  the  lateral  grooving  practically  disappears  and  the  basal 
ring  or  zone,  being  partly  exposed  on  the  grinding  surface,  causes  the 
formation  of  small  vacuities  or  pits  completely  surrounded  by  enamel 
(fig.  3).  These  pits  first  appear  upon  the  lingual  side  of  the  mandi- 
bular molars  and  the  buccal  side  of  the  maxillary  ones,  but  their 
origin  is  mechanical  and  they  are  not  of  fundamental  importance. 

There  is  no  appreciable  deposit  of  cementum  within  the  apices  of 
the  reentrant  angles,  but  the  significance  of  this  fact  is  not  apparent. 
In  the  case  of  the  maxillary  teeth,  the  first  molar  has  an  anterior 
crescent  and  two  closed  triangles  on  each  side.  The  second  has  an 
anterior  crescentic  triangle,  and  an  interior  and  two  exterior  triangles, 
both  normally  closed.  The  third  has  an  anterior  crescentic  division, 
an  internal  triangle,  and,  normally,  two  much  smaller  external  ones, 
the  posterior  of  which  varies  considerably  in  form  and  position. 


1926]  REVISION    OF  GENUS  PHENACOMYS  9 

At  times  this  really  constitutes  a  part  of  the  posterior  loop,  and  this 
whole  is  less  constant  than  any  part  of  the  maxillary  series. 

In  the  mandibular  teeth,  the  first  molar  consists  of  an  anterior 
trefoil  of  a  pattern  too  variable  to  be  of  practical  value  in  diagnosis, 
but  part  of  this  variability  is  often  due  to  wear.  There  are  also  three 
internal  and  two  external  triangles  and  a  posterior  crescent.  All 
these  are  normally  closed,  but  there  is  a  decided  tendency  for  them 
to  remain  open,  and  the  postero-external  one  or  two  is  sometimes 
practically  lacking;  or  to  be  more  exact,  these  may  be  integral  with 
the  corresponding  internal  triangles.  The  second  has  three  internal 
and  two  external  triangles.  The  third  usually  consists  of  three 
simple  internal  loops,  highly  characteristic  of  the  genus;  but  there 
is  sometimes  a  very  small  external,  open  triangle  formed  medially. 

In  the  maxillary  series,  the  internal  reentrant  angles  are  only 
slightly  deeper  than  are  the  external;  but  in  the  mandibular  series 
the  difference  is  much  more  pronounced,  to  such  an  extent  that  this 
point  constitutes  one  of  the  most  important  generic  characters. 

The  roots  of  the  mandibular  incisors  of  Phenacomys  extend  pos- 
teriorly between  the  roots  of  the  second  and  third  molars,  terminat- 
ing upon  the  buccal  side  of  the  rami,  but  at  a  level  below  the  alveolar 
borders.  This  condition  is  also  present  in  Evotomys  and,  to  a  lesser 
extent,  in  Ondatra. 

VARIATION 

Mice  of  the  ungava  group  are  probably  rather  uniform  over  wide 
areas,  with  changes  of  character  gradually  accomplished.  In  the 
case  of  the  short-tailed  mountain  forms,  however,  the  colonies  for 
the  most  part  are  well  segregated  upon  detached  mountain  ranges 
or  parts  of  such  ranges,  separated  by  strips  of  low-zone  country; 
and  variation,  especially  in  coloration,  is  comparatively  large  and 
abrupt.  In  fact,  it  must  be  emphasized  that  variation  of  color  is 
so  extreme  in  some  cases,  even  among  animals  taken  in  the  same 
locality,  that  this  character  is  almost  useless  for  differentiation 
between  some  subspecies.  The  skulls  of  animals  from  segregated 
colonies  also  vary  considerably,  even  though  the  areas  may  be 
closely  contiguous,  and  extreme  caution  must  be  used  in  their  assig- 
nation, especially  when  material  is  scanty. 

Broadly  speaking,  the  molar  pattern  within  the  genus  is  uniform, 
but  although  the  width  and  proportions  of  the  molars  are  subspecifi- 
cally  dependable,  the  details  of  their  patterns  can  not  be  considered 
reliable.  Variation  is  excessive  in  certain  particulars,  and  no  solid 
basis  of  relationship  could  be  established  upon  this  character.  It  is 
considered  wiser,  therefore,  not  to  accentuate  the  importance  of 
minute  average  differences  in  the  molar  patterns  over  what  is  felt  to 
be  their  real  relational  worth. 

The  difference  observable  in  the  extent  of  wear  of  the  molar  crowns 
does  not  seem  to  be  invariably  due  to  age,  but  undoubtedly  depends 
also  upon  the  nature  of  the  food  to  which  the  animal  has  been  accus- 
tomed. A  relatively  tough  or  gritty  food  would  cause  more  than 
ordinary  wear,  and  hence  it  is  sometimes  difficult  to  distinguish  by 
this  character  a  comparatively  young  adult  from  one  that  might  be 
very  much  older. 


10  NORTH    AMERICAN    FAUNA  [No.  48 

Immaturity  is  indicated  by  the  rounded  and  weak  appearance  of 
the  skull,  by  the  failure  of  the  portion  of  the  molars  beyond  the 
alveolar  border  to  show  indications  of  roots,  and  by  the  fact  that 
usually,  but  not  always,  the  pelage  of  the  partly  grown  animal  is 
darker  and  more  slaty-colored  than  in  the  adult.  These,  except  for 
the  tooth  characters,  are  the  same  indications  of  immaturity  as  are 
usually  found  in  the  genus  Micrptus.  As  compared  with  adults, 
juvenile  crania  are  weak,  rounded,  and  unridged,  and  they  appear 
to  have  relatively  small  bullae,  short  incisive  foramina,  a  wide  inter- 
orbital  region,  compressed  zygomata,  and  short  rostra.  Racial  dif- 
ferences are  rather  late  developing  in  the  skulls  of  forms  that  are  at 
all  closely  related,  and  for  this  reason  immature  crania  are  of  little 
value  in  diagnoses.  / 

Much  attention  has  been  given  to  obtaining  a  correct  understand- 
ing of  the  extent  of  individual  variation  occurring  among  the 
Phenacomys  from  a  single  locality,  and  of  the  characters  most  con- 
stant in  the  various  races.  The  former  has  been  obtained  by  study- 
ing the  topotype  series  of  mackenzii.  These  consist  of  14  specimens 
that  are  fully  adult,  and  with  the  exception  of  a  single  individual  in 
ragged  pelage,  exhibit  great  uniformity  in  coloration,  but  there  is 

some  variation  in  size,  as  is 
AW>  cJJILL  «J[tlu»       the    case    with   all    micro- 

tines.  It  has  been  found 
that  size  and  shape  of  the 
nasals  and  the  adjoining 
ascending  branches  of  the 
premaxillae,  the  interpari- 

Fig.  4.— Palates  of  Phenacomys,  selected  to  show       etajs     an(J    ^e    palatal    and 
extremes   of   variation  withm  the  genus.      P.   i.  '       .  .  r 

oiympicus    (No.   9037O,   p.   lotigicaudus    (No.      pterygoid   regions  are  un- 

uS4)'  and  P"  maCkenZU  (N°'  110624)"    En"     trustworthy  Characters  on 

which  to  place  depend- 
ence. The  interorbital  width  is  usually  constant  in  the  case  of 
animals  of  comparable  ages,  as  is  the  degree  of  interorbital  ridging, 
but  there  are  occasional  exceptions.  The  most  uniform  and  reliable 
characters  are  size  of  skull,  size  and  shape  of  brain  case,  supraorbital 
development,  interorbital  characters,  rostrum  (taken  as  a  whole), 
incisive  foramina,  bullae,  and  size  of  the  cheek  teeth.  In  addition, 
the  occasionally  exceptional  development  of  some  other  character, 
as  curvature  of  the  upper  incisors  or  the  infero-supefior  depth  of  the 
jugals,  proves  of  value  in  specific  or  group  discrimination.  The  de- 
gree of  development  of  the  median  process  of  the  palate  is  not  at  all 
constant,  but  its  average  is  sometimes  significant.     (See  fig.  4.) 

PROBABLE    AFFINITIES 

The  molar  teeth  of  the  genus  Phenacomys  are  more  completely 
rooted  than  are  those  of  any  other  recent  microtine,  for  although 
this  character  is  shared  by  the  genera  Evotomys  and  Ondatra,  the 
teeth  of  these  last  do  not  become  rooted  until  a  more  advanced  age 
is  attained,  and  the  fangs  never  become  completely  closed.  This 
is  considered  to  be  a  primitive  character,  and  taking  the  teeth  as  a 
criterion  of  the  greatest  importance  in  judging  the  systematic  posi- 
tion of  the  genus,  as  is  usually  done,  Phenacomys  is  indicated  as  be- 
ing the  most  primitive  of  recent  Microtinae. 


1926]  REVISION    OF   GENUS  PHENACOMYS  11 

Such  argument  can  be  accepted  only  tentatively,  however,  for 
there  are  other  factors  to  be  considered  in  determining  the  evolu- 
tional age  of  the  genus.  There  is  strong,  if  not  conclusive  evidence, 
though,  that  the  ancestors  of  the  microtines  had  rooted  teeth.  In 
considering  the  existing  state  of  affairs,  the  possibility  must  be  en- 
tertained of  an  animal,  directly  ancestral  to  Phenacomys,  having  de- 
veloped rootless  teeth  from  an  original  stock  with  rooted  molars, 
and  the  present  genus  developed  from  that.  This  rooted-rootless- 
rooted  theory,  being  a  case  of  the  modification  of  a  part,  not  the 
loss  of  one,  is  a  possibility,  but  the  evidence  so  far  available  does  not 
encourage  its  acceptance. 

Most  of  the  genera  of  the  Microtinae  now  have  molars  growing 
from  a  persistent  pulp  (hypsodont),  and  this  is  believed  (Hinton  in 
Barrett-Hamilton,  1913)  to  be  a  development  brought  about  by  the 
gradual  adoption  of  tougher  and  more  fibrous  foods,  which  would 
increase  the  wear  of  the  crowns  and  stimulate  the  teeth  to  a  longer 
period  of  growth.  An  investigation  of  the  teeth  of  animals  of 
various  ages,  including  embryos,  of  a  number  of  genera  of  the  Crice- 
tidae  (to  be  discussed  at  greater  length  in  a  future  paper)  indi- 
cates that  there  is  no  radical  difference  in  the  development  of  these 
two  types  of  molars,  for  whatever  the  cause,  it  seems  to  be  due 
merely  to  a  stimulation  of  growth  of  the  teeth,  greatly  prolonged  in 
the  case  of  the  microtines,  beyond  the  usual  period.  The  same  stages 
of  tooth  growth,  generally  speaking,  that  occur  in  Phenacomys  may 
be  observed  in  Neotoma,  but  at  a  very  much  earlier  state  of  develop- 
ment in  the  latter,  and  the  growth  of  fangs  seems  to  be  the  natural 
sequence  to  the  atrophy  of  the  dental  pulp. 

Rooted  (brachyodont)  molars  are  more  suitable  for  crushing  soft 
and  brittle  foods,  as  seeds.  It  thus  seems  probable,  but  is  as  yet  un- 
proved, that  the  food  of  Phenacomys  on  the  whole  is  of  a  character 
more  nearly  comparable  with  that  of  the  majority  of  the  Cricetinae 
than  it  is  with  that  of  Microtus. 

No  fossil  remains  of  Phenacomys  have  been  discovered,  nor  any- 
thing that  may  be  considered  as  having  been  truly  ancestral  to  the 
genus;  but  knowledge  of  the  subfamily  indicates  the  likelihood 
that  Phenacomys  is  descended  from  a  pretype  in  which  the  outer 
and  inner  reentrant  angles  of  the  lower  teeth  were  of  practically 
equal  depth,  the  angles  less  sharp,  the  triangles  more  open,  and  the 
enamel  of  the  concave  and  convex  sides  of  the  prisms  of  equal  thick- 
ness. Furthermore,  it  is  very  possible,  and  even  probable,  that  the 
genus  originated  in  North  America.  The  fact  that  it  now  occupies 
a  boreal  habitat,  however,  is  no.  reason  for  assuming  that  it  has 
always  been  confined  to  such  surroundings.  The  present  facts  of 
its  distribution  point  to  the  probability  that  it,  like  the  lemmings,  is 
usually  unable  to  withstand  much  competition,  and  that  it  has  had 
to  give  way  from  time  to  time  before  the  advance  of  more  adaptable 
species  of  meadow  mice.  This  probably  accounts  for  its  presence 
in,  or  restriction  to,  such  situations  as  the  heather  beds  of  the  high 
mountains,  or  the  dense  forests  of  the  Pacific  coast — localities  where 
it  should  encounter  little  competition  ecologically.  Occasionally, 
where  conditions  are  especially  favorable,  and  there  seems  to  be 
"room  for  all,"  this  genus  may  be  found  closely  associated,  and  ap- 
parently on  favorable  terms,  with  both  Microtus  and  Evotomys. 


12  NORTH  AMERICAN  FAUNA  [No.  48 

Summarizing  the  characters  by  which  the  systematic  position  of 
the  genus  may  best  be  judged,  it  is  considered  that  although  the 
rooted  nature  of  the  molars  is  a  character  that  is  often  considered 
primitive,  it  does  not  necessarily  indicate  more  in  this  case  than  that 
the  genus  has  never  experienced  the  necessity  for  faster-growing 
teeth.  The  great  depth  of  the  inner  reentrant  angles  of  the  lower 
teeth  and  the  difference  between  the  thickness  of  the  enamel  on  the 
anterior  and  posterior  faces  of  the  prisms  is  considered  to  indicate  a 
high  degree  of  specialization  for  a  certain  food  or  manner  of  feeding. 
These  details,  coupled  with  distributional  facts,  indicate  that  the 
Phenacomys  type  of  microtine  is  of  decided  antiquity,  and  in  some 
ways  the  genus  Phenacomys  may  be  considered  primitive,  but  the 
enamel  pattern  is  highly  specialised. 

The  majority  of  species  of  Phenacomys  are  terrestrial,  but  two  of 
them  are  arboreal.  The  most  apparent  and  probably  the  most  impor- 
tant difference  between  these  two  groups  is  in  the  length  and  degree 
of  hairiness  of  the  tail.  From  this  it  is  impossible  to  draw  profitable 
conclusions  as  regards  ancestry.  Although  there  are  many  excep- 
tions, nonsaltatorial,  terrestrial  rodents  living  in  a  nonmountainous 
habitat  in  which  there  is  only  low  cover,  usually  have  tails  that  are 
rather  short  for  their  respective  families  or  groups,  while  those  liv- 
ing in  forests,  or  where  the  brush  is  very  heavy,  have  longer  tails. 
It  is  judged  that  this  modification  follows  the  choice  of  habitat.  The 
terrestrial  Phenacomys  albipes,  living  in  a  densely  forested  region, 
is  in  a  number  of  characters  intermediate  between  the  shorter-tailed 
forms  and  the  two  arboreal  species,  and  the  fact  that  these  two  are 
arboreal  is  probably  correlated  with  the  hairiness  of  their  tails.  It  is 
impossible  now,  however,  to  be  sure  whether  albipes  is  derived  from 
a  hairy-tailed,  arboreal  ancestor,  or  from  a  short-tailed,  terrestrial 
one  that  originally  inhabited  low  cover.  It  is  probable,  though  by 
no  means  certain,  that  all  of  the  species  were  derived  from  a  homo- 
geneous ancestral  stock,  but  it  now  seems  impossible  to  speculate  with 
any  degree  of  profit  on  whether  this  ancestor  was  arboreal  or  ter- 
restrial. 

GROUPS 

The  genus  Phenacomys  as  at  present  known  is  naturally  divisible 
into  four  groups,  as  follows : 

(1)  The  intermedins  group,  comprising  the  short-tailed  races 
without  distinctly  yellowish  faces:  to  be  found  in  the  mountainous 
sections  of  the  West. 

(2)  The  'ungava  group,  containing  all  forms  characterized  chiefly 
by  the  distinctly  yellowish  color  of  the  face:  occurring  in  Canada 
east  of  the  Kocky  Mountain  region. 

(3)  The  albipes  group,  containing  a  single  species,  evidently  ter- 
restrial, but  characterized  by  a  long  tail,  which  is  only  slightly 
hairy ;  with  a  sooty  nose,  and  long,  weak  skull :  occurring  along  the 
Pacific  coast  of  northern  California  and  Oregon. 

(4)  The  longicaudns  group,  holding  two  arboreal  species  that  maj' 
be  known  by  lomr,  rather  heavy  tails,  more  hairy  than  those  of  other 
species ;  and  cranially,  by  sharply  decurved  incisors :  found  in  for- 
ests of  the  humid  coast-district  of  northern  California  and  Oregon. 


1926] 


REVISION   OF   GENUS  PHENACOMYS  13 

List  of  Species  and  Subspecies,  with  Their  Type  Localities 


Phenacomys  intermedins  group : 

Phenacomys    intermedins     intermedins 

Merriam Twenty   miles   north-northwest   of 

Kamloops,      British      Columbia, 
Canada. 
intermedius   levis  A.   B.   Howell—   Saint  Marys  Lake,  Teton  County, 

Mont. 

intermedius    olympicus    Elliot Happy  Lake,  Olympic  Mountains, 

Wash. 
intermedius  celsus  A.  B.  Howell Muir    Meadow,    Tuolumne    Mead- 
ows, Yosemite  Park,  Calif. 
Phenacomys  ungara  group : 

Phenacomys  ungava  ungava  Merriam__    Fort  Chimo,  Ungava,  Canada. 

unga-va  crassus   Bangs Rigolet,  Hamilton  Inlet,  Labrador. 

mackenzii    Preble Fort  Smith,  Mackenzie,  Canada. 

Phenacomys  albipes  group : 

Phenacomys  albipes  Merriam Areata,    Humboldt   County,    Calif. 

Phenacomys  longicaudus  group: 

Phenacomys  longicaudus  True Marshfield,   Coos   County,   Oreg. 

silvicola   A.    B.    Howell Five  miles  southeast  of  Tillamook, 

Oreg. 

Key  to  Species  and  Subspecies 

a1  Tail  much  less  than  50  mm. 

b '  Face  not  pronouncedly  yellow ;  rostrum  but  slightly  depressed. 
c1  Skull  large;   condylobasilar  length  averaging  more  than  24 
mm. ;  rostrum  robust. 
d1  Coloration    light,    incisive   foramina    small;    molars    very 

wide P-  i-  celsus  (p.  23). 

d2  Coloration   dark;    incisive  foramina   larger;    molars   very 

narrow P-  i-  olympicus   (p.  21). 

oa  Skull  smaller;  condylobasilar  length  averaging  less  than  24 
mm. ;  rostrum  light. 
d i  Skull  moderate  ;  condylobasilar  length  averaging  more  than 
23   mm. ;    zygomatic    breadth    averaging   more    than   15 

mm P-  i-  intermedius  (p.  15). 

d,2  Skull  small  and  light;  condylobasilar  length  averaging  less 
than   23   mm. ;    zygomatic    breadth    averaging   less   than 

15  mm__     P.  i.  levis  (p.  20). 

b  2  Face  pronouncedly  yellow ;  rostrum  noticeably  depressed. 
c1  Coloration    bright;    size    medium;    incisive    formina    small 

P.  u.  ungava  (p.  25). 
c2  Coloration  duller. 

d 1  Size  large ;  condylobasilar  length  averaging  more  than  24 

mm.;  coloration  darker.     (Labrador) P.  u.  crassus  (p.  27). 

d 2  Size  small ;   condylobasilar  length  averaging  less  than  23 
mm.;  coloration  lighter,  grayer   (central  Canada) 

P.  mackenzii  (p.  28). 
a1  Tail  much  more  than  50  mm. 
b  l  Incisors  strongly  decurved ;  tail  thick,  quite  hairy ;   arboreal. 
c1  Coloration  reddish;  brain  case  relatively  unridged ;  nose  not 

sooty -P.  longicaudus    (p.  32). 

c2  Coloration    brownish;     brain     case     strongly     ridged;     nose 

sooty P.  silvicola  (p.  34). 

b 2  Incisors  not  strongly  decurved  ;  tail  slender,  scantily  haired ; 

terrestrial P.  albipes   (p.  30). 

DESCRIPTIONS  OF  SPECIES  AND  SUBSPECIES 

Phenacomys  intermedius  Group 
(Characters  under  the   species) 

Remarks. — The  P.  intermedius  group,  including  all  of  the  short- 
tailed  forms  from  mountainous  areas  of  the  West,  may  at  once  be  dis- 

93890°— 26 3 


14 


NORTH   AMERICAN   FAUNA 


[Mo.  48 


tinguished  from  the  P.  ungava  group  by  the  lack  in  the  former  of  a 
pronouncedly  yellowish  nose.  The  skulls  of  P.  mackemii  have  a 
narrower  interorbital  measurement  than  have  any  of  the  forms  of 
P.  intermedins,  and  this  region  is  more  heavily  ridged. 

PHENACOMYS    INTERMEDIUS    Meubiam 

Geographic  distribution. — Mountainous  country  of  western  North 
America,  from  northern  British  Columbia  to  the  southern  Sierra 


Fig.    5. — Distribution    of    the   Phenacomys    intermedins    group : 

1.  P.  i.  intermedins.  3.  P.  i.  olympicus. 

2.  P.  i.  leiHs.  4.  P.  i.  eels  us. 
Circle  shows  type  locality  of  subspecies 

Nevada  in  California,  and  the  southern  Rocky  Mountains  in  north- 
ern New  Mexico  (fig.  5). 

General  characters. — This  species  is  distinguished  from  other  short- 
tailed  Phenacomys  by  the  lack  of  a  decidedly  yellowish  nose.  The 
coloration  varies  from  palest  to  darkest  of  the  genus,  and  in  size, 
from  smallest  to  large.  The  skull  varies  accordingly,  but  it  is  rather 
lightly  ridged  interorbitally. 


1926]  REVISION    OF   GENUS  PHENACOMYS  15 

On  the  whole,  the  palates  of  mice  of  this  group  average  simpler, 
with  less  well-defined  median  processes  than  have  the  yellow-nosed 
phenacomys,  but  there  is  much  subspecific  and  individual  variation 
in  this  regard.     Mammae :  Inguinal,  2:2;  pectoral,  2 : 2. 

Size  least  and  skull  lightest  in  the  northeastern  portion  of  its 
range  (P.  i.  lewis),  increasing  in  size  towards  the  west  and  south, 
and  culminating  in  the  large,  dark,  heavy-skulled  Washington  race 
(P.  i.  olynvpicus) .  As  a  rule,  the  animals  living  in  regions  of  slight 
precipitation  are  pale.  There  is  a  marked  increase  in  the  size  of  the 
rostrum  in  the  case  of  the  two  westernmost  races. 

PHENACOMYS  INTERMEDIUS  INTERMEDIUS  Mkbbiam 

Rocky  Mountain  Phenacomys 

(PI.  2,  figs.  1,  2,  and  6) 

Phenacomys  intermedins  Merriam,  North  Amer.  Fauna  No.  2,  p.  32,  October  30, 
1889. 

Phenacomys  orophilus  Merriam,  North  Amer.  Fauna  No.  5,  p.  65,  July  30, 
1891.  Type  collected  at  10,500  feet  altitude,  east  slope  of  Lemhi  Mountains, 
Idaho,  August  28,  1890,  by  Merriam  and  Bailey ;  female  immature ;  No.  Iff  If 
U.  S.  Nat.  Mus.  (Biol.  Surv.  coll.)  ;  original  number,  1710. 

Phenacomys  truei  Allen,  Bui.  Amer.  Mus.  Nat.  Hist.,  6:  331,  November  7, 
1894.  Type  collected  in  Black  Hills  (=Laramie  Mountains),  Wyo.,  August  10, 
1857,  by  W.  A.  Hammond ;  immature ;  No.  VWVV? ;  U.  S.  Nat.  Mus. ;  original 
number,  109. 

Phenacomys  oramontis  Rhoads,  Amer.  Nat.,  29 :  941,  October,  1895.  Type  col- 
lected on  Mount  Baker  Range,  British  Columbia,  at  6,000  feet  altitude 
(=Lumsden  (sometimes  spelled  Lihumitson)  Mountain,  now  called  Church 
Mountain,  just  north  of  the  International  Boundary  near  longitude  121°  50' 
west),  August  6,  1895,  by  A.  C.  Brooks;  male  adult;  No.  9354,  Acad.  Nat. 
Sci.  Philadelphia  (formerly  No.  2354,  S.  N.  Rhoads  coll.)  ;  original  number, 
428. 

Phenacomys  preolei  Merriam,  Proc.  Biol.  Soc.  Washington,  11 :  45,  March  16, 
1897.  Type  collected  at  about  9,000  feet  altitude,  Longs  Peak,  Colo.,  August 
12,  1895,  by  E.  A.  Preble;  male  immature;  No.  74513,  U.  S.  Nat.  Mus.  (Biol. 
Surv.  coll.)  ;  original  number,   647. 

Phenacomys  eonstaolei  Allen,  Bui.  Amer.  Mus.  Nat.  Hist.,  12:  4,  March  4,  1899. 
Type  collected  at  Telegraph  Creek,  British  Columbia,  August  25,  1898 
[=1897],  by  A.  J.  Stone;  female  adult;  No.  14406,  Amer,  Mus.  Nat.  Hist. 

Type. — Collected  on  basaltic  plateau  about  20  miles  north-north- 
west of  Kamloops,  British  Columbia,  altitude  5,500  feet,  October  2, 
1888,  by  George  M.  Dawson;  immature;  No.  780,  Museum  of  the 
Geological  and  Natural  History  Survey  of  Canada. 

Geographic  distribution. — British  Columbia  west  of  the  eastern 
divide  of  the  Rocky  Mountains,  northeastern  Washington,  Idaho, 
eastern  and  southern  Oregon,  and  northern  California ;  thence  in  the 
mountains  through  southern  Montana,  Wyoming,  Colorado,  and  into 
northern  New  Mexico    (fig.  5,  1). 

Habitat. — Canadian  and  Hudsonian  Zones,  mostly  in  open,  grassy 
parks. 

General  characters. — A  phenacomys  normally  of  a  rather  pale 
coloration  and  the  most  "  medium  "  form  of  this  group,  with  no 
characters,  either  external  or  internal,  accentuated. 

Color. — The  range  of  coloration,  even  in  the  same  locality,  is 
often  great,  and  varies  from  tawny  olive  to  the  Saccardo  umber  of 
Ridgway,  with  various  blendings  of  the  two.    The  plumbeous  bases 


16  NORTH   AMERICAN   FAUNA  [No.  48 

of  the  hairs  show  through  to  a  greater  extent  in  the  case  of  the 
darker  individuals.  The  underparts  are  whitish,  occasionally  with 
a  tinge  of  buffy.  Nose  not  pronouncedly  tinged  with  yellow.  This 
is  the  only  species  of  the  genus  of  which  thoroughly  satisfactory 
specimens  in  undoubted  winter  pelage  are  available.  This,  in  a 
December  specimen  from  Colorado  and  two  October  skins  from 
Idaho,  is  much  grayer  (between  drab  and  hair  brown  of  Ridgway) 
and  with  little  of  the  brownish  cast  to  be  found  in  summer  skins. 
During  cold  weather  the  feet  are  whiter,  because  of  the  longer  hairs, 
and  the  tail  is  then  uniformly  whitish,  instead  of  bicolor.  Young 
animals  are  usually  darker  than  adults  from  the  same  place,  and  are 
more  plumbeous,  both  above  and  below. 

Skull. — The  superior  aspect  of  the  skull  is  essentially  medium 
in  the  development  of  all  characters — rostrum,  interorbital,  and 
brain  case.  Inferiorly,  the  incisive  foramina  and  bullae  are  pro- 
portionately rather  small,  and  the  palatal  projection  or  spine  is 
sometimes  present,  but  as  often  practically  absent.  There  is  varia- 
tion in  skull  characters,  even  in  the  case  of  animals  from  the  same 
locality,  but  not  to  such  an  extent  in  animals  considered  to  be  typical 
of  the  race  as  to  cause  overlapping  with  surrounding  forms  herein 
recognized. 

Teeth. — -Maxillary  teeth  of  a  pattern  normal  to  the  genus,  with 
posterior  loop  of  third  molar  tending  to  vary  in  antero-posterior 
length.  The  anterior  trefoil  of  the  first  lower  molar  varies  from 
an  excessive  development  of  the  reentrant  angles,  causing  an  almost 
complete  tripartite  division,  to  a  condition  in  which  the  trefoil  has 
no  anterior  loop  whatever,  and  the  exterior  one  is  but  slightly 
indicated.  The  third  lower  molar  usually  has  a  complete  external 
triangle.  Occasionally  there  is  a  tendency  for  the  triangles,  espe- 
cially the  external  ones  of  the  first  lower  molar,  to  remain  open. 

Measurments. — Collectors'  measurements  of  adults:  Male  from 
Hazelton,  British  Columbia:  Total  length,  141;  tail,  27;  hind  foot, 
18.5.  Female  from  Nelson,  British  Columbia:  142;  37;  18.  Skin 
from  Okanagan,  British  Columbia:  142;  37;  17.  Five  males,  Bear- 
tooth  Mountains,  Mont.:  145  (142-147);  30  (26-34);  17  (17-18). 
Two  females,  Beartooth  Mountains,  Mont.:  152;  34;  18.  Skull : 
Average  of  six  adults  from  Beartooth  Mountains,  Mont. :  Condy- 
lobasilar  length,  23.8  (23.5-24.1)  ;  nasals,  8  (7.6-8.4)  ;  interorbital 
breadth,  3.6  (3.5-3.7)  ;  zygomatic  breadth,  15.2  (14.9-15.7) ;  lamb- 
doidal  width,  11.4  (11.2-11.5)  ;  incisive  foramina,  4.7  (4.6-4.9)  ; 
maxillary  tooth  row,  6  (5.7-6.2) ;  height,  9.3  (8.9-9.7). 

Remarks. — The  typical  subspecies  intemiedms  can  best  be  de- 
scribed by  the  word  composite,  for  the  animals  comprising  it  are, 
in  large  part,  from  isolated  mountain  colonies,  most  of  which  vary 
in  some  slight  particular  from  what  is  considered  to  be  typical. 
To  one  not  familiar  with  the  material,  it  will  doubtless  seem  re- 
markable that  the  range  of  this  form  is  so  extended.  It  can  not 
be  denied  that  when  far  more  specimens  are  available  than  are 
now  at  hand  it  may  possibly  prove  practicable  to  recognize  one  or 
more  additional  forms  within  this  area.  At  present,  however,  this 
can  not  be  done  without  naming  a  number  of  new  and  very  poorly 
defined  races. 


1926]  REVISION    OF    GENUS  PHENACOMYS  17 

It  is  desired  to  emphasize  strongly  the  belief  that  with  the  present 
material  these  slight  variations  can  not  be  assigned  a  geographical 
foundation,  and  animals  from  the  same  spot  often  vary  to  a  greater 
extent  from  one  another  than  from  still  others  taken  hundreds  of 
miles  away.  Coloration,  except  within  rather  wide  limits,  is  un- 
reliable, and  there  seems  often  to  be  a  sort  of  dichromatism.  Un- 
fortunately, there  are  no  perfect  adult  skulls  from  southern  British 
Columbia,  the  majority  of  specimens  having  the  occipital  region 
broken.  Measurements  of  a  series  of  skulls  from  southern  Montana 
are  given,  however,  so  as  to  demonstrate  the  great  extent  of  varia- 
tion occurring  at  one  locality.  For  the  most  part,  the  material 
is  from  scattered  areas,  and  although  some  of  the  variations  may 
later  prove  to  be  constant,  the  specimens  at  hand  indicate  that 
they  are  not.  There  seems  to  be  no  other  solution  but  to  recognize 
intermedins  as  a  composite  race,  with  many  local  variations  too 
slight  and  inconstant  to  be  worthy  of  names,  but  yet  a  form  suf- 
ficiently homologous  to  be  readily  distinguishable  from  other  con- 
tiguous races  now  recognized.  For  the  foregoing  reasons,  it  may  be 
desirable  to  discuss  these  variations  at  some  length. 

The  type  of  this  form  and  of  the  genus'  is  an  immature  individual 
of  which  the  unstuffed  (but  not  "flat ")  skin  is  pressed  between  two 
pieces  of  glass,  and  the  skull  is  badly  smashed,  with  only  the  anterior 
portion  remaining.  The  characters  assigned  to  the  species  when  it 
was  described  consisted  of  the  considerable  breadth  of  the  inter- 
orbital  (3.8  mm.),  the  expanded  terminations  of  the  ascending 
branches  of  the  premaxillae,  and  the  pattern  of  the  anterior  lower 
molar.  None  of  these  points  are  distinctive,  for  there  are  other 
crania  from  widely  separated  localities,  both  adult  and  immature, 
with  interorbital  measurement  greater  than  this.  There  is  slight 
chance  that  the  width  of  the  premaxillary  branches  is  significant  in 
this  case,  for  broader  ones  are  found  in  topotypes  of  mackenzii  and 
others,  and  there  is  great  variation  in  this  respect  within  almost 
every  series  of  phenacomys.  Maximum  width  of  these  branches 
usually  occurs  in  immature  skulls,  but  sometimes  in  adults  also,  their 
breadth  being  in  inverse  ratio  to  the  width  of  the  posterior  portion 
of  the  nasals,  and  this  last  is  often  variable.  The  molar  pattern  of 
the  type  of  intermedins  was  supposed  to  be  distinctive  because  the 
anterior- trefoil  of  the  first  lower  molar  is  so  deeply  cut  by  reentrant 
angles  that  an  outer  triangle  is  wholly  isolated  and  an  inner  one 
nearly  cut  off.  It  can  not  be  denied  that  in  the  type  this  is  carried 
to  a  degree  slightly  more  pronounced  than  is  to  be  found  in  any 
other  specimen  at  hand  from  British  Columbia  but  this  character 
has  been  found  to  be  so  extremely  variable  that  it  is  worthless  for 
diagnosis  in  other  cases ;  so  it  is  inferred  that  it  should  be  considered 
the  same  in  this.  Finally,  the  skin,  as  well  as  can  be  told  through  the 
glass,  appears  indistinguishable  from  a  number  of  other  skins  taken 
both  to  the  north  and  to  the  south  of  it,  even  though  the  pelage 
may  well  be  a  trifle  less  dense  than  is  usually  the  case. 

Although  there  are  no  topotypes  of  intermedins,  there  is  every 
reason  for  assuming  that  these  would  not  prove  different  from  in- 
dividuals from  surrounding  areas,  and  it  is  felt  that  a  fairly 
good  understanding  of  the  characters  of  the  form  has  been  obtained 


18  NORTH   AMERICAN   FAUNA  *\9- 


■48 


from  a  study  of  the  material  available  from  contiguous  localities, 
even  though  few  of  the  skulls  are  undamaged. 

It  is  unfortunate  that  neither  the  type  nor  topotypes  (except 
two  fragmentary  skulls  of  juveniles)  of  "  eonstableV  are  available 
at  this  time.  Edward  A.  Preble  4  examined  this  material  several 
years  ago  and  came  to  the  conclusion  that  not  only  are  the  specimens 
clearly  referable  to  the  intermedins  rather  than  to  the  ungava  group, 
as  stated  by  the  describer,  but  that  there  are  no  skull  characters  dis- 
tinguishing this  so-called  species  from  intermedins  or  "  orophilus" 
and  the  only  difference  existing  is  its  slightly  darker  color  in  com- 
parison with  "  orophilus."  As  the  type  of  the  latter  is  lighter  than 
most  specimens  of  intermedins,  no  reliance  is  placed  on  this  dis- 
similarity, and  it  is  deemed  better  to  place  the  name  in  synonymy 
for  the  present.  Incidentally  Preble  advises  the  writer  that  the  date 
of  collection  should  have  been  stated  as  1897  instead  of  1898. 

Specimens  from  the  Mount  Baker  Range  of  British  Columbia,  de- 
scribed by  Rhoads  under  the  name  "  oramontis"  are  indistinguish- 
able in  color  from  animals  of  the  Cascade  Mountains  of  Washing- 
ton, but  the  short  skulls  with  weak  rostra  certainly  more  nearly  ap- 
proach intermedins.  After  these  comparisons  were  made,  a  single 
specimen  from  the  Washington  portion  of  the  Mount  Baker  Range 
was  identified  as  olympicns.  Subsequent  material  from  the  same 
region  showing  variable  characters,  however,  indicates  that  although 
selected  specimens  are  fairly  intermediate  in  character  between 
olympicns  and  intermedins,  the  wiser  course  is  to  allocate  them  all 
to  the  latter  race.  It  should  be  mentioned  that  the  type  of  "  ora- 
montis  "  has  a  tail  generously  tipped  with  white,  but  so  have  several 
Washington  individuals. 

There  is  a  small  series  of  skins  available  from  southern  British 
Columbia,  but  unfortunately  the  skulls  are  damaged,  though  not  to 
an  extent  to  make  them  useless  for  study;  and  most  of  them  are 
without  measurements.  Two  of  these  skins  are  a  shade  darker  than 
any  of  this  race  from  the  United  States,  but  four  other  specimens 
from  the  same  locality  are  almost  at  the  opposite  limit  of  bright- 
ness. 

An  old  male  from  near  Mullan,  Idaho,  has  an  unusually  narrow, 
weak  skull  with  constricted  brain  case,  which  may  be  interpreted 
as  showing  a  tendency  toward  levis.  The  skin  is  notable -because 
of  the  excessively  worn  condition  of  the  pelage  and,  therefore,  its 
superficial  darkening. 

Merriam  described  "  orophilus  "  from  the  Salmon  River  Moun- 
tains of  Idaho,  but  on  current  maps  this  name  is  restricted  to  the 
western  part  of  the  range,  the  portion  from  which  the  tj^pe  came 
now  being  known  as  the  Lemhi  Mountains.  As  far  as  can  be  told 
from  the  subadult  type,  the  skull  has  no  peculiar  characters  what- 
ever. The  skin  is  rather  light  in  color,  but  can  be  matched  by 
animals  from  several  other  distant  localities.  In  comparing  speci- 
mens with  the  type  of  " orophilus"  there  have  often  been  used  two 
immatures  from  Sawtooth  City,  which  is  virtually  in  a  part  of  the 
Salmon  River  Mountains.     These  are  paler  than  the  type,  but  are 

4  Preble,  Edward  Alexander.  Discussion  of  species  of  Phenacomys  occurring  in 
British  Columbia.      [Unpublished  manuscript.] 


192C1 


REVISION    OF   GENUS   PHENACOMYS  19 


hardly  comparable,  as  they  are  in  winter  pelage.  After  a  careful 
consideration  of  the  question  involved,  it  is  felt  that  this  name  must 
be  placed  in  synonymy. 

As  one  becomes  more  familiar  with  the  fauna  of  the  Crater  Lake 
region  of  southern  Oregon,  he  is  increasingly  impressed  with  the 
fact  that  it  has  a  well-marked  relationship  with  that  of  the  Great 
Basin  area;  but,  nevertheless,  it  was  somewhat  surprising  to  find 
that  the  Phenacomys  not  only  of  this  locality  but  also  of  northern 
California  must  be  called  intermedins.  It  is  certain  that  specimens 
can  be  placed  neither  with  the  form  directly  to  the  northward,  nor 
the  one  to  the  south.  In  coloration  they  are  entirely  similar  to 
intermedins,  and  in  cranial  characters  as  well,  except  for  a  very 
slight  increase  in  the  size  of  the  bullae. 

The  skull  of  the  only  adult  (barely  so)  from  the  Wallowa-Blue 
Mountain  country  shows  a  greater  departure  from  what  is  con- 
sidered typical  intermedins  than  does  any  other  individual  of  this 
race  examined.  This  is  shown  by  the  rather  broad  and  stubby 
rostrum,  and  by  the  peculiar,  though  not  greatly  developed,  ridging 
of  the  broad  interorbital  and  the  very  wide  sulcus  between.  With 
but  the  single  specimen  it  is  impossible  to  judge  whether  this  is  a 
character  constant  throughout  the  region  or  merely  fortuitous.  It 
is  certainly  closer  to  intermedins  than  to  anything  else  now  recog- 
nized, and  is  so  placed. 

Specimens  from  southern  Montana  have  a  slight  accentuation  of 
the  interorbital  ridging,  and  some  of  the  skulls  have  rostra  that  are 
a  little  more  robust,  and  bullae  a  shade  larger  than  usual,  but  there  is 
so  much  variation  in  this  character,  even  at  the  same  locality,  that 
it  is  impossible  to  give  it  much  significance. 

Allen  described  "truei"  (1894)  from  a  damaged  immature  sup- 
posed to  have  come  from  the  Black  Hills  of  South  Dakota,  in  which 
case  the  specimen  might  have  been  expected  to  belong  to  a  new 
race.  It  was  later  ascertained  that  the  true  locality  was  the  Black 
Hills,  now  known  as  Laramie  Mountains,  Wyo. 

The  type  of  "  prehleV  Merriam  is  a  very  buffy  example,  immature, 
with  the  ascending  branches  of  the  maxillae  very  wide.  The  latter 
character  is  found  to  be  without  significance,  as  mentioned  elsewhere, 
and  the  skin  can  be  practically  matched  by  another  from  Okanagan, 
British  Columbia.  It  is  very  closely  approached,  also,  by  two  others 
from  the  same  locality  and  by  New  Mexico  individuals  as  well; 
hence,  it  can  hardly  be  considered  a  valid  race. 

Specimens  examined. — Total  number,  85,  as  follows : 

British.  Columbia:  Chapa-atan  River,  mountains  near  the  head  of,  1; 
Glacier,  1 5 ;  Hazelton,  3 6 ;  Kamloops,  20  miles  north  northwest  of,  1  T 
(type)  ;  Mount  Baker  Range,  3  s  (including  type  of  "  oramontis  ")  ;  Nel- 
son, 1;  Okanagan,  68;  Rossland,  l9;  Telegraph  Creek,  2°  (broken  skulls 
only). 

California:  Fort  Crook,  1;  Mount  Shasta,  3. 

Colorado:  Buffalo  Pass,  Jackson  County,  l10;  Fairplay  (skull  only),  1; 
Lake  Moraine,  El  Paso  County,  l10;  Longs  Peak,  1  (type  of  "  preblei")  ; 
Nederland,  1  u. 

B  Amer.  Mus.  Nat.  Hist. 

0  Mus.   Vert.  Zoot. 

7  Victoria  Mem.  Mus. 

3  Mus.  Comp.  Zool.,  2  ;  Acad.  Nat.  Sci.  Philadelphia,  1. 

*  Provincial  Mus. 

10  E.  R.  Warren  coll. 

11  Acad.  Nat.  Sci.  Philadelphia. 


20  NORTH  AMERICAN  FAUNA  [No.  48 

Idaho:  Cabinet  Mountains,  3;  Moscow,  8  miles  northeast  of,  1;  Mullan, 
east  of,  l12;  Salmon  River  Mountains  (=Lemhi  Mountains),  4  (including 
type  of  "  orophiliis")  ;   Sawtooth  City,  2. 

Montana:  Beartooth  Mountains,  14;  Big  Snowy  Mountains,  1. 

New  Mexico:  Pecos  Baldy,  1;  Santa  Fe,  1;  Taos  Mountains,  1;  Twining,  1. 

Oregon:  Blue  Mountains,  1;  Crater  Lake,  2;  Diamond  Lake,  1;  Wallowa 
Lake,   4. 

"Washington :  Loomis,  1 12 ;  Round  Top  Mountain,  Pend  Oreille  County,  1  w ; 
Mount  Baker,  3 13. 

Wyoming:  Beartooth  Lake,  9;  Laramie  Mountains  (=Black  Hills),  3  (in- 
cluding type  of  "truei")  ;  Merna,  1;  Yellowstone  Park,  1. 

PHENACOMYS  INTERMEDIUS  LEVIS  A.  B.   Howell 
Alberta   Phenacomys 

•        (P1./2,  fig.  3) 

Phenacomys   intermedins    levis   A.    B.    Howell,    Proc.    Biol.    Soc.    Washington, 
36:  157,  1923. 

Type. — Collected  at  Saint  Marys  Lake,  Teton  County,  Mont., 
May  23,  1895,  by  Arthur  H.  Howell;  male  adult;  No.  72405,  U.  S. 
Nat.  Mus.  (Biol.  Surv.  coll.)  ;  original  number,  13. 

Geographic  distribution. — Upon  the  eastern  slope  of  the  Rocky 
Mountains  at  least  from  central  Alberta  south  to  Teton  County, 
Mont.  (fig.  5,0). 

Habitat. — Canadian  and  Hudsonian  Zones  in  rather  open  grassy  or 
mossy  situations. 

General  characters. — A  subspecies  differing  but  slightly  in  color 
from  many  specimens  of  intermedins,  but  skull  weak  and  smaller 
than  any  other  form.    Bullae  and  rostrum  relatively  large. 

Color. — Topotypes  of  this  form  have  the  basal  tone  close  to  the 
clrab  of  Ridgway,  with  the  tips  of  the  dorsal  hairs  brown,  the  exact 
shade  differing  apparently  with  age — duller  and  darker  in  the  case 
of  old  animals,  and  brighter  and  lighter  in  the  younger  ones  (large 
immatures).  The  underparts  are  grayish,  occasionally  with  a  faint 
tinge  of  buffy.  The  feet  are  pure  white,  except  when  darkened  by 
grease,  and  the  tail  is  indistinctly  bicolor.  Young  animals  are  darker 
and  more  plumbeous. 

/Skull. — Weak  and  smallest  of  the  genus,  with  little  ridging  and 
variable  interorbital  width;  but  this  last  is  sometimes  considerable. 
Rostrum  and  bullae  relatively  large,  and  incisive  foramina  usually 
short.     Palatal  spine  slightly  developed. 

Dentition. — Normal  for  the  genus,  but  in  the  maxillary  teeth  the 
second  external  triangle  of  the  third  molar,  although  variable,  tends 
to  disappear  to  a  greater  extent  than  usual.  Of  the  mandibular  teeth, 
the  trefoil  of  the  first  molar  is  variable  and  there  is  always  an 
external  triangle  to  the  third  molar,  although  this  is  not  invariably 
closed.  With  this  exception,  there  seems  to  be  no  tendency  for  the 
triangles  of  the  teeth  to  remain  open. 

Measurements. — Collectors'  measurements  of  adult  specimens  from 
the  type  locality  are  as  follows :  Of  two  males :  Total  length,  135 
tail,  34;  hind  foot,  17.  Of  6  females:  139  (132-146);  33  (30-36) 
18  (17-18).  Skulls  of  two  adult  male  topotypes  average  as  follows 
Condylobasilar  length,  22.3 ;  nasals,  7.1 ;  interorbital  breadth,  3.6 
zygomatic  breadth,  14.1;  lambdoidal  width,  11.3;  incisive  foramina, 

UD.  R.  Dickey  coll.  13  Wash.  State  College. 


1026]  REVISION    OF   GENUS   PHENACOMYS  21 

4.3;  maxillary  tooth  row,  5.8;  height,  8.7.  Of  five  adult  females: 
Condylobasilar  length,  22.6;  nasals,  7.3;  interorbital  breadth,  3.6; 
zygomatic  breadth.  13.9;  lambdoidal  width,  10.9;  incisive  foramina, 
4.5;  maxillary  tooth  row,  5.8;  height,  8.3. 

Remarks. — One  of  the  unexpected  results  of  the  present  study 
was  the  discovery  of  this  new  race  apparently  confined  to  the  vicin- 
ity of  the  eastern  slope  of  the  easternmost  range  of  the  Rocky  Moun- 
tains in  Alberta  and  northern  Montana.  Although  it  differs  in  some 
respects  from  most  skins  of  intermedins,  the  coloration  of  the  latter 
is  subject  to  such  a  range  of  variation  that  slight  reliance  can  be 
placed  thereon.  On  the  whole,  topotypes  of  levis  are  almost  indis- 
tinguishable in  coloration  from  the  type  of  " orophUus"  and  the 
series  of  skins  from  the  Beartooth  Mountains  of  Montana,  except 
that  the  present  form  has  less  overwash  of  dark-tipped  hairs.  The 
skull,  however,  presents  differences  that  necessitate  the  separation 
of  these  animals  from  any  race  heretofore  known.  Compared  with 
intermedins  it  has  a  skull  much  smaller,  with  a  rostrum  that,  al- 
though relatively  slightly  shorter,  seems  to  be  a  trifle  more  robust. 
The  bullae  are  also  relatively  larger  than  those  of  the  southern 
neighbor.    In  most  but  not  all  cases  the  incisive  foramina  are  smaller. 

The  difference  in  the  coloration  of  the  nose  is  sufficient  to  distin- 
guish levis  from  mackemii  at  a  glance ;  the  latter  has  a  larger  skull 
with  proportionately  narrower  interorbital,  heavily  ridged. 

The  animals  from  central  Alberta,  included  here,  are  undeniably 
somewhat  darker  than  typical,  approaching  in  this  respect  the  dark- 
est intermedins,  but  their  skulls  can  not  be  so  assigned.  Although 
some  of  them  exhibit' slight  variation,  as  in  the  shape  of  the  rostrum, 
this  is  not  directly  towards  intermedins,  and  they  clearly  belong 
with  levis.  With  one  exception  the  skulls  of  the  southwestern  Al- 
berta specimens  are  either  missing  or  badly  damaged ;  hence  identi- 
fication is  made  largely  on  geographical  grounds. 

Specimens  examined. — Total  number,  43  as  follows: 
Alberta:  Banff,  214;  Braggs  Creek,  l15;  Crows  Nest  Pass,  l18;  Fiddle  Creek, 
1  M  ;  Fishing  Lake,  1 ;  Head  of  Smoky  River,  S 17 ;  Jasper  House,  1 15 ;  Moose 
Mountain   (=Indian  Head),  21S;  Mount  Forgetmenot,  1 1G  ( ?,  no  skull); 
Waterton  Lakes  Park,  5 15. 
British  Columbia :  Moose  River,  3. 

Montana:   Kintla  Lake,  1;  Many  Glaciers,  1 ;  Midvale,  1 ;  Saint  Marys  Lake, 
13  ls  (including  type)  ;  Summit,  1. 

,     PHENACOMYS  INTERMEDIUS  OLYMPICUS  Elliot 
Olympic  Phenacomys 
(PL   2,   fig.   4) 

Phenacomys  olympicus  Elliott,   Field   Columb.   Mus.,   Pub.   30,   Zool.    Ser.,   1 : 

225,  February,  1S99. 
Hicrotus   {Lagurus)   pumilus  Elliot,   Field  Columb.  Mus.,  Pub.  30.  Zool.   Ser., 

1 :   226,   February,   1899.     Type  from  Happy   Lake,   Clallam  County,   Wash. 

Male  juvenile ;   No.   6269,    Field   Columb.    Mus. ;    collected   bv   D.    G.    Elliot, 

September  2,  1898. 

Type. — Collected  at  Happy  Lake  at  5,000  feet  altitude,  Olympic 
Mountains,  Wash.,  August  14,  1898,  by  D.  G.  Elliot;  male  adult; 
No.-V78/,  Field  Columbian  Museum. 


"  Mus.  Comp.  Zool.  «  U.  S.  Nat.  Mus.,  7  ;  Provincial  Mus.,  1. 

"Victoria  Mem.  Mus.  lS  U.  S.  Nat.  Mus.,  12;  Amer.  Mus.  Nat.  Hist,  1. 

16  Acad.  Nat.   Sci.   Philadelphia. 

93890°—  26 4 


22  NORTH   AMERICAN   FAUNA  [No.  48 

Geographic  distribution. — In  the  Hudsonian  Zone  of  the  Olympic 
and  Cascade  Mountains  of  Washington,  and  as  far  south  as  central 
Oregon    (fig.  5,  3). 

Habitat. — Upper  Canadian  and  Hudsonian  Zones  about  logs  and 
brush  at  the  eclge  of  timber  and  among  the  grass  and  heather  of  the 
higher  parks. 

General  characters. — The  darkest  species  of  short-tailed  phena- 
comys,  with  skull  large,  heavy  and  well  angled,  and  rostrum  heavy. 
No  difference  in  size  between  the  sexes  is  indicated. 

Color. — It  is  odd  that  of  nine  topotypes  taken  subsequent  to  1920, 
only  one  individual  is  as  light  in  color  as  all  of  those,  numbering 
eight,  taken  prior  to  1900.  Part  of  this,  but  not  all,  may  be  due 
to  age,  the  majority  of  the  paler  ones  being  subadult,  whereas  the 
others,  slightly  darker  than  the  drab  of  Ridgway,  are  mostly  old 
adults  with  short  pelage.  These  latter,  which  are  assumed  to  be 
most  typical  of  the  race,  have  very  little  brown  in  the  pelage,  while 
the  lighter  animals  have  much  brown  and  are  slightly  paler  than 
the  darkest  intermedins.  All  are  clear  gray  below — rather  dark — 
with  white  feet,  and  tails  bicolored  in  varying  degree. 

Skull. — Large  and  heavy,  with  robust  rostrum  and  quite  promi- 
nent ridging,  but  brain  case  not  large.  Supraorbital  processes 
prominent ;  incisive  foramina  broad  but  short,  and  bullae  moderately 
large.  The  supraoccipital  is  more  nearly  at  right  angles  to  the 
axis  of  the  skull  (less  sloping)  than  is  usual  in  the  genus.  Palatal 
spine  present  or  absent. 

Dentition. — The  molars  of  this  race  are  vecy  small  and  narrow. 
The  only  character  of  the  enamel  pattern  worthy  of  note  is  the 
trefoil  of  the  anterior  lower  molar,  which  is  proportionately  larger, 
and  usually  more  fully  expanded,  than  in  any  other  race. 

Measurements. — Collectors'  measurements  of  six  adult  topotypes 
are :  Total  length,  154 ;  tail,  43 ;  hind  foot,  20.  Skull  measurements 
of  five  adult  males  from  the  Olympic  Mountains  are :  Condylo- 
basilar  length,  24.5;  nasals,  7.6;  interorbital  breadth,  3.8;  zj^gomatic 
breadth,  15.4;  lambdoidal  width,  11.7;  incisive  foramina,  4.7; 
maxillary  tooth  row,  6.3 ;  height,  9.6. 

Remarks. — On  the  whole  olympicus  is  the  darkest  and  dullest 
form  of  the  genus,  but  there  is  considerable  variation  in  this  regard, 
as  previously  mentioned,  and  it  may  be  well  nigh  impossible  to 
identify  the  lighter  individuals  by  color  alone.  Skulls  of  this  race 
may  be  told  almost  at  a  glance  from  those  of  intermedins  by  the 
large  size  and  robust  rostrum,  broad  interorbital  usually  well  ridged 
in  adults,  and  less  sloping  supraoccipital.  Inferiorly  the  incisive 
foramina  are  broader  but  not  longer;  the  bullae  are  larger  and  the 
molars  narrower. 

Skins  and  skulls  from  Mount  Rainier  are  practically  indistin- 
guishable from  topotypes  except  for  narrower  incisive  foramina  and 
molars  a  shade  wider. 

Of  two  skins  from  Mount  Adams,  one  is  exceedingly  rusty,  and 
differs  from  the  other  by  very  much  the  same  degree  that  the  type 
of  "  prebleV  does  from  darker  examples  of  intermedins. 

Grayest  and  darkest  of  all  the  specimens  is  a  female  from  Three 
Sisters,  Oreg.,  and  its  skull  is  also  much  closer  to  olympicus  than 
to  any  other.    Another  adult  female  from  the  same  locality,  how- 


1926]  REVISION   OF  GENUS  PHENACOMYS  23 

ever,  is  not  only  closer  in  color  to  intermedins,  but  its  skull  is  con- 
siderably weaker  and  smaller  than  any  specimen  of  that  form,  and 
hence  not  at  all  like  olympicns.  As  there  are  no  other  adults  from 
the  Cascades  of  Oregon  north  of  the  Crater  Lake  region,  and  as  the 
writer  dislikes  exceedingly  to  assign  two  specimens  from  the  same 
locality  to  different  subspecies,  it  is  considered  better  to  class  both 
of  these  individuals  as  olympicns  for  the  present,  although  it  is 
freely  admitted  that  other  disposition  could  be  made  of  at  least  one 
of  them  with  equal  propriety. 

"  Microtus  (Lagurus)  pumilus"  described  by  Elliot  (1899a), 
proved  to  be  a  juvenile  Phenacomys. 

Specimens  examined. — Total  number,  60,  as  follows: 

Oregon:  Deschutes  River,  l19^  Mount  Hood,  1;  Three  Sisters,  420. 

Washington:  Boulder  Lake,  l21;  Buck  Creek,  1;  Cat  Creek,  Olympic  Moun- 
tains, 10 " ;  Dosewallips,  1 ;  Happy  Lake,  Olympic  Mountains,  7 a 
(including  the  type,  and  the  type  of  "pumUus")',  Canyon  Creek, 
Olympic  Mountains,  1 23 ;  Mount  Adams,  2 ;  Mount  Rainier,  21 2* ; 
Olympic  Mountains,  9 ;  Snoqualmie  Pass,  1 25. 

PHENACOMYS  INTERMEDIUS  CELSUS  A.  B.  Howell 

Sierran  Phenacomys 

(PI.  2,  fig.  5) 

Phenacomys  intermedins  celsus  A.   B.   Howell,   Proc.   Biol.   Soc.   Washington, 
36:15S,  1923. 

Type. — Collected  in  Muir  Meadow  at  9,300  feet  altitude,  Tuolumne 
Meadows,  Yosemite  National  Park,  Calif.,  August  13,  1901,  by  J.  H. 
Gaut;  male  adult;  No.  109103,  U.  S.  Nat.  Mus.  (Biol.  Surv.  coll.); 
original  number,  250. 

Geographic  distribution. — The  Sierra  Nevada  of  California  from 
the  Lake  Tahoe  region  south  probably  as  far  as  Tulare  County 

(fig.  M). 

Habitat. — Chiefly  among  low  cover  such  as  grassy  meadows  and 
patches  of  heather  in  high  Hudsonian  Zone. 

General  characters. — A  very  pale  form,  slightly  smaller  than 
Olympic  us,  with  large  skull  but  faintly  ridged  and  with  a  robust 
rostrum.  Incisive  foramina  very  short  and  narrow,  and  molars 
very  stout.     Sexes  of  equal  size. 

Color. — Rather  close  to  the  wood  brown  of  Ridgway  but  some^ 
what  variable.  In  typical  skins  there  are  relatively  few  dark- 
tipped  hairs  and  there  is  no  tendency  towards  an  accentuation  of 
grayness  anteriorly.  Underparts  whitish,  ordinarily  with  a  strong 
tinge  of  buffy.    Tail  bicolored  in  varying  degree. 

Skull. — The  skull  of  celsus  is  large  with  well-expanded  brain  case, 
and  the  rostrum  robust,  though  not  long.  Temporal  ridges  poorly 
developed.  Incisive  foramina  very  small,  and  molariform  teeth 
very  broad  and  heavy.    Palatal  spine  practically  absent. 

19  Game  Dept.  Oregon. 

20  U.  S.  Nat.  Mus.,  3  ;  Game  Dept.  Oregon,  1. 

21  Field  Mus.  Nat.  Hist. 

22  U.  S.  Nat.  Mus.,  9  ;  Wash.  State  College,  1. 

23  Wash.    State   College. 

34  U.  S.  Nat.  Mus.,  IS  ;  Wash.  State  College,  3. 

35  D.  R.  Dickey  coll. 


24 


NORTH   AMEEICAN   FAUNA 


[No.  48 


Dentition. — The  enamel  pattern  is  perfectly  normal  except  for  its 
great  width,  and  the  trefoil  of  the  first  lower  molar  is  even  more 
variable  than  usual. 

Measurements. — Average  of  five  adults  from  the  Yosemite  Park: 
Total  length,  148 ;  tail,  39 ;  hind  foot,  18.  Skulk :  Average  measure- 
ments of  three  adults  from  this  region :  Condylobasilar  length,  24.1 ; 
nasals,  7.8;  interorbital  breadth,  4;  zygomatic  breadth,  15.8;  lamb- 
doidal  width,  11.9 ;  incisive  foramina,  4.3 ;  maxillary  tooth  row,  6.2 ; 
height,  9.4. 

Remarks. — The  relationship  of  celsus  is  apparently  with  olympi- 
cus^ and  it  is  undoubtedly  nearer  that  race  at  the  present  day  than 
to  intermedins.  As  definite  infusion  of  Great  Basin-Rocky  Moun- 
tain tendencies,  however,  is  exhibited  by  the  examples  from  northern 
California  and  southern  Oregon,  and  the  ranges  of  the  Washington 
and  central  California  races  are  thereby  separated,  the  similarity  of 


Fig.  6. — Distribution  of  the  Phenacomys  ungava  group: 

1.  P.  mackenzii. 

2.  P.  u.  ungava. 

3.  P.  u.  crussus. 

Typo    locality    of   each   form    shown    by    circle. 

characters  of  oh/mpicus  and  celsus  may  be  largely  fortuitious.  It  is 
therefore  probable  that  such  similarity  is  the  result  of  evolutional 
stimuli  that  happened  to  be  the  same  in  the  two  areas  in  question. 
Although  celsus,  however,  is  readily  distinguishable  and  intergrada- 
tion  can  not  occur  with  other  races  because  of  low-zone  barriers  to 
the  north,  the  trinomial  had  better  be  employed  for  it. 

Typical  examples  of  celsus  are  evidently  somewhat  smaller  and 
much  paler  than  olympicus,'  in  fact,  in  coloration  they  are  compar- 
able to  the  palest  intermedins,  but  with  less  buffiness  than  is  usually 
exhibited  by  the  latter.  The  skulls  need  comparison  only  with  olym- 
picus, than  which  they  have  a  larger  brain  case,  stouter  though  no 
longer  rostrum,  and  very  little  indication  of  interorbital  ridging. 
The  incisive  foramina  are  considerably  smaller ;  indeed,  they  are 
relatively  smaller  than  in  any  other  race,  and  the  molars  are  much 
heavier.  It  is  worthy  of  note  that,  proportional  to  size,  olympicus 
has  the  lightest,  narrowest  molars  of  the  genus,  and  celsus  the  widest. 
The  latter  has  bullae  a  shade  the  larger. 


1926]  •    REVISION"    OF   GENUS   PHENACOMYS  25 

Two  adult  skins  with  good  skulls  from  the  vicinage  of  Lake  Tahoe, 
Calif.,  are  a  trifle  darker,  and  the  rostra  are  definitely  longer  and  not 
quite  so  robust  as  in  typical  examples.  On  these  characters  they  are 
separable  from  Yosemite  specimens,  but  it  certainly  is  not  advisable 
to  differentiate  the  animals  of  these  regions  subspecifically  on  the 
basis  of  the  present  material. 

Specimens  examined. — Total  number,  12,  as  follows: 

California:  Echo,  32S;  Fletcher  Creek,  Yosemite  Park,  l27;  Lyell  Canyon, 
1 27 ;  Mono  Pass,  1 ;  Mount  Lyell  2 ;  Muir  Meadow,  Tuolumne  Meadows,  2 
(including  the  type)  ;  Pyramid  Peak,  l28;  Ten  Lakes,  Yosemite  Park,  l2t. 

Phenacomys  ungava   Group 

Geographic  distribution. — Nonmountainous  portions  of  the  Cana- 
dian and  Hudsonian  Zones  of  Canada.     (See  map,  fig.  6.) 

General  characters. — A  group  the  members  of  which  are  chiefly 
characterized  by  a  distinctly  yellowish  nasal  area;  skull  well  ridged 
and  palate  normally  with  a  well-defined,  rather  blunt,  median  proc- 
ess, but  with  some  variation  in  the  degree  of  development.  Rostrum 
markedly  depressed.  Size  greatest  and  color  darkest  in  Labrador; 
smallest  and  palest  in  Alberta. 

PHENACOMYS  UNGAVA  Mekriam 

Geographic  distribution. — The  Canadian  and  Hudsonian  Zones  of 
Labrador,  Quebec,  and  Ontario  at  least  as  far  west  as  the  north 
shore  of  Lake  Superior  (fig.  6,  2  and  3). 

General  character's. — A  species  whose  external  characters  are  not 
well  understood,  on  account  of  the  poor  quality  of  the  available 
material  representing  the  typical  race.  A  medium  to  large  phena- 
comys of  bright  to  dark  coloration,  with  no  decided  tendency  toward 
an  increase  of  grayness  anteriorly.  Skull  medium  to  large,  with 
rather  large  bullae.    Larger  and  evidently  darker  eastward. 

PHENACOMYS  UNGAVA  UNGAVA  Mekriam 

Ungava  Phenacomys 

(PI.  3,  fig.  1) 

Phenacomys  ungava  Merriam,  North  Amer.   Fauna  No.  2,  p.  35,   October  30, 

1889. 
Phenacomys  celatus  Merriam,   North  Amer.   Fauna   No.  2,  p.  33,   October  30, 

1889.     Type  from  Godbout,  Quebec,  Canada.     Male  adult ;  No.  186486,  U.  S. 

Nat.  Mus.  (formerly  No.   fffi,  Merriam  coll.)  ;  collected  by  N.  A.  Comeau, 

June  10,  1886. 
Phenacomys  latimanus  Merriam,  North  Amer.  Fauna  No.  2,  p.  34,  October  30, 

1889.     Type    from    Fort    Chimo,    Ungava,    Canada.     Male    immature;    No. 

186487,  U.  S.  Nat.  Mus.  (No.  fiff  Merriam  coll)  ;  collected  by  L.  M.  Turner. 

Type. — Collected  at  Fort  Chimo,  Ungava,  Canada,  by  L.  M. 
Turner;  male  old  adult;  No.  186488,  U.  S.  Nat.  Mus.  (No.  ^-ff 
Merriam  coll.). 

Geographic  distribution. — Probably  suitable  places  throughout  the 
whole  of  Quebec,  including  Ungava,  and  at  least  as  far  west  as 
central  Ontario  (fig.  6,  2). 

M  Mus.   Comp.   Zool. 

27  Mus.  Vert.  Zool. 

28  Field  Mus.  Nat.  Hist. 


26  NORTH   AMERICAN   FAUNA  [No.  48 

General  characters. — A  medium-sized,  short-tailed,  yellow-nosed 
species  of  bright  coloration :  incisive  foramina  small. 

Color. — The  type  specimen  of  this  form  is  of  a  bright  chestnut 
brown,  close  to  the  Prout  brown  of  Ridgway  but  a  shade  brighter 
on  the  dorsal  surface,  with  the  underparts  pale  buffy  gray,  all  hairs 
plumbeous  at  base ;  tail  indistinctly  bicolor,  and  feet  pale  buff.  Face 
somewhat  paler  and  pronouncedly  yellowish,  this  being  brightest  on 
the  nose. 

Skull. — Of  medium  size,  with  rather  short  rostrum,  small  incisive 
foramina  and  with  relatively  large  brain  case,  broad  posteriorly. 

Dentition. — The  molar  pattern  of  the  specimens  examined  exhibits 
no  peculiarities.  The  anterior  trefoil  of  the  first  lower  molar  is 
quite  variable,  as  is  usual  in  this7 genus. 

Measurements. — Of  the  following  three  alcoholics  before  making 
over  into  study  skins,  as  given  by  Miller:  Type  of  ungava:  Total 
length,  138;  tail,  31;  hind  foot,  19.  Type  of  "  latimanus  " :  116;  28; 
18.  Type  of  "celatus":  130;  32;  17.5.  Subadult  male  from  On- 
tario (in  flesh)  :  138;  33;  19.  Skull  (type)  :  Condylobasilar  length, 
23.6;  nasals,  7.4;  interorbital  breadth,  3.7;  zygomatic  breadth,  15; 
lambdoidal  width,  11.7;  incisive  foramina,  4.6;  maxillary  tooth  row, 
6 ;  height,  9.2.  Type  of  "  celatus  " :  Condylobasilar  length,  23.5 ; 
nasals,  7.2 ;  interorbital  breadth,  3.6 ;  zygomatic  breadth,  14.5 ;  lamb- 
doidal width,  11.4;  incisive  foramina,  4.7;  maxillary  tooth  row,  5.6; 
height,  9. 

Remarks. — The  material  from  the  range  assigned  to  this  species, 
consisting  only  of  the  types  of  ungava,  "  latimanus"  "  celatus"  and 
two  other  immature  examples  from  Godbout,  is  too  scanty  to  admit  of 
the  final  working  out  of  its  relationship.  All  five  of  these  specimens 
were  originally  alcoholics,  so  neither  the  coloration  nor  the  external 
proportions  are  reliable.  The  three  above  types  are  entirely  indis- 
tinguishable from  one  another  in  color,  and  are  of  an  unusually 
bright  shade  of  brown  such  as  is  found  in  no  other  race  of  this 
genus,  but  this  condition  may  well  be  due  to  the  action  of  the  alcohol. 
The  only  external  differences  are  that  of  size,  undoubtedly  due  partly 
to  age  and  partly  to  shrinking  by  the  preservative;  and  the  notice- 
able broadness  of  the  forefeet  of  "  latimanus"  but  this  last  seems  to 
be  occasioned  by  drying  in  an  extended  position.  The  types  of  both 
ungava  and  "  latimanus''''  are  from  Fort  Chimo.  The  former  is  an 
old  individual  and  the  latter  an  immature;  therefore  the  skull  of 
"  latimanus  "  is  the  smaller,  especially  in  the  rostral  region,  and  with- 
out interorbital  ridging,  but  these  differences  are  such  as  are  nor- 
mally to  be  found  in  animals  of  corresponding  ages,  and  it  is  con- 
sidered extremely  unlikely  that  they  can  be  referable  to  different 
races. 

The  skull  of  the  type  of  ';  celatus  "  is  of  about  the  same  age  as  that 
of  ungava,  but  a  trifle  smaller  and  slightly  more  ridged;  the  differ- 
ence is  slight  and  well  within  the  range  of  individual  variation,  as 
exhibited  in  series  of  other  forms  of  the  genus  from  a  single  locality. 
An  immature  from  Godbout,  originally  alcoholic,  was  "  made  up  " 
very  small,  but  even  taking  into  consideration  the  consequent  crowd- 
ing of  the  dorsal  hairs,  it  is  distinctly  darker. 

The  immature  male  from  Ontario  is  paler  and  grayer  than  Que- 
bec examples,  and  the  tail  is  more  sharply  bicolor.    Until  adult  skulls 


1926]  BEVISION   OF   GENUS  PHENACOMYS  27 

are  available,  or  topotypes  of  ungava  have  been  taken,  the  signifi- 
cance of  this  difference  can  not  be  interpreted! 

Phenacomys  "  latimanus  "  is  now  placed  in  synonymy  rather  than 
y/ngava  because  the  former  is  not  adult  and  is  a  much  less  desirable 
specimen  to  be  the  type  of  a  species.  For  the  use  of  the  name  ungava 
in  preference  to  that  of  "  celatus"  see  Miller  (1897  b,  p.  77). 

Specimens  examined. — Total  number,  7,  as  follows : 

Ontario:  Frantz,  l29;  Peninsula  (north  shore  of  Lake  Superior),  Is0. 
Quebec:  Fort  Chimo,  2  (type  of  ungava  and  type  of  "latimanus")  ;  God- 
bout,  3   (including  type  of  "  celatus"). 

PHENACOMYS  UNGAVA  CRASSUS  Bangs 

Labrador  Phenacomys 

(PI.  3,  fig.  2) 

Phenacomys  celatus  crassus  Bangs,  Proc.  New  Eng.  Zool.  Club,  2 :  39,  1900. 

Type. — Collected  at  Rigolet,  Hamilton  Inlet,  Labrador,  August 
15,  1895,  by  C.  H.  Goldthwaite;  male  old  adult;  No.  3959,  E.  A.  and 
O.  Bangs  coll.  (now  in  Mus.  Comp.  Zool.). 

Geographic  distribution. — Southern  Labrador:  Limits  of  range 
unknown  (fig.  6,  3). 

General  characters. — A  large,  rather  dull-colored  race  with  long, 
well-ridged  skull,  heavy  rostrum,  and  large  incisive  foramina. 

Color. — Above  brownish,  close  to  the  snuff  brown  of  Riclgway  but 
a  trifle  darker,  with  a  rather  circumscribed  nasal  area  pronouncedly 
yellowish.  Underparts  grayish  without  any  tinge  of  buffiness  and 
tail  quite  sharply  bicolor.  The  young  are  darker  and  less  brownish 
than  the  normal  adult,  with  the  plumbeous  hair  bases  showing  more. 

Skull. — Large  and  long  with  heavy  rostrum;  interorbital  ridging 
and  supraorbital  processes  well  developed.  Incisive  foramina  and 
bullae  large. 

Dentition.— The  enamel  pattern  of  the  molars  is  normal  with  only 
slight  variation  in  the  form  of  the  trefoil  of  the  anterior  lower 
molar.  In  one  or  two  examples  of  topotypes  there  is  a  tendency 
for  the  triangles  of  the  lower  teeth  to  remain  open. 

Measurements. — Three  adult  males  from  Labrador:  Total  length, 
153;  tail,  38;  hind  foot,  20.  Three  adult  females  from  Labrador: 
151 ;  36 ;  16.  Skull :  Average  measurements  of  the  three  best  adults 
from  Labrador :  Condylobasilar  length,  24.4 ;  nasals,  7.6 ;  interorbital 
breadth,  3.5;  zygomatic  breadth,  15.2;  lambdoidal  width,  11.4;  in- 
cisive foramina,  5.1;  maxillary  tooth  row,  5.8;  height,  9.3. 

Remarks. — A  well-marked  race  but  unlikely  to  prove  to  be  specifi- 
cally distince  from  ungava.  From  the  latter  it  differs  in  being  more 
soberly  colored  dorsally,  grayer  (less  buffy)  beneath,  tail  more 
sharply  bicolor,  darker  above,  and  in  having  the  yellow  area  of  the 
face  somewhat  more  restricted.  Longer  feet  also  seem  to  be  indi- 
cated, but  this  difference  may  be  illusory  because  of  all  typical 
specimens  of  ungava  available  having  been  in  spirits.  The  same 
cause  may  be  partially,  though  probably  not  wholly,  responsible  for 
the  fact  that  these  Labrador  skins  have  more  prominent  ears  with 

30  W.  E.  Saunders  coll.  so  Mus.  Comp.  Zool. 


28  NORTH   AMERICAN   FAUNA  [No.  48 

a  lighter  growth  of  hair.  The  skulls  of  crassus  are  larger  and 
longer  than  those  of  ungava,  with  rostrum  proportionately  longer 
and  much  more  robust.  The  jugals  are  deeper  supero-inferiorly  and 
the  postorbital  borders  slightly  more  square.  There  is  some  little 
difference  in  the  interparietals,  but  they  vary,  and  this  part  of  the 
skull  is  usually  of  little  value  in  diagnosis,  as  it  is  extremely  variable 
in  most  forms.  The  incisive  foramina  are  longer  and  wider,  espe- 
cially anteriorly,  in  the  case  of  the  Labrador  animals,  and  the  bullae 
distinctly  larger. 

Of  the  two  skulls  from  Grosswater  Bay  (on  Hamilton  Inlet),  one 
is  badly  broken  and  the  other  is  little  better.  What  is  left  shows  it 
to  be  a  very  old  male  of  a  remarkable  size,  and  far  larger  than  any 
other  specimen  of  this  race.  / 

Specimens  examined. — Total  number,  16,  as  follows: 

Labrador:  Grosswater  Bay,  2  (skulls  only)  ;  Hamilton  Inlet,  13 81;  L'Anse 
au  Loup,  1 32. 

PHENACOMYS  MACKENZII  Preble 

Mackenzie  Phenacomys 

(PI.  2,  fig.  1;  pi.  3,  fig.  3) 

Phenacomys  mackeiizii  Preble,  Proc.  Biol.  Soc.  Washington,  15 :  182,  1902. 

Type. — Collected  at  Fort  Smith  (near  Athabaska-Mackenzie 
boundary  line),  Slave  River,  Canada,  June  29,  1901,  by  E.  A.  and 
A.  E.  Preble;  male  old  adult;  No.  110625,  U.  S.  Nat.  Mus.  (Biol. 
Surv.  coll.)  ;  original  number,  4271. 

Geographic  distribution. — The  territory  east  of  the  mountains  in 
Alberta,  north  almost  to  Great  Bear  Lake,  and  east  to  Hudson  Bay 

(fig.  M). 

Habitat. — Canadian  and  Hudsonian  Zones;  in  strips  of  forest 
along  the  bottom  lands,  and  grassy  glades. 

General  characters. — A  rather  small  phenacomys  with  short  tail 
and  small  feet  compared  with  other  members  of  the  ungava  group. 
Coloration  brightest  on  rump  and  palest  anteriorly;  face  yellowish; 
rostrum  markedly  depressed. 

Color. — Rather  pale,  the  coloration  of  the  rump  being  close  to  the 
snuff  brown  of  Ridgway,  but  paler  and  grayer  anteriorly.  Under- 
pays grayish  white,  superficially  darkened  by  the  plumbeous  bases 
of  the  hairs  showing  through,  but  without  buffmess.  Facial  area 
yellowish  and  feet  pale;  bicoloration  of  the  tail  fairly  well  defined. 
Juveniles  are  slightly  darker  and  duller. 

Skull. — Rather  small,  narrow  interorbitally  and  with  slender  ros- 
trum. Bullae  small  and  molars  light.  Rostrum  more  decidedly  de- 
pressed than  in  any  other  species. 

Dentition. — There  is  no  peculiarity  of  molar  pattern  in  this  race. 
The  trefoil  of  the  anterior  lower  molar  is  variable,  as  is  usual  in  the 
genus  (fig.  7). 

Measurements. — Collectors'  figures  for  adult  topotypes  are  as  fol- 
lows :  Eleven  males :  Total  length,  138  (133-144)  ;  tail,  32  (29-34)  ; 

si  Mus.  Comp.  Zool.,  12;  U.  S.  Nat.  Mus.,  1.  a*  Mus.  Comp.  Zool. 


1926]  EEVISION   OF   GENUS   PHENACOMYS  29 

hind  foot,  17  (17-18).  Three  females:  143;  32;  17.  Skulls  of  11 
adult  topotypes  average  as  follows:  Condylobasilar  length,  22.7; 
nasals,  7.4;  interorbital  breadth,  3.1;  zygomatic  breadth,  14;  lamb- 
doidal  width,  11.1;  incisive  foramina,  4.5;  maxillary  tooth  row,  5.7; 
height,  9.1. 

Remarks. — The  relationship  of  the  present  form  lies  with  ungava, 
and  it  is  thought  that  the  difference  between  them  will  finally  prove 
to  be  only  subspecific  in  degree.  This  as  yet,  however,  can  not  be 
demonstrated,  and  so  it  seems  better  to  retain  the  binomial  for  the 
present. 

As  previously  mentioned,  no  confidence  can  be  placed  in  the  exter- 
nal characters  of  ungava  as  embodied  in  existing  skins,  but  in  com- 
parison with  them,  topotypes  of  mackenzii  are  very  much  duller 
and  considerably  paler,  especially  anteriorly,  with  underparts  clear 
gray  instead  of  being  tinged  with  buffiness.  In  coloration,  skins  of 
mackenzii  are  much  more  comparable  to  crassus,  from  which  they 
differ  only  slightly,  being  a  trifle  paler,  grayer  on  the  head,  tail 
darker  and  face  slightty  less  yellowish.  They  are  also  smaller  in  all 
dimensions.  The  skulls  of  mackenzii,  however,  differ  from  ungava 
in  a  direction  opposite  to  that  shown  by  crassus.  The  first  mentioned 
may  be  distinguished  from  the 
others  by  the  degree  to  which  the 
rostrum  is  depressed. 

Measurements  of  available  skins 
indicate  that  the   females   are   ap- 
preciably larger  than  the  males,  but        tfiG.  7.— First  lower  molars,   right- 
this  disparity  in  size  does  not  extend  £"£■  **£?*  °f  FnheTcom£, J5££ 

.        -i       ,-  J      rri.  .    .    .  ..  kenzn,  selected  to  snow   extremes 

to  the   Skulls.       Ihe  material   IS   ade-  of  variation  In  the  anterior  trefoil 

quate,  however,  to  demonstrate  that  tte  t&&***TrL?Ul°*™  °f 

the    interorbital    ridging    is    much 

more  pronounced  in  the  case  of  males — to  such  an  extent  that  the  sex 
of  an  adult  may  usually  be  told  from  the  skulls  alone.  The  single 
exception  to  this  is  the  specimen  from  Fort  McMurray,  marked  male ; 
but  as  it  is  adult  and  totally  without  ridging,  the  correctness  of  sex 
determination  is  doubtful. 

The  adult  from  Fort  Churchill,  a  female,  has  slightly  larger  bullae 
but  is  otherwise  normal. 

A  subadult  female  from  Muskeg  Creek,  west-central  Alberta,  is 
very  aberrant  in  coloration,  being  a  bright  brown  without  the  usual 
tinge  of  grayish.  Its  yellowish  nose  places  it  in  the  present  group, 
but  the  damaged  skull,  too  young  to  exhibit  pronounced  characters, 
cuts  short  further  speculation. 

The  specimen  from  Red  Deer  River  is  considerably  paler  than  any 
other  at  hand,  but  its  skull  is  smashed  to  bits. 

Specimens  examined. — Total  number,  42,  as  follows : 

Alberta:  Athabaska  Landing,  1;  Athabaska  River,  1;  Fort  McMurray,  1"; 

Muskeg  Creek,  1 ;  Pierre  au  Calumet,  1 ;  Red  Deer  River,  1 ** ;  Slave  River 

(10  miles  below  Peace  River),  1. 
Keewatin:  Fort  Churchill,  3. 
Mackenzie:  Fort  Resolution,  3;  Fort  Smith,  25   (including  type);   Lake 

Saint  Croix,  1. 
Saskatchewan:  Crackingstone  Point,  Athabaska  Lake,  3M. 

83  Mus.  Comp.  Zool. 

M  Field  Mus.  Nat.  Hist. 

86  Mus.  Comp.  Zool.,  2 ;  U.  S.  Nat.  Mus..  1. 

93890°— 26 ^5 


30 


NORTH   AMERICAN   FAUNA 


[No.  48 


Phenacomys  albipes   Group 

(Characters  under  the  species) 

PHENACOMYS  ALBIPES  Mekriam 

Coast  Phenacomys 

(PI.  3,  fig.  4) 

Phenacomys  albipes  Merriam,  Proc.  Biol.  Soc.  Washington,  14 :  125,  1901. 

Type. — Collected  at  Areata,  Humboldt  County,  Calif.,  May  24, 
1899,  by  W.  K.  Fisher;  male  adult;  No.  97236,  U.  S.  Nat.  Mus.  (Biol. 

Surv.  coll.)  ;  original 
number,  821. 

Geographical  dis- 
tribution.— Occurs  in 
a  coastal  strip  of  un- 
known width  from 
Areata,  Humboldt 
County,  Calif.,  north 
to  the  vicinity  of 
the  Columbia  River 
probably,  and  east  as 
far  as  Vida,  Oreg. 
(fig.  8). 

Habitat. — Usually 
the  vicinity  of  little 
streams  in  the  humid 
coast  forests.  Evi- 
dently terrestrial. 

General  charac- 
ters.— A  phenacomys 
of  a  dark,  rich  brown 
with  long  tail  rather 
scantily  haired. 
Skull  relatively  long, 
slender,  and  rounded. 
Color. — The  dorsal 
coloration  of  this 
species  is  of  a  darker, 
richer  brown  (close 
to  the  Prout  brown 
of  Ridgway)  than 
any  other  member 
of  the  genus,  and 
there  is  a  plentiful  admixture  of  black-tipped  hairs.  The  pelage  is 
everywhere  plumbeous  at  base,  and  the  ventral  surface  is  clear  gray 
in  spring  skins,  but  those  taken  in  fall  have  this  part  strongly 
washed  with  pinkish  buff.  The  tail  is  distinctly  bicolor  and  the 
face  sooty.  In  some  specimens  the  ear  tips  are  practically  naked 
and  in  others  they  are  covered  with  fine  black  hairs.  The  feet 
are  rather  light,  their  exact  shade  depending,  as  usual,  upon  the 
quantity  of  grease  present  in  the  dried  skin,  but  they  are  no  paler 
than  in  most  other  species,  if  as  pale. 


Fig.  8. — Distribution  of  Phenacomys  albipes. 
in  circle 


Type  locality 


1926]  REVISION    OF   GENUS  PHENACOMYS  31 

Skull. — The  cranium  of  this  species  is  strongly  characterized  by 
its  relative  slendemess,  especially  anteriorly.  It  is  high  and  full 
posteriorly;  there  is  no  elevation  of  the  interorbital  region,  and  the 
zygomatic  expansion  is  slight.  The  skull  is  perhaps  as  distinctive 
as  that  of  any  member  of  the  genus. 

Dentition. — The  incisors  of  albipes  are  only  moderately  curved — 
about  as  in  the  short-tailed  races — and  the  molariform  teeth  are 
somewhat  light  and  narrow,  though  less  so  than  in  longicaudus. 
The  anterior  trefoil  of  the  first  lower  molar  is  even  more  variable 
than  usual,  and  there  is  often  considerable  and  unusual  distortion  of 
the  first  outer  triangle  of  the  second  lower  molar,  this  then  being 
compressed  and  twisted,  and  remaining  open.  The  third  lower  molar 
is  usually  simple,  with  distinct  crenulation  upon  the  buccal  side,  but 
in  two  instances  there  is  a  small  outer  triangle  indicated,  and  there 
is  a  distinct  tendency  for  the  anterior  loop  to  be  reduced  in  size. 

Measurements. — Average  of  six  adult  males:  Total  length,  171 
(165-181)  ;  tail,  63  (62-71)  ;  hind  foot,  19.5  (19-20).  Subadult  fe- 
male: 159;  67;  21.  Skull:  Average  of  five  adult  males  (two  imper- 
fect) :  Condylobasilar  length,  23.2 ;  nasals,  7.6 ;  interorbital  breadth, 
3.6;  zygomatic  breadth,  13.9;  lambdoidal  width,  11.4;  incisive  foram- 
ina, 4.8 ;  maxillary  tooth  row,  6.2 ;  height,  9.3. 

Remarks. — This  species  can  be  confused  with  no  other  member  of 
the  genus,  but  it  bears  a  striking  superficial  resemblance  to  Microtus 
mordax  angusticeps.  The  length  of  its  tail  at  once  distinguishes  it 
from  other  terrestrial  phenacomys,  and  the  lesser  diameter  and  un- 
pronounced  hairiness  of  this  member,  from  the  two  arboreal  forms. 
It  is  totally  different  in  color  from  longicaudus,  and  although  the 
coloration  is  close  to  that  of  silvicola,  the  only  specimens  of  the  lat- 
ter have  much  longer  pelage  of  a  different  quality. 

The  brain  case  of  the  males  is  strikingly  full  and  rounded,  espe- 
cially the  posterior  portion,  which  is  well  elevated.  The  frontal 
region  is  noticeably  more  depressed  than  in  the  arboreal,  or  indeed 
any  other,  species.  The  rostrum  is  comparatively  long  and  slender, 
and  the  zygomatic  processes  of  the  maxillae  are  weaker  and  more 
sloping  than  in  other  forms.  The  incisive  foramina  are  slightly 
longer  and  narrower  posteriorly  than  in  either  arboreal  species,  and 
the  palatal  pits  are  a  trifle  smaller  in  their  antero-posterior  dimension. 
The  molariform  teeth  are  very  narrow,  but  the  tooth  rows  are  long. 

In  the  present  state  of  knowledge,  the  relationship  of  albipes  is  in 
some  doubt.  Whether  or  not  the  degree  of  curvature  of  the  incisors 
of  the  two  arboreal  species  is  a  character  recently  brought  about  by 
food  habits,  or  whether  it  is  of  deeper  significance,  it  is,  at  any  rate, 
an  excellent  group  character,  and  one  that  is  not  shared  by  albipes. 
Both  the  latter  and  silvicola,  however,  have  pronouncedly  sooty 
faces,  and  this  may  or  may  not  be  of  more  than  simple  specific  signi- 
ficance. No  positive  assertions  can  be  made,  but  it  is  felt  that  the 
points  of  external  resemblance  of  albipes  to  the  longicaudus  group 
are  not  so  profound  as  might  at  first  be  imagined. 

Specimens  examined. — Total  number,  8,  as  follows: 

California:  Areata,  1  (the  type)  ;  Orick,  1 35. 
Oregon:  Blaine,  2s6;  Netarts,  337;  Vida,  1  * 

SB  D.  R.  Dickey  coll.  OT  S.  G.  Jewett  coll. 

36  D.  R.  Dickey  coll.,  1;  A.  Walker  coll.,  1.  ss  Game  Dept.  Oregon. 


32 


NORTH    AMERICAN    FAUNA 


[No.  48 


Phenacomys  longicaudus   Group 

Geographic  distribution. — Forests  of  the  humid  coast  district  from 
Mendocino  County ,  Calif .,  north  probably  to  the  Columbia  River  (fig.  9 ) . 

General  charac- 
ters.— Species  chiefly 
arboreal  in  habitat 
and  characterized  by 
long,  stout  tails, 
black  above,  and 
relatively  heavily 
haired,  and  long 
toes  fitted  for  climb- 
ing. Upper  incisors 
more  sharply  de- 
curved  than  in  other 
groups. 

PHENACOMYS  LON- 
GICAUDUS   True 

Red  Tree  Mouse 

[PI.  1   (frontispiece)  ; 
pi.  3,  fig.  5] 

Phenacomys  longicau- 
dus True,  Proc.  U.  S. 
Nat.  Mus.,  13 :  303, 
1890. 

Type.  —  Collected 
at  Marshfield,  Coos 
County,  Oreg.,  Au- 
gust, 1890; 
ture ; 
U.  S.  Nat.  Mus 

Geographic  distri- 
bution.— Locally  in 
humid  coast  district 
from  Mendocino 
County,  Calif.,  into 
central  Oregon,  and 
possibly  to  the  Co- 
lumbia Kiver  (fig. 
9,1).  _ 

Habitat. — Largely 
arboreal  and  found 
practically  always  in 
coniferous  trees  (ex- 
cept redwoods). 

General  charac- 
ters.— A  bright  red- 
dish species  with  long,  blackish,  rather  hairy  tail  and  long  toes. 
Skull  short  and  thickset  with  post-orbital  processes  well  developed. 
Incisors  sharply  decurved. 

Color. — Above,  of  a  uniform  cinnamon,  close  to  the  ochraceous 
tawny  of  Ridgway,  many  of  the  hairs  sparingly  tipped  with  black; 


lmma- 
TVr»  '      19Q71 

^  "•         ^5  8  8  3> 


Fig.  9. — Distribution  of  the  Phenac-omys  longicaudus  group: 

1.  General    distribution   of  P.    longicaudus,  as   shown   by 
specimens    examined. 

2.  Distribution  of  P.  sttvicola  ;  known,  from  two  localities 
only. 

Circles  show  type  localities  of  species 


1926]  REVISION    OP   GENUS  PHENACOMYS  33 

slightly  paler  on  the  sides.  Pelage  everywhere  plumbeous  at  base, 
showing  through  the  whitish  of  the  underparts  to  some  extent.  Tail 
blackish  and  rather  plentifully  haired,  and  the  ears  and  feet  spar- 
ingly covered  with  hairs  of  the  same  color  as  is  the  dorsal  surface. 
Late  spring  and  summer  adults,  as  well  as  juveniles,  are  much  duller, 
and  because  of  the  wearing  away  of  the  tips  of  the  hairs  in  the  case 
of  these  adults,  the  plumbeous  bases  of  the  hairs  show  through  to  a 
great  extent,  darkening  and  dulling  the  general  tone.  The  oldest 
two  specimens  have  many  pale  and  whitish  hairs  dorsally,  and  in 
the  Mount  Sanhedrin  skin  this  is  so  pronounced  that  a  distinctly 
grizzled  appearance  is  produced. 

Skull. — Brain  case  rather  short  and  squarish,  maxillary  processes 
of  the  zygoma  flaring  slightly  but  jugals  compressed  and  parallel, 
the  whole  giving  a  squat,  robust  appearance  to  the  skull.  Postorbital 
processes  of  the  squamosals  exceedingly  well  developed.  The  shape 
of  the  interpterygoid  fossa  is  not  at  all  constant  and  varies  from  U 
to  V  shaped.  The  ascending  branches  of  the  maxillae  usually  project 
a  full  millimeter  beyond  the  nasals  and  are  very  narrow. 

Dentition. — The  maxillary  dentition  of  loncftcaudms  presents  no 
peculiarities  except  for  the  presence  of  a  tendency  for  the  posterior 
outer  triangle  of  the  third  molar  to  be 
larger  and  more  completely  segregated 
from  the  posterior  loop  than  is  usual  in  the 
short-tailed  forms.  In  the  mandibular 
teeth  there  is  little  or  no  tendency  toward 
the  formation  of  an  external  triangle  on 

,,         ,,  .     t  t  m,  ...  P       ,,  Fig.  10. — First  lower  mo- 

tile  third    molar,      ine    variation   in   the  iarS,    left-hand    series, 

pattern    of   the    first    molar    is    so    great,  cLZT7£ie,*t e *&% 

especially  in  its  anterior  portion,  that  no  show  extremes  of  vari- 

t  -i    ^  i  -i-i  •■_      j>  ation     in     enamel    pat- 

dependency  can  be  placed  upon  its  iorm  tern.    Enlarged 

(fig.  10). 

Measurements. — The  averages  of  adults  of  the  two  sexes  is  as 
follows:  Six  males:  Total  length,  166  (158-176);  tail,  67  (60-72); 
hind  foot,  20  (19-21).  Five  females:  182  (170-187);  73  (66-83); 
21  (21-22).  Skull:  Averages  of  six  adult  males  as  follows:  Con- 
dylobasilar  length,  21.9;  nasals,  7.2;  interorbital  breadth,  3.5;  zygo- 
matic breadth,  14;  lambdoidal  width,  11.5;  incisive  foramina,  4.6; 
maxillary  tooth  row,  5.7;  height,  9.  The  averages  of  five  skulls  of 
adult  females  are  as  follows:  Condylobasilar  length,  22.5;  nasals,  7.8; 
interorbital  breadth,  3.3 ;  zygomatic  breadth,  14.2 ;  lambdoidal  width, 
11.9;  incisive  foramina,  4.6;  maxillary  tooth  row,  5.8;  height,  9.4. 

Remarks. — A  very  distinct  species,  needing  comparison  with  no 
other.  Its  reddish  coloration  is  distinctive,  and  its  short,  robust  skull 
is  readily  distinguishable  from  those  of  others.  The  degree  of  hairi- 
ness of  the  tail  and  the  arboreal  habitat  is  shared  with  silvicola,  from 
which  longicaudus  differs  externally  only  in  its  much  redder  color 
and  smaller  size,  ears  that  are  apparently  more  conspicuous,  pelage 
that  is  shorter  and  slightly  harsher,  and  face  without  sootiness. 
There  are  available  no  thoroughly  adult  skulls  from  Oregon  but  the 
material  seems  to  indicate  that  the  species  varies  little,  if  any,  with- 
in its  range. 

S.  G.  Jewett  informed  the  writer  that  there  are  specimens  of 
longicajudus  in  the  University  of  Oregon  from  Spencer  Butte,  near 


34  NORTH   AMERICAN   FAUNA  [No.  48 

Eugene,  and  that  unmistakable  evidence  was  found  in  old  nests  near 
the  Bonneville  Fish  Hatchery,  at  Bonneville  on  the  Columbia  River. 
Though  this  sign  may  possibly  have  been  of  longicamdus,  it  is  con- 
sidered more  likely  to  have  been  of  silvicola. 

Measurements  both  of  skins  and  skulls  indicate  that  there  is 
a  difference  in  size  between  the  sexes,  in  favor  of  the  females,  too 
great  to  be  ignored.  It  is  consequently  inferred  that  there  is  a 
difference  of  size  between  the  sexes  in  silvicola  as  well. 

It  seems  that  the  females  of  longicaudus  always  build  larger, 
more  readily  discernible,  and  probably  lower  nests  than  the  arbo- 
real shelters  of  the  males.  Recent  data,  discussed  in  Part  II,  point 
to  the  hypothesis  that  the  adult  males  normally  live  in  terrestrial 
holes  or  under  rubbish  at  the  bases  of  the  food  trees  and  probably 
construct  small,  temporary,  arboreal  nests  only  when  they  have 
found  females  that  are  ready  for  their  attention.  This  difference 
in  habits  according  to  sex  seems  to  be  the  reason  that  females  so 
far  outnumber  males  in  collection.  In  fact,  the  six  adult  males 
mentioned  above  seem  to  be  the  only  specimens,  whereas  a  con- 
siderably greater  number  of  females  and  young  have  been  available 
than  were  actually  assembled  during  the  present  study. 

Specimens  examined. — Total  number,  40,  as  follows : 

California:  Bridgeville,  239;   Carlotta,  28  M;  Mendocino  City,  l41;  Mount 

Sanhedrin,  1 42. 
Oregon:  Agness,  S43;  Eugene,  1;  Gold  Beach  (20  mi.  east),  2;  Marshfield,  1 

(the  type)  ;  Meadow,  1. 

PHENACOMYS  SILVICOLA  A.  B.  Howell 

Forest  Tree  Mouse 

(PI.  3,  fig.  6) 

Phenacomys  silvicolus,  A.  B.  Howell,  Jour.  Mamm.,  2 :  98,  1921. 

Type. — Collected  5  miles  southeast  of  Tillamook,  Tillamook 
County,  Oreg.,  October  25,  1916,  by  Peter  P.  Walker ;  female  young 
adult ;  No.  1214,  S.  G.  Jewett  coll. ;  original  number,  40,  A.  Walker 
coll. 

Geographic  distribution. — Known  only  from  the  type  locality, 
and  from  Corvallis,  Oreg.;  undoubtedly  confined  to  the  forested 
area  of  the  humid  coast  belt  (fig.  9,  £). 

Habitat. — Chiefly  arboreal. 

General  characters. — A  large  species  with  sooty  nose,  a  long  tail 
rather  well  haired,  long  pelage  of  a  warm  tone  of  brown,  and  long 
toes.  Skull  rather  weak  but  with  lateral  ridges  of  the  brain  case 
better  developed  than  in  any  other  form  of  the  genus.  Incisors 
sharply  decurved. 

Coloi\ — The  dorsal  coloration  of  the  type  is  of  a  warm  brown, 
close  to  the  cinnamon  brown  of  Ridgway,  with  many  of  the  hairs 
tipped  with  black;  a  trifle  paler  on  the  sides.  The  ventral  surface 
is  whitish,  these  and  all  other  body  hairs  being  plumbeous  prox- 

39  D.  R.  Dickey  coll. 

*°  D.  R.  Dickey  coll.,  2  :  Mus.  Vert.  Zool.,  3  ;  Mus.  Comp.  Zool.,  5  ;  U.  S.  Nat.  Mus.,  IS. 

41  Mus.  Vert.  Zool. 

42  Acad.   Nat.   Sci.   Philadelphia. 
«S.  G.  Jewett  coll. 


1926]  KEVISION"    OF   GENUS   PHENACOMYS  35 

imally.  The  hairs  of  the  tail  are  blackish,  darker  on  the  upper 
side,  and  the  nasal  area  is  noticeably  sooty  in  contrast  to  the  remain- 
der of  the  face. 

Skull. — The  brain  case  is  very  flat,  especially  posteriorly,  and  the 
temporal  ridges  unusually  pronounced,  being  largely  parallel;  but 
the  rest  of  the  skull  is  weak  in  appearance.  The  ascending  branches 
of  the  premaxillae  resemble  those  of  longicaudus  in  the  degree  to 
which  they  project  beyond  the  nasals. 

Dentition. — The  incisors  of  this  species  are  short  and  light,  and 
if  anything  more  sharply  decurved  than  in  longicaudus.  The  molars 
of  the  type  are  unusually  heavy  and  broad,  and  the  triangles  of  the 
lower  teeth  tend  to  remain  open  to  a  marked  degree,  but  in  the  second 
specimen  these  characters  are  less  pronounced.  The  two  topotypes 
were  examined  too  late  to  compare  them  with  the  type. 

Measurements. — Type:  Total  length,  191;  tail,  81;  hind  foot,  22. 
Male:  193;  87;  20.  Skull  (type)  :  Condylobasilar  length,  23.3;  inter- 
orbital  breadth,  3.1;  zygomatic  breadth,  14.4;  lambdoiclal  width,  11.9; 
incisive  foramina,  4.4 ;  maxillary  tooth  row,  6 ;  height,  9. 

Remarks. — Coloration,  combined  with  sooty  face  and  the  char- 
acter of  the  tail,  readily  distinguishes  this  species  externally,  while 
the  presence  of  the  parallel  temporal  ridges  is  sufficient  to  charac- 
terize the  skull.  Furthermore,  it  may  be  distinguished  from  longi- 
caudus by  the  almost  entire  suppression  of  the  postorbital  processes. 
A  unique  character  possessed  by  the  two  arboreal  species  is  the  great 
extent  to  which  the  upper  incisors  are  decurved,  and  the  ascending 
branches  of  the  premaxillae  project  beyond  the  nasals  to  a  greater 
degree  in  these  two  than  in  any  other  species  of  the  genus.  For  this 
reason,  as  well  as  because  of  the  close  similarity  of  external  points, 
silvicolai  is  here  placed  with  longicaudus.  It  is  of  interest  to  note, 
however,  that  in  several  less  noticeable  characters,  as  sootiness  of 
nasal  area,  the  resemblance  is  closest  to  albipes. 

In  some  respects,  the  case  of  silvicola  is  a  puzzling  one.  The  first 
impulse  is  to  consider  it  as  a  subspecies  of  longicaudus.  A  closer 
scrutiny  of  the  evidence,  however,  argues  for  full  specific  separation 
of  the  two,  at  least  for  the  present. 

The  type  was  found  dead  on  a  log  in  dense,  virgin  forest,  and  the 
second  specimen  was  taken  from  a  nest  in  a  tree. 

Specimens  examined :  Total  number,  4,  as  follows : 

Oregon :  Tillamook,  3  u ;  Corvallis,  1 45. 

**  S.  G.  Jewett  coll.,  1   (the  type)  ;  A.  Walker  coll.,  2. 
46  Oregon  Agr.  College  coll. 


Explanation  of  Plate  2 
[All  skulls  about  twice  natural  size] 

Fig.  1. — Phenacomys   intermedins   intermedins,   female,    Beartooth   Mountains, 

Mont.   (No.  66727,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  2. — Phenacomys  intermedins  intermedins,  female,  Nelson,  British  Columbia 

(No.  69052,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  3. — Phenacomys  intermedins  levis  (type),  male,  Saint  Marys  Lake,  Mont. 

(No.  72405,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  4. — Phenacomys    intermedins    olympicus,    male,    Soleduc    River,    Olympic 

Mountains,  Wash.    (No.  90375,  U.   S.  Nat.  Mus.,  Biological   Survey 

collection). 
Fig.  5. — Phenacomys  intermedins  celsus,  male,  Yosemite  National  Park,  Calif. 

(No.  23118,  Mus.  Vert.  Zool.). 
Fig.  6. — Phenacomys   intermedins    intermedins,    female,    Beartooth    Mountains, 

Mont.  (No.  66727,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 
Fig.  7. — Phenacomys  mackenzw,   male,    Fort    Smith,   Mackenzie,   Canada    (No. 

110437,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 

36 


North  American  Fauna  No.  48,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  2. 


Skulls  of  Phenacomys 


North  American  Fauna  No.  48,  U.  S.  Dept.  Agr.,  Biological  Survey 


Plate  3. 


Skulls  of  Phenacomys 


Explanation  of  Plate  3 
[All  skulls  about  twice  natural  size] 

Fig.  1. — Phenacomys  ungava  ungava  (type),  male,  Fort  Chirno,  Ungava,  Canada 
(No.  1864S7,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 

Fig.  2. — Phenacomys  ungava  crassus,  female,  Hamilton  Inlet,  Labrador    (No. 
242334,  U.  S.  Nat.  Mus.). 

Fig.  3. — Phenacomys  mackensn,   male,    Fort   Smith,   Mackenzie,   Canada    ( No. 
110437,  U.  S.  Nat.  Mus.,  Biological  Survey  collection). 

Fig.  4. — Phenacomys  albipes,  male,  Netarts,  Oreg.   (No.  789,  S.  G.  Jewett  col- 
lection). 

Fig.  5. — Phenacomys  longicaudus,  female,  Carlotta,  Calif.    (No.  2063SO,  U.   S. 
Nat.  Mus.,  Biological  Survey  collection). 

Fig.  6. — Phenacomys  silvicola    (type),  female,  Tillamook  County,   Oreg.    (No. 
1214,  S.  G.  Jewett  collection). 

37 


II.  LIFE  HISTORY  OF  THE  RED  TREE  MOUSE 
PHENACOMYS  LONGICAUDUS  TRUE 

By  A.  Brazier  Howell 


INTRODUCTION" 

In  accordance  with,  a  general  plan  for  the  study  both  in  the  field 
and  in  the  laboratory  of  the  rodent  subfamily  Microtinae,  particular 
attention  has  been  paid,  as  opportunity  has  offered,  to  the  red 
phenacoinys  {Phenacomys  longicaudus) — which  may  be  more  popu- 
larly known  as  the  red  tree  mouse  (see  frontispiece) — of  the  more 
southern  portion  of  the  humid  faunal  area  of  the  Pacific  coast.  Op- 
portunity for  this  work  occurred  during  a  part  of  July,  1919,  when 
it  was  prosecuted  by  the  writer  merely  in  conjunction  with  investiga- 
tions regarding  other  mammals  of  the  region,  and  in  July  and 
August,  1923,  when,  on  a  special  trip  made  by  him  for  this  purpose, 
a  life-history  study  was  undertaken  as  circumstances  permitted.  In 
an  excellent  report  on  this  species  by  TV.  P.  Taylor  (1915b),  there 
was  much  valuable  information  concerning  its  life  habits.  The 
present  contribution,  however,  is  believed  to  be  justified  in  that  it 
approaches  the  subject  from  a  somewhat  different  angle.  The  illu- 
strations in  Plates  1  and  6,  made  from  photographs  taken  by  the 
author,  are  of  captive  mice  placed  amid  natural  surroundings. 

Although  every  effort  has  been  made  to  avoid  theorizing,  it  has 
been  necessary  to  advance  some  hypotheses,  for  the  reason  that  it  has 
not  been  possible  to  observe  all  desired  phases  of  the  life  history  of 
the  species.  It  is  believed  that  in  the  light  of  present  knowledge 
these  are  correct ;  but  the  cases  in  point  have  been  indicated  and  the 
reader  may  draw  independent  conclusions  if  he  desires.1 

Phenacomys  longicaudus  is  a  small  rodent  largely  of  arboreal 
habits,  is  related  to  the  common  meadow  mouse,  and  occurs  through- 
out a  portion  of  the  humid  coast  belt  in  California  and  Oregon.  Its 
distribution  is  discontinuous  to  a  certain  extent,  but  it  is  found 
locally  in  some  numbers  in  forested  areas  of  Douglas  and  grand  firs. 
In  these  trees  it  largely  makes  its  home,  descending  to  the  ground 
when  it  must  do  so  to  reach  some  other  tree  or  for  other  activity  of 
an  unknown  extent.  Although  it  may  eat  the  tenderest  bark,  its 
chief  food  in  the  wild  state  consists  of  the  fleshy  portion  of  fir 
needles,  the  fibrous  part  of  which  is  not  consumed  but  is  used  almost 
exclusively  for  constructing  the  nests.  It  gathers  twigs  at  night  and 
piles  a  supply  upon  the  nest  for  consumption  during  the  day. 

1  In  this  work  the  writer  has  had  the  active  cooperation  of  H.  E.  Wilder,  who  during 
his  residence  of  13  years  in  this  region  has  had  better  opportunity  to  become  familiar 
with  the  habits  of  the  tree  mice  than  anyone  else.  It  is  strongly  felt  that  his  name 
should  have  appeared  as  coauthor  of  the  present  contribution,  but  as  he  has  declined  this 
course  grateful  acknowledgment  is  made  of  his  invaluable  aid  in  gathering  the  data  pre- 
sented and  for  criticism  of  the  manuscript, 

39 


40  NORTH   AMERICAN"   FAUNA  [No.  48 

Structurally,  tree  mice  exhibit  only  slight  modifications  for  an 
arboreal  habitat,  at  least  as  far  as  may  be  stated  without  careful 
anatomical  studies.  The  length  and  hairiness  of  the  tail  and  the 
stout  feet  with  long  claws  are  apparently  the  only  characters  that 
have  developed  in  this  direction.  The  writer's  opinion,  however, 
is  that  the  slightness  of  this  structural  modification  is  not  due  to  an 
arboreal  life  for  a  short  length  of  time,  geologically  speaking,  but 
rather  to  the  probability  that  the  species  has  had  very  little  com- 
petition ecologically  and  few  predaceans  with  which  to  contend. 

These  mice  are  now  confined  to  a  rather  humid  area,  for  the  most 
part  covered  with  luxuriant  forests.  As  dense,  humid  forests  were 
in  remote  times  far  more  extensive  than  at  present,  it  is  not  un- 
likely that  arboreal  mice  identical  with  or  closely  related  to  the 
species  under  discussion  were  once  of  much  greater  importance  in 
the  American  fauna  than  is  the  remnant  now  surviving. 

Distributional    Factors 

The  geographic  range  of  Phenacomys  longicaudus  is  indicated  on 
the  map  (fig.  9).  Although  the  range  is  discontinuous  from  the 
very  nature  of  the  case,  the  mice  may  very  well  be  more  uniformly 
distributed  over  suitably  timbered  territory  than  has  yet  been 
proved.  In  general,  the  species  occupies  the  humid  coast  f  aunal  area 
from  Mendocino  County,  Calif.,  north  through  the  corresponding 
portion  of  Oregon.  There  is  apparently  no  reason  why  it  should  not 
occur  in  similar  parts  of  Washington  as  well,  but  it  has  never  been 
detected  within  the  boundaries  of  that  State. 

It  is  difficult  to  make  precise  statements  regarding  the  distribu- 
tional preferences  of  the  species.  It  is  not  to  be  found  in  pure 
stands  of  redwood,  for  it  will  not  feed  upon  this  tree;  but  it  does 
occur  in  its  favorite  trees  along  the  edges  of  these  pure  stands,  and 
where  there  is  mixed  timber.  In  Oregon  the  mice  are  presumed  to 
occur  here  and  there  throughout  most  of  the  coastal  belt  where  this  is 
suitably  forested ;  but  stations  of  capture  are  lacking  in  the  extreme 
northwestern  portions  of  the  State.  The  easternmost  records  are 
from  localities  with  considerably  greater  tendencies  toward  aridity 
than  is  the  case  over  the  greater  portion  of  the  range  of  the  species. 
It  occurs  in  both  the  Transition  and  Canadian  Zones,  but  in  certain 
parts  of  the  coast  belt  where  it  is  to  be  found  between  densely  tim- 
bered hillsides  and  gulches  on  the  one  hand,  and  bare  or  brushy 
slopes  and  bottom  lands  on  the  other,  many  zonal  indicators  nor- 
mally dependable  are  here  so  inextricably  interwoven  that  one  is 
forced  to  be  cautious  in  making  exact  statements. 

Within  the  general  area  studied  the  tree  mice  may  occur  wherever 
there  is  a  stand,  not  too  dense,  either  of  Douglas  or  grand  fir  or 
both.  This  condition,  as  previously  indicated,  is  often  to  be  found 
upon  the  borders  of  dense  redwood  forests,  notably  upon  the 
shoulders  of  southward-facing  hills  (pi.  4).  Elsewhere  than  in 
redwood  country  their  preferences,  according  to  information  pub- 
lished by  others,  may  be  more  difficult  of  proper  interpretation. 
One  distributional  factor  of  paramount  importance  is  that  of  ac- 
cessibility. The  species  will  seldom  or  never  be  found  in  naturally 
isolated  patches  of  timber,  even  though  these  be  of  comparatively 
large  extent.     Theoretically  at  least  a  considerable  expanse  of  brushy 


1926]  LIFE   HISTORY   OF   PHENACOMYS   LONGICAUDUS  41 

or  open  land  without  suitable  trees  is  an  absolute  barrier  to  the 
species.  As  with  all  animals,  there  are  doubtless  other  distribu- 
tional factors  not  discernible,  but  which  nevertheless  operate  to 
prevent  the  tree  mice  from  occupying  certain  territory  that  would 
seem  ideal  for  them.  So  far  as  the  writer's  experience  goes,  how- 
ever, as  well  as  that  of  others  with  whom  he  has  talked,  the  species 
is  indiscriminate  in  its  choice  of  north  or  south  slopes,  hilly  or  level 
ground. 

The  technical  description  of  the  species  and  details  of  its  relation- 
ships are  given  in  Part  I. 

Investigational    Methods 

The  methods  employed  in  the  present  investigation  may  be  divided 
into  observation,  experimentation,  and  deduction.  Especially  dur- 
ing 1923  the  most  careful  observations  were  made  of  all  phases 
of  the  life  and  ecology  of  the  mice  that  it  was  possible  to  discover, 
and  their  habits  carefully  studied.  An  area  was  selected  and  every 
accessible  nest  within  it,  and  many  without,  was  dissected.  Sev- 
eral of  the  mice  were  caught  and  kept  in  captivity  for  varying 
lengths  of  time,  and  feeding  and  other  experiments  conducted. 
As  the  mice  are  normally  abroad  only  at  night,  certain  details  of 
their  habits  can  only  be  deduced,  sometimes  with  the  aid  of  captive 
individuals.  Though  it  is  admitted  that  the  actions  of  captives 
are  not  perfectly  normal,  data  from  such  a  source  are  of  much 
value  when  so  stated.  They  are  often  the  best  to  be  had,  and  need 
only  be  accepted  by  the  reader  for  what  he  deems  they  are  worth. 
It  must  also  be  borne  in  mind  that  the  writer  has  studied  Phena- 
comys  longicaudus  in  detail  in  only  one  part  of  its  range,  and  it  is 
very  possible,  or  probable,  that  many  of  the  conclusions  reached 
will  not  be  found  to  apply  with  equal  force  to  the  mice  of  all  other 
sections. 

AREA  INVESTIGATED 

The  present  study  was  made  chiefly  on  the  hillside  surrounding 
the  home  of  H.  E.  Wilder,  1  mile  east  of  Carlotta,  Humboldt  County, 
Calif.  This  hillside  faces  toward  the  south,  and  the  lower  portion 
consists  of  open  pasture  and  meadowland  adjoining  the  cultivated 
fields  of  the  valley  of  the  Van  Duzen  River  (pi.  4).  Above  this 
there  is  considerable  brush,  clumps,  or  single  specimens  of  rather 
small  and  occasional  large  conifers,  with  a  few  stumps  showing 
where  others  have  been  cut  off.  This  strip,  extending  in  some  places 
to  the  floor  of  the  valley,  and  in  others  for  a  distance  of  200  to  300 
feet  up  the  hillside,  is  interspersed  with  small  patches  and  tongues  of 
open  grassland ;  but  this  is  not  true  "  cut-over  "  land,  Mr.  Wilder 
says.  Rather  is  it  a  natural,  transitional  strip  in  which  many  trees 
remain  small,  located  between  the  originally  open  or  somewhat 
brushy  bottom  lands  and  the  dense  redwood  forest  of  the  higher 
hills  and  upper  gulches. 

The  tract  studied  (fig.  11)  is  perhaps  5  acres  in  extent;  but  the 
mice  are  probably  as  numerous  in  another  tract  of  about  the  same 
size  immediately  beyond  a  small  gulch  to  the  east,  where  the  trees 
average  larger.  They  also  occur  in  scattered,  outlying  clumps  of 
trees  beyond  the  confines  of  these  areas.     Eastward  they  were  ob- 


42 


NORTH    AMERICAN    FAUNA 


[No.  48 


tained  several  miles  up  the  river,  and  Wilder  has  found  others  else- 
where in  this  general  region.  Nowhere  near  by  are  they  so  numerous, 
at  least  in  the  smaller  trees,  as  upon  the  area  studied,  and  it  was 
selected  as  much  for  this  reason  as  because  it  is  the  most  accessible. 
The  trees  occupied  are  also  smaller  than  usual,  and  most  of  the 
specimens  in  the  collections  of  North  America  have  come  from  just 
this  spot. 

Throughout  their  range  these  mice  usually   frequent  the  taller 
trees,  and  for  this  reason  it  is  extremely  difficult  to  estimate  their 


o 

Grand   Ff 

r 

• 

Douglas 

Fir 

WA 

Brush 

X 

Nests 

GARDEN 


HOUSE  BARN 


Fig.   11.. — Diagram  of  area  near   Carlotta,   Calif.,   harboring  colony  of  Phcnacomys 
longicaudus,  showing  major  details,  with  positions  of  nests  and  nesting  trees 

numbers,  for  an  immense  Douglas  fir  might  harbor  a  dozen  nests  that 
can  not  be  recognized  as  such  from  the  ground.  In  cutting  a  number 
of  these  trees  for  firewood,  Wilder  found  that  practically  every 
one  contained  the  nest  of  a  mouse.  It  is  logical  to  assume,  therefore, 
that  a  large  proportion  of  trees  in  localities  favored  by  the  mice  are 
occupied,  and  it  seems  probable  that  the  majority  of  specimens  have 
been  collected  from  smaller  trees  simply  because  nests  in  lofty  situa- 
tions are  well  nigh  inaccessible.    It  may  well  be  that  very  often  such 


North  American  Fauna  No.  48,  U.  S.  Dept  Agr.,  Biological  Survey. 


Plate  4. 


North  American  Fauna  No.  48,  U.  S.  Dept.  Agr.,  Biological  Survey. 


PLATE  5. 


Typical  Nesting  Tree  of  Phenacomys  longicaudus 

Douglas  fir  (Pscndotsuga  taxifolia)  in  the  Carlotta,  Calif.,  area,  in  which  were  nests  numbered  4,  5, 
and  6  in  the  text.    Approximate  location  of  nests  shown  by  arrows 


1926]  LIFE   HISTORY  OF   PHENACOMYS  LONGICAUDUS  43 

lesser  growth  harbors  the  overflow  population  from  more  lofty 
neighbors.  On  the  other  hand,  Wilder  states  that  he  has  walked  for 
many  miles  beneath  large  trees  that  seemed  eminently  suitable  as 
habitations  of  the  mice,  without  seeing  a  sign  of  their  presence,  al- 
though he  usually  had  them  in  mind.  This  evidence  further  ac- 
centuates the  degree  to  which  the  species  is  localized. 

It  seems  safe  to  state  that  in  no  place  yet  investigated  by  a  mam- 
malogist  are  the  tree  mice  so  numerous  as  they  originally  were  at 
Carlotta.  In  two  successive  mornings  during  1913  Wilder  and  C.  I. 
Clay  captured  more  than  50  individuals  on  this  hillside,  and  fully  as 
many  more  have  since  been  obtained  there.  They  are  not  now  so 
abundant  as  at  first,  although  few  have  been  collected  in  the  past 
three  or  four  years,  and  Wilder  considers  that  they  are  slow  to  re- 
cover from  any  check. 

The  trees2  within  the  area  covered  by  the  investigation  (fig.  11) 
consist  of — 

Pseudotsuga  taxifolia Douglas  fir  (pi.  5). 

Abies  grandis Grand  fir. 

Sequoia  sempervirens Redwood. 

Arbutus  menziesii Madrone. 

Pasania  densiflora Tan  oak. 

Umbellularia  californica California  bay  laurel. 

The  first  two  of  these  predominate,  and  are  the  only  species  of  tree 
indicated  on  the  diagram.  Among  scattered  individuals  are  small 
redwoods,  but  there  are  several  very  large  madrones. 

The  k'  brush,"  as  indicated  on  the  diagram,  is  in  the  form  of  rather 
open  thickets  and  clumps,  from  8  to  15  feet  tall,  of — 

Ceanothus  thyrsifiorus California  lilac. 

Corylus  rostrata  californica Hazel. 

Holodiscus  discolor  ariaefolia Spiraea,  or  arrowwood. 

NESTS 

The  largest  conifers  in  this  area  could  be  prospected  for  nests  only 
from  the  ground  and  many  nests  surely  escaped  detection.  Simi- 
larly, it  is  doubtful  whether  all  the  smaller,  inconspicuous  nests  in 
the  tops  of  the  denser  trees,  even  when  these  were  small  enough 
to  be  readily  explored,  were  discovered.  The  approximate  position 
of  each  of  the  nests  that  could  be  found  and  reached  is  shown  in  the 
diagram  by  a  cross,  and  by  a  number  when  the  nest  could  be  ex- 
amined. Letters  denote  inaccessible  bunches  of  debris  which  seemed 
to  be  the  nests  of  mice  when  viewed  through  binoculars  from  the 
ground. 

Nest  No.  1. — In  a  10-inch  grand  fir  16  feet  from  the  ground  and  8  feet  from 
the  trunk  on  a  slender  limb;  at  an  angle,  taken  with  a  compass,  of  100°,  south 
of  east.  This  nest  was  excessively  flattened  and  about  4  inches  deep  by  16  in 
diameter,  with  many  green  twigs  on  top.  Occupied  by  a  fully  grown  female 
without  sign  of  having  suckled  young,  which  when  released  ran  off  over  the 
surface  of  the  ground  without  attempting  to  enter  the  numerous  holes  and 
crannies  beneath  fallen  logs. 

Nest  No.  2. — In  an  8-inch  Douglas  fir  30  feet  from  the  ground  and  4  feet  from 
the  trunk.  It  was  at  an  angle  of  230°,  southwest  from  the  trunk,  upon  a 
slender  branch  midway  toward  another  somewhat  smaller  fir.  The  nest  was 
fairly  new  and  loosely  put  together ;  consequently,  entry  was  possible  almost 
anywhere  and  the  passages  were  difficult  to  define.  There  was  a  great  quan- 
tity of  green  twigs  on  the  nest  and  a  wad  of  them  pulled  into  the  burrow  from 

2  All  botanical  identifications  have  been  kindly  furnished  by  Mrs.  H.  E.  Wilder. 


44  NORTH   AMERICAN   FAUNA  [No.  48 

the  top  toward  the  center.  The  whole  mass  was  about  8  inches  deep  by  10 
in  diameter,  and  far  down  was  the  true  nest  chamber,  in  which  was  a  single 
young  about  3  days  old.  Female  not  seen  until  it  had  left  the  nest. 
When  released  upon  the  tree  trunk  it  ran  half  way  to  the  nest,  crossed  over 
to  the  next  tree,  then  back  at  a  level  above  the  nest,  and  off  out  of  sight. 

Nest  No.  3. — This  nest  could  not  be  reached  but  was  investigated  by  poking 
it  with  a  pole  from  beneath.  It  was  14  feet  above  the  ground  in  a  12-inch 
Douglas  fir  on  the  hillside,  and  15  feet  from  the  trunk,  at  an  angle  of  140°, 
southeast.  This  was  a  medium-sized  nest,  about  6  inches  deep  and  10  by  12 
in  diameter,  with  green  twigs  on  top.  There  was  much  debris  under  it,  indi- 
cating that  it  had  been  at  least  partially  destroyed  and  then  rebuilt.  The 
mouse,  which  must  have  been  a  female,  judging  from  the  size  of  the  nest,  ran 
out  and  then  up  through  the  foliage. 

Nest  No.  4. — Situated  with  the  two  following  nests  in  a  large,  2-foot  Douglas 
fir  (see  pis.  5  and  6),  10  feet  above/ the  ground  and  14  feet  from  the  trunk, 
at  an  angle  of  180°,  south.  This  was  a  large  nest  and  rather  new,  8  inches 
deep  and  12  by  14  in  diameter  but  was  unoccupied. 

Nest  No.  5—  In  the  same  tree  with  Nos.  4  and  6,  50  feet  above  the  ground 
and  15  from  the  trunk,  at  an  angle  of  20°,  east  of  north.  This  was  a  small 
nest,  looking  much  like  that  of  a  male,  but  unoccupied. 

Nest  No.  6. — In  the  same  tree  with  the  last  two,  25  feet  above  the  ground 
and  12  from  the  trunk,  at  an  angle  of  160°,  east  of  south.  This  was  rather 
large,  perhaps  12  inches  in  diameter,  with  many  green  twigs  on  top,  but  noth- 
ing appeared  when  it  was  shaken  to  pieces. 

Nest  No.  7. — In  a  grand  fir  about  2  feet  in  diameter,  35  feet  above  the  ground 
and  15  from  the  trunk,  at  an  angle  of  240°,  south  of  west.  This  was  a  medium- 
sized  nest,  apparently  with  green  twigs  upon  it,  but  inaccessible.  There  was 
communication  only  with  surrounding  hazel  and  spiraea  brush. 

Nest  No.  8. — In  the  same  tree  as  No.  7,  50  feet  above  the  ground  and  15  from 
the  trunk,  at  an  angle  of  190°,  south.  Slightly  smaller  than  No.  7,  and  also  in- 
accessible, but  with  many  green  twigs  on  the  ground  beneath. 

Nest  No.  9. — In  a  2-foot  Douglas  fir,  40  feet  up  and  20  from  the  trunk,  at  an 
angle  of  40°,  northeast.  A  medium-sized  nest,  10  or  12  inches  in  diameter, 
but  old  and  abandoned. 

Nest  No.  10. — In  the  same  tree  as  No.  9,  35  feet  above  the  ground  and  15  feet 
out  on  a  limb,  at  an  angle  of  200°,  west  of  south.  This  was  a  medium  to  large 
nest,  a  foot  or  more  in  diameter,  with  plenty  of  green  twigs  on  top.  No  mouse 
appeared,  however,  when  it  was  shaken  down. 

Nest  No.  11. — In  a  14-inch  Douglas  fir,  50  feet  above  the  ground  and  5  from 
the  trunk,  at  an  angle  of  330°,  north  of  northwest ;  4  inches  deep  and  10  by  15 
inches  in  diameter.  It  was  in  a  thick  bunch  of  twigs  and  very  shallow,  with 
the  nest  chamber  barely  covered,  but  well  protected  above  by  live  twigs.  Occu- 
pied by  an  adult  female. 

Nest  No.  12. — In  a  grand  fir  2  feet  in  diameter,  about  45  feet  above  the  ground 
and  3  from  the  trunk,  at  an  angle  of  290°,  north  of  west.  The  nest  was  a  few 
inches  from  a  3-inch  limb,  between  a  flat  piece  of  dense,  live,  parasitic  growth 
above  and  a  dead  platform  of  the  same  below.  It  was  exceedingly  flat,  being 
3  inches  deep  and  10  by  12  in  diameter.     Occupied  by  an  adult  female. 

Nest  No.  13. — In  a  2-foot  grand  fir  about  45  feet  up  at  an  angle  of  140°, 
southeast,  not  more  than  20  feet  from,  and  precisely  on  the  same  level  with, 
nest  No.  12.  At  about  38  feet  from  the  ground  the  old  trunk  had  been  severed 
10  years  before  in  order  to  make  it  more  branching,  and  the  nest  was  a  foot 
from  the  several  small  shoots  resulting.  It  was  a  mere  platform,  slightly 
cupped,  \y-2  inches  deep  and  4  by  5  in  diameter,  unroofed  but  well  sheltered 
immediately  above  by  a  dense  little  cluster  of  twigs.  When  the  writer  was 
yet  3  feet  away  the  mouse  sprang  into  space  and  descended  without  having 
its  fall  broken  by  intervening  twigs.  It  was  lying  perfectly  still  when  picked 
up,  but  fully  recovered  in  a  few  seconds.  It  proved  to  be  an  adult  female 
with  signs  of  having  suckled  young.  Its  nest  must  have  been  temporary,  and 
was  surprisingly  small  for  an  adult  female;  but  it  could  hardly  have  been  the 
beginning  of  a  family  nest,  for  there  was  not  sufficient  foundation  for  the 
support  of  one. 

Nest  No.  H. — In  a  grand  fir  10  inches  thick  but  only  30  feet  high  because 
the  top  was  once  cut  off.  This  operation  has  been  followed  by  the  usual 
concentric  new  growth  and  little  platform,  upon  which  was  the  nest,  20  feet 
above  the  ground.  It  was  a  large,  family  affair,  10  inches  deep  and  12  by 
14  inches  in  diameter,  and  comparatively  new,  with  an  unusual  number  of 


1926]  LIFE   HISTORY   OF   PHENACOMYS  LONGICATJDUS  45 

stripped  twigs  entering  into  its  composition.3  There  was  communication  only 
with  low  hazel  brush  on  two  sides,  thence  to  a  small  redwood  and  a  grand  fir ; 
but  this  whole  clump  is  well  segregated  from  the  nearest  brush  25  feet  across 
a  road,  and  an  animal  reaching  it  would  have  to  travel  over  bare  ground 
for  at  least  that  distance. 

Nests  a,  b,  and  c. — These  were  in  a  large  Douglas  fir  well  segregated  from 
surrounding  growth  because  of  the  great  height  of  the  limbs.  The  three  nests, 
or  what  appeared  from  the  ground  to  be  nests,  were  at  heights  ranging  from 
60  to  100  feet  or  more.  Numerous  twig  cuttings  on  the  ground  beneath  made 
it  certain  that  this  tree  was  inhabited  by  at  least  one  mouse,  and  possibly 
more. 

Nest  d. — This  was  the  remains,  10  feet  up  in  a  small  Douglas  fir,  of  a  very 
old  and  very  large  nest  which  Wilder  says  was  inhabited  several  years  ago ; 
but  when  observed  it  was  merely  a  substantial  platform  of  punk  and  fine 
debris  with  a  few  twigs  intermingled,  the  whole  about  12  by  15  inches  in 
diameter  and  3  or  4  inches  deep. 

It  can  not  be  said  that  every  nest  within  the  area  surveyed  was 
discovered  and  studied.  It  is  certain  that  a  number  in  the  largest 
trees  escaped  observation,  and  it  is  extremely  probable — indeed, 
practically  certain — that  several  nests  situated  even  at  moderate 
heights  did  also,  especially  the  smaller  ones  of  males.  It  would 
not  be  at  all  surprising  if  these  undiscovered  nests  should  constitute 
40  per  cent  of  the  total  within  the  area  and  if  the  tree-mouse  popula- 
tion should  be  nearly  double  that  indicated  by  the  investigations. 

Construction  and  Site  of  Nests 

The  average  measurements  of  the  15  nests  dissected  were  as  fol- 
lows: Height  above  ground,  31  feet;  distance  out  from  trunk,  9 
feet,  at  an  angle  of  109°,  barely  east  of  south.  It  is  believed,  how- 
ever, that  this  evidence  of  the  propensity  of  tree  mice  to  construct 
nests  on  the  south  sides  of  trees  should  not  be  accepted  without 
reservation  until  substantiated  by  observations  in  other  districts. 
It  is  fully  as  likely  that  the  southern  aspect  was  selected  merely 
because  that  happened  to  be  the  downhill  side  as  for  any  other 
reason.  Similarly,  no  preference  seems  to  have  been  shown  as 
regards  open  or  densely  shaded  situations,  sites  evidently  having 
been  chosen  at  random  or  with  a  view  to  obtaining  the  most  suitable 
foundation. 

Only  two  of  the  nests  dissected  were  placed  against  the  trunks 
of  trees,  the  remainder  being  at  varying  distances  away.  The 
majority  of  the  nests  discovered  by  Taylor  in  Mendocino  County 
were  situated  immediately  adjacent  to  the  trunks,  as  also  were 
those  found  by  Shelton  in  Oregon  (Taylor,  1915b,  p.  145),  but  these 
were  built  chiefly  upon  old  squirrel  nests,  so  that  the  situation  had 
to  a  considerable  extent  already  been  chosen  for  the  mice. 

In  the  Carlotta  district  typical  family  nests  "were  about  12  inches 
in  diameter  by  9  in  depth,  while  the  arboreal  nests  of  the  males  and 
subadult  females  were  smaller.  Barely  were  they  larger,  the  most 
bulky  one  which  the  writer  has  seen  being  hardly  twice  that  mass; 
but  Taylor  states  that  nests  observed  by  his  Museum  of  Vertebrate 
Zoology  party  varied  in  dimensions  from  about  18  inches  in  length, 
breadth,  and  height,  to  3  feet  in  the  horizontal  diameter  and  2  or  3 
feet  in  the  vertical.    Evidentty,  in  districts  in  which  such  enormous 

3  In  January,  1924,  Wilder  discovered  the  small,  inconspicuous  nest  of  a  male  well  out 
toward  the  end  of  a  limb  on  this  tree.  It  may  or  may  not  have  been  there  during  the 
visit  of  the  writer. 


46  NORTH  AMERICAN   FAUNA  [No.  48 

structures  occur  certain  conditions  are  more  favorable  for  the  long 
occupancy  of  a  nest  than  about  Carlotta.  As  long  as  a  nest  is  un- 
disturbed it  will  naturally  grow  in  size  because  of  the  daily  accumu- 
lation of  food  refuse.  If  it  does  not  increase  to  considerable  dimen- 
sions, it  is  either  because  the  mice  object  to  nests  of  such  size  and 
abandon  them  after  they  reach  a  certain  stage,  or  else  because  some 
agency  that  may  be  classed  as  an  enemy  destroys  the  nests  before 
these  have  time  to  grow  to  great  size. 

Arboreal  nests  may  be  placed  at  any  height  above  the  ground  and 
in  trees  of  any  size.  Wilder  has  found  two  nests — of  both  male  and 
female — a  few  feet  above  the  ground  in  fir  saplings  well  segregated 
from  other  trees,  and  in  larger  growth  they  occur  up  to  a  great 
height.  z 

Nests  of  the  tree  mice  in  the  Carlotta  district  are  normally  con- 
structed entirely  by  the  animals  themselves,  and  consist  only  of  the 
discarded  portion  of  the  fir  needles  used  for  food,  among  which 
may  occur  a  few  of  the  stripped  twigs  from  which  the  needles  were 
obtained.  Very  rarely  one  may  find  that  a  mouse  has  utilized  as  the 
foundation  of  a  nest  a  chance  accumulation  of  fallen  twigs  and  rub- 
bish which  has  lodged  in  a  favorable  situation,  or  a  clump  of  para- 
sitic growth  may  be  similarly  employed. 

Indications  were  found  farther  east  along  the  Van  Duzen  River, 
where  squirrels  {Sciurus)  are  more  plentiful,  that  the  abandoned 
nests  of  these  rodents  are  occasionally  used  as  a  scaffolding  on  which 
homes  of  the  tree  mice  are  constructed.  Taylor  (1915b,  p.  146) 
records  that  near  the  Hearst  and  Lierly  ranch,  Mendocino  County, 
Calif.,  all  nests  of  the  tree  mice  were  in  old  homes  of  gray  squirrels 
and  that  some  were  of  large  proportions.  Undoubtedly  the  nests  of 
the  larger  animals  had  first  been  abandoned  and  then  were  readily 
appropriated  as  foundations  on  which  the  more  specialized  homes  of 
the  mice  were  erected.  This  indicates  that  the  tree  mice  are  ready  to 
adapt  themselves  to  circumstances  to  some  extent,  and  that  they  may 
build  upon  anything  that  offers  suitable  foundation;  but  it  is  not 
unlikely  that  they  are  very  slow  to  adopt  any  such  innovation. 

Of  the  15  nesting  sites  studied  in  detail,  9  were  in  Douglas  and  6 
in  grand  firs,  the  former  evidently  being  more  often  favored  where 
they  are  equally  abundant;  but  preference  probably  varies  to  some 
extent  according  to  locality.  Clay  has  taken  several  nests  in  Sitka 
spruces  (Picea  sitchensis)  (Taylor,  1915b,  p.  140),  but  this  ex- 
perience seems  to  be  unique  among  observers,  and  it  is  not  known  to 
what  extent  this  tree  is  so  utilized.  He  has  stated  in  a  letter  that 
these  nest  trees  were  situated  close  to  Douglas  firs  and  it  may  well 
be  that  the  mice  did  not  use  the  spruces  as  food.  In  a  letter  to  the 
Biological  Survey  he  has  also  stated  that  he  observed  nests  in  both 
myrtle  and  redwood  trees,  but  it  is  probable  that  in  these  cases  condi- 
tions were  similar  to  the  single  instance  observed  by  Wilder  of  a  nest 
in  a  small  redwood,  the  branches  of  which  interlaced  with  those  of  a 
fir.     The  latter  undoubtedly  furnished  the  food  supply. 

Nests  sometimes  appear  unexpectedly  in  situations  where  there  were 
none  before,  according  to  Wilder,  so  that  old  material  must  at  times 
be  utilized  in  the  construction  of  a  new  home.  In  fact,  this  must  be 
the  usual  procedure  when  it  is  at  all  possible,  for  it  would  take 
several  days  for  sufficient  fresh  refuse  to  accumulate  from  an  animal's 


North  American  Fauna  No.  48,  U.  S.  Dept.  Agr.,  Biological  Survey. 


Plate  6. 


Fig.  I. — Female  Red  Tree  Mouse  at  Nest 

The  nest,  No.  4  in  the  tree  in  Plate  5,  was  partiy  dissected 


Fig.  2. — Female  Red  Tree  Mouse  in  Fir  Tree 

Photographed  on  drooping  limb  of  Douglas  fir,  near  Carlotta,  Calif. 


North  American  Fauna  No.  48,  U.  S.  Dept.  Agr.,  Biological  Survey 


Plate  7. 


Fig.   I. — Young  of  Red  Tree  Mouse,  About  18  Days  Old 


iMr 

'  ^ 

WmM 

j/j    s^ 

fa.   ^ 

Is 

If  1^31 

M" 

^H 

™5| 

^wp? 

rWwvrf 

2 

Fig.  2. — Day's  Food  of  One  Red  Tree  Mouse 

Twigs  of  Douglas  fir  aggregating  70  inches  and  weighing  19  grams,  and  beneath  them  a  pile  of 
needles  weighing  7.9  grams,  stripped  from  70  inches  of  similar  twigs  by  one  individual  in 
24  hours 


Fig.  3.— Cross  Section  of  Fir  Needle 

From  Douglas  fir  (Pscudotsuga  taiifolia) ;  greatly  magnified,     a,  Lateral  resin  ducts;  6,  vascular  bundles 


192G] 


LIFE   HISTORY   OF   PHENACOMYS  LONGICAUDUS  47 


food  to  afford  it  protection  and  shelter.  It  may  then  be  assumed 
that  after  a  site  has  been  selected,  material  from  some  abandoned 
structure  is  transported  in  the  mouth,  as  the  writer  has  often  ob- 
served captives  do,  and  a  small  platform  of  this  is  deposited.  As 
this  grows  to  sufficient  size  the  animal  scratches  surplus  material 
upward  on  all  sides,  until  finally,  by  twisting  the  body  first  to  one 
side  and  then  the  other  and  scratching  above  the  head,  a  roof  is 
formed.  The  further  construction  of  a  larger  home  is  merely  inci- 
dental to  this.  There  is  no  definite  plan — no  interweaving  of  twigs, 
nor  even  so  much  uniformity  as  is  often  to  be  found  in  other  mam- 
malian tree  nests — apparently  nothing  but  a  haphazard  dropping  of 
nesting  material  in  all  directions,  and  a  pushing  and  pulling  to  fill 
up  vacant  spaces. 

In  the  usual  nest,  at  least  during  the  drier  months,  the  nest  cham- 
ber proper  is  near  the  top.  and  rises  as  the  whole  structure  increases 
in  height  and  the  floor  is  elevated  by  the  gradual  addition  of  a  part  of 
the  daily  food  refuse.  Such  a  nest  chamber,  with  its  slight  covering, 
will  be  placed  beneath  any  available  shelter,  as  a  limb  or  dense  little 
cluster  of  twigs.  In  some  cases  this  chamber  is  buried  deep  in  the 
whole  mass,  probably  by  the  occupant's  dropping  food  refuse  upon 
the  top. 

In  structures  of  sufficient  size  there  is,  in  addition  to  the  nest 
proper,  a  second  small  chamber  used  for  excretory  purposes.  It 
can  not  be  stated  at  just  what  stage  this  is  added,  but  from  the 
actions  of  captive  animals  it  would  seem  that  when  the  nest  is  of 
sufficient  size,  the  original  chamber  is  often  used  for  depositing  the 
feces  and  that  new  sleeping  quarters  are  formed.  During  prolonged 
rains  the  animals  undoubtedly  seek  the  lowest,  driest  parts  of  the 
nests. 

In  the  matter  of  passages,  as  in  other  respects,  the  nests  seem  to 
follow  no  fixed  plan,  but  the  runways  are  formed  or  abandoned, 
especially  in  loosely  built  structures,  as  the  animal  desires.  There 
is  practically  always  an  exit  immediately  above  for  easy  access  to 
the  diurnal  food  pile,  and  another  at  an  inconspicuous  point  of 
egress,  as  directly  beneath  a  small,  protruding  branch.  The  degree 
of  compactness  of  a  nest  depends  partly  upon  whether  it  is  new  or 
old,  and  probably  to  a  great  extent  upon  whether  its  basic  structure 
has  recently  been  altered  by  the  mouse  or  some  extraneous  agent. 

There  may  be  many  exceptions  to  the  rule,  but  in  the  writer's 
experience  the  large  nests  and  those  of  medium  size  belong  to  fe- 
males, whereas  the  arboreal  nests  of  the  males  are  more  likely  to 
be  small,  often  only  a  ball  of  material  little  larger  than  one's  fist. 
Furthermore,  in  addition  to  being  less  conspicuous  on  account  of 
size,  the  nests  of  males  are  usually  better  hidden — either  higher,  in 
the  tops  of  the  trees,  or  farther  out  upon  the  limbs. 

In  pulling  nests  to  pieces  one  may  often  note  small  insects  crawling 
about  in  profusion,  which  the  uninitiated  are  inclined  to  pronounce 
parasitic.  These,  according  to  H.  E.  Ewing,  of  the  Bureau  of  Ento- 
mology, are  the  young  of  one  of  the  species  of  millipeds  belonging 
to  the  group  that  is  scaly  and  hairy.  He  states  that  these  are  often 
found  in  the  nests  of  mice  and  in  other  places  where  there  is  fecal 
and  decaying  vegetable  matter ;  hence,  they  are  not  parasites  of  the 


48  NORTH   AMERICAN   FAUNA  [No,  48 

mice.  The  writer  has  noted  mites,  but  no  fleas,  on  mice  previously 
collected,  but  unfortunately  none  of  these  were  preserved  and  his 
captive  specimens  are  apparently  free  from  them. 

YOUNG 

The  young  (pi.  7,  fig.  1)  are  probably  brought  forth  practically 
throughout  the  year.  Wilder  has  often  found  them  in  the  nests 
during  January  and  February;  but  they  are  perhaps  more  common 
in  spring,  summer,  and  early  fall.  The  average  number  to  a  litter 
is  two,  although  there  are  often  one  or  three,  and  they  are  naked, 
blind,  and  helpless  at  birth,  as  one  would  expect. 

Repeated  efforts  to  ascertain  the  length  of  the  gestation  period 
thus  far  have  been  unsuccessful.  After  trying  in  vain  to  breed  two 
females  that  were  reared  in  captivity,  the  writer  left  them  per- 
manently with  the  male  on  February  13.  Some  six  weeks  later,  or 
10  days  before  a  family  of  young  appeared,  the  gravid  condition  of 
one  female  was  plainly  apparent  and  this  one  was  then  segregated 
from  the  others.  A  few  days  later  it  was  noted  that  her  nest  was 
growing  in  size,  until  finally  it  was  more  than  twice  as  large  as  the 
single  nest  shared  by  the  male  and  the  other  female. 

On  April  7,  when  the  female  was  at  the  approximate  age  of  8l/£ 
months,  two  young  were  born.  An  endeavor  was  made  to  breed 
the  dam  on  the  day  following,  and  weekly  thereafter,  -but  unsuccess- 
fully. When  the  pair  was  placed  together  in  a  fresh  cage  the  fe- 
male was  totally  indifferent  to  the  decided  attentions  of  the  male; 
but  when  either  the  male  or  the  other  female  was  introduced  into 
the  cage  with  the  young — even  when  these  were  a  month  old — the 
mother  viciously  attacked  the  intruder  and  speedy  interference 
was  necessary. 

At  birth  the  young  were  indistinguishable  from  newly  born 
meadow  mice.  Four  days  later  there  was  a  distinct  suggestion  of 
fuzz  all  over  their  bodies  and  the  tails  were  conspicuously  blackish. 
By  the  fifteenth  day  they  were  completely  clothed  in  a  short  coat  of 
the  usual  coloration.  On  this  date  the  female  inflicted  on  the  writer 
her  first  severe  bite,  drawing  blood  from  the  finger,  and  thereafter 
was  so  belligerent  whenever  the  nest  was  invaded  that  no  more 
chances  were  taken,  but  she  was  poked  away  from  the  young  with 
the  aid  of  a  pencil  whenever  it  was  desired  to  examine  them.  What 
scant  hair  is  present  upon  the  tail  is  ochraceous — it  is  the  skin  of  the 
tail  that  is  black.  There  are  also  dusky  areas  of  skin  upon  the 
naked  ears,  the  entire  hind  feet,  and  about  the  ankles  of  the  fore- 
feet. 

Until  2  weeks  of  age  the  young  were  usually  firmly  attached  to  the 
nipples  of  the  dam  whenever  they  were  disturbed,  and  so  tenaciously 
did  they  retain  their  grip  that  it  was  with  a  deal  of  trouble  that  they 
could  be  detached.  They  dragged  along  after  the  female  when  she 
ran  about  the  cage  endeavoring  to  elude  capture,  and  if  one  of  them 
were  lifted,  it  would  pull  the  female  along  with  it  while  she  kicked 
about,  and  she  could  break  loose  only  by  grabbing  something  station- 
ary with  her  forefeet  and  pulling  vigorously.  After  the  age  of  2 
weeks  the  young  relinquished  this  habit. 


1926] 


LIFE    HISTORY   OF   PHENACOMYS  LONGICAUDUS 


49 


During  the  nestling  stage  it  was  noted  that  these  mice  have  a 
strong  propensity  to  remain  where  placed.  They  will  crawl  about 
over  one's  hand  after  the  eyes  have  opened,  and  at  this  stage  their 
clinging  ability  develops  rapidly,  but  until  well  grown  they  are 
willing  to  sit  quietly  in  the  pan  of  the  balance  while  being  weighed. 
None  were  detected  leaving  the  nest  until  they  were  29  days  old,  al- 
though they  could  have  accomplished  this  feat  merely  by  crawling 
over  a  wire  partition  1  inch  high.  All  this  is  in  marked  contrast 
with  the  actions  of  the  young  of  Peromyscus,  which  develop  decided 
restlessness,  at  least  when  disturbed,  well  before  the  eyes  have 
opened. 

That  sedentary  habits  in  the  young  of  Phenacomys  are  of  advan- 
tage to  the  species  is  easily  understood,  and  this  has  undoubtedly 
been  developed  by  its  arboreal  habitat.  If  they  were  prone  to  be- 
come agile  and  inquisitive  at  an  early  age,  large  numbers  would  most 
assuredly  meet  with  disaster  by  falling  from  the  nests. 

On  the  assumption  that  the  mice  were  born  April  7,  although  it 
is  possible  that  they  may  have  put  in  an  appearance  late  the  pre- 
vious evening,  the  eyes  had  fully  opened  at  the  age  of  19  days,  on 
April  26.  The  writer  is  familiar  with  the  timed  stages  of  growth 
of  very  few  small  mammals,  but  the  length  of  the  sightless  period  in 
this  species  seems  to  be  remarkable.  Young  meadow  mice,  for  in- 
stance, are  able  to  shift  for  themselves  at  less  than  3  weeks  of  age, 
and  even  such  a  large  mammal  as  the  domestic  rabbit  is  more  pre- 
cocious as  regards  duration  of  the  period  of  sightlessness,  although 
the  latter  case  is  not  strictly  comparable. 

An  attempt  was  made  to  weigh  the  young  mice  every  second  day 
at  the  same  hour,  but  continuity  was  unavoidably  interrupted  on 
one  occasion  for  three  days.  There  was  no  means  of  distinguishing 
between  the  two  individuals,  nor  could  their  sex  be  determined.  In 
fact,  one  can  hardly  be  positive  regarding  sex  until  an  individual 
has  passed  the  juvenile  stage.  At  all  ages  the  males  are  far  less 
developed  sexually  than  in  Microtus,  and  it  is  even  very  often  diffi- 
cult to  be  positive  of  the  sex  of  an  adult  at  first  glance. 

Table  1. — Increasing  iceights,  in  grams,  from  date  of  birth  of.  two  juveniles  of 
Phenacomys  longicaudus,  bom  in  captivity 


Date 


Apr.  7_. 
Apr.  9.. 
Apr.  11 
Apr.  15 
Apr.  16 
Apr.  18 
Apr.  20 
Apr.  22 


Weights 

No.  1 

No.  2 

3.1 

3.1 

3.1 

3.4 

3.1 

3.2 

4.1 

4.6    ! 

4.5 

4.8 

5.0 

5.3 

5.4 

5.5 

6.1 

6.2 

Date 


Apr.  24 
Apr.  26 
Apr.  28 
Apr.  30 
May  2. 
May  4. 
May  6. 


Weights 


No.  1         No.  2 


6.4 
7.0 
7.4 
8.2 
9.2 
10.5 
11.2 


6.5 
7.0 
7.4 
8.7 
9.7 
10.7 
11.5 


It  is  seen  from  the  data  in  Table  1  that  the  young  mice  did 
not  double  their  natal  weights  until  16  days  old,  nor  triple  these 
weights  until  more  than  25  days  old.  After  three  weeks  the  gain 
was  rapid,  and  this  fact  suggests  that  this  was  perhaps  the  time 
when  they  began  eating  an  appreciable  quantity  of  solid  food  from 


50  NORTH   AMERICAN  FAUNA  [No.  48 

the  needles  kept  on  or  in  the  nest  by  the  parent.  At  the  age  of 
27  days  there  occurred  a  sudden  change  in  the  attitude  of  these 
youngsters,  for  they  then  refused  to  remain  in-  one  place  but  would 
scramble  out  of  the  pan  of  the  balance,  out  of  a  cardboard  box 
with  sides  2  inches  high,  and  would  even  leap  to  the  floor  and 
make  off  with  almost  the  agility  and  speed  of  an  adult.  They 
were  not  detected  out  of  the  nest,  however,  until  the  29th  daj7, 
although  they  may  well  have  ventured  on  short  excursions  earlier. 
At  this  age  they  at  once  began  spending  much  time  out  of  the  nest, 
and  ate  fir  needles  freely. 

The  conclusion  reached  from  observation  of  the  growth  of  the 
young  is  that  development  is  remarkably  slow,  a  state  of  affairs 
doubtless  brought  about  by  habits  that  are  largely  arboreal. 

On  one  occasion  in  one  nest  were  a  single  juvenile  about  3  weeks 
old  and  another  immature  mouse,  perhaps  two-thirds  grown.  The 
latter  had  nonfunctional  mammary  glands  and  was  too  undeveloped 
to  have  borne  young  at  any  rate.  This  is  the  only  certain  evi- 
dence at  hand  that  one  litter  may  remain  with  the  adult  female 
until  after  another  has  arrived.  In  this  case  it  was  probably  a 
state  of  affairs  tolerated  by  the  mother  only  because  both  litters 
were  small  and  hence  did  not  crowd  the  nest.  Young  larger  than 
about  half  grown  are  seldom  found  with  an  adult  female;  but 
there  are  no  data  at  hand  regarding  the  movements  of  immatures 
immediately  after  they  abandon  the  maternal  abode.  The  pos- 
sibility must  be  considered,  however,  that  they  then  live  a  partly 
terrestrial  existence,  as  seems  to  be  the  case  with  adult  males,  until 
the  approach  of  sexual  maturity  urges  the  females  into  the  trees. 

No  other  microtine  is  known  to  have  litters  of  such  small  size, 
and  this  is  a  further  indication  that  the  adverse  life  conditions 
encountered  by  the  species  are  not  severe.  Although  more  than 
one  family  of  young  a  year  is  probably  the  rule,  these  undoubtedly 
appear  with  far  less  frequency  than  is  the  case  with  meadow  mice. 

FOOD 

Although  Phenacomys  longicaudus  is  not  highly  specialized  in 
structure  it  is  exceedingly  so  in  habits.  The  normal  food  apparently 
consists  solely  of  the  fleshy  part  of  the  needles  and  the  bark  from  the 
tenderest  growth  of  the  twigs  of  fir  trees,  and  in  captivity  it  will 
accept  no  items  other  than  coniferous  even  though  reduced  to  dire 
straits  of  hunger.  Experiments  with  various  fare,  including  cereals, 
fruits,  vegetables,  grasses,  and  twigs  of  deciduous  trees  and  of  red- 
wood, failed  to  show  that  any  of  these  was  sampled  or  even  noticed. 

The  mouse's  manner  of  obtaining  food  is  to  cut  from  the  tree 
tender,  terminal  twigs  3  to  8  inches  long.  Some  of  these  are  in- 
advertently dropped  to  the  ground,  many  are  probably  eaten  upon 
the  spot,  and  some  are  carried  to  the  nest,  if  near  by,  for  more 
leisurely  consumption.  A  supply  is  kept  on  top  of  the  nest  as  a 
diurnal  larder,  and  the  refuse  from  it  constitutes  the  building  ma- 
terial. There  is  no  unnecessary  waste  in  eating,  and  every  needle 
is  eaten  from  a  twig  one  after  another,  no  portion  being  dropped 
until  the  edible  part  is  consumed. 

The  mouse  ordinarily  approaches  a  twig,  already  cut,  and  bites 
off  a  needle  at  the  very  base.    Occasionally  with  one  foot,  but  more 


1926] 


LIFE   HISTORY   OF   PHENACOMYS  LONGICAUDUS  51 


often  with  two,  it  grasps  the  needle  as  we  would  an  ear  of  corn. 
Sometimes  this  is  run  between  the  lips  with  a  single  motion  so 
rapidly  as  to  defy  analysis  of  the  movements.  Again,  more  leisurely 
tactics  are  employed  and  one  may  clearly  note  the  animal  neatly  split 
off  one  side  of  the  needle  and  eat  the  smaller  portion  from  one 
"  hand,"  while  holding  the  remainder  with  the  other.  The  "  midrib  " 
is  then  split  off  and  discarded,  and  the  remainder  quickly  consumed. 
Thus,  from  each  needle  there  is  discarded  a  medullary  core  in  the 
form  of  a  slender,  greenish  thread,  and  it  is  of  these  that  the  nests  are 
wholly  composed.  Taylor  (1915b)  states  that  "  all  the  fleshy  sub- 
stance of  the  needle  including  the  vascular  bundles  is  eaten  away  by 
the  tree  mouse,  the  only  portions  left  being  the  two  resin  ducts  which 
traverse  the  entire  length  of  each  needle."  Such  may  possibly  be  the 
case  at  times  or  with  certain  mice,  but  the  writer's  investigations  of 
the  subject,  including  microscopical  examinations  of  cross  sections 
of  the  needles  and  of  a  reasonable  quantity  of  the  threadlike  refuse, 
cleanly  indicate  that  the  lateral  resin  ducts  (a,  in  pi.  7,  fig.  3)  are 
eaten  with  the  remainder  of  the  fleshy  portion,  and  that  the  tougher, 
central,  vascular  bundles  (b),  together  with  a  small  but  varying 
quantity  of  surrounding  tissue,  constitute  the  portion  discarded. 

A  great  deal  of  this  provender  is  consumed,  probably  because  it 
is  rather  low  in  nutrient  value.  For  the  same  reason,  digestion  must 
be  rapid  in  order  to  care  for  the  quantity.  Food  is  invariably 
available  within  a  few  inches,  or  at  most  a  few  feet,  of  the  arboreal 
nests  of  the  mice*  Like  all  mammals  whose  food  is  always  to  be 
had  in  quantity,  they  have  lost  all  ability  to  withstand  hunger  and 
will  exhibit  signs  of  distress  within  an  hour  or  two  after  having 
been  deprived  of  food.  It  is  for  this  reason  that  a  supply  of  twigs 
is  piled  upon  the  nest  each  night,  so  that  the  inmate  shall  not  want 
for  food  during  the  ensuing  day. 

The  writer  has  ascertained,  by  an  exact  arrangement  of  twigs 
upon  a  nest  during  the  morning  and  subsequent  examinations 
throughout  the  day,  that  the  mice  draw  from  their  supply  at 
frequent  intervals,  pulling  the  twigs  down  into  the  nest  one  by  one 
as  desired.  The  quantity  so  stored  varies  from  3  or  4  to  12  or  15 
twigs,  and  they  are  either  piled  in  haphazard  fashion  upon  the  nest, 
as  is  usual,  or  a  part  of  them,  in  a  tight  little  bundle,  is  pulled 
within  one  of  the  entrances.  Very  rarely  are  any  of  these  twigs 
wilted  to  the  slightest  degree,  and  as  a.  surplus  above  the  needs  of 
the  animal  must  often  be  gathered,  all  but  the  freshest  twigs  must 
be  dropped  from  the  nest. 

The  following  feeding  experiments  were  conducted  with  a  sub- 
adult  female  about  four-fifths  grown  and  weighing  20  grams,  se- 
lected in  preference  to  older  subjects  because  she  was  by  all  odds 
superior  in  tractability  and  because  the  only  other  fully  adult  ani- 
mal available  at  the  time  was  with  a  litter  of  young : 

Three  terminal  twigs  were  selected,  each  6  inches  long,  from  a 
large  branch  of  a  Douglas  fir,  and  a  count  made  of  the  needles 
upon  them.  The  results  were  209,  213,  and  218,  for  which  a  con- 
servative average  of  210  is  taken.  It  is  important  here  to  note  that 
the  number  of  needles  upon  such  a  twig  varies  widely  for  different 
trees,  and  according  to  the  light  received  by  the  portion  of  the  tree 
from  which  taken.    All  twigs  used  in  these  experiments  were  from 


52  NORTH  AMERICAN   FAUNA  [No.  48 

the  same  part  of  this  one  fir  and  were  strictly  comparable  in  condi- 
tion and  size.  The  method  followed  was  to  remove  all  food  from 
the  cage  and  then  to  introduce  fresh  twigs  of  an  aggregate  number 
of  inches,  carefully  measured.  After  a  certain  number  of  hours  the 
quantity  remaining  was  measured  and  discarded,  and  a  fresh  supply 
of  measured  twigs  introduced.  Tests  were  made  upon  this  indi- 
vidual, then  and  subsequently  in  good  health,  for  two  successive 
periods  of  24  hours  each,  with  the  results  given  in  Table  2. 

Table  2. — Lengths,  in  inches,  of  tioigs  of  Douglas  fir  (Pseudotsuga  taxifolia) 
consumed  by  one  red  tree  mouse  in  comparable  periods  of  successive  dags 


Period 

/ 

First  day 

Second  day 

12  m.  to  6  p.  m 

Inches 
15.75 
34.00 
16.25 

Inches 
17.25 

40.00 

6  a.  m.  to  12  m 

13.50 

Total  (24  hours) 

.66.00 

70.75 

Of  the  first  lot,  52  per  cent,  and  of  the  second,  57  per  cent  of  the 
totals  was  eaten  from  6  p.  m.  to  6  a.  m.,  and  at  least  some  of  this 
slight  disparity  is  probably  due  to  the  fact  that  as  the  animals  take 
nearly  all  of  their  exercise  during  the  hours  of  darkness,  they  would 
then  be  more  voracious.  A  total  of  66  inches  of  these  twigs  sup- 
ports about  2,310  needles,  weighing  17.5  grams ;  and  of  70.75  inches, 
2,476  needles,  weighing  19  grams  (see  pi.  7,  fig.  2).  It  is  thus  seen 
that  a  subadult  mouse  eats  about  100  fir  needles  an  hour,  day  and 
night.  In  the  case  of  an  old  female  with  a  family,  this  number 
must  be  augmented  considerably.  The  fresh  food  refuse  could  not 
be  weighed,  for  it  is  indistinguishable  from  the  older,  more  desic- 
cated material  of  which  the  nest  is  formed,  and  if  deprived  of  a 
nest  the  mouse  would  soon  have  fretted  itself  to  death.  It  is  be- 
lieved, however,  that  the  weight  of  this  refuse  is  fully  offset  by  the 
quantity  of  bark  eaten  from  the  twig  tips,  of  which  no  account 
was  taken  during  the  investigation.  Hence,  it  seems  probable  that 
the  normal  tree  mouse  will  consume  about  nine-tenths  of  its  weight 
in  food  every  24  hours.  For  the  sake  of  comparison,  it  may  be  well 
to  mention  that  a  larger  animal — an  apparently  full-grown  male — 
after  having  been  in  captivity  for  several  weeks,  weighed  27  grams. 
Additional  weights  of  animals  fully  grown  at  the  time  of  capture  are 
not  available. 

Three  adult  or  subadult  mice,  which  the  writer  kept  in  captivity 
for  some  time,  showed  a  marked  preference  for  needles  of  Douglas  fir 
to  those  of  grand  fir.  Tests  were  made  by  placing  equal  portions 
of  twigs  of  the  two  species  with  an  animal,  and  invariably  the 
Douglas  fir  was  consumed  practically  completely  before  the  other 
would  be  more  than  nibbled ;  but  the  grand  fir  would  then  be  eaten 
readily.  This  preference  was  really  so  marked  that  it  was  greatly 
puzzling,  for  individuals  were  found  in  segregated  grand  firs  that 
could  obtain  other  provender  only  by  crossing  open  patches  of 
ground,  and  grand-fir  twigs  were  found  upon  the  nests.  The  writer 
is  therefore  forced  to  accept  the  hypothesis,  since  substantiated 


1926]  LIFE   HISTORY   OF   PHENACOMYS  LONGICAUDUS  53 

through  experiments  with  captive  mice,  that  the  tastes  of  individual 
mice  differ  to  some  extent,  and  that  all  those  kept  alive  in  captivity 
happened  to  prefer  the  Douglas-fir  needles. 

Clay  (Taylor,  1915b,  p.  145)  took  several  mice  from  nests  in  Sitka 
spruces,  as  previously  mentioned,  and  states  that  these  trees  were 
so  close  to  Douglas  firs  that  the  food  supply  may  have  been  obtained 
from  the  latter.  The  mice  may  or  may  not  have  been  feeding  on 
spruce  needles. 

Clay  (Taylor,  1915b,  p.  145)  also  found,  near  Maplecreek,  Hum- 
boldt County,  certain  trees  that  had  bushed  out  as  if  the  tops 
had  previously  been  killed,  and  that  invariably  contained  the  nest  of 
a  tree  mouse  and  furrowlike  marks  at  the  points  of  damage,  a  con- 
dition inferred  to  have  been  caused  by  Pkenacomys.  Careful  search 
was  made  for  any  such  condition  near  Carlotta,  and  although  two 
nests  were  located  at  points  where  the  tops  of  the  trees  had  long  since 
been  broken  in  some  manner,  and  new  leaders  had  consequently  started, 
no  sign  was  found  of  recent  injury  to  the  bark.  There  is  a  possibility 
that  the  damage  discovered  by  Clay  was  caused  by  gray  squirrels 
or  rodents  other  than  the  mice,  yet  it  seems  not  unlikely  that  the  mice 
of  that  particular  area  may  have  developed  a  preference  for  feeding 
largely  on  bark. 

That  such  change  in  diet  may  be  brought  about  seems  to  be  proved 
by  captive  tree  mice  which  the  writer  has  had  in  southern  California. 
At  first,  clippings  from  every  available  species  of  conifer  grown  in 
Pasadena  as  ornamentals  were  offered  to  them,  including  3  species  of 
pine,  2  of  spruce,  1  fir,  3  of  cypress,  a  redwood,  and  several  cedars. 
The  needles  of  Abies  nordmaimiana — a  native  fir  of  the  Caucasus — 
were  eaten  sparingly,  but  seemed  not  to  be  greatly  relished.  All  the 
others  were  utterly  ignored  except  the  deodar  (Cednts  deodara),  and 
this  was  adopted  as  a  permanent  and  exclusive  diet,  one  upon  which 
the  mice  will  evidently  thrive  indefinitely.  Several  months  subse- 
quently, however,  the  same  mice  welcomed  twigs  from  the  exotic  fir 
and  ate  them  to  a  limited  extent  only  in  preference  to  other  fare, 
but  they  soon  grew  tired  of  them  and  returned  to  the  deodar.  The 
needles  of  the  latter  grow  in  clusters  of  two  dozen.  The  mice  bite  off 
one  cluster  at  a  time,  and  consume  several  needles  and  drop  the 
remainder,  to  which  they  may  or  may  not  return.  Hence,  there  is 
much  waste  and  it  is  impossible  to  estimate  the  quantity  eaten.  The 
needles,  unlike  those  of  Douglas  fir,  have  no  central  vascular  bundles, 
but  are  roughly  circular  in  transverse  section,  with  succulent  medul- 
lary tissue  and  a  somewhat  more  fibrous  cortex.  There  is  no  part  of 
this  discarded,  but  the  mice  munch  the  needles,  held  often  in  one  paw, 
as  we  would  stalks  of  celery.  The  bark,  however,  seems  to  be  as 
much  favored  as  the  needles  and  it  is  stripped  clean  even  from  year- 
old  twigs  as  large  as  a  pencil.  It  is  loosened  chiefly  with  the  teeth, 
but  the  forefeet  sometimes  aid  in  pulling  a  section  from  a  twig. 

A  male  which  Wilder  sent  the  writer  during  January  would  have 
nothing  to  do  with  the  deodar,  even  after  it  had  finally  been  reduced 
to  sore  straits  of  hunger,  and  it  was  necessary  for  weeks  to  keep 
it  supplied  with  twigs  of  the  exotic  fir  previously  mentioned,  with 
the  needles  of  which  it  seemed  well  content,  though  none  of  the 
bark  was  consumed.     When  these  are  eaten,  the  center  of  the  needle 


54  NORTH   AMERICAN   FAUNA  [No.  48 

is  bitten  through,  rapidly  split  the  entire  length,  and  the  medullary 
core  then  skilfully  taken  from. the  half  to  which  it  was  attached. 
This  is  dropped  and  the  two  remaining  sections  are  munched. 
Finally,  after  persistent  effort,  this  mouse  became  reconciled  to  the 
deodar. 

After  the  mice  in  captivity  had  thrived  for  two  months  upon  a 
diet  of  deodar,  twigs  of  the  big-cone  spruce  (Pseudotsuga  macro- 
carpa)  from  the  San  Bernardino  Mountains  of  southern  California 
were  offered.  They  did  not  prove  acceptable  as  long  as  deodar 
was  available,  but  when  the  latter  was  removed  the  spruce  needles 
were  at  once  eaten  in  the  usual  manner,  while  the  bark  was  left 
untouched.  When  these  facts  are  taken  into  consideration,  together 
with  the  indicated  probability  that  individual  wild  mice  may  prefer 
either  Douglas  or  grand  fir,  it  does  not  seem  at  all  improbable  that 
the  tree  mice  of  certain  districts  may  prove  regularly  to  feed  upon 
bark.  The  resulting  damage,  however,  is  at  present  entirely  too 
restricted  and  unimportant  to  merit  economic  consideration. 

A  study  of  the  species  at  once  indicates  that  the  tree  mice  have  no 
regular  water  supply,  and  the  trees  they  inhabit  may,  in  fact,  be  at 
any  distance  from  water.  That  they  require  very  little  moisture 
is  indicated  by  the  fact  that  the  quantity  of  urine  voided  is  very 
small.  Cages  in  which  the  animals  were  kept  remained  fresh  and 
clean  almost  indefinitely  without  being  emptied,  a  state  of  affairs 
which  would  not  be  possible  were  Microtis  the  captive,  without  a 
liberal  supply  of  some  absorbent.  In  much  of  the  range  of  the  tree 
mice,  heavy  fogs  are  very  frequent  and  the  consequent  condensation 
upon  the  foliage  is  probably  used  to  some  extent  at  the  season  when 
rains  do  not  occur.  Such  free  water  does  not  seem  to  be  absolutely 
necessary,  however,  and  it  is  believed  that  the  species  can  at  all  times 
obtain  enough  water  from  the  succulent  new  growth  of  the  trees  to 
meet  their  moderate  needs. 

Mice  kept  in  captivity  by  the  writer  were  usually  fed  perfectly  dry 
twigs,  but  about  once  every  week  these  were  dipped  in  water,  in  case 
such  might  be  relished.  For  a  long  time  there  was  no  evidence  that 
this  attention  was  appreciated;  but  once,  during  an  especially  hot 
spell  in  September,  and  on  a  number  of  occasions  thereafter,  an  indi- 
vidual was  seen  to  lap  a  few  drops  from  the  needles,  and  one  would 
occasionally  lick  a  wet  finger.  The  writer  did  not  succeed  in  get- 
ting the  mice  to  drink  water  from  an  open  container,  but  taking  his 
cue  from  seeing  them  lick  wet  needles,  he  contrived  a  small,  inverted 
bottle  from  which  the  mice  could  get  water  by  lapping  at  the  end  of 
a  slender  glass  tube.  To  this  they  had  regular  though  sparing  re- 
course. As  no  evaporation  from  this  receptacle  could  take  place  and 
there  was  no  waste,  the  quantity  of  water  used  by  the  mice  was 
readily  ascertained,  and  found  to  average  6  grams  a  day  for  each 
mouse  (for  two  subadults  weighing  17  and  19  grams).  This  is  de- 
cidedly less  than  the  similar  demands  of  Microtus,  as  observed  in  the 
case  of  captives. 

This  evidence  substantiates  the  deductions  of  Bailey  (1923)  that 
some  rodents  obtain  sufficient  moisture  from  their  regular  food  to 
carry  on  normal  bodily  functions  without  recourse  to  a  supply  of 
free  water,  but  that  the  same  animals  may  use  liberal  quantities  of 
water  when  this  is  to  be  had.    Nineteen  grams  of  fresh  needles  from 


1926]  LIFE   HISTORY   OF   PHENACOMYS  LONGICAUDUS  55 

the  exotic  fir  in  Pasadena — probably  comparable  in  texture  to  those 
of  the  Douglas  fir — were  air-driecl  for  a  week  and  then  heated  in  an 
oven  at  a  temperature  of  250°  F.  for  one  hour.  They  were  then  found 
to  weigh  8.7  grams.  Not  all  of  this  loss  in  weight  can  be  due  to 
evaporation  of  moisture  content,  for  the  volatile  oils  were  also  driven 
off  during  the  heating  process.  The  evidence  shows,  then,  that  a 
nearly  full-grown  tree  mouse  probably  requires  a  minimum  moisture 
supply  of  about  11  grams  a  day  to  maintain  itself  in  good  health, 
but  that  as  much  as  17  grains  a  day  may  be  used  when  available. 

The  feces  were  abundant,  to  conform  to  the  large  bulk  of  food 
eaten,  but  rather  dry  in  consistency. 

Although  the  species  may  eat  other  substances  upon  occasion,  as 
insects  or  seeds,  the  writer  very  much  doubts  that  they  touch  any- 
thing under  normal  conditions  other  than  the  needles  and  tender 
bark  of  their  favorite  trees. 

Efforts  of  the  writer  to  trap  these  mice  have  never  been  successful, 
with  the  exception  of  two  individuals  caught  by  setting  unbaited 
mouse  traps  upon  the  limbs  of  trees  beneath  the  nests.  As  it  is 
probably  impossible  to  offer  any  bait  that  would  attract  the  mice,  the 
specimens  taken  in  traps  would  be  merely  individuals  which  hap- 
pened to  run  across  the  trigger. 

HABITS 

From  lack  of  observation  in  a  number  of  diverse  localities,  no 
statements  regarding  the  preferences  of  Phenacomys  longicaudus  as 
to  slope  exposure  can  at  present  carry  much  weight.  As  mentioned, 
the  preference  within  the  area  studied  appears  to  be  for  the  south 
side  of  the  tree  but  this  may  not  apply  to  other  parts  of  the  range. 
Taylor  (1915b)  found  the  mice  living  in  grand  firs  at  Mendocino 
City;  but  they  have  usually  been  recorded  from  the  Douglas  firs. 

Among  the  specimens  heretofore  collected  there  is  a  great  pre- 
ponderance of  females,  the  exact  ratio  between  the  sexes  being  un- 
determined. Adult  males  especially  are  greatly  in  the  minority,  and 
Taylor  (1915a,  1915b)  is  of  the  opinion  that  this  numerical  discrep- 
ancy is  the  actual  state  of  affairs  at  birth.  In  perhaps  80  per  cent  of 
the  microtines  which  the  writer  has  examined,  females  have  out- 
numbered males,  and  it  seems  certain  that  any  such  possible  natal 
difference  between  the  sexes  is  not  substantially  greater  in  the 
present  species  than  among  other  forms  of  this  subfamily.  One  fact 
accounting  for  the  greater  number  of  females  so  far  captured  is  that 
the  males  often  elude  the  collector,  because  they  seem  to  be  more 
wary  and  their  arboreal  nests  are  both  smaller  and  better  concealed. 
Another  is  that  the  life  habits  of  adult  males  are  apparently  radically 
different  from  those  of  adult  females. 

Startling  as  this  last  assertion  may  seem,  recent  discoveries  by 
Wilder  point  to  this  conclusion.  Several  years  ago  he  noted  a  hole 
near  the  base  of  a  decayed  stump  beneath  a  live  fir,  and  digging 
into  it,  discovered  a  tree  mouse  in  a  small  nest  of  the  usual  com- 
position; but  the  animal  escaped  and  he  was  unable  to  ascertain  its 
sex.  He  supposed  at  the  time  that  it  had  been  blown  or  frightened 
from  the  tree  above  and  had  sought  temporary  shelter  near  the 
ground.  On  another  occasion  he  was  clearing  away  debris  prepara- 
tory to  spreading  his  blankets  when  a  red  tree  mouse  ran  from 


56  NORTH  AMERICAN  FAUNA  [No.  48 

between  his  hands.  It  proved  to  be  a  male.  He  did  not  know 
whether  it  had  been  beneath  the  rubbish  or  had  fallen  from  the 
trees  above. 

Late  one  fall  a  red  mouse  took  up  residence  in  a  woodpile  at 
the  foot  of  a  fir  tree,  and  remained  there  all  winter  in  a  nest  of  ordi- 
nary construction.  This  was  also  a  male,  but  the  writer  was  unaware 
until  recently  of  the  sex  of  these  specimens  captured  on  the  ground. 
Some  time  ago  Wilder's  cat  brought  in  another  male.  Under  dif- 
ferent dates  he  has  written  that  early  in  February,  1924,  "while 
working  back  of  the  barn  I  noticed  the  pup  digging  in  some  rubbish 
at  the  foot  of  one  of  the  trees  where  you  camped  last  summer,  and 
soon  a  red  mouse  ran  out.  I  caught  it — a  male."  Again,  "February 
16,  while  cutting  brush  out  back  of  the  garden  I  noted  our  '  mouse- 
dale '  barking  and  digging  at  the  base  of  a  fir  tree  and  upon  going 
to  him,  found  that  he  was  following  a  shallow  burrow,  like  that  of 
a  meadow  mouse,  and  he  soon  dug  up  the  nest  of  a  red  mouse — the 
usual  ball  of  midribs  of  fir  needles.  Following  the  burrow  about  a 
foot  farther  he  threw  out  a  fine,  big  red  mouse,  a  male.  It  was  plain 
that  this  mouse  was  regularly  living  in  the  burrow."  Under  date  of 
March  9,  1924,  he  wrote,  "The  dog  located  a  red  mouse  under  the 
remains  of  an  old  rat  nest.  The  pup  followed  it  along  a  runway  in 
the  loose  rubbish  to  a  hole  under  the  tree,  where  he  could  dig  no 
farther,  but  was  close  to  the  mouse  and  much  excited." 

The  facts  seem  to  indicate  that  most  males,  after  attaining  sexual 
maturity,  spend  the  major  portion  of  their  time  below  ground,  or 
at  least  beneath  rubbish,  foraging  among  the  branches  of  the 
trees  at  night.  At  certain  times,  which  probably  depend  upon  sexual 
impulse,  they  do  construct  small,  temporary,  or  arboreal  nests. 

In  hunting  for  the  mice  one  searches  for  their  nests,  which  often 
are  visible  from  the  ground,  especially  when  situated  in  trees  of 
moderate  size.  Green  fir  twigs  on  the  nest  are  a  sure  sign  that  mice 
are  in  either  that  tree  or  an  adjoining  one,  usually  in  the  nest  itself, 
and  a  further  indication  of  their  presence  is  the  finding  of  a  moderate 
number  of  short,  green  twigs  on  the  ground  beneath.  In  the  case  of 
lofty  or  especially  dense  trees,  the  presence  of  such  twigs  on  the 
ground  is  of  use  to  the  investigator.  Of  less  practical  value  is  the 
presence  of  droppings,  for  the  surface  is  rarely  of  such  nature  as  to 
permit  these  to  be  found  without  difficulty.  Such  evidence  is  usually 
present  each  morning,  however,  upon  table  and  sleeping  bag  when 
one  is  encamped  beneath  a  tree  in  which  they  live. 

Any  opinion  regarding  the  abundance  of  tree  mice  in  a  given  area 
is  little  more  than  a  haphazard  guess.  A  densely  forested  area,  of 
course,  could  support  a  greater  population  than  thinly  wooded  slopes, 
as  at  Carlotta.  An  estimate  of  the  number  of  tree  mice  to  the  acre 
or  square  mile  would  be  utterly  without  value  under  present  circum- 
stances. 

One  who  is  familiar  with  the  tree  mouse  is  prone  to  refer  to  an 
aggregation  of  individuals  in  any  well-defined  area  as  a  colony; 
but  the  animals  really  have  no  colonizing  tendency  and  are  not  at  all 
sociable  in  the  strict  sense  of  the  word.  Their  gradual  dispersal 
over  a  territory,  usually  of  limited  extent,  in  which  they  experience 
favorable  conditions  is  in  such  marked  contrast  to  a  closely  adjacent 
territory,  which  may  be  unsuited  for  lack  of  proper  trees  or  some 


1926]  LIFE   HISTORY   OF   PHENACOMYS  LONGICAUDUS  57 

other  reason,  that  tree-mouse  populations  seem  upon  casual  examina- 
tion, to  be  colonial  in  character.  In  reality  the  mice  are  of  a  mark- 
edly solitary  disposition,  the  females  usually  being  truculent  and  in- 
tolerant of  other  individuals. 

The  only  evidence  indicating  that  two  adults  may  occupy  the  same 
nest  is  that  offered  by  Taylor  (1915b,  p.  153),  who  states  that  Clay 
once  caught  a  female  in  a  nest  from  which  another  mouse,  assumed 
to  be  a  male,  escaped.  As  the  writer  has  found  a  large,  immature 
female,  with  no  signs  of  having  suckled,  in  the  nest  with  a  young 
mouse,  the  mother  of  which  undoubtedly  had  been  present  but  had 
escaped,  he  considers  that  Taylor's  example  was  of  this  kind  and 
that  a  female  will  not  tolerate  another  adult  even  in  the  same  tree 
with  her,  if  this  be  small,  except  during  the  mating  period.  It  is 
probable,  however,  that  large  trees  may  often  permanently  harbor 
more  than  a  single  individual.  At  times,  which  it  is  thought  last 
only  during  the  breeding  period,  the  small,  evidently  temporary  nest 
of  a  male  may  be  found  several  feet  directly  above  the  more  sub- 
stantial family  home  of  its  mate,  but  each  animal  occupies  its  re- 
spective domicile  during  the  day. 

The  present  species  is  nocturnal,  but  has  no  aversion  to  moderate 
light.  The  urge  for  eating  very  frequently  necessitates  diurnal  ac- 
tivity within  the  nest,  and  to  a  certain  extent  upon  it,  for  captive 
individuals  would  habitually  crawl  upon  the  pile  of  food  twigs  for 
the  purpose  of  stripping  off  a  supply  of  fodder  at  all  hours  of  the 
day.  The  writer  seriously  doubts,  however,  whether  mice  if  undis- 
turbed ever  voluntarily  leave  the  nests  during  daylight. 

A  factor  that  probably  largely  inhibits  diurnal  activity  is  the 
lack  of  suitable  cover  upon  the  branches.  To  survive  the  dangers 
of  foraging  by  daylight  an  arboreal  rodent  would  have  to  be  ex- 
tremely quick  and  decisive  in  its  actions,  which  tree  mice  are  not. 
The  most  tractable  and  confiding  of  the  individuals  that  the  writer 
has  kept  in  captivity  was  not  at  all  averse  at  any  hour  to  exercising 
on  its  wheel  in  his  presence,  in  this  respect  being  entirely  comparable 
to  the  average  meadow  mouse — an  animal  normally  living  amid 
ample  cover  and  showing  a  propensity  for  being  fully  as  diurnal  as 
nocturnal.  The  average  tree  mouse,  however,  is  relatively  inactive 
as  long  as  there  is  light,  either  natural  or  artificial,  even  when  raised 
from  birth  in  captivity. 

OTHER  BEHAVIOR 

As  anyone  who  has  kept  them  in  captivity  knows,  even  mice  possess 
individuality  to  a  surprising  extent,  and  it  is  difficult  to  predict  just 
what  any  example  will  do  under  a  given  set  of  circumstances.  Some 
tree  mice  will  leap  into  space  before  the  nest  is  touched,  apparently 
regardless  of  where  they  may  land;  others  will  tarry  until  the 
structure  has  been  half  demolished;  a  few  will  hide  until  there 
remains  only  a  handful  of  material  and  one  has  already  pronounced 
the  nest  uninhabited.  Those  individuals  which  leap  boldly  from  the 
nest  are  somewhat  in  the  minority,  the  remainder  preferring  to  slip 
out  and  away  at  the  most  unexpected  time  and  spot,  run  or  leap  a 
short  distance  and  then  remain  motionless  upon  the  chance  of  escap- 
ing observation.    At  this  method  of  escape  the  mice  are  experts,  and 


58  NORTH   AMERICAN   FAUNA  [No.  48 

if  more  than  an  occasional  specimen  is  to  be  secured  it  is  necessary 
to  have  a  companion  on  the  ground  beneath. 

Having  sprung  to  a  slender  dead  twig  several  feet  below  the  nest, 
the  animal  will  usually  remain  motionless  for  a  few  moments.  On 
being  approached  it  will  teeter  and  balance,  endeavoring  to  crawl  to 
an  adjoining  twig  several  feet  below,  which  it  will  perhaps  clutch 
with  a  single  foot.  If  it  has  the  chance  to  get  among  the  denser 
needle  growth,  it  will  try  to  escape  through  this;  but  if  jarred  to 
the  ground  it  will  make  off  at  a  poor  gait,  which  enables  one  to  catch 
it  with  ease. 

Some  individuals  will  not  bite  even  while  being  seized,  whereas 
others,  especially  males  and  old  females,  .may  struggle  and  are  very 
ready  to  nip  the  captor.  After  being  "  cuddled  "  in  the  hands  for  a 
few  moments  some  will  quiet  down  and  run  over  one's  arms  without 
making  any  determined  efforts  at  escape,  while  others  leap  to  the 
ground  repeatedly. 

Some  will  run  over  the  surface  of  the  ground,  taking  no  notice 
of  holes  beneath  logs  and  rubbish ;  others  slip  into  the  first  cranny 
they  come  upon.  When  placed  upon  the  trunk  of  a  tree  they 
"  shin  "  or  hitch  themselves  upward  as  would  a  boy,  often  making 
not  more  than  a  couple  of  feet  in  three  seconds,  with  frequent  short 
pauses  for  a  look  about. 

Although  tree  mice  are  gifted  with  a  certain  sort  of  acrobatic 
agility,  one  is  continually  impressed  with  their  slowness,  clumsiness, 
and  apparent  helplessness,  and  feels  that  the  average  white-footed 
mouse  {Peromyscus)  or  chipmunk  (Eutamias)  would  be  far  quicker 
and  more  agile  in  any  situation  except  upon  the  slenderest  twigs. 
The  chief  ground  on  which  to  attribute  to  Phenacomys  an  arboreal 
expertness  seems  to  be  its  cautiousness  and  the  sure-footed  manner 
with  which  it  grasps  with  widespread  toes  all  objects.  One  really 
can  not  escape  the  conviction  that  the  species  has  survived,  not  be- 
cause of  equipment  that  is  efficient  for  coping  with  its  environment, 
but  rather  for  the  reason  that  it  has  occupied  a  habitat  in  which  it 
encounters  little  competition,  and  hence,  that  it  has  been  enabled  to 
survive  even  while  its  agility  and  speed  have  not  improved. 

It  is  certain  that  even  the  females  descend  to  the  ground  upon 
occasion,  but  it  is  doubtful  whether  the  average  individual  of  this 
sex  does  so  very  often. 

So  far  as  the  writer  is  aware,  no  other  naturalist  has  succeeded 
in  keeping  Phenacomys  longicaudus  long  in  captivity.  In  all  other 
attempts,  including  his  own  early  ones,  the  animals  died  within 
a  day  or  two.  It  is  thought  that  this  has  been  due  to  intolerance  of 
the  species  for  all  conditions  except  such  as  closely  approximate  those 
to  which  it  is  accustomed,  and  to  misguided  efforts  of  the  captors  to 
furnish  warm  nests  of  cotton.  If  the  regular  nesting  material  be 
withheld,  especially  in  new  and  strange  surroundings,  the  mice  soon 
fret  themselves  to  death.  If,  however,  the  mice  be  supplied  with 
an  abundance  of  their  usual  nesting  material  and  of  their  natural 
food,  it  is  easy  to  maintain  them  in  a  captive  state.  After  having 
become  accustomed  to  confinement  they  are  somewhat  more  tolerant 
of  any  irregularity. 

Most  individuals  are  content  to  remain  relatively  quiet  during  the 
day,  but  their  periods  of  sleep  are  of  short  duration  and  they  f  re- 


1926]  LIFE   HISTORY   OF   PHENACOMYS   LONGICAUDTJS  59 

quently  climb  among  and  feed  upon  the  twigs  constituting  the  food 
supply.  The  smaller  twigs  are  bitten  off,  carried  to  the  nest,  and 
there  stripped  of  bark.  The  nest,  in  a  wire  pocket  in  one  corner  of 
the  cage,  is  not  usually  arched  over  but  is  left  open,  provided  that 
it  is  shielded  in  some  way  a  few  inches  above  from  a  too  direct  view 
of  persons  moving  about  the  room.  The  animals  sleep  after  the 
usual  rodent  mode — squatted  upon  the  hind  feet  with  the  head  tucked 
between  the  toes ;  but  they  are  quick  to  awaken,  and  resent  the  close 
approach  of  a  peering  face,  pulling  the  loose  needles  up  over  their 
sides  and  backs. 

If  there  is  no  revolving  wheel  in  the  cage  the  mice  spend  a  large 
portion  of  the  night  in  scrambling  about,  upside  down,  on  the  upper 
side  of  the  wire  cover  to  the  cage,  and  it  is  while  thus  engaged  that 
their  special  kind  of  agility  is  displayed  to  best  advantage.  Though 
not  particularly  quick,  they  will  then  turn  and  twist  in  a  variety  of 
intricate  evolutions,  "  skinning  the  cat  "  suspended  by  the  hind  feet, 
twirling  about  suspended  by  a  single  leg,  and  performing  .  many 
other  feats.  Such  play  is  apparently  abandoned  entirely  when  a 
revolving  wheel  is  given  them.  The  mice  will  discover  the  purpose 
of  this  after  a  very  few  minutes  of  investigation,  and  thereafter, 
during  the  hours  of  darkness,  the  slight  noise  of  the  twirling  wheel 
will  be  almost  incessant.  In  fact,  there  is  nothing  capricious  about 
-this  form  of  exercise,  and  it  is  often  sustained  without  pause  for 
such  a  length  of  time  that  one  is  forced  to  the  conclusion  that  in 
nature  tree  mice  are  more  wide  ranging  and  active  during  the  hours 
of  darkness  than  one  would  give  them  credit  for  judging  by  other 
criteria. 

One  of  the  twin  females  that  the  writer  raised  from  birth  took 
weeks  to  get  over  the  habit  of  leaping  from  his  hand  every  time  it 
was  picked  up ;  the  other  would  invariably  slightly  nip  a  finger 
whenever  it  was  poked  into  the  nest.  The  latter  trait  is  rather  com- 
mon, captive  individuals  usually  showing  resentment  of  an  intrusion 
of  the  nest  either  in  this  manner  or  by  rapidly  pushing  against  the 
offending  finger  with  the  forefeet.  At  times  a  mouse  will  begin 
assiduously  licking  one's  hand  in  a  very  droll  manner,  the  attraction 
possibly  being  a  slight  taste  of  saltiness. 

ENEMIES 

Enemies  of  tree  mice  may  be  considered  to  include  adverse  climatic 
conditions.  Some  of  these  conditions  are  very  difficult  to  interpret, 
for  there  may  be  several  critical  factors  which  can  not  be  known. 
Thus,  we  do  not  know  why  the  coastal  region  of  Washington,  ap- 
parently eminently  suitable,  is  entirely  uninhabited  by  the  species, 
although  we  can  readily  understand  that  the  winter  cold  of  a 
higher  altitude  and  a  greater  distance  from  the  sea  might  prevent 
it  from  having  taken  up  a  residence>  in  the  extensive  forests  of  the 
more  mountainous  regions  of  the  Pacific  Coast  States. 

It  is  probable  that  any  normal  dry  cold  experienced  within  the 
habitat  of  the  species  is  readily  endured,  and  captive  individuals 
apparently  do  not  suffer  during  rather  high  summer  temperatures; 
but  it  is  not  unlikely  that  there  is  considerable  discomfort,  and  very 
possibly  some  mortality,  at  least  among  the  younger  animals,  during 
the  severe  rainstorms  of  the  winter  months.    The  nests  are  often  so 


60  NORTH   AMERICAN   FAUNA  [No.  48 

shallow,  exposed,  and  pervious  to  a  soaking  rain  as  to  indicate  that 
a  hard  winter  storm,  might  reduce  half  the  tree-mouse  population 
to  a  state  of  dripping  misery ;  but  the  writer  has  had  no  experience 
with  them  at  that  season.  In  the  experience  of  Wilder,  the  lower 
parts  of  the  nests  usually  remain  snug  and  fairly  dry,  and  he  has 
found  only  one  individual  that  appeared  disturbed  by  a  sodden  con- 
dition of  its  home. 

There  is  comparatively  little  evidence  of  the  depredations  of  active 
enemies  among  tree  mice.  The  marten  is  apparently  absent  from  the 
territory  inhabited  by  them,  else  the  mice  would  quickly  be 
exterminated,  and  this  carnivore  may  very  well  constitute  one  reason 
why  the  range  of  the  Phenacomys  is  not  extended  in  certain  direc- 
tions. It  is  logical  to  assume  that  raccoons  must  occasionally  blunder 
upon  sa  nest,  and  bobcats  possibly  destroy  many  mice  when  other 
food  is  scarce;  but  the  species  has  no  other  mammalian  enemies  of 
which  the  writer  knows. 

Raptorial  birds  must  be  the  chief  active  enemy  with  which  tree 
mice  have  to  contend.  Hawks  can  not  do  them  much  damage;  but 
owls,  especially  the  screech  owl,  would  be  attracted  by  the  mice  for- 
aging among  the  foliage  of  a  tree.  Fortunately  for  the  mice,  these 
little  owls  are  not  particularly  abundant.  Whether  they  have 
learned  to  pull  a  nest  to  pieces  after  seeing  the  fleeing  owner  escape 
into  it  remains  for  some  other  investigator  to  discover. 

The  sudden  disappearance  of  a  nest  long  established  and  the 
equally  sudden  appearance  of  a  nest  in  another  portion  of  the  same 
tree  or  in  an  adjacent  one  for  a  long  time  puzzled  Wilder.  One  day, 
however,  while  he  was  standing  beneath  a  tree,  bits  of  nest  came 
falling  about  him,  and  looking  up,  he  discovered  a  crested  jay 
(Cyanocitta)  busily  engaged  in  demolishing  the  home  of  a  tree 
mouse.  He  has  since  learned  that  this  is  a  regular  procedure  and 
that  while  one  jay  is  thus  engaged  in  the  tree  above,  a  companion 
will  often  remain  on  the  ground  beneath  carefully  searching  the 
falling  debris  for  young  mice.  This  must  constitute  a  real  drain 
upon  the  Phenacomys  population,  and  perhaps  the  jay  is  the  most 
serious  enemy  of  the  tree  mice,  especially  during  years  which  are 
unfavorable  for  ihe  production  of  other  food  for  the  jay,  and  this 
bird  might  well  prove  to  be  the  critical  factor  operating  for  the 
extermination  of  the  mice  over  certain  areas. 


BIBLIOGRAPHY   OF  PHENACOMYS 

The  following  list  includes  the  principal  titles  referring  to  the 
genus  Phenacotnys.  The  majority  of  the  titles  have  reference  to  the 
subject  matter  of  Part  I  of  this  report. 

Axlen,  Joel  Asaph. 

1894.  Descriptions  of  ten  new  North  American  mammals,  and  remarks  on 
others.     Bui.  Amer.  Mus.  Nat.  Hist.  6:  317-332,  November  7. 

Original    description    of    Phenacomys    truei    [=P.    intermedins], 
p.  331. 
1899.  On  mammals  from  the  Northwest  Territory  collected  by  Mr.  A.  J. 
Stone.     Bui.  Amer.  Mus.  Nat.  Hist.  12:  1-9,  March  4. 

Original  description  of  Phenacomys  coiistaolei  [==P.  intermedins}, 
p.  4.  Tvpe  erroneously  stated  to  have  been  collected  August  25, 
1898.     The  year  was  really  1897. 


192G] 


LIFE    HISTORY   OF   PHENACOMYS  LONGICAUDUS  61 


Allen,  Joel  Asaph — Continued. 

1903  Mammals  collected  in  Alaska  and  northern  British,  Columbia  by  the 
Andrew  J.  Stone  Expedition  of  1902.  Bui.  Amer.  Mus.  Nat.  Hist. 
19:  521-567,  October  10. 

Discusses  characters  of  Plwnaoomys  consiablei  [=P.  intermedins], 
p.  553-554 

Bailey,   Vernon. 

1900.  Revision  of  American  voles  of  the  genus  Miorotus.  North  Amer. 
Fauna  No.  17,  88  p.,  illus.,  June  6. 

Type  of  Microtus    {Lagurus)    pumilus   Elliot    (Field   Columb.   Mus. 
Tub.   30,  Zool.    Ser.   1:   226),   proved  by  examination  to  be  a  Phena- 
comys  [=P.  i.  olympicus],  p.   68. 
1915.  Discovery  of  the  tree  mouse.     Oreg.  Sportsman  3 :  147-149,  July. 

Account  of  discovery  of  nest-building  habits  of  Phenacomys  longir 
caudus. 

1917.  Wild  animals.     [U.   S.]   Dept.  Int.,  Natl.  Park  Serv.,  General  infor- 

mation regarding  Yellowstone  National  Park,  season  of  1917,  p. 
44-54.     (Reprinted  191S  and  succeeding  years.) 

Phenacomys  orophilus  [=P.  intermedins]  recorded. 

1918.  Wild  animals.     [U.  S.l  Dept.  Int.,  Natl.  Park  Serv.,  General  infor- 

mation regarding  Glacier  National  Park,  season  of  1918,  p.  43-52. 
(Reprinted  1919.) 

Phenacomys  orophilus  [=P.  intermedins']  recorded. 

1919.  Wild  animals  of  Glacier  National  Park.     The  Mammals.     [Unnum- 

bered pub.l      [U.  S.]   Dept.  Int.,  Natl.  Park  Serv.,  p.  1-102,  illus. 
Dated  1918,  but  issued  January  10,  1919. 

Account  of  Phenacomys  orophilus   [=P.  intermedins],  p.  59-60. 
1923.  Sources  of  water  supply  for  desert  animals.     Sci.  Mo.  17  :  66-86,  illus., 
July. 

Bangs,  Otttram. 

1897.  On  a  small  collection  of  mammals  from  Hamilton  Inlet,  Labrador. 
Proc.  Biol.  Soc.  Washington  11 :  235-240,  illus.,  September  17. 

Records  specimens  later  described  as  Phenacomys  celatus  crassus. 
1S98.  A  list  of  the  mammals  of  Labrador.    Amer.  Nat.  32 :  489-507,  illus. 
Phenacomys  latimanus,  from  Fort  Chimo,  referred  to  on  p.  495. 
1900.  Three  new  rodents  from  southern  Labrador.     Proc.  N.  E.  Zool.  Club 
2 :  35-41,  September  20. 

Original  description  of  Phenacomys  celatus  crassus,  from  Rigo- 
let,  Hamilton  inlet,  Labrador,  p.  39. 

Barrett-Hamilton,  Gerald  Edwin  Hamilton. 

1913.  A  history  of  British  mammals.  Part  14,  p.  361-408,  illus.  London, 
August,  1913.  (Parts  issued  at  irregular  intervals,  March,  1910, 
to  October,  1921.) 

"Although  the  actual  ancestors  of  the  Microtinae  have  not  yet 
been  discovered,  there  can  be  no  doubt  that  their  cheek-teeth  were  of 
a  brachyodont,  tubercular  type"    (p.   385). 

Brooks,  Allan  Cyril. 

1902.  Mammals  of  the  Chilliwack  District,  British  Columbia.  Ottawa  Nat. 
15:  239-244,  February. 

Phenacomys  orophilus  recorded  from  "  Lumsden  Mountain."  Ref- 
erence is  to  type  of  P.  oramontis,  described  by  Rhoads  from  "  Mount 
Baker  Range,"   p.  241. 

Cary,  Merritt. 

1911.  A  biological  survey  of  Colorado.     North  Amer.  Fauna  No.  33,  256  p., 

illus.,  August  17. 

Discusses  distribution  and  status  of  genus  Phenacomys  within  the 
State,  p.  119-120. 

Elliot.  Daniel  Giratjd. 

1898.  Lists  of  mammals  collected  by  W.  W.  Price  and  assistants,  chiefly  in 
California.  Field  Columb.  Mus.  Pub.  27,  Zool.  Ser.  1 :  193-216, 
March. 

Phenacomys  orophilus  [=P.  intermedins]  recorded  from  Pyramid 
Peak,  Eldorado   County,  California,  p.  204. 


62  NORTH   AMERICAN   FAUNA  [No.  48 

Elliot,  Daniel  Giraud — Continued. 

1899a.  Preliminary  descriptions  of  new  rodents  from  the  Olympic  Moun- 
tains. Field  Columb.  Mus.  Pub.  30,  Zool,  Ser.  1:255-228,  Febru- 
ary 1. 

Original  descriptions  of  Phenacomys  olympicus,  and  Microtus 
(Layurus)  pumilus  [=  P.  i.  olympicus],  p.  225-226. 

1899b.  Catalogue  of  mammals  from  the  Olympic  Mountains,  Washington, 
with  descriptions  of  new  species.  Field  Columb.  Mus.  Pub.  32, 
Zool.  Ser.  1 :  239-276,  illus.,  March  4. 

Phenacomys  olympicus  treated,  p.  255 ;  and  Microtus  pumilus 
[=  Phenacomys  i.  olympicus],  p.  258. 

1907.  A  catalogue  of  the  collection  of  mammals  in  the  Field  Columbian 

Museum.    Field  Columb.  Mus.  Pub.  115,  Zool.  Ser.,  v.  8,  694  p.,  illus. 

Phenacomys  olympicus  listed  from  Happy  Lake  and  Boulder  Lake, 
Washington,  p.   282. 

Grinnell,  Joseph.  , 

1913.  A   distributional   list   of   the  mammals   of   California.      Proc.    Calif. 

Acad.  Sci.  (4)  3:  265-390,  illus.,  August  28. 

Phenacomys  longicaudus  recorded  from  Mount  Sanhedrin,  p.  316. 

Hinton,  Martin  Alister  Campbell. 

1910.  Preliminary  account  of  the  British  fossil  voles  and  lemmings,  with 
some  remarks  on  the  pleistocene  climate  and  geography.  Proc. 
Geol.  Assoc.  [London]  21 :  489-507. 

Discussion  of  remains  of  voles  with  fanged  molars  from  the  forest 
bed  horizon,  and  description  of  new  species  of  the  genus  Mimomys. 

Hollister,  Ned. 

1913.  Mammals  of  the  Alpine  Club  expedition  to  the  Mount  Robson  region. 
Canad.  Alpine  Jour.,  Spec.  Number,  p.  1-44,  illus.,  February  17. 

Series  of  specimens  from  the  Mount  Robson  region  referred  to 
Phenacomys  intermedins,  on  p.  20.  Type  of  P.  intermedins  referred 
to  and  other  species  discussed. 

Howell,  Alfred  Brazier. 

1920.  A  second  record  of  Phenacomys  albipes  in  California,  with  a  discus- 

sion of  the  species.     Jour.  Mammal.  1 :  242-243,  November. 

A  specimen  recorded  from  Orick,  Calif. 

1921.  Description  of  a  new  species  of  Phenacomys  from  Oregon.     Jour. 

Mammal.  2 :  98-100,  illus.,  May  2. 

Original  description  of  Phenacomys  silvicola. 

1923.  Descriptions  of  two  new  rodents  of  the  genus  Phenacomys.  Proc. 
Biol.  Soc.  Washington  36 :  157-158,  May  1. 

Original  descriptions  of  Phenacomys  intermedins  levis  from  St. 
Marys  Lake,  Mont.  ;  and  P.  i.  eels  us,  from  Tuolumne  Meadows, 
Yosemite  National  Park,  Calif. 

Jewett,  Stanley  Gordon. 

1915.  Discovery  of  a  rare  rodent.     Oreg.  Sportsman  3 :  37-38. 

Specimen  of  P.  albipes  recorded  from  Vida,  Lane  County,  Oreg. 

1920.  Notes  on  two  species  of  Phenacomys  in  Oregon.  Jour.  Mammal.  1: 
165-168,  August. 

Notes  on  habits  of  Phenacomys  longicaudus  and  P.  albipes. 

1923.  A  breeding  record  of  Phenacomys  longicaudus.  Jour.  Mammal.  4 : 
125,  May. 

Records  the  finding  of  young  of  this  species. 
Lyon,    Marcus    Ward,    jr. 

1907.  Notes  on  mammals  collected  at  Mount  Rainier,  Wash.  Smithsn. 
Misc.  Collect.  50:  89-92,  June  27.     (Quart,  issue,  v.  4,  pt.  1.) 

P.  oronhilus  [=P.  i.  olympicus]  recorded  from  Paradise  Park, 
Mount  Rainier,  Wash.,  p.  91. 

Lyon,  Marcus  Ward,  jr.,  and  Wilfred  Hudson  Osgood. 

1909.  Catalogue  of  the  type-specimens  of  mammals  in  the  United  States 
National  Museum,  including  the  Biological  Survey  collection. 
U.  S.  Nat.  Mus.  Bui.  62,  325  p. 

Details  of  condition  of  type  specimens  of  Phenacomys  albipes, 
P.  longicaudus,  P.  mackensii,  P.  orophUus  [=P.  intermedins],  P. 
preblei  [=P.  intermedins],  and  P.  truei  [=P.  intermedins],  p. 
86-97. 


1926] 


LIFE   HISTORY   OF   PHENACOMYS  LONGICAUDUS  63 


Major,  C.  I.  Forsyth. 

1902.  [Exhibition  of,  and  remarks  npon,  some  jaws  and  teeth  of  Pliocene 
voles  (Mimomys,  gen.  nov.).]  Proc.  Zool.  Soc.  London  1:  102-107, 
illus.,  February  18.     (Also  reported  in  Nature  65:  431,  March  6.) 

The  name  Mimomys  proposed  for  certain  voles  with  rooted  molars 
from'  the  Pliocene  of  Italy  and  England,  including:  Miorotus  inter- 
medins (Newton)   referred  by  some  authors  to  Phenacomys. 

Mereiam,  Clinton  Hart. 

1889.  Description  of  a  new  genus  {Phenacomys)  and  four  new  species  of 
Arvicolinae.  In  his  Descriptions  of  fourteen  new  species  and  one 
new  genus  of  North  American  mammals,  North  Amer.  Fauna  No. 
2,  p.  27-35,  illus. 

Original    descriptions    of    genus    Phenacomys,   and    species    Phena- 
comys intermedins,  P.  celatws  [=P.  ungara],  P.  lutimanus  [=P.  un- 
gava],  and  P.  ungava. 
1891.  Results  of  a  biological  reconnoissance  of  south-central  Idaho.     North 
Amer.  Fauna  No.  5,  p.  1-108,  illus.,  July  30. 

Original  description  of  Phenacomys  orophilus   [=P.  intermedins], 
p.  65-60. 
1S92.  The  geographic  distribution  of  life  in  North  America  with   special 
reference  to  the  mammalia.     Proc.  Biol.  Soc.  Washington  7 :  1-64, 
April. 

Arvicola  borealis  Richardson  provisionally  referred  to  genus 
Phenacomys. 

1897a.  The  mammals  of  Mount  Mazama,  Oregon.  Mazama  1 :  204-230, 
October. 

P.  orophilus  [=P.  intermedins]  recorded  from  several  localities  in 
Oregon,  p.  221. 

1897b.  Phenacomys  preblei,  a  new  vole  from  the  mountains  of  Colorado. 
Proc.  Biol.  Soc.  Washington  11:  45,  March  16. 

P.  preblei  [=P.  intermedins]  described  from  Twin  or  Lilies  Teak, 
near  Longs  Peak,  Colo. 

1899.  Results  of  a  Biological  Survey  of  Mount  Shasta,  California.     North 

Amer.  Fauna  No.  16,  179  p.,  illus.,  October  2S. 

Phenacomys    orophilus    [=P.    intermedins]    recorded    from    Mount 
Shasta,  p.  95. 
1901.  Two  new   rodents  from  northwestern   California.     Proc.    Biol.    Soc. 
Washington  14 :  125-126,  July  19. 

Original  description  of  Phenacomys  albipes  from  Redwoods  near 
Areata,  Humboldt  Bay,  Calif. 

Miller,  Gerrit  Smith,  jr. 

1896.  Genera  and  subgenera  of  voles  and  lemmings.  North  Amer.  Fauna 
No.  12,  84  p.,  illus.,  July  23. 

Defines  and  discusses  genus  Phenacomys,  p.  40-42. 
1897a.  Notes  on  the  mammals  of  Ontario.     Proc.   Boston  Soc.  Nat.  Hist. 
28:  1-44. 

Phenacomys  latimanus  [=P.  ungava]  recorded  from  Teninsula 
Harbor,   north  shore   of  Lake  Superior. 

1897b.  Synopsis  of  the  voles  of  the  genus  Phenacomys.  Proc.  Biol.  Soc. 
Washington  11 :  77-87,  April  21. 

1900.  Key  to  the  land  mammals  of  northeastern  North  America.     N.   Y. 

State  Mus.  Bui.  S;  59-160,  October. 

Brief  treatment  of  Phenacomys  latimanus  [=P.  ungava],  p.  108. 

Nehring,  Karl  Wilhelm  Alfred. 

1894.  [Review  of  North  American  Fauna  Numbers  1-5].  Naturw. 
Wchnschr.   [Berlin]   9:346. 

Assigns  to  Phenacomys  remains  from  cave  in  Hungary,  p.  346. 

Preble,  Edward  Alexander. 

1902a.  Descriptions  of  new  species  of  Synaptomys  and  Phenacomys  from 
Mackenzie,  Canada.  Proc.  Biol.  Soc.  Washington  15 :  181-182, 
August  6. 

Original  description  of  Phenacomys  mackensii,  p.  182. 


64  NORTH   AMERICAN   FAUNA  [No.  48 

Preble,  Edward  Alexander — Continued. 

1902b.  A  biological  investigation  of  the  Hudson  Bay  Region.  North  Amer. 
Fauna  No.  22,  140  p.,  illus.,  October  31. 

P.  mackensii  recorded  from  Fort  Churchill,  Hudson  Bay,  p.  50. 

1908.  A  biological  investigation  of  the  Atbabaska-Mackenzie  Region.  North 
Amer.  Fauna  No.  27,  574  p.,  illus.,  October  26. 

Discusses    characters    and    ranges    of    Phenacomys    mackensii   and 
P.  intermedins  in  this  region,  p.  177-178. 

Rhoads,  Samuel  Nicholson. 

1S95.  Additions  to  the  mammal  fauna  of  British  Columbia.  Amer.  Nat. 
29:940-942,  October. 

Original  description  of  Phenacomys  oramontis  [=P.  intermedins], 
p.  941. 

Saunders,  William  Edwin. 

1920.  Phenacomys  intermedins  from  High  River,  Alberta.  Canad.  Field 
Nat.  34 :  17.  / 

Phenacomys  intermedins  recorded  from  High  River,  Alberta. 

1926.  Phenacomys  ungava.     Canad.  Field  Nat.  40:18. 
Two  specimens  taken  at  Franz,  Ontario. 
Stone,  Witmer. 

1904.  On  a  collection  of  birds  and  mammals  from  Mount  Sanhedrin,  Cali- 
fornia.    Proc.  Acad.  Nat.  Sci.  Phila.  56 :  576-5S0. 

Specimen     of     Phenacomys     longicaudus     recorded     from     Mount 
Sanhedrin. 

Taylor,  Walter  Penn. 

1915a.  Relative   numbers   in   sexes   of  the   tree   mouse.     Oreg.    Sportsman 

3 :  150-151,  July. 
1915b.  Description  of  a  new  subgenus   (Arborimus)   of  Phenacomys,  with 

a    contribution    to   knowledge    of   the   habits   and    distribution  of 

Phenacomys   longicaudus  True.     Proc.   California  Acad.   Sci.    (4) 

5 :  111-161,  illus.,  December  30. 

Todd,  Aurelius. 

1891.  An  Oregon  mouse.     West  Amer.  Scientist  7 :  240-242,  September. 

Discusses  habits  of  Phenacomys  longicaudus. 

True,  Frederick  William. 

1890.  Description  of  a  new  species  of  mouse,  Phenacomys  longicaudus,  from 
Oregon.     Proc.  U.  S.  Nat.  Mus.  13:  303-304,  November  15. 

Original  description  of  Phenacomys  longicaudus  from  Marshfield. 
Coos  County,  Oreg. 

Warren,  Edward  Royal. 

1906.  The  mammals  of  Colorado.  Colo.  Col.  Pub.  11 :  225-274  (Gen.  Ser.  19 : 
Sci.  Ser.  46). 

Reference    to   Phenacomys  prcblei.  [=P.   intermedins]    from    Twin 
Peak,  near  Longs  Peak,  Colo..,  p.  249. 

1908.  Further  notes  on  the  mammals  of  Colorado.  Colo.  Col.  Pub.  (Engi- 
neering Ser.)  1:61-89  (Gen.  Ser.  33). 

Phenacomys    orophilns     [=P.    intermedins]     recorded    from    Lake 
Moraine,  p.  76. 
1912.  Notes  on  the  distribution  of  some  Colorado  mammals.    Proc.  Biol.  Soc. 
Washington  25 :  3-7,  January  23. 

Phenacomys  orophilns    [=P.   intermedins]    recorded   from    Buffalo 
Pass,  p.  4. 

Wight.  Howard  Marshall. 

1925.  Notes  on  the  tree  mouse,  Phenacomys  silvicola.  Jour.  Mammal.  6: 
282-283,  November. 

Records  Phenacomys  silvicola  from  ridge  between  Willamette  Val- 
ley and  the  Coast  Range,  Oreg.     First  record  east  of  this  range. 

Young,  Robert  Thompson. 

1908.  Notes  on  the  distribution  of  Colorado  mammals,  with  a  description 
of  a  new  species  of  bat  (Eptesicus  pallidus)  from  Boulder.  Proc. 
Acad.  Nat.  Sci.  Phila.  60:  403-409,  July. 

Phenacomys    preblei     I— P.    intermedins]     recorded    from    North 
Boulder  Creek,  Colo. 


INDEX 


Abies  grandis,  43. 

nordmanniana,  53. 
Acknowldgments,  3. 
albipes,  Phenacoruys,  5,  12,  30-31. 
angusticeps,  Microtus  mordax,  31. 
Arborimus,  5. 
Arbutus  menziesii,  43. 
ariaefolia,  Holodiscus  discolor,  43. 
Arrowwood,  43. 
Arvicola  intermedius,  5. 

Big-cone  spruce,  54. 

California  bay  laurel,  43. 

lilac,  43. 
californica,  Corylus  rostrata,  43. 

Umbellularia,  43. 
Ceanotbus  thyrsiflorus,  43. 
celatus,  Pbenacomys,  4. 
celsus,  Pbenacomys  intermedius,  7,  23- 

25. 
constablei,  Phenacomys,  5,  15,  18. 
Corylus  rostrata  californica,  43. 
crassus,  Pbenacomys  ungava,  27-28. 
Crested  jay,  60. 
Cricetinae,  11. 
Cyanocitta,  60. 

densiflora,  Pasania,  43. 
Deodar,  53. 

Distribution,   geograpbic,   1. 
Douglas  fir,  43,  52. 

Evotomys,  6,  10. 

Fir,  Douglas,  43,  52. 

grand,  43. 
Food,  50. 

Geograpbic  distribution,  1. 
Grand  fir,  43. 
grandis  Abies,  43. 
Groups,  12. 

albipes,   30-31. 

intermedius,  13-25. 

longicandus,  32-35. 

ungava,  7,  9,  25-29. 

Habits,  1,  39-60. 

Hazel,  43. 

History,  4. 

Holodiscus  discolor  ariaefolia,  43. 

intermedius,  Arvicola,  5. 

Pbenacomys  intermedius,  4,  15-20. 

Jay,  crested,  60. 


latimanus,  Pbenacomys,  4. 

Laurel,  California  bay,  43. 

Lemming  mice,  1. 

levis,  Pbenacomys  intermedius,  20-21. 

Life  history,  39-60. 

Lilac,  California,  43. 

longicaudus,  Phenacomys,  4,  5,  6,  7,  32- 

34,  39-60. 
Lumsden    (Lihumitson)    Mountain,  15, 

61. 

mackenzii,  Pbenacomys,  2S-29. 
macrocarpa,  Pseudotsuga,  54. 
Madrone.  43. 
Material,  3. 
Measurements,  4. 
menziesii,  Arbutus,  43. 
Microtus,  5,  6,  7,  11. 

(Lagurus)  pumilus,  5. 

mordax  angusticeps,  31. 
Mimomys,  5. 

Nests,  43. 
nordmanniana,  Abies,  53. 

Oak,  tan,  43. 

olympicus,  Phenacomys  intermedius,  5, 

7,  21-23. 
Ondatra,  6,  9,  10. 
oramontis,  Phenacomys,  5,  15. 
orophilus,  Phenacomys,  5,  15. 

Palates  (figured),  10. 
Pasania  densiflora,  43. 
Phenacomys,    Alberta,   20-21. 

coast,  30-32. 

forest  tree,  34-35. 

Labrador,    27-28. 

Mackenzie,  28-29. 

Olympic,  21-23. 

red   tree,   32-34. 

life  history,  39-60. 

Rocky  Mountain,  15-20. 

Sierran,  23-25. 
Phenacomys  albipes,  5,  12,  30-31. 

celatus,  4. 

celsus,  7,  23-25. 

constablei,  5,  15,  18. 

crassus,  27-28. 

intermedius,  4,  15-20. 

latimanus,  4. 

levis,  20-21. 

longicaudus,  4,  5,  6,  7,  32-34, 

mackenzii,   28-29. 

olympicus,  5,  7,  21-23, 

oramontis,  5,  15. 

orophilus,  5,  15. 

preblei,  5,  15. 

65 


66 


NORTH    AMERICAN    FAUNA 


[No.  48,  1926] 


Phenacomys — Continued. 

silvicola,  34-35. 

truei,  5,  15. 

ungava,  4,  7,  25=27. 
Picea  sitchensis,  46. 
preblei,  Phenacomys,  5,  15. 
Pseudotsuga  macroearpa,  54. 

taxifolia,  43,  52. 

Redwood,  43. 

Sequoia  sempervirens,  43. 
silvicola,  Phenacomys,  34-35. 
sitchensis,  Picea,  46. 
Sitka  spruce,  46. 
Skull,  7. 


Spiraea,  43. 
Spruce,  big-cone,  54. 
Sitka,  46. 

Tan  oak,  43. 

taxifolia,  Pseudotsuga,  43,  52. 

Teeth,  7. 

thyrsiflorus,  Ceanothus,  43. 

Tree  mouse,  forest,  34-35. 

red,  1,  32-34,  39. 
truei,  Phenacomys,  5,  15. 
Type  localities,  13. 

Umbellularia  californica,  43. 
ungava,  Phenacomys  ungava,  4,  7,  25- 
/    27. 


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