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74 


HARVARD    UNIVERSITY 

Library  of  the 

Museum  of 

Comparative  Zoology 


NORTH  AMERICAN  QUATERNARY  CANIS 


Frontispiece 

Three  species  of  Canis.  Top,  Canis  lupus  (the  gray  wolf,  photo  by  L. 
David  Mech).  Middle,  Canis  rufus  (the  red  wolf,  photo  by  Curtis 
Carley).  Bottom,  Canis  latrans  (the  coyote,  photo  by  Tom  Smylie). 
All  photographs  courtesy  of  the  Fish  and  Wildlife  Service,  Depart- 
ment of  the  Interior. 


NORTH  AMERICAN  QUATERNARY  CAMS 


RONALD  M.  NOWAK 

Staff  Specialist 

Office  of  Endangered  Species 

U.S.  Fish  and  Wildlife  Service 

Department  of  the  Interior 

Washington,  D.C.  20240 


MONOGRAPH 

OF  THE 
MUSEUM  OF  NATURAL  HISTORY,  UNIVERSITY  OF  KANSAS 

NUMRER  6 

1979 


NORTH  AMERICAN  QUATERNARY  CAN1S 


MONOGRAPH  OF  THE  MUSEUM  OF  NATURAL  HISTORY 

THE  UNIVERSITY  OF  KANSAS 

Number  6,  pages  1-154,  text  figures  1-55 
September  1, 1979 


MUS.  COMP.  ZOOL" 
LIBRARY 

JUN  1  8  1980 

HARVARD 

UNIVERSITY 


Editor:  E.  O.  Wiley 


Copyrighted 

By 

Museum  of  Natural  History 

The  University  of  Kansas 

Lawrence,  Kansas  66045 

U.S.A. 

ISBN:  0-89338-007-5 


Printed 

By 

The  University  of  Kansas  Printing  Service 

Lawrence,  Kansas 

U.S.A. 


CONTENTS 


INTRODUCTION    _  1 

Acknowledgements    2 

Methods   .— 4 

Age  and  Secondary  Sexual  Variation  6 

HISTORY  AND  STATISTICAL  ANALYSIS  OF  RECENT  POPULATIONS  7 

Comparison  of  Known  Series  of  Dogs,  Wolves,  and  Coyotes  7 

Systematic  Problems  in  the  Northeast  12 

Systematic  Problems  in  the  Southeast  _.. 24 

SYSTEMATIC  DESCRIPTIONS  66 

Genus  Canis  Linnaeus   66 

Canis  cedazoensis  Mooser  and  Dalquest 68 

Canis  Jcpophagus  Johnston  68 

Canis  Jatrans  Say  73 

Canis  edwardii  Gazin  82 

Canis  rufns  Audubon  and  Bachman 85 

Canis  armbrusteri  Gidley 90 

Canis  lupus  Linnaeus  93 

Canis  familiaris  Linnaeus 102 

Canis  dims  Leidy   -  106 

SUMMARY 118 

LITERATURE  CITED  121 

ADDENDUM  136 

APPENDIX  A  138 

APPENDIX  B   144 

APPENDIX  C  150 


INTRODUCTION 


According  to  the  revisionary  work  of  An- 
derson (1943),  Goldman  (1937,  1944),  and 
Jackson  ( 1951 ) ,  three  living  species  of  wild 
Cards  occur  in  North  America:  Cards  la- 
trans,  the  coyote;  C.  rufits,  the  red  wolf;  and 
C.  lupus,  the  gray  wolf.  Although  this  taxo- 
nomic  arrangement  has  been  generally  ac- 
cepted, some  questions  have  arisen  concern- 
ing matters  not  fully  explained  by  the 
revisions,  and  certain  newly  recognized  phe- 
nomena  within   canid  populations. 

Much  attention,  especially  since  1960,  has 
been  directed  toward  the  systematics  of  Cards 
in  the  eastern  half  of  North  America.  There 
has  been  controversy  regarding  the  taxonomic 
status  of  the  wolves  (C.  lupus  hjcaon  and 
subspecies  of  C.  rufus)  originally  found  there, 
and  of  the  populations  of  Canis  presently 
inhabiting  the  region  (roughly,  east  of  100°W 
and  south  of  50°N).  The  production  of  fully 
fertile  hybrids,  of  common  occurrence  among 
captive  Canis  (Gray,  1972),  has  been  sug- 
gested as  having  affected  wild  Canis  in  the 
eastern  part  of  the  continent. 

In  addition  to  the  questions  concerning 
living  Canis,  there  are  problems  involving  the 
paleontological  history  of  the  genus  in  North 
America.  Although  many  fossil  specimens 
have  been  described,  not  all  of  them  have 
been  assigned  to  particular  lineages  ancestral 
to  living  populations.  Of  special  interest,  be- 
cause it  is  the  only  fossil  kind  represented  by 
what  a  modern  mammalogist  would  call  a 
good  series,  is  the  dire  wolf,  Canis  dims. 

My  aim  in  studying  Canis  was  to  examine 
large  series  of  specimens  from  throughout 
North  America,  in  order  to  obtain  a  clearer 
understanding  of  the  systematic  relationships 
between  the  species  represented.  I  hoped  to 
get  an  idea  of  the  extent  of  variation  within 
the  Recent  wolves  found  in  the  western  and 
northern  parts  of  the  continent,  and  to  deter- 
mine the  relative  positions  of  C.  dints,  C. 
rufus,  and  C.  lupus  hjcaon.  I  wanted  also  to 
ascertain,  as  well  as  possible,  the  origins  and 


relationships  of  the  presently  existing  popu- 
lations of  Canis  in  the  east.  Partly  from 
study  of  populations,  I  hoped  to  distinguish 
and  more  accurately  delineate  (morphologi- 
cally, geologically,  geographically)  the  living 
and  extinct  species  of  North  American  Canis. 

Because  of  their  recognized  taxonomic 
value,  abundance  in  museum  collections,  pa- 
leontological preservation,  and  relative  ease 
of  handling,  I  used  skulls  as  the  primary  ma- 
terial of  my  study.  Approximately  5,000  spec- 
imens were  examined. 

The  first  main  part  of  the  paper  consists 
of  an  historical  sketch  and  a  statistical  analy- 
sis based  on  those  populations  represented  by 
large  series  of  complete  skulls.  This  analysis 
serves  to  delineate  special  groups  and  to  as- 
sess the  probable  origin  and  relationship  of 
questionable  populations.  The  BMD07M 
program  of  multivariate  analysis  was  a  pri- 
mary method  employed  in  this  study.  The 
second  main  part  of  the  paper  consists  of 
descriptions  of  each  recognized  species  of 
North  American  Canis.  Some  of  the  speci- 
mens discussed,  including  many  of  the  fossils, 
could  not  be  used  in  multivariate  analysis,  but 
the  descriptions  are  supported  in  part  by  uni- 
variate and  bivariate  statistics. 

Collections  cited  in  this  paper  are  repre- 
sented by  the  following  abbreviations: 
AMNH,  American  Museum  of  Natural  His- 
tory; ANSP,  Academy  of  Natural  Sciences, 
Philadelphia;  CM,  Carnegie  Museum;  CNM, 
National  Museum  of  Canada;  FGS,  Florida 
Geological  Survey;  FM,  Field  Museum;  ISM, 
Illinois  State  Museum;  KU,  University  of 
Kansas  Museum  of  Natural  History;  LACM, 
Los  Angeles  County  Museum  of  Natural  His- 
tory; LPI,  Louisiana  Polytechnic  Institute 
Department  of  Zoology;  LSUMZ,  Louisiana 
State  University  Museum  of  Zoology;  MCZ, 
Harvard  University  Museum  of  Comparative 
Zoology;  MSU,  Michigan  State  University 
Museum;  NYEC,  New  York  Department  of 
Environmental   Conservation;    PPM,   Panhan- 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


die  Plains  Museum;  PUWL,  Purdue  Univer- 
sity Wildlife  Laboratory;  QWS,  Quebec  Wild- 
life Service;  ROM,  Royal  Ontario  Museum; 
SD,  San  Diego  Natural  History  Museum; 
SMUMP,  Southern  Methodist  University  Mu- 
seum of  Paleontology;  SR,  Sul  Ross  State  Uni- 
versity Department  of  Biology;  TM,  Texas 
Memorial  Museum;  UAlb,  University  of  Al- 
berta Department  of  Zoology;  UAriz,  Univer- 
sity of  Arizona  Department  of  Biological 
Sciences  and  Laboratory  of  Paleontology; 
UArk,  University  of  Arkansas  Department  of 
Zoology;  UCMP,  University  of  California 
Museum  of  Paleontology;  UCMVZ,  University 
of  California  Museum  of  Vertebrate  Zoology; 
UColo,  University  of  Colorado  Museum;  UF, 
University  of  Florida  State  Museum;  UI,  Uni- 
versity of  Illinois  Museum  of  Natural  History; 
UMMP,  University  of  Michigan  Museum  of 
Paleontology;  UMMZ,  University  of  Michi- 
gan Museum  of  Zoology;  UMinn,  University 
of  Minnesota  Museum  of  Natural  History; 
UN,  University  of  Nebraska  State  Museum; 
UO,  University  of  Oklahoma  Museum; 
USFWS,  United  States  Fish  and  Wildlife 
Service  field  collections;  USNM,  United 
States  National  Museum  of  Natural  History; 
VFG,  Vermont  Fish  and  Game  Department. 
A  few  other  collections  are  spelled  out  in  the 
text.  Other  common  abbreviations  in  this 
paper  include  "C."  for  Canis  and  "A."  for 
Aenocyon. 

This  paper  is  a  slightly  modified  version 
of  a  Ph.D.  dissertation  (Nowak,  1973)  sub- 
mitted to  the  University  of  Kansas  in  1973. 
Since  that  year  substantial  new  information 
has  become  available,  as  for  example  through 
Kurten's  ( 1974 )  study  of  fossil  coyotes,  Kole- 
nosky  and  Standfield's  (1975)  analysis  of 
wolves  in  Ontario,  and  Mooser  and  Dal- 
quest's  ( 1975 )  description  of  a  new  species 
of  North  American  Pleistocene  Canis.  In 
addition,  a  number  of  specimens  have  been 
collected  recently  in  southeastern  Texas,  and 
this  material  allows  an  updating  of  the  status 
of  the  red  wolf  in  that  area.  Although  I  have 
devoted   some   space  to  the  newly  available 


views  and  data,  they  have  not,  in  all  cases, 
received  the  same  degree  of  attention  shown 
the  earlier  material.  Other  differences  be- 
tween my  dissertation  and  this  paper  include 
the  dropping  in  the  latter  of  several  figures 
and  tables  of  measurements,  the  relegation 
of  the  statistical  analysis  of  the  dire  wolf  to 
the  section  entitled  "Systematic  Descriptions," 
and  the  correction  of  several  errors.  I  have 
not  cited  my  dissertation  as  a  reference  for 
this  paper,  except  in  a  few  instances  in  which 
mention  of  the  contrast  between  the  two 
seemed  warranted. 

Acknowledgements 

It  is  impossible  to  express  my  full  appre- 
ciation to  all  who  assisted  me.  I  must,  how- 
ever, single  out  Professor  E.  Raymond  Hall, 
Museum  of  Natural  History,  University  of 
Kansas,  who  was  my  major  advisor  until  his 
retirement  in  May  1972,  and  who  then  vol- 
untarily continued  to  act  in  this  capacity. 
Professor  Hall  initially  suggested  that  I  do 
graduate  work  at  the  University  of  Kansas, 
and  it  was  his  idea  for  me  to  make  a  study 
of  the  relationships  between  species  of  North 
American  Quaternary  Canis.  On  countless 
occasions  he  provided  me  with  assistance  and 
advice  regarding  my  dissertation  and  my 
general  program  of  work  at  the  University. 
I  consider  it  a  rare  honor  to  have  been  among 
his  students. 

Practically  all  of  the  other  instructors,  and 
many  of  the  students,  with  whom  I  have  been 
associated  at  the  University  of  Kansas,  have 
at  one  time  or  another  given  me  some  help 
that  eventually  contributed  to  this  paper.  I 
thank  them  all,  but  specifically  want  to  men- 
tion Professors  Robert  S.  Hoffmann,  Robert 
M.  Mengel.  and  Craig  C.  Black  (now  of  the 
Carnegie  Museum).  I  also  am  grateful  to 
Professor  Peter  M.  Neely,  Associate  Director 
of  the  University  Computer  Center,  who  took 
much  of  his  time  to  explain  the  processes  and 
results  of  the  BMD07M  computer  program. 
Although   I   could  not  have   effectivelv  used 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


BMD07M  without  Professor  Neely's  help,  I 
take  full  responsibility  for  the  application 
and  interpretation  of  this  program  with  re- 
gard to  the  problems  of  my  study.  Additional 
valuable  assistance  on  the  use  and  under- 
standing of  computers  was  provided  by  my 
fellow  student,  Alberto  Cadena. 

The  research  required  in  preparation  of 
this  paper  necessitated  considerable  travel  to 
museums  and  other  localities  throughout 
North  America.  I  therefore  am  especially 
grateful  to  those  organizations  that  aided  me 
in  this  regard.  The  Theodore  Roosevelt  Me- 
morial Fund  of  the  American  Museum  of 
Natural  History,  and  the  National  Science 
Foundation  each  made  a  direct  grant  for 
travel  and  related  expenses.  The  Committee 
on  Systematics  and  Evolutionary  Biology, 
University  of  Kansas,  provided  travel  grants 
in  1970  and  1971,  a  research  assistantship  in 
the  summer  of  1972,  and  also  a  traineeship  for 
the  academic  year  1971-1972. 

No  progress  could  have  been  made  in  my 
research  had  it  not  been  for  the  cooperation 
of  numerous  persons  who  generously  assisted 
me  in  the  examination  of  specimens  and  asso- 
ciated materials  in  their  care.  I  want  to 
especially  thank  John  L.  Paradiso,  Bird  and 
Mammal  Laboratories,  United  States  National 
Museum  of  Natural  History  (now  of  the 
Office  of  Endangered  Species,  U.S.  Fish  and 
Wildlife  Service).  During  the  four  months 
that  my  wife  and  I  worked  at  the  National 
Museum,  he  aided  us  in  every  way  possible 
and  spent  a  great  deal  of  his  own  time  to  see 
that  we  were  well  provided  for  both  in  and 
out  of  the  Museum.  John  and  I  actually  have 
been  in  close  communication  regarding  Canis 
since  1965.  Many  of  the  views  expressed  in 
this  paper  were  developed  jointly  with  him 
in  the  course  of  years  of  pleasant  study,  con- 
versation,   and    correspondence. 

I  am  also  grateful  to  the  following  per- 
sons who  either  sent  me  specimens  on  loan 
or  assisted  me  when  I  visited  their  areas: 
Sydney  Anderson,  American  Museum  of  Nat- 
ural History;  Rollin  H.  Baker,  The  Museum, 


Michigan  State  University;  Troy  L.  Best,  Mu- 
seum of  Zoology,  University  of  Oklahoma; 
Elmer  C.  Birney,  Museum  of  Natural  History, 
University  of  Minnesota;  Ben  Day,  Vermont 
Fish  and  Game  Department;  Diana  Van  El- 
sacker.  University  of  Colorado  Museum; 
David  E.  Fortsch,  Los  Angeles  County  Mu- 
seum of  Natural  History;  Philip  S.  Gipson, 
Department  of  Zoology,  University  of  Arkan- 
sas; John  W.  Goertz,  Department  of  Zoology, 
Louisiana  Polytechnic  Institute;  John  E.  Guil- 
day,  Carnegie  Museum;  C.  R.  Harington, 
National  Museum  of  Canada;  Billy  R.  Harri- 
son, Panhandle  Plains  Museum;  Claude  W. 
Hibbard,  Museum  of  Paleontology,  Univer- 
sity of  Michigan;  Donald  F.  Hoffmeister, 
Museum  of  Natural  History,  University  of 
Illinois;  Emmet  T.  Hooper,  Museum  of  Zool- 
ogy, University  of  Michigan;  J.  H.  Hutchison, 
Museum  of  Paleontology,  University  of  Cali- 
fornia; Frederick  F.  Knowlton,  U.S.  Fish  and 
Wildlife  Service;  Barbara  Lawrence,  Museum 
of  Comparative  Zoology,  Harvard  University; 
Everett  H.  Lindsay,  Laboratoiy  of  Paleontol- 
ogy, University  of  Arizona;  George  H.  Low- 
ery,  Jr.,  Museum  of  Zoology,  Louisiana  State 
University;  Ernest  L.  Lundelius,  Jr.,  Texas 
Memorial  Museum;  Larry  D.  Martin,  Museum 
of  Natural  History,  University  of  Kansas; 
John  D.  Newsom,  Louisiana  State  Univer- 
sity Cooperative  Wildlife  Research  Unit; 
Robert  T.  Orr,  California  Academy  of  Sci- 
ences; N.  Panter,  Department  of  Zoology, 
University  of  Alberta;  Paul  W.  Parmalee, 
Illinois  State  Museum;  Oliver  P.  Pearson, 
Museum  of  Vertebrate  Zoology,  University 
of  California;  Randolph  L.  Peterson,  Royal 
Ontario  Museum;  Charles  Pichette,  Quebec 
Wildlife  Service;  Douglas  H.  Pimlott,  Univer- 
sity of  Toronto;  Clayton  E.  Ray,  U.S.  Na- 
tional Museum  of  Natural  History;  Richard 
L.  Reynolds,  Los  Angeles  County  Museum 
of  Natural  History;  Horace  G.  Richards, 
Academy  of  Natural  Sciences,  Philadelphia; 
Glynn  Riley,  Jr.,  U.S.  Fish  and  Wildlife 
Service;  C.  B.  Bobbins,  Department  of  Bio- 
logical Sciences,  University  of  Arizona;  Den- 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


nis  N.  Russell,  Texas  Parks  and  Wildlife 
Department;  Donald  Schierbaum,  New  York 
Department  of  Environmental  Conservation; 
James  F.  Scudday,  Department  of  Riology, 
Sul  Ross  State  University;  Beryl  E.  Taylor 
and  Richard  H.  Tedford,  American  Museum 
of  Natural  History;  Gilmer  Voss,  San  Diego 
Natural  History  Museum;  S.  David  Webb, 
University  of  Florida  State  Museum;  J.  Wil- 
liam Yon,  Florida  Geological  Survey;  Phillip 
M.  Youngman,  National  Museum  of  Canada; 
and  Curtis  J.  Carley,  U.S.  Fish  and  Wildlife 
Service. 

These  acknowledgements  would  not  be 
complete  without  the  names  of  my  parents, 
Jacob  and  Esther  Nowak,  New  Orleans,  Lou- 
isiana. Throughout  the  course  of  my  research 
they  were  always  ready  and  willing  to  pro- 
vide any  assistance,  whether  requested  or  not. 

I  finally  wish  to  express  my  gratitude  to 
my  wife,  Thu.  Although  having  only  re- 
cently arrived  in  the  United  States,  and  with 
an  incomplete  command  of  the  English  lan- 
guage, she  served  as  an  indispensable  full 
time  assistant,  especially  in  the  recording  of 
data.  Subsequently,  she  prepared  the  base 
maps  and  parts  of  the  figures  herein. 

Methods 

As  a  primary  statistical  tool  I  employed 
the  Biomedical  computer  program,  number 
07M,  stepwise  discriminant  analysis  (Dixon, 
1970).  This  method  is  a  modified  version  of 
multivariate  discriminant  function  analysis, 
as  used  previously  in  the  study  of  Canis  and 
explained  in  detail  by  Jolicoeur  (1959),  Giles 
( 1960 ) ,  Lawrence  and  Bossert  ( 1967 ) ,  and 
Gipson  (1972).  The  BMD07M  program  in- 
volves a  procedure  known  as  canonical  analy- 
sis, as  discussed  by  Rao  ( 1952 )  and  Seal 
(1964). 

In  multivariate  analysis  a  series  of  varia- 
bles from  an  individual  specimen  are  consid- 
ered together  to  determine  the  position  of 
that  specimen  relative  to  other  specimens.  In 
its    simplest   form   this    procedure    resembles 


that  of  a  scatter  diagram  in  which  the  loca- 
tion of  a  specimen  on  a  two  dimensional 
graph  is  determined  by  its  position  along 
both  a  vertical  and  horizontal  axis,  each 
representing  a  single  variable.  The  multi- 
variate analysis,  through  a  process  of  matrix 
inversion,  can  consider  numerous  variables, 
but  plots  the  results  in  the  same  form  of  a 
two  dimensional  graph. 

The  BMD07M  program  requires  that  at 
least  two  designated  groups  of  individuals  be 
entered  into  the  analysis.  The  variables  are 
tested  one  at  a  time  for  their  ability  to  dis- 
tinguish between  the  groups.  If  any  variable 
is  found  to  have  too  low  a  discriminatory 
power,  that  variable  is  rejected  and  not  con- 
sidered in  the  analysis.  The  effects  of  cor- 
relation among  the  variables  are  eliminated 
in  this  program  by  a  process  of  eigenvalue 
extraction. 

On  the  basis  of  the  variables  selected,  the 
designated  groups  are  separated  as  well  as  is 
possible.  The  statistical  distance  between 
groups  (D2  of  Mahalonobis),  calculated  from 
the  combined  variables,  may  be  printed  out 
if  desired.  In  addition,  each  individual  speci- 
men is  given  a  D2  distance  from  each  group, 
and  is  assigned  canonical  coordinates  to  plot 
its  position  relative  to  all  other  specimens. 
If  the  variables  employed  have  effectively 
distinguished  the  groups,  the  specimens 
within  a  particular  group  will  be  nearer  to 
each  other  than  to  the  specimens  of  other 
groups.  Once  definite  groups  have  been  es- 
tablished, specimens  of  questionable  identity 
may  be  individually  entered  into  the  analysis 
to  determine  their  position  relative  to  the 
groups  and  hence  their  possible  taxonomic 
affinity. 

For  use  in  multivariate  analysis,  the  15 
measurements  listed  in  appendix  B  were  se- 
lected. These  measurements  were  considered 
to  represent  all  of  the  main  dimensions  of  the 
skull  plus  those  of  three  of  the  more  diag- 
nostic teeth.  Additional  measurements,  es- 
pecially of  the  teeth,  which  are  individually 
of  diagnostic  value,  could  have  been  added. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


But  large  series  of  specimens  were  desirable, 
and  so  measurements  of  parts  too  often 
missing  or  defective  were  excluded.  Also  it 
was  reasoned  that  the  15  utilized  measure- 
ments would  adequately  express  the  major 
functions  of  the  skull.  Because  the  entire 
mandible  was  occasionally  missing  from 
specimens,  measurements  of  the  lower  jaw 
and  teeth  were  omitted  in  multivariate  analy- 
sis. Several  test  runs  of  the  program,  involv- 
ing as  many  as  35  measurements,  including 
those  of  the  mandible,  did  not  seem  to  pro- 
duce results  different  from  those  that  follow, 
nor  to  noticeably  increase  the  discriminatory 
ability  of  the  analysis. 

Lawrence  and  Bossert  (1967)  divided 
each  of  15  measurements  by  greatest  length 
of  skull  and  entered  their  analysis  with  the 
resulting  series  of  fractions,  intending  thereby 
to  eliminate  size  as  a  discriminating  factor. 
Actually  there  are  various  expressions  of  size 
of  a  skull,  and  dividing  by  any  one  of  them 
may  produce  different  results.  Furthermore, 
it  is  questionable  whether  any  attempt  should 
be  made  to  eliminate  the  size  factor,  because 
it  appears  to  be  a  definite  biological  factor, 
at  least  in  distinguishing  the  wild  species  of 
North  American  Canis.  Certain  skulls,  repre- 
senting two  kinds  of  Canis  that  would  not 
ordinarily  be  confused  because  of  size  differ- 
ences, may  have  similar  proportions  of  great- 
est length  to  other  measurements. 

Therefore  raw  measurements  were  used 
in  most  of  the  following  calculations.  This 
procedure  considers  the  size  of  each  meas- 
urement simultaneously  as  a  factor  in  classi- 
fying a  specimen.  Since  the  sizes  of  the 
various  measurements  may  vary  at  different 
rates  between  different  species,  proportion  is 
also  a  factor  in  the  analysis.  It  is  true  that  an 
unusually  large  or  small  specimen  may  be 
assigned  to  a  group  other  than  that  which 
its  proportions  indicate,  but  such  occurrences 
are  rare.  In  any  case,  my  tests  of  this  par- 
ticular computer  program,  using  the  selected 
15  measurements,  revealed  that  in  most  in- 
stances variables  based  on  raw  measurements 


and  fractions  of  greatest  length  of  skull  pro- 
duced similar  depictments  of  relationship, 
but  that  the  raw  measurements  gave  a  wider 
separation  between  groups. 

With  one  major  exception  (dogs,  see  be- 
low), different  analyses  were  used  for  males 
and  females,  and  it  was  found  that  such  a 
procedure  usually  produced  wider  separa- 
tion between  groups  than  was  achieved  by 
combining  sexes.  This  wider  separation  oc- 
curred regardless  of  whether  raw  measure- 
ments or  fractions  of  greatest  length  were 
used  as  variables. 

The  sex  of  some  of  the  skulls  utilized  in 
statistical  analysis  was  unknown,  and  these 
skulls  were  assigned  to  male  or  female  cate- 
gories on  the  basis  of  size  and  the  other 
factors  explained  below.  Fortunately,  ex- 
cepting domestic  dogs,  each  of  the  major 
standard  groups,  against  which  other  material 
was  tested,  consisted  predominantly  of  speci- 
mens of  known  sex.  In  the  subsequent  pages, 
when  a  sample  size  of  one  sex  is  listed,  it  is 
followed  by  the  number  (in  parentheses)  of 
specimens  in  the  series  (if  any)  for  which 
sex  had  not  been  recorded,  but  which  were 
judged  to  belong  to  that  sex. 

In  the  statistical  analyses,  specimens  of 
domestic  dogs  (C.  familiaris)  were  not  sepa- 
rated by  sex.  Dog  skulls  are  poorly  repre- 
sented in  museum  collections,  compared  with 
skulls  of  wild  Canis,  and  less  than  half  of  the 
50  specimens  of  C.  familiaris  used  in  my 
analyses  were  of  known  sex.  Individual  vari- 
ation in  this  species  is  so  great  that  it  tends 
to  obscure  sexual  differences  in  the  morphol- 
ogy of  the  skull.  Consequently,  dogs  of  male, 
female,  and  unknown  sex  were  combined  in 
one  group. 

In  addition  to  multivariate  analyses,  tables 
of  measurements,  with  means,  extremes, 
standard  deviations,  and  coefficients  of  varia- 
tion, are  provided  in  appendix  B.  In  some 
cases  I  also  have  drawn  ratio  diagrams  that 
depict  differences  in  size  and  proportion  be- 
tween the  specimens  of  various  groups. 
Whereas    multivariate    analysis    demonstrates 


6 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


the  collective  results  of  such  differences,  the 
ratio  diagram  permits  visualization  of  how 
each  group  differs  in  individual  measure- 
ments. Simpson  ( 1941 )  explained  this  method 
in  detail  in  the  course  of  his  account  of 
Pleistocene  felines.  Briefly,  raw  statistics  (in- 
dividual measurements,  means  of  a  series, 
etc.),  taken  on  two  or  more  specimens  or 
series,  are  converted  to  their  logarithms.  One 
of  the  specimens  or  series  is  taken  as  a  stand- 
ard, and  the  difference  is  found  between  the 
logs  of  its  individual  measurements  or  means 
and  the  respective  logs  of  the  other  speci- 
mens or  series.  In  diagramming,  the  standard 
values  are  all  plotted  in  a  vertical  line  repre- 
senting the  zero  point,  and  the  respective 
values  of  the  other  specimens  or  series  are 
plotted  at  a  horizontal  distance  from  the 
standard  values,  representing  the  difference 
between  the  two  values. 

Many  skulls  were  examined  for  which  the 
complete  set  of  15  measurements,  required  in 
multivariate  analysis,  could  not  be  obtained 
because  of  damage,  wear,  or  missing  parts. 
Except  for  fossil  material,  data  from  such 
specimens  were  not  incorporated  in  the  ratio 
diagrams  or  statistical  tables  (appendix  B). 
Therefore,  the  groups  represented  in  the  mul- 
tivariate analyses,  ratio  diagrams,  and  tables 
are  all  of  identical  composition.  Specimens 
not  used  for  the  calculation  of  statistics  did 
not  appear  to  differ  from  the  main  series. 

I  finally  want  to  make  it  clear  that  I  used 
multivariate  analysis  in  a  supporting  and 
demonstrative  role,  rather  than  as  a  problem 
solver  in  itself.  The  analysis  did  not  provide 
any  major  conclusions  that  were  not  apparent 
from  more  conventional  methods  of  examina- 
tion, but  it  did  allow  the  efficient  evaluation 
of  many  data,  and  the  objective,  graphical 
portrayal  of  a  complex  situation. 

Age  and  Secondary  Sexual  Variation 

The  aging  process  in  Canis  was  described 
by  Goldman  (1944:400-401),  Jackson  (1951: 
250-251),     Miller,     Christensen     and     Evans 


(1965:652-653).  and  Mech  (1970:139-143). 
Gier  (196S:54-55)  showed  how  to  estimate 
the  age  of  C.  latrans  by  examination  of  wear 
on  the  incisor  and  canine  teeth.  Linhart  and 
Knowlton  (  1967)  demonstrated  a  method  of 
aging  coyotes  through  evaluation  of  cemen- 
tum  layers  in  the  canine  teeth. 

By  the  age  of  six  months  in  Canis,  the 
permanent  dentition,  except  for  the  canine 
teeth,  is  fully  in  place,  and  the  skull  has 
reached  approximately  90  percent  of  its  even- 
tual total  length.  Complete  emergence  of 
the  canines,  and  maximum  dimensions  of  the 
skull,  however,  are  not  attained  until  about 
12  months  in  coyotes  and  15  months  in  larger 
gray  wolves.  Therefore,  for  the  calculation  of 
statistics  in  the  following  sections  of  this 
paper,  I  used  only  skulls  of  animals  estimated 
to  be  at  least  12  months  old,  and  did  not  use 
some  wolves  that  were  under  15  months  old. 
The  males  of  Canis  average  larger  than 
the  females  in  every  measurable  dimension 
of  the  skull,  but  there  is  extensive  overlap 
between  the  two  sexes.  Males  have  propor- 
tionally broader  rostra  and  higher  sagittal 
crests.  In  many  female  coyotes  the  sagittal 
crest  is  flattened,  and  the  temporal  ridges 
that  usually  coalesce  in  males  are  in  some 
females  Urate. 

Statistical  comparison  was  made  of  meas- 
urements of  skulls  of  97  male  and  61  female 
C.  latrans  testes  from  Colorado  and  Idaho, 
and  of  skulls  of  51  male  and  35  female  C. 
lupus  mogollonensis,  youngi  and  irremotus 
from  the  mountainous  region  of  the  western 
United  States.  In  this  particular  test,  only 
specimens  of  known  sex  were  used.  Each 
species  was  examined  separately  and  a  large 
overlap  of  the  two  sexes  was  found  in  all  15 
of  the  measurements  considered.  Males  av- 
eraged larger  in  each  measurement,  however, 
and  analysis  of  variance  and  STP  tests 
showed  a  significant  difference  (p  less  than 
.05)  between  the  males  and  females  of  each 
species  in  all  measurements  except  postorbi- 
tal  constriction  of  braincase. 


HISTORY  AND  STATISTICAL  ANALYSIS 
OF  RECENT  POPULATIONS 

Comparison'  of  Known  Series  of  Dogs,  Wolves,  and  Coyotes 


The  questions  to  be  considered  in  this 
paper  concern  primarily  eastern  North  Amer- 
ica and  fossil  history.  Is  the  red  wolf  of  the 
southeastern  United  States  a  full  species,  a 
subspecies  of  the  gray  wolf,  a  subspecies  of 
the  coyote,  or  a  hybrid  between  C.  lupus 
and  C.  latrans?  What  is  the  origin  and  affin- 
ity of  the  populations  of  Canis  that  recently 
have  become  established  in  much  of  the 
eastern  half  of  the  continent?  Is  hybridiza- 
tion a  major  factor  in  the  situation?  Is  the 
Pleistocene  dire  wolf  completely  distinguish- 
able from  the  modern  gray  wolf,  and  is  it 
possibly   ancestral   to   the  latter? 

Before  attempting  to  answer  these  and 
other  questions,  it  would  be  advisable  to 
delineate  the  perimeters  of  those  populations 
that  seem  best  to  represent  recognizable  spe- 
cies. Throughout  most  of  that  part  of  North 
America  in  which  the  coyote,  gray  wolf,  and 
domestic  dog  are  found  together,  they  are 
easily  distinguishable  and  usually  behave  to- 
ward one  another  as  species.  The  gray  wolf 
once  occurred  in  all  of  North  America  except 
for  parts  of  the  southeastern  United  States, 
most  of  the  state  of  California,  Baja  Califor- 
nia and  the  coastal  lowlands  of  Mexico,  and 
the  region  south  of  central  Mexico,  ( Goldman, 
1944:414).  A  record  of  C.  lupus  baileyi  from 
Tequisistlan,  Oaxaca,  southern  Mexico 
(Goodwin,  1969:224)  seems  to  have  been 
based  on  questionable  evidence.  The  coyote 
was  originally  found  throughout  most  of  the 
western  half  of  the  continent,  and  its  range 
in  the  northeast  extended  as  far  as  the  upper 
Great  Lakes  (Young,  1951:29).  The  domes- 
tic dog,  C.  familiaris,  has  long  occurred  in 
all  parts  of  the  continent,  almost  always  in 
association  with  man.  All  dogs  may  have 
descended  from  a  small  southwest  Asian  sub- 
species of  C.  lupus  that  was  domesticated  10 


to  12  thousand  years  ago  (Scott,  196S).  The 
dogs  of  the  American  Indians  were  appar- 
ently introduced  into  the  New  World  by  man, 
and  do  not  seem  to  have  been  influenced  by 
interbreeding  with  native  species  of  North 
American  Canis  (Allen.  1920;  Haag,  1948). 
Specimens  of  the  earliest  known  domestic 
dogs  on  the  continent  were  described  from  a 
site  in  Lemhi  County,  Idaho  dated  at  10,400- 
11,500  B.P.  They  reportedly  already  possess 
the  typical  characters  of  C.  familiaris  (Law- 
rence, 1966,  1968). 

Only  in  the  eastern  part  of  North  America 
do  hybridization  and  modification  of  the  orig- 
inal populations  appear  to  be  of  possible 
significance.  A  few  isolated  instances  in 
which  C.  familiaris  hybridized  with  either 
C.  lupus  or  C.  latrans  in  other  regions,  have 
been  reported  (Young,  1944:180-210;  Men- 
gel,  1971;  Gray,  1972),  but  such  cases  do  not 
seem  to  have  had  lasting  effect  on  popula- 
tions. No  instances  of  interbreeding  between 
C.  lupus  and  C.  latrans  in  the  western  half 
of  the  continent  have  yet  been  reported. 
Therefore,  it  is  reasonable  to  consider  C. 
lupus  and  C.  latrans  of  the  western  and 
northern  parts  of  the  continent  as  consisting 
of  natural,  unmodified  populations  that  may 
confidently  be  used  as  a  basis  on  which  to 
test  more  questionable  populations. 

For  an  initial  analysis  of  known  groups,  I 
decided  upon  using  skulls  of  C.  latrans  and 
C.  lupus  that  had  been  collected  not  later 
than  1925  in  the  mountainous  region  of  the 
west.  The  gray  wolf  sample  included  57(6) 
males  and  37(2)  females  (parentheses  con- 
tain numbers  of  specimens  in  the  series  for 
which  sex  had  not  been  recorded,  but  which 
were  judged  to  belong  to  the  particular  sex 
indicated;  see  p.  5).  This  group  consisted 
of  all  skulls  of  adult  C.  lupus  mogollonensis, 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


ijoungi,  and  irrcmotus  in  the  U.S.  National 
Museum  of  Natural  History,  upon  which  the 
15  necessary  measurements  could  be  made, 
except  for  two  specimens  taken  after  1925 
(see  appendix  A,  part  1). 

Of  coyotes,  97  male  and  61  female  skulls, 
all  of  the  subspecies  C.  latrans  lestes,  were 
utilized  in  the  initial  test  (see  appendix  A, 
part  2).  These  specimens  comprised  the  en- 
tire National  Museum  collection  of  Idaho  and 
Colorado  adult  lestes  of  known  sex,  taken 
prior  to  1926,  except  for  skulls  upon  which 
all  of  the  needed  15  measurements  could  not 
be  made. 

The  selection  of  these  particular  speci- 
mens as  standard  comparative  material  had 
the  following  advantages :  ( 1 )  the  wolf  and 
coyote  had  long  been  sympatric  in  the  region, 
and  thus  theoretically  would  have  evolved 
the  maximum  amount  of  differential  charac- 
ters reflecting  their  separate  ecological  niches; 
(2)  the  region  had  a  minimum  human  (and 
presumably  domestic  dog)  population;  (3) 
the  time  period  was  one  in  which  both  the 
wolf  and  coyote  were  common  (most  of  the 
specimens  represent  the  first  few  years  of 
Federal  predator  control  work  which  began 
in  1915);  and  (4)  the  subspecies  of  both  C. 
lupus  and  C.  latrans  do  not  exhibit  extremes 
of  size  or  other  characters  within  their  re- 
spective  species. 

For  a  sample  of  domestic  dogs,  only  those 
skulls  were  selected  which,  while  known  to 
be  C.  familiaris,  were  superficially  nearest  to 
those  of  C.  lupus  or  C.  latrans  in  appearance. 
The  extremes  of  domestication  represented 
by  broad-skulled  dogs  (as  bulldogs),  narrow- 
skulled  dogs  (as  Russian  wolfhounds),  and 
dogs  having  greatly  reduced  rostra  (as  pugs) 
were  avoided.  Extremely  small  dogs,  those 
in  which  the  skull  was  less  than  150  milli- 
meters in  greatest  length,  also  were  not  used. 
Specimens  utilized  in  the  sample  included  1 
Eskimo  dog,  5  Irish  wolfhounds,  3  German 
shepherds,  2  sheep  dogs,  2  Newfoundlands. 
1  doberman  pinscher,  1  greyhound,  1  great 
Dane,    1   mastiff,    1   Irish   setter,    1   beagle,    1 


-   LATRANS 


LUPUS 


FAMILIARIS 


— I 1 r- 

-4    -3    -2 


-i 1 1- 

2      3      4 


5 
4 
3 
2 
I 

0 
-I 
-2 
-3 
-4 
-5 
-6 


LATRANS 


LUPUS 


FAMILIARIS 


■3    -2 


3     4       5      6 


Fig.  1. — Graphical  results  of  multivariate  analy- 
ses comparing  samples  of  C.  lupus  and  C.  latrans 
from  the  mountainous  region  of  western  North  Amer- 
ica, and  C.  familiaris.  Only  the  margins  of  the  range 
of  variation  of  each  species  are  shown.  In  this  and 
in  all  subsequent  portrayals  of  multivariate  analyses, 
the  numbers  along  the  vertical  and  horizontal  axes 
are  canonical  coordinates.  These  coordinates  are  used 
to  indicate  relative  position,  and  do  not  represent 
any  material  values.  In  this  figure  and  subsequent 
portrayals  of  analyses,  males  are  shown  above  and 
females  are  shown  below. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


Fig.  2. — Range  map  of  C.  lupus  in  North  America  showing  localities  (black  dots)  of  specimens  used  in  the 
statistical  analyses  of  this  paper.    The  numbers  on  the  map  represent  recognized  subspecies,  as  follows: 


1.  C.  I.  alces 

2.  C.  I.  arctos 

3.  C.  Z.  baileyi 

4.  C.  I.  beothucus 

5.  C.  I.  bernardi 

6.  C.  /.  columbianus 


7.  C.  I.  crassodon 

8.  C.  I.  fuscus 

9.  C.  /.  hudsonicus 

10.  C.  /.  griseoalbus 

11.  C.  ?.  irremotus 

12.  C.  /.  labradorius 


13.  C.  /.  Zigoni 

14.  C.  /.  lycaon 

15.  C.  Z.  mackenzii 

16.  C.  /.  manningi 

17.  C.  /.  mogollorwnsis 

18.  C.  /.  monstrabilis 


19.  C.  Z.  nubilus 

20.  C.  Z.  occidentalis 

21.  C.  Z.  orion 

22.  C.  Z.  pamba-sileus 

23.  C.  Z.  tundrarum 

24.  C.  Z.  youngi 


The  solid  lines  indicate  subspecific  boundaries.  The  dashed  line  in  southeastern  Ontario  shows  Standfield's 
(1970)  division  between  his  "Ontario  type"  and  "Algonquin  type"  of  C.  lupus  lycaon.  Because  of  the  scale 
of  the  map,  it  was  not  possible  to  plot  all  localities  in  crowded  areas. 


10 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


scale  of  miles 


Fig.  3. — Range  map  of  C.  latrans  showing  localities  of  specimens   used  in  the   statistical  analyses   of  this 
paper.    The  numbers  on  the  map  represent  recognized  subspecies,  as  follows: 

1.  C.  I.  cagottis  7.  C.  /.  impavidus  13.  C.  /.  microdon 

2.  C.  I.  clepticus  8.  C.  /.  incolatus  14.  C.  I.  ochropus 

3.  C.  /.  dickeyi  9.  C.  I.  jamesi  15.  C.  /.  pcninsulae 

4.  C.  Z.  frustror  10.  C.  /.  latrans  16.  C.  /.  f<?xen.rfs 

5.  C.  /.  goldmani  11.  C.  /.  /c.sres  17.  C.  /.  thamnos 

6.  C.  /.  hondurensis  12.  C.  /.  mearnsi  18.  C.  /.  umpquensis 

The  solid  lines  indicate  subspecies  boundaries.    Because  of  the  scale  of  the  map,  it  was  not  possible  to  plot  all 
localities  in  crowded  areas.    Certain   additional  localities  of  C.  latrans  are  shown  in  Figs.  14,  25,  and  31. 


19.  C.  /.  vigilis 

20.  C.  I.  "var."  (Law- 
rence and  Bossert, 
1969) 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


11 


basset  hound,  and  30  dogs  of  unknown  or 
mixed  breed  (see  appendix  A,  part  3).  Of 
these  50  specimens,  only  11  males  and  9  fe- 
males had  been  previously  identified  as  to 
sex.  For  the  reasons  explained  above,  do- 
mestic dogs  of  male,  female,  and  unknown 
sex  were  combined  into  a  single  sample. 

The  graphical  results  of  multivariate  anal- 
yses of  males  and  females  of  the  initial  sam- 
ples are  depicted  in  figure  1.  For  both  sexes 
there  is  complete  separation  between  all 
three  species.  Such  an  arrangement  could 
be  expected,  and  it  may  serve  as  a  sound 
basis  on  which  to  evaluate  other  specimens. 

The  next  step  was  to  compare  skulls  taken 
elsewhere  in  northern  and  western  North 
America  to  the  above  series  of  reliably  dis- 
tinguished specimens.  Skulls  of  176(27) 
males  and  114(33)  females,  previously  identi- 
fied as  C.  lupus,  and  of  69(2)  males  and 
50(4)  females,  identified  as  C.  latrans,  were 
tested  individually  against  the  three  known 
groups  (see  appendix  A,  parts  4  and  5).  The 
maps  in  figures  2  and  3  show  localities  of  all 
specimens  of  gray  wolves  and  coyotes.  Fig- 
ure 4  shows  the  results  of  multivariate  analy- 
ses. Nearly  all  of  the  newly  added  material 
falls  within  the  range  of  variation  of  the  ap- 
propriate original  sample,  or  at  least  is  closer 
to  this  range  than  to  that  of  other  species. 

Only  five  of  these  specimens  (see  appen- 
dix A,  part  6)  seem  confusing  as  to  identity. 
Three  skulls,  previously  identified  as  C.  lupus 
baileyi,  are  statistically  and  morphologically 
intermediate  to  known  samples  of  female  gray 
wolves  and  coyotes.  Canis  lupus  baileyi,  the 
smallest  subspecies  of  North  American  gray 
wolf,  shared  its  entire  range  with  C.  latrans, 
and  interbreeding  between  the  two  might 
have  been  possible  under  certain  conditions. 
I  thus  consider  these  three  specimens  as  prob- 
able hybrids  of  C.  lupus  and  C.  latrans,  and 
henceforth  have  not  used  them  in  the  forma- 
tion of  samples  of  either  parent  species. 

Two  females  from  the  Sacramento  Moun- 
tains of  New  Mexico,  previously  identified  as 
C.   lupus  monstrabilis,   appear   both   visually 


Fig.  4. — Statistical  positions  of  individual  native, 
\vi!d-caught  specimens  from  throughout  northern  and 
western  North  America,  relative  to  ranges  of  variation 
of  the  series  of  C.  lupus,  C.  latrans,  and  C.  familiaris 
shown  in  Fig.  1.  The  black  dots  represent  individuals 
of  C.  lupus  and  C.  latrans,  all  of  which  fall  close  to 
the  appropriate  range  of  variation.  The  letter  B 
represents  specimens  throught  to  be  hybrids  between 
C.  lupus  baileyi  and  C.  latrans;  the  letter  M  repre- 
sents specimens  thought  to  be  hybrids  between  C. 
lupus  monstrabilis  and  C.  familiaris.  Males  are  above, 
females  below;  numbers  along  axes  are  canonical  co- 
ordinates. 


12 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


LUPUS 


4 
3 
2 
I 

0 
-I 
-2 
-3 
-A 
-5 
-6 
-7 
-8 


FAMILIARIS 


-5-4-3-2-IOI2345 

Fie.  5. — Multivariate  comparison  of  all  specimens 
of  C.  lupus  and  C.  latrans  from  northern  and  western 
North  America,  and  C.  familiaris.  Only  the  margins 
of  the  range  of  variation  of  each  species  are  shown. 
Males  are  above,  females  below. 

and  statistically  (Fig.  4)  to  be  intermediate 
to  the  gray  wolf  and  domestic  dog.  At  least 
two  other  skulls   collected  in  New  Mexico, 


but  not  complete  enough  for  inclusion  in  the 
multivariate  analysis,  also  suggest  the  oc- 
currence of  hybridization  between  the  two 
species.  Nonetheless,  the  preponderance  of 
material  that  can  be  clearly  identified  as 
either  C.  latrans  or  C.  lupus,  indicates  that  in 
those  regions  hitherto  discussed  the  relation- 
ships of  these  two  canids  to  one  another  and 
to  C.  familiaris  were  those  of  normal  species. 
All  of  the  northern  and  western  speci- 
mens, except  the  five  considered  to  be  hy- 
brids, were  incorporated  with  the  appropriate 
standard  samples  of  C.  lupus  or  C.  latrans, 
and  these  two  groups  along  with  the  sample 
of  50  domestic  dogs  were  tested  in  new  multi- 
variate analyses.  The  graphical  results  de- 
picted in  figure  5  once  again  indicate  clear 
separation  between  the  three  species.  Meas- 
urements for  the  total  series  of  dogs,  and  of 
western  and  northern  wolves  and  coyotes  are 
listed  in  appendix  B  (parts  1  and  2).  The 
means  of  these  measurements  (of  males 
only  for  C.  lupus  and  C.  latrans)  are  com- 
pared in  the  ratio  diagram  in  figure  6. 

Systematic  Problems  in 
the  Northeast 

Decline  of  the  Gray  Wolf 

According  to  Goldman  ( 1944 ) ,  a  single 
subspecies  of  gray  wolf,  C.  lupus  lycaon,  orig- 
inally occupied  the  region  from  eastern  Min- 
nesota to  the  Atlantic,  and  from  northern 
Ontario  to  parts  of  the  southeastern  United 
States.  Considering  the  enormity  of  this 
range,  however,  and  the  problems  associated 
with  the  systematics  of  Canis  in  eastern  North 
America,  Goldman  used  relatively  few  speci- 
mens for  describing  the  situation.  From  the 
entire  region  south  of  Lakes  Michigan  and 
Erie  he  assigned  to  lycaon  only  four  complete 
skulls  and  one  mandibular  ramus. 

Standfield  ( 1970 )  reported  the  presence 
of  two  distinct  kinds  of  lycaon  in  Ontario, 
which  he  designated  the  "Ontario  type"  and 
the  "Algonquin  type."  The  former  was  said 
to  occur  mainly  in  the  boreal  forests  north 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


13 


-20      -.16        -.12      -  08       -.04        0 
1 1 — i 1 


-i 1 1 1 1 r 

LATRANS 


LUPUS 


FAMILIARIS 


.70 


80 


90 


100 


Fig.  6. — Ratio  diagram  comparing  means  of  total 
samples  of  C.  lupus  and  C.  latrans  from  northern  and 
western  North  America  (males  only  are  shown),  and 
C.  familiaris  (dashed  line).  Vertically  arranged  num- 
bers represent  the  measurements  so  numbered  in  Ap- 
pendix B.  A  log  difference  scale  is  provided  above, 
and  a  ratio  scale  below  the  diagram. 

and  northeast  of  Lake  Superior,  to  be  larger, 
and  to  vary  in  pelage  from  pure  white  to  jet 
black.  The  latter  was  said  to  dccur  in  the 
deciduous  forests  east  and  southeast  of  Lakes 
Superior  and  Huron,  to  be  smaller,  and  to  be 
invariably  gray-brown.  Kolenosky  and  Stand- 
field  (1975)  made  a  multiple  discriminant 
analysis  comparing  105  skulls  of  the  "Algon- 
quin type"  and  122  skulls  of  the  "Ontario 
type"  (now  referred  to  as  the  "Boreal  type"), 
and  concluded  that  there  were  significant 
differences  between  the  two.  These  authors 
(p.  71)  reported  that  "the  ranges  of  the  two 
types  overlap  throughout  a  broad  band 
across  eastcentral  Ontario,  but  there  is  no 
conclusive   evidence   of   their   interbreeding." 


Mech  and  Frenzel  ( 1971 )  suggested  that 
the  present  population  of  wolves  in  north- 
eastern Minnesota  consists  at  least  in  part  of 
the  subspecies  C.  lupus  nubilus.  This  idea 
was  based  on  their  observations  of  black 
wolves  and  white  wolves  in  the  area.  Indi- 
viduals of  these  colors  were  common  among 
nubilus,  but  were  not  reported  in  cited  ob- 
servations of  lycaon  in  eastern  Ontario. 

Whatever  their  original  systematic  status, 
northeastern  wolves  have  suffered  a  drastic 
loss  in  numbers  and  range  because  of  perse- 
cution by  Caucasian  man.  The  species  prac- 
tically disappeared  south  of  the  St.  Lawrence 
River  between  1850  and  1900  (Peterson, 
1966:200;  Goodwin,  1936),  and  was  gone 
from  southern  Michigan  and  Wisconsin  early 
in  the  twentieth  century  (Jackson,  1961:293; 
Arnold,  1952).  Wolves  were  reported  to  be 
present  in  moderate  numbers  in  the  upper 
peninsula  of  Michigan  by  Stebler  ( 1944 ) , 
and  in  northern  Wisconsin  by  Schorger 
(1942).  In  the  1950"s,  however,  the  wolf 
populations  of  these  two  states  declined 
sharply.  Jackson  (1961:293)  estimated  50 
wolves  to  be  present  in  northern  Wisconsin, 
and  Keener  (1970)  reported  that  none  still 
survived  in  the  area.  For  the  upper  penin- 
sula of  Michigan,  recent  numerical  estimates 
have  been  20  (Smits,  1963),  less  than  12 
(Douglass,  1970),  and  about  six  (Hendrick- 
son  and  Robinson,  1975).  In  March  1974 
four  wolves  from  northern  Minnesota  were 
released  in  upper  Michigan,  but  by  Septem- 
ber all  had  been  killed  through  human 
agency  (Weise,  et  o/.,  1975).  A  viable  group 
of  about  20  to  30  individuals,  however,  has 
maintained  itself  on  Isle  Royale  in  Lake 
Superior  since  the  late  1940's  (Mech,  1966; 
Wolfe  and  Allen,  1973). 

The  only  major  population  of  C.  lupus 
to  be  found  anywhere  in  the  United  States 
south  of  Canada  is  that  in  northern  Minne- 
sota. An  estimated  1,000  to  1,200  individuals 
are  reported  to  exist  in  the  area  and  they  are 
said  to  be  in  no  immediate  danger  of  extir- 
pation (Mech,  1977). 


14 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


Despite  intensive  control  measures,  wolves 
reportedly  still  occupy  all  of  the  forested 
parts  of  Ontario,  even  areas  close  to  Toronto 
and  Ottawa.  They  do  not,  however,  enter 
the  settled  agricultural  sections  of  that  prov- 
ince (Clarke,  1970).  Here  and  in  other  parts 
of  the  northeast,  there  evidently  has  been  a 
correlation  between  the  decline  of  the  gray 
wolf  and  the  intensity  of  human  population 
and  agricultural  development. 

Rise  of  the  Coyote 

At  the  time  the  first  white  settlers  arrived 
in  North  America,  coyotes  were  apparently 
confined  to  open  plains  and  more  arid  re- 
gions, mainly  in  the  western  half  of  the  con- 
tinent (Young,  1951;  Seton,  1929).  The 
original  range  of  the  species  did,  however, 
follow  the  prairie  peninsula  through  the  mid- 
western  states,  at  least  as  far  as  northeastern 
Indiana  (Mumford,  1969:85).  According  to 
Jackson  (1961:285)  coyotes  were  undoubt- 
edly present  in  southern  Wisconsin  when  the 
early  explorers  arrived,  and  a  few  may  have 
inhabited  the  northern  part  of  the  state.  The 
name  C.  latrans  thamnos  was  applied  by 
Jackson  ( 1949 )  to  the  coyote  of  the  north- 
eastern portion  of  the  range  of  the  species. 

Sometime  after  the  middle  of  the  nine- 
teenth century,  coyotes  began  appearing  to 
the  north,  east,  and  south  of  the  prairies,  and 
by  the  mid-twentieth  century  they  existed  in 
large  numbers  beyond  their  original  range 
(De  Vos,  1964;  Mech,  1959,  1961;  Young, 
1951).  Many  coyotes  had  escaped  from  cap- 
tivity, or  had  been  deliberately  released  by 
sport  or  bounty  hunters.  Some  of  these  ani- 
mals formed  local  breeding  populations  that 
maintained  themselves  over  a  period  of  time. 
Records  that  seem  attributable  to  such  intro- 
ductions are  as  follows. 

Florida.— Palm  Beach  County,  1925;  Col- 
lier, Monroe,  and  Marion  counties  (Young, 
1951:15);  DeSoto  County,  1933  (Sherman, 
1937);  Polk  County,  1962  (Cunningham  and 
Dunford,  1970). 


Alabama.— Barbour  County,  1924-1929 
(Young,  1951:16);  Bazemore,  Fayette  Coun- 
ty, 1956;  Huntsville  Arsenal,  Madison  County, 
1961   (Holliman,  1963). 

Georgia.— North  Georgia,  1929;  Haber- 
sham County,  1930;  Ware  County  (Young, 
1951:15). 

South  Carolina. — Aiken  County;  Edisto 
Island,  Colleton  County,  1924  (Golley,  1966: 
141). 

North  Carolina.— Gaston  County,  1938 
(Young,  1951:14);  Swain  County,  1947  (Lin- 
zey  and  Linzey,  1968). 

Tennessee. — Grand  Junction,  Hardeman 
County;  Maury  County,  1930;  McCains, 
Maury  County,  1931  (Kellogg,  1939:267); 
Hickman  and  Maury  counties,  early  1930's 
(Young,  1951:15);  Benton,  Hickman,  and  Se- 
quatchie counties  (Schultz,  et  ah,  1954:205); 
Tennessee  National  Wildlife  Refuge,  Henry 
County,  1951  (Schultz,  1955). 

Kentucky. — Near  Fayette-Clark  county 
line,  1953  (Gale  and  Pierce,  1954). 

Virginia. — Rockingham,  Highland,  and 
Grayson  counties  (Handley  and  Patton,  1947: 
140);  Tazewell  and  Lee  counties,  1968-1969; 
near  Mossy  Creek,  Augusta  County,  1970 
(Carpenter,   1971). 

West  Virginia. — Tucker  County  ( Hand- 
Ian,  1946). 

Maryland. — 5  mi.  NW  Poolsville,  Mont- 
gomery County,  1921  (Jackson,  1922);  out- 
skirts of  Baltimore,  1931  (Redington,  1931: 
27);  Cecil  County,  1961  (Paradiso,  1969: 
134). 

New  Jersey. — Near  Ringoes,  Hunterdon 
County,  1938  (Young,  1951:16);  near  Fishing 
Creek,  Cape  May  County,  1948  (Ulmer, 
1949). 

Pennsylvania. — Clinton  County,  1915-1916 
(Shoemaker,  1917:11);  near  Flowing  Spring, 
Blair  County,  1907  (Gifford  and  Whitebread, 
1951:46);  Sheshequin  Township,  Bradford 
County,  1939;  Chestnut  Hill  section,  Phila- 
delphia, 1942  (Ulmer,  1949);  Beaver  and 
Forest  counties,  1946-1947  (Richmond  and 
Rosland,  1949:34). 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


15 


New  York. — Near  Ithaca,  Tompkins  Coun- 
ty, 1920  (Seton,  1929:369);  Ontario  County, 
about  192S  (Rump,  1941:415). 

Massachusetts. — Vicinity  of  Amherst, 
Hampshire  County,  1936  (Warfel,  1937). 

Maine.— Near  Portland,  1932  (Aldous, 
1939). 

Some  other  occurrences  in  New  England 
and  New  York  probably  represent  range  ex- 
tensions and  are  discussed  below.  But  it  is 
difficult  to  determine  whether  some  records 
should  be  designated  as  introductions  or 
range  extensions. 

The  actual  eastward  extension  of  the  coy- 
ote's range  has  seemingly  been  slowest  in  the 
region  between  Michigan  and  the  Ohio  Val- 
ley, possibly  because  of  the  intensity  of  hu- 
man population  there.  Some  sources,  such  as 
Seton  (1929:368)  even  suggest  that  the  coy- 
ote had  been  exterminated  in  part  of  this 
region,  although  Mumford  (1969:85)  doubt- 
ed that  the  species  had  ever  been  extirpated 
in  Indiana.  Jackson  (1961:285)  noted  a  re- 
versal of  the  original  situation  in  Wisconsin, 
in  that  coyotes  had  become  much  more  com- 
mon in  the  northern  half  of  the  state  than  in 
the  southern  part.  Burt  ( 1946 )  observed  that 
the  species  was  rare  in  the  southern  part  of 
the  lower  peninsula  of  Michigan.  Hoffmeister 
and  Mohr  (1957:119)  stated:  "The  coyote 
occurs  in  much  of  Illinois,  but  it  is  not  com- 
mon anywhere  in  the  state."  And  Mumford 
( 1969 :  84 )  reported  coyotes  to  -  be  present 
throughout  Indiana,  but  not  to  be  common. 
Hamilton  (1943:178)  indicated  that  coyotes 
were  of  sporadic  occurrence  in  most  of  west- 
ern Ohio,  and  that  they  had  been  established 
in  Logan  County  for  12  years.  Other  occur- 
rences in  the  western  counties  of  Ohio  were 
reported  by  Negus  (1948),  Whitacre  (1948), 
Young  (1951:15),  and  Goodpaster  and  Hoff- 
meister (1968).  Wilson  (1976)  mentioned 
the  presence  of  a  sparse  coyote  population  in 
western  Kentucky. 

The  coyote's  range  seems  to  have  ex- 
panded mainly  to  the  northeast  of  the  orig- 


inal distribution  in  the  prairie  peninsula.  At 
the  base  of  the  lower  peninsula  of  Michigan, 
individuals  were  reported  in  Rerrien  County 
in  1900-1901  (Wood  and  Dice,  1924)  and  in 
Washtenaw  County  in  1905  and  1910  (Wood, 
1922).  A  specimen  was  taken  in  Genesee 
County,  in  the  east-central  part  of  the  state 
in  1917  ( Wood  and  Dice,  1924 ) ,  and  an  indi- 
vidual was  reported  in  Charlevoix  County, 
at  the  northern  tip  of  the  lower  peninsula,  in 
1919-1921    (Dice,   1925). 

Coyotes  had  also  entered  the  upper  penin- 
sula of  Michigan  by  the  early  twentieth  cen- 
tury. Shiras  (1921:166)  reported:  "In  the 
past  fifteen  years  the  coyote  unexpectedly  ap- 
peared in  northern  Wisconsin  and  Michigan, 
coming  from  Minnesota.  It  has  since  become 
very  numerous."  Wood  and  Dice  (1924) 
listed  occurrences  in  five  upper  peninsula 
counties  between  1912  and  1915. 

Even  earlier,  coyotes  had  begun  to  move 
north  in  Minnesota.  According  to  Bailey 
( 1929 )  they  first  appeared  in  Sherburne 
County  in  1875,  and  Calm  (1921)  reported 
them  to  be  more  common  than  wolves  in 
Itasca  County. 

Snyder  (1938)  thought  that  coyotes  were 
present  in  the  western  Rainy  River  District, 
Ontario  in  1S90.  Peterson  (1966:197)  noted 
that  in  Ontario  before  1900  the  species  was 
restricted  to  Rainy  River  and  western  Kenora 
districts.  Krefting  (1969)  suggested  that  coy- 
otes moved  to  Isle  Royalc  in  Lake  Superior 
sometime  prior  to  1912-1913,  from  the  Sibley 
Peninsula  area  of  Ontario,  where  they  had 
arrived  about  1900.  Coyotes  continued  to  in- 
habit Isle  Royale  until  gray  wolves,  in  a  re- 
versal of  the  general  trend  in  the  northeast, 
occupied  the  island  in  the  1940's. 

From  western  Ontario,  coyotes  spread 
eastward  above  Lake  Superior,  and  northward 
toward  Hudson  Ray.  They  also  apparently 
crossed  the  St.  Clair  River  into  southeastern 
Ontario  where  the  first  specimen  was  taken, 
north  of  Thedford.  Lambton  County,  in  1919. 
In  1943  specimens  were  collected  in  Essex, 
Peterborough,  and  Carleton  counties,  Ontario 


16 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


(Anderson,  1946),  and  by  1956  the  species 
was  reported  to  occur  nearly  throughout  the 
province  (Peterson,  1957).  The  first  record 
for  Quebec  was  a  specimen  taken  near  Lusk- 
ville,  Gatineau  County  in  1944  (Rand,  1945). 
According  to  Wolfram  (1964)  the  first  coyote 
to  be  found  in  New  Brunswick  was  killed 
near  Sussex  in  195S.  Subsequent  expansion  of 
the  species  in  Quebec,  as  far  as  the  Gaspe 
Peninsula,  and  in  New  Brunswick,  was  re- 
ported by  Georges  ( 1976 ) . 

Bromley  (1956)  summarized  the  history 
of  the  coyote  in  northern  New  York.  Indi- 
viduals were  shot  in  the  St.  Lawrence  River 
area  in  1925  and  in  Franklin  County  in  the 
mid-1930's.  The  species  was  said  to  have 
achieved  a  good  foothold  in  the  early  1940's 
when  its  range  included  several  areas  of  the 
Adirondack  Mountains.  By  the  late  1940's 
coyotes  had  spread  throughout  the  Adiron- 
dacks,  and  in  the  early  1950's  they  occupied 
the  entire  northern  third  of  the  state.  Subse- 
quently, according  to  Severinghaus  ( 1974a, 
1974b),  there  have  been  records  from  much 
of  southern  New  York.  An  estimate  of  5,000- 
15,000  coyotes  for  the  state  was  published  by 
Marvinney  (1976). 

Coyotes  apparently  are  continuing  to 
move  down  the  Appalachian  Mountains  from 
the  northeast.  They  are  currently  reported  to 
be  rare,  but  widely  distributed  in  Pennsyl- 
vania (John  L.  George,  Department  of  Wild- 
life Management,  Pennsylvania  State  Univer- 
sity, pers.  comm. ),  and  their  "sporadic  pres- 
ence" in  West  Virginia  was  discussed  by 
Taylor,  Counts,  and  Mills  (1976).  In  May 
1976  a  specimen  was  taken  near  Nestorville. 
Barbour  County  in  northeastern  West  Vir- 
ginia, and  I  found  the  skull  to  closely  resem- 
ble those  of  some  New  England  coyotes. 

In  1936  a  coyote  reportedly  was  killed  in 
Argyle  Township,  Penobscot  County,  Maine. 
Over  the  next  two  years  11  other  individuals 
were  taken  in  that  vicinity,  most  of  which 
were  considered  to  be  hybrids  between  C. 
latrans  and  C.  familiaris  (Aldous,  1939).  A 
specimen    identified    as    C.    latrans    thamnos 


was  taken  in  Lower  Enchanted  Township, 
Somerset  County,  Maine  in  1961  (Carson, 
1962).  The  subsequent  occupation  of  nearly 
the  entire  state  by  the  species  has  been  docu- 
mented by  Richens  and  Hugie  (1974)  and 
Teer  (1975).  The  Maine  Department  of  In- 
land Fisheries  and  Wildlife  (1976)  has  pub- 
lished an  estimate  of  from  1,500  to  5,500 
coyotes  in  the  state. 

Coyotes  were  first  reported  in  Vermont  in 
1942  and  have  since  been  taken  in  all  14 
counties  of  the  state.  The  first  record  in  New 
Hampshire  was  one  shot  near  Holdemess, 
Grafton  County  in  1944,  and  the  first  in 
Massachusetts  was  shot  near  Otis,  Berkshire 
County  in  1957.  Three  more  were  trapped  in 
Massachusetts  on  the  Prescott  Peninsula  of 
Quabbin  Reservoir  in  1958,  and  one  was  shot 
near  Grafton,  Worcester  County  in  1959.  Four 
individuals,  believed  to  be  coyotes  or  hybrids 
between  C.  latrans  and  C.  familiaris  ("coy- 
dogs"),  were  taken  in  western  Connecticut 
from  1957  to  1963  (Pringle,  1960,  1963). 
Silver  and  Silver  (1969:Fig.  30)  depicted  oc- 
currences throughout  Vermont.  New  Hamp- 
shire, and  Massachusetts. 

Difficulties  in  Identifying  Northeastern  Canis 

Although  the  recent  presence  of  wild 
Canis  in  the  northeast  does  appear  to  repre- 
sent primarily  an  extension  of  the  range  of 
the  species  C.  latrans,  the  exact  identity  of 
certain  individuals  and  populations  has  been 
open  to  question.  Perhaps  because  coyotes 
had  never  before  occurred  in  the  region,  their 
initial  appearance  was  something  of  a  mys- 
tery and  the  cause  of  excitement  among  the 
public  and  local  wildlife  officials.  Some  per- 
sons believed  that  wolves  were  returning  to 
areas  in  which  they  had  been  exterminated 
long  before,  while  others  considered  the  new 
canids  to  be  wild  dogs  or  coy-dogs.  And  it 
is  likely  that  each  of  these  three  kinds  of  ani- 
mals contributed  in  some  part  to  the  mystery. 
An  actual  specimen  of  C.  lupus  was  killed  in 
Fulton  County,  New  York  in  1968  (Paradiso 
and    Schierbaum,    1969).     Another    wolf,    of 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


17 


unsuaUy  large  size  and  probably  not  a  native 
animal,  was  killed  north  of  Kemptville,  Carle- 
ton  County,  Ontario,  about  30  miles  from  the 
New  York  border,  in  1962.  According  to 
Bump  (1941)  an  undetermined  number  of 
gray  wolves  escaped  from  captivity  in  south- 
ern Franklin  County,  New  York  about  1930. 
Silver  and  Silver  (1969)  said  that  four  gray 
wolves  were  imported  to  the  vicinity  of  Croy- 
don, Sullivan  County,  New  Hampshire,  and 
that  after  the  last  died  in  1914  there  were 
regular  reports  of  wolflike  animals  in  that 
area. 

Completely  feral  dogs  seem  to  be  uncom- 
mon, but  do  occur  on  occasion,  and  are  some- 
times the  cause,  directly  or  indirectly,  of  re- 
ported wolves  and  coyotes.  Carson  (1962) 
referred  to  several  cases  of  C.  familiaris  living 
and  breeding  in  the  wild  in  Maine.  McKnight 
(1964:48)  reported  that  a  pack  of  wild  dogs 
inhabited  a  den  near  Hopkinton,  New  Hamp- 
shire for  six  years  prior  to  1959.  And  Nesbitt 
( 1975 )  made  a  five-year  study  of  a  feral  pack 
on  Crab  Orchard  National  Wildlife  Refuge, 
Illinois. 

Aldous  (1939)  and  Carson  (1962)  re- 
ferred to  the  taking  of  numerous  coy-dogs  in 
Maine.  Wetzel  and  Penner  (1962)  reported 
the  collection  of  two  specimens  of  coy-dogs 
in  Litchfield  County,  Connecticut.  Cook 
(1952)  discussed  the  presence  of  such  ani- 
mals in  New  York.  Wolfram  (1964)  cited  a 
report  that  20  percent  of  the  coyotes  in  On- 
tario were  actually  coy-dogs.  Paul  (1970) 
stated  that  20  percent  of  the  carcasses  of  wild 
Cards  found  in  Illinois  were  identified  as  coy- 
dogs.  In  tracing  the  spread  of  coyotelike 
animals  in  New  York,  Severinghaus  ( 1974a, 
1974b)  noted  that  the  first  litters  in  a  newly 
occupied  area  were  almost  always  obvious 
coy-dogs. 

Such  records  apply  to  wild-caught  animals 
that  were  presumed  to  be  hybrids  between 
C.  latrans  and  C.  familiaris  on  the  basis  of 
morphological  characters.  Cases  in  which 
known  hybrids  were  born  in  captivity  under 
controlled  conditions  were  discussed  by  Dice 


(1942),  Young  (1951:123),  Kennelly  and 
Roberts  (1969),  Gier  (1968),  and  Mengel 
( 1971 ) .  The  latter  three  of  these  authors 
reported  such  hybrids  to  be  fertile. 

Mengel  (1971)  reviewed  the  subject  of 
hybridization  between  C.  latrans  and  C.  fa- 
miliaris, and  presented  information  on  his 
own  experiments.  He  noted  that  whereas 
coyotes  normally  mate  from  late  January  to 
March,  and  usually  give  birth  in  the  spring, 
coy-dogs  have  been  observed  to  mate  from 
October  to  December,  and  to  give  birth  in 
the  winter.  This  phase  shift  in  the  breeding 
cycle  was  held  to  be  a  barrier  restricting  the 
interbreeding  of  coy-dogs  with  C.  latrans, 
and  hence  preventing  the  introgression  of 
domestic  dog  genes  into  the  wild  coyote  pop- 
ulation. Furthermore,  Mengel  pointed  out 
that  since  the  offspring  of  coy-dogs  would  be 
born  under  harsh  winter  conditions,  and  since 
male  coy-dogs  do  not  demonstrate  the  same 
tendency  to  parental  care  as  male  coyotes, 
the  hybrid  pups  would  be  unlikely  to  survive. 
These  factors  taken  together  seemingly  would 
prevent  the  establishment  of  a  population  of 
canids  of  mixed  coyote  and  dog  ancestry. 
Gier  (1968)  and  Kennelly  and  Roberts  (1969) 
also  reported  the  shift  in  breeding  time 
among  coy-dogs.  Iljin  (1941)  found  a  parallel 
situation  in  captive  hybrids  of  C.  lupus  and 
C.  familiaris. 

Gipson  (1972),  Gipson,  Sealander,  and 
Dunn  (1974),  and  Gipson,  Gipson,  and  Sea- 
lander  ( 1975 )  concluded  that  in  Arkansas 
introgression  of  dog  genes  into  the  coyote 
population  could  and  did  occur.  This  view 
was  based  in  part  on  the  fact  that  38  of  284 
skulls  of  wild  canids  recently  collected  in 
Arkansas,  were  shown  by  multivariate  analy- 
sis to  be  morphologically  intermediate  to  dog 
and  coyote  populations,  and  thus  were  desig- 
nated coy-dogs.  There  was  no  evidence,  how- 
ever, that  any  of  these  individuals  were  other 
than  first  generation  offspring.  It  also  was 
found  that  some  male  coyotes  in  Arkansas 
were  physiologically  in  breeding  condition  in 
late  November  and  December.    On  the  basis 


18 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


of  examination  of  female  reproductive  tracts, 
however,  the  earliest  reported  actual  mating 
was  17  February. 

Freeman  (1976:40-42)  pointed  out  that 
in  Oklahoma,  at  least,  winter  birth  would  not 
seriously  affect  the  offspring  of  coy-dogs, 
since  weather  conditions  were  not  so  severe 
in  that  area.  He  also  suggested  that  lack  of 
care  by  the  male  parent  would  not  necessarily 
result  in  loss  of  the  litter,  since  female  coy- 
otes often  successfully  rear  young  after  losing 
their  mate. 

Silver  and  Silver  ( 1969 )  raised  and  stud- 
ied a  litter  of  five  coyotelike  canids  dug  from 
a  den  in  Croydon,  Sullivan  County,  New 
Hampshire,  and  50  of  their  descendents.  Two 
hybrid  litters  were  born  to  one  of  the  orig- 
inal females  crossed  with  domestic  dogs.  The 
hybrids  were  fertile  and  produced  two  litters, 
but  once  again  it  was  found  that  breeding 
occurred  three  to  four  months  earlier  than  in 
wild  Cams,  and  that  males  did  not  assist  in 
rearing  the  young.  Considering  these  factors, 
and  also  the  disadvantages  faced  by  pups 
born  in  midwinter,  Silver  and  Silver  thought 
it  unlikely  that  eoy-dogs  could  establish  or 
merge  with  a  wild  population.  On  the  basis 
of  behavioral  and  physical  studies  of  these 
various  animals,  it  was  concluded  that  the 
present  population  of  New  England  Canis  is 
of  predominantly  coyote  ancestry,  but  that 
some  dog  and/ or  wolf  genes  had  been  intro- 
duced in  the  past.  Such  genes,  however,  had 
become  well  integrated  so  that  the  population 
was  now  stabilized  and  breeding  true  to 
type.  Therefore,  Silver  and  Silver  thought 
that  wild  Canis  in  New  England  should  not 
be  considered  as  hybrids,  but  as  a  kind  of 
coyote. 

Lawrence  and  Bossert  ( 1969 )  subjected 
31  skulls  of  New  England  Canis  to  linear  dis- 
criminant analysis.  Of  these  specimens,  16 
were  offspring  of  the  captive  canids  studied 
by  Silver  and  Silver,  and  15  were  killed  wild 
in  New  Hampshire,  Vermont,  and  Massachu- 
setts. In  the  analysis,  most  of  the  specimens 
fell  between  known  samples  of  C.  lupus  and 


C.  latrans,  a  few  overlapped  with  C.  latrans, 
and  a  few  also  approached  the  range  of  C. 
familiaris.  The  positions  of  these  wild  indi- 
viduals were,  however,  substantially  different 
from  those  of  21  known,  captive  raised  Fi 
and  FL»  coyote-dog  hybrids.  Lawrence  and 
Bossert,  like  Silver  and  Silver,  concluded  that 
the  New  England  population  was  predomi- 
nantly coyote,  probably  with  some  dog/ wolf 
ancestry.  A  series  of  32  skulls  of  C.  latrans 
thamnos  from  Minnesota  also  demonstrated 
a  shift  away  from  typical  C.  latrans  toward 
both  C.  familiaris  and  C.  lupus,  but  not  to  the 
extent  found  in  New  England  Canis.  Hence 
the  New  England  population  was  considered 
to  represent  the  development  of  a  trend  that 
had  begun  at  an  earlier  time  through  the 
introduction  of  wolf  or  dog  genes  into  the 
coyote  population.  The  designations  "Canis 
latrans  var."  and  "eastern  coyote"  were  used 
for  the  wild  population  of  New  England 
Canis. 

Chambers,  et  al.  (1974)  used  the  same 
technique  as  Lawrence  and  Bossert  (1969) 
on  nearly  150  recently  collected  specimens 
from  New  York,  and  came  to  the  same  con- 
clusion regarding  the  systematic  position  of 
the  population.  Still  another  statistical  analy- 
sis of  skulls  was  done  on  Maine  material  by 
Hilton  (1977)  who  found  that,  except  for 
five  dogs,  all  specimens  received  from  1968 
to  1975  could  be  identified  as  eastern  coyotes. 

As  previously  mentioned,  Mengel  ( 1971 ) 
thought  that  the  introduction  of  dog  genes 
into  a  population  of  wild  Canis  would  be  un- 
likely. He  therefore  suggested  that  the  pres- 
ent population  of  New  England  Canis  was 
the  result  of  the  introgression  of  genes  only 
from  the  wolf  ( C.  lupus  hjcaon )  into  the  wild 
coyote  population.  This  view  was  supported 
in  part  by  a  discriminant  function  analysis  in 
which  skulls  of  55  coyotes  taken  in  north- 
eastern Kansas,  12  known  captive  raised  coy- 
ote-dog hybrids,  and  13  presumptive  wild 
coy-dogs  were  compared  to  the  same  samples 
of  Canis  used  by  Lawrence  and  Bossert 
( 1969 ) .    Since   domestic   dogs   are   abundant 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


19 


in  northeastern  Kansas,  and  since  coy-dogs 
arc  regularly  reported  in  the  area,  Mengel 
argued  that  the  local  coyote  population 
should  demonstrate  the  same  variability  as 
that  of  New  England,  if  introgression  from 
C.  familiaris  were  possible.  Because  in  the 
analysis  the  55  coyotes  fell  close  to  the  orig- 
inal coyote  sample,  and  because  the  positions 
of  the  coyote-dog  hybrids  substantially  dif- 
fered from  those  of  the  Kansas  coyotes,  Men- 
gel  concluded  that  introgression  from  C. 
familiaris  to  C.  latrans  was  not  occurring. 
Canis  lupus,  long  extinct  in  Kansas,  but  still 
present  in  parts  of  the  northeast,  was  thought 
to  be  the  only  source  of  the  genes  that  had 
modified  the  northeastern  coyote  population. 
Known  cases  of  hybridization  between  C. 
lupus  and  C.  latrans  are  rarer  than  those 
between  other  pairs  of  species  of  Canis 
(Gray,  1972).  Indeed,  until  recently  the  only 
suggestion  that  such  a  cross  was  possible  was 
the  statement  by  Young  (1951:124)  that  two 
specimens  of  supposed  coyote-wolf  hybrids, 
born  in  captivity,  were  in  the  Royal  Ontario 
Museum.  Kolenosky  (1971)  reported  that  in 
May  1969  and  May  1970,  litters  of  five  healthy 
pups  were  born  to  a  female  C.  lupus  lycaon, 
captured  in  Algonquin  Provincial  Park,  and  a 
male  C.  latrans,  taken  in  York  County,  On- 
tario. Kolenosky  and  Standfield  ( 1975 )  added 
that  a  subsequent  mating  of  Fi  siblings  pro- 
duced a  litter  of  four  pups.  They  also  ob- 
served that  the  members  of  the  Fj  generation 
were  similar  in  appearance  to  many  speci- 
mens of  a  small  kind  of  wolf,  designated  the 
"Tweed  type,"  that  had  been  collected  along 
the  southern  limits  of  the  range  of  their  "Al- 
gonquin type"  of  C.  lupus  lycaon  in  south- 
eastern Ontario.  They  stated  that  evidence 
was  mounting  that  the  "Tweed  type"  had 
originated  from  hybridization  between  C.  la- 
trans and  C.  lupus  lycaon  in  the  wild. 

Examination  of  Specimens 

With  the  above  background  in  mind,  I 
examined  all  skulls  of  wild  Canis  from  the 
northeastern   United  States  and  southeastern 


LUPUS 


LATRANS 


LUPUS 


FAMILIARIS 


-5   -4    -3    -2     -I      0 


2      3 


Fie.  7. — Multivariate  comparison  of  individual 
specimens  of  C.  lupus  lycaon  to  the  ranges  of  varia- 
tion of  the  total  series  shown  in  Fig.  5.  Black  dots, 
statistical  positions  of  lycaon  from  Michigan,  Minne- 
sota, Wisconsin,  and  western  Ontario;  triangles,  posi- 
tions of  lycaon  from  southeastern  Ontario  and  south- 
ern Quebec;  squares,  positions  of  two  specimens 
thought  to  be  hybrids  between  C.  lupus  lycaon  and 
C.  familiaris.    Males  are  above,  females  below. 


20 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


Canada  that  were  available  to  me.  The  pre- 
viously established  total  series  of  northern 
and  western  wolves  and  coyotes,  and  the 
series  of  50  domestic  dogs,  were  used  as 
standard  groups  against  which  the  northeast- 
ern material  was  tested  (see  pp.  11-12). 

Each  skull  previously  identified  by  others 
as  C.  lupus  tycoon,  except  the  six  to  be  dis- 
cussed later,  were  compared  individually  to 
the  three  standard  groups  by  multivariate 
analysis.  Most  of  these  skulls  fall  within  the 
range  of  variation  of  C.  lupus,  but,  as  could 
be  expected  considering  their  relatively  small 
over-all  size  and  narrow  proportions,  they 
demonstrate  a  shift  toward  C.  latrans  (Fig. 
7).  Two  males  from  the  upper  peninsula  of 
Michigan  (see  appendix  A,  part  6)  have  a 
statistical  position  intermediate  to  those  of 
C.  lupus  and  C.  familiaris,  and  also  appear 
by  eye  to  be  of  mixed  blood.  These  two 
skulls,  therefore,  are  considered  to  represent 
hybrids,  and  henceforth  are  not  used  in  the 
analyses. 

Of  the  remaining  specimens  represented 
in  figure  7,  a  group  of  72  were  collected  in 
the  upper  peninsula  of  Michigan,  northern 
Wisconsin,  northern  Minnesota,  and  that  part 
of  Ontario  to  the  north  and  west  of  Stand- 
field's  (1970)  line  separating  the  "Algonquin 
type"  and  "Ontario  type"  of  C.  lupus  hjcaon; 
and  a  group  of  31  were  obtained  in  south- 
eastern Ontario  and  southern  Quebec  (see 
appendix  A,  part  7.  and  Fig.  2).  The  multi- 
variate distribution  of  these  two  groups  shows 
overlap  and  does  not  suggest  any  sharp  dis- 
tinction. Wolves  from  the  more  westerly  re- 
gion are  larger,  and  have  relatively  broader 
rostra  and  frontal  shields,  and  smaller  second 
upper  molars  (see  appendix  R,  part  3).  In 
these  and  other  characters  the  western  hjcaon 
do  appear  to  fall  between  the  eastern  hjcaon 
and  C.  lupus  nubilus  from  the  Great  Plains. 
Thus  Mech  and  Frenzel  ( 1971 )  could  have  a 
case  in  suggesting  the  survival  of  nubilus  in 
Minnesota.  Throughout  the  entire  region  in 
question,    however,   we   seem   to   be   dealing 


4  r 


LATRANS 


LUPUS 


4HATRANS 

3 

2 

I 

0 
-I 
-2 
-3 
-4 
-5 
-6 
-7 


-5    -4     -3    -2-10       I 


3     4 


Fig.  8. — Multivariate  comparison  of  all  specimens 
of  C.  lupus  (including  hjcaon),  specimens  of  C.  la- 
trans  from  northern  and  western  North  America,  and 
C.  familiaris.  Only  the  margins  of  the  range  of  varia- 
tion of  each  species  are  shown.  Males  are  above, 
females  below. 

only  with  minor  and  gradual  variation,  when 
considering  the  species  C.  lupus  as  a  whole. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


21 


These  findings  are  not  intended  to  contra- 
diet  the  conclusion  of  Kolenosky  and  Stand- 
field  (1975)  that  there  is  a  significant  statisti- 
cal difference  between  specimens  of  C.  lupus 
hjcaon  from  various  parts  of  Ontario.  Those 
authors  dealt  primarily  with  a  question  of 
intraspecific  variation.  My  studies  tend  to 
show  only  that  material  referred  to  lycaon 
is  not  more  noticeably  distinctive  from  C. 
lupus  in  general  than  is  material  referred  to 
most  other  named  subspecies  of  the  gray 
wolf. 

Satisfied  that  the  103  skulls  of  lycaon 
discussed  above  were  representative  of  the 
species  C.  lupus,  I  incorporated  them  into 
my  gray  wolf  samples.  Graphical  results  of 
multivariate  analyses  comparing  these  total 
samples  with  the  standard  samples  of  do- 
mestic dogs  and  coyotes  are  shown  in  figure  8. 

All  available  skulls,  previously  identified 
in  collections  as  coyotes  (with  one  exception 
to  be  discussed  later),  and  that  had  been 
taken  in  the  range  delineated  by  Jackson 
(1951:266-267)  for  C.  latrans  thamnos  (plus 
extreme  southern  Quebec),  were  individually 
tested  against  the  three  standard  groups  of 
C.  lupus  (including  lycaon),  C.  latrans,  and 
C.  familiaris.  The  specimens  included  80(22) 
males  and  50(13)  females  from  Manitoba, 
North  Dakota,  Minnesota,  Iowa,  Illinois,  In- 
diana, Michigan,  Wisconsin,  Ontario,  and 
Quebec  (see  appendix  A,  part  8).  Relative 
multivariate  positions  are  depicted  in  figure 
9.  Nearly  all  specimens  fall  within  or  near 
the  total  range  of  variation  of  C.  latrans,  but 
are  concentrated  closer  to  C.  lupus  than  is  the 
standard  coyote  sample.  One  skull,  obtained 
in  March  Township,  Ontario,  appears  to  rep- 
resent a  coyote-dog  hybrid.  The  overwhelm- 
ing statistical  affinity  of  thamnos  to  the  stand- 
ard sample  of  C.  latrans  suggests  that  the 
coyote  has  established  itself  in  essentially 
unmodified  form  in  much  of  the  northeast. 

In  addition  to  the  above,  I  was  able  to 
examine  a  number  of  specimens  from  the 
extreme  northeastern  United  States  where 
the    wild    canid    population    was    designated 


LUPUS 


LATRANS 


LUPUS 


-i 1 1 1 r~ 

-6    -5    -4     -3    -2 


-i r- 

-i      0 


-i 1 1 1- 

12      3     4 


Fig.  9. — Multivariate  comparison  of  individual 
specimens  of  C.  latrans  thamnos  to  the  ranges  of 
variation  of  the  total  series  of  C.  lupus,  C.  latrans, 
and  C.  familiaris  shown  in  Fig.  8.  Black  dots,  sta- 
tistical positions  of  thamnos;  square,  position  of  prob- 
able hybrid  between  C.  latrans  thamnos  and  C.  fa- 
miliaris.   Males  are  above,  females  below. 


22 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


"C.  latrans  var."  by  Lawrence  and  Bossert 
(1969).  Wild-caught  animals  were  repre- 
sented by  skulls  of  25(5)  males  and  20(2) 
females  (see  appendix  A,  part  9).  Multivari- 
ate positions,  relative  to  the  three  standard 
groups,  are  depicted  in  figure  10. 

Figure  10  also  shows  the  statistical  posi- 
tions of  nine  known  coyote-dog  hybrids 
(some  of  those  discussed  by  Mengel,  1971; 
all  in  KU),  and  the  positions  of  15  wild- 
caught  individuals  (all  of  those  in  USNM 
and  KU  that  could  be  utilized)  previously 
identified  by  others  as  suspected  coy-dogs. 

The  specimens  of  wild  northeastern  Canis 
are  statistically  proximal  to  the  standard  sam- 
ple of  C.  latrans,  but  demonstrate  a  pro- 
nounced shift  toward  C.  lupus.  By  eye,  most 
of  these  skulls  appear  coyotelike,  though  on 
the  average  they  are  larger  than  C.  latrans 
and  have  relatively  broader  rostra  and  frontal 
shields.  Both  visually  and  statistically,  five 
individuals  stand  out  from  the  others  and 
have  multivariate  positions  approaching  the 
limits  of  the  sample  of  C.  familiaris.  My  data 
thus  support  the  view  of  Lawrence  and  Bos- 
sert (1969)  that  both  domestic  dog  and  gray 
wolf  genes  have  influenced  Canis  in  the 
northeast.  Nonetheless,  the  amount  of  graph- 
ical separation  between  the  main  clusters  of 
northeastern  canids  and  the  few  that  are 
scattered  in  the  direction  of  C.  familiaris, 
suggests  to  me  that  two  separate  phenomena 
are  involved.  First,  a  limited  amount  of  hy- 
bridization between  wild  Canis  and  domestic 
dogs  has  occurred,  but  there  has  been  no 
large-scale  shift  in  characters,  among  the 
wild  population  as  a  whole,  in  the  direction 
of  C.  familiaris.  Secondly,  the  predominantly 
coyotelike  population  of  northeastern  Canis 
evidently  has  sustained  the  introgression  of 
genes  from  the  gray  wolf.  There  are  not  yet 
enough  data  to  determine  exactly  when  and 
where  this  process  began,  but  apparently 
wolf  genes  are  now  incorporated  in  the  north- 
eastern coyote. 

The  relationships  among  the  populations 
of  Canis  in  the  northeast  and  in  other  regions 


LUPUS 


4 
3 
2 
I 

0 
-I 
-2 
-3 
-4 
-5 
-6 
-7 


-5    -4     -3    -2 

LATRANS 


-6    -5    -4 


-1 r- 

-3    -2 


-I 


I 


2      3 


Fie.  10. — Multivariate  comparison  of  various  in- 
dividual specimens  to  the  ranges  of  variation  of  the 
total  series  of  C.  lupus,  C.  latrans,  and  C.  familiaris 
shown  in  Fig.  8.  Black  dots,  statistical  positions  of 
specimens  of  wild-caught  individuals  from  New  Eng- 
land, New  York,  and  Pennsylvania;  open  circles, 
wild-caught  animals  previously  reported  as  coy-dogs; 
H,  captive  born  coy-dogs;  triangles,  specimens  from 
southeastern  Canada  thought  to  be  hybrids  between 
C.  lupus  and  C.  latrans;  M,  specimens  in  ROM 
thought  to  be  from  captive  wolf-coyote  hybrids. 
Males  are  above,  females  below. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


23 


TARLE  1 

Statistical  distance,  D2,  between  populations  of  Canis 
in  the  northeast  and  elsewhere. 


C.  familiaris 

A.  Males 

C.  lupus 68.2 

C.  lupus  lycaon  61.7 

C.   latrans  _ 55.8 

C.  latrans  thamnos  ... 53.3 

Northeast  U.S.  Canis  48.4 

B.  Females 

C.  lupus    51.6 

C.  lupus  hjcaon  46.0 

C.    latrans               56.6 

C.  latrans  thamnos  55.8 

Northeast  U.S.  Canis  47.4 


C.  lupus 


C.  lupus 
hjcaon 


C.  latr 


C.  latrans 
thamnos 


4.1 

68.1 

48.3 

63.2 

42.2 

2.0 

49.2 

30.2 

5.4 

3.2 


4.6 

56.8 

39.3 

53.6 

34.9 

2.2 

41.3 

25.1 

3.5 

2.1 


hitherto  discussed,  are  summarized  in  table  1. 
Data  from  those  specimens  considered  to 
represent  hybrids  between  wild  Canis  and 
C.  familiaris  were  not  used  in  the  calculation 
of  the  statistics  shown.  Northern  and  western 
wolves  demonstrate  close  relationship  with 
the  subspecies  C.  lupus  hjcaon  of  the  north- 
east; and  there  is  affinity  between  northern 
and  western  coyotes,  C.  latrans  thamnos,  and 
the  populations  of  Canis  in  the  extreme  north- 
eastern United  States.  This  latter  group  and 
C.  lupus  hjcaon  are  separated  by  less  than 
half  the  statistical  distance  found  between 
the  main  groups  of  northern  and  western 
wolves  and  coyotes. 

I  above  postponed  discussion  of  six  speci- 
mens previously  identified  in  collections  as 
C.  lupus  lycaon,  and  one  identified  as  C. 
latrans  tliamnos  (see  appendix  A,  part  6). 
Each  of  these  skulls  appeared  to  be  of  un- 
usual size  or  to  have  other  characters  sug- 
gesting hybridization  between  C.  lupus  and 
C.  latrans.  Multivariate  positions  are  shown 
in  figure  10. 

In  addition  to  the  above,  I  examined  two 
skulls  (ROM  31-9-15-1  and  31-9-15-2)  re- 
ported to  represent  the  hybrid  offspring  of  a 
captive  wolf  and  coyote   (Young,   1951:124; 


Mengel,  1971:333).  The  labels  of  these  speci- 
mens read  in  part:  "From  Ontario  stock." 
Multivariate  positions  of  each,  relative  to  the 
three  standard  series,  are  plotted  in  figure  10 
(since  ROM  31-9-15-2  lacked  both  PI,  it 
would  not  ordinarily  have  been  included  in 
a  multivariate  analysis,  but  because  of  its 
special  interest  I  used  an  estimate  of  alveolar 
length  of  upper  toothrow). 

If  considered  together,  the  positions  of 
these  nine  specimens  bridge  the  gap  be- 
tween the  statistical  limits  of  the  standard 
sample  of  C.  lupus  and  the  positions  of  the 
individual  specimens  from  the  northeastern 
United  States.  It  is  thus  reasonable  to  sup- 
pose that  hybridization  between  wolf  and 
coyote  in  southeastern  Canada  has,  and  prob- 
ably still  is  permitting  the  flow  of  genes  from 
one  species  to  the  other.  Evidently  wolf 
genes  have  spread  through  much  of  the  coy- 
ote population,  and  resulting  wolflike  char- 
acters have  been  phenotypically  expressed  in 
Canis  of  the  extreme  northeastern  United 
States.  This  introgression  may  have  assisted 
the  coyote  population  of  the  region  to  adapt 
to  and  flourish  in  an  environment  far  from 
the  original  prairie  habitat  of  C.  latrans. 


24 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


Systematic  Problems  in 
the  Southeast 

Background 

When  Europeans  first  entered  the  south- 
eastern part  of  North  America,  they  found 
animals  that  appeared  to  be  closely  related  to 
the  wolves  of  the  Old  World.  For  example, 
Catesby  (1771:xxvi),  writing  of  Florida  and 
the  Carolinas,  noted:  "The  Wolves  in  Amer- 
ica are  like  those  of  Europe,  in  shape  and 
colour,  but  are  somewhat  smaller." 

Bartram  (1791:199),  the  first  author  to 
apply  a  binomial  to  Canis  in  the  southeastern 
region,  referred  to  the  black  wolves  that  he 
saw  in  Florida  as  "lupus  niger."  Harlan 
(1825:82)  used  the  name  Canis  lycaon  for 
these  same  animals,  but  also  for  black  wolves 
inhabiting  mountainous  areas  of  North  Amer- 
ica and  Europe.  The  designation  Canis  ly- 
caon originally  had  been  given  to  a  black 
wolf  from  the  vicinity  of  Quebec  by  von 
Schreber  in  1775  (Goldman,  1944:437-440), 
and  was  restricted  by  Miller  (1912b)  to  the 
wolves  of  eastern  Canada  and  the  north- 
eastern United  States.  Richardson  (1829:70) 
termed  the  black  wolves  that  he  saw  on  the 
banks  of  the  Mackenzie  and  Saskatchewan 
rivers  of  western  Canada  as  "Canis  lupus 
occidentalis  var.  E.  Lupus  ater."  He  said, 
however,  that  the  same  animals  occurred 
throughout  North  America,  and  he  included 
Bartram's  Florida  wolves  under  this  name. 
It  thus  seems  that  early  naturalists  named 
kinds  of  wolves  largely  on  the  basis  of  color, 
and  considered  the  names  to  apply  where- 
ever  the  particular  colors  were  found.  Since 
it  was  eventually  demonstrated  that  colora- 
tion of  wolves  in  nearly  all  parts  of  North 
America  is  highly  variable  and  of  minimal 
taxonomic  value  (Young,  1944:59-66;  Gold- 
man, 1944:401),  these  early  writings  are  use- 
less to  an  understanding  of  the  problems 
presently  under  review. 

Apparently  Audubon  and  Bachman  (1851: 
126,  240)  were  the  first  authors  to  set  definite 
bounds  to  the  ranges  of  named  kinds  of  North 


American  wolves,  and  to  suggest  that  in  the 
southern  United  States  there  existed  wolves 
structurally  different  from  those  in  other  re- 
gions. They  kept  Richardson's  designation  in 
their  description  of  Canis  lupus  var.  Arer,  the 
"Black  American  Wolf,"  but  recorded  this 
kind  only  from  Florida,  South  Carolina, 
North  Carolina,  Kentucky,  southern  Indiana, 
southern  Missouri,  Louisiana,  and  northern 
Texas.  They  also  discussed  Canis  lupus,  var. 
Rufus,  the  "Red  Texan  Wolf,"  which  they 
said  ranged  from  northern  Arkansas,  through 
Texas,  and  into  Mexico.  In  their  description 
of  this  animal  Audubon  and  Bachman  men- 
tioned the  long  legs,  pointed  nose,  and  slen- 
der proportions.  They  noted  that  although 
the  reddish  shade  predominated  in  Texas, 
other  colors  also  were  represented  there,  and 
that  the  wolves  of  different  colors  freely  in- 
terbred. They  stated  that  except  for  Canis 
latrans,  the  coyote,  all  of  the  wolves  that  they 
described  were  only  varieties  of  one  species. 

Audubon  and  Bachman's  delineation  of 
Canis  in  the  southern  United  States  was  gen- 
erally accepted  by  biologists.  But  toward  the 
end  of  the  nineteenth  century  finer  taxonomic 
splitting  became  more  fashionable,  and  Bangs 
(1898)  designated  the  Florida  wolf  as  a  full 
species,  Canis  ater.  Bailey  (1905)  question- 
abb  referred  to  the  large,  dark  wolf  of  east 
Texas  as  C.  ater,  and  expressed  hope  that 
specimens  could  be  obtained  to  confirm  its 
status.  He  also  recognized  Audubon  and 
Bachman's  red  wolf  as  a  full  species  with 
the  name  Canis  rufus,  and  assigned  it  a  range 
in  southern  and  central  Texas.  Bailey  dis- 
tinguished the  larger  gray  wolf  (C.  griseus= 
C.  lupus)  of  western  Texas  from  C.  rufus, 
and  referred  to  the  latter  as  "a  large  coyote 
or  small  wolf." 

Miller  (1912a)  explained  that  the  name 
ater  was  technically  unavailable,  and  he  des- 
ignated the  Florida  wolf  as  C.  floridanus. 
This  name  then  became  generally  accepted 
for  wolves  in  the  forested  areas  of  the  south- 
eastern United  States,  while  C.  rufus  con- 
tinued to  be  recognized  in  central  and  south- 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


25 


em  Texas.  At  this  time,  however,  there  had 
not  yet  been  an  attempt  to  associate  C.  rufus 
and  C.  floriclanus  into  one  group  that  demon- 
strated characters  different  from  those  of  all 
other  North  American  wolves. 

Goldman  (1937)  combined  the  wolves  of 
the  south-central  and  southeastern  United 
States  into  a  single  species,  C.  rufas,  that  he 
considered  distinct  from  all  other  North 
American  wolves.  He  combined  the  latter 
into  the  species  C.  lupus.  Goldman  said  that 
C.  rufus  "exhibits  a  departure  from  the  true 
wolves,  and  in  cranial  and  dental  characters 
approaches  the  coyotes."  He  listed  the  names 
C.  rufus  rufus  for  the  Texas  subspecies,  and 
C.  r.  floridanus  for  the  eastern  race,  and  he 
also  described  C.  r.  gregoryi,  a  new  sub- 
species in  the  lower  Mississippi  Valley. 
Shortly  thereafter,  Harper  (1942)  pointed 
out  that  Bartram's  ( 1791 )  term  for  the  Flor- 
ida wolf  was  actually  the  earliest  name  to 
have  been  applied  in  the  southeast,  and  the 
specific  designation  of  the  wolves  in  the 
region  then  technically  became  C.  niger.  But 
the  International  Commission  on  Zoological 
Nomenclature  (1957,  opinion  447)  rejected 
Bartram's  technical  terms,  and  Hall  (1965) 
listed  C.  rufus  as  the  proper  name.  There- 
fore, the  trinomials  applied  by  Goldman 
( 19.37 )  are  presently  considered  valid  and 
are  used  in  subsequent  discussion. 

Goldman  (1944:481)  observed  that  the 
two  eastern  subspecies  of  C.  rufus  exhibited 
a  remarkable  approach  in  size  and  general 
proportions  to  the  eastern  gray  wolf,  C.  lupus 
hjcaon,  but  that  there  existed  several  specific 
cranial  differences.  On  the  other  hand,  he 
said  that  the  subspecies  C.  r.  rufus  in  central 
Texas  and  Oklahoma  was  so  small  and  in 
general  characters  agreed  so  closely  with  C. 
latrans,  that  some  specimens  were  difficult  to 
distinguish.  He  suggested  the  possibility  of 
hybridization  between  C.  rufus  and  C.  latrans 
in  some  localities  in  Texas. 

Goldman  (1944)  used  the  vernacular  "red 
wolf"  for  the  species  C.  rufus,  presumably  on 
tlie  basis  of  Audubon  and  Bachman's  descrip- 


tion of  the  "Red  Texan  Wolf."  This  term, 
however,  is  not  found  in  any  of  the  early 
literature  discussing  wolves  in  states  east  of 
Texas.  Kellogg  (1915:41)  said  that  trappers 
in  Cherokee  County,  southeastern  Kansas,  did 
refer  to  "Red  Wolves"  in  the  area.  But  other- 
wise this  popular  term  seems  to  have  been 
restricted  to  parts  of  Texas  until  Goldman 
introduced  its  use  throughout  the  range  of 
C.  rufus.  This  appellation  may  be  unfortu- 
nate, because  although  the  rufous  element  in 
the  fur  sometimes  stand?  out,  the  "red  barn- 
roof  paint"  color  mentioned  by  Young  (1946: 
36)  seldom  shows  up.  Early  records  indicate 
that  a  dark-colored  or  entirely  black  phase 
was  locally  common  in  the  eastern  forests. 
But  most  available  specimens  of  "red  wolves" 
actually  exhibit  a  typical  wild  canid  color 
pattern,  consisting  of  an  agouti  gray  or 
brown,  interspersed  with  black  hairs,  espe- 
cially on  the  back,  and  with  the  muzzle,  ears, 
and  outer  surfaces  of  the  limbs  tending  to- 
ward a  tawny  color. 

Lawrence  and  Bossert  (1967)  thought 
that  separation  of  C.  rufus  as  a  distinct  spe- 
cies rested  too  heavily  on  the  small  red  wolves 
of  central  Texas  where  hybridization  with  the 
coyote  may  have  been  a  factor.  They  said 
that  if  the  study  of  wolves  in  the  south  had 
been  based  on  adequate  series  of  specimens 
from  Florida,  separation  of  C.  rufus  from  C. 
lupus  would  have  been  highly  unlikely.  In  a 
multivariate  analysis  they  compared  all  avail- 
able skulls  of  wolves  collected  before  1920  in 
Louisiana,  Alabama,  and  Florida,  including 
one  Florida  specimen  assigned  by  Goldman 
to  C.  lupus  hjcaon  (according  to  Lawrence 
and  Bossert,  1975:81,  a  total  of  12  specimens 
were  in  this  sample),  with  series  of  20  adult 
skulls  each  of  C.  familiaris,  C.  lupus  (various 
North  American  subspecies),  C.  lupus  hjcaon, 
and  C.  latrans.  They  found  C.  lupus  hjcaon 
and  C.  rufus  (the  early  southeastern  ma- 
terial) both  to  overlap  with  C.  lupus,  but  to 
be  distinct  from  each  other.  All  three  of 
these  populations  formed  a  cluster  distinct 
from    C.    latrans,    with    C.    rufus    being    the 


26 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


farthest  removed.  Lawrence  and  Bossert  in- 
terpreted this  analysis  as  demonstrating  that 
early  populations  described  as  C.  rufus,  "east 
of  the  range  of  Canis  latrans,  are  a  local  form 
of  Canis  lupus,  not  a  distinct  species  of  wolf." 

Lawrence  and  Bossert  ( 1975 )  repeated 
the  above  interpretation.  They  again  ob- 
served that  their  12  early  specimens  from 
Louisiana  and  eastward,  which  they  now  des- 
ignated "floridanus,"  and  their  series  of  C. 
lupus  lycaon,  were  both  less  coyotelike  than 
their  general  sample  of  C.  lupus.  Lawrence 
and  Bossert  ( 1975 )  also  evaluated  a  series 
of  30  specimens  (designated  as  "gregoryi") 
taken  in  the  1920's  in  Arkansas,  south  of  the 
Arkansas  River.  This  group  was  found  to  be 
nearer  in  D2  distance  to  the  general  sample 
of  C.  lupus  and  to  C.  lupus  lycaon,  than  to 
"floridanus." 

The  suggestions  by  Lawrence  and  Bossert 
(1967,  1975)  — (1)  that  pre-1920  eastern  C. 
rufus  is  not  more  than  subspecifically  distinct 
from  C.  lupus,  (2)  that  C.  rufus  and  C.  lupus 
lycaon  are  less  coyotelike  than  C.  lupus  in 
general,  and  ( 3 )  that  Arkansas  red  wolves  are 
closer  to  C.  luptis  than  to  Louisiana  red 
wolves — are  not  in  agreement  with  my  own 
findings.  These  suggestions,  at  least  in  part, 
also  seem  not  to  correspond  well  with  results 
obtained  by  some  other  workers. 

Paradiso  ( 1968 )  argued  that  the  relatively 
small  samples  used  by  Lawrence  and  Bossert 
( 1967 )  did  not  adequately  represent  the  vari- 
ability shown  by  Canis,  and  thus  that  con- 
specificity  of  the  red  and  gray  wolves  had 
not  been  demonstrated.  He  also  pointed  out 
several  cranial  and  dental  characters  in  which 
C.  rufus  resembled  C.  latrans  more  than  it  did 
C.  lupus.  Paradiso  and  Nowak  (1972a)  com- 
pared data  on  213  skulls  of  C.  rufus,  214  of 
C.  lupus,  and  336  of  C.  latrans,  and  concluded 
that  the  red  wolf  was  a  distinct  species.  In 
size  and  proportion  C.  rufus  fell  between  C. 
latrans  and  C.  lupus,  but  was  nearer  to  the 
latter.  Development  of  certain  dental  char- 
acters in  C.  rufus  suggested  affinity  to  the 
coyote.     Atkins    and    Dillon    (1971),    on   the 


basis  of  a  morphological  study  of  the  cerebel- 
lum, considered  the  red  and  gray  wolves  to 
be  in  the  same  group,  distinct  from  other 
Canis,  but  that  C.  rufus  was  a  valid  species. 
Shaw  (1975)  evaluated  data  on  the  behavior, 
ecology,  vocalizations,  allelic  frequency,  and 
morphology  of  a  living  population  of  C.  rufus 
gregoryi  in  southeastern  Texas,  and  con- 
cluded that  the  population  represented  a  spe- 
cies distinct  from  both  C.  lupus  and  C.  la- 
trans. 

Three  recent  studies,  aimed  primarily  at 
identifying  newly  collected  specimens  of  wild 
Canis  in  the  south-central  states,  and  each 
employing  a  different  method  of  multivariate 
analysis,  also  have  tended  to  uphold  the  spe- 
cific status  of  C.  rufus.  Gipson,  Sealander, 
and  Dunn  (1974),  who  used  a  single  meas- 
urement and  five  separate  ratios  as  variables, 
compared  as  groups  the  skulls  of  40  C.  la- 
trans, 34  C.  latrans  x  C.  familiaris,  31  C.  fa- 
miliaris, 37  C.  lupus  lycaon,  40  C.  lupus  from 
the  Great  Plains  region,  and  40  C.  rufus  taken 
in  Arkansas  prior  to  1925.  They  found  the 
red  wolf  sample  to  be  statistically  intermedi- 
ate to  the  coyote  and  gray  wolf  samples,  and 
reported  their  results  to  support  recognition 
of  the  red  wolf  as  a  distinct  species.  Freeman 
( 1976 ) ,  who  used  15  raw  measurements  as 
variables,  compared  as  groups  the  skulls  of 
40  C.  latrans,  12  C.  latrans  x  C.  familiaris,  24 
C.  familiaris,  43  C.  lupus  from  the  Great 
Plains  region,  and  44  C.  rufus  taken  in  Arkan- 
sas prior  to  1925.  The  graphical  results  of  his 
analysis  indicate  that  the  red  wolf,  again,  is 
statistically  between  the  coyote  and  gray  wolf. 
Unlike  Gipson,  Sealander,  and  Dunn  (1974), 
however,  who  found  some  overlap  between 
samples  of  C.  lupus  and  C.  rufus.  Freeman 
obtained  complete  statistical  separation  of  the 
two  species.  Elder  and  Hayden  (1977), 
whose  variables  were  proportions  obtained  by 
dividing  14  measurements  by  greatest  length 
of  skull  ( in  accordance  with  Lawrence  and 
Bossert,  1967),  compared  as  groups  the  skulls 
of  29  C.  latrans,  30  C.  familiaris,  and  IS  C. 
lupus  from   widely  scattered  regions.    Then, 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


27 


27  skulls  of  C.  rufus  taken  in  Missouri  in  the 
1920's  and  early  1930's  were  compared  as 
individuals  to  these  three  groups.  None  of 
the  red  wolves  fell  within  the  limits  of  the 
groups,  but  they  were  distributed  statistically 
between  the  coyote  and  gray  wolf  samples. 

Mech  (1970:25,  285,  351)  suggested  that 
the  differing  views  on  the  status  of  C.  rufus 
could  be  reconciled  by  recognition  of  the  red 
wolf  as  a  hybrid  population  that  could  be 
properly  known  as  C.  lupus  x  C.  latrans.  Re- 
gardless of  the  taxonomic  disputes,  it  is  now 
generally  agreed  that  the  original  populations 
of  Canis  inhabiting  the  southeastern  quarter 
of  the  United  States  have  all  but  disappeared, 
and  have  been  replaced  in  large  part  by 
another  kind  of  Canis  (McCarley,  1962;  Para- 
diso  and  Nowak,  1972a;  Pimlott  and  Joslin, 
1968). 

Examination  of  Earliest  Available 
Eastern  Material 

In  my  own  study,  I  wished  to  consider  all 
available  specimens  from  south  of  Lakes 
Michigan  and  Erie,  and  east  of  the  Mississippi 
River.  Goldman  ( 1944 )  examined  only  four 
skulls  (including  one  damaged  and  one  im- 
mature) and  one  mandibular  ramus,  assigned 
to  C.  lupus  Tycoon,  from  this  region.  He  also 
had  only  two  skulls  of  C.  rufus  gregonji  (both 
subadults),  from  Indiana  and  Illinois,  and 
two  of  C.  r.  floridanus  (one  damaged),  from 
Alabama  and  Florida.  Some  other  workers 
have  considered  this  small  collection  to  be 
insufficient  for  an  assessment  of  the  relation- 
ships among  the  original  populations  in  the 
region,  particularly  for  settling  the  question 
of  whether  C.  rufus  and  C.  lupus  intergraded 
in  the  Ohio  Valley  and  along  the  central 
Atlantic  coast.  Unfortunately,  there  are 
scarcely  any  additional  skulls  available,  and 
there  doubtless  never  will  be,  because  the 
native  wolves  of  the  region  were  exterminated 
long  ago.  Excavation  of  fossil  and  aboriginal 
sites  offers  some  hope,  but  most  material  from 
such  sources   that  I   have  examined  is  frag- 


LUPUS 


LATRANS 


-6    -5    -4 


- r~ 
0 


2      3 


Fie.  11. — Multivariate  comparison  of  individual 
specimens  of  C.  rufus  to  the  ranges  of  variation  of 
the  total  series  of  C.  lupus,  C.  latrans,  and  C.  famil- 
iaris  shown  in  Fig.  8.  L,  pre-1920  specimens  from 
Louisiana;  A,  specimen  taken  in  1917  in  Alabama; 
C,  specimen  found  in  Alabama  cave  deposit;  F, 
Florida  specimen;  K,  specimen  from  Garvin  County, 
Oklahoma;  P,  Pennsylvania  specimen;  T,  southeastern 
Texas  specimen.  Males  are  above,  females  below. 
The  Florida,  Oklahoma,  and  Pennsylvania  specimens 
are  compared  both  with  male  and  female  series. 


28 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


mentary  and  of  limited  value.    This  material 
is   discussed  later  in   the  paper. 

Lawrence  and  Bossert  ( 1967)  limited  their 
sample  of  C.  rafu.s  to  specimens  collected 
before  1920,  believing  that  such  material  was 
the  least  likely  to  show  influence  from  C. 
latrans.  As  an  initial  step,  this  paper  also 
considers  material  taken  prior  to  1920.  But 
from  this  period,  and  from  east  of  the  Missis- 
sippi River,  there  are  available  only  four 
skulls  that  arc  adult  and  unbroken,  and  hence 
suitable  for  inclusion  in  multivariate  analysis. 
Two  of  these  specimens  are  unknown  as  to 
sex,  but  since  the  smallest  of  the  four  is  a 
known  male,  I  compared  each  of  them  indi- 
vidually with  the  previously  compiled  sam- 
ples of  50  C.  familiaris,  166  male  C.  latrans, 
and  294  male  C.  lupus  ( including  61  C.  lupus 
hjcaon ) .  Their  relative  multivariate  positions 
are  plotted  in  figure  11  (the  two  skulls  of 
unknown  sex  are  also  compared  to  appro- 
priate female  samples).  The  positions  of  the 
four  are  beyond  the  range  of  variation  of  any 
of  the  known  samples,  but  are  distributed 
between  C.  lupus  and  C.  latrans.    Measure- 


ments of  these  skulls  are  listed  in  appendix 
B  (part  4),  and  other  details  are  given  in 
table  2. 

Although  these  four  specimens  are  indica- 
tive of  the  former  presence  in  the  eastern 
United  States  of  a  kind  of  wolf  different  from 
any  subspecies  of  C.  lupus,  their  value  is  lim- 
ited by  missing  data.  A  better  collection  of 
pre-1920  material  was  taken  in  Louisiana, 
and  was  identified  by  Goldman  as  C.  rufus 
gregoryi.  The  Louisiana  material  suitable  for 
inclusion  in  my  analysis  consists  of  7(2) 
males  and  2(1)  females  (parentheses  contain 
numbers  of  specimens  in  the  series  for  which 
sex  had  not  been  recorded,  but  which  were 
judged  to  belong  to  the  particular  sex  in- 
dicated; see  p.  5).  A  single  skull  of  a 
female  C.  rufus  gregoryi,  taken  in  extreme 
southeastern  Texas  in  1906,  also  is  available. 

As  depicted  in  figure  11,  the  nine  Louisi- 
ana specimens,  and  the  one  from  southeastern 
Texas,  have  multivariate  positions  interme- 
diate to  the  graphical  limits  of  C.  lupus  and 
C.  latrans.  Seven  of  these  skulls  have  indi- 
vidual   D-   distances   farther   from    C.    lupus 


TABLE  2 
Data  on  early  specimens  of  C.  rufus. 


Collection                                                                                                                                                 D"  from  D=  from 

and  number            Sex                                                 Locality  Date          C.  lupus  C.  latrans 

ANSP  2261             male?        Pennsylvania    _ pre-1859      27.2  23.0 

MCZ  11179            male?        Florida"   1854?           24.5  37.8 

USNM  223936       male          12  mi.  S  Cherokee,  Colbert  County,  Alabama  1917              14.5  30.6 

USNM  348063        male          Fern  Cave,  Jackson  County,  Alabama00  ....              43.8  26.4 

USNM  132229        male          Mer  Rouge,  Morehouse  Parish,  Louisiana  1904              25.6  24.9 

USNM   136834       male          23  mi.  SW  Tallulah,  Madison  Parish,  Louisiana  __  1905              18.1  28.1 

USNM   137125       male          20  mi.  SW  Vidalia,  Concordia  Parish,  Louisiana  ....  1905              37.9  27.2 

MCZ  9114               male          Mer  Rouge,  Morehouse  Parish,  Louisiana  1898              23.2  22.8 

USNM   136731        male          18  mi.  SW  Tallulah,  Madison  Parish,  Louisiana  ....  1905              46.1  52.3 

USNM   133687        male?        15  mi.  NW  Tallulah,  Madison  Parish,  Louisiana  ....  1904              38.4  38.1 

USNM  133688       male?        10  mi.  SW  Floyd,  West  Carroll  Parish,  Louisiana  1904             29.7  37.5 

USNM   136105       female       15  mi.  SW  Tallulah,  Madison  Parish,  Louisiana  ....  1904              22.4  19.4 

USNM  234227       female?     12  mi.  N  Avery  Island,  Iberia  Parish,  Louisiana  ....  1919              31.7  19.1 

USNM   147701       female      Kountze,  Hardin  County,  Texas  1906             38.0  25.2 

"Apparently  with  regard  to  this  specimen,  Barbour  (1944:142)  wrote:  "we  have  in  the  Agassiz  Museum  in 
Cambridge  a  skull  of  a  fine  old  adult  animal  which  was  said  to  have  been  killed  in  1S54  in  the  region  where 
the  city  of  Miami  now  stands." 

"  This  specimen  was  found  in  a  cave;  its  state  of  preservation  suggested  that  it  lived  in  Recent  times  (Para- 
diso  and  Nowak,  1973). 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


29 


-5    -4     -3  -2 


Fig.  12. — Multivariate  positions  (black  dots)  of 
individual  pre-1920  males  of  C.  rufus  from  Louisi- 
ana, relative  to  series  of  males  of  C.  lupus,  C.  latrans, 
and  C.  familiaris.  The  variables  used  in  this  par- 
ticular analysis  were  derived  by  dividing  the  meas- 
urements by  greatest  length  of  skull. 

than  from  C.  latrans  (table  2).  A  separate 
analysis  of  males  only,  in  which  the  employed 
variables  were  fractions  derived  by  dividing 
each  of  the  other  measurements  by  greatest 
length  (in  accordance  with  Lawrence  and 
Bossert,  1967),  produced  much  the  same  pic- 
ture (Fig.  12). 

Measurements  of  the  pre-1920  Louisiana 
material  are  listed  in  appendix  B  (part  4), 
and  other  details  are  given  in  table  2.  In 
figure  13,  specimens  of  males  are  compared 
with  the  series  of  C.  lupus  and  C.  latrans  in 
a  ratio  diagram.  The  Louisiana  skulls  ap- 
proach those  of  C.  lupus  in  over-all  length, 
but  are  comparatively  small  in  most  other 
dimensions,  a  notable  exception  being  their 
large  M2.  In  general  the  proportions  of  C. 
rufus  gregoryi  seem  intermediate  to  those  of 
C.  lupus  and  C.  latrans,  but  the  frontal  shield 
in  the  Louisiana  material  is  even  narrower, 
relatively,  than  in  the  coyote,  and  the  post- 
orbital  constriction  is  narrower,  relatively, 
than  in  the  gray  wolf.  The  results  of  these 
analyses   indicate   that   C.   rufus  morphologi- 


cally resembles  C.  lupus  lycaon  more  than  it 
does  any  other  subspecies  of  gray  wolf. 

We  are  still  left  with  what  Lawrence  and 
Bossert  (1967)  called  "the  biologically  diffi- 
cult problem  of  reconciling  the  existence  of 
two  similarly-sized  forms  of  wolf  in  one  con- 
tinuous habitat."  Actually,  because  of  the 
limited  number  of  specimens,  there  may 
never  be  indisputable  proof  that  the  red  and 
gray  wolves  did  not  undergo  intergradation 
in  the  eastern  forests.  The  existence,  how- 
ever, of  14  complete  skulls  collected  before 
1920  in  this  region,  that  do  not  overlap  in 


-.16 


.12   -.08   -.04   0 


LATRANS 


RUFUS  LUPUS 


70 


80 


.90 


1.00 


Fig.  13. — Ratio  diagram  comparing  means  of 
total  series  of  C.  lupus,  series  of  C.  latrans  from 
northern  and  western  North  America,  and  two  series 
of  C.  rufus  (males  only  are  shown  for  all  series).  The 
solid  line  under  rufus  represents  the  mean  values  of 
pre-1920  Louisiana  specimens;  the  dashed  line  rep- 
resents the  mean  values  of  specimens  taken  from 
1919  to  1929  in  the  south-central  states.  The  verti- 
cally arranged  numbers  correspond  to  the  measure- 
ments so  numbered  in  appendix  B.  A  log  difference 
scale  is  provided  above,  and  a  ratio  scale  below  the 
diagram. 


30 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


statistical  position  with  series  of  482  speci- 
mens of  C.  lupus,  including  103  tycoon,  sug- 
gests to  me  that  the  population  represented 
by  these  skulls  is  more  than  subspecifically 
different  from  the  gray  wolf.  Furthermore, 
the  slender  proportions  of  the  red  wolf  indi- 
cate that  its  prey  averaged  smaller  than  that 
of  the  gray  wolf,  and  that  its  ecological  niche 
may  have  approached  that  of  the  coyote 
which  did  not  exist  in  the  eastern  forests 
when  white  settlers  first  arrived.  Possibly, 
when  C.  lupus  entered  the  northeastern  for- 
ests it  underwent  a  degree  of  parallel  evo- 
lution with  C.  rufus,  just  to  the  south.  The 
subspecies  C.  lupus  lycaon  may  even  have 
been  in  the  process  of  replacing  C.  rufus, 
when  the  white  man  interfered.  And  it  is 
reasonable  to  think,  considering  the  ease  with 
which  interbreeding  occurs  in  Canis,  that 
hybridization  between  C.  lupus  hjcaon  and 
C.  rufus  did  occur,  and  that  a  zone  of  intro- 
gression  may  have  developed  that  tended  to 
modify  one  or  both  populations.  The  advent 
of  the  white  man  would  probably  have  stim- 
ulated the  spread  of  such  hybridization,  just 
as  it  seems  to  have  encouraged  interbreeding 
between  C.  rufus  and  C.  latrans  in  Texas 
(Paradiso  and  Nowak,  1972a).  Be  that  as  it 
may,  available  early  specimens  from  the 
southeastern  United  States  can  all  be  sepa- 
rated from  known  series  of  C.  lupus,  and 
seem  to  represent  a  different  species,  C.  rufus. 

The  exact  distribution  of  C.  lupus  and  C. 
rufus,  and  the  extent  to  which  their  ranges 
overlapped  in  eastern  North  America,  will 
probably  never  be  known.  But  if  one  Florida 
specimen,  called  C.  lupus  lycaon  by  Goldman 
(see  table  2),  is  in  fact  C.  rufus  as  I  have 
tried  to  show,  then  there  is  no  longer  any 
confirmation  of  the  original  presence  of  the 
gray  wolf  in  Florida.  And  if  correct,  identifi- 
cation of  a  Pennsylvania  specimen  as  C.  rufus 
rather  than  lycaon  means  that  the  range  of 
the  red  wolf  once  extended  farther  to  the 
northeast  than  was  formerly  thought. 

As  explained  by  Paradiso  and  Nowak 
(1972a:7-8),  Goldman  was  also  incorrect  in 


his  reasons  for  stating  that  the  range  of  the 
red  wolf  in  the  Mississippi  Valley  once  ex- 
tended as  far  north  as  Warsaw,  Hancock 
County,  Illinois;  and  Wabash  County,  Indi- 
ana. The  specimen  supposedly  from  the  for- 
mer locality  had  been  in  possession  of  an 
animal  dealer,  and  there  is  no  evidence  that 
it  represents  a  native  wolf  of  the  area.  The 
other  specimen  was  actually  collected  in  the 
Wabash  River  area  of  southwestern  Indiana, 
not  in  Wabash  County  father  north.  Both 
skulls  appear  to  be  referable  to  C.  rufus,  but 
they  are  subadults  and  hence  not  suitable  for 
use  in  multivariate  comparisons  made  on  the 
basis  of  adults. 

Examination  of  Material  Collected  from 

1919  to  1929  in  the  South-central 

United  States 

In  the  course  of  Federal  predator  control 
work  in  the  south-central  United  States,  from 
1919  to  1929,  a  large  number  of  specimens  of 
Canis  were  taken.  Most  of  these  were  re- 
ferred by  Goldman  (1944)  to  C.  rufus  gre- 
goryi,  the  "Mississippi  Valley  Red  Wolf." 
Lawrence  and  Bossert  (1967)  did  not  use 
this  material  in  their  attempt  to  define  the 
red  wolf,  because  of  the  possibility  that  some 
of  the  specimens  represented  hybridization 
between  red  wolf  and  coyote.  Paradiso  and 
Nowak  ( 1972a ) ,  however,  noted  that  al- 
though a  few  of  these  specimens  appeared  to 
be  hybrids,  the  remainder  demonstrated  the 
continued  survival  of  the  species  C.  rufus. 

Since  many  of  the  specimens  assigned  to 
C.  r.  grcgoryi  may  indeed  represent  the  red 
wolf  in  unmodified  form,  they  should  be  con- 
sidered in  any  effort  to  ascertain  the  relation- 
ship between  C.  rufus  and  C.  lupus.  But 
since  some  of  the  specimens  may  actually  be 
the  result  of  hybridization  between  C.  rufus 
and  C.  latrans,  their  inclusion  with  unmodi- 
fied C.  rufus  in  statistical  compilation  could 
result  in  a  misunderstanding  of  the  situation. 
There  is  no  completely  objective  method  of 
separating  red  wolves  from  red  wolf-coyote 
hybrids,   but   earlier  work    (McCarley,    1962; 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


31 


Fig.  14. — Map  showing  localities  of  specimens  taken  prior  to  1930  in  the  south-central  United  States. 
Triangles,  C.  rufus;  black  dots,  specimens  originally  identified  as  C.  latrans;  squares,  specimens  originally  iden- 
tified as  C.  rujus,  but  considered  in  this  paper  to  represent  hybridization  between  red  wolf  and  coyote. 
See  figures  18-23  for  more  details.  The  dotted  line  shows  the  western  limits  of  oak  forest  in  central  Texas. 
The  dashed  line  divides  localities  of  specimens  previously  identified  as  C.  rujus  gregoryi  (east)  from  those 
previously  identified  as  C.  rufus  rujus  (west).  Note:  because  of  the  scale  of  the  map  it  was  not  possible 
to  plot  all  localities  in  crowded  areas. 


Paradiso,  1968)  indicated  that  skulls  of  C.  r. 
gregoryi  almost  invariably  have  a  greater 
maximum  length  than  those  of  C.  latrans.  In 
my  preliminary  examination  of  the  series 
identified  as  C.  rufus  gregoryi,  I  noticed  15 
adult  skulls  from  Arkansas  and  eastern  Okla- 
homa that  seemed  comparatively  short  (less 
than  215  millimeters  in  greatest  length  for 
males,  and  less  than  210  millimeters  for  fe- 
males), and  which  were  within  or  near  the 
size  range  of  my  standard  series  of  western 
coyotes.  These  15  specimens  were  considered 
the    most    likely    to    represent    hybridization, 


and  hence  were  not  directly  compared  to  the 
standard  series  of  C.  lupus.  They  are  dealt 
with  later  in  this  account. 

Of  the  remaining  specimens  collected 
from  1919  to  1929  that  Goldman  assigned  to 
gregoryi,  those  that  could  be  subjected  to 
multivariate  analysis  were  63(1)  males  and 
52(1)  females  (see  Fig.  14;  and  appendix  A, 
part  10).  Of  these  115  skulls,  64  were  from 
Arkansas,  19  were  from  southern  Missouri, 
29  were  from  southeastern  Oklahoma,  and 
three  were  taken  in  northern  or  western 
Louisiana  in   1928    (a  specimen  collected  in 


32 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


" LATRANS 


LUPUS 


4 
3 
2 

I 

0 
-I 
-2 
-3 
-4 
-5 
-6 
-7 


LATRANS 


-6    -5-4-3-2-1      0       I       2      3     4 

Fig.  15. — Multivariate  positions  (black  dots)  of 
individual  specimens  of  C.  rufus  taken  from  1919  to 
1929  in  the  south-central  United  States,  relative  to 
the  ranges  of  variation  of  the  total  series  of  C.  lupus, 
C.  latrans,  and  C.  familiaris  shown  in  Fig.  8.  Males 
above,  females  below. 

extreme  southern  Louisiana  in  1919  was  in- 
cluded with  the  early  eastern  material  cov- 
ered above;  see  pp.  2S-30). 


TABLE  3 

Statistical  distance,  D2,  between  C.  lupus, 
C.  latrans,  and  C.  rufus. 


C.  lupus 

C 

latrans 

A.    Males 

C.   latrans  

C.   rufus    . 

71.9 

18.6 

22.2 

B.  Females 

C.  latrans  

C.   rufus  

63.4 

16.9 

18.8 

These  115  skulls  were  compared  individ- 
ually, by  multivariate  analysis,  to  the  standard 
series  of  C.  lupus,  C.  latrans,  and  C.  familiaris. 
Their  distribution  (Fig.  15),  like  that  of  the 
earlier  material  from  the  southeast  (Fig.  11), 
is  intermediate  to  the  graphical  limits  of  C. 
latrans  and  C.  lupus.  There  is  no  overlap 
with  the  coyote  sample,  but  three  male  and 
three  female  specimens,  all  from  the  Ozark 
region  of  Arkansas  and  Missouri,  fall  within 
the  range  of  variation  of  the  gray  wolf.  There- 
fore, on  the  basis  of  a  multivariate  analysis 
involving  15  cranial  measurements,  complete 
separation  of  C.  lupus  and  C.  rufus  is  not 
possible.  The  amount  of  overlap  between  the 
two,  however,  is  much  less  than  that  found 
among  the  various  subspecies  of  C.  lupus. 

Measurements  of  these  115  gregoryi  are 
listed  in  appendix  B  (part  5).  Means  of  males 
are  compared  with  those  of  other  series  in  a 
ratio  diagram  (Fig.  13).  Size  and  proportion 
in  both  the  early  red  wolf  and  the  later  series 
of  gregoryi  are  seen  to  match  closely,  and  to 
differ  from  C.  lupus  and  C.  latrans.  There  is 
no  indication  that  the  later  and  more  nor- 
therly concentrated  series  of  gregoryi  is  sub- 
stantially different  or  any  more  coyotelike 
than  the  early  Louisiana  material.  These  two 
series  thus  were  combined  into  a  single  group 
that  was  considered  as  a  standard  sample  of 
C.  rufus.  The  D-  statistics  comparing  the 
three  total  standard  samples  of  C.  lupus,  C. 
latrans,  and  C.  rufus,  were  calculated  and 
are  shown  in  table  3.    The  intermediate  val- 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


33 


5- 
4- 
3- 

2- 


-2- 
-3- 
-4- 

-5- 

-6 
-7 

-8 
-9 


-4     -3   -2     -I 


— i— 
0 


-i 1 1 1 r- 

2      3     4      5      6 


Fig.  16. — Multivariate  comparison  of  886  individ- 
ual specimens  of  Canis  from  the  south-central  United 
States,  to  the  ranges  of  variation  of  the  total  series 
of  pre-1930  C.  rufus  gregoryi,  the  total  series  of  C. 
latrans  from  northern  and  western  North  America, 
and  C.  familiaris.   The  dashed  lines  show  the  margins 

ues  of  the  red  wolf  are  almost  as  near  to 
those  of  the  coyote  as  to  those  of  the  gray 
wolf. 

Influence  of  the  Domestic  Dog  on  Wild  Canis 
of  the  Southeast 

Since  it  has  been  concluded  that  C.  rufus 
is  specifically  distinct  from  C.  lupus,  and 
since  there  is  no  evidence  that  C.  lupus 
existed  in  the  southeastern  United  States  in 
the  twentieth  century,  the  gray  wolf  need  not 
be  considered  further  in  discussion  of  the 
systematic  problems  of  this  region.  Hence- 
forth, this  section  of  the  paper  deals  primarily 
with  the  relationships  between  C.  rufus  and 
C.  latrans,  and  with  their  comparative  status 


-3 


■I 


- r— 

0 


I 


— 1 1 1- 

2      3     4 


of  the  statistical  distribution  of  879  of  the  individual 
specimens.  The  seven  exceptions  are  from  Arkansas 
( A ) ;  Louisiana  ( L ) ;  Missouri  (  M ) ;  Lavaca  County, 
Texas  (T);  and  Van  Zandt  County,  Texas  (Z).  See 
appendix  A  (part  11)  for  additional  details.  Males 
are  right,  females  left. 

in  the  southeast.  Before  proceeding,  however, 
the  question  of  interbreeding  between  C. 
familiaris  and  wild  Canis  should  be  covered. 
In  order  to  evaluate  the  influence  of  the 
domestic  dog,  the  three  previously  estab- 
lished samples  of  C.  familiaris,  C.  latrans,  and 
C.  rufus  were  compared  as  groups  with  886 
other  individual  skulls  collected  in  the  south- 
eastern and  south-central  states.  All  of  these 
specimens  had  been  previously  identified  in 
collections  as  wild  Canis,  and  they  include 
the  members  of  nearly  all  of  the  samples 
referred  to  in  the  following  account  of  the 
southeastern  situation.  The  graphical  results 
of  the  analyses  are  shown  in  figure  16.  There 
is  no  statistical  blending  of  wild  southeastern 


34 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


Canis  and  C.  familiaris,  and  no  suggestion  of 
massive  introgression  from  the  domestic  dog 
into  any  of  the  wild  populations.  Only  seven 
specimens  have  statistical  positions  or  D2 
values  indicating  hybridization  involving  C. 
familiaris   (see  appendix  A,  part   11). 

Pre-1930  Relationship  of  Red  Wolf  and 

Coyote  in  Arkansas,  Missouri,  and 

Oklahoma 

With  the  gray  wolf  and  domestic  dog 
eliminated  from  the  picture,  we  now  are 
dealing  exclusively  with  the  relationship  of 
the  red  wolf  and  coyote.  As  an  initial  step, 
standard  series  of  both  these  species  were 
compared  with  each  other  by  multivariate 
analysis.  For  the  red  wolf,  these  series  con- 
sisted of  the  previously  used  70(3)  males 
and  55(2)  females  collected  before  1930  and 
identified  as  C.  rufus  gregoryi.  For  the  coy- 
ote, the  previously  established  standard  sam- 
ple of  western  material  was  used,  except  that 
one  male  specimen  was  withheld  for  consid- 
eration at  a  later  point.  The  comparative 
series  of  C.  latrans  thus  consisted  of  165(2) 
males  and  111(4)  females.  The  results  of 
these  analyses,  demonstrating  clear  separa- 
tion between  the  two  species,  are  depicted  in 
figure  17. 

All  remaining  group  comparisons  involv- 
ing problems  in  the  southeast  are  based  on 
the  statistical  distribution  of  C.  rufus  and  C. 
latrans  shown  in  figure  17.  All  other  speci- 
mens, mostly  those  collected  farther  to  the 
west  or  later  in  time  than  the  standard  sam- 
ple of  C.  rufus,  are  evaluated  on  the  basis  of 
their  relative  distance  from  this  sample  and 
from  the  standard  sample  of  C.  latrans.  There 
is  no  objective  manner  of  setting  definite 
limits  to  the  ranges  of  variation  shown  by 
red  wolves,  coyotes,  and  hybrids  between  the 
two.  It  would  therefore  be  meaningless  to 
attempt  to  assign  each  and  every  specimen 
to  one  of  these  three  categories.  Therefore  I 
decided  to  examine  each  group  of  specimens, 
collected  in  a  given  area  and  over  a  certain 


RUFUS 


3 

2 

I 

0 
-I 
-2 
-3  - 


-3 


-I 


i 
0 


I 


3     4      5      6 


Fie.  17. — Multivariate  comparison  of  the  series  of 
pre-1930  C.  rufus  gregoryi  and  the  series  of  C.  latrans 
from  northern  and  western  North  America.  Only 
the  margins  of  the  range  of  variation  of  each  species 
are  shown.  Males  are  above,  females  below;  C.  rufus 
to  the  right,  C.  latrans  to  the  left.  All  subsequent 
graphs  of  multivariate  analyses  that  involve  the 
southeastern  or  south-central  United  States  are  based 
on  the  illustrations  shown  here.  The  numbers  along 
the  vertical  and  horizontal  axes  are  canonical  coordi- 
nates. The  position  of  a  single  female  C.  rufus  is  not 
included  within  the  range  of  variation  shown  for 
that  species.  The  coordinates  of  this  specimen  fall 
beyond  the  coverage  of  the  computer  plot,  though 
not  in  the  direction  of  the  range  of  variation  of  C. 
latrans.  The  specimen  is  large  and  might  possibly 
represent  a  mislabeled  male.  The  position  of  this 
specimen  is  among  those  plotted  for  females  in  Fig. 
15. 

period,  and  to  try  to  interpret  relative  affinity 
on  the  basis  of  the  total  group  position. 

We  have  already  seen  that  a  large  group 
of  specimens  identified  as  C.  rufus  gregoryi 
may  be  clearly  distinguished  from  a  large 
series  of  western  coyotes.  The  great  majority 
of  these  specimens  referred  to  gregoryi  were 
collected  between  1919  and  1929  in  the 
Ozark-Ouachita  uplands  of  Arkansas,   south- 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


35 


em  Missouri,  and  eastern  Oklahoma.  Coyotes 
also  were  found  along  the  western  and  north- 
ern edges  of  this  region,  but  the  original 
limits  of  their  range  are  unknown.  According 
to  Audubon  and  Bachman  (1851:152),  C. 
latrans  was  "well  known  throughout  the  west- 
ern part  of  the  States  of  Arkansas  and  Mis- 
souri." 

The  coyotes  of  this  region  are  called  C. 
latrans  frustror.  This  name  was  given  by 
Woodhouse  ( 1851 )  to  a  canid  from  the 
Cimarron  River,  about  100  miles  west  of  Fort 
Gibson,  Oklahoma.  For  a  while,  there  was 
uncertainty  regarding  whether  frustror  ap- 
plied to  the  coyote  or  to  the  larger  wolf 
group.  Merriam  (1897)  considered  the  holo- 
type  to  be  a  coyote,  but  Bailey  (1905:175) 
subsequently  noted:  "A  series  of  topotypes 
of  frustror  secured  since  at  Red  Fork,  Ind.  T., 
shows  it  to  be  a  widely  different  species, 
more  nearly  related  to  Canis  rufus."  Bailey 
seems  not  to  have  considered  the  possibility 
that  both  red  wolf  and  coyote  could  have  oc- 
curred together  in  the  same  vicinity.  Jackson 
(1951:271)  considered  frustror  a  subspecies 
of  C.  latrans  with  a  range  from  eastern  Kan- 
sas and  Missouri  south  to  the  Texas  Gulf 
coast. 

Also,  according  to  Jackson  (1951:274): 
"A  very  few  specimens  from  the  Ozark  re- 
gion of  Arkansas  and  Missouri  superficially 
hint  that  there  may  be  possible  hybridization, 
but  probably  not  intergradation,  with  the 
Mississippi  Valley  red  wolf,  Canisniger  gre- 
goryi  Goldman,  in  that  region."  And  Law- 
rence and  Bossert  ( 1967 )  found  a  series  of 
eight  skulls,  collected  in  1921  at  Fallsville, 
Newton  County,  Arkansas,  and  identified  as 
C.  rufus  gregonji,  to  "span  the  whole  range 
of  variation  from  coyote  to  wolf." 

As  stated  above,  I  did  not  include  15 
skulls,  identified  as  gregonji,  in  my  standard 
sample  of  C.  rufus,  because  they  seemed  un- 
usually small  in  greatest  length.  These  3(1) 
males  and  12(1)  females  were  all  obtained 
in  Arkansas  and  eastern  Oklahoma  between 
1919   and   1929    (see   appendix   A,   part    12). 


They  were  compared  individually  to  the 
standard  red  wolf  and  coyote  series,  and  their 
relative  statistical  positions  are  plotted  in 
figure  IS.  Twelve  of  the  specimens  cluster 
within  or  near  the  range  of  variation  of  C. 
rufus.  It  is  impossible  to  say  whether  these 
specimens  are  small  gregonji  or  represent  ge- 
netic influence  from  C.  latrans.  But  the  fact 
that  their  positions  are  concentrated  close  to 
C.  rufus,  and  do  not  form  an  evenly  distrib- 
uted bridge  between  red  wolf  and  coyote, 
suggests  the  former  possibility.  Although 
these  12  skulls  are  short,  compared  to  most 
C.  rufus,  multivariate  evaluation  of  all  meas- 
urements indicates  affinity  to  this  species.  Of 
the  other  three  specimens,  one  taken  at  Falls- 
ville,   Newton    County,    Arkansas,    occupies 


I-  3 

LATRANS 


RUFUS 


□ 


i 


~\ 1 — r- 

2      3     4 


-i r- 

5     6 


Fig.  18. — Multivariate  positions  of  certain  indi- 
vidual specimens  relative  to  the  ranges  of  variation 
of  the  series  of  C.  rufus  and  C.  latrans  shown  in  Fig. 
17.  Black  dots,  C.  latrans  frustror  taken  prior  to 
1930  in  Missouri;  triangles  and  squares,  specimens 
originally  identified  as  C.  rufus  gregoryi  that  have 
unusually  short  greatest  lengths.  The  squares  indi- 
cate specimens  that  probably  represent  C.  latrans  or 
red  wolf-coyote  hybrids. 


36 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


such  an  intermediate  position  (D2  from  coy- 
ote 23.4,  from  red  wolf  24.7)  that  it  almost 
certainly  represents  a  hybrid.  The  two  re- 
maining specimens  appear  to  be  coyotes. 

All  of  the  questionable  specimens  were 
obtained  in  Arkansas  and  eastern  Oklahoma. 
In  southeastern  Texas,  Louisiana,  and  states 
farther  east,  the  only  specimens  of  wild  Cards 
taken  prior  to  1930  are  clearly  referable  to 
C.  rufus.  In  Missouri,  however,  in  addition 
to  the  19  specimens  that  were  identified  as 
C.  rufus  gregoryi  and  used  in  my  standard 
sample  of  red  wolves,  nine  skulls  were  col- 
lected from  1923  to  1925  and  originally  iden- 
tified as  C.  latrans  frustror  (see  appendix  A, 
part  13 ) .  They  were  taken  in  the  same  period 
and  in  approximately  the  same  area  of  the 
southeastern  part  of  the  state  (see  Fig.  14) 
as  the  series  of  gregoryi.  The  multivariate 
positions  of  these  specimens  all  fall  within 
the  range  of  variation  of  the  standard  coyote 
sample  (Fig.  18).  Therefore,  as  first  pointed 
out  by  Paradiso  and  Nowak  (1972a),  there 
is  direct  evidence  that  C.  rufus  and  C.  latrans 
occurred  sympatrically  in  this  area,  without 
intergradation  or  hybridization. 

Until  now,  this  section  of  the  paper  has 
dealt  only  with  red  wolves  identified  as  C.  r. 
gregoryi,  and  not  with  C.  r.  rufus.  Most  of 
the  pre- 1930  specimens  that  Goldman  as- 
signed to  the  latter  named  subspecies  were 
collected  in  Texas,  but  a  few  were  taken  in 
Oklahoma  and  northwestern  Arkansas.  In 
addition,  Hall  and  Kelson  (1952:340-341) 
assigned  two  specimens  taken  in  1923  and 
1924  at  Reeds  Spring,  Stone  County,  south- 
western Missouri,  to  C.  r.  rufus,  although 
Goldman  had  listed  them  as  C.  r.  gregoryi. 
McCarley  (1962)  suggested  that  the  taxon 
C.  r.  rufus  might  actually  represent  the  result 
of  hybridization  between  C.  r.  gregoryi  and 
C.  latrans,  but  Paradiso  and  Nowak  (1972a) 
continued  to  recognize  its  validity. 

Since  specimens  assigned  to  C.  r.  rufus 
are  on  the  whole  smaller  and  more  narrowly 
proportioned,  and  hence  more  coyotelike  than 
most    specimens    of   gregoryi,    there   was    no 


need  herein  to  depict  a  direct  comparison  of 
C.  r.  rufus  with  the  standard  sample  of  C. 
lupus.  But  one  skull  (USNM  8098),  obtained 
sometime  in  the  nineteenth  century  at  Chero- 
kee Town,  in  what  is  now  Garvin  County, 
central  Oklahoma,  within  the  designated 
range  of  C.  r.  rufus,  is  of  special  interest.  It 
was  placed  in  the  National  Museum's  collec- 
tion of  C.  lupus  nuhilus,  and  was  not  in- 
cluded by  Goldman  (1944)  in  his  list  of 
specimens  examined.  Nevertheless  he  did 
handwrite  "C.  rufus"  on  the  specimen  tag. 
My  comparison  of  this  skull  with  standard 
series  of  C.  lupus,  C.  latrans,  and  C.  familiaris 
shows  that  its  position  is  close  to  that  of  other 
old  southeastern  material  identified  as  C. 
rufus  (Fig.  11). 


s 

LATRANS 


RUFUS 


□ 


i 


3     4 


Fig.  19. — Multivariate  positions  of  certain  indi- 
vidual specimens  relative  to  the  ranges  of  variation  of 
the  series  of  C.  rufus  and  C.  latrans  shown  in  Fig.  17. 
Squares  and  triangles,  specimens  from  eastern  Okla- 
homa and  Arkansas  originally  identified  as  C.  rufus 
rufus;  S,  specimens  from  Reeds  Spring,  Stone  County, 
Missouri;  K,  specimen  from  Garvin  County,  Okla- 
homa; black  dots,  C.  latrans  from  western  and  central 
Oklahoma;  E,  C.  latrans  from  eastern  Oklahoma. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


37 


Figure  19  depicts  the  multivariate  posi- 
tions of  eight  males  identified  as  C.  r.  rufus 
(including  USNM  8098).  Only  males  of  ru- 
fus were  available  from  Arkansas,  Oklahoma, 
and  Missouri  (see  appendix  A,  part  14).  Six 
fall  well  within  the  limits  of  C.  rufus,  whereas 
two  of  the  Oklahoma  skulls  have  intermediate 
positions  (D-  from  C.  rufus  28.S  and  31.4, 
from  C.  latrans  26.8  and  31.3).  Although 
available  data  are  thus  inconclusive  regarding 
the  status  of  C.  r.  rufus  in  this  region,  some 
of  the  animals  obtained  here  are  indistin- 
guishable from  red  wolves  taken  farther  east. 
A  number  of  skulls  identified  as  C.  latrans 
also  were  taken  prior  to  1930  in  Oklahoma, 
mostly  in  the  western  and  central  parts  of  the 
state,  and  in  an  adjoining  county  of  Texas 
(see  Fig.  14;  appendix  A,  part  15).  Figure 
19  shows  the  relative  statistical  positions  of 
15(7)  males  and  12(6)  females  to  fall  mainly 
within  the  limits  of  the  standard  sample  of 
western  C.  latrans.  Two  females  from  Creek 
and  Tulsa  counties,  northeastern  Oklahoma, 
are  also  included  within  these  limits.  Inter- 
estingly, of  a  series  of  three  specimens  taken 
at  Red  Fork,  Tulsa  County,  Oklahoma,  in 
1904-1905,  one  is  statistically  identical  to  the 
coyote,  one  is  identical  to  the  red  wolf,  and 
one  is  intermediate  in  characters. 

Although  there  is  evidence  of  early  hy- 
bridization between  C.  rufus  and  C.  latrans 
at  certain  localities  in  Arkansas  and  eastern 
Oklahoma,  genetic  exchange  appears  to  have 
remained  very  limited  before  1930.  Nearly 
all  specimens  taken  until  then  in  the  lower 
Mississippi  Valley,  and  identified  as  C.  rufus, 
are  statistically  separate  from  western  coyotes 
and  from  coyotes  taken  in  the  south-central 
United  States.  Hence  the  sum  of  available 
evidence  indicates  that  the  species  C.  rufus. 
in  essentially  unmodified  form,  survived 
through  the  1920's  in  this  region.  Rut  by  that 
period  the  species  was  under  heavy  pressure 
from  man  and  already  was  making  its  last 
stand  in  the  Ozark-Ouachita  uplands.  Refore 
proceeding  with  a  discussion  of  the  red  wolf's 
decline,  however,  it  is  necessary  first  to  de- 


scribe   the    complex    pre-1930    situation    in 
Texas. 

History  of  Texas  Canis 

As  we  have  seen,  Audubon  and  Rachman 
(1851)  wrote  that  the  "Red  Texan  Wolf" 
occurred  from  northern  Arkansas  through 
Texas  and  into  Mexico,  although  the  only 
specific  place  they  mentioned  was  15  miles 
west  of  Austin,  where  Goldman  (1944:488) 
fixed  the  type  locality  for  C.  r.  rufus.  Other 
early  naturalists  reported  on  the  presence  of 
wolves  in  this  same  general  area,  and,  like 
Audubon  and  Bachman,  observed  consider- 
able local  variation  in  color.  For  example, 
Roemer  (1849:80),  referring  to  an  1846  visit 
to  a  plantation  in  what  is  now  Colorado 
County,  Texas,  wrote  that  "the  owner  of  the 
farm  offered  several  wolf  skins  for  sale.  He 
had  taken  them  from  wolves  recently  caught 
in  steel  traps.  The  pelts  were  of  various  col- 
ors, one  black,  the  other  yellow,  and  still 
another  greyish  brown.  The  farmer  informed 
us  that  such  variation  in  color  was  quite  com- 
mon among  the  larger  wolves.  They  were 
very  plentiful  in  the  forest  surrounding  his 
house  and  a  number  of  hogs  had  been  killed 
by  them." 

References  are  few  as  to  how  far  east  in 
Texas  the  coyote  originally  occurred,  but 
there  is  no  evidence  of  its  former  presence 
beyond  the  prairies.  One  early  source 
(Fisher,  1841:33)  discussing  Rrazoria  County 
in  1840,  stated:  "The  large  black  wolf 
abounds  in  the  country;  but  the  small  prairie 
wolf  of  the  western  states,  I  think,  is  seldom, 
if  ever,  found  so  far  south." 

Bailey's  detailed  biological  survey  of  Texas 
(1905:171-177)  included  the  following  refer- 
ences to  Canis  in  the  state. 

C.  lupus. — Still  common  over  most  of  the  plains 
and  mountain  country  of  western  Texas,  mainly  west 
of  the  one  hundredth  meridian. 

C.  latrans. — More  or  less  common  over  at  least 
middle  and  southern  Texas  and  apparently  eastward 
on  strips  of  prairie  as  far  as  Gainesville  [Cooke 
County]  and  Richmond  [Fort  Bend  County].  There 
are  vague  reports  of  a  small  wolf  occurring  farther 
east  on  the  coast  prairie  even  to  the  border  of  Louisi- 


38 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


ana,  but  specimens  are  needed  before  these  reports 
can  be  associated  with  definite  species.  East  of  the 
semiarid  mesquite  region  coyotes  are  rare  and  prob- 
ably mere  stragglers.  True  to  their  name  of  prairie 
wolf,  they  do  not  enter  the  timbered  country  to  any 
extent,  although  at  home  in  the  scrub  oak,  juniper, 
mesquite,  and  chaparral,  as  well  as  over  the  open 
prairies  of  the  southern  part  of  the  state. 

C.  rufus. — A  definite  range  can  be  assigned  the 
species,  covering  the  whole  of  southern  Texas  north 
to  the  mouth  of  the  Pecos  and  the  mouth  of  the 
Colorado,  and  still  farther  north  along  the  strip  of 
mesquite  country  east  of  the  plains,  approximately 
covering  the  semiarid  part  of  the  Lower  Sonoran 
zone.  As  yet  there  are  no  specimens  to  show  whether 
these  wolves  extend  into  the  more  arid  region  west 
of  the  Pecos.  While  apparently  nowhere  overlapping 
the  range  of  the  larger,  lighter-colored  'lobo'  or 
'loafer'  [C.  lupus]  of  the  plains,  they  take  its  place 
to  the  south  and  east  as  soon  as  the  plains  break 
down  and  the  scrub  oak  and  mesquite  country  begins, 
but  their  whole  range  is  shared  with  the  coyote.  The 
ranchmen  invariably  distinguish  between  them  and 
coyotes,  and  with  good  reason,  for  the  wolves  kill 
young  cattle,  goats,  and  colts  with  as  much  regularity 
as  the  coyotes  kill  sheep.  While  paying  a  bounty  of 
SI  or  S2  for  coyotes,  the  ranchmen  usually  pay  S10 
or  $20  for  red  wolves. 

Although  Bailey  considered  C.  rufus  to 
occupy  all  of  southern  Texas,  and  to  share  its 
entire  range  with  C.  Jatrans,  some  modifica- 
tion of  this  delineation  is  suggested  by  the 
Bureau  of  Biological  Survey  field  reports  (on 
file,  National  Fish  and  Wildlife  Laboratories, 
U.S.  National  Museum  of  Natural  History) 
upon  which  his  published  work  was  based. 
These  reports  (made  by  Bailey,  J.  H.  Gaut. 
A.  H.  Howell,  W.  Lloyd,  and  H.  C.  Ober- 
holser  from  1891  to  1905)  were  prepared  at 
a  time  when  little  was  known  about  the  sys- 
tematics  of  Canis  in  southern  Texas.  They 
refer  to  C.  rufus  as  either  a  large  species  of 
coyote  or  a  wolf.  But  more  important  than 
the  names  applied  is  the  fact  that  the  pres- 
ence of  both  this  larger  species  of  Canis  and 
a  distinct  smaller  species  (C.  latrans)  was 
generally  recognized.  The  reports  suggest 
that  the  larger  species,  or  the  animal  now 
referred  to  as  the  red  wolf,  was  originally 
common  only  along  the  Texas  coast  above 
Nueces  Bay,  and  that  C.  latrans  was  rare  or 
entirely  absent  from  this  area.  Farther  in- 
land, in  the  more  arid  parts  of  southern 
Texas,  the  red  wolf  progressively  diminished 


in  numbers,  whereas  the  coyote  became 
abundant.  The  reports  do  not  provide  any 
insight  on  the  original  situation  farther  north 
in  the  Edwards  Plateau  area. 

Examination  of  Pre-1930  Material 
from  South  Texas 

The  main  problem  presented  by  the  above 
information  is  the  relationship  between  the 
red  wolf  and  coyote  where  their  ranges  met 
along  the  Texas  coast,  and  farther  inland 
toward  the  Rio  Grande  Valley.  Unfortu- 
nately, the  number  and  distribution  of  avail- 
able pre-1930  specimens  are  not  sufficient  for 
thorough  analysis  of  the  situation.  Although 
a  useful  series  of  coyotes  was  collected  in 
south  Texas,  few  skulls  were  saved  from  the 
critical  areas  immediately  to  the  north  of 
Nueces  Bay  and  inland  toward  San  Antonio. 

A  group  of  eight  skulls,  previously  identi- 
fied as  C.  rufus  rufus,  was  taken  in  1900  and 


3 
2 
I 

0 
-I 
-2 
-3 


3 

LATRANS 


RUFUS 


A 


-3    -2 


-I 


0 


I 


Fie.  20. — Multivariate  positions  of  pre-1930  speci- 
mens from  southern  Texas  relative  to  series  of  C. 
rufus  and  C.  latrans.  Triangles,  C.  rufus  rufus;  black 
dots,  C.  latrans  tcxcnsis. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


39 


1904  near  the  Texas  coast  in  Calhoun,  Colo- 
rado, and  Liberty  counties  (see  appendix  A, 
part  16).  The  multivariate  positions  of  these 
specimens  are  shown  in  figure  20.  Clearly, 
these  specimens  have  close  affinity  with  those 
of  the  standard  red  wolf  sample,  which  were 
taken  farther  east  and  identified  as  C.  rufus 
gregoryi,  but  they  are  smaller  and  more  nar- 
rowly proportioned.  In  other  words,  they  fit 
Goldman's  (1944:487)  description  of  the  sub- 
species C.  r.  rufus.  A  specimen  taken  in  1906 
in  Hardin  County,  extreme  southeastern 
Texas,  and  identified  as  C.  r.  gregoryi,  was 
incorporated  above  in  my  standard  red  wolf 
sample.  There  are  no  other  usable,  pre-1930 
specimens  of  red  wolves  obtained  in  southern 
and  coastal  Texas. 

Skulls  of  31(5)  males  and  26(8)  females 
labeled  as  C.  latrans  texensis  were  taken  from 
1891  to  1918  in  south  Texas  (see  map,  Fig. 
14;  appendix  A,  part  17),  more  than  half 
coming  from  Nueces  County.  Most  of  these 
specimens  fall  within  the  range  of  variation 
of  western  C.  latrans  (Fig.  20).  Although 
three  Nueces  County  specimens  have  statisti- 
cal positions  that  are  removed  in  the  direction 
of  the  standard  red  wolf  sample,  their  D2 
values  confirm  closer  affinity  to  C.  latrans. 
Therefore,  although  there  is  suggestion  of 
limited  hybridization,  early  specimens  from 
south  Texas  may  be  separated  into  two  dis- 
tinct groups,  representing  the  species  C.  la- 
trans and  C.  rufus. 

Examination  of  Pre-1930  Material  from 
Central  Texas 

Early  specimens  from  the  Edwards  Pla- 
teau area  of  central  Texas  are  not  so  easily 
divisible  into  recognized  species,  and  have 
been  the  cause  of  much  confusion.  A  report 
by  Allen  (1896:75-76)  is  pertinent,  since  it 
contains  both  the  earliest  account  of  possible 
hybridization  in  the  area,  and  an  early  refer- 
ence to  the  presence  there  of  three  kinds  of 
Canis.  Allen's  informant,  H.  P.  Attwater,  said 
that  Canis  lupus  was  "formerly  common  in 
Bexar  County,  but  I  have  not  heard  of  their 


occurrence  here  for  several  years.  They  are 
still  found  in  the  broken,  hilly  country  north- 
west of  San  Antonio,  particularly  in  Edwards 
County."  And,  regarding  C.  latrans,  he  stated: 
"In  Kerr  County  and  adjoining  counties  they 
are  the  'thorn  in  the  side,'  of  the  sheepmen. 
Mr.  Lacey  [Howard  Lacey,  a  rancher  said  to 
be  a  careful  and  reliable  observer]  says  that 
the  Coyotes  of  that  region  are  different  from 
the  Coyotes  of  the  prairies,  being  much 
larger.  They  are  believed  by  the  ranchmen 
to  be  a  cross  between  the  'Lobo'  (Wolf)  and 
the  Coyote.  Two  years  ago,  when  the  bounty 
act  was  in  force,  the  regular  'Lobo'  price  was 
allowed  for  the  large  Coyotes  of  the  rocky 
region  to  the  northwestward  of  San  Antonio." 

With  regard  to  the  red  wolf  and  coyote 
in  central  Texas,  Goldman  (1944:481)  re- 
ported: "Specimens  collected  in  the  vicinity 
of  Llano,  Tex.,  include  typical  examples  of 
both  species  and  individuals  not  sharply  dis- 
tinctive of  either.  Close  approach  in  essential 
details  and  the  apparent  absence  of  any  in- 
variable unit  character  suggests  the  possibil- 
ity of  hybridism  in  some  localities  in  Texas." 
McCarley  (1962)  considered  a  series  of  eight 
skulls  from  the  vicinity  of  Llano  to  suggest 
the  occurrence  of  hybridization.  Paradiso  and 
Nowak  (1972a)  concluded  that  specimens 
taken  throughout  the  Edwards  Plateau  area 
of  Texas  between  1890  and  1918  represented 
a  hybrid  swarm  of  C.  rufus  x  C.  latrans.  For 
purposes  of  this  paper,  all  available  early 
material  from  central  Texas,  as  well  as  that 
collected  in  western  Texas  in  the  same  period, 
was  re-examined  and  subjected  to  more  criti- 
cal statistical  evaluation. 

To  facilitate  an  interpretation  of  the  situa- 
tion, the  central  and  west  Texas  material  was 
divided  on  an  arbitraiy  geographic  basis.  Fig- 
ure 21  shows  multivariate  positions  of  speci- 
mens collected  in  Texas  counties  to  the  west 
and  just  north  of  Tom  Green  County;  figure 
22  does  the  same  for  skulls  taken  in  Tom 
Green  County;  and  figure  23  shows  positions 
of  specimens  taken  in  the  following  counties 
of  central  Texas:    Blanco,   Burnet,   Coleman, 


40 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


-3  - 


8 

LATRANS 


-3    -2 


-I 


- r- 
0 


I 


H 1- 

3     4 


RUFUS 


western  Texas,  and  into  Tom  Green  County. 
Goldman  (1944)  assigned  to  C.  r.  riifus 
a  male  from  22  miles  north  of  Sheffield,  Pecos 
County,  which  is  about  100  miles  farther  west 
than  the  closest  other  record  of  C.  rufus 
listed  by  him.  Paradiso  and  Nowak  ( 1972a ) 
considered  the  skull  to  represent  a  coyote 
and  to  hardly  be  distinguishable  from  some 
specimens  of  C.  latrans  lestes.  While  making 
a  preliminary  examination  of  my  standard 
coyote  sample,  I  noticed  one  large  skull  of 
lestes  from  Bountiful,  Conejos  County,  south- 
ern Colorado,  that  closely  resembled  the 
Pecos  County  specimen.  The  Colorado  skull 
was  withheld  from  the  coyote  series  in  the 
multivariate  analysis  comparing  the  standard 
samples  of  C.  rufus  and  C.  latrans.  Its  rela- 
tive position  is  plotted  in  figure  21  and  dem- 
onstrates that  a  few  western  coyotes  are  sta- 
tistically well  removed  from  normal  C.  latrans 
in    the    direction    of    C.    rufus.     The    Pecos 


Fig.  21. — Multivariate  positions  of  certain  indi- 
vidual specimens  relative  to  series  of  C.  rufus  and 
C.  latrans.  Black  dots,  C.  latrans  texcnsis  from  west- 
ern Texas;  P,  specimen  from  Pecos  County,  Texas 
originally  identified  by  Goldman  (1944)  as  C.  rufus 
rufus,  but  considered  in  this  paper  to  probably  rep- 
resent C.  latrans  texensis;  C,  specimen  of  C.  latrans 
lestes  from  Conejos  County,  Colorado. 

Concho,  Edwards,  Gillespie,  Kerr,  Llano,  Mc- 
Culloch,  Menard,  San  Saba,  and  Sutton  (see 
also  Fig.  14;  appendix  A,  parts  18-20).  All 
material  was  collected  from  1915  to  1918, 
except  for  the  specimens  from  Edwards,  Gil- 
lespie, and  Kerr  counties,  which  were  taken 
from  1899  to  1906. 

Some  30  males  and  15  females  from  west- 
ern Texas,  each  previously  identified  as  C. 
latrans  texensis,  fall  within  or  near  the  range 
of  variation  of  the  standard  coyote  sample. 
The  skulls  of  36  males  and  14  females  taken 
farther  east  in  Tom  Green  County,  each  also 
previously  identified  as  texensis,  show  the 
same  kind  of  statistical  distribution.  Hence 
there  appears  to  be  no  clinal  shift,  in  the 
direction  of  C.  rufus,  as  the  range  of  C.  la- 
trans passes  from  the  western  states,  across 


3 
2 
I 

0 
-I 
-2 
-3 

3 
2 
I 

0 
-I 
-2 
-3 


I-    S 

LATRANS 


— i r 

-3     -2 


9 


-3    -2     -I 


0 


RUFUS 


Fig.  22. — Multivariate  positions  (black  dots)  of 
individual  pre-1930  specimens  of  C.  latrans  texensis 
from  Tom  Green  County,  Texas,  relative  to  series  of 
C.  rufus  and  C.  latrans. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


41 


County  specimen,  the  position  of  which  is 
close  to  that  of  the  Colorado  skull,  could  be 
such  a  coyote.  The  D2  values  of  both  skulls 
indicate  closer  affinity  to  C.  latrans  than  to 
C.  rufus,  but  there  is  no  way  of  definitely 
determining  whether  the  Pecos  County  speci- 
men represents  some  genetic  influence  from 
the  red  wolf.  The  skin  of  this  specimen  was 
examined,  but  could  not  be  distinguished 
from  those  of  other  coyotes  taken  in  central 
and  western  Texas. 

Central  Texas  is  a  land  of  major  natural 
transition.  Here  the  high  plains  break  down 
into  the  rough  country  of  the  Edwards  Pla- 
teau; the  arid  Sonoran  area  of  the  Lower 
Austral  Life-zone  passes  into  the  humid  Aus- 
troriparian  area;  and  typical  prairie  and  des- 
ert vegetation  merge  with  eastern  deciduous 
forest.  Regarding  the  transition  of  the  vege- 
tation pattern  on  the  Edwards  Plateau,  Bray 
(1904:14)  noted  that  "conditions  vary  enough 
to  give  in  some  places,  as  in  well-watered  and 
sheltered  canyons,  a  relatively  luxuriant 
growth,  while  in  other  situations,  as  upon 
stony  arid  slopes,  there  is  the  scantiest  vege- 
tation." 

At  the  turn  of  the  century,  central  Texas 
also  was  undergoing  pronounced  changes  be- 
cause of  the  influence  of  man.  Overgrazing 
by  cattle  and  sheep,  and  the  frequent  occur- 
rence of  fire,  had  suppressed  the  native  prai- 
ries and  permitted  the  spread  of  mesquite. 
Simultaneously,  clearing  and  cultivation  along 
the  river  bottoms  had  deprived  the  area  of 
its  richest  forest  growth  (Bray,  1904).  And, 
of  course,  the  hand  of  man  was  turned  heav- 
ily against  predatory  animals  such  as  wolves 
and  coyotes. 

The  specimens  of  Cards  collected  in  cen- 
tral Texas  were  originally  identified  as  C. 
rufus  rufus,  C.  latrans  texensis,  and  C.  latrans 
frustror.  Jackson  (1951:271,  279)  considered 
this  last  named  subspecies  to  range  north  of 
Nueces  Bay,  and  to  the  eastern  edge  of  the 
Edwards  Plateau  where  he  recorded  speci- 
mens from  Blanco,  Burnet,  Llano,  and  San 
Saba   counties.    Jackson   wrote   that   texensis 


occurred  to  the  south  of  Nueces  Bay,  but  that 
its  range  extended  northwestward  into  cen- 
tral Texas  (he  listed  specimens  there  from 
Coleman,  Concho.  Gillespie,  Kerr,  McCul- 
loch,  Menard,  and  Sutton  counties).  Gold- 
man (1944:488-489)  thought  that  C.  rufus 
rufus  occurred  throughout  central  Texas,  and 
he  recorded  pre-1930  specimens  from  Burnet, 
Edwards,  Kerr.  Llano,  and  McCulloch  coun- 
ties. 

Those  specimens  from  central  Texas  that 
could  be  subjected  to  multivariate  analysis  in- 
cluded animals  identified  as  the  following: 
C.  latrans  texensis,  male — 13,  female — 12(2); 
C.  latrans  frustror,  male — 9,  female — 15;  C. 
rufus  rufus,  male — 21(1),  female — 5.  Locali- 
ties are  plotted  on  the  map  in  figure  14,  and 
statistical  positions  are  shown  in  figure  23. 
Also  shown  are  positions  of  the  14  skulls  of 
C.  lupus  monstrabilis  collected  in  central  and 
western  Texas,  each  of  which  was  used  above 
in  my  standard  gray  wolf  sample  (most  were 
collected  from  1900  to  1920).  Their  positions 
are  plotted  here  to  emphasize  that,  although 
C.  lupus  was  present  in  the  area,  it  was  dis- 
tinct from  the  animals  being  called  red 
wolves,  and  was  not  a  factor  in  the  problems 
under  discussion. 

The  multivariate  positions  of  the  central 
Texas  skulls  identified  as  texensis,  frustror, 
and  rufus  form  a  statistical  bridge  between 
the  ranges  of  variation  of  standard  C.  latrans 
and  C.  rufus.  There  is  no  meaningful  place 
to  draw  a  line  separating  these  skulls  into 
coyotes  and  red  wolves,  and  we  apparently 
are  dealing  with  some  sort  of  blending  be- 
tween the  two.  Specimens  collected  in  cer- 
tain restricted  localities,  such  as  those  from 
the  vicinity  of  Llano  and  Burnet,  by  them- 
selves bridge  the  statistical  gap  between  red 
wolves  and  coyotes.  After  an  evaluation  of 
these  statistics,  a  visual  re-examination  of  the 
skulls,  and  a  check  of  available  skins  from 
central  Texas,  it  must  be  concluded  that  the 
original  identification  of  these  specimens  was 
arbitrary  and  based  on  an  incomplete  under- 
standing of  the  situation.    In  this  regard  it  is 


42 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


3 
2 
I 

0 
-I 
-2 
-3 


.    LATRANS 


RUFUS 


m 


m 
m 


m 


m 


m 


m 


-4 


~i 1 r 

2      3     4 


5 


6 


—r- 
7 


— 1- 
9 


0 


8 


10 


3 

2 

I 

0- 
-I- 

-2" 
-3- 


LATRANS 


RUFUS 


m 


m 


m 
m 

m 


m 


n 1 r 

2       3      4 


5 


6 


7 


~i r- 

8      9 


-4     -3     -2 


0       I 


10 


Fig.  23. — Multivariate  positions  of  certain  individual  pre-1930  specimens  from  Texas,  relative  to  series  of 
C.  rufus  and  C.  latrans.  Black  dots,  specimens  from  central  Texas  originally  identified  as  C.  latrans  texensis 
or  C.  latrans  frustror;  squares,  specimens  from  central  Texas  originally  identified  as  C.  rufus  rufus;  M,  speci- 
mens of  C.  lupus  monstrabilis  from  central  and  western  Texas.    Males  are  above,  females  below. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


43 


interesting  to  note  an  unusual  sex  ratio  among 
the  available  specimens  of  rufus  and  frustror 
from  the  area.  For  specimens  originally  iden- 
tified as  frustror,  females  outnumber  males 
15  to  9;  but  for  skulls  previously  referred  to 
rufus,  males  outnumber  females  20  to  5.  This 
suggests  to  me  that  the  original  identification 
of  the  specimens  was  based  partly  on  the  bias 
of  assigning  larger  skulls,  predominantly 
males,  to  rufus,  and  smaller  skulls,  predomi- 
nantly females,  to  frustror. 

No  other  group  of  specimens,  at  least 
among  those  collected  before  1930,  falls  to 
such  an  extent  between  the  statistical  limits 
of  C.  latrans  and  C.  rufus.  This  condition 
could  have  resulted  from  one  of  the  following 
factors:  (1)  intergradation  in  central  Texas 
between  the  small  western  coyote  and  the 
larger  eastern  red  wolf;  (2)  long-term  inter- 
breeding, and  the  production  of  a  hybrid 
zone  serving  as  a  bridge  for  the  flow  of  genes 
from  one  species  to  the  other;  (3)  short-term 
interbreeding  caused  possibly  by  drastic  al- 
teration of  the  environment. 

I  do  not  think  that  the  statistical  distribu- 
tion of  specimens  is  indicative  of  intergrada- 
tion and  hence  the  conspecificity  of  red  wolf 
and  coyote.  Recognized  subspecies  of  C. 
latrans  intergrade  throughout  western  North 
America  and  show  no  morphological  overlap 
with  red  wolves.  There  is  no  evidence  of  a 
cline  of  characters  approaching  those  of 
standard  C.  rufus,  even  among  specimens 
taken  as  far  east  as  Tom  Green  County, 
Texas.  The  sudden  breakdown  found  in  the 
limited  area  of  central  Texas  is  highly  atypi- 
cal of  North  American  Canis,  and  does  not 
represent  normal  subspecific  intergradation. 
Furthermore,  as  we  have  seen,  there  is  no 
suggestion  of  original  intergradation  in  any 
other  area  where  the  ranges  of  C.  rufus  and 
C.  latrans  met,  and  there  is  direct  evidence 
of  the  sympatric  occurrence  of  the  two  spe- 
cies in  Missouri  (see  pp.  35-36). 

While  I  therefore  consider  hybridization 
responsible  for  the  situation  in  central  Texas, 
there  is  no  reason  to  believe  that  the  process 


had  been  occurring  over  a  long  period.  Had 
interbreeding  been  going  on  for  many  years 
or  centuries,  its  effects  seemingly  would  have 
spread  beyond  central  Texas. 

Available  evidence  favors  the  theory  that 
hybridization  between  C.  rufus  and  C.  latrans 
was  of  relatively  short-term  occurrence,  and 
probably  had  begun  in  the  latter  part  of  the 
nineteenth  century  in  response  to  man's  dis- 
ruption of  the  habitat  and  his  persecution  of 
native  wild  canid  populations.  This  is  in 
accord  with  earlier  statements  by  McCarley 
(1962)  and  Paradiso  and  Nowak  (1972a). 
Since  central  Texas  was  an  area  of  natural 
transition,  where  the  eastern  red  wolf  and 
western  coyote  would  have  overlapped  in 
range,  if  not  in  habitat,  opportunities  for 
interbreeding  undoubtedly  occurred.  Once 
man  altered  the  environment,  and  at  the  same 
time  attacked  the  wolves  and  coyotes,  ecologi- 
cal and  behavioral  isolation  might  have  brok- 
en down,  and  large-scale  hybridization  be- 
gun. 

According  to  Mayr  (1963:128):  "By  far 
the  most  frequent  cause  of  hybridization  in 
animals  is  the  breakdown  of  habitat  barriers, 
mostly  as  a  result  of  human  interference." 
Mayr  (1963:118-121)  used  the  term  "hybrid 
swarm"  to  describe  populations,  in  such  areas 
of  breakdown  and  interbreeding,  that  form 
a  continuous  bridge  between  two  parent 
species.  This  term  seems  applicable  to  the 
population  of  Canis  in  central  Texas. 

The  exact  original  limits  of  the  ranges 
of  C.  rufus  and  C.  latrans  in  central  Texas  are 
unknown,  partly  because  the  hybridization 
factor  has  obscured  the  picture.  The  red 
wolf,  however,  usually  is  considered  a  species 
of  the  eastern  forests.  In  this  regard  it  is  in- 
teresting to  note  that  the  area  of  central  Texas 
under  discussion  falls  just  within  the  extreme 
western  boundary  of  eastern  forest  elements 
(see  Fig.  14).  Indeed,  oak  trees  first  appear 
immediately  to  the  east  and  south  of  Tom 
Green  County. 

At  this  point  I  would  like  to  summarize 
briefly  the  early  status  of  Canis  in  the  south- 


44 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


east,  as  evaluated  on  the  basis  of  specimens 
collected  prior  to  1930.  The  red  wolf,  a  spe- 
cies distinct  from  the  gray  wolf  of  northern 
and  western  North  America,  was  found  along 
the  Atlantic  coast  and  westward  probably  to 
the  edge  of  the  prairies.  Large  series  of  spe- 
cimens show  that  before  1930  the  red  wolf 
in  the  lower  Mississippi  Valley  was  easily 
separable  from  the  western  coyote.  The  two 
species  occurred  sympatrically,  or  at  least 
showed  no  tendency  to  intergrade  in  most 
areas  where  their  ranges  approached.  A  small 
percentage  of  specimens  taken  in  eastern 
Oklahoma  and  Arkansas  suggest  that  limited 
hybridization  had  occurred  at  certain  locali- 
ties in  those  states.  In  central  Texas,  how- 
ever, more  extensive  interbreeding  had  re- 
sulted in  the  formation  of  a  hybrid  swarm 
between  the  two  species. 

Survival  of  the  Red  Wolf  from  the 
1930's  to  1950's 

Although  the  red  wolf,  in  unmodified 
form,  seems  to  have  survived  in  the  lower 
Mississippi  Valley  through  the  1920's,  the 
species  had  been  under  heavy  human  pressure 
throughout  most  of  its  range.  The  course  of 
its  decline  was  discussed  by  me  ( 1967,  1970, 
1972,  1974),  Russell  and  Shaw  (1972),  and 
Young  ( 1944 ) .  Wolves  disappeared  from 
Pennsylvania,  Maryland,  West  Virginia,  Vir- 
ginia, North  Carolina,  South  Carolina,  Ten- 
nessee, Kentucky,  Ohio,  Indiana,  and  Illinois 
without  specimens  having  been  saved  which 
would  have  enabled  us  to  determine  their 
systematic  status. 

Farther  to  the  south,  wolves,  probably 
only  C.  rufus,  did  not  survive  much  longer. 
Harper  (1927:315-317)  said  that  the  last 
known  kill  of  C.  r.  floridanus  in  the  Okefino- 
kee  Swamp  of  Georgia  took  place  about  1908, 
although  there  were  later  reports.  There 
seem  to  be  no  other  definite  records  of  wolves 
in  Georgia  after  1900.  According  to  Chap- 
man (1894:345)  wolves  in  Florida  were  al- 
ready "on  the  verge  of  extinction."  The  last 
reported   occurrence   in   the   state,    listed   by 


Young  (1944:25),  was  in  1903  near  the  Ever- 
glades. H.  H.  Bailey  (1930)  wrote  that  a 
wolf  was  killed  about  1918  or  1920  on  the 
Osceola  side  of  the  Kissimee  River. 

Howell  (1921:30)  reported  that  wolves  in 
Alabama  were  "on  the  verge  of  extinction. 
Their  last  stronghold  appears  to  be  the  rough, 
hilly  country  stretching  from  Walker  County 
northwestward  to  Colbert  County."  The  last 
recorded  kill  took  place  south  of  Cherokee, 
Colbert  County  in  1917.  Apparently,  how- 
ever, wolves  held  out  for  a  while  longer  in 
the  state.  In  1937  and  1938  reports  by  the 
Predator  and  Rodent  Control  branch  of  the 
U.S.  Bureau  of  Biological  Survey  (on  file  at 
the  offices  of  the  Division  of  Animal  Damage 
Control,  U.S.  Fish  and  Wildlife  Service, 
Washington,  D.C. )  it  is  said  that  in  Alabama 
nine  wolves  were  killed  in  1937  and  three  in 
1938.  Holliman  (1963:242)  reported  a  speci- 
men of  C.  rufus  collected  at  Livingston,  Sum- 
ter County,  western  Alabama  in  1944.  The 
multivariate  position  of  its  skull  falls  within 
the  statistical  limits  of  the  standard  red  wolf 
sample  ( Fig.  24 ) . 

Wolves  apparently  disappeared  from  most 
of  the  higher  country  of  Mississippi  at  an 
early  date,  but  survived  until  comparatively 
recently  along  the  Mississippi  River  and  Gulf 
Coast.  Jenkins  (1933:125-127)  reported  "but 
few  wolves  in  Mississippi  from  1892  to  the 
present  time."  He  added  that  during  the 
great  flood  of  1927  many  wolves  crossed  into 
southwestern  Mississippi  from  Louisiana.  Ac- 
cording to  Young  (1944:29),  these  wolves 
were  still  present  in  1932  and  were  preying 
on  livestock.  Goldman  (1944:485)  reported, 
but  did  not  examine,  a  specimen  of  C.  rufus 
gregoryi  taken  at  Biloxi  in  1931.  I  found  this 
skull  to  fall  squarely  within  the  range  of  vain 
ation  of  standard  C.  rufus  (Fig.  24). 

Very  little  information  is  available  con- 
cerning the  possible  former  occurrence  of 
the  red  wolf  in  Kansas.  Lantz  (1905)  re- 
ferred to  "Canis  ater"  as  "Once  abundant. 
Still  found  in  a  few  scattered  sections  of  the 
state."    Cockrum   (1952:229)   cited  reports  of 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


45 


RUFUS 


Fig.  24. — Multivariate  positions  of  specimens  of 
C.  rufus,  and  two  hybrids,  taken  from  the  1930's  to 
the  1950's.  A,  Alabama  gregoryi;  K,  Arkansas  gre- 
goryi;  L,  Louisiana  gregoryi;  M,  Mississippi  gregoryi; 
O,  Oklahoma  gregoryi;  T,  Texas  gregoryi;  R,  Texas 
rufus;  B,  specimen  of  C.  r.  rufus  taken  in  1954  in 
Brazoria  County,  Texas;  D,  specimens  of  C.  r.  rufus 
taken  in  Madison  County,  Texas  (originally  identified 
as  C.  latrans,  but  considered  in  this  paper  to  repre- 
sent C.  rufus);  S,  specimen  of  C.  r.  rufus  taken  in 
southern  Harris  County,  Texas  (originally  identified 
as  C.  latrans,  but  considered  in  this  paper  to  repre- 
sent C.  rufus);  N,  specimens  of  probable  red  wolf- 
coyote  hybrids  taken  in  northern  Harris  County, 
Texas. 

a  red  wolf  killed  in  Cherokee  County,  ex- 
treme southeastern  Kansas,  in  1908  or  1909, 
and  of  one  heard  howling  there  in  1915. 

The  situation  in  the  Ozark-Ouachita  up- 
lands, and  adjacent  areas,  is  most  confusing 
in  the  1930's  and  1940's,  and  is  discussed  in 
detail  in  the  next  subsection  of  this  paper.  I 
will  first  deal  with  those  areas  in  which  the 
red  wolf  seems  to  have  survived  past  1930  in 
unmodified  form. 

Six  skulls  obtained  in  northern  Louisiana 
from  1935  to  1940  are  indicative  of  the  con- 
tinued presence  of  C.  rufus  in  that  state  (Fig. 


24).  No  other  specimens  of  wild  Canis  taken 
between  1930  and  1950  arc  available  from 
Louisiana.  All  evidence  suggests  that  up 
until  the  last  few  years  of  this  period  the  red 
wolf  maintained  moderate  numbers  over 
much  of  Louisiana.  Ry  the  early  1950's,  how- 
ever, the  species  had  been  decimated  in 
most  areas,  except  for  the  eastern  bottom 
lands  and  southern  marshes  of  the  state 
(Nowak,  1967).  McCarley  (1962)  reported 
specimens  of  C.  rufus  obtained  in  1956  and 
1957  in  Terrebonne  and  Madison  parishes, 
respectively.  The  skull  taken  in  Terrebonne 
Parish,  southern  Louisiana,  falls  within  the 
statistical  limits  of  the  red  wolf  (Fig.  24). 
The  skull  from  Madison  Parish,  northeastern 
Louisiana,  was  not  suitable  for  multivariate 
analysis,  but  does  represent  C.  rufus. 

According  to  Gipson  (1972:4),  red  wolves 
were  present  in  southern  Arkansas  as  late  as 
the  1950's.  A  specimen  taken  in  1942  on  the 
Union-Columbia  county  line,  just  north  of  the 
Louisiana  border,  has  a  multivariate  position 
within  the  limits  of  C.  rufus  (Fig.  24). 

A  male  and  a  female  specimen  taken  in 
1936  near  Rattiest,  McCurtain  County,  ex- 
treme southeastern  Oklahoma  were  reported 
by  McCarley  (1962)  to  be  red  wolves.  These 
two  skulls  are  statistically  well  removed  from 
my  standard  coyote  sample  (Fig.  24),  and 
probably  indicate  continued  survival  of  C. 
rufus  in  the  area. 

The  southeastern  part  of  Texas,  especially 
the  area  known  as  the  Rig  Thicket,  seems  to 
have  been  one  of  the  last  major  refuges  for 
the  red  wolf.  Goldman  ( 1944:4S6-489)  re- 
corded post-1930  specimens  of  C.  rufus  gre- 
goryi from  Hardin,  Newton,  and  Polk  coun- 
ties; and  C.  r.  rufus  from  Rrazoria,  Rrazos, 
Liberty,  Montgomery,  and  Walker  counties. 
In  addition,  Jackson  (1951:275)  listed  speci- 
mens of  C.  latrans  frustror  from  Harris 
County,  in  the  same  area.  Most  of  this  ma- 
terial, plus  two  other  skulls  in  the  National 
Museum  from  Madison  County  that  had  been 
mistakenly  labeled  as  C.  latrans  texensis,  were 
subjected   to   multivariate   analysis.    The   re- 


46 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


suits  (Fig.  24)  confirm  the  survival  of  the 
red  wolf  in  southeastern  Texas.  As  in  other 
cases,  specimens  identified  as  C.  r.  rufus  tend 
to  show  some  statistical  approach  to  C.  la- 
trans.  The  two  Madison  County  specimens 
apparently  represent  C.  r.  rufus.  One  of  the 
Harris  County  skulls  was  collected  at  Genoa 
on  the  south  side  of  Houston.  Although  orig- 
inally identified  as  C.  latrans  frustror,  it  falls 
within  the  range  of  variation  of  the  standard 
red  wolf  sample,  and  I  think  it  actually  repre- 
sents C.  rufus.  The  other  two  Harris  County 
skulls  were  taken  at  Humble,  on  the  north 
side  of  Houston,  and  their  intermediate  sta- 


tistical positions  suggest  genetic  influence  of 
the  coyote  at  that  locality.  All  of  the  above 
listed  Texas  specimens  were  collected  be- 
tween 1930  and  1943,  and  I  examined  very 
little  material  taken  over  the  next  20  years  in 
the  state.  The  skull  of  a  female  taken  in 
eastern  Brazoria  County  in  1954  has  a  multi- 
variate position  suggesting  the  survival  of 
C.  r.  rufus  in  the  area. 

In  summary,  it  appears  that  the  red  wolf 
continued  to  exist  in  parts  of  its  former  range 
from  the  1930's  to  the  1950's.  No  usable  speci- 
mens of  wild  Canis,  other  than  those  men- 
tioned in  this  subsection  of  the  paper,  were 


100 

90 

39 

! 

\ 

• 
[ 

\             '                     1 
3             •                    \C^ 

39 

_%_B_5__^  «J ' 

|*« 

• 

• 

3 
• 

• 

A    A 

< 

f 

•□n5 

A           ) 

A 

A) 

30 

I        A         / 

N.                            • 

A  A  A 
A  AA. 

aaaa 

a 

4          ~~k  ^Xj^)/*^0- 

30 

0                          200 

i                / 

1 

scale  of  miles 

100 

90 

Fig.  25. — Map  showing  localities  of  specimens  taken  from  the  1930's  to  the  1950's  in  the  south-central 
United  States.  Triangles,  C.  rufus;  black  dots,  specimens  originally  identified  as  C.  latrans;  squares,  speci- 
mens originally  identified  as  C.  rufus,  but  considered  in  this  paper  to  represent  hybridization  between  red 
wolf  and  coyote  (also  two  specimens  taken  in  northern  Harris  County,  Texas  originally  identified  as  C.  la- 
trans). See  Figs.  24,  26,  27,  28,  29,  and  30  for  more  details.  Note:  because  of  the  scale  of  the  map  it  was  not 
possible  to  plot  all  localities  in  crowded  areas. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


47 


collected  in  the  areas  under  discussion.  Al- 
though the  distribution  of  the  red  wolf  was 
certainly  not  stable  through  the  entire  period, 
it  seems  safe  to  say  that  about  1940  the  range 
of  unmodified  C.  rufus  extended  from  extreme 
southeastern  Oklahoma  and  southeastern 
Texas,  across  Louisiana,  southern  Arkansas, 
and  southern  Mississippi,  and  into  western 
Alabama  .  Localities  of  the  available  speci- 
mens from  this  region  are  listed  in  appendix 
A  (part  21),  and  are  plotted  in  figure  25. 

Increase  of  Hybridization  from  the 
19.30's  to  1950's 

Specimens  collected  prior  to  1930  indicate 
the  effects  of  hybridization  between  C.  riifus 
and  C.  latrans  only  in  central  Texas,  and  at 
a  few  localities  in  Arkansas  and  eastern  Okla- 
homa. In  the  following  decades,  specimens  of 
apparent  mixed  genetic  origin  were  taken 
over  larger  areas.  Information  from  the  1930's 
and  early  1940's  seems  important  to  a  full 
understanding  of  the  phenomena  then  engulf- 
ing populations  of  southern  Canis,  but,  un- 
fortunately, comparatively  few  specimens 
were  saved  from  the  areas  of  greatest  interest. 

As  explained  earlier,  specimens  taken 
in  1900  in  Calhoun  County,  on  the  middle 
Gulf  Coast  of  Texas,  have  multivariate  posi- 
tions within  or  near  the  limits  of  standard  C. 
rufus.  No  other  specimens  of  wild  Canis  were 
collected  in  this  area  until  after  1930.  Jack- 
son (1951:275)  listed  specimens  of  C.  latrans 
frustror  from  Aransas,  Refugio,  and  Victoria 
counties.  Goldman  (1944:488-489)  stated 
that  specimens  of  C.  rufus  rufus  had  been 
collected  in  the  same  area.  Skulls  of  10  males 
obtained  from  1936  to  1942  in  these  three 
adjoining  counties,  were  suitable  for  analysis 
(see  Fig.  25;  appendix  A,  part  22).  Five  had 
been  referred  originally  to  C.  rufus  and  five 
to  C.  latrans.  As  shown  by  the  positions 
plotted  in  figure  26  these  previous  designa- 
tions are  questionable  and  some  genetic  ex- 
change seems  to  have  occurred. 


RUFUS 


Fig.  26. — Multivariate  positions  of  individuals 
taken  from  1936  to  1942  in  Aransas,  Refugio,  and 
Victoria  counties  on  the  central  coast  of  Texas. 
Squares,  specimens  originally  identified  as  C.  rufus; 
black  dots,  specimens  originally  identified  as  C. 
latrans. 


Few  specimens  from  farther  inland  in 
southern  and  central  Texas  are  available  from 
this  period.  No  skulls  at  all  were  saved  from 
the  central  Texas  counties  that  were  appar- 
ently occupied  by  a  hybrid  swarm  of  C.  rufus 
x  C.  latrans  prior  to  1920  (see  pp.  41-43). 
Probably,  as  is  shown  in  the  next  subsection 
of  the  paper,  all  wild  Canis  had  been  extermi- 
nated in  this  area  by  the  1930's.  Three  males 
of  C.  /.  texensis,  taken  in  1942  in  Bexar  Coun- 
ty, have  multivariate  positions  within  the 
range  of  C.  latrans  (Fig.  27). 

Farther  north,  a  series  of  specimens  was 
taken  from  1930  to  1942  in  the  area  between 
the  Colorado  and  Red  rivers  (see  Fig.  25; 
appendix  A,  part  23).  Records  of  both  C. 
rufus  and  C.  latrans  were  listed  from  various 
localities  in  this  area  by  Goldman  (1944)  and 
Jackson  ( 1951 ) ,  but  I  can  see  no  basis  for 
separating  these  specimens  into  two  species. 
I  subjected  18  of  the  skulls  to  multivariate 
analysis  and  found  most  to  fall  within  or  near 
the  range  of  variation  of  standard  C.  latrans 
(Fig.  27).  The  more  intermediate  positions 
of  two  females  (D2  from  coyote  36.7  and 
22.8;  from  red  wolf  39.3  and  25.6)  suggest 
genetic  influence  from  C.  rufus. 


48 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


LATRANS 


RUFUS 


~i      i      i      i      i      i      i      i      i      i      i      i 

-4-3-2-10       1        2      3      4      5       6      7 


□ 


I 


-| 1 1- 

2      3     4 


Fig.  27. — Multivariate  positions  of  individuals 
taken  in  Bexar  County,  Texas  (t),  and  in  north- 
central  Texas  from  1930  to  1942.  Black  dots,  speci- 
mens originally  identified  as  C.  latrans;  squares, 
specimens  originally  identified  as  C.   rufus. 

From  1933  to  1942  a  large  series  of  skulls 
was  collected  on  the  Wichita  Mountains  Na- 
tional Wildlife  Refuge,  Comanche  County, 
southwestern  Oklahoma  (see  Fig.  25;  appen- 
dix A,  part  24).  Nearly  all  of  these  speci- 
mens were  originally  identified  as  C.  latrans 
frustror,  and  this  designation  is  supported  by 
statistical  positions  of  22  males  and  25  fe- 
males (Fig.  28).  Goldman  (1944:488)  listed 
a  single  specimen  of  a  male  from  this  group 
as  C.  rufus  rufus.  I,  however,  was  not  able 
to  distinguish  this  skull  from  those  of  some 
large  coyotes,  and  its  multivariate  position 
(Fig.  28)  suggests  affinity  with  C.  latrans. 
Therefore,  if  the  red  wolf  ever  did  occur  as 
far  west  in  Oklahoma  as  Comanche  County, 
it  apparently  disappeared  before  the  1930's. 
Nonetheless,  the  presence  of  coyotes,  some  of 


which  had  perhaps  received  an  introgression 
of  genes  from  C.  rufus,  caused  persons  to 
think  that  wolves  still  inhabited  the  area. 
According  to  Halloran  and  Glass  (1959:363): 
"The  Texas  red  wolf  reaches  the  western  edge 
of  its  range  in  the  Wichita  Mountains.  Most 
of  the  canids  present  on  the  refuge  are  coy- 
otes, but  the  situation  is  clouded  by  the  local 
custom  of  calling  everything  larger  than  a 
small  coyote  a  wolf."  And  Duck  and  Fletcher 
(1945:128)  reported  that  C.  rufus  rufus  was 
being  taken  over  most  of  the  state. 

Several  other  specimens,  identified  as  C.  I. 
frustror,  were  taken  in  central  and  north- 
eastern Oklahoma  in   1932.    Those  usable  in 


3 
2 

I 

0 
-I 
-2 
-3 

3 
2 

I 

0 

-I 
-2 
-3  - 


\-  s 
LATRANS 


RUFUS 


~l 1- 

5      6 


-4 


-i 1 r 

-3    -2     -I 


-T- 
0 


3     4 


Fig.  28. — Multivariate  positions  of  individuals 
taken  in  Oklahoma  from  1932  to  1942,  relative  to 
series  of  C.  rufus  and  C.  latrans.  Black  dots,  C. 
latrans  frustror  from  Comanche  County;  square, 
specimen  from  Comanche  County  identified  by  Gold- 
man (1944)  as  C.  rufus,  but  considered  in  this  paper 
to  represent  C.  latrans;  v,  specimens  from  Cleveland 
County;  E,  coyotes  from  eastern  Oklahoma;  triangle, 
specimen  from  Atoka  County  apparently  representing 
C.   r.  rufus. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


49 


multivariate  analysis  include  three  males  from 
Cleveland  County,  one  male  from  Cherokee 
County,  and  two  females  from  Osage  County 
(see  appendix  A,  part  25).  These  six  speci- 
mens demonstrate  statistical  approach  to  C. 
ntfus  (Fig.  28).  A  single  female,  taken  the 
same  year  in  Atoka  County,  southeastern 
Oklahoma,  and  identified  originally  as  C.  r. 
rufus,  seems  to  be  within  the  range  of  varia- 
tion of  other  specimens  assigned  to  that  sub- 
species (Fig.  28).  Two  other  skulls  (in 
USNM)  collected  in  1932  in  eastern  Okla- 
homa are  a  female  from  Cherokee  County 
identified  as  C.  r.  rufus,  and  a  male  from 
LeFlore  County  assigned  by  Goldman  to 
C.  r.  gregoryi.  Neither  one  is  suitable  for 
multivariate  analysis,  but  both  appear  to  be 
intermediate  in  characters  between  the  stand- 
ard samples  of  C.  rufus  and  C.  htrans.  As 
mentioned  in  the  last  subsection  of  this  paper, 
two  skulls  of  red  wolves  were  collected  in 
1936  in  McCurtain  County,  southeastern 
Oklahoma. 

The  distribution  of  specimens  thus  sug- 
gests that  by  the  1930's  the  red  wolf  had 
become  restricted  to  parts  of  southeastern 
Oklahoma.  A  few  skulls  taken  in  the  north- 
eastern and  central  areas  of  the  state,  how- 
ever, may  represent  the  results  of  interbreed- 
ing between  C.  rufus  and  C.  latrans  (see  Fig. 
25). 

Specimens  discussed  earlier  in  the  paper 
demonstrate  the  sympatric  occurrence  of  C. 
rufus  and  C.  latrans  in  southern  Missouri 
prior  to  1930.  Ry  that  year,  however,  the  red 
wolf  had  been  decimated  by  government  and 
private  hunting  (Nowak,  1970;  Sampson, 
1961).  In  contrast,  the  coyote  seems  to  have 
maintained  its  numbers  and  increased  its 
range.  One  possible  indication  of  subsequent 
hybridization  in  the  area  was  provided  by 
Rennitt  and  Nagel  (1937:168)  who  wrote: 
"It  is  difficult  to  outline  exactly  the  range  of 
these  two  species  in  Missouri,  since  so  many 
observers  cannot  tell  them   apart." 


According  to  Sampson  (1961),  the  last 
pure  specimen  of  a  Missouri  red  wolf  was 
collected  in  1932  in  Dade  County.  This  speci- 
men, however,  was  not  listed  by  either  Gold- 
man (1944)  or  Jackson  (1951),  and  its  multi- 
variate position  (Fig.  29)  is  well  within  the 
range  of  variation  of  C.  latrans.  A  specimen 
collected  the  same  year  in  Iron  County  was 
listed  by  Goldman  (1944:486)  as  gregoryi, 
and  was  considered  by  Paradiso  and  Nowak 
(1972a:  11)  possibly  to  be  a  red  wolf.  At 
present,  however,  I  think  the  skull  represents 
hybridization  involving  C.  familiaris  (see  Fig. 
16). 

Sampson  ( 1961 )  reported  that  in  1941 
and  1942  a  series  of  171  specimens  was  taken 
throughout   Missouri,   partly  in  order  to  de- 


3 
2 
I 

0 
-I 

-2 
-31- 


8 

LATRANS 


RUFUS 


□ 


"i r 

-3    -2 


~i 1 1 

5       6      7 


3 
2 

I 

0 

-I 
-2 
-3  - 


9 


-3    -2 


0 


Fig.  29. — Multivariate  positions  of  individuals 
taken  in  Missouri  from  1932  to  1942,  relative  to  series 
of  C.  rufus  and  C.  latrans.  Black  dots,  specimens 
originally  identified  as  C.  latrans;  d,  specimen  from 
Dade  County  listed  as  C.  rufus  by  Sampson  (1961); 
squares,  specimens  from  Ozark  and  Oregon  counties 
identified  as  C.  r.  rufus  by  Leopold  and  Hall  (1945). 


50 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


termine  whether  the  red  wolf  was  still  present 
in  the  state.  He  wrote  that  44  of  the  skulls 
were  sent  to  the  U.S.  National  Museum,  but 
suggested  that  these  included  all  specimens 
thought  most  likely  to  represent  C.  rufus. 
Hence  the  sample  sent  in  probably  already 
was  biased  in  favor  of  larger  and  more  wolf- 
like individuals.  In  any  case,  the  Museum 
reportedly  identified  the  specimens  as  39  coy- 
otes, one  dog,  one  coy-dog,  and  one  red  wolf- 
coyote  hybrid.  Jackson  (1951:274)  referred 
most  of  these  specimens  to  C.  latrans  frustror, 
and  listed  records  all  across  southern  Mis- 
souri. Of  the  specimens  taken  in  1941  and 
1942  in  the  southern  half  of  Missouri,  eight 
males  and  seven  females  were  suitable  for 
multivariate  analysis  (see  Fig.  25;  appendix 
A,  part  26).  Their  relative  positions  are 
shown  in  figure  29,  and  indicate  close  affinity 
with  C.  latrans,  but  possibly  some  genetic 
influence  from  C.  rufus. 

Elder  and  Hayden  (1977)  evaluated  20 
specimens  of  Canis  from  Missouri,  deposited 
in  the  collection  of  the  University  of  Missouri, 
and  reported  one  to  be  C.  familiaris  and  two 
to  be  coy-dogs.  Of  the  others,  four  taken 
between  1945  and  1950  in  the  Ozarks  were 
identified  as  C.  rufus,  nine  taken  mostly  from 
1940  to  1955  were  identified  as  C.  latrans, 
and  four  taken  between  1946  and  1949  were 
considered  to  probably  be  red  wolf-coyote 
hybrids.  The  graphical  results  of  this  analysis 
suggest  to  me  that  these  last  17  specimens 
represent  a  single  interbreeding  population, 
and  not  that  the  red  wolf  survived  as  a  dis- 
tinct entity  in  Missouri  until  1950.  The  fact 
that  Elder  and  Hayden  also  identified  the 
two  skulls  mentioned  in  the  next  sentence  as 
C.  rufus,  suggests  that  their  accepted  limits 
for  this  species  were  less  restrictive  than  my 
own. 

Leopold  and  Hall  (1945)  referred  to  C.  r. 
rufus  two  specimens  of  males,  taken  in  1941 
and  1942,  respectively,  in  Ozark  and  Oregon 
counties,  extreme  southern  Missouri.  In  a 
1952  letter  cited  by  Sampson  (1961),  Leopold 
wrote:    "In  all  respects  these  two  specimens 


are  intermediate  between  typical  coyotes  and 
typical  red  wolf."  Multivariate  analysis  of 
the  skulls  (Fig.  29)  supports  this  statement. 
Leopold  and  Hall  had  assigned  the  specimens 
to  C.  r.  rufus  on  the  basis  of  their  small  size, 
even  though  they  were  obtained  within  the 
range  designated  for  C.  r.  gregoryi  by  Gold- 
man (1944:484-486).  Since  Goldman  had 
assigned  two  male  specimens  from  Reeds 
Spring,  Stone  County,  Missouri  to  gregoryi, 
an  awkward  situation  developed  in  which 
material  of  rufus  was  reported  from  farther 
to  the  east.  Hall  and  Kelson  (1952:340-341) 
technically  solved  this  problem  by  stating 
that  the  two  Stone  County  skulls  resembled 
C.  r.  rufus  in  small  size  and  cranial  charac- 
ters, and  by  referring  them  to  this  subspecies. 
The  issue,  however,  is  a  temporal  as  well  as 
a  geographic  one.  The  Stone  County  material 
was  taken  in  1923  and  1924,  and  as  we  have 
seen,  falls  right  in  the  statistical  center  of  a 
group  of  70  skulls  of  C.  r.  gregoryi  (Fig.  19). 
The  Ozark  and  Oregon  County  specimens 
were  taken  almost  20  years  later  at  a  time 
when  the  red  wolf  had  been  nearly  or  com- 
pletely exterminated  in  Missouri.  They  are 
statistically  beyond  standard  C.  rufus,  and 
should,  I  think,  be  looked  upon  as  represent- 
ing hybridization  between  C.  rufus  and  C. 
latrans,  rather  than  a  particular  subspecies  of 
the  red  wolf. 

A  related  problem  seems  to  exist  just  to 
the  south  in  Arkansas.  Locality  records  from 
this  state  listed  by  Goldman  (1944:485-488) 
include  42  for  C.  rufus  gregoryi  and  6  for 
C.  r.  rufus.  If  the  adjacent  marginal  records 
for  each  subspecies  are  plotted  on  a  map,  as 
they  were  by  Hall  and  Kelson  (1959:852), 
an  awkward  picture  emerges.  Several  of  the 
localities  from  which  rufus  was  reported  are 
seen  to  be  nearly  surrounded  by  the  desig- 
nated range  of  gregoryi.  There  are  no  geo- 
graphic barriers  that  might  account  for  such 
a  distribution,  but  once  again  the  time  ele- 
ment seems  to  be  a  factor.  As  best  as  I  can 
determine,  all  but  one  of  the  specimens  as- 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


51 


signed  by  Goldman  to  gregoryi  were  col- 
lected prior  to  1930.  In  contrast,  there  is  only 
one  locality  from  which  specimens  assigned 
to  rufus  were  taken  before  1930.  This  locality 
is  Boxley,  Newton  County,  which  is  not  one 
of  those  surrounded  by  the  range  of  gregoryi. 
A  skull  taken  in  1922  at  this  site  falls  within 
the  standard  range  of  variation  of  C.  rufus 
(Fig.  19).  The  localities  that  are  encircled 
by  the  range  of  gregoryi  are  Raspberry  and 
Hector,  both  in  Pope  County.  Of  the  four 
specimens  recorded  by  Goldman  from  these 
places,  two  are  not  usable  in  my  analyses,  one 
has  been  identified  by  me  as  a  dog  hybrid 
(Fig.  16),  and  the  statistical  position  of  one 
is  shown  in  figure  30.  Of  the  other  post- 1930 
specimens  listed  by  Goldman,  only  one  could 
be  subjected  to  analysis,  and  its  position  is 
also  depicted  in  figure  30.  The  specimens 
evaluated    in    the    analysis,    as    well    as    the 


RUFUS 


Fig.  30. — Multivariate  positions  of  individuals 
taken  in  Arkansas  from  1932  to  1951,  relative  to 
series  of  C.  rufus  and  C.  latrans.  Black  dots,  speci- 
mens originally  identified  as  C.  latrans;  squares, 
specimens  originally  identified  as  C.  rufus. 


others,  suggest  a  more  pronounced  influence 
by  C.   latrans  after  1930. 

Jackson  (1951:274)  recorded  C.  latrans 
frustror  from  a  number  of  localities  in  western 
and  northern  Arkansas.  Most  of  the  speci- 
mens involved  were  collected  between  1932 
and  1942,  but  not  all  are  suitable  for  analysis. 
The  positions  of  those  that  were  statistically 
evaluated  are  plotted  in  figure  30.  A  few 
other  skulls,  taken  as  late  as  1951  in  the  same 
areas,  are  also  depicted.  Arkansas  specimens 
taken  in  this  period  are  listed  in  appendix  A 
(part  27),  and  localities  are  shown  in  figure 
25.  The  number  of  specimens  is  hardly 
enough  to  allow  a  full  understanding  of  the 
situation,  but  their  distribution  suggests  that 
after  1930  C.  rufus  was  no  longer  prevalent 
in  Arkansas,  and  that  animals  of  more  inter- 
mediate or  coyotelike  characters  were  begin- 
ning to  predominate. 

The  Empty  Zone  and  Expansion  of 
Modified  Canis 

The  maps  in  figures  14  and  25  show  many 
areas  in  which  no  records  of  Canis  are  plot- 
ted. Of  course,  material  from  states  west  of 
Texas,  and  north  of  Oklahoma  and  Missouri, 
was  available,  but  was  not  considered  appli- 
cable to  the  problems  under  discussion.  The 
records  that  are  shown  seem  to  form  a  rough- 
ly circular  pattern  around  a  blank  area  cen- 
tered in  northeastern  Texas.  It  could  be 
argued  that  this  area  is  not  represented  only 
because  no  one  ever  collected  there,  or  be- 
cause no  material  from  there  was  preserved. 
But  it  should  be  understood  that  almost  all 
specimens  discussed  in  this  section  of  the 
paper  were  taken  in  the  course  of  Federal 
predator  control  efforts.  This  control  work 
was  done  in  areas  where  complaints  existed, 
and  thus  is  indicative  of  the  presence  and 
abundance  of  wild  Canis.  To  be  sure,  only 
a  small  percentage  of  the  animals  killed  were 
sent  as  specimens  to  the  National  Museum, 
and  collecting  was  not  continued  on  a  regular 
geographic  or  temporal  basis.  Nonetheless, 
the  available  material  may  serve  as  one  relia- 


52 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


ble  means  of  evaluating  the  situation  over  a 
lengthy  period.  Therefore,  the  constant  lack 
of  specimens  from  the  area  in  question  leads 
me  to  think  that  no  significant  population  of 
wild  Canis  had  survived  there  into  the  twen- 
tieth century. 

Bailey  (1905:172)  stated:  "The  black  wolf 
is  reported  from  a  few  localities  in  the  tim- 
bered region  of  eastern  Texas,  but  in  most 
cases  as  'common  years  ago,  now  very  rare 
or  extinct.' "  But  in  another  publication,  per- 
haps because  it  was  written  for  different 
purposes,  Bailey  (1907:13)  stated:  "In  the 
timbered  region  of  eastern  Texas,  especially 
in  the  extensive  swamps  and  bottoms,  the 
black  wolf  is  still  abundant  and  very  de- 
structive to  cattle  and  hogs,  while  it  renders 
sheep  raising  practically  impossible."  Bailey's 
own  field  reports  from  1904  (on  file,  National 
Fish  and  Wildlife  Laboratory,  U.S.  National 
Museum  of  Natural  History)  do  indicate  that 
in  the  area  of  the  Big  Thicket  in  Hardin  and 
Liberty  counties,  wolves  were  still  common. 
Strecker  (1926:16)  reported  wolves  in  ex- 
treme eastern  Texas  to  be  "now  almost  or 
quite  extinct."  Apparently  then,  by  the  early 
twentieth  century  wolves  had  been  wiped  out 
in  most  of  the  higher  country  of  eastern  Texas, 
but  continued  to  live  in  the  bottom  land 
swamps,  especially  in  the  area  of  the  Big 
Thicket. 

To  the  north  of  the  Big  Thicket,  it  seems 
as  though  there  was  actually  a  zone  nearly  or 
completely  empty  of  wild  Canis.  This  idea 
is  supported  by  the  existence  of  certain  maps 
showing  the  distribution  of  Canis  in  Texas  at 
various  times.  One  of  these,  prepared  by 
Bailey  (1907:7),  depicts  a  large  area  of  north- 
central  Texas  as  being  free  of  the  presence  of 
wolves.  The  status  of  coyotes  in  the  area, 
however,  is  not  indicated. 

In  the  annual  report  for  fiscal  1931  of  the 
Texas  District  of  the  U.S.  Bureau  of  Biologi- 
cal Survey  (on  file,  offices  of  U.S.  Fish  and 
Wildlife  Service,  San  Antonio),  there  is  a 
map  showing  the  "Present  Infestation  of 
Predatory    Animals."     The    term    "predatory 


animals"  is  vague,  but  since  coyotes  and 
wolves  were  usually  the  main  target  of  Fed- 
eral control  programs,  especially  in  Texas, 
it  is  safe  to  assume  that  canids  are  being  re- 
ferred to.  The  map  indicates  that  predators 
were  absent  in  the  eastern  part  of  Texas, 
except  along  the  coast,  in  the  vicinity  of  the 
Big  Thicket,  and  immediately  south  of  the 
Red  River.  In  a  large  area  of  central  Texas, 
including  all  of  those  counties  thought  occu- 
pied by  a  hybrid  swarm  of  Canis  prior  to 
1920,  the  map  shows  predatory  animals  to  be 
"under  control"  in  1931.  This  term  may  be 
taken  to  mean  nearly,  if  not  completely,  ex- 
terminated. There  is  evidence  that  all  wild 
Canis  had  been  decimated  in  central  Texas 
by  this  time.  According  to  Gabrielson  (1936: 
64),  sheep  raisers  in  this  area  sustained  losses 
of  ten  percent  to  predators  in  1915,  but,  be- 
cause of  Federal  control  programs,  had  losses 
of  only  a  fraction  of  one  percent  in  1935. 
Several  of  the  reports  cited  by  Russell  and 
Shaw  (1971a)  indicated  that  by  the  late 
1930's  coyotes  and  wolves  had  been  com- 
pletely exterminated  in  some  central  Texas 
counties. 

Another  map,  drawn  by  Russell  and  Shaw 
(1971a),  was  based  on  reports  made  in  1940 
by  field  personnel  of  the  Texas  Game,  Fish 
and  Oyster  Commission.  The  range  of  the 
red  wolf  at  this  time  was  shown  to  extend  all 
along  the  Gulf  Coast  north  of  Nueces  Bay, 
to  extend  inland  in  southeastern  Texas  up  the 
Trinity  and  Neches  river  basins,  and  to  in- 
clude strips  along  the  Red,  Sabine,  and  Sul- 
phur rivers  in  extreme  northeastern  Texas. 
Most  of  the  reports  cited  from  northeastern 
and  inland  Texas,  however,  state  that  wolves 
were  very  rare  in  these  areas.  The  only  major 
area  of  abundance  indicated  by  the  reports 
cited  was  the  southeastern  corner  of  the  state. 
In  this  regard  there  is  agreement  with  the 
1931  Biological  Survey  map  and  the  available 
specimens  (see  pp.  45-46). 

Although  Russell  and  Shaw  dealt  pri- 
marily with  C.  rufas,  reports  by  Texas  game 
officials  regarding  the  status  of  coyotes  were 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


53 


also  made  between  1940  and  1942  (on  file, 
offices  of  Texas  Parks  and  Wildlife  Depart- 
ment, Austin).  These  reports  indicate  the 
presence  of  coyotes  only  in  those  areas  south 
of  San  Antonio,  west  of  the  Pecos  River,  and 
northwest  of  Fort  Worth. 

The  sum  of  evidence  from  three  different 
sources  (specimens.  Federal  reports,  and 
Texas  government  reports)  suggests  that  the 
following  situation  existed  in  Texas  in  the 
1930's  and  early  1940's.  Red  wolves  were 
present  in  moderate  numbers  in  the  south- 
eastern corner  of  the  state  and  along  the 
central  Gulf  Coast.  In  the  latter  area  hybridi- 
zation with  coyotes  was  probably  occurring. 
Coyotes  themselves  were  common  in  the 
southern,  western,  and  northern  parts  of 
Texas.  In  a  large  area  centered  in  north- 
eastern Texas,  however,  all  kinds  of  wild 
Canis  seem  to  have  been  rare,  if  not  totally 
absent.  I  think  that  this  empty  zone  may 
have  been  significant  for  two  reasons :  ( 1 ) 
it  served  as  an  actual  barrier  that  for  a  time 
limited  contact,  and  probably  interbreeding, 
between  C.  latrans  to  the  north  and  west,  and 
C.  rufus  in  southeast  Texas,  and  hence  may 
have  assisted  in  maintaining  an  unmodified 
red  wolf  population  in  the  latter  area;  and 
(2)  it  formed  an  empty  niche  for  predators 
that  awaited  reoccupation  by  suitable  canids. 

This  void  apparently  began  to  fill  by  the 
1940s.  As  discussed  in  the  previous  subsec- 
tion of  this  paper,  coyotes  and  animals  inter- 
mediate to  C.  latrans  and  C.  rufus  seemed  by 
that  time  to  be  increasing  in  Oklahoma, 
southern  Missouri,  and  Arkansas.  In  this 
period  there  are  several  references  to  the 
presence  of  wild  Canis  in  Arkansas.  Black 
( 1936 )  wrote  that  the  "timber  wolf"  was  rare 
in  the  northwestern  part  of  the  state,  and  he 
did  not  mention  the  presence  of  coyotes. 
Dellinger  and  Black  (1940)  listed  several 
records  of  coyotes,  and  also  remarked: 
'Wolves  are  becoming  rather  common  in  the 
Ozarks."  The  Arkansas  Game  and  Fish  Com- 
mission (1951:96-99)  reported  that  coyotes 
had  been  extending  their  range  and  could  be 


found  as  far  east  as  the  central  part  of  the 
state.  Wolves,  which  also  had  reportedly  in- 
creased, were  said  to  be  most  common  in  the 
Ozark  area,  and  second  most  common  in 
southwestern  Arkansas.  But  in  this  last  named 
account,  wolves  and  coyotes  were  discussed 
together  and  perhaps  were  being  confused. 
Furthermore,  it  was  stated  that  the  Missis- 
sippi Valley  wolf,  C.  r.  gregoryi,  had  become 
rare  and  should  not  be  totally  exterminated. 
This  implies  recognition  that  the  canids  re- 
ported to  be  common  were  actually  some- 
thing other  than  the  original  native  wolves 
of  most  of  Arkansas.  In  contrast  to  this  view- 
point, Sealander  (1956:279)  reported:  "The 
race  gregoryi  occurs  throughout  the  State  and 
is  quite  numerous  in  some  counties.  It  evi- 
dently has  largely  replaced  the  race  rufus 
over  its  former  range  in  Arkansas."  Sea- 
lander,  however,  informed  me  (pers.  comm.) 
that  this  statement  had  been  based  partly  on 
Goldman's  (1944:487)  report  that  rufus  had 
become  restricted  to  parts  of  central  and 
southern  Texas.  Since  rufus  was  thus  not  sup- 
posed to  exist  in  Arkansas,  but  since  canids 
thought  to  be  red  wolves  were  certainly  pres- 
ent, Sealander  had  to  refer  the  latter  to 
gregoryi. 

These  references  suggest  an  increase  in 
wild  Canis  in  Arkansas  between  the  1930's 
and  1950's,  with  the  animals  involved  be- 
lieved to  be  wolves.  None  of  these  reports, 
however,  was  based  on  a  thorough  examina- 
tion of  specimens.  The  few  specimens  that 
are  available  from  this  period  suggest  that 
the  original  wolf  population  of  Arkansas  was 
being  replaced  by  more  coyotelike  animals 
(Fig.  30).  Later,  Paradiso  (1966)  reported 
that  three  specimens  taken  in  1964  in  Chicot 
County,  extreme  southeastern  Arkansas,  rep- 
resented a  range  extension  of  C.  latrans  frus- 
tror.  The  first  study  of  large  series  of  Arkan- 
sas specimens  collected  since  1930  was 
described  by  Gipson  (1972),  and  Gipson, 
Sealander,  and  Dunn  (1974).  On  the  basis 
of  multivariate  analysis  of  284  adult  skulls 
taken  from  1968  to  1971,  they  concluded  that 


54 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


the  existing  population  of  Cams  in  the  state 
was  predominantly  coyote,  though  with  some 
red  wolf  influence.  The  essentially  coyotelike 
character  of  this  population  also  is  recognized 
in  the  next  subsection  of  my  paper.  There- 
fore, the  canids  that  were  said  to  be  common 
in  Arkansas  through  the  1950's  could  not  have 
been  red  wolves. 

While  true  red  wolves  were  being  deci- 
mated in  the  Ozark-Ouachita  uplands  in  the 
1920's,  coyotes  probably  moved  in  and  in- 
creased in  numbers,  partially  filling  the  va- 
cant ecological  niche.  Surviving  red  wolves 
probably  interbred  with  coyotes  and  pro- 
duced some  of  the  intermediate  specimens 
discussed  in  the  previous  subsection.  These 
hybrids  served  as  a  bridge  for  the  introgres- 
sion  of  genes  from  C.  rufus  into  C.  latrans. 
Much  of  the  coyote  population  was  thus 
modified,  probably  in  a  manner  that  favored 
its  continued  expansion  into  a  woodland  habi- 
tat. Of  course,  lumbering  and  agricultural 
practices  may  also  have  assisted  the  spread 
of  C.  latrans  into  the  south-central  states 
(McCarley,  1962;  Gipson,  1972). 

The  increase  of  coyotes  in  Arkansas,  and 
undoubtedly  also  in  Oklahoma,  was  followed 
by  their  large-scale  build  up  in  the  previously 
empty  zone  centered  in  northeastern  Texas. 
Halloran  (1959,  1960)  provided  records  that 
almost  completely  block  in  northeastern 
Texas.  Although  he  thought  that  these  rec- 
ords depicted  the  occurrence  of  black-colored 
red  wolves,  the  animals  in  question  were  cer- 
tainly members  of  the  expanding  population 
of  modified  coyotes. 

As  Halloran  explained,  the  existence  of 
black  animals  was  thought  to  indicate  the 
presence  of  red  wolves,  rather  than  coyotes. 
Whereas  C.  rufus  had  a  locally  common  black 
phase,  only  one  record  of  a  black  coyote  was 
known  to  Young  (1951:52).  Halloran  (1958) 
also  used  records  of  black  canids  in  an  effort 
to  plot  the  distribution  of  C.  rufus  in  Okla- 
homa. In  recent  years,  however,  there  has 
been  widespread  recognition  of  the  existence 
of  black  coyotes  in  the  south-central  United 


States.  Halloran  himself  (1963)  reported  one 
from  Comanche  County,  Oklahoma.  Dal- 
quest  ( 1968 )  wrote  that  a  specimen  of  C. 
latrans  tcxensis  from  Wilbarger  County, 
Texas  was  in  the  black  color  phase.  Pimlott 
and  Joslin  (1968)  attempted  to  locate  black 
animals  in  Arkansas,  hoping  that  these  would 
be  C.  rufus,  but  the  four  that  they  found 
were  coyotes  of  medium  size.  Of  the  284 
Arkansas  skulls  analyzed  by  Gipson  (1976), 
24  were  from  black  animals.  Of  these,  12 
were  identified  as  coyotes,  six  as  coyote-dog 
hybrids,  five  as  coyote-red  wolf  hybrids,  and 
one  as  a  domestic  dog.  Freeman  (1976:14) 
reported  that  12  of  121  Oklahoma  specimens 
were  black  or  very  dark,  and  that  the  skulls 
of  eight  of  these  were  identified  as  coyotes, 
two  as  coyote-dog  hybrids,  and  two  as  coyote- 
red  wolf  hybrids.  Elder  and  Hayden  (1977) 
stated  that  of  the  seven  Missouri  specimens 
they  considered  to  be  red  wolves  or  coyote- 
red  wolf  hybrids,  five  were  black  or  had  been 
associated  with  black  animals. 

All  recent  records  of  black  coyotes  have 
come  from  the  former  range  of  C.  rufus  (ex- 
cept for  one  reported  in  Michigan  by  Ozoga 
and  Harger,  1966).  Although  it  could  be 
argued  that  this  phenomenon  is  the  result  of 
preservation  of  a  newly  favorable  mutation, 
I  consider  it  further  evidence  of  the  recent 
introgrcssion  of  genes  from  C.  rufus  into 
C.  latrans. 

The  coyotelike  population  apparently  in- 
creased rapidly  in  the  1950's.  McCarley 
( 1959 )  reported  C.  latrans  to  be  common  in 
most  areas  of  east  Texas.  He  said  also  that 
the  red  wolf  was  either  extirpated  or  ex- 
tremely rare  in  the  area.  McCarley  ( 1962 ) 
reported  data  on  110  skulls  collected  since 
1948  in  east  Texas,  eastern  and  central  Okla- 
homa, and  Arkansas.  Observing  that  over-all 
skull  size,  as  represented  by  greatest  length 
and  zygomatic  width,  was  the  only  consistent 
character  separating  red  wolves  and  coyotes, 
he  referred  all  but  one  of  the  specimens  to 
C.  latrans  frustror  (the  exception  was  a  skull 
from  an  unknown  Arkansas  locality,  assigned 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


55 


Fig.  31. — Map  showing  localities  of  specimens  taken  from  1961  to  1971  in  the  south-central  United 
States.  Triangles,  C.  rufus;  squares  and  open  circles,  specimens  apparently  representing  hybridization  between 
C.  rufus  and  C.  latrans  (see  Figs.  37-39);  black  dots,  specimens  of  the  modified  population  of  C.  latrans  (see 
Figs.  32-36).  Note:  because  of  the  scale  of  the  map,  it  was  not  possible  to  plot  all  localities  in  crowded 
areas,  especially  along  the  Texas  coast. 


to  C.  r.  rufus).  McCarley  concluded  that  coy- 
otes had  replaced  red  wolves  in  Oklahoma, 
Arkansas,  and  east  Texas,  and  that  hybridiza- 
tion between  the  two  may  have  occurred. 
He  thought  that  C.  rufus  was  extant  only  in 
a  few  isolated  parts  of  eastern  and  southern 
Louisiana,  on  the  basis  of  two  skulls  obtained 
there  in  1956  and  1957  (see  p.  45). 

Evidently  red  wolves  were  present  over 
much  of  Louisiana  through  the  1940's  (Low- 
ery,  1943;  Nowak,  1967;  St.  Amant,  1959). 
By  the  early  1950's,  however,  their  numbers 
had  been  greatly  reduced  by  government 
trapping  and  private  hunting.  Their  last 
major  concentrations  were  in  the  bottom  lands 


along  the  Mississippi  River  in  the  eastern 
part  of  the  state,  and  in  the  southern  coastal 
marshes  and  prairies.  For  a  brief  period,  the 
northwestern  and  north-central  parts  of  Lou- 
isiana were  apparently  left  without  a  sig- 
nificant population  of  wild  Canis,  and,  in 
effect,  these  areas  became  an  extension  of  the 
"empty  zone"  in  adjacent  northeastern  Texas. 
But  just  after  the  heaviest  period  of  wolf 
trapping,  from  1947  to  1952,  coyotelike  ani- 
mals were  reported  to  be  moving  into  the 
northwestern  part  of  Louisiana.  These  canids 
subsequently  spread  over  most  of  northern 
and  central  Louisiana,  and  continued  to  in- 


56 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


crease  in  numbers  each  year  (Nowak,  1967, 
1970;  Paradiso,  1966;  Wilson,  1967). 

McCarley's  (1959,  1962)  reports  were  the 
first  published  indications  that  the  species 
C.  nifus  might  be  in  serious  trouble  through- 
out its  entire  range.  Until  then,  there  had 
been  a  general  belief  that  red  wolves  were 
common  over  large  areas.  Actually,  Goldman 
(1937)  had  written  that  both  the  subspecies 
floridanus  and  rufiis  might  be  extinct,  but 
subsequently  ( 1944 )  he  reported  the  latter  to 
be  present  in  parts  of  central  and  southern 
Texas.  Young  (1946:43)  thought  that  red 
wolves  could  survive  indefinitely  in  large 
areas  of  Texas  and  Louisiana  where  they 
would  be  in  no  direct  conflict  with  man. 
Later  Federal  reports  told  of  large  and  even 
increasing  numbers  of  red  wolves  in  Texas, 
Arkansas,  and  Oklahoma.  Obviously  such  re- 
ports were  based  on  the  movement  of  coyotes 
into  areas  formerly  occupied  by  C.  rufus,  but 
as  late  as  1964  some  authorities  considered 
the  species  to  be  in  no  danger  (see  Nowak, 
1967,  1970,  1972).  By  that  year,  however, 
the  precarious  status  of  the  red  wolf  was 
generally  recognized,  and  efforts  were  under- 
way to  evaluate  the  threats  both  from  man 
and  from  interbreeding. 

Examination  of  Post- 1960  Material  from 

Oklahoma,  Arkansas,  Louisiana,  and 

Inland  East  Texas 

The  disappearance  of  C.  rufus  from  most 
of  its  former  range,  and  its  partial  replace- 
ment by  other  canids,  motivated  the  collect- 
ing of  many  new  specimens.  Nearly  all  of 
this  material  was  taken  in  the  normal  course 
of  predator  control  work  by  Federal  and  state 
employees.  For  convenience,  the  following 
discussion  is  arranged  on  a  partly  arbitrary 
geographic  basis.  Localities  of  all  specimens 
discussed  in  this  subsection  (that  were  ex- 
amined by  me)  are  plotted  on  the  map  in 
figure  31,  and  listed  in  appendix  A  (parts 
28-32). 

Oklahoma. — Skulls  of  12(4)  males  and 
13(3)  females,  taken  in  1965  in  southeastern 


Oklahoma,  all  fall  within  or  near  the  statisti- 
cal limits  of  standard  C.  latrans  (Fig.  32). 
These  specimens  were  obtained  in  the  same 
area  from  which  a  large  series  of  C.  rufus 
had  been  taken  prior  to  1930.  After  a  multi- 
variate analysis  of  138  skulls  taken  in  1975 
and  1976,  and  114  collected  prior  to  1975, 
mostly  from  1953  to  1970,  Freeman  (1976: 
13-14,  28,  33,  47-48)  concluded  that  the  cur- 
rent Oklahoma  population  of  wild  Canis  was 
essentially  coyotelike.  Of  his  total  of  252 
specimens,  203  were  identified  as  C.  latrans, 
one  as  C.  familiaris,  33  as  coy-dogs,  and  15 
as  intermediate  to  C.  latrans  and  C.  rufus. 
There  was  no  significant  difference  between 
the  older  and  newer  groups,  except  in  the 
southeastern  part  of  the  state  where  skulls 
taken  in  1975  and  1976  were  found  to  be 
smaller.  Freeman  suggested,  and  I  agree, 
that  the  genetic  influence  of  the  red  wolf  in 
eastern  Oklahoma  had  declined  to  the  point 


I-  8 

LATRANS 


RUFUS 


-3  - 


-1 1 1 1~- 

2      3      4       5 


-4     -3    -2 


0 


Fie.  32. — Multivariate  positions  (black  dots)  of 
individual  post- 1960  specimens  from  southeastern 
Oklahoma,  relative  to  the  ranges  of  variation  of  the 
series  of  C.   rufus  and  C.  latrans  shown  in  Fig.   17. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


57 


at  which  only  an  occasional  individual   ex- 
hibited any  characters  of  C.  rufus. 

Northern  Arkansas. — Statistical  positions 
of  20  males  taken  in  1969  and  1970  in  the 
Ozark  area  of  northern  Arkansas  (Conway, 
Franklin,  Newton,  Pope,  and  Van  Buren 
counties ) ,  also  were  found  to  lie  mainly  with- 
in the  range  of  variation  of  C.  latrans  (Fig. 
33).  Two  specimens,  however,  have  D2  val- 
ues slightly  closer  to  C.  rufus,  thus  indicating 
the  continued  genetic  influence  of  this  spe- 
cies. No  female  specimens  from  northern 
Arkansas  were  tested  by  multivariate  analysis, 
but  20  skulls  from  the  area  were  examined, 
and  most  were  indistinguishable  from  those 
of  western  coyotes.  None  exceeded  200  milli- 
meters in  greatest  length  or  100  millimeters 
in  zygomatic  width.  Gipson,  Sealander,  and 
Dunn  (1974)  found  small  "pockets  of  red 
wolf  influence"  in  the  Ozark  and  Ouachita 
mountains,  but  did  not  refer  any  particular 
specimens  from  those  areas  to  C.  rufus.   Pirn- 


il-  8 

LATRANS 


RUFUS 


"l 1 1 1 r~ 

2      3      4       5       6 


Fig.  33. — Multivariate  positions  (black  dots)  of 
individual  post-1960  specimens  from  northern  Ar- 
kansas, relative  to  series  of  C.   rufus  and  C.   latrans. 


8 

LATRANS 


RUFUS 


~i 1 r 

2      3     4 


Fig.  34. — Multivariate  positions  (black  dots)  of 
individual  post-1960  specimens  from  southern  Ar- 
kansas, relative  to  series  of  C.  rufus  and  C.  latrans. 

lott  and  Joslin  (1968)  heard  what  they  con- 
sidered probably  to  be  wolves  in  the  Ozark 
National  Forest  in  1964,  but  found  only  coy- 
otes there  in  1965.  Subsequently,  a  group  of 
investigators  from  Arkansas  Polytechnic  Col- 
lege continued  attempts  to  locate  red  wolves 
in  the  Ozarks,  both  through  examination  of 
skulls  and  elicitation  of  howling  responses 
(see  Nowak,  1970).  Although  some  wolf  like 
calls  were  heard,  no  conclusive  evidence  of 
the  presence  of  C.  rufus  was  obtained  (Henri 
D.  Crawley,  pers.  comm. ).  There  is  little 
doubt  that  true  red  wolves  have  now  disap- 
peared throughout  the  Ozark-Ouachita  up- 
lands. 

Southern  Arkansas. — The  multivariate  po- 
sitions of  skulls  of  52(6)  males  and  11(3) 
females,  taken  from  1964  to  1970  in  southern 
Arkansas,  are  depicted  in  figure  34.  Most  of 
these  fall  within  or  near  the  range  of  varia- 
tion of  C.  latrans,  but  there  is  a  pronounced 
over-all  shift  toward  the  standard  red  wolf 


58 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


sample.  Four  individuals  have  D2  values 
nearer  to  C.  rufus  than  to  C.  latrans.  An  ad- 
ditional 20  skulls  of  females  from  the  area, 
examined,  but  not  tested  by  multivariate  anal- 
ysis, appear  to  follow  the  same  pattern.  Gip- 
son,  Sealander,  and  Dunn  (1974)  reported 
that  two  specimens  from  southern  Arkansas 
were  not  significantly  different  from  C.  rufus, 
and  that  there  was  a  strong  genetic  influence 
from  the  red  wolf  in  the  area.  They  added, 
however,  that  the  red  wolf,  in  pure  form, 
probably  no  longer  existed  in  Arkansas,  and 
that  its  genes  had  been  incorporated  into  a 
predominantly  coyote  population.  Of  the  284 
skulls  they  subjected  to  multivariate  analysis, 
only  these  two  were  classified  as  red  wolves, 
208  were  identified  as  coyotes,  27  were  said 
to  be  intermediate  to  C.  rufus  and  C.  latrans, 
and  the  remainder  were  considered  to  repre- 
sent wild  dogs  or  dog  hybrids.  The  fact  that 
Arkansas  Canis  is  more  wolflike  in  the  south- 
ern part  of  the  state  may  reflect  longer  sur- 
vival of  C.  rufus  there,  and  hence  more  recent 
introgression  from  individuals  of  that  species. 
Gipson  (1972:50-51)  suggested  that  a  "zone 
of  red  wolf  influence"  in  the  south-central 
part  of  the  state  might  have  resulted  from  the 
establishment  of  an  unofficial  refuge  in  that 
area,  in  which  red  wolves  were  released  in 
the  late  1950's. 

Louisiana.— Skulls  of  22(17)  males  and 
19(17)  females,  taken  from  1963  to  1969  in 
Louisiana,  demonstrate  similar  statistical  dis- 
tributions to  those  formed  by  the  southern 
Arkansas  material  (Fig.  35).  Goertz,  Fitz- 
gerald, and  Nowak  (1975)  concluded  that 
155  skulls  collected  in  Louisiana  from  1963 
to  1973  represented  an  essentially  coyotelike 
population  that  had  been  slightly  modified 
through  introduction  of  genes  from  C.  rufus. 
Elements  of  this  same  population  apparently 
now  have  spread  across  Mississippi  and  into 
Alabama  (Cahalane,  1964;  Paradiso,  1966; 
Linzey,  1971;  Wolfe,  1972).  Recent  informa- 
tion indicates  that  coyotelike  animals  are  now 
also  established  in  southwestern  and  south- 
central  Tennessee  ( D.  W.  Yambert,  Tennessee 


V  8 

LATRANS  • 


RUFUS 


0 

-I 
-2 


-3  - 


$ 


-4     -3    -2 


0 


3     4 


n r- 

5     6 


Fig.  35. — Multivariate  positions  (black  dots)  of 
individual  post-1960  specimens  from  Louisiana,  rela- 
tive to  series  of  C.  rufus  and  C.  latrans.  The  triangle 
shows  the  position  of  a  specimen  of  C.  rufus  gregoryi 
collected  in  1964  in  St.  Landry  Parish. 

Wildlife  Resources  Agency,  pers.  comm.), 
and  in  southwestern  Georgia  (Daniel  W. 
Speake,  Cooperative  Wildlife  Research  Unit, 
Auburn  University,  pers.  comm.).  Red  wolves 
seem  to  have  been  extirpated  throughout  this 
region,  except  possibly  in  southern  Louisiana. 
In  1964  Pimlott  and  Joslin  (1968)  heard 
small  groups  of  wolves  in  the  northeastern 
part  of  the  state  and  in  adjoining  sections  of 
Mississippi,  but  there  have  been  no  subse- 
quent records  from  that  area.  Persistent  re- 
ports of  red  wolves  have  come  from  the 
coastal  marshes  in  Cameron  and  Vermilion 
parishes,  but  I  have  examined  no  specimens 
from  this  area.  In  1976,  personnel  of  the 
U.S.  Fish  and  Wildlife  Service  carried  out 
field  studies  in  Cameron  Parish  at  the  ex- 
treme southwestern  corner  of  Louisiana,  and 
live-captured  several  animals  that  appeared 
to    be    red    wolves.     Coyotelike    individuals, 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


59 


however,  also  were  reported  from  this  area. 
A  skull,  obtained  in  1964  near  Washington, 
St.  Landry  Parish,  south-central  Louisiana, 
appears  distinct  from  other  recent  material 
taken  in  the  state.  According  to  Pimlott  and 
Joslin  (1968:383),  this  specimen  was  identi- 
fied as  C.  rufus  gregoryi  by  Barbara  Law- 
rence. Although  the  skull  has  several  dental 
anomalies,  its  multivariate  position  (Fig.  35) 
supports  this  identification.  Red  wolves  have 
received  legal  protection  in  Louisiana  since 
July  1970  (Nowak,  1971). 

Inland  East  Texas. — Figure  36  shows  the 
statistical  positions  of  77(6)  males  and  42(6) 
females,  collected  from  1964  to  1971  in  coun- 
ties of  east  Texas  more  than  100  miles  inland. 
Paradiso  ( 1968 )  examined  most  of  this  ma- 
terial earlier,  and  reported  that  it  represented 
an  interbreeding  population  bridging  the  size 
gap  between  red  wolves  and  coyotes.  Para- 
diso, however,  had  statistically  combined  the 
inland  material  with  a  number  of  specimens 


-3  - 


s 

LATRANS 


RUFUS 


-3    -2 


0 


3     4 


Fig.  36. — Multivariate  positions  (black  dots)  of 
individual  post-1960  specimens  from  inland  east 
Texas,   relative  to  series  of  C.   rufus  and  C.   latrans. 


taken  along  the  Gulf  Coast.  Later,  Paradiso 
and  Nowak  ( 1972a ) ,  also  having  combined 
inland  and  some  coastal  samples,  came  to 
about  the  same  conclusion.  They  thought 
that  the  hybrid  swarm,  established  at  the  turn 
of  the  century  in  central  Texas,  had  expanded 
eastward,  and  by  the  1960's  had  engulfed 
most  of  east  Texas  and  adjacent  parts  of 
Louisiana  and  Arkansas.  This  hypothesis  no 
longer  seems  fully  tenable.  It  was  pointed 
out  above  (pp.  52-53)  that  wild  Canis  in 
central  Texas  had  been  nearly  extirpated  by 
the  1930's,  a  period  prior  to  the  build  up  of 
Canis  in  east  Texas.  Thus  the  population  of 
the  former  area  could  hardly  have  been  the 
source  for  that  of  the  latter.  Furthermore, 
when  intensively  analyzed,  recent  east  Texas 
material  does  not  show  exactly  the  same  kind 
of  statistical  distribution  as  that  of  the  earlier 
specimens.  The  present  population  is  pre- 
dominantly coyotelike,  and  does  not  form  an 
even  distribution  bridging  the  gap  between 
C.  rufus  and  C.  latrans  (compare  Figs.  23 
and  36).  Two  specimens  do  fall  within  the 
range  of  variation  of  standard  C.  rufus,  a  male 
from  Lamar  County  and  a  female  from  Ham- 
ilton County.  There  is  no  apparent  correla- 
tion between  the  geographical  and  statistical 
distributions  of  the  inland  east  Texas  material. 
More  wolflike  and  more  coyotelike  specimens 
occur  throughout  the  area,  sometimes  to- 
gether at  the  same  locality  (see  also  appendix 
B,  part  6). 

The  large  samples  from  inland  east  Texas, 
Louisiana,  and  southern  Arkansas  all  have 
similar  multivariate  distributions,  and  appar- 
ently represent  a  single  population  with  a 
common  origin.  The  region  occupied  by  this 
population  corresponds  in  large  part  to  the 
zone  in  which  Canis  was  rare  or  absent  over 
much  of  the  first  half  of  the  century.  Its  geo- 
graphical distribution  also  lies  mainly  along 
the  southern  fringe  of  the  area  in  which  speci- 
mens of  hybrid  characters  appeared  in  the 
1930's  and  1940's.  Such  hybridization  led  to 
the  introgression  of  C.  latrans,  and  probably 


60 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


also  to  the  genetic  swamping  of  any  red 
wolves  in  the  area  that  had  survived  man's 
onslaught.  After  having  been  modified 
through  introgression  from  C.  rufus,  the  coy- 
ote population  expanded  southward  and 
eastward  across  the  south-central  states.  Ad- 
ditional hybridization  probably  occurred  as 
remnant  pockets  of  red  wolves  were  encoun- 
tered. I  do  not  think,  however,  that  the  cur- 
rent population  of  wild  Canis  in  this  region 
should  be  spoken  of  as  a  "hybrid  swarm." 
The  specimens  demonstrate  a  much  greater 
affinity  with  C.  latrans  than  with  C.  rufus  or 
any  other  species,  and  may  be  properly  re- 
ferred to  as  coyotes.  The  subspecies  consid- 
ered by  Jackson  (1951:271)  to  inhabit  the 
most  proximal  geographic  area  was  C.  latrans 
frustror.  His  description  of  this  subspecies 
as  the  "largest  coyote"  was  based  in  part  on 
the  examination  of  specimens  that  probably 
represented  introgression  from  C.  rufus. 

Examination  of  Post-1960  Material 
from  Coastal  Texas 

The  situation  within  100  miles  of  the  Texas 
Gulf  Coast  is  markedly  different  from  that 
found  farther  inland.  Red  wolves  survived 
in  the  southeastern  corner  of  Texas,  east  of 
the  Rrazos  River,  through  the  1930's  and 
1940's  (pp.  45-46).  Some  skulls  taken  farther 
south  along  the  coast  also  seem  to  resemble 
those  of  C.  rufus.  Halloran  (1961)  reported 
that  five  supposed  red  wolves  were  collected 
on  the  Aransas  National  Wildlife  Refuge  in 
1956,  but  the  listed  weights  were  much  less 
than  those  of  specimens  that  he  said  had 
been  taken  in  1939-1940.  Davis  (1966:112- 
113)  depicted  the  recent  range  of  C.  rufus 
as  extending  all  along  the  Gulf  Coast  from 
the  Louisiana  border  to  Raffin  Bay,  but  he 
observed  that  the  species  was  "on  the  verge 
of  extinction."  Beezley  (1967)  wrote  that 
red  wolves  were  present  on  the  coastal  prai- 
ries from  Jefferson  to  San  Patricio  counties, 
and  farther  inland  in  Harris,  Wharton,  Colo- 
rado, Lavaca,  and  Victoria  counties. 

Once    again,    my    analysis   of   material   is 


divided  on  a  partly  arbitrary  basis.  The  lo- 
calities, many  known  only  to  county,  are 
plotted  in  figure  31  and  listed  in  appendix  A 
(parts  33-38). 

Kenedy  County. — One  of  the  most  inter- 
esting of  the  coastal  specimens  was  taken  in 
December  1961  near  Armstrong,  Kenedy 
County,  not  far  north  of  the  Mexican  border. 
The  skin  and  skeleton  were  reported  as  C. 
rufus  by  Paradiso  ( 1965),  but  with  no  further 
comment.  The  statistical  position  of  the  large 
skull  falls  well  within  the  limits  of  the  stand- 
ard red  wolf  sample  (Fig.  37).  And  yet  the 
specimen  was  taken  in  an  area  beyond  the 
southern  edge  of  the  range  assigned  by  Gold- 
man (1944:487)  to  C.  r.  rufus.  Other  ma- 
terial, taken  south  of  Nueces  Bay  in  an  earlier 
period,  does  not  approach  the  Kenedy  County 
skull  in  size  or  other  characters,  and  appears 


Fie.  37. — Multivariate  positions  (squares)  of  in- 
dividual post- 1960  specimens  from  the  central  Gulf 
Coast  of  Texas,  relative  to  series  of  C.  rufus  and 
C.  latrans.  The  triangle  shows  the  position  of  an 
apparent  specimen  of  C.  rufus  collected  in  1961  in 
Kenedy  County,  extreme  southern  Texas. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


61 


to  represent  only  C.  latrans  (see  Fig.  20). 
No  additional  recent  specimens  from  the  area 
are  available,  but  further  investigation  may 
yet  reveal  evidence  of  the  presence  of  a 
small,  isolated  red  wolf  population. 

Central  Gulf  Coast. — Figure  37  shows  the 
relative  positions  of  skulls  of  24(7)  males  and 
18(5)  females,  taken  from  1962  to  1969  in  an 
area  including  Calhoun,  Victoria,  Matagorda, 
Lavaca,  Colorado,  Austin,  and  Fort  Rend 
counties.  No  character  clines  are  apparent 
in  this  area,  and  several  counties  were  the 
source  of  both  wolflike  and  coyotelike  speci- 
mens. Most  fall  between  the  extreme  limits 
of  C.  rufus  and  C.  latrans,  and  their  statistical 
distribution  is  similar  to  that  shown  by  the 
former  hybrid  swarm  of  central  Texas  (Fig. 
23).  Apparently,  significant  numbers  of  red 
wolves  survived  on  the  central  coast  of  Texas 
longer  than  farther  inland.  Coyotes  have  now 
occupied  the  same  area  and  hybridized  with 
the  wolves,  and  there  is  no  way  of  separating 
the  two  parent  species.  Specimens  taken  in 
the  1930's  and  1940's  (Fig.  26)  indicate  that 
coyotes  were  then  already  beginning  to  pre- 
dominate, but  evidently  red  wolf  influence 
persisted.  Possibly,  small  groups  of  relatively 
unmodified  C.  rufus  still  are  present  in  the 
area,  but  it  would  be  difficult  to  pinpoint 
their  locations. 

Harris  County. — The  vicinity  of  the  Ad- 
dicks  Reservoir,  just  west  of  Houston,  was  the 


RUFUS 


RUFUS 


Fig.  38. — Multivariate  positions  (squares)  of  in- 
dividual post-1960  specimens  from  Harris  County, 
Texas,  relative  to  series  of  C.   rufus  and  C.  latrans. 


Fig.  39. — Multivariate  positions  (squares)  of  in- 
dividual post-1960  specimens  from  western  Brazoria 
County,  Texas,  relative  to  series  of  C.  rufus  and  C. 
latrans. 


source  of  12(2)  males  taken  from  1964  to 
1971.  Seven  of  these  have  a  smaller  D2  sepa- 
ration from  C.  rufus  than  from  C.  latrans. 
The  over-all  statistical  distribution  (Fig.  38) 
suggests  that  the  coastal  hybrid  population  is 
established  in  the  area,  but  that  red  wolf  in- 
fluence still  is  strong.  On  the  basis  of  investi- 
gation of  the  ecology,  behavior,  morphology, 
and  allelic  frequency  of  canids  taken  mainly 
in  southeastern  Harris  County,  Shaw  (1975: 
95)  suggested  that  the  population  there  rep- 
resented hybridization  between  C.  rufus  and 
C.  latrans. 

Western  Brazoria  County. — From  1968  to 
1971,  17(13)  males  and  14(10)  females  were 
collected  in  the  vicinity  of  the  Clemens  Prison 
Farm,  south  of  the  town  of  Rrazoria  and  west 
of  the  Brazos  River.  Their  statistical  distribu- 
tion (Fig.  39)  is  indicative  of  a  hybrid  popu- 
lation bridging  the  gap  between  C.  rufus  and 
C.  latrans. 


62 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


Eastern  Brazoria  County. — Paradiso  and 
Nowak  ( 1972a )  reported  that  typical  C.  rufus 
rufus  still  lived  in  Brazoria  County  east  of  the 
Brazos  River.  Russell  and  Shaw  (1971b)  lo- 
cated red  wolves  in  the  same  area,  through 
elicitation  of  howling  responses.  A  total  of 
11(5)  male  and  9(4)  female  adult  skulls, 
some  of  them  not  available  for  Paradiso  and 
Nowak's  earlier  work,  were  tested  by  multi- 
variate analysis.  All  of  this  material  was  ob- 
tained between  1966  and  1971  in  the  east- 
central  part  of  the  county,  in  the  vicinity  of 
Angleton,  Liverpool,  and  Hopkins  Mound. 
Statistical  positions  (Fig.  40)  are  concen- 
trated along  the  lower  limits  of  the  standard 
red  wolf  sample,  not  unlike  those  of  older 
material  assigned  to  the  subspecies  C.  r. 
rufus.  Four  of  the  specimens,  however,  ac- 
tually have  a  greater  D2  separation  from  C. 
rufus  than  from  C.  latrans.  While  it  thus  is 
evident  that  gene  flow  from  coyotes  has  had 
an  effect  in  this  area,  the  influence  of  C.  rufus 


RUFUS 


Fig.  40. — Multivariate  positions  (triangles)  of  in- 
dividual specimens  taken  from  1966  to  1971  in  east- 
ern Brazoria  County,  Texas,  relative  to  series  of  C. 
rufus  and  C.  latrans. 


seems  stronger  here  than  farther  west  on  the 
Gulf  Coast.  Several  other  skulls  from  eastern 
Brazoria  County,  not  suitable  for  multivariate 
analysis,  were  examined  and  found  to  possess 
typical  red  wolf  characters.  One  of  these 
skulls  was  241.0  millimeters  in  greatest  length. 
Recent  reports  from  field  personnel,  however, 
suggest  that  red  wolf  influence  in  this  area 
has  declined,  and  conservation  efforts  there 
have  been  largely  discontinued. 

Jasper,  Tyler,  Liberty,  and  ivestern  Cham- 
bers counties. — Red  wolves  held  out  for  many 
years  in  the  area  of  the  Big  Thicket  to  the 
northeast  of  Houston.  Currently,  however,  it 
is  not  likely  that  the  species,  in  unmodified 
form,  is  present  in  the  area.  According  to 
recent  reports,  it  seems  that  the  expanding 
coyote  population  has  occupied  the  wood- 
lands of  southeastern  Texas  in  moderate 
numbers  (Russell  and  Shaw,  1971b;  Glynn 
Riley,  U.S.  Fish  and  Wildlife  Service,  pers. 
comm.).  The  status  of  wild  Canis  here  is  not 
well  understood,  and  I  have  examined  no 
specimens  from  within  the  borders  of  the  Big 
Thicket  itself.  The  positions  of  3(1)  males 
taken  in  1970  on  the  eastern  side  of  the 
Thicket,  two  near  Fred,  Tyler  County,  and 
one  near  New  Blox,  Jasper  County,  indicate 
that  the  influence  of  C.  rufus  is  still  strong 
in  the  area  (Fig.  41).  Skulls  of  11  males  and 
10  females  were  taken  from  1965  to  1971 
along  the  lower  edge  of  the  Big  Thicket  in 
southern  Liberty  and  western  Chambers 
counties.  This  area  contains  a  mixture  of 
coastal  prairie  and  woodland  habitat.  Some 
of  the  specimens  are  indistinguishable  from 
red  wolves  collected  many  years  ago  in  the 
south-central  states,  but  others  are  coyotelike 
or  intermediate  in  characters  (Fig.  41).  Ap- 
parently then,  coyotes  have  spread  through 
east  Texas,  and  have  come  into  contact  with 
surviving  red  wolves  on  the  coastal  prairies 
where   hybridization  probably  has  occurred. 

Jefferson  and  eastern  Chambers  counties. 
— Even  while  the  disappearance  of  red  wolves 
over  most  of  their  former  range  was  being 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


63 


RUFUS 


Fig.  41. — Multivariate  positions  of  individual 
specimens  taken  from  1965  to  1971  in  Jasper  and 
Tyler  counties,  Texas  (open  circles),  and  in  southern 
Liberty  and  western  Chambers  counties  (squares), 
relative  to  series  of  C.  rufus  and  C.  latrans. 

generally  acknowledged,  there  were  continu- 
ous reports  of  their  survival  on  the  coastal 
prairies  and  marshes  of  extreme  southeastern 
Texas  between  Galveston  Bay  and  Sabine 
Lake.  Paradiso  (1965)  reported  that  seven 
large  skulls,  collected  in  1963  or  early  1964 
on  the  Anahuac  National  Wildlife  Refuge, 
southeastern  Chambers  County,  were  unques- 
tionably those  of  C.  rufus.  Pimlott  and  Joslin 
(1968)  wrote  that  in  1965  they  located  three 
packs  of  red  wolves  in  the  area  east  of  Gal- 
veston Bay  between  Highway  73  and  the 
Gulf  Coast.  Paradiso  and  Nowak  (1972a) 
considered  that  specimens  collected  between 
1963  and  1969  demonstrated  the  survival  of 
the  subspecies  C.  rufus  gregoryi  in  the  area. 
For  the  purposes  of  this  paper,  skulls  of 
15(3)  males  and  four  females  taken  from 
1963  to  1970  were  tested  by  multivariate 
analysis.    All  skulls  are  from  animals  killed 


in  eastern  Chambers  and  southern  Jefferson 
counties,  the  area  between  Galveston  Bay  and 
Sabine  Lake,  and  south  of  the  line  formed  by 
U.S.  Interstate  Highway  10  and  Texas  High- 
way 73.  The  relative  statistical  positions  of 
these  specimens  (Fig.  42)  fall  predominantly 
within  the  range  of  variation  of  standard  C. 
rufus,  and  confirm  the  presence  of  the  species, 
in  apparently  unmodified  form,  in  extreme 
southeastern  Texas  (see  also  measurements 
in  appendix  B,  part  7). 

The  suggestion  that  the  red  wolf  survived 
in  southeastern  Texas  until  about  1970  was 
supported  by  field  studies  carried  out  in  that 
area.  Glynn  Riley,  an  agent  of  the  U.S.  Fish 
and  Wildlife  Service,  who  was  locally  respon- 
sible for  red  wolf  management  from  1969  to 
1973,  said  (pers.  comm.)  that  the  species  was 
present  in  moderate  to  high  numbers  in 
Chambers  and  Jefferson  counties.   In  Septem- 


A    RUFUS 


n r- 

2      3 


-3  - 


-3    -2     -I 


0 


I 


3     4 


Fig.  42. — Multivariate  positions  (triangles)  of  in- 
dividual specimens  taken  from  1963  to  1970  in  Jef- 
ferson and  eastern  Chambers  counties,  extreme  south- 
eastern Texas,  relative  to  series  of  C.  rufus  and  C. 
latrans. 


64 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


ber  1972  he  estimated  the  population  south 
of  Interstate  Highway  10  to  consist  of  about 
150  individuals.  Russell  and  Shaw  (1971b) 
wrote  that  in  1970  they  located  a  dense  red 
wolf  population  in  southern  Chambers  and 
Jefferson  counties,  through  elicitation  of 
howling  responses.  Shaw  (1975)  reported  a 
relatively  high  population  density  of  C.  rufus 
gregonji  in  the  area  during  his  field  work  from 
June  1971  to  September  1972,  and  found  the 
animals  so  identified  to  have  gross  morpho- 
logical and  behavioral  characters  distinct  from 
those  of  C.  latrans. 

Probably  an  important  factor  in  the  sur- 
vival of  these  wolves  was  their  geographic 
location.  Far  to  the  south  along  the  Gulf 
Coast,  they  were  well  removed  from  the  orig- 
inal and  subsequently  expanding  coyote  pop- 
ulation. The  temporary  existence  in  eastern 
Texas  of  a  large  zone  in  which  wild  Cam's 
was  rare  or  absent  (see  pp.  52-53),  helped 
to  prevent  genetic  exchange  between  coyotes 
to  the  north  and  the  surviving  wolves  on  the 
coast.  To  the  west,  Galveston  Bay  and  the 
Houston  metropolitan  area  limited  the  spread 
of  the  hybridization  process  that  engulfed 
most  of  the  coast. 

In  my  dissertation  (Nowak,  1973:146)  I 
expressed  hope  that  conservation  efforts 
might  help  to  maintain  the  unmodified  popu- 
lation of  C.  rufus  in  southeastern  Texas.  Even 
before  I  wrote  those  words,  however,  the 
situation  probably  had  deteriorated  beyond 
the  point  of  practical  control.  Indeed,  there 
is  good  evidence  that  canids  with  coyotelike 
characteristics  were  already  in  Jefferson  and 
eastern  Chambers  counties  in  the  1960's,  and 
that  the  skulls  of  such  animals  may  never 
have  been  submitted  for  examination,  be- 
cause persons  obtaining  specimens  thought 
that  only  "wolves"  were  wanted  (Carley  and 
McCarley,  1976).  This  bias  probably  was 
not  an  overriding  factor,  however,  since  both 
wolflike  and  coyotelike  skulls  were  received 
from  many  other  parts  of  Texas  (see  Figs. 
36-41).  While  it  is  likely  that  a  portion  of 
the  population  of  wild  Canis  in  Jefferson  and 


eastern  Chambers  counties  in  the  1960's  did 
not  represent  unmodified  C.  rufus,  I  continue 
to  think  that  this  portion  was  small  relative 
to  other  areas  of  coastal  Texas. 

In  any  event,  the  situation  does  appear  to 
have  worsened  in  the  late  1960's  and  early 
1970's.  Conservation  efforts  by  the  U.S.  Fish 
and  Wildlife  Service,  including  some  trapping 
of  non-wolflike  canids  in  the  vicinity,  did  not 
prevent  the  continued  decline  of  the  red  wolf. 
By  1974,  if  not  sooner,  field  personnel  recog- 
nized that  a  substantial  number  of  the  wild 
canids  south  of  Interstate  Highway  10  were 
not  wolves  (Curtis  J.  Carley,  Project  Leader, 
Red  Wolf  Recovery  Program;  pers.  comm.). 
Skulls  of  13  males  and  12  females,  collected 
from  1973  to  1975  in  the  same  area  as  those 
represented  in  figure  42,  were  evaluated  by 
multivariate  analysis.  The  statistical  positions 
of  these  new  specimens  are  shown  in  figure 


RUFUS 


Fig.  4.3. — Multivariate  positions  (squares)  of  in- 
dividual specimens  taken  from  1973  to  1975  in  Jef- 
ferson and  eastern  Chambers  counties,  extreme  south- 
eastern Texas,  relative  to  series  of  C.  rufus  and 
C.  latrans. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


65 


43,  and  the  general  pattern  resembles  that  of 
material  taken  just  to  the  north  from  1965 
to  1971  (Fig.  41).  Thus  it  is  evident  that 
eoyotelike  animals  have  now  completely  oc- 
cupied the  former  range  of  C,  ruftts  in  Texas, 
and  that  no  red  wolf  populations  (though 
probably  some  individuals)  remain  free  from 
significant  genetic  influence  by  C.  latrans. 

When  it  was  recognized  that  the  last  red 
wolf  population  was  being  swamped  through 
hybridization,  the  U.S.  Fish  and  Wildlife 
Service  accelerated  a  program  to  live-capture 
some  individuals  that  did  appear  to  represent 
pure  C.  nifus,  and  to  place  these  animals  in 
a  breeding  pool  at  the  Point  Defiance  Zoo  at 
Tacoma,  Washington.  In  November  1977 
this  facility  contained  29  adults,  all  wild- 
caught  (including  three  from  Cameron  Par- 
ish, Louisiana),  and  13  young  produced  dur- 


ing the  spring  of  1977  (Norman  R.  Winnick, 
Director,  Red  Wolf  Captive  Rreeding  Pro- 
gram, Point  Defiance  Zoo;  pers.  comm.).  On 
13  December  1976  a  pair  from  this  group  had 
been  released  on  Rulls  Island,  Cape  Romain 
National  Wildlife  Refuge,  South  Carolina. 
Both  animals  were  recaptured  on  22  Decem- 
ber 1976  after  one  had  crossed  to  the  main- 
land, and  the  female  subsequently  died  of 
natural  causes.  Another  release  was  planned 
for  January  1978.  Although  Bulls  Island  is 
too  small  (5,000  acres)  to  support  a  viable 
pack  of  wolves,  it  is  hoped  that  procedures 
can  be  developed  there  that  will  aid  possible 
future  reintroductions  in  other  parts  of  the 
original  range  of  C.  rufus.  At  present,  how- 
ever, the  wild  wolf  populations  that  once  in- 
habited the  southeastern  quarter  of  North 
America  are  not  in  existence. 


SYSTEMATIC  DESCRIPTIONS 


ORDER  CARNIVORA 
FAMILY  CANIDAE 
Genus  Canis  Linnaeus 

1758.  Canis  Linnaeus,  Systema  Naturae,  10th 
ed.,  p.  38.  Type,  Canis  familiaris  Lin- 
naeus. 

1816.  Thos  Oken,  Oken's  Lehrbuch  der  Na- 
turgeschichte,  pt.  3  (Zoologie),  sect. 
2,  p.  1037.  Type,  Thos  vulgaris  Oken 
(=Canis  aureus  Linnaeus). 

1839.  Lyciscus  Hamilton-Smith,  in  The  nat- 
uralist's library  (edit.  Jardine),  25:160. 
Type,  Canis  latrans  Say. 

1918.  Aenocyon  Merriam,  Univ.  California 
Publ.  Bull.  Dept.  Geol..  10:532.  20 
April.  Type,  Canis  dims  Leidy,  subse- 
quent designation  by  Hay,  Second  bib- 
liography and  catalogue  of  the  fossil 
Vertebrata  of  North  America,  2:501, 
27  January  1930. 

Geological  distribution. — Middle  Pliocene 
to  Recent. 

Geographical  distribution. — In  historical 
time,  the  species  of  Canis  have  been  distrib- 
uted as  follows: 

Canis  familiaris  Linnaeus,  1758  (including 
C.  dingo  Blumenbach,  1780,  of  Australia),  the 
domestic  dog,  throughout  the  world,  usually 
in  association  with  man; 

Canis  lupus  Linneaus,  1758,  the  gray  or 
timber  wolf,  in  most  of  North  America  and 
Eurasia,  and  on  associated  northern  islands; 

Canis  rufus  Audubon  and  Bachman,  1851, 
the  red  or  southern  wolf,  in  southeastern 
North  America; 

Canis  latrans  Say,  1823,  the  coyote  or  prai- 
rie wolf,  primarily  in  western  North  America; 

Canis  aureus  Linnaeus,  1758  (including 
C.  lupaster  Hcmprich  and  Ehrenberg,  1832, 
of  northern  Africa),  the  golden  jackal,  from 
central  Africa  to  India  and  southeastern  Eu- 
rope; 

Canis  adustus  Sundevall,   1846,   the  side- 


striped  jackal,  in  eastern,  southern,  and  west- 
ern Africa; 

Canis  mesomelas  Schreber,  1778,  the 
black-backed  jackal,  in  eastern  and  southern 
Africa;  and 

Canis  simensis  Ruppell,  1835,  the  Abys- 
sinian wolf,  in  the  mountains  of  Ethiopia. 

If  most  species  assigned  to  the  genus 
Dusicyon  are  actually  referable  to  Canis,  as 
suggested  by  Langguth  ( 1975 ) ,  the  Recent 
geographical  distribution  of  Canis  would  in- 
clude much  of  South  America.  Pleistocene 
fossils  of  Canis  have  been  reported  from 
South  America,  as  well  as  from  North  Amer- 
ica, Eurasia,  and  Africa. 

Relationship  with  other  Recent  genera. — 
The  Recent  Canidae  often  have  been  divided 
into  three  subfamilies:  Caninae,  with  the 
genera  Canis,  Alopcx,  Fennecus,  Vulpes, 
Urocyon,  Dusicyon.  Chrysocyon,  Atelocynus, 
Cerdocyon,  and  Nyctereutes;  Simocyoninae, 
with  the  genera  Cuon,  Lycaon,  and  Speothos; 
and  Otocyoninae,  with  the  genus  Otocyon. 
Van  Valen  (1964)  suggested  that  Otocyon 
does  not  warrant  subfamilial  distinction  from 
the  Caninae,  and  most  workers  seem  to  be 
following  this  viewpoint.  Clutton-Brock,  Cor- 
bett,  and  Hills  (1976)  proposed  abandoning 
the  use  of  all  subfamily  names  for  the  Recent 
Canidae.  These  authors  also  placed  Urocyon 
and  Fennecus  within  the  genus  Vulpes,  and, 
like  Osgood  (1934)  and  Simpson  (1945:109), 
included  Cerdocyon  and  Atelocynus  in  the 
genus  Dusicyon.  Langguth  (1975),  however, 
retained  the  genera  Cerdocyon  and  Atelocy- 
nus, put  all  but  one  of  the  species  of  Du- 
sicyon in  the  genus  Canis,  and  placed  the 
remaining  species  (Dusicyon  vetulus)  in  the 
genus  Lycalopcx. 

Osgood  (1934)  prepared  a  systematic  key 
in  an  attempt  to  demonstrate  the  common 
affinity  of  all  South  American  Caninae  and 
their  collective  distinction  from  Canis.  The 
South  American  genera  (including  Urocyon) 


66 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


67 


were  said  to  be  characterized  by  a  reduced 
upper  carnassial,  with  a  length  always  less 
than  the  combined  length  of  the  two  upper 
molars;  an  occipital  shield  depressed  at  the 
apex;  a  usually  flattened  or  lyrate  sagittal 
crest  (except  in  Chrysocyon);  relatively  small 
upper  incisors,  the  inner  pair  imperfectly  or 
not  trifid;  and  long,  slender  canine  teeth. 
These  characters,  however,  may  be  only  rela- 
tive expressions  of  trends  found  throughout 
the  Caninae.  For  example,  in  some  North 
American  coyotes  the  saggital  crest  is  flat- 
tened and  lyrate,  and  P4  is  shorter  than  the 
combined  length  of  Ml  and  M2. 

Study  of  canid  karyotypes  thus  far  seems 
to  support  the  composition  of  the  genus  Canis 
as  set  forth  above  under  "geographical  dis- 
tribution." Each  species  of  Canis  that  has 
been  analyzed  (C.  familiaris,  C.  dingo,  C. 
lupus,  C.  rufus,  C.  latrans,  and  C.  aureus)  has 
a  diploid  chromosome  number  of  78.  Species 
of  other  genera  within  the  Caninae,  including 
Vulpes,  Fennecus,  Alopex,  Urocyon,  Chryso- 
cyon,  Nyctereutes,  and  Otocyon,  have  been 
found  to  have  diploid  numbers  different  from 
that  of  Canis.  The  species  Dusicyon  vetulus 
(placed  in  the  genus  Lycalopex  by  Lang- 
guth,  1975)  has  a  diploid  number  of  76 
(Wurster  and  Benirschke,  1968;  Chiarelli, 
1975). 

The  production  of  hybrids  provides  fur- 
ther evidence  of  the  validity  of  the  presently 
accepted  structure  of  the  genus  Canis.  Viable 
hybrids  have  been  reported  between  C.  fa- 
miliaris and  C.  lupus,  C.  familiaris  and  C. 
latrans,  C.  familiaris  and  C.  aureus,  C.  famili- 
aris and  C.  dingo,  C.  latrans  and  C.  aureus, 
C.  latrans  and  C.  lupus,  C.  lupus  and  C. 
dingo  (Gray,  1972:45-51),  and  C.  latrans  and 
C.  rufus  (this  paper).  There  are  few  data  on 
interbreeding  between  members  of  Canis  and 
other  recognized  genera,  and  no  evidence  that 
viable  hybrids  ever  have  been  produced  (see 
Chiarelli,  1975:50). 

Remarks. — Determination  of  the  actual 
origin  of  the  genus  Canis  is  not  within  the 
scope  of  this  paper.    One  widely  held  view 


is  that  the  genus  Tomarctus  of  the  Miocene 
and  early  Pliocene  is  near  the  ancestral  line 
of  Canis  and  other  modern  Caninae.  Simpson 
(1945:109,  222)  placed  Tomarctus  within  the 
Caninae,  but  noted  that  it  might  represent 
the  ancestry  of  another  subfamily.  Matthew 
(1930:131)  and  Vanderhoof  and  Gregory 
(1940:145)  considered  Tomarctus  to  occupy 
a  central  position  from  which  both  the  various 
borophagine  dogs  and  the  modern  canines 
arose.  Green  (1948)  concluded  that  certain 
species  of  Tomarctus  were  in  the  direct  line 
of  the  borophagines,  whereas  other  species 
probably  gave  rise  to  Canis  and  possibly  to 
Vulpes.  Williams  (1962)  referred  Tomarctus 
to  the  Borophaginae  and  placed  all  but  one 
of  its  named  species  in  the  genus  Aelurodon. 
He  suggested  that  some  lesser  known  canine 
of  the  late  Miocene  or  early  Pliocene,  such  as 
Leptocyon  vafer  (Leidy),  was  the  ancestor  of 
the  Quaternary  Caninae.  Presumably  then, 
according  to  this  interpretation,  the  lineage 
leading  to  the  living  Caninae  has  been  dis- 
tinct from  other  subfamilial  groupings  at  least 
since  the  late  Miocene,  and  Canis  has  de- 
scended from  small,  foxlike  ancestors.  At  pres- 
ent, the  number  of  reported  Pliocene  canines 
is  small,  and  the  particular  point  at  which 
Canis  arose  has  not  been  established. 

It  is  true  that  many  North  American  speci- 
mens from  the  Pliocene  and  even  earlier 
times,  some  of  them  representing  animals 
larger  than  a  modern  wolf,  were  named  as 
species  of  Canis.  Nearly  all  of  these  names, 
however,  now  have  been  referred  to  various 
other  genera,  and  they  seem  to  have  no  bear- 
ing on  the  lineage  of  Canis  (Hay,  1902:769- 
776,  and  1929-1930:488-512  may  be  consulted 
for  these  named  kinds  and  their  places  of 
description).  Most  recently,  the  species  Canis 
texanus  Troxell,  1915  was  referred  to  the 
genus  Protocyon  by  J.  L.  Kraglievich  (1952: 
621). 

Only  three  Pliocene  species  of  Canis  are 
now  known  from  North  America.  These  in- 
clude C.  davisii  from  southwestern  Nevada 
(Merriam,    1911:242-243)     and    southeastern 


68 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


Oregon  (Shotwell,  1970:70),  and  C.  condoni 
from  northeastern  Oregon  (Shotwell,  1956: 
733).  Shotwell  (1970)  noted  that  additional 
material  might  prove  these  two  species  to  be 
synonymous.  Canis  lepophagus,  discussed  in 
detail  below,  is  known  from  a  number  of  late 
Pliocene  and  early  Pleistocene  sites. 

Of  the  various  specific  names  applied  to 
North  American  Pleistocene  fossils  of  Canis, 
the  following  17  have  not  hitherto  been 
placed  in  other  genera  or  formally  reduced 
to  synonyms  or  subspecies: 

C.  cedazoensis  Mooser  and  Dalquest, 

1975* 
C.  lepophagus  Johnston,  1938° 
C.  latrans  Say,  1823° 
C.  andersoni  Merriam,  1910 
C.  riviveronis  Hay,  1917 
C.  caneloensis  Skinner,  1942 
C.  irvingtonensis  Savage,  1951 
C.  edwardii  Gazin,  1942° 
C.  rufus  Audubon  and  Bachman,  1851° 
C.  priscolatrans  Cope,  1899 
C.  armbrusteri  Gidley,  1913° 
C.  lupus  Linnaeus,  1758° 
C.  milled  Merriam,  1912 
C.  familiaris  Linnaeus,  1758° 
C.  petrolei  Stock,  1938 
C.  dims  Leidy,  1858° 
C.  ayersi  Sellards,  1916. 

The  status  of  all  of  these  names  is  dis- 
cussed in  the  following  pages.  Each  of  those 
marked  above  with  an  asterisk  ( ° )  is  main- 
tained as  a  specific  name  in  this  paper. 

Canis  cedazoensis  Mooser  and  Dalquest 

1975.    Canis    cedazoensis    Mooser    and    Dal- 
quest, Jour.  Mamm.,  56:787. 

Holotype. — Right  maxillary  fragment  con- 
taining P3  to  Ml  and  the  alveolus  of  M2;  no. 
9780,  Midwestern  State  University  Depart- 
ment of  Biology;  Arroyo  Cedazo,  3  kilometers 
SE  City  of  Aguascalientes,  Aguascalientes, 
Mexico. 

Geological  distribution. — Early  Ranchola- 
brean. 


Geographical  distribution. — Known  only 
from  the  type  locality. 

Description. — A  small  canid,  larger  than 
any  North  American  fox,  but  smaller  than  C. 
latrans;  Ml  relatively  small,  with  pronounced 
buccal  cingulum  (for  more  detail,  see  Mooser 
and  Dalquest,  1975:787-788). 

Comparison  with  C.  latrans. — Smaller;  Ml 
with  relatively  smaller  medial  section. 

Remarks. — The  single  available  specimen 
is  too  fragmentary,  and  the  teeth  too  heavily 
worn,  to  allow  the  same  kind  of  account  given 
to  other  species  in  this  paper.  In  general  I 
am  in  accord  with  the  description  and  com- 
parisons provided  by  Mooser  and  Dalquest 
(1975:787-788),  and  with  their  conclusion 
that  the  specimen  does  not  represent  C.  la- 
trans. 

Record  of  occurrences. — Type  locality; 
early  Rancholabrean  (probably  Illinoian); 
specimen  examined:  holotype,  Midwestern 
State  University  Department  of  Biology  9780 
(formerly  no.  FC  634  in  collection  of  O. 
Mooser);  measurement  in  appendix  B  (part 
8). 

Evolutionary  position. — On  the  basis  of 
the  one  available  specimen,  the  relationships 
of  C.  cedazoensis  can  not  be  carefully  as- 
sessed. The  species  might  represent  an  aber- 
rant line  that  separated  from  C.  latrans  in  the 
Irvingtonian,  or  it  could  be  a  surviving  ele- 
ment of  one  of  the  smaller-sized  populations 
of  the  Blancan  C.  lepophagus. 

Canis  lepophagus  Johnston 

1938.    Canis     lepophagus     Johnston,     Amer. 
Jour.  Sci.,  ser.  5,  35:383. 

Holotype. — Skull  without  mandibles;  no. 
W.  T.  881,  Panhandle  Plains  Museum; 
stratum  no.  2,  Harold  Ranch,  North  Cita  Can- 
yon, center  of  west  half  of  sec.  164,  block  6, 
Randall  County,  Texas. 

Geological  distribution. — Blancan. 

Geographical  distribution. — Known  from 
California,  Florida,  Idaho,  Kansas,  Nebraska, 
and  Texas. 

Description. — A    small    coyotelike    canid; 


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NOWAK:    NORTH  AMERICAN  QUATERNARY  CAN1S 


skull  small  with  mostly  narrow  proportions; 
rostrum  elongated  and  narrow;  braincase  rela- 
tively small  and  little  inflated  dorsoposte- 
riorly;  frontal  and  supraoccipital  shields  rela- 
tively broad;  sagittal  crest  prominent;  mandi- 
ble long  and  narrow;  premolars  and  molars 
with  trenchant,  laterally  compressed  cusps; 
P4  with  prominent  deuterocone;  Ml  with 
pronounced  buccal  cingulum;  p4  with  promi- 
nent second  cusp. 

Comparison  with  C.  latrans. — Skull  aver- 
aging smaller,  but  overlapping  in  most  meas- 
ured dimensions;  braincase  relatively  smaller 
and  less  inflated  dorsoposteriorly;  frontal  and 
supraoccipital  shields  relatively  broader;  sa- 
gittal crest  more  prominent;  some  mandibles 
relatively  deeper;  dentition  usually  more 
crowded  and  with  more  trenchant,  laterally 
compressed  cusps;  metaconule  on  Ml  less 
separated  from  protocone;  p4  usually  with 
more  prominent  second  cusp,  and  with  third 
cusp  and  posteromedial  cingulum  more  re- 
duced (if  they  are  present);  anterior  margin 
of  ml  usually  more  nearly  vertical.  Bjork 
(1970:14)  and  Kurten  (1974)  discussed  a 
number  of  postcranial  differences  between 
the  two  species. 

Other  comparison. — See  account  of  C. 
edwardii. 

Remarks.—  Johnston  (1938)  wrote  that  C. 
lepophagus  differed  from  C.  latrans  in  having 
more  prominent  sagittal  and  lambdoidal 
crests,  less  expanded  braincase,  deeper  man- 
dible, and  lesser  distance  between  premolars. 
Subsequently,  other  workers  assigned  addi- 
tional specimens  to  C.  lepophagus,  but  not 
all  of  this  material  shared  the  characters  cited 
by  Johnston.  Hibbard  (1938:243)  wrote  that 
a  specimen  from  the  Rexroad  fauna,  which 
he  later  ( 1941a: 268-269)  referred  to  C.  le- 
pophagus, represented  a  "small  dog  with  a 
light  tapering  ramus."  Hibbard  (1941a) 
stated  that  C.  lepophagus  from  this  fauna 
was  characterized  by  small  size  and  relatively 
narrow  teeth.  Fine  (1964)  wrote  that  a  man- 
dibular fragment,  assigned  later  by  Bjork 
(1970:14)    to   C.    lepophagus,   was   "from    a 


lightly  built  coyote  having  a  jaw  quite  long 
for  its  depth  as  compared  with  those  of  Re- 
cent specimens."  Bjork  (1970)  described  C. 
lepophagus  from  the  Hagerman  local  fauna 
as  having  a  long  and  slender  lower  jaw,  and 
dentition  more  slender  than  that  of  C.  latrans. 
Giles  (1960)  subjected  six  mandibles  of  C. 
lepophagus  from  Cita  Canyon  to  a  multi- 
variate analysis  involving  five  measurements. 
The  results  suggested  to  him  that  C.  lepoph- 
agus was  subspecifically,  but  not  specifically, 
distinct  from  Recent  C.  latrans. 

According  to  Kurten  (1974:27-28),  "Canis 
lepophagus  of  the  Blancan  differs  from  the 
Recent  and  Rancholabrean  C.  latrans  in  its 
shorter  distal  limb  segments,  more  tapering 
snout,  relatively  larger  M2  (this  character  is 
however  retained  in  some  Recent  popula- 
tions), narrower  p2-p4,  and  shorter  ml  (rela- 
tive to  its  own  width,  and  also  to  the  length 
of  p4)."  He  considered  all  coyotelike  ma- 
terial from  the  Blancan,  except  from  the  latest 
part  of  this  age,  to  represent  C.  lepophagus, 
and  that  this  species  formed  a  stage  in  the 
evolutionary  line  leading  to  C.  latrans.  Ma- 
terial from  most  of  the  localities  listed  below 
under  "record  of  occurrences,"  was  referred 
by  Kurten  to  C.  lepophagus. 

I  recognize  most  material  previously  as- 
signed by  others  to  C.  lepophagus  as  repre- 
senting a  single  variable  species,  separate 
from  and  ancestral  to  C.  latrans.  Hibbard 
(1941b)  thought  that  C.  lepophagus  had  as 
large  a  geographic  range  as  Recent  C.  latrans. 
Considering  this  likelihood,  plus  the  long 
period  of  time  through  which  material  of  C. 
lepophagus  is  distributed,  considerable  varia- 
tion would  be  expected.  The  three  skulls 
from  Cita  Canyon  reported  by  Johnston 
(1938),  plus  one  other  obtained  at  the  same 
site,  are  the  only  specimens  of  C.  lepophagus 
upon  which  meaningful  measurements  can  be 
made  of  elements  other  than  the  teeth  and 
lower  jaws.  And,  as  listed  above,  there  are 
differences  between  these  skulls  and  those  of 
C.  latrans.  The  mandibles  of  C.  lepophagus 
from  Cita  Canyon  overlap  in  size  and  pro- 


70 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


portion  those  of  Recent  C.  latrans  (Fig.  44), 
but  average  deeper.  Most  specimens  from 
other  sites  are  mandibles,  some  of  them  rela- 
tively shallow.  Differentiating  characters  of 
the  lower  dentition,   as  listed  in  the  above 


88 

- 

o 

87 

- 

• 

o  o 

86 

•      o          o   o 
•              •  o 

85 

• 
•         o     s   O 

84 

o.o  o0          o 
•    ,oo  •  s    • 

83 

o  •      o         c 
•  °    oc»  -°      o    ° 

82 

O    O    C         o 

81 

80 

#      b„  •o«c»  o 

79 

•• 

78 

••        ••    •    *o 

• 

•  •      •      c 

77 

• 

•• 

• 

••      • 

76 

• 
• 

c      •  • 

75 

•  •  • 

74 

•  ..   c 

73 

.• 

• 

72 

• 
s 

71 

s    s        • 

i 

14 

15    16     17     18    19    20   21    22 

Fig.  44. — Scatter  diagram  comparing  length  and 
depth  of  mandible  in  certain  specimens  of  C.  le- 
pophagus  and  C.  latrans.  The  vertical  axis  indicates 
measurement  of  the  distance  from  the  anterior  edge 
of  the  alveolus  of  pi  to  the  posterior  edge  of  the 
alveolus  of  m3.  The  horizontal  a\is  indicates  meas- 
urement of  the  minimum  depth  from  the  dorsal  sur- 
face of  the  mandible  between  p3  and  p4  to  the 
ventral  surface  of  the  mandible.  Black  dots,  Recent 
C.  latrans  testes;  open  circles,  Pleistocene  C.  latrans 
orcutti;  c,  C.  lepophagus  from  Cita  Canyon;  s,  C. 
lepophagus  from  other  sites;  v,  C.  latrans  irvingtonen- 
sis. 


"comparison  with  C.  latrans,"  are  present  in 
most  of  the  Cita  Canyon  mandibles  and  in 
most  of  the  other  specimens  that  can  be  eval- 
uated. These  characters  are  matched  in 
some  individuals  of  C.  latrans,  but  their  pres- 
ence in  specimens  of  C.  lepophagus  from 
several  major  sites  suggests  affinity  among 
the  populations  represented. 

The  type  locality  of  C.  lepophagus,  Cita 
Canyon,  now  is  recognized  to  contain  an 
Aftonian  (Hibbard,  1970:414)  or  late  Blancan 
(Kurten,  1974:5)  fauna,  and  thus  is  among 
the  youngest  sites  from  which  the  species 
has  been  reported.  Since  no  specimens  from 
contemporary  sites  are  complete  enough  for 
definite  assignment  to  C.  latrans,  but  since 
some  do  have  characters  typical  of  C.  lepoph- 
agus, all  coyotelike  material  from  the  Blancan 
is  referred  below  to  C.  lepophagus. 

Record  of  occurrences. — The  following 
list  is  arranged  alphabetically  by  state,  and 
geographically  (north  to  south,  west  to  east) 
within  states.  Specimens  examined  by  me 
are  identified  by  element,  museum  number, 
or  both;  and  selected  measurements  are  found 
in  appendix  B  (part  9)  and  appendix  C  (part 
1).  Occurrences  also  are  shown  on  the  map 
in  figure  45. 

CALIFORNIA.— Sacramento  Valley,  Sacramento 
River,  Tehama  County;  late  Pliocene;  as  Canis  (Van- 
derhoof,  1933:383);  ml,  UCMP  29828. 

FLORIDA.— Santa  Fe  River  IB,  Gilchrist  County; 
Blancan  (Nebraskan);  as  C.  cf.  lepophagus  (Webb, 
1974b:  17);  five  mandibular  fragments,  UF  10423, 
10424,  10836,  10837,  10858.  Two  of  these  speci- 
mens are  comparatively  large  and  deep,  and  sug- 
gestive of  the  Cita  Canyon  material. 

IDAHO. — Grand  View,  Owyhee  County;  late 
Blancan  (Hibbard,  et  al.,  1965);  as  "Canis  n.  sp." 
(Schultz,  1938a:297);  two  mandibular  fragments, 
LACM  118-1246,  1343.  The  specimens  are  com- 
paratively large  and  deep;  the  p4  has  a  prominent 
second  cusp,  but  no  third  cusp  or  posteromedial 
cingulum. 

Hagerman  local  fauna,  Twin  Falls  County;  early 
Blancan  (late  Pliocene);  as  C.  lepophagus  (Bjork, 
1970:13-16);  four  maxillary  fragments,  USNM 
25136,  UMMP  V52280,  V54995,  V56401;  seven  man- 
dibular fragments,  USNM  25131,  UMMP  V45222, 
V50249,  V53519,  V53817,  V53910,  V56282;  two 
Ml,  UMMP  V50000,  V56034;  Ml  fragment,  UMMP 
V57016;  p2,  UMMP  V56809;  two  p3,  UMMP 
V50008,  V51052;  p4  fragment,  UMMP  V53817;  ml, 


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71 


Fig.  45. — Map   showing  localities  of  C.   lepophagus    (black   dots)    and    C.    armbrusteri    (triangles).     Be- 
cause of  the  scale  of  the  map,  it  was  not  possible  to  plot  all  localities  in  crowded  areas. 


USNM  25132;  ml  fragment,  UMMP  V52757;  post- 
cranial  elements,  UMMP.  As  explained  by  Bjork 
(1970),  the  Hagerman  material  indicates  an  animal 
smaller  than   the  average  Recent  C.   latrans. 

KANSAS. — Rexroad   fauna,   Meade   County   State 
Park,  Meade  County;  early  Blancan   (late  Pliocene); 


as  C.  lepophagus  and  Canis  sp.  ( Hibbard,  1941a: 
268-269);  three  mandibular  fragments,  KU  3914, 
4602,  4603;  Ml,  m2,  KU  3915;  P4,  Ml,  UMMP 
37132.  The  mandibles  are  relatively  small  and  shal- 
low, and  most  of  the  teeth  are  small  with  trenchant 
cusps.     Hibbard   wrote   that   large   size   distinguished 


72 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


an  Ml  and  m2,  found  at  the  site,  from  C.  lepoph- 
agus.  Although  these  teeth  are  larger  than  the  others 
in  the  Rexroad  fauna,  they  are  within  the  size  range 
of  C.  lepophagus  from  other  sites. 

Rexroad  fauna,  Keefe  Canyon,  Meade  County; 
early  Blancan  (late  Pliocene);  as  C.  lepophagus 
(Hibbard  and  Riggs,   1949:838);  ml,  KU  7692. 

Deer  Park  local  fauna,  Meade  County;  late 
Blancan  (Aftonian);  as  Canis  sp.  (Hibbard,  1956: 
172);  two  ml,   UMMP  31945. 

NEBRASKA.— Sand  Draw,  6  mi.  N  Ainsworth, 
Brown  County;  late  Blancan  (Aftonian:  Hibbard, 
et  ah,  1965);  as  C.  cf.  latrans  ( McGrew,  1944:53), 
as  C.  lepophagus  (Skinner  and  Hibbard,   1972:107). 

Broadwater  quarry,  north  side  of  North  Platte 
River,  Morrill  County;  late  Blancan  (Aftonian);  as 
"Canis  sp.  (near  C.  latrans  Say)"  (Barbour  and 
Schultz,  1937:4),  as  "Canis  sp.  (Coyote)"  (Schultz 
and  Stout,  1948:563);  two  skulls,  UN  26111,  26112; 
mandibular  fragment,  UN  26116;  P4,  Ml,  M2,  UN 
26113.  Assignment  herein  to  C.  lepophagus  is  un- 
certain on  the  basis  of  morphology  alone,  as  the 
cranial  parts  are  in  too  poor  a  condition  for  evalua- 
tion of  specific  characters.  A  measurement  of  depth 
on  the  mandible  is  not  possible,  but  it  is  relatively 
thick  lateromedially,  and  has  the  massive  appearance 
attributed  to  C.  lepophagus  by  Johnston  (1938).  The 
teeth  are  approximately  the  same  size  and  have 
the  same  structure  as  those  of  the  Cita  Canyon  speci- 
mens. 

Lisco  quarry,  north  side  of  North  Platte  River, 
Garden  County;  late  Blancan  (Aftonian)-,  as  "Canis 
sp.  (Coyote)"  (Schultz  and  Stout,  1948:563);  skull, 
UN  26107;  mandible,  UN  26114.  This  material  offers 
some  evidence  for  recognizing  that  skulls  of  the  Cita 
Canyon  kind,  as  well  as  small  mandibles  from  other 
sites,  represent  the  same  species.  The  Lisco  mandi- 
ble is  comparatively  small  and  shallow,  with  trench- 
ant, laterally  compressed  cusps  on  the  teeth,  and 
with  a  high  second  cusp  on  p4.  The  skull  also  is 
small,  but  has  a  prominent  sagittal  crest,  as  do  the 
Cita   Canyon   skulls. 

TEXAS.— Red  Coral,  Proctor  Ranch,  Oldham 
County;  late  Blancan;  "within  the  variation  range 
of  C.  lepophagus"   (Kurten,   1974:5). 

Cita  Canyon,  Randall  County;  late  Blancan  (Af- 
tonian: Hibbard,  1970:414);  as  C.  lepophagus 
(Johnston,  1938);  four  skulls,  maxillary  fragment, 
32  mandibles  and  fragments  thereof,  various  isolated 
teeth  and  postcranial  elements,  PPM. 

Blanco  local  fauna,  near  Mount  Blanco,  Crosby 
County;  Blancan  (latest  Pliocene  or  earliest  Pleisto- 
cene); as  C.  lepophagus  (Dalquest,  1975:22,  47). 

Beck  Ranch,  Scurry  County;  early  Blancan;  as 
C.   lepophagus    (Kurten,    1974:5). 

Red  Light  Bolson,  southeastern  Hudspeth  County; 
Blancan  (Nebraskan);  as  C.  cf.  lepophagus  (Aker- 
sten,  1970:86;  1972:19-24). 

Evolutionary  position. — The  small  size 
and  the  dental  characters  of  some  specimens 
of  C.  lepophagus  suggest  affinity  to  the  Old 


World  jackals  and  to  the  foxes  (Vulpes).  The 
other  two  recognized  species  of  Pliocene 
Canis,  C.  davisii  and  C.  condoni,  also  were 
described  as  being  foxlike  in  size  and  certain 
other  details.  Possibly  then,  the  lineage  of 
Canis  separated  from  that  of  Vulpes  in  the 
early  or  middle  Pliocene.  Older  specimens 
of  C.  lepophagus  are  small  and  especially 
indicative  of  descent  from  foxlike  ancestors. 
The  species  C.  latrans  probably  evolved  from 
within  the  complex  assigned  above  to  C. 
lepophagus.  Perhaps  the  shallow-jawed  ma- 
terial represents  the  ancestral  element  of  C. 
latrans,  whereas  the  more  massive  Cita  Can- 
yon specimens  represent  another  line. 

In  my  dissertation  (Nowak,  1973:164),  I 
agreed  with  Johnston  ( 1938 )  in  recognizing 
C.  lepophagus  as  an  ancestral  coyote,  but 
unlike  him  I  did  not  consider  it  to  be  in  the 
direct  phylogenetic  line  of  the  modern  wolves. 
My  opinion  was  based  in  part  on  acceptance 
of  a  late  Blancan  age  for  the  type  of  C. 
edwardii  and  several  other  early  specimens 
of  the  wolf  line.  Since  the  most  wolflike 
specimens  of  C.  lepophagus,  especially  those 
from  Cita  Canyon,  also  were  late  Blancan  in 
age,  but  were  morphologically  distinct  from 
the  supposedly  contemporary  wolf  material, 
they  could  hardly  have  been  ancestral  to  C. 
edwardii.  New  information  (Johnson,  Op- 
dyke,  and  Lindsay,  1975;  Kurten,  1974)  sug- 
gests that  the  type  of  C.  edwardii,  as  well  as 
most  other  early  North  American  specimens 
that  apparently  represent  the  wolf  line,  ac- 
tually are  Irvingtonian  in  age.  The  one  ex- 
ception is  a  Blancan  specimen  from  Mifiaca 
Mesa,  Chihuahua,  which  consists  of  a  mandi- 
ble about  the  size  of  that  of  a  small  wolf, 
but  which  has  certain  dental  characters  re- 
sembling those  of  some  specimens  of  C.  le- 
pophagus (see  account  of  C.  edwardii  be- 
low). There  thus  is  a  possibility  that  this 
mandible  represents  a  transitional  phase 
through  which  the  wolf  line  did  evolve  from 
a  population  of  C.  lepophagus. 


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NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


73 


Canis  latrans  Say 

1823.    Canis  latrans  Say,  in  Long,  Account  of 
an  expedition  from   Pittsburgh  to  the 
Rocky    Mountains    performed    in    the 
years  1819  and  1820,  1:168.    Type  lo- 
cality. Engineer  Cantonment,  about  12 
mi.   SE  present  town  of  Blair,  Wash- 
ington County,  Nebraska,  on  west  bank 
of  Missouri  River. 
1910.    Canis  orcutti  Merriam,   Univ.   Califor- 
nia   Publ.    Bull.    Dept.    Geol.,    5:391. 
Type  from  Rancho  La  Brea,  Los  An- 
geles  County,   California.    Valid   as   a 
subspecies  of  C.  latrans. 
1910.    Canis  andersoni  Merriam,  Univ.  Cali- 
fornia Publ.  Bull.   Dept.  Geol.,  5:393. 
Type  from  Rancho  La  Brea,  Los  An- 
geles  County,  California.    A  synonym 
of  C.  latrans  orcutti. 
1917.    Canis   riviveronis   Hay,    Florida   State 
Geol.    Surv.    Rept.,    9:59.     Type    from 
Vero,    Indian    River   County,    Florida. 
Valid  as  a  subspecies  of  C.  latrans. 
1942.    Canis  caneloensis  Skinner,  Bull.  Amer. 
Mus.    Nat.   Hist.,   80:163.    Type   from 
Papago     Springs     Cave,     Santa     Cruz 
County,  Arizona.   Valid  as  a  subspecies 
of  C.  latrans. 
1951.    Canis  irvingtonensis  Savage,  Univ.  Cal- 
ifornia   Publ.    Bull.    Dept.    Geol.    Sci., 
28:231.    Type  from  Irvington,  Alameda 
County,    California.     Valid    as    a    sub- 
species of  C.  latrans. 
1961.    Canis    latrans    harriscrooki    Slaughter, 
Jour.  Mamm.,  42:505.  Type  from  Lew- 
isville    site,     Denton    County,    Texas. 
Valid  as  a  subspecies  of  C.  latrans. 

In  addition  to  those  listed  above,  22  names 
based  on  Recent  specimens  are  available  for 
use  at  the  subspecific  level.  These  are  to  be 
found  in  the  systematic  revision  by  Jackson 
( 1951 ) ,  and  also  are  listed  by  Hall  and  Kel- 
son  (1959:843-846). 

Type. — None  designated. 

Geological  distribution. — Irvingtonian  to 
Recent. 


Geographical  distribution. — Pleistocene 
and  early  Recent  records  from  Alberta,  Alas- 
ka, Arizona,  Arkansas,  California,  Colorado, 
Florida,  Idaho,  Illinois,  Indiana,  Iowa,  Kan- 
sas, Maryland,  Mississippi,  Missouri,  Ne- 
braska, Nevada,  New  Mexico,  Oklahoma, 
Oregon,  Pennsylvania,  Texas,  Utah,  Wiscon- 
sin, Wyoming,  Aguascalientes,  Estado  de 
Mexico,  Nuevo  Leon,  Oaxaca,  and  Puebla; 
original  range  in  historical  time  included 
most  of  western  half  of  North  America  and 
the  plains  region  as  far  east  as  southern 
Wisconsin,  northwestern  Indiana,  western  Ar- 
kansas, and  central  Texas;  see  also  "'remarks" 
below. 

Description. — Size    small    to   medium    for 
the   genus;    skull   usually   small   with    mostly 
narrow  proportions;   rostrum  elongated,  nar- 
row,  and  shallow;   braincase  relatively  large 
and     well     inflated     dorsoposteriorly,     often 
broader  at  level  of  parietotemporal  sutures 
than   at  base;   postorbital   constriction   broad 
lateromedially    and    short    anteroposteriorly; 
zygomata    usually    slender    and    not    widely 
spreading;  orbits  large;  frontals  usually  only 
slightly  elevated   above   rostrum,   not  promi- 
nently convex,  and  forming  relatively  narrow 
and  flat  shield;  temporal  ridges   usually  not 
sharp,   seldom  obscuring  frontal  suture,   and 
sometimes  forming  lyrate  pattern  over  brain- 
case;   sagittal  crest   seldom   prominent,   often 
thickened     lateromedially,      sometimes     flat- 
tened;    supraoccipital     shield     comparatively 
small;  external  side  of  occipital  usually  with 
thin-walled    projection    just    above    foramen 
magnum;  tympanic  bullae  well  inflated;  man- 
dible long,  narrow,  and  shallow,  ventral  edge 
often  rounded;   incisors  relatively   small,  up- 
per canines  prominent,  thin  anteroposteriorly, 
their  alveoli  set  low  in  premaxillae,  their  ven- 
tral tips   normally  extending   below   level   of 
anterior    mental    foramina    when    jaws    are 
closed;    premolars    with    trenchant,    laterally 
compressed   cusps;    P4   with   prominent   deu- 
terocone  and  lingual  cingulum;  Ml  with  rela- 
tively   large,    deeply   sculptured   medial    sec- 
tion,   prominent    metaconule    well    separated 


74 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


from  protocone,  usually  with  pronounced 
buccal  and  anterior  cingula;  M2  relatively 
large,  cusps  well  developed;  p2  usually  lack- 
ing posterior  cusp:  p.3  sometimes  with  second 
and  third  cusp;  p4  with  second  cusp,  and  usu- 
ally with  pronounced  third  cusp  and  postero- 
medial cingulum  extending  behind  third  cusp; 
m2  and  talonid  of  ml  relatively  large,  with 
trenchant  cusps.  For  details  on  pelage  and 
postcranial  skeleton,  see  Jackson  ( 1951 ) ; 
Grinnell,  Dixon,  and  Linsdale  (1937);  Hilde- 
brand  (1952a,  1952b,  1954);  and  Merriam 
(1912). 

Comparisons. — See  accounts  of  C.  ceda- 
zoensis,  C.  lepophagus,  C.  rufus,  C.  lupus, 
and  C.  familiaris. 

Remarks. — When  the  white  man  first  ar- 
rived in  North  America,  C.  latrans  had  a  wide 
distribution,  primarily  in  the  western  half  of 
the  continent.  The  exact  southern,  northern, 
and  eastern  limits  of  its  former  range  are  not 
known.  Young  (1951:29)  thought  that  the 
species  was  originally  found  only  as  far  as 
central  Mexico,  and  that  movement  farther 
to  the  south  occurred  after  the  introduction 
of  livestock  in  the  region.  But  Jackson  ( 1951) 
recognized  the  presence  of  three  separate 
subspecies  in  Central  America,  and  it  is  ques- 
tionable whether  all  of  these  could  have  come 
into  existence  in  only  400  years.  Also,  accord- 
ing to  Young  ( 1951 ) ,  the  coyote  did  not  be- 
come established  in  northwestern  Canada  and 
Alaska  until  the  nineteenth  and  twentieth 
centuries.  But  Jackson  (1951:265)  suspected 
that  the  subspecies  C.  latrans  incolatus  was 
a  very  long  time  resident  of  Alaska. 

Information  provided  in  the  first  main 
part  of  this  paper  indicated  that  the  original 
range  of  the  coyote  extended  at  least  as  far 
east  as  southern  Wisconsin,  northwestern  In- 
diana, western  Arkansas,  and  central  Texas. 
Skeletal  remains,  identified  as  C.  latrans, 
have  been  reported  from  archeological  ex- 
cavations in  Crawford  County,  Illinois  (Par- 
malee  and  Stephens,  1972);  Will  County, 
Illinois  (Parmalee,  1962b);  Madison  County, 
Illinois    (Parmalee,    1959a);    Phelps    County, 


Missouri  (Parmalee,  1965);  and  Washington 
County,  Arkansas  (Morrison,  1970).  Indian 
sites  from  farther  east  have  apparently  not 
yielded  specimens  of  C.  latrans,  but  in  the 
late  Pleistocene  the  species  ranged  as  far  east 
as  Pennsylvania  and  Florida.  Man's  extermi- 
nation of  the  larger  wolves,  and  disruption  of 
the  environment,  contributed  to  an  expansion 
of  the  coyote's  range  since  the  mid-nineteenth 
century.  The  species  is  now  established  in 
Ontario,  southern  Quebec,  New  England, 
New  York,  the  Ohio  Valley,  and  the  lower 
Mississippi  Valley;  and  introduced  individuals 
have  been  reported  from  all  states  of  the 
southeast. 

Specimens,  especially  skulls,  of  C.  latrans 
can  almost  always  be  easily  separated  from 
those  of  wolves.  Hybridization  under  com- 
pletely natural  conditions  seems  to  have  oc- 
curred rarely,  if  ever.  The  three  specimens 
described  on  page  11  are  the  only  ones 
known  from  the  western  half  of  the  continent 
that  appear  to  represent  hybridization  be- 
tween C.  lupus  and  C.  latrans.  More  exten- 
sive interbreeding,  probably  caused  by  man's 
influence,  has  taken  place  in  southeastern 
Canada.  This  process  has  permitted  the  in- 
trogression  of  wolf  genes  into  the  expanding 
coyote  population  of  the  region,  and  has 
resulted  in  a  modification  of  that  population 
(see  pp.  21-23).  Lawrence  and  Bossert 
( 1969 )  referred  to  the  existing  coyotes  of  the 
northeastern  United  States  as  "C.  latrans  var." 
Specimens  taken  there  in  recent  years  are 
variable,  but,  in  comparison  with  coyotes 
from  farther  west,  are  often  characterized  by 
larger  size,  more  massive  skull,  broader  and 
more  elevated  frontal  shield,  more  prominent 
sagittal  and  lamboidal  crests,  and  teeth  with 
less  trenchant  cusps. 

A  related  process  in  the  south-central 
states  has  allowed  introgression  of  genes  from 
C.  rufus  into  the  coyote  population  that  occu- 
pied the  region  in  the  twentieth  century.  In- 
dividuals of  this  population  generally  closely 
resemble  coyotes  from  farther  north  and  west 
in  proportions  and  dentition,  but  are  charac- 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


75 


terized  by  larger  skull,  more  prominent  sagit- 
tal and  lambdoidal  crests,  and  more  elongate 
postorbital   constriction. 

Kurten  (1974)  grouped  all  coyotelike  ma- 
terial from  the  Irvingtonian  and  latest  Blan- 
can  of  North  America  under  the  name  Canis 
priscolatrans,  which  he  considered  to  repre- 
sent an  intermediate  stage  in  the  evolutionary 
transition  from  C.  lepophagus  to  C.  latrans. 
He  acknowledged  that  there  were  difficulties 
in  drawing  specific  boundaries  in  this  se- 
quence, but  thought  that  differences  in  limb 
proportions,  between  Rancholabrean  C.  la- 
trans  and  available  Irvingtonian  specimens, 
supported  a  separation  of  species  at  that 
point.  In  addition,  C.  priscolatrans  was  re- 
ported to  be  considerably  larger,  on  the  av- 
erage, than  C.  latrans  or  C.  lepophagus,  and 
to  differ  from  those  species  in  certain  dental 
proportions. 

It  is  important  to  note  that  Kurten's 
(1974)  delineation  of  C.  priscolatrans  was 
based  in  part  on  his  inclusion  within  that 
species  of  several  large  specimens  which  had 
been  assigned  by  me  (Nowak,  1973)  or 
others  to  the  wolf  line.  Among  this  material 
is  the  type  of  C.  priscolatrans,  which  I  con- 
sidered not  more  than  subspecifically  distinct 
from  C.  rufus;  the  type  of  C.  edwardii,  which 
Gazin  (1942)  and  I  considered  to  be  closely 
related  to  C.  rufus;  and  specimens  from  the 
Inglis  IA  fauna  in  Florida  which  were  re- 
ferred to  C.  rufus  by  Webb  (1974b:  17).  I 
continue  to  think  that  this  material  repre- 
sents the  wolf  line,  and  that  several  other 
(but  certainly  not  all)  Irvingtonian  or  late 
Blancan  specimens,  assigned  by  Kurten  to 
the  coyote  line,  probably  are  referable  to  C. 
rufus  or  C.  edwardii.  If  such  specimens  are 
removed  from  consideration,  we  find  that  the 
Irvingtonian  coyotes  are  not  notably  larger 
than  those  of  the  Rancholabrean.  Although 
I  did  not  make  a  detailed  analysis  of  limb 
bones,  as  did  Kurten,  he  apparently  evaluated 
only  three  specimens  designated  by  him  as 
C.  priscolatrans,  one  of  which  was  part  of  the 
Inglis  material  referred  to  C.  rufus  by  Webb 


(1974b:  17),  and  another  of  which  was  from 
a  partial  skeleton,  recovered  at  Arkalon  gravel 
pit  in  Kansas,  which  did  not  include  cranial 
portions.  I  do  not  consider  that  this  sample, 
or  the  analysis  based  thereon,  can  substan- 
tially support  a  specific  division  of  the  coy- 
ote line  at  the  Irvingtonian-Rancholabrean 
boundary,  and  I  continue  to  think  that  this 
line  was  represented  in  the  Irvingtonian  only 
by  the  species  C.  latrans. 

Kurten  (1974:12,  27)  also  stated  that  the 
species  Canis  arnensis  of  the  Villafranchian 
of  Europe  closely  resembled  the  North  Amer- 
ican material  he  referred  to  C.  priscolatrans, 
but  that  the  former  averaged  smaller.  He 
observed  that  future  studies  might  demon- 
strate that  these  two  species  formed  part  of 
a  single  Holarctic  coyote  population,  in  which 
case  C.  arnensis  would  have  to  be  regarded  as 
a  synonym  or  subspecies  of  C.  priscolatrans. 
I  would  not  agree  with  the  use  of  the  name 
C.  priscolatrans  for  such  a  population,  but 
since  I  have  not  examined  any  specimens  of 
C.  arnensis  I  can  not  comment  on  the  question 
of  their  relationship  to  North  American  Canis. 

Fossil  record. — The  following  list  is  ar- 
ranged alphabetically  by  state  and  province, 
and  geographically  (north  to  south,  west  to 
east)  within  states,  except  that  Alberta  is 
listed  first  and  Mexican  states  last.  Speci- 
mens examined  by  me  are  identified  by  ele- 
ment, museum  number,  or  both;  and  selected 
measurements  are  found  in  appendix  B  (part 
10)  and  appendix  C  (part  2).  Occurrences 
also  are  shown  on  the  map  in  figure  46. 

ALBERTA. — Medicine  Hat;  Kansan,  Sangamon, 
Wisconsin;  as  C.  cf.  latrans  (Churcher,  1969b:180; 
Kurten,  1974:7,  10). 

ALASKA. — Cripple  Creek  Mine,  near  Fairbanks; 
Wisconsin;  as  C.   latrans   (Guthrie,   1968:352). 

ARIZONA. — Anita,  Coconino  County;  early  Irv- 
ingtonian ( Richard  H.  Tedford,  American  Museum 
of  Natural  History,  pers.  comm.);  as  C.  latrans  (Hay, 
1921:633);  CI,  USNM. 

Ventana  Cave,  Papago  Indian  Reservation,  Pima 
County;  Wisconsin  (Hibbard,  1958);  as  C.  latrans 
(Colbert,    1950:132). 

Papago  Springs  Cave,  southeast  of  Sonoita,  Santa 
Cruz  County;  Wisconsin  (Hibbard,  1958);  as  C. 
caneloensis  (Skinner,   1942:163),  as  C.  latrans  cane- 


76 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


Fig.  46. — Map  showing  localities   (black  dots)    of  fossil  C.   latrans.    Because  of  the  scale  of  the  map,  it 
was  not  possible  to  plot  all  localities  in  crowded  areas. 


loensis  (Slaughter,  1966b:480),  as  C.  latrans  (An- 
derson, 1968:22);  skull  without  mandibles,  AMNH 
42800.  Skinner  originally  distinguished  caneloensis 
as  a  species,  on  the  basis  of  its  following  characters, 
as  compared  to  the  living  C.  estor  ( =C.  latrans 
mearnsi ) :     proportionally   wider   face,    larger   bullae, 


wider  P4,  less  prominent  hypocone  on  Ml,  and 
wider  M2  with  less  developed  hypocone  and  proto- 
conule.  Now,  however,  C.  estor  and  all  other  named 
kinds  of  Recent  coyotes  have  been  arranged  as  sub- 
species of  C.  latrans  (Jackson,  1951),  and  it  seems 
unlikely    that    these    differences    presently    would    be 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


77 


considered  more  than  subspecific.  Slaughter  (1966b: 
480)  demonstrated  that  the  relative  facial  width  of 
caneloensis  fell  within  the  range  of  variation  of  Recent 
C.  latrans.  The  additional  characters  listed  by  Skin- 
ner are  matched  in  large  series  of  Recent  coyotes. 
The  interruption  of  the  ridge  extending  anteriorly 
from  the  hypocone  of  Ml  is  unusual,  but  does  occur 
rarely  in  modern  coyotes.  In  the  structure  of  the 
occipital,  braincase,  frontals,  and  sagittal  crest,  and 
in  all  other  features  that  can  be  evaluated,  the  skull 
is  not  different  from  typical  western  C.  latrans.  I 
thus  concur  with  Slaughter  in  reducing  caneloensis 
to  subspecific  rank. 

Murray  Springs,  1  mi.  W  Lewis  Spring  on  San 
Pedro  River,  Cochise  County;  late  Pleistocene;  man- 
dibular fragment,   UAriz  2406. 

Double  Adobe,  Cochise  County;  late  Wisconsin; 
as   C.   latrans    (Kurten,    1974:8). 

ARKANSAS.— Eddy  Bluff  shelter,  near  Spring- 
dale,  Washington  County;  early  Recent;  as  C.  latrans 
(Morrison,   1970). 

Conard  fissure,  15  mi.  S  Harrison,  Newton  County; 
Illinoian  (Kurten,  1963:100);  as  C.  latrans  (James 
H.  Quinn,  Department  of  Geology,  University  of 
Arkansas,  pers.  comm.). 

Peccary  Cave,  eastern  Newton  County;  early 
Recent;  as  "coyote"  (Quinn,  1972:93). 

CALIFORNIA.— Samwel  Cave,  Shasta  Lake, 
Shasta  County;  Wisconsin  ( Hibbard,  1958);  as  C. 
latrans   (Graham,   1959). 

Hawver  Cave,  5  mi.  E  Auburn,  El  Dorado  Coun- 
ty; Wisconsin  (Hibbard,  1958);  as  C.  ochropus 
(Stock,  1918:479),  as  C.  latrans  (Anderson,  1968: 
22);  cranial  fragment,  UCMP  11041. 

Teichart  gravel  pit,  Sacramento  County;  late 
Pleistocene;   mandibular  fragment,  UCMP  85379. 

Murphys,  Calaveras  County;  Pleistocene;  as  C. 
latrans  (Whitney,  1879:246),  as  "C.  latrans?"  ( Mer- 
riam,  1903:290). 

Irvington,  Alameda  County;  Irvingtonian;  as  C. 
irvingtonensis  (Savage,  1951:231);  two  mandibular 
fragments,  UCMP  38748,  38805;  cranial  fragment, 
UCMP  56090;  radius,  UCMP  38804.  According  to 
Savage,  irvingtonensis  has  a  relatively  deeper  hori- 
zontal ramus,  and  relatively  wider  and  more  closely 
spaced  premolars  than  C.  latrans.  He  compared  the 
two  mandibles  to  79  specimens  of  Recent  western 
coyotes,  and  to  30  jaws  of  Pleistocene  C.  latrans 
from  Rancho  La  Brea.  He  listed  eleven  dental 
measurements  of  irvingtonensis,  but  none  on  the 
comparative  material,  and  also  none  to  demonstrate 
the  proportional  depth  of  the  Irvington  jaws.  Al- 
though I  agree  that  these  mandibles  have  a  more 
massive  appearance  than  those  of  most  other  coy- 
otes, I  find  all  measurements  to  fall  within  the 
range  of  variation  of  Recent  and  Pleistocene  C. 
latrans  from  western  North  America.  The  relative 
depth  of  the  horizontal  ramus  and  width  of  the  teeth 
also  overlap  the  corresponding  dimensions  in  C. 
latrans  (Figs.  44,  47).  The  premolars  are  unusually 
close  together,  but  their  spacing  varies  in  Cams.  In 
development  of  the  posterior  cusps  and  cingulum 
on  p4,  and  in  other  features  that  can  be  evaluated, 


10 
9 
8h 


20    21     22    23    24    25  26 

Fig.  47. — Scatter  diagram  comparing  crown 
length  of  ml  (horizontal  axis)  and  crown  width  of 
ml  (vertical  axis),  in  40  specimens  of  Pleistocene 
C.  latrans  orcutti  from  Rancho  La  Brea  (range  of 
variation  indicated  by  solid  line)  and  two  specimens 
of  C.  latrans  irvingtonensis   (v). 

the  Irvington  material  matches  series  of  C.  latrans. 
Consequently  I  do  not  consider  this  material  to  rep- 
resent a  separate  species.  Savage's  name  may  be 
tentatively  maintained  as  a  subspecific  designation, 
C.  latrans  irvingtonensis,  until  sufficient  material  is 
available  to  more  fully  evaluate  the  situation. 

Tranquility,  Fresno  County;  late  Wisconsin;  as 
C.  latrans  (Kurten,   1974:8). 

McKittrick  tar  seeps,  Kern  County;  Wisconsin; 
as  C.  latrans  orcutti  (Schultz,  1938b:  165;  Giles, 
1960:385);  27  skulls  and  cranial  fragments,  16  man- 
dibles and  fragments  thereof,  LACM.  Although  he 
recognized  considerable  variation  in  cranial  and  den- 
tal characters,  Schultz  referred  all  McKittrick  coyotes 
to  the  subspecies  orcutti.  He  observed  that  some  of 
the  skulls  were  characterized  by  comparatively  more 
massive  dentition,  larger  size,  and  a  broader  muzzle 
than  is  to  be  seen  in  the  living  California  subspecies, 
C.  latrans  ochropus. 

Maricopa  Brea,  near  Maricopa,  Kern  County; 
Wisconsin;  16  skulls  and  cranial  fragments,  23  man- 
dibles and  fragments  thereof,  various  isolated  teeth, 
LACM.  This  hitherto  unreported  collection  closely 
resembles  that  of  C.  latrans  orcutti  from  Rancho  La 
Brea  and   McKittrick. 

Carpinteria  asphalt,  Santa  Barbara  County;  Wis- 
consin (Hibbard,  1958);  as  "resembling  C.  latrans" 
(Wilson,  1933:68),  as  C.  latrans  (Anderson,  1968: 
22). 

Rancho  La  Brea,  Los  Angeles,  Los  Angeles 
County;  Wisconsin;  as  C.  ochropus  orcutti  (Mer- 
riam,  1912:255),  as  C.  latrans  orcutti  (Giles,  1960); 
60  skulls  and  50  mandibles,  LACM.  Compared  to 
those  of  Recent  coyotes,  Merriam  stated  that  skulls 
of  orcutti  averaged  larger,  and  had  broader  palates 
and  zygomata,  deeper  and  thicker  mandibles,  and 
thicker  carnassials.  Giles  (1960)  found  no  significant 
difference  between  samples  of  orcutti  from  Rancho 
La  Brea  and  McKittrick.  He  also  reported  that  the 
statistical  separation  among  the  Recent  subspecies, 
testes,  mearnsi,  and  ochropus,  was  considerably  less 
than  that  between  orcutti  and  any  one  of  them.  My 
own  findings  concur  with  those  of  Merriam  and 
Giles.  The  skulls  of  orcutti  average  larger  in  all  meas- 
urable  dimensions   than   those   of   Recent   subspecies, 


78 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


04 


ORCUTTI 


I 
2 

3 

4 

5 

6 

7 

8 

9 

10 

II 

12 

13 

14 

15 


..00       1.10 


Fig.  48. — Ratio  diagram  comparing  means  of  C. 
lot  ram  orcutti  from  Rancho  La  Brea  and  the  total 
series  of  Recent  C.  latrans  from  northern  and  western 
North  America  (vertical  line).  Vertically  arranged 
numbers  represent  the  measurements  so  numbered 
in  appendix  B.  A  log  difference  scale  is  provided 
above,  and  a  ratio  scale  below  the  diagram.  The 
Recent  series  consisted  of  the  combined  male  and 
female  samples  (n=277).  Sample  sizes  for  the  vari- 
ous measurements  of  Rancho  La  Brea  coyotes  were 
as  indicated  in  appendix  B   (part  10). 

and  several  individuals  from  Rancho  La  Brea  are 
larger  than  the  largest  Recent  specimens  that  I 
examined.  Comparisons  of  proportion  are  provided 
in  figures  44,  48,  and  49.  In  addition,  orcutti  differs 
from  Recent  C.  latrans  in  usually  having  more  convex 
frontals,  a  more  prominent  sagittal  crest  (but  not 
matching  that  of  C.  lepophagus),  a  broader  supra- 
occipital  shield,  and  temporal  ridges  that  join  an- 
terior, rather  than  posterior,  to  the  coronal  suture. 
In  size  and  all  other  characters  there  is  overlap  with 
series  of  living  coyotes,  and  I  agree  with  earlier 
authors  in  recognizing  orcutti  as  a  local  late  Pleisto- 
cene population  of  C.  latrans.  The  fact  that  the 
coyotes  in  this  group  were  large  may  be  correlated 
with  the  sympatric  presence  of  a  much  larger  kind 
of  wolf  (C.  dims)  than  the  kind  (C.  lupus)  that 
shared  the  historical  range  of  C.  latrans.  Hence  the 
morphology  and  ecological  niche  of  the  late  Pleisto- 
cene coyotes  may  have  been  shifted  toward  those  of 
wolves.  Merriam  (1912:258)  noted  that  a  few 
cranial  fragments  from  Rancho  La  Brea  suggested 
the  presence  of  a  smaller  coyote,  more  like  that  found 


presently  in  the  area,  in  addition  to  orcutti.  Schultz 
( 1938b:  164- 168)  also  considered  that  two  subspecies 
might  be  represented  in  the  McKittrick  deposits.  But 
each  of  these  authors  decided  that  it  would  be  best 
to  refer  all  specimens  of  C.  latrans  from  the  respec- 
tive sites  to  a  single  subspecies.  I  found  one  adult 
skull  from  Rancho  La  Brea,  which  was  strikingly 
smaller  than  the  others,  but  considering  the  variation 
that  can  be  expected  in  a  local  population  of  Recent 
Canis,  there  is  no  valid  reason  why  this  single  skull 
should  be  taxonomically  distinguished  from  C.  /. 
orcutti.  And  we  would  expect  variation  in  the  tar 
pit  fauna  to  be  especially  great  since  the  deposits 
were  formed  over  thousands  of  years.  Both  in  state 
of  preservation  and  in  abundance,  the  collections 
from  Rancho  La  Brea  contain  the  most  useful  series 
of  Pleistocene  C.  latrans.  The  number  of  individual 
animals  represented  in  the  Los  Angeles  County  Mu- 
seum was  counted  at  200  by  Stock  (1929)  and  at 
239  by  Marcus    (1960). 

Rancho  La  Brea,  Los  Angeles,  Los  Angeles 
County;  Wisconsin;  as  C.  andersoni  (Merriam,  1910: 
393;  1912:260);  skull  without  mandibles,  UCMP 
12249.  Merriam  specifically  distinguished  this  speci- 
men from  C.  latrans  primarily  because  of  its  rela- 
tively short  and  broad  rostrum.  lackson  (1951:232) 
thought  it  possible  that  additional  material  might 
show  andersoni  to  be  synonymous  with  the  living  C. 
latrans  clcpticus  of  Baja  and  southern  California. 
Giles  (1960)  also  questioned  the  specific  status  of 
andersoni,  and  found  statistical  similarity  between 
andersoni  and  four  skulls  of  clepticus  from  San  Diego 
County,  California.  Slaughter  ( 1966a:479-48I ) 
found  the  rostral  proportions  of  andersoni  to  be  com- 
parable to  those  of  a  Pleistocene  skull  from  the  Lau- 
bach  Cave,  Williamson  County,  Texas,  which  he 
thought  might  be  referable  to  C.  latrans  harriscrooki. 
Anderson  (1968:24)  suggested  that  andersoni  was 
probably  a  subspecies  of  C.  latrans.  The  name  C. 
andersoni  here  is  synonymized  under  C.  latrans,  and 
is  referred  to  the  Rancho  La  Brea  population  of  C.  /. 
orctitti.  The  single  skull  upon  which  Merriam  based 
his  description,  is  from  a  juvenile,  perhaps  five 
months  old  at  time  of  death.  Although  the  rostrum 
is  indeed  relatively  broader  than  in  nearly  all  other 
available  specimens  of  C.  latrans  (Fig.  49),  this 
condition  may  be  accounted  for  in  part  by  age  or  by 
retarded  development  lengthwise.  There  are  several 
other  visible  abnormalities  that  might  contribute  to 
the  aberrant  appearance  of  the  skull.  The  alveolus 
for  the  left  PI  is  missing,  and  the  alveolus  for  the 
right  PI  crowds  that  for  the  right  P2.  The  whole 
skull  seems  to  be  slightly  twisted  out  of  line,  and 
there  is  an  unusual  inflation  of  the  dorsal  surface  of 
the  rostrum  immediately  anterior  to  the  orbits.  The 
characters  of  the  occipital,  braincase,  frontals,  and 
sagittal  crest  are  those  of  normal  C.  latrans,  and 
confirm  that  the  specimen  is  a  coyote,  not  a  "coyote- 
like wolf"  as  stated  by  Merriam   (1912:260). 

Harbor  freeway,  Los  Angeles,  Los  Angeles 
County;  Wisconsin;  as  C.  cf.  latrans  (Miller,  1971: 
54). 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


79 


68 
66 
64 
62 
60 
58 
56 
54 
52 
50^ 


O      O 

O    o 
ho  o 

OOo  • 


o< 


>o 


ad    • 


'  c    •  o  •  •  •  • 


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62    64    66  68    70  72   74   76  78 

Fig.  49. — Scatter  diagram  comparing  alveolar 
length  of  maxillary  toothrow  (horizontal  axis)  to 
maximum  crown  width  across  upper  cheek  teeth 
(vertical  axis)  in  certain  specimens  of  C.  latrans 
(see  appendix  B  for  full  description  of  measure- 
ments). Black  dots,  Recent  C.  latrans  lestes;  open 
circles,  Pleistocene  C.  latrans  orcutti  from  Rancho 
La  Brea;  c,  C.  latrans  canelocnsis;  d,  C.  latrans  hon- 
durensis;  h,  C.  latrans  harriscrooki;  a,  C.  andersoni 
(  =  C.  latrans  orcutti). 

La  Mirada,  Los  Angeles  County;  Wisconsin;  as 
C.   cf.   latrans   (Miller,   1971:49). 

Costeau  pit,  2  mi.  S  El  Toro,  Orange  County; 
Wisconsin;  as  C.  cf.  latrans  (Miller,  1971:17)-  Ml 
LACM  1S220. 

Vallecito  Creek,  San  Diego  County;  Irvingtonian 
(Hibbard,  et  al,  1965);  six  mandibular,  fragments, 
various  postcranial  fragments,  LACM. 

COLORADO. — Chimney  Rock  animal  trap,  Lari- 
mer County;  late  Pleistocene  or  early  Recent;  as  C. 
latrans   ( Hager,   1972:65). 

FLORIDA.— Ichetucknee  River,  Columbia  Coun- 
ty; Wisconsin;  as  C.  latrans  (Webb,  1974b:  17).  Ac- 
cording to  Kurten  (1974:10)  a  left  mandible  (UF 
1151)  from  this  site  represents  a  "medium  to  large" 
C.  latrans.  Martin  and  Webb  (1974:128),  however, 
stated  that  the  same  specimen  "belongs  to  a  smaller 
individual  of  the  same  general  character"  as  material 
from  Devil's  Den,  which  they  referred  to  C.  familiaris 
(see  below).  In  my  own  opinion  UF  1151  represents 
C.  familiaris,  but  Webb  (1974b:  17)  may  have  based 
his  report  of  C.  latrans  on  other  material.  In  my 
dissertation  ( Nowak,  1973:204)  I  associated  another 
mandible  ( UF  11517)  with  the  Ichetucknee  River 
site,  but  that  specimen  actually  is  from  Devil's  Den. 


Haile  XIIB,  Alachua  County;  Rancholabrean; 
mandibular  fragment,  UF. 

Devil's  Den,  near  Williston,  Levy  County;  late 
Wisconsin  or  early  Recent  (7,000-8,000  B.P.);  three 
mandibular  fragments,  UF  11514,  11515,  11517. 
Subsequent  to  my  assignment  of  these  specimens  to 
C.  latrans  (Nowak,  1973:204-205),  they,  along  with 
additional  material  from  the  site,  were  referred  to 
C.  familiaris  by  Martin  and  Webb  (1974:127-128). 
The  suggestion  by  these  same  authors,  that  canid 
specimens  from  the  Ichetucknee  River,  Seminole 
Field,  and  Melbourne  localities  represent  C.  famili- 
aris, rather  than  C.  latrans,  has  raised  questions 
about  the  over-all  status  of  the  coyote  line  in  the 
Wisconsin   and   early   Recent   of   Florida. 

Reddick  IA,  Marion  County;  Sangamon    (Webb, 
1974b:13);  as  C.  latrans  (Gut  and  Ray,  1964:325)! 
Withlacoochee  River  VIIA,  Citrus  County;  Sanga- 
mon; as  C.  latrans  (Webb,  1974b:  13,  17).  ' 

Seminole  Field,  near  St.  Petersburg,  Pinellas 
County;  Wisconsin  (Hibbard,  et  al,  1965);  as  C. 
cf.  riviveronis  (Simpson,  1929a:573),  as  C.  familiaris 
(Martin  and  Webb,   1974:128). 

Lake  Cutaline,  Pinellas  County;  late  Pleistocene; 
mandibular   fragment,   UF. 

Phillipi  Creek-Fruitville  Ditch,  7  mi.  E  Sarasota, 
Sarasota  County;  Wisconsin;  as  C.  cf.  riviveronis 
(Simpson,  1929b:  275). 

Melbourne,  Brevard  County;  Wisconsin  (Hibbard, 
et  al,  1965);  as  C.  riviveronis  (Gazin,  1950:12; 
Simpson,  1929b:268),  as  C.  cf.  latrans  (Ray,  1958: 
433),  as  C.  familiaris  (Martin  and  Webb,  1974: 
128);  rostral  fragment,  MCZ  5909;  mandibular  frag- 
ment, isolated  teeth,  USNM. 

Vero  (stratum  3),  Indian  River  County;  late 
Wisconsin  (Webb,  1974b:13);  as  C.  cf.  latrans 
(Sellards,  1916:157),  as  C.  riviveronis  (Hay,  1917a: 
59),  as  C.  latrans  (Weigel,  1962:38);  maxilla  with 
P4,  FGS  7036.  In  his  description  of  C.  riviveronis, 
Hay  wrote  that  the  specimen  differed  from  those  of 
C.  latrans  in  having  a  relatively  shorter  anterior  lobe 
of  P4,  and  a  relatively  greater  transverse  extent  of 
the  sockets  of  Ml  and  M2.  The  measurements  he 
provided,  however,  were  only  slightly  different  from 
those  of  four  comparative  specimens  of  C.  latrans. 
Ray  (1958:433)  considered  that  there  was  no  basis 
upon  which  C.  riviveronis  could  be  distinguished 
from  C.  latrans.  I  agree,  but  the  Vero  specimen  and 
others  of  C.  latrans  from  Florida  average  smaller 
than  western  coyotes,  possibly  because  of  the  sym- 
patic presence  of  a  small  wolf   (C.   rufus). 

IDAHO. — Jaguar  Cave,  Beaverhead  Mountains, 
Lemhi  County;  late  Wisconsin  (C-14  dates:  10,370 
±350  and  11,580±250  B.P.);  as  C.  latrans  (Kurten 
and  Anderson,  1972:24). 

Moonshiner  Cave,  Bingham  County;  late  Wiscon- 
sin or  early  Recent;  as  C.  latrans  (Kurten  and  An- 
derson,   1972:37). 

Middle  Butte  Cave,  Bingham  County;  early  Re- 
cent; as  C.   latrans   (Kurten,   1974:9). 

American  Falls,  Power  County;  Rancholabrean 
(Hibbard,  et  al,   1965),  Illinoian   (Kurten,   1974:7); 


80 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


as  C.  latrans  (Kurten,  1974:7).  According  to  Kur- 
ten,  two  mandibles  from  this  locality  are  "large." 
With  respect  to  material  from  the  same  site,  Gazin 
(1935:298)  stated:  "An  incomplete  humerus,  a  tibia 
and  a  third  metatarsal  are  recognized  as  belonging 
to  a  dog  somewhat  smaller  than  Canis  occidentalis 
but  larger  than  a  coyote."  The  possibility  that  all 
of  this  material  represents  the  lineage  of  C.  rujus, 
should  not  be  overlooked. 

Rainbow  Beach  local  fauna,  American  Falls  Res- 
ervoir, Power  County;  Wisconsin  (C-14  dates: 
21,500±700  and  31,300±2,300  B.P.);  as  C.  latrans 
(McDonald   and   Anderson,    1975:26). 

Twin  Falls,  Twin  Falls  County;  Wisconsin;  as 
C.  latrans  (Kurten,  1974:9). 

ILLINOIS.— Galena,  Jo  Daviess  County;  Wis- 
consin (Kurten,  1974:10);  as  C.  latrans  (Hay,  1923: 
337). 

Polecat  Creek  gravel  pits,  1  mi.  S  Ashmore,  Coles 
County;  late  Wisconsin  ( Hibbard,  et  ah,  1965);  as 
C.  latrans  (Galbreath,  1938:306,  311). 

INDIANA. — Boone  County;  Wisconsin  (Kurten, 
1974:10);  as  C.  latrans  (Cope  and  Wortman,  1884: 
7;  Lyon,  1936:150). 

IOWA. — Dubuque,  Dubuque  County;  Wiscon- 
sin; as  C.  latrans  (Kurten,   1974:9). 

KANSAS. — Borchers  local  fauna,  sec.  21,  T33S 
R28W,  Meade  County;  Yarmouthian;  as  C.  cf.  la- 
trans (Getz,  1960:363). 

Adams  local  fauna,  north  of  Cimarron  River, 
Meade  County;  early  Illinoian  (Hibbard,  1970);  as 
C.   latrans   ( Schultz,   1969:30). 

Butler  Spring  local  fauna,  15  mi.  SSW  Meade, 
Meade  County;  late  Illinoian;  as  C.  cf.  latrans  (Hib- 
bard and  Taylor,   1960:178). 

Cragin  Quarry  local  fauna,  north  of  Cimarron 
River,  Meade  County;  Sangamon;  as  C.  cf.  latrans 
(Hibbard   and   Taylor,    1960:178). 

MARYLAND.— Cumberland  Cave,  4  mi.  NW 
Cumberland,  Allegany  County;  Illinoian  ( Kurten, 
1963:100);  as  C.  cf.  priscolatrans  (Gidley  and 
Gazin,  1938:23);  cranial  fragment,  USNM  7660. 
The  specimen  is  in  poor  condition,  and  represents 
an  animal  that  may  not  have  been  a  year  old.  The 
skull  is  the  size  of  that  of  a  large  Recent  C.  latrans, 
and  the  braincase  and  postorbital  constriction  are 
coyotelike  in  shape.  The  frontals  are  more  convex 
than  in  most  Recent  C.  latrans. 

MISSISSIPPI.— Vicksburg  (south  of),  Warren 
County;  "Wisconsin?";  as  C.  latrans  (Kurten,  1974: 
10). 

MISSOURI.— Brynjulfson  Caves,  6  mi.  SSE  Co- 
lumbia, Boone  County;  late  Wisconsin  (about  10,000 
B.P.);  as  C.  latrans  (Parmalee  and  Oesch,  1972:29). 

Younger's  Cave,  St.  Clair  County;  early  Recent; 
as  C.  latrans  (Kurten,  1974:10);  two  mandibles,  KU 
5952,  7072. 

Herculaneum  (near),  Jefferson  County;  Wiscon- 
sin (Hibbard,  et  ah,  1965);  as  C.  latrans  (Olson, 
1940:42). 

Bat  Cave,  8  km.  NW  Waynesville,  Pulaski  Coun- 


ty; late  Wisconsin  (10,000-16,000  B.P.);  as  C.  la- 
trans   (Hawksley,   Reynolds,  and  Foley,   1973:77). 

Zoo  Cave,  1  mi.  ENE  Hilda,  Taney  County;  early 
Recent  (less  than  9,000  B.P.);  as  C.  latrans  (Hood 
and  Hawksley,   1975:24;   Saunders,   1977:14). 

NEBRASKA. — Hay  Springs  quarry,  Sheridan 
County;  probably  Illinoian  (Hibbard,  1958);  as  C. 
cf.  latrans  (Matthew,  1902;  Schultz,  1934:369). 

Rushville  fossil  quarry,  Sheridan  County;  Yar- 
mouthian (Schultz  and  Martin,  1970);  as  "Canis 
sp. — Coyote"  (Schultz  and  Tanner,  1957:71);  man- 
dibular fragment,  UN  2913. 

Mullen,  Cherry  County;  late  Irvingtonian  (Kur- 
ten, 1974:7);  as  C.  latrans  (Martin,  1972:174); 
mandible,   UN  26115. 

Schmidt  gravel  pit,  West  Point,  Cuming  County; 
late  Pleistocene;   skull  without  mandibles,  UN   2909. 

Allen  site,  Frontier  County;  late  Wisconsin;  as 
C.  latrans  (Schultz,  Martin,  and  Tanner,   1970:120). 

Angus  fossil  quarry,  Nuckolls  County;  Yar- 
mouthian; as  C.  latrans  (Schultz  and  Tanner,  1957: 
67). 

NEVADA.— Tule  Springs,  Clark  County;  late 
Wisconsin;  as  C.  latrans  (Mawby,  1967). 

NEW  MEXICO.— Isleta  Caves,  8  mi.  W  Isleta, 
Bernalillo  County;  late  Wisconsin;  as  C.  latrans 
(Harris  and  Findley,   1964:115). 

Blackwater  Draw,  near  Clovis,  Curry  County; 
Wisconsin  (Lundelius,  1967:301);  CI,  Ml,  TM  937- 
896,  937-897. 

Shelter  Cave,  near  Las  Cruces,  Dona  Ana  County; 
late   Pleistocene;   skull   without   mandibles,   LACM. 

Conkling  Cavern,  near  Las  Cruces,  Dona  Ana 
County;  late  Pleistocene;  skull,  LACM  1634;  man- 
dible, LACM  1631. 

Burnet  Cave,  50  mi.  W  Carlsbad,  Eddy  County; 
Wisconsin  (Hibbard,  1958);  as  C.  latrans  lestes  and 
C.  microdon  (Schultz  and  Howard,  1935:284);  man- 
dibular fragment,  CI,  P4,  UN   13454,   13455. 

Dark  Canyon  Cave,  Eddy  County;  late  Pleisto- 
cene; mandible,  LACM. 

Dry  Cave,  15  mi.  W  Carlsbad,  Eddy  County;  Wis- 
consin (Kurten,  1974:8);  as  C.  latrans  (Harris,  1970: 
14). 

OKLAHOMA. — Berends  local  fauna,  near  Gate, 
Beaver  County;  probably  Illinoian  ( Hibbard  and 
Taylor,  1960:57);  as  "a  canid  the  size  of  a  coyote" 
(Rinker  and  Hibbard,  1952:101),  as  C.  latrans 
(Starrett,  1956:1187);  mandibular  fragment,  UMMP 
33319. 

Alton,  Ottawa  County;  Wisconsin  (Kurten,  1974: 
9);  as  C.  latrans  (Hay,  1920:129);  axis,  USNM 
9131. 

OREGON. — Fossil  Lake,  Lake  County;  early  or 
middle  Wisconsin  (Allison,  1966:32);  as  C.  lestes 
(Elftman,  1931:7);  three  mandibular  fragments, 
AMNH  8584,  8585,  8586. 

PENNSYLVANIA.— Frankstown  Cave,  Blair 
County;  Wisconsin  (Hibbard,  1958);  as  C.  priscola- 
trans (Peterson,  1926:283);  two  mandibular  frag- 
ments (from  same  individual),  CI,  CM  11027. 
Peterson  thought  that  the  material  indicated  an  ani- 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


81 


mal  about  the  same  size  as  that  represented  by  the 
type  of  C.  priscolatrans  Cope,  1899  (which  in  this 
paper  is  referred  to  a  subspecies  of  C.  rufus).  The 
mandibles  and  teeth  are  large,  but  fall  within  the 
size  range  of  Recent  and  Pleistocene  C.  latrans  from 
western  North  America.  As  noted  by  Peterson,  there 
is  a  prominent  posterior  cusp  on  the  p2  of  each 
mandible.  In  this  respect,  the  Frankstown  mandibles 
resemble  those  of  C.  latrans  harriscrooki  Slaughter, 
1961   from  the  late  Pleistocene  of  Texas. 

TEXAS. — Rock  Creek,  Briscoe  County;  Kansan 
(Hibbard,  1970);  as  C.  priscolatrans  (Troxell,  1915: 
628,  634).  The  description  and  illustration  provided 
by  Troxell  indicate  that  an  Ml  from  this  site  is  best 
referred  to  C.   latrans,  not  C.  rufus  priscolatrans. 

Quitaque  local  fauna,  Motley  County;  Wisconsin 
(Kurten,  1974:9);  as  C.  latrans  ( Dalquest,  1964: 
501). 

Howard  Ranch  local  fauna,  Groesbeck  Creek, 
northwest  of  Quanah,  Hardeman  County;  Wisconsin 
(C-14  date:  16,775+565  B.P.);  as  C.  latrans  (Dal- 
quest, 1965:71). 

Wichita  Falls,  Wichita  County;  early  Recent;  as 
C.   latrans    (Dalquest,    1961:75). 

Lubbock  Reservoir,  Lubbock  County;  Wisconsin 
(Lundelius,  1967:302);  as  C.  latrans  (Kurten,  1974: 
9);   mandibular   fragment,   TM. 

Slaton  quarry,  5  mi.  N  Slaton,  Lubbock  County; 
Sangamon  (Hibbard,  1970);  as  near  C.  latrans  (Dal- 
quest, 1967:9). 

Gilliland  local  fauna,  Knox  County;  Irvingtonian 
(late  Kansan:  Hibbard,  et  al.,  1965);  as  C.  cf. 
latrans    (Hibbard   and   Dalquest,    1966:20). 

Benjamin  Franklin  local  fauna,  North  Sulphur 
River,  Delta  County;  late  Wisconsin  (11,000  B.P.); 
as  C.  latrans  (Slaughter  and  Hoover,  1963:141). 
Two  mandibular  fragments  were  reported  not  to 
have  a  posterior  cusp  on  p2,  as  typical  of  C.  latrans 
harriscrooki  ( see  following  accounts ) .  The  material 
was  thought  to  represent  a  more  modern  coyote  that 
was  either  a  replacement  for  or  a  descendent  of  the 
older  harriscrooki. 

Clear  Creek  local  fauna,  north  of  Denton,  Den- 
ton County;  Wisconsin  (C-14  date:  28,840±4,740 
B.P.;  Hibbard,  et  al,  1965);  as  C.  latrans  cf.  harris- 
crooki  (Slaughter  and  Ritchie,  1963:125). 

Lewisville  site,  Denton  County;  Sangamon  or 
Wisconsin  interstadial;  as  C.  latrans  harriscrooki 
(Slaughter,  1961);  cast  of  type,  mandible,  SMUMP 
60315.  This  subspecies  originally  was  called  "wolf 
like,"  and  was  distinguished  from  other  coyotes  by 
its  well  developed  posterior  cusp  on  p2,  longer  tooth- 
row  relative  to  depth  of  mandible,  reduced  distance 
between  premolars,  and  more  vertical  ascending 
ramus.  Slaughter  examined  52  mandibles  of  Recent 
C.  latrans  from  the  United  States,  and  found  none 
with  a  posterior  cusp  on  p2.  He  said,  however,  that 
this  cusp  was  present  on  two  specimens  from  San 
Luis  Potosi,  Mexico,  and  on  one  from  Archaga,  Hon- 
duras. The  same  condition  existed  in  late  Pleistocene 
specimens  from  Brazos  County  and  Ingleside,  Texas, 
which    Slaughter    referred    to    harriscrooki.     Another 


mandible,  collected  at  a  Pleistocene  site  in  Hender- 
son County,  Texas,  also  was  assigned  to  this  sub- 
species, on  the  basis  of  its  more  vertical  ascending 
ramus.  Slaughter  speculated  that  harriscrooki  might 
be  a  southern  kind  of  coyote  that  could  have  in- 
habited Texas  only  in  an  interglacial  or  interstadial. 
Possible  affinity  to  Recent  C.  I.  hondurensis  of  Hon- 
duras was  implied,  and  a  jaw  of  that  subspecies 
reportedly  had  the  angle  of  ascending  ramus  about 
the  same  as  in  harriscrooki.  I  examined  six  speci- 
mens of  C.  I.  hondurensis  ( 1  in  AMNH,  1  in  KU, 
2  in  MCZ,  2  in  USNM),  of  which  five  had  a  promi- 
nent posterior  cusp  on  p2,  and  one  had  the  cusp 
slightly  developed.  In  contrast,  only  six  of  250 
Recent  specimens  from  the  western  United  States, 
and  only  one  of  40  Pleistocene  specimens  from 
Rancho  La  Brea,  which  could  be  checked  for  this 
character,  had  any  trace  of  the  cusp.  Hondurensis 
originally  was  reported  to  have  a  relatively  broad 
palate,  and  I  found  such  a  condition  to  exist  in  most 
skulls  of  that  subspecies  that  I  examined  (Fig.  49). 
Slaughter  (1966b)  reported  that  a  Pleistocene  skull 
from  Laubach  Cave,  Williamson  County,  Texas, 
which  he  said  might  be  referable  to  harriscrooki, 
had  a  relatively  broader  palate  than  Recent  C.  la- 
trans. Therefore,  it  seems  a  reasonable  hypothesis 
that  a  warmth-adapted  coyote,  with  certain  more 
wolflike  characters  than  typical  Recent  C.  latrans, 
was  found  in  Texas  during  part  of  the  Pleistocene, 
and  might  still  be  represented  by  the  living  coyote 
of  Honduras. 

Moore  Pit  local  fauna,  Dallas,  Dallas  County; 
Sangamon;  as  C.  latrans  cf.  harriscrooki  (Slaughter, 
1966a:481;  1966b:79). 

Trinity  River  terraces,  2.5  mi.  NW  Trinidad, 
Henderson  County;  Wisconsin;  as  "Canis  sp. — Coy- 
ote" (Stovall  and  McAnulty,  1950:233),  as  C.  latrans 
harriscrooki   (Slaughter,   1961:509). 

Clamp  Cave,  San  Saba  County;  early  Recent;  as 
C.  latrans  (Lundelius,  1967:293). 

Carson  Holloway  Ranch,  San  Saba  County;  Wis- 
consin; as  C.   latrans   (Kurten,   1974:9). 

Miller's  Cave  (Travertine  unit),  Llano  County; 
early   Recent;    as   C.    latrans    (Lundelius,    1967:293). 

Longhorn  Cavern,  8.5  mi.  S  Burnet,  Burnet 
County;  late  Wisconsin  (Hibbard,  et  al,  1965);  as 
Canis  (Semken,  1961),  as  C.  latrans  (Lundelius, 
1967:293). 

Laubach  Cave,  Georgetown,  Williamson  County; 
Wisconsin  (Kurten,  1974:9);  as  C.  latrans,  possibly 
C.  /.  harriscrooki  (Slaughter,  1966a:479-481 );  skull 
without  mandibles,  SMUMP  61269.  As  Slaughter 
reported,  the  specimen  has  a  relatively  broad  facial 
width  as  compared  to  most  C.  latrans.  Some  Recent 
and  Pleistocene  specimens,  however,  especially  those 
assigned  to  C.  latrans  hondurensis,  approach  the 
Laubach  Cave  specimen  in  relative  broadness  (Fig. 
49).  Nonetheless,  assignment  to  harriscrooki  seems 
a  reasonable  procedure  (see  account  of  Lewisville 
site,  above ) . 

Cage  gravel  pit,  5  mi.  N  Cameron,  Milam  Coun- 


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MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


ty;  late  Pleistocene;  "closely  related  to  C.  latrans" 
(Hay,    1927:291). 

Brazos  County;  late  Pleistocene;  as  "a  coyote, 
Canis  sp."  (Peterson,  1946:166),  as  C.  latrans  harris- 
crooki  (Slaughter,  1961:509). 

Levi  shelter,  Travis  County;  early  Recent;  as  C. 
latrans  ( Lundelius,   1967:293). 

Schulze  Cave,  28  mi.  NE  Rock  Springs,  Edwards 
County;  Wisconsin  or  early  Recent;  "probably  re- 
ferable to  C.  latrans  harriscrooki  or  .  .  .  intermediate 
between  that  extinct  race  and  the  modern  coyote" 
(Dalquest,  Roth,  and  Judd,   1969:255-257). 

Klein  Cave,  12  mi.  WSW  Mountain  Home,  Kerr 
County;  late  Wisconsin;  as  C.  latrans  (Roth,  1972: 
78). 

Cave  Without  a  Name,  Kendall  County;  late 
Wisconsin  (C-14  date:  10,900+190  B.P.);  as  C. 
latrans    (Lundelius,    1967:293). 

Wunderlich  site,  Comal  County;  early  Recent; 
as  C.   latrans   (Lundelius,   1967:293). 

Friesenhahn  Cave,  near  Bulverde,  Bexar  County; 
Wisconsin;  as  C.  latrans  (Hay,  1920:141;  Lundelius, 
1960:38);  four  mandibles,  TM  933-670,  933-1622, 
933-2454,  933-3398.  No  posterior  cusp  was  present 
on  the  p2  of  the  one  specimen  that  could  be  checked 
for  this  character. 

Ingleside  gravel  pit,  San  Patricio  County;  Wis- 
consin; as  C.  latrans  harriscrooki  (Slaughter,  1961: 
509);  as  C.  latrans  (Lundelius,  1972:20). 

UTAH. — Silver  Creek  local  fauna,  5  mi.  N  Park 
City,  Summit  County;  late  Sangamon  to  early  Wis- 
consin; as  "Canis?  latrans"  (Miller,  1976:401). 

WISCONSIN.— Blue  Mounds,  Dane  County  (or 
"Iowa  lead  region");  late  Pleistocene;  as  C.  latrans. 
O.  P.  Hay  apparently  wrote  of  this  same  material 
three  different  times,  stating  ( 1 )  that  it  was  prob- 
ably from  Iowa  (1914:491);  (2)  that  it  was  prob- 
ably not  from  Blue  Mounds,  but  from  another  crevice 
(1918:347);  and  (3)  that  it  was  found  at  Blue 
Mounds    (1923:341). 

WYOMING.— Little  Box  Elder  Cave,  west  of 
Douglas,  Converse  County;  Wisconsin;  as  C.  latrans 
(Anderson,  1968,  24). 

Bell  Cave,  Albany  County;  Wisconsin  to  early 
Recent;  as  C.  latrans  (Anderson,  1974:81). 

AGUASCALIENTES.— Cedazo  local  fauna,  near 
City  of  Aguascalientes;  early  Rancholabrean  (prob- 
ably  Illinoian);  as  C.  latrans  ( Mooser  and  Dalquest, 
1975:786). 

ESTADO  DE  MEXICO.— Tequixquiac  (near); 
late  Pleistocene;  as  "C.  cf.  ocropus"  (Furlong,  1925: 
139,  152),  as  "C.  ocrupus"  (Alvarez,  1965:27). 
Both  of  these  authors  apparently  were  referring  to 
C.  ochropus,  a  name  for  the  living  coyote  of  Cali- 
fornia, that  has  been  arranged  as  a  subspecies  of 
C.   latrans. 

NUEVO  LEON. — San  Josecito  Cave,  near  Aram- 
berri;  Wisconsin;  as  C.  latrans  ( Kurten,  1974:7);  6 
cranial  fragments  and  22  mandibles,  LACM.  The 
upper  cranial  elements  can  not  be  fully  evaluated 
because  of  their  poor  condition,  but  seem  not  to 
differ  from  typical  C.   latrans.    Measurements  of  the 


mandibles  approach  those  of  specimens  from  Rancho 
La  Brea.  None  of  the  mandibles  have  a  p2  with  a 
posterior  cusp.  Age  has  not  yet  been  reliably  deter- 
mined for  the  mammalian  fauna  of  this  site,  but 
Jakway  (1958:326)  suggested  that  it  was  approxi- 
mately as  old  as  the  fauna  of  Papago  Springs  Cave, 
Arizona  (Wisconsin),  and  older  than  that  of  Rancho 
La  Brea. 

OAXACA.— Monte  Flor  Cave,  2  km.  NE  Valle 
Nacional;  early  Recent;  as  C.  latrans  (Alvarez,  1963). 

PUEBLA. — Valsequillo,  near  Puebla;  late  Pleisto- 
cene; as  C.  latrans  (Kurten,  1967:173). 

Evolutionary  position. — The  species  C. 
latrans  apparently  arose  from  certain  popula- 
tions within  the  species  C.  lepophagus  in  the 
Blancan.  Subsequently,  there  appears  to  have 
been  relatively  little  change  in  the  coyote 
line,  at  least  with  respect  to  the  skull,  but 
some  Pleistocene  populations  became  larger 
and  more  massive  than  most,  if  not  all  Recent 
subspecies. 

Canis  edwardii  Gazin 
1942.    Canis  edwardii  Gazin,  Proc.  U.S.  Natl. 

Mus.,  92:499. 
1954.    Canis   lupus   baileyi,   Hoffmeister   and 

Goodpaster,     Illinois     Biol.     Monogr., 

24:34. 

Holotype. — Skull  with  mandibles;  no. 
12S62,  U.S.  Natl.  Mus.;  about  two  miles  north- 
east by  east  of  Curtis  Ranch  House,  San 
Pedro  Valley,   Cochise  County,  Arizona. 

Geological  distribution. — Late  Blancan  to 
early   Irvingtonian. 

Geographical  distribution. — Known  from 
Arizona,  Kansas,  Oregon,  Texas,  and  Chihua- 
hua. 

Description. — A  medium-sized  canid  re- 
sembling C.  rufus  in  most  observable  char- 
acters; skull  medium-sized  with  mostly  nar- 
row proportions;  rostrum  elongated  and 
narrow;  braincase  relatively  small  and  mod- 
erately inflated  dorsoposteriorly;  postorbital 
constriction  elongated,  broad  lateromedially; 
zygomata  slender,  not  deep;  frontals  moder- 
ately elevated  above  rostrum,  not  prominently 
convex;  sagittal  crest  prominent;  mandible 
long,  slender,  and  shallow,  with  ascending 
ramus  set  at  comparatively  high  angle  to  the 


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83 


vertical;  teeth  relatively  large  and  set  closely 
together  in  jaws;  upper  canines  prominent, 
thin  anteroposteriorly;  P4  with  moderately 
developed  deuterocone;  Ml  with  relatively 
large,  deeply  sculptured  medial  section,  and 
pronounced  buccal  cingulum;  p2  lacking  pos- 
terior cusp;  p3  usually  with  second  cusp;  p4 
usually  with  second  and  third  cusp,  and  pro- 
nounced posteromedial  cingulum  extending 
behind  third  cusp. 

Comparison  with  C.  lepophagus. — Larger 
and  relatively  broader  in  all  measurable  di- 
mensions; teeth  usually  with  less  trenchant 
cusps. 

Comparison  with  C.  rtifus. — Close  resem- 
blance in  size  and  proportions  of  skull;  post- 
orbital  constriction  of  braincase  relatively 
broader;  ascending  ramus  of  mandible  set  at 
more  acute  angle  to  the  vertical;  Ml  with 
broader  medial  section  and  more  prominent 
buccal  cingulum. 

Remarks. — Hoffmeister  and  Goodpaster 
(1954:34)  considered  the  name  C.  edwardii 
a  synonym  of  C.  lupus  baileyi,  the  small  gray 
wolf  found  in  southern  Arizona  in  historic 
time.  Actually,  this  Pleistocene  wolf  can  not 
be  referred  to  a  living  subspecies.  As  is  dis- 
cussed later  in  this  paper,  C.  lupus  seems 
not  even  to  have  entered  North  America 
until  the  Illinoian,  and  the  Curtis  Ranch 
fauna  is  early  Irvingtonian  in  age.  Although, 
as  explained  by  Hoffmeister  and  Goodpaster, 
there  is  little  difference  in  over-all  size  be- 
tween the  type  of  C.  edwardii  and  some  speci- 
mens of  baileyi,  most  measurable  dimensions 
of  C.  edwardii  are  much  smaller  than  the 
means  of  those  of  baileyi.  Moreover,  the  ros- 
trum of  C.  edivardii  is  narrower  and  the  man- 
dible shallower,  than  those  of  any  skull  of 
baileyi  examined  by  me.  The  Ml  of  C.  ed- 
wardii differs  greatly  from  that  of  any  modern 
C.  lupus,  in  having  a  pronounced  buccal 
cingulum  and  a  relatively  large,  deeply  sculp- 
tured medial  section.  In  these  and  other  fea- 
tures, the  skull  of  C.  edwardii  approaches 
that  of  C.  rufus,  and,  as  realized  by  Gazin, 
that  species  is  the  only  one  to  which  the  fossil 


need  be  critically  compared.  Indeed,  the  red 
wolf  probably  is  a  direct  descendent  or  im- 
mediate relative  of  C.  edwardii,  and  the  latter 
eventually  may  be  shown  to  be  only  a  syno- 
nym or  subspecies  of  C.  rufus. 

In  my  dissertation  (Nowak,  1973:208,  229- 
230),  I  had  indicated  that  C.  edwardii  was 
known  only  from  the  type  locality,  but  that 
several  other  early  wolf  specimens  might  be 
referable  to  this  species,  rather  than  to  C. 
rufus.  Additional  material,  subsequently 
made  available  to  me  through  the  kindness  of 
Richard  H.  Tedford  and  Beryl  E.  Taylor  at 
the  American  Museum  of  Natural  History, 
suggests  a  greater  range  for  C.  edwardii  and 
also  provides  a  basis  for  assigning  some  pre- 
viously examined  specimens  to  this  species. 
Of  particular  importance  in  this  regard  is  a 
skull  from  the  Rome  Beds,  Oregon,  which 
allows  direct  comparison  between  its  own 
parts  and  those  of  material  from  the  type 
locality  and  from  the  Gilliland  local  fauna, 
Texas.  Cranial  elements  from  the  latter  site 
appear  to  have  about  the  same  size  and  shape 
as  those  of  the  Rome  Beds  specimen.  The 
type  of  C.  edwardii.  as  well  as  a  maxillary 
fragment  from  the  same  locality,  not  men- 
tioned in  Gazin's  (1942)  original  description, 
have  an  Ml  with  a  remarkably  pronounced 
buccal  cingulum,  and  the  Rome  Beds  speci- 
men shares  this  character.  This  cingulum  is 
never  so  prominent  in  C.  rufus  as  in  these 
three  specimens,  and  this  factor  is  one  reason 
for  not  now  synonymizing  C.  eduardii  with 
the  red  wolf. 

Kurten  (1974)  considered  the  type  of  C. 
edwardii  to  be  one  of  the  large  Irvingtonian 
and  late  Blancan  coyotes  which  he  grouped 
under  the  name  C.  priscolatrans  ( see  p. 
75).  Although  Kurten  correctly  associ- 
ated C.  edwardii  with  C.  priscolatrans,  I 
think  that  both  names  represent  the  lineage 
of  the  wolf,  rather  than  the  coyote. 

Record  of  occurrences. — The  following 
list  is  arranged  alphabetically  by  state,  except 
that  Chihuahua  is  placed  last.  Specimens  ex- 
amined   by    me    are    identified    by    element, 


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MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


museum  number,  or  both;  and  selected  meas- 
urements are  found  in  appendix  B  (part  11) 
and  appendix  C  (part  3).  Occurrences  also 
are  shown  on  the  map  in  figure  50. 

ARIZONA. — Anita,  Coconino  County;  early  Irv- 
ingtonian  ( Richard  H.  Tedford,  American  Museum 
of  Natural  History,  pers.  comm.);  as  "C.  nubilus?" 
(Hay,  1921:632);  two  mandibular  fragments,  USNM 
10210  R  and  C.  As  reported  by  Hay,  the  most  com- 
plete ramus,  in  comparison  with  that  of  Recent  C. 
lupus,  is  lower  and  thinner,  and  has  thinner  teeth. 
The  proportion  of  length  to  depth  actually  is  unlike 
that  found  in  any  specimen  of  C.  lupus,  but  is  close 
to  that  in  some  specimens  of  Recent  C.  rufus.  The 
other  ramus  also  is  shallow,  but  is  not  complete 
enough  for  full  evaluation.  Still  another  mandibular 
fragment  was  found  at  the  site,  and  was  question- 
ably referred  to  C.  lupus  by  Hay.  This  specimen  is 
much  larger  and  deeper  than  the  other  two,  and 
probably  represents  C.  armbrusteri.  Its  presence  at 
Anita,  together  with  the  other  specimens,  suggests 
that  by  the  early  Irvingtonian  there  already  had 
been  a  divergence  between  the  line  of  small  primi- 
tive wolves  (C.  edwardii  and  C.  rufus)  and  the  line 
leading  to  the  larger  wolves  of  the  late  Quaternary. 
Assignment  of  the  Anita  material  to  C.  edwardii, 
rather  than  to  C.  rufus,  is  arbitrary  because  ade- 
quate samples  are  unavailable. 

Curtis  Ranch,  San  Pedro  Valley,  Cochise  County; 
early  Irvingtonian  (Johnson,  Opdyke,  and  Lindsay, 
1975);  as  C.  edwardii  (Gazin,  1942:499);  skull 
with  mandibles,  USNM  12862;  maxillary  fragment 
with  P3-M2,  USNM  12864. 

KANSAS. — Arkalon  gravel  pit,  south  side  of 
Cimarron  River,  Seward  County;  Irvingtonian  (late 
Kansan ) ;  "The  humerus,  femur  and  other  elements 
are  the  length  of  those  of  a  large  Canis  latrans  Say. 
The  bones  are  heavy  and  nearly  as  large  in  diameter 
as  those  of  Canis  lupus  Linne  (Hibbard,  1953:115). 

Cudahy  fauna,  Rig  Springs  Ranch,  Meade  Coun- 
ty; Irvingtonian  (late  Kansan:  Hibbard,  et  al.,  1965); 
as  Canis  sp.  (Getz,  1960:361).  An  astragalus  was 
reported  to  be  from  a  canid  the  size  of  a  small  wolf 
and  larger  than  a  coyote. 

OREGON. — Rome  Reds,  Malheur  County;  Irving- 
tonian; as  C.  priscolatrans  (Kurten,  1974:6);  par- 
tial skull  without  mandibles,  USNM  23898  (in  U.S. 
Geological  Survey  collections,  Menlo  Park,  Cali- 
fornia). The  specimen  is  approximately  the  same 
size  as  the  type  of  C.  edwardii.  The  teeth  are  larger 
than   those   of   nearly   all   coyotes   examined. 

TEXAS. — Gilliland  local  fauna,  Knox  County; 
Irvingtonian  (late  Kansan:  Hibbard,  et  al.,  1965); 
as  C.  cf.  lupus  (Hibbard  and  Dalquest,  1966:20); 
rostral  fragment,  UMMP  46483;  parietal,  UMMP 
46460.  Hibbard  and  Dalquest  thought  these  frag- 
ments to  represent  "a  canid  the  size  of  the  gray 
wolf,"  but  actually  the  specimens  are  smaller  than 
the  corresponding  parts  of  any  skull  of  C.  lupus 
examined  by  me.    They  clearly  are  not  referable  to 


C.  latrans  or  C.  lepophagus,  and  in  features  that 
can  be  evaluated  they  closely  resemble  the  specimen 
of  C.  edwardii  from  Rome  Reds,  and  also  fall  well 
within  the  range  of  variation  of  C.  rufus.  Hibbard 
(pers.  comm.)  came  to  consider  the  Gilliland  fauna 
to  be  pre-Kansan,  and,  if  so,  these  specimens  repre- 
sent one  of  the  earliest  known  occurrences  of  a  wolf 
in  North  America. 

CHIHUAHUA.— Mina  Erupcion,  90  mi.  SSE 
Juarez;  Pleistocene;  as  Canis  sp.  (Eaton,  1923:233). 
Eaton  wrote  that  six  vertebrae  from  an  adult  animal 
were  smaller  than  those  of  C.  lupus,  but  larger  than 
those  of  C.  latrans.  He  implied  affinity  to  C.  prisco- 
latrans Cope,  1899. 

Miiiaca  Mesa,  approximately  100  mi.  W,  10  mi. 
S  City  of  Chihuahua;  Rlancan  (Kurten,  1974:6); 
mandibular  fragment,  LACM  105/149.  Although 
Kurten  associated  this  specimen  with  the  coyote  line, 
it  is  larger  than  any  mandible  of  C.  lepophagus  or 
C.  latrans  examined  by  me.  The  specimen  does, 
however,  resemble  most  available  material  of  C. 
lepophagus  in  the  pronounced  development  of  the 
second  cusp  of  p4,  and  the  reduced  development  of 
the  third  cusp.  In  addition,  unlike  most  wolves,  p3 
(as  well  as  p2)  lacks  posterior  cusps.  While  referral 
to  C.  edwardii  still  seems  most  appropriate,  the 
possibility  remains  that  this  specimen  represents  a 
transitional  phase  through  which  the  wolf  line  evolved 
from  C.  lepophagus. 

Evolutionary  position. — Canis  edwardii 
may  represent  the  first  unquestionable  ap- 
pearance of  a  wolf  in  North  America.  Ma- 
terial referable  to  this  species  is  on  the  aver- 
age older  and  more  primitive  in  characters 
than  that  assigned  to  other  Pleistocene  species 
of  wolves.  Perhaps  C.  edwardii  descended 
from  a  late  Blancan  population  of  C.  lepopha- 
gus, but  we  also  can  not  rule  out  the  alterna- 
tive that  the  wolf  and  coyote  lines  had  been 
distinct  at  an  earlier  time.  The  presence  of 
a  large  wolf  (C.  armbrusteri)  in  the  early 
Irvingtonian,  as  well  as  C.  edwardii  and  C. 
rufus,  suggests  that  radiation  of  the  wolf 
group  had  been  in  progress  for  a  considerable 
period.  This  radiation  may  have  been  asso- 
ciated with  the  initial  glacial  advances  of  the 
Pleistocene,  and  also  with  the  simultaneous 
extinction  of  the  large  borophagine  dogs. 

The  early  history  of  the  wolves,  and  their 
exact  relationships  with  the  coyotes,  can  not 
now  be  assessed  because  of  the  scarcity  of 
fossil  Canis  in  the  Blancan  and  Hemphillian 
(middle  Pliocene).  A  single  mandibular  frag- 
ment    (UN    2908)     from    the    Hemphillian 


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85 


Mitchell  Creek  Ash  Hollow  formation.  Fron- 
tier County,  Nebraska,  has  measurable  di- 
mensions close  to  those  of  C.  edwardii,  and 
appears  to  represent  a  wolf.  Because  of  the 
poor  condition  of  this  specimen,  and  its  re- 
moval in  time  from  the  scope  of  this  paper, 
I  do  not  now  refer  it  to  a  particular  species. 

Canis  etruscus,  a  wolf  resembling  C.  ed- 
wardii, was  present  in  the  early  Pleistocene 
of  Europe  (Kurten,  1968:109).  Thus,  a  group 
of  small,  relatively  unspecialized  wolves,  re- 
taining some  coyotelike  characters,  seems  to 
have  become  widespread  at  this  time.  This 
group  apparently  formed  the  basic  stock  from 
which  the  larger  wolves  of  the  late  Quater- 
nary descended.  Kurten  (1968:108-109) 
thought  that  C.  etruscus  probably  gave  rise 
to  C.  lupus  mosbachensis  of  the  middle  Pleis- 
tocene, from  which  in  turn  modern  C.  lupus 
developed.  In  the  New  World,  C.  edwardii 
seems  to  have  been  close  to  the  line  from 
which  arose  the  larger  C.  armbrusteri  and  C. 
dirus. 

The  geographical  distribution  of  C.  ed- 
wardii (Fig.  50)  appears  to  have  been  con- 
centrated in  the  southwestern  quarter  of  the 
continent,  while  the  closely  related  C.  rufus 
occupied  the  southeast.  The  latter  was  able 
to  survive,  but  C.  edwardii  eventually  disap- 
peared, perhaps  because  of  changing  habitat 
conditions  or  competition  with  C.  latrans 
and/or  C.  lupus. 

Canis  rufus  Audubon  and  Bachman 

1791.  Lupus  niger  Bartram,  Travels,  p.  199. 
Not  available  because  Bartram  was  not 
consistently  binomial  (according  to 
Int.  Comm.  Zool.  Nomen.,  1957,  opin- 
ion 447 ) . 

1851.  Canis  lupus  var.  Rufus  Audubon  and 
Bachman,  Quadrupeds  of  North  Amer- 
ica, 2:240.  Type  locality,  15  mi.  W 
Austin,  Texas  (Goldman,  Jour.  Mamm., 
18:38,  1937). 

1899.  Canis  priscolatrans  Cope,  Jour.  Acad. 
Nat.  Sci.,  Philadelphia,  ser.  2,  9:227. 
Type  from  Port  Kennedy  deposit,  Up- 


per Merion  Township,  Montgomery 
County,  Pennsylvania.  Valid  as  a  sub- 
species of  C.  rufus. 

1905.  Canis  rufus,  Bailey,  N.  Amer.  Fauna, 
25.174. 

1912.  Canis  floridanus  Miller,  Proc.  Biol.  Soc. 
Washington,  25:95.  Type  from  Horse 
Landing,  about  12  mi.  S  Palatka,  Put- 
nam County,  Florida.  Valid  as  a  sub- 
species of  C.  rufus. 

1937.  Canis  rufus  gregonji  Goldman,  Jour. 
Mamm.,  18:44.  Type  from  Macks 
Bayou,  3  mi.  E  Tensas  River,  18  mi. 
SW  Tallulah,  Madison  Parish,  Louisi- 
ana. 

1942.  Canis  niger,  Harper,  Jour.  Mamm.,  23: 
339. 

1965.  Canis  rufus,  Hall,  Univ.  Kansas  Mus. 
Nat.  Hist.  Misc.  Publ.,  no.  43,  p.  13. 

Recent  subspecies  revised  by  Goldman 
(1944);  subspecies  listed  and  distribution 
mapped  by  Hall  and  Kelson  (1959:851-852), 
and  by  this  paper  (Fig.  50). 

Type. — None  designated. 

Geological  distribution. — Early  Irvington- 
ian  to  Recent. 

Geographical  distribution. — Pleistocene 
and  early  Recent  records  from  Arkansas,  Flor- 
ida, Pennsylvania,  Texas,  and  Estado  de  Mex- 
ico; historical  range  confined  to  southeastern 
quarter  of  North  America,  from  central  Texas 
to  Atlantic,  and  from  Gulf  of  Mexico  north 
to  southern  Pennsylvania,  Ohio  Valley,  and 
southeastern  Kansas;  presently  found  only  in 
extreme  southeastern  Texas  and  southern 
Louisiana. 

Description. — Medium-sized  for  the  genus; 
skull  medium-sized  with  narrow  proportions; 
rostrum  elongated  and  narrow;  braincase  rel- 
atively small  and  not  much  inflated  dorso- 
posteriorly;  postorbital  constriction  elongated, 
narrow  lateromedially,  lateral  margins  often 
appearing  parallel  when  viewed  from  above; 
zygomata  usually  slender  and  not  widely 
spreading;  orbits  usually  large;  frontals  usu- 
ally moderately  elevated  above  rostrum,  not 


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MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


prominently  convex,  and  forming  a  relatively 
narrow  and  flat  shield;  temporal  ridges  often 
sharp,  often  obscuring  frontal  suture,  and 
usually  joining  anterior  to  coronal  suture; 
sagittal  crest  prominent  and  sharp  dorsally; 
supraoccipital  shield  moderately  large;  ex- 
ternal side  of  occipital  often  well  ossified; 
tympanic  bullae  usually  well  inflated;  mandi- 
ble long,  narrow,  and  shallow,  ventral  edge 
usually  not  notably  convex  when  viewed  from 
side;  incisors  often  relatively  small;  upper 
canines  prominent,  thin  anteroposteriorly, 
ventral  tips  usually  extending  below  level  of 
anterior  mental  foramina  when  jaws  are 
closed;  premolars  with  trenchant,  laterally 
compressed  cusps;  P4  usually  with  prominent 
deuterocone  and  lingual  cingulum;  Ml  often 
having  relatively  large,  deeply  sculptured 
medial  section,  the  metaconule  usually  promi- 
nent and  well  separated  from  protocone,  buc- 
cal and  anterior  cingula  usually  pronounced; 
M2  relatively  large,  cusps  well  developed;  p2 
occasionally  with  a  posterior  cusp;  p3  some- 
times with  second  and  third  cusp;  p4  with 
second  cusp,  usually  with  a  moderately  de- 
veloped third  cusp  and  posteromedial  cingu- 
lum extending  behind  third  cusp;  m2  and 
talonid  of  ml  relatively  large,  with  moder- 
ately trenchant  cusps.  For  details  on  pelage 
and  postcranial  skeleton,  see  Goldman 
(1944),  Young  (1946:36),  and  Paradiso  and 
Nowak  (1972b). 

Comparison  with  C.  latrans. — Skull  larger 
and  relatively  broader  in  most  dimensions; 
rostrum  usually  relatively  broader  and  deep- 
er; braincase  relatively  smaller  and  not  so 
much  inflated,  never  broader  at  level  of  pa- 
rietotemporal sutures  than  at  base;  postor- 
bital  constriction  narrower  and  more  elon- 
gated; zygomata  deeper  and  more  widely 
spreading;  jugal  more  deeply  inserted  in  max- 
illa; frontals  often  more  elevated  above  ros- 
trum; temporal  ridges  usually  sharper,  more 
often  obscuring  frontal  suture,  and  usually 
joining  anterior,  rather  than  posterior,  to 
coronal  suture;  sagittal  crest  more  prominent; 
supraoccipital  shield  broader,  projecting  far- 


ther posteriorly;  external  side  of  occipital 
more  ossified,  more  often  lacking  projection 
dorsal  to  foramen  magnum;  occipital  con- 
dyles usually  extending  farther  transversely; 
mandible  usually  relatively  thicker  and  deep- 
er; premolars  set  more  closely  together  in 
jaws;  upper  canines  thicker  anteroposteriorly, 
usually  not  extending  so  far  ventrally;  pre- 
molars usually  relatively  broader  with  less 
trenchant  cusps;  deuterocone  and  lingual 
cingulum  of  P4  usually  less  prominent;  Ml 
usually  with  relatively  smaller  medial  section, 
and  less  prominent  hypocone,  metaconule, 
and  buccal  cingulum;  p2  more  often  with 
posterior  cusp;  posteromedial  cingulum  on 
p4  usually  less  prominent;  metaconid  of  ml 
less  pronounced,  not  projecting  so  far  me- 
dially; m2  and  talonid  of  ml  with  less  trench- 
ant cusps.  Atkins  and  Dillon  ( 1971 )  listed 
differences  between  C.  rufus  and  C.  latrans 
in  the  morphology  of  the  cerebellum;  Russell 
and  Shaw  (1972)  and  Jackson  (1951:240) 
discussed  distinguishing  characters  in  external 
appearance. 

Comparison  with  C.  lupus. — Cranial  dif- 
ferences usually  or  more  often  apparent  are 
as  follows:  skull  smaller  and  relatively  nar- 
rower in  most  dimensions;  rostrum  narrower; 
braincase  relatively  deeper;  lateral  margins 
of  postorbital  constriction  appearing  more 
nearly  parallel  when  viewed  from  above,  not 
rising  so  steeply  into  frontal  region;  zygomata 
more  slender,  not  so  deep,  not  so  widely 
spreading;  orbits  relatively  larger;  frontals 
less  elevated  above  rostrum,  less  convex,  and 
forming  a  flatter  and  relatively  narrower 
shield;  tympanic  bullae  more  inflated;  mandi- 
ble shallower;  incisors  smaller;  upper  canines 
thinner  anteroposteriorly,  extending  more 
ventrally;  premolars  narrower;  P4  with  more 
prominent  deuterocone,  its  root  appearing  to 
pass  more  vertically  into  palate;  Ml  with 
relatively  larger,  more  deeply  sculptured  me- 
dial section,  and  more  prominent  metaconule 
and  buccal  cingulum;  M2  relatively  larger; 
p4  more  often  with  third  cusp  and  postero- 
medial cingulum  extending  behind  third  cusp; 


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87 


talonid  of  ml  relatively  larger  with  more 
trenchant  cusps.  Atkins  and  Dillon  (1971) 
listed  differences  between  C.  rufus  and  C. 
lupus  in  the  morphology  of  the  cerebellum; 
Goldman  (1944),  and  Paradiso  and  Nowak 
(1972b)  discussed  differences  in  pelage  and 
external  appearance. 

Other  comparisons. — See  accounts  of  C. 
edwardii  and  C.  armbrusteri. 

Remarks. — Two  critical  problems  concern- 
ing the  systematics  of  C.  rufus  are:  (1)  its 
original  relationship  with  C.  lupus,  particu- 
larly the  question  of  whether  the  two  inter- 
graded  in  the  forests  of  the  eastern  United 
States;  and  (2)  the  relationship  in  historic 
time  between  C.  rufus  and  C.  latrans  in  the 
south-central  United  States. 

Although  it  is  sometimes  difficult  to  sepa- 
rate specimens  of  red  and  gray  wolves,  the 
previous  part  of  this  paper  showed  that  multi- 
variate analysis  could  distinguish  nearly  all 
skulls  of  C.  rufus,  including  all  taken  before 
1920  in  the  eastern  United  States,  from  large 
series  of  C.  lupus.  In  addition  to  the  meas- 
urements used  in  multivariate  analysis,  the 
characters  listed  in  the  above  "comparison 
with  C.  lupus"  usually  serve  to  distinguish 
the  two  species.  Problems  may  still  arise, 
especially  if  complete  skulls  are  not  available. 

In  nearly  all  measurements  and  other  fea- 
tures in  which  C.  rufus  differs  from  C.  lupus, 
the  former  approaches  C.  latrans.  Indeed, 
available  specimens  of  the  red  wolf  almost 
bridge  the  morphological  gap  between  the 
proximal  extremes  of  the  other  two  species. 
Hybrid  origin  for  C.  rufus  thus  seems  to  be 
one  possibility,  but  there  are  other  solutions 
to  the  problem.  The  most  reasonable  expla- 
nation is  that  C.  rufus  represents  a  primitive 
line  of  wolves  that  has  undergone  less  change 
than  C.  lupus,  and  has  thus  retained  more 
characters  found  in  the  ancestral  stock  from 
which  both  wolves  and  coyotes  arose. 

The  frontal  shield  and  postorbital  con- 
striction are  the  only  parts  of  the  reo1  wolf's 
skull  that  often  do  not  have  a  form  interme- 
diate between  that  of  typical  C.  latrans  and 


C.  lupus.  The  postorbital  constriction  in  C. 
rufus  is  sometimes  relatively  narrower  than 
in  both  the  gray  wolf  and  coyote,  and  the 
elongated  lateral  margins  often  appear  paral- 
lel when  viewed  from  above,  unlike  the  nor- 
mal condition  in  either  of  the  other  two 
species.  The  frontal  shield  of  C.  rufus  is  also 
relatively  narrower,  and  in  some  specimens 
has  a  more  flattened  aspect  than  in  either 
C.  lupus  or  C.  latrans.  These  characters,  to- 
gether with  a  prominently  rising  sagittal  crest, 
give  a  unique  appearance  to  certain  speci- 
mens of  C.  rufus,  including  both  some  pre- 
1920  and  some  post- 1960  individuals. 

Lawrence  and  Bossert  ( 1967 )  considered 
that  if  initial  study  of  the  red  wolf  had  been 
based  on  adequate  series  from  the  southeast- 
ern United  States  (rather  than  from  Texas), 
C.  rufus  and  C.  lupus  probably  would  not 
have  been  taxonomically  separated.  As  we 
have  seen,  however  (pp.  25-2S),  complete 
skulls  taken  prior  to  1920  in  Louisiana  and 
eastward  are  rare,  and  can  all  be  distin- 
guished from  those  of  C.  lupus.  In  addition 
to  the  14  skulls  listed  in  table  2,  Goldman 
(1944)  assigned  two  other  early  specimens 
to  C.  rufus.  One  of  these,  a  subadult  female 
taken  in  1832  on  the  Wabash  River,  Indiana 
(in  AMNH),  was  assigned  to  C.  rufus  gre- 
goryi.  I  agree  with  this  designation,  as  the 
specimen  is  comparatively  small  and  narrow- 
proportioned,  and  has  the  dental  characters 
normally  associated  with  the  red  wolf.  The 
other  skull  (in  USNM),  the  type  of  C.  rufus 
floridamis,  was  taken  in  1890  on  the  St.  Johns 
River,  Putnam  County,  Florida.  This  speci- 
men is  difficult  to  evaluate  because  the  pos- 
terior part  is  missing,  and  there  is  also  a 
dental  anomaly  in  that  M2  on  both  sides  is 
missing.  Nonetheless,  the  specimen  seems  to 
be  within  the  morphological  range  of  other 
skulls  of  C.  rufus. 

Archeological  sites  and  other  Recent  de- 
posits in  the  eastern  United  States  have 
yielded  various  specimens  of  wolves,  but  few 
in  good  condition.   Such  specimens  examined 


88 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


by  me,  which  seem  best  referred  to  C.  rufus, 
include  the  following. 

Banks  site,  1.5  mi.  N  Clarksdale,  Crittenden 
County,  Arkansas;  as  C.  lupus  (Parmalee,  1959c:6); 
mandible,  ISM. 

Blain  site,  west  bank  Scioto  River,  south  of  Chilli- 
cothe,  Ross  County,  Ohio;  as  C.  lupus  (Parmalee 
and  Shane,   1970:198);  maxillary  fragment,  ISM. 

New  Paris  Sinkhole  No.  2,  Bedford  County, 
Pennsylvania;  as  C.  lupus  hjcaon  (Guilday  and  Ben- 
der, 1958:134);  incomplete  skull  and  pair  of  mandi- 
bles, CM. 

Eschelman  site,  3  mi.  S  Columbia,  Lancaster 
County,  Pennsylvania;  as  C.  lupus  (Guilday,  Parma- 
lee, and  Tanner,  1962:64);  cranial  fragment  and  pair 
of  mandibles,  CM. 

Buffalo  Village  site,  Putnam  County,  West  Vir- 
ginia; as  C.  lupus  (Guilday,  1971:9);  three  mandib- 
ular fragments  and  isolated  ml,  CM. 

Lauderdale  Indian  mound,  Washington  County, 
Virginia;  isolated  Ml  (collection  of  Ronald  M. 
Nowak). 

Crow  Island  Indian  midden,  Jackson  County,  Ala- 
bama; mandible,  UMMZ.  Recently,  Barkalow  (1976: 
25-26)  reported  this  specimen,  and  material  from 
two  other  sites  in  the  \icinity,  to  be  either  C.  rufus 
or  C.  lupus. 

Jungerman  site,  Indian  River,  just  south  of  south- 
ern tip  of  Merritt  Island,  Brevard  County,  Florida; 
as  C.  cf.  niger   (Wing,  1963:52);  ml,  m2,  UF. 

Nichol's  Hammock,  .7  mi.  NE  Princeton,  Dade 
County,  Florida;  sinkhole  with  contemporary  fauna; 
as  C.  niger  (Hirschfeld,  1968:180);  mandible,  UF. 

Bullen  and  Benson  (1967)  reported  the 
discovery  of  three  cut  and  perforated  canid 
jaws  on  Tick  Island,  Florida.  Although  refer- 
ral to  C.  rufus  is  a  possibility,  the  fragmentary 
nature  of  the  material  (as  illustrated  by  Bul- 
len and  Benson)  would  make  assignment  to 
any  particular  species  of  Canis  difficult. 

Webb  and  Baby  (1957:61-71)  described 
three  specimens  of  wolves  from  the  Wright 
Mounds,  Montgomery  County,  Kentucky; 
near  New  Liberty,  Owen  County,  Kentucky; 
and  the  Wolford  Mounds,  Pickaway  County, 
Ohio.  The  most  complete  of  these  specimens, 
from  the  Wright  Mounds,  was  a  spatula- 
shaped  artifact  cut  from  the  upper  jaws  and 
palate  of  a  wolf.  The  specimen  now  has  ap- 
parently been  lost,  but  a  published  photo- 
graph suggests  that  the  skull  represented  had 
a  narrow  rostrum,  and  may  have  belonged  to 
a  red  wolf. 

Fossil   record. — The  following  list   is   ar- 


ranged alphabetically  by  state,  except  that 
Estado  de  Mexico  is  placed  last.  Specimens 
examined  by  me  are  identified  by  element, 
museum  number,  or  both;  and  selected  meas- 
urements are  found  in  appendix  B  (part  12) 
and  appendix  C  (part  4).  Occurrences  also 
are  shown  on  the  map  in  figure  50. 

ARKANSAS— Eddy  Bluff  shelter,  near  Spring- 
dale,  Washington  County;  early  Recent;  as  C.  rufus 
(Morrison,  1970);  maxillary  fragment,  UArk.  The 
fossil  closely  matches  series  of  modern  C.  rufus  in 
size  and  other  characters. 

FLORIDA.— Haile  VILA,  Alachua  County;  Sanga- 
mon (Webb,  1974b:  13);  cranial  fragment,  UF.  The 
specimen  is  larger  than  comparative  material  of  C. 
latrans  and  is  smaller  than  C.  lupus  or  C.  dirus.  In 
size,  and  in  characters  of  the  frontal  region  and 
dentition,  the  specimen  is  well  within  the  range  of 
variation  of  Recent  C.  rufus.  Martin  (1974:77) 
compared  measurements  of  the  P4  (incorrectly  la- 
beled as  p4  in  his  figure  3.13)  of  nine  specimens 
from  this  site,  to  those  of  other  wolves,  and  stated 
that  either  C.  lupus  or  C.  rufus  was  represented. 
According  to  his  scatter  diagram,  the  measurements 
of  the  Haile  material  are  substantially  closer  to  those 
of  C.  rufus  than  to  those  of  C.  lupus. 

Devil's  Den,  near  Williston,  Levy  County;  late 
Wisconsin  or  early  Recent  (7,000-8,000  B.P.);  as 
C.   rufus   (Martin  and  Webb,  1974:126). 

Inglis  IA,  Citrus  County;  early  Irvingtonian;  as 
C.  cf.  niger  (Klein,  1971:17),  as  C.  rufus  (Webb, 
1974b:  17),  as  C.  lupus  (Martin,  1974:72),  as  C. 
priscolatrans  ( Kurten,  1974:6);  right  and  left  maxil- 
lary fragments,  UF  18046;  P4,  UF  18049;  cl,  UF 
18052;  Ml,  UF  19406;  ml,  UF  19404;  two  mandib- 
ular fragments,  UF  19323,  19324.  This  material 
indicates  the  presence  of  a  canid  close  in  size  and 
dental  characters  to  the  wolf  (C.  rufus)  that  inhab- 
ited the  southeast  in  historical  time.  Several  of  the 
specimens  are  larger  than  those  of  any  Recent  or 
late  Pleistocene  coyote  examined  by  me.  The  first 
upper  molars  in  this  series  have  deeply  sculptured 
medial  sections,  as  does  the  type  of  C.  priscolatrans, 
but  neither  the  Inglis  nor  Port  Kennedy  specimens 
have  the  buccal  cingulum  on  Ml  as  strongly  devel- 
oped as  in  C.  edwardii.  Klein  (1971:17-18)  ob- 
served that  the  measurements  of  the  Inglis  speci- 
mens approached  those  of  C.  edwardii  from  Curtis 
Ranch,  and  that  these  specimens  indicated  a  wolf 
very  close  to,  if  not  conspecific  with,  C.  rufus. 

Crystal  River  Power  Plant,  Citrus  County;  San- 
gamon; maxillary  fragment,  UF  17074.  Kurten 
(1974:10)  assigned  another  maxilla  from  this  site 
to  C.  latrans,  but  noted  that  it  was  "large."  Possibly 
that  specimen  should  be  referred  to  C.  rufus. 

Melbourne,  Brevard  County;  Wisconsin  (Hib- 
bard,  et  al.,  1965);  as  C.  cf.  lupus  (Ray,  1958:434), 
as  C.  rufus  (Webb,  1974b:  17);  mandibular  frag- 
ment, MCZ  17789.  As  explained  by  Ray,  C.  lupus 
and  C.  rufus  can  not  always  be  distinguished  on  the 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


1       ' 


scale  of  m.les 


Fig.  50. — Map  showing  localities  of  C.  rufus  from  archeological  sites  (triangles),  fossil  C.  rufus  (black 
dots),  and  C.  edwardii  (squares).  The  solid  lines  show  the  distribution  of  Recent  subspecies:  C.  rufus  rufus 
(R),  C.  rufus  gregoryi  (G),  and  C.  rufus  floridanus  (F).  Recause  of  the  scale  of  the  map,  it  was  not  possi- 
ble to  plot  all  localities  in  crowded  areas. 


basis  of  the  mandible  and  lower  teeth.    I  tentatively  it   has    a    posteromedial    cingulum    extending   behind 

refer  the  Melbourne  specimen  to  C.  rufus  because  it  the  third  cusp  of  p4. 

is  within  the  size  range  of  that  species,  and  because  Vero,     Indian     River     Country;     late     Wisconsin 


90 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


(Webb,  1974b:13);  as  C.  cf.  niger  (Weigel,  1962: 
37).  Only  a  PI  and  the  anterior  half  of  a  P4  were 
so  referred,  and  Weigel  considered  the  material  too 
meager  for  positive  identification.  Webb  (1974b:17), 
however,  listed  C.   rufus  for  this  site. 

PENNSYLVANIA.— Port  Kennedy  deposit,  Upper 
Merion  Township,  Montgomery  County;  Irvingtonian 
(probably  Yarmouthian:  Hihbard,  1958);  as  C. 
priscolatrans  (Cope,  1899:227);  P4,  Ml,  M2,  p4, 
ANSP  57-58.  Cope  regarded  the  upper  teeth  as 
"the  type  of  a  distinct  species,  having  important 
points  of  resemblance  to  the  coyote"  ( although  his 
publication  listed  the  premolar  as  a  PI,  the  tooth 
actually  is  a  P4).  Cope  also  noted:  "The  forms  of 
the  cusps  and  cingula  in  this  species  are  like  those 
of  the  corresponding  teeth  of  the  coyote,  except  as 
to  the  conules.  The  size  is  that  of  the  large,  but  not 
largest  wolves."  This  description  agrees  well  with 
that  of  C.  rufus  by  Goldman  (1944),  and  Paradiso 
and  Nowak  (1972a),  who  considered  the  red  wolf 
to  have  coyotelike  teeth,  but  to  approach  C.  lupus  in 
size.  A  very  few  specimens  of  Recent  and  late 
Pleistocene  C.  latrans  have  teeth  as  large  as  those 
found  at  Port  Kennedy,  but  I  disagree  with  Kurten's 
(1974)  suggestion  that  the  type  of  C.  priscolatrans 
represents  a  coyote  ancestral  to  modern  C.  latrans. 
In  contrast,  the  Port  Kennedy  teeth  fall  well  within 
the  range  of  variation  of  Recent  C.  rufus,  and  are 
nearly  equal  in  size  to  those  of  C.  edwardii. 

TEXAS. — Miller's  Cave,  Llano  County;  late  Wis- 
consin; as  Cains  sp.  (Patton,  1963:31).  According 
to  Patton,  a  single  m2  from  the  site  was  larger  than 
those  of  coyotes,  and  slightly  larger  than  that  of  one 
available  specimen  of  C.  rufus. 

Buffalo  Bayou,  Houston,  Harris  County;  late 
Pleistocene;  as  "Canis  sp.  cf.  lupins  [sic]  baileyi" 
( Du  Bar  and  Clopine,  1961:99).  Since  the  gray 
wolf  is  not  known  to  have  occurred  near  Houston 
in  Recent  time,  since  the  nearest  geographical  sub- 
species to  Houston  is  not  baileyi,  and  since  the 
specific  name  was  not  correctly  spelled,  there  is 
reason  to  suspect  that  identification  of  the  pertinent 
material  was  not  made  carefully.  The  specimen 
apparently  has  been  lost,  but  I  think  that  any  late 
Pleistocene  remains  of  small  wolves  in  the  area  would 
be  referable  to  C.  rufus. 

ESTADO  DE  MEXICO.— Upper  Becerra  forma- 
tion, northwest  of  Puente  del  Gallo,  Valley  of  Te- 
quixquiac;  Sangamon  or  Wisconsin;  as  Canis  sp. 
(Hibbard,  1955:52).  According  to  Hibbard,  a  man- 
dible from  the  site  "is  smaller  than  Canis  lupus 
Linnaeus  and  appears  closely  related  to  Canis  niger 
( Bartram ) ." 

Evolutionary  position. — Modern  C.  rufus 
apparently  represents  a  comparatively  un- 
modified surviving  line  of  the  primitive  stock 
of  small  wolves  that  had  developed  by  the 
early  Pleistocene.  The  red  wolf  evolved  from 
C.  edwardii,  or  a  close  relative,  and  then 
remained  in  North  America  through  the  mid- 


dle and  late  Quaternary.  The  gray  wolf  prob- 
ably evolved  from  a  branch  of  the  same  stock, 
but  one  that  had  entered  the  Old  World  and 
become  isolated  there  through  factors  asso- 
ciated with  glaciation.  While  C.  lupus  devel- 
oped in  Eurasia  and  eventually  became  the 
only  species  of  wolf  throughout  most  of  the 
Northern  Hemisphere,  and  while  C.  dims 
underwent  its  sudden  rise  and  fall  in  the 
New  World,  the  smaller  C.  rufus  held  on  to 
its  niche  in  the  southern  forests  and  marshes. 

Goldman  (1944:399)  wrote  that  certain 
Pleistocene  remains  from  Rancho  La  Rrea 
suggested  the  presence  there  of  a  species  with 
relationship  to  C.  rufus.  All  of  the  specimens 
of  wild  Canis  from  Rancho  La  Brea  that  I 
examined,  however,  could  be  referred  to  C. 
dints,  C.  lupus,  or  C.  latrans. 

There  has  been  a  suggestion  that  C.  rufus 
evolved  from  a  coyotelike  ancestor  that  had 
become  isolated  by  glaciation  in  a  Florida 
refugium  (Nowak,  1970:84).  This  hypothesis 
no  longer  is  tenable  in  the  light  of  the  above 
outlined  evolutionary  sequence  of  the  red 
wolf.  Furthermore,  the  subspecies  of  coyote 
(C.  latrans  riviveronis)  that  inhabited  Florida 
was  small,  and  survived  into  early  Recent 
time,  and  hence  could  not  have  given  rise  to 
the  much  larger  C.  rufus  which  already  was 
present  in  Florida  by  the  Irvingtonian. 

Canis  armbrusteri  Gidley 

1913.    Canis   armbrusteri   Gidley,    Proc.    U.S. 
Natl.  Mus.,  46:98. 

Type. — Portion  of  a  left  lower  jaw  con- 
taining p4  to  m2;  no.  7662,  U.S.  Natl.  Mus.; 
Cumberland  Cave,  about  4  mi.  NW  Cumber- 
land, Allegany  County,  Maryland. 

Geological  distribution. — Early  (?)  Irv- 
ingtonian to  early  Rancholabrean. 

Geographical  distribution. — Known  from 
Maryland  and  Florida,  with  possible  records 
from  Arizona,  California,  Nebraska,  Pennsyl- 
vania, South  Carolina,  and  Texas. 

Description. — Size  large  for  the  genus; 
skull  usually  large  and  relatively  narrow  in 


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91 


most  proportions;  rostrum  elongated  and  nar- 
row; braincase  moderately  inflated  dorsally; 
zygomata  usually  deep  and  broadly  spread- 
ing; frontals  moderately  elevated  above  ros- 
trum, not  prominently  convex,  and  forming 
relatively  narrow  shield;  sagittal  crest  promi- 
nent and  sharp  dorsally;  supraoccipital  shield 
large;  tympanic  bullae  notably  large  and  well 
inflated;  mandible  long,  moderate  in  depth; 
teeth  comparatively  large;  P4  usually  with 
prominent  deuterocone;  Ml  having  relatively 
large,  deeply  sculptured  medial  section,  and 
pronounced  buccal  cingulum;  M2  relatively 
large;  p2  and  p3  lacking  posterior  cusps  in 
available  specimens;  p4  having  second  and 
third  cusps,  and  pronounced  posteromedial 
cingulum  extending  well  behind  third  cusp: 
talonid  of  ml  relatively  large. 

Comparison  with  C.  rufus. — Usually  much 
larger;  postorbital  constriction  with  lateral 
margins  not  parallel;  zygomata  usually  deep- 
er and  more  broadly  flaring;  mandible  rela- 
tively deeper;  teeth,  especially  carnassials, 
sometimes  relatively  larger;  p4  having  more 
pronounced  posteromedial  cingulum. 

Comparison  with  C.  lupus. — Skull  usually 
narrower  in  most  proportions;  rostrum  rela- 
tively longer  and  narrower;  braincase  usually 
more  inflated  dorsally;  frontals  less  convex 
and  usually  forming  relatively  narrower 
shield;  tympanic  bullae  larger  and  more  in- 
flated; P4  usually  having  more  prominent 
deuterocone;  Ml  having  relatively  larger, 
more  deeply  sculptured  medial  section,  and 
pronounced  buccal  cingulum;  M2  relatively 
larger;  p4  with  third  cusp,  and  pronounced 
posteromedial  cingulum  extending  behind 
third  cusp;  ml  with  relatively  larger  talonid. 

Comparison  with  C.  dims. — Usually 
smaller;  skull  narrower  in  most  proportions; 
rostrum  relatively  longer  and  much  narrower; 
braincase  more  inflated  dorsally;  postorbital 
constriction  not  rising  so  steeply  into  frontal 
region;  frontal  shield  much  narrower;  sagittal 
crest  usually  less  prominent;  supraoccipital 
shield  broader  and  not  projecting  so  far  pos- 
teriorly; tympanic  bullae  larger  and  more  in- 


flated; postpalatine  foramina  more  anteriorly 
placed  ( arrangement  of  the  optic  and  anterior 
lacerated  foramina  can  not  be  evaluated  in 
available  specimens  of  C.  armbrusteri);  an- 
terior parts  of  vertical  plates  of  palatines 
flaring  less  broadly  (placement  of  the  vomer 
can  not  be  evaluated);  mandible  usually 
shallower;  P4  usually  relatively  smaller  with 
more  prominent  deuterocone;  Ml  having  rel- 
atively larger,  more  deeply  sculptured  medial 
section,  more  reduced  paracone  and  meta- 
cone,  more  prominent  hypoeone  with  its  an- 
terior ridge  extending  around  protocone,  and 
more  pronounced  buccal  cingulum;  M2  rela- 
tively larger;  p4  usually  similar;  ml  smaller 
with  relatively  larger  talonid. 

Remarks. — Gidley's  (1913)  original  de- 
scription of  C.  armbrusteri  was  based  on  three 
lower  jaws  from  Cumberland  Cave,  which 
reportedly  differed  from  those  of  C.  lupus  in 
having  relatively  greater  depth,  smaller  ca- 
nines, p2  and  p3  without  posterior  cusps,  p4 
with  a  third  cusp  and  posterior  cingulum, 
and  ml  with  a  larger  heel.  Not  all  of  these 
characters  can  now  be  considered  diagnostic, 
but  on  the  whole  Gidley's  distinction  of  C. 
armbrusteri  was  borne  out  by  the  discovery 
of  additional  material.  The  upper  teeth  of 
the  species  were  first  described  by  Patterson 
(1932),  who,  like  Gidley,  noted  certain  coy- 
otelike characters.  Gidley  and  Gazin  (1938: 
15-23)  discussed  a  number  of  skulls  and  man- 
dibles from  Cumberland  Cave,  which  sup- 
ported the  continued  recognition  of  C.  arm- 
brusteri. 

Martin  (1974:76)  suggested  that  C.  arm- 
brusteri is  synonymous  with  C.  lupus,  and 
that  specimens  from  Cumberland  Cave  are 
closely  matched  by  skulls  of  large,  northern 
gray  wolves.  I  disagree  with  this  interpreta- 
tion for  reasons  provided  in  the  above  com- 
parison of  the  two  species.  Probably  of  great- 
est value  in  distinguishing  C.  armbrusteri 
from  Recent  C.  lupus  is  the  presence  in  the 
former  of  a  pronounced  buccal  cingulum  on 
the  Ml,  and  a  posteromedial  cingulum  ex- 
tending well   behind  the   third   cusp   on   p4. 


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MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


These  characters  are  conspicuous  in  speci- 
mens both  from  Cumberland  Cave  and  Flor- 
ida. Also,  the  bullae  of  C.  armbrusteri  are 
more  inflated  than  those  of  C.  lupus,  and,  in 
fact,  are  larger  than  those  of  any  other  species 
of  Cards, 

Goldman  (1944:399)  thought  that  the 
Cumberland  Cave  wolf  appeared  closely  al- 
lied to  the  red  wolf.  In  most  proportions  and 
dental  characters,  C.  armbrusteri  does  ap- 
proach Recent  C.  rufus,  and  there  would  be 
a  basis  for  considering  it  a  giant  Pleistocene 
red  wolf.  The  two  species  are  easily  distin- 
guished by  size  and  other  characters,  how- 
ever, and  there  is  some  evidence  that  they 
occurred  together  at  certain  Pleistocene  lo- 
calities. 

Whereas  some  specimens  of  C.  armbru- 
steri seem  not  very  different  from  C.  rufus, 
others  are  nearly  as  large  as  C.  dims.  Al- 
though the  range  of  variation  shown  by  C. 
armbrusteri  is  not  unusually  great  for  a  spe- 
cies of  Canis,  the  available  material  does  span 
much  of  the  moi-phological  gap  between  C. 
rufus  and  C.  dims,  and  may  represent  a  part 
of  the  evolutionary  sequence  through  which 
the  dire  wolf  developed  from  more  primitive 
stock.  Martin  (1974:75)  reported  that  one  of 
the  Cumberland  Cave  skulls  (USNM  11886) 
has  an  inion  projection  as  pronounced  and 
hooked  as  in  C.  dims,  and  might  represent 
a  population  beginning  to  develop  into  that 
species.  In  my  own  opinion  this  specimen 
does  match  some  skulls  of  C.  dirus  in  size 
and  height  of  sagittal  crest,  but  not  in  projec- 
tion of  inion.  Furthermore,  in  all  distinguish- 
ing characters  of  the  dentition  that  can  be 
evaluated,  the  specimen  is  unlike  C.  dirus. 

Record  of  occurrences. — Only  the  remains 
from  Cumberland  Cave  and  two  Florida  sites 
are  complete  enough  for  reliable  assignment 
to  C.  armbrusteri.  Various  other  fragments  of 
large  wolves  have  been  reported  from  pre- 
Illinoian  sites,  and  are  not  definitely  referable 
to  C.  lupus  or  C.  dirus  on  a  moiphological  or 
chronological  basis  (specimens  with  the  typi- 
cal  characters   of  these  two   species   do   not 


appear  in  North  America  until  the  Illinoian). 
The  earlier  material  may  represent  the  line- 
age of  C.  armbrusteri  and  is  listed  at  this 
point.  The  following  list  is  arranged  alpha- 
betically by  state;  specimens  examined  by  me 
are  identified  by  element,  museum  number, 
or  both;  and  selected  measurements  are  found 
in  appendix  R  (part  13)  and  appendix  C 
(part  5).  Occurrences  also  are  shown  on  the 
map  in  figure  45. 

ARIZONA. — Anita,  Coconino  County;  early  Irv- 
ingtonian  ( Richard  H.  Tedford.  American  Museum 
of  Natural  History,  pers.  comm.);  as  "C.  nuhilus?" 
(Hay,  1921:632);  mandibular  fragment,  USNM 
10210  A.  Of  three  such  specimens  from  this  site, 
two  are  referred  above  to  C.  edwardii.  The  third 
mandible  is  much  larger  and  deeper,  and  is  almost 
identical  in  size  and  proportion  with  certain  speci- 
mens from  Cumberland  Cave. 

CALIFORNIA. — Irvington,  Alameda  County; 
Irvingtonian;  as  C.  cf.  dirus  (Savage,  1951:230). 
The  few  pertinent  fragments  from  this  site  do  not 
seem  adequate  for  identification,  but  do  represent  a 
large  canid  and  may  be  referable  to  C.  armbrusteri. 

FLORIDA.— McCleod  lime  rock  mine,  2.5  mi.  N 
Williston,  Levy  County;  Irvingtonian;  cranial  frag- 
ment, AMNH  67286;  two  maxillary  fragments  (prob- 
ably from  same  individual),  AMNH  67287-67288; 
two  mandibular  fragments  ( probably  from  same  in- 
dividual), AMNH  67289-67290;  mandibular  frag- 
ment, AMNH  67291.  The  specimens  are  large,  but 
do  not  match  C.  dims  in  size  or  other  critical  char- 
acters. Referral  to  C.  armbrusteri  is  supported  by 
the  presence  of  a  pronounced  buccal  cingulum  on 
Ml,  a  prominent  deuterocone  on  P4,  a  pronounced 
posteromedial  cingulum  on  p4,  and  a  relatively 
shallow  mandible. 

Coleman  IIA  local  fauna,  Sumter  County;  Irving- 
tonian; as  C.  lupus  (Martin,  1974:76);  skull  without 
mandibles,  UF  11519;  cranial  fragment,  maxillary 
fragment,  and  three  mandibular  fragments,  UF 
11520;  mandibular  fragment,  two  ml,  UF  12121; 
mandibular  fragment,  UF  11518;  two  P4,  UF  12114; 
various  teeth  and  postcranial  elements,  UF.  Al- 
though Martin  (1974:75)  considered  one  skull  of 
C.  armbrusteri  from  Cumberland  Cave  (USNM 
7994 )  to  be  "essentially  identical"  to  the  Coleman 
skull  UF  11519,  he  assigned  these  and  other  speci- 
mens from  both  sites  to  C.  lupus.  For  reasons  stated 
above  I  recognize  C.  armbrusteri  as  a  distinct  spe- 
cies, probably  most  closely  related  to  C.  rufus,  and 
the  Coleman  material  seems  best  referred  to  C. 
armbrusteri.  Some  of  the  specimens  are  compara- 
tively small,  but  most  dimensions  fall  within  the  size 
range  for  the  Cumberland  material.  Other  char- 
acters in  which  the  Coleman  wolves  resemble  C. 
armbrusteri,  rather  than  C.  lupus,  include:  the  pro- 
nounced buccal  cingulum,  and  relatively  large,  deeply 
sculptured     medial    section    on     Ml;    the    prominent 


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93 


deuterocone  on  P4;  the  pronounced  posteromedial 
cingulum  extending  well  behind  the  third  cusp  on 
p4;  and  the  large,  well  inflated  bullae  of  the  one 
specimen  on  which  they  could  he  evaluated. 

MARYLAND.— Cumberland  Cave,  4  mi.  NW 
Cumberland,  Allegany  County;  Illinoian  (Kurten, 
1963:100);  as  C.  armbrusteri  (Gidley,  1913:98;  Gid- 
ley  and  Gazin,  1938:15),  as  C.  lupus  (Martin,  1974: 
76);  eight  skulls,  USNM  7994,  8144,  11881,  11883, 
11885,  11886,  11887,  12288;  13  mandibular  fragments, 
USNM  7482,  7661,  8144,  8168,  8169,  8172,  11881, 
11882,  11887,  11888,  12290,  12293,  12295;  P4, 
USNM   12289;   Ml,  M2,  FM  P14790. 

NEBRASKA. — Rushville  fossil  quarry,  Sheridan 
County;  Yarmouthian  ( Schultz  and  Martin,  1970); 
as  C.  dirus  nebrascensis  Frick  ( Schultz  and  Tanner, 
1957:71);  maxillary  fragment,  UN  25691.  The  fea- 
tures of  the  Ml  in  this  specimen  are  not  those  of 
C.  dims,  but  fall  within  the  range  of  variation  shown 
by  C.  armbrusteri. 

Angus  fossil  quarry,  Nuckolls  County;  Yarmouth- 
ian; as  C.  dirus  nebrascensis  Frick  ( Schultz  and 
Martin,  1970:347).  According  to  Larry  D.  Martin 
( Department  of  Systematies  and  Ecology,  University 
of  Kansas,  pers.  comm. ),  this  record  is  based  on  an 
ulna,  but  Merriam  (1912:236)  reported  that  in  C. 
dirus  this  element  shows  no  sharp  distinguishing  char- 
acters. 

PENNSYLVANIA.— Port  Kennedy  deposit,  Up- 
per Merion  Township,  Montgomery  County;  Irving- 
tonian  (probably  Yarmouthian:  Hibbard,  1958);  as 
possibly  C.  indianensis  (Cope,  1899:227).  Cope 
reported  three  postcranial  elements  to  be  larger  than 
those  of  any  wolf  known  to  him.  Such  material  is 
not  reliable  in  the  identification  of  the  dire  wolf, 
and  the  record  herein  is  listed  under  C.  armbrusteri. 

SOUTH  CAROLINA.— Ashley  River,  Charleston 
County;  Pleistocene;  as  C.  occidentalis  (Hay,  1923: 
365).  A  mandibular  fragment  with  p4  was  compared 
by  Hay  to  C.  dirus  and  C.  lupus,  and  was  found  to 
be  closer  in  size  to  the  latter.  Unfortunately,  the 
single  specimen  of  C.  lupus  used  by  Hay  (USNM 
9001 )  is  the  largest  skull  of  that  species  that  I  have 
examined,  and  thus  is  hardly  typical.  It  -seems  un- 
likely that  gray  wolves  of  this  size  ever  occurred  as 
far  to  the  southeast  as  Charleston.  But  the  measure- 
ments of  depth  of  jaw  ( 28.0  millimeters )  and  length 
of  p4  (18.5  millimeters),  listed  by  Hay  for  the 
Ashley  River  specimen,  are  almost  identical  to  those 
of  several  specimens  of  C.  armbrusteri  from  Cum- 
berland Cave  and  Florida. 

TEXAS. — Rock  Creek,  Briscoe  County;  Kansan 
(Hibbard,  1970);  as  C.  dirus  (Troxe.ll,  1915:633). 
Troxell  referred  a  tibia  and  several  other  postcranial 
elements  to  C.  dirus,  solely  on  the  basis  of  size. 
According  to  Stock  and  Lance  (1948),  however,  the 
body  of  C.  dirus  was  small  relative  to  its  skull.  Thus 
size  would  not  be  a  reliable  character  in  distinguish- 
ing the  postcranial  skeleton  of  the  dire  wolf  from 
that  of  other  large  species. 


Evolutionary  position. — Cards  armbrusteri 
is  one  of  several  large  species  that  arose  from 
the  basal  stock  of  primitive  wolves  repre- 
sented by  C.  edwardii  and  C.  rufus.  Descent 
could  have  been  directly  from  either  of  these 
latter  two  species.  The  presence  of  speci- 
mens of  large  wolves  at  several  early  Irving- 
tonian  sites  suggests  that  divergence  between 
the  lineages  of  C.  armbrusteri  and  C.  rufus 
occurred  early  in  the  Pleistocene,  and  is  evi- 
dence for  a  lengthy  independent  evolution  of 
the  wolf  group.  Unfortunately,  this  early 
material  is  so  fragmentary  that  it  is  impossible 
to  determine  how  many  species  of  wolves  are 
represented.  Specimens  clearly  showing  typi- 
cal characters  of  C.  lupus  and  C.  dirus  do 
not  appear  in  North  America  until  the  Illi- 
noian, at  which  time  the  less  specialized  C. 
armbrusteri  was  still  present.  Therefore,  it  is 
reasonable  to  suppose  that  before  then  C. 
armbrusteri  was  the  only  large  wolf  in  North 
America,  and  that  it  may  have  occurred  over 
much  of  the  continent.  In  the  Illinoian,  C. 
armbrusteri  might  have  become  restricted  to 
the  east  by  the  initial  movement  of  circum- 
polar  C.  lupus  into  the  plains  and  western 
mountains.  Canis  armbrusteri  disappeared 
by  the  end  of  the  Illinoian,  but  we  do  not 
know  if  its  lineage  ended  then  or  if  it  gave 
rise  to  C.  dirus,  as  suggested  by  Martin 
(1974:76).  This  latter  hypothesis  is  not  ade- 
quately supported  by  available  morphologi- 
cal evidence,  and  perhaps  C.  dirus  was  a 
replacement  for,  rather  than  a  descendent  of, 
C.  armbrusteri. 

Canis  lupus  Linnaeus 

1758.    Canis    lupus    Linnaeus,    Systema    Na- 

utrae,   10th  ed.,  p.  39.    Type  locality, 

Sweden. 
1910.    Canis    occidentalis   furlongi    Merriam, 

Univ.     California     Publ.     Bull.     Dept. 

Geol.,   5:393.    Type  from   Rancho   La 

Brea,  Los  Angeles  County,  California. 

Valid   as   a  subspecies   of  C.   lupus. 
1912.    Canis    milled    Merriam,    Mem.    Univ. 

California,   1:247.    Type  from  Rancho 


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MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


La  Brea,  Los  Angeles  County,  Califor- 
nia.   A  synonym  of  C.  lupus  furlongi. 
1918.    Aenocyon  milleri,  Merriam,  Univ.  Cali- 
fornia Publ.  Bull.  Dept.  Geol.,  10:533. 

In  addition  to  those  listed  above,  24  names 
based  on  North  American  Recent  specimens 
are  available  for  use  at  the  subspecific  level. 
These  are  to  be  found  in  the  systematic  re- 
vision by  Goldman  (1944),  and  also  are 
listed  by  Hall  and  Kelson  (1959:847-851). 

Type. — None  designated. 

Geological  distribution. — Late  Irvington- 
ian  to  Recent  in  North  America. 

Geographical  distribution. — Pleistocene 
and  early  Recent  records  from  Alberta,  Sas- 
katchewan, Yukon,  Alaska,  Arizona,  Arkan- 
sas, California,  Colorado,  Georgia,  Idaho,  Illi- 
nois, Kansas,  Michigan,  Minnesota,  Nebraska, 
Nevada,  New  Mexico,  Oklahoma,  Oregon, 
Pennsylvania,  Texas,  Virginia,  Wisconsin, 
Wyoming,  Nuevo  Leon,  and  many  localities 
in  Eurasia.  Historical  range  throughout  Eu- 
rasia, except  tropical  forests  of  southeastern 
corner;  throughout  North  America,  except 
parts  of  southeastern  quarter,  southern  and 
coastal  Mexico,  Central  America,  Baja  Cali- 
fornia, and  most  of  California;  and  on  most 
adjacent  continental  islands.  Presently  extir- 
pated in  many  areas  settled  by  man,  includ- 
ing most  of  Europe  and  the  48  southern  con- 
tinental states  of  the  United  States. 

Description. — Size  large  for  the  genus; 
skull  usually  large  with  mostly  broad  propor- 
tions; rostrum  elongated,  usually  relatively 
broad  and  deep;  braincase  relatively  small, 
not  much  inflated  dorsoposteriorly;  postor- 
bital  constriction  elongated,  narrow  latero- 
medially;  zygomata  thick,  deep,  broadly  flar- 
ing; orbits  relatively  small;  frontals  usually 
well  elevated  above  rostrum,  prominendy 
convex,  forming  relatively  broad  shield;  tem- 
poral ridges  sharp,  often  obscuring  frontal 
suture,  usually  joining  anterior  to  coronal  su- 
ture; sagittal  crest  prominent,  sharp  dorsally; 
supraoccipital  shield  large;  external  side  of 
occipital  well  ossified;  tympanic  bullae  usu- 


ally moderate  in  size,  not  much  inflated;  man- 
dible thick  and  deep,  ventral  margin  not  con- 
vex when  viewed  from  side,  toothrow  bowed 
outward  in  center;  incisors  relatively  large; 
upper  canines  prominent,  thick  anteropos- 
teriorly,  alveoli  set  relatively  high  in  premax- 
illae,  ventral  tips  usually  not  extending  to 
level  of  anterior  mental  foramina  when  jaws 
are  closed;  premolars  relatively  broad;  P4 
usually  lacking  prominent  deuterocone  and 
lingual  cingulum;  Ml  having  relatively  large 
paracone  and  metacone,  relatively  small  me- 
dial section  without  trenchant  cusps,  the 
metaconule  reduced  and  not  well  separated 
from  protocone;  Ml  lacking  pronounced  buc- 
cal cingulum;  M2  usually  relatively  small; 
p2  often  with  posterior  cusp;  p3  usually  with 
second  and  third  cusp;  p4  with  second  cusp, 
sometimes  lacking  third  cusp,  usually  with- 
out posteromedial  cingulum  extending  be- 
hind third  cusp;  ml  relatively  broad,  usually 
having  relatively  small  talonid.  For  details 
on  pelage  and  postcranial  skeleton  see  Gold- 
man (1944),  Iljin  (1941),  Mech  (1970),  and 
Hildebrand  (1952a,  1952b,  1954). 

Comparison  with  C.  latrans. — Usually 
much  larger;  skull  larger  and  relatively  broad- 
er in  most  dimensions;  rostrum  relatively 
broader  and  deeper,  especially  in  posterior 
half,  flaring  out  more  anterolaterally;  brain- 
case  relatively  smaller,  less  inflated  dorsally, 
never  broader  at  level  of  parietotemporal  su- 
tures than  at  base;  postorbital  constriction 
narrower,  more  elongated,  rising  more  steeply 
into  frontal  region;  zygomata  deeper,  thicker, 
more  broadly  flaring;  orbits  relatively  smaller; 
frontals  more  elevated  above  rostrum,  more 
depressed  medially,  more  prominendy  con- 
vex, forming  broader  shield;  temporal  ridges 
sharper,  more  often  obscuring  frontal  suture, 
and  joining  anterior,  rather  than  posterior,  to 
coronal  suture;  sagittal  crest  more  prominent; 
supraoccipital  shield  broader,  projecting  far- 
ther posteriorly;  external  side  of  occipital 
more  ossified,  seldom  with  any  trace  of  thin- 
walled  projection  dorsal  to  foramen  mag- 
num;   occipital    condyles    extending    farther 


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NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


95 


transversely;  tympanic  bullae  usually  less  in- 
flated; mandible  thicker  and  deeper,  ventral 
edge  less  convex  when  viewed  from  side, 
toothrow  more  bowed  outward  in  center; 
incisors  larger,  extending  farther  transverse- 
ly; upper  canines  thicker  anteroposteriorly, 
their  alveoli  set  more  dorsally  in  premaxillae, 
not  extending  so  far  ventrally;  premolars 
broader  with  less  trenchant  cusps,  usually 
more  closely  set  in  jaws;  P4  with  deuterocone 
and  lingual  cingulum  much  less  prominent  or 
absent;  Ml  having  relatively  larger  paracone 
and  metacone,  relatively  smaller  medial  sec- 
tion with  less  trenchant  cusps,  hypocone  less 
prominent,  metaconule  smaller  and  less  dis- 
tinct from  protocone,  protoconule  often  less 
distinct  from  protocone,  anterior  cingulum 
less  pronounced,  buccal  cingulum  less  pro- 
nounced or  absent;  M2  usually  relatively 
(and  occasionally  absolutely)  smaller  with 
less  trenchant  cusps;  p2  more  often  with  pos- 
terior cusp;  p4  with  posterior  cusps  more 
reduced,  more  often  lacking  well  developed 
third  cusp  and  posteromedial  cingulum;  ml 
relatively  broader,  metaconid  less  prominent 
and  not  projecting  so  far  medially,  heel  rela- 
tively smaller;  m2  and  talonid  of  ml  with 
less  trenchant  cusps.  Hildebrand  (1952a, 
1954)  discussed  differences  between  the  post- 
cranial  skeletons  of  C.  lupus  and  C.  latrans; 
Atkins  and  Dillon  ( 1971 )  listed  distinguish- 
ing features  of  the  cerebellum. 

Other  comparisons. — See  accounts  of  C. 
rufus,  C.  armbrusteri,  C.  familiaris,  and  C. 
dims. 

Remarks. — The  gray  wolf  is  probably  the 
most  widely  distributed  and  most  naturally 
successful  species  of  Canis  ever  to  exist.  Its 
size,  intelligence,  and  social  nature  are  singu- 
larly adapted  for  its  role  as  the  major  preda- 
tor of  northern  ungulates.  The  systematics  of 
the  species  have  long  been  a  source  of  con- 
fusion, and  are  still  not  completely  under- 
stood. Especially  difficult  problems  involve 
the  status  of  the  small  Recent  subspecies  or 
species  of  wolves  that  existed  all  along  the 
southern  margins  of  the  range  of  C.  lupus. 


Imaizumi  (1970a,  1970b)  recently  raised  the 
extinct  Japanese  wolf,  hodophilax,  back  to  the 
level  of  a  full  species.  The  wolf  of  China  and 
central  Asia  (chanco),  and  of  India  and  the 
Near  East  (pallipes)  are  probably  not  more 
than  subspecifically  distinct  from  C.  lupus, 
but  adequate  series  of  specimens  from  these 
vast  regions  never  have  been  studied  in  de- 
tail. On  the  basis  of  cranial  measurements 
provided  by  Pocock  (1935:671),  and  the  few 
specimens  that  I  have  seen,  pallipes  seems  to 
be  a  highly  variable  entity  with  a  cranial  size 
range  bridging  the  gap  between  North  Amer- 
ican C.  lupus  and  C.  latrans.  Even  more  in- 
teresting in  this  regard  is  arabs  of  southern 
Arabia,  of  which  the  cranial  measurements 
listed  by  Harrison  (1968:203)  indicate  an  ani- 
mal averaging  not  much  larger  than  C.  la- 
trans. Lawrence  (1966:57)  suggested  that 
arabs  may  have  been  influenced  by  hybridi- 
zation with  C.  familiaris.  Harrison  (1973: 
190),  however,  reported  that  all  available 
skulls  of  arabs  could  be  distinguished  by  the 
relatively  greater  size  and  inflation  of  their 
bullae.  Two  other  southern  subspecies  listed 
by  Ellerman  and  Morrison-Scott  (1951:218- 
220),  C.  lupus  italicus  of  Italy,  and  C.  I.  sig- 
natus  of  Spain,  had  been  synonymized  under 
C.  /.  lupus  of  most  of  Eurasia  by  Pocock 
(1935).  One  more  named  subspecies,  C.  I. 
deitanus,  was  based  only  on  two  live  animals 
from  southeastern  Spain.  Miller  (1912c:315) 
noted  that  they  had  a  "general  appearance 
much  as  in  C.  aureus."  On  the  basis  of  this 
description,  Pocock  (1935:653)  suggested  the 
possibility  that  deitanus  was  a  representative 
of  the  North  African  jackal.  The  question  ap- 
parently never  has  been  resolved. 

For  North  America  no  attempt  has  been 
made  to  go  beyond  previous  studies  in  assess- 
ing the  intraspecific  relationships  within  C. 
lupus.  All  of  the  names  and  their  areas  of 
application,  summarized  by  Hall  and  Kelson 
(1959:847-851),  are  maintained  in  this  paper. 
Some  comment,  however,  is  necessary  regard- 
ing a  confusing  situation  on  the  Arctic  is- 
lands. Anderson  (1943)  and  Goldman  (1944) 


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MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


considered  that  four  subspecies  inhabited  this 
region  (see  map,  Fig.  2).  Manning  and  Mac- 
pherson  (1958).  following  extensive  statistical 
analysis,  concluded  that  the  kind  of  wolf 
represented  by  a  series  of  eight  skulls  (in- 
cluding only  two  adults)  collected  in  1914- 
1916  on  Banks  Island,  and  described  as  C. 
lupus  bernardi  by  Anderson  (1943),  had  been 
replaced  by  a  different  kind  of  wolf,  repre- 
sented by  16  specimens  collected  in  1953- 
1955,  that  seemed  closest  to  C.  I.  arctos  of 
Prince  Patrick  and  Ellesmere  islands.  I  did 
not  measure  the  series  of  specimens  taken  on 
Banks  Island  in  1914-1916,  but  skulls  of  six 
males  and  two  females  collected  there  in 
1953-1955  were  suitable  for  inclusion  in  multi- 
variate analyses.  The  series  of  males  demon- 
strates a  consistently  high  statistical  distance 
from  each  subspecies  of  gray  wolf,  including 
arctos.  The  most  striking  character  of  the 
recently  collected  Banks  Island  skulls  is  their 
great  maximum  width  across  the  upper  cheek 
teeth.  In  all  but  one  of  these  specimens  this 
width  actually  exceeds  the  alveolar  length 
from  PI  to  M2.  In  most  other  skulls  of  C. 
lupus,  including  all  but  one  of  the  21  arctos 
that  I  measured,  the  length  was  greater  than 
the  width.  The  Banks  Island  skulls  also  differ 
from  arctos,  and  most  other  subspecies  of  C. 
lupus,  in  their  greater  width  of  frontal  shield. 
Although  I  agree  with  Manning  and  Mac- 
pherson  (1958:43)  that  the  more  recently 
collected  skulls  from  Banks  Island  differ  from 
Anderson's  description  of  C.  /.  bernardi,  I  am 
not  so  certain  that  these  specimens  may  be 
"assigned  to  C.  I.  arctos  with  confidence." 

The  most  critical  problem  that  concerned 
the  Recent  wolves  of  the  New  World  was  the 
relationship  of  C.  lupus  with  C.  rufus  of  the 
southeast.  Information  provided  in  the  pre- 
vious part  of  this  paper,  and  in  the  account 
of  C.  rufus  in  this  part,  has  to  me  confirmed 
the  specific  status  of  the  red  wolf.  Nonethe- 
less, the  paucity  of  available  material  from 
the  eastern  United  States  gives  an  incomplete 
picture  of  the  original  situation  in  that  region. 
In  the  above  account  of  C.  rufus,  I  discuss  a 


number  of  eastern  specimens  which  seem 
best  referred  to  that  species.  Various  other 
fragments  from  the  east,  including  many  from 
archeological  sites  and  not  listed  by  Gold- 
man (1944)  or  Hall  and  Kelson  (1959),  prob- 
ably represent  C.  lupus  hjcaon.  Such  speci- 
mens examined  by  me  (indicated  by  ele- 
ment), or  reported  by  others,  include  the 
following. 

Tick  Creek  Cave  site,  12  mi.  W  Rolla,  Phelps 
County,  Missouri;  as  C.  lupus  (Parmalee,  1965:19). 
Parmalee  also  reported  that  some  remains  from  this 
site  may  represent  C.  rufus. 

Bell  site,  5  mi.  W.  Oshgosh,  Winnebago  County, 
Wisconsin;  as  C.  lupus   (Parmalee,   1963:61). 

Raddatz  rock  shelter,  centra]  Sauk  County,  Wis- 
consin; as  C.  lupus  (Parmalee,  1959b:85);  maxillary 
fragment,  mandibular  fragment,  ISM. 

Moccasin  Bluff  site,  west  of  Buchanan,  Berrien 
County,  Michigan;  as  "wolf"  (Cleland,  1966:205). 

Anker  site,  Cook  County,  Illinois;  as  C.  lupus 
(Parmalee,  1959a:91);  two  maxillary  fragments,  pre- 
maxillary  fragment,  two  mandibular  fragments  (prob- 
ably all  from  same  individual),  ISM. 

Fisher  site,  south  bank  Des  Plaines  River,  Will 
County,  Illinois;  as  C.  lupus  (Parmalee,  1962b:402); 
mandibular  fragment,  ISM. 

Kingston  Lake  site,  15  mi.  SW  Peoria,  Peoria 
County,  Illinois;   as  C.   lupus   (Parmalee,   1962a:10). 

Hummel  Camp  site,  1  mi.  S  London  Mills,  Fulton 
County,  Illinois;  as  C.  nubilus  ( Cole  and  Deuel, 
1937:265). 

Weaver  site,  Fulton  County,  Illinois;  as  C.  lupus 
(Parmalee,  1959a:  91). 

Clear  Lake  site,  Tazewell  County,  Illinois;  as  C. 
lupus  (Parmalee,  1959a:91);  maxillary  fragment, 
mandibular  fragment,  ISM. 

Busch  Estate  site.  Pike  County,  Illinois;  as  C. 
lupus  (Parmalee,   1959a:91). 

Knight  site,  Calhoun  Countv,  Illinois;  as  C.  lupus 
(Parmalee,  1959a:91). 

Snyders  site,  Calhoun  County,  Illinois;  as  C. 
lupus    (Parmalee,   1959a: 91). 

Apple  Creek  site,  Greene  County,  Illinois;  man- 
dibular fragment,  ISM. 

Cahokia  site,  near  East  St.  Louis,  Madison  Coun- 
ty,  Illinois;   as   C.   lupus    (Parmalee,    1957:239). 

Palestine  site,  Palestine,  Crawford  County,  Illi- 
nois; as  C.  lupus  (Parmalee  and  Stephens,  1972:71); 
mandibular  fragment,  ISM. 

Sugar  Camp  Hill  site,  Williamson  County,  Illi- 
nois; as  C.  lupus  (Parmalee,  1959a:91). 

Fifield  site,  Porter  County,  Indiana;  as  "gray 
wolf?  Canis  lupus"    (Parmalee,   1972:205). 

Breck  Smith  Cave,  8  mi.  W  Lexington,  Fayette 
County,  Kentucky;  as  "wolf"   (Miller,  1922). 

Cirico  Mound,  near  Citico  Creek,  Hamilton 
County,   Tennessee;   as   C.   rufus  floridanus   (Kellogg, 


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NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


97 


1939:267),  as  C.  lupus  lycaon  (Goldman,  1944: 
441);  mandibular  fragment,  USNM. 

Madisonville  ancient  cemetery,  Cincinnati  vicin- 
ity, Hamilton  County,  Ohio;  as  C.  lupus  (Langdon, 
1881:299). 

Hobson  site,  near  Middleport,  Meigs  County, 
Ohio;   as  C.   lupus   (Murphy,   1968:12). 

Fairchance  Mound,  Moundsville,  Marshall  Coun- 
tv,  West  Virginia;  as  C.  cf.  lupus  (Guilday  and 
Tanner,  1966:42). 

Mount  Carbon  site,  3.5  mi.  SW  Montgomery, 
Fayette  County,  West  Virginia;  as  C.  lupus  (Guil- 
day and   Tanner,   1965:2);   ml,   CM. 

Doepkin's  Farm  site,  U.S.  Hwy.  50,  Anne  Arun- 
del County,  Maryland;  maxillary  fragment,  Doep- 
kin's  Farm   collection. 

Quaker  State  Rockshelter,  3  mi.  SE  Franklin, 
Venango  County,  Pennsylvania;  as  C.  lupus  (Guil- 
day  and  Tanner,    1962:134). 

Sheep  Rock  shelter,  west  bank  Raystown  branch 
Juanita  River,  Huntingdon  County,  Pennsylvania;  as 
C.  lupus   (Guilday  and  Parmalee,   1965:38). 

Johnston  site,  Indiana  County,  Pennsylvania; 
maxillary  fragment,  CM. 

Hartley  site,  Greene  County,  Pennsylvania;  man- 
dibular fragment,  CM. 

Eschelman  site,  3  mi.  S  Columbia,  Lancaster 
County,  Pennsylvania;  as  C.  lupus  (Guilday,  Parma- 
lee, and  Tanner,  1962:64);  three  mandibular  frag- 
ments, CM.  The  red  wolf,  C.  rufus,  is  also  repre- 
sented  by   material   from   this   site. 

Lewiston  Mound,  Lewiston,  Niagara  County, 
New  York;   as  "wolf"    (Ritchie,   1969:218). 

Garoga  site,  Fulton  County,  New  York;  maxillary 
fragment,   two   mandibular  fragments,   CM. 

Frontenac  Island,  Cayuga  Lake,  Cayuga  County, 
New  York;  as  C.  lupus  (Ritchie,  1969:106);  maxil- 
lary fragment,  CM. 

Lamoka  Lake  site,  near  Tyrone,  Schuyler  County, 
New  York;  as  C.  lupus  (Guilday,   1969:55). 

Sawyer's  Island,  near  Boothbay,  Lincoln  County, 
Maine;  as  C.  occidentalis  ( Loomis  and  Young,  1912: 
27). 

In  addition  to  the  above  records,  Man- 
ville  and  Sturtevant  ( 1966)  reported  the  pres- 
ence of  two  Indian  artifacts,  containing  parts 
of  wolf  skulls,  in  the  collection  of  the  Skok- 
Ioster  Castle  Museum  in  Sweden.  The  speci- 
mens had  probably  been  obtained  from  In- 
dians near  the  Swedish  colony  on  the  Dela- 
ware River,  or  the  Dutch  colony  on  the  Hud- 
son River.  The  material  was  identified  as  C. 
lupus  lycaon,  and  the  measurements  provided 
indicate  that  the  gray  wolf,  rather  than  the 
red  wolf,  is  represented. 

Whereas  there  is  sometimes  difficulty  in 
distinguishing  specimens  of  C.  lupus  and  C. 


rufus,  cranial  material  of  C.  lupus  and  C.  la- 
trans  can  always  be  separated.  The  clear  dis- 
tinction of  the  two  species  was  recognized  by 
American  taxonomists  at  least  as  early  as 
Audubon  and  Bachman  (1851).  Baird  (1857: 
104)  adequately  described  some  of  the  major 
cranial  differences  between  the  gray  wolf  and 
coyote.  Cope  (1879:184)  was  the  first  to 
point  out  the  discriminating  features  of  the 
cusps  on  the  medial  section  of  Ml.  Gidley 
(1913:98-102)  listed  what  he  considered  to 
be  diagnostic  characters  of  the  lower  denti- 
tion, but,  as  explained  by  Jackson  (1951:242), 
these  characters  are  not  always  reliable.  The 
most  thorough  discussion  of  the  differences 
in  proportions  and  other  characters,  between 
the  skulls  of  C.  lupus  and  C.  latrans,  was  that 
provided  by  Lawrence  and  Bossert  ( 1967 ) . 

The  coyote  and  gray  wolf  shared  a  large 
part  of  their  respective  ranges  in  North  Amer- 
ica, but  hybridization  under  completely  nat- 
ural conditions  occurred  rarely,  if  ever.  In- 
terbreeding in  eastern  Canada,  caused  largely 
by  recent  human  environmental  disruption, 
has  resulted  in  the  production  of  some  speci- 
mens with  intermediate  characters. 

Fossils  of  wolves,  other  than  C.  dims,  are 
comparatively  rare  in  North  America,  and  it 
sometimes  is  difficult  to  determine  what  spe- 
cies are  represented.  Martin  (1974:76)  con- 
sidered C.  armbrusteri  of  Maryland  and  Flor- 
ida to  be  synonymous  with  C.  lupus,  but,  as 
explained  previously,  the  two  are  distinct. 
One  named  Pleistocene  species  that  now  can 
be  synonymized  with  a  subspecies  of  C.  lupus 
is  C.  (Aenocyon)  milleri  Merriam  from 
Rancho  La  Brea.  The  single  specimen  on 
which  the  species  was  based  falls  within  the 
morphological  range  of  C.  lupus,  and  seems 
best  referred  to  C.  lupus  furlongi  (see  ac- 
count of  Rancho  La  Brea,  below). 

Fossil  record. — Most  fossil  wolf  material 
is  so  fragmentary  that  determination  as  to 
species  is  difficult.  The  first  specimens  that 
show  the  specific  characters  of  C.  lupus  ap- 
pear in  Illinoian  deposits.  Several  pre-Illi- 
noian  fragments  that  had  been   referred  by 


98 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


Fig.  51. — Map  showing  localities   (black  dots)   of  fossil  C.  lupus.    Because  of  the  scale  of  the  map,  it  was 
not  possible  to  plot  all  localities  in  crowded  areas. 


others  to  this  species,  are  discussed  above  in 
the  accounts  of  C.  edwardii,  C.  rufus,  and  C. 
armbrusteri.  The  following  list  contains  ad- 
ditional literary  references  to  fossil  C.  lupus. 


The  list  is  arranged  alphabetically  by  state 
and  province,  and  geographically  (north  to 
south,  west  to  east)  within  states  and  prov- 
inces,   except    that    Canadian    provinces    are 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


99 


listed  first  and  Mexican  states  last.  Specimens 
examined  by  me  are  identified  by  element, 
museum  number,  or  both;  and  selected  meas- 
urements are  found  in  appendix  B  (part  14) 
and  appendix  C  (part  6).  Occurrences  also 
are  shown  on  the  map  in  figure  51. 

ALBERTA— Medicine  Hat  (north  of);  early  Re- 
cent;  as   C.   lupus    (Churcher,   1969b:  181). 

Medicine  Hat;  Sangamon;  as  C.  lupus  (Churcher, 
1970:62). 

Island  Bluff,  near  Medicine  Hat;  Sangamon;  as 
C.    lupus    (Churcher,    1969b:  181). 

Mitchell  Bluff,  near  Medicine  Hat;  late  Pleisto- 
cene;  as  C.   lupus   (Churcher,   1969a :2). 

SASKATCHEWAN— Fort  Qu'Apelle;  late  Pleis- 
tocene; tibia,  CNM  12178  (see  also  Khan,  1970:13). 

YUKON. — Old  Crow  area;  late  Pleistocene;  as 
"wolf"  (Geist,  1955:1702);  mandibular  fragment, 
CNM  17311. 

Hunker  Creek  vicinity,  Klondike  River;  late  Pleis- 
tocene; as  "Canis  (wolf)"  (Quackenbush,  1909:127); 
skull  without  mandibles,  CNM  9929.  The  specimen 
shares  certain  characters  with  Recent  wolves  of  the 
Arctic  islands,  including  a  broad  rostrum,  crowded 
toothrow,  and  relatively  large  carnassials. 

Gold  Run  Creek,  30  mi.  SE  Dawson;  Wisconsin 
(C-14  dates:  22,200  and  32,250  B.P.);  as  C.  cf. 
lupus   (Harington  and  Clulow,   1973:699). 

Quartz  Creek;  late  Pleistocene;  cranial  fragment, 
CNM  17311. 

ALASKA. — Historic  Bluff,  entrance  to  Esch- 
scholtz  Bay;  late  Pleistocene;  as  "Canis  (wolf)" 
(Quackenbush,  1909:97);  maxillary  fragment  and 
other  cranial  elements,  AMNH  13753. 

Buckland  River,  southeast  of  Eschscholtz  Bay; 
late  Pleistocene;  as  "Canis  (wolf)"  (Quackenbush, 
1909:120). 

Fairbanks  (near);  Illinoian;  as  "Canis  sp.  (wolf)" 
(Pewe    and    Hopkins,    1967:267). 

Fairbanks  Creek  Mine,  near  Fairbanks;  Wiscon- 
sin; as  C.  lupus  (Guthrie,  1968:352). 

Engineer  Creek  Mine,  near  Fairbanks;  Wiscon- 
sin; as  C.  lupus  (Guthrie,  1968:352). 

Gold  Hill  Mine,  near  Fairbanks;  Wisconsin;  as 
C.    lupus    (Guthrie,    1968:352). 

Cripple  Creek  Mine,  near  Fairbanks;  Wisconsin; 
as  C.  lupus  (Guthrie,  1968:352). 

ARIZONA. — Ventana  Cave,  Papago  Indian  Res- 
ervation, Pima  County;  Wisconsin  (Hibbard,  1958); 
as  C.  lupus  (Colbert,  1950:132). 

Papago  Springs  Cave,  southeast  of  Sonoita,  Santa 
Cruz  County;  Wisconsin  (Hibbard,  1958);  as  C. 
nubilus    (Skinner,    1942:164). 

ARKANSAS. — Conard  fissure,  15  mi.  S  Harrison, 
Newton  County;  Illinoian  ( Kurten,  1963:100);  as 
"C.  occidentalis?"  (Brown,  1908:182);  cranial  frag- 
ment, isolated  teeth,  postcranial  fragments,  AMNH 
11762;  mandibular  fragment,  AMNH   11761. 

CALIFORNIA.— Samw  el  Cave,  Shasta  Lake, 
Shasta    County;    Wisconsin     (Hibbard,    1958);    "the 


specimen  appears  to  resemble  the  northern  wolves; 
for  example  C.  /.  pambasileus,  rather  than  specimens 
from  the  southern  part  of  the  range  of  C.  lupus" 
(Graham,  1959:58). 

Potter  Creek  Cave,  1  mi.  SE  Baird,  Shasta  Coun- 
ty; Wisconsin  (Hibbard,  1958);  as  C.  lupus  (Kurten 
and  Anderson,  1972:37);  mandibular  fragment, 
UCMP  5018. 

McKittrick  tar  seeps,  Kern  County;  Wisconsin; 
as  "gray  or  timber  wolf"  (Sternberg,  1928:226),  as 
Aenocijon  near  mitteri  ( Schultz,  1938b:169).  Stern- 
berg reported  that  he  had  collected  ten  specimens 
of  gray  wolves  at  McKittrick,  but  Schultz  did  not 
mention  the  presence  of  C.  lupus  or  C.  furlongi  at 
the  site.  Schultz  did  note  that  two  Ml  seemed  best 
referred  to  Aenocyon  milleri,  which  in  this  paper  is 
considered  a  synonym  of  C.  lupus  furlongi. 

Maricopa  Brea,  near  Maricopa,  Kern  County; 
Wisconsin;  skull  and  mandible,  LACM  18419;  skull 
without  mandibles,  LACM  21921;  maxillary  and 
mandibular  fragments,  LACM  18798;  three  maxillary 
fragments,  LACM  20531  and  two  unnumbered; 
cranial  fragment,  LACM;  two  mandibular  fragments 
LACM  17890,  22288;  Ml,  LACM.  These  specimens 
were  identified  by  me  from  among  a  larger  number 
of  specimens  of  C.  dirus  in  the  hitherto  unreported 
Maricopa  Brea  collection. 

Rancho  La  Brea,  Los  Angeles,  Los  Angeles  Coun- 
ty; Wisconsin;  as  C.  occidentalis  furlongi  (Merriam 
1910:393;  1912:251;  Schultz,  1938b:163;  Goldman! 
1944:399;  Anderson,  1968:26),  as  C.  furlongi  (Hay, 
1927:184;  Stock,  1956:33),  as  C.  lupus  (Hibbard, 
1958:18);  eight  skulls  without  mandibles,  LACM 
2300-44,  2300-56,  2300-353,  2300-384,  2600-1, 
2600-5,  236(315),  one  unnumbered;  cranial  frag- 
ment, LACM;  two  mandibles,  LACM  2301-L476, 
2301-L495;  incomplete  skull,  UCMP  19792;  maxil- 
lary fragments,  UCMP  10733;  maxillary  fragment, 
mandibular  fragment,  UCMP  11283.  Merriam  (1910) 
considered  that  certain  specimens  from  Rancho  La 
Brea  represented  an  animal  closely  related  to  the 
modern  gray  wolf.  His  name  for  this  animal,  C. 
occidentalis  furlongi,  indicated  his  recognition  of  it 
as  a  subspecies  of  the  North  American  gray  wolf 
which  was  then  (1910)  often  referred  to  as  C.  occi- 
dentalis. Miller  (1912b),  however,  restricted  the 
name  occidentalis  to  the  interior  forests  of  northern 
Canada,  and  Hay  (1927:184)  considered  it  improb- 
able that  a  subspecies  of  occidentalis  ever  would 
have  been  present  in  southern  California.  Subse- 
quently, various  authors  either  followed  Hay  in  list- 
ing furlongi  as  a  full  species,  or  continued  to  use  the 
trinomial  C.  occidentalis  furlongi.  Since  all  of  the 
Recent  gray  wolves  of  North  America,  including 
occidentalis,  were  arranged  as  subspecies  of  C.  lupus 
by  Goldman  (1944),  and  since  Merriam's  original 
intention  was  obviously  to  recognize  furlongi  as  a 
subspecies  of  gray  wolf,  the  proper  name  for  the 
animal  in  question  would  be  C.  lupus  furlongi.  Mer- 
riam (1910,  1912)  based  his  descriptions  of  furlongi 
on  three  fragmentary  specimens  in  the  University  of 
California    Museum.     The    material   was    said   to    be 


100 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


smaller  than  that  of  the  much  more  abundant  C. 
dims  from  Rancho  La  Brea,  and  to  be  characterized 
by  a  more  prominent  hypocone  on  Ml.  Subsequently, 
Stock  (1929:20)  reported  that  20  individuals  of 
furlongi  from  Rancho  La  Brea  were  represented  in 
the  collections  of  the  Los  Angeles  County  Museum, 
but  later  (1956:33)  he  wrote  that  "only  eight  speci- 
mens have  been  recognized  in  the  Museum  collec- 
tions." Considering  that  taxonomists  of  the  early 
twentieth  century  sometimes  named  species  on  the 
basis  of  less  critical  analysis  than  is  usual  today,  it 
might  be  tempting  to  write  off  the  few  specimens  of 
furlongi  as  small,  aberrant  examples  of  C.  dims  (of 
which  1,646  individuals  were  reported  to  be  repre- 
sented in  the  LACM  collections  from  Rancho  La 
Brea  by  Marcus,  1960:5).  There  is  no  question, 
however,  that  C.  lupus  is  also  present.  The  speci- 
mens of  C.  dints  from  Rancho  La  Brea  are  remark- 
ably consistent  in  certain  critical  characters,  and 
while  going  through  the  unlabeled  collection  of 
wolves  at  the  Los  Angeles  County  Museum,  I  found 
the  eight  above  listed  skulls  of  C.  lupus  to  stand  out 
clearly  from  the  others.  I  think  that  Stock  (1956: 
33)  must  have  been  referring  to  the  same  eight 
specimens.  Each  of  these  skulls  is  from  an  adult 
animal,  and  has  each  of  the  following  characters  for 
which  it  can  be  evaluated:  f rentals  depressed  me- 
dially; temporal  ridges  sharp;  orbital  angle  under 
47°;  supraoccipital  shield  broad,  not  projecting  far 
posteriorly;  postpalatine  foramina  placed  well  anterior 
to  posterior  edges  of  P4;  optic  foramen  and  anterior 
lacerated  foramen  well  separated;  vertical  plates  of 
palatines  not  broadly  flaring  anteriorly;  posterior  end 
of  vomer  extending  well  behind  posterior  nasal  open- 
ing; Ml  with  large  hypocone,  its  ridge  extending 
completely  or  almost  completely  around  anterior 
base  of  protocone.  These  characters,  along  with 
over-all  moderate  size,  distinguish  the  LACM  speci- 
mens, as  well  as  those  in  the  UCMP,  from  C.  dirus. 
In  addition  to  these  skulls,  I  found  a  cranial  frag- 
ment having  the  characters  of  C.  lupus,  and  two 
mandibles  characterized  by  small  size,  a  prominent 
posterior  cusp  on  p2,  no  posterior  cingulum  on  p4, 
and  a  relatively  high  set  heel  on  ml.  Most  of  the 
specimens  of  furlongi  have  comparatively  large  teeth, 
especially  carnassials,  a  broad  rostrum,  and  a  rela- 
tively broad  frontal  shield.  These  characters  are 
shared  by  C.  dirus,  and  might  suggest  that  furlongi 
represents  an  evolutionary  transition  between  C. 
dims  and  C.  lupus.  But  there  is  no  chronological  evi- 
dence to  support  this  view;  material  of  both  species 
was  found  together  in  the  same  pits  at  Rancho  La 
Brea.  Furthermore,  in  the  great  majority  of  char- 
acters, the  material  from  this  site  shows  no  tendency 
toward  blending;  each  specimen  can  be  unquestion- 
ably referred  to  either  C.  lupus  or  C.  dims.  Large 
carnassials,  and  a  broad  rostrum  and  frontal  shield 
are  also  present  in  specimens  of  modern  gray  wolves 
of  the  Arctic  islands  (see  "remarks,"  above).  Mac- 
pherson  (1965:164)  hypothesized  that  wolves  with 
such  characters  had  been  isolated  by  late  Pleistocene 
glaciation  in   a  Pearyland  refugium,   and  had  subse- 


quently spread  back  across  the  Arctic.  Possibly  this 
population  had  once  occupied  a  large  northern  area, 
and  had  then  been  driven  by  the  Wisconsin  glacia- 
tion both  northeastward  into  Pearyland,  and  south- 
westward  as  far  as  Rancho  La  Brea.  Not  all  speci- 
mens of  C.  lupus  from  this  site  are  as  massive  as 
Recent  Arctic  wolves,  and,  as  noted  by  Merriam 
(1912:253)  there  is  considerable  variation  in  size 
of  teeth. 

Rancho  La  Brea,  Los  Angeles,  Los  Angeles  Coun- 
ty; Wisconsin;  as  C.  milleri  (Merriam,  1912:24),  as 
Aenocyon  milleri  (Merriam,  1918:533),  as  C.  lupus 
(Martin,  1974:76);  skull  with  mandibles,  UCMP 
11257.  According  to  Merriam's  original  description, 
the  single  known  specimen  from  Rancho  La  Brea  is 
intermediate  in  characters  between  C.  lupus  and 
C.  dims.  He  compared  the  specimen  most  critically 
with  C.  dirus,  from  which  it  was  said  to  differ  in 
having  lesser  size,  a  smaller  frontal  shield,  lower 
sagittal  crest,  less  overhang  of  inion,  more  anteriorly 
placed  postpalatine  foramina,  and  more  prominent 
hypocone  on  Ml.  Merriam  reported  milleri  to  have 
a  much  broader  palate  and  much  more  massive 
dentition  than  C.  lupus.  Later  (1918)  he  observed 
that  the  characters  of  milleri  justified  placing  it  to- 
gether with  dims  in  the  new  genus  Aenocyon.  Sub- 
sequently, according  to  Stock,  Lance,  and  Nigra 
(1946:109),  the  validity  of  milleri  as  a  species  was 
questioned,  but  they  did  not  indicate  whether  refer- 
ral to  C.  dirus  or  C.  lupus  was  being  considered. 
Martin  (1974:76)  recognized  C.  milleri  as  a  synonym 
of  C.  lupus.  Had  Merriam  been  able  to  examine  the 
eight  skulls  of  C.  lupus  furlongi  from  Rancho  La 
Brea  in  the  Los  Angeles  County  Museum,  and  a 
series  of  Recent  C.  lupus  from  the  Arctic  islands,  he 
might  not  have  established  milleri  as  a  separate 
species.  As  he  himself  observed  (1912:247),  the 
combination  of  characters  found  in  milleri  is  ap- 
proached most  closely  in  C.  I.  furlongi,  and  several 
of  the  LACM  specimens  are  almost  identical  to 
milleri.  In  its  relatively  large  carnassials  and  unusu- 
ally broad  rostrum,  milleri  resembles  some  of  the 
living  wolves  of  the  Arctic.  Merriam's  (1918)  asso- 
ciation of  the  specimen  with  the  genus  Aenocyon 
was  a  mistake,  as  the  specimen  differs  from  the 
dire  wolf  in  all  of  the  critical  characters  mentioned 
above  in  the  discussion  of  C.   lupus  furlongi. 

Schuiling  Cave,  2  mi.  SE  Newberry,  San  Ber- 
nardino County;  late  Pleistocene;  as  C.  cf.  lupus 
(Downs,  ct  al.,   1959:9). 

COLORADO.— Chimney  Rock  animal  trap,  Lari- 
mer County;  late  Pleistocene  or  early  Recent;  as 
C.   lupus    (Hager,   1972:65). 

GEORGIA.— Ladds,  near  Cartersville,  Bartow 
Countv;  late  Pleistocene;  as  C.  cf.  lupus  ( Rav,  1967: 
133);  Ml,  USNM  23698. 

IDAHO. — Jaguar  Cave,  Beaverhead  Mountains, 
Lemhi  Countv;  late  Wisconsin  (C-14  dates:  10,370 
±350  and  11,580±250  B.P.);  as  C.  lupus  (Kurten 
and  Anderson,   1972:24). 

Moonshiner  Cave,  Bingham  County;  late  Wiscon- 
sin or  early  Recent;  as  C.  lupus  ( Kurten  and  Ander- 
son, 1972:37). 


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101 


ILLINOIS.— Polecat  Creek  gravel  pits,  1  mi.  S 
Ashmore,  Coles  County;  late  Wisconsin  ( Hibbard, 
et  al,  1965);  as  C.  lupus  (Galbreath,  1938:309). 

KANSAS. — Goodland,  Sherman  County;  late 
Pleistocene  or  early  Recent;  as  C.  lupus  (Williston, 
1898:93;  Hibbard,  Frye,  and  Leonard,  1944:10); 
skull  with  mandibles,  KU  2851.  The  specimen  re- 
sembles skulls  of  C.  lupus  baileyi  from  the  south- 
west, and  is  smaller  than  skulls  of  C.  I.  nubilus  col- 
lected on  the  Great  Plains  in  historical  time. 

MICHIGAN.— Millington,  Tuscola  County;  late 
Wisconsin;  as  C.  lupus  (Wilson,  1967:211);  pair  of 
mandibles,   upper  incisors,   UMMP  33770. 

MINNESOTA.— Itasca  bison  site,  Clearwater 
County;  late  Pleistocene;  as  "wolf"  (Shay,  1963:48). 

MISSOURI.— Brynjulfson  Caves,  6  mi.  SSE  Co- 
lumbia, Boone  County;  late  Wisconsin  (about  10,000 
B.P.);  as  C.  lupus   (Parmalee  and  Oesch,   1972:29). 

NEBRASKA.— Hay  Springs  quarry,  Sheridan 
County;  probably  Illinoian  (Hibbard,  1958);  as  C. 
cf.  Occident alis  (Matthew,  1918;  Schultz,  1934:369); 
rostral  fragment,  AMNH  (Frick  Collection)  25511; 
mandibular  fragment,  UN  2912. 

Mullen,  Cherry  County;  late  Irvingtonian  (Kur- 
ten,  1974:7);  as  Aenocyon  dims  (Martin,  1972:174); 
maxillary  fragment,  UN  39337;  mandibular  frag- 
ment, UN  26117.  In  size  and  other  characters,  these 
specimens  resemble  C.  lupus,  not  C.  dims. 

Freedom  (near),  Frontier  County;  late  Pleisto- 
cene;   mandible,    UN   2911. 

Republican  River,  1  mi.  S  Guide  Rock,  Webster 
County;  late  Pleistocene  or  early  Recent;  cranial 
fragments,   USNM   18749. 

NEVADA. — Lake  Lahontan,  near  Fallon,  Church- 
ill County;  early  Recent;  as  C.  lupus  (Morrison, 
1964:73). 

Smith  Creek  Cave,  Baker,  White  Pine  County; 
late   Pleistocene;    cranial   fragment,    LACM   7190. 

NEW  MEXICO.— Isleta  Caves,  8  mi.  W  Isleta, 
Bernalillo  County;  late  Wisconsin;  as  C.  cf.  lupus 
(Harris  and  Findley,  1964:115),  as  C.  lupus  (An- 
derson, 1968:22). 

Blackwater  Draw,  near  Clovis,  Curry  County; 
Wisconsin  ( Lundelius,  1967:301);  crushed  skull, 
TM  937-521;  mandible,  TM  937-895;  two  Ml,  TM 
937-885,   937-905. 

Burnet  Cave,  50  mi.  W  Carlsbad,  Eddy  County; 
Wisconsin  (Hibbard,  1958);  as  C.  nubilus  (Schultz 
and  Howard,   1935:284);   mandible,  UN   14004. 

Hermit's  Cave,  east  slope  of  Guadalupe  Moun- 
tains, Eddy  County;  late  Wisconsin  (C-14  dates: 
11,850±350  and  12,900±350  B.P.;  Schultz,  Martin, 
and  Tanner,  1970:119);  two  maxillary  fragments  and 
two  mandibular  fragments  (probably  all  from  same 
individual),  UN  19211,  19217,  19218,  19220;  pair 
of  mandibles,  UN  19216.  These  specimens  were 
associated  with  a  man-made  hearth  from  which  the 
C-14  dates  were  obtained.  At  least  one  of  the  frag- 
ments is  charred,  as  from  fire,  and  the  single  skull, 
apparently  represented  by  the  first  four  fragments 
listed  above,  may  have  been  deliberately  broken 
apart.      Schultz,     Martin,     and    Tanner     (1970)     re- 


corded only  C.  dims  from  Hermit's  Cave,  and  that 
species  is  present,  but  the  five  specimens  listed  above 
unquestionably   represent   C.    lupus. 

Dark  Canyon  Cave,  Eddy  County;  late  Pleisto- 
cene;  mandible,   LACM   1644. 

OKLAHOMA.— Selman  Cave  system,  7  mi.  SW 
Freedom,  Woodward  County;  Recent,  as  C.  lupus 
(Black  and  Best,  1972);  mandible,  collection  of 
Troy  L.  Best. 

Afton,  Ottawa  County;  Wisconsin  (Kurten,  1974: 
9);  as  C.  nubilus  (Hay,  1920:129);  skull  with  man- 
dibles, USNM  196943;  three  mandibles,  USNM 
196946,  196947,  196948;  P4,  CI,  USNM  9128;  eight 
canine  teeth,  USNM  9129;  premaxillary  fragment, 
P4,   Ml,  p4,   USNM  9130. 

Arkansas  River,  Le  Flore  County;  Pleistocene; 
cranial   fragment,   AMNH   32669. 

OREGON.— Bend  (near),  Deschutes  County; 
late  Pleistocene;  tibia,  CNM  12178. 

Fossil  Lake,  Lake  County;  early  or  middle  Wis- 
consin (Allison,  1966:32);  as  C.  cf.  occidentalis 
(Elftman,    1931:7). 

PENNSYLVANIA.— Crystal  Hill  Cave,  3  mi.  W 
Stroudsburg,  Monroe  County;  late  Pleistocene  or 
early  Recent;  as  C.  lupus  (Leidy,  1889;  Hay,  1923- 
310). 

TEXAS. — Lubbock  Reservoir,  Lubbock  County; 
Wisconsin  (Lundelius,  1967:302);  mandible,  TM 
892-255. 

Schulze  Cave,  28  mi.  NE  Rock  Springs,  Edwards 
County;  Wisconsin  or  early  Recent;  as  C.  cf.  lupus 
(Dalquest,    Roth,   and   Judd,    1969:256). 

VIRGINIA.— Natural  Chimneys,  1  mi.  N  Mt. 
Solon,  Augusta  County;  late  Wisconsin  (ca.  10,000- 
15,000  B.P.);  as  C.  cf.  lupus  (Guilday,  1962:94). 

WISCONSIN.— Blue  Mounds,  Dane  County;  late 
Pleistocene;  as  C.  occidentalis  (Hay,  1918:347-  1923- 
341). 

WYOMING.— Little  Box  Elder  Cave,  west  of 
Douglas,  Converse  County;  Wisconsin;  as  C.  lupus 
(Anderson,  1968:25);  two  mandibles  from  subadult 
individual,   UColo  22287,  24683. 

Bell  Cave,  Albany  County;  Wisconsin  to  early 
Recent;  as  C.  lupus  (Anderson,  1974:81). 

NUEVO  LEON. — San  Josecito  Cave,  near  Aram- 
berri;  Wisconsin;  skull  without  mandibles,  LACM 
192-3017;  mandibular  fragment,  LACM  192-28338. 
In  the  large  collection  of  canid  material  from  San 
Josecito  Cave,  the  gray  wolf  is  represented  only  by 
these  two  specimens  which  probably  belonged  to 
the  same  individual.  The  skull  is  much  smaller  than 
those  of  C.  dims  from  the  same  site,  and,  indeed,  is 
the  smallest  skull  of  an  adult  C.  lupus  that  I  have 
examined.  I  am  indebted  to  Richard  L.  Reynolds 
of  the  Los  Angeles  County  Museum  for  recognizing 
the  presence  of  these  specimens  in  the  collection 
from  San  Josecito  Cave,  and  for  loaning  them  to  me. 

Evolutionary  position. — The  North  Amer- 
ican gray  wolf,  like  many  of  our  other  larger 
mammals,  appears  to  be  a  comparatively  late 
immigrant    from    the    Old    World.     Kurten 


102 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


(1968:109-110)  traced  the  probable  evolution 
of  the  species  in  Europe  from  a  relatively 
small  ancestor.  The  primitive  stock  that  gave 
rise  to  C.  lupus  in  the  Old  World  was  in  all 
likelihood  the  same  that  is  represented  in 
North  America  by  C.  edwardii  and  early  spec- 
imens of  C.  rufus.  At  various  times  in  the 
Pleistocene,  factors  associated  with  glaciation 
presumably  divided  this  widespread  basic 
stock,  and  permitted  development  of  several 
species.  While  C.  rufus,  C.  armbrusteri,  and 
C.  dims  evolved  in  the  New  World,  C.  lupus 
arose  in  Eurasia  and  apparently  became  the 
species  of  wolf  most  suited  for  the  challeng- 
ing environment  of  the  late  Quaternary.  We 
do  not  know  at  what  point  the  gray  wolf 
completed  its  occupation  of  Eurasia  and  be- 
gan to  move  across  the  Bering  Strait,  but  con- 
ceivably this  could  have  been  as  early  as  the 
Kansan  glaciation.  A  few  pre-Illinpian  frag- 
ments had  been  questionably  assigned  to  C. 
lupus  by  previous  authors,  but  at  present 
these  specimens  seem  best  referred  to  C.  ru- 
fus or  C.  armbrusteri.  The  earliest  material 
that  clearly  displays  the  specific  characters  of 
C.  lupus  is  that  from  the  Illinoian  deposits  at 
Hay  Springs  and  Mullen  in  Nebraska.  Illi- 
noian specimens  also  have  been  reported  from 
the  Conard  fissure,  Arkansas,  and  from  near 
Fairbanks,  Alaska.  It  can  be  reasonably  as- 
sumed that  the  species  was  able  to  cross  the 
Bering  Land  Bridge  in  the  Illinoian,  and 
that  it  eventually  established  itself  in  some 
parts  of  North  America.  Glacial  movement 
may  have  been  responsible  for  the  initial  ap- 
pearance of  C.  lupus  in  the  central  United 
States. 

The  only  known  Sangamon  records  of  the 
species  in  North  America  are  based  on  a  few 
fragments  collected  near  Medicine  Hat,  Al- 
berta (Churcher,  1969b:lSl;  1970:62).  Pos- 
sibly the  cold-adapted  C.  lupus  had  with- 
drawn from  more  southerly  regions  during 
that  interglacial  period.  The  number  and 
distribution  of  Wisconsin  records  is  much 
greater,  but  the  fossil  history  of  the  species  is 
comparatively  poor,  and  few  occurrences  are 


represented    by    well    preserved    cranial   ma- 
terial. 

So  fragmentary  the  material,  so  variable 
the  existing  gray  wolf,  and  so  incomplete  our 
understanding  of  Pleistocene  chronology,  that 
it  is  difficult  to  assess  the  factors  leading  to 
the  present  situation.  A  number  of  the  speci- 
mens discussed  in  the  above  list  do  not  ap- 
pear to  differ  significantly  from  specimens  of 
C.  lupus  taken  in  the  same  areas  in  historic 
time.  The  most  interesting  Wisconsin  speci- 
mens are  the  massive  skulls  from  Rancho  La 
Brea  and  the  Yukon,  that  resemble  the  skulls 
of  modern  Arctic  wolves.  A  population  of 
wolves  with  such  skulls  may  have  been  wide- 
spread in  the  north  at  the  beginning  of  the 
Wisconsin,  or,  more  likely,  at  the  beginning 
of  the  last  major  stadial.  The  glacial  move- 
ment may  then  have  split  the  population,  one 
element  moving  into  the  Pearyland  refugium 
where  it  survived,  and  the  other  element  be- 
ing driven  southward  where  it  either  became 
extinct  or  was  eventually  absorbed  into  other 
populations  of  C.  lupus.  A  few  of  the  speci- 
mens from  Rancho  La  Brea  are  small,  and 
there  are  also  unusually  small  skulls  of  C. 
lupus  from  Goodland,  Kansas  and  San  Jose- 
cito  Cave,  Nuevo  Leon.  These  specimens  may 
represent  the  result  of  character  displacement 
following  an  initial  late  Pleistocene  or  early 
Recent  movement  of  C.  lupus  into  areas 
where  C.  dims  still  predominated.  Eventu- 
ally, C.  lupus  prevailed  over  the  dire  wolf, 
either  through  competition  or  because  of  ex- 
ternal factors,  and  established  itself  as  the 
major  large  predator  of  most  of  North  Amer- 
ica. 

Canis  familiaris  Linnaeus 

175S.  Canis  familiaris  Linnaeus,  Systema 
Naturae,  10th  ed.,  p.  38. 

1938.  Canis  petrolei  Stock,  Bull.  S.  California 
Acad.  Sci.,  37:50.  Type  from  Rancho 
La  Brea,  Los  Angeles  County,  Califor- 
nia. The  type  and  only  known  speci- 
men, originally  described  by  Stock  as 
"a  coyote-like  wolf  jaw,"  probably  rep- 


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NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


103 


resents  C.  familiaris.  A  study  on  this 
and  other  specimens  of  early  domestic 
dogs  from  Rancho  La  Brea  is  being 
made  by  Richard  L.  Reynolds  of  the 
Los  Angeles  County  Museum. 

Type. — None  designated. 

Geological  distribution. — Late  Ranchola- 
brean  to  Recent. 

Geographical  distribution. — World-wide  in 
association  with  man. 

Description. — Exceptionally  variable  in 
size  and  other  characters  because  of  the  in- 
fluence of  domestication.  Less  specialized 
breeds  characterized  as  follows:  moderate 
size;  skull  medium-sized,  relatively  broad  in 
most  proportions;  rostrum  usually  relatively 
short,  broad,  and  deep;  braincase  relatively 
small,  not  much  inflated  dorosposteriorly, 
broadly  based,  set  low  relative  to  other  parts 
of  skull;  postorbital  constriction  usually  elon- 
gated, broad  lateromedially,  rising  very 
steeply  into  frontal  region;  zygomata  rela- 
tively thick,  deep,  moderately  flaring;  orbits 
relatively  small;  frontals  rising  steeply  above 
rostrum,  prominently  convex,  forming  broad 
shield  with  bulging  postorbital  processes; 
temporal  ridges  usually  sharp,  often  obscur- 
ing frontal  suture,  often  joining  anterior  to 
coronal  suture;  sagittal  crest  sometimes  prom- 
inent; supraoccipital  shield  small,  not  project- 
ing far  posteriorly;  external  side  of  occipital 
well  ossified;  tympanic  bullae  usually  small, 
not  much  inflated;  medial  part  of  posterior 
margin  of  palate  often  extending  well  behind 
toothrow;  mandible  thick,  deep,  ventral  edge 
often  convex  when  viewed  from  side,  tooth- 
row  bowed  out  prominently  in  center;  ascend- 
ing ramus  sometimes  curving  dorsoposterior- 
ly;  teeth  relatively  small,  usually  lacking 
trenchant  cusps,  usually  widely  spaced  in 
jaws;  upper  canines  thick  anteroposteriorly, 
short  dorsoventrally;  P4  usually  lacking  prom- 
inent deuterocone  and  lingual  cingulum;  Ml 
having  relatively  large  paracone  and  meta- 
cone,  relatively  small  medial  section,  usually 
lacking  buccal  cingulum;   M2  usually  small; 


p2  and  p3  sometimes  with  posterior  cusps; 
p4  with  second  cusp,  sometimes  lacking  third 
cusp,  usually  without  posteromedial  cingulum 
extending  behind  third  cusp;  ml  relatively 
broad,  usually  with  relatively  small  talonid; 
m3  occasionally  absent.  For  details  on  pelage 
and  postcranial  skeleton  see  Haag  (1948); 
Iljin  (1941);  Miller,  Christensen,  and  Evans 
(1965);  and  Scott  and  Fuller  (1965). 

Comparison  with  C.  latrans. — Often  close 
in    size;    skull   usually    relatively   broader    in 
most  dimensions;   rostrum  relatively  broader 
and  deeper  ( a  few  specialized  breeds,  such  as 
collies    and    Russian    wolfhounds,    may   have 
relatively    longer    and    narrower    rostra   than 
coyotes);    braincase    relatively    smaller,    less 
inflated    dorsoposteriorly,    broader    ventrally, 
set  lower  relative  to  other  parts  of  skull  ( some 
breeds  with  smaller  skulls  than  coyotes,  may 
have     relatively     well     inflated     braincases, 
broader  at  level  of  parietotemporal  sutures 
than   at   base);   postorbital   constriction  nar- 
rower,   more    elongate,    rising    much    more 
steeply  into  frontal  region;  zygomata  thicker, 
deeper,   usually  more  broadly  flaring;   orbits 
usually  relatively  smaller;  frontals  rising  much 
more  steeply  above  rostrum,  more  depressed 
medially,     more     prominently     convex,     and 
forming  broader,  more  bulging  shield;  orbital 
angle  greater;  temporal  ridges  usually  sharp- 
er; more  often  obscuring  frontal  suture,  more 
often   joining   anterior,   rather  than  posterior 
to  coronal  suture;  sagittal  crest  often  more 
prominent    (may  be  absent  in  some  smaller 
breeds),    usually    sloping    more    posteroven- 
trally;  supraoccipital  shield  often  not  project- 
ing so  far  posteriorly;  external  side  of  occipi- 
tal  more   ossified,   more    often   lacking   thin- 
walled  projection  dorsal  to  foramen  magnum 
(present  in  some  small  individuals);  tympanic 
bullae  relatively  smaller,  usually  more  rugose 
and     much    less    inflated;    palate    relatively 
broader,    central    part    of    posterior    margin 
sometimes   extending   well   behind   toothrow; 
distance  between  toothrow  and  bulla  relative- 
ly   longer;    mandible    relatively    thicker    and 
deeper,  ventral  margin  sometimes  more  con- 


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MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


vex  when  viewed  from  side,  toothrow  more 
bowed  outward  in  center;  teeth  usually  rela- 
tively smaller  with  less  trenchant  cusps,  more 
widely  spaced  in  jaws  (some  specimens  have 
crowded  teeth);  upper  canines  relatively 
thicker  anteroposteriorly  and  much  shorter 
dorsoventrally;  P4  more  often  lacking  promi- 
nent deuterocone  and  lingual  cingulum;  Ml 
having  relatively  larger  paracone  and  meta- 
cone,  relatively  smaller  medial  section,  more 
often  lacking  buccal  cingulum;  M2  relatively 
much  smaller;  p2  more  often  having  posterior 
cusp;  p4  having  posterior  cusps  more  reduced, 
more  often  lacking  well  developed  third  cusp 
and  posteromedial  cingulum;  ml  relatively 
broader,  metaconid  less  prominent  and  not 
projecting  so  far  medially;  m2  and  talonid  of 
ml  usually  relatively  smaller  with  less  trench- 
ant cusps;  dental  and  other  anomalies  more 
common.  Atkins  and  Dillon  (1971)  listed 
differences  between  C.  familiaris  and  C.  la- 
trans  in  the  morphology  of  the  cerebellum. 

Comparison  wit!}  C.  lupus. — Usually  much 
smaller;  skull  usually  much  smaller  ( some 
breeds,  as  Irish  wolfhounds  and  great  Danes, 
may  have  skulls  larger  than  those  of  most 
wolves ) ;  rostrum  usually  relatively  shorter; 
braincase  usually  broader  based,  set  relative- 
ly lower;  postorbital  constriction  usually  rela- 
tively broader  and  rising  more  steeply  into 
frontal  region;  zygomata  usually  shallower, 
not  so  broadly  flaring;  frontals  usually  rising 
more  steeply  above  rostrum,  more  depressed 
medially,  more  prominently  convex,  and  form- 
ing relatively  higher,  broader,  and  more  bulg- 
ing shield;  orbital  angle  usually  greater;  tem- 
poral ridges  more  often  joining  posterior, 
rather  than  anterior  to  coronal  suture;  sagit- 
tal crest  usually  less  prominent  (may  be 
higher  in  some  large  breeds);  supraoccipital 
shield  much  smaller,  not  projecting  so  far 
posteriorly;  tympanic  bullae  usually  smaller, 
less  inflated;  central  part  of  posterior  margin 
of  palate  more  often  extending  behind  tooth- 
row  (especially  in  wolf-sized  individuals); 
distance  between  bulla  and  toothrow  usually 
relatively  larger;  mandible  usually  relatively 


thicker,  ventral  edge  more  often  convex  when 
viewed  from  side  (especially  in  wolf -sized  in- 
dividuals), toothrow  usually  more  bowed  out- 
ward in  center;  teeth,  including  incisors,  of 
larger  individuals  relatively  much  smaller  and 
more  widely  spaced  in  jaws;  dental  and  other 
anomalies  more  common.  Atkins  and  Dillon 
( 1971 )  listed  differences  between  C.  famili- 
aris  and  C.  lupus  in  the  morphology  of  the 
cerebellum;  Hildebrand  (1952b),  Iljin  (1941: 
377-379),  and  Young  (1944:179)  discussed 
distinguishing  features  of  external  appear- 
ance. 

Remarks.- — An  account  of  C.  familiaris  is 
herein  included  primarily  for  comparative 
purposes.  The  origin  of  the  domestic  dog, 
and  its  fossil  and  archeological  history  in 
North  America,  are  not  within  the  scope  of 
this  paper.  Most  authorities  now  think  that 
the  domestic  dog  was  derived  from  one  of  the 
small  Eurasian  subspecies  of  C.  lupus  (Deg- 
erb0l,  1961;  Lawrence,  1966;  Olsen  and  Ol- 
sen,  1977;  Reed,  1961;  Scott,  1968;  Trouessart, 
1911).  Skaggs  (1946:345)  suggested  that  In- 
dian dogs  found  at  a  site  in  Kentucky  may 
have  had  a  coyotelike  ancestor,  but  Allen 
(1920:440)  thought  that  all  American  aborig- 
inal dogs  had  been  introduced  from  the  Old 
World.  The  oldest  known  remains  of  C.  fa- 
miliaris in  North  America,  dated  at  about 
10,400-11,500  B.P.,  were  obtained  from  Jaguar 
Cave  in  Lemhi  County,  Idaho  (Lawrence, 
1966,  196S). 

Attempts  to  describe  the  cranial  charac- 
ters that  distinguish  C.  familiaris  and  C.  lu- 
pus have  been  common  in  the  literature  of 
several  nations  for  many  years.  One  of  the 
earliest  and  most  detailed  of  these  efforts  was 
that  of  Series  (1835),  who  listed  the  follow- 
ing characters  of  the  dog,  as  compared  to 
those  of  the  wolf:  braincase  more  broadly 
based;  frontals  rising  more  steeply,  with 
broader  postorbital  processes;  sagittal  crest 
less  prominent;  supraoccipital  shield  smaller; 
canine  teeth  shorter  dorsoventrally;  carnas- 
sials  shorter  anteroposteriorly;  mandible 
thicker  with  more  convex  ventral  surface;  m3 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


105 


TABLE  4 

Orbital  angle  (in  degrees)  of  C.  latrans,  C.  lupus, 
C.  familiaris,  and  C.  dims. 


lower 
extreme 


upper 
extreme 


sample 
size 


C.  latrans,  western  U.S.  .._ 

C.  lupus,  western  U.S 

C.   familiaris   — _ 

C.  dirus,  Rancho  La  Brea 


42.8 

36 

50 

53 

42.8 

38 

49 

76 

52.9 

40 

64 

58 

53.1 

48 

60 

75 

occasionally  lacking.  A  few  other  features 
mentioned  by  Serres  do  not  now  seem  as  re- 
liable as  those  listed  here. 

Reynolds  (1909:22-23)  compiled  a  list  of 
cranial  characters,  mostly  those  cited  earlier 
by  Serres,  that  had  been  used  by  various 
authors  to  separate  the  dog  and  wolf.  He 
reported  the  most  important  character  to  be 
that  the  plane  of  the  eye  socket  is  more 
obliquely  inclined  to  the  brow  (the  orbito- 
frontal  or  orbital  angle  is  less)  in  the  wolf. 
The  method  of  measuring  this  angle  was 
illustrated  by  Iljin  (1941:387)  and  Mech 
(1970:27).  I  took  the  measurement  on  many 
specimens  and  found  it  to  be  reliable  in  sepa- 
rating skulls  of  dogs  from  those  of  both 
wolves  and  coyotes  (see  table  4). 

Other  characters  suggested  as  useful  in 
distinguishing  skulls  of  dogs,  as  compared  to 
those  of  wolves,  are:  smaller  average  size, 
relatively  smaller  teeth,  less  inflated  bullae 
(Miller,  1912c:313);  projection  of  palate  be- 
hind M2  (Allen,  1920:436);  less  acute  angle 
of  zygomatic  process  of  maxillary,  smaller 
bullae  (Iljin,  1941:390);  braincase  more  ossi- 
fied and  joining  rostrum  at  greater  angle 
(lower  set  relative  to  other  parts  of  skull), 
interorbital  region  more  elongated  (Lawrence 
and  Bossert,  1967:225);  mandible  relatively 
thicker  lateromedially   (Lawrence,   1968). 

The  cranial  differences  between  C.  latrans 
and  C.  familiaris  received  less  attention  until 
relatively  recently.  Coues  (1873)  and  Pack- 
ard (1885)  were  both  struck  by  what  they 
considered  to  be  close  resemblance  in  external 
appearance  of  the  coyote  and  the  American 


Indian  dog.  But  Allen  (1920:434-435,  450) 
pointed  out  that  no  one  had  yet  made  a  care- 
ful comparison  of  dog  and  coyote  skulls.  He 
reported  that  dogs  had  a  smaller  heel  of  ml, 
less  trenchant  cusps  on  the  molars  and  pre- 
molars, and  a  more  prominent  outward  bend 
of  the  lower  toothrow  at  the  junction  of  the 
molar  and  premolar  series. 

Other  characters  proposed  as  useful  in 
distinguishing  skulls  of  domestic  dogs,  as 
compared  to  those  of  coyotes,  are:  more  in- 
flated frontal  sinuses,  greater  orbital  angle, 
more  obtuse  angle  in  maxillo-jugal  suture, 
smaller  tympanic  bullae,  more  widely  spaced 
teeth,  smaller  medial  section  of  Ml  (Hall, 
1943 ) ;  broader  rostrum  and  more  widely 
spaced  incisors  (Burt,  1946:61-62);  smaller 
ratio  obtained  by  dividing  the  distance  be- 
tween the  inner  margins  of  the  alveoli  of  PI 
by  the  distance  from  the  anterior  margin  of 
the  alveolus  of  PI  to  the  posterior  margin  of 
the  alveolus  of  M2  (Howard,  1949);  canine 
teeth  shorter  dorsoventrally  and  thicker  an- 
teroposteriorly  (Jackson,  1951:242);  relatively 
broader  base  of  braincase  and  deeper  mandi- 
ble ( Bee  and  Hall,  1951 ) ;  deeper  and  thicker 
mandible   (Lawrence,  1968). 

A  problem  associated  with  attempts  to  dis- 
tinguish between  C.  lupus  and  C.  familiaris 
is  the  role  of  domestication  in  modifying  the 
phenotype  of  the  skull.  The  most  critical 
question  is  whether  nutritional  or  other  fac- 
tors involved  in  captivity  cause  the  develop- 
ing skull  of  a  wolf  to  take  on  characters  nor- 
mally found  in  the  dog.  Studies  have  demon- 
strated that  wolves  raised  in  captivity  some- 


106 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


times  show  such  changes  as  a  shortening  of 
the  jaws,  overlapping  of  the  teeth,  more 
steeply  raised  forehead,  and  general  decrease 
in  size.  This  subject  was  discussed  in  detail 
by  Degerbol  (1961),  Iljin  (1941:390-392), 
Lawrence  (1966).  Scott  (1968:246-247),  and 
various  other  authors  cited  therein.  Thus  far 
all  of  the  investigations  have  concerned  what 
happens  to  wolves  in  captivity,  but  it  has  not 
yet  been  determined  whether  C.  familiaris 
could  take  on  wolflike  characters  under  feral 
conditions. 

Interbreeding  of  domestic  dogs  with 
wolves  or  coyotes  has  occasionally  occurred, 
and  was  discussed  in  the  first  main  part  of 
this  paper.  Hybridization  between  C.  lupus 
and  C.  familiaris  has  long  been  accepted  (see 
reviews  by  Allen,  1920:433-434;  Iljin,  1941: 
360-361;  and  Young,  1944:180-210).  Some 
early  naturalists,  such  as  Coues  (1873) 
thought  hybridization  between  dog  and  coy- 
ote to  be  common  in  the  west,  but  Allen 
( 1920 )  refuted  this  view  on  the  basis  of 
cranial  evidence.  Later,  however,  specimens 
began  appearing  in  the  eastern  states,  that 
were  considered  to  represent  hybrids  between 
the  two  species  (coy-dogs).  Mengel  (1971) 
has  reviewed  this  subject  in  detail. 

Canis  dirus  Leidy 

1854.  Canis  primaevus  Leidy,  Proc.  Acad. 
Nat.  Sci.,  Philadelphia,  7:200.  Not  C. 
primaevus  of  Hodgson,  1833. 

1855.  Canis  clirus  Leidy,  Proc.  Acad.  Nat. 
Sci.,  Philadelphia,  1858,  p.  21.  Type 
from  banks  of  Ohio  River  below  Evans- 
ville,  Vanderburgh  County,  Indiana. 

1869.  Canis  indianensis  Leidy,  Jour.  Acad. 
Nat.  Sci.,  Philadelphia,  7:368.  An  in- 
advertant  renaming  of  C.  dirus. 

1876.  Canis  mississippiensis  J.  A.  Allen, 
Amer.  Jour.  Sci.,  ser.  3,  11:49.  Type 
from  Lead  Region  of  Upper  Missis- 
sippi. 

1884.  Canis  lupus,  Cope  and  Wortman,  Ann. 
Rept.  State  Geol.  Indiana,  14:9. 


1912.  Canis  dirus,  Merriam,  Mem.  Univ.  Cal- 
ifornia, 1:218. 

1916.  Canis  ayersi  Sellards,  Ann.  Rept.  Flor- 
ida Geol.  Surv.,  8:152.  Type  from 
Vero,  Indian  River  County,  Florida. 

1918.  Aenocyon  dims,  Merriam,  Univ.  Cali- 
fornia Publ.  Bull.  Dept.  Geol.,  10:533. 

1918.  Aenocyon  ayersi,  Merriam,  Univ.  Cali- 
fornia Publ.  Bull.  Dept.  Geol.,  10:533. 

1929.  Canis  ( Aenocyon )  ayersi,  Simpson, 
Bull.  Amer.  Mus.  Nat.  Hist.,  56:572. 

1946.  Canis  (Aenocyon)  dirus,  Stock,  Lance, 
and  Nigra,  Bull.  S.  California  Acad. 
Sci.,  45:109. 

1962.  Canis  ayersi,  Weigel,  Florida  Geol. 
Surv.  Spec.  Publ.,  no.  10,  p.  37. 

1972.  Canis  dirus,  Knrten  and  Anderson, 
Tebiwa,  15:37. 

1974.  Canis  dirus,  R.  A.  Martin,  in  Pleisto- 
cene mammals  of  Florida  (edit. 
Webb),  p.  73. 

Type. — Left  maxillary  fragment  with  P2- 
M2;  no.  11614,  Acad.  Nat.  Sci.,  Philadelphia; 
banks  of  Ohio  River  below  Evansville,  Van- 
derburgh County,  Indiana. 

Geological  distrilmtion. — Rancholabrean 
to  early  Recent. 

Geographical  distribution. — Known  from 
Alberta,  Arizona,  Arkansas.  California,  Flor- 
ida, Idaho,  Illinois,  Indiana,  Kansas,  Ken- 
tucky, Louisiana,  Missouri,  Nebraska,  Ne- 
vada, New  Mexico,  Oklahoma,  Oregon,  Penn- 
sylvania, Tennessee,  Texas,  Utah,  Virginia, 
West  Virginia,  Wisconsin,  Aguascalientes,  Ja- 
lisco (possibly),  Estado  de  Mexico,  Nuevo 
Leon,  Puebla,  and  northern  Peru. 

Description. — Size  large  for  the  genus; 
skull  averaging  largest  in  genus,  with  mostly 
broad  proportions;  rostrum  elongated,  rela- 
tively broad  and  deep;  braincase  relatively 
small,  not  much  inflated  dorsoposteriorly; 
postorbital  constriction  elongated,  narrow 
lateromedially,  rising  steeply  into  frontal  re- 
gion; zygomata  thick,  deep,  broadly  flaring; 
orbits  relatively  small;  frontals  usually  well 
elevated   above  rostrum,   moderately  convex. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


107 


forming  broad  shield;  temporal  ridges  usu- 
ally not  sharp,  seldom  obscuring  frontal  su- 
ture, usually  joining  at  or  anterior  to  coronal 
suture;  sagittal  crest  prominent,  sharp  dor- 
sally;  supraoccipital  shield  narrow,  pro- 
jecting far  posteriorly;  external  side  of  occipi- 
tal well  ossified;  tympanic  bullae  usually 
moderate  in  size,  not  well  inflated;  vertical 
plates  of  palatines  flaring  broadly  anteriorly: 
posterior  end  of  vomer  usually  extending  only 
slightly  behind  posterior  nasal  opening;  post- 
palatine  foramina  usually  opposite  posterior 
ends  of  P4;  optic  foramen  and  anterior  lacer- 
ated foramen  normally  close  together  in  com- 
mon pit;  mandible  thick  and  deep,  ventral 
margin  not  convex  when  viewed  from  side, 
toothrow  bowed  outward  in  center;  incisors 
relatively  large;  upper  canines  prominent, 
thick  anteroposteriorly;  premolars  relatively 
broad;  P4  relatively  large,  usually  lacking 
prominent  deuterocone  and  lingual  cingulum; 
Ml  having  relatively  large  paracone  and  met- 
acone,  relatively  small  medial  section  with- 
out trenchant  cusps,  reduced  hypocone  with 
ridge  seldom  extending  anteriorly  around 
base  of  protocone,  and  sometimes  with  pro- 
nounced buccal  cingulum;  M2  relatively 
small;  p2  usually  lacking  posterior  cusp;  p3 
often  with  second  and  third  cusps;  p4  usually 
having  second  and  third  cusps,  and  pro- 
nounced posteromedial  cingulum  extending 
behind  third  cusp;  ml  relatively  large,  usu- 
ally having  relatively  small,  low-set  talonid. 
For  additional  details,  and  description  of 
postcranial  skeleton,  see  Merriam  (1912); 
Stock,  Lance,  and  Nigra  (1946);  Nigra  and 
Lance  (1947);  Stock  and  Lance  (1948);  and 
Galbreath   (1964). 

Comparison  with  C.  lupus. — Skull  usually 
larger;  posterior  part  of  rostrum  usually  not 
so  deep;  postorbital  constriction  rising  more 
steeply  into  frontal  region;  frontals  less  con- 
vex, less  depressed  medially,  forming  rela- 
tively broader  and  flatter  shield;  orbital  angle 
greater;  temporal  ridges  smoother,  less  often 
obscuring  frontal  suture;  sagittal  crest  usu- 
ally   more    prominent;    supraoccipital    shield 


narrower,  projecting  farther  posteriorly,  more 
often  having  posteroventral  hook;  vertical 
plates  of  palatines  flaring  more  broadly  an- 
teriorly; posterior  end  of  vomer  not  extending 
so  far  behind  posterior  nasal  opening;  post- 
palatine  foramina  usually  set  more  posterior- 
ly; optic  foramen  and  anterior  lacerated 
foramen  closer  together;  mandible  usually 
relatively  thicker  and  deeper;  upper  canines 
usually  relatively  smaller;  Ml  having  more 
reduced  hypocone,  its  anterior  ridge  much 
less  often  extending  anteriorly  around  base  of 
protocone,  and  more  often  with  buccal  cingu- 
lum; p2  more  often  lacking  posterior  cusp; 
p4  with  posterior  margin  of  second  cusp  usu- 
ally sloping  anteroventrally  rather  than  pos- 
teroventrally,  more  often  with  third  cusp  and 
posteromedial  cingulum  extending  behind 
third  cusp;  ml  usually  relatively  larger,  usu- 
ally with  relatively  smaller,  narrower,  lower 
set  talonid.  For  additional  details,  and  com- 
parison of  postcranial  elements  of  C.  dims 
and  C.  lupus,  see  Merriam  (1912);  Stock, 
Lance,  and  Nigra  ( 1946 ) ;  and  Stock  and 
Lance  (1948). 

Other  comparison. — See  account  of  C. 
armbrusteri. 

Remarks. — The  nomenclatural  confusion 
regarding  the  type  specimen,  and  other  early 
reported  material  of  dire  wolves,  was  resolved 
by  Merriam  (1912:218-221).  He  correctly 
determined  that  Cams  dims  was  the  proper 
name  for  all  specimens  that  had  been  re- 
ported up  to  the  time  of  his  study. 

In  his  original  description  of  Canis  primae- 
vus  (=Canis  dims),  Leidy  (1854:200)  noted: 
"Certain  naturalists  may  regard  the  fossil  as 
an  indication  of  a  variety  only  of  the  Canis 
lupus,  and  of  the  correctness  of  such  a  view 
I  shall  not  attempt  to  decide."  Subsequently, 
Cope  and  Wortman  (1884:10)  wrote  that  "it 
is  impossible  to  admit  this  fossil  to  the  rank 
of  a  distinct  and  well  defined  species,  but  it 
appears,  in  our  judgement,  to  be  but  a  variety 
which  has  a  living  representative  in  the 
mountains  of  Oregon,  today."  Nonetheless, 
Leidv's  original  recognition  of  the  distinctness 


108 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


of  the  dire  wolf  was  borne  out  when  a  large 
number  of  specimens  became  available. 

Rancho  La  Brea  has  yielded  by  far  the 
greatest  amount  of  dire  wolf  material.  In 
fact,  there  are  few  complete  skulls  from  all 
other  localities  combined.   Thus,  for  purposes 


10 

9- 

8- 

7  - 

6 

5- 

4  - 

3 

2 

I 

0 
-I 
-2 
-3 


D 


D 


D 


D 


D 


D 


D 


D 


D 


N 


Y 
I 


B 


0 

I     sxinp       . 
kayrra°tNkl 

S    KHT  dQt  n 

A  R     FHBST  H    I 

N    F    LO   ,  P    F  LA 
SB,AlYMs 


0 


A 

T 


0 


My 

F 


N 


F, 


M 


-1 1 r- 

-4     -3    -2 


■I 


— r 
0 


Fig.  52. — Graphical  results  of  multivariate  analy- 
sis comparing  a  group  of  62  skulls  of  C.  dims  from 
Rancho  La  Brea  (D),  with  a  total  of  467  Recent 
C.  lupus  divided  into  the  following  subspecific 
groups:  C.  /.  arctos  (A),  C.  /.  baileyi  (B),  C.  /. 
fuscus  (F),  C.  /.  hudsonicus  (H),  C.  /.  irremotus 
(I),  C.  /.  mackenzii  (K),  C.  I.  ligoni  (L),  C.  /.  mo- 
goUonensis  (M),  C.  /.  nubilus  (N),  C.  /.  occidentalis 
(O),  C.  /.  pambasileus  (P),  C.  /.  lycaon  (R),  C.  I. 
monstrabilis  (S),  C.  /.  tundrarum  (T),  C.  /.  Ijcrnardi 
(X),  C.  /.  youngi  (Y).  As  in  previous  multivariate 
analyses  in  this  paper,  15  measurements  of  the  skull 
were  used,  but  in  this  case  sexes  were  combined  in 
each  group.  Only  marginal  positions  of  individuals 
of  each  group  are  plotted. 


of  statistical  comparison,  I  have  restricted  my 
sample  of  C.  dims  to  62  specimens  from 
Rancho  La  Brea  that  were  complete  enough 
for  use  in  multivariate  analysis.  These  were 
all  unknown  as  to  sex,  and  so  were  tested 
against  the  combined  male  and  female  sam- 
ples of  each  subspecies  of  C.  lupus  for  which 
more  than  five  specimens  were  available  (a 
total  of  467  specimens).  Figure  52  shows  the 
resulting  ranges  of  variation  of  each  group, 
and  the  complete  separation  of  the  dire  wolf 
and  the  gray  wolf.  Measurements  of  the 
series  of  C.  dims  are  listed  in  appendix  B 
(part  15),  and  their  means  are  compared 
with  those  of  C.  lupus  in  figure  53.  The  skull 
of  the  dire  wolf  is  seen  to  be  much  larger 
than  that  of  C.  lupus,  and  to  be  proportion- 
ally broader  at  the  canines  and  frontal 
shield,  but  not  so  deep,  relatively,  between 
the  toothrow  and  orbit.  The  P4  is  propor- 
tionally much  longer,  but  the  upper  canine 
and  M2  have  relatively  smaller  diameters. 

The  lack  of  multivariate  overlap  between 
C.  dims  and  C.  lupus,  and  the  striking  differ- 
ences in  size  and  proportion,  lead  me  to  no 
other  conclusion  than  that  the  two  must  be 
treated  as  distinct  species.  There  also  is  no 
statistical  evidence  to  suggest  that  the  dire 
wolf  is  ancestral  to  the  living  gray  wolf. 

Although  the  material  from  Rancho  La 
Brea  is  by  far  the  most  abundant  from  any 
one  locality,  well  preserved  upper  cranial  ele- 
ments from  other  sites  in  California,  Mexico, 
Texas,  Missouri,  and  Kentucky  has  confirmed 
that  the  same  species,  with  the  same  well 
marked  characters,  was  broadly  distributed 
in  the  late  Pleistocene.  Among  the  most  re- 
liable cranial  characters  that  distinguish  C. 
dims  from  other  wild  species  of  Canis  are: 
large  over-all  size,  relatively  broad  frontal 
shield,  large  orbital  angle  (see  table  4),  nar- 
row supraoccipital  shield  projecting  far  pos- 
teriorly, vertical  plates  of  palatines  flaring 
broadly  anteriorly,  postpalatine  foramina  set 
relatively  far  posteriorly,  optic  foramen  and 
anterior  lacerated  foramen  close  together  in 
common   pit,   large   carnassial   teeth,   and  re- 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


109 


0 


.04        .08 


LUPUS      DIRUS 


1.00      1.10       1.20    1.30 

Fig.  53. — Ratio  diagram  comparing  means  of  C. 
dirus  (n=62)  and  the  total  sample  of  Recent  C. 
lupus  (combined  sexes,  n=482).  Vertically  arranged 
numbers  represent  the  measurements  so  numbered 
in  appendix  B.  A  log  difference  scale  is  provided 
above,  and  a  ratio  scale  below  the  diagram. 

duced  hypocone   on    Ml.    I   examined   more 
than  500  skulls  from  Rancho  La  Brea  in  the 


Los  Angeles  County  Museum,  and  found  few 
that  lacked  more  than  one  of  these  characters. 
There  was  no  difficulty  in  assigning  most  of 
this  material  to  C.  dims,  and  picking  out  the 
eight  skulls  in  the  LACM  collection  that  are 
referable  to  C.  lupus  (see  pp.  99-100).  The 
specimens  from  Rancho  La  Brea  represent  a 
population  that  was  geographically  local,  but 
the  deposits  were  laid  down  over  many  thou- 
sands of  years,  and  thus  considerable  intra- 
specific  variation  would  be  expected  at  the 
site. 

Sellards  (1916:152-157)  described  Canis 
ayersi  from  Vero,  Florida  as  belonging  to  the 
same  group  as  C.  dims,  but  being  specifically 
distinct.  The  single  skull  on  which  he  based 
his  description  was  thought  to  have  a  rela- 
tively narrower  rostrum  than  the  skulls  found 
at  Rancho  La  Brea.  But  the  rostrum  of 
ayersi  had  apparently  been  damaged  prior  to 
its  recovery,  and  since  Sellards'  study  some 
restorative  work  has  been  carried  out.  A  full 
evaluation  is  now  impossible,  but  the  skull  is 
clearly  narrower  than  most  specimens  from 
Rancho  La  Brea.  In  addition,  some  of  the 
mandibles  from  Florida  that  I  examined  are 
longer  than  any  from  the  southwest.  Dire 
wolves  in  Florida  may  therefore  have  tended 
to  have  relatively  long  and  narrow  jaws,  but 
the  differences  are  slight  and  I  do  not  con- 
sider them  to  be  of  specific  value.  In  all 
other  characters,  as  acknowledged  by  Sel- 
lards, the  Florida  material  resembles  C.  dims 
from  Rancho  La  Brea. 

In  describing  C.  ayersi,  Sellards  (1916: 
156)  noted  that  there  was  no  effective  barrier 
between  Florida  and  Evansville,  Indiana,  the 
type  locality  of  C.  dims,  and  that  the  type  of 
C.  dirus,  a  maxillary  fragment,  might  even- 
tually be  shown  to  represent  the  same  species 
as  found  in  Florida.  Olson  (1940:44)  sus- 
spected  that  the  Indiana  and  Florida  speci- 
mens were  probably  referable  to  the  same 
species,  C.  dirus  on  the  basis  of  nomencla- 
tural  priority,  and  that  a  new  name  might  be 
needed  for  material  from  California.  Re- 
cently, a  very  broad  skull  of  a  dire  wolf  was 


110 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


obtained  at  Welsh  Cave,  Woodford  County, 
Kentucky,  only  150  miles  from  Evansville, 
and  was  assigned  to  C.  dims  by  Guilday, 
Hamilton,  and  McCrady  (1971:274).  In  ros- 
tral proportions  this  specimen  resembles 
skulls  from  Rancho  La  Brea  more  than  it  does 
the  type  of  C.  ayersi,  and  it  thus  supports 
recognition  of  the  type  of  C.  dims  and  speci- 
mens from  Rancho  La  Brea  as  representing 
the  same  species. 

Martin  (1974:73)  reported  that  a  skull 
from  Reddick,  Marion  County,  Florida  was 
similar  in  proportion  to  specimens  from 
Rancho  La  Brea,  and  he  thus  implied  that 
there  was  morphological  overlap  between  the 
two  populations  represented.  Martin  sug- 
gested that  the  type  of  C.  ayersi,  as  well  as 
all  other  dire  wolf  material  from  Florida,  was 
referable  to  C.  dims,  and  I  have  followed  the 
same  course  in  this  paper.  I  have  not  re- 
tained ayersi  as  a  subspecies,  and  presently 
prefer  to  recognize  C.  dims  as  a  widespread 
monotypic  species. 

In  a  popular  article,  Frick  (1930:79)  listed 
two  new  subspecific  names,  Aenocyon  dims 
alaskensis  from  the  pre-tundra  fauna  of 
Alaska-Yukon,  and  A.  dims  nebrascensis  from 
the  Sheridan  fauna  of  Nebraska.  No  type 
specimens  or  exact  localities  were  desig- 
nated, and  no  descriptions  were  provided. 
Nonetheless,  several  authors  referred  to  sup- 
posed dire  wolves  from  Nebraska  as  nebra- 
scensis Frick  (Schultz  and  Stout,  1948:565; 
Schultz  and  Tanner,  1957:71;  Schultz  and 
Martin,  1970:347).  According  to  Schultz 
(1934:369),  there  were  specimens  of  A.  d. 
nebrascensis  Frick  from  Hay  Springs,  Sheri- 
dan County,  Nebraska  in  the  Frick  collection 
of  the  American  Museum  of  Natural  History. 
No  one,  however,  discussed  particular  speci- 
mens, and  the  original  faunal  list  from  this 
site  had  included  only  the  gray  wolf  (Mat- 
thew, 1918).  The  specimens  of  wolves  from 
Hay  Springs  that  I  examined  are  referable  to 
C.  lupus,  and  are  discussed  above  in  the  ac- 
count of  that  species.  The  only  specimens 
of  wolves  reported  from  specific  localities  in 


Alaska  also  have  been  referred  to  C.  lupus, 
and  it  is  unlikely  that  the  dire  wolf  ever  oc- 
curred so  far  north.  I  therefore  consider 
Frick's  names  as  nornena  nuda  and  have  not 
employed  them  in  this  paper. 

Merriam  (1918)  created  the  genus  Aeno- 
cyon to  include  what  he  thought  were  three 
species:  A.  dims,  A.  ayersi,  and  A.  milleri. 
Goldman  (1944:400)  observed  that  the  cra- 
nial and  dental  details  of  the  dire  wolf  justi- 
fied recognition  of  Aenocyon  as  a  genus  or 
subgenus.  Stock,  Lance,  and  Nigra  (1946: 
109)  formally  used  Aenocyon  as  a  subgenus 
for  C.  ( A. )  dims.  Various  other  authors  have 
either  followed  Merriam,  employed  Aenocyon 
as  a  subgenus,  or  ignored  the  term  (see  "Rec- 
ord of  occurrences,"  below). 

As  pointed  out  above,  milleri  and  ayersi 
are  synonyms  of  C.  lupus  furlongi  and  C. 
dims,  respectively,  and  thus  there  is  only  one 
species  of  dire  wolf,  C.  dims.  Although  this 
species  has  the  most  pronounced  specific 
characters  of  any  North  American  member  of 
the  genus  Canis,  these  characters  do  not 
seem  to  be  of  a  higher  order  than  those  dis- 
tinguishing other  species.  Furthermore,  there 
is  no  evidence  that  the  lineage  of  the  dire 
wolf  was  long  separate  from  that  of  other 
Canis.  Indeed,  C.  dims  seemingly  was  the 
last  wild  species  of  Canis  to  appear  in  North 
America,  and  it  probably  descended  from  the 
same  basic  stock  that  gave  rise  to  other 
wolves.  Thus  I  have  not  employed  Aenocyon 
as  a  genus  or  subgenus,  and  have  synony- 
mized  it  under  Canis. 

Record  of  occurrences. — The  dire  wolf  is 
represented  by  more  fossil  material  than  any 
other  species  of  Canis,  and  yet  its  known 
geological  range  is  relatively  short,  being  re- 
stricted to  the  Rancholabrean  and  early  Re- 
cent. A  number  of  large  canids  from  older 
sites  were  assigned  to  C.  dims  by  previous 
authors,  but  in  such  cases  identification  was 
incorrect  or  material  was  inadequate  to  allow 
careful  evaluation.  Irvingtonian  records  from 
Irvington,  California  (Savage,  1951:230); 
Rushville  fossil  quarry   (Schultz  and  Tanner, 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


111 


Fig.  54. — Map  showing  localities    (black   dots)    of  C.    dints.     Because    of    the    scale    of    the    map,    it    was 
not  possible  to  plot  all  localities  in  crowded  areas. 


1957:71)  and  Angus  fossil  quarry  (Schultz 
and  Martin,  1970:347),  Nebraska;  Port  Ken- 
nedy, Pennsylvania  (Cope,  1899:227);  and 
Rock  Creek,   Texas    (Troxell,   1915:633)    are 


discussed  above  in  the  account  of  C.  arm- 
brusteri.  And  records  from  Hay  Springs 
(Schultz,  1934:369;  Schultz  and  Stout,  1948: 
563;  Schultz  and  Tanner,  1957:71)  and  Mul- 


112 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


len  (Martin,  1972:174),  Nebraska  are  dis- 
cussed above  in  the  account  of  C.  lupus.  In 
addition,  the  dire  wolf  was  associated  with 
the  Blancan  Broadwater  quarry  site,  Ne- 
braska (Barbour  and  Schultz,  1937:4),  and 
Blanco  fauna,  Texas  (Vanderhoof,  1937).  No 
particular  specimens,  however,  were  dis- 
cussed, and  subsequent  papers  on  the  Broad- 
water quarry  (Schultz  and  Stout,  1945:234; 
1948:563;  Hibbard,  1970:414)  and  Blanco 
fauna  (Meade,  1945;  Johnston  and  Savage, 
1955:36-37;  Dalquest,  1975)  did  not  mention 
the  presence  of  C.  dims.  Perhaps  the  initial 
reports  of  C.  dims  had  been  based  mistakenly 
on  remains  of  Borophagus. 

The  following  list  is  arranged  alphabeti- 
cally by  state  and  province,  and  geographi- 
cally (north  to  south,  west  to  east)  within 
states  and  provinces,  except  that  Latin  Amer- 
ican areas  are  placed  last.  Specimens  exam- 
ined by  me  are  identified  by  element,  mu- 
seum number,  or  both;  and  selected  measure- 
ments are  found  in  appendix  B  (part  15)  and 
appendix  C  (part  7).  Occurrences  also  are 
shown  on  the  map  in  figure  54. 

ALBERTA.— Castleguard  icefield,  Banff  National 
Park;  late  Pleistocene;  as  C.  dirus  (Cowan,  1954:44). 
This  record  is  based  on  a  single  lower  canine  tooth 
found   lying  on  the  surface. 

Medicine  Hat;  Sangamon;  as  C.  dirus  (Churcher, 
1970:63). 

ARIZONA. — Ventana  Cave,  Papago  Indian  Res- 
ervation, Pima  County;  Wisconsin  (Hibbard,  1958); 
as  C.  dims   (Colbert,   1950:132). 

Murray  Springs,  1  mi.  W  Lewis  Spring  on  San 
Pedro  River,  Cochise  County;  late  Pleistocene;  two 
mandibular  fragments,  UAriz  4394,  4395. 

Whitewater  Draw,  near  Douglas,  Cochise  Coun- 
ty; early  Recent;  as  C.  dirus   (Hester,   1960:69). 

ARKANSAS. — Peccary  Cave,  eastern  Newton 
County;  early  Recent;  as  C.  dirus  (Davis,  1969:164; 
Quinn,  1972:92). 

CALIFORNIA.— Samwel  Cave,  Shasta  Lake, 
Shasta  County;  Wisconsin  (Hibbard,  1958);  as  C. 
dirus  (Kurten  and  Anderson,  1972:37);  mandibular 
fragment,  UCMP  9566. 

Potter  Creek  Cave,  1  mi.  SE  Baird,  Shasta  Coun- 
ty; Wisconsin  (Hibbard,  1958);  as  C.  indianensis 
(Sinclair,  1904:17),  as  C.  dirus  (Anderson,  1968: 
22). 

Hawver  Cave,  5  mi.  E  Auburn,  El  Dorado  Coun- 
ty; Wisconsin  (Hibbard,  1958);  as  C.  near  dims 
(Stock,  1918:478);  as  C.  dirus  (Anderson,  1968:22). 


Cool  quarry.  El  Dorado  County;  late  Pleistocene; 
mandibular  fragment,  UCMP  38328. 

Teichart  gravel  pit,  Sacramento  County;  late 
Pleistocene;   mandibular  fragment,  UCMP  85380. 

Arroyo  Las  Positas,  Alameda  County;  Pleistocene; 
mandible,   FM   PM664. 

Livermore  Valley,  near  San  Leandro,  Alameda 
County;  Pleistocene;  as  C.  indianensis  ( Leidy,  1873: 
230),  as  C.  dirus  (Merriam,  1912:244). 

Oil  Springs,  Tulare  County;  Pleistocene;  as  C. 
indianensis  (Merriam,  1903:288),  as  C.  dirus  (Mer- 
riam, 1912:244). 

McKittrick  tar  seeps,  Kern  County;  Wisconsin;  as 
A.  dims  (Merriam  and  Stock,  1921;  Schultz,  1938b: 
169);  two  complete  skulls,  LACM;  six  maxillary 
fragments,  LACM;  13  mandibles,  LACM;  three  man- 
dibles, UCMP. 

Maricopa  Brea,  near  Maricopa,  Kern  County; 
Wisconsin  (C-14  date:  13,860  B.P.);  as  C.  dirus 
(Shakespear,  1975);  three  complete  skulls  with  man- 
dibles, six  other  mostly  complete  skulls;  10  cranial 
and  maxillary  fragments;  20  mostly  complete  mandi- 
bles; many  other  fragments  and  isolated  teeth,  all 
in  LACM. 

Carpinteria  asphalt,  Santa  Barbara  County;  Wis- 
consin (Hibbard,  1958);  as  A.  near  dirus  (Wilson, 
1933:69). 

Rancho  La  Brea,  Los  Angeles,  Los  Angeles  Coun- 
ty; Wisconsin;  as  "C.  indianensis  (?)"  (Merriam, 
1906),  as  C.  dirus  (Merriam,  1912:218;  Marcus, 
1960:2),  as  A.  dims  (Merriam,  1918:533;  Stock, 
1929:286),  as  C.  (A.)  dirus  (Stock,  Lance,  and 
Nigra,  1946:109;  Nigra  and  Lance,  1947:26;  Stock 
and  Lance,  1948:79);  520  complete  or  mostly  com- 
plete skulls  without  associated  mandibles,  LACM; 
one  skull,  AMNH;  two  skulls,  KU;  five  skulls,  USNM; 
ten  skulls,  UCMP;  90  complete  mandibles,  LACM; 
three  mandibles,  AMNH;  three  mandibles,  USNM; 
five  mandibles,  UCMP;  numerous  fragments  and  iso- 
lated teeth,  LACM.  The  dire  wolf  material  from 
Rancho  La  Brea  is  the  most  abundant  of  any  large 
mammal  from  any  fossil  site  in  North  America.  The 
number  of  individual  wolves  represented  in  the  col- 
lection of  the  Los  Angeles  County  Museum  was 
estimated  at  2,000  by  Stock,  and  was  counted  at 
1,646  by  Marcus  (1960).  The  amount  of  well  pre- 
served material  offers  an  unparalleled  opportunity  for 
studies  of  variation  in  a  local  population  of  canids, 
but  as  yet  little  has  been  done  in  this  regard.  Nigra 
and  Lance  (1947)  found  the  average  size  of  meta- 
podials  of  C.  dirus  to  differ  between  the  five  major 
tar  pits  from  which  remains  were  recovered.  Popula- 
tion studies  of  this  kind  are  qualified  by  the  prob- 
ability that  the  different  pits  were  active  at  different 
times,  and  that  each  was  active  over  a  lengthy 
period  in  which  chronological  mixing  of  elements 
occurred.  Radiocarbon  dates  based  directly  on  speci- 
mens of  C.  dirus  from  Rancho  La  Brea  were  given 
as  9,860±550  and  10,710±320  B.P.  by  Miller  ( 1968: 
14).  It  is  likely,  however,  that  remains  of  wolves 
were  deposited  at  the  site  over  thousands  of  years.  I 
found  the  skulls  of  C.   dirus  to  be  remarkably  con- 


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NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


113 


sistent  in  the  critical  characters  that  distinguish  the 
species,  and  on  the  whole  I  agree  with  the  detailed 
description  provided  by  Merriam  in  1912.  Studies 
of  postcranial  elements  from  Rancho  La  Brea  have 
indicated  that  the  dire  wolf  had  a  relatively  stockier 
body,  lighter  limbs,  and  shorter  feet  than  modern 
C.  lupus  (Merriam,  1912:236;  Stock,  Lance,  and 
Nigra,   1946;   Stock  and  Lance,   1948:79). 

Harold  Beds,  5  mi.  SE  Palmdale,  Los  Angeles 
County;  Pleistocene;  three  metapodials,  USNM  13085. 

Harbor  freeway,  Los  Angeles,  Los  Angeles  Coun- 
ty;  Wisconsin;   as  C.   cf.   dirus   (Miller,   1971:54). 

San  Pedro,  Los  Angeles  County;  Wisconsin;  as 
C.  cf.  dirus  (Miller,  1971:45). 

La  Mirada,  Los  Angeles  County;  Wisconsin;  as 
C.  cf.  dirus  (Miller,  1971:49). 

Newport  Bay  Mesa,  Orange  County;  Wisconsin; 
as  C.  cf.  dirus  (Miller,  1971:34). 

Costeau  pit,  2  mi.  S  El  Toro,  Orange  County; 
Wisconsin;  as  C.  cf.  dirus   (Miller,  1971:17). 

FLORIDA.— Aucilla  River  IA,  Jefferson  County; 
Wisconsin;   as   C.   dirus   (Webb,    1974b:  17). 

Ichetucknee  River,  Columbia  County;  Wisconsin; 
as  C.  dirus  (Webb,  1974b:  17);  maxillary  fragment, 
UF  8006;  three  mandibular  fragments,  UF  8005, 
12899,  17717;  three  canine  teeth,  UF  1995,  8214, 
8215;  postcranial  fragments,  MCZ  18347-18349.  The 
mandibles  resemble  those  of  dire  wolves  from  Rancho 
La  Brea,  but  are  larger. 

Santa  Fe  River  IIA,  Gilchrist  County;  Rancho- 
labrean  (Webb,  1974b:31;  apparently  incorrectly 
designated  as  late  Irvingtonian  on  p.  13);  as  C.  dirus 
(Webb,  1974b:  17). 

Hornsby  Springs,  near  High  Springs,  Alachua 
County;  Wisconsin;  as  A.  mjersi  (Bader,  1957:71); 
maxillarv  fragment,  UF  3988;  mandibular  fragment, 
UF  3987. 

Haile  VIIIA,  Alachua  County;  Sangamon;  as  C. 
dirus   (Webb,   1974b:  17). 

Arredondo  IB,  4  mi.  SW  Gainesville,  Alachua 
County;  Sangamon;  as  A.  ayersi  (Bader,  1957:54), 
as  C.  dirus  (Webb,  1974b:  17). 

Devil's  Den,  near  Williston,  Levy  County;  late 
Wisconsin  or  early  Recent  (7,000-8,000  B.P.);  as 
C.  dims  (Martin  and  Webb,  1974:126);  incomplete 
sk-ull,  UF  7996. 

Wekiva  River,  Levy  County;  late  Pleistocene; 
mandibular  fragment,  UF  14204. 

Reddick  IA,  Marion  County;  Sangamon  (Webb, 
1974b:13);  as  C.  ayersi  (Gut,  1939),  as  C.  (A.) 
ayersi  (Gut  and  Ray,  1964),  as  C.  dirus  (Martin, 
1974:73);  crushed  skull  with  mandibles,  UF  2923; 
two  crushed  skulls  without  mandibles,  UF  3081,  one 
unnumbered;  mandibular  fragment,  UF;  two  isolated 
Ml,  isolated  M2,  P4,  ml,  UF;  P4,  MCZ. 

Eichelberger  Cave,  2  mi.  SW  Belleview,  Marion 
County;  late  Pleistocene;  cast  of  pair  of  mandibles, 
MCZ  7349;  two  mandibular  fragments  ( probably 
from  same  individual),  UF  1622,  1623;  isolated 
teeth,  UF.  The  mandibles  and  lower  carnassials  are 
the   largest  that  I   examined. 

Sabertooth    Cave,    1    mi.    NW    Lecanto,    Citrus 


County;  Wisconsin;  as  C.  ayersi  (Simpson,  1928:9), 
as  C.  dirus  (Webb,  1974b:17). 

Rock  Springs,  Orange  County;  Sangamon;  as  C. 
dims    (Webb,    1974b:  17). 

Seminole  Field,  near  St.  Petersburg,  Pinellas 
County;  Wisconsin  (Hibbard,  et  al,  1965);  as  C. 
(A.)  ayersi  (Simpson,  1929a:572),  as  C.  dirus 
(Webb,  1974b:  17);  mandibular  fragment,  AMNH 
23568;  Ml,  AMNH  23582;  M2,  AMNH  23569;  two 
ml,  AMNH  23565,  23567;  various  other  fragmentary 
teeth,  AMNH.  Simpson  reported  that  a  large  and 
a  small  kind  of  canid  were  represented  both  at  Semi- 
nole Field  and  Sabertooth  Cave.  All  of  the  material 
that  I  examined  appears  to  be  within  the  range  of 
variation  of  C.  dirus.  A  small  P4  listed  by  Simpson 
may  possibly  have  belonged  to  a  large  red  wolf. 
Webb  (1974b:17),  and  Martin  and  Webb  (1974: 
128)  reported  C.  familiaris  from  Seminole  Field.  A 
C-14  date  of  only  2,040±90  B.P.  for  this  site  was 
questioned  by  Hester   (1960). 

Melbourne,  Brevard  County;  Wisconsin  ( Hib- 
bard, et  al.,  1965);  as  C.  (A.)  cf.  ayersi  (Gazin, 
1950:400),  as  A.  cf.  ayersi  (Ray,  1958:433),  as 
C.  dirus  (Webb,  1974b:  17);  mandible,  USNM 
12946;   two  isolated  P4,  two   Ml,   four  ml,   USNM. 

Sebastian  Canal,  Brevard  County;  Wisconsin;  as 
C.  dirus  (Webb,  1974b:  17). 

Vero  (stratum  2),  Indian  River  County;  late 
Wisconsin  (Webb,  1974b:13);  as  C.  ayersi  (Sellards, 
1916:152;  Weigel,  1962:37),  as  C.  dirus  (Martin, 
1974:73);  skull  without  mandibles,  FGS  7166.  The 
status  of  C.  ayersi,  the  type  of  which  was  obtained 
at  Vero,  is  discussed  in  the  above  "remarks." 

Bradenton,  Manatee  County;  Sangamon  (Webb, 
1974b:  13);  maxillary  fragment,  UF  3276;  mandibular 
fragment,  UF  2259. 

Phillipi  Creek-Fruitville  Ditch,  7  mi.  E  Sarasota, 
Sarasota  County;  Wisconsin;  as  C.  ayersi  (Simpson, 
1929b:275). 

IDAHO. — Jaguar  Cave,  Beaverhead  Mountains, 
Lemhi  County;  late  Wisconsin  (C-14  dates:  10,370 
±350  and  11,580±250  B.P.);  as  C.  cf.  dirus  (Kur- 
ten  and  Anderson,   1972:24). 

American  Falls,  Power  County;  Rancholabrean 
(Hibbard,  et  al,  1965),  Illinoian  (Kurten,  1974:7); 
as  ef.  A.  dirus  (Gazin,  1935:298),  as  C.  (A.)  dirus 
(Hopkins,  Bonnichsen,  and  Fortsch,  1969:3).  Ga- 
zin's  original  faunal  list  for  this  site  stated  only  that 
distal  portions  of  two  humeri  and  an  abraded  phal- 
ange could  not  be  distinguished  from  corresponding 
parts  of  dire  wolves  from  Rancho  La  Brea.  This 
material  is  not  reliable  in  the  identification  of  C. 
dirus,   and  might   represent  some   other  large   canid. 

Rainbow  Beach  local  fauna,  American  Falls  Res- 
ervoir, Power  County;  Wisconsin  (C-14  dates: 
21,500±700  and  31,300±2,300  B.P.);  as  C.  dirus 
(McDonald  and  Anderson,   1975:26). 

ILLINOIS. — Galena,  Jo  Daviess  County;  Wiscon- 
sin (Kurten,  1974:10);  as  C.  or  A.  mississippiensis 
(Hay,  1923:337). 

INDIANA. — Ohio  River,  below  Evansville,  Van- 
derburgh  County;   late  Pleistocene;   as   C.   primaevus 


114 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


(Leidy,  1854:200;  1856:167),  as  C.  dims  (Leidy, 
1858:21;  Merriam,  1912:240),  as  C.  indianensis 
(Leidy,  1869:368),  as  C.  lupus  (Cope  and  Wort- 
man,  1884:9),  as  A.  dims  (Hay,  1923:204);  maxil- 
lary fragment,  ANSP  11614.  The  nomenclatural  his- 
tory of  the  type  specimen  of  C.  dirus  was  reviewed 
by  Merriam  (1912:218-221).  As  he  noted  (1912: 
240-241),  the  type  resembles  specimens  from  Rancho 
La  Brea  in  the  reduction  of  the  hypocone  of  Ml, 
as  well  as  in  other  features  that  can  be  evaluated. 
There  is  a  pronounced  buccal  cingulum  on  the  Ml, 
as  found  in  some  specimens  of  C.  dints,  and  the 
teeth  are  within  the  size  range  of  those  from  Cali- 
fornia. 

KANSAS. — Twelve  Mile  Creek,  Logan  County; 
Pleistocene;  as  C.  occidcntalis  (Hay,  1924:143,  165); 
P4,   M2,  p4,  KU  392. 

Pendennis,  Lane  County;  Pleistocene;  as  C.  occi- 
dentalis  (Hay,  1924:71);  ml,  KU  393. 

Cragin  Quarry  local  fauna,  north  of  Cimarron 
River,  Meade  Countv;  Sangamon;  as  "C.  occiden- 
talis?"  (Hay,  1917b:48),  as  A.  dims  ( Hibbard,  1939: 
464;  1949:84;  Hibbard  and  Taylor,  1960:178),  as 
C.    dirus    (Schultz,    1969:53);    mandible,    KU    4613. 

KENTUCKY.— Welsh  Cave,  3.5  mi.  SW  Troy, 
Woodford  County;  late  Wisconsin  (ca.  13,000  B.P.); 
as  C.  dirus  (Guilday,  Hamilton,  and  McCrady,  1971: 
274);  cast  of  skull  without  mandibles,  CM  12625; 
cast  of  mandible  from  different  individual,  CM 
12625a.  The  specimens  are  among  the  most  complete 
that  have  been  collected  at  sites  outside  of  the 
southwest,  and  in  all  characters  they  resemble  speci- 
mens from  Rancho  La  Brea. 

LOUISIANA.— Avery  Island,  Iberia  Parish;  late 
Pleistocene;  as  C.  dirus  (Gagliano,   1967:40). 

MISSOURI.— Brynjulfson  Caves,  6  mi.  SSE  Co- 
lumbia, Boone  County;  late  Wisconsin  (about  10,000 
B.P.);  as  C.  dirus  (Parmalee  and  Oesch,  1972:31); 
isolated  teeth,  ISM.  On  the  basis  of  radii,  Parmalee 
and  Oesch  reported  that  the  wolves  from  this  site 
were  slightly  larger  than  the  huge  individual  found 
at  Powder  Mill  Creek  Cave   (see  below). 

Cherokee  Cave,  St.  Louis,  St.  Louis  County;  late 
Pleistocene  (Webster,  1964);  as  Canis  (Simpson, 
1949:16).  Simpson  reported  that  eight  metapodials 
were  larger  than  those  of  C.  dirus,  but  he  did  not 
refer  them  to  a  species.  The  measurements  he  listed 
are  much  greater  than  the  means  given  by  Nigra  and 
Lance  (1947)  for  the  same  elements  of  C.  dirus 
from  Rancho  La  Brea;  but  the  size  of  more  recently 
collected  metapodials  of  Missouri  C.  dirus,  found  in 
association  with  cranial  material,  is  close  to  that  of 
the  Cherokee  Cave  specimens  (Galbreath,  1964; 
Hawksley,  1963).  Galbreath 's  specimen  (see  ac- 
count of  Powder  Mill  Creek  Cave,  below)  reportedly 
represented  a  female,  and  a  male  of  the  same  popu- 
lation probably  would  have  had  metapodials  as  large 
as  those  found  in  St.  Louis. 

Herculaneum  (near),  Jefferson  County;  Wiscon- 
sin (Hibbard,  et  al,  1965);  as  C.  dirus  (Olson, 
1940:42);  P4,  M2,  FM  WC1736. 

Carroll    Cave,    Camden    County;    Wisconsin;    as 


A.  dirus  (Hawksley,   1963),  as  C.  dints  (Hawksley, 
1965:79). 

Perkins  Cave,  Camden  County;  Wisconsin;  as 
C.  cf.  dirus  (Hawksley,  1965:82). 

Bat  Cave,  8  km.  NW  Waynesville,  Pulaski  Coun- 
ty; late  Wisconsin  (10,000-16,000  B.P.);  as  A.  dints 
(Hawksley,  1963),  as  C.  dirus  (Hawksley,  1965:81; 
Hawksley,  Reynolds,  and  Foley,  1973:72-77).  Ac- 
cording to  these  last  authors,  data  from  Bat  Cave 
tended  to  bear  out  Galbreath's  (1964)  suggestion 
that  dire  wolves  from  Missouri  were  larger  than  those 
from  Rancho  La  Brea  (see  account  of  Powder  Mill 
Creek  Cave,  below ) . 

Cox  Cave,  Pulaski  County;  late  Pleistocene;  "pos- 
sibly Canis  dints"   ( Mehl,   1962:44). 

Bushwacker  Cave,  Pulaski  County;  Wisconsin; 
as  C.  dirus  (Hawksley,  Reynolds,  and  Foley,  1973: 
73). 

Powder  Mill  Creek  Cave,  Shannon  County;  late 
Wisconsin  (C-14  date:  13,170±600  B.P.);  as  C. 
(A.)  dirus  (Galbreath,  1964).  Galbreath  reported 
the  discovery  of  most  of  the  skeleton,  but  not  in- 
cluding the  upper  parts  of  the  skull,  of  a  large 
female  dire  wolf.  Most  of  the  postcranial  measure- 
ments were  found  to  exceed  those  of  even  the  largest 
reported  specimens  of  C.  dims  from  Rancho  La  Brea 
(as  listed  by  Merriam,  1912;  Nigra  and  Lance,  1947; 
and  Stock  and  Lance,  1948).  Galbreath  thus  con- 
sidered the  limbs  and  feet  of  the  Missouri  individual 
to  be  relatively  larger  than  those  of  specimens  from 
the  tar  pits.  The  size  of  this  individual,  and  of  others 
reported  by  Hawksley  (1963),  led  Galbreath  to 
suggest  the  possibility  that  Missouri  dire  wolves 
averaged  larger  than  those  of  California. 

Zoo  Cave,  1  mi.  ENE  Hilda,  Taney  County;  late 
Wisconsin  (9,000-13,000  B.P.);  as  C.  dirus  (Hood 
and  Hawksley,  1975:25,  28).  According  to  these 
authors  the  material  from  this  site  represented  an 
adult  that  was  "quite  small  by  Missouri  standards, 
closely  approaching  the  size  of  Rancho  La  Brea 
specimens." 

NEBRASKA. — Heckendorf  gravel  pit,  Stanton 
County;  late  Pleistocene;  cranial  fragment,  UN  2911. 

NEVADA.— Gypsum  Cave,  16  mi.  E  Las  Vegas, 
Clark  County;  late  Wisconsin;  as  "Canis  or  Aenocyon 
sp."  (Harrington,  1933:192),  as  C.  dints  (Hester, 
1960:69). 

NEW  MEXICO.— Conkling  Cavern,  near  Las 
Cruces,  Dona  Ana  County;  late  Pleistocene;  mandi- 
ble, LACM. 

Hermit's  Cave,  east  slope  of  Guadalupe  Moun- 
tains, Eddy  County;  late  Wisconsin  (C-14  dates: 
11,850±350  and  12,900±350  B.P.);  as  C.  dirus 
(Schultz,  Martin,  and  Tanner,  1970);  maxillary  frag- 
ment, UN  19212;  cranial  fragment,  UN  19215;  man- 
dibular  fragment,   UN    19213. 

OKLAHOMA. — Marlow,  Stephens  County;  late 
Pleistocene;   skull  with  mandibles,  USNM   10278. 

OREGON.— Willamette  Valley,  near  Woodburn, 
Marion  County;  late  Pleistocene;  as  A.  dirus  (Pack- 
ard, 1950:89). 

Fossil  Lake,  Lake  County;  early  or  middle  Wis- 


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115 


consin   (Allison,  1966:32);  as  C.  cf.  dims  (Elftman, 
1931:5). 

PENNSYLVANIA.— Frankstown  Cave,  Blair 
County;  Wisconsin  (Hibbard,  1958);  as  C.  dims 
(Peterson,  1926:282);  maxillary  fragment,  CM 
11023;  three  mandibular  fragments,  CM  11022, 
11024,  11026.  Although  the  Frankstown  material 
is  referable  to  C.  dims,  there  is  some  approach  in 
dental  characters  to  C.  armbrusteri  from  Cumberland 
Cave,  only  about  50  miles  away.  More  accurate  age 
estimates  of  these  two  sites  would  be  desirable,  so 
that  we  might  evaluate  the  idea  of  the  Frankstown 
specimens  representing  a  transition  between  C.  arm- 
brusteri and  C.  dims. 

TENNESSEE. — Jewell  Cave,  near  Ruskin,  Dick- 
son County;  late  Pleistocene;  as  "wolf"  (Barr,  1961: 
178),  as  C.  dims  (Corgan,   1976). 

Robinson  Cave,  8  mi.  SW  Livingston,  Overton 
County;  Wisconsin;  as  C.  dims  (Guilday,  Hamilton, 
and  McCrady,  1969:60). 

Whitesburg,  Hamblen  County;  late  Pleistocene; 
as  "A.  ayersi?"  (Hay,  1921:95);  isolated  incisors  and 
premolars,  USNM   8997. 

TEXAS. — Tule  Canyon,  Briscoe  County;  Pleisto- 
cene; as  C.  indianensis  (Cope,  1895:453),  as  C.  dims 
(Merriam,  1912:242). 

Slaton  quarry,  5  mi.  N  Slaton,  Lubbock  County; 
Sangamon  (Hibbard,  1970);  as  Aenocyon  sp.  (Dal- 
quest,  1967:10). 

Pemberton  Hill,  Denton  County;  Sangamon;  as 
Aenocyon  sp.  (Slaughter,  et  al,  1962:17). 

Moore  Pit  local  fauna,  Dallas,  Dallas  County; 
Sangamon;  as  A.  cf.  dims   (Slaughter,   1966b:79). 

Williams  Cave,  southern  end  of  Guadalupe  Moun- 
tains, Culberson  County;  late  Pleistocene;  as  C.  dims 
(Ayer,  1936:608). 

Scharbauer  site,  south  of  Midland,  Midland 
County;  Wisconsin;  as  C.  dims  or  ayersi  (Wendorf, 
Krieger,  and  Albritton,   1955:113). 

Clamp  Cave,  San  Saba  County;  early  Recent; 
as  A.  dims  ( Lundelius,  1967:293). 

Laubach  Cave,  Georgetown,  Williamson  County; 
Wisconsin  (Kurten,  1974:9);  as  C.  cf.  dims  (Slaugh- 
ter, 1966a:481). 

Levi  shelter,  Travis  County;  late  Wisconsin;  as 
A.  dims  (Lundelius,   1967:293). 

Friesenhahn  Cave,  near  Bulverde,  Bexar  County; 
Wisconsin;  as  "A.  dims?"  (Hay,  1921:141),  as  A. 
dims   (Lundelius,    1960:38). 

Kincaid  shelter,  Uvalde  County;  late  Pleistocene; 
as  A.  dims  (Lundelius,  1967:293). 

Blanco  Creek,  Bee  County;  late  Pleistocene;  as 
A.  ayersi   (Sellards,  1940:1636). 

Ingleside  gravel  pit,  San  Patricio  County;  Wiscon- 
sin; as  A.  dims  (Lundelius,  1962),  as  C.  dims  (Lun- 
delius, 1972:12);  skull  without  mandibles,  mandibu- 
lar fragment,  Ml,  various  other  fragments,  TM.  The 
skull  is  crushed,  but  appears  to  be  the  largest  of 
C.  dims  that  I  examined.  Lundelius  (1972:12,  20) 
observed  that  two  skulls  from  the  Ingleside  fauna 
resembled  the  type  of  C.  ayersi  more  than  they  did 
skulls  from  Rancho  La  Brea,  but  that  there  was  no 


basis    for    considering    C.    ayersi    a    separate    species 
from  C.  dims. 

UTAH.— Silver  Creek  local  fauna,  5  mi.  N  Park 
City,  Summit  County;  late  Sangamon  to  early  Wis- 
consin; as  C.  cf.  dims  (Miller,  1976:401). 

VIRGINIA.— Clark's  Cave,  12  km.  SW  Williams- 
ville,  Bath  County;  late  Wisconsin  (less  than  10,000 
B.P.);  as  C.  cf.  dims  (Guilday,   1977:69). 

WEST  VIRGINIA. — Rennick,  Greenbrier  County; 
late  Pleistocene;  mandible,  CM  24327. 

WISCONSIN.— Blue  Mounds,  Dane  County;  late 
Pleistocene;  as  C.  mississippiensis  (Allen,  1876:49; 
Hay  1914:484;  1923:342),  as  C.  dims  (Merriam, 
1912:221);  four  limb  bones,  MCZ  10988-10991. 
Allen  originally  referred  to  this  record  as  being  from 
only  the  "Lead  Region  of  Upper  Mississippi,"  but 
Hay  (1923:342)  restricted  the  locality  to  Blue 
Mounds.  Allen  thought  that  the  great  size  of  the 
bones  warranted  their  referral  to  a  distinct  species, 
but  he  compared  them  only  to  a  single  small  indi- 
vidual of  C.  lupus.  Hay's  continued  recognition  of 
C.  mississippiensis  also  was  based  on  scanty  compara- 
tive material.  The  measurements  of  length  listed  by 
Allen  actually  fall  within  the  size  range  of  both 
C.  lupus  and  C.  dims  as  given  by  Stock  and  Lance 
(1948:82),  but  are  closer  to  the  means  of  the  latter 
species.  It  seems  best  for  now  to  follow  Merriam 
(1912)  in  synonymizing  C.  mississippiensis  under 
C.  dims. 

AGUASCALIENTES.— Cedazo  local  fauna,  near 
City  of  Aguascalientes;  early  Rancholabrean  (prob- 
ably Illinoian);  as  C.  dims  (Mooser  and  Dalquest, 
1975:788);  four  mandibular  fragments,  Midwestern 
State  University  Department  of  Biology  9781-9784. 
These  specimens,  clearly  referable  to  C.  dims,  pro- 
vide the  only  well  supported  record  of  a  pre-Sanga- 
mon  dire  wolf.  Actually,  however,  there  is  some 
question  about  the  age  of  the  fauna,  as  Mooser  and 
Dalquest  (1975:783)  had  stated:  "Early  Ranchola- 
brean age  (Savage,  1951)  is  indicated.  We  think 
the  Cedazo  local  fauna  could  be  as  old  as  Yarmouth- 
ian  or  as  young  as  Sangamon,  but  favor  Illinoian 
Age." 

JALISCO.— Lago  de  Chapala;  late  Pleistocene; 
as  "Canis  sp.,  large  wolf"  (Downs,  1958). 

ESTADO  DE  MEXICO.— Tequixquiac  (near); 
late  Pleistocene;  as  C.  dims  (Merriam,  1912:243), 
as  A.  dims  (Furlong,  1925:152;  Maldonado-Koerdell, 
1955);  cast  of  cranial  fragment,  UCMP  27615. 

NUEVO  LEON. — San  Josecito  Cave,  near  Aram- 
berri;  Wisconsin;  as  Aenocyon  (Russell,  1960:541); 
two  skulls,  LACM  3106,  9795;  27  cranial  and  maxil- 
lary fragments,  LACM;  30  mandibular  fragments, 
LACM.  This  large  amount  of  material  represents  a 
population  not  differing  in  characters  from  that  of 
Rancho  La  Brea. 

PUEBLA. — Valsequillo,  near  Puebla;  late  Pleisto- 
cene; as  C.   (A.)   dims  (Thenius,  1970:59). 

PERU.— La  Brea,  30  mi.  SE  Talara  (northern 
part  of  Peru,  not  mapped  in  Fig.  54);  late  Pleisto- 
cene; as  C.  (A.)  dims  (Churcher,  1959). 


116 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


Evolutionary  position. — The  species  C. 
dints  is  known  only  from  the  late  Pleistocene 
and  early  Recent,  and  is  the  most  common 
fossil  wolf  of  that  period.  The  dire  wolf  was 
not  an  ancestral  species,  but  rather  a  highly 
specialized  animal,  well  adapted  for  life  in 
the  megafaunal  community  of  its  time.  In 
its  large  size,  broad  proportions,  large  teeth, 
and  other  critical  characters,  it  stood  on  the 
opposite  end  of  the  evolutionary  line  from 
the  small  species  of  Canis  of  the  Pliocene  and 
early  Pleistocene.  Its  initial  appearance  in 
the  southern  part  of  the  continent  may  be  cor- 
related with  a  northern  withdrawal  of  C. 
lupus.  Hay  (1927:192)  speculated  that  C. 
dims  was  restricted  to  a  more  southerly  range 
by  the  presence  of  the  gray  wolf  in  the  north. 

Both  the  geographic  and  phylogenetic 
origin  of  the  dire  wolf  are  unknown.  Kurten 
(1968:109)  suggested  that  C.  falconeri,  a 
large  wolf  of  the  early  Pleistocene  of  Europe, 
might  be  related  to  C.  dims  of  the  New 
World.  But  there  is  no  chronological  or  geo- 
graphic evidence  to  support  recognition  of  a 
connection  between  the  two,  and  the  meas- 
urements listed  by  Del  Campana  (1913:220- 
229)  indicate  that  the  skull  of  C.  falconeri 
did  not  closely  approach  that  of  C.  dims  in 
size. 

Martin  (1974:76)  suggested  that  a  popu- 
lation represented  by  C.  armbrusteri  of  Cum- 
berland Cave,  Maryland  may  have  given  rise 
to  C.  dims.  This  view  is  reasonable  in  that 
the  disappearance  of  C.  armbrusteri  in  the 
Illinoian  coincided  with  the  initial  appear- 
ance of  C.  dirus  (see  account  of  C.  armbrus- 
teri). Moreover,  some  specimens  of  C.  arm- 
brusteri approach  those  of  C.  dirus  in  size, 
and  the  two  species  share  other  characters 
such  as  in  the  morphology  of  the  lower  pre- 
molars. As  yet,  however,  there  is  no  conclu- 
sive evidence  to  indicate  immediate  relation- 
ship between  C.  armbrusteri  and  the  dire 
wolf. 

Apparently  C.  dims  developed  exclusively 
in  the  Western  Hemisphere,  and  its  ancestry 
probably    lies    in    the    basic    stock    of    small 


wolves  represented  by  C.  edwardii  and  C. 
rufus.  At  some  point  in  the  Pleistocene,  an 
element  of  this  stock,  comprised  of  individ- 
uals resembling  either  C.  eduardii,  C.  rufus, 
or  C.  armbrusteri,  must  have  become  isolated 
and  begun  separate  evolution.  But  how  was 
C.  dirus  able  to  appear  suddenly  all  across 
North  America  in  the  late  Pleistocene,  with 
the  most  distinctive  set  of  characters  in  the 
genus  Canis  already  fully  developed? 

One  hypothesis  that  can  not  now  be  disre- 
garded is  that  the  dire  wolf  arose  and  devel- 
oped in  South  America.  There  are  several 
pieces  of  evidence  to  support  this  idea.  First, 
C.  dirus  has  been  reported  from  South  Amer- 
ica, specifically  from  the  La  Brea  tar  pits  near 
Talara,  Peru  (Churcher,  1959).  The  species 
also  is  known  from  several  sites  in  Mexico, 
and  at  one  time  probably  was  distributed 
throughout  that  country  and  Central  America. 
Intriguingly,  the  earliest  occurrence  of  the 
species,  that  is  supported  by  good  evidence, 
also  is  among  the  most  southerly  (see  ac- 
count of  Cedazo  local  fauna,  Aguascalientes, 
above).  The  known  range  of  the  dire  wolf 
(Fig.  54)  suggests  a  southern,  warmth  adapt- 
ed species,  in  contrast  to  the  boreal  C.  lupus. 
The  primitive  stock  of  small  wolves,  repre- 
sented by  the  living  C.  rufus,  also  seems  to 
have  been  warmth  adapted  to  some  degree. 
Factors  associated  with  one  of  the  glaciations 
may  have  driven  an  element  of  this  stock 
into  South  America  where  it  eventually 
evolved  into  C.  dirus.  Possibly  the  Sangamon 
interglacial  afforded  the  opportunity  for  re- 
invasion  of  much  of  North  America.  Further 
evidence  is  offered  by  available  information 
on  large  South  American  fossil  Canis  (L. 
Kraglievich,  1928;  J.  L.  Kraglievich,  1952: 
63).  One  specimen  in  particular,  C.  nehringi 
from  the  province  of  Buenos  Aires  in  Argen- 
tina, appears  to  have  points  of  resemblance 
to  C.  dims.  L.  Kraglievich's  photographs  and 
measurements  show  that  the  skull  of  C. 
nehringi  shares  at  least  the  following  char- 
acters with  C.  dirus:  large  size  and  massive 
proportions,   broad  frontal  shield,   prominent 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


117 


sagittal  crest,  narrow  supraoccipital  shield 
projecting  far  posteriorly,  vertical  plates  of 
palatines  flaring  broadly  anteriorly,  postpala- 
tine  foramina  opposite  posterior  ends  of  P4, 
and  relatively  large  carnassial  teeth.  Of 
course  proof  of  relationship  between  C.  dints 
and  C.  nehringi  would  not  in  itself  establish 
South  American  origin  for  the  dire  wolf. 

Although  its  origin  remains  a  mystery,  the 
dire  wolf  was  clearly  a  common  mammal  of 
the  North  American  late  Pleistocene.  The 
nature  of  its  fossil  remains  suggests  that  it 
was  found  primarily  in  open  lowlands,  and 
was  a  predator  of  its  contemporary  large 
herbivores.  The  extinction  of  most  of  this 
megafauna  at  the  close  of  the  Pleistocene,  for 
any  or  all  of  the  reasons  discussed  by  Martin 
and  Wright  ( 1967 ) ,  probably  also  signaled 
the  end  of  the  dire  wolf.  An  additional  factor 
in  the  extinction  of  C.  dims  may  have  been  a 
renewed  influx  of  gray  wolves  following  the 


withdrawal  of  the  Wisconsin  ice  sheet.  A 
general  consensus  among  authors  who  have 
speculated  on  the  behavior  of  C.  dims  is 
that  it  was  a  powerful  creature,  but  was 
slower  and  possibly  not  so  alert  as  C.  lupus 
(Matthew,  1916;  Merriam,  1912:218;  Scott, 
1937:578;  Stock,  1956:32;  Stock  and  Lance, 
1948).  The  dire  wolf  may  not  have  been  so 
well  adapted  in  the  pursuit  of  the  predomi- 
nantly smaller,  swifter  herbivores  that  sur- 
vived through  the  Recent,  and  it  may  have 
lost  in  competition  with  the  gray  wolf.  The 
sympatric  occurrence  of  the  two  species  is 
demonstrated  by  good  cranial  material  from 
Rancho  La  Rrea,  the  Maricoa  Brea,  San 
Josecito  Cave,  and  Hermit's  Cave.  Fossils  of 
both  species  also  have  been  reported  from 
Fossil  Lake,  Jaguar  Cave,  Samwel  Cave,  Pot- 
ter Creek  Cave,  Ventana  Cave,  Blue  Mounds, 
Brynjulfson  Caves,  and  Medicine  Hat. 


SUMMARY 


Systematic  problems  in  the  genus  Cants 
center  on  its  paleontological  history  and  on 
Recent  populations  in  eastern  North  America. 
In  order  to  investigate  these  matters,  approxi- 
mately 5,000  specimens  were  examined.  Many 
of  these  were  complete  skulls,  15  measure- 
ments of  which  were  utilized  in  the  BMD07M 
computer  program  of  multivariate  analysis. 
For  statistical  purposes,  material  was  sepa- 
rated by  sex,  except  for  fossils  and  specimens 
of  domestic  dogs. 

The  gray  wolf  (Canis  lupus)  and  the  coy- 
ote (Canis  latrans)  are  readily  distinguished 
from  one  another,  and  from  the  domestic  dog 
(Canis  familiaris).  By  multivariate  analysis, 
only  five  skulls  of  wild  Canis  from  northern 
and  western  North  America  appeared  to  rep- 
resent hybrids.  The  remaining  379  specimens 
of  C.  lupus  and  277  of  C.  latrans  from  these 
regions,  along  with  a  series  of  50  C.  familiaris, 
were  used  as  standard  groups  with  which  to 
compare  individuals  taken  in  the  east.  The 
subspecies  C.  lupus  hjcaon,  which  has  been 
nearly  exterminated  in  the  eastern  United 
States,  still  survives  in  the  upper  Great  Lakes 
region,  as  well  as  in  southeastern  Canada. 
Nearly  all  specimens  that  had  been  previously 
identified  as  hjcaon  showed  close  statistical 
affinity  to  the  standard  sample  of  C.  lupus, 
and  thus  were  combined  with  that  sample. 

Available  information  indicates  that  by 
1900  the  coyote  had  begun  to  extend  its  range 
to  the  east  and  north  of  the  prairies.  The 
subspecies  C.  latrans  thamnos,  of  the  north- 
central  United  States  and  southeastern  Can- 
ada, is  statistically  close  to  western  C.  latrans. 
A  few  specimens,  however,  suggest  that  lim- 
ited wolf-coyote  hybridization  has  occurred 
recently  in  southern  Ontario  and  Quebec,  and 
has  allowed  introgression  of  genes  from  C. 
lupus  into  C.  latrans.  As  a  result,  the  multi- 
variate position  of  the  coyote  population  now 
expanding  through  the  northeastern  United 
States  is  shifted  in  the  direction  of  the  wolf. 


Hybridization  of  C.  latrans  and  C.  familiaris 
also  has  taken  place,  but  has  not  had  sub- 
stantial effect  on  wild  species  of  Canis. 

In  historical  time,  the  red  wolf  (C.  rufus) 
inhabited  the  region  from  central  Texas  to 
the  Atlantic,  and  from  the  Gulf  Coast  to  the 
Ohio  Valley  and  Pennsylvania.  The  14  earli- 
est available  specimens,  from  the  part  of  this 
region  that  was  well  separated  from  the  orig- 
inal range  of  the  coyote,  show  no  statistical 
overlap  with  the  standard  samples  of  482  C. 
lupus  (including  103  hjcaon),  277  C.  latrans, 
and  50  C.  familiaris.  An  additional  115  skulls 
collected  from  1919  to  1929  in  Arkansas,  Lou- 
isiana, southern  Missouri,  and  eastern  Okla- 
homa, and  previously  identified  as  C.  rufus 
gregoryi,  have  almost  the  same  multivariate 
distribution.  These  skulls,  plus  most  of  the 
older  specimens,  were  combined  to  make  a 
standard  red  wolf  sample  of  125  individuals. 
This  sample  and  the  standard  coyote  sample 
were  used  to  compare  all  other  southeastern 
material. 

Series  of  specimens  taken  prior  to  1930 
indicate  that  hybridization  between  C.  rufus 
and  C.  latrans  generally  was  uncommon 
where  their  ranges  approached  or  overlapped, 
except  in  the  Edwards  Plateau  area  of  central 
Texas.  Material  from  that  area  forms  a  sta- 
tistical bridge  between  the  ranges  of  varia- 
tion of  the  two  standard  samples.  Subse- 
quently, as  the  red  wolf  became  rare,  hybridi- 
zation increased  along  the  Texas  coast,  and 
in  north-central  Texas,  eastern  Oklahoma, 
southern  Missouri,  and  Arkansas.  This  inter- 
breeding apparently  allowed  introgression  of 
red  wolf  genes  into  the  expanding  coyote 
population,  which  by  the  1960's  had  become 
established  in  most  inland  areas  of  the  south- 
central  states.  This  population  is  essentially 
coyotelike,  but  is  shifted  statistically  in  the 
direction  of  the  red  wolf,  and  contains  a  few 
individuals  that  are  phenotypically  close  to 
C.  rufus. 


118 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


119 


Material  collected  in  the  1960's  and  1970's 
shows  that  the  genetic  influence  of  the  red 
wolf  remained  strong  within  100  miles  of  the 
Texas  coast.  Samples  from  most  localities 
there  fall  mainly  between  the  statistical  dis- 
tributions of  C.  rufas  and  C.  latrans.  Until 
about  1970,  an  unmodified  population  of  the 
red    wolf    survived    in    extreme    southeastern 


Texas    and    probably    in    adjacent    parts    of 
Louisiana. 

The  genus  Canis  apparently  arose  by  the 
middle  Pliocene  (Hemphillian),  and  its  sub- 
sequent hypothetical  evolution  is  shown  in 
figure  55.  The  relationships  of  C.  cedazoen- 
sis  of  Mexico  are  not  well  understood,  but 
otherwise  the  North  American  species  can  be 


OLD 
WORLD 


NEW    WORLD 


RECENT 


RANCHO- 
LABREAN 


SI  ME  AD  AU 


LATRANS 


IRVING- 
TONIAN 


BLANCAN 


CEDAZO- 
ENSIS 


--?-- 


RUFUS 


ARMBRUSTERI 


EDWARD  I 


I 

OLD 
WORLD 

i 

FAMIL- 
LUPUS  IARIS 

DIRUS        %  __J 


'LEPO- 

Iphagus 


HEMP- 
HILLIAN 


J 

r 


--?-■ 


Ietruscus 

1 


Fig.  55. — Hypothetical  phylogenetic  diagram  of  the  evolution  of  Canis.  Dashed  lines  indicate  possible 
lineages  in  which  fossil  evidence  is  lacking.  Names  of  species  are  placed  at  the  latest  levels  at  which  those 
species  are  known.  Only  names  of  species  recognized  in  this  paper  are  shown.  Horizontal  and  vertical  dis- 
tances are  not  necessarily  to  scale,  and  do  not  indicate  degree  of  affinity.  The  following  species  are  abbre- 
viated: C.  simensis  (SI),  C.  mesomelas  (ME),  C.  adustus  (AD),  C.  aureus  (AU).  Question  marks  (?) 
indicate  alternative  lineages. 


120 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


separated  into  coyote  and  wolf  groups.  The 
Blancan  C.  lepophagus  was  a  variable  entity, 
with  some  individuals  resembling  small  coy- 
otes, and  others  having  certain  wolflike  char- 
acters. It  is  known  from  15  localities,  from 
Florida  to  Idaho.  The  species  C.  latrans  prob- 
ably descended  from  certain  populations  of 
C.  lepophagus  by  the  end  of  the  Blancan,  and 
subsequently  there  appears  to  have  been 
little  change  in  the  coyote  line.  Some  Pleisto- 
cene coyotes,  especially  those  of  the  late 
Rancholabrean  of  California,  became  larger 
than  modern  C.  latrans,  but  others  were  about 
the  same  size  or  smaller.  Fossil  C.  latrans  has 
been  reported  from  109  localities  across  North 
America,  from  Florida  to  Alaska,  and  from 
Oaxaca  to  Pennsylvania. 

It  is  not  clear  whether  the  wolf  group  de- 
scended from  some  population  of  C.  lepopha- 
gus, or  was  already  distinct  from  the  coyote 
line  throughout  the  Blancan.  Several  species 
of  wolves  appeared  by  the  early  Irvingtonian, 
the  first  of  which  was  probably  C.  edwardii. 
This  was  a  small  species,  known  from  a  few 
localities  in  the  southwestern  quarter  of 
North  America.  An  immediate  relative,  C. 
rufus  of  the  southeast,  continued  to  represent 
the  primitive  stock  of  wolves  until  Recent 
times.  Still  another  Irvingtonian  wolf,  C. 
armbrusteri,  was  much  larger,  but  resembled 
the  red  wolf  in  certain  dental  characters  and 
skull  proportions.  It  is  known  by  good  ma- 
terial only  from  Maryland  and  Florida, 
though  fragmentary  remains  from  elsewhere 
suggest  that  it  may  once  have  occurred  all 
across  the  continent. 

During  the  early  Pleistocene,  an  element 


of  the  primitive  stock  of  small  wolves  appar- 
ently entered  Eurasia  where  it  gave  rise  to 
C.  lupus.  In  the  course  of  the  Illinoian  gla- 
ciation  this  species  probably  moved  into 
North  America,  where  its  fossils  have  been 
reported  from  58  localities.  Some  Ranchola- 
brean gray  wolves  were  remarkably  small, 
but  others,  particularly  those  of  Rancho  La 
Brea,  had  massive  skulls  resembling  those  of 
some  modern  Arctic  wolves. 

The  extinct  C.  dims  did  not  appear  in 
North  America  until  the  Rancholabrean,  and 
may  have  originated  in  South  America  or 
descended  from  C.  armbrusteri.  This  large, 
highly  specialized  species  was  not  ancestral 
to  modern  wolves,  and  its  skull  is  easily  dis- 
tinguished from  that  of  C.  lupus.  By  multi- 
variate analysis,  there  was  no  overlap  be- 
tween 62  specimens  of  C.  dims  from  Rancho 
La  Brea  and  467  specimens  of  Recent  C. 
lupus.  The  dire  wolf  has  been  reported  from 
96  localities  and  apparently  was  common 
throughout  that  part  of  North  America  to 
the  south  of  Canada. 

The  generic  name  Aenocijon,  sometimes 
applied  to  the  dire  wolf,  is  here  synonymized 
under  Canis.  The  following  names  no  longer 
are  considered  to  represent  separate  species: 
C.  caneloensis,  C.  irvingtonensis,  and  C.  rivi- 
veronis  ( all  are  fossil  subspecies  of  C.  la- 
trans); C.  andersoni  (a  synonym  of  C.  latrans 
orcutti);  C.  petrolei  (a  synonym  of  C.  famil- 
iaris);  C.  priscolatrans  (a  fossil  subspecies  of 
C.  rufus);  C.  milleri  (a  synonym  of  C.  lupus 
furlongi);  and  C.  ayersi  (a  synonym  of  C. 
dims ) . 


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p.  389. 
Vanderhoof,  V.  L.,  and  J.  T.  Gregory 

1940.    A    review   of   the    genus    Aelurodon.     Univ. 
California  Publ.  Bull.   Dept.  Geol.,  25:143- 
164. 
Warfel,  H.  E. 

1937.    A  coyote  in   Hampshire  County,  Massachu- 
setts.  Jour.  Mamm.,  18:241. 
Webb,  S.  D.  (ed.) 

1974a.  Pleistocene  mammals  of  Florida.    University 
Presses  of  Florida,  Gainesville,  x-f-270  pp. 
Webb,  S.  D. 

1974b.  Chronology    of    Florida    Pleistocene    mam- 
mals.   In  Webb   (1974a),  pp.  5-31. 
Webb,  W.  S.,  and  R.  S.  Baby 

1957.    The  Adena  People.    No.  2.    Ohio  Hist.  Soc, 
Ohio  State  Univ.   Press,  xi-(-123  pp. 
Webster,  D. 

1964.  Cherokee  Cave  bone  deposit.  Missouri 
Speleol.,  16:79-86. 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


135 


Weigel,  R.  D. 

1962.    Fossil  vertebrates  of  Vero,  Florida.    Florida 
Geol.  Surv.  Spec.  Publ.,  no.  10,  vii-f  59  pp. 
Weise,   T.   F.,   W.   L.    Robinson,   R.   A.   Hook,   and 
L.  D.  Mech 

1975.  An  experimental  translocation  of  the  eastern 
timber  wolf.  Audubon  Conserv.  Rept.,  no. 
5,  28  pp. 

Wendobf,  F.,  A.  D.  Krieger,  and  C.  C.  Albbitton 
1955.    The  Midland  discovery.    Univ.  Texas  Press, 
viii+139  pp. 
Wetzel,  R.  M.,  and  L.  R.  Penner 

1962.    Coydog  in  Connecticut.    Jour.  Mamm.,  43: 
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Whit  acre,  D. 

1948.    The  mysterious  coyote  pack  of  Ohio.    Ohio 
Conserv.  Bull.,  12(3):29. 
Whitney,  J.  D. 

1879.    The    animal    remains,    not    human,    of    the 
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Williams,  C.  T. 

1962.  Classification  of  the  Borophaginae  (Cani- 
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WlLLISTON,  S.  W. 

1898.    The   Pleistocene   of   Kansas.     Trans.    Kansas 
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Wilson,  J. 

1976.  Bobcats,  bears  and  coyotes  ....  are  they 
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Wilson,  B.  L. 

1967.    The    Pleistocene    vertebrates    of    Michigan. 
Papers  Michigan  Acad.  Sci.,  Arts,  and  Let- 
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Wilson,  R.  W. 

1933.    Pleistocene  mammalian  fauna  from  the  Car- 
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Wilson,  W.  C. 

1967.  Food  habits  of  the  coyote,  Canis  latrans,  in 
Louisiana.  Unpublished  M.S.  thesis,  Louisi- 
ana State  Univ.,  ix+49  pp. 


Winc,  E.  S. 

1963.  Vertebrates  from  the  Jungerman  and  Good- 
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Contrib.  Florida  State  Mus.,  Soc.  Sci.,  10: 
51-60. 

Wolfe,  J.  L. 

1972.  Wolves  in  Mississippi?  Mississippi  Game 
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Wolfe,  M.  L.,  and  D.  L.  Allen 

1973.  Continued  studies  of  the  status,  socializa- 
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1967  to  1970.    Jour.  Mamm.,  54:611-635. 

Wolfram,  G. 

1964.  Coyotes:  the  silent  invaders.  Canadian 
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Wood,  N.  A. 

1922.    The  mammals  of  Washtenaw  County,  Mich- 
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Wood,  N.  A.,  and  L.  R.  Dice 

1924.  Records  of  the  distribution  of  Michigan 
mammals.  Papers  Michigan  Acad.  Sci., 
Arts,  and  Letters,  3:425-469. 

WOODHOUSE,  S.  W. 

1851.  The  North  American  jackal — Canis  frustror. 
Proc.   Acad.   Nat.   Sci.,   Philadelphia,  ser.   1, 

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WURSTER,  D.  H.,  AND  K.  BeNTRSCHKE 

1968.  Comparative  cytogenetic  studies  in  the  or- 
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Young,  S.  P. 

1944.  History,  life  habits,  economic  status,  and 
control.  Part  I  in  Young,  S.  P.,  and  E.  A. 
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Amer.  Wildl.  Inst.,  Washington,  D.C.,  pp. 
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1946.  The  wolf  in  North  American  history.  Cax- 
ton  Printers,  Ltd.,  Caldwell,  Ohio,   149  pp. 

1951.  History,  life  habits,  economic  status,  and 
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Jackson,  The  clever  coyote,  Wildl.  Mgmt. 
Inst.,   Washington,   D.C.,   pp.    1-226. 


Addendum 


Subsequent  to  preparation  of  the  galley 
proof  of  this  paper,  the  following  information 
came  to  my  attention. 

Canis  Jepophagus. — Bjork  (1974)  assigned 
a  newly  discovered  specimen  from  the  Wen- 
dell Fox  pasture  locality  of  the  Rexroad  fauna, 
Meade  County,  Kansas,  to  this  species.  Dal- 
quest  (1978)  listed  "Canis  cf.  lepophagus" 
from  the  Beck  Ranch  local  fauna,  Scurry 
County,  Texas. 

Canis  latrans. — Parmalee,  Munson,  and 
Guilday  (1978)  reported  specimens  of  "C. 
latrans  Say — Coyote?"  from  the  Harrodsburg 
Crevice,  Monroe  County,  Indiana.  Although 
a  radiocarbon  analysis  made  on  bones  from 
this  site  provided  a  date  of  25,050 ±660  B.P., 
some  of  the  fauna!  components  suggested 
that  a  Sangamon  age  was  more  likely.  Corner 
(1977)  reported  this  species  from  a  Ranchola- 
brean  fauna,  4.5  mi.  W  McCook,  Red  Willow 
County,  Nebraska.  Grayson  (1977)  reported 
"Canis  cf.  latrans  Coyote"  from  zones  dated 
6,500-9,500  B.P.  in  the  Dirty  Shame  Rock- 
shelter,  Malheur  County,  Oregon.  Martin, 
Gilbert,  and  Adams  ( 1977 )  listed  this  species 
from  the  late  Pleistocene  Natural  Trap  Cave, 
Big  Horn  County,  Wyoming. 

Canis  rufus. — On  5  January  1978  the  U.S. 
Fish  and  Wildlife  Service  made  a  second  re- 
lease of  a  pair  of  wild-caught  red  wolves  on 
Bulls  Island,  Cape  Romain  National  Wildlife 
Refuge,  South  Carolina.  The  animals  appar- 
ently adapted  well,  and  did  not  leave  the 
vicinity  of  Bulls  Island  and  a  small  adjacent 
island,  until  they  were  recaptured  (un- 
harmed )  on  19  October  and  1  November  1978 
to  terminate  the  experiment.  In  November 
1978,  the  breeding  colony  at  Tacoma,  Wash- 


ington and  a  second  facility  at  Winnie,  Texas 
contained  a  total  of  31  wild-caught  animals 
thought  to  be  red  wolves,  5  surviving  young 
produced  in  the  spring  of  1977,  and  15  young 
produced  in  the  spring  of  1978. 

Canis  lupus. — Corner  ( 1977 )  reported  this 
species  from  a  Rancholabrean  fauna,  4.5  mi. 
W  McCook,  Red  Willow  County,  Nebraska. 
Martin,  Gilbert,  and  Adams  (1977)  listed 
"Canis  sp.  (wolf)"  from  the  late  Pleistocene 
Natural  Trap  Cave,  Big  Horn  County,  Wyom- 
ing. 

Canis  familiaris. — Arredondo  and  Varona 
( 1974 )  described  Cubacyon  transversidens,  a 
new  genus  and  species  of  canid  from  a  Pleis- 
tocene site  in  western  Cuba.  Based  on  the 
published  description  and  an  examination  of 
specimens  of  domestic  dogs,  E.  Raymond  Hall 
(Museum  of  Natural  History,  Univ.  Kansas; 
pers.  comm.)  considers  Cubacyon  transver- 
sidens to  be  a  synonym  of  Canis  familiaris, 
and  I  agree  with  this  assessment.  Beebe 
(1978)  reported  a  specimen  of  C.  familiaris 
from  the  Old  Crow  River  Basin  of  the  north- 
ern Yukon,  with  a  minimum  age  of  20,000 
B.P.,  and  observed:  "The  highly  evolved 
morphology  of  the  specimen  suggests  a  much 
earlier  time  of  domestication." 

Canis  dims. — Parmalee,  Munson,  and 
Guilday  ( 1978 )  reported  specimens  of  "Canis 
cf.  dims"  from  the  Harrodsburg  Crevice, 
Monroe  County,  Indiana  (see  above  para- 
graph on  C.  latrans),  and  from  the  Guy 
Wilson  Cave,  Sullivan  County,  Tennessee. 
According  to  Berta  and  Marshall  (1978),  fos- 
sils referrable  to  C.  dims  have  been  reported 
in  South  America  from  Talara,  Peru;  Tarija, 
Bolivia;  and  Muaco,  Venezuela. 


Literature  Cited 


Arredondo,  O.,  and  L.  S.  Varona 

1974.    Nuevos  genero  y  especie  de  mamifero  (Car- 
nivora:    Canidae)  del  Cuaternario  de  Cuba. 
Poeyana   (Havana),  no.  131,  12  pp. 
Beebe,  B.  F. 

1978.    Two  new  Pleistocene  mammal  species  from 
Beringia.     Amer.    Quat.    Assoc,    Abstr.    5th 
Bien.  Mtg.,  p.  159. 
Berta,  A.,  and  L.  G.  Marshall 

1978.    South    American    Carnivora.     In    Westphal, 


F.   (ed.),  Fossilium  catalogus,  I:    Animalia, 
W.  Junk,  The  Hague,  part  125,  ix  +  48  pp. 

Bjork,  P.  R. 

1974.  Additional  carnivores  from  the  Rexroad  For- 
mation (upper  Pliocene)  of  southwestern 
Kansas.    Trans.  Kansas  Acad.  Sci.,  76:24-38. 

Corner,  R.  G. 

1977.  A  late  Pleistocene-Holocene  vertebrate  fauna 
from  Red  Willow  County,  Nebraska.  Trans. 
Nebraska  Acad.  Sci.,  4:77-93. 


136 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAN1S 


137 


Dalquest,  W.  W. 

1978.    Early  Blancan  mammals  of  the  Beck  Ranch 
local  fauna  of  Texas.   Jour.  Mamm.,  59:269- 
298. 
Grayson,  D.  K. 

1977.  Paleoclimatic  implications  of  the  Dirty 
Shame  Rockshelter  mammalian  fauna.  Teb- 
iwa,  no.  9,  26  pp. 


Martin,  L.  D.,  B.  M.  Gilbert  and  D.  B.  Adams 

1977.  A  cheetah-like  cat  in  the  North  American 
Pleistocene.    Science,  195:981-982. 

PARMALEE,   P.   W.,   P.   J.    MUNSON,   AND  J.   E.   GUDLDAY 

1978.  The  Pleistocene  mammalian  fauna  of  Har- 
rodsburg  Crevice,  Monroe  County,  Indiana. 
Natl.   Speleol.   Soc.  Bull.,  40:64-75. 


APPENDIX  A 


The  following  list  provides  details  on  samples 
used  in  multivariate  analyses.  Specimens  are  in  the 
USNM  unless  otherwise  indicated. 

1.  Canis  lupus  taken  not  later  than  1925  in  the 
mountainous  region  of  western   North  America. 

C.  I.  irremotus 

ALBERTA.— 25  mi.   SE   Lethbridge,    1. 

IDAHO.— Bannock  Co.:  10  mi.  E  Pocatello,  1; 
Tyhee  Basin,  1.  Caribou  Co.:  Soda  Springs,  2.  Clark 
Co.:  Argora,  1.  Lemhi  Co.:  Leadore,  1;  10  mi.  S 
Leadore,  1. 

MONTANA. — No  precise  locality,  1.  Beaverhead 
Co.:  Dillon,  1.  Carbon  Co.:  Red  Lodge,  1.  Carter 
Co.:  Ridge,  1.  Cascade  Co.:  Belt,  1.  Powder  River 
Co.:  Kruger,  1.  Rosebud  Co.:  Ingomar,  1;  Lame 
Deer,  1. 

WYOMING.— Campbell  Co.:  Gilette,  1.  Con- 
verse Co.:  Glenrock,  1;  Lost  Springs,  1.  Fremont 
Co.:  Lenore,  1;  Split  Rock,  1.  Johnson  Co.:  Barber, 
1.  Sublette  Co.:  Cora,  1;  Pinedale,  1.  Sheridan  Co.: 
Arvada,  2.  Teton  Co.:  Elk,  3;  Kelly,  1.  Yellowstone 
National  Park,  1. 

C.  I.  mogollonensis 

ARIZONA. — No  precise  locality,  1.  Apache  Co.: 
Escudilla  Mts.,  3.  Greenlee  Co.:  Clifton,  1;  15  mi. 
SW  Alma,  New  Mexico,  1.  Maricopa  Co.:  Aguila,  1. 
Navajo  Co.:    Cibecue,   1;  Heber,   1. 

NEW  MEXICO.— Catron  Co.:  Datil  Mts.,  1;  Gila 
National  Forest,  6;  Luna,  1;  15  mi.  SE  Reserve,  1. 
Grant  Co.:  head  of  Mimbres  River,  1;  Silver  City,  1. 
Sierra  Co.:  Fairview,  1;  Chlorida,  5;  Monticello,  1. 
Socorro  Co.:    Magdalena,  1. 

C.  I.  youngi 

COLORADO. — Mesa  Co.:  no  precise  locality,  1; 
Glade  Park,  2;  West  Creek,  1.  Pueblo  Co.:  25  mi. 
NW  Pueblo,  1.  Rio  Blanco  Co.:  Piceance,  2;  Sul- 
phur,  1;  Turman's  Creek,   1. 

NEW  MEXICO.— Rio  Arriba  Co.:  Abiquiu,  2; 
Canjilon,  1;  Dulce,  2;  El  Vado,  1;  Hayes,  2.  San 
Juan  Co.:  La  Plata,  1.  Sandoval  Co.:  Cuba,  3; 
Senorita,  1.  Santa  Fe  Co.:  Lamy,  1.  Valencia  Co.: 
San  Mateo,  1. 

UTAH. — No  precise  locality,  1.  Box  Elder  Co.: 
Grouse  Creek,  1.  Duchesne  Co.:  Duchesne,  1.  San 
Juan  Co.:    10  mi.  NW  Monticello,  2. 

WYOMING.— No  precise  locality,  2.  Laramie 
Co.:  Federal,  2.  Sweetwater  Co.:  Rock  Springs,  1. 
Not  located,  Black  Tail  Creek,  1. 

2.  Canis  latrans  lestes  taken  not  later  than  1925 
in  the  mountainous  region  of  western  North  America. 

COLORADO.— Conejos  Co.:  Bountiful,  1;  Ceni- 
cro,  4;  La  Jara,  4;  Rio  Grande,  1.  Delta  Co.:  Cedar 
Edge,  1;  Grand  Mesa,  2.  Garfield  Co.:  Austin,  2; 
East  Salt  Creek,  2;  Salt  Creek,  1.  Grand  Co.: 
Kremmling,  4.  Larimer  Co.:  Arkins,  3;  Loveland, 
1.  Mesa  Co.:  Mesa,  1.  Moffatt  Co.:  Craig,  2.  Park 
Co.:     South    Park,   2;   Tarryall,   3.     Rio   Blanco   Co.: 


Piceance,  6.  Rio  Grande  Co.:  Monte  Vista,  24. 
Routt  Co.:  Battle  Creek,  4;  Russell  Springs,  4; 
Steamboat  Springs,  2.  Summit  Co.:  Gore  Range,  1. 
IDAHO.— Ada  Co.:  Boise,  1.  Bannock  Co.:  no 
precise  locality,  1;  Chesterfield,  1;  McCammon,  1; 
Pocatello,  3;  Tyhee  Basin,  2.  Bingham  Co.:  Alridge, 
2;  Cerro  Grande,  4;  Ft.  Hall,  2.  Blaine  Co.:  Saw- 
tooth National  Forest,  3.  Bonneville  Co.:  John  Gray's 
Lake,  1.  Boundary  Co.:  Schnoors,  1.  Canyon  Co.: 
Bowmont,  4.  Caribou  Co.:  Preuss  Mts.,  2.  Cassia 
Co.:  Almo,  1;  Oakley,  6.  Clark  Co.:  Dubois,  1; 
Kilgore,  1;  Medicine  Lodge  Creek,  4.  Custer  Co.: 
Bigfoot  River,  2.  Elmore  Co.:  Arrow  Rock,  1. 
Gooding  Co.:  Gooding,  1.  Goodnow  Co.:  Bliss,  1. 
Idaho  Co.:  Orangeville,  1;  Rice  Creek,  2;  West  Lake, 
7;  White  Bird,  1.  Lemhi  Co.:  Leadore,  2;  Leesburg, 
1;  Salmon,  3.  Lewis  Co.:  Forest,  1;  Salmon  River, 
2.  Lincoln  Co.:  Shoshone,  1.  Owyhee  Co.:  Grand 
View,  1;  Grassmere,  1;  Hot  Springs,  2;  Three  Creek, 
8.  Payette  Co.:  French,  1.  Pegram  Co.:  Bear  Lake, 
3. 

3.  Canis  familiaris.— 50  (10  in  KU,  6  in  MCZ, 
1  in  ROM,  5  in  UArk,  3  in  USFWS). 

4.  Canis  lupus  from  northern  and  western  North 
America  (other  than  as  listed  in  1  above). 

C.  I.  alces 

ALASKA. — Kachemak   Bay,    Kenai    Peninsula,   2. 

C.  I.  arctos 

NORTHWEST  TERRITORIES.— Ellesmere  Is- 
land: Bear  Peninsula,  1  (CNM);  Eureka  Sound,  6 
(CNM);  Griese  Fjord,  2  (CNM);  Hare  Fjord,  1 
(CNM);  Slidre  Fjord,  Foshien  Peninsula,  8  (6  in 
CNM).  Graham  Island:  Norwegian  Bay,  1  (CNM). 
Prince  Patrick  Island:  Cherie  Bay,  1;  Mould  Bay, 
1  (CNM). 

C.  I.  baileyi 

ARIZONA. — Cochise  Co.:  Huachuca  Mts.,  1. 
Pima  Co.:    5  mi.  SE  Arivaca,  1;  Helvetia,  2. 

NEW  MEXICO.— Dona  Ana  Co.:  Hatch,  1. 
Grant  Co.:  Cloverdale,  2;  Hatchita,  4.  Hidalgo  Co.: 
Animas,  1;  30  mi.  SE  Animas,  2;  35  mi.  SE  Animas, 
1;  Animas  Mts.,  1;  Animas  Peak  1  (KU);  San  Luis 
Valley,  1. 

TEXAS.— Brewster  Co.:  10  mi.  S  Alpine,  1 
(SR).  Jeff  Davis  Co.:  Fort  Davis,  1.  Pecos  Co.: 
near  Longfellow,  1  (SR). 

CHIHUAHUA.— Colonia  Garcia,  1;  Colonia 
Juarez,  1;  near  corner  adjoining  Sonora,  Arizona, 
and  New  Mexico,  3. 

SONORA.— Sierra  Pinto  Mts.,  1. 

C.  I.  beothucus 

NEWFOUNDLAND  (Island).— No  precise  lo- 
cality, 3  (2  in  MCZ). 

C.   /.   bernardi    (including  all  specimens  from   Banks 
Island) 


138 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


139 


NORTHWEST  TERRITORIES  (all  in  CNM).— 
Banks  Island:  no  precise  locality,  1;  North  Adam 
River,  3;  Big  River,  1;  Egg  River,  1;  25  mi.  E  Sachs 
Harbor,  2. 

C.  I.  crassodon 

BRITISH  COLUMBIA.— Vancouver  Island: 
Quatsino  Sound,  2. 

C.  I.  fuscus 

BRITISH   COLUMBIA.— No   precise   locality,    1. 

OREGON. — Clackamas  Co.:  Clackamas  Lake,  1 
(SD).  Curry  Co.:  Rogue  River,  2.  Douglas  Co.: 
Tiller,  2;  Glide,  2.  Jackson  Co.:  25  mi.  NE  Ashland, 
1;  Peavine  Mt.,  3  (SD).  Lake  Co.:  Sycan,  1.  Lane 
Co.:  20  mi.  S  Oakridge,  1  (SD).  Linn  Co.:  Cas- 
cadia,  2. 

WASHINGTON.— Jefferson  Co.:  22  mi.  S  Port 
Angeles,  1. 

C.  I.  hudsonicus 

NORTHWEST  TERRITORIES.— Aberdeen  Lake, 
5  (CNM);  Beaver  Hill  Lake,  3  (CNM);  Cape  Ful- 
lerton,  1  (AMNH);  Hudson  Bay,  1  (AMNH);  Nuel- 
tin  Lake,  1  (CNM);  Red  River,  1  (KU);  head  of 
Schultz  Lake,  2;  Simon's  Lake,  1;  Thelon  River,  1; 
Wajer  River,  1  (AMNH). 

C.  I.  labradorius 

NEWFOUNDLAND  ( LABRADOR )  .—Porcu- 
pine, 1. 

QUEBEC    (UNGAVA).— No   precise   locality,    1. 

C.  I.  ligoni 

ALASKA. — Conclusion  Island,  1;  Ketchikan,  1; 
Kuiu  Island,  2;  Kupreanof  Island,  2;  Prince  of  Wales 
Island,  2;  Revillagigedo  Island,   1;   Wrangell,  6. 

C.  I.  mackenzii 

NORTHWEST  TERRITORIES.— Amundsen 
Gulf,  1  (CNM);  south  side  of  Coronation  Gulf,  1; 
Port  Epworth  Harbor,  1  (CNM);  head  of  Hood 
River,  1  (CNM);  Mackenzie  Delta,  3  (CNM);  Rae 
River,  1  (CNM). 

C.  I.  manningi 

NORTHWEST  TERRITORIES.— Baffin  Island: 
no  pricise  locality,  2  ( 1  in  CNM,  1  in  collection  of 
Douglas  H.  Pimlott);  Pangnirtung  Fjord,  1   (CNM). 

C.  I.  monstrabilis 

NEW  MEXICO.— No  precise  locality,  1.  Otero 
Co.:    Elk,  2;  Sacramento  Mts.,  2;  Mayhill,   1. 

TEXAS.— Crockett  Co.:  Ozona,  2.  Culbertson 
Co.:  Guadalupe  Mts.,  1.  Jack  Co.:  Fort  Richardson, 
1.  Kimble  Co.:  1.  Presidio  Co.:  40  mi.  SW  Marfa, 
1.  Reagan  Co.:  Big  Lake,  1.  Upton  Co.:  Rankin, 
6.    Ward  Co.:    Monahans,  1. 

C.  I.  nubilus 

MANITOBA.— Southeast  of  Carberry,  1  (CNM); 
Duck  Mountain,  2;  Riding  Mountain  National  Park, 
3  (CNM). 

COLORADO.— Bent  Co.:  3. 

KANSAS.— Gove  Co.:  3  mi.  W  Castle  Rock,  1. 
Trego  Co.:  near  Castle  Rock,  1   (KU). 


MINNESOTA.— Becker  Co.:  25  mi.  N  Detroit 
Lakes,  1. 

NEBRASKA.— Platte  River,  3.  Kearny  Co.:  Ft. 
Kearny,  3. 

NEW  MEXICO.— Guadalupe  Co.:  Santa  Rosa, 
1.  Lincoln  Co.:  40  mi.  SE  Corona,  1.  Socorro  Co.: 
Carthage,  3.    Torrence  Co.:    Mountain  Air,  1. 

NORTH  DAKOTA.— Billings  Co.:  Medora,  2. 
Golden  Valley  Co.:    near  Beach,  1. 

OKLAHOMA.— Panhandle  area,  1  (AMNH). 
Comanche  Co.:  Wichita  Mountains  National  Wild- 
life Refuge,  1. 

SOUTH  DAKOTA.— No  precise  locality,  1.  Cus- 
ter Co.:  Folsom,  1.  Harding  Co.:  20  mi.  NE  Buffalo, 
1.  Meade  Co.:  Faith,  1.  Pennington  Co.:  Imlay,  1. 
Ziebach  Co.:    Red  Elm,  1. 

WYOMING.— Converse  Co.:  Douglas,  2.  Na- 
trona Co.:    Natrona,  2. 

C.  I.  occidentalis 

ALBERTA. — Edmonton,  1;  Simonette  River,  1 
(UAlb);  30  mi.  N  Whitecourt,  3  (UAlb);  50  mi.  N 
Whitecourt,  3  (UAlb);  Wood  Buffalo  National  Park, 

4  (AMNH). 

BRITISH  COLUMBIA.— Barking  Horse  River,  2 
(KU);  upper  Henry  River,  1. 

NORTHWEST   TERRITORIES.— Artillery   Lake, 

5  (4  in  CNM);  Aylmer  Lake,  1  (AMNH);  Fort 
Good  Hope,  1;  Fort  Simpson,  1;  Fort  Smith,  1;  Great 
Bear  Lake,  1  (AMNH);  52  mi.  up  Keele  River,  1 
(CNM);  Nahanni  Butte,  10  (CNM);  mouth  of 
Netla  River,  1  (CNM);  Salt  Plains,  5  (CNM);  Slave 
River,  4  (CNM). 

YUKON. — 40  mi.  SE  Crow  Base,  3;  north  fork 
McMillan  River,  1;  Pelly  Lakes,  4;  White  River,  4 
(ROM). 

C.  I.  orion 

GREENLAND.— No  precise  locality,  2  (AMNH). 

C.  /.  pambasileus 

ALASKA. — No  precise  locality,  1.  Anaktuvak 
Pass,  7  mi.  N  Tolugak,  1;  Big  Delta  River,  1;  Cold 
Bay,  1;  Fairbanks,  1;  100  mi.  N  Fairbanks,  1;  Fare- 
well Mts.,  1;  Gold  Creek  (near  head,  above  Curry), 
1;  Jarvis  Creek,  1;  upper  John  River,  7;  Little  Delta 
River,  1;  Mt.  Hayes,  3;  Nome,  1;  Savage  River,  1; 
Sushana  River,  2;  Tanana  River,  2;  Teklanika  River, 
2;  Teller,  1;  Tolugak  Lake,  1;  Yukon  River,  35  mi. 
below  Beaver,  1. 

YUKON. — No  precise  locality,  1;  Hoole  Canyon, 
1. 

C.  I.  tundrarum 

ALASKA. — No  precise  locality,  1;  Noatak  River, 
2;  Pitmega  River,  Cape  Sabine,  1;  Point  Barrow,  1; 
Umiat,  2;  upper  Meade  River,  1  (UCMVZ);  Wahoo 
Lake,  Brooks  Range,  1   (KU). 

C.  I.  youngi 

CALIFORNIA. — San  Bernardino  Co.:  12  mi.  W 
Lanfair,  1  (UCMVZ). 

5.  Canis  latrans  from  northern  and  western  North 
America  (other  than  as  listed  in  part  2). 


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NO.  6 


C.  I.  incolatus 

ALASKA.— Big  Delta  River,  12;  Copper  River 
Flats,  1;  Eagle  River,  4;  Fairbanks,  2;  Mt.  Hayes,  4; 
Tanana,  1. 

C.  I.  latrans 

WYOMING.— Albany  Co.:  lelm,  1;  Laramie,  22; 
Red  Mts.,  1.  Carbon  Co.:  Shirley,  1.  Converse  Co.: 
Douglas,  4.  Crook  Co.:  Manville,  4;  Sundance,  4. 
Laramie  Co.:  Federal,  4.  Natrona  Co.:  Casper,  1. 
Sheridan  Co.:   Arvada,  1. 

C.  I.  mearnsi 

ARIZONA. — Apache  Co.:  Marsh  Lake,  1;  Spring- 
erville,  1.  Coconino  Co.:  Anderson  Mesa,  1;  Bright 
Angel  Spring,  1;  Flagstaff,  1;  Fredonia,  3;  Kaibab 
National  Forest,  1;  Ryan,  6;  Tuba,  4.  Graham  Co.: 
Chiricahua  Ranch,  1.  Mojave  Co.:  Trumbull  Mts., 
1.  Navajo  Co.:  Antelope  Springs,  1;  Ft.  Apache,  4. 
Yuma  Co.:  Gila  Mts.,  1;  Tinajas  Altas,  1;  Tule  Tanks, 
1. 

C.  I.  texensis 

NEW  MEXICO.— Bernalillo  Co.:  Isleta,  1.  Eddy 
Co.:  Salt  Valley,  2.  Lincoln  Co.:  Callo  Canyon,  3. 
Otero  Co.:  Cienega,  3;  Lincoln  National  Forest,  3. 
San  Juan  Co.:  Fruitland,  1.  San  Miguel  Co.:  Pecos, 
1.  Santa  Fe  Co.:  Lamy,  2.  Socorro  Co.:  Carthage, 
6;  San  Andres  Mts.,  1.  Torrance  Co.:  Manzano  Mts., 
1;  Mesa  Jiminez,  1. 

6.  Suspected  hybrids. 

Canis  lupus  x  Canis  latrans 

ARIZONA.— Not  located,  Lanks,  1. 

CHIHUAHUA.— Colonia  Garcia,  1  (MCZ). 

VERACRUZ.— Orizaba,  1  (MCZ). 

ONTARIO.— Captives,  2  (ROM).  Lanark  Co.: 
Sherbrooke,  1  ( ROM ) .  Nipissing  District:  Preston, 
1  (ROM). 

QUEBEC. — Gatineau  Co.:  northern  part,  1 
(QWS);  central  part,  1  (QWS);  Gracefield,  1 
(CNM,  originally  identified  as  C.  latrans  thamnos) , 
Papineau  Co.:  Montebello,  1  (ROM).  Pontiac  Co.: 
Head  Lake,  1  (CNM). 

Canis  lupus  x  Canis  familiaris 

MICHIGAN.— Luce  Co.:  McMillan,  1  (UMMZ). 
Schoolcraft  Co.:   Cusino,  1. 

NEW  MEXICO.— Otero  Co.:  Sacramento  Mts., 
2. 

7.  Canis  lupus  lycaon. 
Western  Group 

ONTARIO.— A/goma  District:  Batchwana  Bay, 
1  (UCMVZ);  McMahon  Tvvp.,  1  (ROM).  Cochrane 
Dist.:  Kapukasing,  1  (collection  of  Douglas  H.  Pim- 
lott).  Kenora  Dist.:  no  precise  locality,  1  (ROM); 
Ball  Lake,  1  (CNM);  Eagle  Lake,  1  (ROM);  100 
mi.  W  Fort  William,  1  (CNM);  Kenora,  1  (ROM); 
Whitefish  Bay,  2  (ROM).  Parry  Sound  Dist.:  Bur- 
ton Twp.,  1  (ROM);  Carling  Twp.,  1  (ROM). 
Rainy  River  Dist.:  Quetico,  3  (UI).  Thunder  Bay 
Dist.:    Hurkett,   1    (ROM);  Killala  Lake,  2   (ROM); 


Lake  Leopard,  1  (ROM);  Lape  Nipigon,  2  (CNM); 
north  shore  of  Lake  Superior,  1  (CNM);  Silver  Islet 

1  (ROM). 

MICHIGAN.— Alger  Co.:  southern  part,  1 
(UMMZ);  Grand  Marais,  1  (UMMZ);  14  mi.  SW 
Grand  Marais,  1  (UMMZ);  25  mi.  NE  Munising,  1 
(UMMZ).  Baraga  Co.:  Phcshika  River,  1  (UMMZ); 
Sec.  6,  T50N,  R31W,  1  (MSU).  Chippewa  Co.: 
north  shore  of  Whitefish  Bay,  1  (UMMZ);  Sec.  35, 
T47N,  R5W,  2  (MSU).  Delta  Co.:  West  Escanaba 
River,  1.  Dickinson  Co.:  no  precise  locality,  1; 
Randville,  1;  West  Escanaba  River,  1.  Gogebic  Co.: 
Iron  River,  1  (UMMZ);  Marinesco,  1  (UMMZ); 
Presque  Island,  2 (UMMZ);  7  mi.  N  Watersmeet,  1 
(MSU).  Houghton  Co.:  Kenton,  1.  Luce  Co.:  north 
of  Newberry,  1.  Marquette  Co.:  30  mi.  NW  Mar- 
quette, 2.  Ontonagon  Co.:  Calderwood,  1.  School- 
craft Co.:    1. 

MINNESOTA.— Beltrami  Co.:  Red  Lake  Na- 
tional Wildlife  Refuge,  1  (UMinn).  Cook  Co.:  no 
precise  locality,  3  (AMNH);  near  Dunn  Lake,  2 
(UMinn);   Horland,   1    (UMinn).    Koochiching  Co.: 

2  ( 1  in  TM,  1  in  UMinn).  Lake  Co.:  Clearwater 
Lake,  1;  Eskwagama  Lake,  1;  Hart  Lake,  1;  Horse 
River,  1;  South  Fowl  Lake,  1.  Lake  of  the  Woods 
Co.:  Baudette,  1  (UMinn);  12  mi.  S  Williams,  1 
(UMinn).  St.  Louis  Co.:  Duluth,  1  (AMNH);  Ely, 
6  (  UMinn ) ;  Four  Town  Lake,  1  ( UI ) .  Sherburne 
Co.:   Elk  River,  1. 

WISCONSIN.— Vilas  Co.:    Eagle  River,   1. 

Eastern  Group 

ONTARIO. — Nipissing  Dist.:  Algonquin  Pro- 
vincial Park,  7  (3  in  CNM,  3  in  ROM);  Bishop 
Twp.,  1  (ROM);  Clancy  Twp.,  1  (KU);  Lake  Ni- 
pissing, 1  (ROM);  Preston,  1  (ROM);  Whitney, 
1    (UCMVZ).    Peterborough   Co.:    north   of  Apsley, 

1  (CNM).    Renfrew  Co.:    Dacre,  1   (ROM). 

QUEBEC. — Southern  part,  no  precise  locality,  2 
(QWS).    Gatineau  Co.:    Aylwin,  1  ( CNM );  Lucerne, 

2  (UCMVZ).  Labelle  Co.:  Boyer,  1  (QWS);  La- 
coste,  1  (QWS);  Mont  Laurier,  1  (QWS);  Nomin- 
ingu,  1  (QWS);  Ste.  Veronique,  1  (QWS);  Val- 
Barrette,  1  (QWS).  Papineau  Co.:  Montebello,  2 
(ROM).  Pontiac  Co.:  near  Cabonga  Reservoir,  2 
(QWS);  Jim's  Lake,  1  (CNM).  Temiscamingue 
Co.:   40  mi.  NE  Mattawa,  1. 

8.  Canis  latrans  thamnos. 

MANITOBA.— Carman,  4;   Duck  Mountain,   1. 

ONTARIO.— Algoma  Dist.:  Dean  Lake,  1 
(ROM);  Prince,  1  (ROM);  Tarbutt,  4  (ROM); 
Wolford,  1  (ROM).  Greg  Co.:  Markdale,  1  (ROM). 
Huron  Dist.:  Zurich,  1  (ROM).  Kenora  Dist.:  Ox- 
drift,  1  (ROM).  Kent  Co.:  Chatham,  1  (ROM). 
Lambton  Co.:  Thedford,  1  (CNM).  Lanark  Co.: 
Sherbrooke,  1  (ROM).  Nipissing  Dist.:  Algonquin 
Provincial  Park,  2  (CNM).  Norfolk  Co.:  1  (ROM). 
Parry  Sound  Dist.:  Monteith,  1  (ROM).  Peter- 
borough Co.:  Lakefield,  1  (CNM).  Rainy  River 
Dist.:   Pinewood,  1  (ROM). 

QUEBEC— No  precise  locality,  2  (QWS). 
Bcauce    Co.:     Beauceville,     1     (QWS).     Charlevoix 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAWS 


141 


Co.:  Baie  St.  Paul,  1  (QWS).  U Islet  Co.:  St.  Au- 
bert,  1  (QWS).  Maskinonge  Co.:  St.  Leon,  1 
(QWS).    Portneuf  Co.:    Valcartier,  1   (QWS). 

ILLINOIS.— Lake  Co.:  Camp  Logan,  1  (FM). 
McClean  Co.:  LeRoy,  1.  Marshall  Co.:  9  mi.  W 
Henry,  1. 

INDIANA.— Clinton  Co.:  Jefferson,  1  (PUWL). 
Jasper  Co.:  McCoysburg,  1.  Newton  Co.:  5  mi.  S 
Roselawn,  1  (PUWL).  Tippecanoe  Co.:  West 
Point,  1  (PUWL). 

IOWA.— Adair  Co.:  Richland,  2  (KU).  Appa- 
noose Co.:  Moravia,  1  (KU).  Monroe  Co.:  2  mi. 
N  Avery,  2  (KU). 

MICHIGAN.— Alcona  Co.:  Aldair,  1  (UMMZ). 
Alger  Co.:  Miners  River,  1.  Baraga  Co.:  no  precise 
locality,  2  (MSU);  Baraga,  1  (UMMZ).  Barry  Co.: 
1  (UMMZ).  Cheboygan  Co.:  Beaugrand,  1 
(UMMZ).  Chippewa  Co.:  Brimley,  1  (UMMZ); 
7  mi.  NW  Pickford,  1  (UMMZ);  Race,  1  (UMMZ). 
Clinton  Co.:  St.  Johns,  1  (MSU).  Crawford  Co.: 
Hanson  Game  Refuge,  1  (UMMZ).  Delta  Co.:  Bark 
River,  1;  Rapid  River,  1.  Dickinson  Co.:  Cedar 
River,  1  (UMMZ).  Gogebic  Co.:  Ironwood,  2 
(UMMZ);  Montreal  River,  1  (UMMZ).  Houghton 
Co.:  Isle  Royale,  3  (UMMZ).  Ingham  Co.:  1 
(MSU).  Iron  Co.:  3  (MSU).  Jackson  Co.:  Liberty, 
1  (UMMZ).  Marquette  Co.:  Negaunee,  2;  Yalmar, 
1  (UMMZ).  Menominee  Co.:  Cedar  River,  2 
(UMMZ);  Dagett,  1;  Ingalls,  4  (UMMZ);  Michiga- 
mee  River,  1;  Whitney,  2  (UMMZ);  Wilson,  2 
(UMMZ).  Montcalm  Co.:  1  (MSU).  Ontonagon 
Co.:  no  precise  locality,  1  (UMMZ);  Ewen,  1 
(UMMZ).  St.  Clair  Co.:  1  (MSU).  Schoolcraft 
Co.:  no  precise  locality,  2  (MSU);  Manistique,  1 
(UMMZ).  Washtenaw  Co.:  Dexter,  1  (UMMZ). 
Not  located,  Warheim,  1   (UMMZ). 

MINNESOTA. — Beltrami  Co.:  no  precise  local- 
ity, 4  (UMinn);  Red  Lake  National  Wildlife  Refuge, 
1  (UMinn).  Isanti  Co.:  1  (UMinn).  Lake  Co.: 
Fernberg,  1  (UMinn).  Lake  of  the  Woods  Co.: 
no  precise  locality,  5  (UMinn);  Norris  Camp,  2 
(UMinn).  Pennington  Co.:  1  (UMinn).  Pine  Co.: 
1    (UMinn).     Sherburne  Co.:    Elk  River,   5. 

NORTH  DAKOTA.— Benson  Co.:  Ft.  Totten,  1; 
Sully  Hill  National  Park,  1. 

WISCONSIN.— Ashland  Co.:  Basswood  Island, 
Apostle  Islands,  1.  Forest  Co.:  Crandon,  4  (MSU); 
Wabeno,  1  (MSU).  Iron  Co.:  Kenosa,  1  (FM). 
Vilas  Co.:  Eagle  River,  1.  Walworth  Co.:  Delavan, 
1. 

9.  Wild  Canis  from  the  Northeastern  United 
States. 

MAINE. — Franklin  Co.:  Rangley,  1.  Kennebec 
Co.:   Monmouth,  1. 

MASSACHUSETTS.— Berkshire  Co.:  Otis,  1 
(MCZ).  Franklin  Co.:  Colrain,  1  (MCZ);  Leyden, 
1  (MCZ). 

NEW  HAMPSHIRE.— Coos  Co.:  Lancaster,  1 
(MCZ);  Stewartstown,  1  (MCZ).  Hillsborough  Co.: 
Temple,  2  (MCZ).  Merrimack  Co.:  Boscawen,  1 
(MCZ).    Sullivan  Co.:    Croydon,  1   (MCZ). 


NEW  YORK.— No  precise  locality,  1  (NYEC). 
Franklin  Co.:  Faust,  1  (NYEC);  Santa  Clara,  2 
(NYEC).  Lewis  Co.:  4  (NYEC).  Oneida  Co.: 
Hawkinsville,  2  (NYEC);  Woodgate,  1  (NYEC). 
Oswego  Co.:  Fulton,  1  (NYEC).  Schenectady  Co.: 
1  (NYEC).   Yates  Co.:   1. 

PENNSYLVANIA.— Clearfield  Co.:  Clearfield,  1. 
Potter  Co.:   1. 

VERMONT. — No  precise  locality,  1.  Addison 
Co.:  Granville,  1  (VFG).  Chittenden  Co.:  Shel- 
burne,  1  (VFG).  Orange  Co.:  Brookfield,  1  (VFG). 
Orleans  Co.:  Barton,  2  ( 1  in  VFG);  Glover,  1 
(VFG);  Jay,  1  (VFG);  Troy,  2  (VFG).  Rutland 
Co.:  no  precise  locality,  1  (VFG);  Middletown 
Springs,  1  (VFG);  Sudbury,  1  (VFG).  Washington 
Co.:  Berlin,  1  (VFG);  Montpelier,  1  (VFG).  Wind- 
ham Co.:  Brookline,  2  (MCZ);  Wardsboro,  1 
(MCZ). 

10.  Canis  rufus  gregoryi,  1919-1929  (for  data  on 
earlier  material  see  table  2). 

ARKANSAS.— Boone  Co.:  Bergman,  1.  Cle- 
burne Co.:  Almond,  1.  Dallas  Co.:  Carthage,  1. 
Garland  Co.:  Crystal  Springs,  1;  Lonsdale,  2  ( 1  in 
MCZ).  Marion  Co.:  Mull,  1.  Newton  Co.:  Falls- 
ville,  7;  Lurton,  2.  Perry  Co.:  Ava,  1;  Cedar,  1. 
Polk  Co.:  Egger,  1;  12  mi.  NE  Egger,  2;  10  mi.  W 
Egger,  1;  Mena,  1;  Shady,  1.  Pope  Co.:  Mill  Creek, 
2;  Simpson,  5;  Solo,  1.  Pulaski  Co.:  Femsdale,  5; 
Pinnacle,  2.  Saline  Co.:  Isaac,  6.  Scoff  Co.:  Blue 
Ball,  4;  Cardiff,  2;  4  mi.  S  Parks,  1.  Yell  Co.:  8  mi. 
NW  Aly,  1;  Onyx,  8;  Stillwater,  3. 

LOUISIANA. — Beauregard  Parish:  near  Sabine 
River,  2.   Madison  Pa.:    1. 

MISSOURI.— Carter  Co.:  Barren,  4.  Crawford 
Co.:  Cook  Station,  3.  Howell  Co.:  West  Plains,  2. 
Iron  Co.:  Arcadia,  2.  Ripley  Co.:  Gatewood,  3. 
Stone  Co.:  3.  Texas  Co.:  Tyrone,  1.  Wayne  Co.: 
Upalika,  1. 

OKLAHOMA. — Le  Flore  Co.:  Octavia,  1;  Page, 
3;  Talihina,  1.  McCurtain  Co.:  Bethel,  3;  Broken 
Bow,  7;  Sherwood,  4;  Smithville,  7.  Pushmataha 
Co.:    Cedar  Creek,  1;  Fewell,   1;  Nashoba,   1, 

11.  Southeastern  specimens  that  suggest  hybridi- 
zation with  Canis  familiaris. 

ARKANSAS.— Pope  Co.:    4  mi.   S   Raspberry,   1. 

LOUISIANA. — Northern  part,  no  precise  locality, 
1  (LPI).  Jackson  Pa.:  1  (LPI).  Winn  Pa.:  Sikes, 
1. 

MISSOURI.— Iron  Co.:    4  mi.  S  Sabula,  1. 

TEXAS.— Lavaca  Co.:  20  mi.  S  Hallettsville,  1. 
Van  Zandt  Co.:   1. 

12.  Specimens,  originally  identified  as  C.  rufus 
gregoryi,  with  short  greatest  lengths. 

ARKANSAS.— Marion  Co.:  Mull,  1.  Newton 
Co.:  Fallsville,  2.  Perry  Co.:  Ava,  1;  8  mi.  W  Wye, 
2.  Pope  Co.:  Simpson,  1.  Pulaski  Co.:  Fernsdale, 
1.   Sf.  Francis  Co.:   Forrest,  1. 

OKLAHOMA.— Le  Flore  Co.:  Octavia,  1;  Page, 
1.  McCurtain  Co.:  Bethel,  1;  Broken  Bow,  1;  Smith- 
ville, 2. 


142 


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NO.  6 


13.  Canis  latrans,  pre-1930,  southern  Missouri. — 
Carter  Co.:  Barren,  2.  Phelps  Co.:  Rolla,  5.  Saline 
Co.:  4  mi.  N  Napton,  1  (MCZ).  Texas  Co.:  Tyrone, 
1. 

14.  Pre-1930  specimens  originally  identified  as 
Canis  rufus  rufus. 

ARKANSAS.— Newton  Co.:    Boxley,  1. 

OKLAHOMA.— Atoka  Co.:  near  Atoka,  2.  Gar- 
vin Co.:  Cherokee  Town,  40  mi.  N  Ardmore,  1. 
Tulsa  Co.:    Red  Fork,  2. 

MISSOURI.— Sfone   Co.:     Reeds   Springs,   2. 

15.  Pre-1930  specimens  originally  identified  as 
Canis  latrans. 

OKLAHOMA.— Canadian  Co.:  Calumet,  5.  Co- 
manche Co.:  Cache,  3.  Creek  Co.:  Manford,  1. 
Custer  Co.:  Anthon,  1;  Butler,  5.  Tillman  Co.: 
Frederick,  9.    Tulsa  Co.:    Red  Fork,  1. 

TEXAS.— Hemphill  Co.:    2. 

16.  Canis  rufus  rufus,  1900  and  1904,  coastal 
Texas. — Calhoun  Co.:  O'Connorsport,  4;  7  mi.  SW 
Port  Lavaca,  2.  Colorado  Co.:  Frelsburg,  1.  Liberty 
Co.:  6  mi.  N  Dayton,  1. 

17.  Canis  latrans  texensis,  pre-1930,  southern 
Texas. — Frio  Co.:  Frio  Town,  1;  11  mi.  W  Frio 
Town,  1;  20  mi.  W  Frio  Town,  2;  8  mi.  SW  Frio 
Town,  4;  9  mi.  S  Moore,  1;  Pearsall,  7;  5  mi.  E 
Pearsall,  1;  20  mi.  W  Pearsall,  3.  Nueces  Co.:  Cor- 
pus Christi,  27;  45  mi.  SW  Corpus  Christi,  3;  Nueces 
Bay,  2;  San  Diego,  1.  Uvalde  Co.:  Sabinal,  1;  10 
mi.  N  Sabinal,  1;  5  mi.  S  Sabinal,  1.  Zavala  Co.: 
12  mi.  NE  Batesville,  1. 

18.  Canis  latrans  texensis,  pre-1930,  western 
Texas. — Brewster  Co.:  Alpine,  3.  Coke  Co.:  10  mi. 
N  Water  Valley,  1.  Crockett  Co.:  Ozona,  3;  9  mi. 
W  Ozona,  1;  12  mi.  NW  Ozona,  5.  Pecos  Co.:  Shef- 
field, 1.  Reagan  Co.:  Big  Lake,  4;  3  mi.  N  Big 
Lake,  1;  25  mi.  E  Big  Lake,  2;  12  mi.  S  Big  Lake, 
2.  Sterling  Co.:  Broome,  1;  Sterling  City,  3;  30  mi. 
S  Sterling  City,  1.  Upton  Co.:  Rankin,  7;  10  mi.  SW 
Rankin,  9.    Nolan  Co.:    Sweetwater,  1. 

19.  Canis  latrans  texensis,  pre-1930,  Tom  Green 
County,  Texas. — Carlsbad,  1;  6  mi.  NE  Carlsbad,  1; 
15  mi.  NE  Carlsbad,  2;  Christoval,  4;  6  mi.  NE 
Christoval,  1;  10  mi.  NE  Christoval,  1;  15  mi.  NE 
Christoval,  2;  20  mi.  NE  Christoval,  1;  Mereta,  1; 
San  Angelo,  25;  15  mi.  W  San  Angelo,  2;  Water 
Valley,  9. 

20.  Specimens  from  central  Texas,  pre-1930. — 
Blanco  Co.:  Blanco,  2;  Round  Mt.,  1.  Burnet  Co.: 
Burnet,  1;  5  mi.  E  Fairland,  1;  Marble  Falls,  6;  6 
mi.  S  Marble  Falls,  1.  Coleman  Co.:  16  mi.  N  Cole- 
man, 1.  Concho  Co.:  5  mi.  N  Pasche,  1.  Edwards 
Co.:  Nueces  River,  1.  Gillespie  Co.:  2.  Kerr  Co.: 
no  precise  locality,  4;  Kerrville,  1.  Llano  Co.:  no 
precise  locality,  4;  Baby  Head,  1;  22  mi.  S  Bird 
Range,  1;  Castell,  7;  Click,  2;  Llano,  7;  20  mi.  N 
Llano,    1;    15   mi.   E    Llano,    1;   20   mi.   S   Llano,   3; 


7  mi.  NW  Llano,  2;  Vallev  Springs,  2.  McCulloch 
Co.:  Brady,  3;  13  mi.  SW  Brady,  1;  5  mi.  SE  Doole, 
1.  Menard  Co.:  Callan,  1;  Ft.  McKavett,  1;  Menard, 
10.  San  Saba  Co.:  Cherokee,  2.  Sutton  Co.:  So- 
nora,  1;  25  mi.  W  Sonora,  2. 

21.  Canis  rufus,  1930's-1950's. 
C.  r.  gregonji 

ALABAMA. — Sumter  Co.:    Livingston,    1. 

ARKANSAS. — Union-Columbia   county  line,    1. 

LOUISIANA— La  Salle  Pa.:  Little  River,  1 
(LSUMZ).  Madison  Pa.:  Tallulah  Reservation,  2 
(LUSMZ).  Terrebonne  Pa.:  near  Houma,  1 
(LSUMZ).    Winn  Pa.:    3. 

MISSISSIPPI.— Harrison  Co.:  Biloxi,  1   (AMNH). 

OKLAHOMA.— McCurtain  Co.:  near  Battiest,  2 
(UArk). 

TEXAS. — Hardin  Co.:  no  precise  locality,  1; 
Honey  Island,  1  (UAriz);  Kountze,  1.  Newton  Co.: 
1.  Polk  Co.:  southern  part,  1;  Carmona,  1 
(UCMVZ);  near  Wakefield,  2. 

C.  r.  rufus 

TEXAS.— Brazoria  Co.:    12  mi.  S,  4  mi.  E  Alvin, 

I  (KU);  Angleton,  1;  9  mi.  NE  Angleton,  2;  5  mi. 
E  Angleton,  1;  12  mi.  E  Angleton,  1.  Brazos  Co.: 
15  mi.  S  Bryan,  1.  Harris  Co.:  Genoa,  1.  Liberty 
Co.:    Cleveland,  1;   1.5  mi.  N  Rye,  2.    Madison  Co.: 

II  mi.  SE  Madisonville,  2.  Montgomery  Co.:  Porter, 
2;  Security,  2.    Walker  Co.:    New  Waverly,  1. 

22.  Specimens   from   the   central   coast   of  Texas, 

1936-1942.— Aransas  Co.:  Aransas  National  Wildlife 
Refuge,  5.    Refugio  Co.:   22  mi.  E  Refugio,  1;  12  mi. 

5  Tivoli,  1;  7  mi.  S  Woodsboro,  1.  Victoria  Co.: 
Bloomington,  1;  6  mi.  S  Bloomington,  1. 

23.  Specimens  from  northern  Texas,  1930-1942. — 
Eastland  Co.:  Cisco,  1.  Jack  Co.:  Henry  Lewis 
Ranch,  2;  25  mi.  NW  Jacksboro,  1.  Palo  Pinto  Co.: 
no  precise  locality,  1;  6  mi.  NE  Graford,  1.  Parker 
Co.:  1.  Shackelford  Co.:  2.  Throckmorton  Co.:  5. 
Wilbarger  Co.:     16  mi.   SE   Vernon,    1.    Young  Co.: 

6  mi.  NE  Murray,  3  ( also  examined,  3  specimens 
from  near  San  Antonio,  Bexar  Co.). 

24.  Canis  latrans  frustror,  Wichita  Mountains  Na- 
tional Wildlife  Refuge,  Comanche  County,  south- 
western  Oklahoma,   1933-1942.--47. 

25.  Specimens  from  central  and  northeastern 
Oklahoma,  1932. — Cherokee  Co.:  1.  Cleveland  Co.: 
Noble,  3.  Osage  Co.:  2  (also  examined,  one  speci- 
men of  C.  r.  rufus  from  Redden,  Atoka  Co.). 

26.  Specimens  from  southern  Missouri,  1941-1942. 
— Christian  Co.:  1.  Crawford  Co.:  1.  Taney  Co.: 
3.  Texas  Co.:  1.  Vernon  Co.:  9  (also  examined, 
one  specimen  from  Dade  Co.,  collected  1932;  one 
from  3  mi.  N  Thomasville,  Oregon  Co.  (UCMVZ), 
collected  1942;  and  one  from  5  mi.  N  Gainesville, 
Ozark  Co.    (UCMVZ),  collected  1941). 

27.  Specimens  from  Arkansas,  1930-1951. — No 
precise   locality,    1    (UArk).     Benton   Co.:     Cherokee 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


143 


City,  3;  Siloam  Springs,  1;  Springtown,  1.  Howard 
Co.:  Umpire,  1,  Lawrence  Co.:  1.  Stone  Co.:  State 
Game  Refuge,  1  (UArk).  Washington  Co.:  Sum- 
mers, 2;  Devil's  Den  State  Park,  2   (UArk). 

28.  Specimens  from  southeastern  Oklahoma,  post- 
1960. — Bryan  Co.:  4.  Choctaw  Co.:  8.  McCurtain 
Co.:    6.    Pushmataha  Co.:    7. 

29.  Specimens  from  northern  Arkansas,  post- 1960 
(all  in  UArk).— Conway  Co.:  8.  Franklin  Co.:  2. 
Newton  Co.:    2.    Pope  Co.:    1.    Van  Buren  Co.:    7. 

30.  Specimens  from  southern  Arkansas,  post-1960. 
—Calhoun  Co.:  1  (UArk).  Chicot  Co.:  3.  Clark 
Co.:  1.  Hempstead  Co.:  26  (15  in  UArk).  Hot 
Springs  Co.:  4  (UArk).  Howard  Co.:  1  (UArk). 
Little  River  Co.:     14.    Miller  Co.:    6.    Nevada  Co.: 

4  (UArk).    Sevier  Co.:    3  (UArk). 

31.  Specimens  from  Louisiana,  post-1960. — Beau- 
regard Pa.:  near  Merryville,  8.  Bienville  Pa.:  2  ( 1 
in  LPI,  1  in  LSUMZ).  Bossier  Pa.:  1  (LPI).  Con- 
cordia Pa.:  Ferriday,  1.  De  Soto  Pa.:  3  (LPI). 
East  Carroll  Pa.:  4  mi.  N  Transvlvania,  1  (LPI). 
Jackson  Pa.:  7  (LPI).  Natchitoches  Pa.:  4  (LPI). 
Red  River  Pa.:  1.  St.  Landry  Pa.:  Thistlewaite 
Game  Management  Area,  3  (LSUMZ).  Union  Pa.: 
3  mi.  S  Farmerville,  1  (LPI);  2  mi.  N  Farmerville, 
1  (LPI).  Webster  Pa.:  1  (LPI).  Wed  Baton  Rouge 
Pa.:    2  mi.  W  Addis,  1   (LSUMZ).    Winn  Pa.:    2  mi. 

5  Brewster's  Mill,  5  (LPI);  4  mi.  E  Dodson,  1 
(LPI)  (also  examined,  one  specimen,  apparently  C. 
rufus  gregoryi,  from  near  Washington,  St.  Landry 
Pa.,  collection  of  Douglas  H.  Pimlott). 

32.  Specimens  from  inland  east  Texas,  post- 1960. 
—Bell  Co.:  2.  Bosque  Co.:  2.  Bowie  Co.:  3. 
Cherokee  Co.:  12.  Collin  Co.:  2.  Delta  Co.:  2. 
Denton  Co.:  16.  Freestone  Co.:  7.  Grayson  Co.: 
14.  Hamilton  Co.:  3.  Hopkins  Co.:  4.  Hunt  Co.: 
11.    Johnson  Co.:    8.    Lamar  Co.:    7.    Leon  Co.:    10. 


Limestone  Co.:  2.  Milam  Co.:  5.  Morris  Co.:  3. 
Rusk  Co.:    2.    Smith  Co.:    4. 

33.  Specimens   from   the   central   coast   of  Texas, 

post-1960.— Austin  Co.:  7  mi.  NW  Sealy,  2  (MSU). 
Calhoun  Co.:  7.  Colorado  Co.:  14  (MSU).  Fort 
Bend  Co.:  3.  Lavaca  Co.:  7.  Matagorda  Co.:  8. 
Victoria  Co.:  1  (USFWS)  (also  examined,  one  speci- 
men, apparently  C.  rufus  rufus,  from  near  Armstrong, 
Kenedy  Co.,   collection   of  Russell  E.   Mumford). 

34.  Specimens  from  the  vicinity  of  the  Addicks 
Reservoir,  Harris  Co.,  Texas,  post-1960. — 12  (7  in 
USFWS). 

35.  Specimens  from  the  vicinity  of  the  Clemens 
Prison  Farm,  western  Brazoria  Co.,  Texas,  post-1960. 
—31  (11  in  USFWS). 

36.  Specimens  from  the  eastern  part  of  Brazoria 
Co.,  Texas,  post-1960.— 15  mi.  S  Alvin,  1  (USFWS); 
5  mi.  E  Angleton,  5;  7  mi.  N  Angleton,  1  (USFWS); 
Graham  Ranch,  1;  near  Hoskins  Mound,  10  (7  in 
USFWS);  Liverpool,  1;  Stringfellow  Ranch,   1. 

37.  Specimens  from  the  vicinity  of  the  Big 
Thicket  southeastern  Texas,  post-1960. — Chambers 
Co.:  7  mi.  E  Baytown,  4  (USFWS).  Jasper  Co.: 
near  New  Blox,  1  (USFWS).  Liberty  Co.:  no  pre- 
cise locality,  3;  1  mi.  S  Ames,  1  (USFWS);  4  mi. 
S  Ames,  1  (USFWS);  5  mi.  S  Dayton,  2  (USFWS); 
2  mi.  E  Devers,  2  (USFWS);  5  mi.  N  Liberty,  2 
(USFWS);  15  mi.  E  Liberty,  2  (USFWS);  3  mi. 
S  Raywood,  1  (USFWS);  5  mi.  S  Raywood,  3 
(USFWS).    Tyler  Co.:    near  Fred,  2  (USFWS). 

38.  Canis  rufus  gregoryi,  southeastern  Texas, 
1963-1970.— Chambers  Co.:  Anahuac  National  Wild- 
life Refuge  and  vicinity,  6  (2  in  USFWS);  Barrows 
Ranch,  1  (UO);  Canada  Ranch,  2  (USFWS); 
Double  Bayou,  4;  Logan  Ranch,  2;  Monroe  City,  1; 
Smith  Point,  1  (USFWS).  Jefferson  Co.:  near  Port 
Arthur,  2. 


APPENDIX  B 


This  appendix  provides  measurements  for  some 
of  the  key  series  used  in  multivariate  analysis,  and 
for  some  of  the  fossil  specimens  examined.  The  num- 
bered parts  of  the  appendix  (left  margin)  are  the 
same  as  referred  to  in  the  text.  The  numbers  along 
the  tops  of  the  columns  correspond  to  the  numbers 
of  the  15  measurements  described  below.  If  no 
sample  size  (n)  is  indicated,  or  if  an  asterisk  (•) 
follows  the  sample  size,  then  the  figures  shown  are 
actual  measurements  of  individuals.  Otherwise,  the 
five  horizontal  rows  under  the  designation  and  sam- 
ple size  (n)  of  the  series  are  mean,  lower  extreme, 
upper  extreme,  standard  deviation,  and  coefficient 
of  variation.  Figures  in  parentheses,  following  the 
sample  size  of  most  series  for  which  sex  is  designated, 
represent  the  number  of  specimens  in  the  sample 
that  were  unknown  as  to  sex,  but  which  were 
judged  to  belong  to  the  category  indicated. 

Descriptions  of  Measurements 

1.  Greatest  length. — Length  from  anterior  tip  of 
premaxillae  to  posterior  point  of  inion. 

2.  Zygomatic  width. — Greatest  distance  across 
zygomata. 

3.  Braincase  width. — Maximum  breadth  of 
braincase  across  level  of  parietotemporal  sutures. 

4.  Alveolar  length  of  maxillary  toothrow. — Dis- 
tance from  anterior  edge  of  alveolus  of  PI  to  pos- 
terior edge  of  alveolus  of  M2. 


5.  Maximum  crown  width  across  upper  cheek 
teeth. — Greatest  breadth  between  outer  sides  of  most 
widely  separated  upper  teeth   (P4  or  Ml). 

6.  Palatal  width  at  PI. — Minimum  width  be- 
tween inner  margins  of  alveoli  of  first  upper  pre- 
molars. 

7.  Width  at  CI. — Greatest  breadth  across  max- 
illae at  outer  edges  of  alveoli  of  canines. 

8.  Width  of  frontal  shield. — Maximum  breadth 
across  postorbital  processes  of  frontals. 

9.  Postorbital  constriction. — Least  width  across 
frontals   at   constriction   behind   postorbital   processes. 

10.  Length  from  toothrow  to  bulla. — Minimum 
distance  from  posterior  edge  of  alveolus  of  M2  to  de- 
pression in  front  of  bulla  at  base  of  muscular  process. 

11.  Height  from  maxillary  toothrow  to  orbit. — 
Minimum  distance  from  outer  alveolar  margin  of  Ml 
to  most  ventral  point  of  orbit. 

12.  Depth  of  jugal. — Minimum  depth  of  jugal 
anterior  to  postorbital  process,  at  right  angle  to  its 
anteroposterior  axis. 

13.  Diameter  of  CI. — Maximum  anteroposterior 
width  of  upper  canine  at  base  of  enamel. 

14.  Crown  length  of  P4. — Maximum  anteropos- 
terior length  of  crown  measured  on  outer  side. 

15.  Crou;n  width  of  M2. — Maximum  transverse 
diameter  from  outermost  point  to  innermost  point  of 
crown. 


MEASUREMENTS 


1            2 

3 

4 

5 

6 

7 

8          9         10        11 

12        13        14        15 

C.  familiaris,  n: 

=50 

217.2     112.4 

58.95 

70.22 

68.12 

29.76 

41.64 

60.94  39.21  60.24  33.61 

15.25  11.21  19.28  10.91 

151.0       84.0 

50.5 

52.5 

51.5 

21.5 

30.0 

40.5     32.2     33.6     20.5 

10.1       8.4     14.4       7.7 

285.0     154.0 

65.0 

88.0 

85.5 

42.3 

59.0 

87.4     44.8     88.0     53.5 

23.6     14.0     22.7     13.0 

30.88     12.91       3.07       8.19       7.33       4.96       6.42       9.78     3.17  11.24     6.78     2.67     1.47     1.66     1.27 
14.27     11.48       5.21     11.66     10.76     16.67     15.42     16.09     8.08  18.66  20.17  17.51   13.11     8.61   11.64 


2.       Total  sample  of  northern  and  western  C.  lupus,  male,  n=233(33) 


259.6 

141.1 

65.92 

86.63 

82.20 

31.89 

48.27 

65.41  41.46 

66.46 

40.91 

19.56 

14.57 

25.92 

13.82 

235.0 

126.0 

58.8 

76.5 

72.2 

26.3 

40.2 

55.1     31.0 

57.0 

33.0 

14.5 

10.9 

22.2 

11.4 

293.0 

164.0 

71.8 

98.4 

94.0 

39.1 

55.0 

76.9     49.0 

78.5 

50.8 

24.1 

17.2 

30.5 

16.7 

12.27 

6.11 

2.46 

4.17 

3.76 

2.09 

2.73 

4.72     3.10 

4.56 

2.96 

1.53 

1.21 

1.42 

.99 

4.73 

4.33 

3.73 

4.81 

4.57 

6.55 

5.66 

7.22     7.45 

6.86 

7.24 

7.82 

8.30 

5.48 

7.16 

Total  sample  of  northern  and  western  C.  lupus, 

female 

n=146(33) 

247.7 

133.5 

64.89 

83.70 

78.30 

30.53 

45.73 

61.35  40.46 

62.68 

38.50 

18.23 

13.53 

24.79 

13.44 

224.0 

120.0 

59.2 

73.7 

70.2 

24.3 

39.3 

50.6     34.1 

54.0 

32.4 

14.8 

11.4 

22.2 

11.2 

278.0 

154.0 

71.3 

95.1 

90.3 

37.6 

53.9 

73.5     48.5 

75.8 

45.8 

23.4 

15.9 

28.2 

16.3 

12.18 

6.62 

2.69 

3.78 

3.60 

2.10 

2.59 

4.18     2.86 

4.26 

2.82 

1.35 

.97 

1.26 

.91 

4.92 

4.96 

4.14 

4.52 

4.56 

6.88 

5.66 

6.81     7.07 

6.80 

7.32 

7.40 

7.17 

5.08 

6.77 

Total  sample  of  northern  and  western  C 

'.  latrans,  male, 

n=166(2) 

197.1 

99.4 

57.79 

70.04 

56.53 

20.06 

30.91 

46.77  34.10 

46.96 

26.06 

12.15 

9.35  20.38 

11.81 

178.0 

88.0 

52.0 

61.4 

49.7 

17.5 

27.1 

36.1     29.1 

39.5 

21.2 

9.1 

7.7 

17.6 

9.7 

213.0 

109.0 

63.6 

78.4 

62.4 

29.7 

35.5 

54.6     39.7 

52.8 

29.6 

14.3 

11.0 

22.8 

13.8 

7.28 

3.96 

2.00 

2.85 

2.40 

1.14 

1.40 

3.13     2.13 

2.61 

1.63 

.88 

.62 

.96 

.64 

3.69 

3.98 

3.46 

4.07 

4.24 

5.68 

4.53 

6.69     6.25 

5.56 

6.25 

7.24 

6.63 

4.71 

5.42 

144 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


145 


1  234  5  6789 

Total  sample  of  northern  and  western  C.  latrans,  female,  n=rlll(4) 
188.1       95.0       56.80     67.39     54.19     19.47     29.44     44.81   33.74 
172.0       87.0       53.7       60.7       48.8       16.7       26.7       38.7     28.7 
204.0     106.0       60.2       74.0       60.4       22.5       32.4       53.6     40.9 
6.57       3.45       1.59       2.56       2.22       1.12       1.55       3.12     2.18 


10 


11 


12        13 


44.49  24.75 

39.0  21.9 

51.3  28.7 

2.36  1.46 


11.66 

9.7 

13.8 


8.78 
7.7 
10.0 
.56 


14 

19.60 

17.7 

21.6 


15 

11.52 
10.4 
13.0 
.55 


3.49       3.63       2.80       3.80       4.10       5.75       5.26       6.96     6.46     5.30     5.90     7.36     6.38     4.39     4.77 


C.  lupus  hjcaon,  western  group,  male,  n=42(ll) 


253.2 
238.0 
274.0 
7.35 
2.90 


136.7 
125.0 
150.0 
5.00 
3.66 


66.38 
59.8 
72.0 
2.99 
4.51 


84.39 
77.5 
90.3 
2.71 
3.21 


79.42 
73.0 
85.5 
2.80 
3.52 


30.31 
26.7 
34.3 
1.93 
6.38 


45.75 
41.6 
50.6 
2.10 
4.59 


C.  lupus  hjcaon,  western  group,  female,  n=30(8) 


241.2 
224.0 
268.0 
10.03 
4.16 


129.0 
121.0 
142.0 
5.36 
4.15 


64.34 

59.0 

69.3 

2.67 
4.16 


81.07 
72.4 
87.5 
3.34 
4.12 


75.14 
70.5 
81.0 
2.86 
3.81 


28.81 
26.0 
32.5 
1.64 
5.71 


C.  lupus  hjcaon,  eastern  group,  male,  n=19(4) 

947  1        1 .14  1  fift  95       R9  fid       77  85       98  TO 


247.1 

237.0 
255.0 
5.96 
2.41 


134.1 
128.0 
140.0 
3.59 
2.68 


63.25 
58.3 
68.0 
2.49 
3.93 


82.69 
78.5 
87.7 
2.48 
2.99 


77.85 
74.2 
84.3 
2.71 
3.48 


28.59 
26.0 
32.0 
1.47 

5.14 


42.47 
39.5 
47.3 
1.79 
4.22 


44.26 
40.8 
47.6 
1.69 

3.81 


C.  lupus  hjcaon,  eastern  group,  female,  n=12(2) 


231.4 
223.0 
241.0 
6.64 
2.87 


125.0 
116.0 
132.0 
4.79 

3.82 


62.68 
60.5 
66.0 
1.70 
2.71 


79.15 
75.0 
83.5 
2.92 
3.69 


73.76 
69.0 
78.3 
3.20 
4.34 


26.39 
23.5 
30.0 
2.17 
8.21 


40.76 
37.6 
44.3 
2.08 
5.11 


235.0 
250.0 
235.0 
228.0 
246.0 
240.0 
234.0 
224.0 
250.0 
247.0 
240.0 


121.0 
128.0 
126.0 
115.0 
120.0 
122.0 
114.0 
122.0 
126.0 
124.0 
120.0 


64.1 
65.4 
60.6 
64.1 
61.1 
64.1 
61.1 
60.9 
62.6 
65.7 
66.4 


81.0 
78.5 
79.5 
82.5 
78.9 
80.4 
77.5 
82.8 
82.6 
80.2 


71.1 
71.0 
66.0 
70.0 
72.5 
70.7 
70.1 
71.2 
71.0 
69.4 


25.1 
26.6 
27.2 
24.0 
28.5 
26.7 
27.4 
28.1 
25.8 
26.1 
26.7 


38.3 
43.8 
42.3 
39.0 
42.5 
40.4 
39.9 
40.5 
39.4 
40.2 
40.2 


64.52 
54.1 
76.5 
5.61 
8.69 


60.43 
49.4 
89.7 
4.81 
7.96 


60.99 
49.4 
72.8 
5.52 
9.07 


56.58 
51.9 
60.6 
2.86 
5.05 


41.56 
35.3 
46.3 
2.57 
6.18 


40.14 

34.9 

46.1 

2.76 
6.88 


65.70  38.65 

60.0  34.0 

76.0  45.0 

3.25  2.35 

4.95  6.07 


61.21 
51.7 
70.0 
4.26 
6.96 


36.04 
32.0 
40.9 
2.20 
6.10 


18.72 
16.3 
22.8 
1.27 
6.80 


17.24 
15.6 
20.8 
1.20 
6.98 


36.93 
35.0 
42.5 
2.21 
5.99 


59.32  3.5.10 

54.6  32.3 

64.4  37.7 

3.30  1.96 

5.57  5.58 


4.       Early  specimens  of  C.  rufus,  as  listed  in  table  2,  male,  n 

84.0       70.0 


11(4)° 
56.7     41.3 


61.9 
57.4 
58.9 
53.0 
52.4 
49.8 
57.6 
58.2 
56.5 
52.5 


43.3 
37.7 
41.3 
35.4 
35.7 
38.5 
38.3 
32.3 
37.0 
38.3 


Early  specimens  of  C.  rufus,  as  listed  in  table  2,  female,  n=3(  1)° 
222.0     114.0       60.5       75.0       66.0       24.4       37.3       48.4     31.5 
230.0     117.0       60.7       79.5       65.0       25.5       38.8       54.0     38.3 
222.0     115.0       64.1       77.3       64.6       25.5       37.0       52.3     37.3 


59.9 
63.8 
57.5 
57.5 
60.3 
62.7 
61.7 
56.3 
60.6 
62.7 
62.0 


55.7 
58.4 
54.5 


5.       C.  rufus  gregoryi,  south-central  United  States,  1919-1929,  male,  n=63(  1) 


232.6 

121.2 

61.93 

78.98 

69.37 

26.21 

40.00 

218.0 

110.0 

58.3 

72.6 

63.6 

22.3 

35.7 

261.0 

138.0 

68.0 

86.8 

75.3 

32.0 

47.2 

8.76 

5.93 

2.10 

2.79 

2.73 

1.95 

2.24 

3.77 

4.89 

3.39 

3.53 

3.93 

7.45 

5.61 

55.52  37.33  58.32 

47.2  32.0  51.8 

62.1  42.5  65.4 

3.73     2.40  3.45 

6.71     6.43  5.91 


34.5 
37.6 
35.3 
34.3 
35.2 
34.2 
34.0 
33.2 
32.2 
35.0 
32.5 


33.6 
32.3 
31.3 


33.64 
29.1 
38.0 
2.24 
6.65 


16.3 
18.3 
18.4 
18.3 
15.8 
15.8 
16.8 
16.0 
15.4 
16.2 
17.5 


15.2 
13.7 
14.5 


15.78 
13.3 
18.5 
1.26 
7.96 


13.61 

11.8 

15.5 

.99 
7.30 


25.19 
23.7 
27.4 
.93 
3.70 


39.78  63.34  37.78  18.15 

36.0  59.0  34.3     16.3 

44.9  69.5  42.2     20.0 

2.77     2.86  1.89     1.08 

6.95     4.51  5.00     5.93 


12.46  23.56 
10.4     21.1 
13.9     26.0 
.89     1.27 

7.18     5.38 


13.09  24.55 
12.0     22.6 
14.8     27.5 
.80     1.20 

6.12     4.88 


16.54 
14.0 
18.5 
1.49 
9.01 


11.98 
11.3 

12.7 

.50 
4.21 


22.67 

21.3 

24.2 

.93 
4.10 


11.1 
12.6 
13.7 
11.2 
13.6 
12.2 
12.4 
11.4 
11.1 
11.2 
13.5 


24.5 
25.2 
23.8 
23.5 
24.8 
23.6 
23.8 
24.6 
23.8 
24.8 
24.5 


14.02 
12.8 

15.7 

.67 
4.79 


13.39 
11.0 
14.8 
.83 
6.18 


14.25 
13.4 
15.7 
.66 
4.64 


13.64 

13.2 

14.3 

.35 
2.61 


13.1 
14.2 
13.5 
14.4 
13.9 
13.7 
14.4 
13.8 
14.6 
15.6 
13.1 


10.5  22.1  12.0 
10.1  22.3  13.0 
11.3     22.2     13.4 


11.93  23.66 
10.5     21.4 
13.2     26.0 
.73     1.00 
6.10     4.23 


13.68 

10.6 

16.0 

.82 
5.99 


146  MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY  NO.  6 

1  2            3            4            5            6            7            8          9         10        11  12  13  14  15 
C.  rufus  grcgoryi,  south-central  United  States,  1919-1929,  female,  n=52(l) 

220.9  115.4  61.14     75.15     66.78     25.32     37.99     52.93  37.93  53.89  31.66  14.84  11.12  22.31  1.3.29 

210.0  108.0  57.6       68.5       61.6       21.2       33.4       42.7     30.8     50.4     27.3  12.0  9.6  20.0  11.7 

245.0  130.0  64.8       80.5       74.7       29.7       45.0       63.0     41.5     66.1     36.1  17.3  12.9  24.4  14.7 

5.58  4.40       1.89       2.58       2.85       1.98       2.23       4.20     2.44     2.66     1.72  1.02  .71  1.08  .72 

2.53  3.81       3.11       3.45       4.27       7.62       5.87       7.94     6.42     4.93     5.33  6.88  6.36  4.84  5.42 


6.       Specimens  from  inland  eastern  Texas,  post- 1960,  male,  n=77(6) 

rtr>r>  o         l  A  A   n  p^o  m         ^l    r\r\         Pfl  r»n         m    C/D         oo  on         AC\  ere"     Off 


206.8  104.2  58.61  71.90  59.22  21.56  33.32  49.55  35.57  50.74  27.91   13.15  10.05  21.04  12.29 

192.0  96.0  54.6  65.5  53.3  18.8  29.5  43.5     29.6     44.0     24.3     10.8  8.8     19.2  10.3 

221.0  112.0  63.7  76.8  64.8  24.6  36.3  61.5     40.2     56.8     30.5     15.1  11.5     23.0  13.6 

5.62  3.45       2.11  2.14  1.91  1.25  1.42       3.59     2.12     2.64     1.44       .97  .56       .85  .64 

2.72  3.31       3.60  2.98  3.22  5.80  4.26       7.24     5.96     5.21     5.17     7.35  5.56     4.02  5.23 

Specimens  from  inland  eastern  Texas,  post-1960,  female,  n=42(6) 

198.0  99.52  58.01  69.81  57.39  20.59  31.75  47.36  35.66  47.86  26.60  12.41  9.51  20.43  12.31 

180.0  91.0  54.2  64.7  53.6  18.6  29.1  42.1     30.1     42.2     22.3     10.7  8.4     18.5  10.5 

214.0  109.0  62.0  75.5  62.8  23.1  37.8  56.0     39.8     54.8     30.8     15.5  11.5     23.0  13.6 

7.51  3.78       1.61  2.77  1.94  1.08  1.65       3.23     2.01     3.20     1.77     1.01  .64       .96  .72 

3.80  3.79       2.77  3.97  3.39  5.24  5.19       6.83     5.63     6.69     6.65     8.15  6.70     4.72  5.85 

Specimens  from  Jefferson  and  eastern  Chambers  counties,  Texas,   1963-1970,  male,  n=15(3) 

233.7  119.1  60.63  76.73  68.05  26.59  39.59  50.99  35.79  60.23  33.31   15.07  11.31  22.27  13.86 

218.0  105.0  56.0  67.0  61.5  24.0  33.8  41.3     27.5     52.5     28.3     11.8  10.0     21.1  12.0 

247.0  130.0  64.7  82.6  73.6  29.2  43.0  58.0     40.0     65.0     36.8     16.7  12.5     23.3  14.8 

6.98  5.95       2.39  3.76  3.06  1.71  2.31       4.14     3.44     3.23     2.30     1.23  .64       .79  .80 

2.99  4.99       3.95  4.90  4.50  6.43  5.83       8.12     9.60     5.36     6.90     8.15  5.63     3.55  5.79 

Specimens  from  Jefferson  and  eastern  Chambers  counties,  Texas,  1963-1970,  female,  n=4° 

222.0  112.0  58.7  75.0  65.5  27.7  38.5  47.7     33.5     55.0     33.6     15.5  11.0     21.1  12.3 

224.0  111.0  61.8  77.1  63.0  24.0  35.8  52.5     39.1     56.7     33.7     14.7  10.6     21.1  13.2 

220.0  111.0  58.8  76.1  66.8  25.4  35.4  48.5     35.7     58.0     30.8     14.0  10.1     21.1  13.9 

225.0  110.0  59.6  74.0  63.8  24.1  38.3  45.8     33.8     55.5     31.6     15.3  10.8     21.2  12.5 


8.  C.  cedazoensis,  Cedazo,  Aguascalientes 

9.  C.  lepophagus,  Hagerman,  Idaho 

C.  lepophagus,  Rexroad  fauna,  Kansas 

C.  lepophagus,  Broadwater,  Nebraska 
C.  lepophagus,  Lisco,  Nebraska 


16.9 


17.0     10.3 
16.0     10.0 


19.1       .... 

16.9       _.. 

—      10.2 


19.1     11.5 
19.6     12.3 


17.9     11.0 


C.  lepophagus,  Cita  Canyon,  Texas 

194.0     103.0         ....        68.6         ...  20.5         ...  54.0       ....  11.8  ....        _ 

187.0         ....        54.3       64.7         _._  _..  ....        ....  26.6  ....  8.7  19.0     11.3 

- -          --  _  57.2  35.0     51.0  ....  ....  ....  _..        _ 

190.0       99.0       50.0       70.0         _..  20.0         „  48.0  33.0       _  25.0  12.0  20.0     12.0 

74.5         __  ....          _..  ....  ....        _..  ....  ....  8.8  20.7     12.0 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


147 


10. 


12  3  4  5  6 

C.  latrans,  Papago  Springs  Cave,  Arizona 
174.0       99.0       56.9       67.1       57.1       20.2 

C.  latrans,  McKittrick,  California 


10 


11 


12 


13        14 


15 


30.0       40.6     32.9     43.1     25.4     12.6 


19.6     11.3 


'98.6 

58.3 

74.5 

59.5 

47.0 

38.2      _ 

._      25.2 

11.5       _.      22.0 

12.6 

62.4 

76.5 

65.0 

24.5 



_      27.6 

....      22.3 

12.3 

210.0 

77.0 

64.4 

22.5 

35.5 

55.0 

36.0       . 

_      28.0 

13.1       ....      22.5 

12.6 

219.0 

110.0 

58.4 

77.5 

64.5 

21.5 

53.1 

35.5 

...      28.8 

14.0       ....      24.0 

13.8 

207.0 

72.5 

61.1 

23.0 

33.3 







....      10.3     21.3 

12.3 

206.0 

61.5 

75.5 

61.8 

20.0 

33.5 

56.0 

36.5      - 

._ 

....      22.5 

— 

C.  latrans,  Maricopa,  California 
204.0        —       59.0      73.5      56.2 

C.  latrans,  Rancho  La  Brea,  California 


21.8       32.8 


20.1 


n=44 

n=36 

n=42 

n=49 

n=21 

n=44 

n=38 

n=44 

205.5 

106.7 

60.54 

72.54 

61.20 

22.18 

34.29 

51.21 

185.0 

90.0 

55.7 

65.5 

50.0 

17.3 

29.8 

44.0 

222.0 

116.0 

65.1 

78.0 

67.4 

25.5 

38.3 

61.5 

9.03 

5.58 

1.76 

3.23 

4.31 

1.70 

2.00 

4.14 

4.39 

5.23 

2.91 

4.45 

7.05 

7.68 

5.83 

8.0S 

;47  n=40  n=40  n=36   n=8 

1  36.73  48.16  27.05  13.07     9.88 

32.9     44.0     22.0     10.2       9.0 

42.0     52.8     32.3     15.2     10.9 

2.12     2.55     2.33     1.10       .53 

5.78     5.30     8.63     8.41     5.32 


C.  latrans,  Rancho  La  Brea,  California  (unusually  small  specimen) 

179.0       90.0       56.5       64.0       50.8       19.6       28.7       41.1     34.9     41.8     22.5 


10.2 


Type  of  C.  andersoni 

173.0       93.0       57.5       63.5       56.0 

11.  C.  edwardii,  Curtis  Ranch,  Arizona 

__      118.0         __        77.0       65.4       22.2 

C.  edwardii,  Rome  Beds,  Oregon 

59.0      78.8      68.0      19.5 

12.  C.  rufus,  Eddy  Bluff  shelter,  Arkansas 


C.  rufus,  Haile  VIIA,  Florida 
78.0 


35.1       38.3     33.3     40.0     24.5     12.2       .._ 

....      .  _      29.1     13.7     12.4 

50.8     38.8       ....        _...        __      11.3 


14.5 


C.  rufus,  Inglis  IA,  Florida 

C.  rufus,  Port  Kennedy,  Pennsylvania 

13.       C.  armbrusteri,  McCleod,  Florida 

C.  armbrusteri,  Coleman  IIA,  Florida 

__      133.0       66.4       85.6       78.2       24.0 


63.0     45.2 


35.0     18.5 


n=43 

n=33 

21.05 

11.84 

18.2 

10.2 

23.5 

13.6 

1.29 

.89 

6.14 

7.54 

18.0 

10.2 

20.5 

_._ 

24.0 

12.6 

24.0 

13.7 

23.0 

13.1 

20.9 

13.3 

22.1 

22.8 

— 

24.0 

12.3 

26.4 

14.9 

26.0 

15.5 

26.1 

13.9 

28.8 

26.5 

148 


MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 


NO.  6 


14. 


12            3            4            5            6  7 
C.  armbrusteri,  Cumberland  Cave,  Maryland 

258.0     128.0         ....        96.0       82.2       27.0  43.5 
64.8         ....        78.1 

....      150.0         ....        95.0       87.0       29.0  45.8 

285.0     161.0       75.0       98.0       80.0       30.7  49.0 


10 


11        12        13        14 


15 


270.0 


94.0 


65.0 


83.5 


61.5     50.0  61.7     34.5  17.2 

59.8     44.5  51.0       .... 

68.0     44.0       

70.0     43.0  77.0     45.0  21.0 

....  65.0     35.0  .... 

59.0     39.8  60.0     35.6  19.0 


C.  armbrusteri,  Rushville,  Nebraska 

C.  lupus,  Hunker  Creek,  Yukon 
257.0         ....        61.0       88.7 

C.  lupus,  Maricopa,  California 
258.0     144.0       69.0       87.5 
255.0 


....      135.0 
C.  lupus,  Rancho  La  Brea,  California 


35.8       52.0       57.5     40.0     66.3     42.7     20.0 


33.8 
32.5 


48.2  '    64.0     39.0 


41.6     18.8 


28.0 

26.6 

14.1 

10.5 

12.8 

28.7 

1577 

27.9 

15.5 

28.9 

15.2 

28.0 

17.0 

29.5 

17.1 

28.9 

15.2 

— 

30.0 

15.5 

— 

28.5 

13.7 

26.6 

13.7 

26.6 

12.5 

26.2 

28.5 

23.3 

13.1 

28.1 

12.2 

240.0 

138.0 

62.7 

81.5 

79.7 

33.6 

48.1 

74.3 

46.0 

62.7 

38.0 

25.0       . 

-      26.8 

14.1 



140.0 



82.7 

80.3 

28.5 

44.0 

64.0 

44.0 

39.0 

18.3       . 

...     24.8 

14.2 

123.0 

66.9 

78.7 

72.3 

29.0 

58.9 

42.2 

56.0 

35.6 

16.0 

_      23.0 

12.5 

230.0 

128.0 

66.0 

77.8 

78.8 

29.0 

45.0 

53.0 

42.7 

56.3 

37.0 

19.0 

...      26.8 

13.1 

265.0 

150.0 

71.4 

90.0 

85.7 

33.3 

49.4 

68.0 

48.2 

66.7 

39.5 

21.0       . 

_      29.0 

14.3 

269.0 

147.0 

69.2 

89.9 

36.1 

52.0 

75.0 

41.3 

70.5 

44.5 

22.1       . 

...      26.8 

13.2 

252.0 



62.7 

82.5 

79.9 

33.5 

44.6 

63.8 

39.5 

19.7       . 

...      25.1 

13.8 



148.0 

69.0 

94.0 

94.5 

38.0 

68.3 

43.7 

70.6 

44.3 

23.5       . 

...      29.2 

12.8 

248.0 

140.0 

64.5 

84.0 

83.2 

31.0 

46.8 

57.5 

41.5 

62.0 





_      27.6 

13.9 

.... 

— 

— 

.... 

.... 

— 

— 

— 

— 

.... 

— 

— 

_      23.5 
_      25.5 

12.5 

Type  of  C.  milleri 

247.0 

137.0 

66.7 

83.0 

89.6 

33.5 

49.8 

63.9 

44.3 

58.6 

39.0 

18.8       _ 

_      28.6 

13.2 

C.  lupus,  Goodland,  Kansas 
223.1     126.0       67.0       76.5 

C.  lupus,  Hay  Springs,  Nebraska 


73.0       30.0       43.6       59.0     34.0     54.5     34.0     15.0     13.3     23.1     14.0 


C.  lupus,  Hermit's  Cave,  New  Mexico 

C.  lupus,  San  Josecito  Cave,  Nuevo  Leon 
216.0     125.0       63.0       76.5       75.0       26.5 

15.       C.  dims,  McKittrick,  California 

309.0     170.0       72.5     106.5       99.0       40.0 


_        _..        ....        ....        __  _  27.4  13.6 

....  14.9  27.9  14.6 

40.0       54.0     40.5     50.5     32.4     17.7  11.9  25.0  12.4 

58.0       78.5     47.8     75.5     49.4     24.0  15.8  33.0  15.6 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CAMS 


149 


12  3  4 

C.  dims,  Maricopa,  California 


311.0 
298.0 
300.0 

310.0 
318.0 
290.0 


174.0       76.0 


160.0 
168.0 

158.0 


74.5 

80.0 

73.0 


106.1 
100.0 
102.5 
100.0 
101.0 
105.0 
101.0 


96.7 


95.2 
97.0 


44.3 
38.0 
39.0 
40.0 
40.0 
39.0 
38.0 


C.  dims.  Rancho  La  Brea,  California,  n=62 


294.8 
258.0 
316.0 
11.31 
3.84 


163.3 
148.0 
177.0 
7.15 

4.38 


74.73 
64.0 
83.0 
3.08 
4.12 


99.99 
85.0 
107.0 
4.18 
4.17 


96.15 
87.7 
104.0 
3.92 
4.08 


39.27 
35.0 
45.3 
2.36 
6.01 


69.0 
59.0 
60.0 

65.0 
58.0 


58.02 
52.0 
65.5 
3.30 
5.68 


10 


90.8     52.5     77.5 
93.0       .... 


82.5 
95.0 
87.0 
89.0 


47.9 
54.0 

47.0 


75.0 


11 

47.0 

45.2 
43.8 
43.0 

45.0 


12 

27.0 

24.5 
22.5 
21.5 
22.0 
21.5 
22.8 


13 


14 


17.2     32.3 


31.8 


83.45 
73.4 
100.0 
5.63 
6.75 


49,33 
43.5 
54.4 
2.13 
4.32 


72.43 
63.5 
77.5 
3.54 
4.89 


42.39 
36.6 
48.5 
2.66 
6.38 


C.  dims,  Hornsby  Springs,  Florida 


C.  dims,  Reddick  IA,  Florida 


C.  dims,  Melbourne,  Florida 


C.  dims,  Bradenton,  Florida 

C.  dims,  Ohio  River,  Indiana 

C.  dims,  Twelve  Mile  Creek,  Kansas 

C.  dims,  Welsh  Cave,  Kentucky 

309.0     180.0       78.0     104.3     100.7       39.0       59.1     104.0     57.3     82.0     45.5     25.0 

C.  dims,  Herculaneum,  Missouri 


C.  dims,  Hermit's  Cave,  New  Mexico 


40.4     20.7 


C.  dims,  Ingleside,  Texas 
333.0     179.0       79.0     110.0 


44.0 


53.0       ....      48.8     27.0 


32.5 
31.0 


21.75 
18.8 
26.5 
1.66 

7.63 


C.  dims,  Marlow,  Oklahoma 

310.0     170.0       78.5     111.5     102.5       37.0       60.0     100.0     50.0     79.0     49.0     24.0     18.0 


C.  dims,  San  Josecito  Cave,  Nuevo  Leon 

—      161.0       72.0     102.0     101.2       37.6  _      49  2  45  5     21 4 

297.0     169.0       76.0     104.3     103.0       37.2       58.8       94.0     54.2     74.0     44.9     21.0     14.! 


15 


14.9 
16.0 
15.4 
14.4 


15.66 
13.5 
17.5 
1.15 

7.34 


31.75 
28.7 
35.3 
1.38 
4.35 


15.15 
13.1 
17.0 
.90 

5.94 


29.6     14.5 
32.2       _._ 


30.7  16.5 

30.8  15.5 


32.0 
30.0 


30.1  15.1 

35.5  14.0 

30.0  15.4 

30.5  15.7 

32.5  15.6 

33.0  15.4 

31.0  .._ 

35.5  17.5 


33.6  15.4 

33.7  15.0 


APPENDIX  C 


The  numbers  along  the  top  of  each  of  the  following  columns  correspond  to  four  measurements  of  the  man- 
dible and  lower  dentition:  (1)  distance  from  anterior  edge  of  alveolus  of  pi  to  posterior  edge  of  alveolus  of 
m3;  (2)  minimum  depth  from  dorsal  surface  of  mandible  between  p3  and  p4  to  ventral  surface  of  mandible; 
(3)  crown  length  of  p4;  (4)  crown  length  of  ml.  If  a  sample  size  (n)  is  listed  for  a  series,  then  the  five 
horizontal  rows  under  that  figure  are  mean,  lower  extreme,  upper  extreme,  standard  deviation,  and  coefficient 
of  variation.  If  no  sample  size  is  given,  the  numbers  shown  are  actual  measurements  of  individuals,  rather 
than  means,  etc. 


1.    C.  lepoplmgus 

Santa  Fe  River,  Florida 


Grand  View,  Idaho 
Hagerman,  Idaho 


Rexroad  fauna,  Kansas 

Broadwater,    Nebraska 
Lisco,  Nebraska  


Cita  Canyon,  Texas 


2.    C.  latrans 

Recent,  western  U.S.,  male 


Recent,  western  U.S.,  female 


Irvington,   California 
McKittrick,   California 


4 


78.0 

15.9 

12.9 

22.5 

83.5 

19.4 

12.3 

22.6 

.... 

20.0 

82.5 

19.1 

20.8 

— 

18.7 

13.5 

22.4 

19.0 

17.1 

13.2 



70.5 

17.5 

12.3 

20.8 

70.0 

14.0 

11.5 

18.5 

78.7 

16.0 





— 

— 

.... 

18.1 

75.5 

— 

20.5 

70.0 

14.5 

11.1 

17.9 

n=14 

n=14 

n=16 

n=13 

78.61 

18.16 

13.08 

21.21 

73.2 

15.1 

12.1 

19.5 

82.0 

21.3 

14.0 

23.0 

2.63 

1.48 

.52 

1.15 

3.35 

8.14 

3.94 

5.42 

n=99 

n=99 

n=99 

n=99 

79.78 

17.10 

12.58 

21.94 

71.4 

14.3 

10.7 

19.5 

88.5 

19.9 

14.0 

24.3 

3.26 

1.14 

.70 

1.01 

4.10 

6.66 

5.58 

4.60 

n=99 

n=99 

n=99 

n=99 

76.35 

15.98 

12.08 

21.10 

69.1 

12.7 

10.8 

18.6 

82.3 

19.0 

13.9 

23.4 

2.71 

1.14 

.60 

.90 

3.55 

7.12 

4.96 

4.26 

79.9 

19.3 

13.8 

23.9 

78.9 

16.9 

13.4 

21.7 

n=10 

n=10 

n=14 

n=16 

82.24 

18.94 

12.78 

22.78 

77.0 

17.4 

11.7 

21.0 

88.5 

20.4 

14.1 

24.6 

3.82 

1.14 

.70 

1.11 

4.64 

6.02 

5.51 

4.88 

150 


1979 


NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS 


151 


l 

Maricopa,   California   n=10 

82.80 
80.0 
85.0 
1.81 
2.19 

Rancho  La  Brea,  California  n=41 

81.77 
77.0 
87.5 
2.52 
3.08 

Vallecito  Creek,  California 85.5 

Haile  XIIB,  Florida  __ 78.1 

Devil's  Den,  Florida _..        72.1 

Lake  Cutaline,   Florida  

Rushville,  Nebraska  _ 

Mullen,  Nebraska  73.6 

Frankstown   Cave,   Pennsylvania   83.5 

Lewisville,  Texas  _. 82.0 

Friesenhahn  Cave,  Texas 78.6 

75.5 

76.5 

San  Josecito  Cave,  Nuevo  Leon  : n  =  12 

80.25 

76.0 

84.0 

2.61 
3.25 


C.  edwardii 

Anita,   Arizona   

Curtis  Ranch,  Arizona  .. 

Minaca  Mesa,  Chihuahua 


n=10 

n=15 

n=22 

18.15 

12.75 

23.13 

17.0 

11.6 

21.1 

19.8 

13.8 

25.0 

1.00 

.55 

.84 

5.55 

4.33 

3.62 

n=41 

n=37 

n=40 

18.69 

12.56 

22.44 

16.0 

11.3 

21.0 

21.1 

13.7 

25.2 

1.30 

.62 

1.00 

6.97 

4.90 

4.45 

18.9 

13.1 

22.0 

16.8 

.... 

.... 

15.2 

20.2 

17.5 

21.5 

96.5 
87.2 
91.5 


18.0 


17.5 

19.6 

17.8 

18.0 
16.0 
15.8 
15.5 

n=15 

18.01 

16.1 

19.9 

1.24 

6.90 


19.5 
20.0 


11.7 

11.6 

13.7 
13.8 


12.3 
12.1 
11.4 

n=10 
12.96 
12.0 
14.1 

.69 
5.30 


16.0 
15.2 
14.5 


21.4 
20.4 
23.3 

23.8 

21.4 
20.4 

20.7 

n=9 

22.44 
20.8 
24.8 
1.44 
6.43 

27.5 
24.8 
23.3 


Hemphillian  specimen  from  Ash  Hollow  formation,  Nebraska 

4.    C.  rufus 

Recent,    male    .._  


Recent,  female 


82.0 


20.3 


13.5 


25.0 


n  =  64 

n=64 

n=64 

n=64 

89.66 

21.60 

14.40 

25.68 

83.2 

19.2 

13.2 

23.4 

99.5 

24.9 

16.0 

28.1 

3.37 

1.31 

.80 

1.04 

3.77 

6.08 

5.56 

4.05 

n=61 

n=61 

n=61 

n=61 

86.05 

20.98 

13.94 

24.52 

79.6 

18.1 

12.3 

22.1 

93.6 

25.2 

15.4 

27.1 

2.78 

1.47 

.72 

1.11 

3.24 

7.00 

5.16 

4.54 

152  MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY  NO.  6 


Inglis  IA,  Florida 

Melbourne,   Florida  

Port  Kennedy,   Pennsylvania 

5.    C.  armbrusteri 

Anita,   Arizona  

McCleod,  Florida 


Coleman  IIA,  Florida 


Cumberland  Cave,  Maryland 


6.    C.  lupus 

Recent,  western  U.S.,  male 


Recent,  western  U.S.,  female 


Maricopa,   California 


Rancho  La  Brea,  California 

Type  of  C.  milleri 

Goodland,   Kansas   

Millington,  Michigan  

Hay  Springs,  Nebraska 

Mullen,    Nebraska    _ 

Hermit's  Cave,  New  Mexico 


1 

2 

3 

4 

27.5 

87.5 

20.7 

15.6 

26.0 

94.0 

25.2 

15.0 

26.3 

— 

.... 

15.8 

— 

105.5 

27.5 

18.0 

102.5 

23.6 

16.4 

31.3 

113.0 

27.5 

17.8 

32.0 

27.4 





28.5 







30.9 

— 

.... 

28.0 
32.0 

104.0 

27.0 

17.3 

29.4 



18.2 



110.0 

28.2 

17.6 

30.5 

106.5 

28.2 

18.4 

30.0 

105.0 

28.7 

17.7 

30.5 

30.7 

.... 



30.9 

— - 

— 

— 

30.9 

n=62 

n=62 

n=62 

n=62 

95.31 

26.62 

15.68 

28.53 

86.7 

23.0 

13.6 

26.0 

104.0 

31.0 

17.0 

31.5 

3.44 

1.66 

.73 

1.20 

3.61 

6.25 

4.68 

4.22 

n=47 

n=47 

n=47 

n=47 

92.15 

24.90 

15.04 

27.07 

84.7 

22.5 

12.7 

25.1 

97.5 

27.7 

16.5 

30.0 

2.66 

1.29 

.74 

1.00 

2.88 

5.17 

4.95 

3.70 

17.5 

30.2 

27.5 

16.3 

29.1 

15.5 

28.4 

97.7 

27.5 

16.1 

30.5 

99.9 

29.1 

18.2 

31.2 

95.5 

26.8 

16.8 

29.0 

95.0 

25.8 

18.0 

32.0 

84.3 

22.1 

15.0 

26.3 

101.0 

28.0 

15.0 

30.0 

98.0 

29.0 

17.4 

29.9 

100.0 

30.0 

17.3 

29.3 

96.0 

30.6 

16.6 

30.2 

95.0 

24.6 

15.8 

29.9 

San  Josecito  Cave,  Nuevo  Leon 84.0  23.0  14.3  26.5 


1979                       NOWAK:    NORTH  AMERICAN  QUATERNARY  CANIS  153 

12  3  4 
7.    C.  dims 

Murray  Springs,  Arizona 110.0             ....  18.6  34.8 

18.4  33.9 

Cool  quarry,  California ... .            31.5  19.0 

Teichart  gravel  pit,  California  20.8 

Arroyo  Las  Positas,  California 110.0           31.9  19.0  34.3 

McKittrick,  California n=7          n=7  n=13  n=15 

113.40         33.73  20.08  35.33 

106.5           32.0  18.8  33.7 

123.0           35.0  21.7  37.0 

5.57          1.25  .89  .98 

4.91           3.71  4.41  2.78 

Maricopa,   California   n=10        n=12  n=18  n=17 

113.2           34.01  19.87  35.01 

108.0           29.8  18.0  33.5 

119.5           36.9  21.5  37.5 

3.82          2.38  1.04  1.47 

3.36           7.01  5.23  4.19 

Rancho  La  Brea,  California  n=73        n=73  n=73  n=73 

110.64         31.81  19.48  34.25 

102.0           25.5  17.9  31.8 

117.5           36.5  20.6  38.5 

3.31           1.96  .66  1.44 

2.99           6.17  3.36  4.20 

Ichetucknee  River,  Florida       117.0           32.5  20.3  35.5 

117.0           31.1  19.8  36.0 
116.5           33.2 

Hornsby  Springs,  Florida  _. . 116.0           29.0  18.3  35.5 

Reddick  IA,  Florida  ....              ....  19.3  34.4 

21.3  37.5 

Eichelberger  Cave,   Florida .._ 125.0           32.1  20.8  37.7 

34.2  20.9  37.6 

Melbourne,   Florida  112.5           31.6  19.3  35.3 

33.5 
34.8 
35.9 

Bradenton,    Florida   ..._ 32.5  18.4  36.2 

Twelve  Mile  Creek,  Kansas  ....              18.0 

Pendennis,  Kansas  ... .              .—              34.0 

Cragin  Quarry,  Kansas 108.5           30.5  18.3 

Welsh  Cave,  Kentucky 113.5           29.6  19.6  36.3 

Conkling  Cavern,  New  Mexico  32.1  20.4  35.2 

Hermit's  Cave,  New  Mexico 118.0             35.5 

Marlow,   Oklahoma _ 120.0           39.0  20.2  36.0 

Frankstown  Cave,  Pennsylvania  _ 108.6           30.0  17.9  32.0 

110.5           29.2  17.7  32.6 


154  MONOGRAPH  MUSEUM  OF  NATURAL  HISTORY 

1  2 

Ingleside,  Texas       118.0  34.0 

Rennick.   West   Virginia   .....  107.0  28.2 

Cedazo,  Aguascalientes  112.0 

33.0 
33.5 

San  Josecito  Cave.  Nuevo  Leon  - - n=9  n=10 

110.0  33.23 

107.0  30.8 

118.2  36.0 

3.32  1.63 

3.02  4.92 


NO.  6 


3 

4 

20.1 

36.0 

18.0 

31.8 

34.5 

34.5 

n=14 

n=18 

19.52 

34.64 

17.8 

32.7 

20.3 

36.5 

.71 

1.00 

3.64 

2.89 

Date  Due 


QE882.C15   N94    1979 

North    American  Quitcmar)   <   mis 

Harvard   MCZ    Library  \ll>uri 


3   2044   062   338   033