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CALIFORNIA 
FISH- GAME 

"CONSERVATION  OF  WILDLIFE  THROUGH  EDUCATION" 


California  Fish  and  Game  is  a  journal  devoted  to  the  conservation  of  wild- 
life. If  its  contents  are  reproduced  elsewhere,  the  authors  and  the  California 
Department  of  Fish  and  Game  would  appreciate  being  acknowledged. 

Subscriptions  may  be  obtained  at  the  rate  of  $5  per  year  by  placing  an 
order  with  the  California  Department  of  Fish  and  Game,  1416  Ninth  Street, 
Sacramento,  California  95814.  Money  orders  and  checks  should  be  made  out 
to  California  Department  of  Fish  and  Game.  Inquiries  regarding  paid  sub- 
scriptions should  be  directed  to  the  Editor. 


Complimentary  subscriptions  are  granted,  on  a  limited  basis,  to  libraries, 
scientific  and  educational  institutions,  conservation  agencies,  and  on  exchange. 
Complimentary  subscriptions  must  be  renewed  annually  by  returning  the  post- 
card enclosed  with  each  October  issue. 


Please  direct  correspondence  to: 

Kenneth  A.  Hashagen,  Jr.,  Editor 
California  Fish  and  Game 
1416  Ninth  Street 
Sacramento,  California  95814 


u 


VOLUME  67 


JANUARY  1981 


NUMBER  1 


Published  Quarterly  by 

STATE  OF  CALIFORNIA 

THE  RESOURCES  AGENCY 

DEPARTMENT  OF  FISH  AND  GAME 

— IDA— 


STATE  OF  CALIFORNIA 
EDMUND  G.  BROWN  JR.,  Governor 


THE  RESOURCES  AGENCY 
HUEY  D.  JOHNSON,  Secretary  for  Resources 


FISH  AND  GAME  COMMISSION 

ABEL  G.  GALLETTI,  President 
Los  Angeles 

RAYMOND  F.  DASMANN,  Vice  President  ELIZABETH  L.  VENRICK,  Ph.D.,  Member 
Nevada  City  Cardiff 

SHERMAN  CHICKERING,  Member  NORMAN  B.  LIVERMORE,  JR.,  Member 
San  Francisco  San  Rafael 


DEPARTMENT  OF  FISH  AND  GAME 
E.  C.  FULLERTON,  Director 

1416  9th  Street 
Sacramento  95814 


CALIFORNIA  FISH  AND  GAME 

Editorial  Staff 

Articles  appearing  in  this  issue  were  edited  by: 

Inland  Fisheries Kenneth  A.  Hashagen,  Jr. 

Marine  Resources  J.  R.  Raymond  Ally 

Wildlife Ronald  M.  Jurek 

Editor-in-Chief Kenneth  A.  Hashagen,  Jr. 


CONTENTS 


Page 


A  List  of  the  Freshwater  and  Anadromous  Fishes  of  California 

Leo  Shapovalov,  Almo  J.  Cordone,  and  William  A.  Dill        4 

Electrophoretic,  Morphometries  and  Meristic  Studies  of  Sub- 
populations  of  Northern  Anchovy,  Engraulis  mordax 
Andrew  M.  Vrooman,  Pedro  A.  Paloma,  and  James  R.  Zweifel       39 

Denning  Characteristics  of  Black  Bears,  Ursus  americanus,  in 
the  San  Bernardino  Mountains  of  Southern  California 
Harold  J.  Novick,  John  M.  Siperek,  and  Glenn  R.  Stewart      52 

Notes 

Update  of  the  Estimated  Mortality  Rate  of  Engraulis  mordax'm 

Southern  California  Doyle  Hanan       62 

First  Record  of  Dextrality  in  the  California  Tonguefish,  Sym- 

phurus  atricauda,  with  a  Second  Report  of  Ambicoloration 
Edward  L.  Telders      65 

Book  Reviews  69 


4  CALIFORNIA  FISH  AND  CAME 

Calif.  Fish  and  Came  67  ( 1 ) :  4-38     1 981 

A  LIST  OF  THE  FRESHWATER  AND  ANADROMOUS  FISHES 

OF  CALIFORNIA1 

LEO  SHAPOVALOV2,  ALMO  J.  CORDONE,  AND  WILLIAM  A.  DILL3 

Inland  Fisheries  Branch 

California  Department  of  Fish  and  Game 

1416  Ninth  Street 

Sacramento,  California  95814 

TABLE  OF  CONTENTS 

Page 

ABSTRACT 4 

INTRODUCTION  4 

PURPOSE 5 

SCIENTIFIC  NAMES  5 

COMMON  NAMES 7 

SCOPE 9 

Marine  Fishes  Successfully  Introduced  into  the  Salton  Sea 9 

Forms  and  Names  New  to  the  Main  List  Since  1959 10 

Forms  and  Names  Removed  from  the  Main  List  Since  1959  21 

REVISED  MAIN  LIST 23 

Native  Species  and  Established  Exotic  Species 23 

REVISED  SUPPLEMENTARY  LISTS 27 

Native  Species — Extinct  in  California 27 

Exotic  Species — Unsuccessfully  Introduced  or  of  Uncertain  Status 29 

ACKNOWLEDGMENTS 35 

REFERENCES  35 

This  list  is  the  second  revision  of  the  check  list  of  the  freshwater,  anadromous,  and 
euryhaline  fishes  of  California  published  by  Shapovalov  and  Dill  (1950)  and  first 
revised  by  Shapovalov,  Dill,  and  Cordone  (1959).  The  present  list  consists  of  a  main 
list  of  native  and  established  exotic  species  and  five  supplementary  lists:  (i)  native 
species  extinct  in  California,  (ii)  exotic  species  unsuccessfully  introduced  or  of 
uncertain  status,  (iii)  marine  fishes  successfully  introduced  into  the  Salton  Sea,  (iv) 
forms  and  names  new  to  the  main  list  since  1959,  and  (v)  forms  and  names  removed 
from  the  main  list  since  1959.  The  main  list  is  composed  of  124  full  species,  compris- 
ing 66  native  freshwater  and  anadromous  species,  13  native  euryhaline  or  marine 
species  which  occasionally  penetrate  into  fresh  water,  and  45  introduced  species. 
The  124  species  comprise  25  families  and  64  genera. 

INTRODUCTION 

Two  previous  editions  of  this  list  have  been  published  (Shapovalov  and  Dill 
1950;  Shapovalov,  Dill,  and  Cordone  1959).  Since  publication  of  the  1959  list, 
many  changes  have  occurred  in  both  the  composition  of  this  fauna  and  the 
nomenclature  applied  to  many  of  its  fishes. 

1  Accepted  for  publication  September  1980. 

2  Retired. 

3  Currently:  Fishery  Consultant,  730  North  Campus  Way,  Davis,  CA  95616. 


A  LIST  OF  CALIFORNIA  FISHES  5 

First,  a  number  of  fishes  have  been  introduced  into  the  State.  Some  of  these 
have  been  introduced  by  the  California  Department  of  Fish  and  Game  as  part 
of  its  research  and  management  program.  Others  have  been  introduced  illegally, 
either  deliberately  or  inadvertently,  especially  by  sportsmen  and  aquarists,  or 
through  escape  from  ornamental  fish  farms. 

Second,  some  forms  have  become  extinct  in  State  waters. 

Third,  some  new  forms  have  been  described  and  the  taxonomic  or  nomen- 
clatural  status  of  a  number  of  others  has  been  revised.  Some  of  these  revisions 
have  been  made  in  the  direction  of  condensation,  simplification,  and  uniformiza- 
tion  of  group  names;  others  have  been  in  the  opposite  direction  of  greater 
diversification.  With  full  recognition  that  opinions  on  taxonomy  and  nomencla- 
ture may  differ  decidedly,  we  have  attempted  to  include  in  the  list  all  revisions 
that  have  been  proposed  in  scientific  publications  and  not  subsequently  refuted. 

The  list  itself  is  preceded  by  several  introductory  sections.  Those  entitled 
"Scientific  Names"  and  "Common  Names",  which  are  of  a  background  nature, 
are  printed  here  with  little  change  from  our  previous  list. 

PURPOSE 

Two  major  objectives  in  publishing  a  check  list  of  California  freshwater  and 
anadromous  fishes  were  cited  in  the  1950  edition  and  reiterated  in  the  first 
revision  (1959).  These  were  to:  (i)  establish  the  basis  for  compilation  of  a 
detailed  handbook  of  these  fishes,  and  (ii)  promote  stability  and  uniformity  in 
both  their  common  and  scientific  names.  Publication  of  a  key  to  these  species 
by  Kimsey  and  Fisk  (1960)  and  especially,  the  publication  of  "Inland  Fishes  of 
California"  by  Moyle  (1976),  have  aided  in  achievement  of  the  first  goal.  The 
second  objective  has  neared  achievement  with  regard  to  common  or  vernacular 
names.  However,  uniformity  in  the  nomenclature  of  scientific  names  continues 
as  a  never-to-be  attained  goal. 

This  list,  like  the  previous  ones,  will  of  course  become  obsolete  in  time,  and 
another  edition  will  be  necessary.  We  suggest  that  its  future  authors,  or  any  who 
propose  to  publish  local,  state,  or  nationwide  lists,  can  materially  advance  stabil- 
ity in  fish  nomenclature  by  attempting  to  resolve  differences  through  consulta- 
tion with  those  who  have  authored  existing  lists.  We  have  done  this  consistently, 
have  invariably  met  with  cooperation,  and  have  thereby  resolved  most  nomen- 
clatural  problems. 

SCIENTIFIC  NAMES 

In  scientific  naming,  stability  is  largely  dependent  upon  the  thoroughness  and 
care  of  the  taxonomist.  Any  proposed  revisions  must  be  carefully  evaluated.  For 
example,  Schultz  (1957:48-49)  stated: 

"The  evaluation  of  generic  characters  and  recognition  of  genera  is  possible 
only  when  a  comprehensive  study  is  made  of  a  family  on  a  world-wide  basis 
and  when  there  is  established  the  nature  of  the  similarities  and  differences 
among  groups  of  species.  .  . 

"The  problem  of  how  far  to  progress  nomenclatorially  in  recognizing  generic 
categories  must  be  resolved  in  a  practical  manner  so  that  biologists  are  not 
presented  with  a  confusion  of  ill-defined  genera.  Usually  this  confusion  and  lack 
of  agreement  among  ichthyologists  and  fishery  biologists  results  from  inadequate 
studies  of  a  family.  Obviously,  no  dependable  solution  is  possible  on  how  many 


6  CALIFORMA  FISH  A\D  CAME 

genera  and  subgenera  to  recognize  in  a  t'amilv  until  the  zoological  relationships 
of  all  its  species  have  been  adequatek  compared  morphologically,  physiologi- 
calk ,  and  as  to  habits.  So  doubt,  after  this  work  has  been  done,  a  middle  of  the 
road  or  even  a  conservatke  attitude  on  the  number  of  phyletic  lines  to  name 
would  meet  \sith  general  acceptance.  Too  often  in  ichthyology  there  is  a  tend- 
encv  either  to  unite  genera  without  adequate  study  or  to  establish  new  genera 
without  an\  attempt  to  review  the  family.  The  least  confusion  results  if  the 
present  status  of  each  genus  in  a  family  is  retained  until  such  time  as  it  is 
thoroughk  studied." 

We  are  in  accord  with  this  opinion  but  believe  that  the  ideas  expressed  are 
applicable  to  species  and  subspecies  as  well.  Subspecies  in  particular  are  subject 
to  much  lumping  and  partitioning,  at  times  without  secure  evidence.  Some 
ichth\ ologists  have  questioned  the  existence  of  certain  forms  in  our  list  while, 
on  the  other  hand,  the\  ha\e  proposed  hitherto  unknown  forms  for  inclusion. 
In  almost  everv  case,  we  have  let  the  decision  hinge  on  the  appearance  in  the 
literature  of  substantiating  data.  The  publication  of  new  scientific  names  and 
elimination  of  familiar  ones  without  sufficient  supporting  evidence  simply  cre- 
ates further  confusion  in  fish  nomenclature. 

Bailey  1956:328-329  has  given  considerable  thought  to  the  problem  of 
subspecies:  ...  the  common  taxonomic  practice  of  dividing  geographically 
variable  species  into  named  races,  or  subspecies,  has  been  subjected  to  critical 
scrutiny.  It  has  been  noted  that  the  pattern  of  geographic  variation  in  some 
species  takes  the  form  of  a  rather  gradual  and  progressive  gradient,  termed  a 
c  ne.  it  is  now  agreed  bv  manv  taxonomists  that  despite  the  high  biological 
significance  of  this  tvpe  of  variation  it  is  undesirable  to  assign  subspecific  names 
on  the  basis  of  clinal  gradients.  .  . 

"Commonk  the  differences  between  geographic  subspecies  are  slight  and  are 
best  expressed  as  average  conditions  applving  to  a  considerable  fraction  of 
individuals,  but  not  to  all.  It  is  my  revised  opinion  that  acceptable  subspecies 
should  evidence  high  uniformitv  over  the  respective  ranges  and  should  differ 
one  from  another  with  high  constancy.  Zones  of  intergradation  should  be  rather 
narrow .  If  thev  are  wide  the  variation  merges  insensibly  into  a  clinal  gradient.  .  . 

"The  ichthvologist,  in  studying  material,  often  perceives  differences  among 
populations  from  v  anous  parts  of  the  geographic  range  of  a  species.  Such  discov- 
eries mav  presage  the  definition  of  validlv  recognizable  subspecies.  The  prema- 
ture use  of  such  information  w  ithout  publication  of  the  full  data  is  disconcerting 
to  other  ,*, or<ers,  who  are  unable  to  evaluate  the  basis  for  the  action.  The 
different  stocks  sometimes  turn  out  to  be  fully  distinct  species.  .  ." 

■Another  excellent  discussion  of  the  subject  which  supplements  the  above 
statements  was  presented  bv  Bailey,  Winn,  and  Smith  (1954:148-150).  The 
following  excerpt  seems  particularly  pertinent: 

Manv  clinal  v  anations  in  the  morphology  of  fishes  may  be  caused  partly  or 
wholly  bv  gradients  of  environmental  factors,  especially  temperature.  The  as- 
sumption that  all  taxonomic  characters,  such  as  meristic  counts,  are  governed 
solek  bv  genetic  factors  is  no  longer  tenable.  .  .  Whether  the  gradient  is  caused 
dv  hereditv  or  the  env  ironment,  we  reject  the  practice  of  establishing  subspecies 
on  characters  that  show  clinal  variation.  Furthermore,  the  insistence  that  a  dine 
be  a  perfectly  smooth  gradient,  we  regard  only  as  an  academic  problem.  Minor 
irregularities  are  to  be  anticipated  because  of  local  genetic  emphasis,  sampling 


A  LIST  OF  CALIFORNIA  FISHES  7 

errors,  environmental  variations  that  impose  structural  change,  and  other  vagar- 
ies." 

We  concur  in  the  statements  above  and  in  keeping  with  them  have  emploved 
binomials  instead  of  trinomials  wherever  sufficient  published  evidence  exists  to 
show  that  a  dine  truly  exists.  This  has  been  done,  for  example,  for  \otemigonus 
crysoleucas  'Hart  1952:  33-38,  77;  Bailey  et  al.  1954:  123-124,  149  I;  and  Ic- 
talurus  punctatus  'Bailey  et  al.  1954:  130).  Subspecific  partitioning  of  many 
species  in  the  main  list  may  be  of  questionable  validity;  however,  we  retain  the 
status  quo  and  await  the  publication  of  evidence  showing  whether  the  trinomials 
are  justified. 

Scientific  names  used  in  this  list  conform  to  the  provisions  of  the  International 
Code  of  Zoological  Nomenclature,  1964,  and  subsequent  amendments. 

Space  does  not  permit  an  explanation  of  each  change  in  scientific  names  used 
in  bringing  this  list  up  to  date.  However,  most  of  the  major  changes  are  discussed 
in  appropriate  text  sections.  Recourse  to  the  references  will  provide  further 
details.  Some  of  the  more  important  relativelv  recent  references  include:  Miller 
(1958),  Bond  '1961),  Walker,  Whitnev,  and  Barlow  '1961  I,  Bailev  and  Bond 
(1963),  Rosen  and  Bailey  '1963i,  Bailev  and  Uyeno  '1964>,  Smith  M966). 
Hubbs  (1967),  Kljukanov  I  1970),  Hopkirk  '1973i,  Ross  1 97*3  ,  Movie  I  1976 
and  Hubbs,  Follett,  and  Dempster  <1979>. 

COMMON  NAMES 

Stability  in  common  names  can  best  be  achieved  bv  adhering  closelv  to  a 
workable  set  of  criteria,  as  outlined  below. 

The  selection  of  common  names  for  fishes  in  this  list  is  complicated  bv  two 
somewhat  paradoxical  factors:  the  multiplicity  of  names  which  have  alreadv 
been  applied  to  certain  species  and,  in  the  case  of  certain  other  forms,  the  dearth 
of  common  names.  Thus,  members  of  the  genus  C\pnnodon  have  been  called 
by  such  varied  names  as  desert  minnow,  desert  killitish,  pursv  minnow,  pvgmv 
fish,  and  pupfish.  Converselv,  a  large  number  of  native  cvprinids  are  so  similar 
and  indistinctive  in  appearance  that  the  layman  has  never  recognized  their 
specific  differences  nor  called  them  bv  anv  name  other  than  the  rather  general 
ones,  such  as  chub  or  shiner.  This  list  attempts  to  reconcile  such  difficulties  bv 
assigning  one  official  common  name  to  each  species  and  subspecies. 

The  basic  rules  or  criteria  for  the  selection  of  common  names  remain  essential- 
ly identical  with  those  presented  in  our  prev  ious  lists.  Such  guides  are  necessarv 
to  prevent  arbitrary  selection  based  on  personal  preference,  and  have  again 
proved  of  practical  value  in  the  objectiv e  establishment  of  the  rev  ised  common 
names.  Insofar  as  possible,  we  have  adhered  to  them,  as  follows: 

1.  Names  should  agree  with  those  in  actual  common  use;  or  when  there  is 
no  common  or  vernacular  use,  with  those  in  published  literature.  Stnctlv 
"book  names"  should  be  avoided. 

2.  Names  should  agree,  if  possible,  with  those  in  other  authoritative  lists, 
especiallv  those  of  the  Committee  on  Names  of  Fishes  of  the  American 
Fisheries  Societv   i  Robins  et  al.  1980)  and  Hubbs  et  al.  (1979 

3.  Names  should  indicate  relationship  and  not  confuse  it. 

4.  Names  should  be  descriptive. 

5.  Preference  should  be  given  to  names  which  are  short,  distinctive,  interest- 
ing, catchy,  romantic,  or  euphonious. 


8  CALIFORNIA  FISH  AND  GAME 

Each  of  these  qualifications  has  exceptions  which  make  it  useless  by  itself. 
Therefore,  each  principle  listed  above  should  be  read  as  though  it  were  prefaced 
by  the  words,  "Other  considerations  being  equal  .  .  ."  For  example,  the  name 
Sacramento  perch  does  not  meet  either  Rule  3  or  4  above,  since  this  species 
(Archoplites  interruptus)  is  not  a  true  perch.  However,  since  this  name  is  so 
commonly  used  (Rule  1 )  and  since  it  agrees  fully  with  the  name  used  in  lists 
such  as  those  cited  in  Rule  2,  it  would  be  foolish  to  select  another. 

Aside  from  such  considerations,  in  this  revision,  as  in  the  previous  one,  we 
have  attempted  continued  advancement  of  the  twin  ideals  of  stability  for  individ- 
ual names  and  the  designation  of  relationships  through  the  selection  of  common 
names  according  to  a  definite  plan.  Such  aims,  long  recognized  by  ornithologists, 
are  well  exemplified  by  the  names  listed  in  "The  Distribution  of  the  Birds  of 
California"  (Grinnell  and  Miller  1944).  Thus,  in  our  list,  wherever  possible  the 
same  basic  common  name  has  been  given  to  all  members  of  a  single  genus,  with 
prefixes  added  to  that  common  name  for  each  full  species  of  that  genus.  In  the 
case  of  subspecies,  additional  prefixes  have  been  added  to  the  specific  name. 
For  example,  all  members  of  the  genus  Gila  have  been  termed  chub,  members 
of  the  Gila  bicolor  group  have  been  termed  tui  chub,  and  each  subspecies  of 
that  group  is  further  designated  by  an  additional  term  such  as  Mohave  for  G.  b. 
mohavensis,  the  Mohave  tui  chub. 

It  should  be  noted  that  this  method  will  permit  the  retention  of  at  least  part 
of  the  common  name  even  if  the  species  or  subspecies  undergoes  a  revision 
which  will  change  the  scientific  name.  This,  in  part,  answers  the  criticism  of  the 
Committee  on  Names  of  Fishes  of  the  American  Fisheries  Society  (Robins  et  al. 
1980):  "The  practice  of  applying  a  name  to  each  genus,  a  modifying  name  for 
each  species,  and  still  another  modifier  for  each  subspecies,  while  appealing  in 
its  simplicity,  has  the  defect  of  inflexibility."  Further,  "If  a  fish  is  transferred  from 
genus  to  genus,  or  shifted  from  species  to  subspecies  or  vice  versa,  the  common 
name  should  nevertheless  remain  unaffected.  It  is  not  a  primary  function  of 
common  names  to  indicate  relationship." 

We  contend,  nevertheless,  that  an  important  and  vital  function  of  common 
names  is  to  reveal  rather  than  confuse  relationships.  It  is  quite  true  that  some 
of  the  most  deeply  rooted  vernaculars  are  completely  misleading;  little  can  now 
be  done  in  these  cases  to  establish  meaningful  names.  Furthermore,  when  a 
name  is  entered  in  an  official  list  it  should  not  be  changed  unless  there  are 
important  reasons  to  do  so.  However,  changing  a  name  to  demonstrate  the 
proper  relationship  of  a  form  known  to  professional  fisheries  people  but  unfamil- 
iar to  laymen  does  not  present  a  serious  problem  and  to  us  is  justifiable.  In  any 
event,  long  usage  of  both  the  first  and  present  revisions  has  shown  that  the 
system  is  workable  and  has  meaning,  with  no  major  difficulties  encountered. 

Some  authors;  e.g.,  Robins  et  al.  (1980)  and  Alden  H.  Miller  (Grinnell  and 
Miller  1944),  believe  that  generally  only  full  species  deserve  common  names. 
Nevertheless,  we  have  listed  common  names  for  each  subspecies,  with  full 
recognition  that  a  number  of  them  may  not  endure.  One  reason  prompting  this 
decision  is  that  certain  subspecies  have  been  distinguished  as  entities  almost 
from  the  beginning,  and  it  would  seem  unfortunate  to  obscure  (through  omis- 
sion) such  names  as  Paiute  or  Kamloops. 

It  should  also  be  noted  that  a  number  of  systematists  have  disagreed  with 
certain  of  our  groupings;  e.g.,  that  for  the  native  trouts,  in  which  assignment  to 


A  LIST  OF  CALIFORNIA  FISHES  9 

specific  or  subspecific  status  is,  in  some  instances,  original  with  the  authors. 
However,  a  firm  nomenclature  has  never  been  developed  for  some  of  these 
plastic  groups.  And,  as  we  have  stated  before,  even  after  some  decided  changes 
in  scientific  nomenclature,  most  of  our  common  names  can  still  be  retained  with 
enough  recognizable  parts  to  promote  stability. 

SCOPE 

The  main  list  covers  both  native  and  successfully  established  exotic  species. 
The  supplementary  lists  include  native  species  believed  to  be  extinct  in  California 
and  exotic  species  unsuccessfully  introduced  or  of  uncertain  occurrence. 

We  have  attempted  to  include  all  native  forms  whose  occurrence  has  been 
reported  and  not  disproved  in  the  literature,  as  well  as  those  verified  through 
examination  of  collections.  The  existence  of  some  of  these  as  valid  species  or 
subspecies  {Catostomus  occidentalis  lacusanserinus,  for  example)  has  been 
questioned  by  some  workers.  Our  criterion  for  inclusion  of  such  forms  is  very 
simple:  we  have  tried  to  include  all  forms  whose  taxonomic  identity  has  not  yet 
been  disproved  in  published  literature. 

Possibly  certain  other  records  of  occurrence  are  based  on  misidentification. 
Possibly  some  of  the  native  species  are  no  longer  a  part  of  our  fauna.  Native 
forms  which  now  appear  to  be  extinct  in  State  waters  include  Salvelinus  malma, 
S.  confluentus,  Gila  crassicauda,  G  elegans,  Pogonichthys  ciscoides,  Ptycho- 
cheilus  lucius,  Cyprinodon  nevadensis  calidae,  and  C.  n.  shoshone.  It  is  practical- 
ly impossible,  however,  to  prove  or  disprove  such  suppositions.  Hence,  in  the 
case  of  the  native  species  it  has  been  thought  best  to  err  on  the  side  of  inclusive- 
ness  and  continue  them  in  the  main  list.  On  the  other  hand,  only  those  exotic 
or  introduced  species  of  which  breeding  populations  are  known  to  have  sur- 
vived are  included  in  this  list. 

Fishes  recorded  only  from  outside  California  have  not  been  included  even  if 
the  stream  in  question  flows  into  or  out  of  this  State;  e.g.,  the  Klamath  and 
Truckee  rivers.  However,  in  the  case  of  the  Colorado  River,  a  boundary  stream, 
fishes  recorded  from  the  Arizona  side  of  the  stream  have  been  included. 

Hybrids  have  also  been  omitted.  Both  interspecific  and  intergeneric  hybrids 
of  a  number  of  the  species  listed  have  been  recorded  from  the  natural  waters 
of  California  (see,  for  example,  Hubbs  and  Miller  1943). 

Marine  Fishes  Successfully  Introduced  into  the  Salton  Sea 
Most  of  the  fishes  in  the  main  list  are  strictly  freshwater  or  anadromous.  For 
the  sake  of  completeness,  we  have  also  listed  those  marine  and  brackishwater 
species  which  we  know  have  penetrated  into  fresh  water.  Strictly  marine  species 
from  the  Gulf  of  California  which  have  been  introduced  into  and  have  success- 
fully spawned  in  the  Salton  Sea,  an  inland  body  of  water  with  salinity  exceeding 
that  of  ocean  water  are,  however,  omitted  from  the  main  list.  They  are  included 
below,  since  they  have  established  breeding  populations  in  an  inland  body  of 
water.  The  history  of  these  introductions  by  the  California  Department  of  Fish 
and  Game  has  been  related  by  Anon.  (1958)  and  Walker  et  al.  (1961 ). 

HAEMULIDAE— grunt  family 
Anisotremus  davidsonii  (Steindachner) — sargo 

Introduced  in  1951.  The  first  sargo  known  to  have  been  spawned  in  the  Sea, 
a  juvenile  young-of-the-year,  was  taken  in  October  1956.  The  first  verified  catch 
2—81475 


10  CALIFORNIA  FISH  AND  GAME 

of  an  adult  was  made  on  17  September  1958.  Since  then  sargo  up  to  305  mm 
in  length  have  been  taken  in  considerable  numbers  by  sport  fishermen. 

SCIAENIDAE— croaker  family 
Bairdiella  icistia  (Jordan  and  Gilbert) — bairdiella 

First  introduced  in  October  1950,  the  population  of  bairdiella  is  now  very 
large. 

Cynoscion  xanthulus  Jordan  and  Gilbert — orangemouth  corvina 

First  introduced  in  October  1950,  it  is  now  present  in  large  numbers,  and  like 
the  sargo  and  bairdiella,  should  remain  so  unless  the  salinity  of  the  Sea  becomes 
too  high. 

The  shortfin  corvina,  Cynoscion  parvipinnis,  also  introduced  in  1950,  estab- 
lished a  breeding  population  but  has  not  been  observed  for  a  number  of  years. 

Forms  and  Names  New  to  the  Main  List  Since  1959 
Numerous  changes  in  scientific  and  common  names  have  taken  place  since 
the  1959  check  list  was  prepared.  Changes  involving  common  names  and  minor 
revisions  in  scientific  names  are  not  discussed.  Forms  and  scientific  names  not 
listed  in  or  differing  from  those  listed  in  the  1959  check  list  are  included  in  this 
revised  edition,  with  a  brief  explanation  for  their  inclusion.  Included  are  19 
species  and  subspecies  of  exotic  fishes  which  have  become  established  in  Cali- 
fornia waters  since  1959. 

Although  the  California  freshwater  fish  fauna  has  been  studied  for  many  years, 
some  undiscovered  species  may  remain.  Collecting  in  coastal  fresh  waters  may 
uncover  additional  euryhaline  forms.  Taxonomists  may  be  expected  to  continue 
to  describe  new  forms  but  at  a  lesser  rate  than  in  the  past.  For  example,  some 
taxonomists  have  recognized  a  trout  from  northern  California  as  a  distinct  spe- 
cies and  have  proposed  the  common  name  of  redband  trout,  but  have  not  yet 
published  a  scientific  name  (Hoopaugh  1974).  The  escape  or  release  into  the 
wild  of  tropical  and  other  ornamental  fishes  may  be  anticipated  and  some  of 
these  may  become  established. 

And,  although  such  activities  have  a  much  lower  priority  now  than  in  the  past, 
the  introduction  of  exotic  game  and  forage  fishes  by  the  California  Department 
of  Fish  and  Game  may  also  result  in  addition  of  other  species.  The  fish  manage- 
ment program  of  the  Inland  Fisheries  Branch  includes  an  evaluation  of  the 
various  aquatic  habitats  and  what  might  constitute  the  most  suitable  game 
and /or  forage  species,  either  native  or  exotic,  for  them.  Each  potential  import 
is  thoroughly  studied  and  screened  to  insure  against  detriment  to  existing  aquatic 
resources. 

PETROMYZONTIDAE— lamprey  family 
Lampetra  folletti  (Vladykov  and  Kott) — Modoc  brook  lamprey 

Vladykov  and  Kott  (19766)  described  this  nonparasitic  species  of  lamprey 
from  the  Klamath  River  system  in  Modoc  County,  California,  as  Entosphenus 
folletti.  We  follow  Hubbs  (1971)  in  treating  Entosphenus  as  a  subgenus  of 
Lampetra. 


A  LIST  OF  CALIFORNIA  FISHES  1 1 

Lampetra  hubbsi  (Vladykov  and  Kott) — Kern  brook  lamprey 

Vladykov  and  Kott  (1976a)  described  this  nonparasitic  species  of  lamprey 
from  the  Friant-Kern  Canal,  east  of  Delano,  San  Joaquin  Valley,  as  Entosphenus 
hubbsi.  We  follow  Hubbs  (1971)  in  treating  Entosphenus  as  a  subgenus  of 
Lampetra. 

Lampetra  lethophaga  Hubbs — Pit-Klamath  brook  lamprey 

The  addition  of  this  species  is  based  on  its  description  by  Hubbs  (1971 ).  It 
is  found  in  the  drainage  basin  of  the  Pit  River  in  northeastern  California,  and  in 
the  upper  Klamath  River  in  south-central  Oregon.  In  the  past  it  has  been  misiden- 
tified  as  Lampetra  planeri  and  Entosphenus  tridentatus. 

Lampetra  pacifica  Vladykov — Pacific  brook  lamprey 

This  small,  nonparasitic  lamprey  was  described  as  a  new  species  by  Vladykov 
(1973).  In  California,  it  is  recorded  from  various  streams  in  the  Sacramento-San 
Joaquin  River  system.  It  is  quite  similar  to  L.  richardsoni and  may  not  be  specifi- 
cally distinct  from  it.  Before  1973  it  had  frequently  been  recorded  as  L.  planeri 
or  L.  richardsoni. 

Lampetra  richardsoni  Vladykov  and  Follett — western  brook  lamprey 

Vladykov  and  Follett  (1965)  described  this  new  nonparasitic  species  of  lam- 
prey from  "...  streams  of  British  Columbia,  Washington,  Oregon,  and  possi- 
bly Alaska".  Follett  subsequently  informed  J.  D.  Hopkirk  that  the  range  of  the 
western  brook  lamprey  was  more  recently  known  to  include  California  ( Hopkirk 
1973:20).  Various  authors  had  previously  listed  it  as  L.  planeri,  the  name  used 
in  our  1959  check  list,  but  now  removed  from  our  main  list. 

Lampetra  tridentata  (Gairdner) — Pacific  lamprey 

The  Pacific  lamprey  was  listed  as  Entosphenus  tridentatus  in  our  1950  and 
1959  check  lists,  but  we  now  follow  Hubbs  (1971 )  in  treating  Entosphenus  as 
a  subgenus  of  Lampetra. 

ACIPENSERIDAE— sturgeon  family 
Acipenser  medirostris  medirostris  Ayres — American  green  sturgeon 

We  follow  Lindberg  and  Legeza  (1965:33)  in  recognizing  this  subspecies.  In 
our  1959  check  list  we  listed  only  the  full  species,  Acipenser  medirostris  Ayres. 

CLUPEIDAE— herring  family 
Clupea  harengus  /?a//a5/'/ Valenciennes — Pacific  herring 

In  our  1959  list  the  Pacific  herring  was  listed  as  Clupea  pallasii.  However, 
Svetovidov  (1952)  has  shown  that  this  form  is  actually  a  subspecies  of  C. 
harengus. 

OSMERI DAE— smelt  family 
Hypomesus  nipponensis  McAllister — freshwater  smelt 

This  species  was  introduced  into  California  from  Japan  as  a  forage  fish  (air 
shipment  of  eggs)  in  1959  (Wales  1962).  At  the  time  it  was  misidentified  as  H. 
olidus.  This  strictly  freshwater  species  has  since  become  firmly  established  in  at 
least  several  waters  in  California. 


12  CALIFORNIA  FISH  AND  CAME 

Hypomesus  transpacificus  McAllister — delta  smelt 

In  his  revision  of  the  smelt  family,  McAllister  (1963)  described  this  new 
species,  known  only  from  the  lower  parts  of  the  Sacramento  and  San  Joaquin 
rivers.  It  had  previously  been  referred  to  in  the  literature  as  Hypomesus  olidus, 
the  name  we  used  in  our  1959  check  list. 

McAllister  described  two  subspecies,  H.  transpacificus  transpacificus  and  H. 
transpacificus  nipponensis.  However,  we  follow  Kljukanov  (1970)  in  treating  the 
two  as  distinct  species. 

COREGONIDAE— whitefish  family 
Prosopium  williamsoni  (Girard) — mountain  whitefish 

Our  1959  list  placed  this  species  in  the  genus  Coregonus.  We  now  follow 
Norden  (1961 ),  who  described  the  characters  separating  the  two  genera. 

SALMONIDAE — salmon  and  trout  family 
Salmo  clarkii  pleuriticus  Cope — Colorado  River  cutthroat  trout 

This  subspecies  was  dropped  from  the  main  list  in  our  1959  check  list  because 
published  reports  of  its  occurrence  in  the  Salton  Sea  were  dubious.  The  reported 
specimens  may  have  been  misidentified;  in  any  case,  if  correctly  identified  they 
almost  certainly  consisted  of  individuals  washed  into  the  basin  from  the  Colo- 
rado River  many  years  ago.  No  specimens  from  the  Salton  Sea  are  known  to  exist 
in  any  collections. 

On  1 1  September  1 974,  the  California  Department  of  Fish  and  Game  collected 
21  specimens  of  this  subspecies  from  the  lower  three  of  the  five  Williamson 
Lakes  of  the  southern  Sierra  Nevada.  These  trout  were  descendant  from  a  1931 
plant  of  Colorado  River  cutthroat  trout  fry  hatched  from  eggs  taken  from  Trap- 
per's Lake,  Colorado  (Gold,  Gall,  and  Nicola  1978). 

Salvelinus  confluentus  (Suckley) — bull  trout 

Although  the  view  that  the  Dolly  Varden,  Salvelinus  malma,  is  the  only  recog- 
nizable member  of  the  genus  in  the  American  northwest  has  been  widely  ac- 
cepted, the  subject  has  been  a  matter  of  some  controversy  for  over  a  century. 
Morton  (1970)  concluded  that  S.  malma  was  the  only  valid  species  and  that 
there  were  no  valid  subspecies.  More  recently,  Cavender  (1978)  presented 
morphometric,  meristic,  osteological,  and  distributional  evidence  to  support  his 
view  that  there  are  two  widely  distributed  forms  of  Salvelinus  native  to  the 
western  United  States  and  Canada:  the  Dolly  Varden,  S.  malma,  and  the  bull 
trout,  S.  confluentus.  He  records  both  species  from  the  McCloud  River  drainage 
in  California,  although  his  only  record  from  there  of  S.  malma  consists  of  two 
specimens  in  the  National  Museum  of  Natural  History  (then  U.S.  National 
Museum)  labeled  as  having  been  sent  by  Livingston  Stone  from  the  McCloud 
River  in  1877.  It  is  on  the  basis  of  this  publication  that  we  have  included  both 
species  in  our  main  list,  even  though  we  think  it  virtually  inconceivable  that  both 
species  could  have  coexisted  within  the  confines  of  the  McCloud  River. 

CYPRINIDAE — carp  or  minnow  family 
Gila  bicolor  (Girard) — tui  chub 

Bailey  and  Uyeno  (1964)  changed  the  name  of  this  species  from  Siphateles 
bicolor,  the  name  used  in  our  1959  check  list,  to  Gila  bicolor. 


A  LIST  OF  CALIFORNIA  FISHES  13 

Gila  bicolor  mohavensis  (Snyder) — Mohave  tui  chub 

Although  this  fish  had  been  accorded  full  species  rank  for  many  years,  Miller 
( 1 973 )  regarded  it  as  a  subspecies  because  he  was  unable  to  discover  characters 
that  would  separate  it  specifically  from  all  populations  of  Gila  bicolor  in  the 
Lahontan  Basin. 

Gila  bicolor  snyderi  Miller — Owens  tui  chub 

This  subspecies  was  described  by  Miller  (1973).  In  our  previous  check  list  it 
was  listed  as  Siphateles  bicolor  obesus.  It  is  confined  to  the  isolated  Owens 
Valley  in  eastern  California. 

Gila  bicolor  thalassina  (Cope) — Goose  Lake  tui  chub 

This  subspecies  was  not  included  in  the  1950  and  1959  check  lists  because  of 
the  belief  that  it  was  extinct  in  Goose  Lake,  Modoc  County  (Hubbs  and  Miller 
1948:70-71 ).  A  prolonged  drought  (1929-1934),  when  Goose  Lake  was  virtually 
dry,  may  have  led  Hubbs  and  Miller  to  this  conclusion.  Recent  collections  made 
by  T.J.  Mills  (Calif.  Dep.  Fish  and  Game,  pers.  commun.)  revealed  that  this  chub 
is  once  again  abundant  in  Goose  Lake.  Its  identity  as  G  b.  thalassina  was 
confirmed  by  C.  E.  Bond  (15  August  1978  letter  to  T.  J.  Mills). 

Gila  bicolor  vaccaceps  Bills  and  Bond — Cowhead  Lake  tui  chub 

Tui  chubs  from  Cowhead  Lake,  Modoc  County,  were  first  recognized  as 
distinct  by  Hubbs  and  Miller  (1948)  and  ultimately  described  by  Bills  and  Bond 
(1980).  The  Lake  is  now  dry  and  the  chubs  are  confined  to  the  small  outlet 
slough. 

Gila  coerulea  (Girard) — blue  chub 

This  species,  from  the  Klamath  River  system,  was  listed  in  our  1959  check  list 
as  Gila  bicolor.  Bailey  and  Uyeno  (1964)  have  explained  why  it  should  be  called 
G  coerulea. 

Gila  elegans  Baird  and  Girard — bonytail  chub 

In  our  1959  check  list  we  used  the  name  Gila  robusta,  and  treated  the  form 
from  the  Colorado  River  as  a  subspecies,  G  robusta  elegans.  G  robusta  elegans 
is  regarded  as  having  specific  status  by  Minckley  and  Deacon  (1968)  and 
Hopkirk  (1973:32). 

Hesperoleucus  symmetricus  mitrulus  Snyder — upper  Pit  western  roach 
Hesperoleucus   symmetricus   navarroensis  Snyder — Navarro    western 

roach 
Hesperoleucus  symmetricus  parvipinnis  Snyder — Gualala  western  roach 
Hesperoleucus  symmetricus  venustus  Snyder — Venus  western  roach 

In  our  1959  check  list  these  subspecies  were  accorded  full  specific  rank.  We 
now  concur  with  Moyle  (1976:180)  and  Hubbs  et  al.  (1979)  that  they  should 
be  treated  as  subspecies  of  H.  symmetricus.  Hopkirk  (1973:  48-51)  discusses 
some  of  the  taxonomic  problems  involved  and  the  need  for  a  thorough  revision 
of  the  genus. 


14  CALIFORNIA  FISH  AND  CAME 

Lavinia  exilicauda  chi  Hopkirk — Clear  Lake  hitch 

Hopkirk  (1973:55-56)  described  this  subspecies  from  Clear  Lake  in  central 
California,  separating  it  from  Lavinia  exilicauda  exilicauda  of  previous  authors. 
He  remarked  that  it  ".  .  .  is  a  lake-adapted  subspecies  with  a  high  number  of 
gill  rakers.  In  this  respect,  it  agrees  with  Pogonichthys  ciscoides  and  Hysterocar- 
pus  traskii  lagunae  from  Clear  Lake  basin." 

Pogonichthys  ciscoides  Hopkirk — Clear  Lake  splittail 

Hopkirk  (1973:30-31 )  described  this  species  from  Clear  Lake  in  central  Cali- 
fornia, distinguishing  it  from  Pogonichthys  macrolepidotus  of  previous  authors. 
He  noted  that  it  ".  .  .  is  a  lake-adapted  species  with  fine  gill  rakers,  terete  body, 
terminal  mouth,  and  small  fins." 

CATOSTOM I  DAE— sucker  family 
Catostomus  fumeiventris  Miller — Owens  sucker 

This  species  was  described  by  Miller  (1973).  Originally  confined  to  the  Ow- 
ens Valley  in  eastern  California,  it  has  been  introduced  into  June  Lake  in  the 
Mono  Lake  Basin,  and  possibly  into  the  Santa  Clara  River  Basin  by  way  of  the 
Los  Angeles  Aqueduct. 

Catostomus  luxatus  (Cope) — Lost  River  sucker 

We  follow  Hubbs  et  al.  (1979)  in  placing  the  species  listed  in  our  1959  edition 
as  Deltistes  luxatus  in  the  genus  Catostomus. 

Catostomus  occidentalis  humboldtianus  Snyder — Humboldt   western 

sucker 
Catostomus  occidentalis  mniotiltus  Snyder — Monterey  western  sucker 

These  subspecies  were  treated  as  full  species  in  our  1959  list.  They  are  current- 
ly recognized  as  subspecies  of  Catostomus  occidentalis  (Hopkirk  1973:69; 
Moyle  1976:214;  Hubbs  et  al.  1979). 

Catostomus  platyrhynchus  (Cope) — mountain  sucker 

In  our  1959  check  list  we  listed  Pantosteus  lahontan,  Lahontan  mountain 
sucker.  Smith  (1966)  united  Pantosteus  platyrhynchus and  P.  lahontan  as  Catos- 
tomus platyrhynchus. 

Catostomus  santaanae  (Snyder) — Santa  Ana  sucker 

In  our  1959  check  list  this  species  was  listed  as  Pantosteus  santaanae  Snyder. 
Smith  (1966)  relegated  Pantosteus  to  a  subgenus  of  Catostomus. 

COBITIDIDAE— loach  family 
Misgurnus  anguillicaudatus  (Cantor) — oriental  weatherfish 

On  12  April  1968,  J.  A.  St.  Amant  collected  loaches  in  a  portion  of  the 
Westminster  flood  control  channel,  Orange  County  (St.  Amant  and  Hoover 
1969).  Identified  as  Misgurnus  anguillicaudatus  by  C.  L.  Hubbs,  this  was  the  first 
verified  record  of  free-living  loaches  in  California.  Their  source  is  believed  to  be 
the  Pacific  Goldfish  Farm,  from  which  some  loaches  escaped  into  the  channel 
as  early  as  the  1930's.  A  thriving  population  was  present  upstream  from  the 
original  collection  site  in  1977  and  another  population  was  discovered  in  the 
adjacent  Bolsa  Chica  Channel  in  1979  (F.  G.  Hoover,  pers.  commun.). 


A  LIST  OF  CALIFORNIA  FISHES  1  5 

ICTALURIDAE — North  American  freshwater  catfish  family 
Ictalurus  furcatus  (Lesueur) — blue  catfish 

The  blue  catfish  is  presently  established  in  four  reservoirs  and  several  ponds 
in  San  Diego  and  Riverside  counties  and  several  ponds  at  the  Imperial  Wildlife 
Area  in  Imperial  County.  The  initial  plant  of  blue  catfish  in  California  was  made 
by  the  California  Department  of  Fish  and  Game  in  October  1966,  when  1,758 
fish  from  Stuttgart,  Arkansas,  were  released  in  Lake  Jennings,  San  Diego  County 
(Richardson  et  al.  1970).  A  single  1.7-kg  specimen  was  collected  from  the  San 
Joaquin  River  near  Mossdale,  San  Joaquin  County,  in  December  1978  by  the 
Department's  Bay-Delta  Study  (Taylor  1980).  Currently  about  20  commercial 
fish  farmers  in  California  are  licensed  to  rear  and  sell  this  species. 

Pylodictis  olivaris  (Rafinesque) — flathead  catfish 

A  collection  of  four  young-of-the-year  specimens  from  the  Highline  Canal  and 
its  tributaries,  near  Niland,  Imperial  County,  constituted  the  first  California  record 
for  this  species  (Bottroff,  St.  Amant,  and  Parker  1969).  They  were  probably 
progeny  from  the  original  introduction  by  the  Arizona  Game  and  Fish  Depart- 
ment of  600  fish  into  the  Colorado  River  above  Imperial  Dam.  The  flathead 
catfish  is  now  common  in  the  Colorado  River  and  adjacent  waters  from  Imperial 
Dam  upstream  to  Headgate  Rock  Dam  near  the  town  of  Parker.  It  is  also 
common  in  the  All  American  Canal  system,  including  various  drains  and  canals 
in  Imperial  Valley. 

CYPRINODONTIDAE— killifish  family 
Cyprinodon  milleri  LaBounty  and  Deacon — Cottonball  Marsh  pupfish 

LaBounty  and  Deacon  (1972)  described  this  pupfish  from  Cottonball  Marsh, 
located  in  an  isolated  sector  of  the  northwest  portion  of  Death  Valley.  Previously 
these  pupfish  had  been  considered  to  be  a  population  of  C.  salinus. 

Lucania  parva  (Baird) — rainwater  killifish 

Hubbs  and  Miller  (1965)  describe  the  establishment  of  this  cyprinodont  in 
streams  and  sloughs  tributary  to  San  Francisco  Bay  and  in  Irvine  Lake,  Orange 
County.  With  respect  to  the  Bay,  where  it  was  first  recorded  in  1958,  the  authors 
state,  "It  is  obvious  that  Lucania  parva  has  become  well  established  about  San 
Francisco  Bay  and  contiguous  waters,  with  vast  increase  in  numbers  and  in 
range."  However,  only  a  few  specimens  (three  in  November  1963  and  six  in 
June  1964)  were  taken  from  Irvine  Lake  and  the  status  of  this  population  is 
unknown.  Another  population  was  discovered  in  1976  in  Arroyo  Seco  Creek,  a 
tributary  of  Vail  Lake,  Riverside  County  (McCoid  and  St.  Amant  1980). 

POECILIIDAE— livebearer  family 
Poecilia  latipinna  (Lesueur) — sailfin  molly 

In  our  1959  check  list  we  listed  Mollienesia  latipinna.  Mollienesia  was  synony- 
mized  with  Poecilia  by  Rosen  and  Bailey  (1963).  The  1959  report  mentioned 
that  this  species  was  established  in  canals  and  ditches  tributary  to  the  Salton  Sea. 
It  is  now  by  far  the  most  abundant  species  in  these  habitats,  as  well  as  in  the 
shallow  margins  of  the  Sea  itself  (Black  1980). 


16  CALIFORNIA  FISH  AND  CAME 

Poecilia  mexicana  mexicana  (Steindachner) — Orizaba  shortfin  molly 

The  Orizaba  shortfin  molly  has  been  established  in  the  Salton  Sea  area  for 
many  years.  It  was  first  reported  in  1964  from  a  small  pond  and  its  tributary  about 
8  km  north  of  the  Salton  Sea  (St.  Amant  1966).  Further  collections  were  made 
in  this  general  area  in  subsequent  years. 

Populations  of  shortfin  mollies  have  persisted  in  scattered  locations  in  the 
drains  and  natural  watercourses  entering  the  Salton  Sea  and  in  the  margins  of 
the  Sea  itself  (Black  1980).  Although  much  less  abundant  and  widespread  here 
than  the  sailfin  molly,  Poecilia  latipinna,  it  may  nevertheless  be  considered  a 
permanent  member  of  the  fish  fauna  in  these  waters. 

Poeciliopsis  gracilis  (Heckel) — Porthole  livebearer 

Mearns  (1975)  reported  the  collection  of  four  specimens  of  this  species  on 
27  July  1974,  from  an  irrigation  canal  near  Mecca,  Riverside  County.  He  suggest- 
ed the  common  name  porthole  livebearer.  The  specimens  were  identified  by  C. 
L.  Hubbs.  Later  in  the  year  Mearns  collected  additional  specimens  at  the  same 
site.  The  presence  of  recently  born  young,  the  wide  range  of  sizes,  and  the 
persistence  of  the  fish  for  at  least  a  4-month  period  suggested  that  P.  gracilis  was 
a  reproducing  resident  of  this  canal.  Introduction  was  presumably  through  direct 
release  by  aquarists  or  escapement  from  a  nearby  tropical  fish  farm.  Additional 
collections  of  this  species  from  the  same  canal  have  been  made  as  late  as  1980 
(J.  A.  St.  Amant,  pers.  commun.). 

ATHERINIDAE— silverside  family 
Menidia  audens  Hay — Mississippi  silverside 

The  Mississippi  silverside  was  introduced  into  the  Blue  Lakes  and  Clear  Lake 
in  Lake  County  in  1967  to  test  its  effectiveness  in  controlling  the  Clear  Lake  gnat 
andchironomid  midges  (Cook  and  Moore  1970).  These  fish  were  obtained  from 
Lake  Texoma,  Oklahoma.  The  Blue  Lakes  plant  was  authorized  by  the  Fish  and 
Game  Commission  whereas  the  Clear  Lake  plant  was  not.  About  6,000  fish  were 
released  in  Upper  Blue  Lake  and  3,000  in  Lower  Blue  Lake  and  Clear  Lake. 
Within  a  year  progeny  from  the  original  plant  were  abundant  in  the  last  two 
waters,  and  since  then  a  virtual  population  explosion  of  silversides  has  taken 
place. 

A  combination  of  experimental  introductions  by  the  Department  of  Fish  and 
Game,  illegal  introductions  by  bait  fishermen,  and  dispersal  via  man-made  wa- 
terways has  resulted  in  wide  distribution  of  this  species.  Moyle,  Fisher,  and  Li 
(1974)  reported  the  presence  of  silversides  in  Putah  and  Cache  creeks  in  Yolo 
County  and  in  eight  reservoirs  and  ponds  in  Alameda  and  Santa  Clara  counties. 
Collections  described  by  Meinz  and  Mecum  (1977)  demonstrated  the  occur- 
rence of  an  abundant,  reproducing  population  in  the  Sacramento-San  Joaquin 
Delta.  From  here  they  have  ready  access  to  the  California  Aqueduct,  the  Delta- 
Mendota  Canal,  and  associated  water  storage  and  conveyance  systems  and 
eventually  southern  California  reservoirs. 

SYNGNATHIDAE— pipefish  family 
Syngnathus  leptorhynchus  Girard — bay  pipefish 

The  bay  pipefish  has  been  recorded  from  the  mouth  of  the  San  Lorenzo  River, 
Santa  Cruz  County,  and  from  the  Navarro  River,  Mendocino  County  (Moyle 
1976:283). 


A  LIST  OF  CALIFORNIA  FISHES  17 

COTTIDAE— sculpin  family 
Cottus  perplexus  Gilbert  and  Evermann — reticulate  sculpin 

A  collection  of  reticulate  sculpins  was  made  from  the  Middle  Fork  of  the 
Applegate  River  (Rogue  River  drainage)  in  California  on  2  March  1971,  by  F. 
H.  Everest  and  recorded  by  Bond  ( 1973) .  Cottus  perplexus  is  the  most  abundant 
representative  of  the  genus  in  the  Rogue.  It  is  not  known  from  coastal  streams 
south  of  the  Rogue. 

Cottus  pitensis  Bailey  and  Bond — Pit  sculpin 

Bailey  and  Bond  ( 1 963 )  described  this  sculpin  as  a  new  species.  This  common 
species  of  the  Pit  river  system  in  northeastern  California  had  been  collected 
frequently  over  the  years  but  had  been  considered  to  be  Cottus  gulosus,  except 
by  Bond  (1961 ),  who  treated  it  as  an  undescribed  species. 

PERCICHTHYIDAE— temperate  bass  family 
Morone  chrysops  (Rafinesque) — white  bass 

Von  Geldern  (1966)  described  the  original  introductions  of  white  bass  into 
California  by  the  California  Department  of  Fish  and  Game,  under  the  name 
Roccus  chrysops.  We  follow  Robins  et  al.  (1980)  and  others  in  placing  this 
species  in  the  genus  Morone. 

About  160  fingerlings  were  planted  in  Nacimiento  Reservoir,  San  Luis  Obispo 
County,  in  November  1965  and  64  adults  were  released  into  the  same  water  in 
February  1 966.  The  fingerlings  were  obtained  from  Lake  McConaughy  in  Nebras- 
ka and  the  adults  from  Tenkiller  Reservoir  in  Oklahoma.  Additional  plants  in 
Nacimiento  included  600  yearlings  and  adults  in  July  1968  from  Lahontan  Reser- 
voir in  Nevada  and  200  adults  in  February  1967  from  Utah  Lake  in  Utah.  The 
Nacimiento  population  is  now  well  established. 

The  California  Department  of  Fish  and  Game  and  the  Arizona  Game  and  Fish 
Department  cooperated  in  a  series  of  plants  of  white  bass  in  the  lower  Colorado 
River  in  1968  and  1969.  However,  the  species  failed  to  become  established  in 
this  location. 

The  popularity  of  white  bass  at  Nacimiento  Reservoir  has  led  to  illegal  intro- 
ductions into  other  waters  of  the  State.  One  such  water  is  Kaweah  Reservoir, 
Tulare  County,  where  it  is  firmly  established. 

Morone  saxatilis  (Walbaum) — striped  bass 

In  the  1959  list  this  species  was  listed  as  Roccus  saxatilis.  We  follow  Robins 
et  al.  (1980)  and  others  in  placing  it  in  the  genus  Morone. 

CENTRARCHIDAE— sunfish  family 
Lepomis  gulosus  (Cuvier) — warmouth 

The  warmouth  was  designated  Chaenobryttus gulosus  in  our  1 959  list.  Howev- 
er, for  reasons  described  by  Bailey  et  al.  (1970:75),  we  believe  that  gulosus 
should  be  regarded  as  a  species  of  Lepomis. 

Lepomis  macrochirus  purpurescens  Cope — southeastern  bluegill 

In  June  1975,  88  adult  southeastern  bluegill  were  stocked  in  Perris  Lake, 


18  CALIFORNIA  FISH  AND  CAME 

Riverside  County,  by  the  California  Department  of  Fish  and  Game  (Henry 
1979).  They  were  obtained  through  the  cooperation  of  the  Florida  Game  and 
Fresh  Water  Fish  Commission  from  one  of  its  hatcheries.  They  have  reproduced 
and  are  firmly  established.  Specimens  collected  from  Perris  Lake  have  been 
stocked  in  several  small  ponds  for  experimental  purposes  and  use  as  broodstock 
for  future  plants. 

Micropterus  coosae  Hubbs  and  Bailey — redeye  bass 

Kimsey  (1954)  recorded  the  original  importation  into  California  of  40  redeye 
bass  for  use  as  broodstock  by  the  California  Department  of  Fish  and  Game  at 
Central  Valleys  Hatchery,  Elk  Grove,  California.  In  reviewing  the  history  and 
status  of  this  introduction  (Kimsey  1957)  concluded,  "No  redeye  bass  were 
planted  in  the  open  waters  of  the  State  and  none  are  now  present  in  California." 

A  second  attempt  to  establish  the  redeye  bass  in  California  was  successful 
(Goodson  1966).  Broodstock  imported  from  Tennessee  and  Georgia  in  the 
spring  of  1968  spawned  successfully  at  Central  Valleys  Hatchery,  and  their 
progeny  were  stocked  in  seven  widely  separated  waters:  Lake  Oroville,  Butte 
County;  Alder  Creek,  Sacramento  County;  South  Fork  Stanislaus  River,  Tuol- 
umne County;  Dry  Creek,  Nevada  County;  Santa  Ana  River,  Riverside  County; 
Sisquoc  River,  Santa  Barbara  County;  and  Santa  Margarita  River,  San  Diego 
County.  Several  thousand  fingerlings  and  yearlings  were  stocked  in  these  waters. 
It  appears  that  only  the  Lake  Oroville  and  South  Fork  Stanislaus  River  popula- 
tions are  firmly  established  (Lambert  1980).  The  remainder  apparently  did  not 
survive. 

Micropterus  punctulatus  henshalli  Hubbs  and  Bailey — Alabama  spotted 
bass 

This  subspecies  is  thriving  in  Perris  Lake,  Riverside  County.  The  original  intro- 
duction consisted  of  94  2-year-olds  stocked  by  the  California  Department  of  Fish 
and  Game  in  January  1974  (Brown,  Aasen,  and  von  Geldern  1977).  An  addition- 
al 29  fish  were  taken  to  the  Department's  Central  Valleys  Hatchery  to  provide 
a  broodstock.  These  spotted  bass  were  collected  by  the  Alabama  Department 
of  Conservation  and  Natural  Resources  from  Lewis  Smith  Lake,  Alabama. 

Reproduction  of  the  bass  held  at  Central  Valleys  Hatchery  provided  fish  for 
a  second  introduction  into  Perris  Lake  in  August  1974.  In  late  1974  between  2,000 
and  3,000  fingerlings  from  this  hatchery  were  stocked  in  Millerton  Lake,  Fresno 
County.  In  early  1975,  this  plant  was  supplemented  with  150  adults  collected 
from  Perris  Lake.  Another  300  adults  and  subadults  collected  from  Perris  Lake 
in  March  and  April  1977  were  released  in  San  Vicente  Reservoir,  San  Diego 
County.  Both  the  Millerton  and  San  Vicente  populations  are  successfully  estab- 
lished. Additional  bass  from  Perris  have  since  been  stocked  in  New  Hogan 
Reservoir,  Calaveras  County;  Lake  Isabella,  Kern  County;  and  Lake  Oroville, 
Butte  County. 

Micropterus  salmoides  salmoides   (Lacepede) — northern   largemouth 
bass 


A  LIST  OF  CALIFORNIA  FISHES  19 

Micropterus  salmoides  floridanus  (Lesueur) — Florida  largemouth  bass 
The  nominate  subspecies  is  the  form  widely  distributed  throughout  the  State. 
The  Florida  largemouth  bass  was  imported  into  California  in  May  1959.  A  ship- 
ment of  about  20,400  fingerlings  from  Holt  State  Fish  Hatchery  near  Pensacola, 
Florida,  was  planted  in  upper  Otay  Reservoir,  San  Diego  County,  on  an  experi- 
mental basis  (Sasaki  1961;  Bottroff  and  Lembeck  1978).  A  self-sustaining  popula- 
tion was  soon  established  and  transplants  were  made  to  other  San  Diego  County 
reservoirs.  It  is  now  established  in  other  waters  in  the  State. 

PERCIDAE— perch  family 
Percina  macrolepida  Stevenson — bigscale  logperch 

In  our  1959  check  list  we  listed  and  described  the  introduction  of  Percina 
caprodes,  the  logperch,  into  California.  Since  then,  Stevenson  (1971 )  described 
the  bigscale  logperch  from  Texas.  Subsequent  examination  of  specimens  from 
California  revealed  them  to  be  P.  macrolepida  rather  than  P.  caprodes  (Sturgess 
1976). 

EMBIOTOCIDAE— surfperch  family 
Hysterocarpus  traskii  traskii  Gibbons — Sacramento  tule  perch 
Hysterocarpus  traskii  lagunae  Hopkirk — Clear  Lake  tule  perch 
Hysterocarpus  traskii  porno  Hopkirk — Russian  River  tule  perch 

Hopkirk  (1973:83-92)  revised  the  genus  Hysterocarpus.  He  described  the  tule 
perch  from  the  Russian  River  as  a  new  subspecies  and  remarked,  "The  subspe- 
cies porno  is  adapted  for  existence  in  small  rivers.  In  body  shape  and  in  certain 
meristic  characters,  it  represents  an  evolutionary  parallel,  not  a  relative,  of  the 
nominate  subspecies."  In  his  description  of  the  new  subspecies  of  tule  perch 
from  the  Clear  Lake  Basin  in  central  California,  Hopkirk  noted  that  it  ".  .  .  is 
adapted  for  pelagic  or  lacustrine  existence,  as  evidenced  by  the  attenuate  body, 
higher  number  of  gill  rakers,  and  silvery  coloration."  Remaining  populations  in 
the  State  are  apparently  referable  to  the  nominate  subspecies. 

CICHLIDAE— cichlid  family 
Tilapia  mossambica  Peters — Mozambique  tilapia 

The  first  breeding  population  of  this  tilapia  species  in  California  was  discov- 
ered in  1964  in  a  small  pond  and  its  tributary  near  the  Salton  Sea  in  Imperial 
County  (St.  Amant  1966) .  This  population,  which  may  no  longer  exist,  originated 
from  a  nearby  tropical  fish  farm  (Sargent's).  Subsequent  authorized  introduc- 
tions in  various  ponds  and  waterways  in  the  late  1960's  and  early  1970's  for 
mosquito  and  aquatic  weed  control,  plus  unauthorized  introductions  and  natural 
movement  of  fish  from  one  area  to  another,  have  culminated  in  the  establish- 
ment of  the  Mozambique  tilapia  in  southern  California. 

Hoover  and  St.  Amant  (1970)  observed  free-living  populations  of  T.  mossam- 
bica'm  irrigation  canals  and  drains  in  Bard  Valley,  Imperial  County,  in  1968.  They 
remain  abundant  there  as  well  as  in  similar  habitat  in  the  Palo  Verde  Valley, 
Imperial  and  Riverside  counties.  Isolated  populations  have  been  reported  from 
drains  in  the  Imperial  Valley,  Imperial  County,  and  the  Coachella  Valley,  River- 
side County.  Lake  Elsinore  in  Riverside  County  and  the  Salton  Sea  support 
abundant,  reproducing  populations.  The  identity  of  this  tilapia  from  the  Salton 


20  CALIFORNIA  FISH  AND  CAME 

Sea,  however,  remains  uncertain,  having  been  variously  identified  as  T.  mossam- 
bica  or  T.  aurea. 

In  recent  years  T.  mossambica  has  established  breeding  populations  in  a  series 
of  watercourses  entering  the  Pacific  Ocean  in  Orange  and  Los  Angeles  counties 
( Knaggs  1 977 ) .  They  are  concentrated  in  the  estuarine  portions  of  various  flood 
control  channels  and  channelized  river  beds  such  as  the  Los  Angeles,  Santa  Ana, 
and  San  Gabriel  rivers. 

Tilapia  zillii  (Gervais) — redbelly  tilapia 

The  redbelly  tilapia  was  one  of  three  tilapia  species  authorized  by  the  Fish  and 
Game  Commission  in  1971  for  use  in  California.  Its  purported  ability  to  control 
aquatic  weeds  was  responsible  for  the  interest  in  this  species.  During  the  early 
1970's,  it  was  stocked  in  several  ponds  in  central  California  and  in  numerous 
ponds,  canals,  and  drains  in  southern  California.  Except  for  the  very  southeastern 
corner  of  the  State,  it  was  believed  that  T.  z/////could  not  survive  winter  tempera- 
tures and  that  small  fish  would  have  to  be  introduced  periodically  to  achieve 
weed  control.  Howver,  until  killed  by  the  exceptionally  cold  winter  of  1972-73, 
they  overwintered  in  the  central  California  ponds.  It  was  this  unexpected  toler- 
ance to  cold  temperatures  that  prompted  the  Fish  and  Game  Commission  in 
1974  to  place  the  redbelly  tilapia  on  the  prohibited  species  list  for  that  portion 
of  the  State  north  of  the  Tehachapi  Mountains. 

Stocking  in  southern  California,  on  the  other  hand,  led  to  the  permanent 
establishment  of  T.  zillii  and  the  likelihood  of  further  spread  of  this  highly 
adaptable  species.  They  are  abundant  and  breeding  in  all  drains  entering  the 
Salton  Sea  and  are  also  abundant  in  the  Sea  itself  (Black  1980).  They  aie  likely 
to  be  encountered  in  certain  canals  and  ditches  in  Bard  and  Imperial  valleys, 
Imperial  County,  and  in  the  Coachella  Valley,  Riverside  County.  Breeding  popu- 
lations have  been  discovered  in  four  backwaters  of  the  Colorado  River  down- 
stream from  the  Palo  Verde  Diversion  Dam  and  in  Lake  Cahuilla,  Riverside 
County.  Two  specimens  have  been  reported  from  the  marine  environment  near 
Huntington  Beach  and  in  Newport  Bay,  Orange  County  (Knaggs  1977). 

GOBIIDAE— goby  family 
Acanthogobius  flavimanus  (Temminck  and  Schlegel) — yellowfin  goby 

This  species  was  first  collected  by  personnel  of  the  California  Department  of 
Fish  and  Game  in  the  San  Joaquin  River  off  Prisoners  Point  on  18  January  1963 
(Brittan,  Albrecht,  and  Hopkirk  1963).  It  soon  spread  rapidly  (Brittan  etal.  1970) 
and  is  now  widely  established  in  the  Sacramento-San  Joaquin  Delta,  the  San 
Francisco  Bay  area,  and  various  coastal  lagoons.  The  origin  of  these  fish  is  not 
known;  they  may  have  been  carried  in  a  ship's  seawater  system. 

Tridentiger  trigonocephalus  (Gill) — chameleon  goby 

Miller  and  Lea  (1972),  who  list  this  species  as  occurring  in  the  shallows  of 
both  Los  Angeles  Harbor  and  San  Francisco  Bay,  state  that  it  was  inadvertently 
introduced  from  the  Orient.  Moyle  (1976:344)  remarks  that  it,  ".  .  .  has  not  yet 
been  collected  in  fresh  water  in  California  but  can  be  expected  there,  since  it 
occurs  in  brackish  Lake  Merritt  in  Oakland  and  in  the  lower  reaches  of  streams 
in  its  native  Asia."  Hubbsand  Miller  (1965:44),  however,  refer  to  data  indicating 
that  Lake  Merritt  is  a  freshwater  lake,  although  it  connects  directly  with  San 
Francisco  Bay. 


A  LIST  OF  CALIFORNIA  FISHES  21 

Forms  and  Names  Removed  from  the  Main  List  Since  1959 

PETROMYZONTI DAE— lamprey  family 
Lampetra  planeri  (Bloch) — brook  lamprey 

Several  different  species  of  "brook  lampreys"  in  California  have  been  listed 
or  identified  as  Lampetra  planeri  and  we  included  this  species  in  our  1959  list. 
It  should  be  removed  from  California  faunal  lists  as  it  is  a  European  form  not 
found  in  North  America  (W.  I.  Follett,  pers.  commun.). 

OSMERI DAE— smelt  family 
Hypomesus  olidus  (Pallas) — pond  smelt 

The  fish  we  listed  in  our  1959  check  list  under  this  name  has  since  been 
described  as  a  new  species,  H.  transpacificus,  by  McAllister  (1963). 

SALMONIDAE — salmon  and  trout  family 
Salmo  clarkii  evermanni  Jordan  and  Grinnell — San  Gorgonio  cutthroat 

trout 

After  finding  a  record  that  cutthroat  trout  from  Lake  Tahoe  had  been  planted 
in  the  southern  California  stream  from  which  Salmo  evermanni  was  later  ob- 
tained, Benson  and  Behnke  (1961)  closely  compared  the  "type"  and  two 
"cotypes"  of  evermanni  with  specimens  of  Salmo  clarkii  henshawi  from  Lake 
Tahoe.  They  found  no  significant  differences  and  concluded  that  evermanniwas 
a  synonym. 

Salmo  gairdnerii  regalis  Snyder — royal  silver  rainbow  trout 

La  Rivers  (1962)  questioned  the  taxonomic  validity  of  both  5.  g.  regalis  of  Lake 
Tahoe  and  S.  g.  smaragdus  of  Pyramid  Lake.  He  argues  convincingly  against  the 
acceptance  of  these  rainbow  subspecies  as  Great  Basin  endemics,  believing  that 
the  specimens  examined  by  Snyder  (1914,  1918)  were  probably  either  intro- 
duced rainbow  or  rainbow-cutthroat  hybrids.  Widespread  stocking  of  rainbow 
trout  in  the  Lahontan  system  beginning  in  the  1860's  was  likely  the  original 
source  of  these  specimens. 

One  of  us  (Cordone)  collected  226  rainbow  trout  from  the  limnetic  zone  of 
Lake  Tahoe  in  the  early  1960's.  Seventy-three  of  these  were  marked  fish,  survi- 
vors from  plants  of  hatchery-reared  rainbow.  Many  of  these  specimens,  both 
marked  and  unmarked,  possessed  the  phenotypic  appearance  of  the  royal  silver 
trout  noted  by  Snyder  (1918),  "It  is  distinguished  by  the  absence  of  spots,  by 
the  blue  or  green  dorsal  surface,  the  silvery  sides  and  white  belly,  and  the  loose 
scales  which,  when  the  fish  is  caught,  adhere  to  the  fingers  like  bits  of  foil." 
Behnke  ( 1 972 )  examined  some  of  these  specimens  and  concluded,  "The  silvery, 
smoltlike  appearance,  supposedly  diagnostic  for  S.  regalis  can  be  duplicated  by 
hatchery  rainbow  trout  after  a  period  of  life  in  Lake  Tahoe." 

CYPRINIDAE — carp  or  minnow  family 
Pimephales  promelas  confertus  (Girard) — southwestern  fathead  min- 
now 

We  follow  Taylor  (1954:42)  and  Vandermeer  (1966:465)  in  not  recognizing 
subspecies  in  Pimephales  promelas,  primarily  because  most  of  the  variation  over 

3—81475 


22  CALIFORNIA  FISH  AND  GAME 

its  range  appears  to  be  clinal.  Even  if  subspecies  were  recognized,  the  popula- 
tions of  the  fathead  minnow  in  California  are  from  such  diverse  out-of-state 
localities  that  it  would  be  difficult  to  single  out  subspecies. 

Plagopterus  argentissimus  Cope — woundfin 

Inclusion  of  this  spiny-rayed  cyprinid  in  our  previous  check  lists  was  based  on 
its  occurrence  in  the  Gila  River  to  its  mouth  at  Yuma,  just  across  the  Colorado 
River  from  California  (Gilbert  and  Scofield  1898).  It  has  now,  however,  been 
removed  from  the  present  list  because  it  has  not  been  taken  even  in  the  lower 
Gila  River  since  1894  (Miller  and  Lowe  1964),  is  known  today  only  from  the 
Virgin  River  system  (Miller  and  Hubbs  1960;  Minckley  1973:115),  and  there  are 
no  records  of  its  actual  occurence  in  California.  It  may  be  noted,  however,  that 
Miller  and  Lowe  (1964)  state  that  it  has  been  used  as  a  baitfish  on  the  "lower 
Colorado  River". 

Rhinichthys  osculus  carringtonii  (Cope) — Pacific  speckled  dace 

W.  I.  Follett  (pers.  commun.)  states:  "We  are  not  recognizing  Rhinichthys 
osculus  carringtonii  (originally  described  from  Warm  Springs,  Box  Elder  County, 
Utah)  as  occurring  in  California.  Dr.  Hubbs  now  regards  as  a  misidentification 
Agosia  nubila  carringtonii  Culver  and  Hubbs,  1917,  Lorquinia,  1(2):83,  from 
Santa  Ana  River,  California."  On  this  basis  we  are  dropping  this  form  from  our 
list. 

Siphateles  bicolor  formosus  (Girard) — Sacramento  tui  chub 

If  this  were  a  valid  subspecies,  its  current  name  would  be  Gila  bicolor  formosa. 
Moyle  (1976:164)  comments  on  it  as  follows:  "The  name  G  b.  formosa  was 
originally  applied  to  tui  chubs  that  were  supposed  to  have  lived  in  the  Sacra- 
mento-San Joaquin  Valley.  Since  only  a  few  poorly  preserved  specimens  of  the 
form  are  known,  the  subspecies  may  be  based  on  a  mislabeled  collection  (C. 
L.  Hubbs,  pers.  commun.)."  For  these  reasons,  we  are  dropping  this  form  from 
the  main  list. 

CATOSTOM I  DAE— sucker  family 
Catostomus  latipinnis  Baird  and  Girard — flannel  mouth  sucker 

This  species,  native  to  the  Colorado  River  system,  is  now  found  only  in  Salt 
River  Canyon,  the  Virgin  River,  and  the  mainstem  Colorado  River  upstream  from 
Lake  Mead  (Minckley  1973:157).  Like  Plagopterus  argentissimus,  it  may  never 
have  occurred  in  the  California  portion  of  the  Colorado  River  except  for  an 
occasional  specimen  washed  down  from  upstream  waters. 

Ictiobus  cyprinella  (Valenciennes) — bigmouth  buffalo 

This  exotic  species  was  included  in  the  first  two  lists  on  the  basis  of  its 
occurrence  in  several  reservoirs  of  the  Los  Angeles  Aqueduct  system  in  Los 
Angeles  and  Inyo  counties  following  its  illegal  introduction  in  the  1940's,  presum- 
ably by  commercial  fishermen  ( Evans  1 950) .  However,  none  has  been  collected 
from  these  waters  since  the  late  1960's  and  they  probably  no  longer  exist  in  the 
State  (F.  G.  Hoover,  pers.  commun.).  Since  this  species,  along  with  the  black 
buffalo,  Ictiobus  niger,  and  the  smallmouth  buffalo,  Ictiobus  bubalus,  are  present 


A  LIST  OF  CALIFORNIA  FISHES  23 

in  Arizona  waters,  they  may  be  expected  on  occasion  to  find  their  way  into  the 
lower  Colorado  River  and  connected  waters.  On  the  basis  of  a  photograph,  C. 
L  Hubbs  and  J.  A.  St.  Amant  identified  a  specimen  collected  from  a  waterway 
in  southern  California  in  1969  as  /.  bubalus. 

Pantosteus  lahontan  Rutter — Lahontan  mountain-sucker 

Smith  (1966)  united  Pantosteus  lahontan  and  P.  platyrhynchus  as  Catostomus 
platyrhynchus,  which  replaces  P.  lahontan  in  our  present  list. 

ICTALURIDAE — North  American  freshwater  catfish  family 
Ictalurus  me/as  me/as  (Rafinesque) — northern  black  bullhead 
Ictalurus  natalis  natalis  (Lesueur) — northern  yellow  bullhead 
Ictalurus  nebulosus  nebulosus  (Lesueur) — northern  brown  bullhead 

We  follow  Hubbs  etal.  (1979)  and  Bailey  (1956:328-329;  pers.  commun.)  in 
dropping  recognition  of  these  trinomials.  They  probably  represent  only  clinal 
variations. 

CENTRARCHIDAE— sunfish  family 
Micropterus  dolomieu  dolomieu  Lacepede — northern  smallmouth  bass 
We  follow  Hubbs  etal.  (1979)  and  Bailey  (1956:328-329;  pers.  commun.)  in 
dropping  recognition  of  this  trinomial.  It  probably  represents  only  clinal  varia- 
tion. 

ELEOTRIDIDAE— sleeper  family 
Eleotris  picta  Kner  and  Steindachner — spotted  sleeper 

This  species  was  added  to  the  1 959  list  on  the  basis  of  a  single  specimen  caught 
by  a  fisherman  at  the  canal  spillway  between  Winterhaven  and  the  Colorado 
River  in  Imperial  County  on  16  April  1952  (Hubbs  1953).  However,  none  has 
been  taken  from  California  waters  since  that  time  (Minckley  1973:259;  Moyle 
1976:70). 

REVISED  MAIN  LIST 
Native  Species  and  Established  Exotic  Species 

This  revised  list  consists  of  124  full  species,  which  may  be  subdivided  as 
follows:  66  native  freshwater  and  anadromous  species  (including  6  which  are 
probably  extinct),  13  native  euryhaline  or  marine  species  which  occasionally 
penetrate  into  fresh  water,  and  45  introduced  species.  The  124  species  comprise 
25  families  and  64  genera. 

Species  which  have  been  introduced  into  California  waters  are  denoted  by  an 
asterisk  ( * ) ,  marine  or  euryhaline  fishes  which  occur  occasionally  in  fresh  water 
by  an  "O",  and  extinct  species  by  a  dagger  (f). 

PETROMYZONTIDAE — lamprey  family 

1.  Lampetra  ayresii  (Giinther) — river  lamprey 

2.  Lampetra  folletti  (Vladykov  and  Kott) — Modoc  brook  lamprey 

3.  Lampetra  hubbsi  (Vladykov  and  Kott) — Kern  brook  lamprey 

4.  Lampetra  lethophaga  Hubbs — Pit-Klamath  brook  lamprey 

5.  Lampetra  pacifica  Vladykov — Pacific  brook  lamprey 

6.  Lampetra  richardsoni  Vladykov  and  Follett — western  brook  lamprey 

7.  Lampetra  tridentata  (Gairdner) — Pacific  lamprey 


24  CALIFORNIA  FISH  AND  CAME 

ACIPENSERIDAE— sturgeon  family 

8.  Acipenser  medirosths  Ayres — -green  sturgeon 

8a.     Acipenser  medirostris  medirosths  Ayres — American  green  sturgeon 

9.  Acipenser  transmontanus  Richardson — white  sturgeon 

ELOPIDAE — tenpounder  family 

10.  Elops  affinis  Regan — machete  O 

CLUPEIDAE — herring  family 

11.  Alosa  sapidissima  (Wilson) — American  shad  * 

12.  Clupea  harengus  Linnaeus — herring  O 

12a.  Clupea  harengus  pallasii 'Valenciennes — Pacific  herring  O 

13.  Dorosoma  petenense  (Gunther) — threadfin  shad  * 

OSMERIDAE— smelt  family 

14.  Hypomesus  nipponensis  McAllister — freshwater  smelt  * 

15.  Hypomesus  pretiosus  (Cirard) — surf  smelt  O 

16.  Hypomesus  transpacificus  McAllister — delta  smelt 

17.  Spirinchus  thaleichthys  (Ayres) — longfin  smelt  O 

1 8.  Thaleichthys  pacificus  ( Richardson )  — eulachon 

COREGONIDAE— whitefish  family 

19.  Prosopium  williamsoni  (Cirard) — mountain  whitefish 

SALMONIDAE — salmon  and  trout  family 

20.  Oncorhynchus  gorbuscha  (Walbaum) — pink  salmon 

21.  Oncorhynchus  keta  (Walbaum) — chum  salmon 

22.  Oncorhynchus  kisutch  (Walbaum) — coho  salmon  (silver  salmon) 

23.  Oncorhynchus  nerka   (Walbaum) — sockeye  salmon   (anadromous  form);  kokanee  salmon 
(freshwater  form  *) 

24.  Oncorhynchus  tshawytscha  (Walbaum) — chinook  salmon  (king  salmon) 

25.  Salmo  aguabonita  Jordan — golden  trout 

25a.  Salmo  aguabonita  aguabonita  Jordan — South  Fork  Kern  golden  trout 
25b.  Salmo  aguabonita  whitei  Evermann — Little  Kern  golden  trout 

26.  Salmo  clarkii  Richardson — cutthroat  trout 

26a.  Salmo  clarkii  clarkii  Richardson — coast  cutthroat  trout 
26b.  Salmo  clarkii  henshawi  G\\\  and  Jordan — Lahontan  cutthroat  trout 
26c.  Salmo  clarkii  pleuriticus  Cope — Colorado  River  cutthroat  trout 
26d.  Salmo  clarkii  se/en iris  Snyder — Paiute  cutthroat  trout 

27.  Salmo  gairdnerii  Richardson — rainbow  trout 

27a.  Salmo  gairdnerii  gairdnerii  Richardson — steelhead  rainbow  trout 
27b.  Salmo  gairdnerii  aquilarum  Snyder — Eagle  Lake  rainbow  trout 
27c.  Salmo  gairdnerii  gilberti  Jordan — Kern  River  rainbow  trout 
27d.  Salmo  gairdnerii  kam/oops  (Jordan) — Kamloops  rainbow  trout  * 
27e.  Salmo  gairdnerii  stonei  Jordan — Shasta  rainbow  trout 

28.  Salmo  trutta  Linnaeus — brown  trout  * 

29.  Salvelinus  confluentus  (Suckley) — bull  trout  t 

30.  Salvelinus  fontinalis  (Mitchill) — brook  trout  * 

31.  Salvelinus  malma  (Walbaum) — Dolly  Varden  ] 

32.  Salvelinus  namaycush  (Walbaum) — lake  trout  * 

32a.  Salvelinus  namaycush  namaycush  (Walbaum) — common  lake  trout  * 

CYPRINIDAE — carp  or  minnow  family 

33.  Carassius  auratus  (Linnaeus) — goldfish  * 

34.  Cyprinus  carpio  Linnaeus — common  carp  * 

35.  Gila  bicolor  (Girard) — tui  chub 

35a.  Cila  bicolor  bicolor  (Cirard) — Klamath  tui  chub 

35b.  Cila  bicolor  mohavensis  (Snyder) — Mohave  tui  chub 

35c.  Cila  bicolor  obesa  (Cirard) — Lahontan  coarseraker  tui  chub 

35d.  Gila  bicolor pectinifer  (Snyder) — Lahontan  fineraker  tui  chub 

35e.  Gila  bicolor  snyderi  Miller— Owens  tui  chub 

35f.  Gila  bicolor  thalassina  (Cope) — Goose  Lake  tui  chub 

35g.  Gila  bicolor  vaccaceps  Bills  and  Bond — Cowhead  Lake  tui  chub 


A  LIST  OF  CALIFORNIA  FISHES  25 

36.  Cila  coerulea  (Girard) — blue  chub 

37.  Gila  crassicauda  (Baird  and  Girard) — thicktail  chub  t 

38.  Gila  elegans  Baird  and  Girard — bonytail  chub  t 

39.  Cila  orcuttii  (Eigenmann  and  Eigenmann) — arroyo  chub 

40.  Hesperoleucus  symmetricus  (Baird  and  Girard) — western  roach 

40a.  Hesperoleucus  symmetricus  symmetricus   (Baird  and  Girard) — Sacramento  western 

roach 
40b.  Hesperoleucus  symmetricus  mitru/us  Snyder — upper  Pit  western  roach 
40c.  Hesperoleucus  symmetricus  navarroensis  Snyder — Navarro  western  roach 
40d.  Hesperoleucus  symmetricus  parvipinnis  Snyder — Gualala  western  roach 
40e.  Hesperoleucus  symmetricus  subditus  Snyder — Monterey  western  roach 
40f.  Hesperoleucus  symmetricus  venustus  Snyder — Venus  western  roach 

41.  Lavinia  exilicauda  Baird  and  Girard — hitch 

41a.  Lavinia  exilicauda  exilicauda  Baird  and  Girard — Sacramento  hitch 
41b.  Lavinia  exilicauda  chi  Hopkirk — Clear  Lake  hitch 
41c.  Lavinia  exilicauda  harengus  Girard — Monterey  hitch 

42.  Mylopharodon  conocephalus  (Baird  and  Girard) — hardhead 

43.  Notemigonus  crysoleucas  (Mitchill) — golden  shiner  * 

44.  Notropis  lutrensis  (Baird  and  Girard) — red  shiner  * 

45.  Orthodon  microlepidotus  (Ayres)— Sacramento  blackfish 

46.  Pimephales  promelas  Rafinesque — fathead  minnow  * 

47.  Pogonichthys  ciscoides  Hopkirk — Clear  Lake  splittail  t 

48.  Pogonichthys  macrolepidotus  (Ayres) — Sacramento  splittail 

49.  Ptychocheilus  grandis  (Ayres) — Sacramento  squawfish 

50.  Ptychocheilus  lucius  Girard — Colorado  squawfish  f 

51.  Rhinichthys  osculus  (Girard) — speckled  dace 

51a.  Rhinichthys  osculus  klamathensis  (Evermann  and  Meek)— Klamath  speckled  dace 
51b.  Rhinichthys  osculus  nevadensis  Gilbert— Amargosa  speckled  dace 
51c.  Rhinichthys  osculus  robustus  (Rutter)— Lahontan  speckled  dace 

52.  Richardsonius  egregius  (Girard) — Lahontan  redside 

53.  Tinea  tinea  (Linnaeus) — tench  * 

CATOSTOMIDAE— sucker  family 

54.  Catostomus  fumeiventris  Miller — Owens  sucker 

55.  Catostomus  luxatus  (Cope) — Lost  River  sucker 

56.  Catostomus  microps  Rutter — Modoc  sucker 

57.  Catostomus  occidentalis  Ayres — western  sucker 

57a.  Catostomus  occidentalis  occidentalis  Ayres — Sacramento  western  sucker 
57b.  Catostomus  occidentalis  humboldtianus  Snyder — Humboldt  western  sucker 
57c.  Catostomus  occidentalis  lacusanserinus  Fowler — Goose  Lake  western  sucker 
57d.  Catostomus  occidentalis  mniotiltus  Snyder— Monterey  western  sucker 

58.  Catostomus  p/atyrhynchus  (Cope) — mountain  sucker 

59.  Catostomus  rimicu/us  Gilbert  and  Snyder — Klamath  smallscale  sucker 

60.  Catostomus  santaanae  (Snyder) — Santa  Ana  sucker 

61.  Catostomus  snyderi  CWben — Klamath  largescale  sucker 

62.  Catostomus  tahoensis  Gill  and  Jordan — Tahoe  sucker 

63.  Chasmistes  brevirostris  Cope — shortnose  sucker 

64.  Xyrauchen  texanus  (Abbott) — humpback  sucker 

COBITIDIDAE— loach  family 

65.  Misgurnus  anguillicaudatus  (Cantor) — Oriental  weatherfish  * 

ICTALURIDAE — North  American  freshwater  catfish  family 

66.  Icta/urus  catus  (Linnaeus) — white  catfish  * 

67.  Icta/urus  furcatus  (Lesueur) — blue  catfish  * 

68.  Icta/urus  melas  (Rafinesque) — black  bullhead  * 

69.  Icta/urus  nata/is  (Lesueur) — yellow  bullhead  * 

70.  Icta/urus  nebulosus  (Lesueur) — brown  bullhead  * 

71.  Icta/urus  punctatus  (Rafinesque) — channel  catfish  * 

72.  Py/odictis  o/ivaris  (Rafinesque) — flathead  catfish  * 


26  CALIFORNIA  FISH  AND  CAME 

CYPRINODONTIDAE— killifish  family 

73.  Cyprinodon  macu/arius  Baird  and  Cirard — desert  pupfish 

74.  Cyprinodon  milleri  LaBounty  and  Deacon — Cottonball  Marsh  pupfish 

75.  Cyprinodon  nevadensis  Eigenmann  and  Eigenmann — Nevada  pupfish 

75a.  Cyprinodon  nevadensis  nevadensis  Eigenmann  and  Eigenmann — Saratoga  Nevada  pupfish 
75b.  Cyprinodon  nevadensis  amargosae  Miller — Amargosa  Nevada  pupfish 
75c.  Cyprinodon  nevadensis  calidae  Miller — Tecopa  Nevada  pupfish  f 
75d.   Cyprinodon  nevadensis  shoshone  Miller — Shoshone  Nevada  pupfish  f 

76.  Cyprinodon  radiosus  Miller — Owens  pupfish 

77.  Cyprinodon  salinus  Miller — Salt  Creek  pupfish 

78.  Fundulus  parvipinnis  Girard — California  killifish 

78a.  Fundulus  parvipinnis  parvipinnis — southern  California  killifish 

79.  Lucania  parva  (Baird  and  Girard) — rainwater  killifish  * 

POECILIIDAE — livebearer  family 

80.  Cambusia  affinis  (Baird  and  Girard) — mosquitofish  * 

80a.  Cambusia  affinis  affinis  (Baird  and  Girard) — western  mosquitofish  * 

81.  Poecilia  latipinna  (Lesueur) — sailfin  molly  * 

82.  Poecilia  mexicana  Steindachner — shortfin  molly  * 

82a.  Poecilia  mexicana  mexicana  Steindachner — Orizaba  shortfin  molly  * 

83.  Poeciliopsis  gracilis  (Heckel) — porthole  livebearer* 

ATHERINIDAE— silverside  family 

84.  Atherinops  affinis  (Ay res) — topsmelt  O 

85.  Menidia  audens  Hay — Mississippi  silverside  * 

GASTEROSTEIDAE— stickleback  family 

86.  Casterosteus  acu/eatus  Linnaeus — threespine  stickleback 

86a.  Casterosteus  acu/eatus  aculeatus  Linnaeus — armored  threespine  stickleback 

86b.  Casterosteus  acu/eatus  microcephalus  Girard — semiarmored  threespine  stickleback 

86c.  Casterosteus  aculeatus  williamsoni  Girard — unarmored  threespine  stickleback 

SYNGNATHIDAE— pipefish  family 

87.  Syngnathus  leptorhynchus  Girard — bay  pipefish  O 

COTTIDAE— sculpin  family 

88.  Clinocottus  acuticeps  (Gilbert) — sharpnose  sculpin  O 

89.  Cottus  aleuticus  Gilbert — coastrange  sculpin 

90.  Cottus  asper  Richardson — prickly  sculpin 

91 .  Cottus  asperrimus  Rutter — rough  sculpin 

92.  Cottus  beldingii  Eigenmann  and  Eigenmann — Paiute  sculpin 

93.  Cottus  gulosus  (Girard) — riffle  sculpin 

94.  Cottus  klamathensis  Gilbert — marbled  sculpin 

95.  Cottus  perplexus  Gilbert  and  Evermann — reticulate  sculpin 

96.  Cottus  pitensis  Bailey  and  Bond — Pit  sculpin 

97.  Leptocottus  armatus  Girard — Pacific  staghorn  sculpin  O 

97a.  Leptocottus  armatus  armatus  Girard — northern  Pacific  staghorn  sculpin  O 
97b.  Leptocottus  armatus  australis  Hubbs — southern  Pacific  staghorn  sculpin  O 

PERCICHTHYIDAE— temperate  bass  family 

98.  Morone  chrysops  (Rafinesque) — white  bass  * 

99.  Morone  saxatilis  (Walbaum) — striped  bass* 

CENTRARCHIDAE— sunfish  family 

100.  Archoplites  interruptus  (Girard) — Sacramento  perch 

101.  Lepomis  cyanellus  Rafinesque — green  sunfish  * 

102.  Lepomis  gibbosus  (Linnaeus) — pumpkinseed  * 

103.  Lepomis  gulosus  (Cuvier) — warmouth  * 

104.  Lepomis  macrochirus  Rafinesque — bluegill  * 

104a.  Lepomis  macrochirus  macrochirus  Rafinesque — northern  bluegill  * 
104b.  Lepomis  macrochirus  purpurescens  Cope — southeastern  bluegill  * 

105.  Lepomis  microlophus  (Gunther) — redear  sunfish  * 

106.  Micropterus  coosae  Hubbs  and  Bailey — redeye  bass  * 


A  LIST  OF  CALIFORNIA  FISHES  27 

107.  Micropterus  dolomieu  Lacepede — smallmouth  bass  * 

108.  Micropterus  punctulatus  (Rafinesque) — spotted  bass* 

108a.  Micropterus  punctulatus  punctulatus  fRafinesque) — northern  spotted  bass* 
108b.  Micropterus  punctulatus  henshalli  Hubbs  and  Bailey — Alabama  spotted  bass  * 

109.  Micropterus  salmoides  (Lacepede) — largemouth  bass  * 

109a.  Micropterus  salmoides  salmoides  (Lacepede) — northern  largemouth  bass* 
109b.  Micropterus  salmoides  floridanus  (Lesueur) — Florida  largemouth  bass  * 

110.  Pomoxis  annularis  Rafinesque — white  crappie  * 

111.  Pomoxis  nigromaculatus  (Lesueur) — black  crappie  * 

PERCIDAE — perch  family 

112.  Perca  flavescens  (Mitchill) — yellow  perch  * 

113.  Percina  macrolepida  Stevenson — bigscale  logperch  * 

EMBIOTOCIDAE — surfperch  family 

114.  Cymatogaster  aggregata  Gibbons — shiner  perch  O 

115.  Hysterocarpus  traskii  Gibbons — tule  perch 

1 1 5a.  Hysterocarpus  traskii  traskii  Gibbons — Sacramento  tule  perch 
115b.  Hysterocarpus  traskii  lagunae  Hopkirk — Clear  Lake  tule  perch 
115c.  Hysterocarpus  traskii  porno  Hopkirk — Russian  River  tule  perch 

CICHLIDAE — cichlid  family 

116.  Tilapia  mossambica  (Peters) — Mozambique  tilapia  * 

117.  Tilapia  zillii  (Gervais)—  redbelly  tilapia  * 

MUGILIDAE — gray  mullet  family 

118.  Mugil  cephalus  Linnaeus — striped  mullet  O 

GOBIIDAE — goby  family 

119.  Acanthogobius  flavimanus  (Temminck  and  Schlegel) — yellowfin  goby  * 

120.  Clevelandia  ios  (Jordan  and  Gilbert) — arrow  goby  O 

121.  Eucyclogobius  newberryi  (Girard) — tidewater  goby 

122.  Cillichthys  mirabilis  Cooper — longjaw  mudsucker  O 

123.  Tridentiger  trigonocephaly  (Gill) — chameleon  goby  O  * 

PLEURONECTIDAE — righteye  flounder  family 

124.  Platichthys  stellatus  (Pallas)— starry  flounder  O 

124a.  Platichthys  stellatus  rugosus  Girard — southern  starry  flounder  O 

REVISED  SUPPLEMENTARY  LISTS 
Native  Species — Extinct  in  California 

We  have  included  in  this  section  only  those  native  species  which,  at  least 
according  to  the  literature,  at  one  time  were  well  established.  Not  included  are 
the  woundfin,  Plagopterus  argentissimus,  and  the  flannelmouth  sucker,  Catos- 
tomus  latipinnis,  which  rarely,  if  ever,  entered  California  waters.  To  avoid  confu- 
sion, we  have  also  omitted,  both  from  this  and  the  main  list,  the  Clear  Lake 
minnow  which  was  described  by  Hopkirk  (1973:57-59)  as  Endemichthys  gran- 
dipinnis,  from  specimens  last  collected  in  1 939  and  1 940.  He  observed  that  it  was 
apparently  extinct.  He  is  now  reconsidering  its  generic  allocation  (J.  D.  Hopkirk, 
pers.  commun.). 

Excluding  the  above,  we  believe  that  the  following  eight  native  fishes  no 
longer  exist  in  California. 

SALMONIDAE — salmon  and  trout  family 
Salvelinus  confluentus  (Suckley) — bull  trout 
Salvelinus  ma/ma  (Walbaum) — Dolly  Varden 

These  species  (there  is  some  question  that  at  one  time  both  existed  in  the 
McCloud  River)  have  likely  become  extinct  in  California  as  a  result  of  man-made 


28  CALIFORNIA  FISH  AND  GAME 

environmental  changes  and  the  introduction  of  exotic  trout  into  the  McCloud 
River  drainage.  The  last  known  specimens,  probably  bull  trout,  were  taken  in 
1975  (Movie  1976:146).  Intensive  sampling  of  the  McCloud  River  and  its  tribu- 
taries in  recent  years  has  failed  to  locate  either  species  (S.  J.  Nicola,  pers. 
commun). 

CYPRINIDAE — carp  or  minnow  family 
Gila  crassicauda  (Baird  and  Girard) — thicktail  chub 

This  chub  was  once  common  in  the  Central  Valley,  Clear  Lake  in  Lake  County, 
and  at  least  one  tributary  to  south  San  Francisco  Bay.  A  combination  of  man- 
caused  habitat  changes  and  the  introduction  of  exotic  fishes  has  led  to  its 
apparent  extinction  (Miller  1963).  The  last  known  specimen  was  taken  in  1957 
from  Steamboat  Slough  in  the  Sacramento  River  Delta  (Calif.  Dep.  Fish  and 
Game  1978).  A  report  to  Moyle  (1976:172)  that  a  specimen  was  collected  from 
Cache  Slough,  near  Rio  Vista,  in  1 958  was  in  error  ( P.  B.  Moyle,  pers.  commun. ) . 

Gila  elegans  Baird  and  Girard — bonytail  chub 

This  species,  listed  in  our  1959  list  as  Gila  robusta  elegans,  Colorado  River 
bonytail  chub,  has  not  been  found  in  the  California  portion  of  the  Colorado  River 
in  recent  years  and  may  be  considered  extinct  in  the  State  (Colorado  River 
Wildlife  Council  1977;  Calif.  Dep.  Fish  and  Game  1978). 

Pogonichthys  ciscoides  Hopkirk — Clear  Lake  splittail 

It  was  not  until  Hopkirk  (1973)  published  the  results  of  his  studies  that  the 
Clear  Lake  splittail  was  recognized  as  a  distinct  species.  By  this  time  it  was 
probably  already  extinct,  since  none  had  been  collected  since  the  late  1960's. 
Cook,  Moore,  and  Conners  (1966)  described  the  early  history  of  the  species. 
It  was  very  abundant  until  the  early  1940's,  when  it  declined  drastically,  and 
occasional  resurgences  did  nothing  to  halt  the  overall  decline.  Habitat  destruc- 
tion and  exotic  fishes  are  believed  responsible  for  its  extinction. 

Ptychocheilus  lucius  Girard — Colorado  squawfish 

Although  still  present  in  a  few  localities  in  the  upper  Colorado  River  drainage, 
the  Colorado  squawfish  apparently  has  become  extinct  in  California  waters. 
Once  abundant  in  the  lower  Colorado  River,  it  was  probably  already  extinct  in 
this  area  by  the  early  1960's  (Moyle  1976:195).  It  has  not  been  collected  there 
since  1952  (Calif.  Dep.  Fish  and  Game  1978).  Environmental  degradation  and 
exotic  fishes  are  again  believed  responsible  for  the  loss. 

CYPRINODONTIDAE— killifish  family 
Cyprinodon  nevadensis  calidae  Miller — Tecopa  Nevada  pupfish 

This  subspecies,  originally  from  north  and  south  Tecopa  Hot  Springs,  Inyo 
County,  has  become  extinct  in  recent  years  (Moyle  1976:256)  as  a  result  of 
activities  by  man  which  led  to  destruction  of  its  habitat. 

Cyprinodon  nevadensis  shoshone  Miller — Shoshone  Nevada  pupfish 

This  subspecies,  from  Shoshone  Springs,  Inyo  County,  like  C.n.  calidae,  has 
also  become  extinct  in  recent  years  (Moyle  1976:256)  as  a  result  of  activities 
by  man  leading  to  destruction  of  its  habitat. 


A  LIST  OF  CALIFORNIA  FISHES  29 

Exotic  Species — Unsuccessfully  Introduced  or  of  Uncertain  Status 

It  is  extremely  difficult  to  establish  rigid  criteria  for  the  inclusion  or  exclusion 
of  fishes  in  the  list  that  follows.  Some  situations  are  obvious.  For  example,  we 
have  included  a  species  in  this  list  whenever  it  was  introduced  as  part  of  a 
planned  program  or  was  known  to  have  had  a  large  escapement  of  the  species, 
say  from  a  tropical  fish  farm,  even  if  subsequent  investigations  have  failed  to 
locate  it.  On  the  other  hand,  if  only  a  single  specimen  or  a  very  few  specimens, 
even  if  positively  identified,  were  recorded,  we  have  omitted  such  species  from 
the  main  list  but  have  tried  to  mention  them  below.  Obviously,  these  are  judg- 
mental assessments. 

The  occurrence  of  a  single  or  a  few  specimens  of  tropical  or  other  ornamental 
fishes  probably  represents  releases  by  home  aquarists.  Brittan  and  Grossman 
(1979)  describe  a  specimen  of  pacu,  Colossoma  sp.,  native  to  South  America, 
caught  by  an  angler  in  1977  from  the  Sacramento  River  in  Yolo  County.  Another 
pacu  was  reportedly  taken  from  the  California  Aqueduct  in  1979  (Calif.  Dep.  Fish 
and  Game,  Region  5  monthly  report  for  November  1979).  Minckley  (1973:185) 
refers  to  a  specimen  of  walking  catfish,  Clarias  batrachus,  taken  by  an  angler 
from  the  All  American  Canal  in  Imperial  County  west  of  Yuma,  Arizona.  Another 
specimen  was  taken  by  an  angler  from  Legg  Lake,  Los  Angeles  County  (J.  A.  St. 
Amant,  pers.  commun.).  A  South  American  aruana,  Osteoglossum  bicirrhosum, 
was  caught  by  an  angler  in  Lake  Berryessa  (Calif.  Dep.  Fish  and  Game,  Region 
3  news  release  for  1 8  June  1 972 ) .  Two  mature  tiger  barbs,  Barbus  tetrazona,  were 
collected  in  1963  from  the  small  stream  flowing  from  Warm  Springs  Sanctuary 
in  Owens  Valley,  Inyo  County  (Naiman  and  Pister  1974).  None  has  been  taken 
since  then,  despite  repeated  collecting  efforts. 

Escapements  and  releases  from  ornamental  fish  farms  apparently  have  been 
the  source  of  a  number  of  established  exotics,  such  as  Misgurnus  anguil- 
licaudatus,  Poecilia  /at/pinna,  P.  mexicana,  and  Poeciliopsis  gracilis.  Other  orna- 
mental species  have  escaped  but  in  small  numbers,  and  fortunately  have  not 
established  permanent  populations.  For  example,  among  the  exotics  collected 
by  St.  Amant  and  Hoover  (1969)  from  the  Westminster  flood  control  channel 
in  Orange  County  in  1968  were  the  guppy,  Lebistes  reticulatus;  green  swordtail, 
Xiphophorus  hellerii;  southern  platyfish,  X.  maculatus;  variable  platyfish,  X. 
variatus;  molly,  Poecilia  sphenops;  zebra  danio,  Brachydanio  rerio;  and  angel- 
fish,  Pterophyllum  sp.  None  of  these  has  since  been  taken  in  this  channel,  despite 
repeated  collecting  attempts.  Mearns  (1975)  took  a  specimen  of  Xiphophorus 
hellerii "in  1974  from  a  drain  to  the  Salton  Sea,  and  G.  F.  Black  (pers.  commun.) 
collected  another  from  the  same  drain  in  1979. 

The  1 959  supplementary  list  included  1 4  species  of  exotic  bait  fishes  that  were 
being  used  along  the  Colorado  River  (Miller  1952).  None  of  these  has  become 
established  in  California  and  apparently  they  are  no  longer  being  used  as  bait 
in  this  area,  so  we  have  deleted  them  from  the  list  that  follows. 

The  exotic  fishes  listed  below  fall  into  several  groups: 

1.  Fishes  known  to  have  been  introduced  but  which  have  not  survived;  e.g., 
No.  2. 

2.  Fishes  reported,  possibly  erroneously,  to  have  been  introduced,  but  which 
have  not  survived;  e.g.,  No.  9. 

3.  Fishes  which  have  been  reported  from  this  State  but  whose  identification 
is  questioned  by  the  authors;  e.g.,  No.  21. 


30  CALIFORNIA  FISH  AND  CAME 

4.  Fishes  which  have  not  been  recorded  from  the  State  for  many  years;  e.g., 
No.  24. 

As  will  be  seen  by  our  annotations,  we  know  of  no  demonstrable  evidence 
that  any  of  them  are  successfully  established  in  the  fresh  waters  of  California 
today. 

As  the  general  sources  for  the  history  and  lack  of  success  of  most  of  these 
introductions  are  fairly  well  known,  there  is  little  point  in  listing  all  the  references 
concerning  the  status  of  thest  fishes.  We  have  alluded  to  specific  literature  only 
when  our  opinion  differs  from  that  of  the  authors  cited,  or  when  such  inclusion 
serves  to  clarify  the  exact  status  of  the  species. 

ANG U I LLI DAE— freshwater  eel  family 

1.  Anguilla  rostrata  (Lesueur) — American  eel 

Introduced  in  1874,  1879,  and  1882.  There  are  no  authentic  records  of 
survival.  However,  an  occasional  eel  is  collected  from  various  waters  in  the 
State.  Skinner  ( 1 971 )  reported  the  capture  of  two  eels  from  the  Sacramento- 
San  Joaquin  Delta.  The  first,  taken  in  1964,  was  identified  by  C.  L.  Hubbs  as 
an  American  eel.  The  second,  caught  in  1969,  was  identified  as  a  European 
eel,  Anguilla  anguilla  Linnaeus,  by  W.  I.  Follett.  Skinner  suggested  that  the 
most  logical  explanation  for  the  occurrence  of  both  eels  is  that  they  were 
transported  from  abroad  in  the  ballast  of  commercial  ships.  In  1978  an 
unidentified  species  of  Anguilla  was  captured  in  the  Los  Angeles  River  (J. 
A.  St.  Amant,  pers.  commun.). 

PLECOGLOSSIDAE— ayu  family 

2.  Plecoglossus  altivelis  Temminck  and  Schlegel — ayu 

Large  numbers  of  eggs  and  fry  of  this  native  Japanese  food  and  sport 
species  were  stocked  in  California  on  the  recommendation  of  Dr.  John  W. 
DeWitt,  Professor  of  Fisheries  at  Humboldt  State  University,  Areata.  Follow- 
ing approval  from  the  Fish  and  Game  Commission,  plants  of  this  species 
were  made  annually  from  1961  through  1965.  About  3,845,000  eggs  and  fry 
were  stocked  during  this  period:  200,000  eggs  and  fry  in  Morris  Lake,  Men- 
docino County;  395,000  eggs  in  Ruth  Reservoir,  Trinity  County;  and  3,250,- 
000  eggs  and  fry  in  the  Eel  River  below  Fortuna,  Humboldt  County  (J.  W. 
DeWitt,  pers.  commun.).  No  survivors  were  reported. 

COREGONIDAE— whitefish  family 

3.  Coregonus  clupeaformis  (Mitchill) — lake  whitefish 

3a.  Coregonus  clupeaformis  clupeaformis  (Mitchill) — Great  Lakes  white- 
fish 
All  introductions  of  this  whitefish  were  made  during  the  last  century.  Even 
the  few  old  reports  of  recapture  (circa  1 880)  are  considered  highly  dubious. 

4 .  Prosopium  gemmiferum  (Snyder)  —  Bonneville  cisco 

In  January  of  1 964,  1 965,  and  1 966,  21 ,506  spawning  Bonneville  cisco  and 
about  250,000  cisco  eggs  were  collected  from  Bear  Lake,  Utah-Idaho,  and 
transported  to  LakeTahoe  (Frantz  and  Cordone  1965, 1967).  About  205,000 
green  eggs,  3,000  eyed  eggs  and  alevins,  and  15,888  ripe  adults  were 
released  in  Lake  Tahoe  over  the  3-year  span.  None  is  known  to  have 
survived. 


A  LIST  OF  CALIFORNIA  FISHES  31 

SALMON  I  DAE— salmon  and  trout  family 

5.  Salmo  clarkii  Richardson — cutthroat  trout 

5a.  Salmo  clarkii  lewisi  (Girard) — Yellowstone  cutthroat  trout 

Several  shipments  of  cutthroat  trout  eggs  have  been  brought  in  from  other 
states,  and  plants  made  in  California  waters.  It  is  probable  that  most  of  them 
were  S.  c.  lewisi.  There  are  no  records  of  survival. 

6.  Salmo  salar  Linnaeus — Atlantic  salmon   (anadromous  form);  landlocked 
Atlantic  salmon  (freshwater  form) 

Both  forms  have  been  planted  several  times.  The  old  records  of  their 
survival  may  be  dubious;  there  are  no  authentic  recent  records. 

7.  Thymallus  arcticus  (Pallas) — Arctic  grayling 

Several  early  attempts  were  made  to  introduce  this  form,  and  it  apparently 
met  with  a  brief  success  in  Yosemite  National  Park  following  plants  made 
during  the  1929-1933  period.  However,  the  last  authentic  report  of  its  sur- 
vival there  (in  Grayling  Lake)  appears  to  have  been  in  1934. 

More  recently,  the  California  Department  of  Fish  and  Game  imported 
large  numbers  of  grayling  eggs  from  Arizona  and  Wyoming.  Resultant  fry 
and  fingerlings  were  stocked  in  one  stream  and  57  high  mountain  lakes 
scattered  from  the  southern  Sierra  Nevada  into  northern  California.  Approx- 
imately 156,000  fish  were  released  during  the  period  1969  to  1975.  Good 
survival  and  growth  were  documented  at  many  of  these  waters  but  actual 
reproduction  has  not  been  confirmed. 

ESOCIDAE— pike  family 

8.  Esox  americanus  Gmelin — redfin  pickerel 

8a.  Esox  americanus  vermiculatus  Lesueur — grass  pickerel 

9.  Esox  lucius  Linnaeus — northern  pike 

£  lucius  was  supposedly  introduced  in  1891,  but  one  of  the  fish  resulting 
from  this  shipment  was  identified  in  1896  as  £  vermiculatus  (now  £  a. 
vermiculatus) .  Possibly  both  species  were  included.  There  are  no  records 
of  capture  of  either  species  after  1896. 

10.  Esox  masquinongy  Mitchill — muskellunge 

10a.  Esox  masquinongy  ohioensis  Kirtland — Ohio  muskellunge 

Introduced  into  Lake  Merced,  San  Francisco  County,  in  1893.  None  sur- 
vived. 

CHANIDAE— milkfish  family 

1 1 .  Chanos  chanos  ( Forsskal ) — milkfish 

Milkfish  from  the  Hawaiian  Islands  were  planted  in  a  stream  in  Solano 
County  in  1877.  There  are  no  records  of  their  survival  there.  The  species  is 
an  ocean  fish  which  occasionally  enters  fresh  water. 

CYPRINIDAE — carp  or  minnow  family 

1 2.  Ctenopharyngodon  idella  (Valenciennes) — grass  carp 

Illegal  introductions  of  grass  carp  into  California  have  been  made  in  the 
past  and  may  still  be  continuing.  Despite  the  fact  that  this  species  of  Chinese 
carp  is  officially  prohibited  in  the  State,  and  thus  may  not  be  imported, 


32  CALIFORNIA  FISH  AND  CAME 

transported,  or  possessed,  some  farm  pond  owners  have  been  importing 
grass  carp  from  commercial  fish  farmers  in  Arkansas  and  Pennsylvania.  The 
Department  has  thus  far  uncovered  four  instances  of  grass  carp  introduc- 
tions: 12  fingerlings  were  released  in  a  small  pond  in  Ventura  County  in  1975, 
48  fingerlings  were  planted  in  a  small  pond  in  El  Dorado  County  in  1975, 
2,800  fingerlings  and  200  0.34-kg  fish  were  released  in  seven  ponds  on  a 
ranch  in  Napa  County  in  1975,  and  20  grass  carp  fingerlings  were  stocked 
in  a  small  pond  in  Mendocino  County  in  1978.  The  latter  plant  apparently 
did  not  survive  the  trip  from  Pennsylvania,  but  the  remaining  lots  from 
Arkansas  survived  and  were  healthy  and  growing  rapidly  until  they  were 
removed  by  the  Department. 

In  May  1980  about  850  hybrids  of  female  grass  carp  and  males  of  another 
Chinese  carp,  the  bighead  carp,  Aristichthys  nobilis,  were  released  in  several 
man-made  waterways  in  the  Coachella  Valley.  Further  releases  are  an- 
ticipated as  part  of  a  study  to  assess  the  aquatic  weed  control  potential  of 
this  hybrid. 

ICTALURIDAE — North  American  freshwater  catfish  family 

1 3 .  Ictalurus  platycephalus  ( G  i  rard )  — flat  bu  1 1  head 

On  the  basis  of  a  survey  made  in  1925,  Coleman  (1930)  recorded  "The 
Great  Blue,  or  Forked-Tail  Cat — Ictalurus  furcatus,  Cuv.  and  Vincen.,"  and 
"The  Brown-Spotted  Cat — Ameirus  [sic]  platycephalus,  Girard,"  from  Clear 
Lake,  Lake  County.  Neither  has  been  recorded  from  the  Lake  since  that  time, 
despite  extensive  collecting.  We  believe  that  Coleman  confused  Ictalurus 
catus  (found  in  Clear  Lake  and  often  called  "forked-tail  catfish"  or  "blue 
cat")  with  his  "furcatus" .  We  suspect  that  his  record  of  /.  platycephalus  is 
based  upon  his  erroneous  interpretation  of  fishermen's  reports. 

ORYZI I  DAE— tooth-carp  family 

14.  Oryzias  latipes  (Temminck  and  Schlegel) — medaka 

The  statements  by  Snyder  (1935),  "It  has  been  found  in  San  Francisquito 
Creek",  and  Coates  (1942:185),  ".  .  .  this  fish  has  been  turned  loose  in 
.  .  .  parts  of  California,  where  it  is  reported  to  be  thriving",  are  the  sole 
bases  for  its  admission  to  this  list.  In  a  conversation  with  Snyder  on  21  March 
1943,  he  told  us  (Dill)  that  some  of  his  students  had  collected  this  form  in 
San  Francisquito  Creek,  Santa  Clara  County.  He  did  not  recall  the  date  or 
other  circumstances. 

CYPRINODONTIDAE— killifish  family 

15.  Cynolebias  bellottii  Steindachner — Argentine  pearlfish 

This  was  the  most  widely  used  of  the  so-called  "annual  fishes"  stocked 
in  several  locations  in  the  State,  principally  in  Butte,  Kern,  and  Riverside 
counties,  for  mosquito  control  purposes.  Bay  ( 1 966)  described  the  first  field 
tests  with  this  species  at  the  University  of  California,  Riverside.  Survivors  of 
the  tests  persisted  in  the  Riverside  ponds  for  5  years  despite  repeated  flood- 
ings  and  dryings  but  finally  died  out  (E.  F.  Legner,  Univ.  Calif.,  Riverside, 
pers.  commun.).  Additional  field  tests  with  the  Argentine  pearlfish  were 
described  by  E.  C.  Bay  (pers.  commun.).  Tests  in  experimental  ponds  were 
conducted  in  1966  and  1967  in  Kern  and  Butte  counties.  The  species  failed 


A  LIST  OF  CALIFORNIA  FISHES  33 

to  become  established. 

Experimental  rice  plots  and  ponds  on  the  grounds  of  the  Butte  County 
Mosquito  Abatement  District  were  the  sites  of  tests  conducted  in  1973  and 
1974  using  the  black  pearlfish,  Cynolebias  nigripinnis,  and  White's  pearlfish, 
Cynolebias  white/  (K.  J.  Hiscox,  Butte  County  Mosquito  Abatement  Dist., 
pers.  commun.).  The  fish  did  not  reproduce  and  the  study  was  terminated. 

POECILIIDAE— livebearer  family 
Gambusia  affinis  holbrooki  Girard — eastern  mosquitofish 

The  eastern  mosquitofish  has  been  widely  distributed  in  the  public  waters 
of  California  by  various  mosquito  abatement  districts  (E.  F.  Legner  and  K. 
J.  Hiscox,  pers.  commun.).  It  is  believed  to  be  more  tolerant  of  colder 
temperatures  than  the  western  mosquitofish.  The  two  subspecies  hybridize 
readily  and  in  California  collections  of  pure  G  a.  holbrooki  have  yet  to  be 
made  in  the  wild. 
Lebistes  reticulatus  (Peters) — guppy 

Besides  the  almost  certain  release  of  guppies  by  tropical  fish  fanciers, 
guppies  have  been  stocked  on  numerous  occasions  in  wastewater  treatment 
ponds  throughout  the  State  where  access  to  public  waters  is  possible  (K.  J. 
Hiscox,  pers.  commun. ) .  In  1 968  the  Fish  and  Game  Commission  approved 
a  request  by  the  University  of  California,  Riverside,  to  stock  guppies  in  dairy 
and  poultry  waste  lagoons  in  San  Bernardino  County  (E.  C.  Bay,  pers. 
commun.).  Also  in  1968,  the  Commission  permitted  the  Kings  Mosquito 
Abatement  District  to  release  guppies  in  lower  Mill  Creek  in  Tulare  and  Kings 
counties.  None  of  the  foregoing  introductions  led  to  the  establishment  of 
permanent  populations.  However,  wild  populations  can  be  anticipated  in 
suitable  areas  with  year-round  warmwater  temperatures. 
Rivulus  hartii  (Boulenger) — Trinidad  rivulus 

St.  Amant  (1970)  first  observed  and  collected  this  species  in  a  small  ditch 
near  a  tropical  fish  farm  in  Imperial  County  in  1967.  It  was  identified  by  C. 
L.  Hubbs.  Additional  specimens  were  collected  in  1968  and  both  adults  and 
juveniles  were  taken  in  1969.  The  population  has  since  disappeared. 
Xiphophorus  variatus  (Meek) — variable  platyfish 

St.  Amant  and  Sharp  (1971 )  collected  approximately  200  adult  and  juve- 
nile Xiphophorus  variatus,  native  to  Mexico,  from  a  drain  ditch  6.4  km  east 
of  Oasis,  Riverside  County,  on  24  December  1969.  C.  L  Hubbs  confirmed 
the  identification.  This  was  the  first  record  of  an  established  population,  but 
it  has  since  died  out. 

ATHERINIDAE— silverside  family 

Labidesthes  sicculus  (Cope) — brook  silverside 

The  brook  silverside  was  one  of  five  species  authorized  by  the  Fish  and 
Game  Commission  in  1963-64  for  introduction  into  experimental  ponds 
beside  Clear  Lake.  These  ponds,  plus  a  deep  well,  were  constructed  in  1963 
by  the  Lake  County  Mosquito  Abatement  District  "...  for  the  express 
purpose  of  evaluating  experimental  fishes  and  their  influence  on  biological 
productivity"  (Cook  1968).  The  Labidesthes,  obtained  from  Ohio,  did  well 
in  one  pond  for  3  years  and  reproduced,  but  then  died  out  from  unknown 
causes. 


34  CALIFORNIA  FISH  AND  CAME 

CENTRARCHIDAE— sunfish  family4 

21.  Ambloplites  rupestris  (Rafinesque) — rock  bass 

It  is  recorded  in  the  literature  as  having  been  introduced  in  1874  and  again 
in  1 891 ,  and  another  record  of  a  plant  of  "rock  bass"  in  1917  was  furnished 
by  E.  H.  Clidden  of  the  then  California  Division  of  Fish  and  Game.  Brief 
statements  by  Neale  (1931 )  and  Anon.  (1934)  as  to  its  limited  success  in 
California,  and  its  occasional  listing  in  State  fish  rescue  records  up  to  1939, 
are  the  only  bases  for  belief  that  this  fish  ever  endured.  The  terminology 
used  in  these  rescue  records  (published  in  the  Biennial  Reports  of  the 
California  Division  of  Fish  and  Game)  has  often  been  inexact.  We  have 
been  unable  to  find  a  single  verifiable  record  of  the  occurrence  of  the  rock 
bass  in  California. 

22.  Enneacanthus  gloriosus  (Holbrook) — bluespotted  sunfish 

This  species  is  listed  in  the  accession  list  for  Steinhart  Aquarium  as  having 
been  collected  in  March  1931  in  the  vicinity  of  Willows,  California.  The 
identification  was  made  by  Alvin  Seale,  but  the  specimens  were  not  saved. 
We  believe  this  to  be  a  misidentification. 

23.  Lepomis  macrochirus  Rafinesque — bluegill 

23a.  Lepomis  macrochirus  speciosus  (Baird  and  Girard) — southwestern 
bluegill 
According  to  Miller  (1952),  "The  southwestern  bluegill  .  .  .  is  also  now 
evidently  established  in  the  Colorado  River  through  introduction  .  .  .  {fide 
C.  L.  Hubbs  in  letter  of  10  May  1951,  to  R.  D.  Beland,  and  letter  from  Beland 
of  23  August  1951  to  W.  A.  Dill)."  Its  current  status  is  unknown. 

PERCI DAE— perch  family 

24.  Stizostedion  vitreum  (Mitchill) — walleye 

Miller  (1967)  summarized  the  history  of  walleye  introductions  in  Califor- 
nia. The  first  introduction  occurred  in  1 874,  when  1 6  fish  from  the  Missiquoi 
River  in  Vermont  were  stocked  in  the  Sacramento  River  near  Sacramento. 
One  was  caught  by  an  angler  but  nothing  further  was  recorded  from  the 
plant. 

The  second  attempt  spanned  the  years  1959  to  1963,  when  the  California 
Department  of  Fish  and  Game,  through  the  cooperation  of  the  Minnesota 
Conservation  Department,  secured  large  numbers  of  eggs  from  walleye 
captured  in  the  Detroit  River,  Minnesota.  About  5,350,000  fry  and  34,590 
fingerlings  were  stocked  in  five  southern  California  warmwater  reservoirs  in 
1959,  1960,  1962,  and  1963.  These  plants  were  successful  in  that  good 
survival  and  growth  were  experienced,  but  anticipated  angling  benefits  did 
not  accrue  and  the  program  was  abandoned.  Natural  spawning  did  not  take 
place  and  the  original  plants  gradually  died  out. 


4  " Lepomis  euryorus  McKay" .  Seale  (1930)  lists  "Sunfish,  Eupomotis  euryoris"  in  an  article  entitled,  "List  of  twenty 
fresh  water  fishes  found  in  California  that  may  be  used  in  small  aquariums  or  garden  pools."  The  Steinhart 
Aquarium  accession  list  for  1931  records  " Apomotis  euryorus"  as  collected  near  Willows,  California.  The 
identification  was  made  by  Alvin  Seale;  the  specimens  were  not  saved.  Hubbs  and  Hubbs  (1932)  have  proved 
that  the  nominal  species  "Lepomis  euryorus"  is  a  hybrid  between  Lepomis  cyanellus  and  Lepomis  gibbosus. 
Both  of  these  species  are  resident  in  California  but  L.  gibbosus  has  not  yet  been  recorded  from  near  Willows 
nor  do  we  have  any  records  of  its  presence  in  the  State  as  early  as  1930  or  1931. 


A  LIST  OF  CALIFORNIA  FISHES  35 

CICHLIDAE— cichlid  family 

25.   Cichlasoma  beam'  (Jordan) — green  guapote 

A  well-established  population  of  this  species  was  discovered  in  1975  in 
several  small  ponds  adjacent  to  Putah  Creek  in  Solano  County  by  A.  D. 
Castro,  Aquarist  with  the  California  Academy  of  Sciences  ( pers.  commun. ) . 
Identification  was  made  by  W.  I.  Follett.  Sampling  in  1979  did  not  uncover 
any  specimens  and  some  of  the  ponds  were  dry,  so  apparently  the  species 
did  not  survive  (R.  L.  Reavis,  Calif.  Dep.  Fish  and  Game,  pers.  commun.). 

ACKNOWLEDGMENTS 
We  are  indebted  to  the  following  individuals  for  their  interest  and  cooperation: 
Reeve  M.  Bailey,  Lillian  J.  Dempster,  W.  I.  Follett,  J.  D.  Hopkirk,  the  late  Carl  L. 
Hubbs,  Robert  N.  Lea,  Robert  R.  Miller,  and  Peter  B.  Moyle.  We  appreciate  the 
criticisms  by  these  and  other  scientists  and  have  incorporated  many  of  their 
suggestions  in  the  final  list.  We  have  not,  however,  been  able  to  reconcile  all  our 
differences,  so  one  should  not  assume  that  these  scientists  are  in  complete 
agreement  with  all  of  the  names  listed  here. 

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5  Actually  published  March  28,  1974. 


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Norden,  C.  R.  1961.  Comparative  osteology  of  representative  salmonid  fishes,  with  particular  reference  to  the 
grayling  (Thymallus  arcticus)  and  its  phylogeny.  Can.,  Fish.  Res.  Bd.,  J.  18(51:679-791. 

Richardson,  W.  M.,  J.  A.  St.  Amant,  L.  J.  Bottroff,  and  W.  L.  Parker.  1970.  Introduction  of  blue  catfish  into  California. 
Calif.  Fish  Came  56(4):31 1-312. 


38  CALIFORNIA  FISH  AND  CAME 

Robins,  C.  R.,  R.  M.  Bailey,  C.  E.  Bond,  j.  R.  Brooker,  E.  A.  Lachner,  R.  N.  Lea,  and  W.  B.  Scott.  1980.  A  list  of 
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Spec.  Publ.  no.  12,  174  p. 

Rosen,  D.  E.,  and  R.  M.  Bailey.  1%3.  The  poeciliid  fishes  (Cyprinodontiformes),  their  structure,  zoogeography  and 
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Ross,  S.  T.  1973.  The  systematics  of  Casterosteus  aculeatus  (Pisces:  Casterosteidae)  in  central  and  southern 
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113,  204  p. 


SUBPOPULATIONS  OF  NORTHERN  ANCHOVY  39 

Calif.  Fish  and  Came  67(1 ) :  39-5 1     1 981 

ELECTROPHORETIC,  MORPHOMETRY,  AND  MERISTIC 

STUDIES  OF  SUBPOPULATIONS  OF  NORTHERN 

A  N  C  H  O V Y,  ENGRA  UL  IS  MORDAX ' 

ANDREW  M.  VROOMAN  \  PEDRO  A.  PALOMA,  AND  JAMES  R.  ZWEIFEL3 

National  Oceanic  and  Atomospheric  Administration 

National  Marine  Fisheries  Service 

Southwest  Fisheries  Center 

P.O.  Box  271 

La  Jolla,  California  92038 

We  investigated  the  population  structure  of  northern  anchovy  found  between 
southern  Baja  California  and  Newport,  Oregon.  We  used  electrophoretic,  morpho- 
metric,  and  meristic  methods  in  our  studies,  and  the  results  indicate  the  presence 
of  three  distinct  anchovy  subpopulations. 

INTRODUCTION 

Hubbs  (1925)  and  McHugh(1951)  found  subpopulations  of  the  northern 
anchovy  along  the  west  coast  of  the  United  States  and  Mexico.  For  more 
effective  management  of  the  growing  United  States  and  Mexican  anchovy  fish- 
eries, knowledge  of  the  number  of  subpopulations  and  how  they  are  distributed 
geographically  is  necessary,  as  is  a  feasible  method  of  readily  distinguishing  the 
subpopulations.  In  this  study  we  used  electrophoretic  methods  to  distinguish 
subpopulations  and  delineate  their  geographical  range;  morphometric  and  mer- 
istic comparisons  were  made  between  these  subpopulations. 

Transferrin  Electrophoresis 
Transferrin  is  the  vertebrate  blood  serum  protein  responsible  for  binding  iron. 
Transferrin  polymorphism  has  been  reported  in  a  variety  of  teleost  fishes  by 
several  authors  including  Creyssel  et  al.  (1964),  Moller  (1966),  Moller  and 
Naevdal  (1966),  Barrett  and  Tsuyuki  (1967),  Fujinoand  Kang  (1968),  and  Utter 
(1969). 

Morphometries 
Hubbs  (1925)  found  small  morphometric  differences  in  samples  of  Engraulis 
mordax  collected  from  San  Francisco  to  southern  California.  He  also  described 
a  distinct  subspecies,  Engraulis  mordax  nanus,  which  he  found  inhabiting  the 
brackish  waters  of  San  Francisco  Bay.  We  were  unable  to  collect  the  bay 
anchovy. 

Meristics 
McHugh  (1951)  found  three  subpopulations  of  northern  anchovy:  one  off 
British  Columbia  to  northern  California,  one  off  southern  California  and  northern 
Baja  California,  and  one  off  central  and  southern  Baja  California.  He  based  his 
conclusion  on  the  mean  values  he  found  in  five  different  meristic  characters. 
Hubbs  (1925)  found  a  distinct  difference  in  vertebral  numbers  when  he  corn- 


Accepted  for  publication,  September,  1980 


2  Retired.  Current  address:  P.O.  Box  22461,  La  Jolla,  CA  92122 

3  Current  address:  National  Marine  Fisheries  Service,  75  Virginia  Beach  Drive,  Miami,  FL  33149 


40 


CALIFORNIA  FISH  AND  CAME 


pared  open  ocean  anchovies  with  bay  anchovies  from  San  Francisco  Bay.  He 
also  found  small  differences  in  vertebral  numbers  between  samples  of  open 
ocean  anchovies  from  San  Francisco  to  southern  California. 

MATERIALS  AND  METHODS 
Transferrin  Electrophoresis 

Anchovies  were  collected  from  Newport,  Oregon,  to  the  southern  end  of  Baja 
California  (Table  1 ) .  Availability  of  samples  was  limited  since  the  anchovies  had 
to  be  kept  alive  until  the  blood  samples  were  taken;  dead  or  preserved  fish  could 
not  be  used.  The  samples  came  primarily  from  commercial  live  bait  vendors  and 
from  short-duration  surface  hauls  made  with  a  midwater  trawl.  We  tried  to 
obtain  50  to  100  fish  per  sample,  but  this  was  frequently  impossible.  In  a  few 
cases,  two  smaller  samples  taken  very  closely  together  in  time  and  space  were 
combined  into  one;  other  samples  which  contained  less  than  35  readable  trans- 
ferrin types  and  could  not  be  combined  were  not  used  in  the  population  analysis. 

TABLE  1.     Sampling  Data  for  Northern  Anchovy  Subpopulation  Genetic  Testing  and 
Percent  Occurence  of  Transferrin  Alleles  in  the  Samples 


Location 
Site 

1 Newport,  Oregon 

2 Newport,  Oregon 

3 Eureka,  California 

4 Salt  Point,  California 

5 Monterey,  California 


6 San  Francisco,  California. 

7 San  Francisco,  California. 

8 Monterey,  California 

9 Monterey,  California 

10 Newport,  California 

11 San  Diego,  California 

12 San  Diego,  California 

13 Ensenada,  Mexico 

14 Ensenada,  Mexico 

15 Todos  Santos  Is,  Mexico. 

16 30°  50.5'N 

17 30°  17'N 

18 30°  12'N 

19 30°09'N 

20 29'33'N 


21 28°  33.2'N 

22 27*55.5'N 

23 27*52.5'N 

24 27°06.0'N 

25 2r04.0'N 

26 24°  30.0'N 


Percent  ti 

ansferrin 

/umber 

alleles 

A 

Date               t 

offish 

TfA 

Tf 

Tfc 

Tf° 

Northern  subpopulat 

ion 

July  1969 

84 

81.6 

8.3 

5.4 

4.8 

July  1970* 

66 

78.0 

13.6 

5.3 

3.0 

July  1970 

38 

77.6 

14.5 

5.3 

2.6 

July  1970 

54 

80.6 

13.0 

3.7 

3.4 

November  1969 

106 

76.9 

12.3 

5.7 

5.2 

Central  subpopulation 

April  1968 

48 

85.4 

7.3 

7.3 

0 

May  1968 

64 

72.7 

16.4 

10.9 

0 

May  1968 

87 

76.4 

13.8 

9.8 

0 

October  1967 

54 

82.4 

7.4 

10.2 

0 

August  1968 

94 

80.8 

10.1 

8.0 

1.1 

July  1968 

47 

81.9 

9.6 

7.4 

1.1 

July  1968 

100 

80.5 

12.0 

7.5 

0 

May  1968 

37 

82.4 

9.5 

8.1 

0 

July  1968 

94 

80.8 

11.2 

8.0 

0 

August  1969 

48 

83.3 

11.5 

5.2 

0 

August  1969 

43 

82.6 

11.6 

4.7 

1.2 

January  1969 

67 

84.3 

11.9 

3.7 

0 

January  1969 

70 

84.3 

10.0 

5.7 

0 

March  1968 

43 

84.9 

7.0 

7.0 

1.2 

November  1967 

36 

80.6 

11.1 

8.3 

0 

Southern  subpopulat 

ion 

November  1967 

83 

91.1 

4.8 

4.2 

0 

November  1967 

64 

90.6 

5.5 

3.9 

0 

November  1967 

87 

87.9 

6.3 

5.8 

0 

November  1967 

75 

88.7 

6.7 

4.7 

0 

November  1967 

72 

87.5 

6.9 

5.6 

0 

November  1967 

72 

88.2 

6.2 

5.6 

0 

SUBPOPULATIONS  OF  NORTHERN  ANCHOVY  41 

We  collected  blood  samples  from  live  fish  by  inserting  a  heparinized  capillary 
tube  through  the  gill  opening  into  the  dorsal  aorta.  The  tube  was  allowed  to  flow 
full  of  blood  and  then  was  sealed  on  the  bottom  with  a  bit  of  clay.  Filled  capillary 
tubes  were  then  centrifuged  at  about  2000  g for  5  min.  When  samples  were  not 
electrophoresed  immediately,  they  were  frozen  with  dry  ice  and  stored  at  0°  C. 

When  a  sample  was  ready  to  be  electrophoresed,  the  thawed  capillary  tube 
was  broken  off  at  the  interface  of  the  serum  and  red  cells;  the  cells  were 
discarded.  A  piece  of  absorbent  paper  was  touched  to  the  end  of  the  capillary 
tube  to  absorb  the  serum  until  a  colum  of  liquid  33  mm  long  remained;  this  was 
equivalent  to  25  jllI.  The  25  jal  of  serum  were  mixed  with  10  /xl  of  radioactive 
Fe59  and  allowed  to  incubate  for  at  least  10  min.  A  slot  cut  in  the  starch  gel  was 
filled  with  the  mixture  and  electrophoresed  for  1  h  40  min  at  150  v  in  a  horizon- 
tal, thin  layer,  starch  gel  apparatus.  After  electrophoresis  was  complete,  we 
prepared  autoradiographs  of  the  gels  using  a  modification  of  the  method  of 
Giblett,  Hickman,  and  Smithies  (1959). 

We  examined  the  hypothesis  that  each  band  represented  a  specific  transferrin, 
controlled  by  a  different  autosomal  allele  at  a  single  locus.  First,  we  verified  that 
"artificial  heterozygotes"  produced  by  mixing  equal  parts  of  sera  from  the 
appropriate  homozygous  types  produced  electrophoretic  patterns  indistinguish- 
able from  the  natural  heterozygous  types.  Secondly,  we  examined  the  statistical 
distributions  of  phenotypes  in  populations  thought  to  be  in  equilibrium  with 
respect  to  the  alleles  found. 

The  frequency  of  occurrence  of  the  transferrin  alleles  in  anchovy  samples  was 
calculated  as  1/N  (Oa+Va^Oq),  where  i  =  A,  B,  C,  D  (representing  alleles)  and 
j  z^z  i.  For  examle,  Oaa  is  the  number  of  phenotypes  AA  observed  and  N  is  the 
total  number  of  fish  in  the  sample.  Allocation  of  samples  to  the  subpopulations 
was  determined  by  cluster  analysis  (Sneath  and  Sokal  1973)  of  the  percentage 
distributions  of  alleles  for  each  sampling  site.  The  clustering  sequence  was  ob- 
tained by  identifying  the  two  sites  most  alike,  combine  the  two  and  clustering 
with  the  next  most  similar  site,  etc.  The  computer  program  used  was  BMDP2M 
written  at  the  Health  Sciences  Computer  Facility,  University  of  California,  Los 
Angeles.  Clustering  was  by  Euclidean  distance  (the  square  root  of  the  sums  of 
squares  of  differences  between  percent  alleles). 

Morphometries 
Morphometric  measurements  were  made  with  a  vernier  caliper  on  formalin 
preserved  anchovies  which  had  been  classified  to  subpopulation  by  transferrin 
gene  frequencies.  Head  length,  eye  diameter,  snout  to  post-orbital  margin,  head 
depth,  and  body  depth  were  measured.  Allometric  regressions  (In  y  =  a  +  b 
In  x)  were  calculated  for  each  of  the  five  morphometric  measurements,  where 
x  is  the  standard  length  (sl). 

Meristics 

We  took  all  meristic  counts  from  formalin  preserved  samples  which  we  had 
classified  as  northern,  central,  or  southern  subpopulation  anchovies  on  the  basis 
of  transferrin  gene  frequencies.  Counts  were  made  from  x-ray  plates  with  the  aid 
of  a  binocular  dissecting  microscope.  Vertebrae,  anal  fin  rays,  and  dorsal  fin  rays 
were  counted.  The  vertebral  counts  did  not  include  the  basioccipital  nor  the 
hypural. 


42  CALIFORNIA  FISH  AND  GAME 

RESULTS  AND  DISCUSSION 
Transferrin  Electrophoresis 

We  found  that  transferrin  polymorphism  in  the  northern  anchovy  orginates  in 
a  genetic  system  of  four  co-dominant  autosomal  alleles,  each  controlling  the 
formation  of  a  single  protein  with  a  specific  anodal  migration  rate  when  electro- 
phoresed  in  starch  gel.  The  four  iron-binding  protein  bands  were  designated  A, 
B,  C,  and  D.  The  migration  distance  in  a  standard  run  was  23.4  mm  for  band 
A,  21.1  mm  for  band  B,  19.0  mm  for  band  C,  and  16.2  mm  for  band  D  (Figure 
1). 

Northern  (sites  1-5)  and  southern  (sites  21-16)  groups  were  clearly  distin- 
guished by  cluster  analysis  (Figure  2)  of  the  transferrin  alleles'  percentage  of 
occurrence  (Table  1 ).  A  central  group  (site  6  and  sites  9-20)  was  also  evident. 
However,  samples  taken  at  sites  7  (San  Francisco)  and  8  (Monterey)  during 
May  of  1968  were  distinct  from  all  groups.  These  samples  were  anomalous  in 
that  they  were  not  intermediate  between  the  major  groupings  but  rather  repre- 
sented extreme  levels  for  all  alleles;  thus  a  mixture  of  populations  or  interbreed- 
ing does  not  suffice  as  a  rational  explanation.  These  anomalies  are  believed  to 
be  due  to  occasional  indistinct  separation  of  the  first  three  bands  within  the  gel. 
The  absence  of  the  D  allele  separates  these  samples  from  the  northern  group  and 
the  relative  frequencies  of  the  B  and  C  alleles  separates  them  from  the  southern 
group;  thus  we  included  them  within  the  central  subpopulation.  There  is  an 
overlap  in  the  geographical  range  of  samples  attributed  to  the  northern  and 
central  subpopulations;  the  southernmost  sample  from  the  northern  subpopula- 
tion was  taken  in  Monterey  in  November  1969,  and  the  northernmost  samples 
from  the  central  subpopulation  were  taken  from  San  Francisco  Bay  in  April  and 
May  1968,  an  overlap  of  about  70  nautical  miles.  This  does  not  mean  that  the 
two  subpopulations  were  necessarily  present  in  these  areas  at  the  same  time; 
instead,  both  subpopulations  may  tend  to  move  north  in  the  spring  and  summer 
and  return  toward  the  south  in  the  fall  and  winter.  Anchovy  tagging  studies 
conducted  by  California  Department  of  Fish  and  Game  support  the  north  and 
south  movements  (Haugen,  Messersmith,  and  Wickwire  1969). 

The  northern  subpopulation  was  distinguished  from  the  other  two  by  the 
Tf°  allele  which  was  not  found  in  the  southern  subpopulation,  was  rare  in  the 
central  subpopulation  (0.2%),  but  occurred  at  a  rate  of  4.02%  in  the  northern 
subpopulation  (Table  2).  The  central  subpopulation  was  distinguished  from  the 
southern  one  by  the  frequency  of  occurrence  of  Tf*  and  Tf*  alleles.  Tf*  occurred 
at  a  rate  of  88.96%  in  the  southern  subpopulation  compared  to  81.17%  in  the 
central  one;  Tf*  occurred  at  1 1 .0%  in  the  central  subpopulation  and  only  6.07% 
in  the  southern  one.  Chi-square  goodness  of  fit  tests  on  observed  numbers  of 
phenotypes  for  the  three  subpopulations  versus  the  expected  numbers  calculat- 
ed from  the  Hardy-Weinberg  equilibrium  formula  (Table  2)  support  the  four- 
allele  hypothesis. 

Similarity  or  dissimilarity  of  the  subpopulations  was  judged  on  the  basis  of  the 
observed  phenotypic  distributions  with  northern-central  and  central-southern 
differences  treated  separately.  We  found  the  rare  allele  to  be  important  in 
discriminating  the  northern  subpopulation,  whereas  the  predominant  alleles 
provided  the  discriminatory  power  for  the  central  and  southern  subpopulations 
(Table  3).  To  avoid  difficulties  with  expectations  in  the  statistical  tests,  all 


SUBPOPULATIONS  OF  NORTHERN  ANCHOVY 


43 


combinations  of  the  Tf°  allele  were  grouped  for  the  chi-square  tests.  The  results 
of  the  chi-square  tests  for  independence  [KO  — E)2/E]  are  as  follows: 

North-central  X2  =  61.99,  d.f.  =  6;  P  <  .005 
Central-southern  X2  =  27.88;  d.f.  =  6;  P  <  .005 

Both  indicate  highly  significant  differences  between  subpopulations. 


Transferrin     Types 
AA     AB      AC      AD      BB       BC      BD     CC      CD     DD 


mm 
20. 

10- 


+ 


FIGURE   1.     Transferrin  pattern  types  found  in  northern  anchovy. 


TABLE  2.  Gene  Frequencies  in  the  Three  Northern  Anchovy  Subpopulations  and  the  Ob- 
served and  Expected  Number  of  Phenotypes.  The  Expected  Numbers  were  Cal- 
culated from  the  Hardy-Weinberg  Equilibrium  Formula 

Allele  Northern  (N)  Central  (C)  Southern  (S) 

TfA  78.88%  81.17%  88.96% 

Tf'  11.92  11.00  6.07 

TFC  5.17  7.56  4.97 

TfD  4.02  0.27  0.00 

Phenotype  Expected  Observed  Expected  Observed  Expected  Observed 

AA 216.52          215  614.05          619  358.51           360 

AB  65.47             67  166.40           161  48.93             47 

AC 28.39             30  114.44           111  40.03             39 

AD 22.09            22  4.05              3  0.00              0 

BB 4.95              4  11.27            13  1.67              2 

BC  4.29              4  15.50             17  2.73              4 

BD 3.34              4  0.55              1  0.00              0 

CC 0.93              0  5.33              6  1.12              1 

CD 1.45              2  0.38              1  0.00              0 

DD 0.56              0  0.01              0  0.00              0 

X£  =  0.36;  d.f.  =  5;  P  >  0.995 
X£  =  1 .38;  d.f.  =  4;  P  >  0.750 
X|  =  0.78;  d.f.  =  3;  P  >  0.750 


44 


CALIFORNIA  FISH  AND  CAME 


SITE 


[I 


I 

5 
4 

3 

2 

21 

22 

24 

25 
26 

23 

15 

18 

16 

17 

9 

10 

II 

13 

14 

20 

12 

19 

6 

8 

7 

FIGURE  2.  Cluster  tree  diagram  of  sample  sites  by  proportion  of  transferrin  alleles.  Clustering 
sequence  is  determined  by  the  distance  of  the  vertical  bars  from  the  solid  vertical  line.  Example: 
23  and  26  are  most  similar,  then  14  and  20,  etc.  Subpopulations  identified  are  northern  (1-5), 
southern  (21-26)  and  central  (6-20). 


Thus,  the  transferrin  data  support  the  conclusion  of  McHugh  ( 1 951 )  that  there 
are  three  subpopulations  of  northern  anchovies  in  the  area.  The  close  agreement 
in  numbers  of  transferrin  types  observed  with  the  expected  numbers  (Table  2) 
calculated  from  the  Hardy-Weinberg  equilibrium  formula  indicates  three  geneti- 
cally distinct  subpopulations  with  little  or  no  interbreeding;  a  northern  one  from 
about  Monterey  north,  a  central  one  from  San  Francisco  to  about  29°  N  lat,  and 
a  southern  one  south  of  29°  N  (Figure  3). 


SUBPOPULATIONS  OF  NORTHERN  ANCHOVY 


45 


FIGURE     3.     Distribution  of  three  northern  anchovy  subpopulations  based 

on  transferrin  allele  frequencies. 


46 


CALIFORNIA  FISH  AND  CAME 


TABLE  3.     Observed   and    Expected    Numbers   of   Phenotypes   Assuming   No   Differences 
Between  Subpopulations 

Phenotype 


AD+BD+ 

AA 

AB 

AC 

BB 

BC 

CC 

CD+DD 

Total 

0 

215 

67 

30 

4 

4 

0 

28 

348 

E 

226.7 

62.0 

38.3 

4.6 

5.7 

1.6 

9.0 

O 

619 

161 

111 

13 

17 

6 

5 

932 

Central 

E 

607.2 

166.0 

102.7 

12.4 

15.3 

4.4 

24.0 

TOTAL 

834 

228 

141 

17 

21 

6 

33 
AD+BD+ 

1280 

AA 

AB 

AC 

BB 

BC 

CC 

CD+DD 

Total 

0 

619 

161 

111 

13 

17 

6 

5 

932 

Central 

E 

658.8 

140 

1009 

10.1 

14.1 

4.73 

3.4 

0 

360 

47 

39 

2 

4 

1 

0 

453 

Southern 

E 

320.2 

68 

49.1 

4.9 

6.9 

2.3 

1.6 

TOTAL 

979 

208 

150 

15 

21 

7 

5 

1385 

Morphometries 
Morphometric  measurements  were  taken  on  613  fish  (Table  4).  Except  for 
eye  diameter,  there  was  no  evidence  that  the  slope  coefficients  b  differed  among 
the  subpopulations  (Table  5)  test  for  parallel  lines.  The  slope  for  eye  diameter 
in  the  northern  group  differed  from  those  of  the  southern  and  central,  but  the 
latter  two  showed  no  statistical  difference.  Only  body  depth  indicated  direct 
proportionality,  i.e.,  b  was  not  significantly  different  from  unity. 

TABLE  4.     Sampling  Data  for  Northern  Anchovy  Morphometric  and  Meristic  Testing 

Mean  standard  Standard 

Subpopulation  N       length  (mm)  Range  deviation 

Northern 206  97.20  77-135  10.01 

Central 225  104.47  70-136  15.39 

Southern 182  90.11  50-116  13.65 

Because  the  slope  coefficients  for  eye  diameter  differed  among  subpopula- 
tions, no  test  for  a  common  regression  line  was  performed  for  eye  diameter.  All 
other  tests  for  a  common  regression  line,  i.e.,  the  same  slope  and  intercept,  were 
highly  significant  (P  <  0.01).  In  each  instance,  the  more  similar  of  the  two 
subpopulations  were  also  tested  for  a  common  relationship.  Body  depth  indicat- 
ed no  difference  between  the  southern  and  central  subpopulations.  Otherwise 
all  differences  between  subpopulations  were  significant  at  P  <  0.05.  The  analysis 
of  covariance  is  not  entirely  appropriate  for  morphometric  data  since  both 
variates  are  subject  to  error,  especially  for  field  sampling  where  the  size  range 
of  the  samples  is  rarely  the  same  and  cannot  be  controlled.  It  is  well  to  insist 
on  a  conservative  level  of  statistical  significance;  therefore,  we  calculated  the 
estimated  morphometric  measurements  for  a  70-,  100-,  and  130-mm  anchovy 
from  each  subpopulation  (Table  6).  Southern  subpopulation  anchovies  showed 
a  distinctly  longer  head,  larger  eye,  and  longer  snout  to  post-orbit  than  did  either 
central  or  northern  ones.  Northern  subpopulation  anchovies  exhibited  a  deeper 
body,  and  northern  and  southern  subpopulation  anchovies  showed  a  slightly 
deeper  head  than  did  those  of  the  central  stock.  Average  morphometric  meas- 


SUBPOPULATIONS  OF  NORTHERN  ANCHOVY 


47 


urements  calculated  for  1 0-mm  intervals  from  70  to  1 20  mm  show  the  consistent 
pattern  of  differences  between  the  subpopulations  at  all  sizes  (Table  7). 

TABLE  5.     Covariance  Analysis  for  the  Three  Northern  Anchovy  Subpopulations: 
SL  =  Standard  Length 


Subpopulation 

Southern 

Central 

Northern 

Southern 

Central 

Northern 

Southern 

Central 

Northern 

Southern 

Central 

Northern 

Southern 

Central 

Northern 


InUd 
InUd 
InUd 

InUd 
InUd 
InUd 

InUd 
InUd 
InU 

InUc 
InU, 

InUo 

In  Ud  ■ 
InU  : 
InU,  = 


Head  depth  (hd) 
-1.637  +  .967  In  LM 
-1.661  +  .968  In  lu 
-1.570  +  .955  In  La 

Body  depth  (bd) 
-1.982  +  1.052  In  Lu 
-1.928  +  1,039  In  L* 
-1.770  +  1.012  In  L* 

Eye  diameter  fed) 
-1,551  +.753  In  L«. 
-1.366  +  .692  In  L* 
-2.235  +  .882  In  L* 
Snout  to  post-orbit  (po) 
-1.273  +  .823  In  La 
-1.313  +  .816lnLu 
-1.042  +  .755  In  L* 

Head  length  (hi) 
-0.850  +  .932  In  L» 
-1.027  +  .955  In  La. 
-0.784  +  .900  In  L* 


Test  for  common 
regression  line 


fw  =  12.79* 


Fw  =  16.00' 


ftM7  =  125.04 


Test  for 
parallel  lines 


>  2.607  " 


r  2,407 


F2.«o7  =  8.37* 


F2.407  —  2.76*' 


Fmo  -  158.92*  f7jm  =  2.56*  * 


'  Significant  P  ^  .01 
"  Not  significant 


TABLE  6.     Estimated  Morphometric  Measurements  of  70,  100,  and  130  mm  Standard  Length 
Northern  Anchovies  Expressed  as  Percent  of  Standard  Length 

Head          Eye  Snout  to  Head  Body 

Length    Subpopulation                                                  length  diameter  post-orbit  depth  depth 

Northern  29.8            6.5            12.4            17.2  17.9 

70    Central 29.6            6.9            12.3             16.6*  17.2 

Southern 31.9*          7.4*          13.2*  16.9  17.2 

Northern  28.8  6.2  11.4  16.9  18.0* 

100     Central 29.1  6.2  11.5  16.4*  17.4 

Southern 31.1*  6.8*  12.4*  16.7  17.5 

Northern  28.1  6.0  10.7  16.7  18.0* 

130     Central 28.8  5.7  11.0  16.2*  17.6 

Southern 30.6*  6.4*  11.8*  16.6  17.7 

*  Significant  difference  (P  <  0.01)  between  subpopulations  within  length  group. 

Hubbs  (1925)  also  reported  longer  head  length  (31.9%  sl)  for  a  San  Fran- 
cisco Bay  subspecies  Engraulis  mordax  nanus  which  also  had  a  greater  body 
depth  (19.7%  sl)  than  did  the  open  ocean  anchovies  (18.1%  sl). 

Mais  (1974)  reported  that  southern  subpopulation  anchovies  are  much  small- 
er than  central  stock  anchovies.  Of  the  2,332  fish  he  measured  from  96  samples 
collected  in  more  than  51/2  yr  south  of  lat  28°30'  N,  less  than  10%  exceeded  106 


48 


CALIFORNIA  FISH  AND  CAME 


mm  total  length  (the  minimum  legal  limit  of  the  California  anchovy  reduction 
fishery),  while  79%  of  the  central  stock  anchovies  were  106  mm  or  greater. 
Southern  anchovies  were  significantly  smaller  than  central  ones  at  all  ages  and 
nearly  attained  their  maximum  length  by  age  3,  while  central  subpopulation 
anchovies  continued  to  grow  for  at  least  3  more  years. 

TABLE  7.     Average  Morphometric  Measurements  (mm)  in  Three  Northern  Anchovy  Sub- 
populations  in  10  mm  Intervals  of  Standard  Length;  N  =  Northern; 
C  =  Central;  S  =  Southern  Subpopulations. 


Standard 

Body 

Head 

Head 

Snout- 

Eye 

Number  of 

Interval 

length 

depth 

depth 

length 

postorbital 

diameter  observations 

N 

0 

70-  79 

C 

75.5 

12.8 

12.4 

22.4 

9.2 

5.2 

24 

S 

75.6 

13.0 

13.0 

24.9 

10.3 

5.9 

34 

N 

86.2 

15.9 

15.0 

25.2 

10.3 

5.5 

42 

80-  89 

C 

84.0 

14.9 

14.6 

24.9 

10.1 

5.5 

3 

S 

83.7 

14.4 

14.0 

26.7 

10.9 

6.1 

36 

N 

92.9 

16.4 

15.5 

27.0 

10.8 

5.8 

91 

90-  99 

C 

95.4 

16.7 

15.8 

27.7 

11.1 

5.9 

62 

S 

95.0 

16.5 

15.8 

29.4 

11.8 

6.4 

51 

100-109 

N 

105.1 

18.9 

17.7 

30.0 

11.8 

6.4 

33 

C 

103.6 

18.2 

16.8 

29.7 

11.6 

6.3 

51 

s 

103.5 

18.3 

17.3 

31.7 

12.6 

6.8 

42 

110-119 

N 

112.1 

20.5 

19.1 

31.9 

12.5 

6.9 

37 

C 

114.2 

20.1 

18.9 

33.1 

12.9 

6.8 

37 

s 

111.6 

19.8 

18.6 

33.9 

13.4 

7.3 

10 

Meristics 
Vertebrae 

Northern  subpopulation  anchovies  had  the  greatest  mean  number  of  vertebral 
centra  (Table  8).  The  mean  for  the  central  subpopulation  was  significantly  less 
than  that  of  the  northern  subpopulation  (d  =  0.46;  Fli404  =  41.83;  p  <  0.001 ). 
This  was  also  the  case  for  the  southern  subpopulation  with  regard  to  the  northern 
one  (d  =  0.43;  F1/386  =  49.44;  p  <  0.001 ).  There  was  no  significant  difference 
between  central  and  southern  subpopulation  mean  number  of  vertebrae  (d  = 
0.03;  F1/380  =  0.16;  p  <  0.25). 

Hubbs  (1925)  reported  a  mean  number  of  44.73  vertebrae  for  offshore  north- 
ern anchovies  off  San  Francisco,  which  is  very  close  to  the  44.75  we  found  for 
the  northern  subpopulation.  When  we  partitioned  McHugh's  (1951,  Tables  2 
and  3)  vertebral  data  into  probable  subpopulations  (northern,  central,  or  south- 
ern) merely  on  the  basis  of  location  of  capture,  we  calculated  his  northern 
subpopulation  samples  to  have  a  mean  of  44.74  vertebrae,  again  in  good  agree- 
ment with  ours.  His  southern  subpopulation  samples  had  a  mean  of  44.32 
vertebrae,  identical  to  ours. 


SUBPOPULATIONS  OF  NORTHERN  ANCHOVY 


49 


TABLE  8.     Meristic  Analysis  of  the  Three  Subpopulations  of  Northern  Anchovies:  x  =  Mean, 

S  =  Standard  Deviation,  and  S  =  Standard  Error  of  Mean 

No.  Range  x  S  Sr 

Vertebrae 

Northern  subpopulation 206  43-46  44.75  0.6325  0.0441 

Central  subpopulation 200  42-46  44.29  0.7994  0.0565 

Southern  subpopulation 182  42-45  44.32  0.5734  0.0425 

Anal  fin  rays 

Northern  subpopulation 

Male 136  20-24  22.18  0.9043  0.0775 

Female _70  20-25  22.20  0.9869  0.1180 

TOTAL 206  20-25  22.19  0.9308  0.0649 

Central  subpopulation 

Male 94  19-25  22.43  1.1499  0.1186 

Female _I06  19-25  22.36  0.9481  0.0921 

TOTAL 200  19-25  22.39  1.0456  0.0739 

Southern  subpopulation 

Male 109  20-25  22.53  1.0850  0.1039 

Female _64  20-25  22.64  1.0445  0.1306 

TOTAL 173  20-25  22.58  1.0686  0.0808 

Dorsal  fin  rays 

Northern  subpopulation 

Male 136  15-18  16.26  0.5962  0.0511 

Female _70  15-17  16.43  0.6272  0.0750 

TOTAL 206  15-18  16.32  0.6108  0.0426 

Central  subpopulation 

Male 94  15-18  16.46  0.6336  0.0654 

Female JI06  15-18  16.37  0.6666  0.0647 

TOTAL 200  15-18  16.41  0.6512  0.0460 

Southern  subpopulation 

Male 110  15-18  16.35  0.6146  0.0586 

Female _65  15-17  16.43  0.6116  0.0759 

TOTAL 180*  15-18  16.37  0.6075  0.0453 

*  Includes  five  juveniles. 


When  we  compared  the  central  subpopulations,  however,  we  calculated  a 
mean  of  44.88  vertebrae  for  his  samples,  which  is  0.59  greater  than  ours.  His  data 
indicated  a  high  degree  of  variability  from  month  to  month  and  year  to  year.  For 
instance,  his  data  for  the  mean  number  of  vertebral  centra  in  anchovy  post- 
larvae  off  southern  California  (McHugh  1951,  Table  4)  was  44.21  in  1945,44.69 
in  1947,  44.84  in  1948,  and  44.65  in  1949. 

Anal  Fin  Rays 

McHugh  (1951 )  reported  strong  evidence  for  sexual  dimorphism  in  the  num- 
ber of  anal  fin  rays,  with  those  of  males  exceeding  those  of  females  by  0.1 3  rays. 
Our  data  did  not  indicate  such  dimorphism.  When  all  of  our  samples  were 


50 


CALIFORNIA  FISH  AND  CAME 


combined  according  to  sex,  fin  rays  of  females  exceeded  those  of  males  by  only 
0.02  rays.  When  each  subpopulation  was  tested  separately  for  sexual  dimor- 
phism, the  greatest  difference  found  (Table  8)  was  in  the  southern  subpopula- 
tion where  females  exceeded  males  by  0.11  anal  fin  rays,  which  was  not 
significant  (F1171  =  0.43;  p  >  0.25). 

When  we  compared  both  males  and  females  combined  for  each  of  the  three 
subpopulations,  we  found  that  the  northern  subpopulation  had  a  mean  anal  fin 
ray  count  0.20  less  than  that  of  the  central  subpopulation;  the  difference  was 
significant  (F1404  =  4.18;  p  <  0.05).  The  mean  number  of  anal  rays  for  the 
northern  subpopulation  was  0.39  fewer  than  that  for  the  southern  subpopulation, 
which  was  highly  significant  (FU79  =  14.33;  p  <  0.001 ).  Central  and  southern 
subpopulations  differed  by  0.19  rays,  which  was  not  significant  (F1373  =  2.93; 
p  <  0.10). 

When  we  partitioned  McHugh's  (1951 )  anal  fin  ray  data  into  their  probable 
subpopulations  on  the  basis  of  locality,  we  found  his  mean  anal  fin  ray  count 
for  the  combined  northern  subpopulation  samples  to  be  only  0.03  fewer  than 
ours.  His  southern  subpopulation  mean  ray  count  was  only  0.1 6  fewer  than  ours, 
but,  as  with  vertebrae,  there  was  a  large  difference  in  the  central  subpopulation, 
with  his  mean  count  being  0.36  greater  than  ours  (Table  9). 

TABLE  9.     Mean  Numbers  of  Anal  Fin  Rays  From  McHugh  (1951,  Table  11,  12  and  13) 
Compared  with  Those  of  This  Study:  N  =  Number,  x  =  Mean 


This 

McHugh  (1951) 

study 

Adult 

Adult 

Combined  Combined 

males 

females 

Young 

total 

total 

Northern  subpopulation 

N 

105 

87 

105 

297 

206 

x 

22.21 

21.94 

22.28 

22.16 

22.19 

Central  subpopulation 

N 

284 

418 

404 

1,106 

200 

x  

22.81 

22.66 

22.79 

22.75 

22.39 

Southern  subpopulation 

N 

41 

49 

100 

190 

175 

x  

22.83 

22.55 

22.18 

22.42 

22.58 

Dorsal  Fin  Rays 

McHugh  (1951 )  noted  a  probable  sexual  dimorphism  in  mean  dorsal  fin  ray 
counts;  males  had  a  grand  mean  difference  of  0.12  count  greater  than  that  of 
females.  Males  in  our  samples  averaged  fewer  dorsal  rays  that  did  females  in 
both  northern  and  southern  subpopulations  but  more  rays  than  did  females  in 
the  central  subpopulation  (Table  8).  Since  the  differences  between  the  sexes 
were  not  significant  and  were  not  consistent  in  direction,  we  concluded  that  the 
variations  were  random  and  we  might  combine  the  data  for  males  and  females 
when  comparing  the  three  subpopulations. 

Central  subpopulation  anchovies  had  the  largest  mean  number  of  dorsal  fin 
rays,  0.09  greater  than  that  of  the  northern  stock  and  0.04  greater  than  that  of 
the  southern  stock,  but  these  differences  are  not  significant  ( northern  vs.  central: 
F1.904  =  2.27,  p  >  0.10;  northern  vs.  southern:  Fli384  =  0.83,  p  >  0.25;  central 
vs.  southern:  FU78  =  0.33,  p  >  0.25). 


SUBPOPULATIONS  OF  NORTHERN  ANCHOVY  51 

SUMMARY  AND  CONCLUSION 

We  found  three  distinct  subpopulations  of  northern  anchovies  inhabiting  the 
coastal  waters  between  Newport,  Oregon  and  the  southern  end  of  Baja  Califor- 
nia, Mexico:  northern,  between  Newport  and  Monterey;  central,  between  San 
Francisco  and  lat  29°  N  and  southern,  south  of;  lat  29°  N.  There  was  an  overlap 
of  about  70  nautical  miles  for  the  northern  and  central  subpopulations  (Figure 
3).  Our  conclusion  was  based  on  our  transferrin  electrophoresis  study,  and 
supports  McHugh's  ( 1951 )  conclusion  of  three  subpopulations.  Our  morphom- 
etric  and  meristic  work  also  supports  our  genetic  findings. 

Given  a  sample  of  anchovies  from  the  southern  subpopulation  range,  our 
studies  showed  that  it  could  be  identified  as  such  if  the  mean  head  length,  snout 
length,  and  eye  diameter  were  greater  than  those  of  the  northern  and  central 
subpopulations,  and  if  the  mean  standard  length  of  the  sample  (at  all  ages)  were 
signficantly  less  than  that  of  the  other  two  subpopulations.  A  sample  of  ancho- 
vies from  the  northern  subpopulation  range  could  be  identified  as  such  if  it  had 
i)  a  greater  mean  number  of  vertebrae  and  fewer  anal  fin  rays  than  either  central 
or  southern  subpopulation  anchovies,  and  ii)  if  the  mean  head  depth  were 
greater  than  that  of  central  subpopulation  anchovies.  However,  any  conclusion 
on  subpopulations  involving  meristic  counts  should  take  into  consideration 
McHugh's  ( 1 951 )  work  showing  a  high  degree  of  variability  in  these  parameters 
from  year  to  year  and  even  from  month  to  month. 

ACKNOWLEDGMENTS 

We  wish  to  thank  P.  Arasmith  for  her  valuable  assistance  with  the  tedious 
laboratory  work.  We  also  thank  K.  Mais  and  the  crew  of  the  California  Depart- 
ment of  Fish  and  Game  research  vessel  ALASKA  for  providing  all  of  the  trawl- 
caught  anchovy  samples. 

REFERENCES 

Barrett,  I.,  and  H.  Tsuyuki.  1967.  Serum  transferrin  polymorphism  in  some  scrombroid  fishes.  Copeia,  (3):551-557. 
Creyssel,  R.,  P.  Silberzan,  G.  Richard,  and  Y.  Manual.  1964.  Etude  du  serum  de  carpe  (Cyprinus  carpio)  par 

electrophorses  en  gel  d'  amidon.  Bull.  Soc.  Chim.  Biol.,  46:149-159. 
Fujino,  K.,  and  T.  Kang.  1968.  Transferrin  groups  of  tunas.  Genetics,  59:79-91. 
Giblett,  E.  R.,  C.  G.  Hickman,  and  O.  Smithies.  1959.  Serum  transferrins.  Nature,  183:1589-1590. 
Haugen,  C.  W.,  J.  D.  Messersmith,  and  R.  H.  Wickwire.  1969.  Progress  report  on  anchovy  tagging  off  California, 

March  1966  through  May  1966.  Calif.  Dept.  Fish  and  Game,  Fish.  Bull.  (147):75-86. 
Hubbs,  C.  L.  1925.  Racial  and  seasonal  variation  in  the  Pacific  herring,  California  sardine  and  California  anchovy. 

Calif.  Dept.  Fish  and  Game,  Fish.  Bull.,  (8):1-23. 
Mais,  K.  F.  1974.  Pelagic  fish  surveys  in  the  California  Current.  Calif.  Dept.  Fish  and  Game,  Fish.  Bull.,  (162):1-79. 
McHugh,  ).  L.  1951.  Meristic  variations  and  populations  of  northern  anchovy  (Engraulis  mordax).  Scripps  Inst. 

Oceanogr.  Bull.,  6(3):123-160. 
Moller,  D.  1966.  Polymorphism  of  serum  transferrin  in  cod.  Fisk.  Dir.  Skr.  HavUnders.,  14:51-60. 
Moller,  D.,  and  G.  Naevdal.  1966.  Serum  transferrins  of  some  gadoid  fishes.  Nature,  210:  317-318. 
Sneath,  P.  H.  A.,  and  R.  R.  Sokal.  1973.  Numerical  taxonomy:  The  principles  and  practice  of  numerical  classification. 

W.  H.  Freeman  and  Co.,  San  Francisco,  573  p. 

Utter,  F.  M.  1969.  Biochemical  polymorphisms  in  the  Pacific  hake  (Merluccius  productus) .  a,  esterase  polymor- 
phism in  vitreous  fluids;  b,  lactate  dehydrogenase  isozymes;  c,  transferrin  variats.  Dissertation.  Univ.  Calif., 
Davis.  60  p. 


52  CALIFORNIA  FISH  AND  CAME 

Calif.  Fish  and  Came  67  ( 1 ) :  52-61      1 981 

DENNING  CHARACTERISTICS  OF  BLACK  BEARS,  URSUS 

AMERICANUS,  IN  THE  SAN  BERNARDINO  MOUNTAINS 

OF  SOUTHERN  CALIFORNIA  ] 

HAROLD  J.  NOVICK, 7  JOHN  M.  SIPEREK, 3  and  GLENN  R.  STEWART 

California  State  Polytechnic  University,  Pomona 

3801  W.  Temple  Avenue 

Pomona,  California  91768 

Denning  information  was  obtained  from  field  studies  of  nine  black  bears  during 
two  winters.  Seven  dens  were  examined.  They  were  located  in  areas  of  steep  terrain 
and  minimal  human  disturbance.  All  of  these  dens  were  dug  under  large  boulders 
or  beneath  the  bases  of  trees.  Six  were  located  in  areas  where  the  Canyon  Oak  Series 
was  dominant  or  co-dominant;  the  other  was  located  in  the  Ponderosa  Pine  Series. 
The  mean  denning  period  of  seven  males  was  from  mid-December  to  mid-March. 
The  range  of  denning  periods  of  all  bears  was  late  October  to  early  April.  Bears 
denned  significantly  longer  and  emerged  later  in  the  wet  winter  of  1977-78  than  in 
the  relatively  dry  winter  of  1976-77.  The  cumulative  effects  of  weather  probably 
caused  these  differences. 

INTRODUCTION 

Throughout  their  geographic  range,  black  bears  vary  greatly  in  denning  habits 
and  habitat  preferences.  The  majority  of  investigations  have  been  done  in  re- 
gions with  moderate  to  severe  winters.  Some  recent  studies  (Poelker  and  Hart- 
well  1973;  Lindzey  and  Meslow  1976;  Hamilton  and  Marchinton,  1980;  LeCount 
1980)  have  reported  on  areas  with  relatively  mild  winters.  Many  researchers, 
including  Erickson  (1965);  Jonkel  and  Cowan  (1971 );  Craighead  and  Craighead 
(1972);  Lindzey  and  Meslow  (1976);  LeCount  (1980);  and  Reynolds  and  Bee- 
cham  (1980),  have  determined  the  length  and  dates  of  denning.  Various  factors 
responsible  for  wide  variations  in  denning  habits  have  been  documented,  includ- 
ing food  availability,  physical  condition  of  bears,  and  cumulative  effects  of 
weather.  Only  one  study  (LeCount,  1980)  has  described  denning  times  and  den 
site  selection  in  the  climatically  mild  Southwest. 

Black  bears  were  introduced  into  southern  California  in  1933  (Burghduff 
1935).  However,  no  information  concerning  the  ecology  of  these  bears  had 
been  gathered  until  1974,  when  we  began  a  long-term  study  of  the  San  Bernar- 
dino Mountain  population.  That  study's  primary  objectives  were  to  determine 
food  habits  (Boyer  1976),  habitat  utilization  (Novick  1979),  and  physical  char- 
acteristics (Siperek  1979)  of  the  bears.  Concurrently,  we  conducted  a  3-year 
investigation  of  denning  characteristics  of  bears  in  this  population.  The  purposes 
of  this  study,  reported  here,  were  to  determine  den  site  characteristics  and  time 
and  duration  of  denning. 

STUDY  AREA 
The  study  area  encompasses  approximately  170  km  2  of  the  Banning  Canyon 
and  Mill  Creek  drainages  and  lies  in  the  southeastern  portion  of  the  San  Bernar- 
dino Mountains  (Figure  1).  Topography  is  characterized  by  deep,  rocky  can- 

1  Accepted  for  publication  August  1980. 

2  Current  address:  California  Department  of  Fish  and  Came,  350  Golden  Shore,  Long  Beach,  California,  90802. 

3  Current  address:  California  Department  of  Fish  and  Came,  P.O.  Box  607,  Red  Bluff,  California,  96080. 


BLACK  BEAR  DENNING  CHARACTERISTICS 


53 


yons  and  steep  ridges,  with  many  slopes  exceeding  45  degrees.  Elevations  range 
from  1,200  to  over  2,750  m.  A  Mediterranean  climate  of  infrequent  winter  rains 
and  pronounced  summer  drought  is  characteristic.  Annual  precipitation  in  the 
form  of  rain  or  snow  at  the  Mill  Creek  Ranger  Station  ranges  between  21  and 
104  cm  and  averages  49  cm.  Snow  cover  is  common  at  the  higher  elevations 
from  about  late  December  to  mid-March.  However,  at  lower  elevations  it  does 
not  remain  long,  especially  on  southern  exposures.  Average  temperatures  range 
from  3°C  during  midwinter  to  over  35°C  in  summer. 


KILOMETERS 


FIGURE  1.     Geographic  location  of  the  study  area  in  the  San  Bernardino  Mountains,  California.  Den 
locations  are  indicated  by  + 

Climate,  topography,  and  fire  history  influence  the  type,  distribution  and 
abundance  of  plant  communities  present.  The  area  has  a  heterogeneous  mixture 
of  Conifer  Forest,  Woodland,  and  Chaparral  Formations  ( Derby  etal.  1978).  The 
relative  amounts  of  these  formations  within  the  study  area  are  approximately 
38%,  24%,  and  29%,  respectively.  Other  habitats  occupying  the  remaining  9% 
are  Barren,  Grassland,  Agriculture,  and  Riparian  Series. 

The  Conifer  Forest  Formation  is  found  from  1,600  to  over  2,750  m  elevation. 
Lodgepole  Pine,  Pinus  murrayana;  Sugar  Pine,  P.  lambertiana;  and  White  Fir, 
Abies  concolor,  Series  are  found  in  the  higher  elevations;  Mixed  Conifer,  Coulter 
Pine,  P.  coulteri,  and  Bigcone  Douglas  Fir,  Pseudotsuga  macrocarpa,  Series  at 
the  lower  elevations. 


54  CALIFORNIA  FISH  AND  CAME 

In  the  Woodland  Formation,  Canyon  Oak,  Quercus  chrysolepis,  Series  is 
found  from  1,600  to  2,450  m  elevation  on  southern  exposures  and  from  1,200 
to  1,700  m  on  northern  exposures.  This  series  occupies  the  broad  interface 
between  the  Chaparral  and  Conifer  Forest  Formations.  It  contains  a  heterogene- 
ous mixture  of  canyon  oak;  interior  live  oak,  Q.  wislizenii;  California  black  oak, 
Q.  kelloggii;  and  a  few  scattered  conifers.  The  Black  Oak  Series  is  found  in  more 
mesic  conditions  from  1,450  to  2,100  m  elevation.  The  Interior  Live  Oak  Series 
occurs  in  more  xeric,  lower  elevations,  usually  below  or  in  association  with 
canyon  oak.  Key  black  bear  foods,  such  as  acorns  Quercus  spp.;  western 
chokecherry,  Prunus  virginiana;  coffeeberry,  Rhamnus  californica;  holly-leaved 
cherry,  Prunus  ilicifolia;  and  manzanita,  Arctostaphylos  spp.,  are  present  in  the 
Woodland  Formation. 

The  Chaparral  Formation  is  found  below  1,650  m  and  includes  the  Ceanothus, 
Ceanothus  spp. /Manzanita  Series  and  the  Chamise,  Adenostoma  fasciculatum, 
Series.  The  latter  is  generally  below  1,400  m. 

METHODS  AND  MATERIALS 

Bears  were  captured  in  a  culvert  trap  or  in  Aldrich  foot  snares  (Novick  1979, 
Siperek  1979)  and  immobilized  with  CI-744,  an  experimental  drug  (Stewart, 
Siperek,  and  Wheeler  1980).  Upon  successful  immobilization  of  each  bear, 
pertinent  information  was  collected,  including  weight,  measurements,  and 
health.  A  third  premolar  was  extracted  for  age  determination  (Stoneberg  and 
Jonkel  1966).  Radio  telemetry  collars  (Telonics  of  Mesa,  Arizona)  were  at- 
tached to  nine  bears  between  May  1976  and  December  1977.  Surveillance  of 
bears  was  achieved  by  ground  and  fixed-wing  aircraft  monitoring.  Dens  were 
located  in  early  winter  and  again  in  the  spring.  Information  was  recorded  on 
slope,  aspect,  elevation,  percent  cover,  and  habitat  type.  Monitoring  was  con- 
ducted primarily  from  the  fall  1976  to  the  spring  1978,  but  a  female  bear  was 
monitored  in  winter  1978-79,  also. 

Entrance  and  emergence  of  bears  were  carefully  monitored.  However,  peri- 
ods ranging  from  3  to  16  days  elapsed  between  monitoring  days,  and  the  exact 
dates  of  entrance  and  emergence  were  not  known  in  most  cases.  Thus,  arbitrary 
dates  were  assigned,  the  day  having  the  lowest  maximum  temperature  and 
highest  precipitation  being  selected  for  the  entrance  date  and  the  day  having  the 
highest  maximum  temperature  being  selected  for  the  emergence  date.  This 
procedure  is  based  on  Lindzey  and  Meslow's  (1976)  correlation  of  pre-  and 
post-denning  behavior  with  daily  weather,  principally  temperature  and  precipi- 
tation. 

All  temperature  and  precipitation  data  presented  here  are  those  recorded  by 
the  U.S.  Forest  Service's  ranger  station  at  Mill  Creek  ( Figure  1 ;  elevation:  762  m ) . 
Student's  t-test  was  used  for  statistical  comparisons  (Zar  1974). 

RESULTS 
Denning  information  was  obtained  from  nine  bears.  During  the  winter  of 
1976-77,  three  den  sites  were  located,  and  an  additional  five  sites  were  discov- 
ered the  following  winter.  One  den  was  occupied  by  the  same  bear  (880)  both 
winters.  Throughout  the  winter  of  1976-77,  a  3-year-old  male  (A483)  moved 


BLACK  BEAR  DENNING  CHARACTERISTICS 


55 


several  times  and  apparently  did  not  have  a  permanent  den.  This  bear  fled  upon 
a  close  approach,  and  a  search  of  the  immediate  area  revealed  only  several  day 
beds. 

Denning  Periods 

Most  bears  denned  from  December  until  March,  with  a  range  for  all  bears  of 
late  October  to  early  April  (Table  1 ).  The  mean  denning  period  of  seven  males 
in  the  two  winters  was  93  days,  and  mean  entrance  and  emergence  dates  were 
15  December  and  15  March,  respectively. 


TABLE  1. 

Age,  Sex,  and  Denn 

ng  Dates  of  Black  Bears  in  the  San  Bernardino 

Mountains 

Denning 

Sex 

Age 
(1976) 

Year 
(winter  of) 

Denning 

dates 

period 

Bear  No. 

Entrance 

Emergence 

(days) 

880 

M 

5 

1976-77 

24  Dec. 

1  Mar. 

68 

1977-78 

30  Nov. 

9  Mar. 

100 

882 

M 

2  (est.) 

1977-78 

20  Dec. 

6  Apr. 

108 

883 

M 

8 

1 976-77 3 

10  Dec. 

1  Mar 

82 

884 

M 

10 

1976-77 

24  Dec. 

8  Mar. 

75 

885 

M 

1 

1977-78 

20  Dec. 

24  Mar. 

95 

886 

F 

5 

1976-77' 

31  Oct. 

7  Apr. 

159 

1 977-78 2' 3 

9  Dec. 

24  Mar. 

106 

890 

M 

3(est.) 

1977-78 

20  Dec. 

28  Mar. 

99 

A483 

M 

3 

1 976-77 3 

unknown 

unknown 

— 

A489 

M 

3 

1976-77 3 

unknown 

1  Mar. 

— 

1977-78 

30  Nov. 

24  Mar. 

115 

1  Pregnant 

2  Denned  with  cub/yearlings 

3  Den  site  not  precisely  located 

During  the  winter  of  1976-77,  a  pregnant  female  (886)  entered  her  den  much 
earlier  and  emerged  much  later  than  did  the  three  males  monitored  the  same 
winter  (Table  1 ).  The  following  winter,  the  same  female  was  not  pregnant  but 
had  two  cubs/yearlings.  Her  denning  times  were  similar  to  those  of  five  males 
monitored.  Due  to  the  variation  in  the  denning  behavior  of  bear  886,  data 
collected  on  her  were  omitted  from  calculations  of  denning  period  averages. 

During  the  winter  of  1976-77,  males  entered  their  dens  between  10  and  24 
December  (n  =  3,  x  =  19  December).  During  the  winter  of  1977-78,  denning 
by  males  commenced  from  30  November  to  20  December  (n  =  5,  x  =  12 
December).  The  mean  duration  of  denning  varied  from  75  (n  =  3,  range  68-82) 
to  103  days  (n  =  5,  range  95-115)  for  the  winters  of  1976-77  and  1977-78 
respectively.  This  difference  is  significant  (P<0.01).  Emergence  dates  ranged 
from  1  to  8  March  (n  =  4,  x  =  3  March)  and  9  March  to  6  April  (n  =  5,  x 
~  =  24  March)  for  the  same  winters. 

Den  Characteristics 
Seven  den  sites  were  precisely  located  (Table  2).  Heavy  snows  during  the 
winter  of  1977-78  hampered  efforts  to  find  the  exact  den  site  of  the  female  bear; 
nevertheless,  aerial  and  close  range  ground  tracking  confined  the  site  to  within 
400  m.2 


56 


CALIFORNIA  FISH  AND  CAME 


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BLACK  BEAR  DENNING  CHARACTERISTICS  57 

All  bears  constructed  dens  in  areas  with  minimal  human  disturbance.  These 
sites  had  southern  or  southeastern  exposures  at  elevations  from  1,920  to  2,469 
m  (x  =  2,248  m).  In  the  winter  of  1976-77,  the  female  utilized  a  den  having 
a  northern  exposure  at  an  elevation  of  1,554  m,  only  40  m  below  an  infrequently 
used  fire  road.  The  average  estimated  slope  for  all  dens  was  49  deg  ( range:  30-60 
deg). 

Dens  were  excavated  under  standing  trees  or  huge  granite  boulders.  The  den 
of  bear  880  was  the  only  one  with  the  entrance  on  the  uphill  side  of  the  tree. 
Den  dimensions  recorded  at  three  sites,  along  with  visual  observations  at  other 
sites,  indicate  that  the  size  of  a  den  is  just  large  enough  to  accommodate  the 
bear.  In  these  three  cases,  entrance  height  averaged  50%  less  than  shoulder 
height.  Our  observations  on  den  dimensions  relative  to  bear  size  are  similar  to 
those  of  Craighead  and  Craighead  (1972).  Small  amounts  of  nesting  material 
found  in  dens  consisted  of  shredded  twigs,  bark,  leaves,  or  needles.  r 

The  most  common  habitat  type  for  den  sites  was  the  Canyon  Oak  Series,  or 
this  series  co-dominant  with  a  conifer  series.  Six  of  the  seven  den  sites  at  least 
partially  contained  the  Canyon  Oak  Series.  Understory  vegetation  in  the  immedi- 
ate vicinity  of  the  dens  was  not  dense  and  had  a  sparse,  brushy  appearance.  It 
was  composed  primarily  of  young  canyon  oak,  mountain  mahogany,  Cercocar- 
pus  betuloides,  coffeeberry,  and  to  a  lesser  extent,  ceanothus  and  manzanita. 

Overstory  cover  at  den  sites  usually  was  either  dense  (75%-95%)  or  sparse 
(5%-10%).  Overstory  cover  for  the  surrounding  areas  was  more  uniform, 
ranging  from  20-65%.  Dens  at  lower  elevations  usually  had  very  dense  cover, 
which  in  part  is  due  to  their  location  near  canyon  bottoms,  but  also  is  due  to 
the  nature  of  the  Canyon  Oak  Series.  The  Canyon  Oak  Series  typically  has  a 
dense  overstory  cover  at  lower  elevations,  where  it  is  dominant.  At  higher 
elevations  the  canopy  cover  is  more  open  due  to  the  increased  slope,  elevation, 
and  the  presence  of  conifer  co-dominants. 

DISCUSSION 

Winter  dormancy  allows  black  bears  to  survive  in  regions  having  severe 
climatic  conditions  and  associated  food  scarcity.  Still,  bears  den  in  regions  with 
mild  winters  and  available  food  sources  (LeCount  1980).  Internal  mechanisms 
controlling  this  phenomenon  probably  are  inherent. 

Comparisons  With  Other  Studies 
For  denning  dates  of  black  bears  in  other  states,  most  reports  give  approximate 
ranges  and  a  few  give  mean  denning  dates;  a  direct  comparison  with  our  study 
results  is  difficult.  In  addition,  monitoring  techniques  available  to  us  are  more 
refined  than  techniques  used  by  earlier  researchers,  so  we  were  able  to  obtain 
more  precise  information.  Despite  these  differences,  it  is  evident  that  denning 
periods  in  regions  with  moderate  or  severe  winters  differ  from  those  in  regions 
having  mild  winters  (Table  3).  Also,  bears  denned  later  and  for  a  shorter  period 
in  our  study  area  than  in  all  other  regions  except  North  Carolina.  Even  in  areas 
of  the  West  having  relatively  mild  winters,  such  as  Arizona  and  the  coast  of 
Washington,  bears  have  earlier  and  longer  denning  periods  than  we  recorded. 
Bears  in  this  study  had  an  average  denning  period  of  3  months,  while  investiga- 
tors in  other  regions  report  dormancy  lasting  from  4  to  6  months.  The  corre- 
spondence between  denning  dates  in  our  area  and  North  Carolina  is  most  likely 


58  CALIFORNIA  FISH  AND  GAME 

due  to  similar  climatic  conditions.  North  Carolina  is  reported  to  have  mild 
winters  with  infrequent  snowfall  (Hamilton  and  Marchinton  1980).  Southern 
California  has  only  slightly  more  harsh  winters,  suggesting  that  black  bears 
maintain  a  minimum  denning  period.  Although  factors  governing  denning 
behavior  are  not  fully  understood,  data  obtained  during  this  investigation  support 
those  of  Lindzey  and  Meslow  (1976).  These  researchers  believe  that  cumulative 
effects  of  weather,  principally  precipitation  and  daily  maximum  temperatures 
(rather  than  food  availability),  are  the  most  influential  factors  affecting  the 
timing  of  denning. 

TABLE  3.     Comparison  of  Approximate  Denning  Dates  for  Black  Bears 

State  Entrance  dates  Emergence  dates  Reference 

Alaska Late  October  April  or  later  Erickson  (1965) 

Arizona Early  Nov.  to  Dec.  Late  March  to  early  April       LeCount  (1980) 

Colorado Early  to  mid-Nov.  Mid-March  to  mid-April         Gilbert  (1952) 

Idaho Late  Oct.  to  early  Nov.  Mid  to  late  April  Amstrup  and  Beecham 

(1976) 

Maine Early  December  Early  April  Spencer  (1955) 

Montana Late  October  Mid-April  to  mid-May  Jonkel  and  Cowan 

(1971) 
North  Carolina Early  to  late  Dec.  Late  March  Hamilton  and  Marchinton 

(1980) 
Washington  Late  October  to  Nov.  Mid  to  late  March  Lindzey  and  Meslow 

(1976) 

California  Mid-December  Mid-March  This  study. 

(Range:  late  Oct.*  (Range:  early  March 

to  late  December)  to  early  April) 

*  Represented  by  one  pregnant  female. 

Yearly  Differences  in  Denning 

Even  in  the  same  area,  yearly  differences  in  denning  behavior  were  noted.  The 
winters  of  1 976-77  and  1 977-78  differed  considerably.  In  1 976-77  California  was 
in  the  midst  of  a  drought,  with  low  precipitation  (30  cm  in  the  study  area)  and 
mild  temperatures.  Prior  to  the  first  major  storm  of  the  year  (on  30  December), 
daily  maximum  temperatures  at  Mill  Creek  fluctuated  slightly  (15-23°C)  and 
dropped  slowly  during  a  1 -month  period.  During  this  period,  all  males  denned 
6  to  20  days  before  the  storm  arrived.  At  the  beginning  of  the  severe  winter  of 
1977-78,  which  had  unusually  heavy  precipitation  (87  cm),  daily  maximum 
temperatures  were  erratic  (13-31°C)  and  dropped  relatively  fast  during  the 
month  before  the  first  storm  arrived.  All  bears  denned  17  days  before  to  3  days 
after  the  start  of  the  first  snowfall  on  17  December.  Although  these  two  winters 
were  remarkably  different,  the  onset  of  denning  did  not  differ  significantly  (P 
>  0.05).  However,  storm  activity  appeared  to  initiate  denning  slightly  earlier  in 
1977. 

It  appears  that  duration  of  denning  is  merely  a  function  of  the  time  of  emer- 
gence. We  found  that  emergence  from  denning  is  slightly  earlier  than  other 
studies  report.  These  differences  are  probably  influenced  by  the  warm  Mediter- 
ranean climate,  which  characterizes  southern  California.  During  this  study,  the 
time  of  emergence  was  significantly  different  for  the  two  springs  ( P  <  0.01 ) .  The 
first  spring  of  monitoring  had  little  precipitation  and  all  bears  came  out  during 
a  warm  trend,  when  daily  maximum  temperatures  were  fluctuating  between 


BLACK  BEAR  DENNING  CHARACTERISTICS  59 

12-22°C.  The  following  spring  had  erratic  temperatures  (daily  maxima  9-28°C) 
and  weekly  storms  during  emergence.  This  weather  pattern,  characterized  by  an 
appreciable  amount  of  precipitation  in  late  February  and  March,  probably 
delayed  emergence  of  most  bears  in  1 978.  Lindzey  and  Meslow  ( 1 976)  state  that 
emergence  of  bears  from  dens  is  a  response  to  a  general  warming  trend  during 
a  period  of  increased  day  length.  In  regions  with  mild  climates,  we  believe  the 
severity  of  the  winter  also  influences  the  time  of  emergence  and  the  duration, 
but  not  the  onset,  of  denning. 

Age  and  Sex  Differences  in  Denning  Patterns 

It  is  notable  that  in  the  exceptionally  mild  winter  of  1976-77,  one  subadult 
male  (A483)  either  did  not  den  or  did  not  have  prolonged  denning.  This  is  not 
surprising,  however,  because  Hamilton  and  Marchinton  (1980)  found  in  North 
Carolina  that  an  adult  male  had  the  shortest  period  of  inactivity  and  two  imma- 
ture males  remained  active  throughout  midwinter.  It  is  unclear  whether  this 
behavior  is  related  to  a  specific  sex  or  age  class.  The  time  and  duration  of 
denning  for  two  yearling/subadult  bears  was  not  significantly  different  (P> 
0.05)  from  adult  male  bears,  a  characteristic  also  described  by  Lindzey  and 
Meslow  (1976).  Still,  most  bears  den  regardless  of  the  mildness  of  the  winter. 

The  behavior  of  the  one  female  bear  monitored  throughout  this  study  provides 
some  interesting  comparisons  with  the  behavior  of  other  bears.  When  this  bear 
was  pregnant  in  the  winter  of  1976-77,  she  entered  her  den  in  late  October.  This 
early  denning  was  not  influenced  by  stormy  weather.  During  this  winter,  she 
denned  approximately  2  months  longer  than  male  bears.  Her  denning  times  with 
cubs/yearlings  at  a  different  location  the  following  winter  was  not  significantly 
different  (P>0.05)  from  those  of  five  adult  males  monitored.  When  pregnant 
again  in  the  winter  of  1978-79,  this  female  reoccupied  the  den  she  had  used  in 
1976-77.  She  entered  at  about  the  same  time  as  did  the  males,  but  emerged  later 
as  in  1976-77.  Several  researchers  have  reported  that  pregnant  female  bears 
enter  dens  earlier  and  emerge  later  than  non-pregnant  females  and  males.  Craig- 
head and  Craighead  1972;  Amstrup  and  Beecham  1976;  Lindzey  and  Meslow 
1976;  LeCount  1980;  and  Reynolds  and  Beecham  1980.  However,  Amstrup  and 
Beecham  (1976)  also  felt  that  females  with  yearlings  were  last  to  emerge  from 
dens.  These  reports  and  our  observations  suggest  that  denning  behavior  of 
females  is  quite  variable. 

Den  Site  Characteristics 

Many  authors  have  reported  dens  to  be  under  large  boulders,  fallen  logs, 
dense  vegetation,  bases  of  dead  and  living  trees,  in  excavations  on  hillsides,  and 
in  tree  cavities  several  meters  above  the  ground  (Jonkel  and  Cowan  1971; 
Erickson  1965;  Hamilton  and  Marchinton  1980;  and  LeCount  1980;  and  Pelton, 
Beeman,  and  Eagar  1980).  In  our  study  area,  dens  were  most  frequently  dug 
beneath  large  boulders.  The  ease  of  digging  in  loose  granitic  soil  and  the  abun- 
dance of  large  boulders  in  most  canyons  contribute  to  making  fhis  type  of  den 
readily  available.  Also,  the  stable  micro-climate  to  be  expec'  ;  in  these  dens 
could  make  them  preferable  to  other  types  of  dens. 

Dens  were  generally  located  in  remote  areas  with  steep  terrain,  where  there 
was  little  human  disturbance.  Most  dens  were  located  within  100  m  of  a  creek 
bottom.  This  is  probably  due  to  a  number  of  factors.  For  example,  there  are  many 


60  CALIFORNIA  FISH  AND  GAME 

den  sites  (i.e.,  large  boulders)  near  canyon  bottoms.  Also,  the  Canyon  Oak 
Series  provides  thermal  and  escape  cover,  and  the  availability  of  water  may  be 
important  to  bears  upon  emergence  from  their  dens. 

Importance  of  the  Canyon  Oak  Series 
The  Canyon  Oak  Series,  while  occupying  only  1 6%  of  the  study  area,  was  the 
habTtat  chosen  for  most  dens.  This  series  supplies  most  fall  food  items.  In  re- 
sponse to  the  phenological  progression  of  coffeeberry  and  various  acorn  crops 
during  the  fall  months,  bears  often  were  found  from  middle  to  high  elevations 
in  their  normal  home  ranges.  Coffeeberry  and  acorns  are  the  most  important  fall 
foods  ( Boyer  1 976 ) .  Most  bears  denned  at  significantly  ( P  <  0.001 )  higher  eleva- 
tion (x  =  2,248  m)  than  where  they  were  active  in  previous  seasonal  ranges 
(Novick  1979). 

We  suggest  that  the  moderate  to  dense  overstory  cover  provided  by  the 
Canyon  Oak  Series  keeps  the  den  site  cooler  than  it  would  be  in  more  exposed 
locations.  On  southern  exposures  below  2,400  m,  snow  does  not  accumulate. 
A  well-developed  canopy  may  help  compensate  for  warm  winter  temperatures, 
particularly  at  lower  denning  elevations,  thus  meeting  thermal  requirements  for 
denning. 

Factors  influencing  den  site  preferences  of  black  bears  are  complicated.  There 
appears  to  be  a  complex  relationship  between  available  den  sites  which  meet 
their  thermal  requirements  (high  elevation  or  moderate  to  dense  overstory 
cover),  areas  with  minimal  human  disturbance  (remote  areas  with  steep  ter- 
rain), and  the  location  of  fall  food  items  (coffeeberry  and  acorns)  within  their 
home  ranges.  The  Canyon  Oak  Series  meets  these  conditions  and  probably  is 
the  most  important  habitat  type  for  black  bear  den  locations  in  the  San  Bernar- 
dino Mountains. 

ACKNOWLEDGMENTS 

We  thank  E.  T.  Roche,  professor,  California  State  Polytechnic  University, 
Pomona,  for  the  preparation  and  sectioning  of  teeth  for  age  determination.  We 
are  especially  grateful  to  the  San  Bernardino  and  Riverside  County  Fish  and 
Game  commissions  and  the  Southern  Council  of  Conservation  Clubs  for  partially 
funding  this  research.  The  assistance  of  the  U.S.  Forest  Service  and  the  California 
Department  of  Fish  and  Game  is  very  much  appreciated.  K.  Boyer,  biologist,  U.S. 
Forest  Service,  and  L.  Puckett,  biologist,  California  Department  of  Fish  and 
Game,  are  thanked  for  critically  reading  this  manuscript.  Finally,  we  thank  our 
wives  and  the  numerous  other  people  who  have  helped  throughout  this  study. 

REFERENCES 

Amstrup,  S.C.,  and  J.  Beecham.  1976.  Activity  patterns  of  radio-collared  black  bears  in  Idaho.  J.  Wildl.  Manage., 

40<2):34C-348. 
Boyer,  K.B.  1976.  Food  habits  of  black  bears  (Ursus  americanus)  in  the  Banning  Canyon  area  of  San  Bernardino 

National  Forest.  M.S.  Thesis.  Calif.  State  Polytech.  Univ.,  Pomona.  63p. 

Burghduff,  A.E.  1935.  Black  bears  released  in  southern  California.  Calif.  Fish  Game,  21(1):83-84. 

Craighead,  F.C.,  and  ).J.  Craighead.  1972.  Grizzly  bear  prehibernation  and  denning  activities  as  determined  by 
radiotracking.  Wildl.  Monogr.,  32.  35p. 

Derby,  J.,  I.  Parker,  T.  Paysen,  V.  Bleich,  H.  Black,  J.  Mincks,  and  B.  Harvey.  1978.  Vegetation  classification  system 
for  southern  California.  U.S.  Forest  Service  and  California  Dept.  of  Fish  and  Game,  Interagency  Publ.  44p. 

Erickson,  AW.  1965.  The  black  bear  in  Alaska — its  ecology  and  management.  Alaska  Dept.  of  Fish  and  Game.  Fed. 
Aid  in  Wildl.  Restoration  Project  Report.  Project  W-6R-5,  Work  Plan  7.  19p. 


BLACK  BEAR  DENNING  CHARACTERISTICS  61 

Gilbert,  D.L.  1952.  Bear  studies.  Colo.  Came  and  Fish  Dept.  Fed.  Aid  Quart.  Jan.:26-31. 

Hamilton,  R.J.,  and  R.L.  Marchinton.  1980.  Denning  and  related  activities  of  black  bears  in  the  coastal  plain  of  North 

Carolina.  Pages  121  to  126  inC  Martinka,  and  K.L.  McArthur,  eds.  Bears — their  biology  and  management. 

February  1977.  Kalispell,  Mt. 

Jonkel,  C.J.,  and  I.  McT.  Cowan.  1971.  The  black  bear  in  the  spruce-fir  forest.  Wildl.  Monogr.,  27.  57p. 

LeCount,  A.L  1980.  Some  aspects  of  black  bear  ecology  in  the  Arizona  chaparral.  Pages  175  to  179//7C.J.  Martinka, 
and  K.L.  McArthur,  eds.  Bears — their  biology  and  management.  February  1977.  Kalispell,  Mt. 

Lindzey,  F.C.,  and  E.C.  Meslow.  1976.  Winter  dormancy  in  black  bears  in  southwestern  Washington.  J.  Wildl. 
Manage.,  40(3):408-^15. 

Novick,  H.J.  1979.  Home  range  and  habitat  preferences  of  black  bears  ( Ursus  americanus)  in  the  San  Bernardino 
Mountains  of  southern  California.  Thesis.  Calif.  State  Polytech.  Univ.,  Pomona.  1-58. 

Pelton,  M.R.,  L.E.  Beeman,  and  D.C.  Eagar.  1980.  Den  selection  by  black  bears  in  the  Great  Smoky  Mountains 
National  Park.  Pages  149  to  151  m  C.|.  Martinka  and  K.L.  McArthur,  eds.  Bears — their  biology  and  manage- 
ment. February  1977.  Kalispell,  Mt. 

Poelker,  R.J.,  and  H.D.  Hartwell.  1973.  The  black  bear  of  Washington.  Wash.  State  Game  Dept.  Biol.  Bull., 

(14):1-180. 
Reynolds,  D.,  and  J.  Beecham.  1980.  Home  range  activities  and  reproduction  of  black  bears  in  west-central  Idaho. 

Pages  181  to  190  inC\.  Martinka,  and  K.L.  McArthur,  eds.  Bears — their  biology  and  management.  February 

1977.  Kalispell,  Mt. 

Siperek,  J.M.  1979.  Physical  characteristics  and  blood  analysis  of  black  bears  (Ursus  americanus)  in  the  San 
Bernardino  Mountains  of  southern  California.  Thesis.  Calif.  State  Polytech.  Univ.,  Pomona.  1-63. 

Spencer,  H.E.  1955.  The  black  bear  and  its  status  in  Maine.  Maine  Dep.  Inland  Fisheries  and  Game,  Game  Div. 
Bull.,  (4):1-55. 

Stewart,  G.R.,  |.M.  Siperek,  and  V.R.  Wheeler.  1980.  Use  of  the  cataleptoid  anesthetic  CI-744  for  chemical  restraint 
of  black  bears.  Pages  57  to  61  otC.J.  Martinka,  and  K.L.  McArthur,  eds.  Bears — their  biology  and  management. 
February  1977.  Kalispell,  Mt. 

Stoneberg,  R.P.,  and  C.J.  Jonkel.  1966.  Age  determination  of  black  bears  by  cementum  layers.  J.  Wildl.  Manage., 
30(2):41 1^*14. 

Zar,  J.H.  1974.  Biostatistical  analysis.  Prentice-Hall,  Inc.,  Englewood  Cliffs,  N.J.  620p. 


62  CALIFORNIA  FISH  AND  GAME 

NOTES 

UPDATE  OF  THE  ESTIMATED  MORTALITY  RATE  OF 
ENGRAULIS  MORDAWH  SOUTHERN  CALIFORNIA 

INTRODUCTION 

The  central  subpopulation  of  northern  anchovy,  Engraulis  mordax,  found 
along  the  west  coast  of  North  America  from  0°  N  to  38°  N  and  concentrated  in 
Southern  California  Bight,  is  subject  to  an  extensive  reduction  and  bait  fishery. 
The  management  plan  for  the  U.S.  anchovy  fishery  ( Pacific  Fishery  Management 
Council  1978)  required  by  the  Fishery  Conservation  and  Management  Act  of 
1 976  ( Public  Law  94-265 )  is  based  on  existing  knowledge  of  population  parame- 
ters. Annual  mortality  {a)  and  instantaneous  total  mortality  (Z)  are  two  of  the 
parameters  used  in  the  plan  and  are  based  on  estimates  of  MacCall  (1974). 
Using  the  catch  curve  analysis  method  developed  by  Chapman  and  Robson 
(Chapman  and  Robson  1960;  Robson  and  Chapman  1961 ),  MacCall  arrived  at 
an  average  Z  of  1.09  and  an  average  a  of  66.5%  for  the  central  subpopulation. 
I  have  updated  the  estimate  of  Z  by  including  more  recent  data  and  have 
examined  the  time  series  for  any  recent  changes  or  long-term  trends  in  the 
parameter  values. 

MATERIALS  AND  METHODS 
The  analysis  is  based  on  data  reports  of  the  California  Department  of  Fish  and 
Game,  Pelagic  Fish  Investigations  Sea  Survey  Project,  from  October  1966  to 
November  1979  (Mais  1969a,  b;  1971  a,  b,  c,  1972, 1973,  1974, 1975, 1976, 1977, 
1978,  1979,  1980).  Also  included  are  data  from  one  cruise  in  1980  (K.  Mais, 
Marine  Biologist,  Calif.  Dept.  Fish  and  Game,  pers.  commun.).  Catch  curves  are 
derived  from  year-class  frequencies  of  anchovies  in  the  midwater  trawl  stations. 
Annual  mortality  rates  and  the  corresponding  instantaneous  total  mortality  rates 
are  calculated  for  each  cruise  using  the  Chapman-Robson  method  as  applied  by 
MacCall  (1974).  The  best  estimate  of  Z  is  then  the  mean  value  over  all  the 
cruises  and  a  is  determined  from  that  mean  value  (Table  1). 

RESULTS  AND  DISCUSSION 

The  estimate  of  Z  is  calculated  to  be  0.97  (s  =  0.38)  which  corresponds  to 
a  62.1%  annual  mortality.  Although  the  assumptions  necessary  to  use  the  Chap- 
man-Robson analysis  are  poorly  satisified  due  to  large  fluctuation  in  recruitment 
and  the  likelihood  of  increasing  mortality  with  age,  these  deficiencies  are  offset 
by  averaging  the  values  of  Z  over  the  14-year  period. 

When  the  values  of  Z  from  each  cruise  are  plotted  against  time  (Figure  1 ), 
the  between  sample  variance  becomes  apparent  and  may  be  due  to  high  varia- 
bility in  recruitment  of  year  classes  or  relative  year  class  strengths.  A  10-year 
decreasing  trend  in  the  values  of  Z  is  dramatically  reversed  after  1976  (Figure 
1 ).  This  4-year  increase  in  Z  since  1976  is  concerning,  since  it  coincides  with  a 
sharp  decrease  of  older  anchovies  in  the  commercial  catch  (J.  Sunada,  Marine 
Biologist,  Calif.  Dept.  Fish  and  Game,  pers.  commun.)  and  a  decline  in  the  total 
U.S.  catch.  Studies  have  not  shown,  however,  if  the  manifestations  are  natural 
fluctuations  in  the  anchovy  population  or  responses  to  outside  stimuli  such  as 
environmental  change,  predators,  or  competitors. 


MORTALITY  RATE  ESTIMATES 


63 


TABLE  1.     Mortality  Rate  Estimates  Based  on  Sea  Survey  Data  for 

Southern  California. 


Number 
Cruise  sampled ' 

66A8 106 

67A2 60 

68A4 145 

68A8 128 

68A9 55 

69A6 105 

69A8 146 

69A1 1  127 

70A1  92 

70A4 145 

70A7 1 1 1 

71 A1  85 

71  A3 92 

71A7 162 

72A3 98 

72A9 1 1 5 

73A2 169 

73A3 156 

Number 
Cruise  sampled  * 

73A8 236 

74A3 246 

74A9 240 

75A1  243 

75A2 90 

75A5 73 

75A6 289 

76A3 307 

76A4B 155 

76A7 123 

76A9 216 

77A3 277 

77A6 81 

77A13 167 

78A2 57 

78A3 174 

79A1  133 

79A2 91 

80A1  100 

Instantaneous 
mortality 
rated) 

1966-1976 0.97 

1976-1980 0.97 

1966-1980 0.97 

*  Number  of  fish  2  years  old  or  older. 


Annual 

Instantaneous 

mortality 

mortality 

rate  (a) 

rate  (Z) 

.60 

.90 

.66 

1.07 

.53 

.75 

.69 

1.17 

.62 

.95 

.66 

1.07 

.68 

1.13 

.72 

1.26 

.75 

1.35 

.73 

1.30 

.67 

1.11 

.71 

1.20 

.61 

.94 

.66 

1.06 

.60 

.90 

.84 

1.79 

.75 

1.36 

.56 

.82 

Annual 

Instantaneous 

mortality 

mortality 

rate  (a) 

rate  (Z) 

.59 

.90 

.55 

.79 

.57 

.84 

.55 

.80 

.54 

.76 

.63 

.98 

.52 

.73 

.41 

.53 

.48 

.64 

.45 

.60 

.40 

.50 

.44 

.58 

.55 

.79 

.47 

.64 

.62 

.94 

.46 

.61 

.72 

1.26 

.64 

1.07 

.84 

1.83 

Annual 

Standard 

mortality 

deviation 

rate  (a) 

0.28 

0.621 

0.42 

0.621 

0.31 

0.621 

64 


CALIFORNIA  FISH  AND  CAME 


1967      1968     1969     1970      1971      1972     1973      1974     1975     1976     1977     1978     1979     I960 

YEARS 

FIGURE  1 .     The  values  of  instantaneous  mortality  rate  are  plotted  for  each  of  the  sea  survey  cruises 
for  the  period  October  1966  to  February  1980. 

ACKNOWLEDGMENTS 
I  wish  to  thank  Alec  MacCall  (CFG),  Kenneth  Mais  (CFG),  and  Gary  Stauffer 
(NMFS)  for  their  help  in  this  study.  Funds  were  provided  by  the  Southwest 
Fisheries  Center,  National  Marine  Fisheries  Service. 

REFERENCES 

Chapman,  D.G.,  and  D.S.  Robson.  1960.  The  analysis  of  a  catch  curve.  Biometrics,  16:  354-368. 

MacCall,  A.D.  1974.  The  mortality  rate  of  Engraulis  mordax  in  Southern  California.  Mar.  Res.  Comm.,  Calif.  Coop. 

Oceanic  Fish.  Invest.  Rept.  17:  131-135. 
Mais,  K.F.  1969a.  California  Department  of  Fish  and  Game  fisheries  resources  sea  survey  cruises,  1966.  Calif.  Mar. 

Res.  Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (16):  1-85. 
19696.  California  Department  of  Fish  and  Came  fisheries  resources  sea  survey  cruises,  1967.  Calif.  Mar. 

Res.,  Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (17):  1-106. 
1971a.  California  Department  of  Fish  and  Came  fisheries  resources  sea  survey  cruises,  1968.  Calif.  Mar. 

Res.  Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (18):  1-181. 
19716.  California  Department  of  Fish  and  Came  fisheries  resources  sea  survey  cruises,  1969.  Calif.  Mar. 

Res.  Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (19):  1-131. 
1971c.  California  Department  of  Fish  and  Game  fisheries  resources  sea  survey  cruises,  1970.  Calif.  Mar. 

Res.  Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (20):  1-139. 
1972.  California  Department  of  Fish  and  Game  fisheries  resources  sea  survey  cruises,  1971 .  Calif.  Mar.  Res. 

Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (21):  1-132. 
1973.  California  Department  of  Fish  and  Game  fisheries  resources  sea  survey  cruises,  1972.  Calif.  Mar.  Res. 

Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (22):  1-88. 
1974.  California  Department  of  Fish  and  Game  fisheries  resources  sea  survey  cruises,  1973.  Calif.  Mar.  Res. 

Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (23):  1-113. 
1975.  California  Department  of  Fish  and  Game  fisheries  resources  sea  survey  cruises,  1974.  Calif.  Mar.  Res. 

Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (24):  1-86. 
1976.  California  Department  of  Fish  and  Game  fisheries  resources  sea  survey  cruises,  1975.  Calif.  Mar.  Res. 

Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (25):  1-122. 
1977.  California  Department  of  Fish  and  Game  fisheries  resources  sea  survey  cruises,  1976.  Calif.  Mar.  Res. 


Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (26):  1-131. 


NOTES  65 

1978.  California  Department  of  Fish  and  Came  fisheries  resources  sea  survey  cruises,  1977.  Calif.  Mar.  Res. 

Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (27):  1-126. 

1979.  California  Department  of  Fish  and  Came  fisheries  resources  sea  survey  cruises,  1978.  Calif.  Mar.  Res. 

Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (28):  1-52. 

1980.  California  Department  of  Fish  and  Game  fisheries  resources  sea  survey  cruises,  1979.  Calif.  Mar.  Res. 


Comm.,  Calif.  Coop.  Oceanic  Fish.  Invest.,  Data  Rept.  (29):  1-60. 
Pacific  Fishery  Management  Council.  1978.  Northern  anchovy  fishery  management  plan.  Federal  Register  43  ( 141 ): 

31655-31783. 
Robson,  D.  S,  and  DC  Chapman.  1961.  Catch  curves  and  mortality  rates.  Amer.  Fish.  Soc.  Trans,  90(2):  181-189. 

— Doyle  Hanan,  California  Department  of  Fish  and  Came,  c/o  Southwest  Fisher- 
ies Center,  P.O.  Box  271,  La  Jolla,  California  92038.  Accepted  for  publication 
July  1980. 


FIRST  RECORD  OF  DEXTRALITY  IN  THE  CALIFORNIA 
TONGUEFISH  SYMPHURUS  ATRICAUDA,  WITH  A  SEC- 
OND REPORT  OF  AMBICOLORATION. 

The  California  tonguefish,  Symphurus  atricauda  (Jordan  and  Gilbert),  is  a 
member  of  the  family  Cynoglossidae,  which  are  characteristically  sinistral  (left- 
eyed).  A  dextral  (right-eyed)  California  tonguefish  was  captured  during  a  trawl- 
ing study  near  Long  Beach,  California  (lat  33°  43'  16"  N,  long  118°  09'  09"  W). 
The  specimen  was  collected  at  10  fathoms  using  an  8-ft  otter  trawl  while  aboard 
the  NAUTILUS.  The  trawl  was  taken  between  1750  and  1810  hours  on  28  January 
1979.  The  specimen  had  a  standard  length  of  1 17  mm,  as  measured  from  the  tip 
of  the  snout  to  the  end  of  the  fleshy  portion  of  the  tail.  It  had  a  damp  weight 
of  25.09  g  and  a  displaced  volume  of  18.5  ml.  Radiographs  indicated  no  unusual 
skeletal  features  other  than  that  of  an  apparent  mirror  image  of  normal,  left-eyed 
specimens  (Figure  1  top).  Fin  formulas  for  this  specimen  were:  D98,  A80,  with 
13  caudal  and  4  pelvic  fin  rays.  Caudal  fin  ray  count  was  determined  using  the 
method  described  by  Menon  (1977).  There  were  120  scales  in  a  longitudinal 
series  from  the  head  to  the  tail,  and  49  scales  in  a  maximum  dorsal  to  ventral 
diagonal  cross  series.  All  counts  were  within  normal  limits  for  the  species  (Jor- 
dan and  Evermann  1 896) .  The  liver  was  on  the  left  side  of  the  abdominal  cavity, 
and  the  intestine  was  on  the  right.  Therefore,  the  viscera  retained  the  typical 
orientation  for  the  family  Cynoglossidae.  The  specimen  is  now  in  the  California 
State  University  Long  Beach  fish  collection  (collection  #  790128). 

Developmental  anomalies  in  the  California  tonguefish  are  apparently  rare.  The 
only  previous  record  of  an  anomaly  was  that  of  a  partially  ambicolored  speci- 
men (Haaker  1973).  I  received  a  California  tonguefish  which  also  shows  partial 
ambicoloration.  The  specimen  had  pigment  on  less  than  half  of  the  posterior 
blind  side  (Figure  1  bottom).  The  pigment  was  continuous  and  of  the  same 
density  as  that  of  the  eyed  side.  This  specimen  was  trawled  off  the  Dume  Canyon 
number  1  station  of  the  Southern  California  Coastal  Water  Research  Project  (lat 
33°  59'  30"  W,  long  118°  49'  30"  N),  on  15  November  1979.  The  only  other 
reports  of  ambicoloration  in  tonguefishes  are  for  S.  plagiusa  (Dawson  1962, 
Dahlberg  1970a),  and  S.  diomedianus  (Moe  1968). 


66  CALIFORNIA  FISH  AND  GAME 

A  dextral  5.  atricauda  was  briefly  mentioned  by  Mahadeva  (1956  unpubl.), 
but  the  specimen  was  undescribed  and  no  data  provided.  Therefore,  I  feel  that 
this  is  the  first  documented  report  of  reversal  for  the  California  tonguefish. 
Reversal  in  the  Cynglossidae  is  not  common.  The  first  incidence  of  reversal 
reported  was  for  S.  plagiusa  (Linneaus)  trapped  off  Louisiana  by  Chabanaud 
(1948).  A  second  reversed  S.  plagiusa  was  trawled  from  Duplin  River,  Georgia 


FIGURE  1 .  The  specimen  of  Symphurus  atricauda  at  the  top  of  the  photograph  is  the  first  reported 
instance  of  reversal  for  the  species.  The  center  individual  shows  the  normal  sinistral  condition. 
The  bottom  individual  is  the  second  reported  incidence  of  ambicoloration  for  the  species. 

(Dahlberg  1979/?).  The  only  other  record  of  reversal  in  the  tonguefishes  was  a 
reversed  and  partially  ambicolorate  S.  diomedianus  (Coode  and  Bean),  from 
the  Gulf  coast  of  Florida  (Moe  1968).  Because  both  Chabanaud's  and  Moe's 
specimens  lacked  pelvic  fins,  they  hypothesized  that  reversal  may  inhibit  the 
development  of  pelvic  fins  (Chabanaud  1948,  Moe  1968).  However,  Dahlberg 
(1970/?)  reported  that,  "The  presence  of  reversed,  but  otherwise  normal  pelvic 
fins  in  my  specimen  does  not  support  their  conclusion."  Normal  California 
tonguefish  have  only  one  pelvic  fin,  and  it  is  found  on  the  left  (-eyed)  side.  In 
this  reversed  specimen  the  pelvic  fin  was  on  the  right  (-eyed)  side;  the  fin  was 
otherwise  normal.  This  also  does  not  support  the  pelvic  fin  inhibition  theory. 
There  is  insufficient  evidence  at  this  time  to  indicate  any  true  connection  with 
the  inhibition  of  pelvic  fin  development  and  reversal. 

Norman  (1934)  and  Dawson  (1962)  both  suggested  that  the  Cynoglossids 
and  Soleids  are  highly  specialized.  Work  by  Dawson  (1962),  and  Haaker  and 
Lane  ( 1 973 )  suggested  that  the  more  primitive  groups  of  flatfishes  show  a  higher 
occurrence  of  anomalies  than  those  groups  which  are  more  specialized.  If  this 
were  true,  Cynoglossids  and  Soleids  would  show  a  lower  incidence  of  anomalies 


*   NOTES  67 

than  the  more  primitive  flatfishes,  such  as  the  Bothids.  This  trend  was  supported 
by  Dawson  ( 1 962 ) ,  who  found  a  higher  incidence  of  anomalies  for  Bothids  than 
for  Cynoglossids  and  Soleids.  Haaker  and  Lane  (1973)  reported  a  higher  occur- 
rence of  anomalies  for  the  bothid,  Paralichthys  californicus,  than  for  the  pleuro- 
nectid,  Hypsopsetta  quttulata,  and  cited  this  as  evidence  that  the  Pleuronectids 
are  more  specialized  than  Bothids.  The  rarity  of  reported  anomalies  in  the 
tonguefishes  appear  to  further  support  the  suggestion  that  Cynoglossids  are 
indeed  highly  specialized. 

ACKNOWLEDGMENTS 

My  thanks  to  R.  Bray  for  his  help  and  advice  during  this  study.  Also,  may 
appreciation  to  the  Southern  California  Coastal  Water  Research  Project  for  the 
donation  of  the  ambicolored  specimen. 

REFERENCES 

Chabanaud,  P.  1948.  Description  d'un  Symphurus  totalement  inverse'.  France,  Soc.  Zool.,  Bull.  73:134-136. 
Dahlberg,  M.D.  1970a.  Frequencies  of  abnormalities  in  Georgia  estuarine  fishes.  Am.  Fish.  Soc.,  Trans.,  99(1 1:95- 

97. 
19706.  A  completely  reversed  blackcheek  tonguefish,  Symphurus  plagiusa,  from  Duplin,  Georgia.  Chesa- 
peake Sci„  1970(21:260-261. 
Dawson,  C.E.  1962.  Notes  on  anomalous  american  Heterosomata  with  descriptions  of  five  new  records.  Copeia, 

1962(1  ):138-146. 
Haaker,  PL.  1973.  Ambicoloration  in  some  California  flatfishes.  Calif.  Fish  Game,  59(4):299-304. 
Haaker,  P.L.,  and  E.D.  Lane.  1973.  Frequencies  of  anomalies  in  a  Bothid  (Paralichthys  californicus)  and  a  Pleuro- 

nectid  (Hypsopsetta  guttu/ata)  flatfish.  Copeia,  1973(11:22-25. 
Jordan,  D.S.,  and  B.W.  Evermann.  1896.  The  fishes  of  North  and  Middle  America.  U.S.Nat.  Mus.,  Bull.  47(31:2707- 

2708. 
Mahadeva,  N.  A  review  of  the  tonguefishes  of  the  Eastern  Pacific,  with  descriptions  of  six  new  species.  Los  Angeles, 

CA:  Univ.  of  California,  Los  Angles;  1956.  272p.  Dissertation. 
Menon,  A.G.K.  1 977.  A  systematicd  monograph  of  the  tonguesoles  of  the  Genus  Cynoglossus  Hamilton-Buchanon, 

(Pisces:  Cynoglassidae).  Smithsonian  Contributions  to  Zoology.  No. 238,  129pp. 
Moe,  M.A.,  Jr.  1968.  A  reversed  partially  ambicolorate  tonguesole,  Symphurus  diomedianus,  from  the  Gulf  of 

Mexico.  Copeia,  1968(1 1:1 72. 
Norman,  JR.  1934.  A  systematic  monograph  of  the  flatfishes  (Heterosomata).  Br.  Mus.  Nat.  Hist.,  London.  459p. 

— Eduard  L.  Telders;  Department  of  Biology,  California  State  University,  Long 
Beach;  Long  Beach,  California  90840.  Accepted  for  publication  September 
1980. 


68  CALIFORNIA  FISH  AND  CAME 

BOOK  REVIEWS 

The  George  Reserve  Deer  Herd 

By  Dale  R.  McCullough;  University  of  Michigan  Press,  Ann  Arbor,  Ml;  1979;  271  p.;  $16.00. 

Seldom  are  wildlife  managers  and  other  applied  ecologists  offered  such  a  detailed  data  set, 
analysis,  and  evaluation  as  that  provided  in  this  volume  by  Dr.  McCullough.  The  book  is  prefaced 
by  the  author  as  a  progress  report  synthesizing  investigations  conducted  on  a  southern  Michigan 
white-tailed  deer  population  since  the  1930's.  Despite  the  continuing  nature  of  experiments  on  the 
study  area,  results  and  conclusions  discussed  in  the  book  provide  an  innovative  approach  to 
management  concepts  for  large  ungulate  through  detailed  analysis  of  empirical  data.  This  book  is 
clearly  not  intended  for  the  layman,  but,  as  stated  by  the  author,  "there  is  no  reason  why  any 
intelligent  person  could  not  comprehend  the  material." 

The  basis  for  the  research  reported  on  by  McCullough  involves  combining  white-tailed  deer 
ecology  with  a  conceptual  ecosystem  model  to  demonstrate  model  function  and  describe  the 
George  Reserve  population.  Specifically,  deer  population  dynamics  are  illustrated  through  the  use 
of  models  for  production,  recruitment,  mortality,  and  yield.  Although  data  were  obtained  from 
white-tailed  deer,  the  ecosystem  hypothesis  tested  produced  conclusions  applicable  to  other  sub- 
climax  ungulate  species.  The  author  presents  an  excellent  discussion  of  the  theory  of  carrying 
capacity  and  problems  associated  with  usages  of  the  term  in  wildlife  management. 

The  George  Reserve  Deer  Herd  emphasizes  sport  hunting  as  the  primary  management  tool  for 
K-selected  ungulate  species.  McCullough  presents  challenges  to  a  number  of  intuitive  beliefs  related 
to  manipulating  sex  and  age  structure  and  effects  on  population  yield  and  stability.  An  excellent 
analysis  of  the  integration  of  social  and  biological  factors  influencing  sport  hunting  is  presented  in 
the  chapter  dealing  with  management  concepts.  The  technically  sound,  thorough  evaluation  of  the 
subject  matter  makes  this  book  a  valuable  contribution  in  the  field  of  large  mammal  ecology  and 
management. —  Terry  M.  Mansfield 

How  to  Build  a  Freshwater  Artificial  Reef — Second  Edition 

By  Eric  D.  Prince,  O.  Eugene  Moughan,  and  Paul  Brouha;  Sea  Grant  at  Virginia  Tech,  Extension  Division, 

Virginia  Polytechnic  and  State  University,  Blacksburg,  VA  24061;  14  pp;  illustrated;  $1.00. 

This  concisely  written  pamphlet  decribes  some  of  the  problems  which  might  be  encountered 
during  the  emplacement  of  artificial  freshwater  reefs.  It  offers  guidelines  for  various  construction 
phases  and  summarizes  the  state-of-the-art  literature  on  the  subject.  The  authors  touch  on  such 
pertinent  topics  as  the  physical  and  biological  need  for  reefs,  various  legal  considerations  and  how 
they  apply  to  various  levels  of  government,  and  benefits,  costs,  and  longevity  of  various  reef  types. 

Photographs  and  illustrations  of  reefs  made  from  scrap  tires,  brush,  wooden  stakes,  vitrified  clay 
pipe,  and  other  materials  clearly  demonstrate  the  practical  applications  of  these  materials. 

Although  artificial  reefs  are  admittedly  not  a  panacea  to  every  fisheries  management  problem,  this 
publication  will  prove  useful  to  private  farm  pond  owners  and  professional  fisheries  biologists 
alike. — Larry  E.  Week 

The  Black  Bear  in  Modern  North  America 

By  Dale  Burk;  Boone  and  Crockett  Club  and  the  Amwell  Press,  Clinton,  New  Jersey;  1979;  300  p. 

Black  bear  management  in  the  last  2  decades  has  gone  from  solving  local  pest  problems  to 
concerned  international  cooperation  in  the  interest  of  the  species.  This  book  is  the  proceedings  of 
a  workshop  on  bear  status  and  management  attended  by  bear  biologists  from  the  United  States, 
Canada,  and  Mexico.  Workshop  chairman  Alan  Stoken  cites  a  threefold  purpose:  to  review  the  status 
of  the  bear,  to  develop  policy  statements,  and  to  publish  the  transcripts.  The  result  is  a  reference 
for  the  desk  of  bear  researchers,  laymen,  and  professionals. 

In  the  first  portion  of  the  book,  Editor  Dale  Burk  has  put  together  an  orderly  geopolitical  arrange- 
ment of  regional  statements  on  the  bear's  status.  The  entertaining  comparative  discussion  of  brown 
and  black  bears  is  followed  by  the  most  important  and  final  section.  Resource  managers  are  led 
regionally  into  the  complex  environmental  interrelationships  within  which  the  species  must  be 
managed.  Workshop  participants  suggest  management  action  based  on  range  environmental  condi- 
tions rather  than  on  isolated  political  districts. 

As  with  their  earlier  bighorn  sheep  publication,  the  sponsoring  Boone  and  Crockett  Club  has  done 
wildlife  a  service. — Larry  Sitton 


REVIEWS  69 

Estimation  of  Density  from  Line  Transect  Sampling  of  Biological  Populations 

By  Kenneth  P.  Burnham,  David  R.  Anderson,  and  Jeffrey  L.  Lake;  The  Wildlife  Society,  Inc.,  Washington, 

D.C.;  202  pp.;  $4.00. 

The  difficulty  of  accurately  estimating  the  density  of  animals  has  led  to  the  development  of  a 
variety  of  estimators.  This  Wildlife  Monograph  proposes  line  transect  sampling  with  a  Fourier  series 
estimator  providing  the  probability  density  function. 

The  authors  have  the  commendable  goals  of  combining  theory  with  practice;  providing  statisti- 
cians with  the  underlying  theory  of  the  methods  presented  and  biologists  with  reliable,  practical 
procedures  for  design,  execution,  and  analysis  of  field  studies.  Unfortunately  the  task  of  including 
material  at  a  level  meaningful  to  each  discipline  has  forced  the  structure  of  the  Monograph  into 
numerous  parts  and  appendixes  and  necessitated  the  inclusion  of  a  Reader's  Guide  which  recom- 
mends which  parts  to  read  for  practitioners  of  the  different  levels  of  the  appropriate  professions. 
Perhaps  it  is  time  to  acknowledge  that  most  individuals  trained  or  employed  as  biologists  are  not, 
as  the  authors  assumed  in  their  preface,  ".  .  .  familiar  with  such  concepts  as  random  variables, 
estimators,  sampling  variance,  confidence  intervals,  bias,  and  chi-square  test  statistics".  As  moral 
philosophers  have  pointed  out,  "you  can't  make  an  is  from  an  ought". 

Nevertheless,  the  Monograph  is  a  particularly  comprehensive  reference  on  line  transect  sampling 
and  with  the  guidance  provided  by  this  treatise,  biologists  seeking  to  estimate  the  density  of  objects 
in  a  sampled  area  should  be  able  to  make  conceptually  sound  and  explicitly  accurate  density 
estimates. 

There  are  occasional  lapses  in  this  generally  lucid  work  which  are  confusing,  such  as  the  para- 
graphs on  page  14  under  the  hearing  "Assumptions"  which  are  promptly  followed  by  the  disclaimer, 
"These  are  not  to  be  considered  as  assumptions."  Also  the  choice  of  print  styles  and  parameter 
names  might  have  been  better  coordinated.  This  reviewer  found  it  extremely  slow  going  when  trying 
to  read  a  paragraph  in  which  the  parameter  "a"  was  discussed,  because  the  article  "a"  occurred 
equally  often  and  was  only  distinguishable  by  careful  attention  to  context. 

These  quibbles  aside,  this  Monograph  is  going  to  be  extremely  useful  to  biologists  who  are 
increasingly  dealing  with  nongame  and  endangered  species  for  which  the  change-in-ratio  methods 
which  depend  on  harvest  are  difficult  to  apply. — Earle  W.  Cummings 

Salmon  Fishers  of  the  Columbia 

By  Courtland  L.  Smith;  Oregon  State  University  Press,  Corvallis,  OR;  128  pp;  illustrated;  $15.00. 

At  times,  trying  to  read  Salmon  Fishers  of  the  Columbia  was  as  difficult  as  staying  awake  during 
a  post-lunch  lecture  in  a  warm  hall.  Dullness  aside,  this  fairly  short  book  provides  a  good  overview 
of  the  history  of  the  Columbia  River  salmon  fishery,  from  aboriginal  times  to  the  early  1970's. 

Drawing  on  the  historical  record,  anthropologist-author  Courtland  Smith  has  documented  the  rise 
and  decline  of  the  canned  salmon  industry,  with  special  emphasis  on  the  competition  within  and 
between  different  user  groups  that  has  existed  from  almost  the  beginning  of  the  industry.  The 
numerous  catch  and  pack  statistics,  while  not  the  highlight  of  the  book,  are  necessary  for  understand- 
ing of  the  history  of  this  fishery.  These  are  balanced  with  frequent  interesting  recitations  from  old 
newspapers,  legislative  records,  association  minutes,  and  even  a  couple  of  native  American  legends. 

One  minor  irritant,  to  me,  was  the  author's  predominant  use  of  the  current,  awkward-sounding, 
terms,  "fisher"  and  "native  American."  They  seemed  contrived,  especially  when  "fisherman"  and 
"Indian"  slipped  in  occasionally. 

To  my  knowledge,  Salmon  Fishers  of  the  Columbia  is  the  most  comprehensive  book  on  the 
subject.  As  such,  it  would  be  valuable  reading  for  those  interested  in  Pacific  Coast  salmon  fisheries 
or  history. — David  A.  Hoopaugh 

Sampling  Design  and  Statistical  Methods  for  Environmental  Biologists 

By  Roger  H.  Green;  John  Wiley  &  Sons,  Inc.  New  York,  1979;  257  p.  $19.95. 

The  author's  stated  purpose  "is  to  provide  biologists  with  a  compact  guide  to  the  principles  and 
options  for  sampling  and  statistical  analysis  methods  in  environmental  studies,"  and  "tie  together 
a  methodology  that  already  exists  but  is  widely  scattered  throughout  many  books  and  journals."  Dr. 
Green  has  avoided  reproducing  that  literature  except  where  it  is  not  widely  available.  Biologists, 
particularly  those  with  ready  access  to  consulting  statisticians,  would  prefer  that  the  book  be  made 
more  compact. 

This  book  will  be  valuable  to  anyone  responsible  for  the  design  or  supervision  of  research  or 
monitoring  projects,  and  not  already  thoroughly  familiar  with  sampling  design.  Those  responsible 
for  advising  biologists  will  get  ideas  for  more  efficient  communication  with  biologists.  It  is  the 
author's  goal  to  bridge  the  gap  between  statisticians  and  biologists.  His  own  research  has  been  in 


70  CALIFORNIA  FISH  AND  GAME 

aquatic  biology,  as  is  much  Fish  and  Came  work.  He  has,  however,  advised  students  of  terrestrial 
systems,  and  found  the  problems  and  questions  to  be  the  same  regardless  of  the  species. 

Biologists  long  skeptical  of  statistical  methods  will  appreciate  Dr.  Green's  warning  that  ".  .  .  the 
biologically  defined  objective  should  dominate  and  utilize  the  statistics  rather  than  the  reverse." 
Because  the  book's  organization  parallels  the  chronology  of  project  design,  I'm  sure  biologists  with 
weak  math  backgrounds  will  be  tempted  to  use  this  book  to  cookbook  their  way  through  projects. 
Many  past  projects  would  have  benefited  from  such  an  approach.  Dr.  Green  does  not  recommend 
that,  but  rather  an  understanding  and  application  of  principles.  Psychologists  have  found  that 
philosophical  changes  occur  after,  not  before,  behavioral  changes.  Biologists  can  therefore  develop 
an  understanding  of  statistical  principles  while  cookbooking  their  way  through  a  project.  This  use 
of  the  book  is  not  likely  to  result  in  great  harm  because  Dr.  Green  warns  the  reader  of  critical  points 
at  which  a  statistician  must  be  consulted.  Consulting  time  will  be  reduced,  made  more  valuable,  and 
be  less  frustrating  if  biologists  will  follow  the  methods  outlined  in  this  book  to  develop  an  understand- 
ing of  what  they  want  to  do  before  consulting  a  statistician. — James  E.  Hardwick 

The  Hawaiian  Goose 

By  Janet  Kear  and  A.  J.  Berger;  Buteo  Books,  P.  O.  Box  481,  Vermillion,  South  Dakota  57069.  1980;  154 

pps.;  $30.00. 

A  comprehensive  treatise  covering  three  timely  subjects:  1.  Life  history  and  biology  of  the  Nene 
Goose,  2.  Captive  rearing  of  endangered  waterfowl,  and  3.  Success  and  problems  associated  with 
artificial  propagation  as  a  method  for  augmenting  endangered  wildlife  populations. 

I  have  never  read  a  more  detailed  account  of  a  propagation  effort  for  waterfowl.  Due  to  the 
endangered  status  of  the  Nene,  the  worldwide  interest  of  aviculturalists  and  the  progressive  individu- 
als involved  in  the  propagation  program,  of  which  Peter  Scott  of  the  Wildfowl  Trust  is  the  most  noted, 
meticulous  record  keeping  accounts  were  made  for  virtually  every  egg  and  individual  in  the  propaga- 
tion program.  Sections  include  Historical  Background,  Morphology,  Ecology,  Causes  for  Decline, 
and  Behavior. 

The  price  of  $30.00  and  the  specific  nature  of  this  book  might  put  it  beyond  reach  or  interest  for 
most  biologists  as  a  general  reference  book.  It  does,  however,  serve  as  an  excellent  high  quality 
reference  book  dealing  with  rehabilitation  and  research  needs  of  endangered  wildlife,  as  well  as 
comprehensive  work  on  the  Nene. — Dan  Connolly 

Population  Dynamics — Alternative  Models 

By  Bertram  G.  Murray  Jr.;  Published  by  Academic  Press,  Inc.,  Ill  5th  Ave.,  New  York,  N.Y.  1979;  212 

p;  $24.00. 

This  book  attempts  to  present  a  new  paradigm  of  population  dynamics,  one  which  rejects  the 
linearly  density  dependent  assumptions  of  the  logistic  model.  Murray  proposes  a  class  of  "density 
independent"  models,  wherein  per  capita  rate  of  increase  is  constant  and  independent  of  density 
up  to  a  population  size  above  which  this  rate  declines  due  to  limiting  factors.  His  thesis  is  that  these 
limiting  factors  need  not  invoke  "density  dependence,"  which  he  has  interpreted  in  the  very  narrow 
sense  of  linear  changes  in  per  capita  rate  of  increase.  Some  of  his  proposed  limiting  mechanisms 
are  reasonable;  others  appear  to  beg  the  question,  expecially  in  the  case  of  food  limitation  (p.  68). 

The  logistic  model,  which  Murray  rejects,  assumes  that  maximum  population  growth  rate  or  net 
productivity  occurs  at  one-half  of  the  maximum  equilibrium  abundance.  His  alternative  models 
characteristically  result  in  maximum  population  growth  at  greater  than  one-half  of  the  maximum 
equilibrium  abundance.  This  is  consistent  with  current,  independent  thought  regarding  population 
dynamics  and  management  of  large  mammals.  On  the  other  hand,  numerous  small  organisms,  such 
as  fish,  have  shown  maximum  net  productivity  to  occur  at  less  than  one-half  maximum  equilibrium 
abundance.  The  proposed  alternative  models  are  unable  (nor  is  the  logistic  model)  to  produce  this 
property  because  their  per  capita  growth  rates  curve  in  the  wrong  direction:  Murray's  curves  are 
necessarily  convex.  Unfortunately,  Murray  does  not  discuss  this  fundamental  limitation  of  his  mod- 
els. 

This  is  a  provocative  book  and  is  enjoyable  reading,  although  annoyingly  pedantic  in  places.  It 
forces  the  reader  to  re-examine  his  views  on  mechanisms  regulating  the  abundance  of  animals, 
which  is  a  valuable  exercise  whether  or  not  those  views  are  modified  as  a  result.  I  particularly 
enjoyed  the  criticism  of  "r-selection  and  K-selection"  wherein  Murray  shows  the  the  circularity  and 
disutility  of  this  concept.  The  book  approaches  population  dynamics  from  a  life-table  viewpoint,  and 
demonstrates  the  strength  of  the  method  even  when  used  in  a  qualitative  rather  than  quantitative 
application.  I  recommend  that  this  book  be  read  by  biologists  with  a  background  in  population 
dynamics,  against  which  it  can  be  evaluated.  I  do  not  recommend  it  as  a  text  for  a  person  seeking 
an  introduction  to  the  subject,  because  of  the  book's  polemical  nature. — Alec  D.  MacCall 


REVIEWS  71 

The  Freshwater  Fishes  of  Alaska 

By  James  E.  Morrow;  Alaska  Northwest  Publishing  Co.,  Anchorage  AK:  1980;  xv  +  248  p.,  $24.95. 

Dr.  Morrow  has  done  an  excellent  job  of  summarizing  the  current  knowledge  of  the  freshwater 
fishes  of  Alaska.  The  56  species  described  include  the  known  freshwater,  anadromous,  and  euryha- 
line  species  that  have  been  collected  in  the  freshwaters  of  Alaska.  This  a  straight  forward,  no  frills 
book  that  covers  the  basic  information  for  each  of  the  species  described. 

The  book  begins  with  a  key  to  the  families  of  fishes,  which  refers  the  reader  to  the  appropriate 
chapter  for  the  family.  Each  chapter  covers  a  separate  family  (subfamilies  in  the  case  of  the 
Salmonidae)  with  a  key  to  the  species  within  it.  Particularly  useful  is  an  illustrated  key  for  identifying 
juvenile  salmonids. 

Each  chapter  has  a  brief  description  of  the  family,  then  detailed  descriptions  for  each  of  the 
species.  An  identical  format  is  followed  for  each  species:  a  brief  paragraph  of  distinctive  characters, 
followed  by  a  detailed  taxonomic  description  of  the  species;  a  section  on  range  and  abundance, 
covering  the  total  range  and  distribution  as  well  as  that  for  Alaska;  a  detailed  description  of  the 
species  habits  and  finally,  its  importance  to  man.  Each  chapter  ends  with  one  or  more  black  and 
white  locator  maps  covering  Alaska  and  the  adjacent  area  of  Canada,  on  which  are  crosshatched 
the  areas  of  distribution  of  the  species.  The  same  base  map  is  used  in  all  cases,  which  simplifies 
comparison  between  species. 

Probably  the  greatest  feature  of  this  book  is  the  illustrations.  There  are  63  pages  of  outstanding 
photographs  and  paintings,  including  30  plates  of  watercolor  and  carbon  dust  illustrations  by  Marion 
J.  Dalen.  The  details  in  Mrs.  Dalen's  illustrations  have  to  be  seen  to  be  appreciated.  Line  drawings 
are  used  in  each  of  the  species  sections. 

One  feature  that  will  be  appreciated  by  the  more  mature  reader  is  the  printing — it  is  clear  and 
sharp,  even  the  finest  is  easily  read. 

The  only  drawback  that  I  could  find  with  the  book  is  the  lack  of  a  hard  cover.  The  present  day 
cost  of  printing,  especially  the  many  color  illustrations,  no  doubt  precluded  this. — Don  A.  LaFrance 

Inland  Fishes  of  Washington 

By  Richard  S.  Wydoski  and  Richard  R.  Whitney;  University  of  Washington  Press,  Seattle  and  London;  1979; 

xxxii   +   220  p;  illustrated;  $8.95  paper,  $17.50  cloth. 

This  ranks  with  the  best  of  the  state  or  regional  ichthyology  books.  It  was  designed  as  "...  a 
handbook  for  everyone  interested  in  fish",  and  I  believe  it  admirably  achieves  this  goal.  All  the  basic 
subjects  are  covered  in  concise,  clear  language.  Included  are  sections  on  the  drainages,  geology,  and 
topography  of  Washington  with  emphasis  on  how  they  influence  fish  distribution;  conservation  and 
management;  family  and  species  keys;  life  history  accounts,  which  include  distinguishing  characteris- 
tics, distribution,  habits  and  habitat,  age  and  growth,  reproduction,  and  food  habits;  and  references. 
The  appendix  consists  of  a  checklist  of  Washington  rivers,  and  checklists  of  Idaho  and  Oregon  fishes 
not  included  in  the  text.  An  extensive  reference  section  and  a  detailed  index  complete  the  book. 

One  of  the  outstanding  features  of  this  book  is  the  excellent  color  plates  for  75  of  the  species 
described  in  the  text.  With  four  exceptions,  the  photographs  are  of  fresh  specimens  taken  soon  after 
capture.  The  very  effective  methods  and  materials  are  described  in  the  appendix.  Numerous  nicely 
executed  line  drawings  also  enhance  this  volume. 

I  take  exception  to  some  of  the  statements.  For  example,  the  authors  maintain  that  compared  with 
other  species  of  trout,  brown  trout  survive  and  thrive  in  warmer  waters  and  are  more  tolerant  of 
turbid  waters  and  lower  oxygen  levels.  This  certainly  isn't  the  case  in  California,  where  rainbow  trout 
replace  brown  trout  in  marginal  waters. 

The  flaws,  however,  are  few  and  are  dwarfed  by  the  overall  excellence  of  this  book.  I  recommend 
it  to  anyone  sincerely  interested  in  freshwater  fishes — A/mo  J.  Cordone 


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