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The systematics of the prickly sculpin, cottus asper : an investigation of genetic and non-genetic variation.. Krejsa, Richard Joseph 1965

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The  University  of B r i t i s h  Columbia  FACULTY OF GRADUATE STUDIES  PROGRAMME OF THE FINAL ORAL  EXAMINATION  FOR THE  DEGREE OF  DOCTOR OF PHILOSOPHY  of  RICHARD JOSEPH KREJSA  B.Sc,  M i c h i g a n S t a t e C o l l e g e , 1954  M.A., U n i v e r s i t y  of C a l i f o r n i a , Los A n g e l e s , 1958  TUESDAY, JUNE 8th,1965, AT 4:00 P.M. IN ROOM 3332, BIOLOGICAL SCIENCES BUILDING  COMMITTEE IN CHARGE Chairman: 1= McT. Cowan D< H. C h i t t y H. D. F i s h e r W. S. Hoar  External  C. C. L i n d s e y G„ E. Rouse N. J . Wilimovsky P. A. Dehnel.  Examiner:  Dr. F e r r i s Neave  Pacific Biological Station Nanaimo, B „ G. '  THE SYSTEMATICS OF THE PRICKLY SCULPI.N, COTTUS ASPER, AN INVESTIGATION OF GENETIC AND ""NON-GENETIC VARIATION WITHIN A POLYTYPIC SPECIES, ABSTRACT The p r i c k l y sculpin., Cottus jaspjar, i s a p o l y t y p i c s p e c i e s c h a r a c t e r i s t i c a l l y r e p r e s e n t e d by very p r i c k l y , fresh-water spawning " i n l a n d " forms., and l e s s p r i c k l y , b r a c k i s h - w a t e r spawning " c o a s t a l " forms. I t s widespread geographic d i s t r i b u t i o n , pronounced p h e n o t y p i c v a r i a b i l i t y , and the e f f e c t s of the c o n t r a s t i n g e n v i r o n ments i n which i t occurs are the s u b j e c t of t h i s i n v e s t i g a t i o n , . A complete nom.encl.at.ural h i s t o r y and synonymy of the s p e c i e s f o r the p e r i o d 1836-1936 i s presented. M o r p h o l o g i c a l evidence i s i n t e r p r e t e d i n the l i g h t of f i e l d and l a b o r a t o r y s t u d i e s of the m i g r a t i o n and r e p r o d u c t i v e b e h a v i o u r . D i f f e r e n c e s found i n d i s t r i b u t i o n and i n t e n s i t y of body p r i c k l e s ( m o d i f i e d s c a l e s ) , g e o g r a p h i c a l d i s t r i b u tion," and m i g r a t o r y behaviour, support, the c o n t e n t i o n t h a t the p o l y t y p y of "coastal." and " i n l a n d " forms of C o t t u s asper has a g e n e t i c b a s i s , Other m o r p h o l o g i c a l evidence i s e q u i v o c a l i n support of t h i s i n t e r p r e t a t i o n . Differences i n f i n ray counts, e.g, p e c t o r a l r a y s , between " c o a s t a l " and " i n l a n d " forms are c o r r e l a t e d p o s i t i v e l y w i t h the presence or absence of s a l i n i t y i n the environment. However, it. i s not known whether such d i f f e r e n c e s are the r e s u l t of i n d i v i d u a l m o d i f i c a t i o n s induced by the local environment, or due to long term g e n e t i c f i x a t i o n . Evidence from s t u d i e s of c l o s e l y - r e l a t e d s p e c i e s supports the i n t e r p r e t a t i o n t h a t Gj__ti_s asper i s p o l y t y p i c s p e c i e s which has become, and i s i n the process of becoming, m o d i f i e d i n t o s e v e r a l c h a r a c t e r i s t i c genotypes. E v o l u t i o n w i t h i n t h i s "asper s p e c i e s group" probably o c c u r r e d i n t h r e e stages, each c o r r e l a t e d w i t h past g e o l o g i c h i s t o r y . During the f i r s t stage, perhaps i n the P l i o c e n e or L a t e Miocene, " c o a s t a l " and " i n l a n d " forms were d e r i v e d from an a n c e s t r a l marine c o t t i d . The second stage, d u r i n g the P l e i s t o c e n e , was c h a r a c t e r i z e d by fragmentat i o n of the gene pool of the " i n l a n d " form i n t o a s e r i e s of g e o g r a p h i c a l l y i s o l a t e d p o p u l a t i o n s which have s i n c e evolved i n t o valid'--species. The t h i r d stage, i n Recent Time s i n c e the r e t r e a t of the C o r d i l l e r a n g l a c i e r s , has  occurred p r i m a r i l y w i t h i n the n o r t h e r n r e p r e s e n t a t i v e s of the " c o a s t a l " form.. At. l e a s t two d e r i v a t i v e s can now be d i s t i n g u i s h e d w i t h i n the " c o a s t a l " form,  GRADUATE STUDIES  Field  of Study;  Zoology  F u n c t i o n a l Morphology of F i s h e s Marine Zoogeography Terrestrial  Related  Zoogeography  C  c  C„  J„ C,  Lindsey Briggs  M, D. V. Udvardy  Studies:  Paleontology Philosophical  V. J . O k u l i t c h Problems  B. Savery  PUBLICATIONS K r e j s a , R.J. 1960. The e a s t e r n , t r o p i c a l P a c i f i c f i s h e s of the genus B l e n n i o l u s , i n c l u d i n g a new i s l a n d endemic, Copeia, 1960, No. 4, 322-336. M c A l l i s t e r , D.E. and K r e j s a , R.J. 1961. Placement of the p r o w - f i s h e s , Z a p r o r i d a e , i n the s u p e r f a m i l y S t i c h a e o i d a e . Nat. H i s t . Pap., Nat. Mus. Canada, I I , 1-4. K r e j s a , R . J 1964, R e p r o d u c t i v e b e h a v i o r and Sexual d i dimorphism i n the manacled s c u l p i n , S y n c h i r u s g i l l i Bean. Copeia, 1964, No, 2, 448-450. C  THE SYSTEMATICS  OF THE PRICKLY  SCULPIN,  COTTUS  ASPER;  AN I N V E S T I G A T I O N OF GENE T I C AND N O N - G E N E T I C VARIATION WITHIN A P O L Y T Y P I C  SPECIES  by  RICHARD J O S E P H B . S o , Michigan M.A. , U n i v e r s i t y  A THESIS  of  State  KREJSA College,  California,  Los  195U  Angeles,  SUBMITTED IN P A R T I A L F U L F I L L M E N T OF  THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF in  the  PHILOSOPHY  Department of ZOOLOGY  We accept this thesis as conforming to  the  required  THE UNIVERSITY  standard  OF B R I T I S H  JUNE,  1965  COLUMBIA  1958  In the  r e q u i r e m e n t s f o r an  British  mission  for reference  for extensive  p u r p o s e s may  be  cation  of  written  Department  of.  by  for  study*  the  0  Head o f my  o o Columbia,  une  I^ 4  S  fulfilment  University  shall  I further  agree for  that  of •  per-  scholarly  Department  shall  of  make i t f r e e l y  or  t h a t , c o p y i n g or  f i n a n c i a l gain  ?  8  Library  the  this thesis  permission*  The U n i v e r s i t y of B r i t i s h Vancouver 8 Canada Date  the  c o p y i n g of  in partial  degree at  I t i s understood  this thesis  w i t h o u t my  that  and  granted  representatives.  this thesis  advanced  Columbia, I agree  available  his  presenting  not  be  by publi-  allowed  ii ABSTRACT The p r i c k l y sculpin, Cottus asper, i s a polytypic species c h a r a c t e r i s t i c a l l y represented by very p r i c k l y , fresh-water spawning "inland" forms, and l e s s p r i c k l y , brackish-water spawning "coastal" forms.  I t s widespread geo-  graphic d i s t r i b u t i o n , pronounced phenotypic v a r i a b i l i t y , and the e f f e c t s of the contrasting environments vestigation.  i n which i t occurs are the subject of t h i s i n -  A complete nomenclatural history and synonymy of the species *  for the period 1836-1936 i s presented.  Morphological evidence i s interpreted  in the l i g h t of f i e l d and laboratory studies of the migration and reproductive behavior. Differences found i n d i s t r i b u t i o n and i n t e n s i t y of body p r i c k l e s (modified s c a l e s ) , geographical d i s t r i b u t i o n , and migratory behavior, support the contention that the polytypy of "coastal" and "inland" forms of Cottus asper has a genetic basis.  Other morphological evidence i s equivocal in  support of t h i s interpretation.  Differences i n f i n ray counts, e.g. ,  pectoral rays, between "coastal" and "inland" forms are correlated p o s i t i v e l y with the presence or absence of s a l i n i t y i n the environment.  However, i t i s  not known whether such differences are the result of i n d i v i d u a l modifications induced by the l o c a l environment, or due to long term genetic f i x a t i o n . Evidence from studies of closely-related species supports the i n t e r pretation that Cottus asper i s a polytypic species which has become, and i s in the process of becoming, modified into^several c h a r a c t e r i s t i c  genotypes.  Evolution within t h i s '?asper species group",probably occurred i n three stages, each correlated with past geologic history. During the f i r s t stage, perhaps i n the Pliocene or Late Miocene, " c o a s t a l " and "inland" forms were derived from an ancestral marine c o t t i d . The second stage, during the Pleistocene, was characterized by fragmentation of the gene pool of the "inland" form into a series of geographically i s o l a t e d populations which have since evolved into v a l i d species.  The t h i r d  iii stage, in Recent Time since the retreat of the Cordilleran glaciers, has occurred primarily within the northern representatives of the "coastal" form  0  At least two derivatives can now be distinguished within the "coastal" form.  TABLE OF CONTENTS  ABS TRACT  o  f  l  o  e  e  TABLE OF CON TEN TS LXST OF TABLES  0  *  o  o  o  o  o  o  o  o  e  o  o  o  o  o  o  o  o  o  o  0  0  0  0  0  0  4  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  LXST OF FX GURES  o  o  o  o  o  e  o  o  o  o  o  o  o  o  o  o  o  o  o  o  ACKN OWLEDGEMENTS  o  e  o  e  o  e  o  o  o  o  o  o  o  o  o  o  o  o  o  e  INTRODUCTION  o.• 0 o 0 o o 0 a e 0 0 0 0 0 0 0 0 0 0 e o .  L i t e r a t u r e Re view  o  e  e  o  o  o  o  o  o  o  o  o  o  o  o  o  o  MORPHOLOGICAL ANALYSIS OF PRICKLY SCULPIN POPULATIONS Materials  and Methods  .  o  o  a  o  o  o  Museum C o l l e c t i o n s  o  o  o  o  o  o  . o  o  o»»»*t»*»»»**».» , . . o o . o o o « « « <> «  M a t e r i a l Examined  <,  « . . » . . » o . « >«  Counts and Measurements  C r i t i q u e o f t h e Methods p f D e t e r m i n i n g P r i c k l i n g D i s t r i b u t i o n and I n t e n s i t y . . « o o o o <> « « o S t a t i s t i c a l Methods O  O  O  O  O  V  O  O  Ran ge  Results;  0  O  0  0  o  .  o  o  .  O  O  O  O  O  O  O  o  o  o  O  O  o  O  o  o  O  o  O  o  O  . o . o o o « o o o o o  Nomenclatural H i s t o r y Distribution  o  O  o  o  o  o  «  o  o  o  o  o  o  o  o  »  o  o  o  o  o  0  0  0  0  0  0  0  0  0  0  0  0  0  0  9  0  0  0  A n a l y s i s o f Geographic V a r i a t i o n « , « «-  Prickling Distribution  o  , 0 . 0 0 0 0 0 0 0 0 0 0  Prickling Intensity  o  o  o  o  o  o  o  o  o  o  o  o  o  o  P e c t o r a l F i n Asymmetry . . . . 0 « 0 « « 0 0 . 0 P e c t o r a l F i n Ray Count Spiny D o r s a l Soft Dorsal  Fin Fin  Combined D o r s a l  8  > . • . 00  0  0 0 0 0 0 0 0 0 . 0 0 0 0 0 0 0 o  o  o  o  o  o  -  .  o  o  o  e  F i n Elements . . « ' . . .  o  o  o  o  . . .  o  «  V  Alldl PlTl  o  PsXVIC P i n  o  c  o  e  o  a  o  o  o  o  o  o  «  o  «  o  o  o  f  l  o  o  o  e  A  o  A  O  O  *  o  o  o  O  o  e  o  o  o  o  <  o  j  o  o  o  o  e  (  o  »  3S  e  9  o  o  ^0  L a t i t u d i n a l Variation i n Ranges of Mean Fin Ray Counts , . . .  40  Lateral Line  43  O  0  o  e  o  6  e  .  .  o  o  o  o  o  o  o  o  .  .  Discussion of Meristic Variation Within C. asper  .  0  .  0  0  <,.„...  0  .  0  Summary of Morphological Analysis . . . . . . . . . « < > . . . . . STUDIES ON THE LIFE HISTORY OF THE PRICKLY SCULPIN . . . . . . . . . . .  43 49 51  Present Knowledge of Spawning Migration i n Cottus asper . . . . .  51  Spawning Period and Temperature Relationships . . . . . . . . . .  52  Materials and Methods .  55  Sampling  c  ,  r  *  =  c  (  ?  •  .  .  .  .  o  •  o  .  .  .  .  .  .  S a l i n i t y Tolerance i n Prickly Sculpin Eggs DlSCUSSXOn  o  .  e  o  o  o  e  o  .  o  o  o  '  o  «  •  .  .  •  •  t t t » »  L o c a l i t i e s and Study Streams  Results of F i e l d Studies  «  .  e  t  i  .  .  3  ?  .  .  .  .  .  58  .  . . . . . . .  e  G  55  s  .  f  l  .  e  0  .  o  64  0  Summary of L i f e History Studies . . . . . . . . . « . « . . . . . DISCUSSION  o  o  ~  .  .  o  o  o  o  e  o  o  o  6  6  6  .  6  0  0  0  0  0  0  0  0  0  0  0  62  65  0  65  0  Morphological Variation Within„Cottus asper . . . . . .  66  The Phylogeny of Cottus asper and Closely-Related Species . . . .  70  Stage  l o  o  o  .  e  o  o  e  e  e  o  o  o  o  o  o  o  e  o  o  o  State  l i e  0  0  6  0  0  0  0  0  0  0  0  0  0  0  0  0  0  0  .  0  0  o  o  .  0  0  0  0  0  0  0  0  0  .  9  6  0  0  4  .  0  o  o  .  .  o  .  .  o  6  o  «  .  .  .  .  0  .  .  e  .  «  «  0  6  0  .  0  0  0  0  0  0  6  0  0  6  6  6  6  0  0  Stage' IIIo LITERATURE CITED  6  o  o  APPENDIX  0  TahleS 1.  0  0  0  0  0  0  0  0  0  0  0  0  0  0  6  0  6  9  6  0  0  0  0  .  0  0  0  0  0  0  .  0  0  0  0  o  o  6  o  9  0  o  .  0  0  72  .  .  a  >  e  0  .  79 83 90  9  0  74  0  90  0  L i s t of C o l l e c t i o n s , L o c a l i t i e s , and Other Pertinent Information  2.  0  o  O  6  0  O  6  .  «  «  6  .  .  6  O  «  «  .  6  O  0  90  Weighted Mean Percentage P r i c k l i n g and Resorption for Individual Populations of Cottus asper Used i n Compiling Table I  o  .  6  0  6  .  e  o  o  o  o  .  o  .  .  6  .  6  0  9  98  3,  Non-Weighted Mean Percentage  Virtual Prickling  in Each o f 1 4 Body Sections f o r A l l Populations (Specimens 4„  ^60  mm  S  0  <  L  i  )  o  o  o  o  o  o  0  o  o  o  o  o  o  o  o  8  o  a  o  o  o  »  »  100  . 102  , « . . .  o  o  o  o  a  o  a  o  o  o  o  o  a  o  o  104  o  o  107  Summary of Meristic Counts (Means and Ranges) f o r A l l Populations of Cottus asper  7o  o  Pectoral Fin Asyfflme-try i n Cottus asper by Populations  60  o  Mean Relative P r i c k l i n g Intensity Index Values for A l l Populations o f Cottus asper  5  o  o  o  o  o  o  o  o  a  o  o  o  a  e  Monthly Mean Temperature i n °C» f o r Selected Coastal and Inland L o c a l i t i e s ,  e  o  o  o  o  s  o  o  a  o  a  t  o  o  a  s  s  a  109  vii LIST OF TABLES Table I  Title Mean Percentage P r i c k l i n g and Resorption Among Aggregate Populations of Cottus asper i . . . . . . . . . . .  II  27  Mean Relative P r i c k l i n g Intensity f o r Populations of Cottus asper Aggregated by L o c a l i t y and P r i c k l i n g Affinity.  III  Skin Section " j "  . . . . . .  30  Incidence of Occurrence, Direction, and Amount o f Pectoral Fin Asymmetry Within and Among Aggregated Populations of Cottus asper  IV  . . . . .  . . . . . 32  Lateral Line Pored-Scale Counts by Size Groups f o r Selected Representative Populations of Cottus asper  V  44  Summary o f Variable Characteristics f o r Selected Phenotypes of Cottus asper  VI  . . . .  . . . . . .  . . . . . . . . .  0  . 67  Summary of M e r i s t i c Data f o r Cottus asper and Other Species i n ithe "asper species group"  °°  .  0  0  9  .  6  0  0  .  77  viii LIST OF FIGURES Figure 1.  Title "Skin Map" Used f o r Recording D i s t r i b u t i o n and Intensity o f P r i c k l i n g i n Cottus asper  2.  . . .  0  » •  Areas o f P r i c k l i n g and Areas o f Resorptions A.  in Coastal Cottus asper . . . . . . . . . .  Bo  i n Inland Cottus asper  9 .. •  o o o o o o  3.  D i s t r i b u t i o n a l Range of Cottus asper . . . . . . . . . . . .  4.  V i r t u a l , Resorbed, and Apparent P r i c k l i n g i n CottUS aSper o o ' o . f l . o o . o . . . Mean Pectoral Fin Ray Count by Latitude  6  Mean Spiny Dorsal Fin Ray Count by Latitude  9 25  . . . . . . . . . . .  5. 0  7  26  . . . . , . . . . * . .  34 36  7.  Mean Soft Dorsal Fin Ray Count by Latitude . , . . . . . .  8.  Hubbs-Hubbs Plot of Soft Dorsal Fin Ray Counts f o r  .  37  Inland Population Samples Pooled into Aggregates of Two Degrees Latitude  . . . . . o . . . . . . . . . . . . .  38  Mean Combined Dorsal Fin Ray Count by Latitude . . . . . . .  39  10.  Mean Anal Fin Ray Count by Latitude  41  11.  Approximate Ranges o f Mean Fin Ray Counts Plotted  9.  . . . . . . . . . . . .  Against Latitude f o r Inland, Coastal, Coastal Lake, and Coastal Derivative Populations 12.  . . . . . . . .  42  Monthly Mean Temperatures i n °C. Arranged by Latitude for Coastal and Inland L o c a l i t i e s Encompassing the D i s t r i b u t i o n a l Range o f Cottus asper . . . . . . . . . . . .  13.  54§  Collection L o c a l i t i e s f o r Spawning Populations o f Cottus asper Used i n Life History Studies  . . . . . . . . .  14.  Collection L o c a l i t i e s Within L i t t l e Campbell R.  15.  Collection S i t e s , Tidal Influence, and Stream Gradients i n the L i t t l e Campbell R.  . . . . . .  . . . . . . . . . . . .  56 57  59  Figure 16„  Title Monthly Distribution of Young-of-the-Year  Cottus asper  Within the L i t t l e Campbell R„ Taken from Pooled Bi = Weekly Samples ,17.  Distribution of Yearling, Sub-Adult, and Adult Cottus asper Within the L i t t l e Campbell R. . . . . . . . .  18.  61  o o o o . o o o o o e o o o o o , , . . . . .  .  63  Known Ranges of Various Phenotypes of Cottus asper Plotted Against the D i s t r i b u t i o n a l Ranges of CloselyRelated Species i n the "asper species group" . . . . . . . .  19.  Phylogeny  of Cottus asper.  the Pliocene (? ) 20.  o o o  O  Phylogeny of Cottus asper.  Stage I. O  O  O  O  O  O  Evolution in 0  Stage I I ,  0  0  0  0  0  e o a o o  Phylogeny of Cottus asper.  a  73  Evolution of "asper  species group" During the Pleistocene 21.  71  78  Stage I I I . Evolution o f  Coastal Form Following Retreat of Cordilleran Glaciers During Recent Time . . , , , e  s  '. , . , . . , , , , « . . .  80  X  ACKNOWLEDGEMENTS I wish to express my indebtedness t o the many persons and organizations who have given me encouragement and assistance during the period o f t h i s i n vestigation.  The B r i t i s h Columbia Fish and Game Branch provided f i n a n c i a l  support during the summer of 1960 and also the much appreciated assistance of Dr. G. F. Hartman and Mr. C„ A„ G i l l i n f i e l d studies.  The National Research  Council of Canada provided studentships i n 1961 and 1962. The Vancouver Public Aquarium provided research space for l i f e history studies and I wish to thank the Curator and s t a f f f o r a l l t h e i r help and kindnesses. Almost every graduate student enrolled i n the Institute of F i s h e r i e s volunteered assistance i n f i e l d c o l l e c t i o n s a t one time or another during the period 1960-1963.  I o f f e r a c o l l e c t i v e thanks to a l l  0  Dr. C. Co Lindsey's assistance in supervision o f the e n t i r e i n v e s t i gation cannot be measured i n words.  His door was always open to me and I  took advantage o f that fact on innumerable occasions. Drs. N. J . Wilimovsky provided encouragement and many h e l p f u l suggestions and c r i t i c i s m s throughout the i n v e s t i g a t i o n , and above a l l , friendship. Dr. R. H, Rosenblatt provided much i n s p i r a t i o n during the period o f h i s (  . . .  ...  v i s i t i n Van'couver  0  Dr. Carl E. Bond shared his wealth o f knowledge about the members o f the genus Cottus. Committee members Drs  0  W. S, Hoar, E, C. Black, P. A, Dehnel, and  D. Wo Chitty read the manuscript and offered helpful suggestions and c r i t i cisms throughout the study. Mrso Mary Williams typed the f i n a l copy and her perserverance i s greatly appreciated.  xi  Finally, I wish to offer my sincere gratitude for the help and encouragement of my wife Julie, who, while patiently tolerating a l l the indignities encompassed within the t i t l e "wife of graduate student", found time to act as literary c r i t i c , typist, and friendo  I INTRODUCTION The role o f the environment  versus the role of heredity i n determining  the phenotypic variations found among and within natural populations has long been a problem o f central interest to students of evolution. Because fishes apparently undergo modification by t h e i r  environment,  they have proved to be useful i n the study of phenotypic v a r i a t i o n .  For  example, several authors have shown that non-genetic morphological or func*t i o n a l variation i n fishes can be caused by maternal care and diet p r i o r t o egg deposition, and to the e f f e c t s of environment development, and a f t e r hatching.  during f e r t i l i z a t i o n and  In spite of such documentation many  systematic studies t a c i t l y assume that differences discovered between i n * dividuals, or within and/or among series of specimens, are genetic. In  some species, breeding experiments have shown that, under controlled  conditions, the i n t e g r i t y o f the parental phenotype i s maintained i n the F^ generation. mined.  Such r e s u l t s have been interpreted as being g e n e t i c a l l y deter-  Except f o r a few notable examples, there have been no studies on  fishes which have attempted to correlate the morphological and physiological evidence and t o r e l a t e both to the l i f e history o f the species concerned. Such an approach i s attempted in. t h i s investigation. The p r i c k l y sculpin ranges geographically over about 3000 miles of P a c i f i c North Temperate coastline and inland as f a r as 300 miles.  Structural  modifications correlated with l a t i t u d i n a l changes i n temperature 0.& s a l i n i t y should, i f present, be revealed by morphological analysis.  Like the s t i c k l e -  back Gasterosteus aculeatus, and the darter Etheostoma (Boleosoma) nigrum, the p r i c k l y sculpin e x i s t s i n two modes of morphological v a r i a b i l i t y ?  one,  a fresh-water spawner, has extensive squamation on certain regions of the body; the other, a brackish-water spawner, has l i t t l e or none. sculpin eggs are spawned n a t u r a l l y i n environments at  least one major f a c t o r , i . e , ,  Since p r i c k l y  which are known to vary i n  s a l i n i t y , and since, under contrasting  environmental circumstances, i t shows so many similarities with the stickleback, the prickly sculpin i s an excellent subject for variational studies. Prior to the present study, regional systematic treatments of this widespread species resulted in a proliferation of generic and specific taxa, a l l referable to Cottus asper.  A complete nomenclatural history and synonymy  of the species for the period 1836-1936 i s presented.  Morphological evidence  based.on examination of 1604 specimens from 190 collections throughout the range of the species i s correlated with that from f i e l d and laboratory studies of the migratory and reproductive behavior. The thesis i s presented in two sections corresponding to the morphological and l i f e history approaches followed in this investigation.  A -  general discussion considers the morphological and ecological variation within the species and attempts to reconstruct the phylogeny of Cottus asper and closely-related species. Literature Review Hubbs (1943) has listed many reasons why fishes are particularly wellsuited for determination of genetic vs. environmental responses.  He has also  summarized (1926) much of the early work especially in regard to the structural consequences of varying developmental rates.  Numerous workers have  found that different morphological or meristic series in fishes are "plastic," i.e., sensitive to environmental influence before egg deposition, during fertilization, during development, and after hatching. Taning (1944) ascribed at least one instance of mass vertebral deformities and low egg survival to an accident befalling the female trout before the eggs were removed from her.  :  Hubbs and Stavenhagen (1958) have  shown that low survival of different egg lots was caused by carotenoid deficiency in the maternal diet.' Kinne (1962) reviewed some of the recent literature on structural and functional properties of fish egg membranes and suggested that the spawning  3 medium affects the physical properties of the immediate environment of the embryo and that these induced properties "tend to persist in later different salinities."  On the basis of his own studies, of Cyprinodon macularius, he  interprets such conditioning by the spawning medium, leading to irreversible functional or structural adjustment in the embryo, as a special case of imprinting, or "irreversible non-genetic adaptation." Battle (1929); Mottley (1934); Taning (1944); Gabriel (1944); Heuts (1949); Lindsey (1954);, and Seymoilr (1959), have demonstrated the effects of environmental influences, especially temperature, during different developmental stages prior to hatching. Strawn (1961) found that the diet of newly-hatched larvae can affect the morpbGi&gy'*  Lindsey (1954) found that certain meristic characters are  thermolabile up to 20 days after hatching. Lagler and Bailey (1947) studied the inheritance of differential morphological characters in two subspecies of Etheostoma (Boleosoma) nigrum, a darter.  The main differences between the subspecies l i e in the extent of  the squamation on the nape, cheek, and breast, one being typically naked or with few scales, and the other being well, i f not f u l l y , scaled. Bailey found that the.F  Lagler and  generation of each subspecies, when reared and  hatched under uniform environmental conditions in the laboratory, maintained the integrity of the parental phenotype.  They concluded that these char-  acters were genetically fixed. «.•••....•• After a vast and careful examination of some 9000 individuals, Bertin (1925) concluded that in populations of the stickleback,. Gasterosteus aculeatus, throughout Europe, most of the variable characters, especially the lateral body plates, are direct individual adaptations to environmental influences, especially temperature.  He described four forms based primarily  on body plate numbers. Heuts (1947a), however, showed that, at least in Belgian sticklebacks,  4 only two main modes of variability exist, each correlated with definite ecological characteristics, each with differences in breeding habits and distribution.  He also showed (1947b) that certain physiological characters  are correlated with morphological ones and he attempted to demonstrate that natural selection favors different complexes of genes in the different ecological niches in which the species occurs.  He considers the two types  of sticklebacks to be valid genetic races. Robins and Miller (1957) have called attention to the fact that because variation i s so marked and haphazard in the sculpins of the genus Cottus, interpretation of the species limits within the group i s frequently d i f f i c u l t . They consider Cottus to be one of the=most perplexing groups of North American freshwater fishes.  In considering a character such as prickles on the body,  they state that so much variation occurs between populations that i t s geographic consistency in one or two forms i s d i f f i c u l t to interpret. They also consider that a character such as "armature, like body prickles, may greatly assist in the identification of a species at any one locality, but i s often unreliable for relating or distinguishing allopatric forms."  MORPHOLOGICAL ANALYSIS OF PRICKLY SCULPIN POPULATIONS Materials and Methods Museum Collections. - Specimens examined include those in the following institutional collections (abbreviations listed are used in the text)! AB  Bureau of Commercial Fisheries, Biological Laboratory, Auke Bay, Alaska  BC  University of British Columbia, Vancouver, Canada  CAS  California Academy of Sciences, San Francisco  NMC  National Museum of Canada, Ottawa, Ontario  OSC  Oregon State University, Corvallis  SNHM Natural History Museum, Stanford University, Palo Alto, California (SU)  5 UMMZ University of Michigan Museum of Zoology, Ann Arbor USNM  United States National Museum, Washington, D, C,  UW  University of Washington, Seattle Additional non-catalogued British Columbia material used in this study  was made available through the personal f i e l d collecting of the following individuals.  Abbreviations used are those which preface field collection  numbers s K-  R. J, Krejsas Little Campbell R. , and miscellaneous others  A-  G. F. Hartman, D. W. Wilkie, and C. A. Gills South Alouette R.  V-  " "  "  " "  SW-  " "  »  R. J. Krejsa,  S-  J. S, Nelson, and P. J. McCarts Peace R. drainage.  11  «•• " " " " "  " "  . Vedder R, s Sweltzer Cr,  Material Examined, - A total of 1604 specimens from 190 collections was utilized in this study.  Data for 158 specimens, from 16 collections, was  culled from the published accounts of Snyder (1905, 1908b, and 1913), and Rutter (1908).  Counts and measurements for 63 specimens, from 6 collections,  were taken by Dr. D. E. McAllister of the National Museum of Canada, The remaining 1383 specimens were examined by the author. A l i s t of collections  B  together with pertinent data, i s presented in  Appendix Table 1, Counts and Measurements. - Measurements were taken to within 0.1 mm using break-arm dividers with one flat point and one needle point, in accordance with the methods of Hubbs and Lagler (1958). Counts for median fins and pelvic fins follow the methods outlined by Robins and Miller (1957).  Other counts were taken as follows.  Pectoral Fin Rayss Counts from both the right and l e f t side were summed by population and then divided by the total number of fins.  Pectoral  fin asymmetry in Cottus asper was investigated to ascertain the incidence of occurrence, the direction, and the amount of asymmetry in this species, and  6 to determine whether individual populations, population aggregates, or geographical regions could be characterized by these criteria. Distribution of Prickles on the Bodyi  The areal distribution of body  prickles i s defined as the area of skin surface covered, or formerly covered, by prickles.  It was recorded as follows.  A superficial median incision was  made through the dorsum and venter of a single adult specimen of Cottus asper 80 mm in standard length (S.L.).  A transverse incision was made ventrolater-  a l ly from the median supratemporal pore to a point medial and posterior to the pelvic f i n base.  The skin was then peeled off the left side of the body  as far posterior as the hypural fold where i t was excised. It was then placed f l a t on a sheet of paper and a tracing made around the perimeter.  The resulting blank outline tracing was then subdivided into  14 lettered sections based on morphological landmarks.  The subdivided put-  line of the original skin hereinafter referred to as the "skin map,"  was  stenciled onto sample work sheets on which a l l counts and measurements were recorded (Fig. IB), To estimate the areal distribution of prickles, a specimen i s examined under the binocular dissecting microscope and the perimeter of the prickly area on the left side i s outlined by scratching the mucous layer with a sharp needle point.  This "apparent" outline is copied onto the work sheet  skin map,with colored ink.  The mucus i s then scraped from the entire l e f t  side with the aid of a syringe-needle jet of compressed a i r , exposing those partially-resorbed prickles which do not project through the mucous layer. As w i l l be discussed later, some prickles in Cottus asper are completely resorbed during maturation.  Therefore, further treatment is necessary to  expose the extent of the prickling in i t s entirety, i.e., the "virtual" prickling.  Vigorous rubbing or massaging of the skin with the air needle  will expose the areas formerly covered by prickles. This technique presumably causes the breakdown of the sub-epidermal chromatophores.  The  7  A  LATERAL  •DORSAL  ASPECT  ASPECT,  LEFT  SIDE  F i g u r e lo — "Skin Map" Used f o r R e c o r d i n g D i s t r i b u t i o n and I n t e n s i t y P r i c k l i n g i n C o t t u s aspero  of  8  melanin granules thus released become aggregated i n the empty dermal p r i c k l e pockets and the areas of resorption become v i s i b l e .  Thus resurrected, the  o r i g i n a l or " v i r t u a l " a r e a l d i s t r i b u t i o n of p r i c k l i n g can be recorded-in ink of another color over the "apparent" p r i c k l i n g d i s t r i b u t i o n already recorded on the skin map  ( F i g . 2).  Since each of the 11 lettered sections of the skin map represents a f r a c t i o n of the t o t a l body surface on the l e f t side of the animal, a polar planimeter was used to determine the t o t a l area occupied by each section. Sectional areas were then converted into a set of f r a c t i o n a l constants, each : representing a percentage of the t o t a l area of the skin map.  These constants  were used i n calculations of the absolute p r i c k l i n g i n each section. For s t a t i s t i c a l treatment, the t o t a l areal d i s t r i b u t i o n of p r i c k l e s was tabulated as followss  A copy of the skin map was permanently printed onto a  piece of transparent p l a s t i c .  On t h i s overlay, each of the lettered sections  was further partitioned into subdivisions of 5, 10, and 25%.  The p l a s t i c  overlay was matched with the completed skin map on the work sheet and the percentage area covered by p r i c k l e s i n each numbered section was v i s u a l l y estimated to within 5%.  The estimates f o r each section were then transcribed  and summed f o r a l l specimens i n the sample and a mean percentage "apparent" and mean percentage " v i r t u a l " p r i c k l i n g was determined.  M u l t i p l i c a t i o n of  each of these sectional mean percentages by the appropriate f r a c t i o n a l constant (previously determined) resulted i n a product which i s an estimate of the mean percentage absolute p r i c k l i n g ( " v i r t u a l " and/or "apparent") f o r each section. The sum of a l l 14 sectional products i s an estimate of the mean per^ centage areal p r i c k l i n g ("virtual'' or "apparent") f o r the population sample. Subtraction of the population mean percentage "apparent" areal p r i c k l i n g from the population mean percentage " v i r t u a l " areal p r i c k l i n g r e s u l t s i n a remainder which represents the population mean percentage absolute /respcption  AREA OF RESORPTION  VIRTUAL  PRICKLING  Figure 2 «—Areas of Prickling and Areas of Resorptions  A. in Coastal Cottus asper; B, in Inland Cottus asper.  10 for any given sample. The mean percentage r e l a t i v e resorption i n any given population sample i s obtained by the formulas Mean % r e l a t i v e resorption = 100% - inn/Mean  V"Apparent"\  \Mean % " V i r t u a l " / An examination of the condition of the gonads i n more than 1000  speci-  mens indicated that Cottus asper usually becomes reproductively mature at a size of 60 mm  Selio  On this b a s i s  treatment of a l l p r i c k l i n g data was  9  divided into groups of specimens smaller than 60  9  or equal to and larger than,  mm. P r i c k l i n g Intensity;  The size of p r i c k l e s in Cottus asper varies  d i r e c t l y with the' size of the animal.  However, the i n t e n s i t y , i . e , , the  number per unit area, does not appear to be a d i r e c t function of s i z e ,  It  varies among individuals of a l l sizes and even on d i f f e r e n t sections of the same i n d i v i d u a l ,  Because of the i m p r a c t i c a l i t y of recording absolute p r i c k l e  numbers f o r large numbers of specimens, a r e l a t i v e index of range 0 through 5 was established.  In t h i s index zero indicated no p r i c k l i n g , and f i v e i n -  dicated the most intense p r i c k l i n g . When a single p r i c k l e was found i n an otherwise naked area, i t was given an a r b i t r a r y value of 0.5,  For s t a t i s t i c a l purppses, a single isolate  was considered to have an a r e a l d i s t r i b u t i o n which was proportional to the lettered body section of the skin map  i n which i t was found, i . e . , 4% i n  sections 'a' and 'b'; 10% i n sections 'd', 'e , 'h , 'k', and 'n'; and 1  i n sections '1' and 'm'.  1  5.5%  I f more than 4 p r i c k l e s were found i n an i s o l a t e d  patch, an index value of "1" was assigned and the areal d i s t r i b u t i o n  was  estimated to the nearest unit percentage. To determine the nature of d i f f e r e n t i a l mean i n t e n s i t y index values, sample counts of p r i c k l e s were taken from sq, cm, sections of skin adjudged to have index values of one and f i v e .  In rounded numbers, an index value of  11 "1" i s equivalent to 400 p r i c k l e s per sq. cm,,  whereas a value of "5" i s  equivalent to an estimated 2500 p r i c k l e s per sq. cm.  Mean values were i n t e r -  polated from these counts. Lateral Lines  Previous workers (Robins and M i l l e r , 1957) have con-  sidered the l a t e r a l l i n e as incomplete i f the number of pored-scales did not reach the hypural f o l d .  The number of completely pored l a t e r a l l i n e scales  varies with size and among populations.  Close examination of sculpins r e -  veals that although a specimen may have an "incomplete" l a t e r a l l i n e ( i . e . , pored-scale count), unenclosed neuromast h i l l o c k s are present and r e a d i l y visible.  Thus the most r e l i a b l e measure of s e r i a l r e p e t i t i o n along the  l a t e r a l l i n e , f o r a l l size categories other than young-of-the-year,  is a  count of the complete pored-scales plus the p a r t i a l l y , or completely unenclosed neuromast h i l l o c k s which appear sequentially out onto the caudal f i n .  along the trunk and  In p r a c t i c e , t h i s count was made only to the  hypural f o l d and the mean number of pored-scales plus h i l l o c k s was considered to represent a "complete" l a t e r a l l i n e count.  "Complete" counts were made;  for less than one half of the specimens examined in t h i s study and, in most small specimens, the count was l i s t e d only as "incomplete." c o l l e c t i o n s i s few i n which a l l size ranges from juvenile presented.  The number i n which "complete"  The number of  to adult are re>-  l a t e r a l l i n e counts have been  taken i s even fewer. Critique of the Methods of Determining P r i c k l i n g Distribution Intensity.  and  - The method of determining d i s t r i b u t i o n proved unreliable  extremely o l d and/or badly preserved specimens.  on  In some the skin was so  desiccated that massage was impossible without damage to the specimen.  In  others, a l l the pigment had been bleached a f t e r years of preservation and no amount of massaging would make the empty p r i c k l e pockets appear.  In such  specimens only the "apparent" p r i c k l i n g d i s t r i b u t i o n was determined recorded.  Estimates of p r i c k l i n g d i s t r i b u t i o n on badly preserved or  and  12 desiccated specimens were not used in compiling population means. Tiny prickles in peripheral areas of immature specimens, and partially resorbed prickles on heavily pigmented mature specimens were d i f f i c u l t to see, even with the aid of the microscope.  Therefore the estimates of  "apparent" prickling both in mature and immature specimens are less than the "virtual" prickling in immature specimens.  The accuracy of the visual  estimate in peripheral areas was checked against alizarin-stained specimens, and was found to be within -10% on specimens > 60t>,mm, and within ±5% in specimens < 60  mm.  While 60 mm appears to be a valid estimate of the average size at which most Cottus asper attain reproductive maturity, not. a l l spawn at that size. Some populations, e.g., from Queen Charlotte Is., are known to have females which spawn at a size of only 42 mm_  S.L., while in other populations, a few  females do not spawn u n t i l a size of 70 mm is attained. The discrepancy in Table I between the mean percentage "virtual" prickling and the slightly lower (from 2-7%)  "apparent" prickling in fishes < 60 mm i s attributed to.;  this cause. Another discrepancy in the data reported in Table I is that mean differences of from 4 to 10% occur between the "virtual" prickling percentages in specimens < 60 mm, and those >60  mm from the same population. These  differences are attributed to the inability to "resurrect" a l l the prickle pockets completely. Statistical Methods. - Most of the raw data has been converted to means or percentages.  Obvious differences were not tested. When a comparison of  means was thought necessary, the two most divergent values were examined first.  If no significant differences were revealed, the less divergent o  means were not tested. The single-classification  test, or the Student  t-test, was used at the 5% c r i t i c a l level to determine acceptance or rejection of an hypothesis of equal means. Graphical presentation of certain  13 meristic characters i s according to the format of Hubbs and Hubbs (1953), Synonymy It has been almost 130 years since Cottus asper was f i r s t described by Sir John Richardson,  Early revisionary work by Girard in 1851 and 1852  incomplete because of lack of specimens.  was  Recent regional works (Robins and  Miller, 1957; McAllister, 1957; McAllister and Lindsey, 1959; and Bond, 1963) have included the species as part of a geographical area or river drainage system, but there has never been a comprehensive treatment of the species throughout i t s entire range.  In the period 1836-1936, 38 reports of Cottus  asper were cited in the literature.  In the same period, 32 additional cita-?  tions occurred which were either misidentifications or synonyms properlyreferable to Cottus asper.  In none of the systematic treatments published  since the original description in 1836 has there been a synonymy containing more than 7 citations.  McAllister (1957) listed 15 citations in his un- >  published M.A, thesis. The present synonymy, consisting of 70 citations published during the period 1836 to 19536, i s thought to be complete.  Since 1936, the species has  been cited in numerous fishery journals and publications.  Therefore, only  the major systematic, or otherwise noteworthy, citations occurring in the past 29 years have been recorded. Cottus asper Richardson, 1836 Cottus asper. -  Richardson, 1836s295, p i , 95, Fig. 1 (original description  and figure; Columbia R,; collected by Dr, Gairdner (probably near Fort Vancouver, Washington Territory)),  Storer, 18468.260, and  1846bs8 (northwestern coast of N. America).  Girard, 1850s409, and  1851asl89 (discusses propriety of present nomenclature).  Eigenmann,  1895&118 (abundant in Fraser system from tidewater to 1900 feet; Mission, Sicamous, Kamloops, and Griffin L., British Columbia; and Umatilla, Oregon).  Seale, 1896s854 (Lake Washington),  Gilbert and  14 Evermann, 1895:201 (description; comparison to Sacramento R. form; Walla Walla R. at Wallula, and Lake Washington, Washington). Gilbert, 1896:418 (description; stream entering Departure Bay, Vancouver Island).  Jordan and Evermann, 1896:439 (synonymy; streams  of the Cascade Range, from Vancouver Island to Oregon). 1898:1 (Columbia R.; notes absence in Klamath R.).  Gilbert,  Jordan and  Evermann, 1898:1944 (description; synonymy; Walla Walla; Departure Bay; about Port Townsend; streams of the Cascade Range, from Vancouver Island to Oregon). Washington). ington).  Evermann and Meek, 1898:83 (Lake  Meek, 1899:231 (Lake Southerland, Olympic Pen., Wash-  Snyder, 1905:337 (description; a f f i n i t i e s ; habitat pre-  ference; San Franciscito, Madera, San Antonio, Guadalupe, Coyote, and Alameda Creeks, a l l flowing into San Francisco Bay). Evermann and Goldsborough, 1907a:306 (characters; prickling descriptions; Deep Bay, Naha R., and Steelhead Cr,, Loring, Alaska; Hunter Bay, Yes Bay, and McDonald L., Alaska).  Evermann and Goldsborough,  1907b:110 (Fraser R, at Mission, Shuswap, L. at Sicamous, Thompson R. at Kamloops, and Griffin L . R u t t e r , 1908:145 (Cottopsis parvus :  f i r s t placed in synonymy with asper; synonymy, in part, except Uranidea semiscaber (sic) centropleura Eigenmann and Eigeniiann; summary of 9 localities in Sacramento R, system). (Russian R., California)*  Snyder, 1908a:269  Snyder, 1908b:184 (characters; prickling  description; summary of A l localities: from Lake Washington, Columbia and Sacramento R., and river basins in between). (head of Chilkoot L., Alaska),  Nichols, 1909:172  Evermann and Latimer, 1910:138 (4  localities in Marin Co., and 2 localities  in San Francisco Bay,  California; 12 localities in Olympic Pen., Washington). 1913:72 (characters; Pajaro R., California). (Papermill and Walker Creeks, California),  Snyder,  Snyder, 1916:381 Kermode, 1917:20  15 (Hanceville, B r i t i s h Columbia (Chilcotin R« drainage)),,  Jordan,  1919;249 (designates Cottus asper Richardson, as the orthotype of Cottopsis Girard),  Bean and Weed, 1920g76 (mouth of Fraser R,).  Hubbs, 1921s7 (re-identification of San Luis Cr, California, speci= 9  mens misidentified by Jordan as C, gulosus (1895sl41); range ex= tension to Ventura R,, California),, flowing into Puget Sound).  Crawford, 1927sl77 (streams  Schultz, 1929s48 (listing only),  Schultz, 1930;14 (most streams and lakes of western Washington), Jordan, Evermann, and Clark, 1930g383 (synonymy; streams of the Cascade Range, southeastern Alaska to Oregon; south to Sacramento R,), Kermode, 1931.19 (Cowichan L,„ Vancouver Island),  Evermann and  Clark, 1931s56 (summary of 32 recorded localities in California), Schultz and DeLacy  s  1936as 128 (synonymy; coastal streams from Alaska  to Ventura Co,, California; freshwater and brackish water; review of most records from Puget Sound to Oregon, and addition of 26 more localities),  Schultz and DeLacy 1936bs213 (additional synonymy; 8  3 new localities),  Schultz, 1936sl79 (keys to species of Cottus;  coastal streams from Alaska to Ventura Co and brackish),  08  California; freshwater  Dymond_ 1936.71 (description; throughout souths  western British Columbia including southern Vancouver Island; 16 8  localities listed),  Sumner, 1942'gl=>25 (common i n tidewater areas  along the Oregon coast),  Hubbs and Wallis  s  of "Cottus sp." recorded by D i l l , 1946s54), 1949sl4 (comparison with Cottus hubbsi), 1950s387 (listing only), California).  1948s141 (identification Bailey and Dimick_ Shapovalov and D i l l ,  Wilimovsky, 1954s285 (southeast Alaska to  Robeck et a l , 1954sB=65 (Columbia R, g  8  above Trinidad,  Washington; cited as "prickly sculpins Cottus sp,", these may include Cottus rhotheus in part,)  Lindsey, 1956s777 (Pacific Slope of  No America from Alaska to California; British Columbia mainland from  16 Columbia, Fraser, and Skeena systems, Stikine R, Peace R.  headwaters, and  (Summit L., Heart' L,, Angusmac Cr,, and McLeod L . ) ) ,  Robins and M i l l e r , 1957:229 (Cottopsis parvus again removed from synonymy of C. gulosus ),  Lindsey, 1957:657 ( B r i t i s h Columbia:  Columbia R.; Fraser R.; Skeena R,; Coast drainages south of Skeena; Nass R.; Stikine R.; Peace R,).  Wilimovsky, 1958:62 (key to species  in Alaska; southeast Alaska to C a l i f o r n i a ) . Cordone, 1959 ( l i s t i n g only),  Shapovalov, D i l l , and  C a r l , Clemens, and Lindsey, 1959:158  (description; P a c i f i c drainages from Chilkoot L,, Alaska to Ventura R, , C a l i f o r n i a ,  In B r i t i s h Columbia: lakes and r i v e r s of the  Columbia, Fraser, Dean, Skeena, Nass, and Stikine systems; coastal r i v e r s of the mainland and Vancouver Island, and Queen Charlotte Islands; headwaters of Peace R, system from Summit L, to McLeod L,). McAllister and Lindsey, 1959:70 (description; synonymy; i n t r a s p e c i f i c v a r i a t i o n ; l o c a l i t i e s as i n C a r l , Clemens, and Lindsey,1959),  M c A l l i s t e r , 1960:42 ( c o l l e c t i o n in s a l t water, Pt, Atkinson,  B r i t i s h Columbia).  Bond, 1961:36 (key to species i n Oregon;  p r i c k l i n g v a r i a t i o n ; Columbia R, drainage).  Bond, 1963:79 (synonymy  l i f e history observations; oxygen, temperature, and s a l i n i t y t o l e r ance of adults; f i s h associates; habitat preference; l i s t s 35 new c o l l e c t i o n l o c a l i t i e s i n Coastal Oregon, 30 l o c a l i t i e s from Columbia R. drainage i n Oregon, and also some lakes i n the southwest corner of Rainer National Park, Washington),  Bailey and Bond, 1963:19  (recognition of several species groups within Cottus i n western N. America; characters and l i s t of species i n the "asper species group"). Centridermichthys asper. - Richardson, 1844:76 (River Oregon = Columbia R.),  Gunther, 1860:170 (description; synonymy; fresh waters of the  Oregon and Washington T e r r i t o r i e s ) .  Lord, 1866a:130 [ l i f e  history  . 17 spawning behavior, i n part; Puget Sound; "streams flowing through the Sumass and Chilukweyuk p r a i r i e s " (Sumas R  0  and Chilliwack  R,  (?)), B r i t i s h Columbia; i n part, a l l streams east and west of the Cascades], Cottopsis asper. - Girard, 1851bs303 (introduction of Cottopsis  gen,  nov.; synonymy; l i m i t e d to River Oregon (= Columbia R,)).  Girard,  1851csl85 (not seen).  Girard, 1852s61 ( d e f i n i t i o n of Cottopsis  gen,  nov., based on Richardson's description of Cottus asper; synonymy; Columbia R.).  Girard, 1859s51 (description; synonymy; based on 8  specimens from Astoria and Fort Dalles, Oregon, and Fort Steilacoom, Puget Sound, Washington T e r r i t o r y ) .  Suckley (in Cooper and Suckley,  1859) 1959:351 (description; synonymy; small fresh water streams emptying into Puget Sound; Ft. Steilacoom; and Columbia R, 200 miles above mouth),  Suckley, 1860.351 (description; synonymy; small fresh-  water streams emptying into Puget Sound; Ft, Steilacoom; and Columbia R, 200 miles above mouth),  Jordan and Jouy, 1882s5 (Puget Sound;  Columbia R,; Mare Island and Sacramento R,, C a l i f o r n i a ) , Uranidea aspera. = Jordan and G i l b e r t , 1883s694 (description; synonymy; streams west of the Sierra Nevada and Cascade Mountains),  Jordan,-  1885sll0 (sub-genus Cottopsis and a l i s t of species therein), Trachidermus richardsoni, - He eke 1,, 1840 ;162 (synonymy; Columbia R.), (not Cottus richardsoni of Agassiz  B  1850).  Cottopsis parvus. ••<• Girard, 1856bsl44 ( o r i g i n a l description; Presidio (on San Francisco  Bay), C a l i f o r n i a ) .  Girard, 1857sll  P r e s i d i o , and Monterey, C a l i f o r n i a ) ,  (description;  Girard, 1859s54 (description;  synonymy; Monterey, Presidio, Fort Reading, and Petaluma, C a l i f o r n i a ) , Cooper, 1868s492.(listing Cottopsis  asper).  only),  Jordan1877§5  (as the young of  18 Centridermichthys parvus. - Gunther, 1860:170 (descriptionj synonymy; fresh waters of C a l i f o r n i a ) .  Lord, 1866b:352 ( l i s t i n g ; "frequenting  the same l o c a l i t i e s as..." Centridermichthys asper3. Uranidea aspera var. parvus. - Jordan and G i l b e r t , 1883:694 (Sacramento  R. forms),  Cottus gulosus parvus. - Jordan and Evermann, 1898:1945, and Jordan, Evermann, and Clark, 1930:383.(Cottopsis parvus: Monterey, Presidio, Fort Reading, and Petaluma, C a l i f o r n i a ) . Uranidea gulosa,  in part. - Jordan and G i l b e r t , 1883 :695  a l l specimens from "Vancouver's  (misidentifications:  Island" and probably those from  "about Port Townsend," c f . Jordan and Evermann, 1898:1944),  Kermode,  1909:87 ( l i s t i n g only; probable m i s i d e n t i f i c a t i o n : Shawnigan L., Vancouver Island). Cottus gulosus, i n part. - Jordan, 1895:141 ( m i s i d e n t i f i c a t i o n : San Luis Cr., near A v i l a , C a l i f o r n i a ,  c f . Hubbs, 1921:7).  Jordan and Evermann,-  1898:1945 ( m i s i d e n t i f i c a t i o n : a l l specimens from San Franciscito Cr., Santa Clara Co., C a l i f o r n i a ) . " Jordan, Evermann, and Clark"*. 1930':383 (probable m i s i d e n t i f i c a t i o n s : specimens from Loring and Boca de Quadra, Alaska).  Evermann and Clark, 1931:57 (misidentifications:  Presidio, Monterey, Fort Reading, Petaluma, and San Luis Cr,, California),  Evermann and Clark, 1931:12, 13 (misidentifications:  Monterey, P r e s i d i o , Fort Reading, and Petaluma),  Bean and Weed,  1920:76 (questionable i d e n t i f i c a t i o n : 4 specimens from V i c t o r i a , Vancouver Island, British,Columbia).  Wilimovsky, 1954:285 (doubts  v a l i d i t y of southeast Alaska record), Centridermichthys gulosus. - Lord, 1866b:352 ( l i s t i n g ; "frequenting the same l o c a l i t i e s as..."  Centridermichthys asper).  Cottus sp. - D i l l , 1946:54 (San Joaquin R, near Friant, C a l i f o r n i a : f i c a t i o n as asper by Hubbs and Wallis, 1948:141).  identi-  19 Nomenclatural History The s p e c i f i c name asper i s currently well founded i n the genus Cottus to which i t was o r i g i n a l l y designated by Richardson i n 1836,  t  But, as shown  in the preceding synonymy, the binomen was extremely unstable f o r the f i r s t 100 years a f t e r . i t s introduction,,  After an i n i t i a l period of uncertainty  regarding i t s a f f i n i t y to marine or to freshwater Cottoids, three main nomenclatural d i f f i c u l t i e s are encountered? as Cottus gulosus (Girard)j a  n  d  the often-repeated m i s i d e n t i f i c a t i o n  the f a i l u r e to recognize that Cottopsis parvus  Cottus asper are conspecific; and the f a i l u r e t o recognize the s p e c i f i c  relationship of asper to other species i n the genus Cottus, The almost immediate placement  of asper into Trachidermus by Heckel  (1840), and then into Centridermichthys by Richardson (1844), r e f l e c t s the early opinion that asper was more closely a l l i e d to the marine Cottoids, Girard (1851b; 1852) recognized i t s a f f i n i t i e s with the freshwater genus Cottus, but distinguished i t from that genus by erecting the genus Cottopsis, based on the presence of palatine teeth and the "skin beset with p r i c k l e s , Instead.of being smooth and s c a i e l e s s "  Lacking any specimens, Girard de-  0  fined Cottopsis on the basis of Richardson's o r i g i n a l description but, on p  0  63, he misquoted Richardson e s p e c i a l l y i n regard to the absence of scales,,  Girard's lack of specimens proved unfortunate since soon thereafter (1856b) he named and described Cottopsis" parvus C a l i f o r n i a , comparing i t not w i t h C Girard, also newly described (1856a) In his l a t e r report (1859) session, from the Columbia  9  e  from the Presidio (in San Francisco),  asper, but with Cottopsis^ gulosus B  from the San Joaquin R,,  California,  Girard had 8 specimens of C,. asper i n h i s posR, and Puget Sound,  Obviously he again f a i l e d to .  recognize the c o n s p e c i f i c i t y of asper and parvus and he followed Richardson's o r i g i n a l description rather than compare them with specimens of parvus, which he seems to have reserved f o r comparison with gulosus, Jordan (1877) referred^, in passing, to Cottopsis parvus as the young of  20 C. asper,  Jordan and Jouy (1882) however, listed specimens of Cottopsis  asper from Mare Island and Sacramento R,, California, and from Puget Sound and the Columbia R,  Lass than a year later, Jordan and Gilbert (1883)  placed asper into the genus Uranidea DeKay, subgenus Cottopsis, based on the presence of palatine teeth, and the g i l l membranes being broadly united to the isthmus.  In the same report, they refer to the Sacramento R, form of • > ,  U, aspera as "var, parvus, smaller in size, paler in color and with the interorbital space concave, narrower than eye," Eigenmann (1895) used the valid name to describe specimens from the Fraser and Columbia R,, as also did Gilbert and Evermann (18.95), who suggested that the nominal species was separable "at least subspecifically from the Sacramento River form,"  Seale (1896), and Gilbert (1896), used the  valid name for northern specimens.  But obviously, Jordan (1895), and Jordan  and Evermann (1896), s t i l l thought in terms of a distinct Californian^species (gulosus),  and a distinct northern species (asper).  Jordan misidentified  a specimen of asper from San Luis Cr.?near Avila, California, as gulosus. Jordan and Evermann listed the range of the nominal species from Vancouver Island to Oregon, and of gulosus, from California Coast Range streams and inland in the San Joaquin R. Gilbert (1896) referred to Cottus asper of the Columbia and Cottus gulosus_ of the Sacramento as "two species so extremely similar that i t is d i f f i c u l t to distinguish them,"  Jordan and Evermann  (1898) repeated the suggestion of Gilbert and Evermann that the nominal species i s separable, at least subspecifically, from the Sacramento R, form "Cottus gulosus." That Jordan* and Evermann perceived neither the conspecific relationship °f parvus to asp_er nor the limits of the valid species Cottus gulosus, be9  comes more fully evident on the next page (p, 1945) of the 1898 report. Their description of Cottus gulosus (Girard) i s taken from misidentified specimens of C. asper collected in San Franciscito Cr,, Santa Clara Co.,  California.  These were large specimens "3 to 7 inches i n length" and, most  s i g n i f i c a n t l y , the count f o r anal rays i s given as "A, 16 to 18." these characters separate asper from gulosus.  Both of  Furthermore, they include  Cottopsis parvus Girard, from Monterey, the Presidio, Fort Reading, and Petaluma, C a l i f o r n i a , i n the synonymy of gulosus. Snyder (1905) collected and correctly i d e n t i f i e d Cottus asper from the same l o c a l i t y , San Franciscito Cr.  He was probably the only one of his time  to understand and explain the true r e l a t i o n s h i p s of asper, parvus, and gulosus.,  On p. 337, he states?  Recent authors have i d e n t i f i e d the common Sacramento form which represents the Cottus asper of the Columbia River with the Cottopsis gulosus of Girard. They have sometimes considered the Sacramento form as i d e n t i c a l with C. asper and have placed the name gulosus i n the synonymy of the l a t t e r . At- other times they have considered the species as a s l i g h t l y d i f f e r e n t i a t e d form worthy of recognition i n nomenclature, and have used the name gulosus to designate i t . The former view concerning the species i s probably correct. The association of the name gulosus with i t , however, i s without warrant. The l a t t e r belongs to a species e a s i l y distinguished from C. asper d i f f e r i n g notably i n having a much shorter anal f i n . There are usually fewer dorsal spines and r a y s a more limited d i s t r i b u t i o n of p r i c k l e s , and an almost uniform absence of palatime teeth. In C. asger the dorsal has 8 to 10 spines and 19 to 22 a r t i c u l a t e d rays, Fhe anal 16 to 18 rays, while i n C, gulosus the dorsal has 7 to 9 spines, 17 to 18 rays, the anal 12 to 14 rays, B  B  —  1  Snyder then continues with a note on habitat preference: In i t s d i s t r i b u t i o n C. asper appears to be largely confined to the lower courses of the streams", being e s p e c i a l l y abundant near t i d e water, while C. gulosus i s found further up, where the water i s c l e a r and the current rapid Rutter (1908) c o r r e c t l y placed Cottopsis parvus into the synonymy of Cottus asper, presumably recognizing that the two were conspecific. However, he i n c o r r e c t l y synonymized Uranidea semiscabra centropleura Eigenmann and  ;  Eigenmann, which i s properly referable to Cottus gulosus. Snyder was the f i r s t to consider a s e r i e s of specimens of the nominal species throughout i t s entire range, as then known, and, in the same issue of the B u l l . Bur. Fisheries i n which Rutter had c o r r e c t l y synonymized parvus, he noted the extreme variation of p r i c k l i n g investment.  While recognizing  22 the variation between streams, he also noted that the p r i c k l i n g v a r i a t i o n i s common among individuals from the same stream. Although e x p l i c i t l y aware of Snyder's comments on asper and gulosus, Evermann and Goldsborough  (1907&)identified 16 specimens of gulosus from,  Loring and Boca de Quadra, Alaska, The r e l i a b i l i t y of these i d e n t i f i c a t i o n s i s questionable.  Kermode's l i s t i n g (1909) of Uranidea gulosa from Shawnigan  L,, Vancouver Island, i s probably a m i s i d e n t i f i c a t i o n of £, asper. Snyder (1913j 1916) again recorded the occurrence of C, asper and gulosus i n d i f f e r i n g habitats of the same stream,  Hubbs (1921) recognized  Jordan's e a r l i e r m i s i d e n t i f i c a t i o n of gulosus from San Luis Cr,, C a l i f o r n i a . He also commented on the v a r i a b i l i t y of p r i c k l i n g in £. asper from several streams. It would seem that with the accession of Snyder's insight into the problem, the v a l i d name was destined f o r s t a b i l i t y .  However, Jordan,  Evermann, and Clark (1930) repeated the e a r l i e r error of Jordan and Evermann (1898) by including Cottopsis parvus as a synonym of Cottus gulosus. similar manner, they also incorporated the e r r o r of Evermann and  In a  Goldsborough  (1907a):,, previously c i t e d , by including the misidentif ied specimens of asper from Loring and Boca de Quadra, Alaska i n the l i s t of records f o r gulosus,In the same work, Jordan and Evermann extended the range of asper (cited in 1898 as "streams of the Cascade Range, from Vancouver Island to Oregon") by appending the phrase "south to Sacramento River,"  Evermann and Clark (1931)  also retained Cottopsis parvus in the synonymy of C. gulosus and perpetuated Jordan's m i s i d e n t i f i c a t i o n of the San Luis Cr. gulosus, which Hubbs had c o r r e c t l y r e - i d e n t i f i e d as asper ten years before (1921),  >  Schultz and DeLacy's catalogue (1936) included a comprehensive l i s t i n g of Washington and Oregon l o c a l i t i e s f o r C, asper.  However, some of the  l o c a l i t i e s are doubtful since they frequently misidentified C, asper as gulosus and/or perplexus.  They also i n c o r r e c t l y maintained the presence of  23 C_. gulosus i n Alaska. Robins and M i l l e r (1957) presumably overlooked the e a r l i e r c i t a t i o n of Rutter (1908) and removed Cottopsis parvus from the synonymy of gulosus, placing i t i n synonymy with asper, supposedly f o r the f i r s t time. McAllister and Lindsey (1959) f i r s t suggested the probable existence of "coastal" and "non-coastal" populations of Cottus ajper on the basis of morphological and, perhaps, behavioral differences. Bond (1961) hinted at the p o s s i b i l i t y of polytypy in Cottus asper when he stated i n his key that the body i s "well covered with p r i c k l e s , e s p e c i a l l y in inland waters and in young individuals from coastal waters.  Bond (1963)  gives the most comprehensive treatment yet recorded f o r Cottus asper and 12 other species i n the genus.  His- study  8  however, i s more concerned with  <  i n t e r s p e c i f i c rather than i n t r a s p e c i f i c r e l a t i o n s h i p s within the genus, <.*•., Bailey and.Bond (1963) have indicated t h e i r concern f o r the supraspecifie relationships within the genus Cottus by t h e i r recognition of several species groups, one of which i s the "asper species group." Distribution Range. - P a c i f i c Slope Drainage of North America? Seward, Alaska, to Ventura R..  Coastal streams from  C a l i f o r n i a ; lakes and streams of the Queen  ;  Charlotte Islands and Vancouver Island; and a l l major P a c i f i c drainages from the headwaters of the Stikine R. in B r i t i s h Columbia, to the Kern R._  San  Joaquin R. drainage. C a l i f o r n i a n - The-following are exceptions? Fraser R. .in the area of the Rocky Mountain Trench, east of Prince George, B r i t i s h (area not yet c o l l e c t e d ) ; Kootenay Lake drainage of the Columbia  Columbia  R, in B r i t i s h  Columbia; Upper Snake R. of the Columbia R, drainage i n Washington and Oregon; Middle Fork of the Willamette R. in Oregon, above Oakridge; Klamath R, Basin in Oregon; and Sacramento R, drainages above Lake Shasta, C a l i f o r n i a ,  Arctic  Slope Drainage of North America? Headwaters of the Peace R. in B r i t i s h Columbia?' from Summit L. to McLeod L. Crooked R. drainage; from Tacheeda L., 9  24 Parsnip R. drainage; from Tchentlo L.* Nation R, drainage. The present d i s t r i b u t i o n a l range of Cottus asper i s i l l u s t r a t e d i n F i g . 3. Results;  Analysis of Geographic Variation  P r i c k l i n g Distribution. - As previously defined (p. 7 ) » the " v i r t u a l " p r i c k l i n g represents the sum of two component percentages, i . e . , the "apparent" p r i c k l i n g and the resorbed p r i c k l i n g .  F i g . 4 i l l u s t r a t e s graphically  that mature specimens from populations of C_, asper from coastal l o c a l i t i e s are considerably less p r i c k l e d , both apparently and v i r t u a l l y , than inland localities.  However, most inland populations from the Peace, Nass, S t i k i n e ,  and parts of the Skeena R. drainage systems are coastal i n their p r i c k l i n g affinities. derivative."  Hereafter, these populations w i l l be referred t o as "coastal Likewise, certain populations from lakes on Vancouver Id, and  the Queen Charlotte Islands (Q C,I.) are also coastal in t h e i r p r i c k l i n g 0  affinities.  Fig, 4 a l s o indicates that the population from  Carquinez  S t r a i t , at the mouth of the Sacramento R. i n San Pable Bay, C a l i f o r n i a , has a mean " v i r t u a l " p r i c k l i n g percentage-greater  than that of any coastal popu-  l a t i o n , and lower than a l l but one of the inland populations.  For comparative  purposes, the Carquinez S t r a i t population w i l l hereafter be referred to as the "intergrade" population. Table I summarizes the mean percentage a r e a l p r i c k l i n g and resorption f o r a l l populations grouped into aggregates according to l o c a l i t y and prickling affinity. If the data f o r specimens under 60 mm S,L, are considered separately, there i s a s i g n i f i c a n t difference i n mean percentage p r i c k l i n g , both "apparent" and " v i r t u a l , " between the coastal and inland populations.  Ex-  pressed i n terms of a r a t i o between coastal-inland populations, a r a t i o of "apparent" p r i c k l i n g 1 s 1.8, and " v i r t u a l " p r i c k l i n g 1 ? 1,5, i s obtained. The aggregate means f o r the "coastal d e r i v a t i v e " populations are e s s e n t i a l l y  Figure 3.—Distributional Range of Cottus asper.  26  Figure 4„—Virtual, Resorbed, and Apparent Prickling in Cottus asper.  TABLE I.  Mean Percentage Prickling and Resorption Among Aggregate Populations of Cottus asper.  Aggregate Population Designation  Virtual  2 Coastal Derivative  Inland * 1  Apparent  it  1 Coastal  Intergrade  Specimens <60 mm S.L.  3  Virtual  Specimens ^60 mm S,L, % Resorption Apparent Relative Absolute  43.5 (9)  36,0 (12)  35.0 (16)  17.8 (19)  48.9  17.2  44.8 (6)  41,8 ( 6)  34,7 ( 6)  24.2 ( 6)  32.0  10,5  60.1 (1)  57,1 ( 1)  52,9 ( 1)  35,6 ( 1)  32.8  17.3  67,2 (16)  65.3 (18)  63.2 (18)  53.2 (19)  16.1  10.0  Brackets enclose numbers of population means in determination of aggregate mean percentages. 1  Populations 1-31, l i s t e d in Appendix Table 1  2  Populations 32-43, listed in Appendix Table 1  3  Population 73, listed in Appendix Table 1  4  Populations 44-72, l i s t e d in Appendix Table 1  28 the same as those f o r the c o a s t a l , and they d i f f e r s i g n i f i c a n t l y from the aggregate means f o r the inland populations. population  The mean for' the intergrade  l i e s closer to the inland aggregate.  When specimens larger than 60 mm S.L. are considered, the same magnitude of difference i s noted between the coastal and inland " v i r t u a l " percentages, i . e . , a r a t i o of 1 s 1.8, but the "apparent" p r i c k l i n g percentages are even more divergent,  having a r a t i o of 1 s 3,  constant, the divergent  Since the " v i r t u a l " r a t i o remains  "apparent" r a t i o s indicate a d i f f e r e n t i a l percentage  resorption between mature specimens from coastal and inland  populations.  This d i f f e r e n t i a l i s further suggested by the inverse relationship which e x i s t s between the mean percentage "apparent" p r i c k l i n g and the mean percentage r e l a t i v e resorption.  It indicates that i n coastal populations a  greater percentage o f the " v i r t u a l " p r i c k l i n g area i s being resorbed than i n inland populations.  The r e a l i t y of t h i s d i f f e r e n t i a l resorption i s shown i n  the figures f o r mean percentage absolute resorption which indicate that, i n coastal forms, resorption occurs over almost twice as much area as i n inland forms.  The d i f f e r e n t i a l can be expressed i n terms of a coastal-inland r a t i o ,  i«e©9 Xe 7 t Xe P r i c k l i n g and resorption percentages on which Table I i s based are given for a l l populations i n Appendix Table 2. Population analysis of the non-weighted mean percentage " v i r t u a l " p r i c k l i n g i n each of the 14 body sections, of specimens ^ 60 mm S.L., i n dicates that i n coastal and "coastal derivative"populations,  body sections  'a', 'k', '1', and 'n' are completely naked except f o r the rare occurrence of an isolated p r i c k l e .  S i m i l a r l y , in coastal and "coastal d e r i v a t i v e " popu-  l a t i o n s , only small portions of sections prickled.  'b', 'd', ' e , 'h', and 'm' are ?  When compared to the mean percentage " v i r t u a l " p r i c k l i n g of inland  populations,  a l l nine of these sections are s i g n i f i c a n t l y d i f f e r e n t .  Further-  more, i n the remaining f i v e sections C c ' , ' f ' , 'g», ' i ' , and ' j ' ) , the mean  29 percentage " v i r t u a l " p r i c k l i n g i s always higher i n the inland  populations.  These r e s u l t s corroborate the suggestion, made for the aggregate  population  means, that the areal d i s t r i b u t i o n of p r i c k l i n g i n coastal and inland popul a t i o n s of C. asper i s d i f f e r e n t .  Data for t h i s analysis are presented i n  Appendix Table 3. P r i c k l i n g Intensity. - In general, the i n t e n s i t y of p r i c k l i n g , i . e . , the number of p r i c k l e s per unit area, i s greater i n inland f i s h than i n coastal f i s h .  In coastal populations,  where p r i c k l e s are almost never pre-  sent i n sections 'a', 'k', '1', and 'n', no comparison of i n t e n s i t y can be made.  Rarely, however, when an isolated p r i c k l e or group of prickles occurs  in one of these sections, the i n t e n s i t y i s always l e s s than that of the corresponding section in an inland.population.  The same holds true f o r  sections 'b', 'd*, 'e', 'h', and.''m'.f The comparison of i n t e n s i t y must therefore be made i n one of the sections i n which p r i c k l e s are always present on both inland and coastal specimens,- Section tive.  ' j ' was chosen as representa-  Table I I shows that the mean r e l a t i v e i n t e n s i t y i s greater in the  inland populations than i n coastal  or "coastal derivatives,"  Interpolation  of mean i n t e n s i t y values given i n Table II into approximate number of prickles per sq, cm. (cf. Methods) y i e l d s counts of ca. 1325 p r i c k l e s per sq. cm. i n coastal forms (2.94), and ca. 1820 per sq, cm, i n inland forms/fo.se). Expressed i n terms of differences between coastal and inland i n t e n s i t i e s , a . r a t i o of 1 ; 1.4 i s obtained.  Comparison to the r a t i o obtained f o r area:  p r i c k l i n g resorption, I.e.", 1,7 ; 1^ indicates that even though inland f i s h have more p r i c k l e s per unit area, more resorption basis) occurs i n coastal f i s h .  (on a p r i c k l e f o r p r i c k l e  Mean r e l a t i v e i n t e n s i t y values for i n d i v i d u a l  populations of Cottus asper are given i n Appendix Table 4, Table II also shows that i n mean intensity, of p r i c k l i n g , the "coastal derivative" populations aggregate i s near the coastal mean whereas the "intergrade" population  i s closest to the inland mean.  I f the mean i n t e n s i t y  TABLE II.  Mean Relative Prickling Intensity for Populations of Cottus asper Aggregated by,Locality and Prickling Affinity.  Skin Section ' j * .  Aggregate  Specimens <60 mm S.L,  Specimens ^60 mm S.L.  Population  Sample  Intensity  Sample  Designation  Size  Mean (Range)  Size  136 (13)  2.94 (1-5)  235 (18)  2,64 (1-5)  103 (11)  2.74 (1-4)  43 ( 7)  2.95 (1-5)  Intergrade  8(1)  3,75 (3-5)  20 ( 1)  3.45 (2-4)  Inland **.  238 (21)  3.86 (2-5)  265 (19)  4.20 (3-5)  Coastal  1  ft  2 Coastal Derivative 3  Intensity Mean (Range) ft  * Brackets enclose numbers of populations included in aggregate sample. 1  Populations 1-31, listed in Appendix Table 1  2  Populations 32-43, listed in Appendix Table 1  3  Population 73 only, listed in Appendix Table 1  4  Populations 44-72, listed in Appendix Table 1  31 values of certain coastal lake forms, i.e., Home L., Buttle L., Skidegate L., and Chilkoot L., are considered (Appendix Table 4), they are found to be slightly lower than the coastal means.  These data corroborate the differ-  ences already shown to exist in mean areal distribution of prickling between coastal and inland populations. • Pectoral Fin Asymmetry, - Of 59 populations investigated for pectoral f i n asymmetry, 56 had some specimens with asymmetrical pectoral f i n element counts.  Of 1163 paired counts, from 59 populations, 23.6% (275 pairs) were  asymmetrical  (Table III). This percentage incidence of occurrence i s similar  to the percentages reported for other cottids by Hubbs and Hubbs (1945)., Of the 56 populations of Cottus asper exhibiting asymmetry, 39 (69.6%) were s i n i s t r a l , 11 (19.6%) were dextral, and 6 (10,7%) had equal numbers of right and l e f t asymmetrical fins.-  This apparent direction of asymmetry  within populations i s not statistically significant since the sample sizes of asymmetrical fins within each population were very small.  However, if-,  the asymmetrical fins are grouped according to locality and supposed de-  r  rivation (based on prickling), or pooled for a l l populations (Table III, columns 5 6 6), the overall direction of asymmetry in Cottus asper is signi2  ficantly sinistral ( X = 21.56, cC--= .05, 1 d.f., for the pooled sample). While Cottus asper i s significantly sinistral in the overall direction of i t s asymmetry, the amount of sinistrality within populations varies from 0 to 100%.  When the populations are grouped according to locality and  supposed derivation (Table 111^ col. 5), the amount of sinistrality for 24 coastal populations i s 66.7%; for 12 coastal derivative populations, 69.2%; and for 23 inland populations, 60.8%. These differences among aggregate populations are not significant.  Additional analysis of data on a north-  south basis revealed no significant differences in direction or amount of asymmetry. Total pectoral f i n counts on which Table III i s based are given for a l l  TABLE III.  Incidence of Occurrence, Direction, and Amount of Pectoral Fin Asymmetry Within and Among Aggregated Populations of Cottus asper.  Locality ft  No. Pairs  Asymmetrical  ftft  Sinistral  Dextral  367 (79,8)  93 (20.2)  62 (66.7)  31 (33.3)  113 (74.4)  39 (25.6)  27 (69.2)  12 (30.8)  551  408 (74.0)  143 (26.0)  87 (60.8)  56 (39.2)  1163  888 (76.4)  275 (23.6)  176 (64,0)  99 (36.0)  Coastal  460  Coastal Derivative  152  Inland Pooled Total  Symmetrical  -  Includes three coastal island lake populations aft Bra eke t s en close aggregat e mean percent age s  33 populations in Appendix Table 5. It has been shown for several species of fish that the lowermost pectoral ray i s shorter on the side which has the higher count (Hubbs and Hubbs, 1945} Hubbs, 1963).  Cottus asper appears to follow this rule.  Of  69 specimens, from 17 populations having asymmetrical pectoral f i n counts, 66 had a shorter lowermost ray on the side with the higher count, and only 3 had the lowermost rays equal on each side. While Cottus. asper demonstrates a significant sinistrality in the .  .j  direction of i t s pectoral f i n asymmetry, the amount of this s i n i s t r a l i t y within and among populations cannot be used to separate individual populations or geographically aggregated groups of populations. Pectoral Fin Ray Count. - The mean number of pectoral f i n rays i n creases progressively from south to north in the inland populations (Figs, 5 & 11).  Coastal populations show a similar increase northward only to about  42-44° N. Latitude where the mean pectoral f i n ray counts diverge from the inland counts and decrease toward-the north.  Mean counts for the "coastal  derivative" population are found to be equal to or less than inland population means, but always higher than those of the coastal means, at the same latitudes.  The means of certain coastal island lake and coastal main-  land lake populations (Buttle, Home, Skidegate.and  Owikeno L., cf.  Appendix Table 1) are almost a f u l l count higher than the means of nearby stream populations from which they probably were derived.  Data from  Carquinez Strait "intergrade" population and two additional populations from the San Francisco-San Pablo Bay area, thought to be intergrades, (cf, Appendix Table 1, population numbers 74 6 76) were compared.  At the low  latitudes in which they occur, their mean pectoral ray counts were indistinguishable from either coastal-or inland means. Spiny Dorsal Fin. - The average spine number is clustered about nine for a l l coastal populations except:the southernmost California populations  34  I  60 - O 58 56 54  "o  52 50 I _ l_48  I  I  I  o o  z  4  O  AA  K  J  O  40 A  AAA  e  A  4 A  A  O  0G  O  I  A  A G O  42  I  I  ,^A®  o  o 44  34  I  iA A A  Q46  36  I  o o e o o oo ®  cr  38  I  I  J  A  o o  o I  I  16  _L  L.  O  COASTAL  ®  COASTAL DERIVATIVE  @  COASTAL  A  INLAND  A INTERGRADE  I  PECTORAL  LAKE  17  RAYS  Figure 5„—Mean Pectoral Fin Ray Count by Latitude,  I  35 which are s l i g h t l y lower ( F i g , 5 )  0  The San Joaquin-Sacramento. R« inland  populations have a mean count o f eight or s l i g h t l y higher.. This wide d i v e r gence tends to accentuate a s l i g h t , but d i s c e r n i b l e , geographic c l i n e f o r the inland populations as a whole.  The mean counts are s l i g h t l y higher than nine  in the high l a t i t u d e s , s l i g h t l y lower i n the mid-latitudes, and much lower i n the low l a t i t u d e s (cf. F i g . 11), A comparison of the ranges (Appendix Table 6) indicates high counts of nine to eleven i n the north, and low counts o f seven to nine i n the south of the d i s t r i b u t i o n a l range of the species. : The intermediate positions of the three "intergrade" populations within the San Francisco-San Pablo Bay area should be noted.  The means of the "coastal de-  r i v a t i v e ' populations tend to be closer to those o f the inland populations. 1  Softs Dorsal F i n . - The mean soft-ray number varies considerably with latitude.  Counts are s l i g h t l y higher than 20 i n the high l a t i t u d e s , closer  to 21 in the mid-latitudes, and between 19 and 20 i n the low l a t i t u d e s .  As  can be seen i n F i g . 7, the inland Sacramento and, e s p e c i a l l y , the San Joaquin R. populations have considerably lower mean counts than t h e i r coastal counterparts at the same l a t i t u d e s .  I f the soft-ray counts f o r a l l inland popu-  lations are summed into aggregates o f 2-degrees latitude and plotted according to the Hubbs-Hubbs format ( F i g . 8), the V-shaped geographic c l i n e within the means and the ranges becomes immediately  apparent (cf. F i g . 11),  Combined Dorsal Fin Elements, - Combining the spiny and soft dorsal elements tends to minimize the variation i n means at the middle of the range, while accentuating the divergence between coastal and inland populations i n the highland, e s p e c i a l l y , the low latitudes (Fig. 9), The divergence between the San Joaquin-Sacramento populations and t h e i r coastal counterparts at the same l a t i t u d e i s e s p e c i a l l y enhanced (cf. F i g . 11), The intermediate p o s i tion of the intergrade population means and the position of the "coastal der i v a t i v e " means w i l l be discussed l a t e r . Anal F i n . - The mean number of anal rays varies around 17 f o r a l l  36  0  60  58  O  -  56 54  O  OO -  ^ A  (X)  52 50  HI cr  O z  o  -  48  O  46  A  A  o  M  44 42 40 38  o  36 34  o  o O o A O o o O o  O  COASTAL  ®  COASTAL DERIVATIVE  O  COASTAL  A  INLAND  LAKE  A INTERGRADE  10  8  DORSAL  SPINES  Figure 6,—Mean Spiny Dorsal Fin Ray Count by Latitude,  60 58  i  —i  0  O  •  O  1  i  1  56  O A #>A  54 52  (8)  •  -  A G  A 50  A.^e  x JZ48 cr Q  4 6  z  o  A  -  e  <* A  -  44  &  4)  A  A  0  0  42  -  40  -  38  -  36 34  A A A  A  A  - G  0 °  I  £  COASTAL LAKE  A INLAND 1  ^~  20  SOFT  COASTAL  ® C O A S T A L DERIVATIVE  ,o 1  19  i  03  A  A  A 1  -  4  A  1  o  A  DORSAL  Figure 7.—Mean Soft Dorsal Fin Ray Count by Latitude.  I  21  RAYS  -1-  A INTERGRADE  22  38 Standarcy—^  fMean  Deviation^  h  .•  A  ,-2 Standard of  the  Errors  Mean  - - 7 /  Range-'''  57h  55 53f  51 49  cr X  O  45h  43 41 39 37 358  19  SOFT  20  DORSAL  21  RAYS  Figure 8,—Hubbs-Hubbs Plot of Soft Dorsal Fin Ray Counts for Inland Population; Samples Pooled into Aggregates of.Two Degrees Latitude,  22  T  60  r  r  1  o o  58 56  <8>  54  o  o  e  52  e  50  H  4  8  o  or  0 Z  46 A  •  O  O  o 44  O  o  40  A  A A  38  O  36 27  _i  L  O  9,—Mean Combined Dorsal  A  o  O O  O  28  C O M B I N E D Figure  G ?  O  42  34  O  o  29  J  D O R S A L  F i n Ray Count by Latitude,  L  O  COASTAL  ®  COASTAL DERIVATIVE  ©  COASTAL LAKE  A  INLAND  A  INTERGRADE  30 E L E M E N T S  40 populations (Fig. 10).  There is a tendency for the mean to be slightly  lower in the northern and southern ends of the distributional range, whereas in the mid-latitudes the means are clustered about 17.  This pattern tends  loosely to follow the V-shaped curve observed in the number of soft dorsal rays (cf. Fig. 11). Pelvic Fin. - The pelvic f i n almost invariable consists of 1 spiny plus 4 segmented rays.  Among more than 1000 specimens examined for this charac-  ter, there were only 20 deviants from-the usual left-right count of 4-4, , In one specimen both fins, and another specimen, the l e f t f i n was missing. In neither specimen was there any apparent injury.  Of the remaining 18, 7  deviants (4 damaged) were on the left side, and 11 (4 damaged) on the right. Thus even in the most stable meristic character, rare deviations occur, A summary of meristic counts for a l l populations i s given in Appendix Table 6.  ,  s  Latitudinal Variation in Ranges of Mean Fin Ray Counts. - Figure 11 is derived from each of the Figures: - 5 , 6, 7, 9, and 10.  When the means of  the inland and the coastal populations are enclosed by sets of lines approximating the range of means, a family of curves representing a l l the meristic data for a l l latitudes i s generated.  Several trends are apparent in  Figure l i s 1)  The curves for inland and coastal population  means, though over-  lapping in some areas, are widely divergent in others; 2)  A l l curves, except for the spiny dorsal and the inland pectoral  means, demonstrate a characteristic V-shaped pattern with an inflection occurring at about 42-44° N. Latitude; 3)  For pectoral fins, the greatest amount of divergence between  coastal and inland means exists in the high latitudes.  The opposite i s true  for the dorsal f i n elements; and 4)  In areas where coastal and inland curves are divergent, the means  41  -i  1  r  -i  60  1  1  o  ,  J  T  r  1  1  r  o  o  58-  _  r  56-  <8>  o  <2>  54-  52  8  50  o8  48 h-  e ^° 0  *46  AO  O o  4 4  |  G  40 38  O O  42  o  cP  34  6  A  o A  A  o o  36  A  _l  o  A N A L  I  J_  O  COASTAL  ®  COASTAL DERIVATIVE  0  COASTAL L A K E  A INLAND A INTERGRADE  18  17  16  .  F I N  R A Y S  Figure 10,—-Mean Anal Fin Ray Count by Latitude,  15  16  10  8  17  Spiny Dorsal  Pectoral  18  20  22  26  28  Soft  Combined  Dorsal  Dorsal  30  16  17  Anal  Figure 11.—Approximate Ranges of Mean Fin Ray Counts Plotted Against Latitude for Inland ( Coastal ( ) and Coastal Lake and Coastal Derivative CciiiiBiiiiii) Populations. Curves f i t t e d by eye from Figures 5, 6 7, 9, and 10. B  g  18  43 for coastal lake and "coastal derivative" populations always approximate the inland rather than the coastal curve. The interpretation of these curves and the similarity of their characteristic V-shape to curves generated from laboratory rearing experiments under controlled temperatures will be considered in the discussion on meristic variation within Cottus asper (below). Lateral Line. - The number of lateral line pored-scales varies directly with s i z e up to about 60 mm S.L., the average size at which most Cottus asper f i r s t become reproductively mature (Table IV),  In most specimens  larger than 60 mm, the pored-scale count approaches, or equals, the "comvi  plete" lateral line count.  One obvious exception i s in a spawning popu-  lation collected from the estuary of a small stream in Juskatla Inlet, Queen Charlotte Islands, in which many of the small specimens 40-49 mnu S.L. were in spawning condition. Their pored-scale counts approached or , r  equalled the mean "complete" lateral line count (Table IV). Another exception to the generalization i s that afforded by specimens from the coastal lake populations found on the British Columbia or Alaska mainland, or on the offshore islands.  Most mature specimens are characterized by their low  pored-scale counts (Table IV).  Of the coastal lake populations, small  specimens were available only from Skidegate L.  A l l of these were im-  mature and a l l had low pored-scale counts, but the "complete" counts of this, and other coastal lake populations, were equivalent to the mean counts for other coastal and inland populations. Tentative analysis of mean "complete" lateral line counts indicates a tendency toward higher counts in the north.  Paucity of counts throughout  the entire range of Cottus asper precludes any further comparisons at this; time. Discussion of.Meristic Variation Within Cottus asper Lindsey (1961; 1962) has reviewed literature which demonstrates that  TABLE IV.  Lateral Line Pored-Scale  Population  Size  Saltery Cr. A-laska **"complete"  ^  6  0  Counts by Size Groups f o r Selected Representative  1 7  1 8  1 9  2 0  2 1  2 2  2 3  2 4  2 5  2 6 _  e n o  • o e s  oo  «o  o  o  o  ==>  — <•  — o  oo  oo  o—  oo  oo  o  o  o  ==  oo  oo  oo  =o  oo  = =  oo  oo  oo  ==  oo  ==  -o  =o  =  o  2 7  2 8 ^  2 9  —  = o  o  3 0  3 1  3 2 _  33  3 4  Populations o f C. asper  3 5  _  3 6 _  11  =  3 8  3 9  4 0  3  1  4 1 _  9  3  3  2  4 2 _  —  4 3 _  .  »  ~  *  X = 4 0 . 0 6  Cultus L o B.C. "complete" X = 4 0 . 2 7  5 0 - 6 0 4 0 = 4 9 3 0 = 3 9 2 0 = 2 9  Graham I d . QCI Juskatla I n l e t "complete " g  X = 3 9 . 3 3  5 * 6 0  X = 3 9 . 7 Q  Buttle L . Van. Id. "complete"  =,=  —  ==  —  =-  —  —  ==. —  1  ~  "  "  oo  oo  oo  < = c  oo  oo  _  oo  —  -  o o  - - •  =  ^  1  =  oo  —  4 0 = 4 9  = -  ==  oo  «=  6  0  —  =  —  —  -  = =  ==  ==  ==  oo  oo  o  oo  = =,  oo  oo  oo  o  —  *  ~ =  o  o —  o  oo  oo  oo  o  — o  oo  o  —  —  ™-  —=  —  — —  —=•  —  —  == 1  ——  3 0 = 3 9  =  = ~  1  —  2  1  1  —  -=  •»=  —  1  ==  =•=  —  o t t o  1  —  ==  OQ  ="•= ==  —  ==.  =«=  •=•=  ==.  OO  1  —  3  —  ,  — ~  —  —  1  1 1  2  2  -=  1  OO  1  1  1  2  M  1  IB<B  „„ 1  —=  —  1  2  1  2  1  1  —  4  1 3  3  2  3 4  3  1 9  5  1 5  =-  3  2  ~  <=„  - =  3  ==  =•"=  ==  2  ==  —  1  1  - —  -=  —\  1  == 2 - - .  1  —  1  —  •<=> «-.<=  o o  1  OS  —=  —  4  3  J = « =  =~  2  ==  ==  >=•=  —  = =====  ==  o c a ,  —  •="• —  «=-  3  CO  .  =  1  =—  1  ~  ==  ==  — =  = • =  ==  ==•  ==  ==  •==  —«=  0 CB-3  ova-  • > < > COB  e-o  ==  B O  = —  DCS  4 0 - 4 9  - -  X = 3 9 . 7 1  Horne L . , Van. Id. "complete"  2 ^ — > •  —  —  <==.  —  —  —  — ~  2  1  ==  1  5 0 = 5 9  6  —  ==•  ==•=  *  4 0 = 4 9  >  OO  -=  ~=  3 0 = 3 9  >  1  •=»  — —  =  5 0 = 5 9  2 0 = 2 9  Moresby Id., QCI Skidgate L . "complete''  «  >  6  0  oo  .  _  —  ~  —  1  1  2  2  2  " 2  -=  —  - = =  = =  .,  5T=39o45  * "complete" counts are mean counts o f l a t e r a l l i n e pored-scales plus unenclosed neuromast h i l l o c k s . ** Saltery Cr. represents a t y p i c a l coastal populationi Cultus L. represents a t y p i c a l inland population; Juskatla Inlet represents a coastal i s l a n d stream population; Skidegate,Buttie, and Horne L, represent coastal lake populations  f  45 in fishes reared experimentally over a wide range of temperatures, a Vshaped, or inverted V-shaped, curve i s t y p i c a l f o r meristic s e r i e s , esp e c i a l l y vertebrae. Generally, an intermediate temperature produces the minimum count, but sometimes, a maximum count.  Lindsey (1962) states that  the i n f l e c t i o n in the curve might not be discovered i f a narrow range of temperature were used and that demonstration of a V-shaped, rather than a simple direct or inverse, r e l a t i o n s h i p between meristic counts and temperatures sometimes requires rearing at extremes close to the upper or lower l e v e l of temperature tolerance of the species. Such V-shaped curves have seldom been shown to exist i n natural f i s h populations.  That they e x i s t i n Cottus asper, a l a t i t u d i n a l l y wide-ranging  species, might be interpreted to indicate that, at the ends of i t s d i s t r i butional range, the p r i c k l y sculpin's eggs are subjected to developmental temperatures close to the upper and lower l e v e l s of tolerance.  However,.we  have elsewhere indicated (p. 53) that the usual spawning range f o r Cottus ..' asper i s from 8 to 13° C,  It i s assumed that i n a l l populations spawning  i s i n i t i a t e d within t h i s 5° range of temperature which proceeds in somewhat of a thermal "wave" from south to north i n both inland and coastal l o c a l i t i e s ( F i g , 5). This i s not to say that they a l l spawn at the same temperature within that range.  Furthermore, because the monthly rate of increase  of ambient temperature i s greater in the north (Fig, 12, and Appendix Table 7), the duration of exposure to any given temperature within the 5° temperature range may be shorter.  Eggs subjected to these conditions would pre-  sumably have a f a s t e r development than eggs which developed under r e l a t i v e l y more thermostable conditions such as occur in the south.  Low meristic  counts are generally (although not invariably) associated with f a s t e r rates of development.  Based on the information given above, northern eggs should  produce offspring with low meristic counts.  But when plotted against l a t i -  tude, the mean pectoral f i n ray counts of inland Cottus asper populations  46 produce a linear curve which i s typically c l i n a l , i.e., with more rays in the high latitudes, fewer in the low latitudes.  The coastal population  means demonstrate a V-shaped curve with low counts in the high and low latitudes, and higher counts in the mid latitudes.  Such results are enig-  matic. Whether a temperature range of only 5° C. i s enough to e l i c i t a Vshaped response has not been experimentally confirmed for Cottus asper, but Lindsey (1962) obtained such a curve in the mean counts of pectoral f i n rays in the stickleback, Gasterosteus aculeatus, over a developmental range of 15 to 20° C. It i s apparent from Fig. 11 that V-shaped curves are characteristic for the median soft f i n ray counts of Cottus asper whereas for the paired pectoral fins, only the coastal populations show a V-shaped curve.  The in-  land means appear to be the only exception, at least in the high latitudes. The existence of such a divergence between coastal and inland means indicates that temperature alone is not the determining factor. Interpretation of the divergent results between coastal and inland means i s facilitated by an analysis of mean pectoral f i n ray counts for coastal lake and coastal derivative populations. In the former, the mean counts do not f a l l , as would be expected, on the curve for coastal populations but they are directly in line with, and indistinguishable from, the higher means of inland populations. In the derivative populations, the means are found to be equal to, or slightly less than, the inland means but always greater than the coastal means at the same latitude (Fig, 5), This further suggests that temperature alone cannot be the determining factor. There i s only one universal factor which, at any given latitude, varies between coastal populations and a l l others. This factor i s salinity and/or the lack of access to i t in the spawning population. At least two of the four coastal lake populations are restricted to spawning in fresh water  47 because of man-made or natural barriers, i.e., Horne L. and Buttle L. on Vancouver Island.  Even though the mouth of Skidegate L. on Moresby Island,  QCI, is only about 10 miles from the sea, the population of Cottus asper from which the meristic data were sampled, is known to spawn in fresh water inlet streams.  The spawning locality of the Owikeno  L., B. C, population  i s unknown. That of the Chilkoot L., Alaska, population is also unknown but the f i n ray counts are typically inland whereas the prickling pattern is definitely coastal in i t s affinity.  It may therefore be supposed that  spawning occurs in fresh water. It may also be presumed that a l l the coastal derivative forms now spawn in fresh water.  Therefore, in the high latitudes, increased-mean  pectoral fin ray counts in coastal lake or "coastal derivative" populations, and in inland populations, is correlated with the lack of salinity in the environment. Any discussion.of the nature of the differences between coastal and a l l other populations is a matter of conjecture with the evidence now at hand. Two opposing hypothesis can be postulated: 1)  The observed differences reflect environmentally induced changes  of the individual phenotypes within each population; or 2)  The differences have become genetically fixed in the populations  over a long period of time. Consideration of these alternatives rests on an interpretation of the relationship of the coastal lake and'coastal derivative" populations to the main coastal population.  In the areal distribution and intensity of their  prickling, the coastal lake and "coastal derivative" forms show very close affinity with the coastal populations.  Furthermore, because of the glacial  history of the coastal islands on which they are found, the near-naked forms from Buttle L., Horne L., and Skidegate L,, can be assumed to have been derived from coastal stream forms which colonized the island after the retreat  H8  of the Cordilleran ice sheet which covered the area during the late Wisconsin Glaciation.  The probability that they were colonized by inland  forms is remote since the nearest inland forms would presumably have been found in unglaciated areas of the lower Columbia R. basin (cf. Fig, 20), In similar manner, the occurrence of the "coastal derivative" populations only in parts of northern British Columbia relatively recently exposed by the retreating glaciers argues that these populations are probably derived from coastal ancestors which followed the retreat of the glaciers northward and then inland.  It can be assumed that they are more recent than  their coastal lake counterparts,.and perhaps even more recent than those i n land forms which followed the retreat of the ice in the Fraser R. drainage (cf. final discussion, p. 7 9 ) ; . As was previously stated, the increased mean pectoral f i n ray counts in coastal lake and "coastal derivative" populations i s correlated with a lack of salinity.  One can explain the phenotypic changes in pectoral f i n  ray counts by postulating an environmental inhibition or alteration of the metabolic pathway usually traversed by the gene which controls the phenotypic expression.  The lack of salinity in the environment might produce  such an alteration. But, since not a l l of the "coastal derivative" means are equal to those of the inland populations, i t can also be argued that,., because of a selective pressure caused by the absence of salinity in the environment, the coastal lake populations have become, and the "coastal derivative" populations are becoming, genotypically modified toward the means characteristic of the inland forms of Cottus asper.  Such modification  might be envisioned in terms of a mutation in one of the steps in the metabolic pathway of the controlling gene. Whatever the cause of the differences, the evidence at hand is not sufficient to determine i t s nature.  now  A definitive experiment regarding the  nature of the differences would consist of taking the eggs of coastal lake  49 or'coastal derivative" forms and rearing them in brackish water using fresh water reared eggs as a control.  If the mean f i n ray counts of the F^ gener-  ation were significantly lower than the control means, the result could be interpreted as being due to.environmental induction rather than to long term genetic fixation.  In either event, the mechanisms involved in the -  actual modification of the phenotype and/or genotype would s t i l l remain a matter for more refined genetic experiment. Summary of Morphological Analysis 1.  Cottus asper i s found in a l l major Pacific Slope drainages from  Seward, Alaska, to Ventura, California.  On the Artie Slope drainage i t s  distribution is limited to the Upper MacKenzie R. drainage only in the headwaters of the Peace R. in British Columbia. 2.  Some of the prickles found in the skin of Cottus asper are re-  sorbed during reproductive maturation which occurs at about 60 mm  S.L.  Because of this, a l l population means were calculated for groups of specimens less than or greater than 60 mm S.L. 3.  Estimates of the percentage of total body surface covered with  prickles are calculated from the formula: % "virtual" prickling = % "apparent" prickling + % area of . 4.  ;  -  resorbed prickling.  Coastal populations of Cottus asper are, "apparently" and  "virtually," less prickled than inland populations. Differences in "apparent" prickling between coastal and inland populations are more pronounced in specimens ^ 60 mm S.L., i.e., a coastal-inland ratio of 1 : 3 exists. 5.  Inland populations of £. asper from the headwaters of the Peace,  Ness, Stikine, and Skeena R. systems in northern British Columbia, are coastal in their prickling affinities and are referred to as "coastal de^r rivative" populations. 6.  A population of C. asper from the mouth of the Sacramento R, in  50 C a l i f o r n i a i s intermediate i n i t s p r i c k l i n g a f f i n i t i e s and i t i s designated as an "intergrade" population. 7.  Asymmetrical p e c t o r a l f i n element counts were found i n 56 of 59  populations of £. asper. .were asymmetrical.  Of 1163 paired counts, from 59 populations, 23.6%  When the asymmetrical  f i n s from a l l populations were  pooled, the o v e r a l l d i r e c t i o n of asymmetry was s i g n i f i c a n t l y s i n i s t r a l ( 6 4 % s i n i s t r a l , 36% dextral). 8.  Differences i n amount of s i n i s t r a l i t y within and among aggregate-  populations are not s i g n i f i c a n t . 9.  No c l i n a l trends are evident.  Mean p e c t o r a l f i n ray counts f o r a l l populations are similar in the  low and mid-latitudes.of the d i s t r i b u t i o n a l range, but are widely divergent between coastal and inland populations i n the high l a t i t u d e s . 10.  Means of "coastal derivative"populations are equal to or l e s s than  those of inland population means but always higher than those of coastal means at the same l a t i t u d e , 11.  Means of coastal lake forms are a f u l l count higher than the means  of coastal stream populations from which they were probably derived.  They  are indistinguishable.from inland means at the same l a t i t u d e . 12.  When plotted against l a t i t u d e , mean f i n ray counts produce curves  which are different f o r inland and coastal populations. 13.  For pectoral f i n s , the, greatest amount of divergence between i n *  land and coastal means e x i s t s i n the high l a t i t u d e s .  The opposite i s true  for the dorsal f i n elements, 14.  Except.for the spiny dorsal and the inland pectoral means, a l l  meristic curves demonstrate a c h a r a c t e r i s t i c V-shaped pattern with the i n f l e c t i o n occurring at about 42 to 44°~N, Latitude, 15.  The p e l v i c f i n almost invariably consists of 1 spiny plus 4 s e g -  mented rays, 16.  Counts of the l a t e r a l l i n e pored-scales are extremely variable  51 with size up t o about 60 mm S.L,  In Cottus asper, a "complete"  lateral  l i n e count cannot be considered r e l i a b l e unless a count i s made of the number of pored-scales plus the number of unenclosed neuromast h i l l o c k s ,  STUDIES ON THE LIFE HISTORY OF THE PRICKLY SCULPIN Present Knowledge of Spawning Migration i n Cottus asper The existence of a seaward spawning migration of p r i c k l y sculpins i n coastal streams has been suggested, or implied, f o r at least 30 years (Taft, 1934; Pritchard, 1936}  Sumner,;1953; Shapovalov and Taft,  Hunter, 1959; and McAllister and Lindsey, 1959).  1954;  Although some of these  authors have observed the presence of reproductively mature Cottus asper in the i n t e r t i d a l areas of coastal streams, none has demonstrated  that  i n t e r t i d a l spawning a c t u a l l y occurs. Populations of Cottus asper occurring i n lakes and streams f a r enough inland to preclude the p o s s i b i l i t y of an annual seaward spawning migration are presumed to spawn i n fresh water  0  The approximate or exact spawning  s i t e s of some of these populations have now  been determined from the pre-  sence of larvae (Nicola L., B, C.), and egg clusters or gravid females (Pothole L. near M e r r i t t , B.C.). 8  Other l o c a l i t i e s are close enough to the  sea to imply the existence of a short seaward migration on the part of the Cottus asper populations l i v i n g therein, but access to the sea i s prevented by natural or manmade b a r r i e r s , e.g.. Buttle L„ and Horne L. on Vancouver Island  s  B.C.  Spawning of p r i c k l y sculpins i n these areas i s necessarily  r e s t r i c t e d to fresh water. S t i l l o t h e r ' l o c a l i t i e s , frequented by migratory salmonids, are close enough to the sea to imply a seaward migration on the part of Cottus asper, but i t does not occur.  For example, the spawning s i t e s of the following,  p r i c k l y sculpin populations i n the lower Fraser Valley i n B r i t i s h  Columbia  52 are known from capture of gravid f i s h and/or egg masses: South Alduette R.; Kenworthy Cr. and Chilqua Slough (both are i n l e t streams to Hatzic L.); Squakum L.; and Cultus L.  In addition, spawning f i s h have been captured i n  i n l e t streams of S k i d e g a t e i n the Queen Charlotte Islands, along with migratory juvenile salmonids. miles from the sea.  The outlet of Skidegate;L. i s only about 13  A newly-hatched larva of Cottus asper (?) has been  taken in a plankton net i n the Second Narrows region of Owikeno L. (about 30 miles from the sea), on the coast of central B r i t i s h Columbia.  It i s  presumed that the parents spawned in fresh water. Spawning Period and Temperature  Relationships. - Egg clusters were  collected from several l o c a l i t i e s in-the lower Eraser v a l l e y and i n the L i t t l e Campbell R. Gonads were examined i n over 1100 of the preserved s p e c i mens from a l l l a t i t u d e s .  These-data indicate that egg deposition begins i n  the south of the d i s t r i b u t i o n range (low l a t i t u d e s ) i n February, and progresses northward u n t i l late July.  Males usually a t t a i n f u l l reproductive  maturity about a month before, and remain in spawning condition f o r almost a month a f t e r , the period of oviposition in females.  Gravid females have  been found over a <+~week period i n Squakum L., and 6 weeks i n the L i t t l e Campbell R„  Ripe males have been taken over an 8 to 12 week period, r e -  spectively, i n these same l o c a l i t i e s . The e a r l i e s t date on which a ripe male, i n n u p t i a l dress and with flowing sperm, was collected i s February 6th, i n San Francisco Bay.  The e a r l i e s t  c o l l e c t i o n of gravid females was in Waddell Cr., C a l i f o r n i a , on February 2<+th, In the north end of the range, gravid females,were taken as l a t e as June 20th in Petersen Cr., near Juneau, Alaska, and on July 22nd i n streams entering Juskatla I n l e t , Queen Charlotte Islands.  Gravid females have also been  collected from Middle R., near Takla L., on June 28th, and from Meziadin L., B.C., on July 25th. F i e l d records and personal observation indicate that natural spawning  53 temperature range from 8 to 13° C, Egg rearing experiments a t various temperatures resulted in complete mortality in 18° C., and less than 50% survival at 15° C, The annual mean range of temperatures experienced by inland populations of Cottus asper i s twice as great as that experienced by coastal populations at the same latitude (Appendix Table 7),  The monthly mean temperature range  between northern and southern localities i s from 8.8 to 19,5° C. on the coast, and from 9.6 to 29.6° C. inland (Appendix Table 7). A p r i o r i , one might expect that inland populations would experience a greater range of developmental temperatures than do coastal populations. ever, they do not.  Apparently, how-  When monthly mean temperatures, representing inland and  coastal localities encompassing the distributional range of Cottus asper« are plotted against latitude, the mean temperature differences between i n land and coastal localities during the spawning period are almost negligible (Fig, 12).  In fact, the common temperature range of 8 to 13° C. (shaded  bar, Fig, 12) can be followed as a thermal "wave" progressing from the south in February, to the north in June,  Approximate spawning times, determined  from examinations of gonadal condition in over a thousand specimens in a l l latitudes, are in general agreement with this south-north progression a l though the latitudinal range over-which spawning occurs in March, April,-and May i s remarkably consistent (Fig. 12). Theoretically, inland forms have a shorter period of exposure to spawning temperature of 8 to 13° C,, thanido coastal forms (Fig, 12),  This has  been born out by f i e l d data from the;two most frequently collected spawning sites, i,e,, Squakum L,, and Little Campbell R. According to Fig. 12, the inception of spawning in inland streams should lag behind that of coastal streams at similar latitudes.  This i s because  upstream or inland areas remain; colder for a longer period than coastal areas.  This i s apparently true in the lower Fraser valley.  For example,  LEGEND; Coastal (• •); Inland (o-—-o); Empirical Spawning Temperature Range ( 8 - l 3 ° C . ) E Latitudinal Range of Probable Spawning in: Coastal 1\ \ \ \ \ l ; Inland K / / / \ \ and Both IX Y Y Y M Forms. State of Maturation: VII = Near Spawning, Vlll= Spawning Imminent, Eggs and Sperm Flowing; IX= Spent. Figure 12.-Monthly/Mean Temperatures i n °C„ Arranged by Latitude f o r Coastal and Inland L o c a l i t i e s Encompassing the D i s t r i b u t i o n a l Range o f Cottus asper. Values obtained from Appendix Table 7. S o l i d or f i l l e d v e r t i c a l b a r s l e f t side o f graph, indicate actual ranges o f l a t i t u d e over which specimens have been found i n given state o f maturity. 8  55 the following localities are a l l within 15' of 49° N. Latitude, and gravid females and/or eggs have been collected from them during the following dates; March 7th to May 10th in Little Campbell R. (122° 46' W. Long.); March 25th to April 15th in S. Alauette R. (122° 35' W. Long.); and April 30th to May 27th in Squakum L. (122° 00' W. Long.). Some of the implications of these temperature and spawning data have a l ready been discussed*(p. 45), >s  Materials and Methods To document the supposed occurrence and to determine the success of  intertidal spawning of prickly sculpins in: the coastal streams, the Little Campbell R. was chosen as a study stream.  In1960-1961, a series of 18  permanent collecting sites (Fig. 14) was sampled at bi-weekly intervals for a period of one year, and at monthly intervals for an additional six months. The lower reaches of the river, stations C-l to C-3, were also sampled  •  several times in late Winter and early Spring of 1962 and 1963, to obtain live specimens for experimental studies, A 3 mm mesh woven nylon seine, 3 meters wide x 2 meters deep, was mounted on collapsible telescoping aluminum poles and used for a l l field collections.  Salinities were measured with three salinity hydrometers,  densities; 1.000 to 1.011; 1.010 to 1,021; and 1,020 to 1.031 sp, gr, of NaCl. Sampling Localities and Study Streams The primary study area was the Little Campbell R, (Campbell Cr.), which is approximately 15 miles long and empties into Semiahmoo Bay between White Rock, British Columbia, and Blaine, Washington (inset of Fig. 13), The stream drainage consists of an area of approximately 28 square miles. Collection-sites are shown in Fig, 14.  Station 0-1 is located on a  sand-mud flat outside the main river channel. Station C-0 is located at the mouth, in the main channel. Station C-l is located about 50 yds, inside the  Figure 1 3 . — C o l l e c t i o n L o c a l i t i e s f o r Spawning Populations of Cottus asper Used i n Life History Studies.  Figure 1 4 . — C o l l e c t i o n L o c a l i t i e s Within L i t t l e Campbell R.  58 mouth and Station C-1A  about 75 yds. inside the mouth.  ed 1/4 mile, and C-3 i s 1/3 mile, from the mouth.  Station C-2 i s l o c a t -  Fig. 15 shows c o l l e c t i o n  s i t e s i n r e l a t i o n to s a l i n i t y gradients and elevation. Stations C-0  through C-9 are a l l subject to t i d a l fluctuations i n  depth, whereas stations C-0 through C-3 are t i d a l l y inundated with mixohaline waters.  Barnacles (Balanus sp.) are found at stations C-0  through  C-3, and permanent beds of the oyster, Crassostrea gigas, are located between Stations C-l and C-1A,  and at C-2,  Typical f i s h associates i n areas  C-0 through C-3 are Leptocottus armatus and Platichthys s t e l l a t u s ; Oligocottus maculosus and Clinocottus acuticeps are commonly found up to Station C-2, Additional l i v e specimens f o r use i n the experimental studies were collected from the following l o c a l i t i e s (Fig, 13)s  N i l e Cr, and Big  Qualicum R,,  Vancouver Island; South Ald;uette R,;  Kenworthy and Edwards Cr,  (Hatzic L o ) ;  Squakum L, (Lake Erroch); and Sweltzer Cr, (outlet of Cultus  L.) Results of F i e l d Studies The p r i c k l y sculpin i s d i s t r i b u t e d p r i m a r i l y i n the lower four miles of the L i t t l e Campbell R, (Fig, 217),  From late February to e a r l y March, the  p r i c k l y sculpin undertakes a migration downstream to the estuary.  The only  area in the lower four miles of stream i n which a suitable spawning substrate (large cobbles, f l a t rocks) occurs i s a stretch about 100 yards long lying one quarter mile upstream from the mouth (Station C-2, Figs, 14 S 15), The males, which come into spawning condition e a r l i e r i n the season than-the females, select nesting s i t e s under large cobbles or f l a t rocks i n areas of the stream bed with current v e l o c i t i e s equal to or l e s s than one cubic foot/ second (at low t i d e ) .  It i s apparently important that the substrate surface  be r e l a t i v e l y rough in texture since the adhesive eggs adhere only tempor a r i l y to a smooth surface.  Old automobile exhaust pipes, or muffler tubes,  A L T I T U D E  cn  O  6S  cn  IN  O  F E E T  cn  60 are preferred nesting sites when available in the environment  (as they are  in the Little Campbell R.), Females aggregate upstream (about Station C-3) above the main spawning area and then move individually onto the spawning beds where they are courted by the males both outside.and inside the nest. After the male selects a female to occupy his nest, further courtship and pre-spawning behavior occurs within the nest.  The adhesive eggs are laid in a j e l l y -  enclosed cluster on the ceiling of the spawning chamber.  Ovariotomy of  preserved gravid females from throughout the range of C. asper yielded counts of 336 mature ova in a 4.9,5 mm S.L. female to 5,652 mature ova in a female 119.5 mm S.L. was spent. female. 4000.  The largest female examined was 192 mm S.L,, but she  A conservative estimate would be about 10,000 eggs for this  Egg numbers, in masses collected in the f i e l d , varied from 700 to However, one male may court and successfully mate with as many as 10  different females.  As many as 10 egg masses, in varying stages of develop-  ment from newly-fertilized to near-hatching, have been found in the nest of a single brooding male.  An estimated 25,000-30,000 eggs were present in  this one nest. After spawning, the spent females leave, or are chased from, the nests and they again aggregate above the spawning areas and begin feeding.  The  males remain in the nests, fanning and protecting the eggs, and do not eat until hatching is completed.  Much of the courtship and pre-spawning be-  havior, as well as most of the paternal brooding behavior, has been documented but w i l l not be reported here. Fig. 16 illustrates numbers and distribution of Cottus asper young-ofthe-year, 12-25 mm S.L., taken in a total of 700 seine hauls.  In late  Spring and throughout the Summer, the young-of-the-year are found in great numbers around and below the spawning site.  The concentration i s greatest  around Station C-1A where there i s a bed of fine, pea-size gravel adjacent  CO 23  III  O  1  £ hCO  O  o o | o IT o I- o 1  9  9  x  o o o o o  o o o o o o  X  o o o o  o o o o o  X  o o o o  o o o o o o  o X  o o o o  o o o o o  X  o o o o o  o o o o o  X  X  X  X  X  X  X  o o  o o X  X X  x  X  o o o  X  X  X  o  o o o o  X  o  X  LU  o  4  s 7 — 6 f" 5 CO 8  o o-4 o 3  o  O O  9  2  CO m I >  o  O  * -J —J LU Op  3 2 IA |, Q, a, 0-1  CO  °2  CD  LEGEND;  X= NO collection made;  0=Collectlon, but no fish; #=1-6 fish; • = 7 - 1 8 ; W = l 9 - 3 6 ; W = 3 7 - 6 0 , l\\\\\M  Intertidal Zone;  W=6l-90  EMnnsnninSpawning Zone; ^ ^ ^ B l I S p a w n i n g Season.  Figure 16„—Monthly D i s t r i b u t i o n of Young-of-the-Year Cottus asper Within the L i t t l e Campbell R Pooled Bi-Weekly Samples,  e  Taken from  £  62 to a large oyster bed. In mid-Summer, there i s a definite upstream migration of the young-ofthe year.  In both 1960 and 1961, the increasing abundance of young-of-the-  year at Stations C-4 and C-5 was.correlated with the decreasing abundance of specimens in the estuarine areas of the river (Fig. 16). Fig. 17 illustrates the disjunct distribution of yearlings, sub-adult, and adult prickly sculpins in the Little Campbell R.  Especially in Spring,  1961, an increased number of Cottus asper were present in the lower reaches of the river, around the spawning site.  Over the 18-month sampling period,  no Cottus asper were captured in stations upstream of C-22,  With three  exceptions, none was taken in the fast flowing, high gradient area of the stream below C-20 and above C - l l (Fig. 15),  This area i s densely populated  with the coastrange sculpin, Cottus aleuticus. Newly metamorphosed young-of-the-year are found in abundance around and especially below the main spawning beds.  Laboratory studies on the behavior  of Cottus asper larvae, done in extension of the.experimental salinity tolerance experiments, indicate that, at 12° C,, the larvae 5 to 7 mm in total length begin swimming immediately upon hatching.  They remain pelagic,  as lightly-pigmented, transparent larvae, for a period of 30 to 35 days before metamorphosing and settling to the bottom. Salinity Tolerance in Prickly Sculpin Eggs Attempts to rear a r t i f i c i a l l y fertilized eggs beyond the c r i t i c a l yolksac resorption stage were unsuccessful. Although a few sculpins were reared to a point where meristic series and prickling patterns were fixed, sample sizes were too small to determine whether the salinity of the fertilization and incubation medium was correlated with f i n ray numbers, or the intensity and distribution of prickles. Alternate c r i t e r i a , based on the percentage hatching success of " i n land" and "coastal" eggs in brackish and fresh water, indicated a phenomenon  CO 23Z  O  r:2r  0  p  0  0  •• • 0  CO , 11 2  O  • • • • X  9  0 0  0  X  0  0  0 0  •  X  0 0  •  0 0  0 0  0 0  X  X  X  X  0  0  X  X  0  0  0  0  0  0  X  0  X  0 0 X  0 0  •  0  • 0  •0 X  0 0 0  X  X  X  • •  •0  •0  0  X  0  X  X  X  X  X  X  X  0  _  X  0  X  X  9 8 71  0  x•  0 0  •-• _  6 5  m  4  #  3  o  LU  CO  m 1  >  O <Q CO  OS CD  <  J  J  A  S  0  I960  N  D  J  F  M  A  M  J  J  A  S  O  1961  LEGEND: X-NO collection made; OCollection, but no fish; •=l-6 fish; •=7-18; W=l9-36, W = 37-60; W = 6 h 9 0 1 \ \ \ \ \ \i Intertidal Zone;  EinnnnsnnniSpawning Zone; M l i j i l i i f f l Spawning Season.  Figure 1 7 . — D i s t r i b u t i o n of Yearling, Sub-Adult, and Adult Cottus asper within the L i t t l e Campbell R,  w  64 of non-genetic adaptation on the part of the "inland" eggs.  The low hatching  success of "inland" eggs which had been f e r t i l i z e d and incubated in brackish water could be s i g n i f i c a n t l y increased by f i r s t f e r t i l i z i n g them in brackish water.  It was also found that acclimation of "inland" parents to brackish  water at least three days p r i o r to f e r t i l i z a t i o n s i g n i f i c a n t l y increased the percentage hatching success of t h e i r eggs over those eggs from parents which had been maintained i n fresh water p r i o r to f e r t i l i z a t i o n .  Data f o r these  egg s a l i n i t y tolerance experiments i s on f i l e i n the Institute of F i s h e r i e s , Department of Zoology, University of B r i t i s h  Columbia.  Discussion This study confirms the existence of a downstream spawning migration of the p r i c k l y sculpins i n the L i t t l e Campbell R.  It not only documents the  occurrence of i n t e r t i d a l spawning in Cottus asper, but indicates that t h i s spawning i s successful. Figure 16 shows a lag of about two months in the appearance of youngof=the~year C, asper, 12 to 25 mm March.  S.L., a f t e r the f i r s t recorded spawning.in  Water temperatures i n March are normally from 8 to 10° C„  Eggs  spawned early i n March probably have an incubation period several days longer than the 15=16 ratory.  day period found to be t y p i c a l at 12° C, i n the labo-  The lag shown i n Figure 16 i s probably due t o an incubation period  of three weeks followed by a pelagic l a r v a l stage of 4 to 5 weeks.  Plankton  tows taken during l a t e A p r i l i n the shallow back-eddies of the stream have captured a few larvae 9 to 10 mm S.L,  (not recorded in F i g , 16),  An upstream migration of adults precedes that of the young=of-the-year in late Summer,  This i s probably related to the food habits of the two  groups and also to the fact that a l a t e r return of the  young-of-the-year  coincides with lower water l e v e l s i n the stream, when reduced v e l o c i t y f a c i l i t a t e s easier access upstream.  65 Summary of Life History Studies 1.  The study stream, the Little Campbell R., i s a small coastal  stream, the lower one half mile of which i s subject to t i d a l inundation of mixohaline waters. 2.  In late Winter and early Spring, the adults and juvenile prickly  sculpins migrate downstream to the estuarine region of the Little Campbell R., the only region in the lower four miles in which suitable spawning substrate is available« 3.  Males set up nesting sites under large cobbles and rocks and  courtship occurs both outside and within the nest* 4. Spawning occurs from March throughout early May. 5.  Newly-hatched larvae begin swimming immediately and remain pelagic  for a period of 30 to 35 days before metamorphosing and settling to the bottom. 6. In May, metamorphosed young-of-the-year (approximately 12 mm S.L.) begin appearing only in those collections taken in the estuarine portion of the river.  They occur in great abundance until September when the numbers  decrease in the estuary and increase in upstream, non-estuarine  waters.  7. During the non-migratory phase of i t s l i f e history, the Cottus asper population in the Little Campbell R. i s distributed disjunctly in the low gradient, low velocity, portions of the stream.  DISCUSSION The purpose of this investigation was to determine which characters in Cottus asper were genetically determined and which were due to nongenetic modification.  Properly answered, these questions could help the  systematist to a better understanding of the selective forces that produce variations within a species.  This knowledge would, in turn, facilitate a  66 more stable classification of taxa. Morphological Variation Within Cottus asper The interpretation of the morphological variation will be considered in the light of experimental and l i f e history data obtained in this investigation.  Table V summarizes several of the variable characteristics for  aggregate populations of Cottus asper. Coastal vs. Inland Populations. - The data presented in Tables I and V and in Figure 4, reveal consistent differences in prickling patterns 8  which coincide exactly with the geographic locality of collections.  Coastal  fish have less area of the body covered by prickles (ratio 1 t 1.8), and less prickles per unit area (ratio 1 : 1,4) than inland fish.  The existence  and maintenance of such a remarkable discontinuity of characters between closely related but geographically distinct populations, distributed over a wide range of latitudes, must be interpreted as being genetically determined.  Furthermore, these populations, which can be separated on the basis  of prickling patterns and geographic distribution, can also be separated by other characters. For example, consider the migratory behavior. Weakly-prickled Cottus asper living in coastal streams having open access to' the sea undertake a downstream migration to the estuarine regions where they spawn. Densely-prickled Cottus asper l i v i n g in distant inland ;, streams, where access to the sea i s almost impossible, undertake only local migratory movements. But densely-prickled forms living in some inland lakes and streams relatively close to the sea, where access to the sea i s open and relatively easy, do not migrate seaward.  They also undertake only local  movements and spawn within the fresh,water system in which they are found. These primary differences in behavior patterns correlated with distinct prickling patterns further corroborate the contention that coastal and i n land forms of Cottus asper are genetically distinct. The evidence based on meristic counts i s equivocal. When plotted  TABLE V»  Summary of Variable Characteristics for-Selected Phenotypes of Cottus asper,,  Phenotype Descriptions  Distribut ion s  COASTAL  Streams of Pacific Coast from Alaska to California  Northern COASTAL DERIVATIVE  River drainages north of Fraser R, drainage in British Columbia  Northern COASTAL LAKE  Southern iNTERGRADE  Island and mainland lakes close to sea i n British Columbia and Alaska  Mouth of Sac= ramento R» in San Pablo and San Francisco Bays  INLAND  A l l major inland drainage sys= terns on Pacific Slope except drainages north of Fraser River.,  Prickling Virtual Areas  Low  Low  Lowest  "-.Higher  Highest  Intensity!  Low  Low  Lowest  Higher  Highest  % Resorptions  High  Lower  Lower  High  Lowest  Pectoral Ray Numbers @ High Latitude  Low  Higher  Highest  Combined Dorsals at Low Latitudes  Highest  Meristics Highest  Higher  Low  TABLE V (cont'd).  Summary of Variable Characteristics for Selected Phenotypes of Cottus asper.  Phenotype Description:  COASTAL  Migratory Behavior:  Seaward Migration  Natural Spawning Environment:  Brackish  Egg Survival in other than natural media:  Excellent; as in natural medium  Northern COASTAL DERIVATIVE  ?  Presumably Fresh  ?  Northern COASTAL LAKE  Southern INTERGRADE  ?  Presumably Fresh  ?  ?  Presumably Brackish  ?  INLAND  Local Migratory movement  Fresh  Poor; s i g n i f i cantly less except i f parents or eggs preacclimated  69  against latitude, the means for inland and coastal populations form V-shaped curves which, though overlapping in certain areas, are widely divergent in others (Fig. 11).  For example, the pectoral rays of inland and coastal  populations are similar in the low latitudes but at a point about 42-44° N, Lat., the coastal means begin to decrease, whereas the inland means continue to increase, in f i n ray numbers.  Such divergence in mean counts be-  tween coastal and inland populations can not be explained on the basis of temperature alone.  Salinity in the spawning environment i s the only factor  which, at any given latitude, can be shown to vary directly between inland and coastal populations. The nature of the differences between coastal forms and a l l the populations probably derived from them, or between coastal and inland forms, i s a matter of conjecture.  Phenotypic modifications now shown to exist can, on  the basis of present evidence, be interpreted either as being individual changes induced by the local environment, or, as having become genetically fixed in the populations over long periods of time.  Theoretical arguments  for each interpretation have been presented ( p, 48 )» but the f i n a l determination of the underlying nature of existing differences in f i n ray counts awaits the test of successful rearing, under controlled conditions, to a stage where f i n ray numbers are fixed. Derivative and Intergrade Populations, - Certain sub-groups of prickly sculpins have been found which are morphologically distinctive but which have close affinity to either the coastal or inland forms.  An analysis of  these populations corroborates the contention that, at least in prickling pattern and migratory behavior, coastal and inland forms of Cottus asper are genetically discrete. Even though the coastal lake and "coastal derivative" populations have prickling patterns similar or identical to coastal forms, the absolute percentage resorption is more like that of the inland forms (Tables I and V),  70 This might be expected since presumably they no longer spawn in brackish water and the osmotic stresses experienced while spawning in fresh water must be different. The prickling pattern of the southern "intergrade" population (Carquinez Str.,Calif.) i s similar to that of the densely prickled inland population forms from which i t may have been derived. The amount of prickling resorption, however, approaches that of the coastal percentage.  If we assume, as  we have above, that the percentage resorption i s greater in brackish water this would indicate that they spawn in estuarine conditions.  In their mean  f i n ray counts, other populations from the San Pablo-San Francisco Bay area are intermediate between the coastal and inland populations. With the i n formation now available, i t would be d i f f i c u l t to determine whether these are true inland derivatives which have colonized the streams of the Bay region or whether they are intergrades between inland and coastal forms. , :  The Phylogeny of Cottus asper and Closely-Related Species The prickly sculpin, Cottus asper, i s a polytypic species. Figure 18 shows the known range of various phenotypes of Cottus asper, determined in this study, plotted against the distributional ranges of closely-related species which comprise the "asper species group" proposed by Bailey and Bond (1963).  From a taxonomic viewpoint several interpretations of this polytypy  can be made: 1)  Cottus asper i s a phenotypically variable species within which can be recognized 4 or 5 races; or  2)  Cottus asper represents a species complex comprised of at least two sibling species, each with one or more derivative subspecies (or races i f the phenotypic differences can be shown to be environmentally induced).  3)  Cottus asper represents a polytypic species complex in which a spectrum of evolutionary grades, from incipient subspecies through  gure 18c— Known Ranges of Various Phenotypes of Cottus asper Plotted Against the D i s t r i b u t i o n a l Ranges Closely-Related Species i n the "asper species group".  72 f u l l species, can be traced. Evidence presented i n t h i s study favors the l a t t e r interpretation. The genus Cottus i s a fresh water derivative of the predominantly marine family Cottidae.  The genus Cottus i s most closely related to the  A s i a t i c marine genus Trachidermus, represented on the P a c i f i c Coast of N, America by Leptocottus, a c l o s e l y - a l l i e d genus.  Although t h e i r i n t e r p r e t a -  t i o n s of relationships within the genus Cottus were not always accurate, I concur with the e a r l i e r opinion of Jordan and Evermann (1898, p. 1942) that, of the species of Cottus occupying the P a c i f i c Slope, Cottus asper appears to be more closely a l l i e d to Trachidermus. It has been concluded that differences i n p r i c k l i n g patterns and migratory behavior between the geographically separated coastal and inland populations of Cottus asper are genetic (p.66 ).  The discontinuity i n such  conservative characteristics indicates that the i s o l a t i o n of the inland populations from the coastal gene pool has been of long-standing, perhaps since the Pliocene or late Miocene.  Evidence to support t h i s speculations i s  based on the geologic history of the area now occupied by Cottus asper and closely-related species i n the "asper,!species group" discussed below. Stage I. - I t must be assumed that at sometime in the distant past, i.e.,  perhaps Pliocene or e a r l i e r , the ancestral form of Cottus asper was  derived from a marine ancestor similar to Trachidermus.  After this i n i t i a l  separation, some of the ancestral Cottus asper penetrated inland and, through geological agencies and/or b i o l o g i c a l processes, became isolated from the coastal gene pool.  Such an i s o l a t i o n i s represented schematically as Stage I  in the evolution of Cottus asper ( F i g . 19).  The dashed arrows indicate  probable routes of invasion of coastal types inland v i a e i t h e r the R. or the Sacramento R. system.  Columbia  During the Pleistocene, access to the north  was blocked by the Cordilleran ice sheet, lobes of which advanced and r e treated several times.  The l i m i t s ^hown in F i g , 18 indicates the maximum  73  Figure 19,~Phylogeny of Cottus asper. Stage I, Evolution in the Pliocene (?)„ Similar pattern of background indicates continuous range,  7 4  southward extent of ice cover during the late Pleistocene; e a r l i e r advances were somewhat further southward, Zoogeographically, the disjunct populations of Cottus asper and  "asper  species group" represented i n F i g , 18 are members of one of three great P a c i f i c Slope fresh water ichthyofaunae, i . e , , the Columbia; the Klamath; and the Sacramento, the basins  B  It i s here proposed that at some time or times before  e s p e c i a l l y the Klamath, became completely i s o l a t e d from one  another by alternate periods of floods and d e s i c c a t i o n , lava flows, and mountain b u i l d i n g , a continuous exchange of fauna was possible.  The geo-  l o g i c a l evidence of contemporaneous connections between the (Pleistocene) p l u v i a l lake basins i s wanting, but the ichthyological evidence c a l l s f o r such a connection some time in the past (Hubbs and M i l l e r , 1948),  For ex-  ample, the cyprinid genus Ptychocheilus i s absent from the Klamath system but closely related species are present in the Columbia system to the north, and i n the Sacramento system to the south.  The sucker, Catostomus snyderi,  and l o c a l representatives of the dace Rhinichthys occurring in the Klamath basin may  be related to the forms in the Columbia and the Sacramento (Hubbs  and M i l l e r , 1948), Stage I I . - The fluctuating regimen of the p l u v i a l lakes i n the Great Basin has been correlated approximately with the advances and retreats of the Cordilleran g l a c i e r s ( F l i n t , 1957;  Hubbs and M i l l e r , 1948).  It i s here  suggested that during t h i s period of extensive flooding and desiccation, the gene pool of the previously continuous inland population of Cottus asper divided up into a series of  was  geographically i s o l a t e d populations, some i n  the Klamath basin, some in the upper Sacramento, and some in the Deschutes R. drainage of the Columbia.  To-the south of Klamath, Cottus gulosus  evolved as a f a s t water, upstream derivative of inland Cottus asper and spread southward and out to the P a c i f i c Coast,  Among other d i f f e r e n c e s , the  fecundity of gulosus i s intermediate between the high egg complement of  75 asper and the low complements of non-related species of Cottus (Bond, 1963).  Perhaps one of the most intensive selective factors impinging on  the i n c i p i e n t gulosus was that favoring a large egg which would produce a non-pelagic larva i n the f a s t water environment (Cottus asper has pelagic larvae). Cottus p i t e n s i s , a disjunct r e l i c t form from the upper Sacramento R. above Lake Shasta, and from the upper P i t R. and Goose L, drainage, i s thought to be derived from gulosus (Bailey and Bond, 1963j Bond, 1963). To the north of Klamath, Cottus perplexus was probably derived from inland asper as the northern ecotypic counterpart of gulosus, in so f a r as i t also prefers upstream areas and has an intermediate egg complement.  By  an unknown connection, perhaps through the outlet of P l u v i a l Fort Rock L. which contains f o s s i l s of possible faunal relationship with the Klamath Lakes, Lake Lahontan, Lake Bonneville, and the Columbia River (Hubbs and M i l l e r , 1948), perplexus worked i t s way into the Deschutes R, and down to the Columbia R. where i t spread both upstream and downstream (Fig, 20), Cottus marginatus  i s probably a r e l i c t form of a formerly more widespread  d i s t r i b u t i o n of perplexus (Bond, 1963). E c o l o g i c a l l y variable, and closely related to perplexus, i s Cottus klamathensis, a disjunct r e l i c t found in Klamath L,, Oregon and P i t R, C a l i f o r n i a ( G i l b e r t , 1898} Bailey and Bond, 1963}  Bailey and Dimick, 1949}  and Bond, 1963).  Robins and M i l l e r ,  1957;  A Pleistocene connection between  the Klamath, Goose L,, and Pit R, basins has been suggested by Hubbs and M i l l e r (1948), and Robins and M i l l e r (1957).  The fecundity of klamathensis  i s as high as i n asper, and other characters suggest a close relationship with, and an early derivation from, the ancestral asper or perplexus.  For  example, Gilbert (1898, p, 11) commented on the resorption of body p r i c k l e s occurring i n klamathensis;  "These, [the p r i c k l e s of the young] become grad-  u a l l y absorbed with age, adults being nearly or quite naked,"  This i s the  76 f i r s t recorded observation on prickling resorption and i t had been completely overlooked by later authors. The suggestion of Robins and Miller (1957) that Cottus princeps i s a long-standing derivative of asper i s probably correct.  The relationship is  probably closest to the ancestral inland asper. Meristic evidence that the "asper species group" could have been derived from an ancestral inland form of Cottus asper i s presented in Table VI. With few exceptions, the ranges of counts for a l l f i n rays  for a l l of the  species f a l l within the range of counts typical of the present populations of inland Cottus asper.  Such a<correlation may be interpreted to indicate  that the genotype of the inland asper has a wide reaction range. A schematic representing the proposed second stage of the phylogeny of Cottus asper is presented in Fig. 20. Additional consideration of Cottus perplexus and C, gulosus w i l l now be given.  In i t s coastal distribution, Cottus perplexus i s widespread in  drainages of Washington and Oregon but does not cross the California border (Fig. 18).  On the other hand, Cottus gulosus i s found somewhat north and  south of San Francisco, but only disjunctly in northern California and parts of the Oregon coast.  When they occur*in the same stream, perplexus i s  generally, though not always, found in faster water, whereas gulosus occurs somewhat downstream in slightly slower velocity waters (Bond, 1963). Bond (personal communication) has stated that in areas where asper, gulosus, and perplexus occur in the same stream* a very thin slice of allopatry i s cut! However, where upstream migration to the upper portion of streams has been blocked by impassable f a l l s , perplexus i s often found above the f a l l s whereas gulosus is always found below (Bond, 1963).  This indicates, perhaps,  that in streams where both species are present, gulosus is the most recent arrival.  Both species have been found to be salt tolerant of at least  2H °/oo S. (Bond, 1963).  Cottus asper i s tolerant of long periods of  TABLE VI.  Summary of M e r i s t i c Data f o r Cottus asper and Other Species i n the "asper species group,,"  Species  D 1  D 2  D +D 1 2  asper (coastal)  8-10 (11)  19-22 (23)  (26) 27-32  A  P 1  P 2  \  n  marginatus perplexus klamathansis  15-18  2  4  2 13-16 2  2  2  18-20  13-15  asper (inland)  7-10  (16) 18-22 (23)  princeps  6-8  19-23  qulosus  5, 6 7- 9  3 16- 19 17- 19  2  26-32  2  15-18 (20)  26-30  2  (6) 8- 9 1 d o )  1  (16) 17-18 ( 1 9 )  15-18  2  14-16 13-15 12-14 (±2) 13-15  3 6  6  pitensis  3 2  18-20  (6) 7-8  (3) 4  14-16  17-19 7-8  14-18  .  Superscript Notation; 1) Bailey and Bond, 1963; 2) Robins and-Miller, 1957; 5) Snyder, 1905; 6) Snyder, 1913. A l l data on asper from t h i s study.  14-16 (13) 14-19 (19) 14-16  2  1 (14) 15-16 (17) 14-16 1 (12) 13-15 (16)  2 4 2 4 (3) 4 2 4  6  5  1  1 (13) 14-15 (16) 14-16  1  3) Bond, 1961;  (3) 4  1  4 ) Schultz, 1936;  78  Figure 20c=-Phylogeny of Cottus asper, Stage I I , Evolution of "asper species group" During the Pleistocene,  79 immersion in higher salinities, i.e., greater than 31 °/oo S. (personal observation; and Bond, 1963).  The salt tolerance of other "asper complex"  species i s unknown. Stage III. - By late Pleistocene, most of the speciation within the inland Cottus asper complex probably had occurred. But another surge of evolutionary activity was yet to* occur, primarily within the coastal population (Fig. 21). With the f i n a l  retreat of the Puget and Juan de Fuca lobes of the  Cordilleran ice sheet (late Vashon Stade) about 13,500 years before present (Armstrong, et a l , 1965), coastal Cottus asper began to move northward along the newly exposed coastline of British Columbia.  With the more or  less contemporaneous retreat of the Okanagan and Spokane lobes from the Columbia R. basin, inland Cottus asper from the lower Columbia advanced northward into Canada and, perhaps at that time, colonized the interior Fraser R. system through meltwater channels then in existence. Coastal Cottus asper now exists in three phenotypic forms in areas of the Pacific Slope formerly covered by the Cordilleran ice sheet: 1)  The typically prickled*.estuarine spawning , coastal form with lower pectoral f i n ray counts;  2)  a typically prickled, fresh water spawning derivative form with high pectoral f i n ray counts, found far inland in the Stikine, Nass, Peace, and parts of the Skeena R. drainages; and  3)  a very weakly-prickled, fresh water spawning derivative form with high pectoral f i n ray counts* found in certain mainland and i s land lakes.  These "coastal derivative" forms probably followed the recent retreat of glacial lobes up narrow valleys in northern British Columbia.  These  forms are necessarily the most recent invaders, arriving, perhaps, even later than the upper Fraser R. forms of the inland Cottus asper from which  80  Figure 21„—Phylogeny of Cottus asper. Stage IIIo Evolution of Coastal Form Following Retreat of Cordilleran Glaciers During Recent Time,  81 they are d i s t i n c t . Lindsey (1956) presented d i s t r i b u t i o n a l evidence that the following primary fresh water species:  Catostomus macrocheilus; Mylocheilus caurinum;  Ptychocheilus oregonense; and Richardsonius balteatus; plus Cottus asper, a euryhaline species, had penetrated into the Peace R. system from the Skeena or Fraser R. system.  " A l l are present in the Columbia, Fraser, and Skeena  systems, a l l are absent from the Yukon, and a l l have been taken i n the Mackenzie system only from the upper portion of the Peace drainage."  The  r e s t r i c t e d range of these species i n the Peace R. system indicates t h e i r r e l a t i v e l y recent a r r i v a l . The present study indicates that the Peace R, representatives of Cottus asper are coastal derivatives which must have penetrated inland v i a the Stikine, Nass, or upper Skeena R.  The inland form of Cottus asper has  apparently f a i l e d to penetrate northward beyond the l i m i t s of the upper Fraser R, drainage.  The exchange probably occurred and the connection be-  came closed before the inland Cottus asper arrived on the scene.  Evidence  from the d i s t r i b u t i o n of other primary freshwater f i s h e s bears t h i s out. Several species got only as f a r north as the inland asper.  For example, the  leopard dace, Rhinichthys f a l c a t u s , i s a Columbia R, form which has pene-. trated northward (presumably  following the same routes as inland asper) as  f a r as Middle River near Takla L. ,• about 120 miles northwest of Prince George, B.C.  The bridgelip sucker, Catostomus columbianus,  another Columbia  R. form, has been taken i n the Salmon-R, system just north of Prince George. Extensive c o l l e c t i n g in the areas north of Prince George indicates that both species have f a i l e d to penetrate into the Peace or the Skeena R, systems. The p r i c k l y sculpin, Cottus asper, i s interpreted as being a polytypic species which has become, and i s in the process of becoming, modified into several c h a r a c t e r i s t i c genotypes.  The dichotomy between "inland" and  "coastal" forms i s b a s i c a l l y ? subspecific in nature.  C l a s s i f i c a t i o n of the  82  "asper species group" i s beyond the limit of a system of binomial nomenclature « Modifications in some of the earlier derivatives of the inland form have presumably become genetically fixed, and various authorities have assigned them f u l l specific rank.  More recently derived forms of the  coastal Cottus asper , such as occur in the coastal lakes of British Columbia and Alaska, or in the British Columbia river drainage systems north of the Fraser R,, are modified sufficiently to perhaps warrant classification as subspecies, but the genetic nature of the modification has not been determined.  It  is n o t known Whether the marked differences i n phenotype o f the  "coastal derivative" or*coastal lake forms are results of a phenomenon which occurs in one generation, o r over a long period of time.  If the changes are  not environmentally induced, then genetic fixation can be assumed to have occurred within the past 10,000 years or less.  83  LITERATURE CITED  Armstrong, E„, D„ R, Crandell, D, J , Easterbrook, and J , B, Noble, 1965, Late Pleistocene stratigraphy and chronology i n southwestern B r i t i s h Columbia and northwestern Washington, Geol, Soc, Amer, B u l l , 76:321-330, Bailey, R. M., and M. F, Dimick, 1949, Cottus hubbsi, a new c o t t i d f i s h from the Columbia River system i n Washington and Idaho, Occ. Pap, Mus, Zool, Univ. Mich. 513s1=18, Bailey, R. M., and C. E, Bond, 1963, Four new species of freshwater sculpins, genus Cottus, from western North America, Occ, Pap, Mus, Zool, . Univ, Mich, 634:1-27, Battle, H. I. 1929. E f f e c t s of extreme temperatures and s a l i n i t i e s on the develppment of Enchelopus cimbrius (L„). Contr, Canad. B i o l , and Fish. N. S, 6:107-192. Bean, B. A,, and A. C. Weed. 1920. Notes on a c o l l e c t i o n of fishes from Vancouver Island, B, C. Trans. Roy. Soc. Can, (3) 13(1919):69-83, Bert i n , L. 1925. Recherches bionomiques, biometriques et syste'matiques sur l e s Epinoches. Ann. Inst. Oce'anograph, Monaco, N, S, 2, fasc, 1. Bond, C. E. 1961. Keys t o Oregon freshwater fishes, Oregon St. Univ., Tech. B u l l , 58:1-42,  Agr, Exp. Sta,,  —,„ 1963. Distribution and ecology o f freshwater sculpins, genus Cottus in Oregon. Ph.D. t h e s i s , Univ. Mich. 1-186, C a r l , C, G,, W. A. Clemens, and C. C. Lindsey. 1959, The fresh-water fishes of B r i t i s h Columbia, B r i t . Col. Prov, Mus, Handb, 5(3rd Ed. Rev,) sl-192. Cooper, J . G, 1868, Fishes, In: The Natural Wealth of C a l i f o r n i a , (Cooper, Jo G„, and T, F. Cronise). H. H. Bancroft and Co., San Francisco, 1868 (Chap. 7):487-498, Crawford, D. R, 1927, Extension of the range o f Cottus aleuticus. 1927(1960):177-178,  Copeia  D i l l , W, A. 1946. A preliminary report on the fishery of Millerton Lake, C a l i f o r n i a . C a l i f . Fish and Game 32(2):49-70, Dymond, J . R. 1936. Some fresh-water fishes of B r i t i s h Columbia, Rep, B. C. Comm. Fish, (for 1935):60-73. Eigenmann, C, H. 1895. Results of explorations i n western Canada and northwestern United States. Bui. U. S. Fish Comm. 14(for 1894:101-132, Evermann, B, W,, and S. E, Meek. 1898, A report upon salmon investigations in the Columbia River Basin and elsewhere on the P a c i f i c Coast in 1896, B u l l . U. S. F i s h Comm. 17(for 1897):15-84.  84 — a n d Eo L Goldsborough,, Bur, Fish, (1906)s219»360o 0  1907a,  The f i s h e s of Alaska, B u l l , U, S  0  — a n d ~=—„ 1907b„ A check l i s t o f the freshwater fishes o f Canada,) Proc, B i o l , Soc, Wash, 20s89~120, and H, B, Latimer, 1910, On a c o l l e c t i o n of f i s h e s from the Olympic Peninsula, together with notes on other West Coast species, P r o c B i o l . Soc. Wash, 23sl31-140, 8  =  and H, W, Clark. 1931, A d i s t r i b u t i o n a l l i s t of the species of freshwater fishes known t o occur i n C a l i f o r n i a . C a l i f , Div, Fish and Game, Fish B u l l , 35 81-67, 1  F l i n t , Ro F„ 1957. G l a c i a l geology and the Pleistocene epoch, and Sons, Inc,, New York, 195781=589,  John Wiley  g a b r i e l , M. L, 1944. Factors a f f e c t i n g the number and form of vertebrae i n Fundulus h e t e r o c l i t u s . J. Exp, Zool, 95 s105-147 0  G i l b e r t , C, H,, and B, W, Evermann, 1895, A report upon investigations i n the Columbia River Basin, with descriptions of four new species of fishes. B u l l , U, S„ Fish Comm. 14(for 1894)s169-207, — — o 1896, The ichthyological c o l l e c t i o n s o f the U, S, Fish Commission Steamer Albatross during the years 1890 and 1891, Rep, U, S. Comm. Fish and Fish, (for 1893), Appendix 6s393=476. 1898, Fishes of the Klamath Basin, 1897)sl-13. 0  B u l l , U„ S, Fish Comm. 17(for  Girard, C. 1850, A monograph of the freshwater Cottus of North Proc, Amer, Assoc, Adv, S c i . 2$409=411, —•—, 1851a, On the genus Cottus Auct, 3(1848-1851)$183-190, —  America,  Proc. Boston Soc, Nat. Hist,  . 1851b. Some a d d i t i o n a l observations on the nomenclature and c l a s s i f i c i a t i o n of the fishes of the genus Cottus. Proc. Boston,Soc, Nat, Hist, 3(1848-1851)8302=305, .. 1851c, Revision du genre Cottus des Auteurs, Sci. Nat, 12(1851)s185 (not seen).  Nouv, Mem, Soc, Helvet,  — - o 1852, Contributions to the natural h i s t o r y of the freshwater f i s h e s of North America, I ^ A monograph of the c o t t o i d s Smiths, Contr. Knowledge 3(art, 3)sl=88. 0  —  1856a, Descriptions of new f i s h e s , collected by Dr, A, L, Heermann, n a t u r a l i s t attached to the Survey of the P a c i f i c Railroad Route, under Lieut, R, S o Williamson, U.S.A. Proc, Acad, Nat. S c i , P h i l a . 7(18541855)s129-141,  0  —--„ 1856b. Observations upon a c o l l e c t i o n of f i s h e s made on the P a c i f i c Coast df the United States, by Lieut, W. P. Trowbridge, U,S,A,, f o r the Smithsonian I n s t i t u t i o n , Proc, Acad. Nat. S c i . Phila, 7(1854-1855)sl42-156.  85 —,-=.„ 1857 Report upon f i s h e s c o l l e c t e d on the Survey, 6(Part IV, No, l ) s 9=34, Ins House of Representatives, 33rd Congo, 2nd Sess „ Ex, Dod No 91o Reports of explorations and surveys f o r a r a i l r o a d route from the M i s s i s s i p p i River to the P a c i f i c Ocean V o l 6s Report of Lieut, Henry L Abbot upon explorations f o r a r a i l r o a d route from the Sacramento Valley to the Columbia River, made by L i e u t o R, So Williamson, assisted by Lieuto Henry L„ Abbot* 1851=1855 Wash Do C„ 0  0  0  0  0  0  0  0  _—... 1859 Fishes, Ins General report upon, the zoology of the several P a c i f i c r a i l r o a d routes U S Pac R Ro E x p l o and Surv 10(part 4)s 1=400, Also found as Part IV, 33rd Congo, 2nd Sess,, Senate Ex, Doc, 78. 0  0  0  0  0  0  0  GSnther, A C L G I860, Catalogue of the Acanthopterygian Fishes i n the c o l l e c t i o n of the B r i t i s h Museum, 2sl=548, 0  0  0  0  Heckel, J o J o 1840„ Ichthyologische Beitrage zu den Familien der Cottoiden, Scorpaenoiden, Gobioiden und Cyprinoiden Ann Wien, Mus, Naturgeshichte 0  0  2(1840)5143-161.  Heuts, Mo J o 1947a, The phenotypic v a r i a b i l i t y of Gasterosteus aculeatus Lo populations in Belguim, Meded, Kon. Vlaam, Akad, Wetenschap, 9(25)s 5=63, 1947b, Experimental studies on the adaptive evolution i n Gasterosteus aculeatus L. Evolution 1(1,2).89-102. ..... i949 aculeatus. 0  0  Racial divergence i n f i n ray v a r i a t i o n patterns i n Gasterosteus J , Genet, 49(3)sl83=191,  Hubbs, C, L, 1921 1929(9)s7-8,  0  Notes on Cottus asper and Cottus aleuticus, " ™ ~ _  Copeia  —--, 1926, The s t r u c t u r a l consequences of modifications of the developmental rate i n f i s h e s , considered i n reference t o certain problems of evolution, Amer. N a t u r a l i s t 60s57=81, —... 1943, C r i t e r i a for subspecies, species, and generja»as determined researches on f i s h e s , Ann, N, Y„ Acad, S c i , *44(art. 2)sl09=121, 0  by  ?  and L, C, Hubbs, 1945, B i l a t e r a l asymmetry and b i l a t e r a l v a r i a t i o n in f i s h e s . Pap, Mich, Acad, S c i . 30s229-310,  ...—=.j  and Ro Ro M i l l e r , 1948. The Great Basin, I I , The zoological e v i dence. Correlation between f i s h d i s t r i b u t i o n and hydrographic history i n the desert basins of western United States, B u l l , Univ.. Utah 38(20) [ B i o l , Ser, 10(7)]S17-166. - - - — , and Oo L, W a l l i s 1948 The native f i s h fauna of Yosemite National Park and i t s preservation. Yosemite Nature Notes 27(12)s131-144, 0  a  - — a n d C, Hubbs, 1953, An improved graphical analysis and of series of samples. Systo Zool, 2(2)s49=57,  comparison  — — , and K. F, Lagler, 1958» Fishes of the Great Lakes Region, Cranbrook Inst, S c i , 26 51=213,  Bull,  86 o — — 1 9 6 3 Chaetodon aya and related deep=living butterflyfishess t h e i r v a r i a t i o n , d i s t r i b u t i o n and synonymy,, B u l l , Mar, S c i , Gulf Caribb, 13(1):133-192. 0  Hubbs, C o , and L, Stavenhagen 1958, E f f e c t s of maternal carotenoid def i c i e n c y on the v i a b i l i t y of darter (Osteichthyes) offspring, Physiol, Zool, 31(4),280=283, 0  Hunter, J, G, 1959, Survival and production o f pink and chum salmon in a coastal stream, J . Fish, Res, Bd, Can. 16(6):835-886, Jordan, D. S. 1877. Contributions to North American ichthyology, No, 2 B u l l . U. S. Nat, Mus. 10s1-120. —  , and P. L. Jouy. 1882. Check l i s t o f duplicate fishes from the P a c i f i c Coast of North America, distributed by the Smithsonian I n s t i t u t i o n in behalf of the United States National Museum. Proc. U. S. Nat. Mus. 4(1881)sl-18.  — — a n d C. H, G i l b e r t . 1883, Synopsis of the fishes of North America, B u l l . U. S. Nat. Mus. 16(1882)sl=l018. _1885 A catalogue of the fishes known to inhabit the waters of North America, north of the Tropic of Cancer, with notes on the species discovered in 1883 and 1884, In_: Rep, Comm, Fish and Fish, ( f o r 1884); 1-185. Also appeared ass App, E, Rep. U. S, Comm. Fish, 1885, (1887; 789-973. 0  0  ——>. 1895. Notes on the fresh-water species of San Luis Obispo County, C a l i f o r n i a . B u l l , U. S„ Fish Comm. 14(for 1894)sl41=142, — — , and B„ W. Evermann, 1896, A check-list o f the fishes and f i s h - l i k e vertebrates of North and Middle America. Rep. Comm,, U, S, Comm, Fish and Fish, ( f o r 1895), App, 5s207-584, — . ^ and B. W. Evermann, 1898. The fishes of North and Middle America. Part I I . B u l l . U. S. Nat. Mus. 47:1241-2183, — —  1919, The^genera of f i s h e s . Part I I . From Agassis to Bleeker, 18331858, twenty-six years, with the accepted type of each. Leland Stanford Jr. Uniy, Publ., Univ. Ser. 1919:1-284, 0  — B . W. Evermann, and H, W. Clark, 1930. Check l i s t of the fishes and f i s h l i k e vertebrates of North and Middle America north of the northern boundary of Venezuela and Colombia, Rep, U, S, Comm, Fish, ( f o r 1928), part 2:1=670. Kermode, F, 1909, V i s i t o r s guide t o the natural history and ethnological collections i n the P r o v i n c i a l Museum, Prov, Mus, Nat, Hist, and Ethn., V i c t o r i a , B. C. 1909:1-92. —.—, 1917. Fishes. In_: Rep. Prov, Mus. Nat. Hist,, Prov, B, C, (1916): 18-22. — — . 1931. Accessions: Ichthyology. In_: Rep. Prov, Mus, Nat. H i s t , , Prov. B. C. (1930):17-23„  87 Kinne, 0 1962 Irreversible nongenetic adaptation, Physiol, 5,265=282, 0  0  Comp Biochem, 0  Lagler, K, F , and R, M, Bailey, 1947, The genetic f i x i t y of d i f f e r e n t i a l characters i n subspecies of the percid f i s h , Boleosoma nigrum, Copeia 1947(1).50=59, 0  Lindsey, C, C, 1954, Temperature-controlled paradise f i s h Macropodus opercularis ( L ) 0  0  meristic variation in the Canada J , Zool, 32 287=98.  1956, Distribution and taxonomy of f i s h e s i n the Mackenzie Drainage of B r i t i s h Columbia, J , Fish.iRes, Bd, Can, 13(6)2759=789, 1957 Possible e f f e c t s of water diversions on f i s h d i s t r i b u t i o n in B r i t i s h Columbia, J, Fish, Res, Bd, Can, 14(4),651=668, 0  1961, The bearing of experimental m e r i s t i c studies on r a c i a l analys i s of f i s h populations, Proc, Ninth Pac. S c i , Cong, 1967 2 54-58, 1962, Experimental study of meristic variation in a population of threespine sticklebacks, Gasterosteus aculeatus, Canad, J, Zool, 40(1962) % 272=312, Lord, J, K, 1866a, The n a t u r a l i s t in Vancouver Island and B r i t i s h Columbia, Richard Bentley, London, 1866, 121=358. 1866b, The n a t u r a l i s t in Vancouver Island and B r i t i s h Columbia, Richard Bentley, London, 1866, 2 21=375. 0  M c A l l i s t e r , D, E. 1957, The systematics of the freshwater sculpins of B r i t i s h Columbia, M,A. Thesis, Univ, B r i t i s h Columbia Depto Zool, A p r i l , 1957sl=109, and C, C, Lindsey, 1959, (Cottus) of B r i t i s h Columbia,  Systematics of the freshwater sculpins Nat, Mus, Can, Cont, Zool,, B u l l . 172266-89,  «,«..„«. i960. L i s t of the marine fishes of Canada, 168, B i o l , Ser, 62 21=76. 0  Nat, Mus,  Can,  Bull.  Meek, S„ E„ 1899. Notes on a c o l l e c t i o n of cold=blooded vertebrates from the Olympic Mountains. F i e l d Mus. Nat, Hist, Publ. 31, Zool, Ser, (1897)1 (12)2225=236, Mottley, C. McC. 1934, The e f f e c t of temperature during the devedopment on the numbers of scales in the Kamloops trout, Salroo kamloops Jord. Contr, Canad, B i o l , , N„ S. 8s254=263, Nichols, J. T. 21sl71-174,  A small c o l l e c t i o n of Alaskan f i s h e s ,  Proc, B i o l , Soc, Wash,  Pritchard, A, L. 1936, Stomach content analysis of fishes preying upon the young of P a c i f i c Salmon during the f r y migration at McClinton Creek, Masset Inlet, B r i t i s h Columbia. Canad. F i e l d - N a t u r a l i s t 50(6)2 104-105, Richardson, S i r J, 1836, Fauna Boreali-Americana, Richard Bentley, London, Part 3 21-327,  Part 3,  The  fish.  88  =„ 1844, Ichthyology Ins Voyage o f the H, M. S, "Sulphur", under the command of Captain S i r Edward Belcher during the years 1836=42, (R, B, Hinds, ed,), Not seen, 0  9  Robeck, G, G,, C, Henderson, and R, C, Palange, 1954 Water q u a l i t y Studies on the Columbia River, U, S, Dept. HEW, Pub, Health Svc,, R, A, Taft San. Engr. Center, C i n c i n a t t i 1954sl=99 plus 4 Appendices, 0  Robins, C, R, and R. R, M i l l e r , 1957, C l a s s i f i c a t i o n , variation and d i s t r i b u t i o n of the s c u l p i n s , genus Cottus, inhabiting the P a c i f i c Slope waters i n C a l i f o r n i a and southern Oregon, with a key t o the species, C a l i f . Fish and Game 43(3)s213=233, Rutter, C, 1908, The f i s h e s of the Sacramento=SanJoaquin Basin, with a study of t h e i r d i s t r i b u t i o n and v a r i a t i o n . B u l l , U, S, Bur. Fish, 27(for 1907)sl03=152. Schultz, L o P. Washington,  1929, Check l i s t o f the fresh=water f i s h e s o f Oregon and Publ. F i s h , , Univ. Wash, C o l l , Fish. 2(4)s43-50,  -—-—, 1930, Notes on the species o f Cottus i n western Washington, 1930(1);14-15. '  Copeia  and A. C, DeLacy, 1936a, Fishes of the American Northwest, A catalogue o f the fishes o f Washington and Oregon, with d i s t r i b u t i o n a l records and a bibliography. Installment 3, Mid-Pacific Mag, (April-June) 1936sl27-142. ..._. and =, 1936b, Fishes of the American Northwest. Mid-Pacific Mag, (July-Sept,) 1936S211-226. (  Installment 4,  1936. Keys t o the f i s h e s of Washington, Oregon and closely adjoining regions. Univ. Wash, Publ, B i o l . 2(4)sl03=228. Seale, A. 1896. L i s t o f fresh-water fishes c o l l e c t e d in the v i c i n i t y of Seattle, Washington, by Edwin C. Starks, Proc, C a l i f , Acad, S c i , ( 2 ) 5 (for 1895) part 2s852-854. Seymour, A. 1959, E f f e c t s of temperature upon the formation o f vertebrae and f i n rays i n young chinook salmon. Trans, Am. F i s h . Soc. 88J58-69, Shapovalov, L, and W. A, D i l l , 1950, A check l i s t of the fresh-water and anadromous fishes o f C a l i f o r n i a , C a l i f , F i s h and Game 36(4)$382-391, -—-«-•, and A, C, Taft, 1954. The l i f e h i s t o r i e s of the s t e e l head rainbow trout (Salmo gairdneri gairdneri) and s i l v e r salmon (Onchorhychus kisutch), C a l i f . Dept. Fish and Game, Fish B u l l . 98s1=375, -  H  w. A, D i l l , and A, J, Cordone, 1959, A revised check l i s t of the freshwater and anadromous f i s h e s of C a l i f o r n i a , C a l i f , F i s h and Game .. 45(3)sl59-180, g  Snyder, J . 0, 1905, Notes on the f i s h e s o f the streams flowing into San Francisco Bay. Rep, Bur. Fish, (for 1904), App. 5s327-338,  89  1908a, The fauna of the Russian River California, and i t s relation to that of the Sacramento, Science, N , S 27s269-271, 9  0  „ 1908b, California,  The fishes of the coastal streams of Oregon and northern Bull, U. S, Bur. Fish, 27(for 1907).153-189.  ——. .1913, The fishes of the streams tributary to Monterey Bay, California. Bull, U, S„ Bur. Fish, 32(for 1912)s47-72, 1916. Fishes of the streams tributary to Tomales Bay, U, S. Bur, Fish, 34(for 1914).375-381,  Bull,  Storer, D. H, 1846a, A synopsis of the fishes of North America, Mem, Amer. Acad. Arts and Sci., N, S, 2:253-550, —, 1846b, A synopsis of the fishes of North America, Metcalf and Co., Cambridge, 1846s1-298, Strawn, K, 1961. A comparison of meristic means and variances of wild and laboratory-raised samples of the fishes, Etheostoma grahami and E. lepidum (Percidae). Texas J. Sci. 13(2)sl27-159. ~° Suckley, G, 1859, Report upon the fishes collected on the Survey, Chap, l i s Report upon the fishes exclusive of the Salmonidaes 350-368, Ins The natural history of Washington Territory, ^Cooper, J . G,, and Suckley, G, Baillere Brothers, New York, Part 111:399, 1960. Report upon the fishes collected on the Survey, Chap 2s Report upon the fishes exclusive of the Salmonidaw, U, S, Pac, R, R, Expl. and Surv,, (for 1853-55), 12(2) part 3 (5):350-368, Also appears as: H. Rep. Ex, Doc, 56, Wash., 1860:350-368. Sumner, F, H, 1942, A reconnaissance survey of Oregon coastal streams. Rep. Oregon St, Game Comm., 1-25 (mimeo). In_2 Bond, C, E, 1963:81. ——•. 1953. Migrations of salmonids in Sand Creek, Oregon, Fish, Soc. 83(1952).139-150. Taft, A, C. 1943 California steelhead experiments. 64:248-251,  Trans, Am.  Trans, Am, Fish, Soc,  Taning, A, V, 1944, Experiments on meristic and other characters in fishes, I, Medd, Komm. Dan. Fisk. Havunders, ll(3)sl-66, , 1952, Experimental study of meristic characters in fishes, Rev, Cambridge Phil. Soc. 27:169-193, Wilimoysky, N. J. 1954. List of the fishes of Alaska, 4:279-294, —  , 1958. Provisional keys to the fishes of Alaska, Wildlife Svc,, Fish, Res, Lab., Juneau. 1958:1-113.  Biol,  Stan, Ich, Bull, U, S, Fish and  Appendix Table 1 Pop, No.  0  L i s t o f C o l l e c t i o n s , L o c a l i t i e s , and Other Pertinent Information.  General Locale C o l l e c t i o n No 0  Specific Locality  .60 mm S.L,  :  60 mm S.L,  NoLato  W.Long,  COASTAL POPULATIONS Alaska Seward area: BC 64-311 "ditch a t a i r p o r t " NMC 61-133 "lagoon at north edge of Seward" Haines area: USNM 76408 "head o f Chilkoot L," Juneau area, Lynn Canal, Favorite Channel:. Petersen Creek, 12 mi, N,of Juneau AB 61-100  AB 61-489  "  v.-,..-.-" "  60°07' 149°24» 1(56, 0) 9(28, 2-38,6) 59°23» 135°35« 8( 62,9-125,0) 58°22' 134°36« 2(53,9-59,5)  8( 73.2« 119.2) 2( 70,1• 77.8) 8( 78.0•142.1)  AB 61-490 " .. ". Loring area, Revillagigedo Id. s 55°35' 14(112,8= •166.2) USNM 64130-49 Loring 2( 87,5= •f,'94.0) USNM 60801 Deep Bay 3(115.4= •135.0) USNM 60371 Mouth of River 2( 68,9• 76,0) USNM 60316 Naha River Kasaan Bay, Prince of Wales Id. 55°00« Stream o f Karta Bay 2(111.6- •116.7) SU 23989 35' Saltery -Cr, j Paul's Bight, Skowl Arm 11( 83.9< •139.0) AB'62-76 25' AB 62-77 2(149,5« •154.5) Saltery Cr, AB 63-167 3( 94,3= 112.1) Saltery Cr,, 100 yds, up from mouth B r i t i s h Columbia Queen Charlotte 53°00« 6. Is. . Graham Id.. Juskatla Inlet , of Juskatla 10(28,5-56,6) 9( 60.2- 90.2) 37' BC 60-437 Small stream about 2 mi. W 59(26.4-59,6) 11( 62,2-136,2) BC 60-439 Mouth, o f stream 3 mi, W, of Juskatla 36« It ; ... I I I I .'. It It II II II . 38(32.7-59.5) 6( 61,5- 83.3) BC 60-440 Queen Charlotte Is, , Moresby Id, : 53°06» 7, Creek-entering Skidegate L, 23(25, ff-5?.&) 20( 61,0-132.2) BC 60-420 12(25, 3-59,1) 6( 62^.3- 98.5) BC 60-424 Shore o f Skidegate L, Princess Royal Id.: 52°57' 2(122.0-123,5) BC 53-214 Bloomfield L,, N.E, s i d e , Laredo Inlet King I d . , Ocean F a l l s area: 52°06' 9. 4(63 ,1-70,8?* 16( 72.4- 85.7^* BC 54-434* Port John L. 51°40' 10, Rivers Inlet area: BC 56-629 Owikeno L.„ 2nd Narrows* 10(none l i s t e d ) *Counts taken by D.E. M c A l l i s t e r ; **Lengths given are T o t a l Lengths, 1  131°40«  132°00 35' 30* e  132°00» 23* 25° 132°00'  128°50« 127°45' 126°40'  Appendix Table 1, List of Collections, etc. (cont'd). Campbell R. area: Vancouver Id 11. Buttle L, BC 54-114 Buttle L, BC 64-312 Buttle L„ BC 64-313 Upper Campbell L. BC 64-314 Qualicum area: 12, Vancouver Id Horne L, BC 54-39 Qualicuin area: 13. Vancouver Id Nile Cr. , 100 paces from mouth K61-13 South ends Vancouver Id .14. BC 63-1462 Nitinat L, 15, Gambler Id,, Howe Sounds Outlet Cr. 100 f t , from mouth BC 58-410 16. White Rock area, Little Campbell R, K 60-52 Stn, C-22 II K 60-40 n K 60-71 II K 60-88 II K 60-128 II K 60-169 II K 60.-27 c- 2 tt K 60-12 it K 63-6 II c-•3 K 60.-80 ti K 60-101 it K 60-116 II K 60-139 17. Saturna Id, Strait of Georgia; Lye 11 Harbor Cr, BC 56-81  19. 20. 21.  Clatsop Co. , Cannon Beach areas UMMZ 93427 Mouth of Elk Cr, Coos Co. Lakeside, Ten Mile Lake BC 63-1089 Ten Mile L. outlet wier Curry Co.j Sixes areas SU 9270 Flores R„ Curry Co,, Winchuck Area, Winchuck R,s OSC 198 2 mi, upstream from U,S, Hwy 101  1(42.5)  5(49,4-59.6) 14(39,9-59.5) 17(23,0-59,9)  1(  39,5)  33(13,0-58,5) 5(55.0-58,6) 5(51,6-59,5) 2(57,5-58,5) 10(28,4-39,8)** 10(21.7-59,4)  9  106.5) 49°47' 125°40' K 2( 71.0- 76.0) 2( ca75-117.5) 132.7 ) 49°55' 125°40' 1( 49°20' 124°40' 11(108.1-132,0) 49°25' 124°40' 17 ( 61,5- 91.0) 48°40' 124°50' 63,089.2) IK 49°25' 123°23' 16 (60.5-129,3) 4 9 ° — 122°— 2( 89.7- 90,2) " 04' ii 1+0' 10( 77.5-107.7) 2( 92,8-127.5) 3( 93.0-102, 4) 2( 80,7-111,0) 2( 80,0- 89,5) 9( 60,6- 71, 8) " 00' " 46' 3( 64.0- 88.5) 6( 81,5-118,5) 2( 70,1- 71.6) " 01' « 45' 9( 60.5-97,3) 7( 60,4- 73.3) 9( 63,5- 83.2) 48°48' 123°10'  5( 68,4-100.2 ) 33( 60.3-114,1)  3(44,2-55,3) 6(36,3-50.9)  9( 77,0-154,0)  45°54' 123°58' 43°35« 123°10' 42°55» 124°30« 42°00« 124°15«  Appendix Table 1,  List of Collections  9  etc  (cont'd),,  California ife Del Norte Co, Crescent City.areas SU 23598 Point St. George CAS 18025 Wilson Cr., at bridge 1-05, U,S„ 101 3(31,7-397) CAS 18028 Smith R., a t bridge on U,S, 199 Del Norte Co.s 23, 14(33,3=59,8) SU 9272 Mouth of Klamath R,, at Requa Mendocino Co,, Navarro area, Navarro R,s 24a, CAS 19066 Indian Cr,, bridge on Hwy, 128, Philo SU 35131=3 N, f o r k , o.2 mi, E, o f Flynn Cr, SU 38016 Albion R., 2 mi. below Comptche 1(49,6) Marin Co., Tomales Bay drain,, Lagunitas (Papermill) Crtt 25a. CAS 186I6A S,P„ Taylor St, Park, a t Camp Taylor 6(22,7-44,4) SU 40849 Near Point Reyes Station 26. Santa•Cruz Co,, Waddell Cr.s CAS 20857 3 mi. S.W, of Ano Nuevo Point CAS 20862 " " " " " " " 27, Santa Cruz Co., So quel areas 1(54,7) CAS 19086 Soquel Cr,, a t Soquel SU 4816 Soquel Cr, San Luis Obispo Co., San Simeon areas 28. 4(22,9=42,5) SU 15183 San Simeon Cr, Santa Barbara Co., Lompoc areas 29, SU 40323 Santa Ynez R, 30. Santa Barbara Co,, Gaviotas 15(21,7-47.4) UMMZ 131728 Gaviota Cr., at mouth 31. Ventura Co., Ventura R*s UMMZ 13173ft 6.4 mi. N.E. o f Ventura UMMZ 132891 1/2 mi. from Foster Park, towards Ventura 22,  9  K K  62,3) 84,7)  41°50« 124°15« II it I I it " 55 • it II 41°34  e  124°10«  39°09  e  123°34  38°00  e  122°45»  37°06  9  122°17»  16( 62,0=115,0) 8  ) 1( 63.0 3( 71.6=106.5) 1( 93,0) 24( 96,0=143,2) 14( 95,8=143.1) 36°57« 12£°55« 4( 64.9=105.2) I35°35» 121°12  8  2(124,9=127,2) 34°41' 120? 35' 2( 76,8= 86.3) 34°24» 120°15' 34°10' 119°20« 1(113,5 ) 2(119,0=151.1)  Appendix Table 1.  List of Collections,  etCo  (cont'd),,  COASTAL DERIVATIVE POPULATIONS B r i t i s h Columbia 32. 33.  34.  35. 36.  37. 38,. 39. 40. 41. 42. 43.  Stikine R„ drainage s BC 54=485 Cold Fish Lake BC 54=485 Cold Fish Lake* Nass R. drainage, Meziadin Lakes BC 55=221 At outlet BC 55=221=A At outlet BC 56=536 O f f McLeod Creek Skeena R, drainage, Babine Lake s BC 55=452 Wright Bay BC 55=453 Opposite Donald's Landing BC 56=455 At Topley Landing Fraser R. drainage, Takla Lake outlets BC 54=249 Middle R. Mackenzie R. drainage. Nation Lakes areas BC 64=102 Tchentlo L. i n l e t stream mouth BC 64=101 Tchentlo L, i n l e t end Mackenzie R. drainage, Mansori Creek areas BC«56£458 Outlet of Wolverine L, Mackenzie R drainage, Parsnip R, watersheds BC-61=464 Tacheeda Lake Mackenzie R. , Crooked R. watersheds BC 54-477 McLeod Lake c Mackenzie R., Crooked R„ watersheds S-98 Bear L. N. of Summit L. Mackenzie R,, Crooked R« watersheds BC 61-463 Hart L. o f f Hart Highway Mackenzie R. , Crooked R, watersheds BC 56-362 Summit L, , N. of Prince George BC 54-459 Summit L., N„ o f Prince George Dean R. drainage, C h i l c o t i n areas BC 56-483 Nimpo L„  56°40 3(48,6-66,7)^ \Wi  2(77,7-^82,5) 1(81,7 >*% ) 8(81,0=109.1) 5(66,2- 82.-0) 11(73,2= 87,2)  12(27.4-35.2)  128°40«  55°03" 129°10»  54°50  9  126°0=05«  54°55  8  125°05  8  55°12  9  125°00  5  55°37  9  124°25  9  54°42  B  122°30°  55°00  9  123°00  9  4(61,5=141,0) 54 28 0  9  122°47  9  54°15  9  122°40'  1(60,6 9(15.8=34.4) 6(34.2=54.3)  9  )  3(61,1= 96,9) 4(65,5= 83,5)  3(41.2=54.7) 3(33.1-47.0) 13(31.0=58,0)  11(65,0=133,3)  0  13(34,0=58.5)  2(65,6-666,2)  15(13,8-23,0) 3(48,5=59,9)  2(79,5- 80.1) 1(25.9 ) 11(10.7=50,8) 52°20« 125°10 15(12,5-34,4)  9  Appendix Table l  n  L i s t o f C o l l e c t i o n s , e t c . (cont'd),,  INLAND POPULATIONS B r i t i s h Columbia Skeena R. Drainages Lakelse R. , at Terrace . 44, BC 56- •411 ' 45„ Bulkley R , 1 mi. below Bulkley BC 56- #53 Fraser R„, Nechako R. drainages 46. BC 55-10 Fraser L., 2 mi from West end 47. BC 54-162 E u l a t a z e l l a L „ Deep Cr. 43. BC 54-367 Nadsilnich L., St. Joseph's Cr. Fraser Thompson R. drainages , 49. BC 54-242 Kamloops L. 50. BC 55-139 Pothole L o u t l e t (Quilchena Cr.) Fraser Fraser Valley drainages 51„ K mi s c, Squak urn L o u t l e t stream 0  e  52,  53.  54,  II  it  K 62-19 BC 54-197 BC 54-196 BC 54-195 V-100 V-150 V-203 BC 59-612 BC 54-43 BC 54-427 BC 59-504 BC 62-70 K misc. K 63-16 K 63-17 SW-1 BC 56-564 BC 54-212 BC 54-209 BC 56-120 BC 55-313 K misc.  it  .  II  ti  It  r~ I I  II  t l .  .  ._  II  .  II  15(16,8-55,6) 6(43,5-57,9) 9(53,6=67,0)  l(468 il ) 54°05' 124°50« 15( 62,2=105.5) 53°42' 123940' 53°45" 122°50»  11(35.0-59,0)  t  ti  15( 60,5=153,6) 54°21« 128°35 9( 67,1=130,7) 54°28 126°30'  1(47.6  )  II  r  It  ...  II  Vedder R. Stn. V-6, at end o f Ford Rd, 15(37.6-58,6) II  ti  II  ... i t  .  it  it  tt  it  II.  n v-4, Vedder R, Canal Vedder Canal, above highway bridge Vedder R, it  II it  Cultus L. Cultus L. , outlet wier i n Sweltzer Cr, II  tt  it  ti  -  t i  II  it  it  tt  -511  II  It  tt tt  Sweltzer Cr., 1/4 mi, below Cultus L, Hatzic L., Hatzic Slough Swan Point tt  ti  II  tt  II  it  it  tt  II  it  !  7( 9( 1( 7(  73,8=104,8 68,4=111,9 68,8 73,9-119,0  2(33,5=43,0)  Squakum L, ii  c  (  50°45« 120°40' 10(14,6-25,7)ft* 10( 60,0- 88,7) 49°55» 120°30» 12(25,3-58.Q)  7  II  14(40,8-55,9) 3(34,4-50,4)  Edwards Or, " " mouth ti  tt  13(17,6-51,0) 6(24,2-32,2) 1(20,9 ) 13(32,2-50.5T  5(103 3(113 U3( 6 1 1 ( 97 2(116 3(113 3Q.09 2(106 1(104 5( 62 5 ( 74, 3 ( 72, 2 ( 74, 2 ( 70, 4 ( 64, 9 ( 75,  118,5 •143,8 •100,1  49°14' 122°00'  -F  49°06' 122°00  8  •129,0 •148,2 128,1 •140,0  •105,7 > 96,5 • 77,0 • 94,3 •116,0 • 70,1 •112.9 3(104.5-•130,2  1(138,2 2( 66,7- 68.0  49°04' 121°5-9»  49°10» 122°14'  Appendix Table 1,  List of Collections, etc, (cont'd).  S„ Alouette R, Stn, A-8, 14th Ave, A-51 it it ti A-122 tt A-6, 5th Ave, A-153 it ti tt ti ti A-220 it tt it ti ti K 63-12 tt ti A-7, 8th Ave, Columbia R drainage, Revel stoke areai Williamson L, BC 54-369 56, BC 5 9-424 tt it off diving float Columbia R„ drainage, Grand Forks area Christina L, 57„ BC 54-124 ti it 1 mio from S, end BC 56-534 Columbia R, , Okanagan R„ drainage s Osoyoos L, 58, BC 53-187 Washington-Oregon Columbia R„ drainage s Moses L o , Grant Co,, Wash, USNM 104608 59 McNary Dam, Benton Co,, Wash, UW 14614 UW 14747 Umatilla R, mouth, Umatilla Co,, Ore, OSC misc, 55,  3(30,0-54,2) 13(15,0-59,6)  0  OSC OSC CAS  "• " (IoUcM,  tt  II  it  II  tt  ti  Columbia R, drainage, Portland and Vicinity? OSC misc. Abernathy Cr,, at mouth, Cowlitz Co, 61, Kalama R 1/4 mi, above mouth Coweman R at mouth ti tt 100 yds, upstream " " Lewis R, j E . f o r k at U,S,'99, Clark Co, below U,S, 99 bridge it Washougal R, , 1 mi, above mouth " " " " , below old Hwy 30 bridge " Vista Slough, Big Rock, Multnomah Co, Sandy R, below old Hwy 30 bridge " Sandy R» Clackamas R„, near mouth, Clackamas Co, 0a  14(27,8-39,8)  3( 73,5- 82,6) 2( 95,4-103,4) 3(100,4-123,0) 5( 64,5-107,7) 8( 65,5- 97.0) 13( 86,0-123,6)  49°14' 122°35  50°59' 118°11 49°09' 118°11  3(34,4-42,6) 11(19,8-34,4)  1( 73,3  23(36,3-59*9)  6( 60,5- 79.8)  20(18,8-26, 6) 1(59,5 ) 3( 70,4-161,5 1(58,9 ) 4( 76,8-100,3 2(125,4-169,2 1(149,5 3( 93,8-186,0 1( 62,2  )  49°11' 119°30  47°06» 119°19' 45 55" 119°20 0  l  45°20 -122°10 46°02 122°52' 9  2(118,8-119,5 1(113,5 1(105,6 3( 88,0-172,0 4(112,5-164,5 1(121,7 3( 94,6-106,0 1(146,5 1( 97,2 2(110.9-121.5 1( 91,4 1(175.0  8  (  Appendix Table 1 ,  L i s t of C o l l e c t i o n s , e t c , (cont'd),  Columbia R o mouth, Washington Shore, P a c i f i c Co,: UMMZ 9 3 4 2 5 Columbia R at end of Chinook P i e r 62, OSC misco Chinook R. OSC misCo Chinook R , , above tide gate SU 3 8 0 1 9 Columbia R, , 1 / 2 mi. W. of Megler Ferry Columbia , mouth, Oregon Shore, Clatsop Co,s SU 3 8 0 1 8 Columbia R, at A s t o r i a 63o  6 (  0  SU  3 8 0 1 7  "  "  "  1 ( 2 4 , 6 )  e  46°11'  1 2 3 ° 5 0  s  4 4 ° 1 5 =  1 2 3 ° 1 0 =  30"  1 5 "  9 7 , 5 - 1 5 0 , 6 )  2 (  8 6 , 5 - 1 2 8 , 2 )  2(  6 0 , 1 - 1 1 2 , 7 )  7 ( 1 2 7 , 5 - 1 5 4 , 0 ) 1 1 ( 1 0 0 , 8 = 1 3 1 , 5 ) 2(  87,8=  8 9 , 8 )  4 ( 1 1 9 , 3 - 1 3 4 , 5 )  2 ( 1 5 0 , 3 = 1 5 0 , 7 ) 2 ( 1 2 3 , 5 - 1 3 3 , 5 ) 1 ( 5 3 , 4 ) 1 ( 3 4 , 5 )  1 1 ( 1 7 , 5 - 5 6 , 1 )  California Sacramento R. drainage, Shasta Co,s 2 ( 2 8 , 5 = 3 1 , 0 ) 66a, CAS 1 8 0 8 1 O'Brien Cr,,. emptying i n Shasta L, Clear Cr„, a few mi, S, o f Redding UMMZ 9 3 4 2 8 Fort Reading (on Cow Cr») 66b, USNM 2 9 5 Sacramento R, drainage, Tehama and Glenn Co,s Sacramento R,, v a l l i t t l e below Red B l u f f 6 7 , UMMZ 9 3 4 2 9 ' UMMZ 9 3 4 3 0 Thomas Cr. .at R i c h f i e l d UMMZ 9 3 4 3 1 Stony C r , near Orland Sacramento R, drainage, Sacramento area s 68o OSC m£sc. Sacramento Brickyard Pond, S, o f c i t y San Joaquin R, d r a i n a g e , San Joaquin Go,; 6 9 , CAS 2 0 8 8 7 San Joaquin R,(exact l o c a l i t y unknown) 10(31,5-56,5) San Joaquin R, drainage, Stanislaus Co.; 4 ( 2 7 , 5 - 4 4 , 9 ) 7 0 , CAS 1 7 9 6 6 I r r i g a t i o n d i t c h on Hwy 1 3 2 , 1 mi, E, 3 ( 2 5 , 0 - 5 0 . 5 ) CAS 1 9 0 9 8 o f Empire. " " " " " " " San Joaquin R, drainage, Madera and Feesno Co.; 7 1 , CAS 2 0 2 5 2 Madera Canal, presumably from M i l l e r t o n 1 ( 3 0 , 0 OSC misc. San Joaquin R, at Mendota 0  1 2 3 ° 5 7  2 ( 1 1 6 , 5 - 1 1 8 , 0 )  "  OSC 7 6 2 Walluski R . , 6 mi, S.E. of A s t o r i a OSC misco Big Creek, at mouth Willamette Ro drainage, C o r v a l l i s areas Overflow slough at C o r v a l l i s OSC 1 8 9 64 o C o r v a l l i s Sand 6 Gravel P i t s , along R, OSC misc. Pothole, Willamette R, at C o r v a l l i s OSC 4 0 2 Stn, 5 , above C o r v a l l i s OSC 5702 Stn, 6 , Harrisburg OSC misc. Stn. 7 , Peoria OSC misco Willamette R, drainage, S o E „ o f Eugene, Lane Co,; , Lookout Point Reservoir, Landax 6 5 . OSC 7 4 4  46°17«  K  7 3 , 3  2(  66.8=  1 0 (  1(  6 5 , 0 =  78,4  ) 7 4 . 6 ) s  122°40'  4 0 ° 4 8  8  1 2 2 ° 1 6  8 4 , 9 )  )  3( $6*7=105,2) 1( 69,71( 99,0 1( 71,8  4 3 ° 4 5  II  3  0  «  II  !  it  ti 2 8 ' " 11*'  ) i40°09' 122°13' ) o » i22°13' ) 3 9 0 4 5 s 122°13 3 9  5 6  9  2( 65,5-100,7)  38°30' 121°30'  4(61,0-84,0)^370^59 37°35»  121020' 120°55'  1  1(120,9  K  72,1  )ca37°00' 1 1 9 ° 5 0 ' ) 37040* 1 2 0 ° 2 0 '  to cn  Appendix Table 1,  L i s t of C o l l e c t i o n s , e t c (cont'd),,  San Joaquin R. drainage Fresno and Tulare Co.s ? i 1(46.5 ) 72. CAS 25541 Kings R. , at Dutch John Cut 36°15' 119°16" CAS 20904 Kaweah R., near V i s a l i a 3(34.5-58.3) INTERGRADE POPULATIONS 38°00' 122°10 Sacramento R. drainage. Contra Costa Co., Carquinez Straits 8(35.0-57.5) 20(60.0-183.4) 73. UMMZ 142373 ca 1/2 mi.E, o f Eckley, S. of D i l l o n Pt 38°ir 122°40 San Pablo Bay area, Sonoma Co., Californias 2(30.9-39.5) 2(71.8-109,5) 74. USNM 296 Petaluma Additional Material From the L i t e r a t u r e , Used Separately, or i n Combination, with Populations L i s t e d Above Sacramento R. drainages 2 specimens 40°30» 122°10 75. Rutter, 1908sl45 Sacramento R. at Redding 1 " 40°21» 122°09 " " mouth o f Battle Cr. 1 40°09' 122°13 " r " at Red B l u f f 1 " " at Jacinto 9 " • ••" " Arcade Cr., at Arcade 1 " 39?05 121°35 Feather R, at Marysville San Francisco Bay, San Mateo Co.s 34 specimens 37°24' 122'°0'9 76. Snyder, 1905s338 San Francisquito Cr, Santa Cruz Co,, Watsonville areas 25 specimens 77. Snyder, 1913s72 Pajaro R„ aeoSO* i21°50 Marin Co., Tomales Bay areas 10 specimens 25b, Snyder, 1908b$185 Papermill Gr. 38°00* i22 45 Mendocino Co., Point Arena areas 6 specimens 78. Snyder, 1908bsl85 Garcia R. 38°55» 123?45 Mendocino Co,, Navarro areas 11 specimens 24b. Snyder, 1908bsi85 Navarro R. 39°09» 123°34 Humboldt Co. , Areata areas 6 specimens 79. Snyder, 1908bs185 Mad:R„ 4Q°55' 124°08 Curry Co. Oregon, Gold Beach areas 8 specimens 80. Snyder, 1908bsl85 Rogue R,, near mouth 42°26' 124°23 Douglas Co., Reedsport areas 16 specimens 81. Snyder, 1908bsl85 Takenitch R. 43°45' 124°07' Tillamook Co., Nestucca Bay areas 10 specimens 82. Snyder, 1908bsl85 Nestucca R, 45°10« 123°57' King Co., Washington, Seattle areas 17 specimens 83. Snyder, 1908bsl85 Lake Washington 47°30» 122°15' Additional Materials UMMZ 131771, Kern R,, C a l i f o r n i a 1 specimen (not used) 35°-i i g o . _ B  9  f  u  Appendix Table 2. Population No. Coastal 1 2 3 4 5 6 7 11 12 13 14 15 16 18 19 20-21 25a 26 28 29 30 31 X Coastal Derivative 32 33 34 35 36 37 38 40 41 X  Weighted Mean Percentage P r i c k l i n g and Resorption f o r Individual Populations o f Cottus aSper Used i n Compiling fable I and Eigure 4. Specimens S=60 mm S.L. Specimens £i60 mm S.L. % Resorption Virtual Apparent Virtual Apparent n Relative Absolute n „_ — — — 37.9 34.2 1* — 17.0 49,9 8 34.1 17,1 33.6 14.0 19.6 18 58.2 34.6 17.8 2 — 9 20,2 41.1 14.1 34.^3 — 16 40.8 24,7 39.5 16.1 9 40,5 42.0 40.3 21,1 48.0 19.4 11 18 26.2 11,7 32.1 20.5 55,3 14.5 12 5 13.7 3.5 74,6 10.2 27.1 11 10,3 62.1 16.8 17 44.5 41.1 24.4 21,8 22.7 51.1 5 43.4 11 27.3 49.3 42.9 37.0 16.1 14 16, 43.0 54.0 51.0 24.8 42.3 18.2 17 29 44.2 41.4 27.5 12.8 40.3 31.8 16 5 31.0 15.4 15.6 50.1 49.7 41.6 10 — -«. 33 47.0 34.0 27,7 13.0 — 9 16.5 30.6 9 — ;j — 1* — 24,2 41.7 6 24 30,4 57,5 12.9 17.5 — —• -2 14,4 35.4 6 — 2 i7.6 -rs — --?* -— — 44.8 43.8 15 3 29.5 13.0 16.5 55.8 35.0 17.8 17.2 48.9 43.5 36.0  ~  5 12 6 13 13 6 — —  — 49.5 49.0 42.4 42.9 34.5 50.8 — 44 , 8  2  48.6 46,2 38.4 40.5 29.1 48,3 41,8  1 4 — 11 2  *  1 2 —  15.0 37.8 35.4 43.8 — 34.8 36.1 49.3 40.5 34.7  8.9 25.4 22.3 38.6 23.4 17.2 27.7 31.6 24.2  40.3 32.9 37.0 11.8 — 32, 8 52.4 43.8 21.8 32.0  6.1 12.4 13.1 5.2 — 11,4 18.9 21.6 8.9 10,5  Appendix Table 2 (cont'd).  Specimens <C 60 mm S, L,  Population No, Inland 44 45 46 47 50 51 52 53 54 55 56 57 58 60 61 62 63 64 65 66=68 69=72  Weighted Mean Percentage Prickling and Resorption for Individual Populations of Cottus asper Used in Compiling Table I and Figure 4,  n  Virtual  15 3 4 6 12 11 15 13 20 12 14 8 24 1*  67,7 69,3 65,1 65,4 61,4 65,4 56,4 63,1 77,6 69,8 52,3 66,9 66,5 84,2  ==  •  Apparent 64, 8 65,8 61,6 60, 7 56,4 60,6 53,5 61.0 73,3 67,0 51,1 66,3 65.1 81,2 —  67.0  2 ™  X  2 11 2 23 ==  81,9 74,5 71,5 67,2  77,4 80,6 74,5 68,0 65,3  60,1  57,1  Specimens I> 60 mm S.L, n  Virtual  14  64,1 65,0 62,6 59,9 59,6 55,3 59,9 58,3 73.5 67,4 51,0 61,3 67,1 69,5 67,3 58, 8 60,7  Apparent  % Resorption Relative Absolute 16,0 32,0 3,6 9,4 16.3 16,6 12.1 14.7 7,5 20,7 11.8 1,7 10,1 16.9 17,1 28,4 7,1  10,2 20, 8 2,2 5.2 9.7 9.2 7.3 8,6 5.5 13,9 6,0 1,0 6.8 11,7 11,5 16,7 4,3  ==  76,7 63, 3 63,3 63,2  53.9 44,2 60,4 54,3 49,9 46.1 52,6 49.7 68,0 53,5 45,0 60,3 60,3 57,8 55,8 42,1 56,4 57,0 65.8 46,2 52,6 53,2  14.2 26.9 12,7 16,1  10,9 17,1 7„7 10,0  20  52,9  35,6  32.8  17,3  9  *  1* 15 10 31 25 15 21 20 13 1* 6 7 15 6 11 7 10 6 7  = =  Intergrade 73  8  Single specimens not included in means (X)  Appendix Table 3.  Non-weighted Mean % V i r t u a l P r i c k l i n g i n Each of 14 Body Sections (Specimens> 60 mm S.L.)  Population , Sample No. Size  a  b  Lettered Body Sections (from Skin Map, F i g . 1) e d c f h I .g  "f  k  m  1  n.  Coastal 2 3 4 5 6 7 11 12 13 14 15 16 18 19 20-21 25a 26 28 29 31  8 18 ;8 16 9 18 5 11 17 11 16 6 5 33 9 1 24 2 2 3  _ = — . . —  —  0.9 0^9  — —  — • '  —  0.7  ft  6.9 8.9 4.7 - =  —  ft* 4.0 3.0 16.5 -  -  70.6 65,9 65.6 78.8 86.7 56.9 2.4 32.7 90.9 90.0 90.3 85.8 26.0 92,3 (6.7)  o o  —  3.1 10.6  1.9 —  —  —  —  7.8 3.3 4.7  — = = " - =  20.6 9.1 20.0 ... ==w  8.9 —  o ~  —  2.8  1.1  12.0 9.5 14.7 5.8 4.0 27.4 (10.0) •  50,4  — - o  —  —  35,6 31,1 25,6 41.2 28.3 26.1 18,0 14.1 53.2 56,8 50,3 40.8 46.0 65.9 (39,4) 15,0 20,6 -  -  5.0 5,0'  53.3  72.5 32.5 55.6 60, 3 42,8 53.6 72,8 41,1 51,1 75.9 44.7 c66,6 85,6 .41.1 66.1 48,9 32.2 37.5 25.0 25.0 17.0 47,7 ) 30.9 41.4 .88.8 11.8 43.5 69.7 84, 5 2.3 42.3 67,3 86,6 2.1 43.4 64.4 38.3 67.5 8,0 J6 50', 0/ 41.0 47.0 87.0 3.5 43,2 74,1 (21,1) (35.6) (11.1) 50.0 35,0 50,0 57.9 52,3 36.2 20.0 17.5 35.0 30.0 56.7 43.3 61.7 o o  r  -  v  -  '  =  —  -  OO  a .  — —  2.2 1.1  — _  oo  oo  — —  oo  o o  o —  oo  o o  o o  1.2*  oo  oo  o o  =.=  o o  ——  oo  -  — -  '••  „o  o o  '  — —  o o  o  —  OO  oo  OO  oo  -o  o-  —  _ »  .  • !»—•?"!• ;;  1.9 5.8 10.0 5.3  oo o o  o o  — o o o o o o  OO  =  . —  -  —  n  —  Coastal Derivative  <  32 33 34 35 37 38 41  - .  2 19 1 4 11 2 2  2.6 —  1.0  ft  oo  2.2 5.4 _ «  —  7.0  32.5 87.9 80.0 97.5 83.6 90.0 90.0  Oo  2.6  6.8  -  -  -  23.8  —  -  18.8 0.4 20.0 —  15.0 46.1 65.0 56.2 44,5 37.5 52.5  25.0 83.4 65.0 93.8 7,6.8 80.0 82.5  —  15,0 •  — — ——.  25.0 32.6 35.0 38.8 32.3 30.0 37.5  15.0 57.6 50.0 62.5 48.2 55.0 62.5  2.5 —  ft  — -  5,0 2.5  — —  Appendix Table 3 (cont'd). Population. Sample Size No.  Non-weighted Mean % V i r t u a l P r i c k l i n g i n Each o f 11 Body Sections (Specimens ^ 6 0 mm S.L.) b  a  Lettered Body Sections (from Skin Map, F i g . 1) e c d f h i g  j  k  1  m  n  Inland.  11 15 16 17 5Q 51 52 53 51 55 56 57 58 60 61 62 63 61 65 66-68 69-72  11 9 i 15 10, 31 25 15 21 20 13 1 6 7 15 6 11 7 10 6 7  33.3 31.6 75.0 23.7 10.0 26.3 15.8 11.3 82.6 75.7 6.3 60.0 77.5 68.3 87.0 26.7 51.8 (11.3) 91.5 61.0 17.1  +  68.9 99.6 52.8 100.0 70.0 100.0 51.0 99.0 65.0 ibo.o 28.0 97.7 65.6 99.0 13.5 98.7 93,8 100.0 91,8 100.0 15.3 98.8 75.0 100.0 91.2 100.0 92.1 99.3 95.3 98.7 13.3 100.0 89.5 91.1 (78.6) (93.6) 100.0 100.0 91.2 100.0 82.8 95.0  56.8 68.3 65.0 59.3 16.5 31.6 .11. 0 29.3 65.0 61,5 38.8 70.0 72.5  100.0 60.6 100.0 57,3 99.0 82.3 91.H 71.3 100. d 95.2 62.3 100.0 100.0  98.6 98.9 86.1 100.0 100.0 100.0 99.0 89.0 92,5 100.0 81.0 95,5 91.8 97,2 85.7 95.3 99.5 99.5 99.2 100.0 72,7 91.9 100.0 100.0 100,0 99,2 50.7 98.6 97,1> 100,0 37.3 93.3! 100.0 96.0 15.0 75.0 89,2 98.3 27.7 95.1 90.9 (25.7) (97.9) (95.0) (79,3) 69.5 100.0 100,0 100.0 57.5 97.5 100.0 100,0 31.1 80.0 98.6 93.3  57.5 56.7 55,0 50.3 17.0 53.5 55,0 13.7 55,7 61.7 69,5 57.0 57.0 69.6 45.8 50.0 35.0 53.3 51.2 35.0 60.7 33.0 60.3 52.5 26.7 22.7 60.0 (7.1) (60.0) 19.5 72.0 19.2 50.8 5?.a 33.6  91.8 10.7 97.2 22.8 85.0 =90.7 25,3 98.0 6,0 ==. 83.1 87.2 6.6 87.0 1.0 96.1 31.1 95,2 20,2 80.0 0.8) 10,0 85,0 93,3 20.8 91.1 20.0 88.7 10,0 1.7 83.3 78.6 1.1 (73.6) (2.8) 96.0 . 30.5 89.2 11.7 90.0 11.1  21.8 3.6 32.2 19.1 -- 10.0 17.3 13.0 -15.2 -1.1 =-' 15,7 2.8 12.9 5.0 --• 21.8 5.5 Tt: ), 2.8 10.0 27.2 5.0 — 17.1 15,7 0.7 30.3 1.7 20.8 i.i 11.8 (0.7) (9.3) 7.0 59.0 8.0 — ' 11.7 — 3.6 — 1.2  mm~  <-,<» •»B  _»  •»«  --  ?  Intergrade  73  37.5 60.6 25,0 53.0 35,5 12.6 29,8  20  25.2  50.0  96.5  38.0  55.2 : .  8.1,2  93.5  17,5  16.5  77.8  2.0  — —  •  8. 8 —  <_  Represents only 1 or 2 p r i c k l e s i n t h i s body section, '* One specimen i n t h i s sample was i n a l l respects t y p i c a l l y "inland" i n i t s p r i c k l i n g , t Data enclosed by brackets indicate mean "apparent" p r i c k l i n g only. 1 These mean percentages have not been,weighted by m u l t i p l i c a t i o n body section.  of f r a c t i o n a l constants appropriate to each  102  Appendix Table 4.  Mean Relative P r i c k l i n g Intensity Index Values For A l l Populations of Cottus asper.  Population  Standard Length < 6 0 mm  Standard Length ^ 6 0 mm  Sample  Sample  Intensity  Size  Mean  2  1.0  2  2.0  eee o o  10 12  2,7 1.6  (2-3) (1-2)  Range  Size  Intensity Mean  Range  2.9 2.3 2.4 3.1 2,9 1.2 3.0 1.0 1.5 2.6 2.6 3.8 3.2 1.6 3.4 3.2 4 2.0 3  (2-3) (1-3) (1-3) (2-4) (2-4) (1-2)  Coastal 1 2 3 4 5 6 7 8 11 12 13 14 15 16 18 19 20-21 25a 26 28 30 31  :  5 14 17 28 10  3.0 2.8 3.8 3>6 2.7  (2-4) (2-4) (3-5) (1-5) (1-4)  9 6  1.9 3.8  (1-5) (3-5)  6 15  2.8 2.9  (2-3) (2-3)  Range  8 18 10 16 9 18 2 2 11 17 11 16 26 5 33 6 1* 24,  2 3 (1,0-3,8)  (1.0-3.8)  ooooo 0 0 0 6 0  (1-3) (1-5) (1-4) (2-5) (2-5) (1-3) (1-5) (2-4) 4 0 0 o 0  (1-3) (2-3)  Coastal Derivative 32 33 34 35 36 37 38 39 40 41 42 43  2 19 15 12 6 13 13 14 6 11 13 Range  3.3 3.0 3.0 3.2 2.1 1.9 3.0 2.9 2.6 (1.9-3.3)  (1-2) (2-3)  4  1.5 3.0 3.0 4.0  11 2  2,9 2.5  (2-3) (2-3)  3 3.0  ooo VW' ooooo  (2-4) 0 0 0 0 0  ooooo  (3.5)  0 0 0 0 0  (2-4) (1-3) (1-2) (2-4) (1-4)  (2.5-4.0)  103  Appendix Table 4 (cont'd).  Population No.  Mean Relative P r i c k l i n g Intensity Index Values For A l l Populations of Cottus asper.  Standard Length <c60 mm  Standard Length ^ 6 0 mm  Sample  Sample  Intensity  Intensity  Size  Mean  Range  Size  Mean  Range  15 3 15 6 12 11 15 13 20 16 14 14 24  3.7 3.7 3.0 3.7 3.3 5.0 3.1 3.8 4.3 3.9 3.2 3.6 4.5 3.6 5  U-4)  14  3.9 4.0 4 3.7 3.4 4.6 4.2 4.3 4.9 3.9 3,3 .4 4.7  (3-5)  Inland 44 45 46. 47 50 51 ,2 53 54 55 56 57 58 59 60 61 62 63 64 65 66-68 69-72 5  2 0  *  .1  4.5  (3-4)  U-4)  U-4) (2-4) 0 4 0 0 6  (3-4) (2-4) (4-5) (2-5) t2-4) (2-5) (3-5) (3-4) ooooo  2 11 2 23 Range  4.5 4.2 4.5 4.2 (3.0-5.0)  *  1* 15 10 31 25 15 21 20 1 3  *  1* 6  —mta  (4-5) (4-5) (4-5) (4-5)  7 21 9 18 7 10 7 6  (3-5)  20  (4-5) - — - -  —  9  4.3 4.3 4,0 4.5 4.9 4.2 4.0 3.6 (3.3-4,9)  •  s o t s  • • • 0  (3-4) (3-4) (4-5) (3-5) (4-5) (4-5) (3-4) (3-4) ooooo (4-5) ~ (4-5) (3-5) (3-5) (4-5) (4-5) (4-5) (3-5) (3-4)  Intergrade 73  8  3.8  Single specimens not used i n computing means  3.4  (2-4)  Appendix Table 5,. Population No.  Pectoral F i n Asymmetry i n Cottus asper, by Population.  Sample Size (Pairs)  Symmetrical No. % Pairs  Asymmetrical No, % Pairs  T o t a l Ray Count Greater Greater on L e f t on Right  Direction o f Asymmetry Within Population S i n i s t r a l Dextral Equal  Coastal ,1 2 3 4 5 6 7 8 11 12 13 14 15 16 18 19 20 21 23 25a 26 28 29,31 30  Totals  10 8 20 21 18 19 30 2 5 11 22 24 33 72 15 33 12  i  30 7 35 8 5 15  9 7 14 18 16 15 20 2 3 9 15 17 29 60 13 30 8 4 26 4 28 4 2 14  90.0 87.5 70.0 85.7 88.9 78.9 66.7 100,0 60.0 81,8 68.2 70.8 87.8 83.3 86.7 90,9 67.7 80.0 86„7 57.1 80.0 50.0 40.0 93.3  4 10 0 2 2 7 7 4 12 2 3 4 1 4 3 7 4 3 1  46.0  367  79.8  93  kS  l 6 3 2  10.0 12.5 30.0 14.3 11.1 21.1 33.3  1 5 3 1 2 9  -  —  40.0 18.2 31, 8 29.2 12.2 16.7 13.3 9.1 33.3 20.0 13.3 42.9 20.0 50.0 60.0 6.7  20.2  1 5 5 3 7  -  2 3 1 3 1 5 3 2 —  62  —  1 1  -  1 2 1  -  2 1 2 2 1 5 2 1 1  -  1 2 2 1 1 1 31  -  + +  +  -  _  _  . + -  -  -  + + +  -  + + +.. +• :  -  +  -  + +  + -  —  +  -  t  15(65.2)  -  -  + +  -  .+ -  -  -  -  -  —  5(21.7) a (13.1)  Appendix Table 5 (cont'd). Population No.  Sample Size (Pairs)  Pectoral F i n Asymmetry i n Cottus asper, by Population Symmetrical No. % Pairs  Asymmetrical No. % Pairs  T o t a l Ray Count Greater Greater on Left on Right  Direction o f Asymmetry Within Populations Sinistral Dextral Equal  Inland 26 12 16 21 22 42 40 27 41 37 27 15 30 18 16 21 19 19 8 22 11 30 28  44 45 46 47 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66-68 69-72 73 Totals  5  5  1  89.6 75.0 62.5 81.0 40.9 <76.2 85.0 74.1 73.2 78.4 63.0 46.7 73.3 50.0 68.8 71.4 100.0 73.7 87.5 77.3 % 63.6 86.6 .26 75.0 .'21 26 9 10 17 9 32 34 20 30 29 17 7 22 9 11 15 19 14 7 17  -.408  74.0  1  3 3 6 4 13 10 6 7 11 8 10 8 8 9 5 6 0 5 1 5 4 4 7  10.4 25,0 37.5 19.0 59.1 23.8 15.0 25.1 26.8 21.6 37.0 53.3 26.7 50.0 21.2 28.6 26.3 12.5 22.7 36.4 13.4 25.0  3 1 2 2 4 5  2  2  + +  143  26.0  87  56  16(72.7)  2 3 4  -  5 6 5 6  §  6 4 5 3 6  <:8  6  -  2 (4 8 4 1 1 5 2 '6 3 5 3 2 ;  :  r  •-  —  -  3 2  -  + + +  -  + + + + +  —  + +  -  +  +  -  + +  + +  -  -  •  -  -  +  -  —  -  -  +  —  —  -  -  5(22.7) 1(4.6)  Appendix Table 5 (cont'd). Population Ho.  Sample Size (Pairs)  Pectoral Fin Asymmetry in Cottus asper, by Population. . Symmetrical No. % Pairs  Asymmetrical No. % Pairs  Total Ray Count Greater Greater on Left on Right  Direction of Asymmetry Hi thin Population*; Sinistral Dextral Equal  Coastal Derivative 2 22 19 16 6 23 15 15 7 2 11 . w-  32 33 34 35 36 37 38 39 40 41 42 43.  Totals  Grand Totals  S 71.4 100.0 2 90.9 10 10 . 71.4  2 4 5 5 2 7 4 3 2 0 1 4  100.0 18.2 26.3 21.2 -33.0 30.'4 26. 7 20.0 28.6  0 18 14 11 4 16 11 12  81.8 73.7 68.8 66.7 69.6 73.3  80.O  2 4 2 3 1 4 3 3 2  3 2 1 3 1  -  —  _  9.1 28.6  1 2  -  +  2  -  -  + + -  +  +  -  + + + +  -  152  113  74.4  39  25.6  27  12  8(72.7)  1163  888  76.4  275  23.6  176  99  39(69.6)  -  +  -  -  -  —  -  -  _  _ -  +  1( 9.1) 2(18.2)  11(19.6) 6(10.7  Appendix Table 6. Pop. No.  No,Specs.  1.  10 8 20 21 18 19 30 2 20 10 6 11 22 25 33 72 10 15 33 12 6 30 5 5 17 35 5 8 5 15 6 24 19 16 6 24  <& •  3. 4. 5. 6. 7, 8. 9. 10. l i e  12. 13, 14. 15, 16 o 17, 18. 19, 20. 21. 22, 23. 24a. 25a S b. 26. 27. 28, 29.31. 30. 32. 33. 34. 35. 36, 37.  Summary o f M e r i s t i c Counts (Means and Ranges) f o r a l l Populations of Cottus asper,D  Dl 8,80 9.88 8.95 8.95 9,06 8.79 9.13 9.00 9,10 9,00 9.00 8.82 8,95 9,08 9,06 8.99 8,90 9.00 8.82 9,00 9,00 9.00 9,00 9.00 9.00 9,03 8.60 8,75 8.80 8.67 9.00 9.46 9.05 9,19  (8-9.) (9=11) (9-10) (8-10) (9=10) (8=9 ) (9-10) (9 ) (9=10) (9 ) (9 ) (8-9 ) (8=9 j (8=10) (8=10) (8=10) (8=9 ) (9 ) (8=10) (9 ) (9 ) (9 ) (9 7 (9 ) (9 ) (8-10) (8-9 ) (8-9 ) (8-10) (8=9 ) (8-10) (9-11) (9-10) (9-10)  --  9.42 (9-10)  20,70 20,50 19,85 20,38 20,50 19, Nt 20,60 20.50 20.70 21,00 20,83 20,27 20,73 20.32 20,76 20,53 20,80 20,67 20,27 20,42 20.83 20,47 20,60 20,80 20,14 20.23 20.00 19.50 20.00 19,73 22.17 20,70 20.21 20,25  ...  A  2  (20=22) (20-21) (19-21) (20-21) (20-21) (19=21) (20=21) (20-21) (20=22) (20-22) (20=21) (20=21) (20-22) (19=21) (20=21) (19=21) (20-21) (20=21) (18-21) (20=21) (20-21) (20=21) (20=21) (20-22) (20-21) (19-21) (20 ) (19-20) (20 ) (19-20) (22-23) (20-21) (20-21) (20-21)  <...  20.54 (20-22)  16.70 16,88 16,35 16,67 17,11 16,37 16,40 17,00 17,00 17.00 16,50 16,64 17,18 16,44 16,88 16,80 17,00 16,87 16,76 16,75 17,00 16,87 17,20 16,80 16,00 16,49 16,60 16.50 16,80 16,47 17,67 16,75 16.42 17,00  Pi  (16=18) (16-17) (16-17) (15-18) (16-18) (16-17) (15=17) (17 ) (16=18) (16=18) (16-17) (16=17) (16=18) (15=18) (16=18) (16-18) (16-18) (16=18) (16-18) (16=18) (16=18) (16=18) (17-18) (16=18) (15-17)^ (15=18) (16-17) (16-17) (16-17) (15-17) (17-18) (16-18) (15-17) (16-18)  ...  16.96 (16-18)  15,30 15,75 15,50 15,95 15.22 15,63 16,63 16,00 15.84 16,60 16,50 17,00 15,95 15,38 15,55 15,89 16,20 15,93 16,45 16,75 16.20 16,10 16,40 15,60 16,14 16,46 16,20 16,38 15,60 16,00 16,67 16,09 16,53 16,25 16,50 16,39  left  (15-16) (15-16) (15-16) (15=17) (15-16) (15-16) (16-17) (16 ) (15-17) (16-18) (16=17) (16=18) (15-17) (15-16) (15-16) (15-17) (16-17) (15-16) (16-17) (16-18) (16-17) (15-17) (16-17) (15-16) (15-17) (16-17) (16-17) (15-17) (15-16) (15-17) (16-17) (16-17) (16-17) (16-17) (16-17) (15-17)  Pj_ r i g h t 15,20 15,88 15,30 15,81 15,22 15,63 16,37 16,00  15=16) 15=16) 15=16) 15=16) 15=16) 15=16) 16=17) 16 )  16,83 17,00 15,82 15,24 15,48 15,86  16=17) 17 ) 14=16) 15=16) 15=16) 15=17)  16,07 16,42 16,58 16,00 16,00 16,20 15,60 16,29 16,37 16,00 16,12 15,40 16,07  16=17) 16=17) 16=18) 16 ) 14=17) 16-17)  15,91 16.58 16.19 16,50 16,38  15- 17) 16=18) 16-17) 16-17) 15-17)  lS=i&)  16=17) 15-17) 15-17) 15=17) 15=16) 15-17)  D  l  +  D  29,50 30,38 28,80 29,33 29,56 28,63 29.73 29,50 2 9, 80 30,00 29,83 29,09 29,68 29,40 29,82 29<,f2 2 9.70 29,67 29,09 29.42 29,83 2 9. ,45 29,60 29,80 29,14 29,26 28,60 28,25 28.67 28.40 31.17 30.17 29,26 29.44  2 (29-31) (2 9=32) (28=30) (28=30) (29=30) (28=30) (29=31) (29=30) (29-32) (29,31) (29-30) (28=30) (29=31) (28,31) (2 9=31) (28=31) (29=30) (29=30) (26=30) (29=30) (29=30) (29=30) (29=30) (29=31) (29=30) (28=30) (28=29) (27-29) (28=30) (27=29) (30=32) (29=31) (29-31) (29=31)  29,96 (29-32)  Appendix Table 6, Pop. No, 38, 39, 40, 41, 42, 43, 44, 45, 46, 47, 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64, 65. 66a. 66b. &V1 o 6*f.c  70?'= 7i|p  72|* 73>  Summary o f Meristic Counts (Means and Ranges) f o r a l l Populations of Cottus a s p e r ( c o n t ' d). p  No, Specs, 15 15 7 2 12 15 29 12 16 21 9  10-  22 42 40 28 41 37 27 15 30 20 16 21 13 24 9 22 3 3 3 ?-2 14 8 3 4 28  Pl left  9,13 (9-10) 9.27 (9-10)  --  9.50 9,17 9.61 9.03 9.00 9.25 9.10 9,22 8,80 8.52 8,98 9.00 9,00 8.98 8,95 8.89 8.80 8.87 9.06 8.69 8.81 9.08 9.00 8.89 8.68 8.00 8.00 8.00 8.00 8.21 8.00 8.00 8.00 8.82  (9-10) (9-10) (8-10) (9-10) (9 ) (9-10) (8-10) (9-10) (8-9)) (7-10) (8-10) (9 ) (8-10) (8-9 ) (8-9 ) (8-9 ) (8-9 ) (8-9 ) (8-10) (8-9 ) (8-10) (9-10)s? (8-10) (8-9 ) (8-9 ) (8 ) (8 ) (8 ) (8 ) (7-9 ) (8 ) (8 ) (8 - ) (8-10)  20.47 (20-21) 20.40 (20-21)  16,93 16.80 —_ 20.00 (20 ) 16.50 20,17 (20-21) 16,42 20,31 (20-21) 16,27 20,27 (19-21) 16,59 2;0,17 (20-21) 16,83 20,31 (20-21) 16,25 20,71 (20-22) 16,52 20.00 (19-21) 16,12 20,70 (20-22) 17,10 20,64 (16-23) 16,67 20.76 (20-21) 17,31 20.82 (20-22) . 17,02 20,82 (20-21) 16, t i 16,98 20.90 (19-22) 16,89 20,70 (19-22) 17,18 20.89 (20-22) 17,00 20.93 (20-22) 16,27 20.13 (19-22) 16,81 20.31 (19-21) 16,81 20.87 (2® £2 2) 17,24 20.69 (19-22) 17.15 20*69) (20-22) 16.58 20.29 (19-21) 17.00 20.56 (20-21) 17,18 20,23 (19-21) 16.67 19.67 (19-20) 17,00 20,00 (20 ) 16.33 19.67 (18-21) 17,50 20.00 (20 ) 16,64 (19 ) 19.00 17,00 19.12 (18-20) 16,67 (19-20) 19.33 16,50 18,75 (18-20) 16,54 19.93 (19-21)  (16-18) (16-18) (16-17) (15-18) (16-18) (15-17) (16-18) (15-17) (15-17) (16-17) (16-18) (15-17) (16-20) (16-18) (1-6-.13) (15-18)® (16-18) (16-18) (16-18) (15-17) (16-18) (15-18) (16-19) (16-18) (15-18) (16-18) (16-18) (16-17) (17 ) (16-17) (17-18) (16-18) (16-18) (16-17) (16-17) (15-18)  16,731 (16KL7) 16,27 (15-17) 17.43 (17-18) 17,50 (17-18) 16,17 (16-17) 16,67 (16-17) 16,86 (16-18) 16,75 (15-18) 16,50 (15-18) 16,29 (15-17) 16,90 17,14 16,93 16.60 16,93 16,44 16,43 16,30 16,67 16,73 16,67 16,44 16,29 16,00 16,25 1.6,00 16,14 16.00 16.00 15.67 16.00 15,64 16,50 16,33 16.00 16,21  (16-17) (13-18) (14-18) (16-18) (16-18) (15-18) (16-17) (15=17) (14-18) (16=18) (14-18) (16-17) (15-18) (15-17) (16-17) (15-17) (15-17) (15-17) (16 ) (14-17) (16 ) (15-16) (16-17) (16-17) (15-17) (15-17)  ?l right 16,67 16.07 17,14 17.50 16,00 16,64 16,83 16,50 16,38 16,45  16- 18) 15-17) 17- 18) 17-18) 16 ) 16- 17) 16-17/) 15-18) 15=18) 15=17)  17,50 16,86 16,50 .16,74 16,39 16.32 16,41 16,53 16,80 16,16 16,38 15,86 16,00 -16,21 15.89 16,18 15.67 16,33 16.00 15,00 15,50 16,25 16,33 16,00 16,11  17=18) 14=19) 16=17) 16-18) 15=17) 15=17) 16=17) 14=17) 16=18) 14=18) 15=17) 15=17) 15=17) 15=17) 15=16) 16=17) 15-16) 16- 17) 15- 17) 14-16) 14-16) 16- 17) 16-17) 15-17) 15-17)  Dl + 2 29,60 29-31) 29.67 29-31) D  29,50 29.34 29/92 29,31 29.18 29,56 29,81 29,22 29,50 29.14 29,74 29,83 29,82 29.88 29,65 29,78 29.73 29,00 29 .45 29,53 29.50 29,77 29,29 29,45 28.91 27,67 2 8.00 27,67 28,00 27,21 27,12 27,33 26,75 28,75 f  29-30) 29-31) 29-31) 28=30) 29-30) 29-31) 28=31) 29=30) 29=30) 28- 32) 2 9-32) 29- 31) 28=31) 28=31) 28=31) 28=31) 29=31) 27- 31) 28=31) 28- 31) 28=31) 2 9=31) 2 8-31) 28=30) 28-30) 27-28) 28 ) 26-28) 28 ) 26-28) 26- 28) 27-28) 26- 28) 27- 31)  Appendix Table 7, Locality  Monthly Mean Temperature i n °C f o r Selected Coastal and Inland Localities,,  Latitude  J  F  M  A  - 5,9  - 2,6  + 1,4  4,2 7,2  M  S  0  7,8  3,0 8,7  D  Annual Range  - 3,9  - 7,5  18,8  5,1  2,7  12,2  7.3  •>5.2  12,3  J  J  A  6,3  10,2  11,3  10,9  6,3  9,8  12,0  13.4  13,9  12,0  9,6  12,4  14,3  15,9  15,7  14,4  10.9  13,3  14,4  14,3  14,7  15.9  15,3  13.6  — .  ='=  N  Coastal Valdez  61° 07'  N .  - 7.1  Prince Rupert  54° 17'  N.  1.7  Victoria  48° 25'  No  3.6  San Francisco  37° 47'  No  Santa Barbara San Diego  10.0  ,2,9 5,7 11,0  11,8 32° 44'  N-S Range  N.  11.9 —  12,5  19,1  _=  12,3  13,4  14,8  16,2  17,8  19,4  19.3  17,4  14,8  li,5  18,4  - 8,9  - 0,4  8,0  19,0  —  —  :  10,9 —  5.9 7,3  21,7  20.9  18.6  15.6  13,2  9,7  11,8  13,1  15,6  -19,5  20.7  13,6  14,8  12,8  9,0  1.1  -13,7  -22,1  .36.8  9,9  4,8  - 2.7  - 7,5  24.8  2,7  -1,1  25.3  8,8  21,0  9,4  Inland Watson L o  60° 07'  -22.0  Prince George  53° 54'  - 9,9  6,6  - 1.4  + 4,6  9.9  13.4  14,9  13,7  Spokane  47° 37'  - 4.4  +0.4  3.8  8,7  13,1  16,3  20,9  20,0  15,4  9,2  Red  40° 09"  6,9  9,2  11.4  14,6  19,8  23,0  27,1  25,4  23,5  17,4  11.6  7,7  20.2  38° 35'  7.6  10.0  12,2  16,3  18,0  21,3  23.9  23,2  22,3  17,6  12,2  8,3  16,3  —  19.9  Bluff  Sacramento Fresno N-S Range  27,4  7,5 29.6  28.4  21.1  15,9  11,8  9,6. 12.6  -  —  12,6  14,5  16,5  —  25,9  *10 year-averages, 1941-1950 (World Weather Records, U„S, Weather Bur, Wash., D,C, 1959.1-1361)  30,4  —  

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