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Phylogenetic analysis of the cottid genus Artedius (Teleostei:Scorpaeniformes) Begle, Douglas P. 1984

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C. 1  PHYLOGENETIC ANALYSIS OF THE COTTID GENUS ARTEDIUS (TELEOSTEI:SCORPAENI FORMES) by DOUGLAS P. BEGLE B.Sc,  Stanford U n i v e r s i t y ,  1980  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Department of Zoology) We accept t h i s t h e s i s as conforming to the r e q u i r e d standard  THE UNIVERSITY OF BRITISH COLUMBIA February 1984 (c)  Douglas P. Begle, 1984  In  presenting  requirements  this for  thesis  an  B r i t i s h Columbia,  it  freely  for  available  that  or  understood  that  financial  by  for  his  that  reference  for  may b e  or  her  shall  Date  the  University  and  study.  I  copying of  granted  by  the  p u b l i c a t i o n of  not  be  this  It  this  allowed without  Columbia  make  further  head  representatives.  of  The U n i v e r s i t y o f B r i t i s h 1956 Main M a l l Vancouver, Canada V6T 1Y3  at  the  Library shall  permission.  Department  f u l f i l m e n t of  the  extensive  copying or  gain  degree  agree  purposes  department  for  I  permission  scholarly  partial  advanced  of  agree  in  thesis  of  my  is thesis my  written  i i  ABSTRACT  The  genus  revised.  Artedius  Artedius  lateralis,  1926)  Evidence  of the  presented. the  (Cottidae:Scorpaeniformes)  hankinsoni  is  synonymized  with  s i n c e the d e f i n i n g c h a r a c t e r s of A r t e d i u s  (Hubbs,  of  Girard  fail  to  separate  diphyletic  of  Artedius  hankinsoni  i t from A r t e d i u s  condition  is  lateralis.  Artedius  Girard  Based on the c h a r a c t e r a n a l y s i s i n t h i s study,  seven  nominal  s p e c i e s are hypothesized  is five  to represent a  monophyletic group (A. c o r a l l i n u s , A. l a t e r a l i s , A. h a r r i n g t o n i , A. f e n e s t r a l i s , and A. n o t o s p i l o t u s ) h e r e a f t e r Artedius  sensu  creaseri  and  strictu.  Artedius  synapomorphies  of  Ruscarius Jordan meanyi  and  Two  meanyi,  Artedius  and Starks  Artedius do  not  species,  share  sensu s t r i c t u ,  Evidence  Phylogenetic Artedius,  analysis  Ruscarius,  the  01igocottus,  Orthonopias,  Icelinus  cladograms.  These d i f f e r  The  of  and  of  the  Ruscarius  i s presented  f o r the  to be more c l o s e l y  i t i s to A r t e d i u s . adult  data  (including  Clinocottus,  Hemilepidotus)  Chitonotus,  yielded  three  only i n the placement of Chitonotus.  r e s u l t s from t h i s study are compared  (1947).  as  t h e r e f o r e the genus  monophyly of Ruscarius, which i s hypothesized and I c e l i n u s than  to  Artedius  any  i s r e s u r r e c t e d to i n c l u d e  Ruscar ius c r e a s e r i .  r e l a t e d to Chitonotus  referred  A l a r v a l data matrix  with  those  of  Bolin  f o r Washington's (1982) study i s  r e c o n s t r u c t e d and three l a r v a l cladograms are presented. F i n a l l y , an a n a l y s i s of taxonomic congruence using  the consensus technique  of Adams (1972).  is  attempted  Two t r e e s were  prepared, one showing the consensus showing  the  consensus  of  the  of  the  larval  adult  trees.  consensus t r e e was then compared with the a d u l t to  give  an o v e r a l l consensus t r e e .  recognizes Clinocottus Icelinus. the  removal  two  generic  and  groups:  Oliqocottus,  one  Ruscarius i n a clade  Artedius with  The  one  larval  consensus  tree  T h i s f i n a l consensus t r e e composed  of  Artedius,  and one composed of Ruscarius and  Thus both a d u l t and l a r v a l from  trees,  classifications  support  of c r e a s e r i and meanyi to the genus Icelinus  i n c l u d i n g A r t e d i u s , 01iqocottus  separate  and C l i n o c o t t u s .  from  the  clade  iv  TABLE OF CONTENTS  ABSTRACT  i i  LIST OF FIGURES  vii  ACKNOWLEDGEMENTS  ix  INTRODUCTION  1  HISTORICAL REVIEW  2  METHODS  9  Adult Specimens  9  L a r v a l Data  9  Meristics  10  Systematic Method  10  Comparison of C l a s s i f i c a t i o n s  18  Adults  18  Larvae  18  Taxonomic Congruence The Genus A r t e d i u s G i r a r d  19 1856  Taxonomic R e v i s i o n Artificial  key to A r t e d i u s  D i a g n o s i s of A r t e d i u s Artedius corallinus  21 21 21 22 24  Synonymy  24  Diagnosis  25  Specimens Examined  25  Artedius  fenestralis  26  Synonymy  26  Diagnosis  28  Specimens Examined  28  Artedius harringtoni  29  Synonymy  29  Diagnosis  31  Specimens Examined  31  Artedius l a t e r a l i s  32  Synonymy  32  Diagnosis  35  Specimens Examined  :  35  Artedius notospilotus  36  Synonymy  37  Diagnosis  38  Specimens Examined  39  The genus Ruscarius  Jordan and Starks  Taxonomic R e v i s i o n Artificial  key to Ruscarius  Ruscar ius d i a g n o s i s Ruscar ius c reaser i  1895  40 40 40 41 42  Synonymy  42  Diagnosis  43  Specimens Examined  43  Ruscar ius meanyi  44  Synonymy  44  Diagnosis  45  Specimens Examined  45  vi  PHYLOGENETIC List  ANALYSIS  47  of Adult Characters  47  L i s t of L a r v a l Characters  49  Adult Character  49 .  Analysis  RESULTS  82  Adult Trees  82  Monophyly and R e l a t i o n s h i p s of A r t e d i u s  82  Monophyly of Ruscar ius  83  A l t e r n a t i v e t r e e s - the r e l a t i o n s h i p s of Ruscarius L a r v a l Trees COMPARISON OF CLASSIFICATIONS  .. 84 86 88  Adults  88  Larvae  91  CONGRUENCE OF LARVAL AND ADULT CLASSIFICATIONS  96  SUGGESTIONS FOR FUTURE STUDIES  100  LITERATURE CITED  102  Appendix  115  1. Adult Data Matrix  Appendix 2. L a r v a l Data Matrix  116  vii  LIST OF FIGURES  Figure  1. A r t e d i u s c o r a l l i n u s  118  F i g u r e 2. A r t e d i u s f e n e s t r a l i s  120  F i g u r e 3. A r t e d i u s h a r r i n g t o n i  122  F i g u r e 4. A r t e d i u s l a t e r a l i s  124  F i g u r e 5. A r t e d i u s n o t o s p i l o t u s  126  F i g u r e 6. Ruscar ius c r e a s e r i  128  F i g u r e 7. Ruscar ius meanyi  130  F i g u r e 8. Orthonopias t r i a c i s  132  F i g u r e 9. Side view of s c a l e r i d g e (schematic)  134  Figure  10. Chin pigmentation p a t t e r n s  136  Figure  11. Top view of s c a l e r i d g e (schematic)  138  Figure  12. P t e r o t i c  140  Figure  13. Adult Wagner t r e e 1  142  Figure  14. Adult Wagner t r e e 2  144  Figure  15. Adult Wagner t r e e 3  146  Figure  16. Cladogram of A r t e d i u s sensu s t r i c t u  148  Figure  17. Cladogram of Ruscar ius  150  Figure  18. Adams consensus t r e e f o r a d u l t t r e e s  152  Figure  19. L a r v a l Wagner t r e e 1  154  F i g u r e 20. L a r v a l Wagner t r e e 2  156  F i g u r e 21. L a r v a l Wagner t r e e 3  158  F i g u r e 22. Adams consensus t r e e f o r l a r v a l t r e e s  160  F i g u r e 23. P h y l o g e n e t i c t r e e of B o l i n  162  Figure  24.  Wagner  flange  tree  calculated  (1947) in  the  absence  of  viii  Ruscarius meanyi  164  Figure  25. Dichotomous l a r v a l  Figure  26. Polytomous l a r v a l t r e e from Washington  Figure  27. Adams consensus t r e e of a d u l t and  trees  t r e e of Washington  (1982). ..166 (1982). .168  larval  Adams 170  ix  ACKNOWLEDGEMENTS I  wish  to thank the f o l l o w i n g f o r l o a n i n g specimens  t h e i r c a r e : Dr. (SIO), R.  Dr.  S.H.  Lavenberg  Pietsch  W.E.  Eschmeyer  Dr.  Weitzman (USNM), Dr.  (LACM), Dr.  (UW).  (CAS),  Mr.  D.E.  Robert  R.R.  McAllister  Carveth  R.  was  Rosenblatt  Miller  (NMC)  under  (UM), Dr.  and  Dr.  especially  T.  helpful  regarding the UBC f i s h c o l l e c t i o n and f r e e l y shared much of h i s e x t e n s i v e knowledge of C o t t i d s . This  study  was  provided p a r t i a l enough to provided  begun  financial  oversee  the  some f i n a n c i a l  under  Dr.  support.  completion support.  was l a r g e l y made p o s s i b l e by a  Dr. of  people  introduced  me  friendship  and  study.  deserve to  the  University  special  project  UBC.  also thesis  Fellowship  Dr.  systematics  at c r i t i c a l  Daniel and  Brooks  provided  times throughout the  Much of what I l e a r n e d from him I l e a r n e d o u t s i d e of the I cannot  thank  enough f o r h i s f r i e n d s h i p and support throughout my stay at Kate  Shaw  encouragement for  and  (1982-1984).  classroom i n innumerable impromptu d i s c u s s i o n s . him  McPhail was kind  Graduate  thanks.  phylogenetic  encouragement  J.D.  Wilimovsky who  The completion of t h i s  from the U n i v e r s i t y of B r i t i s h Columbia Two  N.J.  helping  evolutionary  also  provided  much-needed  friendship  throughout the course of t h i s study. me  understand  many  topics  in  Kate  I thank her  s y s t e m a t i c s and  theory and f o r uncounted hours spent j u s t  I a l s o thank her f o r the c o u n t l e s s hours of c r i t i c a l about t h e s i s  and  talking.  discussion  topics.  Shaw  and  Andrew  Simons  were  kind  enough to read  X  e a r l i e r d r a f t s of t h i s t h e s i s . Ms.  Maggie Hampong.  The cladograms were prepared  by  Most of a l l I thank my mother who p r o v i d e d  u n f l a g g i n g support throughout my time at U.B.C, without which I would not have begun t h i s e n t e r p r i s e , much l e s s f i n i s h e d i t . $3.21, $3.23T  $SIGNOFF  1  INTRODUCTION  The seven are  Cottidae  genera  a r e a S c o r p a e n i f o r m f a m i l y composed o f  (Nelson,  of  the  monophyletic  status  contributing  to  systematic  studies  taxonomic  poorly  i s the  A major f a c t o r  absence  Even i f t h e C o t t i d a e  our  present this  knowledge  study  has  r e v i s i o n of t h e genus A r t e d i u s , r e l a t i o n s h i p s of t h e s p e c i e s  and a d u l t  The Islands  of four  Baja  analyses natural  the  genera  goals:  is  (1)  a  within  Artedius, t o other  (3)  Cottid  data  from  stages.  California.  museum c o l l e c t i o n s a n d a l a r g e available  sound  (2) d e f i n i t i o n of t h e  of t h e r e l a t i o n s h i p of A r t e d i u s  genus A r t e d i u s G i r a r d to  of  is a  g e n e r a , and (4) a t e s t of taxonomic congruence u s i n g larval  and i t s  w i t h i n those genera a r e i n v a l i d .  incomplete,  examination  defined  The  t h a t a number o f g e n e r a a r e n o t m o n o p h y l e t i c  that  genealogical an  confusion  species.  t h a t many s p e c i e s  relatively  are  of the genera, i n c l u d i n g taxonomic  i ti s likely  Given  family  genera  California.  h a s n o t been d e m o n s t r a t e d .  this  the appropriate  taxon, and  F o r t y of t h e s i x t y - s e v e n  found between t h e A l e u t i a n I s l a n d s and Baja  taxonomic l i m i t s  of  1976).  sixty-  f o r study-.  i s d i s t r i b u t e d from t h e A l e u t i a n s The genus i s w e l l - r e p r e s e n t e d i n collection  of  larvae  is  also  2  HISTORICAL REVIEW  The  genus  accomodate two (previously  Artedius  was  s p e c i e s of  scaled  Artedius  Ayres  and Hemilepidotus  1854  Ayres'  In 1882, and  notospilotus  notospilotus  less  "represent  a  in  they  remark  distinct  In  that  the  variety."  v a r i e t y " as a d i s t i n c t  However,  due  the  large  cottid  683).  "northern  t r a n s f e r r e d to and  Calcilepidotus  lateralis  - though the  lateralis  the genus I c e l u s Kroyer,  other  (p.  to  l a t e r a l i s by G i r a r d i n  name).  scaly"  A. n o t o s p i l o t u s ,  In 1883  species their  of  t h e i r previous  were  d e s c r i p t i o n of  northern  specimens  they recognized  species, Artedius  number  i n the  which  this  fenestralis.  s p e c i e s that had  been  from A r t e d i u s s i n c e G i r a r d e r e c t e d the genus,  they d e s p a i r e d of s e p a r a t i n g A r t e d i u s from I c e l u s , and to  to  lateralis  and G i l b e r t p l a c e d both A r t e d i u s  family I c e l i d a e , along with or  (  Artedius  nebulosus G i r a r d 1856  unpublished  Jordan  Artedius  "more  cottids,  d e s c r i b e d as Scorpaenichthys  1854), and  l a t t e r was  e s t a b l i s h e d by G i r a r d in 1856  deferred  (1882a) d e c i s i o n p l a c i n g a l l A r t e d i u s s p e c i e s  in I c e l u s . In 1895 Puget  Jordan  sound.  Lockington  Its  1882,  and  Starks d e s c r i b e d Ruscarius  affinities  apparently  from which i t was  also based  removed on  Artedius  its  preopercular  rougher,  armature.  Jordan  f e n e s t r a l i s to a new more  l a y with  distinguished  back and weaker p r e o p e r c u l a r armature.  scaly  meanyi  by  from  Chitonotus a  scalier  and'Starks  (1895)  genus, A s t r o l y t e s ,  cranium  and  A r t e d i u s n o t o s p i l o t u s was  stronger subsequently  3  t r a n s f e r r e d to A s t r o l y t e s i n 1896 by Jordan and Evermann. In  1896  asperulus  Starks  described  two  new  species,  Artedius  and A x y r i a s h a r r i n q t o n i , both from Puget Sound.  Bean  and Weed (1920) l a t e r d e s c r i b e d a new genus and s p e c i e s from the same r e g i o n , P t e r y q i o c o t t u s macouni. the  male  of  Axyrias  This  harrinqtoni  they  Starks,  suspected  with  was  which i t was  synonymized by B o l i n i n 1944. Regan (1913), i n h i s treatment of fishes  with  preopercle, and  the  third  forming  Astrolytes  suborbital  the  Scleroparei  (those  extended backward onto the  a " s u b o r b i t a l s t a y " ) , placed  both  Artedius  i n the f a m i l y C o t t i d a e and p l a c e d the f a m i l y i n  the d i v i s i o n C o t t i f o r m e s .  No mention was made of Ruscar i u s , but  both I c e l u s and I c e l i n u s were a l s o i n c l u d e d i n the C o t t i d a e . Within  h i s series Cottiformes,  C o t t i d a e to those Astrolytes,  genera that  Axyrias,  lacked  Ruscarius,  were u n i t e d with Chitonotus,  Jordan  scales.  Ruscariops,  Orthonopias,  I c e l i n u s , and twenty-seven other  (1923) r e s t r i c t e d the Thus  Artedius,  and P t e r y g i o c o t t u s  Stelqistrum ,  Icelus,  s c a l e d genera and p l a c e d  i n the  Icelidae. No new A r t e d i u s - 1 i k e s p e c i e s were d e s c r i b e d u n t i l Hubbs  described  Artedius. considered  three  He p l a c e d each s p e c i e s P a r a r t e d i u s hankinsoni  Loma, C a l i f o r n i a ) and specimen, !a.  Allartedius  in  a  monotypic  was  species, related  Ruscar iops  r e l a t e d to genus.  He  (a s i n g l e specimen from Point corallinus  (also  from Point Lobos, Monterey County) c l o s e l y  The t h i r d  County),  s p e c i e s that were apparently  1926 when  creaseri  (from  a  single  related to San  Diego  to Ruscar ius meanyi of Jordan and Starks  4  (1895).  Hubbs favored  A. n o t o s p i l o t u s  monotypic genera and  from A s t r o l y t e s and  in 1926  placed  he  removed  i t i n t o a new  genus:  Parastrolytes. In  1940,  Artedius The  L.S.  and  Berg r e v e r t e d  i t s r e l a t i v e s and  f a m i l y Ice.l idae  he  to Regan's (1913) treatment  placed  reserved  them back in the  for  Icelus.  members of Jordan's I c e l i d a e were reassigned One  year  later,  in  1941,  Taranets  comprehensive treatment of the anatomy and Cottidae.  He  d i v i d e d the S c l e r o p a r e i  erected within  the C o t t i d a e  r e l a t i o n s h i p s of  the  i n t o twelve f a m i l i e s and  thirteen subfamilies.  bony  and  l a c k i n g an anal p a p i l l a  those  with  an  (Oligocottini).  The  Parastrolytes, Ruscariops  anal  were  common along In  possession the  lateral  addition,  body."  (p.4).  Ruscar i u s ,  f i n bases. the  plates  are  This  and  lacking  subfamily  those  with  (Artediini), bony  Artedius,  a subfamily,  plates  Allartedius,  Ruscarius  and  I c e l i n a e , along  T h i s he  justified  by  with their  rows of bony p l a t e s which occur one  at the base of the  usually  is  S t e l g i strum  between the  definition,  O l i g o c o t t i n a e along  dorsal  found in other  description  the area  By h i s own  subfamily  I eelinae.  l i n e , and  Ruscar iops,  scales covering  to  two  groups:  Orthonopias.  Stelgistrum. of  and  included  and  removed  I c e l u s , I c e l i n u s and  papilla  former  Axyr i a s  generic  His  large  and  Cottidae. his  two  (scales)  remaining  completed  O l i g o c o t t i n a e contained plates  Cottidae.  The  to the  of  hardly all  lateral  fin.  p a r t s of  accurate, possess  l i n e and  these three  "one  the  and  not  as  rows of dorsal  genera belong  with A r t e d i u s  the  in  i n the  5  The  last  A. d e l a c y i , They c i t e d  s p e c i e s to be assigned  d e s c r i b e d from Alaska by Hubbs and its  thicker  distinguishing  in  Except  for  (A. a s p e r u l u s ,  lips  from  from Alaska at that  and  snout  A r t e d i u s l a t e r a l i s which was  not known  Artedius  which  contained  three  species  A. d e l a c y i , and A. l a t e r a l i s ) , a l l the  remaining  remained  separate  Ruscariops,  until  Ruscariops  Artedius,  and  Axyr i a s  placed  Neither  nor  delacyi  A. d e l a c y i .  members  This  was  (1947) presented of  the  genus  and  its  included  in  a branching Artedius,  i s the only p h y l o g e n e t i c  a d u l t c h a r a c t e r s for A r t e d i u s  fenestralis  A. h a n k i n s o n i ) .  Ruscar ius meanyi  of the C a l i f o r n i a  r e t a i n e d as  Artedius  and  Bolin  Artedius  and  A. l a t e r a l i s  analysis.  He  (A. h a r r i n q t o n i )  (A. c o r a l l i n u s ,  Artedius  when  Parartedius,  (A. c r e a s e r i ) , A s t r o l y t e s , (A.  A. n o t o s p i l o t u s ) ,  taxonomic  1944,  Astrolytes,  in synonymy with A r t e d i u s f e n e s t r a l i s .  subgenera  1941.  features  P a r a s t r o l y t e s and Axyr i a s i n t o  and  S c h u l t z in  was  time.  gathered  asperulus  steeper  Artedius  as  genera were monotypic and Bolin  to the genus  the  diagram  including  dendrogram based on  close  relatives  (Fig.  23) . No until  further  1963,  specimens  when Rosenblatt  of  Ruscar ius meanyi were c o l l e c t e d  and W i l k i e c o l l e c t e d nine  in B r i t i s h Columbia.  Redescribing  to  on  Artedius  Girard  G i r a r d ' s genus. (Peden  It  and Wilson,  Sound (Moulton,  1977)  was  the  the s p e c i e s , they  basis  referred i t  of B o l i n ' s r e d e f i n i t i o n of  subsequently  collected  1976), B r i t i s h Columbia and C a l i f o r n i a  specimens  (Lea,  from  Alaska  (Peden, 1972), Puget 1974).  6  Since Hubbs and S c h u l t z ' to  separate  putative  Hubbard and  Reeder  (1941) d e f i n i n g c h a r a c t e r s  A. d e l a c y i  (1965)  and  from Quast  A r t e d i u s d e l a c y i was not d i s t i n c t Recent c l a s s i f i c a t i o n of Berg (1940). place Artedius  Nelson  Alaskan  A. l a t e r a l i s ,  (1968)  concluded  from A r t e d i u s  schemes have d i f f e r e d  litle  (1976) and Greenwood et a l .  while  that  lateralis. from t h a t  (1966) both  i n the C o t t i d a e along with 0 1 i g o c o t t u s  Clinocottus G i l l ,  failed  G i r a r d and  I c e l u s i s r e t a i n e d as the only member of  the I c e l i d a e . In  recent  years  morphology of l a r v a l studies  a  cottids  have  attempted  reconstructing  phylogeny.  phenetically larvae  derived  are  known.  Orthonopias,  great  deal  has to  of  accumulated,  use  these  Richardson  states  of  Her  character  and  "group  one"  data  recognized  includes  and 0 1 i g o c o t t u s .  spines,  different  s t a t e s do not p e r s i s t  which do p e r s i s t  only  two in six  Artedius,  I c e l u s and I c e l i n u s  Par i c e l i n u s ,  Triglops  These groupings were p r i m a r i l y based on shared  preopercular  pigmentation,  larval  on the  groups w i t h i n the c o t t i d genera f o r which  Clinocottus  Chitonotus.  but  (1981)  were i n c l u d e d i n "group two", along with and  information  in adults,  snout  diverticula in adults. were  used  and of  body  the  shape,  gut. These  Meristic characters, f o r the  purpose  of  ident i f i c a t i o n . Richardson's  (1981)  "group  one", i n c l u d i n g A r t e d i u s , was  c h a r a c t e r i z e d by a unique p r e o p e r c u l a r likelihood are  indeed  synapomorphic homologous.  spine  pattern,  in  f o r that group, i f the shared However,  no  attempt  was  all  states  made  to  7  establish  any  inter-generic  d i d attempt such Oligocottus. state  (one  an  She  analysis  1,3  spine, two  the  state.  sister  for  soft  an 1,2  state.  of  an  pelvic  have  clade  clade  and  would be the s i s t e r  A. meanyi)  plus  an group  Oligocottus  CIinocottus.  First,  the  genus  there has  species.  Artedius  been  some  U s u a l l y t h i s has  s p e c i e s now  been  some  to  Artedius  resolved.  The  primarily  last  because  on the  basis  (1947)  list  and  of of  in  recognizing  i s Artedius hankinsoni. later  as  to  c o n s t i t u t e a monophyletic group. group  problems have emerged.  been r e s o l v e d by c l o s e r study  in q u e s t i o n  question  three  difficulty  of t h i s s p e c i e s i s d i s c u s s e d has  and  A l l other A r t e d i u s  A. meanyi-Ieelinus  ( l e s s A. c r e a s e r i and  Within  only  CIinocottus  r a y s ) , A. meanyi should be removed to  A. meanyi-Ieelinus-A. c r e a s e r i  plus  (1982)  Under t h i s scheme, A. c r e a s e r i would then become  group  of A r t e d i u s  Artedius,  Washington  suggested t h a t , on the b a s i s of an 1,2  I c e l i n u s , which a l s o has an  relationships.  other  two  i n t h i s study.  and  The  the  status  Second, there  whether  or not these  T h i r d , the  relation  species of  that  s c a l e d C o t t i d genera has not been  problems  researchers  have  characters.  defining  l a r g e head, normal s t r u c t u r e of  remained  unsolved  have attempted to j u s t i f y  plesiomorphic characters  valid  the  Witness  Artedius: pelvic  fins  groups Bolin's  "comparatively and  by  the  unadvanced anus." These  characters,  taken  as  whole,  supposedly  serve  d i f f e r e n t i a t e the genus at hand from other genera d e f i n e d similar  fashion.  However,  no one  in  synapomorphic c h a r a c t e r  to a has  8  ever been proposed of A r t e d i u s • species 1920,  to serve as j u s t i f i c a t i o n  T h i s omission has r e s u l t e d  to and from I c e l u s ; Howe  actually  and  be  1978) that  in Artedius; by  synapomorphy  different  i s recognized  The  obvious  1982).  1965,  Washington  1966; Rosen,  (Nelson,  do,  taxa  (Farris,  1979a,b).  therefore timeless abstractions. in  natural  1981).  i n the c h a r a c t e r s  by  1975, 1979,  1981,  synapomorphies  at  Plesiomorphic c h a r a c t e r s  they  inevitably  1982; Rosen  1974).  lead  to  Such taxa are c l a s s e s and  Such taxa  cannot  participate  processes and should be avoided by those  to d i s c o v e r the p a t t e r n of n a t u r a l 1981,  no  i f the d e l i n e a t i o n of n a t u r a l taxa i s our goal  1971, 1972, 1973, 1979), as  paraphyletic  (Wiley,  1974; Wiley,  A n a t u r a l group must be supported  not  When  groupings and schemes of  justification  the a p p r o p r i a t e l e v e l of u n i v e r s a l i t y . will  (1982).  i s t o t e s t the hypothesis of monophyly  of A r t e d i u s by s e a r c h i n g f o r (Hennig,  should  (3) the r e f e r r a l of  o b j e c t i v e l y be defended  solution  (Bean and Weed,  Orthonopias  and  Ruscarius meanyi to I c e l i n u s  r e l a t i o n s h i p cannot  i n : (1) the t r a n s f e r of  (2) the suggestions  Richardson,  included  f o r the monophyly  order  (Wiley,  attempting  1977, 1979,  9  METHODS  Adult  Specimens  All  specimens  or 50%) or ethanol prepared  using  were preserved (70%).  the  i n e i t h e r isopropanol  C l e a r e d and  method  of  stained  Dingerkus  (37.5%  specimens  and  Uhler  (1977).  C l e a r e d and s t a i n e d specimens were s t o r e d i n 100% g l y c e r i n thymol and  added  to  r e t a r d decomposition.  with  a  Nikon  with  Photographs of c l e a r e d  s t a i n e d f i s h were made through a Bausch and Lomb  microscope,  were  dissecting  F 35 mm camera and Nikon microscope  adapter. I n s t i t u t i o n s p r o v i d i n g specimens were: of  Sciences  California  Academy  (CAS and SU), Los Angeles County Museum of N a t u r a l  History  (LACM),  Division  (SIO), U n i v e r s i t y of B r i t i s h Columbia F i s h Museum (BC),  National  Museum  of Zoology (UW),  (UMMZ),  Scripps  Institution  of  Oceanography  Fish  of Canada (NMC), U n i v e r s i t y of Michigan Museum University  of  Washington  Fish  Collection  and the United S t a t e s N a t i o n a l Museum (USNM).  L a r v a l Data  Information  on l a r v a l morphology was taken from Washington  10  (1982).  L a r v a l specimens  were  not  examined  in  the  present  study.  Meristics  For elements  both  preserved  were counted.  and c l e a r e d and s t a i n e d specimens a l l  The one e x c e p t i o n  was  the  caudal  fin  count which i n c l u d e d only p r i n c i p a l r a y s .  Systematic Method  "If  we  find  facilitated another, quoted  that  by one  the ascertainment of the order of nature i s terminology or one  set of symbols rather than  i t i s our c l e a r duty to use the former."  i n Jordan and K e l l o g g ,  method  which  evolution.  Phylogeny  are i n business genealogical  b i o l o g y (sensu Nelson, best  to  1970)  (Hennig,  1966)  we must choose  that  a l l o w s us to i n f e r the h i s t o r i c a l course of i s the h i s t o r y of l i f e and that discover.  relationships  primary problem  Huxley,  1907.  In c o n s t r u c t i n g a general r e f e r e n c e system for comparative  T.H.  is justifying  This  of  involves  natural  taxa.  i s what we  specifying  the  O b v i o u s l y , the  the n a t u r a l s t a t u s of  the  groups  11  at  hand.  As  Wiley  (1979)  s u f f i c i e n t c r i t e r i o n of Characters,  natural  the  status  only is  necessary  unique  which are our only d i r e c t l y observable  only  secondary  If we  are  to  principles  link  (sensu  characters Hempel,  of  a  genealogy  1965)  their  group's  to  genealogy.  relationships.  we  need  as  necessary  and  n a t u r a l s t a t u s : ontogeny and  c h a r a c t e r s . remained  throughout ontogeny and  only  necessary  but  s t a t u s of a group. of ontogenesis move  from  also  that  Since they can  which we  the  we  Put  concept  need  If  all  through  constitute  evidence  not  for the n a t u r a l  during  bridge  the  course  principles  to  ( c h a r a c t e r s ) to that which  simply,  of  1978).  would  be modified  observe  (genealogy).  from  they  sufficient  and phylogenesis  wish to i n f e r divorced  descent,  (Wiley,  two  character  descent  of p h y l o g e n e t i c  bridge  sufficient  modification during phylogenetic  the course  are  since characters exhibit  use  unchanged  and  evidence,  r e f l e c t i o n s of unique h i s t o r i c a l  phenomena p r e c l u d i n g criteria  noted,  characters  evolutionary  cannot  homology  we be  (Wiley,  1981). Our  method  novelties (Hennig,  should  which 1966,  serve  Wiley,  involve to  1979,  a  search  define 1981).  for  evolutionary  n a t u r a l (monophyletic) taxa Such c h a r a c t e r s  are  at the l e v e l of a n a l y s i s under study,  termed  apomorphies  if,  hypothesized  to be the d e r i v e d s t a t e of an a n c e s t r a l , r e l a t i v e l y  g e n e r a l i z e d homologue.  An apomorphic c h a r a c t e r  one  an autapomorphy.  taxon  is  termed  termed synapomorphies.  Only  in  are  only  Shared apomorphies are  synapomorphies  d e l i m i t n a t u r a l , monophyletic groups.  found  they  may  be  used  to  12  The  relatively  generalized  termed a plesiomorphy symplesiomorphy  state  of  that  and when shared by two  (Hennig  1966).  or  homologue i s more  monophyletic  Thus at some l e v e l of u n i v e r s a l i t y a l l c h a r a c t e r  are apomorphic, d e l i m i t i n g a n a t u r a l taxon. to  diagnose  a  N o t i c e that a l l c h a r a c t e r s when  they a r e e v o l u t i o n a r y n o v e l t i e s , d e l i m i t a n a t u r a l , group.  taxa,  They cannot  states be used  other taxa at a l e v e l of u n i v e r s a l i t y at which they  are not apomorphic  (Hennig,  apomorphy and plesiomorphy  1966, Wiley,  1979, 1981).  Note that  are r e l a t i v e terms and are meaningful  only i f the l e v e l of u n i v e r s a l i t y being c o n s i d e r e d  is  strictly  specified. The c r i t i c a l p a r t of any such a n a l y s i s i s the determination of  hypothesized  character.  plesiomorphic  Hennig  holomorphological  g e o l o g i c a l precedence  their  the  first  and  ontogenetic  methods:  chorological  (out-group  comparison),  precedence.  Of  these  and t h i r d have g e n e r a l l y been d i s c a r d e d due to  r e l i a n c e on a p r i o r i  character  apomorphic s t a t e s f o r each  (1966) d i s c u s s e s four  (biogeographical),  four,  and  change  in  assumptions  time and space.  about  the  pattern  of  These assumptions may or  may not be j u s t i f i e d , depending on the p a r t i c u l a r  circumstances.  Because  analyses  of  this,  outgroup  preferred.  In  outgroup  the outgroup  are judged  and  ontogenetic  a n a l y s i s , c h a r a c t e r s t a t e s shared  to be p l e s i o m o r p h i c at that l e v e l ,  the a l t e r n a t i v e s t a t e i s hypothesized Nelson's  (1978)  from von Baer's states  give  to  be  apomorphic.  are with while In  f o r m u l a t i o n , the ontogenetic c r i t e r i o n f o l l o w s law which  rise  to  states  that  relatively  generalized  s p e c i a l i z e d ones as ontogeny progresses.  13  T h i s method has of  two  character  effectively character  immediate drawbacks:  states  refute  von  analysis  in p r e s s ) .  Nelson  ontogenetic  and  instances  Baer's  suspect (1978)  analysis  comparison, but Wiley  to  in  proceed  outgroups.  Kluge,  of  in cases of  1984  f u r t h e r confound c h a r a c t e r  derived characters  appear p r i m i t i v e , again  outgroup  attained  pointed  paedomorphosis can  that  non-terminal  only be  (1982)  attendant  formulation,  independently  a n a l y s i s can  Fink  the  in p r e s s ;  his  (1980) noted that  addition  de-differentiation  making  (Brooks, 1984  a d d i t i o n meaningful c h a r a c t e r resort  of  "law"  claimed,  can  non-terminal  by  out  that  a n a l y s i s by making  n e c e s s i t a t i n g the  use  of outgroups as a check. P o l a r i z i n g characters more  apomorphic  states  These m u l t i s t a t e establishing  becomes more d i f f i c u l t exist  characters transformation  distribution  on  a  a  given  present  d i s t i n g u i s h e d four types of their  at  two  or  l e v e l of a n a l y s i s .  special  series.  multistate  when  difficulties Mickevich  characters  previously-derived  (1983)  based  cladogram.  in  upon These  are: 1. (a  Additive characters,  parsimony  possible.  leads best  technique - see F a r r i s 1974  These allow 2.  to  that  one  in  state  characters, was  state tree.  which  optimization  for transformation  r e s o l u t i o n of subsections  Convergent  optimization  for a d e s c r i p t i o n ) i s  of a c h a r a c t e r  characters,  multiple p o s s i b i l i t i e s  these allow  likely  construction  Non-additive  3.  f o r which F a r r i s  derived  in  series.  At  of a t r e e . which  it  is  equally  from d i f f e r e n t c o n d i t i o n s .  14  These are of minimal use 4.  D i s j o i n t c h a r a c t e r s , in which no t r a n s f o r m a t i o n  be hypothesized although  in the r e s o l u t i o n of t r e e s .  to  link  whole  transformation  subsets  series  may  of  character  the  allow s e c t i o n s of the t r e e to  be  These may  resolved,  r e l a t i o n s h i p s between these  the  states,  be e s t a b l i s h e d w i t h i n  i n d i v i d u a l subsets. but  can  s e c t i o n s remain  ambiguous. Once c h a r a c t e r a n a l y s i s i s constructed, l i n k i n g s i s t e r immediate  common  taxa  ancestor)  complete  (non-homologous  cladogram  (those hypothesized  same  similarity).  best  Parsimony  considerations  i t i s the l e a s t force  us  to  to  a  preference  due  to  In such an event,  the  characters  (the  It  refuted  explanation.  e v o l u t i o n a r y taxonomy of Mayr (1942, 1969) to  satisfy.  and adaptive divergence (phylogeny) from a use  of  and  "classical"  Simpson  (1945,  similarity  r e s u l t s from attempts to i n f e r a p a t t e r n  particular  from apomorphic information and, at  that  T h e i r r e l i a n c e on o v e r a l l  overall similarity  applied s t r i c t l y  This  set.  i s p r e c i s e l y t h i s parsimony requirement  fails  hypothesis.  for the t r e e with the minimum p o s s i b l e  number of steps for a p a r t i c u l a r data  The  Rarely  minimize the number of  c o n f l i c t s , each of which r e q u i r e s an ad hoc  1961)  an  number of steps or c h a r a c t e r s t a t e changes) i s chosen as  r e p r e s e n t i n g the data;  leads  be  to share  relationships  cladogram with the l e a s t number of c o n f l i c t i n g fewest  may  by means of synapomorphies.  do a l l the c h a r a c t e r s i n d i c a t e the homoplasy  a  the  level  process fails  (Darwinian to separate  evolution). plesiomorphic  as a r e s u l t , c h a r a c t e r s are at  which  they  constitute  not a  15  critical  test  of  the  notion of process was pattern  was  hypothesis  at  the mold i n t o  fitted,  resulting  hand (Wiley  which  the  We  i n p a r a p h y l e t i c groups  nature,  If we  are seeking to  which  discover  on  (grades) attempts  r e c o n s t r u c t the g e n e a l o g i c a l h i s t o r y of monophyletic must then ask:  The  information  which are not d e f e n s i b l e c o n s t r u c t s i n a system to  1981).  the  groups.  order  in  i s t h i s the a p p r o p r i a t e methodology?  T h i s p a t t e r n / p r o c e s s dichotomy i n s y s t e m a t i c s i s r e f l e c t i v e of  a  h i g h e r - l e v e l s t r u c t u r e / f u n c t i o n controversy  and C r a c r a f t ,  1981).  understanding examination thought  that  arises  not; though h i s he  properties  is,  discussion  f i n d s an and  with  process  an  centers  on  the  considered  are  to  understanding  a l l , it  (functions) factors  such  of  which  explanatory u n r e l a t e d to  a  an aims group  produced  be of  studied them  Churchland  prior (1982)  study  of  as  approach to be f a u l t y at  analyzing  of them.  the  discussed  properties  (structures,  it  reasons.  attributes  r e f e r e n c e to the may  which may and  the  "top-down".  f o r two  common  e n t i t i e s without Secondly,  emergent  contingent upon a  (1982)  e f f o r t s on p e r c e i v e d groupings the  an  isomorphism with n a t u r a l science and i n  structural analysis.  F i r s t of  can  Churchland  f a i l i n g s of a f u n c t i o n a l i s t approach which he l a b e l s He  Eldredge  of a p a r t i c u l a r process be a t t a i n e d without  function  priori  question  of the p a r t s ( p a t t e r n s ) involved?  cognition, both,  The  (see  by  focus in fact  our be  extension,  genealogy) of t h e i r members.  "In  the  event,  and  as  in our own  h i s t o r y , such  inquiry  1  (bottom-up) deepens, and changes, our it  i s we  denied  central  stipulative issue,  ignorance."  a  threaten  to  1982,  functionalist  be used  process,  we  (Churchland,  Within cannot  given  for c r i t i c a l  a  priori  are passed  with  make  a  paradigm,  tests  apomorphic.  particular  of  competing  process  of  (Wiley, 1981  Notions used  of  to  our  own  hypotheses  of  the p r o c e s s - l e v e l  change,  p.  only  that  that c h a r a c t e r s 78).  Characters  level  at  functional  which  efficiency  group.  be  known  as  phenetics  phylogeny,  t h i s amalgam of numerical trap  evolutionary phylogeny. statistical  (grouping taxonomy,  pretend  by  does  Since phenetic techniques methods  never  intended  and  today  Sokal,  (usually)  to  techniques has  overall  which  systematics  (Sneath  s i n c e i t does  same  not  in  Ironically,  the  of  to j u s t i f y the n a t u r a l s t a t u s of a  Another e x i s t i n g method in use come  a  systematic i n f o r m a t i o n  m o d i f i c a t i o n ) occurs and  (adaptiveness) are never  of  C l a d i s t i c a n a l y s i s , on the other hand,  on, m o d i f i e d or not  are  initial  status  prison  are used to d e l i m i t n a t u r a l groups only at the they  is  italics his).  does not assume any p a r t i c u l a r (descent  inquiry  of the domain of phenomena  s i n c e these p a t t e r n s a l r e a d y r e f l e c t  b i a s of the s y s t e m a t i s t .  change  i f such  i f our p r i m i t i v e  the  functional specification then  But  s i g n i f i c a n c e , and  conceptions are e x p l i c i t l y  at  i n i t i a l conception of what  should be t r y i n g to s i m u l a t e . any  6  claim arose for  to  from use  1973). estimate  fallen  similarity)  has  as  into has  reconstruct pre-existing  in  biological  17  s y s t e m a t i c s , i t i s h a r d l y s u r p r i s i n g that they f a i l the  separate  d e r i v e d from the g e n e r a l i z e d components in t h e i r measures of  overall the  to  similarity.  If the d e r i v e d component i s not separated,  genealogy of l i f e  1977).  Only  will  escape  largely  unnoticed  (Farris  i f there i s no homoplasy and homogeneous r a t e s of  change during any given time p e r i o d w i l l phenetic a n a l y s i s the  same c l a s s i f i c a t i o n  analysis  on  the  phylogeny even  a  change.  The  Wagner  program  (1983) was  •in  and  i s described  used  the  homoplasy  Cladistic  for estimating  and  PHYSYS package  differential  of F a r r i s  It i s  entirely in F a r r i s  phylogenetic (1970) and  are  invariably  i n t h i s study i n two  the  of Washington  (1982).  generate a consensus t r e e taxonomic  congruence.  in  Farris,  same.  These  The  i n s t a n c e s : to check the  cladogram d e r i v e d by hand and to re-analyze the re-coded data  and  In any event, the r e s u l t s of a Wagner  a n a l y s i s and hand computation was  of  in d e t a i l  Kluge and Eckardt (1970).  program  analysis.  used to analyze the data matrix obtained by  Hennigian c h a r a c t e r a n a l y s i s . nature  cladistic  other hand, i s a powerful t o o l  i n the presence  r a t e s of  Mickevich  as  yield  larval  two r e s u l t s were then used to  (sensu Adams, 1972)  i n an a n a l y s i s  of  18  Comparison of C l a s s i f i c a t i o n s  Adults A  phylogenetic  tree  a n a l y s i s of the C o t t i d a e .  was Two  extracted  from  Bolin's  measures were employed to  the t r e e ' s v a l i d i t y as an h y p o t h e s i s of phylogeny. character  data  a  Secondly,  classification  to  account  with  a  data  matrix  from  Manhattan  (where  distance  matrix  taxa i s the number of  taxa  for  divided  by  the  the  calculating  the  cophenetic  t h i s s p e c i e s was present  compare not  converted to  state  a  differences  number of c h a r a c t e r s ) .  correlation meanyi  d e l e t e d f o r the purposes  data  the Manhattan d i s t a n c e  character  Since B o l i n d i d not i n c l u d e R u s c a r i u s  (1947)  the  Wagner t r e e and B o l i n ' s t r e e were assessed f o r goodness by  the  which i t was  from t h i s study was  between  Firstly,  s i n c e i t i s not p o s s i b l e to d i r e c t l y  c o n s t r u c t e d , the data matrix  between two  assess  from t h i s study were mapped onto B o l i n ' s  t r e e and the number of steps needed counted.  (1947)  Both of  coefficient in  his  of comparison  my  fit (R).  analysis from  the  it  was  analysis.  Larvae  Before  a  congruence  necessary to ensure  study  could  be  attempted  that the best t r e e f o r the l a r v a l  data  had  19  been  calculated.  difficult one  had  s i n c e Washington  desideratum  tree  and  the  with  two  trees  matrix  and  A Wagner  trees  and from  which  was  i m p l i e d polytomous t r e e which r e s u l t e d from  were  then  ( i . e . by counting  Manhattan d i s t a n c e  derived  the dichotomous cladogram  c o l l a p s i n g a l l of the unsupported branches to  trees  initially  from the body of the t e x t .  compared  Washington's (1982) study:  These  was  then performed on the r e c o n s t r u c t e d data matrix  resulting  presented  this  (1982) p u b l i s h e d no data  to be r e c o n s t r u c t e d  a n a l y s i s was the  Satisfying  compared  the  node  in the same way  the number of steps and  below.  as the a d u l t  by  fit  to  the  matrix).  Taxonomic Congruence If  there  is  congruence w i l l sets out data  only  one  genealogy  of  n e c e s s a r i l y be a consequence of any  logically  sources  truly  may  to d i s c o v e r that  allow  history,  life,  then  method which  though  different  r e c o n s t r u c t i o n of that h i s t o r y to g r e a t e r  or l e s s e r degrees. Taxonomic provides of  congruence,  as  defined  a measure of the degree to which  by  Mickevich  the  (1978),  "classifications  the organisms remain s t a b l e as v a r i o u s l i n e s of evidence  considered". method  and  This s t a b i l i t y  i s o f t e n c i t e d as being  another competing a l g o r i t h m Mickevich  1978,  classification information  i s a consequence of  lies  should i  \  1980). in never  The hypothesis be  classificatory  the s o l e property  (Farris,  1971,  importance testing:  are  1982; of new  of one  Fink a  or  1979, stable  sources  of  i n t e r p r e t e d as r e f u t i n g previous  20  hypotheses actually  of  relationship  contradictory  These d i f f e r e n t different different  (Farris  sources may  life  unless  stages,  the  1971;  new  Mickevich  be of the same  such  as  larvae  type  c l a i m i n g to  provide  comparative  evolutionary  the  best  general  biology as  well  as  congruence  tests  there i s no way  only  instead  portrays  observations"  (Mickevich  the  system  for  1978), one  that  (Fink,  1979;  method  for  Eckhardt,  1982; and  Miyamoto,  1981;  1972;  a n a l y s i s of a d u l t s .  order, s u i t e of  To  date,  systematics provides the most  et  Barbour, 1981;  1981;  Mundinger,  larval  natural  such t e s t s of congruence.  Baverstock  and  Farris,  Polhemus, 1980).  determining  f i n d i n g maximum congruence (Andersen,  Jensen  Mickevich  of  i d i o s y n c r a c i e s of one  these have shown that p h y l o g e n e t i c  the  morphology.  1980).  There have been a few  using  and  possible  or  and  from  a d u l t s , or from  (Mickevich,  represents a r e p e a t a b l e ,  Smith,  data  reference  whether c l a s s i f i c a t i o n  B a i r d and  1980).  1980).  "Without  stable  is  i s a l s o a r e q u i s i t e property of methods  represents a l l characters Mickevich,  1978, of  and  types of c h a r a c t e r s , such as behavior  Taxonomic s t a b i l i t y  information  Mickevich 1979;  al.,  1979;  Mickevich, and  Hood 1978,  data  of  Washington  (1982) and  1976;  1981;  T h i s study attempts a congruence the  and 1980;  Johnson,  Schuh and F a r r i s ,  1979;  Shuh  analysis present  21  THE GENUS ARTEDIUS GIRARD 1856  Taxonomic R e v i s i o n  A r t e d i u s G i r a r d , 1856, p.  134.  (genotype by subsequent d e s i g n a t i o n of Jordan and Evermann 1896 Scorpaenichthys  lateralis  A s t r o l y t e s Jordan and S t a r k s , 1895 Axyr i a s S t a r k s , 1896 P t e r y g i o c o t t u s Bean and Weed, 1920 A l l a r t e d i u s Hubbs, 1926 P a r a r t e d i u s Hubbs, 1926 P a r a s t r o l y t e s Hubbs, 1926  Artificial  1a.  key to A r t e d i u s  S c a l e s present 2a.  on the o c c i p u t  2.  P r e o r b i t a l c i r r u s present Artedius harringtoni.  2b.  No p r e o r b i t a l  cirrus  3.  Girard)  22  3a.  Scales  extend under a n t e r i o r o r b i t Artedius  3b.  No s c a l e s under a n t e r i o r o r b i t Artedius  1b.  fenestralis.  notospilotus.  No s c a l e s on the o c c i p u t 4a.  Cirri  present  4.  above upper l i p  Artedius c o r a l l i n u s .  4b.  Diagnosis  of  No c i r r i  above upper l i p  Artedius  lateralis.  Artedius  R e c a l l that B o l i n c h a r a c t e r i z e d attempt at d i a g n o s i s head,  (the  only  on record) as f o l l o w s : "comparatively  such large  normal s t r u c t u r e of the p e l v i c f i n s and by the unadvanced  anus."  (1947, p.  equips  one  description. diagnostic alone.  Artedius  to  161) Even  slight  f i t literally In  this  characters  light,  familiarity  with  Cottids  dozens of genera i n t o the above the  apomorphic  following for  Artedius  is  a  list  of  sensu s t r i c t o  23  3.  Scale ridge c u r v i n g to become p a r a l l e l with b a s a l p l a t e .  4.  G e n e r a l l y darker  surface  interrupted  background by  pigmentation  of  c i r c u l a r areas of l i g h t e r  (same as v e n t r a l body s u r f a c e , not white spots of These  lighter  areas. area  pigmentation  of  anal  closer  background  ventral  f i n . Margins  pigmentation  producing each  Orthonopias).  a  circle  of  i n t e r r u p t i n g darker  head,  scalloped of  light  extending  blotches  of  pigmentation  backwards  onto  of p t e r o t i c  long,  membrane.  27.  P o s t c l e i t h r a absent.  35.  Scale r i d g e a small s e m i c i r c l e . Triangular  extending  pigmentation  to v e n t r a l edge of dark  specimens, a p a t t e r n of c i r c u l a r  surface  branchiostegal  45.  of  body  c r i s p and w e l l - d e f i n e d .  In preserved  light  larger  L a r g e s t , at margin, incomplete, above  12.  circles  lateral  at l e a s t  flange  at  posterior  edge  to p o s t e r i o r edge of cranium.  24  Artedius  corallinus  Artedius c o r a l l i n u s  (Hubbs 1926) ( F i g .  1)  Range: Orcas Island  (Washington) t o Baja C a l i f o r n i a .  H a b i t a t : Rocky i n t e r t i d a l areas t o 70 f t .  Synonymy  A l l a r t e d i u s c o r a l l i n u s Hubbs, 1926, p.  8; Point Lobos,  Monterey County, C a l i f o r n i a . Artedius c o r a l l i n u s Bolin, first  Baja  description, p.  record);  126, f i g . p.  12  1944, p.  126 ( i n key, a l s o 128 (name o n l y ) ;  (description,  53, f i g .  distribution);  checklist);  20 ( i n key,  Eschmeyer,  Herald  and  Fitch  and  Howe and Richardson, 1978,  ( i n key, plus d i a g n o s i s , synonymy, m e r i s t i c  Hubbs, F o l l e t t and Dempster, 1979, p.  18.  63  d i s t r i b u t i o n , r e l a t i o n s h i p s ) ; M i l l e r and Lea, 1972,  Lavenberg, 1975, p. p.  Bolin,  1937, p.  18 (name  variation);  only-California  Hamman, 1983, p.  161, p i .  25  Diagnosi s  20.  Cardiform t e e t h on d e n t a r i e s ,  premaxillae,  palatines  and  vomer. 26.  Two  t o s i x t e e n simple paddle-shaped f l a t c i r r i  upper l i p along  i t s length.  maxilla  forward,  Usually  two  not  small  These l o c a t e d anywhere  just  above  from end of  d i s t r i b u t e d evenly on each s i d e of head.  cirri  on  either  side  of  premaxillary  symphysi s. 22.  43-46  rows of s c a l e s along body, extending from o r i g i n of  first  dorsal  f i n to base of u l t i m a t e or penultimate d o r s a l  soft  ray.  Spec imens Examined UMMZ  14168  1  (holotype)  Mexico:Baja; CAS uncat.  Monterey,  Pt.  1 CA:Monterey; CAS  Verdes; CAS 28838 1 CA:Mendocino; CAS uncat.  Lobos;  CAS 19453 1  W51-74  1 CA:Palos  1 Mexico:Baja; CAS  39853 1 CA:Monterey; CAS Acc.#1972:23 1 CA:Monterey; CAS 48958 2 CA:Orange  Co.;  SU  29536  1  CA:Monterey; SU 48970 7 CA:Orange  County; SU 35365 1 CA:Monterey; SU 40881 W53-395  Lobos;  LACM  1 Mexico:Coronado I s l a n d ; LACM 1989 2 Mexico:San M a r t i n  I s l a n d ; LACM W70-16 48 CA:Monterey; LACM Co.;  1 CA:Pt.  LACM  9421-6  W65-26  13  CA:Ventura  9 Mexico:San N i c o l a s I s l a n d ; LACM 31301-1 34  CA:Diablo Cove; LACMW67-152 2 Mexico:Punta Banda; LACM 1 CA:Humboldt Co.  31937-8  26  Artedius  Artedius fenestralis  fenestralis  (Jordan and G i l b e r t  Range: A l e u t i a n I s l a n d s to southern H a b i t a t : I n t e r t i d a l to 180  1882) ( F i g .  2)  California.  feet.  Synonymy  A r t e d i u s n o t o s p i l o t u s Jordan and Jouy, 1882, p. 27416); Bean, 1882a, p.  250; 1882b, p.  471  (not of  I c e l u s n o t o s p i l o t u s Jordan and G i l b e r t , ( the "northern v a r i e t y " , Icelus (no  locality  not of  690.  1882b, p.  973.  1883, p.  577  Girard).  f e n e s t r a l i s Jordan and G i l b e r t ,  given)  (Commencement  and  Bolin,  Bay,  Washington);  1944, p.  relationships);  Wilby  1949);  ( l i f e history, (depth  48,  Jordan,  fig.18  Wilimovsky,  checklist); McAllister, and  Girard).  1882b, p.  A r t e d i u s f e n e s t r a l i s Jordan and G i l b e r t ,  only);  6 (No.  1960, p.  Clemens  1887,  p.  (description, 1954,  41  (name  (name  (name  distribution  only  only--from  -  Alaska Clemens  298, f i g .  186  d i s t r i b u t i o n ) ; MacPhee and Clemens, 1962, p.  33  distribution);  and Wilby, 1961," p.  898  Wilimovsky,  Alaska c h e c k l i s t ) ; Delacy, M i l l e r ,  1963, and  p.  Borton,  182 (name only 1972,  p.  15  27  (name  only);  Miller  and Lea, 1972, p., 126. F i g . p.  key and d i s t r i b u t i o n ) ; Alaska c h e c k l i s t ) ; 1973,  p.  1975,  128  (distribution-  key,  1972, p.  Blackburn, 1973, ( l a r v a e :  (name  Cottid  distribution);  only);  Peden  B r i t i s h Columbia);  and Dempster, 1979, p. Pearcy  Bay,  Oregon);  sp.  2 ) ; Washington,  and Wilson,  Richardson  1976, p. 235  and  Pearcy  and  19  (name  Myers, 1979, p.  variation); only  1982,  162, p i .  (larvae);  Jordan  and  Gilbert  Goldsborough,  A l a s k a ) ; S t a r k s , 1911, p. and  checklist);  Burke,  Eschmeyer,  California  977  Herald  188  298  (description, 36;  Kincaid  synonymy);  (distribution relationships); 1919, p.  30  (not of G i r a r d )  (in  1. 553, ( v i c i n i t y  of  Jordan and Evermann, 1898, p. 1903  A s t r o l y t e s n o t o s p i l o t u s Jordan and Evermann, 1900, 689a.  and  456 (name o n l y ) ;  (description,  A r t e d i u s a s p e r u l u s S t a r k s , 1896, p. Port Ludlow, Washington);  Artedius  1899 ( i n key, and  1899, p.  1907, p.  1912, p.  Hubbs 1926, p.  Hubbs,  Jordan and S t a r k s , 1895, p. 807  and Thompson, 1905, p. and  13 ( i n  212 ( l a r v a e - Yaquina  (Puget Sound); Jordan and Evermann, 1898a, p. distribution);  1977,  18.  Astrolytes fenestralis  Evermannn  -  Richardson and Washington, 1980, ( l a r v a e :  Hamman, 1983, p.  Gilbert  4 ) ; Hart,  F i t c h and Lavenberg,  d i a g n o s i s , synonymy, and m e r i s t i c  checklist);  Gilbert  20 (name only -  A r t e d i u s sp. 2 ) ; Howe and Richardson, 1978, p.  plus  Follett  and H a l l ,  478 ( l i f e h i s t o r y ,  p.  (larvae:  Quast  127 ( i n  fig.  28  Diagnosis  16.  Three to ten simple  short c i r r i  i n a l i n e on top of each of  t r a n s v e r s e head t u b e r c l e s . 17.  No c i r r i  28.  A  l o c a t e d along s u b o r b i t a l stay.  s i n g l e row of s c a l e s extending  forward  t o a p o i n t under  a n t e r i o r margin of o r b i t . 30.  Body s c a l e s c o n t i n u i n g  caudal  peduncle,  covering  onto  the d o r s a l  area  extending  surface  of the  from end of second  d o r s a l base t o o r i g i n of caudal f i n .  Spec imens Examined  USNM 27206 1 (Holotype) WA:Puget Sound; BC 63-252 2 63-245  Alaska;  BC  3 A l a s k a ; BC 53-74 12 B u r r a r d I n l e t ; BC 63-154 4 A l a s k a ;  BC 62-989 5 A l a s k a ; BC 62-555 4 A l a s k a ; BC 60-226 3 BC:Vancouver Id.;  BC 62-291 2 BC:Gardner Canal; BC 62-589 7 Alaska; BC 61-501  1 Alaska:Lynn 1  BCrJoassa  Canal; BC 53-232a 1 BC:Joassa Channel; BC Channel; BC 59-484 2 Alaska:Zachar  BC:Gale Passage;,BC AlaskarPt.  61-497  3  Alaska:Auke  Bay; BC 53-210 2  Bay;  BC  62-574  2  Armstrong; BC 61-495 2 Alaska:Auke Bay? BC 54-447 6  BC:Saturna I d . ; BC 53-86 5 BC:English Bay; BC Juan  53-232b  55-359  2  WArSan  I d . ; BC 62-881 6 BC:Sooke; BC 62-637 7 BC:Howe Sd.; BC 62-  731 2 BC:Saxe  P t . ; BC  CA:Sonoma,Bodega  53-156  3  BC:Echo  Bay; SU 16676 15 CArDel  Bay;  CAS  17765  6  Norte Co.; CAS 40354 7  29  Oregon; CAS Perpetua;  16662 10 CA:Del SU  Norte  Co.;  40882 1 CArHumboldt Bay;  CAS  LACM 4391  17208  UW  WA:San  Juan  Island;  3  OR:Cape  LACM 3.1700-4 5 CA:Diablo  Cove; LACM 4339 1 WArSkagit Co.; 4  40403  1 WArSkagit  Co.;  5450 15 WA:Edmonds; UW  980  UW 5  WA:West S e a t t l e .  Artedius  Artedius  harrinqtoni  (Starks  harringtoni  1896)  Range: Kodiak I s l a n d to southern Habitat:  Intertidal  especially  to  70  (Fig.  3)  California  feet,  usually  in  rocky  areas,  abundant in kelp beds.  Synonymy  Axyrias ( v i c i n i t y of  Port  1898a,  904  30  pi.  (in checklist);  harrinqtoni Ludlow, ( i n key, Schultz  Starks,  1896,  Washington); and and  p.  Jordan  554, and  p.  78  74  Evermann  d i s t r i b u t i o n ) ; Kincaid, DeLacy, 1936,  pi.  1919,  p.  (annotated  checklist) . Axyr i a s  harringtoni i  Bean  and  Weed,  1920,  p.  72  30  (except mature males). P t e r y q i o c o t t u s macouni Bean and Weed, 1920, p. 3 (Ucluelet,  B r i t i s h Columbia);  Jordan,  1923, p.  73, p i .  212  (name  only). Artedius (description,  harringtoni  distribution);  Clemens and Wilby  296.  185  Clemens,  33  Lea,  1972, p.  128,  history, only);  1973,  (larvae:  Peden and Wilson, Richardson,  Pearcy,  Richardson, meristic  (name  only  Cottid  1978, p.  1976  Richardson, Richardson (larvae);  -  and  and  Puget Sound); Miller  and  -  Alaska  checklist);  235  1975, p.  (distribution-  473 ( l i f e 128 (name British  A r t e d i u s sp.1); Richardson  Artedius  sp.  1); Howe and  Hubbs, F o l l e t t , and Dempster, 1979, p.  Yaquina  1980,  1961, p.  14 ( i n key, p l u s d i a g n o s i s , synonymy, and  variation);  (larvae  -  6); Hart, 1973, p.  p.  (larvae:  (name  MacPhee  15 (name o n l y ) ;  1977, ( l a r v a e :  1977,  41  128 ( i n key, and d i s t r i b u t i o n ) ;  (name only - C a l i f o r n i a c h e c k l i s t ) ; 212  history);  d i s t r i b u t i o n ) ; F i t c h and Lavenberg,  Columbia); and  20  1960, p.  distribution  1972, p.  f i g . p.  Quast and Hall,1972, p. Blackburn,  life  (depth  DeLacy, M i l l e r and Borton,  45, f i g . 17  1949); Clemens and Wilby,  (distribution,  1962, p.  1944, p.  McAllister,  only--from fig.  Bolin,  Bay  (larval  Washington,  Pearcy and Myers,  Oregon);  1979, p.  Richardson, Laroche, and  distribution 1980,  19  -  (larvae);  Oregon  Coast);  Washington, 1982,  Eschmeyer, H e r a l d and Hamman, 1983, p.  162, p i .  18.  31  Diagnosis  20.  In a d u l t males, t e e t h on  those  on  dentaries  and  numerous d i s t i n c t i v e l y  vomer  and  premaxillae  longer canine  palatines  irregularly  teeth  on  cardiform; canine,  with  upper  and  both  lower jaws. 21.  Seven b r a n c h i o s t e g a l  22.  43 to 46 s c a l e rows above l a t e r a l  23.  Incised  membrane  rays, one  of anal  a d d i t i o n a l on c e r a t o h y a l . line.  f i n convex in males, concave in  females. 24.  In males, anal f i n covered  l i g h t pigmentation  on a darker  with an hexagonal l a t t i c e w o r k of background.  25.  Penis present,  i n form of small cone.  30.  D o r s a l body s c a l e s c o n t i n u i n g onto d o r s a l s u r f a c e of  caudal  peduncle. 31.  One  very l a r g e c i r r u s  a n t e r i o r to o r b i t ,  plumose in  mature  ma1e s . 32.  No  cirri  the l a t e r a l 34.  in the region between the p e c t o r a l f i n base  and  line.  Seven to twelve simple  c o n f i n e d to p r e o p e r c u l a r  cirri  along p r e o p e r c u l a r  margin, not  spines.  Specimens Examined  SU 5047 1 (Holotype)  WA:Port  Ludlow;  BC  63-936  1  BCrJervis  32  Inlet;  BC 65-43 16 AK:Kodiak I s l a n d ; BC 61-301 3 BC:Bute I n l e t ;  BC 62-637 6 BC:Howe Sound; BC 65-43 x AK:Kodiak I d . ; CAS 29521 5 CA:Mendocino; CAS 29488 8 CA:Mendocino; CAS 25889  5  CA:Pacific  Grove; LACM 31937-8 1 CArHumboldt Co., T r i n i d a d ; LACM 31938-12 8 CA:Del  Norte Co.; LACM 1679 12 CA:San L u i s Obispo; LACM 7909 20  CA:San L u i s Obispo; UW 18020 s e r . WA:San Juan I s l a n d ; UW 3020 4 WArCape Johnson; UW 17208 4 WA; UW UW 14303 1 WA:San Juan  Island;  lateralis  ( G i r a r d 1854) ( F i g . 4)  Range: Kodiak I s l a n d to Baja Habitat:  Juan  Island.  Artedius  Artedius l a t e r a l i s  18020 6 WA:San  Intertidal  to  California. 45  feet.  Especially  abundant  in  t idepools. Synonymy  Scorpaenichthys  lateralis  (Monterey and San L u i s Obispo,  Girard,  1854  p.  145  California).  A r t e d i u s l a t e r a l i s G i r a r d , 1856, p.  134; 1857, p.  14,  33  (not p l a t e 22a), f i g s . synonymy);  Jordan  checklist);  and  Jordan,  807;  190  Greeley,  only)  1899,  p.  1940,  1919,  figs.  p.  McAllister,  (color  first  (description,  1949); and  and  Clemens,  ( i n key, and d i s t r i b u t i o n ) ;  key, 1975  also p.  Wilson,  M i l l e r and  checklist);  1976, p.  235 ( d i s t r i b u t i o n -  1978,  variation);  p.  55,  in  f i g . 21  41 (name only  Wilby,  1962, 507  Lea,  1961, p.  p.  33  (depth  (distribution,  1975  1972,  485  1972, p. p.  126,  Quast and H a l l , 1972,  Hart,  and d i s t r i b u t i o n ) ;  128 (name o n l y ) ; M a r l i a v e ,  Richardson, meristic  l i f e history  only,  DeLacy, M i l l e r and Borton,  fig.  20 (name only - Alaska  p.99 (name  r e c o r d ) ; Quast, 1968, p.  in checklist);  p.  1911, p.  (name  1960, p.  Clemens  v e r i f i e d Alaskan  distribution);  126  1913,  1944, p.  15 (name only, p.  description,  Starks,  30  d i s t r i b t i o n ) ; Hubbard and Reeder, 1965, p. synonymy,  ( i n key, and  58-74 (eggs, l a r v a l development);  key, and d i s t r i b u t i o n ) ;  MacPhee  437 (name o n l y , i n  72 (not 24765); Hubbs 1926,  (in  f i g . 187;  only - i n  Eigenmann and  1902  p.  4; B o l i n ,  - from Clemens and Wilby  ;  Halkett,  Bean and Weed, 1920, p.  Budd,  (name  only);  Hubbs and S c h u l t z , 1941, p.  299,  6  19  20 ( name  Kincaid,  174 ( d i a g n o s i s ,  Jordan and S t a r k s , 1895,  1896,  relationships);  in checklist);  7;  (name  Evermann,  Osgood, 1910, p.  checklist); p.  and  (description,  only,  p.  Jordan and Evermann, 1898, p.  distribution); habitat);  1881,  355 ( d i s t r i b u t i o n ) ;  Jordan  checklist);  Jouy,  1887, p.898  Eigenmann,1892, p. p.  5, 6; Gunther, 1860, p.  1973, p.  481 ( i n  F i t c h and Lavenberg, (larvae);  Peden  and  B r i t i s h Columbia); Howe and  14 ( i n key, plus d i a g n o s i s , synonymy and  Hubbs, F o l l e t t , and Dempster, 1979, p.  19  34  (name  only,  in  California  checklist);  Washington,  ( l a r v a e ) ; Eschmeyer, Herald and Hamman, 1983,  p.  I c e l i n u s l a t e r a l i s Jordan and G i l b e r t , P a r a r t e d i u s hankinsoni  Hubbs, 1926,  162,  pi.  1883,  p.  1982,  p.  18. 689.  4 (Point  Loma,  California). Artedius 1947,  p.  162;  d e l a c y i Hubbs and  Wilimovsky, 1954,  p.  Schultz, 285  1941,  (name  p.4.;  only  Bolin,  -  Alaska  checklist). A r t e d i u s hankinsoni  B o l i n , 1944,  p.  57,  fig.  22.  Comments: Artedius  hankinsoni  lateralis.  i s here f o r m a l l y synonymized with A.  hankinsoni,  two  f e a t u r e s were c i t e d as being d i a g n o s t i c : the low  s c a l e count  and  the m u l t i f i d upper p r e o p e r c u l a r  new  In Hubbs' (1926) c h a r a c t e r i z a t i o n of  Artedius  characters,  Hubbs had  but  includes  preopercular right  A. hankinsoni  remainder  variation Apart remains. study  s p e c i e s in A r t e d i u s , which  i s considerable v a r i a t i o n  spine w i t h i n each s p e c i e s , indeed spines examined  of in  specimen d i s p l a y e d anything The  the  B o l i n (1944) adds no  not.  In A r t e d i u s , there  l e f t and  spine.  are  one  individual.  this  study,  worthy of being  indistinguishable  found in A r t e d i u s  yielded  a  few  of  upper  even between the In  none  the  the save  called  putative the  type  "multifid".  from the normal range of  lateralis.  from the spine, only the low Examination  in  larger  additional  s c a l e count on the  series "A.  of l a t e r a l i s  hankinsoni"  body  in t h i s  types,  in  35  a d d i t i o n to numerous specimens intermediate between A. l a t e r a l i s and  A. h a n k i n s o n i .  A l l of  these  specimens,  i n c l u d e d , are i n d i s t i n g u i s h a b l e from A. l a t e r a l i s respect.  The e x i s t e n c e of these  represent  unique  i n possessing  other  leads to the c o n c l u s i o n that  A. l a t e r a l i s  development i s incomplete.  i n every  i n t e r m e d i a t e s , coupled with the  r e l a t i v e s c a r c i t y of A. h a n k i n s o n i , A. hankinsoni  A. hankinsoni  The type  in  which  the  scale  specimen seems t o have been  a m u l t i f i d preopercular spine, a feature  not shared by other p u t a t i v e A. h a n k i n s o n i .  Diagnosi s  22.  Twenty-four to twenty-six  surface,  extending  penultimate 29.  from  rows of s c a l e s along d o r s a l  second  or antepenultimate  or  third  dorsal  body  spine  to  d o r s a l s o f t ray.  No s c a l e s on body behind p e c t o r a l f i n base.  Spec imens Examined  UMMZ 126491 1 A K t d e l a c y i holotype;  1  AKtdelacyi  USNM 117494 1 A K t d e l a c y i d e l a c y i paratype;  UMMZ 55001  1 hankinsoni  paratype;  UMMZ  126490  holotype; BC 53-295 16 CA:Morro Bay;  BC  62-589  4  BCtSooke; BC 62-731 2 BC:Saxe P t . ; BC 53-38 1 BC:Nanaimo; BC 53-  36  232  1  BCrJoassa Channel; BC 53-40 2 WArCape Johnson; BC 53-296  32 CA:San L u i s Obispo; BCrSaturna  Island;  BC  BC  53-88  59-286  6 1  BCrNanaimo;  54-452  2  BCrBurrard I n l e t ; BC 62-42 1  BC:Saturna I s l a n d ; BC 54-448 2 BC:Saturna BCrStanley  BC  Island;  BC  54-96  1  Park; BC 53-88 8 BCrNanaimo; BC 60-238 4 Washington;  CAS 50393 17 CArSan N i c h o l a s I d . ; CAS 27316 16 CA:Duxbury  Reef;  CAS 50391 13 CA:Carmel;  CAS 50392 12 CArSanta Barbara; CAS 50388  4  68852  OR:Cape  Arago;  SU  MexicorBaja; LACM W71-9 Light;  LACM  8310  4  CA:Duxbury  3 CArMonterey; LACM 21132  1 CA:Palos Verdes ( h a n k i n s o n i ) ;  CA:San L u i s Obispo; LACM 33709-3 6 CA:Arena Mexico:Baja,  Santo  Reef; LACM 23612 1  Tomas;  LACM  876  9 OR:Yaquina LACM 1679 14  Cove; LACM  21133  2  1 C A : P a c i f i c Grove; LACM  31860-2 23 CA:San L u i s Obispo; UW 3019 16  WArCape  Johnson;  UW  3020  4 WArCape Johnson; UW 2312 4 C l e a r e d and s t a i n e d ; UW 15741  ser.  AKrKodiak I s l a n d  (delacyi);  Artedius  Artedius notospilotus Girard  Ranger  UW  17412 5 WArSan Juan  notospilotus  1856 ( F i g . 5)  Puget Sound (Washington) to Baja C a l i f o r n i a .  Habitatr  I n t e r t i d a l to 170 f e e t .  Island.  37  Synonymy  C a l c y l e p i d o t u s l a t e r a l i s Ayres, 1855, p.  77  (not of  Girard). Hemilepidotus nebulosus G i r a r d ,  1856, p.  134.  (refers  to Ayres p r e v i o u s l y unpublished name). Artedius Bay,  notospilotus  California);  p.71;  Gill  1857, p.  1862, p.  535,  454 ( i n c h e c k l i s t ,  p.  6 (name only, i n c h e c k l i s t ) ;  part);  Eigenmann  and  Bolin,  1944, p.  50  checklist  pi.  279 (name o n l y ) ;  p.  relationships);  Girard,  1856, p. 24,  Jordan,  ( i n key, Miller  and  Borton,  (name o n l y ) ;  Howe and Richardson, meristic  19 (name  1978, p.  figs. on  5, 6 (specimen no.  t h i s specimen o n l y ) ;  Girard,  Bean  and  Hubbs, F o l l e t t , and  1858, p.  Weed,  1975, p.  15 ( i n key, plus  163, p i .  366, c o l l e c t e d  15 ( i n  127, f i g . p.  in California  Eschmeyer, Herald and Hamman, 1983, p. lateralis  1972, p.  Lavenberg,  variation);  only,  distribution,  1972, p.  128  898 ( i n  355 ( d i s t r i b u t i o n ) ;  description,  ( i n key, and d i s t r i b u t i o n ) ; F i t c h and  Artedius  1887, p.  Eigenmann, 1892, p.  126  Dempster, 1979, p.  5,6; 1858,  and d i s t r i b u t i o n ) ; Jordan and Jouy, 1881,  DeLacy,  synonymy,  figs.  Jordan and G i l b e r t , 1881,  : PUget Sound); M i l l e r and Lea,  diagnosis,  134 (Tomales  checklist);  18. 71, p i .  22b,  by Ayres, f i g u r e  based  1920, p.  72  (27645  only) . Icelus (in  part).  n o t o s p i l o t u s Jordan and G i l b e r t ,  1882b, p. 690  38  A s t r o l y t e s n o t o s p i l o t u s Jordan 436  ( i n key, and d i s t r i b u t i o n ) ;  (not f i g . Puget  689a); Jordan,  Sound);  Starks  record of Jordan 653,  fig.  in Jordan  551  1905,  and  1898a, p. p.  442,  Morris,  and G i l b e r t ,  and Evermann,  1881,  1899, fig.  1907, p.  382  p.  61);  1900,  1896,  p.  fig.  689  (mature male,  219  (name only,  Jordan,  1925,  (mature male, Puget Sound, same specimen  p.  figured  1 905) . P a r a s t r o l y t e s n o t o s p i l o t u s Hubbs, 1926,  p.  2.  Diagnosis  10.  A d d i t i o n a l s h o r t , rounded s p i n e l e t s e n c r u s t i n g two  or three  main upper p r e o p e r c u l a r s p i n e s . 18.  In a d u l t s , lower  p r e o p e r c u l a r spines s e r r a t e d at  posterior  edge, extended d o r s o - v e n t r a l l y along p r e o p e r c u l a r margin. 19.  Six  to  twelve  irregularly  spaced  pores  in groove bounded  a n t e r i o r l y by ethmoid hump, p o s t e r i o r l y by o r b i t a l  margin,  not  d i v i d e d evenly by m i d l i n e . 39.  In  adults,  i r r e g u l a r groups of 40.  head  knoblets.  S e r r a t i o n s present  border  of  t u b e r c l e s r a i s e d d i r s a l l y , adorned with  supracleithrum.  on  posterior  posttemporal  and  upper  39  Specimens Examined  USNM  329  1 (Holotype) CA:Tomales Bay; USNM 27206 1 ( O r i g i n a l l y  syntype of A r t e d i u s  fenestralis);  USNM 26865 6 CA:Santa  Barbara;  CAS 27317 17 CArDuxbury Reef; CAS 50390 1 CA:San F r a n c i s c o ; 50389  2  Mexico:Baja;  CAS  W53-398 1 CA:Los Angeles; CAS Acc.#  1958-VI:9 1 San F r a n c i s c o ; CAS Acc.# 1972-1:24 1 13469  1  CA:San  Francisco;  No  Data;  'H52-168  4  MexicorPlaya  Maria;  SIO  74-122  Obispo; 1  Conception; SIO 63-1052 1 Mexico:Bahia San Q u i n t i n ; SIO 2  Mexico:Bahia  San  Q u i n t i n ; SIO 73-101  CAS  SIO 55-34 1 CA:San L u i s Obispo; SIO  H52-164 1 MexicotPunta Rocosa; SIO 55-105 8 CA:San L u i s SIO  CAS  Quintin;  SIO  12 CA:E1 Segundo.  63-1056  CA:Pt. 63-1054  2 Mexico:Bahia San  40  THE GENUS RUSCARIUS JORDAN AND STARRS 1895  Taxonomic R e v i s i o n Ruscarius  Jordan  and S t a r k s ,  1895.  Artedius  Bolin  (in part),  1944.  Artificial  key to Ruscarius  1a.  Scales on eye Ruscarius meanyi.  1b.  No s c a l e s on eye Ruscarius  creaseri. i  Based both  on  formerly  separate  the  phylogenetic  a n a l y s i s , meanyi and c r e a s e r i ,  included in Artedius,  monophyletic  lineage,  are  as  c h a r a c t e r s which are hypothesized  shown  they  to  share  comprise  none  a  of the  to be apomorphic f o r A r t e d i u s .  Conversely,  the c h a r a c t e r s t a t e s u n i t i n g meanyi and c r e a s e r i are  not  with  shared  originally Ruscarius,  Artedius  described  by Jordan  and  and Starks  Ruscarius  have never been diagnosed  a d i a g n o s i s i s here p r o v i d e d , of  strictu.  Since  meanyi  was  i n 1895 i n the genus  i t i s here proposed that t h i s genus be r e s u r r e c t e d to  i n c l u d e Ruscarius meanyi species  sensu  the s p e c i e s .  creaseri.  These  as a monophyletic u n i t .  i n a d d i t i o n t o diagnoses  two Such  f o r each  41  Ruscarius Jordan  Ruscarius  3.  Scale  ridge  almost  perpendicular  L a t e r a l body s u r f a c e covered  darker  interrupted  to b a s a l p l a t e , c t e n i i  by  with l i g h t well-defined  s t i p p l i n g of areas  without  any  A small patch of ten to twenty s c a l e s l o c a t e d j u s t caudad to axilla.  Upper p r e o p e r c u l a r  spine b i f i d ,  spine narrow and  f o r k i n g only  extended c a u d a l l y .  Adult males with dusky black  jaw,  t h r o a t , v e n t r a l body s u r f a c e and a l l f i n s except S c a t t e r e d s c a l e s on snout  hump and  28.  above ascending  Single  row  of  This  in p o s t e r i o r quarter of i t s l e n g t h .  11.  6.  darker  pigmentation.  d o r s a l edge of 7.  1895.  little.  pigmentation,  5.  Starks,  diagnosis  c u r v i n g very 4.  and  covering  lower  caudal.  above upper l i p , on t i p of ethmoid  processes  scales  pigmentation  of p r e m a x i l l a e .  extending  forward  under a n t e r i o r  orbit. 31.  One  to four c i r r i  l o c a t e d at upper a n t e r i o r o r b i t a l  margin.  42  Ruscarius  Ruscar ius c r e a s e r i  (Hubbs  creaseri  1926) ( F i g . 6)  Range: Carmel Bay ( C a l i f o r n i a ) to c e n t r a l  Baja C a l i f o r n i a .  H a b i t a t : I n t e r t i d a l to 90 f e e t .  Synonymy  Ruscariops c r e a s e r i Hubbs, 1926, p. San  Diego  County.  Paratypes  from  White  12.  (Bird  Rock,  P o i n t , Los Angeles  County, and Point Lobos, Monterey County, C a l i f o r n i a ) . Artedius (description, f i g . p.  Bolin,  relationships);  1944, p.  Miller  and  43,  fig.  and  Lavenberg,  128 (name o n l y ) ; Howe and Richardson, 1978, p.  plus  diagnosis,  F o l l e t t and Dempster, checklist);  synonymy and m e r i s t i c 1979, p.  Washington,  Hamman, 1983, p.  1982,  162, p i . 18.  18  (name  16  Lea, 1972, p. 126,  126, ( i n key, and d i s t r i b u t i o n ) ; F i t c h  1975, p. key,  creaseri  13 ( i n  v a r i a t i o n ) ; Hubbs, only  -  California  ( l a r v a e ) ; Eschmeyer, H e r a l d and  43  Diagnosis  9.  Three to  posterior  ten  edge  simple  of  opercle,  makes up i t s p o s t e r i o r 33.  cirri  in  a  cluster  at  dorsalmost  j u s t anterad to f l e s h y  flap  which  margin.  One t o ten simple c i r r i  in  cluster  anterad  to  uppermost  preopercular spine. 32.  One to three simple c i r r i  p e c t o r a l base and l a t e r a l  i n c l u s t e r midway between top of  line.  Specimens Examined  UMMZ 141866 1 B i r d Rock,  San  Mexico:Baja;  2 C A r B i r d Rock, San Diego; CAS 19671 1  CAS  19800  Diego  (holotype);  BC  63-979  Mexico:Baja; CAS 50119 1 Mexico:Baja; CAS 19516 2 CArSan CAS  19797 2 CArSan Diego; CAS 25382 1 CArSanta C a t a l i n a  CAS 19626 3 MexicorBaja; LACM 6587-5 39 4917  3  CArSan  Luis  32045-3 4 MexicorCedros  Obispo; Island;  LACM 32041-9 1 MexicorCedros  CArPalos  3  Diego; Island;  Verdes;  LACM  LACM 32082-1 3 MexicorBaja; LACM LACM  Island.  32053-11  3  MexicorBaja;  44  Ruscarius  Ruscarius  meanyi  meanyi Jordan and Starks  Range: Alaska  to Arena Cove  Habitat: I n t e r t i d a l  to 269  1895 ( F i g .  7)  (California). feet.  Synonymy  Ruscarius  meanyi  Jordan  and S t a r k s ,  1895, p.805, p i .  80 (Port Orchard Puget Sound, Washington); Jordan and 1898,  p.  (name only, (checklist  1908 ( i n key, and d i s t r i b u t i o n ) ;  Halkett,  in c h e c k l i s t ) ;  p.  -  checklist);  Puget  Schultz,  Artedius (redescription,  and  Sound);  1972,  1936, p.  meanyi  p.  1928  29, (name  fig.  66  only,  in  p.  Blackburn,  1973,  (larvae:  life  20  history,  record);  first  o n l y ) ; Peden and Wilson,  (name  British  and  1976, p.  Wilkie,  Columbia  1963  record);  - B r i t i s h Columbia); Quast  only  Cottid  and  3);  -  Alaska  Hart,  distribution);  Fitch  and  177 (key).  168 ( d i s t r i b u t i o n  1972,  California  Hubbs,  Rosenblatt  distribution:  Hall,  (figure,  1919,  1913, p.99  S c h u l t z and De Lacy, 1936, p.127 ( i n c h e c k l i s t ,  distribution);  Peden,  Kincaid,  Evermann,  Lavenberg,  checklist);  1973,  Lea,  1974  1975, p.  235, ( f i r s t  Alaskan  p.  483 (first  128 (name record);  45  Moulton,  1977  Richardson, Pearcy, 1978,  1977,  1977, p.14  variation); only,  (Puget  Sound:  (larvae:  (larvae:  habitat,  Icelus  Icelus  sp.  sp.  diagnosis,  Hubbs,  and  Herald and Hamman, 1983, p.  1); Richardson  synonymy,  1982,  163, p i .  and  meristic  Dempster, 1979, p.  i n C a l i f o r n i a c h e c k l i s t ) ; Richardson  ( l a r v a e : I c e l u s spp.); Washington,  distribution);  1);. Howe and Richardson,  ( i n key, plus Follett,  depth  19 (name  and Washington, 1980, (larvae);  Eschmeyer,  18.  Diagnos i s  1.  In a d u l t males, u r o g e n t i a l opening abuts o r i g i n of anal f i n .  8.  One spine and two s o f t  rays i n p e l v i c  37.  Maximum a d u l t s i z e 50 mm.  38.  In a d u l t males, f i r s t  44.  No c i r r i  47.  Scales  covering  on n a s a l cover  fins.  three anal rays thickened  and s h o r t .  spine.  upper  one-fifth  of  s u r f a c e of eye,  never  pupil.  Spec imens Examined  SU 3127 2 WA:Cotypes; CAS 37281 1 BC:Saanich I n l e t ; CAS 27695  2  CA:Mendocino; BC 62-495 1 BC:Sooke; LACM 38248-1 3 BC:Blind Bay; LACM  38247-1  3  BC:Raymond  I s l a n d ; NMC 77-0148 27 BC:Klaquack  46  Channel; NMC  68-0373 1 BC:Queen C h a r l o t t e  I s l a n d s ; SIO 63-599  BC:Howe Sound; SIO 73-227-55 1 CArArena Cove; UW 20721 I s l a n d ; UW 20720 1 WA:Seattle; UW uncat.  2  1 WA:Shaw  2 c l e a r e d and s t a i n e d .  47  PHYLOGENETIC ANALYSIS  L i s t of Adult  1.  P o s i t i o n of anus  2.  Shape of snout  3.  Form of s c a l e r i d g e  4.  Body c o l o r p a t t e r n  5.  S c a l e s above a x i l l a  6.  S c a l e s on  7.  Shape of upper p r e o p e r c u l a r  8.  Number p e l v i c  9.  Cirri  Characters  snout spine  rays  on o p e r c l e  10.  S p i n e l e t s on main p r e o p e r c u l a r  11.  Male c o l o r  12.  Chin c o l o r a t i o n  13.  Mandibular pore p a t t e r n  14.  Pores on l a t e r a l  15.  Form of head s c a l e s  16.  Cirri  on t r a n s v e r s e head t u b e r c l e s  17.  Cirri  on s u b o r b i t a l stay  18.  Form of p r e o p e r c u l a r  19.  Nasal  20.  Form of t e e t h  21 .  Branchiostegal  22.  Number of s c a l e rows above l a t e r a l  line  spine  scales  spine  pores  number line  48  23.  Anal  f i n membrane  24.  Anal  f i n pigmentation  25.  Penis  26.  Cirri  27.  Postcleithra  28.  S c a l e s under a n t e r i o r of  29.  S c a l e s behind  30.  S c a l e s on caudal  31.  Preorbital  32.  Cirri  above  33.  Cirri  anterad  34.  Cirri  on p r e o p e r c u l a r  35.  S c a l e r i d g e shape  36.  Scale r i d g e placement  37.  Adult  38.  Anal  39.  Form of head t u b e r c l e s  40.  S e r r a t i o n s on posttemporal  41.  S i z e of c i r c l e s on body  42.  White t h r o a t  43.  White spots on body  44.  Cirri  45.  Pterotic  46.  O s s i f i c a t i o n of o p e r c l e  47.  S c a l e s on  on upper l i p  orbit  axilla peduncle  cirri axilla to upper p r e o p e r c u l a r  spine  margin  size ray form  (males)  pigmentation  on n a s a l  spine  flange  eye  and  supracleithrum  49  List  of L a r v a l Characters  1.  Number of p r e o p e r c u l a r  spines  2.  R e l a t i v e s i z e of p r e o p e r c u l a r  3.  Basal p r e o p e r c u l a r  4.  Inner  5.  Skin bubble on nape  6.  D o r s a l gut d i v e r t i c u l a  7.  Parietal  8.  Nape melanophores  9.  Snout shape  spines  shelf preopercular  spines  spines  10.  Hindgut  11.  Number of p e l v i c  length rays  Adult Character  Character  State  1.  0:  spines  Analysis  P o s i t i o n of anus.  u r o g e n i t a l opening l o c a t e d s l i g h t l y anterad  to o r i g i n  of anal f i n base. State  1: i n a d u l t males, u r o g e n i t a l opening abuts o r i g i n of anal  fin. An Triglops,  anterior Blepsias  anus and  is  found  in  Orthonopias.  adults  of  Ruscarius  Clinocottus, c r e a s e r i and  50  A r t e d i u s sensu s t r i c t o have the piesiomorphic 0).  The  c o n d i t i o n in Ruscarius  any  of  the  outgroups  and  Character  2.  meanyi ( s t a t e 1) i s not  is  autapomorphic for Ruscarius  therefore  snout  (fig.  Snout  (state  and  to be  to upper  edge  of  orbit  to  width.  steep  in  lip  shape in the c o t t i d s  profile;  about  distance  one-half  i s highly variable.  snout, but a very  genera, i n c l u d i n g Orthonopias, 1).  hypothesized  width  from of  8)  have a long, f l a t a few  to o r b i t a l  short  a n t e r i o r edge of o r b i t orbit.  found in  Snout shape.  upper l i p about equal 1:  (state  meanyi.  State 0: snout long; d i s t a n c e from a n t e r i o r  State  condition  Artedius  sensu  A majority  short snout i s found in only Gymnocanthus and  s t r i c t u and  Ocynectes  Ruscar ius have  longer  snouts ( s t a t e 0).  Character  3.  Form of s c a l e r i d g e .  (fig.  State 0: s c a l e r i d g e almost p e r p e n d i c u l a r c t e n i i c u r v i n g to an angle State  State plate  little  (fig.  plate,  with  9a).  to b a s a l p l a t e , c t e n i i  9b).  2: no s c a l e r i d g e ; c t e n i i (fig.  to b a s a l  p a r a l l e l with p l a t e ( f i g .  1: s c a l e r i d g e almost p e r p e n d i c u l a r  c u r v i n g very  9)  originating directly  from basal  9c).  State 3: s c a l e r i d g e c u r v i n g  to  become  parallel  with  plate,  51  ctenii ridge  radiating (fig.  from  dorsal,  radiating  from  to  become  caudal  l a t e r a l edge of r i d g e without (fig.  and  caudal margins of  9d).  S t a t e 4: s c a l e r i d g e c u r v i n g ctenii  ventral  parallel  margin  ctenii,  of r i d g e only.  forming  a  The  s c a l e s in c o t t i d s are not  are  consisting  from  which  of  projections.  a  basal  is  found  to  is  be  not  hemilepidotus  plesiomorphic.  or  four  complete  ctenii  plate arise d i r e c t l y 1 i s found  and  from the base, with no  to  hypothesized which  for to be  includes  be  arise  various from State  Orthonopias, scale in  As  ridge  result,  sensu  synapomorphic  for  harrinqtoni,  State  2  is  intervening  is  for  Ruscar i u s .  hypothesized State  subgenus  only  in  the  ridge. and By  to  be  4  is  Artedius,  Artedius l a t e r a l i s ,  found  in  the  creaseri,  stricto. the  and  height as i t  a  Ruscarius  synapomorphic  Artedius  Artedius  Artedius c o r a l l i n u s .  structures  which l i e on or near the m i d l i n e of  comparison with the outgroups, s t a t e 3 synapomorphic  and  decreases  in Ruscar i u s meanyi and  hypothesized  bony  arise directly  The  approaches the m i d l i n e of the b a s a l p l a t e .  is  shelf  i n t e r v e n i n g r i d g e (spinous r i d g e ) .  in Hemilepidotus  hypothesized  Orthonopias  plate  These p r o j e c t i o n s ( c t e n i i ) may  the p l a t e , or from an  State  distinct  the usual t e l e o s t e a n c y c l o i d they  three  Antero-  ctenii.  or c t e n o i d bony r i d g e s c a l e , r a t h e r  is  plate,  9d).  S t a t e 5: no s c a l e r i d g e , no  0  with  and  Chitonotus  puqetensi s. Although  this  character  serves  to  distinguish  several  52  monophyletic  l i n e a g e s , i t has a n o n - a d d i t i v e d i s t r i b u t i o n at the  higher l e v e l of a n a l y s i s and i s t h e r e f o r e determining  inter-generic  regards to the Icelinus.  lineage  of  relationships,  including  little  value  particularly  Ruscarius,  in with  Chitonotus,  and  There are nine e q u a l l y - l i k e l y t r a n s f o r m a t i o n s e r i e s ,  none of which can be favored at p r e s e n t .  Character 4.  Body c o l o r p a t t e r n .  State 0: i n preserved specimens, pectoral  (Figs.  1-8)  the area bounded a n t e r i o r l y  f i n 'base, p o s t e r i o r l y by caudal f i n , d o r s a l l y by rows  of body s c a l e s , v e n t r a l l y by a l i n e d o r s a l to a n a l with  uninterrupted  darker  pigment.  a l t e r n a t i n g s t r i p e s of r e l a t i v e l y (fig.  8).  State  1:  lateral  body  generally  surface  pigmentation  darker  interrupted (the  lighter  latter  (figs.  The  Margins  darker  pigment  s t i p p l i n g of  background pigmentation by the  circular same  largest c i r c l e s ,  incomplete, producing a base.  and  scalloped  areas  6 and 7 ) .  areas  of l a t e r a l  lacking  darker  as v e n t r a l body s u r f a c e ) .  L i g h t e r c i r c l e s become l a r g e r c l o s e r to pigmentation.  may be arranged i n  i n t e r r u p t e d by w e l l - d e f i n e d c i r c u l a r  without any darker pigmentation 2:  This  f i n covered  body s u r f a c e covered with l i g h t  darker pigmentation,  State  by  ventral  edge  of  dark  at the v e n t r a l margin, are  appearance  above  anal f i n  of each c i r c l e c r i s p and w e l l - d e f i n e d ( f i g s .  1-  5). The  distinctive  " A r t e d i u s " p a t t e r n of  pigmentation  (state  53  2) i s found only i n A r t e d i u s sensu s t r i c t u and not i n any of the outgroups.  It  i s t h e r e f o r e p o s t u l a t e d to be synapomorphic f o r  the genus A r t e d i u s .  A s i m i l a r type of c o l o r a t i o n p a t t e r n occurs  in the c o t t i d genus Myoxocephalus, but on c l o s e examination  this  i s seen to be a p a t t e r n of i r r e g u l a r yellow b l o t c h e s on a darker background.  These b l o t c h e s continue to the anal f i n base and  not extend above the l a t e r a l be a d i f f e r e n t blotches  of  are a l s o found  crisp  i n diameter  in A r t e d i u s .  Irregular  pigment, s i m i l a r to the A r t e d i u s c i r c l e s ,  in C l i n o c o t t u s .  characteristically decrease  l i n e : hence t h i s i s hypothesized to  s t a t e from that observed lighter  do  However, these do not  outline  have  the  of A r t e d i u s c i r c l e s and  they  as they continue down to the a n a l f i n .  In  0 1 i g o c o t t u s , the p a t t e r n i s much the same as i n C l i n o c o t t u s : the margins  of  the  ventrally.  spots are i n d i s t i n c t  and they decrease  In 0 1 i g o c o t t u s , though, the middle p o r t i o n  in size of  each  spot i s covered with darker black s p e c k l e s . In  Chitonotus,  l a t e r a l body s u r f a c e pigment, line,  with  areas  is  are  and  covered  widely-spaced  this stippling  These  Icelinus,  i s broken  circular  Ruscarius  with  a  stippling  melanophores. up  by  varies  within  areas  without  Icelinus,  areas of l i g h t pigment, though in  Artedius.  roughly a l i g n e d  In  of  black lateral  pigment.  in I c e l i n u s and Chi tonotus, and  with  creaseri.  Icelinus  I_. f ilamentosus and p o s s i b l y J_. burchami having roughly  those  1 ) , the  Below the  v e r t i c a l bars i n R u s c a r i u s meanyi and Ruscarius pattern  (state  they  Icelinus  are  more  cavifrons  form The  b o r e a l i s, circular  irregular  than  these seem to be  i n v e r t i c a l bars, much as i n Ruscar i u s .  54  T h i s c h a r a c t e r has a d i s j o i n t two  transformation  this  hypothesis  subseries:  seems  to  distribution  0-1-2  lie  in  and  of  I t s main use i n  supporting  monophyly of Ruscarius and A r t e d i u s sensu  Character 5.  3-4.  consisting  the  separate  strictu.  S c a l e s above a x i l l a .  State 0: no s c a l e s l o c a t e d above a x i l l a . State  1:  a  small  patch  of ten to twenty s c a l e s l o c a t e d j u s t  caudad to d o r s a l edge of p e c t o r a l  fin  base,  not  arranged  in  rows. This is  in a  patch of s c a l e s ( i n Ruscarius c r e a s e r i and R. meanyi) different  Hemilepidotus  one of p a r a l l e l is  not  Stelgistrum,  from  hemilepidotus,  above the a x i l l a .  patch  location  The  the  which  is  scale behind  patch  found  in  the f i n and  not  other p a t t e r n of s c a l e s behind the f i n i s  rows, not a  patch  (see  found  in  Artedius,  Ricuzenius  or  Chitonotus  character Orthonopias, pugetensi s,  39).  This  Icelinus, and  is  p o s t u l a t e d to be synapomorphic f o r R u s c a r i u s .  Character 6.  S c a l e s on  snout.  State 0: no s c a l e s on  snout.  State  s c a l e s on snout above upper l i p , at t i p of  1:  scattered  ethmoid hump and above ascending processes of p r e m a x i l l a e . S c a l e s are found on R. c r e a s e r i  as  well  the as  two  snout other  in  Ruscar ius genera  meanyi  and  (Chitonotus  and  55  Ricuzenius).  F a r r i s o p t i m i z a t i o n leads to a hypothesis  independent  origins  for  this  7.  Character  State  upper  c a u d a l l y , having  (fourth)  This  spine  bifid,  preopercular  spine  not  extended  spine narrow and extended  forking  only  in  caudally.  p o s t e r i o r quarter of i t s  6 and 7 ) .  (figs.  A narrow, c a u d a l l y elongated s e v e r a l c o t t i d genera. and  preopercular  spine.  two to three prongs.  1: upper p r e o p e r c u l a r  Porocottus  These  i n A r t e d i u s .sensu s t r i c t u .  State  length  and C h i t o n o t u s .  Shape of upper p r e o p e r c u l a r  0:  two  f e a t u r e : once i n Ricuzenius and  once i n the l i n e a g e l e a d i n g to Ruscarius s c a l e s are not found  of  spine i s found i n  A long, unbranched spine i s  Myoxocephalus.  spine  preopercular  are  found  typical  of  V a r i a t i o n s on the " a n t l e r - l i k e " in  Chi tonotus,  Gymnocanthus,  L e p t o c o t t u s , Enophrys, I c e l i n u s and the Japanese genera S t l e n q i s and  Daruma.  arisen  The  number of times  this  i n the family cannot be estimated  hypothesis  including  time being,  then, only the s t a t e found  Ruscar i u s c r e a s e r i derived  trait.  8.  until  a  phylogenetic  a l l c o t t o i d genera i s generated. i n Ruscar ius  ( s t a t e 1) can be hypothesized A l l of  Number of p e l v i c  rays.  For the  meanyi  and  to be a unique,  the s p e c i e s i n A r t e d i u s sensu  have s t a t e 0.  Character  " a n t l e r - l i k e " spine has  strictu  56  State 0: one  spine, three s o f t  State  spine, two  1: one  There has meanyi by  been some  actually  its  small  has size  two  rays.  contention  two  the  c l e a r i n g and  rays i n h i s specimens.  had  three.  specimens,  Washington  Ruscar ius meanyi i n t h i s study Without  Ruscarius  distinguishing Lea  (1974) found  Richardson  found only one also  (1981) r e -  with two:  re-examined  staining.  She  and  of the one  staining,  thickened  rays  stained  found them to have two and  the  Lea's  found two  I examined c l e a r e d and  clearing  mistake the branched end  of  staining.  (1982)  t h i s time c l e a r i n g and  three.  whether  rays, p r i m a r i l y caused  Howe and  in a l l of the f i s h she examined.  never  to  difficulty  examined Lea's (1974) specimens and rest  as  or three p e l v i c  and  i n d i v i d u a l rays without only  soft  rays.  rays,  i t i s easy to  ray for two  normal  rays. Two  pelvic  Icelinus  and  Ruscarius this  are  also  Stlenqis.  characteristic  Washington  meanyi be p l a c e d  character.  arisen  rays  in a taxon  (1982) with  However, the s t a t e 1,2  independently  of  Icelinus  sister  taxon to Ruscar ius meanyi, and,  the  independent  (1982).  origin  of  R e f e r r i n g to Ruscarius  s t a i n e d specimens have 1,2 is  this  greatly  thickened  and  two.  state  meanyi, she  pelvic  (1,3) is  f u r t h e r , that  f i n rays.  An  that on  to have  I c e l i n u s , since  a n a l y s i s i n d i c a t e s that Ruscar ius c reaser i  i s the s i s t e r taxon to these  based  i s hypothesized  in Ruscar ius meanyi and  genera  suggested  present  ( a l s o 1,3)  the  the the  Chitonotus  indication  of  comes from Washington s t a t e d : " c l e a r e d and The  outermost  ray  branched at the t i p in a l l specimens  57  examined" (1982, p. Icelinus, 172,  "both  italics  148).  By c o n t r a s t , she observed  f i n rays  mine).  are  These two o b s e r v a t i o n s  s t a t e s i n Ruscarius meanyi and homologous.  relatively  Icelinus  that,  in  short and f i n e " (p. suggest  are  that the 1,2  not  structurally  A l l of the s p e c i e s i n A r t e d i u s sensu s t r i c t u have  the s t a t e 1,3 ( s t a t e 0 ) .  Character  9.  Cirri  on o p e r c l e .  S t a t e 0: one to three c i r r i  clustered  at  dorsalmost  posterior  edge of o p e r c l e , j u s t a n t e r i o r to f l e s h y f l a p which makes up i t s p o s t e r i o r margin. State  1: as i n s t a t e 0; but three to ten c i r r i  S t a t e 2: no c i r r i In  three  on o p e r c l e .  Orthonopias,  Ruscarius  Clinocottus,  01igocottus,  meanyi and A r t e d i u s sensu s t r i c t o ,  cirri  on  plesiomorphic Ruscar i u s  the  state.  creaseri  autapomorphic. secondarily  If  lost  opercle,  The p o s s e s s i o n (state Chi tonotus  these  cirri,  there  Icelinus, are  which i s hypothesized  1)  of up t o ten c i r r i is  is  hypothesized hypothesized  one  to  to be the here  by  to  be  to  have  t h i s c h a r a c t e r has the a d d i t i v e  transformation  s e r i e s 0-1-2.  Character  S p i n e l e t s on main p r e o p e r c u l a r  10.  per c l u s t e r .  spines.  State 0: absent. State  1: a d d i t i o n a l s h o r t , rounded s p i n e l e t s e n c r u s t i n g  two  or  58  three main upper p r e o p e r c u l a r These  additional  spines  spinelets  ( f i g . 5).  (state  1),  found  in  A r t e d i u s n o t o s p i l o t u s , are absent from the r e s t of .the examined (which a l l have s t a t e 0) and are t h e r e f o r e  adult  cottids  hypothesized  to be autapomorphic.  Character  State  11.  0:  in  Male c o l o r .  preserved  specimens, v e n t r a l surface of male the  same as female: b r a n c h i o s t e g a l membranes, v e n t r a l body, and f i n s light  colored.  State  1: males with a  lower  jaw,  dusky  branchiostegal  black  pigmentation  a darker,  In  almost black band along  a l l of  coloration females,  the  outgroups  i s either v i r t u a l l y or  (state 0).  the  membrane, and v e n t r a l body s u r f a c e .  In a d d i t i o n , a l l of the f i n s are dusky, with having  covering  and  the  first  dorsal  i t s d o r s a l margin. in  Artedius,  the  the same f o r males as i t  body  i s for  at most only the v e n t r a l f i n s and throat are dusky  Therefore,  the o v e r a l l dusky c o l o r a t i o n of males  Ruscar ius meanyi and Ruscarius  c r e a s e r i i s hypothesized  of  to be an  apomorphic s t a t e ( s t a t e 1).  Character  12.  Chin c o l o r a t i o n .  State 0: no p a t t e r n of c i r c l e s on v e n t r a l s u r f a c e of t h r o a t . State  1:  in  preserved  areas  of  lighter  specimens, a p a t t e r n of l a r g e c i r c u l a r  background  pigmentation  interrupting  dark  59  stippling,  sometimes  extending  b r a n c h i o s t e g a l membrane ( f i g . State 2: i n preserved of  light  anteriormost  10a,b).  pigmentation  Orthonopias;  caudally  outgroup  with  small  not  a d i s t i n c t l y pigmented t h r o a t i s  however, i t s arrangement of white v e r m i c u l a t i o n s on  strictu,  in  Artedius  which c o n s i s t s of an absence of darker  pigment,  the presence of white pigment superimposed  dark  background.  Chitonotus, (state  Ricuzenius,  0).  Ruscarius  of t h r o a t  pigment  Artedius  sensu  CIinocottus,  onto  the  extent, while  meanyi  and  a  uniformly  01igocottus,  stricto.  In  R. c r e a s e r i have s c a t t e r e d  to .be  state  1,  fenestralis,  branchiostegal  the  membrane  i n s t a t e s 2 and 3 i t covers  Artedius  at a l l  The presence of l i g h t e r c i r c u l a r  i s hypothesized  n o t o s p i l o t u s and A r t e d i u s  membrane.  on  and S t e l g i s t r u m lack any t h r o a t pigmentation  s t i p p l i n g on the t h r o a t .  extend  to  ( f i g . I0d).  a dark background i s not the same as the c o n d i t i o n sensu  darker  ( f i g . 10c).  State 3: as i n s t a t e 2, but c i r c l e s very only  blotches  interrupting  on v e n t r a l s u r f a c e of head, extending  b r a n c h i o s t e g a l membrane  The  p o r t i o n of  specimens, a p a t t e r n of c i r c u l a r  background  pigmentation  onto  harringtoni  has  small c i r c l e s of l i g h t e r pigmentation  synapomorphic found  in  pigment to  (state  for  Artedius does  not  any a p p r e c i a b l e  virtually a l l  state  areas  of the  2, while the very 3)  are  found  in  A. c o r a l l i n u s and A. l a t e r a l i s . This  character  has a non-additive  d i s t r i b u t i o n w i t h i n the  genus A r t e d i u s , and i s t h e r e f o r e of l i t t l e value the  relationships  therein.  However, i t serves  in elucidating to d i s t i n g u i s h  60  two  subgroups  including the  Artedius  lineage  lateralis. 2-3;  within  State  0:  Artedius  at  A d d i t i o n a l l y , two  to  twenty  midline.  One  anterior midline cluster McAllister  (rarely  which  of  minute  all  0-1-  share  state  pores  on  10c,d).  pores on each s i d e , o c c u r r i n g clustered  around  or two m i d l i n e pores l o c a t e d c a u d a l l y to (figs.  genera.  Orthonopias,  one)  to lower l i p ( f i g s .  10a,b).  (1968) i l l u s t r a t e d  pores i n many c o t t i d here  Artedius  s e r i e s are e q u a l l y l i k e l y :  s i n g l y or i n c l u s t e r s , plus four to ten pores ventral  and  p o s t e r i o r end of m a x i l l a , three s i n g l e  v e n t r a l m i d l i n e , j u s t caudad ten  corallinus  Mandibular pore p a t t e r n .  beginning  1:  Astrolytes,  0-1-2  pores on each s i d e .  State  subgenus  and A r t e d i u s n o t o s p i l o t u s , and  Three t r a n s f o r m a t i o n  13.  the  fenestralis  including  0-1-3-2; and  character  Artedius;  of  mandibular  Based on h i s study and  examination  Ruscarius, 0,  the  the  pattern  C l i n o c o t t u s and pattern  found  01igocottus, in  Artedius  f e n e s t r a l i s and A r t e d i u s n o t o s p i l o t u s ( s t a t e 1) i s p o s t u l a t e d to be  apomorphic.  Character  State  14.  Pores on l a t e r a l  line scales.  0: one pore l o c a t e d near p o s t e r i o r margin  line plate.  of each  lateral  61  State of  1: one  to three e x t r a pores l o c a t e d on ten to  last thirty  lateral  line scales.  twenty-five  E x t r a pores minute, e a s i l y  d i s t i n g u i s h a b l e from main pore l o c a t e d at  posterior  margin  of  scale. These  e x t r a pores are not  found in Ruscarius,  Oligocottus, C l i n o c o t t u s , Chitonotus, In  a l l of these genera, there  scale  ( s t a t e 0).  hypothesized  Character  State  15.  be  apomorphic  Artedius  State  2: no  directly  are  found only  line  therefore in Artedius  notospilotus.  from blunter  margin  of  basal  scales  r a d i a t i n g in scale  plate.  than c t e n i i of body s c a l e s .  h a r r i n g t o n i , Ruscar ius meany i ,  genera such as onto  the  markedly d i f f e r e n t Therefore  heads of A r t e d i u s  A.  lateral  s c a l e s on head.  Artedius  extending  group  1)  1: s c a l e s on o c c i p u t deeply embedded, c t e n i i  In  0).  pore per  Form of head s c a l e s .  C t e n i i much s h o r t e r and  other  are  Stelqistrum.  body.  directions  State  and  or  0: s c a l e s on head not d i s t i n g u i s h a b l e in form from  covering  all  i s only one  E x t r a , minute pores ( s t a t e  to  f e n e s t r a l i s and  Icelinus  Orthonopias,  Ricuzenius  and  head,  scales  the  in form from the  the  comparison,  l a t e r a l i s have no  Stelgistrum  A.  apomorphic. scales  on  the  scales  with  head.  not  (state  ( s t a t e 1) on by  corallinus Some  and  scales  body  n o t o s p i l o t u s are, Artedius  the  creaseri  on the o c c i p u t are  s c a l e s on  embedded, s t e l l a t e  f e n e s t r a l i s and  R.  the outand  Gymnocanthus  62  species  also  have bony " s c a l e s " on the head, but these c o n s i s t  of c o n i c a l p o i n t s a r i s i n g p e r p e n d i c u l a r a r i s i n g at an angle sensu  strictu.  to a  basal  plate,  not  from the margin of the p l a t e , as i n A r t e d i u s  T h i s c h a r a c t e r has the a d d i t i v e t r a n s f o r m a t i o n  s e r i e s 0-1-2.  Character  16.  Cirri  on t r a n s v e r s e head t u b e r c l e s .  State 0: one to three simple tubercles  on  head,  cirri  tubercles  just  anterad  in transverse  to  four  bony  l i n e p o s t e r i o r to  o r b i t a l margins. State  1 : three to ten c i r r i . l o c a t e d on each t u b e r c l e , i n a l i n e . The  three  plesiomorphic  cirri  on each t u b e r c l e .  s p e c i e s with s t a t e 1 . and and  Ruscarius C.  have  The remainder of A r t e d i u s s t a t e 0.  but  in  these  packed bunch. tubercle  in  to  strictu  C l i n o c o t t u s embryum, C.  analis,  number they  of  cirri  on  f e n e s t r a l i s and are i n a  fenestrali s  these  are not arranged  The presence of up to ten c i r r i Artedius  one  sensu  species  f a s h i o n as i n A r t e d i u s  is  A r t e d i u s f e n e s t r a l i s i s the only  q l o b i c e p s a l s o have a high  tubercles, linear  condition for tubercle c i r r i  is  in a  closely-  ( s t a t e 1) on each  interpreted  to  be  autapomorphic.  Character  17.  Cirri  on s u b o r b i t a l s t a y .  State 0: one t o two simple  cirri  on  suborbital  stay,  midway  between p o s t e r i o r end of p r e o p e r c l e and p o s t e r i o r end of o r b i t .  63  State  1: as  in s t a t e 0, but one  to two  cirri  s u b o r b i t a l stay at a p o i n t p o s t e r i o r to end to  two  point  simple  cirri  j u s t anterad  State 2: no c i r r i  at p o s t e r i o r end  there and  are  i s one  none ( s t a t e 2).  A.  the a n t e r i o r and This s i t u a t i o n two  in C l i n o c o t t u s and  State 0: State  1:  18.  bony  The  adults,  lower  from rounded to sharp,  autapomorphic.  at both  (state  1).  form  three  0.  always  simple.  margin  (fig.  7).  preopercular  spines  1)  varies  However, the range of v a r i a t i o n  the  Ruscarius  s t r i c t u have s t a t e  has  spines.  never branched or e l a b o r a t e d  of  (state  This character  s e r r a t e d at the p o s t e r i o r edge, extended  i n c o t t i d genera.  notospilotus  Artedius  s e r i e s 0-1-2.  Form of p r e o p e r c u l a r  form of the  serrated  stay  none,  i s , by comparison with the outgroups, apomorphic.  d o r s o - v e n t r a l l y along p r e o p e r c u l a r  greatly  the  i n a d u l t s , rounded to sharp, in  loss.  c o r a l l i n u s have, in a d d i t i o n , c i r r i  p o s t e r i o r ends of  the  Oligocottus  A r t e d i u s f e n e s t r a l i s a l s o has  the a d d i t i v e t r a n s f o r m a t i o n  The  spine.  c i r r u s l o c a t e d midway along  Ruscar ius s p e c i e s both have s t a t e 0.  Character  one  on s u b o r b i t a l s t a y .  ( s t a t e 0), while  and  plus  of  of s u b o r b i t a l stay, at a  t h i s i s here i n t e r p r e t e d to be a secondary  lateralis  The  of o r b i t ,  to uppermost p r e o p e r c u l a r  In Orthonopias there s u b o r b i t a l stay  at a n t e r i o r end  preopercular  is and  therefore  spines  in any in  postulated  runs way.  Artedius to  the remainder of A r t e d i u s  be sensu  64  Character  19.  State 0: two bounded  Nasal  to four  anteriorly  evenly  to two  1: s i x to twelve  d i v i d e d by  pores  located  n a s a l pores, Chitonotus  two and  p o s t e r i o r l y by a n t e r i o r  irregularly  spaced  pores,  CIinocottus  20.  In  I c e l i n u s , there are u s u a l l y two.  evenly  pores  spaced  01iqocottus, Possession  of  ( s t a t e 1) i s i n t e r p r e t e d  to  for A r t e d i u s n o t o s p i l o t u s .  remainder of A r t e d i u s sensu s t r i c t u have s t a t e  Ruscarius  and  the  0.  Form of t e e t h .  State 0: t e e t h on d e n t a r i e s , p r e m a x i l l a e ,  State  not  there are four evenly  on e i t h e r s i d e of the m i d l i n e .  autapomorphic  villiform  groove  on e i t h e r s i d e of m i d l i n e .  numerous i r r e g u l a r l y arranged  Character  in  midline.  In Orthonopias and  be  spaced  by ethmoid hump and  o r b i t a l margin, one State  pores.  vomer and  p a l a t i n e s in  bands.  1: c a r d i f o r m t e e t h on d e n t a r i e s ,  premaxillae,  vomer  and  palatines. State 2: those  in a d u l t males, t e e t h on vomer and  on  premaxillae  numerous d i s t i n c t i v e l y  and  dentaries  longer canine  palatines cardiform,  i r r e g u l a r l y canine,  teeth  on  both  upper  with and  lower jaws. In  Ruscarius,  Chitonotus never  and  elongate  Orthonopias,  S t e l q i s t r u m the t e e t h or  canine  (state  CIinocottus, are 0).  uniformly The  01igocottus, villiform,  c a r d i f o r m bands of  65  t e e t h in A r t e d i u s c o r a l l i n u s to  be  autapomorphic.  a d u l t male A r t e d i u s  ( s t a t e 1) are t h e r e f o r e  F u r t h e r , the  harringtoni  are  autapomorphic f o r that s p e c i e s . transformation  s e r i e s 0-1-2.  Character  Branchiostegal  21.  "fangs" also  postulated  ( s t a t e 2) found i n interpreted  T h i s c h a r a c t e r has  to  be  the a d d i t i v e  number.  State 0: s i x . State  1: seven, one In  all  a d d i t i o n a l on  c o t t o i d s , ' there  ceratohyal.  are  s i x b r a n c h i o s t e g a l s on each  s i d e , save f o r the p s y c h r o l u t i d s , which have seven (two epihyal,  f i v e on the c e r a t o h y a l ) and  in a separate  family.  Artedius  its  and  At  the  immediate  of  been  derived  occurrence nature  and  is  including of seven  interpreted  Artedius  i s apparently  found  in  Artedius  l a t e r a l i s : however, the frequency  very  low  and  the  true  frequency  i s unknown.  Character  22.  to  s e p a r a t e l y from the seven in p s y c h r o l u t i d s .  Seven b r a n c h i o s t e g a l s are a l s o i n f r e q u e n t l y fenestralis  analysis  taxa, p o s s e s s i o n  b r a n c h i o s t e g a l s by A. h a r r i n q t o n i ( s t a t e 1) have  the  are p l a c e d by some authors  level  sister  on  Number of s c a l e rows above the l a t e r a l  State 0: 29 to 38  rows.  State  1: 24 to 26  rows.  State 2: 43 to 46  rows.  line.  of in  66  In  Artedius  £enestralis,  Chitonotus,  Orthonopias  S t e l g i s t r u m , there are from twenty-nine to t h i r t y - e i g h t scales  on  the  body  o r i g i n of the f i r s t dorsal  base  above  the l a t e r a l  the  second  rows found i n  A r t e d i u s c o r a l l i n u s and A. h a r r i n g t o n i ( s t a t e  2)  is  therefore  counts  found i n  be  Artedius l a t e r a l i s hypothesized  to  apomorphic.  and have  a d d i t i v e transformation  Character  23.  high  end of of  to  The  of  number  hypothesized  0).  rows  l i n e , counted from the  d o r s a l f i n to the caudal  (state  and  Artedius arisen  The  lower  notospilotus  twice.  This  (state  1) are  c h a r a c t e r has the  s e r i e s 0-1-2.  Membrane of anal f i n .  State 0: i n c i s e d membrane of  anal  f i n concave  in  males  and  females. State  1:  in females  i n c i s e d membrane of a n a l f i n convex i n males, concave (fig.3).  In males of a l l of the outgroups examined, as Ruscar ius  and  Artedius  sensu s t r i c t u  well  as i n  save A. harr ington i , the  i n c i s e d membrane of the anal f i n i s concave i n  both  males  and  females ( s t a t e 0 ) . The c o n d i t i o n i n A r t e d i u s h a r r i n q t o n i , where the membrane i s convex i n males ( s t a t e 1), i s hypothesized  to be  autapomorphic.  Character  State  0:  24.  anal  Male anal f i n  pigmentation.  f i n dusky, sometimes with a l t e r n a t i n g l i g h t and  67  dark  areas.  State  1: anal f i n covered  l a c k i n g pigmentation In  on darker  Orthonopias  dusky, while narrow  band  with hexagonal  and  in Chitonotus  background.  of  areas  (fig.3).  Ruscar i u s ,  the anal f i n i s u n i f o r m l y  the  area  dusky  is  confined  to  a  set in s l i g h t l y from the d i s t a l margin of the f i n .  A- p a t t e r n of l i g h t and  dark s t r i p e s  other  such  cottid  latticework  genera  Myoxocephalus ( a l l s t a t e 0).  as  is  also  found  Hemilepidotus,  Therefore  in  several  01igocottus  and  the p a t t e r n of hexagonal  l a t t i c e w o r k on the a n a l f i n of A r t e d i u s h a r r i n q t o n i ( s t a t e 1) i s hypothesized  Character  to be autapomorphic.  25.  Penis.  State 0: absent. State  1: present,  s t a t e 2: present,  in form of small cone ( f i g . 3 ) much longer, with or without  elaborate  distal  processes. Modifications the C o t t i d a e , and "penis".  Clinocottus, to  Nautichthys  the u r o g e n i t a l p a p i l l a  differences  ranges  from  Icelinus, delicate  the  in  penis  massive  curved  Pseudoblennius, filaments  oculof asc i a t u s  and  in  complex  structures  in  label  of  s t r u c t u r e in v a r i o u s  and  Eurymen,  01 i g o c o t t u s  extremes are the long, slender c o n i c a l penes the  are widespread i n  are u s u a l l y lumped under the generic  However,  c o t t i d genera  paradoxus  of  Chi tonotus,  cylinders  •Psychrolutes Bero  elegans,  Between in  in  Radulinus Ocynectes,  these and and  68  Gymnocanthus.  In the  here, C h i t o n o t u s has neither Bolin  immediate a unique and  Hemilepidotus  in 1941  nor  penis  The  female  analysis,  within  the  no  family.  penis  ( s t a t e 2),  of  penis  but  (though  Orthonopias  has  an  presence of a s h o r t , c a u d a l l y - d i r e c t e d  in A r t e d i u s  as  consideration  have any  harr i n g t o n i  ( s t a t e 1) i s t e n t a t i v e l y  hypothesized to be an autapomorphic t r a i t of  under  elaborate  Orthonopias  r e p o r t s that the  extensible oviduct). conical  outgroups  at the  current  obvious s t r u c t u r a l homologue can  Ruscar ius  and  the  remainder  of  be  level found  Artedius  sensu s t r i c t u a l l lack penes ( s t a t e 0).  Character  State State  26.  Cirri  on upper l i p .  0: none. 1:  two  to  s i x t e e n simple paddle-shaped c i r r i  upper l i p , l o c a t e d anywhere from end distributed  evenly  premaxillary  symphysis.  The Artedius  are not  found in Ruscar i u s ,  Oligocottus  ( s t a t e 0).  lip  The  or  smaller  only other  i s Dasycottus s e t i g e r .  to Not  cirri  at  above the upper l i p  in  Chitonotus,  Stelgistrum  or  other  These c i r r i , l i p , and  as opposed to the paddle-shaped c i r r i  since  Orthonopias,  c o t t i d with c i r r i  found above the p o s t e r i o r p o r t i o n of the slender,  anterad.  c o r a l l i n u s ( s t a t e 1) appears to be autapomorphic,  CIinocottus,  upper  maxilla  U s u a l l y two  presence of paddle-shaped c i r r i  these c i r r i  Artedius  on each s i d e .  of  dorsad  members of above  the  though are  only  are  long  and  of A. c o r a l l i n u s .  69  Character  27.  Postcleithra.  State  0: p o s t c l e i t h r a present.  State  1: p o s t c l e i t h r a absent. The  postcleithra  are  present  absent in A r t e d i u s  lateralis,  A. c o r a l l i n u s  A. n o t o s p i l o t u s .  and  p o s t c l e i t h r a has  occurred  survey  available  of  the  n i g e r , three and  A.  in  the outgroups, but  fenestralis,  cottids  revealed  s p e c i e s of C l i n o c o t t u s  C. q l o b i c e p s ) , G i l b e r t i d i a  A. h a r r i n q t o n i ,  Apparently  s e v e r a l times w i t h i n  the  the  loss  of  family.  A  that Myoxocephalus  (C. a c u t i c e p s ,  C. embryum,  sigalutes, Ascelichthys  (which a l s o l a c k s p e l v i c f i n s ) and  are  in P s y c h r o l u t e s  rhodorus  paradoxus a l l  lack p o s t c l e i t h r a .  Character  28.  Scales  State  0: none.  State  1: a s i n g l e row  under a n t e r i o r of  of s c a l e s extending  under a n t e r i o r margin of In Orthonopias and  orbit. Chitonotus pugetensis,  Stelgistrum  s t e j n e g e r i , Ricuzenius  f e n e s t r a l i s , Ruscar ius meanyi and the  anterior  sensu s t r i c t u have no  optimization  point  the  ( s t a t e 0).  Artedius  a  continue forward under the a n t e r i o r p o r t i o n of  orbit  under  to  the  do  scales  forward  the s c a l e s on  head  Artedius  not  orbit.  based  orbit  R.  ( s t a t e 1).  s c a l e s in  this  pinetorum ,  creaseri The  do  have  remainder of  region.  Farris  on the most parsimonious cladogram suggests  four o r i g i n s f o r t h i s f e a t u r e :  once  in  Stelgistrum,  once  in  70  Ricuzenius,  once  in  Artedius  fenestralis,  and  once  i n the  l i n e a g e l e a d i n g to Ruscar i u s . t  Character 29.  S c a l e s behind  axilla.  State 0: none. State 1: two extending  to three p a r a l l e l  from  top  to  rows of  ventral  five  end  to  twenty  scales  of p e c t o r a l f i n , c l o s e l y  c o n f i n e d to f i n base. S c a t t e r e d s c a l e s are found in the area behind the a x i l l a i n Gymnocanthus, Scattered Pacific In  Ricuzen i u s ,  scales  or  prickles  of  the  preceding  c l o s e l y c o n f i n e d to Artedius,  and  the  extend  o p t i m i z a t i o n suggests that  was  confined  to  subsequently l o s t  in A.  and  also  Furc i n a ,  Hemilepidotus.  found i n the and  western  Pseudoblennius.  genera, however, the s c a l e s are not  pectoral caudally  fin  base  along  as the  they body.  i n A r t e d i u s sensu s t r i c t u , a  the p e c t o r a l base ( s t a t e twice, once i n A.  are  in  Farris row  of  1) arose once and  notospilotus  and  once  lateralis.  Character 30.  State  are  genera Bero, Ale i c h t h y s ,  all  scales  Stelgistrum,  0:  body  S c a l e s on caudal peduncle.  s c a l e s c o n t i n u i n g onto d o r s a l s u r f a c e of caudal  peduncle, c o v e r i n g area extending from end of second d o r s a l base to o r i g i n of f i r s t State  d o r s a l caudal  1: s c a l e s absent  raylet.  from d o r s a l s u r f a c e of caudal peduncle.  71  S c a l e s on the  caudal  Stelqistrum, Ricuzenius, They  are  absent  in  peduncle  Orthonopias, Artedius  A. n o t o s p i l o t u s ( s t a t e 1). is  0)  are  and Chitonotus  corallinus,  found  in  pugetensis.  A. l a t e r a l i s  and  The most parsimonious i n t e r p r e t a t i o n  that they were l o s t twice  including  (state  A. c o r a l l i n u s  in A r t e d i u s - once i n and  A. l a t e r a l i s ,  the  and  lineage once  in  margin  of  A. n o t o s p i l o t u s .  Character  31.  Preorbital c i r r i .  State 0: none. State  1: one to four c i r r i  orbit  ( f i g s 6,7).  State  2:  one  very  l o c a t e d at upper a n t e r i o r  l a r g e c i r r u s a n t e r i o r to o r b i t , plumose in  mature males ( f i g . 3). Preorbital c i r r i Chitonotus  are not found i n I e e l i n u s , Orthonopias  pugetensis  (state  0).  However they are present in  A r t e d i u s h a r r i n q t o n i ( s t a t e 2), Ruscarius, all  species  of 0 1 i q o c o t t u s  Jordania  ( a l l s t a t e 1).  zonope  At present  parsimonious i n t e r p r e t a t i o n i s that they arose in  or  and  the most  four times:  once  J o r d a n i a , once i n A r t e d i u s harr i n g t o n i , once i n Ruscar i u s and  once  in  01iqocottus.  A. h a r r i n q t o n i  s p e c i e s with p r e o r b i t a l c i r r i . transformation  s e r i e s 0-1-2.  Character  Cirri  32.  above  only  Artedius  T h i s c h a r a c t e r has the  additive  axilla.  is  the  72  State  0:  one  to three simple  between p e c t o r a l f i n base and State  in c l u s t e r at p o i n t midway  lateral  l i n e scale  row.  1: none. One  to two  Artedius  Icelinus  cirri  except  state:  C. a c u t i c e p s ) ,  The  the  Ruscarius  axilla  (state  0)  in  meanyi,  Orthonopias,  all  Icelinus  fimbriatus  and  F a r r i s o p t i m i z a t i o n p o s t u l a t e s four o r i g i n s once  once  once in I c e l i n u s , i f correct.  above  C. a c u t i c e p s ,  oculatus.  this  cirri  appear  harrinqtoni,  Clinocottus  for  cirri  in  in  Clinocottus  (with  a  loss  in  A. h a r r i n q t o n i , once i n R. meanyi,  Bolin's  remainder  (1947)  of  phylogenetic  diagram  A r t e d i u s sensu s t r i c t u  lack  and is  these  ( s t a t e 1).  Character  33.  Cirri  anterad  to upper p r e o p e r c u l a r  spine.  in  to  State 0: none. State  1: one  to  preopercular  ten  cirri  cluster  anterad  s p i n e , at margin of o p e r c l e and  uppermost  preopercle.  By outgroup comparison, the c l u s t e r of up to ten c i r r i in  front  (state  of the upper p r e o p e r c u l a r  1) i s autapomorphic, as  R. meanyi,  Artedius  01igocottus, (all  sensu  Chitonotus,  these  strictu,  Icelinus,  spine  in Ruscar ius c r e a s e r i  cirri  34.  Cirri  on p r e o p e r c u l a r  do  not  Orthonopias, Ricuzen ius  s t a t e 0).  Character  just  margin.  or  occur  in  Clinocottus, S t e l g i strum  73  0:  State  one  preopercular State  to  five  simple  not c o n f i n e d to p r e o p e r c u l a r Artedius  sensu  cirri  on  each p r e o p e r c u l a r  the  cirri  to  each  of  along p r e o p e r c u l a r  except  A. h a r r i n g t o n i ,  in  outgroups, there are from one t o f i v e  preopercular  spine  margin,  spines.  strictu  Ruscar ius and i n the other simple  confined  spines.  1: seven to twelve simple  In  cirri  (state 0 ) .  margin, with one or more on The d e r i v e d c o n d i t i o n ( s t a t e  1) found i n A r t e d i u s h a r r i n g t o n i , with seven to twelve c i r r i not confined  to the s p i n e s ,  i s hypothesized  Character  35.  i s shared  with C l i n o c o t t u s embryum,  to have a r i s e n twice w i t h i n the c o t t i d s .  Scale r i d g e shape.  State 0 : s c a l e r i d g e a broad curve State  and  ( f i g . 11a).  1: s c a l e ridge a p o i n t e d a r c h  (figs.  State 2: s c a l e ridge a s e m i c i r c l e ( f i g s .  11b,11c,11d). 11e,11f,11g,11h).  State 3 : no s c a l e r i d g e . The  plesiomorphic  state  I c e l i n u s , and Orthonopias. Chitonotus, aligned Artedius other  though  that found i n Hemilepidotus,  State  1  i t lacks a ridge  i n the same p a t t e r n . sensu  is  stricto  The  (state  is  found  Ruscarius.  ( s t a t e 3 ) , has i t s c t e n i i  semicircular 2)  in  arrangement  i s not found i n any of the  s c a l e d outgroups, and i s hypothesized  to be synapomorphic.  T h i s c h a r a c t e r has a d i s j o i n t d i s t r i b u t i o n and i t s in  this  study  i s severely l i m i t e d .  of  utility  Although a t r a n s f o r m a t i o n  s e r i e s cannot be e s t a b l i s h e d , i t nonetheless  provides  support  74  for  the  monophyly  the l i n e a g e  Character  of two  lineages: Artedius  i n c l u d i n g Ruscar ius and  36.  sensu s t r i c t o  and  Chitonotus.  Scale r i d g e placement.  State 0: s c a l e ridge not  reaching  margins of basal  plate  (fig.  11a). State  1: s c a l e ridge reaching  (fig. State  11b). 2: s c a l e ridge reaching  extending In  Hemilepidotus ,  be p l e s i o m o r p h i c . lateral  the  State  sensu  pugetensis,  reach  the  1,  only  stricto,  to  the  the  T h i s c o n d i t i o n i s hypothesized  to  the  ridge  reaching  the  anterior  Ruscar i u s ,  to  edge,  and  a n t e r i o r margin.  Hemilepidotus,  extending  only to  the  and  anterior  having edge  the  antero-  i n Orthonopias. is  Icelinus.  State  found  in  Chitonotus  ctenii  extending  Having the s c a l e ridge  the margins of the p l a t e i s hypothesized  relative  not  s c a l e r i d g e does not extend to  though i t l a c k s a r i d g e , a l s o has  way  only,  11c-11h).  edge of the p l a t e , i s found only  2, the r i d g e reaching Artedius  a n t e r i o r edge of p l a t e  to l a t e r a l edge ( f i g s .  edges of the p l a t e ( s t a t e 0).  all  a n t e r o l a t e r a l edge of basal p l a t e  to be  the c t e n i i is  apomorphic or the  hypothesized  ridge to  be  apomorphic r e l a t i v e to the c o n d i t i o n in Orthonopias. This 2.  Its  character main  Orthonopias Unfortunately  use  from no  has  an a d d i t i v e t r a n s f o r m a t i o n  lies the other  in  rest  separating  series,  Hemilepidotus  0-1and  of the genera under c o n s i d e r a t i o n .  character  shares  this  distribution.  75  Further c o r r o b o r a t i o n i s needed.  Character  37.  Adult  size.  State 0: maximum g r e a t e r than 130 mm standard State  1: maximum l e s s than 50 mm standard Adult  size  in  the  outgroups,  whole, i s g e n e r a l l y w e l l over in Myoxocephalus ( s t a t e 0). size  of  cottid,  Artedius and  meanyi  i s therefore  i n the c o t t i d a e as a  ranging  The small (state  length.  and  100 mm,  length.  up to about 750 mm  ( l e s s than 50  mm)  adult  1) i s not found i n any other  tentatively  hypothesized  to  be  autapomorphic.  Character  38.  Form of anal  State 0: f i r s t State  1:  three anal rays unmodified i n a d u l t males.  first  three anal rays thickened and short  l e n g t h of the f i f t h  ray)  Many c o t t i d s have notably 0 1 i g o c o t t u s two  are  at  least  ( f i g . 7). modified  anal  rays  in  in  the  as  long as, i f not longer  f i n membrane i t s e l f .  thickened,  meanyi ( s t a t e their  length  1), the  than,  males, In these  thickened, the t h i r d ,  set o f f by  a  The r e s t of the outgroups  under c o n s i d e r a t i o n do not have m o d i f i e d In Ruscarius  mature  two anal rays are s u b s t a n t i a l l y  unmodified, r a y . In 0. s n y d e r i , they are a l s o notch  (50% of the  maculosus and O l i g o c o t t u s s n y d e r i .  species the f i r s t  and  rays.  anal rays i n the males.  first  two  anal  rays  are  about h a l f of the l e n g t h of the f i f t h  76  (the f i r s t is  the  normal) r a y . The t h i r d ray, which i s not  same  length  as the f i r s t  two, while  approximately 75% the l e n g t h of the f i f t h . a  distinct  the f o u r t h ray i s  This  to  represent  and  to be autapomorphic f o r R. meanyi at the  thickened,  is  interpreted  s t a t e from that found i n 0 1 i g o c o t t u s , current  level  of  analysis.  Character  39.  Form of head t u b e r c l e s .  State 0: head t u b e r c l e s unmodified. State  1:  tubercles  raised  dorsally,  adorned  with  irregular  groups of k n o b l e t s . Porocottus, also  have  occur  in  Ereunias,  elaborate the  notospilotus  same (state  Dasycottus, I c e l u s and  bony s t r u c t u r e s on the head, none of which regions 1).  as  ridge',  those  found  in  Artedius  In Myoxocephalus, these are e i t h e r i n  the form of l o n g i t u d i n a l r i d g e s 'fronto-parietal  Myoxocephalus  of  Bolin  various  forms  (along  1947), such as i n M. s c o r p i u s ,  M. s c o r p i o i d e s , M. octodecimspinosus, and M. s t e l l e r i , are short s e c t i o n s of ridge with erose d i s t a l M. q u a d r i c o r n i s  and  M. g r o e n l a n d i c u s .  ridge  (which  is  from rounded bases and are structures  in  other  they  Although  four  of the  the  fronto-  not r a i s e d i n A r t e d i u s ) , they a r i s e globose  cottid  t h e r e f o r e not c o n s i d e r e d  or  s u r f a c e s , found i n  t u b e r c l e s i n A r t e d i u s n o t o s p i l o t u s are a l i g n e d with parietal  the  to be  A. n o t o s p i l o t u s i s hypothesized  genera  distally. and  homologous,  The  bony  A. n o t o s p i l o t u s and  the  to be autapomorphic.  state  head are in  77  Character  40.  Serrations  on  the  posttemporal  and  s u p r a c l e i thrum.  State 0: State  absent.  1 : s e r r a t i o n s present  upper border of In  Clinocottus,  Hemilepidotus ,  Icelinus  supracleithrum  are  State  41.  of  posttemporal  01igocottus,  Ruscar ius (state  serrations  Size of body  no  tiny  l a t e r a l body s u r f a c e 1:  and  the  Chitonotus,  posttemporal  0).  The  (state  1)  presence is  and in  therefore  tiny  State 2: as These  of  (figs.  6,7).  circles  Artedius,  A. c o r a l l i n u s  they A.  lighter  not are  (figs.  pigmentation  1,2). 3,4,5).  found in the outgroups ( s t a t e 0). found  lateralis  (state  background  c i r c l e s dense ( f i g s .  A. c o r a l l i n u s ,  dense  l i g h t e r pigment s c a t t e r e d over  body s u r f a c e  are  and  A. l a t e r a l i s and particularly  of  in s t a t e 1, but circles  circles.  circles  s c a t t e r e d over l a t e r a l  Within  of  to be autapomorphic.  0:  State  and  unmodified  Artedius notospilotus  Character  posterior  supracleithrum.  Orthonopias,  postulated  on  only  (state  in 1).  A. harr i n g t o n i , The  i n which the small  2), i s hypothesized  c o n d i t i o n in circles  are  to be a f u r t h e r  derived condition. This character,  because of  its  non-additive  distribution  78  within  Artedius,  generic  relationships.  the  sister  cannot  group  as  strong evidence  for intra-  I t can, however, c o n t r i b u t e support  status  l a t e r a l i s , as w e l l as the A. l a t e r a l i s ,  serve  of  for  A r t e d i u s c o r a l l i n u s and A r t e d i u s  integrity  A. c o r a l l i n u s ,  of  and  the  clade  containing  A. h a r r i n q t o n i .  Three  transformation  s e r i e s are e q u a l l y l i k e l y : 0-1-2, 0-2-1 and 0-1  character  White throat c o l o r a t i o n .  42.  State 0: none. State  1: white spots present  on t h r o a t , i n r e g u l a r p a t t e r n ( f i g .  I0e). The the  presence of white v e r m i c u l a t i o n s on  throat  in  Orthonopias  (statel)  is  the  underside  hypothesized  autapomorphic, as i t i s not found i n any of the other in Ruscarius spots  or i n A r t e d i u s sensu s t r i c t u  appear  in  (all  to  Character  state  0).  The  a r e g u l a r p a t t e r n , with a "Y" j u s t behind the  c a u d a l l y along  43.  be  outgroups,  median mandibular pore, and a s e r i e s of r e g u l a r l y grouped proceeding  of  the b r a n c h i o s t e g a l  spots  membrane.  White spots on body.  State 0: absent. State  1: present In  the  ( f i g . 8).  area  which  in  Artedius  Orthonopias has a s e r i e s of white spots  is  covered  superimposed  by c i r c l e s , over  the  79  u n d e r l y i n g darker distinctly ventral darker the  pigmentation  lighter  body  ( s t a t e 1).  that the l i g h t background pigmentation  surface,  so  that,  body  indistinct,  in  color.  contrast  A r t e d i u s sensu s t r i c t u  Character  44.  Cirri  State 0: a simple  The to  of  the  beyond the v e n t r a l margin of  pigment, the white c i r c l e s are  lighter  T h i s white pigment i s  clearly  margins  the  visible  of  the  against  circles  sharply-defined  circles  are of  (state 0).  on n a s a l  spine.  slender c i r r u s a r i s i n g  from base of each n a s a l  spine. State  1: no c i r r u s on each s p i n e . In  Orthonopias  and  Chitonotus  slender c i r r u s on each n a s a l spine  pugetensis, (state  0).  there This  found i n Ruscar ius c r e a s e r i , A r t e d i u s h a r r i n g t o n i , A. and to  A. be  lateralis. the  Possession  plesiomorphic  optimization  suggests  Ruscar ius meanyi, in the A. n o t o s p i l o t u s , Clinocottus four  once  of a n a s a l c i r r u s i s  condition  losses  (state  stem each  for  leading in  (C. g l o b i c e p s ) , and  for  to  01igocottus at l e a s t  A.  is  also  corallinus hypothesized  Artedius.  1)  i s a long  Farris  t h i s f e a t u r e in fenestrali s  and  (0. maculosus) and  three  times  for  the  I c e l i n u s s p e c i e s which lack i t (l_. burchami, I_. c a v i f rons,  I_. q u a d r i s e r i a t u s , I_. t e n u i s ) , correct.  Character  45.  Pterotic flange.  if  Bolin's  (1947)  diagram  is  80  State not  0: t r i a n g u l a r flange at p o s t e r i o r edge of p t e r o t i c  reaching p o s t e r i o r edge of cranium  State  1: f l a n g e long, extending  (fig.  (fig.  short,  12a).  to p o s t e r i o r margin  of  cranium  12b).  State  2:  as  in s t a t e 1, but d i s t a l edge of lower flange r a i s e d  to form a r i d g e p e r p e n d i c u l a r  to  plane  of  main  flange  (fig.  12c). The  posterior  flange  of  the  p a r a l l e l t r i a n g u l a r elements which direction.  The  Icelinus, meanyi,  lower  Hemilepidotus, R. c r e a s e r i  and  s h o r t , never extending (state  of  0).  the  pterotic extend  two  state  (state  lateralis,  c o n d i t i o n and considered  ventro-caudal  i s always the l o n g e s t .  In  Oligocottus,  Ruscar ius  Orthonopias,  this  i s always  to the  posterior  flange  edge  1)  to  is  end  i s hypothesized State 2,  hypothesized  to  of  the  cranium  strong evidence  Character  O s s i f i c a t i o n of o p e r c l e .  State 0: o p e r c l e completely 1: ventro-caudal  neurocranium.  to be synapomorphic f o r in  be apomorphic at that l e v e l . provide  the  A. c o r a l l i n u s  represent  series is additive  46.  of  found  transformation  State  two  In A r t e d i u s sensu s t r i c t o t h i s f l a n g e i s very much  A r t e d i u s sensu s t r i c t o . A.  a  of  Clinocottus,  longer, c o n t i n u i n g to the p o s t e r i o r This  in  consists  and  a further derived  This  for these  character  is  groups as  the  (0-1-2).  ossified.  t h i r d of o p e r c l e not  In C I i n o c o t t u s , 0 1 i q o c o t t u s  and  ossified.  A r t e d i u s sensu s t r i c t o ,  the  81  ventro-caudal  section  ossified  does  and  Chitonotus,  of  not  the take  Orthonopias,  meanyi  and  ossified  ( s t a t e 0).  opercle up  is  alizarin  Hemilepidotus,  R. c r e a s e r i , State  the  never  completely  red ( s t a t e 1).  Icelinus,  opercle  is  1 i s hypothesized  Ruscarius  always to  be  In  completely a  derived  in c r e s c e n t  covering  trait.  Character  47.  S c a l e s on  State 0: no s c a l e s on State  eye.  eye.  1: s c a l e s on s u r f a c e of eye,  upper f i f t h of eye, Scales  on  Ruscar ius meanyi  never c o v e r i n g  the  arranged pupil.  eye are found i n Chitonotus  ( s t a t e 1).  i s one  the  and  leading  Ruscar ius c r e a s e r i . sensu s t r i c t u  to  Ruscarius  These s c a l e s  ( s t a t e 0).  and  F a r r i s o p t i m i z a t i o n p o s t u l a t e s that  the most parsimonious e x p l a n a t i o n stem  pugetensis  are  of a s i n g l e o r i g i n ,  Chitonotus, not  found  in  with a l o s s in in  Artedius  82  RESULTS  Adult  Monophyly and  The 13,  14,  R e l a t i o n s h i p s of  Artedius  Wagner program produced three t r e e s of 86 15), each with a c o n s i s t e n c y  recognizes nominal  a  monophyletic  Artedius  A. h a r r i n g t o n i , and  Trees  species  tree  c l a d e composed of f i v e of the  seven  (A. n o t o s p i l o t u s ,  in  and a  stricto  (form  of  scale  coloration), (pterotic  is  ridge),  27  separate  4  (see f i g .  Artedius,  A. n o t o s p i l o t u s  two  p l u s A.  color),  scales), and  a  32  (body  A. l a t e r a l i s ,  Within  12  with  separate  closely  12  (chin  ( s c a l e r i d g e shape), and  clades  45  are  identifiable:  by  synapomorphies  (chin  line pores), 44  (cirri  A. h a r r i n g t o n i ,  synapomorphies  clade,  pattern),  f e n e s t r a l i s supported  (chin c o l o r a t i o n ) and  the l a t t e r  not  meanyi  16).  (extra l a t e r a l  including  R.  monophyly of A r t e d i u s  color  ( c i r r i . a b o v e a x i l l a ) , and  clade  pattern),  14  clade,  The  in c h a r a c t e r s 3 (body c o l o r p a t t e r n ) , 12 (male  fenestralis,  by s i x synapomorphic c h a r a c t e r s : 3  ( p o s t c l e i t h r a ) , 35  flange)  Within  supported  A.  A. c o r a l l i n u s ) .  r e l a t e d to the remainder of A r t e d i u s . sensu  of 76.74.  (figs.  Each  A. l a t e r a l i s ,  R. c r e a s e r i are p l a c e d  index  steps  coloration), 15  11  (form of head  on nasal s p i n e ) ;  A. c o r a l l i n u s  and  in c h a r a c t e r s 4 (body c o l o r 41  ( s i z e of c i r c l e s on  A. c o r a l l i n u s  and  A.  body).  lateralis  are  83  sister  species,  coloration), axilla), (see  sharing apomorphic s t a t e s i n c h a r a c t e r s  17  (cirri  on  suborbital  12 (chin  s t a y ) , 32 ( c i r r i  41 ( s i z e of c i r c l e s on body) and 45  (pterotic  above flange)  f i g . 16). Each  of  Clinocottus  in  supporting character  the  three  one  t r e e s p l a c e d A r t e d i u s , O l i q o c o t t u s and  monophyletic  t h i s p a r t i c u l a r clade 49  Clinocottus  (ossification  are  hypothesized  mainly on the b a s i s of t h e i r genera  lack  genealogical  scales  this  no  refutation  apparent.  For present  these  two  opercle).  to  be s i s t e r  lack of s c a l e s .  evidence  Oliqocottus taxa  Since many  not strong  i s possible  aspects  of  Oliqocottus-Clinocottus-Artedius  but  are  study cottid  evidence f o r character  a l t e r n a t i v e s are  be s u f f i c i e n t the  to  cladogram  clade,  clade)  and  in this  In the absence of f u r t h e r  purposes i t should  01igocottus-Clinocottus  The  i s s l i g h t and c o n s i s t s only of  i s obviously  relationship.  evidence  that  of  lineage.  note (the  and  provisional  the  and  require  further t e s t i n g .  Monophyly of Ruscar i us  Ruscar ius meanyi and Ruscar ius c r e a s e r i do the  monophyletic  lineage  made  up  not  of the other  belong  five  s p e c i e s , although they do c o n s t i t u t e a separate c l a d e own  as evidenced by c h a r a c t e r s  color)  (see f i g .  Artedius of  their  3 (form of s c a l e r i d g e ) , 4 (body  c o l o r p a t t e r n ) , 7 (shape of upper (male  to  17).  preopercular Thus  as  spine)  Washington  and  11  (1982)  84  suggested the genus A r t e d i u s  sensu B o l i n (1947) i s  demonstrably  diphyletic.  A l t e r n a t i v e t r e e s - the r e l a t i o n s h i p s of Ruscar ius  The  only ambiguity present  concerns the Ruscarius  r e l a t i o n s h i p s of Chitonotus,  creaseri,  Artedius,  especially  Clinocottus,  parsimonious differing  in the t r e e s f o r the adult  in  only  i n the placement of Ruscarius  is  and  lineage  Three  of  equally  I c e l i n u s are p l a c e d  Chitonotus:  the  01igocottus-Clinocottus-Artedius  sister  lineage  lineage,  s e q u e n t i a l l y below  to  the  while Chitonotus  Ruscar ius  on  the  and tree  13). 2.  Chitonotus  l i n e a g e , while I c e l i n u s (Orthonopias 1ineage  and  (fig. 3.  plus is  Ruscarius  placed  Hemilepidotus)  Chitonotus i s placed  i n c r e a s i n g l y plesiomorphic Unfortunately, level  that  Icelinus  form  between and  the  as the  monophyletic  the  basal  genera  Chitonotus-Ruscarius  lineage,  Ruscarius  and  the only c h a r a c t e r s  includes  R. meanyi,  are non-additive  sister  characters  taxon  followed  Icelinus ( f i g .  R. c r e a s e r i ,  the  by  the  15).  Chitonotus,  multistate characters,  at t h i s l e v e l ,  to  which are d e r i v e d at  useful in e l u c i d a t i n g r e l a t i o n s h i p s .  binary  a  14).  01igocottus-Clinocottus-Artedius  very  r e l a t i o n to the  01igocottus.  and  meanyi,  a l t e r n a t i v e s emerge from the Wagner a n a l y s i s , each  1.  (fig.  Ruscarius  data  In  which, are  the  absence  the and not of  the branching sequence remains  85  ambiguous. as  a  T h i s ambiguity appears i n the Adams  trichotomy,  from which stem Chitonotus,  Artedius-01igocottus-Clinocottus clade Although there placement  of  i s ambiguity  Chitonotus,  c l o s e r to Chitonotus clade  than  and  i t i s to  suggests the r e v e r s e . to  the  given  Chitonotus  leaving  the  to  only  Chitonotus  i n the  appears to be  the A r t e d i u s - Q l i g o c o t t u s - C l i n o c o t t u s  Icelinus.  Washington's (1982) diagram  there  two  i s no evidence  of  shares  6  as  with  Chitonotus  on  9  and  i n another  a  useful  which 15, 16,  cannot  In b i n a r y  (characters  homoplasious  character  and the  tree  Ruscar ius  c h a r a c t e r s at hand.  be  also  Adams  f o r one of the three t r e e s ( f i g s .  has losses, i n  hypothesized  the  genus  17) over another, s i n c e the p o s i t i o n resolved  Ruscarius  tree  ( f i g . 18).  in  At present  base a preference  consensus  be  characters, 17)  and i s  ( c h a r a c t e r 47),  binary  character.  I c e l i n u s apomorphic s t a t e s i n two  m u l t i s t a t e c h a r a c t e r s which have d i s j o i n t d i s t r i b u t i o n s over the e n t i r e t r e e ( c h a r a c t e r s 4 and 35) and has character  which i s n o n - a d d i t i v e  additive character  a  state  in  another  (character 3 ) . T h i s leaves the  36, f o r which i t shares  the  same  state  as  Ruscar i u s , I c e l i n u s and A r t e d i u s . F i n a l l y , Hemilepidotus and Orthonopias remain unresolved at the  base  of  plesiomorphic of the taxa especially The the data  the  phylogenetic  hypothesis,  since  they  are  f o r a l l of the c h a r a c t e r s with respect to the r e s t at  hand.  Orthonopias  i s highly  autapomorphic,  i n body c o l o r a t i o n ( c h a r a c t e r s 45 and 46). Wagner  program produced only one t r e e ( f i g . 24)  set excluding  R. meanyi  to  make  i t comparable  (for to  86  Bolin's fully  1947  tree).  resolved,  A r t e d i u s , 01igocottus  with the same branching sequence as i n the t r e e s  in which R. meanyi was i n c l u d e d . placed with  together the  R. c r e a s e r i and Chitonotus a r e  as a separate l i n e a g e  sharing a common  Artedius-Qligocottus-Clinocottus  plesiomorphic  r e l a t i v e to  Hemilepidotus  and C l i n o c o t t u s are  and  a l l of  Orthonopias  the  are  clade.  above  again  ancestor  Icelinus i s  taxa.  Finally,  relegated  to a b a s a l  polytomy.  L a r v a l Trees  Three t r e e s of seventeen steps emerged from  from  the  (1982)  collapsed  exactly  Washington's  genera,  although  from an unresolved R. meanyi, resulting  (1982)  One  tree  branching  Artedius, 01igocottus, trichotomy.  trees  (fig.  and  23)  extracted have  a  19) d u p l i c a t e s  sequences  within  her  and C l i n o c o t t u s emerge  The part of the  tree  including  and I c e l i n u s i s the same as i n the t r e e in  Washington's  ( f i g . 25).  second t r e e  (fig.  in the c o l l a p s e d t r e e , as i s the clade.  A l l three  from c o l l a p s i n g unsupported branches  the  22,  None of these t r e e s i s i d e n t i c a l to  tree.  R. c r e a s e r i  dichotomous t r e e In  study.  index of 82.35.  Washington's  21,  Wagner a n a l y s i s of the l a r v a l data  Washington's  consistency  (figs.  01igocottus  20), C l i n o c o t t u s  i s r e s o l v e d as  R. meanyi-R. c r e a s e r i - I c e l i n u s  i s now the s i s t e r  taxon to A r t e d i u s ,  A. h a r r i n g t o n i , which emerges from a trichotomy with  (less  Clinocottus  87  and  the 0 1 i g o c o t t u s - A r t e d i u s The  t h i r d and f i n a l  01igocottus This  and  branch  lineages: of  from  Clinocottus  Artedius  l a r v a l cladogram ( f i g .  Artedius  arises  lineage).  (less a  trichotomy  and  R. meanyi-R. c r e a s e r i - I c e l i n u s as  i n the f i r s t  tree derived  two t r e e s .  since  it  not  analysis  does  Artedius  CIinocottus are  species  acut i c e p s .  i s the  include  a l l of  the  R. meanyi  Adams  consensus  species  not  and the  resolved.  The  demonstrate, however, that R. meanyi and  under  i n c l u d i n g the  consideration.  or R. c r e a s e r i .  Adams consensus t r e e f o r the  three  rest  01igocottus  a r e i n f a c t more c l o s e l y r e l a t e d  either  The  the l a r v a l t r e e i s of l i t t l e use,  R. c r e a s e r i do not belong to the l i n e a g e the  other  of the t r e e i s the same  r e l a t i o n s h i p s of those i t does i n c l u d e are larval  two  l a r v a l Wagner t r e e s .  to A r t e d i u s ,  does  24  has  and a clade made up  CIinocottus portion  also  i n one l i n e a g e .  involving  acuticeps)  Figure  from these three  With respect  harringtoni)  ( l e s s C.  harringtoni  21)  to  Artedius  of and than  T h i s can be seen i n the  larval  trees  (fig.  22).  I c e l i n u s p l u s R. meanyi p l u s R. c r e a s e r i c o n s t i t u t e one r e s o l v e d clade, branches  while  emerging  fenestralis Artedius  the  plus  remainder in  a  Artedius  of  the  polytomy: harringtoni  t r e e i s composed of f i v e 01igocottus, plus  Artedius  h a r r i n g t o n i , C l i n o c o t t u s acut i c e p s , and. the  of C l i n o c o t t u s .  Artedius type 3,  remainder  88  COMPARISON OF  CLASSIFICATIONS  Adults  The t r e e p u b l i s h e d steps  more  R. meanyi  than  (fig.  characters  the 24)  from  study  raises  s i x to  are  23) r e q u i r e s  five  Wagner t r e e c a l c u l a t e d i n the absence of  and  t h i r t y - f o u r percent. this  by B o l i n (1947) ( f i g .  the  seventeen,  number  of  homoplasious  or from twelve percent to  In a d d i t i o n , when the c h a r a c t e r  mapped  onto B o l i n ' s  from  (1947) t r e e , s i x branches  remain unsupported.  Although  failing  my data as w e l l as my t r e e d i d , based on my  to represent  he  data  can  hardly  be  faulted  for  a n a l y s i s I cannot accept h i s h y p o t h e s i s of r e l a t i o n s h i p s . Compared the c h a r a c t e r  to  the Manhattan d i s t a n c e matrix c a l c u l a t e d from  data s e t , the Wagner tree has an R value  whereas B o l i n ' s t r e e has an R value tree  better  represents  of 0.45.  the data i n t h i s study.  The lies  in  Ruscar ius  (1944) c h a r a c t e r  main  difference  the  relationship  creaseri.  i d e n t i f i a b l e p a i r s of s i s t e r fenestralis-Artedius lateralis-Artedius  study  includes  between  h i s t r e e and my wagner tree Artedius  studies  corallinus. species  Bolin  harrinqtoni  recognize  species within  notospilotus,  h a r r i n q t o n i as the s i s t e r  Interestingly,  data.  between  Both  0.97,  C l e a r l y the Wagner  as f a r as c o u l d be determined from h i s t e x t , my most of B o l i n ' s  of  two  Artedius:  and  easily Artedius  and  Artedius  placed  Artedius  to A. c r e a s e r i , i n the absence  89  of  Ruscarius  meanyi.  A. h a r r i n q t o n i similar  and  reliable  have  justification  A. c r e a s e r i  distribution  characters  The  of  been  together  scales.  lay  discovered,  this  relationship.  The  present  The  recognized,  Artedius  two  pairs  01iqocottus  remaining cirri  Finally,  of A r t e d i u s s i s t e r  but A r t e d i u s harr i n g t o n i i s p l a c e d c l o s e s t  to  In the absence of  Artedius  entirely.  The  sensu  str i c t o .  to  be  removed including  the  formerly  sister  sister  s p e c i e s of A r t e d i u s sensu B o l i n , Orthonopias t r i a c i s , to a basal trichotomy  The  is  lineage  C l i n o c o t t u s i s hypothesized  Artedius  and  to  Chitonotus,  of are  group  removed  he  species  genus and  a  taxon of A r t e d i u s .  c o r a l l i n u s plus Artedius l a t e r a l i s .  the  not  no p r e o r b i t a l  R. meanyi, A. c r e a s e r i i s placed with Chitonotus from  the  a n a l y s i s o f f e r s q u i t e a d i f f e r e n t hypothesis  relationships. again  is  The  from R. c r e a s e r i and A. h a r r i n g t o n i .  placed Orthonopias as the s i s t e r  in  corroborative  character  A r t e d i u s s p e c i e s with reduced squamation and  placing  mainly  Since no other  i n d i c a t o r of p h y l o g e n e t i c  are separated  for  with Hemilepidotus,  I c e l i n u s , Ruscar i u s , O l i g o c o t t u s ,  hypothesized is  separate  from  Clinocottus,  and  Artedius• Contrary  to  Bolin's  (1947)  claim  c l e a r l y d e r i v e d from the A r t e d i u s l i n e " shows  that  the  characters  which  (p.  Bolin  Orthonopias with A r t e d i u s are plesiomorphic therefore  inadmissable  as evidence  that  "Orthonopias i s  162).  This  (1947) used to u n i t e at  that  level  f o r s i s t e r group s t a t u s .  a d d i t i o n , Orthonopias i s h i g h l y autapomorphic, e s p e c i a l l y form of the p e l v i c  fins  study  and In  i n the  in the male, the form of the body s c a l e s  90  and  i n the p a t t e r n of  yield  little  the other and  body  coloration.  taxa under c o n s i d e r a t i o n (1978)  i n t h i s study, although  suggestion  A r t e d i u s appears to be t o t a l l y  that  similar  root  together,  t h a t , w i t h i n the C o t t i d a e , there squamation.  This  assumption  w i t h i n the C o t t i d a e . trends  cannot  empirically particular utility the show  be  The  seem  is  missed  is  phylogenetic  hypothesis.  such a g e n e r a l i z a t i o n has  Artedius,  A.  lateralis  though admittedly forms  as  the  R. meanyi  as  reduction  that by  look  the  like  removal  separate  Orthonopias at the unresolved from A r t e d i u s .  of  that  such  In  and  i f any,  within  the placement, non-scaled  "Artedius" R.  genus group  base  These two  of  of  of  s p e c i e s which  totally  an  and this  placement  scalation  A. c o r a l l i n u s ,  01iqocottus-Clinocottus-Artedius  removed  species  lineage of the s c a l e d genus A r t e d i u s .  things  a  the  little,  Witness the  p r o v i s i o n a l , of two  evidenced to  of  and  still  sister  a d d i t i o n , not a l l Artedius,  degree  very  assumption  i n the absence of a l o g i c a l l y  " s c a l y " A r t e d i u s h a r r i n g t o n i with the the two highest  stem  i s a trend towards r e d u c t i o n in  in r e c o n s t r u c t i n g genealogy.  the  to  i n p l a c i n g forms with  i s then used to order  identified  instance,  i n c l u d e d in  on the a p r i o r i  point that  constructed  Howe  analysis is certainly  of the problem l i e s  s c a l a t i o n patterns  be  (1947) a n a l y s i s  from methodology, s i n c e h i s c h a r a c t e r The  it  of  unsupported.  shortcomings of B o l i n ' s  detailed.  characteristics  evidence for placement of Orthonopias with any  Richardson's  The  Such  creaseri  outside  and  the  the  tree  r e s u l t s provide  placement diagram  In are and the of far  an example of  91  the  futility  of grouping  by  overall  t r y i n g to e s t a b l i s h g e n e a l o g i c a l  similarity  when  one  is  relationships.  Larvae None  of  the  mininum-length t r e e s  from the Wagner a n a l y s i s are the Washington  (1982) ( f i g s .  supposedly  derived  same  will  resolved that  be  those  presented  from an "unrooted Wagner a n a l y s i s " . derived  from  r e f e r r e d to as the c o l l a p s e d  or dichotomous t r e e .  this  as  19, 20, and 21)  25 and 26), one of which ( f i g .  f o l l o w i n g comparisons, the two t r e e s study  (figs.  dichotomous  Inspection  of  relationships  25) i s In the larval  t r e e and the f u l l y her  t r e e has no c h a r a c t e r  the complete r e s o l u t i o n of  the  by  data  reveals  justification for  within  Artedius  and  Clinocottus. The  justification  mysterious. her  from  fully  including  A. c r e a s e r i  and  in  a  remains  depicted  lineage  in  separate  A. h a r r i n g t o n i .  Washington removed A. c r e a s e r i to a l i n e a g e A. meanyi,  tree  are i n f a c t those proposed by B o l i n in 1947, who  A. l a t e r a l i s and A. f e n e s t r a l i s that  resolved  I t i s p o s s i b l e that the r e l a t i o n s h i p s  cladogram  placed  f o r her  with  Since  Icelinus  and  t h i s would lead to the placement of A. l a t e r a l i s and  A. f e n e s t r a l i s A. h a r r i n g t o n i . clouds the i s s u e .  in  a  monophyletic  However, This  the  group  problematical  apart  from  Artedius  type 3  i s e i t h e r A. c o r a l l i n u s , A.  or a combination of both, and  i t s inclusion  in  notospilotus the  analysis  92  results  in  arising  Bolin  justification Clinocottus  (1947),  Washington other  type  3,  A. f e n e s t r a l i s and A. l a t e r a l i s  from a trichotomy.  The within  Artedius  character  (1982)  A. h a r r i n q t o n i A. l a t e r a l i s  the  does tree  data  in plus  and  resolution  branching  not for  of  sequence  relationships  correspond  this  depicted  a  genus.  A.  trichotomy type  3_.  In  including Cl inocottus  i n v o l v i n g C.  by  to h i s t r e e , the only the  the body of her t e x t , A r t e d i u s  acut i c e p s p l u s a trichotomy and C.  the  i n the dichotomous t r e e cannot be found with  since  published  for  absence  of  c o l l a p s e s to  A. f e n e s t r a l i s , c o l l a p s e s to C.  a n a l i s,  C.  embryum  recalvus.  As with the a d u l t t r e e of B o l i n , the goodness of f i t of the larval  tree  presented  that of the Wagner t r e e s  Table 1.  by Washington i s c o n s i d e r a b l y  l e s s than  (see t a b l e 1).  Comparison of f i t of l a r v a l  trees.  R-value  Wagner t r e e 1  96.96  Wagner t r e e 2  97. 1 0  Wagner t r e e 3  96.57  Washington  (1982) dichotomous  76.43  Washington  (1982) polytomous  68.99  One of the Wagner t r e e s ( f i g .  19) reproduces  exactly  the  93  branching sequence w i t h i n genera shown in Washington's c o l l a p s e d tree  (fig.  26).  Clinocottus  However, in my  emerge from a trichotomy and  Washington's t r e e . my  inability  she  of  or,  may  matrix  from  r e s o l v e d as i n  be o f f e r e d to e x p l a i n erred  Washington's  in the t e x t , but  in  the  text;  the  I coded them as  i f I coded the data c o r r e c t l y and sound, her  and  the  a n a l y t i c a l technique  original is  not,  a Wagner (parsimony) method.  First,  using  data  a n a l y s i s was  as claimed,  notable  a  are not  trees: I seriously  were p o o r l y analyzed  intended;  character  Three e x p l a n a t i o n s  to reproduce her  preparation characters  tree, Artedius, 01igocottus  Washington's  exception, outgroup  these are  Washington  analysis.  above  apparently  a n a l y s i s of the c h a r a c t e r s . (1982) Given  suspicion.  polarized  by  use  polarized the  One of  the  information  character, the  With  one  characters in the  number  text  11,  she  common-equals-primitive  criterion. "This  condition  (rounded  snout)  is  probably  p r i m i t i v e c o n d i t i o n r e l a t i v e to l a r v a e of A r t e d i u s , and  01igocottus,  divergent  of  C o t t i d s and  the p o i n t e d  snout appears  172).  seems,  It  to  be  Scorpaeniformes. a  derived  171). two  "snout  (therefore) (p.  though, that the a l t e r n a t i v e c h a r a c t e r  state  length  Unfortunately, that  several  condition."  a l s o e x h i b i t s a widespread d i s t r i b u t i o n , comment:  Clinocottus,  because i t i s widespread i n ( l a r v a e of)  genera  the  which  caused  her  i s v a r i a b l e in the outgroup taxa."  t h i s character  (snout  length)  is  one  to (p. of  i s used to j u s t i f y the monophyly of a group i n c l u d i n g  I c e l i n u s , A. c r e a s e r i , and  A. meanyi.  It  must  be  noted  here  94  that  the  shape of the  to snout  shape  analysis).  in  The  snout in l a r v a l  adult  other  forms  larval  (character character  group's monophyly (basal p r e o p e r c u l a r outgroup comparison, but A  single  character,  not  forms does not 2  correspond  in  the  adult  used to j u s t i f y  spine)  was  this  polarized  by  a l l I c e l i n u s s p e c i e s were examined.  p e l v i c f i n ray number, was  used to  justify  the group A. meanyi p l u s I c e l i n u s . Next l e t us consider the  data  offered  as  on  Washington  11 c h a r a c t e r s  (although the  the p o s s i b i l i t y (1982)  cladogram).  these,  The  is  little  analysis  doubt of  straightforward. coding  In  phylogenetic  the  in  for  interest in  her  the  binary  included  of  an  additional  tree. step  to  intended. also  alternative  reproduce  the  These i n v a r i a b l y relative  to  the  trees  had  the same  topology as the t r e e s in Washington's t e x t .  I can  only  conclude  Wagner coding  scheme w i l l y i e l d her  analysis. of her  Although  I  of  to  able  was  fairness,  attempt  and  characters,  characters  collapsed  None  (1982)  these  that no coding  scheme.  code  analysis  of these were not  multistate  tried,  sequence  coding  Washington  that these were coded as she  r e s u l t e d in a t r e e with one original  not  seven are b i n a r y c h a r a c t e r s  state  the  schemes were  branching  did  Again assuming that Washington was  determine the plesiomorphic there  the  s t a t e d , two  Of  four are m u l t i s t a t e .  intended.  to be used in  f o r reasons not  that I  I  am  t r e e when  satisfied,  data r e s u l t s i n the most can  genera as i n her  reproduce  (fig.  collapsed tree  19)  (fig.  though,  subjected that my  parsimonious the 26)  a  original  cladogram.  same topology there does  to  not  within seem  95  to  be  support  i n her  character  data  O l i g o c o t t u s and C l i n o c o t t u s as s i s t e r Although Richardson's relationships  within  no  diagram,  branching  Ruscarius  (1981)  f o r the placement of  taxa.  discussion  of  intergeneric  the c o t t i d s was p r e l i m i n a r y and presented Washington  meanyi was i n c l u d e d  (1982)  pointed  out  that  i n Richardson's (1981) study under  " I c e l u s spp.", which makes p o s s i b l e a comparison of Richardson's (1981) phenetic of  her  groupings with the r e s u l t s from t h i s study.  groups  are  of  particular  interest:  Artedius-01igocottus-Clinocottus-Orthonopias group  including  points  of  Chitonotus,  agreement  Icelus  emerge:  Artedius~Clinocottus-01igocottus  and  the group  separation from  notion that R. meanyi does not belong with All  three  phylogenetic  of  the  larval  (1) the  group and Icelinus.  Two  (2) the Two major of  Icelinus  the  and the  Artedius.  analyses  (one p h e n e t i c ,  one  i n name only and one Wagner) agree that R. meanyi  (or R. meanyi and common ancestor  R. c r e a s e r i )  with  does  not  the r e s t of A r t e d i u s  share  sensu  an  immediate  stricto.  96  CONGRUENCE OF Figure  27  LARVAL AND  shows  the  ADULT CLASSIFICATIONS  consensus  tree  consensus of the a d u l t Adams t r e e and (note  that  in the  f i n a l consensus).  from  the  the  derived larval  Adams  only those taxa common to both analyses The  i n a b i l i t y of the  l a r g e amount  of  from  are  Despite which merit recognize  in  that  trend  the  discussion. the  Both  closest  or O l i g o c o t t u s of the  in c o t t i d  toward  adult  is  In the context  may and  of my  characters  that  does  limited.  r e l a t i v e s of  have  scalation  set;  dermal  the  reduction  pattern  the  same  is  This  in  is  family  a  similar  there  the c o t t i d s characters  hypothesized to be c l o s e l y r e l a t e d to the  that  Oligocottus  Artedius, latter.  no As a  C l i n o c o t t u s , are  Orthonopias,  has  are  distribution.  and  to  unsealed  of t h i s trend  hypothesizes  (although  general  (see B o l i n ,  polarizing  study  prickles).  generally  important  A widespread  of s c a l e s w i t h i n  i t s use  Artedius bony  there  trees  is either  grouped with other  completely unsealed genera, 01igocottus closest  This  r e l a t i o n s h i p s , as  with  be a u s e f u l concept, but is  Adams  Artedius  a n a l y s i s , the use  in s o r t i n g out  r u l e of thumb the  taxa  of  of s c a l e s w i t h i n  forms.  limited u t i l i t y  larval  l a t t e r genera i s s c a l e d .  Unsealed forms are t h e r e f o r e  general  data  areas of agreement  plus C l i n o c o t t u s .  systematics  reduction  and  relative  1947).  corroborative  stems  l a r v a l t r e e to r e s o l v e r e l a t i o n s h i p s .  shortcomings, there are two  neither  assumption  present  are needed.  that  01igocottus  tree,  amgibuity  T h i s ambiguity i s a l s o a f u n c t i o n of the s i z e of the more c h a r a c t e r s  the  two the  rimensis  which has is  a  not  97  The  second  major  area of agreement between the a d u l t and  l a r v a l Adams t r e e s i s the h y p o t h e s i s that Ruscar ius Ruscarius  creaseri,  formerly i n A r t e d i u s , do not i n f a c t  with the r e s t of the nominal i s unable  Washington's  her  of Ruscar i u s .  of R. meanyi,  R. c r e a s e r i ,  i t d i d serve to focus a t t e n t i o n on a c e n t r a l  s y s t e m a t i c s : monophyly. study  is  With r e s p e c t to  instructive.  a t t e n t i o n on the lack of Artedius•  At  the  tree  (1982) a n a l y s i s c o n f l i c t e d with the  a d u l t a n a l y s i s i n i t s placement Icelinus,  and  belong  A r t e d i u s , although the l a r v a l  to completely r e s o l v e the placement  Although  meanyi  The  methodology,  larval  justification  and  issue i n however,  study served to focus  f o r the  monophyly  of  l e v e l at which there was c h a r a c t e r support,  Washington's study d i d recognize that R. meanyi and  R. c r e a s e r i  d i d not belong with the r e s t of A r t e d i u s sensu s t r i c t u , which i s something  that the e v o l u t i o n a r y taxonomic study of B o l i n  (1947)  f a i l e d to uncover. A strict (1981)  congruence study using the r e s u l t s  was  not  possible,  due  i d e n t i f i c a t i o n and the d i f f e r e n t worth  noting:  adult  and  larval  One  i s not  point  in is  suggested  monophyletic,  a  cladograms remove R. c r e a s e r i and  R. meanyi from A r t e d i u s sensu s t r i c t u .  test  uncertainties  taxa examined.  R. meanyi  Richardson  supported by the a n a l y s i s based on a d u l t c h a r a c t e r s .  Both  nature  the  even t h i s a d m i t t e d l y phenetic grouping  that an A r t e d i u s i n c l u d i n g result  to  of  of  much  for further  Due  of the l a r v a l cladogram, congruence.  Still,  to  the  unresolved  i t i s not p o s s i b l e to  in  the  area  that jjs  r e s o l v e d i n both l a r v a l and a d u l t a n a l y s e s , there i s congruence.  98  In  the  presence  of  a great d e a l of noise i n the l a r v a l  phylogenetic analysis  provided  the  placement  of  Ruscarius  analysis.  The  only other systematic  its  relatives  relative  (Bolin,  1947)  same  hypothesis  data,  for  the  to A r t e d i u s as d i d the a d u l t treatment  of  Artedius  d i d not recognize the  and  diphyletic  c o n d i t i o n of A r t e d i u s . Congruence comparing  of  classifications  theories  of  p r e d i c t i o n s regarding theories.  can  regarding  of the l a r v a l  (1982), l i k e De  predictions.  They  predominately  to the  490).  natural  I_f  morphological environment  than  the  that  action  larvae  different strict  natural  selection should with  alone be  of  (1958)  characters  the  obvious  Charlesworth any  specific  p a t t e r n s are  selection shapes  If  those  Beer  larval  make  better  phylogeny.  (1982,  the  s e l e c t i v e regimes, we  then  with  compare  l i f e h i s t o r y stages  such  would not p r e d i c t congruence.  " s e l e c t i v e regime" e x p l a n a t i o n grows more convoluted found.  The  Wiley  (1983,  also  and  p.  organism,  correlated we  due  and a d u l t s , which are presumably subjected to v a s t l y  time congruence i s  predict  De  phylogenetic  of  explicit  there seems to be  face  Beer (1958), do not  in  from  forms to t h e i r h a b i t a t s .  state  change  in  when  derived  congruence.  c l a s s i f i c a t i o n s d e r i v e d from d i f f e r e n t as  be  that c l a s s i f i c a t i o n s based on a d u l t and  adaptation  important  change  Within c u r r e n t e v o l u t i o n a r y theory  were most o f t e n congruent, even  et a l .  also  evolutionary  congruence  a range of statements found  is  Brooks  congruence,  since  it  non-equilibrium Brooks  and  explicitly  framework  Wiley,  1983)  recognizes  A  each of does the  99  constraining  r o l e of h i s t o r y .  of d i f f e r e n t data E v e n t u a l l y we  sets r e f l e c t i n g will  the p a t t e r n of order notions of  Congruence i s then a consequence  in  reach a p o i n t where our nature  about c a u s a l processes.  process  the same phylogenetic  forces  us  E i t h e r we  to  history.  understanding of re-examine  r e s t r u c t u r e our  our ideas  to accomodate f e a t u r e s of the n a t u r a l world such as  congruence, or we  abandon our  former ideas e n t i r e l y  something that b e t t e r accounts f o r the  data.  in favor  of  100  SUGGESTIONS FOR  Now  that  strictu,  a  there  examined.  phylogeny are  other  Firstly,  we  FUTURE STUDIES  is  available  important  can  map  shape  and  size.  g e n e r a l i z e d to t e s t  This  Artedius  questions  that  morphometric  cladogram i n order to i d e n t i f y h i s t o r i c a l body  for  mapping technique  changes  can  a  might be c a l l e d cladogram  " h i s t o r i c a l ecology".  makes  relationships,  possible  leading  to  in fact  in be  respect  to  The a v a i l a b i l i t y of  analysis  studies  be  character.  Other types of analyses are a l s o p o s s i b l e with what  can  data onto t h i s  trends i n  f o r trends i n any continuous  sensu  of  host-parasite  of c o e v o l u t i o n i n g e n e r a l .  With t h i s cladogram, f i e l d e c o l o g i s t s  now  background f o r s t u d i e s of s p e c i a t i o n .  Community e c o l o g i s t s have  a  vital  piece  of  information  have  necessary  in  the  necessary  estimating  h i s t o r i c a l component of e c o l o g i c a l a s s o c i a t i o n s .  Biogeographers  can study the r e l a t i o n s h i p between the h i s t o r y of these the h i s t o r y of the areas In the l i g h t of the higher-level  question  in which they situation  in  In how  evolutionary-taxonomic  can reasonably be asked the  problem  interrelationships  at  of any hand  of  and  live. Artedius,  the  obvious  many of the other  has a robust estimate of phylogeny remained obscure, of  fish  a r i s e s : which other c o t t i d genera are i n  need of p h y l o g e n e t i c a n a l y s i s ?  face  the  study?  taxon, it  Icelinus,  Of course  even in the  t h i s question  not j u s t c o t t i d s .  seems  critical  Chitonotus,  genera  that  Within the  Ruscar i u s ,  0 1 i g o c o t t u s , C l i n o c o t t u s and A r t e d i u s be examined, e s p e c i a l l y i n  101  the  light  of  the  meagre  evidence  for  A r t e d i u s , C l i n o c o t t u s and 0 1 i g o c o t t u s . commonly been grouped been  forthcoming.  the r e l a t i o n s h i p s of  These three genera  have  together, but l i t t l e c h a r a c t e r support Icelinus  is  probably  next  in  line  has for  thorough study, f o l l o w e d by C l i n o c o t t u s and 0 1 i g o c o t t u s . Finally, examination,  other since  not a very r e l i a b l e Our  knowledge  and western benefit  of  scaled  genera  i t i s becoming apparent indicator  of  phylogenetic  suggestions  analytical  Icelus  that  "scaliness" is  intergeneric  merit  relationships.  to  support.  that  those i n S o v i e t waters,  analysis.  C o t t i d a e as i t stands today i s not but  as  v i r t u a l l y a l l of the c o t t i d s i n the northern  Pacific, particularly  from  such  a  would  I t i s p o s s i b l e that the monophyletic  assemblage,  e f f e c t have as yet been made without  102  LITERATURE CITED  Adams, E.N. 1972. Consensus techniques and the comparison of taxonomic t r e e s . S y s t . Z o o l . 21:390-397. Andersen, N.M. 1979. P h y l o g e n e t i c i n f e r e n c e as a p p l i e d to the study of e v o l u t i o n a r y d i v e r s i f i c a t i o n of semiaquatic bugs (Hemiptera: Gerromorpha). S y s t . Z o o l . 28:554-578. Ayres, W.O. 1855. 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I d e n t i f i c a t i o n and s y s t e m a t i c s of l a r v a e of A r t e d i u s , C l i n o c o t t u s , and 01igocottus ( S c o r p a e n i f o r m e s : C o t t i d a e ) . M.Sc. T h e s i s , Oregon State U n i v e r s i t y . 192 pp. Wiley,  E.O. 1975. K a r l R. Popper, s y s t e m a t i c s , and c l a s i f i c a t i o n : a r e p l y to Walter Bock and other e v o l u t i o n a r y taxonomists. S y s t . Zool. 24:233-242.  Wiley,  E.O. 1978. the e v o l u t i o n a r y s p e c i e s concept S y s t . Z o o l . 27:17-26.  Wiley,  E.O. 1979. An annotated Linnean h i e r a r c h y , with comments on n a t u r a l taxa and competing systems. S y s t . Z o o l . 28:308337.  Wiley,  E.O. 1981. Phylogenet i c s . The Theory and Pract i c e of Phylogenetic Systematics. John Wiley and Sons, New York. 439 pp.  Wiley,  E.O. 1982. Convex groups and c o n s i s t e n t S y s t . Botany 6:346-358.  Wiley,  E.O., and D.R. Brooks. 1982. V i c t i m s of h i s t o r y - a n o n e q u i l i b r i u m approach to e v o l u t i o n . S y s t . Z o o l . 31:1-24.  reconsidered.  classifications.  Wilimovsky, N.J. 1954. L i s t of the f i s h e s of A l a s k a . I c h t h . B u l l . 4(5):279-294.  Stanford  Wilimovsky, N.J. 1958. P r o v i s i o n a l keys to the f i s h e s of A l a s k a . U.S. F i s h and W i l d l i f e Serv. MS. 113 pp. Wilimovsky, N.J. 1963. Inshore f i s h fauna of the A l e u t i a n a r c h i p e l a g o . Proc. f o u r t e e n t h Alask. S c i . Conf.: 172-190.  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 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0  0 1 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 0 0 1 0 0 0 0  o  1 1 0 0 0 0  Hemilepidotus Orthonopias  1 0 1 2 1 1 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 2 1 1 0 0 0 0 0 1 0 0 1  R.meanyi  0 0 1 2 1 1 1 0 2 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 1 0 1 2 0 0 0 0 0 0 0 0 0 0 0  R.ereaseM  0 0 2 1 0 1 0 0 1 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 1 2 0 0 0 0 0 0 0 0 0 0 1  Chitonotus  0 0 0 1 0 0 0 1 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 2 0 0 0 0 0 0 0 0 0 0 0  IeelInus  0 0 3 3 0 0 0 0 0 0 0 1 1 1 1 1 2 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 0 0 2 2 0 0 0 0 0 0 0 1 t 1 0  A.fenestra!4s  0 0 3 3 0 0 0 0 0 1 0 1 1 1 1 0 0 1 1 0 0 2 0 0 0 0 1 0 0 0 0 1 0 0 2 2 0 0 1 1 0 0 0 1 1 1 0  A.notospilotus  0 0 4 3 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 1 1 1 1 1 1 0 1 0 1 0 2 0 0 1 2 2 0 0 0 0 1 0 0 0 1 1 0  A.herrtngtont  0 0 4 3 0 0 0 0 0 0 0 3 0 0 0 0 1 0 0 2 0 1 0 0 0 1 1 0 1 0 0 1 0 0 2 2 0 0 0 0 2 0 0 0 2 1 0  A.coral 1inus  0 0 4 3 0 0 0 0 0 0 0 3 0 0 0 0 1 0 0 0 0 2 0 0 0 0 1 0 0 0 0 1 0 0 2 2 0 0 0 0 2 0 0 0 2 1 0  A . l a t e r a l is  0 0 S 4 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 3 3 0 0 0 0 0 0 0 0 0 1 0  01igocottus  0 0 3 4 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 3 3 0 0 0 0 0 0 0 0 0 1 0  Clinocottus  > W Z O •-<  M  > a a r  >  > W  APPENDIX 2.  LARVAL DATA MATRIX  0  0  0  0  0  0  0  0  0  0  Ancestor  0  0  1  0  0  0  O  1  0  1  Icelinus  0  0  1  0  0  0  0  1  0  1  Meanyi  0  0  1  0  0  0  0  1  0  0  Creaser1  1  2  0  0  0  0  2  0  1  0  A.harr1ngtoni  1  2  0  0  0  2  1  0  1  0  A . l a t e r a l 1s  1  2  0  0  0  2  1  0  1  0  A.fenestra 1 1 s  1  2  0  0  0  2  1  0  1  0  A.type 3  1  1  0  2  0  0  1  0  1  0  C.globiceps  1  1  0  2  o  0  0  0  1  0  C.embryum  1  1  0  2  0  0  0  0  1  0  C.anali s  1  1  0  2  0  0  0  0  1  0  C.recalvus  1  1  0  1  0  0  2  0  1  0  C.acut iceps  1  1  0  0  1  1  1  0  1  0  0.snyder i  !  o  0  1  1  1  o  !  0  0.maculosus  1 17  HE STUDENT WANTED TO LEAVE A BLANK PAGE BETWEEN THE END OF THE TEXT AND THE BEGINNING F THE APPENDIX MATERIAL,  118  Figure  1.  Artedius c o r a l l i n u s after Bolin  (1944).  119  1 20  F i g u r e 2.  A r t e d i u s f e n e s t r a l i s from B o l i n  (1944).  Fig. 2  122  F i g u r e 3.  Artedius harringtoni  after Bolin  (1944).  123  Fig.  3  1 24  Figure 4.  Artedius l a t e r a l i s  from B o l i n  (1944).  125  Fig. 4  1 26  Figure 5.  A r t e d i u s n o t o s p i l o t u s from B o l i n (1944).  1 28  F i g u r e 6.  Ruscarius c r e a s e r i  from B o l i n  (1944).  129  Fig. 6  130  F i g u r e 7.  Ruscar ius meanyi from Jordan and Evermann  (1895).  131  Fig. 7  1 32  F i g u r e 8.  Orthonopias t r i a c i s  from B o l i n  (1944).  Fig. 8  134  F i g u r e 9. Side view of s c a l e r i d g e s t a t e 0. F i g u r e 9b, s t a t e 1. F i g u r e 9d, s t a t e s 3 and 4.  (schematic). F i g u r e 9a, F i g u r e 9c, s t a t e 2.  F i g . 9c  F i g . 9d  135a  Ctenii  r  F i g . 9b  1 36  Figure  10. Chin pigmentation p a t t e r n s . . F i g u r e 10a, A r t e d i u s f e n e s t r a l i s (CAS 40354, 55.1 mm). F i g u r e 10b, A r t e d i u s n o t o s p i l o t u s (SIO 63-1054, 56.0 mm). Figure 10c, A r t e d i u s h a r r i n g t o n i (CAS 29521, 62.9 mm). Figure I0d, A r t e d i u s l a t e r a l i s (UMMZ 42958, 62.9 mm). Figure I0e, Orthonopias t r i a c i s (SU 18132, 54.8 mm). Areas outlined in f i g s . I0a-I0d i n d i c a t e l i g h t background pigmentation. Areas o u t l i n e d i n f i g . 1Oe i n d i c a t e white pigment.  Fig. 10a  Fig. 10b  137b  Fig. 10c  137c  Fig. 10d  137d  Fig. 10e  1 38  Figure  11. Top view of s c a l e ridge (schematic). Figure 11a, Hemilepidotus hemilepidotus. F i g u r e 11b, Chitonotus p u g e t e n s i s . F i g u r e 11c, Ruscarius c r e a s e r i . F i g u r e 11d, Ruscarius meanyi. F i g u r e 11e, A r t e d i u s fenestralis. F i g u r e 11f, A r t e d i u s n o t o s p i l o t u s . F i g u r e 11g, A r t e d i u s h a r r i n g t o n i . F i g u r e 11h, A r t e d i u s lateralis.  139a  Fig. 11d  139b  F i g . 11f  1 40  Figure  12. Pterotic flange. F i g u r e 12a, Chitonotus pugetensis (UW uncat., 46.1 mm). F i g u r e 12b, A r t e d i u s f e n e s t r a l i s (BC57-210, 55.5 mm). F i g u r e 12c, A r t e d i u s c o r a l l i n u s (CAS 48970, 62.3 mm). A b b r e v i a t i o n s : PT p t e r o t i c , PA - p a r i e t a l , EP - e p i o t i c , SO s u p r a o c c i p i t a l , EX - e x o c c i p i t a l , TB - t a b u l a r s , SP sphenotic, EC - ethmoid c a r t i l a g e , VO - vomer, NA n a s a l , ME - mesethmoid, LE - l a t e r a l ethmoid, FR frontal.  Fig. 12a  141a  Fig. 12b  Fig. 12c  1 42  Figure  13.  Adult Wagner t r e e  1.  1 44  Figure  14.  Adult Wagner t r e e  2.  ''<3  •''6 *4 /  .  \  V  % "a.  o  146  Figure  15.  Adult Wagner t r e e  3.  147  Figure  16. Cladogram of A r t e d i u s sensu s t r i c t u . C h a r a c t e r s : 3 - form of s c a l e r i d g e . 4 - body c o l o r pattern. 10 - s p i n e l e t s on main p r e o p e r c u l a r s p i n e . 12 - c h i n c o l o r a t i o n . 13 - mandibular pore p a t t e r n . 14 - pores on l a t e r a l l i n e s c a l e s . 15 - form of head scales. 16 - c i r r i on t r a n s v e r s e head t u b e r c l e s . 17 - c i r r i on s u b o r b i t a l stay. 18 - form of p r e o p e r c u l a r spines. 19 - nasal pores. 20 - form of t e e t h . 21 b r a n c h i o s t e g a l number. 22 - number of s c a l e rows above lateral line. 23 - anal f i n membrane. 24 - anal f i n pigmentation. 25 - p e n i s . 26 - c i r r i on upper l i p . 27 - p o s t c l e i t h r a . 28 - s c a l e s under a n t e r i o r o r b i t . 29 - s c a l e s behind a x i l l a . 30 - s c a l e s on caudal peduncle. 31 - p r e o r b i t a l c i r r i . 32 - c i r r i above axilla. 34 - c i r r i on p r e o p e r c u l a r margin. 35 s c a l e r i d g e shape. 39 - form of head t u b e r c l e s . 40 s e r r a t i o n s on p o s t t e m p o r a l / s u p r a c l e i t h r u m . 41 - s i z e of c i r c l e s on body. 45 - p t e r o t i c f l a n g e .  149  1 50  Figure  17. Cladogram of R u s c a r i u s . Characters: 1 p o s i t i o n of anus. 3 - form of s c a l e r i d g e . 4 - body color pattern. 5 - s c a l e s above a x i l l a . 6 - scales on snout. 7 - shape of upper p r e o p e r c u l a r s p i n e . 8 number of p e l v i c r a y s . 9 - c i r r i on o p e r c l e . 28 s c a l e s under a n t e r i o r o r b i t . 31 - p r e o r b i t a l c i r r i . 32 - c i r r i above a x i l l a . 33 - c i r r i anterad to upper preopercular spine. 35 - s c a l e r i d g e shape. 37 adult s i z e . 38 - anal ray form (males). 44 - c i r r i on n a s a l s p i n e . 47 - s c a l e s on eye.  1 52  Figure  18.  Adams consensus t r e e f o r a d u l t  trees.  153  1 54  Figure  19.  L a r v a l Wagner t r e e  1.  155  1 56  Figure  20.  L a r v a l Wagner t r e e  2.  157  1 58  Figure 21.  Larval. Wagner tree 3.  159  160  Figure  22.  Adams c o n s e n s u s t r e e  for l a r v a l  trees.  161  Figure 23.  Phylogenetic  t r e e of B o l i n (1947)  163  1 64  Figure 24. Wagner t r e e c a l c u l a t e d i n the absence of R u s c a r i u s meanyi.  1 66  Figure 25.  Dichotomous l a r v a l t r e e of Washington  (1982).  1 6 7  168  Figure 26.  Polytomous l a r v a l t r e e from Washington (1982).  169  1 70  F i g u r e 27. Adams consensus t r e e of a d u l t and l a r v a l Adams trees.  171  

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