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The genus Martes (Mustelidae) in North America: |b its distribution, variation, classification, phylogeny… Hagmeier, Edwin Moyer 1955

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THE GENUS MARTES (MUSTELIDAE) IN NORTH AMERICA: ITS DISTRIBUTION, VARIATION, CLASSIFICATION, PHYLOGENY AND RELATIONSHIP TO OLD WORLD FORMS  by Edwin M. Hagmeier  A Thesis Submitted i n P a r t i a l Fulfilment of the Requirements f o r the Degree of DOCTOR OF PHILOSOPHY i n the Department of Zoology  We accept this thesis as conforming to the standard required from candidates f o r the degree of Doctor of Philosophy  Members of the Department of Zoology  THE UNIVERSITY OF BRITISH COLUMBIA September  1955  F a c u l t y  o f  G r a d u a t e  P R O G R A M M E  Jffinai  S t u d i e s  O F  T H E  ^xxatttinatimt for ttye of Jioxtor of  ]^t$xtz  ^Irilrfsoijliy  EDWIN HAGMEIER B . A . M . A . A U G U S T I N  T H E  ( Q u e e n ' s ) 1 9 5 0  ( B r i t i s h C o l u m b i a ) 1 9 5 2 8 t h ,  1 9 5 5 , a t  B I O L O G I C A L R o o m C O M M I T T E E H .  I . M c T . ,"  3 : 0 0  S C I E N C E S  p . m . B U I L D I N G  1 0 7 A I N  F . ANGUS,  C H A R G E Chairman  COWAN  V . J . OKULITCH  H . C . GUNNING  P . FORD  K. GRAHAM  J . G . SPAULDING  G . J . SPENCER  V.  External  Examiner  N a t i o n a l M u s e u m  —  KRAJINA  R . J . RUSSELL  o l  C a n a d a  ABSTRACT T h e  G e n u s Martes  ( M u s t e l i d a e ) i n N o r t h A m e r i c a : i t s D i s t r i b u t i o n ,  V a r i a t i o n , C l a s s i f i c a t i o n , P h y l o g e n y t o O l d  T h r e e T h e  M. foina,  M.  martes,  o f o n l y o n e , M.  melampus  pennanti;  a n d M.  T w o  melampus r e a s o n  M.  t h e s e c o n d , Pekania,  americana,  o f o n e , M.  U n t i l r e c e n t l y M. americana  r e c e n t w o r k  a n d M. caurina,  o f P . L . W r i g h t  a r e d i s t i n c t i v e m o r p h o l o g i c a l l y ,  M.  martes,  M.  a l l b e l o n g  t o o n e  b e e n  c o n s i d e r e d  o n e  zibellina  s p e c i e s .  i n t e r g r a d e  s p e c i e s , c o n s i s t i n g o f  a n  w a sc o n s i d e r e d t o c o n s a n d t h i r t e e n o r s o s u b s p e  i n d i c a t e s t h a t w h i l e t h e y  americana  t h e t w o  a t t h e p o i n t  " s p e c i e s "  w h e r e  r a n g e s m e e t a n d m u s t b e c o n s i d e r e d a s i n g l e s p e c i e s . Martes h a s  flavigula  a r e s o c l o s e l y r e l a t e d m o r p h o l o g i c a l l  t o b e l i e v e t h a t t h e y  t w o s p e c i e s , M. americana T h e  zibellina;  s p e c i e s o c c u r i n N o r t h A m e r i c a , n a m e l y M.  M. pennanti-  M.  guatkinsi.  W i t h i n t h e s u b g e n u s Martes,  g o o d  t h e w o r l d t o d a y .  a n d t h e t h i r d , Charronia,  w i t h s o m e t i m e s a s e c o n d , M.  t h e r e a p p e a r s  e x i s ti n  c o n s i s t s o f t h e f o l l o w i n g s p e c i e s :  M.  a n d p o s s i b l y M.  R e l a t i o n s h i p  F o r m s  t h e g e n u s Martes  s u b g e n e r a o f  f i r s t , Martes,  W o r l d  a n d  t h r e e  s u b s p e c i e s .  t h e i r pennanti  T h e  c o n c e p t  satisfactory. c o n t i n u a , T h i s ,  It  o f  t h e  lacks  s u b s p e c i e s p r o v e s  reality,  i t p o s s e s s e s n o  t o g e t h e r w i t h  it involves  l o w e r  m a n y  the  l i m i t , a n d  t h e c l i n a l n a t u r e  l e a d s t o t h e c o n c l u s i o n  i n  r e s p e c t s t o  arbitrary  o f  u n -  partitioning  of  i t i s d e t e r m i n e d  o f v a r i a t i o n i n  t h a t m a r t e n  b e  t h e  N e w  d e d u c t i v e l y .  m a r t e n  W o r l d  a n d  s h o u l d  f i s h e r b e  c o n -  s i d e r e d a s r e p r e s e n t e d b y o n l y t w o s u b s p e c i e s ( M . a. americana a. caurina), T h e  t h ef i s h e r b y o n e s p e c i e s , a n d n o n a m e d s u b s p e c i e s . d i s t r i b u t i o n o f  d i s t r i b u t i o n p r e c i s e i n  o f  t h e  m a r t e n  n o r t h e r n  f i s h e r t h a n  i n  a n d  f i s h e r c o r r e s p o n d s  e v e r g r e e n  f o r e s t s . T h e  o f b o t h  a r e k n o w n i n g , t w o a n d  o f  t h e m  o r  t h e  A m e r i c a .  N o r t h i n  t h e  t h e  o f  t h e  i c e s h e e t , a n d  A l a s k a  o f t h e s e r e f u g i a e x c e p t a m e l i o r a t i o n  N e w  t h e y  a n d  o f ) , o f w h i c h t h a t  ( o r e v o l v e d  i s l e s s  f o s s i l s p e c i e s  f i v e a r e s t i l ll i v m o d e r n  m a r t e n s  t h e r e ) l a t e i n  t h e  e n v i r o n m e n t s  i n  P l e i s t o c e n e . f o u n d  e a s t e r n U n i t e d C o a s t Y u k o n .  a n d  t o  t h e  h a b i t a b l e S t a t e s , t h e  C a s c a d e F i s h e r  t h e A l a s k a - Y u k o n  m i g r a t e d  T w e n t y - e i g h t  I t a p p e a r s  W o r l d  P l e i s t o c e n e m a r t e n  i c e s h e e t , t h e  d i s t r i b u t i o n  t h e  a r e f i r s t r e c o r d e d f r o m  W o r l d s .  i n  f o r e s t r e f u g i a o f s o u t h  s o u t h  N e w  a r e d i s p o s e d  e a r l y i n  D u r i n g  a n d  s y n o n y m s  f i s h e r s a r r i v e d  P l i o c e n e  t h e  t h e O l d  ( w h e n  c l o s e l y t o  m a r t e n .  F o s s i l s r e f e r a b l e t o t h e g e n u s Martes M i o c e n e  a n d  M o u n t a i n s  p r e s u m a b l y  o n e .  r e g i o n s  R o c k y  o f  W i t h  M o u n t a i n s s o u t h  o c c u r r e d  o f i n  t h e a l l  p o s t - g l a c i a l c l i m a t i c  t h e i r p r e s e n t  o c c u r r e n c e .  t h e  GRADUATE STUDIES F i e l d o f S t u d y :  Z o o l o g y  V e r t e b r a t e Z o o l o g y —  I .M c T . C o w a n  I n v e r t e b r a t e Z o o l o g y — O t h e r  W . A . C l e m e n s  S t u d i e s : V e r t e b r a t e P a l a e o n t o l o g y —  J .T . G r e g o r y  ii  TABLE OF CONTENTS Page LIST OF ILLUSTRATIONS  V  INTRODUCTION  1  CLASSIFICATION  6  THE GENUS MARTES  6 6  HISTORY OF CLASSIFICATION TAXONOMIC POSITION  12  THE SUBGENERA OF MARTES  14  HISTORY OF CLASSIFICATION  14  TAXONOMIC CHARACTERS  16 18  THE SPECIES OF MARTES HISTORY OF CLASSIFICATION  18  In Eurasia  18  In North America  22  Marten  22  Fisher  32  SYNOPSIS OF THE SPECIES AND SUBSPECIES OF MARTEN .. 37 Martes martes  41  Martes foina  46  Martes z i b e l l i n a  .  53  Martes melampus  62  Martes americana (including caurina)  65  Martes pennanti  69  Martes f l a v i g u l a  70  RELATIONSHIPS OF THE MARTEN  79 /continued  iii CLASSIFICATION (continued)  page 90  THE MARTENS OF THE NEW WORLD LIFE HISTORY  90  VARIATION  94  Nongeographic v a r i a t i o n  94  Methods of study of geographic v a r i a t i o n . 106 110  MARTENS Distribution  110  Synoptic examination  129  "Martes americana"  130  "Martes americana americana  130  "Martes americana brumalis"  143  "Martes americana atrata"  148  "Martes americana a b i e t i c o l a "  153  "Martes americana actuosa" (including"boria")  159  "Martes americana kenaiensis"  173  "Martes americana abietinoides"  178 187  "Martes caurina" "Martes caurina caurina"  188  "Martes caurina origenes"  203  "Martes caurina humboldtensis"  208  "Martes caurina sierrae"  212  "Martes caurina  219  Vancouverensis"  "Martes caurina nesophlla"  223  Summary  229  "Species" of marten  230 /continued  iv CLASSIFICATION (continued)  Page  FISHERS  242  Distribution  242  Synoptic examination  262  "Martes pennanti pennanti"  262  "Martes pennanti columbiana"  274  "Martes pennanti p a c i f i c a "  2 9 7  CLASSIFICATION OF MARTEN AND FISHER .. PHYLOGENY  289 .. .  298  PALAEONTOLOGY  298  BIOGEOGRAPHY  312  INTRODUCTION  312  REFUGIA OF THE WISCONSIN  327  POST-WISCONSIN MIGRATION  340  MARTEN AND FISHER  342  SUMMARY  348  LITERATURE CITED  354  APPENDICES  439  BRONGERSMA'S INDICES  439  SYNONYMY  441  CRANIAL MEASUREMENTS  463  SOURCES OF LOCALITY RECORDS  466  ABSTRACT  468  V  ILLUSTRATIONS Figures 1 and 2. 2, 3» 4, 5 and 6. 7. 8. 9.  Facing Page The skulls of the species of the world's marten  42  The d i s t r i b u t i o n of Eurasian marten  42  Ratio of canine width to b u l l a length i n the skulls of the world's marten  86  Pie diagrams depicting certain c h a r a c t e r i s t i c s of the skulls of the world's marten  88  Evidence of intergradation between "Martes americana" and "Martes caurina" i n Idaho and Montana, after Wright 1953  235  10.  Evidence of intergradation between "Martes americana" and "Martes caurina" i n Manning Park, B r i t i s h Columbia 235  11.  D i s t r i b u t i o n of the boundary separating "Martes americana" from "Martes caurina"  236  12.  The d i s t r i b u t i o n of the f o s s i l species of marten  308  13.  14.  A hypothesized phylogeny of the Recent species of marten  312  North America i n Mankato time  327  15. 16, North America during the Pleistocene and 17. 18. The d i s t r i b u t i o n of Pleistocene forest f o s s i l s . . 19. The r e l a t i v e densities of marten i n North America 20. Pdst-Mankato migration of marten and f i s h e r i n North America 21.  V a r i a t i o n i n M. a. americana and i t s possible r e l a t i o n to climate  ENCLOSURE A. Geographic d i s t r i b u t i o n of condylobasal length i n marten. ENCLOSURE B. D i s t r i b u t i o n of marten i n North America ENCLOSURE C.' D i s t r i b u t i o n of f i s h e r i n North America  32? 329 333 346 348  1  INTRODUCTION The genus Martes constitutes the fishers and  martens.  Seven or eight species are considered to occur i n the world today, including the pine and stone marten, the sable, yellow b e l l i e d and Japanese marten i n the old world, and the American marten and f i s h e r i n the new.  No revisionary work has been  done on the new world forms i n h a l f a century, and i t i s the purpose of t h i s paper to f i l l the need.  Yeager (1941) points  out that of the fur bearing mammals he considered, only badger, otter and raccoon have received less attention i n the l i t e r a t u r e than the wolverine, f i s h e r and marten together, so that i t i s apparent that much future work i s needed before these animals can be considered r e l a t i v e l y well understood. Myers (1952) suggests that systematic papers should make the problems being solved c l e a r , hence the following statement of the problem.  This i s an attempt to define the l i m i t s  of the genus concerned, to study i t s geographic v a r i a t i o n i n North America, to devise a useable c l a s s i f i c a t i o n from t h i s , to determine the relationship of the new world species to those of the old world, and to devise a plausible phylogeny f o r the whole genus, with emphasis here on the American  forms.  This study i s unusual i n that i t attempts to bridge two points of view concerning marten c l a s s i f i c a t i o n which necessitates the use of certain a r b i t r a r y standards of usage, with which the reader must be f a m i l i a r to understand what follows.  2  The older c l a s s i f i c a t i o n considers North American marten and f i s h e r to be made up of three species containing between them sixteen subspecies, whereas I attempt to advance the view that a truer conception of r e a l i t y i s to recognize only two and two  subspecies.  species,  These two ideas are not by any means s e l f -  exclusive, but merely a d i f f e r e n t i n t e r p r e t a t i o n of the same thing.  C l a s s i f i c a t i o n i s always a r b i t r a r y and since whatever  view accepted i s a matter of opinion, I am bound to present both views, the older scheme and my own. a means by which the reader may  In the following, as  recognize what c l a s s i f i c a t i o n i s  • referred to at any given place i n the text, the term species, subspecies  (or form or race) and the trinomial or i t s components  are always, when referred to i n the older sense, placed i n Inverted commas.  The use of inverted commas f o r the older  scheme i s not meant to imply derogatory scheme could have been reversed. ing the new  treatment of i t .  The  Since, however, I am champion-  plan, i t seems l o g i c a l to employ the above method.  To orientate the reader then, the old scheme of c l a s s i f i cation i s as follows: "Martes americana" "M.  a. americana"  "M.  a. atrata"  "M.  a. brumalis"  "M.  a. a b i e t i c o l a "  "M.  a. actuosa"  "M.  a. kenaiensis"  "M.  a. abietinoides"  "Martes caurina" "M. c. caurina" "M. c. orieenes" H  M. c. s i e r r a e "  "M. c. humboldtensis" "M. c. vancouverensis" "M. c. nesophila" "Martes pennanti" "M. p. pennanti" "M. p. columbiana" "M. p. p a c i f i c a " The new scheme I o f f e r , a s i m p l i f i c a t i o n of the above, i s as f o l l o w s : Martes americana M.  a. americana  M. a. caurina Martes pennanti This study i s based upon the examination of 3301 marten and f i s h e r s k u l l s of which measurements were taken on 1698. s p e c i f i c d i s t r i b u t i o n of these i s as f o l l o w s : Martes americana americana: 1803 examined, 1044 measured M. a. c a u r i n a:  936 examined,  390 measured  M. pennanti:  352 examined,  206 measured  M. f l a v i g u l a and g u a t k i n s i :  32 examined,  2 measured  M. f o i n a :  81 examined,  19 measured  M. martes ;  12 examined,  7 measured  M. z i b e l l i n a :  77 examined,  20 measured  M. melampus:  8 examined,  3 measured  The  4  Besides these a representative number of skins of each were examined.  The specimens examined are i n the collections of the  following:  University of B r i t i s h Columbia, B r i t i s h Columbia Prov-  i n c i a l Museum, National Museum of Canada, Royal Ontario Museum of Zoology, University of Michigan, Carnegie Museum, Museum of Vertebrate Zoology i n Berkeley, American Museum of Natural History, United States National Museum, United States B i o l o g i c a l Survey, U n i v e r s i t y of Utah, Museum of Comparative Zoology i n Cambridge, and Peabody Museum, Yale U n i v e r s i t y .  Besides these,  I have been allowed to examine specimens belonging to Messrs R. Y. Edwards, W. Cottle, R. Webb, J . Bryant, K. Racey and Doctors P. L. Wright and L. E. Yeager.  Mr. D. Flook and Mr. C. Lensinck  have donated specimens of t h e i r own to me, and Mr. E. McEwen provided me with measurements of specimens i n his possession. It i s pleasant to name the i n d i v i d u a l s who have provided assistance during the course of the study.  They include the  custodians of the above named c o l l e c t i o n s , e s p e c i a l l y Drs. D. Johnson, and H. Tate, and Miss V i o l a Schantz.  Dr. Johnson also  loaned specimens and transcribed portions of his rare copy of Kerr's Animal Kingdom f o r my use.  Dr. P. L. Wright, besides  allowing me to examine his specimens, sent me a manuscript  copy  of his paper on marten conspecifity a year before i t s p u b l i c a t i o n . Mr. K. Racey provided valuable advice on western martens.  Dr. L.  Yeager cleaned a large series of skulls so that I might examine them.  Mr. C a l v i n Lensinck and Mr. Donald Flook both made g i f t s  of marten skulls from Alaska and Mackenzie. provided material on marten d i s t r i b u t i o n .  Mr. Lensinck also Dr. G. G. Simpson  5  provided information on nomenclature. Dr. I. McT. Cowan i s d i r e c t l y responsible f o r the i n s t i g a t i o n of this study i n that he f i r s t suggested to me,  the problem  and has guided me throughout, though he must not be held  responsible f o r i t s e r r o r s .  Both he and Dr. W. Clemens have  been a source of ready information and encouragement;  I am  obliged to both too, for having assisted i n obtaining funds to assist me i n the work.  These funds were provided by the B r i t i s h  Columbia Sugar Refining Company and the B r i t i s h Columbia Research Council. Further assistance has been provided by the following: Drs. Peter Larkin, K. Graham, V. Krajina, J . Spencer, M.  Udvardi,  J . T. Gregory, N. Preble, G. Swanson, L. Butler, E. L. Coekrum, Messrs. R. Denney, J . Bryant, A. Cameron, R. W. Sutton, F. W.  Fay,  J . S. Tener, F. C. K l e i n s c h n i t z , B. S. Wright, D. Pimlott, and Miss Helenette S i l v e r .  Mrs. E. F. Bonoff helped i n the transla-  t i o n of Russian l i t e r a t u r e . The academic year 1953-54 was University.  spent i n study at Yale  For t h i s , o b l i g a t i o n i s owed to Drs. E. S. Deevey,  J r . , J . S. Nicholas and Mr. G. E.  Hutchinson.  I t i s a pleasure to acknowledge the help given by my wife, E l i z a b e t h Hagmeier, whose assistance during the preparation of this thesis has been i n c a l c u l a b l e .  6 CLASSIFICATION A.  THE GENTJS MARTES HISTORY OF CLASSIFICATION The post-Linnaean synonymy of the generic term Martes  as applied to martens and f i s h e r s , follows:  1758  Mustela Linnaeus, v o l . 1, p. 45.  1758  Z i b e l l i n e Buffon and D'Aubenton, v o l . 7, p. 309.  1765  Pekan Buffon and D'Aubenton, v o l . 1, p. 304 and p i . 42.  1765  Fouine Buffon and D'Aubenton, v o l . 13, p. l 6 l .  1771  Fisher.Pennant, p.  1774  Pekan Charlevoix, v o l . 3 , p. 134.  1775  Martes F r i s c h , p. 11.  1777  Mustella Scopoli, p. 491.  1792  Martes, P i n e l , v o l . 1, p. 55.  Type Martes  domestica  P i n e l = Martes f o i n a Erxleben.  1800  V i v e r r a Shaw, v o l . 1, p. 414.  1816  Tavra Oken , p. 1001.  1820  Martes, Nilsson, v o l . 1, p. 38.  1829  Z i b e l l i n a Kaup, v o l . 1, p. 31.  1843  G a l a d i c i t i s Smith, W. Jardines Nat. Library, v o l . 35> p t . l , p. 167.  1847  Putorius Boitard, v o l . 8, p. 12 (Diet. Univ. H i s t . Nat. 8:)  1865  Charronia Gray, p. 108.  1865  Foina Gray, p. 108.  1865  Pekania Gray, p. 107.  1865  Gulo "H. Smith", according to Gray 1865: 108.  1873  Aelurodon Cope, p. 1.  7 1904, Palaeogale Trouessart, p. 201. 1907, P u s t e l l a M i l l a r d , p. 1040 (Journ. Bombay Wat. H i s t . Soc. 1911,  170.  Martes. Thomas, p. 139.  1924, P l i o n i c t i s Matthew, v o l . 21, p.  135.  1928, Lamprogale Ognev, p. 26. 1937* Charionia Thorn, p. 317 (Journ. Bombay Nat. H i s t . Soc. Ray, i n 1693  39). was the f i r s t to use the name Martes i n a generic  sense, according to Trouessart (1897).  Linnaeus, however, used  the  term Mustela f o r the genus including with i t the weasels.  For  this reason i t took a long while f o r agreement to be reached  as to the proper designation f o r the genus of martens i n the s t r i c t sense.  During the 18th and 19th centuries, Mustela was  held to be the proper generic term, the weasels being named P u t o r i u s b u t since the f i r s t decade of the 20th century, Martes has been used, Mustela being reserved f o r the weasels, minks and ermines, and Putorius as genus or subgenus f o r the black-footed f e r r e t Mustela (Putorius) nigrepes. Alston, i n 1879> was the f i r s t to interest himself i n the  problem.  He wrote that most writers used the term Mustela  for the martens on the ground that Cuvier (1817) had done so. Alston concluded that the term was used not as Linnaean genus, but rather as "sous-genres" and hence was inapplicable.  The  f i r s t proper naming of martens was done by Nilsson i n 1820, used the term Martes.  who  Thus, concludes Alston, the proper term  8 should be Martes Nilsson 1820.  He adds that certain authors  have used the term Martes i n the past, not after Nilsson, but from Cuvier i n 1797*  This i s incorrect, he states, since the  word used was merely the p l u r a l form of the French Martre. Flower and Lydekker i n 1891 state that while Cuvier (1817) used the terra Mustela as a sub-genus f o r the martens, usual practice allows them consideration as a f u l l generic term.  They conclude  that martens should be named Mustela Cuvier 1817. Trouessart (1898-99) followed Alston i n refuting Mustela Linnaeus as the proper term, and retaining Martes Nilsson. A l l e n i n 1902 (a) gave consideration of Oken's (1816) use of Tavra f o r the genus name, and while considering him scarcely Linnaean, nowhere completely condemned him.  Sherborn i n 1902-  1933 gave authority f o r Martes to P i n e l (1792). In 1904, Palmer, i n his Index Generum, attributed the name Martes to F r i s c h (1775)*  Thomas and M i l l e r i n 1905 con-  demned Palmer's usage of Frisch's term.  They state that while  F r i s c h appears s u p e r f i c i a l l y binomal, he i s not, and often used phrases f o r h i s generic terms, and that h i s terminology was thus unacceptable. Thomas, i n 1911?  reviewing the c o n f l i c t i n g claims of  the terms Mustela and Martes as applying to the martens, concluded that only Martes was v a l i d .  He said that Linnaeus chose  many of h i s terms from Gesner (1606 and 1617-1621) and that by going to Gesner one could determine what "type" Linnaeus was thinking of i n his application of names. Mustela (Systema Natura 10, p. 45),  Concerning the genus  the type, he concluded, i s  9 by tautonymy Mustela erminea ("Mustela" Gesner, misquoted as Mustela v u l g a r i s ) .  In t h i s instance alone of the Gesner  quotations i n the Systema, he said, i s a second name attached to the primary one, f o r which tautonymy i s claimed.  This proved  to be a misquotation by Linnaeus, f o r while Linnaeus wrote "Mustela v u l g a r i s " Gesner, Gesner actually wrote only "Mustela" i n the German e d i t i o n quoted by Linnaeus, while i n the L a t i n edition he wrote "Mustela proprie s i c d i c t a " i n c o n t r a d i s t i n c t i o n to his "Mustelis d l v e r s i s " , which included the marten and the polecat} Thomas concluded that the Mustela referred to by both Gesner and Linnaeus was c l e a r l y the ermine, and that i t must be considered the type species.  Almost a l l writers since Thomas  have used Martes, under various a u t h o r i t i e s , f o r the martens, r e s t r i c t i n g Mustela to the weasels and ermines. Heider, Kuhlgatz, Hesse, Schulze and Kukenthal  (1926-  35) assert that a reference to Martes was made by Brisson i n 1762, but assert that neither he nor P l n e l can be considered legitimate authorities.  Wagler's (1830) usage of the term was  used f o r Herpestes (a v l v e r r i d ) and concluded that Nilsson (1820) must be considered the proper authority.  Ognev i n 1931 credited  the term to P i n e l , as did Neave (1930-1940). Brongersma, i n 1941, reviewed the problem of the use of the term Mustela and concluded that four d i f f e r e n t species were involved as types.  De B l a i n e v i l l e ( 1 8 4 1 ) , he asserts, employed  Mustela f o i n a as an anatomical type, even though i t was not mentioned by Linnaeus (see above).  Chenu and Desmarest (date  uncertain, but probably 1852 or 1853> P« 263) named M. martes as  1G type, although not i n a nomenclature^ sense. (p. 60) named Mustela martes.  Coues i n 1877  This type was also used by  Cuvier (1817, pp. 142-149), M i l l e r and Rehn (1901, p. 226), P a l mer (1904, p. 436) and E l l i o t  (1905b, p. 419).  E l l i o t i n 1901  (p. 33) mentioned Mustela l u t r a as the type, but t h i s apparently was a lapsus, since on page 352 he mentioned Mustela l u t r a as the type of the genus Lutra Erxleben.  Thomas i n 1911 > as pointed  out above, named Mustela erminea Linnaeus the type of the genus. Brongersma concluded that i f the International Regulations were to be c l o s e l y followed, Coues* designation (M. martes) would have to be maintained over that of Thomas (M. erminea). Since, however, Thomas' usage had been employed by most workers for t h i r t y years, much confusion would be created by a change and the author expressed the hope that the Commission would pass an opinion making Mustela erminea the type of the genus Mustela, so that the martens might r e t a i n the name Martes.  Brongersma be-  lieved the authority f o r the l a t t e r term should be attributed to Pinel. In 1945, Simpson i n h i s C l a s s i f i c a t i o n of Mammals reverted to Palmer's usage of F r i s c h as the authority f o r Martes. a l e t t e r dated 1951 he writes: Palmer....  In  "I took Martes F r i s c h 1775 from  Since F r i s c h i s doubtfully or not binomial, the  more usual and probable reference i s Martes P i n e l . . . .  There  i s , however, also some question as to Pinel's absolute p u r i t y ... so perhaps i t should be Martes Nilsson 1820.  I f Nilsson should  not pass, e i t h e r , then the name i s l o s t because the next reference, Wagler, 1830, uses Martes f o r a group of v i v e r r i d s . . .  11 Hershkovitz  (1948a) says that Palmer i n 1941  i n private  conversation denied the v a l i d i t y of F r i s c h ' s names, which Hershkovitz did also.  The same author (Hershkovitz 1948b) reasserted  his opinion, saying that F r i s c h was  non-Linnaean, and persisted  only because Palmer and A l l e n ( B u l l . Amer. Mus. 16: 13-22) had employed them.  In 1949  Nat. H i s t .  1902,  the same author denied  the v a l i d i t y of Oken's (1816) use of Tavra f o r the martens, saying the author was  non-Linnaean.  Ellerraan and Morrison-Scott  (1951)  suggest that Brisson's names could not be considered Linnaean and that Oken's names f a l l into much the same category. Frisch's terminology  they report that the International Commission  i n Paris i n July, 1948, Zool. Nomenclature 1950, this journal.  Concerning  declared his work to be unavailable ( B u l l . 4:549).  I have been unable to examine  To summarize the above, i t appears that by  strict  application of International Regulation, Mustela should be used as the generic term f o r the martens but that common usage and consideration of types gives the term Martes p r i o r i t y (Brongersma 1941,  Thomas 1911).  Although F r i s c h was  the f i r s t to use  the  term, his names have been declared I l l e g i t i m a t e (Ellerman and Morrison-Scott 195D  and the authority f o r the name must l i e with  any of Brisson (1?62), P i n e l (1792) or Nilsson (1820).  Brisson  has generally been considered unavailable, and so i t appears l i k e l y that P i n e l w i l l r e t a i n permanent authority f o r the term. For these reasons I accept Martes P i n e l as designating the genus. In 1918,  Pocock, on the basis of the structure of the  baculum of the Indian and Malaysian M. guatklnsi, separated  species M. f l a v i g u l a and  them as the d i s t i n c t genus Charronia.  12  Ogney i n 1928 showed that t h i s name was preoccupied by Charonia G i s t e l 1848,  p. 559, f o r a genus of Molluscs, and replaced i t  with the name Lamprogale.  Ognev i n 1931 and Pocock i n 1936(b)  accepted the change but Howell 1929, A l l e n 1938 and 1940, Simpson 1945 and Tate 1947 retained Charronla. A l l e n and Simpson i n s i s t ing  that the change was not merited.  Most recent authors,  however, refuse to accept Pocock s d i s t i n c t i o n as generic, and 1  Ognev ( 1 9 3 D , Carter H i l l and Tate (1944), Bobrinskoy e t . a l . (1944) and Ellerman and Morrison-Scott ( 1 9 5 D consider Charronia or Lamprogale merely a subgenus of the genus Martes, which i n t e r p r e t a t i o n i s accepted i n t h i s paper. TAXONOMIC POSITION The martens and f i s h e r s (Martes) have i n v a r i a b l y been placed w i t h i n the family Mustelidae, although how t h i s family l i e s r e l a t i v e to the Arctoids and Aeluroids has never been p r e c i s e l y determined.  For convenience  i t has generally been considered to  l i e nearer the former, however. Within the Mustelidae a great many subfamilies have been devised, e s p e c i a l l y by Pocock, although most authors e.g. Simpson and Ellerman, Morrison-Scott name only four or f i v e .  Martes i s  generally placed i n the subfamily Mustelinae, although some,,for example Pocock and A l l e n (1938 and 1940), name both the Martinae and Mustelinae, the f i r s t f o r the martens, the l a s t f o r the weasels  etc.  For greater d e t a i l see Simpson (1945). For ordinal characters of the family Mustelidae, and the  sub-family Mustelinae see M i l l e r 1912a, Pocock 1921a and b, A l l e n 1938 and 1940, S t i l e s and Baker 1935.  13  About a dozen recent genera of Mustelinae are considered to e x i s t today (Simpson 1945) but other authors have named more.  About another twenty to twenty-five genera have been  named as f o s s i l forms but again there i s great among authors.  disagreement  Keys to the recent genera are given by Gray  1865, M i l l e r 1912a, Pocock 1921b, A l l e n 1938-40, Dammerman 1940, Ognev 1931 and others.  Keys to the f o s s i l forms (now  outdated)  as w e l l as the recent ones are given by Matthew 1924 and Winge 1941.  The characters of the genus Martes have been given by  many authors, i n c l u d i n g the f o l l o w i n g , Flower and Lydekker M i l l e r 1912a, Pocock 1921a and 1921b, Ognev 1931»  1891,  S t i l e s and  Baker 1935> H a l l 1936, Dammerman 1940, and Winge 1941. There f o l l o w s a summary of the characters of the genus, compiled from the above:  are M u s t e l i d carnivores w i t h body l o n g ,  slender and f l e x i b l e , though l e s s so than i n Mustela; what t r i a n g u l a r , muzzle pointed;  head some-  limbs s h o r t , f e e t rounded,  toes s h o r t , f i v e toes on each f o o t ;  t a i l moderately l o n g , more  or l e s s bushy; s k u l l l o n g , narrow and ranging i n b a s i l a r l e n g t h from 60 to 115 mm.;  f a c i a l angle s l i g h t ;  tympanic b u l l a e moder-  a t e l y i n f l a t e d , w i t h t h i n w a l l s , not i n c l o s e contact w i t h parao c c i p i t a l processes;  p a l a t e extending behind l a s t upper molars;  the b r a i n although l a r g e , i s not e x c e s s i v e l y so;  the a n t e r i o r  edge of the o r b i t , although s l i g h t l y pushed forward, i s not g r e a t l y d i s p l a c e d ; the j u g u l a r process i s d i s t i n c t ; formula ^ i l t ] ^  inner moiety of M  1  dental  l a r g e , at l e a s t as l a r g e as  outer w i t h two i n t e r n a l cusps (protocone and an accessory cusp) and a cingulum;  P " w i t h a s i n g l e deuterocone, t h i s forming a 4  14 d i s t i n c t i n t e r n a l lobe; width;  lower  length of whole tooth greater than i t s  with evident though small metaconid and entoconid  (thus with a t o t a l of 5 cusps);  t r i g o n i d always longer than  talonid, the l a t t e r semibasened. B.  THE SUBGENERA OF MARTES HISTORY OF CLASSIFICATION Although Cuvier (1817) and certain other authors notably  Lydekker (1885) used subgenera to separate the martens from the weasels, interest here must be r e s t r i c t e d to subgeneric d i v i s i o n within the martens themselves. The f i r s t  to interest himself i n the problem was Gray,  who i n 1865 and I869 named four subgenera, which with a modern interpretation of the species constituting  them, follows:  Genus Martes: Subgenus Martes: Martes martes, M. melampus, M.' z i b e l l i n a and M. americana. Subgenus Pekania: Subgenus Foina:  Martes pennanti.  Martes foina.  Subgenus Charronia:  Martes f l a v i g u l a and M. guatkinsi.  Pocock i n 1918, as has been described e a r l i e r , reasserted the v a l i d i t y of Gray's subgenera Martes and Charronia, naming Charronia a f u l l genus and basing d i s t i n c t i o n on baculum structure. Pekania, as a subgenus, was not considered but i n North America at least has continued to persist as a taxonomic u n i t . was not upheld by Pocock, nor has i t persisted.  Foina  Ognev, i n 1928  (as discussed e a r l i e r ) changed Charronia to Lamprogale.  Certain  15 authors, namely Ognev (1928), Ognev (193D  and Pocock 1936b  accept Lamprogale, but most (Howell 1929 > A l l e n 1938 Simpson 1945» Tate 1947 Charronia.  and  1940,  and EllermariiMorrison-Scott 195D  retain  Howell, Simpson and Tate give Charronia f u l l generic  status, while Carter H i l l and Tate (1944) and Ellerman, MorrisonOgnev 1931» A l l e n 1938-40 and Bob-  Scott consider i t a subgenus. rinskoy 1944  give Lamprogale subgeneric status only.  In 1931 H a l l named Martes gazini from the t e r t i a r y of North America and erected the new subgenus Tomictis f o r i t s use. S i m i l a r l y i n 1946 Shikama named the subgenus Ten f o r Martes ten from the Pleistocene of Japan. We f i n d then, that the following are the subgenera that have been named to constitute the genus Martes. Subgenus Martes P i n e l 1792,  vol.  1, p. 55 f i d e Gray  1865 and 1869. Subgenus Pekahia Subgenus Charronia p. 307,  Gray 1865,  p. 107.  Gray 1865,  p. 108,  Pocock  1918,  synonymous with Lamprogale Ognev 1928, p. 26  and 30, considered by some a genus. Subgenus Foina  Gray 1865> p. 108.  Subgenus Tomictis  H a l l 1931 > P« 156.  Subgenus Ten Shikama 1946, p. Only three of these subgenera have been held v a l i d by students of recent mammals (e.g. Ognev 1931» Bobrinskoy 1944, M i l l e r 1924, Anderson 1946, Ellerman Morrison-Scott 195D•  These  with t h e i r constituent species are as follows: Subgenus Martes: Martes martes, M. foina, M. z i b e l l i n a , M. melampus and M. americana. Subgenus Charronia = Lamprogale: Martes f l a v i g u l a  16  and M. g u a t k i n s i . Subgenus Pekania; Martes pennanti.  Because t h i s c l a s s i f i c a t i o n i s c u r r e n t l y accepted i t i s the one used i n the present paper. Although only two subgenera of f o s s i l martens have been named, i t appears l i k e l y that a c r i t i c a l examination of f o s s i l members of the genus would r e s u l t i n the naming of s e v e r a l t o many more.  Nor i s i t known w i t h a few exceptions what f o s s i l  species are represented i n the subgenera of recent martens. TAXONOMIC CHARACTERS I t i s g e n e r a l l y stated that nine species of marten e x i s t i n the world today, these being "Martes americana", "caurina", martes, f o i n a , z i b e l l i n a , k i n s i , and melampus.  pennanti, f l a v i g u l a , guat-  F o r reasons given elsewhere, however,  t h i s paper assumes the number to be only seven, by considering "M. americana" and "c a u r i n a" a s i n g l e species (M. americana), and M. f l a v i g u l a and M. g u a t k i n s i another.  As stated e a r l i e r ,  these are grouped i n t o the f o l l o w i n g subgenera: Subgenera Martes: zibellina,  Martes americana, martes, f o i n a ,  and melampus  Subgenus: Pekania: Martes pennanti Subgenus C h a r r o n i a ( F l a v i g u l a ) : Martes f l a v i g u l a The c h a r a c t e r i s t i c s of the subgenera Martes and Charr o n i a have been c a r e f u l l y considered by Pocock (1918), Ognev (19330, Pocock (1936b) and Bobrinskoy (1944).  The d i s t i n c t i v e  nature o f the subgenus Pekania has never been considered seriousl y and i n f a c t i s not so c l e a r cut as i s that between the f i r s t  17  two named groups.  A summary of the c h a r a c t e r i s t i c s of the  subgenera of martens follows. 1.  Subgenus Charronia = Lamprogale.  yellow above; groove;  Colour varied black and  upper l i p not divided by a d i s t i n c t v e r t i c a l  baculum d i s t i n c t i v e , the d i s t a l end curved somewhat  abruptly upwards and backwards i n the form of a hook, the t i p s l i g h t l y expanded to produce four processes, each with a rounded condyle-like head. Distribution:  through the mountainous regions of northern India,  northwards through China to Amur, and southward through Malaysia to Sumatra, Java and Borneo, also i n the h i l l s of southern India. 2.  Subgenus Martes:  Colouration generally monotone, brownish,  but often with a l i g h t e r or yellowish spot on breast; divided by a d i s t i n c t v e r t i c a l groove;  upper l i p  baculum not as above,  gradually i n c l i n e d upwards and forwards at i t s d i s t a l end, where i t divides into two processes which remain separate, or which may unite. Distribution: 3.  a l l of h o l a r c t i c Europe, A s i a and North America.  Subgenus Pekania:  Colouration usually monotone, brownish,  with chest spot absent or reduced, usually whitish; divided by a d i s t i n c t v e r t i c a l groove; genus Martes,.  upper l i p  baculum as i n the sub-  I t i s so l i k e the preceding subgenus that i t i s  best distinguished by i t s r e l a t i v e l y larger s i z e , the length of i t s head and body being more than 570 mm.  instead of l e s s and  i t s condylpbasal length being more than 95 nun* instead of l e s s . Distribution:  h o l a r c t i c North America.  18  C.  THE SPECIES OF MARTES HISTORY OF CLASSIFICATION In Eurasia Adequate h i s t o r i e s of the development of the present  concepts of the c l a s s i f i c a t i o n of the genus Martes are not a v a i l able, although attempts of varying degree have been made by de B l a i n v i l l e (1841), von Martens (1870) as summarized by Coues (1877) and Alston (1879)•  Much of what follows i s collated  from these. It appears that the ancient Greeks were the f i r s t to recognize and name what we now place i n the genera Martes, Mustela and Putorius.  Three names were used f o r these, i n  t h e i r Anglicized form, I c t i s , Gale and Mustela, but these terms were used rather indiscriminately.  A r i s t o t l e compared Gale to  I c t i s , saying that the l a t t e r had a thicker pelage and a paler belly.  Elsewhere he said that Gale of the forest waged war  against r a t s .  De B l a i n v i l l e believed that by Gale A r i s t o t l e  meant weasel and by I c t i s marten, and that the d i s t i n c t i o n of the two had been made by then.  Gesner and l a t e r authors r e f e r  to the marten even more d i s t i n c t l y .  I t i s believed, however,  that these writers s t i l l confused the polecat (Putorius) with the weasel (Mustela) because Aristophanes used the word Gale to refer to an unclean woman, and Herodotus referred to a Gale of A f r i c a , which was l i k e l y a V i v e r r i d . The Roman writers did not use the older Greek terminology, developing rather t h e i r own, e.g. Martes and Mustela, i n t h e i r present form.  Thus the o r i g i n of these words i s nearly  unknown, or at least obscure.  In. spite of t h i s these are the  19 terms used by modern zoologists.  In Leviticus 11, 29 Moses  l i s t e d a number of animals that were considered impure.  Among  these was the Choleb which many translators have interpreted as Gale or Mustela.  De B l a i n v i l l e believes t h i s was the stone  marten (Martes foina) of today. It i s uncertain who f i r s t made the d i s t i n c t i o n between the two species of European marten (Martes f o i n a and the pine marten Martes martes).  De B l a i n v i l l e attributes the f i r s t d i s -  t i n c t i o n to P l i n y , who while transcribing A r i s t o t l e ' s statements referred to above, substituted the word V i v e r r a f o r Gale and Mustela f o r I c t i s .  Ever since, he said, Mustela has appertained  primarily to the stone marten.  Viverra, i t i s assumed, was  meant by P l i n y to be the pine marten.  M a r t i a l replaced the  term V i v e r r a with Martes, though providing no description to either.  Most l a t e r authors, e s p e c i a l l y Agricola and Gesner,  maintained the d i s t i n c t i o n , most applying Martes to the pine marten, Mustela to the stone marten, and Putorius and I c t i s to the weasels and polecats. Von Martens and Alston (1879) however, considered neither A r i s t o t l e nor P l i n y to have recognized the d i s t i n c t i o n seen by de B l a i n v i l l e .  I t may have f i r s t been done by Albertus  Magnus (who mentioned both Martarus or Martes fagorum and Martes abietum).  Just as l i k e l y though, they believed, the d i s t i n c -  t i o n was not r e a l l y made u n t i l as l a t e as the time of Buffon and Linnaeus.  The c r a n i a l and dental characters by which the pine  and stone martens could be distinguished were f i r s t recognized and pointed out by Hensel i n 1853 and further elaborated by  Blasius i n 1857 1865,  (Saugethier  Coues 1877,  Ognev 1931,  Deutschlands, pp. 211-214), Gray  Alston 1879,  M i l l e r 1912a, Reynolds  1912,  Bobrinskoy et al.1944, Rode and Didier 1944,  among  others. The o r i g i n of the name Mustela i s believed by von Martens to originate i n the word Mus.  Sundevall believed further that i t  came from the Greek word f o r hunt, since Palladlus referred to the mouse hunting a b i l i t y of the animal and since i t i s s t i l l called Moustelle i n Nice and Lorraine.  Aubert and Wimmer, said  von Martens, concluded, however, that a l l the older  references  are r e a l l y to the stone marten i n the s t r i c t sense, since i t occurs throughout Greece (see also Rolleston 1868) called I c t i s there.  and i s s t i l l  The o r i g i n of the word Martes i s s t i l l  obscure. Kerr (1792) considered,the Satherius of A r i s t o t l e to have been the f i r s t reference to the sable, and Shitkov agrees.  (1940)  Pallas (1811) and Heptner (1934) attributed the name  to Albertus Magnus, however, and de B l a i n v i l l e (1841) to A g r i c o l a . The f i r s t statement of the d i s t i n c t i v e marks of the species made by Aldrovandi  was  and Ambrosini (according to de B l a i n v i l l e ) and  further elaborated by Brandt (1855), Gray (1865  and 1869)  and  others named above. The  s p e c i f i c names of the three species of martens d i s -  cussed previously were considered by von Martens as follows: Z i b e l l i n a has been known since the l a t t e r half of the middle ages under many variations such as sabelus, z i b e l l i n a , zebel, s e b e l l , sable, sobol, soboli and samur. zebel i s as old as the 9th  Graff says the Germanic term  century.  21 Martes now occurs i n nearly a l l Germanic and Romantic languages as marta, martre, mart, martora, martin, marten, marter, mard, maar, meardtr and martarus.  The word was  first  used by M a r t i a l as Martes, probably being taken from some nonRomantic language. Foina i s present i n nearly a l l Romantic languages. Forms include fuina, f u l n t r a , foulne, f a i n a , faguino, fahino, f ayna, fagina, farveina, f i e r n a , feb, feh-wamme and fagorum. The C e l t i c , Slavonic, F i n n i s h , Russian, and Magyarish words f o r the  animal appear to be of t o t a l l y d i f f e r e n t o r i g i n than the  L a t i n term f o i n a . Martes martes has at times undergone various s p e c i f i c designations, these including s y l v e s t r i s , s v l v a t i e a , v u l g a r i s and ableturn. fagorum.  M. f o i n a has been named v a r i o u s l y domestica and  I t was not recognized by Linnaeus, who did not see  i t near his native Upsalla. Ray introduced the concept of the genus and i t was he who grouped the several species of marten then known into a single genus.  He gave to t h i s the generic name Martes.  Later  species of A s i a t i c marten were described by Boddaert i n 1735 (Martes f l a v i g u l a , the yellow b e l l i e d marten) and by Wagner i n 1841 (M. melampus, the Japanese marten). Buffon l a t e r described many discoveries from America "Pekan","Martre", "Vison", etc., the f i r s t two of which are d i s cussed subsequently.  22 In North America Marten: The c l a s s i f i c a t i o n of species of European marten was f a i r l y well understood by the time Buffon and d'Aubenton (1758 and 1765) described the New World marten and i t was with these that they were f i r s t compared. Buffon employed French names rather than L a t i n binomials and hence h i s terminology has never been preserved i n use. Pennant i n 1771 and again i n 1781 f i r s t used  Linnaean  terminology to describe the New World forms, saying that two species of marten occurred there; Mustela z i b e l l i n a .  these being Mustela martes and  Pennant said he had been informed by Dr.  Pallas that no sables (M. z i b e l l i n a ) occur northeast of the "Anadyr" River i n the country of the "Ichutki" Indians. In 1772 Forster named the American marten Mustela martes after that name of Linnaeus. Mustela z i b e l l i n a .  P a l l a s (1776-1780) named i t  In 1792 Kerr renamed i t Mustela z i b e l l i n a  americana being the f i r s t to apply the epithet americana to the New World form.  He also named two other forms, Mustela martes  and Mustela z i b e l l i n a alba as being present.  Thomas i n 1879 and  A l l e n i n 1895 both believe Kerr should be given authority f o r the present usage of the current name Martes americana, but most subsequent authorities have attributed i t to Turton.  A l l e n says  that Turton copied most of his names from Kerr and that the l a t t e r ' s work i s "mere trash" compared to that of Kerr.  For  purposes of the present paper, I consider Kerr rather than Turton the authority f o r the epithet americana as applied to martens.  23  Turton i n 1802 named the marten as Mustela  americana.  Coues i n 1877 considered Turton's 1806 e d i t i o n the authority of the s p e c i f i c name americana and although Rhoads (1902) pointed out that Turton f i r s t used the combination i n h i s 1802 edition and although Sherborn does the same, most authorities refer the authorship to the 1806 volume.  I have examined both  editions of Turton's work, arid find Rhoad's designation and not Coues  1  to be correct.  Nevertheless f o r reason given previously,  authority i s given to Kerr f o r the name and not to Turton i n either of h i s volumes.  Gray 1865 and 1869 and Trouessart  erroneously give authority f o r the name to a non-existent 1803 edition of Turton. Ord i n 1815 named three American species, namely Mustela martes, M. z i b e l l i n a and M. americanus.  Rafinesque  (1819a and b) described a new species, Mustela vulpina from "the upper Missouri" which Coues (1877) concluded i s r e a l l y a white-tailed s e m i - a l b i n i s t i c i n d i v i d u a l of the t y p i c a l form. Kuhl i n 1820 named a form as Mustela leucopus which Coues (1877) considered a white-footed a l b i n l s t i c form.  In 1823 Cuvier  named a form Mustela huro, which Coues again explains as merely a l i g h t coloured Individual of the t y p i c a l eastern marten. Harlan (1825) considered two forms, Mustela z i b e l l i n a and Mustela martes, to occur and confused these martens with the mink, Mustela vlson, which he considered i d e n t i c a l .  Godman (1828)  named two forms, Mustela martes and Mustela z i b e l l i n a .  American  pine marten, he states, d i f f e r from t h e i r European equivalent i n possessing a yellow rather than white breast.  The sable i s  24 much paler than most marten, he says, a specimen collected by Lewis and Clark being "bleached". G r i f f i t h Smith and Pidgeon (1827) apply the name Mustela l e u c o t i s , this l i k e l y being a corruption of Kuhl's term leucopus.  Waterhouse (1$28) named only one form, Mustela z i b -  b e l l i n a , though ten years l a t e r Waterhouse (1838) considered z i b e l l i n a to be absent from the continent and to be replaced by Mustela martes and M. foina.  Richardson i n 1829 believed only  Mustela martes to be present, as did Gapper i n 1830.  The l a t t e r ,  however, pointed out that d i s t i n c t i o n s existed between the Old and New World forms.  Godman (1831) named Mustela z i b e l l i n a as  the only form present, while Emmons i n 1840, De Kay i n 1842 and Linsley i n the same year named Mustela martes.  Gray (1843)  named the marten Martes leucopus, using Frisch's generic name f o r the f i r s t time.  Smith i n the same year described three  forms, Martes z i b e l l i n a , Martes huro, and Martes l e u c o t i s , and Schinz (1845) named Mustela vulpine. Mustela leucopus and Mustela huro.  Gesner i n 1847 grouped a l l as Mustela martis [ s i c ] .  Audubon and Bachman (1845-48 and 1851-54) concluded that a l l martens they had examined were referrable to Mustela martes and denied the presence of the sable on the continent. Brandt i n 1855> however, compared skins of A s i a t i c and American marten and concluded that the two were almost i d e n t i c a l . The American form he named Mustela z i b e l l i n a var. americana. B i l l i n g s (1857) said that three species of martens occurred i n the New World but did not name them.  25  Baird (1857 and 1859)» after considering Brandt's views, expressed disagreement with him, concluding that two forms existed i n America, one with a whitish head, l y i n g near to the sable, and one with a darkish head near to the pine marten but both probably s p e c i f i c a l l y d i s t i n c t from the corresponding Old World forms.  The name he used for,both was Mustela americana.  Ross i n 1861 (a and b) concluded that two species occurred, these being Mustela americanus and M. z e b i l l i n a [ s i c ] .  He believed  the Mackenzie River form was only a colour v a r i e t y of the A s i a t i c sable. A l l workers up to t h i s period worked primarily on colour and structure of pelage i n t h e i r attempts to determine the species of marten present i n the New World.  For t h i s reason i t i s  not surprising that they referred American forms to e i t h e r the pine marten (Martes martes) , to the sable (M. z i b e l l i n a ) or named a l b i n i s t i c variants as d i s t i n c t species (M. huro, M. leucopus). Before a better understanding of the status of the New World forms could be determined, i t was necessary to examine c r a n i a l and dental c h a r a c t e r i s t i c s .  One of the f i r s t to do t h i s was Gray,  who i n 1865 and again i n 1869 concluded that the l a s t upper molar i n New World forms was small and quadrate, whereas i n Old World forms i t was large, massive and nearly twice as long on the inner as on the outer side.  So long as one remembers that Gray had  no specimens from the C o r d i l l e r a and west coast of North America and that to these h i s d i s t i n c t i o n does not hold, Gray's separation is valid.  He further divided "Martes americana" into three  races, which as subspecies have been held as v a l i d to the present.  26 These are: v a r i e t y huro from Fort F r a n k l i n (the equivalent of "Martes americana actuosa");  v a r i e t y abietinoldes (now "Martes  americana abietinoides"); and v a r i e t y leucopus (now "M. americana americana")*  Gray's work may be considered to mark the  end of the exploratory period and the beginning of the s c i e n t i f i c one, insofar as American marten are concerned. A l l e n , i n 1869, examined many Old and New World skins and skulls and concluded that a l l belonged to a single circumpolar species, with possibly one or two well marked continental races.  Thus he named a l l marten, including the American mar-  tens, Mustela martes.  Ames i n 1874-, however, named two  species  as being present, Mustela americana and Mustela martinus.  He  gave no description of e i t h e r . In I876 (a) A l l e n reviewed h i s older paper and retracted the views i t contained.  He agrees to the d i s t i n c t i o n s made by  Gray between the Old and New World forms and adds that while the character was inadequate f o r the separation of M. f o i n a and M. americana the two could be distinguished by differences i n the second lower molar which i n a l l the Old World forms possesses an inner cusp lacking i n the American form.  Rhoads (1902) points  out, however, that this d i s t i n c t i o n i s i n v a l i d , or at least not constant, and my own studies have l e d me to the same conclusions. A l l e n also pointed out, quite c o r r e c t l y , that American marten increase i n size from south to north i n the fashion stipulated by Bergmann.  A l l these he named Mustela americana.  Coues i n 1877 monographed the Mustelid genera including the genus of martens.  He agreed with Gray's separation of Old  27  and New World forms and through detailed comparison named other distinguishing c h a r a c t e r i s t i c s .  He concluded that M.  americana l a y nearest to the sable, of a l l Old World forms, a relationship not much doubted since.  He disagreed with Gray's  d i v i s i o n of the species into three v a r i e t i e s , saying that they represented i n d i v i d u a l v a r i a t i o n only. to be one species Mustela americana.  A l l forms he considered In his 1884 catalogue,  True l i s t e d Coues one species, M. americana, without comment, as did T y r r e l l i n 1889. With Merriam, i n I890, began the t h i r d phase i n the history of marten c l a s s i f i c a t i o n .  His impress upon mammalian  taxonomy i n North America was extreme and the publication of his description of the species Mustela caurina resulted i n what may be termed the phase of taxonomic s p l i t t i n g .  From 1890  u n t i l recently the martens have been divided into t h i r t e e n or fourteen forms, a l l but one s t i l l considered v a l i d .  Much tax-  onomic s p l i t t i n g of the period i s to be deplored, but i t i s a fact that Merriam's description of caurina from the C o r d i l l e r a of North America marks the most basic d i v i s i o n of marten that can be made.  Merriam believed M. americana to l i e nearest to  M. z i b e l l i n a of a l l Eurasian forms. Flower and Lydekker (189D regress to the conclusion that Mustela americana l a y very near to M. martes and M. z i b e l l i n a , and was d i f f i c u l t to distinguish from them.  Herrlck  (1892) likewise emphasizes t h i s lack of d i s t i n c t i v e n e s s , as does Lydekker I896.  28 Poland (1892) erroneously names the marten Mustela martes, saying i t i s indistinguishable from the Old World form and very nearly inseparable from M. f o i n a .  In spite of t h i s ,  he provides a correct view of geographic v a r i a t i o n i n marten, showing extreme insight into the problem. said, are large and pale; uniformly coloured; furred;  Alaskan martens, he  C a l i f o r n i a n animals are small and  the Labrador marten i s large and r i c h l y  those from southern Canada and the northeastern United  States are small and more l i g h t l y furred.  These are apt des-  criptions of the current forms "actuosa", " s i e r r a e " , "brumalls", and "americana".  He concluded that "each d i s t r i c t of North  America has i t s own peculiar type". In 1897 Bangs described as a d i s t i n c t species Mustela atrata, the Newfoundland marten.  The following year, the same  author described another from Labrador, Mustela brumalls. 1900  In  Osgood named Mustela americana actuosa from Alaska, Yukon  and Mackenzie.  In 1901  he described Mustela nesophila from  the Queen Charlotte Islands. Elliot  (1901a) named f i v e forms, Mustela americana,  Mustela caurina, M. c. atrata, M. c. brumalis and M. c. actuosa. M i l l e r and Rehn (1901) l i s t them as M. americana, M. a. actuosa, M.  atrata, M. brumalis and M. caurina.  In 1902  Preble described  Mustela americana a b l e t i c o l a from Manitoba, and Rhoads i n  1902  Mustela caurina origenes from Colorado.  revis-  Rhoads' work was  ionary i n nature and he named the following forms:  M.  americana  (with the subspecies abietinoides, actuosa and brumalis), atrata and M. caurina with i t s new  subspecies origenes.  M. Elliot  29  i n 1903(a) described Mustela americana kenaiensis from the Kenai, Alaska, and i n 1905(a) M. b o r i a from Mackenzie. l i s t e d s i x species of martens:  Elliot  (1905b)  Mustela americana (with subspec-  ies a b l e t i c o l a , abietinoides, actuosa and brumalis), M. kenaiensis, M. atrata, M. b o r i a and M. nesoohila. synonymized M. b o r i a with M. a. actuosa.  In 1908 Preble  G r i n n e l l and Storer,  i n 1916, described Martes caurina sierrae from the S i e r r a Nevadas and G r i n n e l l and Dixon i n 1926, Martes caurina humboldtensis from the Coast Mountains of C a l i f o r n i a and M. c. vancouverensis Vancouver Island.  from  These l a t t e r mark the l a s t forms of recent  North American marten to be described. The systematic catalogues of the period 1912 to the present show a progressive attempt on the part of systematists to group the known forms of marten into r e a l i s t i c s p e c i f i c groups. M i l l e r i n 1912(b) l i s t e d s i x species:  Martes americana (with  subspecies americana, kenaiensis, a b i e t i c o l a , abietinoides and actuosa), M. atrata, M. b o r i a , M. brumalis, M. caurina (with subspecies caurina and origenes) and M. nesophila. 1917  Anthony i n  named four species, using M i l l e r ' s c l a s s i f i c a t i o n but omit-  ting M. b o r i a , origenes and nesophila. M i l l e r , i n his Catalogue of Mammals, 1924, again l i s t e d s i x species, they being now Martes americana (with subspecies americana, kenaiensis, a b i e t i c o l a , abietinoides and actuosa), M. atrata, M. b o r i a , M. brumalis, M. caurina (with subspecies caurina, sierrae and origenes) and Martes nesophila.  Anthony,  i n h i s Field-book (1928) used M i l l e r ' s c l a s s i f i c a t i o n , with the exception of M. b o r i a which he omitted.  I t would appear that  30  G r i n n e l l and Dixon (1926) were the f i r s t to suspect that only two major types (species) of martens occurred i n North America. They wrote that "the material as yet available f o r study  indi-  cates the existence of two d i s t i n c t species of marten i n northwestern America —  Mustela caurina, with short bullae and  other  minor characters, on the P a c i f i c coast from Admiralty Island south to east-central C a l i f o r n i a ;  and Mustela americana, with  long bullae, etc., i n the i n t e r i o r of the continent and west at the north clear to the Alaska Peninsula.  Each of these  contains several well marked geographic races".  A l l e n (194-2)  employed the most modern c l a s s i f i c a t i o n of a l l , the one which most systematists now  concur.  species  with  He also named only  two  species, Martes americana and Martes caurina, the f i r s t with the subspecies,americana, kenalensis, a b i e t i c o l a , abietlnoides, actuosa, atrata, b o r i a and brumalis;  the second with the sub-  species caurina, s i e r r a e , origenes, nesophila, vancouverensis and humboldtensis.  Anderson, i n his Catalogue of Canadian  forms (194-6) followed A l l e n except that he omitted M. a. b o r i a and the American forms M. c. s i e r r a e , origenes and humboldtensis and raised M. atrata to s p e c i f i c status.  Miller  Kellogg (1955) l i s t a l l of Allen's subspecies and name three species, "M.  americana", "M.  and  except " b o r i a " .  caurina", and"M.  atrata". The r e a l i z a t i o n that only two major forms of marten exist on the continent was  l i k e l y recognized by some systematists  long before A l l e n put the idea into p r i n t .  Just how  these  two  forms (M. caurina and M. americana) were related to each other  31  (i.e.  whether d i s t i n c t species or subspecies) has only recently  been made clear. In 1936  H a l l wrote that " c r i t i c a l study of the described  species and subspecies of true martens i n North America (subgenus Martes) almost c e r t a i n l y would show that the named forms belong to no more than two, In 1942  and possibly only to one, f u l l species".  A l l e n published h i s heretofor discussed volume i n which  he grouped a l l as subspecies of the two species M. americana and M. caurina.  In 1946  H a l l again wrote "I do not know whether  caurina i s a species d i s t i n c t from americana and the species name should be regarded as tentative pending a r e v i s i o n of the genus".  Rand i n 1948(b) l i s t e d these as the only two  of marten occurring i n Canada.  Dalquest,  species  the same year, studied  the problem and concluded that "east of the Rocky Mountains the ranges of the two species approach closely but each retains d i s t i n c t i v e characters....  Gn the basis of the evidence at  hand, the two should be considered f u l l species u n t i l p o s i t i v e proof of intergradation i s established." that "my  Durrant i n 1952  said  studies of martens cause me to doubt that M. caurina  i s d i s t i n c t from Martes americana", but added no d e t a i l . It remained f o r Wright (1950 and 1953) proof of c o n s p e c i f i c i t y demanded by Dalquest.  to provide  the  He showed that  the two underwent complete intergradation i n western Montana and central Idaho.  On the basis of t h i s , Wright l i s t e d a l l t h i r t e e n  forms of marten as subspecies of the single species Martes americana.  In view of the v i t a l nature of Wright's work, i t i s  reviewed i n greater d e t a i l elsewhere i n t h i s paper.  32  Fisher: Because f i s h e r occur only i n the New World and were discovered after the European forms were r e l a t i v e l y well known, the problems of t h e i r nomenclature  and c l a s s i f i c a t i o n are not  the same as were those of New World martens. Buffon, i n 1765> published the f i r s t d e s c r i p t i o n of the Fisher and l a t e r the name was used by Brisson.  In 1771  Pennant named and described not only Buffon's Fisher but another form he named Pekan.  Concerning the f i r s t of these, he expressed  b e l i e f that i t might be i d e n t i c a l to the sable of Asia, although i n 1781 he retracted t h i s view.  Concerning the second, he said  that he could not separate i t from the Vison of Buffon. Commenting on t h i s s i t u a t i o n Coues (1877) f i d e Baird (1857 and 1859) writes that Buffon, Brisson and Pennant a l l "described the animal from the same specimen - one i n the cabinet of M. Aubry of P a r i s . . . .  Pennant's account of his F i s h e r i s  unmistakeable, but he describes i n addition, the Pekan of Buffon, not recognizing i n i t the same species.  These two accounts  furnished f o r many years the bases of a l l the s c i e n t i f i c binomial names imposed by various authors." Erxleben, i n 1777 gave f o r the f i r s t time a Linnaean binomial to the f i s h e r , naming i t Mustela pennantli, although very shortly l a t e r Schreber (1778) renamed i t M. canadensis. Thus not only were two animals named from the same specimen, but two names were applied at nearly the same time to the same species. This s i t u a t i o n provided much of the nomenclatural problem associated with the h i s t o r y of f i s h e r taxonomy.  I n 1785, Boddaert  33  used the name M. melanorhyncha f o r the f i s h e r and Kerr i n 1792 named two forms, M. canadensis and M. z i b e l l i n a nigra (both f o r the f i s h e r ) .  Shaw (1800) changed the generic name to Viverra,  naming two forms, V. piscator and V. canadensis.  Turton (1802)  named Mustela nigra and M. canadensis and used the same nomenclature again i n 1806.  In 1815 Ord used the names M. nigra  and M. hudsonius and used the term M. canadensis f o r the otter (Lutra canadensis) (Rhoads 1894-). duced the name M. m f a .  Desmarest (1820-1822) i n t r o -  In 1825 Harlan named only one form,  c a l l i n g t h i s M. canadensis, while the following year Godman (1828) named a single form M. pennanti [ s i c ] .  Lesson ( 1 8 2 ) 7  named three species, a l l referable to the f i s h e r . M. pennanti, M. canadensis, and M. p i s c a t o r i a . Smith and Pidgeon, i n 1827, followed Godman s 1  These being Griffith  nomenclature.  Fischer (1829) used the term M. godmani (corrupted by Gray 1865 to M. goodmanii).  Richardson (1829) named two forms, these  being M. canadensis and a pale variant M. canadensis var. alba. Swainson 1835, f o r the f i r s t time employed the generic name Martes f o r the f i s h e r , naming i t Martes canadensis.  Emmons  In 1840 remarked that e a r l i e r d i v i s i o n of the f i s h e r into two species was due to colour v a r i a t i o n and grouped them together as one form, Mustela canadensis.  Smith i n 1843 named three  species, Martes canadensis, M. nigra and M. pennantii.  Gray  i n 1843 reduced a l l to the single species Martes canadensis. Schinz 1845 and Audubon and Bachman (1845-1848 and 1851-1854) * named one form Mustela canadensis. Baird i n 1857 and 1859 was the f i r s t to attempt to systematize the chaotic nomenclatural state of the f i s h e r .  He  34 makes c l e a r t h a t , as pointed out p r e v i o u s l y , Buffon, B r i s s o n and Pennant a l l named t h e i r f i s h e r s from the same specimen. Buffon named h i s Pekan, as d i d B r i s s o n , but Pennant named both Pekan and F i s h e r , not knowing that both were the r e s u l t of exami n a t i o n of the same specimen. B a i r d then went on to the problem of p r i o r i t y of date between the two names, canadensis and pennant! applied to the species of Buffon and Pennant. 1777  "While Erxleben's work i s dated  and volume 3 of Schreber's work i s dated 1778,  Baird f e l t  i t p o s s i b l e that p a r t s of Schreber's work may have been issued p r i o r to 1777  and states as evidence that Erxleben quotes the  p l a t e s of Schreber i n v a r i o u s p l a c e s .  Nonetheless, i n l i e u of  b e t t e r evidence, B a i r d accepted Erxleben as having p r i o r c l a i m . On page 455 of E r x l e b e n , i s described a Mustela canadensis, the synonomy of which includes both the Vison and Pekan of Buffon, but which i n a c t u a l f a c t describes only the mink (Mustela v i s o n ) .  On page 470, however, Erxleben described  under M. pennant11 the F i s h e r of Pennant and concluded that the proper name f o r the species i s Mustela p e n n a n t i i E r x l e b e n .  He  suggests elsewhere that two species, the f i s h e r and the b l a c k c a t , might e x i s t , however. I n i860 Godman described both Mustela p e n n a n t i i and M. canadensis but expressed b e l i e f i n Pennant's i d e a that the l a t t e r was named f o r a mink s k i n .  Gray i n 1865 used the binom-  i a l Martes p e n n a n t i i , employing modern terminology f o r the f i r s t time. Coues (1877) continued the c l a r i f i c a t i o n of nomenclature  35 begun by Baird.  He concluded that Erxleben 1777 (Mustela  pennanti), Shaw 1800 (Viverra p i s c a t o r) , Turton 1802 and 1806 (Mustela nigra), Lesson 1 8 2 7 (Mustela piscatoria) and Boddaert 1785 (Mustela melanorhyncha) a l l named t h e i r forms after the Fisher of Pennant.  Schreber 1778 (Mustela canadensis) named  his after the Pekan of Buffon. but applied i t to Mustela vison.  Erxleben named a M. canadensis Shaw named a V i v e r r a piscator  whose o r i g i n Coues could not determine.  Turton named a Mustela  nigra f o r both Buffon's Pekan and the otter (Lutra) which he considered i d e n t i c a l .  Like Baird, he concluded that the f i s h -  er's name should r i g h t l y be Mustela pennantii Erxleben.  Else-  where, however, Coues drops the double i ending of pennantii, reducing i t to pennanti.  The current use of the shortened  epithet may be said to date from Coues. True i n 1884, used Coues* term, M. pennanti, as did T y r r e l l 1889. Rhoads (1894) considered the nomenclature! problem and concluded as did Baird and Coues, that the proper name was Mustela pennantii.  In I 8 9 8 , however, i n his r e v i s i o n of the  species, he employed the name M. canadensis Schreber.  Contrary  to e a r l i e r opinions he concluded that since Erxleben made reference to Schreber's plates, and also on Sherborn's opinion, that Schreber was published (or the I l l u s t r a t i o n s at least) prior to the issuance of Erxleben's work i n 1777 and that he thus gained p r i o r i t y .  Rhoads, i n this paper, described a new  'subspecies from the P a c i f i c coast, M. c. p a c i f i c a .  This has  been the f i r s t named subspecies to persist i n the l i t e r a t u r e . A l l e n , i n I898, replied to Rhoads decision, saying that since  36 Schreber's plate name M. canadensis i s preceded by the plate name M. l u t r a canadensis, i t preoccupies the epithet canadensis as applied to the f i s h e r .  Hence A l l e n concluded, the proper  name f o r f i s h e r was Erxleben's M. pennanti. In 1899 E l l i o t used the contracted epithet penanti, t h i s probably being a lapsus, M i l l e r 1900, M i l l e r and Rehn 1901 and E l l i o t 1901,  returning to the orthodox pennanti.  1910 used the term pennantii.  Bangs i n  In 1912(b) M i l l e r using Thomas'  (1911) precedence, made i t Martes pennanti, which form i t has held to the present, with few exceptions (see Jordan 1929, Warren 1942  f o r exceptions).  In 1935 Goldman named the t h i r d and l a s t subspecies to be accepted by modern workers.  This was M. p. columbiana  from  the Rocky Mountains, which intergraded i n the P r a i r i e Provinces, he believed with t y p i c a l M. pennanti.  A l l three ^subspecies"  are l i s t e d i n M i l l e r and Remington's (1955) catalogue. Some c r i t i c i s m has been directed towards the d i v i s i o n of f i s h e r into subspecies.  Grinnell i n 1933  and i n 1937  (Grin-  n e l l , Dixon and Linsdale 1937) wrote that after c r i t i c a l examination of the specimens i n the United States National Museum, he was unable to separate M. pennanti p a c i f i c a from M. p. pennanti and concluded they were a l l to be named simply M. pennanti. H a l l , i n 1936, working with fewer specimens, came to the same conclusion.  None of them considered M. p. columbiana,  of  course, which was not described u n t i l 1935 (Grinnell's l a t e r manuscripts being completed  before this date).  In 1950 (Hemming 1950) redrafted A r t i c l e 14 of the  37  I n t e r n a t i o n a l Rules such that t r i v i a l names based on a male person's surname should " c o n s i s t o f the exact surname ... t o which should be added ... the t e r m i n a t i o n i n the g e n i t i v e case ... - i " .  Thus the e p i t h e t pennanti i s given v a l i d i t y  over the o l d e r form p e n n a n t i i by the Regulations. The c u r r e n t l y accepted c l a s s i f i c a t i o n of the f i s h e r s i s as f o l l o w s : "Martes pennanti pennanti" "M. p. p a c i f i c a " "M. p. columbiana"  Erxleben 1777.  Rhoads 1898. Goldman 1935.  SYSOPSIS OF THE SPECIES AND SUBSPECIES OF MARTEN Comparisons of the d i s t i n c t i v e q u a l i t i e s of the various species of marten have been made i n the past by the f o l l o w i n g authors:  A l l e n 1876(A), A l l e n 1938-40, A l s t o n 1879, B l a n f o r d  1888-91, Bobrinskoy et a l . 1944, Bonhote 1901, Brongersma 194-1, Coues 1877, de B l a i n v i l l e 1841, de Vos 1952, D i d i e r 1947, Ellerman, Morrison-Scott 1951, Flower 1885, Gerrard 1862, 1865 and 1869, H a l l 1926, Kneeland 1859, Merriam 1890,  Gray  Miller  1912(a), Ognev 1925, Ognev 1931, Owen,1853, Pocock 1918, Pocock 1921(a and b ) , Pocock 1936(b), Reynolds 1912, Rode and D i d i e r 1944, Schmidt 1943, Tate 1947, and Bannikov 1953.  A t no time,  however, have more than three o r four species been compared simultaneously.  The synopsis provided below, prepared  through  reference t o the above named authors and by examination of s p e c i mens, attempts to help remedy t h i s l a c k . For those i n t e r e s t e d i n the general anatomy of the martens the reader i s r e f e r r e d t o the w r i t i n g s of de B l a i n v i l l e  38 (1841), Reynolds (1912), H a l l (1926 a and b), and Cooper  (1954).  Dental terminology i s given i n an i l l u s t r a t i o n by Gregory ( 1 9 5 D  f i g . 20.08. I have no way of knowing how c a r e f u l l y the s p e c i f i c d i s t i n c t i o n of species of marten i n a b i o l o g i c a l sense has been determined.  It i s l i k e l y that the species of the  Pekania and Charronia are t r u l y species.  subgenera  Within the remaining  subgenus Martes, the s i t u a t i o n i s less l i k e l y .  The sympatricity  of the ranges of M. martes and M. f o i n a suggests t h e i r d i s t i n c t specificity.  S t r e u l i (1932) says that not a single hybrid or  intermediate was found among several hundred specimens of the two forms examined although Schmidt (1943) says Severtzov reported hybrids i n the Tienschan Mountains.  Rode and Didier (1944)  report that 42 skulls of the two species were d i s t i n c t i v e i n every case.  Severtzov (I876) considered M. intermedia i n t e r -  mediate between M. f o i n a and M. martes but he apparently worked only with skins.  Ellerman, Morrison-Scott (195D  conclude,  however, that the two species are not always e a s i l y d i s t i n g u i s h able, which suggests the p o s s i b i l i t y that Intermediates may sometimes occur. Martes z i b e l l i n a has a range that barely overlaps that of M. martes;  i t i s uncertain that i t i s i n contact with that  of M. f o i n a .  although Schmidt (1943) c i t e s Severtzov as saying  that i n the Tienschan Mountains M. foina has been crossed with the sable.  The c r a n i a l c h a r a c t e r i s t i c s of a l l three are marked  i n the specimens I have examined, but Ponomarev i n 1946, recorded that he obtained hybrids from M. z i b e l l i n a and M. martes crosses.  39 Whether t h i s situation occurs under natural conditions I do not know. Martes melampus has always been considered a d i s t i n c t species.  Ellerman, Morrison-Scott (1951) say that while i t s  coloration i s near M. z i b e l l i n a the s k u l l i s nearer M. f o i n a and M. martes and conclude that i t i s an " i s o l a t e d and v a l i d species". Its true status i s unknown to me, since older students have never considered i t s s p e c i f i c i t y i n a c r i t i c a l  fashion.  It has never been demonstrated that North American martens are s p e c i f i c a l l y d i s t i n c t from Old World forms.  M. amer-  icana americana i s morphologically d i s t i n c t from M. z i b e l l i n a , i t s nearest Old World neighbour, but t h i s cannot be said to be marked i n the relationship of M. americana caurina to M» z i b e l lina.  These two can almost always be separated, but t h e i r d i s -  tinctions are small, f a r smaller i n fact than exist between the two true subspecies M. americana americana and M. a. caurina. It i s quite possible that the A s i a t i c and American forms w i l l eventually be shown to be p o t e n t i a l l y capable of interbreeding, as Rausch (1953) and Bobrinskoy (1944) believe to be the case i n many Siberian-Alaskan forms.  I f this i s found to be so, M.  am-  ericana w i l l be reduced nomenclaturally to a subspecies of M. z i b e l l i n a or of M. martes.  The relationship between A s i a t i c  and American forms i s considered elsewhere i n greater d e t a i l . In order to f a c i l i t a t e the i d e n t i f i c a t i o n of unknown s k u l l s , a key i s provided below, which i s based almost completely on dental c h a r a c t e r i s t i c s .  I t makes no attempt to denote r e l a -  tionship and i s completely a r t i f i c i a l . a series of indices devised by Brongersma  I t i s based l a r g e l y upon (1941) which were  40 enlarged upon during the course of the present study. indices are given i n d e t a i l i n Appendix A.  These  The key considers  M. f l a v i g u l a and M. guatkinsi to be conspecific, and further, takes M. americana to subspecies. 1.  Width of inner lobe of P P  4  4  x 100/width of trenchant part of  more than 65  2  l b . Less than 65  7  2a. Greatest width of canines at base x 100/bulla length more than 100.5  3  2b. Less than 100.5  5  3a. Mesial length of M  x 100/breadth of M  1  1  more than  55  4  3b. Less than 55  M. f l a v i g u l a (including M. guatkinsi)  4a. Width of inner lobe of P  x 100/length of P  more than  28  M. melampus  4b. Less than 28  M. americana caurina  5a. L a t e r a l length of M than 79  1  x 100/mesial length of M  1  M. americana  more americana  5b. Less than 79  6  6a. Greatest width of canines at base x 100/bulla length more than 95.5 6b. Less than 95.5 7a. Breadth of M  1  M. martes M. z i b e l l i n a  x 100/length pf P4 more than 8 9 . . M. f o i n a  7b. Less than 89  M. pennanti  Where the key proves inadequate f o r the i d e n t i f i c a t i o n of specimens, I suggest reference be made to the s t a t i s t i c s on which i t i s based, given i n Appendix A;  even when identification  41 appears f a i r l y certain, I would suggest that the reader make comparison to these.  Other keys to a r e s t r i c t e d number of  species are given by M i l l e r 1912(a), Ognev 1931» and Bobrinskoy et a l . , 1944.  Diagrams of the skulls of the various species of  marten are given on Figures 1 and 2. Maps showing the d i s t r i b u t i o n of Eurasian forms are given on Figures 3, 4, 5 and 6}  these have been prepared from  many sources, but c h i e f l y from the maps of Kozhantschikov (1930), Ognev (1931), S t i r t o n (1939), Schmidt (1943) and Bobrinskoy et a l . (1944) and from the accounts of Sowerby (1922, 1930), A l l e n (19381940), Chasen (1940) Brongersma (1941), Harper (1945), and E l l e r man, Morrison-Scott (1951). Synonymies f o r the species and subspecies considered'in this paper are, because of t h e i r lengthy nature, summarized alphabetically  i n Appendix B.  The measurements of condylobasal  length provided are meant to give some idea of geographic v a r i a tion.  They have been compiled from the specimens examined and  the l i t e r a t u r e .  Because the samples are small, only the means,  observed extremes, and sample size are given, measurements i n centimeters.  being  The c l a s s i f i c a t i o n that follows i s that of  Ellerman, Morrison-Scott (1951 and 1953) and Chasen (1940). Other recent c l a s s i f i c a t i o n s of importance are given by M i l l e r 1912(a), Ognev 1925, Ognev 1931, Pocock 1936(b), Schmidt 1943, Bobrinskoy et al.., 1944, and Harper 1945. Martes martes (Linnaeus .1758) = M. s y l v e s t r i s , vulgaris  and ableturn of authors (see Appendix B f o r d e t a i l ) .  Type l o c a l i t y : Range:  svlvatica,  Upsala, Sweden.  B r i t i s h Islands including  Ireland, Norway, Sweden,  Figures 1 and  2.  (To face page 42)  Figures 1 and  2.  The skulls of the species of the world's marten, i n ventral aspect. From sources referred to i n the text, and from o r i g i n a l material.  C A U RINA  AMERICANA  MELAMPUS  PENNANTI  FOINA  FLAVIGULA  MARTES  ZIBELLINA  Figures 3?  4, 5 and  (To face page 42)  6.  Figures 3? 4 , 5 and 6.  The d i s t r i b u t i o n of Eurasian marten. Collated from Kozhantschikov (1930), Ognev ( 1 9 3 D , Schmidt (1943), Bobrinskoy et a l (1944) and many other sources. Type l o c a l i t i e s are given f o r the c l a s s i f i c a t i o n s of Ellerman, Morrison-Scott (1951 and 1953) and Chasen (1940).  42 Denmark, France, Belgium, Holland, Switzerland, I t a l y , northern Spain, B a l e a r i c Islands, Sardinia, Bohemia, Poland, to Russia and the very southwestern parts of S i b e r i a , to the Obe and Eptischa Rivers, the White Sea and the Caucasus, also A s i a Minor and P e r s i a (Bobrinskoy 1944 and Ellerman, Morrison-Scott 1951). Sowerby (1922-30) l i s t s them from Heilungkiang and K i r i n i n Manchuria, and Sowerby (1916) from southwest Kansu and northwest Brongersma: (1941) records a specimen very near to  Ssuchuan.  M. martes from Nepal, possibly another from Tibet. Coues 1877, pl» 3 ;  S k u l l diagrams and photographs: Alston 1879, f i g . 2}  Reynolds 1912, p i . 2;  f i g s . 76, 77, 78, 79 and 80; 601  and 602;  Ognev 1931, v o l . 2, pp. 593, 600,  Schmidt 1943, f i g s . 8 and 9}  f i g s . 357, 358, 359, and 360.  M i l l e r 1912(a)  Rode and Didier 1944,  Other sources are given by Ognev  1931, v o l . 2 .  Specimens examined:  twelve from Europe  This forest dweller i s a t y p i c a l occupant of the o l d hardwood or coniferous second growth f o r e s t s . of up to 2000 meters i n the Caucasus. and b e r r i e s . i n trees.  I t l i v e s i n woods  I t feeds on nuts, insects  The den i s b u i l t i n a cavity 5 to 30 meters high Infrequently i t uses the dens of wild pigs.  The  period of heat begins i n February and reaches i t s peak i n July. Gestation l a s t s f o r eight months (270 to 285 days).  The l i t t e r  of two to s i x , i s born i n A p r i l or May (Bobrinskoy 1944, Schmidt 1943).  Geographic v a r i a t i o n , according to Bobrinskoy (1944)  c h i e f l y expressed i n size and colour.  Jurgenson (1951) says  much geographic v a r i a t i o n i n s k u l l and teeth i s associated with  4-3 the type of food  eaten.  Characteristics:  More active than M. f o i n a ;  (males weighing 1200-1650  grams, females 800-1350 grams);  long and slim with short limbs; than 57 cm.; slatey; tips;  smaller and l i g h t e r body  length of head and body l e s s  f u r f i n e , s o f t , brown below and above, r a r e l y  outer fur r i c h brown, under fur red grey with red yellow top of head same colour as spine;  often with a l i g h t  coloured throat patch of yellow, which extends behind rear of forelegs;  upper l i p with d i s t i n c t v e r t i c a l groove;  t a i l bushy,  rather short, being considerably longer (without t i p hairs) than one half the length of head and trunk (Ellerman, Morrison-Scott 1951 say only 49$ length of head and trunk);  t a i l vertebrae  said to be v a r i o u s l y 20-22, 17, 18-19, 18, 10+, 18-33, 17-19, 18-19 (Bobrinskoy et a l . 1944-, Ognev 1931, Gray 1869, de B l a i n v i l l e 1841, Flower 1885, Reynolds 1912, Schmidt 1943, Gerrard 1862);  h a i r endings of t a i l long, being more than quarter length  of t a i l ; hair;  t i p of t a i l rounded;  soles of feet covered with t h i c k  s k u l l massive, z i b e l l i n a shaped, i . e. long, narrow, and  rather produced, t h i n and deep when seen from behind;  s k u l l and  braincase less massive, e s p e c i a l l y i n the posterior part; lobasal length 70 to 90 mm.;  condy-  zygomatic width about 48 mm.,  the  zygomatic width being rather more than half condylobasal length; zygomatic arches highest at rear, whence they slope suddenly down and forward, being longer and narrower; only r a r e l y completely narrow and elongate;  absent;  s a g i t t a l crest heavy*  rostrum with p a r a l l e l sides,  anterior nares oval i n shape;  with moderate central c o n s t r i c t i o n ;  nasal bones  f r o n t a l p r o f i l e only s l i g h t l y  sloping, concavity of p r o f i l e not very pronounced;  supraorbital  44 c o n s t r i c t i o n moderate, pointed, rather deep and always present; post o r b i t a l process nearly midway between point of greatest c o n s t r i c t i o n of cranium and the anterior root of zygoma, usually weak, occasionally absent;  postorbltal  region ( i n front of  point of greatest constriction) short and convex; t i v e l y narrow and long (about 4.3  cm. i n length), often with a  distince azygos notch on rear surface; r e l a t i v e l y large;  palate r e l a -  anteorbital foramen  bullae moderate i n length and rather widely  spaced, so that twice the distance between carotid canals i s greater than the greatest length of the bullae;  the distance  between the outer edges of the jugular foramina i s shorter than the greatest length of the bullae;  distance between the carotid  canals equals about half the distance between the foramen lacerum posteriore and the eustachian foramen;  b u l l a containing three  or more chambers, with p a r t i t i o n s strongly developed;  mastoid  process extends somewhat beyond the edge of the meatus;  general  d e n t i t i o n rather strong; upper premolars not crowded and s t r a i g h t ; 3 PM strongly convex on inner surface, strongly concave on outer;  4  r e l a t i v e l y large PM  with thick inner cusp, i t s width being  to i t s length and nearly equal to width of trenchant trenchant portion as long as M  1  i s wide;  equal  part,  M" i s large, massive, 3  inner moiety large - twice as large as outer moiety and one longer, waist separating the two strongly constricted;  M^  third with  a s l i g h t l y developed inner tubercle at base of main cusp ( i . e . hypoconid);  M2 with inner cusp only s l i g h t l y developed;  bacu-  lum of the Martes type gradually bending upward throughout i t s d i s t a l t h i r d , 35 to 45 mm.  long ending either In two  vertical  45 prongs, or these being closed to form a transverse perforation (according to Pocock 1918 always forked, which conclusion I find to be i n c o r r e c t ) . Martes martes martes (Linnaeus 1758) Type l o c a l i t y : Range:  Upsala, Sweden.  Europe north of the Mediterranean east as f a r as the  White Sea, Kiev and Vetebsk i n A s i a .  Large and dark i n colour  (Bobrinskoy et a l . 1944). Condylobasal length:  males 8.66 (8.41-9.00) n = 6;  females 7.90  (7.78 - 8.16) n = 3 . Martes martes borealis Kutznetzov 1941 (according to Bobrinskoy et a l . 1944). Type l o c a l i t y : not known. Range:  northern parts of European Russia, excluding the Kola  Peninsula. Very l i k e M. m. ruthena, but with winter pelage l i g h t e r (Bobrinskoy et a l . 1944). Martes martes latinorum (Barrett-Hamilton 1904) Type l o c a l i t y : Range:  Nurri Mountains, Sardinia.  I t a l y , Sardinia, Balearic Islands (Ellerman, Morrison-  Scott 1951)•  As i n M. m. martes but with pelage l i g h t e r i n  colour, throat patch yellower ( M i l l e r 1912a). Condylobasal length:  males 8.66 (8.50 - 8.80) n  s  3;  7.89 (7.70 - 8.30) n = 6. Martes martes lorenzl Type l o c a l i t y :  Ognev 1926.  Storojevala, Kuban D i s t r i c t , Caucasia.  females  46 Range:  the forested area of the whole Caucasus.  Large, as i n  M. m. martes. but with winter f u r coarse and thick, dark brown with a c h a r a c t e r i s t i c red-olive cast (Ognev 1926, Bobrinskoy et a l . 1944).  Condylobasal length: 7.89  (7.70 - 8.30)  males 8.53 (8.41 - 8.66) n = 4;  females  n = 6.  Martes martes n o t i a l i s (Cavazza 1912) Type l o c a l i t y :  South of Abruzzi, southern I t a l y .  Range:  Smaller, paler than M. m. martes. with l e s s  Italy.  dense pelage (Cavazza 1912). Martes martes ruthena Type l o c a l i t y : Range:  Ognev 1926.  Dmitrovsk S u b d i s t r i c t , Moscow Government, Russia.  central part of European Russia.  Smaller than M.  m.  martes and l i g h t e r i n colour with a reddish sandy cast (Ognev 1926, Bobrinskoy et a l . 1944). Condylobasal length: 7.57  (7.11 - 8.01)  males 8.20 (7.89 - 8.66) n = 22;  females  n = 11.  Martes martes uralensis Kutznetzov 1944 (according to Bobrinskoy et a l . 1944).  Type l o c a l i t y : Range:  unknown.  the whole of the Ural Range.  Colour very much as i n  M. m. ruthena and borealis but larger i n size than e i t h e r (Bobrinskoy et a l . 1944). Martes f o i n a (Erxleben 1777) Type l o c a l i t y :  =  domestica, alba, fagorum.  Germany.  Range: Europe, except southern Spain and I t a l y and the B r i t i s h  47 Isles;  present i n northern Spain and I t a l y , Bosnia, France,  Belgium, Holland, Germany, Denmark, Switzerland, Crete, Poland, Finland, Ukraine, Crimea, Caucasus, Transcaucasus, Russian and Eastern Turkestan, northwards to the A l t a i , A s i a Minor, Persia, Afghanistan, Syria, Palestine, Baluchistan, Kashmir, Himalayas and the Tibetan Plateau, Punjab, Chinese Turkestan, Mongolia, Manchuria,  and probably parts of northern China, northern C h i l i ,  Shansi and into West China, probably not much further south than Shansi and Szechwan.  (Sowerby 1922-30, Sowerby 1916, Ognev  1931, A l l e n 1938-40, Brongersma 1941, Bobrinskoy et a l . 1944, Tate 1947, Ellerman, Morrison-Scott 1951, Specimens examined:  81 from Europe and A s i a .  S k u l l diagrams and photographs: p i . G4 and G13; pt. 5, P i . 15;  Bronn 1874-90;  Festa 1914, p i . 1;  625, 626, 627, and 628; f i g s . 8 and 9;  A l s t o n 1879, f i g . l j  Miller  Ognev 1931, v o l . 2, pp. 623,  Migulin 1938, p. 167;  Bobrinskoy et a l . 1944, f i g . 77;  Didier 1944, pp. 127 and 128; 367 and 368;  de B l a i n v i l l e 1841,  Klassen und Ordnungen v o l . 1,  Coues 1877, p i . 4;  1912a, f i g . 80;  and Bannikov 1953).  Schmidt 1943, Rode and  Baumann 1949, f i g s . 365, 366,  Bannikov 1953, P« 40.  Other sources are named  by Ognev 1931, v o l . 2. This marten i s unlike M. martes i n that while i t i s found i n woods, i t also occurs i n barren rock s l i d e s .  I t s den  i s made i n crevices i n the rock and unlike M. martes i t does not avoid human habitation, often l i v i n g i n abandoned dwellings. It feeds on rodents, b i r d s , birds' eggs, frogs, various insects,  48 berries, and grapes. 3000 meters.  It l i v e s i n the h i l l s to an a l t i t u d e of  Time and length of the period of reproduction i s  as In H. martes (Bobrinskoy et a l . 1944).  Asdell (1946) says  that i n England i t mates i n July and August and that the gestat i o n period i s 8.5 to 9 . 6 months. ford (1888  - 189D  nine weeks.  In India, according to Blan-  mating occurs i n February and gestation l a s t s  This i n t e r p r e t a t i o n may be a result of lack of  knowledge concerning delay i n implantation of the blastocyst, but A s d e l l suggests that i f i t i s correct that delayed implantat i o n occurs only i n the northern part of the animal's range and not i n the southern, i t may be a function of seasonal daylength. According to Bobrinskoy i n d i v i d u a l v a r i a t i o n i n M. f o i n a i s very great, while geographic v a r i a t i o n i s only s l i g h t . Characteristics:  A sluggish animal, being r e l a t i v e l y large and  heavy (male weighing 1700 - 2100 gms.,  females 1100 - 1500 gms.);  body broad and compact with long limbs; less than 57 cm.; often s l a t y ;  length of head and body  fur coarse, less s o f t , brown above and below,  outer fur d u l l grey brown, under fur greyish white;  top of the head same colour as spine;  usually with a l i g h t e r  coloured throat patch, usually white or creamy, not extending between legs but rather forking, each branch running down the front legs;  upper l i p with d i s t i n c t v e r t i c a l groove;  t a i l bushy,  rather short, though somewhat longer than i n M. martes, without hair t i p s , considerably longer than one half head and body length (Ellerman, Morrison-Scott say actually 49$ length);  of head and trunk  t a i l vertebrae various-stated to be 17 - 19, 18 and  20 - 22 (Schmidt 1943,  de B l a i n v i l l e 1841,  Bobrinskoy 1944  and  49 Ognev 1 9 3 D ;  h a i r endings of t a i l long, being more than half  length of t a i l ; hair;  t i p of t a i l pointed;  s k u l l l i g h t , f l a v i g u l a shaped ( i . e . swollen, f l a t t e n e d ,  with short nose); posterior part;  s k u l l broad, more massive, e s p e c i a l l y i n condylobasal  length 70 - 80 mm. usually l e s s  than 83 mm.,  zygomatic width 48 mm.,  condylobasal  length;  est  soles of feet lacking  being much more than h a l f  zygomatic arches regularly curved, broad-  and highest near t h e i r middle, broader and shorter;  crest absent or very weak; verging sides;  sagittal  rostrum wide and short, with con-  anterior nares heart shaped;  nasal bones  strongly constricted;  f r o n t a l p r o f i l e more sloping, concavity  much more pronounced;  supraorbital c o n s t r i c t i o n s l i g h t , f l a t ,  blunt and often absent;  p o s t o r b i t a l process much nearer point  of greatest c o n s t r i c t i o n than anterior root of zygoma; present, strong, coming to a d i s t i n c t point;  always  p o s t o r b i t a l region  (in front of point of greatest constriction) long and s t r a i g h t ; palate r e l a t i v e l y broad and short ( 35 nmu long), truncated at posterior margin;  a n t o r b i t a l foramen r e l a t i v e l y small;  case broader, less extended;  brain-  bullae short and widely spaced so  that the distance between carotid canals i s greater than the greatest length of the bullae;  the distance between the outer  edges of the jugular foramina i s the same or more than the greatest  length of b u l l a ;  distance between carotid canals equals two-  thirds the distance between foramen lacerum posterior and eustachian foramen;  b u l l a with 3 or more chambers, the p a r t i -  tions l e s s well developed than i n M. martes; extends somewhat beyond edge of meatus;  mastoid  process  general d e n t i t i o n  50  weaker, upper premolars crowded, often placed diagonally;  PM  2  4 evenly biconvex, somewhat small;  PM  with inner cusp small  ( t o t a l width l e s s than length, and barely h a l f width of trenchant p a r t ) ;  inner cusp usually placed diagonally to axis of  trenchant part;  length of PM  4  1 1  greater than width of M ; M  small, inner lobe small, hardly l a r g e r than outer lobe; s t r i c t i o n of waist weak;  con-  M^ with a well developed hypoconid;  M£ likewise with an inner cusp present and well developed;  bacu-  lum of the Martes type, bending gradually upward throughout i t s d i s t a l t h i r d , 56 to 59 nun. long, occasionally with two  vertical  prongs at t i p , these most often united to produce a transverse perforation. Martes f o i n a f o i n a (Erxleben). Range:  Type l o c a l i t y :  German;  Europe, except southern Spain, probably eastwards into  Russia (Ellerman, Morrison-Scott  1951)•  The type form, with  longer hairs of back tipped with sepia, the general hue of upper parts drab ( M i l l e r 1912a). Condylobasal  males 8.19  length:  7.77 (7.40 - 8.00)  - 8.46)  n = 5;  females  n = 5.  Martes f o i n a bosniaca Type l o c a l i t y :  (8.90  Bosnia,  Brass 1911  ( i n Aus der Reich der Pelze)  Yugoslavia.  Range and c h a r a c t e r i s t i c s uncertain. Martes f o i n a bunites (Bate 1906). Type l o c a l i t y :  Kontopalo, Kania, Crete.  Range:  Near to M. f . intermedia but with shorter f u r  Crete.  51  of  a d u l l e r , more uniform colour, without any gloss, t a i l less  bushy and shorter (Bate 1906). Condylobasal length: 7.00  males 7.72  (7.64  - 7.84)  n = 2;  females  1.  n =  Martes f o i n a intermedia Type l o c a l i t y :  (Severtzov  1873)  Basin of the Chu<, - T a l l a s and Naryn, eastern  Turkestan. Range:  Russian and Chinese Turkestan, Tianshan, Afghanistan,  Baluchistan, western P e r s i a and Kashmir (Ellerman, Morrison-Scott 1951).  Bannikov (1953) says i t occurs i n Mongolia.  d i s t i n c t i v e of a l l subspecies of M. foina;  The most  compared to M. f .  foina are small i n s i z e , l i g h t i n colour and with comparatively l i t t l e f u r (Bobrinskoy et al.. 1944). males 9.04  Condylobasal length: females 7.48  (7.37  - 7.80)  Martes f o i n a kozlovi Type l o c a l i t y : Range:  Kam  Ognev  (7.60 - 8.46)  n =  15}  n = 9. 1931  ( v a l l e y of River Mekong, eastern T i b e t .  the Tibetan Plateau and Himalayas.  Characteristics un-  certain. Condylobasal length:  males 8.10  Martes f o i n a mediterranea Type l o c a l i t y :  n = l'j  females 7»52  n =  1.  (Barrett-Hamilton I898)  S i e r r a de Jerez, Cadiz, Spain.  Range: southern Spain.  Compared to M. f . f o i n a with the longer  hairs of back tipped with l i g h t yellowish brown, the general hue of the upper parts l i g h t e r , yellower, and l e s s drab ( M i l l e r 1912a).  52  Martes f o i n a m l l l e r i  F e s t a 1914.  Type l o c a l i t y : A g h i o s I s i d i r o s , I s l a n d o f Rhodes, e a s t e r n Mediterranean. Range:  I s l a n d o f Rhodes.  Martes f o i n a n e h r i n g i  Characteristics uncertain.  ( S a t u n i n 1906), i n M i t t . Kauk. Mus. T i f l i s  2 : 120,292. Type l o c a l i t y :  T I f l i s , Transcaucasia.  Range and c h a r a c t e r i s t i c s u n c e r t a i n . Condylobasal length:  males 8.30 (8.10 - 8.51)  n = 4;  females  7.58 (7.21 - 7.90) n = 4. Martes f o i n a r o s a n o w i M a r t i n o 1917 • Type l o c a l i t y :  north-western slope o f the Chatyrdag Mountains,  Crimea. Range:  Crimean P e n i n s u l a .  Condylobasal length:  Characteristics uncertain.  male 7.87 n = 1;  f e m a l e s 7.57 (7.42 - 7.68)  n * 3. Martes f o i n a s y r i a c a Type l o c a l i t y :  ( N e h r i n g 1902)  Wadi S y r ( w h i c h runs i n t o Wadi K e f r e n , a t r i b u t a r y  of t h e l o w e r J o r d a n ) , S y r i a . Range and c h a r a c t e r i s t i c s u n c e r t a i n . Martes f o i n a t o u f o e u s (Hodgson 1842). I n t h e J o u r n a l o f t h e A s i a t i c S o c i e t y o f B e n g a l , I I , p. 2 8 1 ) . Type l o c a l i t y : Range:  Llasa, Tibet.  T i b e t , Himalayas and A f g h a n i s t a n ( J e r d o n 1 8 7 4 ) . C o l o u r  of head and body above i s l i g h t y e l l o w i s h g r a y ; are  the extremities  b l a c k i s h , t h e c h i n , t h r o a t and b r e a s t pure w h i t e ( H o r s f i e l d  53  1851). Kozlovi.  Possibly quite similar to M. f . intermedia and M. f . Ellerman, Morrison-Scott remark:  "Despite Pocock's  contention that t h i s i s a l l i e d to M. melampus, i t looks much more l i k e M. foina.  I t s range i s adjacent to that of foina,  very f a r from melampus.  From notes l e f t by him, Chaworth-  Musters evidently intended to treat i t as f o i n a . Pocock (1941, p. 322 footnote).  See also  We cannot trace that the form  'kansuensis' noted by him on t h i s page was ever described." Martes z i b e l l i n a (Linnaeus 1758) Type l o c a l i t y :  "Asia s e p t e n t r i o n a l i " ;  according to Ognev (1925)  t h i s must be redesignated, after Gmelin, as "Surroundings of Tobolsk, Tomsk Government, S i b e r i a . " Range:  S i b e r i a , from the middle of Pechora to the lower Leni,  and from the A l t a i to the Ussuri region, also Kamchatka, Sakhaline, northern Japan (Hokkaido and the K u r i l e s ) , Korea, Manchuria, northern Mongolia and Tannu-tuba.  Now highly f r a g -  mented (Bannikov 1953, Bobrinskoy et a l . 1944) and with range much reduced (Harper 1945).  According to Jerdon ( 1 8 7 4 ) , Blyth  took a specimen i n Tibet he was i n c l i n e d to i d e n t i f y as M. z i b e l l i n a (Bobrinskoy et. a l . 1944, Sowerby 1922-30, Ellerman, Morrison-Scott 1953, Kozhantschikov 1930, Ognev 1931).  Zhitkov  (1937) t i t l e s a paper "On the former d i s t r i b u t i o n of the sable i n Europe"; whether i t occured here formerly I do not know, not having examined the paper.  Harper (1945) says that i n the past  i t has extended as f a r west as the Kola Peninsula or even Lapland. Specimens examined:  77 from A s i a .  54  Skull diagrams and photographs: G13;  Ognev 1925, p i . 26;  593 and 594; Bobrinskoy p. 40.  de B l a i n v i l l e 1841, p i . G7 and  Ognev 1931, v o l . 2, pp. 563, 587, 591,  et a l . 1944, f i g . 77;  Bannikov 1953?  Other sources are named by Ognev 1931, v o l . 2. This animal i s a forest dweller, usually l i v i n g i n  h i l l y country interspersed with moss covered rock slopes.  All  of i t s range i s characterized by the presence of spruce and pine (i.e.  t y p i c a l taiga forest) and where these are absent, so i s  the sable.  I t i s nocturnal, and spends more time on the ground  than do Martes f o i n a and M. martes.  I t s food i s c h i e f l y small  nuts, b e r r i e s and a l l small animals to the size of a rabbit or grouse.  The den i s made i n the crevices of roots and rocks.  In February they undergo a "False drive" which l a s t s f o r three or four weeks. by mating.  In July i s the r e a l "drive" which i s followed  The gestation period i s nine to nine and a half  months (270 - 285 days).  The young number two to four, r a r e l y  f i v e or s i x and are born i n A p r i l , or e a r l y May. blind. for  They are born  On the 31st to 34th day, t h e i r eyes open.  They suckle  about two months and i n the s i x t h or eighth week f i r s t leave  the nest.  By August they are f u l l y grown.  Maturity i s reached  i n the second year, according to one author, the t h i r d year according to another and the f i f t h according to a t h i r d .  "False  drive" occurs i n the f a l l , reaching i t s extremity i n October (Bobrinskoy et a l . 1944, Kozhantschikov  1930, Schmidt 1943,  Ponomarev 1938). V a r i a t i o n i n the sable i s rather great.  Ponomarev  (1938) says that within a given region albinism, semi-;albinism, white spotting, and a general mottling occur.  On the Island  55 of F e k l i s t o n white headedness occurs, which he attributed to the a c t i v i t y of a single recessive gene.  Normal sable colour, he  believed i s a t y p i c a l quantitative character controlled by seve r a l genes.  The colour and shape of the throat patch he  attributed to a single gene. Geographic v a r i a t i o n i s likewise marked.  According to  the same author the center of darkest colouration i s found on the northeast coast of Lake B a i k a l . Kozhantshikov (1930) reported that the characters assigned to the continental subspecies of sable are v a r i a b l e and i n s i g n i f i c a n t but that i n s u l a r forms and those from Kamtschadka were quite d i s t i n c t i v e but probably derived from the continental form.  He concluded  that there were e s s e n t i a l l y only two types  (subspecies ?) of sable - the continental and those  remaining.  Ognev (1925) came to somewhat d i f f e r e n t conclusions. colour, at l e a s t , he concluded, occurs.  In  decided geographic v a r i a t i o n  According to him, the animals of the mountains are small,  with very dark pelage, the underfur being c h a r a c t e r i s t i c a l l y slaty. The sables of the forest and low country are l a r g e r , with paler fur, less slaty underfur, and generally l e s s dense, less soft pelage. est  The sable of the very northern tundra-taiga i s the l a r g -  sable of a l l but i t s f u r i s coarse and l i g h t , the underfur  yellowish. Bobrinskoy et a l . (1944) state that the chief characteri s t i c s marked by geographic v a r i a b i l i t y are s i z e , colour of f u r , and q u a l i t y of f u r .  The largest sables, he says, are found i n  two l o c a l i t i e s , one being the western parts of i t s range ( A l t a i  56  and the Ob lowlands), the other i n Kamtschadka.  Medium and  small s i z e d sables occur i n c e n t r a l S i b e r i a and i n the AmuroUssuriski d i s t r i c t s .  Colour and s i l k i n e s s of f u r change  correspondingly and are e v i d e n t l y conditioned by the continenta l i t y of the climate.  The darkest and s i l k i e s t l i v e i n the  Transbaikal and southern Y a k u t i a regions, whereas to the east and west of these p l a c e s , the f u r g r a d u a l l y becomes l i g h t e r coarser5  and  however, i n the mountainous regions the colour again  turns somewhat darker.  I t i s i n t e r e s t i n g to note that i n  n e i t h e r the sable nor the European martens does Bergmann's Rule appear to s t r i c t l y apply (above named authors and Schmidt 194-3). Characteristics:  An a c t i v e form;  weight u n c e r t a i n , but prob-  ably more than i n M. martes and M. f o i n a ; body l e s s than 57 cm.;  length of head and  f u r u s u a l l y f i n e , s o f t , brown both above  and below, o f t e n grading to b l a c k i s h , underfur brown to yellowish,  to s l a t e and s t e e l grey;  than spine;  top of head l i g h t e r i n colour  throat patch o f t e n absent, or at l e a s t very s m a l l ,  r a r e l y reaching to fore l e g s ;  dusky salmon i n colour;  l i p w i t h d i s t i n c t v e r t i c a l groove;  upper  t a i l s h o r t , w i t h h a i r end-  ings l e s s than one h a l f l e n g t h of head and trunk and b a r e l y reaching to end of outstretched l e g s ;  t a i l about f i v e to seven  inches l o n g ;  t a i l vertebrae v a r i o u s l y numbered as 15 to 16,  18, 16, 12-15  (Bobrinskoy et a l . 1944,  13,  Ognev 1931j Gray 1869,  de B l a i n v i l l e 1841, Flower 1885, Schmidt 1943, Owen 1853); endings of t a i l l e s s than quarter t a i l l e n g t h ;  hair  skull light,  never massive, elongate as i n M. americana americana, not short as i n M. f o i n a , narrow and deeper w i t h nose somewhat produced;  57  condylobasal length 70 - 90 mm., width about 48 mm.,  r a r e l y 96 mm.;  zygomatic  about one half condylobasal length;  zygo-  matic arches highest at rear, whence they slope suddenly downward and forward, tend to be narrow, long; only r a r e l y absent; and long;  s a g i t t a l crest heavy,  rostrum sides convergent, but very narrow  nasal bones with only s l i g h t c o n s t r i c t i o n ;  frontal  p r o f i l e not steeply sloping, concavity of p r o f i l e not pronounced; supraorbital c o n s t r i c t i o n moderate, pointed, rather deep, always present;  p o s t o r b i t a l process midway between point of greatest  c o n s t r i c t i o n and anterior root of zygoma, i s usually weak, often lacking;  p o s t o r b i t a l region short and convex;  palate r e l a t i v e l y  narrow and long, often with a d i s t i n c t azygos notch on i t s post e r i o r edge; and extended;  anteorbital foramen small;  braincase high, narrow  bullae long and placed close together so that  twice the distance between carotid canals equals the greatest length of the b u l l a ;  the distance between the outer edges of  the jugular foramen i s l e s s than the greatest length of the b u l l a ; distance between carotid canals i s less than one h a l f the distance between the foramen lacerum posteriore and the eustachian foramen; mastoid process f a i l to extend beyond edge of meatus; dentition weak;  upper premolars not crowded;  general  PM^ rather large,  4 strongly convex on inner, strongly concave on outer side;  PM  with inner cusp placed at right angles to trenchant part, large and thick, but with width less than t o t a l length, although nearly  4 equal to width of trenchant part: molar;  PM  longer than width of upper  M^ wide, inner moiety large (as i n M. martes) twice as  large as outer part and one t h i r d longer, waist strongly  58  constricted;  with well developed inner tubercle (hypoconid)  at base of main cusp;  unlike M. martes and M. foina the l a s t  several caudal vertebrae a l l possess transverse process (according to Schmidt 194-3) baculum 38 mm. long, of the Martes type, bending gradually upwards throughout i t s d i s t a l t h i r d , ending often i n a biforked t i p , according to Ognev (1931) hut equally often the ends of bifurcations uniting to produce a transverse perforation. Martes z i b e l l i n a z i b e l l i n a  (Linnaeus 1758)  Type l o c a l i t y : "Surroundings of Tobolsk, Tomsk Government, S i b e r i a " (Ognev 1925). Range:  Pechora basin, northern Urals, and Ob P l a i n (Ognev 1925)  and 1931» Bobrinskoy et a l . 1944, Ellerman, Morrison-Scott 1951). With long nose, short braincase.  Colour d u l l and pale even i n  dark i n d i v i d u a l s , varying from cinnamon drab to dark brown, the underfur retaining i t s lightness of t i n t .  In the Urals tends  to be smaller, with darker denser f u r , with a large bright throat patch (Ognev 1925). Condylobasal length;  7.83  males 8.59 (8.39 - 8.92) n = 6;  females  (7.73 - 7.93) n = 4.  Martes z i b e l l i n a arsen.levi  Kuznetzov 1941 (according to Bobrin-  skoy et a l . 1944). Type l o c a l i t y :  Ussuri Basin, eastern S i b e r i a .  Range and c h a r a c t e r i s t i c s uncertain. Martes z i b e l l i n a averini Type l o c a l i t y :  Bashanov 1943.  Katon-Karagai region, southern A l t a i .  59  Range and c h a r a c t e r i s t i c s uncertain (Ellerman, Morrison-Scott 1953), but occurs i n Mongolia, according to Bannikov (1953). Martes z i b e l l i n a brachyura Type l o c a l i t y : Range:  (Temminck  1844)  Yeso, Japan.  Hokkaido Island, Japan, and the K u r i l e Islands.  I n f e r i o r to the continental forms i n fineness and length of fur;  back and t a i l dark brown, sides and limbs l i g h t e r ;  of feet long, concealing the claws;  t a i l short,  hair  only, 3»5  inches i n length (Temminck). Martes z i b e l l i n a hamgyenensis  Kishida 1927  (In Dobuts Zasshi  Tokyo 39 : 509). Type l o c a l i t y : Range:  Korea.  so f a r as known, the range of the species i s Korea,  possibly north to near the type l o c a l i t y of M. z. arsen.ievi. Characteristics uncertain. Martes z i b e l l i n a kamtshadalica Mus.  ( B i r u l a 1918)  i n Comptes Rendus  Zool. Acad. S c i . Petrograd, v o l . 821)  Type l o c a l i t y : Range:  Kamtchadka.  the Kamtchadkan Peninsula.  A strongly characterized  sable, being much larger and with more robust d e n t i t i o n than any other subspecies.  Colour ranges from warm sepia to mars  brown, throat patch variable (Ognev 1925)• Condylobasal 8.08  (7.86  length;  - 8.41)  n =  males 8.95 (8.50 - 9*50) n = 22;  females  15.  Martes z i b e l l i n a princeps  ( B i r u l a 1922)  Acad. S c i . , St. Petersberg  22:)  In the Ann. Mus.  Zool.  60  Type l o c a l i t y : Range:  Baragusin Mountains,  Transbaikalia.  the mountain forests of Transbaikalia, i . e . i n the  Baragusin H i l l s and the branches of the Stanavoy Mountains, through parts of Mongolia. Witim Rivers.  and  Formerly between the Olekma and  A medium sized sable, near that found i n the  Sajan Mountains, but with much longer bullae and braincase.  Fur  long, dense and soft, of a dark black-brown, with underfur b l u i s h grey and with brown at base of t i p . i n v i s i b l e (Ognev  Condylobasal length: 7.48  (7.43  -  7.53)  Bannikov  1925,  males  Range:  1953).  8.10  (7*82  -  8.35)  n =  9}  females  n = 5.  Martes z i b e l l i n a sahalinensis Type l o c a l i t y :  Throat patch reduced, often  Ognev 1 9 2 5  Wedernikovo, Sakhalin Island.  Sakhalin Island.  S k u l l d i s t i n c t i v e , fine and weak, with  strong contraction i n the i n t e r o r b i t a l region and with short bullae.  Dentition weak.  Colour, as i n M. z. kamtshadallca,  but commonly more cinnamon (Ognev Condylobasal length: 7.41  (7.26  -  7.59)  1925).  males 7 . 9 7 ( 7 . 6 3 - 8 . 2 0 ) n = 1 3 ;  n =  females  6.  Martes z i b e l l i n a sa.lanensis Ognev 1 9 2 5 . Type l o c a l i t y :  Orsyba River, northern Sayan Mountains,  middle  Siberia. Range:  the mountain forests of northwestern Mongolia including  the Usa and Kasyr Rivers. i a l regions.  S k u l l short, both i n nasal and cran-  Colour dark brown, the underfur pale yellowish.  Throat patch varies from dusky brown to b r i l l i a n t salmon (Ognev 1925,  A l l e n 1 9 3 8 - 4 0 , Bannikov 1953).  61  Condylobasal length: 7.36  males 8.24 (8.16 - 8.40) n = 6;  females  (7.32 - 7-40) n = 2.  Martes z i b e l l i n a schantarica Kuznetzov 1941  (according to Bobrin-  skoy et a l . 1944). Type l o c a l i t y : Range:  Shantar Islands, lower Amur, eastern S i b e r i a .  the Shantar Islands. According to Bobrinskoy i s very  l i k e M. z. sahalinensls. Martes z i b e l l i n a tungusensis  Kuznetzov 1941  (according to  Bobrinskoy et a l . 1944). Type l o c a l i t y :  Basins of the Nizhaya and Podkamennaya Tungush,  middle S i b e r i a . Range:  presumably as the type l o c a l i t y .  Characteristics  un-  certain. Martes z i b e l l i n a yeniseensis Type l o c a l i t y :  Ognev 1925.  Forest on p l a i n along Yenesei River, Krasnoiarsk  d i s t r i c t , eastern S i b e r i a . Range:  Taiga between the Angerra and Sayan f o o t h i l l s  et a l . 1944, Ellerman, Morrison-Scott 1951, to M. z. z i b e l l i n a  (Bobrinskoy  Ognev 1925).  Near  but with shorter nasal region, longer c r a n i a l  region, wider zygomatic arches and p o s t o r b i t a l processes.  The  colour of the fur i s a more dusky warm brown. Condylobasal length: 7.43  males 8.27 (8.00 - 8.70) n = 13;  (7.20 - 7.70) n = 7.  females  Ognev (1925) suggested that another  d i s t i n c t , but un-named, subspecies was to be found i n the Tunguska River region, near Turuhansk.  Bobribskoy 1944 suggests d i s -  t i n c t i v e forms may occur i n the A l t a i and i n southeastern Yakutia.  62  Martes melampus (Wagner 1841) Type l o c a l i t y : Range:  Southern Hondo, Japan (Thomas 1905a)  Hondo, Shikoku, K i u s h i i and Tsushima Islands, Japan and  western Korea. Specimens examined:  eight from Asia.  Skull diagrams and photographs:  Temminck 1844, f i g s . 3 and 4.  This appears to be a poorly understood animal;  at  least I have been able to locate almost nothing on i t s structure, l i f e history, ecology or true relationships to the other forms. Some data may possibly be obtained from Okada 1938, or Kuroda 1940 which papers I have not seen. have t h i s to say:  Ellerman, Morrison-Scott  "In the London material, t h i s has the t a i l  an average about 4 4 - 4 7 per cent of the head and body length (resembling z i b e l l i n a therefore i n rather short t a i l ) ;  a white  throat patch seems f a i r l y constant, and, at least i n winter, the  head tends to be paler than the back, a l l characters remin-  iscent of z i b e l l i n a except the throat patch.  But the bullae  seem to be d e f i n i t e l y of the m a r t e s — f o i n a type, and do not seem to resemble those of z i b e l l i n a .  The forelimbs are c l e a r l y con-  trasted blackish, more so than i n our z i b e l l i n a skins, therefore the  conclusion has been reached that melampus i s an i s o l a t e d and  v a l i d species, p a r t l y combining the characters of the other two groups. the as:  So f a r as colour i s concerned, i t i n no way  subgenus Charronia...."  resembles  Described by Gray 1865 and 1869  f u r s o f t , brown or yellow brown, underside scarcely paler,  shoulder and outside of thigh blacker;  feet blackish; head,  chin and upper part of throat dark red brown; from orbit to nose.  blackish streak  Throat and sides of neck yellow or white,  63  crown paler. Length of head and body l e s s than 57  Characteristics.  fur a r i c h golden brown to a deep brown; i n colour than the spine; i n colour; groove;  cm.;  top of head l i g h t e r  throat patch usually present, white  upper l i p presumably with a d i s t i n c t v e r t i c a l  t a i l short, with h a i r endings about 44 to 47 per cent  length of head and body;  s k u l l very massive, f o i n a shaped, or  perhaps intermediate between f o i n a and martes, not quite so broad, low and short as i n M. f o i n a . Condylobasal length 84 to 85 mm.; h a l f condylobasal  length, averaging  zygomatic width greater than about 48 mm.;  zygomatic  arches highest at rear, whence they slope suddenly downward and forward;  r e l a t i v e l y broad and short;  r a r e l y absent; and short;  s a g i t t a l crest strong,  rostrum sides convergent, the rostrum being wide  nasal bone's with no c o n s t r i c t i o n ; nasal p r o f i l e  rather sloping, with pronounced concavity; striction slight;  supraorbital con-  p o s t o r b i t a l process near middle or s l i g h t l y  nearer point of greatest c o n s t r i c t i o n than anterior root of zygoma, i s strong, coming to a d i s t i n c t point. region long and straight; and short (86 mm.)  flated;  palate r e l a t i v e l y broad (.09  mm.)  with d i s t i n c t azygos notch on rear surface;  anteorbital foramen small; extended;  Postorbital '  braincase broad, not r e l a t i v e l y  bullae short, as i n M. f o i n a , and rather l e s s i n -  widely spaced, so that the distance between the outer  edges of the jugular foramen i s greater than the greatest length of the bullae;  so that the distance between the carotid canals  equals about five-sevenths of the distance between foramen lacerum posteriore and the eustachian tube foramen;  mastoid  64 process extends somewhat beyond edge of meatus;  general denti-  t i o n stronger; upper pre-molars sometimes rather crowded; 3 4 PM convex on inner side, concave on outer; PM with inner cusp heavy, i t s width equalling i t s length and equalling threefourths of the width of the trenchant part placed diagonally to 4 1 axis of main cusp; length of PM greater than width of M ; 1 M  large, massive, inner moiety large, intermediate between  that of M. martes and M. foina, one and a half to two times as large as outer moiety, and twice as long; well developed;  M2 the same;  Mi with inner cusp  baculum presumably of the Martes  type, though as f a r as I know, never p o s i t i v e l y determined as such. Martes melampus melampus Type l o c a l i t y :  (Wagner 1841)  Japan (Wagner 1841), Southern Hondo, Japan  (Thomas 1905a, b, and c ) . Range:  Hondo, Shikoku and Kiushiu Islands, Japan.  With a r i c h  golden brown pelage, the back usually much browner and with a bright orange throat patch (Thomas 1905a, b, and c ) . basal length:  males 8.52  (8.50 - 8.55)  n = 2;  Condylo-  females  7.50  n = 1. Martes melampus coreensis Type l o c a l i t y : Range:  Korea.  Kuroda and Mori 1923.  Tenan, southern Chusei d i s t r i c t , Korea. Near to M. m. melampus but general colour of  upper parts yellow washed with fox-red instead of r i c h orange yellow and the ashy white of head extending farther to the nape instead of confined to the forehead.  Separated from  65  M. m. t s u e n s i s by much p a l e r c o l o u r a t i o n throughout. Martes melampus tsuensis (Thomas 1897) Type l o c a l i t y : Range:  Kamoze, Tsuehima I s l a n d s , Japan.  Tsuehima I s l a n d s , Japan.  S i m i l a r to the t y p i c a l form  i n a l l characters but colour of f u r .  General colour above i n  winter d i r t y y e l l o w i s h brown instead of r i c h golden brown; muzzle and l i p s b l a c k instead of brown; instead of yellow.  throat patch white  In summer pelage much darker than the t y p i c a l  form, a l l brown parts being b l a c k as are the whole of the l i m b s ; throat patch yellow (Thomas 1897). Condylobasal length:  females 7.80 n - 1.  Martes americana ('.."Kerr.: 1792) i n c l u d i n g "M. caurina" (Merriam 1890) Type l o c a l i t y : Range:  eastern North America.  H o l a r c t i c North America. Because t h i s species w i l l be t r e a t e d i n d e t a i l e l s e -  where, only those aspects needed f o r comparison to E u r a s i a n spec i e s and the f i s h e r w i l l be u t i l i z e d here.  Because the two sub-  species to be considered d i f f e r m o r p h o l o g i c a l l y , comparison w i l l be made f o r both of these. Characteristics: limbs.  A c t i v e forms, body long and s l i m w i t h short  Length of head and body l e s s than 57 cm.;  weighs about  700 - 1200 gms. i n males, 600 - 800 gms. i n females;  outer f u r .,  dark brown, under f u r longer i n americana than i n c a u r i n a , l i g h t yellow i n both;  top of head l i g h t e r i n colour than i s spine;  throat patch i n americana u s u a l l y present, v a r y i n g from white to  66 orange, but usually yellow, i n caurina, always present, usually large (extending to base of forelegs), yellow or orange; l i p with d i s t i n c t v e r t i c a l groove;  upper  t a i l i n americana short with  hair endings about one half a head and body length;  i n caurina,  t a i l somewhat shorter equalling less than one half the head and body length;  t a i l vertebrae average 17 (Cooper, 1953) i n ameriAccording to H a l l (1926a) i t possesses an  cana, 14 i n caurina. abdominal skin gland.  S k u l l i n americana l i g h t , narrow, elon-  gate^ nose rather produced; i n posterior part;  braincase not massive, e s p e c i a l l y  i n caurina more massive, broader, f l a t t e r ,  nose less produced;  braincase more massive, e s p e c i a l l y i n pos-  t e r i o r part; condylobasal length 67 mm. to 88 mm.;  zygomatic width about 48  mm. i n both, i n americana being about one h a l f condylobasal length, i n caurina s l i g h t l y more;  arches i n americana highest  at rear, whence they slope suddenly downward and forward, being narrow and long; est  i n caurina regularly curved, broadest and high-  near t h e i r middle, broader and shorter;  s a g i t t a l crest i n  americana only r a r e l y absent, i n caurina often weak or absent; rostrum, i n americana narrow and long, r o s t r a l sides p a r a l l e l ; i n caurina short and wide, sides convergent;  i n americana anter-  i o r nares oval, nasal bones with moderate c o n s t r i c t i o n ; p r o f i l e rather sloping, concavity pronounced;  frontal  i n caurina  anterior nares more heartshaped, nasal bones with no c o n s t r i c t i o n , f r o n t a l p r o f i l e less sloping, concavity less pronounced;  supra-  o r b i t a l c o n s t r i c t i o n i n americana moderate, always present, i n caurina, slight, sometimes absent;  p o s t o r b i t a l process i n ameri-  cana midway between point of greatest c o n s t r i c t i o n and anterior  67  root of zygoma, usually weak; convex;  i n caurina  p o s t o r b i t a l region short and  much nearer point of greatest c o n s t r i c t i o n  than anterior root of zygoma and stronger, p o s t o r b i t a l region long and s t r a i g h t ;  palate r e l a t i v e l y long and narrow, often  with azygos notch on posterior surface i n americana, i n caurina shorter and broader, usually truncate behind; foramen r e l a t i v e l y large, i n both;  anteorbital  braincase markedly elongate,  more so than i n any other marten, i n americana; broader and lower i n caurina;  less extended,  bullae i n americana long, narrow,  highly i n f l a t e d and situated near to each other, so that twice the distance between the carotid canals i s the same or s l i g h t l y less than the greatest length of b u l l a ;  distance between the  outer edges of the jugular foramen i s l e s s than the greatest length of bullae;  distance between the carotid canals i s one  half the distance between the foramen lacerum posteriore and the eustachian foramen;  i n caurina the bullae are less i n f l a t e d ,  shorter, broader and more widely separated  so that of the three  measurements above,the f i r s t two are greater than the greatest length of b u l l a , and the distance between the carotid canals equals two-thirds the distance between the other two  foramina;  i n americana the mastoid process does not extend beyond the meatus, i n caurina i t does;  general d e n t i t i o n i n americana  weaker, upper premolars only occasionally crowded, i n caurina stronger, premolars often crowded; on inner, concave on outer side;  PM  i n both PM 4  strongly convex  i n americana with inner  cusp small, width being much less than length and only h a l f the width of the trenchant part, placed nearly at right angles to  68 main axis of tooth, i s as long as M  1  i s wide;  i n caurina PM  4  has the inner cusp large and thick, i t s width nearly equal i t s length and more than one half the width of trenchant part and placed diagonally to main axis of tooth, not as long as M wide;  1  is  M^" i n americana small, quadrate, always smaller than i n  Old World forms, inner moiety small, very l i t t l e larger or longer than outer moiety and one t h i r d smaller than i n caurina, i n the 1 l a t t e r form M  i s large, massive, inner moiety one and a half  to two times as large as outer moiety and twice as long;  M  with an inner cusp present i n both, more often absent i n americana than i n caurina;  M  2  with inner cusp weak or absent i n  americana, present and well developed i n caurina;  baculum i n  both of the Martes type, bending gradually upwards throughout i t s d i s t a l t h i r d , ending i n two prongs, or more, usually uniting to produce a transverse perforation. Two subspecies are accepted i n t h i s paper. Martes americana americana ('.Kerr,. 1792) Type l o c a l i t y : eastern North America. fiange:  North America west to the Coast Range, excepting the  Rocky Mountains south of central Idaho and Montana;  also New-  foundland, Cape Breton Island and Prince Edward Island. Martes americana caurina Type l o c a l i t y : Range:  (Merriam 1890)  Gray's Harbor, Chehalis County, Washington.  the Coast Mountains, Coast Ranges, Cascade Mountains,  S i e r r a Nevada and the Rocky Mountains, south of central Idaho and Montana, also Vancouver Island, the Alexander Archipelago and the Queen Charlotte Islands.  69  Martes pennanti Type l o c a l i t y : Range:  (Erxleben  1777)  Eastern Canada  H o l a r c t i c North America As i n the case of M. americana, only those aspects  important i n comparison to other species are made here. Characteristics:  Active;  elongate with short limbs;  of head and body usually greater than 57 cm. to 62 cm.);  (ranging from 50  weight from 1500 to 4500 grams;  soft than i n the martens;  length  f u r coarse, less  colour dark brown to blackish; outer  fur brown-black with often a white subterminal area on each hair; white;  usually without a throat patch, when present small and generally darker below than above;  t a i l , without hair  t i p s about one half the length of head and body, at l e a s t t h i r d the length;  t a i l r e l a t i v e l y long, with variously 20 to  21 vertebrae (Kneeland 1859» Gerrard 1862); (1926a)  lacking an abdominal skin gland;  although large; duced;  according to H a l l  s k u l l not massive,  s k u l l elongate, narrow, deep;  braincase not massive e s p e c i a l l y behind;  length usually more than 95 mm., zygomatic width from 55 to 60 mm., dylobasal length;  nose rather procondylobasal  varying from 92 to 122  arches highest at rear, whence they slope  crest strong, only r a r e l y absent;  stricted;  mm.;  being more than one half con-  suddenly downward and forward, are broader and shorter;  sides p a r a l l e l ;  one  rostrum long, narrow;  anterior nares oval;  sagittal rostral  nasal bones s l i g h t l y con-  f r o n t a l p r o f i l e rather sloping, concavity pronounced;  supraorbital c o n s t r i c t i o n moderate, always present;  postorbital  process about midway between point of greatest c o n s t r i c t i o n and anterior root of zygoma, usually weak;  p o s t o r b i t a l region ( i n  70  front of point of greatest constriction) short and convex; palate r e l a t i v e l y long (65 mm.)  and narrow (10 mm.) usually  with d i s t i n c t azygos process on rear; braincase l i g h t , extended;  antorbital foramen small  bullae long, but rather widely  spaced so that twice the distance between the carotid canals i s greater than the greatest length of b u l l a ;  the distance  between the outer edges of the jugular foramen i s the same or s l i g h t l y more than the greatest length of b u l l a ;  the distance  between the carotid canals equals two thirds the distance between the foramen lacerum posteriore and the eustachian foramen;  the mastoid process extends beyond the edge of the meatus  general d e n t i t i o n weak;  upper premolars not crowded;  PM^ con-  4  vex on inner side, concave on outer;  PM  with inner cusp small  width less than i t s length and only half that of trenchant part placed diagonally to main axis of tooth, i s longer than M i s 1  wide;  M  1  i s l i g h t , quadrate but with inner moiety small as i n  M. americana americana, only s l i g h t l y larger than outer lobe; waist moderately constricted; M^ has well developed inner cusp at base of main cusp;  M  2  likewise with strong inner cusp;  baculum of the Martes type, gradually bending upward throughout i t s d i s t a l t h i r d , and with either two prongs at the t i p , or these united to form a transverse perforation. Of supposedly three subspecies, here not accepted. Martes f l a v i g u l a (Boddaert 1785) = Charronia f l a v i g u l a  Pocock  1918 = Lamprogale f l a v i g u l a Ognev 1928 = Martes guatkinsi  Hors  f i e l d 1851. Type l o c a l i t y :  Unknown, but considered by Pocock to have been  71  fixed by t r a d i t i o n as Nepal. Range:  The Amur and Ussuri regions of eastern S i b e r i a , south  through most of China, Tibet, Burma, Assam, thence westwards to Kashmir and North-West F r o n t i e r , and south into  Indo-China,  Siam, Malay States, Formosa, Sumatra, Java, Borneo and Banka; also the N i l g i r i H i l l s , Coorg and Travancore, southern India (Sowerby 1916, Ellerman, Morrison-Scott 1951,  Brongersma 194-1).  Brongersma adds that other unverified records exist i n the l i t e r a t u r e f o r Palawan, Great Naturna, Balabac, and  Calamianes,  Cuyo, Subu, Sibutu, the Paternosters and Sebuko Islands. Jerdon (1874-) says i t occurs on Ceylon, but l a t e r authors do not authenticate t h i s . Specimens examined:  32 from A s i a .  S k u l l diagrams and photographs:  Blanford 1888-91, p.  Ognev 1931, v o l . 2, pp. 635, 636 and 637; Brongersma 194-1, p i . 4 and f i g . 1; pp. 51 and 52.  157;  Pocock 1936b, f i g . 2;  Colbert and Hooijer 1953,  Other sources are given by Ognev 1931, v o l . 2.  According to A l l e n (1938-40) i t i s an animal of wooded mountainous country, f a i r l y well d i s t r i b u t e d over most of southern  and central China south of the Gobi.  The same author  records t h e i r feeding on honey bees, which are apparently a favourite food.  They are also reported to feed on nectar and  to run down and k i l l fawns of the barking deer.  See also  Appelman 1940, and Hutton 194-9, hot examined during t h i s study. Characteristics:  Length of head and body usually more than 57  cm. (Varying from 50 to 80 cm.); above and yellow below;  general colour black-brown  underchin white, chest, neck and under  72  b e l l y yellow, occasionally tending to white;  throat patch  almost always extending so as to include the b e l l y ; never with v e r t i c a l groove;  upper l i p  t a i l long, but not bushy, i t s  length (without hair t i p s ) being two thirds the head and body length;  t a i l vertebrae 24 (Blanford 1888-91, Owen 1853);  of feet covered with hair (excepting i n i n s u l a r forms); massive, swollen, broad and f l a t , with short nose; length 90 to 110 mm., width about 60 mm., length;  skull  condylobasal  occasionally as small as 70 mm.;  zygomatic  always greater than one half condylobasal  arches regularly curved, broadest and highest at t h e i r  middle, broader and short;  s a g i t t a l crest absent or very weak;  rostrum wide and short with convergent strongly constricted; pronounced; present;  soles  sides;  nasal bones  f r o n t a l p r o f i l e not steep, concavity not  supraorbital", c o n s t r i c t i o n moderate, pointed, always  p o s t o r b i t a l process about midway between point of  greatest c o n s t r i c t i o n and anterior root of zygoma, often absent, always weak;  p o s t o r b i t a l region short and convex;  and short, truncated at hinder margin;  palate wide  anteorbital foramen small;  braincase massive, e s p e c i a l l y at hinder end; bullae short and widely spaced so that twice the distance between the carotid canals or the distance between the outer edges of the jugular foramina are both greater than the greatest length of b u l l a ;  dis-  tance between the carotid canals i s two thirds the distance between the foramen lacerum posteriore and the eustachian foramen;  bulla  with only two chambers, never three or more as i n M. martes or M. foina;  mastoid processes do not extend beyond the edge of  the meatus; crowded;  general dentition weaker;  PM  J  upper premolars not  strongly convex on inner, concave on outer side;  73  PM  with inner cusp small, width less than length and much less  than width of trenchant part, placed at right angles to axis of main part, longer than width of M ; 1  M small to moderate i n size, 1  inner moiety small, of about same length and size as outer moiety, waist only s l i g h t l y constricted; lower molars are unknown to me;  the d e t a i l s of the  baculum d i s t i n c t i v e , being  e a s i l y separated from a l l other species of the genus, as pointed out by Pocock, curving up abruptly i n i t s d i s t a l s i x t h , i t s t i p being nearly v e r t i c a l , length 65 - 80 mm.,  the t i p i s divided  into four subsymetrically arranged processes, each with a rounded condyle-like head (for d e t a i l s see Pocock 1918,  1936b and  1941,  and Ognev 1931)• The two most recent c l a s s i f i c a t i o n s of the species ( i n cluding M. guatkinsi) have been made by Pocock (1936b), Morrison-Scott  (195D  and Chasen (1940).  A l l of these  Ellerman, authors,  as well as considering M. f l a v i g u l a and M. guatkinsi d i s t i n c t , divide M. f l a v i g u l a into eight subspecies terminology f o r two of them).  (using quite d i s t i n c t  The c l a s s i f i c a t i o n of the l a t t e r  two authors i s given below, subjugating M. guatkinsi to subspecif i c status. Martes f l a v i g u l a f l a v i g u l a (Boddaert Type l o c a l i t y : Range:  1785)  fixed by Pocock as Nepal.  Kashmir to Tibet and upper Burma and southern  north to Shensi, Kensu (Ellerman, Morrison-Scott 1936b). of M^,  1951)  China, Pocock  Characterized by the narrowness of the inner lobe which i s the same width throughout and i n no way widened  or flattened on i t s innermost margin (Bonhote 1901).  Muzzle  7  and top of head black or dark brown;  shoulders and back tawny  brown to nearly white.  Abdomen brown to whitish;  yellow to nearly white;  h a i r on rear foot extending back to  and beyond the plantar pad. hair.  4  throat r i c h  In India the soles are without  (Jerdon I874, Bonhote 1901, Pocock 1936b).  Condylobasal length, males 10.06 (9.05 - 10.60) n = 19; females 8.97  (8.20 - 9.70) n = 20.  Martes f l a v i g u l a aterrima  (Pallas l 8 l l ) = M. f . borealis Radde  (1862) of Pocock. Type l o c a l i t y :  between the Uth and Amur Rivers, eastern S i b e r i a ,  northern China and Korea (Ellerman, Morrison-Scott 1951? 1936b).  Pocock  Rather larger than t y p i c a l f l a v i g u l a and the colour  of the back i s much more yellow, the colour of the throat much more whitish (Bonhote 1901, Pocock 1936b).  According to Howell  (1929) i n d i v i d u a l colour v a r i a t i o n i s exceedingly high i n the area occupied by t h i s subspecies. Condylobasal length:  males 10.72  (10.20 - 11.00) n = 6;  females 9.63 (9.25 - 10.03) n = 2. Martes f l a v i g u l a chrysospila Swinhoe 1866 = M. f . xanthospila Swinhoe (1870) of Pocock. Type l o c a l i t y : Range:  Mountain forests of Formosa.  Formosa.  Described by Bonhote (1901) as intermediate  between the southern forms (peninsularis and henrici) and the t y p i c a l form to the north. long f u r ;  Small i n s i z e , with only moderately  d i f f e r s from t y p i c a l f l a v i g u l a by the head being  deeper brown;  back of the foreleg with a whitish patch (Bonhote  75  1901, Pocock 1936b). Condylobasal  length:  males 9.20 n = 1;  females 8.20 (7.60 -  8.80) n = 2. Martes f l a v i g u l a h e n r i c i Type l o c a l i t y : Range:  (Schinz 1845)  Sumatra.  Sumatra, Banka and part of Borneo (Chasen 1940).  According t o Pocock (1936b) occurs only on Sumatra.  I n short-  ness of f u r and naked s o l e s , resembles.peninsularis;  separated  from i t by i t s smaller s k u l l and darker c o l o u r a t i o n , e s p e c i a l l y on the abdomen (Bonhote 1901, Pocock 1936b).  Brongersma (1940)  says a specimen of Mustela l u t r e o l i n a from Sumatra i n the Leiden Museum was o r i g i n a l l y m i s - i d e n t i f i e d as M. f . h e n r i c i .  Whether  t h i s casts doubt on the v a l i d i t y of the subspecies, o r of the presence of the species on the i s l a n d I do not know. Condylobasal 8.10  length:  males 9.23 (8.75 - 9.70) n = 4;  females  n = 1.  Martes f l a v i g u l a indochinensis Type l o c a l i t y :  Kloss 1916.  Klong Menao, southeastern Slam (Ellerman,  Morrison-Scott 1 9 5 D , Kohtak, southeastern Siam (Pocock 1936b). Range:  Siam, northern Tenasserim and Annam (Pocock 1936b).  D i s t i n g u i s h e d from t y p i c a l f l a v i g u l a and aterrima by a naked area of s k i n between the lobes of the p l a n t a r pad on the hindf o o t , by the shorter winter coat, w i t h a more y e l l o w i s h t i n t , and by the p a l e r patch beneath the eye (Pocock 1936b). Condylobasal  length:  males 10.00 (10.00 - 10.00) n = 2;  females 9.03 (8.80 - 9.20) n = 3 .  7$ Martes f l a v i g u l a peninsularis Type l o c a l i t y :  (Bonhote 1901).  Bankachon or Bankasun, V i c t o r i a Point, Tennas-  serim. Range:  southern Tenasserim through the Malay Peninsula (Pocock  1936b, Ellerman, Morrison-Scott 1951).  Similar to indochinensis  i n having the skin above the hind plantar pads naked, but d i f f e r s i n having the head brown, not black, shoulders and back less yellow and the abdomen dark brown.  The coat i s short and thin,  varying l i t t l e seasonally (Bonhote 1901, Pocock 1936b). lobasal length:  males 9.75 ( ( . 7 0 - 9-80) n = 2;  Condy-  females 8.67  (8.40 - 9.00) n = 4. (Pocock 1936a).  Martes f l a v i g u l a robinsoni Type l o c a l i t y ; Range:  Tjbodas, Java.  Java (Pocock 1936a).  Distinguished from the Malayan  and other i n s u l a r forms i n having the crown, nape and shoulders buffy grey and the l i n e separating the white from the brown on the cheek poorly delineated (Pocock 1936 a and b). females 9.10  Condylobasal length: Martes f l a v i g u l a saba Type l o c a l i t y : Range:  n = 1.  Chasen and Kloss (1932).  near Sandakan, B r i t i s h North Borneo (Pocock 1936b)  Parts of Borneo.  Distinguished from h e n r i c i by i t s  s l i g h t l y smaller s k u l l and i t s s l i g h t l y smaller carnassial teeth (Pocock 1936b). Condylobasal length: 8.00 (7.80 - 8.20)  males 8.82 (8.60 - 8.95) n = 4; n = 4.  females  77 Martes f l a v i g u l a guatkinsi H o r s f i e l d 1851 = M. guatkinsi of authors. Type l o c a l i t y :  Madras, India.  Range: N i l g i r i H i l l s , Coorg and Travancore, southern India. The baculum, according to Pocock (1936b)  d i f f e r s somewhat from  that of the other subspecies of M. f l a v i g u l a i n that the four d i s t a l processes are quite short, the l e f t of the anterior pair being represented merely by a small tubercle.  Skull distinc-  t i v e i n being depressed, as i n an otter's, much f l a t t e r and l e s s convex on i t s dorsal surface than i n the other subspecies °f f l a v i g u l a , the f r o n t a l region being less swollen, with the occiput less sloped and the zygomata less arched. absent.  PM"*" may  be  Colour dark, as i n M. f. saba uniformly dark above;  unlike saba metatarsal area above the plantar pad not naked (Bonhote 1901, Pocock 1936b). Condylobasal length: 9.00)  males 8.90 n = 1;  females 8.60 (8.20 -  n = 2. Six  to eight subspecies of Martes f l a v i g u l a have been  described from China, Mongolia and eastern S i b e r i a . his  Pocock i n  1936(b) r e v i s i o n , reduced the number of these to two,  but  Howell 1929, A l l e n 1938-1940 and Brongersma 1941 say even these two f a l l within the range of i n d i v i d u a l v a r i a t i o n and name a l l of  them simply as M. f l a v i g u l a f l a v i g u l a admitting the possi-  b i l i t y of the existence of the northern race, M. f . aterrima. According to A l l e n , Thomas has concluded that M. f . indochinensis i s an i n v a l i d form and Brongersma (1941) concluded that three insular forms ( h e n r i c i , saba and roblnsoni) a l l f e l l  78  within the range of i n d i v i d u a l v a r i a t i o n and represented a single subspecies M.~f. Brongersma).  only  h e n r i c i (called M. f. l a s i o t i s by  These forms are supposedly characterized by  s l i g h t l y smaller size, thinner pelage and naked f e e t . Martes f l a v i g u l a guatkinsi has been to the present considered  a d i s t i n c t species, i n part at least because i t i s  isolated geographically from M. f l a v i g u l a .  Jerdon (1874) says,  however, that while H o r s f i e l d applied the name to the Indian race, o r i g i n a l l y i t was  south  given to a Himalayan specimen.  Tate 19.4-7 says that M. guatkinsi i s only "sometimes treated as a separate  species".  Pocock (1936b and 194-1) considered  species d i s t i n c t and pointed out what he considered distinctions. gula into two  the  important  L i t t l e i s gained, however, by s p l i t t i n g M.  flavi-  species and I have here considered i t a single form.  To summarize the above, i t appears that while Pocock (1936b, 1941), Ellerman, Morrison-Scott  (195D  and Chasen (1940)  consider nine forms of the species to e x i s t , the recent tendency has been towards s i m p l i f i c a t i o n and these forms may to f i v e as follows: Martes f l a v i g u l a f l a v i g u l a Martes f l a v i g u l a chrysospila Martes f l a v i g u l a h e n r i c i Martes f l a v i g u l a peninsularls Martes f l a v i g u l a guatkinsi  be reduced  79 RELATIONSHIPS OF THE MARTENS The eventual answer to the puzzling problem of the relationship of the world's marten w i l l be answered, l i k e l y through the study of genetics and paleontology. concerning the l a t t e r i s given elsewhere.  Some material  Concerning the  genetics of marten, I know only that E r l i c h (1949)"reports  that  M. f o i n a l i k e most other carnivores, has a 2n chromosome number of 38 with heterogamic (x - y type) males.  I presume that a l l  species of the genus possess the same sort of chromosome structure . This discussion i s concerned p r i m a r i l y with a comparison of the morphology of the various species.  It should be read i n  conjunction with the discussions elsewhere concerned with f o s s i l marten,zoogeography, etc. I have elsewhere discussed the " s p e c i f i c " status of the various species of marten i n the world. Severtsov  I have pointed out that  (I876) considered the Tienschan Mountains of Eurasia  to be the "homeland" of a l l marten and that here M. f o i n a had been crossed with M. martes and M» z i b e l l i n a , but that most other workers have found few or no hybrids elsewhere. Schmidt (1943) concluded M. martes and M. z i b e l l i n a was  that the separation between  considerably greater than that  between M. martes and M. f o i n a .  His d i s t i n c t i o n would thus be:  A.  M.  zibellina  B.  M. martes and M. f o i n a .  Bonhote (1901) related M. f l a v i g u l a upon the basis of i t s s k u l l conformation  to M. f o i n a .  Pocock's work on the baculum  80  of these species l e d to the relegation of M. f l a v i g u l a to a d i s t i n c t subgenus.  When considering the New World forms i n  r e l a t i o n to the Old, we may immediately distinguish the f i s h e r and thus arrive at: A.  M. z i b e l l i n a  B.  M. martes and M. foina  C.  M. f l a v i g u l a (including M. guatkinsi)  D.  M. pennanti  We have not yet considered the relationship of M. a. americana and M. a. caurina to these, and i t i s this problem that we are primarily concerned with here. the various species here suggests a clue.  The "ecology" of Marshall 194-2,  c o r r e c t l y I believe, points out that M. americana, M. martes and M. z i b e l l i n a have, according to available information, so similar a pattern of behaviour and response as to be e s s e n t i a l l y the same.  M. foina, M. f l a v i g u l a and M. pennanti are, on the other  hand, quite s i g n i f i c a n t l y different i n t h e i r biology from each other and from the preceding, and hence might be grouped thus: A.  M. americana, M. martes and M. z i b e l l i n a  B.  M. foina  C  M. f l a v i g u l a  D.  M. pennanti  This scheme d i f f e r s somewhat from that above with regard to M. americana, M. foina, M. martes and M. z i b e l l i n a . It i s of interest to compare them to that given by Gray (1865 and 1869), the l a s t worker to so c l a s s i f y a l l of the world's species of marten.  His scheme follows:  81 A.  M. martes, M. z i b e l l i n a , M. americana and M. melampus  B.  M. foina  C.  M. f l a v i g u l a  D.  M. pennanti  Aside from the introduction of the species M» melampus, not yet considered, i t w i l l be seen that Gray's scheme l i e s nearest to the ecological one of Marshall and the one here accepted.  The c l a s s i f i c a t i o n currently accepted by taxonomists  i s e s s e n t i a l l y t h i s , so that M. f o i n a i s grouped with M. martes M. z i b e l l i n a , M. americana and M. melampus as the subgenus Martes; M. pennanti as the subgenus Pekania, and M. f l a v i g u l a (including M. guatkinsi) the subgenus or genus Charronia or F l a v i g u l a .  It  i s thus within the subgenus Martes that we must search primarily for r e l a t i o n s h i p .  I think we may accept that M. f l a v i g u l a (upon  the basis of i t s peculiar baculum) and M. pennanti (upon the basis of i t s size and biology) are d i s t i n c t i v e from a l l others, although the degree of difference between the subgenera Charronia and Martes i s probably greater and more fundamental than that between Pekania and Martes. i n the past.  There has been disagreement here  Poland (1892) believed M. pennanti to be c l o s e l y  related to M. f l a v i g u l a , so much so that he called the l a t t e r the "Afghan Fisher" and remarked on the disjunction i n the d i s t r i b u t i o n of the two.  Schmidt (194-3) considered M. f l a v i g u l a  the A s i a t i c counterpart of M. pennanti, but these workers did not know of, or i n Schmidt's case did not take seriously, the fundamental d i s t i n c t i o n of the baculum i n M. f l a v i g u l a and t h e i r suggested relationship cannot be accepted.  82  Returning to the subgenus Martes, I can o f f e r l i t t l e concerning M. melampus,  Ellerman, Morrison-Scott (195D  say-  that i t s colouration i s near M. z i b e l l i n a , but that i t s s k u l l i s nearer M. f o i n a and M. martes, and they conclude that i t i s an "isolated and v a l i d species".  Other d e t a i l i s given subsequently.  It has already been shown that early workers i n North America often considered North American marten to be merely representatives of M. z i b e l l i n a or M. martes but never of M. f o i n a or M. melampus.  Between 1771  and 1869> of those workers  who  named New World marten referable to Old World species, eleven referred them to M. z i b e l l i n a and fourteen to M. martes ( f i v e of these r e f e r r i n g them to both species).  The honours then, have  l a i n about equally between M. martes and M. z i b e l l i n a and not to other species.  We may devise from t h i s again a scheme of r e l a -  tionship: A.  1. Martes martes, M. z i b e l l i n a and M. americana 2. M. f o i n a 3 . M. melampus (possibly with M. foina)  B.  1. M. pennanti  C.  1. M. f l a v i g u l a  The e a r l i e r workers on the problem, Gray (1865), Brandt (1855), Baird (1857), A l l e n (1869),  conclusions referred to above.  and Coues (1077) came to the  Gray distinguished M. americana  americana from the Old World forms on the shape and size of the upper molar. pelage.  Brandt and A l l e n employed only c h a r a c t e r i s t i c s of  Baird employed both c r a n i a l and pelage c h a r a c t e r i s t i c s  as did Coues.  A l l related M. americana to M. martes or M. zib-  e l l i n a and Coues, the l a s t of the group, reviewed the findings  83  of the e a r l i e r workers and found closest relationship to M.  zib-  e l l i n a of the two. Coues and the e a r l i e r workers were unfortunate i n that they did not recognize that a d i s t i n c t i v e marten occurred on the west coast of North America (M. a. caurina) and i t must be assumed, b u i l t their case around representatives of M. a. americana.  In 1890 Merriam described caurina and said that i n c r a n i a l  and dental characters i t departed from M. a. americana i n the direction  of M. z i b e l l i n a . Rhoads (1902) reviewed the problem i n some d e t a i l .  M.a.  americana he considered t o t a l l y d i s t i n c t from a l l other forms on the basis of the smallness of the inner lobe of M"*", as f i r s t outlined by Gray and the smallness of the lower molar and the small size of the inner cusp of M . 2  In general shape of s k u l l M. a.  caurina was nearest to M. f o i n a but i n the large size of the inner lobe of the upper molar and the large size of the lower f i r s t molar t h i s form was very similar  to M. martes and M. z i b -  b e l l i n a , not at a l l to M. americana americana to the east.  In  shape of s k u l l , then, Rhoads related M. z i b e l l i n a to M. americana and M. martes;  M. foina to M. a. caurina.  On the basis of  dentition, however, M. a. americana was d i s t i n c t , and M. a. caurr i n a , M. martes and M. z i b e l l i n a closely itself distinctive.  related and M. foina  He concluded of M. a. caurina that " i t s  homologies connect i t f a r more closely with the Eurasian than the American fM. a. americana] type of marten.... isolated  member of the martes - z i b e l l i n a group".  [ I t ] i s an  84 Lydekker (1901  - 1904)  stated that he believed the  marten of North America were so nearly related to M. martes and M. z i b e l l i n a that they should possibly be considered v a r i e t i e s of these. Ognev (193D  He added no d e t a i l . reviewing  M. z i b e l l i n a lay intermediate  the marten of Russia said that  i n c r a n i a l and dental  i s t i c s between M. martes and "M. c r i p t i o n of "M.  only  americana".  character-  Subsequent des-  americana" makes i t clear that Ognev was r e f e r -  ring to M. a. americana and that he was  giving no  consideration  to M. a. caurina. A l l e n i n 1938-40 said only that "Probably ... the sable (martes z i b e l l i n a ) ... finds i t s counterpart  In the Ameri-  can Pine Martens (M. americana and races), but the Beech Marten (M. foina) i s not represented My own  i n the New  World".  i n t e r p r e t a t i o n of the relationship of martens,  based upon examination of s k u l l s of the species involved, leads me to conclude that the above writers are nowhere r e a l l y wrong, and that M. a. caurina i s most nearly related to M. martes and M. z i b e l l i n a .  M. a. americana i s d i s t i n c t i v e with some features  a l l y i n g i t to M. z i b e l l i n a and that M. melampus i s to some degree related to martes and  zibellina.  When I f i r s t began my study of s k u l l s of a l l the species of marten, I was types of s k u l l existed.  impressed by the fact that two At the one extreme was  basic  the short,  broad, and rounded s k u l l , with short widely spaced bullae, and at the other, the long c l o s e l y spaced bullae.  The species of  marten, placed i n order, going from the former type to the  85 l a t t e r , would f a l l i n t h i s sequence:  M. melampus, M. pennanti,  M. f o i n a , M. f l a v i g u l a , M. a. caurina, M. martes, M. and M. a. americana (see Figure 7).  zibellina  The greatest d i v i s i o n  between the two groups occurs between M. melampus and M. pennanti and M. martes and M. z i b e l l i n a .  I f e l t that I could at that  point divide martens into two basic d i v i s i o n s .  I was  puzzled,  however, by the fact that both M. f l a v i g u l a and M. pennanti f e l l so c l e a r l y within the f i r s t group and yet be so d i s t i n c t i v e wise as to merit subgeneric status.  other-  Perhaps M. pennanti should  not be so distinguished, but M. f l a v i g u l a i n view of i t s d i s t i n c tive baculum, biology, d i s t r i b u t i o n and antiquity, as shows i n the palaeontological record could not be so ignored.  It seemed  that the two schemes of c l a s s i f i c a t i o n were mutually exclusive. In attempting to resolve the problem, help was  given by Alston  (1879) who remarked on the following findings of Blasius, with which Alston himself agreed:  that the differences of proportion  i n the skulls of M. martes and M. f o i n a are less conspicuous when a s k u l l of an aged M. f o i n a i s compared to that of a young M. martes, than when individuals of the same age were contrasted. The implication that was  made was  that at a younger stage M. martes  i s more s i m i l a r to M. f o i n a than i t is- l a t e r on i n i t s l i f e . I myself have been fortunate enough to examine f i v e skulls of very young marten (perhaps two or three months old)  two  referable on geographic grounds to M. a. americana and three to M. a. caurina.  I have been led to conclude from these that at  t h i s very young stage, the skulls of the two races are nearly indistinguishable, and what i s more, both are marked to a greatly  Figure 7 (To face page  86.)  F i g u r e 7«  R a t i o of canine width to b u l l a l e n g t h i n the s k u l l s of the world's marten. This ratio approximates an index combining the c h a r a c t e r i s t i c s of s k u l l l e n g t h , height and breadth.  SKULL  LONG, N A R R O W ,  HIGH,  TOGETHER  86 exaggerated degree by a s k u l l conformation of the foina - caurina type with short widely spaced bullae, broad low braincase and short rather broad rostrum, rather than the elongate s k u l l of the z i b e l l i n a americana type. I conclude that the short, broad and low s k u l l i s chara c t e r i s t i c of a l l martens i n t h e i r youth;  that i n some (e.g.  M. pennanti, foina, f l a v i g u l a , caurina and martes) i t i s l o s t only to a s l i g h t degree with maturity.  In the remaining species  (M. z i b e l l i n a and M. americana americana) while i n youth they have the same s k u l l type, with maturity i t becomes elongate, high, narrow, and with long bullae.  Thus, again r e f e r r i n g to  Figure 7, i t i s hypothesized that the pennanti - melampus group are neotenic;  that i s , the development of the s k u l l i s either  primitive or retarded, while the americana - z i b e l l i n a end of the scale i s a result of hypermorphosis  or acceleration i n devel-  opment, or possibly i t i s a primitive condition ( i . e . and hypermorphosis  are mutually exclusive conditions;  neoteny which of  the two exists i s not known). While I have no means at present of testing t h i s hypothesis f o r the time being I accept i t . accepted, certain conclusions follow.  I f i t i s to be  The most important of  these i s t h i s , that while a l l species possess skull types that f a l l within two extreme kinds, forming a gradient, two of these do not belong, because of other d i s t i n c t i o n s (M. pennanti and M. f l a v i g u l a ) .  I t follows that the neotenic or hypermorphotic  condition has arisen several times i n quite d i s t i n c t i v e groups;  87  thus we may make a c l a s s i f i c a t i o n of species, something l i k e this:  S k u l l broad and short Subgenus Martes:  S k u l l long and narrow  melampus, foina, caurina, martes, z i b e l l i n a , americana  Subgenus Pekania:  pennanti  Subgenus F l a v i g u l a :  flavigula  We come now to the problem:  i s the sequence shown i n  s k u l l shape within the subgenus Martes a r e a l one? t h i s one must consider d e n t i t i o n .  To attempt  Figure 8 depicts pie d i a -  grams of r a t i o s , given i n percentage, made from measurements taken from teeth of the various species.  These are e s s e n t i a l l y  the r a t i o s devised by Brongersma (1941) and referred to previously.  These diagrams show basic s i m i l a r i t y i n the combinations  of the s i x dental r a t i o s depicted f o r the following species: M. melampus, M. a. caurina, M. martes and M. z i b e l l i n a .  Of  these M. a. caurina i s apparently the most d i s t i n c t i v e .  M. a.  americana and M. f o i n a are t o t a l l y d i s t i n c t i v e , and strangely, are to a surprising degree s i m i l a r i n dental conformation though d i s t i n c t i v e i n s k u l l shape.  I assume that these relationships  are r e a l and i n view of the lack of other evidence, that within the subgenus Martes, M. melampus, M. a. caurina, M. martes and M. z i b e l l i n a are c l o s e l y related species.  M. f o i n a and M. a.  americana are d i s t i n c t i v e i n dental c h a r a c t e r i s t i c s and within  Figure 8 face page 88)  Figure 8.  Pie diagrams depicting c e r t a i n c h a r a c t e r i s t i c s of the skulls of the world's marten. The indices used are, with the exception of one, those devised by Brongersma (1941). The values graphed are not absolute, but rather percentages, the smallest mean equalling 0 i n the pie cut, the largest 100. The pie cuts are as follows: 1.  Width of inner lobe of P x 100 / width of trenchant part of P  2.  Width of inner lobe of P length of P  3.  Breadth of M  4.  Lateral length of M length of M  5.  Mesial length of M  6.  Breadth of P  7.  Canine width at base x 100 / b u l l a length  4  4  4  4  1  4  x 100 /  x 100 / length of  1  1  P  4  x 100/mesial x 100 / breadth of  / length of  P  M  1  4  88 themselves are similar i n t h i s respect, but are so d i f f e r e n t c r a n i a l l y and i n t h e i r biology as to be l i k e l y d i s t i n c t i v e within themselves.  On zoogeographic grounds, and pelage  structure and colour, M. a. americana shows a degree of r e l a tionship with M. z i b e l l i n a and M. martes, but for the time being i t i s here considered otherwise.  Thus a scheme of re-  lationship within the subgenus might be drawn up thus: S k u l l broad, short P  4  Skull long, narrow  small, with  small inner lobe;  M  1  with  foina  americana  small inner moiety 4  P  large, with  large inner lobe; M^" with  melampus  caurina martes  zibellina  large inner moietsy k h i e r a r c h i c a l c l a s s i f i c a t i o n designed to show these relationships may be prepared as follows.  I t must be remembered  that t h i s does not attempt to represent phylogeny, though to a degree i t might.  For the time being i t attempts rather to  depict morphological s i m i l a r i t y of crania.  89  Subgenus Martes:  1.  M. melampus, americana caurina, martes and z i b e l l i n a  2.  M. f o i n a  3.  M. americana  Subgenus Pekania: 1.  M. pennanti  Subgenus Charronia: 1.  M. f l a v i g u l a  The baculum of M. pennanti i s so similar to that of the species of the subgenus Martes, that i t s relationship to that subgenus must be considered much closer than i s the r e l a tionship between the subgenera Martes and Charronia. There i s one problem more to be considered.  The e v i -  dence appears such that M. a. americana and M. a. caurina are as morphologically d i s t i n c t between themselves as either i s compared to any of the Old World species.  In f a c t , M. a. caurina  appears much more c l o s e l y related to M. melampus, M. martes or M. z i b e l l i n a than M. a. americana does to M. a. caurina.  And  yet, as Wright has shown, M. a. americana and M. a. caurina interbreed, and must be considered to represent the same b i o l o g i cal species.  I f t h i s i s so, one wonders how i t i s that the  Eurasian species, so much more s i m i l a r morphologically, have maintained t h e i r s p e c i f i c d i s t i n c t i o n i n the eyes of systematists, It has occurred to me that M. martes and M. z i b e l l i n a may possibly interbreed where t h e i r ranges meet i n north central Russia and that M. z i b e l l i n a and M. melampus interbreed where t h e i r ranges meet i n Korea.  It i s s i g n i f i c a n t that the ranges  of these species are nowhere sympatric.  One wonders, too, i f  90  M. a. caurina and hence M. a. americana might not prove to be conspecific i n the same sense with M. z i b e l l i n a .  I f t h i s were  shown to be so, i t would reduce the number of species of marten i n the world from eight as here interpreted to a maximum of four as follows: M. martes (including M. z i b e l l i n a , M. melampus and M. americana) M. foina M. f l a v i g u l a (including M. guatkinsi) M. pennanti.  D.  THE MARTENS OF THE NEW WORLD LIFE HISTORY B r i e f accounts of the l i f e h i s t o r i e s of the European  forms have been given i n the pages preceding.  Schmidt (194-3)  assures us that the natural history, reproductory and food habits of Martes foina, martes and z i b e l l i n a are i n every respect almost i d e n t i c a l to those of M. americana and M. pennanti.  Thus i t  would appear that what i s b r i e f l y stated below may be attributed also to the Eurasian forms named above. Martes americana:  B i r t h occurs usually i n A p r i l , less often i n  May and June ( l a Beree, Brassard and Bernard 1939» Schmidt 1943, Lensink 1953, Pearson and Enders 1944 a and b ) .  According to  Pearson and Enders (1944a) t h i s period can be shortened by three or four months by means of a r t i f i c i a l l i g h t i n g , and since the long gestation period has been shown by the above authors to be caused by delayed implantation of the blastocyst, they concluded  91 that implantation i s controlled by day length i n spring.  Oddly  enough, although the long gestation period has only recently been determined, Kerr i n 1792 wrote that the period was said to be nine months.  The l i t t e r size i s one to s i x , averaging 2.6  to 3*0 (Brassard and Bernard 1939, Markley and Bassett 1942, Marshall 1951, Schmidt 1943, Lensink 1953).  The animals are  born toothless, and the permanent adult d e n t i t i o n i s attained by the 16th to 18th week (Brassard and Bernard 1939, Schmidt 1943). Both above authors give detailed descriptions of the sequence by which adult dentition i s attained.  The young leave the nest at  the e a r l i e s t by the 44th day, but usually by the eighth week, t h i s being most often sometime i n mid-July (Brassard and Bernard 1939,  Schmidt 1943).  The f u l l adult weight i s approximated by  the end of the t h i r d month (Rand 1948a, GrinnelL, Dixon and Linsdale 1937, Bailey 1936, Brassard and Bernard 1939) though the animal continues to f i l l out u n t i l i t s t h i r d year ( l a Beree 1941). The animals f i r s t mate i n t h e i r second year, giving b i r t h to t h e i r f i r s t l i t t e r i n t h e i r t h i r d year ( l a Beree 1941, Markley and Bassett 1942, Marshall 1951,  Schmidt 1943).  According to  Lensink they occasionally mate f o r the f i r s t time at f i f t e e n months, usually at twenty-seven months of age. considered polygamous (Marshall 1951,  They are usually  Schmidt 1943).  One l i t t e r  i s produced every year thereafter u n t i l death, the oldest known animal being seventeen years old when i t died ( l a Beree 1941, Seton 1925-1928, Lensink 1953).  Growth curves f o r the early  stages of marten l i f e are given by Markley and Bassett 1942, Remington 1952, and Schmidt 1943.  92 As remarked on e a r l i e r , the animals occur i n greatest numbers i n coniferous f o r e s t s , especially those dominated by spruce (Picea).  Densities of the animal per unit area vary from  l o c a l i t y to l o c a l i t y .  The area covered by individuals appears  to vary with sex, season and age.  Marshall (195D believed the  winter range of marten to be about ten to f i f t e e n miles.  Males  appear to hunt over a wider area than females (Yeager 1950). Lensink (1953), reviewing the l i t e r a t u r e of de Vos (1952) and Yeager (1950), who had both reviewed the e a r l i e r l i t e r a t u r e , and including with these his own figures from i n t e r i o r Alaska and figures based on skulls used i n the present study, found that 159 males were trapped f o r every 100 females.  Various authors  have, however, suggested that this difference may only be a result of the wider cruising radius of the male and not a r e a l deviation from a 1 : 1 r a t i o .  Quick (1953) i n northern B r i t i s h  Columbia found the ratio i n 1947 to be 1 : 1, i n 1948 1.2 : 1. Lampio (1951) likewise found a nearly 1 : 1 r a t i o to exist i n M. martes i n Finland.  Lensink (1953) concluded that nearly 50$  of the marten population was made up of immature  (non-breeding)  animals. Lensink (1953) reviews the l i t e r a t u r e on foods eaten by Marten (Cowan and Mackay 1950, Marshall 1946, Newby 1952, Remington 1950 and others) adding his own observations to these. Microtines comprised by f a r the largest proportion of food taken, except i n Washington where s q u i r r e l s appear the most eaten. Other foods include animals to the size of rabbits, insects and berries.  See also Newby and Hawley (1954) not examined during  t h i s study.  93  Martes pennanti: B i r t h occurs from March 15 to the end of A p r i l (Douglas 194-3, l a Beree 1941, Hodgson 1937,  Seton 1925-  1928, Enders and Pearson 1943a and b, Schmidt 1943, H a l l 1942), as i n marten probably varying with l a t i t u d e .  Mating occurs  from three days to eighteen days after b i r t h , probably about a week as a r u l e .  Dates of b i r t h given i n the l i t e r a t u r e vary  from March 15 to A p r i l 27 (Douglas 1943, l a Beree 1941, Hodgson 1937, Enders and Pearson 1943a and b, Schmidt 1943, H a l l 1942). The gestation period i s stated to vary from 210 to 370 days, averaging about 350 days (Douglas 1943, l a Beree 1941, Hodgson 1937, Pearson and Enders 1944a and b, H a l l 1942, Schmidt de Vos 1952).  1943,  According to Enders and Pearson 1943a and b,  the period of delayed b l a s t o c y s t i c implantation i s nine or more months.  The l i t t e r size i s said to be one to f i v e , averaging  two or three (Douglas 1943, Hodgson 1937, 1952, Seton 1925-1928.  Schmidt 1943, de Vos  The features coincident to the a t t a i n -  ing of adult dentition have yet to be worked out, but i n view of the s i m i l a r i t y of the other features of the l i f e h i s t o r y to that of the martens i t i s probably quite s i m i l a r .  The young leave  the nest at nine to ten weeks (Schmidt 1943) and are f u l l sized by seven months (Hodgson 1937)•  Mating f i r s t occurs during the  second year, the f i r s t l i t t e r being born i n the t h i r d (Douglas 1943, Hodgson 1937, H a l l 1942).  The animals are said to be  polygamous (Douglas 1943). Fisher tend to occur i n lower altitudes than marten (Edwards 1950, de Vos 1952).  De Vos considers i n rather great  d e t a i l the habitat requirements of the animal.  The animals  94 appear to vary i n density geographically, staying generally within an area of two to s i x square miles (de Vos 1952, Quick 1953).  Unlike marten, there appear to be more females than  males per unit area, trapping figures giving 49 males to 51 f e males (de Vos 1952) and one male to two females (Quick 1953). Animals captured by f i s h e r as food are l a r g e l y grouse, red s q u i r r e l s , hares, shrews, porcupines and others (de Vos 1952, Quick 1953).  VARIATION Non-geographic variations There are several kinds of non-geographic v a r i a t i o n , namely that which occurs by sex, by age, by season and that which i s c h a r a c t e r i s t i c of individuals of the same sex and age, here termed i n d i v i d u a l v a r i a t i o n , whether genetic or otherwise. Since any attempt to describe geographic v a r i a t i o n must be preceded by some understanding of the noh-geographic sort, so that the two may be distinguished a b r i e f account follows.  Burt  (1953) records the taking of about 400 marten from Fort Nelson B r i t i s h Columbia, which should be of use to anyone studying nongeographic v a r i a t i o n . Sexual v a r i a t i o n :  We may consider here only the more obvious  of the secondary sexual c h a r a c t e r i s t i c s . these i s s i z e .  The most important of  The males i n a l l species of martens and fishers  are considerably larger than the females.  For martens i n terms  of weight, i t may be stated as a general rule that the female weighs approximately two-thirds that of the male.  The pelage  95  of the male i s thicker, with longer hairs and a greater elas1952,  t i c i t y to the fur (see Brassard and Bernard 1939? de Vos Markley and Bassett 1942, Lensink 1953? Lampio 1951).  The  under f u r anterior to the urethral aperture on the midline of the b e l l y l i e s forward, rather than backward i n the male (Lampio 1951,  Lensink 1953).  The skulls of the males are p l a i n l y heavier and more robust than those of the females.  The canines and the dentition  i n general i s heavier and the s a g i t t a l crest more l i k e l y to be highly developed i n those forms where i t occurs.  For Martes  americana americana and M. a. caurina the measurements of large samples of six s k u l l c h a r a c t e r i s t i c s a r b i t r a r i l y chosen were compared by sex and i t was found that a l l of these averaged  1.10  2  times l a r g e r i n the male than i n the female, and the X  probabil-  i t y that these averages came from the same supply was f a r beyond the 99$ l e v e l .  The characters employed were condylobasal  length, r o s t r a l width, upper tooth row, b u l l a length, r o s t r a l width / b u l l a length and upper molar widthplus length.  The use  of the above index would allow one to predict the s k u l l s i z e and c h a r a c t e r i s t i c s of both sexes of any unknown form of North American marten, i f only one sex i s known.  No comparable  were made f o r Eurasian martens or M. pennanti. the figure i s about 1.6 compared to 1.1  comparisons H a l l (1934) says  i n marten.  Most of the specimens of both marten and f i s h e r used i n t h i s study were already designated to sex.  Where the sex of an  animal was not known, i t was i d e n t i f i e d by comparison to bar graphs of condylobasal length and other measurements drawn up  96  from larger samples of known sex, from the same general area. Age V a r i a t i o n :  As marten advance from maturity (second  year)  into old age, the following changes have been noted to occur: the condylobasal length increases (Lensink 1953» Wright 1953), the zygomatic breadth increases (Grinnell, Dixon and Linsdale 1937? Wright 1953)? the braincase i s said to decrease i n size ( G r i n n e l l , et a l . , 1937)»  the p o s t o r b i t a l c o n s t r i c t i o n narrows  (Grinnell et a l . , 1937"? Wright 1953), the rostrum narrows and lengthens  (Grinnell et a l . , 1937)>  the naso-maxillary sutures  fuse (other sutures ankylose very early) (Grinnell et a l . , 1937> Rhoads 1902, Lensink 1953? Marshall 1 9 5 D > the p o s t o r b i t a l process becomes more prominent (Grinnell et a l . , Lensink, Marshall), the baculum increases i n weight (Marshall). During the early stages of l i f e ,  age may be obtained by  use of growth curves (Markley and Bassett 194-2, Remington 1952, Schmidt 194-3) or to the eighteenth week by the sequence of tooth eruption (Brassard and Bernard 1939, Schmidt 1943). with "M. c. caurina" of central Idaho, Marshall (1951)  Working concluded  that the presence of a s a g i t t a l crest i n a female or one longer i  than 2 mm. i n a male implied sexual maturity (two years or older). Likewise Marshall concluded that a baculum heavier than 100 mg. implied sexual maturity. Lensink (1953), working on the marten of i n t e r i o r Alaska, concluded that he was able to t e l l the age of males to four years, of  females to f i v e years by means of weight of baculum and  height and length of s a g i t t a l c r e s t . i s reproduced below.  His table of measurements  97  Males Age  Baculum weight (mgs.)  0 - 1  Less than 130  1-2  130 - 205  2 - 3  206 - 260  Over 4 Over 5  Height of s a g i t t a l crest (mm.)  Length of s a g i t t a l Crest (mm.)  0  0  Less than 16  1-20  16 - 19  25 - 40  More than 260  More than 18  41 - 50  More than 260  More than 18  More than 45  Length of sagitt a l crest (mm.)  Height of s a g i t t a l crest (mm.)  Females Age  Min. separation of temporal muscle scars ( mm.)  0 - 1  0  0  More than 5  1 - 2  0  0  Less than 5  2 - 3  1-20  0  0  3-4  25-37  0  0  Over 5  More than 38  0  0  Lensink's work comes closest to providing a means to age marten skulls, and i n systematic work, to break samples into age subsamples.  However, one result of the present study has  been to show that marten from different l o c a l i t i e s possess bacula of s i g n i f i c a n t l y different weights at maturity and acquire sagitt a l crests of different sizes at d i f f e r e n t  ages.  At the beginning of t h i s study, twenty-four characteri s t i c s were selected  from a population of marten of r e s t r i c t e d  geographic extent (Banff region, Alberta).  These were broken  into two age groups by use of the following single  characteristic:  98  fusion or lack of fusion of the naso-maxillary suture.  (fide Rhoads 1902) were consid-  fusion was complete, the animals ered adult;  Where  where unfused, immature.  Wright (1953) has done  the same and i n common with him I found that the following characters varied s i g n i f i c a n t l y between the two age groups: and condylobasal length of s k u l l ;  greatest  zygomatic breadth, p o s t o r b i t a l  c o n s t r i c t i o n , postorbital width, vomerine width, width of i n c i s o r row at base, r o s t r a l width and length and lower jaw length. Mastoid breadth, i n t e r o r b i t a l breadth, height of cranium, size of auditory bullae, and size of teeth did not vary (Wright  found  no v a r i a t i o n also i n r o s t r a l width, concerning which see above, b a s i l a r length, p a l a t a l length). Of the twelve measurements used i n the present  study,  those found to change with age were measured only i n adults, the remainder on both adult and immatures, as follows: 1.  Condylobasal length.  2.  Rostral width.  3.  Upper tooth row. Adults and immatures.  4.  Canine width at base.  5.  B u l l a length.  6.  Upper molar width.  7.  Upper molar inner moiety length.  8.  P a l a t a l length.  9.  Mastoid width,  Adults only.  Adults only.  Adults only.  Adults and immatures. Adults and immatures. Adults and immatures.  Adults and immatures. adults and immatures.  10.  Height of s k u l l at b u l l a e .  11.  Length of l a s t upper premolar.  12.  P a l a t a l width.  Adults only.  Adults only.  Adults and immatures.  99 Details of the above measurements and those given below for f i s h e r are given i n Appendix C.  In general, where large  samples were available, only the adult portion (by the nasomaxillary test) were measured. In f i s h e r , the following c h a r a c t e r i s t i c s are known to change as old age i s attained: (Rhoads 1903);  lightening i n pelage colour  an increase i n condylobasal length (de Vos  1952);  increase i n the weight of the s k u l l , the male about 25$? the female about 9% (Grinnell et a l . ) ; an outward bowing of the zygomatic arches (Coues 1877? Thomas 1886, G r i n n e l l et a l . , de Vos);  the p o s t o r b i t a l c o n s t r i c t i o n narrows (Coues, Thomas,  G r i n n e l l et a l . ) ; the lambdoidal (Coues);  crest increases i n size  the sutures of the nasal region and basi-cranium  ose (Coues, Grinnel e_t a l . , de Vos); (Thomas);  ankyl-  the canines become longer  the i n t e r o r b i t a l width increases (de Vos);  the sag-  i t t a l crest increases i n height and length (Coues, Thomas, Grinnel et a l . ) ; the mandible increases with depth (Hall 194-2); and the baculum increases i n weight (de Vos).  According  Vos a male with no s a g i t t a l crest i s j u v e n i l e , with one than 62 mm.  to de less  long i s adult, and with one longer i s old adult.  Of females, only old adults bear a crest. As i n the case of marten, a large number of characteri s t i c s were measured f o r a sample limited geographically (Stuart Lake, B r i t i s h Columbia), these being broken into ^subadult"  and  "adult" on the basis of degree of fusion of the naso-maxillaries. No s i g n i f i c a n t v a r i a t i o n was  found i n the upper tooth  row,  100 mastoid width, p a l a t a l length, p r e o r b i t a l and p o s t o r b i t a l widths, and size of teeth.  These measurements were:henceforth  taken on a l l "adult" and "immature" animals.  The  remaining  characters were found to vary with age, and were measured only on "adults".  These included condylobasal length, canine width  at base, r o s t r a l width, b u l l a length, and palate length. As i n marten, where large samples were available, only "adults" were measured. Rhoads i n 1898 expressed  the b e l i e f that the white  patches on the breasts of f i s h e r s were l o s t with age.  In  1903  he suggested further, that the oldest specimens are the l i g h t e s t coloured, some becoming nearly white. Seasonal v a r i a t i o n :  This kind of v a r i a t i o n i s concerned c h i e f l y  with v a r i a t i o n i n the condition of f u r .  Surprisingly l i t t l e  i s known concerning the moulting of marten and f i s h e r . Grinnell, Dixon and Linsdale (1937) reported one complete annual moult i n the marten of the S i e r r a Nevada. slowly through the autumn months. the end of August.  This proceeded  It begins i n the t a i l , about  They also report a loss of hair with no  renewal i n the spring months, f o r summer skins are l e s s dense i n pelage than are the winter ones.  Summer and winter skins, they  reported, are c l o s e l y alike i n colouration* the former being only s l i g h t l y the paler of the  two.  Markley and Bassett 1942  reported that during the summer  moult the colour changed from "dark brown to l i g h t tan".. moult began on the t a i l t i p , then moved to the legs, the dorsal region and the face, i n that order.  The mid-  101 According to Schmidt annual moults.  (194-3) European marten undergo two  The hair begins to f a l l at the end of A p r i l or  beginning of May and l a s t s u n t i l the beginning of June. ing  begins on the face and works back to the t a i l .  coat i s darker than that of the winter.  Moult-  The summer  Pregnant females end  t h e i r summer moult much e a r l i e r than non-breeding  ones.  The f a l l moult begins i n August, and ends by November. Here the order of hair loss i s reversed;  as i n Grinnell*s  animals, the f i r s t part affected being the t a i l , the l a s t the neck and head. G r i n n e l l et a l . concluded that C a l i f o r n i a f i s h e r show only one annual moult, t h i s occurring slowly, i n the f a l l .  With  the advance of summer a s l i g h t fading of tones appears. It  seems l i k e l y then, that f i s h e r and marten undergo a  f a l l moult, and a spring loss of h a i r , at l e a s t .  My own detailed  examination of skins has been l a r g e l y r e s t r i c t e d to specimens taken from the Canadian Rockies.  These show a summer pelage f a r  darker and thinner than the winter one.  The f a l l moult  appears  to occur considerably l a t e r than i t does i n the C a l i f o r n i a n specimens examined, time of moulting l i k e l y being, as Bissonnett (1944) shows i n weasels, a function of day length. In  general, where I have attempted comparison of skins,  I have divided them into three groups, summer, winter (by date of capture) and moulting (by appearance).  Comparison has then been  made within each of the classes "summer" and "winter". Individual v a r i a t i o n : for  Individual v a r i a t i o n has been recorded  a long time i n martens and i s known to be extreme, e s p e c i a l l y  102  with regard to colour of pelage.  A l l writers, however, have  f a i l e d to d i s t i n g u i s h between v a r i a t i o n that might be due to sex, age or season, which makes the s i t u a t i o n a d i f f i c u l t  one  to i n t e r p r e t . In 1 8 7 5 Cartwright wrote that the marten "varies the most i n colour, i n the same region, of any animal;  some are  l i g h t yellow, some are dark brown, and others are black".  Mac-  farlane ( 1 9 0 5 ) wrote that he had "seen several albino examples and also a considerable number of bright yellow and dark orange coloured martens  p a r t i c u l a r l y while stationed i n the d i s -  t r i c t s of Mackenzie River and Athabasca".  Hardy i n 1 9 1 0 record-  ed the taking of a very dark specimen from Maine.  It was  "the  darkest furred one I ever saw south of the St. Lawrence" he reported, adding that he had examined over 2 0 , 0 0 0 pelts from the region, but none so dark as t h i s .  Cory ( 1 9 1 2 ) remarks that i n  eastern marten the brown colour i s l i g h t e r or darker i n different i n d i v i d u a l s , the throat patch varying from orange to yellowish white. Markley and Bassett 194-2 reporting on marten of Idaho :  and Wyoming, say that "evident at once i s a great i n d i v i d u a l v a r i a t i o n i n extent of the throat patch. i r r e g u l a r , i n several specimens was  Its shape, always  r e s t r i c t e d to a number of  detached spots on the lower throat and breast, while on others i t became a s o l i d patch extending from the upper throat to the abdominal region.  On a prime winter pelt the t a i l and legs  became nearly black on some i n d i v i d u a l s . "  103  A detailed account i s given by de Vos (1952).  The  colour of the f u r , he concludes, i s "extremely v a r i a b l e " .  He  presents a series of skins, selected from a c o l l e c t i o n of 450 pelts trapped from November 1st to March 31st of two consecutive years.  These varied from "a very pale yellow-orange marten of  a p a r t l y cromistic, p a r t l y a l b i n i s t i c type, "with an e n t i r e l y white head" and with white underparts, feet and t a i l the same colour of the upper parts and with no throat patch to a "very dark marten with a large number of white guard hairs  the  underparts ... of a similar colour ... with a large orange throat patch."  He also records a dark, possibly melanistic  specimen, but no albinos, though Ponomarev (1938) found them i n Russian sables.  Ponomarev reported also a tendency toward  geographic colour v a r i a b i l i t y and de Vos likewise found the marten of Algonquin Park, Ontario, to be, on the average, s l i g h t l y paler.  De Vos concluded that "the v a r i a b i l i t y i n a l o c a l  series indicates that one should be on his guard when making comparisons between series from d i f f e r e n t l o c a l i t i e s , and that one preferably should compare large series."  Wright i n 1953»  working with 300 specimens designated M. a. abietinoides from Montana and Idaho, likewise found great v a r i a b i l i t y ;  dorsally  they varied from I s a b e l l a color or Mouse Grey to Mummy Brown, and v e n t r a l l y from I s a b e l l a or Mouse Grey to nearly black.  The  throat patch varied from almost complete absence to f i v e or s i x inches square and from Pale Orange-Yellow to Capucine Yellow or Deep Chrome (colours from Ridgway, 1912).  104 Size v a r i a t i o n , within a limited area, appears to be rather less than one might expect on s u p e r f i c i a l examination of skins and s k u l l s .  A sample of 100 skins taken from Ontario, when  divided into male and female, but not into age groups, give the following average c o e f f i c i e n t s of v a r i a b i l i t y : 4.2$;  t a i l length, 6.1$;  hind foot 6.0$;  t o t a l length  and weight 13.2$.  Another sample of 21 "adult" males from near Banff, Alberta, when measured f o r twenty-four s k u l l c h a r a c t e r i s t i c s , gave an average c o e f f i c i e n t of v a r i a b i l i t y of 3 . 3 $ , with a maximum much higher i n only one c h a r a c t e r i s t i c (7*5$ f o r p o s t o r b i t a l width). I believe that f i s h e r s show rather l e s s v a r i a b i l i t y than do martens.  De Vos (1952) examined 200 skins from Ontario  taken between l a t e November and l a t e March.  He remarked  "Colour v a r i a t i o n s between individuals are f a i r l y s t r i k i n g . such v a r i a t i o n s could be detected between l o c a l  No  populations.  The o v e r a l l colour of the dorsum varies from pale grey to p r a c t i c a l l y black ....  This i s true f o r both sexes, although the  gray colour predominates i n males. be noticed occasionally....  A reddish brown tinge can  On the v e n t r a l side there i s not  much v a r i a t i o n between i n d i v i d u a l s , although l i g h t backed specimens are s l i g h t l y paler than dark ones.  No large throat patches  ... were observed, although a few i n d i v i d u a l s had one or a few white spots on the throat.  The largest white patch i s usually  found on the b e l l y , anterior to the anus.  In some animals  white spots are present i n the a x i l l a s of the front legs which may occasionally be fused on the mid v e n t r a l l i n e and extend a n t e r i o r l y f o r a short distance."  105  Audubon and Bachman (1851-1854) say they had examined a specimen that was  "nearly white with a brown head".  Elliot  (1901a) says they vary from glossy black to gray or grayish white on the upper parts.  G r i n n e l l , Dixon and Linsdale (1937)  working with C a l i f o r n i a n animals, however, concluded colour v a r i a t i o n s were s l i g h t , being expressed of white on the  that  c h i e f l y i n amount  underparts.  Size v a r i a t i o n i s about the same as i n marten. 1882-188