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Systematics of Arnica, subgenus Austromontana and a new subgenus, Calarnica (Asteraceae : Senecioneae) Straley, Gerald Bane 1980

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SYSTEMATICA OF ARNICA, SUBGENUS AUSTROMONTANA AND A NEW  SUBGENUS, CALARNICA  (ASTERACEAE:SENECIONEAE)  by GERALD BANE STRALEY B.Sc,  V i r g i n i a Polytechnic  I n s t i t u t e , 1968  M . S c , Ohio U n i v e r s i t y , 1974 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS OF THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES (Department o f Botany)  We accept t h i s t h e s i s as conforming to the r e q u i r e d  standard  THE UNIVERSITY OF BRITISH COLUMBIA  March 1980  ©  G e r a l d Bane S t r a l e y , 1980  In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make i t freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives.  I t is understood that copying or publication  of this thesis for financial gain shall not be allowed without my written permission.  Department nf  Botany  The University of British Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 26 March 1980  ABSTRACT  Seven s p e c i e s a r e r e c o g n i z e d species  i n a new  subgenus C a l a r n i c a based on a c r i t i c a l  t i v e r e v i s i o n of the s p e c i e s . representing  i n A r n i c a subgenus Austromontana and  Chromosome numbers a r e g i v e n  h y b r i d i z a t i o n s are given.  l e a f m i n e r s , achene f e e d e r s , cordifolia,  breeding  and  pentaploid,  complex of d i s c o i d p o p u l a t i o n s  floret  and  feeders are presented.  hexaploid  ly r e s t r i c t e d  to s e r p e n t i n e  morphic s p e c i e s , has l a r g e l y s e x u a l and serpentine  s o i l s and  autonomously a p o m i c t i c ,  s e x u a l and  sexual, d i p l o i d  Arnica l a t i f o l i a ,  endemic, and species.  and  species,  A r n i c a g r a c i l i s now races.  The  polyremained cernua,  probably exists largely  subgenus C a l a r n i c a  of A r n i c a  t h r o u g h A r n i c a venosa, the l a t t e r g i v i n g r i s e to A r n i c a v i s c o s a .  ii  pseudo-  to A r n i c a  to A r n i c a g r a c i l i s ,  triploid  diploids.  to a  pseudogamous  the most  given r i s e  from subgenus Austromontana by way  species are r a r e sexual  rise  from A r n i c a d i s c o i d e a , i s l a r g e -  A r n i c a l a t i f o l i a has  as a s e r i e s of autonomously a p o m i c t i c  two  given  a r i s e n from A r n i c a c o r d i f o l i a , a l t h o u g h i t has  diploid.  possibly derived  I t has  i s a s e x u a l d i p l o i d , w i t h a few  populations.  through h y b r i d i z a t i o n with other  was  populations,  to a l e s s e r degree  populations.  tetraploid,  Arnica spathulata, derived  gamous t e t r a p l o i d a p o m i c t i c  the  Arnica  t r e a t e d here as a s i n g l e p o l y m o r p h i c  A r n i c a d i s c o i d e a , w i t h b o t h d i p l o i d and races.  artifi-  I n t e r r e l a t i o n s h i p s of i n s e c t p o l l i n a t o r s ,  to A r n i c a n e v a d e n s i s which i s t e t r a p l o i d and  apomictic  nevadensis.  the a n c e s t r a l s p e c i e s c o n s i s t s l a r g e l y of t e t r a p l o i d  triploid,  populations  s t u d i e s , and  which a r e e i t h e r autonomous or pseudogamous a p o m i c t s , and diploid,  conserva-  f o r 91  a l l s p e c i e s , i n c l u d i n g the f i r s t r e p o r t s f o r A r n i c a  R e s u l t s of a p o m i x i s , v e g e t a t i v e r e p r o d u c t i o n , cial  r e v i e w and  two  spathulata These l a t t e r  iii  TABLE OF  CONTENTS  page Abstract  •  i i  Frontispiece  i i i  L i s t of T a b l e s  vi  L i s t of F i g u r e s  vii  Acknowledgement  x  Introduction  1  .  H i s t o r i c a l Background  2  Arnica - Tribal Position  2  Taxonomic H i s t o r y  3  Methods and  M a t e r i a l s - General  6  D i s c u s s i o n of C h a r a c t e r s Habit  and  7  Branching P a t t e r n  7  Underground P a r t s - Roots, Rhizomes, Caudices  8  Leaves  15  Odors of Leaves  16  Vestiture  21  Capitula  32  Phyllaries  32  F l o r e t s - Ray,  D i s c , and  Ampliate  35  Pappus  44  Achenes  45  D i s t r i b u t i o n and  Ecology  Reproductive Biology Phenology  •  56  *  56 56  iv  page Cytology  58  Apomlxis  60  Pollen Stainability  89  Seed P r o d u c t i o n  95  Seed Germination  96  Seedlings  98  V e g e t a t i v e Reproduction  101  C u l t i v a t i o n of S p e c i e s  I l l  Hybridization Sympatric  112 Occurrence and N a t u r a l H y b r i d i z a t i o n  A r t i f i c i a l Hybridization Entomological  "...  112 115  Interrelationships  120  Leaf Miners  121  F l o r e t Feeders  121  Achene Feeders  126  UV P a t t e r n s  127  Pollinators  130  Systematic  Treatment  A r t i f i c i a l Key  135  to Subgenera  135  S p e c i e s Recognized A r t i f i c i a l Key F i e l d Key  136  to Austromontana and  to Austromontana and  C a l a r n i c a Species  C a l a r n i c a Species  136 138  Taxonomic Treatment  140  Phylogenetic  250  Relationships  Summary  258  Literature Cited  261  Appendices  265 v  LIST OF TABLES pages I. II. III. IV. V. VI. VII. VIII. IX. X.  Comparison o f mature achenes.  55  P r e v i o u s l y r e p o r t e d chromosome numbers  63,64  New Chromosome numbers.  65-70  C o m p a t i b i l i t y and A p o m i c t i c Experiments  86-88  Rhizome Growth.  110  Sympatric Occurrence o f S p e c i e s  114  A r t i f i c i a l Hybridizations  116,117  Leaf M i n e r s .  122  Floret  125  Feeders  Pollinators.  132-134  vi  LIST OF  FIGURES  pages 1-9.  Schematic diagrams of h a b i t and average b r a n c h i n g .  10-25. 26-126.  9-14  Average l e a v e s .  17-20  Comparison of v e s t i t u r e t y p e s on f o u r v e g e t a t i v e p a r t s .  23-31  127-136.  Average i n v o l u c r a l b r a c t s .  33,34  137-142.  T y p i c a l ray f l o r e t s .  36,37  143-152.  Typical disc florets.  38-41  153-156.  Ampliate f l o r e t s  42,43  157-165.  Pappus.  45-48  166-186.  Achene s u r f a c e s .  49-54  187-194.  M e i o t i c chromosomes, m i c r o s p o r o c y t e s ,  195-207.  Chromosomes.  71-82  208-211.  E x p e r i m e n t a l methods.  84,85  212-221.  Pollen stainability.  91-94  222-227.  Seedlings.  99,100  228-231.  Rhizome growth.  232.  of Arnica discoidea  and p o l l e n g r a i n s .  61,62  102-109  Summary o f a r t i f i c i a l c r o s s e s .  118  233-240.  Leaf Mines o f Phytomyza s p e c i e s i n l e a v e s o f A r n i c a .  241-247.  UV p a t t e r n s  i n A r n i c a heads and t y p i c a l  123,124  heads of 3 species.128,129  248.  D i s t r i b u t i o n map o f A r n i c a c o r d i f o l i a .  145,146  249.  Habit  147,148  250-259.  of A r n i c a c o r d i f o l i a .  V a r i a t i o n i n leaves of Arnica c o r d i f o l i a ,  260-267.  V a r i a t i o n i n leaves of A r n i c a c o r d i f o l i a ,  268-276.  V a r i a t i o n i n leaves of Arnica c o r d i f o l i a ,  large-leaved form.  149-151  a l p i n e form.  153,154  disturbed roadside  vii  form.  155,156  L i s t of F i g u r e s , c o n t ' d .  277-281.  Habitats  pages  and h a b i t o f A r n i c a c o r d i f o l i a  and A r n i c a nevadensis.  of p u t a t i v e Arnica c o r d i f o l i a  X Arnica l a t i f o l i a  158,159  282.  Habit  h y b r i d . 161,162  283.  D i s t r i b u t i o n map o f A r n i c a n e v a d e n s i s .  165,166  284.  Habit  167,168  285.  D i s t r i b u t i o n map o f A r n i c a l a t i f o l i a .  174,175  286.  Habit  176,177  of A r n i c a nevadensis.  of A r n i c a l a t i f o l i a .  287-298.  V a r i a t i o n i n l e a v e s of A r n i c a l a t i f o l i a ,  s m a l l form.  299-304.  V a r i a t i o n i n shortened r a y f l o r e t s i n A r n i c a l a t i f o l i a .  179,180 181,182  305.  D i s t r i b u t i o n map o f A r n i c a g r a c i l i s and A r n i c a d i s c o i d e a .  186,187  306.  Habit  188,189  of A r n i c a g r a c i l i s .  307-318.  V a r i a t i o n i n l e a v e s o f A r n i c a g r a c i l i s , s m a l l form.  192,193  319-332.  V a r i a t i o n i n l e a v e s o f A r n i c a g r a c i l i s , l a r g e r form.  194,195  333.  D i s t r i b u t i o n map o f A r n i c a cernua and A r n i c a s p a t h u l a t a .  198,199  334.  Habit  200,201  o f A r n i c a cernua.  335-338.  Habitats  339-355.  V a r i a t i o n i n l e a v e s o f A r n i c a cernua, s m a l l form.  356-365.  V a r i a t i o n i n l e a v e s o f A r n i c a cernua, l a r g e form, sun form.207,208  366-370.  V a r i a t i o n i n l e a v e s o f A r n i c a cernua, l a r g e form, shade form.  371.  Habit  372-375.  376-385. 386.  and h a b i t o f A r n i c a cernua.  203,204  of A r n i c a d i s c o i d e a .  205,206  209,210 214,215  H a b i t a t s and h a b i t o f A r n i c a d i s c o i d e a and A r n i c a spathulata  217,218  V a r i a t i o n i n leaves of A r n i c a d i s c o i d e a .  219,220  Habit of Arnica spathulata.  224,225  387-398.  V a r i a t i o n i n l e a v e s of A r n i c a s p a t h u l a t a ,  s m a l l form.  226,227  399-407.  V a r i a t i o n i n l e a v e s o f A r n i c a s p a t h u l a t a , l a r g e form.  228,229  viii  L i s t of F i g u r e s ,  cont'd.  408.  D i s t r i b u t i o n of A r n i c a v i s c o s a .  233,234  409.  Habit of Arnica v i s c o s a .  236,237  410- 414.  H a b i t a t and h a b i t o f A r n i c a v i s c o s a .  238,239  415.  D i s t r i b u t i o n map o f A r n i c a venosa.  242,243  416.  H a b i t o f A r n i c a venosa.  244,245  417- 421.  Habitat  and h a b i t o f A r n i c a venosa.  422.  Phylogeny of A r n i c a  423.  Phylogeny o f A r n i c a s p e c i e s i n subgenera  246,247  Subgenera.  254,255 Austromontana and C a l a r n i c a .  ix  256,257  ACKNOWLEDGEMENTS  I would l i k e to thank my  a d v i s o r y committee, Bruce A. Bohm, Fred  Ganders, C h r i s t o p h e r J . Marchant, J u d i t h A. Myers, and my Roy  L. T a y l o r , f o r t h e i r h e l p f u l comments and  For c o l l e c t i o n s of seeds and  major a d v i s o r ,  c r i t i c i s m s during t h i s  study.  p l a n t s I thank Bruce A. Bohm, R i c h a r d A. G o r n a l l ,  G e r a l d i n e A. Guppy, Mary F. Jackson, Marchant, K e v i n W.  R.  Timothy A. Johns, C h r i s t o p h e r J .  N i c h o l l s , and A l l a n A. Rose.  For a s s i s t a n c e i n l o c a t i n g  r a r e r s p e c i e s I thank Susan Horner and numerous o t h e r members of the U. N a t i o n a l F o r e s t S e r v i c e , Michigan  N a t u r a l Resource Department, and U. S.  Canadian N a t i o n a l Park S e r v i c e s f o r t h e i r c o o p e r a t i o n i n a l l o w i n g me collect  i n t h e i r areas.  Claude E. Garton, was  Veto and  micrographs and  Sylvia Taylor.  F. C. Thompson, Robbin W. t i o n was  other photographic  L. T a y l o r .  techniques  I n s e c t s were determined by Graham C. D. Thorp, R i c h a r d L. Westcott,  done by P a t r i c i a M.  f o r her many hours of work. thank Roy  Roy  Eckel.  I thank my  I  Griffiths,  the L a t i n d e s c r i p -  c a r e f u l t y p i s t , B e t t y Reubart,  encouragement and  financial  to  support  E n g i n e e r i n g Research C o u n c i l of Canada o p e r a t i n g  g r a n t number 67-5705. I would l i k e to thank the c u r a t o r s of the f o l l o w i n g h e r b a r i a f o r a l l o w i n g me  with  by L a s z l o  L a s t l y and most i m p o r t a n t l y , I would l i k e  L. T a y l o r f o r h i s c o n t i n u e d  through N a t u r a l S c i e n c e s and  and  and  to  Many hours of p l e a s u r a b l e c o l l e c t i n g were spent  R. P a t r i c k H a r r i s o n , A l l a n A. Rose and  a s s i s t e d w i t h SEM  S.  to examine specimens of A r n i c a under t h e i r  Brigham Young U n i v e r s i t y  (BRY)  C a l i f o r n i a Academy of S c i e n c e s  (CAS)  Dudley Herbarium, S t a n f o r d U n i v e r s i t y x  (DS)  care:  C e n t r a l Washington  State  University  (ELRG)  Humboldt S t a t e U n i v e r s i t y (HSC) U n i v e r s i t y o f Idaho (ID) Jepson Herbarium, U n i v e r s i t y o f C a l i f o r n i a  (JEPS)  Kathryn Kalmbach Herbarium, Denver B o t a n i c Gardens (KHD) U n i v e r s i t y o f C a l i f o r n i a , Los Angeles (LA) Lakehead U n i v e r s i t y  (LKHD)  Museum o f N o r t h e r n A r i z o n a Missouri  Botanical  (MNA)  Garden  (MO)  U n i v e r s i t y of Montana (MONTU) Northern Arizona U n i v e r s i t y California  Polytechnic  (NAU)  State  U n i v e r s i t y (OBI)  U n i v e r s i t y o f Oregon (ORE) Pomona C o l l e g e  (POM)  P a c i f i c Union C o l l e g e (PUA) Rancho Santa Ana B o t a n i c a l  Garden (RSA)  Santa B a r b a r a County Museum (SBM) Southern Oregon C o l l e g e (SOC) U n i v e r s i t y o f B r i t i s h Columbia (UBC) U n i v e r s i t y of C a l i f o r n i a , B e r k e l e y (UC) U n i v e r s i t y of C a l i f o r n i a , Utah S t a t e  Santa B a r b a r a (UCSB)  U n i v e r s i t y (UTC)  U n i v e r s i t y of V i c t o r i a  (UVIC)  B r i t i s h Columbia P r o v i n c i a l Museum (V) Willamette University  (WILLU)  xi  Washington S t a t e U n i v e r s i t y U n i v e r s i t y of Washington  (WS)  (WTU)  C r a t e r Lake N a t i o n a l Park  xii  1  INTRODUCTION  A r n i c a i s a c i r c u m p o l a r genus o f q u e s t i o n a b l e t r i b a l p o s i t i o n i n the A s t e r a c e a e , o c c u r r i n g from the n o r t h e r n l i m i t s o f v a s c u l a r p l a n t growth at 85° n o r t h l a t i t u d e , w i t h f o u r major  southern r a d i i of d i s t r i b u t i o n ;  south  i n A s i a to Japan; i n Europe t o N o r t h e r n J u g o s l a v i a and N o r t h e r n S p a i n ; i n E a s t e r n N o r t h America a l o n g t h e A t l a n t i c Coast and Piedmont  to N o r t h e r n  F l o r i d a ; and i n Western N o r t h A m e r i c a , where i t r e a c h e s i t s g r e a t e s t sity,  t o N o r t h e r n New  Mexico and S o u t h e r n  diver-  California.  In the e a r l y y e a r s of Western N o r t h American e x p l o r a t i o n a v a s t number of s p e c i e s names were p r o p o s e d , c u l m i n a t i n g i n t h e F l o r a of North America i n which Rydberg groups.  (1927) r e c o g n i z e d  Rydberg  presumably  107 s p e c i e s i n N o r t h America i n 17 s u b g e n e r i c  d i d n o t i n t e n d t h e s e groups t o denote  or s e c t i o n s , but r a t h e r c o n v e n i e n t groups as an a i d i n i d e n t i f y i n g A much more c o n s e r v a t i v e t r e a t m e n t , and t h e o n l y comprehensive monograph, based l a r g e l y on morphology, which he r e c o g n i z e d 32 s p e c i e s N o r t h America. Arctica sonis  Maguire was  was  proposed by Maguire  subgenera species.  worldwide (1943), i n  (Appendix I ) o f . w h i c h 27 a r e from Western  the f i r s t . t o d i v i d e the genus i n t o 5 subgenera,  (7 s p e c i e s ) , Andropurpurea  (3 s p e c i e s ) , Montana (2 s p e c i e s ) , Chamis-  (7 s p e c i e s ) , and Austromontana  (13  species).  S i n c e Maguire's works (1943, 1947), t h e genus has r e c i e v e d l i t t l e t h e r study o t h e r than a few r e p o r t s o f chromosome numbers 1963,1967; S t r o t h e r , 1973;  fur-  (Ornduff , et a l . ,  S t r a l e y , 1979), and an e x c e l l e n t survey of apo-  m i x i s i n the genus by B a r k e r (1966, u n p u b l i s h e d ) .  A r e c e n t monograph of the  N o r t h American s p e c i e s ( E d i g e r and B a r k l e y , 1978)  i s based l a r g e l y on Maguire  (1943) w i t h the r e p r o d u c t i v e i n f o r m a t i o n . o f B a r k e r (1966). Barkley  (1978) r e c o g n i z e 23 s p e c i e s and 26 i n f r a s p e c i f i c  E d i g e r and  t a x a (Appendix I I ) .  There remain few Montana, and  Chamissonis,  w i t h i n the l a t t e r to be  taxonomic problems i n the subgenera Andropurpurea, a l t h o u g h a r e - e v a l u a t i o n of i n f r a s p e c i f i c  subgenus seems w o r t h w h i l e .  i n the subgenus A r c t i c a .  t i o n of these s p e c i e s and  The  greatest d i f f i c u l t i e s  However, owing to the c i r c u m p o l a r  seem  distribu-  t h e i r r e l a t i v e i n a c c e s s i b i l i t y , p a r t i c u l a r l y of  A s i a n t a x a , a comprehensive s y s t e m a t i c be done.  The  taxa  study  subgenus Austromontana was  the  of subgenus A r c t i c a remains to  s e l e c t e d f o r the p r e s e n t  study  be-  cause of unanswered taxonomic problems i n t h i s subgenus f o l l o w i n g Maguire's monograph and 1947;  the number of taxonomic changes proposed s i n c e then  C r o n q u i s t , 1955,  t i a l l y due  1960;  E d i g e r and  to c u l t u r a l problems, Barker  Barkley,  1978).  of Austromontana s p e c i e s than other  s t a t e s ( B a r k e r , 1966,  p. 66),  montana have proved subgenera."  Furthermore,  species belonging  subgenera.  the  He  to the subgenus A u s t r o -  to be l e s s amenable to i n v e s t i g a t i o n than t h o s e  of other  L a s t l y , a l l t h e s p e c i e s i n t h i s subgenus a r e r e l a t i v e l y  a c c e s s i b l e i n Western N o r t h  par-  (1966) d i d l e s s i n v e s t i g a t i o n of  reproductive behavior  "The  (Maguire,  easily  America.  HISTORICAL BACKGROUND  T r i b a l P o s i t i o n of A r n i c a  Although  i t has  l o n g been p l a c e d i n the Senecioneae, A r n i c a , w i t h i t s  broad  r a y s , o p p o s i t e l e a v e s , s e t o s e r e c e p t a c l e , base chromosome number of  x=19,  and  a p o m i c t i c element, does not  f i t conveniently i n this  Nordenstam (1977) suggests  either a close a f f i n i t y  t h a t p o s s i b l y a new  the " A r n i c e a e " ,  tribe,  some e l e v e n other genera.  tribe.  to the H e l i a n t h e a e  or  should be e r e c t e d f o r A r n i c a and  Baag^e (1977) p r e s e n t s e v i d e n c e  c h a r a c t e r s f o r i n c l u s i o n i n the H e l i a n t h e a e .  Chemical  data  of l i g u l e (Siegler  microet a l . ,  3  1974;  Robins, 1977)  supports the anomalous p o s i t i o n i n the Senecioneae.  However, Turner and P o w e l l (1977) b e l i e v e t h a t A r n i c a should be l e f t Senecioneae. in  i n the  The pappus of c a p i l l a r y b r i s t l e s i s p a r t i c u l a r l y out of p l a c e  the Heliantheae.  The s i t u a t i o n i s w e l l summarized  by C r o n q u i s t  (1977,  p. 148), " I t seems l i k e l y t h a t t h e s e two genera A r n i c a and Doronicum c o n t i n u e to be an embarrassment z a t i o n s i n the f a m i l y . "  to f u t u r e as w e l l as p r e s e n t t r i b a l  will organi-  F u r t h e r r e l a t i o n s h i p s of A r n i c a to Whitneya  (Sierra  Nevada), A r n i c a s t r u m (Northwest M e x i c o ) , Jamesianthus (Southeast U n i t e d S t a t e s ) , and M a l l o p a t o p u s (Japan) and p o s s i b l y o t h e r genera need more c r i t i cal  e v a l u a t i o n b e f o r e they may  t r i b e or p l a c e d i n a new  be a s s i g n e d w i t h c o n f i d e n c e to an  existing  tribe.  Taxonomic H i s t o r y  Maguire (1943) g i v e s an e x c e l l e n t h i s t o r i c a l account of t h e genus as a whole.  While l i t t l e more w i l l be d i s c u s s e d c o n c e r n i n g the remaining f o u r  subgenera  ( A r c t i c a , Montana, Andropurpurea, and Chamissonis) a more d e t a i l e d  account of the t a x a i n Maguire's subgenus Austromontana, subgenera, Austromontana  t r e a t e d here as  and C a l a r n i c a , seems a p p r o p r i a t e .  The f i r s t name proposed f o r a s p e c i e s i n Austromontana was f o l i a by Bongard at  Sitka, Alaska.  Arnica  lati-  (1832) which he d e s c r i b e d from m a t e r i a l c o l l e c t e d by Mertens T h i s proved to be the f i r s t  a p p l i e d t o what a r e now of  two  i n a long s e r i e s of names  i n t e r p r e t e d as a l l v a r i a n t s or even t y p i c a l  t h e most polymorphic s p e c i e s i n t h e subgenus,  specimens  i f n o t i n t h e whole genus.  Hooker (1834) i n the F l o r a B o r e a l i - A m e r i c a n a named A r n i c a  cordifolia  from the Rocky Mountains, a g a i n the f i r s t name o f many t o be a p p l i e d polymorphic, and the most widespread s p e c i e s i n t h i s subgenus.  to a  He a l s o named  4  A r n i c a m e n z i e s i i from the "Northwest Coast of America", the f i r s t s p e c i e s now  considered  Nuttall  as  synonyms of A r n i c a  (1841) d e s c r i b e d  a large-leaved  e a s t e r n Oregon, A r n i c a m a c r o p h y l l a , now of t y p i c a l A r n i c a c o r d i f o l i a .  The  p l a n t from the B l u e Mountains of merely a l a r g e - l e a v e d  s y l v i s prope Monterey".  latifolia,  ot genuina and  m e n z i e s i i under A r n i c a  Herder  (1867) r e c o g n i z e d  3 a n g u s t i f o l i a , and  and  considered  Arnica c o r d i f o l i a var.  California The  synonymous w i t h  eradiata  (now  (Gray, 1878)  and  (1897, 1900,  1917,  names, 17 of which a r e now  relegated  Arnica l a t i f o l i a .  Ediger  and  Arnica  saw  good s p e c i e s  today.  the g r e a t e s t number of new  t o synonymy under o t h e r  synonymous w i t h  and  i n the p r e s e n t  considered  Greene (1900, 1901,  1902,  1910)  a l t h o u g h Greene's A r n i c a t o m e n t e l l a  i t a variety  of  l i k e w i s e proposed 16  remains a q u e s t i o n a b l y  a  others  from Southwestern Oregon r e t a i n e d still  Only  r e t a i n e d as  study, a l t h o u g h  1978)  names, w i t h o n l y A r n i c a s p a t h u l a t a  new  5 o l d e r names.  Barkley,  names  species.  1927), i n a s e r i e s of p a p e r s , proposed 18  good s p e c i e s by Maguire (1943) and 1955;  parviflora  A r n i c a n e v a d e n s i s from L a s s e n ' s Peak,  to 1927  considered  three  r a r e A r n i c a v i s c o s a from  A r n i c a . g r a c i l i s from the Rocky Mountains of Western Montana was  (Cronquist,  Arnica  i n subgenus C h a m i s s o n i s ) .  (Gray, 1884), b o t h of which a r e c o n s i d e r e d  t h r e e decades from 1897  names, A r n i c a  both = A r n i c a d i s c o i d e a )  d i s t i n c t i v e and  proposed, most of which a r e now Rydberg  v a r i e t i e s of  Gray (1884) proposed  other  (= A r n i c a d i v e r s i f o l i a  However, he d i d name the v e r y Shasta, C a l i f o r n i a  by Theodor Hartweg  latifolia.  l a t i f o l i a var. v i s c i d u l a  Mt.  two  named  synonymized Hooker's A r n i c a  In the S y n o p t i c a l F l o r a of N o r t h America, Asa names which a r e now  form  r a y l e s s taxon i n the group was  A r n i c a d i s c o i d e a by Bentham (1849) from m a t e r i a l c o l l e c t e d "in  those  latifolia.  considered  first  of  new  today,  valid  entity  5  (see d i s c u s s i o n under A r n i c a  nevadensis).  Howell (1900) named the s e r p e n t i n e J o s e p h i n e County, Oregon, A r n i c a cernua.  s o i l endemic which he c o l l e c t e d i n I t i s one of the most  of t h e r a d i a t e s p e c i e s , as w e l l as one o f the r a r e s t .  distinctive  Nelson (1901, 1909)  proposed the names A r n i c a p l a t y p h y l l a (= A r n i c a l a t i f o l i a ) ,  Arnica  Columbiana,  A r n i c a a r c a n a (both = A r n i c a g r a c i l i s ) , and A r n i c a p a n i c u l a t a (? = A r n i c a cordifolia;  see d i s c u s s i o n under t h a t s p e c i e s ) .  Jones (1910) proposed v a r i e t y g r a c i l i s of A r n i c a b e t o n i c a e f o l i a Greene (= A r n i c a l a t i f o l i a ) .  Hall  (1915) named the r a r e but d i s t i n c t i v e A r n i c a  venosa which he c o l l e c t e d i n Shasta County, C a l i f o r n i a . Arnica eripoda  (= A r n i c a l a t i f o l i a ) .  Piper  Gandoger (1918) named  (1920) named A r n i c a  from Skeena, B r i t i s h Columbia, another o f t h e l a r g e - l e a v e d A r n i c a c o r d i f o l i a , and A r n i c a a p h a n a c t i s a form of A r n i c a Fernald  latifolia. f o r p l a n t s from the  of M i c h i g a n , f a r d i s j u n c t from Western North American  Austromontana s p e c i e s .  I t was c o n s i d e r e d  a subspecies  of A r n i c a c o r d i f o l i a by  Maguire (1943) and i s not g i v e n taxonomic s t a t u s by E d i g e r or i n the p r e s e n t Williams  shade forms o f  from Mt. Baker, Washington, which i s  (1935) proposed the name A r n i c a w h i t n e y i  Keewenaw P e n i n s u l a  andersonii  and B a r k l e y  (1978)  treatment.  (1935) p l a c e d  Greene's A r n i c a t e u c r i i f o l i a as a v a r i e t y o f  A r n i c a l a t i f o l i a and S t . John (1937) named A r n i c a h a r d i n a e (= A r n i c a  cordi-_  f o l i a ) from Benewah County, Idaho, t h e l a s t new name proposed i n t h e subgenus. Maguire (1943), i n h i s worldwide monograph o f the genus, presented t h e most c o n s e r v a t i v e and  treatment o f t h e genus to that date,  r e c o g n i z i n g 13 s p e c i e s  6 i n f r a s p e c i f i c taxa i n Austromontana (see Appendix I ) .  been l a r g e l y accepted  H i s treatment has  by l a t e r workers and f o r the most p a r t t h e f i n d i n g s o f  6 the present study support h i s work. e n t i r e l y on morphological understanding  Although h i s work was based almost  characters, he seems to have had an e x c e l l e n t  of the genus.  However, the greatest shortcoming of h i s work  i s that h i s keys are long and often very d i f f i c u l t to use. L i t t l e taxonomic work has been done since Maguire's monograph other than some new combinations and a r b i t r a r y lumping of species.  I n a study of Great  Basin p l a n t s Maguire (1947) recognized four v a r i e t i e s of Arnica c o r d i f o l i a subsp. genuina: Cronquist  v a r i e t i e s c o r d i f o l i a , maefophylla,  pumila, and h u m i l i s .  (1955), i n the F l o r a of the P a c i f i c Northwest, considered  Arnica g r a c i l i s a v a r i e t y of Arnica l a t i f o l i a and he lumped Arnica p a r v i f l o r a and Arnica g r a y i under the new combined name Arnica discoidea var. eradiata (A. Gray) Cronquist.  He l a t e r (Cronquist, 1958) changed the status of Arnica  p a r v i f l o r a subsp. a l a t a (Rydb.) Maguire to Arnica discoidea var. a l a t a (Rydb.) Cronquist. In the most recent c l a s s i c a l monograph of the genus f o r North American F l o r a , Ediger and Barkley (1978) b a s i c a l l y f o l l o w Maguire's work except that they use Cronquist's new combinations and that they t r e a t Maguire's subspecies as v a r i e t i e s .  They recognize 9 species with 8 i n f r a s p e c i f i c taxa i n  Austromontana (see Appendix II) . METHODS AND MATERIALS - GENERAL F i e l d c o l l e c t i o n s were made from n a t u r a l populations of Arnica species from B r i t i s h Columbia and A l b e r t a to Ontario and Michigan and south t o Colorado and Southern C a l i f o r n i a during several extensive c o l l e c t i n g t r i p s from the f a l l of 1976 through the summer of 1979.  Some plants or seeds were a l s o  obtained from other c o l l e c t o r s and from b o t a n i c a l gardens.  7 Small, whole heads were f i x e d 95%  ethanol : 3 p a r t s chloroform :  transferred  to 95%  ethanol  and  i n the f i e l d  i n Carnoy s o l u t i o n (6 p a r t s  1 p a r t g l a c i a l a c e t i c a c i d ) and  refrigerated for cytological  later  study.  L i v e p l a n t s or seeds of many p o p u l a t i o n s were c o l l e c t e d f o r f u r t h e r in  the experimental  garden, greenhouse, and  r e c o r d s were made i n the f i e l d f e e d e r s on the A r n i c a s . and  f u r t h e r study.  growth chamber.  of p o l l i n a t o r s and  Insect  other  Observations  insect v i s i t o r s  Rhizome growth was  A morphological  measured f o r a number of  r e - e v a l u a t i o n was  taxonomic treatment p r e s e n t e d  and  the subgenera Austromontana and  Habit  i n these  and Branching  All  populations.  i n the  other  at  UBC.  f e a t u r e s used i n e l u c i d a t i n g taxa i n  C a l a r n i c a and  i n determining  phylogenetic  subgenera were the f o l l o w i n g :  s p e c i e s i n Austromontana a r e herbaceous p e r e n n i a l s a r i s i n g s u r f a c e to s e v e r a l c e n t i m e t e r s  These have a t h i c k woody caudex a few  leaves  conservative  CHARACTERS  from  below the s u r f a c e .  s p e c i e s i n C a l a r n i c a a r e a l s o herbaceous p e r e n n i a l s but  surface.  field  Patterns  rhizomes near the s o i l two  made r e s u l t i n g  i n s e c t s are deposited  DISCUSSION OF  relationships  and  herein.  Voucher specimens of p l a n t s and  Among the m o r p h o l o g i c a l  and  specimens were c o l l e c t e d f o r i d e n t i f i c a t i o n  Specimens from more than 30 h e r b a r i a were s t u d i e d to supplement work.  study  to s e v e r a l c e n t i m e t e r s  The  they l a c k rhizomes. below the  soil  Only the r h i z o m a t o u s s p e c i e s produce s t e r i l e b a s a l r o s e t t e s of  (innovations).  P l a n t s may  near ground l e v e l or h i g h e r on  be unbranched above ground to much-branched  the p l a n t .  In the non-rhizomatous s p e c i e s ,  and  others with  s h o r t rhizomes (as A r n i c a g r a c i l i s ) ,  branched from below the ground l e v e l p r o d u c i n g or nodding and  from one  to many per  representation  of h a b i t and  stem.  the p l a n t may  a clump.  Figures 1 - 9  average branching  Heads may  be be  erect  show a s c h e m a t i c  patterns.  Underground P a r t s - Rhizomes, C a u d i c e s , and"Roots  All slender  species (1-2  mm  i n Austromontana are rhizomatous. thick),  d a r k brown, and  n a n t l y terminal branches.  (especially Arnica g r a c i l i s ) and  unbranched or w i t h a few  and  In some of  e c o t y p e s of o t h e r s  (Arnica  A r n i c a cernua) s c a l e s a r e more numerous or appear to  g i v e the rhizome a much more t h i c k e n e d  of most s p e c i e s , b e i n g markedly g r e a t e r than i n o t h e r s  Table V).  stem bases from  w i t h i n a s p e c i e s or p o p u l a t i o n  W i t h few  ious a r t i f i c i a l in  Rhizomes spread  and  deeper rhizomes (6-10  cm)  (e.g., A r n i c a c o r d i -  Arnica l a t i f o l i a ) .  (See in  from y e a r l y a v e r a g e  under  var-  production  field. centimeters  i n most f o r e s t and  d r y h a b i t a t s , some p o p u l a t i o n s ,  populations  average y e a r l y rhizome p r o d u c t i o n  h o r i z o n t a l l y a few  s u r v a c e i n most s p e c i e s  of  rhizomes a r e u s u a l l y l o n g e r  c u l t u r a l regimes v a r i e d l i t t l e i n the  pre-  However, l e n g t h seems to be l a r g e l y g e n e t i c a l l y  exceptions  the same p o p u l a t i o n  be.  appearance.  i n some s p e c i e s  ( e . g . , A r n i c a g r a c i l i s and  shady than i n sunny h a b i t a t s . controlled.  species  cordifolia,  Rhizome l e n g t h i s r e l a t i v e l y c o n s i s t e n t w i t h i n a m a j o r i t y  folia)  predomi-  the s h o r t - r h i z o m e d  These s c a l e s , i n a d d i t i o n to p e r s i s t e n t l e a f bases and vious years,  rhizomes a r e u s u a l l y  They a r e naked or s p a r s e l y c l o t h e d w i t h p a i r s of  u s u a l l y i n p e r s i s t e n t brown s c a l e s .  Arnica l a t i f o l i a ,  The  below the  soil  meadow h a b i t a t s .  e s p e c i a l l y of A r n i c a s p a t h u l a t a ,  with v e r t i c a l extensions  or  duff  However, i n tend  which branch g i v i n g  to have rise  9  F i g u r e s 1-3. of A r n i c a  Schematic diagrams of h a b i t and average b r a n c h i n g p a t t e r n  species.  1.  Arnica  gracilis  2.  Arnica  latifolia  3.  A r n i c a cernua  11  F i g u r e s 4-7. of A r n i c a  Schematic diagrams of h a b i t and average b r a n c h i n g p a t t e r n  species.  4.  Arnica  cordifolia  5.  A r n i c a nevadensis  6.  Arnica discoidea  7.  Arnica  spathulata  13 F i g u r e s 8, 9. of A r n i c a  Schematic diagrams o f h a b i t and average branching  species.  8.  Arnica viscosa  9.  A r n i c a venosa  pattern  15  t o h o r i z o n t a l rhizomes ( F i g u r e 7) c l o s e r to the ground Roots a r e s p a r s e l y produced along branching  0.5-1  mm  t h i c k and  branched. tend  the rhizome, except toward  t e r m i n a l o r i g i n of a e r i a l p o r t i o n s .  on the s h o r t e r - r h i z o m e d  s p e c i e s and  ecotypes.  to be t h i c k e r (1-2  Primary r o o t s are u s u a l l y simple  e s p e c i a l l y i n serpentine  as new  or  sparsely  s o i l s the  roots  mm).  Rhizomes g e n e r a l l y remain a l i v e f o r about 3 years w i t h dying  the  There are more r o o t s produced  20-30 cm l o n g , r a t h e r v e r t i c a l and  In d r i e r h a b i t a t s and  surface.  rhizomes grow i n the  the o l d e s t growth  spring.  In C a l a r n i c a b o t h s p e c i e s l a c k rhizomes but r a t h e r p o s s e s s a t h i c k woody p e r e n n i a l caudex, 4-5  cm  are produced each year. ces f o r s e v e r a l y e a r s t h i c k (5-6 mm)  and  i n diameter from which u s u a l l y many f l o w e r i n g The  bases of o l d f l o w e r i n g  g i v i n g them a rough c h a r a c t e r .  long  (to 40 cm  or more) and  tend  stems  stems remain on the The  r o o t s are  caudi-  quite  to be q u i t e v e r t i c a l  and  unbranched.  Leaves  All  s p e c i e s of Austromontana produce b a s a l r o s e t t e s of 4-6  usually long-petiolate leaves.  Flowering  membraneous s c a l e s at the base, grading i n t o s m a l l lower c a u l i n e l e a v e s .  p a i r s of  stems have o p p o s i t e p a i r s of brown  g r a d u a l l y , or more o f t e n  There are t y p i c a l l y 2-4  abruptly,  p a i r s of  opposite  c a u l i n e l e a v e s , the l a r g e s t produced toward the lower or m i d d l e p o r t i o n of stem, becoming s m a l l e r and florescence.  b r a c t - l i k e and  sometimes a l t e r n a t e below the i n -  C a u l i n e l e a v e s are s e s s i l e to long p e t i o l a t e , w i t h  o f t e n winged, r a r e l y wider than the b l a d e s . p o s i t i o n of the l e a v e s on the f l o w e r i n g l e a v e s are  held.  the  the p e t i o l e s  There i s much v a r i a t i o n i n d i s -  stem as w e l l as the angle a t which  the  16  Leaf cordate  o u t l i n e ranges from l a n c e o l a t e t o b r o a d l y  bases and acute to rounded t i p s .  s e r r a t e , dentate,  Margins may be e n t i r e t o c r e n a t e ,  or doubly s e r r a t e , dentate,  o f t e n combinations o f these  ovate, w i t h acute to  or crenate,  o f t e n u n d u l a t e , and  characters.  Leaves a r e one o f the most r e l i a b l e c h a r a c t e r s  i n separating  taxa  ( F i g u r e s 10-25), a l t h o u g h they a r e one o f the most v a r i a b l e c h a r a c t e r s some t a x a .  There i s u s u a l l y much i n t e r p o p u l a t i o n a l c o n s i s t e n c y ,  within  although  t h e r e may be a g r e a t d e a l of p h e n o t y p i c v a r i a t i o n w i t h i n a s i n g l e p o p u l a t i o n . Thick,  r e l a t i v e l y narrow sun-leaves o r t h i n , r e l a t i v e l y broad  are o f t e n found w i t h i n p o p u l a t i o n s in Arnica l a t i f o l i a and  shade-leaves  growing i n open f o r e s t h a b i t a t s , e s p e c i a l l y  and A r n i c a cernua.  from p l a n t s growing i n s e r p e n t i n e  Leaves, e s p e c i a l l y i n sunny l o c a t i o n s s o i l s , are often flushed with red or  purple. There a r e no b a s a l r o s e t t e s i n subgenus C a l a r n i c a . the f l o w e r i n g  stems a r e v e r y prominent and o f t e n s e v e r a l (6-12) p a i r s which  grade g r a d u a l l y i n t o c a u l i n e l e a v e s . with  S c a l e s a t t h e base o f  Stems of t h i s subgenus a r e much l e a f i e r ,  7-15 o r more p a i r s o f s e s s i l e ovate l e a v e s on t h e primary a x i s and  smaller  l e a v e s and/or b r a c t s on numerous l a t e r a l branches and i n the l e a f y  inflorescence.  Upper b r a c t s a r e u s u a l l y sub-opposite o r a l t e r n a t e .  Odors o f Herbage  Although d i f f i c u l t with  to d e s c r i b e q u a l i t a t i v e l y , pungent odors a s s o c i a t e d  those s p e c i e s b e a r i n g  rating  some s p e c i e s .  glandular  h a i r s a r e e x c e l l e n t c h a r a c t e r s f o r sepa-  While odors a r e p a r t i c u l a r l y u s e f u l i n t h e f i e l d ,  herbarium specimens o f c e r t a i n s p e c i e s r e t a i n t h e i r odor f o r many y e a r s , bly Arnica viscosa.  nota-  17  F i g u r e s 10-17.  Average B a s a l and C a u l i n e Leaves of A r n i c a S-1676, WA,  Species.  10.  B a s a l l e a f of A r n i c a l a t i f o l i a  C l a l l a m County.  11.  C a u l i n e l e a f of A r n i c a l a t i f o l i a  12.  B a s a l l e a f o f A r n i c a c o r d i f o l i a S-1819, OR, Wasco County.  13.  C a u l i n e l e a f of A r n i c a c o r d i f o l i a S-1354, BC, B o t a n i e V a l l e y .  14.  B a s a l l e a f of A r n i c a cernua S-1743, OR, J o s e p h i n e County.  15.  C a u l i n e l e a f of A r n i c a cernua Sweetser, _s._n. (ORE), OR, J o s e p h i n e  S-1654, BC, Tod Mountain.  County. 16.  B a s a l l e a f o f A r n i c a g r a c i l i s S-1603, AB, Lake L o u i s e .  17.  Cauline l e a f of A r n i c a g r a c i l i s C h r i s t  19463 ( I D ) , Elmore County.  18  19  F i g u r e s 18-25.  Average b a s a l and c a u l i n e l e a v e s of A r n i c a  18.  B a s a l l e a f of A r n i c a s p a t h u l a t a  19.  Cauline  species.  S-1381, OR, J o s e p h i n e County.  l e a f o f A r n i c a s p a t h u l a t a D e t l i n g 4688 (ORE), OR, J o s e p h i n e  County. 20.  B a s a l l e a f o f A r n i c a d i s c o i d e a S-1789, CA, Marin County.  21.  Cauline  22.  B a s a l l e a f o f A r n i c a n e v a d e n s i s S-1910, NV, Washoe County.  23.  C a u l i n e l e a f of A r n i c a n e v a d e n s i s S-1910, NV, Washoe County.  24.  Cauline  l e a f of A r n i c a v i s c o s a Muth 733 (PUA), CA, S i s k i y o u County.  25.  Cauline  l e a f o f A r n i c a venosa S-1793, CA, Shasta County.  l e a f o f A r n i c a d i s c o i d e a Copeland s.n. (ORE), CA, B u t t e County.  20  All  s p e c i e s have odors when the l e a v e s a r e b r u i s e d except  l a t i f o l i a and A r n i c a cernua.  Rarely Arnica l a t i f o l i a ,  Arnica  o r suspected  hybrids  of i t and A r n i c a c o r d i f o l i a a r e s l i g h t l y f r a g r a n t , e s p e c i a l l y i n younger, r a p i d l y growing l e a v e s . grant-leaved of odors. related  The odor i s l o s t  i n older leaves.  Among the f r a -  s p e c i e s t h e r e a r e c o n s i s t e n t d i f f e r e n c e s i n q u a l i t y and degree  A r n i c a v i s c o s a has by f a r t h e s t r o n g e s t odor, w h i l e the c l o s e l y  s p e c i e s , A r n i c a venosa, has o n l y a s l i g h t f r a g r a n c e , a t l e a s t i n t h e  few p o p u l a t i o n s s t u d i e d i n the f i e l d .  There a l s o seem t o be c o n s i s t e n t d i f -  f e r e n c e s i n the odors o f c l o s e l y r e l a t e d s p e c i e s , as i n A r n i c a c o r d i f o l i a and A r n i c a nevadensis,  although  p o p u l a t i o n s of each o f these  t h e r e i s some v a r i a t i o n  i n odor i n d i f f e r e n t  species.  Vestiture  Maguire (1943, p. 403) comments on the use o f pubescence as a c r i t i c a l c h a r a c t e r i n work on Doronicum by C a v i l l i e r by I l j i n alone.  (1907) and the Russian  (1926), b u t he c a u t i o n s a g a i n s t r e l y i n g  Arnicas  too h e a v i l y on t h i s  Pubescence, o r the l a c k t h e r e o f , i s a good m o r p h o l o g i c a l  character  character,  e s p e c i a l l y i n t h e f i e l d , when used i n c o n j u n c t i o n w i t h other c h a r a c t e r s f o r determining  A r n i c a taxa.  With few e x c e p t i o n s  a l l p a r t s o f a l l s p e c i e s a r e to  some degree pubescent, most o f t e n densely s o . There a r e two b a s i c h a i r types glandular.  Each o f these  i n A r n i c a , s e p t a t e g l a n d u l a r and non-  types commonly occur  i n two t y p e s , l o n g (1-2 mm)  and  s h o r t (0.1-0.2 mm), although  The  s t i p i t a t e g l a n d u l a r h a i r s have s e p t a t e s t a l k s u s u a l l y two c e l l s t h i c k  except  t h e r e a r e numerous i n t e r m e d i a t e  j u s t beneath t h e g l a n d where they a r e o n e - c e l l e d .  posed o f two to many c e l l s .  examples.  The g l a n d i s com-  A m a j o r i t y o f the s t i p i t a t e g l a n d u l a r h a i r s a r e  about 1 mm long on s t a l k s composed o f 6-8 c e l l s , o r they a r e q u i t e s h o r t  22  (0.1-0.2 mm) The  with  s t a l k s 2-3 c e l l s  long.  degree o f g l a n d u l a r pubescence i m p a r t s  "clammy" and a r o m a t i c  character  a more o r l e s s v i s c i d o r  to t h e l e a v e s and stems o f c e r t a i n s p e c i e s ,  e s p e c i a l l y Arnica c o r d i f o l i a , Arnica nevadensis, c h a r a c t e r alone makes them r e a d i l y  separable  largely  l a c k i n g g l a n d u l a r h a i r s , most n o t a b l y  folia.  In Arnica g r a c i l i s  i n the f i e l d  than 0.1 mm  appearance under 10X m a g n i f i c a t i o n .  species lati-  i s the short, nearly  l o n g g i v i n g t h e l e a v e s a rough  In the other  species bearing  glandular  t h e r e a r e m i x t u r e s o f both the l o n g and s h o r t h a i r s , except i n A r n i c a  v i s c o s a which has o n l y l o n g g l a n d u l a r  hairs.  N o n - g l a n d u l a r s e p t a t e h a i r s a r e always one c e l l bases and they a r e u s u a l l y s t r a i g h t ,  the exceptions  t h i c k except a t t h e i r being  o f t h e stems of some s p e c i e s , e s p e c i a l l y A r n i c a c e r n u a . two  from those  This  A r n i c a cernua and A r n i c a  t h e predominant h a i r type  s e s s i l e , glandular h a i r less  hairs  and A r n i c a v i s c o s a .  s i z e s of non-glandular  ( F i g u r e 62).  h a i r s a r e seen, l o n g ones 1-2 (-3) mm  s h o r t ones 0.3-0.5 mm l o n g , w i t h g l a n d u l a r h a i r s imparts  on t h e lower p a r t  some i n t e r m e d i a t e s .  Again  long and  The number o f non-  a s o f t n e s s to t h e l e a v e s o f A r n i c a c o r d i f o l i a ,  Arnica  d i s c o i d e a , and A r n i c a venosa. In c e r t a i n s p e c i e s o r p o p u l a t i o n s notably  the y e l l o w - t i p p e d  the g l a n d u l a r h a i r s are c o l o r e d ,  h a i r s o f A r n i c a v i s c o s a and the p u r p l e - t i p p e d o r  based h a i r s o f A r n i c a s p a t h u l a t a and t o a l e s s e r degree A r n i c a d i s c o i d e a . F i g u r e s 26 to 126 show t h e v e s t i t u r e t y p e s the s p e c i e s i n t h e s e dance o f v e s t i t u r e .  of a l l vegetative parts of  two subgenera, but w i t h no i n d i c a t i o n o f r e l a t i v e abun-  23  F i g u r e s 26-126.  Comparison of V e s t i t u r e Types on Four V e g e t a t i v e of A r n i c a S p e c i e s .  Parts  24  25  Leaf, adaxial Taxon  cordifolialatifolia natural hybrid  Leaf, abaxial  26 Lower Cauline Taxon  gracilis  nevadensis  cernua  Upper cauline, Periclinum, P h y l l a r i e s  27  Leaf, adaxial  Leaf, abaxial  Taxon  cernua  none  none  28  29  spathulata  30  31  Leaf, adaxial  Leaf, abaxial  Taxori  1mm  viscosa  (§) venosa  32  Capitula  Heads v a r y from t u r b i n a t e to campanulate i n the few-flowered ( i . e . , A r n i c a v i s c o s a and the r e m a i n i n g s p e c i e s .  Arnica g r a c i l i s )  to b r o a d l y  h e m i s p h e r i c i n most of  They are l e s s than 1 cm broad i n the d i s c o i d A r n i c a  v i s c o s a to n e a r l y 6 cm broad i n t e r m i n a l r a d i a t e heads of A r n i c a Heads i n d i f f e r e n t  s p e c i e s may  be r a d i a t e w i t h a s i n g l e row  f l o r e t s or d i s c o i d , l a c k i n g ray f l o r e t s . t h e r e may  be  the n o r m a l l y mally  1-3  elongated  In the n o r m a l l y  r a y - l i k e (ampliate)  florets  r a d i a t e s p e c i e s the r a y f l o r e t s may  heads per  species  cordifolia.  of m a r g i n a l  d i s c o i d species  ( F i g u r e s 154-156) and  be absent.  stem, w i t h a l a r g e r t e r m i n a l one  and  some p o p u l a t i o n s  lata.  of A r n i c a l a t i f o l i a ,  Heads are h e l d e r e c t at a n t h e s i s  a p a i r or several  t e n t l y nodding i n bud at  A r n i c a d i s c o i d e a , and i n most s p e c i e s .  stem as  Arnica  They are  i n A r n i c a cernua and A r n i c a venosa and  a 45-60° angle at a n t h e s i s  in  There are n o r -  p a i r s o f l a t e r a l heads, w i t h r a r e l y as many as 30 o r more heads per in  ray  spathu-  consis-  are o f t e n h e l d  i n A r n i c a cernua, A r n i c a c o r d i f o l i a , and  Arnica  venosa.  Phyllaries  Average number, s i z e and shown i n F i g u r e s 127-136. within populations,  shape of i n v o l u c r a l b r a c t s  ( p h y l l a r i e s ) are  These are v a r i a b l e c h a r a c t e r s w i t h i n s p e c i e s  but o f f e r some c o n s i s t e n t d i f f e r e n c e s i n some t a x a .  are l a n c e o l a t e to ovate w i t h acute t i p s , except A r n i c a cernua, A r n i c a l a t a , and  A r n i c a d i s c o i d e a which o f t e n have rounded t i p s .  species.  Arnica l a t i f o l i a  All  spathu-  P h y l l a r i e s are  c o n s i s t e n t l y s m a l l e r and narrower i n A r n i c a v i s c o s a and A r n i c a g r a c i l i s i n other  and  P u t a t i v e n a t u r a l h y b r i d s between A r n i c a c o r d i f o l i a c o n s i s t e n t l y have more p h y l l a r i e s than e i t h e r  parent.  than  and  33  F i g u r e s 127-136.  Average I n v o l u c r a l B r a c t s o f A r n i c a  127.  A r n i c a n e v a d e n s i s S-1870, CA, S i e r r a  128.  A r n i c a c o r d i f o l i a S-1343, BC, B o t a n i e V a l l e y .  129.  Arnica c o r d i f o l i a  X Arnica l a t i f o l i a  S-1475, MT, R a v a l l i  County.  (putative n a t u r a l hybrid)  County.  S-1500, WA,  Species.  130.  Arnica l a t i f o l i a  Chelan County.  131.  A r n i c a g r a c i l i s S-1605, AB, Moraine Lake.  132.  A r n i c a cernua S-1386, OR, J o s e p h i n e County.  133.  A r n i c a d i s c o i d e a S-1825, WA,  134.  A r n i c a s p a t h u l a t a S-1380, OR, J o s e p h i n e County.  135.  A r n i c a venosa S-1940, CA, Shasta County.  136.  A r n i c a venosa S-1411, CA, S i s k i y o u  Klickitat  County.  County.  34  illlllhllll I U 11111) 11 |  iMIIIIIlUli® l l l l l l l t l l l l l ( S )  ililiiil®  35  Margins of p h y l l a r i e s are u s u a l l y e n t i r e , a l t h o u g h i n r a r e cases they have a few  irregular teeth.  Bases of the p h y l l a r i e s and  the p e r i c l i n u m  ( a r e a of  attachment of p h y l l a r i e s to the peduncle) are u s u a l l y the most densely  pubes-  cent p a r t o f most p l a n t s and o f t e n have the l o n g e s t h a i r s on the p l a n t .  Ray  Florets  A l l ray f l o r e t s c o n s i s t of a s h o r t tube and 137-142). general  The  tube and  lower s u r f a c e of the limb  o u t l i n e of the limb  more f r e q u e n t l y one  l a r g e f l a t t e n e d limb  are v a r i a b l y pubescent.  specimens.  The  to s e v e r a l c h a r a c t e r i s t i c t e e t h .  The  s i z e s and  shapes of  These a r e  c h a r a c t e r s , not always apparent i n p r e s s e d  es-  herbarium  number of rays v a r i e s from t y p i c a l l y f i v e o r r a r e l y l e s s i n  depauperate heads to o c c a s i o n a l l y twenty or more per head. bright yellow  The  i s l i n e a r oblong to ovate w i t h a rounded t i p or  these t e e t h are used i n d i s t i n g u i s h i n g between some s p e c i e s . p e c i a l l y good f i e l d  (Figures  and  p i s t i l l a t e w i t h an elongated  Rays are  s t y l e extending  always  beyond the  tube.  Disc F l o r e t s  Heads c o n t a i n as few  as 9-10  to more than 180  disc florets.  m o s t l y t u b u l a r or goblet-shaped ( F i g u r e s 143-153). t i v e r i n g at the base where the tube a t t a c h e s about midway up p o i n t the limb  There i s o f t e n a d i s t i n c -  to the ovary and  another r i n g  the tube where the f i l a m e n t s are a t t a c h e d w i t h i n . f l a r e s out, a b r u p t l y  At the d i s t a l end  of the limb are f i v e r e c u r v e d  f l o r e t s are f r a g r a n t .  lobes.  this  i n which the'y are w h i t e or cream-colored.  Disc f l o r e t s are  N e c t a r i s produced and  They are always b r i g h t y e l l o w  translucent.  At  i n some cases, v e r y g r a d u a l l y i n o t h e r s .  f e c t w i t h f i v e a n t h e r s u n i t e d around the p i s t i l .  often nearly  These are  perthe  except i n A r n i c a v i s c o s a  W a l l s of the tube and  D i s c f l o r e t s are s p a r s e l y to densely  limb  are  h a i r y on  the  36  F i g u r e s 137-142.  T y p i c a l r a y f l o r e t s o f A r n i c a subgenus  137.  A r n i c a c o r d i f o l i a S-1353, BC, B o t a n i e V a l l e y .  138.  Arnica c o r d i f o l i a X Arnica l a t i f o l i a  Austromontana.  h y b r i d S-1474, MT,  County. 139.  Arnica l a t i f o l i a  S-1680, BC, Manning Park.  140.  A r n i c a cernua S-1386, OR, Josephine County.  141.  A r n i c a n e v a d e n s i s S-1870, CA, S i e r r a  142.  A r n i c a g r a c i l i s S-1605, AB, Moraine Lake.  County.  Ravalli  37  38  F i g u r e s 143-148.  T y p i c a l D i s c F l o r e t s o f A r n i c a S p e c i e s i n Subgenus  Austromontana. 143.  A r n i c a c o r d i f o l i a S-1353, BC, B o t a n i e V a l l e y .  144.  Arnica c o r d i f o l i a X Arnica l a t i f o l i a  S-1474 ( p u t a t i v e n a t u r a l h y b r i d ) ,  MT, R a v a l l i County. 145.  Arnica l a t i f o l i a  S-1476, MT, M i s s o u l a County.  146.  A r n i c a nevadensis S-1870, CA, S i e r r a  147.  A r n i c a g r a c i l i s S-1605, AB, Moraine Lake.  148.  A r n i c a cernua S-1386, OR, Josephine County.  County.  39  40  F i g u r e s 149-152.  T y p i c a l D i s c F l o r e t s o f A r n i c a S p e c i e s i n Subgenera  Austromontana and C a l a r n i c a . 149.  A r n i c a d i s c o i d e a S-1789, CA, Marin County.  150.  Arnica spathulata  151.  A r n i c a v i s c o s a S-1411, CA, S i s k i y o u County.  152.  A r n i c a venosa S-1793, CA, Shasta County.  S-1742, OR, J o s e p h i n e County.  41  42  F i g u r e s 153-156.  F l o r e t s of A r n i c a d i s c o i d e a .  153.  T y p i c a l d i s c f l o r e t , WA,  Klickitat  County, S-1825.  154.  Marginal ampliate f l o r e t  from same head.  155.  E l o n g a t e d m a r g i n a l f l o r e t w i t h a b o r t i v e a n t h e r s , CA,  Santa  Barbara  County, Smith 8330 (SBM). 156.  R a y - l i k e m a r g i n a l f l o r e t w i t h a b o r t i v e a n t h e r s , CA, County, Nelson  & Anderson 843  (HSC).  Mendocino  43  44  tube o r throughout  ( e s p e c i a l l y i n A r n i c a venosa).  There a r e always a few  h a i r s on the t i p s o f t h e l o b e s as w e l l as d i s t i n c t i v e p a p i l l a e on t h e i r surface.  The f l o r e t s a r e u s u a l l y uniform  throughout the head.  i n s i z e and shape and symmetrical  I n many of the d i s c o i d heads o f A r n i c a d i s c o i d e a the  m a r g i n a l f l o r e t s a r e asymmetrical, known as ampliate much elongated  inner  florets  ( F i g u r e 155) o r  and r a y - l i k e w i t h a b o r t i v e anthers ( F i g u r e s 154, 156).  Pappus  The  pappus i s composed o f a r i n g o r r i n g s o f about 27-65 m u l t i c e l l u l a r  b r i s t l e s with  l a t e r a l setae.  The l e n g t h of b r i s t l e s v a r i e s a c c o r d i n g  l e n g t h o f t h e f l o r e t , but t h e l e n g t h o f l a t e r a l setae t e n t w i t h i n some s p e c i e s o r s p e c i e s groups. these setae  i n t h r e e groups a c c o r d i n g  0.1-0.2 mm;  subplumose, 0.2-0.35 mm;  consis-  Maguire (1943, p. 402) t r e a t s  to length:  b a r b e l l a t e , with  and plumose, 0.35-0.6 mm.  i n these two subgenera f a l l w i t h i n the f i r s t The  i s relatively  t o the  setae Most  taxa  two groups ( F i g u r e s 157-165, 186).  pappus o f the whole head forms a round b a l l 1-3 cm i n diameter upon matur-  i t y of t h e achenes.  C o l o r o f t h e mature pappus i s u s u a l l y pure white i n most  s p e c i e s o r o f t e n tawny i n A r n i c a n e v a d e n s i s , A r n i c a v i s c o s a , and A r n i c a gracilis.  Achenes  The  f o r separating  some o f the c l o s e -  l y r e l a t e d s p e c i e s , i s the v e s t i t u r e of the achene s u r f a c e  ( F i g u r e s 166-185).  Table and  best  single reproductive  character  I g i v e s a comparison o f mature achenes.  They a r e u s u a l l y 5-8 mm  long  1 mm wide a t t h e i r w i d e s t p o i n t , the d i s t a l end, l i n e a r oblong to c l a v a t e ,  weakly to s t r o n g l y 5-10 a n g l e d .  The s u r f a c e i s dark grey o r brown t o b l a c k  45  F i g u r e s 157-161. of  Photographs w i t h l i g h t microscope o f near m i d - l e n g t h  pappus.  157.  A r n i c a g r a c i l i s S-1603, AB, Lake  Louise.  158.  A r n i c a l a t i f o l i a S-1162, WA,  159.  A r n i c a cernua S-1772, CA, Humboldt County.  160.  Arnica c o r d i f o l i a  161.  A r n i c a n e v a d e n s i s S-1870, CA, S i e r r a  Whatcom County.  S-1939, CA, Mono County. County.  47  Figures of  162-165.  Photographs w i t h l i g h t microscope of near m i d - l e n g t h  pappus.  162.  A r n i c a d i s c o i d e a D e t l i n g 6050 (ORE), CA, S i s k i y o u  163.  Arnica spathulata  164.  A r n i c a venosa S-1940, CA, Shasta County.  165.  A r n i c a v i s c o s a S-1411, CA, S i s k i y o u  S-1381, OR, J o s e p h i n e County.  County.  County.  48  49  F i g u r e s 166-171.  Scanning E l e c t r o n Micrographs o f Achene S u r f  (mid-length) o f A r n i c a  species.  166.  Arnica c o r d i f o l i a  S-1353, BC, B o t a n i e V a l l e y , X65.  167.  same, X130.  168.  A r n i c a n e v a d e n s i s S-1870, CA, S i e r r a County, X65.  169.  same, X130.  170.  A r n i c a d i s c o i d e a S-1825, WA,  171.  same, X130.  Klickitat  County, X65.  50  51  F i g u r e s 172-177.  Scanning E l e c t r o n Micrographs o f Achene Surf  (mid-length) o f A r n i c a  species.  172.  A r n i c a cernua S-1772, CA, Humboldt County, X65.  173.  same, X130.  174.  A r n i c a l a t i f o l i a S-1476, MT, M i s s o u l a County, X65.  175.  same, X130.  176.  A r n i c a g r a c i l i s S-1562, BC, Monroe Lake, X65.  177.  same, X130.  52  53  F i g u r e s 178-184.  Scanning E l e c t r o n Micrographs of Achene S u r f a c e s  (mid-length) o f A r n i c a  species.  178.  A r n i c a s p a t h u l a t a S-1380, OR, J o s e p h i n e County,  179.  same, X130.  180.  A r n i c a venosa S-1940, CA, Shasta County,  181.  same, X130.  182.  A r n i c a v i s c o s a S-1411, CA, S i s k i y o u County,  183.  same, X130.  184.  Arnica g r a c i l i s  S-1562, BC, Monroe Lake, X250.  185.  Photomicrograph  w i t h l i g h t m i c r o s c o p e , achene s u r f a c e o f A r n i c a  cordifolia 186.  S-1487, WA,  Okanogan County,  X65.  X65.  X500.  Scanning e l e c t r o n micrograph o f pappus b r i s t l e Arnica cordifolia  X65.  (mid-length) of  S-1353, BC, B o t a n i e V a l l e y , X575.  TABLE I COMPARISON OF MATURE ACHENES OF ARNICA SPECIES  Taxon  Color  Length  (mm)  Width D i s t a l l y (mm)  Double H a i r s  Glandular Hairs  cordifolia  dark grey  (6-)7-9(-10)  1+  u n i f o r m throughout o r especially distally  uniform  latifolia  grey-brown  ( -)6-8(-10)  1-  none o r few d i s t a l l y  none o r few d i s t a l l y  gracilis  black  (4-)5-6(-7)  1  none or v e r y few distally  none o r few throughout o r many d i s t a l l y  cernua  dark grey  6-8  1+  many throughout o r distally  none  nevadensis  dark grey to b l a c k  (4-)5-8  1+  few throughout o r distally  uniform  discoidea  dark grey  6- 8  1+  few to many throughout  few  spathulata  black  7- 9  1  none to few throughout  few throughout o r distally  venosa  grey  many throughout  none  viscosa  dark grey  none  uniform l o n g - s t i p i t a t e throughout  5  . 6-8 5-6  1.5-2 1  throughout  throughout  throughout  56  and m i n u t e l y c r e s t e d and u n d u l a t e ( F i g u r e 184), a l t h o u g h t h i s i s u s u a l l y obscured by the s u r f a c e h a i r s . the base where the achene was located d i s t a l l y  There i s a t h i c k white r i n g , the annulus, a t a t t a c h e d to the r e c e p t a c l e and another  to which the pappus i s a t t a c h e d .  s h o r t e r and broader than those of the d i s c f l o r e t s . few to many double-pronged  Achenes of ray f l o r e t s  ( F i g u r e 183)  are  Most achene c o a t s have  o r "duplex" h a i r s ( F i g u r e 185), e s p e c i a l l y  l y , i n a d d i t i o n to l o n g s t i p i t a t e glandular h a i r s .  ring  distal-  to n e a r l y s e s s i l e ( F i g u r e  184)  The p o s i t i o n and r e l a t i v e abundance or s p a r s i t y of these  h a i r s i s used i n t h i s study as an important d i a g n o s t i c c h a r a c t e r .  DISTRIBUTION AND  Both o f these subgenera  ECOLOGY  are l a r g e l y Western C o r d i l l e r a n ( i n the broad  sense) w i t h e a s t e r n e x t e n s i o n s o r d i s j u n c t p o p u l a t i o n s i n t o the Cypress  Hills  of Saskatchewan, B l a c k H i l l s of South Dakota, and around Lake S u p e r i o r .  None  of these s p e c i e s i s t r u l y a r c t i c as a r e some of the s p e c i e s i n o t h e r  subgenera.  They o c c u r i n a v a r i e t y of h a b i t a t s from open a l p i n e o r mountain meadows, f o r e s t edges, open f o r e s t s , d i s t u r b e d r o a d s i d e banks, s t a b i l i z e d rock and s t a b i l i z e d burned-over  a r e a s , but m o s t l y c o o l , mesic h a b i t a t s .  A r n i c a venosa i s a p l a n t of the hot, dry f o o t h i l l s of the upper R i v e r V a l l e y of N o r t h e r n C a l i f o r n i a . s p a t h u l a t a are r e s t r i c t e d viscosa i s limited  Two  slides,  Only  Sacramento  s p e c i e s , A r n i c a cernua and  to o r u s u a l l y found on s e r p e n t i n e s o i l s .  Arnica Arnica  to h i g h pumice s l o p e s .  REPRODUCTIVE BIOLOGY  Phenology  Being montane or a l p i n e p l a n t s , a l l s p e c i e s o f Austromontana b e g i n  new  57 innovation growth and flowering stem elongation as soon as spring or summer snows have melted. may  In c o a s t a l low mountains of C a l i f o r n i a and Oregon t h i s  occur i n A p r i l , while at higher a l t i t u d e s and l a t i t u d e s growth may  begin  as l a t e as September or October (or probably not at a l l some years where snow cover i s excessive).  High a l p i n e populations often have elongated  flowering  stems under the melting snow or have stems emerging through the snow. In areas where there i s t y p i c a l l y dry weather f o l l o w i n g the spring snowmelt, as occurs east of the Cascade Mountains and i n the Siskiyou and Klamath Mountains, there i s u s u a l l y a b r i e f flowering period of a few weeks with no more l a t e r a l flower buds being produced a f t e r the monocephalous or few-headed stems have flowered.  This i s the common condition i n most species.  h a b i t a t s populations of the same species or other species may branch and flower f o r months.  In wetter  continue to  Other species, e s p e c i a l l y Arnica l a t i f o l i a ,  produce only one " f l u s h " of flowers regardless of the weather conditions thereafter. In a growth chamber under conditions of 16 hour days, 25°C, and adequate moisture and f e r t i l i z e r , populations of several species, Arnica c o r d i f o l i a , Arnica d i s c o i d e a , A r n i c a venosa, and e s p e c i a l l y Arnica v i s c o s a continued to produce l a t e r a l branches and new growth from rhizomes and flowered l y f o r more than a year.  continuous-  Other experimental populations of Arnica c o r d i f o l i a ,  Arnica d i s c o i d e a , Arnica spathulata, Arnica g r a c i l i s , Arnica nevadensis, and e s p e c i a l l y Arnica l a t i f o l i a , produced only one " f l u s h " of heads.  Although  leaves remained green f o r months and new vegetative growth was produced, no more flower buds were produced regardless of the growth regime.  Only a f t e r  several weeks of v e r n a l i z a t i o n and then i n c r e a s i n g l y warmer temperatures and increased l i g h t were new  flowering stems produced.  58  A c t u a l f l o w e r i n g time from the opening of the f i r s t to t h e opening o f the l a s t d i s c f l o r e t may few-flowered flowered  d i s c o r ray  be as b r i e f as two  s p e c i e s (e.g., A r n i c a g r a c i l i s )  floret  days i n the  to more than a week i n t h e  larger  species.  During  dry weather l a t e r a l buds o f t e n a b o r t and  y e a r s a l l the f l o w e r buds a b o r t , as observed Arnica c o r d i f o l i a l a t i o n s had  at l e a s t d u r i n g some  w i t h numerous p o p u l a t i o n s  i n c e n t r a l Wyoming d u r i n g June 1978.  f l o w e r i n g stems but no  of  Here, v e r y l a r g e popu-  flowers.  Achenes r i p e n w i t h i n a few weeks of f l o w e r i n g and a r e blown away. rhizome growth begins  about the time of f l o w e r i n g w i t h  emerging d u r i n g o r j u s t a f t e r f l o w e r i n g . and  soon d i s a p p e a r s  until fall  except  i n wetter  innovations  A l l above-ground growth becomes dry  h a b i t a t s where p l a n t s may  remain green  frost.  In C a l a r n i c a the stems become much elongated f l o w e r s are produced. flowering.  the new  New  and o f t e n branched  S e v e r a l weeks o r months may  Flowering may  before  e l a p s e from emergence to  be r a t h e r b r i e f w i t h few heads per stem i n A r n i c a  venosa or l o n g e r f o r the much-branched A r n i c a v i s c o s a .  Cytology  For chromosomal study,  f i x e d anthers  g l a s s s l i d e , a drop of aceto-carmine was w i t h i r o n needles purple.  f o r about 1-3  from s m a l l buds were e x c i s e d on a added, and  minutes u n t i l  the aceto-carmine became dark  A drop o f Hoyer's medium (Beeks, 1955)  permanent, a cover a flame and  s l i p was  squashed.  added, and  the anthers were macerated  was  added to make the  the p r e p a r a t i o n was  warmed b r i e f l y  Cover s l i p s were l a t e r r i n g e d w i t h diaphane.  t i o n s of chromosome number and m e i o t i c behavior  slide  of m i c r o s p o r o c y t e s  over  Observa-  were made  w i t h a p o s i t i v e phase-contrast microscope.  T a p e t a l c e l l s were n o t used f o r  chromosome number d e t e r m i n a t i o n s s i n c e numerous o b s e r v a t i o n s o f h i g h p o l y p l o i d numbers were made i n t a p e t a l c e l l s s u r r o u n d i n g d i p l o i d o r t e t r a p l o i d sporocytes.  Some chromosome number d e t e r m i n a t i o n s were made from  micro-  root-tips  from p o t t e d p l a n t s grown i n the greenhouse or growth chamber o r from achenes germinated  on moist f i l t e r paper.  The r o o t - t i p s were soaked i n a s u p e r s a t u -  r a t e d aqueous s o l u t i o n o f p a r a d i c h l o r o b e n z e n e  f o r 1-2 hours ( S t r o t h e r , 1969)  b e f o r e f i x i n g and s t a i n i n g as f o r f l o w e r bud m a t e r i a l . A r n i c a chromosomes a r e s m a l l ( c a . 5 Um), they do n o t f i x w e l l , and meiotic associations are often d i f f i c u l t  to i n t e r p r e t , r e s u l t i n g  the p r e v i o u s r e p o r t s b e i n g approximations microsporocytes  o r probable e r r o r s .  from a number o f squashes had to be observed  counts o r p a i r i n g was determined.  i n many o f  O f t e n many before accurate  Although d i p l o i d s u s u a l l y form r e g u l a r  b i v a l e n t s d u r i n g metaphase I o f m e i o s i s ( F i g u r e 194), i n the h i g h e r p l o i d y levels  there i s u s u a l l y i r r e g u l a r b e h a v i o r , w i t h v a r i a b l e numbers o f u n i v a -  l e n t s , b i v a l e n t s , m u l t i v a l e n t s , and c h a i n s p r e s e n t a t metaphase I and l a g g i n g chromosomes and b r i d g e f o r m a t i o n d u r i n g anaphase I ( F i g u r e 187) , as has been >. noted p r e v i o u s l y by O r n d u f f , e t a l . (1963, 1967) and Barker Table I I l i s t s  (1966).  a l l p r e v i o u s l y r e p o r t e d chromosome numbers f o r these two  subgenera, a number o f which a r e a p p r o x i m a t i o n s .  There i s o n l y one p r e v i o u s  r e p o r t f o r A r n i c a cernua and A r n i c a venosa and two each f o r A r n i c a g r a c i l i s , A r n i c a s p a t h u l a t a , and A r n i c a v i s c o s a .  Only f o r the two widespread  A r n i c a c o r d i f o l i a and A r n i c a l a t i f o l i a ,  are there s u f f i c i e n t previous reports  to  show t r e n d s i n p l o i d y l e v e l s .  species,  Table I I I shows new chromosome numbers ob-  served d u r i n g t h i s study, more than twice the number of a l l p r e v i o u s r e p o r t s . Counts f o r A r n i c a n e v a d e n s i s  are presented f o r the f i r s t  time.  F i g u r e s 187-  60  189  and  193,  194  show photographs of m e i o t i c chromosomes of A r n i c a  A r n i c a c o r d i f o l i a , and  Arnica discoidea.  gracilis,  F i g u r e s 195-207 show camera l u c i d a  drawings of a l l p l o i d y l e v e l s observed d u r i n g t h i s study, of a l l s p e c i e s recognized. To  summarize known chromosome numbers, f i v e s p e c i e s i n these  are w h o l l y  diploid.  Arnica l a t i f o l i a  one  t e t r a p l o i d r e p o r t e d ) , and  Both d i p l o i d and  Arnica discoidea. p l o i d and  These are A r n i c a v i s c o s a , A r n i c a venosa, A r n i c a cernua,  (one  ploid reported).  subgenera  Arnica spathulata  t e t r a p l o i d populations  Arnica g r a c i l i s  i s predominately  (one  tetra-  are known f o r  t r i p l o i d , w i t h one  tetra-  h e x a p l o i d p r e v i o u s l y r e p o r t e d , a l t h o u g h voucher specimens from  these r e p o r t s have n o t been seen. counts from t h r e e p o p u l a t i o n s . predominately  A r n i c a n e v a d e n s i s i s t e t r a p l o i d , based  A r n i c a c o r d i f o l i a i s the most v a r i a b l e , b e i n g  t e t r a p l o i d , w i t h a few  p l o i d populations  on  d i p l o i d , t r i p l o i d , p e n t a p l o i d , and  hexa-  known.  Apomixis  To determine i f p l a n t s were a p o m i c t i c , heads were bagged w i t h c l o t h bags ( F i g u r e 211)  before  the r a y s opened on r a d i a t e s p e c i e s , or b e f o r e  d i s c s opened on d i s c o i d s p e c i e s . a r e s e l f - i n c o m p a t i b l e , and However, i t was  t h e r e i s a d d i t i o n a l data  achenes from bagged heads were indeed  anthers  to stigmas,  outer  Barker (1966) showed t h a t s e x u a l d i p l o i d s to support  his findings.  assumed t h a t known p o l y p l o i d p o p u l a t i o n s producing  amphimicts ( s e x u a l p l a n t s ) .  the  apomicts r a t h e r than  To i n s u r e t h a t p o l l e n was  viable  self-compatible  being  t r a n s f e r r e d from  heads were g e n t l y rubbed on s e v e r a l s u c c e s s i v e days,  the f l o r e t s matured and  stigmas emerged from t h e surrounding  the achenes matured each was  checked f o r c o n t e n t s .  anthers.  A r n i c a achenes a r e  as  When  61  F i g u r e s 187-194. Grains 187.  of A r n i c a  M e i o t i c Chromosomes, M i c r o s p o r o c y t e s ,  and P o l l e n  Species.  A r n i c a g r a c i l i s S-1613, BC, Peyto Lake, anaphase I (upper t r i p l o i d with  l a g g i n g chromosomes.  188.  Late  189.  L a t e anaphase I I of same.  190.  " Q u i n t a d " and "Sextad" o f same.  191.  A r n i c a l a t i f o l i a S-1961, BC, Nancy Greene P r o v i n c i a l Park,  anaphase I o f same.  p o l l e n grains stained with cotton blue two u n s t a i n e d 192.  cell),  i n lactophenol,  showing  grains.  A r n i c a nevadensis S-1910, NV, Washoe County, SEM o f p o l l e n g r a i n s showing l a r g e and s m a l l g r a i n s , X1200.  193.  A r n i c a c o r d i f o l i a S-1416, CA, S i e r r a County, t e t r a p l o i d with  194.  37 chromosomes a t each p o l e o f the c e l l  and one l a g g i n g  A r n i c a d i s c o i d e a S-1787, CA, Napa County, d i p l o i d p a i r s of chromosomes a t metaphase I .  (2n=76) pair.  (2n=38) w i t h 19  Bars = ca 50 Jim.  63  TABLE I I .  P r e v i o u s l y Reported Chromosome Numbers f o r A r n i c a i n the Subgenera Austromontana and C a l a r n i c a .  Taxon  Sporophytic Number  Locality  cernua  38  OR,  cordifolia  76  BC, Lambly  cordifolia  76  AB  cordifolia  76  SK, Cypress  Species  Reference  Josephine  Co.  Creek  Barker,  1966  T a y l o r & T a y l o r , 1977 Ornduff, et a l . , 1967  Hills  T a y l o r & Brockman, 19(  P r o v i n c i a l Park cordifolia  ca76  WA,  Stevens  cordifolia  72-76  OR,  Lake Co.  cordifolia  ca76  ID, Lewis Co.  cordifolia  38  MT,  Co.  ca90+6  CO, L a P l a t a  cordifolia  ca95+4  CO, Huerfana  cordifolia  ca95  CO, Larimer  cordifolia  60  discoidea  ca72-76  Co.  Co. Co.  C u l t . , USSR A r c t o -  OR,  1972  Ornduff, et a l . , 1967  Co.  A l p i n e Bot.  1966  Strother,  Missoula  cordifolia  Barker,  Schaack, et a l . , 1974 Ornduff, et a l . , 1967 O r n d u f f , et a l . , 1967 Love & Kapoor, Zhukova,  1968  1967  Gdn.  Josephine  Co.  Ornduff, et a l . , 1963  (as p a r v i f l o r a ) discoidea  38  CA, T r i n i t y  38  CA, Santa Cruz Co.  Ornduff, et a l . , 1967  discoidea  76  CA, Lake Co.  Ornduff, et a l . , 1963  discoidea  76  CA, Orange  Ornduff, et a l . , 1963  Co.  Barker,  1966  (as p a r v i f l o r a ) discoidea (as p a r v i f l o r a )  gracilis  call4  AB, Mt.  Co.  Temple  Love & Love,  1964  64  Table I I c o n t ' d . Taxon  gracilis latifolia  Sporophytic Number ca76 38  Locality  ID, V a l l e y Co.  Ornduff, e t a l . , 1967  BC, Queen C h a r l o t t e  T a y l o r & M u l l i g a n , 1968  Is., latifolia  38  Reference  Graham I s .  BC, Queen C h a r l o t t e  T a y l o r & M u l l i g a n , 1968  I s . , Moresby I s . latifolia  38  AB  latifolia  38  WA,  latifolia  38  WA, Whatcom  latifolia  38  WA,  latifolia  38  MT, R a v a l l i  latifolia  ca76  MT, Lake Co.  P o w e l l , et a l . , 1974  spathulata  ca76  OR, Josephine Co.  B a r k e r , 1966  spathulata  38  CA, Del Norte Co.  Ornduff, et a l . , 1967  venosa  38  CA, Shasta Co.  B a r k e r , 1966  viscosa  38  CA, S i s k i y o u Co.  B a r k e r , 1966  viscosa  38  CA, T r i n i t y  S t r o t h e r , 1973  Ornduff, e t a l . , 1967 K i t t i t a s Co. Co.  Lewis Co. Co.  Co.  Ornduff, e t a l . , 1963 Ornduff, e t a l . , 1963 B a r k e r , 1966 Schaack, e t a l . , 1974  65  TABLE I I I .  New Chromosome Numbers f o r A r n i c a Austromontana and C a l a r n i c a .  Taxon  Sporophytic Number  cernua  38  S p e c i e s i n the Subgenera  Locality  C o l l e c t i o n '.  CA, Humboldt Co.  S-1396  Horse Mtn. cernua  38  CA, Humboldt Co.  S-1772  west o f Hoopa cordifolia  76  YT,  cordifolia  76  BC, B o t a n i e  Valley  S-1342  cordifolia  76  BC, B o t a n i e  Valley  S-1353  cordifolia  76  BC, F o u n t a i n  cordifolia  76  BC, Bonanza Pass  S-1549  cordifolia  57  BC, southeast of  S-1594  Rancheria  S-1853  Valley  S-1360  Windermere cordifolia  57  BC, Tod Mtn.  S-1652  cordifolia  76  BC, Hat  S-2009  cordifolia  76  BC, Sunday Summit  S-2014  cordifolia  57  BC, O l a l l a  S-2022  cordifolia  76  BC, Apex Mountain  S-2025  cordifolia  76  BC, n o r t h o f Kamloops  S-2028  76  WA,  S-1494  cordifolia  i  Creek  Okanogan Co.,  S l a t e Lake cordifolia  114  WA,  Okanogan Co.,  S-1497  S l a t e Lake cordifolia  76  WA,  Okanogan Co.,  S l a t e Lake  S-1498  66  Table I I I cont'd. Taxon  cordifolia  Sporophytic Number  76  Locality  WA,  C o l l e c t i o n Number  K l i c k i t a t Co. ,  S-1822  north of L y l e cordifolia  76  WA,  Klickitat  Co.,  S-1823  Appleton cordifolia  76  OR, Wasco Co.,  S-1819  Pine Grove cordifolia  76  OR, Wasco Co.,  S-1820  P i n e Grove cordifolia  76  CA, S i e r r a Co.,  S-1416  Calpine cordifolia  76  CA, P l a c e r Co.,  S-1425  Lake Tahoe cordifolia  76  CA, S i s k i y o u Co.,  S-1797  west of Meiss Lake cordifolia  76  CA, Mono Co. ,  S-1937  Saddlebag Lake cordifolia  57  CA, Mono Co.,  S-1939  Saddlebag Lake cordifolia  114  cordifolia  AB, V e r m i l l i o n Pass  S-1601  76  AB, Lake L o u i s e  S-1604  cordifolia  76  AB, Moraine Lake  S-1610  cordifolia  76  AB, Peyto Lake  S-1616  cordifolia  76  ID, B o n n e v i l l e Co.,  S-1836  west o f Aspen, WY  67  T a b l e . I l l cont'd. Taxon  cordifolia  Sporophytic Number  Locality  76  MT, M i s s o u l a Co.,  C o l l e c t i o n '.  S-1477  P a t t e e Canyon cordifolia  57  WY,  Teton Co.,  S-1464  Grand Teton Nat. Park cordifolia  57  WY,  Teton Co.,  S-1465  F l a g g Ranch cordifolia  76  WY,  Park Co., Y e l l o w s t o n e  R i v e r , Upper cordifolia  76  cordifolia  76  Falls  SD, Lawrence Co., Deadwood  S-1837  S-1842  Gulch  CO, Routt Co.,  S-1437  Rabbit Ears Pass cordifolia  76  CO, Routt Co.,  S-1438  Rabbit E a r s Pass cordifolia  76  CO, Grand  Co.,  S-1440  Co.,  S-1443  Muddy Pass cordifolia  76  CO, Grand Winter  cordifolia  57  CO, Grand Grand  cordifolia  76  Park  76  S-2035  Lake  CO, C l e a r Creek Co., Berthoud  cordifolia  Co.,  Pass  CO, B o u l d e r Co., Nederland  S-1458  S-1460  68  Table.Ill Taxon  cordifolia  cont'd. Sporophytic Number 76  Locality  C o l l e c t i o n Number  MI, Keweenaw Co.,  S-1846  Copper Harbor cordifolia  76  MI, Keweenaw Co.  S-2005  cordifolia  57  C u l t . , BC, UBC B o t . Gdn.,  S-1512  o r i g . from BC, Kootenay D i s t . cordifolia  57  C u l t . , BC, UBC B o t . Gdn.  S-1513  discoidea  76  WA,  Co., Glenwood  S-1825  discoidea  76  OR, Wasco Co., Warm S p r i n g s  S-1821  Klickitat  Indian discoidea  38  CA, Mendocino Co., Little  discoidea  38  Reservation  River  CA, Napa Co., south o f Napa-Lake  discoidea  38  S-1775  S-1787  Co. l i n e  CA, Marin Co.,  S-1789  Mt. Tamalpias gracilis  57  BC, Monroe Lake  S-1562  gracilis  57  AB, Lake L o u i s e  S-1603  gracilis  57  AB, Moraine Lake  S-1605  gracilis  57  AB, Peyto Lake  S-1613  gracilis  57  C u l t . , BC, UBC B o t . Gdn.  S-1762  latifolia  38  BC, Mt. Seymour  S-1503  latifolia  38  BC, Mt. Seymour  S-1504  latifolia  38  BC, Vancouver I s l a n d  S-1529  Forbidden  Plateau  6.9  Table I I I c o n t ' d . Taxon  latifolia  Sporophytic Number 38  Locality  Collection 1  BC, Vancouver I s l a n d  S-1538  Forbidden P l a t e a u latifolia  38  BC, Bonanza Pass  S-1548  latifolia  38  BC, Kootenay Pass  S-1561  latifolia  38  BC, Manning Park  S-1680  latifolia  38  BC, Nancy Greene Prov. Pk.  S-1961  latifolia  38  BC, B l a c k Tusk  S-2031  latifolia  38  BC, Cypress Bowl  S-2041  latifolia  38  WA, Whatcom Co.,  S-1161  Mt. Baker latifolia  38  WA,  Chelan Co.,  S-1499  Washington Pass latifolia  38  WA,  Chelan Co.,  Rainy  S-1500  Pass  latifolia  38  AB, Peyto Lake  S-1612  latifolia  38  AB, south o f Grand P r a i r i e  S-1855  latifolia  38  MT, M i s s o u l a Co.,  S-1476  P a t t e e Canyon nevadensis  76  CA, S i e r r a Co.,  S-1870  S i e r r a Buttes nevadensis  76  CA, Mono Co., Saddlebag Lk.  S-1938  nevadensis  76  NV, Washoe Co., Mt. Rose  S-1910  spathulata  38  OR, Josephine Co., K i r b y  S-1380  spathulata  38  OR, Josephine Co., e a s t o f  S-1381  Cave J u n c t i o n  70  Table I I I cont'd. Taxon  spathulata  Sporophytic Number 38  Locality  Collection  OR, J o s e p h i n e Co.,  S-1385  O'Brien spathulata  38  OR, J o s e p h i n e Co., Selma  S-1740  spathulata  38  OR, J o s e p h i n e Co., n o r t h  S-1742  of spathulata  38  O'Brien  CA, D e l Norte Co.,  S-1390  northwest of Gasquet spathulata  38  CA, D e l Norte Co., e a s t of  spathulata  38  S-1751  Gasquet  CA, D e l Norte Co.,  S-1392  northwest o f Gasquet venosa  38  CA, Shasta Co.,  S-1791  Shasta B a l l y venosa  38  CA, Shasta Co., South Fork  S-1792  Lookout Mtn. Road venosa  38  CA, Shasta Co., Gibson  S-1793  viscosa  38  OR, C r a t e r Lake Nat. Park,  S-1946  G a r f i e l d Peak viscosa  38  CA, S i s k i y o u  Co., Mt. Shasta  S-1411  P u t a t i v e Natural Hybrids cordifolia-  38  latifolia cordifolialatifolia  MT, R a v a l l i Co., C h a r l e s  S-1474  Waters Campground 38  MT, R a v a l l i Co., C h a r l e s Waters Campground  S-1475  Number  71  F i g u r e 195.  A r n i c a cernua S-1772, CA, Humboldt Co.  Microsporocyte,  Metaphase I p l a t e , d i p l o i d w i t h 19 p a i r s o f chromosomes. F i g u r e 196.  A r n i c a l a t i f o l i a S-1529, BC, Vancouver I s l a n d , Forbidden Plateau.  M i c r o s p o r o c y t e , metaphase I p l a t e , d i p l o i d  19 p a i r s o f chromosomes.  with  72  73  F i g u r e 197.  A r n i c a c o r d i f o l i a S-1819, OR, Wasco County.  Microsporocyte,  l a t e anaphase I , t e t r a p l o i d w i t h 38 chromosomes a t each p o l e . F i g u r e 198.  A r n i c a c o r d i f o l i a S-1601, AB, V e r m i l l i o n Pass.  Micro-  s p o r o c y t e , metaphase I p l a t e , h e x a p l o i d w i t h v a r i a b l e u n i v a l e n t s , b i v a l e n t s , and t r i v a l e n t s .  74  ®  75  F i g u r e 199.  A r n i c a c o r d i f o l i a X A r n i c a l a t i f o l i a S-1475, p u t a t i v e n a t u r a l h y b r i d , MT, R a v a l l i diakinesis,  F i g u r e 200.  diploid  County.  Microsporocyte,  w i t h 19 p a i r s of chromosomes.  A r n i c a c o r d i f o l i a S-1594, BC, E. of Windermere. Microsporocyte,  l a t e anaphase I , t r i p l o i d w i t h  30  chromosomes a t one p o l e o f c e l l and 27 a t the o t h e r .  76  77  F i g u r e 201.  A r n i c a g r a c i l i s S-1613, AB,  Peyto Lake.  Microsporocyte,  metaphase I p l a t e , t r i p l o i d , w i t h v a r i a b l e u n i v a l e n t s , b i v a l e n t s , and F i g u r e 202.  trivalents.  A r n i c a n e v a d e n s i s S-1870, CA, Microsporocyte,  S i e r r a County.  anaphase I , t e t r a p l o i d , w i t h 76  chromosomes.  79  Figure  203.  Arnica  spathulata  S-1392, OR, Josephine County.  M i c r o s p o r o c y t e , metaphase I p l a t e , d i p l o i d w i t h 19 p a i r s of chromosomes. Figure  204.  Arnica  discoidea  S-1787, CA, Napa County.  Microsporocyte,  metaphase I p l a t e , d i p l o i d w i t h 19 p a i r s o f chromosomes. Figure  205.  Arnica discoidea  S-1821, OR, Wasco County.  Microsporocyte,  metaphase I p l a t e , t e t r a p l o i d w i t h 38 p a i r s of chromosomes.  80  81  F i g u r e 206.  A r n i c a v i s c o s a S-1411, CA, S i s k i y o u County.  Microsporocyte,  d i a k i n e s i s , d i p l o i d w i t h 19 p a i r s o f chromosomes. F i g u r e 207.  A r n i c a venosa S-1793, CA, Shasta County.  Microsporocyte,  metaphase I p a l t e , d i p l o i d w i t h 19 p a i r s of chromosomes.  82  83  produced  r e g a r d l e s s of the presence  o r absence of a v i a b l e seed w i t h i n .  These empty achenes are o f t e n as l a r g e as v i a b l e achenes and appear to be v i a b l e u n t i l examined c l o s e l y or u n t i l A c c o r d i n g to Barker  the achene coat i s removed.  (1966) the two  common types of apomixis  found i n  Arnica are: Autonomous apomixis  - Seed p r o d u c t i o n without p o l l i n a t i o n o r  fertilization. Pseudogamous apomixis p o l a r n u c l e i are f e r t i l i z e d  - Seed p r o d u c t i o n does not occur u n l e s s to produce endosperm.  the  Embryo development i s  parthenogenetic. In used  a d d i t i o n to the bagging  to determine  experiments,  autonomous apomicts.  two  e m a s c u l a t i o n methods were  In r a d i a t e p o p u l a t i o n s d i s c  florets  were c a r e f u l l y removed from heads b e f o r e the d i s c s opened and b e f o r e the r a y s had  elongated  ( F i g u r e 209) .  The  head was  then bagged, l e a v i n g the  ray  f l o r e t s to mature i n the absence of p o l l e n .  pistillate  On o t h e r heads, e i t h e r  r a d i a t e o r d i s c o i d , a t the same stage of development, the whole top of head from j u s t above the ovary was  the  removed w i t h a r a z o r b l a d e , thus removing  a l l a n t h e r s , stigmas, and the upper p o r t i o n o f the s t y l e s . then bagged and a l l o w e d t o mature.  Although  The heads were  severe, t h i s treatment  seems to  have no d e t r i m e n t a l e f f e c t s on v i a b l e achene p r o d u c t i o n i n autonomous apomicts. Table  The r e s u l t s of these bagging  and  e m a s c u l a t i o n procedures  a r e shown i n  IV. The b r e e d i n g systems of the n i n e s p e c i e s a r e : Sexual ( s e l f - i n c o m p a t i b l e ) A r n i c a cernua  (Barker,  1966)  A r n i c a d i s c o i d e a p o p u l a t i o n s ( B a r k e r , 1966,  as A r n i c a p a r v i f l o r a )  84  F i g u r e s 208-211. F i g u r e 208.  Experimental  methods.  Arnicas i n cultivation  i n experimental  f i e l d , UBC B o t a n i c a l  Garden N u r s e r y . F i g u r e 209.  Head of A r n i c a c o r d i f o l i a S-1497, WA, disc florets  F i g u r e 210.  removed.  Growth Chamber w i t h p l a n t s i n f l o w e r , s e e d l i n g s i n p o t s , and seeds germinating  F i g u r e 211.  Okanogan County, w i t h  in petri  A r n i c a head bagged.  dishes.  85  TABLE IV. Taxon & C o l l . //  Compatibility Diploid Chromosome //  and Apomixis Experiments w i t h A r n i c a  species.  it of Heads Selfed  // of Heads Emasculated  it of Achenes Produced  // of Achenes With Seeds  69%  4  -  111  5  4.5 17.2  Pollen Stainability  % of Ac With S.  cordifolia  1342  76  cordifolia  1343  1  89  4  -  221  38  cordifolia  1343  1  89  -  3  101  0  0  cordifolia  1353  76  83  2  -  112  7  6.2  cordifolia  1353  76  83  -  2  55  8  14.5  cordifolia  1359  1  38  2  -  96  13  13.5  cordifolia  1359  1  38  -  1  7  0  0  cordifolia  1360  76  42  4  -  157  2  1.3  cordifolia  1416  76  24  3  -  272  67  24.6  cordifolia  1458  76  82  1  -  189  0  0  cordifolia  1458  76  82  -  1  71  0  0  cordifolia  1477  76  32  6  -  371  24  cordifolia  1477  76  32  -  2  46  0  0  cordifolia  1487  ?  49  1  -  52  9  17.3  cordifolia  1494  76  94  3  -  218  0  0  cordifolia  1494  76  94  -  2  52  0  0  cordifolia  1497  114  84  8  —  485  100  6.5  20.6  T a b l e IV cont'd • Taxon & Coll. #  Diploid Chromosome #  Pollen Stainability  // of Heads Selfed  // of Heads Emasculated  // o f Achenes Produced  // o f Achenes With Seeds  % of Achenes With Seeds  c o r d i f o l i a 2005  76  81  3  -  314  4  1.3  c o r d i f o l i a 2005  76  81  -  1  7  0  0  d i s c o i d e a 1821  76  76  1  -  49  0  0  d i s c o i d e a 1825  76  77  4  -  129  15  d i s c o i d e a 1825  76  77  -  1  21  0  gracilis  1562  57  0  10  -  222  187  84.2  g r a c i l i s .1562  57  0  -  4  72  50  69.4  g r a c i l i s '. 1605  57  3  4  -  93  64  68.8  gracilis  57  3  -  2  54  18  33.8  l a t i f o l i a 1499  38  87  1  -  19  0  0  latifolia  1499  38  87  -  1  36  0  0  l a t i f o l i a 1503  38  97  10  -  301  0  0  l a t i f o l i a 1504  38  97  2  -  69  0  0  l a t i f o l i a 1504  38  97  -  3  178  0  0  latifolia  1561  38  75  2  -  57  0  0  l a t i f o l i a 1961  38  90  1  _  28  0  0  1605  •  11.6 0  T a b l e IV c o n t ' d . Taxon & C o l l . //  Diploid Chromosome #  Pollen Stainability  # of Heads Selfed  // of Heads Emasculated  # of Achenes Produced  // o f Achenes With Seeds  % of Achenes W i t h Seeds  l a t i f o l i a 1961  38  90  -  1  8  0  n e v a d e n s i s 1870  76  43  2  -  122  59  48.4  n e v a d e n s i s 1938  76  3  3  -  155  65  41.9  n e v a d e n s i s 1938  76  3  -  3  74  23  31.1  s p a t h u l a t a 1380  38  99  3  -  113  0  0  venosa 1940  38  100  4  108  0  0  v i s c o s a 1411  38  100  20  -  302  0  0  12  -  1570  0  0  20  -  2051  0  0  0  Hybrids c o r d . - l a t . 1474  38  c o r d . - l a t . 1475  38  7  7  34  89 Arnica  latifolia  Arnica spathulata  populations  A r n i c a venosa Arnica viscosa Autonomous Apomicts Arnica c o r d i f o l i a  populations  Arnica g r a c i l i s Arnica  nevadensis  Pseudogamous Apomicts Arnica c o r d i f o l i a  populations  Arnica discoidea populations Arnica spathulata populations  Pollen  1966)  Stainability  For p o l l e n s t a i n a b i l i t y one  (Barker,  or more f l o r e t s  specimen was  t e s t s , contents  of f r e s h l y d e h i s c e d anthers  of a p l a n t from a w i l d - c o l l e c t e d p o p u l a t i o n or herbarium  tapped out on a g l a s s s l i d e .  P o l l e n g r a i n s were s t a i n e d w i t h  drop of c o t t o n b l u e - l a c t o p h e n o l and a cover s l i p was  added.  of  scored a f t e r 24  a 200-grain  random sample f o r each p o p u l a t i o n was  Large, well-formed t h a t d i d not (Figure The 212-221. few  of  The  stainability  g r a i n s s t a i n e d dark b l u e were s c o r e d as "good".  s t a i n or were s m a l l and misshapened were scored as  a  hours. Those  "bad"  191). range of p o l l e n s t a i n a b i l i t y  f o r each s p e c i e s i s shown i n F i g u r e s  S e v e r a l of the s p e c i e s a r e so r a r e i n n a t u r e and  specimens c o u l d be Barker  i n herbaria that  sampled.  (1966) d i s c u s s e d the c o r r e l a t i o n between p l o i d y l e v e l ,  and p o l l e n s t a i n a b i l i t y .  The  data p r e s e n t e d  here support  apomixis,  his findings.  90  There a r e some d i s t i n c t t r e n d s among c e r t a i n s p e c i e s . predominately cernua,  The e x c l u s i v e l y o r  d i p l o i d , sexual s p e c i e s i n c l u d i n g A r n i c a l a t i f o l i a ,  Arnica  A r n i c a s p a t h u l a t a , A r n i c a venosa, and A r n i c a v i s c o s a a l l show v e r y  h i g h p o l l e n s t a i n a b i l i t y , mostly  g r e a t e r than 90%.  A second group o f s p e c i e s i n c l u d e s A r n i c a g r a c i l i s and A r n i c a which have mostly  l e s s than 10% s t a i n a b l e p o l l e n .  or presumed to be autonomous apomicts and m o s t l y  nevadensis  These two a r e known to be  or e n t i r e l y p o l y p l o i d s .  A  number o f p o p u l a t i o n s of A r n i c a g r a c i l i s sampled had no p o l l e n produced i n any of  florets.  I n known t r i p l o i d  p o p u l a t i o n s of A r n i c a g r a c i l i s t h e p r o d u c t i o n  s m a l l and u n s t a i n a b l e normal s i z e d p o l l e n g r a i n s ( F i g u r e s 191, 192) can be  f o l l o w e d from i r r e g u l a r p a i r i n g o f chromosomes to the p r o d u c t i o n o f more f o u r groups o f chromosomes a f t e r anaphase I I , the normal t e t r a d stage 189).  (Figure  W a l l s o f t e n form around these " q u i n t a d s " o r " s e x t a d s " r e s u l t i n g i n  micro p o l l e n g r a i n s and normal s i z e d g r a i n s ( F i g u r e s 187-192). large grains apparently lack a f u l l to  than  develop  Even  these  complement o f chromosomes and o f t e n  fail  into a viable, or at least stainable, pollen grain.  A t h i r d group i n c l u d e s A r n i c a c o r d i f o l i a and the c l o s e l y r e l a t e d A r n i c a d i s c o i d e a i n which p o l l e n s t a i n a b i l i t y seem to i n d i c a t e a n y t h i n g  i s e r r a t i c from 0-100% and does not  s p e c i f i c a l l y about the p l o i d y l e v e l o r  apomixis.  In A r n i c a d i s c o i d e a t h e r e a r e both d i p l o i d p o p u l a t i o n s w i t h h i g h p o l l e n s t a i n a b i l i t y and t e t r a p l o i d p o p u l a t i o n s w i t h lower p o l l e n s t a i n a b i l i t y .  In A r n i c a  c o r d i f o l i a no d e f i n i t e s e x u a l p o p u l a t i o n s a r e known and f o r t h e known p o l y p l o i d populations pollen s t a i n a b i l i t y  i s variable.  The b e s t assumption f o r  A r n i c a c o r d i f o l i a i s that lower p o l l e n s t a i n a b i l i t y p r o b a b l y  represents  nomous a p o m i c t i c p o p u l a t i o n s and the h i g h e r p o l l e n s t a i n a b i l i t y pseudogamous a p o m i c t i c p o p u l a t i o n s .  auto-  represents  The few d i p l o i d p o p u l a t i o n s which a r e  91  jures 212-221.  Range of p o l l e n s t a i n a b i l i t y sampled.  f o r Arnica  populations  cordifolia  .At:.  t • • • • ••••••• ••••••••••  •••• •  Number of populations  latifolia  ••••• •••••••• •••••  cordifolia-latifolia hybrids  10  20  ^0  "  40  % Pollen Stainability  50  60  @ )  70  80  90  100  nevadensis  •••• •  Number of populations  cernua  1  -i  (216  r-  ^  r-  grac ilis  : :.  »»  M 1  •  r  10  r-  ~l  ^  "I  1—  T  ©  (  r-  20  -i  r-  30  40  % Pollen Stainability  50  -i  60  70  80  1  r-  90  100  discoidea  T  99  1-  Q  ©  QQ  • ••  spathulata  e  e  ~1  •  r-  ••••••• •  @ )  •  Number of populations  •  •• ••••  venosa  -r—  1  r-  -i  r-  -i  r-  -i  1  —i  r-  1  1  viscosa  10  20  30  40  < Pollen Stainability  '  50  ~i  60  70  80  1—  90  100  95  known t o e x i s t may be s e x u a l , although  they have n o t been i n v e s t i g a t e d  experimentally.  Seed  Production  Seed p r o d u c t i o n both even i n a p o m i c t i c r a c e s .  i n n a t u r e and i n c u l t i v a t i o n i s o f t e n q u i t e low Although  s e v e r a l dozen to a hundred o r more achenes  are produced i n each head, many o f these a r e empty. of  A l s o , i n many p o p u l a t i o n s  most o r a l l s p e c i e s the heads a r e i n f e s t e d w i t h one to s e v e r a l l a r v a e o f  achene-feeding  d i p t e r a n s which o f t e n f e e d on most o r a l l o f t h e achenes i n a  head b e f o r e the pappus expands and the seeds a r e d i s p e r s e d .  (See f u r t h e r  d i s c u s s i o n under E n t o m o l o g i c a l R e l a t i o n s h i p s ) . Seed p r o d u c t i o n i n c u l t i v a t i o n where t h e r e were no d i p t e r a n i n f e s t a t i o n was s t i l l lis  low except  i n the autonomous a p o m i c t i c p o p u l a t i o n s o f A r n i c a g r a c i -  and A r n i c a n e v a d e n s i s .  Of achenes examined from 21 heads o f A r n i c a  g r a c i l i s 54-100% (mostly g r e a t e r than E i g h t heads o f A r n i c a nevadensis achenes.  80%) o f the achenes c o n t a i n e d  had 0-70% (mostly g r e a t e r than 60%) f u l l  In t h e s e x u a l p o p u l a t i o n s o f A r n i c a l a t i f o l i a  than 20%) o f achenes from 25 heads examined c o n t a i n e d l e c t e d achenes o f A r n i c a cernua have been observed, from 5 heads from one p o p u l a t i o n .  seed.  0-72% (mostly  seeds.  less  Only w i l d  w i t h 6-14% f u l l  col-  achenes  Of the s e x u a l , s e l f - i n c o m p a t i b l e p o p u l a -  t i o n s o f A r n i c a s p a t h u l a t a , A r n i c a v i s c o s a , and A r n i c a venosa, o n l y one each has been grown s u c c e s s f u l l y to the f l o w e r i n g and f r u i t i n g no  d a t a o f seed p r o d u c t i o n a r e a v a i l a b l e .  in  the w i l d had 47% f u l l There i s poor seed  stage.  Therefore,  One head o f A r n i c a venosa  collected  achenes. s e t i n c u l t i v a t i o n o f the predominately  a p o m i c t i c p o p u l a t i o n s of A r n i c a c o r d i f o l i a and A r n i c a d i s c o i d e a .  pseudogamous Of more  than  96  50 heads o f A r n i c a c o r d i f o l i a examined, seed p r o d u c t i o n was 0-64% (mostly l e s s than 30%) o f achene p r o d u c t i o n . achenes.  W i l d c o l l e c t e d achenes had 0-29% f u l l  I n A r n i c a d i s c o i d e a 0-20% o f t h e achenes from 16 heads examined  were  full.  Seed  Germination  Young s e e d l i n g s i n t h e i r a l p i n e o r montaine environment d i s a d v a n t a g e i f t h e achenes germinated  immediately  would be a t a  i n l a t e summer o r f a l l  a f t e r f a l l i n g o r b e i n g c a r r i e d from t h e parent p l a n t .  I t was suspected t h a t  the achenes needed a v e r n a l i z a t i o n o r s c a r i f i c a t i o n of the achene coat b e f o r e the seeds would germinate. spring  Germination p r o b a b l y o c c u r s i n n a t u r e a f t e r the  snow-melt.  Achenes c o n t a i n i n g seeds were s u b j e c t e d to numerous schemes i n an e f f o r t to  induce g e r m i n a t i o n , i n c l u d i n g the f o l l o w i n g : 1.  P l a n t e d i n p o t s of s o i l o u t - o f - d o o r s i n t h e f a l l and a l l o w e d to  o v e r w i n t e r under " n a t u r a l " ( i . e . , Vancouver w i n t e r ) 2.  conditions.  S t o r e d d r y or i n moist sphagnum i n t h e r e f r i g e r a t o r  (0°C) or d r y a t room temperature  (4°C) o r f r e e z e r  ( c a 20°C) f o r s i x weeks to s e v e r a l months  before p l a n t i n g . 3. filter  Achenes p l a c e d immediately a f t e r d i s p e r s a l o r a f t e r s t o r a g e on m o i s t paper i n p e t r i d i s h e s i n l i g h t  (16 hours d a y l i g h t ) a t 25°C.  (Figure  210) . 4.  Same as #3 except t o t a l  darkness.  5.  Achene c o a t s c a r e f u l l y removed a f t e r they had been soaked i n  d i s t i l l e d water f o r 24 hours, then p l a c e d on m o i s t f i l t e r d a y l i g h t a t 25°C.  paper  i n 16 hours  97  The r e s u l t s o f these v a r i o u s t e s t s have been mixed and t h e r e a r e s t i l l some unanswered q u e s t i o n s c o n c e r n i n g seed g e r m i n a t i o n .  The g r e a t e s t success  was had w i t h soaked achenes, w i t h achene coats removed, and then p l a c i n g t h e seeds on moist  f i l t e r paper under l o n g days a t 25°C.  seed germinate  on moist f i l t e r paper b e f o r e the achene coat was removed.  A f t e r removal,  t h e h y p o c o t y l o f t e n began to e l o n g a t e w i t h i n 12 hours.  Using  In no cases d i d any  the above method t h e r e was n e a r l y 100% g e r m i n a t i o n o f seeds from  some p o p u l a t i o n s of some s p e c i e s , e s p e c i a l l y A r n i c a c o r d i f o l i a , A r n i c a g r a c i l i s , and A r n i c a v i s c o s a .  With o t h e r p o p u l a t i o n s of the same s p e c i e s o r o t h e r  species, e s p e c i a l l y A r n i c a c o r d i f o l i a , A r n i c a nevadensis, A r n i c a d i s c o i d e a , and A r n i c a s p a t h u l a t a , none o f t h e seeds germinated method.  w i t h t h i s , o r any other  In some cases a few days a f t e r t h e achene coat was removed, the  c o t y l e d o n s r e f l e x e d , but t h e r e was no f u r t h e r s i g n o f growth, even a f t e r months.  I n y e t o t h e r p o p u l a t i o n s of A r n i c a c o r d i f o l i a , and e s p e c i a l l y A r n i c a  venosa, a few seeds germinated  immediately,  w h i l e o t h e r s remained plump and  a p p a r e n t l y capable o f g e r m i n a t i o n f o r up to s i x months, y e t t h e r e were no s i g n s o f h y p o c o t y l or e p i c o t y l e l o n g a t i o n .  O f t e n d u r i n g the l o n g e r t e s t s the  d i s h e s became i n f e s t e d w i t h mold o r a l g a e , so t h a t the seeds had to be d i s carded . V e r n a l i z a t i o n i s n o t needed f o r g e r m i n a t i o n .  Under a r t i f i c i a l  g e r m i n a t i o n was n o t enhanced by c o o l i n g o r f r e e z i n g t h e achenes.  conditions  However,  s c a r i f i c a t i o n of t h e achene c o a t s seems to be e s s e n t i a l f o r g e r m i n a t i o n . Achenes p l a n t e d d i r e c t l y i n s o i l spring.  o u t - o f - d o o r s d i d n o t germinate  The achenes were v e r y d i f f i c u l t  to l o c a t e to determine  were v i a b l e o r i f they had decomposed d u r i n g t h e w i n t e r .  the f o l l o w i n g i f they  still  98  Seedlings  Of more than 200 p o p u l a t i o n s recognized  in this  Seedlings planted  grown from seeds germinated i n the growth chamber were t r a n s standard  s t a r t e d to d e v e l o p .  p o t t i n g medium as soon as the f i r s t  young s e e d l i n g s was r e l a t i v e l y s e e d l i n g s were v e r y  pair  These were grown i n t h e growth chamber  under the same growing c o n d i t i o n s as f o r g e r m i n a t i o n .  of l e a v e s .  i n c l u d i n g a l l taxa  s t u d y , no s e e d l i n g s were o b s e r v e d .  into a s t e r i l i z e d  of t r u e l e a v e s  studied i n the f i e l d ,  high f o r the f i r s t  M o r t a l i t y rate of the  few weeks.  Most h e a l t h y  slow growing, t a k i n g 1-3 months to develop t h r e e p a i r s  Seedlings  of the serpentine  p o t t i n g medium and i n s e r p e n t i n e  endemic A r n i c a cernua grown both i n  s o i l grew e q u a l l y s l o w l y ,  the b a s a l  reaching  o n l y 2-3 cm i n d i a m e t e r and w i t h  months.  O t h e r s , e s p e c i a l l y A r n i c a c o r d i f o l i a , A r n i c a v i s c o s a , and A r n i c a  venosa grew q u i c k l y a f t e r month ( F i g u r e 278) . and  3-4 p a i r s o f l e a v e s  after  rosette  the t h i r d o r f o u r t h month and f l o w e r e d  6-8  by t h e s i x t h  One s e e d l i n g o f A r n i c a v i s c o s a had become much branched  produced more than 100 f l o w e r s  by the e i g h t h month.  l a t i f o l i a and A r n i c a g r a c i l i s produced robust growth w i t h i n 6 months but no f l o w e r i n g  Seedlings  of A r n i c a  b a s a l r o s e t t e s and some rhizome  stems were produced  Two-month o l d s e e d l i n g s grown i n a growth chamber a r e shown i n F i g u r e s 222-227.  Even when q u i t e young  (by t h e f i r s t  the s e e d l i n g s assume t h e t y p i c a l c h a r a c t e r s leaf  p a i r of true leaves)  o f t h e more mature p l a n t , i n  shape, r o o t s , and e s p e c i a l l y i n pubescence, a l t h o u g h the s e e d l i n g s a r e  o f t e n more pubescent when young than they a r e l a t e r .  A f t e r a few weeks t h e  Austromontana s p e c i e s remain as a b a s a l r o s e t t e w i t h no stem while  many o f  the C a l a r n i c a s p e c i e s  elongation,  form a d e f i n i t e stem by the time the second o r  t h i r d p a i r o f t r u e l e a v e s have d e v e l o p e d .  99  F i g u r e s 222-227.  A r n i c a S e e d l i n g s ca 2 months a f t e r  germination,  grown i n growth chamber. F i g u r e 222.  A r n i c a c o r d i f o l i a S-1497, WA,  Okanogan County.  F i g u r e 223.  A r n i c a l a t i f o l i a S-1855, AB, south of Grande  F i g u r e 224.  A r n i c a g r a c i l i s S-1562, BC, Monroe Lake.  F i g u r e 225.  A r n i c a cernua S-1772, OR, J o s e p h i n e County.  F i g u r e 226.  A r n i c a v i s c o s a S-1411, CA, S i s k i y o u County.  F i g u r e 227.  A r n i c a venosa S-1940, CA, Shasta County.  Prairie.  100  101  Vegetative  Reproduction  Considering  the low seed p r o d u c t i o n , i n f e s t a t i o n s o f achene f e e d i n g  D i p t e r a , and poor g e r m i n a t i o n in  o f seeds, v e g e t a t i v e r e p r o d u c t i o n by rhizomes  s p e c i e s of subgenus Austromontana i s c r i t i c a l f o r t h e i r s u c c e s s f u l c o l o n i -  zation. Little Arnicas.  o r no i n f o r m a t i o n e x i s t s on v e g e t a t i v e r e p r o d u c t i o n p o t e n t i a l i n  Herbarium specimens a r e o f l i t t l e v a l u e .  n i c e herbarium specimens ( i . e . , sheet)  Although  A r n i c a s make  they f i t c o n v e n i e n t l y on a standard  and they a r e f r e q u e n t l y c o l l e c t e d and a r e w e l l - r e p r e s e n t e d  i n herbaria,  p l a n t s a r e u s u a l l y p u l l e d from t h e ground r a t h e r than dug, r e s u l t i n g rhizomes being l e f t behind.  herbarium  i n the  Specimens w i t h rhizomes u s u a l l y show o n l y one  season's growth o r a p o r t i o n t h e r e o f .  No known p r e v i o u s l y c o l l e c t e d s p e c i -  mens of t h e two s p e c i e s i n subgenus C a l a r n i c a had enough o f the underground p a r t s present  t o determine i f they were o r were not rhizomatous.  Records o f annual rhizome growth have been kept on p o p u l a t i o n s of a l l s p e c i e s seen i n the f i e l d of  some p o p u l a t i o n s  i n a d d i t i o n to s e v e r a l years o f r e c o r d s of growth  in cultivation  ( F i g u r e s 228-231, Table V ) .  have been supplemented by measurements from herbarium specimens. stances t h e average growth i s lower than the median. ing  These s t u d i e s In most i n -  A few p o p u l a t i o n s  grow-  i n r e l a t i v e l y dense shade o r i n v e r y l o o s e s o i l s tend to have t h e ex-  tremely  long rhizomes.  Examples a r e the low e l e v a t i o n p o p u l a t i o n s o f A r n i c a  g r a c i l i s from f o r e s t s a t Monroe Lake, BC ( S t r a l e y 1562) w i t h rhizomes up to 6 cm l o n g , whereas a m a j o r i t y o f the p o p u l a t i o n s as r e p r e s e n t e d by those  from  h i g h a l p i n e rock s c r e e s l o p e s o f B a n f f and J a s p e r N a t i o n a l P a r k s , AB ( S t r a l e y 1603,  1605, 1613) average o n l y 1-2 cm o f growth per year.  102  F i g u r e 228.  Rhizome growth measured  S-1512, UBC B o t a n i c a l Garden, N a t i v e Kootenay D i s t r i c t , BC.  over f o u r y e a r s , A r n i c a Garden.  cordifolia  Plant o r i g i n a l l y  from  20cm  Original Plant-1976 Growth -1977 1978 1979  104  F i g u r e 229. S-1162, WA,  Rhizome  growth measured over 4 y e a r s , A r n i c a  Whatcom County, Mt. Baker.  latifolia  105  106  F i g u r e 230.  Rhizome growth measured o v e r f o u r y e a r s , A r n i c a cernua  S-1772, CA, Humboldt  County, Hoopa Mountain.  2cm  Original Plant - 1975 Growth -1976 1977 1978  108  F i g u r e 231.  Rhizome growth measured  S-1605, AB, Moraine Lake.  over 3 y e a r s , A r n i c a  gracilis  109  • - Original Plant -1977 O - 1978 A  - 1979  110  TABLE V.  Y e a r l y Rhizome Growth i n A r n i c a , Subgenus Austromontana  Taxon  Annual Growth (cm)  cernua  (l-)4-15(-20)  6  cordifolia  2-20(-60)  10  discoidea  5-15(-22)  Annual Average (cm)  Breeding System  Species.  Average Seed V i a b i l i t y  Sexual  14%  Apomictic  30%  7  Sexual and Apomictic  20%  Apomictic  80%  gracilis  (0.5-)l-2(-6)  1.5  latifolia  (0.5-)l-7(-10)  4  Sexual  20%  nevadensis  (l-)3-12  6  Apomictic  60%  spathulata  (1-)3-7(-14)  5  Sexual and Apomictic  ?  Ill  There seems to be no d i s t i n c t and  c o r r e l a t i o n between rhizome  s u c c e s s f u l seed s e t o r between the rhizome p r o d u c t i o n  s e x u a l compared t o a p o m i c t i c  species.  production  o f predominately  However, there a r e some c o n s i s t e n t  d i f f e r e n c e s between average rhizome growth of some o f t h e c l o s e l y species.  related  The average growth i n A r n i c a c o r d i f o l i a , f o r i n s t a n c e , i s 2.5 times  that of Arnica l a t i f o l i a  and t h e growth o f A r n i c a g r a c i l i s  i s l e s s than h a l f  as g r e a t as t h a t of A r n i c a l a t i f o l i a .  The d e r i v e d s p e c i e s  ( A r n i c a cernua and  A r n i c a g r a c i l i s from A r n i c a l a t i f o l i a ,  and A r n i c a d i s c o i d e a , A r n i c a  spathulata  and A r n i c a n e v a d e n s i s from A r n i c a c o r d i f o l i a ) a l l have lower rhizome product i o n than do t h e i r presumed a n c e s t r a l s p e c i e s . S e v e r a l o f these s p e c i e s f r e q u e n t l y invade l a n d s l i d e s , burned-over and  other disturbed h a b i t a t s .  P a r t i c u l a r l y i n Arnica c o r d i f o l i a the y e a r l y  rhizome growth r a t e s shown here c o u l d e a s i l y e x p l a i n the v a s t a r e a by a s i n g l e p o p u l a t i o n one  areas,  coverage o f an  a f t e r hundreds o r thousands of y e a r s , from o n l y  s u c c e s s f u l achene i n t r o d u c t i o n .  CULTIVATION OF ARNICA SPECIES  Arnica populations in  have been t r a n s p l a n t e d from n a t u r e o r grown from seed  the growth chamber, greenhouse, and experimental  field  a t UBC w i t h  mixed  results. Some s p e c i e s , e s p e c i a l l y A r n i c a c o r d i f o l i a , A r n i c a n e v a d e n s i s , and A r n i c a g r a c i l i s have been most e a s i l y t r a n s p l a n t e d , w i t h near 100% s u c c e s s , most p o p u l a t i o n s  flowering f r e e l y after transplanting.  Arnica  A r n i c a d i s c o i d e a , and A r n i c a v i s c o s a have given mixed r e s u l t s . tions transplanted  e a s i l y and f l o w e r e d  and w i t h  spathulata, Some p o p u l a -  seasonally f o r several years.  Others  e i t h e r d i d n o t s u r v i v e o r have l i v e d f o r o n l y a few months and f l o w e r e d  poorly  112  or not  at a l l .  The  widespread A r n i c a l a t i f o l i a u s u a l l y s u r v i v e s moving, but  flowers very poorly i n future years, rhizomes i n c u l t i v a t i o n .  The  successfully transplanted. the experimental flowered  a l t h o u g h i t reproduces abundantly  serpentine  A few  endemic A r n i c a cernua has  A few  s e e d l i n g s have grown very s o i l - l e s s p o t t i n g mixture.  achenes have germinated and  slowly both i n serpentine Two  not  been  p l a n t s have s u r v i v e d f o r a year or two  garden but have become p r o g r e s s i v e l y s m a l l e r and  in cultivation.  by  the  s o i l s and  have never  resulting  in a  p l a n t s from d i f f e r e n t p o p u l a t i o n s  A r n i c a venosa have been u n s u c c e s s f u l l y t r a n s p l a n t e d  from the w i l d .  standard of the  q u i c k l y and  transplanted  flowered  In a l l cases,  cases,  on  to a s o i l - l e s s p o t t i n g mixture have grown  freely in cultivation.  the tops d i e d back to the rhizomes when p l a n t s were  from the w i l d w h i l e  rare  However,  u n l i k e A r n i c a cernua, s e e d l i n g s of A r n i c a venosa from achenes germinated f i l t e r paper and  in  i n flower or j u s t before or a f t e r .  t h e r e were new  sprouts  dug  However, i n most  produced from the rhizomes w i t h i n a few weeks  of t r a n s p l a n t i n g , o f t e n producing  f l o w e r i n g stems s e v e r a l weeks l a t e r .  HYBRIDIZATION  Sympatric Occurrence and  Natural Hybridization  There i s v e r y l i t t l e evidence of r e c e n t h y b r i d i z a t i o n o c c u r r i n g i n n a t u r e between any other  of the s p e c i e s i n Austromontana and  subgenera.  There are a number of p o p u l a t i o n s  c h a r a c t e r s between A r n i c a c o r d i f o l i a and w i t h one  C a l a r n i c a , or w i t h  or e i t h e r parent.  exhibiting  Arnica l a t i f o l i a ,  These are c o n s i d e r e d  species i n  intermediate  u s u a l l y not  growing  old s t a b i l i z e d hybrids,  a l l y apomicts, and w i l l be d i s c u s s e d under A r n i c a c o r d i f o l i a .  usu-  113  Because o f d i f f e r e n t e c o l o g i c a l p r e f e r e n c e s , most of t h e n i n e i n these two  subgenera do not occur s y m p a t r i c a l l y .  Where two  species  or more s p e c i e s  do grow i n c l o s e p r o x i m i t y o r a t l e a s t c l o s e enough f o r p o l l e n exchange by i n s e c t p o l l i n a t o r s (see Table V I ) , they o f t e n have l i t t l e o r no o v e r l a p i n f l o w e r i n g times.  T h i s was  observed  i n p o p u l a t i o n s of A r n i c a d i s c o i d e a and  A r n i c a c o r d i f o l i a growing s y m p a t r i c a l l y i n Wasco County, OR 1821)  and K l i c k i t a t  County, WA  ( S t r a l e y 1825,  l a t i o n s the A r n i c a c o r d i f o l i a was was  j u s t beginning  to f l o w e r .  s p a t h u l a t a ( S t r a l e y 1385, of 1977  and  1978.  i n Josephine  j u s t beginning  County, OR was  to f l o w e r .  together  i n Del Norte County, CA  mention of any  evidence  of h y b r i d i z a t i o n .  In most of the mixed p o p u l a t i o n s  ally  observed  These two (Klipfel  Arnica  in  May  the  s p e c i e s have a l s o  770,  HSC),  with  no  there are a l s o d i f f e r e n c e s i n p l o i d y  systems, and v i a b l e p o l l e n p r o d u c t i o n , a c t i n g s i n g l y o r usu-  i n combination  as b a r r i e r s to n a t u r a l h y b r i d i z a t i o n .  A specimen from Custer County, ID ( C r o n q u i s t 3375, UTC)  represents  apparent h y b r i d p o p u l a t i o n growing w i t h both p a r e n t s , A r n i c a g r a c i l i s Arnica l o n g i f o l i a . t a l types.  popu-  and  j u s t f i n i s h i n g f l o w e r i n g as  been c o l l e c t e d  l e v e l s , breeding  In both of these  A mixed p o p u l a t i o n o f A r n i c a cernua  1386)  1820,  f i n i s h i n g f l o w e r i n g as the A r n i c a d i s c o i d e a  The A r n i c a cernua was  A r n i c a s p a t h u l a t a was  1826).  (Straley  The  specimen i s q u i t e i n t e r m e d i a t e between the two  an  and paren-  T h i s i s the o n l y herbarium specimen seen of what appears to be  h y b r i d p o p u l a t i o n between two (Austromontana and  distinct  Chamissonis).  s p e c i e s i n two  different  subgenera  a  114  TABLE V I .  Sympatric p o p u l a t i o n s o f A r n i c a subgenus Austromontana s p e c i e s w i t h no evidence o f n a t u r a l h y b r i d s .  Collection Number  Locality  Pollen Stainability  Diploid Chromosome Number  Breeding System  s p a t h u l a t a S -1385  OR, Josephine Co.  100%  38  sexual  s.-1386  OR, Josephine Co.  98%  -  sexual  c o r d i f o l i a S--1573  BC, Yahk Mountain  30%  -  apomictic  latifolia  s--1576  BC, Yahk Mountain  100%  -  sexual  gracilis  S--1603  AB, Lake L o u i s e  0%  57  apomictic  c o r d i f o l i a S--1604  AB, Lake L o u i s e  52%  76  apomictic  latifolia  S--1612  AB, Peyto Lake  99%  38  sexual  gracilis  S--1613  AB, Peyto Lake  2%  57  apomictic  c o r d i f o l i a S--1616  AB, Peyto Lake  32%  76  apomictic  c o r d i f o l i a S--1653  BC, Tod Mountain  74%  apomictic  latifolia  S--1654  BC, Tod Mountain  95%  -  c o r d i f o l i a S--1820  OR, Wasco County  85%  76  apomictic  discoidea  s--1821  OR, Wasco County  76%  76  apomictic  discoidea  s--1825  WA, K l i c k i t a t Co.  77%  76  apomictic  cordifolia  s--1826  WA, K l i c k i t a t Co.  80%  -  apomictic  nevadensis  s--1938  CA, Mono County  3%  76  apomictic  cordifolia  s--1938  CA, Mono County  8%  57  apomictic  cernua  sexual  115 A r t i f i c i a l Hybridization  R e c i p r o c a l c r o s s e s between a l l the s p e c i e s i n these two subgenera a r e complicated  by s e v e r a l f a c t o r s .  The d i f f i c u l t y  o r p o p u l a t i o n s has been p r e v i o u s l y d i s c u s s e d . c u r r e n t l y i n c u l t i v a t i o n i s o f t e n a problem.  i n c u l t i v a t i o n of some s p e c i e s Getting plants to flower  I n the e x p e r i m e n t a l  con-  garden under  n a t u r a l s p r i n g c o n d i t i o n s , most s p e c i e s produced o n l y one s e t o f f l o w e r i n g stems and t h e f l o w e r i n g d u r a t i o n was q u i t e b r i e f , e s p e c i a l l y i n A r n i c a l a t i folia.  While most of the montane and a l p i n e s p e c i e s f l o w e r a t the same time,  A r n i c a v i s c o s a takes  s e v e r a l months f o r t h e f l o w e r stems to e l o n g a t e ,  f l o w e r i n g much l a t e r than most o t h e r s p e c i e s .  Under c o n t r o l l e d  thus  conditions of  the growth chamber A r n i c a v i s c o s a , A r n i c a venosa, and some p o p u l a t i o n s o f o t h e r s p e c i e s c o u l d be kept f l o w e r i n g f o r many months, e n a b l i n g  experimental  c r o s s e s w i t h these and o t h e r more r a p i d l y f l o w e r i n g s p e c i e s . In r a d i a t e forms, the r a y f l o r e t s may be used as the female p a r e n t i n c r o s s e s , but t h e r e c i p r o c a l c r o s s e s cannot be made u s i n g t h e d i s c o i d w i t h t h e i r complete f l o r e t s as the female p a r e n t , without d i s c o i d form i s s e l f - i n c o m p a t i b l e .  Emasculation  f i r s t proving the  of the s m a l l d i s c o i d  b e f o r e a n t h e s i s i s t e d i o u s and o f t e n r e s u l t s i n damage to t h e stigma of  t h e whole  forms  florets or loss  floret.  P o l y p l o i d y and apomixis a r e f u r t h e r c o m p l i c a t i o n s , w i t h  sexual  p o p u l a t i o n s n o t known i n s e v e r a l s p e c i e s , e s p e c i a l l y A r n i c a c o r d i f o l i a , Arnica nevadensis,  and A r n i c a g r a c i l i s .  used as t h e female parent fertility The  Not o n l y can these s p e c i e s n o t be  i n r e c i p r o c a l c r o s s e s , but w i t h v e r y low p o l l e n  i n many o f them, they cannot be used as a p o l l e n source f o r c r o s s e s . r e s u l t s o f a l l r e c i p r o c a l c r o s s e s a r e g i v e n i n T a b l e V I I and  summarized i n F i g u r e 232.  TABLE V I I . A r t i f i c i a l H y b r i d i z a t i o n s Using Sexual P o p u l a t i o n s (known or  supposed) of A r n i c a S p e c i e s .  Taxon & Collection # ( t f Parent)  Achenes Produced  Diploid Taxon & Chromosome // C o l l e c t i o n // ((^ Parent)  Diploid Chromosome #  Heads Crossed  Achenes With Seeds  l a t i f o l i a 1499  38  c o r d i f o l i a 1497  76  3  9  0  latifolia  1504  38  c o r d i f o l i a 1512  57  2  13  7  latifolia  1961  38  c o r d i f o l i a 2009  76  2  64  0  latifolia  1961  38  d i s c o i d e a 1821  76  1  56  0  l a t i f o l i a 1961  38  d i v e r s i f o l i a 1419  76  3  118  3  l a t i f o l i a 1961  38  g r a c i l i s 1605  57  1  28  0  l a t i f o l i a 1961  38  v i s c o s a 1411  38  1  53  0  l a t i f o l i a 1503  38  m o l l i s 1515  ?  1  8  0  latifolia  38  m o l l i s 1515  1  1  9  0  s p a t h u l a t a 1380  38  c o r d i f o l i a 1477  76  2  69  3  s p a t h u l a t a 1380  38  c o r d i f o l i a 1497  76  O z.  37  0  venosa 1940  38  c o r d i f o l i a 2005  76  1  53  0  venosa 1940  38  c o r d . - l a t . hyb. 1475  38  1  25  0  venosa 1940  38  l a t i f o l i a 2041  38  1  26  0  venosa 1940  38  nevadensis 1938  76  1  26  0  venosa 1940  38  v i s c o s a 1411  38  1  27  0  1504  Table VII cont'd. Taxon & Collection # (0* Parent)  Diploid Chromosome  v i s c o s a 1411  38  c o r d i f o l i a 1343  v i s c o s a 1411  38  v i s c o s a 1411  Taxon & C o l l e c t i o n // (0_ Parent)  Diploid Chromosome #  Heads Crossed  Achenes Produced  ?  2  34  5  c o r d i f o l i a 2005  76  1  22  0  38  l a t i f o l i a 1680  38  1  16  0  v i s c o s a 1411  38  l a t i f o l i a 1961  38  1  19  5  v i s c o s a 1411  38  l a t i f o l i a 2041  38  1  13  2  v i s c o s a 1411  38  c o r d . - l a t . hyb. 1474  38  2  45  0  v i s c o s a 1411  38  c o r d . - l a t . hyb. 1475  38  3  55  0  v i s c o s a 1411  38  d i s c o i d e a 1825  76  5  110  9  v i s c o s a 1411  38  nevadensis 1870  76  2  41  0  v i s c o s a 1411  38  venosa  38  2  36  2  c o r d . - l a t . 1474  38  s p a t h u l a t a 1380  38  1  14  0  cord. - l a t .  1474  38  v i s c o s a 1411  38  1  17  0  cord. - l a t .  1475  38  c o r d i f o l i a 1353  76  1  61  0  cord. - l a t .  1475  38  d i s c o i d e a 1825  76  2  39  0  cord. - l a t .  1475  38  s p a t h u l a t a 1380  38  1  5  0  cord.'- l a t .  1475  38  v i s c o s a 1411  38  3  40  0  1940  Achenes W i t h Seeds  Hybrids  118  F i g u r e 232.  Summary of a l l a r t i f i c i a l  c r o s s e s attempted between  i n subgenus Austromontana and C a l a r n i c a . crosses.  Dotted l i n e s , u n s u c c e s s f u l  Solid  crosses.  species  lines indicate successful  119  120  Many of t h e s e e d l i n g s from c r o s s e s were v e r y weak and d i d n o t s u r v i v e beyond a few days o r weeks, o r d i d n o t s u r v i v e the t r a n s p l a n t i n g from dishes  to s o i l .  As o f t h i s w r i t i n g s e v e r a l s e e d l i n g s a r e s t i l l v e r y much about t h e i r c h a r a c t e r s except  q u i t e young.  The l e a v e s a r e d e n s e l y  flowered.  Seedlings of  intermediate vegetative  characters  g l a n d u l a r as i n A r n i c a v i s c o s a ,  but w i t h toothed margins as i n A r n i c a venosa. between A r n i c a l a t i f o l i a  too young to determine  some v e g e t a t i v e ones.  A r n i c a v i s c o s a X A r n i c a venosa do e x h i b i t while  petri  One s e e d l i n g from a c r o s s  and A r n i c a v i s c o s a has grown r a p i d l y and has  I t i s intermediate  i n most r e s p e c t s .  The l e a v e s a r e l a r g e , b u t  n e a r l y s e s s i l e and t h e r e a r e some g l a n d u l a r h a i r s and the odor o f A r n i c a v i s c o s a , y e t greener  than  t h i s parent.  The few heads were narrow b u t b r i g h t  y e l l o w and w i t h a few s h o r t , t w i s t e d r a y f l o r e t s . the achenes have a few g l a n d u l a r  The pappus i s tawny and  hairs.  ENTOMOLOGICAL INTERRELATIONSHIPS  Although of  many o f t h e A r n i c a s have showy heads and a r e a conspicuous  the montane-alpine-arctic  f l o r a , very l i t t l e  has been r e c o r d e d  part  on p o l l i n a -  t i o n o r any o t h e r p l a n t - i n s e c t r e l a t i o n s h i p s , o t h e r than work done by Griffiths  (1974) on h o s t s p e c i f i c Phytomyza f l y l e a f m i n e r s .  Kunth (1908) has compiled  r e c o r d s o f a few i n s e c t v i s i t o r s  to A r n i c a  montana (subgenus Montana), and some N o r t h American s p e c i e s c u l t i v a t e d t a n i c a l gardens i n Europe. "bees" o r " f l i e s " ,  etc.  Many o f these p o l l i n a t o r s were l i s t e d  o n l y as  He r e c o r d s A r n i c a montana as b e i n g autogamous o r  geitonogamous ( p o l l i n a t e d by a d j a c e n t f i n d i n g s of Barker  i n bo-  florets  i n the same h e a d ) , a l t h o u g h t h e  (1966) and i n t h i s c u r r e n t work suggest  that a l l sexual  121  p o p u l a t i o n s of A r n i c a s a r e p r o b a b l y s e l f - i n c o m p a t i b l e .  Leaf  Miners  Griffiths  (1974) r e c o r d s  seven s p e c i e s of Phytomyza  which form mines i n the l e a v e s of A r n i c a s p e c i e s . these are c o n f i n e d to the genus A r n i c a . s p e c i f i c Phytomyza.  With o n l y one  exception a l l  However, t h e r e seems to be no  species-  Some o f the Phytomyza s p e c i e s can be i d e n t i f i e d from  p a t t e r n of the mine, as they t u n n e l below the During  (Diptera:Agromyzidae)  epidermis.  t h i s study mines were seen on a l l s p e c i e s in.subgenus Austromon-  t a n a , a l t h o u g h none have been o b s e r v e d  on e i t h e r s p e c i e s of C a l a r n i c a .  In  most i n s t a n c e s the mines occupy h a l f or l e s s of the l e a f by the time the vae  pupate.  There a r e some s e v e r e  i n f e s t a t i o n s observed  l e a v e s on the p l a n t s have t h e whole s u r f a c e mined by All  the  a r e shown i n F i g u r e s 233-240. A r n i c a cernua  No  i n which most of  s e v e r a l m i n e r s per  c o l l e c t i o n s i d e n t i f i e d a r e g i v e n i n T a b l e V I I I , and  larthe  leaf.  r e p r e s e n t a t i v e mines  a d u l t f l i e s have been b r e d  o r A r n i c a s p a t h u l a t a , so the s p e c i e s i n f e s t i n g  from  either  those A r n i c a s  are  n o t known.  Floret  Feeders  There a r e o c c a s i o n a l l a r g e h o s t - s p e c i f i c o r omnivorous i n c l u d i n g i n c h worms (Geometriidae) Guenee ( N o c t u i d a e ) , which may  herbivores  or cutworms, e s p e c i a l l y Autographa  devour l e a v e s , heads and  florets.  a l s o a number of s m a l l e r h e r b i v o r e s which f e e d on A r n i c a f l o r e t s . h a l f of the p o p u l a t i o n s o b s e r v e d s p e c i e s had  one  i n the f i e l d o f b o t h r a d i a t e and  o r many s m a l l b e e t l e s ( B u p r e s t i d a e , m o s t l y  A n t h a x i a ) f e e d i n g on  the r a y and/or d i s c f l o r e t s  simplex  There are At  least  discoid  i n the genus  (Table IX).  In some extreme  122  TABLE V I I I .  Phytomyza  A r n i c a specimens c o l l e c t e d  oreas G r i f . o r Phytomyza  A r n i c a c o r d i f o l i a S-1472, MT, Arnica  w i t h l e a f mines o f Phytomyza  arhicivora  Sehgal  Lewis & C l a r k County  c o r d i f o l i a S-1553, BC, Nancy Green P r o v i n c i a l  Park  A r n i c a c o r d i f o l i a S-1604, AB, Lake L o u i s e A r n i c a c o r d i f o l i a S-1616, AB, Peyto Lake A r n i c a c o r d i f o l i a S-1819, OR, Wasco County A r n i c a c o r d i f o l i a S-1826, WA,  K l i c k i t a t County  A r n i c a c o r d i f o l i a Bohm 1109, BC, Arnica  Osoyoos  c o r d i f o l i a Guppy, _s.n_., BC, B o t a n i e V a l l e y  Arnica l a t i f o l i a  S-1676, WA,  Arnica l a t i f o l i a  S-1680, BC, Manning Park  Clallam  A r n i c a n e v a d e n s i s S-1870, CA, S i e r r a  Phytomyza  County  saximontana G r i f .  Arnica l a t i f o l i a  Phytomyza  County  S-1538, BC, Vancouver I s l a n d , Forbidden P l a t e a u  species  A r n i c a cernua S-1396, CA, Humboldt Arnica  s p a t h u l a t a S-1385, OR,  County  J o s e p h i n e County  species.  123  F i g u r e s 233-240.  Leaf Mines o f Phytomyza  Species i n leaves of A r n i c a .  F i g u r e s 233-237.  Mines o f P. o r e a s o r P. a r n i c i v o r a .  F i g u r e 233.  Arnica l a t i f o l i a  S-1680, BC, Manning Park.  F i g u r e 234.  Arnica l a t i f o l i a  S-1676, WA,  F i g u r e 235.  A r n i c a c o r d i f o l i a S-1819, OR, Wasco County.  F i g u r e 236.  A r n i c a c o r d i f o l i a S-1604, AB, Lake L o u i s e .  F i g u r e 237.  A r n i c a nevadensis S-1870, CA, S i e r r a County.  F i g u r e 238.  Mine of P_. saximontana on A r n i c a l a t i f o l i a Vancouver I s l a n d , Forbidden  F i g u r e 239.  S-1538, BC,  Plateau.  U n i d e n t i f i e d b l o t c h mine on A r n i c a cernua S-1396, CA, Humboldt  F i g u r e 240.  C l a l l a m County.  County.  U n i d e n t i f i e d b l o t c h mine on A r n i c a s p a t h u l a t a S-1385, OR, Josephine County.  124  125  TABLE IX.  B u p r e s t i d B e e t l e s C o l l e c t e d F e e d i n g on A r n i c a S p e c i e s .  A r n i c a c o r d i f o l i a S-1594, BC, Windermere A n t h a x i a expansa L e c . - 2 A r n i c a c o r d i f o l i a S-1819, OR, Wasco County, P i n e  Grove  A n t h a x i a expansa L e c . - 1 A n t h a x i a aenescens Casey - 1 A r n i c a c o r d i f o l i a S-2022, BC, Apex Mountain A n t h a x i a expansa L e c . - 2 A r n i c a c o r d i f o l i a S-1842, SC, Lawrence  County, Deadwood  Gulch  A n t h a x i a expansa L e c . - 7 A r n i c a c o r d i f o l i a S-1797, CA, S i s k i y o u  County, Meiss Lake  A n t h a x i a sp. - 2 A r n i c a s p a t h u l a t a S-1381, OR, J o s e p h i n e Acmaeodera  County, Cave J u n c t i o n  connexa L e c . - 1  Anthaxia C a l i f o r n i a  Obenb. -1  A r n i c a venosa S-1793, CA, S h a s t a County, Gibson Anthaxia C a l i f o r n i a A n t h a x i a sp. - 1  Obenb. - 8  126  cases,  as i n A r n i c a  County, CO,  c o r d i f o l i a near Windermere, BC,  S-2035, the i n f e s t a t i o n was  S-1594, and  i n Grand  so severe t h a t a l l the r a y s had  eaten, g i v i n g the p l a n t s a c l o s e s u p e r f i c i a l resemblance to A r n i c a Severe i n f e s t a t i o n s c o u l d a f f e c t p o l l i n a t o r v i s i t a t i o n due reflectance-absorbance The  Buprestid  from one  patterns  b e e t l e s are mostly too s m a l l and  p l a n t to another, thus are not  discoidea.  t o changed  (see f u r t h e r d i s c u s s i o n under UV  been  UV  section).  g l a b r o u s to t r a n s f e r p o l l e n  l i k e l y aiding i n pollination.  Achene Feeders  As species  discussed  under "Seed P r o d u c t i o n "  many p o p u l a t i o n s  (observed i n a l l s p e c i e s except A r n i c a v i s c o s a )  l a r v a e of one The  of most or a l l  are  infested with  o r more s m a l l f l i e s of the genus Xenochaeta ( T e p h r i t i d a e ) .  young l a r v a e b e g i n f e e d i n g on  b e g i n to expand.  The  the o v a r i e s b e f o r e  l a r v a e have u s u a l l y pupated by  mature, occupying the remains of s e v e r a l achenes.  the f i r s t  florets  the time the achenes  The  remaining achenes are undamaged.  pappus on the v i a b l e achenes i s damanged to the extent The  uninfested  r a t h e r than being Frequently  achenes may  open  unit  the pappus blown i n the wind.  the l a r v a e of the t e p h r i t i d s are  l a r v a e of Pteromalus wasps ( P t e r o m a l i d a e ) ,  the  t h a t i t does not  drop to the ground as a s o l i d  c a r r i e d away i n d i v i d u a l l y by  of  In more severe i n f e s t a t i o n s ,  which o c c u r commonly, most or a l l o f the achenes are e i t h e r eaten o r  properly.  are  I n minor i n f e s t a t i o n s the  l a r v a e u s u a l l y f e e d upon o n l y h a l f a dozen o r fewer achenes i n the c e n t e r the head.  the  themselves i n f e s t e d w i t h  so t h a t the a d u l t wasps, r a t h e r  than a d u l t t e p h r i t i d s , emerge from the pupal c a s e s . u s u a l l y i n f e s t e d enough t h a t they are not v i a b l e .  The  achenes are  still  127  U l t r a v i o l e t R e f l e c t a n c e - A b s o r b a n c e o f A r n i c a Heads  Photographs o f A r n i c a heads were taken o u t - o f - d o o r s on a sunny day w i t h a  tripod-mounted M i n o l t a s i n g l e - l e n s e r e f l e x camera u s i n g Kodak T r i - X Pan  b l a c k and w h i t e f i l m  (ASA 400).  At the same d i s t a n c e exposures were taken  w i t h a Wratten 18A f i l t e r a t t a c h e d and the camera stopped-down 3-4 f - s t o p s to compensate f o r the lower l i g h t p e n e t r a t i o n through the opaque f i l t e r . Wratten 18A f i l t e r through. the is  a l l o w s o n l y UV l i g h t  The  i n the 350-400 nm range to pass  A r n i c a heads a r e l a r g e enough t h a t the v a r i a t i o n i n f o c a l l e n g t h o f  l e n s w i t h and without the f i l t e r seen a t t h i s d i s t a n c e  ( c a 30 cm).  i s not c r i t i c a l ,  a l t h o u g h some d i s t o r t i o n  At d i s t a n c e s c l o s e r than t h i s the d i f -  f e r e n c e i n f o c a l l e n g t h must be compensated f o r ( H i l l ,  1977;  Kennedy and  Ganders, 1979). The UV r e f l e c t a n c e - a b s o r b a n c e p a t t e r n i n A r n i c a heads, which appear c o m p l e t e l y y e l l o w to the human eye ( F i g u r e 241), appear l i k e those o f Monoptilon bellioides  (Kennedy and Ganders, 1979) i n which t h e d i s c f l o r e t s  UV l i g h t and appear dark, w h i l e t h e r a y f l o r e t s r e f l e c t light  ( F i g u r e 242).  absorb  UV and appear s o l i d  T h i s i s u n l i k e the p a t t e r n i n many other genera o f t h e  A s t e r a c e a e i n which p a r t o f the r a y f l o r e t s r e f l e c t UV and an i n n e r p o r t i o n of  the r a y absorbs UV as i n Rudbeckia spp. (Abrahamson and McCrea, 1977),  L a s t h e n i a chrysostoma (Kennedy and Ganders, 1979), o r C o r e o p s i s g i g a n t e a (Scogin et a l . , In  1977).  A r n i c a , a f t e r t h e heads have been i n f l o w e r a few days and a l l the  d i s c s have opened and have p r o b a b l y been p o l l i n a t e d , t h e rays b e g i n to absorb UV and t h e whole head appears dark ( F i g u r e 242). i n UV p a t t e r n has been noted by S c o g i n e t a l . and  This p o s t - p o l l i n a t i o n  change  (1977) f o r C o r e o p s i s g i g a n t e a  a s i m i l a r change was noted i n o l d e r herbarium specimens o f t h a t  species.  128  F i g u r e s 241-247. Heads o f Three F i g u r e 241.  UV R e f l e c t a n c e - A b s o r b a n c e i n A r n i c a heads and T y p i c a l  Species.  Arnica c o r d i f o l i a X Arnica l a t i f o l i a  (putative hybrid)  S-1474, MT, R i v a l l i County, photographed i n n a t u r a l sunlight. F i g u r e 242.  Same photographed w i t h Wratten 18A f i l t e r showing d a r k , a b s o r p t i v e UV d i s c s and l i g h t , r e f l e c t i v e UV rays i n two r e c e n t l y opened heads on l e f t  and r i g h t ,  and dark,  absorptive  d i s c s and r a y s o f two p o s t - p o l l i n a t i o n c e n t e r heads. F i g u r e 243.  A r n i c a d i s c o i d e a S-1825, WA, K l i c k i t a t County, photographed in natural sunlight.  F i g u r e 244.  Same photographed w i t h UV f i l t e r showing UV a b s o r p t i v e florets  F i g u r e 245.  (dark).  A r n i c a c o r d i f o l i a S-1497, WA, Okanogan County, v e r y wide-rayed  F i g u r e 246.  population.  Arnica l a t i f o l i a  S-1961, BC, Nancy Greene P r o v i n c i a l Park,  t y p i c a l head w i t h " s q u a r e d - o f f " F i g u r e 247.  disc  rays.  A r n i c a g r a c i l i s S-1603, AB, Lake L o u i s e , t y p i c a l w i t h few, narrow r a y s .  heads  129  130  In the d i s c o i d ( F i g u r e s 243,  s p e c i e s of A r n i c a the head absorbs UV  and  appears dark  244).  Pollinators  An unexpedtedly s m a l l number of i n s e c t p o l l i n a t o r s has been v i s i t i n g A r n i c a p o p u l a t i o n s i n these two  observed  subgenera, a l t h o u g h many hours have  been spent o b s e r v i n g hundreds of p o p u l a t i o n s at a l l hours of the day  through-  out most of the d i s t r i b u t i o n of the s p e c i e s .  popula-  In many of the l a r g e s t  t i o n s w i t h hundreds or thousands of heads i n f l o w e r no p o l l i n a t o r s were observed.  There are s e v e r a l p o s s i b i l i t i e s f o r the l a c k of p o l l i n a t o r s i n  some p o p u l a t i o n s . t h e r e may  In f o r e s t h a b i t a t s where the shade i s r e l a t i v e l y dense  be fewer i n s e c t s than i n the more open h a b i t a t s .  which are l a r g e l y monocephalous, most of the heads may of one  another.  Few  or no p o l l i n a t o r s may  p o l l i n a t e d because of the changing UV  visit  In  populations  open w i t h i n a few  days  a f t e r the heads have been  reflectance-absorbance  patterns.  In  a l p i n e meadows, where t h e r e are many d i f f e r e n t p l a n t s f l o w e r i n g at the same time,  t h e r e may  pollinators.  be a p r e f e r e n c e f o r o t h e r genera or s p e c i e s of f l o w e r s by  Many b u t t e r f l i e s and bees have been observed  f l o w e r s ( i . e . , V a l e r i a n a spp., Lupinus spp., to A r n i c a spp. AB,  In a few  visiting  and Eriogonum spp.)  i n s t a n c e s as on Yahk Mountain, BC,  the  other  i n preference  and Moraine Lake,  t h e r e seemed to be a p o l l i n a t o r p r e f e r e n c e f o r A r n i c a s p e c i e s i n o t h e r  subgenera.  Euphydryas spp.  v i s i t e d Arnica rydbergii,  (Nymphalidae) and H e s p e r i a  spp.  (Hesperiidae)  A r n i c a m o l l i s , and A r n i c a d i v e r s i f o l i a , but  rarely  v i s i t e d A r n i c a l a t i f o l i a , A r n i c a c o r d i f o l i a or A r n i c a g r a c i l i s .  At s l i g h t l y  lower e l e v a t i o n s on Yahk Mountain and near the town of Yahk, BC,  Arnica  f o l i a was  commonly v i s i t e d by s e v e r a l s p e c i e s of Bombus.  These  two  lati-  131  p o p u l a t i o n s a r e t h e o n l y A r n i c a s i n these of i n s e c t p o l l i n a t o r s were  two subgenera where l a r g e numbers  observed.  There seems to be no c o r r e l a t i o n between v i s i t o r s  to a p o m i c t i c  compared  to s e x u a l p o p u l a t i o n s of A r n i c a s or between d i s c o i d and r a d i a t e s p e c i e s , a l though the UV p a t t e r n s a r e d i f f e r e n t i n the two groups. Table X l i s t s  the D i p t e r a , Hymenoptera, and L e p i d o p t e r a  c a p t u r e d on A r n i c a heads.  pollinators  Nomenclature of b u t t e r f l y p o l l i n a t o r s f o l l o w s Howe  (1975), t h a t of t h e o t h e r o r d e r s f o l l o w s B o r r o r e t a l . (1976) above the generic  level.  132  TABLE X.  P o l l i n a t o r s C o l l e c t e d on A r n i c a Heads.  A r n i c a cernua S-1772, CA, Humboldt  County, Hoopa Mountain.  Syrphidae sp. Arnica c o r d i f o l i a  S-1464, WY,  Teton County, Grand Teton N a t i o n a l Park  Lycaenidae M i t o u r a spinetorum (Hewitson) 0_ A r n i c a c o r d i f o l i a S-1648, BC, S h e r i d a n Lake Nymphalidae S p e y e r i a e l e c t a b e a n i (Barnes & Benjamin) 0_ Arnica c o r d i f o l i a  S-1797, CA, S i s k i y o u County, Meiss Lake  Syrphidae Taxomerus  o c c i d e n t a l i s Curran £  A r n i c a c o r d i f o l i a S-1819, OR, Wasco County, P i n e Grove Syrphidae Syrphus o p i n a t o r Osten Sachen 0  71  A r n i c a c o r d i f o l i a S-1823, WA,  Klickitat  County, A p p l e t o n  Syrphidae Sphaerophoria s u l p h u r i p e s (Thomson) 0_ Arnica c o r d i f o l i a  S-1826, WA,  Klickitat  County,  Glenwood  Andrenidae Andrena sp. o" Arnica cordifolia  S-1836, ID, B o n n e v i l l e County, Irwin  Apidae Bombus b i f a r i u s Cresson 0_ Andrenidae Andrena sp.  133  Table.X cont'd. A r n i c a c o r d i f o l i a S-2022, BC, Apex Mountain Halictidae H a l i c t u s sp. A r n i c a c o r d i f o l i a S-2028, BC, Kamloops Megachilidae Megachile sp. A r n i c a c o r d i f o l i a S-2029, BC, S o r r e n t o Syrphidae Eristalis  tenax (L.) ^  A r n i c a d i s c o i d e a S-1777, CA, Napa County,  S. o f Lake/Napa Co. l i n e  Hesperiidae Erynnis propertius Arnica l a t i f o l i a  (Scudder & Burgess) t f  S-1567, BC, Yahk  Apidae Bombus mixtus Cresson 4 0^ 7 \j (=worker c a s t ) Bombus b i f a r i u s  Cresson 2 \*j  P s i t h y r u s f e r n a l d a e F r a n k l i n 1 (f, 1 0_ Anthophoridae Nomada sp. 0_ Syrphidae Eristalis Arnica l a t i f o l i a  tenax (L.) cf S-1574, BC, Yahk Mountain  Apidae Bombus b i f a r i u s  Cresson ^  134  Table X cont'd. Arnica l a t i f o l i a  S-1576, BC, Yahk Mountain  Apidae Bombus melanopygus Nylander tj" Bombus mixtus Cresson 2 cf Nymphalidae Euphydryas Arnica l a t i f o l i a  sp. 5 0^ 2 J S-2031, BC, B l a c k Tusk  Sessidae Vespamina sp. ? A r n i c a s p a t h u l a t a S-1380, OR, Josephine County,  Kirby  Apidae A p i s m e l l i f e r a L. Halictidae H a l i c t u s sp. Arnica,  s p a t h u l a t a S-1740, OR, Josephine County, Apidae Bombus c a l i g i n o s u s ( F r i s o n ) 2 0^  A r n i c a venosa S-1793, CA, Shasta County, H a l i c t i d a e sp.  Gibson  Selma  135  SYSTEMATIC TREATMENT  Genus A r n i c a L. Sp. P l a n t . 884. 1753.  A r t i f i c i a l Key to the Subgenera o f A r n i c a . A.  Anthers p u r p l e ;  i n v o l u c r a l b r a c t s c a l l o u s - t i p p e d ; p l a n t s o f n. B r i t i s h  Columbia and Yukon to A l a s k a AA.  Anthers y e l l o w ; B.  and Japan...Subg. Andropurpurea Maguire  i n v o l u c r a l b r a c t s not c a l l o u s - t i p p e d  B  Leaves a l l s e s s i l e ; p l a n t s from woody caudex C.  Cauline leaves  s t r o n g l y b a s a l l y disposed  heads r a d i a t e ; p l a n t s o f se United Europe CC.  C on f l o w e r i n g  stem;  S t a t e s and c e n t r a l ...Subg. Montana Maguire  C a u l i n e l e a v e s not s t r o n g l y b a s a l l y d i s p o s e d ,  stem l e a f y  throughout; heads d i s c o i d ; r a r e p l a n t s o f sw Oregon and nw C a l i f o r n i a BB.  Subg. C a l a r n i c a S t r a l e y  Leaves mostly p e t i o l a t e ; p l a n t s rhizomatous, producing  basal  rosettes C.  C  Pappus brown o r tawny, subplumose Subg. Chamissonis Maguire  CC.  Pappus white ( r a r e l y tawny), b a r b e l l a t e D.  Leaf  blades  D  r e l a t i v e l y narrow, mostly 3-10 times as  long as wide; p l a n t s mostly densely  tufted  Subg. A r c t i c a Maguire DD.  Leaf blades  r e l a t i v e l y broad, mostly 1-2.5 times as l o n g  as broad; p l a n t s m o s t l y not densely  tufted  Subg. Austromontana Maguire  136  S p e c i e s Recognized  Subgenus Austromontana M a g u i r e . Lectotype: Sect.  Arnica l a t i f o l i a  B r i t t o n i a 4:432. Bongard  E u l a t i f o l i a e Maguire.  Lectotype:  B r i t t o n i a 4:432.  Arnica l a t i f o l i a  1943.  Bongard  1.  A r n i c a c o r d i f o l i a Hooker  2.  A r n i c a n e v a d e n s i s A. Gray  3.  Arnica l a t i f o l i a  4.  Arnica g r a c i l i s  5.  A r n i c a c e r n u a T. H o w e l l  Bongard Rydberg  S e c t . E r a d i a t a e Maguire. Lectotype:  1943.  B r i t t o n i a 4:452.  1943.  A r n i c a d i s c o i d e a Bentham  6.  A r n i c a d i s c o i d e a Bentham  7.  Arnica spathulata  Greene  Subgenus C a l a r n i c a subgenus nov. Type S p e c i e s : A r n i c a v i s c o s a A. Gray 8.  A r n i c a v i s c o s a A. Gray  9.  A r n i c a v e n o s a H.M.  Hall  The f o l l o w i n g keys a r e d e s i g n e d to be u s e f u l f o r a m a j o r i t y of these two subgenera. pressed  f o r b o t h l i v e and  specimens, and a f i e l d key.  Artificial A.  An a r t i f i c i a l key i s p r o v i d e d  o f specimens  Key to t h e S p e c i e s  o f A r n i c a , Subgenera Austromontana and C a l a r n i c a .  Leaves a l l s e s s i l e ; non-rhizomatous p e r e n n i a l s l a c k i n g s t e r i l e r o s e t t e s o f l e a v e s ; heads d i s c o i d B.  Margins o f l e a v e s  toothed;  (Subgenus  basal  Calarnica)  B  2 ° v e i n s prominent; l e a v e s r u g o s e ;  heads nodding i n bud; f l o r e t s b r i g h t y e l l o w ;  achenes  densely  137  s t r i g o s e , non-glandular  9. A r n i c a  venosa  o BB.  Margins  of l e a v e s e n t i r e ; 2  heads e r e c t i n bud;  v e i n s i n c o n s p i c u o u s ; l e a v e s smooth;  f l o r e t s creamy-yellow  o r white; achenes  s t i p i t a t e glandular AA.  8. A r n i c a v i s c o s a  Leaves, a t l e a s t lower ones, p e t i o l a t e ; rhizomatous p e r e n n i a l s p r o d u c i n g s t e r i l e b a s a l r o s e t t e s ; heads r a d i a t e or d i s c o i d  (Subgenus  Austromontana)  B  B. Heads r a d i a t e ( S e c t i o n E u l a t i f o l i a e ) C.  Leaves m o s t l y g l a b r o u s ( i f pubescent scabrous and n o n - g l a n d u l a r ) D.  Heads e r e c t i n bud; top;  then s p a r s e l y D  achenes h i r s u t e from middle to  l e a v e s t h i n ; v e i n s green; of widespread  t r i b u t i o n i n w. DD.  C  Heads nodding pubescent  dis-  N.A  3. A r n i c a  latifolia  i n bud;  achenes s p a r s e l y to d e n s e l y  throughout; l e a v e s t h i c k , l e a t h e r y , v e i n s  r e d d i s h ; r e s t r i c t e d to s e r p e n t i n e s o i l s of sw Oregon and nw CC.  California  5. A r n i c a  Leaves pubescent w i t h g l a n d u l a r and n o n - g l a n d u l a r  cernua hairs..  . D.  D  Pubescence o f l e a v e s g l a n d u l a r - p u b e r u l e n t o n l y ; p l a n t s mostly much-branched and cm t a l l ;  clumped; m o s t l y l e s s than 20  achenes b l a c k , g l a b r o u s to s p a r s e l y 4. A r n i c a  DD.  Pubescence o f l e a v e s g l a n d u l a r s t i p i t a t e and g l a n d u l a r ; p l a n t s m o s t l y unbranched g r e a t e r than 20 cm t a l l ;  pubescent. gracilis non-  and not clumped;  achenes g r e y , mostly d e n s e l y  h i r s u t e a t top o r throughout  E  138  E.  Leaves o f i n n o v a t i o n s and lower c a u l i n e l e a v e s w i t h c o r d i a t e bases; margins m o s t l y toothed; pure white  EE.  1. A r n i c a  pappus  cordifolia  Leaves of i n n o v a t i o n s and lower c a u l i n e l e a v e s w i t h a t t e n u a t e d o r rounded bases; margins m o s t l y e n t i r e ; pappus o f f - w h i t e or tawny 2. A r n i c a n e v a d e n s i s  BB .  Heads d i s c o i d C.  (Section Eradiatae)  Leaves of i n n o v a t i o n s and lower c a u l i n e l e a v e s p l a n t s mostly r e s t r i c t e d and nw  CC.  C spathulate;  to s e r p e n t i n e s o i l s of sw  California  7. A r n i c a s p a t h u l a t a  Leaves of i n n o v a t i o n s and lower c a u l i n e l e a v e s w i t h o r rounded bases; p l a n t s u s u a l l y not o f s e r p e n t i n e s. Washington  t o s. C a l i f o r n i a  cordate soils;  6. A r n i c a d i s c o i d e a  F i e l d Key to the S p e c i e s o f A r n i c a , subgenera Austromontana  A.  Oregon  and C a l a r n i c a .  Leaves d i s t i n c t l y f r a g r a n t when b r u i s e d B.  B  Heads d i s c o i d . C.  C  Leaves b r o a d l y s e s s i l e ; c a u l i n e l e a v e s and b r a c t s g r e a t e r than 6 p a i r s ; b a s a l r o s e t t e s l a c k i n g D.  D  Margins of l e a v e s toothed; p l a n t s p r e d o m i n a t e l y e g l a n d u l a r ; l e a v e s rugose; heads nodding i n bud; f l o r e t s b r i g h t y e l l o w ; r a r e a t low e l e v a t i o n s i n S h a s t a Co., C a l i f o r n i a  DD.  9.  A r n i c a venosa  Margins o f l e a v e s e n t i r e ; p l a n t s s t r o n g l y g l a n d u l a r ; l e a v e s smooth; heads e r e c t i n bud; f l o r e t s white to creamy-yellow; r a r e a t h i g h e l e v a t i o n s i n n.  California  139  and CC.  sw Oregon  8. A r n i c a v i s c o s a  Leaves o f b a s a l r o s e t t e s and lower c a u l i n e l e a v e s  distinctly  p e t i o l a t e ; c a u l i n e l e a v e s and b r a c t s l e s s than 6 p a i r s ; basal r o s e t t e s present D.  D  Lower c a u l i n e l e a v e s  spathulate;  glandular  p u r p l i s h or reddish; of serpentine and nw C a l i f o r n i a DD,  hairs  s o i l s i n sw Oregon  7. A r n i c a  Lower c a u l i n e l e a v e s n o t s p a t h u l a t e ;  spathulata  l e a v e s grey t o  green, l a c k i n g p u r p l i s h o r r e d d i s h h a i r s ; u s u a l l y n o t of s e r p e n t i n e  soils,  s Washington t o s C a l i f o r n i a . . . 6. A r n i c a d i s c o i d e a  BB.  Heads r a d i a t e C.  Leaves appearing  C dark green, l a c k i n g n o n - g l a n d u l a r h a i r s ;  p l a n t s m o s t l y densely  clumped, from s h o r t rhizomes 4. A r n i c a  CC.  Leaves appearing  gracilis  g r e y i s h due t o many n o n - g l a n d u l a r h a i r s ;  p l a n t s m o s t l y n o t clumped, from l o n g rhizomes D.  Basal rosette leaves with  cordate  D  bases; p l a n t s  with  many long n o n - g l a n d u l a r h a i r s (1-2 mm); widespread i n w. N.A. e s p e c i a l l y i n open Pinus ponderosa and Pseudotsuga m e n z i e s i i f o r e s t s . . . 1 . A r n i c a DD.  B a s a l r o s e t t e leaves with attenuated  cordifolia  bases; p l a n t s  l a c k i n g o r w i t h few long n o n - g l a n d u l a r h a i r s ; r a r e s p e c i e s , mostly a t h i g h e l e v a t i o n s i n the S i e r r a Nevada AA.  Leaves n o t a t a l l o r very B.  2. A r n i c a n e v a d e n s i s f a i n t l y f r a g r a n t when b r u i s e d  B  Heads nodding i n bud; p l a n t s dark r e d d i s h o r p u r p l i s h throughout,  140  e s p e c i a l l y along v e i n s ; r a r e s p e c i e s of s e r p e n t i n e sw Oregon and nw BB.  California  Heads e r e c t i n bud;  ...5.  p l a n t s mostly g r e e n i s h  soils in A r n i c a cernua  througout except a t  base of stem; common widespread s p e c i e s not  of s e r p e n t i n e 3. 'Arnica  soils.. latifolia  TAXONOMIC TREATMENT  Subgenus Austromontana Maguire  P e r e n n i a l herbs from naked o r s c a l y rhizomes or r a r e l y a s h o r t woody caudex and  rhizomes; r o o t s s l e n d e r or f l e s h y ; s t e r i l e r o s e t t e s of  (innovations)  produced; r o s e t t e l e a v e s and  l a r g e s t , mostly long p e t i o l e d , decreasing and  lower c a u l i n e l e a v e s u s u a l l y  to c r e n a t e ,  s e r r a t e , dentate and  v e s t i t u r e of stems and glandular;  to a t t e n u a t e ,  ovate to  t i p s acute to rounded, margins e n t i r e  undulate;  leaves subglabrate  stems simple  to much-branched;  to s t r i g o s e , p i l o s e , and  stipitate  i n f l o r e s c e n c e a s o l i t a r y head or corymb; heads r a d i a t e or d i s c o i d ;  i n v o l u c r a l b r a c t s narrowly to b r o a d l y florets linear e l l i p t i c  l a n c e o l a t e , acute to acuminate; r a y  to oblong; d i s c f l o r e t s  achenes r i b b e d , s u b g l a b r a t e  to h i r s u t e and  t u b u l a r or g o b l e t - s h a p e d ;  s t i p i t a t e g l a n d u l a r ; pappus white  ( r a r e l y tawny), b a r b e l l a t e to subplumose.  Section E u l a t i f o l i a e  Maguire  Heads r a d i a t e ; o t h e r w i s e ,  1.  the  i n s i z e upward, becoming b r a c t - l i k e  s e s s i l e below i n f l o r e s c e n c e ; c a u l i n e l e a v e s o p p o s i t e , b l a d e s  l a n c e o l a t e , bases cordate  leaves  A r n i c a c o r d i f o l i a Hooker.  Arnica macrophylla Nutt.  as w i t h  F l . Bor.  T r a n s . Am.  the  Am.  subgenus.  1:33.  P h i l . Soc.  Oregon, Blue Mountains, N u t t a l l (Herb. Hook.,  1834. 7:407. 1841. KEW).  TYPE:  141  Arnica c o r d i f o l i a Hook, v a r . macrophylla (Nutt.) Maguire.  Am. M i d i .  Nat. 37:137. 1947. Arnica g r a n d i f o l i a Greene.  P i t t o n i a 4:173. 1900.  TYPE:  Montana, Bridger  Mountains, J u l y 28, 1896, J.H. Flodman 896 (holotype, ND; isotype, MO) (probable h y b r i d , also l i s t e d under synonymy of Arnica Arnica subcordata Greene.  P i t t o n i a 4:173. 1900.  latifolia).  TYPE: A l b e r t a , Athabasca  River, June 26, 1989, Spreadborough, Geol. Surv. Can. 19644 (UC). Arnica pumila Rydb.  Mem. N.Y. Bot. Gdn. 1:433. 1900.  TYPE:  Colorado,  Gray's Peak, 1872, J . Torrey, s.n. (NY). Arnica c o r d i f o l i a Hook. v a r . pumila (Rydb.) Maguire. Arnica p a r v i f o l i a Greene.  Madrono 6:154. 1942.  PI. Baker. 3:28. 1901. TYPE:  Pass, 3,000 m, July 19, 1901.  Colorado, Marshall  C.F. Baker 515 (holotype, ND; isotypes,  POM, WS). Arnica p a n i c u l a t a A. Nels.  Man. Bot. Rocky Mts. 572. 1909.  TYPE: Wyoming,  Carbon Co., Bridger Peak, moist timber, August 24, 1903, L.N. Gooding 1974 holotype, RM; isotype, MO). Arnica evermannii Greene.  Ottawa Nat. 23:215. 1910.  TYPE:  Idaho, P i t t i t  Lake, 2160 m, August 13, 1895, B.W. Evermann 318 (US). Arnica abortiva Greene.  L e a f l e t s 11:47.  1910.  TYPE: Wyoming, Wind River  Mountains, J u l y 23, 1882, W.H. Forwood, _s.n. (US). Arnica g r a n u l i f e r a Rydb. F l . Rocky Mts. 978. 1917.  TYPE: Montana, L i t t l e  B e l t Mountains, J.H. Flodman 896 (NY). Arnica andersonii P i p e r .  Proc. B i o l . Soc. Wash. 33:106.  1902.  TYPE:  B r i t i s h Columbia, Skeene, E l l i s o n , damp woods, September 11, 1910, j;.R. Anderson 677 (US). Arnica o v a l i s Rydb.  N. Am. F l . 34:338.  1927.  TYPE:  A l b e r t a , Crow Nest Pass  142  July 31,  1897,  Macoun, G e o l .  A r n i c a a u s t i n a e Rydb. C o u n t y , June 1898, A r n i c a h u m i l i s Rydb.  N.  Am. F I .  R.M. N.  Surv.  Can.  34:340.  1927.  A u s t i n & Bruce 2165  Am. F I .  34:341.  " t h e s a d d l e " , 2100 m, Macoun, G e o l . A r n i c a c o r d i f o l i a Hook, v a r . 37:138.  72719 (CAN?) (not  C a l i f o r n i a , Lake  (NY).  1927.  Surv.  TYPE:  seen).  TYPE:  A l b e r t a , Lake L o u i s e ,  Can. 65504  h u m i l i s (Rydb.) M a g u i r e .  (NY). Am. M i d i .  Nat.  1947.  Arnica whitneyi Fern.  Rhodora 3 7 : 3 3 4 .  C o u n t y , Copper H a r b o r , Pease 3579  1935.  TYPE:  M i c h i g a n , Keweenaw  dry deciduous woods, J u l y 4 ,  1934,  Fernald &  (G).  A r n i c a c o r d i f o l i a Hook, s u b s p . w h i t n e y i  (Fern.) Maguire.  Brittonia  4:452.  1943. Arnica hardinae St.  John.  C o u n t y , Lake C h a t c o l e t , isotype,  FI.  SW. Wash. 419.  1937.  TYPE:  I d a h o , Benewah  open woods, (5. Weitman 226 ( h o l o t y p e ,  WASH;  WS).  S t r o n g l y r h i z o m a t o u s p e r e n n i a l h e r b ; rhizomes 2 - 1 0 ( - 6 0 ) cm g r o w t h / y e a r , s l e n d e r 1-2 mm t h i c k , naked except f o r few s c a l e s and o l d l e a f bases toward summit, unbranched or few branches n e a r summit, f o r m i n g l a r g e l o o s e p o p u l a t i o n s of s t e r i l e b a s a l r o s e t t e s and f l o w e r i n g s t e m s ; stems s i n g l e or few,  s i m p l e o r branched n e a r base or t h r o u g h o u t ,  rarely  ( 1 0 - ) 1 5 - 4 0 ( - 5 5 ) cm h i g h ,  s p a r s e l y t o d e n s e l y v i l l o u s and s t i p i t a t e g l a n d u l a r b a s a l l y , more d e n s e l y so h i g h e r , sometimes p u r p l i s h o r r e d d i s h a t b a s e ; i n n o v a t i o n s w i t h 2 - 6 ( - 1 4 ) l o n g - p e t i o l a t e l e a v e s , p e t i o l e s ( 2 - ) 6 - 1 0 ( - 2 0 ) cm l o n g , sometimes n a r r o w l y w i n g e d , b l a d e s o v a t e to b r o a d l y o v a t e or l a n c e o l a t e , (l-)3-6(-9)  ( l - ) 4 - 8 ( - 1 2 ) cm l o n g ,  cm b r o a d , bases c o r d a t e , s u b c o r d a t e , t o sometimes r o u n d e d ,  a c u t e to r o u n d e d , m a r g i n s e n t i r e , s u b e n t i r e , t o u s u a l l y r e g u l a r l y  tips  dentate,  143  s e r r a t e , c r e n a t e , or r a r e l y doubly dentate or s e r r a t e ; c a u l i n e l e a v e s p a i r s , o f t e n reduced b a s a l l y and  becoming b r a c t - l i k e toward summit, lower  cauline leaves u s u a l l y l o n g - p e t i o l a t e (l-)4-8(-10) (-13)  cm  long  cm l o n g , blades  (2-)5-10  (2-)4-6(-9) cm broad, upper p a i r s of c a u l i n e l e a v e s w i t h  g r e s s i v e l y s h o r t e r p e t i o l e s , uppermost p a i r u s u a l l y b r a c t - l i k e and  b r a c t s u s u a l l y ovate to l a n c e o l a t e w i t h t u r e of herbage s p a r s e l y to densely s p a r s e l y to u s u a l l y densely l e a f margins c i l i a t e ; (-35)  p i l o s e and bracts  s u b e n t i r e to e n t i r e margins; v e s t i -  s t r i g o s e o r v i l l o u s and  stipitate  l o n g , peduncle and  s t i p i t a t e g l a n d u l a r ; heads r a d i a t e , 15-25 mm  l a n c e o l a t e , acute to acuminate; r a y f l o r e t s ,  linear e l l i p t i c ,  1-3  s t i p i t a t e glandular; s p a r s e l y to densely g l a n d u l a r upward and  glandular,  i n f l o r e s c e n c e u s u a l l y a s i n g l e head or open corymb of  heads; peduncles (4-)8-15(-20) cm  tube 3-7  upper  s t r i g o s e or v i l l o u s on under s u r f a c e s ,  (7-)10-15(-21) + u n i s e r i a t e , 10-18  en y e l l o w ,  pro-  sessile;  shape and margins of lower c a u l i n e l e a v e s as i n that of i n n o v a t i o n s ,  3-7  2-3(-5)  mm  l o n g , limb  disc florets v i l l o u s and on o u t e r  cm h i g h ; i n v o l u c r a l  l o n g , 2-4  mm  broad, narrowly  ( 5 - ) 8 - l l ( - 1 6 ) p a l e t o dark g o l d -  10-25(-35) mm  t e e t h at t i p , tube and  periclinum  l o n g , 4-7(-10) mm  base of limb  sparsely v i l l o u s  (15-)38-57(-94), t u b u l a r 8-12  s t i p i t a t e glandular  broad,  at base and  mm  long,  sparsely  s u r f a c e of l o b e s ; p o l l e n diameter 40-55 broad, s p a r s e l y to densely  um;  achenes dark grey, 4-10  mm  b i f u r c a t e h a i r s and  s t i p i t a t e glandular h a i r s , e s p e c i a l l y d i s t a l l y ;  few  l o n g , 1 mm  pappus white, b a r b e l l a t e , the l a t e r a l setae apomictic;  Type:  .07-.10 mm  and  h i r s u t e with  l o n g , 38-52 b r i s t l e s ;  2n=38,57,76,95,114.  " A l p i n e woods of the Rocky Mountains", Drummond (Herb. Hook.,  Distribution:  KEW).  A r n i c a c o r d i f o l i a i s common i n many moderately dry h a b i t a t s ,  e s p e c i a l l y i n open P i n u s ponderosa, P i n u s c o n t o r t a , and  Pseudotsuga m e n z i e s i i  144  f o r e s t s o r a l p i n e meadows a t many e l e v a t i o n s from c e n t r a l Yukon and Alaska  s o u t h to N o r t h e r n New  Mexico and  B l a c k H i l l s of South Dakota and Betula  spp.  Figure  248.  Figure  east  to  d i s j u n c t i n A b i e s balsamea, Acer spp.,  f o r e s t s i n Keweenaw Co.,  Illustration:  S o u t h e r n C a l i f o r n i a , and  Eastern  M i c h i g a n and  Thunder Bay,  the  and  Ontario.  249.  T h i s , the most w i d e s p r e a d A r n i c a i n Western N o r t h America, i s u s u a l l y abundant where i t o c c u r s ,  covering vast areas.  I t i s o f t e n the dominant  herbaceous p l a n t i n open f o r e s t s i n the Western  Cordillera.  I n i t s more t y p i c a l form, where i t grows i n open p i n e o r Douglas f i r forests,  i t i s d i s t i n c t i v e with  i t s deeply  s o l i t a r y heads, broad r a y s , and great v a r i a b i l i t y These a r e now of Arnica  h i r s u t e achenes.  i n form, which has  g e n e r a l l y accepted  leaves,  large  habitats there i s  l e d to the naming of a number of the range of  taxa.  variability  cordifolia.  have been g i v e n  large leaved  However, t h i s v a r i a t i o n w i t h t i o n s or populations  growing i n sun, CA,  filtered  roadsides  can be  seen from s i n g l e c o l l e c -  shade, and  dense shade, as i n  s e r i e s o f ecotypes from h i g h  to which the names A r n i c a p u m i l a Rydb. and  h u m i l i s Rydb. have been a p p l i e d .  BC.  S-1797 ( F i g u r e s 250-259).  the o t h e r extreme i s a d i s t i n c t  e n t i r e leaves.  These  A r n i c a a n d e r s o n i i P i p e r from Skeena,  large thin leaves  c o l l e c t i o n s from S i s k i y o u Co.,  disturbed  forms o f t e n o c c u r .  taxonomic r e c o g n i t i o n i n the p a s t , as A r n i c a m a c r o p h y l l a N u t t .  from the B l u e Mtns. of Oregon, and  and  toothed  In o t h e r  as f a l l i n g w i t h i n  I n more shaded l o c a t i o n s v e r y  At  cordate,  They a r e q u i t e s h o r t , w i t h  elevations  Arnica  small, usually  They o f t e n resemble A r n i c a n e v a d e n s i s , from which they can  be  145  F i g u r e 248.  D i s t r i b u t i o n o f A r n i c a c o r d i f o l i a i n Western N o r t h America. P o p u l a t i o n s i n Michigan and O n t a r i o a r e not shown.  146  147  ;ure 249.  A r n i c a c o r d i f o l i a S-1494, WA, X two-thirds.  Okanogan County, hab  148  149  F i g u r e s 250-259.  V a r i a t i o n i n b a s a l l e a v e s from s i n g l e p o p u l a t i o n o f A r n i c a c o r d i f o l i a S-1797, CA, S i s k i y o u Co.  150  151  152  d i s t i n g u i s h e d , often only with d i f f i c u l t y ,  by the w h i t e r , l e s s plumose  n a t u r e of the pappus, and u s u a l l y a g r e a t e r degree of p i l o s i t y of the and  stems than i n A r n i c a nevadensis  ecotypes  populations.  T y p i c a l of these  are c o l l e c t i o n s from S l a t e Lake T r a i l , Okanogan Co.,  ( F i g u r e s 260-267), and  roadside  i n Pinus  Co.,  OR,  S-1820 ( F i g u r e s 268-276).  Co.,  CA,  O e t t i n g e r 1082,  both A r n i c a nevadensis  to one  small S-1498  c o n t o r t a f o r e s t , P i n e Grove, Wasco  Many herbarium specimens from S i s k i y o u  Sawyer 941,  1236  (HSC)  and A r n i c a c o r d i f o l i a .  combine many c h a r a c t e r s of Only where t h e r e a r e s e v e r a l  specimens or where they can be s t u d i e d i n the f i e l d assigned  WA,  leaves  o r the o t h e r of these  can  they be more e a s i l y  species.  L a r g e r , more t y p i c a l A r n i c a c o r d i f o l i a i s s e p a r a b l e from t y p i c a l A r n i c a nevadensis and  i n the g e n e r a l l y narrower, p a l e r r a y s , more densely p i l o s e stems  l e a v e s , more toothed margins, w h i t e r ,  and more densely  s h o r t e r l a t e r a l setae of the pappus,  h i r s u t e achenes o f the former.  A r n i c a c o r d i f o l i a w i t h broader S i s k i y o u Co.,  CA,  S-1797.  densis rays.  The  two  rays  ( F i g u r e 245)  Lake, Mono Co.,  The CA,  S-1937, 1938,  and A r n i c a nevadensis Although  as t y p i c a l A r n i c a neva-  species are a l s o u s u a l l y e c o l o g i c a l l y separate.  two have been observed  Arnica  i n more p r o t e c t e d h a b i t a t s  growing t o g e t h e r a t Saddlebag  w i t h A r n i c a c o r d i f o l i a i n the f o r e s t edge  on more exposed rock  outcrops.  A r n i c a c o r d i f o l i a g e n e r a l l y has o n l y one  o r t h r e e heads p e r  stem, some p o p u l a t i o n s or i n d i v i d u a l s e s p e c i a l l y i n wetter h a b i t a t s , have much branched stems w i t h many heads t e r m i n a t i n g each branch. extreme o f t h i s was Tod Mtn.,  BC,  seen i n one  S-1652.  of  as i n a p o p u l a t i o n from  These rays are as broad  c o r d i f o l i a u s u a l l y grows a t lower e l e v a t i o n s and of open f o r e s t s .  There are some p o p u l a t i o n s  The  greatest  p l a n t w i t h 35 heads from a c o l l e c t i o n  from  153  F i g u r e s 260-267.  V a r i a t i o n i n b a s a l l e a v e s of h i g h e l e v a t i o n p o p u l a t i o n of A r n i c a c o r d i f o l i a S-1498, WA,  Okanogan Co.  154  155  F i g u r e s 268-276.  V a r i a t i o n i n b a s a l l e a v e s from p o p u l a t i o n on d i s t u r b e d r o a d s i d e bank of A r n i c a c o r d i f o l i a S-1820, OR, Wasco Co.  156  157  Other c o l l e c t i o n s w i t h s e v e r a l heads have been named A r n i c a p a n i c u l a t a by N e l s o n ,  a s p e c i e s r e t a i n e d by Maguire (1943), but c o n s i d e r e d a v a r i a n t of  A r n i c a c o r d i f o l i a by E d i g e r & B a r k l e y  (1978).  Maguire (1943) p o i n t s out  p o s s i b i l i t y of t h i s being a h y b r i d between A r n i c a c o r d i f o l i a and p a r r y i (subgenus Chamissonis), apomictic  ( B a r k e r , 1966).  a l t h o u g h both  Arnica  s p e c i e s a r e l a r g e l y or  A r e c e n t h y b r i d between the two  the  wholly  i s unlikely.  It  seems t h a t the c h a r a c t e r s c i t e d c o u l d f a l l w i t h i n the range of v a r i a t i o n of an a p o m i c t i c race of A r n i c a c o r d i f o l i a , and variant  i t i s t r e a t e d i n t h i s work as a  thereof.  Maguire (1943) a l s o gave taxonomic r e c o g n i t i o n to a p o p u l a t i o n of A r n i c a c o r d i f o l i a from Keweenaw Co., S i n c e 1943  Michigan,  t r e a t i n g i t as subsp. w h i t n e y i .  a d d i t i o n a l p o p u l a t i o n s have been d i s c o v e r e d on the n o r t h shore  Lake S u p e r i o r i n S i b l e y P r o v i n c i a l Park, Thunder Bay 15164, 15486 (LKHD). field  D i s t r i c t , Ontario  They are on the average t a l l e r , have more heads (3/  stem), narrower r a y s , and  l e s s pubescence than average A r n i c a c o r d i f o l i a popu-  l a t i o n s i n the Western mountains.  However, t h e r e are numerous p o p u l a t i o n s i n  the West which cannot be d i s t i n g u i s h e d from the Michigan The  species l i s t lation  p o p u l a t i o n i n Michigan i n that state.  i s v e r y r a r e and  I t i s found  as one  and  Ontario  The  spicatum,  l a r g e widely  Acer  d i s p e r s e d popu-  leaves are separable  Eagle Harbor,  saccharum, and B e t u l a  dominant herbaceous cover i n the a r e a i s A s t e r  When n e i t h e r the A s t e r nor  popula-  i s on the endangered  ( o r s e v e r a l s m a l l p o p u l a t i o n s ) between Copper Harbor and  growing i n open A b i e s balsamea, Acer forest.  Garton  Both of these p o p u l a t i o n s have been s t u d i e d i n the  (S-1846, 1849).  tions.  of  spp.  macrophyllus.  the A r n i c a a r e i n f l o w e r , the b a s a l r o s e t t e s of  only w i t h g r e a t d i f f i c u l t y .  The A r n i c a l e a v e s tend  be s m a l l e r , s l i g h t l y g r e y e r , and have a s t r o n g e r odor.  The  A r n i c a may  be  to  158  F i g u r e s 277-281.  H a b i t a t s and H a b i t s of A r n i c a c o r d i f o l i a  and  Arnica  nevadensis. F i g u r e 277.  H a b i t a t of A r n i c a c o r d i f o l i a , WA,  Klickitat  Co.,  Appleton,  (S-1823), open f o r e s t of Pseudotsuga m e n z i e s i i and  Pinus  ponderosa. F i g u r e 278.  Arnica cordifolia, at Hat  F i g u r e 279.  Creek, BC,  (Johns 648)  (putative hybrid),  d i s c o i d head on same stem.  H a b i t a t of A r n i c a n e v a d e n s i s ,  CA,  S i e r r a Co.,  (S-1870), open f o r e s t of Pinus m o n t i c o l a ,  F i g u r e 281.  MT,  C h a r l e s Waters Campground, (S-1475), normal  r a d i a t e head and  and Abies  collected  (S-2009).  Arnica c o r d i f o l i a X Arnica l a t i f o l i a R a v a l l i Co.,  F i g u r e 280.  6-month o l d s e e d l i n g , from seed  Sierra  Buttes,  Tsuga m e r t e n s i a n a ,  sp.  A r n i c a nevadensis,  NV,  Washoe Co.,  Mt.  Rose, (S-1910).  159  160  more common i n the area than now  presumed, but has been o v e r l o o k e d when n o t  i n f l o w e r because o f the s t r o n g resemblance to the A s t e r .  The p l a n t s from  S i b l e y P r o v i n c i a l Park are e c o l o g i c a l l y and m o r p h o l o g i c a l l y q u i t e s i m i l a r the Michigan  ones.  to  T h i s p l a n t should be sought on I s l e Royale i n Lake  S u p e r i o r between the known p o p u l a t i o n s i n Michigan  and  Ontario.  w h i t n e y i Fern, i s not g i v e n s p e c i f i c s t a t u s o r r e c o g n i z e d A r n i c a c o r d i f o l i a i n t h i s treatment,  Arnica  as a s u b s p e c i e s  of  d e s p i t e the d i s j u n c t d i s t r i b u t i o n .  Chromosomally they are l i k e the dominant Western p o l y p l o i d complex (2n=76) and m o r p h o l o g i c a l l y e.g.,  they a r e l e s s d i f f e r e n t than  some of the o t h e r extremes,  the dwarf forms d i s c u s s e d above, or the southern  extremes i n A r i z o n a  ( S c h r e i b e r , _s.n. , Hemmingway, _s.n. MNA) . There a r e numerous i n t e r e s t i n g p o p u l a t i o n s which are i n many morphol o g i c a l aspects  i n t e r m e d i a t e between A r n i c a c o r d i f o l i a and A r n i c a  Most n o t a b l e are s e v e r a l c o l l e c t i o n s from Josephine Savage, _s.n. (ORE), and  Curry  MT,  S-1474, 1475  one  o r e i t h e r of the p a r e n t s , and  probably  ( F i g u r e 282).  Co.,  OR,  Leach 1544,  Co., 1545  OR,  latifolia.  Lang 798  (SOC),  (ORE), R a v a l l i  These p o p u l a t i o n s u s u a l l y do not grow w i t h are considered  to be o l d h y b r i d s which have  become a p o m i c t i c a f t e r h y b r i d i z a t i o n and have r e t a i n e d t h e i r  mediate c h a r a c t e r s .  Co.,  Pollen stainability  ranges from 33%-99%.  chromosome numbers have been o b t a i n e d from two  inter-  Diploid  Montana c o l l e c t i o n s .  They  appear to be s t e r i l e having never produced any v i a b l e achenes i n c u l t i v a t i o n , e i t h e r when s e l f e d or c r o s s e d w i t h o t h e r s p e c i e s o r s i m i l a r  populations.  These p u t a t i v e h y b r i d s are u s u a l l y much l a r g e r and more robust c h a r a c t e r s than e i t h e r p a r e n t . cordifolia. but  The  in a l l  They are s t r o n g l y rhizomatous l i k e A r n i c a  b a s a l l e a v e s and  lower c a u l i n e l e a v e s are u s u a l l y +  the upper c a u l i n e l e a v e s are s e s s i l e .  The  cordate,  l e a v e s are s p a r s e l y pubescent  161  F i g u r e 282.  A r n i c a c o r d i f o l i a X A r n i c a l a t i f o l i a S-1475, p u t a t i v e n a t u r a l h y b r i d , MT,  Ravalli  County.  X  two-thirds.  162  163  and  appear dark green as i n A r n i c a l a t i f o l i a .  However, g l a n d u l a r  h a i r s are  present  throughout g i v i n g the p l a n t s as s t r o n g an odor as i n A r n i c a c o r d i -  folia.  Heads are u s u a l l y l a r g e r than i n e i t h e r p a r e n t ,  25), more d i s c f l o r e t s The  shape of the r a y s  more t r u n c a t e t i p s .  ( n e a r l y 200) ( F i g u r e 138)  i n d i v i d u a l s c l o s e r to one m i c t i c r a c e s and  2.  a r e not  E.L.  are more densely  to  21).  approaches t h a t of A r n i c a l a t i f o l i a ,  with  pubescent  There are numerous o t h e r p o p u l a t i o n s  or the o t h e r p a r e n t .  Gray. Proc. Am.  A r n i c a c h i o n o p h i l a Greene. 1896,  (up to  (up  or  A l l these are c o n s i d e r e d  apo-  g i v e n taxonomic s t a t u s .  A r n i c a n e v a d e n s i s A.  J u l y 20,  and more i n v o l u c r a l b r a c t s  D i s c f l o r e t s ( F i g u r e 144)  throughout than i n e i t h e r p a r e n t .  w i t h more r a y s  Acad. 19:55.  P i t t o n i a 4:171. 1900.  Greene (ND?)  (not  1883.  TYPE:  Nevada, Ruby  Mts.,  seen).  Rhizomatous p e r e n n i a l herb; rhizomes ( l - ) 3 - 1 2 cm  growth/year,  1-2  mm  t h i c k , naked except p a i r s of brown s c a l e s at nodes and  and  o l d l e a f bases toward the summit, unbranched or few  slender,  crowded s c a l e s  branches a p i c a l l y ,  forming s m a l l to l a r g e p o p u l a t i o n s  of s t e r i l e b a s a l r o s e t t e s and  stems; stems s i n g l e or few,  or branched from base or above, (10-)15-30  cm  tall,  very  vations with  simple  s p a r s e l y p i l o s e and + densely  short g l a n d u l a r  (2-)4-6 long p e t i o l a t e l e a v e s , p e t i o l e s 3-5  ovate, rotund,  or e l l i p t i c  rounded to a t t e n u a t e ,  to l a n c e o l a t e , 4-7  cm  cm  l o n g , 2-4  flowering  throughout; long,  blades  cm broad, bases  t i p s acute to rounded, margins e n t i r e , s u b e n t i r e ,  r a r e l y c o a r s e l y s e r r a t e or dentate;  cauline leaves  2-3  inno-  to  p a i r s , lowest p a i r  u s u a l l y w i t h l o n g e s t p e t i o l e s , ( l - ) 2 - 3 ( - 4 ) cm l o n g , sometimes winged to 4 next p a i r u s u a l l y w i t h l a r g e s t b l a d e 3-5.5 p a i r u s u a l l y reduced and  cm  long,  mm,  ( l - ) 2 - 4 cm broad, upper  b r a c t - l i k e , l a n c e o l a t e , shape and margins of  cauline  164  l e a v e s l i k e t h a t of i n n o v a t i o n s ; v e s t i t u r e of herbage s p a r s e l y p i l o s e and  short s t i p i t a t e glandular  throughout, margins c i l i a t e ;  cence u s u a l l y of a s i n g l e head o r open corymb of 3(-7) cm l o n g ; peduncles and and mm  p e r i c l i n u m densely  s p a r s e l y p i l o s e ; heads r a d i a t e 15-20 l o n g , 2-4  5-13,  mm  v i l l o u s and  mm  mm  1-3  l o n g , s p a r s e l y to densely l y d i s t a l l y and  on o u t e r  to b l a c k , 4-9  inflores-  heads; peduncles  long s t i p i t a t e  glandular  to acuminate; ray mm  florets  l o n g , 6-7  mm  broad,  lower back of l i m b , l i n e a r  ellip-  t e e t h a t t i p ; d i s c f l o r e t s 21-51, t u b u l a r 7-10  v i l l o u s and  s t i p i t a t e g l a n d u l a r at base and  s u r f a c e of l o b e s ; p o l l e n diameter 35-45 um, mm  4-12  h i g h ; i n v o l u c r a l b r a c t s 10-15(-18)  l o n g , limb 15-20  s t i p i t a t e g l a n d u l a r on tube and  t i c to b r o a d l y e l l i p t i c ,  dark grey  s h o r t and  wide, n a r r o w l y l a n c e o l a t e , acute  dark golden y e l l o w , tube 2-7  scabrous,  l o n g , to 1 mm  cate h a i r s e s p e c i a l l y d i s t a l l y ,  wide, s p a r s e l y h i r s u t e w i t h  s h o r t to long s t i p i t a t e g l a n d u l a r  sparingachenes bifur-  throughout;  pappus d u l l white to tawny, of 27-50 b r i s t l e s , b a r b e l l a t e to subplumose, l a t e r a l setae  TYPE:  .15-.20 mm;  Mostly  P r i n g l e s.n.  at h i g h e l e v a t i o n s (1500-3500 m)  Tsuga m e r t e n s i a n a , Pinus m o n t i c o l a , of C a l i f o r n i a and Nevada and  and Pinus  (G). i n open f o r e s t s of  c o n t o r t a i n the S i e r r a Nevada  s p a r i n g l y e a s t to the San  Juan Mountains of Utah  and n o r t h i n the Cascade Mountains to n o r t h - c e n t r a l Washington.  Illustration:  Figure  the  a p o m i c t i c ; 2n=76.  C a l i f o r n i a , S i e r r a Nevada, 2700 m,  Distribution:  (Figure  283).  284.  A r n i c a nevadensis i s a d i s t i n c t can be r e a d i l y separated  mm  s p e c i e s i n the S i e r r a Nevada, where i t  from A r n i c a c o r d i f o l i a most o f the time.  i t s other d i s t r i b u t i o n i t i s only separable Arnica c o r d i f o l i a with d i f f i c u l t y ,  In much o f  from h i g h e l e v a t i o n forms of  e s p e c i a l l y from herbarium specimens, as  165  F i g u r e 283.  D i s t r i b u t i o n of A r n i c a  nevadensis.  166  167  gure 284.  A r n i c a nevadensis  S-1910, NV,  Washoe Co.  X two-thirds  168  169  f u r t h e r d i s c u s s e d under t h a t Occasional  populations  species. of A r n i c a r y d b e r g i i growing i n shaded l o c a t i o n s ,  as from H u r r i c a n Ridge, C l a l l a m County, WA that s p e c i e s and  c o u l d e a s i l y be  (S-1677) , are v e r y a t y p i c a l o f  i d e n t i f i e d as A r n i c a n e v a d e n s i s .  v i d u a l s have broad rounded b a s a l and  lower c a u l i n e l e a v e s .  narrower heads, pure w h i t e pappus, and  densely  Many i n d i -  However, the  h i r s u t e achenes r e a d i l y sepa-  r a t e i t from A r n i c a n e v a d e n s i s . In the S i e r r a Nevada, A r n i c a nevadensis i s not growing w i t h Whitneya d e a l b a t a ,  w i t h which i t has  o f t e n been confused.  p e r s i s t e n t , more-papery r a y s on mature achenes and Whitneya are  absence of pappus i n  Greene i s a v e r y dubious s p e c i e s r e t a i n e d as v a l i d  both Maguire (1943) and  genus.  Ediger  & Barkley  (1978).  has not been seen i n n a t u r e by any  Most of the specimens have v e r y  of  US;  i s o t y p e MO),  of the recent workers on  insufficient  data,  the  i n c l u d i n g the  so that r e l o c a t i n g i t i n the f i e l d  California" seems a matter  luck. Maguire (1943) c o n s i d e r e d  or A r n i c a n e v a d e n s i s . and  by  I t i s very r a r e i n c o l l e c -  Type, Purpus 5625, "open woods, middle T u l e R i v e r , s o u t h e a s t e r n (holotype,  The  distinctive.  Arnica tomentella  t i o n s and  i n f r e q u e n t l y found  The  Arnica tomentella  p l a n t s are t a l l e r  r e l a t e d to A r n i c a  cordifolia  than average A r n i c a nevadensis  the i n v o l u c r a l b r a c t s are more pubescent, e s p e c i a l l y near the t i p s .  The  l a t e r a l setae are c l o s e r to those of A r n i c a n e v a d e n s i s , a l t h o u g h i n o l d herbarium specimens the c o l o r of the pappus cannot be determined and  the  shape o f the r a y s i s d i f f i c u l t t o determine. Ediger  & Barkley  (1978) c o n s i d e r A r n i c a t o m e n t e l l a  a possible hybrid  between A r n i c a c o r d i f o l i a and A r n i c a c h a m i s s o n i s v a r . f o l i o s a , p r o b a b l y  based  170  on the a t t e n u a t e  l e a f bases and pubescent i n v o l u c r a l b r a c t t i p s .  However,  the g e n e r a l c h a r a c t e r of t h e pubsecence i s c l o s e to t h a t of A r n i c a  nevadensis,  as are t h e l e a v e s , and  apomictic  i t i s t e n t a t i v e l y p l a c e d here as a probable  r a c e of A r n i c a n e v a d e n s i s ,  3.  Arnica l a t i f o l i a  until  Bongard. Mem.  i t can be found i n the f i e l d  Acad. S t . P e t e r s b .  A r n i c a l a t i f o l i a a genuina Herder. B u l l . Arnica l a t i f o l i a  Soc. Nat.  8 a n g u s t i f o l i a Herder. B u l l .  A r n i c a l a t i f o l i a A. Gray. Syn.  F I . N. Am.  A r n i c a m e n z i e s i i Hooker. F I . Bor.  Am.  VI. 2 : 1 4 7 .  1832.  Mosc. 4 0 : 4 2 4 . 1 8 6 7 .  Soc. Nat.  1:381.  again.  Mosc. 4 0 : 4 2 4 .  1867.  1884.  1:331. 1834.  TYPE:  North West Coast  of America, A. Menzies, _s.n. (KEW). A r n i c a b e t o n i c a e f o l i a Greene. Olympic Mountains, Mt. Arnica t e u c r i i f o l i a  P i t t o n i a 4:163.  S t e e l e , 1 8 0 0 m,  Greene.  1900.  TYPE:  August 1 8 9 5 , _C.V.  P i t t o n i a 4:164.  1900.  d'Alene Mountains, d i v i d e between S t . Joe and  TYPE:  Washington, Piper 2002  (ND).  Idaho, Coeur  Clearwater  Rivers, 1820  g r a s s y mountain s l o p e s , J u l y 1 0 , 1 8 9 5 , _J.I3. L e i b e r g 1 2 2 9 ( h o l o t y p e , i s o t y p e s , MO,  1:171.  teucriifolia  (Greene) L. W i l l i a m s .  Leafl.  1935.  A r n i c a ventorum Greene.  P i t t o n i a 4:173.  1900.  TYPE:  Wyoming, Wind R i v e r  Mountains, Union Pass, August 1 1 , 1 8 9 4 , A. Nelson 836 ( h o l o t y p e , i s o t y p e s , MO,  ND;  WS).  A r n i c a g r a n d i f o l i a Greene.  P i t t o n i a 4:173.  Mountains, J u l y 2 8 , 1 8 9 6 , J^.H. A r n i c a p l a t y p h y l l a A. Nelson. Bot.  1900.  TYPE:  Montana, B r i d g e r  Floodman 896 ( h o l o t y p e , ND; Gaz.  31:407. 1901.  TYPE:  isotype,  MO).  Oregon, Cascade  Mountains, Hood R i v e r , f i r fo r e s t s , J u l y 1 8 , 1 8 9 6 , I J . F . Henderson, (RM) .  US;  POM).  A r n i c a l a t i f o l i a Bong. v a r . West. Bot.  m,  s.n.  171  A r n i c a l a e v i g a t a Greene.  Ottawa Nat. 15:279. 1902.  C h i l l i w a c k V a l l e y , 900 m, by s p r i n g s  TYPE:  British  Columbia,  i n woods, August 15, 1901, J..M. Macoun,  G e o l . Surv. Can. 26926 (ND). A r n i c a a p r i c a Greene.  Ottawa Nat. 15:279. 1902.  TYPE:  British  C h i l l i w a c k V a l l e y , 1050 m, open ground a l o n g s t r e a m l e t s , _J.M. Macoun, G e o l . Surv. Can. 26284 ( h o l o t y p e , Arnica j o n e s i i  Rydb. F l . Rocky Mts. 978. 1917.  J u l y 10, 1901,  ND; i s o t y p e , MO) .  TYPE:  Utah, A l t a , Wasatch  Mountains, 2700 m, J u l y 31, 1879, M.E. Jones 119 ( h o l o t y p e , POM,  NY;  isotypes,  UTC) .  A r n i c a p u b e r u l a Rydb. F l . Rocky Mts. 979. 1917. Louise,  TYPE:  A l b e r t a , head of Lake  J u l y 22, 1904, J_.M. Macoun, G e o l . Surv. Can. 65523 (NY).  A r n i c a e r i o p o d a Gandoger. B u l l .  Soc. B o t . France 6538. 1918.  Cascade Mountains, J u l y 27, 1902, W._C. i s o t y p e s , POM,  Oregon,  US;  MO).  Washington, Mt. Baker, 1915, C^.W. flodmanii  TYPE:  C u s i c k 2914, ( h o l o t y p e ,  A r n i c a a p h a n a c t i s P i p e r . P r o c . B i o l . Soc. Wash. 33:105. 1920.  Arnica  Columbia,  Rydb. N.Am.  TYPE:  Turesson, _s.n. (NY).  F l . 34:334. 1927.  TYPE:  Montana, Madison Range,  Spanish Peaks, 2100-2400 m, J u l y 14, 1896, j;.H. Flodman 898, ( h o l o t y p e , isotype,  NY;  MO).  A r n i c a g l a b r a t a Rydb. N. Am. August 10, 1897, A u s t i n  F l . 34:335. 1927.  TYPE:  & Bruce 1627, ( h o l o t y p e ,  A r n i c a o l i g o l e p i s Rydb. N. Am.  F l . 34:336. 1927.  Skeena R i v e r , H a z e l ton, June 23, 1917,  Oregon, C r a t e r  NY; i s o t y p e s ,  TYPE:  J_.JA.  British  Lake,  POM). Columbia,  Macoun, Geol. Surv. Can. 96048  (NY) . A r n i c a l e p t o c a u l i s Rydb. N. Am.  F l . 34:336. 1927.  TYPE:  British  Columbia,  Vancouver I s l a n d , Mt. Mark, J u l y 25, 1887, _J.M. Macoun, Geol. Surv. Can. 14570 (NY).  172  A r n i c a membranacea Rydb. N. Am. F l . 34:338. 1927. Wimmer, June 13, 1892, E.W. Hammond 231,  TYPE:  Oregon, J a c k s o n Co.,  (NY).  Rhizomatous p e r e n n i a l herb; rhizomes l-7(-10) cm/year, s t o u t , 3-5(-7) mm t h i c k , naked o r densely  clothed with  o l d l e a f bases and dark brown s c a l e s ,  unbranched o r few-branched near summit, forming l o o s e o r dense clumps of s t e r i l e b a s a l r o s e t t e s and f l o w e r i n g stems; stems s i n g l e o r few, simple o r branched above upper p a i r o f l e a v e s / b r a c t s , r a r e l y branched lower, (10-)15-40 cm  tall,  g l a b r a t e to very  s p a r s e l y v i l l o u s throughout, d i s t i n c t l y p u r p l i s h o r  r e d d i s h a t base; i n n o v a t i o n s (2-)4-8(-12) cm, blades long,  with  2-6(-12) l o n g - p e t i o l a t e l e a v e s , p e t i o l e s  narrowly e l l i p t i c  to ovate o r rotund,  (0.5-)2-4(-5) cm broad, bases a t t e n u a t e  r a r e l y cordate,  ( l - ) 3 - 5 ( - 6 ) cm  t o rounded, s u b c o r d a t e , o r  t i p s acute, margins e n t i r e to u s u a l l y r e g u l a r l y  s e r r a t e , o r doubly dentate o r s e r r a t e , o r c r e n a t e ;  dentate,  cauline leaves  2-4(-7)  p a i r s , o f t e n reduced b a s a l l y and toward summit, where b r a c t - l i k e , lower caul i n e leaves mm, b l a d e s ly  s e s s i l e or with  short p e t i o l e s l - 3 ( - 5 )  cm l o n g , o f t e n winged 2-5  2-6(-9) cm l o n g , 1-4(-7) cm wide, upper c a u l i n e l e a v e s  s m a l l e r , almost always s e s s i l e ,  progressive-  shape and margins as i n l e a v e s o f i n n o v a -  t i o n s ; v e s t i t u r e o f herbage g l a b r a t e t o s p a r s e l y ( r a r e l y densely) v i l l o u s , s t r i g o s e , and v e r y  r a r e l y s h o r t s t i p i t a t e g l a n d u l a r , margins o f t e n  ciliate;  i n f l o r e s c e n c e o f a s i n g l e head o r open corymb; peduncles (1-)3-10(-20) cm l o n g , becoming more v i l l o u s upward; p e r i c l i n u m s p a r s e l y to densely heads l - 3 ( - 1 3 ) , r a d i a t e , 10-20 mm h i g h ; u n i s e r i a t e , 8-14 mm l o n g , acuminate; r a y f l o r e t s  i n v o l u c r a l bracts  villous;  (7-)ll-16(-23), +  ( l - ) 2 - 3 mm broad, n a r r o w l y l a n c e o l a t e , acute t o  (5-)8-15(-18), u s u a l l y p a l e y e l l o w ,  (5-)10-25 mm l o n g , 2-5 mm wide, l i n e a r e l l i p t i c , t e e t h , tube and base and back o f limb  truncate  sparsely v i l l o u s ;  tube 3-5 mm,  limb  a t t i p , w i t h 0-3  disc florets  173 (15-)50-90(-124), tubular,  5-10 mm l o n g , s p a r s e l y to d e n s e l y v i l l o u s near  base and on o u t e r s u r f a c e of l o b e s ; p o l l e n diameter 35-50 um; achenes dark grey-brown, tally,  5-10 mm l o n g , 1 mm w i d e ,  r a r e l y throughout,  throughout,  few b i f u r c a t e h a i r s from m i d - p o i n t  dis-  few s h o r t s t i p i t a t e g l a n d u l a r h a i r s d i s t a l l y or  r a r e l y g l a b r o u s ; pappus w h i t e , b a r b e l l a t e ,  the l a t e r a l s e t a e  .05-  .07 mm l o n g , of 43-64 b r i s t l e s ; s e l f - i n c o m p a t i b l e ; 2n=38.  TYPE:  A l a s k a , S i t k a , M e r t e n s , £.n_.  Distribution:  (Herb. B o n g a r d ,  Leningrad).  A r n i c a l a t i f o l i a o c c u r s i n many h a b i t a t s a t v a r y i n g  elevations,  m o s t l y i n c o o l mixed c o n i f e r f o r e s t s i n the mountains from S o u t h e r n A l a s k a t o N o r t h e r n C a l i f o r n i a and C e n t r a l C o l o r a d o . Illustration:  (Figure  285).  F i g u r e 286.  A r n i c a l a t i f o l i a i s one of t h e commonest and most p o l y m o r p h i c of a l l species.  It  u s u a l l y grows i n w e t t e r  l o c a t i o n s than A r n i c a c o r d i f o l i a ,  e s p e c i a l l y i n c o o l f o r e s t s of Pseudotsuga m e n z i e s i i , P i c e a s p p . , and A b i e s s p p . and near stream m a r g i n s . There are so many ecotypes t o w h i c h names have been a p p l i e d t h a t i t d i f f i c u l t to d e s c r i b e the " t y p i c a l " A r n i c a l a t i f o l i a . l i n e l e a v e s , the r a y s a r e u s u a l l y t r u n c a t e stems a r e u s u a l l y g l a b r o u s , throughout  is  Most have s e s s i l e c a u -  (Figure 139),  the l e a v e s and  and the mature achenes a r e u s u a l l y  glabrous  or h i r s u t e o n l y on t h e d i s t a l h a l f .  Among the v a r i a b l e ecotypes are a group of b r o a d - l e a v e d ,  robust  t i o n s w i t h t h i n l e a v e s from shady l o c a t i o n s , and another group of narrower-leaved,  o f t e n h i g h e l e v a t i o n , sun forms w i t h much t h i c k e r  popula-  smaller, leaves.  The extreme c o n d i t i o n of the shade form i s r e p r e s e n t e d i n c o l l e c t i o n s from Mt.  B a k e r , Whatcom C o . , WA,  S - 1 1 5 8 , 1161.  These are v e r y r o b u s t p l a n t s  to  174  ;ure 285.  D i s t r i b u t i o n of A r n i c a locality.  latifolia.  Star indicates  type  175  176  F i g u r e 286.  A r n i c a l a t i f o l i a S-1500, WA,  Chelan Co.  X  two-thirds.  177  178  40 cm  tall,  form has  w i t h t h i n broad l e a v e s and  c o a r s e l y s e r r a t e margins.  r e c e i v e d taxonomic r e c o g n i t i o n i n the past w i t h  This  s e v e r a l names  been a p p l i e d ( A r n i c a p l a t y p h y l l a N e l s . , A r n i c a a p r i c a Greene, A r n i c a gata  Greene).  At the o p p o s i t e  Arnica g r a c i l i s . tall),  clumped, and  branched, w i t h  h a i r s ( o r l a c k i n g h a i r s o f any  s m a l l heads, but  (UBC).  Green Mtn.,  Vancouver I s . , BC,  f o l i a Greene has  l a r g e r and more robust  Every p o s s i b l e combination o f e x i s t s between the l a r g e and  much of the d i s t r i b u t i o n of the  erroneously  One  betonicae-  such form  experi-  intermediate  s m a l l ecotypes from  p a r t of the range f o r p o p u l a t i o n s  have a g r e a t e r p i l o s i t y to the l e a v e s and T a y l o r , et a l . 5993, I l g a c h u z  to the  species.  There i s a tendency i n the n o r t h e r n  stems ( B u t t r i c k 779,  Mtns., BC,  from the Queen C h a r l o t t e I s l a n d s , BC,  all  and  A t l i n Lake,  has  not been seen, a l t h o u g h S-1961, some p l a n t s  rays which were v a r i o u s l y shortened ( F i g u r e s 299-304), compared to (Figure  139).  BC,  UBC).  from Nancy Greene P r o v i n c i a l Park, BC,  rays f o r t h i s s p e c i e s  to  e s p e c i a l l y some c o l l e c t i o n s  A truly discoid condition in Arnica l a t i f o l i a  had  (UBC),  name A r n i c a  the f o l l o w i n g year when t r a n s p l a n t e d  i n d i v i d u a l s and p o p u l a t i o n s  population  include  S-1561 growing on a d i s t u r b e d r o a d s i d e , became much  mental garden i n Vancouver.  i n one  The  been a p p l i e d to these ecotypes i n the p a s t .  from Kootenay Pass, BC,  paler,  K r a j i n a et a l . 4987  Records of A r n i c a g r a c i l i s from Vancouver I s . probably  r e f e r to these s m a l l ecotypes of A r n i c a l a t i f o l i a .  and  A l l e n , _s.ii.  cm  glandular  Examples of these s m a l l ecotypes  Arrowsmith, Vancouver I s . , BC,  ( F i g u r e s 287-298), and  l a c k i n g the  resemble  as 10  type) of A r n i c a g r a c i l i s , and having  b l u n t e r r a y s than A r n i c a g r a c i l i s . c o l l e c t i o n s from Mt.  (as l i t t l e  having  laevi-  extreme are p l a n t s which s u p e r f i c i a l l y  These u s u a l l y a l p i n e p l a n t s are short  large  typical  179  F i g u r e s 287-298.  V a r i a t i o n i n b a s a l l e a v e s of s m a l l a l p i n e form of A r n i c a l a t i f o l i a A l l e n s.n., BC, Mt.  Arrowsmith.  180  181  F i g u r e s 299-304.  V a r i a t i o n i n shortened r a y f l o r e t s from heads on one p l a n t of A r n i c a Nancy Greene Prov. Park.  several  l a t i f o l i a S-1961, BC,  183  The  r e l a t i o n s h i p of A r n i c a l a t i f o l i a  diversifolia  (both  to A r n i c a m o l l i s and  Arnica  i n Subgenus Chamissonis) deserves f u r t h e r i n v e s t i g a t i o n .  Arnica d i v e r s i f o l i a  has  been t r e a t e d i n p r e v i o u s  two  Arnica l a t i f o l i a ,  i n c l u d i n g s e v e r a l c o l l e c t i o n s from G l a c i e r N a t i o n a l  Harvey 5163,  There are p o p u l a t i o n s  hybrid  between the o t h e r  MT,  species.  works as a p o s s i b l e  which s t r o n g l y resemble  5574 (MONTU), which have been determined as A r n i c a  Park,  diversi-  folia. Further  d i s c u s s i o n s of the r e l a t i o n s h i p of t h i s  Arnica c o r d i f o l i a those  4.  and  A r n i c a g r a c i l i s may  species  to A r n i c a  cernua,  be found under the d i s c u s s i o n s  of  species.  A r n i c a g r a c i l i s Rydberg, B u l l . T o r r . Bot.  Arnica betonicaefolia var. g r a c i l i s B i o l . 15:45.  Club 24:297.  (Rydb.) M.E.  1897.  Jones, B u l l .  Univ. Mont.  1910.  A r n i c a l a t i f o l i a Bong, v a r . g r a c i l i s  (Rydb.) Cronq. Vase. P I . Pac.  NW.  5:51.  1955. Arnica  Columbiana A. N e l s . ,  F a l l s , R._S.  Williams  1049  A r n i c a m u l t i f l o r a Greene. Orielle,  Leiberg  234  Bot.  Gaz.  TYPE:  Montana, Columbia  (RM) .  P i t t o n i a 4:162. 1900.  (holotype,  A r n i c a o v a l i f o l i a Greene.  30:200. 1900.  ND;  TYPE:  Idaho, Lake Pend  p a r a t y p e s , MONTU, NY,  P i t t o n i a 4:168. 1900.  TYPE:  WS).  Wyoming, B i g Horn  Mountains, B l a n k i n s h i p , _s._n. (ND) . A r n i c a arcana A. N e l s . , Bot.  Gaz.  Mountains, Doyle Creek, N._L.  37:276. 1904.  Gooding 377  A r n i c a l a c t u c i n a Greene, Ottawa Nat. W.jC.  M c C a l l a 2014  (ND) .  TYPE:  (holotype,  33:215. 1910.  Wyoming, B i g Horn RM;  isotype,  TYPE:  Alberta,  MO). Banff,  184  Rhizomatous, o f t e n clumped p e r e n n i a l herb; rhizome growth 0.5-3(-6) per y e a r , and  s l e n d e r , 1-2  mm  t h i c k , naked on o l d e r p a r t s or w i t h o l d l e a f  bases  brown s c a l e s e s p e c i a l l y toward the summit, unbranched or u s u a l l y s e v e r a l  branched forming clumps of s t e r i l e b a s a l r o s e t t e s and s i n g l e or u s u a l l y s e v e r a l and l e a v e s / b r a c t s , 10-30 g l a n d u l a r , and and  flowering  stems; stems  o f t e n much branched throughout or above upper  cm h i g h ; g l a b r a t e below to s p a r s e l y s h o r t  s p a r s e l y s c a b r i d or s t r i g o s e , becoming densely  stipitate short  stipitate  s e s s i l e g l a n d u l a r above w i t h y e l l o w i s h h a i r s ; i n n o v a t i o n s w i t h  long p e t i o l a t e l e a v e s , p e t i o l e s 0.5-2(-6) cm blades  elliptic  to a t t e n u a t e ,  to obovate, 2-3(-5) cm  p a i r s and  one  long, l-2(-3)  cm  broad, bases  or more p a i r s of o p p o s i t e  b r a c t s , p e t i o l e s 1.5-3(-4) cm l o n g , o f t e n winged 2-3 elliptic,  2-4(-5) cm  l o n g , 1-3.5  acute to rounded, s u b e n t i r e  cm  4-6(-12)  l o n g , r a r e l y n a r r o w l y winged,  o f t e n o b l i q u e , t i p s acute, margins s u b e n t i r e  c a u l i n e l e a v e s , 2-3  mm,  cordate  to s e r r a t e ; lower or a l t e r n a t e  blades  ovate to  broad, bases u s u a l l y a t t e n u a t e d ,  tips  to i r r e g u l a r l y s e r r a t e ; upper c a u l i n e l e a v e s  duced, ovate, u s u a l l y s e s s i l e , o f t e n c o n n a t e - p e r f o l i a t e , 2-3(-6) cm  long,  1-2(-3) cm broad, except uppermost b r a c t - l i k e , o f t e n a l t e r n a t e , 1-2  cm  1-2(-5) mm  broad; v e s t i t u r e of l e a v e s  s t i p i t a t e below, densely r e s c e n c e simple and  longer  s p a r s e l y s t r i g u l o s e and v e r y  s t i p i t a t e glandular;  cm;  heads ( l - ) 3 - l l ,  + u n i s e r i a t e 5-10(-12) mm  p e r i c l i n u m densely  r a d i a t e , 10-15 l o n g , 1-2  l a n c e o l a t e , acute to acuminate; ray f l o r e t s , 5-11, tube 3-4  mm,  acute w i t h  limb 1-3  10-16(-20) mm  b l u n t t e e t h at t i p , g l a n d u l a r and  lower back of limb; and  l o n g , 3-4(-6) mm  disc florets  mm  on o u t e r  long,  infloshort  high; i n -  broad, n a r r o w l y  p a l e to b r i g h t  yellow,  broad, l i n e a r e l l i p t i c , s p a r s e l y v i l l o u s on tube  (9-)16-23(-44) t u b u l a r , 5-6  s p a r s e l y v i l l o u s at base and  mm  re-  short  s h o r t s t i p i t a t e or s e s s i l e s t i p i t a t e above;  or open corymb; peduncles 2-8  v o l u c r a l b r a c t s 6-14,  lar  cm  mm  long,  tips and  glandu-  t i p s of l o b e s ; p o l l e n diameter  185  30-50 um;  achenes dark brown to b l a c k , 4-7  s h o r t s t i p i t a t e g l a n d u l a r throughout and pappus white, Apomictic.  Type: J.H.  mm  few  b a r b e l l a t e , the l a t e r a l setae  l o n g , 1 mm  to  bifurcate hairs d i s t a l l y , .05-107 mm  l o n g , 37-48 b r i s t l e s .  2n=57.  Montana, Madison Range, Spanish Peaks, 1800 Flodman 901  Distribution:  m,  J u l y 14,  1896,  (NY).  A r n i c a g r a c i l i s i s found  s l o p e s i n open f o r e s t s at mid  on a l p i n e s c r e e s l o p e s and  to h i g h e l e v a t i o n s (1000-2400 m)  Rocky Mountains from southern B.C. Figure  wide, g l a b r o u s  and A l b e r t a to Wyoming and  rocky  i n Cascade to Oregon.  305.  Illustration:  Figure  306.  There has been much debate r e g a r d i n g the v a l i d i t y of A r n i c a g r a c i l i s . Maguire (1943) summarized h i s d i s c u s s i o n of the s p e c i e s as "a l o o s e which i s m a i n t a i n e d  as d i s t i n c t  In more r e c e n t f l o r a s B a r k l e y , 1978)  ly  ( C r o n q u i s t , 1955;  the authors  Arnica l a t i f o l i a .  from A r n i c a l a t i f o l i a w i t h  entity  some d i f f i c u l t y . "  T a y l o r & McBryde, 1978;  Ediger &  have t r e a t e d i t as a v a r i e t y of the widespread  In a subgenus (Austromontana) where t h e r e are no  absolute-  c l e a r - c u t s p e c i e s , and t h e r e a r e p o p u l a t i o n s or i n d i v i d u a l s which a r e more  o r l e s s i n t e r m e d i a t e or combine c h a r a c t e r s of two assemblage of p o p u l a t i o n s present  treatment  seems as good as most of the o t h e r s p e c i e s .  i t i s without  h e s i t a t i o n r e t a i n e d as a separate  Where A r n i c a g r a c i l i s grows i n i t s t y p i c a l stabilized bia,  o r more s p e c i e s , t h i s the  species.  form a t h i g h e l e v a t i o n s on  s c r e e s l o p e s , e s p e c i a l l y i n the Rocky Mountains of B r i t i s h  A l b e r t a , and Montana, i t i s densely  In  clumped w i t h l i t t l e  Colum-  rhizome growth  186  F i g u r e 305.  D i s t r i b u t i o n of A r n i c a g r a c i l i s discoidea (closed c i r c l e s ) .  (open c i r c l e s ) and A r n i c a  S t a r s i n d i c a t e type l o c a l i t i e s .  187  188  F i g u r e 306.  A r n i c a g r a c i l i s S-1605.  AB, Moraine Lake.  X  two-thirds.  189  190  each y e a r (S-1603, 1605, 1613).  P l a n t s branch from near ground l e v e l and  o f t e n above each p a i r o f l e a v e s , p r o d u c i n g ( F i g u r e s 307-318).  A t the o t h e r  many heads p e r stem and p e r p l a n t  extreme, i n more shaded h a b i t a t s o f t e n under  Pinus ponderosa o r Pinus c o n t o r t a , as i n s t a b i l i z e d rock s l i d e s a t Monroe Lake, BC (S-1562), the rhizomes a r e much l o n g e r and p l a n t s form open clumps more i n t h e n a t u r e o f A r n i c a l a t i f o l i a are r a r e l y branched, producing  or Arnica c o r d i f o l i a .  These p l a n t s  o n l y one head ( r a r e l y 3) per stem.  shape o f the l e a v e s , r a y f l o r e t s , and the c h a r a c t e r of the s h o r t  Y e t the stipitate  g l a n d u l a r h a i r s i s t y p i c a l o f those p l a n t s from h i g h e r e l e v a t i o n s .  (Figures  319-332). A r n i c a g r a c i l i s and A r n i c a l a t i f o l i a field. leaves  A r n i c a g r a c i l i s has a d i s t i n c t  and v i s c i d n a t u r e to the  a l s o l o o k darker  Arnica g r a c i l i s  leaves  A r n i c a g r a c i l i s a l s o has fewer, narrower, more  ( F i g u r e s 246, 247), l o n g - p e t i o l a t e o v a t e l e a v e s , and broad o v a t e ,  usually sessile bracts.  The  ( e s p e c i a l l y from  green, l o o k and f e e l rough to the touch, and a r e n o t i c e a b l y  rough under 10X m a g n i f i c a t i o n .  latifolia  i s usual-  l a t i t u d e s ) the pubescence i s n o n - g l a n d u l a r o r p r e d o m i n a t e l y so,  i t i s u s u a l l y n o t a t a l l f r a g r a n t when b r u i s e d .  pointed rays  Arnica l a t i f o l i a  and when i t does have pubescence on the l e a v e s  more n o r t h e r n  in  fragrance  caused by t h e g l a n d u l a r h a i r s on the l e a v e s .  l y glabrous  and  a r e e a s i l y d i s t i n g u i s h e d i n the  Very r a r e l y , and o n l y a t h i g h e l e v a t i o n s , does A r n i c a  have the v e r y s h o r t rhizomes t y p i c a l of A r n i c a predominance o f t r i p l o i d a p o m i c t i c  this species.  I t probably  represents  populations  gracilis. has been demonstrated  a complex o f a p o m i c t i c  races  having  o r i g i n a l l y a r i s e n from h y b r i d i z a t i o n between the d i p l o i d A r n i c a l a t i f o l i a and the t e t r a p l o i d races o f A r n i c a m o l l i s o r A r n i c a d i v e r s i f o l i a b o t h i n Subgenus C h a m i s s o n i s ) .  ( t h e l a t t e r two  The g l a n d u l a r pubescence o f l e a v e s and stems,  191  s h o r t s c a l y rhizomes, b l a c k , o f t e n g l a n d u l a r  achenes, and  are a l l c h a r a c t e r i s t i c s s i m i l a r to the l a t t e r are many p o p u l a t i o n s  which are s e p a r a t e d  two  o f t e n tawny pappus  species.  only with  Indeed,  difficulty  there  from these  s p e c i e s , e s p e c i a l l y when u s i n g h e r b a r i u m specimens.  Examples are Moraine  Lake, B a n f f  (S-1605) grows w i t h  N a t i o n a l Park, AB,  where A r n i c a g r a c i l i s  s m a l l form of A r n i c a d i v e r s i f o l i a d i s t i n g u i s h e d i n the f i e l d  by  (S-1606) on a s c r e e  the more v i s c i d  slope.  The  l e a v e s , darker  a  latter is  pappus, and  less  branched h a b i t . Artificial  c r o s s e s between A r n i c a l a t i f o l i a  d i v e r s i f o l i a have been m o s t l y u n s u c c e s s f u l . p o o r l y i n c u l t i v a t i o n and Arnica l a t i f o l i a s i f o l i a has and  as  a few  A r n i c a m o l l i s has  s u r v i v e beyond a few  weeks.  l y produce an A r n i c a g r a c i l i s - l i k e h y b r i d have not There may  grown v e r y  achenes.  However o n l y one  has been s u c c e s s f u l , p r o d u c i n g  Arnica  the male p a r e n t  have produced no  grown w e l l i n c u l t i v a t i o n .  However, t h e s e d i d not  A r n i c a m o l l i s or  c r o s s e s between i t as  the female p a r e n t  Arnica l a t i f o l i a  and  Arnica diver-  c r o s s between  a few So,  and  this,  v i a b l e achenes.  e f f o r t s to  artificial-  been s u c c e s s f u l .  a l s o be confusion' between A r n i c a g r a c i l i s and  Arnica rydbergii  (Subgenus A r c t i c a ) , where they a r e growing i n s i m i l a r h a b i t a t s .  However,  A r n i c a r y d b e r g i i u s u a l l y has more s e s s i l e c a u l i n e l e a v e s which have a  very  d i s t i n c t p a i r of l a t e r a l v e i n s n e a r l y p a r a l l e l  the  l e a v e s a t r i - n e r v e d appearance. o r a n g e - y e l l o w , and  The  to the m i d - r i b ,  giving  pappus i s pure w h i t e , the r a y s are  the achenes a r e d e n s e l y  h i r s u t e throughout,in  darker  Arnica  rydbergii.  5.  A r n i c a cernua T. H o w e l l , F I . NW.  A r n i c a c h a n d l e r i Rydb., N. Co.,  Hoopa I n d i a n  i s o t y p e s MO,  UC).  Am.  Am.  373.  1903.  F I . 34:339. 1927.  Reservation,  June 1901,  H.P.  TYPE:  C a l i f o r n i a , Humboldt  C h a n d l e r 1298  (holotype  NY;  192  F i g u r e s 307-318.  V a r i a t i o n i n b a s a l l e a v e s of a s i n g l e  population  t y p i c a l h i g h - e l e v a t i o n A r n i c a g r a c i l i s S-1603, AB Lake L o u i s e .  194  F i g u r e s 319-332.  V a r i a t i o n i n b a s a l l e a v e s of a s i n g l e  population  of l o w e r - e l e v a t i o n A r n i c a g r a c i l i s S-1562, BC, Monroe Lake.  195  196  Rhizomatous p e r e n n i a l herb forming l a r g e open p o p u l a t i o n s clumps of s t e r i l e b a s a l r o s e t t e s and cm growth/year, ( l - ) 2 - 3 ( - 5 ) mm nodes, and ly  crowded s c a l e s and  flowering  stems; rhizomes  base or h i g h e r , curved  (1-)4-15(-20)  o l d l e a f bases toward summit, simple  stems s i n g l e or few,  simple  mm  or  sparse-  t h i c k , mostly  or r a r e l y branched near  (5-)10-30(-45) cm h i g h , g l a b r a t e to s p a r s e l y v i l l o u s w i t h i n -  h a i r s , p u r p l i s h or r e d d i s h at base or throughout; i n n o v a t i o n s w i t h  long p e t i o l a t e leaves, p e t i o l e s (l-)4-6(-10) (-5)  tight  t h i c k , naked except p a i r s of brown s c a l e s at  to much branched e s p e c i a l l y toward summit; r o o t s 2-3  unbranched, s u c c u l e n t ;  or  cm broad, e l l i p t i c  to ovate o r r o t u n d ,  cm,  blades  4-6(-7) cm  long  crenate,  or l a c i n i a t e and  o f t e n undulate;  2-4  bases rounded to subcordate,  rounded to acute, margins e n t i r e or r e g u l a r l y or i r r e g u l a r l y s e r r a t e ,  4-8  tips  dentate,  c a u l i n e l e a v e s 2-3(-5) p a i r s , the  l a r g e s t p a i r u s u a l l y near base of stem, l o n g p e t i o l a t e , p e t i o l e s (-l)2-4(-6) cm  l o n g , blade (0.5-)2-4(-6) cm  long,  ( l - ) 2 - 4 ( - 5 ) cm broad, p e t i o l e s and  s i z e reduced upward, upper-most b r a c t - l i k e , o f t e n b r o a d l y a l t e r n a t e ; shape and margins of c a u l i n e l e a v e s l e a v e s , a l t h o u g h o f t e n more deeply  s e s s i l e and  s i m i l a r to t h a t of  i n c i s e d , leaves  t h i c k and  blade  often  innovation  succulent  and  o f t e n r e d d i s h or p u r p l i s h e s p e c i a l l y along v e i n s of lower s u r f a c e ; herbage g l a b r a t e to s c a b r i d e s p e c i a l l y on margins and s c a b r i d and  densely  heads, nodding i n bud,  (2-)5-15(-22) cm p i l o s e and  s c a b r i d ; heads 15-25  acute to obtuse; ray f l o r e t s  and  mm  l o n g , 4-8  base of limbs  mm  densely  mm  mm  l o n g , 3-5  (5-)7-9(-14),  an  nodding or e r e c t a t a n t h e s i s , ped-  l o n g , upper peduncle and  7-12(-15), + u n i s e r i a t e , 10-15  limb 10-20  densely  p i l o s e on upper s u r f a c e ; i n f l o r e s c e n c e u s u a l l y a s i n g l e head or  open corymb of 3-5 uncles  a l o n g main v e i n s , r a r e l y  tall, mm  p e r i c l i n u m s p a r s e l y to radiate; i n v o l u c r a l bracts  broad, b r o a d l y  dark y e l l o w ,  broad, l i n e a r e l l i p t i c w i t h  lanceolate,  tube 4-6 1-3  mm  long,  t e e t h at t i p , tubes  s h o r t v i l l o u s ; d i s c f l o r e t s 24-57, t u b u l a r 10-15  mm  197  l o n g , s p a r s e l y to densely  v i l l o u s at base and  l o b e s ; p o l l e n diameter 35-50 Um; s p a r s e l y to u s u a l l y densely  achenes grey, 6-8  mm  l o n g , 1+ mm  broad,  .10-.15 mm  b a r b e l l a t e to subplumose  l o n g ; s e x u a l d i p l o i d , 2n=38.  Type: Oregon, Josephine County, near Waldo, on dry banks, base of Mountains, June 1884,  Distribution:  T. Howell 166  of the Coast Ranges and  c o u n t i e s of Southwestern Oregon (Coos, Curry, t i e s of Northwestern C a l i f o r n i a  Coast  (ORE).  A r n i c a cernua i s r e s t r i c t e d to s e r p e n t i n e  e l e v a t i o n s (400-1800 m)  on  h i r s u t e w i t h b i f u r c a t e h a i r s throughout or from  m i d - p o i n t d i s t a l l y ; pappus pure w h i t e , 41-60 b r i s t l e s , the l a t e r a l s e t a e  s p a r i n g l y toward summit and  s o i l s at  mid-  S i s k i y o u Mountains i n t h r e e  and  Josephine) and  three  (Del Norte, S i s k i y o u , and Humboldt).  counFigure  333.  Illustration:  Figure  This d i s t i n c t  334.  s p e c i e s i s one  common where i t does occur.  of the r a r e s t A r n i c a s , a l t h o u g h i t i s o f t e n  Populations  occupy rock c r e v i c e s or r u b b l e  open exposed h a b i t a t s i n open f o r e s t s of Pinus j e f f r e y i , Garrya f r e m o n t i i , and  Arctostaphylos  spp.,  Calocedrus  decurrens,  or at lower e l e v a t i o n s are  scat-  t e r e d i n open f o r e s t s of P i n u s ponderosa, Pseudotsuga m e n z i e s i i , and cedrus decurrens.  I t i s not known to occur  A r n i c a cernua i s o b v i o u s l y s i m i l a r to t h a t s p e c i e s , but other  o f f serpentine  d i f f e r i n g from i t i n a number of  which are u s u a l l y p u r p l e of A r n i c a l a t i f o l i a  The  being  thick, leathery  f l e s h y nor  most  characters  or red are d i s t i n c t i v e i n A r n i c a cernua.  are never as t h i c k and  Calo-  soils.  d e r i v e d from A r n i c a l a t i f o l i a ,  than t h e i r d i f f e r e n t e c o l o g i c a l n i c h e s .  in  leaves, The  leaves  are they o f t e n more  198 F i g u r e 333.  D i s t r i b u t i o n of A r n i c a s p a t h u l a t a A r n i c a cernua ( c l o s e d c i r c l e s ) . localities.  (open c i r c l e s )  and  S t a r s i n d i c a t e type  199  200  F i g u r e 334.  H a b i t of A r n i c a cernua S-1386. X two-thirds.  OR,  Josephine  201  202  than s l i g h t l y s u f f u s e d w i t h r e d o r p u r p l e .  The nodding o r cernuous buds,  from which A r n i c a cernua gets i t s name, may o r may not become e r e c t a t anthesis.  No specimens o f A r n i c a l a t i f o l i a have been seen w i t h nodding buds,  Both the p h y l l a r i e s ( F i g u r e s 130, 132) and the rays  ( F i g u r e s 139, 140) o f  A r n i c a cernua a r e l a r g e r and broader than those o f A r n i c a l a t i f o l i a .  The  d i s t i n c t t e e t h a t t h e t i p o f t h e r a y f l o r e t s of A r n i c a cernua d i f f e r from the more t r u n c a t e  t i p s o f the A r n i c a l a t i f o l i a  A r n i c a cernua a r e much more densely s m a l l e r achenes o f A r n i c a The  The achenes o f  h i r s u t e d i s t a l l y o r throughout than t h e  latifolia.  s p e c i e s c o n s i s t s o f two f a i r l y d i s t i n c t ecotypes, a s m a l l  ( F i g u r e 336) from exposed rocky in  ray f l o r e t s .  slopes a t higher  c o l l e c t i o n s from Hoopa I n d i a n R e s e r v a t i o n  form  e l e v a t i o n s (750-1450 m) as  and Horse Mountain, both from  Humboldt Co., CA ( S t r a l e y 1396, 1772) and a l a r g e r form ( F i g u r e 337) from lower e l e v a t i o n s (400 m) i n open f o r e s t h a b i t a t s as from near O ' B r i e n , J o s e p h i n e Co., OR ( S t r a l e y 1386).  The s m a l l e r  ecotype i s b a r e l y  rhizomatous,  forming dense clumps w i t h many stems, which o f t e n branch above producing o r more heads p e r stem.  The l e a v e s  tend  3-5  to be s h o r t e r - p e t i o l e d and a r e r e l a -  t i v e l y s m a l l ( F i g u r e s 339-355).  The l a r g e r ecotype i s s t r o n g l y rhizomatous  forming l a r g e " l o o s e " c o l o n i e s .  The stems a r e u s u a l l y produced s i n g l y , a r e  unbranched and c o n s i s t e n t l y produce a s i n g l e head p e r stem.  Within  popula-  t i o n s o f t h e l a r g e r ecotype t h e r e i s a marked d i f f e r e n c e between p l a n t s growing i n n e a r l y f u l l p e t i o l e d leaves  sun i n t h a t they have t h i c k e r , s m a l l e r ,  shorter  ( F i g u r e s 356-365) compared to the t h i n n e r , l a r g e r , l o n g e r  p e t i o l e d l e a v e s o f p l a n t s growing nearby i n r a t h e r dense shade ( F i g u r e s 366370).  Although Rydberg (1927) c o n s i d e r e d  cies, recognizing  the s m a l l ecotype a separate  spe-  i t as A r n i c a c h a n d l e r i , t h e r e a r e many c o l l e c t i o n s w i t h  203  F i g u r e s 335-338. F i g u r e 335.  H a b i t a t and h a b i t of A r n i c a cernua.  H a b i t a t a t summit o f Hoopa-Redwood V a l l e y Road, e l e . 750 Hoopa I n d i a n R e s e r v a t i o n , Humboldt Serpentine  soils  lambertiana,  F i g u r e 336.  i n open f o r e s t of Pinus j e f f r e y i ,  Calocedrus  Arctostaphylos  Co., CA, S t r a l e y 1772.  decurrens,  Pinus  Arbutus m e n z i e s i i , and  spp.  Dense clump w i t h two open heads and many buds at above locality.  F i g u r e 337.  Larger  form w i t h one head per stem i n dense f o r e s t of  P i n u s ponderosa, Calocedrus spp. F i g u r e 338.  decurrens,  a t O'Brien, Josephine Co., OR,  Seed head from above p o p u l a t i o n .  and  Arctostaphylos  S t r a l e y 1386.  m,  205  F i g u r e s 339-355.  V a r i a t i o n i n b a s a l r o s e t t e l e a v e s of A r n i c a S-1772, CA, Humboldt Co., s m a l l ecotype.  cernua  206  207  Figures  356-365.  V a r i a t i o n i n b a s a l r o s e t t e l e a v e s of A r n i c a  cernua  S-1743, OR, Josephine Co., l a r g e ecotype, from p l a n t s growing  i n f u l l sun.  209  Figures  366-370.  V a r i a t i o n i n b a s a l r o s e t t e l e a v e s of A r n i c a  cernua  S-1743, OR, Josephine Co., l a r g e ecotype, from p l a n t s growing  i n shade.  210  211  i n t e r m e d i a t e c h a r a c t e r s , so t h e r e seems t o be no reason populations  separate  to g i v e  these  taxonomic r e c o g n i t i o n .  There are a number of e x c e p t i o n a l c o l l e c t i o n s of t h i s s p e c i e s , i n c l u d i n g extremely  narrow-leaved specimens from Red  Mountain, Curry Co.,  3442 (ORE), w i t h the l a r g e s t c a u l i n e l e a f b l a d e s o n l y 1.5 broad  and  (ORE)  with  and  a c o l l e c t i o n from Tennessee Pass, Josephine  deeply  Curry  Co.,  the lower c a u l i n e l e a f b l a d e s broader lobed. OR,  Leach 2879 (ORE), a l l l e a v e s and  Savage  T h i s p o p u l a t i o n may  Two  represent a  (ORE)  c o n s i s t s of t h r e e f l o w e r i n g stems later  However, the t h i r d specimen on the sheet has  two  an  c h a r a c t e r f o r the genus.  A r n i c a d i s c o i d e a Bentham. PL. Hartw. 319.  A r n i c a p a r v i f l o r a A.  Gray. Proc. Am.  Humboldt Co.,  c h a p p a r a l , 1867,  ( h o l o t y p e , G;  isotype,  1849.  Acad. 7:363. 1867.  Geol. Surv. C a l i f . , H.H.  TYPE:  TYPE:  California,  Bolander  Oregon, Hood R i v e r , 1884,  F l . N. Am.  1:381.  1955.  1884.  B a r r e t t , s..n. (G) .  A r n i c a d i s c o i d e a Bentham v a r . e r a d i a t a (A. Gray) C r o n q u i s t . Vase. P I . 5:49.  6051.  KEW).  A r n i c a c o r d i f o l i a Hook, v a r . e r a d i a t a A. Gray. Syn.  NW.  local  of the pubescent  s e t s of d i s t i n c t whorls of t h r e e l e a v e s r a t h e r than o p p o s i t e p a i r s ,  6.  Creek,  throughout  of the p l a n t s a r e s i m i l a r to most of t h e  c o l l e c t i o n s of the s p e c i e s .  unusual  cm)  cordifolia.  Type specimen, Howell 166  and one b a s a l r o s e t t e .  cm  s.n.  than long (3.5  stems are scabrous  r e f l e c t p o s s i b l e h y b r i d i z a t i o n w i t h one  species, l i k e l y Arnica The  OR,  0.5  In an u n u s u a l l y pubescent specimen from Head S l i d e  g i v i n g the p l a n t s a grey appearance. ecotype o r may  Leach  cm l o n g and  Co.,  (5 cm)  OR,  Pac.  212  A r n i c a g r a y i H e l l e r . Muhlenbergia 1:5. 1884,  1900.  TYPE:  Oregon, Hood R i v e r ,  B a r r e t t , _s.n. (G) .  A r n i c a f a l c o n a r i a Greene. Ottawa Nat. Klickitat  Co.,  F a l c o n V a l l e y , June 27, 1892.  A r n i c a a l a t a Rydb. N. Am. 1865,  23:215. 1910.  J . T o r r e y 258a  A r n i c a p a r v i f l o r a A.  F I . 34:342. 1927.  W.  TYPE:  TYPE: Suksdorf  Washington, 1617  C a l i f o r n i a , Yosemite,  (NY).  Gray subsp. a l a t a  (Rydb.) Maguire. B r i t t o n i a 4:455.  A r n i c a d i s c o i d e a Bentham v a r . a l a t a (Rydb.) C r o n q u i s t . Contr. 5:102.  1943,  Dudley Herb.  1958.  A r n i c a sanhedrensis Co.,  (ND).  Rydb. N. Am.  south of Mt.  isotype,  Sanhedrin,  F I . 34:342. 1927. J u l y 25, 1902,  TYPE:  C a l i f o r n i a , Lake  H e l l e r 5985 ( h o l o t y p e ,  NY;  POM).  Rhizomatous p e r e n n i a l herb; rhizomes 5-15(-22) cm growth/year, 2-5 t h i c k , naked except  f o r p a i r s of brown s c a l e s at nodes and  o l d l e a f bases toward summit, forming r o s e t t e s and  unbranched; stems s i n g l e o r few, (15-)20-45(-65) cm  tall,  crowded s c a l e s and  l a r g e p o p u l a t i o n s of s t e r i l e  f l o w e r i n g stems; r o o t s + s u c c u l e n t , 2-3  mm  mm  thick,  basal  relatively  simple o r branched near base o r  throughout,  s p a r s e l y to d e n s e l y s t r i g o s e , v i l l o u s , and  long  s t i p i t a t e g l a n d u l a r throughout; i n n o v a t i o n s w i t h 4-8(-12) l o n g p e t i o l a t e l e a v e s , p e t i o l e s ( l - ) 4 - 6 ( - 1 0 ) cm l o n g , b l a d e s , ovate o r b r o a d l y ovate  to  broadly  sub-  l a n c e o l a t e , 3-5(-9) cm l o n g , 2-4(-5) cm broad, bases c o r d a t e ,  c o r d a t e , to rounded, o r r a r e l y a t t e n u a t e , t i p s acute s u b e n t i r e to u s u a l l y r e g u l a r l y s e r r a t e , dentate, s e r r a t e or dentate;  to rounded, margins  o r c r e n a t e , or r a r e l y  doubly  c a u l i n e l e a v e s (2-)3-5(-6) p a i r s , o f t e n reduced b a s a l l y  and becoming b r a c t - l i k e toward summit, lower c a u l i n e l e a v e s u s u a l l y the l a r g e s t , l o n g p e t i o l a t e , p e t i o l e s 2-6(-10) cm l o n g , o f t e n winged 2-5(-14)  mm,  213  b l a d e s (3-)4-7(-9) cm p e t i o l e d , and  l o n g , 2-6  cm wide, upper p a i r s p r o g r e s s i v e l y  often subalternate,  t h a t of i n n o v a t i o n s ;  shape and margins of c a u l i n e l e a v e s  v e i n s , and  s p a r s e l y long  stipitate  throughout; i n f l o r e s c e n c e s an open corymb of 3-7(-20) heads, r a r e l y  subradiate,  12-15  l o n g , 2-3(-5) mm  mm  high;  i n v o l u c r a l b r a c t s , 7-12  wide, narrowly to b r o a d l y  f l o r e t s 19-45, t u b u l a r , 8-12 rarely + ligulate,  to 20 mm  mm  mm  l a n c e o l a t e , acute to acuminate;  wide, s p a r s e l y  a t base and  f a c e of l o b e s ; p o l l e n diameter 45-60 um; wide, s p a r s e l y  + u n i s e r i a t e , 10-15  l o n g , m a r g i n a l f l o r e t s o f t e n a m p l i a t e or  l o n g , 5 mm  sparsely s t i p i t a t e glandular  lar  like  v e s t i t u r e of herbage u s u a l l y moderately to d e n s e l y  scabrous and v i l l o u s , e s p e c i a l l y along glandular  shorter  to d e n s e l y v i l l o u s  s p a r i n g l y upward, and  on outer  achenes dark grey, 6-8  to densely h i r s u t e w i t h b i f u r c a t e h a i r s , and  mm  and sur-  long, 1  stipitate  mm  glandu-  throughout, e s p e c i a l l y d i s t a l l y ; pappus w h i t e , b a r b e l l a t e , of 42-64  bristles, lateral  Type:  setae  .05-.15 mm  C a l i f o r n i a , Monterey Co.,  Distribution:  long; apomictic;  2n=38,  76.  Monterey, i n f o r e s t , Hartweg 1805  (KEW).  T h i s s p e c i e s o c c u p i e s a wide v a r i e t y of h a b i t a t s from open  c h a p p a r a l t o c o n i f e r f o r e s t s , near sea l e v e l to 1800 ton ( K l i c k i t a t  County) to Southern C a l i f o r n i a  Illustration:  Figure  m,  from Southern Washing-  (Orange County).  (Figure  305).  371.  There has been much c o n f u s i o n  and  d i f f e r e n c e of o p i n i o n i n the  treatments  o f A r n i c a d i s c o i d e a , the most widespread and most v a r i a b l e of the d i s c o i d species.  A r n i c a d i s c o i d e a , as p r e s e n t e d here, c o n t a i n s  by Maguire (1943) as t h r e e 2 subspecies),  and  elements  recognized  species, Arnica discoidea, Arnica p a r v i f l o r a (with  Arnica grayi.  Ediger  species, Arnica discoidea, with three  & Barkley  varieties.  (1978) t r e a t these as  one  214  F i g u r e 371.  A r n i c a d i s c o i d e a S-1775.  CA,  Mendocino Co.  X  two-thirds.  215  216 At the northern l i m i t of t h e i r d i s t r i b u t i o n there are populations formerly known as Arnica g r a y i H e l l e r , from K l i c k i t a t Co., WA  (S-1825),  Wasco Co., OR (S-1821) , and Hood River Co., OR, B a r r e t t , s.xi.  (G). , growing  with Arnica c o r d i f o l i a .  The leaves of these northern populations tend to be  more densely s t r i g o s e and with somewhat l e s s cordate bases than i n most of the southern populations.  However, there are numerous intermediates through-  out the range of the species, as i n c o l l e c t i o n s from San Mateo Co., CA, Keck 1775  (RSA). V a r i a b i l i t y i n l e a f s i z e , shape, and e s p e c i a l l y i n margins can be seen i n  many populations  (Figures 376-385) and i s p a r t i c u l a r l y notable i n a c o l l e c -  t i o n from Mt. Eddy, S i s k i y o u Co., CA, Copeland 3909 (POM). Many populations throughout the range of the species have marginal l i a t e f l o r e t s (Figure 155) or elongated r a y - l i k e f l o r e t s (Figures 154,  amp156).  This condition i s e s p e c i a l l y pronounced i n c o l l e c t i o n s from Santa Barbara  Co.,  CA, Smith 8330 (SBM), T r i n i t y Co., CA, Baker 328 (HSC), and Eldorado Co.,  CA,  Peirson 9517  (RSA).  S u p e r f i c i a l l y , these c o l l e c t i o n s look very much l i k e  Arnica c o r d i f o l i a and are often m i s i d e n t i f i e d as such.  However, none of the  c o l l e c t i o n s has true female ray f l o r e t s , although they are l i k e l y often funct i o n i n g as female.  Even the elongated marginal f l o r e t s are u s u a l l y + tubular  (Figures 154, 156) and have f u n c t i o n a l or abortive anthers. Arnica discoidea d i f f e r s from Arnica c o r d i f o l i a i n features other than an absence of ray f l o r e t s .  Arnica discoidea i s u s u a l l y more branched, has more  heads (up to 20/stem), i s l e s s strongly rhizomatous, and u s u a l l y prefers d r i e r h a b i t a t s than Arnica c o r d i f o l i a .  The d i s c o i d species also tends to be l e s s  pubescent, or at l e a s t the pubescence tends to be shorter than that of Arnica cordifolia.  Trends toward, and d i f f e r e n c e s between Arnica spathulata and  217  F i g u r e s 372-375.  Habitat  and h a b i t of A r n i c a d i s c o i d e a and  Arnica  spathulata. F i g u r e 372.  Habitat  of A r n i c a d i s c o i d e a S-1825, WA,  Klickitat and  Klickitat  Co.,  R i v e r Gorge, open f o r e s t of Pinus ponderosa  Pseudotsuga m e n z i e s i i .  F i g u r e 373.  same, f l o w e r i n g stems and  basal rosettes.  F i g u r e 374.  same, t h r e e f l o w e r i n g heads and  F i g u r e 375.  A r n i c a s p a t h u l a t a S-1385, OR,  two  seed heads.  Josephine Co.,  O'Brien.  218  219  F i g u r e s 376-385.  V a r i a t i o n i n b a s a l l e a v e s of A r n i c a S-1789, CA, Marin County.  discoidea  220  221  A r n i c a d i s c o i d e a are d i s c u s s e d under A r n i c a There does not  seem to be any  chromosome number, and evenly  c o r r e l a t i o n between d i s t r i b u t i o n ,  s i z e of heads, whether l e a v e s are b a s a l l y d i s p o s e d  d i s t r i b u t e d along  the stem, or achene and  the d i s c o i d p o p u l a t i o n s . coid species recognized as one  spathulata.  pappus f e a t u r e s i n any  These f e a t u r e s have been used to separate i n previous  treatments.  polymorphic s p e c i e s w i t h both d i p l o i d ,  I t seems best  s e x u a l and  of  the  dis-  to t r e a t  tetraploid,  or  these  apomictic  races.  7.  A m i ca. spathuXa tci Greens,  A r n i c a eastwoodiae Rydb. N. Am. Norte Co.,  Pittonia. 3il03.  1896.  F l . 34:343. 1927.  Gasquet, F r e n c h H i l l ,  September 14,  TYPE: 1912,  California,  Del  A. Eastwood 2211  A r n i c a s p a t h u l a t a Greene subsp. eastwoodiae (Rydb.) Maguire.  (NY).  B r i t t o n i a 4:458.  1943. A r n i c a s p a t h u l a t a Greene v a r . eastwoodiae (Rydb.) E d i g e r Fl.  I I . 10:43.  F l . 34:343. 1927.  Mountains, dry western s l o p e , J u l y 11, POM,  and  Oregon, Cascade  C u s i c k 2873 ( h o l o t y p e ,  s h o r t woody caudex 0.5-1.5  cm h i g h , 2-5(-7) cm below s u r f a c e of ground, producing  brown s c a l e s at nodes and m i t , forming  stems; r o o t s r e l a t i v e l y few  and  cm  vertical  cm/year, g e n e r a l l y naked except p a i r s of  crowded s c a l e s and  l a r g e open p o p u l a t i o n s  simple  NY;  US).  h o r i z o n t a l rhizomes ( l - ) 3 - 7 ( - 1 4 )  o r few,  TYPE:  1902,  Rhizomatous p e r e n n i a l herb o f t e n forming broad, 1-2  Am.  1978.  A r n i c a c u s i c k i i Rydb. N. Am.  i s o t y p e s , MO,  & B a r k l e y . N.  some o l d l e a f bases toward sum-  of s t e r i l e b a s a l r o s e t t e s and  s u c c u l e n t , 1-3  mm  flowering  i n diameter; stems s i n g l e  to branched below and above, (15-)25-45(-55) cm t a l l ,  sparsely  222  to densely v i l l o u s and l o n g s t i p i t a t e g l a n d u l a r throughout; i n n o v a t i o n s 4-6(-10) l o n g p e t i o l a t e l e a v e s , p e t i o l e s 2-7 2-7(-12) mm  broad, b l a d e s  with  cm l o n g , w i n g l e s s o r w i t h wings  (2-)4-6(-8) cm l o n g , 1-3.5  cm wide, s p a t h u l a t e to  e l l i p t i c o r obovate, bases a t t e n u a t e or r a r e l y rounded, t i p s rounded to a c u t e , margins e n t i r e to c o a r s e l y and l e a v e s 3-6  p a i r s , b a s a l l y disposed  subopposite,  reduced  evenly s e r r a t e or c r e n a t e ; c a u l i n e  to + evenly d i s t r i b u t e d along  stem, o f t e n  and b r a c t - l i k e upward, where f r e q u e n t l y a l t e r n a t e , w i t h  p e t i o l e s as wide as or wider than b l a d e , l a r g e s t p a i r of c a u l i n e l e a v e s l o n g e s t p e t i o l e s , u s u a l l y near base, p e t i o l e s ( l - ) 3 - 5 ( - 8 ) cm. mm,  blades  wings 2-10(-20)  (2-)3-6(-8) cm l o n g , ( l - ) 2 - 4 ( - 5 ) cm wide, shape and margins as  w i t h l e a v e s of i n n o v a t i o n s , v e i n s u s u a l l y prominent and  r e d d i s h or p u r p l i s h  e s p e c i a l l y on lower s u r f a c e ; v e s t i t u r e of herbage p i l o s e and l o n g  stipitate  g l a n d u l a r throughout, e s p e c i a l l y along v e i n s , margins o f t e n c i l i a t e ; rescence  l o n g , upper peduncles and p e r i c l i n u m l o n g p i l o s e and  t a t e g l a n d u l a r ; heads 15-25  mm  h i g h , d i s c o i d , r a r e l y w i t h ampliate  i n v o l u c r a l b r a c t s 7-13(-17), + u n i s e r i a t e , 10-15  wide, l a n c e o l a t e to b r o a d l y l a n c e o l a t e , obtuse to acute; goblet-shaped v i l l o u s and  to t u b u l a r , 8-11  mm  mm  l a t e , the l a t e r a l incompatible  l o n g , 1 mm setae  distally,  marginal mm  f l o r e t s 18-52, densely  s p a r s e l y upward and on  lobes;  short  few o r no b i f u r c a t e h a i r s  wide, pappus pure white,  .07-.10 mm  stipi-  l o n g , 1-4  achenes b l a c k , dark brown, o r dark grey,  s t i p i t a t e throughout o r from m i d - p o i n t d i s t a l l y , 6-9  mm  l o n g , dark y e l l o w , s p a r s e l y to  s t i p i t a t e g l a n d u l a r toward base and  p o l l e n diameter 35-55 um;  Type:  inflo-  a s i n g l e head o r an open corymb of 3-5(-25) heads, e r e c t ; peduncles  (2-)4-8(-15) cm  florets;  with  37-56 b r i s t l e s , b a r b e l -  l o n g ; a p o m i c t i c p o l y p l o i d and  self-  s e x u a l d i p l o i d ; 2n=38, 76.  Oregon, Douglas County, G l e n d a l e , June 3, 1887,  T. Howell 1200  (US).  223  Distribution:  A r n i c a s p a t h u l a t a occurs on open s e r p e n t i n e o u t c r o p s  f o r e s t s at low  to m i d - e l e v a t i o n s  from Douglas and California.  Curry Counties  Figure  Illustration:  (180-1400 m) i n SW  of Coast  and  o r open  Cascade Ranges  Oregon to S i s k i y o u County i n NW  333.  Figure  386.  T h i s , the r a r e s t of the s p e c i e s i n s e c t i o n E r a d i a t a e i s known from about two  dozen p o p u l a t i o n s l a r g e l y r e s t r i c t e d  to s e r p e n t i n e s o i l s .  Where i t does  occur the p o p u l a t i o n s o f t e n c o n s i s t of s e v e r a l dozen to hundreds of uals.  individ-  Populations or i n d i v i d u a l s are q u i t e v a r i a b l e i n s i z e , branching  l e a f shape, and pubescence.  There i s a s m a l l ecotype to which the name A r n i c a  eastwoodiae Rydb. has been a s s i g n e d . from French H i l l  Rd.,  habit,  These p o p u l a t i o n s , as i n c o l l e c t i o n s  Gasquet, Del Norte Co.,  CA,  (S-1751, p r o b a b l y  the  l o c a l i t y of A r n i c a eastwoodiae) a r e s m a l l e r , have narrower l e a v e s w i t h  type less-  winged p e t i o l e s ( F i g u r e s 387-398), narrower heads, and  s h o r t e r pubescence  most o t h e r p o p u l a t i o n s of the s p e c i e s .  on open  s l o p e s i n mixed f o r e s t s of Pinus ponderosa o r P i n u s j e f f r e y i . t i o n s , with t a l l e r 399-407), and  They a r e found  a t t e n u a t a , Calocedrus  decurrens,  l o n g g l a n d u l a r h a i r s throughout.  Co.,  on more densely  OR,  (S-1380, 1381,  1740).  which a r e i n t e r m e d i a t e  popula-  T y p i c a l p o p u l a t i o n s of  this  Selma, a l l i n  These p o p u l a t i o n s are u s u a l l y found  There are numerous p o p u l a t i o n s  i n some o r a l l of the above c h a r a c t e r s , i n c l u d i n g c o l -  l e c t i o n s from Gasquet, Del Norte Co., (S-1385).  Pinus  f o r e s t e d h i l l s i d e s of Pinus ponderosa, Pseudotsuga m e n z i e s i i ,  Arbutus m e n z i e s i i , and Quercus k e l l o g g i i .  OR,  and  l e a v e s , u s u a l l y winged p e t i o l e s ( F i g u r e s  l a r g e r ecotype a r e c o l l e c t i o n s from K i r b y , Cave J u n c t i o n , and Josephine  serpentine  At the o t h e r extreme are a m a j o r i t y of  stems, broader  than  CA,  (S-1390) and O'Brien,  Josephine  Co.,  224  F i g u r e 386.  A r n i c a s p a t h u l a t a S-1740.  OR,  Josephine Co.  X  two-thirds.  225  226  F i g u r e s 387-398.  V a r i a t i o n i n b a s a l r o s e t t e l e a v e s of A r n i c a S-1751, CA, D e l N o r t e Co., s m a l l e c o t y p e .  spathulata  227  228  Figures  399-407.  V a r i a t i o n i n basal rosette leaves S-1740, OR, Josephine Co., l a r g e  of A r n i c a ecotype.  spathulata  229  230 Occasionally Pleasant leaves seen OR,  Creek,  are  in  Leach  usually  2874  has  which  more  achenes, ulation US).  and has  The  been  often  in  the  been  resembles  some A r n i c a  monograph  of  ymy u n d e r  Arnica  Arnica  the  the  the  spathulata  Ampliate this  species  from  Del  ampliate  or than  Norte  or  is  its  differs its  rhizomes.  cusickii known,  Rydb., only  Gold  from  of  cauline  character Basin,  was  Curry  leaves,  not  Co.,  and  un-  "dry  western  leaves  rhizomes,  herein  of  with  Cascade  variable,  but  the  cordate  bases the  1978),  it  is  characters  as  a  it  florets  seems that  seem t o  one  specimen,  Monumental,  has  been  as  growing with  recent  in  the  synon-  than  the  leaves,  to  fit  best  in  species. b e much r a r e r  Tracy  seen w i t h  typical  closely  most  listed  Only  reported  in  other  synonym of  MO,  the  that  achenes,  POM,  (NY,  pop-  of  so  marginal  discoidea.  are  darker  intermediate  slope  so much  the  and  retained  ray-like  north  in  2873  it  leaves  larger,  A somewhat Cusick  from which  spathulate  heads,  cauline  However,  discoidea  its  disposed  and B a r k l e y ,  was  in  taller  (Ediger  CA, It  in  Arnica  specimens  Arnica  petioles  cauline  several  elongated  a specimen  This  from  disposed  specimens,  is  the  The  vestiture,  in  former  of  not  discoidea.  and  collection  discoidea  genus  in  blades.  resembles  red,  Arnica  basally  Co.,  florets.  The  Oregon."  of  (MO),  the  basally  closely  purple  locality  than  as  vestiture.  character  Southern  in  strongly  most  called  wider  An u n u s u a l  derived.  character  especially  often  has  populations,  O R , Hammond 2 3 0  short-pilose  exact  Mountains,  Co.,  Arnica.  spathulata  certainly  large-leaved  are  (ORE)  dense,  are  and  other  Arnica  the  Jackson  broad  any  in  19424 long  discoid  in  (UC)  ray-like  plants.  231 Subgenus C a l a r n i c a subgen.  nov.  Herbae perennes e r e c t a e  a c a u d i c e l i g n o s u b t e r r a n e o ; rhizomata  radices  succulentae,  v e r t i c a l e s ; caules  plantae  dense v e l s p a r s i m s t i p i t a t o - g l a n d u l o s a e  pappus a l b u s v e s u b i n f u s c a t u s tum  numerus gamatae  Erect perennial + v e r t i c a l and  ramosi; f o l i a c a u l i n a o v a t a ,  carentes; sessile;  et p i l o s a e ; c a p i t u l a e r a d i a t a ;  suffusus, barbellatusve  subplumosus; chromosoma-  n=19.  herbs from woody caudex, non-rhizomatous; r o o t s  unbranched; l a c k i n g s t e r i l e b a s a l r o s e t t e s  fleshy,  (innovations);  flow-  e r i n g stems b r a n c h i n g from base, o r s p a r i n g l y to much-branched above; stems equably l e a f y , c a u l i n e l e a v e s  s e s s i l e , ovate, margins e n t i r e to s e r r a t e , bases  rounded, apex acute to rounded; stems and glandular;  leaves  achenes r i b b e d , h i r s u t e and  8.  A r n i c a v i s c o s a A.  A r n i c a v i s c o s a A.  S i s k i y o u Co., W._L.  Mt.  Greene.  Chrysopsis shastensis Siskiyou  Co.,  Perennial  Mt.  acute;  florets  pappus white to  n=19.  Acad. 13:374.  E r y t h e a 3:48.  1895.  1878. TYPE:  California,  S h a s t a , near the l i m i t of t r e e s , August 4,  Jepson, _s .n_. (not  bases and  stipitate  Gray  Gray. P r o c . Am.  R a i l l a r d e l l a paniculata  lanceolate,  s t i p i t a t e glandular;  tawny, b a r b e l l a t e ; gametic chromosome number  Type:  long  i n f l o r e s c e n c e a s i n g l e head o r much-branched l e a f y corymb; heads  d i s c o i d ; i n v o l u c r a l b r a c t s narrowly to b r o a d l y tubular;  p i l o s e and  1894.  seen) .  Jeps. Man.  F I . PI.  C a l . 1037.  Shasta, Horse Camp, 2400 m,  herb from woody caudex 1-4  W.L.  1927.  TYPE:  Jepson 5 1 i  California, (JEPS).  cm broad, covered w i t h o l d stem  brown s c a l e s ; rhizomes l a c k i n g ; r o o t s  succulent,  mostly v e r t i c a l  232  and unbranched to 40  cm l o n g , 3-6  mm  branched throughout, 25-50 cm t a l l  t h i c k ; stems s o l i t a r y or s e v e r a l , much  at anthesis, sparsely short  g l a n d u l a r and p i l o s e below, densely  stipitate  l o n g s t i p i t a t e g l a n d u l a r and +  pilose  above; i n n o v a t i o n s not produced; c a u l i n e l e a v e s w i t h 6-10(-12) p a i r s , near middle of stem, reduced and  s c a l e - l i k e toward base, reduced and  largest bract-  l i k e above, upper or a l l l e a v e s / b r a c t s o f t e n a l t e r n a t e o r subopposite, gest c a u l i n e l e a v e s 3-5 e l l i p t i c , broadly  cm l o n g , 1.5-3.5 cm broad, n a r r o w l y to  and margins, densely  p i l o s e , e s p e c i a l l y along  long s t i p i t a t e glandular, with yellow-tipped  heads s o l i t a r y t e r m i n a t i n g a l l upper branches; peduncles 0.5-5  cm  wide, n a r r o w l y l a n c e o l a t e , acute; w h i t e to creamy-white, densely achenes grey  mm  hairs; long,  l o n g , 1-3  f l o r e t s 9-32(-54) t u b u l a r , 8-12  acute  veins  p e r i c l i n u m long s t i p i t a t e g l a n d u l a r ; heads e r e c t , d i s c o i d ,  2 cm h i g h ; i n v o l u c r a l b r a c t s , 8-14(-21), + u n i s e r i a t e , 6-12  40-45 um;  broadly  s e s s i l e , margins e n t i r e , bases rounded to acute, t i p s  to obtuse, both s u r f a c e s s p a r s e l y to densely  peduncles and  lar-  mm  1.5mm  long,  l o n g s t i p i t a t e g l a n d u l a r below; p o l l e n diameter  to brown, s p a r s e l y to densely  throughout, s h a l l o w l y r i b b e d , 4-6  mm  l o n g , to 1 mm  tawny, 36-48 b r i s t l e s , b a r b e l l a t e , 5-9  mm  long s t i p i t a t e  glandular  wide, pappus o f f - w h i t e to  long, l a t e r a l setae  .05-.15 mm  long;  s e l f - i n c o m p a t i b l e d i p l o i d ; 2n=38.  TYPE:  C a l i f o r n i a , S i s k i y o u County, Mt.  Shasta,  2400 m,  1877,  J.D.  Hooker &  A. Gray, _s._n. (G) .  Distribution: (1750-2500 m)  T h i s s p e c i e s occurs  on l o o s e pumice s l o p e s at h i g h e l e v a t i o n s  i n C r a t e r Lake N a t i o n a l Park, Klamath County, Oregon and  Western S i s k i y o u and N o r t h e r n T r i n i t y  Counties,  California.  Figure  in  408.  233  F i g u r e 408.  A l l known p o p u l a t i o n s  of A r n i c a v i s c o s a .  @)  235  Illustration:  Figure  409.  A r n i c a v i s c o s a i s the most d i s t i n c t i v e and I t s much-branched h a b i t , s m a l l , e n t i r e , c o i d heads, and  one  of the r a r e s t  Arnicas.  s e s s i l e l e a v e s , narrow, white,  tawny pappus s e t i t a p a r t from a l l o t h e r s p e c i e s .  dis-  I t i s the  o n l y s p e c i e s w i t h white f l o r e t s , a c h a r a c t e r s t r a n g e l y never mentioned i n previous  d e s c r i p t i o n s or monographs.  The  heads a d e f i n i t e y e l l o w i s h c a s t which may  prominent y e l l o w anthers be the reason why  give  the  the white f l o r e t s  have not been observed p r e v i o u s l y . A r n i c a v i s c o s a i s s t r o n g l y v i s c i d throughout w i t h l o n g g l a n d u l a r h a i r s on a l l v e g e t a t i v e p a r t s and on f l o r e t s and s t r o n g odor i s u n l i k e t h a t of any  other A r n i c a .  The  yellow-tipped achenes.  Its  odor i s r e t a i n e d i n  herbarium specimens f o r many y e a r s . Described v i s c o s a was Garfield y e a r s has  from Mt.  Shasta,  California  i n 1887  by Asa  Gray, A r n i c a  long known o n l y from the type l o c a l i t y ( F i g u r e s 410-412) and  Peak i n C r a t e r Lake N a t i o n a l Park, Oregon. i t been noted from a t l e a s t  and  two  localities in  i n the Salmon Mountains and T r i n i t y  Where A r n i c a v i s c o s a has been observed on Mt. t h e r e i s v i r t u a l l y no o t h e r v a s c u l a r p l a n t l i f e .  Shasta  and  Peak  the s o i l below.  able  Both of  a r e q u i t e l a r g e , composed of s e v e r a l hundred p l a n t s .  A study of 25 herbarium specimens and  two  populations  i n the f i e l d  l e s s v a r i a b i l i t y i n t h i s s p e c i e s than i n many of the o t h e r s . tendency f o r upper branches, l e a v e s , and b r a c t s to be ly alternate.  Garfield  Alps.  The v e r y deep r o o t s are  to grow almost v e r t i c a l l y among the l o o s e pumice i n t o these p o p u l a t i o n s  20  f o u r o t h e r p l a c e s i n C r a t e r Lake, f i v e  l o c a l i t i e s i n Western S i s k i y o u County, C a l i f o r n i a , Northern T r i n i t y County, C a l i f o r n i a ,  Only i n the past  from  showed  There i s a g r e a t  subopposite  to complete-  An extreme example of t h i s i s a specimen from Upper E n g l i s h  236  F i g u r e 409.  A r n i c a v i s c o s a S-1411. (Type L o c a l i t y ) .  X  CA,  S i s k i y o u County, Mt.  two-thirds.  Shasta  237  238  F i g u r e s 410-414.  H a b i t a t s and h a b i t of A r n i c a v i s c o s a .  F i g u r e s 410-412.  H a b i t a t of A r n i c a v i s c o s a S-1411. Mt. Shasta  F i g u r e 413.  CA, S i s k i y o u Co.,  (Topotype).  H a b i t a t of A r n i c a v i s c o s a S-1946, OR, Klamath Co., C r a t e r Lake.  F i g u r e 414.  Flowering Mt.  head of A r n i c a v i s c o s a S-1411, CA, S i s k i y o u Co.,  Shasta.  239  240  Lake, S i s k i y o u County, CA, O e t t i n g e r 668  (HSC).  One u n u s u a l l y  pilose  specimen i n which the stem and v e i n s a r e p r e d o m i n a t e l y long p i l o s e than p r e d o m i n a t e l y l o n g s t i p i t a t e g l a n d u l a r as u s u a l was Peak, S i s k i y o u County, CA, Ground  820  ity  of A r n i c a v i s c o s a .  c o l l e c t e d on P r e s t o n  (PUA).  There i s a q u e s t i o n a b l e c o l l e c t i o n by W.L. R a i l l a r d e l l a p a n i c u l a t a by Greene.  rather  It f i t s  As mentioned e a r l i e r  Jepson from Mt. S h a s t a , named  the d e s c r i p t i o n and exact l o c a l there i s v i r t u a l l y no o t h e r  p l a n t l i f e growing near the A r n i c a a t that e l e v a t i o n on Mt. S h a s t a , so  this  c o l l e c t i o n must be t h i s s p e c i e s and i s l i s t e d under the synonymy a l t h o u g h a voucher specimen was  not c i t e d i n the o r i g i n a l d e s c r i p t i o n and has not been  located.  9.  A r n i c a venosa H.M.  Hall.  Univ. of C a l . P u b i s . Bot. 6:174. 1915. '  P e r e n n i a l herb from woody caudex 3-5 cm broad covered w i t h o l d stem bases and dark brown s c a l e s ; rhizomes l a c k i n g ; r o o t s s u c c u l e n t , m o s t l y unbranched, to 30 o r more cm l o n g , 2-5 mm to  t h i c k ; stems s o l i t a r y or few, simple  r a r e l y branched above upper p a i r s of l e a v e s / b r a c t s , 30-50 cm h i g h a t  a n t h e s i s , s p a r s e l y scabrous and p i l o s e below, becoming and s t i p i t a t e g l a n d u l a r above; of  stem, reduced and becoming  more d e n s e l y p i l o s e  c a u l i n e l e a v e s 7-10 p a i r s , l a r g e s t near middle s c a l e - l i k e toward base, and reduced and  l i k e above, upper l e a v e s and b r a c t s o f t e n s u b o p p o s i t e o r a l t e r n a t e , cauline leaves to  bract-  largest  (3-)4-6(-7) cm l o n g , (1.5-)2-3(-4) cm broad, n a r r o w l y  elliptic  b r o a d l y so, b r o a d l y s e s s i l e , margins e v e n l y c o a r s e l y s e r r a t e o r r a r e l y  doubly s e r r a t e , bases rounded, t i p s acute to obtuse, rugose, v e i n s prominent, e s p e c i a l l y on lower s u r f a c e , lower s u r f a c e s p a r s e l y to d e n s e l y p i l o s e  and  s p a r s e l y s t i p i t a t e g l a n d u l a r ; i n f l o r e s c e n c e a s i n g l e head t e r m i n a t i n g main  241  stem and  l a t e r a l branches, i f any;  peduncles 1.5-5  cm  l o n g , peduncles  p e r i c l i n u m d e n s e l y p i l o s e ; heads nodding, e s p e c i a l l y i n bud, cm h i g h ; broadly  i n v o l u c r a l b r a c t s 8-14,  mm  discoid,  l o n g , 4-6  mm  2-2.5 broad,  l a n c e o l a t e , acute to obtuse; f l o r e t s 32-56(-70), t u b u l a r , 8-10  l o n g , dark y e l l o w , pm;  + u n i s e r i a t e , 10-15  and  mm  densely v i l l o u s , e s p e c i a l l y below; p o l l e n diameter 35-40  achenes grey, s t r o n g l y angled and  r i b b e d , 6-8  mm  l o n g , 1.5  mm  broad,  d e n s e l y h i r s u t e w i t h b i f u r c a t e h a i r s ; pappus w h i t e , of 45-55 b r i s t l e s , mm  l o n g , b a r b e l l a t e , the l a t e r a l setae 0.1-0.2 mm  long;  6-10  self-incompatible;  2n=38.  TYPE: _E.F.  C a l i f o r n i a , Shasta County, S a l t Creek, 430  Bab cock 4013  Distribution:  m,  H.M.  Hall &  (UC).  A r n i c a venosa i s found i n f o o t h i l l s  Sacramento R i v e r V a l l e y i n mixed c o n i f e r and County, C a l i f o r n i a .  Illustration:  June 1903,  Figure  Figure  (430-530 m)  Quercus spp.  of  the  f o r e s t of Shasta  415.  416.  T h i s d i s t i n c t i v e s p e c i e s , c l o s e l y r e l a t e d to A r n i c a v i s c o s a , has l e a v e s , nodding buds, y e l l o w t i n g i t apart species.  f l o r e t s , and  from t h a t s p e c i e s .  For many years i t was  T h i s too, i s one known from o n l y two  County, C a l i f o r n i a , I r o n Mountain and ity.  The  exact  a p a u c i t y of g l a n d u l a r  Shasta. and  o f the r a r e r A r n i c a localities  l o c a t i o n of " S a l t Creek" i s not known.  I r o n Mountain i s now  the A r n i c a may  hairs, set-  i n Shasta  S a l t Creek, the l a t t e r the type  S a l t Creeks i n Shasta County, i n c l u d i n g one  which i s now  There are  have been e x t e r m i n a t e d from t h a t l o c a l i t y . the a r e a .  local-  several  p a r t i a l l y under Lake  the s i t e of an a c t i v e copper mining  owners w i l l not a l l o w b o t a n i s t s to search  toothed  Having not  The  operation mine  been c o l l e c t e d  242  F i g u r e 415.  A l l known p o p u l a t i o n s  of A r n i c a venosa.  0 50 100  200km  244  F i g u r e 416.  A r n i c a venosa S-1793.  CA, Shasta County.  X  two-thirds.  245  246  F i g u r e s 417-421.  H a b i t a t and h a b i t of A r n i c a venosa S-1793,  CA,  S h a s t a Co., Gibson. F i g u r e s 417-418.  H a b i t a t , r o a d s i d e bank i n open mixed f o r e s t of Quercus  spp. and  conifers.  F i g u r e 419.  H a b i t of numerous branches from base.  F i g u r e 420.  S t r o n g l y nodding bud.  F i g u r e 421.  Head w i t h f i r s t rugose l e a v e s .  f l o r e t s open and nodding buds,  and  248  for  more than 50 y e a r s , i t was  r e c e n t y e a r s has personnel found, It  to be p o s s i b l y e x t i n c t .  t h e r e been renewed i n t e r e s t by  i n the a r e a to f i n d  a l l i n Shasta i s found  thought  i t , and  Only i n  the N a t i o n a l F o r e s t S e r v i c e  a t l e a s t 5 o t h e r l o c a l i t i e s have been  County.  i n d i s c r e t e p o p u l a t i o n s of a few  individuals  to a few  p l a n t s , i n open f o r e s t s of P i n u s ponderosa, Pinus l a m b e r t i a n a , Pinus u a t a , P i n u s s a b i n i a n a , P s e u d o t s u g a m e n z i e s i i , Quercus k e l l o g g i i , chrysolepis, and  Calocedrus  decurrens,  A r c t o s t a p h y l o s spp.  a l o n g open r o a d s i d e s i n t h e s e f o r e s t s .  d i s t u r b e d r o a d s i d e banks.  and  s t r a g g l e r s may  cultivation  P l a n t s b e g i n to f l o w e r i n mid-  c o n t i n u e to f l o w e r throughout  seedlings continued  atten-  Quercus  Ceanothus  spp.,  P l a n t s s p a r i n g l y c o l o n i z e the to l a t e May  bloom f o r a few weeks b e f o r e t h e h i l l s become v e r y hot and few  dozen  and  dry, although  a  the e a r l y summer months.  In  to f l o w e r f o r many months when grown under  wetter c o n d i t i o n s . Specimens a r e v e r y r a r e i n h e r b a r i a .  Of about 6 specimens seen from 3  l o c a l i t i e s p l u s 3 o t h e r p o p u l a t i o n s s t u d i e d i n the f i e l d and  summer of 1978,  d u r i n g the s p r i n g  t h e r e seems to be v e r y l i t t l e v a r i a b i l i t y  i n the p l a n t s .  249  PHYLOGENY  Maguire (1943) s u g g e s t s  t h a t t h e genus A r n i c a p r o b a b l y  or s u b a r c t i c Western N o r t h America and south. He  He  recognized  considered  spread  in arctic  from t h e r e to the e a s t  and  f i v e subgenera w i t h A r c t i c a as the a n c e s t r a l subgenus.  the l a r g e , h e m i s p h e r i c  or t u r b i n a t e heads and  p e r i c l i n u m as b e i n g p r i m i t i v e c h a r a c t e r s . are considered p r i m i t i v e , F i g u r e 422  arose  then  Maguire's p h y l o g e n e t i c  I f these m o r p h o l o g i c a l  the p o s s i b l e p h y l o g e n e t i c  can be deduced f o r t h e o t h e r  densely  subgenera.  pubescent  characters  scheme shown i n  T h i s i s a m o d i f i c a t i o n of  scheme f o r h i s f i v e subgenera, p l u s my  newly proposed  subgenus C a l a r n i c a . Within  subgenus Austromontana, A r n i c a c o r d i f o l i a i s the p r o b a b l e  between the two  subgenera, A r c t i c a and  u s u a l l y s o l i t a r y heads, and the o t h e r  densely p i l o s e periclinum.  s p e c i e s i n Austromontana and  as shown i n F i g u r e  Austromontana.  link  T h i s s p e c i e s has From A r n i c a  cordifolia,  C a l a r n i c a c o u l d l o g i c a l l y have  evolved  423.  Arnica c o r d i f o l i a occurs p o p u l a t i o n s , from t r i p l o i d  today l a r g e l y as a s e r i e s of a p o m i c t i c p o l y p l o i d  to h e x a p l o i d , w i t h  e v e r , d i p l o i d p o p u l a t i o n s do o c c u r r a r e l y .  t e t r a p l o i d predominating.  Whether o r n o t any  d i p l o i d s are s e x u a l has not been demonstrated.  of  How-  these  Based on p o l l e n q u a l i t y  the u s u a l l y s t r o n g c o r r e l a t i o n of d i p l o i d y w i t h s e x u a l i t y i n the genus, i s a strong p o s s i b i l i t y Barker  t h a t some of these  (1966) h y p o t h e s i z e d  d i p l o i d populations  Sexual  been more common and  past.  widespread  i n the  D i p l o i d A r n i c a c o r d i f o l i a probably habitats.  are  there  recent  d i p l o i d s a r e assumed to have  gave r i s e to A r n i c a l a t i f o l i a  Arnica l a t i f o l i a  and  sexual.  t h a t a p o m i x i s i n the genus i s a r e l a t i v e l y  development ( i n t e r - o r p o s t - g l a c i a l ) .  g e n e r a l l y wetter  large,  in  u s u a l l y has narrower heads, w i t h  250 fewer f l o r e t s , and a l e s s - p i l o s e p e r i c l i n u m than A r n i c a c o r d i f o l i a . characters  are considered  as b e i n g  derived.  Arnica l a t i f o l i a  These  i s the most  polymorphic s p e c i e s i n t h e subgenus, a l t h o u g h i t e x i s t s almost e n t i r e l y as sexual d i p l o i d populations,  w i t h many e c o t y p e s .  The c l o s e r e l a t i o n s h i p o f  these two s p e c i e s  can be seen i n numerous p o p u l a t i o n s  the m o r p h o l o g i c a l  characters  Arnica l a t i f o l i a  species.  has i n t u r n p r o b a b l y  A r n i c a cernua, a l s o a s e x u a l combines many m o r p h o l o g i c a l distinct.  o f both  diploid,  given  i s a rare serpentine  ly  by many a u t h o r s .  r o c k i e r h a b i t a t s than A r n i c a l a t i f o l i a .  these c h a r a c t e r s  as b e i n g  fewer f l o r e t s  advanced.  possibly  populations  Arnica g r a c i l i s  Arnica g r a c i l i s  It  a v a r i e t y of  However, i n my o p i n i o n , most  autonomous apomict, found m o s t l y a t h i g h e r  has much s m a l l e r heads, w i t h  endemic.  but i s e c o l o g i c a l l y  I t i s considered  be e a s i l y d i s t i n g u i s h e d from A r n i c a l a t i f o l i a . a triploid,  soil  species.  d e r i v e d from A r n i c a l a t i f o l i a ,  through h y b r i d i z a t i o n , i s A r n i c a g r a c i l i s .  can  r i s e to two o t h e r  characters of Arnica l a t i f o l i a  A second s p e c i e s p r o b a b l y  Arnica l a t i f o l i a  which combine many of  i s large-  elevations or i n i s more branched and  than A r n i c a l a t i f o l i a .  I interpret  A r n i c a g r a c i l i s has the g l a n d u l a r  pubes-  cence found i n A r n i c a d i v e r s i f o l i a and A r n i c a m o l l i s i n subgenus C h a m i s s o n i s . Both o f these l a t t e r The  s p e c i e s have been shown to have t e t r a p l o i d  chromosome number and morphology o f A r n i c a g r a c i l i s  h y b r i d o r i g i n between d i p l o i d A r n i c a l a t i f o l i a  s u g g e s t s i t may be o f  and t e t r a p l o i d A r n i c a  f o l i a o r A r n i c a m o l l i s , a l t h o u g h attempts t o make t h i s c r o s s have been u n s u c c e s s f u l .  Arnica l a t i f o l i a  elements.  o f t e n grows w i t h  diversi-  artificially  e i t h e r o r both  A r n i c a d i v e r s i f o l i a and A r n i c a m o l l i s , but t h e r e a r e p r e s e n t l y b a r r i e r s to h y b r i d i z a t i o n between t h e s e x u a l of t h e l a t t e r  d i p l o i d and t h e a p o m i c t i c  two s p e c i e s u s u a l l y have v e r y  low p o l l e n  tetraploids.  fertility.  Both  251  A r n i c a n e v a d e n s i s c l o s e l y resembles A r n i c a c o r d i f o l i a i n morphology i n being  a s e r i e s of t e t r a p l o i d autonomous a p o m i c t i c  u s u a l l y e c o l o g i c a l l y d i s t i n c t , being exposed h a b i t a t s . ficult  found a t h i g h e r  There are populations  to a s s i g n to one  s p e c i e s o r the o t h e r .  a r i s e n as s e l e c t i o n s of a p o m i c t i c i z a t i o n with other  of these two  species.  races  races.  The  s h o r t e r pubescence, and  attenuated  suggest t h a t some o t h e r  s p e c i e s may  former i s  e l e v a t i o n s and  i n more  s p e c i e s which are  A r n i c a n e v a d e n s i s may  dif-  have  of A r n i c a c o r d i f o l i a , w i t h o u t  However, the l o n g e r pappus s e t a e ,  and  hybrid-  tawny pappus,  o r rounded l e a f bases of A r n i c a  nevadensis  have been i n v o l v e d i n i t s e a r l y e v o l u t i o n .  Based on morphology, A r n i c a n e v a d e n s i s c o u l d have o r i g i n a t e d as the r e s u l t h y b r i d i z a t i o n between A r n i c a c o r d i f o l i a and  A r n i c a c h a m i s s o n i s i n the  of  sub-  genus Chamissonis. Arnica discoidea d i f f e r s l i t t l e l a c k o f ray f l o r e t s .  The  from A r n i c a c o r d i f o l i a except i n i t s  u s u a l l y , more-branched h a b i t , more heads, and  c o i d c o n d i t i o n of A r n i c a d i s c o i d e a are  i n t e r p r e t e d as d e r i v e d  A r n i c a d i s c o i d e a i s a p l a n t of c h a p a r r a l and of A r n i c a c o r d i f o l i a . be  a serpentine  from any  Another d i s c o i d s p e c i e s , A r n i c a s p a t h u l a t a ,  as b o t h d i p l o i d and  subgenus C a l a r n i c a w i t h  venosa i s considered of the o t h e r  characters.  d r i e r h a b i t a t s than  endemic d e r i v a t i v e of A r n i c a d i s c o i d e a .  c o i d species occur The  other  as d i s t i n c t subgenera.  two  tetraploid  Both of these  dis-  populations.  s p e c i e s , A r n i c a v i s c o s a and  Arnica  from subgenus Austromontana as the l a t t e r i s I consider  subgenus C a l a r n i c a to be  i t s numerous s m a l l heads, s e s s i l e l e a v e s , l a c k of rhizomes, and  tion  to d r i e r h a b i t a t s .  i s unclear.  that  seems to  with  The  dis-  r e l a t i o n s h i p of t h i s subgenus.to o t h e r  I t i s p o s s i b l y d e r i v e d from A r n i c a s p a t h u l a t a .  The  advanced adapta-  subgenera  pappus,  pubescence, and woody caudex o f A r n i c a venosa are c l o s e r to t h a t of  Arnica  252  spathulata  than any  other  species.  A r n i c a v i s c o s a i s probably  e l e v a t i o n d e r i v a t i v e of A r n i c a v e n o s a. consider Arnica v i s c o s a , with most advanced member of the  i t s very  genus.  They are b o t h s e x u a l narrow heads and  a  high-  diploids.  white f l o r e t s ,  I the  253 F i g u r e 422.  Possible Phylogenetic  R e l a t i o n s h i p s of A r n i c a  ( m o d i f i e d a f t e r Maguire, 1943).  Subgenera  254  CALARNICA  AUSTROMONTANA CHAMISSONIS  ARCTICA  ANCESTRAL SPECIES  PHYLOGENETIC RELATIONSHIPS IN ARNICA SUBGENERA  255  F i g u r e 423.  Possible Phylogenetic  R e l a t i o n s h i p s of A r n i c a  i n Subgenera Austromontana and C a l a r n i c a .  Species  256  VISCOSA  VENOSA  CERNUA  CALARNICA  GRACILIS  LATIFOLIA  SPATHULATA  DISCOIDEA  NEVADENSIS CORDIFOLIA  AUSTROMONTANA  257 SUMMARY  A c o n s e r v a t i v e taxonomic t r e a t m e n t seven s p e c i e s and sented.  a new  T h i s treatment  artificial evaluation.  of A r n i c a subgenus Austromontana w i t h  subgenus, C a l a r n i c a , w i t h i s based on f i e l d  hybridizations, insect  two  s p e c i e s has been p r e -  studies, cytology, breeding  i n t e r r e l a t i o n s h i p s , and  systems,  a morphological  In a monograph of t h e genus A r n i c a by Maguire (1943) t h e s e  subgenera were r e c o g n i z e d as one, i n f r a s p e c i f i c taxa.  Austromontana, w i t h 13 s p e c i e s and  retwo  three  I r e c o g n i z e the f o l l o w i n g s p e c i e s i n subgenus A u s t r o -  montana: Sexual, incompatible Arnica l a t i f o l i a Arnica  diploids (one  tetraploid population  reported)  cernua  A r n i c a s p a t h u l a t a (one  t e t r a p l o i d population  reported)  Pseudogamous a p o m i c t i c p o l y p l o i d s Arnica c o r d i f o l i a  (some autonomous apomicts and  Pseudogamous a p o m i c t i c p o l y p l o i d s and  rarely,  sexual, incompatible  diploids) diploids  Arnica discoidea Autonomous a p o m i c t i c p o l y p l o i d s Arnica  gracilis  Arnica  nevadensis  I r e c o g n i z e the f o l l o w i n g s e x u a l , i n c o m p a t i b l e Calarnica: A r n i c a venosa Arnica viscosa  diploids  i n subgenus  258  No two  i n f r a s p e c i f i c taxa are recognized.  o r more r e l a t i v e l y d i s t i n c t  Although  several species  elements, t h e r e i s , i n my  o p i n i o n , too much  o v e r l a p i n c h a r a c t e r s to w a r r a n t g i v i n g them taxonomic r e c o g n i t i o n . a l s o does not o f these  seem to be any  c o n s i s t e n t geographical or e c o l o g i c a l  There isolation  populations.  There i s a s t r o n g c o r r e l a t i o n between h i g h p o l l e n s t a i n a b i l i t y , and  contain  sexuality.  diploidy,  Autonomous a p o m i c t i c p o l y p l o i d s have v e r y low p o l l e n s t a i n -  a b i l i t y , o r p o l l e n i s n o t produced a t a l l . have h i g h e r p o l l e n s t a i n a b i l i t y lower than the s e x u a l  than  Pseudogamous a p o m i c t i c p o l y p l o i d s  the autonomous a p o m i c t i c s , but u s u a l l y  diploids.  Seed s e t and v i a b i l i t y  i s q u i t e low  i n most of t h e s p e c i e s , and  achenes which are produced a r e o f t e n f e d upon by  insect predators.  those On  o t h e r hand, v e g e t a t i v e r e p r o d u c t i o n by rhizomes has been shown to be  the  an  impor-  t a n t f a c t o r i n s h o r t d i s t a n c e d i s p e r s a l o f s e v e r a l of the s p e c i e s i n subgenus Austromontana. There a r e no Every  completely  s p e c i e s , as I r e c o g n i z e  c l e a r - c u t s p e c i e s i n subgenus Austromontana. them, i n t e r g r a d e s i n t o one  o r more o t h e r  species.  In the s e x u a l d i p l o i d s t h e r e a r e a number of l o c a l e c o t y p e s  which a r e  p l a s t i c , responding  I n d i v i d u a l s have  to the l o c a l e n v i r o n m e n t a l  been shown to e x h i b i t m o r p h o l o g i c a l ing environmental  d i f f e r e n c e s from year  conditions i n c u l t i v a t i o n .  a r e r e c o g n i z e d as a s e r i e s o f o u t b r e e d i n g ability, The species.  s i m i l a r morphology, and two  sexual, d i p l o i d  These two  conditions.  Species  to y e a r under  i n these  sexual  vary-  diploids  populations, with high p o l l e n s t a i n -  similar ecological niches.  s p e c i e s i n subgenus C a l a r n i c a are v e r y c l e a r - c u t  r a r e s p e c i e s show v e r y l i t t l e  I n the a p o m i c t i c  very  interpopulational variation.  s p e c i e s , a.number o f a p o m i c t i c r a c e s have r e c e i v e d  taxonomic r e c o g n i t i o n i n t h e p a s t .  In r e a l i t y every p o p u l a t i o n may  be  259  considered a species.  These a r e s e l f - b r e e d i n g e n t i t i e s w i t h b a r r i e r s t o  interbreeding with other morphologically they may have r e c e n t l y e v o l v e d  s i m i l a r p o p u l a t i o n s , even though  from a common a n c e s t o r .  these p o p u l a t i o n s as a s p e c i e s would be c o m p l e t e l y  To t r e a t each o f  impractical.  Other  t e r i a such as p o l l e n s t a i n a b i l i t y o r chromosome numbers a r e o f l i t t l e elucidating and  species.  ecologically  crihelp i n  I t seems b e s t to r e c o g n i z e groups o f m o r p h o l o g i c a l l y  s i m i l a r a p o m i c t i c r a c e s as s p e c i e s .  In one s p e c i e s , A r n i c a d i s c o i d e a , t h e r e a r e both apomictic, t e t r a p l o i d populations.  s e x u a l , d i p l o i d and  The t e t r a p l o i d s a r e p r o b a b l y  r i v a t i v e s , which have n o t become m o r p h o l o g i c a l l y d i s t i n c t . c h a r a c t e r s which can be used to s e p a r a t e counterparts, other  than  I can see no  the t e t r a p l o i d s from t h e i r  t h e i r d i f f e r e n t breeding  as a s i n g l e s p e c i e s w i t h both  r e c e n t de-  s e x u a l and a p o m i c t i c  systems. elements.  diploid  They a r e t r e a t e d  260  LITERATURE CITED  Abrahamson, W. & K.D. McCrea. 1977. 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A r n i c a s p a t h u l a t a Greene subsp. genuina Maguire lib.  A r n i c a s p a t h u l a t a Greene subsp. eastwoodiae (Rydb.) Maguire  12.  A r n i c a venosa H.M.  13.  A r n i c a v i s c o s a A. Gray  Hall  -  265  APPENDIX I I  Taxa r e c o g n i z e d by E d i g e r & B a r k l e y  (1978)  Subgenus Austromontana la.  A r n i c a l a t i f o l i a Bongard v a r . l a t i f o l i a  lb.  A r n i c a l a t i f o l i a Bongard v a r . g r a c i l i s  2.  Arnica  cernua H o w e l l  3.  Arnica  t o m e n t e l l a Greene  4.  A r n i c a nevadensis  (Rydb.)  Cronquist  A. Gray  5a. A r n i c a  c o r d i f o l i a Hooker v a r . c o r d i f o l i a  5b. A r n i c a  c o r d i f o l i a Hooker v a r . pumila  6a. A r n i c a  d i s c o i d e a Bentham v a r . a l a t a  6b. A r n i c a  d i s c o i d e a Bentham v a r . e r a d i a t a  (Rydb.) Maguire (Rydb.)  Cronquist  (A. Gray)  Cronquist  6c. A r n i c a d i s c o i d e a Bentham v a r . d i s c o i d e a 7a. A r n i c a  s p a t h u l a t a Greene v a r . s p a t h u l a t a  7b. A r n i c a  s p a t h u l a t a Greene v a r . eastwoodiae (Rydb.) E d i g e r &  8.  A r n i c a v e n o s a H.M.  9.  A r n i c a v i s c o s a A. Gray  Hall  

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