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Studies in the genus saxifraga in the Pacific Northwest : Part I: taxonomy of saxifraga occidentalis… Krause, David Larkin 1972

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c  STUDIES IN THE GENUS SAXIFRAGA IN THE PART I :  PACIFIC NORTHWEST  TAXONOMY OF SAXIFRAGA OCCIDENTALS AND S. MARSHALLII.  PART I I : NOTES ON SAXIFRAGA OCCIDENTALS AND CLOSELY RELATED SPECIES.  by.  DAVID L. KRAUSE B.Sc,  U n i v e r s i t y of B r i t i s h  Columbia, 1964  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE  i n t h e Department of Botany  We a c c e p t t h i s  t h e s i s as conforming t o the  required standard  THE UNIVERSITY OF BRITISH COLUMBIA December, 1972  <  In p r e s e n t i n g  t h i s t h e s i s in p a r t i a l  f u l f i l m e n t o f the  requirements f o r  an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree the L i b r a r y s h a l l make i t f r e e l y  a v a i l a b l e f o r r e f e r e n c e and  I f u r t h e r agree t h a t p e r m i s s i o n f o r e x t e n s i v e f o r s c h o l a r l y purposes may by h i s r e p r e s e n t a t i v e s .  be g r a n t e d by  thesis for financial  written  permission.  gain  Department o f The U n i v e r s i t y o f B r i t i s h Columbia Vancouver 8, Canada  Date  £7  shall  not  thesis  Department o r  I t i s u n d e r s t o o d that c o p y i n g o r  of t h i s  study.  copying of t h i s  the Head o f my  that  publication  be a l l o w e d w i t h o u t  my  ii.  ABSTRACT  PART I . The  Taxonomy o f S a x i f r a g a o c c i d e n t a l i s  and S_. marshal I i i .  taxonomy o f S a x i f r a g a m a r s h a l l i i and S_. o c c i d e n t a l i s i s  reviewed.  On m o r p h o l o g i c a l , c y t o l o g i c a l and d i s t r i b u t i o n a l e v i d e n c e ,  the new combination S_. m a r s h a l l i i s s p . i d a h o e n s i s i s proposed f o r t h e taxon o r i g i n a l l y r e f e r r e d  t o as S_. i d a h o e n s i s .  Keys t o the t a x a ,  synonomy, d i s t r i b u t i o n a l r e c o r d s , maps, and chromosome numbers are  presented.  PART I I . Notes on S a x i f r a g a o c c i d e n t a l i s  and c l o s e l y r e l a t e d  species.  A s p e c i e s key, d i s t r i b u t i o n maps and some chromosome numbers a r e presented f o r Saxifraga  occidentalis  S. Wats., S_. rhomboidea Greene,  S_. r u f i d u l a (Small) James Macoun, S^. r e f l e x a Hook., and the comb i n e d s p e c i e s S_. n i v a l i s L. - S_. t e n u i s possible  (Wahl.) H. Sm.  t o s e p a r a t e these f i v e t a x a u t i l i z i n g  characters.  certain  Itis qualitative  S a x i f r a g a n i v a l i s and S_. t e n u i s a r e two c l o s e l y  t a x a whose r e l a t i o n s h i p s  a r e unknown.  related  iii. TABLE OF CONTENTS  PART I .  Taxonomy of S a x i f r a g a o c c i d e n t a l i s and S_. m a r s h a l l i i .  Abstract  . i i .  Introduction  1.  M a t e r i a l s and Methods  3.  Taxonomy  6.  Discussion  20.  Literature Cited  24.  Table 1  26.  F i g u r e s 1-5  27.  Figure 6  .28.  Figure 7  29.  PART I I .  Notes on S a x i f r a g a o c c i d e n t a l i s and c l o s e l y r e l a t e d s p e c i e s .  Abstract  i i .  Introduction  30.  Taxonomy  32.  Literature Cited  41.  Table 1  43.  Figure 1  44.  Figure 2  45.  Figure 3  .46.  1. PART I INTRODUCTION  In the P a c i f i c Northwest the r e l a t i o n s h i p s i n the genus S a x i f r a g a are o f t e n d o u b t f u l and  the  names d i f f i c u l t  to a p p l y .  c e r t a i n l y true for Section Boraphila subsection e s t a b l i s h e d by E n g l e r and ment of S a x i f r a g a c e a e .  T h i s statement i s  Nivali-virginienses  Irmscher (1916) i n t h e i r comprehensive  The marked disagreement over the  i n t e r - r e l a t i o n s h i p s of t h i s s u b s e c t i o n are due  partly  treat-  intra-  to the  and  morphological  s i m i l a r i t i e s between e n t i t i e s and p a r t l y to the many s p e c i f i c and  sub-  s p e c i f i c d e s c r i p t i o n s which have r e s u l t e d i n c o n s i d e r a b l e synonomy. One  o f the  problems w i t h i n the group i s the r e l a t i o n s h i p of  the  taxa commonly r e f e r r e d to as S_. o c c i d e n t a l i s S. Wats., Sj_ m a r s h a l l i i Greene, and  S_. i d a h o e n s i s  In 1888  Piper.  Greene p u b l i s h e d the name S^. m a r s h a l l i i f o r a s a x i f r a g e  found i n the Hoopa V a l l e y , Humboldt County, C a l i f o r n i a . Sereno Watson d e s c r i b e d a new  s p e c i e s , jS.  Rocky Mountains of B r i t i s h America  Also i n  o c c i d e n t a l i s , "...  1888,  from the  (Drummond) to B r i t i s h Columbia  and  Vancouver I s l a n d ( L y a l l , Macoun), Oregon ( C u s i c k , Henderson, H o w e l l ) , the n o r t h e r n  S i e r r a Nevada ( C h i c o , Mrs.  specimen was  designated  more than one is  and  J . B i d w e l l , Gray)."  i t i s g e n e r a l l y accepted  s p e c i e s w i t h i n the a r e a c i t e d .  The  No  and  type  that there i s  name S_. o c c i d e n t a l i s  f r e q u e n t l y used f o r the most common of the B r i t i s h Columbia s a x i -  f r a g e s from the s u b s e c t i o n N i v a l i - v i r g i n i e n s e s . d e s c r i b e d a new  In 1900  C. V.  s p e c i e s from w e s t e r n Idaho, S^. i d a h o e n s i s .  Piper  He b e l i e v e d  2.  it its  t o be r e l a t e d  t o S_. o c c i d e n t a l i s  "... n e a r l y g l a b r o u s  and S_. r e f l e x a , b u t d i f f e r i n g by  l e a v e s , much branched i n f l o r e s c e n c e and s m a l l  flowers  ..." E n g l e r and Irmscher (1916) reduced  variety  of  marshallii.  species closely  related  S_. i d a h o e n s i s t o a  Abrams (1944) m a i n t a i n e d  S_. i d a h o e n s i s as a  t o S_. m a r s h a l l i i , b u t s e p a r a t e d from the l a t t e r  on the b a s i s of s m a l l e r f l o w e r s and absence o f y e l l o w spots on t h e p e t a l s . In t h e  l a t e s t treatment  S_. i d a h o e n s i s a v a r i e t y  o f the group, H i t c h c o c k e t a l . (1961) made o f S_. o c c i d e n t a l i s  o t h e r races o f S_. o c c i d e n t a l i s ,  because o f i t s t r a n s i t i o n t o  and because i t i s g e o g r a p h i c a l l y i s o l a t e d  from S_. m a r s h a l l i i . Small  (1905) and Johnson (1923) p u b l i s h e d many  to s u b s e c t i o n N i v a l i - v i r g i n i e n s e s , reduced  t o synonomy, mainly  most  o f which  new s p e c i e s b e l o n g i n g  have s i n c e been  o f S_. o c c i d e n t a l i s .  The p r e s e n t paper i s an attempt t o b r i n g c y t o l o g i c a l , and  d i s t r i b u t i o n a l evidence  S. o c c i d e n t a l i s ,  morphological,  t o g e t h e r t o determine the r e l a t i o n s h i p s  S. m a r s h a l l i i , and S. i d a h o e n s i s .  of  3. MATERIAL AND METHODS  F i e l d work has been done mainly across and n o r t h w e s t e r n U n i t e d whenever p o s s i b l e . of B r i t i s h  The  States. live  southern B r i t i s h  Both l i v e m a t e r i a l and buds were c o l l e c t e d  m a t e r i a l was  grown i n pots a t The U n i v e r s i t y  Columbia.  L i v e c o l l e c t i o n s and h e r b a r i u m specimens were s t u d i e d t i v e l y and q u a l i t a t i v e l y descriptions. processed standard  Columbia  The data  quantita-  f o r the c h a r a c t e r i s t i c s used i n the taxa from the q u a n t i t a t i v e c h a r a c t e r i s t i c s were  f o r mean, m i d p o i n t , range, v a r i a n c e , s t a n d a r d e r r o r , and c o n f i d e n c e  l i m i t s a t the 5%  level.  deviation, Scheffe's  f o r d i f f e r e n c e s between groups u s i n g unequal sample s i z e s was between p o p u l a t i o n s  and between  attempted  taxa.  The q u a n t i t a t i v e measurements were inflorescence.  taken  o f the  l e a f , scape, and  Of these measurements o n l y the means and s t a n d a r d  t i o n s were found t o be u s e f u l .  The means and s t a n d a r d  however, each c h a r a c t e r has such a wide  diagnostic characters.  devia-  d e v i a t i o n s were  used to d e l i m i t the v a r i a b i l i t y of a s i n g l e m o r p h o l o g i c a l  l a p s so much the range i n o t h e r  test  character;  range i n each taxon and o v e r -  taxa t h a t they a r e not u s e f u l  as  G r a p h i c r e p r e s e n t a t i o n u s i n g computerized s c a t t e r  diagrams confirmed t h i s h i g h degree o f o v e r l a p  f o r the q u a n t i t a t i v e  characters. The e v a l u a t i o n of the q u a l i t a t i v e u s e f u l f o r the s e p a r a t i o n o f the v a r i o u s t i v e characters  c h a r a c t e r s was found to be most taxa.  A combination of q u a l i t a -  i s u s u a l l y needed as no s i n g l e c h a r a c t e r c o u l d be used to  4.  separate a l l taxa.  F o r example  the presence  o f y e l l o w spots on t h e  p e t a l s i s d i a g n o s t i c f o r S_. m a r s h a l l i i , b u t t h e absence o f spots on herbarium specimens i s n o t d i a g n o s t i c as the spots commonly f a d e upon drying. The buds c o l l e c t e d i n the f i e l d and taken from m a t e r i a l grown i n pots were f i x e d i n 95% e t h a n o l and g l a c i a l a c e t i c a c i d  (3:1).  Buds  were s t a i n e d u s i n g Snow's carmine t e c h n i q u e (1963) and anthers d i s s e c t e d out and squashed i n h a e m a t o x y l i n (Whittman 1965).  Voucher  specimens f o r chromosome counts a r e kept i n t h e h e r b a r i u m o f The U n i v e r s i t y of B r i t i s h  Columbia.  C o l l e c t i o n d a t a and chromosome counts a r e g i v e n . i n T a b l e 1. Herbarium m a t e r i a l was borrowed from the h e r b a r i a l i s t e d  below.  ALTA  Herbarium, Department  of Botany, U n i v e r s i t y of A l b e r t a , Edmonton.  UAC  Herbarium, Department  of Biology, U n i v e r s i t y of Calgary, A l b e r t a .  CAN  N a t i o n a l Museum o f Canada,  COLO  Herbarium, U n i v e r s i t y o f C o l o r a d o , B o u l d e r , C o l o r a d o .  DAO  P l a n t Research I n s t i t u t e , Department  GH  Gray Herbarium o f Harvard U n i v e r s i t y , Cambridge, Mass.  ID  Department  JEPS  Jepson Herbarium, Department  Ottawa.  of A g r i c u l t u r e ,  Ottawa.  o f Botany, U n i v e r s i t y o f Idaho, Moscow, Idaho. o f Botany, U n i v e r s i t y of C a l i f o r n i a ,  Berkeley. MONTU  U n i v e r s i t y o f Montana, M i s s o u l a ,  Montana.  ND  U n i v e r s i t y of N o t r e Dame, N o t r e Dame, I n d i a n a .  NY  The New York B o t a n i c a l Garden, New York C i t y , New York.  Herbarium o f the U n i v e r s i t y o f Oregon, Eugene, Oregon. Herbarium, Department  o f Botany, Oregon S t a t e  University,  Corvallis. Academy o f N a t u r a l  Sciences, P h i l a d e l p h i a ,  Rocky Mountain Herbarium, U n i v e r s i t y  Pennsylvania.  o f Wyoming, Laramie,  Wyoming. Herbarium o f the U n i v e r s i t y o f C a l i f o r n i a , Department University  of C a l i f o r n i a ,  U.S. N a t i o n a l  Museum  o f Botany,  Berkeley.  (Department o f B o t a n y ) , Smithsonian I n s t i -  t u t i o n , Washington, D.C. Department  of Plant Pathology, State  College  o f Washington,  Pullman, Washington. Herbarium, Department Seattle.  o f Botany, U n i v e r s i t y o f Washington,  6.  TAXONOMY  KEY  A.  TO THE  SPECIES  I n f l o r e s c e n c e open, branches and peduncles u s u a l l y d i v a r i c a t e , p e d i c e l s 2 to 4 times as l o n g as f l o w e r s ; flowers seldom grouped at ends o f b r a n c h e s ; f i l a m e n t s c l a v a t e , o f t e n showy a f t e r p e t a l s f a l l ; w i t h 2 y e l l o w spots  a t base ( o f t e n  inconspicuous  deciduous; s e p a l s r e f l e x e d ; ovary l e s s than 1/5 B.  Ovary almost completely oblong; p e t i o l e s (0.5)  on d r i e d m a t e r i a l ) , inferior.  s u p e r i o r ; l e a v e s oblong-ovate to 1 to 3 times as l o n g as the b l a d e  Western Oregon and N.W.  Ovary from almost completely  s u p e r i o r to 1/5  ovate to rhombic; p e t i o l e s 0.5 blade  length.  to 1.5  Chelan and K i t t i t a s  C e n t r a l Idaho and  a d j a c e n t N.E.  inferior;  (2) times as l o n g  Cos.  length.  marshallii leaves as  of Washington; North  Oregon and w e s t e r n Montana 2. S_. m a r s h a l l i i ssp.  AA.  elliptic-  California 1. S_. m a r s h a l l i i s s p .  BB.  petals  idahoensis  I n f l o r e s c e n c e open to congested, branches and peduncles s p r e a d i n g ascending, flowers  p e d i c e l s e q u a l t o , to twice as l o n g as, the  tending  to be  p e t a l s without to p a r t i a l l y inferior.  flowers;  grouped at ends of branches; f i l a m e n t s  l i n e a r or sometimes s l i g h t l y  clavate,  inconspicuous  British  subulate,  i f petals  b a s a l yellow spots, often p e r s i s t e n t ; sepals  r e f l e x e d i n o l d e r f l o w e r s ; ovary  to  (1/10) 1/5  to  fall;  spreading 1/3  Columbia; Rocky Mountains o f A l b e r t a ; Cypress  Hills  7.  of Saskatchewan; Mt. R a i n i e r a r e a i n Washington; n o r t h e r n Washington and  Idaho; c e n t r a l Idaho; Rocky Mountains i n Montana,  Oregon, and n o r t h e r n Wyoming; and o c c a s i o n a l l y as  1.  f a r s o u t h as New  i n the Rocky Mountains  Mexico...  3. S.  S a x i f r a g a m a r s h a l l i i Greene s s p . m a r s h a l l i i , P i t t o n i a , TYPE C.C. M a r s h a l l s.n., Hoopa V a l l e y , Apr.  1887.  northeastern  Humboldt  occidentalis  1: 159.  Co., C a l i f o r n i a ,  ND!  IS. p e t i o l a t a A.M.  Johnson, Minn. Stud. B i o l .  S c i . 4: 29.  1923.  TYPE T. Howell s.n., Oregon, on wet banks, W o o d v i l l e , May, WTU!  Isotypes:  NY I UC!  Micranthes m a r s h a l l i i S_. m a r s h a l l i i  (Greene) S m a l l , N. Am.  Greene v a r . m a r s h a l l i i  P f l a n z e n r . IV, 117, 1: 36.  ann. 1871.  S_. l a e v i c a u l i s A.M.  f . c r e n a t a E n g l . & Irmsch.  1916. 1923.  TYPE:  E. H a l l  MGB.  Johnson, Minn. Stud. B i o l .  S c i . 4: 26.  P i p e r 5061, Mt. Grayback, Oregon, June 15,  _S. m a r s h a l l i i  1905.  1916.  Johnson, Stud. P I . S c i . 4: 24.  151, Oregon  TYPE:  F l o r a 2 2 ( 2 ) : 140.  f . d e n t a t a E n g l . & Irmsch.  Greene v a r . m a r s h a l l i i  S_. h a l l i i A.M.  1889.  (2) CAN!  P f l a n z e n r . IV, 117, 1: 36. S^. m a r s h a l l i i  1888.  1923.  1904.  Greene v a r . d i v a r i c a t a Peck, L e a f l . West. Bot. 5: 59.  1947.  TYPE:  Peck 18158, Snake  Battle  Creek, Wallowa Co., Oregon, March 28,  Stem f r e q u e n t l y s o l i t a r y  (2-3) from  with reddish purple glandular hairs with  River  Canyon, n e a r mouth o f 1934.  (0.5) 1 t o 3 (4) dm l o n g ,  covered  m u l t i c e l l u l a r heads; l e a v e s  oblong-ovate t o e l l i p t i c - o b l o n g , upper s u r f a c e g l a b r o u s , lower s u r f a c e  8.  o f t e n rusty-tomentose, e s p e c i a l l y s m a l l e r l e a v e s , margins c r e n a t e wide, b l u n t t e e t h  at r i g h t  angles  t o t h e margin, b l a d e  (1) 1.5 t o 3.5 cm b r o a d , t r u n c a t e to s u b t r u n c a t e the  (0.5)  with  2 to 6 cm l o n g ,  a t base, t a p e r i n g i n t o  1 t o 3 times l o n g e r p e t i o l e ; i n f l o r e s c e n c e an open d i f f u s e  p a n i c l e , branches s l e n d e r , d i v a r i c a t e , subtended by s m a l l , b r a c t s which a r e o c c a s i o n a l l y l e a f l i k e and toothed,  linear  p e d i c e l s 3 t o 4 times  l o n g e r than f l o w e r s ; s e p a l s s t r o n g l y r e f l e x e d , o v a t e - l a n c e o l a t e  to oblong-  l a n c e o l a t e , 2 t o 3 times as l o n g as b r o a d ; p e t a l s deciduous, oblong to ovate,  2.5 t o 3.5 mm l o n g , narrowed i n t o a s h o r t d i s t i n c t  claw, 2 y e l l o w  spots on base o f p e t a l , mostly faded i n h e r b a r i u m m a t e r i a l ; f i l a m e n t s p e r s i s t e n t , spreading,  e q u a l t o o r s h o r t e r than t h e p e t a l s , s t r o n g l y  c l a v a t e t o p e t a l o i d , o f t e n showy a f t e r d i s t i n c t t o base, f l u t e d  nectary  petals  fall;  a t base appearing  the 2 (3) c a r p e l s as wavy r i b b o n as  capsules mature, immature c a r p e l s l o n g and t h i n , mature tips divergent;  ovary almost completely  carpels erect,  s u p e r i o r , n_ = 10 (Table 1; F i g . 1 ) .  M o i s t banks b e s i d e streams i n n o r t h western  C a l i f o r n i a and western  Oregon ( F i g . 6 ) . Representative OREGON:  Specimens BENTON Co.: Mary's Peak, C o r v a l l i s , E p l i n g s.n. R., 4 mi from A l l e g a n y ,  Cronquist  (ORE); COOS  Co.:  e. f o r k o f M i l l i c o m e s  6859  (WTU,  WS, ID, OSC, CAN, UC, DAO, UBC, MONTU); Sitkum, Krause and Beamish  30-70 (UBC); Dora, Krause and Beamish 10-70 (UBC); Bear Ck., middle f o r k o f C o q u i l l e R., Krause and Beamish 35-70 (UBC); CURRY Co.:  Two M i l e  Creek, Ih mi s. o f I l l a h e , Baker 5264 (WTU, WS, OSC, UC); F r y e P l a c e , 3 mi s. o f Agness, Baker 3513 (OSC, UC); DOUGLAS Co.:  Gardiner,  Overlander s.n.  (OSC); Mapleton, Krause and Beamish  36-70 (UBC); between N o r t h Bend and  Roseburg, Mason 5290 (UC, OSC); JACKSON Co.: Howell s.n. (NY, UC, 10 mi w.  on wet banks, W o o d v i l l e ,  CAN); Weimer, Hammond 125b  (NY); JOSEPHINE Co.;  of Selma, O v e r l a n d e r s.n. (OSC); LANE Co.:  W i l l a m e t t e Pass  between Dexter and Oakridge, Packard and G i l k e y s.n. (UBC, OSC); w. o f T r i a n g l e Lake, G i l k e y s.n. (WS, WTU); 15-71  (UBC); LINCOLN Co.:  S c o t t s b u r g , Krause and  Waldport-Yachats, Smith s.n. (OSC); wet  s. s i d e Oswego L., N e l s o n 2550 (WS); POLK Co.: Peck 16213  DEL NORTE Co.:  UC); HUMBOLDT Co.:  2.  C., 4 m sw.  Gasquet, Pack and T r a c y 12351  W i l l o w Creek, T r a c y  2 mi s e . of D e v i l ' s H o l e , T r a c y 15511 (UC, JEPS) ; MENDOCINO Co.:  E e l R.,  Mill  cliffs  of B u e l l  (UC).  CALIFORNIA:  8011  Beamish  (DAO,  6644 (UC, WTU); T r i n i t y Summit,  (UC); TRINITY Co.: S a l y e r , T r a c y  R e e l Mt. Creek b l u f f , on s. f o r k o f  T r a c y 16530 (UC) .  S a x i f r a g a m a r s h a l l i i Greene s s p . i d a h o e n s i s  ( P i p e r ) Krause and  Beamish, comb. nov. S_. i d a h o e n s i s P i p e r , B u l l . T o r r e y Bot. Club, 27: 394.  1900.  TYPE:  Henderson 4588, K e n d r i c k , L a t a h Co., Idaho, Apr.  1896.  GH!  ISOTYPE:  NYI' Photograph:  S_. m a r s h a l l i i P i p e r v a r . i d a h o e n s i s P f l a n z e n r . IV, 117, 1: 37. S_. m i c r o c a r p a A.M. TYPE:  M.F.  26,  WSPl  ( P i p e r ) E n g l , and Irmsch.,  1916.  Johnson, Minn. Stud. B i o l . S c i . 4: 28.  Ehod 98a, M i s s o u l a v i c i n i t y .  S_. o c c i d e n t a l i s S. Wats. v a r . i d a h o e n s i s  1923.  Mont. MONTU!  ( P i p e r ) C.L.  Hitch.,  V a s c u l a r P l a n t s o f the P a c i f i c Northwest., Univ. Wash. P r e s s , H i t c h c o c k e t a l . IV.  49.  1961.  10.  Caudex s h o r t and  stout,  rhizome s h o r t , o b l i q u e w i t h  stem, 1.2-2  (2.5)  dm  l e a f blades  from  ovate to e l l i p t i c  1 (2)  flowering  t a l l w i t h dense to s p a r s e p u r p l i s h g l a n d u l a r h a i r s ; to rhombic, glabrous  on upper s u r f a c e  and s p a r s e l y r u s t y tomentose on lower s u r f a c e , margin i r r e g u l a r l y dentate,  blade  from 2.3  to 3 cm l o n g , 1.8  to 2.7  cm wide w i t h  an  attenuate  (obtuse) base; p e t i o l e from s l i g h t l y s h o r t e r than to twice as l o n g blade;  as  i n f l o r e s c e n c e g e n e r a l l y an open, d i f f u s e p a n i c l e , branches  slender,  d i v a r i c a t e , p e d i c e l s from 2 to 3 times as l o n g as f l o w e r s , f l o w e r s i n compact c l u s t e r s at ends of b r a n c h e s ; b r a c t s s m a l l , l i n e a r , p i l o s e , o c c a s i o n a l l y l e a f y and  toothed;  times as l o n g as b r o a d , s l i g h t l y deciduous, oblong to ovate, spots p r e s e n t  glandular-  sepals ovate-lanceolate,  2.5-3  to s t r o n g l y r e f l e x e d a t a n t h e s i s ; p e t a l s  2-3.5  mm  l o n g , narrowed i n t o a claw, 2 y e l l o w  a t base ( r a r e l y seen i n p r e s s e d m a t e r i a l ) ; f i l a m e n t s p e r s i s -  t e n t , c l a v a t e to s t r o n g l y c l a v a t e , about e q u a l l i n g the almost completely nectary present Fig.  seldom  s u p e r i o r to 1/5  a t a n t h e s i s but  p e t a l s ; ovary from  i n f e r i o r ; 2 (3) c a r p e l s , e l o n g a t e  r a r e l y v i s i b l e at m a t u r i t y ; n = 10  thin; (Table  1;  2). Moist  rocky s l o p e s , K i t t i t a s and  Idaho, e s p e c i a l l y i n low drainage  systems; and  l y i n g areas  adjacent  Chelan Cos.,  Washington; western  o f the S t . Joe and  northeast  Clearwater  River  Oregon ( F i g . 6 ) .  Type - A l l the specimens P i p e r examined i n e s t a b l i s h i n g h i s s p e c i e s were from western Idaho.  He s t a t e d t h a t the f i l a m e n t s were  p e r s i s t e n t , a t l e n g t h e x c e e d i n g the p e t a l s . " a s k i n g him  i f the d e s c r i p t i o n of " s u b u l a t e  r e p l i e d " I t i s evident  In 1902  J.K.  f i l a m e n t s " was  t h a t the d e s c r i p t i o n  of  "subulate, S m a l l wrote  an e r r o r .  the f i l a m e n t s as  He  'subulate'  was  a blunder."  ( L e t t e r s on NY herbarium  sheet.)  In h i s type  he a l s o s t a t e d t h a t S_. i d a h o e n s i s was r e l a t e d to dentalis .  description  r e f l e x a and  occi-  As most, e a r l y S_. m a r s h a l l i i specimens from the U n i t e d S t a t e s  w i t h r e f l e x e d s e p a l s were c a l l e d S_. r e f l e x a , t h i s f i t s v e r y w e l l w i t h  the  h y p o t h e s i s t h a t S_. i d a h o e n s i s i s c l o s e l y r e l a t e d to S_. m a r s h a l l i i but  has  some c h a r a c t e r i s t i c s o f S^. o c c i d e n t a l i s . JS. m i c r o c a r p a idahoensis.  appears  t o be a s m a l l e r p l a n t of ;S. m a r s h a l l i i s s p .  Johnson s t a t e s i t i s o n l y found  M i s s o u l a , Montana.  He examined o n l y two  from the type  locality,  specimens.  R e p r e s e n t a t i v e Specimens WASHINGTON: T23N, R18E  S21,  CHELAN Co.: CL.  M o i s t draw above Camas Land Meadow, s. s i d e ,  H i t c h c o c k and C.V.  M u h l i c k 21542 (WTU); rocky  s l o p e s on S t u a r t fiidge above I n g a l l s , J.W. Thompson 8968 (NY,  moist  WTU);  s t e e p rocky h i l l s i d e , n. exposure above Swakane Cr., about 5 mi e. o f d i v i d e t o Nahukum Cr. , C L . s l o p e s on Lookout Mt.,  H i t c h c o c k 23440 (UC, WSP, WTU);  (OSC,  WTU);  8294 (NY, J.  wet  rocky  near Leavenworth, J.W. Thompson 6465 (OSC); above  P e s h a s t i n Cr., ca. 6 mi n. o f Blewett Pass, C L . 4738 (NY, WTU);  moist  H i t c h c o c k and J.S.  g r a n i t e s l o p e s near Leavenworth, J.W. Thompson  Martin 6411  moist s l o p e s of h i g h r i d g e above Thronson Cr., J.W. Thompson UC);  10 mi n. o f Swaux Pass on Hy.  Maze e t a l . 340 (CAN); KITTITAS Co.:  97,  ca. 10 mi sw.  o f Cashmere,  moist rocks a l o n g Boulder  Cr.,  J.W.  Thompson 10422 (WTU); 1 mi below Camp Cr., upper C l e Elum R i v e r V a l l e y ,  A.R.  Kruckeburg  Rd.,  ca. 5 mi n. o f Colochum Pass, J . Maze e t a l . 398 (CAN); on moist  ca.  2 mi nw.  (WTU).  4984 (UC); Sec. 4, T. 20N,  R. 21E,  of M i s s i o n Peak, Wenatchee Mts.,  CL.  on  Ellensburg-Wenatchee  H i t c h c o c k 20346  cliffs  12.  IDAHO:  IDAHO Co.:  V. A l l e n s.n. J.H.  ( I D ) ; 2 mi s. o f Ahsahka, on c l i f f s  C h r i s t and W.W.  6265 (NY, WTU);  Ward 12107  Reed and F.A.  (NY); moist  a t summit of John Beale Rd.,  B a r k l e y 84  along Clearwater  R.,  (NY); Lewiston, moist h i l l s i d e , J.H.  Hapwai, D. McDougal 93  g r a v e l on south c l i f f M.J.  a l o n g the Selway R. near Fenn Ranger S t a t i o n ,  situations i n granite L o l o Nat. F o r e s t .  (MONTU); e. s l o p e of mountain, 4000 f t s. of  mouth o f Sheep Cr., Snake R i v e r Canyon, S. Beinke  131  (WTU); rocky b l u f f s  i n c o n i f e r o u s shade on Meadow Cr., above Selway F a l l s , L. Constance R.C. M.  Rollins  1638  (WS,  Ownbey and R.P.  WTU,  ALTA); n. s l o p e , s. o f Selway R.,  Ownbey 2040 (WTU,  f o r k o f C l e a r w a t e r R.  Small  UC, WS,  NY);  and  Selway  Falls,  4 mi below L o w e l l , M i d d l e  at Three D e v i l s Cr., L. Constance  e t a l . 1083  (WS,  UC);  m o i s t , s o u t h - f a c i n g s l o p e , 1 mi up G r a n i t e Cr., Snake R. Canyon, J . Packard 372  (UC); on r o c k o u t c r o p p i n g near Selway R.,  J . L . McMullen 1487  (UC, I D ) ; LATAH Co.:  4 mi s. of J u l i a e t t a , M. Ownbey and R.P. WS,  NY,  OSC,  water R.,  ALTA); NEZ  PERCE Co.:  near M y r t l e , W.H.  near Culdesac, F.A. C l e a r w a t e r R.,  a l o n g C l e a r w a t e r R.,  Ownbey 2032 (COLO, DAO,  (WS);  Christ  c l i f f w.  H i t c h c o c k and C.V.  OREGON:  WALLOWA Co.:  B a t t l e Cr., M.E. Snake R.  Peck 18161  Canyon n. o f the  moist rocky s l o p e ,  Baker 6606 (UC); t a l u s s l o p e s ,  1 mi e. o f S p a u l d i n g , W.H.  a l o n g S t . Joe R.,  UC,  moist rocky c l i f f , canyon of the  on r d . t o Nez P e r c e , J.H.  on c l i f f s  WTU,  Cr.  s t e e p rocky h i l l s i d e , along the C l e a r -  o f M y r t l e , W.H.  o f Twin L., C.L.  Falls,  moist n. s l o p e a l o n g P o t l a t c h  Baker 6606 (JEPS, WTU);  Warren s.n.  4 mi sw.  2 mi up from Selway  Baker 13909 ( I D ) ; Onfino^  7537 (NY); SHOSHONE Co.:  moist  M u h l i c k 21745 (WTU,  7 mi s. o f Culdesac, J.H. rocky s l o p e , Snake R.  slate WS,  C h r i s t 10958 (ID,  NY)  Canyon near mouth o f  (NY); on b a s a l t ledge a t L i t t l e Creek, mouth of the Salmon R.,  NY);  W.A.  Weber 2103  i n the (COLO);  moist rocky n. h i l l s i d e , MONTANA:  RAVALLI Co.:  B i t t e r Root Mts., K.H. L o s t Horse 1215  Cr. Rd.,  on bedrock  L a c k s c h e w i t z and  Saxifraga occidentalis  S_. saximontana  (MONTU); o f f the  Fageraas 477  S. Wats., P r o c . Am.  Fageraas  L., L o l o Peak, B i t t e r  (MONTU).  Acad. 23: 264. B.C.,  May  17,  1888. 1875.  CAN! E. N e l s o n , E r y t h e a , 7: 168.  N e l s o n 5917, NY!  R.  1264  above C a r l t o n  UC).  above Glen L.,  L a c k s c h e w i t z and R.  J . Macoun s.n., Y a l e Mt.,  Specimen A,  Jones 6295 (WS,  Lackschewitz and R. Fageraas  B i t t e r Root Mts. K.H.  LECTOTYPE:  G.N.  i n mossy c r a c k s on bedrock  (MONTU); MISSOULA Co.:  Root Mts., K.H.  3.  mouth o f Imnaha R.,  1899.  TYPE:  Yancey's Y e l l o w s t o n e Nat. Park, J u l y  ISOTYPES:  WSP!  A. and E.  17,  1899.  RM!  Micranthes o c c i d e n t a l i s  (S. Wats.) S m a l l , N. Am.  F l o r a , 22(2):  144.  1905. M.  allenii 242,  l a t a S m a l l , N. Am.  Photograph: saximontana  S_. a l l e n i i  1905.  TYPE:  1896.  GH!  1905.  TYPE:  O.D.  ISOTYPES:  Allen NY!  WTU! F l o r a 2 2 ( 2 ) : 144.  s_.n. , L y t t o n , B.C.,  M.  F l o r a , 2 2 ( 2 ) : 144.  Goat Mts., Washington, June 27,  US! WS! M.  S m a l l , N. Am.  Apr.  16,  1889.  NY!  ISOTYPES:  John Macoun GH!  CAN!  DAO! (E. N e l s . ) S m a l l , N. Am.  F l o r a , 2 2 ( 2 ) : 145.  (Small) Fedde, J u s t ' s Bot. J a h r e s b e r . 31(1): 613.  S_. l a t a (Small) Fedde, J u s t ' s Bot. J a h r e s b e r . 31(1): 613.  1905. 1906. 1906.  S_. o c c i d e n t a l i s S. Wats. v a r . w a l l o w e n s i s Peck, L e a f l . West. Bot. 4(4)60.  1947.  TYPE:  Peck 18542, moist s l o p e above I c e Lake,  Wallowa Mts., Wallowa Co.,  Oregon, J u l y 4,  1934.  14. S_. o c c i d e n t a l i s S. Wats. v a r . a l l e n i i  C.L. H i t c h c . , V a s c u l a r  of the P a c i f i c Northwest. Univ. Wash. P r e s s , III:  49.  Plants  Hitchc. et a l . ,  1961.  S_. o c c i d e n t a l i s S. Wats. v a r . l a t i p e t i o l a t a H i t c h c , V a s c u l a r  Plants  o f the P a c i f i c Northwest. U n i v . Wash. P r e s s , H i t c h c . e t a l . I l l : £49.  1961.  TYPE:  M.W.  Oregon, June 20, 1915.  Gorman 3561, Saddle Mt., C l a t s o p Co., WTU!  Stems 1 ( 2 - 3 ) , from 0.8 to 2 (2.5) dm t a l l w i t h p u r p l i s h glandular h a i r s ; l e a f blades oval-ovate  dense t o s p a r s e  v a r i a b l e i n s i z e and shape, from  t o rhombic, upper s u r f a c e s g l a b r o u s ,  lower s u r f a c e s sometimes  rusty-tomentose, margins s e r r a t e t o c r e n a t e , b l a d e  from 1.5 to 3 cm l o n g ,  1.2 t o 2.5 cm wide t a p e r i n g i n t o a p e t i o l e e q u a l t o , t o twice as l o n g as the b l a d e ;  i n f l o r e s c e n c e an open p a n i c l e , from c o n s i d e r a b l y c o n s t r i c t e d  w i t h s h o r t e r e c t branches t o open w i t h s p r e a d i n g branches, glomerate t o rounded o r p y r a m i d a l ; flowers and to  p e d i c e l s e q u a l l i n g twice the l e n g t h o f f l o w e r s ,  grouped a t ends o f branches; b r a c t s s m a l l and  ( o r ) rufose-tomentose; s e p a l s ovate t o o b l o n g - o b l a n c e o l a t e ,  toothed  spreading  r e f l e x e d ; p e t a l s w h i t e , immaculate, deciduous o r p e r s i s t e n t , ovate to  oblong, 1.5 to 3.5 cm l o n g w i t h subulate,  linear,  or s l i g h t l y  a distinct  claw o r c l a w l e s s ,  t h e i r length at maturity;  filaments  c l a v a t e , p e r s i s t e n t , e q u a l l i n g the p e t a l s ;  c a r p e l s 2 (3) narrowing g r a d u a l l y i n t o l o n g s t y l e s ,  at  l i n e a r , often  s h a r p l y divergent  ovary from 1/10 to 1/3 i n f e r i o r ; n e c t a r y  a n t h e s i s , b u t d r y i n g a f t e r a n t h e s i s and d i f f i c u l t  f o r 3/4  visible  t o s e e ; n = 10, 19, 28, 29  (Table 1; F i g s . 3 and 4 ) . M o i s t banks and meadows t o rocky  alpine slopes.  B r i t i s h Columbia; Rocky  15.  Mountains i n A l b e r t a ; Cypress H i l l s i n Saskatchewan; area around Mt. R a i n i e r , Washington; n o r t h e r n Washington and Idaho; c e n t r a l Idaho, Rocky Mountains i n Montana and northwest Wyoming and n o r t h e a s t e r n Oregon and o c c a s i o n a l l y the Rocky Mountains as f a r s o u t h as New Mexico  (Fig.  7).  Type - No type specimen has been d e s i g n a t e d f o r J3. o c c i d e n t a l i s . Watson (1888) c i t e d s e v e r a l specimens f o r the range o f S_. o c c i d e n t a l i s "From t h e Rocky Mountains o f B r i t i s h Columbia and Vancouver I s l a n d  America (Drummond) t o B r i t i s h  ( L y a l l , Macoun), Oregon  ( C u s i c k , Henderson,  Howell) and t h e n o r t h e r n S i e r r a Nevada ( C h i c o , Mrs. J . B i d w e l l , G r a y ) . " I t i s g e n e r a l l y a c c e p t e d t h a t the m a t e r i a l from Vancouver I s l a n d and Oregon b e l o n g t o o t h e r s p e c i e s . Island  One o f Macoun's specimens from Vancouver  (Mt. F i n l a y s o n , V . I . , Macoun s.n. DAOl NY! CAN!) has been  nated as the h o l o t y p e o f S_. r u f i d u l a ;  desig-  the m a t e r i a l c o l l e c t e d by Howell  from t h e Columbia R i v e r Gorge i s most l i k e l y S_. r u f i d u l a  (high  cliffs  a l o n g t h e Columbia R i v e r a t Multnomah F a l l s , J . H o w e l l s.n. NDl PHI), and a l l t h e m a t e r i a l from s o u t h e r n Oregon i s most l i k e l y S_. m a r s h a l l i i as j3. o c c i d e n t a l i s does n o t grow as f a r s o u t h as s o u t h e r n Oregon P a s s , J . Howell s_.n. NY!). as  (Grant's  We have chosen one o f Macoun's specimens  the l e c t o t y p e f o r _S. o c c i d e n t a l i s :  Y a l e Mountain, B.C., May 17,  1875,  J . Macoun s.n. CAN! ( F i g . 5, specimen A ) .  Representative BRITISH  Specimens  COLUMBIA:  2 mi ne. o f Salvus on P r i n c e Rupert - T e r r a c e  Highway, J.A. C a l d e r , D.B.O. S a v i l e and J.M. Ferguson 13117 (DAO); P a r a d i s e Mine 15 mi w. o f Windermere,  J.A. C a l d e r and D.B.O. S a v i l e 11232  (DAO); S k a g i t V a l l e y , B.C. - Washington  boundary  to 10 mi n., H. Rhodes  16.  and  C. Brayshaw 6641 (DAO); B l u f f Lake about 12 mi ssw. o f T a t l a Lake  P.O., 51°46'N 124°43'W, J.A. C a l d e r , J.A. Parmelee, R.L. T a y l o r 17263 (DAO); Kimberley, F. Fodor 119 (UBC); a l o n g Hwy. 3, about 2 mi w. of N e l s o n , W.C. M c C a l l a K.I.  8127 (UBC); H o p e - P r i n c e t o n Hwy., n. o f m i l e 126,  Beamish e t a l . 7638 (UBC); B l a c k w a l l , Manning Park, K.I. Beamish  and F. Vrugtman 60792 (UBC, DAO); r.ocky mountain summit,  Liumchen a r e a ,  above c a b i n , C u l t u s L. a r e a , K. Beamish 7910 (UBC); Tod Mt., Kamloops, T.M.C. T a y l o r and A.F. S z c z a w i n s k i  784 (UBC); C o r r i e Ridge, G a r i b a l d i ,  A.E. Beggs s.n. (UBC); Empetrum Ridge, G a r i b a l d i P r o v i n c i a l Park, K.I. Beamish 680176 (UBC); Mt. B r e n t , P e n t i c t o n , J.W. Eastham s.n. (UBC); C a t h e d r a l Lakes, Ashnola D i s t . , T.M.C. T a y l o r 1346 (UBC, WS, UC); P r i n c e t o n , T.M.C. T a y l o r 2053 (WS); M i l e 19, F l a t h e a d Rd., K.I. Beamish 156 (UBC, DAO); Goat Mt., 49°58'N 114°42'W, R.L. T a y l o r and D.M. Ferguson 3039 (DAO); Moosehorn L., 58°10'N 132°07'W, R.J. P i l f r e y 57 (DAO); 3 mi s . o f Camp 7, 18 mi sw. o f Anahim L., D.W. Hyndman and H.W. 27 (DAO); s u b a l p i n e s l o p e s  Tipper  near Lake B o t a n i e , Marble Mts. 5000 f t ,  J.W. Thompson and E.M. Thompson 70 (UC, WS); c r e v i c e s o f r o c k s , Mt. Queert, B.C., Macoun 8375 (CAN); S k a g i t R., Macoun 79347 (CAN); Crow's Nest P a s s , Rocky Mts., J.M. Macoun 20142 (CAN). The f o l l o w i n g f i v e specimens from B r i t i s h Columbia have open i n f l o r e s c e n c e s ,  partially  r e f l e x e d s e p a l s , and l e a f shapes which make them resemble some o f t h e S. m a r s h a l l i i s s p . i d a h o e n s i s  plants  from Idaho:  6 mi n. o f C l e a r -  water S t a t i o n , North Thompson R i v e r a t Bear Creek F a l l s , J.A. C a l d e r , J.A. Parmelee and R.L. T a y l o r 17423 (DAO); 1 mi sw. o f Midway, J . Grant 38 (DAO); Goat R i v e r Canyon on E r i c k s o n Mt. near C r e s t o n , J.A. C a l d e r  17. and D.B.O. S a v i l e 7735 (DAO); 3 mi e. Highway, J.A.  C a l d e r and K.W.  of South S l o c a n  S p i c e r 32877 (DAO).  on  Nelson-Trail  The f o l l o w i n g  plant  had e x c e s s i v e l y l a r g e broad l e a v e s which g i v e i t a s t r a n g e appearance: 38 mi on road from Anahim L. to B e l l a C o o l a , 52°25'N C a l d e r , J.A. Parmelee and R.L. ALBERTA:  125°53'W,  T a y l o r 17338 (DAO);  Shoulder o f W i l c o x Mt.,  (CAN); B a n f f Nat. Park., n e a r l o o k o u t  J a s p e r Nat.  P a r k , E.H. Moss s.n.  p o i n t above Peyto L., W.C.  7073 (ALTA); head o f L. L o u i s e , Rocky Mts., J.M.  Macoun 65316  rocky s l o p e above B e r t h a L., Waterton Lakes Nat. Park, C O . 3618  (ALTA);  Park, W.C. R.A.  M c C a l l a 11673  (CAN);  Rosendahl  (ALTA); Goat Cr., e. f l a n k o f F l a t h e a d Range,  36 mi due n. o f Coleman, E.W. SASKATCHEWAN:  Mountjoy  of I s o l a t i o n  ( I s o l a ) Peak  1 (DAO);  Cypress H i l l s Park., dry h i l l s i d e ,  A.J. Breitung  (DAO); WASHINGTON:  V a l l e y of War (WTU,  WS,  OKANOGAN Co.:  UC); sandy  (WS, WTU);  the  P a s a y t e n R., UC, OSC,  Range, J.W.  m o i s t rocky l e d g e s , Sawtooth  Cr., H. S t . John, W.D.  263  WTU,  McCalla  edge of woods above Red Rock Canyon, Waterton Lakes Nat.  P r i c e 10 (DAO); Savannah Cr. j u s t w.  4212  J.A.  hillside,  Courney and C S .  Packer 3822  Ridge, Ptarmigan C r . , Hidden L s . , O.T.  rocky o u t c r o p below S t a t e Pk.,  at the m i d d l e f o r k o f  Cascade Mts., M. Ownbey and F . C  WS);  Ridge,  Meyer 2299  (ALTA,  exposed open summit of Muckamuck Lookout, T i f f a n y  Thompson 6985 (WTU); near Ruby Grade, on Salmon Cr., Ruby,  C.B.  F i k e r 1183  (WS,  W.N.  Suksdorf 9671  WTU);  SKAMANIA Co.:  Wettenburg,  (WSP); c r e v i c e s o f c l i f f s  P a r k , Mt. R a i n i e r , J.W.  Thompson 7614  Chequash Mts.,  under w a t e r f a l l s , P a r a d i s e  (WTU); SPOKANE Co., damp r o c k y  Edwards  18.  h i l l s i d e near L a t a h C r . , s o u t h e a s t of Spangle, W. WS,  WTU);  Mts.,  YAKIMA Co.:  J.W.  Thompson 9841  IDAHO:  L., Sawtooth  s l o p e s i n Chinook P a s s , Cascade  B o u l d e r Ck.  Canyon, Sawtooth Nat. F o r e s t ,  Thompson 14056 (NY, WTU);  Primitive A r e a ,  BOISE Co.:  A l p i n e Ck.,  C.L. H i t c h c o c k and C.V.  M u h l i c k 9872 (NY); BONNER Co.:  Mt.,  J.H.  Porthill,  p r o t e c t e d s l o p e s , Josephus UC); s a d d l e w.  For., w.  Christ  C.L. H i t c h c o c k and C.V.  and C.V. about  WS);  of  B o u l d e r Mts.,  and E.B.  dry  s l o p e above Storm L., Anaconda Mts.,  Davis 674  (WS);  meadow on  M u h l i c k 9514 DEERLODGE  (UC,  Co.: Muhlick  base o f summit of mountains ne. of l a k e ,  Payson and L.B.  Payson 2059 (NY, UC);  IDAHO Co.:  East  on  f o r k o f Sheep  (WS).  BEAVERHEAD Co.:  C.L. H i t c h c o c k and C.V. Ajax L.,  WS);  Nat.  C.L. H i t c h c o c k and C.V.  d i v i d e between head of Sheep Cr. and head o f the  MONTANA:  Payson 3549  i n semiswampy s o i l i n rocky canyon  L i v i n g s t o n Mine, R.J".  J . Packard 344  moist,  Sawtooth Nat. F o r . , C.L. H i t c h c o c k  WTU,  Cr.,  Continental  CUSTER Co.:  C h a l l i s , C.L. H i t c h c o c k and C.V.  14820 (NY); FREMONT Co.:  Spr.,  White Cloud Range, C h a l l i s  NY,  Henry L., E.B.  (NY):  M u h l i c k 10869 (NY, WTU,  M u h l i c k 10647 (NY, WTU);  10 mi w.  1678  L s . , J . F . Macbride  o f C a s t l e Pk.,  base o f Ryan Pk.,  of A l t u r u s  on rocks 5 mi n. o f E l k L., Sawtooth P r i m i t i v e  C.L. H i t c h c o c k and C.V.  (NY,  nw.  M u h l i c k 10505  A r e a , headwater o f S. Fork P a y e t t e R. above Sacajawea Hot  25 mi w.  (UC,  (WTU);  BLAINE Co.:  Sawtooth Mts., J.W.  (NY, WS);  crags and moist  Suksdorf 8616  on top of O d e l l Peak, P i o n e e r Range,  M u h l i c k 14925 (WTU,  NY,  WS);  C i r q u e above  j u s t below summit of C o n t i n e n t a l D i v i d e , on g r a n i t e t a l u s  and  19.  r o c k , B i t t e r Root Range, C L . H i t c h c o c k and C.V. M u h l i c k WTU);  12679 (WS,  E a s t P i n t l a r Peak, head of P i n t l a r C r . , Anaconda Range, C L .  cock and C.V. M u h l i c k 12846 (WTU, NY, WS);  w.  Hitch-  t a l u s s l o p e above T o r r e y  L., P i o n e e r Range, C L . H i t c h c o c k and C.V. M u h l i c k DEERLODGE Co.:  NY,.  15056 (WTU,  NY);  s l o p e of Mt. T i n y , s. o f Storm L., Anaconda Mts.,  C L . H i t c h c o c k and C.V. M u h l i c k  14860 (WTU, WS) ; FLATHEAD Co.:  r i d g e , Mt. Aeneas, Swan Range, L.H. Harvey 3283 (MONTU); Mt.  on  Cannon  above McDonald C r . , G l a c i e r Nat. Park, L.H. Harvey 6050 (MONTU); i n seep below snow bank on t a l u s s l o p e , Logan Pass, G l a c i e r Nat. Park, L.H. Harvey 6689 (MONTU); h i g h above C L . H i t c h c o c k 18300 (WS, WTU, slide, 11425  and s e . o f McDonald L., M i s s i o n Range,  UC, COLO, NY,  I D ) ; MISSOULA Co.:  g r a n i t e r o c k , 5 mi above Bonner, C L . H i t c h c o c k and C.V.  talus Muhlick  (WTU, NY, WS) ; h mi w. o f Upper H o l l a n d L., F l a t h e a d Range,  H i t c h c o c k 18422 (WTU, WS); POWELL Co.:  CL.  top of Gordon Mt., 6 mi s. B i g  P r a i r i e Ranger S t a t i o n , F l a t h e a d Nat. F o r . , C L . H i t c h c o c k 18848 (WTU). WYOMING:  rocky p a r t of Canyon C r . , Grand Teton Nat. Park, W.C.  4836 (ALTA); on mossy rocks  i n s p r a y of f a l l s ,  Crazy C r . , on Red Lodge-  Cooke C i t y Highway, Park Co. , C L . P o r t e r 5437 (DAO, WTU); hillside,  Cascade Canyon, L. W i l l i a m s 1135  Park, moist  rocky c l i f f s  McCalla  moist  mossy  (ORE, ID); G l a c i e r Nat.  near Cracker L. t r a i l , D.H.  Brant 45 (WTU).  20. DISCUSSION  The  r e l a t i o n s h i p of the  taxa p r e v i o u s l y  m a r s h a l l i i , S_. i d a h o e n s i s , and f e e l that  S^. o c c i d e n t a l i s has  t h i s r e l a t i o n s h i p i s b e s t e x p r e s s e d by  _S. m a r s h a l l i i sis  (Piper)  Greene s s p .  D.L.  Saxifraga  Krause and  K.I.  Beamish, and  areas of Oregon and  ton,  the  following  S_. o c c i d e n t a l i s  Montana.  Saxifraga  adjacent A l b e r t a ,  occasionally  We  taxonomy: idahoen-  S. Wats.  Oregon  i d a h o e n s i s i n Chelan  i n northcentral  Idaho and  Occident alis  and  and  adjacent  grows i n B r i t i s h  Cypress H i l l s of Saskatchewan, Washing-  Idaho, n o r t h e a s t e r n Oregon, the  Wyoming, and  Saxifraga  been d o u b t f u l .  m a r s h a l l i i s s p . m a r s h a l l i i grows i n western  Counties of Washington and  Columbia and  to as  m a r s h a l l i i , J 3 . m a r s h a l l i i Greene ssp.  n o r t h w e s t e r n C a l i f o r n i a , S_. m a r s h a l l i i s s p . Kittitas  referred  Rocky Mountains o f Montana  the Rocky Mountains  as  f a r south as  and New  Mexico. Saxifraga  m a r s h a l l i i ssp.  from J5. o c c i d e n t a l i s .  m a r s h a l l i i can be  However, s s p .  separated quite  i d a h o e n s i s i s much  more  readily  difficult  to s e p a r a t e from S_. o c c i d e n t a l i s because, where t h e i r ranges meet, two  t a x a have c h a r a c t e r i s t i c s which tend to approach each o t h e r .  example^ i n Idaho some s s p . p a r t i a l l y i n f e r i o r ovaries, rhombic l e a v e s and idahoensis. d e n t a l i s by  filaments  that  idahoensis plants  can be  former's r e f l e x e d s e p a l s ,  deciduous n a t u r e of the  petals,  persistent  found which  are not  more congested i n f l o r e s c e n c e s  These s s p . the  i d a h o e n s i s specimens are  yellow  strongly  than  the For  have  clavate,  typicalssp.  s e p a r a t e d from S_. o c c i spots on  filaments,  and  the  petals,  the  long  pedicels  of the f l o w e r s .  We b e l i e v e t h a t s s p . i d a h o e n s i s  to S_. m a r s h a l l i i than t o S_.  i s more c l o s e l y  related  o c c i d e n t a l i s and t h e r e f o r e cannot be c l a s s i -  f i e d as an i n f r a s p e c i f i c taxon o f S_. o c c i d e n t a l i s  ( H i t c h c o c k 1961) even  though t h e r e may have been some i n t e r g r a d a t i o n o f c h a r a c t e r i s t i c s w i t h  this  species. S a x i f r a g a i d a h o e n s i s has been reduced  to a subspecies  o f S_.  m a r s h a l l i i because some specimens o f s s p . i d a h o e n s i s a r e almost g u i s h a b l e from specimens o f s s p . m a r s h a l l i i . p o p u l a t i o n s have been s e p a r a t e d  indistin-  The Washington and Idaho  from the Oregon and C a l i f o r n i a  t i o n as a s u b s p e c i e s because the i n f e r i o r i t y of the o v a r y ,  popula-  open n a t u r e  o f t h e i n f l o r e s c e n c e , shape o f the f i l a m e n t s , and the l e a f shape s l i g h t l y i n most specimens.  The m o r p h o l o g i c a l  differ  characters i n ssp.  i d a h o e n s i s a r e skewed s l i g h t l y  towards those i n S_. o c c i d e n t a l i s .  tendency toward m o r p h o l o g i c a l  s i m i l a r i t y between the l a t t e r two taxa  c o u l d be the r e s u l t o f l i m i t e d gene f l o w between some p r e v i o u s s t a g e i n t h e i r e v o l u t i o n . p r e c l u d e s gene flow a t t h e The presence  present  The  the two taxa a t  Present  c y t o l o g i c a l information  time.  o r absence o f y e l l o w spots on the p e t a l s of f r e s h  m a t e r i a l appears to be a good c h a r a c t e r i s t i c f o r s e p a r a t i n g the subs p e c i e s o f S_. m a r s h a l l i i from S_. o c c i d e n t a l i s .  A l l the l i v e  of s s p . i d a h o e n s i s and s s p . m a r s h a l l i i had y e l l o w s p o t s , y e l l o w spots were observed  collections  whereas no  on c o l l e c t i o n s o f S_. o c c i d e n t a l i s .  Unfor-  t u n a t e l y , t h i s c h a r a c t e r i s t i c i s n o t u s e f u l f o r most d r i e d specimens as the y e l l o w s p o t s f r e q u e n t l y fade upon d r y i n g . The q u a n t i t a t i v e measurements taken from and  the herbarium  both  the l i v e  populations  specimens c o u l d n o t be used t o s e p a r a t e the taxa  22.  because the v a r i a b i l i t y overlapped The  within  f o r a l l measurements  each taxon was  so g r e a t t h a t the  taken.  chromosome counts s t r e n g t h e n  the h y p o t h e s i s  that ssp.  i s more c l o s e l y r e l a t e d to s s p . m a r s h a l l i i than i t i s to j>. As seen i n T a b l e  1, a l l the s s p . m a r s h a l l i i and  t i o n s counted had  10 p a i r s of chromosomes.  S_. o c c i d e n t a l i s w i t h  taxa  idahoensis  o ccidentalis.  ssp. idahoensis  Only one  popula-  population  10 p a i r s o f chromosomes has been  of  r e p o r t e d , and i t  i s from A l b e r t a , w e l l to the n o r t h of the range of s s p . i d a h o e n s i s . chromosome counts o b t a i n e d  f o r S_. o c c i d e n t a l i s i n d i c a t e t h a t the  g e n e t i c s o f t h i s s p e c i e s i s more c o m p l i c a t e d The  d i f f e r e n t counts of jS. o c c i d e n t a l i s c o u l d not be  morphological to be  differences.  More p o p u l a t i o n s  counted to d i s c o v e r the e x t e n t The  one  possibility  The  and  suspected.  c o r r e l a t e d with  of t h i s chromosome v a r i a b i l i t y . S_. o c c i d e n t a l i s i n t o  (1906) and H i t c h c o c k  s e p a r a t i o n of these  range i s q u i t e s a t i s f a c t o r y and,  cyto-  of S_. o c c i d e n t a l i s need  of combining S_. m a r s h a l l i i and  s p e c i e s as suggested by P i p e r  seem s a t i s f a c t o r y .  out,  than p r e v i o u s l y  The  two  as E n g l e r and  (1961) does not  taxa over most of  their  Irmscher (1916) p o i n t  t h e r e i s abundant h y b r i d i z a t i o n between s p e c i e s i n the same s e c t i o n , even between s e c t i o n s .  lumped t o g e t h e r because they would have one  o r two  I f s p e c i e s i n the genus S a x i f r a g a were o c c a s i o n a l l y i n t e r g r a d e , then the genus  extremely complex  superspecies.  S a x i f r a g a o c c i d e n t a l i s i s an extremely v a r i a b l e s p e c i e s . workers, p a r t i c u l a r l y Johnson (1923) and s p e c i e s which have subsequently dentalis .  The  Small  Many  (1905), have d e s c r i b e d  been put i n t o the synonomy o f S_. o c c i -  v a r i a b i l i t y within this species  i s so g r e a t t h a t most  23.  of the combinations o f c h a r a c t e r i s t i c s used t o d i s t i n g u i s h i t s subs p e c i f i c c a t e g o r i e s can be found throughout t h e range o f the s p e c i e s . We f e e l t h a t n o t h i n g would be gained by r e t a i n i n g any s u b s p e c i f i c c a t e g o r i e s f o r S_. o c c i d e n t a l i s as we have been unable to c o r r e l a t e d i s t r i b u t i o n w i t h any o f t h e proposed combinations o f c h a r a c t e r i s t i c s . The p o s s i b l e e x c e p t i o n i s the p o s i t i o n  o f S_. r u f i d u l a .  This  taxon has been c l a s s i f i e d as a s p e c i e s , and as a v a r i e t y and subs p e c i e s o f S_. o c c i d e n t a l i s . still  i n doubt as i n s u f f i c i e n t  The taxonomic p o s i t i o n  o f _S. r u f i d u l a i s  data have, as y e t , been o b t a i n e d .  Some l i v e c o l l e c t i o n s as w e l l as a few h e r b a r i u m specimens s o u t h e r n Oregon  appear  t o be s l i g h t l y d i f f e r e n t from S_. m a r s h a l l i i .  Some o f t h e s e specimens were l a b e l e d S_. h o w e l l i i . enough  material  taxonomic  from  We have n o t seen  o f t h i s taxon t o r e a c h any c o n c l u s i o n s r e g a r d i n g i t s  position.  The s u b p o p u l a t i o n o f S_. m a r s h a l l i i has been changed  from a v a r i e t y  ( E n g l e r and Irmscher 1916) t o a s u b s p e c i e s as our views on the use o f the rank s u b s p e c i e s i n l i e u o f v a r i e t y a r e s i m i l a r to t h o s e o f Raven (1962) as based on the views o f Du R i e t z  (1930) and P e n n e l l  (1949).  24.  LITERATURE CITED  Abrams, L. 1944. S a x i f r a g a . I n I l l u s t r a t e d f l o r a o f the P a c i f i c S t a t e s . V o l . 2. S t a n f o r d Univ. P r e s s , S t a n f o r d Univ., C a l i f , pp. 355-367. Beamish, K. I . 1961. S t u d i e s o f m e i o s i s i n the genus S a x i f r a g a o f the P a c i f i c Northwest. Can. J . B o t . 39: 567-580. Du R i e t z , G. E. 1930. The fundamental u n i t s o f b i o l o g i c a l taxonomy. Svensk. B o t . T i d s k r . 2_4: 333-428. E n g l e r , A., and E. Irmscher. 1916, 1919. S a x i f r a g a c e a e - S a x i f r a g a . In Das P f l a n z e n r e i c h , 4, 117, H. 67 (1916); H. 69 (1919). R e p r i n t e d 1958. Greene, E. L.  1888.  New o r noteworthy s p e c i e s X X I I I .  P i t t o n i a , 1_: 159.  H i t c h c o c k , C. L., and A. C r o n q u i s t . 1961. S a x i f r a g a . I n V a s c u l a r p l a n t s o f the P a c i f i c Northwest, P a r t 3. S a x i f r a g a c e a e t o E r i c a ceae. E d i t e d by C. L. H i t c h c o c k . A. C r o n q u i s t , M. Ownbey, and J . W. Thompson. U n i v e r s i t y o f Washington P r e s s , S e a t t l e , pp. 31-55. Johnson, A. M. 1923. A r e v i s i o n o f the,North American s p e c i e s of the s e c t i o n B o r a p h i l a E n g l e r o f the genus S a x i f r a g a (Tourn.) L. Minn. Stud. B i o l . S c i . _4: 1-110. Packer, J . G. 287.  1968.  IOPB chromosome number r e p o r t s .  P e n n e l l , F. W. 1949. Bot. 36: 19-22.  Taxon, 17(3) :  Toward a simple and c l e a r nomenclature.  P i p e r , C. V. 1900. New and noteworthy northwestern T o r r e y Bot. Club, 27: 394.  plants.  Am. J .  IV.  Bull.  . 1906. S a x i f r a g a . I n F l o r a o f the s t a t e o f Washington. C o n t r i b . U.S. Nat. Herb. 11: 312-317. Raven, P. H. 1962. The s y s t e m a t i c s o f Oenothera subgenus C h y l i s m i a . Univ. C a l i f . P u b l . B o t . 34: 1-122. S m a l l , J . K.  1905.  Micranthes.  N. Am. F l o r a , 2 2 ( 2 ) :  132-148.  Snow, R. 1963. A l c o h o l i c h y d r o c h l o r i c a c i d - c a r m i n e as a s t a i n f o r chromosomes i n squash p r e p a r a t i o n s . S t a i n T e c h n o l . 2_3: 9-13.  t  25.  Watson, S. 1888. XVII. Contributions Acad. Arts, 2_3: 264.  to American botany.  Proc. Am.  Whittman, W. 1965. Aceto-haematoxylin - c h l o r a l hydrate f o r chromosome s t a i n i n g . Stain Technol. 10_(3) : 161-162.  26.  TABLE I .  CHROMOSOME COUNTS Collection Number  Collector  Chromosome numbers  Location  Saxifraga occidentalis 1 Packer  2839(ALTA)  B l a k e s t o n Mt., Waterton Nat. Park, A l b e r t a  n - 10  8224  Botanie V a l l e y , near L y t t o n ,  n - 19  7901  Mt. Lihumitson, S. o f C h i l l i w a c k , B.C.  n - 19  Krause and Beamish  31-71  Yale,  n - 19  Krause  69018  B l a c k w a l l , Manning  Krause  43-70  Mt. Apex, near P e n t i c t o n ,  B.C.  n - 19  Krause  33-70  Dominion Mt., near Salir.o, B.C.  n - 19  Beamish  57501  T c h a i k a z a n R i v e r V a l l e y , B.C.  n - 19  Krause  19-71  Tweedsmuir  n - 28  Krause  16-71  B l u f f L.•,  Krause and Maze  690001  Chehalis,  Krause  42-70  Boulder Mt., N. of Ketchum, Idaho  n - ca 29  Krause  40-70  Bayhorse L . ,  n - ca 29  Krause  20-70  New Meadows,  Beamish, Vrugtroan and S p e r r i n g s 2  Beamish  B.C.  B.C. Park,B.C.  n - 19  park,B.C.  n - 28  B.C.  n - 29  Washington  near C h a l l i s ,  Idaho  n - ca 29  Idaho  Saxifraga m a r s h a l l i i ssp. m a r s h a l l i i Krause and Beamish  30-70  Stikum,  Krause and Beamish  10-70  Dora,  Krause and Beamish  35-70  Bear C r . , M i d d l e f o r k of C o q u i l l e R.,  Krause and Beamish  15-71  Scottsburg,  Krause and Beamish  36-70  Mapleton,  Oregon  n - 10  Oregon  n - 10  Oregon  Oregon  n - 10 n - 10  Oregon  n - 10  Saxifraga m a r s h a l l i i ssp. idahoensis Krause  38-70  S t . Joe R i v e r , Avery,  Krause  5-71  Myrtl-e,  Krause  6-71  Orifino,  Krause  41-70  Winchester,  Krause  32-70  Cow C.r., Hamilton, Montana  n - 10  Krause  32-71  7 mi N. of Leavenworth, Wash.  n - 10  Krause  33-71  I c i c l e C r . , S. of Leavenworth, Wash.  n - 10  Krause  39-70  Mt. T i f f a n y , Washington  n - 10  1 Packer, Taxon Beamish,  17(3):287. 1968.  Can. J . Bot. 39:567-580.  1961.  Idaho  Idaho Idaho Idaho  n - 10 n - 10 n - 10 n - 10  27  V © tt  \  •  \  ©  1  ©  1»  t- »  4*  •  •©  *| N.rUir.il  \\\<»x\  nmn II i M F i g s . 1-4.  Swui "I' l'.ni:i'l;i  —  The. black l i n e represents  2)i.  F i g . 1. Saxifraga m a r s h a l l i i ssp. mar-  s h a l l i i ; n = 10; Sitkum, Oregon; Krause and Beamish 30-70. ssp. idahoensis; n = 10; Winchester, Idaho; D.L.  Krause 41-70. . F i g . 3. _S. o c c i -  dentalis ; n = 28; Tweedsmuir Pk., B.C., Krause 39-71. ii - 29; Cbehalis, Washington; Krause & Maze 690001. type f o r Saxifraga o c c i d e n t a l i s : May  17, 1875,  CAN.  F i g . 2. S_. i n a r s h a l l i i  F i g . 4. S_. o c c i d e n t a l i s F i g . 5. Photograph of l e c t o -  specimen A.J. Macoun s.n.; Yale Mountain,  B.C.;  28.  FIGURE  6.  D i s t r i b u t i o n o f S a x i f r a g a m a r s h a l l i i spp. mar_sh,aJJ ii, S. m a r s h a l l i i ssp i d a h o e n s i s , and S. o c c i d e n t a l i s i n Washington, Idaho, Montana and s o u t h e r n B r i t i s h Columbia. ;  29.  FIGURE  7.  D i s t r i b u t i o n of S a x i f r a g a o c c i d e n t a l i s Columbia.  in  British  30. PART I I INTRODUCTION  The its  d i s t r i b u t i o n and  systematics  c l o s e l y r e l a t e d species  of S a x i f r a g a  in British  occi dentalis  Columbia are  and  p o o r l y known.  Fragmentary treatments have been i n c l u d e d i n the works of C a l d e r Taylor and  (1968), H u l t e n  Henry (1915).  complex as a whole.  (1968), H i t c h c o c k  However, no  dealt with  Johnson (1923)  do not  (1917)  the B r i t i s h  Even the monographic works of E n g l e r and  (1916) , S m a l l (1905) and this  f l o r a has  e t a l (1961), Rydberg  and  Columbia  Irmscher  adequately t r e a t  complex. In B r i t i s h Columbia the S a x i f r a g a o c c i d e n t a l i s complex c o n s i s t s  of the f o l l o w i n g s p e c i e s  of E n g l e r and  Irmscher's  section Boraphila subsection N i v a l i - v i r g i n i e n s e s :  (1916) S a x i f r a g a S_. o c c i d e n t a l i s S.  * Wats., S_. r e f l e x a Hook.; S_. rhomboidea Greene , S_. r u f i d u l a James Macoun, jS. n i v a l i s L., two  and  S_. t e n u i s  (Wahl.) H.  taxa are c l e a r l y d i s t i n g u i s h a b l e from the o t h e r  be e x p l a i n e d  l a t e r , t h e i r r e l a t i o n s h i p to each  Sm.  (Small) The  four but,  other  is s t i l l  Throughout t h i s paper, t h e r e f o r e , they are r e f e r r e d to as the nivalis-tenuis  last as  will  not c l e a r . "S_.  complex".  In an attempt to c l a r i f y  the s y s t e m a t i c  problems i n t h i s  complex, a program of f i e l d and herbarium s t u d i e s was herbarium m a t e r i a l and s o u r c e of m o r p h o l o g i c a l  l i v i n g m a t e r i a l c o l l e c t e d by data.  species  undertaken.  The  the authors were the  Q u a n t i t a t i v e measurements were taken from  Although j ^ . rhomboidea does not  extend as f a r n o r t h as  British  Columbia, i t i s i n c l u d e d h e r e because most f l o r a s g i v e  British  Columbia and A l b e r t a , as i t s n o r t h e r n  limit.  l e a v e s , scapes and i n f l o r e s c e n c e s and were a n a l y z e d  statistically.  However, they proved t o be inadequate t o d e l i m i t the t a x a under s t u d y . A c o m b i n a t i o n o f q u a l i t a t i v e c h a r a c t e r i s t i c s was found t o be the most u s e f u l c r i t e r i o n f o r t h e s e p a r a t i o n o f v a r i o u s t a x a .  These were  used t o e r e c t the d i a g n o s t i c key to the s p e c i e s . As t h i s paper d e a l s p r i m a r i l y w i t h B r i t i s h Columbia, the d i s t r i b u t i o n maps and l i s t s in all  o f r e p r e s e n t a t i v e specimens do n o t i n c l u d e ,  cases, adjacent  areas.  Herbarium m a t e r i a l was examined ALTA, Dept.  from the f o l l o w i n g  herbaria:  o f Botany, U n i v e r s i t y o f A l b e r t a , Edmonton; UAC, Dept. o f  B i o l o g y , U n i v e r s i t y o f C a l g a r y , A l b e r t a ; CAN, N a t i o n a l Museum o f Canada, Ottawa; COLO, U n i v e r s i t y o f C o l o r a d o , B o u l d e r , C o l o r a d o ; DAO, P l a n t Research I n s t i t u t e , Dept. o f A g r i c u l t u r e , Ottawa; ID, Dept. o f Botany, U n i v e r s i t y o f Idaho, Moscow, Idaho; MONTU, U n i v e r s i t y o f Montana, M i s s o u l a , Montana; ND, U n i v e r s i t y o f N o t r e Dame, N o t r e Dame, I n d i a n a ; NY, the New York B o t a n i c a l Garden, New York C i t y , New York; ORE, U n i v e r s i t y o f Oregon, Eugene, Oregon; OSC, Dept. o f Botany, Oregon S t a t e U n i v e r s i t y , C o r v a l l i s , Oregon; UC, U n i v e r s i t y o f C a l i f o r n i a , B e r k e l e y , C a l i f o r n i a ; WS, Dept.  of Plant Pathology, State College of  Washington, Pullman, Washington; and WTU, Dept. o f Botany, U n i v e r s i t y of Washington,  Seattle.  32. TAXONOMY KEY  A.  Ovary 1/2  inferior;  TO THE  f i l a m e n t s s u b u l a t e , e q u a l l i n g the s e p a l s ;  s e p a l s about as broad  as l o n g , a s c e n d i n g ;  glomerate head or a few-flowered B.  Filaments  SPECIES  inflorescence a tight  cyme.  not i n f l e x e d at t i p ; l e a v e s rhombic to s p a t u l a t e ,  t e e t h u s u a l l y o n l y on upper 1/2 u s u a l l y a few-flowered  of l e a f b l a d e ; i n f l o r e s c e n c e  cyme; pubescence of scape s p a r s e  dense, h a i r s w h i t e w i t h p u r p l e g l a n d u l a r t i p s ;  to  circumboreal,  southward i n the h i g h mountains of B r i t i s h Columbia and A l b e r t a ^ particularly  the Rocky Mountains as f a r south  Colorado BB.  Filament  1.  as  S_. n i v a l i s - t e n u i s  t i p s i n f l e x e d ; l e a v e s rhombic w i t h w e l l  complex  developed  c r e n a t e margins over most o f l e a f b l a d e ; i n f l o r e s c e n c e a t i g h t , occasionally interrupted,  glomerate head; pubescence o f scape  dense, h a i r s y e l l o w i s h w i t h y e l l o w g l a n d u l a r t i p s ; from c e n t r a l Montana south AA.  2.  Ovary s u p e r i o r to 1/3  inferior;  filaments subulate  S_. rhomboidea  to c l a v a t e ,  l o n g e r than the s e p a l s ; s e p a l s l o n g e r than b r o a d , s p r e a d i n g  to  s t r o n g l y r e f l e x e d ; i n f l o r e s c e n c e open, o c c a s i o n a l l y t i g h t - g l o m e r a t e . C.  Petals with 2 basal yellow spots; filaments strongly clavate to p e t a l o i d ; s e p a l s s t r o n g l y r e f l e x e d ; ovary s u p e r i o r ; leaves puberulent  completely  (seldom g l a b r a t e ) on both  upper  and lower s u r f a c e ; from f a r n o r t h e r n B r i t i s h Columbia i n t o Alaska,  Yukon  and Northwest T e r r i t o r i e s  3.  S_. r e f l e x a  33.  CC.  P e t a l s immaculate; f i l a m e n t s s u b u l a t e ,  l i n e a r or c l a v a t e ;  sepals spreading  to s l i g h t l y r e f l e x e d ; ovary 1/5  i n f e r i o r ; leaves  glabrous  s u r f a c e , glabrous D.  to rusty-tomentose on  to  lower  on upper s u r f a c e .  I n f l o r e s c e n c e a corymb; l e a v e s c r e n a t e margins w i t h  ovate  to rhombic w i t h  small, well-formed teeth, often  c o p i o u s l y rusty-tomentose on lower s u r f a c e s ; subulate;  Columbia R i v e r Gorge, and  Olympic  Peninsula,  4.  I n f l o r e s c e n c e v a r i o u s , seldom  leaves  to s l i g h t l y c l a v a t e ; from  northern  Columbia south to Washington, Idaho, Montana,  northeast  Oregon and  f a r south as New  1.  flat-topped;  S_. r u f i d u l a  seldom rusty-tomentose on lower s u r f a c e ;  filaments subulate British  the  the Santeam R i v e r V a l l e y ,  Oregon  various,  filaments  i n B r i t i s h Columbia r e s t r i c t e d to Vancouver  I s l a n d , a l s o found i n the  DD.  1/3  o c c a s i o n a l l y i n the Rocky Mts.  Mexico  5.  as  S_. o c c i d e n t a l i s  S a x i f r a g a n i v a l i s - t e n u i s complex The  taxonomy of the _S_. n i v a l i s - t e n u i s complex needs  biosystematic 1961,  study to s e p a r a t e  Love & Love 1951  the taxa i n v o l v e d .  & Jorgensen et a l 1958)  t e n u i s Wahl. i s s u f f i c i e n t l y d i s t i n c t from S_. s p e c i f i c rank, a l t h o u g h o t h e r s  (Hult£n 1968,  extensive  Most workers ( S a v i l e  f e e l t h a t S_. n i v a l i s  var.  n i v a l i s to warrant F l o v i c k 1940)  f e e l i t should  remain a subcategory of S_. n i v a l i s .  Over much of t h e i r range they can  adequately s e p a r a t e d  s i m i l a r to those proposed by  using  criteria  be  Savile  34. (1961):  number of f l o w e r s ; shape o f i n f l o r e s c e n c e ; c o l o u r o f p e t a l s ,  c a l y x and scape; type of pubescence;  and e s p e c i a l l y shape o f l e a v e s .  the Yukon, however, these d i s t i n c t i o n s seem to break down  and  In  the p l a n t s  appear t o i n t e r g r a d e . Cytotaxonomists i n Europe (Hedberg  1967,  (see Love & Love 1961)  and North  Johnson & Packer 1968) have r e p o r t e d S_. n i v a l i s  America to have  e i t h e r 58, 60 o r about 60 chromosomes and J3. n i v a l i s v a r . t e n u i s to have 20 chromosomes.  It i s d i f f i c u l t  t o c o r r e l a t e the d i f f e r e n t counts w i t h  m o r p h o l o g i c a l f e a t u r e s as we have not o b t a i n e d enough counts and the p r e v i o u s authors have not g i v e n the c r i t e r i a they used f o r i d e n t i f i c a tion.  We have o b t a i n e d counts o f 10, 20 and 30 p a i r s of chromosomes f o r  the m a t e r i a l c o l l e c t e d i n the Yukon ( T a b l e 1, F i g . 1A & I E ) . bility  i n the chromosome numbers w i t h i n t h i s complex may  than was  varia-  be more p r e v a l e n t  previously suspected.  The S_. n i v a l i s - t e n u i s complex i s c i r c u m b o r e a l . Rocky Mountains southward  The  o f B r i t i s h Columbia  as f a r as C o l o r a d o .  Columbia and southward  I t i s common i n the  and A l b e r t a y ( F i g . 2) and  The s c a t t e r e d  occasionally  distribution in British  i s p r o b a b l y due t o c o l l e c t i n g anomalies.  B r i t i s h Columbia i t i s s y m p a t r i c w i t h S_^_  occidentalis  i n high  In mountains  but i t can be r e a d i l y s e p a r a t e d from S_. o c c i d e n t a l i s as shown i n the key to s p e c i e s . Some r e p r e s e n t a t i v e  specimens  o f the S_.  niv alis-tenuis  complex i n  A B r i t i s h Columbia and A l b e r t a f o l l o w :  BRITISH COLUMBIA:  summit of O t t e r t a i l P a s s , Rocky Mt. Park, Macoun s.n. A d d i t i o n a l c i t a t i o n s f o r a l l species i n T a b l e 1.  appear  On rocks below  (CAN); A l i c e  Mt.,  with, chromosome counts  35.  n.e. o f T u r n a g a i n L., C a s s i a r D i s t . A.F. S z c z a w i n s k i s.n. (DAO); Murray Range a t n o r t h end o f A z o u z e t t a L., H a r t Highway, J.A. C a l d e r , D.B.O. S a v i l e and J.M. Ferguson 14075 (DAO); Red Rose Tungsten Mine about 9 m i l e s e. of Skeena C r o s s i n g , J.A. C a l d e r , D.B.O. S a v i l e and J.M. F e r guson 15182 and 13369 (DAO); M i l e 75, Haines Rd., T.M.C. T a y l o r , A. S z c z a w i n s k i and M. B e l l 824 (DAO, UBC); M i l e 60, Haines Rd., T.M.C. T a y l o r , A. S z c z a w i n s k i and M. B e l l 1204 (UBC, DAO); M i l e 68, Haines Rd., T.M.C. T a y l o r , A. S z c z a w i n s k i and M. B e l l 896 (DAO, UBC); M i l e 61, Haines Rd., D. K r a u s e , K.I. Beamish and J . L u i t j e n s 681807 (UBC).  ALBERTA:  Snow  L. Pass about 30 m i l e s n. o f B a n f f , J.A. C a l d e r 23996 (DAO); B a n f f Nat. Pk., a l p i n e , wet c l i f f s  above Lake Agnes, A. Harmon 8 (DAO);  Rocky s l o p e , above timber l i n e , J a s p e r Nat. Pk., E.H. Moss 2736a top o f rocky r i d g e about 5300 f t .  (ALTA);  above Cadomin, E.H. Moss 10339 (ALTA).  The f o l l o w i n g sheets c a r r y specimens o f both S_. o c c i d e n t a l i s and the j>. n i v a l i s - t e n u i s C a l d e r 37210 (DAO).  complex.  ALBERTA:  BRITISH COLUMBIA:  Mt. E d i t h C a v e l l , J.A.  Mt. T h o r n h i l l ,  Terrace,  J.A. C a l d e r , D.B.O. S a v i l e and J.M. Ferguson 14888 (DAO); S y l v a n Mine, Hudson Bay Mt., about 9 m i l e s w. o f S m i t h e r s , J.A. C a l d e r , D.B.O. S a v i l e and J.M. Ferguson 14612 (DAO).  2.  The l a s t p o s s i b l y has h y b r i d s .  S a x i f r a g a rhomboidea Greene Most f l o r a s  British  f o r the P a c i f i c Northwest i n c o r r e c t l y g i v e s o u t h e r n  Columbia and A l b e r t a as t h e n o r t h e r n  l i m i t s o f S. rhomboidea  36. distribution.  T h i s i s p r o b a b l y due  some _S. o c c i d e n t a l i s p l a n t s w i t h pubescent  scapes  to the s u p e r f i c i a l resemblance of  t i g h t glomerate  to S_. rhomboidea.  heads and  As shown i n the key  densely  these two  can be s e p a r a t e d r e a d i l y u s i n g f i l a m e n t shape, i n f e r i o r i t y of shape, and  position  of s e p a l s , and  species  ovary,  c o l o u r of the g l a n d u l a r h a i r s on  the  scape. S a x i f r a g a rhomboidea extends (Fig.  3).  o n l y as f a r n o r t h as c e n t r a l Montana  In s o u t h e r n Montana and Wyoming i t i s o c c a s i o n a l l y  w i t h S^. o c c i d e n t a l i s .  The  f o l l o w i n g herbarium  b o t h _S. rhomboidea and  S_. o c c i d e n t a l i s on them:  Wyoming, L i l l i a n W i n i e c k i 67 L.A.  F i t c h s.n.  (WS);  Mts.  sympatric  sheets were found w i t h H e l l Roaring  trail,  at 7000 f t , S h e r i d a n , Montana,  (NY)f Columbia F a l l s , Montana, R.S.  W i l l i a m s.n.  (NY).  The n o r t h e r n d i s t r i b u t i o n of S_. rhomboidea i s shown by the f o l l o w i n g r e p r e s e n t a t i v e specimens from Montana: n.w.  of Red Lodge, C H .  Draper s.n.  G a l l a t i n Canyon near Squaw Cr., W.E. Bozeman B r i d g e Canyon, J.W. s l o p e of King's H i l l , 12337a (WTU, cock 16131  WS); (WTU,  WS,  CARBON Co.:  (UC); GALLATIN Co.: Booth 1514  B l a n k i n s h i p s.n.  Little  Yoho Pk.,  MONTANA:  B e l t Mts.,  (WTU,  4 miles  Bozeman,  WS,  MONTU);  (UC); MEAGHER Co.:  CL.  L i t t l e B e l t Mts.,  H i t c h c o c k and  C.V.  south s i d e , C L .  H i t c h c o c k and C.V.. M u h l i c k  (WTU); s. s i d e of B l a c k B u t t e , G r a v e l l y Range, C L .  Cooke Guard S t a t i o n , 3 m i l e s e. of Cooke C i t y , J . C  1161  WTU,  272  (WS);  Hitch-  W a l l r o c k B a s i n , n.w.  STILLWATER Co.:  Absaroka Nat. F o r . , C L .  of W i l l s a l l , W.N.  n. s l o p e of Mt. H i t c h c o c k and C.V.  Haystack,  12500  H i t c h c o c k 16875 (WTU);  PARK Co.:  WS);  Muhlick  UC); MADISON Co.: . 1 m i l e s. of C r o c k e t t L.  ( D e v i l ' s L.) , G r a v e l l y Range, C L .  (COL,  s.w.  Witt  Suksdorf  head o f Boulder  M u h l i c k 13424a (WTU).  Cr.,  37.  3.  S a x i f r a g a r e f l e x a Hook. S a x i f r a g a r e f l e x a extends southward o n l y i n t o t h e most n o r t h e r l y  p o r t i o n o f B r i t i s h Columbia cut.  ( F i g . 2 ) . I t s taxonomy i s r e l a t i v e l y  clear-  I t can be i d e n t i f i e d by i t s r e f l e x e d s e p a l s , y e l l o w s p o t s on t h e  p e t a l s , and i t s p u b e r u l e n t l e a v e s . S a x i f r a g a r e f l e x a appears t o be most c l o s e l y r e l a t e d t o S^. mars h a l l i i Greene growing i n Oregon,  C a l i f o r n i a , Washington  and Idaho.  have y e l l o w s p o t s on t h e p e t a l s , r e f l e x e d s e p a l s , s u p e r i o r  Both  ovaries,  open i n f l o r e s c e n c e s , and 10 p a i r s o f chromosomes (Krause and Beamish, i n p r e s s ) , w i t h t h e e x c e p t i o n o f one p o p u l a t i o n Creek, Yukon. and  o f S_. r e f l e x a from  Quill  At the Q u i l l Creek s i t e counts were found o f n = 10, n = 20,  (probably i n a h y b r i d ) n = c a . 30 (Table 1 ) . R e p r e s e n t a t i v e specimens  from B r i t i s h Columbia:  F l o r a of Cassiar,  T.M.C. T a y l o r , A. S z c z a w i n s k i and M. B e l l 512, 623, and 498 (UBC); M i l e 87, Haines Rd.,, T.M.C. T a y l o r , A. S z c z a w i n s k i and M. B e l l 932 (UBC); M i l e 81, Haines Rd., T.M.C. T a y l o r , A. S z c z a w i n s k i and M. B e l l 1266  (UBC); B.C. - Yukon boundary,  (CAN); Stonehouse  Cr., M i l e 61, Haines Rd., K.I. Beamish,  and J . L u i t j e n s 681801  4.  59°58*W, M.P. and R.T. P o r s i l d 198  Saxifraga r u f i d u l a  D.L. Krause  (UBC).  (Small) James Macoun*  S a x i f r a g a r u f i d u l a i s found on Vancouver I s l a n d , the Olympic P e n i n s u l a , the Columbia R i v e r Gorge and i n t h e Santeam R i v e r V a l l e y o f Oregon In B r i t i s h Columbia i t i s c o n f i n e d t o Vancouver I s l a n d . distinct  (Fig. 2).  I t i s geographically  from S_. o c c i d e n t a l i s as the l a t t e r i s found on t h e mainland.  The name S a x i f r a g a r u f i d u l a to John Macoun by Beamish  (Small) Macoun was i n c o r r e c t l y  (1967).  attributed  38. In  the Columbia R i v e r Gorge S_. r u f i d u l a appears to i n t e r g r a d e w i t h  S_. o c c i d e n t a l i s and perhaps w i t h to  Much work remains  be done to determine the e x t e n t of the i n t e r g r ^ a d a t i o n . Chromosome counts  o b t a i n e d f o r S_. ruf i d u l a are g i v e n i n T a b l e  On Vancouver I s l a n d i t was but d e t a i l e d study  In the  found  d i f f e r e n c e between the  10-  the 2 9 - p a i r e d p o p u l a t i o n ( F i g . ID).  Columbia R i v e r Gorge, counts  n = 10, n = 19  1.  to have 10 and 29 p a i r s o f chromosomes  r e v e a l e d no m o r p h o l o g i c a l  p a i r e d p o p u l a t i o n s and  of  S_. m a r h s a l l i i .  ( F i g . I B ) , n = 28  f o r S_. ruf i d u l a were o b t a i n e d  ( F i g . IC & F) and n = c a . 28.  The  p l a n t s i n the Gorge are m o r p h o l o g i c a l l y very v a r i a b l e but none of the morphological  d i f f e r e n c e s c o u l d be  c o r r e l a t e d w i t h chromosome numbers.  R e p r e s e n t a t i v e specimens f o r B r i t i s h n.w.  of V i c t o r i a , L.C.  Sherk 302  Columbia f o l l o w :  Mt.  Helmcken,  (DAO); a l o n g s. s i d e of Cameron L.  between P o r t A l b e r n i and P a r k s v i l l e , J.A.  C a l d e r , J.A.  Parmelee,  and  R.L.  T a y l o r 16414  (DAO); near C l i f f s i d e on e a s t s i d e of Shawnigan L.,  J.A.  C a l d e r , J.A.  Parmelee, and R.L.  R.  about 3 m i l e s n. of M i l n e ' s Landing, w.  28420 (DAO); C a s t l e c r a g Mt., J.R. Mt.  T a y l o r 16245 (DAO); along Sooke  Anderson 644^  (DAO); Mt.  Arrowsmith, J.A.  (DAO); Observatory  J.E. U n d e r h i l l 406  Parmelee and R.L.  n. of V i c t o r i a , J.A.  28660 (DAO); above Boston Lk., below Mt.  M.T.  MacKay 429021 (DAO).  MacKay  C a l d e r and  Calder  (DAO);  19431  C a l d e r and M.T.  J.A.  Arm,  Henson s.n.  Taylor  Becher, J.A.  MacKay 32557 (DAO); Nanoose H i l l , Nanoose Bay,  Calder  (DAO); Saanich  Douglas, V i c t o r i a , P.P.  C a l d e r , J.A.  Hill,  of V i c t o r i a , J.A.  and  M.T.  39. 5.  S a x i f r a g a o c c i d e n t a l i s S. Wats. Saxifraga occidentalis  can be s a t i s f a c t o r i l y s e p a r a t e d from  other species of s e c t i o n Boraphila subsection N i v a l i - v i r g i n i e n s e s the c h a r a c t e r s i n the k e y , a l t h o u g h as mentioned  using  e a r l i e r , some s p e c i -  mens w i t h t i g h t glomerate heads have been m i s t a k e n l y i d e n t i f i e d as S_. rhomboidea  (See under S_. rhomboidea) .  The d i s t r i b u t i o n o f S_. o c c i -  d e n t a l i s i s g i v e n i n F i g u r e s 2 and 3. R e p r e s e n t a t i v e specimens  o f S_. o c c i d e n t a l i s w i t h t i g h t  heads, many o f which have been i d e n t i f i e d as S_. rhomboidea, BRITISH COLUMBIA:  follow:  Q u i n i s c o e L., A s h n o l a d i s t r i c t , M.E. B a r r PB69  (UBC); Mt. McLean, near L i l l o o e t , B.C., J.M. Macoun s.n. (CAN); o f M c G i l l i v r a y C r . , Cascade Range, J.M. Macoun s.n. C h i l l i w a c k V a l l e y , W. Spreadbourough Park, L. Holm 428 (UBC). Park, L.H. Harvey  MONTANA:  s.n.  head  (CAN); S l e s s e Mt.,  (CAN); B l a c k w a l l Pk., Manning  R i s i n g Sun Meadows, G l a c i e r Nat.  4773 (MONTU); Logan P a s s , G l a c i e r Nat. Park, G.N. Jones  5525 (UC, WTU); S i y e h P a s s , G l a c i e r LAKE Co.:  glomerate  Nat. Park, S. Bamberg 288 (COL);  s.e. o f McDonald L. , Mission, Range,  (ID, COL, UC); MISSOULA Co.:  C.L. H i t c h c o c k 1830,0  1 m i l e w. o f Gordon Pass, F l a t h e a d Range,  C.L. H i t c h c o c k 18421 (UC, WS) .  ALBERTA:  Shoulder o f W i l c o x Mt.,,  J a s p e r Nat. Park, E.H. Moss 4965 (ALTA); above Sunwapta P a s s , J a s p e r Nat. Park, E.H. Moss 4887 (ALTA); near B l a k i s t o n Brook, Waterton L. Park, E.H.  Moss 10174 (ALTA, CAN); 5 m i l e s w. o f P i n c h e r Cr., S . S . Servey,  1193  (ALTA); Upper S p r i n g C r . , between L i v i n g s t o n and P o r c u p i n e Range  S t n . , c a . 30 m i l e s n. o f L i n d b r e c k , R.G.H. Cormack ,52 (UAC); B a n f f v i c i n i t y , E.S. Dowding s..n. (ALTA); Raspberry Ridge., Kananaskis  Forest  40. P r e s e r v e , B. de V r i e s 2348-64 (DAO, UAC); Opal H i l l s , n.e. o f Rainbow Lodge, M a l i g n e  L., L. J e n k i n s  B a n f f , J.A. C a l d e r 23976 n.w.  7903 (DAO); Snow Ck. Pass,  (DAO); s.w.  of d r i l l s i t e ,  20 m i l e s n. of  B i g h o r n Mts., 4 m i l e s  of South Ram R., j e t . w i t h Nordegg - Cochran F o r e s t r y Rd., C.D.  Bird  7538 (UAC). Not  a l l S_. o c c i d e n t a l i s specimens from n o r t h e r n Montana have t i g h t  glomerate heads.  The f o l l o w i n g  have open i n f l o r e s c e n c e s :  m i l e s n. o f S t . Mary, D. Lynch 6284 (WS); L. S t . Mary, L.W. BEZVERHEAD Co.: 14925  GLACIER  Top of O d e l l Pk., P i o n e e r Range, C.L. H i t c h c o c k and C.V.  (WS); MISSOULA Co.:  h m i l e w. of Upper H o l l a n d L., F l a t h e a d  18422 (WS); 5 m i l e s above Bonner, C.L. H i t c h -  cock and C.V. M u h l i c k  11425  (WS, WTU);  DEERLODGE Co.:  Storm L., Anaconda Mts., C.L. H i t c h c o c k and C.V. M u h l i c k LAKE Co.:  C.L. H i t c h c o c k  Dry s l o p e above 14820  (WS,  High above and s.e. of McDonald L., M i s s i o n Range,  18300 (WTU,  WS).  2.6  S a y l o c 43 (UC);  Range, C.L. H i t c h c o c k  WTU);  Co.:  Muhlick  41. LITERATURE CITED  Beamish, K. I . 1967. A P a c i f i c Coast s a x i f r a g e w i t h 10 p a i r s of chromosomes: M e i o s i s , development of the female gametophyte, and seed p r o d u c t i o n . Can. J . Bot. 45: 1797-1801. C a l d e r , James A. and T a y l o r , Roy L. 1968. S a x i f r a g a . I n , F l o r a of the Queen C h a r l o t t e I s l a n d s . Systematics of t h e . V a s c u l a r P l a n t s . Res. Branch, Canada Dept. A g r i c , Monogr. 4, P a r t 1. Queen's P r i n t e r , Ottawa, p. 380-389. E n g l e r , A. and Irmscher, E. 1916. R e p r i n t 1958. S a x i f r a g a c e a e S a x i f r a g a , S e c t . 1 B o r a p h i l a E n g l . T n , Das P f l a n z e n r e i c h 4, 117, H67: 5-89. F l o v i c k , K. 1940. Chromosome numbers and p o l y p l o i d y w i t h i n the f l o r a of S p i t z b e r g e n . H e r e d i t a s 2J>: 430-440. Hedberg, 0. 1967. Chromosome numbers o f v a s c u l a r p l a n t s from a r c t i c and s u b - a r c t i c North America. Ark. f . Bot. n.s. 6_: 309-326. Henry, J . K. 1915. S a x i f r a g a . I n , F l o r a of Southern B r i t i s h and Vancouver I s l a n d . Gage, T o r o n t o , p. 164-168.  Columbia  H i t c h c o c k , C. L., C r o n q u i s t , A., Ownbey, M., and Thompson, J . W. 1961. Saxifraga. I n , V a s c u l a r P l a n t s o f the P a c i f i c Northwest. P a r t 3. U n i v e r s i t y of Washington P r e s s , S e a t t l e , p. 31-55. H u l t e n , E. 1968. Territories.  Saxifraga. I n , F l o r a o f A l a s k a and N e i g h b o r i n g Stanford University Press, Stanford, p. 563-583.  Johnson, A. M. 1923. A r e v i s i o n o f the North American s p e c i e s o f the s e c t i o n B o r a p h i l a E n g l e r of the genus S a x i f r a g a (Tourn.) L. Minn. Stud. B i o l . S c i . 4_: 1-110. Johnson, A. W. and P a c k e r , J . G. 1968. Chromosome numbers i n the f l o r a o f Ogotoruk Creek, N.W. A l a s k a . Bot. Not. 121: 403-456. J o r g e n s e n , C. A., Sorensen, T. H., and Westergaard, M. 1958. The f l o w e r i n g p l a n t s of Greenland. A t a x o n o m i c a l and c y t o l o g i c a l s u r v e y . • Danske Vidensk. S e l s k . B i o l . S k r . 9 ( 4 ) : 1-172. Krause, D. L. and Beamish, K. I . Taxonomy o f S a x i f r a g a o c c i d e n t a l i s S. Wats, and S. m a r s h a l l i i Greene. Can. J . B o t . I n p r e s s . Lbve, A. and Love, D. 1951. S t u d i e s on the o r i g i n o f the I c e l a n d i c flora. I I . S a x i f ragaceae. Svensk Bot. T i d s k r . 45_: 368-399.  42.  LbVe, A. and Love, D. 1961. Chromosome numbers o f c e n t r a l and n o r t h west European P l a n t s p e c i e s . Opera B o t a h i c a 5_: 1-581. Rydberg, P. A. 1917. Saxifragaceae. and A d j a c e n t P l a i n s . H a f n e r , New  I n , F l o r a o f the Rocky York, p.* 376-391.  S a v i l e , D. B. 0. 1961. The botany of n o r t h w e s t e r n Queen Islands. Can. J . Bot. 39: 909-942. S m a l l , J . K. and Rydberg, P. A. can F l o r a 22(2) : 81-158.  1905.  Saxifragaceae.  Mountains  Elizabeth  I n , North Ameri-  TABLE  1.  C H R O M O S O M E COUNTS F R O M BRITISH C O L U M B I A AND A D J A C E N T A R E A S  Collector  Collection Number  Chromosome numbers  Location  S. n i v a l i s - t e n i u s c o m p l e x  Krause, Beamish and L u i t j e n s  682000  M i l e 101, L a p i e L.,  Krause, B e a m i s h and L u i t j e n s  681807  M i l e 61, H a i n e s Rd., B.C.  2n = 60  Krause, Beamish and L u i t j e n s  681398  S.E. of L a p i e L.,  n = 10  Krause  68  M t . S.S.W. of A l t u r u s L., Idaho  *n = 20  Yukon  Yukon  n = 10  S. r c f l e x a Krause, Beamish and L u i t j e n s  681768  Q u i l l C r . w. of K l u a n e L.,  Yukon  n = 10  Krause, Beamish and L u i t j e n s  681768  Q u i l l C r . w. of K l u a n e L.,  Yukon  n = 20  Krause, Beamish arid L u i t j e n s  681768  Q u i l l C r . w. of K l u a n e L.,  Yukon  2n c a . 30  Krause, Beamish and L u i t j e n s  682001  M i d n i g h t Dome, D a w s o n C i t y ,  Krause, Beamish and L u i t j e n s  681436  N.E. of F i s h L.,  22 - 71  Sooke, B.C.  n = 10  21 - 71  Mt. F i n l a y s o n , B.C.  n = 10  1 - 71  Shawnigan L.,  n = 10  1 - 70  C a m p b e l l R i v e r , B.C.  *n = 29  1 rni w. of Cook at D o g Cr., W a s h i n g t o n  *n = 19  B e a v e r C r . F a l l 10 m i e. of Clatskanie, Oregon  "n = 19  69000 5 7 - 71  Yukon  Whitehorse, Yukon  B.C.  n = 10  n = 10  K r a u s e and M a z e  690006  V i e n t o State P a r k , 8 m i w. of H o o d R., O r e g o n  n = 19  K r a u s e and M a z e  690007  W. of A i n s w o r t h State P a r k , n e a r O e n e o n t a Gorge, Oregon  n = 10  E l k Rock, W i l l a m e t t e R., O r e g o n  n = 10  Krause  9 - 71  Krause  11 - 71  B i g C l i f f Dam, n e a r D e t r o i t , O r e g o n  n = 10  Krause  12-71  5 m i e. of S. S a n t i a m R., L i n n Co., O r e g o n  n - 10  Krause.  4 - 71  Mosier, Oregon  n = c a . 28  Krause  2 - 71  1 mi w. of C l a r k - S k a m a n i a Co. l i n e , C o l u m b i a R. Gorge, Washington  n = 10 n » 28  K r a u s e and B e a m i s h  27 - 70  The D a l l e s , O r e g o n  K r a u s e and B e a m i s h  24 - 70  Troutdale, Oregon  K r a u s e and B e a m i s h  28 - 70  White Salmon, Washington  Gh r oino some n u m b e r s i n bold fact! indicate new count!! fur the taxon.  ' n - 10 n - c a . 28  44.  Figure 1.  A-D.  The black l i n e represents  2u.  F i g . 1A.  Saxifraga n i v a l i s - t e n u i s complex, n = 20, M i l e  101,  Lapie Ls., Yukon, Krause, Beamish and L u i t j e n s 682000; F i g . IB.  S. r u f i d u l a , n = 19, 1 mi w. of  Cook at Dog Cr., Washington, Krause & Maze 690005; F i g . IC.  J3.  r u f i d u l a , n = 28, The D a l l e s ,  Krause & Beamish 27-70; F i g . ID. Campbell River, B.C.,  Oregon,  S. r u f i d u l a , n = 29,  Krause 1-70; F i g . IE.  p r e t i v e drawing of F i g . 1A; F i g . IF.  Inter-  Interpretive  drawing of F i g . IC.  0  • O V  »  ©  .  ©  © •. #  occidentalis, j5. reflexa and j>. rufidula i n British Columbia and from f i e l d work and  some adjacent areas, derived herbarium specimens.  \  I I."  I f I  •  Saxifraga  •  S. occidentalis  rhomboid ea  I  i  Figure 3.  D i s t r i b u t i o n of S a x i f r a g a rhomboidea and S_. o c c i d e n t a l i s i n B r i t i s h Columbia and adjacent areas, derived from f i e l d work and herbarium specimens.  

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