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Variability in Saxifraga ferruginea Graham (Saxifragaceae) Randhawa , Ajit S. 1969

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VARIABILITY IN SAXIFRAGA FERRUGINEA GRAHAM (SAXIFRAGACEAE) by AJIT  S.  RANDHAWA  B. Sc. ( A g r i . ) , Panjab U n i v e r s i t y , I n d i a , 1952 M. Sc. ( A g r i . ) , Panjab U n i v e r s i t y ,  I n d i a , 1955  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR  OF  PHILOSOPHY  I n the Department of BOTANY  We a c c e p t t h i s t h e s i s as conforming to t h e required  standard  THE UNIVERSITY OF BRITISH COLUMBIA A p r i l , 1969  In p r e s e n t i n g  this  thesis  an a d v a n c e d d e g r e e a t the I  for  this  written  for  for extensive  may be g r a n t e d b y  representatives. thesis  It  is  f i n a n c i a l gain  of  The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, Canada  Columbia  shall  the requirements  Columbia,  I agree  r e f e r e n c e and copying  of  this  that copying or  for  that  Study. thesis  t h e Head o f my D e p a r t m e n t  understood  permission.  Department  British  it freely available for  that permission  s c h o l a r l y purposes  by h i s of  the U n i v e r s i t y of  L i b r a r y s h a l l make  f u r t h e r agree  in p a r t i a l f u l f i l m e n t of  or  publication  n o t be a l l o w e d w i t h o u t my  ABSTRACT  S a x i f r a g a f e r r u g i n e a Graham complex i s w i d e l y in  the P a c i f i c northwest and  Different ways.  exhibits considerable v a r i a b i l i t y .  taxonomists have r e c o g n i z e d  the v a r i a b i l i t y  T h e i r d e c i s i o n s were based on m o r p h o l o g i c a l  herbarium specimens and  p l a n t s i n the  1.  in different  s t u d i e s of  field.  The main o b j e c t i v e of the p r e s e n t fold:  distributed  i n v e s t i g a t i o n was  two-  to determine the extent of v a r i a b i l i t y w i t h i n the  complex.  2.  species  to f i n d out whether or not t h e r e i s c o r r e l a t i o n between  d i f f e r e n t c h a r a c t e r i s t i c s to d e l i m i t s u b - s p e c i f i c taxa. study from which i n f o r m a t i o n was  d e r i v e d were:  F i e l d s of  c y t o l o g y , morphology  ( l i v i n g and h e r b a r i u m specimens), embryology, anatomy, chromatography, and c r o s s a b i l i t y i n v o l v i n g d i f f e r e n t p o p u l a t i o n s of the s p e c i e s range grown i n the greenhouse a t the U n i v e r s i t y of B r i t i s h Laboratory  and  field  Columbia.  i n v e s t i g a t i o n s were conducted d u r i n g 1965-1968.  C y t o l o g i c a l o b s e r v a t i o n s and p a i r e d d i p l o i d as w e l l as 19-paired  chromosome counts r e v e a l e d p o l y p l o i d (aneuploid)  populations.  P o l y p l o i d s were i n g e n e r a l more i r r e g u l a r i n m e i o s i s  than the  Embryological  of  s t u d i e s i n d i c a t e d an unusual  behaviour  10-  diploids.  integumentary  and n u c e l l a r l a y e r s , not r e p o r t e d e a r l i e r i n the genus S a x i f r a g a . T h i s phenomenon was  common to a l l p o p u l a t i o n s  Morphological obtained  examined.  s t u d i e s were c a r r i e d out on herbarium specimens  f o r the e n t i r e range. L i v e p l a n t s r e p r e s e n t i n g d i f f e r e n t popu-  l a t i o n s were examined f o r seed,  l e a f , f l o w e r and  pollen characters.  c h a r a c t e r s used i n the keys of p r e v i o u s taxonomists were c a r e f u l l y examined.  The  s p e c i e s was  found  to propagate s e x u a l l y by  seed,  and  The  iii v e g e t a t i v e l y by p l a n t l e t s developed crowns a r i s i n g from the rhizome. presence  i n the i n f l o r e s c e n c e and by  new  Anatomy o f the rhizome showed the  of endodermis which i s not p r e s e n t  i n the peduncle.  a r e common i n l e a v e s , s e p a l s and ovary w a l l s .  In these and  Druses other  a n a t o m i c a l r e s p e c t s the p o p u l a t i o n s were a l i k e . The chromatographic  study gave a s i m i l a r p a t t e r n of p h e n o l i c  compounds f o r a l l the p o p u l a t i o n s examined.  S. f e r r u g i n e a l a c k s  2,hydroxyphenyl a c e t i c a c i d , a compound known to be p r e s e n t o n l y i n one  genus, A s t i l b e , o f The  p o l y p l o i d p o p u l a t i o n s were i n t e r f e r t i l e as were the  but w i t h a reduced  seed  served as c o n t r o l s . diploid  Saxifragaceae.  s e t as compared to i n t r a p o p u l a t i o n c r o s s e s which  The r e c i p r o c a l c r o s s e s between p o l y p l o i d  populations f a i l e d  between p o l y p l o i d and  to set seed.  The b l o c k to gene f l o w  The  d i s t r i b u t i o n of d i p l o i d s and  f e r r u g i n e a appears to be c o r r e l a t e d  G l a c i a t i o n i n the C o r d i l l e r a n G l a c i a l Complex and p o s t - g l a c i a l times. Although  and  d i p l o i d p o p u l a t i o n s suggests a g e n e t i c b a r r i e r  f a v o u r i n g d i f f e r e n t i a t i o n i n time. p l o i d s i n S.  diploids  poly-  to P l e i s t o c e n e plant migration i n  T h i s s i t u a t i o n has been d i s c u s s e d .  a tremendous m o r p h o l o g i c a l  p l a s t i c i t y has been  observed  i n the s p e c i e s complex, the o n l y c o n s i s t e n t d i f f e r e n c e among p o p u l a t i o n s i s the two  chromosomal r a c e s .  The other c h a r a c t e r i s t i c s do not  show any  c o r r e l a t i o n w i t h chromosome number so t h a t the d i p l o i d s and p o l y p l o i d s are m o r p h o l o g i c a l l y i n d i s t i n g u i s h a b l e . d i f f e r e n t i a t i o n may  occur  The c o n c l u s i o n i s t h a t  i n the f u t u r e , e s t a b l i s h m e n t o f  c a t e g o r i e s i s not warranted  a t the p r e s e n t  time.  although  subspecific  iv TABLE; OF CONTENTS Page ABSTRACT  i i  LIST OF TABLES  v i i  LIST OF FIGURES  viii  ACKNOWLEDGEMENT S  x  INTRODUCTION  1  MATERIAL  3  METHODS, OBSERVATIONS, AND RELEVANT LITERATURE  6  CYTOLOGICAL STUDIES  7  Methods  7  O b s e r v a t i o n s and r e l e v a n t l i t e r a t u r e  7  MORPHOLOGY AND THE L I F E CYCLE  11  Methods  11  O b s e r v a t i o n s and r e l e v a n t l i t e r a t u r e  13  Seeds  13  Seed g e r m i n a t i o n  14  Adventitious roots  15  Leaves  15  Pubescence , ,  15  Stomata  16  Flower  16  Plantlets  17  Rhizome The L i f e C y c l e  .18 19  V  Page EMBRYOLOGY  • • • • • . . . . . . . . . . . . . . .  21  Methods  21  O b s e r v a t i o n s and r e l e v a n t l i t e r a t u r e  22  Ovule  22  Megasporogenesis  23  Megagametogenesis  24  Fertilization  25  Endosperm  25  Embryogenesis  26  ANATOMICAL STUDIES .  28  Methods  28  O b s e r v a t i o n s and r e l e v a n t l i t e r a t u r e  29  Root  29  Peduncle  30  Rhizome  31  Leaf  31  Flower  32  Shoot Apex and I n f l o r e s c e n c e  33  CHROMATOGRAPHIC STUDIES  34  Methods  34  O b s e r v a t i o n s and r e l e v a n t l i t e r a t u r e  35  CROSSABILITY  STUDIES  36  Methods  36  Observations  36  GENERAL DISCUSSION  38  vi Page SUMMARY  •  LITERATURE CITED APPENDICES FIGURES  I to V  .V . . . . .  5  8  60 66 72  vii LIST OF TABLES Table I.  Page C o l l e c t i o n s i t e s , voucher specimens and chromosome numbers i n S a x i f r a g a f e r r u g i n e a Graham  II.  Meiosis  i n microsporocytes  3  o f S. f e r r u g i n e a Graham  ( d i p l o i d and p o l y p l o i d ) III.  Pollen v i a b i l i t y  9  i n d i p l o i d and p o l y p l o i d  populations  of S. f e r r u g i n e a Graham IV.  E f f e c t of d i f f e r e n t the g e r m i n a t i o n  V.  VI.  temperatures and p h o t o p e r i o d s on  o f seeds o f S_. f e r r u g i n e a Graham . . .  V a r i a t i o n i n morphological populations  10  characters of d i f f e r e n t  o f S. f e r r u g i n e a Graham  20  UV f l u o r e s c e n c e and c o l o u r r e a c t i o n s o f the p h e n o l i c compounds found i n l e a v e s o f S_. f e r r u g i n e a Graham  VII.  14  Interpopulation crosses  in S. f e r r u g i n e a Graham . . . .  35 37  viii LIST OF FIGURES Figure 1.  Page D i s t r i b u t i o n map  o f S a x i f r a g a f e r r u g i n e a Graham  based on h e r b a r i u m specimens examined  72  2.  M i c r o s p o r o g e n e s i s i n S.  73  3.  G e r m i n a t i o n o f seed and the development o f s e e d l i n g s i n S.  4.  ferruginea  74  V a r i a t i o n i n b a s a l l e a v e s o f jS. f e r r u g i n e a from C a l i f o r n i a , Montana,  5.  ferruginea  Idaho, and A l a s k a  V a r i a t i o n i n b a s a l l e a v e s o f S.  75  f e r r u g i n e a from  B r i t i s h Columbia 6.  76  V a r i a t i o n i n b a s a l l e a v e s o f S. f e r r u g i n e a from Washington and Oregon  7.  77  V a r i a t i o n i n the b r a c t e a l l e a v e s o f jS. f e r r u g i n e a from Oregon, Washington, Idaho, Montana, Alaska,  8.  9.  California,  and B r i t i s h Columbia  78  V a r i a t i o n i n the s i z e and shape o f p e t a l s i n the f l o w e r s o f £>. f e r r u g i n e a ..  79  Vegetative  80  reproduction  i n S>. f e r r u g i n e a  10.  R e p r o d u c t i v e b i o l o g y o f S.  11.  Megasporo-  12.  Embryogenesis i n S.  ferruginea  and megagametogenesis ferruginea  i n j>. f e r r u g i n e a  81 82 83  ix Figure  Page  13.  Anatomy o f J3. f e r r u g i n e a  14.  Anatomy o f the rhizome o f S.  15.  Vasculature  o f the f l o w e r o f j>. f e r r u g i n e a  86  16.  Anatomy o f the shoot apex o f j3. f e r r u g i n e a  87  17.  The t w o - d i r e c t i o n a l chromatogram showing spots o f phenolic spray  18.  ferruginea  compounds developed w i t h  from b a s a l l e a v e s o f S.  85  p-nitroaniline  ferruginea  88  I n t r a s p e c i f i c c r o s s e s between d i f f e r e n t populations  19.  84  o f S.  ferruginea  G l a c i a l map o f western N o r t h America  89 90  X  ACKNOWLEDGEMENTS  I am s i n c e r e l y g r a t e f u l t o Dr. K a t h e r i n e d i r e c t i o n and a d v i c e ; and  assistance  I. Beamish f o r h e r  t o Dr. W. B. S c h o f i e l d f o r h i s p l e a s a n t  company  i n f i e l d work and e s p e c i a l l y f o r long d i s c u s s i o n s and  constructive c r i t i c i s m during  the c o u r s e o f t h i s study; to Dr. T.M.C.  T a y l o r , Dr. Thana B i s a l p u t r a and Dr. E. B. Tregunna f o r t h e i r i n valuable  c r i t i c i s m and a s s i s t a n c e i n p r e p a r a t i o n  to Dr. B. A.  of t h i s manuscript;  Bohm and Dr. K e i t h Moore f o r t h e i r a s s i s t a n c e i n chromato-  g r a p h i c work and to Dr. G. E. Rouse f o r h i s h e l p f u l s u g g e s t i o n s on glaciation; assistance.  to Mr. J . Thorpe f o r h i s m e c h a n i c a l and t e c h n i c a l I a l s o acknowledge the h e l p w i l l i n g l y rendered by  b o t a n i s t s , f o r e s t e r s and n a t u r a l i s t s who c o l l e c t e d p l a n t m a t e r i a l . I am g r a t e f u l to the c u r a t o r s o f many h e r b a r i a f o r the l o a n o f specimens. I wish to r e c o r d my indebtedness to the Department o f Botany, U n i v e r s i t y o f B r i t i s h Columbia, f o r f a c i l i t i e s of t h i s study.  throughout a l l phases  T h i s p r o j e c t was supported by NRC Grant A636 and the  Dean's Committee on Research, U n i v e r s i t y o f B r i t i s h Columbia. I would l i k e  Finally,  to thank my w i f e , Mrs. Raj Randhawa, f o r her support and  assistance during  the r e s e a r c h  and p r e p a r a t i o n  o f the t h e s i s .  INTRODUCTION S a x i f r a g a i s a l a r g e genus comprising Engler  and  Irmscher  (1916).  Due  species according  to a l a r g e amount o f  w i t h i n the genus E n g l e r r e c o g n i z e d f u r t h e r d i v i d e d i n t o groups.  302  variability  f i f t e e n s e c t i o n s , most of them  S a x i f r a g a f e r r u g i n e a Graham b e l o n g s  to h i s s e c t . B o r a p h i l a group  Stellares.  Saxifraga ferruginea i s a perennial species with,  like a l l  members of the group S t e l l a r e s , a r o s e t t e of more or l e s s  succulent  basal leaves.  tall  I t s d i f f u s e i n f l o r e s c e n c e grows 1 to 4 dm.  bears white star-shaped  flowers.  The  each have a p a i r of conspicuous y e l l o w p e t a l s l a c k spots.  described  spots w h i l e  petals  the o t h e r  two  propagules).  p l a n t s of S a x i f r a g a f e r r u g i n e a from which Graham (1829)  t h i s s p e c i e s were c u l t i v a t e d a t the R o y a l B o t a n i c  Edinburgh.  Graham g i v e s n e i t h e r  However, the s p e c i e s i s now the c o a s t  upper t h r e e o f i t s f i v e  and  F l o w e r s are numerous, sometimes p a r t l y to n e a r l y  a l l r e p l a c e d by p l a n t l e t s ( v e g e t a t i v e The  to  type l o c a l i t y nor  western C a l i f o r n i a , and  distribution.  known to range from A l a s k a  through B r i t i s h Columbia, Washington and  Garden,  southward  Oregon to  along  north-  i n the Rocky Mountains from e a s t c e n t r a l  B r i t i s h Columbia south to western Montana and p l a n t u s u a l l y grows i n s u b a l p i n e found a l s o down to sea l e v e l .  Idaho ( F i g . 1).  to a l p i n e l o c a l i t i e s , but  can  The be  I t commonly grows i n r o c k c r e v i c e s on  exposed o u t c r o p s where adequate m o i s t u r e i s a v a i l a b l e i n the blooming season, o f t e n i n the form o f The  seepage.  s p e c i e s complex e x h i b i t s c o n s i d e r a b l e  v a r i a t i o n i n many  c h a r a c t e r s , and  the v a r i a b i l i t y has been r e c o g n i z e d by  taxonomists i n d i f f e r e n t ways. into four species. under one  Small  (1905) s p l i t  s p e c i e s e.g.  (1968), two  the complex  Others r e c o g n i z e d o n l y i n f r a - s p e c i f i c Engler and  Irmscher  Johnson (1923), s i x v a r i e t i e s ; H i t c h c o c k Hulten  different  varieties.  The  (1916), t h r e e  taxa varieties;  e t a l (1961), two  varieties  d e c i s i o n s have been based  on  herbarium specimens a l o n e or on herbarium specimens supplemented f i e l d work.  The  c h a r a c t e r s used a r e the degree of p l a n t l e t  by  forma-  t i o n i n the i n f l o r e s c e n c e , s i z e of f l o w e r s , the dimensions of p e t a l s and  the morphology o f b a s a l and b r a c t e a l l e a v e s .  Such c h a r a c t e r s  have taxonomic v a l u e o n l y i f t h e i r g e n e t i c s t a b i l i t y and mental v a r i a b i l i t y have been c a r e f u l l y i n v e s t i g a t e d and and  i f they c o r r e l a t e i n such a way  as to i d e n t i f y  The main o b j e c t i v e o f the p r e s e n t  environunderstood,  populations.  i n v e s t i g a t i o n i s to d e t e r  mine the extent o f v a r i a b i l i t y w i t h i n the s p e c i e s complex not from the morphology o f l i v i n g and  pressed  only  p l a n t s but a l s o from o t h e r  a s p e c t s of b i o l o g y not p r e v i o u s l y a p p l i e d to S a x i f r a g a f e r r u g i n e a i . e . c y t o l o g y , embryology, anatomy, chromatography, c r o s s a b i l i t y  and  comparative growth response under u n i f o r m growing c o n d i t i o n s . Furthermore t h i s study aims a t d e t e r m i n i n g  whether or not  c o r r e l a t i o n s of c h a r a c t e r s , or of c h a r a c t e r s and would j u s t i f y the e s t a b l i s h m e n t  there  are  d i s t r i b u t i o n , which  o f i n f r a s p e c i f i c taxa.  3 MATERIAL L i v e p l a n t s as w e l l as herbarium specimens were used i n the study o f v a r i a b i l i t y i n S a x i f r a g a f e r r u g i n e a . much o f the s p e c i e s ' range as p o s s i b l e .  F i e l d work covered as  Many p o p u l a t i o n s  were thus  examined i n t h e i r n a t u r a l h a b i t a t s and m a t e r i a l was c o l l e c t e d i n the field  t o be f i x e d , p r e s s e d , o r kept a l i v e .  obtained  through the k i n d c o o p e r a t i o n  n a t u r a l i s t s whose h e l p  P l a n t m a t e r i a l was a l s o  o f s e v e r a l b o t a n i s t s and  i s g r a t e f u l l y acknowledged.  of c o l l e c t i o n s i t e s and r e f e r e n c e s  The complete  list  to voucher specimens a r e g i v e n i n  Table I. Table I C o l l e c t i o n s i t e s , voucher specimens and chromosome numbers i n Saxifraga ferruginea Collection site  Voucher specimen  Chromosome number  ALASKA: 1.  K o d i a k I s . , P i l l a r Mountain.  Randhawa 1586 UBC  n=10  2.  Juneau, Montana Creek.  Randhawa 1587 UBC  n=19  16. Randhawa 1588 UBC  n=19  Randhawa .1589 UBC  n=10  BRITISH COLUMBIA: 3.  P r i n c e Rupert, 36 mi. e a s t on Hwy  4.  Alliford  Bay, Queen C h a r l o t t e I s l a n d s . (Q.C.I.)  5.  Moresby Mountain, Q.C.I.  Randhawa 1590 UBC  n=10  6. 7.  Onward P o i n t , Q.C.I. Randhawa 1591 UBC Carew Bay, Q.C.I. Young i n f l o r e s c e n c e s o n l y c o l l e c t e d by W. B. S c h o f i e l d .  n=10 n=10  8.  Tar I s l a n d .  W . G r i f f i t h s.n.UBC  n=10  9.  Haines Road.  Beamish 681802 UBC  n=19  Suzette  W . G r i f f i t h s.n.UBC  n=10  10.  Bay, Dowager I s .  4 11.  Mt.  12.  Revelstoke.  F.Fodor  s.n. UBC  n=19  Mause Creek, F o r t S t e e l e .  F.Fodor s.n. UBC  n=10  13.  Vancouver I s l a n d , Campbell R i v e r .  Randhawa 1612 UBC  n=19  14.  L i g h t House Park, Vancouver.  Randhawa; 1499 UBC  n=19  15.  Richardson P o i n t ,  Randhawa  1485 UBC  n=19  16.  Vancouver, Mt. Seymour Park.  Randhawa 1498 UBC  n=19  17.  Squamish.  Randhawa 1500 UBC  n=19  Vancouver.  WASHINGTON: 18.  Mt. P i l c h u k , e a s t o f E v e r e t t . Young r o s e t t e s c o l l e c t e d by W. B. S c h o f i e l d .  n<=19  19.  M a r i o n County.  n=10  20.  C r a t e r Lake N a t i o n a l Park.  OREGON: Sherk 62653 UBC R i c h a r d M. Brown s.n. UBC  n=10  Herbarium specimens from the e n t i r e range o f the s p e c i e s were k i n d l y made a v a i l a b l e by t h e c u r a t o r s o f t h e h e r b a r i a o f t h e i n s t i t u t i o n s listed  below: S t a t e C o l l e g e o f Washington Pullman (WSP) U n i v e r s i t y o f Washington  S e a t t l e (WTU)  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 (OSC) U n i v e r s i t y o f M i n n e s o t a Herbarium (MIN) U n i v e r s i t y o f C a l i f o r n i a (UC) Jepson Herbarium, U n i v e r s i t y o f C a l i f o r n i a M i s s o u r i B o t a n i c a l Garden (MO) N a t i o n a l Museum o f Canada (CAN) U n i v e r s i t y o f A l a s k a (ALA)  (JEPS)  P l a n t Research I n s t i t u t e , Ottawa. University of Calgary  (DAO)  (UAC)  U n i v e r s i t y o f A l b e r t a , Edmonton (ALTA) U n i v e r s i t y of B r i t i s h  Columbia  (UBC)  M a t e r i a l from a l l the p o p u l a t i o n s l i s t e d for  chromosome counts.  i n T a b l e I was  L i v e p l a n t s from s e v e r a l p o p u l a t i o n s (#  16, 20) were c u l t i v a t e d a t the U n i v e r s i t y o f B r i t i s h u n i f o r m greenhouse c o n d i t i o n s .  Juneau, P r i n c e Rupert, A l l i f o r d studies.  Columbia  1-4,  under  O b s e r v a t i o n s on l i v e p l a n t s s u p p l e -  mented the survey o f h e r b a r i u m specimens.  used f o r d e t a i l e d  examined  P o p u l a t i o n s from Kodiak,  Bay, Vancouver and M a r i o n County were  6 METHODS, OBSERVATIONS, AND  RELEVANT LITERATURE  T h i s i n v e s t i g a t i o n embodies i n f o r m a t i o n on S a x i f r a g a d e r i v e d from s i x a s p e c t s  ferruginea  o f b i o l o g y : c y t o l o g y , morphology, embryology,  anatomy, chromatography, and c r o s s a b i l i t y .  Obviously  the  techniques  employed f o r the c o l l e c t i o n and p r e p a r a t i o n o f m a t e r i a l were  different  i n each l i n e o f study and t h e r e f o r e they have been i n c l u d e d i n the appropriate general  sections with observations  and r e l e v a n t d i s c u s s i o n .  d i s c u s s i o n i n t e g r a t e s the f i n d i n g s a t the end.  A  7 CYTOLOGICAL STUDIES Methods: Young i n f l o r e s c e n c e s from d i f f e r e n t  l o c a t i o n s (Table I ) were  f i x e d i n Carnoy's e t h y l a l c o h o l - g l a c i a l a c e t i c a c i d Navashin's f l u i d , a b b r e v i a t e d as ' C r a f c o u n t i n g , anther  (Johnson 1940).  M e i o s i s was s t u d i e d i n a l l stages.  o f Mj to T_ i n m i c r o s p o r o c y t e s  Hj. was c o n s i d e r e d normal i f  the chromosomes l a y on the metaphase p l a t e , and was l o s t  i n the cytoplasm.  used i n the study o f p o l l e n v i a b i l i t y , considered v i a b l e .  L a t e r a l views  from both d i p l o i d and p o l y p l o i d p o p u l a t i o n s  were scored f o r i r r e g u l a r i t i e s i n d i v i s i o n .  chromatin  F o r chromosome  smears were s t a i n e d u s i n g 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 -  carmine (Snow 1963).  all  (3:1) or i n  and T^ i f no  Lactophenol-cotton  b l u e s t a i n was  the blue-stained g r a i n s b e i n g  One thousand p o l l e n g r a i n s were counted from 10  f l o w e r s o f 3 t o 5 p l a n t s i n each p o p u l a t i o n examined. The  Vancouver p o p u l a t i o n was s t u d i e d i n g r e a t e s t d e t a i l , and  u n l e s s otherwise  s t a t e d the d e s c r i p t i o n o f m e i o s i s and the f i g u r e s a r e  based on t h i s p o p u l a t i o n .  Other p o p u l a t i o n s were compared w i t h i t to  determine v a r i a b i l i t y . Observations  and r e l e v a n t  literature:  In most o f the m i c r o s p o r o c y t e s a t the  p l a t e ( F i g . 2B).  a l l the chromosomes assemble .  However sometimes chromatin  o u t s i d e the e q u a t o r i a l r e g i o n ( F i g . 2G).  can be  observed  During A_ the movement o f  chromosomes to the p o l e s i s n o t synchronous ( F i g . 2D) b u t a p p a r e n t l y the l a g g i n g chromosomes a r e u s u a l l y i n c l u d e d i n the t e l o p h a s e n u c l e i ( F i g . 2E).  One b i v a l e n t was sometimes observed  as a l a g g a r d which remained  8 on or near the metaphase p l a t e  ( F i g . 2H) and i n such cases the daughter  chromosomes were p r o b a b l y l e f t out o f the c h r o m a t i n were a l s o observed a t A ^  nuclei.  ( F i g . 21).  Laggards or  lost  The r e s u l t o f the  i r r e g u l a r i t i e s o f m e i o s i s became obvious a t the t e t r a d  stage when  i n s t e a d o f , or i n a d d i t i o n t o , the u s u a l f o u r m i c r o s p o r e s t h e r e were one or more m i c r o n u c l e i w i t h i n the m i c r o s p o r o c y t e w a l l are formed  Micronuclei  from l o s t chromatin and t h e i r presence foreshadows  i n the amount o f v i a b l e p o l l e n . produced  ( F i g . 2F).  88.5% v i a b l e p o l l e n .  d u r i n g m e i o s i s i n S.  I n 1967  a reduction  f i v e p l a n t s of t h i s p o p u l a t i o n  Beamish (1961) r e p o r t e d i r r e g u l a r i t i e s  f e r r u g i n e a and r e c o r d e d abnormal m i c r o s p o r e s i n  tetrads. Extreme clumping o f chromatin was T  observed a t MLj. (Fig. 2C) or a t  i n m a t e r i a l c o l l e c t e d from a r o c k outcrop on Mt.  Seymour on a v e r y  warm a f t e r n o o n (110°F t h r e e i n c h e s above the r o c k s u r f a c e ) . clumping was and was  r e p o r t e d i n the j>. montanensis  Similar  p l e x u s by Beamish (1961)  suggested to be due to an environmental e f f e c t ,  such as  temperature. The p o l l e n g r a i n s i n one f l o w e r were observed to germinate i n s i t u and clumps o f p o l l e n were formed by the entwining p o l l e n (Fig.  2N).  (Fig.  2,0).  The development  o f the p o l l e n tubes appeared  A chromosome count o f 19 p a i r s from the Vancouver r e p o r t e d by Beamish (1961).  to be  tubes normal  p o p u l a t i o n was  The p r e s e n t study c o n f i r m s t h a t count.  N i n e t e e n p a i r s of chromosomes f o r another p o p u l a t i o n a t Campbell R i v e r are i l l u s t r a t e d  i n F i g u r e 2A.  9 Observations  and chromosome counts o f the m i c r o s p o r o c y t e s  o f o t h e r p o p u l a t i o n s r e v e a l e d d i p l o i d s as w e l l as p o l y p l o i d s (Table I ) .  A l l o t h e r p o l y p l o i d s , l i k e the Vancouver p o p u l a t i o n  exhibited i r r e g u l a r i t i e s during meiosis. found  t o be m o s t l y r e g u l a r ( F i g . 2M).  M e i o s i s i n d i p l o i d s was  However, even the d i p l o i d s  sometimes were i r r e g u l a r e.g. chromosomes o c c a s i o n a l l y lagged a t Aj (Figs.  2K,Q).  Such i r r e g u l a r i t i e s , a p p a r e n t l y l e a d to the  f o r m a t i o n o f m i c r o n u c l e i i n the t e t r a d s ( F i g s . of Mj. and A^ i n a d i p l o i d  2R,S).  A comparison  ( A l l i f o r d Bay p o p u l a t i o n ) and a p o l y p l o i d  (Vancouver p o p u l a t i o n ) appears i n T a b l e I I .  Chromosome counts o f  C r a t e r Lake, Kodiak I s . , and A l l i f o r d Bay p o p u l a t i o n s a r e i l l u s t r a t e d (Figs.  23, L and P ) . Table I I  Meiosis i n microsporocytes  o f j3. f e r r u g i n e a Graham ( d i p l o i d and p o l y p l o i d )  Metaphase I Population  A l l i f o r d Bay (n=10) Vancouver (n=19)  Plants examined  Anaphase I  Cells examined  Cells with irregularit i e s 7,  Cells examined  3  789  0.6  523  1.3  4  841  2.5  750  5.2  P o l l e n v i a b i l i t y was r e c o r d e d p l o i d populations.  As expected  C e l l s with irregularities %  i n t h r e e d i p l o i d and t h r e e p o l y -  on the b a s i s o f m e i o t i c behaviour the  p o l y p l o i d s produced c o n s i d e r a b l y l e s s v i a b l e p o l l e n than the d i p l o i d s  10 Table I I I i n d i p l o i d and p o l y p l o i d p o p u l a t i o n s o f S. Graham  Pollen v i a b i l i t y  Diploid Population  (n=10)  Polyploid  Kodiak  Alliford  Crater  Prince  Is.  Bay  Lake  Rupert  Plants examined  3  5  4  5  Viable p o l l e n 7o  93.4  (Table  III).  95.4  93.1  81.9  (n=19)  Vancouver  Juneau  5  5  88.5  One n o t a b l e e x c e p t i o n was the group o f 5 p l a n t s  Juneau which had u n e x p e c t e d l y good p o l l e n .  was re-examined.  The l e v e l o f v i a b l e p o l l e n had  from 81.9% i n 1967 t o 90.67. i n 1968.  95.2  from  An i n t e r e s t i n g s i d e l i g h t  on p o l l e n p r o d u c t i o n came i n 1968 when p o l l e n from the P r i n c e population  ferruginea  Rupert increased  11  MORPHOLOGY AND THE L I F E CYCLE Methods: The development to  o f p l a n t s was s t u d i e d from seed g e r m i n a t i o n  the shedding o f seed a t m a t u r i t y .  Seeds were c o l l e c t e d  from  the  Vancouver  p o p u l a t i o n f o r a seed g e r m i n a t i o n experiment.  Space  for  the g e r m i n a t i o n experiment was k i n d l y made a v a i l a b l e i n the  c o n t r o l l e d e n v i r o n m e n t a l chambers by the Phycology, P l a n t S c i e n c e and P l a n t P h y s i o l o g y l a b o r a t o r i e s .  However, o n l y those temperature  c o n d i t i o n s and p h o t o p e r i o d s ( T a b l e IV) c o u l d be used t h a t were designed for  the experiments conducted by the above mentioned  each growth chamber d u p l i c a t e p e t r i 100 seeds on m o i s t f i l t e r d u r i n g the experiment. (petri  paper.  sections.  In  d i s h e s were p l a c e d , each h a v i n g  The f i l t e r papers were kept m o i s t  Seeds were a l s o grown i n complete  darkness  d i s h e s covered w i t h b l a c k tape) a t c o n s t a n t temperatures o f  60°F, 46°F, and 41°F to check the i n f l u e n c e o f l i g h t g e r m i n a t i o n c a p a c i t y o f S.  on the  ferruginea.  In a d d i t i o n to the o b s e r v a t i o n s made on the p r e s s e d  specimens  c o v e r i n g the e n t i r e range, l i v e p l a n t s from Kodiak, Juneau, Rupert, A l l i f o r d  Bay, Onward P o i n t , Vancouver  p o p u l a t i o n s were'cultivated "and  Prince  and M a r i o n County  were examined f o r the f o l l o w i n g  c h a r a c t e r s : l e n g t h and w i d t h o f seeds, l e n g t h o f the ' t u b e r c l e s '  12  o f seeds measured from the c e n t r a l row  on convex s i d e ; shape and  s i z e o f p e t a l s ; s i z e o f p o l l e n g r a i n s ; v a r i a t i o n i n the b a s a l b r a c t e a l l e a v e s w i t h r e s p e c t to s i z e , number and d e n s i t y o f pubescence and  size of h a i r s ;  extent of p l a n t l e t f o r m a t i o n  was  (length o f t e e t h ;  s i z e o f stomata; and  i n the i n f l o r e s c e n c e .  e f f i c i e n c y o f p l a n t l e t s as p r o p a g u l e s ,  and  the  To observe  one b a t c h o f f i v e  plantlets  removed from the i n f l o r e s c e n c e , o f each of t h r e e p l a n t s  taken  from the Vancouver p o p u l a t i o n and  p l a n t e d e a r l y i n the season.  Another l o t o f p l a n t l e t s was  i n t a c t , and was  the  left  the  planted late i n  season. S p e c i a l techniques  used f o r p r e p a r i n g the m a t e r i a l f o r obser-  v a t i o n s a r e g i v e n below:1.  F o r making the permanent mounts of p e t a l s , the  were f i x e d i n a b s o l u t e a l c o h o l and g l a c i a l a c e t i c a c i d f l o w e r s were passed through two  flowers (3:1).  The  changes of a b s o l u t e a l c o h o l and  s t a i n e d f o r 30 to 45 minutes i n a m i x t u r e  of 1 p a r t Orange G  oil  s t a i n e d f l o w e r s were a g a i n  and 4 p a r t s of a b s o l u t e a l c o h o l .  passed through two The  The  changes of a b s o l u t e a l o c h o l , each f o r two  intclove  minutes.  i n d i v i d u a l p e t a l s were then detached and mounted i n e u p a r a l .  Ten p l a n t s were examined from each p o p u l a t i o n t a k i n g three from one 2.  flowers  plant. A random sample of p o l l e n g r a i n s was  taken from t h r e e p l a n t s  13. i n each p o p u l a t i o n and s t a i n e d i n l a c t o p h e n o l - c o t t o n b l u e .  The  s t a i n e d p o l l e n g r a i n s were used f o r measurements. 3.  The o u t l i n e s o f the b a s a l and b r a c t e a l l e a v e s were drawn  d i r e c t l y or from c o n t a c t p r i n t s made on p h o t o s e n s i t i v e paper exposed to ammonia vapour. 4.  O b s e r v a t i o n s on the pubescence o f f u l l y developed b a s a l  l e a v e s were made on b o t h s u r f a c e s o f the l e a f lamina.  Pubescence was  examined a t f o u r p o i n t s , two on each s i d e o f the mid r i b w i t h i n an a r e a 2 of 2 mm .  Ten p l a n t s were sampled from each p o p u l a t i o n by examining  two l e a v e s per p l a n t . 5.  The s i z e o f stomata was measured from the lower e p i d e r m i s  o f l e a v e s taken from t e n p l a n t s i n each p o p u l a t i o n .  The e p i d e r m a l  s t r i p s were s t a i n e d i n s a f r a n i n and f a s t green and mounted i n Canada balsam. O b s e r v a t i o n s and r e l e v a n t Seeds:-  literature:  The seeds a r e t u b e r c u l a t e h a v i n g 10 t o 15 l e n g t h w i s e  rows o f s p i n e - l i k e p r o t u b e r a n c e s .  The t u b e r c l e s ( F i g s . 3A, B) a r e l o n g e s t  (up to 63 microns) i n the m i d d l e rows on the convex s i d e o f the seeds as compared t o the f l a n k i n g rows where they range from mere d o t s to p r o j e c tions of varying lengths. 1030 microns.  The l e n g t h o f seeds v a r i e s from 440 m i c r o n s t o  A random sample o f 1000 seeds weighed 0.0233 grams.  comparative study o f seed c h a r a c t e r s i n s i x p o p u l a t i o n s cated  The  ( T a b l e V) i n d i -  s l i g h t l y l a r g e r seeds i n P r i n c e Rupert and Vancouver  populations.  The seeds from M a r i o n County and Vancouver p o p u l a t i o n s had more prominent t u b e r c l e s than those from Kodiak and A l l i f o r d  Bay p o p u l a t i o n s .  these c h a r a c t e r s a r e h i g h l y v a r i a b l e i n a s i n g l e p o p u l a t i o n .  However  14. Seed g e r m i n a t i o n : Vancouver  G e r m i n a t i o n was  observed i n seeds o f the  p o p u l a t i o n i n p o t s i n the greenhouse  the c o n t r o l l e d environment  chamber.  and i n p e t r i d i s h e s i n  When proper m o i s t u r e i s a v a i l a b l e ,  the seeds s w e l l and the seed c o a t c r a c k s a l o n g the rows o f t u b e r c l e s . The  s t o u t r a d i c l e p i e r c e s through the endosperm and develops i n t o  p r i m a r y r o o t system c o a t i s pushed  ( F i g s . 3C-J).  the  Germination i s e p i g e a l and the seed  o f f by the u n f o l d i n g c o t y l e d o n s .  The c o t y l e d o n s develop  c h l o r o p h y l l and p e r s i s t f o r a l o n g time, growing up to 5mm. i n l e n g t h , until  they a r e crowded and shaded under  the d e v e l o p i n g r o s e t t e .  s e e d l i n g s a r e so extremely s m a l l t h a t i t i s d i f f i c u l t their natural habitat u n t i l  to f i n d  The  them i n  they grow t o an a p p r e c i a b l e s i z e .  The  seed-  l i n g p l a n t s set seed i n about 85 days, i n c l u d i n g the g e r m i n a t i o n time o f 15 days. The r e s u l t s o f the g e r m i n a t i o n experiment a r e p r e s e n t e d i n T a b l e IV. T a b l e IV Effect of different seeds.  temperatures and p h o t o p e r i o d s on the g e r m i n a t i o n o f  Temperature  °F Photoperiod  Germination  Growth Chamber  *Day  1  105  65  8 hours  22.5  2  95  95  8 hours  nil  3  95  80  8 hours  3.5  4  95  65  8 hours  82.0  5  105  65  14 hours  nil  6  95  95  14 hours  nil  7  95  80  14 hours  nil  8  95  65  14 hours  17.0  9  75  65  16 hours  10  60  60  16 hours  58.5  11  46  46  16 hours  84.5  12  41  41  16 hours  nil  Night  7o  * The day temperatures c o r r e s p o n d to the p h o t o p e r i o d . ^ F u n g a l growth i n p e t r i d i s h e s .  **21.0  15. Temperatures o f 46°F to 65°F appear to be near optimum f o r seed g e r m i n a t i o n  among the range o f temperatures a v a i l a b l e .  High  temperatures up to 95°F c o u l d be t o l e r a t e d f o r 8 hours (chamber 4) but  f o r longer p e r i o d s  mental.  Seeds f a i l e d  (chambers 5,8 and 9) they proved h i g h l y d e t r i t o germinate a t 41°F and temperatures above  75°F were a l s o n o t f a v o u r a b l e .  G e r m i n a t i o n i n complete darkness was  comparable to t h a t i n 16 hour p h o t o p e r i o d  a t the same temperatures  except t h a t i t was d e l a y e d by about a week. l i g h t i s n o t important  This indicates that  f o r seed g e r m i n a t i o n .  Lewin (1960) observed  t h a t l i g h t was n o t an e s s e n t i a l f a c t o r f o r the g e r m i n a t i o n  o f seed o f  Saxifraga v i r g i n i e n s i s . A d v e n t i t i o u s Roots:adventitious r o o t s developing  The p r i m a r y r o o t system i s r e i n f o r c e d by from the base o f the r o s e t t e .  v a r y from 0.3 to 1.0 mm. i n diameter and even r e a c h  The r o o t s  1.5 mm. i n some cases.  A d v e n t i t i o u s r o o t s a r e sometimes a l s o i n i t i a t e d i n p l a n t l e t s in s i t u as observed i n the M a r i o n County and Vancouver Leaves:-  populations.  The l e a v e s a r e somewhat s u c c u l e n t , o b l a n c e o l a t e o r  s p a t u l a t e , t h i c k , s h a r p l y toothed  above the m i d d l e , pubescent t o a  g r e a t e r o r l e s s e r e x t e n t , and taper i n t o a s h o r t o r l o n g p e t i o l e . f i r s t young t r u e l e a v e s have o n l y two shallow Successive  The  t e e t h on the l e a f margin.  l e a v e s have 3 t o 15 t e e t h per l e a f .  The shape o f the b a s a l  and b r a c t e a l l e a v e s i s n o t c o n s i s t e n t even w i t h i n a s i n g l e p o p u l a t i o n ( F i g s . 4-7).  Dentation  i s a l s o n o t a s t a b l e c h a r a c t e r as the l e n g t h  of t e e t h i n b a s a l l e a v e s v a r i e s from 1 t o 3 mm. Pubescence:length. l a r head.  The l e a v e s p o s s e s s g l a n d u l a r h a i r s o f v a r i a b l e  The h a i r s a r e m u l t i c e l l u l a r and m u l t i s e r i a t e w i t h a m u l t i c e l l u The d e n s i t y o f pubescence v a r i e s w i t h i n the s p e c i e s from  16 o c c a s i o n a l h a i r s to 31 h a i r s per u n i t a r e a  (2 mm.).  s i t u a t i o n has been observed i n A l l i f o r d Bay  and  The  latter  Onward P o i n t  p o p u l a t i o n s , where the p l a n t s grow on the r o c k s f a c i n g the The h a i r s on the  l e a v e s o f the Onward P o i n t p o p u l a t i o n  however, extremely s h o r t so t h a t the l e a v e s appear to be g l a b r o u s  and r a t h e r glaucous.  pubescence was  maintained  of B r i t i s h Columbia. pubescence and (Table V ) .  v a r i e s from 0.2  to 2.8  but  the h a i r s on  mm.  profuse  examined have  the a b a x i a l s u r f a c e of the  alpine populations glabrous  macroscopically  c h a r a c t e r of  A l l other p o p u l a t i o n s  have a p p a r e n t l y  are,  i n the p l a n t s c u l t i v a t e d at U n i v e r s i t y  almost none on  The  The  sea.  leaves The  scanty leaves  on Queen C h a r l o t t e I s l a n d s a l s o  ( T a y l o r 1968).  The  length of h a i r s  l e a f margin i s always  the p e t i o l a r margin are much longer  ciliated, than those  on  the l a m i n a l margin. Stomata:- The  l e n g t h o f stomata i s v a r i a b l e w i t h i n  A range i n s i z e from 30 to 39 microns was Flower:- P l a n t s f l o w e r d u r i n g germination.  The  The  species.  seed  i n the r o s e t t e and  growth of the p e t a l s i s slow  stages of flower development but near the time of  a n t h e s i s they develop v e r y r a p i d l y . having  i n the  the f i r s t year a f t e r  inflorescence i s initiated  develops i n t o a p a n i c l e of cymes. i n the e a r l i e r  recorded  populations.  The  c o r o l l a i s zygomorphic,  t y p i c a l l y f i v e petals.The upper three p e t a l s are ovate or  s a g i t t a t e with p e t a l lamina  two  and  y e l l o w i s h t r i a n g u l a r spots near the base o f  c l o s e to the c e n t r a l v e i n ( F i g . 8).  p e t a l s are l a n c e o l a t e and i s notable  l a c k spots.  f o r the absence o f spots and  The  The  Onward P o i n t  the  lower  two  population  f o r the presence of o n l y  ovate p e t a l s ( F i g s . 8-19, -20).  P e t a l s i z e v a r i e s from a  minimum o f 4.0 mm. long and 1.9 mm. broad  i n the M a r i o n County  p o p u l a t i o n to 9.0 mm.long and up t o 3.8 mm. broad Bay  population.  i n the S u z e t t e  The Tar I s l a n d p o p u l a t i o n produced  w i t h p e t a l s up to 8.0 mm.long and 4.9 mm.broad.  flowers  The comparison  of s i z e s o f p e t a l s f o r d i f f e r e n t p o p u l a t i o n s a l o n g the c o a s t i s g i v e n i n T a b l e V. and  Flowers w i t h 6 o r 7 p e t a l s a r e n o t uncommon  8 or 9 occur o c c a s i o n a l l y .  The two l a n c e o l a t e s p o t l e s s p e t a l s  a r e sometimes r e p l a c e d by ovate o r s a g i t t a t e s p o t t e d p e t a l s so t h a t a l l p e t a l s a r e then s i m i l a r . cling  tightly  The s e p a l s a r e r e f l e x e d and  to the f l o w e r p e d i c e l by the time the f l o w e r i s h a l f  expanded. The  anthers begin  0.5 mm. long. completely protandry.  to d e h i s c e when the s t y l e s a r e o n l y 0.2 to  The stigma becomes r e c e p t i v e a f t e r  shed from the a n t h e r s .  the p o l l e n i s  The p l a n t , t h e r e f o r e , e x h i b i t s  There i s no d i s t i n c t d i f f e r e n c e i n the s i z e o f p o l l e n  g r a i n s among p o p u l a t i o n s and t h i s v a r i a b i l i t y  therefore  d e t e c t i o n o f d i p l o i d and p o l y p l o i d p o p u l a t i o n s .  The s m a l l w h i t e  f l o w e r s o f S a x i f r a g a f e r r u g i n e a a r e adapted t o d i p t e r o u s but bees have a l s o been observed Plantlets:-  visiting  prevents  pollinators  them.  The i n f l o r e s c e n c e i n most cases develops  some  p l a n t l e t s which r e p l a c e the f l o w e r s and serve as v e g e t a t i v e propagules  ( F i g . 9A,B).  When p l a n t l e t s r e p l a c e f l o w e r s v e r y e a r l y d u r i n g  development, the i n f l o r e s c e n c e then bears more p l a n t l e t s flowers.  than  I n other cases o n l y f l o w e r s a r e produced i n the b e g i n n i n g  o f the season w h i l e the development o f p l a n t l e t s i s delayed.  As  18 observed i n the c l o n e s d u r i n g two y e a r s o f c u l t i v a t i o n ,  the  p r o p o r t i o n o f p l a n t l e t s and f l o w e r s i s v a r i a b l e , w i t h i n  and  between d i p l o i d and p o l y p l o i d p o p u l a t i o n s . Alliford  For example,  Bay and Marion County p o p u l a t i o n s are d i p l o i d  p a t t e r n of p l a n t l e t production i s h i g h l y v a r i a b l e .  though  the  P l a n t l e t s are  r a r e l y seen i n the former whereas the l a t t e r produces  largely  plantlets. The p l a n t l e t s , when detached and p l a n t e d e a r l y i n the season, grow r o o t s w i t h i n a week, bear i n f l o r e s c e n c e s and i n about 73 days.  s e t seed  However p l a n t l e t s detached and p l a n t e d l a t e  i n the season develop i n t o r o s e t t e s w i t h o u t i n i t i a t i n g i n f l o r e s c e n c e , p r o b a b l y due  an  to an u n f a v o u r a b l e p h o t o p e r i o d .  The  c a p a c i t y o f the p l a n t l e t s to develop i n t o f u l l grown p l a n t s w i t h i n f l o r e s c e n c e s i n one house.  Presuming  season has been demonstrated  i n the green-  t h a t the same c a p a c i t y e x i s t s i n n a t u r e , the  p l a n t l e t s can complete a second c y c l e o f r e p r o d u c t i o n i n the same season.  Rabotnov (1946) noted t h a t i n d i v i d u a l s which have  developed v e g e t a t i v e l y have a s h o r t e r of  ' v i r g i l period  1  (beginning  growth to s e x u a l m a t u r i t y ) than those developed from seeds.  Taylor  (1965) found i n Lithophragma  (Saxifragaceae) that p l a n t s  r a i s e d from b u l b i l s reached m a t u r i t y i n a s h o r t e r time than those grown from seed. Rhizome:-  The rhizomes emerge from the base o f the o l d  19  r o s e t t e s and g i v e r i s e  to many new  crowns ( F i g s . 9C, D).  These crowns become anchored by a d v e n t i t i o u s r o o t s . f i v e r o s e t t e s have been observed r e t a i n i n g t h e i r to the p a r e n t r o s e t t e .  Up to  attachment  N o r m a l l y the c o n n e c t i o n s a r e severed  through decay so t h a t each crown i s u l t i m a t e l y e s t a b l i s h e d as an independent p l a n t . The L i f e C y c l e : i s summarized  i n Figure  The r e p r o d u c t i v e 10.  b i o l o g y o f the  species  The i n d i v i d u a l s o f a n a t u r a l popu-  l a t i o n develop from three d i f f e r e n t s o u r c e s : f i r s t ,  sexually  from seed, a p r o c e s s which imparts v a r i a b i l i t y as a r e s u l t o f cross p o l l i n a t i o n ;  second, v e g e t a t i v e l y from p l a n t l e t s ; t h i r d ,  a l s o v e g e t a t i v e l y but from the new  crowns developed from the  rhizomes around the p a r e n t p l a n t .  The l a s t  tuate  the p a r e n t a l  genotype.  two methods perpe-  20 Table V *...  '.' . V a r i a t i o n i n m o r p h o l o g i c a l c h a r a c t e r s o f d i f f e r e n t p o p u l a t i o n s  Plant part  Kodiak  Juneau  Prince Rupert  Alliford Bay Q. C. I.  Length )i  636.6  675.2  728.2  697.2  Breadth v  339.7  340.0  371.1  340.1  Tubercle length /i  21.6  35.2  31.6  26.6  -  8.1  2.6  3.5  16.9  Pubescence few abaxial surfac^mm-.  few  few  Length of h a i r cm.  1.0  1.5  Length of l e a f cm.  3.6  Number of teeth/leaf Length o f t e e t h mm.  Characters  SEED  Pubescence adaxial s u r f aceyAmm.  LEAVES  Length o f stomata u FLOWER PETALS  Length  POLLEN GRAINS  Size i n u  *  mm.  Onward Point Q.C. I.  Vancouver M a r i o n County B.C. Oregon 853.3  644.0  416.2  334.1  50.6  46.7  18.1  3.2  2.5  15.9  17.1  few  1.5  1.9  0.2  1.6  0.5  2.5  4.2  3.7  4.0  4.7  3.9  8.7  5.4  8.5  8.9  10.6  9.9  9.4  1.7  1.5  2.1  1.9  1.4  2.2  1.8  34.5  39.1  35.0  31.9  38.3  30.1  6.0  5.9  5.6  5.8  6.1  5.1  16.6  16.8  18.1  15.6  17.7  16.1  6.0  -  few  Seed c h a r a c t e r s , average o f 90 measurements; Pubescence, average of 80 c o u n t s ; Length o f h a i r s , average o f 200 measurements; L e a f l e n g t h and d e n t a t i o n based on 20 l e a v e s taken from 10 p l a n t s ; Stomata, average o f 100 measurements; P e t a l s , average o f 90 measurements; P o l l e n g r a i n s , average o f 90 measurements.  21  EMBRYOLOGY Methods; Young i n f l o r e s c e n c e s , buds, f l o w e r s , and f r u i t s i n different  stages o f development  a l c o h o l and g l a c i a l a c e t i c a c i d  were f i x e d i n 95 p e r c e n t e t h y l (3:1).  O v a r i e s were d i s s e c t e d  out, except from v e r y s m a l l buds, and passed through a t e r t i a r y butyl alcohol embedded  (TBA) d e h y d r a t i o n s e r i e s .  i n Fisher  The m a t e r i a l was  tissuemat (m. p. 50° - 53°C).  S e c t i o n s were  c u t a t 10 to 12 microns and s t a i n e d w i t h a s a f r a n i n - f a s t combination.  green  Heidenhain"s i r o n h e m a t o x y l i n was used f o r some  e a r l y stages i n embryo sac development. Much o f the work on female development naturally pollinated material.  was done on  However, to check the t i m i n g o f  v a r i o u s s t e p s i n the p r o c e s s a r t i f i c i a l p o l l i n a t i o n s were a l s o made.  Flowers on seven p l a n t s o f the Vancouver p o p u l a t i o n  ^ c u l t i v a t e d a t the U n i v e r s i t y o f B r i t i s h Columbia were emasculated and bagged.  The stigmas o f the emasculated  f l o w e r s were c o n s i d e r e d  to be r e c e p t i v e and were p o l l i n a t e d when they s e c r e t e d a d r o p l e t at t h e i r  surface.  The p o l l i n a t e d f l o w e r s were f i x e d a t v a r i o u s  i n t e r v a l s a f t e r p o l l i n a t i o n i . e . 10 minute f i r s t hour, 6 hour  i n t e r v a l s f o r the  i n t e r v a l s up to 24 hours, then a t 48 hours and  72 hours. S a t i s f a c t o r y s e c t i o n s were d i f f i c u l t t r e a t e d mature seeds.  to o b t a i n from un-  However, when the seeds were s o f t e n e d ^  i n 1:1 g l a c i a l a c e t i c a c i d and 6% hydrogen p e r o x i d e f o r 4 hours  ^ L e a f l e t No. 40.  F o r e s t P r o d u c t s Research Lab. England 1956.  22 a t 60°C p r i o r  to the TBA d e h y d r a t i o n s e r i e s , good s e c t i o n s were  obtained. M a t e r i a l from the Vancouver p o p u l a t i o n was used f o r d e t a i l e d study d u r i n g the y e a r s 1965 and 1966 and the f o l l o w i n g o b s e r v a t i o n s and f i g u r e s a r e based l a r g e l y on t h i s  investigation.  The other f i v e p o p u l a t i o n s were l a t e r examined f o r comparison. O b s e r v a t i o n s and r e l e v a n t Ovule:-  literature  Ovules develop as s m a l l p r o t r u s i o n s o f the  n u c e l l u s i n t o the l o c u l u s o f the ovary when the sporogenous  tissue  of the a n t h e r s i s d i s t i n g u i s h a b l e by i t s d e e p l y s t a i n e d n u c l e i and dense cytoplasm.  When the o v u l e s a r e about 50 to 60 m i c r o n s l o n g  and have been p r o j e c t e d i n t o the c a v i t y by the e l o n g a t i n g  funiculus,  the megasporocyte appears as a c o n s p i c u o u s l y l a r g e c e l l o f the n u c e l l u s under the p a r i e t a l c e l l s i n the hypodermal r e g i o n ( F i g . The o v u l e s a r e thus c r a s s i n u c e l l a t e  (Maheshwari 1950).  11B).  The m i c r o -  s p o r o c y t e s by t h i s time have completed m e i o s i s and formed p o l l e n g r a i n s ( F i g . 11A).  The o v u l e s i n S a x i f r a g a f e r r u g i n e a a r e anatropous.  The young o v u l e s u s u a l l y develop a c u r v a t u r e even b e f o r e m e i o s i s b e g i n s i n the megasporocyte ovules s t i l l  ( F i g . 11D).  However, o c c a s i o n a l l y some  appear o r t h o t r o p i c a t t h i s stage ( F i g . H E ) .  The  growth o f the s i n g l e integument i s more r a p i d on the a b a x i a l than on the a d a x i a l s i d e and g r a d u a l l y the n u c e l l u s i s enveloped except a t the m i c r o p y l e .  At no stage o f development was any i n d i c a t i o n o f a  second integument observed. hieracifolia rufidula  One integument has been r e p o r t e d i n S.  (Wiggins 1959), S. i n t e g r i f o l i a  (Beamish 1967).  f  (Beamish 1963), and S.  The b i t e g m i c c o n d i t i o n has been r e p o r t e d i n  23 granulata  ( J u e l 1907), S. l i g u l a t a  (Pace 1912), and S.  diversifolia  (Saxena 1964). The innermost l a y e r o f the integument and the outermost layer o f the nucellus  are morphologically  alike until  n u c l e a t e stage o f embryo sac development.  The integumentary l a y e r  then becomes c o n s p i c u o u s as a d e e p l y s t a i n e d  s i n g l e - c e l l e d layer.  The c e l l s o f the outermost l a y e r o f the n u c e l l u s i n s i z e and l o s e d e n s i t y . layer  the 4 -  gradually  increase  A f t e r f e r t i l i z a t i o n the integumentary  s t a r t s c o l l a p s i n g so t h a t by about t h e d i v i s i o n o f the embryonal  c e l l i t appears as a dark s t r e a k o f t i s s u e around the l a r g e - c e l l e d nucellar  layer  ( F i g s . 12A-E).  The l a t t e r  i s d i s s o l v e d d u r i n g the  development o f t h e endosperm i n the m a t u r i n g seeds.  Such  modifi-  c a t i o n i n the integumentary and n u c e l l a r l a y e r s has n o t been i n other species  of Saxifraga.  to a l l the p o p u l a t i o n s  the  The megasporocyte undergoes the f i r s t  d i v i s i o n ( F i g . 11F) g i v i n g r i s e to a dyad.  second d i v i s i o n o f m e i o s i s a r e o r i e n t e d  (Fig.  o f development i s common  examined.  Megasporogenesis;meiotic  This pattern  reported  The s p i n d l e s a t  as a 'T*-shaped  11C). M e i o s i s i s g e n e r a l l y r e g u l a r b u t o c c a s i o n a l l y  chromosomes do n o t l i n e up on t h e M^ o r M_^ p l a t e .  tetrad some  The top b a r o f the  ' T' i s always a t an a n g l e t o t h e lower b a r b u t was n o t observed e x a c t l y at r i g h t angles.  I n some c a s e s the o r i e n t a t i o n i s n e a r l y  *T'-shaped t e t r a d was r e p o r t e d and j>. g r a n u l a t a integrifolia  also i n a hybrid  linear.  of Saxifraga  A  decipiens  (Schurhof f 1925), S>. v i r g i n i e n s i s (Chapman 1933), (Beamish 1963), and S. r u f i d u l a (Beamish 1967).  l i n e a r t e t r a d has been observed i n S.  l i g u l a t a , S_. sponhemica,  A  24 S.  cordifolia,  (Wiggins  S.  crassifolia  (Pace 1912), and  S.  hieracifolia  1959). A f t e r m e i o s i s i s complete,  the n o n - f u n c t i o n a l megaspores  s t a r t d i s i n t e g r a t i n g and can be seen as dark s t r e a k s o f less material u n t i l o f development.  structure-  the embryo sac e n t e r s a f a i r l y advanced  The c h a l a z a l megaspore i s f u n c t i o n a l  stage  ( F i g . 11G).  I t g r a d u a l l y i n c r e a s e s i n s i z e w i t h v a c u o l a t i o n u s u a l l y prominent toward the m i c r o p y l a r end.  The c h a l a z a l megaspore i s f u n c t i o n a l als<  i n S.  1959), j>. i n t e g r i f o l i a and  hieracifolia  (Beamish 1963,  (Wiggins  1967).  rufidula i n S.  Twin embryo sacs were not observed  f e r r u g i n e a but two r a d i c l e s were found emerging from a seed, a f a c t which suggests  germinating  the p o s s i b i l i t y o f twin embryo sacs.  Saxena (1963) r e p o r t e d twin embryo sacs i n S. Taylor  S.  diversifolia  and  (1965) i n the r e l a t e d genus Lithophragma. Megagametogenesis:-  ferruginea  The megagametophyte i n S a x i f r a g a  i s the monosporic, 8-nucleate, Polygonum type.  f u n c t i o n a l megaspore d i v i d e s m i t o t i c a l l y to g i v e r i s e to two  The nuclei  which l i e a t o p p o s i t e ends o f the d e v e l o p i n g embryo sac ( F i g . 11H). The next d i v i s i o n g i v e s r i s e to a p a i r o f n u c l e i a t each (Fig.  HI).  The growing embryo sac now  i n the m i d d l e developed The  has one or two  end  large vacuole  as a r e s u l t o f c o a l e s c e n c e o f s m a l l v a c u o l e s  t h i r d d i v i s i o n produces an 8-nucleate embryo sac w i t h a group o f  f o u r n u c l e i a t each end. apparatus,  The r e o r g a n i z a t i o n o f the n u c l e i i n t o  a n t i p o d a l s , and the p o l a r n u c l e i appears  q u i c k l y as the 8-nucleate appear a l i k e a t t h i s  embryo sac was  stage.  egg  to proceed  r a r e l y seen.  A l l the n u c l e  25 Gradually appearance. and  The  the egg apparatus assumes i t s c h a r a c t e r i s t i c egg  c e l l u s u a l l y takes up  a c q u i r e s a rounded o u t l i n e w i t h  the c e n t r a l p o s i t i o n  the n u c l e u s o r i e n t e d  the m i d d l e o f the embryo sac.  The  pear-shaped s y n e r g i d s  at  The  a n t i p o d a l s form an  the s i d e s o f the egg  cell.  t r i o a t the c h a l a z a l end.  The  towards are  located  inconspicuous  p o l a r n u c l e i f u s e to form the f u s i o n  n u c l e u s which l i e s i n the m i d d l e somewhat towards the c h a l a z a l o f the embryo sac. similarity  The mature embryo sac i s 7-nucleate and  o f the n u c l e i i s no  antipodals are short l i v e d , fertilized. reported  longer m a i n t a i n e d  p e r s i s t i n g only u n t i l  the  ( F i g . 11J). the egg^  The  is  Secondary m u l t i p l i c a t i o n o f the a n t i p o d a l s has  i n jS. d i v e r s i f o l i a  (Davis  1966)  but was  end  been  not observed i n  S.  ferruginea. Fertilization:synergid  is visible  A f t e r f e r t i l i z a t i o n of the egg,  ( F i g . 12B).  The  only  d i s a p p e a r a n c e o f the  one  other  s y n e r g i d appears to c o i n c i d e w i t h the e n t r y o f the p o l l e n tube i n t o the embryo sac. at  least u n t i l  S.  integrifolia  The  r e m a i n i n g s y n e r g i d p e r s i s t s f o r a longer  the proembryo i s 3 - c e l l e d . (Beamish 1963)  period,  U n l i k e the s y n e r g i d  the s y n e r g i d o f S.  of  f e r r u g i n e a has  not  been observed to d i v i d e . Endosperm:-After double f e r t i l i z a t i o n has been e f f e c t e d (Fig.  12A),  (Fig.  12B).  the primary endosperm n u c l e u s d i v i d e s b e f o r e The  f i r s t d i v i s i o n i s t r a n s v e r s e and  between the c h a l a z a l n u c l e u s and  a  the  partitionndevelops  the l a r g e r m i c r o p y l a r  nucleus.  c h a l a z a l n u c l e u s produces the s o - c a l l e d b a s a l endosperm by free - nuclear  divisions.  The m i c r o p y l a r  zygote  The  repeated  nucleus also d i v i d e s  and  26  r e d i v i d e s to form a m u l t i - n u c l e a t e d endosperm complex o f f r e e n u c l e i around the d e v e l o p i n g embryo.  A d e v e l o p i n g seed n e a r l y  330  microns broad had about 455 f r e e n u c l e i i n the endosperm complex. At  the advanced stages of embryonic development a d e f i n i t e  o c c u r s i n the endosperm ( F i g .  segmentation  12G,H). Thus the endosperm o f S.  f e r r u g i n e a i s o f the h e l o b i a l type which seems to be the g e n e r a l type i n the genus as r e p o r t e d by Samuelsson (1913), Gaumann (1918), Dahlgren (1930), Beamish and L i n (1965), Beamish (1967) and o t h e r s . Wiggins (1959) c o n s i d e r s the endosperm to be n u c l e a t e from the b e g i n n i n g i n S.  hieracifolia.  Embryogenesis:- A f t e r p o l l i n a t i o n the egg i s f e r t i l i z e d about 24 hours.  in  The p o l l e n tubes were f r e q u e n t l y seen i n the m i c r o -  p y l e , thus the p o l l e n tube e n t r y i s assumed to be g e n e r a l l y by t h a t route.  I n the c o n t r o l l e d p o l l i n a t i o n experiment the embryo was  c e l i e d by48 hours and 3 - c e l l e d by 72 hours a f t e r p o l l i n a t i o n .  The  z y g o t e g i v e s r i s e u l t i m a t e l y to a 4- t o 8- c e l l e d proembryo by t r a n s v e r s e d i v i s i o n s  (Figs.  the  c u n e i f o l i a t h a t the embryonal c e l l  proembryo i s o n l y 3 - c e l l e d , however  d u r i n g development.  I n JS.  integrifolia  vertical  The embryonal c e l l  sometimes d i v i d e s a t the 3 - c e l l e d stage o f the proembryo. (1950) noted i n S.  usually  12 C-E) a l t h o u g h o c c a s i o n a l l y  i n t e r c a l a r y d i v i s i o n s have a l s o been observed.  2-  Lebegue  d i v i d e s when  the proembryo becomes 8 - c e l l e d (Beamish and L i n 1965) the  proembryo may become 7 - c e l l e d b e f o r e d i v i s i o n of the embryonal c e l l . The proembryo i s 6 - c e l l e d i n S_. r u f i d u l a when the embryonal c e l l  divides divide.  (Beamish 1967). In S.  granulata  The b a s a l c e l l  i n S.  (Soueges 1936)  the b a s a l c e l l  d i v i s i o n s t o form 4 to 8  f e r r u g i n e a does not undergoes  cells.  The t e r m i n a l c e l l o f the proembryo  c o n s t i t u t e s the  embryonal c e l l which f i r s t d i v i d e s v e r t i c a l l y second d i v i s i o n i s t r a n s v e r s e  ( F i g . 12F).  to form a quadrant.  The  Another d i v i s i o n  makes i t an o c t a n t and g r a d u a l l y a g l o b u l a r mass o f c e l l s i s formed by repeated  d i v i s i o n s ( F i g . 12G).  embryo becomes h e a r t cotyledonary, (Fig.  121).  As development proceeds the  - shaped i n o u t l i n e ( F i g . 12H) and l a t e r  hypocotylary  the  and r a d i c l e r e g i o n s become d i s t i n g u i s h a b l e  The mature embryo l i e s  by the 4- to 5- l a y e r e d endosperm  s t r a i g h t i n the seed surrounded ( F i g . 12J).  The integument  d e v e l o p s 10 to 15 i r r e g u l a r rows o f t u b e r c l e s as e p i d e r m a l outgrowths. The seed coat  i s c h o c o l a t e brown a t m a t u r i t y .  28 ANATOMICAL STUDIES An o u t s t a n d i n g  study o f the f a m i l y S a x i f r a g a c e a e  c a r r i e d o u t by Thouvenin (1890).  was  He s t r e s s e d the importance o f  a n a t o m i c a l f e a t u r e s o f r o o t , stem and l e a f i n c l a s s i f i c a t i o n . order  t o f i n d o u t whether there a r e s i g n i f i c a n t a n a t o m i c a l  ences among p o p u l a t i o n s  of Saxifraga  ferruginea, material  In  differfrom  Kodiak, Juneau, P r i n c e R u p e r t , A l l i f o r d Bay, Vancouver and M a r i o n County was examined. Methods:rhizomes, flower  Various  developmental stages o f r o o t s , peduncles,  buds, v e g e t a t i v e  were used f o r t h i s i n v e s t i g a t i o n . formalin-aceto-alcohol  shoot a p i c e s and young i n f l o r e s c e n c e s The p l a n t m a t e r i a l was f i x e d i n  i n the f i e l d b u t was l a t e r washed, c u t i n t o  c o n v e n i e n t p i e c e s , a s p i r a t e d and put i n t o c l e a n f i x a t i v e i n the laboratory before dehydration  being  series.  passed through a t e r t i a r y b u t y l a l c o h o l (TBA)  The r o o t s , leaves and young  became t r a n s l u s c e n t  inflorescences  i n 100% TBA i n 24 to 48 hours.  rhizomes and shoot a p i c e s r e q u i r e d  roots,  The  i n F i s h e r tissuemat  l e a v e s and f l o w e r s were s e c t i o n e d  microns; flowering  stems,  72 to 96 hours i n 100% TBA and  a c c o r d i n g l y needed extended time i n p a r a f f i n and wax The m a t e r i a l was embedded  The f l o w e r i n g  (m. p.  infiltration. 53 - 55°C).  The  t r a n s v e r s l y a t 10 t o 12  stems, rhizomes and shoot a p i c e s a t 15 t o 20 microns.  s e c t i o n s were s t a i n e d w i t h a s a f r a n i n - f a s t green combination.  Numerous free-hand s e c t i o n s o f the m a t e r i a l were a l s o examined. For (Gourley boiled  studying  the v e n a t i o n  1930) was used f o r v i t a l  i n basal staining.  leaves, basic  fuchsin  The l e a v e s were then  i n s l i g h t l y a l k a l i n i z e d 85% a l c o h o l to remove the c h l o r o p h y l l  29 and  passed through 95% and a b s o l u t e  ethyl alcohol before being  i n methyl s a l i c y l a t e f o r c l e a r i n g (Johansen 1940). had  r e d v e i n s b u t were hard and b r i t t l e .  leaves  The l e a f p e t i o l e s  a t the base, t h e m i d l e n g t h and the upper r e g i o n where  the lamina broadens o u t .  Flower buds were c l e a r e d i n l a c t i c a c i d  (Simpson 1929) to r e v e a l v a s c u l a t u r e .  O b s e r v a t i o n s made on whole buds  were supplemented by examination o f s e r i a l and  The c l e a r e d  A few days i n g l y c e r i n e made  them p l i a b l e enough to handle f o r o b s e r v a t i o n . were s e c t i o n e d  treated  sections cut l o n g i t u d i n a l l y  transversely. O b s e r v a t i o n s and r e l e v a n t  literature  Some i n t e r e s t i n g a n a t o m i c a l f e a t u r e s were found i n t h i s but  the s i x p o p u l a t i o n s  examined were e s s e n t i a l l y i d e n t i c a l .  species  The  f o l l o w i n g d e s c r i p t i o n , t h e r e f o r e , o u t l i n e s the c h a r a c t e r i s t i c s o f the species. The  Root:- T r a n s v e r s e s e c t i o n s o f young r o o t s , 0.5 mm. t h i c k ,  show t h e e p i d e r m i s c o m p r i s i n g  a s i n g l e layer of i r r e g u l a r c e l l s .  c o r t e x i s parenchymatous and 5 to 7 l a y e r s deep. most l a y e r o f the c o r t e x cortical cells  The  C e l l s o f the i n n e r -  ( t h e endodermis) a r e much s m a l l e r  than  ( F i g . 13F). The p e r i c y c l e i s 2- t o 3-layered.  other  The  xylem i s t r i a r c h o r t e t r a r c h , each primary xylem bundle c o n s i s t i n g o f 3 to 5 t r a c h e a r y  elements.  Phloem a l t e r n a t e s w i t h t h e xylem elements.  An u n d i f f e r e n t i a t e d parenchymatous zone l i e s i n the middle o f the root. Soon a f t e r seed germination,when s e e d l i n g s  are: e s t a b l i s h e d ,  a d v e n t i t i o u s r o o t s a r e i n i t i a t e d a t the base o f the r o s e t t e . r o o t s c o n s t i t u t e the main r o o t  system o f the p l a n t .  These  The a d v e n t i t i o u s  30 r o o t s v a r y from 0.3  mm.to 1.0 mm. or up to 1.5 mm. i n t h i c k n e s s .  o r d e r to study secondary examined.  growth, r o o t s o f d i f f e r e n t t h i c k n e s s e s were  In young r o o t s the f a s c i c u l a r cambium d i f f e r e n t i a t e s  i n s i d e the phloem and, continuous.  extending o u t s i d e the protoxylem,  During e a r l y stages o f secondary  xylem pomfs u n t i l  the cambium becomes c i r c u l a r  phloem i s produced  than secondary  becomes  growth, xylem i s  a c t i v e l y added o p p o s i t e the phloem but v e r y s l o w l y over  secondary  In  the p r o t o -  ( F i g . 13G).  xylem.  Much l e s s  In o l d e r r o o t s cork  cambium d i f f e r e n t i a t e s i n the c o r t e x and g i v e s r i s e to c o r k towards the p e r i p h e r y ( F i g . 13H). as the c o r k l a y e r develops.  tissue  The outer t i s s u e s a r e sloughed o f f  Secondary xylem and c o r k l a y e r s  constitute  the major p a r t o f o l d r o o t s . The Peduncle:c e l l s with a c u t i c l e .  The e p i d e r m i s c o n s i s t s o f a s i n g l e l a y e r of I t i s i n t e r r u p t e d by the stomata and a l s o  gland-tipped m u l t i c e l l u l a r hairs.  The c o r t e x has 3 to 5 l a y e r s o f  chlorenchymatous c e l l s w i t h i n t e r c e l l u l a r  spaces.  I n s i d e the c h l o r e n -  chymatous t i s s u e a r e 1 or 2 l a y e r s o f parenchyma. band of sclerenchyma been r e f e r r e d  e n c l o s e s the s t e l e ( F i g s .  by Thouvenin (1890) and M e t c a l f e and C h a l k demonstration  support arranged  13A,C).  T h i s band  These f i b r e s make up  the i n f l o r e s c e n c e .  (1950).  Lacking casparian  the main mechanical  The v a s c u l a r bundles  the p i t h r a y s extending  Saxifragaceae  band p e r i v a s c u l a r f i b r e s t i s s u e to  are c o l l a t e r a l  i n a r i n g around the parenchymatous p i t h .  a r e separated by  has  of the endodermis, i t would  be a p p r o p r i a t e to c a l l the sclerenchymatous (Esau 1965).  A 3- to 4 - l a y e r e d  to as the p e r i c y c l e i n o t h e r members o f the  s t r i p s or a t l e a s t without  bears  Vascular  towards the c o r t e x .  and  bundles Some o f  the p i t h c e l l s become d i s o r g a n i z e d thus making the p i t h The  Rhizome:- The epidermis  i s s i n g l e l a y e r e d and the c e l l s  a r e s m a l l when compared t o those o f the c o r t e x .  The c o r t e x i s  made up o f 15 to 30 l a y e r s o f parenchymatous c e l l s number o f l a y e r s d e t e r m i n i n g  lacunate.  ( F i g . 13B), the  the t h i c k n e s s o f the rhizome.  A d v e n t i t i o u s r o o t s grow on the young branches o f the rhizomes and can be seen t r a v e r s i n g the c o r t e x a r e observed i n the t r a n s v e r s e rhizome ( F i g . 14D). spicuous  (Figs.  14 A,B).  traces  s e c t i o n s o f the o l d e r p a r t s o f the  The innermost l a y e r o f the c o r t e x i s a con-  t r u e endodermis ( F i g s 14C,E) as evidenced  of c a s p a r i a n s t r i p s .  by the presence  The v a s c u l a r bundles i n the young branches o f  the rhizome a r e observed as two broad bands.  As t h e rhizome i n c r e a s e s  i n t h i c k n e s s , the two v a s c u l a r bundles a p p a r e n t l y four  The l e a f  d i v i d e to form  ( F i g . 14A). With f u r t h e r i n c r e a s e i n t h i c k n e s s the v a s c u l a r  bundles b r a n c h to form as many as 8 ( F i g . 14D). The L e a f : - Lamina: epidermis along  A single layer of c e l l s  comprisesthe  which i s i n t e r r u p t e d by stomata on both s u r f a c e s except  the m i d r i b .  H a i r s a l s o develop from the epidermis.  The h a i r s  a r e m u l t i c e l l u l a r and m u l t i s e r i a t e , w i t h a m u l t i c e l l u l a r head. h a i r a r i s e s from 3 to 4 epidermal  cells.  Most epidermal  cells  Each possess  wavy w a l l s , b u t those a t the m i d r i b have s t r a i g h t w a l l s ; the c e l l s a r e arranged  i n l o n g i t u d i n a l rows, as viewed i n the whole mounts.  The m e s o p h y l l c o n s i s t s o f 3 to 7 l a y e r s o f p a l i s a d e parenchyma and 4 to 6 l a y e r s o f very  spongy parenchyma ( F i g . 13E). The v a s c u l a r  b u n d l e s o f the l e a f a r e n o t surrounded by a bundle sheath  o r endodermi  32 Druses were observed i n the l e a v e s , s e p a l s and i n Saxifraga ferruginea.  M e t c a l f e and  druses to be r a r e i n S a x i f r a g a .  C h a l k (1950) c o n s i d e r e d  However Thouvenin (1890) r e p o r t e d  c a l c i u m o x a l a t e c r y s t a l s as druses i n S. and  as s o l i t a r y n e e d l e s  i n S.  ovary w a l l s  f o r tune i  nipponica.  and j>.  sendaica  These t h r e e s p e c i e s  belong  to s e c t . D i p t e r a Borkhausen. Petiole: s t r u c t u r e but  p e t i o l e i n S.  The  f e r r u g i n e a i s not a  distinct  i s formed from the e x t e n s i o n o f the m i d r i b  w i t h the narrow a t t e n u a t e d  base of the lamina  together  ( F i g . 13D).  toward the lower p o r t i o n o f the narrow p e t i o l e the lamina d i f f e r e n t i a t i o n o f the m e s o p h y l l i n t o p a l i s a d e and In S.  ferruginea only a s i n g l e trace enters  However, lacks  spongy parenchyma.  the l e a f .  Watari  studied p e t i o l a r v a s c u l a r i z a t i o n i n various Saxifraga species r e p o r t e d 3 to 7 l e a f t r a c e s e n t e r i n g the l e a f i n S. s i n g l e l e a f t r a c e i n S. The Flower:- The illustrated the f l o w e r  c e r n u a and  and  f o r t u n e i , but  a  S. c h e r l e r i o i d e s .  s e c t i o n o f a bud  ( F i g . 16G).  Normally  t e n v a s c u l a r b u n d l e s which t r a v e r s e the hypanthium.  F i v e b u n d l e s go to the s e p a l s to form the midveins and to the p e t a l s . The  (1939)  arrangement of the p a r t s o f a flower i s  i n a transverse s t a l k has  the  five  proceed  l a t e r a l v e i n s o f the s e p a l s a r e c o n t r i b u t e d  branches a r i s i n g from the two n e i g h b o u r i n g  p e t a l bundles ( F i g .  by 15A).  Where a p e t a l bundle g i v e s o f f branches to form the l a t e r a l v e i n s o f neighbouring t h i r d branch.  s e p a l s , a stamen o p p o s i t e  the p e t a l a l s o r e c e i v e s  The m i d v e i n o f the p e t a l g i v e s o f f two  i n t o the p e t a l lamina.  a  l a t e r a l branches  Sometimes the l a t e r a l s a l s o produce s h o r t  branches i n broad p e t a l s ( F i g s . 8-1,4).  The  gynoecium has  vascular  33  b u n d l e s i n the p l a c e n t a and o v a r y w a l l . the p l a c e n t a enter  The v a s c u l a r bundles o f  towards the base o f the c a r p e l s i n the c e n t r e ,  a r i s i n g from the v a s c u l a r c y l i n d e r o f the p e d i c e l , each f u r t h e r d i v i d i n g to g i v e two branches to the p l a c e n t a l t i s s u e j u s t below where the f u s i o n o f the two c a r p e l s ends ( F i g s . 15 A,B).  The  w a l l s o f the c a r p e l s r e c e i v e b u n d l e s from each o f t h e v e i n s to s e p a l s and p e t a l s .  A f u r t h e r branching  going  o f some t r a c e s o c c u r s  i n the  w a l l s o f the c a r p e l s . The  Shoot Apex and I n f l o r e s c e n c e : - O b s e r v a t i o n s were made on  numerous l o n g i t u d i n a l s e c t i o n s o f shoot apex and many developmental stages cept;  !  o f young i n f l o r e s c e n c e s . A c c o r d i n g  to the " t u n i c a - c o r p u s "  con-  o f a p i c a l o r g a n i z a t i o n (Esau 1965), a s i n g l e p e r i p h e r a l l a y e r in  S. f e r r u g i n e a comprises the t u n i c a . observed i n t h i s l a y e r .  Only a n t i c l i n a l d i v i s i o n s were  However, b o t h a n t i c l i n a l and p e r i c l i n a l  d i v i s i o n s were viewed i n the c o r p u s , the zone o f c e l l s under the tunica.  The same a p i c a l o r g a n i z a t i o n was m a i n t a i n e d d u r i n g the  t r a n s i t i o n to the r e p r o d u c t i v e s t r a t i f i e d configurations.  stage except t h a t the corpus developed  The young i n f l o r e s c e n c e d i d n o t show  d i f f e r e n t i a t i o n o f p l a n t l e t s i n e a r l y stages  ( F i g s . 16A to D).  However i n the l a t e r p a r t o f the season the replacement o f flower buds by p l a n t l e t s takes p l a c e  ( F i g s . 16E,F).  The p l a n t l e t  formation  i s a t the expense o f the flower buds and the extent o f replacement o f flower buds by the v e g e t a t i v e p l a n t l e t s i s v a r i a b l e i n d i f f e r e n t populations  as w e l l as i n d i f f e r e n t years w i t h i n the same p o p u l a t i o n s .  34 CHROMATOGRAPHIC STUDIES Methods:F r e s h l e a v e s were c o l l e c t e d from Kodiak, Juneau, P r i n c e Rupert, A l l i f o r d Bay, Vancouver and C r a t e r Lake p o p u l a t i o n s to p r e p a r e e x t r a c t s f o r s t u d y i n g v a r i a b i l i t y phenolic acids.  i n the p a t t e r n o f  The p l a n t s were i n a p p r o x i m a t e l y the same  stage o f development, most o f them i n bloom. The t e c h n i q u e o f Ibrahim and Towers (1960) f o r 2 - d i r e c t i o n al  paper chromatography  chopped  used.  Leaves from each p o p u l a t i o n were  i n a b l e n d e r and the e x t r a c t o b t a i n e d i n 807 e t h a n o l .  The  o  e x t r a c t was  evaporated to dryness i n a r o t a r y e v a p o r a t o r .  d r i e d e x t r a c t was filtered  was  taken up i n 80 to 100 ml. o f hot water  through ' c e l i t e ' - a f i l t e r a i d .  acidified  and  The f i l t r a t e was h y d r o l y z e d  w i t h NaOH (pH 10) a t room temperature f o r 24 hours. the ^ e x t r a c t was  The  w i t h HCl to pH 4.  After  hydrolysis  The h y d r o l y z a t e  was  e x t r a c t e d w i t h ether f o r 12 hours and the e t h e r e x t r a c t evaporated to dryness.  The d r i e d e x t r a c t was  s p o t t e d i n Whatman No. from the margins. paper For  d i s s o l v e d i n a l i t t l e e t h a n o l and  1. chromatography  The f i r s t  paper a t a p o i n t  equidistant  s o l v e n t used a l o n g the g r a i n o f the  ( t h e l o n g d i r e c t i o n ) was benzene:  a c e t i c a c i d : water (10:7:3).  the second d i r e c t i o n 27» f o r m i c a c i d was  used.  The f i r s t  solvent  r e q u i r e d 6 to 7 hours to r e a c h w i t h i n an i n c h o f the margin o f the chromatographic paper.  The second s o l v e n t r e a c h e d the d e s i r e d  p o s i t i o n i n 3 to 3.5 hours depending on room temperature.  The  dried  chromatograms were exposed t o ammonia vapour and then examined  under  UV  light  (3660&).  The f l u o r e s c e n t s p o t s were marked.  The  chromato-  35 grams were sprayed  u n i f o r m l y w i t h d i a z o t i z e d p - n i t r o a n i l i n e and  then w i t h IN NaOH.  The c o l o u r e d spots were r e c o r d e d  Observations  and marked.  and r e l e v a n t l i t e r a t u r e :  The chromatograms f o r t h e s i x p o p u l a t i o n s turned o u t to be a l i k e  ( F i g . 17). The f l u o r e s c e n c e and c o l o u r r e a c t i o n s o f t h e  spots a r e g i v e n i n T a b l e VT.  T a b l e VI UV  f l u o r e s c e n c e and c o l o u r r e a c t i o n s o f t h e p h e n o l i c compounds i n l e a v e s o f S. f e r r u g i n e a .  found UV/NH  p-nitroaniline  3  (3660 &)  Compound  No. o f spots  Blue  B l u i s h green  2  Ferulic  Blue  Bluish  2  p-Coumaric a c i d  Pink  1  p-Hydroxy b e n z o i c  L i g h t brownish yellow  2  Caffeic  Brownish green  1  Protocatechuic  Blue  -  purple  B i l l e k and K i n d l e  acid  acid  and  2,hydroxy phenyl  considered  acid  (1962) sampled nine genera o f the f a m i l y  S a x i f r a g a c e a e b u t S a x i f r a g a was n o t i n c l u d e d i n t h e i r recorded  acid  study.  They  a c e t i c a c i d o n l y i n t h e genus A s t i l b e  i t to be diagno s t i c f o r t h i s genus a t that time u n t i l  a d d i t i o n a l evidence  proved i t otherwise.  The p r e l i m i n a r y  identifi-  c a t i o n based on UV f l u o r e s c e n c e and c o l o u r r e a c t i o n s shows t h a t S a x i f r a g a f e r r u g i n e a l a c k s 2,hydroxy phenyl therefore,  still  a c e t i c a c i d which i s ,  d i a g n o s t i c f o r the genus A s t i l b e .  36 CROSSABILITY STUDIES Since the i n f o r m a t i o n d e r i v e d from d i f f e r e n t study i n d i c a t e s a l a r g e amount o f u n c o r r e l a t e d between and w i t h i n p o p u l a t i o n s over  lines of  variability  the range o f t h i s  species  complex, an attempt was made to f i n d o u t whether b a r r i e r s to gene f l o w e x i s t between p o p u l a t i o n s . Methods:C r o s s e s were made i n v o l v i n g M a r i o n County, Washington, Vancouver, A l l i f o r d  Bay,and P r i n c e Rupert p o p u l a t i o n s  cultivated  a t the U n i v e r s i t y o f B r i t i s h Columbia ( F i g . 18). Due to the shortage  o f f l o w e r s the Washington p o p u l a t i o n was used as a male  parent only. and  The K o d i a k p l a n t s d i d n o t bloom i n the greenhouse  thus c o u l d n o t be used i n c r o s s a b i l i t y experiments.  a v a i l a b l e , 3 t o 5 p l a n t s were used from each p o p u l a t i o n .  Where Control  c r o s s e s were a l s o made w i t h i n a l l b u t the Washington p o p u l a t i o n . The  female p a r e n t s were emasculated and bagged.  The f l o w e r s were  p o l l i n a t e d when the stigmas s e c r e t e d a d r o p l e t a t the s u r f a c e . At m a t u r i t y  the d e h i s c i n g f r u i t s were detached and the seeds  collected. Observations:The  i n t r a p o p u l a t i o n c r o s s e s produced seed  freely.  The  r e s u l t s o f the i n t e r p o p u l a t i o n 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 .  37 Table VII Interpopulation crosses  i n Saxifraga  # Flowers pollinated  Crossed populations DIPLOID  ferruginea.  X  DIPLOID  X  M a r i o n County  37  M a r i o n County  X  A l l i f o r d Bay  28  DIPLOID  X  POLYPLOID  X  Vancouver  118  M a r i o n County  X  Vancouver  16  M a r i o n County  X  Washington  22  Alliford  X  P r i n c e Rupert  87  Alliford  Alliford  Bay  Bay  Bay  POLYPLOID X  + +  11 seeds !  DIPLOID  Vancouver  X  Alliford  Bay  152  P r i n c e Rupert  X  Alliford  Bay  82  POLYPLOID X  Seed s e t *  POLYPLOID  Vancouver  X  P r i n c e Rupert  Vancouver  X  Washington  P r i n c e Rupert  X  Vancouver  * + Reduced seed s e t as compared as c o n t r o l s .  136  +  12  +  167  +  to i n t r a p o p u l a t i o n c r o s s e s  serving  - No seed s e t .  The p o l y p l o i d p o p u l a t i o n s  were found to be c r o s s c o m p a t i b l e ,  w i t h a reduced seed s e t as compared was o b t a i n e d  The same r e s u l t  i n the r e c i p r o c a l c r o s s e s between the d i p l o i d  Bay and M a r i o n County p o p u l a t i o n s . d i p l o i d s and p o l y p l o i d s f a i l e d Alliford  t o the c o n t r o l s .  The i n t e r p o p u l a t i o n c r o s s e s among  t o s e t seed except one c r o s s between  Bay and P r i n c e Rupert, from which 11 seeds were  T h i s c r o s s would be c y t o l o g i c a l l y of chromosomes a t m e i o s i s .  Alliford  obtained.  i n t e r e s t i n g f o r studying  the p a i r i n g  However, t h i s c o u l d n o t be s t u d i e d f u r t h e r  w i t h i n the p e r i o d o f the p r e s e n t i n v e s t i g a t i o n .  38 GENERAL DISCUSSION The d e s c r i p t i o n o f S a x i f r a g a f e r r u g i n e a was w r i t t e n byGraham (1829) based on p l a n t s grown a t the Royal B o t a n i c Edinburgh.  Garden  No mention was made o f the type l o c a l i t y but a c c o r d i n g  to S i r John R i c h a r d s o n (1829) the seeds were c o l l e c t e d i n the Rocky Mountains by Thomas Drummo»clwho  accompanied  him as A s s i s t a n t  N a t u r a l i s t on S i r John F r a n k l i n ' s second n o r t h l a n d Drummond ( I n Hooker 1830)  expedition.  d e s c r i b e s h i s j o u r n e y from J a s p e r up  the  Athabaska R i v e r , through the Athabaska Pass i n the Rocky Mountains and down to Boat Encampment on the Columbia R i v e r . i o n s on the r o c k s around a s m a l l l a k e c a l l e d near Athabaska Pass.  He made c o l l e c t -  'Committee Punch Bowl'  The type l o c a l i t y o f j5. f e r r u g i n e a then  appears to be the r o c k s near the l a k e 'Committee Punch Bowl' a t Athabaska Pass, which i s about 52° 20*  l a t . 118° 10'long. ( F i g . 1).  I n the y e a r s s i n c e Drummond's c o l l e c t i o n was made from the type l o c a l i t y , our knowledge o f the d i s t r i b u t i o n o f t h i s has expanded.  species  We now know t h a t i t ranges from n o r t h w e s t e r n  C a l i f o r n i a to A l a s k a a l o n g the c o a s t and the Rocky Mountains  and  t h a t i t grows from sea l e v e l to a l p i n e r e g i o n s . A c c o r d i n g to C l a u s e n , Keck and H i e s e y (1941) w i d e l y d i s t r i b u t e d s p e c i e s tend to d i f f e r e n t i a t e i n t o b i o t y p e s and r a c e s under environmental c o n d i t i o n s .  different  S a x i f r a g a f e r r u g i n e a i s indeed a widespread  s p e c i e s which c o u l d w e l l be expected to comprise a number o f subspecific  taxa.  To r e c o g n i z e such taxa p o p u l a t i o n s should be  examined  throughout the range o f the s p e c i e s , e s p e c i a l l y when e c o l o g i c a l , g e o g r a p h i c a l , and c l i m a t i c f a c t o r s as w e l l as g l a c i a t i o n have  39 offered opportunity  f o r e v o l u t i o n a r y change.  A p r e l i m i n a r y c y t o l o g i c a l examination o f S. f e r r u g i n e a (Beamish 1961)  i n d i c a t e d the chromosome number to be  i n a few p o p u l a t i o n s survey  around Vancouver, B r i t i s h Columbia.  o f many p o p u l a t i o n s  the presence o f 10-paired Randhawa and  Beamish 1968,  r e n c e o f d i p l o i d and  19 p a i r s  throughout i t s range has now as w e l l as 19-paired  races  r e c o g n i t i o n i s not uncommon.  revealed  (Packer  T a y l o r and M u l l i g a n 1968).  p o l y p l o i d r a c e s i n one  A  The  1968,  occur-  s p e c i e s without  formal  I n f r a s p e c i f i c chromosome r a c e s have  been r e p o r t e d i n S a x i f r a g a h i e r a c i f o l i a ,  S. h i r c u l u s , S.  o p p o s i t i f o l i a e t c . ( F l o v i k 1940). M e i o t i c i r r e g u l a r i t i e s and c o u l d be  expected to d i f f e r w i t h  pollen f e r t i l i t y  i n S. f e r r u g i n e a  l e v e l o f p o l y p l o i d y and  in fact,  the p o l y p l o i d s appear to be g e n e r a l l y more i r r e g u l a r than diploids.  However, the change i n p o l l e n f e r t i l i t y  Rupert p l a n t s from 1967  to 1968  v i a b i l i t y i n S. f e r r u g i n e a was  of P r i n c e  suggests an environmental  which b l u r s g e n e t i c d i f f e r e n c e s .  the  effect  Similar fluctuation i n pollen  r e p o r t e d by Beamish (1961).  (1937) working w i t h T r a d e s c a n t i a designed  temperature  fluctuation  experiments which demonstrated that p l a n t s t r a n s f e r r e d from c o l d room (8°C)  the  to a hot chamber (38°C) e x h i b i t e d a b n o r m a l i t i e s  n e a r l y 507„ of the p o l l e n - m o t h e r - c e l l s a week a f t e r abnormalities continued months.  Sax  treatment.  to appear w i t h a low frequency  temperature on  meiosis.  The  for several  S i n c e Sax's experiments, many workers have demonstrated  the adverse e f f e c t Gf  in  40 The  d i s t r i b u t i o n p a t t e r n o f d i p l o i d and  p o l y p l o i d populations  suggests a s i g n i f i c a n t c o r r e l a t i o n w i t h the problem of r e f u g i a i n the C o r d i l l e r a n G l a c i a l Complex and p o s t - g l a c i a l times. Observations  plant migration routes i n  This i s discussed i n d e t a i l  later.  on herbarium specimens of S>. f e r r u g i n e a , r e p r e -  s e n t i n g i t s e n t i r e range, i n d i c a t e a tremendous m o r p h o l o g i c a l ability.  A l l c l a s s i f i c a t i o n s have been based l a r g e l y on  variation. carefully  The  characters u t i l i z e d  to a s s e s s  The  shape and  their  even w i t h i n a s i n g l e p o p u l a t i o n bear 3 to 15 t e e t h which can  anywhere from 1 to 3 mm.in l e n g t h . inconsistent w i t h i n populations  7).  a r e d i s c u s s e d below:  s i z e of the b a s a l l e a v e s i s extremely v a r i a b l e  Furthermore the l e a v e s may  v a r y i n s i z e and  this  i n p r e v i o u s keys were a p p r a i s e d  taxonomic v a l u e and  among d i f f e r e n t p o p u l a t i o n s and 4-6).  vari-  The  (Figs. be  s i z e of stomata i s l i k e w i s e  (Table V).  The b r a c t e a l l e a v e s a l s o  number o f t e e t h even w i t h i n a s i n g l e p o p u l a t i o n ( F i g .  Most p o p u l a t i o n s have scanty pubescence on the l e a v e s , e s p e c i a l l y on  the a b a x i a l s u r f a c e .  However, A l l i f o r d  Bay  and  Onward P o i n t  have a h i g h d e n s i t y of pubescence on b o t h s u r f a c e s of the These two  populations d i f f e r  the A l l i f o r d  Bay  from each other  p o p u l a t i o n having  leaves.  i n the l e n g t h o f h a i r s ,  distinctly  Onward P o i n t p o p u l a t i o n (Table V).  populations  longer h a i r s than the  Even d e n s i t y o f pubescence i s not  c h a r a c t e r i s t i c of a l l Queen C h a r l o t t e I s l a n d p o p u l a t i o n s , some a t h i g h e r a l t i t u d e s having apparently glabrous  leaves  ( T a y l o r 1968).  T y p i c a l l y , f l o w e r s have f i v e p e t a l s , the upper three w i t h two  y e l l o w spots and  the o t h e r s  lanceolate  without  sagittate  41 spots.  However, f l o w e r s o f t e n possess  a l l bear spots. possesses  One  o n l y s a g i t t a t e p e t a l s and  p o p u l a t i o n on the Queen C h a r l o t t e I s l a n d s  ovate p e t a l s without  spots  ( F i g s . 8-19,20).  The  size  of f l o w e r s i s h i g h l y v a r i a b l e , some p o p u l a t i o n s on the c o a s t a l i s l a n d s and  the Queen C h a r l o t t e I s l a n d s b e a r i n g r a t h e r l a r g e  f l o w e r s but  the n e i g h b o u r i n g  different observed a r e not _t*  sizes  ( F i g s . 8-1  i n the shape and  p o p u l a t i o n s i n d i c a t e an a r r a y o f  to 30).  I n s p i t e o f the  s i z e o f f l o w e r s , the  variability  characteristics  s t a b l e enough to be used f o r d e l i m i t a t i o n o f f e r r u g i n e a reproduces  t i v e mechanism i n t h a t few  by an obvious  and  taxa.  efficient  vegeta-  to a l l the f l o w e r s i n an i n f l o r e s c e n c e  a r e f r e q u e n t l y r e p l a c e d by p l a n t l e t s .  Although  plantlet  formation  i n the i n f l o r e s c e n c e i s an a p o m i c t i c phenomenon, i t has not  inter-  f e r e d w i t h the p o t e n t i a l i t y f o r s e x u a l r e p r o d u c t i o n , because where f l o w e r s a r e produced they s e t seed  freely.  S i n c e seed  i n a l l p o p u l a t i o n s i s the r e s u l t o f the s e x u a l p r o c e s s , appear t h a t p l a n t l e t f o r m a t i o n c o u l d be to the p l a n t . set 50 to 150  i t would  sometimes disadvantageous  One : p l a n t l e t can r e p l a c e a f l o w e r which c o u l d have seeds.  However, i f u n f a v o u r a b l e  environmental  d i t i o n s can reduce p o l l e n p r o d u c t i o n , t h e r e may p l a n t l e t p r o d u c t i o n i s an advantage.  be  con-  seasons when  The p l a n t l e t a l r e a d y has  passed the c r i t i c a l p e r i o d o f g e r m i n a t i o n , develop  production  by-  so t h a t one p l a n t l e t might  i n t o a p l a n t w h i l e hundreds o f seeds might never r e s u l t i n  seedlings. j>. f e r r u g i n e a a p p a r e n t l y developed problem o f s u r v i v a l , namely apomixis and  two  approaches to  the  sexual reproduction.  42 Apomixis enables  a b i o t y p e to i n c r e a s e i n numbers which may  most s u c c e s s f u l i n a g i v e n s e t of environmental However, when a f l u c t u a t i o n i n the environmental  conditions. c o n d i t i o n s would  o c c u r , the e n t i r e b i o t y p e w i l l be a d v e r s e l y a f f e c t e d .  Thus  apomixis p r o v i d e s a s h o r t - t e r m advantage to the b i o t y p e . r e p r o d u c t i o n , on the o t h e r hand, imparts v a r i a b i l i t y s e g r e g a t i o n and recombinations  be  which i s a long-term  Sexual  through advantage to  the whole s p e c i e s . C o n t r o l l e d growth chambers should be used to t e s t r e l a t i o n s h i p of environmental and  seed  set.  c o n d i t i o n s to p l a n t l e t  production  The p r o d u c t i o n o f p l a n t l e t s has o f t e n been used i n  c l a s s i f i c a t i o n o f t h i s s p e c i e s complex. on the p l a n t s r a i s e d from seed and p r o p o r t i o n o f p l a n t l e t s and r a t i o of p l a n t l e t s unstable  the  However, o b s e r v a t i o n s  c l o n e s have i n d i c a t e d t h a t  f l o w e r s v a r i e s from year  the  to year.  to; f l o w e r s , t h e r e f o r e , appears to be  The  an  character.  A comparative study of embryology among s i x p o p u l a t i o n s brought out i n t e r e s t i n g i n f o r m a t i o n but no i n seed f o r m a t i o n was  observed.  infraspecific  Seed p r o d u c t i o n i n t h i s s p e c i e s , as  d e s c r i b e d p r e v i o u s l y , i s by a normal s e x u a l p r o c e s s . d i f f e r s from v a r i o u s o t h e r  after  This  species  s p e c i e s of S a x i f r a g a i n s e v e r a l r e s p e c t s  i . e . number o f c e l l s o f the proembryo, behaviour and  variability  of s y n e r g i d s  the f e r t i l i z a t i o n o f the egg, number o f integuments  type o f endosperm.  I t d i f f e r s i n two  °f S a x i f r a g a r e p o r t e d so f a r : integument becomes deeply  first,  before and  d e t a i l s from a l l o t h e r  species  the innermost l a y e r o f  the  s t a i n e d at about the4-nucleate  stage of  43  embryo sac development. and  This layer collapses after  fertilization  i s l a t e r observed as a dark s t r e a k around the n u c e l l u s ; second,  the c e l l s o f the outermost l a y e r o f the n u c e l l u s b e g i n due  to e n l a r g e  to extreme v a c u o l a t i o n when the integumentary l a y e r i s d i f f e r e n -  tiating.  These c e l l s  as the seed matures. populations  lose their  i d e n t i t y and a r e a p p a r e n t l y  These f e a t u r e s , b e i n g c o n s t a n t  a r e o f no use i n d i f f e r e n t i a t i n g  absorbed  over the s i x  subgroups w i t h i n the  s p e c i e s b u t may be u s e f u l i n working out a f f i n i t i e s w i t h i n the genus S a x i f r a g a . Thouvenin (1890) s t r e s s e d the importance o f a n a t o m i c a l of r o o t , stem, and l e a f i n c l a s s i f i c a t i o n . characters i n a l l s i x populations  However,  M e t c a l f e and C h a l k (1950) c o n s i d e r e d t h e i r presence or absence probably  Druses  Since  druses to be r a r e i n S a x i f r a g a ,  c o u l d form a v a l u a b l e  i n the c l a s s i f i c a t i o n o f the genus. and  anatomical  follow a single pattern.  were observed i n the l e a v e s , s e p a l s and ovary w a l l s .  features  character  The anatomy o f the shoot apex  the young i n f l o r e s c e n c e i n d i c a t e s t h a t p l a n t l e t f o r m a t i o n  i s at  the expense o f the flower buds. Chromatographic study was c a r r i e d out to determine whether t h e r e was any c o r r e l a t i o n between m o r p h o l o g i c a l  and c y t o l o g i c a l  v a r i a b i l i t y and the p h e n o l i c a c i d p a t t e r n i n s i x p o p u l a t i o n s . However, the chromatograms f o r a l l the p o p u l a t i o n s were s i m i l a r . T h i s study  i s significant  i n r e c o r d i n g the absence o f 2, hydroxy  phenyl a c e t i c a c i d i n Saxifraga.  B i l l e k and K i n d l e  (1962) found  t h i s compound o n l y i n the genus A s t i l b e among many genera o f Saxifragaceae  they had examined.  2, Hydroxyphenyl a c e t i c a c i d  44 therefore, s t i l l  i s d i a g n o s t i c o f the genus A s t i l b e .  C r o s s e s were made between d i f f e r e n t p o p u l a t i o n s . p l o i d populations  are i n t e r f e r t i l e although  normal seed set was p o l y p l o i d and case o f one  observed.  The  poly-  a r e d u c t i o n from  The r e c i p r o c a l c r o s s e s between  d i p l o i d p o p u l a t i o n s d i d not  c r o s s which y i e l d e d 11 seeds.  s e t seed except i n the T h i s c r o s s should  be  s t u d i e d f u r t h e r to determine whether the embryos i n the seeds a r e h y b r i d t r i p l o i d s or whether the seeds developed due contamination  o r , by any  chance, a r e a p o m i c t i c .  M a r i o n County p o p u l a t i o n and  The  c r o s s between  the i s o l a t e d Queen C h a r l o t t e I s l a n d  p o p u l a t i o n , b o t h d i p l o i d , i s i n t e r e s t i n g f o r two should be  to p o l l e n  studied further: f i r s t ,  reasons  s i n c e the A l l i f o r d  Bay  and plants  r a r e l y produce p l a n t l e t s i n the i n f l o r e s c e n c e and M a r i o n County p l a n t s bear jnostly p l a n t l e t s , the g e n e t i c s t a b i l i t y o f p l a n t l e t flower production generations;  i n the i n f l o r e s c e n c e can be  s t u d i e d i n F^  second, the c o m p a t i b i l i t y of the two  between d i p l o i d and  species.  and  d i p l o i d genomes  suggests t h a t b e f o r e the P l e i s t o c e n e G l a c i a t i o n t h e r e may a s i n g l e widespread d i p l o i d  and  have been  The b l o c k to gene f l o w  polyploid populations  suggests a g e n e t i c  barrier  f a v o u r i n g s u b s p e c i a t i o n i n time. V a r i a b i l i t y observed i n d i f f e r e n t c h a r a c t e r s  suggests t h a t  s i z e of f l o w e r s , dimensions o f p e t a l s , morphology o f b a s a l b r a c t e a l leaves are unstable characters. p o l l e n g r a i n s , and  and  S i z e o f stomata, pubescence,  seeds a l s o y i e l d e d no d i a g n o s t i c f e a t u r e s to  serve as key c h a r a c t e r s f o r d e l i m i t i n g i n f r a s p e c i f i c c a t e g o r i e s i n J5. f e r r u g i n e a .  45 Many c l a s s i f i c a t i o n s o f the _S. f e r r u g i n e a complex have appeared as i s e v i d e n t s p e c i e s was d e s c r i b e d 1eucanthemifolia  synonymy.  A l t h o u g h the  i n 1829, i t was t r e a t e d as a synonym o f S.  i n the 19th century.  i n c l u d e d i t under "S. called i t S.  from the e x t e n s i v e  Torrey  leucanthemifolia  (Lapeyr  leucanthemif o l i a brunoniana.  and Gray (1838-40) ?)". '  Engler  (1872)  Index Kewensis s t a t e s  " S a x i f r a g a f e r r u g i n e a R. Grah...= l e u c a n t h e m i f o l i a . " (Hooker and J a c k s o n 1895).  S m a l l (1896), d i s c u s s i n g the v a l i d i t y o f S.  stated " I t i s apparently  v e r y r a r e and has u s u a l l y been r e f e r r e d to  a v a r i e t y o f S. m i c h a u x i i i n that s p e c i e s . "  S.  ferruginea,  (S. l e u c a n t h e m i f o l i a Michx.)  or included  l e u c a n t h e m i f o l i a Michx. i s now r e g a r d e d as a  synonym o f the e a s t e r n N o r t h American S. m i c h a u x i i endemic t o e a s t e r n U n i t e d  S t a t e s and d i s t i n c t  ( F e r n a l d 1950, Gleason and C r o n q u i s t  B r i t t . which i s  from _S.  ferruginea  1963).  Small (1905) d i v i d e d the complex i n t o f o u r s p e c i e s as summarized i n the key under the genus S p a t u l a r i a :  Inflorescence with b u l b l e t s P l a n t s w i t h a l l the f l o w e r s r e p r e s e n t e d w i t h a s i n g l e flower  terminating  by b u l b l e t s , o r  the main a x i s . S. v r e e l a n d i i  P l a n t s w i t h many f l o w e r s and some b u l b l e t s , a t l e a s t w i t h a flower  teminating  each branch. S.  brunoniana  I n f l o r e s c e n c e without b u l b l e t s Upper p e t a l s 7. 5 to 8. 0 mm. l o n g  Si. newcombei  Upper p e t a l s 3 t o 5 mm.long  S.  ferruginea  46 T h i s key f a i l s even when a p p l i e d to p o p u l a t i o n s mentioned Small.  by  P l a n t l e t f o r m a t i o n i s v e r y common i n the p o p u l a t i o n s o f  Juneau, P r i n c e Rupert and  Vancouver, but, a c c o r d i n g to S m a l l ,  r e g i o n should produce p l a n t s l a c k i n g p l a n t l e t s .  this  Size of p e t a l s i s  l i k e w i s e a v a r i a b l e c h a r a c t e r ( F i g . 8). E n g l e r and Small  Irmscher (1916) u n i t e d a l l e a r l i e r  (1905) and regarded  f e r r u g i n e a Graham.  segregates  of  them as a s i n g l e s p e c i e s , S a x i f r a g a  They d e s c r i b e d t h r e e v a r i e t i e s based on p a t t e r n  o f p l a n t l e t f o r m a t i o n i n the i n f l o r e s c e n c e as i n d i c a t e d i n the summarized key  Var. a.  ( t r a n s l a t e d from L a t i n ) .  t y p i c a Engl.and  P l a n t s without Var. 0. M a c o u n i i Only I and  V.  any b u l b l e t s whatsoever.  E n g l , ani Irmsch. I I branch  floriferous, Var.  Irmsch.  Vreelandii  a p i c e s o f the i n f l o r e s c e n c e  the r e m a i n i n g branches b u l b i l l i f e r o u s . (Small) Engl, and Irmsch.  I n f l o r e s c e n c e b u l b i l l i f e r o u s except a s i n g l e  flower  t e r m i n a t i n g the main a x i s .  E n g l e r and to be and  Irmscher i n d i c a t e d  the d i s t r i b u t i o n o f v a r .  typica  s o u t h e a s t e r n A l a s k a , Queen C h a r l o t t e I s l a n d s , Olympic Mountains,  the n o r t h e r n Rocky Mountains (Idaho).  on p l a n t s from s o u t h e a s t e r n A l a s k a and  P l a n t l e t s f r e q u e n t l y occur  the Rocky Mountains,  thus  c o n t r a d i c t i n g t h e i r "key c h a r a c t e r " s e g r e g a t i n g the t y p i c a v a r i e t y . A l s o the c h a r a c t e r of v a r . V r e e l a n d i i i s not r e s t r i c t e d to Montana  47 as a s i m i l a r p a t t e r n of f l o w e r i n g has been observed i n the populations let  ( C r a t e r Lake and M a r i o n County).  formation  i n var. M a c o u n i i i s h i g h l y v a r i a b l e ,  r e s e m b l i n g the other  two  f e r r u g i n e a Graham.  He  p a t t e r n of  plant-  frequently  varieties.  Johnson (1923) a l s o r e c o g n i z e d split  the  o n l y one  species  c h a r a c t e r i s t i c s which he regarded as and  The  southern  species,  Saxifraga  i n t o s i x v a r i e t i e s based  'constant  on  p e c u l i a r i t i e s of form  habit':  var.  foliacea:-  (based on bracts'.  seven specimens): ' l a r g e l e a f y  var.  grandiflora:-  ( t h r e e specimens): ' u n u s u a l l y flowers.'  large  var. cuneata:-  ( s i n g l e specimen): 'cuneate bases o f leaves'.  var.  diffusa:-  (two specimens); 'elongated narrow l e a v e s and a l o o s e spreading i n f l o r e s c e n c e of s l e n d e r branches.'  var.  s t e l l a r i f o r m i s : - ( s p e c i m e n s from Mt. Hood and Mt. Adams): 'the resemblance o f the l e a v e s w i t h S. stellaris'.  var.  nivea:-  (a s i n g l e herbarium s h e e t ) : 'leaves cm. or l e s s and f l o w e r s l a r g e i n p r o p o r t i o n to p l a n t s ' .  Apparently h i s observations alone.  He was  extensive regions  fully  aware of t h i s  c o l l e c t i o n s and  are to be  1.5  were based on herbarium specimens l i m i t a t i o n when he  s t a t e d "More  a more c a r e f u l survey o f the p a r t i c u l a r  desired."  As p o i n t e d  out above the b a s a l  leaves  48 ( F i g . 4-6),  b r a c t e a l leaves  tremendous v a r i a t i o n and of i n f r a s p e c i f i c Hitchcock (1923) and used by  ( F i g . 7), and  p e t a l s ( F i g . 8) show  can not be r e l i e d on f o r the d e l i m i t a t i o n  taxa i n j5. f e r r u g i n e a . et al  (1961) combined the v a r i e t i e s o f Johnson  recognized  two  varieties.  The  d i s t i n g u i s h i n g character  them i s the p r o p o r t i o n o f p l a n t l e t s i n the i n f l o r e s c e n c e .  However, these w r i t e r s were c e r t a i n l y aware o f the v a r i a b i l i t y i n p l a n t l e t formation  as i s e v i d e n t  from t h e i r c a r e f u l l y worded  key  to the v a r i e t i e s o f J3. f e r r u g i n e a .  F l o w e r s p a r t i a l l y r e p l a c e d by b u l b l e t s ; w i t h southern range o f the s p e c i e s , g r a d u a l l y northward by var.  ferruginea  the more  replaced  var. m a c o u n i i E n g l . & Irmsch.  Flowers a l l (or n e a r l y a l l ) normal; A l a s , along  the c o a s t ,  southward,  to Nanaimo, Vancouver I . , v e r y o c c a s i o n a l l y  elsewhere to the south.  var.  ferruginea  P l a n t s i n c u l t i v a t i o n have shown conspicuous v a r i a t i o n i n the flower: p l a n t l e t proportions Juneau, P r i n c e Rupert and d e s c r i p t i o n o f var. another.  Therefore,  i n the i n f l o r e s c e n c e .  The  plants  from  Vancouver have been observed to f i t the  f e r r u g i n e a i n one  year and  var. macounii i n  h e r b a r i u m mounts o f the specimens c o l l e c t e d from  the same p o p u l a t i o n  i n d i f f e r e n t y e a r s can be c l a s s e d as e i t h e r of  the two  A l s o the p l a n t l e t s u s u a l l y develop q u i t e e a r l y  varieties.  i n the southern p o p u l a t i o n s layed i n northern  but  populations  the development of p l a n t l e t s i s de-  u n t i l f l o w e r i n g i s w e l l advanced.  49 However, t h i s b e h a v i o u r i s not c o n s i s t e n t . be but  i d e n t i f i e d as var.  So the p l a n t s a r e apt to  f e r r u g i n e a when no p l a n t l e t s have developed  l a t e r i n the season the same p l a n t s may produce p l a n t l e t s and  then one would c o n f i d e n t l y put them under var. macounii. An example o f t h i s s i t u a t i o n annotated on a s i n g l e sheet w i l l A collection  (WTU 101205) by K i n c a i d  s t r e s s the p o i n t .  i d e n t i f i e d as S. s t e l l a r i s has  been annotated by E r i c H u l t e n (1936) as S. f e r r u g i n e a var. Engl,  and Irmsch.  as var.  Another a n n o t a t i o n  macounii  appears on the same sheet  ferruginea.  Although a tremendous m o r p h o l o g i c a l v a r i a b i l i t y has been observed i n the s p e c i e s complex, t h e o n l y c o n s i s t e n t d i f f e r e n c e i s the presence o f 10-paired The  d i p l o i d and 19-paired  two chromosomal r a c e s , however, a r e m o r p h o l o g i c a l l y  guishable  and do not warrant f o r m a l  Taking i n t o c o n s i d e r a t i o n  a t present  races.  indistin-  r e c o g n i t i o n a t the p r e s e n t  the evidence o b t a i n e d  l i n e s o f study i t i s concluded t h a t S a x i f r a g a  The  polyploid  time.  from v a r i o u s  f e r r u g i n e a i s best  left  as one s p e c i e s h a v i n g i n f r a s p e c i f i c chromosomal r a c e s .  i n c o m p a t i b i l i t y o f d i p l o i d and p o l y p l o i d p o p u l a t i o n s  on the  mainland suggests a b l o c k i n g o f gene flow which i s l i k e l y to r e s u l t i n the d i f f e r e n t i a t i o n o f r a c e s Charlotte Islands,  i n time.  on Queen  i s o l a t e d from the mainland s i n c e g l a c i a t i o n and  therefore having a smaller  gene p o o l as compared to the mainland  stock, would be expected to evolve taxon.  The p o p u l a t i o n s  faster into a distinguishable  The p o s s i b i l i t y o f such e v o l u t i o n i n i s o l a t i o n has been  suggested by F o s t e r  (1965) i n the c h a r a c t e r s  the n a t i v e mammals o f the Queen C h a r l o t t e  of i n s u l a r races  Islands.  :  among  50 During  the survey  for variability  i n t e r e s t i n g p o i n t s have come up  i n S. f e r r u g i n e a a  few  that deserve some d i s c u s s i o n and  should be pursued f u r t h e r . The  occurrence  of a 10-paired  d i p l o i d and  a 19-paired  p l o i d w i t h i n a s p e c i e s poses the i n t e r e s t i n g problem o f how  polyand  where the p o l y p l o i d s o r i g i n a t e d . The  simplest hypothesis  to e x p l a i n the o r i g i n of t e t r a p l o i d s  from d i p l o i d s i n v o l v e s d i r e c t a u t o p o l y p l o i d y  (Hayman 1960).  According  to MUntzing (1936) a u t o p o l y p l o i d y has p l a y e d a very  important  r o l e i n the e v o l u t i o n of h i g h e r  out  that a u t o p o l y p l o i d y may  aneuploidy  and  important.  b a s i c numbers a r e the most  8,9,10,11,12,13 and  s p e c i e s i n the s e c t . B o r a p h i l a w i t h S.  i s therefore widely  o c c i d e n t a l i s plexus  14 chromosomes. 19 p a i r s e.g.  (Beamish 1961).  s c a t t e r e d i n t h i s genus.  There _S.  Aneuploidy  Polyploids with  f e r r u g i n e a c o u l d have o r i g i n a t e d i n e i t h e r of two ways.  f u s i o n of one and  a l s o pointed  D a r l i n g t o n and W y l i e (1955) have l i s t e d b a s i c numbers i n  i n t e g r i f o l i a and  i n S.  He  have secondary e f f e c t s , o f which  the f o r m a t i o n o f new  the genus S a x i f r a g a to be are other  plants.  normal gamete w i t h another having  subsequent p o l y p l o i d i z a t i o n of the h y b r i d .  have been p r e s e n t  i n the p a s t , but now  that crossed with  the 10-paired  allopolyploid.  The  Second, t h e r e  may  absent, a s p e c i e s w i t h 9 p a i r s  former argument may  to a  19-paired  be more c l o s e to r e a l i t y  s p e c i e s c l o s e l y r e l a t e d to S_.  A r t i f i c i a l c r o s s e s among d i p l o i d s and  be made i n l a r g e numbers i n o r d e r  First,  chromosome l e s s  species g i v i n g r i s e  s i n c e t h e r e i s no r e c o r d of a 9-paired ferruginea.  one  19 p a i r s  polyploids  to o b t a i n h y b r i d s .  should  P a i r i n g behaviour  o f the chromosomes i n the h y b r i d s might p r o v i d e v a l u a b l e tion.  I f hybrid  informa-  seeds do not a t t a i n f u l l development i t w i l l  f e a s i b l e to attempt a r t i f i c i a l experiments should be  c u l t u r e of e x c i s e d embryos.  be  These  supplemented by a r t i f i c i a l p o l y p l o i d i z a t i o n  of d i p l o i d s to e l u c i d a t e the problem o f p o l y p l o i d y i n t h i s  species.  Chromatography has been used s u c c e s s f u l l y i n Asplenium (Smith and L e v i n 1963) p l o i d and  to d e t e c t  the parent  the p o s t u l a t e d a n c e s t r y was  confirmed.  (1963) noted an i n f r a s p e c i f i c c h e m i c a l and  s p e c i e s of the  A l s t o n and  The  chromatographic study on S.  population  f e r r u g i n e a gave one  s i s t e n t p a t t e r n o f p h e n o l i c a c i d s f o r b o t h d i p l o i d and lations.  Turner  variation in Baptisia nuttalliana  suggested the use of chromatography f o r a f u r t h e r  analysis.  poly-  con-  p o l y p l o i d popu-  T h i s t e n t a t i v e l y suggests the o r i g i n o f p o l y p l o i d s by  polyploidy.  However, t h i s i n f o r m a t i o n alone  conclusion.  A survey o f f l a v o n o i d compounds might b r i n g out  ferences  should  and  be made.  auto-  cannot l e a d to a f i r m some d i f -  Jay and L e b r e t o n (1965) examined the  t r i b u t i o n of f l a v o n o i d s i n a l a r g e number of s p e c i e s of S a x i f r a g a __• f e r r u g i n e a was  not  included i n their  but  study.  A second p o i n t of i n t e r e s t i s the d i s t r i b u t i o n p a t t e r n o f two  chromosomal r a c e s .  r e l a t e d d i p l o i d and  the  D i f f e r e n c e s i n the d i s t r i b u t i o n o f c l o s e l y  p o l y p l o i d s p e c i e s have been observed by Hagerup  (1933), T i s c h l e r (1934), S o k o l o v s k a j a Lb*ve (1943) and  dis-  others.  and  Strelkova  (1938), Lbve  Iii . B i s c u t e l l a l a e v i g a t a (Manton 1934)  p l o i d forms have t h e i r h a b i t a t i n c e n t r a l Europe, w i t h i n the which were covered w i t h  i c e during  the g r e a t g l a c i a t i o n ,  and  the  poly-  regions  while  o l d e r d i p l o i d forms have t h e i r h a b i t a t i n the i c e f r e e r e g i o n s  the i n the  52 south.  A n o r t h e r l y d i s t r i b u t i o n of p o l y p l o i d s i s not an  rule.  There are cases where the r e v e r s e  i s true.  absolute  F l o v i k (1940)  f o r example, found d i p l o i d C h r y s o s p l e n i u m tetrandrum i n  Spitzbergen  while  I n Vaccinium  the more southern C.  uliginosum  alternifolium i s tetraploid.  (Hagerup 1933), and  Campanula r o t u n d i f o l i a  (BHcher  1936)  the d i p l o i d s have a more marked a r c t i c d i s t r i b u t i o n than the p o l y p l o i d races.  In Saxifraga f e r r u g i n e a p o l y p l o i d s i n general  t h e r l y d i s t r i b u t i o n on I s l a n d and  the mainland.  Queen C h a r l o t t e I s l a n d s  a r e a s were p a r t l y u n g l a c i a t e d .  The  The  d i p l o i d s discovered  s t r o n g l y favour  et a l 1958, The  a t Kodiak  the i d e a t h a t  d i p l o i d population reported  Packer (1968) c o i n c i d e s w i t h a r e g i o n b e l i e v e d to be (Wilson  show more nor-  these by  unglaciated  P r e s t et a l 1967).  d i s t r i b u t i o n of d i p l o i d s and  p o l y p l o i d s i n j>. f e r r u g i n e a  appears t o be r e l a t e d to P l e i s t o c e n e G l a c i a t i o n i n the C o r d i l l e r a n G l a c i a l Complex and  plant migration  i n p o s t - g l a c i a l times.  C o r d i l l e r a n G l a c i a l Complex:the P l e i s t o c e n e i c e sheet  At  the time of maximum g l a c i a t i o n  extended from the A l a s k a n A l e u t i a n I s l a n d s  the Cascade Mountains n o r t h of the Columbia R i v e r . G l a c i a l Complex was eastern  2350 m i l e s  long and,  s i d e of the Rocky Mountains, was  However some a r e a s remained u n g l a c i a t e d The regions  to  This C o r d i l l e r a n  from Vancouver I s . to 5"50miles wide ( F l i n t  the 1957).  ( F i g . 19).  a v a i l a b l e evidence/about the problem o f f e f u g i a i n v a r i o u s  f a l l i n g w i t h i n the range of S a x i f r a g a f e r r u g i n e a i s mentioned  below. Kodiak I s : -  Kodiak I s l a n d i s the major i s l a n d  southwest of K e n a i P e n i n s u l a  separated  from A l a s k a by  i n the Shelikof  chain Strait.  53 The Kodiak Mountains a r e  t o p o g r a p h i c a l l y and g e o l o g i c a l l y a c o n t i n u -  a t i o n o f the K e n a i Mountains.  They r e v e a l a g l a c i a t e d s u r f a c e  r i d g e s and summits up to ca. 4000 t o 5000 feet e l e v a t i o n .  with  Ice free  c i r q u e s , some a t e l e v a t i o n s as low as 500 to 1000 f e e t and 'U' shaped v a l l e y s a t 4000 f e e t a t t e s t Heusser  (1960) a l t h o u g h  to i n t e n s i v e g l a c i a t i o n .  According  Kodiak I s . and i t s n e i g h b o u r i n g  to  i s l a n d s served  as a l o c u s f o r g l a c i a t i o n ,  the upper h e i g h t s o f the i s l a n d appear to  have stood above the i c e .  According  punctata and  ssp, p a c i f i c a i s a v a l i d  capsule characters, confined  Alaska.  to C a l d e r and S a v i l e (1960) j>.  subspecies,  based on d i s t i n c t  to a narrow a r c along  the G u l f o f  Although they b e l i e v e that t h i s a r e a was c o m p l e t e l y  they c o n s i d e r  the p o s s i b i l i t y  that ssp. p a c i f i c a  leaf  glaciated  s u r v i v e d the l a s t  g l a c i a t i o n on p a r t s o f the c o n t i n e n t a l s h e l f t h a t a r e now below sea level.  I n p a r t i c u l a r A l b a t r o s s Bank and P o r t l o c k Bank, southeast and  northeast  r e s p e c t i v e l y o f Kodiak I s . may w e l l have escaped complete  glaciation. Alaskan  Alexander A r c h i p e l a g o : -  Based on p l a n t c o l l e c t i o n s made  between 5600 f e e t and 6900 f e e t e l e v a t i o n w i t h i n the Juneau I c e F i e l d , Heusser  (1954) concluded  have s u r v i v e d . maximum w h i l e 'little  t h a t o n l y 18 s p e c i e s o f v a s c u l a r p l a n t s would  The e x i s t i n g f l o r a developed m a i n l y d u r i n g some s p e c i e s have migrated  i c e age'.  the thermal  to the nunataks s i n c e the  The upper l i m i t o f g l a c i a t i o n  i n the Juneau I c e  F i e l d n o r t h o f the Taku R i v e r V a l l e y i s 5600 t o 6000 f e e t on the nunataks i n the i n t e r i o r o f the f i e l d and between 5000 to 5500 f e e t on the margin. These f i g u r e s agree w i t h  the g l a c i a l Map o f Canada (Wilson e t a l 1958).  Heusser found S a x i f r a g a f e r r u g i n e a to be w i d e l y  d i s t r i b u t e d from 3500  54  feet  to 5900 f e e t e l e v a t i o n and S. f e r r u g i n e a v a r macounii a t 4500  f e e t e l e v a t i o n i n the Juneau I c e F i e l d . Wrangell,  Petersburg,  Peat s e c t i o n s from  Ketchikan,  S i t k a and Juneau suggest the g l a c i a t i o n o f the  e n t i r e r e g i o n except p o s s i b l y the h i g h e r peaks (Heusser 1952). Queen C h a r l o t t e I s l a n d s : - T h e  Queen C h a r l o t t e I s l a n d s have  a t t r a c t e d s c i e n t i s t s o f v a r i o u s f i e l d s due to t h e i r i s o l a t e d p o s i t i o n . Opinions  d i f f e r about the g l a c i a t i o n o f the i s l a n d s , but evidence i s  accumulating evidence  i n favour o f the e x i s t e n c e o f u n g l a c i a t e d areas.  comes from 3 l i n e s o f study. 1.  B o t a n i c a l Evidence:-  Calder  (1964) e s t i m a t e s  t o t a l v a s c u l a r f l o r a o f the i s l a n d s i s a p p r o x i m a t e l y which 45 t o 65 a r e a l p i n e . degree o f endemism.  470 s p e c i e s o f  There a r e many unique endemic s p e c i e s and a few  S a v i l e 1959, F o s t e r 1965).  among p t e r i d o p h y t e s  t h a t the  Among the a l p i n e s p e c i e s there i s a h i g h  endemic r a c e s among spermatophytes (Mathias and  The  and Constance 1959, C a l d e r  Other i n t e r e s t i n g cases a r e known  ( T a y l o r 1967) and a l a r g e body o f i n f o r m a t i o n has  accumulated i n b r y o p h y t e s ( S c h o f i e l d 1968) i n d i c a t i n g cases o f b i c e n t r i c distribution.  Such b i c e n t r i c d i s t r i b u t i o n i s c o n s i d e r e d  a good  indica-  t i o n of plant r e f u g i a . 2.  Z o o l o g i c a l Evidence:-McCabe and Cowan (1945) p r o v i d e  z o o l o g i c a l evidence  and l i s t many w e l l d e f i n e d s p e c i e s r e s t r i c t e d  Queen C h a r l o t t e I s l a n d s .  They suggest that those animals have  on the i s l a n d s from a p r e g l a c i a l o r i n t e r g l a c i a l stock.  to the  evolved  Foster  (1965)  d i s c u s s i n g the p o s s i b l e paths o f e v o l u t i o n among the n a t i v e mammals o f the Queen C h a r l o t t e I s l a n d s concludes  that ancestors  o f the w e a s e l ,  b e a r , marten and a r a c e o f shrew s u r v i v e d i n the Queen C h a r l o t t e  55 Islands r e f u g i a during  the l a s t g l a c i a t i o n .  The  c h a r a c t e r s of  those  i n s u l a r r a c e s are a r e s u l t o f e v o l u t i o n i n i s o l a t i o n s i n c e the  last  glaciation. 3.  G e o l o g i c a l Evidence:-  g l a c i a t e d areas i n Scandinavia  Dahl (1955) r e f e r r i n g  to the  s t a t e d t h a t where h i g h mountains abut  on deep sea, g l a c i a t i o n i s l i k e l y to have been incomplete and present.  refugia  Such a s i t u a t i o n i s observed on the west c o a s t of Queen  Charlotte Islands. McCabe and and  un-  Brown and Nasmith (1962) quoted statements of  Cowan (1945. p.158), Heusser (1955  S a v i l e (1959  p.64), and  Beebe (1960  p.  p.442, 443,  153)  447),  expressing  Calder  some doubt  about the v a l i d i t y o f the b i o l o g i c a l evidence f o r u n g l a c i a t e d a r e a s Queen C h a r l o t t e I s l a n d s . support  They have presented  complete g l a c i a t i o n but  o f some u n g l a c i a t e d  p h o t o g r a p h i c evidence to  they have a l s o l e f t open the  possibility  areas.  Late Pleistocene Migration:- According  to C r a n d e l l ,  Mullineaux  and Waldron (1958) the r e g i o n south of the g l a c i a l boundary was main source o f p l a n t m i g r a n t s . t r i c u s p i d a t a and  on  The  p l o t t e d d i s t r i b u t i o n of  the  Saxifraga  S. b r o n c h i a l i s ssp. austromontana i n d i c a t e the r a t e s a t  which these p l a n t s r e c o l o n i z e d B r i t i s h Columbia from t h e i r r e f u g i a i n Montana to s o u t h e r n Washington ( C a l d e r and b r o n c h i a l i s ssp, austromontana has coast  S a v i l e 1959b).  t r a v e l l e d f a r t h e r n o r t h near  than i n the i n t e r i o r o f B r i t i s h Columbia.  800 m i l e s  As  A spread  s i n c e the r e t r e a t of i c e shows the e f f i c i e n c y of  even i n h i g h l y d i s s e c t e d c o u n t r y  like British  f a r as i s known at p r e s e n t  l a t i o n s on Vancouver I s . and  Saxifraga the  o f 600  to  migration  Columbia.  the S a x i f r a g a f e r r u g i n e a popu-  on the mainland n o r t h o f the  glacial  56 boundary a r e a l l p o l y p l o i d . the areas rendered  Apparently  the p o l y p l o i d s r e c o l o n i z e d  a v a i l a b l e by the r e c e d i n g g l a c i e r s .  l a t i o n s south o f the g l a c i a l margin a r e d i p l o i d . covered  The popu-  The d i p l o i d s  dis-  on Kodiak I s . , Queen C h a r l o t t e I s l a n d s and o t h e r s c a t t e r e d  locations*: s on the mainland p r o b a b l y single diploid  represent  the components o f a  s p e c i e s widespread b e f o r e the P l e i s t o c e n e G l a c i a t i o n  and now p r e s e n t as r e l i c t escaped g l a c i a t i o n .  populations  i n o r near areas which  The t h i r d p o s s i b i l i t y ,  t h a t the S a x i f r a g a f e r r u g i n e a  p o p u l a t i o n s on the Queen C h a r l o t t e I s l a n d s were d e r i v e d from the mainl a n d s t o c k a f t e r g l a c i a t i o n i s u n l i k e l y f o r two reasons. o r i g i n of a d i p l o i d  i s not expected  F i r s t , the  from a p o l y p l o i d stock.  the p o l l e n p r o f i l e s from Langara I s . (Q.C.I.), K e t c h i k a n , Rupert  and P r i n c e  (Heusser 1955) i n d i c a t e the Langara I s . p r o f i l e to be o l d e r  than the other  two p r o f i l e s  so t h a t p l a n t s a r r i v e d on the mainland much  l a t e r due t o the more prolonged other  Second,  g l a c i a l cover.  The o n l y  possibility  than i t s p e r s i s t e n c e i n the Queen C h a r l o t t e I s l a n d refugium, i s  the d i s p e r s a l and m i g r a t i o n from p o p u l a t i o n s p e r s i s t i n g beyond the g l a c i a l border  i n Washington 600 m i l e s d i s t a n t .  The c o n d i t i o n s o f  such long range d i s p e r s a l a r e so complex t h a t the chances o f a f u l establishment  o f a d i a s p o r e would be h i g h l y remote.  d i p l o i d p o p u l a t i o n s o f Kodiak I s . p r o b a b l y p r e g l a c i a l times and managed  arrived  success-  S i m i l a r l y , the  from the mainland i n  to p e r s i s t d u r i n g g l a c i a t i o n .  The  apparent absence on Queen C h a r l o t t e I s l a n d s o f p o l y p l o i d p o p u l a t i o n s a l s o argues s t r o n g l y a g a i n s t p o s t - p l e i s t o c e n e m i g r a t i o n o f S a x i f r a g a f e r r u g i n e a from the mainland. I f the d i s t r i b u t i o n o f the s p e c i e s i s superimposed on the  C o r d i l l e r a n region:. (Bostock,  M u l l i g a n and Douglas 1957, F i g . 61), i t  i n d i c a t e s t h a t t h e m i g r a t i o n has p r o b a b l y  taken p l a c e a l o n g the  Coast Mountains on the one hand and a l o n g  the P u r c e l l , S e l k i r k ,  Monashee, C a r i b o u and Rocky Mountains on the o t h e r from the unglaciated  southern  region.  Mountains i s p r o b a b l y The by and  uniform  The spread  f a r n o r t h i n the Rocky  b l o c k e d by the L i a r d P l a i n and Hyland  Plateau.  spread o f the s p e c i e s from the c o a s t eastwards i s checked  the I n t e r i o r p l a t e a u i n B r i t i s h Columbia. Skeena Mountains enabled  However, the H a z e l t o n  the m i g r a t i o n from the Coast Mountains  inland. Although the h a b i t a t o f S. f e r r u g i n e a i s v a r i e d i n b o t h a l t i t u d e and l a t i t u d e , t h e p a t t e r n o f v a r i a b i l i t y  i s d i f f u s e and does  not warrant d e l i m i t a t i o n o f s u b s p e c i f i c taxa, a t l e a s t a t the p r e s e n t time.  T h i s l a c k o f d i f f e r e n t i a t i o n a f t e r g l a c i a t i o n might be due to a  slow r a t e o f e v o l u t i o n a r y change o r to the mode o f p r o p a g a t i o n . v a r i a b i l i t y , whatsoever, i s imparted by  the normal s e x u a l p r o c e s s .  The  o n l y by seed p r o d u c t i o n , which i s  However, the s p e c i e s propagates  itself  q u i t e e f f i c i e n t l y through v e g e t a t i v e apomixis ( p l a n t l e t s and rhizomes). The  l a t t e r perpetuates  o n l y the p a r e n t a l genotype.  S i n c e we now know  of the e x i s t e n c e o f g e n e t i c b a r r i e r s to gene f l o w between d i p l o i d and p o l y p l o i d p o p u l a t i o n s , we may expect d i f f e r e n t i a t i o n i n the c o u r s e o f time t h a t w i l l j u s t i f y the r e c o g n i t i o n o f i n f r a s p e c i f i c c a t e g o r i e s .  1  58 SUMMARY V a r i a b i l i t y w i t h i n the S a x i f r a g a f e r r u g i n e a complex i s r e p o r t e d based on evidence from c y t o l o g i c a l , embryological, studies.  anatomical,  morphological,  chromatographic, and  crossability  In a d d i t i o n to herbarium specimens r e p r e s e n t i n g  the  e n t i r e range of the s p e c i e s , l i v i n g p l a n t s were examined from Kodiak I s . and  Juneau ( A l a s k a ) , A l l i f o r d  Bay  and  Onward P o i n t  (Queen C h a r l o t t e I s l a n d s , B r i t i s h Columbia), P r i n c e Rupert Vancouver  (mainland B r i t i s h Columbia), and M a r i o n County  Chromosome counts r e v e a l e d d i p l o i d s and characters  19-paired  two  chromosomal r a c e s ,  polyploids (aneuploids).  embryologically  A l l the p o p u l a t i o n s  identical.  n u c e l l u s i s conspicuous and Anatomically  rhizome but not  has  the outermost l a y e r o f not been r e p o r t e d  innerthe  i n the genus  no d i f f e r e n c e s were observed between popufeatures of  the  the presence of an endodermis i n the  i n the peduncle, the o c c u r r e n c e o f druses i n the  s e p a l s , and  ovary w a l l s , secondary growth i n a d v e n t i t i o u s  r o o t s , a p i c a l o r g a n i z a t i o n o f the shoot and and  Morphlogical  i n v e s t i g a t e d were  However, s e v e r a l i n t e r e s t i n g a n a t o m i c a l  s p e c i e s are r e p o r t e d :  leaves,  10-paired  An u n u s u a l behaviour of the  most l a y e r of the integument and  lations.  (Oregon).  s t u d i e d i n d i c a t e d extreme v a r i a b i l i t y between but  also within populations.  before.  and  young i n f l o r e s c e n c e ,  the course of development o f p l a n t l e t s a t the expense o f  flowers. similar  The  p a t t e r n of d i s t r i b u t i o n o f p h e n o l i c  i n a l l populations  chromatographed.  acids i s  Si. f e r r u g i n e a  lacks  59 2,hydroxy phenyl a c e t i c a c i d , a compound r e p o r t e d by o t h e r workers o n l y i n one genus, A s t i l b e , of S a x i f r a g a c e a e . set no seed i n d i c a t i n g diploids.  However,  Inter-population  crosses  a b l o c k to gene f l o w between p o l y p l o i d s and  i n t e r p o p u l a t i o n c r o s s e s between d i p l o i d s and a l s o  between p o l y p l o i d s a r e s u c c e s s f u l b u t w i t h a reduced seed s e t as compared to i n t r a p o p u l a t i o n c r o s s e s . The d i s t r i b u t i o n o f d i p l o i d s and p o l y p l o i d s suggests a c o r r e l a t i o n with  the P l e i s t o c e n e G l a c i a t i o n i n the C o r d i l l e r a n G l a c i a l  Complex. A l t h o u g h a tremendous m o r p h o l o g i c a l  v a r i a b i l i t y has been  observed, o n l y the d i f f e r e n c e of chromosome numbers and  correlated with d i s t r i b u t i o n .  appear to be m o r p h o l o g i c a l l y ference  i s consistent  The two chromosomal r a c e s , however,  i n d i s t i n g u i s h a b l e , t h e r e f o r e , the d i f -  i n chromosome numbers does n o t j u s t i f y d e l i m i t a t i o n o f i n f r a -  s p e c i f i c c a t e g o r i e s i n S.  f e r r u g i n e a at present.  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(Doklady) de l'Acad. des S c i . de 1* URSS 21: 68-71. Soueges, E.C.R. 1936. Embryogenie des S a x i f r a g a c e e s Developpement de 1'embryon chez l e S a x i f r a g a g r a n u l a t a L. Compt. Rend. Acad, des S c i . P a r i s 202: 240-242. S t e b b i n s , G.L. J r . 1950. V a r i a t i o n and E v o l u t i o n i n P l a n t s . Columbia Univ. P r e s s . N.Y. xx + 643 pp. T a y l o r , R.L. 1965. The genus Lithophragma. (Saxifragaceae) Univ. C a l i f . P u b l . Bot. 37. v i + 122 pp. 1968  P e r s o n a l Communication.  and G.A. M u l l i g a n . 1968. F l o r a of the Queen C h a r l o t t e Islands. P a r t 2. Research Branch. Can. Dept. Agr. Monograph No. 4. P a r t 2: i x + 148 pp. ( S a x i f r a g a f e r r u g i n e a p. 77). T a y l o r , T.M.C. 1967. Mecodium w r i g h t i i i n B r i t i s h Columbia and Am. F e r n . Jour. 57 ( 1 ) : 1-6. Thouvenin, M. 1890. Ann. S c i . Nat.  Recherches sur l a s t r u c t u r e des Bot. Ser. 7. 12: 1-174.  Alaska.  Saxifragaceae.  T i s c h l e r , G. 1934. Die Bedeutung der P o l y p l o i d i e fUr d i e V e r b r e i t u n g der Angiospermen e r l M u t e r t an den A r t e n S c h l e s w i g - H o l s t e i n s , mit A u s b l i c k e n auf andere F l o r e n g e b i e t e . Bot. Jahrb. 67: 1-36. T o r r e y , J . and A. Gray. (1838-40). S a x i f r a g a l e u c a n t h e m i f o l i a In: A f l o r a o f North America V o l . I. W i l e y and Putnam, N.Y. : p.570. W a t a r i , S. 1939. Anatomical s t u d i e s on the l e a v e s of some S a x i f r a gaceous p l a n t s w i t h s p e c i a l r e f e r e n c e to the v a s c u l a r system. J . Fac. S c i . I m p e r i a l Univ. Tokyo, s e c t i o n I I I Botany. 5(3): 195-316. Wiggins, I r a L. 1959. Development of the o v u l e and megagametophyte in Saxifraga h i e r a c i f o l i a . Am. J . Bot. 46: 692-697. W i l s o n , S.T. at a l . 1958. Toronto.  G l a c i a l Map  of Canada. Geol. Assoc.  Canada.  66 APPENDIX I. Specimens r e p r e s e n t e d t o i l l u s t r a t e v a r i a t i o n i n b a s a l l e a v e s o f S. CALIFORNIA:  1.  (shown i n F i g . 4)  ferruginea.  S i s k i y o u County, Salmon Mountains, A n n e t t a C a r t e r  1473 (UC);  2.  S i s k i y o u County, Sanger Peak, C . L . H i t c h c o c k & J . S . M a r t i n 5287 (UC, WTU);  3.  S i s k i y o u County, Mt. D i a b l o , L.C.Wheeler  3147 (UC);  County, Poker F l a t , R. B a c i g a l u p i 7448 (MIN,JEPS);  5.  4.  Siskiyou  S i s k i y o u County,  C a r i b o u Lake, I r a L. Wiggins 13548 (UC,DS,WSP). MONTANA: 2.  1.  M i s s o u l a County, G l a c i e r Lake, C . L . H i t c h c o c k 18201 (UC,WTU,WSP);  R a v a l l i County, St. Mary's Peak, C . L . H i t c h c o c k 17110 (WSP,WTU);  G l a c i e r N a t i o n a l Park, W.C.McCalla Umbach 812 (MIN);  3772 (MIN);  4.  3.  Sperry G l a c i e r , L.M.  5. R a v a l l i County, St. Mary's Lake, C . L . H i t c h c o c k &  C. V.Muhlick 15335 (UC,WTU,WSP). IDAHO:  1.  K o o t e n a i County, P a c k s a d d l e Mountain, J . B . L e i b e r g 468 (MIN); 2.  Boundary County, Smith Creek, G. B. Rossbach & R. P. Rossbach 245 (UC,WTU); 3.  K o o t e n a i County P a c k s a d d l e Mountain, D.T.MacDougal 384 (MIN);  4.  K o o t e n a i County, H i g h Mountains, J.H. Sandberg, s.n. (MIN). ALASKA:  1.  B e r i n g Sea r e g i o n , P o r t E t c h e s , J.M.Macoun s.n. (CAN); 2,3.  P r i n c e o f Wales I s l a n d near Ratz Harbour, J . B . F l e t t 1946 (WTU, WSP); 4.  Juneau I c e F i e l d , Taku B,  C.H. Heusser 214 (OSC);  o f h i g h mountain, Thomas Howell 1621 (UC,MIN); southeast o f Hope, J . A . C a l d e r 6510 (UC,DA0); l o c a l i t y , J.P.Anderson 5564 (MIN); L l o y d A. Spetzman 2608 (CAN,US); J.A. C a l d e r 5267 (DAO,ALA); Eyerdam 7430 (MIN); s.n. (MIN);  8. 9.  Thompson Pass, h i g h a l p i n e ,  11. Shores o f Behm C a n a l , r o c k b l u f f s , M.W.Gorman L a r s e n Bay, E r n e s t Lepage 25071 (CAN); 13. 7207 (MIN,CAN);  A / A . H e l l e r 14947 (WSP)  Bay, W.J.Eyerdam 3548 (UC); Juneau I c e F i e l d , Taku E, T . K i n c a i d s.n. (WTU);  Dearborn s.n. (OSC).  Juneau, r o c k y a l p i n e  K e n a i P e n i n s u l a , Moose Pass V a l l e y ,  Evans I s l a n d , P o r t San Juan, W.J. Eyerdam  ity,  7.  Kenai Peninsula,  10. K u i u I s l a n d , Washington Bay, W.J.  12. K o d i a k I s .  near Sawmill Creek,  6.  5. Yes Bay, Summit  16.  15.  14. S i t k a ,  K n i g h t I s l a n d , Thum  C r a i g , L.Norberg s.n. (UC,CAN);  G.W.Argus 60 (ALA);  18.  17.  Juneau, a l p i n e  19. Afognak I s . , L i t t l e K i t o i Lake, John  local-  67 APPENDIX I I Specimens r e p r e s e n t e d t o i l l u s t r a t e v a r i a t i o n  (shown i n F i g . 5)  i n b a s a l l e a v e s o f S. f e r r u g i n e a i n BRITISH COLUMBIA. 1.  Head o f A z o u z e t t a Lake, on H a r t Highway 97, R. L. T a y l o r and  G. Staudt 4159 (WSP);  2.  Manning Park, B l a c k w e l l Peak, J.A.  C a l d e r & D. B. 0. S a v i l e 10518 Newcombe (CAN); 542  (CAN);  5.  (WTU,CAN,UBC);  4.  (CAN); 10.  9.  3.  K i m s q u i t , mouth o f Dean  Banks I s l a n d , C F . R i v e r , H.M. L a i n g  Haines Road, M i l e 60, T.M. C. T a y l o r e t a l 1207 6.  McCabe 1438 (WTU); (UC,CAN,WTU);  (WTU, DAO) ;  8.  B e l l a C o o l a , r o c k s above timber l i n e , T.T. 7.  Kokanee G l a c i e r Park, J . W. Thompson 14454  Nootka, Vancouver I s . , W. Spreadborough 79351  H a z e l t o n , Rocher de Boule Mountain, J.A. Munro 189 (CAN);  Kokanee G l a c i e r Park, J.W. Thompson 14454 (UC,CAN,WTU);  C h i l l i w a c k Lake, J.M. Macoun 919 (CAN);  11.  12. Sunburst Lake Camp,  Mt. A s s i n i b o i n e , E d i t h Scamman 6645 (CAN);  13. B l a n k e t Creek F a l l s ,  between R e v e l s t o k e and Arrowhead, J.A. C a l d e r & D. B. 0. S a v i l e 8950 (DAO, ALA); 14. A l l i f o r d Bay, Queen C h a r l o t t e I s l a n d s , A. S. Randhawa 1589  (UBC);  15. Mt. Seymour, Vancouver, A. S. Randhawa 1498 (UBC);  16.  Queen C h a r l o t t e I s l a n d s , T r i - t s u l t Mountains, C M . Dawson 34921  (CAN);  17. G l a c i e r , Mt. Abbott, F. K. B u t t e r s , E.W.D. Hoi way 259  (MIN);  18. Lake Cowichan, Vancouver I s . , C O . Rosendahl 1809  (MIN);  19. Squamish, A.S. Randhawa 1500 (UBC); 20. G l a c i e r , F.K.  B u t t e r s & E.W.D. Holway 259 (MIN); A. S. Randhawa 1485  (UBC);  21. Horseshoe Bay,  Vancouver,  22. Onward P o i n t , Queen C h a r l o t t e  I s l a n d s , A.S. Randhawa 1591 (UBC);  23. Mt. Moresby, Queen C h a r l o t t e  I s l a n d s , A. S. Randhawa 1590 (UBC); 24. V a l l e y , F. K. B u t t e r s e t a l 492 (MIN);  S e l k i r k Mountains, Cougar 25. Mt. R e v e l s t o k e , near  Heather Lodge, J.A. C a l d e r and D. B. 0. S a v i l e 10841  (UC, DAO);  26.  Squamish, A. S. Randhawa 1500 (UBC); 27. Tungsten Mine,east o f Skeena C r o s s i n g , J.A. C a l d e r , D. B. 0. S a v i l e & J.M. 28.  Ferguson 13375 (UC, DAO);  S e l k i r k R a n g e , v i c i n i t y o f G l a c i e r , J.Woods 3657 (MIN); 29.  R i c h a r d s o n P o i n t , Vancouver, A. S. Randhawa 1485 (UBC).  68 APPENDIX I I I Specimens r e p r e s e n t e d to i l l u s t r a t e v a r i a t i o n  (shown i n F i g .  6)  i n b a s a l l e a v e s o f S. f e r r u g i n e a .  WASHINGTON:  1.  37 (WSP);  Whatcom County, 33 m i l e s east o f E v e r s o n , L y l e P y e a t t  2. C h e l a n County, west f a c e o f Mt. S t u a r t , A.R.Kruckeberg  2630 (UC,WSP,CAN,WTU); Ohanopeckosh,  3.  Lewis County, 8 m i l e s southeast o f  D. Malcolm 130 (WSP);  Lake, G.N.Jones 2005 (WTU); H.L. Mason 7442 (MIN); (MIN);  7.  5.  6.  K i t t i t a s County, S p e c t a c l e  Whatcom C o u n t y , n o r t h o f Mt. Baker,  Mt. R a i n i e r N a t i o n a l Park, W.C.McCalla  Mt. Adams, J . B . F l e t t s.n. (WTU);  Sandberg & J . B . L e i b e r g  (MIN,WSP)  C . L . H i t c h c o c k 8019 (WTU); Eyerdam 6268 (CAN);  10.  B a s s e t t Maguire 16036 (MIN); P u r e r 7835 (MIN); 15.  9.  8.  Stevens P a s s , J . H.  K i t t i t a s County, Mt. D a n i e l s ,  C l a l l a m County, Deer Lake, W.J.  12.  Whatcom County, T a b l e Mountain,  13. P i e r c e County, Nachess Pass,E.A.  14. Horseshoe b a s i n near S k a g i t Pass, Lake & H u l l Mt. R a i n i e r , W.C.McCalla  5054 (MIN);  16.  Yakima  County, Chinook P a s s , J.W.Thompson 15135 (CAN,WSP ,WTU,UC ,MIN) ; A u s t i n Pass, Mt. Baker, J.W.Thompson 5390 (MIN,M0); Dry  r o c k y p l a c e s , O.D.Allen 199 (UC ,MIN ,WSP);  P a r a d i s e V a l l e y , L.S.Rose 40759 (UC); Col.  5099  11. Okanogan County, n o r t h f o r k o f B r i d g e Creek,  A.D.E. Elmer 691 (MIN,WSP) ;  468 (WSP);  4.  20.  Bob Lookout J.W.Thompson 7256 (UC,WTU);  F o r e s t , W.C.Muenscher  797 (MIN);  J.W.Thompson 14310 (MIN).  22.  19.  18.  17.  Goat Mountain,  P i e r c e County,  Grays Harbor County, 21.  Mt. Baker N a t i o n a l  Snohomish County, Mt. Pugh,  69  APPENDIX I I I (CONTINUED)  OREGON:  1.  J o s e p h i n e County, S i s k i y o u Mts. , C . L . H i t c h c o c k & J . S . M a r t i n  5261 (WSP,UC,WTU); (WSP); 4.  3.  2.  M a r i o n County, J e f f e r s o n Park, Don P e t e r s 196  J o s e p h i n e County, Mt. E l i j a h , R.H.Whittaker  s s l 4 6 (WSP);  Hood R i v e r County, Mt. Hood, J.W. "Thompson 3222 (WTU);  L i n n County, H.M.Gilkey L. E. D e t l i n g 3081 (MIN);  s.n. (OSC);  6.  5.  East  Lane County, O'Leary Mt. ,  7. Clackamas County, southeast s l o p e o f  Z i g z a g Canyon, H. A. Thomas & J.A. Thomas 4331 (UC);  8.  C r a t e r Lake  N a t i o n a l Park, near K e r r Notch, W. H. Baker 6265, (UC,0SC,WTU,WSP); 9. Douglas County, B l a c k Rock Lookout, D. O v e r l a n d e r & H.M.Gilkey s.n. (OSC);  10.  Mt. Hood, P a r a d i s e Park, H.W. R i c k e t t 1635 (SMS)  Hood, P a r a d i s e Park, J.W.Thompson 5107 (MIN); 62653 (UBC)  13.  16.  15.  12. M a r i o n County Sherk  T i l l a m o o k County, Mt. Hebo, L. Constance & A. A. B e e t l e .  2657 (UC,WTU,WSP); (MIN);  11. Mt.  Mt. Hood, banks o f C l a r k Creek, J.W.Thompson 5097  C l a t s o p County, Saddle Mountain, L. E. D e t l i n g 6658 (UC);  C l a t s o p County, Saddle Mountain, Robert Ornduff 6220 (UC).  APPENDIX IV Specimens r e p r e s e n t e d to i l l u s t r a t e v a r i a t i o n  (shown i n F i g . 7)  i n the b r a c t e a l l e a v e s o f S. f e r r u g i n e a .  OREGON: 2.  1.  C l a t s o p County, Saddle Mountain, Robert O r n d u f f 6220 (UC);  C l a t s o p County, Saddle Mountain, L.E. D e t l i n g 6658 (UC);  3.  Clackamas County, southeast s l o p e o f Z i g z a g Canyon, H. A. Thomas & J. H. Thomas 331(UC). WASHINGTON:  1.  K i t t i t a s County, Mt. D a n i e l s , west o f Hyas Lake, C.L.  H i t c h c o c k 8019  (WTU);  Purer 7835 (MIN);  3.  2.  P i e r c e County, summit o f Naches P a s s , E.A.  P i e r c e County, Mt. R a i n i e r N a t i o n a l Park,  P a r a d i s e V a l l e y , L.S.Rose 40759 (UC);  Whatcom County, n o r t h o f Mt.  Baker, H.L.Mason 7442 (MIN);  5.  L u c i l e Rousch s.n. (UC);  K i n g County, Guy Peak, Snoqualmie P a s s ,  J.W.Thompson 9673 (UC); J.W.Thompson 7256 (UC, IDAHO:  6. 7.  Mt.  4.  Baker, j u s t below timber l i n e ,  Grays Harbour County, C o l . Bob Lookout,  WTU).  K o o t e n a i County,Packsaddle Mountain, J . B. L e i b e r g 468  (MIN);  Boundary County, Smith Creek, G. B. Rossbach & R.P. Rossbach 245 3. Idaho County, E l k Mountain, W.H.Baker 12593 (JEPS); County, C.B.Wilson 279  (UC,  4.  2.  (UC,WTU);  Shoshone  WSP).  MONTANA:  1.  G l a c i e r N a t i o n a l Park, near Gunsight Lake, B a s s e t t Maguire  (UC);  2.  R a v a l l i County, St. Mary's Lake, C . L . H i t c h c o c k and C. V.  857  M u h l i c k 15335 (UC,WTU,WSP). CALIFORNIA: 2.  1.  S i s k i y o u County, Poker F l a t , R . B a c i g a l u p i 7448  (MIN,JEPS);  S i s k i y o u County, Sanger Peak, C . L . H i t c h c o c k & J . S . M a r t i n 5287 (UC,  WTU). ALASKA:  1.  K o d i a k I s . , L a r s e n Bay,  E r n e s t Lepage 25071 (CAN);  2.  Raspberry I s . , P o r t V i t a , W.J.Eyerdam 3078 (UC). BRITISH COLUMBIA: (MIN);  2.  1.  Vancouver I s . , Lake Cowichan, C. 0. Rosendahl  Summit o f Mt. R e v e l s t o k e , W.A.Weber 2513  1809  (WSP,MIN,WTU).  71 APPENDIX V. Specimens r e p r e s e n t e d Fig.  1,2.  3,4. Mt.  9,10.  (shown i n  8) i n the s i z e and shape o f p e t a l s i n j>. f e r r u g i n e a .  S u z e t t e Bay, Dowager I s l a n d , B.C.,  (UBC); 5-8.  to i l l u s t r a t e v a r i a t i o n  Tar I s l a n d , B.C.,  William G r i f f i t h  William G r i f f i t h  Seymour, Vancouver, B.C.,  s.n.  s.n. (UBC);  A. S. Randhawa 1498  (UBC);  P i l l a r Mountain, Kodiak I s . , A l a s k a , A. S, Randhawa  1586  (UBC); 11,12. Montana Creek, Juneau, A l a s k a , A. S. Randhawa  1587  (UBC);  13,14.  P r i n c e Rupert, B.C.,  Highway 16, A. S. Randhawa 1588 of E v e r e t t , Washington,  W.B.  (UBC); Schofield  Bay, Queen C h a r l o t t e I s l a n d s , B.C., 19,20.  (UBC); 21,22.  A. S. Randhawa 1590  Randhawa 1500  (UBC);  s.n.  16-18.  62653 (UBC); 28,29.  east  Alliford  A. S. Randhawa 1589  (UBC);  A. S.  Mt. Moresby, Queen C h a r l o t t e  (UBC); 23-25.  26,27.  on  15. Mt. P i l c h u c k ,  Onward P o i n t , Queen C h a r l o t t e I s l a n d s , B.C.,  Randhawa 1591 B.C.,  36 m i l e s e a s t  Squamish, B.C.,  M a r i o n County, Oregon.  Islands,  A. S.  Sherk  Campbell R i v e r , Vancouver I s . , B.C.,  A. S. Randhawa 1612  (UBC); 30. L i g h t House Park, Vancouver,  A. S. Randhawa 1499  (UBC).  B.C.,  72  F i g u r e 1:  D i s t r i b u t i o n map Graham based on examined. The  of Saxifraga ferruginea the herbarium specimens  l a r g e spot r e p r e s e n t s  presumed "Type L o c a l i t y " .  the  73  F i g u r e 2:  Microsporogenesis A.  T  Mt.  X 800.  Regular  C.  D.  F.  800.  X 500.  G.  T e t r a d s , one  with  M^ w i t h chromatin o f f  the metaphase p l a t e , X 800.  H.  L a t e A^ w i t h a  b i v a l e n t a t the metaphase p l a t e , X 800. showing a l a g g a r d , X 450. N a t i o n a l Park, Oregon. X 2000. L & M.  K.  Mt.  E a r l y and  J & K.  X 1600.  M.  C h a r l o t t e I s l a n d s , B.C.  Regular T , X 700.  P-S.  N.  X 950.  M^,  Aj. w i t h l a g g a r d s , X 600.  R.  One  s m a l l and  a t e t r a d , X 600.  800.  L. 0.  N &  Clumps of p o l l e n 0.  Alliford  P.  three large microspores S.  10,  l a t e A_ w i t h l a g g a r d s , X  Seymour, Vancouver, B. C.  p o l l e n g r a i n , X 1000.  A^  C r a t e r Lake  J. Diakinesis, n =  grains germinating i n s i t u ,  Q.  I.  P i l l a r Mountain, Kodiak I s . A l a s k a .  Mj, n = 10,  19i  chromatin  with laggards, X  T , X 800.  micronucleus,  Clumping o f  1500  X  Seymour, Vancouver, B. C. n =  Regular Kj., X 800.  at M_, E.  n = 19,  Campbell R i v e r , Vancouver I s . M ,  B-I, N & 0. B.  i n S a x i f r a g a f e r r u g i n e a Graham.  Germinating Bay,  n = 10, Four  Queen X  1600.  s m a l l and  i n p l a c e of a t e t r a d X  three large microspores ( A l l m a g n i f i c a t i o n s are  600.  i n p l a c e of approximate).  74 F i g u r e 3:  G e r m i n a t i o n o f seed and the development  of  s e e d l i n g s i n S a x i f r a g a f e r r u g i n e a Graham. A & B  X 75,  C to J  X 12.  A.  The g e r m i n a t i n g seed.  B.  Two  C to J .  r a d i c l e s emerging  from the seed.  Stages i n the growth o f s e e d l i n g s .  75 F i g u r e 4:  V a r i a t i o n i n b a s a l leaves of Saxifraga Graham from C a l i f o r n i a , Montana,  ferruginea  Idaho, and  Alaska.  X 0. 7 ( D e t a i l s o f specimens a r e g i v e n i n Appendix I).  \  76 F i g u r e 5:  V a r i a t i o n i n b a s a l leaves of Saxifraga Graham from B r i t i s h Columbia.  X 0.7  specimens a r e g i v e n i n Appendix I I ) .  ferruginea  ( D e t a i l s of  77 F i g u r e 6:  V a r i a t i o n i n b a s a l leaves o f Saxifraga Graham from Washington and Oregon.  ferruginea  X 0.7  o f specimens a r e g i v e n i n Appendix I I I ) .  (Details  WASHINGTON  78 F i g u r e 7:  V a r i a t i o n i n the b r a c t e a l l e a v e s o f S a x i f r a g a f e r r u g i n e a Graham from Oregon, Washington, Idaho, Montana, C a l i f o r n i a , A l a s k a , and B r i t i s h Columbia. X 0.8  ( D e t a i l s o f specimens a r e g i v e n i n Appendix IV),  79 F i g u r e 8:  V a r i a t i o n i n the s i z e and shape o f p e t a l s i n the f l o w e r s o f S a x i f r a g a f e r r u g i n e a Graham. I, 2.  Dowager I s . , B. C. ;  5 to 8. Vancouver, B. C. ; I I , 12. Juneau;  Tar I s . , Q. C. I. ;  9,10.  Kodiak I s . ;  13,14. P r i n c e Rupert;  P i l c h u k , Washington; Q.C.I. ;  3,4.  Vancouver.  Bay,  21,22.  23 to 25. Squamish, B. C. ;  26,27. M a r i o n County, Oregon; R i v e r , Vancouver I s . ;  15. Mt.  16 to 18. A l l i f o r d  19,20. Onward P o i n t , Q. C. I. ;  Mt. Moresby, Q. C. I. ;  X 5.  28,29. Campbell  30. L i g h t House Park,  ( D e t a i l s o f the specimens a r e g i v e n  i n Appendix V).  1 \ W  T i  6  i *  11  2  1  3  7  8  12  13  1  T  10  21  22  l  23  28  24  4  f  14  19  f  ,1  4  15 _________  ^  4 t  20  * »  A  80 F i g u r e 9:  Vegetative repreduction i n Saxifraga ferruginea Graham. A.  P l a n t l e t s detached from the i n f l o r e s c e n c e . X 1.  B.  Young r o s e t t e s developed from p l a n t l e t s . X 1.  C.  F o r m a t i o n o f new crowns from the rhizomes. X 0.75.  D.  A c l u s t e r o f new crowns developed from the rhizomes around a s i n g l e r o s e t t e . X 0.75.  81  F i g u r e 10:  Reproductive Graham.  b i o l o g y of S a x i f r a g a f e r r u g i n e a  Vertical  line  (dashes) i n d i c a t e s steps  i n r e p r o d u c t i o n which may ( l e f t ) and  take p l a c e i n one  the f o l l o w i n g year  (right).  year  > Plant  Rhizome N  Seed  Plant  Plant  Flower  Inflorescence  Plantletrosette  Plantlets  _>Plant  82  Figure' 11:  Megasporo- and megagametogenesis i n JS. f e r r u g i n e a Graham. A.  Megasporocyte d i f f e r e n t i a t e d  i n the n u c e l l u s (arrow,  p o l l e n g r a i n s formed a f t e r m e i o s i s i n the m i c r o s p o r o c y t e s , X 125. (enlargement t i o n at D.  Megasporocyte i n a d e v e l o p i n g o v u l e  from F i g . A ) , foreshadowing  X 275.  Spindle orienta-  a T'-shaped t e t r a d , 1  D, but no c u r v a t u r e o f the body, X 275. G.  The  the d e g e n e r a t i n g spores, X 300.  embryo sac, X 650. J.  C.  X  675.  E. Young o v u l e , same stage as i n  the megasporocyte, X 350. and  X 300.  Young curved o v u l e w i t h megasporocyte and d e v e l o p i n g  integument, Fig.  B.  Seven-nucleate  I.  M^  of  f u n c t i o n a l megaspore H.  Two-nucleate  F o u r - n u c l e a t e embryo sac, X  embryo sac showing egg apparatus,  n u c l e u s and a n t i p o d a l s , X 525. approximate).  F.  700. fusion  ( A l l m a g n i f i c a t i o n s are  83  F i g u r e 12:  Embryogenesis A.  in  f e r r u g i n e a Graham.  Double f e r t i l i z a t i o n completed,  and endosperm n u c l e u s , X 325.  B. Endosperm n u c l e u s  d i v i d i n g b e f o r e the zygote, X 275. proembryo, X 375. E.  D.  ( v e r t i c a l ) o f the embryonal c e l l , cellular  t r u e embryo, X 325.  embryo,  X 100.  J.  C.  Two-celled  T h r e e - c e l l e d proembryo, X  F o u r - c e l l e d proembryo, X 375.  I.  showing zygote  H.  F. F i r s t X 375.  G.  450.  division Multi-  'Heart-shaped  1  Embryo i n an immature seed, X  100.  Embryo i n a mature seed, X 75. Note:  The d a r k - s t a i n i n g  conspicuous i n A-F.  layer i s  The n u c e l l a r l a y e r of e n l a r g e d  c e l l s can be seen i n C-E. approximate).  integumentary  ( A l l magnifications  are  84  Figure  13:  Anatomy o f S a x i f r a g a  f e r r u g i n e a Graham.  A.  Flowering  B.  Rhizome i n c r o s s  C.  Cross s e c t i o n o f the peduncle matic).  D.  stem i n c r o s s  s e c t i o n X 140.  s e c t i o n X 30. (diagram-  X 40.  Whole mount o f a v i t a l l y  s t a i n e d and c l e a r e d  l e a f . X 1.5. E.  Leaf i n cross s e c t i o n .  F.  Young r o o t  G.  Cross s e c t i o n of root  i n cross  X 40.  section.  X 80.  showing secondary growth.  X 80. H.  Cross s e c t i o n o f an o l d r o o t growth and c o r k c e l l s .  c.  cortex,  cc.  cork c e l l s ,  showing secondary  X 80. sx.  secondary xylem.  85  F i g u r e 14:  Anatomy o f the rhizome o f S a x i f r a g a  ferruginea  Graham. A.  C r o s s s e c t i o n o f a young rhizome (diagramm a t i c ) . X 25.  B.  C r o s s s e c t i o n o f young rhizome. X 30.  C.  E n l a r g e d view o f endodermis. X 130.  D.  C r o s s s e c t i o n o f o l d rhizome (diagrammatic), l e a f t r a c e s seen i n the c o r t e x . X 20.  E.  C r o s s s e c t i o n o f o l d rhizome. X 30.  c.  cortex,  end.  x.  xylem,  It.  endodermis, leaf  trace.  ph.  phloem,  86  Figure  15:  Vasculature Graham. A. sb. owb.  o f the flower  of S a x i f r a g a  ferruginea  X 25.  P l a n o f the f l o w e r ; s e p a l bundle,  pb.  B.  L o n g i - s e c t i o n a l view.  p e t a l bundle, stb. stamen bundle,  o v a r y w a l l bundle, p l b .  p l a c e n t a l bundle.  Fig.  15  87  F i g u r e 16:  Anatomy o f the shoot apex o f S a x i f r a g a f e r r u g i n e a Graham ( L o n g i t u d i n a l s e c t i o n s ) . A.  V e g e t a t i v e growing apex. X  B to D.  Young i n f l o r e s c e n c e s i n v a r i o u s stages  development. E.  B. X 95;  C.  X 85;  D.  D i f f e r e n t i a t i o n o f p l a n t l e t s i n the scence. X  F.  95.  95.  inflore-  35.  P l a n t l e t s developed i n p l a c e o f f l o w e r s , p e d i c e l of terminal flower i n c r o s s s e c t i o n . X  G.  X  of  Cross  s e c t i o n o f a f l o w e r bud.  X  35.  30.  88  F i g u r e 17:  The t w o - d i r e c t i o n a l chromatogram showing spots o f p h e n o l i c compounds developed w i t h spray  p-nitroaniline  from b a s a l l e a v e s o f S a x i f r a g a f e r r u g i n e a  Graham.  Fig.  00  m R >  m H  >  O  0  n  FORMIC ACID 8/  17  89 F i g u r e 18:  Intraspecific  c r o s s e s between d i f f e r e n t  populations  o f S a x i f r a g a f e r r u g i n e a Graham. S i n g l e arrows  ^ one way c r o s s , arrow p o i n t i n g to  the female p a r e n t s . Double a r r o w s - * — ^ r e c i p r o c a l c r o s s e s . +  successful crosses.  -  unsuccessful  +?  crosses.  successful cross doubtful. (seed  s e t o n l y i n one c a p s u l e ) .  Washington n = 19  90  F i g u r e 19:  G l a c i a l map  o f western N o r t h America.  G l a c i a t e d areas are  stippled.  Fig.  19  

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